Contributions from the United States National Herbarium

BULLETIN OF THE UNITED STATES NATIONAL MUSEUM

CONTRIBUTIONS FROM THE UNITED STATES NATIONAL HERBARIUM

Votume 35

SYSTEMATIC PLANT STUDIES

Exisworts P. Kituip, Jose Cuatrecasas, Lyman B. Smita,
and JosePH Ewan

SMITHSONIAN INSTITUTION PRESS ¢ WASHINGTON, D.C. ¢ 1968

Publications of the United States National Herbarium

The United States National Herbarium, which was founded by
the Smithsonian Institution, was transferred in the year 1868 to the
Department of Agriculture and continued to be maintained by that
department until July 1, 1896, when it was returned to the official
custody of the Smithsonian Institution. The Department of Agri-
culture, however, continued to publish the series of botanical reports
entitled “Contributions from the United States National Herbarium,”
which it had begun in the year 1890, until, on July 1, 1902, the Na-
tional Museum, in pursuance of an act of Congress, assumed
responsibility for the publication. The first seven volumes of the
series were issued by the Department of Agriculture.

Frank A. Tayor,
I%rector, United States National Museum.

ii

Parr 1.

Parr 2.

Part 3.

Part 4.

Parr 5.

Parr 6.

CONTENTS

Supplemental notes on the American species of
Passifloraceae, with descriptions of new species.
By Ellsworth P. Killip. (Published May 10,
1960) . 2... ee ee

A taxonomic revision of the Humiriaceae. By Jose
Cuatrecasas. (Published April 14, 1961)... .

Origins of the flora of southern Brazil. By Lyman
B. Smith. (Published March 19, 1962)

A Synopsis of the American Velloziaceae. By Lyman
B. Smith. (Published March 19, 1962)

Synopsis of the South American species of Vismia
(Guttiferae). By Joseph Ewan. (Published June 5,
1962)... 2. 2

Cacao and its allies: A taxonomic revision of the
genus Theobroma. By Jose Cuatrecasas. (Pub-
lished August 21,1964)... .. 2.2.2...

Page

1-23

25-214

. 215-249

. 251-292

293-377

379-614
iff

UNITED STATES NATIONAL MUSEUM

CONTRIBUTIONS FROM THE UNITED States NATIONAL HERBARIUM

VoLuME 35, Part 1

SUPPLEMENTAL NOTES ON THE AMERICAN
SPECIES OF PASSIFLORACEAE WITH
DESCRIPTIONS OF NEW SPECIES

By Evitsworts P. Ki.uie

BULLETIN OF THE UNITED States NationaL Museum

SMITHSONIAN INSTITUTION *« WASHINGTON, D.C, « 1960

Preface

This paper was partly prepared by Ellsworth P. Killip, 10 years
after the publication of his monograph of the Passifloraceae in 1938,
to summarize the additional data acquired in the 10-year period,
and to present descriptions of 8 new species. In 1956, Mr. Killip
turned over the manuscript to Dr. José Cuatrecasas to use as he saw
fit in the preparation of a flora of Colombia. It is now published
with the addition of three new species that Mr. Killip indicated as
new but did not describe. Dr. Cuatrecasas wrote descriptions for
these as well as Latin diagnoses for all the new species described by
Mr. Killip. Included are 11 new species and citations of all new
material examined since the publication of the monograph. Dr.
Lyman B. Smith, curator of the division of phanerogams, has care-
fully edited the manuscript.

Jason R. SwALLEN
Head Curator, Department of Botany

SUPPLEMENTAL NOTES ON THE AMERICAN
SPECIES OF PASSIFLORACEAE, WITH
DESCRIPTIONS OF NEW SPECIES

By Extswortu P. Kini

Introduction

In 1938 the Chicago Natural History Museum (then the Field
Museum of Natural History) published a monograph on the American
species of Passifloraceae written by the present author! The prepara-
tion of this monograph covered a period of many years, during which
time I made three visits to Europe and consulted nearly all the type
specimens as well as many others of historic importance. A large
amount of material was generously placed at my disposal by those
in charge of many of the larger herbaria of the world. It is therefore
perhaps not surprising that in the intervening years I found little
reason to revise the treatment of the species presented in the mono-
graph.

The intensive exploration carried on in certain parts of northern
South America, especially in the basins of the Orinoco and Amazon in
Colombia and Venezuela, resulted in the collecting of many Passi-
floraceae in regions little known in 1938. In the course of studying
these specimens, as well as material from Central America and other
parts of South America, I detected several new species, some of
which I described elsewhere. This material also shows interesting
extensions of ranges of certain species, and in some instances permits
a desirable amplification of the descriptions given in the monograph.
It therefore seems advantageous to assemble in one place all this
subsequent information along with descriptions of several new species.
The numbers associated with the species of Passiflora are those of
the monograph.

4. Passiflora gracillima Killip

This species is now represented in the U.S. National Herbarium
by several additional Colombian collections. One specimen from the

'Field Mus. Bot. 19. 1938.

2 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Province of Chiriquf, Panama, was cited; the species was recollected
in the vicinity of Chiriqui Lagoon in 1941 by H. von Wedel (No.
2421). Likewise, a single specimen was listed from Riobamba,
Ecuador. This species is now represented by Steyermark 52932,
which was collected between Cruz Pamba and Loma de Canela,
north of Molleturo, Province of Azuay, Ecuador, at an altitude of
2,315-2,500 m. The collector noted that the fruit is edible and has a
flavor similar to that of Annona.

10. Passiflora gracilis Jacq.

Additional collections are:

VENEZUELA: Bolfvar: Gran Sabana, Cerro Kanaimé Santa Elena, March
1946, Tamayo 3025.
Ecuvapor: Azuay: Between Rio Gamolotal and Rio Norecay, Steyermark

52879,

Both of these collectors noted that the fruit is red. Steyermark
gave “shunshun” as a local name, and stated that the corona fila-
ments are white with a purple streak at the center and that the
seeds are red.

14a. Passiflora ursina Killip & Cuatr., sp, nov. PLaTE 1

Frutex elatus scandens; rami foliosi dense rufo-tomentosi; stipulae
subulatae; folia coriacea trinervia ovato-oblonga vel oblonga basi
truncata vel rotundata apice acuta vel subacuta mucronulata utroque
latere saepe 1-lobata, lobo triangulari brevi mucronulato, subtus
rufo-hirtella vel tomentella, margine integra revolutaque, petiolo
brevi basi duobus glandulis minutis inconspicuis; bracteae liberae
subulatae; calycis tubus petalliformis, sepalis ovatis apice rotundatis
subapicem corniculatis, petalis paulo brevioribus; corona filamentis
biseriatis apice incrassatis exterioribus robustioribus; operculum
membranaceum plicatum erectum breviter fimbriatum; ovarium
glabrum.

Large, scandent shrub; leafy branches moderately thin, flexuose
subterete, densely reddish brown tomentose with acute, subrigid,
spreading hairs, 0.5-1 mm. long; tendrils closely spiral, tomentose;
stipules subulate, hirsute, 5-6 mm. long, 0.5-1 mm. broad; petioles
8-14 mm. long, stout, straight or bent at base, densely rufo-tomentose
(like the branches), the glands small, semiglobular, sessile at base,
concealed by the indument; blades coriaceous, trinerved, ovate-oblong
or oblong, 5-8.2 cm. long, 3-4.5 cm. broad, rounded, truncate or
cordate at base, subtrilobate (or rarely entire) near the apex, the two
lateral lobes very short, form’ng an obtuse mucronulate angle, the
terminal one (the apex) acute or subacute, mucronulate, the margin
entire, revolute, the surface puberulous with spreading, sparse hairs

KILLIP—NOTES ON AMERICAN PASSIFLORACEAE 3

above, pubescent or subtomentose with spreading, long, rufous hairs
beneath, the 3 nerves and veins very prominent beneath, little con-
spicuous above; a few minute, semiglobose glands near the base
beneath; inflorescence cymose, the cymes in pairs with 5-7 flowers,
the rachis less than 1 cm. long; bracts subulate, 3-5 mm. long, up to
0.5 mm. broad, rufo-hirsute; the peduncles thick, straight, rufo-
tomentose, 8-14 mm. long, articulate about the middle or slightly
above the middle; flowers 2.5-3 cm. wide, greenish white; sepals
ovate, obtuse, 1.1-1.3 cm. long, 0.7—-0.9 cm. wide, dorsally more or
less hirsute and keeled, corniculate below the apex, the horn up to
3 mm., hairy; petals membranaceous, ovate, narrowed at both ends,
obtuse, subdentate below the apex, 1-1.1 cm. long, 0.6-0.7 cm. wide;
corona filaments filiform, much dilated at apex, in 2 series, the outer
about 3.5 mm. long, rigidulous, the inner about 2.5 mm. long, thinner
and flexuose; operculum membranaccous, closely plicate, short-
fimbrillate, 1.5 mm. high; limen annular; gynophore absent; ovary
ellipsoid-obovoid, glabrous.

Type in the U.S. National Herbarium, No. 1799118, collected at
Péramo de Santa Lucia, south side of Laguna de la Cocha, Depart-
ment of Narifio, Colombia, at an altitude of 2,900-3,100 m., on
Jan. 9, 1941, by J. Cuatrecasas (No. 11886).

P. ursina is closely related to P. apoda Harms and P. sodiror
Harms, from which it differs in the thick, longer, rufous indument,
in the shorter petioles with sessile, minute basal glands concealed by
the indument, and more conspicuously in the horned sepals. It
differs from P. sodiroi in lacking the gynophore and in the form of
the petals, and from P. apoda by the shorter corona filaments.

26. Passiflora dolichocarpa Killip

Known only from the type specimen at the time of my monograph,
this species is now represented in the U.S. National Herbarium by
the following additional collections:

Mexico: Chiapas: Hacienda Siltepec, Matuda 1892.

GuatTemaua: Jalapa: Between Jalapa and Montafia Miramundo, alt. 1,500-—
2,000 m., Steyermark 32884 (F). Suchitepéquez: Volcdn Santa Clara, alt.
1,250—2,650 m., Steyermark 46761.

46a. Passiflora macrostemma Killip, Field Mus. Bot. 23:217. 1947.

Additional collections are:

GuatremaLa: Chiquimula: Voledn de Quezaltepeque, alt. 1,500-2,000 m.,
Steyermark 31519 (F, type, US photo and fragment), 31281 (F). Jalapa:
Montafia Durazno, alt. 1,400-1,900 m., Steyermark 32985 (F).

4. CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

61. Passiflora lancearia Mast.

Recorded as being confined to Costa Rica, this species was collected
in the Province of Chiriquf, Panama, in 1938 by M. E. Davidson
(No. 336) and in the Province of Bocas del Toro in 1941 by H. von
Wedel (No. 2120).

63. Passiflora panamensis Killip

This species was known at the time of my monograph only from
the type, which came from southern Darién, Panama. ‘The species
has been found in the adjacent coastal part of Colombia, a recent
record being:

Corompra: El] Chocé: Bahfa Solano, dense forest along the Quebrada
Jellita, alt. 50-100 m., Feb. 22, 1939, Killip & Garcia 33482.

69. Passiflora bauhinifolia H.B.K.

P. andreana Mast. was treated as a synonym of P. bauhinifolia in
my monograph. Specimens collected by Garcia-Barriga and Balls in
southwestern Colombia in 1939 show that these two species are
unquestionably distinct. In P. andreana the inside of the sepals and
the petals are purple, reddish purple, or deep violet;-in P. bauhinifolia
these parts are white. The outer corona filaments are slenderer in
P. andreana.

Of the specimens listed in my monograph under P. bauhinifolia,
I have rechecked only those in the U.S. National Herbarium; on this
basis, P. bauhinifolia is common in Ecuador, and P. andreana is
known only from Colombia. P. bauhinifolia is represented in the
U.S. National Herbarium by the following:

Cotomsia: El Valle: Quebrada de la Palma, Rio Bugalagrande Valley,
Cuatrecasas 20935. Putumayo: Laguna de la Cocha, Péramo El Tabano,
Garcia-Barriga 7785A.

Ecuapor: Without locality, Mille. Imbabura: Otovalo, Mezia 7645.
Pichincha: Quito, Humbold & Bonpland (photo of type); Mille 222, s.n.
in 1918. Verde Cruz, Firmin 310. Gudpulo, Stevens 258. Tungurahua:

Bafios, Lehmann, 675, 8020.
Peru: Cajamarca: Pefia Rota, toward Cochabamba 2,800 m., Ochoa 1565.

This last record is apparently the second of this species in Peru.
69a. Passiflora andreana Mast. Journ, Linn. Soc. 20:37, 1883.

Additional collections are:

Coromsia: Narifio: Las Lajas, near Ipiales, André 3478 (photo of type);
Garcta-Barriga 7757A; Balls 7360. Jenoy, 5 km. northwest of Pasto,
alt. 2,460 m., Hwan 15899 (USNA). Between Pasto and Anganoy,
Schultes & Villarreal 7417.

KILLIP—NOTES ON AMERICAN PASSIFLORACEAE 0

100. Passiflora micropetala Mast.

In the monograph I stated that this species was one of the few
maintained as valid by Masters in the Flora Brasiliensis that I knew
only from description. After studying a specimen subsequently col-
lected along the Rio Putumayo, I have decided that it and a speci-
men from the Comisaria of Caqueté, Pérez-Arbeldez 669, which was
cited by me as P. dawei, both represent P. micropetala. The type
locality of this species as given by Masters is: ‘In regione fluvii Japurd,
prov. do Alto Amazonas.” The Rio Japurd is identical with the Rio
Caqueté, and the Putumayo is in the same general area. On his trip
to this river, Martius is known to have collected in present-day
Colombia, and the type may thus have come from this country.

The Pérez collection is sterile, and the more recent one is in fruit
only. Both resemble P. biflora in leaf shape, but the coronal structure
is different according to Masters’ description. P. micropetala there-
fore appears to be known from the following locations:

Cotomsia: Caqueté: Florencia, alt. 420 m., Pérez-Arbeldez 669 (COL, US).
Putumayo: Selva higréfila del Rio Putumayo en Puerto Ospina, alt, 230 m.,

Cuatrecasas 10776 (COL, US).
Brazit or Cotomsia: Rio Japurd (R. Caqueté), Martius (M, type).

111. Passiflora adenopoda DC.

A recent collection of this species by Dugand and Jaramillo (No.
3965) has far more deeply cleft bracts than heretofore observed.
They are deeply palmate-trisect, and the segments are repeatedly
cleft. The largest bract is 2 cm. wide. This unusual variant was
collected in the Department of Cundinamarca, Colombia, at Es-
tacién San Bernardo, between Sasaima and Alban, at an altitude of
1,700-1,800 m.

132. Passiflora bicuspidata (Karst.) Mast.

This species, known hitherto only from the Eastern Cordillera of
Colombia, was recently collected by Steyermark across the border
in Venezuela. His detailed notes on the color of the flowers are worth
reproducing as earlier collectors have described the flower color in
only general terms.

“Calyx rose-lavender, the lobes with more lavender, the tube with
more brick-color; coronal filaments golden; calyx tube below the coro-
nal filaments within dull pale greenish-yellow; filaments purple; an-
thers buff; styles purplish; petals purplish-rose.”’

The collection was made at:

VENEZUELA: Tdchira: Swampy woods between Paramito and Quebrada de

Palmar, at base of Piramo de Tam4, 7 km. above Villapdez, alt. 2,500 m,
Steyermark 57240 (F, US).

6 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

135. Passiflora sanguinolenta Mast.

In the monograph I followed Masters in placing this Ecuadoran
species in the subgenus Psilanthus, but I observed that it appeared
more closely related to P. rubra, of the small section XYerogona of the
subgenus Plectostemma, than to the three known species of Psilanthus.

Later, there were submitted to me, through the courtesy of Harold
N. Moldenke, then of the New York Botanical Garden, a fine series
of specimens of P. sanguinolenta and P. rubra collected by Reinaldo
Espinosa in the Province of Loja. One of P. sanguinolenta, No. 806,
is in fruit, apparently the first time that fruit of this species has been
collected. The transverse ridges of the seeds are rugose, not smooth
as in Xerogona, and I believe that the species is best left in Psilanthus,
even though in vegetative parts (especially in the absence of any
bracts) and shape of the fruit it very closely resembles P. rubra.

To the collections cited in the monograph there may now be added
the following, all from the Province of Loja and deposited in the U.S.
National Herbarium:

Ecuapor: Catacocha, alt. 2,050 m., Espinosa 617. Pucala, north of Loja,
alt. 2,300 m., Espinosa 621. Between San Pedro and Chinchas, about 55
km, west of Loja, alt. 1,200 m., H’spinosa 1314. Aguahedionda, north of

Loja, alt. 2,150 m., Espinosa 806. Desert country between Vilcabamba
and Cachiyacu, alt. 1,600-2,100 m., Steyermark 54397.

137. Passiflora bracteosa Planch. & Linden
Tacsonia infundibularis Mast.

Both these species were based upon Funck and Schlim’s 1381. The
type locality of P. bracteosa was correctly given as La Baja, Colombia;
that of Tacsonia infundibularis was incorrectly recorded as ‘“Vene-
zuela.”” However, this species has recently been collected by Steyer-
mark (No. 57277, at F) in the State of Tachira, Venezuela, at the
base of the Péramo de Tam4, at an altitude of 2,285-2,430 m.

147. Passiflora adulterina L-f.

Although this species is fairly common in the vicinity of Bogoté,
fruiting material was not found, apparently, until 1939. Cuatrecasas
5136, collected between El Delirio and Guadalupe (eastern slope of the
Cordillera Oriental, between 2,900 and 3,220 m. altitude), bears mature
fruit, which may be described as follows:

Fruit broadly ovoid, 7.5 cm. long, about 4.5 cm. in diameter,
apparently reddish brown, mottled with greenish yellow; seeds
obovate, about 9 mm. long, and 4.5 mm. wide, reticulate, dark brown,
lustrous.

KILLIP—NOTES ON AMERICAN PASSIFLORACEAE 7

148a. Passiflora cuatrecasasii Killip, sp. nov. PLATE 2

Suffrutex clata scandens, caule primum lanato denique glabrato;
stipulae subulatae; folia coriacea ovato-lanceolata vel oblongo-
lanceolata plus minusve serrata basi rotundata vel subcordata apice
acuta vel subobtusa subtus lanata, petiolo sub apice 2-3 glandulis;
bracteae liberae ovato-oblongae vel anguste oblongae acutae serru-
latae; calycis tubus cylindricus glaber, sepalis tubo valde brevioribus
oblongis obtusis, petalis sepala subaequalibus; corona annularis
undulata; operculum deflexum denticulatum leviter recurvatum;
ovarium tomentosum.

Vine; stem subangular, lanate when young, the indument at length
evanescent; stipules subulate, up to 1.1 em. long, glabrous, purplish
black below middle, the upper part spongiose-thickened, light yellow-
ish brown, coriaccous, acuminate; petioles up to 2 cm. long, bearing 2
or 3 glands near apex; leaves unlobed, ovate-lanceolate or oblong-
lanceolate, 6-11 cm. long, 3-6.5 cm. wide, acute or subobtuse,
rounded or cordulate at base, serrate or serrulate, penninerved (princi-
pal lateral nerves about 9 to a side), coriaceous, dark green, lustrous,
and glabrous above, densely lanate beneath, except sometimes on the
nerves and veins; peduncles 7-10 em. long, slender, evanescently
lanate, articulate about 1 cm. below the apex; bracts distinct to the
base, ovate-oblong to narrowly oblong, 3-4 cm. long, 0.8-3 cm. wide,
acute, sessile, finely serrulate; calyx tube cylindric, 8-10 cm. long,
1-1.5 cm. wide, greenish at base, red or purplish above, glabrous;
sepals oblong, 3.5-4.5 em. long, 1-1.5 cm. wide, obtuse, dorsally
awned just below the apex (awn about 1.5 mm. long), pinkish purple;
petals subequal to the sepals; corona a single indulate ridge at the
apex of the tube; operculum borne near the base of the tube, 4-7 mm.
long, dependent, the margin slightly recurved and minutely dentic-
ulate; ovary ovoid, densely tomentose.

Type in the U.S. National Herbarium, No. 1796435, collected on the
eastern slope of the Péramo de Guasca, Cordillera Oriental, Depart-
ment of Cundinamarca, Colombia, at an altitude of 3,200-3,300 m. on
June 2, 1940, by J. Cuatrecasas (No. 9479). Also represented by J.
Cuatrecasas 13538 from the same locality, and by Uribe 1061, from
La Rusia, northwest of Duitama, Department of Boyacé, at an
altitude of 2,700 m.

The proposed species probably should be placed nearest P. lanata
in the arrangement adopted in my monograph, though it also has
affinities with P. rugosa and P. leptomischa. From P. lanata it differs
in having a scantier indument on the vegetative parts, a pubescent
ovary, and generally longer peduncles. In P. rugosa the bracts are
coalesced partway and the calyx tube is much shorter. P. leptomischa
is a species with dimorphic foliage and very long (15-25 cm.) peduncles.

512425—60——2

8 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

148b. Passiflora rigidifolia Killip sp. nov. PLaTE 3

Caulis scandens angulatus sursum albopilosus basin versus gla-
bratus; stipulae anguste lineares glanduloso-serrulatae subrevolutae;
folia coriacea lanceolata integra basi rotundata apice acuta praeter
costam supra pilosulam glabra, petiolo albo-piloso 2-3 glandulis parte
media; bracteae liberac lanceolatae integrae; calycis tubus cylindricus
glaber sepalis tubo brevioribus oblongis obtusis, petalis sepalis simi-
libus; corona annularis dentata; operculum deflexum recurvatum
minuteque denticulatum; ovarium tomentosum.

Stem subangular, white-pilose toward the apex, becoming glabrous
below; stipules narrowly linear about 1 cm. long and 1 mm. wide,
acuminate, glandular-serrulate, subrevolute, coriaceous; petioles up
to 7 mm. Jong, 2- or 3-glandular near the middle, white-pilose; leaves
lanceolate, 4-6.5 cm. long, 1-1.8 cm. wide, acute at apex, rounded at
base, entire, revolute (?), penninerved (midnerve impress2d above,
the lateral nerves and veins somewhat elevated on both surfaces),
pilosulous on the midnerve above, otherwise glabrous, coriaccous,
glaucescent beneath; peduncles solitary in the axils, 2-2.5 cm. long,
glabrous or sparingly pilose; bracts borne about 5 mm. below the
apex of the peduncle, free to base, lanceolate, about 2 cm. long and
8 mm. wide, acuminate, sessile, entire, pilose; flowers pale rose; calyx
tube cylindric, 6.5-7 cm. long, 8-9 mm. in diameter, slightly enlarged
at base, glabrous; sepals oblong, about 2.5 cm. long, and 1 cm. wide,
obtuse, dorsally aristate just below apex, the awn 2 mm. long; petals
similar and subequal to the sepals, about 8 mm. wide; corona a dark
ring at the throat of the tube, etuberculate; operculum borne close
to base of tube, 4 mm. long, dependent, the margin recurved and
minutely denticulate; ovary ovoid, densely tomentulose.

Type in the Herbarium of the Royal Botanic Gardens, Kew,
collected in Colombia at an altitude of about 3,900 m., 1894-1896,
by D. Burke (No. 184).

This species is nearest to P. cuatrecasasii but differs in its much
narrower leaves, shorter peduncles, and slenderer calyx tube.

Unfortunately, the locality at which the type was collected is
uncertain. The label reads ‘Flora of Colombia. Locality: Conig,
altitude 13,000 ft., Comm. Messrs. J. Veitch, 1897.” The word
“Conig” is not a Spanish one, and I have been unable to find in any
gazeteer a Colombian name at all similar to it. Conejo, in Antioquia,
on the railroad from Puerto Berrfo to Medellin, suggests itself, but
there is no altitude of 13,000 ft. attainable from there. In Peru there
is a “Cofiec,” but it is not known that Burke collected in that country.

Accompanying this specimen is a letter dated April 4, 1902, from
James Veitch & Sons to Hemsley, in response to a request for infor-
mation about Burke’s collections. Nos. 146, 154, and 167 are listed

KILLIP—NOTES ON AMERICAN PASSIFLORACEAE 9

without localities, but the Veitch letter observes that the preceding
numbers were collected in Antioquia. Nos. 184 and 185 follow both
from ‘‘Conig,’”’ number 185 being given as “Berberis sp (Goudoti ?),
11,000 ft.” Berberis goudoti is known only from the eastern Cordillera
of Colombia.

149. Passiflora truxillensis Planch. & Linden.

This rare species hitherto known only from two collections from
the State of Trujillo, Venezuela, was recently collected in the neighbor-
ing State of Mérida by Steyermark, from whose notes the following
more complete description of the flowers may be presented:

Bracts pale green with pale rose line down the center; calyx tube
pale green at very base, dull rose without, a greenish part in the
middle; sepals deep rose at margin, lavender in the middle; petals
rose-colored throughout; stamen-filaments rose-orchid; ovary green;
styles rose-lavender above, whitish below.

The complete collection data are:

VENEZUELA: Mérida: Rich forest between Mesa del Trapiche and El
Molino, alt. 2,530-2,590 m., Steyermark 56503 (F, US).

154a. Passiflora boyacana Killip sp. nov. PLATE 4

Herba scandens, caule lanato denique glabrescenti; stipulae lineari-
subulatae; folia coriacea profunde trilobata basi subtruncata vel sub-
cordata lobis lineari-lanceolatis subacutis mucronulatis integris, subtus
dense rufo-lanata, petiolo eglanduloso lanato; bracteae liberae integrae
extus lanatae; calycis tubus cylindricus glaber, sepalis petalisque tubo
dimidio brevioribus elliptico-oblongis obtusis mucronulatis; corona
brevem annulum minute tuberculatum reducta; operculum deflexum
recurvatum denticulatum; ovarium glabrum.

Herbaceous vine; stem stout, subterete, about 3.5 mm. in diameter,
lanate, becoming glabrous; stipules linear-subulate, about 1.2 cm.
long; petioles 1-1.5 cm. long, glandless, lanate in lines; leaf-blades
lobed about three-quarters their length, 4-8 cm. along midnerve,
3.5-6.5 em. along lateral nerves, 3-9 cm. between apices of lateral
lobes, (lobes 1-1.5 cm. wide, narrowly lanceolate or the middle one
sometimes elliptic-lanceolate, subacute, mucronulate), subtruncate or
cordulate at the base, entire, coriaceous, essentially glabrous above
with the nerves slightly impressed, densely rufo-lanate beneath;
peduncles solitary, about 4 cm. long, pubescent like the petioles;
bracts borne at base of flower, 2—-2.3 cm. long, 8-10 mm. wide, free to
the base, sessile, entire, deciduous-lanate without, glabrous within;
flowers young; calyx tube cylindric, 4 em. long, glabrous; sepals and
petals equal, about 2 em. long, obtuse, the sepals 7 mm. wide, mucron-
ulate, the petals 1 em. wide; corona reduced to a low, minutely tuber-

10 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

culate ridge; operculum borne at base of tube, dependent, about 4 mm.
long, the margin recurved, denticulate; ovary ellipsoidal, glabrous.

Type in the U.S. National Herbarium, No. 1796049, collected in
paramo thickets, above Puente de Boyacé, Department of Boyaca.
Colombia, at an altitude of 2,800 m. on Feb. 24, 1940, by E. Pérez-
Arbeliez & J. Cuatrecasas (No. 8083).

This species, belonging to the subgenus Tacsonia, keys out to species
154-157 of my monograph of the American Passifloraceae. The
narrow, entire leaf lobes are quite unlike those of P. ampullacea. In
depth of lobation the leaves are rather suggestive of P. trianae and P.
purdier, which are essentially glabrous plants.

155a. Passiflora raimondii Killip, sp. nov. PLaTE 5

Herba scandens, ubique glabra; stipulae setaceae; folia membranacea
trilobata, lobis oblongis vel ovato-oblongis, obtusis, denticulatis, peti-
olis eglandulosis; bracteae liberac, lanceolatae, acutae; calycis tubus
cylindricus, sepalis tubo brevioribus, lineari-oblongis, petalis anguste
linearibus; corona annuliformis, crenulata; operculum deflexum.

Herbaceous vine, glabrous throughout; stem quadrangular, longi-
tudinally ridged, the tendrils very slender; stipules setaceous, about
4 mm. long; petioles slender, 1-3 cm. long, glandless; leaves 3-5 cm.
long, 4.5-8 cm. wide, 3-lobed to below the middle (lobes oblong or
ovate-oblong, up to 3 cm. wide, rounded and sometimes slightly
emarginate at apex, denticulate), cordate at base, membranous, bi-
glandular at base; peduncles solitary, slender, about 5 cm. long; bracts
lanceolate, 1-1.2 cm. long, 4-5 mm. wide at base, acute at apex,
rounded at base, sessile, minutely denticulate, distinct to base, borne
at base of flower; calyx tube cylindric, 3-3.5 em. long, 7-8 mm. in
diameter, narrowed at base; sepals linear-oblong, 2 cm. long, 3-3.5 mm.
wide, obtuse, slightly carinate, the keel terminating in a filiform awn
1 mm. long; petals narrowly linear, 1.5 cm. long, 1.5 mm. wide, sub-
acute; corona a low ridge, crenulate; operculum borne about 5 mm.
above base of tube, dependent, 4 mm. long, minutely denticulate;
gynophore very slender, about 5 cm. long; ovary fusiform.

Type in the Herbarium of the Universidad Nacional Mayor de
San Marcos, Lima, collected at Chorrillos, Department of Cajamarca,
Peru, at an altitude of 2,200 m. on June 8, 1875, by A. Raimondi
(No. 7807). Represented also by A. Raimondi 8241, collected at
Cascas, Province of Contumazé, in the same Department.

This species is most nearly related to P. gracilens, which is rather
common in Peru and extends from Hudnuco southward. The new
species differs in its larger leaves with much broader lobes, its longer
peduncles and larger flowers, and its:lanceolate, not ovate, bracts.

KILLIP—NOTES ON AMERICAN PASSIFLORACEAE 11

The Raimondi specimens were submitted to Harms, who suggested
that they probably belonged to our undescribed species.

159a. Passiflora zamorana Killip sp. nov. PuatE 6

Herba scandens folia supra et flores excepti dense griseo- vel brunneo-
tomentosa; stipulae late ovato-reniformes semiamplexicaules plus
minusve dentatae vel incisae; folia membranacea usque medium
3-lobata basi subcordata lobis oblongo-lanceolatis argute acuminatis
serratis supra pilosula deinde glabrata, petiolo 3-4 glandulis incon-
spicuis praedito; bracteae liberae ovato-lanceolatae serrulatae; calycis
tubus cylindricus glaber, sepalis petalisque ovato-oblongis obtusis tubo
paulo brevioribus; corona circulum minutorum tuberculorum reducta;
operculum deflexum recurvatum; ovarium brunneosericeum.

Herbaceous vine, densely brownish or grayish tomentose throughout
except the upper side of the leaves and the flowers; stem terete;
stipules broadly ovate-reniform, about 1 cm. long and 1 cm. wide
at the base, scarcely oblique, closely semiamplexicaul, irregularly
dentate or laciniate at the apex, the divisions aristate; petioles 2—2.5
cm. long, bearing 3 or 4 obscure glands; leaves 6-9 cm. long, 8-9 cm.
wide, 3-lobed about to the middle (lobes oblong-lanceolate, 2.5-3.5
cm. wide, sharply acuminate, the lateral somewhat ascending),
conspicuously serrate, subcordate, membranous, the upper surface
sparingly pilosulous glabrate; peduncles 12-15 cm. long, rather stout;
bracts free to base, ovate-lanceolate, about 3 cm. long, and 1.7 cm.
wide, acuminate, serrulate, sessile, borne at the apex of the peduncle;
calyx tube cylindric, 7-7.5 cm. long, 1-1.3 em. wide when pressed flat,
slightly enlarged at base, otherwise of uniform diameter, glabrous;
sepals ovate-oblong, 6-6.5 cm. long, about 2.5 cm. wide, obtuse,
dorsally awned just below the apex (awn very slender, about 5 mm.
long), deep salmon-rose, paler within, the midnerve with a pale green
center; petals similar and subequal to the sepals; corona a single
series of minute tubercles; operculum borne near the base of the tube,
about 3 mm. long, dependent, the margin recurved; ovary ovoid,
densely light-brown-sericeous.

Type in the Herbarium of the Chicago Natural History Museum,
No. 1204733, collected in the vicinity of Achupallas, Province of
Santiago-Zamora, Ecuador, at an altitude of 2,500 m. in a forest along
a stream on October 8, 1943, by Julian A. Steyermark (No. 54493).
Duplicate at US.

In the monograph the new species keys out to P. gamesonti because
of the large stipules; however, these are far more deeply fringed in
P. jamesonii. Moreover, P. jamesonii is glabrous throughout and
has shorter peduncles and deeply fringed bracts. In leaf shape,
indument, and length of the peduncles, the new species suggests
P. ampullacea.

12 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

162. Passiflora matthewsii (Mast.) Killip

Masters placed this species among the tacsonias with united bracts,
lobed leaves, and narrowly linear stipules, though in the description
of the species, which was given at some length in the key, he did not
specifically mention the stipules. A specimen recently collected at
the type locality, Chachapoyas, Peru, by Metcalf (Goodspeed Expedi-
tion No. 30784) has foliaceous, semiovate stipules which are deeply
pinnatisect into filiform segments, the body of the stipules sometimes
being reduced to a mere band surrounded by the filiform segments.
Otherwise the specimen agrees perfectly with a photograph that I
took of the type at the Kew Herbarium and with the amplified descrip-
tion given in my monograph. I believe that both Masters and I
mistook either the filiform segments or abortive tendrils for stipules,
that the stipules of P. matthewsii had mostly fallen off the type
material at Kew and the British Museum, and that therefore P.
matthewsit should be placed next to P. roseorum (species 164).

A. H. G. Alston has kindly examined for me the Matthews specimen
at the British Museum, and writes: ‘The stipules appear to have
fallen off our specimen, but there are sometimes young tendrils which
could easily have been taken for stipules. ... On the young parts
there are, however, some filiform hairy segments, which could easily
be the segments of a laciniate stipule.”

166a. Passiflora loxensis Killip & Cuatr., sp. nov. PLATE 7

Suffrutex scandens, caule angulato glabro; stipulae coriaceae ovatae
fimbriato-dentatae; folia coriacea glabra trilobata lobis triangulari-
Ovatis acutis serratis, subtus reticulo venorum valde elevato con-
gesteque incrassato; bracteae ad medium connatae; calycis tubus
eylindricus glaber, sepalis petalisque oblongis tubum subaequilongis;
corona tuberculata; operculum deflexum paulo laciniatum recurva-
tum; ovarium glabrum.

Vine; stem acute-angular, glabrous, lustrous; stipules coriaceous,
glabrous, ovate or lance-ovate, acuminate, oblique and cordate at
base, amplexicaul, fimbriate-dentate, 10-20 mm. long, 4-8 mm. broad;
petioles 1-2 cm. long, bearing 3-4 sessile or short-stipitate glands
about 1 mm. thick; leaf-blades 4-6 em. long, 5.5-9 cm. broad, deeply
3-lobed (lobes triangular-ovate, acute, 2-3.5 cm. long, 1.5-2.8 cm.
wide, the lateral ones subascending), truncate or subrounded at base,
callous-serrate, coriaceous, glabrous, above green, rugulose, the nerves
impressed, the minute reticulate veins prominulous, beneath paler,
the 3 primary nerves thick and prominent, the secondary nerves
prominent, the veins thick forming an incrassate reticulum separating
very narrow, deep, irregular alveolac; peduncles 2.5-4.5 cm. long,
subangular, stout; bracts subcoriaceous, 5-6 em. long, connate to

KILLIP—NOTES ON AMERICAN PASSIFLORACEAE 13

about the middle (the free part oblong, abruptly acuminate, entire,
1.4-2 em. wide), outside glabrous, slightly nervate and reticulate,
inside more or less lanuginous or pilose and conspicuously venose;
flowers lilac-blue; calyx tube cylindric, 7-8 cm. long, about 1 cm.
broad when pressed, of uniform diameter, glabrous; sepals oblong,
obtuse, aristulate below the apex, 6.5-8 cm. long, 1.8-2.6 cm. wide;
petals subequal to the sepals; corona tuberculate; operculum depend-
ent, about 7 mm. long, the margin recurved, more or less fimbriate;
gynophore finely spreading-pubescent; ovary fusiform, glabrous;
immature fruit obovoid-oblong, about 6.5 cm. long, 2 cm. broad;
seeds oblong-ovate, 7-8 4-5 mm., alveolate.

Type in the U.S. National Herbarium, No. 1909202, collected at
Horta-Naque, Province of Loja, Ecuador, at an altitude of 3,200 m.,
on Nov. 8, 1946, by Reinaldo Espinosa (No. £965). Supplementary
sheet, US No. 1909586.

P. lozensis belongs to the subgenus Tacsonia and is closely related
to P. cumbalensis (Karst.) Harms. It differs in the laciniate-dentate,
coriaceous stipules, in the more coriaceous leaves and bracts, and
especially in the structure of the lower surface of the leaves. The
thick veins on the lower surface form a very close reticulum that leaves
very thin but deep and irregular pits between the turgid network.

170. Passiflora psilantha (Sodiro) Killip

This Ecuadorean species has been found in cultivation near Sonsén,
Antioquia, Colombia, by Gutiérrez & Barkley (No. 18C294). The
specimen is deposited in the Herbarium of the Facultad de Agronomfa,
Medellin, Colombia.

173. Passiflora mixta L.f.

On page 297 of my monograph, variations of pubescence in this
wide-spreading species are noted thus:

(1) Leaves, calyx tube, and ovary glabrous.

(2) Leaves glabrous, calyx tube and ovary pubescent.

(3) Under surface of leaves, calyx tube, and ovary pubescent.
No formal varietal names were assigned to these variants though
a fourth, P. mixta var. eriantha, was treated as a variety.

Two specimens received subsequent to the publication of my
monograph exhibit other combinations in the occurrence of indument,
and are worth noting:

(4) Under surface of leaves pubescent, calyx tube glabrous, ovary
pubescent: Ecuador: Cafiar: Tipococha, altitude 3,200 m., Penland
& Summers 1006 (US).

(5) Leaves and calyx tube glabrous, ovary pubescent: Bolivia:
La Paz: Charazani, Province of Mufiecas, altitude 2,700 m., Cardenas
3886 (US).

14 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

177. Passiflora antioquiensis Karst.

In the description given of this species in my monograph, notes
on the peduncle were unfortunately omitted. It is worthwhile also
to amend the description in certain details in view of recent collections
thus:

Leaves unlobed, up to 18 cm. long and 8.5 cm. wide, asymmetrically
2-lobed or 3-lobed to within 2 cm. of the base; peduncles very slender,
almost filiform, 17-60 cm. long, pendulous; flowers up to 14 cm. wide
when expanded.

Additional specimens examined are:

Cotomsia: Antioquia: Yarumal, Daniel 1916 (US). Cauca: El Tambo,
alt. 2,500 m., von Sneidern 1442 (S, US).

180a. Passiflora macrepoda Killip sp. nov. PLaTE 8

Caulis scandens robustus rufo-subvelutinus; stipulae cordato-ovatae
acutae inciso-fimbriatae glabrae; folia subcoriacea pilosula ad medium
trilobata basi subcordata lobis ovatis acutis serratis; bracteae liberae
ovato-lanceolatae subcordatae inciso-fimbriatae; calycis tubus tubu-
loso-campanulatus, sepalis anguste lanceolatis pilosulis apice longe
mucronatis quam tubo duplo longioribus, petalis leviter brevioribus;
corona 3 seriebus filamentis filiformibus et interiore brevi membrana
horizontali subcrenata; operculum deflexum recurvum; gynophorum
tener; ovarium velutinum.

Vine; stem terete, stout, about 5 mm. in diameter, rufo-velutinous;
stipules cordate-ovate, 2-3 cm. long, 1.5-2 em. wide, acute, strongly
oblique at the base, laciniate-fimbriate, membranous, glabrous;
petioles stout, 1.5-3 cm. long, bearing 2 or 3 short stipitate glands at
the apex and 1 or 2 just below the apex, these partially concealed in
a dense tomentum; leaf-blades 3-lobed about to middle, 10-12 cm.
along the midnerve and the lateral nerves, 16-18 cm. between the
apices of the lateral lobes (lobes about 5 cm. wide, aristate), cordulate
at base, serrulate, subcoriaceous, minutely pilosulous on the nerves
above and on the principal nerves beneath cano-puberulent or rufo-
puberulent on the secondary nerves and the veins beneath; peduncles
solitary, stout, 12-14 cm. long, densely rufo-velutinous; bracts free
to base, ovate-lanceolate, 4-5 cm. long, about 2 cm. wide, cordulate,
laciniate-fimbriate, the teeth toward the base the largest; calyx tube
slenderly campanulate, about 2 cm. long, and 1.3 cm. in diameter;
sepals narrowly lanceolate, 4 cm. long, 1 cm. wide, white within,
dorsally keeled toward the apex, the keel terminating in a filiform
awn about 8 mm. long; petals narrowly lanceolate, 3.5-4 cm. long,
white; corona in 4 series, the outer 3 ranks irregular, filamentose, the
filaments filiform, about 1.5 cm. long, close to these ranks a short,
thick, horizontal, subcrenulate membrane; operculum dependent,

KILLIP—NOTES ON AMERICAN PASSIFLORACEAE 15

about 8 mm. long, the margin recurved, short filamentose; gynophore
slender, about 3.5 cm. long; ovary ovoid, white-velutinous.

Type in the U.S. National Herbarium, No. 1803911, collected near
Unduavi, Department of La Paz, Province of Nor Yungas, Bolivia,
at an altitude of 3,000 m., in forest along roadside, on Apr. 15, 1939,
by W. J. Eyerdam (Goodspeed Expedition No. 25379). Duplicates
at University of California and Chicago Natural History Museum.

This species belongs to the subgenus Granadillastrum and because
of its laciniate-fimbriate bracts and general flower structure is evidently
nearest P. weberbaueri and P. trisecta. Both of these have trifoliate
leaves. The indument and general shape of the foliage are suggestive
of P. peduncularis.

189. Passiflora racemosa Brot.

In describing the bracts of this species I said in the monograph:
“bracts setaceous (?), soon deciduous.”” This statement is in error.
In Chase 10125, which was cited, bracts are present, and Masters’
plate in the Flora Brasiliensis shows a bract. They are ovate, about
1 cm. long and 8 mm. wide, tapering slightly at the base, thin-mem-
branous, and pink, and are borne close to the base of the flower.

P. G. Greenway has recently sent me for study a superb lot of
passionflowers from the Herbarium of the East African Agricultural
Research Station, Amani, Tanganyika, nearly all of which are from
plants grown at the Station and are of New World origin. Among
these, No. 5942, collected by Greenway, is P. racemosa. The flowers
are borne singly in the axils of fully developed leaves and are not
in a raceme or on long branches with much reduced leaves. This
plant may be of hybrid origin, or perhaps in a natural state the
flowers may sometimes be solitary.

200. Passiflora seemannii Griseb.

This species has a rather unusual range of distribution. At the
time that my monograph was prepared, the species was known in
its native state by several collections from Panama and from scattered
localities in Colombia. The following additional collections have sub-
sequently been received at the U.S. National Herbarium:

Cotomsia: Cundinamarca: San Francisco, alt. 1,550-1,580 m., Garcta-
Barriga 11051. El Meta: Villavicencio, alt. about 500 m., Killip 34445,
35692; Cuatrecasas 4673.

VENEZUELA: Amazonas: Yavita, L. Williams 14000.

209. Passiflora capparidifolia Killip

Material from British Guiana and the Rio Branco of Brazil was
cited as this species in my monograph. The species is now represented
in the U.S. National Herbarium by the following Venezuelan collec-
tions:

16 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Bolivar: Between Rfo Torono and Salto de Auraima, edge of forest along
Rio Paragua, alt. 275 m., Killip 37539. Selvas de galerfa del Rio Uari,
Gran Sabana, Tamayo 3154.

The first of these collections was in fine flower and the second one
bore young fruit. The description in the monograph may be amplified
thus:

Bracts distinct to the base, broadly ovate, up to 5 cm. long and 3 cm.
wide; sepals greenish white without, red-brown within; petals deep
pinkish red; outermost corona filaments about 1.2 cm. long, banded
with blue and white in upper part, red and white in lower part, the
second series three times longer, blue and white in the upper two-
thirds, red and white in the lower third; young fruit ovoid, about twice
as long as wide, finely pilose, obcordate, about 5 mm. long and 3.5 mm.
wide, reticulate.

210. Passiflora popenovii Killip

At the time that my monograph was prepared the only specimens
of this species available to me were from Ecuador. The species was
also collected in 1938 at El Tambo, Department of Cauca, Colombia,
alt. 1,700 m. by K. von Sneidern (No. 1444), and was deposited in the
Riksmuseet, Stockholm, and in the U.S. National Herbarium.

2lla. Passiflora brevifila Killip, sp. nov. PLATE 9

Frutex scandens omnino glaber, caule subquinqueangulato; stipulae
anguste lineares subfalcatae; folia subcoriacea integra lanceolata vel
oblongo-lanceolata acuminata basi rotundata; bracteae liberae
lanceolatae minute cuspidatae integrae basi subtruncatae, petiolo
prope apicem 4-6 glanduloso; calycis tubus campanulatus, sepalis
petalisque similibus oblongis obtusis; corona filamentis uniseriatis
liguliformibus ad apicem leviter dilatatis; operculum fimbriatum fila-
mentis rigidulis ad basim paulo incrassatis.

Woody vine, glabrous throughout; stem subquinquangular; stipules
narrowly linear, 5-6 mm. long, about 0.8 mm. wide, subfalcate,
coriaceous; petioles 1-2 cm. long, bearing 4-6 minute glands near
the apex, the glands short-stipitate or nearly sessile; leaves lanceolate
or oblong-lanceolate, 12-14 em. long, 5-5.5 cm. wide, acuminate,
rounded at base, entire, penninerved (principal lateral nerves 6 or 7
to a side, arcuate-ascending), conspicuously reticulate beneath, sub-
coriaceous, concolorous, lustrous beneath; peduncles solitary, 2.5-3
cm. long, bracts free to base, lanceolate, 5-7 cm. long, 2-3 cm. wide,
acuminate, minutely cuspidate, subtruncate at base, entire, similar in
texture to the leaves; flowers about 6.5 cm. wide; calyx tube campanu-
late; sepals and petals similar, oblong, about 2.5 cm. long and 9 mm.
wide, obtuse, apparently white; corona filaments in a single series,
narrowly liguliform, slightly thickened at apex, 1 cm. long; operculum

KILLIP—NOTES ON AMERICAN PASSIFLORACEAE 17

a fringe of rather stiff filaments about 3 mm. long, slightly thickened
at base; limen a thick ridge close to the operculum; floor of calyx tube
smooth; ovary ovoid, green; fruit globose, about 6 cm. in diameter,
the exocarp brittle, orange; seeds obovate, strongly flattened, 9-10
mm. long, 5 mm. wide, coarsely reticulate, tridentate at the apex, the
middle tooth the more prominent.

Type in the U.S. National Herbarium, No. 1643741, collected at
Vara Blanca de Sarapiqui, north slope of Central Cordillera, between
Pods and Barba Volcanoes, Costa Rica, at an altitude of 1,890 m., in
February 1938, by A. F. Skutch (No. 3591).

This species belongs to the subgenus Granadilla series Laurifoliae,
where it is the only one with a single-ranked corona. The other
representative of the series in Central America, P. ambigua, not only
has a 5-ranked corona but the ovary is densely brown-tomentose.
Doubtless the closest relative is P. nigradenia, of Bolivia, in which the
corona is 2-ranked and the ovary finely sericeous.

213a. Passiflora killipiana Cuatr., sp. nov. Priate 10

Planta breviter hirtella vel pubescens, pilis patulis rufescentibus;
stipulae lineari-subulatae; folia tenuiter subcoriacea ovato-oblonga
vel elliptico-oblonga basi subrotundata apice acute acuminata margine
remote minuteque crenulata, penninervia, nervis utrinque ferrugineo-
hirtellis vel tomentulosis, venulis reticulatis utrinque prominulis,
petiolo brevi subapicem biglanduloso; bracteae liberae obovato-
oblongae apice rotundatae sursum profunde dentatae; calycis tubus
campanulatus, sepalis oblongis subapicem corniculatis, petalis
brevioribus; corona filamentis 6-7 seriatis exterioribus liguliformibus
valde longis, interioribus mediocribus, intermediis brevibus; operculum
membranaceum profunde fimbriatum; ovarium velutino-tomentosum.

Plant hirtellous or pubescent nearly throughout, the hairs reddish,
short, spreading; stem terete, slightly striate; stipules linear-subulate,
10-13 mm. long, 1 mm. wide, ferruginous-hirtellous, soon deciduous;
petioles 7-11 mm. long, stout, ferrugineo-tomentellous, biglandular
below the apex; blades 9-16 cm. long, 4.5-7.5 em. broad, thin-sub-
coriaceous, ovate-oblong or elliptic-oblong, rounded or obtuse at base,
acutely acuminate at apex, the margin subentire, minutely and
distantly crenulate, hirtellous on the nerves above, more densely
ferruginous-hirtellous on the nerves and the surface puberulous
beneath, the midrib very prominent beneath, the secondary nerves
8-9 pairs, prominent, ascending, the smaller veins reticulate-promin-
ulous on both sides; flowers 8-9 cm. wide, solitary, axillary; peduncles
3-4 cm. long, ferruginous-tomentellous, moderately thick; bracts
thick obovate-oblong subrounded at apex, cuneate at base, coarsely
and deeply toothed on the upper half, densely ferruginous-pubescent,
4.5-5.5 em. long, 2-3 cm. broad; calyx tube cylindric-campanulate,

18 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

7-8 mm. high; sepals membranaceous, pubescent, 3.5-4 cm. long,
1-1.2 cm. wide, oblong, narrowed toward the obtuse apex, dorsally
corniculate, the horn about 5 mm. long; petals smaller, thinner,
oblong, about 3 cm. long, 0.5 cm. wide; corona filaments transversely
colored-banded, in 6-7 series, the outermost narrowly ligulate, about
3 cm. long, 0.8 mm. wide, the innermost ligulate, about 6 mm. long,
those of the intermediate series slender, 1.5-2.5 mm. long; operculum
membranaceous, horizontal, the margin broadly recurved and deeply
laciniate; limen cupular close to the gynophore; ovary ovoid-oblong,
densely velutino-tomentose; gynophore about 1.5 cm. long.

Type in the U.S. National Herbarium No. 1952839, collected in
the vicinity of La Pedrera near Rio Caquet4, Comisaria del Caqueté,
Colombia, in April 1944, by Richard Evans Schultes (No. 5875).

P. killipiana is closely related to P. laurtfolia L., but differs from
it and from the other known species of the section Laurifolia by the
rufous or ferruginous, spreading pubescence or tomentum which
covers several parts of the plant; P. killipiana differs also in the
minute crenature of the leaves and in the thick, toothed bracts.
The size of the different parts of the flowers and bracts and the
relative length of the filaments of the several rows of the corona also
distinguish this species from P. laurifolia and other related species.

The type of this species was examined by Killip in 1951 and classi-
fied as a new species in the section Laurifolia, but neither description
nor name were given. The describer takes this opportunity to
honor the man who did the largest and best work in Passiflora by
naming the new species after him.

216. Passifiora riparia Mast.

Only material from Peru and Brazil was cited in my monograph.
The following specimens from other countries are now in the U.S.
National Herbarium:

British Guiana: Western extremity of Kanuku Mountains, in drainage of
Takutu River, alt. 500 m., A. C. Smith 3157.

CotomsBia: Vaupes: Vicinity of Mitt, Allen 3340. Near Miraflores, alt.
300 m., Gutiérrez & Schultes 754 (Med). Putumayo: Mocoa, Schultes &
E. Smith 2000.

Some specimens recently seen by me have the flowers borne singly
in the leaf axils and not clustered on short, axillary branches as is
typical of the species.
221la. Passiflora pachyantha Killip, Lloydia 2:198. 1939.

This species was based upon a specimen collected by A. C. Smith
(No. 3620) on Mount Iramaikpang, in the northwestern portion of the
Kanuku Mountains, British Guiana. The species belongs to a new
section of the subgenus Granadilla, which comes between series 5,
Laurvfoliae, and series 6, Serratifoliae.

KILLIP—NOTES ON AMERICAN PASSIFLORACEAE 19

232. Passiflora incarnata L.

In the monograph I gave as the distribution of this species: ‘‘Vir-
ginia to Missouri, south to Florida and Texas; also in Bermuda.
Introduced farther north in the United States” I also listed representa-
tive specimens of each state from which I had seen specimens.

Charles C. Deam wrote me that he found P. incarnata in 1899 on a
hill near Wyandotte Cave, Crawford County, Indiana, and collected it
on a bluff of the Ohio River, a half mile above Tell City, Perry County,
Indiana. In his most useful Flora of Indiana (1940), he recorded it
from two additional counties—all these localities being in the southern
part of the State.

F. C. Gates likewise advised me that there are two specimens in the
Kansas State College Herbarium from Labette County and Cherokee
County, Kansas, collected in 1892 by A. S. Hitchcock.

No specimens were listed from these two States in my monograph.
The distribution of this species should therefore be given as Virginia to
southern Illinois and southeast Kansas, south to Florida and Texas.

245a. Passiflora danielii Killip sp. nov. PuatTe 11

Herba scandens, caule teneri glabro; folia subcoriacea ovato-
lanceolata subabrupte acuminata integra basi leviter cordata subtus
breviter adpresseque pilosa, petiolo glabro 2 paribus glandulis pracdito;
calycis tubus late campanulatus, sepalis lineari-lanceolatis, petalis
sepalis similibus obtusis; corona filamentis circa 5-seriatis in duobus
exterioribus seriebus subcapillaribus ceteris filiformibus minuteque
capitellatis; operculum membranaceum erectum ad medium fimbria-
tum; ovarium glabrum.

Herbaceous vine; stem terete, slender, striate, glabrous; stipules
semicordate, 1.5-2 cm. long, 6-9 cm. wide, abruptly cuspidate-
acuminate, attached laterally slightly above the rounded base, the
costa strongly eccentric; petioles 2.5-4 cm. long, glabrous, bearing
slightly above the middle a pair of sessile, laterally compressed, con-
cave glands about 1.5 mm. long and wide and another pair at the apex,
the latter pair sometimes being borne instead at the margin of the blade
close to the petiole; leaf-blades ovate-lanceolate, 10-12 cm. long,
6-6.5 em. wide, subabruptly acuminate, slightly cordulate at the base,
entire, 7-nerved, closely reticulate, subcoriaceous, concolorous, sub-
lustrous and glabrous above, short-pilose on the nerves and veins
beneath, the hairs divaricate, closely appressed to the surface of the
blade; peduncles solitary, terete, 8-9 cm. long, slender, glabrous,
articulate about 5 mm. below the apex; bracts unknown; calyx tube
broadly campanulate, about 6 mm. long, 8-9 mm. in diameter at the
throat, introrse at the base; sepals linear-lanceolate, 2—2.2 cm. long,
about 8 mm. wide at the base, horned dorsally just below the apex,

20 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

the horn 1.2-1.3 cm. long; petals similar to the sepals about 1.3 cm.
long and 5 mm. wide, obtuse; corona filaments apparently in about
5 series, the 2 outer filiform, almost capillary, 1.5—2 cm. long, those
of the succeeding series filiform, about 3 mm. long, minutely capitel-
late; operculum membranous, erect, about 5 mm. long, filamentose to
the middle; nectar ring a low ridge; limen tubular, 4 mm. high, closely
surrounding the base of the gynophore; ovary ovoid, glabrous.

Type in the U.S. National Herbarium, No. 1742750, collected at
Cocornd, Department of Antioquia, Colombia, in August 1938, by
Brother Daniel (No. 1536).

This species belongs to the series Simplicifoliae of the subgenus
Granadilla (species 245-258 of my monograph). The unusual petiolar
glands, which are more like the auricular appendages in P. auriculata,
at once distinguish it from the other species of this series, nearly all
of which have filiform glands. The slender peduncles, which are much
slenderer than the stem, separate it from the four species of the first
group within this series, and the long peduncles distinguish it from the
other Simplicifoliae. The large, conspicuous horns are duplicated
among species of this series only in P. praeacuta.

266. Passiflora picturata Ker

This rare species, whose distribution was recorded as “Surinam to
Para, Brazil” in my monograph, was also collected by the eminent
Colombian botanist, Lorenzo Uribe Uribe, in eastern Colombia.
The specimen in the U.S. National Herbarium bears the following
data:

Cortomsia: Meta: Acacfas, near the Rfo Acaciftas, alt. 450 m., July 29,
1946, Uribe Uribe 1334. Flowers purple.

287. Passiflora menispermifolia H.B.K.

An extreme variant of this species has been collected in Bolivia by
W. J. Eyerdam (Goodspeed Expedition 24718). It was found on
the new road to Todos Santos, 130 km. northeast of Cochabamba,
north side of Rio San Mateo, near Chimore, Province of Chaparé,
Department of Cochabamba, at an altitude of 700 m., hanging from
bushes in a damp thicket. The flowers are noted as cerulean blue.
Here also belongs Buchtien 4652, collected from Antahuacana, De-
partment of Espiritu Santo, at an altitude of 750 m., which, in the
monograph of Passifloraceae, I referred with some doubt to P. rojasii
Hassl.

In this Bolivian variant, the leaves are scarcely lobed, are more
acute, and are of a thinner texture.

KILLIP—NOTES ON AMERICAN PASSIFLORACEAE 21

309. Passiflora pittieri Mast.

This species has been known only from the type material collected
at Santo Domingo, Golfo Dulce, Costa Rica. Additional collections
in the U.S. National Herbarium may now be listed as follows:

Panama: Isla de Coiba, March 1938, Méndes 140. Forests around Pino-
gana, Darién, Pittier 6964.

Cotomsia: Antioquia: Forest at Guapd, 53 km. south of Turbo, April 25,
1945, Haught 4600. Forest near Chigorodo, 40 km. south of Turbo,
Haught 4695.

This record is one of several of species of the Costa Rican lowlands
occurring in the Darién region of Panama and extending into north-
western Colombia.

Pittier 6964 consists of a few flowers only, and I did not discuss the
collection in my monograph.

The leaves of the Méndez and Haught specimens are oblong or
oblong-lanceolate, rather than prevailingly obovate as in the type,
and are proportionately narrower. The largest one is 19 cm. long
and 6.5 cm. wide.

309a. Passiflora cardonae Killip, Brittonia 3:172. 19389.

An additional collection was made at:

VENEZUELA: Bolivar: Cerro de Auyantepui, December 1937, Cardona 284
(VEN).

348. Passiflora longiracemosa Ducke

Known heretofore only from British Guiana and northern Brazil,
this species has been collected in the forest at Salto de Parad, middle
Caura River, State of Bolivar, Venezuela, by Llewelyn Williams
(No. 11369). Though the leaves of this species are variable, I have
seen none heretofore of the shape of those recently collected. They
are broadly ovate-lanceolate, tapering gradually to a blunt tip, and
are about 15 cm. long by 9 cm. wide.

349. Passiflora securiclata Mast.

The intensive exploration that has taken place in the Orinoco Basin
in the last 10 years has led to the collection of much additional material
of this species. I cited in my monograph only six specimens from
Venezuela and none from Colombia. The numerous specimens now
available show a greater variation in the shape and texture of the
leaves. The description of the apex of the leaves as given in my mono-
graph reads, “obtuse, usually emarginate, rarely bluntly short-
acuminate.” Masters, in the original account of the species, used the
term “acute,” though some of the leaves on the type specimen are
rounded or almost truncate at the apex. Several recent specimens
have leaves which are oblong-lanceolate or even lanceolate, tapering

22 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

to an acute apex. In many of them there is a dark band about the
margin of the leaves. The texture varies from thick-coriaceous to
almost membranous. No differences are discernible in the flowers,
and it does not seem desirable to propose any segregates.

The additional material now in the U.S. National Herbarium con-
sists of the following collections:

VENEZUELA: Anzodtegui: Rfo Chive, Pittier 15023. Bolivar: Medio Rfo
Caura, Williams 11618, 11716. Rio Paragua, Cardona 746; Killip 37534.
Amazonas: Puerto Ayacucho, Williams 13125. San Antonio, Rfo
Orinoco, Williams 15069. Tamatama, Williams 15245. Esmeralda,
Williams 15310, 15515.

CotomsBia: Arauca: Matabubosa, Rio Meta, Cuatrecasas 4237. Boyaca:
Rio Guachiria, Haught 2663. Vichada: Curazao, Rfo Meta, Cuatrecasas
4134,

Two of these specimens, Pittier 15023 and Williams 15069, bear
mature fruit, which, apparently not hitherto known, may be described
thus:

Fruit ovoid, about 4.5 cm. long, and 3 cm. wide; the pericarp
glabrous, brittle, yellowish; seeds obovate, 5-6 mm. long, 4-4.5 mm,
wide, flattened, reticulate.

“Parcha”’ is given as the local name for several of the specimens.

352. Passiflora pyrrhantha Harms

Recent collections made are:
CotomBiaA: Meta: Monte de Machadero, Océa, near Villavicencio, Hermann
11120. Sierra de la Macarena, Philipson, Idrobo, & Jaramillo 2225.
Peru: Loreto: Gamitanacocha, Rfo Mazan, J. M. Schuncke 358.

353. Passiflora spinosa (Poepp. & Endl.) Mast.

Fruiting material of this species has been collected for apparently
the first time by Cuatrecasas (No. 6846) at Mitu, Vaupés, Colombia.
The fruit may be described thus:

Fruit ovoid, about 5 cm. long, and 2.5 cm. in diameter, subhex-
agonal, very minutely pilose, ochre; seeds broadly obovate, 7 mm.
long, 5 mm. wide, strongly compressed, coarsely reticulate. The
specimen is in the Herbario Nacional Colombiano.

3. Mitostemma brevifilis Gontsch,

An additional collection was made at Estacas, Campo Grande,
Mato Grosso, Brazil, on Sept. 10, 1936, by Archer & Gehrt (No.
170). This species is known otherwise only from the type material
collected on the Rio Pardo, Rio Grande do Sul (Riedel 535).

4. Dilkea acuminata Mast.

An additional collection was made by Philipson and Idrobo (No.
1820) at Central Mountains, Sierra de la Macarena, Meta, Colombia,
at an altitude of 8,000 m., on Dec. 20, 1949 (BM). ‘Climber; fruit

KILLIP—NOTES ON AMERICAN PASSIFLORACEAE

yellow” are the notations given.

23

This species is otherwise known

only from the type specimen collected at Manfos, State of Amazonas,

Brazil.

List of Exsiccatae

The numbers in parentheses refer to the species number in the text.

Allen 3340 (216).

André 3478 (69a).

Archer & Gehrt 170 (Mitostemma
brevifilis).

Balls 7360 (69a).

Buchtien 4652 (287).

Burke, David, 184 (148b).

Cardenas 3886 (173).

Cardona 284 (309a); 746 (349).

Chase 10125 (189).

Cuatrecasas 4134 (349); 4237 (349);
4673 (200); 5136 (147); 6846 (353);
9479 (148) ; 10776 (100) ; 11886 (14a);
13538 (148a); 20935 (69).

Daniel 1536 (245a); 1916 (177).

Davidson, M. E., 336 (61).

Ducke 34968 (330a).

Espinosa 617 (135); 621 (135); 806
(135); E965 (166A) ; 1314 (135).

Ewan 15899 (69a).

Eyerdam (Goodspeed)
25379 (180a).

Firmin 310 (69).

Funck & Schlim 1381 (137).

Garefa-Barriga 7757A (69a);
(69); 11051 (200).

Greenway 5942 (189).

Gutiérrez & Barkley 18C294 (170).

Gutiérrez & Schultes 754 (216).

Haught 2663 (349); 4600 (309); 4695
(309).

Hermann 11120 (352).

Humboldt & Bonpland (69).

Killip 34445 (200); 35692 (200); 37534
(349); 37539 (209).

Killip & Garcia 33482 (63).

Lehmann 675 (69); 8020 (69).

Matuda 1892 (26).

24718 (287);

77854

Méndez 108 (3809).

Metcalf (Goodspeed) 30784 (162).
Mexia 7645 (69).

Mille s.n. in 1918 (69).

Mille 222 (69).

Ochoa 1565 (69).

Penland & Summers 1006 (173).
Pérez-Arbeldez 669 (100).

Pérez-Arbel4ez & Cuatrecasas 8083
(154a).

Philipson & Idrobo 1820 (Dilkea
acuminata).

Philipson, Idrobo, & Jaramillo 2225
(352).

Pittier 6964 (309); 15023 (349).

Raimondi 7807 (155a); 8241 (155a).

Riedel 535 (Mitostemma brevifilis).

Schultes 5875 (218a).

Schultes & E. Smith 2000 (216).

Schultes & Villarreal 7417 (69a).

Schuncke 358 (352).

Skutch 3591 (211a).

Smith, A. C., 3157 (216) ; 3620 (221a).

Sneidern, von, 1442 (177); 1444 (210).

Stevens 258 (69).

Steyermark 31281 (46a); 31519 (46a);
32884 (26); 32985 (46a); 46761 (26);
52879 (10); 52932 (4); 54397 (138);
54493 (159a); 56503 (149); 57240
(111); 57277 (137).

Tamayo 3025 (10); 3154 (209).

Uribe Uribe 1061 (148a); 1334 (266).

Wedel, H. von, 2120 (61); 2421 (4).

Williams, L., 11369 (348); 11618 (349);
11716 (349) ; 138125 (349) ; 14000 (200) ;
15069 (349) ; 15245 (349); 15310 (349);
15515 (349).

©

CONTR. U.S. NAT. HERBARIUM, VOL. 35

KILLIP, PLATE 1

UNITED STATES NATIONAL MUSEUM

HERBARIO NACIONAL COLOMBIANG

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PASSIFLORA URSINA KILLIP & CUATRECASAS

CONTR. U.S. NAT. HERBARIUM, VOL. 35 KILLIP, PLATE 2

HERBARIO NACIONAL COLOMBIANO

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CONTR. U.S. NAT. HERBARIUM, VOL. 35 KILLIP, PLATE 3

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PASSIFLORA RIGIDIFOLIA KILLIP

CONTR. U.S. NAT. HERBARIUM, VOL. 35 KILLIP, PLATE 4

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CONTR. U.S. NAT. HERBARIUM, VOL. 35 KILLIP, PLATE 5

Museo de Historia Natural de la Universidad de Lima

Fundado en i918
No.

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Director Fundador: Dr. Carlos Rospigliosi Vigil
Herbario Raymondi

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CONTR. U.S. NAT HERBARIUM, VOL. 35

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54493 PLANTS OF ECUADOR

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UNITED STATES NATIONAL MUSEUM

UNIVERSITY OF CALIFORNIA

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PLANTS OF COSTA RICA

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UNITED STATES NATIONAL MUSEUM

CONTRIBUTIONS FROM THE UNITED STATES NATIONAL HERBARIUM

VoLumE 35, Part 2

A TAXONOMIC REVISION OF THE
HUMIRIACEAE

By JosE CuaTREcasas

BULLETIN OF THE UNITED StarEs NatTionaL Museum

SMITHSONIAN INSTITUTION + WASHINGTON, D.C. + 1961

Introduction . a
Historical sketch .
Drift fruit .
Fossil species . Lo
Structure of the fruit. .
Relationships and evolution
Family Humiriacae . .
Tribe Vantaneoideae
Genus Vantanea
Tribe Humirioideae .
Genus Duckesia
Genus Endopleura
Genus Hylocarpa .
Genus Humiria.
Genus Humiriastrum
Genus Schistostemon .
Genus Sacoglottis .
Collections cited
Bibliography .
Index . .

Contents

A TAXONOMIC REVISION OF THE HUMIRIACEAE

Jose CUATRECASAS

Introduction

My special interest in the tropical trees and shrubs of the family
Humiriaceae developed many years ago while I was studying my
own collections, gathered on expeditions sponsored by the regional
Government of El Valle del Cauca, from the Pacific coast of Colombia.
What drew my attention most were the rare fruit collected and
their similarity to the fossil specimens of Sacoglottis cipaconensis
presented to me some years earlier in Bogota by the geologist J. Royo
Gémez. These fossils proved to belong to the genus Vantanea rather
than to Sacoglottis. Notwithstanding the exceptionally good work
of Urban in the ‘Flora Brasiliensis,”’ the existing literature lacked
information on the structure of the fruit, information indispensable
to a more complete taxonomic understanding of the family. In
view of the collections made since Martius’ gigantic work on neo-
tropical botany, some revision of the group seemed necessary. In
1951, while in Chicago, I initiated this revision with the cooperation
of T. Just, who intended to prepare a section on paleobotany in the
planned synopsis; however, the project was discontinued. In 1957,
with the primary purpose of writing the Humiriaceae for the “Flora
of Colombia,’ I started anew with a taxonomic revision of the entire
family; the results of this study are summarized in the present
publication.

For this revision I have used the large collections of the U.S.
National Museum, in the U.S. National Herbarium (US), which have
recently been augmented by the collections formerly in the National
Arboretum. I have also used those of the:

New York Botanical Garden, New York (NY),

Gray Herbarium, Harvard University, Cambridge, Massachusetts (GH),
Arnold Arboretum, Harvard University, Cambridge, Massachusetts (A),
School of Forestry, Yale University, New Haven, Connecticut (Y).
Museum d’Histoire Naturelle, Paris (P),

Royal Botanic Gardens, Kew, Surrey (K),

British Museum (Natural History), London (BM),

Naturhistoriska Museum, Stockholm (8),

Botanische Staatssammlung, Munich (M),

Botanical Museum and Herbarium, Utrecht (U),

Museu Goeldi de Historia Natural, Belém do Pard (MG),

Instituto Agronomico do Norte, Belém do Pard (IAN), and

Instituto de Ciencias Naturales, Bogoté (COL).

26 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

The above-mentioned herbaria have the largest existing collections
of members of Humiriaceae, and their examination has resulted in a
considerably greater understanding of the family than we had pre-
viously. These herbaria combined have a copious representation of
the important historical collections made by Schomburgk, Spruce,
Martius, and others, as well as of more recent collections, among
them those of Ducke, Maguire, and the Boschwezen (Forest Service)
of Surinam. Nearly all types or isotypes of the described species
are found among the material of the herbaria mentioned. The
amount of fruiting material in the modern collections, though not
abundant, is sufficient to allow us to draw new lines in the concept
of the family and its genera. Especially rewarding have been the
amazing Brazilian collections of Adolfo Ducke. The relative impor-
tance of the collections used can be readily seen by examining the
index of collectors at the end of this paper (p. 187).

The present revision is based on the classical method of comparative
morphology. I utilize the fruit structure to a considerably greater
extent than previous authors. The paper concludes that the
Humiriaceae are to be maintained as an independent family with 8
genera, 49 species, and many subspecies, varieties, and forms.

The introduction to this paper begins with a historical sketch of
contributions to our knowledge of the family and a discussion of the
phenomena of drift fruit and fossils. There follows, because of its
taxonomic importance, a discussion of the structure of the fruit
and also a discussion of taxonomic relationships and evolutionary
trends. No data on the cytological structure of any Humiriaceae
are available. The considerations of evolutionary trends are founded
on comparative gross morphology and are merely tentative. The
paleontologic data are lmited to the mention of published species
and to a few taxonomic suggestions.

The main part of the paper is devoted to classification and taxo-
nomic descriptions of the family and the tribes, genera, species, sub-
species, varieties, and forms comprising the family. Included are
plates and figures comparing and contrasting related forms. The
plates are grouped following p. 84. The figures appear in the text
near the species to which they refer.

The citations of collections constitute an important part in a
monographic work and, because of the relative scarcity of locality data,
I consider it useful to publish the information given on the herbarium
labels by each collector. For greater accuracy these data appear in
their original language. In the citations of herbaria where specimens
are deposited, I use the abbreviations of Lanjouw’s ‘Index herbari-
orum.” In citing the photographic series of the Chicago Natural

CUATRECASAS—HUMIRIACEAE 27

History Museum, the abbreviation ‘Photo F.M.” has been used
preceding the number.

I wish to extend my thanks to the directors and curators of the
herbaria who have aided by lending me collections or giving me facilities
for my work, among them especially Albert C. Smith, Director of
the Museum of Natural History of the U.S. National Museum;
Jason R. Swallen, head curator of Botany; Lyman B. Smith, curator
of phanerogams, who has checked the manuscript; Roland Brown,
honorary curator of paleobotany, who has been of great help locating
humiriaceous fossil collections; the artist Russell Zimmermann, who
made all but four of the figures with a complete understanding of my
interpretations; and Paula Gerard and Jane Roller, who made the
accurate drawings of figure 1.

The work for this revision has been carried out at the Department
of Botany of the U.S. National Museum, with the help of a grant
from the National Science Foundation.

HISTORICAL SKETCH

1775: Aublet publishes in his ‘‘Histoire des plantes de la Guiane
francaise” the first descriptions and drawings of Humiriaceae—
Houmiri balsamifera and Vantanea guianensis—two new genera and
species named after the Caribbean names ‘“‘Houmiri” and ‘“Jouantan.”
The first genus is characterized by small flowers with 20 stamens and
the second by large flowers and many stamens. He places the genera
in the class Polyandria and the order Monogynia of Linné’s sexual
system.

1777: Scopoli, like other botanists of the time, finding the names
given by Aublet barbarous, proposes ‘‘Wernisekia” for ‘“Houmini”
(Introd. Hist. Nat.).

1789: Schreber, for the same reason, in his eighth edition of
“Tinné’s genera plantarum” changes “Houmiri” to ‘“Myrodendrum”’
and ‘‘Vantanea” to ‘‘Lemniscia” and writes: ‘‘Aubletiano vero nomen-
claturae quum nimis sit barbara, aliam quin substituerem, me continere
nequivi.” They are included in the Polyandria Monogynia.

1789: Jussieu, trying to Latinize Aublet’s name, gives it the
spelling ‘‘“Houmiria,” in his ‘Genera plantarum.”

1792: Lamark publishes a new species form French Guiana—
Vantanea parviflora.

1800: Willdenow, following Schreber, in his edition of Linnaeus’
“Species plantarum” adopts ‘“‘Lemnescia,”’ thus correcting its spelling,
and, for Vantanea guianensis, he uses ‘“Lemnescia floribunda.”

1805: St. Hilaire completes the ‘‘Latinization” initiated by Jussieu
with the spelling ‘‘“Humiria.” He includes the plant in ‘Classe

28 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Polyandrie”’ and “Ordre Monogynie” (Exposition des familles
naturelles).

1824: Nees and Martius publish the new genus Helleria, based
upon H. obovata, not noticing that it is congeneric with Vantanea
guianensis. The authors place the genus next to Humira and
consider its close relationship to several groups, primarily Aurantiae
and Ternstroemiaceae.

1827: Martius publishes (Nov. Gen. Sp. Pl. Br.) detailed descrip-
tions of Humirium crassifolium and H. floribundum, and gives a new
spelling to the generic name, ‘“Humirium” (proposed in ms. by
Richard). Martius describes the genus Helleria and a new genus,
Sacoglottis, based upon Sacoglottis amazonica, and distinguished by
10 stamens. He includes Sacoglottis in Linnaeus’ Monadelphia-
Decandria, whereas he places Humirium and FHelleria in Monadelphia-
Polyandria. Martius comments on the close relationship between
the three genera and traces their differences with the genera considered
related to Meliaceae, Symplocaceac, and Styracaceae.

1829: Jussieu gives, in St. Hilaire’s “Flora Brasiliae meridionalis,”’
for the first time a taxonomic category (order=family) to these three
genera under the name ‘‘Humiriaccac.” He adds the new species
Humirium montanum, H. parvifolium, and Helleria ovalifolia and
lists Helleria obovata Nees. & Mart.

1880: Lindley (Introd. Nat. Syst. Bot.), considers the tribe
Humiriaceae and their relation to the Aurantiaceae, Diosmeae, and
Rutaceae.

1840: Endlicher in “Genera plantarum” lists this family as “Ordo
CCXXII-Humiriaceae,” following Jussieu and recalling Martius’ con-
siderations about the close relationship between the three genera with
Symplocaceae and Styracaceae, and their separation from the Melaceae.

1842: Casaretto publishes Humirium dentatum.

1843: Bentham publishes four new species of Humirium from the
Guianas—H. densiflorum, H. guianense, H. obovatum, and H. sub-
crenatum.

1848: Planchon, in his studies ‘Sur la famille des Linées,”’ makes
comparisons between Humiriumand Erythroxylon, and the Linaceae, and
shows their close relationship; nevertheless, he considers the Hum-
iriaceae independent but connected with the Linaceae through
Roucheria. The main differential characters given are the thick
connective of the anthers, the intrastaminal free disk, and the single
style. Planchon’s transverse division of the ovary cavities by false
septa are only true of Humiria.

1850: Miquel publishes Humirium surinamensis.

1858: Bentham publishes ‘‘“Notes on Humiriaceae,’”’ which is the
first synthetic account of this family, and which defines with precision

CUATRECASAS-—HUMIRIACEAE 29

and simplicity the “order” and its subdivisions. He identifies Helleria
of Martius with Vantanea; supposedly Martius failed to recognize
the latter genus because of inaccuracies in Aublet’s drawings, which
overlooked the enlarged connective of the anthers. Bentham dis-
tinguishes three genera by the number of stamens: Vantanea with
numerous stamens (75-150); Humirium with 20 stamens, and Saco-
glottis with 10 stamens. He does not consider the structure of the
thecae of the anthers, but uses the number of ovules in the definition
of the genera: Vantanea with two ovules in each cavity of the ovary;
Sacoglottis with one; Humirium with both cases being true. Bentham
lists 4 species of Vantanea (guianensis, minor sp. n., obovata, and
ovalifolia), 12 species of Humirium (crassifolium, guianense, flori-
bundum, montanum, obovatum, cuspidatum sp. n., suberenatum, denta-
tum, balsamiferum, macrophyllum sp. n., densiflorum, and oblongi-
folium sp. n.), and 2 species of Sacoglottis (amazonica and guianensis
sp. n.).

1860: Baillon, in Adansonia, discusses the family with the descrip-
tions of three examples—Sacoglottis amazonica, Humirium arenarium,
and Helleria—that he considers different from the Ericaceae and
Styracaceae.

1862: Baillon describes a new genus based on a West African
species, Aubrya gabonensis. He divides the Humiriaceae into two
series: One with “free stamens” (Vantanea and Aubrya), and the other
with monadelphous stamens. He distinguishes Sacoglottis with 10
stamens and uniovulate ovary cells from Humirium with 20 stamens
and biovulate ovary cells.

1862: Bentham and Hooker define “Ordo XX XV Humiriaceae”
mainly by small sepals, disk, stamens 10-, carnose connective, and
solitary ovules or rarely 2 in each cell. They distinguish 3 genera
according to the number of stamens and disk: Vantanea, numerous
stamens; Humiria, 20 stamens monanthcriferous and sometimes 5
stamens triantheriferous; and Sacoglottis, 10 stamens. Bentham con-
siders this order closely related to the Linaceae series [xonantheae
but different from it because of the enlarged connective of the anthers.

1870: Schnizlein publishes Humirium compactum.

1873: Baillon, in Adansonia, sees no reason for replacing Aublet’s
name “Houmiri.” He compares this form with the Erythroxylaceae,
Hebepetalum, and Ixonanthes (Linaceae) and sees close affinities.
Vantanea parviflora, which Richard (ms.) considered apart (Vantane-
oides), has only strongly contorted corolla and imbricate calyx, features
insufficient to separate it from Vantanea (V. guianensis, with tubulose
calyx). Nor does he see a difference between Helleria and Vantanea.
He writes that the ovary cells are alternipetalous as in ugonia, Hebe-
petalum, and Izonanthes, and that the ovules are pendulous with

30 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

the micropyle pointing upward, almost collateral or superposed, as
in Izonanthes and Erythrozylon. He considers only 1 genus in Hu-
miriaceae, Houmiri, with 15 species distributed in 5 sections: Houmiri
properly (Humirium), Aubrya, Saccoglottis, Vantaneoides, and Van-
tanea. All are included in the family Linaceae, series Houmirieae
Baillon.

1874: Baillon, in “Histoire des plantes,” describes as examples
three types of Humiriaceae and reiterates his viewpoints already men-
tioned in Adansonia. He divides the genus Houmiri into sections:
Sect. Aubrya with 10 free stamens; Sect. Saccoglottis with 10 stamens
and 10 staminodia; Sect. Humirium with 20 stamens; Sect. Vantanea
with 20-30-60 stamens; and Sect. Vantanecides never formally pub-
lished. They form the series Houmiriées in the Linaceae.

1877: Urban publishes a thorough treatment of the family in the
“Flora Brasiliensis” of Martius. His detailed morphologic investi-
gations on the structure of the thecae of the anthers, number of ovules
in the ovary cells, position of the carpels, and number and structure
of stamens are the greatest contribution to the systematics of the
Humiriaceae. Urban finds in these characters the basis for
a new concept of classification. He divides the family into two
primary groups according to the number of cells in the thecae of the
anthers. Thecae with two loculi and many stamens define Vantanea;
unilocular thecae and fewer stamens (10-20) define Humiria and
Sacoglottis. Urban distinguishes Humiria for having biovulate carpels
opposite the petals from Sacoglottis, which has uniovulate carpels
opposite the sepals. He further finds that the sacs of the anthers are
glabrous in Sacoglottis and pilose in Humiria. Contrary to previous
authors (Planchon, Baillon, and Bentham), Urban finds that the
number of stamens is not the differential character between Humiria
and Sacoglottis; Humiria has 20 stamens, but Sacoglottis also includes
species with 20 stamens. Going further in the systematization and
considering the number of stamens and anthers also important, Urban
divides Sacoglottis into three subgenera: I, Humiriastrum, with 20
undivided stamens, 3 species; II, Schistostemon, with 20 stamens, the
larger 5 tridentate bearing 3 anthers, 4 species; III, Eusacoglottis,
with 10 stamens, 3 species. Urban’s system proves to be definitive
because later discoveries and studies reinforce the concepts and taxo-
nomic trends devised by him. All later botanists follow his treatment.
Unfortunately, Urban sees very few fruit and does not use them in his
system. All 19 species that Urban treats are Guianan-Brazilian.
Several new combinations, varieties, and one new species (Sacoglottis
dichotoma) are described in the “Flora Brasiliensis.”’

1878-1893: Urban adds three new species of Sacoglottis and describes
the fruit of a Sacoglottis sent to him by Schwacke and Glaziou. In

CUATRECASAS—HUMIRIACEAE 31

these globose, large fruit with hard endocarp filled with resinous cav-
ities, Urban sees a new difference between Sacoglottis and Humiria,
which confirms his separating the two genera on a basis other than
the number of stamens. In this connection Urban writes the inter-
esting following paragraph: ‘‘The fruit of Humiriaceae are very seldom
in botanical museums. For the preparation of my monograph of the
family, in 1877, I could only obtain fruit from Humiria floribunda
and Sacoglottis oblongifolia. Afterward, I have seen fruit of a Saco-
glottis species collected by Poeppig in the Amazonas, of a Humiria
(or Vantanea?) from the same place (almost the size of a hen’s
egg), and another of a Sacoglottis species found floating on the shores
of Jamaica (see Morris, in Nature, 1889, p. 322). Since no herbarium
specimens of the fruit mentioned were available, it was not possible
to identify the species. The identification of the genera could be
solved only through the analogous structure of the fruit with those
of Humiria floribunda or Sacoglottis oblongifolia. Through the exam-
ination of the fruit it is very satisfactory to see confirmed my view-
point on the separation of Humiria and Sacoglottis. The fruit were
sent to me in dried condition by Schwacke and preserved in alcohol
by Glaziou. My separation between Humiria and Sacoglottis was es-
tablished on the basis of the position of the ovary cells and number
of ovules (in contrast to the formerly prevailing criterion based on
the number of stamens), so that the differences between the two
genera, as I saw them, harmonize with differences in the structure
of the fruit. The fruit of Vantanea, which up to now I have not
seen, could possibly indicate that this genus is more closely related
to Humiria than to Sacoglottis’ (Beibl. Bot. Jahrb. 34:3, 1892).

1890: Reiche, in summarizing the family for the Engler and
Prantl ‘‘Pflanzenfamilien,’”’ follows Urban, the only difference being
the reduction of the three subgenera of Sacoglottis to sections.

1898-1910: Huber describes three new species based on new col-
lections made by either him or Ducke: Sacoglottis uchi, S. duckei, and
Vantanea cupularis.

1921: Hallier, in “Beitrige zur Kenntnis der Linaceae,” follows
Baillon in joining the Humiriaceae with the Linaceae and supports
this viewpoint with much data and long considerations. The main
difference of Humiriaceae from closely related families, such as
Linaceae, is the presence of an intrastaminal free disk, which is usually
lacking in Linaceae. But, he says, Izonanthes also contains an intra-
staminal disk (according to Pierre in I. cochinchinensis).

1922-1938: Ducke, in various papers on Amazon plants, pub-
lishes four new species of Vantanea, five new species of Sacoglottis,
several new varieties, and numerous morphologic, ecologic and geo-
graphic observations about Humiriaceae. In 1938, he publishes

32 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

synopses in key form of the Amazonian Sacoglottis and Vantanea.
Ducke’s studies are mainly based on his field experience and large
collections, and constitute the largest contribution to our knowledge
of the Amazonian flora in this century. Ducke’s most outstanding
discoveries in HZumiriaceae are two species—S. verrucosa and S.
heterocarpa—which in the present treatment are credited the rank of
new genera.

1922: Berry gives the first description and illustration of a fossil
humiriaceous species based on fruit, Sacoglottis tertiaria, from the Pli-
ocene horizon in Bolivia.

1924: Berry publishes the first description and illustration of a
fossil Vantanea, based on well preserved endocarps of the late Eocene
or Oligocene of Cipacén, found at 8,185 ft. elevation in Colombia.
The endocarps are described as having from five to seven seeds, but
what Berry believes to be the seeds (“The stone has imbedded in it
from five to seven large seeds”) are in fact the valves of the endocarps.
This species is named Saccoglottis cipaconensis. Other specimens
from the Lobitos formation of the Upper Eocene in Peru are attributed
to the same species. Another fossil fruit, actually Sacoglottis, from
the same Cipacén Oligocene formation, is described as Vantanea
colombiana.

1928: Malme publishes Sacoglottis mattogrossensis.

1929: Berry in several papers publishes new data on humiriaceous
fossils and describes Saccoglottis cipaconensis var. peruvianus of the
early Tertiary from Belen, Peru, Vantanea compressiformis n. sp.
from the same locality, and Vantanea Sheppardi n. sp. of the Eocene
from Ida Seca in Ecuador.

1931: Gleason, in his account of plants of his and other expedi-
tions in Guiana, publishes one species (H. savannarum) and one
variety (HZ. floribunda var. spathulata).

1931: Winkler publishes a new treatment in the second edition
of the “Pflanzenfamilien.” He follows Baillon and Hallier in con-
sidering this group in the Linaceac, but decides to form with it a
subfamily, ‘“Unterfamilie Humirioideae.” This subfamily is char-
acterized by the intrastaminal disk, the staminal tube, and the thick
connective of the anthers. In the treatment he follows Urban’s
system without making any further contributions to our knowledge.

1933: Reid, in describing Sacoglottis costata, a fossil species of the
Tertiary in Colombia, emphasizes the structure of Sacoglottis fruit
and its peculiar dehiscence.

1933: Hill, in an interesting article dedicated to the study of
different kinds of germinal dehiscence, gives the first existing infor-
mation on how the endocarps of Sacoglottis amazonica and S. gabonensis
are dehiscent at germination: “The devices exhibited by the five-locular

CUATRECASAS—HUMIRIACEAE 33

fruits of Sacoglottis and Aubrya are of similar character, except that
in both the valves are of a more spongy nature and ovate in outline,
rounded at one end and somewhat pointed at the other, deeply hol-
lowed out on the insides to allow sufficient space for the embryos
lying within close to the centre of the fruit” (p. 881).

1984: Macebride describes Vantanea peruviana from Peru.

1935: Siissenguth publishes Humiria cassiquiart from Venezuela.

1935: The Sixth International Botanical Congress in Amsterdam
approves of the inclusion of ‘“Humiria St. Hil.” (1805), as against ‘“Hou-
miri Aubl.,” in the “(Nomina generica conservanda.” ‘The proposal
by Lanjouw and Sprague reads: “The correct name for the type
genus is Houmiri Aubl. (1775). The Latinized form Humiria St. Hil.
(1805) is so widely employed, however, that it seems desirable to
conserve it. Unless this is done, the spelling of the family name will
have to be altered.”

1988: Weyland publishes a morphologic study of fossil endocarps
of Vantanea and describes a new species, Sacoglottis germanica, which,
according to Kirchheimer, is not a Humiriaceae.

1940: Bakhuizen van der Bilt publishes Sacoglottis kaboeriensis
from Surinam.

1943: Stanley publishes Vantanea barbourei, based on an inter-
esting discovery of Barbour that extends the known range of the family
northward to Costa Rica.

1945: WDucke describes another Amazonian species, Sacoglottis
ceratocarpa.

1945: Selling makes the first detailed description of a fossil endocarp
of Humiria after studying its structure and germinal dehiscence in
longitudinal, lingulate valves. He describes the new species Zumaria
bahiensis of the Miocene or Pliocene from the State of Bahia, Brazil,
in which specimens show the imprint of two superimposed seeds in a
fertile cell. The external picture of the fossil, the cross-section, and
the lack of subapical holes are very much suggestive to me of the
genus Vantanea, but the small size of the endocarp and the two seed
impressions incline me to think that Selling is right in considering it
a Humiria. Selling’s work is well illustrated and also gives a picture
of Sacoglottis cipaconensis Berry and its variety peruviana Berry. He
discusses this species’ generic status and transfers it to Humiria, and
sees a variety as a species. He believes that the genus is of consid-
erable age and that its distribution in the Tertiary was even wider
than at the time of the publication of his paper.

1948: Little publishes Humiria procera, based on his collections
from the west coast of Ecuador.

1950: Cuatrecasas publishes Vantanea occidentalis, Sacoglottis
diguensis and its variety, S. melanocarpa, and S. ovicarpa—all of which

34 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

are based upon his own collections in the western rainforests of Colom-
bia. These and Little’s data extend the known range of the family to
the Pacific side of the Andes.

1950: Frées publishes Sacoglottis villosa in Brazil.

1951: Kirchheimer attributes another fossil species to the Humi-
riaceae, Sacoglottis kaysert (Schindehutte) Kirchheimer, which he
transferred from Actinostrobites. This finding is especially important
because the fossil specimens came from Eichelkopf, Homberg Kassel
in Germany, this locality signifying a larger area and greater antiq-
quity in the original distribution of the family. He presents mor-
phological descriptions of several known fossils of Humiriaceae and
critical and comparative remarks on the three known genera, which
he tries to distinguish; but the distinguishing characters used are
mainly the existence or absence of resinous cysts, the presence of one
or two seeds in each fruit-cell, and the marks or vestiges of vascular
bundles. No light is really shed on the actual morphology that
distinguishes the endocarps of the three genera. I have serious
doubts, because of the illustrations, that the newly transferred
Sacoglottis kayseri belongs to the Humiriaceae.

1952: Steyermark publishes Humiria pilosa from Venezuela.

1956: Cuatrecasas publishes Vantanea magdalenensis and a
variety of Sacoglottis from the Magdalena Valley in central Colombia.

Drirt Frurt

Although Aublet’s publication in 1775 constitutes the first con-
tribution to the knowledge of humiriaceous plants, unidentified
fruit of one species of this family had already been known for about
two centuries. These fruit were the woody, ellipsoid endocarps of
Sacoglottis amazonica, which, filled with empty resinous cavities, had
drifted on the ocean currents to the shores of the West Indies and
other islands. Found as far away as the Azores and the British Isles,
these endocarps had puzzled botanists for almost three centuries.
The first written notice of them was published in 1605 by Clusius in
his “Exoticorum libri decem’’ (lib. 2, cap. 4), in which appeared a
coarse drawing of the endocarp and a masterpiece of description.
Clusius’ data were quoted or reproduced by subsequent botanists,
such as J. Johnston, J. Bauhin, and H. Sloane. Sloane in 1696 men-
tioned the abundance of the endocarps found in Jamaica. According
to Morris, the specimens collected by Sloane and deposited in the
British Museum were recognized by Baker in 1889. In 1884 Morris,
at the Palisadoes in Jamaica, collected among other drift fruit those of
Sacoglottis, which he sent to Kew for determination.

In 1887 Kew received another specimen of the same fruit collected
on the shore of Bigborough Bay in southern England. Hillier, assist-

CUATRECASAS—HUMIRIACEAE 35

ant at Kew, noticed the great similarity between the drift fruit and
specimens of Sacoglottis gabonensis, collected by Mann in West Africa;
the latter, however, were smaller and more rounded than those of
Jamaica and England. Oliver, however, thought that they belonged
to Humiria, a genus widespread in South America; he wrote: “Until
we obtain fruits of H. balsamifera we are unable to say whether it is
that species or not.” In order to attract the attention of people able
to help in identifying the Jamaican drift fruit, Morris in 1889 pub-
lished in ‘Nature’ (pp. 322-323) an article with drawings.

This first article of Morris roused widespread interest and resulted
in the discovery of the parental plant of the mysterious fruit. In 1892,
Martius identified it as a species of Sacoglottis. On several islands
and in different countries, people collected and reported the fruit
(from Barbados, Dominica, Mustique, St. Vincent, Grenada). The
most important fruit was found in Trinidad, where J. H. Hart, Super-
intendent of the Botanic Gardens, identified Morris’ fruit with
accurate drawings, by Herman Crueger, a former botanist in Trinidad,
of fruit and flowering specimens of a Trinidad tree.

According to Morris (1895, p.65), Hart wrote in March 1889 as follows:
“T am extremely pleased that you called my attention to the Jamaica
drift-fruit. JI remember the specimens well, and cut several of them
in Jamaica at the time we were packing the set you sent to Kew for
the Botany of the Challenger Expedition. As soon as I read your
article in Nature, I commenced a search among the material in the
herbarium here, and found a drawing of Sacoglottis, by Crueger, with
dissections of the flower and fruit. These made it evident that the
plant which produces the unknown fruit is a native of Trinidad.
Feeling further interested in the matter, I communicated with Mr.
Syl. Devenish, the friend and companion of Crueger on many of his
excursions, and I learnt further particulars as follows. When travel-
ling in the forest at Irois, in the southeastern part of the island, they
found on the beach specimens of the fruit in question. Following up
the stream they came to the tree producing it, from which, I presume,
the drawings were taken. In addition, Mr. Devenish gave me a fruit,
which I now send, to show there can be no mistake in the matter.
This was collected by himself on the spot, so that there can be no
doubt of the identity of the species we are both discussing. Mr.
Devenish states that the tree is very rare. He saw but two in all his
travels through the island. It is known locally as Cojén de Burro. It
is probable that a greater portion of the drift-fruits found in Jamaica
and elsewhere are produced on the mainland of South America, and
are brought down by the flood waters of the Orinoco and the Amazon.”

Oliver at Kew solved the problem when, receiving Crueger’s draw-
ings, he was able to identify them as Sacoglottis amazonica. Morris’

36 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

second article in Nature (1895) is a complete and interesting narration
of the earlier and recent history of the Jamaica drift fruit.

In his book ‘Plants, seeds, and currents,” H. B. Guppy (1917) gives
an account of his own investigations, conducted independently from
those of Morris, on the Jamaica drift fruit. Guppy found endocarps
of Sacoglottis amazonica on the beaches of Colon, Jamaica, and on
Turks Islands, Tobago, and Trinidad. In Trinidad he made the
acquaintance of the botanist Hart, who showed him the parental plant
species of the then mysterious fruit S. amazonica growing in a restricted
area of the island. Guppy’s book summarizes the history of the drift
fruit, and points out that Sloane first recognized this fruit after it
had been thrown up on the shores of the northwest islands of Seotland.
Guppy emphasized the suitability of Sacoglottis endocarps for drifting
great distances, and studied possible courses between the Amazon and
Orinoco estuaries and the West Indies and Europe via the South
Equatorial Current and the Gulf Stream. Stating that Sacoglottis
amazonica was the only species dispersed by the currents, he pointed
out that the currents “would readily transport the fruits in a sound
and effective conditon from tropical West Africa to Brazil, but not
from the tropics of the New World to West Africa.”

Ridley in his book ‘The dispersal of plants” (1930, pp. 203-204)
excluded the Humiriaceae from the sea-dispersed plants “as it has
failed entirely to cross the sca and establish itself anywhere in Trini-
dad.” But I must say that I cannot agree with Ridley’s point of
view and still less with that of Guppy. In my opinion, the Humiriaceae
are a tropical American indigenous family widespread and rich in forms,
whose center of origin was the somewhat elevated lands surrounding
the Amazon Basin before the uplifting of the Andes. It is quite clear
to me that the single African Sacoglottis species, which is very closely
related to S. amazonica, is an offspring of the Amazon stock; its
ancestor at some time in the Tertiary may have found its way along
the Brazilian current to establish itself on the West African coast.
Furthermore, evidence has established the Amazon and Orinoco
origin of the Sacoglottis drift endocarps found in the West Indies and
British Isles, but no examples of Sacoglottis gabonensis have been found
on American shores,

The structure of many humiriaceous fruit provides them with
empty resinous cysts that make them buoyant, a fact that explains
why they drift easily and are often found on the shores of rivers and
seas. On the Pacific coast, these fruit were collected and reported by
I. M. Johnston at San José Island; they were probably brought there
from the western coast of Colombia or from western Costa Rica by
the local drifts originated by the Equatorial Counter Current (see

CUATRECASAS—HUMIRIACEAE 37

Sacoglottis ovicarpa, p. 168). Very often fruit or endocarps of Van-
tanea, Sacoglottis, and Schistostemon have been found in drifts and
drift deposits of the Amazon and Magdalena Rivers. The abundance
of Vantanea endocarps in fossil beds proves the drift origin of these
deposits and therefore the capacity of currents to transport the Van-
tanea fruits (which lack resinous cavities), especially the mountain
rivers with speedy waters.

FossiLs

About 12 or 13 fossil taxa attributed to the Humiriaceae have
been described (see the historical summary above). Except for two
described from European specimens, they are from tropical South
America—Colombia, Peru, Brazil, and Bolivia—where they were
found in Tertiary layers. The most conspicuous species belong to
Vantanea, others belong to Sacoglottis. Much morphologic work on
this family has been done by paleontologists and much literature
exists, but the genera are not well understood, and many of the
published species probably will need transferring. ‘The fossil species
published are:

Vantanea colombiana Berry from Cipacén, Colombia (Oligocene)

V. compressiformis Berry from Belén, Peru (late Eocene)

V. sheppardii Berry from Ecuador (Eocene)

Sacoglottis tertiaria Berry from Bolivia (Pliocene)

S. cipaconensis Berry (Humiria cipaconensis (Berry) Selling) from Cipacén,
Colombia (Eocene or Oligocene)

S. cipaconensis peruviana Berry (Humiria peruviana (Berry) Selling) from
Belén, Peru (early Tertiary)

S. costata Reid from Colombia (Tertiary), probably belonging to Humiri-
astrum

Humiria bahiensis Selling from Brazil (Miocene or Pliocene).

Vantanea wilcoriana Berry, which is based on leaves, has been
recently identified by Roland Brown as Diospyros sp. According to
Kirchheimer, Sacoglottis germanica Weyland is identical to Sypon-
dylostrobus smythit Mueller. On the other hand, I think that Saco-
glottis kaysert Kirchheimer (Actinostrobites kaysert Schindchitte)
does not belong to the Humiriaceae. There is no proof at present
that the Humiriaceae in Tertiary times spread throughout Europe
and Africa; the documentation of living and fossile material supports
Berry’s belief that the Humiriaceae are an old American stock. This
stock evolved and diversified inside the American continent into
eight genera and many species widely spread throughout tropical
American areas. The only West African species, Sacoglottis gabonensis,
probably originated from drift fruit brought to Africa from the
Amazon regions by ocean currents in earlier Tertiary times.

513359—61——2

38 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

A revision of the fossil Humiriaceae is necessary. At present
Father Gustavo Huertas from Bogotdé (Instituto de Ciencias Natu-
rales) is working on this revision.

STRUCTURE OF THE FRUIT

One of the outstanding features of the Humiriaceae is the structure
of the fruit. The fruit is drupaceous with a single stone derived
from a syncarpous ovary with biovular cells as in Vantanea and
Humiria, or uniovular as in all other genera. The exocarp, which
is the whole fleshy layer outside the endocarp, is more or less soft or
hard; thick or thin; easily removable at maturity or very adnate to
the stone; and hard, granulose, or coriaceous when dry. Usually the
surface is smooth but sometimes it is rugate or tuberculate. The
endocarp is very hard, woody, with thick walls. Rarely, it has five
(also four, six, or seven) seminal cavities, because generally only one
or two of the ovary cavities are fertile and developed in the fruit.
The cavities are normally monospermous in genera with uniovular
ovary cells.

According to my observations on endocarps of different. species,
in Vantanea only one of the two ovules develops into a seed, the
fertile cavities being occupied by a single, long, oblong seed. The
few notes on Vantanea with bispermous loculi that I found in the
literature (Selling, 1945, p. 261), are indirect references and not the
result of observations. In Humiria the endocarp develops a complete
cavity for each fertile ovule. When both twin ovules are fertile, we
can see two superposed fertile cavities. Often these two cavities are
marked outside the endocarp by the curved edges of the valves
pinched at the middle. More commonly, only one of the twin ovules
and its cavity are developed, the lower one of a carpel alternating
with the upper one of the next. The sterile cavities are empty,
usually reduced, and sometimes connected with the subapical foramina
of the endocarp. Usually not more than four or five ovules are found
in a Humaria endocarp, more often only two or three, and those of
the lower layer. In mature endocarps frequently only fertile cavities
are present. The sterile ovary cells become filled with woody wall
tissue. Only in Humiria are some vacuous sterile loculi often found.

The most interesting peculiarity of humiriaceous fruit is the type
of germinal dehiscence of the endocarps. No references to experi-
ments on seed germination in this group exist, but the study of the
morphology of the endocarps makes me assume that the single
or the few seeds borne germinate without being liberated. The woody
endocarp serves as protection to the seed and has a special device to
facilitate the liberation of the embryo at germination.

CUATRECASAS—HUMIRIACEAE 39

In Vantanea, Duckesia, and Humiria the endocarp shows conspic-
uous longitudinal lines, or shallow furrows, which mark potential
valves stretching from near the apex to above the base. At full
maturity, these valves can be removed, like a window shutter or
fenestra, from the well-ridged top to the base, continuous with the
wall tissue of the endocarp. In Humiria it is more difficult to take
the valves off artificially. In general sterile valves are more difficult
or impossible to remove. At the time of germination, the valves
are easily pushed away by the pressure of the developing embryo.

In the above-mentioned genera, the valves have an oblong or
tongue shape and have approximately the same width as the alter-
nating costae that separate them. The thick septa of the endocarp
form what, for practical purposes, can be called the costae or ribs
framing the valves. These costae are more or less at the same level
of the valves in the above cited genera. In Hylocarpa the costae
are very prominent and robust, and separate longitudinal grooves.
At the bottom of the grooves the narrower, but also prominent, valves
may be seen. In Endopleura the prominent and winged ribs com-
prise five in the lower part; they fork and form 10 woody ribs or
wings at the 2 upper thirds. At the bottom of the deep furrows,
between the twin costae, lie the narrow, thin, and conspicuous valves.
In Humiriastrum the germinal valves are reduced in size to the upper
half of the endocarp.

On the other hand, the endocarps of Sacoglottis and Schistostemon
lack any conspicuous opening device when inspected superficially.
The woody septa and walls are usually filled with large globose,
resinous cavities, which commonly mark an irregularly bullate
surface. On this surface 5 or 10 longitudinal shallow furrows can
barely be seen. Only when the endocarps are very much washed and
eroded are the 10 thin furrows conspicuous. At late maturity, it is
possible, though not always easy, to remove the broad, thick valves
on the fertile spots from the rest of the endocarp. These valves are
oblong-elliptical, and they cover the space almost from the apex to the
bottom. Laterally they are almost adjacent. The intermediate
costae are thin, and the septa are almost membranaceous (fig. 1).

The endocarps of Vantanea, Humiria, Hylocarpa, Endopleura, and
most species of Humiriastrum are compactly woody, whereas the
other genera have many cysts in the woody tissue of the septa, costae,
and valves, which in cross section appear to have a spongy appearance.
These cavities are secretory sacs, usually empty or with some resinous
powder. They provide the fruit with a very buoyant property, for
which reason Sacoglottis may float and travel for years on rivers and the
sea. The larger development of resinous cysts is parallel to the disap-
pearance of fenestra-like valves and the formation of broad, almost

40 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

frameless valves. In Humiriastrum, most species lack resinous cysts,
and only in a few of them are small resinous cavities conspicuous.

Another feature with a possible connection to the embryo liberation
is the presence in some genera of small germinal holes at the apex of
the endocarp. These holes (foramina) are located on the ribs, near
the top. Sometimes they are very deep and may communicate with
the seminal cavities. Subapical foramina are especially developed in
Humiria and Humiriastrum. They are also present in Duckesia and
Endopleura. |

This type of germinal dehiscence of the endocarps is rare in the plant
kingdom. The only similar cases that I have found are those of
Davidia (Cornaceae) and Tectonia grandis (Verbenaceae). The
Davidia type is the only very close one to Vantanea, the former having
been well described and illustrated by Hill (1933, p. 884, fig. 12), who

Ficure 1.—a-c, Vantanea occidentalis, X 144 (Patifio 12): a, endocarp with uplifted valve;
b, endocarp with valve removed uncovering the seed; c, valve from the inside. d-f,
Sacoglottis amazonica, X 1 (Archer 7964): d, endocarp, showing the dehiscence line;
e, endocarp with uplifted valve, uncovering the seed; f, valve from the inside.

CUATRECASAS—~HUMIRIACEAE Al

was the first botanist to describe the dehiscence of Sacoglottis amazonica
and S. gabonensis (p. 881).

RELATIONSHIPS AND EVOLUTION

The basic features of the Humiriaceae as a whole are: Epitropous
ovules with micropyle facing upward; free and thick petals; stamens
in two or more whorls, united in tube at base; anthers with thick,
fleshy, usually elongated connective; intrastaminal free disk sur-
rounding superior ovary; ovary cells 1-2 ovulate; drupaceous fruit
with woody, thick endocarp of unique structure with germinal
dehiscence and reduction of numbers of seeds to 2 or 1; woody plants
with alternate, simple, coriaceous leaves.

These characters place the family in the order Geraniales, suborder
Geraniineae of Engler. Undoubtedly, as first suggested by Planchon,
the major affinities of the Humiriaceae are found in the woody mem-
bers of the Linaceae and the Erythroxylaceae, but not to such an
extent as to justify their inclusion in the Linaceae, an inclusion
made by Baillon, Hallier, and Winkler. The Humiriaceae form a
very natural, homogeneous, and compact group, well-defined and
perfectly separable from the Linaceae. The morphology of the fruit,
which has shed much light on the definition of the genera in the
present work, also shows a basic structure common to the entire family.
The endocarps of the Humiriaceae are very different from those
of the drupaceous species of the Linaceae. The Linaceae genera
closest to the Humiriaceaze are Izonanthes, Ochthocosmus, and Ctenolo-
phon in having a floral disk, but they differ in that the disk is intra-
staminal and completely free in Humiriaceaec, whereas the stamens
are united to the disk (inside or outside) in the three linaceous genera.
Furthermore, the flower is perigynous and the fruit a septicidal
capsule in Izonanthes and Ochthocosmus; the disk is extrastaminal,
the ovary bicarpelar, and the style divided in Ctenolophon. In all
three the anther never has the fleshy, thick connective, typical of
the Humiriaceae. In linaceous genera with drupaceous fruit (as
in Hebepetalum and Hugonia), the endocarp is formed by several
free pyrena.

The anatomical structure of the Humiriaceae also bespeaks the
family rank, as is stated by Heimsch: “Thus, structurally the Humiri-
aceae are a homogeneous group as evidenced by diffuse porous wood
without growth rings; round chiefly solitary vessels with scalariform
perforations; heterogenous IT A rays; and diffuse parenchyma which
generally shows transitions to the abaxial paratracheal type. For
this reason the family is described separately rather than as a tribe
of the Linaceac as Hallier (33, 34) and Winkler (27) have done.
Nevertheless on the basis of the structure of the secondary xylem,

42 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

the affinities of the Humiriaceae are more with the Linaceae rather
than with any other family” (1942, pp. 96-97, figs. 2-4).

Likewise, Record and Hess write: “In Winkler classification the
Humiriaceae are reduced to the status of a subfamily of the Linaceae,
but this proposal has little support in the anatomy of the woods”
(1943, p. 191). The morphology of the pollen grains differs from that
of the general types in the Linaceae. According to Erdtman (1952),
the pollen grains in the Humiriaceae are slightly similar to those of
certain Celastraceae. The presence of balsamic oils and resins in
the bark and fruit of Humiriaceae also distinguishes them from the
Linaceae. Hutchinson (1926, p. 196) not only considers the Humiri-
aceae an independent family, but even places them in the order Mal-
pighiales, in company with Malpighiaceae and Erythroxylaceae; in
this placement I concur. In the general treatises, noncritical authors
usually follow standard classifications such as Engler’s ‘‘Pflanzenfami-
lien.” Recent floral works dealing with Humiriaceae treat the group
as a family, as do Bakhuizen van den Brink and Pulle in the ‘Flora
of Surinam,” and Exell and Mendoga in the “Flora of Angola.”

Concerning relationships among the Humiriaceae, one must con-
sider the genus Vantanea, with an indefinite number of stamens in
several whorls, as the most ancient of the family. Some other older
characters are the biovular cells, the bilocular anther thecae, and the
narrow, shutterlike valves of the endocarp. The first apparent
trend of variation that we can see in the evolution of this family is
the reduction of the stamens to a definite number (30, 20, 10). Ob-
viously connected with this reduction is the separation of twin anther
sacs in disjunct unilocular thecae and a successive reduction of the
four thecae to two. Another evolutionary trend is the reduction of
the two ovules in each cell to one. Yet another is the appearance
of ribs on the endocarp, a divergence from the almost smooth endo-
carps of the more primitive forms. Very important evolutionary
tendencies are the reduction in length of the germinal opercular
valves or, contrarily, the increase in size of the valves, accompanied
by substantial decrease of the intermediate costae. In some stages
the reduction in size of the valves seems to be compensated by the
appearance of apical holes (foramina). The broadening tendency of
the valves is parallel to the appearance in the woody tissue of the
endocarp of resinous cysts, which are lacking in the primitive forms
(Vantanea, Hylocarpa). ‘The most highly evolved endocarps are more
or less bullate and filled with cysts, inconspicuously costate-furrowed,
and with broad adjacent germinal valves.

The first lines of variation consisting in the reduction of the number
of the stamens with dissociation of the anther twin cells affect all other
genera; notwithstanding, the stage with four anther unilocular cells

CUATRECASAS-—HUMIRIACEAE 43

is kept in only two genera (Duckesia and EHndopleura), all others having
the pollen sacs reduced to two. The other basic trend of variation,
the reduction of ovules in the ovary cavities from two to one, affects
all other genera except Humiria. The reduction in length of the endo-
carp valves is represented by Humiriastrum and Endopleura; these in
turn are provided with subapical foramina. The small genera Duckesia
and Hylocarpa, with lengthwise valvate endocarps, may be considered
as more ancient than Endopleura. Hylocarpa is more evolved than
the other two, on the ground of more simplified anthers (with two
pollen sacs); Hndopleura is more advanced on account of the smaller,
inconspicuous valves. Hylocarpa and Endopleura show, in comparison
to Vantanea, a more advanced form of endocarp in the pronounced
furrows and ribs. Humiriastrum exhibits an intermediate stage having
20 stamens, short valves limited to the upper half of the endocarp,
and subapical foramina. Schistostemon and Sacoglottis have almost
nonfurrowed and nonforaminate, wide-valved endocarps provided
with resinous cysts, features typical of the most advanced structure.
Furthermore, Sacoglottis, having attained the maximum reduction in
the androecium (10 stamens), represents the most evolved genus.
Humiria would remain in an independent line not far from Vantanea
because of having the primitive number of two ovules and narrow en-
docarp valves alternating with costae of the same width (see fig. 2).

Most genera of the Humiriaceae are very natural, that is, real.
Almost all are extremely homogeneous, the trends of evolution of
their species being inconspicuous. However, Vantanea, though also
natural, has some specific variations that reveal evolutionary tend-
encies. The oldest type of the genus probably is represented by a
species with coriaceous, broad leaves; medium-sized flowers; white,
subcoriaceous petals; and smooth or nearly smooth endocarps. This
type includes a group (species 1-6) that cover a wide area from Paré
to the Pacific coast and to Costa Rica. One trend of evolution diverg-
ing from this type could be the reduction in the size and thickness of
the leaves, the size of flowers, the number of stamens, and the en-
docarps, which become smaller and corrugated (species 7 and 8).
Another trend is that the flowers become larger and the endocarps
corrugated or tuberculate (species 9-13). One of these latter species
(V. guianensis) exhibits a more advanced degree of evolution through
its very large flowers, red petals, and strongly corrugated thick en-
docarp with a great reduction of the valves, which are inconspicuous
from the outside. On the other hand, V. parmflora shows another
line of evolution, toward a tomentous floral disk and also corrugated
endocarp.

I see few connections between ecology and evolutionary variation.
The Vantanea fruit possibly are eaten by animals and frequently are

44

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Ficure 2.—Trends of evolution in Humiriaceae: a-g, Androecium, from the multistaminate

flower of Vantanea to 10-staminate Sacoglottis: a, Vantanea; b, Hylocarpa; c, Endopleura;
d, Duckesia, theoretically constructed in two whorls; ¢, Duckesia and Endopleura
in one whorl as it appears; f, Humiria, Humiriastrum, Schistostemon; g, Sacoglottts.
h-l, Anthers: h, Vantanea, 2 bilocular anthers; 1-7, Duckesia, Endopleura, 4 unilocular
disjuncted anthers; &, 4 unilocular anthers with sterile upper sacs, transition to /, as in
all other genera. m-n, Ovary: m, loculi biovular, Vantanea, Humiria; n. loculi
uniovular, the other genera. o-z, Endocarp: o, shape and section of typical Vantanea,
from which derive three tendencies: 1, p, Appearance of apical foramina and reduction
in length of valves as in: g, Humiria; r, Humiriastrum; s, Endopleura. 2, t, Reduction
of valves in width and enlargement of costae asin: u, Vantanea guianensis; 0, Hylocarpa;
w, Endopleura. 3, x, Appearance of resinous cysts, enlargement of valves with reduc-
tion of costae as in: y, Duckesia; z, Sacoglottis, Schistostemon.

transported by rivers. The development of more buoyant endocarps
(in Duckesia, Sacoglottis, and Schistostemon) could be favored in
species more likely to live in low regions and flooded lands. On the
other hand, species of Zumiria, which are more widely spread through-
out the mountains, have small, juicy, fleshy fruit more suited to being
spread by birds; the endocarps lack resinous cavities.

IFramily Humiriaceae

Humiriaceae Jussieu in St. Hil. Fl. Bras. Merid. 2:87. 1829.
Ordo XXXV Humiriaceae, Benth. & Hook. Gen. Pl. 1:246-247. 1862.
Linacearum Series Houmirieae, Baill. Adansonia 10:368, 371. 1873.—Hist.

Pl. 5:51, 56. 1874,

CUATRECASAS—HUMIRIACEAE 45

Humiriaceae (Familia), Urban in Mart. Fl. Bras. 12(2):425-454, pls. 92-96.
1877.

Humiriaceae (Familie), Reiche in Engl. & Prantl, Pflanzenfam. 3(4): 35-37, fig.
32. 1890.

Humirioideae (Unterfamilie), Winkler in Engl. & Harms, Pflanzenfam.
19a:126-130, figs. 58, 59. 1931.

Description: Flowers hermaphroditic, complete, actinomorphic.
Sepals 5, persistent, thick and carnose at base, thinner toward margin,
suborbicular or triangular, more or less connate in tube or cupule,
glabrous, pubescent or tomentous outside, sometimes with marginal
or dorsal glands; estivation quincuncial or imbricated; all same size
or 2 outer ones smaller. Petals 5, deciduous or sometimes persistent,
free, thick or membranaceous, usually 3—5-nervate, oblong, linear or
oblong-lanceolate, acute to obtuse, rarely with gland at top, margin
smooth, sometimes with tooth at 1 side near apex, above glabrous,
below glabrous or pilose, white, greenish white, or yellowish white,
rarely red; estivation contorted, cochlear or quincuncial, indistinctly
dextrorse or sinistrorse. Stamens monadelphous, numerous (in-
definite) and pluriseriate or in definite number, 30-10 and 1-2 seriate.
Filaments filiform (when numerous), slender and flexuose, or thick,
complanate, linear, acute at apex, straight and glabrous or papillose;
at base connate in more or less long tube, alternating in different
lengths, sometimes the 5 alternating with the petals are trifurcate at
apex and triantheriferous. Anthers dorsifixed or subbasifixed; thecae
2, bilocular, laterally attached, ellipsoid-oblong and each cell dehiscing
by longitudinal slit, or 4, unilocular, rounded or ellipsoid disjunct
thecae (2 lateral and 2 basal), dehiscing by detachment, or 2 unilocular,
disjunct, basal, dehiscing by detachment; connective thick, fleshy,
ovoid or lanceolate, obtuse at apex or most commonly produced in
apiculum or linguiform appendix. Sometimes some filaments lack
anther; occasionally smaller, sterile filaments (staminodia) present.
Pollen grains usually 3-colporate, also 4-colporate, suboblate, oblate-
spheroidal, prolate-spheroidal (27X32, 31X35, 33 X3y), exine not
verrucose, tenuisexinous (Erdtman). Intrastaminal free disk girding
ovary, membranaccous or subcoriaceous, tubular or cupular, dentate,
lobate, laciniate or composed of 10-20 free scales. Gynoecium
syncarpous, carpels 5 (rarely 4, 6, or 7), opposite sepals or alternate.
Ovary ovoid or ellipsoid, sessile, 5 (4, 6, 8) septate with axile
placentation, cells uniovulate or biovulate. Style single, entire,
columnar, erect, as long as stamens or shorter, rarely longer. Stigma
narrowly or broadly capitate, 5-lobate or 5-radiate. Ovules
anatropous, epitropous with 2 integuments, pending at inner angle of
ovary cells, micropyle pointing upward, raphe ventral; when 2
ovules present in each cell, superposite and lower one hanging from
longer funiculus. Fruit drupaceous from size of pea to that of mango;

46 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

exocarp hard-fleshy varying from pulpy to fibrous, subcoriaceous tex-
ture. Endocarp (putamen) woody, usually very hard, compact or with
many resin-filled, round cavities, rarely spongious-woody, 5 (4, 6,
7) septate, commonly with only 1-2, rarely 3, 4, or 5 seeds developed;
surface smooth, bullate, rugose, or tuberculate, slightly striate or
strongly costate; with germinal dehiscence, provided with as many
longitudinal opercula or valves as carpels, which may open or be
pushed away by emerging embryo at germination of seed inside fruit.
Often subapical foramina are present. Seeds oblong with double
testa, exterior often adherent to putamen, inner membranaceous,
thin; embryo straight or slightly curved, cotyledons oblong or ovate,
often subcordate at base, radicle half as long, endosperm fleshy
and oily.

Woody, evergreen plants from small shrubs to large trees; wood
reddish, hard, often with balsamic juice. Leaves alternate, simple,
often distichous, coriaceous or subcoriaceous, penninerved, entire,
crenulate or slightly serrate, petiolate or rarely sessile, sometimes
decurrent along branches, often punctate-glandulose near margin on
underside, marginal teeth when young often aculeate, later callous-
tipped. Stipules very small, geminate, deciduous or lacking. In-
florescences axillary, pseudoterminal or rarely terminal, paniculate,
often corymbiform, of dichasial type and trichotomous, but through
reduction often with dichotomous or alternate (cincinnate) branching;
branchlets often with incrassate ends, articulate. Pedicels short,
articulate. Bracts and bracteoles persistent or deciduous, small,
amplectant. Wood, according to Record and Hess: “Heartwood
grayish brown to reddish or purplish brown; distinct but not sharply
demarcated, from the sapwood. Luster usually low. Odorless
and tasteless when dry. Hard and heavy to decidedly so; sp. er.
(air-dry) 0.80 to 1.10; weight 50 to 69 Ibs. per cu. ft.; texture medium
to coarse ; grain mostly irregular; not casy to work; is tough and strong;
finishes smoothly; durability doubtful.” For further technical
characteristics, see Record and Hess, p. 191-192, 1943 ; see also
figure 3.

Disrripution: A typical tropical family with 8 genera, 49 species,
and many subspecies, varieties, and forms. They are important
constituents of the Tropical American rain forests as well as of the
subhygrophytic formations of the savannas, campos, restingas, and
caatingas extending from Costa Rica to southern Brazil. Only one
species is found outside America, on the West African coast. This
distribution is shown in figure 4.

The fossil remains of Vantanea found at the west coast of Peru
(Belén, Piura) indicate not only that a wider distribution of the

CUATRECASAS—-HUMIRIACEAE 47

Ficure 3.—Floral diagrams of Humiriaceae: a, Vantanea; ), Endopleura; c, Duckesta;
d, Hylocarpa; e, Humiria; f, Humiriastrum; g, Schistostemon; h, Sacoglottis; i-p, different
kinds of aestivation of the corolla found in Humiriaceae. In and d the white circles
represent sterile stamens; in other diagrams some of the smaller stamens are often
also sterile.

family existed in the Eocene, but also that the mentioned arid region
was rainy and covered with rain forest during Tertiary times.

According to Croizat (1952, p. 388), the Humiriaceae belong to a
Genorheithrum of Gondwanic origin and reached the Americas from
the east. Andean fossils prove that abundant earlier populations
in that region were deeply disturbed by the uplifting of the Andes,
and that the Humiriaceac existed in the New World long before the
Tertiary.

Economic uses: The bark and wood of some species and varieties of
Humiria produce the ‘“umiri” or ‘“umiry-balsam,”’ with properties
similar to those attributed to the Copaiva and Peruvian balsam.
Little is known about its preparation, trade, and chemical composition
(Wiesner, 1927, p. 1040; Wehmer, 1929, p. 597; Urban, 1887, p. 453).
The exocarp of the fruits of Humiriaceae is more or less fleshy and
in some species edible, as in many Humiria varieties and in some of
Sacoglottis and Vantanea. The exocarp and seeds contain a fatty
oil that in some places of the Amazon is used in the domestic economy.
The oil of “uchi’ has organoleptic properties similar to those of

48 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

45
EQUATORIAL SCALE
0 1000 2000 3000 MILES
[are oe reer eres arrar ae eer ee detbed hd ttn)
0 1000 2000 ©3000 4000 KILOMETERS
. MERCATOR'S. PROJECTION
60 —t —f2————_} 60
d er
el
ie
——_—_oe | fe - ce a Oe Oe een eee ra
‘90 60 30 WEST LONGITUDE f¢) EAST LONGITUDE 30

Ficure 4.—Worldwide distribution of the family Humiriaceae.

olive oil (Pereira Pinto, 1956). The seeds of Sacoglottis gabonensis
contain 54 percent fatty oil (Wehmer, 1929, p. 597; Wiesner, 1927,
p. 743).

The wood of the Humiriaceae is hard and used locally in construc-
tion work. Metcalf and Chalk (1950, pp. 277-279) wrote about its
economic uses thus: “The timbers of this family are dense; they are
of little importance though some species of Humiria are used in tropi-
cal America for heavy construction, wheels, etc.”

Key to the Tribes and Genera of Humiriaceae

1. Stamens 50-180; anthers with 2 bilocular thecae: Tribe VANTANEOIDEAR.
Connective of anthers acute. Carpels opposite sepals, 2-ovulate. Endo-
carp with lingulate valves... 2... 0.0.2.2... 2... .1. Vantanea

1. Stamens 10-30; anthers with unilocular, free thecae: Tribe HumrRioipEAE.

2. Anthers with 4 unilocular thecae; connective acute. Carpels opposite
sepals, l-ovulate.

3. Endocarp spongy-lignose, evenly costate with long, lingulate valves,

resinous-lacunose. Stamens 20-25 ........ . .2. Duckesia

CUATRECASAS—HUMIRIACEAE AQ

3. Endocarp prominently, sharply costate and furrowed with shorter, in-
conspicuous valves at bottom of furrows; compact-woody, not resinous-
lacunose. Stamens 20-30. ........ . .. +3 Endopleura

2. Anthers with 2 unilocular thecae.

4. Stamens 30, anthers with thick, very obtuse connective; thecae basal,

glabrous. Endocarp strongly costate. valvate at furrows, compact-

woody, not resinous-lacunose. ....-.. - . . . «4. Hylocarpa
4, Stamens 10-20, anthers with attenuate, acute connective (very rarely
obtuse).

5. Thecae of anthers basal, pilose. Carpels opposite petals, 2-ovulate.
Endocarp woody, striate, evenly costate-valvate, 5-foraminate at
apex, not resinous-lacunose, valves linear, oblong or lingulate.

5. Humiria

5. Thecae of anthers glabrous. Carpels opposite sepals, 1-ovulate.

6. Stamens 10. Thecae of anthers inferolateral. Endocarp shallowly
or inconspicuously furrowed, not foraminate at apex, resinous-
lacunose, valves broad, adjacent, alternating ribs thin, inconspic-
uous . 2. ee ee ee ee ew ee ew ee + 6 8 Sacoglottis

6. Stamens 20.

7. Episepalous stamens 5, longer, trifurcate at apex, triantheriferous.

Epipetalous stamens 5, medium-sized, entire, monantheriferous.

Ten shorter alternate stamens monantheriferous. Thecae of

anthers inferolateral. Endocarp shallowly or inconspicuously

furrowed, not foraminate at apex, resinous-lacunose, valves
broad, adjacent, alternating ribs inconspicuous,

7. Schistostemon

7, All 20 stamens monantheriferous, 10 episepalous and epipetalous

longer than alternating ones. Thecae of anthers basal. Endo-

carp 5-foraminate at apex, with 5 alternating descending, oblong

and short opercular valves... . . . . - . 6. Humiriastrum

Tribe Vantaneoideae

Vantaneoideae Cuatr., tribus nova Humiriacearum.
Stamina numerosa antheris thecis duobus bilocularibus loculis
rimoso-dehiscentibus. Genus typicum Vantanea Aublet.

1. Vantanea

Vantanea Aubl. Pl. Guian. 1:572, pl. 229. 1775.—Lam. Encycl. 8:334, pl. 471.
1808.—Benth. in Hook. Journ. Bot. Kew Mise. 5:98. 1853.—Benth. &
Hook. Gen. Pl. 1:246. 1862.—Urb. in Mart. Fl. Bras. 12(2):450. 1877.—
Reiche in Engl. & Prantl, Pflanzenfam. 3(4) :37, fig. 32. 1890.—Winkl. in
Engl. & Harms, Pflanzenfam. 19a:106, 128, fig. 59. 1931.—Lemée, Dict.
Descr. Synon. Gen. Phan. 6:832. 1935.

Lemniscia Schreb. Gen. Pl. (8 ed.) 1:358, 1789.

Lemnescia Willd., Sp. P] 2:1172. 1800.—Pers. Syn. Pl. 2:70. 1807.

Helleria Nees & Mart. Nov. Act. Acad. Nat. Cur. 12:38, pl. 7. 1824.— Mart.
Nov. Gen. et Sp. 2:147. 1827.—Juss. in St. Hil. Fl. Bras. Merid. 2:91.
1829.—Endl. Gen. Pl., 1040. 1840.—Baill. Adansonia 1:209. 1860.

50 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Houmiri Sect. Vantanea Baill. Adansonia 10:370. 1870.—Hist. Pl. 5:48,
figs. 96, 97. 1874.

Houmiri Sect. Vantaneoides Baill. Adansonia 10:370. 1870.—Hist. Pl. 5:48.
1874.

Type species: Vantanea guianensis Aublet.

Calyx cupular with 5 sepals more or less united up to a subentire
margin. Petals 5, free, thick, oblong or linear, the estivation con-
torted. Stamens 50-120, sometimes more (up to 180), in 3 or 4
rows, the filaments thin, glabrous, flexuose, inferiorly connate in a
tube surrounding the ovary. Anthers ovate-lanceolate, affixed near
the base; thecae 2, bilocular, ellipsoid, attached at the lower side,
each cell dehiscent by a longitudinal cleft; the connective thick,
ovate-oblong, acuminate, acute, or subobtuse. Disk cupular dentate
or fimbriate, girdling the ovary. Carpels opposite the sepals, biovul-
ate. Ovary 5-locular, the two ovules in each cell anatropous, with

ZIMMERMANN —___.

Figure 5.—a-e, Vantanea parviflora: a, bud, X 214; b, open bud showing the androecium;
c, open flower, X 2% (Ducke 23425); d, anthers, X 10; ¢, gynoecium surrounded by the
disk, X 10 (Schultes F Lopez 9267). f-l, Vantanea minor (Tamayo 3123): f, petal,
x 21; g, bud, X 214; h, androecium, X 2!M; 7, gynoecium surrounded by the disk,
% 2%; 7, longitudinal section showing the staminal tube, disk, and ovary, X 744;
k, anthers, X 10, inner, outer, and lateral view; 2, transection of the ovary, X 7%.
m-n, Vantanea micrantha (Ducke 751): m, open flower, X 244; n, bud, X 214.

CUATRECASAS—HUMIRIACEAE 51

ventral raphe, hanging superposed from the inner angle, the lower
one with long funiculus. Style erect, attaining or exceeding the
stamens. Stigma more or less thickened, 5-lobate. Drupe medium-
sized or large, smooth, ovoid or ellipsoid, the exocarp carnose, sub-
coriaceous when dry, thick or thin. Endocarp woody without resin-
iferous cavities, dehiscent at germination of the seeds by longitudinal,

Z\MMERMANN

Ficure 6.—Vantanea, fruit, < 1: a-c, Vantanea paraensis (Ducke 23430): a, fruit; },
endocarp; c, transection. d-f, Vantanea celativenia (Krukoff 7182): d, fruit; ¢, endo-
carp; f, transection. g-t, Vantanea magdalenensis (Lamb 133): g, fruit; 4, endocarp;
i, transection. j, Vantanea barbourii (Barbour 1018), endocarp. k-l, Vantanea oc-
cidentalis (Patifio 12): k, endocarp; /, transection.

52 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

linear or oblong valves or opercula, which are pushed away by the
emerging radicle of the embryo. Usually only 1 seed develops in
each cell and only 1 (rarely 2 or 3) can be counted in each fruit.
Evergreen trees with coriaceous or subcoriaceous, simple, alternate,
petiolate, or sessile leaves. Inflorescences axillary and terminal,
paniculate, usually dichotomous, also with alternate branching.
Bracts deciduous. (See also figs. 1, 3, and 5-7.)

ZIMMERMANN —__.

Ficure 7.—Vantanea, fruit, X 1: a-b, Vantanea compacta (Ducke 1513): a, fruit; 8,
endocarp. c-d, Vantanea parviflora: c, fruit (Ducke 23425); d, endocarp (Ducke 23426).
e-f, Vantanea micrantha (Ducke 30135): ¢, fruit; f, endocarp. g-h, Vantanea minor
(Bernardi 2613): g, fruit; h, endocarp.

Vantanea comprises 14 species spread throughout the tropical rain
forests, from the Brazilian State of Santa Catarina (its southern
limit) to Costa Rica in Central America. Most of the species are
endemic to limited regions. They grow at low altitudes, though not
in flooded areas, and can be found on mountains up to 800 m. elevation
in the Guianas. Except for V. guianensis, which exhibits large, red,
or red-purple petals, the flowers are white or whitish.

The name “Vantanea” is derived from the Caribbean name
‘fovantan,” which is given in French Guiana to Vantanea guianensis.

Key to the Species of Vantanea

1. Fruit with smooth pericarp.
2. Disk tomentose. Ovary hispid-villose. Drupe ellipsoid, 2.5-2.8>2.2-2.5
cm. Endocarp very rugose, 2.1-2.5%X1.8-2 cm. Petals white.
14. V. parviflora

CUATRECASAS—HUMIRIACEAE 53

2. Disk glabrous.
3. Ovary more or less tomentose or hirsute. Petals white above.

4. Leaf blades small, 3.5-7(-9) X 1.3-2.5(—4) em. oblanceolate or lanceolate-

elliptic, rarely oblong-elliptic. Endocarp rugulose.

10. V. compacta

5. Drupe obovate-ellipsoid, oblong, rounded at apex, narrowed at base,
2.4-2.8%1.5-1.8 em. Endocarp oblong-ellipsoid, about 2.5% 1.4

cm. ... . . 10a. V. compacta subsp. compacta

5. Drupe globose- ovoid, subrounded or obtuse at apex, rounded at
base, 1.6-1.8%X1.4-1.6 em. LEndocarp subglobose-ovoid about
1.7X14em.... . . .10b. V. compacta subsp. microcarpa

4, Leaf blades larger, 4-19 2.5-10 cm., ovate or obovate, also elliptic.

Iindocarp smooth or nearly so (unknown in V, obovata).
6. Petals pubescent or tomentose outside.

7. Petiole 2-4 mm, long. Ovary oblong, glabrous on lower third,
villose above. Disk membranaceous, tubular 1-3 mm, high.
Leaf blades obovate, large (10-19 6.5-10 cm.). Petals 5.5-6
*2.5 mm. Endocarp oblong-ovoid, rounded at base, acute
at apex... Lew ew ew ww ee dL. V. magdalenensis

7. Petiole 6-12 mm. long. Ovary ovoid, long-tomentose-hirsute.
Disk rather thick, cupular, less than 1 mm. high. Leaf blades
elliptic or obovate, small (4-12 X 2.4-8 cm.).

8. Petals 10-13 mm. long. Branchlets of inflorescence densely
tomentose-hirtous. Stamens more than 100, 6-9 mm. long.
2. V. obovata
8. Petals about 7 mm. long. Branchlets of inflorescence short-
tomentulose-hirtellous. Stamens 50-60, 5-7 mm. long.
Endocarp oblong-ellipsoid, obtuse at both ends, 2.71.6 cm.
3. V. barbourii
6. Petals glabrous.

9. Petioles 1-5 mm. long. Ovary densely long-hirsute-lanate.
Petals about 9 mm. long. Drupe ovoid-ellipsoid, 3.51.8 cm.
long. Endocarp elipscia-attennate at both ends, 3-3.3X1.5-
17em.... ee ew ee ee e & Ve occidentalis

9. Petioles 10-20 mm. "Jong.

10. Drupe oblong-ellipsoid, 5-5.32.8-3 em. Endocarp ellipsoid-
oblong, obtuse at both ends, 4.62.5 cm. with 7 valves.
Exocarp 2 mm. thick, Ovary short-velvety-tomentose.
Petals 6-8 mm. long. . . . .. . 5. V. paraensis
10. Drupe ovoid-ellipsoid, 3X1. 6-1. 7 cm. rounded at base, acute
at apex. Endocarp ovoid-ellipsoid, rounded at base, acute
or apiculate at apex, 2.8-3X1.6 em. with 5 valves, Exo-
carp 0.5mm. thick. Flowersunknown . 6. Y. celativenia
3. Ovary glabrous. Petals glabrous.

11. Inflorescence glabrous. Flowers 25-40 mm. long. Petals red, 25-35
mm. long. Stamens 26-35 mm. long, tube 5-8 mm. high. Anther
cells oblong, 0.8 mm. long, the connective with short, acute tip.
Calyx 4 mm. high, lobes glandular outside. Petioles 6-12 mm.
Pedicels 2-5 mm... . .... . -13. V. guianensis

11. Inflorescence more or less densely “tomentulose- hirtellous. Flowers
smaller, petals white or whitish, less than 15 mm. long.

12. Flowers small, petals 4-5 mm. long. Calyx 0.6-0.7 mm. high,
papillose outside, margin ciliate. Leaf blades thin, flexible,

513359—61——_3

54 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

lanceolate or lanceolate-elliptic, 5-12X2-5 cm, Petioles 3-8
mm. long. Disk 1-2 mm. high more or less laciniate. Drupe
medium-sized 20-25 X 14-22 mm. _Endocarp strongly corrugate-
suleate,20X16mm,... . .... .%. V. micrantha
12. Flowers larger, the petals 10-141 mm. Leaf blades rigid, coriaceous.
13. Blades large (10-20X5-10 ecm.). Petioles 6-12 mm. Calyx
pubescent-hirtellous. Disk 1.8 mm. high, laciniate. Anther-
connective elongate, subacute. Drupe large, ellipsoid (5-10
X3.5-4.5 em.). Endocarp anfractuose-rugose.

ll. V. macrocarpa
13. Blades smaller (3-8X1.2-4.5 cm.), rigid. Petiole 1-4 mm.

Calyx glabrous. Disk short (1 mm. high), denticulate.
14. Petals 12-14 mm. long. Stamens 9-12 mm. long, tube 3-4 mm.
high. Anther-connective short, thick, obtuse. Drupe smooth
3.5-5 X 2.2-3 em. Endocarp densely anfractuose-rugose, 3-4
x2-3 em. Petiole1-2mm.long..... .8 V. minor
14. Petals about 11 mm. long. Stamens 7-10 mm. long, tube
1-1.5 mm. high. Anther-connective elongate, subobtuse.
Drupeunknown, Petioles3-4mm.long . . .9. V. peruviana
1. Fruit with strongly tuberculate, warty pericarp, 6-8 cm. long 4-6 cm. broad.

Exocarp 8-10 mm. thick. Endocarp anfractuose-rugose, cavernous.
12. V. tuberculata

1. Vantanea magdalenensis Cuatr., Brittonia 8:195. 1956.

FIGURE 8,a-c

Type: Lamb 133, Colombia, Magdalena valley.

Large tree; terminal branchlets subterete, greenish, lenticellate,
glabrous. Leaves rather thick coriaceous, glabrous. Petioles thick,
2-4 mm. long. Blades entire, obovate, oblong-obovate or obovate-
elliptic, more or less narrowed, obtusely cuneate or subrounded at
base, rounded or very obtuse at apex, 11-19 cm. long, 6.5-10 cm.
broad; brownish above when dry, the midrib conspicuous, secondary
nerves filiform, minor veins obsolete; pale brown beneath and with
very prominent midrib, 8-10 prominent secondary nerves on both
sides, subspreading, near margin arched, ascending, anastomosed,
and slender, minor nerves prominulous and loosely reticulate.

Inflorescences at end of branchlets, paniculate, shorter than upper
leaves; branches and branchlets dichotomously articulate, hirtellous-
tomentose. Bracts deciduous. Pedicels thick, tomentose, about 1
mm. long. Calyx when open 3 mm. diameter, lobes rounded, 1 mm.
long, pubescent outside. Petals obovate-linear, rather thick, 5.5-6
mm, long, 2.6 mm. broad, tomentose outside. Stamens many, about
triseriate, unequal, filaments glabrous, united in a ring 1-2 mm. high.
Anthers minutely ovate-rhomboid, 4-lobed, the connective thick,
attenuate toward the apex. Disk tubular, membranous, eroded at
margin, 1-3 mm. high. Ovary oblong, 3 mm. high, the lower third
glabrous, pubescent above. Styles filiform, glabrous, 2 mm. long.
Drupe subovate-elliptic, rounded at base, slightly narrowed at apex,

CUATRECASAS—HUMIRIACEAE 55

b

Se

Ficure 8.—a, Vantanea magdalenensis, X ¥% (Lamb 133); b, Vantanea magdalenensis,
bud, X 1%; c, Vantanea magdalenensis, petal, X 1%; d, Vantanea obovata (Glaziou
18963); e, Vantanea obovata, bud, X 1%.

about 3.2 cm. long, 2 cm. thick. Exocarp gray when dry, 1 mm. thick,
minutely velutinous-tomentose. Endocarp woody, about 2.8X1.8
cm., broadly rounded at base, acute at apex, surface slightly rugulose,
practically smooth, with 5 marked ribs and 5 oblong, 2.5-em.-long,
5-mm.-wide valves.

V. magdalenensis is the only species of the genus found in the interior
valleys of Colombia. It is a large tree of the upland rain forests. Its
distribution is limited to the Magdalena Valley.

COLOMBIA: SanTanpeEr: Valle del Magdalena, Cimitarra, km. 3 de la carre-
tera del Ermitafio, 24-VII-1954, ‘“‘macabalo,” Lamb 133 (US, holotype; COL,
isotype).

2. Vantanea obovata (Nees & Mart.) Benth. in Hook. Journ. Bot. Kew Misc.
5:99. 1853.—Urb. in Mart. Fl. Bras. 12(2):451. 1877. Figure 8,d-e
Helleria obovata Nees & Mart. in Mart. Nov. Act. Acad. Nat. Cur. 12:40,

pl. 7. 1824.—A. Juss. in St. Hil. Fl. Bras. merid. 2:91, pl. 504. 1829.

Helleria ovalifolia A. Juss. in St. Hil., ibid.: 91. 1829.
Vantanea ovalifolia Benth. in Hook. Journ. Bot. Kew Misc. 5:99. 1853.

Types: Martius, Brazil, Minas Geraes, Tejuco. St. Hilaire,
Bomfin, Brazil, Minas Geraes, Minas Novas (type of Helleria ovalifolia
Juss.).

56 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Tree with subterete, granulate-lenticellate, glabrous or scarcely
puberulous, terminal branchlets. Leaves rigid, coriaceous, glabrous.
Petioles 6-12 mm. long, subterete more or less sulcate above. Leaf
blades entire, obovate or obovate-elliptic, obtusely or acutely cuneate
at base, rounded or very obtuse and often retuse at apex, 4-10 cm.
long, 2.5-6 cm. wide; above lustrous, with slightly prominent main
nerves, the smaller ones less conspicuous; midrib and 8-9 pairs of
secondary nerves prominent beneath, the latter subascendent, curving
and anastomosing near the margin, smaller veins prominulous and
reticulate, often with glandspots on the nerves at the middle.

Inflorescence cymose-paniculate, dichotomous, ending the branch-
lets or in the axils of the upper leaves; peduncle strong, sparingly
puberulous, the branchlets rather thick and densely hirtellous. Pedi-
cels thick, about 2 mm. long, densely hirtulo-pubescent. Bracts ovate,
about 1.5 mm. long, soon deciduous. Calyx about 2 mm. high,
pubescent, with suborbicular, ciliate lobes. Petals white, linear-
oblong, 10-13 mm. long, 2.5-3 mm. wide, densely hirtulo-pubescent
outside with spreading or retrorse hairs, margin glabrous, estivation
contorted. Stamens more than 100; 2-3 seriate, united at base in a
cup; the filaments glabrous, unequal, 6-9 mm. long. Anthers ovate-
lanceolate, 0.8—0.9 mm. long, with fleshy, lanceolate connective and 4
elliptic, basilar lobes. Disk annular, girdling the ovary, glabrous, 1
mm. high. Ovary subglobose, 2 mm. high, densely hirsute-villose,
with 5 biovulate cells. Style thick, erect, pilose only at base, 4-6 mm.
long. Stigmas obtuse. Drupe ovoid, rounded at base, obtuse at
apex, densely tomentous, about 2.2 cm. long, 1.6 cm. diameter.

V. obovata is a small tree at present only known from the States of
Minas Geraes and Bahia in Brazil.

BRAZIL: Minas Gerars: Diamond Dist., common on the Serra Mendanha,
small tree, Gardner 4452 (BM). Estrada de Bom Gosto a Olivenga; arvore,
15-III-1943, Frées 19933 (NY, IAN, US). Carraseo-Chapad&o, Municipio de
Diamantina, muito frequente; flores alvas, arbusto 2 m., 29-IV-1942, Mendes
Magalhées 2117 (IAN). “Minas Geraes,” St. Hilaire B 1705 (P, holo-
type); photo F.M. 35175. Minas Geraes, Catal B, St. Hilaire 1984 bis (P).
Biribiry, arbuste fleur blanc jaundtre, 22-III-1892, Glaziou 18963 (P, US);
photo F.M. 12590 of Gardner 4452. ‘Habitat in alpestribus districtus adaman-
tini prope Tejuco, etc., Provinciae Min. General. Dr. Martius Iter Brasil.,’’ Jul.,
Martius s.n.(M.holotype). Minas Gerais, Dr. Martius Iter Brasil. Observ. 1274,
Martius s.n. (M, 3 sheets). Brasilia in altis Serra do Frio, Provinciae Minarum
Generalium, Observ. 1274 conf. HWirtella polyandra Kunth, Martius s.n. (M).
Brasilia in alpestribus Provinciae Minarum Generalium, Dr. Martius Iter Brasil.,
Martius s.n. (M, isotypes, 2 sheets). Banta: Campos casa Pedra, 27-VII-1914,
Luetzelburg 40 (M). Jussieu quoted for V. obovata the following location, prob-

ably based on St, Hilaire’s collections: ‘‘Nascitur prope vicum Curumatahy in
parte deserta occidentalique Provinciae Minas Geraes quam vocant Certao.”’

CUATRECASAS—-HUMIRIACEAE 57

3. Vantanea barbourii Standl. Trop. Woods 75:5. 1943.—Barbour, ibid.:
7-8. 1943.—Record, ibid., 77:8-9. 1944.—Allen, The rain forests of
Golfo Dulce 351, fig. 20. 1956. Ficure 9,f-h

Type: Dayton & Barbour 3129, Costa Rica, Cartago.

Large tree with angular, somewhat lenticellate, glabrous branchlets.
Leaves coriaceous, firm, glabrous. Petioles 6-10 mm. long, flat
above, thickened at base. Blades entire elliptic or oblong-elliptic,
attenuate at both ends, cuneate base, obtuse but sometimes rotundate
or even emarginate apex, 5-12 cm. long, 2.4-6 cm. wide; above
brownish-green, lustrous, with midrib and secondary nerves flat and
conspicuous, the smaller veins little visible or obsolete; pale-brownish
with prominent midrib beneath, the 7—9 lateral nerves on each side,
prominent, ascendent, curved and connected at the ends, the lesser
veins prominulous, loosely reticulate.

Ficure 9.—a, Vantanea occidentalis, X VY (Cuatr. 19937); b, Vantanea occidentalis, bud,
X 1%; ¢, Vantanea paraensis, X V4 (Ducke 752); d, Vantanea paraensis, bud, X 1%
(Ducke 17782); ¢, Vantanea paraensis, petal, X 133 (Ducke 17782); f, Vantanea barbourit,
x % (Dayton &F Barbour 3129); g, Vantanea barbourii, bud, X 1%; h, Vantanea
barbourti, petal, X 1%.

58 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Inflorescence axillary subterminal, paniculate-cymose, leafy, as
long as the leaves, the peduncle strong, glabrous or subglabrous, the
branchlets angled, dichotomo-furcate, shortly hirtellous. Bracts
caducous. Pedicels 1-2 mm. long, minutely puberulous. Calyx
1-1.5 mm. high, slightly puberulous, deeply lobate, with rounded
ciliate lobes. Petals white, oblong, subacute, attenuate towards the
apex, glabrous inside, adpressed retrorse-pubescent outside, about
7 mm. long and 2.5 mm. wide. Stamens 50-60, glabrous, the fila-
ments 5-7 mm. long, complanate, united at base. Anthers ovate-
lanceolate, about 0.8 mm. long, with 4 small, ellipsoid lobes, the con-
nective thick, acuminate-lanceolate, twice as long as the lobes. Disk
thick, glabrous, 1 mm. high; margin short-denticulate. Ovary
ovoid, 1.5-2 mm. high, densely tomentose-hirsute, 5-loculate, cells
biovulate. Style rigid, glabrous, about 4 mm. long. Drupe ovoid-
oblong, smooth, obtuse at the basc, attenuate at the apex, 2.8-3 cm.
long, 1.8-2 cm. broad. Endocarp ellipsoid-ovoid, oblong, about 2.7
cm. long, 1.6 cm. broad, obtuse at both ends; the valves 5, oblong,
obtuse at apex, about 1.8 cm. long, 4-5 mm. wide.

Of V. barbourii, Paul Allen gives a good illustration and writes
the following first hand data and observations: “Ira chiricana or
Nispero. Tall forest trees, 90-120 ft. in height and up to 5 ft. in
diameter above the basal buttresses, with alternate, short-petiolate,
leathery leaves, the oblong-elliptic blades 3.5 in. in length, with
rounded or cmarginate apices and cuneate bases. The trees are
completely deciduous for a brief period at about the end of the heavy
rains in November or December. The small white flowers are pro-
duced in June, and again from about November until January in
dense terminal or subterminal corymbs, followed by ellipsoidal fruits
about 1 in. in length. The seeds have a very distinctive appearance,
and are frequently seen on the forest floor. The strong, reddish-
brown wood is fairly hard and heavy, and is reported to be very
durable, having been used for bridge timbers on the Pan-American
highway. Locally very common on forested hillsides up to about
2,500 ft. in elevation. Golfo Dulce, without definite locality, H. J.
Marks, s.n. (Yale School of Forestry 16822), Esquinas Forest, 250
ft., Allen 6546, Hills near Palmar Sur, 200 ft., Allen 6681.”

Barbour states: ‘The range of Ira Chiricana, so far as is now known,
is confined to a relatively small area on the rolling hills near San
Isidro, at elevations from 700 to 800 meters above sea level...
[It] . . . has a thin brownish gray bark with longitudinal fissures.
It is usually buttressed at the base, above which it has a good timber
form. Specimens 36 inches or more in diameter with 40 feet or
more of clear length, are not uncommon. Its fairly hard and heavy
reddish brown wood has a good local reputation for strength and

CUATRECASAS—HUMIRIACEAE 59

durability and is being used for bridge timbers on the Inter-American
Highway, which traverses its range.”’

COSTA RICA: San Jos&: 1 mile north of San Isidro del General, % mile west
of Pan-American Highway, right of way, alt. 2,500 ft., rain forest, grows in
patches, tree 70 ft. high, DBH 12 in., size up to 100 ft., usable length 40 ft.,
strongest wood in this locality used for heavy construction, pits in scalariform
arrangement bark brownish gray with longitudinal fissures, 11-VJ-1943, “ira
chiricana,’”’ Dayton & Barbour 3129 (US, holotype; F, isotype). Carraco:
2 km. south of Rio Hermosa, about 8 km. from San Isidro del General, alt. 2,100
ft., mixed virgin forest, tree 75 ft. high, DBH 16 in., 6-IX-1948, “‘nispero,”’
‘Gra chiricana,”’ Barbour 1018 (F, Y).

4. Vantanea occidentalis Cuatr. Trop. Woods 96:40. 1950.
Fiaurss 1; 6,j-l; 9,a—b

Type: Cuatrecasas 19937, Colombia, Valle, Buenaventura.

Large tree; branchlets green, cinnamon-brown when dry, lustrous,
glabrous. Petioles 1-5 mm. long, robust, sulcate above, thickened
at base. Leaf blades coriaceous, entire, cinnamomeus when dry,
elliptic, obovate-elliptic or oblong-elliptic, cuneate at base, abruptly
attenuate, obtuse or shortly acuminate at apex, 7-16 cm. long, 2.5—
8.5 cm. broad; green above, lustrous, with well-marked midrib,
the secondary ones immersed, obsolete; with thick and prominent
midrib beneath, the 8-10 secondary nerves on each side prominent,
ascendent, near the margin arched and united, the minor veins loosely
reticulate, slightly prominulous.

Inflorescences at the end of the branchlets paniculate, shorter
than the upper leaves, many-flowered; branchlets dichotomously
articulate, shortly pubescent; the bracts caducous, not seen by the
author. Pedicels thick, puberulous, 1-2.5 mm. long. Sepals orbicu-
lar, subcoriaceous, glabrous or subglabrous, pale greenish, 6-7 mm.
long with a single gland outside. Petals subcoriaceous, oblong,
rather obtuse, 9 mm. long, 3 mm. wide, glabrous, outside greenish-
white, inside white. Stamens numerous (about 80); filaments
unequal, about 8 mm. long, linear, rather thick, acute at apex, white,
glabrous, united at base. Anthers small, ovate-lanceolate, the
connective thick at base, acute at apex. Disk cupular, 1 mm. high,
striate, minutely dentate, glabrous. Ovary 3-4 mm. high, very
hirsute, with 5 or 6 biovulate cells. Styles filiform, 5 mm. long.
Drupe oblong-ovoid-ellipsoid, narrowed at both ends, about 3.5 cm.
long and 1.8 cm. broad. Endocarp woody, one-celled, about 3 cm.
long and 1.5-1.7 cm. thick, ellipsoid-oblong, attenuate and subacute
at both ends; surface rather smooth, with 5 broad ribs alternating
with 5 oblong, 2-cm. long and 5-mm. wide valves, only one of them
removable. One oblong seed in the single remaining cavity developed,
about 2 cm. long.

60 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

V. occidentalis is a rain forest hardwood tree with a distribution
restricted to the western Colombian coast and Chocé region.

COLOMBIA: Vatie: Bahfa de Buenaventura (Costa del Pacffico), Quebrada
de San Joaquin, 0-10 m., alt.; Arbol grande; hoja coridcea, flexible, verde brillante
haz; capullos verde claros, pétalos blanco-verdosos por fuera, blancos por dentro,
filamentos blancos; drupa verde, dura, ovoideo-aguzada, 3 em. long, 22-II-1946,
Cuatrecasas 19937 (F, holotype; G, VALLE, US, isotypes). Buenaventura,
1945, Patifio 12 (F, US, paratypes). CHocé: Quibdé, alt. 40 m., III-1958,
“chan6,” only fruits from the ground, Ramos & Patifio s.n. (US).

5. Vantanea paraensis Ducke, Arch. Jard. Bot. Rio Janeiro 4:99. 1925.—
Arch. Inst. Biol. Veget. Rio Janeiro, 4:30, 31. 1937.
Fiaurns 6,a-c; 9,c—e

Type: Ducke 17782, Brazil, Pard, Rio Tapajoz.

Large tree; branchlets subterete, glabrous, sparsely lenticellate.
Leaves rigid, thick-coriaceous, glabrous. Petioles 10-15 mm. long,
semiterete, subsulcate above, thickened at base. Blades entire,
obovate-elliptic, cuneate at base, broadly obtuse or subrounded and
occasionally emarginate at apex, 7-12 cm. long, 3-6.5 em. wide;
obscurely yellowish-green above with prominulous midrib and less
visible secondary nerves; pale with thick midrib beneath, the second-
ary nerves 8-10 on each side, prominent, ascending at some distance
from the margin, thinner and arcuately connected, the lesser veins
prominulous and loosely reticulate.

Inflorescence cymose-paniculate, ending the branchlets or axillary
in the upper leaves, corymbiform, dichotomous at summit, rather
smaller than the leaves; peduncle strong, ribbed, slightly hirtellous-
puberulous; branchlets angled, hirtellous-puberulous. Bracts soon
deciduous. Pedicels thick, subterete, minutely puberulous or glabrous.
Calyx 5-lobate, about 0.5 mm. high and 1.5 mm. in diameter, with
rounded, outwardly glabrous lobes, margin minutely ciliate. Petals
white, thickish, oblong, subacute, glabrous, 6-8 mm. long, 2-2.5 mm.
wide above the base, estivation contorted. Stamens about 50, the
filaments glabrous, flexuous, complanate below, shortely united at
base; the anthers ovate-lanceolate, about 0.9 mm. long with 4 ellip-
soid lobes and fleshy, lanceolate, elongate connective. Disk thick,
annular, glabrous, about 1 mm. high with dentate margin. Style
3.5-5 mm. long, only pilose at base, thick but attenuate toward the
apex. Ovary ovoid, about 2 mm. high, furrowed, densely and shortly
velvety-tomentose, 5-celled, the cells biovulate. Stigma obtuse.
Drupe oblong-ellipsoid about 5-5.32.8-3 cm.; exocarp coriaceous
when dry, about 2 mm. thick, glabrous or subglabrous; endocarp
woody, oblong, obtuse at base, thick above, narrowed and acute at
apex or apiculate, 4.6 cm. long, 2.5 em. broad, the surface slightly
rugose; the valves 7, linear-oblong, obtuse at apex, 3-3.5 cm. long,
about 5 mm. broad.

CUATRECASAS—-HUMIRIACEAE 61

V. paraensis is a Brazilian species, known only from the regions of
Manaos and Rio Tapajoz.

BRAZIL: Amazonas: Manaos, silva non inundabili ultra flores; arbor magna
floribus albis, 12-VII-1941, Ducke 752 (IAN, MG, US). Manaossilva terris
altis circa rivum Mindt superiorem; arbor magna, floribus albis suaveolentibus,
22-X-1929, Ducke 23430 (P, 5, U, US). Lago de Badajoz, rio Capitari, terra
firme, alta, floresta alta arvore 25 m., flores brancas, 29-VIII-1950, Frées 26428
(IAN, US). Pard: Rio Tapajoz, circa cataractas infimas loco Bella Vista, silva
non inundabili, arbor magna floribus albis, 24-IX-1922, Ducke 17782 (US, 58, U,
isotypes), photo I’.M. 12591 from Berlin.

6. Vantanea celativenia (Standl.) Cuatr., comb. nov.
Fiaures 6,d-f; 10,a—b
Licania celativenia Standl. Field Mus. Bot. 17:254. 1937.

Type: Krukoff 7182, Brazil, Amazonas, Rio Madeira.

The vegetative parts of this species agree completely with V.
paraensis, at least for the specimens now available. But the fruit
of the type of Standley’s species are so different that they define

NPN
TAY

Ficure 10.—a-b, Vantanea celativenia, X% (Krukoff 7182); c, Vantanea compacta, X iy
(Blanchet 3362); d, Vantanea compacta bud, 1%; ¢, Vantanea compacta, petal,
1%; f, Vantanea micrantha, X % (Ducke 751); g, Vantanea minor, X ¥% (Tamayo 3123).

62 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

another species. The flowers, which are lacking, probably will pro-
vide complementary characterization when found.

Drupe ovoid-ellipsoid, about 3 cm. long and 1.6-1.7 cm. broad,
rounded at the base, acute at apex. Exocarp thin (about 0.5 mm.
thick when dry), minutely papillose and sparsely pilose. Endocarp
woody, ovoid-ellipsoid, more or less oblong, rounded at base, acute
or shortly apiculate at apex, 2.8-3 cm. long, about 1.6 cm. broad,
marked by 5 broad ribs alternating with 5 oblong, appressed valves,
2.2 cm. long, 4 mm. wide.

The distribution of V. celativenia is restricted to the Municipio de
Humayta in the State of Amazonas, Brazil.

BRAZIL: Amazonas: Basin Rio Madeira, Municipio de Humaytd, on plateau
between Rio Livramento and Rio Ipixuna; tree 110 ft. high, on campinarana,
7-18-IX-1934, Krukoff 7182 (US, NY, 5, U, isotype of Licania celativenia).
Municipio of Humaytd, near Tres Casas, on lower terra firma, trees 120 ft. high,
14-IX-11-X-1934, Krukoff 6371 (A, BM, NY, §, U, US).

7. Vantanea micrantha Ducke, Arch Inst. Biol. Veget. Rio Janeiro 4:30. 1938.
Fiaurss 5,m-—n; 7,e-f; 10,f

Type: Ducke 30135, Brazil, Amazonas, Manaos.

Large tree with slender, gray, almost smooth, sparsely lenticellate
branchlets, glabrous or the youngest ones puberulous. Leaves thin-
coriaceous, glabrous. Petioles 3-6 mm. long, sulcate above. Blades
entire, elliptic-oblong or elliptic-lanceolate, broad near the base,
abruptly narrowed and tapering to the petiole, narrowed-acuminate
or cuspidate at apex, 5-12 cm. long, 2-5 cm. broad; green-olivaceous
and with prominulous midrib above, the secondary nerves and veins
very thin; lighter and with prominent midrib beneath, the secondary
nerves 10-12 each side, slender, curved-ascending, prominulous, the
veins reticulate, prominulous; bearing a few seriate glands at the
middle.

Inflorescences cymose-paniculate, axillary and terminal, much
shorter than the leaves, the peduncle and branches minutely hirtellous
or papillose. Bracts early deciduous, ovate-lanceolate, about 1.2 mm.
long. Pedicels thick, 0.5 mm. long. Calyx 0.6-0.7 mm. high, sub-
cupular, slightly 5-lobate, the lobes very obtuse, sparsely papillose out-
side, minutely ciliolate at margin. Petals white, glabrous, oblong,
about 5 mm. long, 1.5 mm. broad at base, the estivation contorted.
Stamens about 50, glabrous, the filaments slender, flexuous, unegual,
2.5-4 mm. long, united at base. Anthers ovate-lanceolate, 0.6—0.7
mm. long, with 4 elliptic lobes, the connective fleshy, elongate, acute
or subacute. Disk glabrous, girdling the ovary, about 1.2 mm. high,
the upper half laciniate. Ovary ovoid, glabrous, striate, 1 mm. high,
with 5 biovulate cells. Styles rigid, glabrous, about 4 mm. long.
Drupe ellipsoid, about 2-2.5%1.4-2.2 cm., yellowish. Endocarp

CUATRECASAS—HUMIRIACEAE 63

ellipsoid-obtuse, woody, strongly costate-corrugate, somewhat re-
sembling the cotyledons of a walnut, about 21.6 cm. Exocarp thin,
oleaginous, sweetish, aromatic.

Characterized by its small, white flowers and small fruit, V. mi-
crantha is only known from the floodless lands of the Manaos region
in Brazil.

BRAZIL: Amazonas: Manaos, silva terris altis circa cataractas fluvi Taruma;
arbor magna floribus albis, 6-VIII-1937, Ducke 30135 (US, P, 8, U, isotypes).

Manoas, silva non inundabili ultra Coloniam Jo&éo Alfredo, arbor magna floribus

albis, 7-XIII-1941, Ducke 751 (GH, MG, IAN, NY, US).

8. Vantanea minor Benth. in Hook. Journ. Bot. Kew Misc. 5:99, 1853.—Urb.
in Mart. Fl. Bras. 12(2):452. 1877. Ficures 5,f-l; 7,g-h; 10,9

Type: Schomburgk 1552, British Guiana.

Medium-size tree with subterete, smooth, glabrous or minutely
and sparsely puberulous branchlets. Leaves coriaceous, rigid, gla-
brous. Petioles very short, 1-2mm.long. Blades entire, obovate or
elliptic-obovate, acute or obtusely cuneate at the base, rounded or
very obtuse at the apex, sometimes emarginate, 3-8.5 cm. long, 1.6-4.5
cm. broad; above lustrous, green, midrib slightly prominulous, minor
veins almost obsolete; cupreous and with prominent midrib beneath;
13-15 secondary nerves on each side, filiform, prominulous, united
near margin, minor veins loosely reticulate, slightly prominulous;
bearing a few seriate glands on each side of midrib.

Inflorescences terminal and axillary-terminal, cymose-paniculate,
with furrowed and minutely puberulous branches. Pedicels thick,
slightly and minutely puberulous, 1.5-2.5 mm. long. Calyx about
2 mm. high, lobes thick, subrotundate, glabrous outside, margin
ciliate. Petals linear-oblong, thickish, white, glabrous, 12-14 mm.
long, 2 mm. broad; estivation contorted. Stamens more than 100,
pluriseriate, 9-12 mm. long, glabrous, united in a 3-4 mm. high tube
at base, free part of filaments filiform. Anthers about 0.6 mm. long
with 4 ellipsoid lobes, the connective thick and obtuse. Disk tubular,
thick with toothed margin, glabrous, 1.2 mm. high, girdling the ovary.
Ovary ovate, glabrous, about 2 mm. high, with 5 biovulate cells, the
ovules superposed. Styles flexuous, about 12 mm. long, glabrous.
Stigma obtuse. Drupe 3.5-5 em. long, 2.2-3 em. broad, oblong-ellip-
soid or ovoid-ellipsoid, rounded at base, obtuse at apex. Exocarp
coriaceous when dry, resinous, reddish-brown, rather smooth, gla-
brous, 2 mm. thick. Endocarp woody, ellipsoid, broadly rounded at
base, abruptly apiculate or obtuse at apex, 3-4 cm. long, 2-3 cm.
broad, strongly anfractuose-rugose, inside slightly cavernous; opercula
5, oblong, obtuse at the apex, about 22 mm. long, 5 mm. broad
(Bernardi 2613) alternating with 5 small foveolae at the top. Ina

64. CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

sectioned specimen (2613) 4 oblong monospermous cavities about
22 mm. long, 3 mm. thick can be seen.

For the description of the fruits, which were unknown before, I used
the Cardona (2362) and Bernardi specimens. Bernardi’s endocarps
are more obtuse at the top than those of the Cardona specimens, but
I have no doubt about their being conspecific and about their identity
with Bentham’s species.

V. minor is endemic in the Guiana region which is called Gran
Sabana in Venezuela and nearby British Guiana.

VENEZUELA: Botivar: Sabana de Iecaburu, entre la reducida faja de
vegetacién lefiosa a lo largo del rfo, aguas negras, alt. 450-850 m.; Arbol 5 m.,
flores blancas, grandes, frutos grandes monospermos, 22—XII-1955, Bernardi
2613 (VEN). Gran Sabana, selvas de galerfa del rfo Urari; arbusto, y a veces
arbol muy frondoso en la selva de galerfa, en matorrales de la serranfa es acha-
parrado y menos frondoso, ITI-1946, Tamayo 3123 (US). Camino del Rfo Tirica
que sale a la Gran Sabana, 900 m. alt., drbol 15 m., X-1947, Cardona 2362 (US).
Rfo Caronf, sitios pedregosos en las orillas del Uaiparu, afluente del Icabaru,
500 m, alt., drbol 10 m., 27-X-1946, Cardona 1912 (US, VEN). Rfo Cuquendn
al sur del Roraima; Blueten weisslich, am Quewewode, II-1910, Ule 8801 (MG,K).

BRITISH GUIANA: No locality, Schomburgk 1552, photo F.M. 12589 of
type in Berlin.

9. Vantanea peruviana Macbr. Candollea, 5: 371. 1934.—Flora of Peru, Field
Mus. Bot. 13:632. 1949, Fiaure 11,c—e

Type: Klug 1130, Peru, Loreto.

Medium-size tree with reddish-brown, lenticellate, glabrous branch-
lets. Leaves rigid-coriaceous, glabrous. Petioles 3-4 mm. long,
thickened at base. Blades entire, obovate or obovate-elliptic, cuneate
at base and decurrent on the petiole, rounded, truncate or even retuse

Figure 11.—a, Vantanea macrocarpa, X¥3 (Ducke 30133); b, Vantanea macrocarpa, bud,
X 1%; ¢, Vantanea peruviana, X ¥ (Klug 1130); d, Vantanea peruviana, bud, X 1% (Klug
1091); ¢, Vantanea peruviana, petal, * 1% (Klug 1091).

CUATRECASAS—HUMIRIACEAE 65

at the apex, 3-8 cm. long, 1.2-4.5 cm. broad; greenish and lustrous
above with marked midrib and almost obsolete veins; brown when dry
beneath, with prominent midrib; secondary nerves 10-12 on both
sides, very slender, almost obsolete, arched and united near the margin,
the lesser veins inconspicuous; bearing a few minute glands spread at
the middle.

Inflorescences in axils of uppermost leaves, forming cymose corymbi-
form pseudoterminal panicle; peduncle and branchlets robust, angled,
hirtello-puberulous. Bracts deciduous. Pedicels thick, glabrous or
subglabrous, 1-1.5 mm. long. Calyx cupular, thick, about 1.2 mm.
high, 5-lobate, lobes subrounded, glabrous outside, the margin mi-
nutely ciliolate. Petals white, thickish, linear, acutish, glabrous,
about 11 mm. long, 1.5-1.8 mm. broad. Stamens about 70, glabrous,
the filaments slender, flexuous, 7-10 mm. long, united in a tube
1-1.5 mm. high. Anthers ovate, 0.6 mm. long, with 4 ellipsoid
lobes, the connective fleshy, oblong, obtuse. Disk tubular, gla-
brous, with toothed margin, girdling the ovary, 1.5 mm. high. Ovary
ellipsoid, glabrous, 1.5 mm. long, with 5 biovulate cells. Styles
erect, glabrous, about 7 mm. long. Fruit unknown.

This species is only known from the type locality in Amazonian
Peru.

PERU: Loreto: Mishuyacu, near Iquitos, alt. 100 m., forest, tree 10 m.,
flowers white, IV-1930, Klug 1130 (isotype, US); tree 12 m. high, flowers white,
II-III-1930, Klug 1091 (US).

10. Vantanea compacta (Schnizl.) Cuatr., comb. nov.
Fiaures 7,a—b; 10,c-e; PLate 1
Humirium compactum Schnizl. Abbild. Nat. Fam. 38:Ordo 222, pl. 222.

1843-1870.
Humirium contractum Moric. Msc. Schnizl. in Urb., Bot. Jahrb, 15, Beibl.

34:3. 1892.
Vantanea contracta Urb., ibid.
Vantanea panniculata Urb. in Mart. Fl. Bras. 12(2):450, pl. 96. 1877.

Type: Blanchet 3362, 3305, Brazil, Bahia.

Medium or large tree; branchlets slender, lustrous, densely lenti-
cellate, glabrous or slightly puberulous. Leaves small, coriaccous,
moderately rigid, glabrous or sometimes sparsely and appressed
puberulous. Petioles 3-6 mm. long, thickened at base. Blades
entire, oblanceolate or lanceolate-elliptic, shortly or longly cuneate
at base, attenuate, acuminate or long-cuspidate at apex, callous-
thickened and slightly revolute at margin, 3.5-7 (-9) em. long,
1.3-2.5 (-4) em. broad; shining above, midrib more or less marked,
other nerves almost inconspicuous; beneath with prominent midrib,
8-9 secondary nerves slightly prominulous on each side, lesser veins
loose and thin.

66 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Inflorescence cymose-paniculate, axillary, small, equaling the
leaves or shorter, axis and branchlets angled, minutely pubescent-
hirtellous. Bracts deciduous. Pedicels thick, minutely hirtellous,
0.5-1.5 mm. long. Calyx cupular, 1 mm. high, scarcely puberulous,
5-lobate; lobes subrotundate (0.3-0.5 mm. long), with minutely
ciliate margin. Petals white, linear-oblong, subobtuse, glabrous,
4-5(-8) mm. long, 1.5-1.8 mm. wide. Stamens about 50-60, filaments
slender, flexuous, 1.5-4(-7) mm. long, united in a 0.6 mm. high ring.
Anthers ellipsoid, about 0.7-0.9 mm. long, with 4 oblong, apiculate
lobes about 0.5-0.6 mm. long, the connective with acute, 0.3-0.4 mm.
long tip. Disk annular, girdling the ovary, glabrous, 1 mm. high
with strongly denticulate margin. Ovary ovoid, 1.5 mm. high,
minutely velvety-sericeous, with 5 biovulate cells. Style 2-3 mm.
long, erect, thickish, pubescent on the lower half. Stigma obtuse.
Drupe oblong-obovate-ellipsoid, rounded at apex, attenuate at base,
2.4-2.8 cm. long, 1.5—-1.8 cm. diameter; the exocarp smooth, 1-1.5 mm.
thick, coriaceous, brittle when dry. Endocarp woody, oblong,
ellipsoid, obtuse at apex, slightly narrowed at base, up to 2.5 X 1.4
em.; the surface rugose with 5 slight furrows toward the base and
5 longitudinal, oblong, germinal valves 1.8-2 em. long, slightly
conspicuous. Only one oblong sced in the single cavity observed.
The drupe may be smaller and globose-ovoid in a subspecies.

Usually the leaves are completely glabrous. Only in one specimen
(Blanchet 85) have I seen very sparse, minute, appressed hairs beneath.
The Ducke 1518 collection, which is preserved at the Museu Goeldi,
has some larger leaves (up to 9 X 4 cm.) than the regular type;
nevertheless, this collection agrees with the other characters of
V. compacta, despite the smaller leaves of the specimens. Blanchet
85 in Paris with regular, small leaves also has one large (8.5 & 4 cm.)
leaf. Ducke 1513 is the only collection of the species that I have
seen with fruit and is the basis for this part of the description.

V. compacta is restricted to the eastern States of Brazil, from
Cearé to Santa Catarina; however, the Pearce collections indicate the
presence of a new subspecies on the eastern slopes of the Bolivian Andes.

The specific epithet published by Schnizlein, is “compactum”’ and
is the correct and valid name for the species; “compactum”’ is used
several times in the detailed explanation of plate 222 of Schnizlein,
the explanation amounting to the original description. Urban
erroneously quoted “Humirium contractum” as the name published
by Schnizlein—a herbarium name presumably handwritten by
Moricand on a Martius herbarium sheet. (which is the type). But
for some unknown reason, when Schnizlein published his very fine
drawing and analysis of the species, he used the name “compactum,”’
and it therefore has priority.

CUATRECASAS—-HUMIRIACEAE 67

The specimens used by Schnizlein were reportedly from the Martius
Herbarium (‘“Saemmtliche Figuren nach Exemplaren aus v. Martius
Herbarium’), but this species is not among the humiriaceous collec-
tions received from Munich. Martius probably did not collect the
species himself, and the specimens that he used were those collected
by Blanchet. The type of V. panniculata, Blanchet 3305, must be
the first collection cited by him. The type of V. contracta (Moric)
Urban is Blanchet 3362, which is in the Paris Herbarium and was
originally labeled with name and analysis by Moricand and cited by
Urban. I think that this same specimen (Blanchet 3362) ought to be
considered the type of Humirium compactum Schniz.

10a(1). Vantanea compacta subsp. compacta var. compacta

BRAZIL: Bauta: Jacobina, Igreja Velha, Blanchet 8362 (holotype, P; isotypes,
Ga, NY); Blanchet 3805 (holotype of V. panniculata, P; isotype, NY).
Camandiea, Blanchet 3837 (P); Blanchet 85 (P). CrarX: Guaramiranga, matta
das collinas ao redor da ville; arvor pequena, 8-VIII-1908, frutos, Ducke 1513
(MG). Rio pe Janerro: Petrépolis ao Retiro, 5-IJI-1880, ‘‘Magaranduba de
folha miuda,” Glaziow 11829 (P); Corcovado, 4-II-1880, Glaziow 11828 (GH,
BM, IAN, NY, P). Alto Macahé, 7-IV-1891, arbrisseau, Glaziou 18182 (P,
US). Sio Pauto: Jardim Botanico, ‘“Aroeirana,” 10—X-1933, 17-III-1982,
Hoehne 29281 (A, GH, NY, 8, P). Santa CaTaRrtiNa: Mata da Azambuja,
Brusque, “guaraparin,” 23-II-1950, Klein 37b (8, US). Brusque, arvore 20
m., flor verde amarelada, “guaraparin,” 23-II-1950, Reitz 3353 (S, US).
Horto Florestal INP, Ibirama, 300 m. alt., mata, arvore 20 m. flor branca,
“ouaraparin,” 1-IIJ-1954, Reitz & Klein 1589 (US, 8, U). Morro da Fazenda,
Itajaf, 50-350 m., mata, “guaraparin,” arvore 20 m., 14-V-1954, Reitz & Klein,
1836 (US); 4-III-1954, Reitz & Klein 1730 (US); arvore, flor branco amarelada,
18-III-1954, Reitz & Klein 1744 (US, U, 8).

10a(2). Wantanea compacta subsp. compacta var. grandiflora (Urb.), Cuatr.,

comb. nov.
Vantanea contracta var. grandiflora Urb. Bot. Jahrb. Engler, Beibl. 34:3.
1893.

Type: Glaziow 16723, Brazil, Rio de Janeiro.

Flowers larger than ordinary; petals 7-8 mm. long; stamens about
80. Leaves puberulous with minute, sparse, appressed hairs beneath.
Inflorescences pauciflorous.

BRAZIL: Rio pe Janerro: Alto Macahé, 5-III-1888, Glaziou 16723 (isotypes
US, P). Alto Macahé, arbor, 11-III-1891, Glaziou 18181 (paratype P). Rio de
Janiero, IIJ-1891, Glaziow 14640 (US).
10b. Vantanea compacta subsp. microcarpa Cuatr., subsp. nov. PLATE 2

Arbor 10-20 m. alta ramis terminalibus tenuibus glabris. Folia
tenuiter coriacea flexibilia oblongo-elliptica basi longe angustata
cuneata apice paulo attenuata obtusiuscula. Petiolus 4-6 mm. longus.
Lamina 5-8 cm. longa 24.5 cm. lata costa praecipue subtus notata
nervis secundariis venulisque leviter conspicuis. Calyx cupularis
sepalis subrotundatis dimidia inferiore parte connatis glaberrimis 1
mm. longis. Petala crassiuscula oblonga sursum angustata subacuta

68 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

4-5 mm. longa 1.5-1.8 mm. lata, aestivatione contorta. Stamina
circa 58 glabra epapillosa filamentis flexuosis 1.5-3.4 mm. longa
tantum basi coalita. Antherae ovoideo-ellipsoideae circa 0.9 mm.
longa, apiculo 0.3-0.4 mm. longo acuto incluso. Discus annularis
argute dentatus 0.8-1 mm. altus. Ovarium ovoideum circa 1.5 mm.
altum minute velutino-sericeum. Stylus 2-2.5 mm. longus glaber.
Stigma anguste capitatum breviter 5-lobatum. Drupa globoso-
ovoidea 1.6-1.8 cm. longa 1.4-1.6 cm. lata basi rotundata apice
obtusissima vel subrotundata, exocarpio laevi in sicco 0.6-1 mm.
crasso. Hndocarpium subgloboso-ovoideum circa 1.7-X1.4 em.
rugulosum basi rotundatum apice obtusum, 5 valvis longitudinalibus
obovato-oblongis circa 1.4 cm. longis basim versus argute exculptis
instructum ; sectione 10 radiatum raro 5 cavitatis saepe tantum 1-2
cavitatis seminiferis monospermis evolutis.

Type in the Kew Herbarium, tree 70-80 ft. high, collected at
“Chailla” (probably Challa), Bolivia, May 1866, by R. Pearce. Para-
type in the Kew Herbarium collected at Moro, Yungas, Bolivia,
January 1866, by R. Pearce; isoparatype also in British Museum.

This subspecies is only known from the Bolivian type localities.

BOLIVIA: “Chailla” according to the collector, but probably ‘“‘Challa”’; tree

70-80 ft.; V-1866, Pearce s.n. (type, K). Yungas, Moro, alt, 500-600 ft., tree
40-50 ft., I-1866, Pearce s.n. (paratype, K, BM).

11. Vantanea macrocarpa Ducke, Arch. Inst. Biol. Veget. Rio Janeiro 1:205.
1935.—Trop. Woods 43:20. 1935. Figures 11,a—b; 12

Type: Ducke 20427, Brazil, Amazonas, Manaos.

Large tree with glabrous greenish brown, densely lenticellate
branchlets. Leaves thick-coriaceous, glabrous. Petioles 8-12 mm.
long, thick, sulcate above. Blades ovate-oblong or elliptic-oblong,
rounded or obtuse at base and decurrent upon the petiole, attenuate
and obtusely acuminate at apex, entire and thickened at the margin,
11-20 em. long, 5-10 em. broad, both sides brownish when dry;

Ficure 12.—Vantanea macrocarpa, X1 (Ducke 2230): a, fruit; b, endocarp

CUATRECASAS—HUMIRIACEAE 69

lustrous above, costa flat, secondary nerves slightly visible, lesser veins
obsolete; with thick midrib beneath, secondary nerves slender but
prominent, 16-18 on each side, subspreading arcuately connected
near the margin; minor veins few and obscure; some minute glands
more or less seriate, conspicuous.

Inflorescence cymose-paniculate, axillary and terminal, much
shorter than the leaves; the peduncle short, stout; the branchlets
thick, articulate, angled, minutely hirtellous. Pedicels thick, short,
about 0.5 mm. long, hirtellous. Bracts and bracteoles deciduous,
thickish, embracing, ovate, obtusiuscule, 2.5-1.5 mm. long, tomentu-
lose outside. Calyx cupular, about 1.5 mm. high, thick, pubescent-
hirtellous, lobate, the lobes subrotundate. Petals white, linear,
subacute, glabrous, about 1 cm. long, 1.5-1.8 mm. wide. Stamens
about 70, the filaments slender, minutely papillose-verruculose,
flexuous, unequal, 6-8 mm. long, at base united in a 2 mm. long tube.
Anthers oblong, about 0.9 mm. long, with 8 basal, ellipsoid lobes, the
connective fleshy, elongate, subacute. Disk cylindric 1.8 mm. high,
glabrous, with dentate-fimbriate margin. Ovary glabrous, 1.5 mm.
high, 5-loculate, cells biovulate. Style erect, glabrous, 7-8 mm. long.
Stigma obtuse. Drupe ellipsoid, rounded at both ends or obtuse at
apex, almost smooth, 4.6-10 em. long, 3.6-4.5 em. broad. Exocarp
coriaceous when dry, about 3 em. thick. Endocarp ellipsoid-ovoid,
rounded at base, obtuse at apex; the specimen seen 4 cm. long by 3 cm.
broad, deeply and anfractuosely alveolate-rugose, the rugosities con-
nected with about 10 large, irregular cavities. The opercula broadly
oblong, about 3.4 cm. long and 9 mm. broad, inconspicuous because
corrugations entangle their margins. Only one seed and one fertile
cavity in the specimen seen, oblong, 2.6 cm. long, 4 mm. thick.

Vantanea macrocarpa, usually a large rain forest tree with white
flowers, is known only from the uplands of the Manaos region in
Brazil. ‘“Uchy rana”’ is its common name.

BRAZIL: Amazonas: Manaos, silva terris altis ultra flores, arbor magna
floribus albis odoratis, 9-III-1937, Ducke 30133 (P, 8, U, US); Ducke 416 (A,
NY, 8S, US). Manaos, prope cataractas flum.; taruma silva non inundabili;
arbor magna floribus albis, 25-1 V-1932, Ducke 20427 (US, P, 8, US, RB, isotypes) ;
Ducke 21357 (Y); photos F.M. 35174 at Paris. Rio Taruma, mata da terra

firme perto da cachoeira alta; arvore grande, flor branca perfumada, 22-IJ-1949,
Ducke 2230 (IAN, MG).

12. Vantanea tuberculata Ducke, Arch. Inst. Biol. Veget. Rio Janeiro 4:30.
1938. Fiaures 13; 14,c

Type: Ducke 30134, Brazil, Amazonas.

Large tree; branchlets brownish, lustrous, glabrous. Leaves coria-
ceous, brownish when dry, glabrous. Petioles thick, 3-4 mm. long.
Blades entire, obovate-elliptic, cuneate at base, obtuse or rotundate

513359—61——-4

70 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

at the apex, 8-11 cm. long, 4.5-6 cm. broad; rather smooth above,
costa slightly distinct, smaller nerves almost obsolete; beneath with
prominent midrib, the secondary nerves about 10 on both sides,
slightly prominulous or almost inconspicuous, near margin arched-
connected, minor veins obsolete. Drupe ellipsoid, rounded at both
ends, 6.6-8 cm. long, 4.6-6 cm. broad. Exocarp very thick, strongly
tuberculate, coriaceous when dry, 8-10 mm. thick. Endocarp heavy,
hard, woody, about 5 cm. long, 3.5 cm. broad (in the specimenseen),

Ficure 13.—Vantanea tuberculata, X1 (Ducke 30134): a, fruit; b, endocarp, apical view;
c, transection,

strongly and deeply anfractuose-rugose, inside with 10 irregular empty
cavities (5 in the valves, 5 larger alternating). The 5 opercula (or
valves) longitudinal, oblong, subacute at apex. Often only 1 to 3
seeds well developed, oblong, protected by the opercula.

V. tuberculata is known only from its type locality in Amazonas,
Brazil. The flowers of the species are unknown, but the structure of
the fruit leaves no doubt as to its inclusion in Vantanea. The endo-
carp is like that of V. macrocarpa, but the thick warty pericarp
distinguishes V. tuberculata from all other species of Vantanea.

BRAZIL: Amazonas: S&o Paulo de Olivenga, Rio Solimées, silva non inun-
dabili, arbor magna, 6-II-1937, Ducke 30134 (isotypes, US, U).

CUATRECASAS—HUMIRIACEAE 71

13. Vantanea guianensis Aubl. Pl. Guian. 1:572. 1775.—Benth. in Hook.
Journ. Bot. Kew Mise. 5:98. 1853.—Urb. in Mart. Fl. Bras. 12(2) 3452,
1877.—Ducke, Arch. Jard. Bot. Rio Janeiro 3:179. 1922; 5:14, pl. 17,
figs. 42a, 42b. 1930.—Arch. Inst. Biol. Veget. Rio Janeiro 4:30, 31. 1937.

Figures 14,a-b; 15; PLate 3
Lemniscia guianensis Gmel. Syst. Nat. 817. 1791.
Lemniscia guianensis Raeuschel Nomencl. Bot. ed. 3; 156. 1797.
Lemnescia floribunda Willd. Sp. Pl. 2:1172. 1800.
Lemniscia floribunda Sprengel, Syst. 2:600. 1825.

Type: Aublet, French Guiane, Comté de Géne.

Large trees with brownish-green, glabrous, subterete lenticellate
branchlets. Leaves coriaceous, flexible, glabrous. Petioles 6-12 mm.
long, flat and sulcate above. Blades entire, elliptic or oblong-elliptic,
obtuse-cuneate at base and decurrent on the petiole, attenuate, shortly
and obtusely acuminate at apex, 6-14 cm. long, 2.7-6 cm. broad; more
or less lustrous above with prominent and thin costa, secondary nerves
slightly prominulous, lesser veins few and almost inconspicuous;
beneath with a thick midrib, 13-15 pairs of prominulous, slightly
ascendent arcuate-anastomosate secondary nerves, the minor veins
reticulate, less conspicuous.

Inflorescences axillary and terminal, shorter than the leaves,
cymose-paniculate, subcorymbiform, rich in flowers, the peduncle
stout, short, subterete, glabrous, the branchlets angled, glabrous.

Ficure 14.—a, Vantanea guianensis, X ¥3 (Ducke 200); b, Vantanea guianensis, bud, X13; ¢,
Vantanea tuberculata, leaf, X% (Ducke 30134); d, Vantanea parviflora, X V% (Ducke
23425).

72 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Bracts soon deciduous, embracing, ovate, about 1 mm. long. Pedicels
rather thick, smooth, glabrous, 2-5 mm. long. Calyx thick, deep-
cupular about 4 mm. long, glabrous, slightly dentate with obtuse teeth,
glandular outside. Petals red or purple, linear, acutish, glabrous,
about 3 cm. (up to 4 cm.) long, 2.5 mm. broad. Stamens 64-80,
glebrous, filaments flexuous, 25-32 mm. long, united at base in 5-7
mm.long tube. Anthers 1 mm. long, oblong-ellipsoid, lobes 4, oblong,
0.8 mm. long, the connective rather thick with a short, acute tip.
Disk tubular, thick, glabrous, 1.5 mm. high with almost smooth
margin, encircling the ovary. Ovary ovoid, about 2 mm. high,

Z\MMERMANN —

Ficure 15.—Vantanea guianensis, X 1 (Ducke 200): a, fruit; b, endocarp; c, transection.

glabrous, 5-celled, the cells biovulate. Styles flexuous, glabrous, 30
mm. long or longer. Stigma obtuse. Drupe ellipsoid-ovoid, almost
smooth, about 6 cm. long, 4.5 em. broad. Exocarp carnose, 3-5 mm.
thick and coriaceous when dry. Endocarp 53.8 cm., hard, woody,
ovoid-ellipsoid, broadly rounded at the base, abruptly and obtusely
apiculate-contracted at the apex, deeply anfractuose-rugose with
encephalouslike aspect; five longitudinal, deep and narrow furrows.
At the bottom of each furrow a hidden longitudinal, long, narrow
operculum (valve). Inside irregular empty cavities alternate with
the furrows. The seeds oblong, about 3.2 cm. long and 0.3 em. thick,

CUATRECASAS—HUMIRIACEAE 73

only 2 or 3 developed, at bottom of the grooves, protected by the
opercula.

V. guianensis is a medium-size or large tree of humid forests on
elevated land, displaying inflorescences of large crimson flowers. Its
fruit are of the size of an apricot with a fleshy, insipid, scarcely fatty
mesocarp and a hard and corrugated endocarp. According to Schom-
burgk, these endocarps cut in halves are used as neck ornaments by
Guiana Indians. Aublet quoted the native name ‘Touantan” as the
Caribbean name for the species, on which he based the generic Latin
name. Urban (p. 453) gave the Caribbean name the spelling ‘‘Schuan-
tan.” In Brazil it is known as “uchi-rana” or “achud-rana.”’

The species is widely distributed in Venezuela, British and French
Guiana, and throughout the middle and lower Amazon Valley.

VENEZUELA: Amazonas: Rio Casiquiare, Rfos Pacimoni-Yatua, occasional
along Rio Yatua, 20-25 km. above Piedra Arauicana, tree 15-25 m., flowers
brilliant pink, 30-IX-1957, Maguire, Wurdack, & Maguire 41640 (NY, US).

BRITISH GUIANA: Roraima, 1842-43, Schomburgk 982 (BM, NY, P).
Corentyn River, Schomburgk 1581 (BM, US). “British Guiana” ex Herb. Ben-
thamianum, Schomburgk 47 (K).

FRENCH GUIANA: Gourdonville, 2-VIII-1914, Benoist 1530 (P). Without
locality Melinon 100 (US). ‘Habitat in America meridionali, specimen e collect:
Aublet: misit Gener: Barron: Dans Banks, a: 1782”; this Aublet specimen may be
considered as the holotype (8).

BRAZIL: Amazonas: Manaos, Estrada do Aleixo, silva non inundabili; arbor
magna floribus roseo purpureis, 20-V-1936, “uchi-rana,”’ Ducke 200 (IAN, NY,
S, US, A). Manaos, silva non inundabili, arbor mediocris, floribus laete purpureis,
31-III-1932, Ducke 23814 (U, S, US). Manaos, beira do Rio Tarumé, t.f. baixa;
arvore 10 m., flores vermelhas, vistosas, sem odor, 7-VIII-1949, Frées 24936
(IAN). Par: Belém, cultivada no IAN; arvore grande, flor muito vistosa, roseo
avermelhada, 25-VIII-1953, Pires 4518 (IAN, NY, U, US). Santa Isabel ad
viam ferream Belém-Braganca, silva humosa; arbor magna floribus coccineis,
10-IX-1922, Ducke 17783 (U). Peixeboi (Belém-Braganga), matta, 21-X-1907,
‘uchi-rana,” Siqueira 8775 (MG, BM, US, P). Carapara (Belém-Braganga),
matta, 6-LX-1908, Museu Goeldi 9664 (MG, BM, P, US). Utinga, Belém, 13-
VIII-1914, Ducke 15451 (MG). Utinga, 13-VIII-1914, Ducke 15415 (MQ).
Belém, Estrada Providencia, Tapand, matta da terra firme, arvore grande, flor
carmesin, 31-X-1944, “uchi-rana,’”’ Ducke 1647 (IAN). Breves, transecto para
inventario forestal 101-1, 7-30-VII-1956, Pires, Frées, & Silva 5380 (IAN). Par4,
sine loco, Jobert 362 (P). Rocky riverside below Fiafun Mount by falls Mapueva
River, headwaters Acarai Mountains; 8 in. X 46 ft. leaning tree with rough bark;
orange brown }4 in. slash yellow splintery wood; leaves alternate, ovate, lanceolate,
glossy, leathery; recurved terminal inflorescences of bright rose pink flowers with
linear petals to 1 in. recurved, caducous and stamens of same colour to 1} in.;
flower shoots erect; leaves hanging; fruit (young) green, ovoid, with terminal
point to which persistent stigma attached, after rested in by ants, Forest Dept.
G. 600 (NY). Distrito Acard Thomé Assu up Rio Acardé, 5 km, river bank in
virgin forest, alt. 35 m., tree 18 m. high, bright cerise flower standing above crown,
very ornamental, 5-VIII-1931, Mexia 6049 (A, BM, GH, NY, U, 8, US). Rio
pE JANEIRO: Restinga de Cabo Frio, 29-XII-1877, Glaziou 10078 (P).

74 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

14. Vantanea parviflora Lam. Journ. Hist. Nat. par. 1:145, pl. 7. 1792—
Urb. in Mart. Fl. Bras. 12(2):454. 1877. Ficures 5,a—e; 7,c-d; 14,d
Vantanea cupularis Huber, Bol. Mus. Goeldi 6:83. 1910.—Ducke, Arch.
Jard. Bot. Rio Janeiro 3:179. 1922.—Arch. Inst. Biol. Veget. Rio Janeiro

4:29. 1937.

Type: Le Blond, French Guiana.

Large tree with subterete, lenticellate, glabrous, rarely puberulous
branchlets. Leaves coriaceous, flexible, glabrous or sometimes
appressed-pilose, or midrib hirtellous beneath. Petiole semiterete,
sulcate above 5-15 cm. long. Leaf blades entire, elliptic, more or less
elongate, obtusely cuneate or cuneate at base, obtuse, emarginate,
obtuse-acuminate or even subrotundate at apex, 5-10 em. long, 2-5.5
cm. broad; greenish-brown or olivaceous above with distinct flat
midrib and prominulous secondary and smaller nerves; beneath
more or less lustrous, cuperous-brownish with prominent midrib,
9-10 slender but prominent ascendent secondary nerves on each
side, arcuate and united near the margin, smaller veins reticulate,
prominulous.

Panicle cymose-dichotomous, corymbiform, axillary or terminal,
shorter than the upper leaves; the axis and branchlets more or less
compressed, pubescent-hirtulous. Bracts ovate, subobtuse, minutely
pubescent, soon deciduous, 1 mm. long. Pedicels about 1-2 mm. long,
moderately thick, minutely pubescent-hirtulous. Calyx broadly
cupular scarcely 1 mm. long, 2.5 mm. diameter, puberulous outside,
the margin ciliolate, entire or slightly undulated. Petals white, linear,
rather thick, 7-8 mm. long, pubescent outside, hairs abundant, retrorse,
subappressed, glabrous inside, estivation contorted. Stamens 80-
120, filaments glabrous, 2-3 seriate, united at base, unequal, 5-7 mm.
long. Anthers about 0.8-1 mm. long, dorsifixed with 4 elliptic-oblong
lobes, the connective with a long, thickish, acute tip. Disk annular,
thick, densely hirto-tomentous, 1-1.5 mm. high, girdling the ovary.
Style columnar, about 3.5 mm. long, subglabrous, only some long,
spreading hairs near its base. Ovary 2.5 mm. high, ellipsoid, villoso-
hispid with long hairs, with 5 biovulate cells. Stigma obtuse, glu-
tinous. Drupe ellipsoid, 2.5-2.8 cm. long, 2.2-2.5 em. broad, becoming
glabrous; the endocarp woody like a nut, rugose, 2.1-2.5 cm. long,
1.8-2 cm. broad, with 5 longitudinal broad ribs and valves.

One specimen from French Guiana, Wachenheim 179 (P), has
globose drupes and endocarp. The drupe is about 24 mm. in diameter,
with a 1-2 mm. thick ectocarp. The endocarp is densely ligneous,
rugose, with 12 sulci, and 6 longitudinal valves. The specimen seen
has only two seminal cavities. This specimen and another closely
related one from French Guiana do not show any other difference from
the Brazilian specimens, for which reason they are considered con-
specific.

CUATRECASAS—-HUMIRIACEAE 15

The typical V. parviflora is completely glabrous except for the
inflorescence. Only a few specimens from Brazil show appressed,
sparse hairs on the underside of the leaf or a slightly hirtellous pubes-
cence on the midrib and petiole; these forms are transitional toward
the variety puberulifolia.

This large or medium-size tree is much spread out in the un-
inundatable rain forests of French Guiana, the Brazilian States of
Para and Amazonas, and the Venezuelan State of Amazonas.

14a. Vantanea parviflora var. parviflora

VENEZUELA: Amazonas: Rio Negro, San Carlos and vicinity, tree, flowers
white, 9-XII-1947, Schultes & Lépez 9267 (COL, IAN, US).

FRENCH GUIANA: “Herb. de la Guyane francaise, année 1863, boise géelette
rouge,” Melinon 100 (BM, P). “Guyane francaise,’ Melinon s.n. (US, several
specimens in P). Arbor 30-40 pedalis in sylvis Guyannae, ex. herb. Claude Richard
sn. (P). Guyane ex Herb. Maire, Herb. Cosson (P). Cayenne, photo F.M.
12592, without collector (from Berlin-Dahlem). Guyane fr. 18-XI-1921, Godebert
Wachenheim 179, (P); 23-VI-1921, G. Wachenheim 489 (P). Route de Saint
Laurend a Cayenne km. 13; terrain sain sablonneux inonde & la raison des pluies,
fleurs blanches, 6-VIII-1953, “koko” (Paramaka), “‘gris-gris-bois gaulette,”’
BAFOG 48M (P). Route de Charvein 4 |’Acarouany km. 3 céte sud et 420 m.
de la route “terrain sain.” Fruits verts ovoides forme et grosseur d’une petite
olive, recouverts d’une pubescence blanchatre par petites grappes 4 la pointe des
rameaux, “boliquin” (Pamaka), “gris-gris rouge,” BAFOG 102M (P). Route
Charvain-Acarouany km. 1; plateau argileux, 10 m. audessus du niveau de l’eau;
fleurs non épanouies petites, boutons rosatres, ovales, 1-2 mm., en bouquet a
Vextremité des rameaux, “adougoue” (Paramaka), 24-I1X-1954, BAFOG 220M
(U). Cédte gauche et 4 10 m., plateau peu élevé, environ 5 m., sabro-argileux assez
profund, fleurs blanchitres en grappes 4 Vextremité des rameaux, étamines
nombreuses, BAFOG 247—-M (U).

BRAZIL: Amazonas: Manaos, Estrada do Aleixo, silva non inundabili, arbor
sat magna floribus albis odore forti, mata da terra firme, 3-IV-1936, 31-VII-42,
Ducke 157 (A, IAN, MG, NY, 8, US). Manaos, silva non inundabili versus
flores; arbor media floribus albis odore forte, 8-VIII-1929, Ducke 23426 (P, S,
U, US). Manaos, Igarapé da Cachoeira Baixa do Tarumd4, terra arenosa, ca-
poeira; arvore 15 m., flor branca, 21-IX-1955, INPA 1809 (MG 21541). Rio
Urubt, igarapé Sangana; terra firme, alta, floresta central, arvore 18 m. X 35
em., 2-X-1949, Frées 25459 (IAN). Rio Urubt, Sao José das Pedras, terra
firme, alta, floresta alta; arvore de 18 m. X 35 cm., flores brancas, 11-IX-1949,
Froes 25185 (IAN). Basin Rio Juru4, near mouth of Rio Embira (tributary of
Rio Tarauaca) tree 110 ft. high, on terra firma, 21-VI-1933, Krukoff 4956 (A,
BM, M, NY, 8, U, US). Rio Solimées, Sao Paulo de Olivenca, arbor sat magna
floribus albis odore fortissimo, 19-VIII-1929, Ducke 23429 (P, 8, U, US). Basin
Rio Madeira, Municipality Humaytd, on plateau between Rio Livramento and
Rio Ipixuna, tree 20 ft. on campinarana alta, Krukoff 7120 (IAN, NY, §, US, U).
Part: Santa Rosa (Belém-Braganga), matta, “uchi-rana,”’ 10—-X-1908, Museu
Goeldi 9723 (MG, BM, P, US). Moéma ad viam ferream Belém-Braganga, 30-
VIII-1908, Huber 9583 (S, U, isotype of Vantanea cupularis Huber); photo
F.M. 12588 at Berlin-Dahlem. Moéma, matta, 17-IX-1908, Museu Goeldit
9670; 30-VIII-1908, Museu Goeldi (Huber) 9583 (P, US). Utinga (Belém-
Braganca), silva non inundabili; arbor magna, floribus albis, matta da terra
firme, 27-VIII-1941, Ducke 781 (IAN, MG, US). Santa Isabel, Belém-Braganga,

76 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

27-IX-1908, Museu Goeldi 9680 (B, US, MG). Utinga-Providencia, 28-VIII-
1914, Ducke 15467 (MG). Belém, Horto Mocambo (Reserva Florestal) » arvore,
VIIT-1951, Pires s.n. (US). Perto do Igarapé Arapij6 municipio de Breves,
2-1-1955, 7-30-VII-1956, Pires, Frées, & Silva 5105, 4954 (IAN). Obidos,
silvis non inundatis, inter lacuna Mamaurt et Castanhal do Prior; arbor media
floribus albisfodoratis, 14-IX-1926, Ducke 20426 (U, 8, US). Juruty Velho,
silva non inundabili, arbor sat magna, 24-XII-1926, Ducke 20428 (US). Trom-
betas, entre Cumindmirfm e Ariramba, 12-X-1913, Ducke 14962 (MG, BM, P,
US). Region Boa Vista, hab. terra firme, “uxy bravo,” 10-X-1932, Capucho
483 (IAN). Without locality, Herb. John Miers 6167 (BM).

14b. Vantanea parviflora var. puberulifolia Cuatr. var. noy.

Cum typo differt: Folia subtus sparse adpresse pilosula. Petiolus
costaque subtus breviter hirtula. Ramuli terminales hirto-puberuli.

Type in the U.S. National Herbarium, No. 1518624, collected in a
nonflooded forest at Fonteboa (Solimées River), State of Amazonas,
Brazil, September 6, 1929, by A. Ducke, No. 23428. Isotypes in
Museum National d’Histoire Naturelle, Phanerogamie, Paris, and
in Naturhistoriska Museum, Stockholm.

This variety is characterized by its appressed puberulous leaves
beneath and short-hirtellous midrib, petioles, and young branchlets.
The typical form of the species has completely glabrous leaves and
branchlets. Only a few specimens from Brazil are very shortly and
scarcely puberulous on the underside of the leaves or have the midrib
puberulous (Ducke 23429, 20428, Schultes &: Lépez 9267, Krukoff 4956,
7120, Museu Goeldi 9723); they are intermediate forms of this variety.
Ducke 23428 has the apical appendix of the anthers more slender and
longer (up to twice as long as the thecae), but this condition is not
characteristic of the variety because examination of many specimens
shows a great variability.

BRAZIL: Amazonas: Rio Solimées, Fonteboa, Ducke 23428 (type). Manaos,
Silva non inundabili; arbor magna floribus albis odore fortissimo, 10-III-1929,
Ducke 23427 (US). Manaos, prope Cachoeira Grande, silva non inundabili;

arbor sat magna, floribus albis, odore fortissimo, 8-VIII-1929, Ducke 23425
(US, S, U).

Tribe Humirioideae

Humirioideae Cuatr., tribus nova Humiriacearum.

Stamina definita (10-30) antheris 2-4 thecis unilocularibus dis-
junctis, basim abscisso-dehiscentibus. Genus typicum Humiria
(Aubl.) St. Hilaire.

2. Duckesia
Duckesia Cuatr., gen. nov.
Sepala suborbiculata basi coalita imbricata. Petala 5 libera
oblonga praefloratione contorta vel cochleari. Stamina 20-25 fila-
mentis subulatis paulo compressis papillosis inaequalibus basi in

CUATRECASAS—HUMIRIACEAE 77

tubum brevem coalitis. Antherae lineari-lanceolatae dorsifixae,
thecis 4 unilocularibus disjunctis globosis vel globoso-ellipsoideis
duobus basilaribus duobus subparte media utroque latere instructis
connectivo acuto, tantum 5 oppositipetalae fertiles. Discus squamis
10 liberis subulatis crassiusculis. Ovarium ovoideum 5-loculare
loculis episepalis uniovulatis, ovulis anatropis raphe ventrali.
Stylus stamina attingens stigma anguste capitato breviter 5-lobato.
Drupa ovoidea sublaevis exocarpio crasso. Endocarpium spongioso-
lignosum resinoso-lacunosum verrucosum subapicem 5-foraminatum
lateraliter longitudinaliterque 5-valvatum. Semina oblonga. Ar-
bores foliis simplicibus persistentibus coriaceis vel subcoriaceis
petiolatis. Inflorescentiae axillares breves cymoso-paniculatae plus
minusve dichotomae. Typus: Sacoglottis verrucosa Ducke.

Sepals 5, suborbicular, imbricate, united in a cup. Petals 5, free,
linear-oblong, thick-membranaceous, the estivation contorted or
cochlear. Stamens 20-25, biseriate, glabrous, the filaments united
at base, complanate, subulate, papillose, alternating in three dimen-
sions. Occasionally some shorter staminodia present. Anthers
linear-lanceolate, glabrous, dorsifixed, usually only 5 fertile; the
connective lanceolate, carnose; the thecae 4, unilocular, dissociated,
subglobose or ellipsoid, 2 attached at the base, the other 2 laterally at
the middle, dehiscing by accidental pulling away. Disk formed by
10 subulate free scales surrounding the ovary. Ovary glabrous,
5-locular, each cell with a single anatropous, pendulous ovule. The
carpels opposite the sepals. Style erect, as high as the stamens.
Stigmas forming a 5-lobate head. Drupe large, ovoid, almost smooth,
with thick exocarp, coriaceous and fragile when dry. Endocarp
strongly verrucose, spongy-lignose, resinous-lacunose, five-foraminate
at the apex, laterally with 5 longitudinal oblong germinal valves or
opercula. Seeds few, oblong. Evergreen trees with alternate, thin-
coriaceous, petiolate, serrate leaves; the teeth ending with an erect
deciduous gland. Inflorescences axillary, small, cymose-paniculate,
furcate below, upwards alternate branching. Bracts persistent.

Duckesia differs from Sacoglottis in the structure of the anthers
with a long connective and four unilocular thecae, two of them being
at the base, the other two being higher up and lateral, and in the
valvate endocarp. The endocarp is strongly rugose-tuberculate,
somewhat corky-woody, and provided with five longitudinal opercula
as is typical in Vantanea. From Vantanea if differs in the four uniloc-
ular thecae of the anthers, the number of stamens, and the spongy
structure of the endocarp full of small resinous cavities. The fertile
stamens, which are opposite the petals, originate from an inner
whorl, which is concrescent with the outer whorl. (See figs. 3, 16, 19.)

This genus is named in honor of Adolpho Ducke, the discoverer

78 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

of the plant and the most outstanding Amazonian botanist and plant-
explorer of this century.

Duckesia has only one species, which is limited to Amazonian
Brazil.

1. Duckesia verrucosa (Ducke) Cuatr., comb. nov.

Sacoglottis verrucosa Ducke, Arch. Jard. Bot. Rio Janeiro 3:177, pl. 10b.
1922; 5:142, pl. 14 figs. 35a, 35b. 1930.—Arch. Inst. Biol. Veget. Rio
Janeiro 4:26. 1987. Figures 16,9-k; 18,b; 19,d—e

Type: Ducke 10815, Brazil, Pardé, Obidos.

Tree with thin, minutely hirtellous-pubescent terminal branchlets.
Leaves small, thin-coriaceous, sessile or subsessile. Petiole 0-1.5
mm. long, rather thick, pubescent. Blade lanceolate, cuneate,
sessile or narrowed to a very short petiole at base, acuminate or
cuspidate at apex, serrate-crenulate at margin, 3-8 cm. long, 1-2.3
cm. broad; above smooth, glabrous or when young with minute hairs
on the conspicuous midrib, the other nerves obsolete; the midrib

Ficure 16.—a-f, Endopleura uchi (Baker 58): a, petal; b, bud; c, bud with 2 front petals
removed, X 5; d, detail of stamens, X 10; ¢, gynoecium and disk, X 10; f, longitudinal
section of ovary X10. g-k, Duckesia verrucosa (Ducke 16325): g, petal, h, bud; 7, bud
with frontal petals removed, X 5; 7, details of stamens and anthers with 2 sterile anthers
right, X10; k, gynoecium and disk, «10. L-p, Hylocarpa heterocarpa (Pires 708); l,
open old flower (with fallen anthers), 5; m, stamens X 10, 2 sterile anthers at right; n,
transection of ovary, X10; 0, gynoecium and disk, X10; p, position of ovules in the
ovary, X5.

CUATRECASAS—-HUMIRIACEAE 79

beneath prominent and minutely pubescent or puberulous, elsewhere
glabrous or scarcely pilose, the 12-14 pairs of secondary nerves
slightly prominulous, ascending, near the margin arcuate-anastomos-
ing, the minor nerves almost obsolete; when young the teeth end with
a linear erect deciduous gland 0.4 mm. long.

Inflorescence axillary, much shorter than the leaves, cymose-
paniculate, divaricate, the branchlets thin, usually alternate, furcate
below. Peduncles and branchlets minutely hirtellous-pubescent.
Bracts ovate-lanceolate, amplectant, 1-0.5 mm. long, puberulous.
Pedicels thick, 0.4-0.5 mm. long, articulate with short peduncles or
sessile. Sepals suborbicular, about 0.6 mm. long, minutely ciliate
at margin. Petals lincar-oblong, subacute, glabrous, about 3.5 mm.
long, 1 mm. broad. Stamens 20-25, glabrous, the filaments 1.7-2.2
mm. long, complanate, subulate, acute, minutely papillose, united
at base; the 5 opposite the petals larger. Anthers linear-lanceolate,
elabrous, 1 mm. long, dorsifixed, only 4-5 of them (those of the
larger filaments) fertile; of 4 thecae 2 short-ellipsoid, basal, the other
two oblong-ellipsoid, lateral on the middle; the connective lanceolate,
subtriquetrous, carnose; the other 15-20 anthers sterile. Some
shorter additional staminodial filaments may also be present. Style
columnar, glabrous, about 1.5 mm. long. Stigma subcapitate-
pyramidal, slightly 5-lobate. Scales of disk 10, subulate, rather
thick, 0.5 mm. long, free. Ovary ovoid, glabrous, about 1 mm.
high, furrowed, 5-locular, the cells uniovulate, the ovules ellipsoid-
oblong, 0.6 mm. long. Drupe short-ovoid or subglobose, about
7 cm. long and 6 cm. in diameter, almost smooth; exocarp 10-13 mm.
thick. Endocarp (Ducke 16764 in US) ovoid, abruptly acute at
apex, 3.5 cm. in diameter, spongy-lignose, densely resinous-lacunose,
acutely verrucose, 5-foraminate below the apex, conspicuously
5-valvate lengthwise. The seeds oblong, 2 seen in the specimen.

Duckesia verrucosa is a large tree of the rain forest of firm land in the
middle Amazonian region. It has been discovered and collected only
by Adolfo Ducke, who writes about it thus: “This species produces
the fruit ‘uchy-curua’ or ‘uchy-coroa’ (‘uchy’ meaning warty, referring
to the protuberances of its stone), which can be found on the markets
of Manaos and Obidos where it is eaten raw. Wood brown-gray
purplish, rather hard and dense, with no known uses.”

BRAZIL: Amazonas: Manaos, Estrada do Aleixo, mata das terras altas
argilosas; arvore muito grande, flor verde con estames amarelos, fruto comestivel,
‘“uchi curtia,” 23-VIII-1947, Ducke 2108 (IAN, MG, US, PU). Tapajoz medio,
Cachocira do Mangabal, “‘uchi-curtia,” 13-JI-1917, Ducke 16764 (MG, US).
Park: Obidos, matta da terra firme, “uchy-curta,”’ 18-X-1913, Ducke 14992

(MG). Obidos, silva non inundabili, arbor magna floribus viridibus odoratis,
‘“uchy-curda,’” 11-VIII-1916, Ducke 10815 (isotypes, NY, S, U). Same data,

80 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Ducke 16325 (MG number; holotype, MG; isotypes, PS), photo F.M. 12603
from Berlin. Territorio do Guaporé, VIII-1952, Silva 416 (IAN).

3. Endopleura
Endopleura Cuatr., gen. nov.

Sepala 5 orbicularia imbricata basi coalita. Petala 5 libera oblonga
praefloratione contorta vel cochleari. Stamina 20-30 filamentis
crassiusculis leviter angulatis papillosis biseriatis inaequalibus 5
longioribus oppositisepalis inferiore parte coalitis, Antherae con-
nectivo crassiusculo elongato acuto, thecis 4 disjunctis subglobosis
2 ad basim utroque latere 2 lateralibus supra basim, interdum 2 vel
omnibus thecis sterilibus. Discus 10 squamis anguste ovato-tri-
angularibus basi coalitis. Ovarium subelobosum 5-loculare loculis
episepalis uniovulatis ovulis anatropis raphe ventrali. Stylus brevis.
Stigma capitatum 5-lobatum. Drupa ellipsoidea exocarpio fibroso-
farinaceo; endocarpio lignoso eminente 5-alato-costato costis basi
excepto bifidis cum sectione figura 10-radiata, 5-valvato ; Mesocarpio
inter costas dense fibroso. Semina oblonga 5 vel saepe tantum 1-3
evoluta. Arbores foliis simplicibus alternis persistentibus coriaceis
vel subcoriaceis petiolatis. Inflorescentiae axillares cymoso-panicu-
latae bracteatae bracteolataeque. Typus: Sacoglottis uchi Huber.

Sepals 5 suborbicular, imbricate, united at base. Petals 5, free,
oblong, thick, the estivation contorted or cochlear. Stamens 20-30
biseriate, glabrous; the filaments thick, angulate, papillose, united at
base, alternating in different dimensions, the largest opposite the
sepals. Anthers dorsifixed with thick, elongate, acute connective,
the thecae 4, unilocular, dissociated, subglobose, 2 attached at base,
other two lateral above the base, sometimes 2 thecae or rarely all 4
sterile; irregular dehiscence when pulling away the sacs. Disk formed
by 10 ovate-triangular, thick scales, united at base. Ovary glabrous,
suborbicular, 5-locular, cells uniovulate, ovules anatropous, pendant
with ventral raphe. Carpels opposite sepals. Style short. Stigmas
lobate-capitate. Drupe large, ellipsoid, with exocarp fibrous-farina-
ceous, coriaceous when dry. Endocarp woody, with 5 apical foveola,
with 5 ribs strongly elevated and divided into 2 except at the end,
forming 10 strong, woody wings which make in section a 10-ray stellate
figure; the mesocarp fills the space between the ribs with a compact
fibrous structure; inconspicuous linear, germinal valves between the
branching ribs at bottom of the furrow. Seeds oblong, rarely all 5
developed, usually only 1-3 becoming ripe. Evergreen trees with
alternate, coriaceous, serrate, petiolate leaves. Inflorescences axillary,
cymose-paniculate, with trichotomous or dichotomous branching.
Bracts persistent. (See figs. 3and 16-18.)

CUATRECASAS—HUMIRIACEAE 81

Endopleura differs from Sacoglottis in the four unilocular thecae of
the anthers, of which two are basal and the other two located above
and laterally, and in the deeply sulcate endocarp, which has five
eminent longitudinal bifid winged ribs. The transversal section of
Endopleura looks like a 10-ray star. Furthermore, the endocarp has
narrow, shorter, inconspicuous valves and lacks resinous cavities.
Endopleura differs from Vantanea in the structure of the anthers, the
number of stamens, and the shape of the endocarp.

The name “Endopleura” is derived from the Greek “endon”’ (inside)
and “pleuron” (rib), the latter referring to the strong ribs of the
endocarp.

This genus has only one species, which is endemic to the Amazon
basin from Manaos to Belém.

1. Endopleura uchi (Huber) Cuatr., comb. nov. Fraures 16,a-f; 17; 18,a

Sacoglottis uchi Huber, Bol. Mus. Paraense. 2:489. 1898.—Bol. Mus.

Goeldi, Paré 4:395, 1904.—Ducke, Arch. Jard. Bot. Rio Janeiro 3:177

1922; 5:142, pl. 14 figs. 34a, 34b. 1930.—Arch. Inst. Biol. Veget. Rio
Janeiro 4:26, 1937.

Type: Huber 1260, Brazil, Para, Belém.

Large tree with glabrous, green, more or less angulate, compressed
young branchlets. Leaves coriaceous, firm or flexible. Petiole 1-2.3
cm. long, semiterete, pubescent or glabrous, sulcate above, more or

Figure 17.—Endopleura uchi, X1 (Ducke 305): a, fruit; b, endocarp; ¢ transection.

less winged at margin. Blade elliptic-oblong or elliptic-lanceolate,
abruptly and obtusely cuneate at base, narrowed-acuminate, often
cuspidate at apex, serrate and slightly thickened at margin, 8-20 cm.
long, 2.3-8 cm. broad; lustrous above with conspicuous lighter midrib,
glabrous or with very minute hairs toward the base, the secondary
nerves pale, prominulous, the veins conspicuously prominulous-
reticulate; dull, glabrous beneath with thick midrib, the 12-14 pairs
of secondary nerves thin, prominent, patulous and curvate-anas-
tomosing, the prominulous veins forming a very conspicuous reticulum.

82 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Inflorescences cymose-paniculate, axillary, much shorter than the
leaves; the 1-3 cm. long peduncles and branchlets compressed,
hirtellous or puberulous, trifurcate below, dichotomous above.
Bracts persistent, ovate, amplectant, about 2 mm. long. Bracteoles
ovate, subacute, hirtellous-puberulous, minute, about 0.5 mm. long,
persistent. Pedicels rather thick, hirtellous-pubescent, 0.2-0.5 mm.
long, articulate with 0-1 mm. long, pubescent peduncles. Sepals
about 0.7 mm. long, orbiculate, hirtellous-pubescent, united at base.
Petals greenish, rather thick, linear-oblong, subacute or subobtuse,

Figure 18.—a, Endopleura uchi (Ducke s.n., US 1693003), floriferous branchlet, x; 2b,
Duckesia verrucosa (Ducke 2108), floriferous branchlets, X 4; ¢, Hylocarpa heterocarpa
(Pires 708), leaf, X14.

glabrous inside, hirtellous-pubescent outside, 3-3.5 mm. long, 1-1.4
mm. broad. Stamens 22-30 (mostly 25), the filaments 1.5-2 mm.
long, thick, angulate, denscly papillose, united in their 4% lower
part, different lengths alternating, the longest with larger anthers
opposite the sepals. Anthers about 0.9 mm. long with 4 globose-
elliptic, 0.2 mm. long, 2 basal and 2 inferior-sided thecae, the con-
nective lanceolate, acute, about 0.7 mm. Jong. Sometimes 2 of the
thecae of the anthers sterile, rarely all 4. Scales of the disk 10, about
0.6 mm. long, triangular, united at base. Ovary glabrous, subglobose,

CUATRECASAS—HUMIRIACEAE &3

0.9-1 mm. high, 5-locular, the cells uniovulate, opposite the sepals.
Style stout, longer than the ovary. Stigma capitate 5-lobate. Drupe
oblong-ellipsoid, 4-6 cm. long, 2-3.8 cm. in diameter, rounded at
both ends; exocarp 1 mm. thick, coriaceous when dry, almost smooth,
reddish-brown; mesocarp 2-3 mm. thick, carnose (granulose when
dry), resinous. Endocarp woody, deeply 5-grooved, with 5 projecting
ribs divided halfway into two, except at one end, giving a 10-radiate
shaped section; the tissue between the ribs very compact, fibrous,
removable at maturity. Seeds most frequently 2-3, oblong, about
30 mm. long, 7 mm. thick.

Endopleura uchi is widely spread in the Amazon Valley, from Pard
to the Purus and Solimées regions. It is a large tree often cultivated
at Belém. Ducke writes about it thus: “This species is called
‘Uchy’ at Belém and is known at the lower Amazonas as ‘uchy pucu,’
‘uchy’ meaning elongated shape. It is found cultivated at the capital
and spontaneous in the forests of uninundated lands between Belém
and Braganga. The so-called ‘uchi-pucw’ of the lower Amazon Basin
belongs to this species, according to a flowering specimen from the
highlands near Cumina-mirim of the Trombetas region. I have also
observed this species in the forests of Volta of Xingu, of Gurupé, on
the ‘serras’ between Almeirin and Faro, and in the middle Tapajoz.
The fruit found by Huber in the forests of the upper Purus also seem
to belong to the species.”

BRAZIL: Amazonas: Manaos, Pensador, mata da terra firme; arvore grande,
flor verde cheirosa, “uchi,” “uchi-pucu,” 10-VIII-1943, Ducke 241 (IAN, MG,
S, NY, US, A). Pensador, arvore grande, flor verde, “uchya,” ‘“uchy-pucu,”
17-VII-1936, Ducke 305 (Y, NY). Pensador, silva non inundabilis, arbor magna
floribus viridibus odoratis, “uchi,” “uchi-puci,’’ 17-VII-1936, Ducke s.n. (US).
Manaos, silva non inundabili, arbor magna floribus viridibus odoratis, ‘uchi,’”’
15-VIII-1931, Ducke 23815 (US, 8, U). Rio Tonantins, terra firme baixa,
arvore de 4 m., flores amareladas, 27-X-1949, Frées 25565 (P, IAN). Region
Boa Vista, habitat terra firme, 8-IX-1932, Capucho 430 (IAN). Par: Pard,
horto bot. “uehi,” 14-VII-1896, Huber 239 (MG); photo F.M. 12602. Belém,
Jard. Bot., I-VI-1908, Huber 940 (M, U). Belém, hort. bot., “uchi,’’? X-1897,
Museu Goeldi 1260 (Manoel Guedes or Huber) (holotype, MG; isotype, US);
Manoel Guedes 1260 (US, MG). Bot. Gard. Mus. Goeldi, a fruit-yielding tree
long known in Pard as “uehi’’; the tree magnificent, becoming very tall with
dense rounded top, 1-VII-1908, Baker 58 (A, GH, NY, US, U). Belém, Jard.
Bot., 1-VI-1908, Baker 9401 (MG); Baker s.n. (M, 8, US). Belém, cultivado,
“uchi,’”? X-1897, Huber 1260 (isotype, US). Belém, culta et circa urbem spon-
tanea, arbor magna, floribus viridibus odoratis, “uehy,” VII-1923, Ducke 17779
(US, P, 8, U). Belém, arvore 108 ft. nfo possue cheiro, as fléres novas so en-
leiadas de cipés, madeira a ser estudada pela Universidade de Yale, ‘‘uchi,”
7-VII-1947, Black 47-1001 (Black project 135) (IAN, NY, U, US, VEN, Y).
Belém, Inst. Agr. Norte, cafezal antigo, “uchi,” folha consistente, 26-IV-1957,
Black 57-19306 (US). Rio Cumindmirim, Castanhal das Pedras, “uchy-puct,”
13-X-13, Ducke 14979 (MG). Rio Xingi, estrada da Volta, ‘uchi,” 20-XII-
1916, Ducke 16641 (MG).

84 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

4. Hylocarpa
Hylocarpa Cuatr., gen. nov.

Sepala 5 rotundata basi coalita imbricata. Petala 5 libera ovato-
oblonga praefloratione cochleari vel contorta. Stamina 30 filamentis
basi breviter coalitis. Antherae dorsifixae oscillantes connectivo
oblongo crasso obtusissimo pistilato vel malleiformi, 2 thecis uni-
locularibus subglobosis ad basim dispositis, tantum 5-15 fertiles.
Ovarium ovoideum 5-loculare loculis oppositisepalis uniovulatis,
ovulis anatropis raphe ventrali. Stylus crassus brevis. Stigma
5-lobatum. Drupa grandis subfussiformis exocarpio crasso sub-
farinaceo. Endocarpium lignosum durum haud resinosum eminente
5-costatum profunde 5-sulcatum fundo sulcis operculo lineari costi-
formi dehiscenti instructo. Semina oblonga. Arbores foliis alternis
coriaceis petiolatis persistentibus. Inflorescentiae cymoso-panicu-
latae axillares breves bracteatae bracteolataeque. Typus: Saco-
glottis heterocarpa Ducke.

Sepals 5, suborbicular, imbricate, united at base. Petals 5, free,
thick, oblong, the estivation cochlear or contorted. Stamens 30,
glabrous, the filaments biseriate, concrescent at base, thick and
papillose except toward the narrowed and pointed apex. Anthers
only 5 to 15 fertile, dorsifixed, the connective thick, very obtuse, club
or hammer shaped, the thecae 2, unilocular, subglobose, basal, de-
hiscing by the insertion line. Disk of 10, linear, thick, free scales.
Ovary ovoid, strigose, 5-locular, the cells uniovulate. The carpels
opposite the sepals. Ovules anatropous with ventral raphe, pendu-
lous. Style thick, very short. Stigma subcapitate, 5-lobate. Drupe
large, subfusiform, the exocarp thick, subfarinaceous. Endocarp
woody, hard, not resinous, prominently 5-costate, the elevated ribs
alternating with deep furrows, each having in the bottom a long
linear germinal operculum protruding as one rib. Seeds oblong,
usually only 1-2 (rarely 5) developed. Evergreen trees with simple
alternate, coriaceous, petiolate leaves. Inflorescences axillary,
dichotomous-paniculate. Bracts soon deciduous.

Hylocarpa differs from Sacoglottis in its large, compact (not resinose)
narrow-valvate-dehiscent endocarp. From this and all other humiri-
aceous genera it further differs in its thick, obtuse, pistillate or hammer
shaped anther-connectives bearing two basal unilocular glabrous
thecae, and in having 30 stamens, of which only 5-15 may be fertile.
Only one species of this genus is known, the Brazilian Hylocarpa
heterocarpa. (See figs. 3, 16, and 18-19.)

The name “Hylocarpa” is derived from the Greek “hyle” and
“hylo” (wood), and karpos”’ (fruit), an allusion to the hard, woody
large endocarp.

CONTR. NAT. HERB. VOL. 35 CUATRECASAS—PLATE 1

Fantanea compacta (Schniz.) Cuatr. (= Mumirtum compactum Schniz., — /fumirium
contractum Nloric.; holotype, Blanchet 3362, P)

CONTR. NAT. HERB. VOL. 35 CUATRECASAS—PLATE 2

Tantanea compacta subsp. microcarpa Cuatr. G(vpe, Pearce, IW)

CONTR, NAT. HERB. VOL. 35 CUATRECASAS—PLATE 3

f: fre eotinihern

Vee Ye

Fantanea guianensts Aubl. (type, Aublet, S)

CONTR. NAT. HERB. VOL. 35 CUATRECASAS—PLATE 4

Humirta balsamifera var. floribunda (Mart.) Cuatr. (= Mumirtum floribundum Mart.s: type

Martius NI)

’

CONTR. NAT. HERB. VOL. 35 CUATRECASAS—PLATE 5

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Humiria balsamifera var. guianen.
& Bergdolt: type,

CONTR. NAT. HERB. VOL. 35 CUATRECASAS—PLATE 6

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Hlumtria balsamtfera var. parvifolia (Juss.) Cuatr.

St. ffilaire, P)

(=Humirium paroifolium Juss.; tvpe,

CONTR. NAT. HERB. VOL. 35 CUATRECASAS—PLATE 7

Humirta halsamifera var. parvifolia (Juss.) Cuatr. (= flumirtum arenartum Guillemin; type,
Gruillemin 205, P)

CONTR. NAT. HERB. VOL. 35 CUATRECASAS—PLATE 8

Ci hy
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Ministerio de Agricultura y Cria
HERBARIO NACIONAL DE VENEZUELA
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Humtria balsamifera var. coriacea Cuatr, (type, Cardona 1823, US)

CONTR. NAT. HERB. VOL. 35 CUATRECASAS—PLATE 9

Harinca garroncfirn (dened / hid
: “fe tA:

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Humiria balsamifera var. guaiquinimana Cuatr. (type, Cardona 1112, US)

CUATRECASAS-PLATE 10

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CONTR. NAT. HERB. VOL. 35

HERBARIUM OF
THE NEW YORK BOTANICAL GARDEN

. a manrs orferr, Amazonas, Venezuela
Lo8otyo Se na. 42760 »
: Huwirea. wurdactha¢ Cumhs. Helena
al Shrub 1-3 m. Fla white, Locally abundant at mare
Sabana % bank of Cano Cumare

of Sal Cumare on
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elev. 125 m. .

THE NEW YOe@n @OTAN'GAL GanDEN

vGuUCrEOs FOR WOOO saerite earns. ree
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Ay SYSAC USE AMO VALE UNIVE Mm TS

fa

Humirta <x

Wurdack and Adderley 42760, US)

eurdackit Cuatr. (type,

CONTR. NAT. HERB. VOL. 35 CUATRECASAS—PLATE 11

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Palade de Amar (
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Humiriastrum villosum (Froes) Cuatr. (= Sacoglottts villosa Froes; tvpe, Proes 22644, TAN)

CONTR. NAT. HERB. VOL. 35

flumirtasirum

subcrenatum

(Benth.) Cuatr. (
Mearitn, IW)

/fumirium

CUATRECASAS—PLATE 12

cubcrenatum Benth.; tyvpe,

CONTR. NAT. HERB. VOL. 35 CUATRECASAS—PLATE 13

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UNITED STATES NATIONAL MUSEUM
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Humiriastrum mapiriense Cuatr. (type, Krukoff 11270, US)

CONTR. NAT. HERB. VOL.. 35 CUATRECASAS—PLATE 14

COSTA RICA

STATES NATIONAL MUsrUM — *

Humiriastrum diguense subsp. costaricense Cuatr. (dilen 5812, US)

CONTR. NAT. HERB. VOL. 35 CUATRECASAS—PLATE 15

- . Sthiatootin Tappesiemae
Lies Hotelypud

Ministerio de Agricultura y Cria
HERBARIO NACIONAL DE VENEZUELA

VOC ORE GAS
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Schistostemon auyantepuyense Cuatr. (Varescht °* Foldats 4673, type, VIEN)

CONTR. NAT. HERB. VOL. 35 CUATRECASAS~— PLATE 16

Schistetiveon lianlatanne
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; “ef Ly x ‘
tap [teeta Cees .
P-s9s8 Htety p “4s

Schistostemon reticulatum subsp. froestt Cuatr. (Frées 21370, NY)

CONTR. NAT.

HERB. VOL. 35

CUATRECASAS—PLATE 17

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Ver.

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f 3

Observactas bet gudertde lo @

if > hue ki :

Coliegit.

Determ, par Date

Schistostemon retusum (Ducke) Cuatr. (= Sacoglottts retusa Ducke; type, Ducke 20131, US)

CONTR. NAT. HERB. VOL.

35

(Benth.)

Cuatr. (
7I7VE, NIG)

}

(

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f

Sai Oelott ry

CUATRECASAS-—PLATE 18

[Sacogfathi Bact Nuher

Pototgpio 4

¥-58

Sik ta to (iwein anaed hell Ay how

(Bint) ‘Cleathh .

ducket Hluber: Type, Ducke

CONTR. NAT. HERB. VOL. 35 CUATRECASAS—PLATE 19

HERD. ROAD REORO. TRAD

Schistostemon densiflorum (Benth.) Cuatr. (= Sacoglottis haboertensis Bakh. f.; type, Bosch-
wesen 2068 UL)

CONTR. NAT. HERB. VOL. 35 CUATRECASAS~—PLATE 20

Sacoglottis amasonica Mart. (Martius. M)

CONTR. NAT. HERB. VOL. 35 CUATRECASAS—PLATE 21

HeL VANE Anca stot

ower ace TSGR ON

Brn. t+

ate ae een

Sacoglottis guianensts {ma. guianensis Benth. (Schomburgk 571, P)

—

Bae

~

CONTR. NAT. HERB. VOL. 35

Lattin rang paves anaty fhadirond,
rf ‘

. prin nto Ye phate cas pa, Tec

ae ae oe

Sacoglottts mattogrossensis {ma. mattogrossensis Malme (.Malme 2237, type, S)

CUATRECASAS—-PLATE 22

a a ‘grt hes

Brose cit, Matte Grosso

Herb. Brasil, Regsell.

Platte Hineris

Ue ecog lelti¢ 6 a og

$ 47 rg aes

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CONTR. NAT. HERB. VOL. 35 CUAT RECASAS—PLATE 23

qm hich
vies gh z
Oy heGpoe

—<_——___-——
SEI PARAENSIS (Mf Gieldi)

PARA (Bragily

Famitte

; ( ucke) é un: _ 7 ae -
MAO Coot... |
Moen 1 veces RY /958 Hefrtiee. .

Sacoglottis mattogrossensis Malme var. subintegra (Ducke) Cuatr. fma. puberula Cuatr.
(Ducke 16286, type, MG)

CUATRECASAS—-PLATE 24

CONTR. NAT. HERB. VOL. 35

a
{

URIOW NEREV IES

IX 113 J - : ,
HERB. ACAD. RHENO-TRAI t/
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SURINAME 8. exerve (F Reserve)
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Sacoglottis eydontoides Cuatr. (type, Boschwesen 6495, UV)

CUATRE CASAS—HUMI RIACEAE 85
|

1. Hylocarpa heterocarpa (Dueke) Cuatr., comb. nov.
Fiaurss 16,l-p, 18,c; 19,a—c-
Sacoglottis heterocarpa Ducke, Arch. Inst. Biol. Veget. Rio Janeiro 4:27,
pl. 2. 1938.
Type: Ducke 30137, Brazil, | mazonas, Rio Curicuriary.
Large tree with glabrous, more or less reddish terminal branchlets.
Leaves large, thick-coriaceous, glabrous. Petiole 3-6 cm. long,

Ficure 19.—a-c, Hylocarpa heterocarpa, *1 (Ducke 30137): a, fruit; b, endocarp showing a
detached valve; c, transection. d-e, Duckesia verrucosa (Ducke 16764): d, endocarp,
view from apex; ¢, transection (see two seeds).

513359—61 5

86 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

semiterete, striate, sulcate above. Blade obovate-elliptic or sub-
obovate, obtusely narrowed at base and decurrent along the petiole,
rounded or very obtuse at apex, slightly crenate or subentire at
margin, 11-17 em long, 7.5-10 cm. broad; above, dark green or brown-
ish, the midrib flat but conspicuous, the secondary nerves and veins
more or less prominulous; beneath brown or reddish when dry, the
midrib thick, striolate, the 14-16 pairs of secondary nerves thin and
prominent, ascending, curved at their insertion, arcuate and anasto-
mosing near the margin, the tertiary nerves and minor veins lax-
reticulate, prominulous.

Inflorescences cymose-paniculate, shorter than the petioles, dichoto-
mous; the short peduncle and the branchlets complanate, whitish-
puberulous. Bracts ovate-triangular, ciliate, deciduous, the upper-
most 0.5 mm. long. Pedicels 0.1-0.2 mm. long, glabrous, articulate
with a short (0-0.5 mm.), hirtcllous peduncle. Sepals 5, orbicular,
united at base, ciliate at margin, glabrous outside, 0.8-1 mm. long.
Petals 5, free, white, rather thick, lincar-oblong, obtuse, glabrous,
about 3.5 mm. long and 1.5—1.8 mm. wide, the estivation cochlear or
contorted. Stamens 30, more or less biseriate, concrescent at base, the
filaments 1.5-1.7 mm. long, rather thick, filiform at the end, broaden-
ing at the middle, densely covered with long papillas. Anthers
dorsifixed, versatile, only 15 or less fertile; the connective carnose,
oblong clavate or hammer-shaped, with 2 unilocular, globose, 0.7-0.8
mm. broad separated thecae at base; sterile anthers oblong, thick,
obtuse at apex, 0.7-0.9 mm. long. Disk perigynous formed by 10
linear, thick, about 3 mm. high scales. Ovary ovoid, slightly 10-
sulcate, strigose, 0.8 mm. long, 5-locular, the cells opposite the sepals,
uniovulate. Style short, rather thick. Stigma 5-lobate. The ovules
anatropous with ventral raphe. Drupe elliptic-subfusiform, about
9.5 em. long, 4.5 cm. broad, attenuate toward the base, abruptly
rounded-constricted at base, narrowed upward, apiculate, the
apiculum about 1 cm. long, more or less curved. Exocarp smooth,
2-3 mm. thick. Endocarp woody, broadly fusiform, truncate at base
(about 1 cm. diameter), middle part about 3.6 cm. in diameter, very
narrow at apex and acutely apiculate, deeply 5-sulcate, 5-costate, the
ribs very thick, prominent and robust, each of the 5 sulci with a germ-
inal operculum or window in form of a 4 em. long, very narrow, com-
pressed rib, only 2-3 (rarely 5) fertile.

According to Ducke, the mature fruit of H. heterocarpa, called
“cumate rana” by the natives, has an easily separable mesocarp, some-
what farinaceous, dry and tasteless. It is a tree of the catingas of the
uppermost part of the Rio Negro basin at the Brazilian corner between
Colombia and Venezuela.

CUATRECASAS—HUMIRIACEAE 87

BRAZIL: Amazonas: Rio Curicuriary, afluente del rio Negro, in silva “‘catinga”’
circa Cataractam Cajd, loco paludoso; arbor sat magna, fructus virides, ‘““ccumaté
da catinga,”’ Ducke 265 (Isotypes, NY, Y). Same locality and data, flores, 18-XI-
1936, fructus, 21-II-1936, Ducke 30187 (isotypes, US, U). Rio Igana, Tunuf, pé
da serra, terreno pedregoso (arenito) ; arvore de 10-15 m., flor branca, 23-X-1947,
J. Murga Pires 708 (IAN).

5. Humiria

Humiria St. Hil. Exp. Fam. 2:374. 1805.—Person, Syn. Pl. 2:70. 1807.—DC.
Prodr. 1:619. 1824.—Benth. & Hook. Gen. Pl. 1:247, in part, 1862.—
Urb. in Mart. Fl. Bras. 12(2):487, tab. 92. 1877.—Reiche in Engl. &
Prantl, Pflanzenfam, 3(4):37, fig. 32. 1890.—Winkl. in Engl. & Harms,
Pflanzenfam. 19a:106, fig. 59. 1931.—Lemée, Dict. Desc. Synon. Gen.
Phan. 3:670. 1931.—Bakhuizen van den Brink in Pulle, Fl. Surin. 8(1):
413. 1941.

Houmiri Aubl. Pl. Guian. 1:564, pl. 225. 1775.—Lam. Encycl. 3:139, 1789.

Wernisekia Scop. Intr. Hist. Nat. 273. 1777.

Myrodendrum, Schreb. Gen. Pl. (8 ed.), 1:358. 1789.

Houmiria Juss. Gen. Pl. 435. 1789.

Humirium Rich. ex Mart. Nov. Gen. Sp. 2:142. 1827.—Endl. Gen. Pl.
1040 (No. 5486). 1840.—Walp. Repert. Bot. Syst. 1:425. 1842.—Benth.
in Hook. London Journ. Bot. 2:373 (in part). 1843.—Benth. in Hook.
Journ. Bot. Kew Misc. 5:100 (in part). 1853.—A. Juss. in St. Hil. Fl.
Bras. Merid. 2:88. 1829.—Muell. in Walp. Ann. Bot. Syst. 4:383 (in part).
1857.—Baill. Adansonia 1:209. 1860; 2:262-264. 1861.

Myrodendron Spreng. Syst. Veg. 2:600. 1840.

Verniseckia Steud. Nom. Bot. (2 ed.), 2:752. 1841.

Houmiri Sect. Humirium Baill, Adansonia 10:370 (in part). 1873.—Hist.
P1. 5:54, figs. 88, 89 (in part). 1874

Wernischeckia Scop. ex Post & Kuntze, Lexicon 288. 1904.

Type species: Humiria balsamifera (Aubl.) St. Hil.

Sepals 5, suborbicular or ovate, imbricate, more or less coalescent
in a cupular calyx. Petals 5, free, thick-membranaceous, oblong or
linear, subacute or obtuse, the estivation cochlear or quincuncial.
Stamens 20, uniseriate, the filaments united in tube on about the
lower half, more or less complanate, densely papillose or muricate,
10 longer and alternating with the shorter 10. Anthers ovoid-
lanceolate, dorsifixed above the base, the connective thickly linguiform
or lanceolate, much longer than the 2 subglobose, hairy thecae inserted
sublaterally on the inside at the base, opening when pulled away.
Disk annular, surrounding the ovary, formed by 20 linear, thick, more
or less united scales. Ovary scarcely pilose at apex or completely
glabrous, 5-locular (rarely 4-locular), each cell with 2 anatropous,
pendulous, superposed ovules. Carpels opposite the petals. Style
columnar, erect, as long as the filaments or longer, more or less
hirsute. Stigmas globose, stellate. Drupe small (not exceeding 16
mm.), ovoid, ellipsoid or oblong; the epicarp thin; the mesocarp
fleshy, usually more or less sweet and aromatic, edible. Endocarp

88 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

woody, ellipsoid or ovoid, finely 10 (rarely 8) striate; the striae
equidistant marking 5 longitudinal, narrow, germinal valves, alter-
nating with 5 small holes at the apex; each cavity of the ovary usually
developing into 2 superposed cavities with 1 seed in each; sometimes
the 2 superposed seeds and cavities present and fertile; at other
times only 1 cavity (generally the lower one) fertile; often the inferior
fertile one alternating with the next superior one; usually 1-4 seeds
fertile, rarely more. Although no written material exists about the
dehiscence and germination of Humiria fruits, their structure makes
me assume that at the moment of germination, the narrow valve
is pushed away by the pressure of the embryo. Seeds commonly
4-1, well developed, 3-5 mm. long, subpyriform-ellipsoid, oblong,
acute at apex, the outer episperm scaly, the inner membranaceous;
the endosperm carnose. Evergreen trees or shrubs; wood hard.
Leaves alternate coriaceous or subcoriaceous, sessile or petiolate, often
more or less decurrent on the branchlets; margin slightly crenate
rarely completely entire, dotted with glands near the margin on the
lower side. Stipules small, falling soon or lacking. Inflorescences
paniculate and corymbiform with apparently alternate branching or
dichotomous, axillary or subterminal. Bracts persistent. (See figs.
3 and 20-23.)

There are only three recognized species of Humiria in this treatment;
one of them with numerous varieties and forms is widely spread
through the rain forests and savanna thickets of tropical South
America. They are important constituents of the above communities
and may become dominant in bush or thicket communities in savannas
and on the Guiana sandstone hills.

The name, ‘‘Tfumiria’” is taken from the Caribbean name “umiri,”
given in French Guiana for /7. balsamifera and is the form conserved
by the Code of International Nomenclature (see page 33).

Key to the Species of Humiria

1. Leaves small or linear.

2. Leaves linear (2.5-10X0.3-0.8 cm.), glabrous. Plant entirely glabrous.
Sepals orbicular, very obtuse . . . ... . 3. H. wurdackii

2. Leaves small (1.5-40.5-1.5 cm.), oblong « or elliptic-oblong, minutely
patulous-pilose, slightly velutinous at both sides, sessile. Young branch-
lets minutely pilose. Sepals subacute... ... . . 2. H. fruticosa

1. Leaves larger, glabrous or only the midrib pubescent beneath, rarely the
blade sparsely hirtellous. Sepals orbicular or very obtuse.

3. Leaves broad (7-144-9.5 em.); blade thick-coriaceous, rigid, oblong-
obovate or obovate-elliptic; petiole stout, broadly winged, folded-amplec-
tant. Drupe ellipsoid, 10-12 * 7-9 mm, Plant entirely glabrous.

4. H. crassifolia

3. Leaves medium-size (4-12X2-6 cm.), small (1.5-50.8-2.5 em.), rarely
larger (up to 18 cm. long); blade subcoriaceous or coriaceous, sessile and

CUATRECASAS—HUMIRIACEAE 89

amplectant or petiolate; petiole flat, narrower, not amplectant. Drupe
oblong-ellipsoid or oblong, 10-14*4-8 mm. Plant glabrous or some-
what pubescent. . ..... 2. . ss... . dL. H. balsamifera

1. Humiria balsamifera (Aubl.) St. Hil., Exp. Fam. 2:374. 1805.—Urb. in
Mart. Fl. Bras, 12(2) :440, tab. 92, fig. I. 1877. DC. Prodr.1:619. 1824.
Figure 20; PLares 4-9

Houmiri balsamifera Aubl. Pl. Guian. 1:564-566 pl. 225. 1775.

Myrodendrum balsamiferum Raecuschel, Nom. Bot. (3 ed.) 156. 1797.

Myrodendrum amplezicaule Willd. Sp. Pl. 2(2):1171. 1800.

Myrodendron amplezicaule Spreng. Syst. veg. 2:600. 1825.

Humirium floribundum Mart. Nov. Gen. Sp. Pl. 2:143-145, pl. 199. 1827.—
Benth. in Hook. Lond. Journ. Bot. 2:374. 1843.—Benth. in Hook. Journ.
Bot. Kew Misc. 5:100. 1853.

Humirium montanum A. Juss. in St. Hil. Fl. Bras. Merid. 2:90. 1829.

Humirium parvifolium A. Juss. in St. Hil., ibid.:89. 1829.

Humirium parviflorum A, Juss. in St. Hil., ibid.:63. 1829.

Humirium guianense Benth. in Hook. London Journ. Bot. 2:374. 1843.—
Hook. Journ. Bot. Kew Mise. 5:100. 1853.

Humirium balsamiferum Benth. in Hook. Journ. Bot. Kew Miscel. 5:102.
1853.

Humirium surinamense Miquel, Stirp. Surinam 86, pl. 24. 1850.

Humirium arenarium Guill. in Baill. Adansonia, 1:208. 1860.

Humirium multiflorum Mart. Spach Suites 17 in Pritz. Icon. Bot. Ind. 560.
1866.

Houmiri arenarium Baill. Hist. Pl. 5:52-53, figs. 88-89. 1874.

Humiria floribunda Mart. ex Urb. in Mart. Fl. Bras. 12(2):438, pl. 92.
1877.—Ducke, Arch. Jard. Bot. Rio Janeiro 3:176. 1922.

Myrodendron petiolatum Mart. ex Urb. in Mart. Fl. Bras. 12(2):438. 1877,
as synonym.

Humirium amplexicaule Mart. ex Urb., ibid.:440, as synonym. 1877.

Humirium ellipticum Klotsch ex Urb., ibid.:438, as synonym. 1877.

Humirium laurinum Klotsch ex Urb., ibid.:439, as synonym. 1877.

Humirium subsessile Spruce ex Urb., ibid.:439, as synonym. 1877.

Humiria Cassiquiari Siissenguth & Bergdolt, Repert. Sp. Nov. Fedde 39:16.
1935.

Humiria savannarum Gleason in Bull. Torrey Club. 58:378. 1931.

Humiria pilosa Steyermark, Fieldiana, Bot. 28:270, 1952.

Humiria floribunda var. guianensis (Benth.) Urb. in Mart. Fl. Bras. 12(2) 7439.
1877.

Humiria floribunda var. laurina Urb. in Mart., ibid.

Humiria floribunda var. montana (Juss.) Urb. in Mart., ibid.:438, 1877.

Humiria floribunda var. parvifolia (Juss.) Urb. in Mart., ibid.

Humiria floribunda var. subsessilis Urb. in Mart., ibid.:439. 1877.

Humiria floribunda var. spathulata Gleason, Bull. Torrey Club. 58:374.
1931.

Type: Aublet, French Guiana.

Small or large tree, in some varieties a low shrub. Branchlets
hirtellous, puberulous or glabrous, more or less compressed, the very
young in the typical form ancipital and more or less winged by
decurrence of the leaf base, in other forms subterete. Leaves sub-
coriaceous or coriaceous, thin or thick, flexible or more usually rigid,

90 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Ficure 20.—Humiria balsamifera (Ducke 23424): a, bud, 5; 6, petal, 5; c, flower, the
petals removed, showing the androecium, X 5; d, detail of staminal tube from the outside,
X10; ¢, anther, X10, front and lateral view; f, ovary, transection, 5; g, gynoecium,
from outside surrounded by the disk, and longitudinal section, 5.

sessile, subsessile or pseudopetiolate. Blade elliptic, obovate or
oblong with variable shapes; in the typical form broad and amplectant
at base, in others more or less attenuate at base, obtuse and sessile
or cuneate and short-petiolate or abruptly contracted into a winged
petiole; the apex rotundate, truncate or obtuse, broad or slightly
attenuate, sometimes mucronulate, often emarginate and mucronate
in the depression; very variable in size, up to 18 cm. long and 7 cm.
broad. Glabrous above or rarely with hirtellous-puberulous, con-
spicuous midrib, lateral nerves slightly apparent; with prominent,
glabrous or hirtellous midrib beneath, secondary nerves spreading,
thin, slightly prominent and conspicuous, 3-5 mm. distant, curved
and anastomosing near the margin, minor nerves parallel and others
laxly reticulate.

Inflorescences axillary and subterminal, cymose-paniculate, usually
corymbiform, 3-10 mm. long, very often aggregate forming very
floriferous compound panicle; peduncle 1.5-6 cm. long, ancipital and
winged in typical forms or simply angulate; branchlets rather stout,
rigid, fastigiate, often ancipital and more or less winged, thinner and
subdichotomous above, more or less densely hirtellous or glabrous.
Bracts triangular or ovate, 0.5-3 mm. long, amplectant. Pedicels
0.5-2 mm. long, thick, glabrous or rarely hirtellous, articulate with
usually puberulous, rigid, 1-2 mm. long peduncles, Calyx cupular,
1-2 mm. high, the sepals rather thick, suborbicular, imbricate, lower
part united, margin ciliolate, glabrous, rarely hirtellous outside.
Petals cochlear or quincuncial in estivation, rather thick, white or
greenish white, glabrous or rarely pubcrulous outside, lanceolate-
linear, subacute or subobtuse at apex, 4.5-7 mm. long, 1-1.6 mm.
broad. Stamens 20, filaments erect, rigid, 4-5 mm. long, alternating
2 sizes, united in lower half, free portion abundantly papillose. An-

CUATRECASAS—HUMIRIACEAE 91

thers ovate-lanceolate, 0.8-1 mm. long, thecae basal, subglobose,
pilose. Disk annular with about 20 linear, glabrous scales united at
base. Ovary ovoid, glabrous except for few hairs at apex, 5-locular,
cells biovulate with superimposed ovules. Style erect, rather thick,
more or less hirtellous, but glabrous near apex. Stigmas 5, thick,
stellate-capitate. Drupe oblong-ellipsoid, 10-14 mm. long, 5-8 mm.
thick, exocarp carnose, glabrous. Endocarp woody, ellipsoid-oblong,
obtuse or rounded at base, attenuate and acute at apex with 5 very
conspicuous foramina around tip; 10 thin and curved furrows longi-
tudinally; showing in transection 5 irregular cells. Seeds variable in
number, few.

IH. balsamifera is a broad specific complex that includes a great
number of different types with all possible transitional forms. Some
of the extreme forms already have received specific denominations,
such as H. floribunda, H. guianensis, H. montana, and H. parvifolia.
In the ‘Flora Brasiliensis,” Urban, understanding the great poly-
morphism of this group, merged the four species into one (ZH.
floribunda) and recognized in it six varieties, Through the study
of the abundant material, I came to the conclusion that all
these varieties are variations of the basic form of H. balsamifera;
furthermore, several new varieties, mostly endemics, have been dis-
covered in recent expeditions, especially in the Guianan highlands.

The most important variations observed in H. balsamifera are con-
cerned with the shape and size of the leaves and the indument. The
leaf blades vary from sessile and amplectant (broadly auriculate) to
narrowed and long petiolate, from obovate and subrotund to oblong,
and from abundantly pubescent on the nerves beneath to glabrous;
the leaf blades vary from thin-subcoriaceous to thick and rigid-
coriaceous, from small (2-4 cm.) to large (up to 18 cm. long), and
from an entire to finely crenulate margin. The young branchlets in
the typical HZ. balsamifera are appressed, ancipital, and winged, but
in other forms the young branchlets become angulate or subterete;
they vary from completely glabrous to hirtellous. The fruit also
varies from ellipsoid to ellipsoid-oblong. The auriculate, amplectant
long leaves are a juvenile form; examination of abundant collections
taken from the same tree at different times shows how this character
changes. Surinam material from the Forest Service, which is rich in
specimens taken from the same trees, shows the little value of the
form that I call attenuata because specimens with broad auriculate
amplectant leaves as well as with attenuate leaf blades are found on
one single tree. Between the extreme forms given as types of varieties,
all possible gradations are found.

The typical H. balsamifera has sessile and amplectant, obovate or
elliptic leaves and winged young branchlets; an adult form with atten-

92 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

uate leaves at base is frequent (fma. attenuata). When the leaves are
narrowed toward the base to being almost or shortly petiolate, we
have the variety floribunda with usually wingless branchlets. The
typical variety guianensis has obovate or broadly elliptic leaves,
abruptly contracted into a petiole. When the blades are oblong and
suddenly petiolate, we have the variety /aurina. Other varieties are
savannarum, with narrow sublanceolate leaves, attenuate at base;
parvifolia, with small, obovate, attenuate-cuneate leaves; coriacea,
with thick, rigid, obovate or elliptic-obovate leaves, attenuate at
base; guarquinimana, being like the former but with suborbicular and
subsessile leaves; stenocarpa, differing in its thinner drupes and
endocarps; awuana, differing from coriacea in its hirtellous petals;
imbaimadaiensis, with its sessile, amplectant, elliptic leaves; pilosa,
with more or less pubescent branchlets and leaves; and minarum,
with a distinct subacute form of elliptic leaves.

Among the many reported names which are locally or regionally
applied to Humiria balsamifera and varieties, the most common and
widely used are “umiri” or “umiry,” in Brazil and French Guiana,
and “blakkaberie” or “blaka-berie” in Surinam.

Key to the Varieties of Humiria balsamifera

1, Leaves sessile or subsessile, broad and amplectant, or broadly attenuate at

base.
2. Leaves elliptic, rounded at both ends, amplectant at base, rigid-coriaceous,
2.5-5 X 1.7-3.2 cm. Branchlets subterete. . . Ik. var. imbaimadaiensis

2. Leaves slightly attenuate toward the base, subrigid or flexible.

3. Leaves broad toward the apex, obovate or obovate-elliptic, rounded or
very obtuse at apex; 6-14 2.5-6 em. Young branchlets flattened
and more or less winged. . . .. .. . . . la. var. balsamifera

4. Leaves broad at base, more or less auriculate, amplectant.

la(Q1). fma. balsamifera

4. Leaves broadly attenuate, subcuneate at base. . la(2). fma. attenuata

3. Leaves narrowed toward the obtuse apex. Branchlets angulate or
subterete, rarely narrow-winged.

5. Leaves rhombo-elliptic-oblong or sublanceolate-elliptic, slightly at-
tenuate, obtuse or rarely auriculate-amplectant at base, 3.5-10X
15-4em. 2... ee . le. var. subsessilis

5. Leaves linear-oblong or subelliptic-oblong, sublanceolate, narrowed
and subacuminate, attenuate-cuneate at base; 5-10 1.5-3.5 em.

lf. var. savannarum

CUATRECASAS—HUMIRIACEAE 93

1. Leaves more or less attenuate at base with petiole or pseudopetiole.
6. Leaves abruptly contracted into a long, winged petiole, subcoriaceous or
subrigid-coriaceous.
7. Leaf blades broadly obovate or oblong-obovate or elliptic or suborbiculate,
3-12X2-6 em. The midrib generally pubescent beneath. Pseu-

dopetiole 0.5-2.5 cm. long... . .. . . « Id. var, guianensis
7. Leaf blades elliptic-oblong, 3.5-7 ‘1. 5- 3cm., glabrous. Pseudopetiole
0.5-1.5 em. long... ... ... .. Je. var. laurina

6. Leaves gradually attenuate at ‘base, more or less cuneate or suddenly
cuneate-contracted with short (1-3 mm.) petiole.
8. Leaves obovate or obovate-elliptic or rarely oblong-elliptic, long-atten-
uate, cuneate at base.

9, Leaves medium-sized or large, 5-12X2-6 em., subcoriaceous or sub-

rigid-coriaceous.. . . . . 2... . . « Lb. var, floribunda

9. Leaves smaller, 1.5-7 cm. long.

10. Leaves subcoriaceous 1.5-4%0.8-2.5 em., the veins slightly
prominulous. Young branchlets angulate, hirtellous-puberulous.

lg. var. parvifolia

10. Leaves thick-coriaceous, rigid or subrigid, the veins prominulous
especially beneath. Young branchlets glabrous or minutely
hirtellous-puberulous.

11. Petals hirtellous. Branchlets glabrous. Leaves 3.5-7X1.8-4
em. Petiole5-7mm. ......... . H. var. iluana
11. Petals glabrous.

12. Drupe oblong-ellipsoid, 10-145-8 mm. Leaves thick-coria-
ceous, 4-7 2.4-5.2 em. Petiole 2-6 mm. Young branch-
lets hirtellous or glabrous... . . . . . Lh. var. coriacea

12. Drupe oblong, 11-14X4-5 mm. Leaves thin-coriaceous
3-5.5X2-3.6 em. Petiole 5-7 mm. Young branchlets
glabrous. ... .. . . . lj. var, stenocarpa

8. Leaves generally subelliptic, short-attenuate at base, the veins prominu-
lous beneath.

13. Leaves suborbicular-elliptic or suborbicular, suddenly cuneate-
attenuate in a thick, broad, short (about 2 mm. long) petiole; the
margin usually entire, almost eglandular; 4-7X8-5.2 cm. Young
branchlets hirtellous.. ..... . . . li. var, guaiquinimana

13. Leaves oblong-elliptic, slightly attenuate at both ends, thin-coriaceous,
rather rigid, 3-5 1.5-2.5 cm.

14. Leaves glabrous, attenuate, subacute or subobtuse at apex, nar-
rowed at base with 1-2 mm. long petiole, the margin crenulate,
gland-dotted beneath. Young branchlets glabrous.

In. var. minarum

14. Leaves hirtellous on the midrib beneath, subrotund or subobtuse
at apex, attenuate at base with 2-3 mm. long petiole; the margin
generally entire, sometimes glandular. Branchlets hirtellous.

lm. var. pilosa

94 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

% 3 (
se 3 h os 1 Nie? ‘NG

YS

0000

Ficure 21.—a-t, Humiria balsamifera, X 1, fruit and endocarp: a, var. balsamifera (Black
49-8369); b, fma. attenuta (Frées 22738); c, fma. attenuata (A. C. Smith 2423); d, var.
floribunda (Black 47-1756); ¢, var. floribunda (Maguire et al. 30987); f, var. floribunda
(Maguire &$ Maguire 35040); g, var. attenuata (Maguire €9 Stahel 24957); h, var,
gutanensis (Maguire (9 Stahel 23654); i, var. guianensis (Maguire t? Politi 28828);
j, var. laurina (Maguire et al. 37632); k, var. subsessilis (Spruce 2454); 1, var. subsessilis
(Cowan tf Wurdack 31472); m, var. subsessilis (Schultes €3 Lopez 9510); n, var. subses-
silts (Williams 13868); 0, var. coriacea (Maguire 24707); , var. coriacea (Maguire
Politi 27627); q, var. coriacea (Maguire €$ Politi 27695); r, var, coriacea (Maguire et
al, 30018); s, var. stenocarpa (Maguire et al. 35882): t, var. stenocarpa (Maguire &9
Maguire 40105); u, H. crassifolia (Maguire &3% Fanshawe 23233); 0, H. balsamifera
var. savannarum (Wurdack & Monachino 41380); , H. balsamifera var. guaiquinimana
(Maguire 32763).

la(1). Humiria balsamifera var. balsamifera fma. balsamifera
FIGURES 21,a; 22,a-c

This variety is widespread throughout the savannas of British
Guiana, Surinam, French Guiana, and the Brazilian States of Ama-
zonas, Rio Branco and Pard, and is relatively common in wet and
inundatable places.

BRITISH GUIANA: Mazaruni Station, tree 60 ft., 12 in. diameter with spread-
ing crown on flat white sand areas by savanna; ripe fruit black, oval, edible,
glossy, pulp sweetish; ‘‘tauaranzu,” 10-I-1942, Fanshawe F715 (NY, U). De-
marara River ‘‘Jouranaro,”’ V-1887, Jenman 3912 (K, NY). Kartabo, small
tree, clearing in second growth, 17-VIII-1920, Bailey 115 (GH). Essequibo
River, Moraballi Creek near Bartica, near sea-level, ‘‘tauroniro” or “twaranru,”
tall tree, 121 ft. high, 344-4 ft. diameter in Wallaba forest, bark thinly fluted,
internodes winged, 5 white petals, numerous united stamens in column (compared
with Aublet’s specimens at B. by Sandwith), 8-X-1929, Sandwith 399 (NY, U,
US, P). Rupununi River, Quimatta, X-1889, Jenman 5672 (K). Without
locality, Rudge s.n. (BM).

CUATRECASAS—HUMIRIACEAE 95

a

>

;

DOO:

Ficure 22.—Humiria balsamifera, leaves: a, var. balsamifera (Herb. Richard, French
Guiana); b, var. balsamifera (French Guiana); ¢, var. balsamifera (French Guiana);
d, fma. attenuata (Schultes (3 Cabrera 15511); e, var. subsessilis (Spruce 2454); f, var.
laurina (Maguire et al. 36354); g, var. guianensis (Schomburgk 270); h, var. gutanensis
(Maguire et al. 41917); 1, var. coriacea (Maguire (9 Fanshawe 23295); j, var. cortacea
(Cardona 1869); k, var. coriacea (Maguire €9 Politi 27627); 1, var. coriacea (Maguire
24443); m, var. imbaimadaiensis (Maguire & Fanshawe 32158); n, var. laurina (Schom-
burgk 560); 0, var. parvifolia (St. Hilaire,‘ H. montanum’’); p, var. parvifolia (St. Hilaire);
q, different types of mucronate tip in H. balsamifera, especially in var. coriacea.

96 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

SURINAM: Boschreserve Brownsberg, Boomnummer 1273, 20-X-1924,
Boschwezen 6670 (U); 21-VII-25, Boschwezen 6907 (U). Savannah near Bown-
sweg, tree, ‘‘blakkaberie,’”’ 13-XI-1933, Lanjouw 1253 (US, U). Boschreserve
Zanderij I, Boschwezen 6234 (U). Zanderij I, Boomnummer 39, Boschwezen 1547,
2765, 2885, 5827 (U); 1935 (U, IAN, US); 2232 (U, US); 3010 (IAN, U). In
maurisie moerasbos op veen langs kreek in savanne bos bij Zanderij I; boompje,
“swietmerie” (Sur), 28-VIII-1954, Lindeman 6541 (U). Boschreserve Sectie O,
Boomnummer 92, Boschwezen 4669 (U, IAN); Boomnummer 845, Boschwezen
2471, 2918, 3934, 4684, 4810, 5412, 5858, 6010, 6068 (U). Zandsavanne W van
Patamacea, 23-I-1949, Lanjouw & Lindeman H8 (U). Blakawaka, 2-II-1921,
Junker 5525 (U). Jodensavanne high savanne side forest, sandy soil, alt. 70 ft.,
16-V-1954, Houtmonster 543A (Y); low forest, soil clay, 70 ft., 15-V-1954,
Houtmonster 541A (Y); 15-V-1954, Houtmonster 542A (Y). Jodensavanne-
Mapane kreek area (Suriname R), op droge zandsavanne bij Blakawatra kreek,
struik, “‘blaka-berie” (Sur.), 17—XII-1954, Lindeman 6880 (U); op droge zand-
savanne aan Blakawatra kreek; hoge struit, 17-XII-1954, ‘“blakaberie’’ (Sur.),
Lindeman 6861 (U); op droge zandsavanne achter Suhoza, struikje 2 m. hoog,
10-VII-1958, Lindeman 4202 (U). Patricksavanne, 9-V-1910, Boschwezen
“Indig.’”’ 212 (U). Via secta ab Moengo tapoe ad Grote Zwiebelzwamp; Klei-
savanne O van km. 65, struik 1.25 m. hoog; vrucht onrijp, groen, “blakaberie”’
(Sur.), “tawararo” (Arow), 29-IX-1948, Lanjouw & Lindeman 573 (U); km.
6.7, struik 2% m., bloem wit, Lanjouw & Lindeman 652 (U). Wajombo R.,
Donderkreek, op Zandsavanne; kleine boom circa 8 m. hoog, over de geheele stam
dicht begroeid met donkergroen loof met eigenaardigen boomvorm, ‘“tawaanroe”’
(Arow), “‘meerie” (Kar), ‘‘blakberrie” (NE), “baslerd,” “bolletrie,” 23-IX-1916,
Stahel & Gonggryp 3570 (U).

FRENCH GUIANA: St. Jean, Concession Bonnet, 24~V-1914, Benoist 1239
(P). Acarouany, 1858, Sagot s.n. (P). Route de l’Acarouany au km. 5, coté
gauche de la route, fleurs blanchdtres en grappes formant corymbe axillaire en
pointe des rameaux, “‘bonga-bita’”’ (Paramaka), 5-XI-1956, BAFOG 7587 (U).
Km 4 coté droit et 4 20 m. de la route, fruit vert, leger-ovales ressemblant a des
petites olives; inflorescences 4 la pointe des rameaux, “bonga-bita,” 29-IX-1954,
BAFOG 228M (U). Cayenne, arbe, “bois rouge,” 14-XII-1955, BAFOG “P.
Bent: 1083 (U). Cayenne, Martins.n. (BM, P). Cayenne, 1857, Sagot s.n. (P).
“Guyane frangaise,” without locality, Melinon 1863 or 1864 (A, GH, MO, NY,
P, US), Melinon 48 (P); Melinon 377 (P, K). Herb. Exp. Col. (P), ex Herb.
Richard (P), ex Herb. Cosson (P); Le Pricur 1838, 1840 (P, K).

BRAZIL: Amazonas: Manaos, Cachoeirinha circa ripas paludosas Igarapé do
Crespo, arbuscula floribus albis, Ducke 23424 (US). Rio Vaupes, Taraqud,
caatinga proxima a serra, arvore 15-20 m., flor branca, frutos comestiveis, “umiri,”
10-XI-1947, Pires 989 (IAN, NY, P, US). Rio Curicuriary, afluente do Rio
Negro, silva ad ripas altas cataractae Caji; arbor mediocris floribus albis, cortice
odorato, “umiry,” 29-II-1936, Ducke 30128 (MO, F, K, US, U, P). AmMap&:
Oiapoque, campo de aviacéo campina arvore pequena, 3-X-1949, Black 49-8313
(IAN). Oiapoque, cafio Inocencia, arvore, 4-X-1949, Black 49-8369 (IAN).
Oyapock Airfield, infrequent, secondary forest, tree 5 m. tall, 7—XII-1954,
Cowan 38700 (NY). Estrada Macap4-Clevelandia, km. 130, arvore flor branca,
27-VIII-1955, Black 55-18577 (IAN). Rio Branco: Caminho de Samauma
a campo de aviagdo de Mucajaf, kms. 14-15 da estrada Boa Vista-Caracaf, arvore
pequena, flor branca, 25-VITI-1951, Black & Magalhdes 51-12954 (IAN, P). Rio
Branco, entre as Fazendas Bom Intento e Capela, flor branca, 31-VITI-1951,
Black 51-13231 (IAN). Pari: Belém, Lago da Agua Preta, 29-X-1914, Ducke
15514, 15515 (MG). Pard, Moss 13 (US); Moss 57 (US, NY). Santa Isabel

CUATRECASAS—HUMIRIACEAE 97

(Estagdo Ferr. Braganga), VI-1908, Museu Goeldi 9419 (MG). Igarapé Game-
leirinha, campos gerais, regiéo do Araguaia, arvore 7 m., flor branca, 17-VI-1953,
Frées 29854 (IAN). Rio Maicurd, estrada Cad-ussi a Serra de Luna, campina
arenosa, muncipio de Monte Alegre, arvore de 5 m., flor esbranquigada, 9-III-
1953, Frées & Filho 29486 (IAN, US). Tefé, mata virgem, arvore 30 m., flor
branca, madeira bonita, vermelha carregada (amago), flor cheirosa, “pau mirim,”
25-VIII-1947, Black 47-1276 (IAN, NY, U, US, VEN). Marajé, Jutuba, matta
de marjem, arvore alta, 21-VII-1902, Huber 2785 (BM, MG). Without data,
Schultes 9431a, 9435 (IAN).

la(2). Humiria balsamifera var. balsamifera fma. attenuata Cuatr., fma. nov
Fiaures 21,b-c,g; 22,d; 23,p

A forma balsamifera typica differt folia basi late attenuata (nec
auriculata) sessilia.

Type in the U.S. National Herbarium, No. 2171627, collected on
quarzite rocks of falls at Raudal Yayacopi (La Playa), 800 ft. altitude,
Rfo Apaporis, Amazonas-Vaupés, Colombia, February 18, 1952, by
Richard Evans Schultes and Isidoro Cabrera, No. 15511.

Ficure 23.—Humiria fruit: a-d, H. balsamifera var. floribunda: a, transection (Maguire
&F Maguire 35040); b, longitudinal section; c, transection (Black 47-1756); d, longitudinal
section. e-g, H. balsamifera var. coriacea: ¢, transection (Maguire et al. 30018); f,
longitudinal section; g, transection (Maguire & Politi 27695); j-k, H. balsamifera var.
subsessilis: j, transection (Williams 13868); k, transection (Cowan &¢ Wurdack 31472).
I-m, H. balsamifera var. guianensis: 1, transection (Hitchcock 16938); m, transection
(Maguire &9 Politi 28828). n, H. balsamifera var. stenocarpa (Maguire & Maguire
40105), transection. o, g, r, H. crassifolia (Maguire 9 Fanshawe 23233): 0, transec-
tion; g, shape of endocarp with oblong and pitched valves corresponding to super-
posed cavities; r, longitudinal section of the same endocarp to show two superposed
cavities and ovules at one side and one ovule in its cavity with sterile upper cavity
at the other side. », H. balsamifera fma. attenuata, longitudinal section of endocarp
showing two superposed cavities and ovules from one carpel and a single ovule
developed in another carpel (Frées 22738).

98 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

This is just a form of the typical H. balsamifera with attenuate
leaves at base. It is found in somewhat dryer parts than the typical
form, in sandy or sandstone places, in savanna remnant forests or
thickets, etc.; the morphologic difference probably depends very much
on local ecological conditions. It is spread from the four Guianas
through the Amazon basin to the Huallaga River in Peru and to
Belém (Pardé) and Goids in Brazil.

COLOMBIA: Vaupfs: Rio Apaporis, raudal Yayacopf (La Playa), quarzite
base, alt. 800 ft., small tree, 18-II-1952, Schultes & Cabrera 15511 (US, holotype);
on sand, small black edible fruit, “wa-too-moo-ko” (Yukuna), 18-VIII-1952,
Schultes & Cabrera 16893 (US).

VENEZUELA: Botfvar: Region de los rfos Ieaburé, Hacha, 450-850 m.,
sabana, drbol aislado en la sabana, corteza rayada, hoja coridcea, 24-XII-1955,
Bernardi 2601 (NY).

BRITISH GUIANA: Mazaruni Station, mixed forest, tree 90 ft. high, 17 in.
diam., bark dark brown, fissured; wood hard, heartwood red, “tawasansu,”
12-V-1933, Tutin 83 (U, BM, US). Basin of Rupununi River, Isherton, “umir”
(Wapisiana), A. C. Smith 2423 (A, F, MO, NY, Y, US, 8). Atkinson Field, at
the east border, tow tree in open dry bush, flowers white, odoriferous, 2-V-1954,
Irwin 246 (US). Lucky Spot, sandbank, tree, flowers white, X-1924, Persaud
191 (F, K, NY). Massaroonie v. Essequebo, “bastard bully-tree,” “towraneroo,”’
Appun 37 (K).

SURINAM: Prope Patrick-Savanna, 5-V-1910, Boschwezen “Indig.” 177 (U).
Boschreserve, Sectie O, 1-XI-1920 Boschwezen 4770 (U); Boschwezen 5850 (U,
IAN). Zanderij 1-X-1942, ‘“‘tawaldenra,” “tawalingro” (Arow), “meri’’ (Ka),
“basrabolletrie,” Stahel 90 (A, IAN, NY, U, Y). Vicinity Arawak Village of
Mata, tree, 18-X-1944, Maguire & Stahel 24957 (NY, U). Boven Sipaliwini,
kamp IV, groote savanne, 2 NB, 56 WL, 23-X-1935, Rombouto 228 (U).

FRENCH GUIANA: “Guyane,” Poiteau s.n. (K).

BRAZIL: Rio Branco: Uferwald bei Boa Vista, Strauch 3-8 m., X-1908,
Ule 7625 (US, K, UC). Igarape Carand, arvore em campo seco, flor branca,
20-VITI-1951, Black 51-12776 (IAN). Amazonas: Rio Demeni, affluente
do Rio Negro, Posto Indigena Genipapo, caatinga, arvore 15 m., 12—X-1952,
Frées 28895 (IAN, US). Rio Negro, Preto Campina, tree 25 ft., whitish flowers,
rather grassy country, sandy soil, 5-XI-1957, Frées 22738 (IAN, NY, US,
VEN). Manaos, shrubby tree, 20 ft., Tate 50 (NY). Part: “Prov. Para,”
Martius s.n. (M). Belém, near Catu, 8 km. northeast of Inst. Agr. do Norte,
tree 20 m., fruits olive-colored, edible, a local drugstore sells a syrup made from
bark for relief from constipation, ‘“mirim,” 15-IX-1942, Barbosa da Silva 73
(IAN, US). Vigia, Campina do Palha, arvore pequena, fruto verde, 29-IX-
1948, Black 48-3249 (IAN, P); solo arenoso, arvore 12 m., flor branea muito
cheirosa, 10-VI-1952, Frées 28454 (IAN, US); bordos e ilhas da matta, arvore
pequeno, flores alvo-cremes, ‘‘umiri,” 29-IV-1953, Lima 53-1273 (IAN). Obidos,
beira do lago, flor branca, 12-VII-1905, Ducke 7213 (BM, MG). Rio Guamé4,
acima de Ourén, porto do Posto Indigena Tombé, arvore grande, flor branca,
matta de varzea, VI-53, Pires & Silva 4629 (NY), 4624 (IAN). Gotks: Carolina,
arvore de terras frescas, Branca, “meirin” (pan-meirim), 11-VIII-1955, Macedo
4034 (IAN, US).

PERU: Lorero: Miahuyaco, near Iquitos 100 m., forest, tree 18 m. high,
flowers white, V-1930, Klug 1315 (F, US). Balsapuerto, 200 m., tree 4 m.,
flowers white, I-1933, Klug 2846 (A, BM, F, GH, MO, NY,S, US). Balsapuerto,

CUATRECASAS—HUMIRIACEAE 99

lower Rio Huallaga basin, 150-350 m., dense forest, 28-30—VIII-1929, Killip &
Smith 28681 (F, NY, US).

lb. Humiria balsamifera var. floribunda (Martius) Cuatr., comb. nov.
Fiaures 21,d-f; 23,a-d; 24,c; PLATE 4

Humirium floribundum Mart. Nov. Gen. Sp. Pl. 2:148, pl. 199. 1827.—
Benth. in Hook. Journ. Bot. Kew Miscel. 5:100. 1853.

Humirium multiflorum Pritz., Ic. Ind. 560. 1866.

Humiria floribunda Mart. Urb. in Mart. Fl. Bras. 12(2):437, 1877.—Ducke,
Arch. Jard. Bot. Rio Janeiro 3:176. 1922.—Arch. Inst. Biol. Veget.
Rio Janeiro 4:25. 1937.

Humirium ellipticum Klotsch ex Urb., ibid., as synonym.

Type: Martius, Brazil, Pard, Xingt (M).

Leaves attenuate toward cuneate base, petiolate or subsessile,
elliptic, obovate-elliptic or somewhat oblong; slightly attenuate and
obtuse, often emarginate, sometimes rounded or truncate at apex;
glabrous but rarely with pubescent midrib beneath; margin usually
minutely crenate, glandular-punctuate beneath or eglandular; 4-12
cm. long, 2-6 cm. broad. The young branchlets generally glabrous
or sometimes puberulous, angulate or subterete, rarely narrowly
winged.

The binomial Humirium multiflorum is the result of an erroneous
quotation by Pritzel, which has never been validly published. Spach
(1846) described two species with a French title followed by the
corresponding Latin binomial; one species is ‘““Humirium de Guiane”
(=Humirium balsamiferum Aubl.), and the other is ‘“Humirium
multiflore’ (=Humirium floribundum Mart.). The adjective ‘“multi-
flore’”’ is here merely the French translation of H. floribundum. Pritzel
erroneously latinized this adjective to a specific name.

The variety floribunda is widely spread throughout the Amazon
Basin, from Venezuela, Guianas, and Surinam to the southern States
of Brazil in Rio de Janeiro. It can be found in the rain forests on
uninundatable ground, as well as in open places in savannas and
campos. According to Ducke, it is common in the State of Paré
on white sandy soils containing some humus and is the typical tree
of the “campinas”; it varies from a small shrub scrambling on the
stony campos of Ariramba to a large tree of 30 m. height in the
forests near Belém. Ducke mentions intermediary forms between
var. floribunda and balsamifera between Belém and Braganga. <Ac-
cording to Ducke, floribunda produces the “umiry balsam’’ which
smells like the Peruvian balsam. ‘This ‘umiry balsam’ is only
produced by the more or less sick old trunks of the forest. I have
never seen this balsam in trees located in open places; it seems to be
due to an illness of the tree probably caused by bacteria. ‘The drupes
of this variety are edible, sweet, and agreeable, especially those from

100 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

y
é

De

ANN
a

Ficure 24.—a-f, Humiria balsamifera: a, var. guaiquinimana, X ¥ (Cardona 1112); b, var;
coriacea, bud, X5 (Maguire {3 Fanshawe 23295); c, var. floribunda, *Y (Riedel s.n.).
d, var. laurina, bud, X5 (Krukoff 1483); e, var. parvifolia, bud, X5 (type of “H.
montanum,” St. Hilaire); f, var. pilosa, leaf, X14 (Steyermark 60289); g, Humiria
fruticosa, X% (Maguire et al. 36580); h, Humiria fruticosa, bud, X5 (Maguire et al.
36580).

the trees of the campos and campinas, whereas I have found those
of the rain forest to be insipid.”

VENEZUELA: Amazonas: Cerro Sipapo, forest near Base Camp, alt. 125 m.,
small tree with greenish white flowers, 28-XII-1948, Maguire & Politi 27974
(NY, US). Serrania Yutaje, Rio Manapiare, Base Camp, alt. 200 m., occasional
at savanna edge 1 km. east of Base, shrub or small tree, 4-10 m. tall, fruit green,
29-I-1953, Maguire & Maguire 35040 (NY, US). Cerro Moriche, occasionally
in sabanita at northwest base of mountain, alt. 150 m., tree 3 m., fruit green,
17-I-1951, Maguire, Cowan, & Wurdack 30987 (NY, US).

CUATRECASAS—HUMIRIACEAE 101

BRITISH GUIANA: Along the Berbice-Rupununi Cattle Trail, Berbice
or Demerara County, 29-V-1919, Abraham 152 (NY).

SURINAM: Jodensavanne-Mapane Kreek area (Suriname R.); op droge
zandsavanne acter Suhoza; struik, bloem wit, 10-VII-1953, Lindeman 4201
(U); op droge zandsavanne aaen Blakawatra, struik, “blakaberie’’ (Sur.), 17-
XII-1954, Lindeman 6862 (U). Via secta ab Moengo tapoe ad Grote Zwiebel-
zwamp; in ritsbos bij km. 14.9; boom 17 m. 18 cm. dik, “blakaberie’’ (Sur.),
25-X-1948, Lanjouw & Lindeman 968 (U).

FRENCH GUIANA: Cayenne, west, frequent small tree in islands of trees
in white sand savannas, 26—X-1954, Cowan & Maguire 38034 (NY). ‘Guiana
fr.,’’ Leblond 441, 402 (P). Donant Herb. 1686, without locality (P).

BRAZIL: Rio Branco: Serra da Malacacheta, “umiry,” Kuhlmann 3509
(US). Uferwald bei Boa Vista, Bl. weisslich, Strauch 3-8 m., X-1908, Ule 7625
(MG). Amazonas: Manaos, margen do Igarapé do Buido; terra firme, capoeira,
arvore de 6 m. alt., flor branea, 31-VIII-1955, INPA 1773 (MG, US). Manaos,
prope Cachoeira alta do Tarumé, silva non inundabili loco humido; arbor mediocris
floribus albis, 25-IV-1932, Ducke 87 (A, Y). Manaos, Colonia Joao Alfredo,
arvore de flor ligeiramente rosea, 17-VIII-1954, JNPA 86 (MG, US). Manaos,
Igarapé do Parque 10, terra firme; arvore de 10 m., flor braneca, 25-V—1955,
INPA 1056 (MG, US). Manaos, Rio Negro, IX-1948, Schultes & Lopez 10339.
Manaos, arvore, “humiry,” 15-II-1945, Frées 20480 (IAN, US). Municipality
of Manaos, Rio Negro basin, along road to Aleixo, campinarana, VIII-IX-1936,
“mirim,” tree 40 ft., trunk 4 in. diameter, flowers white, Krukoff 7928 (A, NY,
BM, 8, U); ‘‘mirim,” tree 45 ft., 5 in. diameter, flowers white, Krukoff 7926 (A,
BM, F, MO, NY, 8, U). Campinarana, Rejias do Alto Ariramba, 20-XII-1906,
Ducke 8029 (BM, MG). Manaos, circa Cachoeira da Mundi, silva non inun-
dabili; arbor parva, flos albis, “umiry,” 2-IX-1942, Ducke 541 (IAN, MG, NY,
8, US). Rio Negro, Aiary-Pird, Rio Aiary, caatinga arenosa, arvore, 5-XI-1945,
Frées 21346 (IAN, NY, US). Rio Negro, prope Barra, 1851, Spruce 1499 (NY,
GH, BM, K, M, US, 8, P). Maranu&o: Cururupi, Lisboa 4099 (US). Graja-
hd, 13-VIII-1909, Lisboa 2327 (US). Ilha de Sao Luiz; reservatorio Socaven;
terra firme, baixa, floresta alta; arvore de 5 m., flor branca, 15-I-1951, Frées
26812 (IAN). Ilha de Saéo Luiz, SAo Sim&o, campinarana, tree 60 ft. 9 in.,
“umiry,’’ 1940, Frées 11813 (NY). Alcdntara; taboleiros descampados de beira
mar, 28-IX-1903, Ducke 440 (BM, MG). Bords du Rio Tocantins, VITI-IX-
1844, Weddell 2361 (P). Amari: Rio Araguari, campos gerais, arvore 7 m.,
flor branea, 22-VII-1951, Frées & Black 27572 (IAN). Par: Santarém, January,
Spruce 928 (K, P). In vicinibus Santarém, VI-1850, Spruce s.n. (P, BM, GH).
Perto da Casa Santa Izabel, municipio de Santarém, campo coberto, arvore
pequena, ‘“umiri,” 28-X-1950, Black & Ledoux 50-10371 (IAN). Caripi, VIII-
1849, Spruce 928, 181 ? (NY), 181 (M). Caripi, on the beach, low round-topped
tree with roughest sweet-scented bark, flowers white, ‘umiri,” WIII-1849,
Spruce 164 (K, P). Rio Arapius, 6-VI-1952, Pires & Silva 4192 (IAN, NY).
In ripis amnis Parae et St. Antonii Herb. Richard (P). Gurupdé, campina do
Igarapé Jucup{; arvore mediana, flor branca, 18—-VIII-1954, Pires & Silva 4702
(IAN, US). Conceigéo do Araqudia; terreno arenoso, imido a beira dos campos
de transicio para a mata dos carrascos; arvore de 10 m., 3-VII-1953, Frées
30093 (IAN, US). Rio Jamaracart, perto do barracéo; arbusto reptante; flor
branca perfumada, 26-V-1957, Black, Egler, et al. 57-19590 (IAN, US). Obidos,
Rio Jaramacurti, sandy and over-flowed field, plant of 50 em., corolla white,
27-V-1957, Egler 261 (MG, US). Belterra, beira do R. Tapajés igapé6; arbustivo
espalhado, flor branca, fruto verde, “umiri,”’ 24-X-1947, Black 47-1756 (IAN,
NY, US, U, VEN). “Iter Brasil. Sept. habitat in ripa ad flumen Xingti ostia

513359—61——_6

102 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

locis arenosis, Provinciae Paraénsis, Mart. Observ. 2692, Myrodendron petiolatum
Mart. tab. 195,” Martiuss.n. (M, holotype of Humirium floribundum). ‘Brasilia
Prov. Pard, Umiri Paraénsibus Martius Observ. 2692,” Martius s.n. (M, isotype
of Humirium foribundum). Crara: Fortaleza, Huber 96 (BM, US). Bemfica,
10-IX-1907, Huber 96 (MG). “Ceard,’’ Allemio & Cysneiros 255 (US). Banta:
Salzman s.n., no locality (P).. Rio pg Janrtro: Restinga de Cabo Frio; arbrisseau,
fleur blanc jaundtre, 20-VI-1877, Glaziou 10437 (P). Inter Rio de Janeiro et
Campos, Sellow 171 (K). Environs de Rio de Janeiro, Weddell 526 (P). Without
locality Sellow sn. (M, NY); Riedel s.n. (P). Versus Colares, Poeppig s.n.
(P). “Amerique Meridionale-Poeppig, 1868 No. 34,” Poeppig 3011 (P). Prov.
of Alagoas, Maceio, IV-1838, Gardner 1263 (BM). Without locality, Poeppig
81 (BM).
le. Humiria balsamifera var. subsessilis (Urb.) Cuatr., comb. nov.
Fiaures 21,k-n; 22,e; 23,j-k
Humiria floribunda var. subsessilis Urb. in Mart. Fl. Bras. 12(2):439. 1877.
Humirium subsessile Spruce ex Urb. in Mart., ibid., as synonym.

Type: Spruce 2454, Brazil, Vaupés.

Leaves subcoriaceous, rigidulous, sublanceolate-elliptic or rhom-
boid-elliptic, slightly attenuate, obtuse, subrotundate, rarely am-
plectant at base, attenuate and obtusely acuminate at apex, some-
times minutely emarginate, the margin crenulate and glandular-
punctate beneath; the midrib densely hirtellous, puberulous or
glabrous; 3.5-10 cm. long, 1.5-4 em. broad. Branchlets wingless,
hirtellous, puberulous, or also glabrous. Pedicels and sepals hirtel-
lous or glabrous. Petals glabrous or puberulous. Drupe ellipsoid-
ovoid, 11-12 X 7.5-8.5 mm.; the endocarp verruculose, 9-11 5-7
mm., acute at apex.

The type has the branchlets, nerves of the leaves beneath, pedicels,
and sepals hirtellous-pubescent; but few of the other collections
(Spruce 2454, Pires 1029) have these features; most of the specimens
lack the indument on pedicels and sepals, whereas in others it becomes
scarce on branchlets and leaves and passes gradually to the glabrous
forms. But the elongate-elliptic, sublanceolate leaves, which are
narrowed toward the obtusely subacuminate apex, are a typical
feature that characterizes a consistent variety spread from the
Vaupés River to the Guiana plateaus. It does not have obovate,
spatulate, or broadly obtuse leaves (not attenuate at apex in adult
stage) as do varieties balsamifera and floribunda.

The range of subsessilis extends from the Brazilian and Colombian
Vaupés to the Negro and Orinoco Rivers and the southern savannas
of Venezuela. Undoubtedly the Humiria population of the west coast
of Colombia refers to the same variety.

COLOMBIA: Vaupés: Rfo Piraparand, tributary of Apaporis, Cafio Timifia,
large bush, flowers white, ‘‘ta-ta-wee-tee-go” (Barasana), 6-IX-1952, Schultes &

Cabrera 17231 (US). Rio Cubiyi, cerro Cafiend4, sabanas 15 millas arriba de la
bocana, alt. 800-900 ft., small tree, bushy, 20 ft., flowers white, 10-XI-1952,

CUATRECASAS—HUMIRIACEAE 103

Schulies & Cabrera 18319, 18371 (US). Rio Cubiyd, alt. 350 m., 9-10-X-1952,
Humbert & Schultes 27364 (US). Rfo Negro, vicinity of Piedra del Cocui, bush,
caatinga, 27-XII-1947, Schultes & Lépez 9510 (IAN, NY). Rio Negro, San
Felipe, alt. 200 m., 18-25-XI-1952, Humbert 27422 (P). Regién de San Felipe
a orillas del Rio Guainfa, 250 m. alt., drbol 4 m., frutos verdes, 21—XI-1948,
Araque & Barkley 18 Va021 (US). Valle, Costa del Pacffico, Bahia de Buenaven-
tura, Quebrada de Aguadulce 0-10 m. alt.; drbol, hoja coridcea, flexible, verde
oscura semibrillante haz, claro envés; perianto verde pdlido, blanquecino,
“oloroso,” 11-XI-1945, Cuatrecasas 19727 (COL, G, U, F).

VENEZUELA: Amazonas: Region of San Fernando de Atabapo, Rfo Orinoco,
frequent on laja on left bank of Cafio Cupueni opposite mouth of Rfo Atabapo,
rounded dark green tree, 15 m. tall, flowers white, 12-XI-1953, Maguire, Wurdack,
& Bunting 36210 (US, NY). Between Sabana Grande and southeast base of
Cerro Duida, alt. 200 m., 23-VITI-1944, Steyermark 57880 (US, F). Rio Orinoco,
Rio Cunucunuma, edge of Culebra savanna, north base of Cerro Duida, alt.
200 m., much branched tree, 10 m., flowers greenish white, 27-XI-1950, Maguire,
Cowan, & Wurdack 29769 (US, NY); 18-XI-1950, Maguire, Cowan, & Wurdack,
29416A (US, NY). Rfo Ventuari, savanna on Cafio Arisa at west base of moun-
tain, alt. 200 m., locally frequent, tree 6 m., fruit green, 16-II-1951, Cowan &
Wurdack 31502 (US, NY); tree to 10 m., fruit purple-black, edible, 15-II-1951,
Cowan & Wurdack 31472 (US, NY). Esmeralda, alt. 325 ft., tree savannas,
flowers white, Tate 330 part (first sheet) (NY); Tate 331 (US, NY). Yavita,
280 m. alt. en terreno rocoso y abierto; drbol 5 m. o mds con copa casi plana,
tronco redondo, corto, 10 cm. did4m., madera clara; fruto monospermo, azulejo o
casi negro de pulpa algo dulce y comible, “‘nifia,’’ 22-I-1942, Llewelyn Williams
13868 (US). Botivar: Soropdn-tepuf, wooded ridge, between quebrada and
waterfall, alt. 1,656-1,980 m., small tree 25 ft.; leaves subcoriaceous, dark green
above and shining, dull green below; ‘‘coporik-warei-yek,” Steyermark 60192 (NY).

BRAZIL: Amazonas: Vaupés, Panure, caatinga; arvore pequena, flor branca,
15-XI-1947, Pires 1029 (IAN, VEN). Panure, IX-1952 to I-1953, Spruce
2457 (GH, BM, NY, MG, P). Prope Panure, IX and XII-1952, Spruce 2454
(isotype of H. floribunda var. subsessilis, BM, F, GH, NY, P, 8). Ipanoro, Rio
Vaupés, between Ipanoré and confluence with Rfo Negro, small tree, flowers
green-white, 14-15-XI-1947, Schultes & Pires 9103A (US, IAN). Rio Xiborem,
weiss, am Ufer, 15 m., 30-VIII-1928, Luetzelburg 24014 (M).

1d. Humiria balsamifera var. guianensis (Benth) Cuatr., comb.nov.
FicurEs 21,h-1; 22,g-h; 23,l-m; PuaTE 5

Humirium guianense Benth. in Hook.London Journ.Bot.2:374. 1843.—
Hook. Journ. Bot. Kew Miscel.5:100. 1853.

Humirium surinamense Miquel, Stirp.Surinam 86, pl. 24. 1850.

Humiria floribunda var. guianensis (Benth.) Urb. in Mart. Fl. Bras.12(2):
439. 1877.

Myrodendron petiolatum Mart. ex Urb., ibid., as synonym.

Humiria Cassiquiari Sussenguth & Bergdolt, Repert. Sp. Nov. Fedde 39:16.
1935.

Type: Schomburgk 270, British Guiana.

Leaves coriaceous or subcoriaceous, broadly obovate or suborbicular,
abruptly contrated into winged petiole, rounded, emarginate or very
obtuse at apex; the midrib generally hirtellous-pubescent beneath,
sometimes glabrous, the margin more or less conspicuously crenulate,

104 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

rarely entire, usually glandulose-punctate beneath; blades 3-12 cm.
long, 2.5-6 cm. broad. Petiole 0.5-2.5 em. long. The young
branchlets pubescent or puberulous, rarely glabrous.

The variety guianensis may be a medium-size tree, but it is most
commonly a small tree or shrub producing small blackish or blue-
black berries that are more or less sweet and flavorous. This variety
is very much spread throughout Surinam, British Guiana, the Amazon
territory of Venezuela, the Rio Negro-Guainia region of Columbia
and Brazil, and Rio Branco in Brazil, with small incursions into the
Para region. It is found in thickets and in marginal forests, and is
also isolated in campos, campinas, caatingas, andsavannas. Although
its typical form is very characteristic, intermediate forms transitional
to variety floribunda are very often found.

H. cassiquiari Suss. & Bergd. cannot be separated from variety guia-
nensis. The type, Lutzelburg 22627, does not differ in any way from
other abundant specimens seen of guianensis, which often have
a retuse or emarginate apex with a mucro in the bottom of the depres-
sion. This natural notch is often widened by some exterior cause,
such as insect eating or some kind of burning. The mucro, with
a glandular appearance at the tip, is seen only in young leaves, for it
usually breaks off. The petals in H. balsamifera sensu lato sometimes
end in a small glandular or pseudoglandular tip. This is more or less
apparent or inconspicuous and cannot be used as a taxonomic charac-
ter. The inflorescences of the typical specimens of H. cassiquiari are
very young and the few buds present are too underdeveloped to typify
the floral characters of a species. In 1 of these buds, 18 stamens
could be counted, but in some normal specimens of H. balsamifera
sensu amplo, I also have observed very rare flowers with some
reductions in the number of stamens.

COLOMBIA: Vaupsis: Raudal Guacamayo en el Rfo Infrida, margen izquierda,
180 m. alt., arbolito 4-5 m., frutos color verde, 4-II-1953, Ferddndez 2148 (COL,
US). Rio Negro, at confluence of Rivers Guainfa and Casiquiare, Cafio Ducuru-
apo, Igarapé Rana, tree, Schultes & Lépez 9363 (F, IAN). Rfo Negro a San
Felipe, alt. 200 m., 183-25-XI-1952, Humbert 27440 (US). Rio Atabapo, bushy
sprawling tree 1-5 m., flowers greenish white, frequent in savanna 1 km. west of
Cacagual, alt. 130 m., 19-XJI-1953, Maguire, Wurdack, & Bunting 36295 (NY,
US).

VENEZUELA: Amazonas: In sylvis humilioribus fluminis Guainia, XI-1954,
arbor ramosa 15-20 ft., Spruce s.n. (K). Rio Guainfa, frequent in sabanita 1 km.
west of La Ceiba, Cafio San Miguel, 2 km. above Limoncito, shrubby tree 2-5 m.
in bud, Maguire, Wurdack, & Keith 41917 (NY, US). Rfo Guainfa, riverine
forest just south of Maroa, tree, flowers white, fruit green, 28-XI-1953, Maguire,
Wurdack, & Bunting 36456 (NY, US). Orinoco, Cerro Yapacana, shrub or tree

to 5m., flowers white, locally frequent in savanna No. 3, northwest base of moun-
tain, alt. 125 m., 1-I-1951, Maguire, Cowan, & Wurdack 30543 (NY, US); occa-

CUATRECASAS—HUMIRIACEAE 105

sionally in savanna No. 1, northwest base of mountain, alt. 125 m., 7-I-1951,
Maguire, Cowan, & Wurdack 30791 (US, VEN). Cerro Sipapo (Paraque), open
shrub savanna on white sand, 3 km. southwest of Base camp, alt. 200 m., 8-II-
1949, Maguire & Politi 28828 (NY, US). Yavita, terreno abierto, arenoso,
128 m. alt., d4rbol pequefio o arbusto 2 m., flores amarillo pdlidas, fruto azulejo
oO casi negro, jugoso; corteza Aspera, rojo oscura, madera rosada, 22-I-1942,
Llewelyn Williams 13903 (US). ‘‘Pacimoni, Fl. Febr. 1854,” ‘Ad flumina
Casiquiari Vasiva et Pacimoni coll. R. Spruce, 1853-54,” Spruce 3409 (P,K,S).
I savanna prope Blaanve Berg, missit de Vacibe 1849, collector ? (8). Cassi-
quiare, Ufer, Laja de Caraga, orange, 4 m. 5-X-1928, Luetzelburg 22627 (M,
holotype of Humiria cassiquiari). Cassiquiare, Buenos Aires, Uferwald, gelb,
6 m., 6-X-1928, Luetzelburg 22561 (M, paratype). Laja de Caraga, Ufer, weiss,
9-X-—1928, Luetzelburg 22575 (M, paratype).

BRITISH GUIANA: Basin of Essequibo River, Kurupukari, slender tree
3 m., petals and filaments white, on burned forest land, 3-X-1937, A. C. Smith
2176 (A, F, NY, S, U, US). Upper Mazaruni River, IX-X-1922, De La Cruz
2210, 2227 (F, GH, MO, NY, US); De La Cruz 2202 (MO, NY). Demerara
River, Malali, 30-X-5-XI-1922, De La Cruz 2644 (MO); De La Cruz 2645
(F, GH, MO, NY, UC, US). Demerara River, Mahaica-Mahaicany, “muri,”
fruit edible, Forest Dept. (Anderson) 559 (K). Demerara, savanna plant, 21-IT-1910,
“muri,’”’ Forest Dept. (Anderson) 506 (K). Lungo il fiume Demerara, “muri bush’’
nella regione delle Canister Fall (Camo IV), 5—XI-1931, “muri,” Beccari s.n. (K).
Demerara, V-1889, Jenman 4883 (BM, K). Waranama Ranch, Wirumi-Ituni
savanna, Nkuri white sand area; a dominant bush in Nkuri-association; glaucous
leaves; small, inconspicuous flowers; fruits green, black when ripe, edible; Martyn
136 (NY). Kaieteur Savanna, Potaro River, IX-X-1881, tree 15 ft., Jenman
1281 (K). Kaieteur Plateau, occasionally shrubby, procumbent, 1 m. high,
clump, flowers white, buds light green, 14-V-1944, Maguire & Fanshawe 23450
(NY, U, US). Rupunini River, Qumatta, X-1889, Jenman 5561 (NY), Jenman
5562 (NY, US). Rupununi River, near Apoteri, alt. 300 ft., small tree with
flat spreading crown up to 20 ft., in savanna caused by fire on white sand soil
liable to inundation, Forest Department 2055 (K). Corentyne River, sand
ridge woods, tree 20 ft. or more, Jenman 299 (P). Savannas east of Itumi, 35
miles south of Makenzie, frequent in margin of forest “islands,” tree, flowers
white, fruit green, 17-I-1955, Cowan 39266 (NY); semi prostrate tree, Cowan
39263 (NY, US). East Coast Water Conservancy, southeast of Georgetown,
sand reef at head of Hoorubia Creek, sandy scrub, 26—XI-1919, Hitchcock 16938
(GH, NY, 8, US). “British Guiana,” Schomburgk 270 (P, NY, US, isotypes),
photo F.M. 35176 at P. “British Guiana, Oct. 19,” shrub 5-12 ft., fruits
greenish white, collector ? (P).

SURINAM: Boschreserve, Sectie 0, 3-VIII-1920, witte bloeme, shrub of
savanna, Pulle 150 (U); 20-XII-1906, Boschwezen 36a (U); 21-II-1918, Bosch-
wezen 3646 (IAN, U); Boomnumer 842, 23-IV-1917, Boschwezen 2816 (U);
20-XII-1916, Boschwezen 2599 (U); 8-VIII-1918, Boschwezen 3947 (U); 2-VIII-
1917, Boschwezen 3040 (U); Zanderij I, 26-IV-1915, Boschwezen 531 (U).
Zanderij I, 10-XI-1921, Boschwezen 5486 (U). Zanderij I savanna a, struik
14% m., vrucht groen, onrijp, “blakaberie,” 10-III-1949, Lanjouw & Lindeman
3259 (U). Zanderij I, savanna, flowers white, shrub, 22-VII-1923, Lanjouw 195
(U, US). Zanderij I, savanna bij vliegveld, struik, bloem groenachtig-wit,
9-IX-1948, Lindeman 258 (U). Zanderij I, savanna voor vliegveld, struik bloem
wit, 9-IX-1948, ‘“‘tauraroe”’ (Arow.), Lanjouw & Lindeman 267, 268 (U). Zanderij

106 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

I, savanna, sand, flowers white, fruit green, shrub (may become a large tree),
“blakkaberie,’ 30-VII-1933, Lanjouw 334 (U). Zanderij I, indianenkamp,
“tawararoe”’ (Arow.), ‘meri’ (Kar.) ‘‘blakkaberie’ (NE), 28-VII-1920, Pulle
52 (U). Zanderij II, zansavanne, 16-III-1949, struik 2-3 m., vrucht groen,
onrijp, ‘‘blakkaberie” (Sur.), Lanjouw & Lindeman 3289 (U). 4anderij I, grass
savanna, frequent, shrub to 3 m., flowers white, 3-VI-1944, Maguire & Stahel
23654 (NY, US, U); Maguire & Stahel 23696 (A, NY, U, F, US, VEN). Forest
of Zanderij, 31-V—-1916, Samuels s.n. (US). Coppename R., Onobisi Kreek,
4-III-1915, Boschwezen 1120 (U). Plantage Berlijn, savanna, flores albi, Focke
1018 (isotype of H. surinamensis, U). Jodensavanne-Mapane Kreek area
(Suriname R), op droge zadsavanne aan Blakawatre kreek, Kruiptruik als rand
om hogere struik groepen, 17—XII-1954, Lindeman 6881 (U); op droge zand-
savanne bijaan Blakawatre Kreek, Kruipstruik, bloemwit, “blakaberie” (Sur.)
17-XII-1954, Lindeman 6882, 6883 (U). Jodensavanne, V~—1909, flores albi,
Focke 1286 (U). Tibitisavanne; savanne bijkm. 3.1 tweede Lijn; struik 4 m.;
bladen meer elliptisch, vrij spits, evenals takken witstaand, ‘“‘blakaberie”’ (Sur.)
11-I-19494, Lanjouw & Lindeman 1798 (U); struik 4 m.; bladen breed en
stomp, steil opgericht aan opgerichte takken, ‘“‘blakaberie” (Sur.), Lanjouw &
Lindeman 1797 (U). Patrichsavanne, 2-X-1909, Boldingh 3886 (U). Via secta
ab Moengo tapoe ad Grote Zwiebelzwamp, in bos bij km. 15.75, boom 12 m,
17 cm. dik; bloem wit, “blakaberie” (Sur.), ‘‘merie”’ (Car.), 20-X-1948, Lanjouw
& Lindeman 911 (U); it is an atypical form. Op natte zandsavanne bij Kopie,
Cassewinica, struik 2-3 mm. hoog, bloem wit, ‘“‘blakaberie’”’ (Sur.), 16-VII-1953,
Lindeman 4381 (U). Savanne bij km. 100 langs spoorlijn; struik 3-4 m. hoog;
vrucht groen, jong, ‘“‘blakaberie” (Sur.), 16-III-1949, Lanjouw & Lindeman 3317
(U). Savanne, 9-VIII, Kuypper 568 (U); 11-X-1911, Kuypper, 33 (U). Without
locality, Hostman 793 (P, BM, GH, F, MO, U); bl. wit, savanna, Wallschlaegel
1393 (U). Splitgerber s.n. (BM).

BRAZIL: Rio Branco: Vista Alegre, in campis, frutex floribus albis odoratis,
fr. nigro edule, “umiry,” Kuhlmann 2894 (US). Amazonas: Rio Igana, Tunuf,
capoeira; arvore de porte muito reduzido, em cumes esteris e pedregosos, 10-V—
1948, Black 48-2589 (IAN, NY, VEN). Rio Igana, Santana, caatinga, arvore
baixa, flor branca, 2-V—1948, Black 48-2512 (IAN); pe da serra, terreno pedrogoso,
arbusto, flores, esbranquicadas, 24-X-1947, Pires 754 (IAN). Manaos, Chapada,
terra firme arenosa, capoeira, arvore de 5 m., flor amarelo esverdeada, perfumada,
19-VIII-1955, INPA 1684 (MG, US). Manaos, Strauch 1-4 m., Blumen weiss-
lich, V-1902, Ule 6142 (K, MG). Alto rio Aracd, subafluente do Rio Negro,
mata alta de caatinga en formacio, terreno plano, solo arenoso, 30-X-1952,
Frées & Addison 29211 (IAN). Itaubal, Rio Aracd, 26-X—-1952, Frdées & Addison
29102, 29096 (IAN). Rio Negro, Preto, Campina tree 25 ft. X10 in., caatinga,
white flowers, 5-XI-1947, Frées 22760 (IAN, NY, U, US, VEN). Preto, Malu-
piry, tree 30 ft. 7 in., greenish white flower, yellow center, sweet smell, open
country campina, 13—XI-1947, Frées 22838 (IAN, NY, US, U); Frées 22842
(IAN, NY, U). Rio Negro Aiary-pird, rio Aiary, arvore 5 m., caatinga arenosa,
5-XI-1945, Frées 213842 (IAN, F, NY, US). Alto rio Aiary, arvore pequena,
flores, alvas forte cheiro agradavel, 5-XI-1945, Frées 21338 (IAN, NY, US).
Campo on River Paduira, 28-VI-1874, flowers white, Traill 80 (K). Manaos,
300 ft., tree with pale flowers slightly perfumed, VIII-1942, Sandeman 2199 (K).
Pard: Lago de Fero, Praia, arbusto grande, “umiry”, 20-VIII-1907, Ducke
8410 (BM, MG). Rio Mapuera, campinara a noreste do Tabolierinho. Arbusto,
“umiry,’”? 12-XII-1907, Ducke 9123 (MG).

CUATRECASAS—HUMIRIACEAE 107

le. Humiria balsamifera var. laurina (Urb.) Cuatr., comb. nov.
Ficures 21,j; 22,f,n; 24,d

Humiria floribunda var. laurina Urb. in Mart. Fl. Bras.12(2):4389. 1877.
Humirium laurinum Klotsch ex Urb. in Mart., ibid., as synonym.
Humiria floribunda var. spathulata Gleason, Bull. Torrey Club 58:374. 1931.

Type: Schomburgk 560, British Guiana.

Leaves elliptic-oblong, abruptly contracted into petiole at base,
rounded, spatulate or obtuse at apex; entire at margin, sometimes
with dotted glands beneath, glabrous; blades 3-9 cm. long, 1-3 cm.
broad; petiole 5-15 mm. long. Inflorescences small, pauciflorous.
Petals puberulous in the typical form, also glabrous. Young branch-
lets glabrous or glabrate.

H, balsamifera var. laurina is spread throughout the Gran Sabana
of Venezuela, British Guiana, entering Colombia through the Llanos,
and Brazil in some places of Paré, Mato Grosso, and Maranhio.
Usually a small tree or shrub growing in open savannas or in thickets,
it becomes a large tree when in the rain forest. The berries are
reported to be black or purple-black when ripe and edible.

COLOMBIA: Vaupfs: Rio Parand Pichuna, 700 ft, bush, VI-1953, Schultes
& Cabrera 19951 (US); low bush, VI-1953, Schultes & Cabrera 19963 (US). Rio
Infrida, Raudal Alto o Mariapiri, margen derecha, 250 m. (cerro del varador),
3-II-1953, Ferndndez 2084 (COL, US), a broad-leaf form tending toward var.
guianensis.

VENEZUELA: Amazonas: Esmeralda, XII-1853, Spruce 3419 (P, K).
Esmeralda, Gran Sabana, section I, alt. 325 feet, 1-XI-1928, Tate 283 (NY);
shrub with red berries, XI-1928, Tate 286 (NY, holotype of H. floribunda var.
spathulata Gl.). Esmeralda, upper Orinoco, 1438 m., sabana abierta; Arbol
pequefio 4-5 m., copa de forma irregular, tupida, tronco redondo, 10 cm., didm.,
corteza gris oscura, aspera, fisurada, liber color castafio claro, albura, también de
tono claro, duramen de castafio a rosado fuerte o colorado; fruto ovoideo y segtin
los nativos de color negro en la madurez y comible, ‘“‘nifia,”” 16-V—1942, Llewelyn
Williams 15418 (US, VEN). Between Esmeralda Savanna and southeastern
base of Cerro Duida, 200 m., 22-VIII-1944, Steyermark 57817 (A, US, VEN).
Sabana Grande, between Esmeralda and base of Cerro Duida, 175 m., abundant
in savanna, shrub 0,.2-2 m. tall, forming colonies, 2-5 m. across, 24-III-1953,
Maguire & Wurdack 34677 (NY, US). Tyler-Duida expedition, rocky top of
Esmeralda Ridge, small tree, 6-X-1929, Tate 209 (NY, US). Rio Guainfa,
Savanna El Venado, left bank of Cafio Pinichin, above Pinichin, alt. 140 m.,
occasional shrub, 0.3-0.7 m., 14-IV—-1953, Maguire & Wurdack 35579 (US, NY);
alt. 120-140 m., frequent, sprawling shrub to 1 m. tall, flowers greenish white,
fruit green, 23—-XI—-1953, Maguire, Wurdack, & Bunting 36354 (NY, US); abundant
2 km. above Pinichfn at margins of savanna, shrub sprawling to 3 m., flowers
greenish white, 10O-X-1957, Maguire, Wurdack, & Keith 41821 (NY, US). Casi-
quiare, Rfo Siapa, occasionally in Sabana Hechimoni on left bank of Cafio Hechi-
moni, shrub 0.5-2 m., flowers white, fruit dark purple, 9-IJ-1954, Maguire,
Wurdack, & Bunting 37632 (NY, US). Orinoco, San Antonio, 121 m., alt.,
arbusto tendido de 60 cm. o mas de altura; fruto pequefio, ovoideo, verdoso;
sitios arenosos en el margen de la sabana, 27-IV-1942, Llewelyn Williams 15052
(US).

108 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

BRITISH GUIANA: Kuyunun, IV-1842, Schomburgk 560 (M, holotype; NY,
US, isotypes). Roraima, Schomburgk 628 (BM, P). Pirara, Schomburgk 346 (P).
Parinari mountains, Schomburgk s.n. (KX), not a typical specimen; Schomburgk 968
(M), not typical. Canje River, Ikuruwa Island (County Berbice), ‘‘tauroniro’’
(Arawak), Forest Depart. (Hohenkerk) 663 (IX).

BRAZIL; Maranudo: Grajaht, beira do Rio, Campo Cerrado; arvore alta,
flor amarella; a fructa é procurada pela pomba amargosa, “umiry,” 13-VITI-1909,
Lisboa 2327 (BM, MG); this collection is a puberulous form with abundant hairi-
ness on inflorescences and the underside of leaves (midrib). Parf: Rio Sao
Manoel, Posto dos Indios Caiabi, arbusto de campo, 8-I-1952, Pires 3877 (IAN,
US). Serra do Cachimbo, 425 m., 14—XTII-1956, Pires, Black, Wurdack, & Silva
6209 (IAN). Marro Grosso: Near Tabajazta, upper Machado River Region
shrub on grassland, flowers white, XI-XII-1931, Krukoff 1483 (A, BM, NY,
P, 5, U).

If. Humiria balsamifera var, sayannarum (Gleason) Cuatr., comb. nov.
FIGURE 21,v

Humiria savannarum Gleason Bull. Torrey Club 58:378. 1981.

Type: Tate 330, second sheet at NY, Venezuela, Amazonas, Esme-
ralda.

Leaves linear-oblong or subelliptic-oblong, narrowed-cuneate at
base, sessile or subpetiolate, 5-10 cm. long, subacuminate and suddenly
obtuse at apex, frequently emarginate and minutely mucronulate, the
margin entire with some or no glands beneath, 5-10 cm. long, 1.5-3.5
em. broad. Petals puberulous outside in the type specimen (the others
lack flowers). Fruit oblong-ellipsoid, about 148 mm.

This variety is a narrow-leaf form of H. balsamifera found on the
savannas of Esmeralda and the low Roraima Mountains. The typical
leaves are narrow-sublanccolate with blunt apex and cuneate base
with a 0-2 mm. long petiole. The forms with smaller and thicker
leaves (e.g., Shomburgk 628) are transitional to variety corzacea.

VENEZUELA; Amazonas: Esmeralda, alt. 325 ft., 2-XI-1929, Tate 330,
second sheet (NY, holotype). Botfvar: Frequent on eastern slopes of Cerro
Marimarota (Cerro La Puerta), 100-250 m., shrub 3-6., fruit green, 26-I-1956,
Wurdack & Monachino 41380 (NY,US).

BRITISH GUIANA: Roraima, 1842-438, Schomburgk 576 (P,K). ‘Br. Guiana,
Oct. 1842,” Schomburgk 845 (P, K).

Ig. Humiria balsamifera var, parvifolia (Juss.) Cuatr. comb. nov,

FIGuRES 22,0—p; 24,e; PLates 6-7
Humirium parvifolium A. Juss. in St. Hil. Fl. Bras. Merid. 2:89, 1829.
Humirium montanum A. Juss. in St. Hil. ibid. 2:90. 1829.
Humirium parviflorum A. Juss. in St. Hil. ibid. 2:63. 1829.
Humirium arenarium Guill. in Baill., Adansonia 1:208, 1861.
Hunurtia floribunda var. parvifolia Urb. in Mart. Fl. Bras, 12(2):438. 1877.
Humiria floribunda var. montana Urb, in Mart. ibia,

Type: St. Hilaire, Brazil, Rio de Janeiro.
Leaves small, subcoriaceous or coriaceous, glabrous, obovate,
obovate-elliptic or oblong-obovate, narrowed toward the base, cuneate,

CUATRECASAS—-HUMIRIACEAE 109

short-petiolate or sessile, rounded, truncate or very obtuse, emarginate
at apex, entire or obsolete-crenulate, the secondary nerves slightly
conspicuous; 1.5-4 (rarely 5) cm. long, 0.8-2.5 cm. broad. Young
branchlets generally angulate or narrow-winged and_hirtellous-
puberulous.

There is no type cited for Humirium arenarium Guillemin (in
schaedula) as was informally described by Baillon. Several speci-
mens of the Paris Herbarium labeled as H. arenarium belong to the
variety parvifolia. The notation ““M. Guillemin Cat. 205, Restingas
de Tocaia, Humirium arenarium Nob” of one specimen must have
been written by Guillemin himself, and I consider it to be the type.

This variety with small leaves is a small tree or shrub, and is fre-
quent in “‘catingas,” “restingas,” and “serras” of eastern Brazil, from
Pernambuco to Matto Grosso and Rio de Janeiro. St. Hilaire noted
that he collected the type of H. parvifolium “prope urbem Cabo Frio
in prov. Rio de Janeiro” and the type of H. montanum “prope vicum
Itambe in montibus provinciae Minas Geraes”’ (pp. 89-90). Curiously,
the same variety (or a very close form) has been found in eastern Peru.

BRAZIL: PernamsBuco: Recife, fazenda Modelo Tigipié, arbusto pequeno,
flores alvo crema perfumadas, 12-III-1950, Lima 1623 (IAN). Gords: Serra
dos Viadeiros, arvore pequena na margen do um corrego, [X-1892, Glaziou 63 (P).
Banta: “Circa Bahiam,” Blanchet 1005 (NY, BM); Blanchet 3144A (P). “In
Provincia Bahiac,’”’ Blanchet 3570 (BM, P). Serra de Acurua, Blanchet 2810 (F,
BM, NY, US, paratypes of H. montanum). Rio San Francisco, Blanchet s.n.
(GH). Saubino, Jacobin, Blanchet 3422 (US, F, paratypes). Inter Vitoria et
Bahia, Sellow 2212, 2228, 180 (NY, US,8, BM). Bompui, Curran 159 (Y). Inter
Campos et Victoria, Sellow s.n. (K). Minas Gerais: ‘Minas, septembre fl,”
St. Hilaire s.n. (P, isotype of Humirium montanum Juss.). Cocais, arvor da
Serra, flor alva, Hoehne 7970 (NY). Chiribiry, arbrisseau, fleurs blanchAtres,
29-III-1892, Glaziow 18962 (P). Serra do Cipé, km. 132, arvore pequena,
6-IV-1951, Black & Magalh@es 51-11790 (IAN). Serra do Cipé, 16-I-1951,
Pires & Black 2961 (IAN). Catingas de Tocaia, XII-1838, “Humirium arenartum
Nob. medium inter H. montanum et H. parvifolium,’’ Guillemin 205 (P, type
of Humirium arenarium Guill.). Rio pe Janrerro: No locality, St. Hilatre
114/5 (P, isotype of Humirium parvifolium Juss.). Rio de Janeiro, Glaziou
10342 (P). Rio de Janeiro, Herb. Brognfard, Herb. Richard, (P, 8). Rio de
Janeiro, Glaziou s.n. (S). Restinga de Maua, 2-V—1891, arbrisseau trés touffu,
Glaziou 18180 (P, F, US, NY); Glaziou 8286 (A, P, 8). Maud, 15-XII-1874,
Glaziou 7765 (P). Restinga da Tijuca, 25-XI-1866, Glaziou 731 (P), 29-VII-
1872; Glaziow 6196 (US, P). Cabo Frio, Praia do Pontal, ao nivel do mar,
17-IV-1952, L. B. Smith 6694 (US). Cabo Frido, 19-IX—1881, Glaziou 12515.
(NY, UC, P). Cabo Frio Co., Arraial do Cabo, Pontal Beach, 1953, Segadas-
Vianna, et al. I-1383 (US); II-III-1951, Mello Filho L, E, 1186; restinga, 3-VI-
1953, arbore 3 m., calice branco esverdeado, corola branca, Segadas-Vianna, et al.
I-439 (US); na fimbria de moita da restinga interna, 13-VIII-1953, substratum
arenoso, arbusto 1 m., caliz verde, petalos brancos, Segadas-Vianna et al. I-821
(US); na restinga interna, 24-V-1953, substratum arenoso, plano, mesofanerofita
comun, en colonias, arbusto ramificado da base em moita arredondada de 1.50
a 2 m., calice verde, corola branca, estambres filete branco, antera branca.,

110 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Segadas-Vianna et al. I-310 (US). Casimiro de Abreu Co., District of Barra de
Sao Jodo, near Barra de Séo Joao Village, restinga interna, 3-IX-1953, calice verde,
corola 5 petalos, branca, estames filete branco, antera branca, Segadas-Vianna et
al, I-907 (US); 5 km. north of Rio das Ostras borough, 5-I X-1953, Segadas-
Vianna et al, I-1416 (US); 1 km. ao sul do Povoado de Rio das Ostras, inland
restinga, 28-V-1953, Segadas- Vianna et al. I-369 (US); 3 km. norte do Povoadode
Rio das Ostras, nas moitas ou isolada na restinga interna, Segadas-Vianna et al.
I-385 (US); 5 km. ao norte do Povoado de Rio das Ostras, 29-V-1953, na restinga
arborescente ate 5 m., 4-IX-1953, Segadas-Vianna et al. I-945 (US). Distrito
Federal, Restinga de Itapeba, Recreio dos Bandeirantes, 1-X-1950, ao nivel do
mar, Segadas-Vianna 3506 (US); 29-X-1950, Segadas-Vianna 3634, 3635 (US).
Recreio dos Bandeirantes, XI-1931, Lutz 681 (US); 22-X-1938, Alston & Lutz
169 (BM). Praia de Sernambetiba, ao nivel do mar, 4-IX-1952, L. B. Smith
6406 (NY, US). Rio de Janeiro, 1836, Gomes s.n. (K). Tocojd, ‘Herbarium
Zucearinii, comm. M. C. Vendob, anno 1809, legit in Brasilia Dr. Pohl,’? Pohl
s.n.(M). Macahé, 1845, Herb. John Miers 8915 (BM). Without locality, Riedel
s.n. (GH, K, M, P, US); Riedel 3570 (U, 8, in part).

PERU: San Martin: Zepalacio, near Moyabama, alt. 1,100 m., mountain
forest, tree 8 m., flowers pale vellow, VIII-1934, Klug 3706 (A, GH, BM, F, MO,
NY, 8, US). In monte Campana prope Tarapoto orientalis, VIII-1856, Spruce
4335 (GH, BM, 8).

lh. Humiria balsamifera var. coriacea Cuatr., var. nov.
Fraures 21,0-r; 22,7-]; 23,e-g; 24,b; PLare 8

Arbuscula vel arbor 4-8 m. alta ramosa ramulis terminalibus
crassis nitidis minute hirto-puberulis vel glabris.

Folia crassa rigideque coriacea. Lamina obovata vel obovato-
elliptica apice rotundata vel obtusa breviter emarginata et in sinu
mucronulata, basi cuneata subite vel gradatim in pseudopetiolum
alatum 2-6 mm. longum attenuata cuneata, margine integra plana
eglandulosa vel paucis glandulis, 4-7 em. longa, 2.4-5.2 em. lata,
utrinque glabra supra viridis nitidula nervo medio signato secundariis
obtuse prominulis venulis reticulatis paulo vel haud conspicuis;subtus
costa crassa elevata nervis secundariis circa 4-5 mm. inter se dis-
tantibus subparallelis prope marginem arcuato-anastomosatis nervis
minoribus parallelis et venulis prominulis anastomosato-reticulatis.

Inflorescentiae cymoso-paniculatae corymbiformes subterminales et
axillares folia aequilongae vel breviores, axi ramulisque angulosi vel
hirto-puberuli; bracteis late ovatis ovato—triangularibus subacutis vel
obtusiusculis amplexicaulibus inferioribus ad 2 mm. longis superioribus
1 mm. longis crassiusculis glabrisque. Ramusculi cymorum ultimi
crassi angulati hirtulo-puberuli breves ad 1 mm. longi. Pedicelli crassi
glabri 0.5-1.5 mm. longi. Calyx cupularis circa 1 mm. altus crassus;
sepalis usque ad medium coalitis depresso-rotundatis glabris margine
minutissime ciliatis excepto. Petala 5 libera, crassiuscula oblonga
subacuta glabra circa 5 mm. longa 1.4 mm. lata. Stamina filamentis
3-4.5 mm. longis infra medium in tubum glabrum coalitis parte libera
minute papillosis; connectivo antheris oblongo linguiformi subacuto

CUATRECASAS—HUMIRIACEAE 111

0.8 mm. longo thecis orbicularibus barbatisque. Squamae disci 20
crassiusculae in tubulum argute dentatum coalitae. Ovarium obo-
vatum glabrum vel apice leviter pilosum 5-loculare loculis 2 ovulis
superpositis pendulis. Stylus 3-4 mm. longus hirtulus. Stigma
capitatum 5-lobatum. Drupa oblongo-ellipsoidea 10-14 mm. longa
5-8 mm. crassa, exocarpio carnoso. Endocarpium durum circa
10 * 5 mm. 10-striatum sublaeve apice leviter attenuatum
5-foraminatum.

Type in the U.S. National Herbarium, No. 1901203, collected in
rocky soil at Mureyena Falls, alt. 800 m., State of Bolivar, Venezuela,
October 10, 1946, by {Félix Cardona, No. 1828. Isotypes in the
Herbarium of the New York Botanical Garden and in the Instituto
Botdnico of Venezuela, Caracas. Paratype at U.S. National Herba-
rium, No. 1909308, collected at Rio Caronf, Bolivar, Venezuela, by
Félix Cardona, No. 1768.

Shrub or small tree with hirtellous or puberulous, sometimes
elabrate, young branchlets. Leaves thick and rigid, coriaceous,
glabrous, obovate or obovate-elliptic, rounded or obtuse, emarginate
and mucronulate at apex, attenuate in winged, 2-6 mm. long petiole
at base, midrib prominent beneath, the secondary nerves and reticulum
slightly conspicuous or obsolete above, prominulous beneath; 4-7 cm.
long, 2.4-5.2 em. broad. Branchlets of the inflorescence hirtellous-
puberulous, the pedicels (0.5-1.5 mm. long) glabrous. Calyx cupular,
glabrous except for the ciliate margin. Petals oblong, subacute,
glabrous. Style 3-4 mm. long, hirtellous. Drupe oblong-ellipsoid,
10-14 < 5-8 mm., the endocarp 10 X 5 mm., striate, 5-foveolate
at apex.

The rigid sclerophyllous texture of the leaves, and their shape and
size are the most distinctive characters of this variety, which seems
to conform to the most xerophytic conditions of the sandstone and
granitic hills of the savanna.

H. balsamifera var. coriacea is endemic to the Guiana hills where it is
very common. It is a characteristic tree or shrub of the vegetation
of these mountains, it being found often abundantly on the slopes
and at the top of the hills where it forms actual stands. It is also a
constituent of the open savanna vegetation. It is especially found at
between 700 and 2,000 m. elevation, but it is present at much lower
altitudes (as, e.g., 350 m.).

VENEZUELA: Amazonas: Cerro Duida, Rio Cunucunuma, Culebra Creek
drainage, alt. 1,500 m., tree to 5 m., flowers white, 19-XI-1950, Maguire, Cowan,
& Wurdack 29541 (NY, US). Ridges of Cafio Negro Basin, alt. 2,000—-2,300 m.,
abundant in scrub sabana, tree 4 m., flowers white, 23-XI-1950, Maguire, Cowan,
& Wurdack 29697 (NY, US). Cerro Duida, Savanna Hills; alt. 1,020-1,200 m.;

shrub 10-15 feet tall, leaves coriaceous, dark green above, dull green below; fruit
blackish, flowers greenish, Steyermark 58288 (NY, US). Summit of Savanna

112 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Hills, bushy tree, Tate 733 (NY, US). Cerro Sipapo (Pardque), savanna breaks,
Peak 1, alt. 2,000 m., shrub or small tree, 12-XII-1948, Maguire & Politi 27627
(US, NY); frequent in marsh at savanna camp, alt. 1,500 m., much branched shrub
to 4m, tall, 15-XII-1948, Maguire & Politi 27695 (NY, US). Serrania Yutaje,
Rfo Manapiare, cerro Coro-Coro, frequent in savannas, alt. 1,500 m., rounded
shrub 1-3 m. tall, flowers white, 2-III-1953, Maguire & Maguire 35453 (NY, US).
Cerro Yutaje, occasional in scrub forest on northwest ridge, alt. 1,400 m., bushy
rounded tree 2-5 m. tall, flowers white, 11-II-1953, Maguire & Maguire 35140
(NY, US). Cerro Yapacana, frequent on summit at alt. 1,200 m., tree to 5 m.,
flowers white, 2-I-1951, Maguire, Cowan, & Wurdack 30622 (US, NY); this tree
is a form having smaller leaves, but it can hardly be separated from var. coriacea.
Rfo Cunuecunuma, Cerro Huachamacari, summit camp at alt. 1,800 m., occasion-
ally near camp, much branched tree 8 m., flowers white, 6-XII-1950, Maguire,
Cowan, & Wurdack 30018 (NY, US). Rfo Ventuari, Serranfa Part, Rio Paru,
Cafio Asisa, alt. 2,000 m., occasionally on summit along west rim, tree 3 m.,
flowers white, 7-II-1951, Cowan & Wurdack 31301 (NY, US); bushy tree 2 m.,
buds turning white, 31-I-1951, Cowan & Wurdack 31090 (NY, US); abundant in
open areas in cumbre, alt. 1,800 m., shrub 1 m., flowers greenish, Kathleen Phelps
& Hitchcock 508 (NY). Cerro Moriche, frequent in cumbre at alt. 1,250 m.,
15-I-1951, Maguire, Cowan, & Wurdack 30918 (NY, US); frequent in montane
savanna at alt. 800 m., tree 5 m., fruit turning brown-red, 14-I-1951, Maguire,
Cowan, & Wurdack 30885 (NY, US). Bottvar: Gran Sabana, between waterfall
at Rue-meru and Divina Pastora on Rfo Cuquenan north of Santa Elena, south
of Mount Roraima, alt. 1,065 m., dry sandy upland; shrub 4-6 ft. tall; leaves
subcoriaceous, deep green above, pale green below; petals white; Steyermark
59186 (NY, US, VEN). Rfo Caronf, playas rocosas, aguas arriba del salto
Kutouamini, 720 m. alt.; 4rbol 5 m., flores gris claro, Cardona 1768 (US, paratype).
Rio Caronf, lugares rocosos frente a los raudales Mureyena, 800 m. alt., drbol 6 m.,
flor amarilla, 10-X-1946, Cardona 1823 (US, holotype; NY, VEN, isotypes).
Rio Caronf, cumbre del cerro Acopdn, 2,200 m. alt., arbusto 6 m., X-1947, Cardona
2269 (US). Lomas pedregosas entre el Cerro Peraf y las fuentes del rio Uaipard
afluente del Icabart, 900 m., alt. arbol 5 m., flores blancas, 20-X-1946, Cardona
1869 (NY, US). Lomas pedregosas del rfo Icaburt, 460 m., arbusto 4 m., V-48,
Cardona 2533 (VEN). Sabanas de Icaburt, hacia El Caribe, 450-850 m. alt., Arbol
aislado 10 m., flores blaneas, 22-XTI-1955, Bernardi 2603 (VEN). Alto Caronf,
rfo Carrao, 350 m. alrededores Salto Hacha, 350 m. alt., en playas, arbolito 4
m., Cardona 2877 (US, F). Auyantepuf, 2,100 m., alt. arbusto 4 m., I-1949,
Cardona 2670 (NY). Mount Auyan-Tepuf, 1,100 m. alt., Tate 1113 (NY, VEN).
Auyantepuf, campamento de Guayaraca 1,100 m., alt. crecen en Barbacenietum,
formacién seca arbustiva, cerca del campamento, IV-1956, Vareschi & Foldats
4563, 4573 (VEN). Chimanté Massif, Torono-tepuf, summit along Cafio Mojado,
between base of upper falls and drop to escarpment at alt. 1,895-1,910 m., tree
4 m., buds green, flowers greenish white, fruit green, locally frequent, 23-II-1955,
Steyermark & Wurdack 1109 (NY, US). Alto Rio Paragua, Cerro Perro, in sand-
stone savannas, Cardona 774 (NY, US, VEN). Cerro Bolfvar, frequent on
summit of West Peak at alt. 750 m., rounded much-branched small tree, flowers
white, 2-XII-1951, Maguire 32686 (NY, US). Cerro Bolivar, Ciudad Piar, 750
m. alt.; variable desde arbusto 2 m. a drbol de 8-12 m., siempre muy ramificado,
troneo libre muy corto; es una de las especies caracteristicas del tope del Cerro
Bolivar I1V-1954, Aristeguieta 2174 (VEN). Sucre: Lago de Guanoco, crece en
pequefias islas formadas por el levantamiento del suelo, arbusto, flores blancas,
VITI-1955, Lasser & Vareschi 3888 (VEN).

CUATRECASAS—-HUMIRIACEAE 113

BRITISH GUIANA: Kaietur Plateau, from bush island, in savanna, frequent,
bushy crowned tree 4-6 m., 4-8 cm., flowers greenish white, 8-V-1944, Maguire &
Fanshawe 23295 (NY, U, US, VEN). Kaietur Plateau, Potaro River, on sand-
stone flat in the open, alt. 1,300 ft., a tree 12 ft., calyx green, corolla greenish
white, Alison 545 (K, P); tree 20 ft. IX—X-1881, Jenman 1023 (KX).

SURINAM: Tafelberg, vicinity Camp No. 1, savanna, alt. 300 m., shrub 4
m., much-branched and rounded, flowers white, 4-VIII-1944, Maguire 24223
(NY, US, U, VEN). Tafelberg, Savanna No. II, frequent, shrub to much-
branched rounded tree of 7 m. height, 12-IX-1944, Maguire 24707 (NY, U, US).
Savanna VII, common shrub or rounded, much-branched-tree to 10 m., 25 cm.
diam., 17-IX-1944, Maguire 24789 (NY, U, US). Savanna VIII, frequent
shrub or bushy small tree to 8 m., 10 cm. diameter, flowers white, extensive rock
and pigmy bush openings, alt. 776 m., 23-VIII-1944, Maguire 24443 (NY, US, U).

BRAZIL: Rfo Branco: Serra Tepequén, dominant on unburnt portion of rim,
alt. 1,000—1,200 m., bushy rounded tree, 5-8 m. tall, leaves glossy green, flowers
white, 4-XII-1954, Maguire & Maguire 40159 (NY, US). Regiéo entre o Rio
Mat e o Canan, arvore de flor esverdeada, 22-XI-1954, INPA 620 (MG).

li. Humiria balsamifera var. guaiquinimana Cuatr., var. nov.
Figure 21,w; 24,a; PLars 9

Arbor parva usque 8 m. alta ramulis terminalibus crassis minute
hirto-pubescentibus deinde glabris griseis fissuratis nitidis. Folia
crasse rigideque coriacea subsessilia. Lamina suborbiculato-ellip-
tica vel suborbiculata apice rotundata vel obtusissima saepe retusa
et emarginata in sinu longe mucronata, basi subite lateque cuneata
subpetiolata contracta, margine integra plana eglandulosa, 4—7 cm.
longa 3-5.2 cm. lata, utrinque glabra supra viridis nitidula nervo
medio signato secundariis obtuse prominulis venulis reticulatis paulo
conspicuis; subtus costa crassa elevata nervis secundariis prominulis
circa 4-5 mm. inter se distantibus subparallelis prope marginem ar-
cuato-anastomosatis nervis minoribus parallelis et venulis prominulis
anastomosato-reticulatis.

Inflorescentiae cymoso-paniculatae subterminales et axillares folia
breviores, axi ramulisque angulosi pubescentes; bracteis late ovatis
amplexicaulibus inferioribus ad 3 mm. longis superioribus 1 mm.
longis crassiusculis glabrisque. Ramusculi cymorum ultimi crassi
pubescentes breves ad 1 mm. longi. Pedicelli brevissimi circa 0.5
mm. longi. Calyx circa 1.2 mm. longus; sepalis usque ad medium
coalitis rotundatis glabris margine minutissime ciliatis excepto.
Petala 5 libera, elliptico-oblonga crassa glabra in alabastro 3.5 mm.
longa. Stamina basi in tubum glabrum coalita, parte libera minute
papillosa; connectivo antheris ovato compresso, thecis orbicularibus
barbatisque. Ovarium glabrum apice leviter piloso excepto.

Type in the U.S. National Herbarium, No. 1878323, collected at
Cerro Guaiquinima at the headwaters of Rio Paragua, alt. 1760 m.,
in the State of Bolfvar, Venezuela, July 15, 1944, by Félix Cardona
(No. 1112).

114 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Small tree with thick, hirtellous terminal branchlets becoming
glabrous. Leaves thick, rigid-coriaceous, glabrous, subsessile, sub-
orbiculate-elliptic or suborbicular, rounded or very obtuse at apex,
often retuse and emarginate, and mucronate, abruptly attenuate,
subpetiolate-cuneate at base, the margin flat, entire, eglandular,
the midrib prominent beneath, secondary nerves and reticulum promi-
nulous beneath. Petals elliptic-oblong, glabrous. The thick, stiff,
glabrous, elliptic or suborbicular leaves only very shortly, abruptly
cuneate-contracted at base (pseudopetiole 1-2 mm. long) and hir-
tellous young branchlets differentiate this variety, which is restricted
to Mount Guaiquinima.

VENEZUELA: Borfvar: Alto Rio Paragua, Cerro Guaiquinima, alt. 1,760
m., 15-VII-1944, Cardona 1112 (US, holotypus); alt. 1740 m., X-1943, Cardona
965 (F, NY). Cerro Guaiquinima, dominant in quebradas near Cumbre Camp,
alt. 1,800 m., densely rounded tree to 8 m. tall, leaves glossy, 25-XII-1951,

Maguire 32763 (NY, US); rounded tree 3-10 m. tall, dominant along west escarp-
ment, alt. 1,800 m., 14-IV-1952, Maguire 33099 (NY, US).

1j.. Humiria balsamifera var. stenocarpa Cuatr., var. nov.
Fiaures 21,s-t; 23,n

Arbor 3-8 m. alta ramulis juvenilibus glabris. Folia tenuiter
coriacea rigidiuscula glabra; lamina obovato-elliptica apice rotundata
vel obtusissima breviterque emarginata et mucronulata, basi subite
cuneata in petiolum alatum 5-7 mm. longum attenuata, costa promi-
nenti nervis secundariis plus minusve prominulis, margine integra
vel levissime remoteque crenata sparsis glandulis vel eglandulosa; 3-
5.5 em. longa 2-3.6 cm. lata. Petala glabra oblongo-attenuate circa
6X 1.5mm. Ovarium glabrum, parcis pilis apice exceptum. Drupa
oblonga apice subacuta 11-14 mm. longa 4-5 mm. crassa in sicco,
exocarpio tenui; endocarpio oblongo utrinque obtuso 11-13.5 mm.
longo 3.8-4.8 mm. crasso, longitudinaliter 10-striato apice minute
5-foraminato.

Type in the U.S. National Herbarium, No. 2270239, collected on
Serra Tepequen, in Rio Branco, Brazil, dominant on rocky slopes
at alt. 1,000-1,200 m., November 29, 1954, by Bassett and Celia
Maguire (No. 40105). Isotype in the Herbarium of the New York
Botanical Garden.

Small much-branched or bushy tree with glabrous branchlets.
Leaves subcoriaceous rather stiff, glabrous, obovate-clliptic, rounded
or very obtuse and emarginate at apex, cuneate at base and attenuate
into petiole 5-7 mm. long; midrib prominent, minor veins more or
less prominulous; 3-5.5 cm. long, 2-3.6 cm. broad. Petals glabrous,
oblong-attenuate toward the apex. Drupe oblong, subacute, 11-14
mm. long 4-5 mm. thick when dry, endocarp oblong, obtuse at both
ends, 11-13.5 mm. long, 3.8-4.8 mm. thick, with 10 thin longitudinal

CUATRECASAS—-HUMIRIACEAE 115

striations and 5 minute foveae at apex. Very closely related to variety
coriacea, it primarily differs in its cylindrical, oblong, narrow
endocarp and its thinner and longer attenuate leaves.

VENEZUELA: Botfvar: Cerro Altamira, 10 km. east of Ciudad Piar, alt.
425-650 m., frequent, bushy tree 3-8 m., flowers greenish white, fruit green,
19-X-1953, Maguire, Wurdack, & Bunting 35882 (NY, US).

BRAZIL: Rio Branco: Serra Tepequena, alt. 1,000-1,200 m., dominant on
rocky slopes, rounded tree 3-8 mm. tall, flowers white, 29-XI-1954, Maguire &
Maguire 40105 (US, holotype; NY).
lk. Humiria balsamifera var. imbaimadaiensis Cuatr., var. nov.

FIGURE 22,m

Frutex ad 30 em. alta, ramulis glabris. Folia crasse rigideque
coriacea sessilia glabra; lamina elliptica apice rotundata sed anguste
emarginata in sinu mucronulata, basi rotundata vel subtruncata plus
minusve auriculata amplectens, margine integra subtus paucis glandu-
lis vel eglandulosa, 2.5-5 em. longa. 1.7-3.2 cm. lata, costa subtus
eminenti basim versus crassa, nervis secundariis venulisque praeser-
tim subtus prominulis. Inflorescentiae folia subaequantes pilis minu-
tissimis patulis sparsis munitae. Petala oblonga obtusiuscula circa
4 mm. longa 1.2 mm. lata dorso sparsis pilis patulis reliqua glabra.
Ovarium glabrum apice paulo piloso excepto. Drupa ellipsoidea,
specimine unico vidit 7 X 5 mm.

Type in the U.S. National Herbarium, No. 2270222, collected at
the Imbaimadai savannas, alt. 550 m., along the Upper Mazaruni
River in British Guiana, October 21, 1951, by Bassett Maguire and
D. B. Fanshawe (No. 32158). Isotype in the Herbarium of the New
York Botanical Garden.

Low shrub with glabrous terminal branchlets. Leaves thick, rigid-
coriaceous, sessile, glabrous, elliptic, rounded and emarginate (sinus
mucronulate) at apex, rounded or subtruncate and more or less auricu-
late, amplectant at base, margin entire; 2.5-5 cm. long, 1.7-3.2 cm.
broad, midrib prominent beneath, secondary nerves and veins pro-
minulous, chiefly beneath. Petals oblong, subobtuse, about 4 mm.
long, with very few hairs outside. Drupe ellipsoid about 7 * 5 mm.
It differs from related variety coriacea in its elliptic, sessile, basally
rounded, subauriculate and amplectant leaves; it is, furthermore, a
low, depressed shrub and has shorter petals with few hairs outside.

BRITISH GUIANA: Upper Mazaruni River, Imbaimadai savannas, alt.
550 m., frequent, depressed shrub to 3 dm. tall and 1 m. across; leaves stiff, leathery ;
flowers greenish white, 21-X-1951, Maguire & Fanshawe 32158 (US, holotype;
NY, isotype).

ll. Humiria balsamifera var, iluana Cuatr., var. nov.

Arbuscula usque ad 8 m. alta ramulis terminalibus glabris. Folia
crassiuscule rigideque coriacea glabra; lamina obovato-elliptica apice

116 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

paulo attenuata obtusa emarginulata minuteque mucronulata, basi
cuneata in latum petiolum alatum crassum 5-7 mm. longum attenuata,
margine integra raris minutis glandulis; costa conspicua subtus inferne
crassa, nervis secundariis crebris venulisque utrinque prominulis;
3.5-7 em. longa 1.8-4 cm. lata. Petala lineari-oblonga subacuta
5.5-6 mm. longa, circa 1.2 mm. lata extus hirtula, pilis sparsis patulis
praedita. Ovarium apice parcis pilis exceptis glabrum. Drupa
oblongo-ellipsoidea basi rotundata, apice obtusiuscula 10-12 mm.
longa 6—-6.5 mm. crassa, exocarpio carnoso in sicco tenui; endocarpio
striatulo.

Type in the U.S. National Herbarlum, No. 2270225, collected on
Mount [lu-tepui, alt. 1,400 m., at Gran Sabana, in the State of Bolivar,
Venezuela, March 18, 1952, by Bassett Maguire (No. 33388). Isotype
in the Herbarium of the New York Botanical Garden. Paratype in
the Herbarium of the New York Botanical Garden, collected between
Uarapata and Enemasio at Gran Sabana, Bolivar, Venezuela, Febru-
ary 8, 1952, by Bassett Maguire (No. 33242).

Small bushy tree or shrub with glabrous terminal branchlets.
Leaves thick, stiffly coriaceous, glabrous, obovate-elliptic, slightly
attenuate and obtuse at apex, more or less emarginate and mucronu-
late, cuneate at base and attenuate into 5-7 mm. long petiole; midrib
prominent beneath, secondary nerves and veins prominulous on
both sides; 3.5-7 cm. long, 1.8-4 cm. broad. The petals linear-
oblong, subacute, sparsely hirtellous outside. Drupe oblong-ellipsoid,
rounded at base, rather obtuse at apex, 10-12 mm. long, 6-6.5 mm.
thick. Variety iluana essentially differs from variety coriacea in its
hirtellous petals.

VENEZUELA: Botfvar: Gran Sabana, Ilu-tepuf, between Uarapata and
Enemasio, common in savannas, alt. 1,000 m., bushy tree or shrub 1-6 m. tall,
8-II-1952, Maguire 33242 (NY, paratype); alt. 1,400 m. near Camp 1, occasional

in low open woodland, rounded tree 8 m., 13—-III-1952, Maguire 33388 (US,
holotype; NY, isotype).

Im. Humiria balsamifera var. pilosa (Steyermark) Cuatr., comb. nov.
Humiria pilosa Steyermark, Fieldiana Bot. 28:270. 1952.
FicureE 24, f

Type: Steyermark 60289, Venezuela, Bolivar, Ptari-tepuf.

Leaves coriaceous, oblong-elliptic, attenuate very little at both
ends, subrounded or obtuse at apex, commonly emarginate and
mucronulate in depression, narrowed to short, winged petiole (2-3
mm. long), entire or slightly crenulate at margin with few minute
glands or eglandular; 3-5 cm. long, 1.5—-2.5 em. broad; above glabrous,
lustrous, smooth with conspicuous midrib; beneath with prominent,
hirtellous midrib, elsewhere puberulous or glabrous, secondary nerves
and veins prominulous. Branchlets of inflorescence and_ pedicels

CUATRECASAS—HUMIRIACEAE 117

hirtellous. Sepals and petals hirtellous outside. Ovary glabrous,
Young branchlets hirtellous-pubescent or hirtellous-puberulous.

Primarily the hirtellous character of the branchlets, pedicels,
calyx, petals, and leaves beneath distinguishes this variety. Steyer-
mark 59621, which undoubtedly belongs to the same taxon, has a
scarcer indument on leaves and branchlets.

VENEZUELA: Botfvar: Ptari-tepui, scrubby forest on rocky portion of
plateau on southeast facing slopes, alt. 1,600 m.; shrub 10 ft. tall; leaves charta-
ceous to subcoriaceous, dark green above, dull paler green below; fruit edible,
oblong, dull purple-red or blackish red, 1-XI-1944, Steyermark 59621 (F, NY).
Vincinity of Misia Kathy Camp on mesa between Ptari-tepuf, alt. 1,615 m.; tree
40 ft; leaves coriaceous, deep green above, paler green below; rich woods, Steyer-
mark 60289 (holotype F, isotype NY).

In. Humiria balsamifera var. minarum Cuatr., var. nov.

Arbuscula ramulis terminalibus glabris. Folia tenuiter coriacea
rigidula glabra. Lamina oblongo-elliptica utrinque attenuata apice
angustata subacuta mucronulata interdum minute emarginata, basi
angustata in brevem petiolum 1-2 mm. longum, margine minute
crenulata et subtus glandulosa; supra tantum costa conspicua; subtus
costa prominenti nervis secundariis et venulis laxe reticulatis pro-
minulis; 3-5.5 em. longa 1.5-2.4 cm. lata. Inflorescentiae folia
subaequales vel breviores, ramulis parcissime pilosulis. Pedicelli
(circa 1.5 mm.) et calyx glabri. Petala glabra oblonga apicem versus
angustata subacuta 5.5-6 mm. longa basi 1.6 mm. lata. Ovarium
apice piloso excepto glabrum.

Type in the U.S. National Herbarium, No. 1592630, collected on
the slope of Serra do Rio Grande, at 1,280 m., near Diamantina,
State of Minas Geraes, Brazil, May 12, 1931, by Ynes Mexia (No.
5815).

Shrub with glabrous terminal branchlets. Leaves thin-coriaceous,
rigidulous, glabrous, oblong-elliptic, attenuate at both ends, narrowed,
subacute and mucronulate (sometimes emarginate) at apex; petiole
1-2 mm. long; margin minutely crenate, glandular-punctate beneath;
midrib prominent beneath, secondary and lax-reticulate veins pro-
minulous; 3.5-5 cm. long, 1.5-2.4 em. broad. Pedicels and calyx
glabrous. Petals glabrous, oblong, subacute at apex.

This variety is very closely related to varieties coriacea, parvifolia,
and pilosa; it is distinguished by its elliptic, subcoriaceous leaves,
which are finely crenulate and glandular beneath and attenuate at
apex; furthermore, it is completely glabrous except for the very
sparse minute hairs at the inflorescence.

BRAZIL: Minas Gerags: Diamantina, slope of Serra do Rio Grande, alt.
1,280 m., among rocks near seepage, shrub 1.5 m., slightly fragrant, greenish
white flowers, Y. Mexia 5815 (US, holotype; A, BM, NY, 5, GH, U, isotypes).

513359—61——7

118 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Moist plains near Cidade Diamantina, shrub about 4 ft. high, VITI-1840, Gardner
4452 bis (BM). No locality, Riedel s.n. (M, S part).

2. Humiria fruticosa Cuatr., sp. nov. Ficure 24,g-h

Frutex ramosus ad 1 m. alta ramulis griseis vel cinereis minute
hirtello-pubescentibus.

Folia brevia coriacea rigida sessilia glauca. Lamina subelliptico-
oblonga vel oblonga, basi subcordata vel rotundata amplectens raro
paulo attenuata obtusaque, apice leviter attenuata obtusa mucronu-
lata, margine integerrima subtus inferne plerumque 3 minutis glandulis
utroque latere reliqua eglandulosa, 1.5-4 cm. longa, 0.5-1.5 cm. lata;
supra pallido-viridis laevis costa pallida tantum notata minutissimis
pilis acutis patulis copiosis pracdita; subtus densiuscule patulo-
pilosula costa elevata nervis secundariis circa 10 utroque latere paulo
ascendentibus paulo prominulis prope marginem curvato-anastomosatis
venulis laxe reticulatis leviter vel haud prominulis conspicuisque.

Inflorescentiae axillares foliis breviores plerumque 3-4 florae,
pedunculo 0.5-1.4 em. longo erecto striolato paulo compresso minute
patulo-piloso, ramulis 1-3 mm. longis rigidis angulatis minute pilosis.
Bracteae ovato-triangulares acutae vel ovato-lanceolatae, 0.6—1.5
mm. longae minute patulo-pubescentes. Pedicelli 1-2 mm. longi
sparse puberuli sursum incrassati. Sepala triangulari-ovata subacuta
crassa 1-1.2 mm. longa extus sparse pilosula margine dense ciliata,
Petala aestivatione quincuncialia, linearia ad apicem attenuata apice
subacuta brevissime mucronulata, crassiuscula sursum dorso minutis-
simis pilis sparsis munita reliqua glabra, circa 6 mm. longa 1.3-1.5
mm. lata. Stamina 20 filamentis crassiusculis papillosis inaequilongis,
3-4.5 mm. longis, tertio inferiore in tubum coalitis. Antherae 1.2-
14 mm. longae connectivo crasso lanceolato acutiusculo glabro
0.9-1 mm. longo, thecis ellipsoideis basalis barbatis circa 0.4 mm.
longis. Discus intrastaminalis crassiusculus dentatus circa 0.8 mm.
longus. Ovarium ovatum apice paulo piloso ceterum glabrum, 1.5
mm. altum 5-loculare loculis biovulatis, ovulis ellipsoideis in loculo
superpositis. Stylus circa 3 mm. longus robustus erectus dense
hirtus. Stigma 5-capitato-lobatum.

Type in the U.S. National Herbarium, No. 2270237, collected at
the base of Cerro Yapacana, alt. 125 m., savannas, in the State of
Amazonas, Venezucla, November 20, 1953, by Bassett Maguire,
John Wurdack, and George Bunting (No. 36580). Isotype in the
Herbarium of New York Botanical Garden.

Sprawling shrub up to 1 m. high with spreading branches and
minutely hirtellous-pubescent branchlets. Leaves small, coriaceous,
rigid, sessile. Blade subelliptic-oblong or oblong, subcordate or
rounded at base, amplectant, slightly attenuate, obtuse and mucronu-
late at apex, margin entire and mostly with three small glands near

CUATRECASAS—HUMIRIACEAE 119

base on lower side; 1.5-4 cm. long, 0.5-1.5 em. broad; nearly smooth
above with abundant minute, pointed, spreading trichomes, pale
midrib conspicuous, other nerves obsolete; densely covered beneath
with minute, patulous trichomes, midrib prominent, 9-10 pairs of
secondary nerves slightly prominulous, somewhat ascending, curvate-
anastomosate near margin, veins lax-reticulate, slightly or not at all
conspicuous.

Inflorescence axillary, shorter than leaves, usually bearing 3-4
flowers, peduncle 0.5-1.4 cm. long, compressed, striate, minutely
spreading-pubescent, branchlets 1-3 mm. long, angulate, minutely
pilose. Bracts ovate-triangular or ovate-lanceolate, 0.6-1.5 mm.
long, minutely spreading-pilose. Pedicels 1-2 mm. long, sparsely
puberulous, thickened toward top. Sepals triangular-ovate, subacute,
thick, 1-1.2 mm. long, sparsely pilose outside, margin densely ciliate.
Petals linear, white, attenuate, subacute and mucronulate, with scarce,
minute hairs outside near apex, about 6 mm. long, 1.3-1.5 mm. broad.
Stamens 20, filaments rather thick, papillose, 3-4.5 mm. long, united
in tube on lower third. Anthers 1.2-1.4 mm. long with lanceolate,
subacute, 0.9-1 mm. long connective; thecae basal, ellipsoid, hairy,
about 0.4 mm. long. Disk about 0.8 mm. high, dentate. Ovary
ovoid, only scarcely pilose at apex, 5-locular, cells biovulate, ovules
ellipsoid, superimposed inside cavities. Style about 3 mm. long, erect,
densely hirtellous. Stigmas 5, capitate-lobate.

H. fruticosa is very distinct from all other species by its shrubby
habit, its longer subacute sepals, its small, narrow, sessile leaves, and
its fine pubescence of minute, patulous, pointed trichomes, which more
or less densely cover the branchlets, inflorescences, and leaves.

This species is highly endemic and well defined, and is limited to the
savannas around the Guiana hill called Cerro Yapacana.

VENEZUELA: Amazonas: Orinoco, Cerro Yapacana, Savanna No. 3, alt.
125 m., northwest base of mountain, frequent, weak shrub to 1 m., 31-XII-1950,
Maguire, Cowan, & Wurdack 30483 (NY, US); shrub to 0.5 m., flowers white, fruit
red, occasional in savanna, 1-I-1951, Maguire, Cowan, & Wurdack 30561 (NY,

US); low sprawling shrub, leaves glaucous, fruit orange, 20-XJ-1953, Maguire,
Wurdack, & Bunting 36580 (holotype, US; isotype, NY).

3. Humiria wurdackii Cuatr., sp. nov. PLATE 10

Frutex 1-3 m. altus ramulis tenuibus tortuosis griseis vel ultimis
badiis, glaberrimis.

Folia coriacea glabra linearia, 2.5-10 cm. longa 3-8 cm. lata, apice
angustata obtusa vel emarginata et mucronulata, mucrone 0.3-0.4
mm. longo calloso deciduo, basim versus ad modum petiolum subala-
tum 4-8 mm. longum angustata, margine integerrima eglandulosa vel
prope margine parce subglanduloso-punctata, supra olivacea subnitida
laevia tantum costa plana conspicua subtus pallide viridia costa

120 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

prominula nervulis minoribus laxe reticulatis paulo conspicuis vel
obsoletis; saepe lamina secundum costam plicata.

Inflorescentiae axillares pauciflorae foliis breviores 1-3 ramis ad 15
mm. longis teneris glaberrimis. Bracteae crassiusculae ovatae obtusac.
amplectentae glabrae 0.5-0.7 mm. longae. Pedicelli 1-1.5 mm. longi
crassi sursum ampliati. Sepala late rotundata circa 1.5 mm. longa
inferne coalita crassa glabra margine minutissime ciliata excepto,
eglandulosa vel raro glandula dorsale munita. Petala alba aestiva-
tione cochlearia crassiuscula glabra linearia apice attenuata minuteque
glanduloso-mucronulata, 5-7 mm. longa 1-1.3 mm. lata. Stamina
20 filamentis crassiusculis papillosis inaequilongis petala non attingen-
tibus dimidia parte in tubum coalitis. Antherae circa 0.8 mm. longae
connectivo crasso lanceolato acutiusculo thecis ellipsoideis basi
breviter barbatis circa 0.4 mm. longis. Discus intrastaminalis an-
nularis crassiusculus glaber 10-lobatus lobis emarginatis circa 1 mm.
altus. Ovarium ovatum sursum pilosulum, circa 2 mm. altum,
5-loculare loculis biovulatis ovulis oblongo-elipsoideis in loculo super-
positis. Stylus erectus robustus hirtellus circa 3 mm. longus. Stigma
5-capitato-lobatum. Drupa ellipsoidea.

Type in the U.S. National Herbarium, No. 2282982, from Venezuela,
Amazonas, Rio Atabapo, 20 km. above San Fernando de Atabapo,
abundant at margin of Sabana Cumare on right bank of Cafio Cumare,
elevation 125 m., collected June 3, 1959, by J. J. Wurdack and LS.
Adderley, No. 42760.

Shrub about 3 m. high with spreading, thin, glabrous branchlets.
Leaves coriaceous, small, linear, entire, usually folded at length, ob-
tusely narrowed and minutely mucronate at apex, narrowed into a
subpetiole at base, 2.5-10 cm. long, 3-8 mm. broad, the midrib con-
spicuous above, prominent beneath, the veins inconspicuous above,
loosely reticulate and slightly prominulous beneath.

Inflorescences short, axillary, usually with 5-10 flowers and 1-3
thin, glabrous branchlets up to 15 mm. long. Bracts ovate, obtuse,
glabrous, 0.5-0.7 mm. long. Pedicels 1-1.5 mm. long, thickening
towers the apex. Sepals broad, rounded, united at the lower half,
1.5 mm. long, glabrous except for the minutely ciliate margin, usually
without glands, sometimes with a rounded gland on the back. Petals
white, thick, glabrous, linear, attenuate at apex, ending with a minute
callose gland, 5-7 mm. long, 1-1.8 mm. wide. Twenty stamens, little,
shorter than the petals, united in the lower half forming a tube, the
filaments rather thick, papillose, unequal in length. Anthers with a
thick, lanceolate, acute connective, the thecae basal, ellipsoid, slightly
hairy. Intrastaminal disk annular, about 10-lobate with emarginate
lobes, 1 mm. high. Ovary hairy at the apex, 5-locular, each cell with

CUATRECASAS—HUMIRIACEAE 121

two superimposed ovules. Style erect, rigid, hairy, about 3 mm.
long. Stigma capitate-lobate. Drupe ellipsoid.

HZ. wurdackii is characterized by its narrow, linear leaves, a unique
feature in the family. This species has some relationship to H. bal-
samifera var. laurina. The species is only known from a single
savanna in the Venezuelan Llanos, where it is abundant.

VENEZUELA, Amazonas: Rio Atabapo, 20 km. above San Fernando de
Atabapo, elev. 125 m., in Sabana Cumare, on right bank of Cafio Cumare.

Shrub 1-3 m., flowers white, locally abundant at margins of the Sabana, J. J.
Wurdack & L. S. Adderley 42760.

4. Humiria crassifolia Mart. ex Urb. in Mart. Fl. Bras. 12(2):441. 1877.
FicureEs 21,u; 23,0,q—-r; 25,a-b
Humirium crassifolium Mart. Nov. Gen. & Sp. Pl. 2:148-144, pl. 198. 1826.
Myriodendrum subvaginale Mart. ex Urb.in Mart. Fl. Bras, 12(2):441. 1877.

Type: Martius, Colombia, Caqueta, Sierra de Araracuara.

Small or medium-size tree with almost smooth, lustrous, glabrous,
rather thick and densely leafy terminal branchlets. Leaves thick-
coriaceous, rigid, glabrous. Petiole 1-2.5 cm. long, stout, winged,
amplectant at base. Blade elliptic-ovate, little oblong, somewhat
attenuate towards the base, slightly narrowed and obtuse or sub-
rotundate at apex, entire and with some very distant gland-spots at
margin; 7-16 cm. long, 4-9.5 cm. broad; above greenish, nitid, with

ZIMMERMANN ’ C
&

Ficure 25.—a-b, Humiria crassifolia: a, X¥% (Maguire &F Fenshawe 23233); b, bud, X3%
(Schultes &% Cabrera 15054); c, Humiriastrum obovatum, X14 (Gleason 729); d-f,
Humiriastrum villosum: d, X% (Froes 22644); e, bud, X34; f, petal. ¥ 314.

122 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

flat and broad midrib, lateral nerves obscurely prominulous, smaller
obsolete; beneath with very thick midrib, 11-12 pairs of spreading
secondary nerves slightly prominulous, near margin arcuate-anasto-
mosate, minor veins parallel and reticulate, little conspicuous.

Inflorescences axillary-subterminal, cymose-paniculate, dichoto-
mous, corymbiform, shorter than leaves; peduncle and branchlets
compressed, glabrous. Bracts amplectant, persistent, ovate or
triangular, acute, 2-1 mm. long. Fertile ultimate branchlets very
short (0.5 mm. long), with bracteole 0.5 mm. long. Pedicels glabrous,
thick, thicker towards apex, about 1 mm. long. Sepals rotundate,
united at base, about 1 mm. long, minutely ciliate at margin, glabrous
outside. Petals white, thick, rigid, linear-oblong, narrowed and
subobtuse at apex, 5 mm. long, 1-1.5 mm. broad, puberulous out-
side. Stamens 20, filaments papillose, 4-5 mm. long, concrescent on
lower half, 10 shorter alternating with longer ones. Anthers about
0.8 mm. long, thecae globose and hairy, connective ovate-lanceolate.
Disk annular, scales linear, acute, united at base. Ovary globose,
about 1 mm. high, glabrous, 5-locular, cells biovulate. Style rather
thick, rigid, 3.5 mm. long, hirtellous. Stigmas 5, oblong-ellipscid,
translucent, connivent-stellate. Drupe ellipsoid, 10-12 mm. long,
7-9 mm. broad. Endocarp obovoid, subrounded at apex, subacute
at base, 10-11 mm. long, 6.5-8 mm. broad, 5-foraminate at apex.

H. crassifolia is only known from the Cerro de Araracuara, Caquetd,
Colombia (type locality), the Cerro Isibuquiri in Vaupés, and the
Kaieteur Plateau in British Guiana. Probably it is spread further
along the isolated mountains of the crystalline Guiana shield.

COLOMBIA: Caquer: “In sylva ad radicem montis Araracoara,’ Martius
s.n. (holotype, M; isotypes, M). Vavupfs: Rfo Cananarf, Cerro Isibuquiri, base
de cuarcita, cerca del pico, 10 ft. tall, flowers white, yellow anthers, I-1952,
Schultes & Cabrera 15054 (US).

BRITISH GUIANA: Kaieteur Plateau, from bush island in savanna, occa-
sional; 6 m. tree, 12 cm, diam.; leaves rigid coriaceous; young fruit oval, green;

seed bony pale brown; 5-V—1944, Maguire & Fanshawe 23233 (NY, US, U,
VEN).

6. Humiriastrum

Humiriastrum (Urb) Cuatr., gen. nov.

Saccoglottis subgen. Humiriastrum Urb. in Mart. Fl. Bras. 12(2):443. 1877.

Saccoglottis Sect. Humiriastrum (Urb.) Reiche in Engl. & Prantl, Pfanzenfam.
3(4):37. 1890.

Sacoglottis Sect. Humiriastrum (Urb.), Winkl. in Engl. & Harms, Pflanzenfam.
19a:128. 19381.

Humirium Benth. in Hook, London Journ. Bot. 2:373. 1843 (in part);
in Hook. Journ. Bot. Kew Gard. Mise. 5:100. 1853.

Sepals 5, suborbicular, imbricate, united at base. Petals 5, free,
thick-membranaceous, linear or oblong, the estivation quincuncial,

CUATRECASAS—HUMIRIACEAE 123

contorted, or cochlear. Stamens 20, in 2 alternating lengths, glab-
rous, filaments connate at base. Anthers ovate-lanceolate or oblong
attached near base, thecae 2, unilocular, ellipsoid or subglobose, basal,
connective thick, more or less lanceolate and acute at apex. Disk a
dentate ring circling ovary or more or less free scales. Ovary 5-locular,
cells uniovulate. Carpels opposite sepals. Ovules anatropous with
ventral raphe, pendant at inner angles of ovary. Style short. Stigma
capitate-lobate. Drupe medium-size or small, ellipsoid or subglobose,
smooth, exocarp carnose, subcoriaceous when dry. Endocarp woody,
usually without resinous cavities, 5 foramina (small holes) around apex
and 5 oblong germinal opercula or valves on upper half. 1-2 semi-
niferous cavities well developed, rarely up to 5. Seeds oblong. Ever-
green trees with coriaceous or subcoriaceous, simple, alternate, peti-
olate leaves, entire or dentate. Stipules small, deciduous or lacking.
Inflorescences axillary or pseudoterminal, paniculate mostly with
trichotomous or dichotomous branching. Bracts persistent or de-
ciduous. (See also figs. 3, 26, and 29.)

Type species: Zumiriastrum cuspidatum (Benth.) Cuatr.

“TWumiriastrum”’ is a derivative name indicating resemblance to
“Humiria.”’

Humiriastrum comprises 12 species spread throughout tropical
South America from the eastern to the western coast going northward
as far as Costa Rica and southward to Rio de Janeiro.

Key to the Species of Humiriastrum

1. Bracts persistent. Petals hispidulous, rarely glabrous. Ovary glabrous,
rarely puberulous.

2. Leaves obovate-spatulate or elliptic-obovate, rounded or truncate at
apex, attenuate at base, sessile or subsessile, margin revolute, pilose
with hirsute midrib beaneath, secondary nerves fine, spreading, promin-
ulous. Branchlets pubescent-hirsute. .... . . 1. H. obovatum

2. Leaves narrowed acuminate or cuspidate at apex, flat, nerves and veins
obsolete or very little conspicuous, firmly coriaceous.

3 Leaf blades more or less villous-hirsute and densely so on midrib, ovate-
acuminate or cuspidate. Terminal branchlets hirsute. Peduncle and
branchlets of inflorescence hirsute, uppermost densely hirtellous.

Sepals hirtellous. . . ........... . . .2. H. villosum
3. Leaf blades glabrous. Terminal branchlets glabrous or rarely puberulous-
hirtellous.

4. Drupe oblong-elliptic, 2.4-3 em. long. Leaf blades ovate-oblong or
elliptic-oblong, narrowed cuspidate. Petals hispidulous. Sepals
shortly puberulous. Ovary sparsely puberulous or glabrous.

3. H. piraparanense

4. Drupe globose, 1.7-2 em. in diameter. Leaf blades ovate or elliptic-
acuminate, abruptly cuspidate, more conspicuously dentate. Petals
hispidulous or glabrous. Sepals glabrous outside. Ovary glabrous.

4. H. cuspidatum

124 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

1. Bracts deciduous.

5. Inflorescence mostly terminal, as long or longer than the leaves, erect,
corymbiform, multifiorous. Leaves glabrous above, sparsely covered
with very minute, thin, appressed, inconspicuous hairs beneath; rather
thick.

6. Young branchlets winged, glabrous. Leaves sessile or subsessile, broadly

ovate or elliptic, rounded or obtuse at base, more or less amplectant.
Ovary glabrous. Petals puberulous. . .. . . . 12. H. procerum
6. Young branchlets subterete or slightly angulate, pubescent or puberulous.
Leaves short-petiolate, subobovate-elliptic or oblong-elliptic, cuncate
at base. Ovary more or less pilose. Petals appressed pubescent or
scarcely puberulous... . .. . . .1L H. diguense

5. Inflorescences mostly axillary, shorter than leaves, usually divaricate-
paniculate. Leaves completely glabrous or spreading-pilose.

7. Terminal branchlets hirsute or puberulous-hirtellous. Peduneles and

branchlets of inflorescences hirsute or hirtellous.

8. Leaf blades softly spreading, pubescent or puberulous beneath,
rigidulous, the midrib densely pubescent above, the secondary
nerves and reticulum = sharply prominulous on both _ sides.
Branchlets very hirsute. Petioles 7-10 mm. long, hirtellous. Pedi-
cels glabrous. Sepals glabrous outside. . . . . 7. H. dentatum

8. Leaf blades glabrous or inconspicuously puberulous, nervation incon-
spicuous above. Sepals minutely pilose outside. Pedicels pilose.

9. Petioles about 1 mm. long; leaf blades inconspicuously puberulous

with minute, thin, subappressed hairs beneath; rigid coriaceous,
secondary nerves slightly conspicuous, reticulum obsolete beneath.
Petals hirtellous . . . . ... . .9. H. subcrenatum

9. Petioles 2-6 mm. long; leaf blades glabrous.

10. Petioles 2-3 mm. long. Leaf blades thin-coriaceous, flexible,
rounded or obtusely cuneate at base, nerves and reticulum
slightly prominulous beneath. Petals puberulous-hirtellous.
Drupe ellipsoid-ovoid, 2-2.5%1.4-1.8 em. . 5. H. excelsum

10. Petioles 3-6 mm. long. Leaf blades rigidulous-coriaceous, acutely
cuneate at base, nerves and reticulum prominent beneath.
Petals glabrous. Drupe globose, 1-5-1.6 cm. in diameter.

10. H. mapiriense
7. Terminal branchlets glabrous. Leaves glabrous.

11. Leaves rounded at apex often abruptly, shortly and obtusely acu-
minate, abruptly and obtusely cuneate at base, subsessile, smooth
above; with thick midrib, numerous thin spreading secondary
nerves and prominulous reticulum beneath. Inflorescences termi-
nal and subterminal, dichotomous, short-hirtellous-puberulous, be-
coming divaricate. Petais sparsely strigose. Drupe black, ovoid,
apiculate, 1.6-1.8X1-1.3 em... .. . 13. H. melanocarpum

11. Leaves attenuate toward apex, acute, acuminate or cuspidate.

12. Petiole 6-11 mm. long. Leaf blades with midrib impressed above
and prominent beneath, nerves and reticulum thin above, sharply
prominent beneath. Peduncle and branchlets of inflorescence
glabrous, rarely puberulous. Pedicels glabrous. Sepals glabrous
outside. Petals glabrous. Ovary glabrous. Drupe ellipsoid-
globose 19%2.2 em... . ... . .8 HL glaziovii

12, Petiole 2-4 mm. long. Leaf blades smooth above, midrib promin-
ulous, other nerves thin or inconspicuous beneath. Peduncle

CUATRECASAS—HUMIRIACEAE 125

and branchlets of inflorescence hirtellous. Pedicels puberulous.
Sepals puberulous outside. Petals puberulous. Ovary hirtellous.
Drupe ellipsoid-ovoid, 2-2.21.2-1.5cem. 6. H. colombianum
1. Humiriastrum obovatum (Benth.) Cuatr., comb. nov. Figure 25,c

Humirium obovatum Benth. in Hook. London Journ. Bot. 2:373. 1843.
Sacoglottis obovaia (Benth.) Urb. in Mart. Fl. Bras. 12(2):443. 1877.

Type: Schomburgk 166, British Guiana.

Tree about 16 m. high with slender, grayish, pubescent-hirsute
terminal branchlets. Leaf blades coriaceous, rigid, obovate or
elliptic-obovate, more or less elongate, rounded or truncate-emarginate
at apex or sometimes obtuse, attenuate toward base, cuneate-sessile
or with very short winged petiole; margin apparently entire, strongly
revolute; 2.5-9 cm. long, 1.4-4 em. broad; green above, nitid, pubes-
cent on midrib and margin, elsewhere glabrous, secondary nerves
and veins scarcely conspicuous; hirtellous beneath with copious,
thin, spreading hairs, midrib thick, hirsute, secondary nerves, 6-8
pairs, subprominent, patulous, near margin arcuate-anastomosate,
veins prominent and lax-reticulate.

Inflorescences corymbose-paniculate, trichotomous or dichotomous,
axillary in upper leaves, shorter than leaves, peduncle and branches
pubescent-hirtellous. Bracts persistent, amplectant, triangular or
ovate, acute, hirsute, 1-0.5 mm. long. Pedicels thick, hirsute, about
0.5 mm. long, articulate with 0.4-1 mm. long, thick, hirtellous
peduneles. Sepals about 0.7 mm. long, hirtellous, connate at base,
apex rotundate. Petals thick, oblong, subacute or subobtuse at
apex, hirtellous, 2 mm. long, 0.8 mm. broad. Stamens about 20,
filaments 0.8-1.2 mm. long, lower part connate. Anthers oblong,
glabrous, about 0.6 mm. long, thecae subglobose, small, connective
thick, sublanceolate. Disk annular, 0.4 mm. high, 20-denticulate,
girdling ovary. Ovary subglobose, glabrous, 0.7-0.8 mm., 5-locular,
cells uniovulate; ovules oblong, 0.4 mm. long. Style thick, about
0.6 mm. long. Stigma capitate, 5-lobate.

H. obovatum is a very well-defined species with a distribution
limited to the British and Venezuelan Guianas.

BRITISH GUIANA: Upper Kamuni, 15—-XII-1908, “hurihi,’’ collector ?
(NY). Upper Kamuni River, “hurihi,” a low spreading savanna tree, bark
used as an antiseptic, XJI-1908, Forest Department (Anderson) 154 (KK). Yam-
pari Creek, 20 miles southwest of Georgetown, swampy land near stream on
alluvial clay, flowers greenish, ripe fruit purplish black and eaten by parrots,
medium-size tree not buttressed with smooth bark and reddish blaze, 29—V—1929,
“hurihi,”’ (Arawak), Forest Department 931 (KK). Sine loco, 1841, Schomburgk
166 (P, isotype). Sine loco, 1841, Schomburgk 135 (Richard) (M). Sine loco,
2nd expedition Schomburgk 825 (P). “Guiana Anglica,’’ Schomburgk 584 (NY).
Sylva ad lacunam Tapacuma, VIII-1848, Schomburgk 1359 (GH, 8, US, isopara-
types). Demerara River, 1887, “honronhi,” Jenman s.n. (NY, U). Butukari,
dense upland forest, 20-21—-VII-1921, Gleason 729 (GH, NY, US).

126 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

VENEZUELA: Botfvar: Regién de los rfos Ieaburti, Hacha y cordillera sin
nombre a 280° de las cabeceras del Rio Hacha, 450-850 m. alt.; selva pluvial o
sabana natural; drbol 30 m., madera roja sin litex ni resina, hoja verde clara,
flores en racimos abiertos, estambres numerosos erectos, 7-I-1956, Bernardi 2814
(NY).

2. Humiriastrum villesum (Frées) Cuatr., comb. nov.
Fiagure 25,d-f; Puats 11
Sacoglottis villosa Frées, Bol. Tec. Inst. Agr. do Norte 20:53. 1950.

Type: Frdées 22644, Brazil, Amazonas, Rio Padauiri.

Medium-size tree with hirsute terminal branches. Leaves coria-
ceous, short-petiolate to subsessile. Blade ovate-acuminate or ovate-
lanceolate, rotundate and abruptly and shortly cuneate at base,
narrowed and acuminate at apex, margin slightly crenate or sub-
entire; 3-6 cm. long, 2-3.8 em. broad; above only prominent and
hirsute midrib conspicuous, elsewhere almost smooth and glabrous;
midrib prominent and hirsute beneath, remaining more or less densely
and softly villous-hirtellous, minor nerves and veins immersed,
obsolete.

Inflorescences axillary and subterminal, half the length of leaves,
paniculate-cymose, lower branches trichotomous or dichotomous,
peduncle 1-1.5 cm. long, straight, densely and spreadingly hirsute,
branchlets densely hirtellous. Bracts persistent, ovate-oblong, hir-
tellous, 1.5-0.5 mm. long. Pedicels thick, 0.4-0.5 mm. long, hirtellous.
Sepals 0.6 mm. long, ovate-orbicular, short-connate at base, hirtellous.
Petals elliptic-oblong, subobtuse, hispidulous, about 3 mm. long and
1-1.3 mm. broad, estivation quincuncial. Stamens 20, the 2 sizes
alternating, filaments complanate, about 1.2 and 1.6 mm. long, gla-
brous lower parts connate in a tube. Anthers 0.6-0.7 mm. long,
thecae ellipsoid, connective carnose, ovate-acuminate. Disk formed
by oblong, bitridentate, glabrous, scarcely adherent scales. Ovary
globose, glabrous, 5-locular, cells uniovulate. Style 0.5 mm. long.
Stigma shortly 5-lobate.

H, villosum is characterized by its general hirsute indument. The
type specimens are densely hirsute, whereas Ducke’s and Humbert’s
collections have a loose indument on leaves and branches; peduncles
and calyx, however, are always abundantly hairy. This species is
found in the upper Amazon Basin in the Rio Negro and Rio Vaupés
regions, and it has also been collected far away in Obidos, in the
State of Para.

COLOMBIA: Vaupks: Rio Cubiyd, afluente del Vaupés, 350 m. alt., 9-10-XI-
1952, Humbert & Schultes 27363 (US, P).

BRAZIL: Amazonas: Cachoeira do Rio Araca, subafluente do Rio Negro,
terra baixa; arvore de 7 metros, a beira do rio, flor branca, 29-X-1952, Frées &

Addison 29144 (IAN). Rio Negro, Paauiry, Rio Pitima, tree 50 ft., 16 in., greenish
yellow flowers, on low land, high forest, border of river of blackish water, clay soil,

CUATRECASAS—-HUMIRIACEAE 127

21-X-1947, Frées 22644 (holotype IAN, isotype P). Par&: Obidos, campinas de
areia ao sul da Serra do Valho-me Deus, 20-VII-1912, Ducke 12030 (MG).

3. Humiriastrum piraparanense Cuatr., sp. nov. Figure 26,e-g

Arbor parva vel media ramulis ultimis subteretibus glabris nitidis
deinde griseis rugulosis. Folia coriacea breviter petiolata glabra.
Petiolus 2-7 mm. longus crassiusculus complanatus anguste alatus.
Lamina oblongo-ovata vel oblongo-elliptica basi subrotundata subite
breviterque cuneato-angustata vel obtuse cuneata, apice longe

ZIMMERMANN, _.

Ficure 26.—Humiriastrum, 1, fruit: a-d, Humiriastrum procerum (Little 6320): a, fruit;
b, endocarp; c, transection at upper third; d, transection at lower third. e-g, Humiria-
strum piraparanense, X1 (Schultes 2 Cabrera 15922): e, fruit; f, endocarp; g, tran-
section. h-j, Humiriastrum excelsum, X1 (Ducke 15459): h, fruit; 7, endocarp; J,
transection. &-l, Humiriastrum colombianum, X1 (Romero Cast. 4942): k, fruit; /,
endocarp. m, Humiriastrum cuspidatum, X 1 (Ducke 30126) transection. n-o, Humiria-
strum cuspidatum var. glabriflorum, 1 (Ducke 23436): n, fruit; 0, endocarp. p-g,
Humiriastrum mapiriense, X1 (Krukoff 11270): p, endocarp; q, fruit. r-t, Humuiria-
strum melanocarpum, X1 (Cuatrecasas 19909): r, transection; s, endocarp; #, fruit.

128 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

angustato-acuminata cuspidata, margine breviter vel obscure serrata
plana, 7-13 cm. longa, 3-6 cm. lata, utrinque sublaevis subopaca
costa crassa sed inmersa vel subtus prominenti bene notata nervis
secundariis 13-16 utroque latere valde tenuibus saepe inconspicuis
prope marginem arcuato-anastomosatis, venulis minoribus tenuissimis
reticulatis haud vel leviter visibilibus.

Inflorescentiae cymoso-paniculatae axillares foliis breviores ped-
unculo puberulo erecto plus minusve compresso apice_ trifurcato
ramis trichotomis vel sursum dichotomo-ramosis, complanatis hir-
tulis; bracteis ovatis amplectentibus ciliolatis inferioribus 1 mm.
longis sursum 0.5 mm., obtusiusculis vel subacutis, persistentibus.
Flores plerumque ternati ad terminationem ramusculis ultimis
hirtulis brevissimis usque 2 mm. longis. Pedicelli crassi brevissimi
minute pubescentes, 0.2-0.3 mm. longi. Calyx ad 0.7-0.8 mm. altus
sepalis basi coalitis, lobis rotundatis margine ciliolatis reliquis his-
pidulo-puberulis. Petala elliptico-oblonga obtusiuscula crassiuscula
viridula extus pubescentia vel puberula circa 3 mm. longa 1.2 mm.
lata acstivatione saepe contorta. Stamina 20 circa 2 mm. longa
glabra filamentis crassiusculis complanatis minute papillosis, tertio
inferiore coalitis 1.7 et 1.2 mm. longis parte libera directa integra
acuta. Antherac circa 0.7 mm. longae glabrae thecis basalis minutis
globosis vel transverse cllipsoideis connectivo crasso angulato multo
longiori lanceolato. Discus in annulo circa 0.5 mm. alto margine
acute 20-dentato ovarium cingens. Ovarium glabrum obovoideum
circa 0.8 mm. altum apice subite in stylum 0.5 mm. longum attenu-
atum, 5-loculare, loculis uniovulatis. Stigma capitatum 5-lobatum.
Drupa ellipsoidea vel ovato-ellipsoidea 2.4-3 cm. longa, 1.8-2.5 cm.
diamitens, laevis nitida. Exocarpium circa 3 mm. crassum coriaceum
in sicco granuloso-resinosum interiore parte magis fibrosum. Endo-
carpium lignosum eresinosum tuberculato-rugosum oblongo-ellip-
soideum 2.2-2.6 cm. longum 1.4-1.7 em. diamitenti, apice subacuta-
tum, subapicem 5 foramina cum 5 operculis oblongis 7-11 mm.
longis descendentibus alternantia, 5 cavitatis seminiferis monospermis
vel tantum 3-1 evolutis. Semina oblonga circa 1.5 em. longa.

Type in U.S. National Herbarium No. 2279921, collected along the
Rio Piraparand tributary of Rio Apaporis in Comisaria del Vaupés,
Colombia, March 9, 1952, by Richard Evans Schultes & Isidoro
Cabrera (No. 15922), specimen with fruit; paratype in U.S. National
Herbarium No. 2270077, collected in Loma Buc-chia, 250-600 m. alt.
at the Rio Piraparan4, tributary of the Apaporis River, Comisaria del
Vaupés, Colombia, August 28-31, 1952, by Hernando Garcia Barriga
(No. 14287), specimens with flowers.

Small or medium-size tree with terete, grayish, nitid, glabrous
branchlets. Leaves coriaceous, rigid, rather thick, glabrous. Petiole

CUATRECASAS—HUMIRIACEAE 129

2-8 mm. long, rather thick, flattened above, narrowly winged on sides.
Blade ovate-oblong, elliptic-oblong, sometimes ovate, subrotundate
and abruptly short-cuneate at base, narrowed, acuminate, and
cuspidate at apex, margin short-serrate and flat, 7-13 cm. long, 3-6
cm. broad, almost smooth on both sides, dull, midrib conspicuously
broad, flat above, prominent below, secondary nerves 13-16 pairs,
very slender and, as well as veins, immersed and inconspicuous.

Inflorescences cymosc-paniculate, axillary, shorter than leaves,
peduncle puberulous, erect, more or less compressed, trifurcate,
branches trichotomous and above dichotomous, complanate, hirtel-
lous. Bracts persistent, amplectant, ovate, obtuse, or subacute
ciliolate at margin, 1-0.5 mm. long. Pedicels thick, short, 0.2-0.3
mm. long, minutely pubescent. Flowers usually ternate on short
(up to 2mm.) hirtellous peduncles. Sepals 0.7-0.8 mm. long, connate
at base, rounded, hispid-puberulous, margin ciliate. Petals rather
thick, greenish, elliptic-oblong, subobtuse, about 3 mm. long, 1.2 mm.
broad, pubescent or puberulous outside. Stamens 20, about 2 mm.
long, the lower third connate, filaments thick, complanate acute,
entire, longer ones about 1.7 mm. long, shorter ones 1.1-1.2 mm. long.
Anthers about 0.7 mm. long, glabrous, minute thecae globose or ellip-
soid, basal, connective thick, angular, lanceolate. Disk annular,
0.5 mm. high, acutely 20-denticulate, girdling ovary. Ovary about
0.8 mm. high, obovoid, glabrous or sparsely pilose, 5-locular, cells
uniovulate. Style 0.5 mm. long. Stigma capitate 5-lobate. Drupe
ellipsoid or ovate-ellipsoid, 2.4-3 em. long, 1.8-2.5 cm. in diameter,
smooth, lustrous. Exocarp about 3 mm. thick, coriaceous when dry,
resinous-granulose becoming fibrous toward interior. Endocarp
woody without resiniferous cavities, tuberculate-rugose, oblong-
ellipsoid, 2.2-2.6 cm. long, 1.4-1.7 cm. in diameter, subacutish at
apex, 5 foramina below apex alternating with 5 oblong, descending,
7-11 mm. long opercula; rarely 5 seminiferous cavities, usually only
3-1, monospermous, seeds oblong, about 1.5 mm. long.

H. piraparanense is very closely related to H. cuspidatum, from
which it differs in its elongate, usually thicker, less markedly dentate
leaves, its puberulous sepals and especially its larger oblong fruit.
In the type specimen the ovary is puberulous.

COLOMBIA: Vaurks: Rfo Piraparand, tributary of Rfo Apaporis, lower
course, small tree, fruit yellow, 9-III-1952, Schultes & Cabrera 15922 (holotype).
Rio Piraparand, Loma Buc-chia, 250-600 em., alt., drbol 20 m., flores amarillas,
28-31-VIII-1952, Garcia Barriga 14287 (paratype).

BRAZIL: Amazonas: Rio Vaupés, Panure caatinga, arvore pequena, flor
branca, 15—-XI-1947, Pires 1080 (IAN). Rio Icgana, Estiréo Santana, caatinga
tipica a margem do rio; arvore 15 m., 22-III-1952, Frées 27985 (IAN) ; specimen

with very good and typical fruit and leaves. Capoeira de Santana, solo are-
noso; arvore 8 m. a margem do rio, 3-IV-1952, Frées 28407 (IAN). Manaos,

130 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

beira do rio Taruma, terra baixa; arvore de 4 m., flores branco-amareladas,
7-VIII-1949, Frées 24924 (IAN).
4. Humiriastrum cuspidatum (Benth.) Cuatr., nov. comb.
Figures 26,m; 27,a-b
Humirium cuspidatum Benth. in Hook. Journ Bot. Kew Mise. 5: 101.
1853.
Sacoglottis cuspidata (Benth.) Urb. in Mart. Fl. Bras. 12(2):444. 1877.—
Ducke, Arch, Jard. Bot. Rio Janeiro 3:178. 1922; 6:39. 1933.—Arch.
Inst. Biol. Veget. Rio Janeiro 4:25, 29. 1937.
Sacogiottis excelsa var. glabriflora Ducke, ibid. 4:25. 19388.

Type: Spruce 1715 and 1915, Brazil, Amazonas, Barra de Rio
Negro.

Medium-size tree with slightly rugose, lenticellate glabrous (in one
variety hirtellous) terminal branchlets. Leaves rigid, coriaceous,
glabrous. Petiole short, thick, broad, and flattened, 2-6 mm. long.
Blade ovate-elliptic, elliptic or obovate-elliptic, cuneate at base
tapering to the petiole, abruptly narrowed, acuminate or caudate at
apex, serrulate-crenate at margin; 4-11 cm. long, 2-6 cm. broad;
midrib broad and flat above, flat or prominent beneath, surface
smooth on both sides lateral nerves immmersed and inconspicuous.

Inflorescences axillary and subterminal, cymose-paniculate, shorter
than leaves, peduncle robust, striolate more or less compressed,
glabrous, (in one variety puberulous), 1-2.5 cm. long, branches
trichotomous and (the superior) dichotomous, spreading, striolate,
very sparsely hirtellous. Bracts ovate, subobtuse or subacute,
glabrous, about 1 mm. long, persistent. Bracteoles ovate, puberulous,
0.8—0.4 mm. long, persistent. Pedicels very short, thick(0.2-0.4 mm.
long). Sepals orbicular, connate at base, glabrous except for ciliate
margin, about 0.6 mm. long. Petals rather thick, greenish, elliptic-
oblong, subobtuse, 2.5-3 mm. long, 1.2-1.5 mm. broad, pubescent,
puberulous or rarely glabrous. Stamens 20, filaments minutely papil-
lose, lower third connate, ten 2—2.2 mm. long alternating with shorter
about 1.7 mm. long. Anthers glabrous, 0.7-0.8 mm. long, connective
thick, ovate-acuminate, thecae short-ellipsoid, basal. Disk formed
by 0.5 mm. long ovate, deeply bidentate, more or less united scales.
Ovary globose glabrous, 5-locular, cells uniovulate; ovules oblong,
about 0.5 mm. long, pendent, with ventral raphe. Style 0.6 mm.
long. Stigma capitate, 5-lobulate. Drupe globose, about 18-20
mm. diameter, exocarp thin and smooth; endocarp globose, rugose,
about 17 mm. diameter, with 5 foramina at apex and 5 elliptic-
oblong, 7 mm. long, descending opercula.

HH. cuspidatum is a species of the upper central Amazonian Basin
(Rio Negro, Vaupés, Manaos region), spreading to southern Venezuela

CUATRECASAS—HUMIRIACEAE 131

Ficure 27.—Humiriastrum: a, H. cuspidatum, X% (Ducke 30126); b, H. cuspidatum,
bud and petal, <3% (Ducke 23434); c, H. cuspidatum var. glabriflorum, Xs (Ducke
243); d, H. cuspidatum var. glabriflorum, petal and bud, X3% (Ducke 23436); ¢, H.
colombianum, X44 (Lamb 141); f, H. colombianum, petal and bud, X 3% (Lamb 141);
g, H. excelsum, X¥% (Ducke 1614); h, H. excelsum, petal and bud, X 3% (Ducke 1614);
i, H. glaziovii var. glaziovii, X % (Ducke 19166); j, H. glaziovii var. glaziovii, bud and
petal, X 3% (Ducke 19166); k, H. glaziovii var. angustifolium, leaf, X ¥4 (Glaziou 16724);
l, H. dentatum, X¥% (Glaziou 18178); m, H. dentatum, bud and petal, X3% (Glaziou
18178).

and to the State of Paré. It grows mostly in inundated places but,
especially its varieties, can also be found on elevated ground. Accord-
ing to Ducke (p. 25, 1938), it is frequent in central parts of Amazonia
along the sandy river banks periodically flooded, the lakes, swamps,
and still-water rivers that are poor in sediments. The mature drupe
(according to Ducke) is globose, 15-25 cm. in diameter, black pur-
plish with thin, juicy, red mesocarp, which has an astringent taste;
when dry it is black and lustrous.

132 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Key to the Varieties of Humiriastrum cuspidatum

1. Young branchlets glabrous. Peduncle of inflorescence glabrous.
2. Petals hispidulous ..........2.2. 2... 4a. var. cuspidatum
2. Petals glabrous .. . .... . 4b. var. glabriflorum
1. Young branchlets pubescent- hirtellous.. Peduncle of inflorescence minutely
puberulous. Petals glabrous or subglabrous. . . 4c. var. subhirtel!um

4a. Humiriastrum cuspidatum var. cuspidatum.

VENEZUELA: Botfvar: Piedra Marimare, east Bank of Rfo Orinoco opposite
head of Isla I] Gallo, tree 18 m., young fruit green, morichal, edge at east base
of Piedra, 2 km. east of river, alt. 100 m., 20-XTI-1955, Wurdack & Monachino
40881 (US).

BRAZIL: Par: Campina entre as Serras do Dedale da Igacaba, 4-IX-1907,
Ducke 8628 (BM, MG). Baixo Yamunda, Lago das dois bocas, beira do campo
alagado, 18-V—-1911, Ducke 11790 (BM, MG). Rio Jamund4; praia Porto Rico,
Municipio de Faro; arvore, fruto verde, “uchirana,”’ “achua,” 15-XI-1950,
Black & Ledouz 50-10783 (IAN, US). Amazonas: Prope Panure ad Rio Vaupxés,
X-1852-I-1853, Spruce 2424 (K, NY, S, P); 2443 (GH, K, P). Rio Negro
prope Barra, VII-1851, Spruce 1715 (isotypes, US, GH, P, NY); 1915 (K, holotype
“Herbarium Hookerianum’’; M, isotype), photo F.M. 12593. Rio Curicuriary,
afluente Rio Negro ad ripas inundabilis; arbor parva vel mediocris, fluoribus
viridibus, fructus violascenti-nigris, 4-X-1935 (fl.), 26-II-1936 (fr.), Ducke 30126
(US, 8, U); Ducke s.n. (IAN). Rio Curicuriary (middle course), small tree,
I-1948, Schultes & Lépez 9701 (US). Rio Apurahu inferior afluente Rio Negro,
silva inundabili: arbor media, floribus viridibus odoratis, 26-VII-1929, Ducke
23434 (US). Rio Urubu, Sucuriju, terra firme baixa; arvore 7 m., flores esbran-
auigadas, 29-IX-1949, Frées 254388 (IAN). Without locality, Schultes 23131
(IAN). Without locality, Martius s.n. (P).

4b. Humiriastrum cuspidatum var. glabriflorum (Ducke) Cuatr., comb.

noy. FIGURES 26,n—0; 27,c—d
Sacoglottis excelsa var. glabriflora Ducke, Arch. Inst. Biol. Veget. Rio Janeiro
4:25. 1938.

Type: Ducke 23436, Brazil, Amazonas, Manaos.

Ramuli hornotini glabri. Pedunculi inflorescentiae et rami majores
glabri, superiores hirtulo-puberuli. Petala glabra.

This variety differs from cuspidatum in its glabrous petals and
the usually smaller leaves. All other characters, including the globose
fruit, completely agree with the available collections of cuspidatum.

BRAZIL: Amazonas: Manaos, prope Cachoeira do Mindi, silva non inunda-
bilis, solo arenoso humoso; arbor magna floribus viridibus odoratis, “achud,”’
22-VII-1936, Ducke 243 (A, NY, 8, US); 8-VII-29, Ducke 23436 (US, P, 8, U,
isotypes). Manaos, Cachoeira Alta do Tarumd, terra umida, arvore de 8 m.,
flor amarela, 11-VIII-1945, INPA 204 (MG. 21546). Manaos, margem do
Igarapé de Forquilha; arvore de copa densa e baixa, flor esverdeada com o
centro castanho, 22-VIII-1955, INPA 1700 (MG 21542), Sao Paulo de
Olivenga, terra firme alta, arvore, I1V-1945, Frées 20803 (IAN, K, NY).

CUATRECASAS—HUMIRIACEAE 133

4c. Wumiriastrum cuspidatum var. subhirtellum Cuatr., var. nov.

Ramuli hornotini hirtuli-pubescenti vel hirtuli-puberuli. Pedunculi
ramique inflorescentiae minute puberuli. Petala glabra vel subglabra.

Type in the Herbarium of Instituto Agronémico do Norte, Belém
do Parad, Brazil, collected on the margin of Rio Urubt, near Sao
Francisco in the State of Amazonas, Brazil, October 4, 1949, by
R. Lemos Frées (No. 25480).

This variety has an uncertain position. The very young fruit are
more or less oblong and their final form is unknown. The leaf charac-
ters agree very much with those of H. cuspidatum and its variety
glabriflorum.

BRAZIL: Amazonas: Rio Urubt, Sdo Francisco, beira do rio, arvore de 5 m.,

Frées 25480 (IAN, holotype). So Francisco, 24-VITI-1949, Frées 24820 (IAN).
Sao Francisco, tierra firme baixa, arvore 8 m., 3-X-1949, Frées 25463 (IAN).

5. Humiriastrum excelsum (Ducke) Cuatr., comb. nov.
Fiacures 26,h-j; 27,g-h

Sacoglottis excelsa Ducke, Arch. Bot. Rio Janeiro 3:178. 1922; 5:143, pl. 14
fig. 41. 1930; 6:39. 1933.—Arch. Inst. Biol. Veget. Rio Janeiro 4:25, 29.
1937.

Type: Ducke 15459, Brazil, Paré, Belém.

Large tree with pubescent-hirtellous or puberulous branchlets.
Leaves thin-coriaccous, flexible, glabrous. Petiole 2-3 mm. long,
puberulous beneath or glabrous. Blade ovate or ovate-elliptic,
broadly cuneate or subrounded at base, more or less abruptly narrowed,
acuminate or cuspidate at apex, flat and slightly crenulate-serrate at
margin; 2.5-9 cm. long, 1.5-5 cm. broad; above with flat midrib,
slender secondary nerves and veins almost obsolete; beneath midrib
prominent, filiform secondary nerves, 10-12 pairs, prominulous, sub-
patulous, near margin reticulate-anastomosate, reticulum prominulous.

Inflorescence axillary, short, cymose-paniculate, mostly trichot-
omous, upwardly dichotomous, peduncle and branchlets minutely
pubescent-hirtellous. Bracts minute, early deciduous. Pedicels rather
thick, pubescent 0.2-0.4 mm. long, articulate with short and minutely
pubescent peduncles. Sepals 0.6-0.8 mm. long, rotundate, minutely
pubescent, margin ciliolate. Petals linear-oblong, attenuate at apex,
subacute, hirtellous-puberulous, about 2.5 mm. long, 1 mm. broad.
Stamens 20, filaments connate at base, about 1.5 mm. long, more or
less papillose. Anthers about 0.8 mm. long, thecae ellipsoid a third
or a fourth of total length, connective lanceolate, acute. Disk
annular, 0.3-0.4 mm. high, dentate. Ovary ovoid, glabrous, 5-
locular, 5-ovulate. Style 0.6 mm. long. Stigma capitate, 5-lobate.
Drupe ellipsoid-ovoid, 2-2.5 cm. long, 1.4-1.8 cm. broad; exocarp
smooth; endocarp woody with 5 foramina at apex and 5 oblong
opercula, about 6 mm. long around apex.

513359—61——8

134 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Hf. excelsum is a large tree of the uninundatable forests of the
Amazon basin. It is frequent in the Belém region (Para) and is also
reported from French Guiana and eastern Peru. The fruit are more
or less oblong and yellowish with oleagineous mesocarp, and sweet
and fragrant when ripe. The tree, according to Ducke, attains a
height of 45 m. and has a brownish-red bark.

FRENCH GUIANA: Riviére “la Compte” rive droite 4 50 m. en Amont du
Saut-Bief chemin minier de Bief et 4 3 km. 500 de la rive Montagne Papillon;
sur le flane d’une montagne, pente raide; sol argileux; “prefontaine,” 29-I-1957,
Bena 1319 (U).

BRAZIL: Pard: Santa Izabel (Belém-Braganca) grand arbre de la forét,
“achud,” 18-IX-1908, Museu Goeldi 9672 (MG, US). Belém, silva non inun-
dabilis; arbor magna vel maxime floribus viridibus oderatus, ‘“achud-rana,”’
16-IX-1922, Ducke 17780 (NY, P, S, US, U), photo F.M. 12597. Belém,
grand arbre de la forét, “achud,” 20-VIII-1914, Ducke 15459 (isotypes, MG, US).
Belém, Catu, mata da terra firme, arvore grande, flor verde, 31—VIII-1944,
“achud,” Ducke 1614 (A, IAN, MG, NY, US). Beira do rio Mapua, entre
Vila Emilia e Boca do Mapua, varzea; arvore com folhas pequenas, 18-VII~-1950,
Black, Frées, & Ledoux 50-9811 (US, IAN).

PERU: Hu4nuco: Tingo Maria, Lote Dianderas (carretera Hudnuco-Pucallpa)
800-900 m.; suelo arcilloso, pendiente mediana, selva luviosa; Arbol 20-30 m.,
40-60 cm. diam., madera dura, de construccion, corteza rojiza, descamdndose
superficialmente, albura clara, duramen rojizo oscuro; relativamente abundante,
“quinilla,” 18-VII-1946, Burgos 85 (Y). Tingo Maria, a 1-2 km. carretera
Hudnuco-Pucallpa, km. 160, 800 m. alt., suelo arcilloso, profundidad mediana;
selva densa; drbol 30 m., 60-100 em. didmetro, flores amarillentas, olorosas;
madera dura, rojiza, usada en postes, durmientes, columnas, ete. Crecen en
grupos + densos y se destacan por su color canela; la corteza se desprende sola en
arboles viejos; regularmente abundante, “hispi,” 28-VITI-1945, Burgos 37 (Y).

6. Humiriastrum colombianum (Cuatr.) Cuatr., status nov.
Figures 26,k-l; 27,e-f
Sacoglottis excelsa var. colombiana Cuatr., Brittonia 8:196. 1956.

Type: Lamb 141, Columbia, Santander, Cimitarra.

Medium-size or large tree, trunk with reddish brown, smooth bark
and reddish, very hard wood. Terminal branchlets slender, brownish,
glabrous. Leaves thin-coriaceous, glabrous. Petiole 2-4 mm. long,
rounded and thickened at base. Blade elliptic or ovate-elliptic,
abruptly cuneate and tapering to petiole at base, abruptly narrowed
and acutely cuspidate at apex, margin erenulate; 4—7 em. long (includ-
ing tail), 1.5-3.5 em. broad; above green with prominulous and con-
spicuous midrib, other nerves immersed, visible or obsolete ; beneath
lightly brownish with prominent and very conspicuous midrib,
secondary nerves, about 9 pairs, extrem ely thin, near margin reticulate-
anastomosate, very slightly prominulous or obsolete, veins obsolete.

Inflorescence cymose-paniculate, axillary and subterminal, shorter
than leaves, lower branching trichotomous, superior dichotomous,
peduncle 1-1.5 cm. long, rigid, striolate, slightly puberulous, branch-

CUATRECASAS—-HUMIRIACEAE 135

lets articulate, more or less spreading hirtellous-puberulous. Bracts
deciduous. Bracteoles ovate, 0.2-0.3 mm. long, puberulous, soon
deciduous. Pedicels very short, thickened, puberulous, 0.2 mm. long;
flowers practically sessile. Sepals 0.4-0.5 mm. long, rounded, mi-
nutely papillose, puberulous, ciliolate. Petals oblong, 2-2.1 mm. long,
about 1 mm. wide, subappressed-puberulous. Stamens 20, filaments
more of less papillose, 10 about 1.1-1.2 mm. long, alternating with
10 shorter ones about 0.7-0.8 mm. long. Anthers oblong-lanceolate,
0.7-0.8 mm. long, thecae short-cllipsoid, basal, connective thick-
lanceolate. Several small, subdentate, 0.2 mm. long 0.1-0.2 mm.
broad, free scales circling ovary and forming disk. Ovary globose,
hispidulous, 5-locular, cells uniovulate. Style about 0.5 mm. long.
Stigma capitate, 5-lobulate. Drupe ellipsoid-ovoid, rounded at base,
abruptly narrowed and subacute or acute at apex, 20-22.5 mm. long,
12-15 mm. broad; exocarp glabrous, smooth thin (0.5 mm.) when dry;
endocarp ovoid-ellipsoid, rounded or subtruncate at base, acutely
acuminate at apex, woody, hard, rugose, barely pentagonal with 5
apical foramina and 5 subapical oblong, 6 mm. long, descending
opercula.

I. colombianum differs from the Amazonian 7. excelsum in its some-
what smaller leaves, which are attenuate and longer cuneate at the
base and much longer cuspidate at the apex (the caudex is very acute,
1-1.5 cm. long, the nervation on the leaves is almost obsolete), in the
glabrous young branchlets, in the free scales of the disk, and in the
hispidulous ovary.

COLOMBIA: Santanper: Regién del Carare (valle del Magdalena), Cimi-
tarra, km. 3 camino del Ermitafio, tree 20 in. diameter, 60 ft. tall, forest canopy,
“aceituno,” 29-VII-1954, Lamb 141 (holotypus, US); 30-VII-1954, Lamb 145
(COL, US); forest tree 25 in. diameter, 50 ft. tall top canopy; wood hard, pink
when fresh; “aceituno”’; 17-VIII-1954, Lamb 170 (COL, US). Barranca Bermeja,
12 leguas al sureste, a 5 km. de la margen derecha del rio Opén, 200 m. alt.; drbol
20 m., madera rojiza, dura, pesada, para polines y pilotes, corteza rojiza un tanto
4spera, aletas basales cortas, ramillas parduscas, ‘“‘aceituno,” 28-IX-1954, Romero
Castafteda 4942 (COL, US). Diez leguas al SE de Barranca Bermeja, 8 km. de la
margen izquierda del rio Opén; drbol 25 m., corteza gris, lisa, pardo rojiza, man-
- chada de gris, albura rosada, corazén rojo, ramillas parduscas, madera dura para
formaletas y entablados, +200 m., “aceituno,’”’ 31-VIII-1954, Romero Castaneda
4785 (COL, US).

H. colombianum is only known from the Magdalena Valley in
Colombia, where it is called “‘aceituno.” Up to now it is also the only
recorded species of the genus from the interior valleys of Colombia.
H. colombianum is a large buttressed tree, the hardwood of which
may be used in construction work; it is frequent in the rain forest on
elevated ground.

136 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

7. Humiriastrum dentatum (Casar.) Cuatr., comb. nov. FiaureE 27,l-m

Humirium dentatum Casar. Nov. Stirp. Bras. Decas IV: 38. 1842.—Benth.
in Hook. Journ. Bot. Kew Mise. 5:102. 1853.

Sacoglottis dentata Urb. in Mart. Fl. Bras. 12(2):444 (in part). 1877.—
Ducke, Arch. Jard. Bot. Rio Janeiro 5:143, pl. 14, fig. 40. 1930.

Type: G. Casaretto, Brazil, ‘from the sandy maritime woods called
restingas in the Province of Rio de Janeiro.”

Tree with pubescent-hirsute terminal branchlets. Leaves coria-
ceous, subrigid. Petiole 7-10 mm. long, narrowly winged, pubescent-
hirtellous, Blade ovate or elliptic-lanceolate, more or less oblong,
narrowed and obtusely short-cuneate at base, attenuate and acuminate
at apex, margin serrate-dentate, slightly revolute, 4-11 cm. long,
1.5-5 cm. broad; above with minutely and densely pubescent midrib,
elsewhere glabrous, secondary nerves filiform, veins forming a con-
spicuous, more or less prominulous reticulum; beneath softly and
sparsely pubescent, prominent striolate midrib, subdensely hirtel-
lous, prominulous secondary nerves about 10 pairs, subascendent,
curvate-anastomosate near margin, veins reticulate and prominulous.

Inflorescence cymose-paniculate, axillary, shorter than leaves,
dichotomous, peduncle and branchlets densely hirtellous. Bracts
deciduous, ovate-oblong, subobtuse, puberulous, 1-0.5 mm. long.
Pedicels thick, glabrous, 0.5-0.8 mm. long, articulate with elabrous
or glabrescent, 0.5-2 mm. long peduncles. Sepals orbicular, connate
at base, glabrous except ciliate margin. Petals rather thick, oblong,
subobtuse, glabrous, about 2.5 mm. long, 1 mm. broad. Stamens 20,
filaments 2-2.5 mm. long, connate at lower third, glabrous. Anthers
glabrous, oblong-lanceolate, connective thick, thecae basal and
oblong. Disk membranaceous, about 0.6 mm. high, short-dentate,
girdling ovary. Ovary ovoid, glabrous, 1 mm. high, 5-locular with
uniovulate cells, ovules elliptic-oblong, about 0.8 mm. long. Style
robust, 0.5-0.6 mm. long. Immature fruit obovate.

Urban included in his “Flora Brasilicnsis” a hirtellous and a glabrous
form in the description of Sacoglottis dentata. The hairy plants
undoubtedly belong to this species and agree with the original descrip-
tion by Casaretto. Urban saw in glabrous plants an unnamed
variety which, I believe, is the species described later by Urban as
Sacoglottis glaziovii. Ihave seen no authentic material of LZ. dentatum,
but it is possible that specimens in Paris without the collector’s name
(from the Drake or Richard Herbaria) belong to the Casaretto
collections.

At present H. dentatum is known only from the Rio de Janciro
region.

BRAZIL: Rio pe JaANneErro, Glaziou 18178 (NY, P), photo F.M. 12595. ‘Herb.
Richard, Humirium dentatum Casar. Bresil F. Nob. 1855” (P). Rio de Janeiro,

CUATRECASAS—-HUMIRIACEAE 137

1851, Anderson s.n. (S). SXo Pavxo: Santos, Sorocaba, I-1875, Mosén 3475
(P, 8). “Herb. E. Drake, Humirium dentatum Casar.” (P)

8. Humiriastrum glaziovii (Urban) Cuatr., comb. nov. FIGURE 27,i-7

Sacoglottis dentata var. Urb. in Mart. Fl. Bras. 12(2):445. 1877.
Sacoglottis glaziovii Urb. Bot. Jahrb. Engler 17:503. 1893.

Type: Glaziou 18964, Brazil, Rio de Janeiro, Nova Friburgo,
Alto Macahé; photo of holotype in Berlin-Dahlem, F.M. 12598.

Small or medium-size tree with glabrous, greenish and _ nitid
young branchlets becoming rugose, brownish, and lenticellate.
Leaves coriaceous, subrigid, glabrous. Petiole 6-11 mm. long.
Blade elliptic-ovate or ovate-lanceolate, obtuse or short-cuneate at
base, abruptly acuminate at apex, the margin scrrate-dentate, flat
or slightly revolute, 3.5-10 em. long, 1.5-5 cm. broad; above nitid
with impressed and thin midrib, lateral nerves and reticulum little
prominent but conspicuous; beneath with prominent midrib, 9-10
pairs of filiform secondary nerves prominulous, subascendent, anas-
tomosate near the margin, minute reticulum prominent.

Inflorescences cymose-paniculate, axillary, much shorter than
leaves. Peduncle and dichotomous (rarely trichotomous) branches
glabrous or smaller branchlets hirtellous-puberulous. Pedicels short
(0.2-0.3 mm.), glabrous, articulate with 0.4-2 mm. long glabrous
peduncle. Sepals about 1 mm. long, rotundate, glabrous except the
minutely ciliate margin. Petals oblong, rather obtuse and thick,
glabrous, 3-3.5 mm. long, 1 mm. broad. Stamens 20, with 2.5-3 mm.
long filaments, glabrous and connate at base. Anthers glabrous,
ovate-lanceolate, thecae oblong and as long as a half to a third of the
acute connective. Disk annular, membranous, girdling ovary,
deeply dentate, 0.6-0.7 mm. high. Ovary ovoid, glabrous, about 8
mm. high, 5-locular with uniovulate cells. Style about 0.7 mm. long.
Stigma capitate. Drupe ellipsoid-globose, smooth, glabrous, about
1922 mm. in diameter.

The concept of H. glaziovii includes the glabrous variety of S.
dentata mentioned by Urban: “Var. ramulis, foliis, inflorescentiis
glaberrimis.” H. glaziovii grows in “restinga’”’ type forests and sec-
ondary woods of the Rio de Janeiro region.

8a. Humiriastrum glaziovii var. glaziovii. Figure 27,i-j

BRAZIL: Rio pz JanErro: Restinga de Mand, 30-XI-1896, avec Mr. Schwacke,
Glaziou 18179 (NY, P, US); Glaziou 18964, photo F.M. 12598. Porto da Estrella,
silvestris siccioribus; arbor parva, floribus viridibus inoodris, in silvula secundaria
sieciora, 17—XI-1925, Ducke 19166 (P, 8, U, US). “Brasilia, Humiria dentata
Cassaretto, Sacoglottis, Ex herbario horti Petropolitani,”’ Riedel s.n. (P, K).

138 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

8b. Humiriastrum glaziovii var. angustifolium Cuatr., var. nov.
Fiaure 27,k

Lamina folii anguste lanceolata basi cuneata crassior coriacea, 2.7—
6.5 cm. longa 1-2 cm. lata.

Type in the U.S. National Herbarium, No. 1123930, collected at
Alto Macahe, State of Rio de Janeiro, Brazil, February 6, 1888, by
M. A. Glaziou (No. 16724). Isotypes at the U.S. National Herbarium,
Nos. 483735 and 287412, in the Kew Herbarium and in the Herbarium
of the Arnold Arboretum.

This variety differs from the typical form only in its narrower,
lanceolate, and firmer leaves. It is only known from the Rio de Janeiro
region and Sdo Paulo in Brazil.

BRAZIL: Sio Pavno: Arvore da Matta da Estacdo Biologica, 5-III-1919,

Hoehne 3021 (NY). Rio pg Janerro: Prov. Nov. Friburgo, Alto Macahé, 6-IT-
1888, Glaziou 16724 (type US, K, A).

9. Humiriastrum subcrenatum (Bentham) Cuatr. PLaTE 12
Humirium subcrenatum Benth in Hook. London Journ. Bot. 2:374, 1843.—
Hook. Journ. Bot. Kew Mise. 5:102. 1853.
Humiria subcrenata Urb. in Mart. Fl. Bras. 12(2):442. 1877.
Sacoglottis subcrenata Urb. Sitz. B. Ges. Naturf. Berl. 5. 1878.

Type: Martin, French Guiana, Cayenne.

Terminal branchlets hirtellous. Leaves rigid coriaceous sub-
sessile; petiole about 1 mm. long hirtellous; blade subelliptic shortly
and obtusely acuminate at apex, cuneate at base, subentire or slightly
crenate except toward the base, 2.6-4.5 em. long, 1.7—2.8 cm. broad;
above lustrous, glabrous or with minute hairs on conspicuous midrib,
veins obsolete; below inconspicuously puberulous by minute, sparse
hairs, midrib prominent, 8-10 pairs of secondary nerves extremely
thin or inconspicuous, minor veins obsolete.

Inflorescences axillary shorter than leaves, cymose-paniculate,
subdichotomous, above branchlets alternate, peduncle 9-12 mm.
long, branchlets angulate and hirtellous; the pedicels 0.2-0.4 mm.
long, hirtellous; bracts and bracteoles soon deciduous; calyx about
0.8 mm. high, quincuncial; sepals free, truncate-rounded, minutely
ciliate at margin and minutely puberulous outside, 0.5 mm. high,
0.8 mm. wide. Petals thickish, linear, acute, hirtellous outside, 2.1—
2.3 mm. long, 0.6mm. broad. Stamens 20, glabrous, filaments united
near base, 2 lengths, 1.1 and 1.4 mm., alternating; anthers thick, acute,
about 0.6 mm. long, thecae minute basal, 0.15 mm. broad. Disk
cupular, 0.7 mm. high, 20-denticulate. Ovary ellipsoid, appressed
from top, minutely hirtellous, 5-loculate, cells uniovulate, ovules
deltoid acute at apex. Style thickish, glabrous, 0.7 mm. long.

H, subcrenatum is only known from the type specimen from French
Guiana. Urban treated this as “species dubia’? under Humiria, but

CUATRECASAS—-HUMIRIACEAE 139

in the following year (1878), after seeing original material sent to him
by Bentham and Oliver, he published the right identification of the
species as Sacoglottis subcrenata in the section Humiriastrum.

FRENCH GUIANA: Cayenne, Martin s.n., Herbarium Hookerianum, holotype
(K).

10. Humiriastrum mapiriense Cuatr., sp. nov.
Fiaures 26,p-q; 28,9; PLate 13

Arbor media ramis terminalibus griseis tenuibus minute puberulo-
hirtulis.

Folia parva coriacea petiolo crassiusculo puberulo 3-6 mm. longo.
Lamina obovata vel rhomboideo-obovata vel oblanceolata basi
valde cuncata in petiolum attenuata apice angustata acutiuscule
acuminata, margine serrato-crenulata, utrinque glabra; 3-4 cm. longa
1.5-2.5 cm. lata supra costa plana visibili ceteris nervis vix obsoletis;
subtus costa elevata nervis secundaris filiformibus 8-9 utroque
latere paulo adscendentibus arcuato-anastomosatis, nervulis venu-
lisque prominulis reticulatis.

Inflorescentiae axillares et subterminales cymoso-paniculatae folia
excedentes dichotomae vel inferne trichotomae pedunculo 1.5—2.5 cm.

Ficure 28.—a-d, Humiriasirum procerum: a, X¥% (Cuatr. 16615); b, section of stem; c,
bud, X 3%; d, petal, X34; ¢, Humiriastrum diguense var. diguense, X Y (Cuatr. 14956);
f, Humiriastrum diguense var. anchicayanum, X14 (Cuatr. 14418); g, Humiriastrum
mapiriense, X % (Buchtien 1518); h, Humiriastrum melanocarpum, X Y% (Cuatr. 19989);
i, Humiriastrum melanocarpum, bud and petal, X 3% (Cuatr. 19989).

140 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

longo tenui striolati minute pubescenti-hirtuli ramulis gracilibus
articulatis minute hispidulis. Bracteae deciduae ovatae circa 1 mm.
longae villosulae. Pedicelli crassiusculi circa 0.3 mm. longi hirtuli.
Sepala ovata circa 0.6 mm. longa margine ciliata dorso hirtula.
Petala (in alabastro 1 mm. longa) elliptico-oblonga glabra aestiva-
tione contorta vel cochlearia. Stamina 20 alternatim inaequalia
filamentis glabris basi coalitis. Antherae oblongae 0.5 mm. longae
(in alabastro) connectivo oblongo-lanceolato. Discus 10 squamis
brevibus liberis instructus. Ovarium glabrum_ 5-loculare loculis
uniovulatis. Stylus brevis. Stigma capitatum 5-lobulatum. Drupa
globosa 15-16 mm. diamitens exocarpio sublaevi tenui (0.5 mm.
crasso in sicco). Endocarpium globosum circa 14 mm. diam. duro-
lignosum resinoso-lacunosum ad apicem 5 minusculis foraminibus,
cum 5 pseudo-operculis oblongis circa 6 mm. longis alternantibus
reliqua superficie rugosa.

Type in the Herbarium of the New York Botanical Garden, col-
lected in Sarampinni near San Carlos, region of Mapiri, Department
of La Paz, Bolivia, alt. 600 m., March 7, 1927, by Otto Buchtien
(No. 1518). Paratype (fruiting specimens) in the U.S. National
Herbarium, No. 1905788, collected at Copacabana, about 10 km.
south of Mapiri, alt. 850-950 m., Province Larecaja, Bolivia, October-—
November 1939 by B. Krukoff (No. 11270).

Medium-size tree with slender, minutely hirtellous-puberulous
terminal branches. Leaves small, coriaceous with 3-6 mm. long,
puberulous petiole. Elade glabrous, obovate or rhomboid-obovate
or oblanceolate, very cuneate at base, narrowed and acutely acuminate
at apex, margin serrate-crenulate; above with flat midrib, other
nerves obsolete; beneath with prominent midrib, filiform secondary
nerves, 8-9 pairs, little ascendent, near margin arcuate-anastomosate,
minor nerves reticulate, prominulous.

Inflorescences axillary and subterminal, cymose-paniculate, longer
than leaves, dichotomous or lower branches trichotomous, peduncle
1.5-2.5 cm. long, striolate and minutely hirtellous, branchlets slender,
articulate, minutely hispidulous. Bracts deciduous, ovate, villous,
about 1mm.long. Pedicels thickened, about 0.3 mm. long, hirtellous.
Sepals ovate, about 0.6 mm. long, ciliate at margin, hirtellous without.
Petals (in bud) 1 mm. long, elliptic-oblong, glabrous, estivation
contorted or cochlear. Stamens 20, alternating in length, filaments
glabrous, connate at base. Anthers oblong, 0.5 mm. long (in bud),
connective oblong-lanceolate. Disk formed by 10 short, free scales.
Ovary glabrous, 5-locular, cells uniovulate. Style short. Stigma
capitate, 5-lobulate. Drupe globose, 15-16 mm. diameter with thin,
rather smooth epicarp (0.5 mm. thick); endocarp spheroid, about
14 mm. in diameter, woody, hard, with 5 minute holes at apex alter-

CUATRECASAS—HUMIRIACEAE 141

nating with 5 descending oblong opercules of about 6 mm. length,
remaining surface rugose.

H. mapiriense is endemic of the eastern slopes of the Bolivian
Andes not exceeding 1,000 m. elevation. It is a medium-size tree
easily recognized by its small, firm leaves and small, globose fruit.

BOLIVIA: La Paz: San Carlos, Sarampinni (Regién de Mapiri), 600 m.

alt., Buchtien 1518 (NY, holotype). Larecaja, Copacabana (south of Mapiri),

850-950 m., alt., Arukoff 11270 (US, paratype).

11. Humiriastrum diguense (Cuatr.) Cuatr., comb. nov. Fiaures 28,e-f; 29
Sacoglottis diguensis Cuatr. Trop. Woods 96:38. 1950.

Type: Cuatrecasas 14956, Colombia, Valle, Rio Digua.

Large tree with 60 em. thick trunk, young branchlets puberulous.
Leaves firm, coriaceous. Petiole very short (1-2 mm. long). Blade
obovate-elliptic, abruptly cuneate at base, rotundate and abruptly
and obtusely acuminate at apex, margin slightly and remotely crenate,

Ficure 29.—Humiriastrum diguense var. anchicayanum (Cuatrecasas 14418): a, bud; ),
petal; c, flower, petals removed, 5; d, detail of stamens, X 10, inside view and out-
side view; ¢, gynoecium and disk, X10; f, longitudinal section of ovary, X10; g,
transection of ovary, 10.

5.5-8.5 em. long, 2.5-5.5 em. broad; above dark green, glabrous,
midrib marked, filiform lateral nerves little conspicuous; beneath with
prominent midrib, lateral nerves ascendent, little prominent or
inconspicuous, spread with minute, fine, strigose, inconspicuous hairs.

Inflorescences terminal, dichotomous-paniculate, corymbiform,
longer than leaves (8-15 cm. long), branches robust, articulate, sub-
angulate, pubescent. Bracts small, ovate, ciliate, pubescent, decidu-
ous. Pedicels very short. Sepals 1-1.2 mm. long, suborbicular,
pubescent. Petals 2.2 mm. long, elliptic-oblong, appressed, estiva-
tion quincuncial. Stamens 20, glabrous, filaments connate at base,
unequal, longer 1.2 mm. alternating with shorter of 0.9 mm. length.
Anthers 0.8 mm. long, connective very thick, ovoid-lanceolate, obtuse,
2 thecae elliptic, basal. Disk formed by 0.2 mm. long scales. Ovary
subpyriform 1 mm. high, hirsute (at base glabrescent). Style short.
Stigma 5-lobate.

Humiriastrum diguense is an important species of the rain forests
of the Pacific slopes of Colombia and can be found from the bills at
low altitude up to about 1,200 m. elevation. It stretches northward

142 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

to Costa Rica where a subspecies is common in the forests near
Esquinas. Its wood is very hard and can be used for construction.

Key to the Subspecies and Varieties of Humiriastrum diguense

1, Leaves broad (5.5-8.5 X 2.5-5.5 em.). Sepals appressed-pubescent outside.
Inflorescences very robust. . . . . .1la. subsp. diguense var. diguense
1, Leaves narrower (3.5-9 X 1.5-4 em.). Sepals glabrous outside. Inflores-
cences more slender.
2. Petals sparsely puberulous. Branchlets puberulous.
llb. subsp. diguense var. anchicayanum
2. Petals subglabrous. Branchlets more densely hirtellous.
Ile. subsp. costaricense

lla. Humiriastrum diguense subsp. diguense var. diguense. FIGURE 28,¢

COLOMBIA: Ex Vatxe: Cordillera Occidental, vertiente occidental, Hoya del
Rio Digua, lado izquierdo; Piedra de Moler, selvas 1,140-1,180 m. alt.; gran
arbol, tallo 60 em. diim., copa grande, superiormente plana, hoja coridcea,
verde oscura, brillante haz; “arraydn negro,’ 20-VIII-1943, Cuatrecasas 14956
(holotype F; isotypes US, COL, G, VALLE).

llb. Wumiriastrum diguense subsp. diguense var. anchicayanum (Cuatt.)
Cuatr., comb. nov. Ficures 28,f; 29
Sacoglottis diguensis var. anchicayana Cuatr. Trop. Woods 96:39. 1950.
Type: Cuatrecasas 14418, Colombia, Valle, Hoya del Anchicay4.
Leaves subovate-lanceolate, acuminate, 5-9 em. long, 2-4 cm. broad.
Branchlets of inflorescence only puberulous. Sepals glabrous exteri-
orly, margin ciliate, about 0.8 mm. long. Petals quincuncial or
cochlear, sparsely pilose, 3-3.2 mm. long, 1.2 mm. broad. Filaments
united in lower third, minutcly papillose, the shorter 1.2 mm., the
longer ones 1.6 mm. Anthers about 0.7 mm. long with carnose,
lanceolate connective and basal, ellipsoid thecac. Disk cupular with
10 long narrow teeth alternating with other irregular shorter ones.
This variety is known only from the type locality in western
Colombia.
COLOMBIA: Vattg, Cordillera Occidental, Hoya del Rfo Anchicay4, entre
Pavas y Miramar, 350-450 m. alt., bosques; Arbol grande, cdliz verde, pétalos

blanco-verdosos, anteras amarillentas, hoja verde oscura, craso-coridcea, 16-IV—
1943, Cuatrecasas 14418 (holotype, F; isotypes, VALLE, COL, G, US).
Ile. Humiriastrum diguense subsp. costaricense Cuatr., subsp. nov.
PuaTE 14
Folia suboblongo-clliptica basi cuncata apice acuminata margine
crenata 3.5-6 crm. longa 1.8-3 cm. lata, petiolo circa 2 mm. longo
basi incrassato, supra nervis paulo visibilibus subtus prominulis et
costa eminenti, pilis tenuibus minutis adpressis sparsis inconspicuis
munita. Inflorescentiac subterminales vel terminales cymoso-panicu-
latae trichotomae sursum dichotomae pedunculis ramisque minute
hirtulis quam 8. diguensi tenuioribus. Sepala glabra margine ciliata,

CUATRECASAS—-HUMIRIACEAE 143

orbicularia, 0.9 mm. longa (in alabastra). Petala oblonga obtusa
parce puberula (in alabastra 1.7 < 0.9 mm.), aestivatione cochlearia,
raro contorta.

Type in the U.S, National Herbarium, No. 2085709, collected in
the Esquinas forest, region between Rio Esquinas and Palmar Sur
de Osa, province Puntarenas, Costa Rica, January 30, 1951, by Paul
H. Allen (No. 5812). Isotype in the Herbarium of Escuela Agricola
Panamericana, El] Zamorano, Costa Rica.

Subspecies costaricense resembles greatly variety anchicayanum,
which seems to be an intermediate form. The Costa Rican plant has
narrower leaves than variety diguense (like anchicayanum), more hirtel-
lous juvenile branches and inflorescences, glabrous sepals except for
the ciliate margin, petals almost glabrous and slender, and trichoto-
mous inflorescenses that are more ascendent and abundantly rami-
fied. The undeveloped stage of the flowers (very small buds) makes
difficult 2 more complete study of the plant, which in the future may
prove to be a different species.

About this subspecies in Costa Rica, Paul Allen states: “Very
tall trees, to 140 ft. in height, with alternate, elliptic-lanceolate,
glabrous leaves, the serrulate blades 1%-2’’ in length, with shortly
acuminate apices and nearly sessile, cuneate bases. The very small
greenish fragrant flowers are produced in terminal cymes in early
January and are soon followed by the oblong or ellipsoidal, drupaceous
fruits which are about 1 in. long. The reddish-brown heartwood is
very hard and heavy, and is reported to be durable in contact with the
ground. It is used locally for fence posts, and would appear to be
suitable for general heavy construction, such as bridge timbers, rail-
road ties, and piling. There is some indication that the wood of this
species may sometimes be confused with that of Vantanea barbourw
and it seems possible that this may be the species represented by the
sawmill sample from San Isidro del General forwarded to the Yale
School of Forestry by John A. Scholten and William F. Barbour in
1944. The species is fairly frequent in the forested hills near Esquinas”
(p. 317, 1956).

COSTA RICA: Puntarenas: Esquinas forest, between Esquinas and Palmar

Sur de Osa, ‘“‘nfspero,”’ tree 140 ft., flowers in bud and young fruit only, P. H.
Allen 5812 (type, US, EAP).
12. Humiriastrum procerum (Little) Cuatr., comb. nov.
Fiaures 26,a—d; 28,a—d
Humiria procera Little, Journ. Washington Acad. Sci. 38:93, fig. 2. 1948.
Sacoglottis procera (Little) Cuatr., Trop. Woods 96:40. 1950.
Type: Little 6412, Ecuador, Esmeraldas, Playa de Oro.
Large tree up to 40 m. high, trunk up to 1.20 m. diameter, buttressed
at base with granular bark and reddish brown, verv hard wood.

144 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Branchlets glabrous, more or less compressed, angulate, marked with
longitudinal wings from decurrent leaves. Leaves rather thick,
rigid, coriaceous, dark green, apparently glabrous, sessile. Blade
elliptic or ovate-elliptic, rotundate or very obtuse and amplectant
at base, abruptly and slightly narrowed or with short and obtuse
acumen 3-8 mm. long at apex, margin almost flat or slightly revolute,
sinuate-crenate, 5.5-12 cm. long, 3-7.5 cm. broad, brownish green
when dry; above lustrous, glabrous with flat conspicuous midrib,
secondary nerves visible and minor veins inconspicuous; beneath
with prominent, more or less carinate midrib, numerous secondary
nerves 2-4 mm. distant, slender, little prominent but conspicuous,
spreading, near margin reticulate-anastomosate, minor veins parallel
and reticulate, puberulous with thin, minute, appressed, inconspicuous
hairs.

Inflorescences cymose-paniculate, corymbiform, terminal or sub-
terminal, as long as leaves, peduncle robust, rigid, more or less
compressed, short-pubescent, trifurcate at apex; branchlets first
trifurcate upwards, dichotomous, compressed pubescent. Bracts
ovate-triangular, rather acute, amplectant, ciliate, little puberulous,
later deciduous. Pedicels very short (0.1 mm.), glabrous, articulate
to glabrous short peduncle (0.1 mm.). Sepals 0.7 mm. long, rounded,
glabrous except ciliate margin. Petals elliptic-oblong, scarcely
puberulous outside, 3-3.5 mm. long, estivation quincuncial. Stamens
20, filaments almost smooth, connate at base, 10 longer 2.6 mm.,
shorter 2mm. long. Anthers about 0.9 mm. long with 2 thecae at the
base (one at each side), globose-ellipsoid, 0.2 mm. long, connective
lanceolate, thick and angulate, cuspidate at apex. Disk 0.6-1 mm.
high. Ovary globose 1 mm. high, pubescent toward apex, glabrous
downward, 5-locular, cells episepal, uniovulate. Style thick, 0.6
mm. long. Stigma capitate, 5-lobate. Drupe oliviform, rounded
at base, subattenuate-subacute at apex, 2.8-3.8 em. long, 1.8-2.3
em. in diameter. Exocarp coriaceous when dry, resinous-granulose,
about 1.5 mm. thick. Endocarp up to 3.5X2 em., woody, without
or with very rare resiniferous cavities, surface rugose and alvcolate, 5
deep holes at apex alternating with 5 oblong descending opercula,
about 8mm. long. Usually 1, often 2, seeds developed.

H. procerum is one of the largest trees of the rain forests of the
Pacific slopes of Colombia and Ecuador. It is frequent at low
elevations on hills and on uninundatable grounds along the rivers.
It has a dense and dark green foliage and a buttressed trunk. The
wood is extremely hard but not resistant to decay. It took two men
2 hours to cut down one of my specimens with an ax (16615). For
this reason the natives resist cutting it, and chant therefore remains
more common in the forests than other more usable or practical

CUATRECASAS—HUMIRIACEAE 145

woods. E. L. Little writes about this species: ‘Large, erect tree of
forest canopy, 30 to 34 m. tall and 40 cm. in diameter, according to
measurements from trees felled for wood samples, but becoming
larger. Heartwood dull red, turning brownish on exposure, hard and
heavy. Wood fairly difficult to cut, rather harsh, readily polished,
strong but brittle, not resistant to decay. Not used much because
of alleged silica content, which dulls saws. Might be used for con-
struction where not exposed to the weather. It is said that the oil
seeds are edible. Common and probably widely distributed in the
wet tropical forest of Esmeraldas” (p. 247, 1948).

COLOMBIA: Eu Vatus: Costa del Pacifico, Rio Calima, La Trojita, loma en
la orilla izquierda, 30-40 m. alt.; gran 4rbol 40 m., 1.20 m. didm., estribos tabulares
grandes; corteza dura, adherida, granulosa, pardo rojiza; madera dura, castafio-
rojiza; hoja coridcea, rigida, verde oscura; capullos verdoso pélidos, “chant,”
frecuente, 1-III-1944, Cuatrecasas 16615 (F, COL, G, US, VALLE, Y). Rfo
Cajambre, Barco, margen izquierda de Agua Clara, Jestis, selva, 40-60 m. alt.;
gran drbol, 35 m., muy coposo, tallo 60 cm. didm. con robustos estribos basilares,
corteza granuloso pulverulenta, pardo rojiza clara (hacia arriba blanquecina) ;
madera rojiza oscura; hoja coridcea, rigida, verde oscura, mate; drupas muy
duras, verdes, del tamafio de oliva pequefia; corteza, madera y hojas aromaticos,
“chant,” 27-IV-1944, Cuatrecasas 17186 (F, COL, G, VALLE, Y).

ECUADOR: EsMERALDAS: Playa de Oro, alt. 65 m., wet tropical forest, slightly
cut, tree 31 m., DBH 42 cm., immature flowers green, “chanul,”’ 1-V-1943,
Little 6412 (holotypus US, isotypi Y); tree 32 m., DBH 42 cm., “chanul,’”’ 1-V—-
1943, Little 6413 (paratypus US, Y). 2 km. south of San Lorenzo, alt. 10 m.,
forest. wet tropical, partly cut; tree 34 m., DBH 40 cm., common; observed at
Quinindé Apr. 9 and seeds collected; good for general construction not in contact
with ground; seeds like pecan; flower buds greenish, twigs winged, “chanul,”
21-IV-1943, Little 6320 (US, Y). Quinindé, alt. 65 m., wet tropical forest
(only fruit, from ground), ‘“chanul,” 9-IV—-1943, Little 6233 (US).

13. Humiriastrum melanocarpum (Cuatr.) Cuatr., comb. nov.
Figures 26,r-t; 28,h-7

Sacoglottis melanocarpa Cuatr., Trop. Woods 96:37. 1950.

Type: Cuatrecasas 19989, Colombia, Valle, Buenaventura.

Large tree with grayish or brownish rugulose terminal branchlets.
Leaves coriaceous, subsessile, glabrous. Blade obovate, cuneate and
tapering to a very short petiole at base, subrotund but abruptly,
shortly and obtusely acuminate, the margin slightly crenate with
small glands; 4-7 cm. long, 2.5-4.5 em. broad, above green, nitid with
conspicuous midrib and obsolete lateral nerves, beneath with promi-
nent midrib, secondary nerves spreading, slightly prominent, near the
margin anastomosate.

Inflorescence axillary, subterminal, cymose-paniculate, 2-4 cm.
wide, peduncle terete, glabrous; branches dichotomous, articulate,

uberulous, 2-6 mm. long; bracts deciduous; pedicels up to 1 mm.
ong. Sepals 0.5 mm. long, almost free, ovate, obtuse, sparsely puber-

146 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

ulous with ciliate margin. Petals oblong, slightly broadened at base.
sparsely pilose, 2.6 mm. long, 1 mm. broad. Stamens 20, glabrous,
longer filaments about 2 mm. long, shorter 1.5 mm. long, alternating
linear, acute, united at base. Anthers 0.7 mm. long, glabrous, cordate-
lanceolate, connective thick, subobtuse, 2 thecae suborbicular, di-
vergent at base. Disk formed by several 0.4 mm. long dentate scales.
Ovary ovate-orbicular, 1.1 mm. high, glabrous, 5-locular or 2-3-
locular through abortion, cells uniovulate. Drupe ovoid or ovoid-
oblong, apiculate, acute, black and lustrous when ripe, 16-18 mm.
long, 10-13 mm. diameter. Endocarp woody, oblong-ovoid sharply
acute at apex with 5 minute foramina alternating with 5 small, oblong
descending opercules not surpassing half of the fruit, 14-15 mm. long.
7-8 mm. in diameter.

A very distinct species endemic to the Pacific coast of Colombia
it is known only from the cited collections.

COLOMBIA: Et Vauie; Costa del Pacifico, Buenaventura, Quebrada de
Aguadulce, 0-5 m. alt.; gran drbol; hoja coridcea, delgada, rigida, verde medio y
brillante haz, verde claro envés; perianto verde amarillento claro; fruto ovoideo.
apiculado, negro brillante, 18X13 mm., “chant,” 24-IIT-1946, Cuatrecasas 1998¢
(F, holotype; COL, G, W, VALLE, isotypes). Buenaventura, Quebrada de San
Joaquin, 0-10 m. alt.; drbol grande; hoja coridcea, subrigida, verde; frutos verdes.

oblongos, apiculados, 16-18X9 mm., ‘‘chand,’”’ 22-II-1946, Cuatrecasas 1990¢
(F, COL, G, W, VALLE, paratypes).

7. Schistostemon

Schistostemon (Urban) Cuatr., gen. nov.

Saccoglottis subgen. Schistostemon Urb. in Mart. Fl. Bras. 12(2):448, 445. 1877

Saccoglotlis Sect. Schistostemon (Urb.), Reiche in Engl. & Prantl, Pflanzenfam.
3(4):37, fig. 32. 1890.

Sacoglottis Sect. Schistostemon (Urb.), Winkl. in Engl. & Harms, ibid., 19a:128,
fig. 59-J. 1931.

Humirium Benth. in Hook. London Journ. Bot. 2:374. 1843 (in part); in
Hook. Journ. Bot. Kew Mise. 5:102. 1853.

Sepals 5, suborbicular, more or less united at base. Petals 5,
free, linear or oblong, thick, estivation quincuncial, cochlear or
contorted. Stamens 20, glabrous, very inequal; 5 opposite sepal
longer, trifureate at apex and triantheriferous, 5 opposite petal
less long, entire and monantheriferous, 10 intermediate shorter, monan
theriferous; filaments more or less complanate and thickish, united in
lower part up to middle. Anthers ovate or ovate-lanceolate, thecae 2
unilocular, ellipsoid or oblong, affixed on lower side, connective thick
more or less lanceolate rarely obtuse. Occasionally lateral anthers o
trifurcate stamens are stcrile. Disk cupular, dentate, or rarely of 1
free scales. Ovary 5-locular, cells uniovulate. Carpels opposite sepals
Ovules anatropous with ventral raphe, pending from inner angles o
ovary. Styles thick, short, up to 1 mm. long. Stigma capitate an

CUATRECASAS—HUMIRIACEAE 147

5-lobate. Drupe rather large, smooth, with thick carnous or sub-
coriaceous (when dry) exocarp. Endocarp woody, more or less bullate
with 10 very narrow, slightly apparent furrows, filled with many
globose resinous cavities; 5 septa in transection irregular and more or
less conspicuous; usually only 1 or 2 seeds developed, apparently
indehiscent, but the germinating embryos pushing away longitudinal
broad valves; these valves going from apex to base of endocarp,
apparently covering its surface; ribs between valves very thin; germ-
inal subapical foramina not observed. Evergreen trees with coria-
ceous or subcoriaceous, simple, alternate, petiolate, or sessile leaves with
usually more or less crenate margin. Inflorescences axillary or sub-
terminal, paniculate, with dichotomous or trichotomous branching.
Bracts persistent. (See also figs. 3, 30, and 31.)

Type species: Schistostemon oblongifolium (Benth.) Cuatr.

The name ‘‘Schistostemon” is derived from the Greek “schistos”’
(split) and “stemon” (filament), an allusion to the five trifurcate
stamens of this genus.

Schistostemon has seven known species and one subspecies mainly
distributed throughout the Guianas and the middle and northwestern
Amazon Basin.

Key to the Species of Schistostemon

1. Young branchlets minutely pilose.

2. Leaves oblong (4 times as long as broad), acuminate, crenate, minutely
papillose and sparsely pilose beneath. Inflorescence very small, di-
chotomous. Sepals and petals pubescent outside. Drupe elongate,
subfusiform or fusiform. Petiole 5-7 mm. long. . .1. S. oblongifolium

2. Leaves ovate, lanceolate or attenuate-elliptic (less than 3 times as long as
broad), subentire, glabrous, not papillose.

3. Connective of the anthers obtuse. Calyx glabrous except for the
minutely ciliate margin. Leaves very rigid, strongly and minutely
reticulate. Petiole 2-5 mm. long. Petals pubescent outside.

2. S. auyantepuiense

3. Connective of the anthers ovate-attenuate, acute. Calyx more or less
pilose and margin long-ciliate. Leaves less rigid with prominulous,
lax reticulum. Petiole2-8mm.long... . . . 3. S.reticulatum

4. Petals 4-4.5 mm. long, appressed- pubescent. Sepals sparsely pube-
scent. Leaves 6-12.5%4.5-7.5 cm., ovate or elliptic. Petiole 3-8
mm.long....... .3a. 8. reticulatum subsp. reticulatum

4. Petals 3.5-4 mm. long, subglabrous, with scarce hairs at apex. Sepals
densely hispidulous. Leaves smaller, 4—9.5 2-5 em., sublanceolate.
Petiole2-5mm.long. ... . .3b. S. reticulatum subsp. froesii

1. Young branchlets glabrous.

5. Petioles 0-1 mm. long. Leaf blade thick-coriaceous, broadly elliptic or
suborbicular, rounded or retuse at apex, margin subentire, sheath (or
very short petiole) very thick, venation lax reticulate and prominulous.
Drupe globose, 3—4.5 cm. diameter, endocarp 2.6-3.4 cm. .4. S. retusum

148 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

5. Petioles 6-14 mm. long. Leaf blade more or less attenuate at apex, more
or less crenate at margin.
6. Leaves minutely and conspicuously reticulate above, secondary nerves
prominulous and veins slightly conspicuous beneath. Petals thick,
glabrous, subobtuse or subacute, deciduous. Sepals broad, subecoria-
eceous. Inflorescences with alternate branches or lower ones dicho-
tomous; peduncle 1-3 em. long. Drupe oblong-ovoid, 3.5-42-2.8
em. Disk withunitedscales. ..... . . .5. S. macrophylium
6. Leaves prominently venose-reticulate on both sides. Inflorescence di-
chotomous. Petals and sepals submembranous, persistent.
7. Petals glabrous. Sepals ciliolate at margin. Peduncle of inflorescence

3.5-6 em. long. Disk with free scales. Drupe unknown.
6. S. dichotomum
7. Petals hirtellous-pubescent with retrorse hairs. Sepals hirtellous pubes-
cent, margin ciliate. Peduncle of inflorescence 1.5-3 em. long.
Disk annular, short, dentate. Drupe ovoid-ellipsoid, 2.62.3 em.
7. S. densiflorum

1. Schistostemon oblongifolium (Benth.) Cuatr., comb. nov.

Ficures 30,h-7; 31,g-i; 32,a

Humirium oblongifolium Benth. in Hook. Journ. Bot. Kew Mise. 5:103. 1853.

Sacoglottis oblongifolia (Benth.) Urb. in Mart. Fl. Bras. 12(2):447 pl. 93.

1877.—Ducke, Arch. Inst. Biol. Veget. Rio Janeiro 4:26,29. 1937.

Type: Spruce s.n., Brazil, Amazonas, Barcellos.

Small tree. Branchlets terete, ochraccous-brownish, lenticellate,
minutely hirtellous-pubescent or puberulous. Leaves firm, coriaceous
with 5-7 mm. long, thickened and more or less puberulous petiole.
Blade elliptic-oblong, sublanceolate, rounded or obtuse at base,
slightly decurrent along petiole, narrowed toward and acuminate at
apex, margin slightly serrate-crenate, 7-17 em. long, 3—-4.8 cm. broad,
acumen obtusely triangular, up to 1.5 em. long; above rather lustrous,
glabrous or flat midrib puberulous toward base, secondary nerves
obscurely prominulous, vein-reticulum lax, more or less conspicuous;
dull beneath, minutely papillose and with abundant, although diffuse,
subappressed hairs, midrib prominent, minutely pubescent or puberu-
lous, 9-12 pairs of spreading and prominent secondary nerves near
margin arcuate-anastomosate, veins forming a rather prominent
reticulum.

Inflorescences axillary, short-cymose-paniculate, dichotomous (rare-
ly trichotomous), peduncle robust, 4-10 mm. long, hirtellous, short]
branches rather thick, hirtellous. Bracts persistent, ovate, obtusish,
amplectant, pubescent, densely ciliate, 1.5-0.5 mm. long. Sepals
1 mm. long, rotund, imbricate, minutely pubescent, densely ciliate.
Petals greenish, rather thick, oblong, attenuate toward apex, subacute,
subappressed-pubescent, 4-5 mm. long, 1.5 mm. broad. Stamens 20,

CUATRECASAS—HUMIRIACEAE 149

Ficure 30.—a-f, Schistostemon retusum (Froes 22747): a, bud; b, petal; c, open flower with
petals removed, <5; d, stamens from outside and one another from inside, X 10; ¢, gynoe-
cium and disk, X 10; f, longitudinal section of ovary, X10. g, Schistostemon reticulatum
subsp. reticulatum (Ducke 23819), opening bud, X5. h-i, Schistostemon oblongifolium
(Froés 21090): h, opening bud, <5; i, detail of staminal tube, the longer stamen from
inside, X10. j-k, Schistostemon reticulatum subsp. froesit (Frées 21370): 7, bud 5;
k, petal, x5,

glabrous, filaments 2.7-3.5 mm. long united in tube up to middle,
5 longer ones short-trifurcate and bearing 3 anthers, 5 medium and
10 smaller alternating, undivided and each with one anther; sterile
filaments often present. Anthers ovate-lanceolate, about 0.9 mm.
long, thecae oblong, about 0.4 mm. long, connective thick, angulate,
lanceolate. Disk rather thick, cupuliform, about 0.6 mm. high, with
dentate margin. Ovary ovoid, glabrous, about 1.4 mm. high, 5-
locular, cells uniovulate. Style thick, 0.5-0.6 mm. long. Stigma
capitate, 5-lobate. Ovules about 0.6 mm. long. Drupe oblong,
subfusiform, 4—4.5 cm. long, 1.3-1.5 em. broad, attenuate at both ends,
apex acute, exocarp glabrous, almost smooth, thick, fibrous, easily
loosening; endocarp woody, subfusiform, rather smooth or slightly
bullate, about 3 * 1.2 cm., obtuse at base, apiculate at apex.

S. oblongifolium is a small tree found along the rivers of the Rio
Negro in Brazil and southern Venezuela.

513359—61——_9

150 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

ax Sy, AY
ASC:
) O20U y

neat

Ficure 31.—a-c, Schistostemon macrophyllum, X1 (Ducke 1175): a, fruit; b, endocarp; c,
transection. d-f, Schistostemon densiflorum, X1 (Persaud 102): d, fruit; e, endocarp;
f,transection. g-i, Schistostemon oblongifoltum, X 1 (Baldwin 3187): g, fruit; h, endocarp;
1, transection. 7, Schistostemon retusum, transection, *1 (Ducke 30131).

VENEZUELA: Amazonas: Rio Negro, prope San Carlos, 1853-54, Spruce 3073
(GH, P, NY). Ad flumen Casiquiari, Vasiva et Pacimoni, 1853-54, Spruce 3094
(NY). “In ripis Vasiva, XII-1853, one specimen taken from another tree with
narrower leaves, looks different from rest’’; “there was no more in flower’’ (Herb.
Benthamianum), Spruce 3194 (BM, K).

BRAZIL: Amazonas: Rio Negro, Serra de Sao Gabriel beira d’agua; arvore
pequena, 1-V—1947, Pires 588 (NY,IAN). Rio Negro, San Gabriel, 18-IX-1928,
Tate 142 (NY, US). Sao Gabriel, Igarapé Curucuhy, 27-XI-1945, Frées 21437
(IAN, K, NY, US). Sao Gabriel, ad ripas periodice inundatas; arbor parva,
floribus viridibus, 28-X-1932 (flores), 16-II-1936 (fructus), Ducke 23817 (US,
P, 8, U). Rio Negro, Jerusalem, terreno arenoso, beira rio nfo inundavel,
arbusto 3 m. em feixe; flores abundantes branco verde perfumativo, 1-VIII-1955,
Frées 21090 (NY, IAN, K, US). Rio Negro, inter Barcellos et San Isabel,
XII-1851, Spruce 1969 (isotypes, NY, M, GH). Rio Negro, mouth of Rio Igana;
tree to 20 ft. high, 5-III-1944, Baldwin 3187 (IAN, US). In flumini Negro
superioris ripis, XII-1854 (Herbarium Benthamianum), Spruce s.n. (holotype, K).
Rio Negro prope San Gabriel do Cachoeiros, VIII-1852, Spruce 2419 (P).

CUATRECASAS—HUMIRIACEAE 151

2. Schistostemon auyantepuiense Cuatr., sp.nov. FiGuRes32,b; 36,o—p; PLare 15

Arbor ramis terminalibus minutissime pilosulis.

Folia rigide coriacea fragilia glabra. Petiolus brevis robustus
inferne incrassatus 2-5 mm. longus. Lamina ovata vel ovato-
elliptica basi obtusa cuneata apice attenuata acuminataque acuta
raro obtusata, margine levissime crenata subintegra glandulis minutis
sparsis basi plus minusve conspicue biglandulosa; supra nitida costa
ampla prominula nervis nervulisque conspicue prominuleque reti-
culatis; subtus costa eminenti, nervis secundariis tenuibus sed pro-
minentibus 8-9 utroque latere arcuato anastomosatis nervulis minutum
reticulum prominentem formantibus. 4-8 cm. longa, 2. 5 cm. lata.

Inflorescentiae axillares subterminales breves cymoso-paniculatae
pedunculo 2-10 mm. longo breviter minuteque hirtulo-pubescenti
ramulis semel dichotomis vel omnibus alternis crassiusculis striatis
minute pilosis. Bracteae triangulares amplectentes persistentes
acutae 1-1.5 mm. longae margine minute ciliolatae, dorso minute
papillosae et sparse pilosulae. Pedicelli crassiusculi minute piloso-

Ficure 32.—a, Schistostemon oblongifolium (Ducke 23817); 6, Schistostemon auyantepuiense
(Vareschi &§ Foldats 4673); c, Schistostemon reticulatum subsp. froesit (Frées 21370);
d, Schistostemon retusum (Froes 22747), X34.
513359—61——_10

152 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

hispiduli 0.5 mm. longi. Calyx 5 sepalis crassis rotundatis margine
tenuibus minuteque ciliatis circa 1.5 mm. longis basi breviter coalitis
dorso minute papillosis et glabris. Petala cochlearia vel contorta
oblonga acutiuscula crassa 4-4.5 mm. longa 1.5 mm. lata dorso
papillosa minuteque pubescentia. Stamina 20 filamentis crassius-
culis complanatis tertio inferiore in tubum connatis 5 longioribus
(3 mm.) oppositisepalis apice trifidis triantheriferis antheris tribus
fertilibus, 5 mediis brevioribus (2.5 mm.) 10 intermedius minoribus
(2.2 mm.). Antherae glabrae ovatae 0.5-0.7 mm. bithecae, thecis
unilocis oblongo ellipsoidcis, connectivo crasso obtuso vel obtusissimo.
Discus membranaceus cylindraccus margine dentatus circa 0.8 mm.
altus. Ovarium pyriforme glabrus in stylum 0.5 mm. longum attenu-
atum 5-loculare loculis uniovulatis. Stigma breviter capitatum 5-
lobulatum.

Type in the Herbarium of Instituto Botanico, Caracas, Venezuela,
collected in montane forests above the Guayaraca camp, about
1,100 m. elevation, on Mount Auyantepui, State of Bolivar,
Venezuela, April 1956, by Volkmar Vareschi and Foldats (No. 4673).

Tree with minutely pilose terminal branchlets. Leaves. stiffly
coriaceous, glabrous. Petiole short, robust, 2-5 mm. long, the base
thickened. Blade ovate or ovate-elliptic, obtusely cuneate at base,
attenuate and acuminate at apex, rarely obtuse, margin slightly
crenate, subentire with minute punctiform glands, more or less
conspicuously biglandular at base; 4-8 cm. long, 2.5-5.2 cm. broad;
lustrous above with broad prominulous midrib, other nerves promin-
ulous, conspicuous, reticulate; midrib prominent beneath, secondary
nerves slender but prominent, 8-9 pairs, arcuate, anastomosate, veins
forming minute prominent reticulum.

Inflorescences axillary, subterminal, short, cymose-paniculate,
peduncle 2-11 mm. long, minutely hirtellous-pubescent, branchlets
first dichotomous or all alternate, rather thick, striate, minutely
pilose. Bracts amplectant, persistent, triangular, acute, 1-1.5 mm.—
long, ciliolate at margin, minutely papillose and sparsely pilose
outside. Pedicels rather thick, minutely pilose-hispidulous, 0.5 mm.
long. Sepals thick, rounded, about 1.5 mm. long, thin and ciliolate
at margin, papillose and glabrous outside, connate at base. Petals
4-4.5 mm. long, 1.5 mm. broad, oblong, subacute, densely papillose
and minutely pubescent; estivation cochlear or contorted. Stamens
20, filaments thick, complanate, lower third connate in tube, 5 longer
(3 mm.) opposite sepals, shortly trifurcate at apex and triantherifer-
ous (3 anthers fertile), 5 intermediate shorter (2.5 mm.) and 10
shortest (2.2 mm.) entire and monantheriferous. Anthers glabrous,
ovate, 0.5-0.7 mm. long, thecae unilocular, oblong-cllipsoid, connec-

CUATRECASAS—HUMIRIACEAE 153

tive thick, obtuse or very obtuse. Disk membranaceous, cylindrical,
about 8 mm. high, margin dentate. Ovary pyriform, glabrous.
Style 0.5 mm. long, attenuate, 5-locular, cells uniovulate. Stigma
capitulate, 5-lobate.

By its smaller and more rigid leaves and almost obovate connective
of its anthers, S. auyantepwiense differs essentially from the closely
related S. reticulatum. It is known only from Auyantepuf, one
of the Venezuelan Guayana hills, where it grows in the low forest of
the upper mountain. S. auyantepuiense is a xeromorphic endemic
characterized by its rigid, brittle, glossy, prominently reticulate
leaves; it is the species of the genus found at the highest elevation.

VENEZUELA: Borfvar: Cerro Auyantepuf, arriba del campamento Guay-
araca, 1,100 m. alt., Vareschi & Foldats 4673 (holotype, VEN).

3. Schistostemon reticulatum (Ducke) Cuatr., nov. comb.
Fiaures 30,9; 33,h
Sacoglottis; reticulata Ducke, Arch. Inst. Biol. Veget. Rio Janeiro 1:206.
1935; 4:26, 29. 1937.
Type: Ducke 23819, Brazil, Amazonas, Sdo Paulo de Olivenga.
Medium-size tree with minutely puberulous terminal branchlets.
Leaves firm, coriaceous, glabrous. Petiole thick, short, 3-8 mm. long.
Blade ovate or elliptic, obtuse or subrotundate at base, abruptly
attenuate, obtusely acuminate at apex, margin slightly crenulate,
6-12.5 cm. long, 4.5-7.5 cm. broad; above nitid, pale green, midrib
slightly prominent, 8-9 pairs of secondary nerves prominulous, sub-
patulous, near margin arcuate-anastomosate, smaller veins promi-
nently reticulate; midrib thick and prominent beneath, secondary
nerves prominent, reticulation lax, more or less prominulous.
Inflorescences axillary, cymose-paniculate, dichotomous at base,
peduncle compressed, striate, short-pubescent, branchlets minutely
pubescent-hirtellous. Bracts amplectant, persistent, ovate-triangular,
inferior obtuse, superior acute, 1-0.5 mm. long. Sepals 1.4 mm. long,
submembranous, imbricate, ovate, obtuse or subrotundate, puber-
ulous, margin ciliate, apex with small callous gland. Petals greenish,
oblong, rather acute, appressed-pubescent, 4—4.5 mm. long, 1-1.5
mm. broad, estivation cochlear. Stamens 20, glabrous, filaments
2.2-3 mm. long, complanate, united in tube at base, 5 longer shortly
trifurcate and triantheriferous. Anthers ovate, about 0.8 mm. long,
thecae elliptic-oblong 0.3-0.4 mm. long, connective ovate, acuminate.
Ovary subglobose, glabrous. Style short (0.4 mm. long), thick.
Stigma thick, capitate, 5-lobate.
Only fruit of subspecies froesii is known. The type specimen looks
glabrous at first sight, but careful examination of the only available
branchlet shows the existence of some minute hairs near the end.

154 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Ficure 33.—a, Schistostemon macrophyllum, <1 (Ducke 23816); b, Schistostemon macro-
phyllum, petal and bud, X34; ¢, Schistostemon dichotomum, *¥%s; d, Schistostemon
dichotomum, bud and petal, X3%; ¢, Schistostemon densiflorum, XM (Schomburgk
543); f, Schistostemon densiflorum, bud and petal, X 3% (Schomburgk 543); g, Schistoste-
mon densiflorum, bud and petal, X3% (Boschweyen 2068, “S. kaboeriensis”); h, Schisto-
stemon reticulatum subsp. reticulatum, leaf, X44 (Ducke 23819).

3a. Schistostemon reticulatum subsp. reticulatum FiauREs 30,9; 33,h
BRAZIL: Amazonas: Sio Paulo de Olivenca (Rio Solimdes), silva non
inundabilis; arbor mediocris floribus viridibus, 3-X—1931, Ducke 23819 (US, iso-

type).
PERU: Loreto: Mishuyacu, near Iquitos, alt. 100 m., tree 15 m., flowers
yellow-green, fruit edible, very ‘‘aromatic,” ‘“parinari,” VII-1930, Alug 1564 (US).

3b. Schistostemon reticulatum subsp. froesii Cuatr., subsp. nov.
Figures 30,j-k; 32,c; PLATE 16

Arbor parva ramulis terminalibus brunneis vel rufescentibus minute
hispidopubescentibus.

Folia firme crasseque coriacea simplicia. Petiolus 2-5 mm. longus
supra planus vel canaliculatus basi vaginatus juvenili pilosulus demum
glabratus. Lamina glabra ovato-lanceolata basi subrotundata et
breviter angusteque cuneata apicem versus gradatim attenuata et
subacuta raro obtusa, margine integra vel levissime sinuata, 4—9.5 cm.

CUATRECASAS—HUMIRIACEAE 155

longa 2—5 cm. lata; supra nitida viridula costa plana nervis secundariis
tertiarisque laxe reticulatis prominulis; subtus costa crassa elevata
nervis secundarius 6—7 utroque latere prominentibus nervulis prominu-
lis laxe reticulatis.

Inflorescentiae breves axillares inferne dichotomae pedunculo 1-6
mm. longo robusto striato minute hispido-pubescenti, ramulis brevibus
dense hispidulis, bracteis bracteolisque ovatis subacutis vel obtusis
pubescenti-hirtulis 1-1.5 mm. longis amplectentibus persistentibus.
Pedicelli crassiusculi hispiduli cirea 0.5 mm. longi. Sepala subro-
tundata 1 mm. longa 1.5 mm. lata basi coalita extus hirtulo-pubes-
centia, margine longe ciliata apice calloso-glanduloso excepto. Petala
crassiuscula oblonga subacuta 3.5—4 mm. longa 1.5 mm. lata subglabra
tantum dorso sursum parce pilosula, praefloratione cochleari. Stamina
20 glabra filamentis margine saepe leviter papillosis complanatis basi
in tubum 1 mm. altum vel ultra coalitis, 5 episepalis longioribus (2.5
mm.) tridentatis triantheriferis (antheris lateralibus sterilibus), 5
epipetalis (2 mm. long.) monantheriferis et 10 alternantibus breviori-
bus monantheriferis (1.5 mm. long.). Antherae ovato-acuminatae
circa 0.7 mm. longae thecis unilocularibus oblongo-ellipsoideis connec-
tivo crasso ovato-acuminato. Discus cupuliformis 0.8 mm. altus
margine denticulatus. Ovarium glabrum subglobosum 5-loculare
loculis uniovulatis. Stylus 0.3-0.4 mm. longum. Stigma capitatum.

Type in the Herbarium of the New York Botanical Garden, col-
lected at Foz de Cairay, Serra de Tunuhy, alt. 500 m., Igana drainage,
State of Amazonas, Brazil, November 13, 1945, by Ricardo de Lemos
Frées (No. 21370). Isotype in the U.S. National Herbarium,
No. 1496431, in Kew Herbarium and in Instituto Argonémico do
Norte.

Small tree with brownish or reddish and minutely hirtellous termi-
nal branchlets. Leaves firm, rather thick-coriaceous with 2-5 mm.
long pilose or glabrate petiole. Leaf blade ovate-lanceolate, rounded
and shortly angustate-cuneate at base, gradually attenuate and sub-
acute (rarely obtuse) at apex, entire or slightly sinuate at margin,
4~9.5>2-5 em.; above lustrous, greenish with flat midrib and promi-
nulous secondary and tertiary nervation; beneath midrib thick,
prominent, 6-7 pairs of secondary nerves prominent, reticulation lax
and prominulous.

Inflorescences short, axillary, dichotomous below, peduncle stout,
striate, minutely hispidulous, branchlets small, densely hispidulous,
bracts persistent, amplectant, ovate, subacute or obtuse, hirtellous.
Pedicels rather thick, hispidulous, about 0.5 mm. long.

Sepals subrounded, 1 mm. long, united at base, hirtellous outside,
ciliate at margin, with callous gland at apex. Petals thickish, ob-
long, subacute, 3.5-41.5 mm., subglabrous, scarcely pilose, up-

156 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

wards outside, estivation cochlear, Stamens 20, glabrous, filaments
complanate, more or less papillose, united in tube 1 mm. high at base,
5 episepalous longer, tridentate, triantheriferous, 5 epipetalous
shorter, uniantheriferous and other 10 still shorter alternating.
Anthers ovate-acuminate, about 0.7 mm., with unilocular oblong-
ellipsoid thecae and carnose, ovate-acuminate connective. Disk
cupuliform 0.8 mm. high, denticulate. Ovary glabrous, subglobose,
5-locular, cells uniovulate. Style 0.3-0.4 mm. long. Stigma capi-
tate, 5-lobate. Ovules anatropous with ventral raphe, suspended.

This subspecies differs from reticulatum in its smaller, more rigid
leaves, which are more cuneate at the base, and in the almost gla-
brous petals and the densely hispidulous sepals. The terminal
branchlets are conspicuously hispid-pubescent seen under the micro-
scope. S.r. froesiit represents a xeromorphic subspecies of S. reticu-
latum and is adapted to the hills near Rio I¢ana.

BRAZIL: Amazonas: Serra de Tunuhy, 500 m. alt., arvore 3 m., Foz do
Cairay, Rio Igana, Frées 21370 (type).

4. Schistostemon retusum (Ducke) Cuatr., comb. nov.
Figures 30,a-f; 31,7; 32,d; Phare 17
Sacoglottis retusa Ducke, Arch. Inst. Biol. Veget. Rio Janeiro 4:26, 29, pl.
2, fig. d. 1938.

Type: Ducke 30131, Brazil, Amazonas, Rio Curicuriary.

Small or medium-size tree with glabrous branchlets. Leaves
stiff, thick-coriaceous, sessile or subsessile with extremely short,
vaginate-thickened petiole. Blades broadly elliptic or suborbicular-
elliptic or slightly obovate, rounded or very obtuse at base, at apex
rounded, often retuse, rarely obtusely attenuate, margin subentire,
6-15 cm. long, 4.5-10 cm. broad; above almost smooth to conspicu-
ously nerved with small prominent midrib and lateral nerves and
veins more or less prominulous; prominent thick midrib beneath,
8-9 secondary nerves on each side, prominent, subspreading, near
margin arcuate-anastomosate, minor veins forming lax and more or
less prominent reticulum.

Inflorescences axillary, cymose-paniculate, much shorter than leaves,
peduncle erect, striolate, little compressed, pubescent-hirtellous, up
to 3 cm. long, branches short, alternate (rarely dichotomous), angulate,
hirtellous-puberulous. Bracts ovate or ovate-oblong, obtuse, amplect-
ant, more or less puberulous, ciliate at margin, 2—0.5 mm. long.
Ultimate branchlets short, hirtellous, 1-2 mm. long with 2-3 flowers.
Pedicels very short (0.2-0.3 mm. long). Sepals thick, rounded, 1-1.2
mm. long, 1.2-1.5 mm. broad, puberulous, margin ciliate and with
minute gland at apex. Petals thick, yellowish white, oblong, atten-
uate at apex, acute, 4.5-5 mm. long, 1.5 mm. broad, more or less

CUATRECASAS—HUMIRIACEAE 157

puberulous, estivation quincuncial. Stamens 20, uniseriate, glabrous,
filaments united in tube at base, 5 episepalous longer (2.5 mm.) shortly
trifurcate and triantheriferous, medial anther larger and fertile, lateral
anthers sterile, 5 epipetalous filaments (2 mm. long) and 10 alternating
and shorter ones (1.5 mm.) entire and monantheriferous. Anthers
dorsifixed, glabrous, 2 thecae basal, ellipsoid, unilocular, connective
thick, ovoid, interiorly carinate, apex acute. Disk membranous,
cupular, 0.8 mm. high, girdling ovary, margin about 20-dentate.
Ovary globose, glabrous, 5-locular, 5-ovulate. Style 0.4 mm. long.
Stigma capitate, 5-lobate. Ovule oblong-ellipsoid, anatropous with
ventral raphe. Drupe globose, 3-4.5 cm. diameter, exocarp nearly
smooth, coriaceous when dry, 3-5 mm. thick. Endocarp globose,
hard, woody, almost smooth, copiously filled with resinous cavities,
2.6-3.4 cm. diameter, usually with 2 oblong seeds.

S. retusum differs from S. reticulatum essentially in its elliptic,
broader, obtuse, sessile, or subsessile leaves with rounded base. The
very short petiole or base of the midrib is much thickened and rounded;
contrarily S. reticulatum has a very apparent and flattened petiole.
The leaf blades are thicker in S. retusum and more attenuate and acute
in S. reticulatum. 'The nerves are immersed and less conspicuous in
the type of S. retusum, but in other specimens that undoubtedly
belong to the species, they are more or less prominent.

S. retusum usually is a small tree found on sandstone elevations or
sandy soils in the upper Rio Negro and Vaupés basins.

COLOMBIA: Vaupés: Cerro de Circasia, 300-400 m. alt., drbol, frutos
grandes, comestibles, 10-X-1939, Cuatrecasas 7203 (COL; US, frag.). Riberas
del Rio Infrida; Raudal Guacamayo, margen izquierda, 180 m.; arbusto 3 m.,
inflorescencias péndulas, frutos péndulos, 4-II-1953, Ferndndez 2142 (COL, US).

BRAZIL: Amazonas: Rio Icana, Santana, caatinga aberta, terreno humido,
arenoso; arbusto formando tougas, frutos comestiveis, 18-XI-1945, Frées 21411
(IAN, K, NY). Rio Igana, Santana, caatinga, arvore pequena, 2—-V—1948,
Black 48-2514 (IAN, NY, U, US, VEN). Rio Curicuriary, afluente Rio Negro
super cataractem Caji ad ripas; arbor submedia, 22-II-1936, Ducke 30131 (US,

isotypes). Rio Preto, Campina, tree 25 ft. 7 in., white-yellow flowers, open
campina, rather grassy country, sandy soil, 5-XI-1947, Frées 22747 (IAN).
5. Schistostemon macrophyllum (Benth.) Cuatr., comb. nov.
Ficures 31,a—-c; 33,a—b; PLATE 18
Humirium macrophyllum Benth. in Hook. Journ. Bot. Kew Misc. 5:102. 1853.
Sacoglottis macrophylla (Benth.) Urb. in Mart. Fl. Bras. 12(2):446. 1877.—
Ducke, Arch. Jard. Bot. Rio Janeiro 6:40. 1933; Arch. Inst. Biol. Veget.
Rio Janeiro 4:26, 29. 1937.
Sacoglottis Ducket Huber, Bul. Mus. Goeldi 5:413. 1909.—Ducke, Arch.
Jard. Bot. Rio Janeiro 5:148, pl. 15, fig. 37. 1922.

Type: Spruce, Brazil, Amazonas, Barra do Rio Negro.
Small tree with greenish-brown, lenticellate, glabrous branchlets.
Leaves firmly coriaceous, glabrous. Petiole 8-14 mm. long, semi-

158 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

terete, flat above, thickened at base. Blade ovate-oblong, rounded
at base, attenuate and acuminate at apex, slightly crenulate at margin,
8-20 em. long, 3.5-7.5 em. broad; lustrous above with broad and
slightly prominent midrib, secondary nerves and reticulum sharply
prominulous; prominent striolate midrib beneath, 10-12 pairs of
secondary nerves obscure-prominent, subascendent, more or less
evanescent near margin, veins subimmersed, slightly conspicuous or
obsolete, small punctiform glands along margin.

Inflorescence cymose-paniculate, axillary, much shorter than leaves,
peduncle stout, compressed, short-pubescent, 1-3 cm. long, bifurcate;
branches short, alternate, pubescent. Bracts coriaceous, persistent,
ovate-triangular, rather obtuse, amplectant, carinate, glabrous but
ciliate margin. Pedicels thick, very short (about 0.3 mm. long).
Sepals thick, broad, orbicular, imbricate, about 1.5 mm. long, ciliate,
glabrous without. Petals thick, greenish, glabrous, oblong, obtusish
or subacute, 4.5-5 mm. long, 1.5-1.8 mm. broad. Stamens 20, gla-
brous, filaments compressed, lower half concrescent in tube, 2.5-4 mm.
long, 5 longer trifurcate and triantheriferous. Anthers ovoid or
rhomboid, about 1 mm. long with elliptic-oblong thecae about 0.5 mm.
long, connective thick, angulate, subdeltoid-lanceolate. Sometimes
sterile filaments found between the stamens. Disk annular-cupular
virdling ovary, formed by 10 bidentate, connate scales, about 0.6—-0.7
mm. long. Ovary subglobose, about 1.2 mm. high, glabrous, 5-
locular, with uniovulate cells. Ovules oblong, about 1 mm. long.
Style columnar, glabrous, 0.8 mm. long. Stigma capitate, 5-lobate.
Drupe subovoid or oblong-ovoid, subrounded at base, abruptly
attenuate at apex, subobtuse or shortly apiculate, smooth, 3.5—4 em.
long, 2—2.8 cm. broad, exocarp carnose (dry coriaceous), thin, endocarp
woody, rather smooth or slightly bullate, with abundant spherical,
resinous cavities.

S. macrophyllum is a frequent small tree on the flooded river margins
of the Manaos and middle Rio Negro regions.

BRAZIL: Amazonas: Manaos, Rio Tarum4, igapé, arvore pequena, flor verde,
12-VIII-1942, Ducke 255 (A, IAN, MG, NY, 5, US). Rio Tarumdé, arvore 7 m.
pendendo para o Rio, flores verde-brancas, 6-VIII-1949, Frées 24916 (IAN);
margen do rio, solo de areia, arvore de 5 m., 26-II-1950, Frées 26071 (IAN).
Margem do Igarapé do Tarumé4; Igapé6 de agua preta, arbusto de 3 m., flor esver-
deada, 22—VI-1955, INPA-1243 (MG—21545). Imgarapé de Cachoeira Alta de
Tarum4, arvore 8 m., flor esverdeada, 25-VIIJ-1954, INPA 244 (MG—21552).
In ripis inundatis Igarapé da Cachoeira Grande, arbor parva floribus viridi-
bus vix odoratis, 7-V—-1933, Ducke 23816 (US, U); margem inundada, arvore
pequena, 17-I-19438, Ducke 1175 (IAN, MG, NY, US). Igarapé do Cachoeira
Grande dos Bilhares, terreno firme arenoso, capoeira, flor esverdeada, estames

amarelos, arvore 6 m., caule tortuoso e muito esgalhado, 26-[X-1955, INPA 2044
(MG-21540); arbusto 3 m., 12—XII-1955, Chagas 3093 (IAN). Igapé6 do

CUATRECASAS—HUMIRIACEAE 159

Tarumazinho, agua preta, arvore caule tortuoso, 15-VII-1955, INPA 1407 (MG-
21544). Manaos, silva paludosa, arbor floribus albidis, 26-VIII-23, Kuhlmann
21029 (US, 8, U); silva paludosa, Ducke 21024, isotype of Sacoglottis ducket
Huber, photo F.M. 12596. Manaos, praia baixa do Rio Negro, arvore pequena,
flor verde, 1-IX-1945, Ducke 1744 (GH, IAN, MG, NY, US). Baixo Rio Negro,
Tanacoera, praia, 26-IV-1911, Ducke 11550 (MG). Rio Negro, Padauiry, Tapera,
tree 30 ft., yellowish white, flowers border of river of whitish water, low land, high
forest, 11-X-1947, Frées 22703 (IAN, NY, US, U). Rio Negro, Barcellos, Beira
de uma ilha no rio, 23-VI-1905, Ducke 7174 (MG, holotype of S. Ducket Huber).
Middle Rio Negro, Barcellos, flood bank, bush, flowers pinkish, 26-IX—-14—X-
1947, Schultes & Lépez 8881 (IAN, US). Rio Negro, Tauapecagu, shrub 20 ft.
growing in clumps, yellowish flowers, border of rivers, sandy soil, Frées 22472
(IAN); ad ripas inundatas fluminis Apuaht inferioris, arbor parva, floribus
viridibus odoratis, 25-VII-1929, Ducke 23432 (8, US). Margen do Igarapé do
Franco, arvore 5 m., frutos esverdeados, terreno arenoso, terra unida capoeira
fechada, 20-II-1956, Chagas & Dionisio 3472 (IAN). Itaubal, Rio Aracd, suba-
fluente do Rio Negro, caatinga, 26-X—1952, Frdées & Addison 29119 (IAN). Prope
Barra, “prov. Rio Negro,” VII-1851, Spruce 1714 (M, isotype). Mouth of Rio
Negro, Igarapé da Colonia at Loge, 21-VII-1874, flowers yellowish, Traill 81 (kK).
6. Schistostemon dichotomum (Urban) Cuatr., comb. nov. Figure 33,c-d

Sacoglottis dichotoma Urb. in Mart. Fl. Bras. 12(2):446. 1877.

Type: Kappler 2144, Surinam, Lava.

Tree with light-brown lenticellate terminal branchlets. Leaves
coriaceous, rigid, glabrous. Petiole thick, semiterete, 6-12 mm.
long, flat above, thickened at base. Blades oblong-ovate or ovate-
elliptic, rounded or somewhat cuneate-decurrent at base, attenuate
at apex, more or less cuspidate, margin slightly crenulate, 10-16 cm.
long, 4-8 cm. broad; lustrous above with almost flat midrib, secondary
nerves filiform and prominent; rather dull beneath with prominent
striolate midrib, 8-10 pairs of secondary nerves slender, prominent
subascendent, curvate and anastomosate with minute impressed
distant glands near margin; minute vein reticulation sharply prominent
and very conspicuous on both sides.

Inflorescences cymose-paniculate, dichotomous, axillary, half as
long as leaves, peduncle slightly compressed, minutely puberulous,
3.5-6 em. long, branchlets minutely pubescent, bracts amplectant,
ovate, obtuse, 1.5-0.5 mm. long, short-ciliolate, persistent. Pedicels
thick, up to 0.5 mm. long. Sepals rather thick, imbricate, sub-
orbicular, glabrous except the minutely ciliate margin, 1 mm. long.
Petals submembranaceous, yellowish, oblong, acute, glabrous, 3-3.5
mm. long, about 1 mm. wide. Stamens 20, glabrous, filaments
complanate, lower part connate, 2.2-3.3 mm. long, 5 longer shortly
trifurcate and triantheriferous at apex. Anthers ovate-rhomboid,
about 0.9 mm. long, thecae ellipsoid-oblong, 0.4-0.5 mm. long,
connective thick, angulate, ovate-lanceolate. Disk cupular, circling
ovary, formed by 10 bidentate loose scales, 0.5—-0.6 mm. long. Ovary

160 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

globose, glabrous, 0.8 mm. high, 5-locular, 5-ovulate. Style rather
thick, about 0.6 mm. long. Stigma capitate 5-lobulate.

S. dichotomum is only known from the region bordering Surinam
and French Guiana.

SURINAM: Ad fluvium Lava, arbor floribus flavis, October, Kappler 2144
(P, isotype). Marowijne River, 1861 (1862 ?), Kappler s.n. (U). Fluvium
Lawa, prope Cottica, X-1903, Versteeg 265 (U).

FRENCH GUIANA: Maroni, ‘‘M. le Dr. Rech, 1862” (P). Collector un-
known, “Guyane francaise” (P).
7. Schistostemon densiflorum (Benth.) Cuatr., comb. nov.
Ficures 31,d-f; 33,e-g; Phare 19
Humirium densiflorum Benth. in Hook. London Journ. Bot. 2:347. 1843.
Sacoglottis densiflora (Benth.) Urb. in Mart. Fl. Bras. 12(2):445. 1877.
Sacoglottis kaboeriensis Bakh. v.d. Brink f. apud Pulle in Rec. Trav. Bot.
Neerl, 37:292. 1940.—Medel. Bot. Mus. Herb. Rijks Univ. 79:292.
1940.— Pulle, Flora of Suriname 3(1):420. 1941.

Type: Schomburgk 543, British Guiana, Quitaro river.

Medium-size tree with terete, more or less lenticellate, glabrous
branchlets. Leaves coriaceous, more or less rigid, glabrous. Petiole
5-12 mm. long, semiterete, thickened at base. Blade ovate or ovate-
oblong, rounded at base, narrowed, subacute or shortly acuminate
at apex, margin slightly and distantly crenate, 7-19 em. long, 3-7.5
em. broad; lustrous above with flattened but conspicuous midrib and
prominulous secondary nerves and reticulum; lustrous beneath with
thick and prominent midrib, secondary nerves 10 pairs, prominulous,
arcuate-ascendent, anastomosate near margin, veins forming minute
prominent reticulum.

Inflorescences cymose-paniculate, axillary, dichotomous, alternate
at the end, peduncle robust, slightly compressed, minutely pubescent,
1.5-3 cm. long, branchlets rather thick, pubescent. Bracts persistent,
amplectant, 1.5-0.5 mm. long, ovate-lanceolate or sublanccolate,
ciliate and more or less puberulous. Pedicels very short (0.1-0.3 mm.).
Sepals ovate-orbicular, united at base, 0.7-0.8 mm. long, hispidulous-
pubescent, ciliate at margin. Petals linear, attenuate at apex,
3.5-4.5 mm. long, 1 mm. broad, hispidulous-pubescent with retro-
flexed hairs. Stamens 20, glabrous, filaments united in lower half,
complanate, 2-3 mm. long, 5 longer short trifurcate at apex, trianther-
iferous, others simple, acute, monantheriferous. Anthers ovoid,
about 0.7 mm. long, thecae ellipsoid, connective carnose, lanceolate.
Disk membranaceous, annular, 0.4 mm. high, dentate at margin.
Ovary glabrous, ovoid, about 1.2 mm. high, 5-locular, cells uni-
ovulate. Style about 1 mm. long. Stigma 5-lobate. Drupe ovoid-
ellipsoid, about 2.6 cm. long, 2.3 cm. diameter, exocarp almost smooth,
coriaceous when dry, 1.5-2 mm. thick. Endocarp ellipsoid, slightly

CUATRECASAS—HUMIRIACEAE 161

bullate, 2-2.3 cm. long, 1.8-2.2 cm. diameter, woody, resinous-
lacunose, usually with one seed.

The type of Sacoglottis kaboeriensis can be distinguished from the
typical Schistostemon densiflorum only by its larger and more flexible
leaves; undoubtedly it only represents a form of younger plants from
a more humid and shady habitat.

S. densiflorum is a tree growing mainly in the British and Dutch
Guiana forests at the margin of the rivers.

BRITISH GUIANA: “In Guiana,” Schomburgk 543 (isotype US, P). Esse-
quibo River, bank near Bartica, small tree, flower pale green; 5 petals, 10-15
stamens cohering below, 4-X—1929, Sandwith, 374 (NY, U). Bartica, XI-1888,
Jenman 4719 (NY); Jenman 2489, IX-1886 (BM). Comaka, Demerara River,
river bank, V-1923, Persaud 288 (F). Ruri, forest, clay soil, near Blue Moun-
tain, tree, VIII-1924, Persaud 102 (F, NY). Corantyne River, medium-size tree,
XI-1879, Jenman 287 (P); X-1879, Jenman 478 (P). County Berbice, Couran-
tyne River between Mapenna and Hubudikuru Creeks, 13-V-1918, ‘‘bukuria”’
(Arawak), Forest Department (Hohenkerk) 404A (K). “British Guyana,” VII-
1879, Thurn s.n. (K).

SURINAM: Corantyne River, Kaboerie, tree J.W.G. No. 138, 26-VI-1916,
Boschwezen 2068 (U, holotype of Sacoglottis kaboeriensis); tree 708, 30-II-1920,
Boschwezen 4933 (U); tree 593, 1-X—1920, Boschwezen 4960 (U).

8. Sacoglottis

Sacoglottis Mart., Nov. Gen. et Sp. Pl. 2:146. 1827.—Benth. in Hook. Journ.
Bot. Kew Miscel. 5:98, 103-104. 1853.—Benth. & Hook. Gen. Pl. 1:247
1862.—Muell. in Walp. Ann. Bot. Syst. 4:385. 1857.—Hemsl. in Hook,
Icon. Pl. 4 ser. 6:P1.2521. 1897.—Lemée, Dic. Deser. Synon. Gen. Phan.
5:919 (in parte). 1934.—Bakhuizen van den Brink in Pulle Fl. Surin.
8(1):417. 1941.

Sacoglottis Sect. Eusaccoglottis (Urban), Winkl. in Engl. & Harms, Pflanzenfam.
19a:128, figs. 58, 59. 1931.

Saccoglottis Endl. Gen. P1.:1040. 1840.—Baill. Adansonia 1:208. 1860;
2:265. 1861.—Ducke, Arch. Inst. Biol. Veget. Rio Janeiro 4:28-29. 1937
(in parte).—Walp. Rep. Bot. Syst. 1:425. 1842.—Hutchinson & Daziel,
Fl. West Trop. Afr. 1:274 fig. 114. 1928.

Saccoglottis Subgen. Eusaccoglottis Urb. in Mart. Fl. Bras. 12(2):442, 448,
pls. 94-I, 95. 1877.

Saccoglottis Sect. Eusaccoglottis (Urb.), Reiche in Engl. & Prantl, Pflanzenfam.
3(4) :37, fig. 32. 1890.

Houmiri Sect. Aubrya (Baill.), Baill. Adansonia 10:370. 1873.—Hist.

Pl. 5:54. 1874.
Houmiri Sect. Saccoglottis (Mart.), Baill. Adansonia 10:370. 1873.—Hist.
Pl. 5:54. 1874.

Aubrya Baill., Adansonia 2:265. 1862,—Oliver, Fl. Trop. Afr. 1:275. 1868.

Type species: Sacoglottis amazonica Martius.
Sepals 5, suborbicular, imbricate in estivation more or less united
at base. Petals 5, free, thick-membranaceous, estivation cochlear

162 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

or quincuncial. Stamens 10, glabrous, 5 opposite sepals longer than
alternate ones, filaments thick, complanate, oblong-lanceolate, united
at base. Anthers ovoid or ovoid-oblong, attached dorsally near
base, thecae 2, unilocular, ellipsoid, affixed at lower side, dehiscing by
detachment, connective ovate-acuminate, acute. Disk cupular,
dentate, girdling ovary. Ovary 5-locular, cells uniovulate, carpels
opposite sepals. Ovules anatropous with ventral raphe, pendulous.
Style as long as or exceeding stamens. Stigma capitate and 5-lobate.
Drupe of medium or large size, smooth, exocarp carnose, subcoriaceous
when dry. Endocarp woody, more or less bullate, globose, filled with
resinous cavities and commonly bearing only 1-2 oblong seeds.
Evergreen trees with simple, alternate, coriaceous or subcoriaceous,
petiolate leaves, with crenate or entire margin. Stipules small,
deciduous or lacking. Inflorescences axillary or subterminal, panicu-
late with dichotomous or trichotomous branching. Bracts persistent
or deciduous. Endocarp woody, more or less bullate with 10 very
narrow, usually slightly apparent furrows; filled with globose, resinous,
vacuous cavities; 5 septa in transection irregular and more or less con-
spicuous; commonly only 1 or 2 oblong seeds developed; apparently
indehiscent but at germination developing embryos push off broad,
thick, oblong-elliptic valves; these valves go almost from apex to
base and have width of two stripes (between every third furrow);
separating ribs being so thin that valves seem to cover whole surface
of endocarp; germinal foramina at apex very rare. (See also figs. 1, 3,
and 35-36.)

Sacoglottis has been spelled by Endlciher and others as ‘Sacco-
glottis,” but the original spelling by Martius is with one “ec.” Sprague
writes: “The generic name seems to have been derived from odxos
(sakos), more usually spelled cdéxxos (sakkos) a sack, and yAwrrs
(glottis) a little tongue in allusion to the tonguelike connective sepa-
rating the two thecae of the anther. There was therefore no ortho-
graphic error in the name Sacoglottis, and it must stand as published
by Martius” (Sprague, 1929).

The genus Sacoglottis, as here understood, comprises eight species
and several varieties and forms, which are spread widely throughout
tropical South America from east of the Andes to the Atlantic
coast; one species also is found on the Pacific drainage of Colombia
and Costa Rica, and another reaches the Atlantic side of this Central
American country. The southern limit in South America is in Rio de
Janeiro. S. gabonensis is the only extra-American species of the
family, and is found in tropical West Africa.

CUATRECASAS—HUMIRIACEAE 163

Key to the Species of Sacoglottis

1. Bracts deciduous.

2. Leaves small, blades up to 3.5-3 cm. long, obovate, firm-coriaceous, smooth
above, secondary nerves scarcely more conspicuous than minute, pro-
minulous reticulum beneath. Petals hispidulous outside. Sepals his-
pidulous-pubescent. .... . 2. ss... 1S. maguirei

2. Leaves larger, blades 6-22 cm. long, secondary nerves prominent and veins
lax-reticulate, prominulous beneath.

3. Drupe narrowly oblong, acute or apiculate, 4-5 em. long, 1.3-1.8 em,
diameter. Inflorescence sessile or subsessile, conglomerate. Calyx
broadly cupular. Petals glabrous. Leaves rigid-coriaceous.

2. S. ceratocarpa

3. Drupe ovoid, ellipsoid or subglobose. Inflorescences conspicuously
pedunculate,

4. Leaves rigid-coriaceous, rather thick, sharply nerved beneath. Drupe
ovoid, 5-5.54-4.5 em., dry exocarp 6-7 mm. thick.
3. S. ovicarpa
4. Leaves thin-coriaceous, flexible. Drupe ellipsoid or subglobose, dry
exocarp 1.5-2.5 mm. thick.

5, Sepals glandular at margin. Drupe oblong-ellipsoid, 5-6 3-3.5 cm.
Petals glabrous, 4-4.5X1 mm. subobtuse. Filaments united for
lower third. Disk dentate ......... 4 S.amazonica

5. Sepals not glandular or with very minute, scarce glands at margin.
Drupe globose-ellipsoid, 2.7-3.5%2.5-3 em. Petals pubescent
outside, 6-7 X 1.5-2 mm., acute. Filaments united briefly at base.
Disk fimbriate. . ...... =... +. + + + & S. gabonensis

1. Bracts persistent.

6. Drupe oblong, narrowed at base, acute or subobtuse at apex, 1.5-3 cm.
long, 0.9-1.2 em. thick. Leaves rigid-coriaceous, with immersed or
slightly conspicuous nerves above and more or less conspicuously minute-
reticulate beneath. . . . ee ew we ee ©)66 S. guianensis

6. Drupe spheroid or subspheroid,

7. Drupe 1.7-2.8 em. diameter; exocarp simple, compact, coriaceous,
resinous-granulose, 1-2 mm. thick; endocarp rather smooth, slightly
bullate and forrowed. Leaves thin-coriaceous, flexible, prominently
lax-reticulate both sides... . ... . . 7 S. mattogrossensis

7. Drupe 1.5-2 cm. diameter; exocarp 3-5 mm, thick, double, outer layer
coriaceous, compact, resinous-granulose, inner part densely fibrous;
endocarp more or less tuberculate. Leaves rigid-coriaceous, with
slightly or not conspicuous nerves above and prominulous, minute
reticulum beneath. . ........ =... . . 8 S. cydonioides

1. Sacoglottis maguirei Cuatr., sp. nov. Figure 34,f

Arbor parva, ramulis ultimis brunnescentibus lenticellatis minutis-
sime hirtulis.

Folia rigida coriacea petiolo 3-6 mm. longo supra canaliculato
subtus semitereti basi incrassato, minute hirtulo-puberulo. Lamina
obovata apice subtruncata vel subrotundata vel obtusissima basi
subite cuneata margine 4-6 dentibus obtusis distantibus utroque
latere, 2-3.5 cm. longa 1.5-3 cm. lata; supra nitida laevis tantum costa

164 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Ficure 34.—a-c, Sagoglottis gabonensis (Koechlin 2632): a, terminal branchlet, 1%; },
bud, X34; ¢, petal X3%4; d-e, Sacoglottis ceratocarpa: d, X% (Frées 21192); e¢, bud,
X 3% (Ducke 12); f, Sacoglottis maguirei, X¥3 (Maguire et al. 30693); g. Sacoglottis
amazonica, X ¥ (Ducke 1723); h, Sacoglottis ovicarpa, X M% (Cuatr. 19998).

conspicua; subtus subnitida costa prominenti, nervis secundariis
paulo distinctis nervulis minutum reticulum prominulum formantibus.

Inflorescentiae axillares foliis breviores, pedunculo 0.8-1.5 cm.
longo, robusto striato minute hirtello, ramis inferne dichotomis bre-
vibus ultimis fertilibus 2-4 mm. longis erectis minute hirtulis. Brac-
teae deciduae. Sepala rotundata 0.5 mm. longa (in alabastro 1.5 mm.
longo), pubescenti-hispidula. Petala oblonga subacuta extus his-
pidula, 1.3 mm. longa (in alabastro). Stamina 10 filamentis glabris
complanatis triangularibus, margine papillosis dimidia parte inferiori
coalitis, brevibus 1.2 mm. longis cum longioribus 1.6 mm. longis
alternantibus. Antherae oblongae 0.5 mm. longae thecis oblongis,
connectivo crasso apice obtusiusculo. Ovarium glabrum. Stylus 0.4
mm. longus. Stigma capitatum 5-lobatum. Fructus satis juvenilis
tantum vidi subglobosus glabrus numerosis cavitatis resinosis in-
structus.

The small, rigid, broadly dentate leaves of S. maguirei are different
from the leaves of all other species of the genus. This species is

CUATRECASAS—HUMIRIACEAE 165

xeromorphic, adjusted to the ecologic conditions of the high Guiana
plateaus.

Type in the U.S. National Herbarium, No. 2270195, collected at
Mount Cerro Yapacana, where it is frequent in cumbre, at an altitude
of 1,200 m., Orinoco River region, State of Amazonas, Venezuela,
January 3, 1951, by Bassett Maguire, Richard S. Cowan, and John J.
Wurdack (No. 30693). Isotype in the Herbarium of the New York
Botanical Garden.

Small tree with brownish, lenticellate, minutely hirtellous terminal
branches. Leaves small, coriaceous, rigid. Petiole 3-6 mm. long,
sulcate above, semiterete below, thickened at base. Blade obovate,
subtruncate, subrotundate or obtuse at apex, abruptly cuneate at
base, distantly and obtusely dentate at margin, 2-3.5 cm. long, 1.5-3
cm. broad; lustrous, smooth with only midrib conspicuous above;
less nitid beneath with prominent midrib, secondary nerves little
distinct and veins forming minute and prominulous reticulum.

Inflorescences axillary, shorter than leaves, peduncle 0.8-1.5 cm.
long, stout, striate, minutely hirtellous; branches dichotomous on
first divisions, short, last ones fertile, 2-4 mm. long, erect and minutely
hirtellous. Bracts deciduous. Sepals rounded, 0.5 mm. long (in
buds 1.5 mm. long), pubescent-hispid. Petals oblong, subacute,
hispidulous, in buds 1.3 mm. long. Stamens 10, filaments glabrous,
complanate, triangular, papillose, lower half concrescent, 5 longer
1.6 mm. long alternating with 5 shorter 1.2 mm. long. Anthers
oblong, 0.5 mm. long with oblong thecae and thick and subobtuse
connective. Ovary glabrous. Style 0.4 mm. long. Stigma capitate
5-lobate. Drupe very young, subglobose, glabrous and with many
resinous cavities.

VENEZUELA: Amazonas: Rio Orinoco, Cerro Yapacana, alt. 1,200 m.,
Maguire, Cowan, & Wurdack 30693 (US, holotype; NY, isotype).

2. Sacoglottis ceratocarpa Ducke, Bol. Tec. Inst. Agr. Norte 4:13. 1945.
Ficures 34, d-e; 35, i-k

Type: Ducke 12 and 1174, Brazil, Amazonas, Manaos.

Medium-size or small tree with hirtellous, soon glabrous, young
branchlets. Leaves rigid, coriaceous, glabrous. Petiole 7-13 mm.
long, robust, subterete, more or less winged, flat above, thickened at
base. Blade ovate-oblong or ovate-elliptic-oblong, rounded or obtuse
at base, attenuate and acuminate or cuspidate at apex, slightly
crenate at margin; 8-22 cm. long, 3.5-10 cm. broad, brownish when
dry; above smooth with almost flat midrib, secondary nerves and
veins obsolete; beneath with prominent midrib, secondary nerves
thin but prominent, 10-12 pairs, subspreading, near margin arcuate,

166 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Ficure 35.—a-c, Sacoglottis amazonica, * 1 (Archer 7964): a, fruit; b, endocarp; ¢, transec-
tion. d-e, Sacoglottis ovicarpa, X1 (Cuatr. 19998): d, fruit; e, transection. f-h,
Sacoglottis gabonensis, X 1(Krukoff 121): f, fruit; g, endocarp; A, transection. i-f,
Sacoglottis ceratocarpa, X1 (Ducke 1174): 7, fruit; 7, endocarp; k, transection.

ascendent, anastomosate, minor veins forming a lax-prominulous
reticulum.

Inflorescences axillary, cymose-paniculate, dichotomous, very short,
sessile, subglomeriform; peduncle almost absent, branchlets reddish,
angulate, short-pubescent, hirtellous. Bracts ovate-triangular, acute,
subglabrous, 0.5-1 mm. long, deciduous. Pedicels very short articu-

CUATRECASAS—-HUMIRIACEAE 167

late to very short peduncles. Sepals about 1.5 mm. long, rather
thick, orbicular, imbricate, glabrous except for ciliate margin. Petals
greenish, thick, glabrous, linear, subacute, about 5 mm. long, 1 mm.
broad. Stamens 10, united in lower half, glabrous, filaments com-
pressed, 2.5 and 3 mm. long alternating. Anthers ovoid-oblong,
about 1.2 mm. long, thecae elliptic about 0.5 mm. long, connective
thick, ovoid-sublanceolate, subacute. Disk membranaceous, denticu-
late ring 0.7-0.8 mm. high. Ovary ovoid, glabrous, 5-locular, cells
uniovulate, attenuate at apex. Style 2-3 mm. long, erect, glabrous.
Stigma capitate, 5-lobulate. Drupe elliptic-oblong-clongate, attenuate
at both ends, subfusiform, apex acute, often apiculate, 4-5 cm. long,
1.3-1.8 cm. diameter, plus apiculum up to 8 mm. long. Exocarp
smooth, subcoriaceous when dry, 1 mm. thick. Endocarp woody,
narrowly oblong, acute at apex, resinous-lacunose, surface slightly
bullate.

S. ceratocarpa is limited to the upper northern Amazon Basin.
Its presently known range extends from the lower Colombian Vaupés
region to the Manaos area.

A. Ducke wrote the following interesting remarks about the dis-
tribution of this species and its relation to S. amazonica: ‘This species
has been confused with S. amazonica since the time of Martius who
cited his S. amazonica not only for the Amazon estuary but also for
the upper Amazon where this species probably does not exist (he
certainly did not know the fruits of both species). The true S.
amazonica grows rather frequently on the shores of streams subjected
to the influence of the Atlantic tide; I observed it near Belém, on
the Islands of Breves, and near Gurup4 at the head of the Amazon
estuary. Its fruit has a thin, more coriaceous than fleshy, mesocarp
and is apt to float; old endocarps can be found every time on the banks
of the rivers around the city of Para. S. ceratocarpa, on the contrary,
crows in the central part of Amazonia in marshy upland forest along
streamlets, and its fruits do not seem apt for water transport, but
they are probably dispersed by forest animals. Their white, more
farinaceous than fleshy, mesocarp is somewhat like that of S. heterocarpa
Ducke, growing in ‘caatinga,’ in the upper Rio Negro basin.”

COLOMBIA: Amazonas-Vaupks: ‘Cafio Oogé-’dja,” Jinogojé, flowers white,
tips orange, “‘nee-saw-kaw’-ké-too” (Makuna), 26-VIII-1952, Schultes & Cabrera
17045 (US). Vaurfis: Between Mitdé and Javaraté, Tipiaca, Igarapé Murutinga,
small tree, 14-24-V-1953, Schultes & Cabrera 19290a (US). Rio Piraparand
(tributary of Rio Apaporis), Cafio Teemeena, 6-IX-1952, bush, flowers yellowish,
6-IX-1952, Schultes & Cabrera 17253 (US). Rio Paca (tributary of Rio Papurf),
Uacaricuari and vicinity, 650 ft., tree, flowers yellowish, 1-3-VI-1953, Schultes &
Cabrera 19519 (US).

BRAZIL: Amazonas: Manaos, Pensador, silva paludosa secus rivulum; arbor
mediocris fructibus maturis viridiflavis, “‘uchy-rana,” 23-I-1943, Ducke 1174

513359—61——-11

168 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

(IAN, NY, US, isotypes; MG, holotype). Pensador, silva paludosa secus rivulum;
arbor mediocris, floribus viridibus, “uchi-rana,’’ 20-VIII-1935, Ducke 12 (isotype,
florum, A, NY, 8, US). Pensador, silva paludosa non inundabili secus rivulum;
arbor mediocris floribus viridibus, ‘‘uchi-rana,”’ 31-VII-1943, Ducke 1301 (IAN,
MG, NY, US, A). Manaos, silva paludosa circa Cachoeira do Mindu, arbor
media floribus viridibus, 15-VII-1929, Ducke 23431 (paratypes, US, U). Manaos,
beira do Rio Tarum4é, Igapo; arvore 5 m., flores verde-brancas, ramos pendentes,
7-VITI-1949, Frées 24934 (IAN). Rio Tarumé, beira alagada perto da cachoeira
baixa, arvore pequena, 11—-I-1949, Guedes 80 (IAN, US); 25-VII-1948, Guedes
58 (IAN, US). Estrada do Tarum4, terreno arenoso, capoeira; flor esverdeada,
arvore 7 m., caule ereto, 4-VIII-1955, INPA 2084 (MG 21539). Igarapé da
Cachoeira do Tarum4, mata de varzea; flor amarelo esverdeada, 2—IX—1955,
INPA 1814 (MG 21547). Manaos, Estrada da Forquilha, terra umida capoeira;
arvore de caule tortuoso, flor verde, 12-VIII-1955, INPA 1638 (MG 21543).
Silva non inundabili ad margines paludosas rivuli, arbor sat magna floribus
viridibus, ‘“uchi-rana,” 31-VIII-1931, Ducke 16, Ducke 16a (Y). Rio Papury,
Vaupés-Rio Negro, terra firme, terreno silicoso, arvore 20 m. 30 em., 17—VIII-
1945, Froes 21192 (NY, US, K, IAN).

3. Sacoglottis ovicarpa Cuatr. Trop. Woods 96:39. 1950.
Ficures 34,h; 35,d—-e

Type: Cuatrecasas 19998, Colombia, Valle, Buenaventura.

Large tree with 50 cm. thick trunk, rugose and dark bark that is
reddish or pink in section, hard, yellowish pink wood, subterete and
glabrous branchlets. Leaves coriaceous, glabrous, reddish brown when
dry. Petiole subterete, 8-14 mm. long, robust, thickened toward base.
Blade elliptic or ovate-elliptic, rounded or obtuse at base, abruptly
narrowed and cuspidate at apex. Margin slightly crenulate, 9-16 cm.
long, 3.5-9 em. broad; above yellowish greenish, lustrous, with
conspicuous midrib and inconspicuous lateral nerves and reticulum;
beneath pale green, midrib prominent, the prominent lateral veins
spreading distantly, 8-10 pairs, arcuate-anastomosate near margin,
minor nerves conspicuously reticulate and prominulous. Drupe ovoid
or ovoid-ellipsoid, approximately form and size of hen egg (5-5.5 X
4—4.5 cm.), greenish, nitid; exocarp carnose, hard and coriaceous when
dry, 6-7 mm. thick; endocarp woody, almost smooth, irregularly
5-septate, abundantly provided with large rounded, nitid, resinous
cavities; usually 1-2 seeds, rarely 5. Inflorescences and flowers
unknown.

Collected by the author on the west coast of Colombia, S. ovicarpa
is a large hardwood tree that probably grows along the entire Pacific
coast in the uninundatable forest from Ecuador to Costa Rica. It
replaces on this side of the continent the Atlantic large-fruiting species
S. amazonica, the Andean chains separating the two species. The
Pittier specimens (16260) from the Cocos Islands on the Pacific coast
of Costa Rica are almost sterile (they have only very young fruit),
but I have almost no doubt that these specimens belong to this spe-
cies. Likewise belonging to S. ovicarpa are the endocarps encoun-

CUATRECASAS—HUMIRIACEAE 169

tered by I. Johnston (1949, p. 52) among the drift deposits at the
San José Island shores (Panama). These fruit, as suggested by
Johnston, undoubtedly originated on the southern Pacific slopes
(Panama to Ecuador) and drifted northward with the oceanic cur-
rent. The fruit of S. ovicarpa differ from those of the very closely
related S. amazonica by being ovoid and having a thicker and tougher
exocarp; the endocarp has the irregularly distributed resinous cysts
separated by thin walls, breakable at the surface, whereas in S. ama-
zonica the resinous cysts are conspicuously arranged in 10 rows with
thicker and stronger woody walls.

COSTA RICA: Cocos Island, Pacific, Le plus grand arbre de I’fle; appelé
“palo de hierro” par les colons; caracteristique pour les foréts, 10-250 m., I-1902,
Pittier 16260 (US). The material is almost sterile including the leafy branchlets
and a few initiations of fruit; more complete collections will be necessary to
assure this identification.

COLOMBIA: Et VaLue: Costa del Pacifico, Bahia de Buenaventura, Quebrada
de Aguadulce, 0-10 m. alt.; gran drbol, hoja coridcea, verde amarillenta oscura;
fruto ovoide-elipsoideo, 5.54.5 cm., exocarpo carnoso-duro, verde brillante, 7-7
mm. grueso, endocarpo lefioso, “guayabito”, 24-II-1946, Cuatrecasas 19998
(holotype F, isotype VALLE). Bahia de Buenaventura, Quebrada de San
Joaquin, 0-10 m. alt.; gran Arbol, tallo 45 cm. didm., corteza sublisa, pardo-
rosada, en seccién rosada o roja, en la base del tallo gris verdosa clara y separable;
madera dura, amarillo-rosada; hoja coridcea rigida, dura, frdgil, verde oscura
brillante haz, verde claro envés, “‘corosillo,’’ 22-II-1946, Cuatrecasas 19927
(F, VALLE). Rio Cajambre, Barco, 5-80 m. Cerro El Sapote en el brazo del
Rio Agua Sucia; drbol 30 m., tallo 50 cm. diam., corteza rugosa, pardo-rojiza,
seccién rosada, zumosa; madera blanco-ocrdcea, zumosa; frutos ovoideos, verdosos,
carnoso-coridceos, endocarpo semilefioso, 6X5 cm., “chanusillo,” ‘“chanosillo,”’
“chanesillo,” 28-IV-1944, Cuatrecasas 17226 (paratypes F, VALLE).

4. Sacoglottis amazonica Mart. Nov. Gen. Sp. Pl. 2:146. 1827; Fl. Bras.
(12)2:449, tab. 95. 1877.—Ducke, Arch. Jard. Bot. Rio Janeiro 3:179.

1922; 5:142, pl. 14, fig. 36. 1930; Arch. Inst. Biol. Veget. Rio Janeiro

4:26 and 29. 1937.—Hemsl. in Hook. Icon. Pl. 4 ser., 6:Pl. 2521. 1897.
Figures 1,d-f; 34,9; 35,a-c; 36,a-g; PLate 20

Type: Martius, Brazil, ‘“Crescit in sylvis secus fluvium Amazonum
et in ripa canalis Tagipuru.”

Medium-size tree with glabrous young branchlets. Leaves thin-
coriaceous, flexible, glabrous. Petiole 5-12 mm. long subterete,
slightly winged, sulcate above, lower portion thickened. Blade
oblong-elliptic sublanceolate, obtusely cuneate or very obtuse at
base, narrowed toward apex and acuminate-cuspidate, slightly crenate
at margin; 6-15 cm. long, 2.5-5.7 cm. broad, acumen about 1 cm.
long; above with flat midrib, secondary nerves and veins when young
more or less prominulous, later almost inconspicuous; beneath with
eminent striolate midrib, secondary nerves, 9-12 pairs, thin but
prominent, subpatulous, extreme ascendent, curvate-anastomosate,
veins laxly reticulate, more or less prominent.

170 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Ficure 36.—a-g, Sacoglottis amazonica (Ducke 1723): a, bud, <5; b, petal, X55 ¢, detail of
staminal tube from outside and one anther from inside, X10; d, gynoecium and disk,
X10; ¢, longitudinal section of ovary; f, transection of ovary, X10; g, seed, X20
h-n, Sacoglottis mattogrossensis fma. subintegra (Ducke 23820): h, bud, 5; 4, j,
petals (more or less developed), 5; &, flower with petals removed X5; J, stamen
dorsal view, and anther inner view, X10; m, gynoecium and disk, X10; n, gynoecium,
longitudinal section. o-p, Schistostemon auyantepuiense (Vareschi &3 Foldats 4673):
0, opening bud, 5; p, anthers, X 10.

Inflorescences cymose-paniculate, axillary, much shorter than leaves,
peduncle subterete, striate, glabrous, branchlets alternate, striolate,
glabrous or upwardly sparsely pilose. Bracts deciduous, ovate-tri-
angular, acute, about 1 mm. long. Pedicels short, thick, glabrous,
often articulate to short (about 1 mm.) peduncles. Sepals 1-1.5 mm.
long, broadly orbicular, imbricate, thickened, glabrous outside, margin
minutely ciliate and with small glands. Petals linear-oblong, sub-
obtuse, glabrous, about 4 mm. long, 1 mm. broad (at base 1.5 mm.
broad), whitish-greenish. Stamens 10, glabrous, filaments com-
planate, smooth, lower third connate, 5 shorter 3 mm. long, lower
ones about 4 mm. long. Anthers ovate-oblong, about 1 mm. long
with half long thecae, connective thickened, angulate, sublanceolate.
Disk girdling ovary, dentate, 0.6-0.7 mm. high. Ovary ovoid, about
1.5 mm. high, glabrous. Style 2.5-3 mm. long. Stigma capitate,
5-lobate. Drupe oblong-ellipsoid, 5-6 cm. long, 3-3.5 em. diameter.

CUATRECASAS—HUMIRIACEAE 171

Exocarp more or less smooth, coriaceous when dry, 1.5-2 mm. thick.
Endocarp slightly and irregularly 10-sulcate and bullate, woody,
filled with resinous cavities, usually with only 1 oblong seed.

Sacoglottis amazonica varies from a large to a small tree present in
flooded forests of the Amazon Basin, chiefly at its estuary. This
species is also recorded from the Orinoco delta, from British and
French Guiana, Trinidad, and some islands of the Lesser Antilles.
Sterile collections from Costa Rica have foliage that agrees completely
with the foliage of this species; however, I consider these collections as
provisionally belonging to this species until flowering and fruiting
material is made available.

The ellipsoid, 4-6 cm. long fruit (dark purplish according to
Broadway) are a remarkable feature of S. amazonica; its endocarp
filled with resinous, vacuous cavities is light and apt to float and be
carried by the rivers and currents. Ducke states that endocarps are
always on the banks of the rivers around the city of Para. But these
endocarps have also been found at many other places. Picked up
by currents at the Amazon and Orinoco deltas, the endocarps have
been drifted to the shores of the West Indies and Central America,
and strangely enough even across the Atlantic Ocean to Great Britain.

The hardwood of S. amazonica is used in house construction. The
most common name for the species is ‘‘uchi-rana” in Brazil, with some
phonetic and spelling variants. In British Guiana it is called “funyu”
and “nabaru”’ by the Guaraos of the Orinoco delta. According to
Devenish the local name in Trinidad is ‘‘cojén de burro.”

COSTA RICA: Limén: Hills above La Florida, alt. 750 ft., rain forest, associated
with “pejiballito,” “pilén,” “plomillo,” “bernabé,’”’ volcanic loam, tree 60 ft.,
DBH 12 in., ‘“campana,”’ 7-V-1943, Dayton & Barbour 3004 (US, Y, 40888),
sterile specimens. Hills 4 miles south of Sequirres, alt. 500 m., north slope, clay
loam, rain forest tree 80 ft., DBH 28 in., usable length 40 ft., “campana,” 7-V-
1943, Merker, Scholtes, & Dayton 3041 (Y, 40842), sterile specimens. These
identifications have to be considered provisional until fruit and flowering speci-
mens from the same region are made available.

TRINIDAD: Palo Seco, washed upon beach, 20-III-1920, Britton, s.n. (NY).
Irois, 28-XI-1916, Broadway 8475 (NY). Cedros, forest, 18-VIII-1896, Lunt
s.n. (BM, US). Irois, VI-1896, Lunt 5984 (NY, K); Hart (NY).

ST. VINCENT: L. Guilding s.n. (K).

VENEZUELA: Detra Amacuro: Antonio Diaz, Cafio Jobure, occasionally
in flooded forest along upper part; tree 15-20 m., buds greenish white; fruit
green, edible, used by Guaraos for diarrhea treatment; ‘“‘nabaru” (Guarauno);
wood heavy used locally in heavy construction, 7-IV-1955, Wurdack 293 (NY,
US, VEN).

BRITISH GUIANA: Rain forest on hilltop 8 miles east of Onoro Creek mouth,
alt. 1000 ft., “funyu,” 30-IX-1952, Guppy 308 (NY).

BRAZIL: Pari: Belém, silva inundata, arbor media floribus viridibus 24-
VIII-1922, Ducke 17781 (S). Belém, Pires & Black 31 (P, U); south forest of
the IAN, “achua-rana,” 2—XII-1942, Archer 7915 (IAN, US); same place,

172 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

13—XII-1942, “uchyrana,” Archer 7964 (IAN, US). Belém, Utinga, igapo,
margen de riachos, 14—X-1916, Ducke 16578 (MG, US, U). Utinga, Igapo
Buiussuquara; arbor parva floribus viridibus, “uchi-rana,” 13-VII-1945, Ducke
1723 (A, IAN, MG, NY, US). Gurupa, margen de um riacho na varzea; arvore
mediana, flor verde; ‘“uchi-rana,” 17-VIII-1918, Ducke s.n. (IAN); Ducke 17221
(BM, US, P, MG). Beira do Rio Guama, entre 8S. Miguel e Acary; arvore
pequena; fruto perto d’agua 7 X 3 cm., verde, flor branca verdusca, 31—-X—1948,
“uchi-rana,” Black & Foster 48-3393 (IAN). Beira do Rio Mapua, entre Vila
Emilia e Boca do Mapua, varzea; arvore, flor branca, 18-VII-1950, Black, Frées,
& Ledoux 50-9810 (US). Rio Curupere, afluente do Rio Abaete, igapo, municipio
de Abaetetuba; arvore, fruto verde, “uchirana’, 19-IX-1952, Araujo 6 (IAN).
Arama, beiro do rio, “uchi-rana,’’ 26-II-1900, Huber 1850 (MG). Brasilia in
itinere amazonico in ripa (canalis) Tagipuru, Obser. 2671, Martius s.n. (M, holotype;
M, isotypes). Amazonas: Esperanga (ad ostium fluminis Javary), silva non
inundabili; arbor magna, floribus viridibus, 30-X-1942, Ducke 1055 (NY, MG,
IAN, US). Municipio Humayta near Tres Casas, on low terra firma, tree 80
ft., 7-X-1934, Krukoff 6506 (GH, NY, U). Brasilia in sylvis ad Egam prov. Rio
Negro, Dr. Martius iter Brasiliensium, Martius s.n. (M, paratypes).

5. Sacoglottis gabonensis (Baill.) Urb. in Mart. Fl. Bras. 12(2):449. 1877.—
Reiche in Engl. & Prantl, Pflanzenfam. 3(4):37, fig. 32. 1890.—Winkler
in Engl. & Harms, Pflanzenfam. 19a:128, fig. 58. 1931.—Hutchinson
& Daziel, Fl. West Trop. Afr. 1:274, fig. 114. 1928.—Exell, Journ. Bot.
65 (Supl. Polypet.):50. 1927 (as S. gabunensis).—Guinea, E., Ensayo
geobotdnico de la Guinea continental espafiola, 300. 1946.—Exell &
Mendoga, Conspectus Fl]. Angolensis 1:249. 1951.

Figures 34,a-c; 35,f-h

Aubrya gabonensis Baill. Adansonia 2:266. 1862.—Oliver, Fl. Trop. Afr.
1:275. 1868.

Houmiri gabonensis Baill. Hist. Pl. 5:52. 1874.

Aubrya occidentalis A. Chevalier, Expl. Bot. Afr. Occident. Fr. 1:94. 1920,
nomen.

Type: Aubry-le Compte, Africa, Gabon.

Large tree with glabrous terminal branchlets. Leaves thin-
coriaceous, flexible, glabrous. Petiole 6-10 mm. long, narrowly
winged toward top, thickened at base. Blade ovate-elliptic or
elliptic-oblong, suddenly and obtusely narrowed at base and slightly
decurrent on petiole, narrowed-acuminate at apex, slightly crenate;
7-14 cm. long, 3-7 cm. broad; flat smooth midrib above, lateral
nerves prominulous but little conspicuous; midrib beneath prominent,
secondary nerves filiform, prominent, about 12 pairs, subpatulous,
curvate-anastomosate near margin, veins rather lax-reticulate,
prominulous.

Inflorescences cymose-paniculate, dichotomous, robust peduncle
and branches minutely pubescent. Pedicels 0.5-1.5 mm. long, thick,
angulate, puberulous or hirtellous-pubescent, articulate to short
peduncle or sessile. Bracts amplectant, persistent, triangular,
carinate, pubescent and ciliate, about 2 mm. long. Sepals 1.5 mm.
long, broadly orbicular, imbricate, puberulous at base, ciliate at

CUATRECASAS—HUMIRIACEAE 173

margin. Petals thick, linear, subacute, subsericeous, 6-7 mm. long,
1.6-2 mm. broad. Stamens 10, filaments thickened, complanate,
connate at base, 5 oppositipetalous oblong, subacute, about 3.5 mm.
long, 0.5-0.6 mm. wide, alternating with 5 longer, linear, 4-5 mm.
long, 0.3-0.4 mm. wide. Anthers with ellipsoid thecae about 0.6
mm. long, connective thick, lanceolate, compressed at upper part,
those of shorter filaments 2 mm. long, those of longer ones 1.5 mm.
long. Disk membranous, 0.8-1.5 mm. high, laciniate. Ovary
ovoid, glabrous, 5-locular, cells opposite petals and uniovulate,
about 2 mm. high. Style thick, erect, 3 mm. long. Drupe short-
ellipsoid or subglobose, 2.7-3.5 cm. long, 2.5-3 cm. broad; exocarp
fibrose-carnose, hard when dry, about 2-2.5 mm. thick; endocarp
woody, slightly 10-sulcate and bullate, with resinous cavities.

Sacoglottis gabonensis is the only species of Humiriaceae found out-
side America. It is a frequent timber tree in the tropical West
African rain forests (mostly flooded forests), throughout the Gulf of
Guinea, from Sierra Leone to northern Angola. The tree is one of
the tallest of the forests, and its timber is hard and mostly used
locally for construction. The subglobose-ellipsoid fruits have edible
exocarps and seeds; the endocarps often are found floating in the
river and on the sea shores. Aubry le Compte gave the native name
of ‘‘djouga” or ‘douga” of Gabon. Exell quoted ‘n’ooca” as a
native name at Sumba, and Exell and Mendoga quoted ‘‘n’coca” as
used in Angola. According to Emilio Guinea, the name from Spanish
Guinea is ‘‘esua.”

AFRICA: LIBERIA: Johnsonville, Dinklage 2973 (A). Vicinity of Firestone
Plantations, along Dukwai River, tree 100 ft.x4 in. much fluted and buttressed,
“daush” (cherry), Cooper 68 (A, BM, GH, NY, US, Y); tree 50 ft. 12 in. to 100
ft. straight bole, great buttressed, ‘‘daush’”’ (cherry), 1929, Cooper 274 (A, BM,
GH, NY, US, Y). Monrovia on clayish ground near the Mesurado River,
medium-size tree, flowers green, the anthers only ochre coloured, Dinklage 2973

(A).

IVORY COAST: Abidjan, “aguapo,’”’ 7-V-1929, Aubreville 92 (A). Tropical
West Africa, Mann 925, 1417 (GH).

SIERRA LEONE: Kambui, forest reserve; big timber tree, bearing fruits,
bark peeled and put in Raphia wine, ‘‘kpou-wuli,’”’ 29-X—1937, Edwarson 181
(BM). Without locality, Afzelius s.n. (BM).

GOLD COAST: Without locality, Vigne 2801 (NY). Western Province
Ankasa, Fuale; tree in wet forest, crown spreading; fruit reported edible; alt.
100 m., ‘“‘nzima,’’ I-1942, Adjimang 4849 (A). Without locality, Oigne 2800
(BM).

NIGERIA: Oban, Talbot 1744 (BM).

CAMEROONS: Bipinde, Urwaldgebiet, Zenker 148 (US); 440 (M); 1249 (A,
BM, M, 8); 1624 (BM, M, NY, 8); 1671 (NY); 1677 (BM, M, 8); 1953 (BM);
2499 (BM, M, 8S); 2760a (BM); 4407 (BM, 8).

FRENCH EQUATORIAL AFRICA: Gabon, 180 km. to the southeast of
Port Gentili; concession of Mr. Marchier on Lake Anengue, between Rivers

174 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Kouroue and Edjiwe; large tree, bole crooked, local lumbermen say this tree
rotten when old, ‘‘ozouya” (N’komi), 21-VI-1930, Krukoff 121 (NY, A, US).
Brazaville, forét de Mayombe—-Holle, 26-II-1954, Herbier I.E.C. No. 4769,
Koechlin 2632 (U).

ANGOLA: Congo Portugues, Sumba, Peco, proximum flumen, Zaire, alt. 0.0
m., 1923, Gossweiler 751 (US); Gossweiler 8707 (BM). Sumba, lower Congo
River, an erect green tree, ‘‘n’ooca,’”’ 15-V—1923, Gossweiler 8751 (BM). At
Belize- Maiombe-Congo Post, a tree 40 m. high with usually somewhat obliquely
ascending trunk, primary branches few and horizontally spreading, flowers
greenish, fruits globular the size of a walnut, highest forest trees at Belize,
Gossweiler 6996 (BM). Belize, on the Luali River, tree evergreen, 25-35 m.,
timber of a dusky mahogany color, leaves rigid, fruits the size of a walnut,
flowers collected 2 years ago from the same tree, common near the official resi-
dence, “niuca,” 138—-II-1918, Gossweiler 8182 (BM).

6. Sacoglottis guianensis Benth. in Hook Journ. Bot. Kew Misc. 5:103.
1853.—Urb. in Mart. Fl. Bras. 12(2):448 (in part), tab. 94, fig. 1, habitat.
1877.—Walp. Ann. Bot. Syst. 4:385. 1857.

Sacoglottis amazonica Benth. in Hook. Journ. Bot. Kew Mise. 5:104. 1853.

Sacoglottis guianensis fma. dolichocarpa Ducke, Arch. Jard. Bot. Rio Janeiro
3:179. 1922; 5:143, pl. 14, fig. 38. 1930.—Arch. Inst. Biol. Veget. Rio
Janeiro 4:27, 1937.

Sacoglottis guianensis var. mator Ducke, Arch. Inst. Biol. Veget. Rio Janeiro
4:27. 1938.

Type: Rob. Schomburgk 574; paratype: Rich. Schomburgk 842,
British Guiana.

Large or medium-size tree with minutely hirtellous-puberulous or
glabrous terminal branches. Leaves coriaceous, firm. Petiole 4-12
mm. long, minutely puberulous or glabrous, sulcate above. Blade
elliptic, ovate-elliptic, subovate or oblong-ovate, rounded, obtuse or
cuneate at base, narrowed, acuminate or cuspidate at apex, slightly
serrate-crenate or subentire at margin; 5-15 cm. long, 3-6 cm. broad;
above nitid, glabrous with flat midrib, lateral nerves and veins little
or not conspicuous; beneath scattered, appressed, minute hairs or
more frequently glabrous, midrib thick, eminent, secondary nerves
10-12 pairs, little ascendent more or less prominulous, near the margin
arcuate-anastomosate, minor nerves finely reticulate slightly
prominulous.

Inflorescences axillary, cymose-paniculate, shorter than flowers,
dichotomous, peduncle and branchlets shortly hirtellous-pubescent.
Bracts amplectant, ovate-triangular, ciliate, 0.5-1 mm. long, per-
sistent. Pedicels very short, (0.1-0.2 mm. long), glabrous, articulate
with 1-3 mm. long peduncles or sessile on pilose or glabrate terminal
branchlets. Sepals ovate-rotundate 6-7 mm. long, puberulous and
minutely ciliate. Petals greenish, rather thick, linear or oblong-
lanceolate, subacute, glabrous or puberulous, 3-4.5 mm. long, 1.5 mm.
broad at base. Stamens 10, filaments complanate, thickened, about
lower half connate, 2.5 and 3 mm. long, alternating. Anthers ovate-

CUATRECASAS—-HUMIRIACEAE 175

lanceolate, 1 mm. long, thecae elliptic about 0.4 mm. long, connective
thick, trigonous-lanceolate. Disk annular, membranous, denticu-
late, 0.5-0.6 mm. high. Ovary globose, glabrous, 5-locular, cells
uniovulate. Style 2-3 mm. long, erect, glabrous. Stigma subcapitate,
5-lobate. Drupe ellipsoid-oblong, attenuate at base, subacute or
subobtuse at apex, 15-30 mm. long, 9-12 mm. in diameter (most
typical 30X10 mm.); exocarp smooth compact, resinous, 1-1.5 mm.
thick; endocarp woody, smooth or very slightly bullate and furrowed,
narrow-oblong, acute at both ends, resinous-lacunose, usually 1-
seeded, rarely with 2 or 3 seeds.

S. guianensis is a widespread polymorphous species throughout
tropical South America, and includes varieties difficult to distinguish
and often mistaken for closely related species. The lack of complete
material for every collection or tree and the fact that the existing
herbarium specimens only bear flowers or fruit make it difficult to
establish correlations between flowering and fruiting characters and
the constancy of them. Through his extensive knowledge of the
Amazonian flora, Ducke was the first to point out the existence of
several varieties and forms among the supposed S. guianensis popu-
lations. Study of abundant material with special attention given
to the fruit as a taxonomic character has made it possible to consider
two new species segregated from the former broader concept of S.
guianensis. A few varieties remain in this species with somewhat
artificial characters due to the lack of fruiting material. For the same
reason the typification of the species cannot be free from error because
the Schomburgk collections are flowering specimens lacking fruit,
which in fact are key characters. Schomburgk 842, 571, and 574,
coincide so much that they seem to belong to one and the same col-
lection; they agree completely with fruiting collections (as for example
the Melinon from French Guiana) having elongated fruit; I there-
fore do not hesitate to consider the oblong fruit the characteristic trait
of S. guianensis. The typical form of this species has pubescent-
hirtellous branches and glabrous petals, but the pubescence of the
branchlets is lacking in some varieties, and the petals become more
or less puberulous in others. The length and thickness of the fruit
show some variation, but the elongate, acutish or subobtuse drupe is
the main character of the species and distinguishes it from the newly
segregated S. cydonioides and S. mattogrossensis. From the latter
one it is furthermore distinguished by its more rigid and thicker, not
conspicuously reticulate leaves.

The variety guianensis in all its forms, typical or glabrous, is
spread throughout the Amazon Basin on elevated (uninundatable)
places in rain forests or in the savanna and campo thickets varying
from large to small-size trees. It is found abundantly in the Amazon

176 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Basin in the States of Amazonas and Paré and spreads through Rio
Branco to the Guianas and Venezuela and to the Apaporis region in
Colombia. In the Guianas it is found on the slopes of the mountains
up to 1,000 m. elevation. Variety maior is almost restricted to the
Manaos region, and variety hispidula is more extended in the Ama-
zonian States of Venezuela and Brazil, it being less frequent in Surinam
and Paré.

Key to the Varieties and Forms of Sacoglottis guianensis

.

1. Petals glabrous. . . oe toe ee eee 6a. var. guianensis
2. Terminal branchlets hirtulous pubescent. . . . 6a(1). fma. guianensis
2. Terminal branchlets glabrous. .......2.2.. 6a(2). fma. glabra

1. Petals hispid-puberulous.

3. Terminal branchlets glabrous. Petioles 6-12 mm. long . 6b. var. maior
3. Terminal branchlets hirtulous-pubescent.
Petioles 4-8 mm. long. ............ Ge. var. hispidula

6a(1). Sacoglottis guianensis var. guianensis fma. guianensis
Figure 37,a-m; 38,b-d; Puatr 21

Sacoglottis guianensis fma. dolichocarpa Ducke, Arch. Jard. Bot. Rio.
Janeiro 3:179. 1922.

VENEZUELA: Amazonas: Cerro Moriche, Rfo Ventuari, tree to 10 m., fruit
green, frequent on lower slopes, alt. 200 m., 16-I-1957, Maguire, Cowan, &
Wurdack 30967 (NY, US); tree 3 m., fruit green, frequent on dry open east slopes,
alt. 300-1,000 m., 18-I-1951, Maguire, Cowan, Wurdack 30844 (US). Savanna
at Santa Barbara, Rfo Orinoco at mouth of Rio Ventuari, alt. 125 m., tree 5 m.
mature, fruit orange, 21-II-1951, Cowan & Wurdack 32022 (US). Borfvar:
Rio Apacard, Municipio Woisaan, vegetacié6n riparia, alt. 500 m., ‘‘Peru-yek,’”’
28-VITI-1954, Bernardi 1571 (NY). Regién de los rios Icaburti, Hacha y
cordillera sin nombre a 280° de las cabeceras del rio Hacha rumbo al sur, 450-850
m. alt.; selva pluvial o sabana natural, 7-I-1956, Bernardi 2813 (VEN, NY).

BRITISH GUIANA; “Guiana anglica,” X-1842, Schomburgk 842, (US,M, para-
type); photo F.M. 12599 in Berlin; Roraima, 1842-43, Schomburgk 571, (P, para-
type); 574 (P, isotype).

SURINAM: Boschreserve, Sectie O, 1—XI-1915, Boschwezen 1166 (U); Boom-
nummer 593, 21-V—1920, Boschwezen 4673 (U); 4-XI-1921, Boschwezen 5430 (U);
16-VITI-1918, Boschwezen 3961 (U); 1-VIII-1917, Boschwezen 3079 (US, U);
24-VIII-1917, Boschwezen 3125 (U, IAN). Suriname-rivier, 24—-X-~1947, Bosch-
wezen 117 (U). Boschreserve Zanderij I, Boomnummer 56, 27-VI-1919,
Boschwezen 4469 (U); 9-VII-1917, Boschwezen 2974 (US, U, IAN); IX-1942,
“doekoelia,” ‘“japopalli,” “gannasagon,” Stahel 18 (IAN. U, NY, A). In monit-
bus qui dicuntur Nassau; in bos bij K 11.2, Boom 30 m, 30 cm. dik, 22-IIT-1949,
Lanjouw & Lindeman 2869 (U). Tafelberg (Table Mountain), frequent, tree
20 m., 30 cm. diameter, flowers greenish fragrant, high mixed walaba forest, base
talus, 24-IX-1944, Maguire 24844 (M, NY).

FRENCH GUIANA: “Herbier de la Guyane, Année 1863,” Melinon s.n. (P).
“Herbier de la Guyane Francaise, Année 1862, No. 11, bois rouge tisane,”’ Melinon
584 (P). ‘‘Guiane frangaise,”’ Le Prieur 253 (P).

BRAZIL: Park: Obidos, Serra da Boa Vista, 24—-XIJ-1913, “achud,”’ Ducke
15234 (MG). Santarém, cerrado, arbusto grande, 16—-XI-1909, Huber 10446

CUATRECASAS—HUMIRIACEAE 177

ZIMMERMANN _

Ficure 37.—a-m, Sacoglottis guianensis var. guianensis, X1, a-g, fruit: a, Ducke 9868;
b, Pires 41; c, Melinon s.n.; d, Black 50-8831; e, Boschwezen 1166; f, Bernardi 1571;
g, Cowan &F Wurdack 32022. h-m, Endocarps: 4, Boschwezen 1166; 1, transection;
j, Bernardi 1571; k, transection; J, Cowan €3 Wurdack 32022; m, transection. n-?,
Sacoglottis mattogrossensis: n, Fruit (Piers 4017); 0, endocarp; 7p, transection. g-s,
Sacoglottis mattogrossensis {ma. subintegra: q, Fruit (Ducke 23820); r, endocarp; s, transec-
tion. t-u, Sacoglottis cydonioides: t, Fruit (Boschwezen 6495); u, transection.

(BMMG). Municipio de Monte Alegre, Rio Maicurd, caminho de Cdaussu a
localidade Balanga, terra firme, arvore 5 m., 16-IX—-1953, Frées 30284 (US). Rio
Tapajoz pres des cataractes du Mangabal, 31-VIII-1916, Ducke 16419 (MG, P, US).
Municipio de Faro, Fazenda Santa Olimpia, Campo Umiri, arvore, pequena,
fruto verde, 6-XI-1950, Black & Ledoux 50—-10553 (US). Lago de Faro, matta da
beira, 16-VIII-1907, Ducke 8368 (MG). Campos a E. de Faro, 27—VIII-1907,
Ducke 8524 (BM, MG). Campina rana, Alto Ariramba, 21—XII-1906, Ducke
8042 (BM, MG). Amazonas: Basin of Rio Madeira, Municipality Humayta on
plateau between Rio Livramento and Rio Ipixuna, tree 60 ft. high on campinarana
alta, 7-18-XI-1934, Krukoff 7082 (U, BM, IAN, A, US, 8, NY). Maués, varzea,
arvore pequena, 30-XI-1946, Pires 41 (IAN). Rio Negro, Preto, Matupiry, tree
35 ft. 6 in., border of river, restinga alta, 14-XI-1947, Frées 22857 (US). Rio
Branco: Caracarahy, road Boa Vista, tree 4 m., whitish flowers, low hard
growth on highland, 3-III-1948, Frées 22940 (IAN). Caminho de Samauma a

178 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

campo de aviagdo de Mucajai, kms. 14-15 da estrada Boa Vista-Caracarai,
arbusto a beira da capoeira, 25-VIII-1951, Black & Magalh&es 51-12972 (IAN).
Boa Vista-Caracarahy, perto da Colonia Fernando Costa, arvore baixinha, fruto
verde, 11-IX~-1951, Black 51-13454 (US, IAN). Gords: Porto Nacional, arvore
da matta, flores amareladas aromaticas, 27-VII-1955, Macedo 3922 (IAN, 8, US).
Without locality, Riedel s.n. (K).
6a(2). Sacoglottis guianensis var. guianensis fma. glabra Cuatr., forma nov.
Ficure 38,a

A forma guianensi typica differt ramulis terminalibus glabris.

Type in the Herbarium of the Instituto Agronomico do Norte,
Belém do Paré, Brazil, collected in the Ilha Collares, State of Par,
Brazil, December 29, 1953, by Ricardo Lemos Frées (No. 30670).

Ficure 38.—a, Sacoglottis guianensis fma. glabra, X¥%4 (Ducke 14872); b, S. guianensis fma.
guianensis, X ¥3 (Schomburgk 842); c, S. guianensis fma. guianensis, leaf, X\ (Bernardi
1571); d, S. guianensis fma. guianensis, bud and petal, X 134 (Ducke 8524); ¢, S.
guianensis var. maior, X 4 (Ducke 23818); f, S. guianensis var. mator, bud and petal,
1% (Ducke 23818); g, S. guianensis var. hispidula, bud and petal, X34 (Maguire
29337); h, S. guianensis var. hispidula, leaf, XY (Maguire 29337); i, S. mattogrossents
fma, mattogrossensis, X¥% (Ducke 2188); j, S. mattogrossensis f{ma. subintegra, X¥%
(Ducke 23820); k, S. cydonioides, X 1%.

CUATRECASAS—HUMIRIACEAE 179

COLOMBIA: Amazonas-Vaupés: Rio Apaporis, Jirijirimo, 250 m., alt., Arbol
15 m., flores verde-amarillentas, 25-26—XI-1951, Garcia Barriga 13681 (U8).

SURINAM: Boschreserve Zanderij I, Boomnummer 56, 8-XII-1951, Bosch-
wezen 1490 (UV).

FRENCH GUIANA: “Herbier de la Guyane, année 1863,’’ Melinons.n. (US, P).
Route de Cayenne au km. 7700 céte gauche de la route, en face de la Pepiniére,
“bofo-oudon” (Paramaka) ‘“mahot-cochon,” “bois cochon” (nomme commercial),
4—XII-1956, BAFOG 7622 (U).

BRAZIL: Park: Santarém, VIII-1950, Spruce 5963 (MG). Santarém,
VIII-1950, Spruce s.n. (MG, BM, 8); Spruce 763 (M); Spruce 1009 (P). ‘Fre-
quent on margin of moist forests, Santarem; I also saw much of it at Obidos,
an./50; spreading tree of 30 ft., young leaves deep red, petals whitish-green, anthers
yellow, fruits said to be very good eating ‘uaxua,’”? Spruce 1009 (Herb.
Benthamianum, K). Region des Campos de |’Ariramba (Trombetas), 30-IX-
1913, “ichud4,” Ducke 14872 (BM MG, P, US). Matta entre Cumindmirim e
Ariramba, 12-X-1913, Ducke 14967 (MG). Monte Alegre, Campo, 11—XII-1908,
Ducke 9868 (MG). Remansao, Rio Tocantins, tree 14 m., on high land, high
forest, west side, 15-IX—-1948, Frées 23494 (US). Rio Pori, afluente del Xingu, re-
gido de levantamento estatistico florestal feito pelo IAN, SPVEA e FAO, arvore 10
m.ao lado de capinarana, 30-XI-1955, Frées 32478 (US). Cambinho de Jubim
para Condeixa, Iiha do Marajé, arvore 12 m., fruto verde, 14-XI-1948, Black 48-3555
(IAN). Vigia, moita de mato em cima de un pequeno monte arvore 10 m., fruto
verde, 24-I-1950, Black 50-8831 (IAN). Collares, matta, umirizal, ‘‘achud,”
15-VIII-1913, Ducke 12656 (MG). Ilha de Collares, terra firme capoeira,
terreno argiloso, municipio de Vigia, arvore 4 m., 29-XII-1953, Frées 30670
(IAN holotype). Vigia, Campina do Palha, ilha de mata; arvore pequena,
““achira,” 21-I-50, Black 50-8685 (IAN). Jaramacani, corola branca, 27-V-57,
Egler 281 (MG, US). Amazonas: Basin Rio Solimées, Municipality Sio Paulo
de Olivenca, basin of creek Belém, tree 100 ft., trunk 2 ft. diameter, terra firma,
high forest, 26-X-11-XII-1936, Krukoff 8757 (U, NY, US, 8, P, BM).

6b. Sacoglottis guianensis var. maior Ducke, Arch. Inst. Biol. Veget. Rio
Janeiro 4:27. 1938.

FIGuRE 38,e-f

Type: Ducke 23818, Brazil, Manaos.

Petala hispidula-puberula. Ramuli glabri. Petiolus 6-12 mm.
longus. Reticulum venosum supra paulo conspicuum. Fructus
verisimiliter oblongus.

Ducke distributed under the type collection (23818) flowering
branches and loose fruits, which he described. I have seen one single
sample of this fruit (at US), and it is ellipsoid, rounded at the base,
pointed at the top, 3.5 cm. long, 2.1 cm. broad; the exocarp is about
2 mm. thick and the endocarp is hard, woody, with resinous cavities.
This fruit was not collected at the same date as the flowering speci-
mens, and I hesitate to admit that they belong together; if they did,
this collection would be a new species. I suspect that the fruit here
referred to belongs to S. macrophylla, a species much spread in the
same region (Manaos) where the Ducke specimens were collected.

BRAZIL: Amazonas: Manaos, silva non inundabili, arbor sat elata floribus
viridibus, 2-X-1932, Ducke 23818 (holotype, RB; isotypes, U, 8, US); loco arenoso

180 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

ad Cachoeira do Mindd, arbor sat magna flor. viridibus, 25-IX-1929, Ducke
23433 (US, 8). Manaos, silva terris altis ultra Flores, arbor sat magna floribus
viridibus, 14-IX-1945, Ducke 1756 (MG, NY, IAN, A, US). Breves, in estuario
amazonico circa Pard; silva non inundata, arbor magna, floribus viridibus,
fructibus globosis, Ducke 17784 (U). Cajatuba, habitat low and high land,
“achud,” 18-I-1932, Montes da Costa 281 (IAN, P). Maranuio: Grajaht, Rio
Mearim, 180 m. alt., arvore alta, flor amarella, Arojado Lisboa 2330 (BM, MG).
PaRA: Peixeboi (Belém-Braganga), ‘‘Parard,’’ 16-VII-1907, Siqueira 8281
(BM, MG, US).

6c. Sacoglottis guianensis var. hispidula Cuatr., var. nov. FIGuRE 38,9-h

Petala extus plus minusve hispidulo-puberula. Ramusculi termi-
nales pubescenti-hirtuli. Petioli 4-8 mm. longi. Fructus ignotus
verisimiliter oblongus.

Type in the U.S. National Herbarium, No. 1660793, collected near
Livramento on Rio Livramento, firm land, Municipality of Humayta,
at Rio Madera Basin, State of Amazonas, Brazil, November 1934, by
B. Krukoff (No. 6653). Isotypes in NY, A, IAN, BM, S, U.

The collections included in this variety show much polymorphism
in the leaf-shape within the broad range of variation of S. guianensis.
The Surinam collection, Maguire 24836, is a form with scanty hairs
on the petals and more rigid and more venose leaves. On the other
hand, Maguire 29337 from the Orinoco is another extreme form with
more hispidulous petals and small, rigid, almost enervate leaves.
Maguire, Cowan, & Wurdack 30844 and 30967 and Froes 22587 are
named as forma guianensis, but they have very few hairs on the petals;
they range in the very intermediate forms.

VENEZUELA: Amazonas: Rfo Orinoco, Culebra savanna, north base of
Cerro Duida, alt. 200 m., medium-size tree, flowers white, occasional at the edges
of savanna, 13-X-1950, Maguire, Cowan, & Wurdack 29416(VEN). Rfo Orinoco
Rio Atabapo, Cafio Temi, 1 hour below Yavita, alt. 125 m., edge of small laja,
medium-size tree with greenish flowers, 20-X-1950, Maguire 29337 (US). Botf-
vaR: occasional along river 1-4 km. above Salto de Humito (25-30 km. from
mouth), tree 9 m., flowers green with yellow anthers, 7-I-1956, Wurdack &
Monachino 41149 (US). Along Rfo Karuai at base of Sororopén-tepu{, west of
La Laja; alt. 1,220 m.; tree 4 ft. tall; leaves subcoriaceous, rich green and very
shining above, pale green below; “pert-yek,” 29-XI-1944, Steyermark 60756
(NY, VEN).

SURINAM: Coppenam River headwaters, Wallaba forest, km. 9, line between
Camps No. 5 and 4, frequent; tree 15 m., 20 em.; flowers greenish, fragant,
23-IX-1944, Maguire 24836 (GH, NY, US, U, VEN).

BRAZIL: Amazonas, Rio Urubt, Cachoeira Iracena, terra firme, alta; floresta
central; arvore 18 m., flores brancas, 22-IX-1949, Frées 25369 (US). Campinha
Rio Breves, IX-1913, Kuhlmann 3510; tree 40 ft., green-whitish flowers, border
of river of whitish water, on low land, high forest, 11-X-1947, Frées 22587 (IAN).
Basin Rfo Madeira, Humayta near Livramento, Krukoff 6653 (type).

CUATRECASAS—HUMIRIACEAE 181

7. Sacoglottis mattogrossensis Malme, Arkiv. Bot. Stockh. 22A, No 7:9.
1928. Figures 37,n-p

Sacoglottis guianensis forma. sphaerocarpa Ducke, Arch. Jard. Bot. Rio
Janeiro 3:178. 1922; 5: pl. 14 figs. 39a-b. 1930.

Type: Malme I1:2237, Brazil, Mato Grosso, Santa Ana da Chapada.

Small or medium-size tree with lenticellate and hirtellous or
glabrous branchlets. Leaves coriaceous, flexible, glabrous or sub-
glabrous. Petiole 5-8 mm. long, semiterete, puberulous or glabrate,
thickened at base. Blade oblong-elliptic or elliptic-lanceolate,
ovate-elliptic, rounded, obtuse or cuneate at base, acutely acuminate
or cuspidate at apex, obtusely serrate at margin or subentire; 5-11 cm.
long, 2-2.5 cm. broad, glabrous or with sparse hairs on midrib be-
neath; midrib prominulous above, prominent beneath, secondary
nerves 8-10 on each side, subascendent, thin and prominent, minor
nerves and veins forming sublax reticulum prominently conspicuous
on both sides. Sometimes, chiefly on young branches, blades bi-
glandular at base.

Inflorescences axillary, small, cymose-paniculate, dichotomous,
peduncle 2-5 cm. long, stout, striate, hirtellous, the branchlets
short, hispidulous. Bracts persistent, amplectant, ovate-triangular,
acutish, minutely puberulous, ciliate, 1-0.6 mm. long. Pedicels
thick, 0.7 mm. long, glabrous. Sepals ovate, rounded at apex,
thick, about 0.6 mm. long, glabrous except for ciliate margin. Petals
linear, narrowed toward apex, subacute, glabrous, about 4 mm. long,
1-1.2 mm. broad at base, the estivation cochlear. Stamens 10,
elabrous, filaments complanate, 2.5 and 3.2 mm. long alternating,
lower part united in tube. Anthers ovate, 0.7-0.9 mm. long, thecae
oblong, connective thick, acute or subacute. Disk about 0.4 mm.
high, annular, rather thick, dentate. Ovary ovoid, glabrous, 0.8 mm.
high. Style 2-3 mm. long. Stigma short-capitate, 5-lobate. Drupe
globose, 17-28 mm. diameter, exocarp compact, resinous, 1-2 mm.
thick, coriaceous and granulose when dry; endocarp woody, almost
smooth, slightly bullate and sulcate, innerly resinous-lacunose, usually
monospermous or with 2 (rarely 3) seeds.

This species has been confused with S. guianensis, from which
Ducke first separated it as a variety based on its globose fruit form.
It furthermore differs from S. guianensis in its thinner, flexible, and
prominently nerved leaves; for this reason I promoted forma sphaero-
carpa Ducke to the rank of a species (type: Kuhlman 2128). After
examining the type of S. mattogrossensis Malme (flowering material),
I realized that the Kuhlmann and Malme plants are conspecific.

S. mattogrossensis is rather a species of the lower Amazon Basin,

182 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

extending from Maranhéo southward to Rio de Janeiro; variety
subintegra goes further west to the Manaos region and is found in
Colombia at the northern end of the rain forest domain.

The species is rather polymorphic, and the following key outlines
the most conspicuous varieties and forms.

Key to the Varieties and Forms of Sacoglottis mattogrossensis

1. Petals glabrous. ........4.. 2... Ta. var. mattogrossensis
2. Branchlets hirtulo-puberulous. . .. . . Ta(1). fma. mattogrossensis
2. Branchletsglabrous. ......4. 4.4.4... . . 7a(2). fma, glabra

1. Petals puberulous. .......4..2. 2... . Tb. var. subintegra
3. Branchlets glabrous. .......... . Wb(l). fma. subintegra
3. Branchlets hirto-puberulous. ...... .. . 7b(2). fma. puberula

7a(1). Sacoglottis mattogrossensis var. mattogrossensis fma. mattogrossensis
FicuReE 38,i; PLATE 22

BRAZIL: Par: Santarém, ilha de matta no campo, ‘achud,” 17-VII-1916,
Ducke 16346 (MG). Obidos, matta, 8-III-1909, arvore pequena, Ducke 10218
(MG); matta de terra firme, 23-IX-1910, Ducke 11050 (MQ); in terreno arenoso,
“achud,” 10-VIII-1916, Ducke 16320 (US, BM, MG). Rio Xingu, em frente
Seuzel, mun. Porto de Méz; regido onde foi feito um levantamento estatistico
florestal pelo IAN, SPVEA e FAO; arvore de 20 m. alt., terra firme, flanco do
planalto, margem esquerda do rio, 18-XI-1955, Frées 32391 (IAN). Monte
Alegre, arbor parva 26-ITI-1928, Kuhlmann 2128 (US, 8, U, isotypes of fma.,
sphaerocarpa Ducke). Soure, Capoeira densa, arvore pequena, fruto verde, polpa
(verde) muito adstringente, amarela, 26-II-1950, Black 50-9058 (US). Serra do
Cachimbo, 425 m., 12-XII-1956, Pires, Black, Wurdack, & Nilo 6140 (IAN).
Alto Tapajés, Vila Nova, perto da Cachoecira do Chacorao, terra firme, campo;
arvore pequena, frutos esfericos amarelos, 24—I-1952, Pires 4017 (IAN, US).
Marannao: Séo Luiz, Granja Barreto, arvore pequena, 26-VI-1949, Murca
Pires 1510 (IAN, NY); high land, tree 4 m. yellow fruit, 14-V—1949, Frées 24297
(IAN). Sao Luiz, Granja Federal, tree 4 m., white flowers, 12-V-1949, Frées
24256 (IAN). Granja Barreto, viveiros de aves, arvore pequena, flor verde,
registro n. 32, “‘parurd,’” 29-X-1948, Ducke 2188 (IAN). Anil, Capueira,
“uachua,” 12-IX-1903, Ducke 363 (US, BM, MG). Rio Branco: Rio Canta,
arbusto ou arbore, ‘‘achua,” 8—X—-1951, Black 51-13843 (US). Marro Grosso:
Santa Anna da Chapada, arbor parva habitu C. salicifolia, in capueira, 10-VIII-
1902, Malme 2237 (S, holotype; 8, isotype, sterile). Rio pr JANEIRO: Rio de
Janeiro ad urbem loco Gavea, VITI-1916, Frazdo 8118 (US), in silvulis siccioribus
prope Porto Estrella, arbor parva fructibus aurantiacis, 23-VIII-1925, Ducke &
Kuhlmann 19165 (US).

7a(2). Sacoglottis mattogrossensis var. mattogrossensis fma. glabra Cuatr.,
fma. nov.

Ramuli terminales glabri.

Type in the U.S. National Herbarium, No. 1040536, collected near
Faro, State of Pardé, Brazil, May 11, 1911, by Adolfo Ducke (No.
11653). Isotypes at BM and MG.

COLOMBIA: Vaupfs: Riberas del Rfo Infrida alrededores de Morichal, cerca

de la boca del rio Papanaua, 200 m. alt., drbol 15 m., frutos color verde, 9-II-1953,
Fernandez 2228 (COL, US).

CUATRECASAS—HUMIRIACEAE 183

BRAZIL: Par&: Soure, ilha do Marajo, Capoeira do campo con caembé;
arvore pequena, fruto verde, inflorescencias velhas, 12-X-1948, Black 48-3453
(IAN); entre os km. 1 e 7 estremo Monte Alegre a CANP; campos cerrados
(“cobertos”), 5-V-1953, Lima 53-1332 (IAN). Belém on lands of IAN, 3 km.
east of Administration Building, near Fazenda Velha, medium-size tree, flowers
green, fruits red, 21-I-1944, Antonio Silva 59 (IAN, US). Faro, “achua,” 11-V-
1911, Ducke 11653 (type US, BM, MG), photo F.M. 35178 from Paris. Portel,
matta da terra firme, arvore de porte mediano, casca castanha, 17-X—-1955,
Williams & Silva 18201 (US). Serra de Arumandube, Almeirim, matta da
Chapade “achud ” 26-VIII-1918, Ducke 17262. Sitio Cagote, Areias-Recife-Pe,
6-X-1949, arvore pequena, flores brancacentas, 6-X-1949, Lima 49-336 (IAN).
Monte Alegre, campo, arvore grande, ‘“achud,” 11-XII-1908, Ducke 9866 (MG).
Belém, Hosp. Dom. Fr., II-05, Huber 6992 (MG). EFB, Santa Izabel, Carapara,
Capoeira, arbusto, flor amarela, 27-XII-1908, Museu Goeldi 10180 (BM, MQ).
PrernamBuco: Prazeres, 13-X-1932, Pickel 591 (GH, NY, US). Ilha Itamaraca,
XII-1857, Gardner 1146 (BM). Iguarassu, 1-X-1887, Ramage s.n. (BM).
Maranuao: Anil, Capoeira, 3-VI-1907, Ducke 519 (MG). So Pauxo: Santos,
in ripa arenosa sicca aprica maris, 18-X-1875, Mosen 3477 (8).

Tb. Sacoglottis mattogrossensis var. subintegra (Ducke) Cuatr., comb. nov.
Sacoglottis guianensis var. subintegra Ducke, Arch. Inst. Biol. Veget. Rio
Janeiro 4:27. 1938.

Tb(1). Sacoglottis mattogrossensis var. subintegra fma. subintegra
Figures 36,h-n; 37,q-s; 38,7

Petla puberula. Ramuli terminales glabri. Fructus globosus
18-28 mm. diamitens sublaevis, exocarpio coriaceo-resinoso 1-2 mm.
crasso, endocarpio leviter undulato, 1.7-2.4 cm. diamitenti, lacunoso-
resinoso monospermo.

Type: Ducke 23820, Brazil, Amazonas, Manaos.

BRAZIL: Amazonas: Manaos, silva terris altis ultra coloniam Jodo Alfredo,
arbor magna floribus viridibus odoratis fructus rubro-aurantiacis, “achua,”
2-VIII-1937, Ducke 23820 (RB, holotype; U, P, 8, US, isotypes); silva primaria
terris altis argillosis, arbor magna viridibus, 23-VII-1943, “ex arbore typica”’
flowers, Ducke 1295 (A, NY, IAN, US, MG). Monte Alegre, regido da Colonia

da Mulata, terra firme, matta virgen, arvore, flor branca, 28-IX~-1950, Frées
30416 (US).

Tb(2). Sacoglottis mattogrossensis var. subintegra fma. puberula Cuatr., fma.
nov. PLATE 23

Ramuli terminales hirto-puberuli.

Type in the Herbarium of Museu Goeldi at Belém do Par4, Brazil,
collected at Canuta, beira da Campina, State of Pard, Brazil, July 21,
1916, by Adolpho Ducke (No. 16286). Isotype in the Herbarium of
British Museum.

BRAZIL: Para: Canuta, Ducke 16286 (MG, holotype; BM, isotype).

8. Sacoglottis cydonioides Cuatr., sp. nov. Figure 38,k; Puate 24

Arbor media vel grandis ramis terminalibus brunneis subteretibus
plus minusve lenticellatis glabris.

184 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Folia crasse vel crassiuscule coriacea rigida glabra. Lamina ovato-
elliptica, ovato-oblonga, oblongo-elliptica vel subovato-lanceolata
apice subite acuminata vel cuspidata, basi rotundata vel obtusa et
abrupte in petiolum robustum subalatum basi incrassatum 3-6 mm.
longum cuneato-angustata, margine plus minusve revoluta obtuse
serrata vel subintegra; 6-13 cm. longa 2.5-6 cm. lata; supra nitida
saepe laevis costa crassiuscule prominula nervis secundariis reticuloque
inmersis obsoletis vel plus minusve conspicuis; subtus costa valde
eminenti nervis secundariis 10-12 utroque latere tenuibus prominulis-
que patulis marginem versus arcuato-anastomosantibus venulis minute
reticulatis bene prominulis vel subinmersis sed conspicuis.

Inflorescentiae axillares et terminales cymoso-paniculatae dichotomo-
ramosae foliis valde breviores pedunculo 0.5-1.5 cm. longo robusto
striato plus minusve complanato puberulo, ramis brevibus angulatis
minute hispidulo-puberulis. Bracteae persistentes amplectentes
ovatae acutiusculae parce ciliatae 1.3-0.5 mm. longae. Pedicelli
crassi 0.4—0.5 mm. longi glabri. Sepala ovata obtusa crassiuscula
extus nitida glabraque margine bene ciliata, 0.6—-0.7 mm. longa.
Petala aestivatione quincuncialia oblonga sursum attenuata subacuta
2.8-3 mm. longa 1 mm. lata dimidia superiore parte minute hispidula.
Stamina 10 glabra filamentis complanatis basi coalitis, parce papillosis
longioribus 1.8-2 mm. longis brevioribus circa 1.4 mm. longis. An-
therae 1 mm. longae, thecis oblongis infra lateralibus 0.4 mm. longis,
connectivo crasse ovato sursum longe acuteque complanato-cuspidato.
Discus 0.5 mm. altus squamis crassiusculis denticulatis coalitis.
Ovarium glabrum ovoideum 5-loculare, in loculis ovule singulo.
Stylus circa 0.5-0.7 mm. longus. Stigma capitatum 5—lobatum.
Drupa sphaeroidea 15-20 mm. diamitens exocarpio laevi vel leviter
granuloso 3-5 mm. crasso duplo, exteriori coriaceo compacto (in sicco)
granuloso-resinoso, interno dense fibroso; endocarpio lignoso resinoso-
lacunoso plus minusve tuberculato, 8-12 mm. diamitenti, 1-3 spermo.
Epicarpium odore Cydoniae.

Type in the Herbarium of the Botanical Museum, Utrecht, Nether-
lands, collected in the Boschreserve Brownsberg in Surinam, June 28,
1924, by Reis (No. 64). Paratypes (flowering specimens) were
collected in the Boschreserve Watramiri, Surinam, Boomnummer
1606, collected by the Forest Service, Boschwezen (No. 4720). in
Botanical Museum, Utrecht, and in Instituto Agronomico do N orte,
Belém do Paré.

Medium-size or large tree with brownish, lenticellate and glabrous
terminal branchlets. Leaves thick-coriaceous, rigid, glabrous. Blade
ovate-elliptic, ovate-oblong, oblong-elliptic or subobovate-lanceolate,
suddenly acuminate or cuspidate at apex, rounded or obtuse at base
and abruptly and shortly tapering into a broad petiole, 3-6 mm. long,

CUATRECASAS—HUMIRIACEAE 185

very thickened at base; 6-13 cm. long, 2.5-6 cm. broad; above lustrous,
usually smooth, with prominulous and broad midrib, immersed
secondary nerves and veins obsolete or more or less conspicuous;
midrib very prominent beneath, 10-12 pairs of spreading secondary
nerves thin, prominulous, near margin arcuate and anastomosing,
reticulum minute and prominulous or sometimes immersed but con-
spicuous.

Inflorescences axillary and terminal, cymose-paniculate, dicho-
tomous, shorter than leaves, peduncle 0.5-1.5 cm. long, stout, striate,
more or less complanate, puberulous, branchlets short, angulate,
minutely hispid-puberulous. Bracts amplectant, persistent, ovate,
subacute, sparsely ciliate, 1.3-0.5 mm. long. Pedicels thick, glabrous,
0.4-0.56 mm. long. Sepals ovate, obtuse, rather thick, glabrous
except for ciliate margin, 0.6-0.7 mm. long. Petals oblong, attenuate
toward apex, subacute, 2.8-3 mm. long, 1 mm. broad, minutely
hispidulous on upper half. Stamens 10, glabrous, filaments com-
planate, united at base, sparsely papillose, longer ones (1.8-2 mm.)
alternating with 5 shorter (1.4 mm.). Anthers 1 mm. long, thecae
narrowly oblong on lower sides, 0.4 mm. long, connective thick,
ovate, compressed and acutely cuspidate at apex. Disk 0.5 mm.
high, scales thick, denticulate, united. Ovary glabrous, ovoid, 5-
locular, cells uniovulate. Style about 0.5-0.7 mm. long. Stigma
capitate, 5-lobate. Drupe globose 15-20 mm. diameter, exocarp
almost smooth or somewhat granular, 3-5 mm. thick and double,
outer layer coriaceous, compact when dry, resinous-granular, inner
layer densely fibrous; endocarp woody, resinous-lacunose, more or
less tuberculate, 8-12 mm. diameter, 1-3 seeds. Epicarp with strong
scent of Cydonia fruits.

S. cydonioides has been mistaken for S. guianensis, and it is difficult
to distinguish in sterile specimens. But the fruit of S. cydonioides
are very different; they have a double exocarp, the inner layer of which
is fibrous and difficult to dissociate from the endocarp. The latter
is more or less tuberculate; the hispid-puberulous petals also differ-
entiate the two species.

S. cydonioides is a medium-size tree from the uninundatable
rain forest ranging from Surinam to French Guiana and northeastern
Brazil. Westward it spreads into the British and Venezuelan Gui-
anas.

VENEZUELA: Botfvar: El Dorado, 80-90 km. al sur, en selvas pluviales;
Arbol 20 m., drupas rojo-ladrillo, perfumadas, 15 mm., hojas discoloras; especie es-
casa, “trompillo,’’ 30-III-1956, Bernardi 3033 (VEN).

BRITISH GUIANA: Cuyuni River, Akarabice Creek in mixed forest, tree
80 ft. high and 15 in. diameter, flowers green, “Duhuria,’”’ 28-VII-1933 (dry
flowers), Tutin 421 (US, BM, U, paratype, flowers).

513359—61—12

186 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

SURINAM: Boschreserve Watramiri, 21-VI-1920, Boschwezen 4720 (para-
type, flowers US, IAN, U); 19-III-1919 Boschwezen 4296 (U); 8-X-1918, Bosch-
wezen 4038 (US) (U); 6-V-1916, Boschwezen 1936 (U). Boschreserve Brownsberg,
28-VI-1924, Boschwezen 6495 (holotype, U). In montibus qui dicuntur Nassau;
in bos bij km. 0.6 Boom 28 inch., 30 em. dik; vrucht oranje, ‘“‘Kwatta sirie”’
(Sur.), “soort buffelhout” (S-D.), Lanjouw & Lindeman 2194 (U). Awarraballi,
Sectie O, XI-1944, Stahel 263 (A, NY, U).

FRENCH GUIANA: Crique Serpent Rive gauche & 800 m. de son embouchure;
terrain marecageuse-sablonneuse; grains (fruit) de la grosseur d’un pois de couleur
vert devenant brundtre & maturité; “bofo-oudou” (Paramaka), 20-VII-1953,
BAFOG 35M (P). Chantier Fosima & 1 km. au Sud de la sicri Margot, terrain
plat et sain; fruits ronds 2-3 cm. jaun-orange 4 maturité ayant un odeur de pomme;
“bofo-oudou”” (Paramaka), ‘“mahot-cochon” (nom commercial), 7—-XII-1953,
BAFOG 124M (P). Route de St. Laurent a Cayenne km. 18 coté droit et 4
20 m. de la route s/terrain sain; fruits verts ronds 10-20 mm., pulpe 2-3 mm.;
noyou trés dur, odeur non définie; “boliquin’”’ (Pamaka) “gris-gris rouge” (nom
commercial) ; les rodins fendus en lamelles de 3 & 5 mm. d’épesseur sur 10-12 cm.
de largeur donnent les ‘‘gaulettes’’ qui servent & cloisonner les “carlets” habita-
tions des établissements en foréts; 10-XII-1953, BAFOG 131M (P). St. Laurent,
II-1956, “bofoudou”’ (Paramaka), BAFOG 347 (U). Route de Cayenne au km.
14.100 coté gauche et a 20 m. de la route; fruits jaundtres, globuleux, 1.5-2 cm.,
odeur de pomme, groupes en grappes axillaires; ‘‘bofo-oudou” (Paramaka),
“mahot cochon” ou “bois cochon’” (noms commerciaux), 29-I-1957, BAFOG
7656 (U). ‘Herbier de la Guyane” Année 1863, Melinon s.n. (P). Cayenne,
Martin s.n. (P, K).

BRAZIL: Par: Trombetas, Rio Amind, matta da terra firme a liste do
Lago Salado, 22-IV-1917, Ducke 16809 (MG). Amapd, Rio Oiapoque, beira do
rio, terra firme, alta; arvore de 6 m., fruto vermelho-salmon, 1—-II-1950, Frées
25783 (IAN). Amapé, Rio Oiapoque; terra firme, alta floresta alta arvore 15 m.,
15-X-1950, Frées 26636 (IAN).

Collections Cited

ApraHaM, A. A.

152 Humiria balsamifera var. flori-
bunda

ApDJIMANG, E. O.
4849 Sacoglottis gabonensis
Arze.ius, A.
s.n. Sacoglottis gabonensis
AtuLEMAO, F., & Cysnerros, F.

255 Humiria balsamifera var. flori-

bunda
ALLEN, P. H.
5812 Humiriastrum diguense subsp.
costaricense

6415 Vantanea barbourii
6681 Vantanea barbourii
Auston, A, H. G., & Lutz, B.
169 Humiria balsamifera var. parvi-
folia
Aurson, R. A.
545 Humiria balsamifera var. cori-
acea
ANDERSON, C. W.

154 Humiriastrum obovatum
506 Humiria balsamifera var. gui-
anensis
559 Humiria balsamifera var. gui-
anensis
ANDERSON, M. J.

s.0o. Humiriastrum dentatum

Appwun, C. F.
37 Humiria balsamifera fma. atte-
nuata
Araqun, J., & BARKLEY, F.

18Va021 Humiria balsamifera var.
subsessilis

Aravso, J. M. P.

6 Sacoglottis amazonica

ArcHer, W. A.

7915 Sacoglottis amazonica
7964 Sacoglottis amazonica

ARISTEGUIETA, L.

2174 Humiria balsamifera var. cori-
acea

AvuBLET, J. B. C. F.

sn. Vantanea guianensis

AUBREVILLE, A.

92 Sacoglottis gabonensis

BAFOG (Bureau AGr. ET FORESTIER
GUYANAIS)

35M _ Sacoglottis cydonioides
48M Vantanea parviflora var. parvi-
flora

Vantanea parviflora var. par-
viflora

Sacoglottis cydonioides

102M

124M

131M _ Sacoglottis cydonioides

220M Vantanea parviflora var. par-
viflora

228M MHunmiria balsamifera fma. bal-
samifera

247M Vantanea parviflora var. par-
viflora

347M _ Sacoglottis cydonioides

1083 Humiria balsamifera fma. bal-
samifera

7587 Humiria balsamifera fma. bal-
samifera

7622 Sacoglottis guianensis fma.
glabra

7656 Sacoglottis cydonioides
Baitey, I. W.

115 Humiria balsamifera fma. bal-
samifera

187

188 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Baxerr, B. O.

s.n. Endopleura uchi
58 Endopleura uchi
9401 Endopleura uchi

BaLpwin, J.

3187 Schistostemon oblongifolium

BaRBOSA DA Sitva, M.

73 Humiria balsamifera fma. atte-
nuata

Barsour, W. R.

1018 Vantanea barbourii

Becca, N.

s.o. Humiria balsamifera var. gui-
anensis

Bena, P.

1319 Humiriastrum excelsum

Benorst, R.

1239 Humiria balsamifera fma. bal-
samifera
1530 Vantanea guianensis

Brrnarpl, A. L.

1571 Sacoglottis guianensis fma.

guianensis

2601 Humiria balsamifera fma. at-
tenuata

2603 Humiria balsamifera var. cori-
acea

2613 Vantanea minor

2813 Sacoglottis guianensis fma,
guianensis

2814 Humiriastrum obovatum

3033 Sacoglottis cydonioides

38820 Vantanea minor

Buack, G. A.

47-1001 Endopleura uchi

47-1276 WHumiria balsamifera fma.
balsamifera

47-1756 Humiria balsamifera var.
floribunda

48-2512 Humiria balsamifera var.
guianensis

48-2514 Schistostemon retusum
48-2589 Humiria balsamifera var.
guianensis

48-3249 Humiria balsamifera fma.
attenuata

48-3453 Sacoglottis mattogrossensis
fma. glabra

48-3555 Sacoglottis guianensis fma.
glabra

49-8313 Humiria balsamifera fma.
balsamifera

49-8369 Humiria balsamifera fma.
balsamifera

50-8685 Sacoglottis guianensis fma.
glabra

50-8831 Sacoglottis guianensis fma.
glabra

50-9058 Sacoglottis mattogrossensis
fma. mattogrossensis

51-12776 Humiria balsamifera fma.
attenuata

51-13231 Humiria balsamifera fma.
balsamifera

51-13454 Sacoglottis guianensis fma.
guianensis

51-13843 Sacoglottis mattogrossensis
fma. mattogrossensis

55-18577 Humiria balsamifera fma.
balsamifera

57-19306 Endopleura uchi

Buack, G. A., & Foster, M. B.
48-3393 Sacoglotis amazonica

Buack, G. A., & Lepoux, P

50-10371 UHumiria balsamifera var.
floribunda

50-10553 Sacoglottis guianensis fma.
guianensis

50-10783 Humiriastrum cuspidatum
var. cuspidatum

Buack, G, A., & Macauyizs, D.

51-11790 Humiria balsamifera var.
parvifolia

5951-12954 Humiria balsamifera fma.
balsamifera

51-12972 Sacoglottis guianensis fma,
guianensis

Buack, G.A., Eater, W., CAVALCANTE

P., & Sinva, A.
57-19590 Humiria balsamifera var.
floribunda

CUATRECASAS—HUMIRIACEAE

Buack, G. A., Frézs, R. L., &
Lxepoux, P.

50-9810 Sacoglottis amazonica
50-9811 Humiriastrum excelsum

BLANCcHET, J. S.

s.n. Humiria balsamifera var. parvi-
folia

85 Vantanea compacta var. compacta

1005 Humiria balsamifera var. parvi-

folia

2810 Humiria balsamifera var. parvi-
folia

3144A Humiria floribunda var. par-
vifolia

3305 Vantanea compacta var. com-
pacta

3362 Vantanea compacta var. com-
pacta

3422 Humiria balsamifera var. par-
vifolia

3570 Humiria balsamifera var. par-
vifolia

3805 Vantanea compacta var. com-
pacta

3837 Vantanea compacta var. com-
pacta

Boupinau, I.

3886 Humiria balsamifera var. guia-
nensis

BoscHWEZEN, B. W., SURINAM FOREST

SERVICE.

36A Humiria balsamifera var. gui-
anensis

117 Sacoglottis guianensis fma. gui-
anensis

177 Humiria balsamifera fma. at-
tenuata

212 WHumiria balsamifera fma. bal-
samifera

531 Humiria balsamifera fma. gui-
anensis

1120 Humiria balsamifera var. gui-
anensis

1166 Sacoglottis guianensis fma.
guianensis

1490 Sacoglottis guianensis fma. glabra

1547 Humiria balsamifera fma. bal-
samifera

1935 Humiria balsamifera fma. bal-

samifera

1936
2068
2232
2471
2599
2765
2816
2885
2918
2974
3010
3040
3079
3125
3646
3934
3947
3961
4038
4296
4469
4669
4673

4684

4720
4770

4810
4933

4960
5412

189

Sacoglottis cydonioides
Schistostemon densiflorum
Humiria balsamifera fma, bal-
samifera

Humiria balsamifera fma. bal-
samifera

Humiria balsamifera var. gui-
anensis

Humiria balsamifera fma. bal-
samifera

Humiria balsamifera var. gui-
anensis

Humiria balsamifera fma. bal-
samifera

Humiria balsamifera fma. bal-
samifera

Sacoglottis guianensis fma. gui-
anensis

Humiria balsamifera fma. bal-
samifera

Humiria balsamifera var. gui-
anensis

Sacoglottis guianensis fma. gui-
anensis

Sacoglottis guianensis fma. gui-
anensis

Humiria balsamifera var. gui-
anensis

Humiria balsamifera fma. bal-
samifera

Humiria balsamifera var. guia-
nensis

Sacoglottis guianensis fma. guia-
nensis

Sacoglottis cydonioides
Sacoglottis cydonioides
Sacoglottis guianensis fma. guia-
nensis

Humiria balsamifera fma. bal-
samifera

Sacoglottis guianensis fma. guia-
nensis

Humiria balsamifera fma. bal]-
samifera

Sacoglottis cydonioides
Humiria balsamifera fma. at-
tenuata

Humiria balsamifera fma.
samifera

Schistostemon densiflorum
Schistostemon densiflorum
Humiria balsamifera fma. bal-
samifera

bal-

8475

1518

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Sacoglottis guianensis fma. guia-
nensis
Humiria balsamifera var. guia-
nensis

Humiria balsamifera fma. bal-
samifera
Humiria balsamifera fma. at-
tenuata

Humiria balsamifera fma. bal-
samifera

Humiria balsamifera fma. bal-
samifera

Humiria balsamifera fma. bal-
samifera

Humiria balsamifera fma.
samifera

Sacoglottis cydonioides
Humiria balsamifera fma.
samifera

Humiria balsamifera fma bal-
samifera

bal-

bal-

Britton, N. L.

Sacoglottis amazonica

Broapway, W. E.

Sacoglottis amazonica

Bucnutien, O.

Humiriastrum mapiriense

Buroos, J. A.

37 Humiriastrum excelsum
85 Humiriastrum excelsum

430
483

774

965

1112

1768

1823

1869

Caprucno, P.

Endopleura uchi
Vantanea parviflora var. parvi-
flora

Carpona, F.

Humiria balsamifera var. cori-
acea
Humiria balsamifera var. guai-
quinimana

Humiria balsamifera var. guai-
quinimana

Humiria balsamifera var. cori-
acea

Humiria balsamifera var. cori-
acea

Humiria balsamifera var. cori-
acea

1912 Vantanea minor

2269 MHumiria balsamifera var. cori-
acea

2362 Vantanea minor

2533 Humiria balsamifera var. cori-
acea

2670 Humiria balsamifera var. cori-
acea

2877 Humiria balsamifera var. cori-
acea

CASARETTO, G.
s.n. Humiriastrum dentatum

CHAGAS

3093 Schistostemon macrophyllum

Cuacas & Dionisio

38472 Schistostemon macrophyllum

Cooper, G. P.

68 Sacoglottis gabonensis
274 Sacoglottis gabonensis

Cowan, R. 8.

38700 Humiria balsamifera fma.
balsamifera

39263 Humiria balsamifera var. gui-
anensis

39266 Humiria balsamifera var. gui-
anensis

Cowan, R.8., & Maauire, B.

38034 Humiria balsamifera var. flori-
bunda

Cowan, R.8., & Wurpack, J. J.

31090 Humiria balsamifera var. co-

riacea

31301 Humiria balsamifera var. co-
riacea

31472 Humiria balsamifera var. sub-
sessilis

31502 Humiria balsamifera var. sub-
sessilis

32022 Sacoglottis guianensis fma gui-
anensis

CUATRECASAS, J.
7203 Schistostemon retusum
14418 Humiriastrum diguense var.

anchicayanum
14956 Humiriastrum diguense

CUATRECASAS—HUMIRIACEAE

16615 Humiriastrum procerum
17186 Humiriastrum procerum
17226 Sacoglottis ovicarpa
19727 Humiria balsamifera var. sub-
sessilis
19909 Humiriastrum melanocarpum
19927 Sacoglottis ovicarpa
19937 Vantanea occidentalis
19989 Humiriastrum melanocarpum
19998 Sacoglottis ovicarpa
Curran, H. M.
159 Humiria balsamifera var. parvi-
folia

Dayton, W. A., & Barsour, W. R.

3004 Sacoglottis amazonica
3129 Vantanea barbourii

De La Cruz, J.S.

2202 Humiria balsamifera var. gui-
anensis

2210 Humiria balsamifera var. gui-
anensis

2227 Humiria balsamifera var. gui-
anensis

2644 Humiria balsamifera var. gui-
anensis

2645 Humiria balsamifera var. gui-
anensis

Dinxuaap, M. J.
2973 Sacoglottis gabonensis
Donant HERB.

1686 Humiria balsamifera var. flori-

bunda
Ducks, A.

s.n. Sacoglottis amazonica

s.n. Humiriastrum cuspidatum var.
cuspidatum

s.n. Endopleura uchi

12 Sacoglottis ceratocarpa
16 Sacoglottis ceratocarpa
16a Sacoglottis ceratocarpa

87 Humiria balsamifera var. flori-
bunda

157 Vantanea parviflora var. parvi-
flora

200 Vantanea guianensis
241 Endopleura uchi

243

255
265
305
363

416
440

519

541

751
752
781

1055
1174
1175
1295

1301
1513

1614
1647
1723
1744
1756

2108
2188

2230
7174
7213
8029
8042
8368

8410

191

Humiriastrum cuspidatum var.
glabriflorum

Schistostemon macrophyllum

Hylocarpa heterocarpa

Endopleura uchi

Sacoglottis mattogrossensis fma.
mattogrossensis

Vantanea macrocarpa

Humiria balsamifera var. flori-
bunda

Sacoglottis mattogrossensis fma.
glabra

Humiria balsamifera var. flori-
bunda

Vantanea micrantha

Vantanea paraensis

Vantanea parviflora var. parvi-
flora
Sacoglottis amazonica
Sacoglottis ceratocarpa
Schistostemon macrophyllum
Sacoglottis mattogrossensis var,
subintegra fma. subintegra
Sacoglottis ceratocarpa
Vantanea compacta var. com-
pacta
Humiriastrum excelsum
Vantanea guianensis
Sacoglottis amazonica
Schistostemon macrophyilum
Sacoglottis guianensis var.
maior
Duckesia verrucosa
Sacoglottis mattogrossensis fma.
mattogrossensis
Vantanea macrocarpa
Schistostemon macrophyllum
Humiria balsamifera fma. at-
tenuata
Humiria balsamifera var. flori-
bunda

Sacoglottis guianensis fma. gui-
anensis

Sacoglottis guianensis fma.
guianensis

Humiria balsamifera var. gui-
anensis

Sacoglottis guianensis fma.
guianensis

Humiriastrum cuspidatum var.
cuspidotum

Humiria balsamifera var. gui-
anensis

192
9866
9868
10218

10815
11050

11550
11653

11790

12030
12656

14872
14962
14967
14979
14992
15234
15415
15451
15459
15467
15514
15515
16286
16320

16325
16346

16419

16578
16641
16764
16809
17221
17262

17779

CONTRIBUTIONS FROM THE

Sacoglottis mattogrossensis fma.
glabra

Sacoglottis guianensis fma.
glabra

Sacoglottis mattogrossensis fma.
mattogrossensis

Duckesia verrucosa

Sacoglottis mattogrossensis fma.
mattogrossensis

Schistostemon macrophyllum

Sacoglottis mattogrossensis fma.
glabra

Humiriastrum cuspidatum var.
cuspidatum

Humiriastrum villosum
Sacoglottis guianensis fma.
glabra

Sacoglattis guianensis fma.
glabra

Vantanea parviflora var. parvi-
flora

Sacoglottis guianensis fma.
glabra

Endopleura uchi

Duckesia verrucosa

Sacoglottis guianensis fma.
guianensis

Vantanea guianensis

Vantanea guianensis

Humiriastrum excelsum

Vantanea parviflora var. parvi-
flora

Humiria balsamifera fma. bal-
samifera

Humiria balsamifera fma. bal-
samifera

Sacoglottis mattogrossensis var.
subintegra fma. puberula

Sacoglottis mattogrossensis fma.
mattogrossensis

Duckesia verrucosa

Sacoglottis mattogrossensis fma.

mattogrossensis
Sacoglottis guianensis fma.
guianensis

Sacoglottis amazonica
Endopleura uchi

Duckesia verrucosa
Sacoglottis cydonioides
Sacoglottis amazonica
Sacoglottis mattogrossensis
fma. glabra

Endopleura uchi

NATIONAL HERBARIUM

17780
17781
17782
17783
17784

19166

20427
20428

21024
21357
23424
23425
23426
23427
23428
23429
23430
23431
23432
23433
23434
23436
23814
23815
23816
23817
23818
23819
23820
30126
30128
30131

30133
30134

Humiriastrum excelsum

Sacoglottis amazonica

Vantanea paraensis

Vantanea guianensis

Sacoglottis guianensis var.

maior

Humiriastrum = glaziovii
glaziovii

Vantanea parviflora var. par-
viflora

Vantanea macrocarpa

Vantanea parviflora var. parvi-
flora

Schistostemon macrophyllum

Vantanea macrocarpa

Hunmiria balsamifera fma. bal-
samifera

Vantanea parviflora var. pu-
berulifolia

Vantanea parviflora var. par-
viflora

Vantanea parviflora var.

puberulifolia

Vantanea parviflora var.

puberulifolia

Vantanea parviflora var. parvi-
flora

var.

Vantanea paraensis
Sacoglottis ceratocarpa
Schistostemon macrophyllum
Sacoglottis guianensis var.
maior
Humiriastrum cuspidatum
var. cuspidatum
Humiriastrum cuspidatum
var. glabriflorum
Vantanea guianensis
Endopleura uchi
Schistostemon macrophyllum
Schistostemon oblongifolium
Sacoglottis guianensis var.
maior
Schistostemon reticulatum
subsp. reticulatum
Sacoglottis mattogrossensis
var. subintegra fma. subintegra
Humiriastrum cuspidatum var,
cuspidatum
Humiria balsamifera fma.
balsamifera
Schistostemon retusum
Vantanea macrocarpa
Vantanea tuberculata

CUATRECASAS—HUMIRIACEAE

30135 Vantanea micrantha
30137 Hylocarpa heterocarpa
Ducks, A., & Kugimann, J. G.

19165 Sacoglottis mattogrossensis
fma, mattogrossensis
Epwarpson, I. E.

181 Sacoglottis gabonensis

Eater, W. A.

261 Humiria balsamifera var. flori-

bunda
281 Sacoglottis guianensis fma. glabra

FaNsHAVE, D. B.

F715 Humiria balsamifera fma. bal-
samifera

FERNANDEZ, A.

2084 Humiria balsamifera var. lau-

rina

2142 Schistostemon retusum

2148 Humiria balsamifera var. gui-
anensis

2228 Sacoglottis mattogrossensis fma.
glabra

Focke, H. C.

1018 Humiria balsamifera var. gui-
anensis

1286 Humiria balsamifera var. gui-
anensis

Forest DeparTMENT, Britisu GUIANA

154 Humiriastrum obovatum
404A Schistostemon densiflorum

506 Humiria balsamifera var. gui-
anensis

559 WHumiria balsamifera var. gui-
anensis

600 Vantanea guianensis

663 Humira balsamifera var. laurina

931 Humiriastrum obovatum

2055 Humiria balsamifera var. gui-
anensis

Frazio, A.

8118 Sacoglottis mattogrossensis fma.
mattogrossensis

11813

19933
20480

20803
21090
21192
21338
21342
21346
21370
21411
21437
22472
22587
22644
22703
22738

22747
22760

22838
22842
22857
22940
23494
24256
24297
24820

24916
24924

24934
24936
25185

193

Fréss, R. L.

Humiria balsamifera var. flori-
bunda
Vantanea obovata

Humiria balsamifera var.
floribunda
Humiriastrum cuspidatum

var. glabriflorum

Schistostemon oblongifolium

Sacoglottis ceratocarpa

Humiria balsamifera var. gui-
anensis

Humiria balsamifera var. gui-
anensis

Humiria balsamifera var. flori-
bunda

Schistostemon
subsp. froesii

Schistostemon retusum

Schistostemon oblongifolium

Schistostemon macrophyllon

Sacoglottis guianensis var.
hispidula

Humiriastrum villosum

Schistostemon macrophyllum

Humiria balsamifera fma. at-
tenuata

Schistostemon retusum

Humiria balsamifera var. gui-
anensis

Humiria balsamifera var. gui-
anensis

Humiria balsamifera var. gui-
anensis

reticulatum

Sacoglottis guianensis fma.
guianensis

Sacoglottis guianensis fma.
guianensis

Sacoglottis guianensis fma.
glabra

Sacoglottis mattogrossensis
fma. mattogrossensis
Sacoglottis | mattogrossensis
fma. mattogrossensis
Humiriastrum cuspidatum

var, subhirtellum
Schistostemon macrophyllum
Humiriastrum piraparanense

Sacoglottis ceratocarpa
Vantanea guianensis

Vantanea parviflora var. par-
viflora

194 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

25369 Sacoglottis guianensis var.
hispidula

25438 Humiriastrum cuspidatum
var. cuspidatum

25459 Vantanea parviflora var. par-
viflora

25463 Humiriastrum cuspidatum

var. subhirtellum
25480 Humiriastrum cuspidatum var.
subhirtellum

25565 Endopleura uchi

25783 Sacoglottis cydonioides

26071 Schistostemon macrophyllum

26428 Vantanea paraensis

26636 Sacoglottis cydonioides

26812 Humiria balsamifera var. flori-
bunda

27985 Humiriastrum piraparanense

28407 Humiriastrum piraparanense

28454 Humiria balsamifera fma. at-
tenuata

28895 Humiria balsamifera fma. at-
tenuata

29854 Humiria balsamifera fma. bal-
samifera

30093 Humiria balsamifera var. flori-
bunda

30284 Sacoglottis guianensis fma. gui-
anensis

30416 Sacoglottis |§mattogrossensis
fma, subintegra

30670 Sacoglottis guianensis fma. gla-
bra

32391 Sacoglottis mattogrossensis

fma. mattogrossensis
32478 Sacoglottis guianensis fma.
glabra

Frées, R. L., & Apprson, G.

29096 Humiria balsamifera var. gui-
anensis

Humiria balsamifera var. gui-
anensis
29119 Schistostemon macrophyllum
29144 Hunmiriastrum villosum
29211 Humiria balsamifera var. gui-

anensis

29102

Frées, R. L., & Brack, G. A.

27572 WHumiria balsamifera var. flori-

bunda

Froées, R. L., & Finuo, J. P.

29486 Humiria balsamifera fma. bal-
samifera

Garcia Barriaa, H.

13681 Sacoglottis
glabra

14287 Humiriastrum piraparanense

guianensis fma.

GARDNER, C. A.

1146 Sacoglottis mattogrossensis fma.
glabra

1263 WHumiria balsamifera var. flori-
bunda

4452 Vantanea obovata

4452 bis Humiria balsamifera var. mi-
narum

Guaziou, A. F. M.

s.n. Humiria balsamifera var. parvi-

folia

63 Humiria balsamifera var. parvi-
folia

731 Humiria balsamifera var. parvi-
folia

6196 Humiria balsamifera var. parvi-
folia

7765 Humiria balsamifera var. parvi-
folia

8286 Humiria balsamifera var. parvi-
folia

10078 Vantanea guianensis

10342 Humiria balsamifera var. par-
vifolia

10437 Humiria balsamifera var. flori-
bunda

11828 Vantanea compacta var. com-
pacta

11829 Vantanea compacta var. com-
pacta

12515 Humiria floribunda var. parvi-
folia

14640 Vantanea compacta var. gran-
diflora

16723 Vantanea compacta var. gran-
diflora

16724 Humiriastrum glaziovii var.
angustifolium

18178 Humiriastrum dentatum

18179 Humiriastrum glaziovii var.

glaziovii

CUATRECASAS—HUMIRIACEAE 195

18180 Humiria balsamifera var. par-
vifolia

18181 Vantanea compacta var. gran-
diflora

18182 Vantanea compacta var. com-
pacta

18962 Humiria balsamifera var. par-
vifolia

18963 Vantanea obovata

18964 Humiriastrum glaziovii var.
glaziovii

Guerason, H. A.

729 Hunmiriastrum obovatum

Gomgs, A. I.

8.0. Humiria balsamifera var. parvi-
folia

GOSSWEILER, J.

751 Sacoglottis gabonensis

6996 Sacoglottis gabonensis
8182 Sacoglottis gabonensis
8707 Sacoglottis gabonensis
8751 Sacoglottis gabonensis

GuEDEs, M.
1260 Endopleura uchi
GuepEs, T,

58 Sacoglottis ceratocarpa
80 Sacoglottis ceratocarpa

GuILpING, L.
s.n. Sacoglottis amazonica
GUILLEMIN, A.

205 Humiria balsamifera var. parvi-
folia

Guppy, N.
308 Sacoglottis amazonica
Hart, J. H.
s.n. Sacoglottis amazonica
Hitcucocr, A. E.

16938 Humiria
guianensis

balsamifera var.

Horune, F. C.

3021 Humiriastrum glaziovii var.
angustifolium

7970 Humiria balsamifera var. parvi-
folia

29281 Vantanea compacta var. com-
pacta

Hostman, W. R.

793 Humiria balsamifera var. gui-
anensis

HoutTMONSTER

541A Humiria balsamifera fma. bal-
samifera

542A Humiria balsamifera fma. bal-
samifera

543A Humiria balsamifera fma. bal-
samifera

Huser, H.

96 Humiria balsamifera var. flori-
bunda

239 Endopleura uchi

940 Endopleura uchi

1260 Endopleura uchi

1850 Sacoglottis amazonica

2785 Humiria balsamifera fma, bal-
samifera

6992 Sacoglottis mattogrossensis fma.
glabra

9583 Vantanea parviflora var. parvi-
flora

10446 Sacoglottis guianensisfma. gui-
anensis

Humsert, H.

27422 WHumiria balsamifera var. sub-
sessilis

27440 Humiria balsamifera var. gui-
anensis

Humsenrt, H., & Scuuttss, R. E.

27363 Humiriastrum villosum
27364 Humiria balsamifera fma. sub-
sessilis

INPA (Instituto NaAcIONAL DE
PrEsquisas DA AMAZONICA, MANAOs)

86 Humiria balsamifera var. flori-
bunda

204 Humiriastrum cuspidatum var.
glabriflorum

244 Schistostemon macrophyllum

620 Humiria balsamifera var. cori-
acea

196
1056
1243
1407
1638
1684
1700
1773
1809
1814

2044
2084

246

287
299

478
1023

1281

2489
3912

4719
4883

5561
5562
5672

8.n.

362

5525

8.0,
2144

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Humiria balsamifera var. flori-
bunda

Schistostemon macrophyllum
Schistostemon macrophyllum
Sacoglottis ceratocarpa
Humiria balsamifera var. gui-
anensis

Humiriastrum cuspidatum var.
glabriflorum

Humiria balsamifera var. flori-
bunda

Vantanea parviflora var. parvi-
flora

Sacoglottis ceratocarpa
Schistostemon macrophyllum
Sacoglottis ceratocarpa

Irwin, H. 8.

Humiria balsamifera fma. at-
tenuata

JENMAN, G. 5S.

Schistostemon densiflorum

Humiria balsamifera var. gui-
anensis

Schistostemon densiflorum
Humiria balsamifera var. cori-
acea

Humiria balsamifera var. gui-

anensis

Schistostemon densiflorum
Humiria balsamifera fma. bal-
samifera

Schistostemon densiflorum
Humiria balsamifera var.
anensis

Humiria balsamifera var. gui-
anensis

Humiria balsamifera var.
anensis

Humiria balsamifera fma. bal-
samifera

Humiriastrum obovatum

gui-

gui-

JoBert, Dr.
Vantanea guianensis
JunxeEr, N. W.

Humiria balsamifera fma.
samifera

bal-

KappuEr, A.

Schistostemon dichotomum
Schistostemon dichotomum

Kiuup, E. P., & Surrg, A. C.

28681

37b

1091
1130
1315
1564
2846

3706

2632
121

1483
4956
6371
6506
6653
7082
7120

7182
7926

7928

8757

11270

2128

2894

3509

Humiria balsamifera fma. at-
tenuata

Kuern, R.

Vantanea compacta var. com-
pacta

Kuva, G.

Vantanea peruviana

Vantanea peruviana

Humiria balsamifera fma. at-
tenuata
Schistostemonreticulatum
subsp. reticulatum

Humiria balsamifera fma. at-
tenuata

Humiria balsamifera var. parvi-
folia

KorEcuHLIn, J.

Sacoglottis gabonensis

Krukorr, B.

Sacoglottis gabonensis
Humiria —_ balsamifera
laurina
Vantanea parviflora var. parvi-
flora
Vantanea celativenia
Sacoglottis amazonica
Sacoglottis guianensis var. his-
pidula
Sacoglottis guianensis fma. gui-
anensis
Vantanea parviflora var. parvi-
flora
Vantanea celativenia
Humiria balsamifera var. flori-
bunda
Humiria balsamifera var. flori-
bunda
Sacoglottis guianensis fma. gla-
bra

Humiriastrum mapiriense

var.

KuuuMan, J. G.

Sacoglottis mattogrossensis
fma. mattogrossensis

Humiria balsamifera var.
guianensis

Humiria balsamifera var.
floribunda

CUATRECASAS—HUMIRIACEAE 197

3510 Sacoglottis guianensis var. his-
pidula
21029 Schistostemon macrophyllum

Kuyrrer, J.

33 Humiria balsamifera var. gui-
anensis

568 Humiria balsamifera var. gui-
anensis

Lamp, F. B.

133 Vantanea magdalenensis

141 Humiriastrum colombianum
145 Humiriastrum colombianum
170 Humiriastrum colombianum

Lansouw, J.

195 Humiria balsamifera var, gui-

anensis

334 Humiria balsamifera var. gui-
anensis

1253 Humiria balsamifera fma, bal-
samifera

Lansouw, J., & Linpeman, J. C,

H8 MHumiria balsamifera fma. bal-

samifera

267 Humiria balsamifera var. gui-
anensis

268 Humiria balsamifera var. gui-
anensis

573 Humiria balsamifera fma. bal-
samifera

652 Humiria balsamifera fma. bal-
samifera

911 Humiria balsamifera var. gui-
anensis

968 Humiria balsamifera var. flori-
bunda

1797 Humiria balsamifera var. gui-
anensis

1798 Humiria balsamifera var. gui-
anensis

2194 Sacoglottis cydonioides
2869 Sacoglottis guianensis fma, gui-

anensis

3259 Humiria balsamifera var. gui-
anensis

3289 Humiria balsamifera var. gui-
anensis

3317 Humiria balsamifera var. gui-
nensis

Lasser, T., & Varescut, V.

3888 Humiria balsamifera var

coriacea
LEBLOND

402 Humiria balsamifera var. flori-
bunda

441 Humiria balsamifera var. flori-
bunda

Le Prieur, M.

253 Sacoglottis guianensis fma. gui-

anensis

1838 Humiria balsamifera fma. bal-
samifera

1840 Humiria balsamifera fma. bal-
samifera

Lima, D, DE

1623 Humiria balsamifera var parvi-

folia

49-336 Sacoglottis mattogrossensis
fma. glabra

53-1273 Humiria balsamifera fma,
attenuata

53-1332 Sacoglottis mattogrossensis
fma. glabra

LINDEMAN, J. C.

258 Humiria balsamifera var. gui-

anensis

4201 Humiria balsamifera var. flori-
bunda

4202 Humiria balsamifera fma, bal-
samifera

4381 Humiria balsamifera var. gui-
anensis

6541 Humiria balsamifera fma. bal-
samifera

6861 Humiria balsamifera fma. bal-
samifera

6862 Humiria balsamifera var. flori-
bunda

6880 Humiria balsamifera fma. bal-
samifera

6881 Humiria balsamifera var. gui-
anensis

6882 Humiria balsamifera var. gui-
anensis

198

6883 Humiria balsamifera var. gui-

anensis
Lispoa, A.

2327 Humiria balsamifera var. flori-
bunda

2327 Humiria balsamifera var. lau-
rina

2330 Sacoglottis guianensis var.
maior

4099 Humiria balsamifera var. flori-
bunda

Lirt.e, E. L.
6233 Humiriastrum procerum

6320
6412
6413

Humiriastrum procerum
Humiriastrum procerum
Humiriastrum procerum

LUETZELBURG, P, v.

40 Vantanea obovata

22561 Humiria balsamifera var, gui-
anensis

22575 Wumiria balsamifera var. gui-
anensis

22627 Humiria balsamifera var. gui-
anensis

24014 Humiria balsamifera var. sub-
sessilis

Lunt, W.

s.n. Sacoglottis amazonica
5984 Sacoglottis amazonica

Lutz, B.

681 Humiria balsamifera var. parvi-
folia

Macepo, A.

3922 Sacoglottis guianensis fma. gui

anensis

4034 Humiria balsmifera fma,
attenuata

Macurirg, B.

24223 Humiria balsamifera var.
coriacea

24443 Humiria balsamifera var.
coriacea

24707 Humiria balsamifera var.
coriacea

24789 Humiria balsamifera var.
coriacea

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

24836 Sacoglottis guianensis
hispidula

24844 Sacoglottis guianensis
guianensis

29337 Sacoglottis guianensis
hispidula

32686 Humiria balsamifera
coriacea

32763 Humiria _ balsamifera
guaiquinimana

33099 Humiria balsamifera
guaiquinimana

33242 Humiria balsamifera
iluana

33388 Humiria balsamifera
iluana

Macurre, B., & Fansuawe, D.

23233 Humiria crassifolia

23295 Humiria balsamifera
coriacea

23450 Humiria balsamifera
guianensis

32158 Humiria balsamifera
imbaimadaiensis

var.
fma,
var.
var.
var.
var.
var.

var.

B.

var.
var.

var.

Macuire, B., & Maaeuire, C.

35040 Humiria balsamifera
floribunda
35140 Humiria balsamifera
coriacea
35453 Humiria balsamifera
coriacea
40105 WHumiria balsamifera
stenocarpa
40159 Humiria balsamifera
coriacea
Macuire, B., & Port,
27627 Humiria balsamifera
coriacea
27695 Humiria balsamifera
coriacea
27974 Humiria balsamifera
floribunda
28828 Humiria balsamifera
guianensis
Macuire, B., & STAHEt,
23654 Humiria balsamifera
guianensis
23696 Humiria balsamifera

guianensis

var,

var.

var.

var.

var.

CUATRECASAS—HUMIRIACEAE 199

24957 Humiria balsamifera fma.

attenuata

Macurire, B., & Wurpack, J. J.

34677 Humiria
laurina

35579 Humiria
laurina

Maaurre, B., Cowan, R. 8., &
Wurpack, J. J.

balsamifera var.

balsamifera var.

29416 Sacoglottis guianensis var. his-

pidula

29416A Humiria balsamifera’ var.
subsessilis

29541 Humiria balsamifera var.
coriacea

29697 Humiria balsamifera var.
coriacea

29769 Humiria balsamifera var.
subsessilis

30018 Humiria balsamifera var.
coriacea

30483 Humiria fruticosa

30543 Humiria balsamifera var.
guianensis

30561 Humiria fruticosa

30622 Humiria balsamifera var.
coriacea

30693 Sacoglottis maguirei

30791 Humiria balsamifera var.
guianensis

30844 Sacoglottis guianensis fma.
guianensis

30885 Humiria balsamifera var.
coriacea

30918 Humiria balsamifera var.
coriacea

30967 Sacoglottis guianensis fma.
guianensis

30987 Humiria balsamifera var.
floribunda

Maguire, B., Wurpack, J. J., &

Buntina, G.
35882 Humiria balsamifera var.
stenocarpa
36210 Humiria balsamifera var.
subsessilis
36295 Humiria balsamifera var.
guianensis

36354 Humiria balsamifera var.

laurina

36456 Humiria balsamifera var.

guianensis

36580 Humiria fruticosa

37632 Humiria balsamifera var.
laurina

Maauire, B., Wurpack, J. J., &

Keita, W.

41821 Humiria balsamifera var,
laurina

41917 Humiria balsamifera var.
guianensis

Macurre, B., Wurpack, J. J., &
Macuirp, C.

41640 Vantanea guianensis

Matmg, G. D.

2237 Sacoglottis mattogrossensis
fma. mattogrossensis

Mann, G.

925 Sacoglottis gabonensis
1417 Sacoglottis gabonensis

Martin, J.

s.n. Humiria balsamifera fma. balsa-
mifera

s.n. Humiriastrum subcrenatum

s.n. Sacoglottis cydonioides

Martius, C. E. P.

s.n. Humiria balsamifera fma. atte-
nuata

s.0. Humiria balsamifera var flori-
bunda

s.n. Humiria crassifolia

s.no. Humiriastrum cuspidatum var.
cuspidatum

s.n. Sacoglottis amazonica

s.0. Vantanea obovata

Martyn, E. B.

136 Humiria balsamifera var. gui-
anensis

ME.utnon, M.

s.n. Humiria balsamifera fma. bal-
samifera

sn. Sacoglottis cydonioides

s.n. Sacoglottis guianensis fma. gla-
bra

200

s.n. Sacoglottis guianensis fma. gui-

anensis

s.n. Vantanea parviflora var. parvi-
flora

48 Humiria balsamifera fma. bal-
samifera

100 Vantanea guianensis (US)

100 Vantanea parviflora var. parvi-
flora (P, BM)

377 Humiria balsamifera fma. bal-
samifera

584 Sacoglottis guianensis fma. gui-
anensis

ME .to Fino, L. E.

1186 Humiria balsamifera var. parvi-
folia

MenpEes MAGALHAES
2117 Vantanea obovata

{urKpER, C. A., Scnottss, J. A., &

Dayton, W. A.
3041 Sacoglottis amazonica
Mex, Y.
5815 Humiria balsamifera var. mi-
narum
6049 Vantanea guianensis
Miers, J.
6167 Vantanea parviflora var. par-
viflora
8915 Humiria balsamifera var. parvi-
folia

Montes pa Costa
281 Sacoglottis guianensis var. maior

Mosg&n, H.
3475 Hunmiriastrum dentatum

38477 Sacoglottis mattogrossensis
fma. glabra
Moss, M.
13. Humiria balsamifera fma. balsa-
mifera
57 Humiria balsamifera fma. balsa-
mifera

MusEv GoELpDI
1260 Endopleura uchi
9419 Humiria balsamifera fma. bal-
samifera

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

9583 Vantanea parviflora var. par-
viflora

9664 Vantanea guianensis

9670 Vantanea parviflora var. par-
viflora

9672 Humiriastrum excelsum

9680 Vantanea parviflora var. par-

viflora
9723 Vantanea parviflora var. par-
viflora
10130 Sacoglottis mattogrossensis
fma. glabra
Orang, C.

2800 Sacoglottis gabonensis
Patifo, V. M.

12 Vantanea occidentalis

Pearce, R.

s.n. WVantanea compacta subsp. mi-
crocarpa

Persaup, A. C.

102 Schistostemon densiflorum

191 Humiria balsamifera fma. atte-
nuata

288 Schistostemon densiflorum

Puetps, K., & Hitcucock, C.

508 Humiria balsamifera var. cori-
acea

Picket, D. B,

591 Sacoglottis mattogrossensis fma,
glabra

Pires, J. Murea
s.n. Vantanea parviflora var. par-
viflora

41 Sacoglottis guianensis fma. guia-
nensis

588 Schistostemon oblongifolium
708 Hylocarpa heterocarpa
754 Humiria balsamifera var. guia-

nensis

989 Humiria balsamifera fma. bal-
samifera

1029 Humiria balsamifera var. sub-
sessilis

1030 Humiriastrum piraparanense

CUATRECASAS—HUMIRIACEAE 201

1510 Sacoglottismattogrossensis fma.

mattogrossensis

3877 Humiria balsamifera — var.
laurina

4017 Sacoglottis mattogrossensis
fma. mattogrossensis

4518 Vantanea guianensis

Pires, J. M., & Buack, G. A.
31 Sacoglottis amazonica
2961 Humiria balsamifera var. par-
vifolia
Pires, J. M., & Siiva, A.

4192 Humiria balsamifera var. flori-

bunda

4624 Humiria balsamiferafma. at-
tenuata

4629 Humiria balsamifera fma at-
tenuata

4702 Humiria balsamifera var. flori-
bunda

Pires, J. M., Buack, G. A., WurDAcK,

J. J., & Nito

6140 Sacoglottis mattogressensis
fma, mattogrossensis

Pires, J. M., Buack, G. A., WuRDACK,
J. J., & Sitva, A.

6209 Humiria
laurina

Pires, J. M., Fréus, R. L., & Siva, A.

4954 Vantanea parviflora var. par-
viflora

5105 Vantanea parviflora var. par-

viflora

Vantanea guianensis

balsamifera var.

5380
Pirrrer, H.

16260 Sacoglottis ovicarpa
Porpria, E.

s.n. Humiria balsamifera var. flori-

bunda
18 Humiria balsamifera var. flori-
bunda
3011 Humiria balsamifera var. flori-
bunda
Pout, Dr.

s.n. Humiria balsamifera var. par-
vifolia
513359—61—_—_13

Porrgay, A.

s.n. Humiria balsamifera fma. at-
tenuata

Putz, A.

52 Humiria, balsamifera var. guia-
nensis

150 Humiria balsamifera var. guia-
nensis

RaMAGE, G. A,

s.n. Sacoglottis mattogrossensis fma.
glabra

Ramos, G., & PatiXo, V. M.
s.n. Vantanea occidentalis
Recu, Dr.
1862 Schistostemon dichotomum
Reitz, P. R.

Vantanea compacta var. com-
pacta

3353

Reitz, P. R., & Kier, R.

1589 Vantanea compacta var. com-

pacta

1730 Vantanea compacta var. com-
pacta

1744 Vantanea compacta var. com-
pacta

1836 Vantanea compacta var. com-
pacta

Ricwarp, L. C.

s.n. Humiria balsamifera fma. bal-
samifera

s.n. Humiria balsamifera var. flori-
bunda

s.n. JHHumiria balsamifera var. parvi-
folia

s.n. Humiriastrum dentatum
s.n. Wantanea parviflora var. parvi-
flora

RIEDEL, L.

s.n. Humiria balsamifera var. flori-
bunda

s.n. Humiria balsamifera var. mi-
narum

s.n. Humiria balsamifera var. parvi-
folia

202 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

s.n. Humiriastrum = glaziovii var.

glaziovii

s.n. Sacoglottis guianensis fma. gui-
anensis

3570 Humiria balsamifera var. parvi-
folia

Romsouto, H. E.
228 Humiria balsamifera fma. at-
tenuata
RomeERo CastaNepa, R.
4785 Humiriastrum colombianum
4942 Humiriastrum colombianum
Rupesr, E.

s.n. Humiria balsamifera fma. bal-
samifera

Sacot, P. A.
s.n. Humiria balsamifera fma. bal-
samifera
Sr. Hivarre, A. DE

s.n. Humiria balsamifera var. parvi-
folia

114/5 Humiria balsamifera var. par-
vifolia

1705 Vantanea obovata

1984 bis Vantanea obovata

SALZMAN, LP.
s.n. Humiria balsamifera var. flori-
bunda
SaMuELs, J. A.
s.n. THumiria balsamifera var. gui-
anensis
SANDEMAN, CHRISTOPHER
2199 Humiria balsamifera var. gui-
anensis
Sanpwitu, N. Y.

374 Schistostemon densiflorum
399 Humiria balsamifera fma. bal-
samifera

ScHOMBURGE, R.

47 Vantanea guianensis
135 Humiriastrum obovatum
166 Humiriastrum obovatum

270 Humiria balsamifera var. gui-
anensis

346 Wumiria balsamifera var. laurina

543 Schistostemon densiflorum

560 Humiria balsamifera var. laurina

571 Sacoglottis guianensis fma. gui-

anensis

574 Sacoglottis guianensis fma. gui-
anensis

576 Humiria balsamifera var. savan-
narum

584 Humiriastrum obovatum

628 Humiria balsamifera var. laurina

825 Humiriastrum obovatum

842 Sacoglottis guianensis fma. gui-
anensis

845 Humiria balsamifera var. savan-
narum

968 Humiria balsamifera var. laurina

982 Vantanea guianensis

1359 Humiriastrum obovatum

1552 Vantanea minor

1581 Vantanea guianensis

s.n. Jumiria balsamifera var. laurina

Scuuttes, R. E.

943la Humiria balsamifera fma. bal-

samifera

9435 Humiria balsamifera fma. balsa-
mifera

23131 Humiriastrum cuspidatum var.
cuspidatum

Scnuttes, R. E., & Caprera, I.

15054 Wumiria crassifolia

15511 Humiria balsamifera fma. at-
tenuata

15922 Humiriastrum piraparanense

16893 Humiria balsamifera fma.
attenuata

17045 Sacoglottis ceratocarpa

17231 Humiria balsamifera var. sub-
sessilis

17253 Sacoglottis ceratocarpa

18319 Humiria balsamifera var. sub-
sessilis

18371 Hunmiria balsamifera var. sub-
sessilis

19290a Sacoglottis ceratocarpa

19519 Sacoglottis ceratocarpa

19951 Humiria balsamifera var.
laurina

CUATRECASAS—HUMIRIACEAE 203

19963 Humiria
laurina

Scuuttss, R. E., & Lépez, F.
8881 Schistostemon macrophyllum

balsamifera var.

9267 Vantanea parviflora var. par-
viflora

9363 Humiria balsamifera var.
guianensis

9510 Humiria balsamifera var. sub-
sessilis

9701 Humiriastrum cuspidatum var.
cuspidatum

10339 Humiria balsamifera var. flori-
bunda

Scuuttses, R. E., & Pires, J. M.

9103A Hunmiria balsamifera var. sub-
sessilis

SeGADAS-VIANNA, F.

3506 Humiria balsamifera var. parvi-

folia

3634 Humiria balsamifera var. parvi-
folia

3635 Humiria balsamifera var. parvi-
folia

Sreaapas-Vianna, F., Dau, L.,
Ormonp, W. T., MAcHLINE,
G. C., & Lorepo, L.

I-310 and I-369 Humiria balsamifera
var. parvifolia
Humiria balsamifera var. par-
vifolia
Humiria balsamifera var. par-
vifolia
Humiria balsamifera var. par-
vifolia
Humiria balsamifera var. par-
vifolia
I-945 Humiria balsamifera var. par-
vifolia
I-1383 Humiria balsamifera var. par-
vifolia
I-1416 Hunmiria balsamifera var. par-
vifolia

I-385
I—439
I-821

I-907

SELLow, F.

s.n. Humiria balsamifera var. flori-
bunda

s.n. Humiria balsamifera var. par-
vifolia

171 Humiria balsamifera var. flori-
bunda

180 Humiria balsamifera var. par-
vifolia

2212 Wumiria balsamifera var. par-
vifolia

2228 Humiria balsamifera var. par-
vifolia

Sriva, A,

59 Sacoglottis mattogrossensis fma.
glabra

Sintva, J. F.
416 Duckesia verrucosa
Srquerra, R.

8281 Sacoglottis guianensis var.

major
8775 Vantanea guianensis
SuitH, A. C.
2176 UHumiria balsamifera var. gui-.
anensis
2423 Humiria balsamifera fma. at-
tenuata
Smity, L. B.
6406 Humiria balsamifera var. parvi-
folia
6694 Humiria balsamifera var. parvi-
folia

SPLITGERBER, F. L.

s.n. Humiria balsamifera var. gui-

anensis
Spruce, R.
s.n. Humiria balsamifera var. flori-
bunda
s.n. Humiria balsamifera var. gui-
anensis

s.n. Sacoglottis guianensis fma. glabra

sn. Schistostemon oblongifolium

164 Humiria balsamifera var. flori-
bunda

181 Humiria balsamifera var. flori-
bunda

763 Sacoglottis guianensis fma. glabra

928 Humiria balsamifera var. flori-
bunda

1009 Sacoglottis guianensis fma. gla-
bra

204

1499 Humiria balsamifera var. flori-
bunda

1714 Schistostemon macrophyllum

1715 Humiriastrum cuspidatum var.
cuspidatum

1915 Humiriastrum cuspidatum var.
cuspidatum

1969 Schistostemon oblongifolium

2419 Schistostemon oblongifolium

2424 Humiriastrum cuspidatum var.
cuspidatum

2443 Humiriastrum cuspidatum var.
cuspidatum

2454 Humiria balsamifera var. sub-
sessilis

2457 Humiria balsamifera var. sub-
sessilis

3073 Schistostemon oblongifolium

3094 Schistostemon oblongifolium

3194 Schistostemon oblongifolium

3409 Humiria balsamifera var. gui-
anensis

3419 Humiria balsamifera var.
laurina

4335 Humiria balsamifera var.
parvifolia

5963 Sacoglottis guianensis fma. gla-
bra

STAHEL, G.

18 Sacoglottis guianensis fma. gui-
anensis

90 Humiria balsamifera fma. atte-
nuata

263 Sacoglottis cydonioides

STanEL, G., & Goneanryp, J. W.

3570 Humiria balsamifera fma. bal-
samifera

STEYERMARK, J. A,

57817 UHumiria balsamifera var. lau-
rina

57880 Humiria balsamifera var. sub-
sessilis

58288 Humiria balsamifera var. cori-
acea,

59186 Humiria balsamifera var. cori-
acea

59621 Humiria balsamifera var. pi-
losa

60192 WHumiria balsamifera var. sub-
sessilis

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

60289 Humiria balsamifera var. pi-
losa

60756 Sacoglottis guianensis var. his-
pidula

STEYERMARK, J. A., & Wurpack, J. J.

1109 Humiria balsamifera var cori-
acea

TaupBot, P, A.
1744 Sacoglottis gabonensis
Tamayo, F.
3123 Vantanea minor
Tate, G. H. H.

50 Humiria balsamifera fma. atte-
nuata

142 Schistostemon oblongifolium

209 Humiria balsamifera var. lau-

rina

283 Humiria balsamifera var. lau-
rina

286 Humiria balsamifera var. lau-
rina

330 Humiria balsamifera var. savan-
narum (2nd sheet)

330 Humiria balsamifera var. sub-
sessilis (1st sheet)

331 Humiria balsamifera var. sub-

sessilis
733 Humiria balsamifera var. cori-
acea
1113 Humiria balsamifera var. cori-
acea
Tuurn, FE. F.

s.n. Schistostemon densiflorum
TRAILL, J. W. H.

80 Humiria balsamifera var. gui-

anensis
81 Schistostemon macrophyllum
Tutin, T. G.
83 Humiria balsamifera fma. atte-
nuata
421 Sacoglottis cydonioides
Uug, E. H. G.
6142 Numiria balsamifera var, gui-
anensis

7625 Humiria balsamifera fma. at
tenuata

CUATRECASAS—HUMIRIACEAE 205

7625 Humiria balsamifera var. flori-
bunda
8801 Vantanea minor

Varescui, V., & Foupats, E.

4573 Humiria balsamifera var. cori-
acea

Schistostemon auyantepuiense
Humiria balsamifera var. cori-

acea

4673
5463

VersteEeeG, G. M.

265 Schistostemon dichotomum

VIGNE, C,
2801 Sacoglottis gabonensis

WacHENHEIM, G.

179 Vantanea parviflora var. par-
viflora

489 Vantanea parviflora var. par-
viflora

WEDDELL, H, A.

526 Humiria balsamifera var. flori-
bunda

2361 Humiria balsamifera var. flori-
bunda

WIuraMs, L.

13868 Humiria balsamifera var. sub-
sessilis

13903 Humiria balsamifera var. gui-
anensis

15052 Humiria balsamifera var. lau-
rina

15418 Humiria balsamifera var. lau-
rina
Wiuirams, L., & Sriva, N. T.

18201 Sacoglottis
fma. glabra

mattogrossensis

WULLSCHLAEGEL, H. R.

1393 Humiria balsamifera var. gui-
anensis

Wourpack, J. J.

293 Sacoglottis amazonica

Worpack, J. J., & Apprrty, L. 8.

42760 Humiria wurdackii

Wourpack, J. J., & Monacwino, J.

40881 Humiriastrum cuspidatum var.
cuspidatum

41149 Sacoglottis guianensis var. his-
pidula

41380 Humiria balsamifera var. sa-
vannarum

ZENKER, G.

148 Sacoglottis gabonensis
440 Sacoglottis gabonensis
1249 Sacoglottis gabonensis
1624 Sacoglottis gabonensis
1671 Sacoglottis gabonensis
1677 Sacoglottis gabonensis
1953 Sacoglottis gabonensis
2499 Sacoglottis gabonensis
2760a Sacoglottis gabonensis
4407 Sacoglottis gabonensis

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Index

Page references to descriptions in Boldface.

Actinostrobites, 34
kayseri, 37
Aubrya, 29, 30, 33, 161
gabonensis, 29, 172
occidentalis, 172
Ctenolophon, 41
Davidia, 40
Diospyros sp., 37
Duckesia, 39, 40, 43 (fig.), 47 (fig.),
48, 76
verrucosa, 78 (fig.), 82 (fig.), 85
(fig.)
Endopleura, 39, 40, 43 (fig.), 47 (fig.),
49, 80
uchi, 78 (fig.), 81 (fig.), 82 (fig.)
Erythroxylon, 28, 30
Eusacoglottis, subg., 30, 161
Hebepetalum, 29, 41
Helleria, 28, 29, 49
obovata, 28, 55
ovalifolia, 28, 55
Hirtella polyandra, 56
Houmiri, 27, 30, 33, 50, 87
arenarium, 89
balsamifera, 27, 89
gabonensis, 172
sect, Aubrya, 161
sect. Saccoglottis, 161
Houmiria, 27, 87
Hugonia, 29, 41
Humiria, 27, 28, 30, 31, 33, 35, 38, 39,
40, 43 (fig), 44, 47 (fig.), 48, 49,
76, 87, 88 (key)
bahiensis, 33, 37
balsamifera, 35, 87, 88, 89, 90
(fig.), 92 (key), 94 (fig.), 95
(fig.), 98, 108
balsamifera attenuata, 91, 92, 94
(fig.), 95 (fig.), 97 (fig.)
balsamifera var, balsamifera, 95
(fig.), 99, 102, 104
balsamifera var. balsamifera fma.
balsamifera, 94 (fig.)
balsamifera var. coriacea, 92, 94
(fig.), 95 (fig.), 97 (fig.), 100
(fig.), 108, 110, 115, 116, 117

Synonyms in Jtalics.

Humiria—Continued

balsamifera var. floribunda, 94
(fig.), 97 (fig), 99, 100 (fig.),
102, 104

balsamifera var. guaiquinimana,
113

balsamifera var. guianensis, 94
(fig.), 95 (fig.), 97 (fig.), 100
(fig.), 103

balsamifera var. iluana, 115

balsamifera var. imbaimadaiensis,
95 (fig.), 115

balsamifera var. laurina, 94 (fig.),
95 (fig.), 100 (fig.), 107, 121

balsamifera var. minarum, 117

balsamifera parvifolia, 95 (fig.),
100 (fig.), 108, 117

balsamifera var. pilosa, 100 (fig.),
116, 117

balsamifera var. savannarum, 94
(fig.), 108

balsamifera var. stenocarpa, 94
(fig.), 97 (fig.), 114

balsamifera var. subsessilis, 94
(fig.), 95 (fig.), 97 (fig.), 102

cassiquiari, 33, 89, 103, 104, 105

cipaconensis, 37

crassifolia, 94 (fig.), 97 (fig.), 121
(fig.)

floribunda, 31, 89, 91, 92, 99

floribunda var. guianensis, 103

floribunda var. laurina, 89, 92, 107

floribunda var. montana, 89, 108

floribunda var. parvifolia, 89, 91,
92, 108 . .

floribunda var. guianensis, 89, 92

floribunda var. spathulata, 32, 89,
107

floribunda var. subsessilis, 89, 102,
103

fruticosa, 100 (fig.), 118

peruviana, 37

pilosa, 34, 89, 116

procera, 33, 143

savannarum, 89, 108

211

212 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Humiria—Continued
subcrenata, 138
wurdackii, 119
Humiriastrum, 30, 39, 40, 43 (fig.), 47

(fig.), 49, 122, 123 (key), 127
(fig.)

colombianum, 127 (fig), 131 (fig),
134

cuspidatum, 127 (fig.), 129, 130,
131 (fig.), 182 (key), 133

cuspidatum var. cuspidatum, 132

cuspidatum var. glabriflorum, 127
(fig.), 181 (fig.), 182, 133

cuspidatum var. subhirtellum, 133

dentatum, 131 (fig.), 136

diguense, 141, 142 (key)

diguense var. anchicayanum, 139
(fig.), 141 (fig.), 142, 143

diguense subsp. costaricense, 142

diguense var. diguense, 139 (fig.),
142

diguense subsp. diguense var.
anchicayanum, 142

diguense subsp. diguense var.
diguense, 142

excelsum, 127 (fig.), 181 (fig.),
133, 1385

glaziovii, 137

glaziovii var. angustifolium, 131

(fig.), 138

glaziovii var. glaziovii, 131 (fig.),
137

mapiriense, 127 (fig.), 139

melanocarpum, 127 (fig.), 139
(fig.), 145

obovatum, 121 (fig.), 125
piraparanense, 127 (fig.), 129
procerum, 127 (fig.), 139 (fig.), 143
subcrenatum, 138
villosum, 121 (fig.), 126
Humirioideae, 48, 76
Humirium, 28, 29, 30, 87, 122, 146
amplexicaule, 89
arenarium, 29, 89, 108, 109
balsamiferum, 29, 89, 99
compactum, 29, 65
contractum, 65
crassifolium, 28, 29, 121
cuspidatum, 29, 130
densiflorum, 28, 29, 160
dentatum, 28, 29, 136
ellipticum, 89, 99
floribundum, 28, 29, 89, 99, 102

Humirium—Continued
guianense, 28, 29, 89, 91, 103
laurinum, 89, 107
macrophyllum, 29, 157
montanum, 28, 29, 89, 91, 100, 108,
109
multiflorum, 89, 99, 108
oblongifolium, 29, 148
obovatum, 28, 29, 125
parviflorum, 89
parvifolium, 28, 29, 108, 109
savannarum, 32, 92
subcrenatum, 28, 29, 138
subsessile, 89, 102
surinamensis, 28, 89, 103, 106
Hylocarpa, 39, 42, 43 (fig.), 47 (fig.),
49, 84
heterocarpa, 78 (fig.), 82 (fig.), 84,
85 (fig.)
Ixonanthes, 29, 30, 31, 41
cochinchinensis, 31
Lemnescia, 27, 49
jloribunda, 27, 71
Lemniscia, 27, 49
floribunda, 71
guianensis, 71
Licania celativenia, 61, 62
Myriodendrum subvaginale, 121
Myrodendron, 87
amplexicaule, 89
petiolatum, 89, 102, 103
Myrodendrum, 27, 87
amplexicaule, 89
balsamiferum, 89
Ochthocosmus, 41
Roucheria, 28
Saccoglottis sect. Eusaccoglottis, 161
Saccoglottis sect. Humiriastrum, 122,
139
Saccogloltis sect. Schistostemon, 146
Saccoglotiis subg. Eusaccoglottis, 161
Saccoglottis, subg. Humiriastrum, 122
Saccoglottis subg. Schistostemon, 146
Sacoglottis, 25, 28, 29, 30, 31, 32, 33,
34, 35, 36, 37, 39, 43 (fig.), 44,
47 (fig.), 49, 77, 81, 84, 161, 163
(key)
amazonica, 28, 29, 32, 34, 35, 36,
40 (fig.), 41, 161, 164 (fig.), 166
(fig.), 167, 168, 169, 170 (fig.),
174
ceratocarpa, 33, 164 (fig.), 165,
166 (fig.)

—

INDEX 213

Sacoglottis—Continued

cipaconensis, 25, 32, 33, 37

cipaconensis var. peruviana, 33, 37

cipaconensis var. peruvianus, 32

costata, 32, 37

cuspidata, 130

cydonioides, 175, 177 (fig.), 178
(fig.), 183, 185

densiflora, 160

dentata, 136

dentata var., 137

dichotoma, 30, 159

diguensis, 33

diguensis var. anchicayana, 142

duckei, 31, 157, 159

excelsa, 133

excelsa var. colombiana, 134

excelsa var. glabriflora, 132

gabonensis, 32, 35, 36, 37, 41, 162,
164 (fig.), 166 (fig.), 172

gabunensis, 172

germanica, 33, 37

glaziovii, 137

guianensis, 29, 174, 176 (key),
181, 185

guianensis fma. dolichocarpa, 174,
176

guianensis var. guianensis, 177
(fig.), 178 (fig.)

guianensis var. guianensis fma.
glabra, 178 (fig.)

guianensis var. guianensis fma.
guianensis, 176

guianensis var. hispidula, 178
(fig.), 180

guianensis var. maior, 174, 178
(fig.), 179

heterocarpa, 32, 84, 85

kaboeriensis, 33, 160, 161

kayseri, 34, 37

macrophylla, 157, 179

maguirei, 164 (fig.)

mattogrossensis, 32, 175, 177 (fig.),
181, 182 (key)

mattogrossensis fma. mattogros-
sensis, 178 (fig.)

mattogrossensis var. mattogros-
sensis fma. glabra, 182

mattogrossensis var. mattogros-
sensis fma. mattogrossensis, 182

mattogrossensis var. subintegra
fma. puberula, 170 (fig.), 177
(fig.), 178 (fig.), 183

Sacoglottis—Continued
mattogrossensis var. subintegra
fma. subintegra, 183
melanocarpa, 33, 145
oblongifolia, 31, 148
obovata, 125
ovicarpa, 33, 37, 164 (fig.), 166
(fig.), 168
procera, 143
reticulata, 153
retusa, 156
subcrenata, 138
tertiaria, 32, 37
uchi, 31, 80, 81
verrucosa, 32, 78
villosa, 34, 126
Sacoglottis sect. Husaccoglottis, 161
Sacoglottis, sect. Humiriastrum, 122
Sacoglottis sect. Schistostemon, 146
Schistostemon, 30, 37, 39, 43, 44 (fig.),
47 (fig.), 49, 146, 147 (key)
auyantepuiense, 151 (fig.), 170
(fig.)
densiflorum, 150 (fig.), 154 (fig.),
160
dichotomum, 154 (fig.), 159
macrophyllum, 150 (fig.), 154
(fig.), 157
oblongifolium, 147, 148, 149 (fig.),
150 (fig.), 151 (fig.)
reticulatum, 149 (fig.), 153, 157
reticulatum subsp. froesii, 149
(fig.), 151 (fig.), 154
reticulatum subsp. reticulatum,
149 (fig.), 154 (fig.)
retusum, 149 (fig.), 150 (fig.), 151
(fig.) 156
Spondylostrobus smythii, 37
Tectonia grandis, 40
Vantanea, 25, 27, 29, 30, 31, 32, 33,
37, 38, 39, 40, 42, 43 (fig.), 47
(fig.), 48, 49, 51, 52 (key), 70,
77, 81
barbourii, 33, 51, 53, 57 (fig.), 143
celativenia, 51 (fig.), 61 (fig.)
colombiana, 32, 37
compacta, 52 (fig.), 61 (fig.), 65
compacta subsp. compacta var.
compacta, 67
compacta subsp. compacta var.
grandiflora, 67
compacta subsp. microcarpa, 67

214

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Vantanea—Continued

compressiformis, 32, 37

contracta, 65

contracta var. grandiflora, 67

cupularis, 31, 74

guianensis, 27, 28, 29, 43 (fig.),
44, 50, 52, 71 (fig.), 72 (fig.)

macrocarpa, 64 (fig.), 68 (fig.), 70

magdalenensis, 34, 51 (fig.), 54,
55 (fig.)

micrantha, 50 (fig.), 52 (fig.), 61
(fig.), 62

minor, 29, 50 (fig.), 52 (fig.), 61
(fig.), 63

obovata, 29, 55 (fig.)

occidentalis, 33, 40 (fig.), 51 (fig.),
57 (fig.), 59

Vantanea—Continued
ovalifolia, 29, 55
panniculaia, 65, 67
paraensis, 51 (fig.), 57 (fig.), 60, 61
parviflora, 27, 29, 44, 50 (fig.), 52

(fig.), 71 (fig.), 74

parviflora var. parviflora, 75
parviflora var. puberulifolia, 75, 76
peruviana, 33, 64 (fig.)
sheppardi, 32, 37
tuberculata, 69, 70 (fig.), 71 (fig.)
wilcoxiana, 37

Vantaneoideae, 48, 49

Vantaneoides, 30, 50

Verniseckia, 87

Wernischeckia, 87

Wernisekia, 27, 87

U.S. GOVERNMENT PRINTING OFFICE: 1961

UNITED STATES NATIONAL MUSEUM

CONTRIBUTIONS FROM THE UNITED STATES NATIONAL HERBARIUM

Vo Lume 35, Parts 3 Anp 4

3. ORIGINS OF THE FLORA OF SOUTHERN BRAZIL

4. A SYNOPSIS OF THE AMERICAN VELLOZIACEAE

By Lyman B. Smite

BULLETIN OF THE UNITED STATES NATIONAL MuSEUM

SMITHSONIAN INSTITUTION e WASHINGTON, D.C. e 1962

Contents

Page

3. Origins of the Flora of Southern Brazil... .........2.. ~ 4215
Maritime Zone ...... ee ee ee 216
Coastal Rain Forest ... 2... ee ee ee B19
Paraguayan Rain Forest .. 2... 2... 2. ee ee ee 228
Northern Campo Flora... . . ...... 2... ee 224
Andean Flora... 2... ee ee ee BBE
Overseas Disjuncts. . . .... ee ee ee 287

4. A Synopsis of the American Velloziaceae ..........2.2.2. 251

Part 3

ORIGINS OF THE FLORA OF SOUTHERN BRAZIL

Lyman B. SMITH

Ever since the first land plants evolved, the face of the earth has
been changing constantly, so that we may safely assume that the
flora of any given region today has arrived from somewhere else. In
the case of land recently risen out of the sea, it is relatively easy to see
whence its flora came, but in the case of a great center of distribution
like the Amazon Basin it is difficult if not impossible to reconstruct
its past.

Southern Brazil, which is to say the states of Sdo Paulo, Parana,
Santa Catarina, and Rio Grande do Sul, is an intermediate case. The
area is not recently exposed except for a narrow coastal fringe, but on
the other hand, most of its plant families have their greatest develop-
ment elsewhere. Thus there are some fairly obvious lines of migration
(fig. 1), and the continuity of some ranges and the discontinuity or
disjunction of others indicate that these migrations have taken place
at different times. The flora of the coastal rain forest is a direct
continuation and attenuation of that centering on Rio de Janeiro,
while the flora of the campos or rolling prairies of the planalto has a
similarly direct origin from the central states of Minas Gerais and
Goids. There is another continuous migration pattern from northern
Argentina and Paraguay eastward and a considerable amount from
the Andes. We have a few wide ranging maritime species extending
from the north into the area, but from due south there is relatively
little evidence of immigration, possibly because this area may not have
emerged from the sea until a later date.

The discontinuous ranges follow much the same migration routes
as the continuous ones from the north, but these are suspect as being
due to gaps in collecting. Of those based on fairly firm evidence the
most important by far are the Andean, with only a few species each
for the Austral, coastal plain, and African.

It may seem presumptuous and reckless to discuss ranges at all on
the basis of such meager evidence as we now have, and probably we

215

216 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

still need several hundred years of collecting and classifying before the
flora of Brazil is really well known. However, there are certain broad
physiographic regions that are readily recognizable even if they do
blend at the boundaries, and while collecting and classifying we become
aware of phytogeographic patterns that correlate with them, like a
picture slowly coming into focus. Furthermore we cannot afford to
wait indefinitely because the rapid advance of agriculture is eliminating
the native flora over much of Brazil.

Maritime Zone

There is always a seacoast and the highly specialized halophytic
flora closely follows its extremely narrow and infinitely long zone.
For thousands of miles one can pick a stretch of Brazilian coast at
random and know that he can find Ipomoea pes-caprae (pl. 1) and
Remirea maritima (fig. 2) on the sand just above high-tide mark or
Rhizophora mangle (pls. 1,2) at the seaward end of an estuary. Higher
up in the restinga or scrub forest zone or on saltmarsh will be
Hibiscus tiliaceus. If we had complete records of these species their
ranges would form an almost continuous line on the map. This
assumption is one of the safest that we will make because maritime
conditions are much the same the world over.

However, the maritime flora of southern Brazil has one unusual
feature when compared to those familiar in the Northern Hemisphere.
So far as present evidence shows, its wide ranging species are all of
tropical derivation and none of circumpolar. The list of such strictly
or predominantly maritime species would include:

Sporobolus virginicus (Gramineae)
Stenotaphrum secundatum

Remirea maritima (Cyperaceae)
Salicornia virginica (Chenopodiaceae)
Canavalia rosea (Leguminosae)
Hibiscus tiliaceus (Malvaceae)
Rhizophora mangle (Rhizophoraceae)
Conocarpus erecta (Combretaceae)
Laguncularia racemosa

Hydrocotyle bonariensis (Umbelliferae)
Ipomoea pes-caprae (Convolvulaceae)
Scaevola plumieri (Goodeniaceae)

Some of the above like Sporobolus virginicus and Stenotaphrum secunda-
tum (fig. 3) although predominantly maritime also occur inland.
Species like Rhizophora mangle on the Ilha de Santa Catarina, reach
their limits in southern Brazil, while others extend beyond, but ap-
parently none are found south of the province of Buenos Aires.
There are a number of subtropical species that are common in the

SMITH—ORIGINS, FLORA OF SOUTHERN BRAZIL 217

MIGRATIONS
m———--———+ CONTINUOUS RANGE i
Beeeee + DISJUNCT RANGE

SCALE
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Ficure 1.—Different types of migration into southern Brazil.

maritime zone from southern Brazil to Uruguay and the province of
Buenos Aires. Among them may be noted:

Triglochin striata (Scheuchzeriaceae)
Androtrichum trigynum (Cyperaceae)
Chenopodium retusum (Chenopodiaceae)
Alternanthera maritima (Amaranthaceae)
Acicarpha spathulata (Calyceraceae)

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

218

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SMITH—ORIGINS, FLORA OF SOUTHERN BRAZIL 219

At first glance one might suppose that some of these arrived from the
south, but in each case checking reveals that more likely the migration
was in the opposite direction or from the west across northern Argen-
tina. Whether southern Argentina was submerged in recent times or
not, its present climate appears to be too arid to allow any trace of
southern connections below the province of Buenos Aires.

The maritime zone is unlike any of those that follow in that it in-
cludes no disjunct ranges. This situation is no more than might be
expected from the continuous and constant ecological conditions that
it provides.

Coastal Rain Forest

The coastal rain forest (pl. 2) extends some 2000 miles or 3200
kilometers from extreme northern Rio Grande do Sul to the Herval and
Tapes Ranges in Rio Grande do Norte. Thus it is about the same
length as the Amazonian rain forest though only a fraction of the
width, A flight from Trinidad to Rio de Janeiro takes several hours
to cross the Amazonian forest but only minutes to cross the coastal.
In general the western boundary of the coastal forest is the height of
land from which the sea is visible from many points. Only in Santa
Catarina does this zone extend inland to a depth of about 100 miles
or 160 kilometers.

In relation to its area the coastal rain forest must be fully as rich
in species as the Amazonian but the representation by families differs
greatly. Some families like the Vochysiaceae (Stafleu, 1948) are ob-
viously remnants of much larger ones of the Amazonian forest, while
others like the Bromeliaceae (Smith, 1934) have formed a new center
in the coastal forest and evolved many new species and genera.
The center of speciation of the coastal rain forest is very close to
Rio de Janeiro, now the state of Guanabara. From there the flora
deteriorates north and south, so that in the area we are considering
the species decrease regularly from Sao Paulo to northern Rio Grande
do Sul.

For example the species of the genus Vriesea (Smith, 1955) in the
Bromeliaceae are distributed from north to south as follows:

Species
Espirito Santo... 1. we ee ee 31
Rio de Janeiro and Guanabara (former Federal District). . 54
Sdo Paulo... . . . ee 31
Parani . 2... we ee 26
Santa Catarina... . ee 28
Rio Grande do Sul .............8. 20884 7

The areas of rain forest are not all commensurate, but the comparison
between the first three is fairly close on this score. Parand has much
less rain forest and Santa Catarina with its westward bulge much

230 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

more, and Rio Grande do Sul only a remnant. If these scores were
to be weighted by areas there would be a fairly regular gradation
from Rio de Janeiro southward. On the other hand, Rio de Janeiro
enjoys the advantage of the greatest amount of collecting, so if this
factor could be weighted also, Rio would not stand quite so high
above the others.

More graphic pictures of the situation, though unfortunately for a
smaller area, are given by Reitz (1958, 1957) for the Palmae (fig. 4)
and Araceae (fig. 5) of Santa Catarina. Both show an overlapping
pattern of southern species limits like wave marks on a strand or
tiles on a roof, presumably due to the differential tolerance of the
various species to minimum temperatures. The point was made dra-
matically when the boom of southward expansion of coffee cultivation
was stopped by a severe winter with unexpected frosts.

Reitz’s map of the Araceae shows another feature of distribution,
the evolution or isolation of local species in a wide ranging genus,
Anthurium, thus bringing in the element of time in migrations. The
other species of Anthurium have expanded southward into Santa
Catarina without essential change, while the ancestral stock of A.
lacerdae and A. pilonense has not only migrated over the same route
but developed new species as well. Whether their present restricted
range is due to newness or shrinking from a wider area, it still indi-
cates an earlier arrival for the parent stock than the species that are

oS Redele*
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ESTADO DE SANTA CATARINA
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Trithrinax brasiliexsis Mart.
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wett4s Futerce edulis Mart.

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eemees Aslrocaryum acileatissinuim (Schott) Bui’.

Dap. Esladeal da Cargratin¢ Carvegrafia Jia

Ficure 4.—Rain forest migration: Palmae. After R. Reitz 1953.

SMITH—ORIGINS, FLORA OF SOUTHERN BRAZIL 221

fe)
°
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DISTRIBUIGAO DAS ARACEAS C
EM SANTA CATARINA

Pistia stratiotes, Asterostigma lividum
© © O {Spathicarpa hastifolia, Philod. bipinnatitidum,
Philodendron ochrostemon var. uleanum,
+ + + {Philodendron imbe. Anthurium scandens,
thurium willdenowii.

jonstera pertusa, Heteropsis rigidifolia,

JAsterostigma t di » Phil dana A
dendron “i ;

Phitodendron = renduxii, Anthurium undatum,

Anthurium acutum, Anthurium gdaudichaudianum ,

jum harrisii var. beyrichienum .

ee0 Euioendron crassinervium, Philodendron cordatum ,
Phifodendron melanorrhizum

Canthurium lacerdoe(}) Anthurium pilonense(2)

Txedercs

Figure 5.—Rain forest migration: Araceae. After R. Reitz 1957.

unchanged. Admittedly there is a possibility that in some cases
species have evolved at different speeds, otherwise we should not
have any primitive forms of life left in the world. However, within
the same genus it would seem less likely for such a difference to be
large. The same comparison may be found between genera in dif-
ferent families, where Vantanea compacta (fig. 6) is the sole repre-
sentative of the Humiriaceae (Cuatrecasas, 1961) from Rio de Janeiro
to Santa Catarina, while Quesnelia (fig. 7) of the Bromeliaceae (Smith,
1955) is represented by several species in Rio de Janeiro and Guana-
bara, a different set in Sao Paulo, and still another in Parandé and
Santa Catarina. The probability is that Vantanea arrived in the
rain forest of Santa Catarina ahead of Quesnelia, the year to year
change of position in migration being more perceptible than the
change of form in evolution. .

The rain forest of southern Brazil also shows evidence of migration
from the Amazonian rain forest (fig. 1), though of course to a much
lesser degree than from the much closer center of Rio de Janeiro.
The smallness of the Amazonian element is easy to understand when
one considers the extensive barrier interposed by the high dry planalto
area that lies between the two rain forests for a minimum distance
of over 700 miles or 1100 kilometers. For many genera and even
some families this climatic barrier is as effective as an ocean. Others,
however, have found a bridge across through the network of gallery

222 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

forests (pl. 4) that break up the area. The extent and effectiveness
of this planalto barrier has fluctuated in the past and is still visibly
fluctuating now. As has long been pointed out, new young tree
growth invades the campo or prairie of the planalto as the climate
grows moister or dies back as it grows drier. This is comparable
to the constantly shifting division between long grass and short
grass prairie in central United States and Canada.

The different patterns of migration from the Amazonian rain forest
would indicate that there have been major fluctuations in the climate
so that the barrier of the planalto has been bridged at different times
for different groups of plants. Beginning with the migrations that
appear most recent because the species extend across northern and
central South America without any noticeable change or at most
with forms that show no geographic correlation, we have (figs. 8-10):

Protium heptaphyllum (Burseraceae—fig. 8, Swart, 1942)
Tapirira guianensis (Anacardiaceae—fig. 9)
Amaioua guianensis (Rubiaceae—fig. 10, Smith & Downs, 1956)

These trees must have migrated by gallery forest or there may
have been continuous broad rain forest between the Amazon and the
east coast under more favorable conditions.

Other species combine Amazonian and Andean distribution and
appear to have spread southward through thinner forest at higher
elevations. Such are (figs. 11-13):

Chlorophora tinctoria (Moracvae—fig, 11)

Casearia silvestris (Flacourtiaceae—fig, 12)

Psychotria carthagenensis (Rubiaceae—fig. 13, Bacigalupo, 1952)
On present evidence they have flanked the planalto rather than crossed
it, but further collecting in the interior could change this picture
considerably.

There are also certain weedy species that are not truly Amazonian
but show much the same overall distribution, not because they are
characteristic of the forest but because they immediately take ad-
vantage of any break in it, however small. Such are (figs. 14-16):

Axonopus compressus (Gramineae—fig. 14, Black, ined.)
Xyrts caroliniana (Xyridaceae—fig. 15, Smith & Downs, ined.)
Polygonum punctatum (Polygonaceae—fig. 16)

At the other extremity of distributional types is the case of the
genus Vochysia (fig. 17). Here nearly the same area has been covered,
but the migration has been so slow that there has been time for the
evolution of a completely new set of species on the planalto and almost
another in the coastal rain forest. There is also the disjunct of Quiina
(fig. 18, Pires, ined.) with one species, Q. glaziovii, on the coast and
the remainder confined to the Amazon basin. This may indicate a
still older migration because of the absence of planalto species, al-
though there is always the possibility of others turning up there.

|

SMITH—ORIGINS, FLORA OF SOUTHERN BRAZIL 223

The migration appears to have taken place at different points all
across the planalto judging by the occurrence of some species over
nearly the entire length of the coastal rain forest. A very few like
Chlorophora tinctoria (fig. 11), Casearia silvestris (fig. 12), and Psy-
chotria carthagenensis (fig. 13) have even connected with the Rio
Paraguay Basin and entered southern Brazil across the top of Rio
Grande do Sul, meeting the final segment of the coastal rain forest in
the northeast corner of the state. However, the Amazonian element
is minor, the Paraguayan Basin having developed a flora of its own
to an even greater extent than has the coastal rain forest.

Paraguayan Rain Forest

The rain forest of the Rio Paraguay Basin is not a continuous area
comparable to the Amazonian or the coastal rain forests but a collec-
tion of gallery forests that are so broad that they merge when large
rivers lie close together as in the Misiones Territory of Argentina.
A comparison of Richards’s description of typical rain forest (Tropical
Rain Forest, pp. 2-7) and Hauman’s of that in Misiones (La Vege-
tacion de la Argentina, pp. 14-41) shows agreement in all details, yet
Richards in his figure 2 follows A. C. Smith and Johnston (Verdoorn,
Plants and Plant Science in Latin America, pp. 12, 13) in ignoring
this area on his map. Hoehne (Ind. Pl. Com. Rondon, p. 12) and
Rambo (Sellowia, No. 7, p. 185) both emphasize the gallery character
of this rain forest and their view is much broader than Hauman’s, so
that Richards after all would seem justified in not including this area
had he not included similar fragments in northern South America.

Be that as it may, the southern part of the Paraguayan forest ex-
tends over eastern Paraguay, Misiones, and the extreme western parts
of Parana, Santa Catarina, and Rio Grande do Sul, and has developed
a very distinctive flora. It has invaded southern Brazil up the Rio
Uruguay and its tributaries, and its migration shows the same over-
lapping pattern of distribution as that of the coastal rain forest but
with an eastward instead of a southward direction. In Santa Catarina
where it is known as ‘‘mato branco”’ the forest is quite obviously ex-
panding up the river valleys and invading the Araucaria and campo
zones (personal field observation and Klein, 1960). Rambo notes the
same movement in Rio Grande do Sul. Of the four hundred species
listed for the upper Rio Uruguay by Rambo (1956a, pp. 191-207)
special mention might be made of the following to illustrate the type
of ranges in the Paraguayan forest flora (figs. 19-26):

Pseudoplantago friesit (Amaranthaceae—fig. 19, Smith & Downs, 1960)
Balfourodendron riedelianum (Rutaceae—fig. 20, Cowan, 1960)
Holocalyx balansae (Leguminosae—fig. 21, Burkart, 1943)

Bernardia pulchella (Euphorbiaceae—fig. 22, Smith & Downs, 1959)

2234 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Euphorbia sciadophila (Kuphorbiaceae—fig. 23)

Diatenopteryx sorbifolia (Sapindaceae—fig. 24, Radlkofer, 1931-34)
Patagonula americana (Boraginaceae—fig, 25)

Dunalia breviflora (Solanaceae—fig. 26, Sleumer, 1950)

Some species have rather restricted ranges like Pseudoplantago
Jriesvt and Holocalyx balansae, while others such as Bernardia pulchella
and Dunalia breviflora have crossed the planalto and the divide and
entered the coastal rain forest. It has been claimed that new or
isolated species do not occur in the Paraguayan forests of southern
Brazil, which would indicate a recent migration. Certainly they
are not so evident as in the coastal rain forest. On the other hand,
such monotypic endemics as Pseudoplantago (fig. 19), Aphaerema
(Flacourtiaceae), and Schenckia (Rubiaceae), would argue a rather
distant origin for the Paraguayan forest flora as a whole.

Northern Campo Flora

The campo or prairie flora of southern Brazil derives in part from
the central Brazilian highland or planalto and in part from the Andes.
It is a crazy quilt cut by a fine dendritic pattern of angiosperm gallery
forest in much of Sao Paulo and Rio Grande do Sul and occurring
only as isolated openings in the Araucaria formation of Parana,
Santa Catarina, and northern Rio Grande do Sul. It is considered
by some to be the oldest of the large formations and the lack of a
well defined pattern of distribution would support the idea. Several
families and many genera center in Minas Gerais on the planalto,
but it is even more difficult than in the Amazonian flora to find species
with a continuous distribution into southern Brazil.

The Velloziaceae (fig. 27), one of the most characteristic families
of Minas Gerais, well illustrates this situation. The plants grow on
dry barren slopes and crests and the species in most cases have small
ranges. ‘The few that reach southern Brazil are isolated species known
from a single locality each. Incidentally this family also illustrates
a fallacy of some plant geographers. Its world range has been shown
as a line enclosing the whole Amazon basin, while its true occurrence
there is limited to a few bare peaks near the edge of it. Similarly
the points indicated for Vellozinceae in the area of the coastal rain
forest all represent bare granite peaks rising above the forest (pl. 3).

Perhaps Dyckia (fig. 28) in the Bromeliaceae is a better example be-
cause 1t ranges over the planalto of all southern Brazil and beyond,
though it does include more species on the coast and in open spots in
the coastal rain forest. Again the species are mostly narrow endemics.
There are 28 species in Minas Gerais against only 2 in Rio de Janciro
and Guanabara showing that it is truly a genus of the planalto, and

SMITH—ORIGINS, FLORA OF SOUTHERN BRAZIL 225

the numbers hold up well southward with 7 in Sao Paulo, 8 each in
Paran& and Santa Catarina, and 9 in Rio Grande do Sul.

The Eriocaulaceae are undoubtedly the most extreme case of high
concentration in Minas Gerais rapidly dwindling out in southern
Brazil as the distribution of the species of Paepalanthus illustrates
(from Moldenke, 1959 a—b, 1960):

Species
Minas Gerais... 1 ee eee 660
Rio de Janeiro and Guanabara . . ... 1... ee es 21
Saio Paulo. . 1... ee ee 27
Parandé ......... Co ee kk ee ke ee 9
Santa Catarina. . . .. . Coe ek ee 8
Rio Grande do Sul...) . ee 5
Paraguay .... .. ee eee Ce . 2
Argentina... 2... ee ee Co ek ee 0
Uruguay 6 6 0

It has been difficult to find single species to illustrate the full sweep
from the central planalto south, which is probably another indication
that the flora is old and that its genera have had time to break up
by evolving new species to fit local conditions. Hither the ranges
are too limited to give a good illustration or they spread out into the
coastal zone taking advantage of open areas and thus are not wholly
typical of the planalto. However, the following give some idea
of the situation (figs. 29-33):

Esterhazya splendida (fig. 29—partly coastal) (Barroso, 1952)
Baccharis anomala (fig. 30—partly coastal)

Baccharis megapotamica (fig. 31—not known to be coastal)
Vernonia nitidula (fig. 32—partly coastal)

Vernonia platensis (fig. 33—not known to be coastal)

Some species like Lobelia camporum are found throughout the campo
of southern Brazil up to the highest points but are indifferent to
altitude and occur also at low levels along the coast. Even such
species of general distribution in the area are disjuncts in a sense
because the campo itself is disjunct and does not provide the means
of a continuous range.

Andean Flora

The genera that have invaded southern Brazil from the Andes
show the greatest diversity in distribution of all that we have studied
so far, indicating the longest time elapsed during the migrations
from first to last. Some genera include single species that bridge the
gap and go far beyond, while others are widely disjunct with few,
localized, and distinct species on either side. The first represent
extremely recent migrations and the second ancient ones and there
are many intermediate cases.

226 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

The recent type of migration is shown by Tillandsia usneoides (fig.
34) of the subgenus Diaphoranthema. Its range is so large that it is
thought of as neotropical. Yet its kindred are all represented and
in some cases confined to a small area in northwestern Argentina and
vicinity (Smith 1934, 1935). Further testimony of its Andean origin
and preference for cool climate is shown by its avoidance of the
Amazon basin and its extension in the United States as far north as
Virginia, the extreme point in the Bromeliaceae by over 500 miles or
900 kilometers. Its recent evolution is shown by its highly specialized
structure, with the inflorescence reduced to a single flower and the
leaves taking over the functions of the lost roots. Everything points
to Tillandsia usneoides having been the latest invader of southern
Brazil from the Andes.

However, most genera of Andean origin do not include single species
extending all across South America and have a maximum distribution
in the east from slightly north of Rio de Janeiro to Uruguay. Almost
all of these have the greater part of their eastern range in the four
southern Brazilian states and many do not extend beyond. Also in
the larger examples of this pattern there is a link across northern
Argentina to the area of distribution in the Andes. The species of
southern Brazil are plants of cool moist climates and have their center
of speciation on the crests of the divide and from there extend west-
ward on the planalto mingling and competing with the flora of the
northern campo zone. The species connecting across northern Argen-
tina are necessarily more tolerant of heat and aridity than those on
either side of them. The ranges of the Andean genera have been
pinched in the middle by the southward push of Amazonian heat and
the northward push of Patagonian aridity (fig. 35). In many cases
the link has been completely broken. This situation makes interesting
comparison with North American disjunctions between east and west
where glaciation has been used as an explanation. The most recent
glaciation in South America preceded flowering plants, so that climate
alone accounts for not only the occurrence of the disjunction but for
its continuance also.

One of the best examples we know of a wide ranging Andean element
in southern Brazil is the genus Hypericum (fig. 36). It has numerous
species endemic to the Andes and 25 found only in eastern South
America (Smith, 1958b). Hypericum connatum by a strange coinci-
dence is the link across, its name referring not to this but to its
perfoliate leaves. In addition to what is indicated on the map, there
is H. mutilum of North America occurring on both sides of South
America apparently from pre-Columbian times, JZ. perforatum
recently naturalized from Kurope, and H. gentianoides, a natural
disjunct with North America.

SMITH—ORIGINS, FLORA OF SOUTHERN BRAZIL 227

The genus Berberis (fig. 37—Schneider, 1904-05) has an overall
appearance in South America much like Hypericum, but has a single
disjunct species, B. montana, occurring in central Chile and on Campo
dos Padres, the highest peak in Santa Catarina. It is thus intermedi-
ate between Hypericum and the genera that are completely disjunct.

Herreria (fig. 38) is unusual in having more species in the east than
in the west (Smith, 1958a) and unlike Hypericum and Berberis does
not occur outside South America. It still has a link across northern
Argentina. Viviania (fig. 39), however, has lost all connection and
has no species common to both areas (Knuth, 1912). Crinodendron
(fig. 40) is similar but its representation in southern Brazil is limited
to C. brasiliense (Reitz & Smith, 1958), endemic to the peak of Campo
dos Padres.

Araucaria (fig. 41) is represented by A. angustifolia in southern
Brazil (pl. 4) and A. araucana in the southern Andes on the Argentine-
Chilean boundary, and by a number of species in Australia and the
southwestern Pacific region. The disjunction is the longest we have
met of this type, some 1100 miles or 1800 kilometers, and at the
same time the most certain. Herbs or low shrubs might be overlooked
in some sheltered spot in the interval, but there is no chance of hiding
the stately pinheiro. It still shows movement as its heliophile progeny
slowly invade the campo, but on the other side it is losing at a much
faster rate to the inroads of the Paraguayan forest and to man (Klein,
1960).

Cordyline (Baker, 1875) of the Liliaceae has a range much like that
of Araucaria except that there is no Andean species known to link
©. dracaenoides (fig. 42) of southern Brazil with the rest of the genus
on the other side of the Pacific. It might equally well be classed with
our overseas disjuncts except for this similarity.

Overseas Disjuncts

Under the title of overseas disjuncts we have a miscellany of odds
and ends, some of which are probably pre-Columbian arrivals in
southern Brazil and others that are not, but are obvious weedy new-
comers. Sometimes it is difficult to decide which. Arenaria groen-
landica on a single peak in Santa Catarina (Smith & Downs, 1960b)
and known elsewhere from Georgia to Greenland in North America
hardly seems likely to be a weed. Hypericum gentianoides of Rio
Grande do Sul, Paraguay, and eastern United States (Smith, 1958b)
might possibly be a recent introduction, but Rambo who found
it in Brazil, thinks it unlikely because of the habitat. Proserpinaca
palustris (Reitz, 1954) of Santa Catarina and Rio Grande do Sul is
indistinguishable from the North American plant. The three taken
together and compared with a more nearly continuous range such as
that of Utricularia subulata make a fairly consistent pattern.

597430—62

2

238 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Links with Africa, whether specific like Xyris anceps and X. capensis
or generic like Piteairnia and Rhipsalis, are indirect, with the South
American center lying to the north of the area we are considering.
The European links, so far as I have been able to ascertain, are all
recent weedy introductions. Rambo has covered much of the details
of this flora (1960) and its origin is too obvious to need discussion here.
If his list is anywhere near complete it would indicate that southern
Brazil is not so badly overrun with weeds as are equivalent latitudes
in North America.

Acknowledgments

The present paper is an enlargement of one presented at the pinth
International Botanical Congress at Montreal in 1959. Unless other-
wise noted the range maps have been compiled from specimens in
the United States National Herbarium, However, as indicated in the
bibliography, I have leaned heavily on monographic and floristic works
where available. Personal observations were made throughout the
planalto of Santa Catarina in 1956-57 on a grant from the National
Science Foundation. The coastal rain forest was studied in 1928-29
on a Sheldon Travelling Fellowship from Harvard University and in
1952 on grants from the Rockefeller Foundation and the Servico
Nacional de Malaria of Brazil. The two most recent expeditions were
made jointly with the Herbario “Barbosa Rodrigues” of Santa
Catarina.

Unless otherwise indicated, the maps shown in this paper are based
on the Goode base map No. 103, copyright 1937 by the University of
Chicago.

229

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SMITH—ORIGINS, FLORA OF SOUTHERN BRAZIL 235

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Ficure 18.—Ancient Amazonian migration: Qutina.

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

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the Andes and southern Brazil.

SMITH—ORIGINS, FLORA OF SOUTHERN BRAZIL

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597430—62——3

243

244 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

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SMITH—ORIGINS, FLORA OF SOUTHERN BRAZIL QA5

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SMITH—ORIGINS, FLORA OF SOUTHERN BRAZIL 247

Ficure 39.—(Top left) Andean migration: Vivianta.

Ficure 40.—(Top right) Andean migration: Crinodendron.

Ficure 41.—(Bottom left) Western Pacific migration: Araucarta.

Ficure 42.—(Bottom right) Western Pacific migration: Cordyline dracaenoides.
Bibliography

Some of the following works are referred to directly in the text,
while others have been used indirectly as background material, but
they all add to our information on the subject.

BacuicaLupo, Nénma Marfa. Las especies argentinas de los géneros Psychotria,
Palicourea y Rudgea (Rubiaceae). Darwiniana, vol. 10, pp. 31-64. 1952.

BaxkeEr, J.G. Revision of the genera and species of Asparagaceae. Journ. Linn.
Soc., vol. 14, pp. 508-632. 1875.

Barroso, G. M. Scrophulariaceae indigenas e exéticas no brasil. Rodriguésia,
vol. 15, No. 27, pp. 9-64. 1952.

Buack, Grorce A. A taxonomic treatment of the genus Aronopus. Doctoral
thesis, The George Washington University.

Burxart, Arturo. Las leguminosas argentinas silvestres y cultivadas, pp.
1-590. 1943.

Casprera, ANGEL Lutio. Manual de la flora de los alrededores de Buenos Aires,
pp. 1-589. 1953.

Carn, StanteEy A. Foundations of plant geography, pp. 1-556. 1944.

Cuopat, R. La végétation du Paraguay, pp. 1-157. 1916.

Cowan, R. 8. Rutaceae of Santa Catarina. Sellowia, No. 12, pp. 79-97. 1960.

Cuarrecasas, J. A taxonomic revision of the Humiriaceae, Contr. U.S. Nat.
Herb., vol. 35, pt. 2, pp. 25-214. 1961.

DANSEREAU, Pierre. Biogeography an ecological perspective, pp. 1-394. 1957.

Frernatp, M. L. Persistence of plants in unglaciated areas of boreal America.
Mem. Amer. Acad. Arts & Sci., vol. 15, pp. 295-317. 1925.

Gurason, H. A. The progress of botanical exploration in tropical South America.
Bull. Torrey Bot. Club, vol. 59, pp. 21-28. 1932.

Havuman, L., BurKart, A., Paropi, L. R., & Caprera, A. L. La Vegetacién
de la Argentina, pp. 1-349. 1947.

Horung, F. C. Phytophysionomia do estado do Matto-Grosso (with map), pp.
1-104. 1923.

Indice bibliogréphico e numérico das plantas colhidas pela Comissao
Rondon, pp. 1-400. 1951.

Kurin, R. M. O aspeto dindmico do pinheiro brasileiro. Sellowia, No. 12, pp.

17-44. 1960.
Knutu, R. Geraniaceae. Engler, Pflanzenreich, tv. Fam. 129, pp. 1-640.
1912.

LAWRENCE, GeorcE H. M. Taxonomy of Vascular Plants, pp. 1-823. 19951.

Martius, K. F. P. von. Flora Brasiliensis, 15 vols. 1804-1906.

Mo.tvEnKE, H. N. A résumé of the Verbenaceae, Avicenniaceae, Stilbaceae,
Symphoremaceae and Eriocaulaceae of the world as to valid taxa, geographic
distribution and synonymy, pp. 1-495. 1959a.

Supplement 1, pp. 1-26. 1959b.

Supplement u, pp. 1-16. 1960.

Prres, J. Murcga. Monografia Quiinaceae, ined.

248 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

RaADLKOFER, L. Sapindaceae. LTngler, Pflanzenreich, 1v. Fam. 165, pp. 1-1539.
1931-34.

RamsBo, Batpuino. A imigragao da selva higréfila no Rio Grande do Sul. Sello-
wia, No. 3, pp. 55-91. 1952.

————. Der Regenwald am oberen Uruguay. Sellowia, No. 7, pp. 183-233.
1956a.

———. A flora fanerogamica dos aparados Riograndenses. Sellowia, No. 7,
pp. 235-298. 1956b.

———. A fisionomia do Rio Grande do Sul. Jesuitas no sul do Brasil, ed. 2,
vol. 6, pp. 1-456. = 1956c.

—-——. Die Europiiischen Unkrauter in Siidbrasilien. Sellowia, No. 12, pp.
45-78. 1960.

Reitz, Rautino. As palmeiras de Santa Catarina e a sua distribuigdo geo-
grafica. An. Bot. Herb. “Barbosa Rodrigues,” No. 5, pp. 233-252. 1953.

a . As Halorrhagaceae de Santa Catarina. Sellowia, No. 6, pp. 237-242.

——-—, Ardceas catarinenses. Sellowia, No. 8, pp. 20-70. 1957.

Rerrz, RAULINO, AND Smith, Lyman B. Crinodendron no Brasil. Sellowia,
No. 9, pp. 19-21. 1958.

Ricwarps, P. W. The tropical rain forest, an ecological study, pp. 1-450. 1952.

Rosas, TEopoRo, AND Carasia, J. P. Breve resefia de la vegetaci6n Paraguaya.
In Verdoorn, Plants and plant science in Latin America, pp. 121-125. 1945.

Sampaio, A. J. pe. Phytogeographia do Brasil, ed. 3, pp. 1-372. 1945.

ScHNEIWER, C. K. Die Gattung Berberis (Muberberis), Vorarbeiten fiir eine
Monographia. Bull. Herb. Boiss. 1. vol. 5, pp. 33-48, 133-148, 391-403,
449-464, 655-670, 800-831. 1904-05.

Scoacan, H. J. The flora of Bic and the Gaspé Peninsula, Quebec. Nat. Mus.
Canada, Bull. 115, pp. 1-399. 1950.

SLEUMER, H. Estudios sobre el genero ‘‘Dunalia” H. B. K. Lilloa, vol. 23, pp.
117-142. 1950.
Smiru, A. C., & Jounsron, I. M. A phytogeographie sketch of Latin America.
In Verdoorn, Plants and plant science in Latin America, pp. 11-18. 1945.
SmitH, Lyman B. Geographical evidence on the lines of evolution in the Bro-
meliaceae. Bot. Jahrb., vol. 66, pp. 446-468. 1934.

—-——~. Studies in the Bromeliaceae—vi. Proc. American Acad. Sci., vol. 70,
pp. 147-220. 1935.

———. The vegetation of Brazil. Jn Verdoorn, Plants and plant science in Latin
America, pp. 297-302. 1945.

———, The Bromeliaceae of Brazil. Smithsonian Mise. Coll., vol. 126, pp.
1-290. 1955.

~-—-—-. Notes on South American phanerogams—1. Journ. Washington Acad.
Sci., vol. 9, pp. 282-284. 1958a.

——-—, Notes on South American phanerogams—11. Journ. Washington Acad.
Sci., vol. 48, pp. 310-314. 1958b.

Situ, Lyman B., anp Downs, Ropert J. Resumo preliminar das Rubidceas
de Santa Catarina. Sellowia, No. 7, pp. 13-86. 1956.

—-——=, Resumo preliminar das Euforbidceas de Santa Catarina. Sellowia,
No. 11, pp. 155-231. 1959.

————, Resumo preliminar das Amaranticeas de Santa Catarina. Sellowia,
No. 12, pp. 99-120. 1960a.

~~-——-, Resumo preliminar das Cariofildceas de Santa Catarina. Sellowia,
No. 12, pp. 121-133. 1960b.

The Xyridaceae of Brazil. (In preparation.)

SMITH—ORIGINS, FLORA OF SOUTHERN BRAZIL 249

SraFLeu, F. A. A monograph of the Vochysiaceae, 1. Salvertia and Vochysia.
Rec. Trav. Bot. Néerl., vol. 41, pp. 397-540. 1948.

Swart, J. J. A monograph of the genus Protiwm and some allied genera (Bur-
seraceae). Rec. Trav. Bot. Néerl., vol. 39, pp. 211-446. 1942.

Wutrr, E. V. Introduction to the historical geography of plants. Bull. Appl.
Bot. Gen. & Pl. Breeding. Suppl. 52. (translation Brissenden, E.): An
introduction to historical plant geography. pp. 1-223. 1943.

Part 4

A SYNOPSIS OF THE AMERICAN VELLOZIACEAE

Lyman B. Smiru

Introduction

The present paper is a necessary corollary to my research on the
origin of the flora of southern Brazil that was supported by a grant
rom the National Science Foundation in 1956-57. Although none
of my 1956-57 collections are represented here, the Velloziaceae are
one of the best examples of the distribution of a family of consistently
heliophile species, and its extension southward from the planalto
of central Brazil provides a pattern to illustrate the movement of
many other groups.

The American species of Velloziaceae have not been organized
since Seubert’s treatment in the “Flora Brasiliensis” in 1847 and
then only sketchily and without complete keys. In 1937, Gocthart
and Henrard published many new species without any key whatever,
but this has only aggravated the problem of classifying new material.
Thus, in order to make the material available for phytogeographic
studies it is first necessary to revise it. A number of American
herbaria have been consulted and photographs of most of the types
of Goethart and Henrard have been obtained from the Berlin-Dahlem
herbarium through the kindness of Dr. G. M. Schulze and of Frau
Christa Metzger, who had them on loan from Leiden. Frau Metzger
is working on material in European herbaria with special emphasis
on leaf anatomy. The present paper was begun without knowledge
of her work, but since all novelties are based on material in American
herbaria it should prove largely supplementary.

In preparing the present paper, I am indebted to the curators
of the following herbaria for the opportunity to study the material in
their care:

Chicago Natural History Museum (F)
Gray Herbarium of Harvard University (GH)
Instituto Agronémico do Norte (IAN)
Museu Paraense “Emilio Goeldi’? (MG)
New York Botanical Garden (NY)
University of California (UC)
United States National Herbarium (US)
251

252 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

ixsiccatae have been cited only for novelties and new combinations,
but many that do not appear have been most helpful in making keys
and studying distribution. My knowledge of the older species has
been much increased by the type photographs of J. Francis Macbride
received from the Chicago Natural History Museum, and N. Y.
Sandwith has kindly communicated locality data of Gardner’s collec-
tions from his notebook at Kew.

Some of the earliest authors to treat the Velloziaceae, like Kunth,
confused the group with the Bromeliaceae, presumably on the basis
of habital resemblances. However, it is clear from the character
of the endosperm as well as of the perianth that the Velloziaceae are
much more closely allied to the Amaryllidaceae. In fact, under the
broad interpretation of the Amaryllidaceae, Baker was doubtless
more logical to include the Velloziaceae as a tribe, Vellozieae, than
was Pax to maintain it as a separate family. In a system such as
Hutchinson’s, the Velloziaceae, as distinguished by stalked placentas
and many-ranked ovules, are reasonably equivalent to the new units
derived from the division of the Liliaceae and Amaryllidaceae.

The primary subdivision of the Velloziaceae has been based on
three different sets of characters according to the authors involved.
Baker and Hutchinson used the presence or absence of an extension
of the perianth-tube above the ovary. This is a distinction which is
striking between extremes but which is so finely intergrading from
species to species in a complete survey that it is quite unworkable.
Pax used the distinction between 6 stamens and more than 6, and
since no species combining the two has yet been found, his system
can be said to be workable. However, it places under Barbacenia
species that are in every other respect indistinguishable from Vellozia,
and furthermore it is quite illogical because species with 9 stamens
have some tepals subtending a single stamen as in Barbacenia and
others subtending more than one as in Vellozia. Seubert used the
form of the filament, which is a clear and logical distinction with good
correlations. For instance, sharp cleavage of leaf-blades and broad
3-lobed stigmas are restricted to Vellozia by Seubert’s classification.
It is difficult to understand how later authors could have overlooked
or disregarded his system.

Velloziaceae

Drude in Schenck, Handb. Bot. 333. 1886; Pax in Engl. & Prantl, Pflanzenfam. 2,
Abt. 5: 125. 1887;ed. 2. 15a: 431. 1930; Hutchinson, Fam. Fl. Pl. ed.
2.2: 678. 1959.

Amaryllideae- Vellozieae D. Don, Edinb. New Philos. Journ. 8: 164. 1830; Baker
in Benth. & Hook. Gen. Pl. 3: 739. 1880.

Family Vellozieae Seubert in Mart. Fl. Bras. 3, pt. 1: 65. 1847.

SMITH—SYNOPSIS OF AMERICAN VELLOZIACEAE 953

Perennial plants; indument highly diverse, often not. strictly
epidermal (Greves, Journ. Bot. 59: 274, 1921); stems (caudex) woody,
fibrous, simple or dichotomously branched, covered with persistent
leaf-sheaths: complete leaves clustered at the end of the stem or its
branches, orass-like: scapes one or more, terminal becoming lateral by
the elongation of the axis; flowers solitary on cach peduncle, perfect,
actinomorphic, brightly colored; perianth-tube equaling to greatly
exceeding the ovary and adnate to it; tepals in 2 series but mostly
very similar, erect to reflexed at anthesis; stamens 6 to numerous
in 6 bundles; filaments terete and simple or flat with 2 lobes or
appendages at the summit, rarely lacking; anthers basifixed to medi-
fixed, opening by longitudinal slits; ovary 3-locular, inferior; style
slender but usually enlarged by the stigmas; ovules numerous in many
rows on stalked placentas; fruit a capsule; seeds numerous; embryo
small; endosperm copious, non-farinaceous.

1, Anthers basifixed; stigma broadly 3-lobed; leaf-blades deciduous along a
straight transverse line; stamens 6 or usually more, the filaments terete and
elongate, unappendaged above .. . ... . . IL. Vellozia

1. Anthers dorsifixed, although sometimes only slightly above the base; stigma
slightly if at all lobed: leaf-blades tardily and irregularly deciduous; stamens
always 6.

2, Filaments very short, terete, unappendaged, the anthers appearing sessile.

2. Barbaceniopsis

2. Filaments evident, flat with two lobes or appendages at the summit.
3. Barbacenia

1. Vellozia

Vellozia Vand. Fl. Lusit. & Brasil. Spec. 32, pl. 2. 1788.
Xerophyta Juss. Gen. 50. 1789.
Campderia A. Rich. Bull. Soe. Philom. 79. 1822.
Radia A. Rich. in Kunth, Syn. Pl. Aequin. 1: 300. 1822.
Schnizleinia Steud. ex Hochstetter in Flora 27: 31. 1844.
Talbotia Balf. in Trans. Bot. Soc. Edinb. 9: 190. 1868.

Plants from small stature to over 4 meters high; caudex mostly
few-many-branched; leaf-blades deciduous along a straight transverse
line; tepals free above the ovary or forming a tube; stamens 6, or
more numerous and in 6 bundles; filaments terete and filamentous.

Type species: Vellozia glabra Mikan.
Keys to the Species of Vellozia

1. Tepals free or obscurely short-connate above the ovary . . Section I. Vellozia
1. Tepals forming a distinct tube above the ovary . . . Section II. Radia

SECTION I. VELLOZIA

1. Ovary (or capsule) smooth and glabrous.
2. Plant nearly stemless; leaves 5-7 cm. long.
3. Stamens subequal; phalanges simple... .. . . 1. V. macedonis
3. Stamens very unequal; phalanges appendaged . 2. V. tenella

254 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

2. Plant distinctly caulescent.
4, Scapes less than 1 em. long; capsule obconic or truncate.
5. Stamens 6; leaf-blades obtuse or acute.

6. Leaf-blades acute, ciliate; sheaths pubescent. . . 3. V. abietina
6. Leaf-blades obtuse, serrulate. ....... 4 Y~ tragacantha
5. Stamens more than 6; leaf-blades retuse. . . . . . . 5. V. pusilla

4. Scapes much more than 1 cm. long.
7. Caudex simple or subsimple; leaf-blades only 2.5 em. long.
6. V. sellovii
7. Caudex branched; leaf-blades at least 8 cm. long.
8. Stamens 6; leaf-blades lanceolate, widest at the middle.
7. V. plicata
8. Stamens more than 6; leaf-blades linear or very narrowly triangular.
9, Capsule alate-trigonous, broadest at the middle, dull; leaf-blades
filiform-acuminate, coarsely brown-ciliate near the base.

8. V. alata
9, Capsule terete to trigonous but not alate, broadest at apex, lustrous.
10. Seapes erect; leaves filiform-acuminate. . . . . 9. V. glabra

10. Scapes recurved; leaves broadly acute to retuse.
10. V. incurvata
1. Ovary (or capsule) at least partially vestite.
11. The ovary vestite primarily on the angles or base.
12. Trichomes of the ovary acute, making the angles scabrous, concolorous
with the ovary.
13. Tepals 60-70 mm. long; leaf-blades obtuse to subacute; old leaf-sheaths
mostly entire, 2.0.0.0. 2 ee ee ee ee 0 V. compacta
13. Tepals 30 mm, long.
14. Old leaf-sheaths mostly entire; leaf-blades acuminate (?).
12. V. verruculosa
14. Old leaf-sheaths splitting into coarse fibers; leaf-blades obtuse to

subacute... ee ee ee ee ee ee ee 6. Y.~ fibrosa
12. Trichomes of the ovary rounded, glandular in character.
15. Leaf-blades acuminate; tepals 40-60 mm. long. . . 14. V. variabilis

15. Leaf-blades subacute to retuse.
16. Glands of the ovary stipitate; leaf-sheaths splitting into fibers.
17. Fibers of the leaf-sheaths straight (?). . . . . 15. V. ambigua
17. Fibers of the leaf-sheaths incurved. . . . . . 16. V. martiana
16. Glands of the ovary sessile.
18. Tepals obtuse and submucronulate; leaf-blades 9-15 mm. wide.
17. V. wettsteinii
18. Tepals acute; leaf-blades not more than 3 mm. wide.
19. Leaf-blades to 18 em. long, 3 mm. wide . 18 V. angustifolia
19. Leaf-blades less than 4 em. long; about 1 mm. wide.
19. V. virgata
11. The ovary completely vestite or laxly and evenly vestite.
20. Indument of the ovary glandular in character.
21. Leaves acuminate or subulate-acuminate.
22. Capsule oblong.
23. Leaf-blades narrow, not over 6 mm, wide; bases of the glands
stout, confluent . 2... . 4... .... . . 20. V. echinata

SMITH—SYNOPSIS OF AMERICAN VELLOZIACEAE 255

23. Leaf-blades 10 mm. or broader.
24, Margins of the leaf-blades pilose; stamens 15.
21. V. fimbriata
24. Margins of the leaf-blades merely scabrous; stamens 30-36.
22. V. intermedia
22. Capsule (or ovary) ovoid, globose, or hemispheric.
25. Ovary broad at apex, hemispheric or broadly truncate.
26. Margins of the leaf-blades pilose; stamens 30.
23. V. barbaceniifolia
26. Margins of the leaf-blades serrulate; stamens 18.
24. V. hemisphaerica
25. Ovary contracted at apex, ovoid, or globose.
27. Scape less than 2 em. long; tepals obovate, rounded, 50 mm.
long 2... . 2. 2... 625. VY. breviscapa
27. Seape elongate.
28. Leaf-blades about 3 mm. wide; caudex slender with few
ranked laxly imbricate leaf-sheaths. . . . 26. V. squalida
28. Leaf-blades 15 mm. or broader; caudex stout with many-
ranked very densely imbricate leaf-sheaths.
29. Tepals 75 mm. long; leaf-blades pubescent beneath; ovary

densely glandular-hirsute . . . . . . . 27. V. aloifolia
29. Tepals 45 mm. long; leaf-blades glabrous; ovary subdensely
and coarsely stipitate-glandular . .. . 28. V. viannae

21. Leaves broadly subacute, obtuse or emarginate.
30. Margins of the leaf-blades entire and glabrous.
31. Ovary rufescent-pilose; leaf-blades gray villous beneath.
29. V. glandulifera
31. Ovary scabrous-glandular; leaf-blades very glutinous.
30. V. scoparia
30. Margins of the leaf-blades serrate, setose, or pilose.
32, Leaf-blades wholly white-pilose, 1.5 mm. wide, ecarinate.
31. V. pilosa
32. Leaf-blades incompletely vestite if at all.
33. Glands of the ovary sessile or subsessile; capsule widest near

the apex.
34. Stamens 6; peduncle less than 1 cm.long. . 32. V. taxifolia
34. Stamens 18-30; peduncle much longer than 1 em.
35. Tepals 2 em. long; stamens 18 .. . . 33. V. declinans
35. Tepals 4 cm. long; stamens 30 (?) . . . 16. V. martiana

33. Glands of the ovary stipitate.
36. Sheaths ciliate throughout with stiff spreading setae.
34. V. barbata
36. Sheaths entire or with a few teeth toward apex.
37. Capsules trigonous or oblong.
38. Sheaths aculeolate toward apex; capsules trigonous,
12-costate; tepals linear-lanceolate, acute.
35. V. pleurocarpa
38. Sheaths entire; capsules oblong; tepals linear-spathulate.
36. V. leptopetala
37. Capsules ellipsoid or globose.
39. Leaf-sheaths densely white-lanate at apex; blades sub-
densely setose beneath; tepals 40-50 mm. long.
37. V. resinosa

256 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

39. Leaf-sheaths glabrous; tepals smaller.
40. Sheaths wholly covered; caudex very short and stout.
38. V. brevifolia
40. Sheaths much exposed toward their apices; caudex

slender.
41. Plants of medium stature, well over 25 em. high;
leaf-blades serrulate . . . .39. V. epidendroides

41. Plants dwarfed, less than 25 cm. high.
42. Leaf-blades straight, 1 mm. wide, laxly vestite

with dark sessile glands. . . . 40. V. minima
42. Leaf-blades contorted, 3 mm. wide, subdensely
setose .. . .... . 41. ¥. streptophylla

20. Indument of the ovary punctulate, acute, or sharply truncate, not
glandular in character.
43, Trichomes of the ovary minute, making it scabrous or granulate and
generally exposing its surface in between.
44. Scape none; capsule hidden in the leaf-sheaths, obconic.
45. Leaf-blades 2.5 cm. long, emarginate . . . . . 32. V. taxifolia
45. Leaf-blades 9-10 em. long, subobtuse. . . . 42. V. cryptantha
44. Scape evident.
46. Ovary ovoid or ovoid-globose.
47. Caudex to 2 meters high, much branched; leaf-blades deeply
retuse, 5 mm. wide; old sheaths not splitting into fibers.
43. V. ramosissima
47. Caudex not over 20 cm. high, simple or few-branched.
48. Leaf-blades obtuse; old sheaths densely imbricate, splitting;

caudex very short... . . .... . 44 V. asperula
48. Leaf-blades retuse; old sheaths much exserted, entire; caudex
to 20cm. high . . .... . 36. V. leptopetala

46. Ovary oblong or oblong-ellipsoid.
49. Tepals 40-50 mm. long.
50. Scape nearly glabrous; ‘epals long-attenuate toward base,

white (?).. . . . . 45. V. punctulata
50. Scape strongly vestite on the upper half; tepals lanceolate,
blue... . tee ew ee es 46, V. bradei

49. Tepals 10-20 mm. ‘long.
51. Margins of the blades entire; sheaths remaining entire;

blades 12 mm. wide, glaucous beneath . . 47. V. piresiana
51. Margins of the blades serrulate; sheaths splitting into coiled
fibers.

52. Leaf-blades 3-4 mm. wide; tepals 20 mm. long.
48. V. ornata

52. Leaf-blades to 7 mm. wide; tepals 10 mm. long.
49. V. granulata

43. Trichomes of the ovary coarse, completely covering it.
53. Leaf-blades acuminate or subulate-acuminate.
54. Scape glabrous or at most slightly scabrous at apex, contrasting
sharply with the densely vestite ovary.

55. Trichomes of the ovary broadly truncate with contiguous
apices. 2... 2 ee ee ee ee 50. V. flavicans

SMITH—SYNOPSIS OF AMERICAN VELLOZIACEAE 257

55. Trichomes acute or obtuse and laterally flattened but not
truncate.

56. Trichomes of the ovary short, ovoid; leaf-sheaths entire.

51. V. glauca

56. Trichomes of the ovary elongate, subulate or laterally

flattened.

57. Leaf-sheaths remaining entire.
58. Blades strongly sulcate . . . . . . 52. V. glochidea
58. Blades nearly even . . . .. . . 53. V. gardneri

57. Leaf-sheaths splitting into fibers.
59. Ovary-trichomes subulate, acute; leaf-blades glabrous

beneath except the keel . . . . . 54. V. erassicaulis
59. Ovary-trichomes much flattened laterally, obtuse; leaf-
blades villous beneath. . . . . . . 55. V. sulphurea

54. Scape strongly vestite on the upper third or half, not contrasting
with the ovary.
60. Leaf-blades 15 mm. wide; tepals 80-120 mm. long; seape muri-
cate toward apex.
61. Scape 7 em. long; ovary-trichomes abruptly attenuate.
56. Y. seubertiana
61. Scape to 40 em. long; ovary trichomes sharply truncate.
57. V. froesii
60. Leaf-blades 4-8 mm. wide (uncertain in V. pumila); scape
2-4.5 cm. long above the leaf-sheaths.
62. Tepals 20mm. long... .... . . . 58. V. hypoxoides
62. Tepals 40-80 mm. long.
63. Sheaths splitting into fibers.
64. Caudex ovoid, 4 cm. long; leaf-margins and keel sub-
densely aculeolate; tepals 40 mm. long. 59. V. pumila
64. Caudex cylindric, elongate; leaf-margins and keel entire

and glabrous; tepals 80 mm. long. . . 60. V. swallenii
63. Sheaths remaining entire; plant stemless; tepals 60 mm.
long. ... ... . . 61. V. bulbosa

53. Leaf-blades broadly subacute, ‘obtuse, or rT emarginate.
65. Scape glabrous, contrasting sharply with the densely vestite ovary.
66. Leaf-sheaths remaining entire; ovary subglobose.
62. V. cinerascens
66. Leaf-sheaths splitting into coarse fibers; ovary clavate-cylindric,
truncate. . ... ... . . 63. V. circinans
65. Scape strongly vestite toward : apex.
67. Trichomes of scape and ovary fine, filamentous.
64. V. dasypus
67. Trichomes of scape and ovary much coarser.
68. Ovary prismatic, its trichomes subulate, more or less
twisted ....., .... . , 65. V. glaziovii
68. Ovary terete or if trigonous, broader at center, base, or apex.
69. Plant essentially stemless.
70. Leaf-blades ciliate-serrate . . . . . 66. V. crinita
70. Leaf-blades merely serrate; ovary acutely trigonous.
67. V. caruncularis

258 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

69. Plant long-caulescent.
71. Scape about equaling the leaves, ca. 13 cm. long.
68. V. gracilis
71. Scape much exceeded by the leaves.
72. The scape trigonouss ovary-trichomes in part
bicuspidate .... .. . . . 69. V. grisea
72. The scape terete; ovary- trichomes seule,
. V. exilis

SECTION If. RADIA

1. Scapes not over 3 em. long.
2. Tepals 2 em. long; leaf-blades 8-5 cm. long; flowers numerous, sessile.
71. V. uaipanensis
2. Tepals 8 em. or longer; leaf-blades to 27 em. long; flowers solitary.
8 Flowers sessile. 2... ee ee ee ee ee ee TH. VV. markgrafii
3. Flowers scapose.
4. Leaf-blades subappressed-pilose beneath, 4mm. wide . 73. V. riedeliana
4. Leaf-blades glabrous except for the ciliate keel and margins, 10 mm.
wide 2... ee ee ee ee ee ee eee Th. Ve maguire
1. Scapes elongate.
5. Perianth-tube not more than 30 mm. long above the ovary.
6. Seape 6-l5em. long... 6. ee ee ee ee ee TS V. candida
6. Seape over 20cm. long... .. . . . +. + + + . 76. V. macrantha
5. Perianth-tube 35-80 mm. long.
7. Leaf-blades 4-7 mm, wide at most.
8. Apex of leaf-blade obtuse; sheaths soon divided into coarse fibers.
77. V. leucanthos
8. Apex of leaf-blade filiform-attenuate.
9, Perianth-tube 35-50 mm. long.
10. Scapes about equaling the leaves; caudex rarely to 4 cm. long.
78. V. cachimbensis
10. Scapes not more than half as long as the leaves; caudex 50-100
cm. long.
11. The perianth-tube 35-40 mm. long. . . 79. V. macarenensis
11. The perianth-tube 45-50 mm, long . . . 80. V. macrosiphonia
9. Perianth-tube 60-80 mm. long.
12. Flowers exceeding the leaves; leaf-sheaths spotted with dark
brown... .. . . .8l. V. maculata
12, Flowers about equaling the leaves s or - shorter.
13. Leaf-blades glabrous beneath; branches including old leaf-
sheaths not over 1 em. in diameter.
14, Trichomes of the ovary ¥ subglobose with a minute black gland
ontheapex. ... .. . . . 82. V. machrisiana
14, Trichomes of the ovary “attenuate toward apex with a broad
peltate gland. . . .. . . . 83. V. uleana
13. Leaf-blades lepidote- _pilose. be nee ith: branches imcluding old
leaf-sheaths 2 cm. or more in diameter . . . (94. V. tubiflora)
7. Leaf-blades 8-20 mm, wide.
15. Seapes less than 10 em, long.
16. Trichomes of the ovary attenuate, the glands distinctly stipitate;
scapes 4-6 cm. long. . . ... ... . . . 84. V. panamensis

SMITH—SYNOPSIS OF AMERICAN VELLOZIACEAE 259

16. Trichomes of the ovary ovoid, the glands subsessile; scapes 7-9
cm. long.
17. Caudex simple or few-branched, 1-2 meters high.
85. V. phantasmagoria
17. Caudex much branched.
18. Perianth-tube densely glandular. . . . . . 86. V. dumitiana
18, Perianth-tube very sparsely glandular . . 87. V. rhynchocarpa
15. Seapes 10-30 cm. long.
19. Leaf-sheaths essentially glabrous; tepals rounded; perianth-tube

75 mm, long. ... . . . . . 88. V. maudeana
19. Leaf-sheaths densely vestite ‘at least bet ween the nerves.
20. Apex of leaf-blades rounded... .... . .89. V. annulata

20. Apex of leaf-blades filiform-acuminate.
21. Flowers exceeding the leaves; leaf-blades densely vestite on
both sides, 20 em. long.
22. Leaf-blades 8 mm. wide, flowers solitary . .90. V. velutinosa
22, Leaf-blades 17 mm. wide; flowers at least 3 together.
91. V. dawsonii
21. Flowers not exceeding the leaves.
23. Leaf-blades lepidote-pilose on both sides; perianth-tube

65mm.long. . . . ee ee es. 2 92. Ve cana

23. Leaf-blades glabrous: above.
24. Perianth-tube ca.40mm.long. .. . .93. V. lithophila
24. Perianth-tube 45-80 mm. long... . .94. V. tubiflora

Section I. Vellozia

1. Vellozia macedonis Woodson, Ann. Mo. Bot. Gard. 37: 398. 1950.
Braziu: Minas Gerais: Mun. Ouro Preto: Saramenha.
2. Vellozia tenella Mart. ex Schult. f. in R. & S. Syst. 7: 293. 1826.
Vellozia graminea Pohl, Pl. Bras. 1: 118, pl. 93. 1827.
Brazit: Minas Gerais: Inficionado. Vila Rica (Ouro Preto).
3. Vellozia abietina Mart. Nov. Gen. & Sp. 1: 14, pl. 6. 1823.
Xerophyta abietina Spreng. Syst. Veg. 4: Cur. Post. 187. 1827.
Braziu: Minas Gerais: Itambé. Serra da Lapa.
4. Vellozia tragacantha Mart. ex Seubert in Mart. Fl. Bras. 3, pt. 1: 75. 1847.
Xerophyta tragacantha Mart. ex Schult. f. in R. & S. Syst. 7: 290. 1826.
Brazit: Minas Gerais: Villa do Rio das Contas. Diamantina. Serra de
Caraga.
5. Vellozia pusilla Pohl, Pl. Bras. 1: 122, pl. 97. 1827.
Braziu: Minas Gerais: Itambé. Serra do Cipé. Diamantina.

Note: Leaves and habit identical with those of V. tazitfolia, but
stamens numerous and ovary smooth.
6. Vellozia sellovii Seubert in Mart. Fl. Bras. 3, pt. 1: 75. 1847.

Braziu: No locality. (?) Minas Gerais: Serra da Piedade.

q. Vellozia plicata Mart. Nov. Gen. & Sp. 1: 16, pl. 9. 1823.
Xerophyta plicata Spreng. Syst. Veg. 4: Cur. Post. 187. 1827.
Braziu: Bahia: Serra de Monte Santo.

597430—62——-4

260 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

8. Vellozia alata L. B. Smith, sp. nov. PLATE 1

A V. glabra Mikan, cui affinis, laminis foliorum basi grosse brun-
neociliatis, capsula alato-trigona viride vix lucida differt.

Caudex simple (?), terete and 5 cm. in diameter including the old
leaf-sheaths; leaves many-ranked, very densely imbricate, the com-
plete ones about 40 at the apex of the stem; sheaths very closely
nerved, suboblong, 7 cm. long, glabrous, the apices recurved with
age but remaining entire; blades linear, filiform-acuminate, 4 dm.
long, 12 mm. wide at base, densely and strongly nerved on both
sides, otherwise even, coarsely brown-ciliate on the slightly thickened
margins but not on the inconspicuous keel, otherwise glabrous; scapes
few or solitary, erect or decurved, to 16 cm. long, trigonous, glabrous;
capsule broadly ellipsoid, 5 cm. long, 3 em. wide, alate-trigonous,
sublustrous, yellow-green when dry; stamens 12 or more; style 5
em. long.

Type in the U.S. National Herbarium, No. 2340342, collected 334 miles from
Hotel Chapeu do Sol, Serra do Cipé, State of Minas Gerais, Brazil, altitude 1110
meters (3700 feet), December 19, 1959, by Bassett Maguire, C. K. Maguire and
J. Murga Pires (No. 44690). Isotype in the New York Botanical Garden.

9. Vellozia glabra Mikan, Delect. Fl. & Faun. Brasil. fase. 2. 1820; Spreng.
Syst. Veg. 3: 338. 1826.
Vellozia Vand. Fl. Lusit. & Brasil. Spec. 32, pl. 2. 1788. Without indica-
tion of species.
Vellozia phalocarpa Pohl, Pl. Brasil. 1: 123, pl. 98. 1827.
?Vellozia crassirama Goeth. & Henr. Blumea 2: 368. 1937.
Braziu: Minas Gerais: Serra do Cipé, Diamantina, Sérro. Itambé. Goids.
Bolivia.
10. Vellozia incurvata Mart. ex Schult. f. in R. & S. Syst. 7: 293. 1826.
Braziut: Minas Gerais.
11. Vellozia compacta Mart. ex Schult. f. in R. & S. Syst. 7: 293. 1826.
Vellozia compacta var. obtusiflora and var. acutiflora Seubert in Mart. FI.
Bras. 3, pt. 1: 77. 1847.
Braziu: Minas Gerais: Piedade, Nova Lima. Serra do Cipé.
12. Vellozia verruculosa Mart. ex Schult. f. in R. & 8, Syst. 7: 293. 1826.
BraziLt: Minas Gerais, Goids: Corumbdé. Natividade. Mato Grosso.
13. Vellozia fibrosa Goeth. & Henr. Blumea 2: 370. 1937.
Braziu: Minas Gerais: Biribiri, near Diamantina.
14. Vellozia variabilis Mart. ex Schult. f. in R. & S. Syst. 7: 293. 1826.
4a. Vellozia variabilis var. variabilis
? Vellozia alutacea Pohl, Pl. Bras. 1: 130. 1827.

Capsule tuberculate on the angles only.

Braziut: Minas Gerais: Lagoa Grande. Goids: Serra Dourada.

14b. Vellozia variabilis var. tuberculata Seubert in Mart. Fl. Bras. 3, pt. 1:
77. 1847.

Capsules with a row of tubercules on each side as well as on angles.

Braziu: Minas Gerais: Nova Lima. Caldas.

SMITH—SYNOPSIS OF AMERICAN VELLOZIACEAE 261

15. Vellozia ambigua Goeth. & Henr. Blumea 2: 364. 1937.
Braziu: Minas Gerais: Itacolumi.

16. Vellozia martiana Goeth. & Henr. Blumea 2: 376. 1937.
Brazit: Minas Gerais: Pico Itabira do Campo.

17. Vellozia wettsteinii Goeth. & Henr. Blumea 2: 383. 1937.
Braziu: Sao Paulo: Between Sacramento and Jaguara.

18. Vellozia angustifolia Goeth. & Henr. Blumea 2: 365. 1937.
Braziu: Goids (?): Morro Cubatéio. Minas Gerais: Diamantina.

19. Vellozia virgata Goeth. & Henr. Blumea 2: 382. 1937.
Braziu: Minas Gerais: Sio José (Jodo?) del Rei. Pico Itabira do Campo.

20. Vellozia echinata Goeth. & Henr. Blumea 2: 369. 1937,
Braziu: Minas Gerais: Caraga.

21. Vellozia fimbriata Goeth. & Henr. Blumea 2: 371. 1937.
Brazit: Minas Gerais: Serra da Lapa.

22. Vellozia intermedia Seubert in Mart. Fl. Bras. 3, pt. 1: 78. 1847.
BraziL: Without locality. ?Minas Gerais: Serra do Cipé.

23. Vellozia barbaceniifolia Seubert in Mart. Fl. Bras, 3, pt. 1: 79. 1847.
Braziu: Without locality.

24. Vellozia hemisphaerica Seubert in Mart. Fl. Bras. 3, pt. 1: 80. 1847.
Braziu: Bahia.

25. Vellozia breviscapa Mart. ex Schult. f. in R. & S. Syst. 7: 293. 1826.
Brazit: Minas Gerais: Diamantina.

26. Vellozia squalida Mart. ex Schult. f. in R. & S. Syst. 7: 292. 1826.
BraziL: Minas Gerais: Serra do Cip6.

27. Vellozia aloifolia Mart. Nov. Gen. & Sp. 1: 15, pl. 7. 1823.
Brazit: Minas Gerais: Serra de Itambé. Tejuco. Diamantina.

28. Vellozia viannae L. B. Smith, sp. nov. PLATE 9, FIGURE 1

A V. variabile Mart. ex Schult. f., cui affinis, ovario omnino regu-
lariterque glanduloso differt.

Caudex simple (?), terete and 5 cm. in diameter (including the old
leaf-sheaths); leaves many-ranked, very densely imbricate; leaf-
sheaths oblong, over 4 cm. long, very densely nerved, dark castaneous
with a thin stramineous margin, the apices recurved with age; leaf-
blades over 20 before falling, linear, filiform-acuminate, imperfectly
known but probably 5 dm. long, 15 mm. or wider, setose-serrate on
the scarcely thickened margins, otherwise glabrous, densely nerved,
the keel broad but not prominent; scapes slender, 12-15 cm. long,
densely dark-glandular toward apex; ovary ellipsoid, 8-10 mm. long,
densely and finely dark-glandular throughout; tepals elliptic, rounded
and apiculate, 45 mm. long, stamens ca. 18, the phalanges appendaged ;
anthers subequal, 11 mm. long.

Type in the U.S. National Herbarium, No. 2323056, collected 6 kilometers
north of Palacio, Serra do Cip6, Municipio of Jaboticatubas, State of Minas

Gerais, Brazil, October 1953, by F. Segadas-Vianna and J. Lorédo, Jr. (No.
Serra-II, 1103).

262 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

29. Vellozia glandulifera Goeth. & Henr. Blumea 2: 372. 1937.
Brazit: Minas Gerais: Cachoeira.
30. Vellozia scoparia Goeth. & Henr. Blumea 2: 380. 1937.
Braziu: Minas Gerais: Parauna and Tejuco. Cachoeira.
31. Vellozia pilosa Goeth. & Henr. Blumea 2: 377. 1937.
Braziu: Minas Gerais: Diamantina.
32. Vellozia taxifolia (Mart. ex Schult. f.) Mart. ex Seubert in Mart. Fl. Bras.
3, pt. 1: 75. 1847.
Xerophyta taxifolia Mart. ex Schult. f. in R. & S. Syst. 7: 291. 1826.
Brazit: Minas Gerais: Serra do Cipé. Serra Frio.
33. Vellozia declinans Goeth. & Henr. Blumea 2: 369. 1937.
Braziu: Minas Gerais: Serra da Lapa.
34. Vellozia barbata Goeth. & Henr. Blumea 2: 366. 1937.
Braziu: Minas Gerais: Diamantina.
30. Vellozia pleurocarpa Goeth. & Henr. Blumea 2: 377. 1937.
Braz: Minas Gerais: Biribiri near Diamantina.
36. Vellozia leptopetala Goeth, & Henr. Blumea 2: 374. 1937.
Braziu: Minas Gerais: Biribiri near Diamantina.

37. Vellozia resinosa Mart. ex Schult. f. in R. & S. Syst. 7: 293. 1826, emend.
L. B. Smith.

Ab omnibus speciebus sectionis Velloziae vaginis foliorum apice
albo-lanatis, laminis subtus subdense setosis, et ovario dense glanduloso
differt.

Caudex simple, very short, terete and 2 cm. in diameter (including
the old leaf-sheaths) ; leaves densely many-ranked; sheaths suboblong,
35 mm. long, dark castaneous with a pale thin margin, lustrous, the
exposed apex densely white-lanate, remaining entire; blades about 12
before falling, suberect, linear, subacute, more or less complicate, 15
cm. long, 10 mm. wide, glabrous above, subdensely setose beneath,
ciliate; scapes 1-2, slender, 13-15 cm. long, densely dark-glandular
toward apex; ovary globose, densely dark-glandular; tepals elliptic,
obtuse, 50 mm. long, purple; stamens ca. 12, 15 mm. long.

Braziu: Minas Gerais: Martius (M, type; F, photo no. 18982). Serra do

Cipé, between km. 111 and 128, alt. 1140 m., December 20, 1959, Maguire 4:
Pires 44694 (NY, US).

38. Vellozia brevifolia Seubert in Mart. Fl. Bras. 3, pt. 1:84, pl. 10, fig.2. 1847.
Braziu: Without locality.
39. Vellozia epidendroides Mart. ex Schult. f. in R. & S. Syst. 7: 292. 1826.
?Vellozia variegata Goeth. & Henr. Blumea 2: 381, 1937.
Braziu: Minas Gerais: Serra do Cip6. Diamantina. Carangola.
40. Vellozia minima Pohl, Pl. Bras. 1: 119, pl. 94. 1827.
Braziu: Minas Gerais: Itambé. Serra do Cipé6. Diamantina. Guinda.
41. Vellozia streptophylla L. B. Smith, sp. nov.
PLaTE 2; PLATE 9, FIGURES 2, 3

A V. minima Pohl, cui affinis, caulibus validioribus, laminis foliorum
valde contortis latioribus longe setosis differt.

SMITH—SYNOPSIS OF AMERICAN VELLOZIACEAE 263

Caudex erect, much branched, less than 2 dm. high, 7 mm. in
diameter; leaves more than 3-ranked, densely imbricate; sheaths
very closely nerved, glabrous, erect, entire in age; blades linear,
narrowly obtuse, 3 cm. long, 3 mm. wide, subdensely setose, at first
erect but recurved and contorted with age, long-persistent; scapes
solitary but soon axillary and long-persistent, 25 mm. long, very
slender, glandular especially toward apex; capsule globose, 5-6 mm.
long, subdensely and evenly glandular; tepals, stamens, and style
unknown.

Type in the New York Botanical Garden, collected in the vicinity of Diaman-
tina, State of Minas Gerais, Brazil, 1840, by George Gardner (No. 5233).

42. Vellozia cryptantha Seubert in Mart. Fl. Bras. 3, pt. 1: 80. 1847.
Brazit Minas Gerais: Diamantina.

43, Vellozia ramosissima L. B. Smith, sp. nov. PLATE 9, FIGURES 4, 5

V. asperula Mart. atque V. leptopetala Goeth. & Henr. affinis, a
priore foliis retusis, earum vaginis vetustis integris, a posteriore
foliorum vaginis vix exsertis, a ambobus caudice maxima ramosissima,
foliorum laminis latioribus distinguenda.

Caudex to 2 meters high, slender, the branches 1 cm. in diameter
including the old leaf-sheaths; leaves many-ranked, very densely
imbricate, the complete ones about 10 at the apex of each branch;
sheaths very closely nerved, broadly ovate, amplexicaul, ca. 4 cm.
long, glabrous, remaining entire; blades linear, deeply and asym-
metrically retuse, 7-9 cm. long, 5 mm. wide, densely nerved, obscurely
serrulate on the margins and keel; scapes solitary, about equaling
the leaves, decurved at apex with age, slender, trigonous, deeply
sulcate, scabrous especially toward apex; capsule stoutly ellipsoid,
12-14 mm. long, subdensely scabrous with fine pale subacute tri-
chomes; tepals linear-spatulate, acute, 23 mm. long; stamens 15-20.

Type in the U.S. National Herbarium, No. 2340340, collected on the Serra do
Cip6 2% miles from Hotel Chapeu de Sol, Minas Gerais, Brazil, altitude 1110

meters, December 19, 1959, by Bassett and Celia K. Maguire and J. Murga Pires
(No. 44666). Isotype in the New York Botanical Garden.

44, Vellozia asperula Mart. Nov. Gen. & Sp. 1: 15, pl. 8. 1823.

44a. Vellozia asperula var. asperula
Vellozia papillosa Pohl, Pl. Bras. 1: 128. 1827.

Leaf-blades 12-20 cm. long, about equaling the flowers to somewhat
exceeding them, 3-4 mm. wide.

Braziu: Minas Gerais: Tres Barras. Tejuco. Itambé.
44b. Vellozia asperula var. filifolia L. B. Smith, var. nov.

A var. asperula foliis valde elongatis flores superantibus, laminis
angustissimis differt.

264 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Leaf-blades over 30 cm. long, much exceeding the flowers, only 1
mm, wide.
Type in the New York Botanical Garden, collected on the Serra do Caraga,

State of Minas Gerais, Brazil, January 18, 1921, by F. C. Hoehne (No. 5025 in
hb. SP).

45. Vellozia punctulata Seubert in Mart. Fl. Bras. 3, pt. 1: 82. 1847.
Braziu: Bahia.

46. Vellozia bradei Schulze-Menz ex Markgraf, Notizblatt 15: 215. 1940.
Braziu: Minas Gerais: East of Montes Claros and north of Grio Mogul.

47. Vellozia piresiana L. B. Smith, sp. nov. PLATE 9, FIGURES 6, 7

A V. ornata Mart. ex Schult. f., cui affinis, foliorum vaginis vix
fissis, laminis integris latis differt.

Caudex branching, terete and 3 cm. in diameter (including the
old leaf-sheaths); leaves many-ranked, very densely imbricate; leaf-
sheaths 4 cm. long, stramineous, lustrous, remaining entire, the
nerves nearly contiguous; blades about 15 before falling, erect, linear,
attenuate to an obtuse asymmetric apex, 18 cm. long, 12 mm. wide,
glabrous, glaucous beneath, the thickened margins entire; scapes
slender, to 13 cm. long above the leaf-sheaths, scabrous especially
toward apex; ovary ellipsoid, subdensely scabrous; tepals 20 mm.
long, violet (! Black & Pires); stamens 15 (?); ovary 13 mm. long.

Type in the Instituto Agronémico do Norte, No. 69981, collected on the Serra

do Cipé at kilometer 132, State of Minas Gerais, Brazil, April 4, 1951, by G. A.
Black and J. M. Pires (No. 51-12157).
48. Vellozia ornata Mart. ex Schult. f. in R. & S. Syst. 7: 293. 1826.
Braziu: Minas Gerais.
49. Vellozia granulata Goeth. & Henr. Blumea 2: 373. 1937.
Braziu: Minas Gerais: Curalinho near Diamantina. Biribiri near Diamantina.
50. Vellozia flavicans Mart. ex Schult. f. in R. & S. Syst. 7: 293. 1826.
Vellozia squamata Pohl, Pl. Bras. 1: 124, pl. 99. 1827.
Brazit. Minas Gerais: Itambé. Goids: Serra Dourada. Anapolis.
51. Vellozia glauca Pohl, Pl. Bras. 1: 125, pl. 100. 1827.
dla. Vellozia glauca var. glauca
Vellozia glauca var, genuina Seubert in Mart. Fl. Bras. 3, pt. 1: 79. 1847.
Vellozia hamosa Pohl ex Seubert in Mart. Fl. Bras. 3, pt. 1: 79. 1847.
Nomen.

_ Scape elongate, completely glabrous.
Brazit: Goids: Aldea Carretéo de Pedro Terceiro. Chapada dos Veadeiros.
Slb. Vellozia glauca var. cujabensis Seubert in Mart. Fl. Bras. 3, pt. 1:79. 1847.
Scape short, furfuraceous-hirsute toward apex.

Braziu: Mato Grosso: Cuyaba.

52. Vellozia glochidea Pohl, Pl. Bras. 1: 129. 1827.
Braziu: Goids: Serra Dourada. Rio Aragudia,

53. Vellozia gardneri Goeth. & Henr. Blumea 2: 371, 1937.
Braziu: Goids: Natividade.

SMITH—SYNOPSIS OF AMERICAN VELLOZIACEAE 265

54. Vellozia crassicaulis Mart. ex Schult. f. in R. & S. Syst. 7: 292. 1826.
Vellozia albiflora Pohl, Pl. Bras. 1: 121, pl. 96. 1827.
Brazit: Minas Gerais: Arraial da Nossa Senhora de Penha. Mendanha.

55. Vellozia sulphurea Pohl, Pl. Bras. 1: 120, pl. 95. 1827.
Braziu: Minas Gerais: Arraial de SAo Jo&io Baptista. Between Tapanhoa-
canga and Padre Bento.

56. Vellozia seubertiana Goeth. & Henr. Blumea 2: 380. 1937.
Brazit: Mato Grosso: Cuyaba; Buriti in the Serra da Chapada.

57. Vellozia froesii L. B. Smith, sp. nov. PLATE 3

A V. seubertiana Goeth. & Henr., cui verisimiliter affinis, pedunculo
elongato, muricibus ovarii acute truncatis differt.

Caudex and leaf-sheaths unknown but undoubtedly large; leaf-
blade linear, filiform-acuminate, to 75 cm. long, 15 mm. wide, an-
trorsely spinulose-serrate on the pale margins, glabrous; scapes slender,
20-40 cm. long, densely spinulose toward apex; ovary ellipsoid, terete,
20 mm. long, completely covered with slenderly cylindric sharply
truncate trichomes; tepals elliptic-spatulate, to 10 cm. long; stamens
numerous, shorter than the style; style 7 cm. long.

Type in the Instituto Agronédmico do Norte, No. 15,405, collected at Mucugé,

Serra da Sincor4, State of Bahia, Brazil, February 1943, by Ricardo de Lemos
Frées (No. 19980). Same, Frées 19979 (NY, paratype).

58. Vellozia hypoxoides L. B. Smith, sp. nov. PLATE 9, FIGURE 8

A V. pumila Goeth. & Henr., cui verisimiliter affinis, foliorum
marginibus carinaque ciliatis, tepalis subduplo minoribus differt.

Caudex simple or once-branched, 4-10 cm. long, terete and 1 cm.
in diameter (including the old leaf-sheaths); leaves densely few-
ranked; leaf-sheaths ca. 2 cm. long, dull, with age splitting into coarse
fibers; leaf-blades about 10 before falling, recurving, linear, acuminate,
10-11 cm. long, 4 mm. wide, mostly conduplicate, finely ciliate on
the margins and keel, otherwise glabrous, densely nerved on both
sides; scapes 1-3, very slender, 3-5 cm. long above the leaf-sheaths,
densely and finely contorted-setose toward apex; ovary ellipsoid, 5
mm. long, covered with subulate contorted stramineous trichomes;
tepals elliptic, obtuse, 2 cm. long, violet; stamens ca. 15, phalanges
unappendaged; capsule to 10 mm. long.

Type in the U.S. National Herbarium, No. 2249220, collected on the Serra

dos Pirineus, Municfpio of Corumbé4, State of Goids, Brazil, February 17, 1956,
by Amaro Macedo (No. 4330). Isotype in the Instituto Agronémico do Norte.

59. Vellozia pumila Goeth. & Henr. Blumea 2: 378. 1937.
Braziu: Goids: Ponte Alta.
60. Vellozia swallenii L. B. Smith, sp. nov. PLATE 9, FrauRES 9, 10
A V. pumila Goeth. & Henr., cui verisimiliter affinis, foliorum
marginibus carinaque integris glabrisque, tepalis subduplo majoribus
differt.

266 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Caudex incompletely known, branched, terete and 2 cm. in diameter
including the old leaf-sheaths; leaves densely few-ranked; leaf-sheaths
ca. 4 cm. long, castaneous, lustrous, soon becoming pectinate-fibrous
by the persistence of fine cross-strands on the separating nerves;
leaf-blades 7-10 before falling, suberect, linear, acuminate, 30 em.
long, 7 mm. wide, entire, wholly glabrous, the margins thickened,
pale; scape solitary, slender, 4 cm. long above the leaf-sheaths, tri-
gonous, densely and finely pale-setose toward apex; ovary ellipsoid,
8 mm. long, covered with pale subulate contorted trichomes; tepals
linear-elliptic (?), 80 mm. long; stamens 3 on the unappendaged
phalange.

Type in the U.S. National Herbarium, No. 1592031, collected on chapada

(dry brushy field), between Barra do Corda and Grajati, State of Maranhio,
Brazil, March 1-5, 1934, by Jason R. Swallen (No. 3617).

61. Vellozia bulbosa L. B. Smith, sp. nov. PLATE 4

A V. pumila Goeth. & Henr., cui verisimiliter affinis, planta acaule,
foliorum vaginis haud fissis, laminis integris glabrisque, tepalis ma-
joribus differt.

Stemless; leaf-bases forming an irregularly ovoid bulb, 25 mm.
long, castaneous, lustrous, densely nerved; blades about 10 before
falling, suberect, curved, lincar, acuminate, to 24 cm. long, 6 mm.
wide, conduplicate, glabrous, the keel and margins pale, thickened,
entire; scapes 1-2, slender, 2-3 cm. long above the leaf-sheaths,
densely setose-scabrous toward apex; ovary obovoid, 7-9 mm. long,
covered with fine subulate contorted stramineous trichomes; tepals
linear-elliptic (?), 60 min. long, pale violet; capsule globose, 18 mm.
in diameter, the trichomes becoming thickened and conical.

Type in the Instituto Agronémieco do Norte, colleeted on rocks in savanna,
Serra do Cachimbo, State of Pard, Brazil, altitude 425 meters, December 17,
1956, by J. M. Pires, G. A. Black, J. J. Wurdack, and N. T. Silva (No. 6423),
62. Vellozia cinerascens Mart. ex Schult. f. in R. & 8S. Syst. 7: 292. 1826.

Braziu: Pernambuco: Capoculo. Santa Isabella.

63. Vellozia circinans Goeth. & Henr. Blumea 2: 367. 1937.

Braziu: Minas Gerais: Sao Joio del Rei. Pico da Sander or Pico du Suspiro.
Cachocira do Campo, Serra da Lapa.

64. Vellozia dasypus Scubert in Mart. Fl. Bras. 3, pt. 1: 81. 1847.

Braziu: Bahia: Jacobina. Sao Salvador. Serra Sincora.

Further collections may show that this is no more than a variety
of V. cinerascens which differs only in its glabrous scape. The sup-
posed difference of stamen number is not borne out in the descriptions
of the two species in “Flora Brasiliensis.”’

65. Vellozia glaziovii Goeth. & Henr. Blumea 2: 372. 1937.
Brazit: Goids: Serra dos Veadeiros.

SMITH—SYNOPSIS OF AMERICAN VELLOZIACEAE 267

66. Vellozia crinita Goeth. & Henr. Blumea 2: 368. 1937.
Brazit: Minas Gerais: Sio Jodo del Rei.

67. Vellozia caruncularis Mart. ex Seubert in Mart. Fl. Bras. 3, pt. 1: 78, pl.
8, fig. 1. 1847.
Braziz: Minas Gerais: Serra do Cipé. Serrado Garimpo. Serra Rola Moga.
68. Vellozia gracilis Seubert 7m Mart. Fl. Bras. 3, pt. 1: 81, pl. 9. 1847.
Braziu: Minas Gerais.
69. Vellozia grisea Goeth. & Henr. Blumea 2: 373. 1937.
Braziu: Between Goids and Cuyaba.
70. Vellozia exilis Goeth. & Henr. Blumea 2: 370. 1937.
Brazit: Goids: Chapadaéo dos Viadeira (?Veadeiros).

Section II. Radia

71. Vellozia uaipanensis (Maguire) L. B. Smith, comb. nov.
Barbacenia uatpanensis Maguire, Mem. N.Y. Bot. Gard. 9: 477, fig. 117.
1957.
VENEZUELA: Bolivar: Uaipdn-tepuf.
72. Vellozia markgrafii Schulze-Menz ex Markgraf, Notizblatt 15: 216. 1940.
Braziu: Minas Gerais: East of Montes Claros.

73. Vellozia riedeliana Goeth. & Henr. Blumea 2: 379. 1937.
Brazit: Minas Gerais: Serra da Lapa.

74. Vellozia maguirei LL. B. Smith, sp. nov. PLATE 5

A V. markgrafit Schulze-Menz ex Markgraf et V. riedeliana Goeth.
& Henr., quibus affinis, foliis subtus glabris, pedunculo majore differt.

Caudex to 1 meter high (! Maguire), amount of branching unknown,
terete and 3 cm. in diameter at apex (including the old leaf-sheaths) ;
leaf-sheaths 4 cm. long, connate in a funnel for 2 cm., lustrous, densely
granular-resinous, densely nerved, the nerves soon separating as
coarse fibers; leaf-blades about 12 before falling, suberect, linear,
filiform-attenuate, to 28 cm. long, 10 mm. wide at base, densely nerved,
resinous, minutely lepidote-ciliate at base on the keel and thickened
margins, otherwise glabrous; scape solitary (?), 3 cm. long, glabrous;
flowers white; perianth-tube slenderly cylindric and 8 cm. long above
the ellipsoid 1 cm. long ovary, 5 mm. in diameter, subdensely glandu-
lar; ovary covered with short cylindric gland-tipt trichomes; tepals
elliptic, to 5 cm. long.

Type in the U.S. National Herbarium, No. 2340339, collected 49 miles from
Diamantina, State of Minas Gerais, Brazil, altitude 1140 meters (3800 feet),

December 22, 1959, by Bassett Maguire, C. K. Maguire and J. Mur¢a Pires (No.
44742). Isotype in the New York Botanical Garden.

75. Vellozia candida Mikan, Delect. Fl. & Faun. Brazil. 2: pl. 1. 1820; text
1825.
Vellozia tertia Spreng. Neu. Entd. 2: 108. 1821.
?Vellozia maritima Vell. Fl. Flum. 219. 1825;Icon.5:pl. 79. 1835. (Rocks,
Island of Parati).
Braziu: Guanabara: Tijuca. Dois Irmfos. Penha. Corcovado.

?268 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

76. Vellozia macrantha Lem. Ill. Hortic. 12: Misc. 41. 1865.
Braziu: Without locality.
77. Vellozia leucanthos Goeth. & Henr. 2: 375. 1937.
Braziut: Minas Gerais: Caraga.
78. Vellozia cachimbensis L. B. Smith, sp. nov. PuaTE 6

A V. macrosiphonia Taub., cui verisimiliter affinis, caudice brevis-
simo, pedunculos folias subaequantibus differt.

Plant stemless or sometimes with a caudex up to 4 cm. long and
2 cm. in diameter (including the old leaf-sheaths); leaves densely
many-ranked; sheaths ca. 2 cm. long, castaneous, lustrous, white,
separating into coarse fibers at apex; blades 20-30 before falling,
suberect to spreading, linear, filiform-attenuate, to 14 cm. long, 6 mm.
wide, densely nerved, glabrous above, typically lepidote-pilose be-
neath, setose-ciliate; scape solitary, very slender, about equaling the
leaves, densely stipitate glandular toward apex; flowers white (! Pires) ;
perianth-tube slenderly cylindric and 45-65 mm. long above the
ellipsoid 5 mm. long ovary, 1-2 mm. in diameter, rather densely
glandular; ovary covered with subsessile glands; tepals 4 cm. long;
stamens 12 (! Pires).

Type in the Instituto Agronémico do Norte, No. 90.910, collected on rocks,

Serra do Cachimbo, State of Pard, Brazil, altitude 425 meters, December 14,
1956, by J. M. Pires, G. A. Black, J. J. Wurdack, and N. T. Silva (No. 6232).

79. Vellozia macarenensis Philipson in Schultes, Fam. Velloz. Colomb. in Rev.
Acad. Colomb. Cienc. 8, No. 32: 461, pl. 1952.
Cotomsia: Meta: Sierra de la Macarena.

80. Vellozia macrosiphonia Taub. Bot. Jahrb. 21: 424. 1896.
Braziu: Goids: Serra dos Pirineus. Serra Dourada. Serra dos Viadeiros.

81. Vellozia maculata Goeth. & Henr. Blumea: 2: 375. 1937.
Brazit: Goids: Cachoeiras da Vargem Grande da Serra da Balisa.

82. Vellozia machrisiana L. B. Smith, sp. nov. PLATE 9, FIGURES 11, 12

A V. leucantho Goeth. & Henr., cui aflinis, foliis scapoque multo
majoribus, foliorum vaginis vetustis haud vel vix fissis differt.

Low, only fruiting material known; caudex simple (?), more or less
prostrate (! Dawson), 15 cm. long, terete, 1 cm. in diameter near
apex; leaves 3-ranked, very densely imbricate, the complete ones
about 12 at the apex of the stem; sheaths sulcate, densely white-
sericeous beneath except for the narrow brown lustrous margin,
scarcely if at all divided with age, the apices of the upper ones strongly
recurved; blades linear, filiform-acuminate, 27 cm. long, 5.5 mm. wide
at base, densely and strongly nerved on both sides, otherwise even,
ciliate toward base on the thickened margins and rounded keel,
otherwise glabrous; scape solitary, extending 18 cm. above the leaf-
sheaths, obtusely trigonous, very sparsely vestite toward apex with
minute stipitate black glands, capsule ellipsoid, 12 mm. long, strongly

SMITH—SYNOPSIS OF AMERICAN VELLOZIACEAE 269

3-lobed, completely covered with minute ovoid gland-tipt trichomes;
remains of the perianth-tube slenderly cylindric, 8 cm. long, sparsely
and minutely sessile-glandular near base.

Type in the U.S. National Herbarium, No. 2280629, collected on rocks in
stream below falls (subject to running water), sandstone area 14 kilometers south

of Veadeiros, region of the Chapada dos Veadeiros, ca. lat. 14°30’ S., long. 47°30’
W., State of Goids, Brasil, April 25, 1956, by E. Yale Dawson (No. 14674).

83. Vellozia uleana_ L. B. Smith, sp. nov. PuLaTE 9, FIGURES 13, 14

A V. leuwcantho Goeth. & Henr., cui affinis, vaginarum nervis sub-
contiguis, foliis in apicem filiformem attenuatis, ovarii glandulis
graciliter stipitatis differt.

Caudex over 23 cm. long, branching at least once, terete and 1
cm. in diameter (including the old leaf-sheaths) ; leaf-sheaths 4 cm.
long, dull, densely nerved, remaining entire and not splitting into
fibers, recoiling at apex; leaf-blades about 12 before falling, suberect
to spreading, linear, filiform-attenuate, to 25 cm. long, 6 mm. wide,
densely nerved, setose-ciliate on the keel and thickened margins when
young, glandular-punctate above, glabrous beneath; scape solitary,
9 cm. long above the leaf-sheaths, slender, subdensely stipitate-
glandular toward apex; flowers white; perianth-tube slenderly cylindric
and 5-7 cm. long above the ellipsoid 8 mm. long ovary, 1.5 mm. in
diameter, subdensely glandular; ovary covered with slender gland-tipt
trichomes; tepals 4 em. long.

Type in the Museu Paraense ‘Emilio Goeldi,’’ No. 13.447, collected on rock
faces of the Serra de Uairary (?Mairarf), Surumd, Territory of Rio Branco,

Brazil, altitude 1000-1200 meters, September 1909, by E. Ule (No. 8372). Iso-
types in the New York Botanical Garden and U.S. National Herbarium.

84. Vellozia panamensis Standl. Journ. Washington Acad. Sci. 15: 457. 1925.
PanaMA: Chiriquf: Cerro Vaca.
85. Vellozia phantasmagoria R. E. Schultes, Bot. Mus. Leafl. Harvard 12: 130,
pl. 19, 20. 1946.
Cotomsia: Vaupés: Mount Chiribiquete.
86. Vellozia dumitiana R. E. Schultes, Mutisia No. 12: 2, pl. 1952.
CotompBia: Vaupés: Cerro Isibukuri.
87. Vellozia rhynchocarpa Goeth. & Henr. Blumea 2: 378. 1937.
Braziu: S40 Paulo: Between Canna Verde and Cajura near Mato Grosso.
88. Vellozia maudeana R. E. Schultes, Bot. Mus. Leafl. Harvard 16: 198, pls.
29, 30. 1954.
CotomBi1A: Vaupés: Mesa de Yambf.
89. Vellozia annulata Goeth. & Henr. Blumea 2: 365. 1937.
Brazit: Goids: Cachoeira da Vargem Grande. Serra dos Veiadadoes
(? Veadeiros).
90. Vellozia velutinosa Goeth. & Henr. Blumea 2: 382. 1937.
Braziu: Goids: Cabeceira (headwaters) do Rio Santa Ana.

270 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

91. Vellozia dawsonii L. B. Smith, sp. nov. PLATE 7

A V. velutinosa Goeth. & Henr., cui verisimiliter affinis, foliorum
laminis subduplo latioribus, floribus haud solitariis differt.

Caudex incompletely known, 25 mm. in diameter (including the old
leaf-sheaths) ; leaves densely many-ranked; sheaths 4 cm. long, densely
white-lanate; blades over 30 before falling, linear-triangular, filiform-
attenuate, 20 cm. long, 17 mm. wide, finely lepidote-pilose on both
sides, flat, densely and minutely ciliate; scapes at least 3 together,
slender, about equaling the leaves, densely stipitate-glandular toward
apex; flowers white; perianth-tube slenderly cylindric, 70-80 mm. long
above the ellipsoid 10 mm. long ovary, 4-5 mm. in diameter, sub-
densely glandular; ovary covered with slenderly stipitate glands;
tepals elliptic, acute, 7 cm. long; anthers 15 mm. long.

Type in the U.S. National Herbarium, No. 2279791, collected on sandstone
outcrop 7 kilometers south of Veadeiros, region of the Chapada dos Veadeiros,

ca. lat. 14°30’ S., long. 47°30’ W., State of Goids, Brasil, April 24, 1956, by E.
Yale Dawson (No. 14580).

92. Vellozia cana Goeth. & Henr. Plumea 2: 367. 1937.
Brazit: Goids: Paranana.

93. Vellozia lithophila R. I. Schultes, Fam. Velloz. Colomb. in Rev. Acad.
Colomb. Cienc. 8, No. 32: 459, pl. 1950.

Cotomsia: Vaupés: Mesa La Lindosa. San José del Guaviare. Cerro Yapo-
boddé. Cerro Kafienddé. Rio Parana Pichuna.

94. Vellozia tubiflora (A. Rich.) H. B. K. Nov. Gen. & Sp. 7: 155. 1824.
Radia tubiflora A. Rich. in Kunth, Syn. Pl. Aeq. 1: 300. 1822,
Barbacenia alexandrinae Rob. Schomb. Barbacenia Alexandrinae 13, pl.

1845; Rob. Schomb. ex Hook. London Journ. Bot. 4: 13. 1845.

Barbacenia tubiflora Jackson, Ind. Kew. 4: 1269. 1895. Wrongly attributed
to Benth. & Hook. Gen. 3: 740. 1883, who made the combination only
inferentially and not validly.

Vellozia alexandrinae Goeth. & Henr. Blumea 2: 363. 1937.

VENEZUELA: Amazonas and Bolivar. Brazil: Paré. British Guiana.

The only distinction encountered in the descriptions of Vellozia
tubiflora and V. alexandrinae is the height and branching of the caudex.
Vellozia tubiflora from along the Rio Orinoco is described as low and
slightly branched while V. alerandrinae from Mount Roraima is 3
to 4 meters high and much branched. However, Dr. J. J. Wurdack,
who has observed V. tubiflora in the type area, assures me that it
varies according to age and attains great size also.

2. Barbaceniopsis

Barbaceniopsis L. B. Smith, gen. nov.

Caulescens, simplex vel ramosus; caudice foliorum vaginis vetustis
omnino obtecto; foliis novellis terminalibus, laminis linearibus, haud

SMITH—SYNOPSIS OF AMERICAN VELLOZIACEAE 271

regulariter deciduis; scapis terminalibus, unifloris; tepalis 6, sub-
aequalibus, tubum epigynum formantibus; staminibus 6; filamentis
brevissimis, teretibus, haud appendiculatis; antheris dorsifixis; ovario
infero; styli parte stigmatifera subcylindrica.

Type species: Barbaceniopsis boliviensis (Baker) L. B. Smith.

Key to the Species of Barbaceniopsis

1. Leaf-blades even, cinereous beneath, 2-2.5 mm. wide; all tepals merely

acute... 1 ee ee ee ee ee ee ee ee we) 6B. boliviensis
1. Leaf-blades carinate, green beneath, 3.5 mm. wide; outer tepals long-
aristate . 2. 1 ee eee ee ee ee ee ed 62 BY vargasiana

1. Barbaceniopsis boliviensis (Baker) L. B. Smith, comb. nov.
PLATE 9, FIGURES 15, 16
Vellozia boliviensis Baker, Mem. Torrey Bot. Club 6: 126. 1896.
Barbacenia boliviensis Hauman, Anal. Mus. Nac. Hist. Nat. Buenos Aires
29: 429. 1917.
Barbacenia castilloni Hauman, Anal. Mus. Nac. Hist. Nat. Buenos Aires 29:
426, pl. 4. 1917.

Bouivia: Cochabamba: Near Cochabamba, Bang 1134 (K, type; US). Tarija:
La Merced near Bermejo, alt. 1700 m., FPiebrig 2505 (F).

ARGENTINA: Salta: Cafayate, alt. 1400 m., November 24, 1949, Araque &
Barkley 19—-Ar-366 (F). Same, alt. 1700 m., February 10, 1953, Hayward 2512
(US). Jujuy: Tumbaya, Volcan to Loma del Tambo, alt. 2500 m., February
22, 1924, Schreiter 2561 (US from hb. Venturi 3445). Tilcara, alt. 2300 m.,
February 20, 1926, Venturi 6847 (US). Capital, Quebrada de Chafii, alt. 1600 m.,
January 30, 1940, Schreiter 10959 (UC). Tucumén: Tafi, El Bafiado, Quebrada
de las Cafias, alt. 2140 m., February 1913, Castillon 3245 (BA). Calchaquies,
El Bafiado, Quebrada de las Cafias, alt. 2200 m., January 10, 1917, Castillon 58
(US).

Note: I have not examined the type of Barbacenia castilloni
Hauman, but other material from the same region shows no essential
difference from Vellozia boliviensis Baker.

2. Barbaceniopsis vargasiana (L. B. Smith) L. B. Smith, comb. nov.
Barbacenia vargasiana L. B. Smith, Bol. Soc. Peru Bot. 1: 18, figs. 1, 2.

1948.
Perv: Cuzco: Anta, between Sisal and Cunyac, Vargas 4883 (GH, type; US).

3. Barbacenia

Barbacenia Vand. FI. Lusit. & Brasil. Spec. 21, pl. 1. 1788.
Pleurostima Raf, Fl. Tellur. 2: 97. 1836.
Visnea Steud. ex Endl. Gen. 173. 1837. Nomen in synonymy.

Plants of small or medium stature; caudex usually simple or few-
branched; leaf-blades breaking off irregularly; tepals forming an
epigynous tube; stamens 6; filaments evident, flat with 2 lobes or
appendages at the summit; anthers dorsifixed; stigmas inconspicuous,
not peltate nor broadly lobed.

Type species: Barbacenia brasiliensis Willd.

272 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Keys to the Species of Barbacenia

1. Perianth-tube not more than twice as long as the ovary.
2. Ovary section of the perianth partly or wholly glabrous, the costae evident

(lacking in B. beauverdit). . . . 2 ee es) 6SUBKEY I

2. Ovary section of the perianth wholly vestite . wee es | ) 6SuBKkey IT

1. Perianth-tube more than twice as long as the ovary. . . . . . Susxery III
SUBKEY I

1. Perianth-tube completely glabrous.
2. Tube of the perianth smooth, ecostate ...... . . .1. B. beauverdii
2. Tube of the perianth costate.
3. Costae of the perianth 18; caudex very short; leaf-blades 11 em. long,

10mm. wide. ... . . .2. B. brachycalyx
3. Costae of the perianth 12; caudex to 30. cm. ‘ong; leaf-blades 7-8 cm.
long, 2.6mm. wide ........4.2.4...... .3. B.caricina

1. Perianth-tube vestite on the costae.
4, Indument of the perianth of pointed trichomes, eglandular.
5. Anthers twice as long as the filaments. ...... .4. B. squamata
5. Anthers exceeded by the filament-appendages.
6. Tepals 40 mm. long, ample; leaf-blades 9 mm. wide . . . 5. B. rogieri
6. Tepals 20-30 mm. long, narrow.
7, Leaf-blades 5mm. wide. ...... 2.2.2... .6. B. gaveensis
7. Leaf-blades 7-9 mm. wide. .... .. . 7. B. seubertiana
4. Indument of the perianth of sessile (verruculose) or of stipitate glands.
8. Anthers equaling or exceeding the filament-appendages.
9. Leaf-blades 2 mm, wide; perianth-tube 8 mm. long.
8. B. stenophylla
9. Leaf-blades 6-18 mm. wide; perianth-tube 13-20 mm. long.
10. Perianth-tube about twice as long as the ovary; leaf-blades 12-18
mm. wide. .... . . . .9 B. inclinata
10. Perianth-tube not more > than half again as s long as the ovary.
11. Scape 35 cm. long; leaf-blades 10-18 mm. wide.
10. B. longiscapa
11. Scape ca. 15 cm. long; leaf-blades 4-11 mm. wide.
12. Leaf-blades 9-11 mm. wide; filaments 8 mm. long.
11. B. flavida
12. Leaf-blades 4-8 mm. wide.
13. Tepals glandular-punctate; costae wholly covered by black
stipitate glands... . . ... . .12. B. irwiniana
13. Tepals glabrous; costae laxly glandular or the intermediate
ones glabrous.
14. Filaments 7-8 mm. long, only slightly shorter than the
anthers; leaves distributed along the stem.
13. B. foliosa
14, Filaments ca. 4 mm. long, about half as long as the anthers;
leaves clustered at apex of stem (perianth-tube described
as glandular, but not so depicted) . . (4, B. squamata)
8. Anthers exceeded by the filament-appendages.
15. Scape 1 cm. long; perianth-tube about twice as long as the ovary and
forming a slender tube above it; anthers medifixed. . 14. B. goethartii

SMITH—SYNOPSIS OF AMERICAN VELLOZIACEAE 273

15. Scape elongate; perianth-tube much less than twice as long as the
ovary and forming a broad cylinder above it.
16. Leaf-blades 6-8 cm. long, only %4—% as long as the scapes, 2-4 mm.
wide; anthers equaling the base of the filament-sinus.
15. B. brevifolia
16. Leaf-blades more than twice as large, more than half as long as the
scape and mostly exceeding it.
17. The leaf-blades entire; costae bearing stipitate glands; style
subulate, scarcely dilated by the sigmas . 16. B. mantiqueirae
17. The leaf-blades denticulate or spinulose-serrate.
18. Tepals 40 mm. long, ample; leaf blades 9 mm. wide (perianth-
tube described as glandular, but not so depicted).
(5. B. rogieri)
18. Tepals 16-20 mm. long, narrow.
19. Leaf-blades with narrow dark margins and keel, denticulate;
stigmas apical, making the style slenderly conical.
17. B. nigrimarginata
19. Leaf-blades concolorous.
20. Stigmas apical on the style, making it clavate; anthers
acuminate, oxceading the base of the sinus; leaf-blades

denticulate. . . . ... . . 18. B. gounelleana

20. Stigmas submedian on nthe style; anthers rounded-apiculate;

leaves spinulose-serrate. . . ... . . 19. B. purpurea
SUBKEY II

1. Indument of the perianth wholly of pointed trichomes, not glandular.
2. Tepals only about % as long as the tube; leaf-blades hirsute beneath.
20. B. schwackei
2. Tepals 1-2 times as long as the tube; leaf-blades glabrous . . 21. B. fulva
1. Indument of the perianth glandular, either sessile or stipitate.
3. Scapes up to 12 per rosette; leaves up to 40; sheaths resinous-conglutinated;
flowers violet; tepals 4-5 mm. long. . . ... . . . 22. B. polyantha
3. Scapes 1-5 per rosette.
4, Leaf-blades uniformly vestite.
5. Ovary trigonous, verrucose ...... .- .... . 23. B. trigona
5. Ovary terete or if subtrigonous then covered with stipitate glands.
6. Flowers yellow; Perianth-tube 12 mm. long, twice as long as the
ovary .. . ee ee ew ww we 1 oe B. globata
6. Flowers red to lilac, or white.
7. Glands of the perianth- -tube sessile; perianth-tube twice as long
asthe ovary .. .... . 25. B. coccinea
7. Glands of the perianth-tube stipitate.
8. Leaf-indument glandular.
9. Tepals 7 mm. long; flowers red. . . .. . 26. B. glutinosa
9. Tepals 17-20 mm. long; flowers white . . 27. B. markgrafii
8. Leaf-indument eglandular.
10. Perianth-tube 25 mm. long; flowers red. . . 28. B. ignea
10. Perianth-tube 20 mm. long; flowers lilac . . 29. B. lilacina
4, Leaf-blades glabrous except for the margins and keel.
11. Glands of the ovary sessile; perianth-tube 50 mm. long; anthers medifixed,
nearly twice as long as the filaments; flowers yellow . 30. B. vandellii

274 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

11. Glands of the ovary stipitate.
12. Anthers equaling the filament-appendages; flowers yellow.
31. B. brasiliensis
12. Anthers exceeding the filament-appendages.
13. Trichomes of the upper scape long, slender, mostly or entirely
eglandular; tepals 9-11 mm. long.
14. Flowers deep lilac; leaf-blades 10 mm. wide . 32. B. hirtiflora
14. Flowers yellow; leaf-blades 7 mm. wide. . 33. B. williamsii
13. Trichomes of the upper scape short, stout, coarsely glandular;
tepals 15-17 mm. long.
15. Epigynous tube subcylindric; perianth-tube 20-23 mm. long.
34. B. flava
15. Epigynous tube campanulate; perianth-tube 25-32 mm. long.
16, Perianth-tube 32 mm. long; caudex 3-4 dm. high.
35. B. glauca
16. Perianth-tube 25 mm. long; caudex very short.
17, Leaf-blades 40 cm. long, 8-10 mm. wide, glaucous.
36. B. itabirensis
17. Leaf-blades 9-16 em. long, 4-7 mm. wide, green (?).
37. B. sellovii

SUBKEY II

1. Npigynous tube infundibuliform, distinctly wider at the mouth than at the

base.
2. Tepals broadly ovate, as broad as tong perianth-tube 35 mm. long; flowers
subsessile . . . . .... . 38 B. macrantha

. Tepals oblong, lanceolate, or clliptic, “much longer than broad.
3. Perianth-tube 70-75 mm. long.
4, Tepals reflexed, 40 mm. long; perianth-indument eglandular.
39. B. rubro-virens
4, Tepals erect, 20 mm. long; perianth-indument glandular.
40. B. riedeliana
3. Perianth-tube 30-55 mm. long.
5. Indument of perianth eglandular; perianth-tube 30 mm. long.
41. B. luzulifelia
5. Indument of perianth glandular; perianth-tube 50-55 mm. long.
6. Leaf-blades 9 em. long, 9 mm. wide; ovary 10 mm. long, %-% of

the perianth-tube.... . .. . . 42. B. viscosissima
6. Leaf-blades 22 em. long, 17 mm. wide: ovary 17 mm. long, % of
the perianth-tube . . . ... . 43. B. paranaensis

1. Epigynous tube slenderly cylindric or very ‘slightly enlarged toward apex.
7. Indument of the perianth subulate or lacking.
8. Flowers sessile.
9. Tepals broadly elliptic, 10 mm. long; flowers yellow . .44. B. exscapa
9. Tepals linear, 24 mm. long; flowers red . . . 45. B. gentianoides
8. Flowers distinctly scapose.
10. Perianth-tube 50-60 mm. long.
11. Ovary % of perianth-tube; scape pilose .. . . .46. B. hilairei
11. Ovary % of perianth-tube.
12. Scapes glabrous; perianth-tube 50 mm. long, sparsely pilose on
the costae... . .... . 47. B. oxytepala
12. Scapes densely tomentellous tow ard apex; perianth-tube 60 mm.
long, densely and evenly gray-tomentellous . . . 48. B. grisea

SMITH—SYNOPSIS OF AMERICAN VELLOZIACEAE 275

10. Perianth-tube 30-40 mm. long.
13. Plant completely glabrous; tepals ovate or lance-ovate, acute.
49. B. glabra
13. Plant vestite.

14. Filaments less than half as long as the anther; tepals erect, 12
mm.long ... .... . .90. B. tomentosa
14, Filaments as long a as or ‘longer than the anther, tepals subspread-

ing to reflexed, 15-26 mm. long.
15. Epigynous tube 10 mm. in diameter; tepals reflexed, obtuse.
51. B. schidigera
15. Epigynous tube, 3-6 mm. in diameter; tepals subspreading,

acute.
16. Perianth-tube densely pubescent; epigynous tube 4-6 mm.
in diameter .... .. . . .41. B. luzulifolia

16. Perianth-tube very sparsely pubescent and appearing
glabrous; epigynous tube 3 mm. in diameter.
52. B. graciliflora
7. Indument of the perianth glandular.
17. Glands of the perianth sessile or subsessile.
18. Leaves glabrous except the margins; perianth-glands sessile; tepals
12 mm.long... . ... . .53. B. ensifolia
18. Leaves evenly pilose; perianth glands subsessile; tepals 23 mm. long.
54. B. glaziovii
17. Glands of the perianth long-stipitate.
19. Scape exceeding the leaves, its glandular indument of 2 types.
20. Perianth-tube 3 times as long as the ovary . . 55. B. conicostigma
20. Perianth-tube 4-5 times aslongas theovary . . .56. B. cuspidata
19. Scape equaling or shorter than the leaves.
21. Leaf-blades glabrous except the keel and margins.
22. Perianth-tube 65 mm. long; flower yellow . . .57. B. longiflora
22. Perianth-tube 30-50 mm. long.
23. The perianth-tube 5 times as fone as the ovary, straight,
ecostate. .... .. . . . 58 B. gardneri
23. The perianth-tube 3 times as long as the ovary, curved, costate.
59. B. curviflora
21. Leaf-blades evenly vestite.
24, Filaments linear.

25. Tepals 30-35 mm. long ........ . . 60. B. fragrans
25. Tepals 18-20 mm. long.
26. Leaf-blades 4mm. wide ....... . . 61. B. tricolor
26. Leaf-blades 10 mm. wide... .. . . . 62. B. blanchetii

24. Filaments broad.

27. Sinus of the filament shallow; filaments quadrate, 4 mm. long.
63. B. mollis

27. Sinus of the filament deep; filaments oblong or broad-based.
28. Indument of the scape white, contrasting sharply with
the dark indument of the perianth-tube; perianth-tube
25mm.long.... .... . 64, B. leucopoda
28. Indument of the scape and ‘perianth-tube uniformly dark;

perianth-tube 30-50 mm. long.
29. Filaments oblong, their sides parallel; anthers 8-9 mm.
long; leaf-blades uniformly setose-glandular.

597430—62——_5

276 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

30. Leaf-blades 9 mm. wide; anthers exceeding the filaments

by less than 4. . . ... . . 65. B. blackii
30. Leaf-blades 51 mm. wide; ‘anthers exceeding the filaments
by %. . .... . 66. B. rubra

29. Filaments broad- based, much contracted upward; anthers
12-15 mm. long, exceeding the filaments by %4 or more.
31. Ovary rounded at base, broadly obovoid.
67. B. damaziana
31. Ovary long-attenuate at base, slenderly obconic.
68. B. bahiana

1. Barbacenia beauverdii Damazio, Bull. Herb. Boiss. I. 7: 595, fig. 1907.
BraziL: Minas Gerais: Serra do Frasdo.

2. Barbacenia brachycalyx Goeth. & Henr. Blumea 2: 340, fig. A. 1937.
Brazit: Goidis: Pichoa to Morro do Espigao.

3. Barbacenia caricina Goeth. & Henr. Blumea 2: 340, fig. B. 1937.
Brazit: Rio de Janeiro: Nova Friburgo to Pedra do Conego.

4. Barbacenia squamata Hook. Bot. Mag. 71: pl. 4136. 1845.

Vellozia squamata Jackson, Ind. Kew. 4: 1269. 1895, wrongly attributed to
Benth. & Hook. Gen. 3:740. 1883, who made the combination only in-
ferentially and not validly.

Brazi_: Rio de Janeiro: Serra dos Orgios. Guanabara: Morro do Flamengo.
5. Barbacenia rogieri hort. ex Moore & Ayres, Mag. Bot. 2: 209, pl. 1850; Lem.
Jard. Fl. 1: pl. 82. 1851.
Sourn America: No locality.
6. Barbacenia gaveensis Goeth. & Henr. Blumea 2: 346. 1937.
Braziu: Guanabara: Gavea. Tijuca.
7. Barbacenia seubertiana Goeth. & Henr. Blumea 2: 360. 1937.

Barbacenia purpurea Wook. var. minor Seubert in Mart. Fl. Bras. 3, pt.

1:68. 1847.
Braziv: Rio de Janeiro: Serra dos Orgaos, Faboinha. Guanabara: Gavea.
8. Barbacenia stenophylla Goeth. & Henr. Blumea 2: 361, fig. F. 1937.
Braziu: Goids: Morro do Salto.
9. Barbacenia inclinata Goeth. & Henr. Blumea 2: 352. 1937.
Braziu: Minas Gerais: Diamantina.
10. Barbacenia longiscapa Goeth. & Henr. Blumea 2: 355. 1937.
Braziut: Minas Gerais: Serra da Lapa.
11. Barbacenia flavida Goeth. & Henr. Blumea 2: 343. 1937.
Brazit: Minas Gerais. Goids: Pichoa to Morro do Espigao.
12. Barbacenia irwiniana L. B. Smith, sp. nov. Pate 10, ricuREs 17, 18
A B. foliosa Goeth. & Henr., cui affinis, tepalis glanduloso-punctatis,
ovarii costis glandulis nigris stipitatis omnino obtectis differt.
Caudex 3 cm. long with roots almost throughout; leaves numerous,
3-ranked; sheaths ovate, 1 cm. long, very closely nerved, glabrous;
blades linear, acuminate, 13-15 cm. long, 4 mm. wide, closely nerved

SMITH—SYNOPSIS OF AMERICAN VELLOZIACEAE 277

on both sides, sparsely denticulate on keel and margins, otherwise
smooth and glabrous; scape solitary, about equaling the leaves,
slender, bearing black stipitate glands especially near apex; perianth-
tube 15 mm. long; ovary ellipsoid, the costae covered with black
stipitate glands; epigynous tube infundibuliform, 4 mm. high; tepals
suberect at anthesis, lanceolate, subacute, 13 mm. long, yellow
(Irwin), reddish when dry, the outer slightly narrower; filaments
6 mm. long, the oblong rounded appendages about twice as long as
the quadrate base; anthers linear, 8 mm. long, slightly exceeding
the filament-appendages.

Type in the U.S. National Herbarium, No. 2324659, collected in narrow crevice
among rocks on steep slope, Pico da Bandeira, Serra Caparao, State of Minas
Gerais, Brazil, at 2700 meters (9000 feet) altitude, March 8, 1959, by H. 8.
Irwin (No. 2803).

13. Barbacenia foliosa Goeth. & Henr. Blumea 2: 344, fig. D. 1937.

BraziL: Guanabara: Tijuca.

14. Barbacenia goethartii Henr. Blumea 2: 350, fig. K. 1937.

Braziu: Minas Gerais: Penha (?).

15. Barbacenia brevifolia Taub. Bot. Jahrb. 12, Beibl. 27: 2. 1890.

Braziu: No locality.

16. Barbacenia mantiqueirae Goeth. & Henr. Blumea 2: 355. 1937.

Braziu: Sao Paulo: Serra da Mantiqueira.

17. Barbacenia nigrimarginata L. B. Smith, sp.nov. Paty 10, rigurms 19-22

A B. gounelleana Beauverd, cui affinis, foliorum vaginis mox in
fibras solutis, laminis nigrimarginatis, filamentorum lobis obtusis,
antheris filamentorum sinum apicalem haud attingentibus differt.

Caudex only about 2 cm. long; leaves very densely imbricate,
glaucous beneath; sheaths 3 em. long, soon dividing into coarse stiff
fibers; blades very narrowly triangular, filiform-acuminate, not at
all contracted at base, 25 cm. long, 9 mm. wide, finely and closely
nerved on both sides, the thickened margins and keel black and
minutely serrulate; scape solitary, 25 cm. long, slender, sulcate, the
upper part obscurely glandular; perianth-tube obconic, 17 mm. long,
very slightly exceeding the ovary, 5 mm. in diameter, diffusely
verruculose; tepals suberect at anthesis, elliptic, obtuse, 22 mm. long,
the outer distinctly narrower; filaments narrowly obovate with short
rounded lobes, about half as long as the tepals; anthers linear, 6
mm. long, not reaching the base of the sinus.

Type in the U.S. National Herbarium, No. 2279780, collected on stony sum-
mit of butte shoulder 5 kilometers west of Veadeiros, region of the Chapada
dos Veadeiros, ca. lat. 14°30’ S., long. 47°30’ W., State of Goids, Brazil, April 29,
1956, by E. Yale Dawson (No. 14717).

18. Barbacenia gounelleana Beauverd, Bull. Herb. Boiss. 11. 7: 704. 1907.

Braziu: Rio de Janeiro: Itatiaia.

278 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

19. Barbacenia purpurea Hook. Bot. Mag. 54: pl. 2777. 1827.
Pleurostima purpurea (Hook.) Raf. Fl. Tellur. 2: 97. 1836.

Brazi_: Guanabara (?): “Neighborhood of Rio de Janeiro.”

20. Barbacenia schwackei Goeth. & Henr. Blumea 2: 359, fig. J. 1937.
Braziu: Minas Gerais: Serra do Cipé,

21. Barbacenia fulva Goeth. & Henr. Blumea 2: 345, fig. E. 1937.
Braziu: Minas Gerais: No locality.

22. Barbacenia polyantha Goeth. & Henr. Blumea 2: 357. 1937.
Braziu: Minas Gerais: Biribiri.

23. Barbacenia trigona Goeth. & Henr. Blumea 2: 362. 1937.
Braziu: Minas Gerais: Milho Verde.

24. Barbacenia globata Goeth. & Henr. Blumea 2: 349. 1937.
Braziu: Minas Gerais: Rio das Pedras.

25. Barbacenia coccinea Mart. ex Schult. f. in R. & S. Syst. 7: 286. 1826.
Braziu: Minas Gerais: Itambé,

26. Barbacenia glutinosa Goeth. & Henr. Blumea 2: 350. 1937.
Braziu: Minas Gerais: Morro da Lapa.

27. Barbacenia markgrafii Schulze- Menz ex Markgraf, Notizblatt 15: 216. 1940.
Braziu: Minas Gerais: Gréo Mogul to Montes Claros.

28. Barbacenia ignea Mart. ex Schult. f. in R. & 8. Syst. 7: 285. 1826.
Braziu: Minas Gerais: Serra Frio. Sérro.

29. Barbacenia lilacina Goeth. & Henr. Blumea 2: 354. 1937.

29a. Barbacenia lilacina var. lilacina

Leaf-blades hirsute; filaments bifid 4-4; flowers lilac.

Braziu: Minas Gerais: Serra do Funil, Rio Paranaha.
29b. Barbacenia lilacina var. pallidiflora Henr. Blumea 2: 354. 1937.

Leaf-blades sparsely hirsute; filaments bifid more than %; flowers
pale lilac.

Braziu: Minas Gerais: Curralinho to Diamantina.
30. Barbacenia vandellii Pohl ex Seubert in Mart. Fl. Bras. 3, pt. 1: 72, pl. 8,
fig. 3. 1847.
Brazit: Minas Gerais: Sao Jofio. Itambé.
31. Barbacenia brasiliensis Willd. Sp. Pl. 2: 227. 1799.

Barbacenia Vand. Fl. Lusit. & Brasil. Spec. 21, pl. 1. 1788; Roemer, Seript.
Hisp. 98, pl. 6, fig. 9. 1796. Without indication of species.

Barbacenia vandelli Schult. f. in R. & S. Syst. 7: 288. 1826. Wrongly
attributed to Roemer who cited the genus and its author without any
specific name.

Barbacenia bicolor Mart. Nov. Gen. & Sp. 1: 19, pl. 13. 1823.

Visnea Steud. ex Endl. Gen. 173. 1837. Nomen in synonymy, without
indication of species.

Braziu: Minas Gerais: Diamantina.
32. Barbacenia hirtiflora Goeth. & Henr. Blumea 2: 351, fig. L. 1987.
Braziu: Minas Gerais: Curralinho to Diamantina.

33. Barbacenia williamsii L. B. Smith, sp. nov. PLaTE 10, FIGURES 23, 24

A B. hirtiflora Goeth. & Henr., cui affinis, floribus aureis, foliorum
laminis angustioribus differt.

SMITH—SYNOPSIS OF AMERICAN VELLOZIACEAE 279

Caudex very short; leaves numerous, persistent; sheaths ovate,
1 cm. long, glabrous, sulcate; blades linear, filiform-acuminate, to
22 cm. long, 7 mm. wide, closely and finely nerved, dull green, glabrous
except for the long-ciliate keel and thickened yellow margins; scapes
1-3, slightly shorter to slightly longer than the leaves, ca. 1 mm. in
diameter, sulcate, densely crisp-pilose with slender red-brown mostly
eglandular trichomes; perianth-tube 25 mm. long, about twice the
ellipsoid ovary and cylindrical above it, 5 mm. in diameter, densely
pilose-glandular; tepals suberect at anthesis, oblong, obtuse, 11 mm.
long; filaments oblong with long acuminate lobes, nearly equaling
the linear anthers.

Type in the Gray Herbarium of Harvard University, collected on iron ore,
Serra da Mutuca, near Belo Horizonte, Municipio of Nova Lima, State of Minas
Gerais, Brazil, altitude 1200-1400 meters, April 7, 1945, by Louis O. Williams

and Vicente Assis (No. 6696). Isotype in the U.S. National Herbarium, No.
1932777.

Braziu: Minas Gerais: Mun. Nova Lima: In rocky slopes, Serra da Mutuca,
February 1945, L. O. Williams 5201 (GH); 5204 (GH). In campo, Serra do
Curral, alt. 1300 m., March 30, 1945, L. O. Williams & V. Assis 6386 (GIT, US).

34. Barbacenia flava Mart. ex Schult. f. in R. & 8. Syst. 7: 286. 1826.

34a. Barbacenia flava var. flava

Caudex to 8 em. long; leaf-blades 13-20 cm. long, 8-12 mm. wide;
scapes equaling the leaves; perianth-tube 16 mm. long; tepals 10-12
mm. long.

Braziu: Minas Gerais: Itambé. Serra do Cipé.
34b. Barbacenia flava var. minor L. B. Smith, var. nov.

A var. flava omnibus partibus valde minoribus differt.

Caudex 1-4 em. long; leaf-blades 8-13 cm. long, to 8 mm. wide;
scapes shorter than the leaves; perianth-tube 15-20 mm. long; tepals
5 mm. long.

Type in the U.S. National Herbarium, No. 2340345, collected 344 miles from
Hotel Chapeu de Sol, Serra do Cipé, Municfpio Jaboticatubas, State of Minas
Gerais, Brazil, altitude 1110 meters (3700 feet), December 19, 1959, by Bassett
Maguire, C. K. Maguire and J. Murga Pires (No. 44665). Isotype in the New
York Botanical Garden. Paratype, No. 44691, same data (NY, US).

35. Barbacenia glauca Mart. ex Schult. f. in R. & 8S. Syst. 7: 288. 1826.

Brazit; Minas Gerais: Itambé.

36. Barbacenia itabirensis Goeth. & Henr. Blumea 2: 353. 1937.

Braziu: Minas Gerais: Pico d’Itabira do Campo.

37. Barbacenia sellovii Goeth. & Henr. Blumea 2: 359. 1937.

Braziu: Minas Gerais: Serra da Mooda (Moeda?). Serra da Piedade.
38. Barbacenia macrantha Lem. Jard. Fl. 4: pl. 390. 1854.

Brazit: Minas Gerais: Serra do Cipé.

39. Barbacenia rubro-virens Mart. Nov. Gen. & Sp. 1: 20, pl. 4, fig. 1. 1823.

BraziL: Minas Gerais: Diamantina.

IS CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

40. Barbacenia riedeliana Goeth. & Henr. Blumea 2: 35819. 37.
BraziLt: Minas Gerais: Serra da Lapa.

41, Barbacenia luzulifolia Mart. ex Schult. f. in R. & S. Syst. 7: 285. 1826.
Brazit: Minas Gerais: Ouro Preto.

42. Barbacenia viscosissima Goeth. & Henr. Blumea 2: 363. 1937.
Brazit: Minas Gerais: Séio Joio. Itacolumi. Ouro Preto (?).

43. Barbacenia paranaensis L. B. Smith, sp. nov. PuLare 10, ricurEs 25, 26

A B. viscosissima Goeth. & Henr., cui affinis, foliis ovarioque multo
majoribus, tepalis ellipticis latioribus, tubo perianthii ovarium triplo
solum longioribus differt.

Caudex over 2 dm. long, terete, 10-15 mm. in diameter at apex
(including leaf-sheaths); leaves densely imbricate; sheaths ovate,
amplexicaul at base, 3 cm. long, sulcate at apex and margins; blades
linear-lanceolate, filiform-acuminate, slightly attenuate toward base,
23-30 cm. long, 17-20 mm. wide, entire, finely nerved, green with the
mid-nerve white above, finely and subdensely glandular-pilose on
both sides; scape solitary, extending 10-12 cm. above the leaf-sheaths,
slender, densely glandular-pilose; perianth-tube 45-55 mm. long, about
3 times as long as the ellipsoid ovary, slightly contracted above the
ovary and narrowly campanulate, 15 mm. in diameter at apex, mi-
nutely stipitate-glandular, green (! Hatschbach); tepals reflexed at
anthesis, broadly elliptic, subacute and apiculate, 17 mm. long; fila-
ments broadly subtriangular with short acute appendages; anthers
linear, subbasifixed, 11 mm. long; style exceeding the stamens, broadly
ovoid at apex with 3 ovate stigmas.

Type in the U.S. National Herbarium, No. 2279892, collected on sandstone
(arenite) bluffs, Fazenda Morungava, Rio de Funil, Muniecfpio Sengés, State of

Parand, Brazil, December 15, 1958, by G. Hatschbach and R. B. Lange (No.
5359). Isotype in Herbdrio Hatschbach.

Braziu: Parand: Sengés: Itararé, Morungava, on shady banks in “campo
cerrado,” alt. 740 m., January 23, 1915, Dusén 16478 (F, GH); December 7, 1915,
Dusén 17379 (F, GH, fruit).

44, Barbacenia exscapa Mart. Nov. Gen. & Sp. 1: 21, pl. 14. 1823.

Braziu: Minas Gerais: Itambé,

45. Barbacenia gentianoides Goeth. & Henr. Blumea 2: 347, fig. M. 1937.
Brazil: Minas Gerais: Rio das Pedras by Valu. Serra da Lapa. As Dattas
to Parauna. Serra do Cipé (?).

46. Barbacenia hilairei Goeth. & Henr. Blumea 2: 351. 1937.
Brazit: Minas Gerais: No locality.

47. Barbacenia oxytepala Goeth. & Henr. Blumea 2: 357. 1937.
Brazit: Minas Gerais: Biribiri to Diamantina.

48. Barbacenia grisea L. B. Smith, sp. nov. PLATE 8

A B. oxytepala Goeth. & Henr., cui affinis, foliis longe ciliatis, scapo
et perigonii tubo densissime adpresseque griseo-tomentellis differt.

SMITH—SYNOPSIS OF AMERICAN VELLOZIACEAE 281

Caudex simple, erect, nearly 2 dm. high, covered with the charred
remains of old leaves and appearing 5 cm. in diameter; leaves many-
ranked, very densely imbricate; sheaths closely sulcate with the nerves
almost contiguous, covered with a thick resinous coat; blades linear,
attenuate to a filiform apex, 22 cm. long, 13 mm. wide, sulcate on both
sides, resinous-punctulate between the nerves, entire, glabrous except
the finely long-ciliate margins; scapes 2, 22-26 cm. long, with the peri-
anth very finely and closely gray-tomentellous especially toward apex,
scarcely glandular; perianth-tube subcylindric, 60 mm. long; tepals and
stamens imperfectly known; ovary 25 mm. long, 5 mm. in diameter,
attenuate at base, greatly enlarged in fruit.

Type in the New York Botanical Garden, collected 49 miles from Diamantina,
State of Minas Gerais, Brasil, altitude 1140 meters (3800 feet), December 22, 1959,
by Bassett Maguire, C. K. Maguire and J. Murga Pires (No. 44745).

49, Barbacenia glabra Goeth. & Henr. Blumea 2: 348, fig. H. 1937.
Brazit: Minas Gerais: No locality.

50. Barbacenia tomentosa Mart. Nov. Gen. & Sp. 1: 18, pl. 11. 1823.
Braziu; Minas Gerais: Ouro Preto.

51. Barbacenia schidigera Lem. Jard. Fl. 2: pl. 198. 1852.
Braziu: Minas Gerais: Pico d’Itacolumi.

52. Barbacenia graciliflora L. B. Smith, sp. nov. PLATE 10, FIGURES 27, 28

A B. schidigera Lem., cui affinis, tubo epigyno gracili, tepalis
subpatentibus acutis differt, et a B. luzulifolia Mart., tubo epigyno
graciliori subglabro differt.

Caudex simple or few-branched, 8-15 cm. long, 1 cm. in diameter
(including the leaf-sheaths); leaves very densely imbricate; sheaths
widely sulcate; blades linear, cuspidate-acuminate, slightly narrowed
toward base, 11-13 cm. long, 9 mm. wide, dull green, glabrous except
for the ciliate margins, rather broadly sulcate with the grooves wider
than the nerves; scapes 1-3, extending 4 cm. above the leaf-sheaths,
ca. 0.7 mm. in diameter, glabrous; perianth-tube 4 cm. long, more
than twice as long as the ellipsoid ovary and contracted above it to a
slender tube 3 mm. in diameter, obscurely costate, very sparsely
pubescent and appearing glabrous, red (! Williams) ; tepals subspread-
ing at anthesis, linear, acute, 26 mm. long, the outer finely pubescent;
filaments linear with minute triangular lobes, 16 mm. long, connate
in a slender tube for most of their length; anthers affixed about ye
above base, 10 mm. long; style slenderly clavate at apex.

Type in the Gray Herbarium of Harvard University, collected on rocks at
edge of stream, sandstone area, Serra de Monjolo, 18 kilometers north of Sérro,
Municipio of Sérro, State of Minas Gerais, Brazil, May 5, 1945, by Louis O.
Williams and Vicente Assis (No. 6816).

282 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

53. Barbacenia ensifolia Mart. ex Schult. f. in R. & S. Syst. 7: 287. 1826.
Braziu: Minas Gerais: Serra da Lapa.

54. Barbacenia glaziovii Goeth. & Henr. Blumea 2: 348. 1937.
Braziu: Minas Gerais: Biribiri to Diamantina.

55. Barbacenia conicostigma Goeth. & Henr. Blumea 2: 341, 1937.
Braziu: Minas Gerais: Serra da Lapa.

56. Barbacenia cuspidata Goeth. & Henr. Blumea 2: 342, fig. C. 1937.
Braziu: Minas Gerais (?): No locality.

57. Barbacenia longiflora Mart. Nov. Gen. & Sp. 1:19, pl. 12. 1823.
Braziu: Minas Gerais: Tejuco, near Diamantina. Guinda.

58. Barbacenia gardneri Seubert in Mart. Fl. Bras. 3, pt. 1: 70, pl. 8, fig. 5.
1847.
Braziu: Minas Gerais: Diamantina. Sérro.

59. Barbacenia curviflora Goeth. & Henr. Blumea 2: 342. 1937.
Brazit: Minas Gerais: Rio das Pedras by Valu.

60. Barbacenia fragrans Goeth. & Henr. Blumea 2: 345. 1937.
Brazit; Minas Gerais: Caldas. Sao Paulo: Sao Jo&io da Boa Vista.

61. Barbacenia tricolor Mart. Nov. Gen. & Sp. 1: 18, pl. 10. 1823.
Braziu: Minas Gerais: Serra da Tiradentes. Serra de Rola Moca. Serra do
Cipé.
62. Barbacenia blanchetii Goeth. & Henr. Blumea 2: 339. 1937.
Brazit: Bahia: “Igreja Velha.” Jacobina. Pougo d’Areia.
63. Barbacenia mollis Goeth. & Henr. Blumea 2: 356. 1937.
63a. Barbacenia mollis var. mollis.

Leaf-blades to 12 cm. long, 12 mm. wide.

Braziu: Minas Gerais: No locality.
63b. Barbacenia mollis var. microphylla L. B. Smith, var. nov.

A var. molli foliis multo minoribus differt.

Leaf-blades to 8 cm. long, 4 mm. wide; flower red (! Williams).

Type in the Gray Herbarium of Harvard University, collected on iron ore
slope, Serra da Mutuca, beyond Barreiro, Municfpio of Nova Lima, State of
Minas Gerais, Brazil, altitude 1400 meters, April 15, 1945, by Louis O. Williams
and Vicente Assis (No. 6641).

64. Barbacenia leucopoda L. B. Smith, sp. nov. PLATE 10, FIGURES 29, 30

A B. damaziana Beauverd, cui aliquid affinis, scapl indumento
pallido, floribus minoribus differt.

Caudex about 6 cm. long; leaves densely imbricate; sheaths broadly
ovate, 2 cm. long, glabrous, rather laxly nerved; blades linear-lanceo-
late, acute, 15 cm. long, 12 mm. wide at the middle, finely and densely
setose-glandular; scape solitary, 9 em. high above the leaf-bases,
slender, densely vestite with fine white gland-tipped trichomes; flow-
ers orange; perianth-tube subcylindric, very slightly enlarged upward,
25 mm. long, about 3 times as long as the ovary, very dark setose-
glandular; tepals suberect at anthesis, elliptic, obtuse, 15 mm. long;
filaments subtriangular, 7 mm. long, their appendages narrow, acute;

SMITH-—SYNOPSIS OF AMERICAN VELLOZIACEAE 283

anthers linear, 9 mm. long, exceeding the filament-appendages by
about %; stigmas small, apical, subcapitate.
Type in the herbarium of the University of California, No. 972105, collected in

campo, Serra do Cipd, Munie{pio of Mato Dentro, State of Minas Gerais, Brazil,
January 15, 1951, by Amaro Macedo (No. 2999).

65. Barbacenia blackii L. B. Smith, sp. nov. PLATE 10, FIGURES 31, 32

A. B. damaziana Beauverd, cui affinis, filamentis oblongis antheris
paulo superatis, foliis latioribus differt.

Caudex about 3 cm. long; leaves numerous, rather persistent,
densely imbricate; sheaths broadly ovate, 1 om. long, the lower half
glabrous except the glandular-ciliate margins, narrowly sulcate, am-
plexicaul at base; blades linear-lanccolate, slightly narrowed toward
base, cuspidate-acuminate, 10-13 cm. long, 9 mm. wide, finely and
densely setose-glandular; scape solitary, 6-10 cm. long, less than 1
mm. in diameter, finely and densely setosc-glandular; perianth-tube
3-4 em. long, slightly constricted and subcylindric above the ellipsoid
ovary and about 3 times as long, red, densely setose-glandular; tepals
suberect at anthesis, elliptic-oblong, obtuse, 14-17 mm. long; filaments
oblong with large lobes surpassing the centers of the anthers; anthers
linear, 8 mm. long, extending only slightly below the filament-sinus.

Type in the U.S. National Herbarium, No. 2221524, collected on the Serra do

Cipé, State of Minas Gerais, Brazil, January 13, 1951, by J. M. Pires and G. A.
Black (No. 2734).

Braziu: Minas Gerais: Mun. Mato Dentro: In campo, alt. 1600 m., January 15,
1951, A. Macedo 2962 (US).

66. Barbacenia rubra L. B. Smith, sp. nov. PLATE 10, FIGURES 33, 34

A B. damaziana Beauverd, cui affinis, tepalis late rotundatis apicula-
tisque, filamentis oblongis, antheris subbasifixis basi filamentorum
haud attingentibus differt.

Caudex unknown except for apex, this about 1 cm. in diameter (in-
cluding leaf-sheaths) ; leaves densely imbricate, sheaths broadly ovate,
ca. 15 mm. long, glabrous below the apex, sulcate, lustrous; blades
very narrowly triangular, filiform-acuminate, 14 cm. long, 5 mm. wide,
finely and densely setose-glandular; scape solitary 10 cm. long, very
slender, setose-glandular; perianth-tube 35 mm. long, cylindric above
the broadly obovoid ovary, 8 mm. in diameter, red (! Williams),
densely setose-glandular; tepals erect at anthesis, oblong, broadly
rounded and apiculate, 20 mm. long; filaments oblong with short lobes,
7 mm. long; anthers linear, 9 mm. long, subbasifixed, extending only
slightly below the filament-sinus.

Type in the Gray Herbarium of Harvard University, collected on rocky slopes,

Serra da Mutuca, Municipio of Nova Lima, State of Minas Gerais, Brazil, Feb-
ruary 1945, by Louis O. Williams (No. 5432a).

284 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

67. Barbacenia damaziana Beauverd, Bull. Herb. Boiss. u. 5: 1078. 1905.
Braziu: Minas Gerais: Pico d’Itacolumi.

68. Barbacenia bahiana L. B. Smith, sp. nov. Pate 10, FrGuREs 35, 36

A B. damaziana Beauverd, cui affinis, ovario basi longe attenuato
anguste obconico differt.

Caudex unknown except for apex, this about 7 mm. in diameter;
leaves 3-ranked, densely imbricate; sheaths closely sulcate with the
nerves almost contiguous; blades linear, attenuate to a filiform apex,
15 cm. long, 5-8 mm. wide, entire, sulcate on both sides, densely vestite
with long acuminate setae and much shorter stipitate glands; scape
solitary, extending 6 cm. above the leaf-sheaths, more glandular than
the leaf-blades; perianth-tube 4 cm. long, subdensely pale glandular,
slenderly cylindric above the ovary, 4 mm. in diameter; ovary slen-
derly obconic; tepals erect at anthesis, oblong, broadly subacute, 15
mm. long, 3 mm. wide, the outer minutely glandular, the inner nearly
glabrous; stamens slightly shorter than the tepals, filaments sub-
triangular with broad base and short appendages; anthers linear,
basifixed, 11 mm. long; style about equaling the tepals, broadly ovoid
at apex with 3 ovate stigmas.

Type in the U.S. National Herbarium, No. 762256, collected in the vicinity of

Machado Portello, State of Bahia, Brazil, June 19-23, 1915, by J. N. Rose and
P. G. Russell (No. 19924).

Excluded and Doubtful Taxa

Barbacenia gracilis hort. ex Baker, Journ. Linn. Soc. 18: 239. 1880. Nomen.
This horticultural name was published in the synonymy of Dasyli-
rion acrotrichum Zucc., a liliaceous species of Mexico.

Vellozia coerulescens hort. Belg. ex Gumbl. Gard. Chron. for 1874. 2: 623,
657. 1874.

According to Jackson, Index Kewensis 4:1173. 1895, this horti-
cultural name is a synonym of Barbacenia purpurea Hook.
Vellozia duidae Steyermark, Fieldiana Bot. 28, no. 1: 157. 1951.

The species is based on sterile material from southern Vene-

zuela. It is related to if not identical with V. tubiflora (A. Rich.)
H.B.K.

Vellozia hirsuta Goeth. & Henr. Blumea 2: 374. 1937.

This species was described from sterile material collected at Biri-
biri, Minas Gerais, Brazil.
Vellozia lanata Pohl, Pl. Bras. 1: 130. 1827.

This species was described from sterile material collected at Caldas
Novas, Goias, Brazil.

SMITH—SYNOPSIS OF AMERICAN VELLOZIACEAE 98h

Vellozia leptophylla Seubert in Mart. Fl. Bras. 3, pt. 1:84. 1847.

This species was based on a sterile collection by Sellow without
any indication of locality. Consequently there is not much hope of
its ever being clarified.

Vellozia scabra Spreng. Syst. 3:338. 1826.

The single line of description reads: ‘‘V. capsula glabra, scapo
hispido,” leaving little hope of ever identifying the species, although
effecting valid publication.

Vellozia squamata auctt. ex Steud. Nom. Bot. ed. 2. 2: 746. 1841. Nomen.

This name was published in the synonymy of Xerophyllum saba-
dilla, 1 Mexican liliaceous species of doubtful identity. It is scarcely
more than evidence of a misdetermination.

Vellozia tomentosa Pohl, Pl. Bras. 1: 130. 1827.

This species was described from sterile material collected between
the crossing of the Rio Jequitinhonha and Calumbi, Minas Gerais,
Brazil.

Vellozia triquetra Pohl, Pl. Bras. 1: 129. 1827.

This species was described from sterile material collected between
Inhumas and Quartel do Teixeira, District of Minas Novas, Minas
Gerais, Brazil.

Supplement

The following novelties were received too late to be included in the
foregoing revision, but are correlated with it by species numbers.
The Latin diagnosis gives the salient differences between each novelty
proposed and its nearest relatives in the key.

3a. Vellozia maxillarioides L. B. Smith, sp. nov. PuaTteE 11, FIGURE 37

A V. abietina Mart., cui affinis, ramis elongatis, vaginis foliorum
lepidibus erectis fimbriatis praeditis, ovario angustissime sulcato
differt.

Plant slender, branched, to 5 dm. high (! Maguire); branches ca.
4 mm. in diameter including the old leaf-bases; leaves few-ranked,
laxly imbricate, the complete ones 2-5 at the apex of the branch;
sheaths ca. 3 cm. long, tubular at base, very densely nerved, yellow-
brown, subdensely vestite with erect white fimbriate scales; blades
linear, abruptly acute and cuspidate, 4 cm. long, 3.5 mm. wide,
densely nerved, lepidote on the thickened yellow margins, elsewhere
glabrous, the keel somewhat prominent beneath; scape single, exactly
equaling the leaf-sheath so that the flower appears sessile, soon lateral;
capsule slenderly obconic, 10 mm. long, very densely and coarsely
ribbed, otherwise even and glabrous.

286 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Type in the U.S. National Herbarium, No. 2369002, collected on sandstone,
slopes and summit of Grao Mogul, State of Minas Gerais, Brazil, altitude 900-
1100 meters, August 17, 1960, by Bassett Maguire, Geraldo Mendes Magalhies
and Celia K. Maguire (No. 49268). Isotype in the New York Botanical Garden.

l6a. Vellozia fruticosa L. B. Smith, sp. nov. PuaTE 11, FicuRES 38, 39

A V. ambigua Goeth. & Henr. et. V. martiana Goeth. & Henr.,
cuibus ex descriptionibus affinis, caudice graciliore, foliorum laminis
glutinosis persistentibus, floribus multo minoribus differt.

Fruticose, much branched (! Maguire); caudex 7-8 mm. in diameter
including old leaf-bases; leaves few-ranked, subdensely imbricate, the
living about 5; sheaths ca. 2 cm. long, tubular at base, densely nerved
except the narrow brown margin, glabrous, not dividing with age;
blades linear, acute and long-cuspidate, 7 cm. long, 3 mm. wide,
sparsely and minutely serrulate on the keel beneath, on the margins,
and in two longitudinal lines above, the living very glutinous, erect to
spreading, the dead reflexed and evidently persistent for several
seasons; scape single, 7-8 cm. long, 0.8 mm. in diameter, nearly even,
sparsely and obscurely glandular toward apex; stamens 18, appendaged
at base; capsule ellipsoid, 8 mm. long, sharply trigonous with several
additional costae, stipitate-glandular on the costae.

Type in the U.S. National Herbarium, No. 2368988, collected on sandstone
outcrop, ca. 3-5 km. east of Serra, along road to Diamantina, State of Minas

Gerais, Brazil, August 9, 1960, by Bassett Maguire, Geraldo Mendes Magalhies
and Celia K. Maguire (No. 49130). Isotype in the New York Botanical Garden,

26a. Vellozia grao-mogulensis L. B. Smith, sp. nov. Pare 11, rigurEs 40-42

V. squalida Mart. ex Schult. f. in systema mea proxima sed habito
caespitoso humile, foliis angustioribus densioribusque, scapo densiu-
scule stipitato-glanduloso differt.

Plant cespitose; caudex short, much branched, ca. 8 mm. in diameter
including old leaf-bases; leaves rather many-ranked, very densely
imbricate, the living ones only 4 or 5 at the apex of the branch; sheaths
17 mm. long, almost completely covered; blades linear, attenuate, to
9 cm. long, 1.8 mm. wide, densely pale-lepidote on both sides at first,
the keel and thickened margins purple-black, the margins serrulate;
scapes 1 or rarely 2, 12-15 cm. long, very slender, subdensely stipitate-
glandular; ovary subglobose, 5 mm. in diameter, much enlarged in
fruit, evenly and subdensely dark-glandular; tepals subequal, elliptic,
obtuse, 17 mm. long, purple; stamens 18, appendaged at base, much
shorter than the tepals.

Type in the U.S. National Herbarium, No. 2368996, collected on sandstone,
Serra Grio Mogul, north base of mountain, State of Minas Gerais, Brazil, alti-
tude 600-700 meters, August 16, 1960, by Bassett’ Maguire, Geraldo Mendes

Magalhaes and Celia K. Maguire (No. 49218). Isotype in the New York Botan-
ical Garden.

SMITH—SYNOPSIS OF AMERICAN VELLOZIACEAE 287

27a. Vellozia bicolor L. B. Smith, sp. nov. PLATE 11, FIGURES 43-45

A V. aloifolia Mart., cui affinis, foliis bicoloribus, ovario grosse
stipitato-glanduloso differt.

Plant 3-5 dm. high (! Maguire); caudex to 5 cm. in diameter
(! Maguire); leaves many-ranked, very densely imbricate, the complete
ones about 9 at the apex of the caudex; sheaths ample, 55 mm. long,
densely nerved, white-lanate at apex, elsewhere glabrous, black with
brown margins, lustrous; blades soon deciduous, linear, acuminate,
30-35 cm. long, 2 em. wide, even, green, and glabrous above, com-
pletely cinereous-lanate beneath, the midnerve impressed above, prom-
inent beneath; scapes 1-2, 25 cm. long, obtusely trigonous, 4 mm. wide,
densely stipitate-glandular especially toward apex; tepals (only
remnants known) glandular-appendaged at base; stamens about 40;
capsule globose, 25 mm. in diameter, densely and coarsely stipitate-
glandular.

Type in the U.S. National Herbarium, No. 2368976, collected on rocky sandy
soil, summit of Serra do Cip6é, road from Hotel Chapeu de Sol between Km. 111
and 120, State of Minas Gerais, Brazil, altitude 1200 meters, August 6, 1960, by

Bassett Maguire, Geraldo Mendes Magalhies and Celia K. Maguire (No. 49037).
Isotype in the New York Botanical Garden. Same, No. 49047 (NY, US).

37a. Vellozia mollis L. B. Smith, sp. nov. Puate 11, rigurEs 46-48

A V. resinosa Mart. ex Schult f., cui affinis, foliis juvenilibus utrinque
molliter denseque albo-tomentosis, ovario e glochidiis dimorphis
dense glanduloso differt.

Caudex nearly 5 cm. in diameter including old leaf-bases; leaves
many-ranked, very densely imbricate, the complete ones about 12;
sheaths ample, 35 mm. long, densely nerved, dark castaneous, lustrous,
the apical half densely white-tomentose; blades linear, acute to sub-
obtuse, 13 cm. long, 8 mm. wide, at first densely white-tomentose, then
becoming more or less glabrous and green above; scapes 1-2 (?), 22
cm. long, obtusely trigonous, densely stipitate-glandular toward apex;
ovary subglobose, densely vestite with two types of glandular tri-
chomes, one short and conical, the other about twice as long with a
setose prolongation; stamens numerous; capsule 10 mm. long.

Type in the U.S. National Herbarium, No. 2368991, collected on sandstone
outcrop, ca. 3-5 km. east of Serra, along road from Conceigéo to Diamantina,
State of Minas Gerais, Brazil, August 9, 1960, by Bassett Maguire, Geraldo

Mendes Magalhaes and Celia K. Maguire (No. 49156). Isotype in the New
York Botanical Garden.

37b. Vellozia spiralis L. B. Smith, sp. nov. Puate 11, rigures 49-51

A V. resinosa Mart. ex Schult. f., cui affinis, vaginis foliorum apice
ciliatis alibi glabris, laminis basi ciliatis alibi glabris differt.

Plant to 2.5 meters high (! Maguire), over 3 cm. in diameter in-
cluding old leaf-bases; leaves few-ranked in a strong spiral, densely

288 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

imbricate, the complete ones about 20; sheaths ample, 4 cm. long,
densely nerved and dark castaneous except for the thin pale margin,
lustrous, disintegrating at apex and the whitish fibers recurving in a
tight coil, coarsely setose-ciliate toward apex, elsewhere glabrous;
blades linear, rounded and retuse, 30 cm. long, 11 mm. wide, setose-
ciliate at extreme base, erect-serrulate on the margins, densely nerved,
otherwise even and glabrous; scapes 3, 12-15 cm. long, 2.6 mm. in
diameter, sulcate, densely and finely stipitate-glandular toward apex;
ovary subglobose, 15 mm. long, subdensely and finely stipitate-
glandular; tepals subequal, elliptic, 45 mm. long, purple; stamens
ca. 40, the phalanges appendaged.

Type in the U.S. National Herbarium, No. 2369000, collected on sandstone,
slopes and summit of Grio Mogul, State of Minas Gerais, Brazil, altitude 900-

1100 meters, August 17, 1960, by Bassett Maguire, Geraldo Mendes Magalhées and
Celia K. Maguire (No. 49261). Isotype in the New York Botanical Garden.

38a. Vellozia coronata L. B. Smith, sp. nov. PuateE 11, FIGURES 52-54

A V. bremfolia Seubert, cui affinis, habito subacaule, foliis flori-
busque majoribus, ovario tubo brevi coronato differt.

Caudex simple or forked, 4 cm. long; 3 cm. in diameter including
the old leaf-bases; leaves very densely imbricate, the complete ones
4-5 at the top of the caudex; sheaths ample, 3 cm. long, dark
castaneous, glabrous, lustrous, the old ones deeply divided; blades
linear, narrowly obtuse and bicarinate near apex, to 16 cm. long,
15 mm. wide, very densely and finely serrulate-ciliate, otherwise
glabrous and nearly even, the midnerve impressed on both sides;
scapes at least 2, 9-15 cm. long, sulcate and subdensely stipitate-
glandular toward apex; stamens 18 or more, appendaged at base;
capsule subglobose, 12 mm. long, subdensely stipitate-glandular,
crowned with a short epigynous tube.

Type in the U.S. National Herbarium, No. 2368985, collected on sandstone
outcrop, ca. 20 km. from Concei¢éo on the road from Conceigéo to Diamantina,
State of Minas Gerais, Brazil, August 9, 1960, by Bassett Maguire, Geraldo

Mendes Magalh&es and Celia K, Maguire (No. 49114). Isotype in the New York
Botanical Garden.

39a. Vellozia epidendroides Mart. ex Schult. f. var. major L. B. Smith, var. nov.

A var. epidendroides habito majore, foliorum vaginis densiore
imbricatis, laminis longioribus differt.

Plant 1.5 meters high (! Maguire); sheaths of the leaves ca. 3 cm.
long, exserted less than 1 cm.; blades 12 cm. long.

Type in the U.S. National Herbarium, No. 2368982, collected on rock exposures,
summit of Serra do Cipé, along road from Hotel Chapeu do Sol, km. 120-140,
State of Minas Gerais, Brazil, altitude 1200-1300 meters, August 8, 1960, Bassett
Maguire, Geraldo Mendes Magalhiies and Celia K. Maguire (No. 49098). Isotype
in the New York Botanical Garden.

SMITH—SYNOPSIS OF AMERICAN VELLOZIACEAE 289

39b. Vellozia epidendroides Mart. ex Schult. f. var. divaricata L. B. Smith,
var. nov.

A var. epidendroides habito ramosiore, ramis divaricatis brevibus,
foliorum vaginis densiore imbricatis differt.

Plant 3 dm. high (! Maguire); branches 3-5 cm. long; sheaths of
the leaves exserted for 5 mm.; blades to 8.5 cm. long.

Type in the New York Botanical Garden, collected on rocks, sandstone terraces
and ridges, summit of Serra do Cipé, km. 112-128 along road from Hotel Chapeu
de Sol, State of Minas Gerais, Brazil, altitude 1200-1300 meters, August 1960,

by Bassett Maguire, Geraldo Mendes Magalhaes and Celia K, Maguire (No.
49064).

39c. Vellozia marcescens L. B. Smith, sp. nov.

A V. epidendroides Mart. ex Schult. f., cui affinis, foliis marcescen-
tibus, laminis reflexis persistentibus plus attenuatis, scapo breviore
differt.

Caudex erect, slender, mostly covered by old leaves; leaves few-
ranked; sheaths tubular, much exposed apically, very densely nerved,
yellow-brown, glabrous, lustrous; blades linear, attenuate to a nar-
rowly obtuse apex, closely nerved beneath, serrulate on the margins,
otherwise even and glabrous, the living few, erect to spreading, the
old blades reflexed, long-persistent; scapes 1-2, soon becoming lateral,
slender, at first shorter than the leaves, densely stipitate-glandular
toward apex; ovary subglobose, densely stipitate-glandular; tepals
appendaged at base; stamens about 24.

Vellozia marcescens var. marcescens Puate 11, rigures 55-57

Planta fere omnibus partibus majoribus, caudice ramoso, scapis
minoribus.

Caudex incompletely known but over 3 dm. high, branched, 5 mm.
in diameter (including the old leaf-sheaths); leaf-blades to 7 cm. long,
7 mm. wide; scapes 15-20 mm. long above the leaf-sheaths.

Type in the U.S. National Herbarium, No. 2368998, collected on sandstone,
slopes and summit of Serra Grao Mogul, Minas Gerais, Brazil, altitude 900-1100

meters, August 17, 1960, by Bassett Maguire, Geraldo Mendes Magalhaes and
Celia K. Maguire (No. 49259). Isotype in the New York Botanical Garden.

Vellozia marcescens var. minor L. B. Smith, var. nov.

A var. marcescente caudice simplici, foliis minoribus, scapis majoribus
differt.

Caudex simple, to 2 dm. high, 3 mm. in diameter including the old
leaf-sheaths; leaf-blades to 45 mm. long, 3.5 mm. wide; scapes to 35
mm. long above the leaf-sheaths, evidently elongating in fruit.

Type in the U.S. National Herbarium, No, 2368997, collected on sandstone,
Serra Griio Mogul, north base of mountain, altitude 600-700 meters, August 16,

290 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

1960, by Bassett Maguire, Geraldo Mendes Magalhdes and Celia K. Maguire
(No. 49219). Isotype in the New York Botanical Garden,

42a. Vellozia ciliata L. B. Smith, sp. nov. PLATE 11, riGuREs 58, 59

A V.cryptantha Seubert, cui affinis, laminis foliorum multo latioribus
basi dense longeque ciliatis, ovario obscure verruculoso differt.

Caudex to 15 cm. long, 1-2-branched (! Maguire), 4-5 em. in
diameter including the leaf-bases; (! Maguire), the complete ones
numerous; leaves many-ranked, very densely imbricate; sheaths
broadly ovate, over 25 mm. long, very densely nerved with a thin
nerveless margin, pale stramineous, glabrous, lustrous; blades linear,
subacute and cuspidate, 20 cm. long, 10 mm. wide, densely ciliate
toward base with long suberect yellow-brown setae, elsewhere glabrous,
the midnerve impressed above, prominent beneath; scapes 1-2 (?),
slenderly clavate, 2 cm. long; capsule obconic, truncate, 15 mm. long,
stramineous, lustrous, obscurely verruculose, hidden by the leaves.

Type in the U.S. National Herbarium, No. 2368992, collected on sandstone
outcrop ca. 3-5 km. east of Serra, along road from Conceigéo to Diamantina,
State of Minas Gerais, Brazil, August 9, 1960, by Bassett Maguire, Geraldo

Mendes Magalhies and Celia K. Maguire (No. 49158). Isotype in the New
York Botanical Garden.

62a. Vellozia pulchra L. B. Smith, sp. nov. PLATE 12, FIGURES 60-62

A V. cinerascente Mart. ex Schult. f., cui verisimiliter affinis, foliis
majoribus glabris, glochidiis ovarii apice subulato-acuminatis haud
divisis differt.

Somewhat branched shrub about 1 meter high (! Maguire); caudex
2 cm. in diameter (including the old leaf-sheaths) near apex; leaves
densely few-ranked; sheaths tubular at base, 3 cm. long, glabrous,
yellow-brown with a narrow white margin where covered, lustrous,
very densely nerved, remaining entire; blades many in the terminal
fascicle, linear-lanceolate, retuse at apex, to 27 cm. long, 15 mm. wide,
closely nerved, serrulate on the keel and margins, otherwise even and
glabrous; scape single, 18 cm. long above the leaf-sheaths, even,
glabrous; ovary broadly ellipsoid 15 mm. long, covered with suberect
slender contorted pale subulate-acuminate trichomes; tepals subequal,
elliptic-oblong, obtuse, 9 cm. long, 3 cm. wide, purple, finely pale-
scabrous toward base; stamens 6; anthers linear, 3 cm. long.

Type in the U.S. National Herbarium, No. 2369003, collected on granite slopes
immediately west of Pedra Azul, State of Minas Gerais, Brazil, August 21, 1960,
by Bassett Maguire, Geraldo Mendes Magalhies and Celia K. Maguire (No.
49289). Isotype in the New York Botanical Garden.
15a. Barbacenia brevifolia Taub, var. recurvata L. B. Smith, var. nov.

A var. brevifolia foliis recurvatis conduplicatis subduplo latioribus,
floribus majoribus differt.

SMITH-——-SYNOPSIS OF AMERICAN VELLOZIACEAE 291

Leaf-blades arching-recurved, conduplicate, ca. 8 mm. wide; flowers
4 cm. long.

Type in the New York Botanical Garden, collected on rocks, Guinda, Municfpio
of Diamantina, State of Minas Gerais, Brazil, November 5, 1937, by H. L. Mello
Barreto (No. 9526).
45a. Barbacenia magalhaesii L. B. Smith, sp. nov.

PLATE 12, FIGURES 63-65

A B. gentianoides Goeth. & Henr., cui valde affinis, foliis flores longe
superantibus subtus glabris, floribus aureis minoribus differt.

Densely cespitose herb, branches seldom more than 10 cm. long
(! Maguire); leaves numerous, persistent; sheaths ovate, 2 cm. long,
amplexicaul, glabrous, sulcate; blades linear, filiform-acuminate, to
16 cm. long, 9 mm. wide, closely and finely nerved, glabrous beneath,
subdensely glandular-tuberculate above, setose-ciliate on the narrow
dark scarcely thickened margins; flowers appearing sessile, glabrous,
yellow (! Maguire); perianth-tube slenderly cylindric, 4 cm. long,
3 mm. in diameter, flaring somewhat at apex; tepals suberect at
anthesis, linear, broadly subacute, 3 cm. long; stamens 25 mm.
long; filaments oblong with very short rounded lobes, high-
connate; anthers attached near apex of filament, nearly basifixed,
linear, 11 mm. long, purple; ovary slenderly ellipsoid, ca. 8 mm. long,
nearly glabrous.

Type in the U.S. National Herbarium, No. 2368981, collected on sandstone
rocks and ledges, road from Hotel Chapeu de Sol, Serra Cipé to Conceigaéo, km.
120-140, State of Minas Gerais, Brazil, altitude 1200-1300 meters, August 8,
1960, by Bassett Maguire, Geraldo Mendes Magalhaes and Celia K. Maguire
(No. 49094). Isotype in the New York Botanical Garden.

Brazit: Minas Gerais: Mun. Jaboticatubas: Serra do Cipé, Conceigéio do
Mato Dentro, Morro do Pilar, road to Pilar, October 16, 1953, F. Segadas Vianna
& J. Lorédo Jr. No. Serra-II-1066 (R, US). Frequent on rocks, sandstone
terraces and ridges, summit of Serra do Cipé, km. 112-128 along road from Hotel
Chapeu, alt. 1200-1300 m., August 7, 1960, Maguire, Magalhies & Maguire
49062 (NY).

45b. Barbacenia sessiliflora L. B. Smith, sp. nov. PLATE 12, FIGURES 66-68

A B. gentianioides Goeth. & Henr., cui parum affinis, foliis angusti-
oribus, tubo perianthii angustissime obconici, filamentorum lobis
triangularibus differt.

Plant only 5-10 cm. high but much branched; leaves numerous,
persistent; sheaths broadly ovate, 7 mm. long, glabrous, sulcate;
blades linear, acuminate, to 9 cm. long, 4 mm. wide, closely and
finely nerved, glabrous except for the setose-ciliate margins; flowers
sessile, red (! Magalhaes); perianth-tube 5 cm. long, very slenderly
obconic above the ovary, 5 mm. in diameter at apex, sparsely vestite
with fine eglandular trichomes; tepals spreading at anthesis, nearly

597430—_62—_6

292 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

uniform, oblong, 15 mm. long; filaments oblong, 4.5 mm. long, the
lobes triangular, acuminate; anthers linear, exceeding the filaments by
5 mm.; ovary obovoid, 10 mm. long.

Type in the New York Botanical Garden, collected on rock, Serra Cata Altas,
Municipio of Santa Barbara, State of Minas Gerais, Brazil, february 2, 1943, by

Geraldo Mendes Magalhies (No. 6400). Cotype: Same data as the type,
Magalhdes 6854 (NY).

59a. Barbacenia albiflora L. B. Smith, sp. nov. PLATE 12, FiguREs 69, 70

A B. conicostigma Goeth. & Henr., cui valde affinis, foliis utrinque
pilis glandulosis biformibus vestitis, floribus albis differt.

Stem short with multiple crown, very viscid (! Maguire); leaves
numerous, persistent; sheaths ovate, over 3 cm. long, closely and finely
nerved, glabrous; blades linear, filiform-acuminate, ca. 30 cm. long,
15 mm. wide, flat, laxly and coarsely setose and densely and minutely
glandular on both sides, the keel and margins inconspicuous; scapes
3 or more, ca. 50 cm. long, much exceeding the leaves, coarsely and
finely glandular intermixed especially toward apex; flowers white
(! Maguire); perianth-tube 4 cm. long, slenderly cylindric and laxly
and coarsely glandular above the ovary, densely and finely glandular
on the ovary; tepals irregularly reflexed at anthesis, 2 cm. long,
strongly dimorphic, the outer narrowly oblong, densely glandular, the
inner elliptic, glabrous; filaments oblong, 8 mm. long, the lobes short,
acuminate; anthers linear, 18 mm. long, extending below the base of
the filament; ovary ellipsoid, 15 mm. long.

Type in the U.S. National Herbarium, No. 2368995, collected on sandstone,
Serra Gréo Mogul, north base of mountain, State of Minas Gerais, Brazil, altitude

600-700 meters, August 16, 1960, by Bassett Maguire, Geraldo Mendes Magalhies,
and Celia K. Maguire (No. 49213). Isotype in the New York Botanical Garden.

CONTR. U.S. NAT. HERBARIUM VOL. 35, PART 4 SMITH -PLATE 1

Vellozia alata . B. Smith (Maguire & Pires 44690).

CONTR. U.S. NAT. HERBARIUM VOL. 35, PART 4 SMITH PLATE 2

FLORA OF BRAZIL
New York Boragical GARDEN

MUSEUM

Vellozia streptophylla \.. B. Smith (Gardner 5233).

CONTR. U.S. NAT. HERBARIUM VOL. 35, PART 4 SMITH—PLATE 3

‘WO Ol

Fellozta froesti L. B. Smith (Froes 19980).

CONTR. U.S. NAT. HERBARIUM VOL. 35, PART 4 SMITH —-PLATE 4

INSTITU TO A

PLANTAS DA AMAZONIA

Vellozia

90914
Serra do Gachimbo, 425m. de altitude.

Planta baiza, flér branoo-viclfoay
(orquidacea); em-savana sSbre pedras,

I.M-Pires, G,A.Hiack,J,J,Wurdack &
N.T-Silva 6423 17-Dez. , 1956

Vellozia bulbosa L. B. Smith (Prres et al. 6423).

CONTR. U.S NAT. HERBARIUM VOL. 35, PART 4 SMITH-—PLATE 5

ROE ANQES BRAZILIAN
359

» Sranil

Vellozia maguirei L. B. Smith (Maguire & Pires 44742).

CONTR. U.S. NAT. HERBARIUM VOL. 35, PART 4 SMITH —-PLATE 6

Y i =
“YY —
or “ d 2

s sa al l
fu

PLANTAS DA AMAZONIA

SOS10

Fellozia cachimbensts LL. B. Smith (Pires et al. 6232),

CONTR. U.S. NAT. HERBARIUM VOL. 35, PART 4 SMITH—PLATE 7

FD STATES NAMONAL MERBAReLN

davsenn L.G. amits

FLORA OF GOIAS, BRAZIL
Los Angeles County Museutt
MACHRIS BRAZILIAN EXPEDITION

Keyion of the Chapada dos Veadeiros ar
W. Long, 4° 40°, 8. Lar, Lae 30"

eadstone cuterop 7 km. south of Vesdeiros

e Dawson No 1¢$80 April 2aMex "1956

Vellozia dawsonii L. B. Smith (Dawson 14580).

CONTR. U.S NAT. HERBARIUM VOL. 35, PART 4 SMITH PLATE 8

Sane, ty Fe ar, ~

ie

Barbacenia grisea LB. Smith (Macuire 22 Pires 44745).

CONTR. U.S. NAT. HERBARIUM VOL. 35, PART 4 SMITH—PLATE 9

Figures 1-16.—-(All figures & 1, except 12 and 14 which are about X 5): 1, Vellozta viannae
L. B. Smith (Segadas-Vianna No. Serra 11-1103), flower. 2, V’. streptophylla L. B. Smith
(Gardner 5233), leaf-blade; 3, fruit. 4, VY. ramosissima L. B. Smith (Maguire & Pires
44666), apex of leaf-blade; 5, fruit. 6, /”. piresiana L. B. Smith (Black & Pires 51-12157),
apex of leaf-blade; 7, fruit. 8, V’. hypoxoides L. B. Smith (Macedo 4330), flower. 9, I’.
swallenit LL. B. Smith (Swallen 3617), flower; 10, phalange of stamens, ventral. 11,
PV. machrisiana L. B. Smith (Dawson 14674), fruit; 12, trichomes of ovary. 13, V. uleana
lL. B. Smith (Wle 8372), flower; 14, trichomes of ovary. 15, Barbaceniopsis boliiensis
(Baker) L. B. Smith, fruit (Schreiter 2561); 16, stamen, lateral (Bang 1134).

SPECIES OF BARBACENIA

Figures 17-36.—(All figures & 1): 17, Barbacenia trwiniana 1. B. Smith (Jrwin 2803),
flower with tepals cut away; 18, stamen, dorsal. 19, B. nizrimarginata, L. B. Smith
(Dawson 14717), section of leaf-blade; 20, flower; 21, stamen, ventral; 22, apex of style.
23, B. williams LL. B. Smith (Williams &F Assis 6696), flower; 24, stamen, ventral. 25,
B. paranaensis L. B. Smith (/atschbach & Lange 5359), flower; 26, stamen, dorsal. 27,
B. gracilifiora L. B. Smith (Williams <8 Assis 6816), flower; 28, stamen, dorsal. 29, BR.
leucopoda 1. B. Smith (Macedo 2999), flower; 30, stamen, ventral. 31, B. blackit L. B.
Smith (Pires & Black 2734), flower; 32, stamen, ventral. 33, B. rubra L. B. Smith
(Williams 5432a), flower; 34, stamen, ventral. 35, B. bahiana L. B. Smith (Rose &
Russell 19924), flower; 36, stamen, dorsal.

SMITH—PLATE 10

CONTR. U.S. NAT. HERBARIUM VOL. 35, PART 4

(For explanation see opposite page)

SPECIES OF VELLOZIA

Ficures 37-59.—(All figures *& 1): 37, Vellozia maxillarioides L. B. Smith (Maguire,
Magalhaes Maguire 49268), apex of branch. 38, V’. fruticosa L. B. Smith (Maguire,
Magalhaes ‘2 Maguire 49130), section of leaf-blade; 39, capsule. 40, 7”. grao-mogulensts
L. B. Smith (Maguire, Magalhaes <2 Maguire 49218), section of leaf-blade; 41, flower; 42,
phalange of stamens. 43, J’. bicolor L. B. Smith (Maguire, Magalhaes & Maguire 49037),
section of leaf-blade; 44, capsule; 45, phalange of stamens (remnant), 46, /. mollis L. B.
Smith (Maguire, Magalhaes ‘2 Maguire 49156), section of leaf-blade; 47, capsule; 48,
phalange of stamens (remnant). 49, J”. spiralis L. B. Smith (Maguire, Magalhaes &
Maguire 49261), apex of leaf-blade; 50, flower; 51, phalange of stamens. 52, V’. coronata
L. B. Smith (Maguire, Magalhaes © Maguire 49114), section of leaf-blade; 53, capsule; 54,
phalange of stamens (remnant). 55, /’. marcescens L. B. Smith var. marcescens (Maguire,
Magalhaes & Maguire 49259), section of leaf-blade; 56, capsule; 57, phalange of stamens
(remnant). 58, V’. ciliata LL. B. Smith (Maguire, Magalhaes Ff Maguire 49158), section of
leaf at junction of blade and sheath; 59, scape and capsule.

CONTR. U.S. NAT. HERBARIUM VOL. 35, PART 4 SMITH—PLATE 11

|

(For explanation see opposite page)

CONTR. U.S. NAT. HERBARIUM VOL. 35, PART 4 SMITH—PLATE 12

Figures 60-70.—(All figures X 1): 60, Vellozia pulchra L. B. Smith (Maguire, Magalhaes &3
Maguire 49289), apex of leaf-blade; 61, flower; 62, stamen. 63, Barbacenia magalhaesti
L. B. Smith (Maguire, Magalhdes ¢% Maguire 49094), section of leaf-blade; 64, flower; 65,
stamens. 66, B. sessilifora L. B. Smith (Magalhaes 6400), section of leaf-blade; 67, flower;

68, stamen, 69, B. albiflora L. B. Smith (Maguire, Magalhaes Maguire 49213), section
of leaf-blade; 70, flower

Index

(Synonyms are in /ftalves.)

217
217
251 (fig.)

Acicarpha spathulata,
Alternanthera maritima,
Amaioua guianensis, 222,
Amaryllidaceae, 252

» 204

Barbacenia— (con.)
flava, 279

var. flava,

var. minor,

279
279)

Amarvllidaceae, Tribe Vellozieae, 252 flavida, 276
Amar yllideae— Vellozieae, 252 foliosa, 276, 277
Androtrichum trigynum, 217 fragrans, 282
Anthurium, 220 fulva, 278
lacerdae, 220 gardneri, 282
pilonense, 220 gaveensis, 276
Aphaerema, 224 gentianoides, 280, 291
Araceae, 220, 221 (fig.) glabra, “8!
Arauearia, 227, 246 (fig.) glauea, 279
angustifolia, 227, 246 (fig.) glaziovii, 282
araucana, 227, 246 (fig.) globata, 278
Arenaria groenlandiea, 227 glutinosa, 278
Axonopus compressus, 222, 233 (fig.) goethartii, 277
Baccharis anomala, 225, 239 (fig.) gouneleana, 277
megapotamica, 225, 240 (fig.) graciliflora, 281
Balfourodendron riedeHanum, 223, 256 gracilis, 284
(fig.) grisea, 281
Barbacenia, 252, 271 hilairei, 280
albiflora, 292 hirtiflora, 278
alexandrinae, 270 ignea, 278
bahiana, 284 inclinata, 276
beauverdii, 276 irwiniana, 276
bicolor, 278 itubirensis, 279
blanchetii, 282 leucopoda, 282

blackii, 283
boliviensis, 271
brachycalyx x, 276
brasiliensis, 271,
brevifolia, 277
rar. brevifolia, 290
var. recurvata, 290
‘aricina, 276
castillont, 271
coccinea, 278
conicostigma, 282, 292
curviflora, 282
cuspidata, 282
damaziana, 282-4
ensifolia, 282
apa, 280

278

exsc

lilacina, 278

var. lilacina, 278

var. pallidiflora, 278
longiflora, 282
longiscapa, 276
luzulifolia, 280, 281
macrantha, 279
magalhaesii, 291
mantiqueirae, 277
markgrafii, 278
mollis, 282

var. microphylla, 2

var. molli, 282
nigrimarginata, 277
oxytepala, 280, 281
paranaensis, 280

R2

&2

VI

Barbacenia— (con, )
polyantha, 278
purpurea, 278, 284

var. minor,
riedeliana, 280
rogieri, 276
rubra, 283
rubro-virens, 280
schidigera, 281
schwackei, 278
sellovii, 279
sessiliflora, 291
276

276

seubertiana,
species, 278
squamata,
stenophylla, 276

tomentosa, 281

tricolor, 282

trigona, 278

tubtflora, 270

uaipanensis, 267

vandellii, 278

vandellit sensu Schult. f., 278
vargasiana, 271

Viscosissima, 280

williamsii, 278

276

Barbaceniopsis, 270
271
vargasiana, 271
Berberis, 227, 244 (fig.)
montana, 227, 244 (fig.)
Bernardia pulehells, 223, 22
Bromeliaceae, 252
Campderia, 253
Canavalia rosea, 216
Casearia silvestris, 222,
Chenopodium retusum,
Chlorophora tinectoria,
(fig.)
Climate effecting disjunction, 242
Conocarpus erecta, 216
Cordyline, 227

boliviensis,

223, 282 (fig.)
217
222, 223, 251

)

(fig.)

dracaenoides, 227, 246 (fig.)
Crinodendron, 227, 24 ( (fig.)
brasiliense, 227, 246 (fig.)

dependens, 246 (fig.)

patagua, 246 (fig.)
Dasylirion acrotrichum, 284
Diatenopterys sorbifolia, 224, 237 (fig.)
Disjunction of ranges, 242 (fig.)
Dunalia breviflora, 224, 237 (fig.)
Dyckia, 224, 238 (fig)

INDEX

Eriocaulaceae, 225
Esterhazya splendida, 225,
Kuphorbia sciadophila, 224
Herreria, 227, 245 (fig.)
stellata, 245 (fig.)
Hibiscus tiliaeeus, 216
Holocalyx balansae, 228, 2
Hydrocotyle bonariensis, 2
Hypericum, 226, 227, 245 (fig.)
)

239 (fig.)
| 237 (fig.)

24, 236 (fiz.)
16

connatum, 296, 243 > (fig.
gentianoides, 226, 227
mutilum, 265
perforatum, 226
Ipomoea pes-caprae, 216

Lagunecularia racemosa, 216
Liliaceae, 252
Lobelia camporum, 225
Migrations, 217 (fig.)
Paepalanthus, 225
Palmae, 220 (fig.)
Patagonula americana, 224, 237 (fig.)
Piteairnia, 228
Pleurostima, 271
purpurea, 278
Polygonum punetatum,
Proserpinaca palustris, 2
Protium heptaphyllum,
Pseudoplantago, 224
friesii, 223, 224, 236 (fig.)
Psychotria carthagenensis, 222
232 (fig.)
Quesnelia, 221, 229 (fig.)
imbricata, 229 (fig.)
Quiina, 222, 235 (fig.)
glaziovil, 222, 235 (fig.)
Radia, 253
Radia, Section of Vellozia,
Radia tubiflora, 270
Remirea maritime, 216,
Rhipsalis, 228
Rhizophora mangle, 216
Salicornia virginica, 216
Scaevola plumieri, 216
Schenekia, 224
Schnizleinia, 253
Sporobolus virginicus, 216
secundatum,

B22, 234 (fig.)
227
YOO

wee,

230 (fig.)

t
t
an

Teri

207

218 (fig.)

Stenotaphrum 216, 218
(fig)

Talbotia, 253

Tapirira guianensis, 222, 230 (fig.)

Tillandsia subg. Diaphoranthema, 226,

241 (fig.)

INDEX

Tillandsia usneoides, 226, 241 (fig.)
Topography effecting disjunction, 242
(fig.)
Triglochin striata, 217
Utricularia subulata, 227
Vantanea, 221
compacta, 221, 229 (fig.)
Vellozia, 252, 253
Vellozia Section Radia, 267
Vellozia Section Vellozia, 259
Vellozia abietina, 259, 285
alata, 260
albiflora, 265
alezandrinae, 270
aloifolia, 261, 287
alutacea, 260
ambigua, 261, 286
angustifolia, 261
annulata, 269
asperula, 263
var. asperula, 263
var. filifolia, 263
barbaceniifolia, 261
barbata, 262
bicolor, 287
boliviensis, 271
bradei, 264
brevifolia, 262, 288
breviscapa, 261
bulbosa, 266
cachimbensis, 268
cana, 270
candida, 267
caruncularis, 267
ciliata, 290
cinerascens, 266, 290
coerulescens, 284
compacta, 260
var. acutiflora, 260
var. obtusiflora, 260
coronata, 288
crassicaulis, 265
crassirama, 260
crinita, 267
cryptantha, 263, 290
dawsonii, 270
declinans, 262
duidae, 284
dumitiana, 269
echinata, 261
epidendroides, 262, 289
var. divaricata, 289
var. epidendroides, 288, 289

Vellozia (con.)

var. major, 288
exilis, 267
fibrosa, 260
fimbriata, 261
flavicans, 264
froesii, 265
fruticosa, 286
gardneri, 264
glabra, 253, 260
glandulifera, 262
glauca, 264

var. cujabensis, 264

var. genuina, 264

var. glauca, 264
glaziovii, 266
glochidea, 264
gracilis, 267
gramined, 259
granulata, 264
grao-mogulensis, 286
grisea, 267
hamosa, 264
hemisphaerica, 261
hirsuta, 284
hypoxoides, 265
incurvata, 260
intermedia, 261
lanata, 284
leptopetala, 262
leptophylla, 285
leucanthos, 268, 269
lithophila, 270
macarenensis, 268
macedonis, 259
machrisiana, 268
macrantha, 268
macrosiphonia, 268
maculata, 268
maguirei, 267
marcescens, 289

var. marcescens, 289

var. minor, 289
maritima, 267
markgrafii, 267
martiana, 261, 286
maudeana, 269
maxillarioides, 285
minima, 262
mollis, 287
ornata, 264
panamensis, 269
phalocarpa, 260

VII

VII

Vellozia— (con.)
phantasmagoria, 269
pilosa, 262
piresiana, 264
pleuroearpa, 262
plicata, 259
pulchra, 290
pumila, 265, 266
punctulata, 264
pusilla, 259
ramosissima, 263
resinosa, 262, 287
rhynchocarpa, 269
riedeliana, 267
scabra, 285
scoparia, 262
sellovii, 259
seubertiana, 265
species, 260
spiralis, 287
squalida, 261, 286
squamata, 264, 276
squamata auctt., 285
streptophylla, 262
sulphurea, 265
swallenii, 265
taxifolia, 262
tenella, 259
tertia, 267
tomentosa, 285
tragacantha, 259

INDEX

Vellozia—(con.)

triquetra, 285

tubiflora, 270, 284

uaipanensis, 267

uleana, 269

variabilis, 260, 261

var. tuberculata, 260
var. variabilis, 260

variegata, 262

velutinosa, 269, 270

verruculosa, 260

viannae, 261

virgata, 261

wettsteinii, 261
Velloziaceae, 224, 238 (fig.), 251, 252
Vellozieae, 252
Vernonia nitidula, 225, 240 (fig.)

platensis, 225, 241 (fig.)
Viviania, 227, 246 (fig.)
Visnea, 271, 278
Vochysia, 222, 234 (fig.)
Vriesea, 219
Xerophyllum sabadilla, 285
Xerophyta, 253

abietina, 259

plicata, 259

taxifolia, 262

tragacantha, 259
Xyris anceps, 228

capensis, 228

caroliniana, 222, 233 (fig.)

U.S. GOVERNMENT PRINTING OFFICE: 1962

UNITED STATES NATIONAL MUSEUM

CONTRIBUTIONS FROM THE UNITED STATES NATIONAL HERBARTUM

Votume 35, Parr 5

SYNOPSIS OF
THE SOUTH AMERICAN SPECIES OF
VISMIA (GUTTIFERAE)

By Josep KWAN

BuLLETIN OF THE Unrrep States NatTionaL Museum

SMITHSONIAN INSTITUTION ¢ WASHINGTON, D.C. * 1962

SYNOPSIS OF THE SOUTH AMERICAN
SPECIES OF VISMIA (GUTTIFERAE)

JosEPH EWAN

Introduction

Vismias are mostly low, leafy-crowned trees or tall shrubs, with op-
posite, broad, poplar-like leaves, and white, green, yellowish or brown-
ish flowers borne in more or less crowded panicles. The flowers recall
those of the genus Hypericum, the St.-John’s-worts, and, indeed,
Vismias were first described as species of Hypericum. The petals are
usually hairy or even woolly, but the amount of hairiness varies in
many instances among individuals of the same species. The usually
black berry of Vismia, subtended by a rather prominent calyx, early
attracted attention and provides a ready means of distinguishing this
genus from Hypericum, with which it agrees in most technical charac-
ters, but which has a dry capsular fruit.

The first Vismias described were based on collections originating
in Brazil and the Guianas. About four-fifths of the species of the
genus are confined to tropical South America and these constitute the
subject of the present synopsis. The few Vismia species known from
Central America have been reviewed in connection with this study;
they are mostly referred to the South American species but present
some special problems. The only species from Mexico and Central
America not otherwise mentioned in this treatment are V. mexicana
Schlecht. and V. camparaguay Sprague. Vismia demonstrates Afro-
American relationships, spanning as it does the South Atlantic Ocean,
with five or six species occupying tropical West Africa, and one species
isolated in the Pugu Range and near Dar es Salam on the coast of
East Africa. Adolph Engler, in the last summary of the genus as a
whole (1925), recognized about 30 species of Vismza but in all proba-
bility there are in the genus between 45 and 50 species. Knowledge
of the ecological relationships and life histories is scanty but there are
data that some species are giant forest trees whereas others are

293

294 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

characteristically early invading “weedy” species in disturbed
vegetation and do not usually attain true tree stature.

The present study was undertaken in 1946 at the suggestion of Mr,
E. P. Killip of the United States National Museum, while I was a
member of the Museum staff, and his continuing interest in the prob-
lem and the placing of supplementary collections in my hands for
study are appreciated. Work on the study was considerably advanced
during the summer of 1947 by a grant from the Smithsonian Institu-
tion. Since then Dr. Elbert Little, Jr., and Dr. Richard Evans
Schultes have made their Colombian collections available. The
kindness of Mr. N. Y. Sandwith in checking types at Kew early in
the progress of this study was particularly valuable. P. J. Eyma’s
careful notes on the types in European herbaria were often useful.
The photographs of type specimens preserved in European herberia
made by Dr. J. Francis Macbride under the Rockefeller Foundation,
together with fragments of authentic material now filed at the Chicago
Museum of Natural History, have been of decisive importance in
several instances. The use of three of these photographs to illustrate
this paper is an appreciated privilege. Opportunity to examine the
Vismia collections in person in European herbaria presented itself in
1954-55 when as a Guggenheim Fellow I visited 15 different institu-
tions primarily for the study of archive materials bearing on the
early botanical contacts between America and Europe. I am indeed
grateful to the John Simon Guggenheim Memorial Foundation for
this opportunity. The only important Vismia series not seen was
that of Martius in the Botanische Staatssammlung, Miinchen, but
duplicates of most of these were examined at Vienna and Paris.

It is a pleasure to acknowledge my indebtedness to several curators
of herbaria in this country and abroad for making their series of
Vismias available to me: Arnold Arboretum, Cambridge, Mass., (A);
British Museum (Natural History), (BM); Academy of Sciences, San
Francisco, Calif., (CAS); Botany School, Cambridge University,
(CGE); Herbario Nacional Colombiano, Bogot&, (COL); Botanical
Museum and Herbarium, Copenhagen, (C); Dudley Herbarium,
Stanford University, (DS); The Royal Botanic Garden, Edinburgh,
(4); Chicago Museum of Natural History, formerly the Field Museum,
(F) ; Herbarium Universitatis Florentinae, Florence, (FI) ;Conservatoire
et Jardin Botanique, Geneva, (G); Herbier de Candolle, Geneva,
(G-DC); Dept. of Botany, Glasgow University, (GL); The Herbarium,
Kew, (K); Rijksherbarium, Leiden, (LL); Missouri Botanical Garden,
(MO); National Arboretum Herbarium, Beltsville, Md., (NA);

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 295

Tulane University, New Orleans, (NO); Fielding Herbarium, Oxford
University (OXF); Muséum National d’Histoire Naturelle, Paris,
(P); Academy of Natural Sciences of Philadelphia, (PH); Natur-
historiska Riksmuseum, Stockholm, (S); Trinity College, Dublin,
(TCD); Botanical Museum and Herbarium, Utrecht, (U); University
of California, Berkeley, (UC); United States National Herbarium,
(US); Naturhistorisches Museum, Vienna, (W); and Botanisches
Institut and Botanisches Garten der Universitat, Vienna, (WU).

The citation of abbreviations following the citation of specimens
indicates that specimens have been studied by the writer in the
herbaria concerned. The few specimens cited without such abbrevia-
tions, mostly types, have not been seen.

Finally, I am sincerely grateful to C. V. Morton for a careful
scrutiny of this paper in editing it for publication.

Taxonomic History

The genus Vismia was established by Domingos Vandelli, Professor
in Lisbon, in a small quarto volume entitled ‘‘Florae lusitanicae et
brasiliensis specimen,” published in Coimbra, Portugal, in 1788.
Prof. Vandelli was first director of the Jardim Botanico da Univer-
sidade, Coimbra, from its founding in 1772 (or 1773?) until 1791.
He was succeeded by the better known botanist Felix de Avellar
Brotero (1744-1828), whose name is commemorated in the name of
the Coimbra garden’s bulletin, Broteroa. An examination of a copy
of Vandelli’s work (1788), in the Arnold Arboretum Library, shows
that the re-publication of this rare volume by J. J. Roemer (Scriptores,
pp. 67-164. 1796) is a verbatim textual copy and, as such, completely
reliable. The original generic description of Vismia appears in Van-
delli (1788) as shown in plate 1.

The illustration of the flower only, in his figure 24 which accompanies
the original description, shows the petals about equal to the sepals,
obovate, spotted, and ciliate all around. The illustration clearly
fixes the generic identity of the plant Vandelli had before him, but its
specific identity can hardly be determined because of its too gen-
eralized nature.

Vismia comprises the baccate-fruited species of Hypericum, includ-
ing Hypericum bacciferum, mentioned in 1648 in Georg Marcgrav’s
Historia naturalis Brasiliae! (p. 96). In 1760 Jacquin proposed
Hypericum cayennense, based on plants from French Guiana. In

! Published posthumously in a separately paged folio work by Willem Piso.

296 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

1775 Aublet described three additional species from French Guiana,
H. sessilifolium, H. latifolium and H. guianense. J. B. C. Frisee
Aublet (1720-1778) arrived in Cayenne on July 23, 1762, and remained
two years collecting plants of the country as he had done during his
previous nine-year sojourn in Mauritius. What is generally believed
to be Aublet’s principal herbarium is preserved at the British Museum
(Natural History) and the Vismias contained therein have been
studied by N. Y.Sandwith. A small collection of Aublet’s uncovered
at Carignan (Ardennes), France, apparently does not contain any
Vismia types.?

In 1764 the elder Linnaeus * published Hypericum petiolatum; it
was credited to Brazil, but without citation of earlier references.
Reichardt, who wrote the notable account of the genus for the Flora
Brasiliensis,* was unable to place Hypericum petiolatum L. Hypericum
petiolatum LL. is quite a different species, moreover, from the /.
petiolatum of Linnaeus the Younger,® a species based upon a collection,
communicated by Mutis, which without doubt originated in Colombia,
Authentic Mutis material is preserved in the United States National
Herbarium and was studied in the preparation of this synopsis.
Hypericum petiolatum Lf. is a homonym, however, and therefore not
available for this Colombian species, which must be called Vismia
lauriformis (Lam.) Choisy. <A little later (1790), Loureiro used the
name /Typericum petiolatum for the third time, quite independently
and without any reference to the two separate uses of the name by
Linnacus, father and son, and he thus created a third homonym.$
Choisy was justifiably in doubt that Lourciro’s name for a plant of
southeast Asia concerned the Brazilian Vismia previously described
as Hypericum petiolatum 1. Merrill, in his commentary” upon the
binomials of Loureiro,’ refers Hypericum petiolatum Lour. to Crato-
rylum ligustrinum (Spach) Blume (Guttiferae), as a synonym. In
1797 Lamarck added two more species to the genus under the generic
name Hypericum, H. rufescens and H. acuminatum. In 1797 Ruiz
and Pavon adopted the name Vismia and established two Peruvian
species under that name, V. glabra and V. tomentosa.

2J. Lanjouw and H. Uittien, Rec. trav. bot. neerl. 37 1; 133-170, 4 pls. 1940.
ISp. Pl. ed. 3. 1102. 1764,

4121: 210, 1878.

5 Supplementum 345. 1781.

6 Fl. Cochinch, 472. 1790 (Willd. reprint 577. 1793).

7 Amer. Philos. Soc. Trans. 24 ?: 268, 1935.

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 297

Subsequent to 1800, additions to Vismea were made, either as
transfers from Hypericum or as proposals under the name Vismia, by
the following 23 botanists:

Persoon (1807) Sagot (1880)
Poiret (1813) Hieronymus (1895)
Choisy (1821) Ruhland (1901)
Humboldt, Bonpland, and Kunth® Huber (1901, 1906)
(1822) Sprague (1905)
St. Hilaire (1827) Rusby (1912)
Schlechtendal and Chamisso (1828) Blake (1918)
Martius (1828) Hochreutiner (1919)
Miquel (1844, 1851) Mngler (1925)
Triana and Planchon (1862) Ducke (1939)
Turezaninow (1863) A. C. Smith (1939)
Reichardt (1878) Cuatrecasas (1946)

Jacques Denys Choisy, son of a minister in Geneva, studied under
Augustus Pyramus de Candolle and prepared a thesis entitled “ Pro-
dromus d’une monographie de la famille des Hypericineés,” which was
published in Geneva in 1821. The presentation copy to his master,
now in the Conservatoire Botanique library, contains a letter from
Choisy dated “1 Jan 1821,” which would indicate that the “little
work,” as Choisy called it, was actually printed in 1820.

Engler summarized the genus in the second edition of Engler and
Prantl’s Die natiirlichen Pflanzenfamilien,® recognizing at that time
the two sections in the genus founded by Reichardt in 1878, Trian-
thera and Huvismia (including Stictopetalum). N. Y. Sandwith, in
his studies of the British Guiana flora,’ revived Miquel’s name
Vismia angusta for a species that was up to that time confused with
either Vismia latifolia (Aubl.) Choisy or V. macrophylla H. B. K.
Pierre Joseph Eyma, a student of Prof. A. Pulle, critically reviewed
the Caribbean South American Vismias in the course of his doctoral
thesis at the University of Utrecht."

British, French, and American botanists have remarked upon the
need for realignment in the genus Vismia. A representative com-
ment is that of L. A. M. Riley who has said that since “many of the old
species [are] imperfectly known or misunderstood,”the genus is in

‘ Kunth wrote the botanical text of the Nov. Gen. et Sp. and perhaps should carry the credit alone, but
for bibliographic purposes and in accordance with historical tradition it is desirable to use the well-estab-
lished designation ‘*H. B. kK.”

9 21:185. 1925.

10 Kew Bull. 1931:174, 1931.

Polygonaceac, Guttiferae, and Lecythidaceae of Surinam. 1932. ef. pp. 48-53. Reprinted in Meded.

Bot. Mus. & Herb. Utrecht 4: 41-46, 1932. His tragic death is related by C. GQ. G.J. Van Steenis in Bull.
Bot. Gard. Buitenzorg, ser. III. 18: 403-406. portr. 1950.

298 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

“urgent need of revision.” ” Sagot made in effect the same comment
in 1880, and S. F. Blake wrote in 1918 that “the genus as a whole . . .
is rather badly in need of a thorough revision.” %

Though Adanson’s characterization of the genus as Caopia in 1763
is clear and unmistakable, founded as it is on the references to Bra-
azilian species in the works of Piso and Marcgrav, the generic name
Vismia has been admitted to the Nomina Generica Conservanda of
the International Code of Botanical Nomenclature, over the prior
name Caopia of Adanson. Vismia cayennensis (Jacq.) Pers. was
designated as the type species.

Economic Uses

Visma as a genus has no known commercial uses as a timber tree,
since the boles are generally too small for lumber or of too poor quality
for cabinet woods. However, some individuals of Vismias in Brazilian
Amazonia attain a height of 25 meters. Among the Andean peasant
farmers the genus is familiar as a conspicuous member of the second-
growth woodland or along the borders of cleared land, where it may
serve as an important species in erosion control because of its quick
growth. It is also a tree of the lowland selva and occasional in the
mountain rain forest. In both of these sites the trees flower more
inconspicuously and lack the rather large panicles of bloom character-
istic of trees growing in the open potreros. Certainly the large
number of vernacular names for the genus attest to the familiarity of
the trees among the natives over its wide range. Some of these
folk names refer to the characteristic yellow or orange resinous sap
which exudes rather freely from the cut stem. This resin is used in
local folk medicines. The juice of the bark of Vismia baccifera is
used in Boyaca, Colombia, by the Indians there to paint their faces
(teste A. Ki. Lawrance). Vismias having the same vernacular name
may be used for different purposes in different but adjacent regions.
Thus, “swinani” was reported to be used as a fish poison by Pichoto,
a Carib native, near Paramaribo, in November 1934 (W. A. Archer,
Ms. report, p. 377, in files of U.S. Agricultural Research Service),
whereas at Carolina, Surinam, “swinani” is not used as a fish poison
but as a medicine (Dec. 16, 1934, W. A. Archer report). Perhaps
it may be found that two different species of Vismia are involved
here under the name “swinani.” The bark of the Vismia is scraped
and the juice used to cure “ota,” a skin disease which Archer sug-
gests may be of a fungal origin. Rudolf Graves, a gardener of Para-
maribo, Surinam, reported to Archer that “pina” was used as a fish
poison there, the bark being scraped into the water. The orange

1 Kew Bull, 1925:134, 1925.
Contrib, Gray Herb, 53:42. 1918.

CONTR. U.S. NAT. HERBARIUM VOL. 35

EWAN-—PLATE 1

PoLYADELPHIA.

~ aurantinn * Laranget-
ra.
Vis. fruét. Corrobo-
rans , anthelmintica ,
pellens.

POLY ANDBRIA.
VISMIA. Fig. 24.

CAL. Per. coriaceum , mo-
nophyllum, inferum, 5-
partituum , perfiftens ,
foliolis ovato-acutis ,
concavis.

COR. Petala 5, ovata ,
intus pilofa.

STAM. filamenta plura in
5 phalanges s: corpora
connata, brevia, plana
incurva, apice multifida,
filamentis capillaribus ,

villofis: Anthere pluri-
mae, fubrotundae facie
externa filamenti fpi-
rae inftar difpofitae, s:
fecundae.

Neéftaria corpufcula s:
Glandulae 5. ovatae , pi-
lofae inter phalanges.

PIST. Germen fubrotun-
dum. Sty/f5. filiformes.
Stigmata peltata.

PER. Capfala 5-locularis,

entagona.

SEM. plurima. B.

Hypericum androfaemum.

~ quadrangulare.

— perforatum. Ipericad.

Vis. vulneraria , refol-
vens, anthelmintica.

— humifufum.

~ crifpum.

—~ montanum.

ORIGINAL DESCRIPTION OF THE GENUS VISMIA VANDELLI IN FLORAE LUSITANICAE
ET BRASILIENSIS SPECIMEN, P. 51. 1788.

CONTR. U.S. NAT. HERBARIUM VOL. 3! EWAN-—-PLATE 2

MOC y at

z “fr ,7
“ Gy. be fe.
/

a cre

VISMIA MACROPHYLLA H. B. K.

CONTR. U.S. NAT. HERBARIUM VOL... 35 EWAN—PLATE 3

Q{GS

th. Typus!

VISMIA PARVIFLORA SCHLECHT. AND CHAM.

CONTR. U.S. NAT. HERBARIUM VOL. 35 EWAN—PLATE 4

VISMIA RUFESCENS (LAM.) PERS PETIOLATE FORM OF V. SESSILIFOLIA

CONTR. U.S. NAT. HERBARIUM VOL. 35 EWAN—PLATE 5

TYPE CS.807 8 OF

er teow s@amiilfokamm Avbiet (Vi rouse

VISMIA SESSILIFOLIA (AUBL.) DC.

CONTR. U.S. NAT. HERBARIUM VOL. 35 EWAN PLATE 6

TYPE Ccliget ice
tyyertoum latifoiian acblet (¥)
a
*. dub oet,

Negative @ koe, 2 ue

Type Specimen
i .

VISMIA LATIFOLIA (AUBL ) CHOISY

CONTR. U.S. NAT. HERBARIUM VOL. 35 EWAN-—PLATE 7

Y

VISMIA BILLBERGIANA BEURL.

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 299

latex of V. baccifera subsp. ferruginea is said to color fabrics with a
fast color called ‘“carate colorado” (teste F. R. Fosberg).

VERNACULAR NAMES

Some of the more frequent vernacular names applied to species
of Vismia though without significant constancy for any one species
include the following:

achiotillo (Venezuela, Colombia, etc.) lacre (Amazonian drainage)
bloodwood (British Guiana) pichirina (northern Peru)
caopia (Brazil) punta de lanza (Colombia)
caparosa (Orinoco drainage) sangrito (Venezuela), Colombia
capianga (State of Bahia, Brazil) swinani (Carib name)

carate (Colombia)

Additional vernacular names will be found in the text below and in
S. J. Record and R. W. Hess’s ‘“Timbers of the New World,” p. 186.
1943.

GENERAL MorpHoLoGy AND DESCRIPTIVE TERMINOLOGY

LEAF: The leaf-blades of Vismia species vary from thick, shining
but concolorous structures on short stout petioles to thin-textured,
deltoid, long-petiolate blades with a strongly bifacial aspect. The
blades are mostly plane but may at times be plicate. Drip-tips are
well developed in forms of Vismia cayennensis. The secondary veins
may be weak to strongly impressed, camptodrome or simply diminish-
ing towards the margin. Dot-like glands may be borne singly and
centrally within an areole of fine veins, two or more may occur within
an areole, or they may be absent altogether. There is no correlation
between glandular-punctate leaves and vittate sepals or petals
occurring together within a given group of species. When a leaf-
blade is glabrous both above and beneath it is described as amphi-
glabrous, a term introduced in my studies of Delphinium (Univ.
Colorado Stud. ser. D. 2:78. 1945) and employed in my revision of
Macrocarpaea and elsewhere. The comparative anatomy of the
leaves of Vismia species should prove an engaging study.

Fiower: The inflorescence of Vismia is generally paniculate, but
the panicle may be supplemented by lateral few-flowered axes borne
in the upper leaf-axils. In Vismia cauliflora, the flowers are borne
directly on the stems on short stalks. The sepals often have a
scarious flange-like margin. The petals are generally comose within
and glabrous on the back; there may be dark glands on one or both
surfaces of the petals, called vittae, in which case the members are
described as vittate. The androecium provides the fundamental
basis for distinguishing subgeneric groupings. The stamens are
connate for about half their length into what I have called a stamen

300 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

column (“staminal bundle” of Eyma,'t “androphora” of Bentham,"
or ‘“phalange” of Reichardt '). The stamen column may persist
as an almost wire-like stiff structure in old flowers, but usually
shrivels after anthesis. The ovoid or globose anthers are included
and dehisce longitudinally. The ovary does not offer significant
differences usable in taxonomy. The five styles are generally coherent
but may at times spread like antennae,

Fruir: Vismia baceifera attracted the attention of the early
systematists by its heavy, fleshy, smooth-coated fruits, which led
to the recognition of the genus as distinct from the capsular-fruited
Hypericums. But most Vismias have rather inconspicuous spherical
to oblong berries with shriveled walls, these sometimes vittate with
wart-like epidermal appendages but usually unmarked.

Systematic ‘Treatment
Genus Vismia

Hypericum pro parte, sensu Jacq. (1760), L. (1764), L.f. (1781), Lamarck (1797).

Caopia Adans, Fam. 2:448. 1763.

Caspia Scop. Intr. 276, 1777.

Vismia Vandelli, Fl. Lus. Bras. Spec. 51, ¢. 3, fig. 24. 1788, nomen conservandum.

Acrossanthes Presl, Bot. Bemerk., in Abhandl. Boehm. Gesell. Wiss. V, 3:452.
1845.

Trees or shrubs, often with orange sap; leaves opposite, the blades
oblong or narrowly ovate to deltoid-ovate, acute to truncate or
cuspidate, short- to long-petiolate, exstipulate, glabrous or stellulate-
tomentose, sometimes ferruginously so, punctate-glandular or without
glands, sometimes lustrous on both surfaces; inflorescences terminal,
simply paniculate or corymbose-paniculate or with supplementary
lateral floriferous branchlets, or at times altogether lateral; sepals
generally ovate, more or less prominently scarious margined; petals
equaling or exceeding the sepals, often oblanceolate and densely
comose within with matted hairs, the petal-blades lineate, vittate, or
eglandular, glabrous on the back; stamens numerous, included, more
or less united half their length into 3 or 5 staminal columns, the
stamens tardily deciduous or persistent into fruiting stage, the anthers
globose or ovoid; ovary 5-celled, each locule with numerous, seldom
few ovules; styles 5, more or less free, at times prominently spreading
in age; stigmas capitate or subcapitate; fruit a berry, the pericarp
thick and fleshy or thin and withering, dark brown or blackish, some-

4 Eymain Pulle, Fl. Surinam 3:66. 1934.

'S Hooker, London Journ, Bot. 2: 371. 1843.
'6 Mart. Fl. Bras, 12!: 195. 1878.

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 301

times vittate with black scab-like glands, the calyx usually persistent
on the fruits.
Tyre (i.e. Standard Species): Vismia cayennensis (Jacq.) Pers.

Key to the Species

Flowers borne in sessile glomerules along the stems; leaf-blades pandurate-obovate,
large, 35-65 cm. long. . . . .. . . . IL ¥. cauliflora
Flowers paniculate; leaf-blades seldom as much as 5 35 cm. long.

Stamen column villous or comose, persisting as a stiff spreading appendage
after flowering; leaf-blades shallowly cordate at base (or merely rounded
in V. lateriflora and at times in V. angusta, | the veins prominently raised
beneath... . . .. . . . SUBKEY A

Stamen column not villous, ¢ or ‘if hairy then soon deciduous after flowering;
leaf-blades not cordate at base, the veins not very strongly raised beneath.

Flowers small, inconspicuous, usually very numerous (50-150 in an inflores-
cence or more, or only 5-8 in V. billbergiana); sepals less than 5 mm.
long, often less than half as long as the petals; inflorescence shorter than
the uppermost subtending leaves and hidden by them; leaf-blades finely
black-glandular-dotted beneath (except in V. rufa) . . . . Supkey B

Flowers larger, generally fewer (mostly 40-80 in an inflorescence, or more
numerous in V. tomentosa and V, lehmannit); sepals 5 mm. long or more,
at least half as long as the petals; inflorescence generally exceeding the
uppermost leaves or not hidden by them; leaf-blades with or without
glandular dots on the lower surface.

Leaf-blades glabrous or glabrate on both surfaces (the immature blades
sometimes puberulent in V. lauriformis) ; sepals 5-6 nm. long, spread-
ing or reflexed in fruit (ef. also V. amazonica and V. pentagyna).

SuBKEY C

Leaf-blades variously pubescent beneath, varying from felty-tomentose to
merely cinereous with a fine close puberulence under a lens (cf. also V.
rusbyi); sepals 7-10 mm. long, more or less erect in fruit (5 mm. long
in V. amazonica and V. pentagyna) . . . . . . . . . . . SuBKEY D

SUBKEY A

Rachises of the panicle only sparsely tomentulose with scattered flocs of stellate
hairs or subglabrous; leaves subglabrous or sparsely dark-tomentulose
beneath, the tomentum not obscuring the fine sessile black dot-like glands,
the blades usually lustrous above. .. . ... . . 2. V. macrophylla

Rachises of the panicle densely rufous-tomentose throughout; leaves more or less
copiously rufous-tomentulose beneath, the tomentum obscuring the glands
(if these are present), the upper surface dull.

Rachises and upper stems heavily rufo-pubescent with long, much-branched
hairs; leaf-blades both apiculate at apex and acute at base, long-petiolate.

3. V. sandwithii

Rachises pubescent but the hairs simple, rarely much-branched; leaf-blades
either apiculate at apex or acute at base but not both, rarely long-petiolate.
Panicles terminal, ample, openly branching; leaf-blades cordate at base,
thick, the principal nerves prominently raised beneath; petals comose,

the hairs matted, wholly obscuring the blade of the petal . 4. V. angusta

302 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Panicles axillary (sometimes both axillary and terminal, rarely terminal
alone), congested, compactly branching; leaf-blades truncate or subcordate
at base, rather thin-textured, the principal nerves conspicuous but not
raised beneath; petals ciliate-comose, the hairs not usually obseuring the
blade of the petal. . 2... ..... 2.4... . .5. V. lateriflora

SUBKEY B

Panicle cymose, few-flowered (5-8); leaf-blade apiculate, the cusp 1 em. long.

30. V. billbergiana

Panicle racemose, generally more than 8-flowered or if fewer then not cymose;
leaf-blade acute but not strictly apiculate.

Under-surface of the leaves rich ferruginous-tomentose; sepals about 4 mm.
long, felty-tomentose; stamen column very short, only 1.0-1.5 mm. long;
leaves felty-tomentose beneath, not finely black-glandular . .6. V. rufa

Under-surface of the leaves variously puberulent but never ferruginous-tomen-
tose; sepals mostly 1.5-2.0 (3.0) mm. long, not felty-rufo-tomentose; stamen
column more than 2 mm. long; leaves finely black-glandular beneath.

Nach stamen column bearing 3 or 5 stamens; leaf-blade ovate, lance-ovate,
or broadly ovate, hairy along the veins beneath; sepals without glands.
Stamen column triandrous; leaf-blades acute at base, hairy chiefly along
the veins beneath; fruit globose, coal-black, small (less than 5 mm. in
diameter) .. . an .. .. . T V. micrantha
Stamen column pentandrous; ‘leaf- blades rounde d or subcordate at base,
puberulent over the whole blade beneath with short, crisp, white hairs;

fruits ovoid, dull black, larger (more than 5 mm. in diameter).
8. V. brasiliensis
Hach stamen column bearing numerous stamens; leaves lanceolate to oblong-
lanceolate, not hairy along the veins beneath; sepals bearing glands.
Sepals 3- to 5-nerved, punctate with a few scattered black, dot-like glands;

petioles 10-12 mm. long... . .. . . .9. V. parviflora
Sepals with a single median raised conflue nt black gland; petioles 2-4 mm.
long . 2... ee ee ee eee ee. 2 10. Ve minutiflora

SUBKEY C

Principal leaf-blades mostly (3) 7-12 cm. wide; petals 8-14 mm. long, mostly
twice as long as the sepals or more (except in glabrous forms of V.
lauriformis).

Sepals perfectly glabrous even in bud (rarely hairy at the tips); leaf-blades

3-7 em. wide, ovate, less often lanceolate, acute at base, shining beneath.

43. V. sprucei
Sepals finely pubescent on the back; leaves 7-12 em. wide, dull beneath.
Leaf-blades lance-ovate, acute at base; flowers not at all showy, the petals
5-7 mm. long; glabrous form of . . . . . . . . 25. V. lauriformis
Leaf-blades broadly ovate or ovate-oblong, rounded at base; flowers some-
times rather showy, the petals 7-9 (14) mm. long.

Leaf-blades ovate-oblong, often punctate, thin-textured, commonly
wrinkling in drying, the veins prominently areolate, the secondary
veins 6-10 pairs.

Petals 11-16 mm. long, very finely lineate for their whole length, sparsely
vittate toward the tips; sepals 8-9 mm. long, the hyaline margin
nearly obsolete; panicles always terminal . . . 23. V. cuatrecasasii

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 303

Petals smaller, 7-10 mm. long, black-glandular toward the tips but not
finely lineate; sepals 4-5 mm. long, the hyaline margin distinct;
panicles either terminal or lateral... .. . . .42. V. obtusa

Leaf-blades broadly ovate, not punctate, firm-textured; veins anastomosing
at the margin, inconspicuously if at all areolate, with usually 12 pairs
of secondary veins or more.

Leaf-blades usually less than 15 cm. long, acuminate at apex, brownish,
golden or silvery beneath, more or less contrasting with the green
upper surface; petals glanduliferous . .. . . .24. V. baccifera

Leaf-blades usually 15-20 cm. long, abruptly acute at apex, sooty or
ashy beneath, hardly contrasting with the dark olive-green upper
surface; petals not glandullferous, usually merely vittate or dark
lineate. . ... ... . .41. V. confertiflora

Principal leaf-blades mostly less than 5 cm. wide: petals less than 8 mm. long,
about half again as long as the sepals.

Sepals rusty or ashy with a close felt-like tomentum on the back (cf. also
V. glabra with often thinly floccose-hairy buds); leaf-blades lanceolate,
gradually acuminate, pinnately veined or areolate; petioles 1-2 cm. long.

Leaf-blades areolate-veined, thin-textured, dull on the upper surface, ovate.
36. V. glaziovii

Leaf-blades pinnately veined, firm-textured (usually not folding or wrinkling
in drying), shining or dull above, generally lanceolate, often narrowly so.
Leaf-blades acute or acuminate, dull or less often shining and dark green
on the upper surface; panicle not compact-pyramidal; widespread

species ... .... .2%. V. guianensis
Leaf-blades often shortly. cuspidate, lustrous upper surface; panicle
compact-pyramidal; eastern Brazil. . . . . .35. V. reichardtiana

Sepals glabrous on the back (sometimes tomentulose in V. plicatifolia); leaf-
blades ovate or lanceolate, the veins usually prominently areolate; petals
always vittate-glandular outside; petioles 1 cm. long or less (1.0-1.5 em.
long in V. glabra subsp. pozuzoensis and V. laevis).

Leaf-blades lanceolate to oval, long-acuminate or merely acute but not
apiculate, plane; fruit ovoid to fusiform, tapering to often persistent
styles.

Leaf-blades 5-7 em. wide, acute, coriaceous, the midrib stout; branchlets
stout, stiffly spreading, the knoblike joints of pedicels prominent.

18. V. laevis

Leaf-blades 3.5-4 cm. wide, long-acuminate, firm but hardly coriaceous,
the midrib not prominent; branchlets slender, the pedicels inconspicu-
ously jointed.

Leaves spreading, the blades lanceolate, shortly acute or rounded at
base; petioles 6-10 mm. long. . . .19. V. glabra subsp. glabra
Leaves ascending, the blades oval, cuneate at base; petioles 10-15 mm.
long ... . .. . .19a. V. glabra subsp. pozuzoensis

Leaf-blades ovate, shortly ‘acute or apiculate, plane or plicate; fruit oblong
to subglobose.

Tree 3-10 (35) m. high; internodes not noticeably short; leaves only
moderately if at all plicate, generally markedly glandular-punctate
beneath, the 1 to few black dot-like glands prominent within each
areole ... . . .. .17. V. cayennensis

Shrub or low tree 2- 30 m, “high; internodes short; leaves strongly plicate,
usually finely glandular-punctate beneath, the glands rather faint in
theareoles. . ............ 4... .20. VY. plicatifolia

304 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

SUBKEY D

Calyx gibbously enlarged below, constricted above, the sepals all erect, the
alternate sepals prominently hyaline-margined; petals included, short, 5-7
mm. long, tardily expanding. . . . res OP v. urceolata

Calyx not urceolate; petals distinctly longer than ‘the sepals, expanding early.

Sepals broadly deltoid, 6-8 mm. wide at base, thick, proportionately narrowly
hyaline-margined, hard or corky-ribbed in fruit; flowers few, mostly 5-12
in a rather compact short-pedunculate panicle. . . .12. V. japurensis

Sepals oblong or ovate, not deltoid; panicle more than 15-flowered or if fewer-
flowered then not compact.

Membranous border of sepals conspicuous, wide, ciliolate; sepals 5 mm. long.
40. V. amazonica
Membranous border of sepals inconspicuous or if evident then not ciliolate;
sepals 7-10 mm. long.
Panicle freely and shortly branching, diffuse, with supplementary short
lateral upper clusters, ample, 100-140-flowered.

Sepals felty-tomentose, the tomentum dense dark chocolate-brown;
flowers 10-12 mm. long; leaf-blades 18-24 cm. long, 10-13 em.
wide, the lower surface rich red-brown, finely hairy (under a lens)
with discrete rufous hairs... . . . .29. V. tomentosa forms

Sepals pale- or yellow-green-pubescent; flowers 8-9 mm. long; leaf-blades
9-15 em. long, 5-7 em, wide, the lower surface silvery or yellowish,
cinereous-puberulent beneath. . . . . . . . . 22. V. lehmannii

Panicle not both freely and compactly branching or if so then less ample,
fewer than 100-flowered, the flowers generally 10 mm. long or more.
Uppermost floral leaves reduced (except in V. tomentosa and V.
mandurr).

Outer surface of sepals permanently felty with a thick tomentum (ef.
also V. lateriflora); petioles of even the uppermost leaves noticeably
long (1-2 em. long); sepals of fruiting calyx reflexed (spreading in
V. panamensis).

Leaves permanently felty-tomentose, thick-textured; sepals tomentose
with a thick raised dark chocolate-brown tomentum,
Leaf-blades narrowly lanceolate or oval, acuminate, 3-5 em. wide,
Waxy-shining above, the veins obscure beneath . 32. V. crassa
Leaf-blades ovate to broadly ovate, shortly acute, 6-11 em, wide,
dull above, the veins prominent beneath, areolate,
29. V. tomentosa
Leaves more or less densely tomentose beneath when young, thinly
tomentose in age, thinner in texture and wrinkling; sepals
tomentose with an appressed brown or gray tomentum.
Leaf-blades gray-tomentulose beneath, oval to elliptic, acute at
base; pubescence of rachises and pedicels straw-colored; panicle
cymose, compactly branching, with 3 axes or more.
26. V. mandurr
Leaf-blades reddish-tomentulose beneath, ovate to oblong-ovate,
rounded at base or if acute not also cinereous; pubescence of
rachises and pedicels rufous; panicle more or less pyramidal
with one principal axis.

Flowers large, the petals nearly twice as long as the sepals, 14
mm, long, 5-6 mm. wide; leaf-blades 15-18 cm. long, glossy
above; endemic species of central Colombia.

28. V. cavanillesiana

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 305

Flowers much smaller, the petals little exceeding or at most 1.5
times as long as the sepals; leaf-blades 7-12 cm. long or
if 15-18 em. long or more then not glossy above.

Leaf-blades 12-22 em. long, 6-10 em, wide; petals hardly ex-
ceeding the sepals; lowland forest of Pacific coast of
Panama and northern South America.

14. V. panamensis

Leaf-blades smaller, 7-12 em. long, 5-8 (10) cm. wide; petals
1.5 times as long as the sepals; species of eastern Brazil.

Leaf-blades broadly ovate, shortly acute; margins of calyx-
lobes vittate with black glands; panicle subtended by
many small supplementary leaves . . 33. V. martiana

Leaf-blades oval to narrowly ovate, acuminate; margin of
calyx-lobes without glands; panicle subtended by few if
any small supplementary leaves . 34. V. magnoliifolia

Outer surface of sepals glabrous to variously puberulent but never with
a thick felt-like tomentum, the veins or wrinkles clearly visible
beneath the pubescence (ef. also V. mandurr); petioles mostly less
than 1 em. long (longer in V. baccifera); sepals of fruiting calyx
spreading or appressed against the fruit (except in V. baccifera).

Mature leaf-blades 3-6 (8) cm. wide.
Leaves densely rufous-tomentose beneath with stellate or branched
hairs at least when young.

Leaf-blades broadly ovate, shortly acute; margins of calyx-lobes
black-glandular-vittate; panicle foliose . . 33. V. martiana

Leaf-blades narrowly ovate, acuminate; margins of calyx-lobes
without glands; panicle without supplementary leaves.

34. V. magnoliifolia
Leaves all merely cinereous or yellowish with a close puberulence
beneath, sometimes appearing glaucous (reddish tomentulose

in forms of V. lauriformis).

Branches of the panicle glabrous, puberulent, or with occasional
flocs of short-branched hairs; leaf-blades ovate, elliptie- or
lance-acuminate, the petioles slender.

Principal leaf-blades ovate to ovate-elliptic, rather abruptly
acuminate, sometimes plicate, not at all faleate; sepals
of the fruiting calyx strongly reflexed.

Petals not vittate; leaves strongly bicolored, dull above,
conspicuously glandular-punctulate beneath.

37. V. rusbyi

Petals vittate; leaves not strongly bicolored, shining above,
not punctulate (or obscurely so on the youngest blades),
appearing more or less glaucous beneath.

38. V. pentagyna

Principal leaf-blades lance-acuminate, more or less long-taper-
ing to aslender tip, not at all plicate; sepals of the fruiting
calyx spreading.

Leaf-blades more or less falcate, dull above, glandular-
punctulate beneath; petals black-glandular-vittate.

16. V. falcata

Leaf-blades ovate or narrowly lanceolate, not at all faleate,
lustrous above, not punctulate beneath; petals not
glandular-vittate.

306 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Panicle many-flowered, compact-pyramidal; veins closed-
areolate, distinct; petals densely villous-tomentose
within. . . .... .. . .35. V. reichardtiana

Panicle few- and loosely-flowered; veins not areolate or
weakly so; petals only thinly tomentose within.

39. V. buchtienii

Branches of the panicle ferruginous-tomentose throughout with

branched hairs; leaves elliptic or lanceolate, not faleate, the
petioles generally short, rather stout.

Leaf-blades oval or elliptic, 8-12 cm. long, 5-8 em. wide, acute
or abruptly acuminate. rather prominently areolate; fruit
usually oblong, the sepals spreading to strongly reflexed.

Leaf-blades tapering equally at both ends or only shortly
acuminate, strictly elliptic, 5-6 em. wide; petioles 1-1.5
cm. long; sepals in fruit strongly reflexed.

31. V. lindeniana

Leaf-blades rather rounded at base, acuminate at apex, 6-8
cm. wide; petioles less than 1 cm. long; sepals in fruit
spreading or somewhat reflexed . .11. V. sessilifolia

Leaf-blades lanceolate, 8-12 cm. long, 3.5-4.5 em. wide, taper-
ing to an acuminate apex, not prominently areolate; fruit
globose, the sepals not strongly reflexed.

21. V. viridiflora
Mature leaf-blades mostly broader, 7-13 cm. wide.
Leaves finely but densely punctate with minute dots . . 13. V. latifolia
Leaves not punctate beneath.
Leaf-blades uniformly rufous-tomentose beneath with stellate or
branched hairs at least when young . 34. V. magnoliifolia
Leaf-blades all merely cinereous or yellowish puberulent beneath

(or thinly tomentulose in V. baccifera).

Flowers not at all showy, the petals 5-7 mm. long; leaf-blades
lance-ovate, acute at base, yellowish or cinereous beneath
with a very fine close puberulence . . 25. V. lauriformis

Flowers rather showy, the petals 9-10 mm. long (7 mm. in V.
baecifera subsp. subcuneata); leaves mostly broadly ovate
(narrowly ovate in V. baccifera subsp. ferruginea), sub-
truncate or rounded at base, ferruginous, cinereous or
canescent beneath.

Leaf-blades uniformly acuminate, tomentulose beneath with
fine but discrete stellate hairs (as seen under a lens);
petals glanduliferous with irregular black dot-like or
welt-like glands. ....... . .24 V. baccifera

Leaf-blades abruptly acute at apex, cinereous-puberulent
beneath with minute simple hairs; petals usually non-
glandular, often evenly lineate with fine dark lines
extending to the tips of the blades . 41. V. confertiflora

1. Vismia cauliflora A. C. Smith, Journ. Arn. Arb. 20: 299. 1939.

Tyre; Along the road to Aleixo, Munic. Mandus, basin of Rio Negro, Ama-
zonas, Brazil, Krukoff 7947 (NY; isotypes, A, BM, G, K, MO, 8, U). Paratype
from Livramento, Munic. Humaytd, basin of Rio Madeira, Amazonas, Brazil,
Krukoff 6976 (A; dupl. paratypes, BM, G, K, 8, U, US).

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 307

ADDITIONAL SPECIMENS EXAMINED:
BRAZIL: Amazonas: Colonia Jofo Alfredo, Mandus, Ducke 930 (US);
Estrada do Aleixo, Ducke 25055 (I).

Shrub-like tree 4-8 m. high, often with few, short branches. In
some particulars this Vismia is the most distinctive species in the
genus, though evidently rarely collected. Its affinities are clearly with
V. angusta, but, as Ducke has commented," the sessile glomerate in-
florescence alone would easily separate it from all other species. The
striking leaves with blades measuring up to 65 cm. long, or longer
than originally described, display an unusual shape recalling those of
Ficus lyrata (F. pandurata); they are evenly narrowed for their lower
half and shallowly cordate at the very narrow base. The common
name is reported as “lacre.’’

2. Vismia macrophylla H. B. K. Nov. Gen. & Sp. 5: 184. 1822. PLATE 2

Caopia macrophylla Kuntze, Rev. Gen. Pl. 1: 59. 1891.

V. macrophylla var. glabrescens Hochr. Ann. Cons. Jard. Genéve 21:53. 1919.
Type from ‘Prope San Gabriel de Cochoeira, ad Rio Negro, Brasiliae
borealis,” Spruce, Jan._Aug. 1852 (isotypes, BM, E, F, FI, Boissier at
G, NY, P, TCD, W).

Type: Banks of Rio Cassiquiare, Amazonas, Venezuela, Humboldt & Bonpland
(P; isotype B, numbered 1151, Photo FM 9497); ef. Sandwith, Kew Bull. 1931:
174.

ADDITIONAL SPECIMENS EXAMINED:

SURINAM: Tawa Creek, Saramacca River headwaters, Maguire 23766 (NY,
U, US); Groningen, 10 May 1916, Samuels (US).

BRITISH GUIANA: Aruka River, Barima, Anderson 40 (NY); Waini
River, Northwest District, de la Cruz 3841 (NY, UC, US); Kamakusa, Upper
Mazaruni River, de la Cruz 2115 (UC, US), 2383 (US), Jenman 5324 (BM, NY);
Rockstone, Gleason 480 (US), 551 (US); Potaro River, Tumatumari, Gleason 340
(US); Essequibo River Basin, A. C. Smith 2162 (U, US); Barima River, de la Cruz
3382 p.p. (US); Upper Rupununi River, de la Cruz 1446 p.p. (US), 1744 (U8);
Malali, Demarara River, de la Cruz 2614 (UC, US); sand reef at head of Hoorubia
Creek, southeast of Georgetown, Hitchcock 16950 (8, US); Demerara River,
Jenman 5035 (BM); Good Hope, Persaud 44 p.p. (F); Berbice, Schomburgk 405
p.p. (CGE, F, FI, OXF, W); Kabakaburi, Pomeroon District, de la Cruz 3255
(NY, UC, US).

VENEZUELA: Zutta: Rio Lora, Pittier 10961 (US). TAcurra: San Félix,
Curran & Haman 1011 (US). Botfvar: Rfo Karuai, west of La Laja, 1220 m.,
‘“‘minchu-warei-yek,”’ Steyermark 60784 (F, US); wooded slopes of Quebrada
O-paru-m4, between Santa Teresita de Kavanayén and Rio Pacairao, 1065-1220
m., Steyermark 60434 (US); Roraima, Schomburgk 560 (P), 815 (BM, W). Ama-
ZONAS: San Carlos, 100 m., Holt & Gehriger 331 (US).

COLOMBIA: Botfvar: Palotal, Romero C. 1158 (US). SANTANDER: Que-
brada Angulo, 4 km. south of Lebrija, 955 m., St. John 20589 (NO, US). Meta:
Near junction of Rio Giiejar and Rio Zanza, Cordillera Macarena, 500 m., Smith
& Idrobo 1464 (US); 20 km. southeast of Villavicencio, Killip 34245 (US); Cano
Ciervo, Sierra Macarena, 600 m., Philipson et al. 2088 (US). Vaupgs: Mitt, Rio

1” Arquiv. Serv. Florest. 1: 34. pl. 13, figs. a, b. 1939.

598830-—62——2

308 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Vaupés, Cuatrecasas & Pérez-Arbeldez 6750 (N Y). Caquets: Solano, near Tres
Esquinas, Little 9783 (NO, US). Amazonas: Leticia, Schultes & Black 46-317
(US). Cxocé: Darien region, Dawe 871 (K); Palestina, Rio San Juan, 5-50 m.,
Cuatrecasas 16903 (US).

BRAZIL: Amazonas: Ega, Poeppig 2905 (F, W); along Rio Castanho, tributary
of Rio Padauiri, Upper Rio Negro Basin, 100-140 m., Cardona 1418 (US); Sao
Paulo do Olivenga, “lacre,”” Ducke 392 (A, F, US); San Gabriel de Cachoeira,
Spruce (F, NY); Cachoeira Caranguejo, Rio Canabury, Holt & Blake 537 (US).
Maro Grosso: Rio Jatuarana, Rio Machado region, Krukoff 1692 (A, 8).

Vismia macrophylla was widely misunderstood up to Sandwith’s
commentary on the species (Kew Bull. 1931: 174), in which he pointed
out that Hochreutiner had misinterpreted the species and introduced
a var. glabrescens “which agrees well with the type [of V. macrophylla].”
The two sheets of the Spruce collection that served as the basis of var.
glabrescens show the distinct black punctate glands of the lower leaf
surface and the thinly tomentulose rachis of the panicle of Vismia
macrophylla.

“Oralli” is the forester’s name for Vismia macrophylla in British
Guiana and one collection (Mazaruni Station, Fanshawe F627, US)
is annotated “low branchy tree peculiar to swampy savannahs on
white sand where it is co-dominant with Tabebuia longipes.”

A West Indian collection of this species labeled in Walker Arnott’s
hand merely “St. Vincents (Dr. Hooker)” in the Glasgow Herbarium
is almost certainly a cultivated specimen grown at the botanic garden
on the island and communicated by Rev. Lansdowne Guilding to
W. J. Hooker with whom he was in active correspondence.

The Triana collection “8,” February 1856, from Llano San Martin,
Villavicencio, Meta, Colombia, 400 m. (BM) cited by Triana and
Planchon ® as Vismia macrophylla, represents the extreme form of
this species. Triana’s No. “7” from the same locality is very puzzling,
combining as it does the flower characters of V. macrophylla with the
leaf shape and vestiture of V. baccifera subsp. dealbata. On the other
hand Fendler 8, from Chagres, at the mouth of the Chagres River,
now in the Panama Canal Zone, represents a less frequent inter-
mediate phase between this species and true Vismia latifolia. This
Fendler collection is the basis of Sprague’s doubt as to the correctness
of Triana and Planchon’s interpretation of V. macrophylla. But,
judging from a photograph, Mendler 8 closely approaches the isotype
collection (formerly at Berlin, but destroyed in World War IT) in
leaf shape, panicle characters, and the rather strong nervation. In
many characters these collections stand between Vismia angusta and
V. macrophylla and were it not for their fascicled stamens might be
taken for a variant of V. tomentosa.

Two Brazilian collections, Ducke 1068 (NY, US), and Corner 63
(NY), from the vicinity of Man&us, Amazonas, are unique in the

18 Ann, Sci. Nat. [V. Bot. 17: 306. 1862.

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 309

abundance and character of the pepperlike black dot glands on the
lower leaf-surfaces, which in these collections are so prominent as to
show clearly through to the upper surface. The leaves furthermore
are of an unusually thin texture for this species.

3.Vismia sandwithii Ewan sp. nov.

Arbor parva vel frutex 5 m. altus vel minus, ramis superioribus
firmis subquadrangularibus, dense rufo-tomentosis, rache atque ramis
conspicue subpannosis; laminis foliorum oblanceolatis vel obovatis,
basi cuneatis vel vix cordatis, apice apiculatis vel caudatis, subtus atro-
punctatis, tomentosis, venulis supra subglabris, nitentibus, 18-30 em.
longis, 7-10 cm. latis; petiolis crassis, 1-2 cm. longis, rufo-tomentosis;
paniculis compactis atque breviter ramosis; sepalis oblongis breviter
acutis, 5-6 mm. longis, viridi-rufidulis dense tomentosis, intus viri-
descentibus, atro-vittatis; petalis oblongo-spathulatis, breviter acutis,
ochroleucis, atro-vittatis, intus dense comosis; staminibus ad anthesin
conspicuis filamentis persistentibus; fructibus immaturis conico-
ovoideis, 1 cm. longis.

Small tree or shrub to 5 m. high, the upper branches stout some-
what quadrangular, densely, almost velvety rufous-tomentose with
long, branched hairs; upper leaves little if at all reduced below the
panicle and surpassing it, all the blades strongly bifacial, dark green,
subglabrous and shining above, light green below, sparsely rufous-
tomentose but densely so along the veins, the blades oblanceolate or
obovate, apiculate at tip or even caudate, acute at base or at most
weakly cordate, 18-30 cm. long, 7-10 cm. wide, the submarginal
veins well-defined; petioles stout, 1-2 cm. long; panicle compact,
shortly branching, 6-9 cm. long, the rachis and branches densely
rufous-tomentose ; sepals oblong, shortly acute, 5-6 mm. long, densely
rufous-tomentose becoming greenish in fruit, greenish within, black-
vittate; petals oblong-spatulate, acute, pale cream, densely comose on
face, black vittate on back; stamen column conspicuous at anthesis,
becoming indurated in fruit; immature fruit conic-ovoid, 1 cm. long,
‘Male green, brown-dotted.”’

Type in the Royal Botanic Garden Herbarium, Kew, collected at Mahdia
River, Potaro River, 107 miles on the Bartica-Potaro road, British Guiana,

8 January 1943, by an unrecorded collector for the Forest Department of British
Guiana (no. 3726); isotype in the New York Botanical Garden.

PARATYPES:

BRITISH GUIANA: Mahdia River, Potaro River, Jan. 21, 1943 (fruit), Forest
Dept. B.G. 3805 (K, NY); 111 miles Potaro River Road, May 10, 1952, Forest
Dept. B.G. 6480 (K,U).

Vismia sandwithii is clearly an extreme morphological development
of the V. macrophylla-angusta alliance. The bifacial leaves, their
shape, venation, and apiculate tips are suggestive of V. sessilifolia
and so this plant may prove to be a hybrid between that species

310 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

and V. macrophylla. Then too, the large floral leaves subtending
the compact and at times lateral panicles suggest V. lateriflora.
Although V. macrophylla and V. angusta represent “exceedingly
variable populations” (cf. Maguire, Bull. Torrey Club 75: 417,
418. 1948), I am not including V. sandwithit within them because
of its possible hybrid origin involving genetic relationships with
V. sessilifolia as well as with V. macrophylla, and because of the
extreme nature of its sepal, leaf, and pubescence differences. Vismia
sandwithit is a small local population of coastal British Guiana where
it has been described as a “very common” small understory tree
or shrub in second-growth forest. One collection, (Morest Dept.
B.G. 6480) mentions its growing on “ironstone gravel,”’ another
(3805), on “white sand,” and the type collection (3726), ‘‘of all
[types of] soils.”’

It is appropriate to associate this British Guiana Vismia with Noel
Yvri Sandwith, of the Royal Botanic Gardens, Kew, who has botanized
in British Guiana and who published on this genus in 1931, and who
in many ways has assisted me in the present revision.

4, Vismia angusta Miq. Linnaea 18: 27. 1844.

?Hypericum reticulatum Poir. in Lam. Encyl. Suppl. 3: 694. 1813. Type
from Cayenne, no collector designated, in Desfontaine Herb. at Paris.
Sagot reports the Poiret type as consisting of a single leaf with no record
of the country of origin. Sagot’s drawing and short memorandum of
the lost type, examined at Paris, are not sufficient to affect the rejection
of Poiret’s name as a nomen dubtum.

2V. reticulata Choisy, Prodr. Monog. Hyper. 34, 1821.

V. macrophylla sensu Benth. in Hook. London Journ. Bot. 2: 371. 1843,
as, for example, Schomburgk 405 (K); non H.B.K.

V. latifolia sensu Reich., in Mart. Fl. Bras. 12'; 208. t. 38. 1878, excluding
var.; non V, latifolia (Aubl.) Choisy.

?V. latifolia var. reticulata Reich. in Mart. Fl. Bras. 12!: 208, 1878, as to
basionym.

Caopia cordata Rusby, Bull. N. Y. Bot. Gard. 8: 105. 1912. Type: Apolo,
Bolivia, 1440 m., R. S. Williams 99 (NY; isotypes, BM, K, US).

V. cordata Blake, Contr. Gray Herb. 53: 41. 1918.

Type: “Surinami ad Osembo in Para, mfense] Aprilis 1842,” Focke 382 (U).

Eyma cited ‘584’ in error.

ADDITIONAL SPECIMENS EXAMINED:

FRENCH GUIANA: Cayenne, Martin (BM); Godebert, Wachenheim 107
(K, US).

SURINAM: Without locality, Hostmann & Kappler 162 (MO, NY, P, 8,
U); Cottica River, Lanjouw 398 (U); Watramiri, B.W. 4543 (NY); Sectie O,
B.W. 2711 (US), 2919 (NY); Zanderij I, B.W. 4862 (NY).

BRITISH GUIANA: Morabelli Creek, Sandwith 395 (K, NY, U); Mora
Landing, Moruka River, de la Cruz 1853 (UC, US); Mazaruni Station, Forest
Dept. B.G. 6469 (tree in open creek swamp 4 in. diam., 40 ft. high) (K); below
Kaieteur, Potaro River, Jenman 977 (K).

EWAN-——-SOUTH AMERICAN SPECIES OF VISMIA 311

VENEZUELA: Borfvar: Rio Karin, Alto Rfo Paragua, ‘uadama-yek,”’
Cardona 1229 (US); Mojasilla, south of La Paragua, 75 m., “sangrito,’’ L. Williams
12773 (US); Rfo Uairén, Sabanas de Santa Elena, Tamayo 2958 (US).

COLOMBIA: Meta: Between Villavicencio and Rio Ocoa, 450 m., Dugand
& Jaramillo 2917 (US); Villavicencio, Apiai, 500 m., Cuatrecasas 4772 (US);
Villavicencio, Schiefer (US). Purumayo: Quebrada del Rio Mulato, Mocoa,
Cuatrecasas 11305 (US). CaquerrtA: Solano, near Tres Esquinas, 200 m., ‘“‘lacre,’”’
Little & Little 9502 (NO, US). Amazonas-Vaupks: Soratama, Rfo Apaporis,
Schultes & Cabrera 12724 (NO, US). Tourma: Faldn, region of ‘“‘Calamonte,”’
1120 m., “puntelanza,” Garcia-Barriga 8377 (US). Antioquia: San Carlos,
Kalbreyer 1374 (C); near Yolombé, Lehmann 40039 (BM, K, US); [Rio] Samand4,
500-1300 m., Lehmann 7547 (US); Mulatos, 50 m., Haught 4885 (NO, US,).
?Cauca: El Chorro, 800 m., March 1853, Triana (COL), same loc. no. ‘5”’ (BM).
Eu VALLE: Quebrada de La Brea, Rio Calima, region of Choeéd, 20-40 m., Cua-
trecasas 21279 (US). CuHocé: Rio Atrato, vicinity of Quibdé6, 400 m., Araque-
Molina & Barkley 19Ch 134 (NO, US, a variable coll.)

PERU: Loreto: Iquitos, ca. 100 m., Sandeman 2273 (K), L. Williams 1385
(F), 1460 (F), 1533 (F), Tessmann 3614 (NY), 5040 (G), Killip & Smith 26923
(US), 27221 (US); Mishuyacu, near Iquitos, “pichirina blanca,’ 100 m., Klug
888 (US); Pebas, Rfo Amazonas, L. Williams 1734 (US), 1748 (US); Maquisapa,
Upper Rio Nanay, L. Williams 1205 (F); Caballo-Cocha, Rio Amazonas, L.
Williams 2194 (US); Gamitanacocha, Rio Mazdn, 100-125 m., Schunke 133 (F,
NA, NY, UG, US).

BRAZIL: Without locality, Sellow 159 (P.) AMazonas: Rio Embira, at Rio
Tarauaca, basin of Rio Jurua, Krukoff (G, K, 8, UC, US); San Jose de Rio Negro,
Riedel 1427 (US). Banta: Without locality, Salzmann (P); Ilheos, Riedel 187
(US), Blanchet 1933 (C, F, FI, G, K, NY, W).

This widely distributed Vismia, a tree 4 to 12 meters high or more,
is characteristic of selva borders or in Amazonian Brazil it grows on
higher ground back from the seasonally inundated lands. Vernacular
names, arising in most instances from the orange sap that bleeds
freely from the cut stems, include: ‘‘ Bloodwood” (British Guiana);
“sanerito” (Amazonian Venezuela, Amazonas); ‘‘puntelanza’”’ (Colom-
bia); “lacre’ (Amazonas). See Eyma”™ for Surinam vernacular
names.

N. Y. Sandwith drew attention (Kew Bull. 1931: 174) to the identity
of Vismia angusta as distinguished from V. macrophylla H. B. K.,
and determined the oldest name for this species, which long passed
as Vismia ferruginea, to be Miquel’s V. angusta. Eyma followed
Sandwith in this conclusion.

The cordate leaf-base is a characteristic feature of Vismia angusta
yet some Peruvian collections have a rounded leaf-base. Further-
more, these non-cordate specimens average to have distinctly smaller
leaves. In British Guiana V. macrophylla likewise has cordate-based
leaves.

1A wrong number. In the Lehmann fieldbook (US), no. 4003 is listed as Masdevallia, an orchid.
20 Pulle, Fl. Surinam 3: 75. 1934.

312

On Gorgona Island, off the Pacific Coast of Colombia (Collenette
588, K, US), Vismia angusta likewise exhibits smaller leaves which
are not prominently cordate as is so characteristic of the species over
its principal continental range. Perhaps this tree of Gorgona Island
and the Peruvian-Colombian transmontane forests merits description
as a subspecies of V. angusta.

Vismia angusta reaches its southern limit in Bolivia, where, as in
the interior of Peru and Colombia, the leaves average smaller as noted
above. Rusby’s Caopia cordata was a collection of Bolivian V.
angusta named for the cordate leaf-bases, which are conspicuous in
the type collection.

Though the name Vismia angusta has been adopted for this dis-
tinctive species there is considerable evidence that the oldest valid
name may prove to be V. reticulata (Poir.) Choisy. The basionym,
Hypericum reticulatum Poir., rests on a good description and the
source of the material, though not documented, might well have been
from typical angusta territory. Neither Sandwith nor Kyma suc-
ceeded in locating the type. In view of the complex nature of this
group of Vismias and the undoubted basis for the adoption of the
name V. angusta, it has seemed in the interest of stability to regard
Hypericum reticulatum as a nomen dubium.

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

TABLE 1—Comparison of the characters of Vismia angusta, macrophylla, and

Vismia macrophylla
Tree 2-5 (or 10) m. high
at flowering time.

Leaves often thin-
textured, ovate to
ovate-lanceolate,
acuminate, weakly
cordate.

Secondary veins promi-
nent or at times rather
faint, esp. toward tip
of blade.

Submarginal veins
indistinct.

Petioles 1.5 cm. long or
more.

Rachis of inflorescence
glabrate in age, with
only flocs of persistent
brown tomentum of
stellate hairs.

sandwithii
Vismia angusta

Tree 5-10 m. high or

more at flowering time.

Leaves generally
leathery, oblong,
shortly acute, dis-
tinctly cordate at the
base (except Gorgona
Island and interior
Colombia and Peru-
Bolivia).

Secondary veins gen-
erally all very promi-
nent, raised below.

Submarginal veins
indistinct.

Petioles usually 1 cm.
long or less.

Rachis of inflorescence
permanently rusty-
tomentose with stel-
late hairs or tardily
deciduous- to floccose-
tomentose.

Vismia sand withii
Shrub or small tree less
than 5 m. high (?), the
stem 5 cm. diam.
Leaves rather thin-
textured, oblanceolate
or obovate, cuneate,
apiculate to caudate,
acute or at most
weakly cordate.

Secondary veins rather
faint toward tip of
blade.

Submarginal veins promi-
nent, well defined.
Petioles 1-2 em. long.

Rachis of inflorescence
conspicuously densely
rufous-tomentose,
branching hairs almost
velvety.

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 313

TABLE 1.—Comparison of the characters of Vismia angusta, macrophylla, and
sandwithii— Continued

Vismia macrophylla Vismia angusta Vismia sandwithii

Flower buds dark fuscous Flower buds light yel- Flower buds rufous.
to almost blackish. lowish brown.

Flowers without odor Flowers fragrant. Flowers without odor (?).
(?).

Petals obovate, greenish, Petals linear-spatulate, Petals oblong-spatulate,
tomentose on inner whitish or cream- acute, whitish, densely
face but not so densely colored, so heavily comose on face, black-
as to obscure the comose as to obscure guttate on back.
blades. the blade, the hairs

often twice as long as
the width of the petal.

Fruit globose, green. Fruit globose to ovoid, Frutt conic-ovoid, pale

green or olive-brown. green,

Sepals 4-5 mm. long; Sepals 4-5 mm. long; Sepals 5-6 mm. long, the
veins usually obscured veins evident beneath tomentum so dense as
by dense tomentum. tomentum. to obscure the vena-

tion.

5. Vismia lateriflora Ducke, Arquiv. Serv. Florest. Rio Janeiro 1: 33. 1939.

LrectotyPE: Esperanga, at the mouth of Rio Javary, Amazonas, Brazil,
Ducke 25054 (isolectotype, K). Syntype: Presidente Marquez Station, on the
Madeira~Mamoré Railway, Mato Grosso, Brazil, Kuhlmann 21223 (duplicate
syntypes, K, US).

ADDITIONAL SPECIMENS EXAMINED:

COLOMBIA: Rio Loretoyacu, Trapecio amazénico, 100 m., Schultes &
Black 8448 (US), 8449 (US).

PERU: [probably Hudnuco but without loc.] Dombey (F). Cuzco: Cerro
San Pedro, 1600 m., Vargas 8504 (US). Lorero: Pebas, L. Williams 1793
(F); La Victoria, Rio Amazonas, L. Williams 2691 (F); Caballo Cocha, L. Wil-
liams 2192 (F), 2364 (F); Soledad, Rio Itaya, 110 m., Killip & Smith 29689
(F, US); Maynas, Poeppig 421 (P); Mishuyacu, Iquitos, 100 m., “pichirina,’’
Ktug 354 (F, US); Iquitos, 100 m., Kilizp & Smith 27085 (F, US); Yurimaguas,
Lower Rio Huallaga, 155-210 m.,.L. Williams 3821 (F), 4899 (F, US), 5003
(F, US), Killip & Smith 27550 (US); San Antonio, Rio Itaya, 110 m., Killip &
Smith 29475 (F, US). HuAnuco: Pampayacu, 1050 m., Macbride 5019 (F).

BRAZIL: Amazonas: Near Tres Casas, basin of Rio Madeira, Munic.
Humayta, Krukoff 6325 (A, G, 8, U, US, W); near Palmares, basin of Rio Soli-
moés, Munic. Sa0 Paulo de Olivenga, Krukoff 8328 (A, G, K, P, 8, U, US).

Vismia lateriflora may prove to be but a seasonal phase of V.
angusta with axillary inflorescences produced through suppression of
the usual terminal panicle, induced perhaps at times of exceptional
water supply or by some other microclimatic factor. Still, the oblong
leaves of V. angusta are dull above, whereas the ovate, acute leaves of
V. lateriflora are more or less shining above as if varnished. Vismia
angusta typically has distinctly cordate leaves, the midrib and second-
aries prominently raised beneath, and marginal or submarginal
areolate venation. Its spherical flower buds, opening tardily, are

314 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

yellowish-green or sulphur-colored on the outside, the petals heavily
yellowish comose, with the hairs often so long as to extend beyond the
petals, forming a kind of tuft in the center of the flower. V. later-
flora, on the other hand, has ovate leaves of a distinctly thinner tex-
ture, a truncate or at most subcordate base, and the nerves not prom-
inently raised beneath, areolate with loop veins well inside the leaf
margin. Moreover, the flower buds in Vismia lateriflora are long-
ovoid, early opening, brownish on the outside, the petals less densely
comose, the hairs not exceeding the petals. The pubescence of the
lower leaf surfaces of the two species is similar but in V. angusta it
has a dusky quality, due to the presence of minute black glands,
one gland to each areole of the lower leaf surface. In V. lateriflora
the thin-tomentulose lower leaf surface is distinctly reddish and,
though the minute dark glands are present in this species as well,
they are obscure. Though the lateral axillary panicles especially
characterize V. lateriflora, both lateral and terminal inflorescences
may occur (e.g¢., Macbride 5019 and Krukoff 8328), or, again, lateral
panicles may be wholly absent and only the terminal ones present.
In this connection, V. ramuliflora, which might from the name be
thought to be an earlier name for this species, is, in fact, a synonym
of V. sessilifolia, of the coast of northern South America.

6. Vismia rufa Cuatrecasas, Rev. Acad. Colomb. Cienc. 7: 47. fig. 1. 1946.

Tyre: Forest, Rio Cajambre, Department of Il Valle, Colombia, 5-80 m., 5-15
May 1944, José Cuatrecasas 17449 (Comision de Bot. del Valle, Cali, Colombia)
(isotypes, NY, US).

ADDITIONAL SPECIMENS EXAMINED:

COLOMBIA: Et Vatue: La Trojita, Rio Calima (region of Chocé), 5-50
m., Cuatrecasas 16614 (US); hills near Quebrada de la Brea, Rio Calima (region of
Choc6), 30-50 m., Cuatrecasas 21102 (US); near Buenaventura, April 8, 1882,
J. Ball (K).

Vismia rufa is a tree 16 to 30 meters tall, the trunk 25 to 40 centi-
meters in diameter, the bark dark reddish-brown, scaly and breaking
into narrow rough plates from longitudinal cracks; wood soft-textured,
red; latex bleeding bright orange-red; leaves thick, leathery, con-
spicuously petiolate, the petioles 12-25 mm. long, subterete or grooved
on the upper side, the blades ovate or ovate-elliptic, shortly acute,
11-18 em. long, 4.5-7 cm. wide, lustrous-shining and glabrous above,
the midrib impressed, strikingly rufous beneath with a heavy felt-like
tomentum; inflorescence much-branched, sometimes few-flowered and
lateral but generally with very numerous small crowded flowers in a
diffuse panicle; sepals early spreading, ovate-oblong, small, 3.5-4.5
mm. long, narrowly hyaline-margined, evenly ferruginous-tomentose
on the back; petals a little longer than the sepals, thick-textured,
spatulate-ovate, rounded at the tips, roseate, vittate with a few dark

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 315

lines; stamens reddish, very short, about half as long as the ovary;
fruit unknown.

This interesting endemic Vismia of the Pacific rain forests of Colom-
bia is most singular for its small flowers, a character not evidently
appreciated by Cuatrecasas as significant when the tree was described.
Its taxonomic position may be near V. brasiliensis, judging from the
five small stamen-columns, the rufous-tomentose leaves, and the ample
panicle of numerous crowded flowers, but the leaves are wholly unlike
that Brazilian species, recalling rather V. cavanallesiana.

7. Vismia micrantha Mart. ex St. Hilaire, Fl. Bras. Merid. 1: 327. 1827; Mart. in

Spix & Mart., Reise Bras. 2: 552. 1828.

Trianthera floribunda Pohl ex Reich. in Mart. Fl. Bras. 12 !: 198. 1878,

pro syn. V. micranthae. Authentic material from Villa Rica, Brazil,
E. Pohl 3738 (F, W).

Caopia micrantha Kuntze, Rev. Gen. Pl. 1: 59. 1891.

Tyre: Syntypes “in sylvis et campis provinciae Minas Geraés prope San
Gabriel, vicum vulgo Catas Altas, urbem Villa Rica, et in monte Serra Negra,”’
presumably referring to St. Hilaire collections, now at Paris. Presumably authen-
tic material for the Spix & Martius description: Without locality, Martius 970
(BM, G, K, L, MO, P, W, WU).

ADDITIONAL SPECIMENS EXAMINED:

BRAZIL: Minas Gerais: Serra de Ouro Preto, en route to Antonio Pereira,
Schenck 3577 (C); Ouro Preto, Riedel 2634 (K); Morro de Cruziero, Ouro Preto,
Damazio 1328 (G); Serro de Cipé, Munic. Santa Luzia, Barreto 1274 (F); ‘Minas
sylvis montosis,”’ Riedel 25 (P); Monte Itacoluni, Santa Luzia, Casaretto 3510 (G);
Monte Capanema, Riedel 1474 (S); Saé Juliad, Santa Luzia, Schwacke & Glaziou
(P); Ressaca, Bello Horizonte, Barreto 2910 (F); Serra do Caraga, Munic. Santa
Barbara, Barreto 2912 (F).

Tree; leaves rather thin, more or less green on both surfaces, the
blades short-ovate, 6-10 cm. long, 3-4 cm. wide, finely glandular-
punctate and often thinly hirtellous beneath, especially along the
veins, with mostly simple hairs; panicle often very compound, rather
compact, the flowers numerous; sepals oblong, rounded, thinnish, with
no or a barely distinct marginal flange, glabrate; petals about twice as
long as the sepals, cream-colored, obovate, tomentulose within, punc-
tate on the back with black glands; stamen column bearing 3 anthers;
ripe fruit globose to short-pyriform, black, 4 mm. long, usually tipped
with 5 persistent styles, the sepals spreading.

Vismia micrantha is easily distinguished from all other species by
the unique character of its stamens, but a certain affinity with V.
brasiliensis is suggested by characters of both the foliage and the
inflorescence. It will be noted that both species differ from the section
Euvismia in having a reduced number of stamens borne on a single
stamen column. These two Vismias are in fact not easily distin-
guished in sterile condition. However, the secondary veins in Vismia
micrantha in anastomosing at or near the margin unite by a weak con-

316 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

necting vein, this often freely branching. In V. brasiliensis, on the
other hand, the secondaries anastomose by a single strong connecting
vein, without branches to the margin. The shape, size, and pu-
bescence of the leaves of the two species are very similar, but on the
whole V. micrantha has thinner leaf-blades than those of V. brasi-
hensis. ‘The tomentulose sepals of V. brasiliensis often contrast easily
with the glabrate sepals of V. micrantha; the fruit of the former is
larger and generally the styles are earlier deciduous than those of
V. micrantha.

8. Vismia brasiliensis Choisy, Prodr. Monog. Hyper. 35. t. 2 1821.

V. longifolia St. Hil. Fl. Bras. Merid. 1: 326. t. 68. 1827. Typx: “In sylvis
caeduis prope tabernam vulgo Rancho de José Henriquez, haud longe
ab urbe Villa Rica, in provincia Minas Geraés; florebat Januario,” Brazil,
St. Hilaire. Not distinguished among the St. Hilaire colls. at Paris.
The illustration though generalized in details is conclusive evidence of its
taxonomic position here.

V. laccifera Mart. in Spix & Mart. Reise Bras. 2: 552. 1828. Type: A
Martius collection, presumably from the vicinity of Villa Rica, in the state
of Minas Gerais, Brazil. St. Hilaire evidently saw the manuscript de-
scription of Martius’ species prior to its publication the following year,
for he refers to it in 1827 (in his Fl. Bras. Merid. 1: 326. 1827). Rei-
chardt evidently believed lacczfera to be a typographical error for baccifera,
and ignored it as a synonym of brasiliensis; however, laccifera was sup-
ported by a full description, which shows it to be a synonym of brasiliensis.

V. brasiliensis var. lasiantha Reich. in Mart. Fl. Bras. 12!: 198.1878. Types:
Sellow 530, from Brazil (Delessert Herb. and Boissier Herb. G, GL, K, L,
W, fragments of isotype ex Geneva, F; Photo FM 9166, of specimen in
Berlin Herbarium).

V. lasiantha Klotzsch ex Reich. in Mart. Fl. Bras. 12 1: 198. 1878, pro syn.

V. brasiliensis var. longifolia Reich. in Mart. Fl. Bras. 121: 198. 1878.

Caopia brasiliensis Kuntze, Rev. Gen. Pl. 1:59. 1891.

Tyre: “In Brasilia’, according to the description but not so labeled, simply
“Mr. Steven, 1820’ (in de Candolle Herb., G). The name ticket is in the hand
of Choisy. The plate, drawn directly from the specimen, is too generalized in
detail, and fails to show the thin pubescence on the lower leaf-surfaces, the
lineate-guttate sepals, and the fine hairs of the pedicels.

ADDITIONAL SPECIMENS EXAMINED:

BRAZIL: SAo PauLo: Penha, in 1839, Guillemin 498 (G), Jan. 1840, Guille-
min 498 (TCD); Jardim Botanico, SAo Paulo, Hoehne 198 (F,S; this no. at A
and G is V. micrantha). Minas Gurats: Jardim Botanico, Bello Horizonte,
Barreto 2909 (F); Serra do Taquaril, Barreto 2908 (F); Matta de Empreza, Barreto
4035 (I); Lagoa Santa, Warming (C); Caa do Campo, Claussen 3 (F), Claussen
4(FI, G, W); Caldas, Regnell III 298 (FI, 8, US, WU); Gongo-soco, Jan. 1825,
Riedel (NY, US); ‘on border of the forest Gongo-Soco,” in 1834, Bunbury (CGE);
without locality, Claussen 134 (BM, K).

Tree; leaves rather thin, generally green on both surfaces, the blades
ovate or short-ovate, 6-10 cm. long, 3-7 cm. wide, finely glandular-

punctate and generally simply hirtellous beneath, especially pubescent
along the veins with both simple and branched hairs; panicle rather

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 317

compactly branching, the flowers numerous, the whole inflorescence
cano-tomentulose; sepals oblong-ovate, bordered by a distinct flange-
like membranous margin, tomentose over the back with a close felt-
like tomentum; petals about 1.5 times as long as the sepals, obovate,
cream-colored, densely tomentulose within, punctate with black
glands on the back; stamen column bearing 5 anthers; ripe fruit
pyriform or globose, acute, dull black, 4-9 mm. long, the styles
persistent.

Vismia brasiliensis comprises two distinguishable geographic races,
a more northern and southern Brazilian race, and each has been named
in the past and maintained as varieties by Reichardt. The Minas
Gerais race, which tends toward longer leaves and only moderately
tomentose flowers, is evidently St. Hilaire’s V. longifolia. The
Sao Paulo race, with short-ovate leaves, recalling those of the garden
apricot in size and texture, and with truly lanuginous flowers, is
what Klotzsch designated in the herbarium as V. lasiantha. Choisy’s
type evidently represents this southern phase, but if so the pubescence
is somewhat deciduous from the fruiting branchlets. Both forms
are inconstant and many more collections from the vast regions of
Brazil are needed before a clear conclusion may be reached.

Vismia brasiliensis has been compared with V. micrantha under the
latter species. The leaves of V. brasiliensis generally have a slightly
developed cordate base, and the blades tend to be plicate, a condition
not seen in V. micrantha.

St. Hilaire listed three Vismias for central Brazil,”! namely, V.
longifolia, V. guianensis, and V. micrantha. When he described V.
longifolia he overlooked Choisy’s recent publication of V. brasiliensis
(1821) or its inclusion in the account of the genus in de Candolle
(1824). V. longifolia was described by St. Hilaire as having ‘“‘stamini-
bus 25-30.” It is singular that if he was describing a phase of V.
brasiliensis he did not refer to the distinctive character of that species,
namely, that each stamen column bears only five stamens. However,
Reichardt aligned V. longifolia with V. brasiliensis, and certainly the
very detailed original description of V. longifolia is not out of harmony
with V. brasiliensis.

It is possible that the “Ubapitanga” of Marcgrav (Hist. Nat.
Brasil. 293) and his ‘“Caopia arbore ejusque facultatibus” (op. cit. 60),
accompanied by a crude line drawing of a thick-leaved plant with
variable-shaped leaves and globose fruits, may represent Vismia
brasiliensis. If so, it constitutes the earliest mention of this Vismia,
or, indeed, of any Brazilian Vismza in literature.

We may be much more certain of the fact that the Vismia described
by Vandelli when he established the genus (1788) was V. brasiliensis,

1 Fl. Bras. Merid. 1:325-328. 1827.

318 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

for he characterized it as having “filamenta plura in 5 phalanges
s[ive] corpora connata.” V. brasiliensis was certainly therefore the
original species of the genus, although it was not so chosen when
Vismia was conserved. Vandelli indicates the origin of his Vismia
by the initial “B,’”’ meaning Brasilia.

9. Vismia parviflora Schlecht. & Cham. Linnaea 3: 119. 1828. PLaTE 3
Caopia parviflora Kuntze, Rev. Gen. Pl. 1: 59. 1891.

Type: ‘Brasilia tropica,’’ Sellow (B, Photo FM 9168, which carries a ticket
reading ‘‘Sello 1365’’) (Isotype CGE).

“VY. Calycibus pellucidis 3-nervibus subpunctatis, petalis punctatis, phalangibus
triandris calyce brevioribs, stylis longis, foliis ellipticis nigro-punctatis, pilis
sparsis.

“In Brasilia tropica collegit Sellow.

“Folia elliptica v. elliptico-lanceolata, acuminata, saepius in ovatum rarius in
obovatam vergentia formam, basis subinde inaequali acuta vel rarius obtusa
rotundata, acumine brevi subtrilineari saepe obliquo, submembranacea, quam in
congeneribus paullo tenuiora, subtus punctulata, punctis opacis nigris prominen-
tibus, obsitaque pilis stellatis rufescentibus sparsis; maxima 4 poll. longa. 114 lata,
petiolo semipollicari. Rami novelli cum petiolis, panicula alabastrisque tomento
tenui rufo-canescente oblitterante obducti. Panicula multiflora, laxiflora, quam
in congeneribus uberior, pyramidata, e ramorum primariorum paribus circiter
sex constans, quam panicula Syringae persicae paulo minor.—Flores brevissime
pedicellati, parvi, expansi 2 lin. circiter metientes. Laciniae calycinae ellipticae,
obtusae, lineam circiter longae, subpellucidae, 3-5 nerves, evittatae, nune punctis
resinosis paucis (rarius 6), in aliis aliter dispositis, notatae, nune omino impunc-
tatae. Petala elliptica, obtusa, calyce longiora, intus villis brevibus parce obsita,
punctis resinosis irregulariter adspersa. Staminum phalanges calyce breviores,
omnino glabrae, triandrae, filamento medio longiori supra tertia parte libero.
Squamae villosae, subspathulatae, phalangibus paulo breviores. Germen glabrum.
Styli 5 longiusculi, filiformes, e calyce demum exserti, stigmate depresso-capitato.
Fructus—?”’

Vismia parviflora is a rare species evidently closely related to V.
micrantha but that species has vittate sepals. Prior to my visit to the
Paris Herbarium I had not seen any collection of this Vismia and rested
my disposition of what is certainly a distinctive species on the original
description, reproduced here, and the Macbride photograph of the
type, Sellow’s collection in the Berlin Herbarium having been de-
stroyed. However, one collection of Vismia parviflora came to light
in Paris: ‘“Minas Gerais, 1816-1821,” St. Hilaire D55 (P). Dwyer ”
identifies this St. Hilaire collection as one taken in 1821-22 when he
visited the provinces of Rio de Janeiro (nos. 1-30), Minas Gerais
(31-587) and Sado Paulo (588-818). There is a second collection,
Minas Gerais, 1841, Claussen 5 (P), which is referred to this species
with reservations. It is notable that no recent collections of Vismia
parviflora, evidently of extreme local occurrence in Brazil, have been
made.

22 Ann. Missouri Bot. Gard. 42:162. 1955.

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 319

10. Vismia minutiflora Ewan, sp. nov.

Arbor parva vel frutex 2.5 m. altus, ramis superioribus gracilibus,
fissuratis, griseo-brunneis, raches atque ramis rufo-tomentulosis, in-
ferioribus glabrescentibus; foliis superioribus non reductis, laminis
omnibus submembranaceis, ovato-lanceolatis, 7-9 (13) cm. longis,
acuminatis vel cuspidatis, ad basim breviter acutis vel subrotundatis,
amphiglabris, infra atro-punctatis, supra venis impressis; petiolis
brevibus, 2-4 mm. longis; paniculis compactis et brevi-ramosis; sepalis
minimis, 1.5-2 mm. longis, flavibus, partibus centralibus atro-pubes-
centibus; petalis breviter oblongis, albis, atro-vittatis, intus sparse
comosis; staminibus ad anthesin exsertis; fructibus ignotis.

Small tree or shrub 2.5 m. high or more, the trunk slender, the
branchlets clothed with thin gray-brown bark splitting into narrow
shreds, rufous-tomentulose with fine branched hairs on young wood
and rachis of the panicle, glabrescent below; leaves little if at all re-
duced at the inflorescence and exceeding it, thin, amphiglabrous except
at the extreme base beneath where weakly stellate-pubescent, finely
black gland-dotted beneath and impressed-venose above, ovate-lan-
ceolate, 7-9 (or 13) cm. long, acuminate to cuspidate, shortly acute or
barely rounded at the base, the petioles slender, short, 2-4 mm. long;
flowers short-pedicellate, the panicle compactly and shortly branched ;
calyx very small, 1.5-2 mm. long, yellowish with a narrow central
black pubescent ridge, spherical in bud; petals short-oblong, white,
vittate with impressed black glands, rather lightly comose within;
stamens exceeding the corolla at anthesis; fruit unknown.

Type in the U.S. National Herbarium, No. 1,795,145, collected at San José del
Guaviare, Rio Guaviare, Intendencia of Meta, Colombia, in savana, alt. 240

meters, Nov. 11, 1939, by José Cuatrecasas (No. 7658); isotypes in the herbaria
of the Royal Botanic Garden, Kew, and the New York Botanical Garden.

PARATYPES:
PERU: Rio Azara, Upper Rfo Marcapata, east of Urcos, 1050 m., Sandeman
3730 (K); Moro, 1200-1500 m., Jan. 1866, Pearce (KX); ‘‘Perou,’”’ Gay 939 (P).

The affinities of Vismia minutiflora are with the section Vismia
and not with the section Trianthera, to which Vismia micrantha be-
longs, as might well be expected upon the basis of the small flowers,
although the combination of characters shown in Vismia minutiflora
are not closely approached by other species of the section Vismia.
The leaves recall Vismia cayennensis but are thinner and more herba-
ceous. The short-pedunculate panicle suggests a species of Sambucus,
an impression further borne out by the small flowers crowded into
small cymules. Individually, the flowers are rather showy for their
white petals which far exceed the exceptionally small calyx.

A Peruvian collection, Klug 2344 (A, US), from the mouth of the Rio
Zubineta, near Florida, Rio Putumayo, Department of Loreto, has

320 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

much larger leaves, 12-14 cm. long, and a more compound diffuse
panicle—quite unlike the Colombian type collection from the Rio
Guaviare. In points of flower structure this Peruvian collection is in
agreement, but the curious callosities on the sepals and the unusual
twisting of the long petals suggests a diseased plant. Aside from the
short-petiolate leaves, Klug 2344 resembles the type collection of
Vismia parviflora in several particulars.

11. Vismia sessilifolia (Aubl.) DC. Prodr. 1: 542. 1824. Puatss 4, 5

Hypericum sessilifolium Aub]. Pl. Guian. 2: 787. t. 312, fig. 2. 1775.

H. rufescens Lam. Encycel. 4: 150. 1796. Type: Aroura, a village opposite
the Isle of Cayenne (cf. Bellin’s map of 1763), French Guiana, 17-21 Nov.,
1785, Stoupy (P, Photo FM 35239, Photo Killip 911).

V. rufescens Pers. Syn. Pl. 2: 86. 1806.

V. rufescens var. sessilifolia Pers. loc. cit.

V. ramuliflora Miq. Stirp. Surinam 88. 1850. Type: “Sylvis umbrosis
regionum interiorum,” Surinam River, Surinam, April 1847, Hostmann &
Kappler 1823 (holotype, U; isotypes, G, 8; Photo FM 35238, of specimen
in Paris Herbarium),

Caopia sessilifolia Kuntze, Rev. Gen. Pl. 1: 59. 1891.

Caopia sessilifolia var. rufescens Kuntze, loc. cit.

Typu: Cayenne, Aublet (holotype, BM, Photo NY; isotype, G-DC). Gleason
64 (US) is a close match for Aublet’s figure.
ADDITIONAL SPECIMENS EXAMINED:

FRENCH GUIANA: Maroni, Wachenheim 58 (E); Karouany, Sagot 65
(BM, 8, W), 69 (P); Godebert, Wachenheim 49 (P); Roura, 1859, Sagot (P).

SURINAM: Patrick Savannah, B. W. 672 (NY, U); Sectie O, Krukoff 12315
(NY); Grote Zwiebelawamp, Lanjouw & Lindeman 1255 (U).

BRITISH GUIANA: Vicinity of Kartabo Station, junction of Mazaruni
and Cuyuni Rivers, Graham 292 (US); Arawak Matope, Cuyuni River, Tuéin 374
(BM, U, US); Tumatumari, Potaro River, Gleason 64 (NY, US); Potaro, 10 miles
south of Potaro Landing, Hitchcock 17389 (US); Waini River, de la Cruz 3730 (UC,
US); Kurupung, Tacoba, Lang & Persaud 286 (F); Barima River, Jenman 7017
(NY); “Roraima,” Schomburgk 917 (BM, CGE, FI, G), certainly from lowlands,
possibly from Venezuelan side of boundary, where essentially all of Schomburgk’s
collections originate.

VENEZUELA: Angostura, Humboldt & Bonpland 1070 (H. & B. Herb., P).

COLOMBIA: Et Cuocé: Istmina, Killip 35453 (US). Cauca: Cérdoba,
Dagua Valley, 30-100 m., Pitter 511 (US). Ex Vauie: Buenaventura, Killip
11734 (US); Road to Buenaventura, 400 m., Triana 6 (COL); Rfo Dagua, 10
km. from Buenaventura, Core 1519 (NO).

BRAZIL: Par: Faro, Campos do Tigre, Ducke 12494 (K).

In Vismia sessilifolia the lower surface of the leaves has a close
fulvous or rufous pubescence with usually a single, raised, dot-like
gland in a single areole; the upper surface is not impressed-reticulate,
as are both V. baceifera and lauriformis. The leaves in this species
are oblong-ovate, shortly acuminate, the petioles short, noticeably
broad, often wing-margined, the blades rather bifacial, glabrous and
shining, green above, reddish-puberulent beneath with a thin deciduous
scattering of fine hairs. The sepals are narrowly ovate, acute,

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 321

tomentulose on the back, about half as long as the narrowly oblong
petals.

Though most often with petioles about 1 cm. long, Vismia sessilifolia
occasionally has subsessile leaves, and it was this form that Aublet
described and illustrated as Hypericum sessilifolium. Gleason
describes his sessile-leaved collection (no. 64) as from a “shrub 15
feet high.’ The types of the two forms, the petiolate rufescens (PI. 3)
and the subsessile sessilifolia (Pl. 4), came from the same district
of Cayenne and I cannot separate the two on any geographic grounds.
Persoon aligned the sessile-leaved plant as a variety of Vismia
rufescens, and de Candolle admitted both as full species and even
placed them some distance apart in his treatment of the genus in the
Prodromus, but it does not seem necessary to give them taxonomic
recognition.

Vismia sessilifolia develops lateral axillary racemes either along
with, or in the absence of, the usual terminal inflorescence, and it is
this latter condition that led Miquel to distinguish V. ramuliflora.
Miquel believed its relationship to be with V. cayennensis, overlooking
V. sessilifolia and Persoon’s V. rufescens. Nearly all the specimens
cited that are in flower bear at least some reduced axillary racemes.
I cannot determine any morphological basis for Miquel’s proposed
species. Incidentally, the Amazonian V. lateriflora, which might be
presumed to belong here, is rather a relative of Vismza angusta.

Vismia sessilifolia is one of the most clearly marked species of the
genus. However, Eyma did not admit it in his account of the flora
of Surinam, an omission the more singular in that Eyma particularly
concerned himself with the typification of species of Guttiferae of
Surinam. ‘Triana’s collection from the vicinity of Buenaventura, 400
meters, agrees well with Choisy’s plate * and with the photograph
of Stoupy’s type at Paris of Hypericum rufescens Lam. Neither
Triana and Planchon, who remark on this Buenaventura collection,
nor Sandwith mention having examined the Stoupy specimen at
Paris.

12. Vismia japurensis Reich. in Mart. Fl. Bras. 121: 209. 4. 39. 1878.
Caopia japurensis Kuntze, Rev. Gen. Pl. 1: 59. 1891.

Tyre: “Prov. do Alto Amazonas in silva ad ripas fluminis Japura,” Martius.
SPECIMENS EXAMINED:
SURINAM: Zanderij I, Maguire & Stahel 25053 (XK, NY, US).
BRITISH GUIANA: Pomeroon River, Pomeroon District, de la Cruz 3149
(F, UC, US), 3170 (F, UC, US); Adaro River, Mazaruni drainage, near Wupaima
Mountain, Pinkus 166 (NA, US); Bootooba, Demerara River, Persaud 16 (F);
Canister Falls, Demerara River, Beccari 16 (FI, K); ‘‘Massaroonie & Cuyounie,”
Appun 277 (K).

23 Prodr. Monog. Hyper. ¢. 7. 1821.

322 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

VENEZUELA: Botfvar: Cerro Sarisarinama, Alto Caurd River, 800 m.,
Cardona 386 (US). Amazonas: Maroa, Rio Guainfa, 127 m., L. Williams
14348 (US); San Carlos, Rio Negro, 100 m., Holt & Gehriger 297 (US), 302 (US),
L. Williams 14478 (US), Holt & Blake 633 (US); Sabana de Rio Sanariapo, 120
m., L. Williams 16003 (US), 13064 (US); Yavita, 128 m., L. Williams 13912
p.p. (US).

COLOMBIA: Meta: Puerto Lépez, 240 m., “papa de lacre,’’ Little &
Little 8252 (NO, US). Vaupfs: Vicinity of Raudal de Yurupari, Gutierrez &
Schultes 931 (MEDEL, US). Caaquerd: Sucre, 1000-1300 m., Cuatrecasas
9047 (US). Amazonas-Vaupfs: Raudal Yayacopi, Rio Apaporis, Schultes &
Cabrera 15404 (NO, US).

BRAZIL: Without locality, Burchell 9697 (K, L), 10042 (K). Amazonas:
Tapurucuara, Rio Negro, Schultes & Lépez 8954 (US); Sio Gabriel, Rio Negro,
Holt & Blake 599 (US); Yucabi, Tate 966 (NY); Mandus, Rio Negro, Ule 5964
(G, L). Pard: Belém, Archer 8053 (US), 8243 (US).

Vismia japurensis is at once recognizable for its deltoid, acute
sepals which are noticeably thick, so that the fruiting calyx is almost
woody. The ovate leaves appear nearly glabrous beneath, but are
actually puberulent with microscopic golden glands. The leaf-base
is nearly shallowly cordate at times, but, unlike Vismia macrophylla
with which it might be confused, the peduncle and rachis of the panicle
are merely puberulent, not at all conspicuously tomentose. The fruits
are few, globose, and more pulpy than those of Vismia macrophylla.
13. Vismia latifolia (Aubl.) Choisy, Prodr. Monog. Hyper. 36. 1821. PLare 6

Hypericum latifolium Aubl. Pl. Guian. 2: 787 and 4: t. 312, fig. 1. 1775.

Caopia latifolia Kuntze, Rev. Gen. Pl. 1:58. 1891,

Typr: Cayenne (BM, photo, NY).

ADDITIONAL SPECIMENS EXAMINED:

SURINAM: Brownsberg, B. W. 1676 (L, US), 2372 (K,NY), 2869 (K,NY);
Zanderij I, B. W. 459 (S), 1404 (IK, NY), 1564 (IK, L, NY); Paramaribo, Split-
gerber 184 (L); Forest of Zandery, Samuels 275 p.p. (L, K).

FRENCH GUIANA: Rowea, Mar. 1859, Sagot (KK); Cayenne, Martin 27
(BM); Maroni, in 1864, Melinon (NY).

Small tree about seven meters high. Apparently a well-known
forest tree in Surinam, judging from the number of vernacular names
mentioned by Eyma in Pulle’s Flora of Surinam, but few collections
seem to have reached herbaria.

Sandwith drew attention (Kew Bull. 1931: 174) to the long persist-
ing confusion of this Vismia with V. angusta. The few collections
seen, however, do not uniformly conform to Aublet’s description of
the leaves as ‘“‘superne viridibus, inferne tomentosis, rufescentibus.”’
The inflorescence is congested and the flowers are small but in the
collections seen the sepals and fruits are blistered with a plant disease
which may have affected their development. A noteworthy character
not stressed in the literature is the narrowly oblong, abruptly acutish
sepals, which form an oblong-ellipsoid flower bud very truncate at the
base.

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 323

14. Vismia panamensis Duchass. & Walp. Linnaea 23: 748. 1850.
Caopia panamensis Kuntze, Rev. Gen. Pl. 1: 59. 1891.

Typs: “Panama,” Duchassaing, probably from the vicinity of Panama City.
SPECIMENS EXAMINED:
CANAL ZONE: Ancén Hill, Gillespie P-12 (DS); woods at Gatin, Hayes
921 (NY).
COLOMBIA: Eu Cuocé: Small quebrada opposite Palestina, Rio San Juan,
0-5 m., Cuatrecasas 21389 (US). En. VauLe: Rio Cajambre, 5-80 m., Cuatrecasas
17120 (US), 17551 (US); Puerto Merizalde, Rfo Naya, 5-20 m., Cuatrecasas 14081
(US); La Trojita, Rio Calima, 5-50 m., Cuatrecasas 16354 (US); La Cuarantena,
Buenaventura Bay, Killip & Cuatrecasas 38976 (US); Quebrada de Aguadulce,
Bahia de Buenaventura, 0-10 m., Cwatrecasas 19738 (US); between Isla de
Golondro and La Amargura, Rio Yurumangui, 10-40 m., Cuatrecasas 16068
(NO, US).
ECUADOR: Esmeraupas: Mangrove swamp, San Lorenzo, near Rfo
Nadadero, “sangre de gallina,” Marrero & Little 6274 (NY, US).

Vismia panamensis is here interpreted as a lowland rain-forest tree
of the Pacific Coast from Panama to Ecuador, having much the
aspect of V. lauriformis on the one hand and of V. baccifera subsp.
ferruginea on the other, and is probably most closely related to the
former. The leaves of V. panamensis are more prominently venose
than those of V. lauriformis, and these together with the leaf axils,
rachises, etc., are much more rufous-tomentellous. V. pana-
mensis is altogether a more heavily pubescent plant than V. lauri-
formis, from the stellate-tomentulose pedicels to the rich rufous-
tomentulose sepals that remain erect in flower. V. panamensis is
distinguished from V. baccifera subsp. ferruginea by the more heavily
tomentose leaves which remain tomentose when the plant is in flower,
and by the petals being only a little longer than the sepals, these
twice as long in V. baccifera subsp. ferruginea.

Triana and Planchon examined a Duchassaing collection at Mont-
pellier, the type of Vismia panamensis, and referred a Triana collec-
tion from Susumuco, Colombia, to that species. I have not seen a
specimen or photograph of a Duchassaing collection, but I have studied
a Triana sheet from Susumuco (now in Meta, near the Cundinamarca
boundary) preserved at Bogoté (COL), which probably is a duplicate
of the collection referred to by Triana and Planchon. Susumuco is
out of the expected range of V. panamensis, but the Triana collection
is a close match for a Panama specimen made by N. J. Andersson,
April 1852 (US). I am suggesting that Triana’s ‘“Susumuco”’ col-
lection of V. panamensis was taken, not in the interior of Colombia
but in the Chocé, where we know he botanized,™ and that a mixture
of labels ensued.

% Cf. A. Dugand, Rev. Acad. Colomb. Cienc. 5: 488, 1944.

598830—62——-3

324 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

15. Vismia urceolata Ewan, sp. nov.

Arbor parva 5 m. alta, ramis gracilibus, apicibus pubcrulentis,
deorsum glabris; laminis foliorum ovatis vel lanceolato-ovatis,
acuminatis, 10-11 cm. longis, 4-5 cm. latis, non crassis, supra obscuris,
infra minute cinereis vel brunneo-puberulentis praecipue secus nervos
tenuiter (vel non) glandulosis, omnibus petiolatis, petiolis saepe reflexis,
prominentibus, ca. 2 cm. longis; paniculis parvis, paucifloris, 4-5 cm.
longis; calycis valde urceolatis vel deorsum gibbosis, 7-8 mm. latis,
apice abrupte acutis, 9-10 mm. longis, deorsum aliquantum glabris, lobis
persistente clausis, tantum erectis ad anthesin, ferrugineo-tomentulosis,
marginibus hyalinis latis, minute atrosanguineo-vittato-glandulosis;
petalis auguste ovatis fulvis, lineatis, intus comosis ad apicem;
staminibus inclusis.

Small tree 5 m. high, with gray smooth bark, the branchlets puber-
ulent above, glabrate below; leaf-blades ovate or lance-ovate, 10-11
em. long, 4-5 cm. wide, rather thin-textured, dull above, finely
cinereous or brownish puberulent beneath, especially along the nerves,
only microscopically glandular-punctulate (if at all), the secondary
veins faintly anastomosing to form a submarginal vein, the petioles
slender, rather prominent, often reflexed, about 2 em. long; flowers
few, loosely clustered in small panicles 4-5 cm. long; calyces strongly
urceolate, enlarged below, 7-8 mm. wide, abruptly tapering to an
acute summit, 9-10 mm. long, glabrate below, the lobes tardily
spreading, and merely erect at anthesis, ferruginous-tomentulose on
the back, the broad hyaline margin prominent, very finely black
vittate-glandular; petals only a little exceeding the calyx, narrowly
ovate, dark brown, finely lineate, comose on the inner face near the
tips; stamens included; fruit unknown.

Type in the U.S. National Herbarium, No. 2,166,544, collected in cutover
flood-plain forest of Rio Meta, Puerto Lépez, Intendencia of Meta, Colombia,
alt. 240 meters, July 28, 1944, by Elbert L. Little, Jr., and Ruby Rice Little
(no. 8275); isotype in the herbarium of Tulane University.

Vismia urceolata, known only from the type collection, is perhaps
restricted to the Nanos of Colombia. The urceolate calyx distin-
guishes this Vismia at once from all other species. From the leaf
characters, habit, and panicle characters it is evidently most closely
related to V. lauriformis of cismontane Colombia.

The collectors record the vernacular name “‘lacre’’ for this species.
I am grateful to Dr. Little for the opportunity to study this and his
other Vismia collections.

16. Vismia falcata Rusby, Descr. S. Amer. Pl. 59. 1920.
V. angustifolia Rusby, Deser. S. Amer. Pl. 59. 1920. Typn: Sacupana,

Venezuela, April 1896, Rusby & Squires 141 (holotype, NY; isotypes, A,
BM, E, G, K, MO, US, W, WU).

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 325

Tyre: Santa Catalina, Lower Orinoco River, Venezuela, May 1896, Rusby &
Squires 142 (holotype, NY; isotypes, BM, K).

ADDITIONAL SPECIMENS EXAMINED:

TRINIDAD: Cumuto, Broadway 5726 (UC); Caroni, Eggers 1420 (US),
Britton & Mendelson 837 (US); Arima, Eggers 1118 (E, UC, US); (?) Mount
Tucouche, Broadway 5291 p.p. (A, UC, young shoots); (?) Blanchisseuse Road,
Broadway 5661 (UC); “Fl. Trinitatis,’’ Sieber 100 (E, G).

TOBAGO: Bacolet, Broadway 4139 (S, US, W); without locality, May 6,
1925, R. O. Williams (NY).

BRITISH GUIANA: Kamakusa, Upper Mazaruni River, de la Cruz 2819
p.p. (UC, US, somewhat atypical); Rockstone, Gleason 481 (US); Barima River,
Jenman 7017 p.p. (NY); Demerara River, Jenman 5029 (NY); without locality,
Schomburgk 182 (GL, US).

VENEZUELA: Botfvar: Roraima, Schomburgk 935 p.p. (CGE); Guayapo,
Bajo Caur4 River, 100 m., “‘sangrito rastrojero,” L. Williams 11754 p.p. (UC,
US); La Paragua, 285 m., Killip 37579 (US).

BRAZIL: Amazonas: (?) San Carlos, Rio Negro, Spruee 3115 (MO, P).

Vismia falcata is marked by its usually faleate peach-like leaves,
borne on slender petioles. The thin, crisp blades measure 3 to 4
times as long as broad, tapering to a slender acuminate tip. The
leaves are finely puberulent, punctulate beneath with black pepper-
dot glands.

Vismia falcata is certainly closely related to V. lauriformis. Some
collections from the State of Bolivar, Venezuela, suggest morphological
points of kinship with V. lauriformis in having broader ovate leaves
and a more diffuse panicle than usual.

Two of the Trinidad collections, Broadway 5291 and 5661, suggest
another and possibly unrecognized Vismia. These specimens show
smaller leaves, long-acuminate at the tips, the surface more shining
above and more suggestive of V. guianensis. It is noteworthy that
Broadway 5291 comes from the summit of Mt. Tucouche, a locality
recognized for its peculiar flora, as yet only partially known.

The Brazilian collection, Spruce 3115, is variable; a sheet in the
Meisner Herbarium (NY) is very much like typical V. falcata, but
another (NY) is broader leaved, like V. guianensis.

17. Vismia cayennensis (Jacq.) Pers. Syn. Pl. 2: 86. 1807.

Hypericum cayennense Jacq. Enum. Pl. Carib. 28. 1760.

H. acuminatum Lam. Encycl. 4: 150. 1796. Type: Herb. Lamarck from
“Aroura, [a precinct of] Cayenne,’ French Guiana, Stoupy (P, Photo
FM 35234, Photo Killip 712). Type labeled “ex roura in 17 au 21 novembre
1785.”

Vismia acuminata Pers. Syn. Pl. 2: 86. 1806.

H. eugeniaefolium Willd. ex. Spreng. Syst. Veg. 3: 351. 1826, pro syn.
Based on Humboldt coll. (no. 1038) from Curichana, Orinoco, Venezuela,
in Willdenow Herb. (Photo FM 9498). Authentic specimen in Humboldt
& Bonpland Herb. (P).

V. latifolia var. acuminata Sagot, Ann. Sci. Nat. VI, Bot. 11: 163. 1881.

Caopia acuminata Kuntze, Rev. Gen. Pl. 1: 59. 1891.

C. cayennensis Kuntze, op. cit. 58.

326 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Vismia floribunda Sprague, Trans. Bot. Soc. (Edinburgh) 22: 428. 1905.
Type: Mocoa, Caqueté [now Com. Putumayo], Colombia, Sprague 401
(holotype, K; isotypes, BM, US).

Type: Not designated by Jacquin, but undoubtedly from Cayenne, French
Guiana (cf. Jacq. Stirp. Amer. 213. 1763).

ADDITIONAL SPECIMENS EXAMINED:

TRINIDAD: Arima, Eggers 1363 (US); Toco Road, Valencia, Britton,
Hazen & Mendelson 1778 (NY, US); Fort St. Georges Hill, Johnston 106 (US);
Ortoire River, Guayaguayare Road, Britton, Freeman & Nowell 2542 (US);
Caroni River, south of Dabadie, Britton & Hazen 724 (US); Caroni North Bank
Road, Britton & Mendelson 824 (US).

TOBAGO: Big River Bridge, Mason Hall, Broadway 3035 (L, NY); Green-
hill, Broadway 4140 (S, US); Cocoawattee, 15 Jan. 1913, Broadway (US); Bacolet
River, Eggers 5731 (S, US).

FRENCH GUIANA: Karouany, Sagot 66 (G, K, NY, 8, W); Cayenne, 1792,
Leblond (A, NY); Cayenne, 1838, Leprieur (K, NY); Godebert, Wachenheim 40
(K, US); vicinity of Cayenne, Broadway 399 (NY, US), 209 (NY).

SURINAM: Raleigh Falls, Coppename River, Lanjouw 972 (NY); Kabel-
station, Lanjouw 1180 (NY); Tamarinal, Cottica River, Linder 105 (NY);
Charlesburg, near Paramaribo, Lanjouw 96 (U); near Oude Ryweg, Paramaribo,
Samuels 212 (A), 213 (S); Juden-Savanne, Kappler 1722 (G, 8); Charlesburg Rift,
3 km. north of Paramaribo, Maguire & Stahel 22779 (NY, US); Tawa Creek,
Maguire 23765 (NY, US); forest of the Station, Groningen, Samuels 112 (A, K);
Scotelweg, “swinani,’’ “pina,” Archer 2660 (NA, 8, US), 2706 (NA, US); without
locality, Hostmann 438 (BM, CGE, DS, G, K, OXF, 8); Sectie O, Maguire &
Stahel 23624 (US), B.W. 635 (US), 1772 (US)

BRITISH GUIANA: Without locality, Schomburgk 607 (CGE, G, GL, K,
S, US); Pirara, Schomburgk 240 (BM, FI); Mabaruma Compound, “bloodwood, ”
Archer 2304 (K, US); Kamakusa, Upper Mazaruni River, Lang 370 (NY):
Isherton, Basin of Rupununi River, A.C. Smith 2443 (K, 8, US).

VENEZUELA: Amacuro: Manoa, Rusby & Squires 143 (US). Anzos-
TEGUI: Cabeceras de Guaraguara, Pittier 14843 (US); Rio Cochama, Mesa de
Guanipa, 240 m., Pittier 14331 (US). Bofvar: Quebrada O-paru-mé, between
Santa Teresita de Kavanayén and Rio Pacairao, 1065-1220 m., “minchu-warei-
yek,”’ Steyermark 60423 (US); Medio Cauré River, Salto de Pard, 170 m.,
L. Williams 11472 (US); Sabana de Guayapo, Bajo Caur4 River, 100 mi., L.
Williams 11863 (US); La Paragua, 70 m., L. Williams 12612 (US); hetw een Ciudad
Bolivar and Rfo Caroni, 100 m. | Steyermark 57579 (US); Gran Sabana, between
Kun and Ruémert, south of Mount Roraima, 1065 m., Steyermark 59142 (US);
vicinity of Tumeremo, 305 m., “sangrito,’”’ Steyermark 60936 (US) ; Cuidad Bolfvar,
Holt & Gehriger 190 (US), Bailey & Bailey 1335 (US). Amazonas: San Carlos
de Rio Negro, 100 m., “lacre blaneo,’”’ L. Williams 14529 (US).

COLOMBIA: Bo.fvar: San Martin de Loba, Curran 67 (US). Vaupfgs:
San Felipe, below confluence of Rfo Guainia and Rio Cassiquiare, Rio Negro, ca.
180 m., “puinave,” “teen-k4,” Schultes, Baker, & Cabrera 18083 (US). Caquerti:
Morelia, 150 m., von Sneidern 1318 (US). AMAZON AS: Leticia, Trapecio amazéni-
co, 100 m., Schultes 8208 (US); Rio Humacayacu, Trapecio amazénico, above 100
m., Schultes 8252 (US). Eu Vauue: Prov. Buenaventura, 1200 m., July 1853,
Triana 3 (BM, COL).

ECUADOR: Santraco-Zamora: Above Valladolid, Rfo Valladolid, 2100-
2400 m., Steyermark 54726 (US).

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 327

PERU: San Marrtfn: Zepelacio, near Moyobamba, 1200-1600 m., Klug
3455 (F, K, 8, US). Loreto: Yurimaguas, Lower Rio Huallaga, 135 m., Killip &
Smith 27580 (US), 28205 (US); Iquitos, 100 m., Killip & Smith 27039 (F, US).

BRAZIL: Amazonas: Near Livramento, on Rio Livramento, Munic.
Humayt4, Krukoff 6865 (A, US); Paranagua, basin of Rio Jurua, Krukoff 4566
(A, US); road to Boa Vista, Caracarahy, Froes 22948 (NY); Ega, Poeppig 2532
(W); Esperanga, mouth of Rio Javari, Ducke 1882 (US); Cucuhy, Rio Negro, 120
m., Holt & Gehriger 373 (US); Amapa, Huber 1219 (G); Mandus, 25 m., Killip &
Smith 30089 (US); near mouth of Rio Embira, tributary of Rio Tarauaca, Krukoff
8479 (8, US); Parintins, Ducke 117 (US); near Palmares, Munic. Séo Paulo de
Olivencga, Krukoff 8479 (S, US). Par: Santarém, August 1850, Spruce (E, G,
NY, TCD).

Vismia cayennensis and V. glabra are very close in nearly all mor-
phological characters. The buds of V. glabra are hairy on the outside,
and those of V. cayennensis glabrous. The most noticeable feature of
V. glabra is that the panicle-branches are pubescent right up to the
peg-like swelling upon which each pedicel articulates; sometimes this
swelling is clothed with a minute fringe of hairs. V. cayennensis is
without this pubescence. The leaves of V. glabra are often larger
than those of its relative but this is not uniformly true.

When Persoon established Vismia cayennensis he did not mention
the earliest use of the epithet by Jacquin, but by followmg back
through the literature by way of Willdenow,” to whom Persoon refers,
the ultimate name-bringing basionym is established.

In Humboldt, Bonpland, and Kunth’s ‘Nova Genera et Species
Plantarum,” V. acuminata (Lam.) Pers. is included on the basis of
a specimen collected near Curichana, on the Rio Orinoco near the
confluence of the Rio Meta. I have examined the Humboldt and
Bonpland collection at Paris and find that it represents almost typical
V. cayennensis. It does not have the long-acuminate leaf-tips of the
form described as acuminata. The sheet bears the nomen nudum
Hypericum eugeniaefolium Willd., indicated as a synonym by Sprengel.
18. Vismia laevis Triana & Planch. Ann. Sci. Nat. IV. Bot. 17: 303. 1862.

Caopia laevis Kuntze, Rev. Gen. Pl. 1: 59, 1891.

Typge: “Foréts du Quindio, alt. 2000 metres, prov. de Mariquita et foréts
d’Antioquia (Tr.).” The first syntype is represented by a specimen “Foréts du
Quindio, 1400 m.,” Triana 262 (K). The second syntype is represented by
“Prov. Antioquia, 2000 [meters ?], n.v. ‘carate,’ Mayo 1852,” Triana 1 (COL,
Photo U.S. Nat. Herb. 2967). The latter is designated as lectotype; isolecto-
types, BM, P, Photo FM 35235.

ADDITIONAL SPECIMENS EXAMINED:

COLOMBIA: Antioquia: Rio Negro, Medellin, 2100 m., Sandeman 5552A
(K), 5712 (K), Archer 313a (US); La Sierra, 2000 m., Archer 1323 (NY, U8);
Llano Grande, near Rio Negro, 2200 m., “punta de lanza,” Daniel 3852 (US);
Santa Elena, 2500 m., Rios C. et al. 619 (US); San Pedro, “‘carate,” Tomas 168

25 Sp. Pl. 3: 1489. 1803.

328 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

(US); La Ceja, Daniel 2197 (US); Alto Catatumbo, north of Yarumal, 2300 m.,
Core 620 (NO). Catpas: Portachuela, Quindio, 1844, Goudot (P). Tota:
El] Libano to Murillo, in subpéramo, 2200-2900 m., Garcia-Barriga 12261 (US).

Vismia laevis, a relative of V. cayennensis, is an endemic species of
central Colombia, characterized by longer, more narrowly lanceolate
leaves than those of its relative, the blades tapering to long-acuminate
tips, the texture distinctly more coriaceous. The sepals are more
prominently hyaline-bordered, with a distinct broad flange. The
pedicels are stout, peg-like, and tend to persist on older fruiting
branchlets. ‘The fruits average larger than those of V. cayennensis
(12-20 mm. long), and are fusiform rather than subspherical, with
conspicuous spreading or reflexed sepals. In certain of these char-
acters V. laevis stands between V. cayennensis and V. glabra.

19. Vismia glabra Ruiz & Pav. Syst. Veg. Peruv. Chil. 183. 1798.

Caopia glabra Kuntze, Rev. Gen. Pl. 1: 59. 1891.

Type: ‘Peru,’ without further indication. An authentic specimen in the
Delessert Herbarium (G, Photo FM 23954) received from Pavon is designated
as lectotype; there is another specimen in the Delessert Herbarium (G, Photo
FM 8005) which is presumably a duplicate. Other authentic collections, perhaps
isotypes: ex Herb. Lambert (BM), ex Herb. Pavon (K), and Herb. Boissier
(G). Possible isotypes are ‘‘Dombey 640” (P) and ‘‘Cochero, Dombey’’ (P).
Another collection Dombey 639, without locality (F) may be a syntype.

ADDITIONAL SPECIMENS EXAMINED:

PERU: Loreto: Above Pongo de Manseriche, 205 m., “pichirina,’”’ Mezia
6306 (CAS, F, 8, UC, US); Pumayacu, Klug 3140 (A); Mishuyacu, near Iquitos,
100 m., Klug 1009 (F, US); Paraiso, Alto Rfo Itaya, 145 m., L. Williams 3366
(F), L. Williams 41 (F), 136 (F), 3310 (US); San Antonio, Rfo Itaya, 110 m.,
Killip & Smith 29410 (US), L. Williams 3324 (F), 3396 (F); Iquitos, 100 m.,
Killip & Smith 27075 (F, US), Tessmann 3659 (G, NY); Pebas, Rfo Amazonas,
L. Williams 1639 (F); Caballo-Cocha, L. Williams 2061 (F); Yurimaguas, Lower
Rio Huallaga, Killip & Smith 27987 (F, US); Lower Rio Nanay, L. Williams
357 (F). San Martin: Upper Rio Huallaga, L. Williams 5826 (F); San Roque,
1350-1500 m., L. Williams 7043 (US), 7044 (US); Tarapoto, Mathews 1310
(CGH, GL), L. Williams 5546 (F). HvuAnuco: La Purisima, Distrito Churu-
bamba, 1750 m., Meria 8232 (F, K, NA, 8, U, UC, US); Pampayaco, Poeppig
1020 (G, W, with remarkable ferruginous raceme branches).

BRAZIL: Amazonas: Sado Paulo de Olivenga, near Palmares, Rio Solimdes
Basin, Krukoff 8479 (A, US); Rio Macauhan, basin of Rio Purus, Rio Acre,
Krukoff 5241 (UC, US); Cobija, Rio Acre Ule 9613 (K, Photo FM 9171 of a
specimen in the Berlin Herbarium), 9639 (K); Seringal, Taraguassu, Ule 9909
(K, Photo FM 9172 of a specimen in the Berlin Herbarium).

Vismia glabra is a polymorphic species, the distinguishable forms
being more or less correlated with the topography of the central
Andes and their adjacent plains. Some forms are so strongly marked
as to their vegetative features as to have been named, although not
published, by Melchior of the Berlin Herbarium. The Amazonas
collections—and there are many in our herbaria—are marked by
unusually thin leaves borne on slender petioles, at more or less right

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 329

angles to the stem and long-pedunculate, loosely-flowered panicles.
However, the flowers do not differ in their technical characters
from those of the usual Vismia glabra of higher elevations in the
Andes. These Amazonas collections commonly display lateral as well
as terminal panicles, recalling V. lateriflora from the same region
in this particular.

Vismia glabra is certainly most closely related to V. cayennensis,
and, indeed, some collections are not easily identified; the differentiae
are discussed in the account of V. cayennensis.

Although there was no mention made of Dombey collections when
the species was described, Dombey 639 (F), consisting of merely a
small but mature leaf, agrees closely with the leaf characters of the
authentic collections preserved at Geneva.
19a. Vismia glabra subsp. pozuzoensis (Engler) Ewan, comb. nov.

V. pozuzoensis Engler, Bot. Jahrb. Engler 58, Beibl. 130: 1. 1923.

Type: Pozuzo, Department of Hudnuco, Peru, N. Esperto (Photo FM 9170,
of a specimen in the Berlin Herbarium).

SPECIMENS EXAMINED:

PERU: Casamarca: Tambillo, Jelski 252 (S, US, W). HuAnuco; Moyo-
bamba, Weberbauer 4515 (G); Cueva Grande, near Pozuzo, Macbride 4763 (F);
Pozuzo, Macbride 4574 (F). Junin: Carpapata, Soukup 3434 (US). Without
locality, Mathews 1310 (BM).

BOLIVIA: Santa Cruz: Yungas de San Mateo, 2500 m., Steinbach 8403

p.p. (8).

Resembling typical Vismia glabra but the leaves strongly ascending,
the blades oval, cuneate at the base, rather thick or firm, plane,
sometimes punctate beneath with pin-point pits, the petioles 10-15
mm. long; panicles loosely fewer-flowered, often lateral as well as
terminal; flower buds often ferruginous-woolly on the outside.

This subspecies is evidently restricted in its range. It is described
as a ‘compact shrub or small tree” of ‘sunny brushy slopes” by J. F.
Macbride, who has collected it at the type locality. Plants that
appear to represent the same subspecies, though geographically at
some distance from the Peruvian stations, have been collected in
the Trapecio amazénico, Department of Amazonas, Colombia: Rio
Loretoyacu, 100 m., Schultes 8264 (US), 6691 (US), Schultes & Black
8389 (US); Rio Boiauasst, 100 m., Schultes 6876 (US).

Hipolito Ruiz describes the location of the pueblo of Puzuzo [sic!]
in some detail in Chapter 38 of his Travels,” and I believe he collected
subsp. pozuzoensis there. In any event there is in the British Museum
(Nat. Hist.) a sheet labeled in Pavon’s (?) hand “Vismia sp. nova del
Peru,” from the herbarium of Lambert, who acquired the Ruiz and
Pavon collection. It is a good example of this subspecies.

40 Cf. Field Mus, Publ. Bot. 21:172-183, 1940,

330 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

20. Vismia plicatifolia Hochr. Ann. Cons. Jard. Genéve 21:54. 1919.

Caopia parvifolia Rusby, Phytologia 1: 65. 1934. Typr: Chuquini, Cordil-

lera Real, 900 m., Bolivia, 17-19 April 1926, Tate 1128 (NY).

TypE: Polo-Polo near Coroico, North Yungas, Bolivia, Buchtien 222 (G,
Holotype, Photo FM 9169, from B; isotypes, C, EH, F). Paratypes: Yungas,
Bang 595 (C, E, F, G, K, L, US).

ADDITIONAL SPECIMENS EXAMINED;

BOLIVIA: Without locality, Bang 293327 (K, US). Coroico, North
Yungas, 2000 m., Mexia 4268 (MO, UC); Polo-Polo, near Coroico, Buchtien 6014
(US); Yungas, 1800 m., Rusby 720 (F, K, US), Bang 621 (US); San Francisco,
Werdermann 2525 (Photo FM 9159 of specimen B); Hacienda Simaco, on road
to Tipuani, 1400 m., Buchtien 5464 (MO, 8, US); Ixiamas (Isiamas), 240 m.,
Cardenas 1947 (IK, NY, US), White 1149 (US); Milluhuaya, south of Coripata,
1300 m., Buchtien 4645 (US); Songo, Bang 835 (BM, E, G, K, US); San José,
1500 m., &. S. Williams 385 (IK, US); Buena Vista, Santa Cruz, 450 m., Steinbach
5081 (NY), 6519 (BM, F, K), 7252 (BM, E, K, MO, U, UC); Yungas de San
Mateo, Steinbach 8403 p.p. (F, K, US); Riberalta, Beni, 300 m., Cardenas 4172
(US).

PERU: San Marrfn: San Roque, L. Williams 7043 p.p. (F).

The type of Caopia parvifolia has small, hardly plicate leaves,
unusual for this species, and weaker, more delicate panicles. Tate’s
collection is unique, but another collection, Steinbach 6519, is tran-
sitional to the typical phase. This Bolivian collection from Santa
Cruz comes from a lower elevation than the Tate specimen. However
a second Steinbach collection (no. 7252) from the same locality is
more nearly typical. In short, C. parvifolia can hardly represent
more than the normal variation to be expected within a species.

Certain Peruvian collections prove difficult to place, showing
characters of both Vismia sprucei and V. cayennensis, as well as
V. plicatifolia. These include: Puerto Bermudez, Junin, 375 m.,
Killip & Smith 26439 (US) and Pumayacu, Loreto, 600-1200 m.,
Klug 3140 (US; but no. 3140 at A and MO is V. glabra).

21. Vismia viridiflora Duchass. ex Triana & Planch. Ann Sci. Nat. IV. Bot. 17:

302. 1862.
Caopia viridiflora Kuntze, Rev. Gen. Pl. 1: 59. 1891.

Type: Panama, Duchassaing (likely from the vicinity of Panama City; cf.
Contr. U.S. Nat. Herb. 27: 44). Paratype: Chagres, on Chagres River [now
Canal Zone], Feb. 9, 1850, Fendler 6 (K, OXF, TCD, US).

ADDITIONAL SPECIMENS EXAMINED:

PANAMA: Without locality, Seemann 466 (BM, K, S); Campana, 600-800
m., Allen 1692 (NY, US); Matachin, Hayes 456 (NY); Ancén Hill, 180 m., R. S.
Williams 7 (NY, US); Bismarck, above Penonome, R. S. Williams 559 (NY,
US). Cana Zone: Bohio, 10-20 m., Mazon 4774 (S, US); Chiva-Chiva Trail,
Piper 5763 (S, US); Rio Tapia, Standley 30688 (K, US); Chagres, Fendler 299 (K).

COLOMBIA: Cuocé: Cordillera Occidental, 1200 m., Triana (P).

Vismia viridiflora has the general appearance of V. cayennensis,
from which it differs in its fine, almost microscopic puberulence on

“7 No locality data supplied by account in Bull, N.Y. Bot. Gard. 4:327. 1907.

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 331

the lower surfaces of the more narrowly lanceolate leaves. The other
distinguishing characters mentioned by Triana and Planchon, and
illustrated by Fendler 6 from Panama, are minor and without taxo-
nomic significance.

22. Vismia lehmannii Hieron. Bot. Jahrb. Engler 20, Beibl. 49:53. 1895.

Caopia lehmannii Hieron. loc. cit.

Typrr: Vicinity of Popaydén, Department of Cauca, Colombia, 1750 m., Lehmann
3551 (isotypes BM, K, US).

ADDITIONAL SPECIMENS EXAMINED:

COLOMBIA: Cauca: Popaydn, Lehmann K74 (probably part of type col-
lection) (K, 8, US), BT1086 (A, K, L); Capilla, Lehmann BT450 (L). Eu
VaLuE: Piedra de Moler, Rio Digua, 900-1180 m., Cuatrecasas 14893 (US).
Caquers: Florencia, 420 m., “acre,” Pérez-Arbeldez 692 (US).

Vismia lehmannii is more closely related to V. baccifera than to V.
cayennensis, contrary to the opinion of Hieronymus. The leaves are
nearly farinose-puberulent, recalling the vestiture of V. confertiflora
Spruce, although smaller than those of that species. From forms of
V. baccifera growing in the Popayan region, this local species may be
identified by the small flowers crowded in a compact panicle.

23. Vismia cuatrecasasii Hwan, sp. nov.

Arbor gracilis 20 m. alta, ramis ad 25 cm. diam., castaneis, inter-
nodiis superioribus aliquantum brevibus, 5-7 cm. longis; laminis
foliorum magnis, ovatis vel oblongo-ovatis, ad basim rotundatis,
12-20 em. longis, 7-10.5 cm. latis, cuspidatis, supra vitreo-viridibus,
glabris, infra puberulentis, sparse atro-punctatis vel eglandulosis,
prominente venosis, coriaceis, atque flexilibus, siccitate submembra-
naceis, omnibus breviter petiolatis, petiolis subalatis, 5-12 mm. longis;
paniculis oblongo-pyramidalibus, cum adjunctis ramis in axillis su-
periorum foliorum, rache atque pedicellis rufo-puberulentis ; calycibus
subcampanulatis, submembranaceis, minute tomentulosis, pruinosis,
sepalis ovatis, acutis, 8-9 mm. longis, divaricatis, interdum paullo
atro-pustulato-glandulosis in margine, marginibus hyalinis atque
subobsoletis; petalis lanceolato-ovatis, obtusis, ochroleucis, extus sub-
tiliter atro-lineatis, apice sparse vittatis, intus dense tomentosis,
11-16 mm. longis; fasciculis staminorum quinque omnibus dense
comosis, apice ramosis.

Slender tree up to 20 m. high, the stems to 25 cm. in diameter,
the bark brown scaly, the upper internodes rather short, 5-7 cm.
long; leaves large, leathery and flexible when fresh, thinner when
dried, all shortly petiolate, the petioles 5-12 mm. long, more or less
wing-margined or channeled above, the blades ovate to oblong-ovate,
rounded at the base, 12-20 em. long, 7—10.5 cm. wide, abruptly acute
at apex, the upper surfaces bright lustrous green when fresh, rather
dull in the herbarium, glabrous, the lower surface finely puberulent,

332 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

weakly punctate with black dot-like glands or at times evidently
without glands, the midrib and secondary veins distinct and wholly
anastomosing in submarginal loops; panicle oblong-pyramidal, at
times with supplementary lateral branches in axils of the uppermost
leaves, the rachises and pedicels ruf ous-puberulent; calyx subcamp-
anulate, thin-textured and early withering, finely but unevenly
tomentulose, more or less pruinose, the sepals ovate, acute, 8-9 mm.
long, spreading, sometimes with a few black pustulate glands on the
margins, the hyaline margin nearly obsolete; petals lance-ovate to
narrowly lanceolate, obtuse, cream-colored, finely lineate with black
pencil markings on outside, weakly vittate toward the summits,
densely tomentose within, 11-16 mm. long; stamen columns 5, all
densely comose, branching about midway into several branches:
fruit unknown.

Type in the U.S. National Herbarium, no. 1,950,013, collected between La
Herradura de Orddéifiez and Pefia de Campotriste, Rfo Calima (region of Chocd),
Department of Fl Valle, Colombia, alt. 5-10 meters, 3 March 1944, by José
Cuatrecasas (no. 16687); isotype in the herbarium of Tulane University.

PARATYPES:

COLOMBIA: En Vane: (Costa del Pacifico): Barco, Rio Cajambre, 5-80
m., Cuatrecasas 17199 (NO, US); between Isla de Golondro and La Amargura,
Rfo Yurumangui, 10-40 m., Cuatrecasas 16049 (US); Puerto Merizalde, Rio Naya,
5-20 m., Cuatrecasas 13954 (US); Silva, Rio Cajambre, 5-80 m., Cuatrecasas
17641 (US); Quebrada de Guapecito, Rio Cajambre, 0-5 m., Cuatrecasas 17694
(US).

Vismia cuatrecasasii is evidently restricted to the lowland rain
forests of the Pacific Coast of Colombia. The collections at hand, all
made by Dr. José Cuatrecasas, are very uniform for the genus. The
taxonomic position of this species is clearly with V. obtusa of Ecuador
and Peru, from which it is at once separable by its larger flowers.
From the form of V. lauriformis that grows in the same region, this
species may be distinguished by the darker red tint of the lower leaf
surface, the pubescence obscuring the secondaries, whereas the dark-
colored veins in V. lauriformis contrast prominently with the red-
brown pubescence.

The distinguished botanist, José Cuatrecasas, is commemorated in
the name of this Vismia; he has made the largest and most. critical
collections not only of Vismia but of the rich Colombian flora in
general.

24. Vismia baccifera (L.) Triana & Planch. Ann Sci. Nat. IV, Bot. 17: 298. 1862.

Hypericum bacciferum L. Mant. 277. 1771.

Vismia guttifera Pers. Syn. Pl. 2: 86. 1807. Based on Hypericum bacciferum

L. An illegitimate renaming.

Caopia baccifera Kuntze, Rev. Gen. Pl. 1: 58. 1891.

Type: Said to be from Mexico, collected by J. C. Mutis, but undoubtedly from
Colombia and likely from the vicinity of Mariquita, where Mutis lived and worked.

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 333

There is an authentic Mutis collection in the U.S. National Herbarium (sheet
no. 1,561,386) and another from Lambert’s Herbarium now in the British Museum
(Natural History). Eyma reported in 1932 that no type exists in the herbarium
of the Linnaean Society, London.

Vismia baccifera is the most polymorphic species of the genus.
Typical plants, which are trees of Colombia, are here associated with
three subspecies, each with a more or less consistent geographic range
but commonly connected by a series of morphologically transitional
forms that prove difficult to separate.

Key to the Subspecies

Leaves strongly bicolored, white-tomentulose beneath, bright lively green above,
generally broadly deltoid-ovate or at times ovate or lance-ovate.
V. baccifera subsp. dealbata
Leaves not strongly bicolored white and green, more or less ferruginous or un-
evenly cinereous or canescent beneath, usually dull dark green above, varying
from cordate-ovate or suborbicular to ovate or even lanceolate, seldom
exactly broadly deltoid-ovate.

Leaves ferruginous-tomentulose beneath, lance-ovate, acuminate to an acute
narrow apex, rounded or often subcordate at the base, the principal blades
averaging 14-18 cm. long; petioles often 2 em. long or more; lowland and
middle elevations of Venezuela, Colombia, and Panama.

V. baccifera subsp. ferruginea

Leaves not ferruginous-tomentulose beneath, varying from cinereous to ochre-
yellow sometimes on same branchlet, chiefly broadly ovate but variable,
generally not lanceolate, if the principal blades as much as 14 em. long
then proportionately broader; petioles less than 2 cm. long; interior and
upland Andes from Colombia to Bolivia.

Petals short, 7-9 mm. long; fruiting panicle rather compact, the branches
short, stout; Peru and Bolivia. . . . V. baccifera subsp. subcuneata
Petals longer, 10 mm. long or more; fruiting panicle diffuse, the branches
spreading, slender; Colombia and northern Ecuador.
Y. baccifera subsp. baccifera
24a. Vismia baccifera subsp. baccifera.

SPECIMENS EXAMINED:

COLOMBIA: Norte pz SanranpErR: Hoya del Rio Margua, Quebrada del
Rio Negro, region of Sarare, 1200-1300 m., Cuatrecasas 12932 (US). CuNDINA-
marca: Between Cachipay and Quebrada del Hueso, 1600-1700 m., Cuatrecasas
13591 (US); 11 km. south of La Palma, 1950 m., Little 7392 (NA, NO).
Antioquia: Angelopolis, 1950 m., Gutierrez & Barkley 17C679 (US). Cuocé:
Nuqui, Romero C. 418 (US). Eu VALue: Below Queremal, Rfo San Juan, Hoya
del Rio Digua, 1300-1500 m., Cuatrecasas 23868 (US); Cordoba, Killip 5079
(NY, US). Cauca: La Capilla, 25 km. n. of Popaydn, 1740 m., Killip 38469
(US); ‘‘Las Guacas,” Morales, 1600-1650 m., Pennell & Killip 6298 (US).

The typical tree of Colombia has broadly ovate rather long-petio-
late leaves, that are not strongly bifacial, but have a pale yellowish
or ashy lower surface, a diffuse panicle whose branches are spreading
at right angles to its rachis, moderately large flowers with white petals,
about half again as long as the sepals, the sepals with a prominent

334 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

membranous flange, and baccate fruits. It is these large (12-13 mm.
in diameter, 15-17 mm. long), heavy fruits that must have suggested
the original specific name “baccifera”’ to Linnaeus, who, indeed, may
have received the manuscript name from Mutis.

There is a fragmentary collection in the Chicago Museum of Natural
History Herbarium (no. 940,497) bearing two labels, one a form
label provided by the Museum d’Histoire Naturelle, Paris, and a
second more important ticket which bears the characteristic mark of
the Willdenovian Herbarium at Berlin and the words “Hy peric[um]
bacciferum [vol.] 3 p. 1440.” The pagination refers to Willdenow’s
Species Plantarum. The leaves of this collection, though fragmentary,
are a remarkably close match for some of the leaves accompanying
Mutis 3718 (US 1,563,038), an authentic sheet of Vismia baccifera.
It is possible that this incompletely labeled collection from Bon-
pland’s herbarium may represent portions of the Mutis material.

24b. Vismia baccifera subsp. dealbata (H.B.K.) Ewan, comb, nov.

V. dealbata H. B. K. Nov. Gen. & Sp. 5:184. ¢. 454, 1822.

Caopia dealbata Kuntze, Rev. Gen. Pl. 1: 59. 1891.

V. hamanit Blake, Contr. Gray Herb. 53: 41. 1918. Type: San Félix-
Tachira, Tachira, Venezuela, 16 May 1917, “lancetillo,’ H. M. Curran
& M. Haman 1010 (GH; isotypes, A, K, NY, P, US, WU). The label of
an isotype reads, “Estacién Tachira,” which is a pueblo in the Rio Té-
chira valley; “San Félix” has not been located in the State of Tachira;
it was transcribed by Blake as ‘San Felipe.”

Type: Labeled “No. 1152. Rio Negro. Cassiquiare” from Rio Negro and
Rio Cassiquiare, Venezuela, Humboldt & Bonpland (P, Killip photo 714).

ADDITIONAL SPECIMENS EXAMINED:

SURINAM: Forest of Zandery, 31 May 1916, Samuels (US), 275 (P).

FRENCH GUIANA: Without locality, in 1792, Leblond (BM); Maroni,
in 1862, Melinon (i, US); Cayenne, Jelski (Kx, P, US).

VENEZUELA: Anzodreaur: Cerro Peonia (Cerro Coroy), northeast of
Bergantin, Steyermark 61394 (F). Caranozo: Buenavista, Linden 1502 (FI,
G). Distr. Feverau: Without locality, Birschel (TCD); Caracas, 1400 m.,
Pittier 9184 (US); El Junquito, 1500 m., Lasser 1122 (US); between Caracas and
La Guaira, 1100-1300 m., Pittier 9558 (NY, US); between La Guaira and Rfo
Grande, ‘“onotillo,” Curran & Haman 985 (paratype of V. hamanii) (A, US).
Mériwa: Tabay, 1900-2200 m., “punta de lanza,” Gehriger 351 (MO, US);
Los Teques, 1140 m., Eggers 13031 (US); Capellania, 1600 m., “lancetillo,”
Tamayo 2436 (UC, US); Mérida, 1700 m., Prttier 12748 (G, US); along Rio
Albarregas, Monte Serpa, 1675-2135 m., “mancha ropa,” Steyermark 55934
(US). TAcnuira: Alto de Lirio, between Bramén and Las Delicias, 1890-2285
m., Steyermark 57440 (US). Trugstiuo: “Andes de Truxillo,” Linden 332 (P).

COLOMBIA: Anrioqufa: Fredonia, Toro 1037 (NY). Boyack: La Chap-
6n, 1050 m., Lawrance 303 (K, MO). Caqueri: Solano, near Tres Esquinas,
200 m., “acre,” Little & Little 9501 (transitional to subsp. ferruginea) (NO, US).
CUNDINAMARCA: 10 km. south of Gachalé, 2150 m., Fosberg & Grant 21966
(NO, US); Finea “La Esmeralda,” Vereda Térriba, San Francisco, 1550 m.,
“punta de lanza,’ Garcia-Barriga 10989 (US); near Sasaima, 1600-1800 m.,
Cuatrecasas 9632 (US); between El Salto and El Colegio, 1680 m., Cuatrecasas

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 335

8242 (US); above El Colegio, 1500-1600 m., Dugand 3688 (US); Estacién
Santana, 1600-1700 m., Dugand & Jaramillo 3977 (US); between Sueva and
Gachet&, 1800-2100 m., Cuatrecasas & Jaramillo 11991 (US). Hurna: Santa
Ana, alt. 1650 m., “acre, small tree to 8 m., 10 cm. D.B.H., bark gray, slightly
fissured; flowers brown; produces mucilage for sealing letters; fruit produces
red coloring, and also medicine for livestock, common in dry forest,’’ Little
& Little 7110 (NO, US). Macpauena: Rio Hacha, Purdie (K); around San
Andrés de la Sierra, western slope of Sierra de Santa Marta, 1100-1300 m.,
Pittier 1651 (US); Santa Marta, Purdie (TCD), 600 m. H. H. Smith 803 (K,
MO, UC, US). Mera: Puerto Lopez, Fosberg 20173 (NA, NO), Little & Little
8273 (NO, US); Villavicencio, toward E1 Parrao, 500 m., Cuatrecasas 4568 (US);
Cabuyaro, Los Llanos, Rio Meta, 235 m., Cuatrecasas 3593 (US); Cano Ciervo,
Sierra de la Macarena, 600 m., Philipson et al. 2092 (BM, US); along Rio Ocoa,
southeast of Villavicencio, 500 m., Killip 34381 (US). Norre pp SANTANDER:
Toledo, 1700-1900 m., Killip & Smith 20037 (US); Rio Pamplonita, between
Cicuta and Pamplona, 700 m., Cuatrecasas & Garcta-Barriga 10177 (US).
SANTANDER: Charta, 2000-2600 m., Killip & Smith 19232 (US); Bucaramanga,
1000 m., Weir 93 (K), “sangrito,” Killip & Smith 16335 (NY, US); 10 km. north
of Bucaramanga, 1500 m., Araque-Molina & Barkley 185215 (US); 18 km. south
of Socorro, 1200 m., St. John 20540 (NO, US); Mesa de los Santos, 1500 m.,
Killip & Smith 15074 (US). Tora: Libano, 1100-1300 m., Pennell 3446 (NY,
US).

BRAZIL: Pari: Santarém, May 1850, Spruce (BM, E, FI, K, OXF), Spruce
766 (K).

This subspecies dealbata has long been treated as a full species
perhaps because of the rather striking whitish lower leaf -surface but
a study of a large series of specimens shows this tree of Venezuela
and northern Colombia with strongly bicolored leaves to be a variable
geographic population of the widespread Vismia baccifera. The
number of intermediate specimens that cannot be assigned to the
subspecies or the typical baccifera is too large to regard this subspecies
as distinct from V. baccifera.

There is a notable collection in the Harvey Herbarium of the
Dudley Herbarium, Stanford University, with a ticket reading
“Vismia guianensis, Pers. DC. 5, Porto Rico, 1844.” which is in
fact this subspecies dealbata. No Vismia is known from the island of
Puerto Rico and I find no record of botanical collections having been
made on the island during the year 1844. The specimen matches
collections of this subspecies from coastal localities in Venezuela.
Henrik Johannes Krebs botanized in Puerto Rico, according to
Urban, and traveled on occasion to both North and South America.
Though Urban is not detailed in his dates, Krebs’ period of activity
supports the conjecture that this specimen was taken, not in Puerto
Rico, but at a Venezuelan port on one of these trips. Duplicates
of this collection are preserved at Dublin and at Geneva (ex
Delessert Herbarium).

336 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

The type of Vismia dealbata H.B.K. at Paris does not closely
resemble the illustration of the species (Pl. 454) accompanying the
original description. Perhaps the artist somewhat idealized the
plant for purposes of illustration, but the leaves are drawn as being
more or less undulate, apparently of a firm, thick texture, and tending
to fold along the midrib, whereas the type shows a plant with plane
leaf-blades of a thin or membranous texture, at least as preserved
in the herbarium. However, the leaves are described as “integerrima,”’
which conforms with the type. The word “subcoriacea”’ might be
interpreted as fortifying the artist’s figure, yet I believe that a firm,
cutinized leaf surface is intended, in spite of the evidently thin texture
of the type specimen. In interpreting the species as distinguished
from its close relatives the phrase “subtus pilis stellulatis tenuissime
albido-tomentosa,” descriptive of the white-tomentose character
of the leaves, is important.

24c. Vismia baccifera subsp. ferruginea (H. B. K.) Ewan, comb, nov.
V. ferruginea H. B. K., Nov. Gen. & Sp. 5: 183. 1821.
Hypericum cuspidatum Willd. ex Spreng. Syst. Veg. 3: 351. 1826, pro syn.
V. ferrugineae.
V. cuspidata Steud. Nom. Bot., ed. 2, 1: 787. 1840, nom. nud. Based on H.
cuspidatum Willd.
Caopia ferruginea Kuntze, Rev. Gen. Pl. 1: 59. 1891.

Type: Orinoco River, between Atures and Maypures, in the present state
of Bolivar, Venezuela, Humboldt «& Bonpland s.n. (Humboldt & Bonpland Her-
barium P, Killip photo 713).

ADDITIONAL SPECIMENS EXAMINED!

VENEZUELA: Forests of Guamita, “cedrillo,” Delgado 132 (US). Cara-
BoBO: Near Tinaquielo, Chardon 137 (US). AMAzonas: Esmeralda, 143 m.,
L. Williams 15375 (US). Bo.fvar: Guayapo, 100 m., L. Williams 11754 p-p.
(K); La Union, Medio Cauré River, 120 m., L. Williams 11689 (US).

COLOMBIA: Antioquta: Salto de Guadalupe, 1600 m., Hodge 6964 (US);
Llano Grande, near Rio Negro, 2200 m., Daniel 3850 (US); Angostura, 2000 m.,
Fosberg 21604 (NO); Angeldpolis, 1950 m., Gutierrez & Barkley 17C680 (US);
Fredonia, Toro 187 (NY); Primavera, Medellin, 1560 m., Oct. 1945, Rutz (US);
Palmitas, 1700 m., Scolnik et al. 532 (US). Botrfvar: San Martin de Loba,
Curran 183 (US). Boyack: Between Moniquiré and Arcabuco, 2150 m., Pérez-
Arbeléez & Cuatrecasas 8168a (US). Cauca: 4 km. north of Popaydn, 1680 m.,
Ewan 15860 (NO, US); “El Ramal” to Rio Sucio, west of Popaydn, 1600-1900
m., “a tree in prairie, petals white,” Pennell & Killip 8146 (NY, US). Cnocé:
Rfo Atrato, near Quibdé, 400 m., Araque-Molina &: Barkley 19ChO47 (NO, US);
between Oveja and Quibdé, Archer 1741 (NY, US); Istmina, Rio San Juan, 75 m.,
Killip 35455 (BM, 8, US); banks of Quebiada Togoroma, Killip & Cuatrecasas
39090 (K, US). CuNDINAMARCA: Fusagasugé, André 1411 (NY); Buenavista
to Pipiral, southeast of Quetame, 1000-1200 m., “lacre,’’ Pennell 1674 (US).
Huiua: 30 km. northwest of Palermo, 2100 m., Little 8727 (N O, US); drainage
of Quebrada de la Cuandinosa, 15-20 km, east of Gigante, 1500 m., Fosberg
19851 (NO, US). Mera: Susumuco to Villavicencio, Triana 12 (BM). San-
TANDER: Quebrada Angulo, 4 km. south of Lebrija, 955 m., ‘“‘manchador,” St.
John 20588 (NO, US). Ex Vatue: Alto de las Brisas, Pichindé, Hoya del Rfo

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 337

Cali, 2050-2100 m., Cuatrecasas 18331 (US); Alto de Miravalle, Pichindé,
2020-2080 m., Cuatrecasas 18327 (US).

The rich red-brown lower surface of the leaves of this subspecies
is the first feature to attract attention. The leaf-blades vary widely
from long-acuminate to shortly ovate; they are not usually as dis-
tinctly truncate at the base as those of subsp. dealbata. This sub-
species is frequent in Colombia and in transmontane Venezuela.

24d. Vismia baccifera subsp. subcuneata (Huber) Ewan, comb. nov.
V. subcuneata Huber, Bol. Mus. Goeldi 4: 588. 1906.

Typr: “Quebrada grande do Cerro de Canchahuaya,” basin of Rio Ucayali,
Junin, Peru, Nov. 13, 1898, Huber 1479.
SPECIMENS EXAMINED:

PERU: Junin: Middle Uca Yali, Tessmann 3279 (G, NY, 8); Schunke
Hacienda, above San Ramén, 1300-1700 m., Schunke A103 (US); Pichis Trail,
Porvenir, 1500-1900 m., Killip & Smith 25911 (F, NY, US). San Martin:
Tingo Maria, 800 m. Allard 21629 (US). San Carlos, Mathews 1309 (CGE, GL,
K); Moyobamba, Mathews 1311 p.p. (CGE).

BOLIVIA: Charopampa, 570 m., Nov. 1907, Buchtien 1907 (L, US); San
Carlos, 850 m., Buchtien 889 (F, US), 890 (US), 890a (US), 750 m., 9 Nov. 1907,
Buchtien 2114 (US), 750 m., 11 Sept. 1907, Buchtien 2114 p.p. (K, US); Mapiri,
Rusby 1810 p.p. (US); Sorata, Bang 1724 p.p. (US), 850 m., 16 Dee. 1926, Buchtien
(US); Tumupasa, R&. S. Williams 339 (K, NY, US), 517 (K), 547 (K, NY); Rio
Chimate, 570 m., Tate 545 (NY).

This subspecies is separated geographically from the other com-
ponents of Vismia baccifera, being restricted to Peru and Bolivia,
and is the most distinct morphologically of its subspecies. The
leaves resemble subsp. dealbata at times in being somewhat bifacial
but less uniformly cinereous, and oblong rather than deltoid. The
crowded panicle is perhaps the most striking difference from other
subspecies of V. baccifera, but in this regard the collections are not
uniform and the lack of other distinctive characters leads me to
conclude that like the other taxa of V. baccifera this subspecies is a
recent variation maintained especially by geographic isolation.

25. Vismia lauriformis (Lam.) Choisy, Prodr. Monog. Hyper. 35, 1821.

Hypericum petiolatum L. f. Suppl. Plant. 345. 1781; non H. petiolatum L.
(1764).

H. lauriforme Lam. Encycl. 4: 152. 1789. Renaming of H. petiolatum
L. f. (1781) non L. (1764).

H. arboreum J. F. Gmel. in L. Syst. Nat., ed. 18, 2: 1156. 1792. A renaming
of H. petiolatum L. f. (1781) non L, (1764). Illegitimate.

H. laurifolium Willd. Sp. Pl. 3: 1440. 1803. A renaming of H. petiolatum
L. f. (1781) non L. (1764). Illegitimate.

Vismia laurifolia Pers. Syn. Pl. 2: 86. 1806. A renaming of Hypericum
lauriforme Lam. Illegitimate.

H. sanguineum L. ex Triana & Planch. Ann. Sci. Nat. IV, Bot. 17: 301.
1862. A manuscript name “fide specim. authentici Mutisiani in herb.
Linn,’”’ now in Herb. Linnaean Soc., London.

338 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Caopia lauriformis Kuntze, Rev. Gen. Pl. 1: 59. 1891.

V. calvescens Gilg & Hieron. Bot. Jahrb. Engler 21: 322. 1895. Typr:
Vicinity of Popaydn, Department of Cauca, Colombia, Stuebel 259. Pre-
sumably lost at Berlin.

Type: “Nova Granada,’ i.e. Colombia, communicated by José Celestino
Mutis (authentic coll. US 1,562,666). The type was taken most probably in the
Department of Cundinamarca. The fruit was described by Linnaeus filius,
but the US sheet is only in early flower, and so it is doubtful if that represents
an exact duplicate of the collection in the Linnaean Herbarium.

ADDITIONAL SPECIMENS EXAMINED:

FRENCH GUIANA: Maroni, 1864, Melinon (NY); Cayenne, in 1839(?),
Leprieur (F).

VENEZUELA: Botfvar: Roraima, Schomburgk 861 (BM, G, NY, P), 552
(FI, G, P); along mesa escarpment between Santa Teresita de Kavanayén and
wooded quebrada about 8 km. northwest of Kavanayén, 1220 m., Steyermark
60481 (US); along Rio Karuai, northwest of Santa Teresita de Kavanayén, 1220
m., “minchuba-rei-yek,”’ Steyermark 60827 (US); Gran Sabana, between Kun and
Uaduara-pard, south of Mount Roraima, 1065-1220 m., Steyermark 59091 (US),
59099 (US). AnzoarEcur: Ijigua, northeast of Bergantin, 600-800 m., “lacre,”’
Steyermark 61234 (US). Sucre: Los Altos, Tamayo 2168 (US).

COLOMBIA: Mera: Villavicencio, Pennell 1403 (N Y); Apiai, Villavicencio,
500 m., Cuatrecasas 4757 (US). ANTIOQUIA: Medellin, Tero 757 (NY); Puerto
Valdivia, Rio Cauca, 240-260 m., Metcalf & Cuatrecasas, 30070 (UC, US).
NoRTE DE SANTANDER: Quebrada de Gibralter, Rio Cubugon, region of Sarare,
320 m., Cuatrecasas 13228 (US). SanranpER: Between Puerto Wilches and
Puerto Santos, Killip & Smith 14852 (NY, US), 14890 (NY, US). Cunpina-
MARCA: La Mesa, 1200 m., Triana (K); Guaduas to Palmar (Quebrada Honda),
road to Guaduero, 1040-1150 m., Garcia-Barriga 11778 (US). Cauca: Rfo
Ortega to San Antonio, 1500 m., Pennell & Killip 7263 (NY); Popaydn, Lehmann
2882 (K), 5543 (K). Touma: Libano, 1100-1300 m., Pennell 3446 p.p. (MO).
Mariquita, Humboldt & Bonpland 1715 (P).

Vismia lauriformis is highly variable over its range and at times is
separable from typical V. baccifera and its subsp. dealbata only with
difficulty. The leaves of V. lauriformis, however, are generally
smaller and glossy above. The acute, at times almost cuneate, leaf-
base is an especially useful character for recognition.

In the Popay4n region of southern Colombia, Department of
Cauca, a peach-leaved form of Vismia lauriformis with narrower
leaf-blades, often acuminate to a long point in the manner of adult
leaves of Eucalyptus globulus, more or less replaces the typical tree
of the more northern districts of Colombia. This form has never
been distinguished nomenclaturally, and it does not seem necessary
to do so at this time, but it illustrates again the local endemism of that
interesting region. Collections representative of this form include:
Popayan, Lehmann 2832 (US), 5543 (S, US); La Capilla, 25 km. north
of Popayan, 1740 m., Killip 38480 (US); Tres Cruces, Popay&n,
Yepes, Araque, & Barkley 18Ca063 (NO, US).

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 339

Vismia lauriformis is related both to V. baccifera and to V. guianen-
sis, especially to V. baccifera subsp. ferruginea, and through V.
falcata forms an almost complete phyletic series to V. guianensts.

26. Vismia mandurr Hieron. Bot. Jahrb. Engler 20, Beibl. 49: 54. 1895.

Caopia mandurr Hieron. loc. cit.

Typr: Three syntypes were originally cited, Lehmann 920, 5593, and 6617, the
originals all probably destroyed in Berlin. Lectotype: Lehmann 5593 (US);
isolectotype K; the original was without locality; according to Lehmann’s field-
book (US), no. 5593 has the data: “Vismia mandurr Hieron. Species of trees up
to 15 m. in height, with large, close, nutant crowns. Leaves dark yellow-green,
shiny. Flowers brownish yellow-green. Fruit of the size and shape of a sparrow’s
egg. Colombia. Grows in moderately dense forests on the mountain slopes
above Popaydn, 1800-2400 m.” In the field-book, the data for 6617 (IX) are:
“Vismia mandurr Hieron. ‘Trees up to 12 m. in height with large crowns. Leaves
leathery, dark green. Flowers brownish green. Colombia. Grows in the dense
forest formations around Paisbamba above Popaydn, 1800-2300 m. Flowers
in March.” No. 920 (not seen) according to Hieronymus had the data: “Crescit
in altiplanitie prope urbem Popaydn, alt. s. m. 1200-2000 m., mense Augusto
florens et fructifera.”

ADDITIONAL SPECIMENS EXAMINED:

COLOMBIA: Meta: Hacienda El Pao, southeast of San Antonio Fort-
alecillas, 1710 m., Little 7982 (NO, US). Huria: Parque Arquelégico, 3 km.
west of San Agustin, 1800 m., Little 7617 (NO); San Agustin, Daniel 4101 (US);
north of Santa Ana, 1800 m., Little 7071 (NO, US). Ex Vauie: La Laguna,
Rio Sanquinini, 1250-1400 m., Cuatrecasas 15678 (NO, US, rather atypical).
Cauca: Quebrada de Santo Domingo, Rio Palo, 2470 m., Cuatrecasas 19373 (US) ;
around Huila, Rio Paez Valley, 1600-1900 m., Pittier 1300 (US); Popayan,
Lehmann K75 (F, K); Cristalores, 10 km. east of Timbio, 2225 m., Grant & Drew
10653 (NO, US); between Aguabonita and Candelaria, Rio San José, region of
Moscopién, 2280-2350 m., Cuatrecasas 23561 (US). NariNo: Piedrancha, 1550
m., Fosberg 21090 (US).

From this large series mostly of recent collections, made in part by
field botanists under the Foreign Economic Administration, it is
desirable to draw up an emended description of this endemic Vismia
of southern Colombia:

Leaves dark yellowish-green, rather densely crowded at the ends
of the nodose branchlets, the internodes generally short; leaf-blades
elliptic or oval, often very variable as to size on a single shoot, at
times punctate beneath, long-petiolate, the petioles about 2 cm. long,
rather stout, stellate-tomentose; panicle cymose, compactly short
branching; sepals early spreading, narrowly ovate, reflexed in fruit,
the hyaline margin distinct but narrow; petals narrowly ovate, green-
ish glandular dotted above, tomentose within; fruits large, 10-15 mm.
long, greenish.

Vismia mandurr is evidently closely related to V. lauriformis, and,
like V. panamensis, probably represents a recent derivative from that

598830—62——4

340 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

variable polymorphic species. V. mandurr displays a large close
crown of dark yellowish-green shining leaves, which are at times
punctate, more elliptic and longer petiolate than those of V. lauri-
formis. Also, the sepals are more heavily tomentulose than in V.
lauriformis, and only becoming thinly so in advanced fruit. The usual
crowded panicle recalls that other endemic of southern Colombia,
V. lehmannw, but that species has more numerous and smaller flowers.
From the label records V. mandurr may become a forest tree 6 to 15
meters high with a D. B. H. of 8 to 30 centimeters. The cinereous
leaves give a glaucous appearance in the field and droop on the tree
from their long petioles like the leaves of the garden peach.

I. R. Fosberg records an interesting bit of folk-lore for this species
from the Department of Narifio to the effect that the tree is said
to grow from the dead body of a worm. ‘The worm shown me was
a scarab larva which had a club-shaped fungus growing from it.”
In alternate years the beetle is supposed to emerge, the other years
the tree” (label accompanying Fosberg 21090).

27. Vismia guianensis (Aubl.) Choisy, Prodr. Monog. Hyper. 34. 1821.

Hypericum guianense Aubl. Pl. Guian. 2: 784. t. 311. 1775.

V. caparosa H. 13. K. Nov. Gen. & Sp. 5: 182. 1822. Typx: “In monte
Hyguerote, prope Buena Vista Caracasanorum,” which is in the vicinity
of Los Teques, Aragua, Venezuela (ef. Kew Bull. 1925: 302) (Photo FM
9160 of specimen in Berlin Herbarium, bearing the number 676); isotype
labeled “676. Caracas prope Buenavista” in Humboldt & Bonpland
Herbarium (P).

V. acuminata var. caparosa Choisy in DC. Prodr. 1: 543. 1824.

Caopia guianensis Lyons, Plant Names Sci. & Pop. ed. 2, 94. 1907.

Type: “‘Sylvis et pratis Caiennae et Guianae,”’ that is, the island of Cayenne
and the mainland of French Guiana. Brazilian references from Marcgrav and
Piso are cited as well, which are discussed below.

ADDITIONAL SPECIMENS EXAMINED:

TRINIDAD: Caroni North Bank Road, Britton & Mendelson 821 (US);
Mount Toeuche, Britton, Hazen, & Mendelson 1336 (US); Long Stretch, 25-26
mile posts, Broadway 6837 (8, US); Blanchisseuse Road, top of Morne Bleu,
Broadway 6208 (US); Aripo Savanna, Britton, Broadway, & Hazen 307 (US);
woods at Ganapo, Eggers 1380 (US); woods near Omora, Eggers 1411 (US);
Ganapo, Eggers 1078 (P, UC, US).

SURINAM: Sectie O, B. W. 159b (NY, US); Forest Reserve, Lanjouw 344
(NY); “reg. inter. ad fl. Surinam,” Hostmann & Kappler 1249 p. p. (S).

FRENCH GUIANA: Godebert, Wachenheim 41 (US); Maroni, 1863,
Melinon (NY, US); Karouany, Sagot 64 (P, 8); Cayenne, 1838, Leprieur (NY).

BRITISH GUIANA: Without locality, Schomburgk 607 (FI, OXF);
drainage of Takutu River, Kanuku Mountains, 600 m., A. C. Smith 3187 (8S, US);
Kurupukari, Essequibo River, A. C. Smith 2158 (MO, 8, US); Mount Iramaik-
pang, Kanuku Mountains, 975 m., A. C. Smith 3653 (US); Malali, Demerara
River, de la Cruz 2705 (US); Assakatta, Northwest District, de la Cruz 4321

*s For a discussion of Cordyceps, the “vegetable fly,’? see John Ramsbottom, Mushrooms and Toadstools
(London, 1953), 149-153.

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 341

(UC, US), 4281 (UC, US); Kamwatta, Pomeroon District, de la Cruz 1213 (US);
Kamakusa, Lang 370 p.p. (NY, US), 337 (NY, US), de la Cruz 2760 (US), 2820
(US), 4175 (US); Akyma, Demerara River, above Wismar, Hitchcock 17415
(S, US); Kalakoon, junction of Mazaruni and Cuyuni Rivers, Graham 147 (US);
Kaieteur Falls, Potaro River, de la Cruz 4471 (UC, US); Wanama River,
de la Cruz 3896 (US); bank of Potaro River, Tumatumari, Gleason 345 (US);
Butukari, Gleason 709 (US); upper Demerara River Jenman 4279 (US), 6278
(NY); vicinity of Bartica, Essequibo River, de la Cruz 2011 (US).

VENEZUELA: Sucre: Aricagua, vicinity of Cristébal Colén, Broadway
555 (US). Boxrfvar: Sabafias de Santa Teresa, Santa Elena, Gran Sabajia,
Tamayo 2804 (US); Paraguara, Velez 2389 (US); 1 Palmar, 300 m., Cardona
2113 (US); Cerro Upuima, Caronf, “uadamdé,” 1300 m., Cardona 2244 (US);
fields near Rfo Tirica, Caronf, Guayana, 500 m., Cardona 2200 (US).

COLOMBIA: Botfvar: San Martin de Loba, Curran 193 (US). Mera:
Villavicencio, 500 m., Killip 34340 (US); Puerto Lépez road, near Villavicencio,
Schiefer 729 (UC, US). Caaquxrrd: Florencia, 400 m., Cuatrecasas 8819 (US);
50 km, southeast of Algeciras, Huila, 1650 m., Little 7732 (NO, US). ANTIOQUIA:
Bocan4, 1900 m., Araque-Molina et al. 350 (US); vicinity of Santa Elena, between
Medellin and Rio Negro, 2500 m., Barkley et al. 387 (US); La Ceja, 2430 m.,
Johnson & Barkley 18C770 (US); Guarne, 2500 m., Gutiérrez V. et al. 125 (US);
Medioluna, 1700 m., Molina 15 (US); Boquerén de Medellin, 2500 m., Barkley
et al. 113 (US); Bello, 1500 m., Molina 21 (NO, US); between Medellin and Rfo
Negro, 2500 m., Killip et al. 39878 (US).

BRAZIL: Amazonas: San Gabriel da Cachoeira, Spruce 2170 p.p. (BE, G,
OXF, 8); San Carlos, Rio Negro, Spruce 3115 p.p. (BM, E, NY, OXF), erron-
eously cited as 2115” by Reichardt; Santa Izabel, Rio Negro, Black 48-2421
(NY). Park: Para, Jul-Aug. 1849, Spruce (BM, TCD). PERNAMBUCO:
Tapera, Pickel 333 (US).

Vismia guianensis is a frequent tree in the Central Cordillera of
Antioquia, where it shows certain morphologic differences. The
sepals are sparingly tomentulose on the back and at times subglabrous.
The leaf-blades are thicker, average broader and more obtuse than the
Guiana collections, and the leaves are often crowded on the ends of the
branchlets. The sepals are very prominently reflexed in fruit. The
flowers are now green, now whitish.

Aublet cited a Marcgrav reference under his Vismia guianensis
which may not represent that species. Marcgrav’s descriptions, often
amounting to characterizations of genera rather than of species, are not
precise enough to add certainty to the establishment of the species.
On geographic grounds there is some question that Maregrav could,
by the known distribution of Brazilian Vismias, have encountered V.
guianensis. However, Aublet’s species is sufficiently validated on
other grounds so that this discordant Marcgrav element does not
vitiate its standing.

The Colombian collections of Vismia guianensis from the head-
waters of the Meta and east of the Andes are notably different in leaf-
shape, being narrower, more acuminate, and more closely set on the
branchlets, recalling certain populations of Vismea lauriformis. From

342 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

their resemblance to leafy shoots of the garden peach this transandean
subspecies may be distinguished as:

27a. Vismia guianensis subsp. persicoides Ewan, subsp. nov.

A V. guianensi subsp. guianensi laminis foliorum lanceolatis, tenuiter
attenuatis, conspicue petiolatis differt.

Type in the herbarium of Tulane University, collected at Soratama, Rfo
Apoporis, between Rfo Pacoa and Rfo Kananari, alt. 250 meters, Amazonas-
Vaupés, Colombia, 15 June 1951, by Richard Evans Schultes and Isidoro Cabrera
(no. 12570) (NO); isotype in the U.S. National Herbarium.

PARATYPES:
COLOMBIA: Amazonas-Vaupts: Soratama, Schultes & Cabrera 12747
(NO, US). Mera: Flood plain forest of Rio Meta, Puerto Lopez, 240 m., Little &
Little 8281 (NO, US).

This subspecies is recorded as a ‘small tree” or ‘Tree 8 m., 12 em.
D. B. H. Bark gray, rough, deeply furrowed. Flowers brownish.”
The leaves are rusty, a little paler above, with a microscopic puberu-
lence beneath, the areoles very small, epunctate. The petals are
long, spatulate, rounded at the tip, and a little black-vittate. The
sepals are ciliolate, with a distinct membranous flange.

28. Vismia cavanillesiana Cuatrecasas, Rev. Acad. Colomb. Cienc. 7: 47. 1946.

Tyre: ‘Abajo de Gabinete en la Hoya del Abra de San Andrés,” Department of
Huila, Colombia, 1900-2100 m., Mar. 24, 1940, collected by José Cuatrecasas (no.
8605) (Isotypes, NY, US).

ADDITIONAL SPECIMENS EXAMINED:
Colombia: Llano de St. Martin, Karsten (W). Cauca: Popaydn, 1700 m.,
Triana 2(BM). CunpInaMarca: Fusagasugdé, 1500 m., Triana 4 (BM). Meta:
‘‘Susumuco et Villavicencio, 400-1000 m.” Triana (G, K, W).

Vismia cavanillesiana is a remarkable endemic in several respects.
The large flowers are almost unique in the genus, the petals being
sparingly glandular-dotted on the outside and heavily villous on the
inside. The leaves are unusual, by their large oval long-petiolate
blades, dark green and glossy above, ferruginous-tomentulose beneath,
with all the nerves prominent and raised beneath. In some respects
the leaves recall V. dindeniana but the leaf characters approach some
forms of V. baceifera and perhaps V. cavanillesiana is most closely
related to that species. Certain Colombian collections of V. tomentosa
(for example, Sprague 265, US) approach this species in their leaf
characters but the flowers are smaller. The heavily rufous-tomentose
sepals ally this Vismia with the Brazilian species Vismia martiana
and V. magnolitfolia, as shown in the key, but the floral characters
are otherwise very different.

A collection transitional between Vismia cavanillesiana and V.
tomentosa and further discussed under the latter species is Cuatrecasas
22276 (US), from above Las Brisas, Monte El Tabor, Department of
El Valle, Cordillera Occidental, 1970-2100 m., Colombia.

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 343

29. Vismia tomentosa Ruiz & Pav. Syst. Veg. Peruv. Chil. 183. 1798.
Caopia tomentosa Kuntze, Rev. Gen. Pl. 1: 59. 1891.

This species, originally inadequately described, may be given an
amplified description on the basis of a good series of collections as
follows: Slender tree 3 to 10-20 m. high, the trunk 10-140 cm., in
diameter breast high, the branchlets slender, the ultimate branchlets
finely red-tomentellous, including the rachises of the panicles; leaves
typically large, little if at all reduced above, the blades broadly ovate,
rarely oblong, rounded to cordate at the base, 15-23 em. long, typically
10-12.5 cm. wide or less, often only 5-8 cm., rather abruptly acumi-
nate at tip or cuspidate, rich red-brown-tomentellous beneath, speck-
led with minute simple hairs in addition, dull coppery-brown above,
the veins 12 to 16 pairs, extending to the margin and ascending,
raised beneath, impressed above, all petiolate, the petioles stout,
1-2.5 em. long; panicle ample and compound, or small and few-
flowered, much shorter than the uppermost leaves, the flowers short-
pedicellate, lustrous chocolate-brown; calyx 7-8 mm. long, the sepals
felty-tomentose with a well-defined marginal flange, this becoming
increasingly evident in fruit, the fruiting sepals spreading or reflexed ;
petals spatulate-obovate, floccose with curling hairs on the inner face,
yellow or pale green, 8-11 mm. long, shining with a silk-like sheen,
lineate or vittate; stamens included; fruit ovoid to conical, acute,
more or less 5-grooved, 10 mm. long, wine red or dark brown.

Tyre: “Perou,” likely from vicinity of either Cuchero or Chinchao, Depart-
ment of Hudnueo. Authentic collections: Moricand Herb. (G, photographed
by Maebride, Photo FM 23957), and ex Lambert Herb. (BM, photographed by
Morton, Photo 8049), and ex Herb. Hooker (K, ticketed ‘‘Chinchao,”’ which is
surely a portion of the same plant as represented by the collection at Geneva).
Cf, Field Mus. Publ. Bot. 21: 78. 1940.

ADDITIONAL SPECIMENS EXAMINED:

COLOMBIA: Purumayo: Mocoa, about Puerto Viejo, 580-600 m., Cua-
trecasas 11385 (US). Vaupés: Rio Kananari and Cerro Isibukuri, 250 m.,
Garcia-Barriga 13784 (US). Amazonas-Vaupés: Raudal de Jirijirimo, Rfo
Apaporis, ‘‘tin-k4,”’ Schultes & Cabrera 14548 (US), 14946 (US); Raudal Yayacopi,
Rio Apaporis, Schultes & Cabrera 15364 (US), 16938 (US); Soratama, Rfo Apa-
poris, 250 m., Schultes & Cabrera 12728 (US); Jinogojé, Rio Apaporis, 210 m.,
Schultes & Cabrera 15669 (US).

ECUADOR: Sanrtaco-Zamora: Along Quebrada Achupallas, 2500-2800
m., Steyermark 54542 (F, US); trail between Mirador and Pailas, 2010-2255 m.,
Steyermark 54288a (sterile, F, US).

PERU: Lorrro: Yurimaguas, Maynas, Poeppig 2421 p.p. (G, L, OXF);
Florida, Rio Putumayo, at mouth of Rio Zubineta, 180 m., Klug 2263 (A, BM,
F, G, K, US); Timbuchi, on Rio Nanay, L. Williams 966 (F); Moyobamba,
Mathews 1311 p.p. (CGE, GL); Manfinfa, on upper Rio Nanay, L. Williams 1144
(F); lower Rio Nanay, L. Williams 571 (F, US) ; Lower Rio Huallaga, 155-210 m.,
L. Williams 4016 (F), 4231 (F, US), 4953 (F), 5002 (F, US). San Martin:
Tarapoto, 750 m., L. Williams 6104 (F). Hvuinuco: Cuchero, Poeppig 1361 p.p.

344 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

(K, P, W); Mirador, between Acomayo and Chanchao, 2400 m., Meria 7760
(BM, K, 8, U, UC, US), Mexia 04138 (UC). Junin: Tarma, Huacapistana,
1800 m., Velarde-Nufiez 781 (US); Huacapistana, 1680 m., Sandeman 4510 (K,OXF).

BOLIVIA: La Paz: Mapiri, 750 m., Rusby 1810 p.p. (F, K).

BRAZIL: Amazonas: Municipality S40 Paulo de Olivenga, basin of Belem
Creek, Krukoff 8728 (A, BM, F, G, MO, 8, U, US); Mandus, Cachoeira Grande,
Ule 8923, 31 May 1936, Ducke 206 (F, US), 22 Nov. 1942, Ducke 206 (A, MO,
8, US). Mandus, Feb.-Mar. 1945, Froes (US); Itapumua, Lower Rio Madeira,
Cooper III (US).

Vismia tomentosa grows in both the high forest or selva back from
the rivers and in the montafia along the river banks. Ynes Mexia
describes the bark as “gray flaky over cinnamon brown’’ and the
juice as “thick, gummy, and brick-red.” It is evidently a tree that
begins flowering when it attains three meters in height and a trunk
diameter of ten centimeters. The flowers are variously described as
yellow or pale green, and the capsule wine-red.

Both Choisy and A. P. de Candolle relegated Vismia tomentosa to
the group of dubious species because the description was too brief to
be readily identifiable. The original description reads:

“YV. foliis ovatis acutis subtus tomentosis, racemis terminalibus.
Flor. Per. et Chil. tom. &.

Arbor quadriorgyalis

Habitat in Peruviae nemoribus versus Cuchero Chinchao, Muja,
Pozuzo et Pillao ad Chacahuassi tractus.

Floret a Julio ad Octobrem.”

The type collection preserved in the Conservatoire botanique de
Genéve bears a label reading in part ‘“Perou M°* Payon. 1827.”
This label, added perhaps at the time the material was received by
Moricand, is overlaid, however, by what appears to be the smaller
original ticket, fortunately preserved, reading “Vismia tomentosa.
Peru.” That this collection is authentic is corroborated by the fact
that the branch is only in bud, a point consistent with Ruiz and
Pavon’s having failed to mention floral characters in the original
description. Contemporary descriptions of Vismia so often included
the glands or vestiture of the petals. The characters of the leaves
and of the panicle are both well matched by Cuatrecasas 11385, from
Colombia, and less closely by Krukoff 8728 from Brazil. Both of
these collections originate in altitudes considerably below that of the
type localities.

Actually the collections here referred to Vismia tomentosa are not
morphologically alike in all their characters, Especially variable
are leaf size and texture, and the degree of persistence of the dis-
tinctive red-brown tomentum of the under-surface. The collections
from the lower Rfo Huallaga are hardly typical, though this may be
due in part to the immaturity of the specimens, in that the leaves are

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 345

smaller, narrower, and pale red-brown beneath. All of the collections
agree in having terminal panicles (auxiliary panicles present in Steyer-
mark 54542), exceeded by the uppermost leaves, and in the singular
rich cinnamon-brown-tomentellous lower leaf surface free from glands,
the broad almost cordate base and the acute tips of the mature leaves.

Of particular interest is the excellent collection from above Las
Brisas, Monte El Tabor, Department of El Valle, Cordillera Occi-
dental, 1970-2100 m., Colombia, made by Cuatrecasas (no. 22276,
US), which suggests Vismia cavanillesiana in its long-petiolate leaves
with prominent impressed nerves and ample many-flowered panicle,
but the flowers of this collection are smaller than those of that species
and in many respects this collection agrees with V. tomentosa.

Of interest are two specimens from central Colombia: San Agustin,
Tolima, Sprague 265 (BM, K, US), and an immature collection made
Sept. 22, 1948, by A. Gartner N., from between Medellin and Rio
Negro, Antioquia, 2300-2500 m. (Herb. Fac. Nat. de Agronomia,
Medellin).

30. Vismia billbergiana Beurl. Vet. Akad. Nya Handl. (Stockholm) 1854: 117.
1856. PLATE 7
Caopia billbergiana Kuntze, Rev. Gen. Pl. 1: 59. 1891.

Evidently a small slender tree or shrub 3 to 6 m. high with almost
vinelike branches; leaves of the same size up to the inflorescence, the
blades thin-textured, ovate and distinctly apiculate to lanceolate
and long-acuminate, the tip 1 cm. long or more, sometimes sub-
orbicular, 8-13 (17) cm. long, 5-7 (9) cm. wide, the venation closed,
areolate, each areole with a single central dot, bifacial, finely pu-
bescent beneath with light brown hairs, dark green and very sparsely
pubescent above witb scattered stellate hairs, the petioles short, 1
cm. long; panicle cymose, the flowers small, few, less than 8 in a few-
branched terminal raceme; sepals narrowly lanceolate, acute, finely
rufous-tomentose, 4-5 mm. long, the flange margin narrow to broad,
with a single submarginal black gland; petals lanceolate or narrowly
ovate, 9 mm. long, 4 mm. wide, acute or rounded, vittate, comose
within; fruit globose, 4-8 mm. long.

Tyre: “In montibus, Porto Bello,” Panama, April 1826, by Johan Immanuel
Billberg (no. 231) (S, photo NO and US).

ADDITIONAL SPECIMENS EXAMINED:

PANAMA: Fat River, Prov. Colén, 10-100 m, Pittier 3876 (C, US); Loma
de la Gloria, Prov. Colén, 10-104 m., Pittier 4238 (US); Porto Bello, Prov. Colén,
5-100 m., Pittier 2437 (US); Frijoles, Canal Zone, Piper 5826 (US), Standley
27488 (US), 27510 (US); Fish Creek lowlands, Prov. Bocas del Toro, vicinity
of Chiriqui Lagoon, von Wedel 2383 (US).

Vismia billbergiana is an endemic of Panama recalling V. sessilifolia
on one hand, and V. tomentosa on the other, but certainly more closely

346 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

related to the latter. The leaves which are variable (as is frequent
in mosaic forming vine shoots) are thinner than those of V. sessilifolia
and even more prominently apiculate. The cymose panicles are
distinctive in this Vismia; the inflorescence is inconspicuous since the
flowers are both few and small. The apiculate thin-textured leaves
and few-flowered racemes recall V. viridiflora of Panama and the
possibility of hybrid origin between that species and V. sessilifolia
is suggested. V. billbergiana has evidently been overlooked both in
the field and in the literature. Standley did not mention the species
even in synonymy in considering the Panama flora.

31. Vismia lindeniana Dene. in Turcz., Bull. Soc. Nat. Moscou 31!: 381. 1858.
Lectotyre: Galipan, Venezuela, 1350 m., Funck & Schlim 101 (Photo FM
23955, fragment of isotype, F).
ADDITIONAL SPECIMENS EXAMINED:

SURINAM: Forest of Zandery, Samuels 275 p.p. (A).

VENEZUELA: Without locality, Fendler 41 (US). Distriro FEDERAL:
Sabafias de Agua Negra, Pittier 13785 (US). Matorrales de Agua Negra, 1400
m., L, Williams 9939 (US). Anzodrecui: Quebrada Seca, northeast of Los
Chorros, east of Bergantin, Steyermark 61528 (F). Monaaas: Forested summit
of mountain northwest of Caripe, 1300-1350 m., “lacre’’ Steyermark 61975 (F, US).

The name “Vismia lindeniana”’ was proposed by Funck, later
taken up by Decaisne, and ultimately published by Turezaninow.
A collection bearing a label with Funck’s manuscript name has been
chosen as the type, although the first cited collection, Linden 13,
from Cerro de Avila, Province of Caracas, alt. 7000 feet, might well
have been designated as the lectotype. However, in choosing types
for photographing Macbride and Killip independently selected the
Funck and Schlim collection in the Delessert and Paris herbaria
respectively.

Vismia lindeniana grows to be a small tree four to six meters high
with a trunk as much as twenty centimeters in diameter.

A unique collection made in 1917 in the vicinity of Perija, State of
Zulia, Venezuela, Tejera 10 (US), with the fruit finely pubescent is
possibly teratological for its relatively large fruits, 15-21 mm. long; it
is doubtfully referred here. The hirsutulose or strigulose leaves with
stiff pustulate-based hairs, are notable, but the shape and the size of
the leaves would place it with Vismia lindeniana.

32. Vismia crassa (Rusby) Blake, Contr. Gray Herb. 53: 41. 1918.
Caopia crassa Rusby, Mem. Torrey Club 4: 204. 1895.

Typre: Yungas, Bolivia, Bang 683 (NY; isotypes,BM, E, F, G, K M, O, US).

ADDITIONAL SPECIMENS EXAMINED:

BOLIVIA: Without locality, Miers 178 (BM), Bang 2931 (BM, C, EF, F,
G, 8, US, W, WU); Incacorral to Paracti, 2200-2400 m., Herzog 2298 (L); Siru-

paya, near Yanacachi, South Yungas, 2150 m., Buchtien 364 (US); San José,
South Yungas, 480 m, 2. S, Williams 239 (BM, NY).

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 347

Vismia crassa is a small tree about four meters high, well marked in
the genus for its noticeably thick, firm, ovate leaf-blades, white-lanate
beneath, smooth and shining above, borne on stout petioles. This
endemic of eastern Bolivia must be allied to Vismia lindeniana of
cismontane Venezuela.

33. Vismia martiana Reich. ex Mart. Fl. Bras. 121: 204. #. 37. 1878.
Caopia martiana Kuntze, Rev. Gen. Pl. 1: 59. 1891.

Type: “Ad urbem Mariana,” Prov. Minas Gerais, Brazil, Martius (cf. Martius,
Observ. 890). The sheet in the Munich Herb. bears a number ‘576” and an
annotation in Reichardt’s hand (Photo FM 19549). It agrees well with the plate
but lacks the flowers illustrated there.

SPECIMENS EXAMINED:

BRAZIL: Rio pr JANEIRO: Therezopolis, Barreto 4013 (F); Nova Fri-
burgo, Glaziou 12465 p.p. (P); Canto Gallo, Peckolt (I, labeled in Reichardt’s
hand). Banta: Ilheos, 1821-1824, Riedel (US). Minas Gerais: Langsdorff
(US). Cran: Serra Araripa, Taquara, von Luetzelburg 26248A (F). Pari:
Belterra, Black 47-937 (NY).

Vismia martiana has a small congested panicle overtopped by the
uppermost leaves which are little reduced and are augmented by leaf-
like bracts. The principal leaves are ovate, thick, dull above and
felty-tomentose beneath, the punctate dots fainter than suggested by
Martius’s Plate 37, fig. 15, and the veinlets of the interspaces between
the secondaries more obscure. From V. magnolifolia this species
differs in its rounded, not acute, leaf-bases, its shorter petioles, 8-12
mm. long rather than 15-20 mm. long, and less ferruginous-tomentose
rachises of the panicles. The persistent stigmas of the fruits are con-
spicuous in both species, and in both the fruiting sepals are spreading.
Reichardt contrasted the few-powered panicle of V. magnolifolia
with the many-flowered panicle of V. martiana, and the vittate sepals
of the former with the evittate sepals of the latter. Neither of these
characters, however, are very useful. Only the exceptional specimen
of V. martiana (for example, Peckolt s. n.) shows a many-flowered
panicle and Sello 1366 (Field Museum photo 9165), cited by Rei-
chardt as V:. magnoliifolia, displays as many flowers as average speci-
mens of V. martiana. The vittate character of the sepals of V.
magnoliifolia is hardly convincing and at least occasional specimens of
V. martiana (for example, Peckolt s. n.) show dark raised lines on the
inner face of the sepals. In short, the two species are indeed closely
related and more study of a larger series of collections than I have seen
may show that only a single species exists. Nothing is known to me
of the ecology of the two Vismias in Brazil. Reichardt describes V.
martiana as becoming a tree whereas V. magnoliifolia is a shrub, sug-
gesting from other instances among Andean species that there may be
habitat preferences.

348 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

34. Vismia magnoliifolia Schlecht. & Cham. Linnaea 3: 118. 1828 (as “mag-
noliaefolia’’).
V. hilairtt Gardn. ex Hook. London Journ. Bot. 2: 334. 1843. Tyrer:
Serra dos Orgios, Rio de Janeiro, 900 m., “dry bushy places,’ January
1837, Brazil, Gardner 329 (OXF, isotypes, BM, E, F, FI, G, K, NY,
P, 8, TCD, US, W).
Caopia magnoliaefolia Kuntze, Rey. Gen. Pl. 1: 59. 1891.
Type: “Brasilia aequinoctiali,’’ Sellow (Photo FM 9165 of specimen in Berlin
Herbarium; probable isotypes, K (but lvs. attacked by fungus!), L)
ADDITIONAL SPECIMENS EXAMINED:
BRAZIL: Without locality, Glaziou 11804 (BM, C, K, NY, P, US). Rio
DE JANEIRO: Serra da Estrella, Glaziou 2946 (P). Minas Gerais: Lagoa Santa,
9 May 1866, Warming (C); without locality, Langsdorff (US); Campos, Itacolumi,
Schenck 3623 (C); Vigosa Agricultural College Grounds, 680 m., Mexia 4186
(NY, UC, US), 4873 (NY, U, UC, US); Ouro Preto, Damazio 1358 (G); Sara-
menha, Macedo 2740 (US); Gongosoco, Dec. 1834, “a large shrub, full of deep
yellow juice similar to gamboge; flowers straw-colored, streaked and speckled
with dark red; very common in the coppice woods and on the skirts of the
forest,”?’ Bunbury (CGE).

The relationships of Vismia magnoliifolia with V. martiana have
been discussed under the latter species. When Gardner published
V. hilairit he correctly removed the plant he was describing from V.
guianensis, but he did not consider its possible relationships with
V. magnoliifolia, of which it seems to represent a narrow-leaved
form; the long-petiolate leaves with acute bases, and the generally
few-flowered panicles are evidences of relationship. Collections of
this narrow-leaved form include, in addition to the Gardner type
collections:

BRAZIL: Without locality, in 1835, Riedel (P); without data, no. 6278
(US), possibly Glaziou. Rio pe JANErRo: Serra dos Orgaos, Gardner 321 (CGE);
Wilkes Exped. (NY, US). Minas Gerais: Without locality, Langsdorff (US
1,573,893), Rzedel (US 1,573,615, possibly same source as last, ef. Reichardt);
Serra do Caraga, Claussen 27 (P).
35.Vismia reichardtiana (Kuntze) Ewan, comb. nov.
V. guttifera Salam. ex Turez. Bull. Soc. Nat. Moscou 31!: 382. 1858; non
Pers. (1807). Syntypes: “Collibus Bahiae,”’ Salzmann, Blanchet 3041.
The Salzmann collection may be designated as lectotype; a duplicate has
been examined at E. Duplicates of the syntype Blanchet 3041 have been
seen at BM, FI, and W. Another collection, Blanchet 3520, is cited by
Turezaninow as a narrower, more acuminate leaved variant (cf. V. bac-
cifera var. angustifolia Reich. below).
V. baccifera sensu Reich. in Mart. Vl. Bras. 12!: 204. 1878; non V. baccifera
(L.) Triana & Planch. (1862). The collections cited as V. baccifera by
Reichardt are: Bahia, Salzmann, Lhotzky; Rio das Contas, Martius;
Ilheos, Jacobina, and Moritiba, Blanchet 990, 18638, 3041, 3520; Piaui,
Gardner 2491; Surinam, Wullschlaegel.

V. baccifera var. angustifolia Reich. loc. cit. No specimens were referred to
the variety as distinguished from the species in original publication.
Lectotype: Jacobina, Bahia, Brazil, Blanchet 3520 (an authentic sheet

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 349

labeled var. angustifolia by Reichardt in Herb. Delessert, G.) (Isolecto-

types: BM, F, FI, G, P, W).

Caopia reichardttana Kuntze, Rev. Gen. Pl. 1: 59. 1891. Based on V. bacci-
fera sensu Reich. non V. baceifera (L.) Triana & Planch.
V. cearensis Huber, Bull. Herb. Boiss. II, 1: 313. (28 Feb.) 1901. Type:

Between Fortaleza and Bemfica, Ceard, Brazil, Huber 92 (Boissier

Herb. G).

V. guaramirangae Huber, loc. cit. Type: Guaramiranga, Serra de Baturité,

Ceard4, Brazil, alt. ca. 700 m., Huber 263 (Boissier Herb. G). Topotype:

25 July 1908, Ducke 21274 (Photo FM 9163, of specimen in Berlin

Herbarium).

Type: No type cited by Kuntze; all the specimens cited as bacczfera by Reich-
ardt are thus syntypes. As lectotype may be selected: Gardner 2491 (US), from
Flores, banks of Rio Gurgea, South Piaui, Brazil, August 1839; isolectotypes:
BM, CGE, FE, F, FI, G, K, NY, OXF, P, W.

ADDITIONAL SPECIMENS EXAMINED:

Braziu: Par: Vicinity of Pard, Baker 172 (BM, E, G, L, P, 8, W);
Belém, Schultes 8671 (US), Silva 162 (US); Thomé Asst, Distr. Acard, 50 m.,
Mexia 5981 (CAS, G, MO, P, 8, U, UC, US), 5988 (US); Garup4, Rio Amazonas,
Killip & Smith 30579 (NY, US); Island of Marajé, Kauffman 7 (US). Crear:
Baturité, Loefgren 91 (S); near Forteleza, Ule 9071 (F, K, L, US); Serra de
Ibiapaba, Campo Grande, Dahlgren 965 (narrow-leaved form, F). PERNAMBUCO:
Without locality, Gardner 989 (BM, CGE, E, FI, NY, OXF, P, 8, US), 946
(GL). Banta: Without locality, Bondar 3017 (F), Blanchet 595 (NY); Ilha de
Cal, “capianga,’”’ Curran 107 (US); Maranhas, Salzmann 234 (CGE). Rio DE
JaNnErRO: Porto d’Estrella, Sellow 185 (L, UC).

Vismia reichardtiana in its typical form is a shrub four to six
meters high of the State of Cearé with shiny, stiff, lanceolate leaves;
it is well exemplified by Ule 9071 from that state. The extreme leaf
form is the narrowly lanceolate V. baccifera var. angustifolia Reich.,
well illustrated by Dahlgren 965, also from Cearfé. This may prove
to be but a developmental state of the species when more ample
collections are available.

Vismia guttifera Salam. (1858) is a clearly identifiable name for
this species, but is invalidated by an earlier use of the same epithet
by Persoon (1807).

The morphological distinctions between Vismia reichardtiana and
V. pentagyna are indicated under the treatment of the latter. Judging
from annotated collections at Florence and Geneva, this is the plant
Choisy called Vismia rufescens.

36. Vismia glaziovii Ruhl. Bot. Jahrb. Engler 30: Beibl. 67: 27. 1901.

“Frutex ramis teretiusculis vel paullum compresso-tetragonis,
pruinoso-canescentibus, cito glabriusculis, internodiis 5-6 cm longis;
foliorum petiolis circiter 1 cm longis, supra leviter canaliculatis,
primum dense incano-puberulis, foliorum lamina ovata, basi rotun-
data, cuspidato-acuminata, apice ipso obtuso instructa, 10-15 cm
longa, paullo infra medium 5-6 cm lata, integerrima subcoriaceo-

350 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

chartacea, supra pallide subolivaceo-vel glaucescenti-viridi, nitida,

glaberrima, subtus tomento in junioribus uberiore, canescente, e

pilis stellatis, hyalinis vel basi spadiceolis formato instructa, penni-

nervi, venis ordinis primarii 12-16 parallelis, 7-14 mm distantibus,
reticulatim anastomosantibus; thyrsis terminalibus et axillaribus,
pedunculatis, 4-8 cm longis, rhachide, ramis, pedicellis dense
incano-, rare subferrugineo-tomentosis; calycis laciniis oblongo-
ellipticis vel lato-lanceolatis, extus densissime et persistenter incano-
tomentosis, intus glabris et 3-5-vittatis, integerrimis, tenuiter
membranaceo-marginatis, planis vel vix concavis; petalis flaves-
centibus calycem dimidio superantibus, evittatis et epunctatis,
obovatis, brevissime unguiculatis, acutiusculis, flabellato-venosis,
flavis (?), extus glabris, intus dense pilis incanis, rigidulis vestitis;
staminibus multipartitis, villosis, calycem superantibus; staminodiis
parvis, ellipticis, obtusiusculis, crassiusculis, praesertim apice longe

villosis, vix 1 mm aequantibus; germine globoso, glabro, 2-2.5 mm

longo; stylis 3 mm _ longis, filiformibus, rectis vel subflexuosis,

capitellatis.’”’

Syntypes: “Brasilia: civit. Goyaz ad Corrego Fundo in sylvis prope Jaragua,
m. Aug. fl. (Glaziou n. 20694), ibidem in sylvis ad Rio Areas, m. Sept. fl. (Glaziou
n. 20695).”” Photo FM 9162, of a specimen in the Berlin Herbarium, bears a
printed label indicating it as Glaziou 20695, the second syntype cited, and a
handwritten ticket reading “Glaziou 20694, Corrego Fundo dans le bois prés de

Jaragua, Goyaz, 23 Aoft 1895; arbuste; fl. jaundtre,” indicating the first syntype.
The plant is certainly authentic, but which syntype it is is uncertain.

IT was first inclined to include the name as a synonym of Vismia
pentagyna or, less satisfactorily, of V. reichardtiana. However, V.
glaziovit comes from an area floristically different from Cearé and, on
the basis of other instances of local Brazilian species studied in this
and other genera, I am tentatively accepting the species pending more
evidence. The leaves, judging from the type collection alone, are
broadly ovate, more like those of V. latifolia than either V. pentagyna
or V. rewchardtiana, and the raceme is more densely crowded and the
small flowers are more numerous. The pubescence of the lower
leaf-surface agrees with that of V. reichardtiana.

37. Vismia rusbyi Ewan, sp. nov.

Planta fruticosa vel arborescens, intricate ramosa, dense foliosa,
ramis superioribus tomentulosis, inferioribus glabrescentibus; laminis
foliorum ovatis vel lanceolato-ovatis,11-13 em. longis, 5-6 cm. latis,
submembranaceis, supra pallidis, venis subtus obscuris, subtus cano-
pubescentibus, punctulatis, venulis secundariis anastomosantibus in
venulam submarginalem; petiolis gracilibus, 10-15 mm. longis;
paniculis cymosis, compactis, 4-5 cm. longis; calycibus brevibus,
divaricatis, sepalis anguste ovatis, 5 mm. longis, sparse tomentulosis,

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 351

valde membranaceis secus margines, atro-vittatis; petalis valde
exsertis, intus glanduloso-vittatis, extus venulis hirsutis, laminis
rufo-tomentosis.

Shrub or small tree with shortly branching leafy branchlets, these
finely tomentulose above, glabrescent below; leaf-blades ovate or
lance-ovate, 11-13 cm. long, 5-6 cm. wide, rather thin-textured, dull
above, the veins obscure, finely brownish or silvery pubescent beneath,
glandular-punctulate, the secondary veins connecting in a submarginal
vein; petioles slender, 10-15 mm. long; flowers crowded in a short
cymose panicle 4-5 em. long; calyces short, the sepals soon spreading,
narrowly ovate, 5mm. long, thinly tomentulose, the thin membranous
border prominent, more or less black glandular-vittate; petals about
1.5 times as long as the calyx, vittate-glandular, the veinlets appressed-
bairy beneath with simple hairs, heavily reddish tomentose; fruit
unknown.

Type in the U. 8S. National Herbarium, no. 1,516,639, collected at San Carlos,
Mapiri region, Bolivia, alt. 850 meters, flowers Dec. 16, 1926, fruit Feb. 21, 1927,
by Otto Buchtien (no. 888) ; isotypes in the herbaria of the Royal Botanic Garden,

Edinburgh, the Chicago Natural History Museum, the Missouri Botanical
Garden, and the New York Botanical Garden.

PARATYPES:

PERU: San Martin: Moyobamba, Mathews 1311 (OXF). HuAnuco:
Cuchero, Poeppig 1361 p.p. (BM, F, OXF). Puno: Santo Domingo, 1550 m.,
McCarroll 99 (NY); (?) 3 km. above Santo Domingo, 1950 m., Metcalf 30643
(US, ef. below).

BOLIVIA: Hacienda Casana, Tipuani Valley, 1400 m., Buchtien 7610 (US).
Tipuani-Guanai, Bang 1695 (E, F, G, K, US, WU). Mapiri, 1500 m., Rusby
722 (BM, EF, F, G, K, MO, P, US). Rurrenabaque, 300 m., Rusby 837 p.p. (K,
NY, US), 1271 (NY, US). Sorata, Bang 1724 p.p. (F), Guanai, 600 m., Rusby
860 (NY). Lake Rogagua, 300 m., Rusby 1667 (NY, US).

Vismia rusbyi is related to V. guianensis and may be considered the
southern Andean representative of that more northern species, from
which it differs in having the petals prominently vittate with black
glands. Furthermore, the leaves are finely pubescent and generally
larger than those of V. guianensis; were it not for the glandular-
punctulate lower surface, collections might be taken for the Bolivian
V. buchtienit.

Though Rusby 722 is unquestionably Vismia rusby: the specimens
of that collection are variable among different herbaria; evidently
specimens were collected from several trees in making up the sets.
For this reason it has seemed preferable to select the more uniform
series of Buchtien 888 as type.

Metcalf 30643 (US) from Peru may prove to be another, perhaps
undescribed species. The leaves are oblong and notably thicker, and
the petioles are longer and thicker, but more technical characters are
wanting.

352 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Tentatively placed here is another Peruvian collection, Stork &
Horton 9582 (F, G, NA), from west and above Puente Durand, north
of Huanuco, 1800 m., Department of Hudnuco, Peru, which has the
vestiture and floral characters of this species but deltoid-ovate, acute
leaves that are unusually coriaceous for V. rusbyi and suggest V.
baceifera subsp. subcuneata. It is described by the collectors as a
“tree to 6 m., bark brown; wood hard.” Evidently Poeppig 1361
from Cuchero, Peru, cited above, is the same form.

As indicated under Vismia pentagyna, there is a close affinity be-
tween that species of eastern Brazil and V. rusbyi, and both species
seem to be extralimital segregates of the more northern typically
glabrate V. guianensis.

38. Vismia pentagyna (Spreng.) Ewan, comb. nov.

Symplocos pentagyna Spreng. Syst. Veg. 3: 340. 1826.

Vismia decipiens Schlecht. & Cham. Linnaea 3: 116. 1828. Renaming of
Symplocos pentagyna Spreng.

V. decipiens var. laurifolia Schlecht. & Cham. loc. cit. Based on Symplocos
pentagyna.

V. decipiens var. pyrifolia Schlecht. & Cham. op. cit. 117. Type: Brazil,
Sellow. Probable isotypes are Sellow 158, from “prov. Bahia inter Bahia
et Victoria,” (E, Kk, L, Photo FM 9161, of a specimen in the Berlin
Herbarium; the latter photograph is labeled var. pyrifolia and may be
part of the holotype).

Acrossanthes lhotzkyanus Presl, Bot. Bemerk. in Abhandl. Boehm. Gesell.
Wiss. V, 3: 453, 1845. Type: Rio de Janeiro, Brazil, Lhotzky (not seen),

Caopia decipiens Kuntze, Rev. Gen. Pl. 1: 59. 1891.

Tyre: Brazil, Sellow.

ADDITIONAL SPECIMENS EXAMINED:

SURINAM: Without locality, Hostmann & Kappler 1249 p.p. (F, FI, G,
NY, P).

BRAZIL: Rio pe Janerro: Glaziou 10341 (K), 13571 (K). AMazonas:
Mandus, Schultes 8086 (US), Corner 65 (NY); Parintins, January 16, 1936,
Ducke 130 p.p. (US); Patua, Rio Negro, Baldwin 3270 (atypical, US), Ule 5963
(G); San Gabriel, Spruce 2170 p.p. (P). Pari: Belém, Dahlgren & Sella 355
(US), Archer 7712 (NY, US). Psernampuco: Victoria, Pickel 3594 (US). Baufa:
Without locality, Blanchet 1862 (FI, OXF, P), Glocker 88 (G); Mount Toboa,
Bomfim, Curran 156 (F, NY, UC, US).

Vismia pentagyna is related to V. rusbyi of the more interior districts
of the Amazonian basin, and perhaps even more closely related to
V. reichardtiana of eastern Brazil. V. pentagyna and V. rusbyi are

contrasted in the key to the species. V. reichardtiana may be
compared with V. pentagyna as follows:

pentagyna reichardtiana
Leaves ovate to ovate-elliptic, at- lanceolate, rounded or acute
tenuate at the base at the base
Panicle small, crowded, sessile larger, loosely flowered, pe-
dunculate
Petals punctate, vittate, usually not punctate, weakly vit-

strongly so tate, if at all

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 353

The description of Symplocos pentagyna Spreng. is brief but adequate.
It reads:

“pentagyna* 15. S [ymplocos] foliis oppositis oblongo-lanceolatis acumi-
natis integerrimis glabris, floribus racemosis 5-gynis. Brasil. Sello.”
The superscript asterisk indicates in the customary manner a species
described as new. Chamisso and Schlechtendal recognized this
Sellow collection as a Vismia and, according to a procedure then
common in systematic botany, renamed it rather appropriately
“Vismia decipiens.” There is no reason for abandoning Sprengel s

original epithet.

It is singular that when A. Brand monographed the Symplocaceae
in Engler’s Das Pflanzenreich (1901) he did not dispose of Sprengel’s
binomial in the section “Species excludendae,” nor mention the
species in the account elsewhere. Evidently the inclusion of the
binomial Acrossanthes lhotskyanus in the Index Kewensis was based
on Reichardt’s citation of it since Presl’s obscure is name not listed
elsewhere.

When describing Vismia decipiens Schlechtendal and Chamisso
recognized two component varieties among Sellow’s material, dif-
ferentiated on the basis of leaves, calyx segments, and petals. Of
these distinctions the most obvious is the shorter-leaves of var.
pyrifolia (leaves 9-10 cm. long) as contrasted with the longer-leaves
of var. laurifolia (leaves 10-16 cm. long). Unfortunately, the num-
ber of sheets available of V. pentagyna are too limited to warrant
conclusions on these varieties.

Reichardt placed Vismia decipiens near V. confertiflora in his
treatment of the Brazilian species, but I do not believe its relation-
ship with that species is very close. Both Visma pentagyna (i.e.
V. decipiens of former accounts) and V. reichardtiana are more closely
related to Vismia guianensis than to V. confertiflora, when one dis-
regards the pubescence character unduly stressed by Reichardt
in his groupings “Rufescentes” and “Dealbatae.”’ The very youngest
leaves of Vismia pentagyna may be rufous-tomentose beneath (e.g.
Curran 156 (F) from Bomfim, Bahia), but the pubescence is early
deciduous and the mature leaves are glabrescent with a very fine
close, at times grayish, puberulence.

39. Vismia buchtienii Ewan, sp. nov.

Arbor nana vel suffrutex 5-9 m. altus, ramis gracilibus, paullo
compressis, glabrescentibus, apicibus rufo-pubescentibus; laminis
foliorum lanceolato-ovatis, acuminatis, 9-15 cm. longis, 3.5-5.5 em.
latis, supra lucidis, infra opacis, argenteo-pubescentibus vel minute
stellato-lepidotis, costa prominente, venis secundariis manifestis,
petiolatis, petiolis longis (12-16 mm.); paniculis ramosis, ramis
divaricatis et patentibus, interdum inflorescentiis adjunctis in axillis

354 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

foliorum supremorum, omnibus rachibus et ramis rufo-tomentellis;
sepalis brevibus, anguste ovatis, vix acutis, cinereo-tomentellis,
4.5-5.0 mm. longis, sepalis alternis anguste marginatis, marginibus
membranaceis vittatis et minute ciliolatis; petalis ovatis, quam sepalis
duplo longioribus, minute lineatis, intus dense tomentosis; staminibus
inclusis; fructibus oblongis vel ovoideis, 6-12 mm. longis.

Low tree or shrub 5-9 m. high, with slender branchlets, these a
little flattened, glabrescent in age, finely rufous-pubescent at the
tips; leaves lance-ovate, acuminate, 9-15 cm. long, 3.5-5.5 cm. wide,
shining above, dull, more or less silvery pubescent or lepidote beneath
with scattered fine stellate hairs, especially along the midrib, the
secondary veins evident, moderately long-petiolate, the petioles
12-16 mm. long; flowers in a short, loosely divaricate panicle with
at times supplementary floriferous branchlets in the uppermost
leaf-axils, the rachis and branchlets rufous-tomentulose; calyx short,
cinereous-tomentulose, the tomentum in low longitudinal ribs, the
sepals narrowly ovate, barely acute, 4.5-5.0 mm. long, the alternate
sepals with narrow dark-vittate membranous margins, finely ciliolate;
petals ovate, about twice as long as the sepals, finely lineate, densely
tomentose within; stamens included; fruit oblong or ovoid, blackish
6-12 mm. long.

Type in the U.S. National Herbarium, no. 1,159,313 collected at San Antonio,

Mapiri region, Bolivia, alt. 850 meters, in December 1907, by Otto Buchtien
(no. 2022) ; isotype in the Rijksherbarium, Leiden.
PARATYPES:

PERU: San Martin: Moyobamba, 800-900 m., Weberbauer 4520 (G), in
1838, Mathews (FI, K); Corico, 1500-1800 m., Dec. 1865, Pearce (K); Tarapoto,
750 m., ZL. Williams 5954 (F); Lamas, 840 m., L. Williams 6344(F).

BOLIVIA: La Paz: Basin of Rio Bopi, San Bartolomé, near Calisaya, 750-
900 m., Krukoff 10215 p.p. (G, K, MO, NY, US); San Carlos, Mapiri region, 850
m., Buchtien 886, (US), 887 (US); Copacabana, about 10 km. south of Mapiri,
Prov. Larecaja, 850-950 m., Krukoff 11042 (F, G, K, S, US); Unduavi, 2400 m.
Rusby 719 (NY). Santa Cruz: Buena Vista, Province of Sara, Steinbach 6527
(BM, E, F, K, MO, NY, 8S, U, UC).

BRAZIL: Maro Grosso: Moore 137 (BM), 609 (BM, E, NY, WU); Santa
Anna da Chapada, Malme 2076 (S, one sheet of leaves pinked by leaf cutting ants!):
Burity, northeast of Cuyaba, 675 m. Collenette 167 (K, NY).

Vismia buchtienti is most closely related to V. rusbyi, which it re-
sembles in its leaf characters, but that species has shorter, more
broadly ovate leaf-blades, which are finely punctate beneath. The
pubescence of the lower leaf-surfaces is very similar; however, V.
rusbyi is finely pubescent with short curling but unbranched hairs.
The sepals are more densely tomentose in V. buchtienii. In V. rusbyi
the secondary veins anastomose in a submarginal loop, whereas in
V. buchtienvi they are free to the margin.

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 355

40. Vismia amazonica Ewan, sp. nov.

Arbor 3-15 m. alta, trunco gracili, 3-5 cm. diam., ramis castaneis,
deorsum glabris, sursum aliquantum rufo-tomentulosis sicut axibus
panicularum; foliis superioribus non reductis, laminis omnibus ovatis,
basi obtusis, apice acuminatis, cuspidatis, vel saepe subcaudatis,
supra glabris et nitentibus, subtus subglabratis, valde sparse rufo-
tomentellis praecipue in venis secundariis, 12-15 cm. longis, 5.5—-8.0
cm. latis, petiolatis, petiolis gracilibus 12-14 mm. longis; paniculis
dense cymiformibus 8-10 (13) em. longis, pedicellis ultimis ca. 5 mm.
longis; floribus parvis; sepalis 5 (6) mm. longis, dense tomentellis,
marginibus membranaceis et angustis 0.5 mm. latis, ciliolatis vel
subciliatis; petalis obovatis ovalibusve, breviter unguiculatis, 5-8
mm. longis, simpliciter lineatis, intus villossissimis viridi-flavis; stam-
inibus inclusis; fructibus ignotis.

Tree 3-15 m. high, the trunk slender, 3-5 cm. in diameter breast
high, the branchlets chestnut-brown and more or less rufous-tomen-
tulose, like the rachis of the inflorescence; leaves little if at all reduced
up to the panicle, the blades all ovate, rounded at the base, acuminate
or cuspidate or often contracted to a slender subcaudate tip, glabrous
and shining above, appearing glabrous below, but actually thinly
rufous-tomentulose beneath, more strongly so along the larger veins,
12-15 cm. long, 5.5-8.0 cm. wide, the petioles slender, 12-14 mm.
long; panicle compactly cymosely branching, 8-10 or 13 cm. long, the
ultimate pedicels about 5 mm. long; flowers small, the sepals mostly
5 (rarely 6) mm. long, closely tomentulose with a narrow membranous
border about 0.5 mm. wide, ciliolate along the whole margin or at
least on the distal half; petals obovate or oval, short-clawed, 5-8
mm. long, simply lineate with fine dark lines, copiously hairy within,
ereenish-yellow, the unopened buds light gray-green; stamens in-
cluded; fruit unknown.

Type in the U.S. National Herbarium, no. 1,461,142, collected at Iquitos,

Department of Loreto, Peru, alt. about 100 meters, Aug. 2-8, 1929, by E. P.
Killip and A. C. Smith (no. 27378).

PARATYPES:

BRITISH GUIANA: Karinyi, Upper Essequebo River, Myers 5764 (K,
panicle exceptionally open).

COLOMBIA: Caquertt: Sucre, banks of Rfo Hacha, 1000 m., Cuatrecasas
9015 (US).

PERU: Lorero: Mishuyacu, near Iquitos, 100 m., Klug 317
(F, US), “pichirina,’ Klug 1524 (US). San Marrfn: Tarapoto, 750 m., L.
Williams 5379 (F, US); Moyobamba, Sandeman 163 (K, OXF).

BOLIVIA: CocuaBaMmBa: Colonia Presidente Busch, Puerto Polonia, Rio
Coni, 14 km. east of San Antonio, 395 m., Cardenas & Cutler 7202 (US).

BRAZIL: Amazonas: Parintins, Ducke 130 p.p. (A, F); near Urucurituba,
Munie. Borba, Krukoff 5946 (BM, G, K,8, U, US). Par&: Upper Rio Cupary,

598830—62——_5

356 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

plateau between Xingu and Tapajos Rivers, Krukoff 1075 (A, BM, G, K, 8, U),
1175 (A, BM, G, K, 8, U); Belém, Utinga, Schultes 8071 (US); Santarem, May
1850, Spruce (CGE, NY, TCD).

Most of the collections of Vismia amazonica have been identified
previously as V. confertiflora. V. amazonica was distinguished from
that species, however, by Eyma, in the Utrecht Herbarium without
proposing a name for it. Its small flowers recall V. méerantha,
although actually they are somewhat larger and intermediate in size
between the V. micrantha group and the Section Huvismia. The
leaves of V. amazonica average definitely smaller than those of V.
confertiflora and are ovate and acuminate rather than ovate-oblong
and obtuse. V.amazonica may be distinguished from V. reichardtiana
of eastern Brazil, which it recalls in its lustrous shining leaves, by
the ovate rather than lanceolate blades. V. amazonica differs from
V. confertifora in that the sepals of the former have the hyaline
border distinctly ciliolate. The upper surface of the leaf-blades in
V. amazonica is yellowish-green and shining, and the leaf-blades are
smaller and often rhomboid, whereas in V. confertiflora they are
generally larger and ovate.

41. Vismia confertiflora Spruce ex Reich. in Mart. Fl. Bras. 121: 205. 1878.

Caopia confertiflora Kuntze, Rev. Gen. Pl. 1: 59. 1891.

? V. gracilis Hieron. Bot. Jahrb. Engler 20. Beibl. 49: 52. 1895. Typm:

Zamora, ‘East Andes of Loja,’’ Keuador, 500-1200 m., Lehmann 7735.
The locality determined by reference to Lehmann’s ms. field notes in the
U.S. National Herbarium, (Isotype, K).

? Caopia gracilis Kuntze, loc. cit.

Type: Vicinity of Santarem, Pard, Brazil, September 1850, Spruce 1087 (iso-
types BM, CGH, E, FI, G, K, NY, OXF, TCD, W).

ADDITIONAL SPECIMENS EXAMINED:

COLOMBIA: Mera: Sabanas de San Juan de Arama, Rio Giiejar, 500 m.,
Idrobo & Schultes 1199 (US). Vaurers: Rfo Piraparand, Schultes & Cabrera
17321 (NO, US), 17363 (US). Amazonas-Vaurfis: Raudal Yayacopi, 240 m.,
Schultes & Cabrera 15365 (US); Raudal Jirijirimo, 270 m., Schultes & Cabrera
14978 (US); Jinogojé, 210 m., Schultes & Cabrera 19830 (US). AMAZONAs:
Loretoyacu River, Trapecio Amazonico, 100 m., Schultes 6668 (US), Schultes &
Black 8542 (US); Cafio Guacayd4, Rfo Miritiparand, Schultes & Cabrera 16272
(NO, US). Purumayo: Mocoa, Sprague 361 (BM, K), Schultes & Cabrera 19070
(US); Rio Putumayo, Puerto Porvenir, near Puerto Ospina, 250 m., Schultes
3401A (US), Schultes & Cabrera 18986 (US); Umbria, 325 m., “pichirina,’’ Klug
1861 (A, F, K, 8, US).

ECUADOR: EK Oro: Along quebrada on south and west slopes of Montafia
Sichicay, near Cachicardn, on a tributary of Rfo Minas Nuevas, above Huertas,
east and northeast of Paccha, 2135-2285 m., “jerigoa,”’ “leaves buff-brown below;
calyx ferruginous-brown without, pale green within; petals green without, white-
hairy within; ovary orange; bark brownish-ruddy, peeling like syeamore or
Myrtaceae, inside of bark orange, staining orange and with gummy resin; wood
white, inferior,’’ Steyermark 54112 (US) [ef. V. gracilis Hieron.]. Losa: André
4600 (K). Napo-Pastaza: Zatzayacu, 400-500 m., Merta 7084 (NA, UC, US),

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 357

7113 (F, NA, NY, UC, US); Puyo, Sydow 876 (US). SantTraGo-ZAMoRA: Along
Quebrada Achupallas, 3000-3500 m., Steyermark 54530 (US) [ef. V. gracilis
Hieron.].

BRAZIL: Amazonas: Santarem, Spruce 766 p.p. (P); Tabatinga, Ducke
1883 (K, US).

The sepals of Vismia confertiflora have a broad hyaline flange
contrasting sharply with the body of the sepal. The leaves have a
silvery, almost lepidote appearance from the very thin puberulence,
distinguishing it from V. obtusa, of the same region, which is wholly
glabrous, and punctate beneath with scattered raised black dots.
Both species may have at times short-acuminate leaf-tips.

Vismia confertiflora is a tree of dense bushy habit about 10 to 15
meters high, with gummy orange-red juice, and yellow petals. The
fruit is evidently green when ripe, to judge from the collectors’ field-
notes. V. tomentosa, another species of the upper Amazon basin,
agrees with V. confertiflora in its narrowly obovate, finely vittate,
yellow petals, but the pubescence of the sepals in V. tomentosa is
truly a tomentum, being denser, more felt-like, than the close fine
puberulence of V. confertiflora. Both have the floral leaves exceeding
the panicles, long-petiolate, broadly ovate principal leaves, prominent
flanges on the evittate sepals in flower, and strongly spreading to
reflexed sepals in fruit. The fruit in V. confertiflora, however, is
green at maturity and in V. tomentosa dark brown. There is some
evidence that the habitats contrast to some degree, V. confertiflora
being a spreading tree of the dense upland forests and V. tomentosa
a taller, more ponderous species of the lowland “high forests.”” The
collections studied of V. confertiflora from British Guiana show smaller
leaves and not so prominent hyaline margins of the sepals. In British
Guiana the species is a shrub or small tree about four meters high.
V. gracilis, of Ecuador, represents a narrow-leaved phase which may
prove distinct when better known.

42. Vismia obtusa Spruce ex Reich. in Mart. Fl. Bras. 121: 207. 1878.

Caopia obtusa Kuntze, Rev. Gen. Pl. 1:59, 1891.

Tyre: “Rio Negro, gap6, Feb. 1851, slender tree 25 ft.,” vicinity of Mandus,
Brazil, Spruce 1352 (Photo FM 19550, of specimen in Munich Herbarium; isotypes,
K, P).

ADDITIONAL SPECIMENS EXAMINED:

COLOMBIA: Hurta: Rio Suaza, 1650 m., Little 8540 (NA, atypical).
Amazonas: Rio Hamacayacu, Trapecio Amazénico, between Amazon and Putu-
mayo watersheds, 100 m., Schultes 8242 (US).

ECUADOR: Picuincna: Mindé, Sydow 297 (US); Santo Domingo de
Colorado, Little 6171 (F, US). Navo-Pastaza: Tena, 400 m., Mexia 7153
(US, NA, UC), 7167 (UC, US). Esmpranpas: San Lorenzo, Little 6330 (F, US).

PERU: Lorero: Gamitanacocha, Rfo Mazén, 100-125 m., Schunke 264
(A, F, NA, UC, US).

358 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

BRAZIL: Amazonas: Banks of Igarapé do Crespo, above Cachoeirinha,
near Mandus, “‘lacre,’’ Ducke 697 (US).

Vismia obtusa occasionally bears axillary panicles, supplementary
to the usual terminal inflorescence, as in the type collection, or in
lieu of it, asin Mexia 7153. When the panicles are axillary rather than
terminal the species suggests V. lateriflora Ducke, which has the
leaves more or less cordate at the base and tomentose beneath. Ynes
Mexfa described this species as a spreading shrub with white flowers
(label of Meata 7153).

Spruce (or Bentham?) may have divided the original collection into
two portions: the February, 1851, “Manaus” collection, and a second,
Dec.—Feb. 1850-51, “Barra” collection (CGE, E, F, G, OXF, TCD,
W) sometimes numbered “1352,” which is a very close match for the
Manaus type.

43. Vismia sprucei Sprague, Trans. Bot. Soc. (Edinburgh) 22: 428. 1905.
Tyre: Vicinity of Panuré, Rio Vaupés, Brazil, Spruce 2601 (K). (Isotypesy
BM, C, F, G, NY, OXF, TCD, W).
ADDITIONAL SPECIMENS EXAMINED:

COLOMBIA: Vaupfs: Guaracapuri Cachoeira, east of Mitd, Rfo Vaupes,
Allen 3375 (US).

ECUADOR: Napo-Pastaza: Rio Pastaza, between Rfo Topo, at Topo, and
Mera, 1158-1675 m., Steyermark 54899 (F, inflorescences both axillary and
lateral, US).

PERU: Loreto: Yurimaguas, Lower Rfo Huallaga, 135-180 m., Poeppig
2411 (W), Killip & Smith 27541 (F, US), Mexia 6078 (BM, CAS, F, G, K, NO,
NY, 8, U, UC, US); Caballo-Cocha, L. Williams 2074 (F); La Victoria, L. Williams
3014 (fF, US); Mishuyacu, 100 m., Klug 745 (US). HuAnuco: Huacachi, near
Mufia, Macbride 4089 (F); Lower Rio Huallaga, L. Williams 3827 (F); Huallaga,
1500-1600 m., Weberbauer 6803 (F, US), ca, 1200 m., Macbride 4229 (F); Chinchao,
Sawada 84 (F); Middle Rfo Ucayali, “pichirina,”’ Tessmann 3278 (F, NY, Photo
FM 9167, of specimen in Berlin Herbarium bearing manuscript name meaning
oval-leaved).

BRAZIL: Amazonas: Matupiry, basin of Rio Jurua, Krukoff 4597 (UC,
US); Borba, Rio Madeira, August 1828, Riedel 1318 (US); Cobija, Rio Acre, Ule
9614 (G, K, L).

Vismia sprucei is well marked by the shining amphiglabrous leaves
with areolate veins. V. obtusa shares the same crowded habit of its
foliage, especially below the inflorescence, but in that species the
blades are rounded or barely acute not apiculate, the upper leaf sur-
face is dull, the petioles are generally longer, and the texture thinner.
The sepals even in bud are glabrous; in V. obtusa, they are tomentose,
the tomentum thinning in age. V. sprucet varies in leaf shape from
oval, the usual typical condition, to lanceolate. Like V. obtusa, it
is constant in having the leaves punctate, with marginal vein-loops.

On the trail to San Ramén, near Yurimaguas, Loreto, Peru, Mrs.
Mexia noted this species (her no. 6078) as a “shrub 3 m. high, many

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 359

branched; white flowers; juice orange-colored, staining. Abundant.”
She recorded the vernacular name as “pichirina.”’

During my early studies of the genus I thought Vismia sprucer
undescribed, overlooking Sprague’s description and the isotype pre-
served in the New York Botanical Garden, and named the presumed
new species for Ynes Mexia (1870-1938). Collections encountered
that are so annotated should be referred to this species.

Uncertain Species or Names

Hypericum lanceolatum Lam. ex Steud. Nom. Bot., ed. 1, 420. 1821. Nom.
nud.

No specimen so labeled was found in the Lamarck Herbarium at
Paris.

Hypericum petiolatum L. Sp. Pl., ed. 3, 1102. 1764.
Caopia petiolata Kuntze, Rev. Gen. Pl. 1: 59. 1891.

Not identifiable. There is no specimen so labeled in the Linnaean
Herbarium, London. The original description is as follows:

‘Hypericum floribus trigynis, foliis ovatis petiolatis integerrimis
subtus tomentosis, caule fruticoso tetragono compresso.

“Habitat in Brasilia.

“Statura & Stamina H. Lasianthi. Caulis tetragonus, obtusus.
Stipulae nullae. Folia Citri, petiolata, acuta, subtus obsolete tomen-
tosa. Corymbus brachiatus. Staminum phalanges oblongae,
maxime spectabiles.”’

Vismia guianensis var. 8 glabrata Choisy in DC. Prodr. 1: 542. 1824.

Based on Hypericum bacciferum Marcgr. bras. 96. fig. 1., a Brazilian
plant, and, in part, on a drawing of a Mexican plant identified by the
citation ‘Moc. et Sesse, Ic. fl. Mex. ined.”’

A supporting collection has not been located in the de Candollean
Herbarium.

Vismia humboldtiana Schlecht. & Cham. Linnaea 3: 118. 1828.

Vismia latifolia H.B.K. Nov. Gen. & Sp. 5: 183. 1822; non V. latifolia
(Aubl.) Choisy (1821).

Typn: Banks of Rio Cassiquiare, Amazonas, Venezuela, Humboldt & Bonpland
(presumably at P).

Vismia humboldtiana Schl. & Cham. is a renaming of V. latifolia
H.B.K., non V. latifolia Choisy, which is based on Hypericum lati-
folium Aublet. The type has not been studied. The original de-
scription is as follows:

“VY. ramulis subpuberulis; foliis ovato-ellipticis, acuminatis, sub-
cordatis, supra nitidis, subtus calycibus tenuissime ferrugineo-
tomentosis; paniculis terminalibus, simplicibus, pedunculatis.

360 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

“Crescit ad ripam fluminis Cassiquiare. Floret Aprili.

“Arbor ramulis compresso-tetragonis, laevibus, glabris, junioribus
tenuissime puberulis. Folia opposita, petiolata, ovato-elliptica,
acuminata, basi rotundata, subcordata, integerrima, reticulato-
venosa, nervo medio venisque primariis subtus prominentibus,
subcoriacea, supra glabra, viridia et nitida, subtus punctulata et
tenuissime ferrugineo-tomentosa, pilis stellulatis, 4-4% pollices longa,
2-2% pollices lata. Petioli semipollicares, tenuissime fuscescenti-
tomentosi. Paniculae terminales, pedunculatae, solitariae, simplices,
breves; pedunculo, pedicellis, rhachi ramisque angulatis, tenuissime
ferrugineo-tomentosis. Flores pedicellati; in specimine nostro non-
dum aperti; pedicellis 2 lineas longis. Calyx quinquepartitus,
externe tenuissime fusco-tomentosus; foliolis ovato-oblongis, acutiu-
sculis, coriaceis, margine membranaceo-diaphanis, subaequalibus.
Petala 5, subrotundo-obovata, externe glabra, interne villosa. Stam-
ina Vismeae, in quinque phalanges coalita. Ovarium ovatum,
glabrum. Styli 5 (?). Fructus desideratur.”’

Vismia jelskii Szyszylowicz, Rozprawy, Akademija Umiejetnésci w Krakowie,
Wydzial Matematyczno-przyodniczy, II, 9: 225. 1895.

The original description, taken from a copy in the National Library
of Medicine, Washington, D.C., is as follows:

“Arborescens. Folia petiolata, petiolis dense nigro punctatis 2-3
cm. longis, laminibus ovato-ellipticis, basi rotundatis vel attenuatis,
12-17 cm. longis, 6-10 cm. latis, integerrimis, coriaceis, supra viridi-
bus, glabris, subtus pruinoso-canescentibus glandulisque prominenti-
bus nigro punctatis. Gemmae axillares stipitatae, glaberrimae.
Thyrsi terminales 4-7 cm. longe pedunculati, prostrati, 8 cm. longi,
8-10 cm. lati, ramis patentibus glabris. Pedicelli 10-20 mm. longi,
media parte articulati bracteolatique. Calyx intus glaber, extrinse-
cus pruinoso canescens, laciniis lanceolatis, 7-8 mm. longis, coriaceis,
margine integerrimo anguste membranaceis, albo pubescentibus.
Petala calyce subduplo longiora, oblongo spathulata, extrinsecus
glabra, intus dense ferrugineo villosa. Staminodia claviformia,
apicem versus pilosa. Staminum phalanges quinque, 8 mm. longae,
multiandrae, filamentis specialibus, capillaceis, denso ferrugineo villo-
sis. Ovarium ovoideum, glabrum, styli quinque, erecto-patentes,
stigmatibus depresso-capitatis. Fructus (immaturus ?) baccatus,
oblongus, 15 mm. longus, 10 mm. latus, glaber, calyce reflexo basi
cinctus.

“Vismiae dealbatae H. B. K. et Vismiae confertiflorae Spr. proxima.

“Cutervo, Jelski no. 253.”

Szyszylowicz indicates (op. cit. 216) that Cutervo is in the Depart-
ment of Cajamarca. The species is not definitely identifiable from the

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 361

description; it may be allied with V. rusbyz, since the description sug-
gests very much the specimen Metcalf 30643, mentioned above under
V. rusbyt. Szyszylowicz work was perhaps issued as a doctoral
dissertation under Zahlbruckner.

Vismia latifolia var. glabrescens Sagot, Ann. Sci. Nat. VI, 11: 163. 1881.
Type: An undesignated specimen in the de Candolle Herbarium, Geneva, ‘“‘sub
nomine V. reticulata Poiret.”

This may represent V. macrophylla H. B. K.
Vismia laxiflora Reich. in Mart. Fl. Bras. 121: 203. 1878.
Caopia laxiflora Kuntze, Rev. Gen. Fl. 1: 59. 1891.

Type: Roraima, British Guiana [actually probably Venezuela], in 1841, R.
Schomburgk 835 [an error for 837] (Isotypes: BM, F, FI, G, K, P, W, Photo FM
32271 of a specimen in the Vienna Herbarium). The type collection represents
an immature, few-flowered plant of almost vinelike habit, unique among hundreds
of collections examined. The corollas are unknown, but the calyx in bud and the
characters of the leaves agree with V. falcata Rusby. No other Vismia has been
collected on or near Roraima, unless a collection of V. falcata so labelled was in
fact taken there [Schomburgk 935 (CGE)] and a collection of V. sesszlifolia
(Schomburgk 917).

Vismia schomburgkiana Klotzsch ex Schomburgk, Reisen in Brit.-Guiana
3: 999. 1848, nom. nud.

“Am oberen Pomeroon auf lichten Waldstellen. Bliiht im Sep-
tember und October. Strauch,’ without a reference to a Schom-
burgk collection number.

Vismia sieberiana Klotzsch ex Schomburgk, loc. cit., nom. nud.

Based on an undesignated Schomburgk collection, identified by the

same phrase as the preceding.

362 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Numbered Specimens Cited

ALLARD, H. A.

21629 baccifera subsp. subcuneata

ALLEN, P. H.

1692 viridiflora
3375 sprucei

ANDERSON, C. W.

40 macrophylla
Anprbk, E.

1411 baccifera subsp. ferruginea
4600 confertiflora

Appwun, C. F.
277 +japurensis

ARAQUE-Mo.ina, J., and BARKLEY
F. A.

188215 baccifera subsp. dealbata
19Ch047__baccifera subsp. ferruginea
19Ch134 angusta

’

ARAQUE-Mo.ina, N., et al.

350 guianensis

ArcHeEr, W. A.

313a_ laevis

1323 laevis

1741 baccifera subsp. ferruginea
2304 cayennensis

2660 cayennensis

2706 cayennensis

7712 pentagyna

8053 japurensis

8243 japurensis

Bartey, L. H., and Barney, E. Z.
1335 cayennensis
Baker, C. F.
172 reichardtiana
Batpwin, J. T.
3270 pentagyna
Bana, M.

595 plicatifolia
621 plicatifolia
683 crassa

835 plicatifolia

1695 rusbyi

1724 p.p. baccifera subsp. subcune-
ata, p.p. rusbyi

2931 crassa

2933 plicatifolia

Bark Ley, F. A., et al.

113 guianensis
387 guianensis

BaRRETO, M.

1274 micrantha
2908 brasiliensis
2909 brasiliensis
2910 micrantha
2912 micrantha
4013 martiana
4035 brasiliensis

Beccari, N.
16 japurensis

BILLBERG, J. I.
231 billbergiana

Buiack, G. A.

2421 guianensis
47-937 martiana
24-2421 guianensis

BLANCHET, J.

595 reichardtiana
1862 pentagyna
1933 angusta

3041 reichardtiana
3520 reichardtiana

Bonpar, G.

3017 reichardtiana

Britton, N. L., et al.

307 guianensis
724 cayennensis
821 guianensis
824 cayennensis
837 falcata

1336 guianensis
1778 cayennensis
2542 cayennensis

EWAN—SOUTH AMERICAN SPECIES OF VISMIA

Broapway, W. E.

399 cayennensis
555 guianensis

3035 cayennensis
4139 falcata
4140 cayennensis
5291 falcata?
5661 falcata?
5726 falcata

6208 guianensis
6837 guianensis

BucutTiEn, O.

222 ~piicatifolia

364 crassa

886 buchtienii

887 buchtienii

888 rusbyi

889 baccifera subsp. subcuneata
890 baccifera subsp. subcuneata

890a_baccifera subsp. subcuneata
1907 baccifera subsp. subcuneata
2022 buchtienii
2114 baccifera subsp. subcuneata
4645 plicatifolia
5464 plicatifolia
6014 plicatifolia
7610 rusbyi

BuRcHELL, W. J.
10042 japurensis

B[oscu] W[EsEN] (Surinam)

459 latifolia
635 cayennensis
672 sessilifolia

1404 latifolia
1564 latifolia
1676 latifolia
1772 cayennensis
2711 angusta
2732 latifolia
2869 = latifolia
2919 angusta
4362 angusta
4543 angusta
CARDENAS, M.

1947 plicatifolia
4172 plicatifolia

CArpenas, M., and Cuter, H.
7202 amazonica

363

Carpona, F.

386 japurensis

1229
1418
2113
2200
2244

angusta
macrophylla
guianensis
guianensis
guianensis

CasARETTO, J. (GIOVANNI)

3510

micrantha

CuHarpon, C. E.

137 baccifera subsp. ferruginea

Cuausen, P.

3 brasiliensis

4 brasiliensis

5 parviflora?

27 magnoliifolia
134 brasiliensis

CouLENETTE, C. L.

(“Sr. GEORGE
EXPEDITION’’)

167 buchtienii
588 angusta (forma)

Cork, E. L.
620 laevis
1519 © sessilifolia
Corner, A.

63 macrophylla
65 pentagyna

3593
4568
4757
4772
7658
8242
8605
8819
9015
9047
9632
11305
11385
12932
13228
13591
13954

CUATRECASAS, J.

baccifera subsp. dealbata
baccifera subsp. dealbata
lauriformis
angusta
minutiflora
baccifera subsp. dealbata
cavanillesiana
guianensis
amazonica
japurensis
baccifera subsp. dealbata
angusta
tomentosa
baccifera
lauriformis
baccifera
cuatrecasasii

364

14081 panamensis
14893 lehmannii
15678 mandurr
16049 cuatrecasasii
16068 panamensis
16354 panamensis
16614 rufa

16687 cuatrecasasii
16903 macrophylla
17120 panamensis
17199 cuatrecasasii
17449 rufa

17551 +panamensis
17641 cuatrecasasii
17694 cuatrecasasii
18327 baccifera subsp. ferruginea
18331 baccifera subsp. ferruginea
19373 mandurr
19738 panamensis
21102 rufa

21279 angusta
21389 panamensis
22276 tomentosa?
23561 mandurr
23868 baccifera

CuaTRecasas, J., and GarRctra-
Barriga, I.

10177 baccifera subsp. dealbata

Cuatrecasas, J., and JARAMILLO, R.

11991 baccifera subsp. dealbata

CuatTrRecasAs, J., and Prrez-
ARBELAEZ, FE.

6750 macrophylla

Curran, HIT. M.

67 cayennensis

107 reichardtiana

156 pentagyna

183 baccifera subsp. ferruginea

Curran, H. M. and Haman, M.
1010 baecifera subsp. dealbata
1011 macrophylla

Damazio, L.
13828 micrantha
1358 magnoliifolia
DaH aren, B, FE.

965 reichardtiana

CONTRIBUTIONS FROM THE

NATIONAL HERBARIUM

DawuaRre_en, B. E., and Seua, E.

305

2197
3850
3852
4101

871

1213
1446

1744
1853
2011
2115
2383
2614
2705
2760
2819
2820
3149
3170
3255
3382
3730
3841
3896
4175
4281
4321
4471

pentagyna
DANIEL, BROTHER

laevis

baccifera suhsp. ferruginea
laevis

mandurr

Dawe, M. T.
macrophylla
DE LA Cruz, J. 58.

guianensis
p.p. macrophylla, p.p. sessili-
folia.
macrophylla
angusta
guianensis
macrophylla
macrophylla
macrophylla
guianensis
guianensis
falcata
guianensis
japurensis
japurensis
macrophylla
macrophylla
sessilifolia
macrophylla
guianensis
guianensis
guianensis
guianensis
guianensis

De.ueapo, I.

baccifera subsp. ferruginea

Domsey, J.

glabra
glabra

Duck, A.

cayennensis

p-p. amazonica, p.p. pentagyna
tomentosa

macrophylla

obtusa

cauliflora

EWAN-—-SOUTH AMERICAN SPECIES OF VISMIA 365

1068 macrophylla
1882 cayennensis
1883 confertiflora
12494 sessilifolia
21274 reichardtiana
25054 laterifiora
25055 = cauliflora

Dueanp, A.
3688 baccifera subsp. dealbata

Dueanp, A., and JARAMILLO, R.

2917 angusta
3977 baccifera subsp. dealbata

Eaaers, H. F. A.
1078 guianensis
1118 falcata
1363 cayennensis
1380 guianensis
1411 guianensis
1420 faleata
5731 cayennensis
13031 baccifera subsp. dealbata

Ewan, J. A.
15860 _ baccifera subsp. ferruginea
FaNnsHAWE, D. B.

F627 macrophylla
FENDLER, A.

6 viridiflora

8 macrophylla
41 lindeniana
299 -viridiflora

Focks, H. C.
382 angusta

Forestry DEPARTMENT B[RITISH]
G[uUIANA]

3726 = sandwithii
3805 sandwithii
6469 angusta

6480 sandwithii

Fosspera, F. R.

19851 _ baccifera subsp. ferruginca
20173 baccifera subsp. dealbata
21604 baccifera subsp. ferruginea

Fossere, F. R., and Grant, M. L.

21966 baccifera subsp. dealbata
Frogs, R. L.

22948 cayennensis

IFuncx, N., and Scuuim, L.

101 lindeniana

Garcia-Barriaea, H.

8377 angusta

10989 < baccifera subsp. dealbata
11778 lauriformis

12261 laevis

13784 tomentosa

GARDNER, G,

321 magnoliifolia
329 magnoliifolia
939 reichardtiana
946 reichardtiana
2491 + reichardtiana

Gay, C.

939 minutiflora

GEHRIGER, W.

351 baccifera subsp. dealbata

GILLESPIE, J. W.

P12 panamensis

Guaziou, A. F. M.

2946 magnoliifolia
10341 pentagyna
11804 magnoliifolia
12465 martiana
13571 pentagyna
20694 = glaziovii
20695 glaziovii

Gueason, H. A.

64 sessilifolia
340 macrophylla
345 guianensis
480 macrophylla
481 falcata

551 macrophylla
709 guianensis

GLocKER, C.

88 pentagyna

366 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

GrauaM, E, H.

147 guianensis
292 sessilifolia

Grant, M. L., and Drew, W. B.
10653 mandurr
GuILLEMIN, J. B. A.

498 brasiliensis

GuTIERREZ, G., and BarKuey, F. A.

17C679 _baccifera subsp. baccifera
17C680__baccifera subsp. ferruginea

GUTIERREZ, G., ET AL.

125 guianensis
931 japurensis

Havant, O.
4885 angusta

Hayss, 8.

456 -viridiflora
921 panamensis

Herzog, T.

2208 crassa

Hitcucock, A. 8.

16950 macrophylla
17389 sessilifolia
17415 guianensis

Hopes, W. H.

6964 baccifera subsp. ferruginea

HoeEugneg, F. C.

198 p.p. brasiliensis, p.p. micrantha

Hott, E. G., and Buaxeg, E. R.

537 macrophylla
599 japurensis
633 japurensis

Hott, E. G., and Grenricer, W.

190 cayennensis
297 japurensis
302 japurensis
331 macrophylla
373 cayennensis

HostmMann, F. W., and Kappuer, A.

162 angusta
438 cayennensis
1249 p.p. guianensis and p.p. pen-
tagyna
1823 | sessilifolia
Huser, J.

92 reichardtiana

263 reichardtiana

1219 cayennensis

1479 baccifera subsp. subeuneata

Humpotpt, A. von, and
Boneuanp, A.

676 guianensis

1038 cayennensis

1070 sessilifolia

1151 macrophylla

1152 baccifera subsp. dealbata

1715 lauriformis

Iproso, J., and Scuutres, R. E.

1199 confertiflora

JELSKI, C. DE
252 glabra subsp. pozuzoensis
253 jelskii (see Appendix J)
JENMAN, G. S.

977 = angusta

4279 guianensis

5029 falcata

5035 macrophylla

5324 macrophylla

6278 guianensis

7017p. p. faleata, p. p. sessilifolia

Jounson, W. M., and BarkKtey, F. A.

18C770_ guianensis
Jounston, J. R.

106 cayennensis
KALBREYER, W.

1374 angusta

Kappier, A.

1722 cayennensis

KauFrMan, E.

7 reichardtiana

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 367

Kiuurp, E. P. 1861 confertiflora
11734 sessilifolia 2263 tomentosa
34245 macrophylla 2344 minutiflora? oe
34381 baccifera subsp. dealbata 3140 p.p. glabra, p.p. plicatifolia?
35453 sessilifolia 3455 cayennensis
35455 baccifera subsp. ferruginea Kruxorr, B. A.
37579 = faleata .
38469 baccifera 1075 amazonica
38480 lauriformis 1175 amazonica

1692 macrophylla

Kiuuip, E. P., Barkuey, F. A., and 4566 cayennensis
DANIEL, Bro. 4597 sprucei

39878 guianensis 4735 angusta

4946 cayennensis
Kiturp, E, P., and Cuatrecasas, J. 5241 glabra
5946 amazonica
6325 lateriflora
6865 cayennensis

38976 panamensis
39090 baccifera subsp. ferruginea

Kiuurp, E. P., and Smiru, A. C. 6976 cauliflora
7947 cauliflora

14852 lauriformis 8328 lateriflora
15074 baccifera subsp. dealbata 8479 p.p. cayennensis, p.p. glabra
16335 baccifera subsp. dealbata 8728 tomentosa
19232 baccifera subsp. dealbata 10215 buchtienii
20037 baccifera subsp. dealbata 11042 buchtienii
25911 baccifera subsp. subcuneata 12315 sessilifolia
26439 _ plicatifolia?
26923 angusta KuniMann, J. G.
27039 cayennensis 21223 lateriflora
27075 glabra
27085 _ lateriflora Lana, H.
27221 angusta 337 guianensis
27378 amazonica 370 p.p. cayennensis, p.p. guianensis
27541 ~sprucei
27550 lateriflora Lane H., and Prersaup, A. C.
27580 cayennensis 286 sessilifolia
27987 glabra
28205 cayennensis Lansouw, J.
29410 glabra 96 cayennensis
29475 = lateriflora 344 guianensis .
29689 lateriflora 398 angusta
30089 cayennensis 972 cayennensis
30579 =reichardtiana 1180 cayennensis

34340 guianensis

Lansouw, J., and LinpEman, J. C.

Kuve, G. 1255. sessilifolia

317 amazonica
354 lateriflora
745 sprucei 1122 baccifera subsp. dealbata
888 angusta
1009 glabra
1524 amazonica 303 baccifera subsp. dealbata

LassER, T.

LawRANcE, A. E.

368 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

LEHMANN, F. C,

2832 lauriformis
3551 lehmannii
4003 angusta
5543 lauriformis
5593 mandurr
6617 mandurr
7547 angusta
7735 confertiflora
BT450 Jehmannii
BT1086_ lehmannii
K74 lehmannii
K75 mandurr

LINDEN, J.

13 lindeniana
332 baccifera subsp. dealbata
1502 baccifera subsp. dealbata

Linper, D. H.

105 cayennensis
Litre, EK. L., Jr.

6171 obtusa

6330 obtusa

7392 baccifera

7617 mandurr

7732 guianensis

7982 mandurr

8540 obtusa

8727 baccifera subsp. ferruginea
9783 macrophylla

Lirtue, KE. L., Jr., and Lirrir, R. R.

7110 baccifera

8252 japurensis

8273 baccifera subsp. dealbata
8275 urceolata

8281 guianensis subsp. persicoides
9501 baccifera subsp. dealbata
9502 angusta

LoEFGREN, A.

91 reichardtiana

LuerzELBura, T. von
26248A martiana

Macsripp, J. F.

4089 sprucei
4229 sprucei
4574 glabra subsp. pozuzoensis

4763 glabra subsp. pozuzoensis
5019 lateriflora

Macepo, A.
2740 magnoliifolia
Maguire, B.

23765 cayennensis
23766 macrophylla

Macurre, B., and STaHEL, G.

22779 cayennensis
23624 cayennensis
29053 japurensis

MatmE, G. O.
2076 buchtienii

Marrero, J. and Lirris, FE. L., Jr.

6274 panamensis

Martin, J.
27 latifolia
Martius, K. F. D. von
576 martiana
970 micrantha
MatTHeEws, A.

1309 baccifera subsp. subcuneata
1310 p.p. glabra, p.p. glabra subsp.
pozuzoensis
1311 p.p. baccifera subsp. subcune-
ata, p.p. rusbyi, p.p. tomentosa
Maxon, W. R.

4774 viridiflora

McCarro tu, D.
99 rusbyi

Metcatr, R. D.
30643 ~rusbyi?
Mercatr, R, D., and Cuarrecasas, J.
30070 lauriforimis

Mexfa, Y.

4138 tomentosa
4186 magnoliifolia
4268 plicatifolia
5981 reichardtiana
5988 reichardtiana

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 369

6078 sprucei
6306 glabra
7084 confertiflora
7113 confertiflora
7153 obtusa
7167 obtusa
7760 tomentosa
8232 glabra
Miers, J.

178 crassa

Mouina, F.

15 guianensis
21 guianensis

Moore, 8.
137 buchtienii
609 buchtienii
Motis, J. C.
3718 baccifera subsp. baccifera

Myers, J. G.

5764 amazonica

PENNELL, I. W.

1403 lauriformis

1674 baccifera subsp. ferruginea

3446 p.p. baccifera subsp. dealbata,
p.p. lauriformis

PENNELL, F. W., and Kiuuip, E. P.
6298 baccifera

7263 lauriformis
8146 baccifera subsp. ferruginea

PENTLAND, J. B.

178 tomentosa

PéRreEz-ARBELAEZ, E.

692 lehmannii
2534 baccifera subsp. ferruginea

P£REz-ARBELAEZ, E., and
CUATRECASAS, J.

8168a_ baccifera subsp. ferruginea

Prersaup, A. C.

16 japurensis
44 macrophylla

Puitipson, W. R., Iproso, J. M., and
JARAMILLO, R.

2088 macrophylla
2092 baccifera subsp. dealbata

Picket, D. B.
3594 pentagyna

Pinkus, A. 8.
166 japurensis

Prrer, C. V.

5763 viridiflora
5826 billbergiana

Pirtier, H.

511 sessilifolia

1300 =mandurr

1651 baccifera subsp. dealbata
2437 ~~ billbergiana

3876 billbergiana

4238 billbergiana

9184 baccifera subsp. dealbata
9558 baccifera subsp. dealbata

10961 macrophylla

12748 baccifera subsp. dealbata
13785 lindeniana

14331 cayennensis

14843 cayennensis

Poerria, EK. F.

421 lateriflora
1020 glabra
1361 p.p. rusbyi, p.p. tomentosa
2411 = sprucei
2421 tomentosa
2532 cayennensis
2905 macrophylla
Pout, FE.

3738 micrantha
REGNELL, A. F,
JII-298 brasiliensis

RIEDEL, L.

25 micrantha
187 angusta

370

1318 sprucei
1427 angusta
1474 micrantha
2634 micrantha

Rfos C., D., Scotnix, R., and
BETANCOURT, A.

619 laevis
Romero C., R.

418 baccifera
1158 macrophylla

Russy, H. H.
719 buchtienii
722 rusbyi
837 rusbyi
860 rusbyi
1271 rusbyi

1667 rusbyi

1810 p.p. baccifera subsp. subcune-
ata, p.p. tomentosa

Russy, H. H., and Squires, R. W.

141 falcata

142 falcata

143 cayennensis
720 pilicatifolia

Saacot, P.

64 guianensis
65 sessilifolia
66 cayennensis

St. Hinairg, A.

D55_—soparviflora

sr. JoHN, H.

20540 baccifera subsp. dealbata
20588 baccifera subsp. ferruginea
20589 macrophylla

SALZMANN, P.

234 reichardtiana

SAMUELS, J. A.

112 cayennensis

212 cayennensis

213 cayennensis

275 p.p. baccifera subsp. dealbata,
p.p. latifolia, p.p. lindeniana

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

SANDEMAN, C.

163. amazonica
2273 + angusta
3730 minutiflora
4510 tomentosa
5552A laevis
5712 laevis
SanpwituH, N.Y.

095 angusta

Sawapa, M,

84 sprucei

Scuenck, H.
3577 micrantha
3623 magnoliifolia
Scuierer, H.

729 guianensis
789 angusta

ScHOMBURGK, R.

182 falcata

240 cayennensis

405 macrophylla
560 macrophylla

607 p.p. cayennensis and p.p.

anensis
837 falcata
861 lauriformis
917 sessilifolia

ScHULTES, R.H., et al.

3041a confertiflora

6668 confertiflora

6691 glabra subsp. pozuzoensis
6876 glabra subsp. pozuzoensis
8071 amazonica

8086 pentagyna

8208 cayennensis

8242 obtusa

8252 cayennensis

8264 glabra subsp. pozuzoensis
8389 glabra subsp. pozuzoensis
8448 lateriflora

8449 lateriflora

8542 confertiflora

8671 reichardtiana

8954 japurensis

gui-

12570 guianensis subsp. persicoides

12724 angusta

EWAN—SOUTH AMERICAN SPECIES OF VISMIA 371

12728 tomentosa

12747 guianensis subsp. persicoides

14548 tomentosa
14946 tomentosa
14978 confertiflora
15364 tomentosa
15365 confertiflora
15404 japurensis
15669 tomentosa
16272 confertiflora
16938 tomentosa
17321 confertiflora
17363 confertiflora
18083 cayennensis
18986 confertiflora
19070 confertiflora
19830 confertiflora
46-317 macrophylla

ScHUNKE, C,

133 angusta
264 obtusa

424 baccifera subsp. subcuneata
A103 baccifera subsp. subcuneata

Scounik, R., ARAQuE-Mo tina, J,

and Barriaa, R.
532 baccifera subsp. ferruginea
SEEMANN, B,
466 viridiflora
Seuiow, F.

158 pentagyna
159 angusta

185 reichardtiana
530 brasiliensis
1365 parviflora
1366 =martiana

SieseEr, F. W.
100 _ falcata

Siuva, A.

162 reichardtiana

SMITH, A. C.

2158 guianensis
2162 macrophylla
2443 cayennensis
3187 guianensis
3653 = =guianensis

598830—62——6

803

Situ, H. H.

baccifera subsp. dealbata

Smiru, S. GaLen, and Iproso, J. M.

1464

1318

3434

184

265
361
401

766

1087
1352
2170
2601

3115

27488
27510

macrophylla

SNEIDERN, K. VON

cayennensis
Soukup, J.
glabra subsp. pozuzoensis
SPLITGERBER, F. L.
latifolia
SpracuB, T. A.

tomentosa
confertiflora
cayennensis

Spruce, R.

p.p. baccifera subsp. dealbata,
p.p. confertiflora

confertiflora

obtusa

p.p. guianensis and p.p. pen-
tagyna

sprucei

p.p. falcata, p.p. guianensis

STANDLEY, P. ©,

billbergiana
billbergiana

30688 viridiflora

5081
6519
6527
7252
8403

STEINBACH, J.

plicatifolia

plicatifolia

buchtienii

plicatifolia

p. p. glabra subsp. pozuzoensis,
p. p. plicatifolia

STEYERMARK, J. A.

54288a tomentosa

54542

tomentosa

54726 cayennensis

54899 sprucei

55204 baccifera subsp. dealbata
55934 baccifera subsp. dealbata
57440 baccifera subsp. dealbata
57579 cayennensis

372 CONTRIBUTIONS FROM
59091 = lauriformis

59099 =lauriformis

59142 cayennensis

60423 cayennensis

60434 macrophylla

60481 lauriformis

60784 macrophylla

60827 lauriformis

60936 cayennensis

61234 lauriformis

61394 baccifera subsp. dealbata
61528 lindeniana

61975 lindeniana

Stork, H. E., and Horron, O. B.
9582 rusbyi?

STUEBEL, A.

259 lauriformis

Syvow, H.
297 obtusa
876 confertiflora

Tamayo, F.
2168 lauriformis
2436 baccifera subsp. dealbata
2804 guianensis
2958 angusta

Tater, G, H. H.

545 baccifera subsp. subcuneata
966 japurensis
1128 plicatifolia

Trsera, EF.
10 lindeniana?

TESSMANN, G.
3278 sprucei
3279 baccifera subsp. subeuneata
3614 angusta
3659 glabra
5040 angusta
Tomas, BrorTHer

168 laevis

Toro, R. A,
187 baccifera subsp. ferruginea
737 ~~ lauriformis

1037 baccifera subsp. dealbata

THE

NATIONAL HERBARIUM

TRIANA, J.

laevis
cavanillesiana
cayennensis
cavanillesiana
angusta
sessilifolia
macrophylla
macrophylla
baccifera subsp. ferruginea
laevis

wo

SO om

Cos!

12
262
Tutin, T. G,

374 — sessilifolia
Us, I

pentagyna
japurensis
tomentosa
reichardtiana
glabra
sprucei
glabra

glabra

Varaas, J.
lateriflora
VELARDE-NuNeEz, O.
tomentosa
VE&LEz, |.
2389 guianensis
WaACHENHEIM, Ci,

40
41

cayennensis
guianensis
49 sessilifolia
58 sessilifolia
107 angusta

WEBERBAUER, A.

4515 glabra subsp. pozuzoensis
4520 buchtienii
6803 sprucei

WEDEL, H. von
2383 billbergiana

WEIR, J.

93 baccifera subsp. dealbata

EWAN-—-SOUTH AMERICAN SPECIES OF VISMIA

WERDERMANN, EF.
2525 _ plicatifolia
Waits, O. FE.
1149 plicatifolia

WituiaMs, L.

41 glabra

136 glabra

357 glabra

571 tomentosa
966 tomentosa
1144 tomentosa
1205 angusta
1385 angusta
1460 angusta
1533 angusta
1639 glabra
1734 angusta
1748 angusta
1793 lateriflora
2061 glabra
2074 sprucei
2192 lateriflora
2194 angusta
2364 lateriflora
2691 lateriflora
3014 sprucei
3310 glabra
3324 glabra
3366 glabra
3396 glabra
3821 lateriflora
3827 sprucei
4016 tomentosa
4231 tomentosa
4899 lateriflora
4953 tomentosa

5002
5008
5379
5546
5826
5954
6104
6344
7043
7044
9939
11472
11689
11754

11863
12612
12773
13064
13912
14348
14478
14529
15375
16003

tomentosa
lateriflora
amazonica
glabra
glabra
buchtienii
tomentosa
buchtienii

303

p.p. glabra, p.p. plicatifolia
glabra

lindeniana

cayennensis

baccifera subsp. ferruginea
p.p. baccifera subsp. — fer-
ruginea and p.p. falcata
cayennensis

cayennensis

angusta

japurensis

japurensis

japurensis

japurensis

cayennensis

baccifera subsp. ferruginea
japurensis

Wiuitams, R. 8.

7 viridiflora
99 angusta

239
339
385
5909

crassa

baccifera subsp. subcuneata
plicatifolia

viridiflora

YeEres-AGREDO, S., ARAQUE-MOLINA,

J., and Barktry, F. A.

18Ca063__lauriformis?

INDEX

Synonyms are in italics and page numbers of principal entries are
in boldface.

achiotillo, 299 carate, 299, 327
Acrossanthes, 300 Caspia, 300
lhotzkyanus, 352, 353 cedrillo, 336
bloodwood, 299, 311, 326 Cratoxylum lingustrinum, 296
Caopia, 298, 299, 300 Delphinium, 299
acuminata, 325 Eucalyptus globulus, 338
baccifera, 332 Euvismia, 297, 315, 356
billbergiana, 345 Ficus lyrata, 307
brasiliensis, 316 pandurata, 307
cayennensis, 325 Hypericum, 293, 295, 297, 300
confertiflora, 356 acuminatum, 296, 325
cordata, 310, 312 arboreum, 337
crassa, 346 bacciferum, 295, 332, 334, 359
dealbata, 334 cayennense, 295, 325
decipiens, 352 cuspidatum, 336
ferruginea, 336 eugeniaefolium, 325, 327
glabra, 328 guianense, 296, 340
gracilis, 356 lanceolatum, 359
gquianensis, 340 latifolium, 296, 322, 359
japurensis, 321 laurifolium, 337
laevis, 327 lauriforme, 337
latifolia, 322 petiolatum, 296, 337, 359
lauriformis, 338 reticulatum, 310, 312
laxiflora, 361 rufescens, 296, 320, 321
lehmanniz, 331 sanguineum, 337
macrophylla, 307 sessilifolium, 296, 320, 321
magnoliaefolia, 348 jerigoa, 356
mandurr, 339 lacre, 299, 308, 311, 324, 331, 334, 336,
martiana, 347 338, 346, 358
micrantha, 315 lacre blanco, 326
obtusa, 357 lancetillo, 334
panamensis, 323 Macrocarpaea, 299
parviflora, 318 manchador, 336
parvifolia, 330 mancha ropa, 334
petiolata, 359 Masdevallia, 311
reichardtiana, 349 minchuba-rei-yek, 338
sessilifolia, 320 minchu-warei-yek, 307, 326
sessilifolia var. rufescens, 320 onotillo, 334
tomentosa, 343 papa de lacre, 322
viridiflora, 330 pichirina, 299, 313, 328, 355, 356, 358
caparosa, 299 pichirina blanca, 311
capianga, 299, 349 pina, 326

375

376 INDEX

puinave, 326
punta de lanza, 299, 334
puntelanza, 311
Sambucus, 319
sangre de gallina, 323
sangrito, 299, 311, 326, 335
sangrito rastrojero, 325
Stictopetalum, section of Vismia, 297
swinani, 298, 299, 326
Symplocos pentagyna, 352, 353
Tabebuia longipes, 308
teen-ka4, 326
tin-kd, 343
Trianthera, section of Vismia, 297, 319
floribunda, 315
uadama, 341
uadama-yek, 311
ubapitanga, 317
Vismia, 300
acuminata, 325, 327
acuminata var. caparosa, 340
amazoniea, 304, 355, 356
angusta, 297, 301, 307, 308, 309,
310, 311, 312, 313, 314, 321, 322
angustifolia, 324
baccifera, 298, 300, 303, 305, 306,
320, 331, 332, 333, 334, 335,
337, 338, 339, 342, 348
baccifera var. angustifolia, 348,
349
baccifera subsp. baccifera, 333
baccifera subsp. dealbata, 308,
333, 334, 335, 337, 338
baccifera subsp. ferruginea, 299,
306, 323, 333, 336, 339
baccifera subsp. subcuneata, 306,
333, 337, 352
billbergiana, 302, 345, 346
brasiliensis, 302, 315, 316, 317,
318
brasiliensis var. lastantha, 316
brasiliensis var. longifolia, 316
buchtienii, 306, 351, 353, 354
calvescens, 338
camparaguay, 293
caparosa, 340
cauliflora, 299, 301, 306
cavanillesiana, 304, 315, 342
cayennensis, 298, 299, 301, 303
319, 321, 325, 327, 328, 329, 330
cearensis, 349
confertiflora, 303, 306, 331, 353,
356, 357, 360

Vismia—Continued

cordata, 310

crassa, 304, 346, 347

cuatrecasasii, 302, 331, 332

cuspidata, 336

dealbata, 334, 336, 360

decipiens, 352, 353

decipiens var. laurifolia, 352, 353

decipiens var. pyrifolia, 352, 353

falcata, 305, 324, 325, 339, 361

ferruginea, 311, 336

floribunda, 326

glabra, 296, 327, 328, 329

glabra subsp. glabra, 303

glabra subsp. pozuzoensis, 303,
329

glaziovii, 303, 349, 350

gractlis, 356, 357

guaramirangae, 349

guianensis, 303, 317, 325, 335,
339, 340, 341, 342, 348, 349,
351, 352, 353

guianensis var. glabrata, 359

guianensis subsp. persicoides, 342

guttifera Pers., 332

quttifera Salam. ex. Turez., 348,
349

hamanit, 334

hilairit, 348

humboldtiana, 359

japurensis, 304, 321, 322

jelskii, 360

laceifera, 316

laevis, 303, 327, 328

lasiantha, 316, 317

lateriflora, 302, 304, 310, 313, 314,
321, 329, 358

latifolia Choisy, 297, 306, 308,
310, 322, 350, 359

latifolia H.B.K., 359

latifolia var. acuminata, 325

latifolia var. glabrescens, 361

latifolia var. reticulata, 310

laurifolia, 337

lauriformis, 296, 302, 305, 306,
320, 323, 324, 325, 332, 337,
338, 339, 340, 341

laxiflora, 361

lehmannii, 304, 331, 340

lindeniana, 306, 342, 346

longifolia, 316, 317

macrophylla, 297, 301, 307, 308,
309, 310, 311, 312, 313, 322, 361

INDEX

Vismia—Continued

macrophylla var. glabrescens, 307,
308

macrophylla sensu Benth., 310

magnoliifolia, 305, 306, 342, 347,
348

mandurr, 304, 305, 339, 340

martiana, 305, 342, 347, 348

mexicana, 293

micrantha, 302, 315, 316, 317,
318, 356

minutiflora, 302, 319

obtusa, 303, 332, 357, 358

panamensis, 304, 305, 323, 339

parviflora, 302, 318, 320

pentagyna, 305, 349, 350, 352,
353

plicatifolia, 303, 330

pozuzoensis, 329

ramuliflora, 314, 320, 321

377

Vismia—Continued

reichardtiana, 303, 306, 348, 349,
350, 352, 353, 356

reticulata, 310, 312, 361

rufa, 302, 314

rufescens, 320, 321, 349

rufescens var. sessilifolia, 320

rusbyi, 305, 350, 351, 352, 354,
361

sandwithii, 301, 309, 310, 312, 313

schomburgkiana, 361

sessilifolia, 306, 309, 310, 314, 320,
321, 345, 346

sieberiana, 361

sprucei, 302, 330, 358, 359

subcuneata, 337

tomentosa, 296, 304, 308, 342, 343,
344, 345, 357

urceolata, 304, 324

viridiflora, 306, 330, 346

U.S, GOVERNMENT PRINTING OFFICE: 1962

UNITED STATES NATIONAL MUSEUM

CONTRIBUTIONS FROM THE UNITED StTaTEs NATIONAL HERBARIUM

VoLuME 35, Part 6

CACAO AND ITS ALLIES
A TAXONOMIC REVISION OF THE
GENUS THEOBROMA

By Jose CuaTRECASAS

BULLETIN OF THE UNITED States NaTIoNAL MusEeuM
SMITHSONIAN INSTITUTION e WASHINGTON, D.C. e 1964

Contents

Page

Introduction. 2... 6 ee ee ee 379
Historical Sketch . 2 2 2 1 1 ke ee ee 383
Morphology . . 1-6 ee et et ee ee ee ee 415
Ecology . . . Lo ee ee ee ee ee ee 430
Geographic Distribution Lk ee ee ee ee ee ee ee 430
Pollination. 2... 0. 0. ee ee ee ee ee ee re 432
Relationships 2... 1 6 eee ee ee ee ee 432
Evolution . 20.0.0. 0. ee ee ee ee ee es 435
Economic Uses... . «6 ee ee ee et ee es wee ee 438
Anatomy of the Wood (Contributed by William L. Stern) Lee ee 439
Pollen Morphology (Contributed by G. Erdtman). . .... ++. 442
Cytology (Contributed by F. W. Cope). . . Loe ees 446
Pollen Incompatibility (Contributed by) F. W. - Cope). Loe es 447
Genus Theobroma. . . . . toe ke ke ee 449
Rejected names and excluded species a 584
Collections cited . 2 6. we ee 589
Bibliography. 2 1. 6 6 ee 599
Index... 1. 1 ee ee ee ee tee ee ee woe ee 607

III

CACAO AND ITS ALLIES

A TAXONOMIC REVISION OF THE GENUS THEOBROMA

Jose CUATRECASAS

Introduction

“Celebrem etiam per universam Americam multique usus
fructum Cacao appellatum.”
Cuiusivus, 1605.
“Cacao nomen barbarum, quo rejecto Theobroma dicta est
arbor, cum fructus basin sternat potioni delicatissimae,
saluberrimae, maxime nutrienti, chocolate mexicanis, Euro-
paeis quondam folis Magnatis propriae (Bpwya rwv Gewv, Vos
Deos feci dixit Deus de imperantibus), licet num_ vilior
facta.”
Linnagvs, Hort. Cliff. 379. 1737.

Theobroma, a genus of the family Sterculiaceae, is particularly
noteworthy because one of its members is the popular “‘cacao tree”’ or
“cocoa tree.’ The uses and cultivation of this outstanding tropical
plant were developed in the western hemisphere by the Mayas in
Central America a long time before Europeans arrived on the con-
tinent. The now universally used name cacao is derived directly
from the Nahuatl “cacahuatl” or “‘cacahoatl,” just as the name of the
popular drink, chocolate, is derived from ‘‘xocoatl” or ‘“‘chocoatl.”’
The economic importance of cacao has given rise to great activity in
several fields of development and research, especially in agronomy.
Historians and anthropologists have also been very much interested
in learning the role played by cacao in the economy and social
relations of the early American populations. There exists today an
extensive literature devoted to the many problems related to cacao.

I saw cacao for the first time in Colombia in 1932, but became
actually interested in the genus in 1939 and the years following, when
I found cacao trees growing wild in the rain forests of the Amazonian
basin. I was fascinated by the unique structure of the flowers of the
cocoa tree, and its extraordinary fruit. My explorations from 1942

379

380 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

to 1947 at the service of the Government of the Valle del Cauca,
mainly in the dense, humid forests of the Pacific coast of Colombia
and western slopes of the Andes, offered me the opportunity to
become better acquainted with wild species of Theobroma. I
published the descriptions and illustrations of four new species
found in that region, and at the same time gathered material for a
monograph of the genus. Subsequently in Chicago and Washington
I continued these studies, using the collections of the museums there
and loans received from important European herbaria. The steadfast
cooperation of V. M. Patiiio has been of great value to me in several
respects, and his explorations have furnished two new species.
Jorge Leén from Turrialba sent me specimens of an outstanding new
species. Collections sent to me for identification by the members of the
English Colombian Cocoa Expedition (1952-1953) very much helped
to broaden my knowledge of many of the species and their distribution.
In 1954, I had the opportunity to study the important Theobroma
collections at the British Museum (Natural History), London; the
Royal Botanic Gardens, Kew; and the Muséum d’Histoire Naturelle,
Paris, where many Theobroma types are preserved. In 1961 (July 11-
15) I was allowed to examine several European collections, at that
time on loan at the Harvard Botanical Museum, which I felt desirable
to see before publishing this revision. On my way to Colombia in
1961 for a general collecting trip, I also visited the living Theobroma
collection of the Imperial College of Agriculture in Trinidad where
some 14 species and several hybrids and varieties are cultivated. I
was thus able to supplement my data on the growing system and
fruits of some of the species I had not seen living before. For the
same purpose [ visited the cacao stations of the Interamerican
Institute of Agronomic Sciences at Turrialba, Costa Rica.

This publication is restricted to the taxonomy of the genus. Many
binomials (see the index) have been published in the past, and there
is great confusion with regard to the names in the herbaria and
literature. A critical revision was necessary in order to establish
the validity of the species and to list synonymy. Although a complete
study of the genus would require much more exploration, because
of the great gaps existing in the herbarium collections, the present
revision of all available materials seems justified. Three critical
treatments of the genus have previously been published; Bernoulli
(1869) recognized 18 species, and Schumann (1886) 11. Chevalier, in
1946, acknowledged 13 species and a few subspecies; he united some
species which should be held separate and on the other hand listed
as different others which are actually synonyms. Twenty-two species
are recognized in the present work. The number of species in the
genus will probably increase in the future, because new explorations

CUATRECASAS—CACAO AND ITS ALLIES 381

in Central as well as South America will undoubtedly bring about
the discovery of new ones.

This revision is based on the classical method of comparative
morphology. I have to a large extent used the structure of the fruit
and the vegetative characters which I found were basic features in
the definition and taxonomy of Theobroma. For a better under-
standing of the genus, some new concepts are also contributed by
G. Erdtman in pollen morphology (pp. 442-446), F. W. Cope in
cytology and incompatibility in cacao (pp. 446-449), and by W. L.
Stern in anatomy (pp. 439-442). These contributions may help in
the understanding of the taxonomic problems of Theobroma, espe-
cially those derived from cultivated varieties. In regard to 7. cacao,
the classification of the various cultivars is provisionally presented in
a conservative way; an understanding of the innumerable existing
forms of cacao will only be possible after long-term genetic research.
Dr. Cope and Dr. Bartley in Trinidad are working in this direction.
Dr. Soria in Turrialba, Costa Rica, is engaged at this time in an
ambitious project of this kind, largely supported by the American
Cocoa Research Institute of Washington.

Due to the nature of this paper, the historical sketch is limited to
the works which have contributed basic new data related to the
taxonomy of Theobroma. The relationships with other groups and
within the genus itself have so far as possible been treated objectively,
with few hypothetical speculations. The specific descriptions are
accompanied by original analytical drawings of the flowers, fruits, and
leaves of almost all the species, and the illustrations, carefully
supervised by the author, can be considered complementary to the
written descriptions. Because of the relatively small number of
existing collections, I think it useful to publish the information on
herbarium specimens given by collectors; these data, except in
special cases, have been translated into English whenever written
in another language. However, the numerous herbarium collections
of Theobroma cacao, mostly from cultivated plants, cannot be iden-
tified as to the variety and are therefore not included in the text but
simply listed in the index. For the citations of herbaria the abbre-
viations of Lanjouw’s “Index Herbariorum”’ have been used. The
abbreviation Photo F.M. is used to indicate the photographic series
of the Chicago Natural History Museum.

The artistic work for most of the illustrations has been carried out
by the artists Christopher Reinecke, Maria Luisa Biganzoli, and
Gil Cuatrecasas. Their work, consisting of about 35 plates, has
been sponsored by the American Cocoa Research Institute; a few
other plates had formerly been made by Gustavo Rojas, artist of the

382 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Comisién Botanica del Valle, Colombia, and by Paula Gerard,
Chicago.

This revision has been done in the Department of Botany of the
U.S. National Museum under a grant from the National Science
Foundation.

I wish to express my thanks to the directors and curators of the
herbaria for their aid in the loan of collections and granting of work
facilities, particularly to Dr. Albert C. Smith, Assistant Secretary of
the Smithsonian Institution; to staff members of the Museum of
Natural History, U.S. National Museum, who have checked the
manuscript: Dr. Jason R. Swallen, head curator of botany, Dr.
Lyman B, Smith, curator of phanerogams, and Conrad V. Morton,
curator of ferns; to Dr. E. P. Imle, director of Research of the
American Cocoa Research Institute; the Ministry of Agriculture of
Colombia; to Mr. N. Y. Sandwith of the Royal Botanic Gardens,
Kew, who helped on several occasions with important data and col-
laborated in the typification of a few species; as well as to Mr. J. E.
Dandy, of the British Museum, whose advice was of great help in
the typification of 7. cacao L. Likewise, I extend my thanks to
Dr. W. Robyns of Brussels, Dr. H. Melchior of Berlin, Prof. H.
Humbert, Prof. A. Aubreville, and Dr. A. Lourteig of the Museum
National d’Histoire Naturelle, Paris, Drs. J. Pablo Leyva and
A Fernandez Pérez of the Instituto de Ciencias N aturales, BogotA;
to my collaborators in Theobroma studies, Dr. Victor M. Manuel
Patifio, Ing. Humberto Guerrero, Dr. Ovidio Barros and Mr. Luis
Willard, all of Colombia, and Dr. L. Aristeguieta, of the Instituto
Botanico, Venezuela. I further thank Drs. Cope and Bartley of the
Imperial College of Trinidad both of whom were helpful in many
ways, and Drs. Leén and J. Soria of Turrialba, Costa Rica.

The collections used for this revision are the following:

Arnold Arboretum, Harvard University, Cambridge, Mass. (A).

Botanical Museum, Harvard University, Cambridge, Mass. (AMES).

Botanisches Museum, Willdenow Herbarium, Berlin (B).

Bailey Hortorium, Ithaca (BH).

British Museum (Natural History) London (BM).

Jardin Botanique de l’Etat, Bruxelles (BR).

Botanical Museum and Herbarium, Copenhagen (C).

Instituto de Ciencias Naturales, Bogota (COL).

Edinburgh Royal Botanic Garden (E).

Chicago Natural History Museum (F).

Conservatoire et Jardin Botaniques, Geneve (G).

Goteborg Botaniska Triidgird, Géteborg (GB).

Gray Herbarium, Harvard University, Cambridge, Mass. (GH).

University of Glasgow, Dept. of Botany (GL).

Systematisch-Geobotanisches Institut der Universitat Gottingen
(GOET).

_ CUATRECASAS—CACAO AND ITS ALLIES 383
Staatsinstitut fir Allgemeine Botanik, Hamburg (HBG).
Instituto Agronémico do Norte, Belem do Parad (IAN).
Royal Botanic Gardens, Kew, Surrey (K).
Rijksherbarium, Leiden (L).
Botanical Museum and Herbarium, Lund (LD).
Komarov Botanical Institute of the Academy of Sciences, Leningrad
(LE).
Botanische Staatssammlung, Munich (M).
Instituto de Biologia, México (MEXU).
Museu Goeldi de Historia Natural, Belem do Pardé (MG).
University of Michigan, University Herbarium, Ann Arbor (MICH).
Missouri Botanical Garden, St. Louis (MO).
New York Botanical Garden (NY).
Muséum d’Histoire Naturelle, Paris (P).
Naturhistoriska Riksmuseum, Stockholm (8).
Imperial College of Tropical Agriculture, Trinidad (TRIN).
Botanical Museum and Herbarium, Utrecht (U).
University of California, Berkeley (UC).
U.S. National Herbarium (US).
Facultad de Agronomfa del Valle, Cali, Colombia (VALLE).
Instituto Botdnico, Venezuela (VEN).
Naturhistorisches Museum, Wien (W).
School of Forestry, Yale University, New Haven, Conn, (Y).

Historical Sketch

1605: First citation of cacao in botanical literature, by Charles de
’Ecluse (Clusius) in chapter XXVIII, of his Exoticorum libri decem,
under the name Cacao fructus. It refers only to the fruit and gives
a poor illustration of cacao seeds. ‘‘Celebrem etiam per universam
Americam multique usus fructum Cacao appellatum.”

1623: K. Bauhin mentions for the first time in his books in his
chapter ‘“Amygdalus” the cocoa plant as “Amygdalis similis
Guatimalensis Avellana Mexicana cujus fructum indigenae Cacao
appellant,” etc. (Pinax Th. Bot. 442).

1630: The first prints of the Hernandez’s Rerum Medicarum Novae
Hispaniae Thesaurus appear in which are given descriptions of the
cocoa tree under its Mexican name cacahoaquahuitl and of four varieties
called guauhcacahoatl, mecacahoatl, xochicacahoatl and tlalcacahoatl
which are distinguished by the fruits diminishing in size from the first
to the last, presumably representing cultivars; the pods were called
cacahoacentli and the useful seeds cacahoatl; he also mentions quauh-
patachtli which undoubtedly refers to Theobroma bicolor. His illus-
tration of the first (fig. 223) is clearly cacao Criollo.

1658: W. Piso describes cocoa ‘De Arbore Cacavifera”’ and repeats
the varieties cited by Hernandez in a long article on cocoa and
chocolate. His drawing also represents the Criollo variety.

384 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

1688: John Ray, in his Historia Plantarum, gives much attention
to cacao and its products in chapter VIII under the title: Cacao Ger.
Cacao sive Cacavate Park. Cacao Americae sive Avellana Mexicana
J. B. Amygdalae similis Guatimalensis C.B. The Cacao Tree. He
explains that there are four kinds of cocoa trees and under the heading
of Cacava quahuitl describes the tree, fruits, and seeds which he
compares to almonds saying that, they are white before ripening and
red when fully ripe.

1696: Plukenet, in his Almagestum Botanicum, classifies cacao as
an almond, under the name Arvor cacavifera Americana, the fruits of
which (folliculi) contain some kind of almonds; in his plate 268,
fig. 3, a leafy branch with a cacao fruit of the Criollo variety is
represented.

1696: Sloane lists Cacao in his catalog of Jamaican plants, with a
long series of quotations from previous authors and travel writers.

1700: The first taxonomic statement is made by Tournefort pub-
lishing the genus Cacao: “Cacao est plantae genus auctore clariss.
Plumerio” with a single species “‘speciem unicam novi.” He gives a
short description and drawings sent to him by Plumier. They dis-
tinguish the 5 sepals, the strangulated petals, and a pistil surrounded
by a laciniate girdle (staminodes) which develops into a ridged and
pointed fruit filled with seeds.

1706: Sibilla Merian gives an illustration of Surinam cacao which
proves to represent clearly the Criollo type (26, t. 26).

1710: Ray in his Methodus Plantarum copies Tournefort’s descrip-
tion and data to define Cacao; there are no changes in the 1733 edition.

1725: Sloane publishes a long article on ‘The Cacao Tree” in his
Voyage to Jamaica (vol. 2) giving the following botanical description
(p. 15): “Out of the Body of the Tree, or Branch comes a very small
Flower, standing on a half Inch long Footstalk, it is made up of 5
Capsular Leaves, 5 crooked Petals, several Stamina, and a Stylus, of
a very pale Purple color, after which follows the Fruit, which when
ripe is as big as one’s Fist, bigger in the Middle than at the Ends,
which are pointed, it has some Swlei and Asperities on its Outside,
is for the most Part of a deep Purple colour, the Shell being about
Half a Crown’s thickness, and containing within it many Kernels
of an oval Shape, each of which is as big as a Pistachia Nut, having
a thin Membrane without which is a mucilaginous Substance in which
it lies. The Nuts themselves are made up of several parts like an
Ox’s Kidney, some Lines being visible on it before broken, and is
hollow within, its Pulp is oyly and bitterish to the Taste, made up of
many Striae, which tend from the Circumference to the Center.”
The plate 160 illustrates a leafy branch with oblong-ovoid fruits,
10-ridged and strongly pointed; there are separate drawings of flowers

CUATRECASAS—CACAO AND ITS ALLIES 385

showing more or less clearly 5 sepals, some petals and (not well defined)
4 or 5 staminode laciniae at the center, but no stamens; seeds, one
isolated, some others together covered with pulp are also illustrated.
The article gives much information about cultivation, varieties,
geographical distribution, and trade in cocoa, and many authors are
cited.

1737: Linnaeus (Genera Plantarum) introduces cacao into Classis
18 of his classification, in Polyadelphia pentandria giving it a new name
Theobroma—meaning “food for the Gods”; the name Cacao given by
Plumier and Tournefort was rejected by Linnaeus as “barbarous.”
By the International Code of Botanical Nomenclature, Linnaeus’
name prevails. The new genus Theobroma was published almost
at the same time in the Genera Plantarum and in Hortus Cliffortianus;
Linnaeus described the flowers as having 5 stamens, 5 petals and
staminodes (folioli nectarii), but only 3 sepals and with 5-celled
anthers instead of 4. Linnaeus probably used dried specimens
and annotations sent to him by Sloane for his description. He
included in Theobroma two species: one with “foliis integerrimis,”
Cacao, the other with “foliis serratis,’’ Guazwma, both also differen-
tiated by their fruits. In the Genera Plantarum, he gives as the
only synonyms and citations Tournefort for Cacao and Plumier for
Guazuma, but in the Hortus Cliffortianus he quotes for Cacao:
Clus., Raj., Tourn., Sloane, Mer., Hern., Pluk., and Bauh. But
Linnaeus found out by himself, with the help of Sloane, the num-
ber and kind of stamens typical of 7. cacao. He writes in Hortus
Cliffortianus: ‘Flores a nullo bene depicti, multo minus descripti
sunt,” and then: “Sloane mihi inspiciendi copiam fecit, videbatur
structura exacte sequentis, ab aliis in universum omnibus diversis-
sima.” His original description of the stamens, given in the Genera
Plantarum (“Filamenta subulata, longitudine nectarii, cui radiorum
instar innata: singula apice quinquefida. Antheris in singulo stamine
quinque, tectis petalo concavo”’), was made on the basis of drawings
or flowers sent to him by Sloane, for which reason the flowers of the
Sloane herbarium have to be considered as the type of Linnaeus’
description. But Linnaeus may have had very scanty material of
the flowers, because he described the anthers as 5-celled instead of
4-celled.

1739: Weinmann writes extensively about cacao and chocolate and
gives a plate (277) which is inspired by Tournefort’s illustration
using very much imagination in painting it. Plate 278, devoted to
Cacao minor, depicts a very deformed kind of pod.

1739: Elizabeth Blackwell depicted cacao (pl. 373) using Miller’s
specimens and suggestions. The fruit is figured as elongate, pointed

386 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

and 10-ridged, although slightly verrucose; it represents the Criollo
type.

17 : Geoffroy in his De Vegetabilibus Exoticis has a long article
(XIX) De Cacao giving detailed descriptions of the tree, fruits, seeds,
and their preparations. He calls the attention to the variability of
the species in size or thickness of the different organs (especially the
leaves and fruits).

1747: Catesby publishes a magnificent colored plate of the cocoa
tree in his Appendix to the Natural History of Carolina, ete. (p.
6, pl. 6); the buds are colored red, the petals yellow, the staminodes
red, and the fruits orange, obovoid-oblong, 10-ridged and warted, and
very pointed ; clearly it is the Criollo type. There is a long description
of the tree with observations taken from Dampier; of the fruit it is
said: “Fruit about the bigness of a swan’s egg, but longer, more
tapering, and ending in a point. The fruit hangs pendant, and when
ripe, has a shell of a purple color, in substance somewhat like that of a
pomegranate, and furrowed from end to end, containing in the middle
many kernels of the size of acorns, inclosed in a mucilaginous sub-
stance...” In a later edition (1771) the color of the fruit was
changed to dark violet.

1749; Linnaeus, in his Materia Medica, includes Theobroma foliis
integerrimis (p. 364) with the previous definition and classifica-
tion; he attributes to it the qualities and virtues of pinguis, subamara,
nutriens, aphrodisiaca, calefaciens.

1753: Linnaeus gives a binomial name to cacao in his first edition
of the Species Plantarum (p. 782): Theobroma Cacao, with the short
specific diagnosis “‘folizs integerrimis.”” He also names a second spe-
cies Theobroma Guazuma with the diagnosis “foli’s serratis.’” To the
bibliographical citations are added: “Mat. Med. 364,” “Geoffr. Mat.
409,” and ‘‘Catesb. car. 3. p. 6. t. 6.” Inasmuch as the first edition
of Linnaeus’ Species Plantarum is the official beginning publication
date for phanerogams by the Code of Nomenclature, Theobroma
cacao receives here its official nomenclatural start, the generic name
being in accordance with the diagnosis given in the corresponding
edition (fifth) of Linnaeus’ Genera Plantarum.

1754: Linnaeus, in the fifth edition of Genera Plantarum, defines
Theobroma and gives it the same classification (Polyadelphia pen-
tandria) as in the first edition. Linnaeus did not improve his knowl-
edge of the genus nor change his concepts of it in later works. In
the third edition of Species Plantarum (1764), and also in the twelfth
edition of Systema Naturae (2: 508. 1767) he keeps the same treatment
of Theobroma as in his earlier publications.

1754: The generic name Cacao is validated, according to the present
Code of Nomenclature, by Miller in his fourth abridged edition of

CUATRECASAS—CACAO AND ITS ALLIES 387

“The Gardeners Dictionary.’”’ As in his popular sixth (1752) and eighth
(1768) and other later editions of the Gardeners Dictionary, Miller
publishes, without or with slight variations, a long article on “Cacao,”
“The Chocolate Nut.” He explains that “This genus of plants was
constituted by Father Plumier, who communicated the characters,
which he had drawn in America, to Dr. Tournefort, who has inserted
it in the Appendix of his Institutions. Dr. Linnaeus has joined this
to the Guazuma of Plumier, under the title of Theobroma, but as the
fruits of these plants are very different from each other, I shall keep
them under different genera. We have but one species of this plant,
which is Cacao.”

1768: Adanson separates Cacao (Theobroma) from Guazuma, but,
while the latter is kept in the family “Les Tilleuls,’’ Cacao is placed in
the family “Les Pistachiers,” side by side with Diosma, Triopteris,
Acaju, Hugonia, etc., far away from its true relationships.

1765: A disciple of Linnaeus, Antonius Hoffmann, presents to the
Swedish Royal College of Medicine the first doctoral thesis ever pro-
posed dealing with a Theobroma subject, ‘““Potus Chocolatae.” It is an
excellent review of the knowledge about the composition and ways to
prepare the chocolate at that time and the nutritional and medical
importance of it. Hoffmann gives a more detailed description of the
cacao plant and the fruit than did Linnaeus, probably inspired in
Geoffroy; he writes: “Fructus magnitudinem et figuram refert Melonis,
sed verrucosus est, decem angulis instructus et superne acuminatus;
dum maturescit, fit colore coccineus atque maculis variegatus flavis;
intus continet nucleos circiter triginta, qui magnitudine Olivas aemul-
antur ac pulpa obteguntur albida, subdulci et amaricante. Olei
magna scatent hic nuclei copia, quod expressum vocatur Pinguedo de
Cacao.” (1769, p. 257.)

1775: A disciple of Linnaeus, Jacobus Alm, in his doctoral disserta-
tion Plantae Surinamenses republished ten years later in Amoenitates
Academicae, emends the Linnaean description of the cacao flowers.
He recognizes sterile stamens (‘stamina alia 5, castrata”) in what
Linnaeus called “nectaria,” alternate with the other ‘‘stamina fertilia
solitaria’”’; he corrects Linnaeus also in seeing the calyx ‘“‘pentaphyllus”’
and the anthers “‘quadriplici.”

1776: Aublet, in his explorations in French Guiana, found
wild species of Theobroma, which he describes extensively in his
Histoire des Plantes de la Guiane. He names them Cacao guianensis
(pl. 275) and Cacao sylvestris (pl. 276); he quotes “cacao” as a Carib-
bean name used. To the cultivated cacao, which he also found, he
gives the new name Cacao sativa. It is unfortunate that Aublet mixed
up elements of three species in describing his two new species; Cacao
sylvestris was pictured from concordant parts of foliage and fruits

388 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

which agree well with the species at present known as 7’. subincanum,
but the flowers mentioned in his description were from TJ. cacao.
Cacao guianensis was described and depicted from branches and
foliage identical to Cacao sylvestris (=T. subincanum) and its flowers
were taken from specimens of 7’. cacao; the fruit is the only different
part, belonging to a third species. Aublet’s descriptions and drawings
are detailed, this being the first publication giving an accurate idea of
the cacao flowers and of the fruits of some wild Theobroma.

1785: Lamarck, in his famous Encyclopédie Méthodique, lists
Cacao as belonging to the family of Cacaoyers characterized by
hermaphrodite and complete flowers with 5 petals, 5 or 10 stamens
and superior, usually 5-celled ovary; other genera of the family were:
Ambroma, Guazuma, Ayenia, Buttneria, and Kleinhovia. He points
out its close relation to Hermannia, Tilia, and the Malvaceae. He
describes three species: Cacao sativa, C. sylvestris, and C. guianensis
with the remarks mostly taken from Aublet.

1789: A. L. Jussieu (Genera Plantarum) places Theobroma in Classis
XIII, Ordo XIV (Malvaceae), of his system including it in his section
V, (bis!) characterized by mixed fertile and sterile stamens connate
at base; here Theobroma is associated with Pentapetes L., Abroma Jacq.,
Guazuma Plum., Melhania Forsk., Dombeya Cav., Assonia Cav.,
and Byttneria L.

1791: Gaertner, in his remarkable book on fruits and seeds, gives a
good description of cacao and drawings of its fruits and seeds, which
represents a Criollo form; undoubtedly, Gaertner had at hand a dried
specimen, for the section of the pod is drawn relatively thin and he
describes it with ‘cortex sublignosus.” The specific name used by
Gaertner, Cacao minus, is a nomenclatural synonym of Theobroma
cacao; Gaertner mentions Theobroma foliis integerrimis Linn., besides
Sloane and Blackwell.

1791: Schreber, in the eighth edition of Linnaeus’ Genera
Plantarum, divides Classis XVIII in two subclassis, Decandria and
Dodecandria. He places Theobroma in Decandria, a new concept,
while Bubroma (a new name for Guazuma) and Abroma are brought
into the Dodecandria. Schreber points out as differences between
Theobroma and Bubroma, that the first has “laminae subrotundae
acuminatae” and “antherae in singulo filamento duae,’”’ while Bubroma
has “laminae semibifidae,” ‘“antherae in singulo filamento tres,”
and ‘“‘capsula non dehiscens muricata.”

1791: Gmelin, in the 13th edition of Linnaeus’ Systema Naturae
(vol. 2, p. 1151), includes Theobroma in Polyadelphia Decandria to-
gether with the genus Abroma. He still lists 7. Cacao and T. Guazuma
as species of Theobroma, adding another: Theobroma guianense, a
new combination based on Aublet’s Cacao guianensis.

CUATRECASAS—CACAO AND ITS ALLIES 389

1796: Salisbury publishes Theobroma celtifolia, which is a synonym
of Guazuma ulmifolia.

1796: Lamarck (Tableau Encyclopédique) publishes illustrations
of Theobroma Cacao, fig. 1 representing flowers, copied from Aublet’s
plate, fig. 2 the smaller form of 7. cacao illustrated by Gaertner, and
fig. 8 a correct drawing of flowering and fruiting branches of a Criollo
cacao (not J. guianense as stated).

1802: Willdenow, in his edition of Linnaeus’ Species Plantarum,
follows the treatment of Schreber, including Theobroma in Poly-
adelphia Decandria and Bubroma and Abroma in Dodecandria. Still
only two species are considered: 7. cacao and T. gujanensis.

1806: Humboldt and Bonpland, in Plantae Aequinoctiales, a mag-
nificent work, publish the first perfect botanical description of a species
of Theobroma. It is supplemented with two plates illustrating leafy
and flowering branches, fruits, seeds, and flowers of Theobroma bicolor,
found by the authors cultivated in Colombia. The drawings show
details of the embryo; the staminodes are wrongly figured as pointed
instead of obtuse.

1808: De Tussac, in Flora Antillarum, writes extensively on the
cacao tree, its cultivation and uses, under the heading Cacao Theo-
broma, giving “Le Cacaoyer Theobrome” as the French and ‘The
Chocolate Tree” as the English name. There is a plate in folium
(pl. XIII) showing illustrations of orange yellowish, 10-ridged fruits,
attenuate at both ends, foliage, floral details, and the embryo. De
Tussac is so enthusiastic about the use of chocolate that he says
“Je chocolat est au corps, ce que le café est a l’ésprit’”’ (p. 103).

1811: Poiret, in the supplement to Lamarck’s Encyclopédie, makes
the new binomial Cacao bicolor, a combination based on the Humboldt
and Bonpland species.

1812: Stokes, in his Botanical Materia Medica, publishes a new
name, Theobroma integerrima, for T. cacao L.

1823: Kunth, in Humboldt, Bonpland, and Kunth, Nova
Genera et Species Plantarum, gives a good description of the genus
Theobroma and of the two species TJ. cacao and T. bicolor; the descrip-
tion of the androecium is entirely correct: ‘‘Filamenta 10, basi in
urceolum connata; quinque petalis opposita dianthera; quinque al-
terna sterilia, lineari subulata. Antherae didymae biloculares, in
petalorum cavitatae reconditae.” Kunth gives a good description
for the family Biittneriaceae R. Brown and for the five sections in
which it is divided (Sterculiaceae, Biittneriaceae verae, Lasiopetaleae,
Hermanniaceae and Dombeyaceae). Theobroma Gointly with Gua-
zuma, Abroma, Glossostemon, Biittneria, Ayenia, and Commersonia)
are placed in the Biittneriaceae verae which have the stamens ‘10-30.
Filamenta magis minusve connata; quinque, laciniis calycinis opposita,

390 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

antheris destituta, alius formae. Antherae didymae, longitudinaliter
dehiscentes.”’

1824: De Candolle, in his Prodromus, accepts also family status
for the Byttneriaceae (as Ordo XXVI) and follows the classification
of Kunth, except for calling the sections divisions. In the tribe
Byttneriaceae he includes the same seven genera, plus another doubt-
fulone. In his treatment of Theobroma he includes five species, three
already known, T. cacao, T. guianensis, and T. bicolor and two new
ones, taken from the Mexican flora of Mocifio and Sessé: 7. angus-
tifolia and T. ovatifolia. Mociiio and Sessé had written an unpub-
lished Flora of Mexico supplemented by a series of illustrations, which
were copied at Geneva by artists hired by De Candolle.

1826: Sprengel, in the sixteenth edition of Linnaeus’ Systema Vege-
tabilium, keeps Theobroma in Polyadelphia (Classis XVIII) and
divides it in three genera: Theobroma with 2-antheriferous stamens
and Bubroma and Abroma with 3-antheriferous stamens. He lists
five species in Theobroma: T. cacao, T. bicolor Humb., T. speciosum
W. herb., 7. ovatifolium Sess., and T. guianense W. The genus
Bubroma, considered synonymous with Guazuma Poir., comprises
five species: B. Guazuma, B. tomentosum, B. polybotryon W., B. grandi-
Jlorum W. herb., and B. Invira W. Of these B. grandiflorum is
actually a Theobroma. In Abroma, Sprengel included two species:
A. augustum L. suppl., and A. fastwosum Salisb. New Theobroma
species in this publication are 7. speciosum and B. grandiflorum.

1827: Descourtilz, in his picturesque Flore Médicale des Antilles,
dedicates much space to the description of the cacao tree “Cacaoyer
cultivé,”” to its origin, cultivation, varieties, uses, etc. Plate 266
represents a leafy branch with flowers, seeds, and a fruit which is of
the Criollo type (10-ridged, very warty, and acute).

1828: Voigt publishes a description of a Theobroma guianensis
without mention of Cacao guianensis Aublet (‘‘fol. acuminatis cor-
datis sublobatis inaequaliter eroso-dentatis, subtus tomentosis, ramis
petiolisque ferrugineo-hirtis, corymbo terminali. Flores albi, parvi).
According to some of the features given (corymbo terminali, flores
albi), the plant described does not belong to Theobroma.

1530: Sweet, in Hortus Brittanicus, listed four species under
Theobroma mentioning a new binomial, 7. caribaea, with no descrip-
tion. It is undoubtedly a name for a form 7’. cacao.

1830: Martius, who observed and collected many Theobromas
on his trips throughout Brazil, says that more species may be found
growing wild in tropical forests; he gives an account with short de-
scriptions of the species found by him in Brazil, of which three are
new species: Theobroma subincanum Mart., T. sylvestre Mart., and

CUATRECASAS—CACAO AND ITS ALLIES 391

T. microcarpum Mart.; the other species listed are T. cacao L. (=T.
sativum Lam.), T. speciosum Willd. ?, and 7. bieolor H. et B.

1831: Don, in A General History of the Dichlamydeous Plants,
places Theobroma in the family Byttneriaceae, tribe Byttnerieae
DC. He gives good descriptions of these groups and of Theobroma,
and short definitions for six species with some special attention to T.
cacao. The other species listed are: 7. guianensis, T. bicolor, T.
angustifolia, T. ovatifolia, and T. sylvestris, the last being a new com-
bination for Cacao sylvestris Aubl.

1831: Martius, in his Reise in Brasilien (p. 1127), explains that
the cocoa from Para and Rio Negro is of a lower, more bitter quality,
because it comes more often from wild cacao trees than from culti-
vated trees. He also says that he found 7’. bicolor growing wild in
Barra do Rio Negro, in Manacurt, and Yapura.

1840: Endlicher, in his Genera Plantarum, gives excellent descrip-
tions for the Ordo CCXI Biittneriaccae and its six tribes, two of them
being new: Eriolaenae and Philippodendrae; the other four are the
same as those of Kunth and De Candolle, except for the Sterculiaceae
which are treated as an order apart united with Bombacaceae and
Helicteraceae. He includes in Biittnerieae DC. the genera Rulingia,
Commersonia, Abroma, Biittneria, Ayenia, Theobroma, and Guazuma.
The genus Glossostemon is placed in the Dombeyaceae.

1847: Dietrich, in his Synopsis Plantarum, describes briefly nine
species of Theobroma and gives a new name, Theobroma Martiana, for
T. sylvestris Mart.

1856: Karsten describes Theobroma glaucum, a new species from
eastern Colombia.

1861-1870: Baillon studies the development of the parts of the
flower in 7. cacao. The primordia of the sepals appear successively
one after another, emerging above a common basal annulus. The
petal primordia appear simultaneously and have the same aspect as in
flowers of most other plants, but when they develop, there appears a
strangulation which divides them into two articulated parts; the
basilar parts are valvate, the upper parts contorted. The staminode
primordia, opposite to the sepals, appear before those of the fertile
stamens, which are opposite to the petals, and develop a simple fila-
ment which divides into two, each branch having an anther whose
two thecae become superimposed. The five primordia of the carpels
opposite to petals have a half-moon shape; they become connate and
develop alternate growths at the joints which are the primordia of the
walls; these are centripetal and progressively divide the ovary into
five cavities. Primordially, the placentation is parietal, becoming
axile when the carpelar walls reach the axis; the ovules develop two
integuments, become anatropous and placed in two rows in each

680-695—64——-2

392 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

cavity, with the raphe, outside facing each other, horizontal. The
embryo at first has ovate-orbicular entire, flat cotyledons; later these
develop folds becoming corrugated. The initial transparent albumen
is absorbed and at the end disappears.

1862: Triana and Planchon, in Prodromus Florae Novogranatensis,
list three species of Theobroma for Colombia (7. cacao, T. bicolor, and
T. glauca) and three species of Herrania (H. pulcherrima, H. albiflora,
and H, lacinizfolia). T. cacao is only given as cultivated.

1862: Bentham and Hooker (Genera Plantarum 1: ix, 216), following
basically the lines of De Candolle, although using other terms, classify
Theobroma in the series Thalamiflorae, cohors VI Malvales, ordo
XXXIT Sterculiaceae and tribus VI Buettnerieae. The chief charac-
ters for the tribe are the hermaphrodite flowers, concave petal base,
and anthers 1-3 alternating with staminodes; it is divided into two
groups: *) the fertile stamens with 2-0 anthers (Glossostemon,
Abroma, Theobroma, Herrania, Guazuma), **) the fertile stamens with
a single anther (Ayenia, Buettneria, Rulingia, Commersonia). With
the main characters of most of the genera already known, good defining
descriptions are given for each of them.

1869: Bernoulli attempts the first monograph of Theobroma.
After a long time of field study in Central America and using the
herbarium collections preserved at Berlin, Kew, and Munich, Ber-
noulli became so well acquainted with the genus that he was able
to draw a very good natural classification of it. The five sections
established by him, based on flower and fruit characters, may be
entirely kept today, strongly reinforced by other more recently known
characters. When he started the study, there were only four or five
species known besides T. cacao; and he writes, ‘Actually nobody knew
these species, because the extremely short diagnoses of the old species
were completely insufficient to determine them, for which reason there
was the greatest confusion in the nomenclature in the herbaria that
I had the opportunity to see.” Bernoulli found that flowers and
fruits give constant characters, whereas the leaves only give some
secondary characters, especially the basal nervation; he also noticed
that in some species there is great variation in the shape and pubes-
cence of the leaves, with all transitions from one to another form
when abundant material is compared. The basic characters used
by Bernoulli for his system are: petal appendix or ligula sessile,
subsessile or stipitate; staminodes erect or reflexed in bud, subulate,
claviform, or petaloid; stamens 2-antheriferous or 3-antheriferous;
fruit, when it was known; calyx 5- or 3-parted with narrow or broader
lobes. With combinations of these features the following five sections
(the descriptions here abridged) result.

CUATRECASAS—CACAO AND ITS ALLIES 393

1. Cacao: Petal ligule stipitate; staminodes erect, subulate; stamens 2-
antheriferous; calyx 5-parted, the laciniae equal. Fruit ovate-
oblong.

2. Oreanthes: Petal appendix subsessile; staminodes erect, subulate; sta-
mens 3-antheriferous; calyx 5-parted, the laciniae equal.

3. Rhytidocarpus: Petal appendix subsessile; staminodes erect, claviform ;
stamens 2-antheriferous; calyx 5-parted, the laciniae equal; fruit
woody.

4. Telmatocarpus: Petal ?; staminodes erect ?, linear-subulate with broad
base; stamens 3-antheriferous; calyx 5-parted, the laciniae equal;
fruit ovate, lacunose.

5. Glossopetalum: Petal ligule stipitate; staminodes reflexed, petaloid;
stamens 3-antheriferous; calyx irregularly 3-5-fid, ‘‘foliaceous’’;
fruit sublignose.

Bernoulli describes 18 species, some of them extensively, others
very briefly, according to the material he had at his disposal. Of
these, 12 species are described as new. In sectio Cacao he includes 4
species: 7. cacao L. and 3 new ones: 7. pentagona, “cacao lagarto”
from Guatemala; J. leiocarpa, ‘“cumacaco” from cultivation in
Guatemala; and J. saltzmanniana from Bahia, Brazil. In sectio
Oreanthes he includes T. speciosa Willd. ex Spreng., and 2 new species:
T. quinquenervia and T. spruceana, both from Brazil. Sectio Rhytido-
carpus with one species, 7’. bicolor Humb. & Bonpl. (synonym, 7.
ovatifolia DC.), and, as doubtful, also 7. glauca Karst. In sectio
Telmatocarpus there is a single species, 7. microcarpa Mart. In the
sectio Glossopetalum, the largest, he describes in some detail 7.
angustifolia DC., T. subincana Mart., T. sylvestris Mart., and as new
species 7. macrantha from Brazil, T. ferruginea from Peru, T’. obovata
from Brazil, 7’. alba from British Guiana, and 7. nitida from Brazil.
About “Cacao guyanensis” Aublet, he writes “bleibt somit eine
vollstandig ungewisse Art. Sie scheint auch von keinem weiteren
Autor gesehen worden zu sein, sondern immer nur nach Aublet citiert
zu Werden.” Bernoulli quoted Herrania as a genus differing from
Theobroma in the habit of the plant and in the 5-6-foliolate digitate
leaves. Bernoulli’s work is illustrated with several drawings. ‘This
monograph was published by the Swiss Society for Natural Sciences
in its new serial of Memoires, vol. 24, no. 3, in 1871. However, a
reprint was issued previously in pamphlet form in 1869, which is
the effective date of publication. (See Pritzel, 1872; Sargent, 1912).
C. G. Bernoulli was born in Basel, Switzerland, Jan. 24, 1834, and
died in San Francisco, Calif., while returning home, May 18, 1878;
he lived in Guatemala from 1858 until 1878 and collected plants and
animals extensively in Central America.

1873: Baillon publishes excellent comparative descriptions of the
genus based on T. cacao in his Histoire des Plantes. He includes it
in the family Malvaceae which includes the Sterculiaceae and Bomba-

394 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

caceae; the family is divided into twelve series, the sixth being the
Buettnerveae with 12 genera: Buetineria, Ayenia, Commersonia,
Rulingia, Theobroma, Herrania, Guazuma, Scaphopetalum, Lepto-
nychia, Abroma, Mazxwellia, and Glossostemon. The main characters
given for the tribe are the usually hooded petal base, the fertile
stamens opposite the petals, the staminodes alternate with the petals,
the bilocular anthers (rarely 3-locular), the plurilocular ovary, and
the capsular or carnose fruit.

1882: D. Morris of Jamaica, in his well-known book on cultivation
of cacao, after describing the cacao plant and fruit, publishes the
first known classification of its varieties based upon the nomenclature
of some of the best estates of Trinidad. He distinguishes two great
classes (p. 12), the second being divided into several varieties:

Class I. Cacao Criollo (Red).

Class II. Cacao Forastero.

var, a. Cundeamor verugoso amarillo (yellow), (rough yellow

Cerasee)
. Cundeamor verugoso colorado (red), (rough red Cerasee)
Liso amarillo (yellow), (smooth yellow)
. Liso colorado (red), (smooth red)
Amelonado amarillo (yellow), (yellow Melon)
Amelonado colorado (red), (red Melon)

. Calabacillo amarillo (yellow), (yellow Calabash)
. Calabacillo colorado (red), (red Calabash)

re hs aae

This is the first valid publication in the nomenclature of the cultivars
of Theobroma cacao. Morris goes on to say that before the “blast”
or plague that almost exterminated cacao plantations in Jamaica
before the end of the seventeenth century, the Criollo class was the
only kind cultivated there. Since then, the Criollo has been entirely
discarded for the hardier Forastero, the Criollo being “now chiefly
confined to the mainland (Venezuela) where its yield, though small,
is considered of great value” (p. 13). This assertion determines that
the nomenclatural type of the Criollo cultivar is the Venezuelan
Criollo. Of the Forastero varieties the best are the Cundeamor, the
yellow kind being preferred for yielding a larger proportion of seeds.
The seeds of Cundeamor “are mostly of the true almond shape—
large, plump and full, of a pale crimson colour in the interior, and
ferment easily.” The Liso variety is closely allied to the former.
The Amelonado is intermediate between Cundeamor and Calaba-
cillo, and is considered as of good quality. The Calabacillo is the
lowest quality and, Morris says, “never cultivated by a judicious
planter; its fruits are small, the seeds flat, angular, intensely bitter
and of dark crimson colour.”

1556; Schumann, in his Vergleichende Bliithenmorphologie, makes
an accurate morphological analysis of eleven genera of Biittnerieae,

CUATRECASAS—CACAO AND ITS ALLIES 395

especially of Biittneria, Ayenia, Commersonia, Rulingia, Guazuma,
Theobroma, and Abroma, and tries to figure out the existing relation-
ships between them and other Sterculiaceae genera. He concludes
that the genera treated have a very close relationship and constitute
a very natural group. Schumann sees in the Sterculiaceae two series,
one of which is the main series (Hauptreihe), with a floral diagram
C;, P;, Std;, A;, G;, where P.A.G. are opposite; these elements are
either normally developed or modified by division or abortion; to
this series belong as the leading tribe, the Biittnerieae, followed by
Sterculieae, Helictereae, Lasiopetaleae and part of Hermannieae
(Melochia, Dicarpidium, Waltheria). The other division of the Ster-
culiaceae differs in having a floral diagram with the carpels opposite
the sepals, this includes the Dombeyeae and Hermannia. He sees
very close relationships between Biittnerieae and Lasiopetalae, which
have to be artificially separated, and also between Biitineria and
Ayenia, between Theobroma and Guazuma, and between the last and
Scaphopetalum and Leptonychia. There is no direct relationship
between the two series of the Sterculiaceae, the Dombeyeae being
close to the Malvaceae. Schumann does not see a way to explain this
and decides not to draw any evolutionary hypotheses.

Schumann describes the calyx of Theobroma as pentamerous or
with three divisions, the sepals often concave and slightly cucullate,
the petals sessile or shortly unguiculate. The inner glands of the
receptacle are pluricellular and stipitate-globose or flagelliform;
their function is unknown. The staminodes are more or less carnose.
The ovules are in 2 rows in the ovary cells, but the seeds become
uniseriate. The pulp of the seeds is mucilaginous and includes fine,
curled, spiralish fibres.

1886: Schumann, in the Flora Brasiliensis, gives a synopsis of
Theobroma, describing with detail and accuracy its recorded species.
But he does not follow the well-based system of Bernoulli. Schumann
includes the genus Herrania in Theobroma as section Herrania, leaving
the other species as section Kutheobroma. He uses the shape of the
petal lamina and the staminodes and the number of anthers in a single
stamen as secondary characters to classify the species using as the
primary character the many-flowered or few-flowered inflorescence, a
very vague feature in many instances. The keyed species in the
first group are T. Cacao, T. bicolor, and T. speciosum, and in the
second group, LT. microcarpum, T. grandiflorum, T. subincanum, and
T. angustifolium. Four species are listed outside of the key as
“dubiae”’: 7. glaucum Karst., T. silvestre Mart., T. Martii Schum.,
and 7’. album Bern. Two excellent illustrations are given for 7. Cacao
and 7. grandiflorum. Schumann reduces to 7. Cacao three Bernoulli
species, 7’. leiocarpum, T. pentagonum, and T. salzmannianum, which

396 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

he thinks were based on insignificant variations of fruits or petals
of this polymorphic species. Also he considers the flowers of Cacao
guianensis Aublet as being 7. Cacao, whereas the foliage and fruit is
possibly 7. subincanum. Bernoulli’s 7. quinquenervium and T.
spruceanum are reduced by Schumann to varieties of 7. spectosum.
Schumann transfers Bubroma grandiflorum Willd. to Theobroma, citing
as a synonym 7. macranthum Bern. T. obovatum Kl. ex Bern. was
not well known to Schumann, who makes it a synonym of T. subin-
canum. In short, the 18 species listed by Bernoulli are reduced by
Schumann to 11, four of these being dubious to him.

Schumann places Theobroma in the family Sterculiaceae, tribe IV
Biittnerieae, characterized mainly by cucullate petals. This is divided
into two subtribes, corresponding exactly to the two groups made by
Bentham and Hooker, but Schumann gives them names: a) Theo-
brominae, defined as having 10 or 15 stamens and the base of petals
cymbiform, and b) Biittnerinae, distinguished by having 5 stamens
and the petal cucullus incurved at apex. TZheobroma is the principal
genus of the first, differing from Guazuma by its baccate fruit, lack of
perisperm, and scarce, mucilaginous endosperm; Guazuma has lignose
fruits, a developed perisperm, and lacks endosperm.

1890: Schumann in his treatment of the Sterculiaceae for Engler
and Prantl’s Die Natiirlichen Pflanzenfamilien divides Theobroma into
three sections: 1) Herrania, 2) Eutheobroma (2-antheriferous stamens)
with 7. cacao and T. bicolor, and 3) Bubroma (3-antheriferous stamens)
with T. angustifolium, T. ovatifolium, T. grandiflorum, and T. subin-
canum. There are good illustrations. Schumann divides the family
into eight tribes, Theobroma belonging to the fifth, Biittnerieae,
characterized by hooded petals. It is divided into subtribe Biittneri-
nae with single stamens and Theobrominae with bundled stamens;
the first includes Rulingia, Commersonia, Biittneria and Ayenia, and
the second Glossostemon, Scaphopetalum, Leptonychia, Abroma, Theo-
broma, and Guazuma.

1890: Gémez de la Maza publishes Theobroma tomentosa, based on
Guazuma tomentosa.

1892: Hart modifies the Morris classification of the varieties of
T. cacao, removing Calabacillo from the Forastero group and making
with it a third class, as follows:

Class I. Criollo or fine thin-skinned.
1. var. a. Amarillo
2. var. b. Colorado
Class II. Forastero or thick-skinned cacao.
3. var. a. Cundeamor verugosa amarillo.
4. var. b. Cundeamor verugosa colorado.

5. var. c. Ordinary amarillo.
6. var. d. Ordinary colorado.

CUATRECASAS—CACAO AND ITS ALLIES 397

7. var. e. Amelonado amarillo.
8. var. f. Amelonado colorado.
Class III. Calabacillo, or small-podded, thick, smooth-skinned, flat-beaned.
9. var. a. Amarillo.
10. var. 6. Colorado.

Hart makes extensive and sound comments on the characteristics
and qualities of each variety, saying, among other considerations, that
“The finest cacao is by general consent admitted to be produced by
the Criollo variety, and this is assumed to be identical or similar in
character to that called the Caracas variety.” (pag. 48). ‘The
characteristics of the Criollo cacao are the thinness of its pod, its
rounded beans and pale colour of the interior of the bean on section.
The leaves of the tree are small when compared with the Forastero
varieties, and the tree itself is not nearly so sturdy and thriving, and
does not produce such regular and abundant crops as the Forastero
and Calabacillo varieties. The skin of the bean itself is thinner, and
the interior has but a small proportion of that bitter flavour which is
characteristic of the unfermented bean of Forastero and especially
that of Calabacillo. The flattest beans are those produced by pods
of the Calabacillo type. The beans of Forastero are intermediate
between these and the rounded form of the Criollo.” (p. 51). Hart
illustrates his important pioneer work with not too good illustrations
of pods and its sections of Amelonado, Calabacillo, Forastero and
Criollo, and with three diagrammatic sections of typical seeds of Criollo,
Forastero and Calabacillo. He adds, however, that there will be
found intermediate forms hardly reconcilable with any of the figures,
so that ‘“‘these are to be taken as representative only of the typical
varieties with some latitude.”

Hart’s illustrations are not representative of the concepts for which
the same nomenclature is generally used outside Trinidad: For
instance, the figure given as Criollo represents the type Cundeamor,
and the models he used to illustrate Calabacillo were not well selected.
These facts explain the comments made some years later by Preuss
about Hart’s somewhat confusing nomenclatural concepts.

1898: John Donnell Smith describes a new Theobroma with yellow
flowers and smooth cylindrical pods which Pittier and Tonduz dis-
covered in the mountains of Costa Rica, T. stmiarum.

1899: Jumelle publishes an excellent monographic compilation on
cacao with a long chapter devoted to the botany following an inter-
esting account of its history. Disregarding the work of Bernoulli,
Jumelle follows Schumann in his treatment of the Sterculiaceae in
the Flora Brasiliensis, declaring that “‘C’est certainement l'étude la
plus compléte et la plus consciencieuse qui ait été faite sur les Theo-
broma.’”’ Then he gives in French Schumann’s complete key for seven

398 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

species. He recognizes that several species*have been published
before, based on weak characters, probably due to variations caused
by cultivation or changes of soil or climate. Jumelle devotes much
attention to the description of 7. cacao and its different varieties.
He follows in this the works, experiences, and classification of Hart.
Chapters with descriptions and accounts are also devoted to T. bicolor
(and separately 7. ovatifolium), T. angustifolium, T. subincanum,
T. grandiflorum, T. speciosum, T. miecrocarpum, and T. glaucum.
Short notes are given to JT. sylvestre Mart., T. Martii Schum., and
T. album Bernoulli. Although excellent as a compilation, Jumelle’s
publication offers nothing original with regard to the systematics of
Theobroma; its many illustrations were taken from Bernoulli,
Schumann, and Hart.

1899; De Wildeman publishes a new binomial, Theobroma Kalagua,
with a description and illustrations of leaves, flowers, and fruit.
Unfortunately, this ‘new species’? was based on separate elements
belonging to different trees and different species from unknown
localities, sent to de Wildeman by a Mr. Ch. Patin, at that time
vice consul of Belgium in Panama. Although the specimens used
for the description were claimed by Patin as having been collected
from a single tree, Patin himself recognized later that they came from
different trees and that the fruits were from 7. simiarum. At that
time, Panama was part of Colombia and that is the reason why the
new species and the specimens were labeled as from Colombia.

1900: In a second work, Hart makes no alterations in his 1892
classification, stating that “after the lapse of some years I still see
no necessity to revise the list.”” But he extended his comments on
characteristics, variations, and properties of the cacao varieties with
new considerations, as for instance: ‘‘Calabacillo is certainly as far
removed from Forastero, as Forastero is from Criollo, as seen in
plantations of the present day, when every intermediate form from
Criollo down to Calabacillo can be seen linking the whole in one
continuous chain of varieties. To properly classify Cacao, we must
first know what the originals were like, and it is clear that at the
present time, it is hard to decide exactly what were the forms assumed
by the older types of Cacao fruit. There is an apparent consensus
of opinion however which points to the thin-skinned and _ bottle-
necked variety as the original Criollo (Spanish for Creole), and this
is quite confirmed by the Criollo being discovered in the virgin forest
of an uncultivated part of Trinidad” (p. 52).

The Criollo as well as the cacao of Java and Ceylon, the Criollo
of Central America and T. pentagonum have the seeds white or almost,
white inside. The best quality Forasteros have the seeds slightly
violet, and the Calabacillo strongly colored. He adds that the best

CUATRECASAS—CACAO AND ITS ALLIES 399

qualities of fruits of Venezuela (e.g., from Ocumare) are distinguished
by the lightness of the seeds, and their shape, although the pod might
belong to the type of Forastero. In Trinidad, in the Estates where
certain strains of cacao from the continent had been introduced, we
find the finest qualities of Forastero. The illustrations, with a
similar nomenclature, are less clear than those of first edition.

1901; Paul Preuss publishes the first important field report ever
written with keen observations on the varieties of cacao, conditions
of cultivation, conditions and ways of preparation, and qualities of
the products in several countries of South and Central America. It
is the report of his trip made during 1899 and 1900, in order to obtain
information on tropical crops for the Colonial-Economic Committee
of the German government. From Surinam he names three varieties
of 7. cacao as cultivated: “Surinam,” also called ‘Porcelaine’’ (cor-
responding to the Amelonado of Trinidad), ‘‘Aligator” (the Cunde-
amor of Venezuela and Trinidad), and “Caracas” (similar to Carupano
or Forastero from Trinidad). He also observed mixed intermediate
forms. From Trinidad and Grenada, where he saw extensive planta-
tions, he describes several cultivars: “Amelonado,” ‘“Calabacillo,”
“Sangre de toro,’ ‘Forastero,” “Criollo’”’; he mentions that Hart
calls Criollo what is called Forastero in Venezuela and that Hart’s
Forastero is the Venezuelan Criollo. But according to Preuss, the
formerly general use of the name Forastero in Trinidad applied to
cacaos other than Criollo. At present, Forastero has the same
meaning as Trinitario, which might have been introduced from the
Venezuelan eastern coast or from the Orinoco region, after the earlier
Trinidad plantations (all of the Criollo type) had been destroyed by
some kind of disease. Preuss goes on to describe the three main
Trinidad varieties using the concepts of Hart: ‘‘Forastero,’”’ ‘“Amelo-
nado,” and ‘“Calabacillo.”” He compares the yellowish variety of
Amelonado to the Guayaquil cacao and the elongated form of Foras-
tero to the Colombian cacao, Of Venezuela, Preuss says that it
is the classic land of Criollo, producing large seeds with thin shell
of the best quality. He distinguishes two main cacao varieties in
Venezuela: 1) Cacao Criollo, 2) Cacao Trinitario. Preuss clearly
distinguishes seven types, but there are also many intermediate
forms.

1. Angoleta.
2. Cundeamor (Cundeamor legitimo, with red shell and Cundeamor amarillo,
with yellow shell).
3. Carupano legitimo (Carupano grande, and Carupano mestizo, red with
yellow).
. Carupano parcho (yellow fruit).

. Carupano Taparito (yellow or brownish).
. Sambito (red or yellow, short, thick, rather smooth fruit).

Ao

400 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

7. Trinitario amargo or ‘‘Cojén de toro” (red or red brownish, smooth, blunt
or shortly attenuate pointed fruit).

He describes the Criollo tree and mentions its three varieties: ‘‘Criollo
legitimo” (the best quality) with deep red, ‘‘Criollo amarillo” with
yellow, and ‘Criollo mestizo”’ with yellow and red shells. The
seeds are white violet in the first and white in the second. Preuss sees
in Ecuador a great uniformity in the cacao fruits as he had observed
in no other country. They are of the Amelonado type, the same type
being cultivated in the Cameroons, St. Thomé, Grenada, and Surinam;
the Ecuadorian type has thicker shells. He mentions four varieties:
“Arriba,” “Balao,” “Machala,” and “Bahia.” He says that cacao
is frequent in the underlayer of the rain forests in Ecuador and that
these wild trees, which cannot be distinguished from those of the
plantations, also produce good fruits, which have very thick shells
and roundish seeds in section. Preuss sees in this wild cacao the
origin of the widespread plantations of the variety Amelonado (p.
247). Theobroma bicolor is also mentioned as a wild cacao (‘‘cacao
blanco,” ‘‘bacao”’) in Ecuador.

From Central America he describes the native variety “Cacao
Lagarto” (Alligator), the other native variety Criollo and two others
introduced, the “Cauca” from Colombia, and ‘‘Trinitario” from
Trinidad; in this Trinitario he distinguishes the three types Forastero,
Amelonado and Calabacillo. There are intermediate forms between
Lagarto and Criollo ‘Cacao del pais” difficult to separate one from
another; they have red and yellow pods with a thin shell and white-
violet or white cotyledons. The seeds of Nicaraguan Criollo are the
largest of all varieties and of the best quality known. Mention is
made and illustrations are given of T. bicolor, T. angustifolium, and
T. pentagonum from Central America. Also drawings of fruits and
seeds of five varieties of T. cacao (Nicaraguan and Venezuelan Criollo,
Calabacillo, ‘‘Cundeamor Legitimo,” and ‘‘Curupano grande’’) are
reproduced.

1902: De Wildeman gives a taxonomic compilation of Theobroma
in his book on tropical cultivated plants. Following Schumann he
divides the genus in JTerrania, Hutheobroma, and Bubroma and uses
Schumann’s key for the species (as in the Flora Brasiliensis), but adds
another species, 7. simiarum Donn. Smith, and accepts 7. pentagonum
as different from 7. cacao; altogether, nine keyed species and four
doubtful additional ones, as by Schumann, besides Herranias, are
treated. For each species, summarized descriptions, native names
and uses, geographic location, and miscellaneous comments, are
given. Speaking of 7. siméarum he writes: ‘Il faut rapporter en
partie 4 cette espéce la plante que nous avons décrite, en 1899, sous le
nom de 7. kalagua, dont la description avait été fait sur de feuilles,

CUATRECASAS—CACAO AND ITS ALLIES 401

des fleurs et des fruits ne provenant pas de la méme plante.”” Some
attention is given to 7. cacao and the differences between its seeds and
those of Herrania mariae, based on anatomical sections and gross
chemical analysis made by Heim. The varieties of Theobroma cacao
are reduced to ten “‘series” distributed in three ‘‘grand” groups—
I) Criollo with two varieties: 1) amarillo, 2) colorado; IJ) Forastero
with varieties: 3) cundeamor verrugosa amarillo, 4) cundeamor
verrugosa colorado, 5) amarillo, 6) colorado, 7) amelonado amarillo,
8) amelonado colorado; III) Calabacillo with varieties: 9) amarillo,
and 10) colorado.

1904: Lignier, in a list of the Caen City Herbarium, mentions
Theobroma sativa, as a Sagot collection from French Guiana without
further comment on the author or synonym; it probably refers to
Cacao sativa Aubl.

1904: Huber, a botanist at the Goeldi Museum, Belém do Para,
inaugurates a new epoch by publishing direct observations on the
botany, ecology, and location of species of Theobroma. Huber found
spontaneous and subspontaneous trees of 7. cacao in several places of
Amazonia and believes that it may really be indigenous in the forests
of the Alto Purtis Rio, Rio Ucayali, and other places down to San-
tarém and Obidos.

1906: Huber extends his explanations on the indigenism of T.
cacao on the alluvian soils of the Rio Alto Purts, in the inundatable
forests around the mouth of Rio Acre, and along the rivers Ucayali,
Japurdé, Jurud, and Madeira. Among observations on other species
(T. microcarpum, T. speciosum, T. obovatum (as T. sylvestre), T.
subincanum, and T. bicolor) he shortly describes a new species from
Peru, Theobroma sinuosum Pavon.

1906: Huber publishes a variety coriaceum of T. speciosum found
in Brazil. At the same time, he mentions having found 7’. cacao
growing wild in a forest near the Canchahuaya Lake.

1908: Chevalier, in his extensive studies on cacao crops in western
Africa, besides detailed descriptions of the cocoa tree and its growth
and ecology in Africa, mentions its variations and describes a new
species, Theobroma sphaerocarpa, distinguished by its globose, almost
smooth fruit 9 to 11 cm. in diameter, which has been cultivated for
a long time on the island of Sao Tomé.

1911: Hart, in the introduction of his book on cacao, gives an
excellent account of the varieties of the cultivated cacao, discussing
the external features of the chief different types and the qualities of
their crops:

“The species known as Theobroma cacao, covers innumerable varie-
ties or forms, differing in shape of pods, in size and vitality of trees,
in bearing capacity, and in colour, shape and quality of the bean.

402 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

The many names under which varieties of this tree (Theobroma
cacao) are known, do not constitute species, but must be merely con-
sidered as varieties of one species. These varieties probably owe
their origin to seed variation and cross-breeding, together with the
local influence of soil and climate, but it would serve no useful purpose
to record the names by which they are known, as these differ in each
district, in each plantation, and in each country where they are
grown”’ (p. 2).

‘We have therefore a classification under Theobroma cacao carrying
fourteen types under three classes, but it must be understood that
these are separated by no definite margin, and that intermediate
forms will be found on estates showing every conceivable form of
variation” (p. 3).

Hart’s classification here amplifies his initial one of 1892, mainly
by broadening the concept of Criollo, in which class he includes types
known under this name outside Trinidad a long time before.

THEOBROMA CACAO

Class I. Criollo
Trinidad Criollo.
(1) Var. a. Amarillo= Yellow, thin-skinned, bottle-necked.
(2) Var. 6b. Colorado=Red, thin-skinned, bottle-necked.
Venezuelan Criollo.
(3) Var. a. Amarillo= Yellow, thick-skinned, high-shouldered, some-
times pointed.
(4) Var. 6. Colorado=Red, thick-skinned, high-shouldered, some-
times pointed.
Nicaraguan Criollo.

(5) Var. a. Amarillo— Yellow
(6) Var. b. Colorado—Red

Thick-skinned, high-shouldered, and
very large beans with light-coloured
interior.

Class II. Forastero

(7) Var. a. Cundeamor verugoso Amarillo = Yellow-warted.

(8) Var. b. Cundeamor verugoso Colorado = Red-warted.

(9) Var. c. Ordinary, or typical Amarillo= Yellow Forastero.
(10) Var. d. Ordinary, or typical Colorado=Red Forastero.
(11) Var. e. Amelonado Amarillo= Yellow, melon shaped.
(12) Var. f. Amelonado Colorado= Red, melon shaped.

Class III. Calabacillo
(13) Var. a. Amarillo= Yellow |Calabacillo, flat-beaned, smooth, thin-
(14) Var. b. Colorado= Red or thick-skinned, and small pods.
THEOBROMA PENTAGONA

(15) Theobroma pentagona= Alligator cacao. Has yellow, much-
warted pods, with five distinctly
raised ribs, and large beans, having
white or light-coloured interior.

CUATRECASAS—CACAO AND ITS ALLIES 403

Hart characterizes the Criollo varieties by their light-colored seeds,
the high quality of the cured product, and the less vigorous growth.
The Forastero has light to dark purple seeds in large rough-ridged
pods; it is rather variable and is a strong grower. Calabacillo is an
inferior but stronger growing tree, having ovoid pods with thin,
solid, dark-colored seeds. “7. pentagona has the largest seeds of
any known species.” ‘In general, however, it is hard to say where
one form begins and another ends.” In fact in most countries,
cacao “consists of a heterogeneous mixture of cross-bred varieties of
one species (7. Cacao) though of late years it is thought possible that
the common species may have become hybridized with 7. pentagona.”

1914 and 1982: Van Hall publishes his well-known book on cacao
which since then has been a textbook and main source of information
for cacao growers and agronomists. In his botanical chapter he
presents some nonoriginal information on the noncultivated Theo-
bromas. He gives excellent comparative descriptions and evaluations
of the most important variations and forms of cultivated cacao.
He follows Morris in recognizing two groups: 1. Criollo; 2. Foras-
tero. He describes seven types or subvarieties of the first: the
Venezuelan, the Ceylonese, the Javan, the Samoan, the Madagas-
carian, the Nicaraguan, and the Surinam Criollos. The subvarieties
of Forastero described are Angoleta, Cundeamor, Amelonado, and
Calabacillo. Theobroma sphaerocarpa Chev. is considered a mere
form of Calabacillo. In 1932, van Hall introduces a new name
T. aspera, transferred from Herranza.

1914: Pittier publishes two new species of Theobroma from Panama:
T. bernowilli * discovered by him in the forests of the Colon province,
and T. purpureum which belongs to Herrania.

1915: Cook, after a thorough morphological field study of T. bicolor
and 7’. cacao, especially of the branching system and the flower and
inflorescence structure, decides that they belong to different genera,
and publishes a new monotypic genus Tribroma with one species, T.
bicolor; it mainly differs by its clusters of three branches and the woody
pericarp in contrast to the 5-branching clusters and fleshy pericarp of
T. cacao. Although all observations by Cook are very sound, he does
not compare 7’. bicolor with other species of Theobroma besides cacao.
~ 1916: Cook publishes in detail and with illustrations the results of
his studies on growth and dimorphism of branches and leaves of 7.
bicolor and T. cacao. He explains the sympodial structure of the
Theobroma stem, the verticillate primary branching, the formation
and succession of upright lateral shoots, which continuing the main
stem, bring the clusters into a lateral position and the alternate

1 Originally so misspelled, although the correct name would have been “bernoullii.””

404 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

branching of lateral branches. Cook describes the two types of leaves:
the long petiolate symmetrical leaves of the seedlings and upright
shoots and the bilateral, asymmetrical, dorsiventral, short-petiolate
leaves of the lateral branches. The structure of inflorescences and
flowers also are studied.

1918: Stahel publishes a precise, illustrated, morphological study
of the structure of the inflorescences of 7. cacao and T. bicolor. These
inflorescences are always formed in the axil of the subtending bract of
a lateral, non- or short-developing bud. The inflorescence in T. cacao
consists of a cincinnous main axis with many short internodes and a
few dichasial final branchlets. In 7. bicolor, the main cincinnous axis
has a few, long internodes, and the lateral branching is dichasial. The
peduncles have a bilateral, the pedicels a radial structure.

1921: Benoist publishes a new species of Theobroma from French
Guiana found by himself, 7. velutinum. It is characterized by large
leaves, velutinous beneath, and by ellipsoid, 5-ridged, velutinous pods.

1923: Standley recognizes three spontaneous species for the Mexican
flora: T. cacao, T. angustifolium, and T. bicolor, and gives interesting
historical, geographical, and economic data.

1925: Ducke publishes firsthand new botanical and ecological data
on several Brazilian species.

1925: Pittier presents his interesting new theory that all existing
forms of cultivated cacao are the result of hybridization between two
initial species: 1) Theobroma cacao L., with elongate, claviform, rugose,
10-ridged pods, containing large, ovoid, white or slightly yellowish
seeds, and 2) 7. letocarpwm Bernoulli, with more or less rounded,
smooth, slightly 5-ridged pods, with flattened, more or less triangular
and dark purplish seeds. The first type is the one commonly known
as “Cacao dulce” or ‘Cacao criollo,’’ the second is commonly known
as Calabacillo or Trinitario and in Guatemala as “Cumacaco.” T.
sphaerocarpum Chev. from Sao Tomé is a hybrid, retrogressive to 7.
leiocarpum. The two typical forms are united through an unlimited
number of intermediate hybrid forms the characteristics of which are
very variable; he says that the present nomenclature of varieties
should be abandoned.

1925: Chevalier, in his “Observations” to the preceding work of
Pittier, accepts basically Pittier’s theory, and says that 7. sphaero-
carpa would be the extreme form of the series with smooth fruits and
that T. leiocarpa is a hybrid between 7. cacao and T. sphaerocarpa.

1926: Pittier answers Chevalier saying that he has never found 7’.
sphaerocarpum in Central America, but that he has found growing
wild in Costa Rica and Panama forms almost identical to 7. /evocarpum
Bernoulli. He recognizes that the forms with small, round pods often
found in the region of Barlovento (Venezuela) are very close to T.

CUATRECASAS—CACAO AND ITS ALLIES 405

sphaerocarpum. Both original species are still found in pure, typical
form and it can be said that in most plantations, next to the “primitive
species’ a number of hybrids are found. ‘‘We have still in Venezuela
cacao plantations with absolute domination of the Criollo type, as, for
instance, it happens at Caruao and Chuao.” There can be observed
over great extensions of land, tree after tree with elongated, claviform,
pointed fruits, which may be reddish or yellow, with rounded seeds and
almost white and insipid cotyledons. Pittier adds that at the time
Linné described T. cacao, the Criollo was the dominant form in
cultivation.

1930: Myers reports about his exploration of the upper Mamaboen
Creek, a tributary of the Coppename River, where wild cacao
was discovered by Stahel ten years earlier. This place is located in
the middle of the Surinam rain forests, 40-50 kms. away from the last
small Indian village. Abundant trees of cacao in a wild state were
found in the lower tree-layer of the forest, under conditions of dense
shade and high humidity; most of the cacao trees were found on the
flooded margins of the river; few were found on higher ground. The
trees were 10 to 25 feet high, and abundantly fruiting; the ripe pods
were bright, light yellow in color, and almost smooth (with little indi-
cation of longitudinal ribs) and with 40 to 50 seeds with deep violet
cotyledons. Myers adds that the fruits were of the Amelonado-
Forastero type. He also comments on information about wild trees in
other parts of Surinam, British Guiana, Brazil, and elsewhere.

1930: Pittier publishes an abridged key to classify the known species
of Theobroma, including Herrania. Following Schumann in Die
Natiirlichen Pflanzenfamilien he divides it into three sections, the first
being Herrania. The other two sections, defined by diantheriferous
(Sect. Hutheobroma) and triantheriferous stamens (Sect. Bubroma),
comprise 13 species, which are distributed in subsections corresponding
to the Bernoullisections. In Hutheobroma he includes subsect. Cacao
with 7. cacao, T. leiocarpum, and T. pentagonum and subsect. Rhyti-
docarpus with T. bicolor and T. Bernouillu. In Bubroma there are
subsect. Telmatocarpus with only 7. microcarpum, subsect. Oreanthes
with 7. spruceanum, T. speciosum, and T. simiarum and subsect.
Glossopetalum with T. angustifolium, T. grandiflorum, T. subincanum,
and J’. sylvestre. The characters given in the key are few and not
always the most typical or correct, some of the species being wrongly
placed (7. Bernowillii, T. simiarum). The binomials 7. glaucum,
T. martii and T. album are considered dubious. Pittier refers to
having received from Cook photographs of two forms of cacao culti-
vated in Peru with small fruits which might be different species; one
has almost spherical, rugose pods called locally “cacao chuncho.”
Pittier says that, with the exception of 7. leiocarpum and T. cacao

406 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

(respectively Calabacillo and Criollo), there do not exist stable forms
of cultivated cacao. Forastero from Trinidad stands between these
original species and can be considered as the result of their crossing.
The Trinidad Forastero seems to be quite different from Venezuelan
and Central American Forastero and probably from the Old World
Forastero. He also says that it does not make sense to talk about
the origin of Forastero, because its forms appear wherever the parental
species are present and successive crossings produce constant varia-
tions. Commenting on the finding of Myers in Surinam, Pittier says
that the wild cacao of the Mamaboen valley belongs to T. leiocarpum.
The same opinion is expressed in connection with Schomburgk’s
spontaneous cacao found in the Rio Branco valley and numerous
references about wild cacao throughout the Venezuelan Guayana.
The cacao with tapering, pointed, ridged and rugose pods with white
cotyledons (7. cacao) has really never been found indigenous to the
east of the Panama isthmus. Pittier found it at the isthmus but says
that the origin of this species is to be sought towards the north, in the
Soconuzco, Chiapas, and Tabasco regions, where the Criollo type
finds the best conditions for its development. Conversely, the planta-
tions of Calabacillo do not go much further west than Costa Rica
where it was recently introduced. Here are to be found the western-
most stations of T. leiocarpum, although the type came from planta-
tions in Guatemala.

1931: Mildbraed publishes a new species 7’. tessmannii, found by
Tessmann in eastern Peru. He relates it to 7. ferrugineum Bernoulli,
from which it differs especially by the long, soft tomentum of the
leaves beneath.

1932: Cheesman, who had been conducting research on cacao corps
for many years, presents a significant account of the economic botany
of Theobroma. He believes that it is unlikely that any of the now
uncultivated species has any direct economic significance. He speaks
about the cultivated species, those of section Cacao, as interfertile
species (7. cacao, T. letocarpum, T. pentagonum). Cheesman says
that ‘the taxonomic status of the group of forms at present included
under the collective term 7’. cacao can only be determined by pro-
longed research, including genetic, and possibly also cytological
studies.” Cheesman agrees with Pittier in considering that more
than one species contributed to build the “cacao complex,” adding
that this idea provides the most helpful way of regarding the ex-
traordinary variation exhibited by the crop. His discussion on the
history, characters, and merits of the varieties is illuminating in many
respects, especially when trying to understand their possible origin.
In practice, he follows, with slight modification, van Hall’s classifica-

CUATRECASAS—CACAO AND ITS ALLIES 407

tion which he says ‘“‘is a compromise between a natural and an artificial
system.”

1932: Pittier insists in his viewpoints on the origin of the cultivated
cacao. Finding that 7. cacao L. is not well typified by a known
variety, Pittier decides to abandon this name and to substitute for it
T. sapidum, but no specific description is given and a type specimen
is not indicated. The Calabacillo seems to have a stronger fertilizing
power than the Criollo which explains why few trees of the first suffice
to alter the plantations of the second.

1933: Ciferri makes a very detailed, critical study of the cultivated
cacaos of Santo Domingo giving a thorough classification with defini-
tion, descriptions, and illustrations of their numerous types. Mor-
phological, taxonomic, historical, and economic comments are given.
Ciferri follows the principles of Pittier in accepting the theory that the
majority of the cultivated cacaos are hybrids of two initial types,
but he considers these two types varieties instead of species. Ciferri
gives the system formal nomenclatural status by publishing the fol-
lowing new varieties: 7. cacao L. emend. var. typica Ciferri; T. cacao L.
emend. var. leiocarpa (Bernoulli) Ciferri, and 7. cacao var. typica X
T. cacao var. leiocarpa = Forasteros. ‘I cacao denominati global-
mente ‘‘Forasteros’”’ sarabbero dunque, secondo l’idea di Pittier, che
moi adottiano in pleno, i meticci tra le due variet&é sunnominate del
T. cacao.” Ciferri’s work is a significant contribution to the knowl-
edge of cacao varieties and their distribution.

1935: Cheesman describes the branching system and dimorphism
in cacao, and studies different ways of vegetative propagation.

1936: Campos Porto publishes information on the species of Theo-
broma cultivated at the botanical garden of Rio de Janeiro, with a
photograph of the inflorescences of JT. speciosum; the other species
referred to are 7’. cacao, T. bicolor, T. grandiflorum, T. subincanum, and
T. microcarpum.

1987: Standley lists and describes five species of Theobroma in his
Flora of Costa Rica plus a Herrania as T. purpureum. He considers
T. simiarum, T. angustifolium, T. bicolor, and T. cacao to be wild in the
forests and 7’. leitocarpum as probably wild.

1937: Pérez Arbélaez publishes a manual for the cacao growers of
Venezuela including a botanical introduction with descriptions of and
information on Venezuelan cultivars; he also gives interesting historical
data.

1938: Pound finds interesting varieties of cacao in his explorations
in South America, especially in the upper Amazon basin.

1938: Bondar publishes a documented book on the cultivation of
cacao in Bahia. He gives comments and a key to the known species
of Theobroma. He considers that the varieties cultivated in Bahia for

680-695—64—8

408 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

many years belong to 7. levocarpum; a great uniformity in the prod-
uct exists, but he believes that recent introductions of Criollo may
well stop this uniformity. He also describes and illustrates several
forms of Forasteros known in the region.

1939: Diels publishes a new species, 7’. calodesmis, found by Hertha
Schultze-Rhonhof in the rain forests of eastern Ecuador, which he
relates to 7. speciosum and T. bernouillii.

1939: Cope’s investigations in Trinidad on agents of pollination
using thrips (Frankliniella parvula Hood), red ants (Wasmannia
auropunctata, Rog.), and aphids (Yoxoptera aurantii B. de Fousc.),
suggests that red ants and thrips are the responsible agents of pollina-
tion in a cacao population at River Estate.

1940: KE. W. Emmart publishes the Badianus Manuscript, the
earliest book ever written on Mexican medicinal plants, the work of
two Aztec Indians, Martinus de la Cruz who composed the work in
Aztec, and Juannes Badianus who translated the text into Latin.
On plate 68, illustrating six plants, plant no. 2 represents ‘Tlapal-
cacauatl,” colored cacao, i.e., “tlapal’” = colored, ‘“‘cacauatl’”’ = cacao,
according to Emmart (p. 273), who adds, ‘This picture is the earliest
illustration of the cacao, Theobroma cacao L., the source of chocolate.”
This interesting, primitive drawing clearly illustrates the Criollo
variety.

1940: Ducke summarizes his experience of many years in Brazilian
cacaos with a new and detailed key to the Brazilian species. He
brings new data into consideration in his classification, as for example
fruit characters that were unknown before. He gives photographic
illustrations and new information, based on direct field observations,
about morphological, phenological, and ecological features of the
species treated, which are T. cacao, speciosum, spruceanum, micro-
carpum, obovatum, subincanum, and grandiflorum. He also includes
in Theobroma the genus Herrania with one species, T. Mariae. Con-
cerning 7. cacao, Ducke recognizes it as indigenous throughout the
central and western Amazonia. Ducke considers 7. leiocarpum
Bernoulli a mere form of 7. cacao and makes the new nomenclatural
combination: 7. cacao L. forma leiocarpum.

1942: Schery publishes a new species, 7. asclepiadiflorum, based
on specimens from Panama.

1944: Cuatrecasas publishes a new species with yellow flowers,
T. cirmolinae, found by the author in the rain forests of the western
slopes of the western Andes in Colombia.

1944: Cheesman makes a thorough examination of the taxonomic
situation in cacao, the most important conclusion being that the whole
assemblage of wild, semiwild, and cultivated cacao constitutes “one
interbreeding population.”” He still supports the main division of

CUATRECASAS—CACAO AND ITS ALLIES 409

cacao into two groups of varieties, Criollo and Forastero. He pro-
poses the new theory that the Criollo, which may occur wild in some
regions from southern Colombia to southern Mexico, may have
originated at the headwaters of the Amazon. He divides this group
into Central American and South American Criollos. The Forasteros
are divided into Amazonian Forasteros, which can be found wild in
Amazonia and which are widespread in cultivation, and the Trini-
tarios, possibly originating from the mingling of South American
Criollo and Amazonian Forastero stocks. Theobroma pentagonum is a
simple form of J. cacao, probably a segregate of the large cross-
fertilized population. The same opinion is expressed with regard to
T. leiocarpum, which, according to Cheesman, does not belong to
Amelonado; it is an aberrant form of the Criollo, for which rea-
son the binomial falls into the strict synonymy of 7. cacao. The
data assembled and arguments of Cheesman are a very valuable
contribution.

1946: Chevalier publishes a monographic revision of Theobroma.
He recognizes 13 species (excluding Herrania) arranged according to
the five sections of Bernoulli. He includes in the first section (Cacao)
only T. cacao, in section II, Oreanthes: T. guianensis and T. spruceana;
in section III, Rhytidocarpus: T. bicolor, T. glauca, and T. Bernowilli;
in section IV, Telmatocarpus only T. microcarpa and in section V,
Glossopetalum: T. sylvestris, T. obovata, T. ferruginea, T. grandiflora,
T. angustifolia, and T. simiarum. In the key (Tableau Analytique),
the species are differently arranged; they are divided in the two sec-
tions of Schumann: Eutheobroma with four species (T. cacao, T.
bicolor, T. Bernouillii, and T. glauca), and Bubroma with the other
nine species. The characters given in the key are not always well
chosen, and some species are misplaced in the sections (as, e.g., 7.
glauca and Bernouillit). The nomenclature and typification of the
species are not always correct; the concepts of 7. speciosa Willd. ex
Spreng., 7. guianensis (Aubl.) Gmel., 7. velutina Benoist, T. syl-
vestris Mart., T. sylvestris (Aubl.) Don., 7. obovata Klotzsch ex
Bernoulli, 7. ferruginea Bernoulli, T. sinuosum Pavén and others
are actually not clarified. Some confusion is also brought with the
new names 7. sagittata Pavén, 7. hastata, and T. undulata. I have
identified T. sagittata Pavén as Herrania nitida and suppose that T.
hastata Cheval. is a lapsus calami for the former and 7. undulata
Cheval. a lapsus calami for T. sinuosum.

Special treatment is devoted to J. cacao, with which Chevalier
had long and sound experience. He considers all cultivated cacaos as
belonging to a single species, J. cacao L., in which four different races
or “species jordaniennes” can be recognized. These races, which
cross among themselves “A l’infini,” can be only distinguished by their

410 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

fruits and seeds. These are lacking in herbaria, and therefore the
races must be studied in the field. Four of the races or Jordanian
species of Chevalier correspond to formerly described species (7°.
sativa, T. leiocarpa, T. pentagona, and T. sphaerocarpa) but he adds a
fifth based only on foliage, 7. sagittata, which, as I have pointed out,
is not a Theobroma. Chevalier’s classification for the “formes jorda-
niennes” of 7. cacao L. follows:

Leaves obovate-oblong, acuminate.
Fruits ovoid-oblong 5-10 ridged, + rugose-bullate, long-attenuate into a

point. . ee Loe . T. sativa
Fresh seeds yellowish white .......... =... Var. leucosperma
Fresh seeds dark violet. . . . . - . + var. melanosperma

Fruits ovoid, rounded at apex, smooth or with 5-10 shallow furrows. T. letocarpa

Fruits globose, more or less smooth, rounded or depressed at apex . 7’. sphaerocarpa

Fruits ovoid-oblong narrowed to the apex, with 5 very prominent ridges.

T. pentagona
Leaves narrowly oblong, or oblong-acuminate, more or less undulate . 7”. sagittata
Chevalier goes on to describe the three first “races,” and declares that
he had insufficient information on the other two. Chevalier uses for
what he calls races, Jordanian species, or Jordanian forms, the same
binomial denomination as for species, for instance, 7. cacao L. forma
T’. sativa which should be T. cacao I. forma cacao according to the
present rules of nomenclature. It seems right to consider the original
species described by Linnaeus as belonging to the Criollo form, but
nothing is clarified by Chevalier using the binomial T. sativa, because
this name was based on 7. cacao L. Chevalier considers his 7. sativa
originally from Central America. To the “Jordanian form” T.
leiocarpa Bernoulli, Chevalier refers the “Cacao creoulo” of Sao Tomé,
the Cumacaco, Calabacillo, and Trinitario, and he supposes it origi-
nated in Guiana and Brazil. Concerning the Jordanon 7’. sphaero-
carpa Chevalier, described on Séo0 Tomé (Africa) plants, very similar
specimens have been found in Venezuela (var. sambito), in the high
Amazonian forests, and at the Rio Marafién. The Jordanon 7
pentagona has never been found wild; it seems to be originally from
Central America. The experience and opinions of Chevalier have
to be taken into account when considering the classification and origin
of cultivated cacaos.

1946: Cuatrecasas publishes 7. capilliferum discovered on the
Pacific coast of Colombia.

1947: Llano Gémez publishes information about the cultivated
cacao in Colombia, with several plates in color representing the
principal types.

1948: Rombouts discusses Theobroma Saltzmaniana Bernoulli,
showing that it might be based on a flower with defective or abnormal

CUATRECASAS—CACAO AND ITS ALLIES 411

petals and therefore cannot be distinguished from other forms of
T. cacao.

1949: Standley and Steyermark consider five species of Theobroma
in their Flora of Guatemala, recognizing T. pentagonum and T.
leiocarpum as different from T. cacao following Bernoulli. T. angustz-
folium is given as cultivated and 7. bicolor as uncertainly native.

1949: Cuatrecasas and Leén describe a new species, 7. mammosum,
collected by Leén as a rarity on the Atlantic coast of Costa Rica.

1950: Holdridge publishes some new information on Mexican and
Central American species of Theobroma, with a key to nine species
and one Herrania. He suggests that 7. pentagonum might be the
original type and source of the cultivated cacao in Mexico and Central
America and that the Criollo types were the product of interbreeding
of T. pentagonum with the South American T’. lecocarpum.

1950-1958: Cuatrecasas publishes 7. stipulatum and T. nemorale
from the rain forests of the Pacific coast of Colombia and T. gilert
from the Pacific range of Ecuador.

1951: Freytag publishes a revision of Guazuma, which helps in the
study of its relationships of Theobroma. The genus is reduced to
four species.

1951: Addison and Miranda Tavares explain the results of their
six-year work in trying to produce hybrids from different 7 heobroma
species. They crossed 7. cacao with all the Amazonian species of
Theobroma, without success, and proceeded then to cross the other
Amazonian species. In 1946, from 719 pollinations of 7. speciosum
on T. cacao, they obtained 29 fruits and 979 seeds, which were mostly
abnormal and did not germinate. Same results were attained by a
few pollinations with Herrania mariae. Among 798 cases of polli-
nation of JZ. microcarpum on T. cacao 11 fruits and 26 seeds were
produced, from which only three seedlings were produced which
grew no more than 10 cm. Similar negative results were produced
from J. cacao X obovatum and T. bicolor X cacao. Some particular
trees of J. cacao were more receptive than others; one of them gave
fruits when submitted to pollination from all other species. When
T. cacao was used as pollinator on 7. microcarpum, T. speciosum, and
H. mariae, no fruits or seeds were obtained. In 1947, another series
of cross-pollinations were made on J. cacao with similar results,
although a few more or less viable hybrids were produced, e.g.,
T. cacao X microcarpum gave 28% fruits, but these decayed after
developing one month.

Better results were attained by Addison and Miranda in crossing
T. grandiflorum and T. obovatum; many hybrid seedlings were pro-
duced and several developed into perfect trees (in 1} years); the
leaves, fruits, and flowers of the hybrids showed intermediate char-

412 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

acters, and the pollen grains were normal and fertile. Well-developed
hybrids between 7. grandiflorum and T. subincanum, T. obovatum and
subincanum, and T. speciosum and T. sylvestre (= spruceanum) were
also produced. In 1948, some fruits were obtained by crossing
I’. cacao with T. grandiflorum, but these gave very few seeds, from
which only few plants developed up to 15 cm.

Addison and Miranda also made grafting experiments with good
results using 7. grandiflorum, T. obovatum and T. subincanum. T.
bicolor, T. speciosum and T. sylvestre (spruceanum) proved to be
another successful grafting group.

During their experiments, Addison & Miranda had the opportunity
of making interesting morphological and physiological observations.
The seeds of Theobroma usually germinate within 15 days. Theobroma
subincanum, obovatum, grandiflorum, microcarpum, and H. mariae
were found to have hypogeous germination, whereas 7’. cacao, sylvestre
(spruceanum), bicolor, and speciosum have it epigeous. The very
young leaves are green in 7. speciosum, sylvestre, bicolor, and micro-
carpum, but they may be green or red in the other species.

1952-1953; The Anglo-Colombian Cacao-Collecting Expedition pub-
lishes reports on its explorations in the search for wild and cultivated
species of Theobroma and Herrania in Colombia. The expedition took
place from June 1952 to October 1953, with the participation of the
British botanists and specialists F. W. Cope, D. J. Taylor, R. E. D.
Baker, P. C. Holliday, and B. G. Bartley. The Colombian botanists
who joined the expedition were H. Garcia Barriga, Canuto Cardona,
R. Romero Castafieda, and Alvaro Fernandez P. The main areas
explored were: (1) parts of the rivers Caqueté, Apaporis, Vaupés,
Negro (Guainfa), Infrida, and their tributaries in the provinces of
Amazonas and Vaupés, from 1°30’ S. to 3° N. and from 67° W. to
71° W.; (2) parts of the rivers Putumayo, Caquet4, and Caguian in the
provinces of Caqueté and Putumayo, from 0°20’ S. to 2° N. and from
74° W. to 77° W.; (3) parts of the trans-Andean provinces of Valle del
Cauca and El Chocé, from 3° N. to 6° N. and between 76° W. and 78°
W.; (4) scattered areas in the provinces of Antioquia, Norte de
Santander, Magdalena, Santander, and Huila.

The expedition made 191 botanical collections, of which 63 were of
living material sent to Trinidad. The well-preserved specimens have
been extremely useful for the study of the species and their
geographical distribution. Twelve indigenous species of Theobroma
were collected (7. calodesmis, microcarpum, subincanum, grandiflorum,
obovatum, capilliferum, gileri, nemorale, cirmolinae, simiarum, stipu-
latum, and chocoense). T. bicolor, always found planted, and 7’. cacao
were also collected. In a very few areas (Rio Caguan, Rio San Miguel)
spontaneous trees of cacao were found inside the forest but under

CUATRECASAS—CACAO AND ITS ALLIES 413

circumstances that make it impossible to say with complete certainty
that these trees were spontaneous. In general the subspontaneous
and planted cacaos found in the southeastern region of Colombia
were of the very uniform Amelonado type. The information and
materials (living and preserved) gathered by this expedition are a
very important contribution to the knowledge of Theobroma.

1954: Ducke makes a revision of his previous synopsis of the
Brazilian species, incorporating new morphological data into an
accurate, precise, well-balanced key. He introduces the character of
the ramification being 5-whorled and 3-whorled in separating 7’. cacao
from the other seven Brazilian species (7. bicolor, T. speciosum,
T. spruceanum, T. microcarpum, T. obovatum, T. subincanum, and
T. grandiflorum). Ducke considers each fertile stamen as the union
of two or three (stamens geminous and trigeminous), and as Addison,
Molina, and Pires had already observed before, characterizes 7.
spruceanum as having geminous stamens. Ducke still retains Herrania
in Theobroma, with two Brazilian species, T. Mariae and T. Ca-
margoanum (Schultes) Ducke. He summarizes the ecology and dis-
tribution of the genus in Brazil, calling it a typical Amazonian genus;
he writes that it is not absent in any place in Amazonia where rain
forest exists.

1956: Cuatrecasas recognizes seven species of Theobroma for the
Flora of Peru: 7. calodesmis, T. grandiflorum (planted), T. obovatum,
T. speciosum, T. subincanum, T. bicolor, and T. cacao subsp. leiocarpum,
which is found spontaneous in the rain forests of Peru.

1958: Schultes publishes the results of his discoveries and research
on Herrania. His synopsis comprises 17 species, eight of them new.
One species spreads northwards to Costa Rica, and the others are
limited to the humid tropics of South America. This monograph
shows the consistent unity of the group Herrania and the consistency
of the characters that may be used to separate it from Theobroma and
other related genera.

1958: Mora Urpi found a much greater variability in 7’. cacao
throughout Mexico and Central America than in South America. In
Central America and southern Mexico there can be found today prac-
tically all the known forms of cacao, for which reason Mora believes
that Central America has been the center of domestication of the
cultivated cacao; historical data also support this theory. He con-
siders cacao as having been probably introduced in South America in
pre-Colombian times. The geographical distribution of the Criollo
type would also prove this theory. The author agrees with Holdridge
in considering the pentagonum form as playing an important role in
the origin of the cultivated hybrid complex; he considers pentagonum
native in Central America and the original and most ancient form

414 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

of 7. cacao, from which, through mutation, introgressive hybridiza-
tion, and geographical isolation, the present population arose (p. 34).

1959: Soria confirms the observations of Mora about the great
variation in the characteristics of shells and seeds of Theobroma cacao
in plantations in Nicaragua. ‘Trying to establish the correlation be-
tween pod shape and color of the seeds, he found that dark-colored
seeds occurred in a large percentage of Criollo type pods, and white
seeds were often found in pods of the Forastero and Calabacillo types.
This agrees with the previous observation by Mora of dark seeds in the
I. pentagonum pod-type. These observations, according to Soria,
show that it is probable that the genetic factors controlling these
characters are independent of each other. Soria sees as reasonable
Holdridge’s theory that Criollo cacao is a result of crosses between 7.
pentagonum and T. letocarpum. But he adds “The possibility cannot
be overlooked, however, that the Criollos originated as mutations in
populations on the periphery of the area of distribution of the species,
the mutations afterwards being fixed and maintained through geo-
graphic isolation and selection. In this case, pentagona could be a
product of mutations in Criollo cacaos.” Soria emphasizes that
pentagonum can be fertilized very easily in either direction by other
types of T. cacao. Mora observed such hybrids as often having con-
spicuous characteristics of pentagonum. ‘All these observations lead
to the conclusion that pentagona is nothing more than one of the ex-
tremes in the variability of the complex of types forming the species
T.. cacao.”

1960: At the River Estate Experiment Station of the Imperial
College of Tropical Agriculture in Trinidad, 13 species of Theobroma
were planted for research and observation. Cope and Bartley suggest
the possible interrelationships of species of Theobroma, distributing
them in two groups: 1) with epigeal germination and growth con-
tinuing from below jorquette, comprising 7. cacao with 3-5-branched
jorquette and 7. bicolor, speciosum and calodesmis with 3-branched
jorquette; 2) hypogeal germination and growth continuing from above
jorquette in 7. microcarpum, grandiflorum, subincanum, obovatum,
angustifolium, mammosum, simiarum, cirmolinae, and nemorale. This
is the first attempt to classify the genus on the basis of germination
and branching.

1960: Cristébal publishes an excellent monograph on Ayenia with
important information concerning the relationships with other genera
of Sterculiaceae.

1960: Leén, in Hardy’s Cacao Manual, summarizes the taxonomy
of Theobroma, recognizing 19 species and considering as doubtful 7.
kalagua, tessmannii, ferruginea, and glauca. He gives abridged de-
scriptions and distribution for 7’. cacao, bicolor, bernouillii, capulifera,

CUATRECASAS—CACAO AND ITS ALLIES 415

calodesmis, asclepiadiflorum, microcarpa, gileri, guianensis (= speciosa
sensu. Chevalier), spruceana, angustifolia, cirmolinae, grandvflora,
mammosa, obovata, simiarum, stipulata, sylvestris (= subincana sensu
Chevalier), and nemoralis. The fruits of 17 species are illustrated.
In the classification he follows more or less Chevalier; under 7’. cacao
he distinguishes three subspecies: sativa (Lam.), letocarpa Bern. and
pentagona (Bern.); the listed cultivars are classified according to van
Hall.

1961: Soria reports on cacao in Mexico, having visited extensive
plantations in Tabasco. Before 1900, the variety cultivated in Mex-
ico was almost exclusively Criollo, but at present that is disappear-
ing, hardier and more productive varieties being substituted. He
observes great variability in the pod form in plantations of old
Criollo, which always have white seeds. Great variation is also
seen at present in the widespread hybrid populations that resemble
the Trinitario of South America, although the Mexican types lack
the red pigmentation of the shells usually exhibited by Trinitario.
(They are mostly whitish green or slightly reddish.)

Morphology

STEM AND BRANCHING (Fig. 1).—There is a dimorphism in the
vegetative organs of Theobroma. The main stem and the adventi-
tious orthotropic shoots have a radial structure, and the normal,
plagiotropic branches are monopodial and dorsiventral. The trunk
is sympodial.

The seedlings have an erect stem with regular, long-petiolate leaves
arranged in phyllotaxy cycles of 5/13, 5/8, or 3/8 (Cook, Baker).
After reaching a height of a few feet the vegetative end of the stem
stops growing and by the way of a cluster of secondary buds it
forks into 3 to 5 spreading branches arranged in a terminal whorl
called a “jorquette” or “fan.” These branches are plagiotropic and
dorsiventral, with alternate, distichous, short-petiolate leaves with
a phyllotaxy of 1/2. Further growth of the stem may now take place
by two different ways: 1. One of the dormant buds axillary to the
branches of the jorquette, and therefore adjacent to the central,
inert, apex of the stem, develops into a new vertical shoot with the
same structure as the main stem and looking like its continuation.
It grows to a limited extent, ending also with a whorl of 3 to 5 branches;
from above this second whorl or jorquette a third shoot is developed
in the same way, forming a third internode of the stem, and so on.
By this way a sympodial main trunk is built, with alternating inter-
nodes and nodes with regularly centered verticils of branches. Since
new terminal shoots are produced above the jorquette next to the

416 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

apex we can call this pseudapical growth. 2. No buds at all develop
above the first whorl of branches but an adventitious lateral one below
it grows into a vigorous upright shoot with a structure similar to that of
the main stem. Although lateral, it forces the node of the jorquette
to one side, and takes progressively the central position of the stem, of
which it will appear to be the continuation. After reaching some
length it forks, ending in a jorquette; a new adventitious shoot is
formed below that jorquette and so successively a sympodial trunk is
built, with alternating internodes and irregular nodes. In this case
the trunk is usually not truly straight and the whorls of branches, in
spite of the fact that these tend to take a circular position around the
stem, are always more inclined to one side, often making a lateral
bunch; the closer to the jorquette the lateral adventitious shoots
originate, the less irregular is the appearance of the sympodium
resulting. We can call this subterminal growth.

The dimorphism of the stems is transmitted by the buds. Those
of the seedlings and upright (orthotropic) shoots produce only, again,
orthotropic shoots (chupons) bearing long-petiolate leaves and pro-
ducing only the plagiotropic, dorsiventral branches of one terminal
jorquette. The buds of the lateral, plagiotropic branches produce
only other plagiotropic branches. Only exceptionally due to special
physiological conditions or following mechanical injuries (e.g., trim-
ming), do plagiotropic branches originate upright shoots (chupons).
More exceptionally the extraordinary formation of alternate plagio-
tropic branches has been observed on upright stems which have
failed to form a jorquette (Baker 1961, p. 9), but this has to be con-
sidered an abnormal case due to unknown special conditions of some
cultivated trees.

The lateral, plagiotropic branches are monopodial and branch by
axillary buds; frequently the growth of a lateral branch bends the
young joint of the primary branch forcing this into an angle, thus
simulating a dichotomous fork; branches may appear several times
forked and are then called “dichotomous” branches.

The stem and branching dimorphism is important in the practice
of propagation and cultivation of Theobroma trees, because only the
trees produced by cuttings of orthotropic stems (chupons) are upright
and regular; conversely, those from plagiotropic (dorsiventral) lateral
branches, branch bilaterally (dorsiventrally) and tend to slant or to
bow (incline), being thus weaker. In cultivated cocoa the formation
of adventitious upright shoots (chupons) on branches and at the base
of the trees is frequently observed; they may be used in practice to
regenerate old trees by pruning, and as cuttings for propagation.
But the production of chupons is always too small to serve for ex-

CUATRECASAS—CACAO AND ITS ALLIES 417

7

N 111 pes

EInezcw

CHR

Ficure 1.—Stem growth in Theobroma trees. a, B, subterminal or subapical growth: a,
adult seedling of T. cacao with its primary stem (bearing long-petiolated, radially
arranged leaves) topped by a whorl of dorsiventral, leafy branches; B, formation of the
sympodial trunk in T. cacao by way of upright adventitious shoots from buds borne
below the terminal verticil (or jorquette). c, p, pseudoterminal growth: c, formation
of the sympodial trunk by developing one of the axillary buds of the terminal whorl
of branches (e.g., sect. Glossopetalum); p, apex of stem topped by a whorl of 3 branches
each with an axillary bud, of which only one will develop (growth above jorquette).

tensive propagation. The two different kinds of stem growth have
taxonomic implications.

418 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Leaves.—There is a dimorphism of leaves correlated with the
dimorphic stems. The leaves are arranged in several phyllotaxic
cycles (5/13, 5/8, 3/8 have been recorded) around the radial, ortho-
tropic stems or shoots, and are distichally alternate (1/2 phyllotaxy)
on plagiotropic (lateral) branches.

The first leaves (on orthotropic stems) are long-petiolate and sym-
metrical. The petiole is elongate and thickened at both ends, forming
a long, cylindrical pulvinus below the lamina and a more tubercular,
shorter one at the base; this normal type of petiole facilitates all
kinds of orientation to the blade.

The leaves of plagiotropic branches (the normal ones of the mature
tree) are short-petiolate and asymmetrical. The petiole is with very
few exceptions reduced to the thickened part of the cushions. The
blade has more or less markedly unequal halves, especially at the
base, which may be extremely asymmetrical.

The blades are simple and pinnatinerved, thick-coriaceous or
chartaceous, with a strong midrib and several alternate, spreading-
ascending, prominent secondary nerves; there are elevated tertiary
nerves. Lesser ones form a small usually conspicuous reticulum.
Often the lowest pair of secondary nerves is somewhat more separated
from the next pair than the others, and may give some impression of
a trinerved base; sometimes there are one or two stronger developed
tertiary nerves, giving some appearance of a 5-nerved or 7-nerved
base, but usually the main costa is much stronger than the secondary
nerves and these are thicker than the tertiary, so that the mainly
pinnate arrangement is always clear. In some cases, as in the primary
leaves of 7’. bicolor, the lateral lower nerves are more clearly arranged
so as to show a 5-7-nerved base, and some botanists describe them as
palmatinerved.

The shape of the blades is often ovate, obovate-elliptic-oblong,
or lanceolate, and usually acuminate at apex and obtuse or rounded
at base, but there is a great deal of variation from one species to
another. The margin is basically entire, but sometimes slightly
sinuate or broadly dentate in adult leaves; the primary leaves may be
coarsely dentate in the upper half. Indument is present except in
a few species; most species have a more or less dense tomentum on the
underside, which may be composed of one, two, or three different kinds
or sizes of stellate hairs. This tomentum may cover the whole lower
surface of the leaves entirely or may cover the areoles between the re-
ticulum leaving all or part of the venation glabrous. The different
kinds of hairs and their distribution supply good taxonomic characters.

INFLORESCENCE (Fig. 2).—The inflorescence is of the definite type.
In some cases it may be a well-branched dichasium as in 7’. bicolor, but
generally, the dichasium is totally or partly reduced to a monochasium

CUATRECASAS—CACAO AND ITS ALLIES 419

Ficure 2.—Inflorescence in Theobroma: a, caulinar inflorescence of T. bernouilli subsp.
capilliferum (Cuatr. 16160). 8, caulinar inflorescence in 7. glaucum (Cope & Hol. 118).
c, diagrammatic terminal inflorescence branch in T. cacao. D, detail of sympodia.
branch of inflorescence in T. cacao, diagrammatic, after Stahel. &, inflorescence of
T. cacao, diagrammatic, after Stahel. F, G, detail of inflorescence in 7. cacao, dia-

grammatic, after Stahel. u, inflorescence of T. bicolor (Klug 2021). 1, diagrammatic
inflorescence of T. bicolor after Stahel.

420 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

PAL Hana
oo: mew itive, .

a wei Byte
SSA

\ ee

eel |

/

4uzravs
() HR arAzZe

[Ficure 3]

CUATRECASAS—CACAO AND ITS ALLIES 4?1

of the cincinnate type. Frequently the branches are very short,
forming nodose, articulated sympodia, in which the internodes are
hardly noticeable and the bracts appear almost imbricate. Usually
the bracts are alternate and the fertile terminal branchlets or peduncles
end with 3 bracteoles and one pedicel, the 3 bracteoles corresponding
to a theoretical terminal dichasium which develops only the central
flower. According to Stahel the peduncles have bilateral, the pedicels
radial structure.

The inflorescences may be axillary, in young branchlets, but more
often are originated on short, woody branchlets on the trunk and
branches; these perennial branchlets form irregular tubercles, some-
times very protuberant, which may form woody branchlets up to
several centimeters in length, producing flowering cymes at their
ends. The flowers are always pedicellate, the pedicels being relatively
long, longer than the reduced branchlets of the cymes.

Catyx (Figs. 3, 4).—The five sepals are valvate and may be almost
free and spreading at anthesis or united from one fourth to one half
or more of their length; the lower united part is cupular, the free parts
are patulous-reflexed at anthesis but finally the whole calyx becomes
reflexed, exposing the inner surface. In some instances, the sepals
unite 2 by 2 simulating a calyx of 3 lobes, two of them twice as broad
asthethird. In thesection Andropetalum the sepals usually are united
by three and two and together form a two-lobate cup. The calyx
is persistent and its remains may often be seen below premature
fruits.

In most cases the sepals are tomentose outside with abundant stel-
late, ochraceous or ferrugineous hairs, but they may also be puberulous
or glabrous, as in species or forms of the sections Telmatocarpus and
Theobroma. In the latter multicellular, glandular, stipitate trichomes
are present. The upper or inner surface of sepals is often glabrous
or may be more or less pubescent. The inner margin always has a

Ficure 3.—Calyx and aestivation in Theobroma: a, B, calyx of T. grandiflorum (Cuatr.
25801) with sepals united by pairs appearing to be trimerous. c, calyx of 7. cacao
(Cuatr. 26004), the sepals spreading, very shortly united at base. pv, E, calyx of T.
nemorale with semispreading and semireflexed sepals, united more than 14, F, G, calyx
of T. chocoense, with reflexed sepals unequally united in }4-)4 their length; the basal
glandular papillas very conspicuous. u, diagrammatic long. section of bud in T.
cirmolinae showing the relative position of flower parts in section Glossopetalum; at
the left side the folded petal (the alternating staminode cut away), at right the staminode
(the alternating petal cut away). 1, globular bud of T. chocoense with valvate aestiva-
tion, pedicel, bracteoles and peduncle apparent (X 2). J, ovoid, elongated bud of
T. velutinum (X 2). x, petals in bud showing the contorted aestivation in T. velutinum
(Benoist 161), X 5. 1, diagrammatic long. section of bud in T. cacao showing the rela-
tive position of flower parts in sections Oreanthes, Theobroma, and Rhytidocarpus.

422 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

aCK

CH. REIN

Figure 4.—a-c, three types of floral diagrams in Theobroma: A, sections Glossopetalum
Andropetalum and Oreanthes (part); B, sections Theobroma, Rhytidocarpus and Oreanthes
(part); c, section Telmatocarpus. p, basic diagrammatic representation of vascular
bundles in the flower of the Byttnerineae; the marginal vessels (bundles) derive from the
vascular branches directed to the petals; the fertile stamens are epipetal, the staminodes
episepal; p, vascularization of petal in continuous line, after Gazet du Chatelier slightly
modified, , vascular bundles in flower of Theobroma (sect. Glossopetalum); bundles
in stamens branching shortly above the base; adapted from Gazet du Chatelier. F,
section of an ovary in Theobroma (X 20). 6G, gynoecium in T. cirmolinae (X 5). 4,
flower in anthesis with spreading sepals in 7. sylvestre (Ducke 7882), X 2. 1, flower in

CUATRECASAS—CACAO AND ITS ALLIES 423

narrow band of extremely minute and dense, whitish, stellate hairs
which join the sepals before anthesis. At the base of the sepals,
inside, there are commonly glandular, stalked trichomes, which may
be very scarce but often are numerous and dense, forming a ring
outside the place of petal insertion. These glandular trichomes are
lacking in some species or very rare and scattered above on the sepals.

Corouua (Figs. 3, 4)——The 5 petals are free and uniform; their
special feature is that they are strangulated into two very different
parts united by a narrow joint. The lower part is cymbiform and
erect, rather carnose and rigid, usually 3—7-(or 1)-nerved and has the
appearance of a hood rounded at the top; in fact, it represents the
claw of the petal, and because of its appearance, it is called the
“hood” (cucullus). The upper part of the petal (the lamina) is flat,
varying in shape from oblong to elliptical or discoid, membranaceous
or very thick, yellow, red or purplish. It is almost sessile and directly
articulate to the apex of the hood or may be supported by a narrowly
laminar pedicel, which is its basal extension ; in some species the lamina
is lacking or is almost reduced to the pedicellar extension (T. mam-
mosum). The petals are dextrorsely contorted in estivation, the
laminae being erect when directly articulated to the hood or horizontal
and reflexed through the folding of the pedicel. In anthesis the
laminae are erect.

Anprorcium (Figs. 3, 4).—This is formed by two verticils, which
are connate in a tube at the base. The sterile outer whorl has 5
petaloid staminodes, which are subulate, oblong, or obovate. and
usually very showy with the same color of the corresponding petal-
laminae. They may be erect or reflexed in estivation, and erect,
spreading like a star, or reflexed in anthesis; they are thick-membra-
naceous or carnose and firm, or when subulate or narrowly lanceolate
they may have a thick, carnose midrib; they may be glabrous, but
more commonly are hairy or covered by minute, muricate trichomes.
The inner whorl consists of 5 fertile stamens, with thick filaments
which are connate to the tube except for a short (1-3 mm.) free part;
the apex is 2- or 3-furcate and each short branch bears an anther. The
filaments are spreading and curved and, being opposite the petals,
each anther is concealed in the cavity of its corresponding opposite
petal-hood. The anthers are 2-celled, and each cell, ellipsoid or almost
globose, is unilocular and opens by a longitudinal slit.

The 2- or 3-antheriferous stamens of Theobroma have been treated by
some botanists as a result of the coalescence of two or three original

anthesis with reflexed sepals, spreading petals and erect staminodes in T. cirmolinae
(Cuatr. 15336), X 2. 4, flower of T. cirmolinae (Cuatr. 15336) initiating anthesis in
apical view (X 2).

680-695—64——4

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

424

TT

ae

‘on

f
’
ul
t
'
,
i
'
'
'
t
‘
'
t
!
1
1
1
'
1
4
1
‘
1
’
'
i)

[Ficure 5]

CUATRECASAS—CACAO AND ITS ALLIES 425

stamens. Nevertheless, I consider them to be bifurcate or trifurcate
original stamens. The anatomical works of Gazet du Chatelier (1940,
p. 278) prove this assertion; at the base of the flower, 5 vascular
bundles proceed to the five staminal filaments, these bundles being
forked above to serve the two short branches of the filament in
T. cacao.

Gynorcium (Figs. 3, 4).—This is of the coeno-syncarpic type,
superior, with carpels opposite to petals. The ovary is 5-celled with
axile placentation and many ovules in two rows in each cavity; it is
ovoid or ellipsoid, more or less markedly 5-ridged and furrowed,
densely stellate-tomentose, or rarely glabrous or covered by stipitate
glands.

Stylodes 5, free or more or less adherent to one another, simulating
a single style, glabrous, usually about twice as long as the ovary, thin
and ending in a punctiform stigmatic apex. The ovules are anatropous
with dorsal raphe and two integuments.

FruiT AND SEED (Figs. 5-7).—The fruit is almost baccate or sub-
drupaceous and indehiscent, the various types differing in the firmness
of the pericarp and in the shape. Almost always there can be dis-
tinguished three layers in the pericarp. In the sections Glossopetalum
and Andropetalum, the fruits are externally rigid, hard, the epicarp
being woody, about 1 to 2 mm. thick, with an outer tomentose epiderm;
the mesocarp is fleshy, differing little in color and firmness from the
adjacent endocarp; the inner surface of the latter is a thin but com-
pact membrane; sometimes, the whole endocarp is reduced to this
membrane. When the fruit is ripe the carnose inner layers decay or
dry, and shrink, but the rigidity of the epicarp maintains absolutely
the size and shape of the fruit, keeping the loose seeds inside if they
have not been accidentally liberated. In the section Lhytidocarpus,
the mesocarp is the rigid, woody layer; the epicarp being thinner and
carnose, although also with an outer tomentose epiderm; the endocarp
is also carnose, and also provided with an inner membrane. In the
section Oreanthes, typified by JT. speciosum, the whole pericarp is
5 to 6 mm. thick; the innermost layer, the endocarp, although very thin,

Ficure 5.—Fruit sections of Theobroma species: a, long. section of fruit of T, simiarum
(Cuatr. 26515A), X %. B, transection of pericarp of a, natural size. c, transection of
pericarp of T. grandiflorum, X 1. pb, transection of pericarp of T. bicolor, X 1. £,
transection of pericarp of T. gileri, X 2. ¥, transection of pericarp of T. speciosum,
x 1. G, transection of pericarp in T. cacao cultivar ‘‘cundiamor” (Cuatr. 26492), X 1.
H, transection of pericarp in 7’. cacao {ma. pentagonum (Cuatr. 26540), X 1. 1, transec-
tion of pericarp in 7. cacao cultivar “‘lagarto” (Cuatr. 26004), X 1. J, transection of
pericarp in Herrania cuatrecasana (Cuatr. 25793), X 2. ed, epiderm; ep, epicarp; m,
mesocarp; en, endocarp; i, interior pelicule limiting the endocarp inside; pu, pulp;
5, seed.

496 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Ficure 6.—Seeds of Theobroma, natural size: a-E, T. grandiflorum (Cuatr. 25780T): a,
front view of seed stripped from pulp; 8, c, p, side and front view of embryo; £, seed
surrounded by its pulpy layer. F-n, 7. simiarum (Cuatr. 26515A):F, seed; c, embryo;
H, embryo in apical view. 1, J, T. mammosum (Cuatr. 25791): seed in lateral and apical

CUATRECASAS—CACAO AND ITS ALLIES 427

is woody and rigid; the mesocarp is thick and fleshy; the epicarp is also
woody but thin and less compact. When the mesocarp dries or decays,
the epicarp shrinks slightly, becoming more or less rugose. In the
section Telmatocarpus the innermost layer, the endocarp, is the most
compact and rigid, although thin; the epicarp is coriaceous and thin
and the mesocarp is thick and fleshy with strong bundles which build
protruding ridges and veins covered by the epicarp. Finally, in
section Theobroma (Cacao) the fruit is almost baccate because the
whole pericarp is carnose; the inner membrane and the outer epiderm
may be very firm but the whole pericarp decays easily; it can also dry
out, being then coriaceous. Usually the three layers are conspicuous,
the epicarp carnose and thick, with glabrous outer epiderm, the
vascular mesocarp papyraceous, rigid, thinly woody, and the endocarp
carnose, more or less thick, with an inner pellicle; in some forms of
cacao the mesocarp may be reduced to a very thin or discontinuous
layer or to isolated vascular bundles (fig. 51); rarely the endocarp is
reduced to the inner pellicle (fig. 5, forma pentagonum). Usually,
the dorsal vascular bundles of each carpel develop into transverse
membranes within the fleshy epicarp connecting with the mesocarpial
layer. Gummy sacs are always present in several parts of the pericarp.

The young fruit, as well as the ovary, has five cavities with the in-
cipient seeds arranged in one or two rows in each cavity. At maturity
the cell walls vanish and the seeds with their thick outer pulpy layer
fill the single cavity, arranged usually in five rows.

The shape and size of the fruit are variable and they are, com-
bined, specific characters except for cultivated cacaos. The fruits
range between 6 and 35 cm. long by 5 to 12 cm. broad and may
contain, when ripe, from 16 to 60 seeds. They may be ellipsoid,
globose, ovoid, oblong, or fusiform, with rounded or attenuate ends,
with completely smooth surface like a potato or marked more or
less with 5 or 10 ridges, or they may be echinate or verrucose. In
all cases they are indehiscent and the liberation of seeds follows the
decay of the shell, which in many cases, as in those with hard, woody
pericarp, may take so long that the seeds have died. The common
natural way of propagation of the seeds is accidental, usually by

view. x-M, T. angustifolium (Cuatr. 25790): x, seed stripped from pulpy layer; 1,
embryo; M, embryo in apical view. n-p, 7. gileri (Cuatr. 26167): nN, seed in apical
view; 0, same laterally; p,embryo. -s, T. speciosum: Q, seed; r, embryo; s, embryo
in transection, showing the folding of cotyledons. t-v, T. bernoutllit subsp. capilliferum
(Cuatr. 17034): 1, seed; vu, embryo; v, embryo in apical view. w-xx, T. cacao fma.
pentagonum (Cuatr. 26004): w, seed; x, embryo; xx, transection of embryo. y, ry, 7’.
cacao fma. pentagonum (Cuatr. 26540): y, transection of seed with episperm and pulp;
yy, seed. 2, 2z, T. cacao cultivar “cundiamor” (Cuatr. 26492): z, seed; 2z, transection

of seed with episperm and pulp.

428 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

animals (mostly monkeys, but also squirrels, rats, and other animals)
which break the pericarp in order to suck the pulp surrounding the
seeds, which may be expelled later in other places, thus disseminating
the seeds (fig. 5).

The fruits, which are commonly called pods in English, mazorcas
in Spanish and cabosses in French, may stay on the tree or fall down
after maturation; in the latter case, they may fall with the peduncle
(e.g., T. bzeolor) or without it (as in 7. grandiflorum). Precise obser-
vations in many species are wanted.

The seeds are ellipsoid, ovoid or amygdaloid, more or less irregularly
compressed, complanate, or terete, and range from 15 to 40 mm. long
and 10 to 22 mm. broad; the integuments or skin form two strata
with an additional outer, thick, gelatinous-pulpy layer surrounding
them. The testa is generally thick and subcoriaceous, with an
external epiderm covered by a thick cuticle, a thick layer of poly-
hedric and mucilaginous cells, and an inner layer of sclerosed cells.
The inner tegument is a thin membrane of several layers of thin-
walled, complanate cells. Inside, the large embryo is composed of
two large, thick, strongly folded and corrugate cotyledons and a
straight, rather thick, terete radicle, the plumule being scarcely
developed. The endosperm at maturity has the shape of a very fine
pellicle, containing scattered cells with calcium oxalate, covering the
embryo outside and between its foldings. The cotyledons possess
an epiderm, often with scattered, stipitate glandular trichomes and
a main cellular tissue rich in starch, fat, aleurone, tannoid and alka-
loidal substances, among these the important theobromine compounds.
In most species the cotyledons are white, but in a few they are violet,
reddish, purplish, being stained by tannins (figs. 5, 6). Germina-
tion may be either epigeous or hypogeous according to the species
(fig. 7).

The pulp surrounding and united to the seeds is white, yellowish,
or yellow, and often sweet and aromatic and palatable, but it may
be also scentless and tasteless to men; it is, however, always appreci-
ated by animals, which hunt the pods, extract the seeds to suck the
pulp, thus disseminating them.

In appropriate conditions the pulp suffers a fermentation process
which separates it from the seed; during that fermentation, very well
known in the case of T. cacao, chemical changes take place inside the
embryo developing a special aroma. In the industry of cacao torre-
faction completes the desired effects of fermentation.

Premature fruits keep their viability for some time provided they
are protected against loss of humidity and stored under suitable
temperatures (20-25° C.); when ripe, the seeds become immediately
ready for germination; they may germinate inside the pods. The

CUATRECASAS—CACAO AND ITS ALLIES 429

nt
x
u
g
uy
&
z
w

Ficure 7.—a, Epigeal germination in Theobroma bernouillii subsp. capilliferum (Cuatr.
17350A). 3, Hypogeal germination in T. grandiflorum (Cuatr., Cope & Bart. 25780A).

germinating power of Theobroma seeds lasts only for a short time, a
few weeks; observations on 7’. cacao have shown a maximum extension
time of viability to about three months, when carefully preserved in
their pods or under special protection. They are extremely sensitive
to the degree of humidity, which has to be kept high, and to low
temperatures. Recent experiments in Turrialba showed that cacao
seeds could not resist low temperatures for even a short time; seeds
exposed for 16 minutes at 8° C. lowered the germinating capacity to
6%, and 4 minutes exposure at 2° C. inhibited germination almost
completely (Hunter, 1959; Hunter & Boroughs, 1961). This may
be the explanation why cacao seeds lose their germinating power

430 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

when transported by airplanes at high altitude (Hunter, personal
communication).
Ecology

Theobroma is a tropical American genus restricted to the lower
tree-story of the evergreen rain forests. The species demand a high
degree of mean annual temperature with narrow oscillations, a con-
stant high humidity, and protection (shade) against direct radiation
and evaporation. Several species are often found at the edges of
rivers or marshes in more or less temporarily flooded areas; others
always grow on elevated, drained places. They like relatively rich
and neutral soils. These conditions are found only in the warm, wet,
forested, equatorial regions between latitudes 18° North and 15°
South, with temperatures of 20° to 30° C., with a minimum of 16° and
maximum of 40°. <A few species grow at higher altitudes up to 1250
meters, being able to withstand minimum temperatures of 14° and
even 12° C. Where Theobroma is at home, the rainfall is from 2000
mm. to 8000 mm. annually or even higher, and is more or less evenly
distributed throughout the year. Theobroma does not resist even
short dry seasons without the protection of dense shade and local
humidity. In cultivation T. cacao can endure less humid climates,
and more open lighter spots especially when irrigated, and somewhat
lower temperatures than the normal optimum. So the area of culti-
vated cacaos may extend far above 20° North and below 20° South
of the Equator. Not only the Theobroma trees but also the seeds are
highly specialized to the humid equatorial ecological conditions. It
is known that the seeds keep for a very short time their capacity for
germination, which often takes place inside of the pod; only under
high humidity and optimum temperatures can they maintain their
viability (see above).

Geographic Distribution

The genus Theobroma is a typical neotropical genus, distributed
throughout the rain forests of the western hemisphere between
latitudes 18° North and 15° South. Some species have a broad range
of distribution, like 7. subincanum, which is spread throughout the
Amazon-Orinoco basins, being one of the most ancient of the genus.
The elevation of the Andes in the carly Tertiary separated populations
of Theobroma previously widespread before, favoring speciation
through isolation. Vicarian species separated by this way are T.
subincanum (east of the Andes) and 7. hylaeum (west) ; T. microcarpum
(east) and 7’. gilert (west). The complexity of the mountains of the
northern part of Colombia through Central America was also an iso-
lating factor which favored speciation in that part of the hemi-
sphere where regional or local endemics are present. Maps 1 and 2
are self-explanatory.

CUATRECASAS—CACAO AND ITS ALLIES 431

50° 40°
/o”
W
— o°
T ”
A.
~
A 20°

Map 1.—Geographical areas of Theobroma sect. Oreanthes: 2, T. sylvestre; 3, T. speciosum;
4, T. velutinum; 5, T. glaucum; T. bernouillii: 6a, subsp. bernouillit; 6b, subsp. as-
clepiadiflorum; 6c, subsp. capilliferum.

a
f

Map 2.—Geographical areas of Theobroma sect. Glossopetalum (10-21) and sect. Andro-
petalum (22). 10, T. angustifolium; 11, T. cirmolinae; 12, T. stipulatum; 13, T. choco-
ense; 14, T. simiarum; 15, T. grandiflorum; 16, T. obovatum; 17, T. subincanum; 18,
T. hylaeum; 19, T. nemorale; 20, T. sinuosum; 21, T. canumanense; 22, T. mammosum.

432 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Pollination

Although transportation of pollen by the wind has had some ac-
ceptance, it seems that only pollination by insects has been proved.
Works by Harland, Stahel, Posnette, Saunders, Cope, Uzel, Jones,
and Pound, have demonstrated that several kinds of flying and crawl-
ing insects are involved in pollen transportation, among them thrips,
ants, midges, and aphids. Experiments by Cope proved that Frank-
liniella parvula Hood and Wasmannia auropunctata Rog. were mostly
responsible for pollen transportation in Trinidad. Saunders found in
Costa Rica that Morctpomyia midges performed pollination in cacao.
The aphid Tozoptera aurantii Fouse is also recorded as a pollinating
agent.

Relationships

The basic features of Theobroma are: Flowers bisexual, pentamer-
ous; sepals valvate; petals strangulated, contorted in bud with
cymbiform, cucullate lower half; 5 stamens opposite to’petals*and 5
evident, alternating staminodes united in a short basal tube; stamens
shortly 2-3-branched; anthers 2-celled; ovary superior, 5-celled;
ovules many, anatropous with 2 integuments; fruit subdrupaceous or
subbaccate; seeds with pulpy envelope; cotyledons folded, corrugate;
evergreen trees with dimorphic branching and dimorphic, entire,
alternate leaves. This diagnosis places the genus in the family
Sterculiaceae (fig. 8).

Theobroma exhibits a unique set of characters which makes it a
very “natural”? genus. However, some of the outstanding features of
its floral structure are also shared by other genera, the most con-
spicuous being the cucullate or concave lower part of the petals which
define the tribe Byttnerieae DC., and determine the close relation-
ships between its members: Byttneria, Ayenia, Rulingia, Commer-
sonia, Theobroma, Guazuma, Herrania, Abroma, Glossostemon, Scapho-
petalum, and Leptonychia. In most of these cases the similarity with
Theobroma in the flower structure (petals, androecium-tube, stami-
nodia, and position of anthers) is so obvious that it was noticed since
early times. The first historical association of Theobroma to another
genus was by Linnaeus who joined it with Guazuma under Poly-
adelphia Pentandria. Lamarck (1785) was the first to make a family
associating Theobroma with Abroma, Guazuma, Ayenia, Byttneria, and
Kleinhovia. Jussieu (1789), associated Theobroma with Abroma,
Guazuma, Byttneria, Dombeya, Assonia, Pentapetes, and Melhania in
sectio V [bis] of ordo XIV. Kunth (1823) was the first to estab-
lish critically the main groups of the Sterculiaceae, one of them the
Byttneriaceae verae including Theobroma, Guazuma, Abroma, Glosso-

CUATRECASAS—CACAO AND ITS ALLIES 433

stemon, Byttneria, Ayenia, and Commersonia. This grouping was
basically followed by DC. (1824), Endlicher (1840), although he
separated the Sterculiaceae from Byttneriaceae, Baillon (1873) in his
series Byttneriées, Bentham & Hooker (1862), who enlarged the
family to 7 tribes, and Schumann (1890) who enlarged it to 8 tribes.
The latter botanist, who made an outstanding contribution to the
comparative morphology and taxonomy of the whole family, did not
alter the concept of the Byttnerieae DC. as presented by Bentham and
Hooker. Recent workers, like Gazet du Chatelier (1940), who made
broad comparative anatomical and morphological studies in the
Sterculiaceae, found good reasons to keep Schumann’s basic taxonomic
approach.

Bentham and Hooker divided the tribe artificially in two groups
which were named by Schumann Theobrominae and Byttnerinae, with
respectively 2-3-antheriferous and 1-antheriferous stamens. On the
other hand, the four genera of Byttnerinae differ from Theobroma also
because Byttneria has spirally convolute cotyledons, short, dentiform
staminodes and linear, rather thick, petal-laminae; Ayenia has very
long, linear petal-claws, trilocular anthers, and spirally convolute
cotyledons; the Old World Commersonia and Rulingia have a pitcher-
shaped petal base and flat cotyledons. From the other Theobrominae
genera, Theobroma is distinguished by the special structure of the
petals, staminodia, and vegetative system; the Persian genus Glos-
sostemon is a shrub with hairy, dentate leaves, ovate-oblong petals,
concave at base, and with many short stamens connate to the basal
part of the staminodes; the Old World Leptonychia differs by its short,
squamiform petals, fertile stamens with filaments much longer than
the staminodes, and flat cotyledons; the west African shrub Scapho-
petalum has exappendiculate petal hoods, a campanulate androecium
with shortly triangular staminodes and sessile 3-grouped anthers; the
tropical American Guazuma differs, besides in the fruit, by the long,
bifid petal appendages, the spirally convolute cotyledons, and the
vegetative structure, the leaves being serrate; Abroma, an oriental
genus spread from eastern India through the Pacific islands to Aus-
tralia, is similar to Theobroma in the floral arrangement but usually
has more developed petal laminae, shorter petaloid staminodes,
subsessile anther groups, flat cotyledons, a different vegetative habit,
and usually cordate, more or less lobate, hairy leaves. Moreover,
all genera mentioned of the Byttnerinae differ from Theobroma by
their capsular, generally dehiscent fruit. Only Theobroma and
Herrania in the tribe have an indehiscent baccate or subdrupaceous
fruit. For this reason, Schumann united them, calling the latter
section Herrania, an arrangement adopted by other botanists, such
as Pittier and Ducke. Nevertheless, Bernoulli, the monographer who

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

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CUATRECASAS—CACAO AND ITS ALLIES 435

went deeply into the genus, and Chevalier in his revision consider
Herrania and Theobroma different genera. R. E. Schultes followed
the same line in monographing, after long experience, the genus
Herrania with 17 species. They are undoubtedly two well-defined,
independent genera.

Herramia, Abroma, Guazuma, and Byttneria surely are the genera
closest to Theobroma. Chromosome number and palynology help to
determine relationships. The chromosome number is identical for
Theobroma and Herrania, 2n = 20 but it is 2n = 16 for Guazuma and
2n = 14 in Byttneria (Cristdébal) ; data for Abroma not available. The
pollen grains are suboblate in Theobroma, prolate in Herrania, prolate-
spherical in Guazuma, and oblate in Abroma.

Because of the similarity of the fruits and the confusion which had
prevailed in the past between Theobroma and Herrania, their differ-
ences are summarized here as follows:

Theobroma. Stem sympodial, with 3-5-verticillate branching;
branches dimorphic; branching copious; leaves dimorphic; leaf-blades
simple, entire; petal lamina more or less rounded to lanceolate, not
more than twice as long as the hood, erect or inflexed and contorted
in aestivation; pollen grains suboblate; cotyledons strongly folded and
corrugate; fruit usually smooth or rugose, angular, seldom strongly
costate; staminal filament symmetrically and shortly 2- or 3-furcate
at apex.

Herrania. Stem monopodial, unbranched, with apical growth;
leaves uniform, 5-9 digitate, long-petiolate, in a terminal, lax cluster;
petal lamina many times longer than the hood, linear, pendulous
in anthesis, involute in aestivation; pollen grains prolate; cotyledons
thick, flat or very slightly folded; fruit usually strongly costate;
staminal filament usually asymmetrically parted in two branches,
one 1-antheriferous, the other 2-antheriferous (fig. 9).

Evolution

The question of how the genus Theobroma may have originated
is a speculative matter on which botanists like Schumann (1886)

Ficure 8.—Genera related to Theobroma: a-H, Guazuma: a, articulated and hooded petal
with bifid appendix, from inside; B, same from outside; c, same in lateral view, X 5;
p, androecium, X 5; £, fertile stamen, X 20; F, sepals, X 2; G, bud; n, gynoecium, X 5
(G. tomentosa, Cuatr. 22942). 1, J, Byttneria: 1, articulated petal from inside, X 10;
J, same laterally. x, flower, X 10. 1, gynoecium, X 10, M, androecium, * 10.
nN, detail of the anther, X 20. 0, sepals, from inside and outside, X 5. p, bud, & 5.
q, carpel of fruit from inside, outside and apical view, X 2 (B. arguta, Cuatr. 8226).
R-Z, Abroma: R, articulated, hooded petal, < 2; s, same in lateral view; T, hood from
inside and v, from outside, X 5; v, sepal, X 2; w, pistil, X 5; x, androecium surround-
ing the gynoecium, X 5; y, base of staminode with laterally attached stamens, X 5;
z, biantheriferous stamen, X 10 (Abroma augusta, Sulit 18880).

436 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

[FicureE 9]

CUATRECASAS—CACAO AND ITS ALLIES 437

did not want to take a stand. Edlin (1935) who developed a theory
on the evolution of the Malvales, considers the family Sterculiaceae
limited to the tribe Sterculieae, all other groups forming the family
Byttneriaceae; he considers the stamens the result of union. Gazet
du Chatelier (1940), after a detailed examination of the Sterculiaceae,
came to the conclusion that there was an original unknown type from
which were derived two diverging groups (subfamilies or families),
the ‘‘Eriolaenées” and the ‘“Buettneriées,’’ but he did not go much
further in his speculative evolution; the stamens of Theobroma are
considered as branched by him.

All the genera of the Byttnerieae are similar and probably originated
at the same time evolving from an original unknown type; they
diversified their flowers and the leaves, probably through mutations
aided by geographical and ecological barriers. The parts have
evolved independently, e.g., Rulingia has undivided fertile stamens,
a more ancient character, but pitcherlike petals, a more evolved one.
Conversely, Leptonychia has simple, more primitive petals but
exhibits branched stamens, a more advanced character. Byttneria
and Guazuma have elaborate, advanced petals but less developed
staminodes; Commersonia, as well as Abroma and Theobroma, have
more advanced petals and staminodes than the other genera.
Scaphopetalum is an example of a more advanced type due to the loss
of the petal lamina and reduction of the staminodes. Even if we can
attribute primitiveness or the contrary to some characters, it is not
possible to draw a lineal series of genera according to antiquity.
Nevertheless, I would venture to say that Theobroma and Herrania
belong to the most modern in the Byttnerieae because of the structure
of the fruit, with thick and partly or totally carnose pericarp and
delicate, short-lived seeds. These may be characters acquired in
the process of evolution and kept by their adaptation to the extremely
hot and humid ecological conditions of the tropical American forests.
It also seems to me that Herrania is a more evolved genus in regard
to the flower, but not in the simplicity of the monopodial, juvenile-

Ficure 9.—a-H, Herrania pulcherrima v. pacifica (Patifio 23): a, articulated and hooded
petal, X 5; B, segment of androecium with a staminode and the adjacent stamens,
X 5;c, bud, X 5; p, stamens, X 10; £, gynoecium, X 5; F, seed, X 1; G, transection of
seed, X 1; H, embryo, X 1. 1-x, H. cuatrecasana:1, embryo; j, seed; x, transection
of embryo, X 1. u-R, Theobroma bicolor (Garcia B. 11178): 1, petal from inside,
outside and laterally, X 5; M, androecium, X 5; N, fertile stamen and part of staminal
tube, X 10; 0, sepal from inside and outside, X 2; Pp, gynoecium, X 5; Q, styles, X 5;
R, bud and pedicel, X 2. s-y, T. sylvestre (Ducke 7882): s, petal from inside and
laterally, X 5; 17, androecium, X 5; u, stamen, X 10; v, bud, X 2; w, gynoecium, X 5;
y, sepal from outside and inside, * 2,

438 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

like, unbranched stems, in the digitate, loosely clustered leaves, and
inseparable, hypogeous cotyledons, due probably to the stringent
ecological conditions of the shadowy underlayer of the humid tropical
forest. Aside from Herrania, the sections of Theobroma (Glossopetalum)
with pseudoterminal growth, may be more primitive than the ones
(Oreanthes, Rhytidocarpus, Theobroma) which have lost the axillary
buds of the jorquette branches, necessitating lateral shoots to continue
erowing. The section Theobroma may be more evolved than the
others on account of the 5-branching system and carnose fruits. The
section Telmatocarpus may be more advanced in another direction
because of the reduction or absence of the petal lamina and the
discontinuity of the vascular, woody system in the pericarp, which is
only partially woody and more vulnerable. The parallelism in evolu-
tion of the sections Theobroma and Telmatocarpus is seen in the
elabrous or almost glabrous leaves, more suited to rain-forest ecology.
The section Rhytidocarpus may be an ancient type with less showy
petals and staminodes, axillary flowers, and a thick-woody pericarp.

No fossils belonging to Theobroma have been recorded.

The geographic distribution does not give any solution to these
questions of evolution because almost all sections are represented at
both sides of the Andes. It seems that the richest region in species
is around Panama and Colombia, where species with a very restricted
area are found, especially if we consider this region extended to
Costa Rica. I feel that Theobroma is a genus with a marvelous set
of characters controlled perhaps by independent genes, which seem-
ingly can combine independently resulting in many different sets of
combinations.

Economic Uses

The seeds of Theobroma are rich in starch (15%), protein (15%),
and oil (50%), for which reason they are considered a substantial food.
Moreover, they have a volatile oil (cacao-essence) which gives an
aromatic flavor and 1.5 to 3% of theobromine, an alkaloid known for
its stimulant properties. Caffeine is also present in Theobroma seeds.
Both alkaloids have been found in the seeds and leaves of 7. bicolor,
cacao, grandiflorum, microcarpum, obovatum, speciosum, sylvestre, and
subincanum (Willaman and Schubert, 1961). The cacao seeds contain
also a red pigment, tanine, and small quantities of malic and tartaric
acids, asparagine, and coline.

It is not necessary to emphasize the economic importance of the
industry in cocoa and chocolate. Most of the wild species of cacao
are often used by the natives, who suck the pulp or prepare refreshing
drinks with the pulp. The seeds of most species may serve for the
preparation of chocolate, but actually only one species has become

CUATRECASAS—CACAO AND ITS ALLIES 439

commercially important in this respect, T. cacao, which is the only one
widely cultivated. An important secondary product from cocoa
seeds is the cocoa butter extracted by pressure during the process of
making chocolate. Cocoa butter is important in cosmetics and
pharmaceutical industries. Cacao extracts and theobromine are
important in medicine because of their cardiotonic and diuretic
properties.

The wood of several Theobroma species is important in local con-
struction and because of its toughness and strength is very much used
in the manufacture of tools and parts of instruments and machines.

Anatomy of the Wood

CONTRIBUTED BY WILLIAM L. STERN!

This study of the wood of Theobroma is based largely on microscope
slides borrowed from the S. J. Record Memorial Collection of woods
at Yale University and from the wood collection of the Imperial
Forestry Institute at the University of Oxford in England.? It is
regrettable that among these slides, only 9 species were present
(table 1). However, the description of the wood probably represents
a fairly good outline of at least the qualitative aspects of the micro-
scopic structure, and is sufficiently complete to enable comparisons
between Theobroma and other genera to be made.

It is evident from this brief study that noticeable variation occurs
in the wood anatomy of different specimens of the same species. In
this regard it is interesting to note that Record and Hess (19438, p.
517) were impressed with the structural variation in rays in different
parts of the same specimen in their study of the woods of Sterculiaceae.
As a whole, however, the wood of Theobroma species does not present
any characters of significant anatomical import which would enable
us to separate them on anatomical grounds. Chattaway (1937) also
found this to be true of the genera she studied in her investigation of
the Sterculiaceae (sensu Edlin 1935).

1 REFERENCES:
Bailey, I. W. The problem of differentiating tracheids, fiber-tracheids, and libriform wood fibers.
Trop. Woods 54:18-23. 1936.
Bentham, G., & Hooker, J. D. Sterculiaceae, in Genera plantarum. 1:214-228. London, 1862.
Chattaway, M. Margaret. The wood of the Sterculiaceae. I, Specialisation of the vertical wood
parenchyma within the sub-family Sterculieae. New Phytol. 31:119-132, 1932,
——. Ray development in the Sterculiaceae. Forestry 7:93-108. 1933.
. The wood anatomy of the family Sterculiaceae. Phil. Trans. Royal Soc. London, Ser. B-Biol.
Sci. 228:313-366. 1937.
Edlin, H. L. A critical revision of certain taxonomic groups of Malvales. New Phytol. 34: 1-20.
1935.
Metcalfe, O. R., & Chalk, L, Anatomy of the dicotyledons. 1:251. Oxford, 1950.
Record, 8. J., & Hess, R. W. Timbers of the New World. p. 517. New Haven, 1943.
1 I would like to thank Dr. Graeme Berlyn of Yale and Dr. L. Chalk of Oxford for their kindnessin making
slides available for study.

680-—695—64——_5

440 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

The imperforate tracheary elements are all fiber-tracheids; that is,
the bordered pits are smaller in diameter than those found in the walls
of vessel elements in the same species. Bordered pits usually show
extended inner apertures; these may be crossed or not. The wall
thickness varies from thick to thin, sometimes even within the same
species (cf. specimens of 7. bicolor).

Pores are distributed mainly in the solitary configuration on the
transverse section (34-86 percent; average 62 percent); radial mul-
tiples are next in abundance (8—47 percent; average 33 percent) and
pore clusters are least abundant (0-10 percent; average 4.5 percent).
In different specimens of the same species, dissimilarities may occur;
for example, in 7. obovatum (Williams 161), solitary pores account for
86 percent of the pores per field, whereas in T. obovatum (Williams 230),
they account for only 57 percent of the pores per field. Perforation
plates are entirely simple. Vessel element end walls form angles from

45° to 80° with the vertical. Intervascular pitting is alternate. The
TABLE 1.—Specimens examined in anatomical analyses
Species of Theobroma Collector and No. Origin Herb. USw Yw FHOw
voucher No. No. No,
angustifolium Cooper & Slater | Panama Y 10595 | 3502
DC. 242
bernowillit Pitt. Pittier 4105 Panama US 30
bicolor H. & B. For. Dept. Br. British F 5632
Hond. Hon-
H. 2192/29(?) duras
bicolor H. & B. “Ford-Brazil Brazil 22075 | 6998
3977’ (?)
bicolor H. & B. UH. Williams Peru 17804 | 7008
149
bicolor H. & B. L, Williams Peru 18176 | 7007
46
cacao L. “T,, 3225 (via South 5703
Hamburg)” America
cacao L. Vigne 2433 Ghana we ; 6898
)
grandiflorum Ll. Williams Peru 17893 | 7001
(Willd.) 2401
Schum.
microcarpum Krukoff 6203 Brazil US 36510
Mart.
obovatum Ll. Williams Peru F 71232 | 7010
Klotzsch ex 161
Bernoulli
obovatum Ll. Williams Peru F 17263 | 7011
Klotzsch ex 230°
Bernoulli
sylvestre Mart. Ducke 103 Brazil Y 21362 | 7009
subincanum Ll. Williams Peru F 17578 | 7000
art.
subincanum Ll. Williams Peru F 18144 | 6999
Mart.

* Abbreviations from W. L. Stern & K. L. Chambers. The citation of wood specimens and herbarium

vouchers in anatomical research.

Taxon. 9: 7-13. 1960.

CUATRECASAS—CACAO AND ITS ALLIES 441

bordered pits are frequently crowded and their outlines markedly
angular. Other times the pits are rounded to elliptical. Vessel—axial
parenchyma pitting and vessel-ray parenchyma pitting generally
follow the pattern of the intervascular pitting. Occasionally pits may
be elongated or slightly irregular. No deposits or tyloses appeared
in any vessels.

Both uniseriate and multiseriate vascular rays occur in each of
the specimens examined. Multiseriate rays may be up to 20 cells
wide (in 7. sylvestre), but a width of 10 to 15 cells is more common.
Uniseriate rays are much lower in height than multiseriate rays; the
latter range from 30 to 230 cells high. There is often evidence of dis-
sociation of these broad, high rays into lower, narrower rays by the
‘Gntrusive action” of fiber-tracheids while the cells are still in a plastic
stage. Many of the multiseriate rays are characterized by the
presence of sheath cells, e.g., in 7. obovatum (Williams 161); however,
they never form complete sheaths about the rays and are rare in some
specimens. Multiseriate rays are heterocellular, with the multi-
seriate portion comprising procumbent cells, and uniseriate, alate
extensions of 1 to several (15+) upright or square cells; uniseriate
rays are usually homocellular composed of square or upright cells
and sometimes occasional procumbent elements. The ray cells are
commonly characterized by deposits of reddish or yellowish, non-
staining materials.

Axial parenchyma occurs in two dispositions: apotracheal, as
diffuse and/or diffuse-in-aggregates arrangements, and paratracheal,
as vasicentric sheaths 1 or 2 cells wide. Sometimes a ladderlike
configuration is formed on the transverse section by short bands of
axial parenchyma which frequently intercept vascular rays (e.g., in
T. angustifolium).

Storying of tissues occurs in the wood of Theobroma, but in its
most highly developed state, it would have to be considered incon-
spicuous. Where it does appear, it is limited in distribution and
confined to the uniseriate rays. In TJ. microcarpum it was also
observed in the axial parenchyma.

Crystals occur in the wood of all species examined. Generally
they are more frequent and conspicuous in the cells of rays, although
they also occur in axial parenchyma cells of some species. In rays,
only crystals of rhomboidal nature were observed except in 7. micro-
carpum where large druses were present exclusively. Crystals in
axial parenchyma cells are mostly rhomboidal, but in some species
small druses occur.

Discusston.—The most significant anatomical studies of the wood
of Sterculiaceae are those of Chattaway (1932, 1933, 1937). Unfor-
tunately, her work is of limited value as a basis for comparison here,

442 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

for she adopted the restricted view of the family proposed by Edlin
(1935) and confined herself to the tribe Sterculieae. It should be
mentioned that Edlin suggested dividing the Sterculiaceae, as treated
by Bentham and Hooker (1862), into two families: Sterculiaceae, to
be restricted to the tribe Sterculieae, and Buettneriaceae, to contain
all other taxa (including Theobroma). Chattaway corroborated
Edlin’s proposals according to the anatomical findings which resulted
from her studies. Nevertheless, it does not seem to me, judging
solely from her summary of the characteristic anatomical features of
the Sterculiaceae (sensu stricto), that the wood anatomy of Theobroma
(which would be eliminated from Sterculiaceae according to Edlin’s
concept) would preclude its being allied with the species upon which
she reported if we used only anatomical bases. There are only two
apparent anatomical differences between Theobroma wood and that
of Sterculiaceae (sensu stricto): Regardless of statements to the
contrary (Metcalfe and Chalk 1950, p. 251), the imperforate elements
in Theobroma wood are not libriform wood fibers, but fiber-tracheids
with small bordered pits (sensu Bailey 1936). Chattaway describes
corresponding cells in Sterculiaceae (sensu stricto) as libriform wood
fibers. Also, she indicates that the rays in Theobroma woods lack
sheath cells (Chattaway 1932, 1937), ‘but are present in the rays of
all genera of the Sterculiaceae except Heritiera.” I cannot agree
that Theobroma rays are totally devoid of these specialized elements.
Although they are of sporadic occurrence, it is relatively easy to
demonstrate them among the rays in any given tangential section.

In summary we can say that Zheobroma woods are characterized
by fiber-tracheids with small bordered pits, mostly solitary pores,
simple perforation plates, alternate intervascular pitting, both homo-
cellular uniseriate rays and heterocellular multiseriate rays with
sheath cells in the same species, both apotracheal and paratracheal
axial parenchyma in the same species, and crystalliferous deposits
which are most abundant in the cells of ray tissue. Storied structure
is present to a limited degree and is confined largely to the uniseriate
rays. Although anatomy is variable within specimens of a given
species, it is not consistently variable to allow for the division of the
genus on anatomical grounds. In my opinion, the wood anatomy of
Theobroma does not differ significantly from that in Edlin’s Stercu-
liaceae as delineated by Chattaway.

Pollen Morphology of Theobroma and Related Genera

CoNTRIBUTED BY G. ERDTMAN 3

Theobroma L. (fig. 10): Pollen grains 3-colporate, peritreme,
suboblate (about 15-22 17.5-25 y).

3 Palynological Laboratory of the Swedish Natural Sclence Research Council, Stockholm.

CUATRECASAS—CACAO AND ITS ALLIES 443

Species investigated: 7. angustifolium Moc. & Sessé (Pittier s.n.):
about ?X23.5 uw; T. bernouilli Pitt. (Pittier 4105): about ?22 u;
T. bicolor Humb. & Bonpl. (Klug 2021): about 16X18 yu; 7. cacao L.
(Calderon 107): about 15X17.5 yw; T. glaucum Karst. (Holliday &
Cope T-118): about 2224.5 yu; T. grandiflorum Schum. (Archer
7549): about 1922.5 u; T. microcarpum Mart. (Archer 7551): about

16.5X21 yw; T. speciosum Willd. var. coriaceum Huber (Rusby 647):
about 2224.5 u.

,

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i 1

1

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‘ i

Ficure 10.—Palynograms, X 1500: a, Herrania pulcherrima v. pacifica Schult.; p, Glos-
sostemon bruguieri DC.; c, Guazuma polybotrya Cav.; p, Theobroma glaucum Karst.;
E, Abroma augusta L.; G. Erdtman & A. L. Nilsson, original.

444 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Examples: 7’. glaucum Karst. (Holliday & Cope T-118): pollen
grains 3-colporate, peritreme (amb circular), oblate spheroidal (about
2224.5 uw). Apocolpium diameter about 18 u. Colpi narrow, about
18 w long. Ora about 3.5ubroad. Exine about 1.5 thick. Sexine
as thick as nexine or slightly thicker, tectate. Tegillum distinctly
undulating. The waves of the tegillum are smoother than those of,
for example, Herrania pulcherrima var. pacifica, but nevertheless they
impart to the pollen surface a reticuloid pattern with muroid ridges
(supported by one or two rows of endosexinous bacula) separated by
luminoid depressions (diameter up to 3 uv). The tegillar bottom of the
latter seems to be supported by stray baculoid rods.

T. microcarpum Mart. (Archer 7551): pollen grains 3-colporate
(amb circular), suboblate (about 16.521 4). Apocolpium diameter
about 13.5 4. Colpi narrow, about 10 u long. Ora lalongate (about
153 uw). Exine about 1.6 » thick. Sexine thicker than nexine,
tectate, undulating (waves not as smooth as in 7. glaucum). Retic-
uloid pattern much as in 7’. glaucum, with more or less irregular
luminoid areas (longest axis up to about 3.5 y).

T. speciosum Willd. var. coriaceum Huber (Rusby 647): pollen
grains 3-colporate, oblate spheroidal (about 2224.5 yw). <A single
4-colporate (loxocolpate) pollen grain seen.

Apocolpium diameter about 15 ». Colpi narrow, about 12 pu long.
Ora lalongate (about 38 yu).

Exine about 1.5 uw thick or a little less. Sexine thicker than nexine,
probably tectate, presenting a reticuloid pattern (OL) with narrow
straight muroid and irregularly polygonal luminoid areas (maximum
diameter of the latter 1.5 «). Muroid areas supported by a single
row of endosexinous bacula.

The pollen grains of Herrania differ from those of Theobroma.

Herrania Goud.: pollen grains 3-colporate, peritreme, prolate
(about 32-35 23-25 pn).

Species investigated: HT. camargoana Schult. (Baker 39): about
34X25 uw; H. cuatrecasana Garcia B. (Cuatrecasas 11168): about
35X24 uw; H. mariae Schum. (Ducke 595 and Martius 318 (type)):
about 33 25 yw; H. puleherrima var. pacifica Schult. (Patiiio 23): about
32X23 yp.

Example: H. pulcherrima var. pacifica Schult.: pollen grains 3-
colporate, peritreme, prolate (about 3223 y).

Apocolpium diameter about 14 ». Colpi about 25 ». Ora about
2.25 pw high, slightly lalongate, their horizontal margins incrassate.

Exine about 2 » thick at poles, 1 » at center of mesocolpia. Sexine
thicker than nexine, tectate. Tegillum undulating, with anastomos-

CUATRECASAS—CACAO AND ITS ALLIES 445

ing, slightly winding, crestlike and slightly carinate folds imparting
a distinct reticuloid LO-pattern to the exine surface. Crests about
1 » broad at the poles, gradually more narrow (about 0.5 u or less) in
mesocolpia. They are supported by a single row of endosexinous
bacula except at the poles, where there are several rows. The lumi-
noid, concave areas between the folds of the tegillum are equally
supported by small endosexinous bacula or baculoid rods (largest and
longest at the poles). The longest diameter of these areas varies
between 2 and 5 u or more.

The pollen grains in Glossostemon bruguieri are somewhat similar
to those in Herrania.

Glossostemon bruguieri DC. (Iraq, Falluja, Haines s.n.): pollen
grains 3-colporate, peritreme, subprolate (85X28 y).

Apocolpium diameter about 8 u. Colpi about 25 yu, constricted at
the equator, ends rounded, margins thickened. Ora lalongate (about
8X1.5 pw).

Exine about 2.1 uw thick at the poles, about 1 » at the equator.
Sexine thicker than nexine, tectate. Tegillum strongly undulating,
forming distinct, anastomosing muroid ridges (about 0.5 wide)
separated by luminoid areas. In the apocolpia and towards the
colpi margins the latter are very small (diameter usually not exceeding
0.5 w); in the mesocolpia they are larger (longest diameter up to 4 y).
The muroid ridges are supported by a single or double row of endo-
sexinous bacula. The tegillar bottom of the luminoid areas is also
supported by small bacula. Bacula in apocolpia considerably longer
than those in mesocolpia.

The pollen grains in Abroma and Guazuma are somewhat similar
to those in Theobroma.

Examples: Abroma auqusta L. (Assam; herb. Riksmus., Stockholm,
marked “no. 370’): pollen grains cf. 3-colporate, peritreme, oblate
(21X29 wu).

Apocolpium diameter about 23 ». Colpi about 5.52.5 u, their
margins incrassate. Ora not very distinct.

Exine about 1 uv thick, tectate, very slightly undulating, presenting
a reticuloid pattern. Muroid ridges low, about 1 » wide, supported
by a double row of endosexinous bacula and enclosing small rounded
luminoid areas 1-2 » wide. Under each of the latter is one or several
endosexinous bacula.

Guazuma polybothra Cav. (Cuba, Boldo s.n.; herb. Madrid, marked
‘no. 94’): pollen grains 3-colporate, peritreme, prolate spheroidal
(18.5X16.5 y).

Apocolpium diameter about 12 yw. Colpi about 12 y long.

Exine about 1 » thick, tectate, undulating, with distinct narrow,

446 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

carinate muroid folds separated by luminoid, concave areas (diameter
less than 1 » in apocolpia as well as in mesocolpia).

Guazuma ulmifolia Lam. (Mexico, Pringle 2570): pollen grains
3-colporate, peritreme, spheroidal (16,).

Apocolpium diameter about 5 w. Colpi about 14 u long, 1 » wide.
Ora lalongate, about 1 » high and 3.5 pw wide.

Exine about 1 uw thick (of the same thickness in apocolpia as in meso-
colpia, probably tectate (tegillum undulating, exhibiting narrow,
muroid ridges separated by luminoid areas less than 1 » in diameter).

Cytology *
CONTRIBUTED BY F. W. Corr

CHROMOSOME NUMBERS IN THEOBROMA SPECIES.—The first pub-
lished count of 2n—20 for 7. cacao, the now accepted figure, was
made by Davie (1933) from studies of mitosis in root-tips. He
noted that “the chromosomes are very small, quite different from
Malvaceous chromosomes. A few show median constrictions. Two
pairs of satellited chromosomes were seen.” In 1935, Davie con-
firmed the diploid number of 20 from studies of meiosis in pollen
mother cells of 7. cacao.

Confirmation of this number has been made for T. cacao by Carletto
(1946), Mufioz Ortega (1948), Simmonds (1954) and Cope (unpub-
lished). The first three authors have also shown twenty to be the
diploid number in other Theobroma species. Carletto counted 20
chromosomes in 7. ‘“Teiocarpa,” T. speciosum, and T. granaiflorum
and Mufioz Ortega in 7. “leiocarpa,” T. ‘“pentagona,” T. bicolor,
T. mierocarpum, T. speciosum, T. simiarum, T. capilliferum, T.
grandiflorum, T. obovatum, T. angustifolium and T. cirmolinae. Sim-
monds confirmed 2n=20 in 7. bicolor and T. angustifolium. Accord-
ing to Mufioz Ortega, the chromosomes throughout the genus show
medial, submedial, and terminal centromeres. The chromosomes are
uniformly small, with size gradations within each species examined.
The largest chromosomes of 7’. cacao are 2 uv in length; the smallest of
T. microcarpum only 0.5 pu long.

¢ REFERENCES:

Carletto, G. M. (1946). O numero de cromosdmios em cacauciros. Bol. Tec. Inst. Cacau Bahia No.
6, 35-39.

Davie, J. H. (1933). Cytological studies in the Malvaceae and related families. Journ. Genet. 28: 33-67.

Davie, J. H. (1935). Chromosome studies in the Malvaceae and certain related families II, Genetica,
17: 487-498.

Mufioz Ortega, J. M. (1948). Estudios cromos6micos en el género Theobroma L, MSS in library of the
Instituto Interamericano de Ciencias Agricolas, Turrialba, Costa Rica.

Simmonds, N. W. (1954). Chromosome behavior in some tropical plants. Heredity, 8: 139-146.

CUATRECASAS—CACAO AND ITS ALLIES 447

Pollen Incompatibility °
CONTRIBUTED BY F. W. Corr

The incidence of self- and cross-incompatibility, and self- and
cross-compatibility in 7. cacao was first discovered by Pound (1932)
when he showed that some trees in Trinidad could not set fruit with
their own pollen nor with one another’s. These self- and cross-incom-
patible trees needed pollen from a self-compatible tree in order to
set fruit. Posnette (1945) discovered cross-compatibility between
self-incompatible types in his studies on cacao trees introduced from
the upper Amazon into Trinidad. The existence of self-incompatible
and self-compatible cacao trees has now been established in nearly all
areas where the species is wild or cultivated.

Cope has shown, in a series of publications, that unlike most other
plant species showing incompatibility the site of the incompatibility
reaction in cacao is in the embryo-sac, and not in the stigma or in
the style. Pollen tubes in incompatible pollinations grow as fast
as those in compatible pollinations and deliver their male gametes
into the embryo-sacs in perfectly normal fashion. It is only when the
male gametes come to lie in contact with their female counterparts that
any abnormality appears (fig. 11). According to the genotype of the
tree or trees involved in an incompatible pollination, either one quarter,
one half, or all encounters between male and female gametes result in
failure of the fusion process. When an incompatibly pollinated cacao
flower falls from the tree 25%, 50%, or 100% of the ovules in the
ovary show nonfusion; in the first two cases the other fertilized
ovules in the same ovary show normal fusion between the gametes to
give a zygote and a triploid primary endosperm nucleus in each.

The genetic system controlling the nonfusion and fusion of gametes
in the embryo-sac of J. cacao is now known. Three complementary
loci appear to be involved, which have been called A, B, and S. The
first two show simple dominance and recessivity; the S locus carries
multiple alleles between which dominance and independence relation-
ships exist. The action of the S locus was first postulated by Knight
and Rogers (1955), based on results obtained from wholly self-incom-
patible material. The need for other loci, to act in a complementary

6 REFERENCES:
Cope, F. W. (1939). Studies in the mechanism of self-incompatibility in cacao I. 8th Ann. Rep. on
Cacao Res. (1939), Trinidad, 20, 21.
(1940). Studies in the mechanism of self-incompatibility in cacao II. 9th Ann. Rep. on Cacao
Res, (1939), Trinidad, 19-23.

— (1958). Incompatibility in Theobroma cacao. Nature, London, 181, 279.
(1959). Incompatibility in Theobroma cacao, A Rep. on Cacao Res., 1957-58, 7-17.
Knight, R., and Rogers, H. H. (1955). Incompatibility in Theobroma cacao. Heredity, 9: 69-77.
Posnette, A. F. (1945). Incompatibility in Amazon cacao. Trop. Agriculture, Trin., 22: 184-187.
Pound, F. J. (1932). Studies in fruitfulnessin cacao. II—Evidence for partial sterility. 1st Ann, Rep.

on Cacao Res. (1931), Trinidad, 24, 25.

448

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Figure 11.—a, Camera lucida drawing of an embryo sac of Theobroma cacao, fixed 24 hours

after pollination: one male gamete (0) is in contact with the egg nucleus (9) and
the second is moving towards the polar nuclei, p; the darkly-staining synergid cell (sy)
has been penetrated by the pollen tube; and starch grains (s) are abundant. sB, camera
lucida drawing of an incompatibility fertilized embryo sac of 7. cacao, 72 hours after
pollination: one male nucleus lies in contact with the egg nucleus (top L.H.); and the
second male nucleus (¢’3) is associated, unfused, with one polar nucleus (pg), the other
polar (p;) having moved away. c, as in B, except that the second male nucleus (<2)
and the two polar nuclei (p; and pg) are all dissociated (co; is the first male gamete
lying in contact with the egg nucleus). , as in B, except that the second male nucleus
is separated from the two coherent polar nuclei. F.W. Cope, original.

CUATRECASAS—CACAO AND ITS ALLIES 449

manner with the S locus, was pointed out by Cope (1958) in order to
explain the emergence of self-incompatible progeny from a cross be-
tween two true-breeding self-compatible parents.

The A and B loci both act before meiosis. When both are at least
heterozygous for the dominant allele, it is believed that a general pre-
cursor substance is produced and on this the S locus acts to produce
very highly specific incompatibility reactions between gametes carry-
ing the same S allele. The S locus acts both before and after meiosis,
the premeiotic action giving the overall sporophytic control of incom-
patibility and the postmeiotic action leading to a gametophytic reac-
tion between gametes.

If one or more of the A, B, and S loci become homozygous for an
inactive allele the self-incompatible condition is lost; the tree is then
self-compatible and cross-compatible with any other cacao genotype.

A few examples of genotypes of the two classes of tree are:

Self-incompatible Self-compatible
AABBS, aaBBS, y aaBBS..t
AaBbS,.y aabbS,.s
AABbS,.: AABBS,.¢
AaBBS, +t aaBbS:.+

where S, and S, are two active 5 alleles and 5, is an inactive amorph
of the 5 series.

Self-incompatible genotypes are also cross-incompatible if they
have common dominant S alleles; if one has alleles independent in
action and one of these is duplicated in the other as a dominant; or
if both genotypes, carrying only alleles of independent action, have one
allele in common.

Genus Theobroma ° L.

Theobroma L. [Gen. Pl. 351. 1737; Hort. Cliff. 379. 1737] Sp. Pl. 782. 1753;
Gen. Pl. ed. 5, 340. 1754; Benth. & Hook. (1862) 225; Bernoulli (1869) 4;
Schumann (1890) 86 pro parte; Schumann (1896) pro parte; Chevalier (1946)

269.
Cacao [Tourn. Inst. 660, t. 444. 1700). Miller, Gard. Dict. Abr. ed. 4th. 1754.

Tribroma Cook, Journ. Washington Acad. Sci. 5:288. 1915.

Typr.—Theobroma cacao L.

Flowers hermaphroditic, pentamerous, pentacyclic, diplostemonous.
Buds globose, ovoid or oblong-ovoid. Sepals 5, valvate in aestivation,
almost free and spreading or more or less united in the lower part,
cupular, or united by pairs into one single and two double lobes, or
rarely in two lobes. Petals 5, dextrorsely contorted in aestivation,
each one strangulated in two halves: 1) a lower part corresponding

6 Theobroma is a neuter name and the genus must be neuter by the present International Code of Nomen-

clature. The feminine endings for the species used by some authors (De Candolle, Bernoulli, Chevalier,
etc.) are corrected in this revision into the neuter form except for the original bibliographic references.

450 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

to the claw, rigid and strongly veined with the shape of a hood (cu-
cullus); 2) an upper part, a flat blade (lamina), articulated to the
inflexed apex of the claw. Androecium in two verticils of five,
united in a tube at base: an outer whorl with 5 sterile, petaloid or
linear staminodes, opposite to the sepals; an inner whor! with 5 fertile
stamens opposite to the petals, the filaments short, minutely 2-3-
branched, each branch with an anther. Anthers hidden inside the
petal-hoods, bilobate (bithecate), the thecae unilocular and dehiscent
by longitudinal clefts. Pollen grains 3-colporate, peritreme, sub-
oblate (about 15-22 x 17.5-25 pw). Gynoecium 5-carpellar, syncarpic,
superior, the carpels opposite to petals, the ovary ovoid, pentagonal,
5-celled with axile placentation, the many ovules in two rows in each
cell. Stylodes 5, connivent, free or more or less united, filiform.
Stigmas apical, short, acute. Ovules anatropous with two integu-
ments and dorsal raphe.

Fruit large, subbaccate or subdrupaceous, indehiscent, ovoid,
ellipsoid or oblong, obtuse or acute, smooth or ridged, rugose or
tuberculate, the pericarp fleshy or hard and partly woody or coriace-
ous, the vascular axis thin and vanishing; seeds usually in five rows,
each one surrounded by a thick, fibrose, pulpy tissue filling the cavity
at maturity, ovoid, ellipsoid, or amygdaloid, the episperm double,
thick, subcoriaceous, the outer layer with a trichomatic and gelati-
nous epiderm developing into a thick, pulpy envelope; embryo straight,
the radicle cylindrical, inferior; cotyledons thick, strongly plicate-
corrugate; endosperm usually reduced to a filmy membrane covering
the cotyledons. Germination epigeous or hypogeous.

Evergreen tree with the apical growth of the stem limited to the
production of a terminal whorl of 3-5 spreading branches; sympodial
growth of the stem attained by adventitious upright subterminal
shoots or by pseudoapical shoots from buds axillary to the apical
branching whorl. Primary branching of stem 3- or 5-verticillate, the
further branching alternate. Leaves simple, entire, penninerved,
persistent, coriaceous, long-petiolate and varied in phyllotaxy on the
primary stems, short-petiolate and distichous on the branches.

Inflorescences dichasial or monochasial (cincinate), axillary or on
reduced tuberculiform branchlets on trunk and larger branches.
Peduncles bracteate, articulate to pedicels.

Pluricellular trichomes in all species, usually as stellate hairs,
rarely simple. Globose, stipitate glands present in some species.

Chromosome number: 2n=20.

Subgeneric classification

The division of the genus Theobroma in five sections by Bernoulli
is the best to date. He used the characters of the petal-lamina
(sessile, stipitate, or lacking), shape of staminodes and their position

CUATRECASAS—CACAO AND ITS ALLIES 451

in the bud, and the number of anthers. A sound combination of these
characters gives five very natural groups. Schumann’s (1886, 1890)
separation of two sections, Theobroma and Bubroma, according to their
2-antheriferous or 3-antheriferous stamens, leads to an unnatural
grouping because the number of anthers for each stamen may vary
in the same section and even in the same species (e.g., 7. glaucum).
For this reason, the combination of Schumann’s with Bernoulli’s
classification made by Pittier (1930) was erroneous, because the section
Oreanthes cannot be placed in either of Schumann’s two groups.
Chevalier (1946) used both classifications but without trying to in-
tegrate them. Ducke (1954), who published the best elaborated key
for 7 Brazilian species, did not pay attention to sections, but he used
the 3- or 5-whorled branching as a new character to distinguish the
species. Another character, the epigeous or hypogeous germination
of the seeds correlated with growth-habit of the tree, was used by
Addison and Tavares (1951) for distinguishing species, and by Cope
and Bartley (1960) in classifying them.

I have applied to the classification the mode of germination, and
the growth and the branching system for all the species, and I have
found the sections founded by Bernoulli to be very much reinforced
by the addition of these vegetative features, and other floral and fruit
characters unknown before. Epigeous germination and subterminal
growth apply to all species of the sections Rhytidocarpus, Oreanthes,
and Theobroma, whereas the other sections exhibit hypogeous germi-
nation and pseudoterminal growing. These vegetative characters
prove to be very important and basic, being uniform for each section,
but like other characters, even though constant within the section,
they are not sufficient to give taxonomic recognition to the two groups
separated by those characters. The Bernoulli sections are all of
similar rank, independent and probably of parallel origin. Only the
new section Andropetalum, based in its extraordinarily broad and
reflexed staminodes, relatively reduced petal-lamina, and gamosepal-
ous calyx, seems to be closer to Glossopetalum than the other sections
to each other. The fruit structure, as explained above, is also impor-
tant in the present classification. The following key will give a clear
idea of the characters of each section. Another artificial key is
added to facilitate the identification of specimens lacking complete
information.

The position of the inflorescences (cauline or axillary), the color,
size, and shape of petal-lamina and staminodes, number of anthers,
shape of sepals and indument, form and size of fruits, the outline,
venation, thickness, and firmness of leaves, and especially the kind of
indumentum they bear, as well as the pilosity on different parts of the
flowers, inflorescences, and branchlets, and the form and caducousness
of stipules are the characters used to distinguish the species.

452 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Key to Sections of Theobroma

1. Cotyledons epigeous at germination; growth of stem by adventitious upright,
lateral-subterminal shoots; staminodes in aestivation erect.

2. Staminodes thick-linear, obtuse; petal-hood 1l-nerved; petal-lamina sub-
sessile; stamens 2-antheriferous; pericarp thick, ridged and nerved, the
mesocarp very hard, woody; Primary branches ternate; leaves tomentose
beneath. ........., Loe ees 1. Rhytidocarpus

(Contains the single species: 7. bicolor)

2. Staminodes linear-subulate or lanceolate, acute; petal-hood 3-nerved.

3. Petal-lamina sessile; stamens 3- or 2-antheriferous; pericarp coriaceous ;
primary branches ternate; leaves tomentose beneath. . 2. Oreanthes

3. Petal-lamina attenuate-stipitate; stamens 2-antheriferous; pericarp firmly
carnose; primary branches quinate; leaves glabrous or puberulous
beneath. . 2... ..... ..... 38. Theobroma

(Contains the single species: T. cacao)
1. Cotyledons hypogeous at germination; growth of stem pseudoapical; primary
branches ternate; stamens 3-antheriferous.

4. Staminodes flexuose in bud, ovate, subulate-caudate; petal-hood 5-nerved;
petal-lamina lacking; pericarp carnose-coriaceous, lignose-ridged and
reticulate; leaves glabrous or puberulous . .. . . 4. Telmatocarpus

4. Staminodes reflexed in bud, obovate-oblong, broadly lanceolate or broadly
obovate, reflexed or erect in anthesis; petal-hood 7-nerved; pericarp
rigid, the epicarp hard, woody; leaves tomentose beneath.

5. Staminodes oblong-obovate or lanceolate, reflexed or erect at anthesis;
petal-lamina broadly developed, Hat stipitate; sepals more or less
united and reflexed. . . . . ... . . 5. Glossopetalum

5. Staminodes broadly obovate, as broad as long; petal-lamina somewhat
reduced, narrow, and plicate; calyx cupular, the sepals united one
half or one third their length. ..... . . . 6. Andropetalum

(Contains the single species: T. mammosum)

KEY TO SECTION OREANTHES

1. Leaves stellate-tomentose beneath on the minor reticulate veins, the areoles
densely, minutely, stellulate-tomentose.
2. Filaments 2-antheriferous; inflorescences small, on the leafy branches; flow-
ers rather small (sepals 7-9 x 2-2.5 mm.); fruits globose-elliptical about
10 x 9cm., glaucous when ripe. Leaves beneath with the quaternary
nerves, minor veins, and areoles covered Py minute tomentum of minute,
thin, white, stellate hairs. . .. ... . . 2. T. sylvestre
2. Filaments 3-antheriferous; inflorescences 0 on the trunk, multiflorous; flowers
rather large (sepals 10-12 x 3.5-4 mm.).

3. Leaves with glabrous primary and secondary nerves beneath or sub-
glabrous with very scattered mediocre stellate hairs and sparse callose
spots; fruit globose-ellipsoid, 10 x § cm., without ribs, shortly tomentose,
yellowish when ripe. . . . ... . .& T. speciosum

3. Leaves softly velutinous beneath, the nerves and veins with abundant
long, thin, patulous stellate hairs; fruits ellipsoid, densely velutinous,
with 5 very prominent ribs... .. ... . . 4 Ty velutinum

1. Leaves with glabrous nerves and veins beneath, or subglabrous with very
sparse, mediocre stellate hairs, only the areoles covered with compact
tomentum of minute, white, stellate hairs.

CUATRECASAS—CACAO AND ITS ALLIES 453

4. Flowers large: petal-lamina suborbicular 5.5-7 x 5-6.5 mm.; petal-hood
5-6 x 2.5-3 mm.; staminodes lanceolate-subulate 10-12 mm. long; fila-
ments 2—3-antheriferous; sepals 12-13 x 3-4 mm.; petal-hood puberulous;
fruit ellipsoid, obtusely pentagonal, attenuate at apex, umbilicate at base,
10-11 x 5-5.5 em. Leaves coriaceous, broadly ovate or ovate-oblong.

5. T. glaucum

4. Flowers smaller: petal-lamina suborbicular, orbicular or elliptic, 2.5-4 mm.
long; staminodes 6-9 mm. long; filaments 2-antheriferous; sepals 8-10 x
3 mm.; petal-hood hirtellous pubescent; fruit ellipsoid-oblong, obtusely
pentagonal, abruptly narrowed at apex, umbilicate at base, constricted
or not above the base, 12-25 x 5.2-Sem. .... . .6. T. bernouillii

KEY TO SECTION TELMATOCARPUS

1. Leaves regularly penninerved; inflorescences on trunk and branches; pedun-
cles 5-25 mm. long; pedicels 7-8 mm. long; fruit peduncles 2-3 cm. long;
sepals stellate-tomentose; petals pilose above, narrowed in the lower third,
the apex blunt or emarginate; staminode-base with very short, thick
hairs; ovary ovoid-ellipsoid, tomentose; fruit, when ripe, ovoid, attenuate
at apex, slightly 5-costate, alveolate, the epicarp densely appressed tomen-
tose, 7.5-11 x 7-9em. 2... ee ee ee 8. T. gileri

1. Leaves at base 3-nerved, the two lateral basal nerves ascending at an acute
angle, the other 2 or 3 pairs of secondary nerves remotely higher; inflores-
cences axillary on young branchlets; peduncles 0.5-1 mm. long; pedicels
0.5-1 mm. long; fruiting peduncles 4-8 mm. long and thick; sepals with
sparse stellate hairs; petals glabrous, gradually attenuate to the base, the
apex acuminate, the acumen acute, 2-dentate; staminode-base with rather
thick, long, flexuose hairs; ovary pyriform, glabrous or sparsely granulate,
rarely sparsely stellate-pilose; fruit, when ripe, ellipsoid-globose, conspicu-
ously 10-costate, reticulate-alveolate, usually 6.5-7 x 6-6.5 cm.

9. T. microcarpum

KEY TO SECTION GLOSSOPETALUM

1. Inflorescences born on the trunk and main branches. Flowers large; stami-
nodes obovate-oblong or oblong-spathulate, 9-11 mm. long, erect in anthesis;
leaves large (20-54 x 8-30 cm.), coriaceous, obtuse at both ends, strongly
nerved and tomentose beneath; stipules coriaceous, persistent; calyx 5-
lobate, cupular at base reflexed.

2. Petal-lamina and staminodes yellow.

3. Fruit ellipsoid-oblong or ovoid-oblong, attenuate at both ends, umbilicate,
obtusely pentagonal, 25-35 x 10-12 cm.; leaves oblong-elliptic or ovate-
elliptic, subvelvety beneath with minute, stellate, flexuous, white hairs
crowded on the areoles and minor veins, and other equal or slightly
larger hairs on the other nerves; stipules oblong-lanceolate, subacute;
petal-lamina subdeltoid-spatulate; staminodes oblong-obovate.

11. T. cirmolinae

3. Fruit ovoid-ellipsoid or ellipsoid, smooth, rounded at both ends or slightly
attenuate at apex, 18-22 x 9-11 cm.; leaves oblong-elliptic or ovate-
elliptic, tomentose beneath with minute, stellate, flexuous, white hairs
crowded on the areoles and minor veins, and other larger, thicker, fer-
ruginous hairs copious on the other nerves; stipules ovate or ovate-
oblong, obtuse; petal-lamina subtriangular-spatulate; staminodes
oblong-obovate .. 2... 2.2 eee eee 12. T. stipulatum

454

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

2. Petal-lamina and staminodes red. Fruits smooth (not ridged).

4. Fruit ellipsoid-oblong, rounded at apex, umbilicate, 16-40 x 6-11 cm.;

leaves obovate-oblong, densely covered beneath with minute, stellate,
white, intricate hairs on the areoles and minor veins, other mediocre,
stellate, ferruginous hairs sparse or copious on nerves, and larger ones
with longer rays usually copious on the major nerves; stipules lanceo-
late; petal-lamina subtrapezoid, attenuate at base; staminodes
oblong-obovate . . . . . 14, T. simiarum

4, Bruit ellipsoid-ovoid, rounded at base, obtuse at apex, 19-20 x 10-11.5

.; leaves oblong-elliptic or ovate, often rugose, densely covered
beneath with minute, stellate, white, intricate hairs on areoles and
minor veins, and other larger ferruginous hairs with longer, spreading
rays abundant on nerves; stipules ovate, rather obtuse; petal-lamina
obovate-deltoid, attenuate at base; staminodes obovate-oblong.

13. T. chocoense

1. Inflorescences small, axillary on foliose branches.
5. Petal-lamina and staminodes yellow; staminodes oblong-obovate, erect.

5.

Calyx trilobate, reflexed; leaves thin, subcoriaceous, subobovate-oblong
or elliptic-oblong (9-25 x 3-9 cm.), attenuate at both ends, acute, cinere-
ous beneath with minute, stellate, whitish, intricate hairs and larger ones
with longer, patulous rays on nerves; stipules membranaceous, subulate,
deciduous; petal-lamina subobovate-spatulate, bilobate, emarginate;
fruit ellipsoid-elongate, 5-angulate, more or less irregularly tuberculate,
10-18 x 6-9em. ..........-.-2--- 10. T. angustifolium

Petal-lamina and staminodes red; staminodes curved, spreading in anthesis.
6. Young branchlets, petioles, and buds covered with a woolly-floccose,

ochraceous or tawny, deciduous tomentum. Adult leaves glaucous

or glauco-cinereous, monotrichous beneath, with the principal and ter-

tiary nerves glabrous, glossy, reddish punctate, the areoles and reticulum
whitish tomentose by minute stellate hairs.

Flowers large; calyx (14-15 mm. long) 3-lobate; staminodes very acute,
lanceolate, spreading (9-13 mm. long); petal-lamina trapezoid-
elliptic, thick, dark red; leaves firm-coriaceous ; stipules subcoriaceous,
persistent; fruits ellipsoid, 16-25 x 10-12 cm., rounded at both ends,
smooth. .... . ... . 15. T. grandiflorum

Flowers smaller; - calyx (about 7 mm. ong) 5-parted; staminodes oblong-
elliptic, rounded at apex (6 x 2.8-3 mm.); petal-lamina suborbicular,
red, rather thick; leaves thin-chartaccous, very asymmetrical at
base; stipules membranaceous, linear-subulate, deciduous; fruit
ellipsoid-obovoid, acute, granulate-tuberculate, 5-7 x 3-4 cm.

16. T. obovatum
Young branchlets, buds and petioles hirsute, or short-tomentose. Calyx
5-parted.

8. Young branchlets, buds, and petioles densely hirsute or hirsute-

tomentose; leaves beneath tomentose-hirsute (especially in young

plants), ferruginous or ochraceous by spreading long-radiate hairs

on the nerves and veins, and with minute, white, intricate ones
covering the surface.

9. Pedicels 5-10 mm. long; peduncles 5-10 mm. long; inflorescences

loose; sepals about 10 mm. long; petal-lamina and staminodes not

ciliate; fruit ellipsoid-pyriform, smooth . . . . 20. T. sinuosum

CUATRECASAS—CACAO AND ITS ALLIES 455

9. Pedicels up to 1 mm. long; peduncles up to 6 mm. long; inflorescences
compact, glomerate; sepals about 7 mm. long; petal-lamina and
staminodes ciliate; fruit unknown. . . . 21. T. canumanense

8. Young branchlets, buds, and petioles densely subappressed-tomentose;
leaves cinereous or ferruginous beneath with minute, stellate, whitish,
intricate hairs covering the areoles and the smallest reticulum and
mediocre, thicker, ferruginous hairs, copious or scattered on the
nerves.

10. Bracteoles narrowly linear.

11. Staminodes scarlet, lanceolate, acute or subacute, 6-7.5 x 2 mm.;
petal-lamina scarlet, orbicular or subrounded, thick, 2-2.5 x
2.2-3 mm.; ovary glabrous and smooth or very sparsely granu-
lar; fruit ellipsoid or oblong-ellipsoid, often slightly attenuate
at base, 7.5-11.5 x 5-6.6 cm., the pericarp tomentose, smooth,
3-4mm.thick whendry ..... .. 17. T. subincanum
11. Staminodes brownish red, obovate-oblong, rounded or subspatu-
late at apex, 5-5.5 x 2-3 mm.; petal-lamina obovate-subrhom-
bic, 4-5 x 4 mm.; ovary densely tomentose; fruit ellipsoid,
rounded bothends,7x4em. ...... . 18.T.hylaeum
10. Bracteoles 3, orbicular, cochlear, embracing the single bud; stami-
nodes brownish red, obovate-oblong, rounded at apex, 6-7.5 x
3.5-4.5 mm.; petal-lamina oblong-elliptic or oblong-obovate,
5-7 x 3-5 mm.; ovary densely tomentose; fruit ellipsoid, rounded
at apex, constricted above the base, 8-10 x 4.5-6 cm., the peri-
carp, when dry, 1-1.8 mm. thick, the epicarp 1 mm. thick,
fragile at maturity ........... 19. T. nemorale

Artificial key to the species

1. Leaves glabrous or puberulous beneath.

2. Leaves firmly coriaceous, regularly penninerved, with 10-14 secondary
nerves each side, usually 20-30 x 7-10 cm.; fruits glabrous, ovate-oblong,
more or less pentagonal or decagonal, with carnose relatively thick peri-
carp; stamens 2-antheriferous; staminodes linear-subulate, erect in bud,
red; petal-lamina spatulate, stipitate, yellowish; jorquette with 5
branches; growth below jorquette. Flowers on trunk and on branchlets.

7. T. cacao

2. Leaves chartaceous with 3-6 pairs of secondary nerves; fruits smaller,
ovoid or subglobose, with hard-costate reticulate tomentose pericarp;
stamens 3-antheriferous; staminodes subulate-subflagelliform, flexuous
in bud; petals lacking laminae or ligular appendages; jorquette with 3
branches; growth above jorquette.

3. Leaves 5-20 x 1.5-8 cm., regularly penninerved with 5 or 6 nerves on

each side, puberulous beneath, the midrib tomentulose; peduncle

5-25 mm. long; pedicels 7-8 mm. long; ovary tomentose; fruit ovoid,

slightly 5-costate and reticulate, 7.5-11 x 7-9 cm., fruiting peduncle

2-3 cm.; flowers on trunk and on branchlets. Sepals tomentulose.

8. T. gileri

3. Leaves 6-16 x 2-7 cm., 3-nerved at base, the two lateral-basal nerves

ascending, quite distant from the other 2 or 3 pairs of secondary

nerves, glabrous or minutely and sparsely puberulous beneath; pe-

duncle 0.5-1 mm. long; pedicel 0.5-1 mm. long; ovary glabrous or

sparsely granulate; fruit ellipsoid or subglobose ellipsoid, strongly
680—695—64——6

456 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

10-costate and reticulate 6.5-7 (-9) x 6-6.5 em.; fruiting peduncle
4-8 mm. long; flowers only on branchlets. . . 9. T. microcarpum
1. Leaves densely stellate-tomentose beneath. Jorquette 3-branched.
4. Leaves thin, chartaceous, tomentose-cinereous beneath. Flowers on ter-
minal, leafy branchlets.

5. Leaves ovate or ovate-oblong, subcordate or cordate at base, cinereous
or ochraceo-cinereous (more or less silvery) beneath, the hairs uniform,
minute, stellate, covering the surface and nerves; stamens 2-antherif-
erous; petal-lamina subsessile; growth below jorquette. Dichotomous
inflorescences on branches; fruit large, ellipsoid with thick-woody,
strongly costate-reticulate and lacunose pericarp; flowers small, red
dish; staminodes thick-linear, obtuse . .... . . 1.T. bicolor

5. Leaves oblong, elliptic-oblong, lanceolate-oblong, regularly pinnate-
nerved, obtuse or cuneate at base, tomentose-cinereous beneath;
stamens 3-antheriferous; petal-lamina stipitate; growth above
jorquette.

6. Leaves obovate-elliptic or obovate-oblong, obtuse and very asym-
metrical at base with 5-7 nerves each side, 7-35 x 3-13 em.; homo-
trichous, covered by minute, dense, white, stellate hairs beneath,
except for the glabrous red-punctate main veins; young vegetative
parts with a floccose, lanate, ochraceous, deciduous indument; fruit
small, ellipsoid-obovoid, tuberculate-warty, 5-7 x 3-4 cm.; calyx
5-lobate. Flowers small; staminodes petaloid, red.

16. T. obovatum

6. Leaves elliptic-lanceolate or oblanceolate, rather oblong, slightly
asymmetrical at base, heterotrichous beneath with dense, white,
minute, stellate hairs and longer, patulous or subpatulous, pale
ochraceous ones on the main nerves; young vegetative parts mi-
nutely tomentose; fruit large; calyx 2- or 3-lobate.

7. Leaves subobovate-oblong or elliptic-oblong or oblanceolate, acute,
9-25 x 3-9 cm., with 6-8 secondary nerves on each side; petal-
lamina and erect staminodes yellow; fruit ellipsoid-oblong, ir-
regularly sulcate tuberculate, 10-18 x 6-9 em.

10. T. angustifolium

7. Leaves elliptic-oblong or sublanceolate, 10-25 x 3.5-8.5 em., with
9-12 secondary nerves each side; petal-lamina small, narrow,
red; staminodes red-purplish, very broad, reflexed, covering the
stamens and petals; fruit ellipsoid-oblong, constricted above the
base and at the top below the apex, 10-20 x 6-8 em.

22. T. mammosum
4. Leaves coriaceous, firmer and more markedly nervose-reticulate than in
the species above.

8. Leaves ovate-oblong or elliptic-oblong, long-caudate, with curved, as-
cending secondary nerves, softly velvety or apparently glabrous and
shining beneath; growth below the jorquette; petal-lamina sessile;
staminodes lanceolate or subulate, erect in bud.

9. Leaves more or less bullate, softly velvety beneath, heterotrichous
with a layer of dense, minute, white, stellate hairs and longer,
thin-rayed, patulous, stellate hairs on the nerves. Flowers purplish
red on trunk; fruit ellipsoid with 5 protuberant ribs, 8-9 x 6-6.3 em.

4. T. velutinum

CUATRECASAS—CACAO AND ITS ALLIES 457

9. Leaves flat, with practically homotrichous, cinereous or whitish
indument beneath, of minute, white, stellate hairs; larger hairs
very rare, the major nerves glabrous.

10. Leaves beneath with glabrous major nerves, the quaternary and
minor veins stellate-tomentulose, the areoles densely whitish
tomentose.

11. Inflorescences small, on leafy branchlets; flowers small, brownish
red; stamens 2-antheriferous; fruit ellipsoid-globose, glaucous,
10x 9m... .... . 2 T. sylvestre

11. Inflorescences large, on the trunk; flowers purplish red, larger than

above; stamens 3-antheriferous fruit globose-ellipsoid, 10 x 8
em., yellowish . ... . .... . .3 T. speciosum

10. Leaves beneath with completely ‘glabrous veins, only the smallest
veins of the reticulum subglabrous with scattered mediocre
hairs, the areoles with very appressed tomentum of minute,
white, stellate hairs, the leaf surface with a glabrous appearance.

12. Flowers 12-13 mm. long; petal-lamina suborbicular 5.5-7 x
5-6.5 mm. long; staminodes lanceolate-subulate, 10-12 mm.
long; stamens usually 3-antheriferous, also 2-antheriferous;
fruit ellipsoid, obtusely pentagonal, 10-11 x 5-5.5 cm.

5. T. glaucum

12. Flowers 8-10 mm. long; petal-lamina elliptic, suborbicular or
orbicular, 2.5-4 mm. long; staminodes 6-9.5 mm. long; sta-
mens 2-antheriferous; fruit ellipsoid-oblong, more or less
pentagonal; 12-25 x 5.2-8 cm... . . . 6. T. bernouillii

8. Leaves broad or oblong with regularly spreading, pinnate, secondary
nerves, cinereous or ferruginous tomentose beneath, with prominent
venation; growth above the jorquette; petal-lamina pedicellate or
very reduced; staminodes broadly oblong. Stamens 3-antheriferous.

13. Inflorescences on leafy branches.

14. Leaves beneath monotrichous, glaucous, covered by minute, white,
intricate, stellate hairs, the main nerves glabrous with scattered
reddish, callous spots; young vegetative parts with ferruginous,
floccose-lanate, deciduous indument; calyx trilobate; fruit large,
ellipsoid, smooth, 16-25 x 10-12cm. ; stipules persistent. Flowers
dark red. .... .... 15. T. grandiflorum

14. Leaves beneath heterotrichous, with minute, white, densely intri-
cate, stellate hairs covering the surface, and larger, thicker,
reddish or ochraceous stellate hairs on the veins; calyx 5-lobate;
fruits ellipsoid, smooth, 7-11.5 x 4-6.6 cm.; stipules caducous.

15. Young branchlets hirsute or hirsute-tomentose; leaves with long,
spreading radiate hairs beneath.

16. Pedicels 5-10 mm. long; peduncles 5-10 mm. long; sepals
about 10 mm. long; staminodes and petals not ciliate.

20. T. sinuosum

16. Pedicels almost lacking (up to 1 mm. long), peduncles up to

6 mm. long; sepals about 7 mm. long; staminodes and petals

ciliate .... ..... 21. T. canumanense

15. Young branchlets and leaves beneath subappressed-tomentose.

17. Bracteoles broadly ovate or orbicular. Staminodes obovate-
oblong, rounded at apex; petal-lamina oblong, obovate spat-
ulate; ovary densely tomentose... .. 19. T. nemorale

17. Bracteoles linear.

458 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

18. Staminodes scarlet, lanceolate, acute, or subacute; petal-
lamina scarlet, orbicular or suborbicular; ovary glabrous
or very sparsely granular. . . . . 17. T. subincanum

18. Staminodes brownish red, obovate-oblong, rounded or sub-
spatulate at apex; petal-lamina obovate, subrhombic;
ovary densely tomentose. ... . . 18. T. hylaeum

13. Inflorescences on trunk and branches. Leaves usually large and thick-
coriaceous, strongly nervose beneath; stipules persistent.
19. Flowers yellow.

20. Indument of leaves beneath subvelvety, the minute stellate hairs
on veins slightly larger than those of the surface; stipules oblong-
lanceolate, subacute; fruits ellipsoid-oblong, obtusely pentag-
onal narrowed at apex, 25-35 x 10-12 cm. . 11. T. cirmolinae

20. Indument of the leaves beneath, tomentose, the hairs on the
veins ferruginous, larger than those of the surface; stipules
ovate, obtuse; fruits ovoid-ellipsoid, rounded at both ends,
smooth, 18-22x9-llem. ...... . .12.T. stipulatum

19. Flowers purple-red.

21. Leaves obovate-oblong with three kinds of hairs beneath (minute,
mediocre, and longer); stipules lanceolate; fruit ellipsoid-
oblong, rounded at apex, 16-40 x 6-llcm. . . 14. T. simiarum

21. Leaves oblong-elliptic or ovate, usually rugose, with two kinds of
hairs beneath (minute and larger); stipules ovate, rather
obtuse; fruit ellipsoid-ovoid, rounded at base, obtuse at apex,
19-20 x 10-115 cm. ........ . .13. T. chocoense

Section I. Rhytidocarpus

Theobroma sect. Rhytidocarpus Bernoulli, Uebers. Art. Theobroma 9. 1869.
Sect. Eutheobroma subsect. Rhytidocarpus (Bernoulli) Pittier, Rev. Bot. Appl.
10(110):779. 1930.

Petal-lamina very shortly stipitate, subsessile. Petal-hood 1-nerved.
Staminodes linear-oblong, obtuse, thick, erect in aestivation. Fila-
ments 2-antheriferous. Fruits subglobose-ellipsoid with hard _peri-
carp, strongly costate and reticulate-nerved, minutely tomentose,
the mesocarp thick-woody, very hard. Cotyledons epigeous at ger-
mination. Leaves beneath, appressed stellate-tomentose. Primary
leaves palmatinerved, regular leaves subpalmatinerved, the base 5-7
Merved. Inflorescences axillary or extra-axillary on leafy branches.
Sympodial growth of stem by orthotropic, adventitious, lateral-
Subterminal shoots. Primary branches ternate, deciduous in age,
leaving a naked stem; leafy crown lax, flat. Secondary branching
dichotomous.

TyPE spEciEs.—Theobroma bicolor Humb. & Bonpl.

A single species known.

1. Theobroma bicolor Humb. & Bonpl. Fiaures 2, 5, 9, 12, 18, 35; Map 3
Theobroma bicolor Humb. et Bonpl. Pl. Aequin. 1:104, pl. 30. 1806; H. B. K.

(1823) 317; Triana & Planch. (1862) 208; Bernoulli (1869) 9, pl. 4; Schu-
mann in Mart. (1886) 73; Jumelle (1899) 21, figs. 10, 11; Preuss (1901)

CUATRECASAS—CACAO AND ITS ALLIES 459

Ficure 12.—Leaves of Theobroma, X ¥%: a, bicolor, from orthotropic branches (Dawe 83),
from above; B, base of same from underside; c, bicolor from plagiotropic (current)
branches (Kill. & Smith 30006); p, speciosum var. coriaceum (Huber 1567); £, velutinum
(Benoist 516).

460 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

251, 255, pls. 3, 4; De Wildeman (1902) 94; Huber (1906a) 274; Standley
(1923) 808; Ducke (1925) 132; (1940) 269, pl. 1, fig. 2; (1954) 13; Standley
(1937) 687; Chevalier (1946) 276; Standl. & Stey. (1949) 422; Holdridge
(1950a) 3; Addison & Travares (1951) pls. 10, 13, fig. 8; Baker, Cope & al.
(1954) fig. 9; Cuatrecasas (1956) 652; Leén (1960) 313, 315 fig.

Theobroma ovatifolia Moc. & Sessé ex DC. Prodr. 1:485. 1824; Icon. FI.
Mex. DC. pil. 113.

Cacao bicolor (Humb. & Bonpl.) Poir. in Lam. Encycl. Méth. Suppl. 2:7.
1811.

Tribroma bicolor (Humb. & Bonpl.) Cook, Journ. Washington Acad. Sci.
5:288. 1915; Contr. U.S. Nat. Herb. 17(8), pls. 46, 47, 48, 49, 60, 62,
54, 1916.

Theobroma cordata Ruiz & Pavén, Fl. Peruv. Chil. vol. 6, ined.

Mar 3.—Geographical distribution of Theobroma bicolor, based on specimens mostly from
planted trees.

Typus.—Humboldt & Bonpland, Colombia (P). Mocifio & Sessé,
Mexico (of 7. ovatifolia).

Commonly a small tree 3-8 m. tall, attaining in high forest a height
of 25-30 m., with rather narrow crown; sympodial growth by lateral,
subterminal, upright shoots; trunk erect with light bark and white
wood; primary branches ternate, dichotomous, spreading; young
branchlets often horizontal or pendulous, more or less flexuous,
subterete, densely and appressed cinereous tomentose with minute

CUATRECASAS—CACAO AND ITS ALLIES 461

stellate hairs; older branches glabrate, smooth, gray; stipules oblong-
lanceolate, 5-8 mm. long, 1.2-2 mm. broad, minutely appressed-
tomentose, more or less persistent.

Leaves subpalmatinerved, firmly chartaceous, green above and
silvery greenish or silvery cinereous, sometimes pale ochraceous
beneath; petiole rather thick, subterete, rigid, minutely appressed-
tomentose, 12-25 mm. long, transversely rimose when old; blade
oblong-ovate or elliptic-ovate, more or less deeply cordate or emargi-
nate, asymmetrical at base, attenuate, abruptly acuminate at apex,
entire or rarely sinuate at the upper margin, 12-34 cm. long, 6-18
cm. broad, the acumen triangular, 6-12 mm. long, glabrous above or
with scattered stellate or furcate hairs, green or when dry, pale
brownish, the main nerves noticeable, the lesser slightly conspicuous,
cinereous beneath, covered with a dense layer of intricate, white,
sericeous, stellate hairs, at base 5-7-nerved, the thicker costa and 2 or
3 main nerves on eachsidestrongly prominent, the interior, basal nerves
upright, ascending, the 1 or 2 exterior basal pairs arched, spreading,
thinner, on the % upper part with about 4 secondary nerves each side,
prominent, ascending, near the margin curved and vanishing, the
tertiary transverse nerves prominent, the lesser prominulous veins
minutely reticulate; leaves of young upright (orthotropic) shoots
larger, symmetrical, long-petiolate, the blades broadly ovate, deeply
cordate, more markedly palmatinerved, 30-50 cm. long, 21-36 cm.
broad, the petiole 10-38 cm. long, thickened-pulvinate at both ends.

Inflorescences axillary or extra-axillary on leafy, juvenile branchlets,
usually 3-6 cm. broad, with very short axis and divaricate, dichasial
and cincinnate branching; branchlets and pedicels angulate, densely
ochraceous or cinereous tomentose; peduncles very short, supporting
an articulate pedicel subtended by a bracteole; pedicels erect, to-
mentose, 3-6 mm. long; bracteoles lanceolate, subacute, rather thick,
more or less curved, densely and minutely tomentose, 1.5-2.5 mm.
long, 0.6-1 mm. broad; buds oblong-ovate, subacute, slightly 5-
angulate, densely adpressed and minutely cinereous or ochraceous
tomentose.

Sepals lanceolate or ovate-lanceolate, acute, shortly connate at
base, spreading and more or less curved-inflexed, 5-6 mm. long, 2-2.5
mm. broad, 3-nerved and sparsely pilose inside, glandular at base,
subappressed stellate-tomentose and reddish outside, the margin
minutely whitish tomentellous.

Petal-hoods 2-2.5 mm. long, 1-1.2 mm. broad, submembranaceous,
whitish rosy, or reddish with darker midrib, oblong-obovate-elliptic,
rounded-cucullate, emarginate, auriculate and incurved apex, glabrous
with a thick trifurcate midrib inside, hirtellous-pubescent with a de-
pressed tomentellous midrib outside; lamina rather carnose, red,

462 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

brownish red or purplish, ovate, rounded at apex, abruptly contracted
at base in short nail, articulate to the claw, hirtellous pubescent, 1-1.2
mm. long, 0.8-1 mm. broad, the nail 0.2—-0.3 mm. long, erect in bud.

Androecium tube 1.5-1.8 mm. long; staminodes 5 mm. Jong, brown-
ish red, usually lighter red toward the base with whitish margin,
carnose, linear-oblong, obtuse or subacute, slightly narrowed at base,
copiously covered with minute, thickish, patulous hairs, erect in bud,
3.5-4.5 mm. long, 0.6-0.8 mm. wide; filaments compressed, curved,
reflexed, glabrous, about 1-1.5 mm. long, shortly 2-furcate, 2-anther-
iferous ; anther lobes ellipsoid, 0.2-0.3 mm. long; ovary obovate-oblong,
sharply 5-costate, greenish white, velutinous-tomentose, 1.8-2 mm.
long, 1-4 mm. broad; styles whitish, about 1.7 mm. long, united, rigid.

Fruits subglobose-ellipsoid, oblong-ellipsoid, or ovoid-ellipsoid,
15-20 (10-25) cm. long, 9-12 (-15) cm. broad, green, when ripe yel-
low or brownish; pericarp hard, strongly 10-costate, the commissural
ribs thick and elevated, the 5 alternate similar but diminishing in
thickness towards the apex, the deep furrows reticulate (ligno-nerved),
deeply lacunose; pericarp composed of three layers: 1) carnose endo-
carp about 2 mm. thick, 2) woody, very hard, ribbed and nerved
mesocarp, 3-6 mm. thick, 3) firm, carnose becoming coriaceous epi-
carp 1-2 mm. thick with an outer, densely stellate-pilose epiderm,
the whole pericarp about 7 mm. thick on the furrows and 14-15 mm.
thick on the ribs; pulp surrounding seeds fibrose, yellowish, sweet,
Scenty; seeds arranged in 5 rows, complanate, ovoid-amygdaliform,
16-30 mm. long, 14-23 mm. wide, 8-13 mm. thick; embryo white;
germination epigeous.

Theobroma bicolor is unique in the genus, easy to recognize by its
small, lax crown of few whorls of horizontal, dichotomous branches
topping a naked stem, and by its leaves, fruits and flowers. The
large papery, firm leaves are whitish silvery beneath and those of
the upright shoots are larger, palmatinerved, cordate, and very long
petiolate. The flowers are small and pale red generally. The large
fruits are strongly ribbed and nerved, with a hard, woody, carved
shell; they keep the green color until ripening, when they become
yellow or brownish, falling from tree.

Common NAmpES.—In English: Patashte. Mexico: Patashte,
patashtle, pataste, petaste, patatle, petaxte, patasht, pataste de sapo,
pataste simarron, cacao malacayo, cacao blanco. Guatemala: Patashte,
pataxte, patasht, balamati, balam (Kekchi), pec (Pokonchi). Costa
Rica: Pataste, pataiste, skar-ub (Bribri), uerba (Terraba), scarbo
(Bribri), carvu (Kabekara), saparén (Estrella), erefa (G@uatuso).
Panama: Pataste, cu-lu-hu (Choko). Colombia: Bacao (general), ca-

CUATRECASAS—CACAO AND ITS ALLIES 463

cao silvestre, cacao marraco, and marraco (in Caquet4). Ecuador:
Patas, cacao blanco. Peru: Macambo, majambo, najambu. Brazil:
Cacau do Peru (Belém), cupuassti, cupua-i, cacau baff, cacao bravo.

The Anglo-Colombian Cocoa Expedition (Baker, 1953) recorded
the following names: heé-a (Maku), (Piraparandé, Taraira); a6 (Ma-
kuna) (lower Piraparand, Popeyacaé) ; la-na-pee-t4-ma-ca-la-chu-na-ni
(Yakuna) (Miritiparan4); ha-ha (Tanimuka) (Guacay4); maraca
bacao (Choc6).

Usrs.—The pulp is frequently eaten or used by natives to prepare
refreshments although its flavor is not very attractive. The seeds
are used in many places like those of cacao, giving a chocolate of
inferior quality; it is also locally used to manufacture pastry and
candy. The seeds of T. bicolor have been commercially mixed some-
times with those of cacao. They are poor in theobromine but they
have a great proportion of a good quality cocoa butter. In Guate-
mala the seeds are known in commerce as “tiger” ““wariba” or “pa-
tashte” cacao (Standley).

According to Llano (1947), in Colombia this species has been tried
for grafting but the bark heals with difficulty.

The hard shells of the pod are used locally, as containers like those
of mate or tutuma (Crescentia cujete). According to Tafalla it is
called “cacao de Castilla” in the Ecuadorian region of Uchiza, where
the shells of the pods are smoothed to be used as bowls.

Distrisution.—Theobroma bicolor is widespread in cultivation
throughout all humid tropical America, from southern Mexico to
Bolivia and Brasil. It is never cultivated extensively, but a few trees
can always be seen where cacao is cultivated, and usually also in
backyards of tropical farms and in secondary growth. It is fre-
quently found also subspontaneous in more or less open thickets. I
have always seen the species in cultivation only. Its truenative place is
uncertain. Probably it originated in Central America where it is
said to be found in primary forests. Miranda (1.c.) writes “se en-
cuentra en acaguales de las selvas altas’’ in Chiapas; Steyermark
found it in dense forests of the region of Ixcan (Huehuetenango) in
Guatemala. Another possible original region is the eastern region of
Peru and Ecuador. Ynes Mexfa recorded it from dense forests in
the Napo-Pastaza river basin. Llewellyn Williams found it spon-
taneous in secondary growths in Loreto. But thus far I have no
assurance of a place where it is incontestably native.

MEXICO: Herbarium Sessé et Mocifio No. 3620, preserved at the Madrid
Botanical Garden ‘Plantae Novae Hispaniae a Sessé, Mocijio, Castillo et Mal-
donado lectae (1787-1795-1804)-Theobroma Patastle Ie. N.” (MA, lectotype of
T. ovatifolium; BM, F, G, isotypes); the MA specimen has foliage, inflorescences
and flowers and may be considered the holotype of Theobroma ovatifolium Mocifio

464 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

et Sessé ex DC.; I select it as the lectotype; it is photograph FM No. 48412.
The Chicago specimen is a duplicate of this and has only one leaf; the very good
British Museum specimen, and two at Geneva are certainly isotypes in spite of
not bearing the name of Sessé and Mocifio and having ‘‘Theobroma Patastle,
N.E.” as only information; some specimens have the indication ‘‘Herb. Pavén,”’
having been sent by Pavén from Madrid. The collections with the number
3621 Sessé et Mocifio et al. (MA) are paratypes and may be duplicates of 3690,
having been numbered recently (in 1935); they include normal foliage and inflo-
rescences and juvenile long-petiolate, broadly cordate leaves (FM photographs
48414 and 48413). The photograph FM 30526 is from plate 485 of the Mocifio
and Sessé Flora Mexicana, No. 113 in De Candolle’s copies.

Oaxaca: Pochutla, Capital Rancho Viejo; tree 15 m., flowers purple, “cacao
malacayo,” Feb. 1941, Reko 6068 (F).

Catapas: Acagoyagua, Escuintla, “pataste de sapo,’ 22 VII 1947, Matuda
16733 (F). Ibidem, “pataste simarrén,”’ fruit only, 6-7 x 4 em., 28 VIII 1947,
Matuda 16840 (F, MEXU). Esperanza, Escuintla, ‘“‘pataste,” cultivated, 11
VIII 1947, Matuda 16690 (BR, F, MICH, MEXU, NY). Palenque, 100-150
m. alt.; tree up to 25 ft. tall; commonly cultivated around Palenque for its fruits;
fruits yellow; seed ground and mixed with cornmeal for making “pazoli’’; flower-
ing and fruiting in May; said to be common in the forests on the slopes above
Chacamax River, V 1937, Ll. Williams 9345 (F, Y).

GUATEMALA: Without locality, “cacao,” 10 V 1914, Davidson s.n. (US).
AuTa VERAPAZ: Trece Aguas, cacao plantation at Cacao, ‘“petaste”’ or ‘‘patashte,”
9 III 1914, Cook & Doyle 50 (US); Trece Aguas, near Finca Sepacuité, ‘“petaxte”’
IV 1905 (leaf from upright stem of young tree), Cook 4 (US). Ibidem, Chirujija
Oxee, near Finca Sepacuité, “balamati,” 25 V 1902, Cook & Griggs 756 (US).
Chama, 900 ft. alt.; tree 25 ft., flowers reddish, fruit large, hard shelled, ‘‘patashte’’
cultivated by Indians; used similarly to cacao, rather sweeter in flavor, 15 VI
1920, H. Johnson 237 (F, US). Cubilquitz, 350 m. alt., VI 1901, von Tuerckheim
7824 (GH, K, NY, US). Vicinity of Sibicte, 370 m. alt., small tree 25 ft. tall,
leaves firmly membranaceous, deep green above, gray silvery beneath; flowers
with grapy-purple calyx, “balam” (Kekchi), “patasht’’ (Spanish), cultivated,
12 III 1942, Steyermark 44941 (F).

HUEHUETENANGO: Sierra de los Cuchumatanes, 150-200 m., between Ixcan
and Rfo Ixcan; tree 30 feet tall, leaves membranaceous, papyraceous, dark green
above, silvery green beneath; dense rich forest, 23 VII 1942, Steyermark 49317
(F, US). Huehuetenango, 1800 m. alt., seeds in market brought here from the
Pacific Coast, Standley 82446 (F).

Santa Rosa: Region of Platanares, between Taxisco and Guazacapdn, 220
m. alt.; wet forested quebrada, simple shrub 3-4 m. tall; escaped here, 3 XII 1940,
Standley 79069 (F).

SUCHITEPEQUEZ: Mazatenango, cultivated, III 1865, Bernoulli 94 (F, G, NY).
Ibidem, III 1865, Bernoulli & Cario 3145 (GOET).

BRITISH HONDURAS: Conservation Forests H. 2192/29 (F, GH, UC).
Stann Creek Valley, Big Eddy Valley; tree 13 inch diameter, fruit 8 inch long,
“‘pataste,” ‘mountain cacao,” 16 XII 1840, Gentle 3464 (F, GH, MICH, MO, NY,
U).

EL SALVADOR: Sonsonate, cultivated, “patashte,’’ ‘‘pataste,’’ 20 IV 22,
Calder6én 23610 (F, GH, MO, NY, US); ibidem, “pataste,”’ 1922, Calderén 627
(GH, NY, US).

COSTA RICA: Peninsula Osa ad Golfo Dulce, circa Puerto Giménez, ad litus;
arbor 6-8 m., floribus purpureis, 15 IV 1930, Cudofontis 92 (WU). Vicinity of
Guapilés (prov. Limén), 300-550 m. alt.; planted tree 25 ft., flowers dull red 12,

CUATRECASAS—CACAO AND ITS ALLIES 465

13 III 1924, Standley 37374 (US). Puerto Giménez de Osa and vicinity, 141V 1930,
Brenes 12333 (Herb. Nac. C.R. 212) (F, NY). Edge of the road to Tuis, 650 m.;
tree with spreading branches, “‘pataste”’ or “‘pataiste,’’ XI 1897, Tonduz 11304 (F).
Tucurrique, grassland at Las Vueltas; tree 40 cm. diam., 15-20 m. high, 635 m.
alt., “pataste,”’ ‘‘pataiste,’’ III 1899 Tonduz 13110 (G, P, US). Hacienda Balti-
more (Limén), 10 m. alt., in a small plantation of this species, 8 VII 1949, Hold-
ridge s.n. (TURRI). La Lola, planted, 6 XI 1961, Cuatrecasas & Paredes 26534
(US).

PANAMA: Bocas del Toro, in Laguna de Chiriquf and its neighborhood,
Pope’s Island, XI, XII 1885, Hart 158 (US). Darién, Headwaters of Rio Chica,
500-750 ft.; tree 35 ft.; flowers dark red; cultivated by Choco Indians, ‘‘cu-lu-hu,”
Allen 4593 (G, MO, NY). Canal Zone, along Cafio Quebrado, 2 XII 1914, Pittier
6883 (GH, NY, US).

TRINIDAD: L’Eranche Est, Sangre Grandre, 10 VII 1929, Boehlmer 12229
(TRIN). Blue Basin, 24-3 miles distant, 21 IX 1928, Lange 12056 (TRIN).
Grounds of I.C.T.A., River Estate Diego Martinez, field 19; fruits 19.3x 13.3 cm.,
yellowish at maturity, specimen from seeds brought from Jinogojé, Apaporis,
Colombia, 31 VIII 1961, Cuatrecasas, Cope, & Bartley 25784T (US); same field;
calyx pale red, hoods with darker midrib, petal-lamina very small, brownish red,
staminodes brownish red, lighter reddish toward the base, with whitish margin,
styles whitish, ovary greenish white, 1 IX 1961, Cuatrecasas & Cope 25795 (US).
Tree from seeds from La Pedrera, Colombia; fruits 16.2 x 14 cm., 31 VIII 61,
Cuatrecasas, Cope, & Bartley 25787T (US). Field 2; fruits 15 x 10, 16 x 10.8,
16.5 x 10, 18.2 x 11.2, peduncle 0.7-0.9 cm., 31 VIII 61, Cuatrecasas, Cope,
& Bartley 25786T (US).

SURINAM: Paramaribo Gardens, cultivated, VI 1910, Stockdale s.n. (K, U).

COLOMBIA: Antioquia: Savaletas, 100-500 m., XII, Lehmann 7909 (K).

Toitmma: Ibagué, II 1916, ‘cacao silvestre,” Dawe 83 (US).

Huiua: Valle del Magdalena, Garzén, IV 1845, Goudot s.n. (G, P).

Ex Va.uE: Palmira, Granja Agricola, 900 m. alt. “bacao,” Duque Jaramillo
1205 (F). Ibidem, X 1943, Llano s.n. (COL, F). Palmira, 925 m., cultivated in
grounds of experiment station (said to have been brought from the Choc6); tall,
erect tree, leaves hanging vertically, almost white beneath, flowers dark red with
maroon staminodes, fruit oblong, green with the furrows strongly rugose, 29 X 1944,
Fosberg 21310 (NY, UC, US). Palmira; tree 4 m., “bacao,” I 1947, Duque Jara-
millo 4403A (COL). Ibidem; tree 3 m., flowers red, ‘‘bacao,’”’ 3 XII 1947, Car-
defiosa, Murgueitio, & Barkley 17C934 (COL, F). Cartago, Goudot s.n. (P, WU).
Pacific Coast, Rfo Calima, La Trojita, left side of river, 5-20 m. alt.; tree 3 m.,
“bacao,” 27 II 1944, Cuatrecasas 16526A (F, VALLE) (fruit only). Buenaven-
tura (and Tumaco) 0-400 m., fl. perpetual, Lehmann 9021 (K, NY).

Cuocé: ‘“Nvelle. Grenade, Chocé,” ‘‘bacao,’’ Humboldt and Bonpland, s.n.
(P, holotype). Bonpland, probably isotype, from Paris (F). Near Istmina, road
to Cértegui, in forest, 75 m. alt.; tree 6 m., flowers red, “bacao,” 3 VII 1944, Garcia
Barriga 11178 (COL, US). Chocé (i Barbacoas), 25 m. alt., “bacao,’”’ IV 1853,
Triana 5333 (-3) (BM, COL). Ciudad Mutis, 27 X 1946, Romero Castafieda
s.n. (COL).

NariNo: Pacific Coast, Amarales, “‘bacao,” 1866, Triana s.n. (BM, BR, COL,
G, K, NY, WU).

Caqueri: Solano, 3 km. SE of Tres Esquinas, on Rio Caquetd, below mouth
of Rio Orteguaza, wet tropical forest of Amazon basin; tree 8 m. high, 10 cm.
DBH,; dark brown, smooth, lichen patches, flowers red; watermelonlike fruit 18
em. long, 10-11 cm. broad, yellow, edible; Indians plant it; secondary lowland
forest, river bank; possibly an escape from cultivation; several cultivated trees

466 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

seen March 13 walk near Rio Caquetd, 2 km. s. of Solano; ‘‘cacao marraco,”’
6 III 1945, Little & Little 9598 (NY, US).

Putumayo: Vicinity of Mocoa;small tree 4 m., sterile; apparently cultivated,
17 III 1958, Holliday & Cope T 77 (COL, TRIN, US).

Vaupts: Rio Negro, near San Felipe, Cafio Mayabo, river level; tree 4—5
years old, introduced on a house site, 27 X 1952, Baker 34 (COL, F, TRIN).
Rio Infrida, right bank below mouth of Cafio Caribe; tree 25 m. high, 1 m. diameter
at ground level, growing on slope just above high watermark; overtopping the
surrounding forest; not thought to be wild. 23 11953, Bartley & Holliday T66
(COL, TRIN, US). Rio Vaupés, opposite confluence with Rfo Papurf, Yavaraté,
Silesian Mission Sio Miguel; trees 3-4 years old from seed brought by Indian
from the interior of Colombia, 20 II 1952, Bartley & Holliday T 47 (COL, TRIN,
U, US). Cafio Umufia, Rfo Piraparand, river level; cultivated tree in Indian
garden, 8IV 1952, Baker & Cope lla (TRIN). Jinogojé, on Rfo Apaporis, small
tree 15 ft. cultivated in Indian garden, 23 VIII 1952, Baker & Cope 2 (TRIN).
Rfo Apaporis, Gino-Gojé, between the rivers Piraparané and Popeyaca, 250 m.
alt.; tree 4 m., leaves white below, 3-11 IX 1952 Garcia Barriga 14416 (US).

Amazonas: Rfo Caquetd4, La Pedrera, river level; tree 30 ft., presumed cul-
tivated, 1 X 1952, Baker & Cope 26 (COL, F, TRIN, US). Rfo Igaraparand,
vicinity of La Chorrera, 180 m. alt.; small tree, cultivated, ‘‘marraca,’”’ 4-10 VI
1942, Schultes 3922 (COL, F).

ECUADOR: Muller s.n. (K). Balao; arbor 40 m., in forest, flowers purplish,
“cacao blanco,” I 1892, Eggers 14244 (A, L, LE, M, US). Rio Sucumbfos, be-
tween Putumayo and Quebrada Teteyé, 260 m., ‘‘cacao’’ (Kofdn), 29 III 1942,
Schultes 3471 (NY).

Napo-Pastaza: Between Tena and Napo; tree about 15 m. high, petals
dark red, staminodia blackish brown, 5 I 1940, Asplund 10271 (S). NE of the
province, Tiputini-Lagartococha, 20 I-5 IJ 1953, Fagerlind & Wibom 2371 (8).
Cantén Napo, near Tena, 400 m. alt., dense forest; tree up to 18 m. high, fruit
called ‘‘Patas,” size of small watermelon, the pulp being eaten and the seeds
cooked or raw much appreciated, 2 II-VI 1985, Mexia 7214 (F, P, UC, US).

BRAZIL: “Brazil, Dr. Martius,” no data (G). Manaquiry forest; 15-30 ft.,
leaves white beneath, flowers dark purple, Spruce s.n. (KX).

Amazonas: Rio Negro, Barcellos, matta, cult.?, 27 VI 1905, Ducke 7202
(BM, MG, US). Ibidem, Ducke 7202-B (BM, MG). “Prov. Rio Negro, in
sylvis, Martius Obs. 28823, Iter-Brasiliensium 319, Martius [862, 863, 864, 865]
(M). Rio Negro, Lagos, ‘‘cacao bravo,’’ 5 VIII 1874, Traill s.n. (GH). Rio
Negro, Schomburgk 870 p.p. (BM, G, GH, GL, K, OXF, P, US). Sad Antonio de
Iga, capueira, ‘“‘cupuasst,’’ 3 VIII 1906, Ducke 7638 (MG). Municipio Sio Paulo
de Olivenga, basin of creek Belem; tree 40 ft. high, trunk 5 inch diam., planted by
Indians 26 X—XII 1936, Krukoff 9019 (A, BM, F, G, K, LE, MICH, MO, NY, P,
US). Manaos, Campos Salles, 15 m., 20 VIII 1928, Luetzelburg 23895 (M).
Manaos, Agricultural Experiment Station, 25 m. alt.; tree 30-35 ft., fruit oblong
on ultimate branches (25 x 10 em.), cult.?, 18 X 1929, Killip & Smith 30006 (GH,
NY, US), fruit collection 681 (US). Experimental Garden of Nord Brazil, culti-
vated and wild in forest, ‘cacao d’Anta,” 20 VIII 1928, Luetzelburg 23065 (M).
Ega, “‘colitur cireum Indorum villas, Oct. 1830,’ Poeppig 2746 p.p. (WU); “In
Bras. tropica fl. Amazonas Oct. 1839,” Poeppig 2746 pp. (GOET). Ega, Poeppig
2746 p.p. (F, LE). Teffé, forest, 29 VI 1906, Ducke 7397 (BM, G, MG, US). Fonte
Béa, firm land, medium-sized tree, ‘“cacau baft,’’ 28 III 1945, Frées 20625 (F,
JAN, K, NY, US). Rio Juru4, Santa Clara, cultivated, ‘‘cupuagd’’ Baum, X 1900,
Ule 5030 (BM, G, HBG). Rio Jurud, Gaviao; flowers purple, ‘“cupua-i,’’ III 1875,
Traill 60 (K, P). Rio Sapo, 21 II 1874, Traill 60 (K). ‘‘Ad oram meridionalem

CUATRECASAS—CACAO AND ITS ALLIES 467

flum. Amazonum ad ostium flum. Solimoes,’”’ VI 1851, Spruce 1609 (BM, K, M,
P, WU).

Ceara: Ceard, VIII-XI 1838, Gardner 870 (G).

Pari: Belém, Horto do Museu Goeldi, arvore 483; medium size tree, granate
flowers, 10 X 1957, Cavalcante 310 (MG, US). Belém, cult., Pires & Black 746
(IAN). Belém, I.A.N. cult., “‘cacau do Peri, 6 XI 1952, Pires 4340 (IAN, NY).

PERU: ‘Theobroma cordata del Perti, sp. nov.,”’ Ruiz & Pavén (BM). ‘‘Peru-
via, Herb-Pavén” (G). Perou, Pavén 617 (G). 1909-1914 Weberbauer 6245.

Loreto: Mishuyacu, near Iquitos, 100 m.; forest, tree 6 m. high, flowers
dark violet and rose, clearing, ‘‘“macambo,’’ V-VI 1930, Klug 1523 (F, NY).
Florida, Rfo Putumayo at mouth of Rio Zubineta, 200 m.; forest clearing, tree
4 m., flowers garnet, ‘‘“macambo,” III-IV 1931, Klug 2021 (A, F, GH, K, MICH,
MO, NY, 8, US). Paraiso, upper river Itaya, 145 m.; “najambu,” 1 X 1929,
Ll. Williams 3346 (F, US). Amazon River, Caballo-Cocha, 6 VIII 1929, Zl.
Williams 2149 (F, GH, US). Maynas, Poeppig (L), Poeppig 18 (BM). Marafién
River from Iquitos to the mouth of Rio Santiago at Pongo de Manseriche, ca.
77°30’ W., 1924, Tessmann 4079 (NY, 8).

Section 2. Oreanthes

Theobroma sect. Oreanthes Bernoulli, Uebers. Art. Theobroma 7, 1869.
Figure 4; Map 1

Sect. Bubroma subsect. Oreanthes (Bernoulli) Pittier, Rev. Bot. Appl. 10(110) :779.
1930.

Petal-lamina sessile, large. Petal-hood 3-nerved. Staminodes
linear-subulate or lanceolate, erect in estivation. Filaments 3- or
2-antheriferous. Fruit more or less angulate or costate with coriaceous,
tomentose pericarp, the endocarp rigid, thin-lignose. Cotyledons
epigeous at germination. Leaves densely, minutely, stellate-
tomentose beneath inside the reticulum. Inflorescences many-
flowered, on the trunk or, in few cases, small and on leafy branches.
Sympodial growth of stem by orthotropic, adventitious, lateral-
subterminal shoots. Primary branches ternate, deciduous when old,
leaving an unusually long, naked stem. Leafy crown loose, flat.
Secondary branching dichotomous.

Tyre species: Theobroma speciosum Willd. ex Spreng.

2. Theobroma sylvestre Mart. Ficures 4, 9, 13, 16, 18; Mar 4; Puare 1

Theobroma sylvestre Mart. in Buchner, Repert. 35:24. 1830; Linnaea, Litt.
Ber. 32. 1831; Bernoulli (1869) 14, pl. 7, fig. 1; Schumann in Mart. (1886)
78 (as T. silvestre); Jumelle (1899) 34; De Wildeman (1902) 98.

Theobroma Spruceana Bernoulli, Uebers. Art. Theobroma 9, pl. 3, fig. 1. 1869;
Ducke (1925) 131; (1940) 270, pl. 3; (1954) 13; Chevalier (1946) 276;
Addison & Tavares (1951), pl. 6, fig. 1, pl. 6, fig. A, pl. 12, fig. 4; Leén (1960)
318, 319, fig.

Theobroma nitida Bernoulli, Uebers. Art. Theobroma 15, pl. 7, fig. 2. 1869.

Theobroma Martii Schum. in Mart. Fl. Bras. 12(3):78. 1886; Jumelle
(1899) 35; De Wildeman (1902) 98.

Theobroma speciosum var. Spruceana (Bernoulli) Schum. in Mart. Fl. Bras.
12(3):75. 1886; Jumelle (1899) 32, fig. 16; De Wildeman (1902) 95.

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

468

nd plagiotropic

B, sylvestre (US-1693105); c, sylvestre (Martius, type).

Ficure 13.—Leaves of Theobroma, X 1%: a, speciosum, from orthotropic a

branches (US-1031918);

CUATRECASAS—-CACAO AND ITS ALLIES 469

50° 49°

io

o*

20°

|

Map 4.—Distribution of Theobroma sylvestre, @; T. bernouilliit subsp. bernouillii @;
T. bernouillii subsp. asclepiadiflorum ©; T. bernouillit subsp. capilliferum CQ; T.
velutinum A.

Typres.—Martius, Brazil, Iter Brasil. 322 (in M, no. 891) (Photo
F. M. 19644). Spruce 166, Brazil, Paré, Obidos (of T. Spruceana).
Martius, Brazil, Iter Brasil. 322 (in M, no. 890) (Photo F. M. 40709)
(of T. nitida).

Small- or medium-sized tree up to 12 m. high, sympodial growth by
lateral, subterminal upright shoots; primary branching ternate;
terminal, leafy and floriferous branches dichotomous, terete, rugulose,
dark brownish, the hornotinous ones pulverulent, stellate-tomen-
tellous, soon glabrate or glabrous; stipules small, soon deciduous.

Leaves distichous, firmly coriaceous; petiole short, robust, appressed
stellate-tomentose, 5-10 mm. long; blades elliptic-oblong with obtusely
cuneate, asymmetrical base, often ovate-oblong or oblong-ovate with
rounded, asymmetrical base, attenuate and acutely acuminate at
apex, the margin entire or slightly sinuose, 12-30 cm. long, 5-8 (—13)
cm. broad, the acumen 1.5-2.5 cm. long, rather lustrous above, pale
brownish when dry, glabrous, the costa and secondary nerves sub-
filiform, prominulous, the other nerves almost unnoticeable, paler
beneath with tawny nervation, the costa very prominent, the second-
ary nerves about 6 pairs, prominent, ascending, decurrent, arching
and uniting near the margin, the inferior pair forming a much more
acute angle, the transverse tertiary nerves distant 5-10 mm. from

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

470

. a
SOS eet
wpe dueeer
-

enya ee .

[FicureE 14]

CUATRECASAS—CACAO AND ITS ALLIES 471

each other, thin and prominent, the lesser veins forming a minute,
prominulous reticulum; midrib, secondary and tertiary nerves gla-
brous, those of the fourth rank subglabrous, the lesser reticulate veins
and the areoles covered by dense, minute, whitish, sericeous tomentum
of stellate hairs.

Inflorescences small, axillary or extra-axillary on small branches;
panicles short, 1-2 cm. long, branched from base, the branchlets 1-8,
fasciculate, crowded, scarcely ramulose, densely stellate-tomentose,
bracteolate at joints, bracteoles ovate, or ovate-lanceolate, 1-2 mm.
long, stellate-tomentose; pedicels 2-5 mm. long, moderately thin,
densely tomentose; sepals shortly united at base, thick, sublanceolate-
oblong, subacute, involute at margin, ferruginous, stellate-tomentose
outside, glabrous inside except for the minutely stellate-tomentose
margin and glandular trichomes at base, curvate and spreading at
anthesis, 7-9 mm. long, 2-2.5 mm. broad.

Petal-hood thick-membranaceous, trinerved, pale and reddish
striate, rugulose, papillose, sparsely pilose outside, oblong-obovate,
cucullate-rounded at apex, 4 mm. long, 2 mm. broad; petal-lamina
brownish red or rose, rotundate-subreniform, minutely crenulate,
rugose, 2 mm. long, 2.2-2.5 mm. broad, suddenly constricted into a
short claw, articulate at base; androecium tube 1.5-2 mm. high;
staminodes brownish red or rose, linear-subulate, abruptly narrowed-
acuminate at apex, the acumen often curled, densely and minutely
pilose-muricate throughout, 5-6 mm. long, 0.7-0.8 mm. wide; fila-
ments 1—1.2 mm. long, glabrous, arched, diantheriferous, the lobes
of the anthers ellipsoid, 0.4 mm. long; ovary ovoid-ellipsoid, 5-ridged,
ferruginous-tomentose, 1.5-2 mm. long, the apex whitish tomentulose;
styles 5, thin, acute, connivent, united only at base.

Fruits elliptic-globose, or subglobose, about 6 x 5.8 cm., rounded at
apex, umbilicate at base, the pericarp coriaceous, minutely and densely
tomentose, glaucous when ripe, about 5 mm. thick, the inner and
outer layer hard, thin, the middle one carnose; seeds ovoid-oblong,
1.4~2.1 cm. long and 0.5 x 0.9 mm. broad; pulp rather sweet, scentless,
white; fruiting peduncle about 1.8 cm. long, 7 mm. thick.

At the Botanische Staatssammlung in Munich there are several
specimens from Martius’ trip to Brazil which bear the annotation

Figure 14.—a-1, Theobroma glaucum (Baker & Cope 11 and 18): a, petal from inside X 5;
B, bud, X 2; c, pistil, X 5; p, petal laterally, X 5; x, styles, X 5; F, androecium,
X 5; G, 2-antheriferous stamen, X 10; un, 3-antheriferous stamen, X 10; 1, sepal from
outside and inside, X 2, K-R, T. speciosum (Archer 7619): x, petal from inside, X 5;
L, styles, X 5; m, bud, X 2; Nn, petal laterally, & 5; 0, androecium, X 5; p, stamen,
X 10; g, sepal from outside and inside, X 2; R, pistil, X 5. s-y, T. velutinum (B. W.
1161): s, petal from inside, X 5; 1, pistil, X 5; U, petal from outside, X 5; v, petal,
laterally, X 5; w, androecium, X 5; x, stamen, X 15; y, sepal from inside and outside,
X 2.

680-695—64—__7

472 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Figure 15.—a-1, Theobroma bernouillii subsp. capilliferum (Holliday 142): a, B, c, peta
from inside, outside, and laterally, X 5; p, androecium, X 5; £, stamen, X 10; F, anther:
X 20; c, pistil, X 5; m, sepal from inside and outside, X 2; 1, bud, X 2. k-R, 7
bernouillii: subsp. bernouillii (Pittier 4105): x, 1, mM, petal from inside, outside an
laterally, X 5; N, androecium, X 5; 0, stamen, X 10; p, anther, X 20; a, sepal fror
inside and outside, X 2; r, pistil, X 5. s-z, T. bernouillii subsp. ascleptadiflorur
(Wedel 1535): s, T, u, petal from inside, outside and laterally, X 5; v, androeciun
x 5; w, stamen, X 10; x, anthers, X 20; y, pistil, X 5; yy, bud, X 2;z, sepal fror
inside and outside, X 2.

CUATRECASAS—CACAO AND ITS ALLIES 473

Ficure 16.—Fruits of Theobroma, X ¥%: a, bernouillii subsp. asclepiadiflorum (Lucas 1);
B, bernouillit subsp. capilliferum (Cuatr. 17034); c, glaucum (Froes 20645); pv, sylvestre
(Froes 20463); £, speciosum; ¥, velutinum (Benoist 516); c, nemorale (Cuatr. 21291).

474 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

“Tter brasiliensis” and then the number 322 at the foot of the label;
they are named by Martius as “T. sylvestris Martius.” All the
specimens agree with each other essentially, although some of them
have oblong leaf blades, slightly asymmetrical or symmetrical at the
base, and others have broad, oblong-ovate blades, very rounded and
asymmetrical at the base; in all of them there is a longer distance
between the basal pair of secondary nerves and the next pair than
between the other pairs of secondary nerves; two of these specimens
bear typical rounded, smooth fruits. One of the specimens which
shows the number 891 on a mounting strip (Photo F. M. 19644) may
be considered as the holotype of 7. sylvestre (lectotype); it has a leafy
branch with the basal half of a fruit, and a loose, broad, obovate-
oblong leaf. Bernoulli, without obvious reasons, considered one of
the specimens [890] (Photo F. M. 40709) a different species and de-
scribed it as 7. nttida Bernoulli. Another of these specimens [871] was
named by Schumann as 7. grandiflorum, and it is mixed with an
authentic 7. grandiflorum specimen on the F. M. 19641 Photograph.
After a close examination of the series of Martius types, it was clear
to me that there do not exist basic differences between them and 7.
spruceanum Bernoulli. Most of Martius specimens have somewhat
thinner and smaller leaves than the current collections, variations
which can be attributed to the habitat and to the age of the collected
branches.

Schumann listed 7. sylvestre Mart. under his ‘Species dubiae.”

The binomial Theobroma Martti was published by Schumann as a
new name for 7. nitida Bernoulli, although due to some typograph-
ical error this name was not quoted as synonym. But Schumann
cited “l.c. 15”? and quoted unchanged the diagnosis given by Ber-
noulli for 7. nitida on page 15. Schumann considered the name a
homonym of his 7. nitidum (Poepp. & Endl.) K. Schum. (Abroma
nitida Poepp. & Endl.).

Theobroma sylvestre has been confused with T. speciosum, which is
usually a larger tree, but its small axillary inflorescences, smaller,
scentless and paler brown-reddish flowers, and its green-bluish fruit
at maturity, distinguish it readily.

ComMon NAMES.—‘‘Cacao azul’ is generally used. Other names
recorded are: Cacau azul, cacaui, cacaut, cacao-ht, cacao rana,
cacau rana, cacau bravo. Cacao azul refers to the color of the
mature fruit (bluish).

Usrs.—No special uses are recorded.

DisrripuTion.—More or less frequent along the Amazon River
and lower part of its tributaries from Santarém to Tonantins, east-
ward from the mouth of the Iga. It grows on elevated ground in
rather dry places, and appears frequently in secondary stands.

CUATRECASAS—CACAO AND ITS ALLIES 475

According to Ducke, it is common around Manaos, especially in
relatively dry thickets on clay soils poor in silica.

BRAZIL: Amazonas: “Martius Iter brasiliensis 322: Theobroma sylvestris
Mart. Observ. 2832, Cacao sylvestris Aubl., Cacao Rana Incol. Habitat in sylvis
ad fl. Solimoes, Prov. Rio Negro,” Martius [887] (Photo F. M. 40708) (M,
syntype of T’. sylvestris Mart.). ‘Martius Iter brasiliensis 322: Theobroma syl-
vestris Mart. in silvis ad Fl. Solimoes,” Martius [891] (Photo F. M. 19644)
(M, lectotype). Ibidem Martius [871] (Photo F. M. 40707) (M, syntype).
Ibidem Martius [888] (M, syntype). Ibidem Martius [889] (M, syntype).
“Martius Iter Brasiliensis 322, in silvis ad fl. Solimoes,” Martius [890] (Photo
F. M. 40709) (M, type of 7. nitida). Manaos, Mata do Aleixo, “cacau azul,”
16 III 1945, Frées 20556 (IAN, NY). Manaos, Colonia Joao Alfredo, rain
forest in noninundatable ground; small tree, flowers pale brown-rose, ‘cacao
azul,’ 6 XII 1935, Ducke 100 (A, F, IAN, K, MG, MO, S, US). Manaos, in
noninundatable forest; rather large tree with weak trunk, flowers brownish-
flesh color, ‘‘cacao azul,” VI 1932, Ducke 103 (F, Y). Manaos, matta, vicin-
ity Igarapé da Cachoeira Grande, elevated ground; small flowers on trunk and
branches, 14 X 1912, Ducke 12187 (G, MG). Fonte Béa, matta, elevated ground;
tree 4 m., 10 cm. diam., “cacao azul,” 5 IV 1945, Frées 20655 (F, K, US). Ibi-
dem, “the Indians say that it was introduced from Japura river,” Frées 20463
(US). ‘Campo experimental do I.A.N., introduced from Rio Negro, Frées
34949 (IAN). Maués; small tree, flowers red, wood hard, “cacaurana,” “cacau
azul,” 30 XI 46, Pires 136 (IAN). Rio Madeira, Rio Canuma, Borba munic-
ipality; tree 4 m., elevated ground, 12 XI 1927, Frées 33788 (IAN). Rio
Demeni, Sumauma, Barcelos municipality; tree 8 m., fruits, flowers rose, 30 IV
1952, Frées 28382 (IAN). Rio Tonantins, Frées 25554 (IAN).

66° 50° 49°

to"

148s_ yiBOF.
Pa hed

if

ones
)

‘YSNILNY

10"

20°

|

Map 5,—Geographical distribution of Theobroma speciosum @ and T. glaucum ©.

Pall

476 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Par&: “In vicinibus Obidos, Prov. Para, Dec. 1849,’ Spruce 166 (Photo
F. M. 40702) (M, holotype and lectotype of Theobroma spruceanum Bernoulli;
isotypes BM, WU). Obidos, “cacao azul,” 7 I 1904, Ducke 4878 (BM, G, MG,
P, US); ibidem, matta, 11 V 1905, Ducke 7216 (MG). Ibidem, rain forest, ele-
vated ground; small tree, fruit green glaucous when ripe, flowers pale brownish
reddish, ‘‘cacao-azul,” 10 I 1920, Ducke 14734 (S, U). Sao Jorge, municipality
of Faro; small tree in rain forest, flower dark red on branches, “cacau azul,”’
11 XI 1950, Black & Ledoux 50-10644 (IAN, UC, U). Oriximind, Las Trombetas,
flowers on trunk and branches, ripe fruit green, “cacao azul,’ ‘“‘cacao-hu,”’ 8 XII
1906, Ducke 7822 (BM, G, MG, US). Lazo de Faro, above Tanaenera, forest on
elevated ground, “cacau azul,” 12 II 1910. Ducke 10669 (MG). Alenquer,
Estrada da Vila do Curud4, municipality of Obidos, sandy ground with vegetation
of Bertholletia and Attalea; tree 2.5 m. in rain forest, 4 III 1953, Frées & Filho
29465 (IAN, U). “Km 23 da BR-17 Est. a direita,” firm land, sandy, natural
forest; flowers yellowish red, edible fruits, 12 X 1955, “cacao bravo,” INPA
(Dionisio) 2125 (IAN).
3. Theobroma speciosum Willd.
Fiaures 5, 12, 13, 14, 16, 18; Mar 5; Puatres 2, 3
Theobroma speciosum Willd. ex Spreng. Syst. Veg. 3:332. 1826; Bernoulli
(1869) 8, pl. 3, fig. 2; Schumann in Mart. (1886) 74; Jumelle (1899) 30,
fig. 16; De Wildeman (1902) 95; Huber (1906a) 273; Ducke (1925) 130;
(1940) 270, pl. 1, fig. 3, pl. 2; (1954) 13; Addison & Tavares (1951), pl. 4,
fig. 2, pl. 6, fig. B, pl. 12, fig. 6; Cuatrecasas (1956) 658.
Theobroma quinquenervia Bernoulli, Uebers. Art. Theobroma 8, pl. 3, fig. 3.
1869.
Theobroma speciosum var. quinquenervia (Bernoulli) Schum. in Mart. Fl.
Bras. 12(3):75. 1886: Jumelle (1899) 32, fig. 16; De Wildeman (1902) 95.
Theobroma speciosum var. coriaceum Huber, Bol. Mus. Goeldi 4:586. 1906.
Theobroma guianensis sensu Chevalier, Rev. Inter. Bot. Appl. 26:274, 1946;
Leén (1960) 318, 319, fig., pro parte, non Gmelin.
Sapokaia brasiliensis Rich. ex Chevalier. Rev. Int. Bot. Appl. 26:275. 1946,
as synonym.

Typres.—Siber: ‘Hoffmannsegg’ in Herbarium Willdenow no.
3680 (B). Spruce 1737, Brazil, Barra do Rio Negro (of 7. quinque-
nervia). Huber 1567, Peru, Ucayali (of var. coriaceum).

Tree up to 15 mm. tall; trunk about 20 cm. in diameter with light
gray, smooth bark; sympodial growth by lateral, subterminal upright
shoots; primary branches ternate, usually furcate, spreading; branch-
lets terete, smooth, more or less pulverulently stellate-pilose, later
glabrate, grayish brown; crown rather narrow; stipules subulate,
short, stellate-tomentose, soon deciduous.

Leaves firmly coriaceous, distichous; petiole robust, subterete,
densely stellate-tomentulose, 8-14 (—-20) mm. long, 2-4 mm. thick;
blades usually large, ovate-oblong or elliptic-oblong, broadly rounded
or very obtuse and asymmetrical at base, exceptionally (in ortho-
tropic branches) cuneate and symmetrical, attenuate near the apex,
abruptly and acutely acuminate, the margin entire or very slightly
sinuose, often slightly revolute, 20-40 cm. long, 7-18 cm. broad, the

477

CUATRECASAS—CACAO AND ITS ALLIES

SY

TY)

SS i

2 +
pan aa ots a
se iratelree

<9 68 Se

Sh

“ENV EY ZF if. \

SNE O : oa sd Ay \E
PALL ia a yf & \
a i. I Pd i rams
ky } i ¢

~, 7 > a i =
, A TANG
*.
P.
ra

Figure 17.—Leaves of Theobroma, X14: a, glaucum, from lateral branches (Bartley

& Holl. 74); B, glaucum, from orthotropic branches (Baker 37); c, bernouillii subsp.

asclepiadiflorum (Wedel 681); p, bernouillii subsp. bernouillii (Pittier 3199); E, bernouillii

subsp. capilliferum (Cuatr. 16160).

478 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

acumen 1.5-2.5 cm. long; lustrous above, green, usually pale brown or
olivaceous when dry, glabrous, or with a few stellate hairs on the
nerves, the costa and main nerves filiform, conspicuous, the minor
veins less noticeable or obsolete, pale cinereous beneath with tawny
nervation, the costa thick, very prominent, the 6-8 pairs of secondary
nerves prominent, subcurvate ascending, near the margin thinner,
curving, decurrent and anastomosing, the inferior pair stronger,
forming together with the midrib a trinerved base, the transverse
tertiary nerves thin but prominent, 5-15 mm. distant from each
other, sometimes the basal more conspicuous, forming together with
main nerves a basally 4- or 5-nerved leaf; minor veins prominulous,
reticulate; main nerves glabrous or subglabrous with sparse, stellate
hairs and scattered callose dots, the tertiary glabrous or subglabrous,
the minor reticulate veins and the areoles covered by dense minute
whitish sericeous tomentum of stellate hairs.

Inflorescences on trunk, forming delicate panicles gathered in many-
flowered bunches borne on woody, short, tuberculose branches,
often very showy with up to 250 dark-red or purplish-red fragrant
flowers; panicles 3-10 cm. long, dichotomously furcate-branched from
the base, the branches thin but rigid, reddish and covered with a
minute, whitish tomentum, the terminal branchlets (peduncles) thin,
flexuous, 10-25 mm. long, 3-bracteolate and articulate to the pedicel
at apex; pedicels slender, 5-20 mm. long, minutely tomentulose;
bracteoles minute, linear or linear-triangular, 1-2 mm. long, 0.2—
0.5 mm. wide, very soon deciduous; buds ovoid, 7-9 mm. high, rather
reddish, with 5 longitudinal, white-tomentose, prominulous com-
missural lines, sparsely stellate-pilose; sepals rather thick, oblong,
subobovate-oblong, attenuate toward the base, abruptly narrowed
and subobtuse at apex, the margin incurved with a minutely, whitish
tomentulose strip inside, the apex shortly cucullate-inflexed, the inside
purplish and glabrous, except for glandular trichomes at base, the
outside sparingly stellate-pilose, usually one free and the others
united one third or almost completely by pairs, 10-12 mm. long,
3.5-4 mm. broad; petal-hoods thick-membranaceous, trinervate,
whitish with red lines, oblong-obovoid, attenuate-clawed at base,
rounded cucullate at apex, with sparse, thin, spreading hairs outside,
6-7 mm. long, about 4 mm. broad; petal-lamina rather thick, red or
dark red, transversely elliptic, subtruncate, slightly emarginate and
mucronulate at apex, abruptly cuneate-attenuate at base, entire or
slightly erose at margin, conspicuously (especially by transmitted
light) reticulate-veined, 5-7.5 mm. long, 7-9 mm. broad; androecium
tube thick, about 2-3.5 mm. high, sparsely stellate-pilose; staminodes
purplish red, subulate, thick, the apex thinner and curled, minutely
muricate-pilose, 5-7 mm. long, 1.2-1.8 mm. wide; filaments glabrous,

479

CUATRECASAS—CACAO AND ITS ALLIES

cS

*

aS
a

ante
VJ

13.4 ue aS

.

ry
ao

Be
Kd
he N

SoA

eae |
PRK? oe

qo

a}

a, Theobroma bicolor (Killip & Smith

T. velutinum (Benoist

D,

T. speciosum (Ule 9629);

c,

D X 20.

and C X 30

ot aan
AN ey Ly
Kos LAP iy
B, ae Sa al
SAREE iN ks

pate SU

==,

We

hoe
as
F ag
i,

AX

.
AR IA
' ER Re Ee . :
RE IG as Ra een Aye,

7]

T. sylvestre (Ducke 100);

B,

?

.—Indument on the underside of leaf in:

B

30006) ;
516). A,

Ficure 18

480 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

about 2 mim. long, curved, dilated at base, minutely 3-furcate at apex,
triantheriferous; anther lobes ellipsoid, about 0.4 mm. long; ovary
ovoid-ellipsoid, 2-3 mm. long, 5-ridged, whitish velvety-tomentose;
styles 5, subfiliform, 1.2-2 mm. long, glabrous, connivent, only united
at base.

Fruit globose-ellipsoid, about 10 cm. long and 7-8 cm. broad, almost
smooth, with 5 more or less conspicuous (when dry depressed) costae,
shortly and densely tomentose-velvety, yellow when ripe; pericarp
about 5-6 mm. thick, the inner layer coriaceous, smooth, very hard,
about 0.5-1 mm. thick, the middle tissue about 3-4 mm., carnose, the
outer layer coriaceous but less hard than the innermost and becoming
rugose after the shrinking of the intermediate layer by drying; seeds
about 20-26, surrounded by whitish, sweet, scentless pulp, ovoid-
oblong or ellipsoid-oblong, 24-26 mm. long, 13-14 mm. broad, 10-
12 mm. thick, the episperm thick (about 1 mm.), coriaceous with the
middle layer becoming gelatinous; embryo white, oblong, covered by
a very thick pellicle, 22-24 mm. long, 10-11.5 mm. broad, 9-10 mm.
thick; germination epigeous.

The leaves of the type of 7. speciosum are long-petiolate, cuneate
and trinerved at the base. Theobroma quinquenervium Bernoulli was
described from a specimen with short-petiolate, broadly oblong
leaves, asymmetrically rounded at the base. In the first type the
margins of the leaves are close and parallel to the prominent, basal
pair of secondary nerves; in the second the margins of the leaves are
broadened and remote from the prominent basal pair of nerves, and
an additional lower tertiary nerve on each side makes the base of the
blade somewhat 5-nerved. The latter type of blade, borne on a short,
stout petiole, is the common one in the species. Leaves with slender,
long petioles, thickened at both ends, and cuneate blades are seldom
found; they appear on young, orthotropic terminal branches. This
dimorphism was already noticed by Huber who first united 7. quin-
quenerrium and T’. speciosum.

The type specimen was collected near Belém de Para by Siber who
was sent on a collecting trip to Brazil by Hoffmannsegg; it is pre-
served in the Berlin-Dahlem Botanical Museum in the Willdenow
Herbarium. I have been able to study this specimen thanks to the
kindness of Prof. Werdermann and Prof. Melchior. The photo-
graph F.M. 9640 is from a specimen at B now destroyed, which agrees
perfectly with the type in the Willdenow Herbarium, of which it was
undoubtedly a duplicate. In his monograph, Chevalier made this
species a synonym of 7’. guianense (Aubl.) Gmel., but this is a con-
fused species, the identity of which is discussed in this paper.

Spruce and Ducke called attention to beauty of this tree in blossom,
according to Spruce “one of the prettiest things I have seen.”

CUATRECASAS—CACAO AND ITS ALLIES 481

ID

y

7

3
2
:
Fe
a iN"
; I.
a

tats
eS

Wy

na

vane

SAR,
Ete

iN,

BAK TS
=e

C

Ficure 19.—a, B, Detail of nervation at the underside of the leaf in: a, Theobroma speciosum
(Ule 9609); B, 7. cacao (Cuatr. 7756). c, p, indument on the underside of the leaves

in: c, T. glaucum (Bart. & Holl. 74); p, 7. bernouillii subsp. capilliferum (Cuatr. 16160).
A and B X 2, Cand D X 25.

482 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Theobroma speciosum can grow to be 15 m. high, with a few whorls
of dichotomous leafy branches near the top of a long branchless stem,
which can bear abundant, large, cauline inflorescences. These may
form large cushions of showy, blood-red, wine-red, or purplish-red
flowers which give off an intense lemon- or orange-skin odor (Ducke).
The outer pulp of the seeds is sweet but scentless. It is said to be easy
to grow in gardens.

CoMMON NAMES.—Cacauf, cacauu. Other recorded names or
different ways of spelling are: cacao-y, cacao-u, cacau, cacaohy,
cacau-i, cacao-i, cacaofllo, cacau-rana, cacao-rana, cacao biaro,
cupuy, cupuyh, cacao do matta, cupurana, cacao azedo, cacao sacha
(Peru), chocolatillo (Bolivia).

Usrs.—The pulp is eaten by natives. The seeds are used very
occasionally to prepare low quality chocolate.

DisrrisutTion.—This species extends throughout the Amazonian
Hylaea except in the northwestern section, from the state of Maranhao
at Cururupu to Acre and Madre de Dios in eastern Bolivia and the
Ucayali River (Loreto) in Peru. It grows on noninundatable ground
in rain forests as well as in not too humid places and it appears also in
secondary growth near towns, but it is never of frequent occurrence.

BRAZIL: “Bresil. Herb-Lusit,’ “Herrania paraensis,”’ “Theobroma subin-
canum Mart.?,” Geoffroy [St. Hilaire?] s.n. (P). ‘“‘Polyadelphia Decandria
Theobroma speciosa foliis acuminatis integerrimis subtus tomentosis. Flores
purpurei. Habitat in Para Brasilia,” ‘Theobroma speciosa (W.) Spreng. Bernoulli
determ.” Hoffmannsegg, W. (B, Herb. Willdenow 3680, holotypus). ‘Ex herb.
Linn. Theobroma speciosa Brasil Col.’ Siber (B), specimen destroyed (photo-
graph F. M. no. 9640); identical with the type in the Willdenow Herbarium.

ParA: Belém, noninundatable forest near city; small tree with dark-red
flowers on trunk, “‘cacaohy,” IX 1936, Ducke 281 (A, F, K, MO, NY, 8S, US).
Belém, Jardim Boténico do Museo Goeldi; slender tree, inflorescence on trunk,
petals dark red, sepals reddish stained, fruit 5-parted, yellow when ripe, “cacau-i,”’
26 VIII 1942, Archer 7619 (IAN, K, NY, USDA). Ibidem, slender tree,
“cacao-i,” 29 X 1942, Archer 7721 (IAN, USDA). Ibidem, small tree, flowers
deep crimson, calyx light pink, leaves deep green above, gray green beneath,
cultivated, 15 VII 1946, Schultes & Silva 8066 (GH). Ibidem, 19 X 1945, Pires
& Black 695 (IAN), 740 (IAN). Braganga, ‘‘cacao claro,” XII 1899, Huber
1748 (G, MG). Taperinha, at Santarém, secondary forest on elevated land;
small tree, red flowers on trunk, “cupuy,’”’ Ginzberger 802 (WU). Rio Curuauna,
Cachoeira do Portao, region of Planalto de Santarém, rain forest, XI 1954, Frées
31414 (IAN). Santarém, hills, “cacao do Matto,” Jobert 903 (P). Near Obidos,
Prov. Pardé, XII 1849, Spruce s.n. (K, Herbarium Hookerianum 1867). Forest
near Obidos, XII 1845; slender tree 40-50 ft., flowers on the naked stem, leaves
on the top of the tree only, ‘‘cacao-rana,”’ Spruce 456 (K, Herbarium Benthami-
anum, 1854, P, BM). Monte Alegre, region of Colonia da Mulata, elevated
ground; tree 8m., red flowers, 28 IX 1953, Frées 30432 (IAN). Breves, VII
1956, Pires, Frées, & Silva 5886 (IAN). Belterra, capoeira, way to Pindobal;
tree 7 m., red flowers on trunk with lemon scent, leaves white beneath, 31 XI 1947,
Black 47-1889 (IAN, NY). Region Tapajés, Boa Vista firm land; tree 4 m.,

CUATRECASAS—CACAO AND ITS ALLIES 483

10 cm. [diam.], flower color wine, fruits on the trunk, eaten by Indians, ‘‘cupuhy,”’
30 VIII 1932, Capucho 397 (IAN, F). Tapajés, Vila Braga; tree, dark-red
flowers borne on the trunk, 27 X 1908, Snethlage 10044/b (MG). Upper Cupary
River, plateau between the Xingi and Tapajdés Rivers; tree 25 ft. high, 2 inches
in diam. breast high, in high forest, flowers red on old wood, ‘“cacaorana,” IX
1931, Krukoff 1117 (A, G, K, NY, P). Ibidem; tree near river shore; flowers
dark red, borne on large branchlets; wood used for ‘‘farinha”’ containers (to keep
it dry), “cacaorana,” Krukoff 1080 (A, BM, G, K, MICH, MO, NY, 5S, U). Rio
Cumina-mirim, forests at NE; ripe fruit yellow, only on trunk, “cacaorana,”’
16 XII 1906, Ducke 7975 (MG). Oriximina, Las Trombetas, ‘cacao rana,”
flowers only on trunk, 8 XII 1906, Ducke 7884 (G, MG).

Amazonas: Near Barra, Prov. Rio Negro, Aug. 1851; forests south of Rio Negro;
tree 20 feet, straight with a whorl of branches only at summit, which are twice
or thrice dichot{omous]; trunk almost completely clad with flowers, which have
fine odor of bruised orange leaves; petals cucullate, claw pink, white limb dull
crimson with dark veins. The subul[ate] processes blood red; one of the prettiest
things I have seen, Spruce 1737 type of Theobroma quinquenervium Bernoulli
(M, holotype, Photo F. M. 40708; isotypes: BM, E, F, G, GH, GOET, K, LD,
LE, OXF, P, WU) (Photo F. M. 9639 from Berlin). Rio Marmellos, near
mouth; flower deep salmon red, buds dark crimson, bark smooth, light gray
white, leaves pale dull beneath, dark green glossy above, branches umbelli-
form at top, trunk 6-7 inches in diameter at base, 20-22 feet tall, inflorescences
200-flowered; seeds give low grade chocolate, 2-12 VIII 1945, ‘‘cacao azedo,”’
Schultes & Cordeiro 6507 (AMES, F, IAN). Amazonas, Rio Capitare, munici-
pality of Codajas, elevated ground, high forest; tree 8 m., red flowers, 3 IX 1950,
Frées 26526 (IAN). Manaos, Estrada do Aleixo; tree 5 m., fruits on trunk,
14 V 1953, Frées 30180 (IAN).

Guapore: Porto Velho, Estrada de Rodagem, Km 8, Viana, disturbed
forest, elevated land; small tree, red flowers with lemon scent, 31 V 1952, Black,
Cordeiro, & Francisco 52-14655 (IAN).

Goras: Margen do Rio Araguaia; tree 4 m., red flowers, 13 VI 1953, Frées
29732.

AcrE (TERRITORIO DEL): Rio Acre, Seringal San Francisco; tree 5-20 m.,
black-purplish flowers on stem, VII 1911, Ule 9609 (G, K, L), Ule 14448 (MG).
Near mouth of Rio Macauhan (tributary of Rio Yaco), Lat. 9°20’ 8., Long. 69° W.,
on firm land; tree 40 feet high, 5 VIII 1933, Krukoff 5295 (A, F, K, LE, M, MICH,
MO, NY, 8, U, UC, US).

Rio pE JANEIRO: Quinta de Sao Christovas, 10 I 1876, Glaziou 9633 (C, P).
Jardim Botdnico, flowers on trunk and main branches, V 1944, Camargo 2395
(IAN).

PERU: Loreto: Region of the middle Ucayali, rain forest, Yarina Cocha,
155 m. alt., elevated ground; tree 12-15 m., 35 cm. thick, the first branch at
7m. from ground, flowers with strong anise scent, sepals red crimson, petals dark
crimson-striped on white ground, stamens less crimson than petals, pistil whitish,
“cacao sacha,” 22 IX 1925, Tessmann 5398 (G, M, 8). Rio Ucayali, Paca, 21
VII 1898, Huber 1567 (Holotype of var. coriaceum Huber, MG; isotypes BM, US),
(photo F, M. No. 1567). 40km. south of Pucallpa, rain forest of the Amazon basin
virgin rain forest on loamy sand, Ellenberg 2565 (U).

BOLIVIA: Km. 7 on road Guayaramerin-Cochuela Esperanza, [Prov. Vaca
Diez, Depto. Beni], 120 m. alt., ‘‘chocolatillo”; growing wild in rain forest; flowers
wine red, borne on cushions along the whole, rather slender trunk, dichotomous
branches slender, pendent, fruits small. I have seen also a specimen in Brasilia,
where it is called “‘cacau-i’’; 9 IX 1954, Patifio s.n. (GH). Junction of Rivero

484 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Beni and Madre de Dios; pulp edible and equal to that of 7. cacao, seeds white,
not used, VIII 1886, Rusby 654 (BM, E, F, G, GH, K, LE, MICH, MO, NY, P,
US). Unduavi, 10,000 ft., Rusby 647 (US).
4. Theobroma velutinum Benoist, Bull. Mus. Hist. Nat. Paris 27: 113. 1921.
Ficures 3, 12, 13, 14, 16, 18; Map 1.
Herrania guianensis Sagot ex Schum. in Mart. Fl. Bras. 12(3):75. 1886.
Theobroma speciosum sensu Uittien in Pulle, Fl. Surinam 3:45. 1932.
Theobroma sp. 4, Uittien in Pulle, Fl. Surinam 3:46. 1932.
Herrania guyanensis Sagot in Chevalier, Rev. Int. Bot. Appl. 26:275. 1946
(as synonym).
Typr.—Benoist 516, French Guiana.
Branches terete, subcinereous, densely appressed stellate-tomentose
or becoming glabrate, grayish brownish; branchlets densely tomentose.
Leaves large, firmly coriaceous; petiole robust, subterete, densely
appressed cinereous-tomentose, when old transversely rimose, 12-14
mim. long; lamina oblong-ovate or ovate-oblong, broadly rounded and
more or less asymmetrical at base, suddenly narrowed and long-
acuminate at apex, the margin slightly revolute, entire or very slightly
sinuose, 28-40 cm. long, 16-21 em. broad, including the acumen, this
about 3 cm. long, pale olivaceous above (when dry), subnitidous,
slightly and broadly bullate, glabrous except for the midrib, this
sparsely pilose towards the base, the costa and secondary nerves thin,
the tertiary nerves filiform, the minor veins less apparent or obsolete,
softly velutinous beneath, ochraceous-cinereous, the midrib very
prominent, the 5-7 pairs of secondary nerves prominent, ascending,
curved near the margin and anastomosing, the longer pair stronger,
more separated from the next, the transverse tertiary nerves promi-
nent, others broadly reticulate, prominulous, the lesser veins forming a
minute, prominulous reticulum, the minor reticulum and its areoles
densely tomentose with minute, white, dense, subappressed, stellate
hairs, the other nerves and the sides of the midrib covered with abun-
dant, long, delicate, stellate hairs with long, thin, spreading rays.
Inflorescences many-flowered, borne on lignose, tuberculate branches
on the trunk, the panicles fasciculate, ramose from the bases, 5-12 cm.
long, the branchlets moderately thin, cinereous, hirtellous-tomentel-
lous with thin, mediocre, stellate hairs, the terminal (peduncles)
thinner with 2 or 3 deciduous bracteoles at the end; pedicels slender,
6-18 mm. long; bracteoles at the joints minutely lanceolate, hirtellous,
deciduous, 1-1.5 mm. long; sepals thick-membranaceous, spreading,
shortly united at base, lanceolate-oblong, subacute, glandular at base,
otherwise glabrous inside, with fine stellate hairs copious on the
outside, the margin minutely tomentulose, about 10 mm. long and
3.5-4.5 mm. broad, usually one free, the others united by pairs.
Petal-hoods 6-7.2 mm. long, about 3 mm. broad, membranaceous,
obovate-oblong, cucullate-rounded at apex, 3-nervate, the median

CUATRECASAS—CACAO AND ITS ALLIES 485

vein furcate, outside sparsely, weakly pilose, the end auriculate-
emarginate, articulate to the erect lamina, 6-7.2 mm. long, about
3 mm. wide; petal-lamina sessile, subtrapezoid, subtruncate or often
slightly sinuate or 3-toothed at apex, abruptly narrowed into a short
claw at base, moderately thick, but venation conspicuous by trans-
mitted light, glabrous, somewhat rugulose-papillose toward the base,
5 mm. long, 6-6.5 mm. broad; androecium-tube about 2 mm. long and
2.4 mm. broad, minutely papillose-pilose; staminodes purplish red,
lanceolate-subulate, thick, suddenly narrowed into a short, crisp,
acute acumen at apex, minutely muricate-pilose, about 6 mm. long
and 1.5 mm. wide; filaments triantheriferous, 2.5 mm. long; ovary
ovoid-oblong, about 2.2 mm. long, 5-ridged, tomentose; styles 5,
connivent, adherent, easily separable, glabrous, 2 mm. long.

Fruits ellipsoid, densely and softly stellate-pilose-velutinous, 5-
costate, the ribs thick, very prominent, the surface smooth, 8.2—-9 cm.
long, 6-6.3 cm. broad, the pericarp 3-4 mm. thick; seeds usually 25-30
in each pod.

This species is closely related to T. speciosum, having very similar
flowers and leaf outline. However, 7. velutinum is very different on
account of the structure of the fruits and theindument. The ellipsoid,
densely velvety fruits have 5 longitudinal, very prominent, typical
ribs, a character only known in this species. The leaves beneath have
a soft, velvety indument of long, thin, stellate or dendroid, more or less
densely distributed hairs on the whole nervation and a lower layer of a
short, dense tomentum of minute, stellate, intricate hairs covering
the areoles and the minor reticulate veins. Also, the terminal
branches are densely tomentose and the inflorescence branchlets and
pedicels have copious, rather long, fine hairs.

Theobroma velutinum is only known from French Guiana and the
neighboring Dutch side of Maroni River valley. The excellent foliage
and fruiting collections made by Benoist and recent other collections
by members of French and Dutch Forest Services, especially the
flowering specimens B. B.S. 1161, have facilitated a complete descrip-
tion of the species. Its inflorescences are cauline, many-flowered,
and often showy, like those of 7. speciosum. The Sagot collection
1206, preserved in several herbaria and consisting only of large
inflorescences, belongs to this species. Sagot named it Herrania
guianensis and left an accurate unpublished description which 1s
attached to the specimens in Paris.

CoMMON NAMES.—Bouchi-cacao, cacao sauvage, cacao.

Usrs._Reported to yield edible seeds comparable to cacao but no
information is given on the quality of the product.

DistripuTion.-—French and Dutch Guiana in the valley of Maroni
River.

486 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

SURINAM: 4 X 1950, B. W. 1161 (U). Placer l’Arver, 31 X 1918, Gonggryp
4108 (U). Flur of Marowyne, Reu naar L. etwa en Tupanahoni, No. 47 Maro-
wyne, 26 XI 1918, Gonggryp 4127 (U). Flur Tapanahoni, III 1922, No. 33
Tapanahoni, Gonggryp 4148 (U).

FRENCH GUIANA: A 1 kilometre plus ou moins du camp de transportés de
Charvein, le long de chemin qui conduit a la leproserie de Acarouany, 8 I 1914,
Benoist 516 (P, holotype, foliage and fruits). Crique Serpent, rive droit 4 1
kilometre de la crique terrain en pente, & proximité immediate d’un ravin rocheuse;
arbuste ayant été repéré au moment de sa fructification en février 1951; abattu
depuis et pourvu de rejets de 3 m. de haut; produit de gousses edtées contenant
des grains comestibles, “‘bouchi cacao’’ (Paramaka), 12 XII 1952, BAFOG 136M
(P). Placean No. 2, Carreau No. 3, route de mana, terrain sablonneaux; fruits
jaundtres, gousses ovoides de 7 4 8 x 10 A 12 cms. déhiscents, sparses sur le trone
de l’arbre, “‘bouchi cacao,” “cacao sauvage,” 19 III 1956, BAFOG 7386 (P).
Karouany, 1858, ‘Flores atropurpurei suaveolentes, fructus ovatus, pentagonus,
breviter tomentosus cacao sativa paulo minor, folia non vidi, flores e ligno
prodeunter, ‘‘cacao,” “Herrania guianensis Sagot,”’ 1858, Sagot 1206 (BM, G,
K, P, U, WU); type of Herrania guianensis Sagot (only inflorescences and flowers
in all the specimens; a long and accurate description made ‘in vivo’ by Sagot
is attached to the specimens at Paris).

5. Theobroma glaucum Karst.
Ficures 2, 10, 14, 16, 17, 19; Map 5; Puarn 4
Theobroma glaucum Karst., Linnaea 28:447. 1856; Triana & Planch. (1862);
Bernoulli (1869) 10; Jumelle (1899) 34; De Wildeman (1902) 97; Chevalier
(1946) 277.
Theobroma calodesmis Diels, Notizbl. Bot. Gart. Berlin 14:336. 1939; Baker,
Cope & al. (1954), figs. 10, 12; Cuatrecasas (1956) 655; Leén (1960) 314,

317, fig.
Types: Karsten s.n. Colombia, San Martin. Schultze-Rhonhof
2312, Ecuador, Papayacu (of 7. calodesmis, formerly in Berlin).
Tree 8-15 m. high; stem up to 30 em. in diameter, with grayish,
inside reddish bark and white wood; sympodial growth by lateral,
subterminal, upright shoots; primary branches ternate, regularly
dichotomous, spreading, deciduous when old, the terminal minutely
stellate-pulverulent with additional simple, spreading hairs, soon
glabrate, smooth, rather shining, dark brown or somewhat purplish;
stipules linear-subulate, 4-5 mm. long, sparsely stellate-pilose, soon
deciduous.

Leaves coriaceous, rather rigid, distichous; petiole robust, densely
ferruginous or brownish tomentose with stellate hairs, transversely
rimose when dry, 0.8-1.8 cm. long; blades oblong-ovate or ovate-
oblong, broad in the lower third, obtusely cuneate at base, narrowed
near the apex, prolonged with a long slender appendage, the margin
entire or slightly sinuate and slightly revolute, 16-36 em. long, 7-13
cm. broad, the acumen 2-3.5 mm. long, shining above, green, pale
olivaceous brown when dry, apparently glabrous but with sparse
mediocre stellate hairs and callose scar-dots on the nerves, the costa
and secondary nerves prominently filiform, the others slender, more or

CUATRECASAS—CACAO AND ITS ALLIES 487

less noticeable, somewhat cinereous beneath, glaucous or pale rosy,
with a glabrous aspect but the rather shining, pale brownish principal
nerves sparsely callose-dotted and with very scarce ferruginous,
stellate hairs, the small veins glabrous, the areoles covered with a
very appressed microscopic tomentum of minute, white, stellate hairs,
the costa very prominent, the 4 or 5 pairs of secondary nerves promi-
nent, the basal one at an acute angle (remote from the margin), ascend-
ing, the others curved-ascending, near the margin becoming slender,
decurrent, curving, anastomosing, the cross-tertiary nerves thinner,
prominent, 3-10 mm. distant from each other, the lesser veins
minutely prominulous-reticulate; leaves of the orthotropic spreading-
puberulous branches long-petiolate, with the blades attenuate-
cuneate at base, the lower pair of nerves very close to the margin,
the petiole slender, 3.5—4 cm. long.

Inflorescences on the trunk, often many-flowered and showy, with
up to 200 flowers, the base woody-tuberculate; branches 3-many, me-
diocre, 4-6 cm. long, furcate-ramose from near the base, the branchlets
fastigiate, angulate, rather rigid, ferruginous-tomentose, the terminal
(peduncles) moderately robust, 3-4 mm. long, articulate to pedicel,
this 5-15 mm. long, striolate, slightly thicker, tomentellous, the
subtending bracteoles minute, ovate-lanceolate, about 1 mm. long,
very soon deciduous; buds ovoid, round at base, subacute at top,
densely stellate-tomentose, 8-9 mm. long, about 6 mm. broad.

Calyx umbilicate; sepals thick, lanceolate-oblong, acute, inflexed
at apex, connate for 2 mm. at base, densely and appressed stellate-
tomentose outside, within minutely, whitish stellate-pilose near the
margin and glandular at base, otherwise subglabrous, 12-13 mm. long,
3.5-4 mm. broad, curved-spreading after anthesis.

Petal-hoods light red, oblong-obovate, shortly unguiculate at base,
rounded cucullate at apex, the end emarginate, biauriculate, articulate
to the lamina, the 3-nerves prominent inside, thin, with spreading,
weak, sparse hairs outside, 5-6 mm. long, 2.5-3 mm. broad; petal-
lamina red crimson, thick, minutely rugose, and more or less trans-
lucid-venulose, glabrous, suborbicular or broadly elliptic, subsessile
at base, abruptly contracted into a short claw, minutely sinuate at
margin, 5.5-7 mm. long, 5-6.5 mm. broad, the claw 0.5 mm. long.

Staminal tube about 2 mm. high and 2.5 mm. in diameter; stami-
nodes red crimson, erect, lanceolate-subulate, acute at apex, fleshy,
minutely muricate-pilose, 10-12 mm. long, 1.4-1.8 mm. wide above
base; filaments flexuose, 2.5-3 mm. long, glabrous, shortly 2 or 3
furcate at apex, bearing 2 or 3 anthers, the loculi ellipsoid, convergent,
0.5-0.6 mm. long; ovary oblong, about 2 mm. long, 5-ridged and sul-
cate, hirsute-tomentose; styles connivent, 3 mm. long, united only
at base.

680-695—64——-8

ASS CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Fruits ellipsoid-oblong, obtusely pentagonal, broad and umbilicate
at base, more or less attenuate and subacute at apex, the pericarp
1 cm. thick, coriaceous, rigid, densely and minutely velutinous-
tomentose, bluish greenish, 11-13 em. long, 5.5-9 cm. broad; seeds
2-2.3 x 1.2-1.4 x 0.9 cm.; fruiting peduncle robust, 7-8 em. long, 1-1.5
cm. thick; germination epigeous.

According to Baker and his associates the cotyledons are white and
the pulp is pale orange and of a very sweet taste.

The type collection of T. calodesmis was destroyed during the war
in Berlin; it was collected by Hertha Schultze-Rhonhof near Papayacu
at about 200 m. altitude on the Bonanza River, a tributary of the
Pastaza River in eastern Ecuador. The description given by Diels
makes it possible to identify his species perfectly with several Amazo-
nian collections from nearby regions of Colombia, Peru, and Brazil.
The sterile type specimen of Theobroma glaucum, collected on the
Llanos de San Martin, agrees perfectly with a specimen that is almost
a topotype, collected by Philipson, Idrobo, and Fern4ndez in the
foothills of the Sierra Macarena, Intendencia del Meta. I have no
doubt that all these collections represent the same species, which
extends from the upper Orinocia to upper Amazon basin on both sides
of the great river.

Diels did not see fruits but gave an accurate description of the
foliage and flowers; he related his species to 7. speciosum on account
of the texture and tomentum of the leaves, and also to 7. bernouillii,
but he says that the leaves are broader, the inflorescences larger, the
flowers larger, and the staminodes longer.

Common NAMES.—Cacao de monte, cacao silvestre, chucti’ (Rio
Papayacu [Schultze-Ronhof]), ‘“bicco” Rio Apaporis).

Usrs.—According to Karsten, the seeds are used as cacao by the
natives, being very similar to the true cacao. Schultze-Rhonhof gives
the indigenous name ‘“chucti”’ for the fruit which, according to her,
is very much appreciated by the natives.

DistripuTion.—This species grows in the upper Amazon basin
(northwestern section) along the rivers Caqueta, Caguan, Putumayo,
Vaupés, Guainfa, Infrida, and Apaporis in Colombia and along the
Colombian boundary with Ecuador, Peru, and in western Brazilian
Amazonas; at the northern end of its range it enters the Orinoco basin
into the Meta drainage in Colombia.

It grows in the humid rain forests from the lower level of the great
rivers to an altitude of 450 m. in forested hills.

COLOMBIA: Mera: Villavicencio, Llanos de San Martin, Karsten, s.n. (WU,
holotype) (photo F. M. 32205). Sierra Macarena, Cano Yerly, 450 m., dense,

humid forest; unbranched tree 10 m. high, bunches of cauliflorous fruits green,
24 XI 1949, Philipson, Idrobo, & Fernandez 1552 (BM, COL, US).

CUATRECASAS—CACAO AND ITS ALLIES 489

Putumayo: Vicinity of Mocoa; tree 6-7 m., growing fully exposed in meadow,
trunk 23 em. thick at base, branched inflorescence (dead) borne on trunk and
branches, pod broadly pentagonal with 5 very shallow furrows, 13 cm. long,
9 cm. diam., blue green in colour, fruit pedicel 8 cm. long, 1.5 cm. diam., with
abscission ring 1.5 cm. from pod base, in colour pale green with fine white hairs,
17 III 1953, Holliday & Cope T/79 (COL, TRIN, US). Ibidem; sterile tree 15 m.
in forest, undoubtedly similar to T/79, shoots from below jorquette, Holliday
& Cope T/79A (COL, TRIN, US). Rio Leguizamo, Laguna Primavera, 3 1V
1953; tall tree 15 m., obviously cauliflorous, Holliday & Cope T/94 (COL, TRIN,
US). Rio Leguizamo; tree 16 m., with old trunk inflorescences, no flowers or
fruits, 5 1V 1953, Holliday & Cope T/96 (COL, TRIN, US).

Caqueth: Rio Cagudn, Camp 4; branched tree 10 m., found in good flowering
condition, flowers in large inflorescences on trunk, 27 IV 1953, Holliday & Cope
T/118 (COL, TRIN, U, US). Ibidem; tree 15 m., branched, two immature pods,
on sloping land near T/114, 26 IV 1953, Holliday & Cope T/115 (TRIN, US).

Vaupsés: Rio Guainfa, near Victorino, river level; tree without flowers or
fruit but with stipules; said by the local Indians to be a type of cacao with small
smooth pods, native in the forest, 3 II 1952, R. EH. D. Baker 37 (TRIN, US).
Rio Infrida, Santa Rosa, 300 m.; tree 15 m., trunk about 30 cm. in diameter,
bark greenish, cortex light red, wood white, no terminal growing point, young
shoots arising from below jorquette, cauliflorous, pod surface 10-ridged, also
with transverse ridges, fruit pedicel about 4 cm. long with abscission layer 3 cm.
from trunk, 25 I 1953, Bartley & Holliday T/69 (COL, TRIN, US). Left bank
of Rfo Infrida, San Joaquin, 200 m. from river bank, 300 m. alt.; tree 10 m.,
trunk base 15-20 cm. in diameter, one dead cymose inflorescence seen, 28 I 1953,
Bartley & Holliday T/70 (COL, TRIN, US). Rio Inirida, Rio Papunana; tree
10 m., trunk about 30 cm. in diam., bluish gray in appearance, bark red, wood
white, three branches at each jorquette, dichotomous branch habit, inflorescences
on upper part of trunk, 18 II 1953, Bartley & Holliday T/74 (COL, TRIN, U, US).

Amazonas: Rio Apaporis, Jinogojé, river level; tree 40-50 ft., 8’’-9’’
diameter at base, jorquettes of 3 branches, subsequent growth from below, ultimate
branches repeatedly bifurcating, flowers in large clusters, sepals, ligules, and
staminodes dark crimson, 8 IX 1952, Baker & Cope 11 (COL, F, TRIN, US).
Ibidem (boundary Amazonas-Vaupés) between Rfo Piraparand and Rio Popeyaca,
250 m., Cafio Unguy4; tree 8 m., calyx red, petals white, purplish red at apex,
“bicco,’’? 3-11 XI 1952, Garcia-Barriga 14380 (COL, US).

BRAZIL: Amazonas: Sao Paulo de Olivenga, on elevated land; tree 10 m.,
12 cm. [in diam.], ‘‘cacau azul,” 18 V 1945, 'rées 20942 (NY). Cidade Fonte Béa
(“introduzida pelos indios do Japurda’’); tree 10-12 m., 12-15 cm. diam., “‘cacau
azul,” “cachu azul,” 41V 1945, F’rées 20645 (K, USDA). B. constant, tree 10 m.,
“cacau azul,” 9 V 1945, Frées 20885 (NY). Macacacain, Rio Jutahi, Barreira
Branca; tree 15-25 feet, small, main branches in whorls of 3, each bifid, 31 I 1875,
Traill 62 (GH, K, P).

6. Theobroma bernouillii Pittier
Ficures 2, 6, 7, 15, 16, 17, 19; Map 4; PuaTE 5
Theobroma bernouillii Pittier, Repert. Sp. Nov. Fedde 13:319. 1914.
Chevalier (1946) 26: 277; Leén (1960) 314.
Theobroma asclepiadiflorum Schery, Ann. Mo. Bot. Gard. 29:360. 1942.
Theobroma capilliferum Cuatr. Rev. Acad. Colomb. Cienc. 6:547, figs. 3a,
4a, pl. 34. 1946.

490 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Typrs.—Pittier 4105, Panama. Wedel 1535, Panama (of T’. asclepia-
diflorum). Cuatrecasas 16160, Colombia, Pacific coast (of TI.
capilliferum).

Tree 15-20 m. high; stem up to 25 cm. in diameter, with grayish-
brown, rugulose bark 5 mm. thick and yellowish-white wood; sympo-
dial growth by lateral, subterminal, upright shoots; primary branches
ternate, grayish brown or blackish brown, dichotomous, the oldest
falling off, leaving the stem naked, the terminal stem leafy, terete,
dark brown, minutely, subappressed tomentulose, when older puber-
ulous or glabrate; stipules linear-oblong, attenuate at apex, acute,
pubescent, about 8 mm. long, 2.5 mm. wide.

Leaves coriaceous, more or less rigid, entire; petiole thick, 4-12 mm.
long, terete, densely and minutely tomentulose, transverse rimose
when dry; blades rather asymmetrically ovate, ovate-oblong, or
elliptic-oblong, obtusely cuneate-attenuate and mostly very asym-
metrical at base, triplinerved, narrowed toward the apex, ending
in a long, linear, acute tip, the margin entire or very slightly sinuate,
slightly revolute, 13-30 cm. long, 5.5-18 em. broad, including the
acumen, this 1.5-6 cm. long, 2-4 mm. broad, green or pale brownish
above when dry, sbining, the main nerves filiform, prominent, the
others reticulate, slightly prominulous, pale greenish ochraceous or
ashy beneath, apparently glabrous but the tawny shining veins
with very sparsely minute stellate hairs and the areolae covered
with very appressed, white tomentum of smaller, microscopic,
entangled stellate hairs, the midrib and 5-7 secondary nerves on each
side very prominent, the basal pair acutely ascending, the others
distally curved-ascending, decurrent and anastomosing near the
margin, the transverse tertiary nerves thin but prominent, the minor
veins prominulous, minutely reticulate.

Inflorescences on the main trunk usually many-flowered, borne
on short, tuberculate, woody branches, the panicles abundant,
4-10 cm. long, spreading, bristly, the axes slender (0.4-1.5 mm.
thick), rigid, striolate, stellate-tomentose in the upper third or rarely
from the base, cymose-bifurcate, corymbiform, the branchlets rigid,
erect at acute angle; peduncles (ultimate branchlets) capillary,
tomentulose, 2-20 mm. long, 3-bracteolate at apex; bracteoles narrowly
linear, 1-2 mm. long, very soon deciduous; pedicels thicker than the
peduncles, about 5-20 mm. long at anthesis, minutely stellate-
tomentellous like the branchlets; buds ovoid, densely and shortly
tomentose.

Flowers crimson; sepals moderately thick, lanceolate-oblong, rather
acute, shortly united at base, often first temporarily united in pairs
but separated later, 8-10 mm. long, 3 mm. wide, reddish, with sparse
slender, flexuous, sericeous hairs inside, ferruginous, rugulose, and

CUATRECASAS—CACAO AND ITS ALLIES 49]

stellate-tomentose outside, minutely tomentulose at the margin
inside, with thick, glandular hairs crowded at the insertion at base;
petal-hood red, oblong-obovate, rounded-cucullate at apex, narrowed
at base, 3-nerved, prominent inside, hirtellous pubescent outside,
4-5 mm. long, about 2 mm. wide, at base 0.6 mm. wide; petal-lamina
crimson, sessile, shortly unguiculate-articulate, moderately thick,
rugulose, glabrous, orbicular or suboricular or elliptic, minutely
crenulate, 2.5-4 mm. long, 3-4 mm. broad; staminal tube about
1.5-2 mm. high; staminodes 6-9 mm. long, erect in bud, purplish red,
sublanceolate-linear-subulate, thick, suddenly narrowed toward the
apex, covered with minute, spreading, acute, conical trichomes;
fertile filaments glabrous, flexuous, about 2.5 mm. long, 2-anther-
iferous; ovary 5-sulcate-costate, tomentose, 1.2 mm. high; styles
five, 2 mm. long, adherent into a column but separable.

Fruit 15-20 (12-25) em. long, 6-7.5 (5.2-8) cm. broad, ellipsoid-
oblong, more or less prismatic, obtusely 5-angulate, abruptly narrowed
subacute or subobtuse at apex, umbilicate at base, more or less con-
stricted near the base or not; pericarp thick, rigid, coriaceous at
maturity, the epicarp and endocarp hard coriaceous, the mesocarp
fleshy, shrinking in drying, dull brown, dense velvety-tomentose ;
seeds compressed-ovoid, 16-22 x 9-14 x 9-11 mm., the testa reddish,
papery, the cotyledons white; pulp white, flavored, acidulous; germi-
nation epigeous.

This species, as here broadly considered, includes heteromorphic
elements described as three species, two of which came from the
Atlantic coast of Panam&, and the third from the Pacific coastal
region of Colombia. The Colombian population (7. capilliferum) is
the best known, being represented by several collections with fruits
and flowers, showing morphological uniformity throughout its area.
The original 7. bernouillit is known only from flowering material of
one collection (the type), which has some minor differences from the
Colombian plants in the shape of the leaves and details of flowers.
The other described Panamanian species, 7. asclepiadiflorum, was
based on discordant elements collected by von Wedel in Water Val-
ley. Schery wrote: “Although fruiting material of this species is
lacking, floral and vegetative characters distinguish it sufficiently to
warrant description as a new spccies.”?’ The glabrous branches and
leaves of the type specimen, which resemble those of 7. cacao, do
not belong to a Theobroma; they belong actually to the Lauraceae.
The inflorescences and flowers are similar to those of T. bernouzllii,
type specimens of which were collected by Pittier not very far away.
The question about what kind of leaves belong to the described
flowers of JT. asclepiadiflorum is answered by the collection Wedel
681 from the same locality, Water Valley; the flowers of this collec-

492 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

tion are identical to those of the type of 7. asclepiadiflorum and the
leaves are exactly like those of 7. bernowillii. There are minor differ-
ences between the flowers of these two types, but considering the
vicinity of the geographic range of these two populations, it may be
safe to consider them mere forms of one species. Fruits from the
type locality of 7’. bernowillia have never been collected, but Allan
Lucas did collect fruits of 7. asclepiadiflorum in Water Valley. The
flowers of T. asclepiadiflorum are almost identical to those of T.
capillferum, but the fruits are smoother and not constricted.

In view of these facts, I consider all these Colombian and Panama-
nian plants to belong to one species; since the three forms are geo-
graphically separated the observed differences warrant subspecific
recognition.

DistTRIBUTION: Pacific coast and the Chocé region of Colombia and
the Atlantic coast of Panama. In the underforest of the tropical
rain forest, from sea level and lowland swamps to heavily forested
hills about 100 m. altitude.

Key to Subspecies of Theobroma bernouillii

1. Leaves broadly ovate or ovate-oblong, very asymmetrical, rigid, 5-12 x 2-6
em. Petal-lamina orbicular 2.5-3.5 mm. long; petal-hood 4-5 x 2 mm.;
staminodes subulate, densely pilosulous, 7-8.5 mm. long; inflorescences
long, with thin branches, the peduncles 5-20 mm. long, the pedicels about
10 (5-20) mm. long; fruits obtusely pentagonal with smooth, conspicuous
ridges, 12-25 x 5-8 em., umbilicate constricted above the base.

6c. subsp. capilliferum

1. Leaves oblong-elliptic, slightly asymmetrical, thinner, less rigid.

2. Staminodes lanceolate, slightly pilose, 5.5-6 mm. long, 1.5-2 mm. broad at
base; petal-lamina suborbicular, 3 x 3-4 mm. long; petal-hood 4-5 x 2-2.5

mm.; peduncles 2-5 mm. long; pedicels 5-8 mm. long; fruit unknown.
6a. subsp. bernouillii
2. Staminodes subulate, densely pilose, 9-10 mm. long, 1.5 mm. broad;
petal-lamina elliptic, 4 x 3 mm.; petal-hood 5 x 3 mm.; peduncles 6-12
mm. long; pedicels 10-12 mm. long; fruit oblong, 17 x 7 cm., very slightly
pentagonal, attenuate at apex, not contracted at base, with a filiform,

impressed, furrow on each inconspicuous ridge.

6b. subsp. asclepiadiflorum

6a. Theobroma bernouillii Pittier subsp. bernouillii

DistriButTion.—Atlantic coast of Panama.

PANAMA: Prov. Colén, in forests near Faté, Loma de la Gloria (Nombre de
Dios), 10-104 m., 4 VIII 1911, Pittier 4105 (US holotype; isotypes BM, BR, C,
F, GH, K) (Photo USNH 3199).

Further collections at the type locality are necessary to know the
nature of the fruits.

CUATRECASAS—CACAO AND ITS ALLIES 493

6b. Theobroma bernouillii Pittier subsp. asclepiadiflorum (Schery) Cuatr.,
stat. nov.
Theobroma asclepiadiflorum Schery, Ann. Mo. Bot. Gard. 29:360. 1942;
Leén (1960) 316, 321, fig. (as. T. bernouillit).

TyprE.—Wedel 1535, second sheet, flowers (MO, lectotype).
DistrisutTion.—Atlantic coast of Panama.

PANAMA: Bocas del Toro: Water Valley, vicinity of Chiriquf Lagoon; tree
90 feet, flowers red, 8 XI 1940, H. von Wedel 1535 (second sheet MO, lectotype) ;
the first sheet of this collection (MO) belongs to the Lauraceae. Bocas del Toro,
Water Valley, 10 IX 1940; tree 80 ft.; flowers maroon red, H. von Wedel 681
(MO). Bocas del Toro, Water Valley, V 1949, Allan Lucas 1 (F, TURRI).

The fruit of this subspecies is known through one specimen brought
by Allan Lucas (TURRI), which is 17 x 7 em., ellipsoid-oblong, very
slightly pentagonal with 5 filiform furrows on the obtuse angles; the
surface is slightly rugose due to the drying; the apex is shortly attenu-
ate and the base is subtruncate and umbilicate. The shape differs
clearly from that of the fruits of subsp. capilliferum and T. glaucum;
it must be a mutant form geographically isolated.
6c. Theobroma bernouillii Pittier subsp. capilliferum (Cuatr.) Cuatr., stat.

nov. PLATE E

Theobroma capilliferum Cuatr. Rev. Acad. Colomb. Cienc. 6:547, figs. 3a,
4a, pl. 84. 1946; Baker, Cope & al. (1954) 18, figs. 17, 18; Leén (1960)
314, 317, fig.

Typr.—Cuatrecasas 16160, Colombia, Pacific coast.

ComMon NAMES.—Chocolate de monte, cacao de monte bravo,
cacao de monte (Colombia).

Usres.—On the Pacific coast of Colombia and in the Choco area,
the fruits are known as wild cacao (chocolate de monte, cacao de
monte, cacao de monte bravo). Their white seeds are considered a
high quality cacao, but the fruits remain abandoned on the trees, the
people not making actual use of them, although monkeys and other
animals break or pierce them, sucking the pulp or eating the seeds.

DistriputTion.—Pacific Coast and the Chocé region of Colombia.
In the under story of the rain forest, from the lowland swamps next
to the mangroves to the forested hills about 100 m. in altitude.

COLOMBIA: Et Vatue: Pacific Coast, Rio Yurumangui, Veneral, swampy
rain forest in Quebrada del Zancudo, 5 m. elevation; tree 15 m., 25 cm. diam. at
base, bark granulate-rugose, brown or grayish, its section 5 mm. thick, producing
abundant mucilage; wood yellowish white; fruits 11-12 x 6 cm. (immature),
ellipsoid-prismatic, thick, umbilicate at base, more or less constricted above the
base, with 5 furrows or flat sides, and 5 well-marked angles, apex acute and
slightly umbilicate, the surface minutely tomentose, greenish ferruginous, pedun-
cles 6-10 cm. long, thick, fruits abundant, hanging on trunk, leaves coriaceous,
rigid, green above, pale green beneath, long-tailed (Cuatr. photographs C-—2202,
2203), 10 II 1944, Cuatrecasas 16160 (VALLE, holotype; isotype, I’). Pacific

494 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Coast, Rio Cajambre, Barco, forest on hill at the right margin of Quebrada de
Agua Clara, 40-100 m. elevation; tree 20 m., stem 25 em. diam., leaves coriaceous,
pale yellowish green above, cinereous beneath, distichous and pendulous, the
lower ones larger and thicker, dry inflorescences with thin, long branchlets,
fruiting peduncles 8-10 cm. long, 10-12 mm. thick, fruits with 5 furrows and 5
thick angles, surface sinuate-rugose, velvety-tomentose, pale tawny, 16-20 em.
long, 6.5-7.5 cm. broad, umbilicate at both ends, usually constricted above the
base, cotyledons whitish, wood yellowish white, 23 IV 1944, Cuatrecasas 17034
(F, VALLE). Same locality and date, seedlings, Cuatrecasas 17034A (F, VALLE).
Rio Cajambre, San Isidro, hill forest on left margin of Quebrada de Veneno,
about 50 m. alt.; tree 20 m., stem 20 cm. diam., bark yellowish white, leaves
coriaceous, rigid, yellowish green above, greenish cinereous beneath, fruits hang-
ing on trunk, 4 V 1944, Cuatrecasas 17350 (F, VALLE). Same locality, seedlings,
Cuatrecasas 17350A (F, VALLE). Rio Calima, Cafio de la Brea, about 5-10
m. elev.; tree 15-20 m. high, 20-25 cm. diam. at base, crimson flowers in dense
cushions on upper portion of trunk only, 29 VI 1943, Holliday T/142 (TRIN,
US). Rio Calima, Estacién Agroforestal, about 5-10 m. alt.; tree 12-15 m. in
forest clearing, crimson flowers in dense cushions on upper part of trunk only,
dry fruit up to 25 x 8 em., ridged, 29 VI 53, Holliday T/145 (TRIN, US).

Cuocé: Lloré, young tree, sterile, 4 VIII 1953, Holliday & Bartley T/177
(TRIN, US). Ibidem; tree 6 m., Holliday & Bartley T/178 (TRIN, US). Rfo
San Juan, Remolino, young tree 2 m., sterile, 1 VIII 53, Holliday & Bartley T/
172 (TRIN, US).

NariNo: South of Tumaco, in heavy rain forest; tree 10 m., fruits on trunk,
olivaceous, 18 X 1955, Romero Castafieda 5405 (COL).

Mar 6,—Location of known spontaneous, wild, populations of Theobroma cacao subsp.
cacao © and subsp. sphaerocarpum @ which may be the origin of the present cultivated
varieties.

CUATRECASAS—CACAO AND ITS ALLIES 495

Section 3. Theobroma

Theobroma sect. Theobroma Fiaurses 3, 4
Theobroma sect. Cacao (Aubl.) Bernoulli, Uebers. Art. Theobroma 4, 1869.

Sect. Eutheobroma subsect. Cacao (Aubl.) Pittier, Rev. Bot. Appl. 10
(110):779. 1930.

Petal-laminas spatulate, long-attenuate stipitate. Petal-hoods 3-
nerved. Staminodes linear-subulate, erect in aestivation. Filaments
2-antheriferous. Fruit ovoid-oblong or ellipsoid, more or less pentag-
onal, the pericarp thick, firmly fleshy glabrous. Cotyledons epigeous
at germination. Leaves glabrous or sparsely pilose. Inflorescences
on the trunk and on leafy branches. Sympodial growth of stem
by orthotropic adventitious, lateral-subterminal shoots. Primary
branches in 5’s, persisting in age.

Type spEcIEs.— Theobroma cacao L.

7. Theobroma cacao L.
Ficurss 1, 2, 3, 5, 6, 20, 21, 22, 23, 24, 25, 26; Map 6; PLaTE 6

Theobroma cacao L. Sp. Pl. 2:782. 1753; Syst. Nat. ed. 12, 2:508. 1767;
H. B. K. (1823) 316; Richard (1845) 183 (1845a) 73; Bernoulli (1869) 5,
pl. 1, 2; Baillon (1884) 792-795, figs.; Schumann in Mart. (1886) 72, pl. 7;
Jumelle (1899) 11, figs. 1-9; Preuss (1901), pls. 1, 2; De Wildeman (1902)
91, figs. 16, 18, 20, 21. 1902; Standley (1923) 805; Ducke (1925) 130;
(1940) 268, pl. 1, fig. 1, (1954) 11; Fawcett & Rendle (1926) 158-160,
fig. 60; Uittien in Pulle, Fl. Surinam 3:45. 1932; Ciferri (1933) 604;
Standley (1937) 688; Chevalier (1946) 269-274, pl. 5; Standley & Stey.
(1949) 423; Holdridge (1950) 4; Addison & Tavares (1951) 25, pl. 7, pl. 13,
fig. 7; Lemée (1952) 379; Baker, Cope & al. (1954) 9-11; Cuatrecasas
(1956) 653; Leén (1960) 311-313.

Cacao Clusius, Exot. Libr. Dec. 55. 1605; Ray (1688) 1670; Sloane (1696)
134; Tournef. (1700) 660, pl. 444; 1700; Merian (1705), pl. 26;
Sloane (1725) 15, pl. 160; Ray (1733) 158; Weinm. (1739) 1-11, pl. 277,
278; Geoffr. (1747) 409; Catesb. (1747) 6, pl. 6; Blackwell (1739) 373;
Hernandez (1630) 79, (1946) 908-914.

Amygdalis similis guatimalensis, Avellana mexicana Bauh. Pinax Theat.
Bot. 442. 1623.

Arvor cacavifera americana Pluk. Almagest. Bot. 40, pl. 268. 1696.

Theobroma foltis integerrimis Linn. Hortus Cliff. 379. 1737.

Cacao guianensis Aubl., Pl. Guian. 2:683, pl. 275, figs. 1-15. 1775, pro parte
(tantum flores).

Cacao sativa Aubl., Pl. Guian. 2:689. 1775; Lam. Encycl. Meth. 1:533, pl.
636. 1797.

Cacao minus Gaertn. Fruct. & Sem. 190, pl. 122. 1791.

Cacao Theobroma de Tussac. FI. Antill. 1:101, pl. 13. 1808.

Theobroma integerrima Stokes, Bot. Med. 4:83. 1812.

Theobroma caribaea Sweet. Hort. Britt. 67. 1830 (nom. nud.)

Theobroma pentagona Bernoulli, Uebers. Art. Theobroma 6-7, pl. 2. 1869;
Preuss (1901) 199, 255, pl. 3, 5; De Wildeman (1902) 94; Standley & Stey.
(1949) 427.

Theobroma leiocarpa Bernoulli, op cit. 6, pl. 2; Standley (1937) 688; Standley
& Stey. (1949) 426.

496 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Theobroma Saltzmanniana Bernoulli, op. cit. 7, pl. 2.

Theobroma Kalagua De Wild., Bull. Herb. Boiss. 7:957, pl. 11. 1899 (tantum
folia, sed lectotypus).

Theobroma sativa (Aubl.) Lign. et Le Bey, Bull. Soc. Linn. Norm. V, 8: 263.
1904; Chevalier (1946) 270.

Theobroma sphacrocarpa Chevalier, Veget. Util. Afr. Trop. Fr. 4:12. 1908.

Theobroma sapidum Pittier, Bol. Soc. Venez. Cienc. Nat. 1: 183. 1932, nom.
nud.

Theobroma cacao var. typica Ciferri, Monogr. 604. 1933.

Theobroma cacao var. letocarpa (Bernoulli) Ciferri, Monogr. 604. 1933.

Theobroma cacao var. typica x T. cacao var. leiocarpa, Ciferri, 604. 1935.

Theobroma cacao forma letocarpum (Bernoulli) Ducke, Rodriguesia 4: 274,
1940.

Theobroma sativa var. leucosperma Chevalier, Bull. Inter. Bot. Appliq. 26:
270. 1946.

Theobroma sativa var. melanosperma Chevalier, loc. cit.

Theobroma cacao subsp. leiocarpum (Bernoulli) Cuatr. in Macbr. Fl. Peru,
Field Mus. Publ. Bot. 13 (3A):654. 1956.

Theobroma cacao subsp. sativa (as (Lam.) Lign. & Le Bey) Leén in Hardy’s
Cacao Man. 312. 1960.

Theobroma cacao subsp. pentagona (Bernoulli) Ledn, loc. cit.

Type.—Sloane Herbarium vol. 5, p. 59 (BM, lectotype); Paratype:
Tournefort pl. 444, fruit-lectotype.

Tree usually 4-8 m. high, rarely taller (up to 20m.), with the
growth of the sympodial stem by subterminal, lateral, u pright shoots
(chupons); primary branching by successive whorls of normally
spreading branches; young branchlets terete, grayish green or brown-
ish, densely or sparsely pubescent, with slender, patulous, acute,
simple or furcate hairs 0.1-0.3 mm. long, later glabrate, more or less
striate, rugulose and sparsely lenticellate; stipules subulate, very
acute, 5-14 mm. long, 0.5-1.5 mm. wide at base, pubescent or pu-
berulous, deciduous.

Leaves coriaceous or chartaccous, more or less rigid, alternate,
distichous on the normal branches, green; petioles pubescent or
tomentose, with simple, acute, slender, rather dense, spreading hairs,
thickened pulvinate at the ends, 1.5-2 (1-3) em. long, on orthotropus
stems 3-10 cm. long; blades subobovate-oblong or elliptic-oblong,
slightly asymmetrical, rounded or obtuse at base, attenuate and
cuspidate at apex, acute or subacute, usually entire or slightly and
irregularly sinuate, green above, pale when dry, glabrous except for
the pubescent or puberulous midrib, the midrib linear, prominent,
the secondary nerves filiform, the fine veins reticulate, often slightly
prominent, lighter green beneath, glabrous or with very sparse,
minute, simple, furcate or stellate hairs, rarely puberulous, the
midrib thick and prominent, the secondary nerves 9-12 each side,
prominent, subpatulous, then ascending, near the margin curving,
slendering, anastomosing, the tertiary nerves prominent, the minor
veins reticulate and prominulous, 15-50 cm. long, 4-15 cm. broad,
the acumen 1~2.5 em. long.

CUATRECASAS—CACAO AND ITS ALLIES 497

CKE

6.44. REIN

Ficure 20.—a-1, Theobroma cacao subsp. cacao fma. pentagonum (Cuatr. 26004): a, B, C,
petal from inside, outside and laterally, X 5;p, androecium, X 5; , stamen, X 10;
r, anther, X 20; c, pistil, X 10; u, bud, 251, sepal from inside and outside, X 2.
x-0, T. cacao subsp. sphaerocarpum (Cuatr. 7756): K, L, M, petal from inside, outside
and laterally, X 5; N, androecium, X 5; 0, stamen, X 10 and anther, X 20; P, pistil,
< 5; a, sepal from inside and outside, X 2. R-¥, T. cacao subsp. cacao (Nelson 2490):
R, S, T, petal from inside, outside and laterally, X 5; androecium, X 5; v, stamen, X 10
and anther, X 20; w, pistil, X 5; bud and pedicel, 2; ¥, sepal from inside and outside,
x 2.

498 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Inflorescences on the trunk and branches, usually borne on small
tubercles, the cymose branchlets short, knotty, persistent, the cymose
peduncles 1-3 mm. long, stellate-pubescent, hirtellous and with
scattered glandular trichomes; bracts ovate or ovate-oblong, amplec-
tant, pubescent; bracteoles ovate-oblong, acute or subacute, 0.5-1.2
mm. long, pubescent, deciduous; pedicels capillary, rigid, pale green,
whitish or reddish, 5-15 mm. long, hirtellous with rather dense, thin,

Ficure 21.—Sketches of fruits of Theobroma cacao from classical and recent literature:
A, in Tournefort plate 444, clearly representing a ‘“‘criollo type”; B, in Sloane, pl. 160;
c, in Chevalier 1946, pl. 5 (T. sphaerocarpum Chev.).

CUATRECASAS—-CACAO AND ITS ALLIES 499

Ficure 22.—a, Fruit of Theobroma cacao, cultivar ‘‘cundiamor” (Cuatr. 26492 from
Colombia). 3, Cross section of same showing structure of pericarp and arrangement
of seeds each surrounded by their pulp. c, 7. cacao subsp. cacao, cultivar ‘“‘criollo”
(“Caldas”) (Cuatr. 26006 from Colombia). p, Section of same. All X }4.

500 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

acute, patulous, stellate or furcate hairs and sparse pluricellular,
glandular, capitate trichomes; buds white, whitish green, lilac, or
reddish, ovoid or oblong-ovoid, acute, 5-7 mm. long, subglabrous or
sparsely puberulous with slender stellate hairs and thicker, glandular,

Figure 23.—a, Fruit of Theobroma cacao subsp. cacao fma. pentagonum (Cuatr. 26004 from
Colombia) with some degree of introgression from fma. cacao. B, transection of the
same showing the great reduction of the structure of pericarp (mesocarp represented
by isolated bundles). c, 7. cacao subsp. cacao fma. pentagonum, typical (Cuatr.
26540 from Costa Rica). pb, transection of the same, showing the pericarp reduced to
one layer. All X 44 Cultivar “lagarto,”

CUATRECASAS—CACAO AND ITS ALLIES 501

Ficure 24.—a, Fruit of Theobroma cacao subsp. sphaerocarpum, cultivar ‘‘amelonado”
(Cuatr. 25805 from Venezuela, Barlovento). 8, transection of the same. c, fruit of
T. cacao cultivar ‘‘angoleta” (Cuatr. 26494 from Colombia). pb, transection of same.
AllX &.

stipitate trichomes; sepals thick-membranaceous, lanceolate or oblong-
lanceolate, acute, white, greenish white, pale violaceous or reddish,
slightly 3-nerved, shortly (0.5-1 mm.) united at base, 5-8 mm. long,
1.5-2 mm. wide, minutely tomentose at margin, outside sparsely
pubescent with stellate and glandulose hairs, or glabrous, inside
glabrous or with rare glandular trichomes; petals contorted in aestiva-
tion, thick-membranaceous, the hood 3-4 mm. long, 5-2 mm. wide,

002 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Mm ee gt

Figure 25.—a, Fruit of Theobroma angustifolium (Cuatr. 25790). 3, fruit of 7. cacao
subsp. cacao fma. lacandonense (Miranda 9299), c, T. cacao subsp. sphaerocarpum
cultivar “‘calabacillo” (Cuatr, 25805P from Venezuela). p, section of the same. £,
fruit of T. cacao subsp. cacao {. leiocarpum from the original drawing by Bernoulli. F,
T. hylaewm (Araque & Barkley 18C745). All X %,

503

CUATRECASAS—CACAO AND ITS ALLIES

{

aky
Hos

ASS

bipulvinate petiole in Theobroma

cacao: A, T. cacao from orthotropic stem (Steyermark 54143); s, 7. cacao from lateral

branches (Cuatr. 7756);

’

?

Cc

Ficure 26.—a-

T. cacao from young, lateral branches (Steyermark 49218).

D, exceptional, broad, rounded leaf of T. cacao, a cultural mutation (Leén 5078).

c,

?

Showing the characteristic elongated

"op be
a &
1]

zo
af
ei
2”
aN
2
Ss
[e)
ad
o

T. microcarpum (Frées 23963 and Holliday & Cope 125).
pubescens and stipules of T. microcarpum (Holliday & Cope 125),

162). oc, X 3; leaves, X \%.

680-695—64——_9

504 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

obovate, rounded at apex, white, 3-nerved, the nerves papillose inside,
the lateral ones very thick, usually purplish or red, the medial promi-
nent only upwardly; petal-lamina pale yellowish, 1.5-2.5 mm. long,
1.5-2 mm. wide, obovate or rhombic-obovate, attenuate at apex,
acuminate or subtruncate and shortly mucronate, rarely blunt, entire
or sinuate at margin, pedicellate at base, the pedicel linear, 3-nerved
and sometimes 4- or 5-nerved at apex, 2-4 mm. long, 0.2-0.5 mm.
wide; androecium tube rather thick, 1.2-1.5 mm. long, glabrous;
staminodes 4-6 mm. long, 0.6-0.7 mm. broad at base, narrowly
subulate, very acute, erect in bud, erect or somewhat flexuous in
anthesis, the middle vein thick, angular, red or purplish, minutely
papillose-pilose, the thin margin whitish, more or less revolute, ciliate,
with slender, flexuose simple hairs; stamens diantheriferous, the
filaments glabrous, patulous or recurved, 1.5—2 mm. long, the anthers
about 0.4 mm., with rounded cells; ovary oblong-ovoid, obtusely
pentagonal or 5-ridged, about 1.5 mm. high, 0.8 mm. thick, glabrous
or usually glandular, covered with more or less copious white or
reddish, pluricellular, stipitate glands; styles 5, glabrous, adherent,
1.5-2.5 mm. long.

Fruit subbaccate, variable in shape, from globose to fusiform and
acute, and with a very smooth to a strongly ridged and rugose or
verrucose surface; pericarp consistently fleshy and thick (5-15 mm.),
and usually made of two, more or less conspicuously different,
carnose layers (epicarp and endocarp) separated by a thin,
ligneous membrane (mesocarp), the endocarp limited by a firm
epidermis inside, the inner wall of the shell, the epicarp sometimes
differing in color and firmness, rich in mucilaginous cells, limited
outside by the hard epidermis of the fruit, the mesocarpic membrane
sometimes reduced to isolated bundles of fibers or lacking, the endocarp
also sometimes lacking.

Seeds (20-40) usually arranged in 5 rows, but sometimes when
large arranged in 4 or 3 rows, the five radial walls initially separating
the 5 cavites in the earlier stages reabsorbed long before the maturation
of the fruit; seeds ovoid, ellipsoid, amygdaloid, more or less complanate
through mutual pressure or almost round in cross section, variable in
size (20-40 mm. long, 12-20 mm. broad), the integuments brown,
subcoriaceous, the surrounding pulp white, sweet, the cotyledons
white, purplish, violet, or intermediate in color.

Theobroma cacao is variable in its characters, especially with regard
to the color, size, and shape of the parts of the flower and the fruits.
These are variations to be expected of an ancient crop spread
throughout a very wide area.

Based on some of these more or less consistent variations, Bernoulli
described three species, 7. pentagonum, T. leiocarpum, and T. saltz-

CUATRECASAS—CACAO AND ITS ALLIES 505

mannianum, and Chevalier T. sphaerocarpum. The shape of the fruit
is the main defining character, except for T. saltzmannianum, which
was based on petal characters, probably from an abnormal specimen.
The few floral characters given for the other three species are irrelevant
or inconstant. Theobroma pentagonum is defined by having elongate,
gradually and acutely narrowed, warty pods which are strongly pen-
tagonal and 5-ridged; it has white seeds. It was described from the
Atlantic coast of Guatemala and is called “‘cacao lagarto.”” Theobroma
leiocarpum was characterized by ovoid pods, almost smooth, with five
very shallow furrows and a glabrous ovary; the color of the seeds
was not stated; it was found in plantations on the Atlantic coast of
Guatemala, and stated to be rather rare. Theobroma sphaerocarpum
was described from cultivated specimens on Sao Tomé island,
western Africa, characterized by its nearly spherical, slightly 10-
furrowed, almost smooth or slightly rough pods, and violet cotyledons.
Schumann considered the three Bernoulli species as mere local varia-
tions of 7. cacao and therefore unworthy of taxonomic consideration.
Some authors have followed Schumann in this view, but there is a
tendency to accept 7. pentagonum as a different species, because of its
characteristic fruit form and thinner pericarp.

For many years there has evidently been confusion in the taxonomy
of cultivated cacaos, the main problem being insufficient knowledge
of the wild populations of 7. cacao. There are many citations of
places in Central and South America where T’. cacao is said to have
been found wild, which may have been true in some cases, but not in
others where we may be dealing with the remains of abandoned
plantations. The discovery of wild cacao by Stahel in the rain
forests of Mamaboen Creek in Surinam (confirmed later by Myers)
and a few other places very distant from populations, is very significant ;
the cacao is of the Amelonado type. Ogilvie also found it in abundance
on Black Creek, a branch of the Essequebo River, and along the
Berbice River in British Guiana. He found it along the Rupunumi
River at Rewa and Quitaro Creeks, at Kuduwini Kassikedju or
Dewar Wow, etc., on the upper Essequebo River, and also on the
Quitari River (according to Myers). Robert Schomburgk also cited
wild cacao in several places in British Guiana on the Cutari, at the
head of the Correntyne River, and also at Quitaro, and Richard
Schomburgk found it on the Upper Pomeroon River. Wild cacaos
also have been found, according to Stahel, in the Upper Oyapok in
French Guiana (Myers), all of the Amelonado type, implying that
this population of the Guiana highland area westwards to the Amazon
valley may have been the home of this cultivated variety. In Brazil
there has been found wild cacao near the Guianas and, according to
Ducke, at the upper Rio Branco, northeast of Obidos, and at Francez

506 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

on the middle Tapajéz; he adds that the area of spontaneous 7. cacao
includes a greater part of the Hylaea and that it is of the form “leio-
carpum.” Preuss (1901, p. 247) found wild cacao in eastern Ecuador,
also of the Amelonado type. Huber found spontaneous cacao in the
upper Purus river, and along the Rio Ucayali, Peru. Pound also
claimed to have found spontaneous cacaos in the upper Amazon
basin. The spontaneous cacao trees found by the Anglo-Colombian
Expedition in the forests of Caguan (Caquet4) and Rio San Miguel
offer some doubt about their indigenousness. I found myself, in
1939, in the rain forests of Rio Guaviare wild trees of 7. cacao, tall
and well developed, but we have to be cautious in accepting them as
indigenous, because the Guaviare River is said to have had more
extensive plantations on its banks in earlier times. An interesting
find was made in British Honduras by Sampson, who encountered
wild trees of the Criollo type in the Southern forests. In Mexico, Miranda
recently found a completely indigenous cacao tree in a forested region
of Chiapas (Selva Lacandona), very distant from any population; the
natives (Lacandones indians) reported that other trees of such cacaos
are found scattered; this cacao has a somewhat scandent stem and
produces small, elongated, pointed, slightly 10-ridged and rugose
fruits. Standley and Steyermark (Fl. Guat. 426) say that in the
lowlands of Guatemala sometimes cacao is found more or less wild in
the forest, especially in Alta Verapaz, and that it is not improbable
that it is native in the wet North Coast region.

Many forms of the cultivated cacaos have received local or regional
names which after the many introductions of cultivars from one
country to another have brought confusion to the complicated system
of cacao types. Morris was the first to make a classification of
cultivars, arranging them in two main groups: Criollo and Forastero.
Hart (see page 396) modified this, making three groups (Criollo,
Forastero, and Calabacillo), later separating T. pentagonum (cultivar
“Lagarto’”’) into another group, as a different species from 7. cacao.
The classification of Hart has been followed by many authors of
cacao treatises until recent times when van Hall, simplifying, went
back to the basic two groups of Morris: Criollo and Forastero.

Pittier, a botanist with much experience both in Central and South
America, recognized the existence of two different, original, spon-
taneous forms of cacao, Criollo with elongated, ridged, pointed fruits
and white cotyledons, and Forastero, with short, roundish, almost
smooth fruit and purplish cotyledons; he believed that they corre-
sponded with two different species, T. leiocarpum generally spread
throughout tropical South America, where it is still found spontaneous,
and TJ. cacao, spontaneous in Central America and which was the

CUATRECASAS—CACAO AND ITS ALLIES 507

source of the prehistoric cultivation and selection of cacao, all of
the Criollo type, in Mexico and Central America. Introduction of
Criollo types in South America and West Indies and conversely of
the smooth type into West Indies and Central America created the
cross-varieties which multiplied with the years. Although there is
much speculation in this, the theory is a workable and reasonable
one, because the available historical data favors the recognition of
the earlier Central American and Mexican cacaos as being of the
Criollo type. The finding of spontaneous cacao of this type in Chiapas
and British Honduras also supports this theory. Another favorable
fact is the uniformity of the Venezuelan Criollo, supposed to have
been introduced from Central America to Venezuela in the earlier
times of the Spanish conquest.

Soria, after visiting (1961) important plantations in Mexico where
new Forastero types have been abundantly introduced in recent times,
recognized that in Mexico before 1900 varieties of the Criollo type
almost exclusively were cultivated. He observed in large, old
(more than 50 years) plantations a great variation in the Criollo type,
but the seeds were always white and the pods, variable in shape,
were always pointed, with surfaces varying from tuberculate to rugose,
from light green, through green, to reddish; the trees were small,
slow growing, and often with fewer branches (5-3 in each whorl) than
is normal in the species; the petal-laminas are bright yellow.

Pittier’s theory was very much welcomed by botanists and cocoa
experts; Chevalier supported it, and Ducke also in its basic idea.
Cheesman adhered to it at first (1932), but later (1944) developed a
new theory that all cultivated Criollo cacaos came ‘from an offshoot
of a general cacao stock in the headwaters of the Amazon carried over
the Andes into southern Colombia, and developed many of their
present characters in association with man.” But historical knowledge
at present can only closely relate cacao to Central American man,
especially to the Mayans and not to the South American Indians.
Central American Indians undoubtedly developed the art of planting
and selecting of cacao through several thousands of years, finally
obtaining the high quality produce which the Spaniards found at the
time of the conquest.

It may be assumed that in early times a natural population of
Theobroma cacao was spread throughout the central part of Amazonia-
Guiana, westward and northward to the south of Mexico; that these
populations developed into two different forms geographically sepa-
rated by the Panama isthmus; and that these two original forms, when
isolated, had sufficiently consistent characters to be recognized as
subspecies. As they intermingle readily by crossing, giving fertile

508 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

and robust hybrids, they cannot be considered distinct species. Both
types in cultivation have originated many mutations, some of them
persistent, thanks to ecological adjustment, selection, and isolation.
Because the cultivation and selection has been very active for some
thousand years in the Central American-Mexican area, it is in these
areas where we find the richest variety of forms. When the sub-
species (with their different forms) interbred the products gave the
great and confusing variety of existing types.

Another theory is that most of the stable forms of cacao might
have originated by mutations from a widespread, uniform original
specific type. In such a way the forms pentagonum, levocarpum,
Criollo, Cundiamor, Angoleta, Nacional Ecuador, Trinitario, etc.,
and their variants could have originated. This theory does not ex-
clude hybridization as a factor in the multiplicity of forms, but its
influence would be secondary instead of basic, as postulated by the
Pittier theory.

Types or THEOBROMA cacao.—The only specimen of 7’. cacao in
the Linnaean herbarium is the number 934-1, but this specimen is
posterior to the publication of Species Plantarum (1753), since it bears
the annotation “Pl. Surin. 1775. p. 13.” The type has to be sought
among the bibliographic references of Linnaeus. In Species Plan-
tarum, Linnaeus refers to Hortus Cliffortianus. At my request,
Mr. Sandwith was kind enough to examine the Hortus Cliffortianus
herbarium at the British Museum and found no specimens of Theo-
broma in it. However, Hortus Cliffortianus page 379 seems to give a
key to the matter, where Linnaeus writes ‘Flores a nullo bene depicti,
multo minus descripti sunt, . . . Sloane mihi inspiciendi copiam
fecit, .. .” and gives the quotation “Hist. Jam. pl. 160.” Linnaeus
was especially preoccupied with the flowers; he wanted to know
exactly the structure in order to be able to place Theobroma in the
right place in his sexual system; the previous literature did not give
him the answer, and neither did the Sloane plate. However, from the
above paragraph we may infer that Sloane sent him flowers at his
request, surely very few, which may be the reason why there are none
of them in the Linnacan herbarium; from Sloane’s flowers Linnaeus
found the floral structure of Theobroma by himself and drafted his
diagnosis. The original flowers (surely dissected) having disappeared,
the corresponding specimens in the Sloane Herbarium have to be
considered the isotype. Messrs. Dandy and Sandwith found the
specimens in the Sloane Herbarium at the British Museum. Mr.
Sandwith writes: ‘We found the corresponding specimen in the
Sloane Herbarium and it is obviously the source of (in fact part of) the
plate, though not identical; there are leaves, detached flowers, frag-

CUATRECASAS—CACAO AND ITS ALLIES 509

ments of fruit and one seed; it is the vol. 5 p. 59.”’ There is the possi-
bility that Sloane lent Linnaeus these specimens, but even if this were
not the case, we may assume that Linnaeus used flowers taken from
this specimen which would thus be an isotype; therefore, the sheet
page 59, volume 5, of the Sloane Herbarium is to be chosen as lecto-
type for the flowers and leaves. According to Sandwith and Dandy
“there is also what appears to be a duplicate, leaves only, in the
Plukenet Herbarium volume of Herb. Sloane. The leaves of both
specimens are reddish brown and glabrous beneath with reticulate
tertiary veins.”

Sloane’s plate 160 is a paratype, but another even more important
paratype is Tournefort’s plate 444. Tournefort is the only reference
given by Linnaeus in his original description of the genus in Genera
Plantarum (1754), and its citation precedes the reference to Sloane
in Hortus Cliffortianus. I propose it as the lectotype for the fruit,
because the Tournefort drawing is perfectly defined, leaving no doubt
that the characters are of the Criollo type. On the other hand, the
fruits shown on Sloane’s plate 160 are not unquestionable, even
though they are very pointed; they are among the variations found
in Criollo populations. But the reasons which compel me to consider
the flower specimens in the Sloane Herbarium as the lectotype do not
apply to the fruit, the origin of which may have been different from
the origin of the flowers and leaves, for Sloane collected in several
places and countries. A fruit typification by the Tournefort plate
fits well the definition of 7. cacao L. sensu stricto given by Pittier
and later authors. It is obvious that all these authors described
cultivars and that the cacao described by the earlier authors was of
the type Criollo, as can be inferred from the illustrations, and also
from the descriptions of Hernandez, Pison, Plukenet, Merian,
Weinmann, Tournefort, Catesby, and Gaertner.

Synonrms.—T. pentagonum Bernoulli was characterized by the
shape of the fruit and by smaller flowers. Last character is inconstant
but the fruit form is a very particular one and constant, fruit always
easily recognized. This type was described in vivo by Bernoulli
from the Atlantic coast of Guatemala; there is no type specimen
of the fruit, but it was well drawn (Pl. 2, fig. [IJ) and the drawing may
be considered the type. It is my belief that 7. pentagonum is just a
cultivar. It crosses easily with other forms of cacao giving inter-
mediate products. Gross morphological study also supports this
view. The pericarp in Theobroma is composed of three visible layers,
one of these being more or less consistently woody; in 7’. cacao the
woody layer is the middle one, the mesocarp. It seems that in
cultivation there is a tendency for the pods to change, the shells

510 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

becoming thinner in the best quality Criollos. This reduction is
extreme in 7. pentagonum which lacks the firm mesocarp and the
fleshy endocarp, the whole pericarp being only half as thick as in
other cultivars; pentagonum trees are also weaker than others. I
have no doubt that pentagonum is a fixed product of mutation selected
for better fruits. I must agree with Soria (1959) when he writes
“Pentagona is nothing more than one of the extremes in the variability
of the complex of types forming the species T. cacao.”” The name
must be kept but only at the rank of forma.

Theobroma levocarpum was also described by Bernoulli from living
specimens in cultivation on the Atlantic coast of Guatemala; there are
no type specimens of the fruits recorded, and so the published drawing
(Pl. 2, fig. IT) must be chosen as the type. This plant was charac-
terized by a glabrous ovary and smaller flowers (an insignificant
feature) and by ovoid, shallowly 5-sulcate, almost smooth fruits. The
color of the cotyledons was not mentioned. For a long time this form
was identified with the widespread South American ‘“‘Calabacillo” or
“Amelonado”’ fruit types, especially since Pittier published his theory,
it being supposed that the Venezuelan ‘‘Calabacillo,” with thick-
shelled, rounded or ellipsoid, obtuse, slightly 10-furrowed fruits, and
violet cotyledons, was 7’. leiocarpum. All workers followed this nomen-
clature, myself included. It was Cheesman (1944, p. 14) who called
attention to the differences between Calabacillo and Central American
T. leiocarpum. The Bernoulli cacao has thin, ovoid attenuate shells,
with only 5 furrows, and plump sceds which are probably white or
light violet. Preuss had written previously that the seeds were
different. I saw Calabacillo in Nicaragua with very light-violaceous
seeds. The recent observation by Soria (1961) in Mexico of the great
variety in external form of Criollo in an old plantation of Criollo
makes it clear that 7’. lecocarpum is a mere segregate form or mutant
from the Criollo type of T. cacao, originating in a similar way to T.
pentagonum. I consider it only as a cultivar.

Theobroma sphaerocarpum was described by Chevalier from cultiva-
tion in Isla Sao Thomé, in the South Atlantic Ocean west of tropical
Africa, conforms perfectly with the “Calabacillo” cultivar of Venezuela
and other South American plantations. It is an extreme form of the
widespread South American subspecies. The type is a preserved fruit
in the Museum in Paris. This name has to take the place of the sub-
species which Pittier and later authors have called T. leiocarpum.

Cacao sativa Aubl. is a nomenclatural synonym of T. cacao, and
cannot be used as a substitute for the latter. Any imprecision implied
by the binomial 7. cacao is implied also in Cacao sativa, and Theobroma
sativum.

CUATRECASAS—CACAO AND ITS ALLIES 511

Theobroma sapidum Pittier may well have been unintentionally
published as a lapsus calami for T. sativum; it corresponds to T.
cacao sensu stricto of Pittier, restricted to the Criollo type. But the
binomial is a nomen nudum, because it was not formally published,
not being accompanied by a description and with no indication of any
type.

Cacao minus Gaertn. was published without mention of specimens
or locality. It agrees well with some forms of the Criollo type. It
cannot be 7. pentagonum because in 7’. pentagonum the ridges are
always very prominent and smooth. The type of the binomial
consists of two fruits identical with the original drawing, labeled
Cacao minus, Gaertner at Paris.

Cacao theobroma de Tussac, T. integerrima Stokes, and T. caribaea
Sweet are nomenclatural synonyms.

Theobroma saltzmannianum was established by Bernoulli, the emar-
ginate petal-laminae being the only difference from T. cacao (ligulae
lamina anguste obovata, apice truncata emarginata). According to
Bernoulli the shape of the ligula was a constant character in 7. cacao
and 7. leiocarpum; having found at Kew some specimens collected by
Salzmann near Bahia, in which he saw the petal-lamina emarginate,
he did not hesitate to make a new species. Schumann could not
identify this character in any of many flowers collected in Bahia by
Salzmann, and inferred that Bernoulli had examined some exceptional,
teratological flower. Rombouts in 1948 (Kew Bull. 1948: 104)
studied in detail the type specimens at Kew and arrived at the same
conclusion as Schumann, that Bernoulli did base his species on an
accidental character. Chevalier had already expressed the same
view (1946, p. 270). I also can confirm the above opinions after
having examined several Salzmann collections at different times and
the type specimen in 1954 at Kew.

Theobroma sagittata Pavon was published by Chevalier in his revision
(1946: 274) as a microspecies of the complex of T. cacao. The bino-
mial, however, is not validly published for lack of the required Latin
diagnosis. Moreover, I have identified the type specimen, which is
preserved in Paris, as a 3-leaflet fragment of a leaf of Herrania nitida
(Poepp.) Schultes. Theobroma hastata, a name mentioned by Cheva-
lier in the footnote on page 273, presumably is a lapsus calami for
T. sagittata.

The varieties leucosperma and melanosperma published by Chevalier
without reference to any type specimens are fortunately not validly
published for lack of Latin diagnoses. It would be very difficult to
ascribe these two supposed varieties to any recognized taxonomic
entity relying only on the seed color.

512 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

CoMMON NAMES.—-Cacao, cocoa, cacao dulce, cacao criollo, cacao
calabacillo, cacao forastero, cacao amelonado, cacao trinitario, cacao
lagarto, and many other adjectives according cultivar varieties and
regional or local cultivar forms. Also: Bizoya, yagabizoya (Reko),
deghy (Otomi), caocauatzaua (Zoque), kako (Mize), cahequa (Taras-
cdn), Chudechu (Otomi) in Mexico after Standley; cacahuatl, caca-
hoatl, cacahoaquabuitl, quauhcacahoatl, mecacahoatl, xochicacahoatl,
tlacacahoatl, cacahoacuahuitl, cacaotlquahuitl (Nawhatl), kicou, kicob,
cuculat, pacxoc, cucuh, caco in Costa Rica and Guatemala; cacao
chuncho in Peru. For Costa Rica and Chiriqui, Pittier (1902) gives
the following Indian names: ko (Térreba), koé (Brunka), kajo (Gua-
tuso), ku& (Guaimi), kno (Penonomé), dolé (Doraske), tsiri (Bribrt),
(Cabécara); according to him the Bribri Indians use the following
names for some varieties of cacao: murt-uak, tsip4-uak, xi-uak, and
betstin-uak; Standley (1937) mentions, also for Costa Rica: kuk
(Rama), tsiri-kurti (Cabécara), kao-kr4 (Brunka) and kau (Tiribé).
Cacao silvestre, cacao de monte, wild cacao, are names often used in
different countries whenever a cacao tree is found apparently spon-
taneously growing. It is a fact that although the other species have
native Indian names, for Theobroma cacao only the name ‘‘cacao”’ is
recorded from the whole of South America, whereas this species has
many indigenous names in Central America.

SUBSPECIFIC DIvis1ons.—A correct classification of all forms will
only be possible after careful and extensive genetic research. In the
meanwhile we cannot do more than use a provisional, conservative
approach, confining the nomenclature to names already used.

The summarized classification given below of cultivated varieties
follows Morris, who was the first to give status to the most popular
common names of cocoa cultivars; the adaptation in general use made
by van Hall is followed in this paper, with few modifications. See also
my reviews of Preuss (1901), and Hart (1892, 1900 and 1911) in the
Historical Sketch above. The reader will find more extensive infor-
mation on this subject in special treatises (van Hall, Baker in Urqu-
hart, Hart, etc.).

Key to subspecies and forms of Theobroma cacao L.

1. Fruit elongate, claviform, fusiform or ovoid-oblong, tapering and pointed, more
or less strongly 10-costate or 5-costate and verrucose; pericarp moderately
thick, the woody mesocarp thin; seeds ovoid or ellipsoid, usually rounded
in cross section; cotyledons white or yellowish white . . 7a. subsp. cacao

2. Fruit 10-costate. . . . 1. f. cacao, lacandonense and unnamed forms.
2. Fruit claviform, strongly 5-costate, the ridges prominent and smooth, the
sides strongly verrucose, the pericarp thinner, lacking the woody mesocarp
and endocarp. . . ... .. 2 £ pentagonum
2. Fruit ovoid, shallowly 5-furrowed, ‘almost smooth, obtusely attenuate at
apex... 1. ee ee ee eee ew ww). 8 ££. leiocarpum

CUATRECASAS—CACAO AND ITS ALLIES 513

1. Fruit ellipsoid, almost globose or more or less oblong, rounded at both ends,
smooth or slightly verrucose, more or less shallowly 10-furrowed; pericarp
very thick, the woody mesocarp firm; seeds ovoid, more or less compressed;
cotyledons purplish or dark violet... ... 7b. subsp. sphaerocarpum

7a(1). Theobroma cacao subsp. cacao Fiaures 20, 21, 22
Cacao minus Gaertn. 1791.
Theobroma sapidum Pittier, 1932.
Theobroma cacao var. typica Ciferri, 1933.
Theobroma sativa var. leucosperma Chevalier, 1946.

Typr.—Sloane Herb. (flowers, leaves) (BM, lectotype); Tournefort
pl. 444 (fruit-lectotype).

Common NAMzES.—The leading name is cacao criollo or criollo.

DistriBuTion.—Originally from Mexico and Central America, and
cultivated more or less extensively in other tropical countries. Cor-

responds to the Criollo cultivars.

Among the collections examined are:

MEXICO: Veracruz: Colonia San Rafael, flowers and pods, January, 1946;
forests from Colima to Chiapas, Olivia Converse 74 (UC). Fortufio, Coatzacoalcos
River, 30-50 m., III 1937, cacao”; tree 15-25 feet tall, crown fairly dense, trunk
branching from base, inner bark reddish or reddish brown, wood light brown,
fruit yellow about 8 in. long, 4 in. wide, often cultivated and growing wild in low-
lands slightly humid or subject to seasonal floods, (Li.) Williams 8457 (F, P).

Oaxaca: Santa Maria de Chimilapa, I 1927, ‘‘wild cacao,”’ Mell 29 (US).

BRITISH HONDURAS: Stann Creek Valley, Mountain Cow Ridge; tree 5 in.
diam., 30 III 1940, ‘‘wild cacao,” Gentle 3292 (MO, NY, US). Middlesex, 200 ft.
alt.; tall tree of upright habit of growth, generally found growing along river banks,
fruits dark red, occasional, 19 XI 1929, specimen with fruits, Schipp 178 (UC).
El Cayo district, Mountain Pine Ridge, 8 V 1931, Bartlett 13108 (F, NY).

7a(2). Theobroma cacao subsp. cacao forma pentagonum (Bernoulli) Cuatr.,
Fiaurss 5, 6, 20, 23

comb. nov.
Theobroma pentagona Bernoulli, Uebers. Art. Theobroma 6-7, pl. 2, fig. III.
1869.
Theobroma cacao subsp. pentagona (Bernoulli) Leén in Hardy’s Cacao Man.
312. 1960.
Typr.—Bernouilli, op. cit., plate 2, fig. III, isotype, Bernoulli 98
(GOET).

ComMMOoN NAMES.—Cacao lagarto, alligator.

Usrs.—One of the best quality cacaos known.

DistrisutTion.—Only known in cultivation, being probably a
mutant cultivar from 7. cacao L. originally from Central America;
mainly cultivated in southern Mexico and Central America, seldom
in other areas. It is a weaker variety, for which reason it is being
displaced by more robust and disease resistant varieties, in spite of

the high quality of its product.
Collections examined (cultivated) :

514 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

GUATEMALA: Mazatenango, ‘cult. Marz. 1865 Theobroma pentagonum n.
sp.?”; specimen with two leaves and one seed, Bernoulli 98 (GOET, isotype)
Retaluleu, Apr. 1878, Bernoulli & Cario 3151 (GOET).

COSTA RICA: Turrialba; fruit red, “lagarto’”’ seed white, 7 XI 1961, Cuatre-
casas & Soria 26540 (US).

COLOMBIA: Eu Vatue: Finca Esmirna, “lagarto rojo’; seeds white, 16 X
1961, Cuatrecasas & Barros 26004 (US); tbidem, “lagarto amarillo’; seeds white,
Cuatrecasas & Barros 26005 (US).

7a(3). Theobroma cacao subsp. cacao forma leiocarpum (Bernoulli) Ducke,

Rodriguesia 4:274. 1940. Figure 25

Theobroma leiocarpa Bernoulli, Uebers. Art. Theobroma 6, 7, pl. 2, fig. IT.
1869: (not T. letocarpum of current authors).

Theobroma cacao var. leiocarpum Ciferri, Real Accad. Ital. Mem. Cl. Sci.
Fis. Mat. e Nat. 4:604. 1933, as to basionym.

Theobroma cacao subsp. letocarpum Cuatr. in Macbride, Field Mus. Publ.

Bot. 13 (8A): 654. 1956, as to basionym.

Typr.—Bernoulli, op. cit., pl. 2, fig. IT; isotype, Bernoulli 97
(GOET).

CoMMON NAME.—Cumacaco (Guatemala).

Usrs.—A current cocoa of high quality.

DistriBpuTIoN.—Atlantic coast of Guatemala and rarely in other
parts of Central America and southern Mexico. Only known in cul-
tivation, probably being a mutant cultivar of TJ. cacao L. The com-
mercial type known as ‘‘Porcelaine Java Criollo” probably represents
this form.

Collections examined (cultivated) :

GUATEMALA: Mazatenango, “cult. Marz. 1865, Theobroma laeve n. sp.?.,”
Bernoullt 97 (GOET, isotype). Retaluleu, «April 1878, Theobroma laeve Bern.,”’
Bernoulli & Cario 3152 (GOET).

Theobroma cacao subsp. cacao forma lacandonense Cuatr., forma nov.
FigurRE 25

Fructus ovoideo-oblongus, acutus, 10-angulatus, 12 cm. longus, 5.3
cm. diam., pericarpio 5 mm. crasso, epidermide dura, epicarpio molli
2-3 mm. crasso, mesocarpio cartilagineo, endocarpio molli circa 2-3
mm. crasso; semina oblonga 20-22 x 14 x 8-10 mm.

Type in the U.S. National Herbarium, No. 2404633, collected near
Caribal Lacanj& (“selva lacandona’’), northeast of Chiapas, Mexico,
in high primary forest (about 50 m. high), 550 m. alt. by F. Miranda
(No. 9299) ; half-vine, about 7 m. tall, with very long branches; trunk
15cm. diam. The Lacandona Indians call it ‘‘cacao.”

Distrinution.—Only known from the type locality. It is a very
interesting variety due to the fact that it is a true wild plant and
therefore a possible ancestor of the present cultivated cacao.

CUATRECASAS—CACAO AND ITS ALLIES 515

MEXICO: Carapas: Selva Lacandona, Caribal Lacanja, 550 m. alt. in primary
forest, X 1960, F. Mtranda 9299 (US, holotype).

7b. Theobroma cacao subsp. sphaerocarpum (Chevalier) Cuatr., comb. nov.

Fieures 20, 24, 25

Theobroma sphaerocarpa Chevalier, Veget. Util. Afr. Trop. Fr. 4:12. 1908.

Theobroma cacao subsp. leiocarpum sensu Cuatr., excl. basionym T’ leiocarpum
Bernoulli.

Theobroma leiocarpum sensu Pittier et auctt., excl. basionym 7, leiocarpum
Bernoulli.

Theobroma cacao var. leiocarpa sensu Ciferri, excl. basionym T. leiocarpum
Bernoulli.

Theobroma cacao f. leiocarpum sensu Ducke, excl. basionym T. letocarpum
Bernoulli.

Typr.—Sado Tomé, Chevalier, preserved fruit (P).

Common NAMES.—Calabacillo, amelonado, Amazonian forastero,
forastero, etc. (South America). Laranja (Sio Tomé, West Africa).

Usrs.—A currently used cocoa of variable quality; the original type
gives the lowest quality of the cultivated varieties.

DistRIBuTION.—Native in South America, found spontaneous in
the Hylaea from the Guianas and middle Amazonia north and west-
ward to the Andes. Its several cultivars and forms are planted
throughout the tropics, especially in South America and Africa. Be-
ing hard and robust trees, and fast-growing plants, they are spreading
steadily in plantations.

Some of the collections examined are:

BRITISH GUIANA: Mataruki River, upper Essequebo; small riparian tree’
Myers 5829 (K).

COLOMBIA: Mera: San Jose del Guaviare, forest on left side of the Guaviare
River, 240 m. elev.; fruits 18 x 7 cm., abundant, apparently spontaneous trees,
in the lower tree-layer of forest, 14 XI 1939, Cuatrecasas 7756 (COL, F, US).
Sierra de la Macarena, train between Giiejar River and Cafio Guapayita, Cafio
Yerli, 500-600 m. elev.; slender tree 35 ft. tall, 20-28 XII 1950, Idrobo & Schultes
784 (COL, US). Ibidem, tree 12 m. tall, 20-28 XII 1950, Idrobo & Schultes 940
(COL, F. US). Ibidem, Cafio Yerli, dense humid forest; shrub 3 m. tall, 25 XI
1949, Philipson, Idrobo, & Ferndndez 1565, 1569 (BM, COL, US).

Amazonas: La Pedrera, Caquetaé River, river level; old trees possibly planted
long ago and not truly wild on the river banks and islands, pods green, ripening
yellow, 20 X 1952, Baker & Cope 15 (COL, F, US, TRIN).

Caqurtd: Rio Cagudén, Cartagena; tree 7-8 m., found distant from river
bank, white-based pod, smooth and scarcely furrowed, all beans pigmented,
20 IV 1953, Cope & Holliday 105 (COL, TRIN, US). Ibidem; tree 8 m., numerous
small fruits smooth, distinctly 10-ridged, somewhat pigmented, very dark purple
beans, 20 IV 1953, Cope & Holliday 107 (COL, TRIN, US).

MacpA.LENna: Poponte, in forest of Magdalena valley; tree 30 ft., leaves dark
green, flowers yellow, fruit red, 16 XII 1924, Cyril Allen 880 (MO). Ibidem;
fruit green yellow, Allen 881 (MQ).

BRAZIL: Amazonas: Esperanga, at mouth of Rio Javary, in noninundatable
rain forest; small tree, flowers whitish, spontaneous “cacao,” 25 IX 1942, Ducke
1095 (IAN, MG, MO, NY, US); ibidem, Ducke 23976 (US). ‘“B.” Constant,

516 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

inundatable ground; tree; ‘‘cacau silvestre,” 9 V 1945, Frées 20882 (IAN, NY);
ibidem, Frées 20573 (IAN, NY). Rio Solimoes, on an island below Tabatinga;
rain forest, slightly inundatable, small tree, flowers white, spontaneous ‘‘cacao,’’
24 IX 1931, Ducke 23970 (P).

PERU: Loreto: Rfo Ucayali, Laguna de Canchahuaya, 28 X 1898, Huber
1392 (MG). A specimen collected by Asplund near Tingo Maria (No. 13408)
may well represent a new species. I did not study this, the material having been
lent to Dr. Schultes, Botanical Museum, Cambridge.

The cultivars of Cacao are grouped as follows:

Fruit elongated and pointed, warty, 10-furrowed, with 5 ridges more
prominent than the alternate, or only 5, sometimes almost smooth
and only attenuate not pointed; shell medium thick, easy to cut;
seeds usually rounded in cross section, white or yellowish white
inside, and slightly bitter in taste. (Figs. 5, 20, 21, 22, 23.) . . CrroLio

Fruit ellipsoid, rounded at both ends or somewhat narrowed toward
the apex, rather smooth, with 10 more or less marked furrows;
shell thick and harder to cut; seeds usually flattened, violet inside,
bitter in taste. (Some varieties with deep furrows or warty surfaces,
and pale violet or white beans, being intermediate forms with
Criollo. (Figs. 5, 6, 20, 22,24,25.).........0.8. FoRASTERO

Crriotto Morris (1882, pp. 12, 13). This is the type developed
and propagated since prehistoric times in Central America and
southern Mexico and which later acquired importance in western
Venezuela (cacao Caracas). It comprises the best qualities of cocoa,
but the plants are vulnerable to diseases. Criollo is widely cultivated
throughout the tropics, the following cultivars being especially
known by the growers: Venezuela criollo, Nicaragua, Java, Ceylon,
Samoa, Madagascar, Surinam, and Porcelaine Criollos. See van Hall
for more information. Among the Criollos has to be included ‘‘Alh-
gator” or ‘“Lagarto” mentioned above as 7’. cacao f. pentagonum. It
seems to me that ‘‘Porcelaine Criollo’’ corresponds to T. cacao f.
leiocarpum. Figs. 5, 20, 21, 22, 23.

Forastero Morris (1882, pp. 12, 18). This group of cultivars
was divided by Hart into two: Calabacillo and Forastero, the former
containing the most typical, shortly ellipsoid, smooth forms. It
comprises the hardiest, easiest and fastest growing types, but also
those with lower qualities of cocoa; most Forasteros are an inter-
breeding product of Calabacillo and Criollo and their segregates.
Some varieties have developed a good combination of characters.
They originated and spread in South America and Trinidad, especially
in the Amazon and Orinoco Valley. The most common known types
of Forastero are the four listed by van Hall: Angoleta, rather deeply
ridged and warty, about twice as long as broad, with a wide base;
Cundeamor, strongly ridged and warty, narrower than half of its
length, rather acute at the apex, constricted above the base; Amelo-
nado, oblong, ellipsoid, obtuse, rather smooth with rather shallow

CUATRECASAS—CACAO AND ITS ALLIES 517

furrows and not constricted above the base; Calabacillo, shortly
ellipsoid or almost round, the surface smooth with very shallow
furrows. Figs. 5, 6, 20, 22, 24, 25.

The Trintarios also are Forasteros with features intermediate to
Criollo, being variable in shape and in seed characters. They prob-
ably originated by interbreeding in the Venezuelan Orinoco basin,
from which they were introduced to Trinidad where they acquired
new genes from old Criollo plantations and developed extensively.
Later they were brought to Venezuela (receiving the name Trinitario),
to Ceylon, and more recently to other countries. Trinitarios are
cocoas with well-balanced conditions of hardness and quality of prod-
uct. Cacao Nacional of Ecuador is another special Forastero of
superior quality. (See van Hall, Baker, etc.)

At the Eighth Inter-American Cacao Conference (1960) an ‘‘Inter-
national Register of Cacao Cultivars” was established, to be organized
under the chairmanship of Dr. B. G. D. Bartley, of Trinidad.

Section 4. Telmatocarpus

Theobroma sect. Telmatocarpus Bernoulli, Uebers. Art. Theobroma 11. 1869.
FIGURE 4
Sect. Bubroma subsect. Telmatocarpus (Bernoulli) Pittier, Rev. Bot. Appl.
10(110):779. 1930.

Petal-lamina lacking. Petal-hood 5-nerved. Staminodes long-cau-
date, flagelliform, ovate-enlarged at base, flexuose in aestivation.
Filaments 3-antheriferous. Fruit ovoid, ellipsoid or globose, the
pericarp thick, costate-nervate-reticulate and lacunose, pilose, or
tomentulose, the endocarp rigid, thin-woody. Leaves beneath pu-
berulous or subglabrous. Germination hypogeous. Inflorescences
axillary on trunk, small, the peduncles solitary or 2 or 3. Main axis
sympodial with pseudoapical growth; orthotropic shoots from axillary
buds of the terminal jorquette. Primary branching ternate.

Type species: Theobroma microcarpum Mart.

8. Theobroma gileri Cuatr. Ficures 5, 6, 26, 27, 28, 29; Map 7
Theobroma gileri Cuatr. Rev. Intern. Bot. Appl. 33:562, fig. 1. 1953; Baker,
Cope & al. (1954) 13, fig. 19; Leén (1960) 316, 315. fig.

Typr.—Esmeraldas, Ecuador, Manuel Giler 162 (flowers), 168
(fruits).

Small tree up to 14 m. high; growth pseudoapical; trunk 20 cm. in
diameter, brownish, the bark orange in section, the wood white,
rather hard; primary branches ternate-verticillate, light brown, warty
lenticellate, glabrous; terminal branchlets thin, the hornotinous
stellate-puberulous or pubescent; stipules narrowly subulate, minutely
stellate-pubescent, 5-8 mm. long, deciduous.

518 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

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oh XQ
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i

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Map 7.—Geographical distribution of Theobroma microcarpum @ and T. gileri ©, the two
vicariant species (eastern and western of the Andes) of the sect. Telmatocarpus.

Leaves distichous, thin-coriaceous; petioles 3-5 mm. long, mediocre,
subterete, stellate-tomentulose, 3-5 mm. long; blade elliptic-lanceolate,
narrowed and caudate at apex, asymmetrically rounded at base,
entire, 6-22.5 cm. long and 1.5-8 cm. broad (usually 7.5-13 x 2-4 cm.)
including the acumen, this acute, 12-25 mm. long, shining above,
sparsely stellate-puberulous when young, glabrous or subglabrous
when adult except for the midrib, sparsely callous-granulate, the
midrib and secondary nerves conspicuous, with scattered appressed
stellate hairs beneath, these more copious on the main nerves, but
the prominent midrib stellate-tomentulose with smaller stellate hairs,
the 5 or 6 secondary nerves each side filiform, conspicuously prominent,
arched-ascending, near the margin decurrent and anastomosing, the
transverse tertiary nerves prominent, the lesser veins prominulous
loosely reticulate.

Inflorescences on tubercular growths on trunk and on the branches,
the cymes few-flowered, often reduced to one flower, the axis very
short, knotty, bracteate; peduncle erect, filiform, 5-25 mm. long,
3-bracteolate at apex, the pedicel 7-8 mm. long, somewhat thickened
toward the apex, scattered pilose; sepals ovate-lanceolate, rather
thick, reddish or purplish, glabrous inside, greenish or ochraceous
and stellate-tomentulose outside, minutely tomentulose at margin,

CUATRECASAS—CACAO AND ITS ALLIES 519

WAINZECK

CA.

Ficure 27,—a-n, Theobroma microcarpum (Ducke 21045): a, 8, c, petal from inside, outside,
and laterally, X 5; p, androecium, X 5; £, stamen, X 10; F, sepal from inside and
outside, X 2; 6, pistil, X 5;", bud, X 2. 1-N, T. gileri (Giler 162): 1, flower on branch;
j, ovary, X 5; x, androecium, X 5; L, stamen, X 10; M, sepal from inside and outside,
X 2; n, petal from inside, outside, and laterally, X 5.

united 1-1.5 mm. at base, about 6 mm. long, 3 mm. broad; petal-
hoods purplish-red obovate, very narrowed in the lower third, rounded-
cucullate, muticous or emarginate at apex, with no appendix, sparsely
covered with slender hairs above, with 5 prominent minutely pilose
nerves inside, 3-3.2 mm. long, 2.2 mm. broad; androecium purplish
red, the tube about 1.2 mm. high, minutely puberulous; staminodes
about 7 mm. long, the base laminar, ovate, 1 mm. long and wide,
with minute, thick hairs, suddenly narrowed at apex into a subulate,
flexuous, flagelliform, glabrous appendix about 6 mm. long; filaments
white, rather thick, glabrous, shortly trifurcate, triantheriferous;
anthers bilobate, the thecae ellipsoid; ovary ovoid or ellipsoid 5-
angulate, densely stellate-tomentose; styles glabrous, about 1 mm.
long, coherent.

Fruit ovoid or ellipsoid at maturity, 7.5-11 cm. long, 7.5-9 cm.
broad, brownish green or olivaceous, appressed stellate-tomentose,
with 5 longitudinal prominulous ribs and irregularly, loosely, reticulate
prominulous nerves, the intermediate surface shallowly depressed ;
pericarp 1 cm. thick, with a pelliclelike epicarp, a thick carnose, very
mucilaginous mesocarp, and an inner, 1 mm. thick, coriaceous,

680—695—64——10

520 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

ligneous, hard endocarp, inside with a sweet, flavorous, ochraceous-
white pulp surrounding the seeds; seeds ovoid or ovoid-oblong, 2.5-3
cm. long, 1.8-2 cm. broad, 1.5 cm. thick, usually 20-25 in a pod, the

Ficure 28.—Theobroma gileri (Giler 162, 168): a, leaf, X 1; B, young, ribbed fruit, X 14;
c, transection of same; p, mature fruit, X 14; £, long. section of same; F, two seeds
stripped from pulp, X 4; G, seed with its pulp, x 4.

CUATRECASAS—CACAO AND ITS ALLIES 521

Ficure 29.—a, B, Detail of indument on the underside of the leaf in: a, Theobroma gileri
(Giler 162); B, T. microcarpum (Krukoff 1644). c, p, detail of nervation at the under-
side of the leaf in: c, 7. angustifolium (Allen 6259); p, T. subincanum (Cuatr. 7277).
AX 35,B xX 25, Cand D X 2.

522 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

cotyledons white, when fresh condition.!. The juvenile fruits are
ellipsoid or oblong-ellipsoid, with 5 very strong, protuberant ribs and
5 less prominent ribs, these sometimes becoming covered by the
fleshy mesocarp at maturity, but often mature fruits are strongly
10-ribbed; fruiting peduncle 2-3 cm. long, about 6 mm. in diameter;
germination hypogeous.

The fruits of 7. gilert are conspicuously larger than those of T.
microcarpum, its Amazonian vicariant; they are more oblong-ellipsoid
or more oblong in shape. The young fruits have 10 very prominent
and thick ribs, 5 of them much stronger than the alternate, and a coarse
reticulum between the ribs. In a perfect, healthy, mature fruit, the
surface becomes slightly 5-ribbed and shallowly reticulate-lacunose,
the fleshy tissue of the mesocarp having filled up the spaces between
the ribs, as is shown in fig. 28. But often mature fruits keep the 10-
ribbed, strongly reticulate shape of the young stage, looking very
similar then to the fruits of 7. bicolor. Both kinds and different
sizes of fruits are usually seen on the same tree. The trees are com-
monly 8-10 m. high in heavy rain forests. The growth is pseudo-
terminal, that is to say, by shoots produced above the terminal whorl
of branches or jorquette. The branching is normally verticillate
(ternate) but often TJ. gilert develops adventitious, upright shoots
which give irregularity to the branching. Fungus diseases may have
some influence in this. Already in 1953 Patifio wrote: ‘Unfor-
tunately, in spite of the abundance of trees, this species does not
seem to offer much commercial prospects, because almost all fruits we
found were diseased. I sent specimens to Quito to identify the respon-
sible fungi,”’ and he says further: “(Almost all fruits we saw suffer a
disease which hardens the mucilaginous tissue making it compact and
hardening the seeds, many are covered by a mashy down, partly or the
whole.” I have seen myself the same, very recently near Villa
Arteaga. Baker et al. wrote: “Seeds of this species were sent to
Trinidad but arrived in a decomposed condition. The fungus
Monilia roreri Ciferri was found infecting fruit of this species.”

ComMMON NAMES.—Chocolate de monte.

Usrs.—Natives used to chew or eat the pulp, which is sweet and
aromatic. They prepare also a chocolate which is said to have a
good taste (Patifio).

DistriputTion.—Restricted to the rain forests of western Colombia
and northwestern Ecuador.

COLOMBIA: Antioquia: Villa Arteaga; tree 10-12 m., 15-18 cm. in diameter

at base, flowers not scen but borne in cushions on trunk and main branches, fruit
about 9 x 7 cm., ridged and reticulate, said to be yellow at maturity, jorquette

1 Holdridge reports them purplish when cut.

CUATRECASAS—CACAO AND ITS ALLIES 523

symmetrical, 22 VII 1953, Holliday & Bartley T./163 (TRIN, US). Ibidem; tree
about 10 m., dry fruit 0-9.5 x 7-7.5 em., no flowers, 24 VII 1923, Holliday &
Bartley T/167 (TRIN, US). Mutatd, in virgin forest on slope above Villa Arteaga,
250 m. alt.; small tree, 8 m. tall, 18 cm. DBH, cauliflorous and ramiflorous, sepals
greenbacked but reddish inside, petals dark blood red, base of central column
reddish, fruit 10-furrowed, elliptical in long section, about 15 cm. long, seeds
purplish inside when cut, 23 IX 1959, Holdridge 5133 (US). Ibidem, about 150
feet alt.; tree 12 m. tall, 6-20 II 1953, Schultes & Cabrera 18695 (US). Villa
Arteaga, Las Caucheras, Villa Agraria, rain forest, 200 m. alt.; tree 15 m. tall,
stem 8 cm. in diam., crown narrow, primary branches ternate, abundant
inflorescence-cushions (now dry, woody) on the trunk and oldest branches, many
old fruits on trunk and fallen, and some green ones, hanging, leaves chartaceous,
firm, fruits 10 x 7 cm., 10 x 6.5, 11 x 7.5, 11 x 8 em., 2 X 1961, Cuatrecasas &
Willard 26167 (COL, US).

ECUADOR: Esmeratpas: Rio Mira, Guadual, near Piguamb{ camp, 27
VII 1953, Giler 162 (holotype, F, 1423965; isotypes, F, US). Ibidem (fruits),
Giler 168 (paratype, F). Ibidem, Giler & Patifio 164, 165, 166 (H. Cuatr.).
Probably at Rio Mira; fruit collected by Acosta Solis & Giler 12392, 12423 (F).
Santo Domingo de los Colorados, cultivated from seeds brought from Lita,
27 IX 1957, Jorge Leén 4832 (TURRI).

9. Theobroma microcarpum Mart. FicurEs 26, 27, 29, 35; Map 7

Theobroma microcarpum Mart. in Buchn. Repert. 35: 24. 1830; Linnaea,
Litt. Bericht. 32. 1831; Bernoulli (1869) 11, pl. 6; Schumann in Mart.
(1886) 75; Jumelle (1899) 32, fig. 17; De Wildeman (1902) 95; Huber
(1906a) 273; Ducke (1925) 131; (1940) 271, pl. 1, fig. 4; (1953) 14; Chevalier
(1946) 277; Addison & Tavares (1951) 25, pl. 9, 12, fig. 5; Baker, Cope
& al. (1954) 12, fig. 11; Leédn (1960) 316, 321, fig.

Typr.—Brazil, Rio Negro, Martius.

Small tree up to 10-20 m. high; stem 20-30 cm. in diameter; growth
pseudoapical; crown small; branches much ramose, ternate at least
when young, brownish rugulose; branchlets thin, the hornotinous
appressed stellate-tomentose, cinereous or pale ochraceous, becoming
glabrous in age; stipules narrow, subulate, pubescent, 2.5-4 mm.
long, soon deciduous.

Leaves chartaceous, light green; petioles mediocre, subterete,
appressed tomentose, when older transversely wrinkled, 4-8 mm. long
(in very young specimens longer, up to 15 mm.); blades triplinerved,
elliptic-oblong or obovate-oblong, attenuate near the base, rounded
or shortly attenuate at apex, abruptly long-acuminate, asymmetri-
cally rounded at base or sometimes rounded one side, the other
cuneate, rarely symmetrically cuneate, 6-18.5 cm. long, 2-7 cm. broad,
including the acumen, this 0.7-2.5 cm. long, the margin entire, with
scattered minute, simple and stellate hairs above and pubescent costa
or glabrous throughout, the main nerves filiform, conspicuous, the loose
reticulum almost obsolete, with sparse, minute, stellate hairs beneath
or subglabrous, the principal nerves more or less minutely tomentellous
or glabrate, the costa and the lower pair of lateral ascending nerves

524 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

prominent, the other 2-4 secondary nerves each side thin but
prominent, ascending, curved near the margin, anastomosing, the
slender, transverse tertiary nerves prominulous and the minor
reticulate veins less prominulous, but conspicuous.

Inflorescences axillary or extra-axillary on young leafy branchlets,
the cymose clusters extremely small, bearing 1-3 flowers, the woody
primary branches short, knotty, bearing ovate, amplectant bracts,
0.6 mm. long and wide; peduncles rather thick, 0.5-1 mm. long,
tomentulose, with 3 subulate, deciduous bracteoles at apex, these
0.6-1 mm. long; pedicels 0.5-1 mm. long, moderately thick, tomentu-
lose; buds ovoid, acute, 4.5 mm. long, 3 mm. broad, with 5 whitish,
minutely tomentulose lines and scattered, minute stellate hairs.

Sepals thick-membranaceous, lanceolate, acute, united about 1 mm.
at base, 5-6 mm. long, 2 mm. broad, with scattered, minute, stellate
hairs outside, minutely, whitish tomentulose at margin, with con-
spicuous midrib and sparse, mediocre, flexuous, glandular hairs
inside; petal-hoods pale brown, thick-membranaceous, glabrous,
oblong-obovate, rounded-cucullate and acuminate at apex, the acumen
triangular, shortly bidentate, exappendiculate, the 5 nerves rather
thick, prominent and minutely papillose inside, often confluent into
pairs near the base; androecium red or reddish, the tube about 1.5
mm. high, 10-furrowed, minutely pilose; staminodes minutely pilose
with a rather thick, oblong-ovate concave base, this 2.5 mm. long and
1 mm. broad with fine flexuose hairs inside, topped by a subulate,
flexuose tail 4-5 mm. long; filaments rather thick, 1 mm. long, shortly
3-furcate, triantheriferous; cells of anthers ellipsoid, about 0.4 mm.
long; ovary 1.5-1.8 mm. long, pyriform, glabrous or sparsely, minutely
stellate-pilose, sharply 10-furrowed-costate; styles 5, united in the lower
fourth, rather thick, acute, glabrous.

Young fruit ellipsoid (+3 cm. long), strongly 10-ribbed, with 5 very
thick and prominent, dorsal, shortly pinnate costae, and 5 smaller,
commissural ones; mature fruit 6.5-7 (-9) em. long, 5.5-6.5 cm.
broad, green yellowish, puberulous, ellipsoid-globose, conspicuously
10-ribbed, the surface between the ribs shallowly alveolate, the
exocarp thick, carnose, padding the hollows between the lignose
underlayer, when dry the ribs and the lignose reticulum become ex-
tremely marked and prominent; seeds more or less compressed, ovoid,
12-14 mm. long, 18-20 mm. broad, and 11-12 mm. thick; fruiting pe-
duncle thick, robust, 4-8 mm. long and broad; cotyledons hypogeous
at germination.

ComMMON NAMES.—Cacauf, cacaurana, cacao rana, cacau bravo,
cabeca de urubti (Brazil). Cacao de monte (Colombia). Me-tré-
ree-moo-ee (Karihona, Upper Apaporis); béo-e (Mirana, Caqueta
River) (Angl-Col. Cocoa Exp., Baker, 1952).

CUATRECASAS—CACAO AND ITS ALLIES 525

Usrs.—None recorded on the use of its sweet scentless pulp or the
seeds.

DistrRisuTION.—In the southern and western upper Amazon
Brasilian region Rios Solimoes, Yapurd, Puris, Madeira, Tapajés,
and western Colombia on Caqueté River. (Baker & Cope.) The
eastern known limit according to Ducke is Rio Tapajés. Thespecimens
around Belém are cultivated. It is frequent in its area and may
become abundant in some places as a significant element of the shad-
owy under layer of rain forests on elevated ground and in moder-
ately inundatable alluvial lands.

COLOMBIA: Amazonas: Rfo Caqueté, La Pedrera, river level; tall tree 15-
20 m., 30 cm. in diam. at base, extensive branch system, jorquettes of seedlings 3-
branched, growth continuing from above, flowers small, petals without ligules,
fruit abnormal due to attack by Marasmius perniciosus, 5 X 1952, Baker & Cope
28 (COL, K, TRIN, US). Ibidem; tall tree 15-20 m., native in forest on the river-
bank, 7 X 1952, Baker & Cope 29 (COL, F, K, TRIN, US). Ibidem; tree 30
feet, 9 inches in diameter, on floodbank, 5 X 1952, Schultes & I. Cabrera 17780
(AMES, US). Rio Caquet4é, Remolino; leaves only from small seedling tree
2.5 m., typical Theobroma habit, jorquettes arising symmetrically, 2 V 1953,
Cope & Holliday T/125 (COL, TRIN, US).

BRAZIL: Amazonas: In sylvis ad Costa de Ubicuna et de Camarocoari, fluv.
Solimoes, prov. Rio Negro (Dr. Martius Iter Brasiliensis, 321”), Martius Observ.
2890, [884] (M, lectotype, photo F. M. 19643). Ibidem, Martius Observ. 2890,
[885, 886] (M, isotypes). Lower Rio Yapurd, Jubar4 matta, 15 IX 1904, Ducke
6773 (BM, MG). Basin of Rio Solimoes, Municipality Sao Paulo de Olivenga,
near Palmares; tree 60 ft. high, trunk 7 inches in diam., terra firma, high land,
11 IX-26 X 1936, Krukoff 8280 (A, BM, F, G, GB, K, LE, MICH, MO, P, §S, U, US,
USDA). Ibidem; mata, caatinga, “cacau bravo,” arvore pequefia, 19 IV 1945,
Frées 20750 (IAN, USDA), 34814 (IAN). Basin Rio Madeira, Municipality Hu-
mayta, near Livramento, on Rio Livramento on varzea land; tree 50 feet high,
“cabeca de Urubt,”’ 12 X-6 XI 1934, Krukoff 6592 (A, BM, F, G, K, LE, MICH,
MO, 8, U, US, USDA, WU). Ibidem, Municipality Humayta, near Tres Casas,
on restinga alta; tree 60 ft. high, 14 IX-11 X 1934, Krukoff 6203 (A, F, G, K, LE,
MO,§, U, US, USDA, Y). Rio Purus, Bom Lugar; ‘cacao rana,’’ IT 1904, Goeldi
4228 (BM, G,MG). Camatian; high forest lowland, border of creek; tree 7 m.
high, 25 cm diam., 24 I 1949, Frées 23963 (IAN, US).

Guapore: Porto velho, Entrada de Redagan, Km. 8, Viana, mata derrum-
bada, terra firme; arvore pequena, 31 V 1952, Black, Cordeiro, & Francisco 52-
14649 (IAN, UC).

Mato Grosso: Machado River region, source of the Jatuarana River; tree
3 feet high in terra firma, ‘“‘cabeca de urubu,” XII 1931, Krukoff 1644 (A, BM, F, G,
K, MICH, MO, P, 8, U, UC).

Part: Belém, Jardim Botanico do Museu Goeldi; medium tree, cultivated,
11 VIII 1942, Archer 7551 (F, IAN, K, USDA). Belém, Horto Botanico Pard
(cultum proven. Rio Purus, Bom Lugar anno 1904), 25 V 1906, Huber 7081 (G, MG).
Ibidem; arbor parva floribus rubescentibus, 4 II 1926, Ducke 21045 (G, GH, K, 8,
U, US). Ibidem; arbor parva, floribus pallide brunnescentibus, ‘“cacaohy,”’
IX 1936, Ducke 283 (A, F, K, MO, 8, US). Ibidem, 21 VII 1944, F. C. Camargo 8
(IAN). Ibidem, 23 XI 1945, Pires & Black 742 (IAN). Rio Guama, near Belém,
“cacao bravo,” IV 1929, Dahlgren & Sella 10 (F, GH). Rio Tapajos, Cachoeira
do Mangabal, beira de assahyzal, 7 IX 1916, Ducke 16466 (BM, G, MG, P, US).

526 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

COSTA RICA: (cult.) Limon, La Lola, experimental station IICA; tree 4 m.
tall, narrow crown, eight years old, first flowers this year, 7 XI 1961, Cuatrecasas
& Paredes 26538 (US).

Section 5. Glossopetalum

Theobroma sectio Glossopetalum Bernoulli, Uebers. Art. Theobroma 11. 1869

Ficures 1, 3, 4; Map 2

Sect. Bubroma Schum. in Engl. & Prantl. Nat. Pflanzenfam. 3(6):89. 1890.

Sect. Bubroma suksect. Glossopetalum (Bernoulli) Pittier, Rev. Bot. Appl. 10
(110):779. 19380.

Petal-laminas obovate, spatulate or trapezoid, stipitate. Petal-
hoods 7-nerved. Staminodes laminar, petaloid, obovate or broadly
lanceolate, curved-reflexed covering the hoods in aestivation, erect or
reflexed in anthesis. Filaments 3-antheriferous. Fruit ellipsoid or
oblong, smooth or more or less angulate or tuberculate, the epicarp
hard, woody, with a tomentose epidermis. Cotyledons hypogeous
at germination. Leaves beneath reticulate-nerved, stellate-tomen-
tose. Inflorescences on the main trunk or on the branches. Main
axis sympodial with pseudoapical growth; orthotropic shoots from
axillary buds of the terminal jorquette. Primary branching ternate.

Typr spEcies.— Theobroma grandiflorum Schumann.

10. Theobroma angustifolium Mogifio & Sessé

Fiaures 6, 25, 29, 30, 31, 37; Mar 8
Theobroma angustifolium Mogiiio et Sessé ex DC. Prodr. 1: 484. 1824; Icon.
Fl. Mexicana ex DC. pl. 112; Bernoulli (1869) 12, pl. 6; Schumann in Mart.
(1886) 77; Donn. Smith in Pittier, Prim. Fl. Costar. 96. 1898; Preuss
(1901) 255, pl. 2,4;De Wildeman (1902) 96, figs. 12,13; Standley (1923)
808; Standley (1937) 687; Chevalier (1946) 282; Standley & Steyermark
(1949) 421; Holdridge (1950a) 3; Allen (1956) 342, pl. 26; Leén (1960)

318, 315, fig.

Typp.—sSessé et Mogifio, Plantae Novae Hispaniae, Herbarium
Florae Mexicanac.

Tree 8-26 m. high; trunk up to 30 cm. in diameter, with smooth
bark and whitish wood; growth pseudoapical; primary branches ternate ;
lower branches horizontal, the higher ascending; branchlets when
young green ochraceous, densely and moderately appressed tomentose,
with very minute, fine, translucid-white stellate hairs intermixed with
other mediocre, fulvous or ochraceous, somewhat thicker, stellate
hairs, when older more or less glabrate, grayish, rugulose; stipules
lanceolate-subulate, acute, broadened at base, above sparsely, below
densely stellate-tomentose, 5-7 (-15) mm. long, about 1 (-2) mm.
broad, deciduous.

Leaves distichous, thin-coriaceous, rather flexible; petioles moder-
ately thick, densely subappressed tomentose, 6-10 mm. long; blades
subobovate-oblong, elliptic-oblong, or oblanceolate, slightly narrowed

CUATRECASAS—CACAO AND ITS ALLIES 527

Map 8.—Geographical distribution of Theobroma angustifolium @ and T. simiarum ©.

to the obtuse and slightly asymmetrical base, attenuate and acuminate
at apex, entire or at the upper part slightly sinuate-dentate, 9-25 cm.
long and 3-9 cm. broad, including the 1-2 cm. long and 3-5 mm. wide
acumen, green above, when adult smooth, glabrous or with few hairs
scattered on the costa, this depressed, filiform, the secondary and
tertiary nerves little conspicuous, light greenish or cinereous beneath,
appressed tomentose, heteortrichous, the surface covered with a dense
layer of white, minute, stellate hairs, and additional, more or less
copious, larger, ochraceous, stellate hairs with longer rays on the main
nerves, the costa very prominent, the 6-8 secondary nerves on each
side thinner and prominent, ascending, near the margin decurrent,
becoming slender, vanishing, the transverse tertiary nerves, thin,
prominulous, 2-5 mm. from each other, the minute reticulum conspic-
uous.

Inflorescences usually abundant on the branchlets, axillary or
extra-axillary, the cymes strongly reduced to a few extremely short
1-3-flowered branchlets; peduncles 0.4-1mm. long, 3-bracteolate;
pedicels erect, rigid, mediocre, densely ochraceous or ferruginous,
ebracteate, 5-10 mm. long; bracteoles very minute (1-0.5 mm. long),
linear, deciduous; buds globose, 7-8 mm. broad, densely ochraceous
tomentose; calyx 8-9 mm. long, reflexed in anthesis, all the sepals
united to 3-4 mm. into a cupular base, in the upper part two united

528 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

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Figure 30.—Detail of indument on the underside of leaf in: a, T. angustifolium (Allen
6259); B, T. cirmolinae (Cuatr. 14897); c, detail of venation in 7. stipulatum (Cuatr.
21339); p, indument in 7. stipulatum (Cuatr. 21339). A, B, and DX 30, C X 2.

Ficure 31.—a-r, Theobroma nemorale (Patino 116): a, B, petal from inside and laterally,
5; c, androecium, X 5; p, stamen, X 10; EF, gynoecium, < 5; F, sepal from inside
and outside, X 2. GM, T. angustifolium (Allen 6341): G, H, petal from inside and
laterally, X 5;1, androecium, * 5; J, stamen, X 10; x, gynoecium, X 5; 1, bud, X 2;

529

sett
wt
se
& SS
Ss

CACAO AND ITS ALLIES

CUATRECASAS

sews, oS

ee “Se
ers
s

en !
SONAL a ome a F

Ain Ne actin pe een ee

[Ficure 31]

Nn, the three

.
.

n, 0, T. nemorale (Cuatr. 26007)

M, sepal from inside and outside, * 2.

o, pedicelled bud surrounded by a

bracteole, the other two removed. p, T. chocoense, staminode, X 5 (Cuatr. 26074).

bracteoles covering the opening flower, X 2

530

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

it Mie

TSR \ wk ,
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[Figure 32]

CUATRECASAS—CACAO AND ITS ALLIES 531

by pairs forming 3 unequal lobes, two of the lobes twice as broad as
the third, ovate, rather obtuse, scarcely puberulous and with minute,
thick, oblong glandular hairs at the base inside, tomentose, greenish
ochraceous or ferruginous outside, each single sepal 4-5 mm. wide.

Petal-hoods thick-membranaceous, yellowish, broadly obovate,
rounded-cucullate at apex, 7-nerved with very sparse, thin hairs
outside, about 5 mm. long, 4 mm. broad; petal-lamina pedicellate,
yellow, thick-membranaceous, 5-nerved and finely veined, subobovate-
spatulate, emarginate at apex with 2 ovate or rounded lobes, atten-
uate towards the base, glabrous, about 5.5 mm. long, 3 mm. broad,
the pedicel linear, 3-nerved, attenuate downward, about 4-5 mm.
long, 1 mm. wide; androecium tube 2.5-3 mm. long, thick; staminodes
laminar, thick-membranaceous, sulphur yellow, glabrous, oblong-
obovate, rounded or subspatulate at apex, with marked venation,
8-11 mm. long, 4.2-5 mm. broad, at base 1.2 mm. broad, when in
bud curved-reflexed, in anthesis erect; filaments glabrous, thick,
curved, 2 mm. long, enlarged and shortly 3-furcate, 3-antheriferous;
anther cells globose; ovary obovoid-oblong, about 1.5 mm. long,
5-sulcate, densely stellate-tomentose; styles 2.5 mm. long, united in a
rigid erect column 1.5 mm. long, ending in 5 slender branches 1 mm.
long.

Fruit unequally oblong-ellipsoid or ovoid-ellipsoid, more or less
pentagonal, slightly attenuate at apex, umbilicate and 5-costate at
base, very irregularly tuberculate-rugose, densely brown tomentose,
the epicarp hard, ligneous, about 1.5 mm. thick, the mesocarp plus
endocarp carnose 5-6 mm. thick, the pulp enveloping the seeds thick,
juicy, aromatic, edible, 10-18 em. long, 6-9 cm. broad; seeds 5-7 in
each fruit compartment, compressed oblong-ovoid, 26-32 mm. long,
16-19 mm. broad, and 14-16 mm. thick, the cotyledons white; ger-
mination hypogeous.

The leaves of T. angustifolium are similar to those of T. nemorale,
but they are narrower, rather lanceolate, and the larger hairs of the
double indument beneath are longer with much finer, longer rays
than those of 7. nemorale. Paul Allen (1956) writes about this tree:
“The young branches, petioles, and veins of the lower leaf surface are
covered with a rather scurfy pale-tan tomentum. The relatively

Ficure 32.—a-c, Theobroma stipulatum (Cuatr. 21339): a, B, petal from inside and later-
ally, X 5;¢, androecium, 5; p,stamen, X 10; , gynoecium, X 5; F, sepal from inside
and outside, X 2; .c, bud, X 2. u-n, 7. simiarum (Tonduz 7313): nu, 1, petal from
inside and laterally, X 5; 5, androecium, X 5; x, stamen, X 10; 1, gynoecium, X 5;
M, sepal from inside and outside, X 2; n, bud supported by pedicel and 3-bracteolate
peduncle, X 2. o-v, T. cirmolinae (Cuatr. 14897): 0, p, petal from inside and lat-
erally, X 5; Q, androecium, X 5; R, stamen, X 10; s, gynoecium, X 5; T, sepal from
inside and outside, X 2; u, pedunculate bud, x 2.

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

532

> i
Lore

ror :

S2——
= e Ze

SLLEFIS
LE go

ee ie

[Ficure 33]

CUATRECASAS—CACAO AND ITS ALLIES 533

large, bright-orange flowers are produced in great profusion in several
successive flowerings from November until February from the axils
of the slender, younger branches, and are followed in August and
September by the large, woody, brown-tomentose, cacaolike pen-
dulous pods which are from 4” to about 7” in length.”

ComMMoN NAMES.—Cacao de mico, cacao silvestre (Costa Rica),
cushta, cacao de la India (Salvador), cacao silvestre (Mexico).
Cacao de mico, cacao meco, coca mono (Nicaragua), soré (Bribi
indians).

Usxs.—It was stated by Standley (1923) that 7. angustifolium was
much planted in southern Mexico, especially in Chiapas as a source
of commercial cacao and that the famous Soconusco cacao was from
this species. This statement is very doubtful, for the seeds of T.
angustifolium are considered at present in that area as of inferior
quality without commercial value.

DistriBuTION.—This species is often planted in Central America
and southern Mexico. It is certainly native in the lowland forests of
the Pacific range of Costa Rica (Allen) and nearby Central American
countries (Holdridge). Standley and Steyermark (1949, 422) say
that the native region of this cacao is unknown, but Tonduz already
in 1891 cited it from the forests of Terraba; Leén (No. 937) writes:
‘Important tree in the regional forests,” and Allen says that ‘it is
locally frequent in lowland forests throughout the area [Golfo Dulce].”’

MEXICO: Direcién de Estudios Biolégicos, Ord. 34-G823 (MEXU). Her-
barium Sessé & Mocifio in M, “18-1, Theobroma Simiarum N. Ic,” Sessé, Mogziio,
Castillo, & Maldonado 3618 (holotype, MA; isotype, F, Photo F. M. 48411).
Sessé & Mogifio s.n. (BM, probable isotype). Copy of the Sessé & Mogifio drawing
at G (Negative F. M. 30527), plate 112 of the DC. published series.

GUATEMALA (cult.): Retaluleu, April, 1877, Bernoulli & Cario 3188 (GOET,
K, 8). Mazatenango, III 1865, cult., Bernoulli 95 (NY, BR). Region of Plata-
nares, between Taxisco and Guazacapén (Dept. Santa Rosa), 220 m. alt., wet
forested quebrada; small tree escaped here, 3 XII 1940, Standley 79068 (F, US).

SALVADOR (cult.): Sonsonate, ‘“‘cushta,’”’ 1922, Calderén 630 (GH, NY, US).
Vicinity of Sonsonate, 220-300 m. alt.; tree 20-30 feet, very dense and narrow
crown, flowers on branches, fruit brown, the pulp edible with very aromatic odor,
seeds give chocolate, grown here only in finca, “‘cushta,’”’ ‘cacao de la India,”
Standley 22317 (GH, MO, NY, US).

Ficure 33.—a-u, Theobroma grandiflorum (Ducke 598): a, B, petal from inside and
laterally, X 5; c, androecium, X 5; p, stamen, X 10; E, gynoecium, X 5; F, styles,
> 5; G, sepals from inside, X 2; u, sepals from outside, X 2. 1-0, T. obovatum (Ducke
265): 1, J, petal from inside and laterally, X 5; x, androecium, X 5; 1, stamen, X 10;
M, gynoecium, X 5; N, sepal from inside and outside, X 2; 0, bud supported by bracte-
olate peduncle and pedicel, X 2. P-w, T. subincanum (Baker & Cope 32 and Holliday
43): p, Q, petal, from inside and laterally, X 5; R, androecium, X 5; s, stamen, X 10;
t, bud, X 2; u, sepal from inside and outside, X 2; v, initiation of fruit, X 5; w, ovary,

x 5.

534 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

NICARAGUA: Quezalguaque, Dept. of Leén, Baker 2102 (A, AMES, C, GH,
G, K, L, MICH, MO, NY, U, UC, US, WU). Chichigalpa, ‘cacao de mico,”
II 1900, Preuss 1381 (UC). Belén, “cacao meco,” ‘coca mono,” or ‘‘monkey
cocoa,” 28 VI 1893, Hart 5381 (K, NY, U).

COSTA RICA: Guanacaste: Upper portion of cafién of Rfo San José, 460-480
m., 12, 13 I 1930, Dodge & Thomas 6399 (F, GH, MICH, MO, UC, US). Nicoya,
300 m. alt., VI 1949, Lopez s.n. (F, TURRI). Ibidem, Pittier sn. (US). Ho-
jancha de Nicoya, 20 m. 4rbol importante en la selva de la regién, ‘‘cacao de mico,”
29 I 1942, Leén 937 (F). Perico, Nicoya, 100 m., I 1954, Leén 4267 (TURRI).
Borde du Rfo Zurquin, 3 1894, Pittier & Durand 8536 (P).

Limon: Cienaguita, near Puerto Limén, 10 m. alt.; small tree with ap-
pressed crown, flowers sulphur yellow, VII 1901, Pittier 16142 (G, US, WU).
La Lola, I.I.A.C.A. experimental station, about 20 m. alt.; tree erect, trunk about
25 cm. diam. at base; pseudoapical growth; primary branches ternate, spreading,
persistent; leaves thin-coriaceous but firm, light green and somewhat glaucous and
cinereous beneath; young fruits thickly tomentose, axillary flowers abundant, now
dry, 6 XI 1961, J. Cuatrecasas & Paredes 26537 (US).

PunTARENAS: Dans le forét a Terraba, 260 m. alt., II 1891, Tonduz 4074
(US). Ibidem; ‘‘cacao de mico,” II 1891, Pittter & Tonduz 4074 (BR, G).
Boruca, Diquis Valley, 1891, Pittter s.n. (US). Tinoco Station, fairly frequent in
swampy forest; tree 80 ft., fruits pendulous, produced from the ends of branches,
“cacao de mico,”’ 13 VII 1951, Allen 6259 (BH, MO, US). Lowland forest near
Palmar Norte; tree 45 feet, flowers bright orange, Allen 6341 (BH, F, MO, US).
Llanuras de Corredor (Golfo Dulce), III 1897, Pitter 11112 (GQ).

PANAMA: Progreso (Chiriquf); small tree 30 feet, 6 inches in diam., with a
fruit like wild cacao except that the husk is smooth like a potato skin, big seeds
with white meat inside, 1927, Cooper & Slater 242 (F, GH, NY, US, Y). Comarca
del Bart, Puerto Armuelles, United Fruit Company farms between Canasco and
Cocos, mostly cutover land with some of the original trees still standing, about
100 feet alt.; tree, fruit resembling a cacao pod; leaves pale bluish, green, beneath,
17 VI 1957, Stern & Chambers 140 (MO, US, Y).

TRINIDAD: Royal Botanic Gardens, Port-of-Spain, LZ. H. Bailey, s.n. (BH).
Ibidem; small tree, flowers orange colour, 8 IX 1918 (originally from Guatemala),
Broadway 8935 (BM, BR, MO, 8). Ibidem; tree 8 m., flowers orange yellow,
10 VII 1953, R. BH. D. Baker s.n. (TRIN). Government House Gardens, 12 III
1929, Williams 12121 (TRIN). St. Augustine, Imperial College of Tropical Agri-
culture; tree 3-4 m. alt., lower branches horizontal, upper ones ascending, bark
granulose, lenticellate, more or less cleft, yellowish brown, abundant, hanging,
very rugose dried fruits, 1 IX 1961, Cuatrecasas & Cope 25789 (US). Ibidem;
tree about 8 m. (9 years old), trunk 20 cm. in diam. at base, primary branches
ternate from near the base, leaves thin-coriaceous, yellowish green, shining above,
greenish cinereous beneath, 1 IX 1961, Cuatrecasas & Cope 27591 (US).

11. Theobroma cirmolinae Cuatr.
FiaureEs 3, 4, 30, 32, 34, 36, 37; Mar 9; PuLats 7
Theobroma cirmolinae Cuatr. Notas Fl. Colomb. VI: 5, fig. 1-8 1944;
Rev. Acad. Colomb. Cienc. 6:32, fig. 1-5. 1944; Llano (1947) 34,
pl. 14; Baker, Cope & al. (1954) 13, fig. 22; Leén (1960) 320, 317, Jig.
Typr.—El Valle, Colombia, Cuatrecasas 14897
Medium-sized or large tree up to 20 m. high; growth pseudoapical;
trunk up to 40 cm. in diameter, branched in the upper third, the bark
dark grayish, somewhat rimose-scaly, under the periderm brown or

CUATRECASAS—CACAO AND ITS ALLIES 535

&0° 70° 60° 50° 40°

10°

E85, QuiB22.
5

H Ww.
ognat wos

2o°

||

Map 9.—Geographical distribution of Theobroma stipulatum @, T. cirmolinae O, T.
chocoense @, T. mammosum ©, T. sinuosum A, and T. canumanense If.

rufous, the wood ochraceous, the hardwood ochraceous reddish, very
hard; branches gray or brownish gray, the primary ternate, the
terminal leafy branchlets tawny or ferruginous, appressed stellate-
tomentose; gum resin flowing easily from bark and wood; stipules
large, persistent, subcoriaceous, oblong-lanceolate, subacute, ochra-
ceous-tomentose, 12-22 mm. long, 3-5 mm. broad at base.

Leaves of the young branchlets (smaller than in adult) thin-
chartaceous, green-ochraceous with scattered stellate hairs above,
green ochraceous or grayish green, appressed stellate-tomentose
beneath; adult leaves large, thick-coriaceous, rigid, shortly petiolate;
petioles very strong, thick, subterete, appressed ochraceous-tomentose,
1-2 cm. long, 6-8 mm. broad; blades oblong-elliptic or ovate-elliptic,
slightly asymmetrical, rounded, cordate, or sharply emarginate at
base, little attenuate, rounded or very obtuse and shortly acuminate
at apex, entire or very slightly sinuate and flat at margin, 26-54 cm.
long, 14-30 cm. broad, ochraceous green above, pale brown when
dry, apparently glabrous but with scattered, appressed stellate
hairs, these more copious on the main nerves, the costa and the
secondary nerves filiform and depressed, the minor venation less
noticeable, velvety-tomentose beneath, the surface rosy-glaucous,
the veins somewhat more ferruginous or rufescent, the costa very
thick and prominent, the 12-14 secondary nerves on each side very

680-695—64——11

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

536

ries
=

Ficure 34.—Theobroma cirmolinae, flowering and fruiting trunk, X %.

CUATRECASAS—CACAO AND ITS ALLIES 537

Ficure 35.—Fruits of Theobroma, * 4: a, bicolor (Llano s.n.); B, stipulatum (Cuatr.
21339); c, subincanum (Little 9544); p, microcarpum (Archer 1551); £, obovatum (Klug
2983).

prominent, subspreading, near the margin arched, decurrent, anasto-
mosing, the transverse tertiary nerves prominent, those of the fourth
rank also prominent, reticulate-anastomosing, the lesser veins forming
a minute, prominulous reticulum, the minor reticulum and areoles
covered by a dense tomentum of intricate, white, sericeous, minute

538 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

stellate hairs, the major nerves with only scattered minute, slightly
larger hairs and abundant, minute reddish, callose warts; base of lamina
7-nerved, the 4 lower nerves small, of lower degree.

Fertile branches perennial on main trunk and big branches, short,
ligneous, tuberculate, prolific in flowering, the short, intricate, cin-

Ficure 36.—Fruits of Theobroma, X 16:"4, simiarum (Cuatr. 26515A); B, simiarum (Cuatr.
26536); c, chocoense (Patifio 115); p, cirmolinae (Cuatr. 15336); £, grandiflorum (Cuatr.
25780T); F, mammosum (Cuatr. 26535).

CUATRECASAS—CACAO AND ITS ALLIES 539

sv

Ficure 37.—Leaves of Theobroma: a, stipulatum (Cuatr. 21339); B, stipulatum, stipules and
base of leaf beneath (21339); c, cirmolinae (Cuatr. 15336); pv, angustifolium (Allen
6259); £, angustifolium (Stand. 22317), showing the stipules. All leaves X 4; 3B, X 36.

540 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

cinnate branches forming cushions up to 12 cm. broad on the trunk;
sympodial branchlets angulate, ramose, bracteolate, forming crowded
panicles up to 8 cm. long, appressed stellate-tomentose or glabrate
when old; bracts 1.5-3 mm. long, 1.5-2.5 mm. broad, subcoriaceous,
persistent, subamplectant, triangular-ovate, tomentose; peduncles
elongate in anthesis, erect, rather thin, ferruginous tomentose, 5-8
mm. long, 3-bracteolate at apex; bracteoles linear, subacute, 3-6 mm.
long, 1-1.5 mm. wide; pedicels in anthesis 15-20 mm. long, thin,
erect, tomentose, ebracteate; buds globose, with 5 commissural ribs,
densely ferruginous tomentose.

Sepals thick, carnose, ovate-triangular, acute, united in the lower
third or fourth, soon umbilicate-reflexed, with the five free lobes
spreading, 10-15 mm. long, 6-7 mm. broad, pale yellow inside,
glabrous, except for the base, with minute, oblong, glandular hairs
at their insertion, yellowish, thick-tomentose outside with ochraceous
or tawny stellate hairs, the margin minutely cinereous-tomentose.

Petal-hoods yellow, thick-membranaceous, elliptic-obovate, rounded-
cucullate at apex, glabrous, with 7 prominent nerves inside, 6-7 mm.
long, 4-5 mm. broad; petal-laminae thick, carnose, glabrous, sub-
triangular-spatulate, slightly 3-undulate or emarginate at apex, about
3.5 mm. long, 2.2 mm. broad, tapering to a pedicel at base; pedicel
about 4 mm. long, 0.6 mm. wide, bidentate at the joint.

Androecium tube thick, 2-2.2 mm. high; staminodes laminar,
rather thick, oblong-spatulate, slightly broadened towards the end,
retuse at apex, erect, glabrous, sulphur yellow, often somewhat
reddish near the base, 10-11 mm. long, 4-5 mm. broad, at base 1.5 mm.
wide; filaments robust, glabrous, 2.2-3 mm. long, shortly 3-furcate,
3-antheriferous, rarely 4-furcate, 4-antheriferous; ovary ovoid, 1.6-2
mm. long, 1.8 mm. broad, 5-costate, tomentose; styles 2 mm. long,
connivent, free up to near the base.

Fruiting peduncle robust, 3-4 cm. long, 8-10 mm. thick, articulate;
young fruits large, fusiform, prismatic with 5 obtuse, prominent ribs
corresponding to the loculi and 5 others, more or less marked, alternate
commissural ones, the base umbilicate, the apex attenuate and obtuse ;
ripe fruit 25-35 cm. long, 10-12 cm. wide, ellipsoid-oblong or obovoid-
oblong, very little narrowed to the umbilicate base, attenuate to the
obtuse apex, the surface obtusely pentagonal, with rounded ridges,
brown or ferruginous, stellate-tomentose; pericarp about 1-1.5 cm.
thick; epicarp 2 mm. thick, very hard, ligneous, the mesocarp and
endocarp carnose, becoming hard and coriaceous when dry; covering
of the seeds fibrous, pulpy, yellowish white and flavorous; seeds striped
from the more or less compressed pulp, ovoid, the testa brown, the
outer tegument light brown, the inner one dark brown, the cotyle-

CUATRECASAS—CACAO AND ITS ALLIES 541

dons reddish, 20-24 mm. long, 15-19 mm. broad, and 9-16 mm. thick;
germination hypogeous.

Common NAMES.—Bacao, cacao de monte, cacao indio.

Uses.—No special uses known besides occasional preparation of
chocolate by the natives. This species is the one which grows at the
highest altitudes; it should be tried as a grafting base, especially in the
coldest zone of cacao production.

DistrisuTion.—Only known from the Pacific slopes of the Andes
in Colombia in the Department of El Valle, between 800 and 1300

meters altitude.

COLOMBIA: Et Va.ue: Western slope of western Cordillera, valley of
Rio Digua, Piedra de Moler, rain forests, 900-1180 m. alt.; tree 20 m., trunk 40 em.
in diam., adult leaves thick-coriaceous, green above, yellowish green beneath,
sepals ferruginous-green, petals yellow, staminodes bright yellow, their bases
reddish, bark and central wood with resine, ‘“‘bacao,”’ 19 VIII 1943, Cuatrecasas
14897 (holotype VALLE; isotypes, F, Y). Valley of Rfo Digua, La Elsa, forests
1000-1200 m. alt.; tree 12 m., branched in the upper part, bark dark gray, almost
smooth, flowers yellow, 9 XI 1943, Cuatrecasas 15336 (F, VALLE, Y, paratypes).
La Elsa, about 800 m. alt.; tree 12-15 m., 30-35 cm. in diam. at base, jorquettes
arising symmetrically, flowers (yellow) and fruit on trunk and main branches,
mature fruiting peduncle 4 cm. long, 1 em. thick, pod 26-28 x 10-11 cm., bluntly
ridged, 23 VI 1953, Holliday 140 (COL, TRIN, US). La Elsa; tree 9-12 m.,
30 cm. in diam. at base, jorquettes arising symmetrically, flowers yellow, fruit on
trunk and main branches, 23 VI 1953, Holliday 139 (TRIN, US). Hoya del Rfo
Sanquininf, left side, La Laguna, forests 1250-1400 m. alt.; tree 20 m. alt., yellow
flowers, fruits large, brown tomentose, ‘‘cacao indio,’”’ 20 XII 1943, Cuatrecasas
15700 (F, VALLE, Y).

12. Theobroma stipulatum Cuatr. Ficures 30, 32, 35, 37; Map 9
Theobroma stipulatum Cuatr. Fieldiana Bot. 27(1):84, fig.7. 1950; Baker,
Cope, et al. (1954) 14, line 15 (as Theobroma sp.); Leén (1960) 322, 315,

fig.

Typr.—Cuatrecasas 21339, Colombia, Chocé.

Large tree to about 30 m. high; growth pseudoapical ; trunk about 45
cm. in diameter, somewhat triangular at base, the bark rugose,
granulate, reddish brown, the wood dark ochraceous, hard; branches
grayish, rugose-squamulose, glabrate, the primary ternate; terminal
branchlets pale ferruginous tomentose, densely covered by stellate or
fasciculate hairs; stipules coriaceous, densely tomentose, pale fer-
ruginous, ovate or ovate-oblong, obtuse, persistent, 8-12 mm. long,
5-9 mm. broad, the terminal up to 25 x 11 mm.

Leaves large, strongly coriaceous; petioles robust, very thick,
short, densely ferruginous tomentose, 5-10 mm. long; blades ovate-
elliptic or elliptic, more or less oblong, rounded, truncate or obtuse at
apex, the usually slightly asymmetrical base emarginate-cordate,
entire or slightly sinuate and flat at margin, 23-45 cm. long, 11-17 cm.
broad, green above, when dry brown or pale brown, slightly rugose or

542 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

almost smooth, with scattered, minute, stellate hairs, the costa and
secondary nerves linear, tomentose, depressed, greenish ochraceous
beneath or when dry ochraceous brown or pale brownish, tomentose,
covered with minute, whitish, intricate, stellate hairs, and other
mediocre, thicker, ferruginous, stellate hairs copiously covering the
veins, the costa very robust and prominent, the secondary nerves
about 12 on each side, very prominent, subascending, near the margin
curved, decurrent, and anastomosing, the transverse tertiary nerves
prominent, parallel, 5-15 mm. distant from each other, the minor
veins prominously reticulate.

Inflorescences cauline, on trunk or main branches, the fertile
branches perennial originating from tubercles; ligneous branches
short, tortuous, intricate, furcate-ramose (dichasial and cincinnate),
bracteate, tomentose, up to 1-3 cm. long; bracts ovate, subcoriaceous,
tomentulose, minute; peduncles solitary, 5-12 mm. long, thin, stellate-
tomentose, 3 bracteolate and 1-flowered at apex; bracteoles linear,
acute, 2-3 mm. long; pedicels erect, thin, densely tomentose, 10-22
mm. long; buds globose, 10-14 mm. in diameter, sublanate-tomentose;
sepals thick, ovate-triangular, glabrous and ochraceous inside, except
for the minute, thick, oblong, glandular hairs at base, densely stellate-
tomentose or sublanate outside, 10-15 mm. long, united in the lower
third, reflexed at anthesis, umbilicate at base; petals yellow, glabrous,
the hoods obovate elliptic, 7-nerved-sulcate, involute at margin,
round-cucullate at apex, about 5 mm. long and 3-4 mm. broad;
petal-laminae yellow, thick, 3-3.5 mm. long, ca. 3-3.5 mm. broad,
suborbicular or subspatulate, slightly retuse at apex, attenuate into a
narrowly linear pedicel at base, this 3.5-4 mm. long.

Androecium tube 1.5-2 mm. long, glabrous; staminodes petaloid,
yellow, thick, glabrous obovate-oblong, subspatulate, rounded at
apex 1-toothed on each side, 10-11 mm. long, 4.5-5 mm. broad, at
base 1.5 mm. broad; filaments rather thick, glabrous, 1.8-2 mm. long,
curved, shortly 3-furcate, 3-antheriferous; anther lobes ellipsoid;
ovary ovoid-oblong, 5-furrowed, densely tomentose-hirsute, about 2
mm. high; styles filiform, glabrous, coherent, about 2 mm. long.

Fruit 17-22 cm. long, 9-11 cm. broad, ovoid-ellipsoid or ellipsoid
and oblong, rounded at base, slightly attenuate, obtuse or rounded at
apex; pericarp hard-coriaceous, rigid, smooth, appressed brown
tomentose, the epicarp woody, about 1.5 mm. thick, the mesocarp
and endocarp carnose, creamy, about 1 cm. thick; seeds compressed
about 20-55, surrounded with pale, yellowish white, soft, scented
pulp, more or less amygdaliform, 20-25 mm. long, 18-21 mm. wide,
and 7-10 mm. thick, the testa subcoriaceous about 0.5 mm. thick;
cotyledons white; germination hypogeous; fruiting peduncles robust,
1-3 cm. long, about 1 cm. thick.

CUATRECASAS—CACAO AND ITS ALLIES 543

Common NAMES.—Chocolate de monte, cacao de monte.

Uszrs.—The seeds are said to yield a good chocolate but they are
only occasionally used by the natives.

DistTRIBUTION.—Restricted to the rain-forested basins of the rivers
San Juan and Atrato (Chocé region) and perhaps Rio Sequién (Narifio)
in western Colombia, where it is of rare occurrence.

COLOMBIA: Antioquia: Villa Arteaga, about 100 m. alt.; tree 12 m., 20 cm.,
diameter base, flowers borne on trunk and main branches, pedicel from base
to bracts 1.6 cm., from bracts to flower 2.2 cm., bracts 3, one usually larger,
abscission layer near bracts, sepals joined about 4 way from base, reflexed when
flower opens, all parts of flower yellow, 7 ridges on inside of petal, staminodes
spatulate, 14 mm. long, 6 mm. wide, 22 VII 1953 Holliday & Bartley T/165 (TRIN,
US). Ibidem, and same tree; Cope Ant. 2 (specimens lost), field annotations
by Cope: “A branched tree, showing the branching of the subincanum group, new
growth from above jorquette; 3 branches.” ‘Interior parts of bud examined
were creamy white; sepals reflexed in opened flower, 7 mm. long x 4 mm. wide,
elliptic, pointed, apparently rather soft and spongy, truncate margins, free almost
to base, inner surface glabrous; petals 5, free, with cup-shaped base and ligule,
base 4 x 4 mm., ligule and strap 5 mm. long, expanded portion spatula-shaped,
staminodes 5, reflexed, oblanceolate, 10 mm. x 3 mm., fruit somewhat ovate in
outline, in cross section, slightly flattened on 5 side, no ridges or furrows, densely
covered with medium brown, stellate hairs, which impart a mealiness to surface,
18 cm. long x 10 cm. diameter, wall with woody outer layer about 1.5 mm. thick,
inner surface very soft, creamy, about 1 cm. thick, 55 seeds embedded in soft,
cream-colored tissue, beans flattened, up to 1 cm. thick, somewhat triangular in
outline, up to 25 mm. long x 21 mm. wide, testa chocolate brown in color, about
0.5 mm. thick, rather leathery, cotyledons pure white, markedly convoluted, pulp
has smell of green bananas.”

Cuocé: Rfo San Juan, right margin of river on low hill near Palestina, about
30 m. altitude; tree 32 m. tall; trunk triangular at base, 45 cm. diam., branchless
up to 25 m. height, bark rugate-granulate reddish brown, wood rather dark
ochraceous, leaves coriaceous, rigid, green above, ochraceous green beneath, when
very young light yellowish green above, young branchlets and stipules pale
ferruginous, fruits oblong-ellipsoid or oblong-subovoid, obtuse, dark brown
tomentose, 17—22 cm. long by 9-10 cm. broad, pericarp rigid woody, ‘‘chocolate
de monte,”’ 28 V 1946,Cuatrecasas 21339 (holotype, F; isotypes, US, VALLE, Y).
Rio Atrato, Lloré; young tree 3 m., sterile, jorquette of three branches, 4 VIII
1953, Holliday & Bartley 175 (TRIN, US).

Narifio: Iscuandé, Rio Sequién, 100 m. alt.; tree 18 m., bark reddish and
smooth, wood cream colored, ‘chocolate,’ 23 XI 55, Romero Castateda 5500
(COL). (A sterile specimen, the identification is doubtful; flowers and fruits
necessary.)

13. Theobroma chocoense Cuatr., sp. nov. Ficurss 3, 31, 36, 38; Mar 9

Arbor 8-15 m., trunco erecto pseudoapicale crescente, cortice sub-
laevi, ramis ternatis robustis patulis superioribus ascendentibus ramulis
alternis juvenilibus dense crasseque lanuginoso-tomentosis viridi-
ochraceis vel viridi-ferrugineis, pilis stellatis intricatis mediocribus
ochraceis densis et pilis fasciculatis longioribus pallidis sparsis tectis,
denique glabratis brunneis nitidis angulosis cicatricosis; stipulae
coriaceae persistentes petiolis longiores ovatae obtusiusculae longi-

544 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

tudinaliter striato-nervatae primum ochraceo-tomentosae, pilis fer-
rugineis mediocribus stellatis, deinde cinereae, pilis stellatis minutis
albidis adpressis munitae, intus glabrae 12-22 mm. longae, 10-15
mm. latae, in ramis valde juvenilibus ovato-lanceolatae 2-25 mm.
longae, 8-9 mm. latae.

Folia alterna rigide coriacea; petiolus brevis crassus robustus viridi-
ochraceus vel viridi-ferrugineus dense lanuginoso-tomentosus, pilis
mediocribus stellatis densis et pilis fasciculatis longissimis (ad 2
mm. longis) pallidis sparsis, 7-14 mm. longus, 8-10 mm. crassus;
lamina oblongo-elliptica vel subovato-elliptica, integra vel levissime
undulata, basi ampla leviter attenuata subsymmetrica rotundata cor-
dato-emarginata, apicem versus paulo attenuata apice obtusa subite
acuteque acuminata 18-42 cm. longa, 9-20 cm. lata, acumine 1-1.5 cm.
longo, supra juventute ochraceo-stellato-tomentosa deinde glabra vel
sparsissimis pilis stellatis mediocribus munita, plus minusve venoso-
rugosa, nervis secundariis filiformibus impressis, nervulis venulisque
plus minusve impressis, superficie lutescenti-viridi in sicco brunne-
scente, subtus valde nervosa viridi-cinerea in sicco ochraceo-ferruginea,
costa crassa valde eminente, nervis secundariis valde prominentibus
12 vel 13 utroque latere paulo ascendentibus ad marginem arcuatis,
nervis tertiaris transversis parallelis prominentibus 3-10 mm. inter se
distantibus nervulis minoribus venulisque bene prominentibus reti-
culatis, indumento heterotricho-tomentoso, superficie alveolorum
venulisque reticuli pilis-stellatis minutissimis tenuibus albidis in-
tricatis dense tectis, nervis alteris copiosis punctis callosis rubris et
copiosis, pilis stellatis majoribus ferrugineis radiis patulis munitis,
in foliis vetustis nervis majoribus saepe glabratis, in prole foliis tenui-
oribus subtus albo-cinereis, pilis minutissimis stellatis albis dense
tecta, in nervulis sparsis in petiolo costa nervisque principalibus
densis pilis stellatis ochraceis radiis patulis gracilibus 1-1.5 mm.
longis basi calloso-rubescente praedita.

Folia prolis orthotropae tenuia petiolata late ovata vel rhomboideo-
ovata sursum crenato-dentata apice acuminata, circa 8 nervis secunda-
riis utroque latere; lamina 12-20 cm. longa, 8.5-12 cm. lata, petiolo
3.5-4 cm. longo, vel latiore; stipulae anguste lanceolatae.

Inflorescentiae in trunco et in ramis majoribus copiosae, tuberculis
lignosis persistentibus orientes, ramulis brevissimis cymosis tomentosis
densos glomerulos floriferos formantibus; bracteae lineares circa 5
mm. longae ferrugineo-tomentosae; pedunculi ad 1 cm. longi tomentosi
apice 3-bracteolati; bracteolae anguste lineares 2—2.5 mm. longae
tomentellae; pedicelli 9-14 mm. longi erecti mediocres stellato-
tomentosi viridi-ochracei; alabastra_ ellipsoideo-(globoso)-depressa
9-10 mm. lata, 7-8 mm. longa, in commissuris 5-costata, minute ochra-
ceo-tomentosa; calyx basi cupularis, intus basi ad marginem insertionis

CUATRECASAS—CACAO AND ITS ALLIES 545

annulo pilis densis crassis glandulosis praeditus, in anthesin reflexus;
sepala 5 crassuiscula ovato-acuta, 10-11 mm. longa, circa 7 mm.
lata, basi 2.5-3 mm. longe in tubum coalita, extus minute ochraceo-
stellato-tomentosa, margine minutissime albo-tomentosa, intus glabra
superne parce pilosula excepta.

Petala glabra, cucullo crassiusculo pallido obovato-suborbiculato
circa 6 mm. longo et lato, basi amplo apice subrotundato depresso
apiculo bidenticulato, venulis 7 extus paulo conspicuis intus promi-
nentibus; lamina crassa rigida rubra obovato-deltoidea apice sub-
truncata, 5-6 mm. longa, 5-6.5 mm. lata, basi attenuata et cum
pediculo circa 3.5 mm. longo articulata.

Androecii tubus crassiusculus glaber circa 1.5 mm. altus; stami-
nodia petaloidea crassiuscula glabra in anthesin erecta obovato-oblonga
apice rotundata margine integra basim versus gradatim attenuata,
13-15 mm. longa, 5-7 mm. lata, basi 3-3.2 mm. lata; stamina fila-
mentis crassiusculis circa 2 mm. longis recurvatis glabris, antheris 3,
lobis bilocularibus loculis ellipsoideis circa 0.6 mm. longis; ovarium
globosum 2.5 mm. diam. dense hirsuto-tomentosum; styli circa 2
mm. longi.

Fructus laevis ellipsoideo-ovoideus basi late rotundatus apice paulo
attenuatus obtusus, 19-20 cm. longus, 10-11.5 cm. latus, pericarpio
crasso duro, extus crasse brunneo-tomentosus, epicarpio circa 2
mm. crasso lignoso; semina ovoidea vel triangulari-ovoidea compressa,
20-23 mm. longa, 16-18 mm. lata, 11-12 mm. crassa.

Type in the U.S. National Herbarium, No. 2402158, collected on the
grounds of the Experimental Station ‘‘Agroforestal del Calima,”
Bajo Rio Calima, at about 35 m. altitude, in the Department El
Valle, Colombia, September 25, 1961, by J. Cuatrecasas and L.
Willard (No. 26074). Flowers from the same tree collected by H.
Guerrero A. (No. 26074) December, 1961. Paratype collected at
the same place with a dry mature fruit by V. M. Patifio (No. 115).

CoMMON NAMES.—Cacao de monte, cacao grande de monte, bacao
de monte.

Distrisution.—Restricted to the Chocéd region, in the heavy
forested valleys of the rivers San Juan and Atrato in western Colombia.
The records of 7’. simiarum from the Calima River by Baker, Cope &
al. (1954, 14) refer to this species.

Theobroma chocoense is closely related to T. simiarum, from which
it differs in its broadly ovoid or ellipsoid pods, in its elliptic or ovate
leaves (instead obovate), in the indument of the lower side of the
leaves, which is composed of two kinds of hairs (minute, entangled,
white, stellate hairs covering the surface and larger ones with long,
spreading rays on the nerves), by the short, ovate, obtuse stipules,
and shorter bracteoles. All these features indicate that Chocoan

546 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

species is distinct from the Costa Rican species. Although the fruit
of T. chocoense is very similar to that of T. stipulatum, the red color
of the flower and the different indument of the leaves readily dis-
tinguish these two species.

COLOMBIA: Eu Vauie: Baja Calima (Chocé region), Rfo Calima, Estacién
Agroforestal del Calima, 30-40 m. alt.; tree about 8 m. high, with apical growing
trunk almost smooth with few fertile tubercles, primary branches ternate, abun-
dantly inflorescence-tuberculate, leaves rigid, medium yellowish green above,
cinereous green beneath, “cacao de monte,” 25 IX 1961, J. Cuatrecasas & L.
Willard 26074 (holotype, US; isotype, COL). Ibidem; flowers purple red, XII
1961, Humberto Guerrero 26074 (US). Ibidem; small tree (about 10 m.) on the
edge of the small creek behind the Palmetum, dry fruit 19 x 11.5 em., I 1953,
V. M. Patifio 115. Ibidem; tree 12-15 m. in forest, fruit 16 x 11 and 20 x 10
cm., quite smooth, no flowers, 29 VI 1953, Holliday 144 (TRIN, US). Rfo
Calima, Cajfio la Brea; young tree 2 m., in forest, sterile, 29 VI 1953, Holliday 143
(TRIN, US).

Cuocé: Rfo San Juan, in front of Palestina, Quebrada de la Sierpe, 0-40 m.
alt.; tree about 14 m. high, trunk 30 cm. diam., leaves rigid, rugose, coriaceous,
deep green above, cinereous green beneath, strongly nerved, fruit ellipsoid,

smooth, velvety tomentose, 20 cm. long, 10 em. wide, the seeds acid, “bacao de
monte,” 13 III 1944, J. Cuatrecasas 16896 (F, VALLE, US).

13a. Theobroma chocoense var. bullatum Cuatr., var. nov.

A var. chocoensi foliis rugosis reticulato-bullatis differt. Lamina
rigida elliptico-oblonga vel paulo obovato-oblonga, magna, usque ad
50 x 21 cm., basi rotundata, apice rotundata breviter subiteque cuspi-
data, juvenilis basim versus plus minusve attenuata apice magis
angustato-acuminata; cucullum petalorum 5-6 mm. longum, 3-4 mm.
latum, pediculo 3.5-4.5 mm. longo; lamina triangulari-spathulata,
2.5-4 mm. longa, 2.5-3 mm. lata; staminodia obovato-oblonga apice
rotundata, circa 10 mm. longa, 4 mm. lata, basi 1.5 mm. lata; sepala
extus tomentosa intus glabra, circa 12 mm. longa, 5 mm. lata, 3 mm.
basi coalita.

Type in the U.S. National Herbarium, No. 2404636, collected in
Quebrada Juan Marfa, a small tributary of Rfo Juradé, at about 500
m. from its mouth, in the Municipality of Nuqui, Department of
Chocé, Colombia, at 50-100 m. altitude, on a hill in a region having a
dry season of about 4 months, 15 II 1955, V. Mf. Patifo 171.

Although at first sight the leaves of this variety look very different
from typical 7. chocoense, no other differences can be found on the
basis of the available material. According to fragmentary data it
seems that the fruit of the Juradé specimens are identical with those
of T. chocoense. The leaves are strongly nerved, reticulate and rugose-
bullate, but some specimens of typical T. chocoense also show rugosity
in the leaves. V.M. Patifio writes about his specimens: ‘Tree 12-14
m. alt., the trunk straight, triangular on the lower half, with sides

CUATRECASAS—CACAO AND ITS ALLIES 547

about 30 cm. broad, tubercular, floral cushions copious on the whole
trunk; the specimens are scattered; no other trees were seen in the
vicinity; the “cuzumbies”’ like its fruits very much, for which reason
it is difficult to find them unbroken; branches few, terminal; leaves
40-50 x 16-21 cm., petiole 1-1.5 cm., 6-11 mm. thick; leaves of very
young specimens dentate in their upper half; fruits on trunk and
branches; the 2 flowers collected were dry but a Choko Indian told
that its natural color is red and that the seeds are purplish; the Choko
name for the species is ‘‘cumaj6”’ or “‘judromajé6.”

To this variety belong the sterile specimens from Rfo Atrato referred
to by Baker, Cope and al. in their Report, page 14 as “Theobroma sp.
(possibly new).”’ Additional flowering and fruiting collections of
this plant are needed to complete our knowledge and to determine
better its taxonomic rank.

Common NAMES.—Chocolate de monte, bacaito de monte, cumajé
(Choké), judromajé (Chokd), according to Victor M. Patifio notes.

Uszs.—The pulp is acid and of pleasant taste, for which reason it is
sought by animals and also by the Indians.

DisTRIBUTION.—Rio Atrato and northwestern Pacific drainage,
Chocé (Colombia).

COLOMBIA: Cuoco: Rfo Atrato, Lloré, about 50 m. alt.; tree 12 m., 4 VIII
1953 Holliday & Bartley T/176 (K, TRIN, US). Municipality Nugquf, Rio Juradé,
Quebrada Juan Marfa, about 500 m. from mouth, 50-100 m. alt., 15 II 1955,
Patifio 171 (US). Ibidem, seedling, Patifio 171A (US); first branches of a
young plant, Patifio 171B (US).

14. Theobroma simiarum Donn. Smith. Fiaures 5, 6, 32, 36, 38, 40; Map 8
Theobroma simiarum Donn. Smith in Pittier, Prim. Fl. Costar. 2: 52. 1898;
Bot. Gaz. 25: 145. 1898; DeWildeman (1902) 97; Standley (1937) 689;
Chevalier (1946) 282; Holdridge (1950a) 2; Allen (1956) 343; Leén (1960)

322, 321, fig.

Typr.—Turrialba, Costa Rica, Tonduz 8373 (= 7313 distributed
J. Donnell Smith). Lectotype: US 1,382,332 [Photo F. M. 40723],
US 471,873 [Photo F. M. 40722]. Syntypes: Puttier & Tonduz
3925, Tonduz 6852, Cooper 10244.

Medium to large tree up to 20 m. tall, with erect, thick trunk
up to 60 cm. in diameter at the triangular base; growth pseudoapical ;
bark more or less rugose; branches spreading, the primary ternate,
the upper ones ascending; young branchlets thick-tomentose, greenish
ferruginous or brownish, covered by three kinds of hairs: 1) abundant
minute squamose stellate hairs with thin rays, 2) mediocre stellate,
longer hairs and 3) scattered large stellate or furcate hairs with very
long white rays, when older glabrate, brownish or grayish; stipules
persistent, longer than the petioles, subcoriaceous, firm, erect, lanceo-

548 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

late or linear, acute, 15-24 mm. long, 3-6 mm. broad, tomentose
outside.

Leaves large, rigid, coriaceous; petiole robust, thick, somewhat
striate, thick-tomentose-ferruginous, 5-12 mm. long, 6-10 mm. thick;
blades obovate-oblong, rounded or very obtuse and abruptly cuspidate
at apex, broad or slightly narrowed, rounded or slightly cordate and
asymmetrical at base, entire or sinuate-dentate in the upper third,
in very young plants obovate-rhomboid, long-acuminate, the upper
half or third repand, acutely dentate, the adult 20-40 cm. long, 8-17
em. broad, the acumen 0-1 cm. long, green above, when dry pale
brown, smooth, glabrous or with scattered stellate hairs on the midrib,
this and the secondary nerves filiform and depressed, the other veins
usually obsolete, light green beneath, when dry cinereous or whitish
but the nervation ferruginous, the costa thick, very prominent, the
9-11 secondary nerves on each side thick and prominent, spreading-
ascending, the lower pair usually at a more acute angle, the transverse
tertiary nerves prominent, separated 5-10 mm. from each other,
the minor nerves and veins prominulous, reticulate; indument hetero-
trichous, the areolar surface and the small reticular veins covered by
a dense tomentum of intricate, minute, white, stellate hairs, the
thicker nerves with sparse mediocre, ferruginous, stellate hairs and
reddish callous scars, the costa densely ferruginous by more or less
copious mediocre hairs and other ochraceous, long ones; in seedlings
the under leaf surface whitish, covered with a white tomentum of
extremely minute, white, stellate hairs, and on the nerves sparse and
on the costa dense ferruginous stellate hairs with long (1 mm.), thin,
patulous rays and red, callous bases.

Inflorescences borne on tubercular protuberances on the trunk;
cymose branches usually cincinnate, ligneous, very short, forming
dense, many-flowered glomerules; branchlets up to 4 cm. long, tor-
tuous, rugose, glabrate, bracteate, the bracts coriaceous, ovate,
subacute or acuminate, tomentulose outside, 1-2 mm. long, 1-1.5 mm.
broad, persistent; hornotinous branchlets short, crowded, ferruginous-
tomentose, covered by fertile, tomentose, imbricate bracts; pe-
duncles axillary, mediocre, densely stellate-pilose-tomentose, 5-15
mm. long, 3-bracteolate at apex, the bracteoles narrow-linear, to-
mentose outside, 3-7 mm. long; pedicels up to 15 mm. long; buds
globose, densely tomentose, above almost 5-angulate; calyx cupular
at center, subglabrous, with a ring of minute, thick, oblong, glandular
hairs at base and with very thin hairs above inside, densely and
thickly stellate-tomentose and ochraceous green or ferruginous outside,
the sepals ovate-acute, about 10-12 mm. long, 6 mm. broad, united
about 2 mm. at base, reflexed at anthesis; petals red, thick, glabrous;
hood yellowish white, obovate-oblong, with 7 prominent nerves inside,

CUATRECASAS—CACAO AND ITS ALLIES 549

Ficure 38.—Leaves of Theobroma: a, simiarum, from orthotropic stem (Stand. 37377);
B, simiarum, from normal lateral branches (Cooper 10244); c, chocoense (Cuatr. 26074);
p, leaf base and stipules of 7. chocoense (Cuatr. 26074). All leaves X 14; D, X %.

550 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

narrowed-unguiculate, and inside reddish at base, 6-8 mm. long, 4—5
mm. wide, rounded-cucullate, inflexed-apiculate at apex, the apiculum
bifid, articulate to the pedicellate lamina; petal-lamina obtrapezoid,
carnose, red, subtruncate at apex, cuneate at base, 3.5-4.6 mm. long,
3.5-5 mm. wide, reflexed in bud, erect in anthesis; pedicel linear,
4-5 mm. long, 1 mm. broad; androecium tube about 2 mm. long,
rather thick, glabrous; staminodes petaloid red, glabrous, obovate-
oblong, rounded at apex, slightly sinuate-dentate in the upper third
or subentire, 11-13 mm. long, 5-7 mm. broad, 1-2 mm. wide at base,
reflexed in bud, erect at anthesis; filaments rather thick, glabrous,
curved, 2 mm. long, shortly 3-furcate, 3-antheriferous; anther lobes
ellipsoid, 0.6-0.7 mm. long; ovary obovate, densely hirsute-tomentose,
about 1.2 mm. high; styles about 1.2 mm. long, thin, acute, united
at base.

Fruiting peduncle short, thick, usually 1-1.5 cm. long; fruit ellip-
soid-oblong, smooth, truncate or very blunt, rarely narrowed at
base, rounded, very obtuse or slightly attenuate at apex, 16-35 cm.
long, 6-10 cm. broad, covered by dense and thick, ferruginous or
brown tomentum; pericarp 11-15 mm. thick with: 1) epicarp woody,
1,5-2 mm. thick, with a tomentose epidermis, 2) mesocarp 5-10 mm.
thick, firmly carnose, pale ochraceous whitish, and 3) endocarp 2-5
mm. thick, softer carnose; seeds 36-60, distributed in 5 more or less
double rows, each one covered by a thick, fibrose, white or rosy-
white, aromatic pulp irregularly ovoid, 20-21 mm. long, 18-17 mm.
broad, and 13-15 mm, thick; cotyledons white; germination hypogeous.

CoMMON NAMES.—Cacao de mico, cacao de mono, teta negra.
Indian names: Kréaku (Guatuso), Nunfsup (Rama), Uirub (Bribrt),
Dzug-mang-ua (Brunka), Ku-gin, Bik (Térraba), according to
Standley, loc. cit., Uir-ub (Bribri) according Pittier.

Uses.—The seeds are said to give a cacao of good quality. They
are occasionally used.

DistripuTion.—Limited to Costa Rica. Infrequently found in
preserved or remnant forests in both Atlantic and Pacific lowlands,
from sea level to 600 m., and exceptionally in higher altitudes up to
900 m.; recorded from the provinces of Limén, Heredia, Puntarenas,
and Cartago. It may extend to Nicaragua and northern Panama,
but no records from these two countries exist at present in herbaria.

In its natural habitat T. simiarum attains a considerable height
and thickness. The trunk is erect and branchless up to several meters.
The inflorescences may appear on the trunk but they are more abun-
dant on the big branches. The fruit is very characteristic because of
its terete, sausage shape. From the closely related Colombian species
L’. stipulatum and T. chocoense it can be distinguished by the elongate

CUATRECASAS—CACAO AND ITS ALLIES 551

shape of the pod; in the two other species the fruit is ovoid or broadly
ellipsoid. From TY. stipulatum it differs also by its red flowers, longer
lanceolate stipules, longer bracteoles, and by the indument of the
leaves. From T. chocoense it differs also by its narrower and longer
stipules, the obovate form of the leaves, by the leaves having beneath
usually three kinds of hairs, and by its longer bracteoles.

COSTA RICA: Carraco: Turrialba, grassland, 570-600 m. alt., XI 1893,
Tonduz 8373 (distributed by Donn. Smith under number 7313) lectotype US;
isolectotypes BR, G, GH, K; Photos F. M. 30722, 40723, 40743. Turrialba, forest
at margin of Rio Reventazén, about 600 m. alt.; tall tree with unbranched trunk
in the lower half, 60 cm. diam. near the subtriangular base, abundant tuberculate
inflorescences on the upper part and big ternate branches, few sausage-shaped
pods, 6 XI 1961, J. Cuatrecasas & Leén 26515A (US). Turrialba, Instituto Inter-
americano C.A., 600 m. alt.; flowers bright red, cult., 4 XI 61, J. Cuatrecasas & J.
Leén 26515 (US). Turrialba, I.I.C.A., 600 m. alt., cult., 14 III 51, J. Leén
3189 (TURRI). Tuis, forests, 670 m. alt., “cacao de mico,’’ XI 1900, Pitter
14016 (Donn. Smith 7731) (GH, K, US). Tucurrique, grassland and forests
around Las Vueltas, about 635 m. alt.; large tree, fruits brown, oval-elongated
30-40 cm. long, XI 1898, Tonduz 12822 (BM, G, LE, P, US), 18222 (M) [erroneous
number].

Limon: La Colombiana Farm of the United Fruit Company, about 70 m.,
wet forest; tree 60 feet or more tall with trunk 2 ft. thick, small crown, flowers
bright red in bunches on trunk, fruit sausage-shaped, hairy, 1 foot long or more,
said to be rare here, seeds give good cacao, 6, 7 III 1924, Standley 36822 (US). 28
miles on the railroad from Puerto Limén, towards Rio Barbilla, in marginal
forest, about 60 m. alt.; tall tree, flowers flame red, 12 V 1930, Cufodontis 599
(WU). Along Rio Reventazén, below farmhouse Finca Castilla, 30 m. alt., in
Gynerium sagittatum thickets, 27 VII 1936, Dodge & Goerger 9420 (MO). Palm
swamp between Rio Reventazén and Rio Parismina, on Castilla Farm, 2 IV
1930, Dodge & Nevermann 7164 (MO). La Lola, farm of the I.I.C.A., about 40
m. alt., cult.; trees about 10 m. high, trunk 20-30 cm. diam., triangular at base,
flowers scarlet, fruits oblong 15.5 x 7 cm., 18 x 8.7, 19.5 x 8.5, 22.7 x 7.7, 23.3 x
8.4, 25.7 x 9.3 em., with thick brown tomentum, 6 XI 1961, Cuatrecasas & Paredes
26536 (US). Vicinity of Gudpiles, 300-500 m., seedling, III 1924, Standley 37377
(US).

Herepia: La Concepcién, Llanuras de Santa Clara, 250 m. alt., “cacao de
mico,” II 1896, Donn. Smith 6457 (BM, US). Santa Clara; 6-7 m. high tree,
“cacao de mono,” IX 1896, Cooper 10244 (syntype, US) (Photo F. M. 40724).

PuNnTARENAS: Terraba, 260 m. alt., forests; caulinar flowers, large cylindrical
fruits (30-35 by 10 cm.), ‘cacao de mico,” II 1891, Pittier & Tonduz 3925
(syntype, BR). Boruca, forests 466 m. alt., III 1892, Tonduz 6852 (syntype, US).

TRINIDAD (cult.): Imperial College of Tropical Agriculture, Cuatrecasas &
Cope 25792 (US), 25794 (US).

SURINAM (cult.): Paramaribo, culture garden; tree 10 m., 15 cm. diam.,
almost horizontal branches, flowers bursting in dense clusters from the trunk
and old branches, calyx rusty brown, ligula and staminodes shining lacquer red,
8 IV 1954, Lindeman 5725 (U). Ibidem; seeds round, cotyledons white, germina-
tion hypogaeic, VIII 1956, van Suchtelen, s.n. (US).

BRASIL (cult.): Pard, cultivated and said to be introduced from Venezuela;
small tree with purplish flowers, II V 1957, Pires 6575 (IAN).

680-695—64——_12

552 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

15. Theobroma grandiflorum (Willd. ex Spreng.) Schum.
Fiaures 3, 5, 6, 7, 33, 36, 39, 40; Map 10; Pars 8

Theobroma grandiflorum (Willd. ex Spreng.) Schum. in Mart. Fl. Bras.
12(3):76, pl. 8, 1886; Jumelle (1899) 28, figs. 14, 15; DeWildeman
(1902) 95; Ducke (1925) 131; Ducke (1940) 272, pl. 4, fig. 2; Chevalier
(1946) 281; Addison & Tavares (1951) 25, pl. 1, fig. 1, pl. 2, fig. A, pl. 14,
fig. 10; Ducke (1953) 11; Baker, Cope & al. (1954) 13, fig. 14; Cuatrecasas
(1956) 656; Leén (1960) 320, 319, fig.

Bubroma grandiflorum Willd. ex Spreng. Syst. Veg. 3:332. 1826.

Guazuma grandiflora (Spreng.) Don, Hist. Dichl. 1:523. 1831.

Theobroma speciosum Willd.?, sensu Mart. in Buchn. Repert. Pharm. 35:24.
1830; Linnaea Litt. Bericht 32. 1831, non Willd.

Theobroma macrantha Bernoulli, Uebers. Art. Theobroma 11. 1869.

Theobroma silvestre Spruce ex Schum. in Mart. Fl. Bras. 12(3):76. 1886,
as synonym.

Typr.—Siber 4, “Hoffmannseg” in Herbarium Willdenow No.
14352 (B). Spruce 1822 (syntype of 7. macrantha Bernoulli and
neotype of B. grandiflorum Willd.).

Medium-sized tree, usually 6-10 m. high, reaching up to 18 m.;
growth pseudoapical; trunk up to 25-30 cm. in diameter, the bark
grayish, granulose, more or less wrinkled, internally rosy or reddish,
the wood pale; branches robust, spreading, the superior ascending, the
primary ternate; branchlets terete, the terminal rather thick, densely
and abundantly lanate-tomentose, the indument ferruginous, floccose,

$9" 49°

io”

o*

Map 10.—Geographical distribution of Theobroma obovatum @ and T. grandiflorum ©.

503

CACAO AND ITS ALLIES

CUATRECASAS

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Ficure 39.—Leaves of Theobroma, X ¥%: a, obovatum (Poeppig s.n.); B, grandiflorum (Cuatr.

257801); c, grandiflorum (Killip & Smith 30011); p, hylaeum (Araque & Barkley 18C745);

E, subincanum (Baker 38).

554 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

more or less deciduous, the hairs stellate or fasciculate with long,
intricate rays, caducous, the surface fulvous, with exfoliating rhyti-
dome, becoming gray, glabrous, and reticulate-rimose; stipules sub-
coriaceous, rigid, oblong or lanceolate-oblong, obtuse or subacute,
nervate-striate, abundantly lanate-tomentose, the tomentum more or
less deciduous, 10-20 mm. long, 3-6 mm. broad, persistent.

Leaves firmly coriaceous, medium sized or large; petiole thick, terete,
with a dense, thick, ferruginous, lanate tomentum, 7-14 mm. long,
2-5 mm. thick; blades oblong, subobovate-oblong or subelliptic-
oblong, more or less attenuate toward the base, this obtuse, rounded,
usually emarginate, or subcordate and slightly irregular, abruptly
attenuate and acutely acuminate at apex, the margin entire or slightly
dentate-sinuate toward the apex, 20-35 (15-60) em. long, 6-11 (5-16)
cm. broad, the acumen 1-2.5 cm. long, glabrous above, green, more
or less shining, brownish olivaceous or tabacine when dry, the midrib
and secondary nerves filiform, depressed, the tertiary ones slightly
marked or obsolete, greenish cinereous, glaucescent, or pale rosy
beneath, the costa very prominent, the 9 or 10 pairs of parallel second-
ary nerves prominent, subascending, near the margin thinner, curved,
anastomosing, the lowest pair usually forming a more acute angle
and more distally separated from the next, the filiform tertiary nerves
prominent, transverse, parallel, the minor nerves and veins forming
a fine prominulous reticulum, the costa, secondary, and tertiary nerves
glabrous, nitidous, with sparse reddish, callose dots, the reticulum
and areoles minutely tomentose, covered by white, minute, intricate,
dense, stellate hairs.

Inflorescences small, axillary and extra-axillary on leafy branches,
the short cymes reduced to 3-5 flowers or fewer, the branchlets
extremely short, ferruginous tomentose, the peduncles robust, 2-5
mm. long, 3-bracteolate at apex, the bracteoles narrowly linear,
tomentose, 3-4 mm. long; pedicels robust, rather thick, tomentose,
ebracteate, 5-20 mm. long.

Calyx subcymbiform; sepals firm, thick, carnose, ovate-oblong,
subacute, about 14-15 mm. long, 6-8 mm. broad, 1.5 mm. thick,
united in the lower third, but often the five separated to near the base
in two pairs and one free (25+25+S), the margin involute, the apex
minutely inflexed, thickly stellate-tomentose outside, ochraceous
green or ferruginous, rosy or reddish inside, shining, minutely,
sparsely, whitish pubescent, at base with minute, thick glandular
trichomes, the margin densely and minutely whitish tomentose.

Petal-hoods thick, carnose, whitish or yellowish, often with red
lines, obovate, rounded-cucullate at apex, 7-nerved, rugulose without,
pubescent, glabrous within, near the base reddish, 6-7 mm. long, 4-6
mm. broad; petal-lamina dark red or crimson, pedicellate, thick,

CUATRECASAS—CACAO AND ITS ALLIES 555

carnose-coriaceous, trapezoid-elliptical, more or less truncate or slightly
retuse, obcordate, minutely puberulous, 4-9 mm. long, 4.5-8.5 mm.
wide, abruptly contracted at base into a 4.5-7 mm. long, 1.5 mm.
broad pedicel.

Androecium tube 2.5 mm. high, sparsely pilose; staminodes reflexed
in bud, spreading in anthesis, crimson, dark red or purplish red,
lanceolate, very acute, thick, but somewhat flattened with a marked
midrib, 9-15 mm. long, 2-2.5 mm. broad throughout, pilose, especially
outside, the abundant hairs rather long, thin, flexuous; filaments
1.7-2 mm. long, thick, pilosulous, very shortly 3-furcate, 3-anther-
iferous; lobes of the anthers broadly elliptic, 1 mm. long; ovary
pentagonous-obovate, densely whitish hirsute-tomentose; styles 2
mm. long, connivent, free to the base.

Fruits falling from tree at maturity, without peduncle, densely
covered with a brown tomentum, large, smooth, ellipsoid or obovoid-
ellipsoid, rounded at both ends or, rarely, slightly attenuate at apex,
umbilicate and conically excavate at base, 16x10—-25x12 cm.; pericarp
about 1 em. thick, with: 1) epicarp hard, woody, about 2 mm. thick,
covered with a thin, tomentose epidermis, 2) mesoendocarp about 7
mm. thick, softly carnose at maturity with a thin, but firm, inner
pellicle limiting the seed cavity; seeds about 50, 5-seriate, each
surrounded by a yellowish, acidulous and distinctively scented, fibrous
pulp, the inner tegument a delicate pellicle, the testa subcoriaceous,
light brown, striped from the remains of the pulp, ovoid, or ellipsoid-
ovoid, more or less flattened, 20-30 mm. long, 20-25 mm. broad,
10-12 mm. thick; embryo white marbled, 19-23 mm. long, 16-20 mm.
broad, 9-12 mm. thick; cotyledons white; germination hypogeous.

The type of Bubroma grandiflorum Willd. ex Sprengel is the specimen
No. 14352 of the Willdenow Herbarium in Berlin, received from the
Hoffmannsegg Herbarium, collected in Brazil by Siber. This species
was transferred to Guazuma by G. Don who probably did not see the
plant, his description being taken from Sprengel. Schumann found
out that B. grandiflorum is synonymous with T. macranthum described
by Bernoulli many years later. The short diagnosis given by Sprengel
agrees with 7’. macranthum and the description of the leaves (‘‘amplis
oblongis abrupte acuminatis integerrimis”’) disagrees with any known
species of Guazuma, which always have serrate leaves. Freitag, who
listed Bubroma grandiflorum as synonym of Guazuma wulmifolia
(p. 216), did not see the type. In 1952, Dr. Mildbraed of the Berlin
Herbarium wrote me about this type specimen: ‘‘Im Herbar Willdenow
fehlte unerklirlicherweise der Bogen 14352 mit Bubroma grandiflorum.”
Schumann had seen the type in the Willdenow Herbarium, for which
reason I accept his judgment as final regarding the synonymy
established by him. Accordingly, I propose Spruce 1822 as a neotype

556 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

of B. grandiflorum Willd. ex Spreng., as it is the first collection cited
by Schumann and likewise a syntype of 7. macranthum Bernoulli.

ComMMOoN NAMES.—Cupasst, with some variations in the spelling or
pronunciation throughout its area of cultivation, cupuacu, cupt-asst,
copuassii, cupai-aci. Ducke quoted cupti do matto for wild speci-
mens in the middle Tapajoz River. The Anglo Colombian Cocoa Ex-
pedition recorded the following indigenous names (Baker, 1952):
Win-cheék-chéo-ai (Puinave), Infrida-Guaviare; bawk-pom (Maku),
Piraparand, Taraira; maga (Barasana), Upper Piraparand; fiee-aw
(Tanimuka), Guacaya; cupu-uassi (Brazil-Portug.); ba-dja-na-hoo
(Makuna), Lower Piraparana.

Usus.—The natives like to eat the acid and agreeably scented pulp
which covers the seeds, for which reason cupuassti is very much culti-
vated or planted in the state of Para and the eastern section of Ama-
zonas. ‘This pulp is used to prepare soft drinks (vinho do cupuassti)
and different kinds of preserves and candy which are exported from
Para and Maranhio. The taste of the pulp is sui generis.
L. Williams compares it with that of guand&bana. Patifio found the
odor of it similar to that of the “mate” (Crescentia cujete) when be-
ginning its fermentation. The fruits are very much liked by animals,
especially by monkeys (macacos), which very often empty the pods
to sip the pulp, contributing to the dissemination of the seeds.

DistrisutTion.—The known natural area of 7. grandiflorum is
the southern half of state of Pard, Brazil, and adjacent Amazonian
Maranhao. It has been found wild only in the rain-forest, on ele-
vated ground in the middle Tapajéz River region (waterfalls of Man-
gabal and of Itapacurdi, Ducke), Tocantins River (railroad of Alcobaca,
Ducke), Guamé River between Ourem and Braganca (Huber), Xingt
River between Victoria and Altamira (Ducke), and Anapt River
(LeCointe). The tree is always scarce in its natural area.

This medium-sized tree with large leaves, the largest: flowers in the
genus, and the largest pods among the Brazilian cacaos, is frequently
planted or cultivated throughout the states of Paré, Maranhao, and
the eastern part of Amazonas to Manaos. It is also occasionally
planted outside Brazil in warm lowlands of other tropical American
countries, such as Colombia, Venezuela, Ecuador, and Costa Rica.
It is found also in tropical botanical and agricultural gardens.

MARTINIQUE: From seeds from Cayenne, L. C. Richard, s.n. (P).

BRITISH GUIANA: “Herbarium Benthamianum,” Schomburgk, s.n. (K).

VENEZUELA: Amazonas: Capihuara, Alto Casiquiare, 120 m. alt.; culti-
vated, small tree up to 8-10 m., subrounded crown, trunk 25 cm. diam., gray
bark, inner rose or reddish, wood pale, pulp of fruit used to prepare beverages

with similar taste to the guandbana, “cupuast,” 28 V 1942, Ll. Williams 15615
(F, US, VEN).

CUATRECASAS—CACAO AND ITS ALLIES 557

COLOMBIA: Vaupés: Monfort; tree 6 m., sepals fleshy, petals maroon,
23 IX 1943, P. Allen 3105 (COL, MO, US). Rfo Vaupés, opposite confluence
with Rfo Papurf, Yavaraté, Salesian Mission Sio Miguel; tree about 3 years old,
growing in full sunlight, beginning to flower, no fruit, 20 II 1952, Bartley &
Holliday T-46 (COL, US). Rio Piraparan4, near confluence with Rfo Apaporis,
river level; young tree 3-4 m., cultivated in Indian garden, 24 VIII 1952, Baker
& Cope 5 (TRIN).

Amazonas: Rfo Caquet&, La Pedrera, river level; cultivated tree, rather
exposed, in the garden of the Orfanatorio, 19 IX 1952, R. H. D. Baker 16 (COL, F,
TRIN, US). La Pedrera, cultivated, highland; bushy tree 12 ft. tall, petals
purple red, calyx light golden brown, staminodes yellowish, 7 X 1952, Schultes &
Cabrera 17781 (US). Rfo Ricapuya, tributary of Rio Apaporis, river level; 2-3
years, young cultivated tree in Indian garden, 25 VIII 1952, Baker & Cope 6
(COL, F, TRIN, US). Leticia, 22 VIII 1946, Black & Schultes 46-61 (AMES,
F, IAN, NY, U, VEN); tree 8 m., “cupu-asst,” cult., 24 IX 1946, Black &
Schultes 46-III (IAN). Trapecio Amazénico, Amazon River, Leticia, 100 m.
alt.; cultivated, ‘“cupuassi,” IX 1946, Schultes 8178 (AMES, F, US).

BRAZIL: ‘Catal. Geogr. Pl. Bras. Trop.,”’ Burchell 9467 (GH, K, P). Ibidem,
Burchell 9375 (syntype of Theobroma macrantha Bernoulli, K). ‘Amazon region,”’
H. A. Wickham, s.n. (K). ‘Brazil Cameta,” Herb. Hanbury 9471 (K).

Amazonas: Rio Negro, Sao Gabriel; arvore 8 m., planta antiga dos sitios,
“cupu-assu,’”’ 27 XII 1945, Frées 21556 (IAN, K, NY, USDA). Upper Rio Negro
basin, Mouth of Rio Xie, cultivated; small tree, staminodes and ligules deep red,
rest of flower pink, ‘‘cupu-uassu,’”’ 29 XI-7 XII 1947, Schultes & Lépez 9204
(AMES, F, IAN, US). Rio Negro, prope Barra; ‘shrub 12-15 ft., flowers only
on ramuli, solitary normally ascendant, calyx pinkish within, petals crimson,
cucullate, bases yellow white, coronal scales red’; the Munich specimen bears
the number 83 on a mounting tape (F. M. Photo 40705), Oct. 1851, Spruce 1822
(neotype of B. grandiflorum, M; isosyntypes of T. macrantha Bernoulli, BM,
E, G, GH, K, LE, LD, NY, OXF, P, WU). Manaos, Horto Experimental, 20
m., “cupai-agi,” 19 XII 1923, Luetzelburg 22007 (M, NY, WU). Manaos, Agri-
cultural Experiment Station, 25 m. alt., cultivated; tree 35-40 ft., inflorescences
on main trunk, sepals green without, pink within, petals red, 13 X 1929, Killip &
Smith 30011 (NY, US). Manaos, Estrada do Aleixo, firm land; tree 6 m., flower
red, edible fruit, ‘“cupuagd,’ 16 IX 1955, Francisco (INPA) 1966 (IAN).
Manaos, Hacienda Brasil, 15 m. alt., “‘cupu-agd,” Luetzelburg 23287 (M). Ma-
naos, VIII 1906, ‘‘cupu-assti,” Labroy, (P). Three days upstream from Manaos,
300 ft. alt.; grown in semishade; flowers cream colored or dark crimson, grow-
ing out of the bark, faintly scented, Sandeman 2333 (K). Parandé de Matitins,
Rio Putumayo, Iga, between Tarapacd and its mouth (Santo Antonio do Iga,
100 m. alt.); small treelet; calyx very fleshy, green yellow; staminodes white,
laciniae flesh red turning brown on fertilization, “cupuassd,” 11-18 IX 1946,
Schultes & Black 8146 (F, IAN). Municipality of Mahacapuru, Solinos River
region, terra firma, Lago do Italiano; tree 25 ft. high, 4 inches diam., ‘‘cupu-
asst,’ Krukoff 1274 (A, BM, G, K, MICH, MO, NY, P, 8). Tefé; tree 7 m.,
flower brownish rose, cult. ‘‘cupuaci,”’ Black 47-1502 (IAN).

MaranuaAo: Turyassu, Igapo-wald; Baum 6-15m., Krone gelblich, filaments
rot-violett, 31 X 1923, Snethlage 300 (F, GH, US).

Acre (Territorio Federal): Brasilea, in a house in front of the Bolivian town
Cobijia, 150 m. alt., “‘copuasst,”’ cultivated, 5 IX 1954, Patio 163 (GH).

Park: Belém, northeast woods of the Instituto Agronémico do Norte;
infected with witches broom, ‘“‘cupu-assu,’’ 30 X 1942, Archer 7734 (IAN, NY,
US). Belém, Botanic Garden of the Museu Goeldi; native of the lower Amazon

558 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

country; the tree becomes much larger than the cacao and the top is relatively
narrower; the fruit resembles the pod of cacao but is much larger and the pulp
surrounding the seeds is most delicious; Dr. Huber speaks of it as the most im-
portant native fruit of Pard; “cupuagi,”’ 25 VI 1908, C. F. Baker 62 (A, C, E,
GH, LE, MICH, MO, P, U, US, NY). Museu Goeldi, 23 XI 1945, Pires & Black
744 (IAN). Jardim Botdnico do Museu Goeldi; large tree, calyx with red marks
inside base, spatulate portion petal dark red, fruit edible, ‘‘cupuacd,” 11 VIII
1942, Archer 7549 (F, IAN, K, US). Ibidem; small tree, leaves gray both sur-
faces, flowers large, cauliflorous, calyx white yellow, staminodes deep red, petals
pinkish, very showy, a profusion of flowers, cultivated, 15 VII 1946, Schultes
8065 (AMES, US). Horto Botdnico do Pard, XI 1903, ‘“cupuassi,” Huber 4008
(BM, G). Ibidem, ‘“‘cupuasst,’’ XI 1903, Siquetros 4008 (MG). Belém, “in
urbe cultum; arbor parva floribus brunnescentibus centro albido,” ‘“cupuasst,”
16 X 1940, Ducke 598 (F, IAN, MG, MO, NY, US). Ibidem, III V 1929, DaAl-
gren & Sella 438, 634 (F), “cupu-assu,” Dahlgren & Sella 733, 739 (F, GH, US).
Vicinity of Pard, 1 VII 1908, C. F. Baker 421 (BM, C, F, L, U, WU). In sylva
prope Barba et alibi, VIII 1828, Riedel 1373 (A, LE, S$, US). ‘Brasilia, Borbar,”’
Riedel, sn. (OXF). Boa Vista, on the Tapajés River, Aramanahy River,
“cupuassu,”’ Monteiro de Costa 121 (F). Taperinha bei Santarem, kultiviert;
aus dem Fructen wird Marmelade bereitet, kleiner Baum, ‘‘Cupu-asst,’”’ 18 IV
1927, Ginzberger & Zerner 800 (F, WU). “Habitat in sylvis udis umbrosis ad
Para, Dr. Martius Iter Brasil. Jul. 323,’’ Martius [874] (M). ‘Prov. Paraensis
ad Para, Dr. Martius Iter Brasil. 323,’’ Martius [875], [876] (M) (Photo F. M.
40706). Ibidem, Martius [873] (M) mixed with 7. guianense in Photo F. M.
19641). Pard, “Bresil-Martius,” Martius (G, P). Tapana, near Pard, woods;
30-40 ft. tall, appendages purplish red, clearing, fruit edible, 29 X 1929, Killip
& Smith 30320 (NY, US). Forét des collines du Mangabal, moyen Tapajés,
Cachoeira do Mangabal, “cupi do matto,” 5 IX 1916, Ducke 16458 (BM, G,
MG, P).

Rio DE JANEIRO: Rio Janeiro, cultivated, “cupti assu,’’ Glaziou 9643 (C, P).

PERU: Lorero: Caballo Cocha, on the Amazon River, ‘“cupuassut,” VIII
1929, Ll. Williams 2401 (F).

TRINIDAD: St. Augustine, Imperial College of Tropical Agriculture, River
State Diego Martinez, Field 2; trunk 20 cm. diam. at base, leaves coriaceous
dark green above, green beneath, fruits ellipsoid, densely tomentose, brown
ferruginous, 11 x 17, 11.5 x 18.5, 11.5 x 16.5, 12 x 19, 12 x 19.5, 12.5 x19 em., with
47-50 seeds, pulp yellowish with special scent, cultivated, brought from Belém
do Pard, 31 VIII 1961, Cuatrecasas, Cope, & Bartley 25780 T (US). Ibidem; tree
with larger and slightly attenuate fruits at apex, 10.7 x 22, 11.1 x 21.5, 11.8 x 25
cm., 31 VIII 1961, Cuatrecasas, Cope, & Bartley 25781 T (US). Saman Plot, culti-
vated from seeds brought from Pard; trunk 30 cm. diam., the diameter reducing
progressively upwards from one cluster of branches to the next, primary branching
ternate, from the base, apical growth, bark greenish brown, granulate-lenticellate,
minutely rimose, sepals 1.5-2 mm. thick, greenish ochraceous outside, whitish
and pink at base inside, hoods white yellowish or dirty whitish, the margin and
the base inside purplish, lamina carnose, dark red, later red brown, staminodes
star-patulous, red purplish becoming red brownish, styles united whitish, ovary
white, hirsute-tomentose, 2 IX 1961, Cuatrecasas & Cope 25801 (US).

CUATRECASAS—CACAO AND ITS ALLIES 559

16. Theobroma obovatum Klotzsch ex Bernoulli
FiauRes 33, 35, 39, 40; Map 10
Theobroma obovatum Klotsch ex Bernoulli, Uebers. Art. Theobroma 14, pl.
7, fig. 8. 1869; Ducke (1935) 132; (1940) 271, pl. 6, fig. 1; Chevalier (1946)
280; Addison & Tavares (1951) 25, pl. 1, fig. 2, pl. 2, fig. B, pl. 3, fig. 2,
pl. 4, fig. B, pl. 11, fig. 1; Ducke (1953) 10; Cuatrecasas (1956) 657; Baker,
Cope & al. (1954) 12, fig. 15; Leén (1960) 322, 319, fig.
Theobroma sylvestre sensu Huber, Bull. Herb. Boiss. II, 6: 273. 1906, non
Mart.

Typs.—Maynas, Peru, Poeppig.

Rather small tree up to 15 m. high; growth pseudoapical; trunk 10-
25 cm. in diameter; primary branches ternate, the inferior horizontal,
the upper ascending; terminal branchlets slender, when young
ochraceo-ferruginous, densely lanate-tomentose, the intricate, long,
stellate hairs more or less floccose, deciduous, when older glabrate,
pale brown or pale gray, the rhitidome somewhat scaly; stipules
narrow-linear, subulate, acute, tomentose, 3-5 mm. long, 0.5-1 mm.
wide, soon deciduous.

Leaves chartaceous, flexible, variable in size; petiole mediocre,
subterete, transversely rimose, densely and thickly lanate-tomentose,
ferruginous, when old the indument appressed, grayish, 4-15 mm. long;
blades obovate-elliptic or obovate-oblong, more or less narrowed to the
very asymmetrical and rounded base, more abruptly attenuate and
cuspidate at apex, entire or slightly sinuate at margin, 7 x 3 to 38x 13
cm., including the 0.5-3 cm. long acumen, varying very much in size
on the same branch, when very young stellate-pilose above, then
glabrous, pale olivaceous or pale brownish, shining, the main nerves
depressed, filiform, the others obsolete, pale ochraceous or pale cinere-
ous beneath, glaucescent, the costa very prominent, the 5-7 secondary
nerves on each side prominent, curved-ascending, near the margin
arched anastomosing, the lower pair usually forming an acute angle,
ascending and more distally separated from the others, the transverse
tertiary nerves thin but prominent, the minor veins forming a promi-
nulous conspicuous reticulum, the costa and the principal lateral
nerves covered when young, mainly towards the base, by a floccose
deciduous indument of ochraceous, entangled, stellate hairs, in time
becoming glabrous, shining, marked with minute callose, reddish,
sparse dots, the tertiary nerves also glabrous, the minor veins, reticu-
lum, and areoles covered by minute, white, stellate hairs forming an
appressed, whitish tomentum.

Inflorescences very small, axillary or in exfoliated, thin branchlets,
the axis and branchlets of the cymes very reduced, ochraceous-
ferruginous, lanuginose-tomentose; peduncles 2~7 mm. long, 3-bracteo-
late at apex; ‘pedicels 3-8 mm. long, somewhat thicker; bracteoles

560 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

linear, 1.5-2 mm. long; buds globose, also ochraceous-lanuginose and
woolly-tomentose.

Sepals thick, ovate, subacute, slightly involute-marginate, about
6-7 mm. long, 3-4 mm. broad, united 1 mm. at base, rosy or reddish
inside, subglabrous, glandular at base, 3-5 nerved, the margin mi-
nutely whitish tomentose, rather woolly outside, stellate-tomentose,
spreading in anthesis; petal-hoods yellowish or reddish obovate,
rounded cucullate and slightly retuse at apex, with 7 prominent nerves
and copious, spreading, minute hairs inside, subglabrous with few,
slender, flexuous hairs outside, 3-3.5 mm. long, 2.5 mm. broad; petal-
lamina deep red, rather thick, suborbicular, often retuse at apex,
abruptly contracted into a pedicel at base, pilose at margin, and with
very sparse, flexuous, slender hairs on the inner side, 3.5 mm. long,
4 mm. broad; pedicel red, 0.6 mm. wide with sparse, slender hairs,
2.5 mm. long.

Androecium tube about 1.5 m. high, glabrous; staminodes laminar,
thick-membranaceous, deep red, oblong-elliptic, rounded and often
emarginate at apex, attenuate near the base, with a conspicuous medial
nerve and thin, flexuous, subspreading hairs distributed throughout,
especially on the margins, 5-6 mm. long, 2.5-3 mm. broad; filaments
thick, glabrous, 1.5 mm. long, shortly 3-furcate, 3-antheriferous; anther
lobes ellipsoid, 0.4—0.5 mm. long; ovary ovoid, 1.5 mm. long, densely
tomentose-hirsute; styles glabrous, 5 mm. long, united only at base.

Fruit obovoid-ellipsoid, rounded at apex, contracted at base,
greenish, when ripe brown yellowish, 5-7 cm. long, 3-4 cm. broad,
the pericarp thin-coriaceous densely covered with acute, hard warts
and sparse stellate bairs, when dry about 1.5 mm. thick; seeds 16
mm. long, 9 mm. broad; germination hypogeous.

ComMon NAMES.—Cabeca de urubti is the most common and
widespread. Also, the following have been locally recorded:
copu-ai, cupu-curtia, cupurana, cacao de macao, urubt-acain,
cabeca de Umbu.

Win-cheék (Puinave), Infrida-Guaviare; ma-oo-hee-rée (Ka-
buyart) (Rio Cananarf) (Angl. Col. Cocoa Exped., Baker 1952).

DistRInuTION.—Spread throughout the western part of the Amazon
basin, on elevated ground and humid, fertile soil of rain forests. In
Brazil it is frequent in the western half of Amazonia, the easternmost
localities known being Teffé, on Rio Solimoes and Rio Jau, a tributary
of the Rio Negro (Ducke). The Anglo-Colombian Cacao Expedition
found it (although not abundantly) in the rivers Caguidn, upper and
lower Caqueté, and the Putumayo (Report, 1954, 12, 13). In Peru
goes as far as the lower parts of Rfo Huallaga and Rio Ucayali, Rfo
Itaya, and also Putumayo.

CUATRECASAS—CACAO AND ITS ALLIES 561

a

3 oAnT, 5

PES
S a
oh. ie es

=

n aed
Bitsy

ay,

Ficure 40.—Detail of indument on the underside of leafin: a, Theobroma simiarum (Stand-
ley 37377) from seedlings; 8, 7. simiarum (Pittier 7731), from adult branches; c, 7.

grandiflorum (Killip & Smith 30011); v, 7. obovatum (Ducke 265). a X 20, BX 30,
cand p xX 40.

562 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Theobroma obovatum is a rather small tree with a crowded, leafy
crown, easy to recognize by its small, ellipsoid-obovoid fruits con-
stricted at the neck, with thin, fragile, granulate or verrucose pericarp,
usually green but becoming brownish yellow when ripe. Also
characteristic are its light, papery, asymmetrical, oblong-obovate
leaves with a close, white or cinereous monotrichous tomentum
beneath; when very young the upper side, the principal nerves as
well as the petioles, stipules, and twigs, are covered with an ochraceous
brown or orange, easily removed down, composed of long, deciduous,
stellate hairs. The fruit when ripe separates from the pedicel and
falls.

Schumann did not know the species, for it is lacking in his her-
barium; he included the name 7. obovatum as a synonym of 7. sub-
incanum in his treatment in the Flora Brasiliensis (p. 77).

Huber knew the species well, although he called it 7. sylvestre,
confusing and misinterpreting Aublets figure of Cacao sylvestris.
Ducke also knew 7’. obovatum and its distribution very well, but he
thought that 7. sylvestre Mart. was a synonym.

COLOMBIA: Purumayo: Rio Caucayd, Rio Leguizamo, Laguna Primavera:

tree 15 m., 3 IV 1953, Holliday & Cope T/90 (COL, TRIN, US). Ibidem; tree
10 m., flowering, on elevated land above river, 5 IV 1953, Holliday & Cope T/95
(COL, TRIN, U, US).

Caqugta: Rfo Cagudn, camp 4, 27 IV 1953, Holliday & Cope T/119 (COL,
TRIN, US). Ibidem; found on sloping land, 30 ft., above river, tree 6 m.,
jorquettes arising symmetrically, 26 IV 1953, Holliday & Cope T/114 (COL,
TRIN, US).

AMAZONAS: Rfo Caqueté, La Pedrera, river level; tree 10 m., native in forest
on the riverbank, 7 X 1952, Baker & Cope 30 (COL, F, TRIN, US). Ibidem;
tree 7-8 m. in forest river-bank, presumed native, jorquettes regularly of three
branches growth continuing from above, young twigs and leaves with caducous
fuzz, flowers pale crimson, 5 X 1952, Baker & Cope 27 (COL, F, TRIN, US).
Ibidem; large tree 30 ft. high; calyx lobes light outside and pinkish inside, petals
dark purplish red, diameter 9-10 inches, 5 X 1952, Schultes & Cabrera 17775 (US).
Rio Caquetdé, Remolino; tree 6-8 m., no flowers but had two ripe pods, 2 V 1953,
Holliday & Cope T/123 (COL, TRIN, US). Trapecio Amazénico, Loretoyacu
River, 100 m. alt., XI 1945, Schultes 6921 (F).

BRAZIL: Pari: Belém, Jardim Botdénico do Museu Goeldi; small tree, “cacau
cabega de urubu,” cultivated, 11 VIII 1942, Archer 7537 (USDA,IAN). Ibidem;
arvore no. 482, 22 XII 1958, Cavalcante 339 (MG, US). Ibidem, Pires & Black,
s.n. (BH), 743 (IAN). Ibidem (Rio Purtis, loco dicto Bom Logar oriuntur, J.
Huber anno 1904 accedit); arbor parva floribus atrorubris, fructus maturitate
flavidis, ““cabega de urubu,” 4 IT 1926, Ducke 21044 (G, K, P, U, US). Ibidem,
Ducke 265 (A, F, K, MO, 8, US). Belém, cultivado in Instituto Agronémico
do Norte, ‘‘cabega de urubt,” Pires, Nilo, & Silva 4339 (IAN, UC, US).

Amazonas: San Antonio do Iga, matta, 27 IX 1906, Ducke 7704 (MG).
On the Rio Negro, Schomburgk 870 p.p. (1). Municipality Sao Paulo de Olivenca,
near Palmares; tree 35 ft. high, trunk 4 inches diam., terra firma highland, 11
IX-26 X 1936, Krukoff 8275 (A, BM, FE, F, G, K, MICH, MO, P, S, U, US,
USDA, Y). Basin Rio Madeira, Municipality Humayta, near Tres Casas, on

CUATRECASAS—CACAO AND ITS ALLIES 563

varzea land, shrub 25 ft. high, ‘‘cabeca de umbd,” 14 IX-11 X 1934, Krukoff
6263 (A, BM, F, GOET, K, LE, MICH, MO, 5, U, US, Y). ‘Amazonas Ega,”
Poeppig 2746 p.p. (LE). ‘Brasilia in sylvis circum Ega, 1831,” Poeppig, s.n. (WU).
Amazonas; Bocca et Teffé, matta, ‘‘cabeca de urubti,” 27 X 1904, Ducke 6823
(BM, G, MG, P, US). Teffé, matta virgen; tree, flower white, ‘‘copu-ai,’’ 221X
1947, Black 47-1496 (IAN). Igarapé Jandiatuba, lowland, border of the
river, 14 I 1949, Frées 23926 (US, IAN). Arredodes de Fonte Béa, terra firma
alta; arvore media, ‘‘cupurana,” “‘cupu-curua,” 12 IV 1945, Frdées, 20646 (F, K,
IAN, USDA). Rio Jurud, Juru4 Miry; Baum 3-9 m., Blumen dunkelpurpurn,
VIII 1901, Ule 5637 (G, HBG, K, MG, L). Purts, Monte Verde, “cacao de
macaco,”’ II 1904, Goeldi 4226 (MG). Rio Acre: Antimari, matta, 31 III 1904,
Huber 4295 (BM, G, MG, P, US). Upper Amazon, Paranary; low tree, petals
purple, stamens yellowish, staminodia yellow, ‘‘urubt-acaim,” 20 X 1924, Traill
61 (K, P).

Acre: Basin Rio Purtis, near mouth of Rio Macauhdn (tributary of Rio
Yaco), Lat. 9°20’ 5, Long. 69° W, on terra firma; tree 35 ft. high, 9 VIII 1933,
Krukoff 5388 (A, G, K, MICH, MO, 8, U, US). Ibidem; shrub 40 feet high,
3 IX 1933, Krukoff 5759 (A, BM, F, G, K, MICH, MO, 8, U, US).

Marto Grosso: Machado River region, source of the Jatuarana River; tree
45 ft. high in terra firma, ‘““cupuarana,” XII 1931, Krukoff 1668 (A, BM, G, K,
F, MICH, MO, 8, U, UC).

GuaporeE: Porto Velho, km. 8, matta on elevated ground; small tree, 17
VI 1952, Silva 155 (IAN).

PERU: Loreto: Along Rfo Itaya, Rfo Masana, 8 V 1929, Li. Williams 161
(F). Forest of Rio Itaya, 3 V 1929, Ll. Williams 230 (F, US). Lower Rfo
Huallaga, Puerto Arturo, Yurimaguas, 155-210 m., medium-sized tree, forest,
20 XI 1929, Ll. Williams 5268 (F, US). Balsa-puerto, 220 m., forest; tree 4 m.,
flowers wine red, IV 1933 Klug 2983 (A, BM, G, GH, K, F, MO, 8S, US). May-
nas, in sylvis circum Yurimaguas, 1831, Poeppig 1845 (G). Maynas 1831,
Poeppig 2352 p.p. (lectotype, WU; isosyntypes, F, GOET, G, LE, P). Stromgebiet
des Ucayali, von 10° S bis zur Miindung, 1923, Tessmann 3433 (G, S). Rain
forest of the Amazon basin, 230 m., 40 km. south of Pucallpa, 24 VII 1957,
Ellenberg 2551 (L). Bank of Rfo Putumayo, opposite Puerto Leguizamo; sucker
of large flowering tree, leaves rather large, 7 IV 53, Holliday & Cope T/98 (COL,
TRIN, US).

TRINIDAD (cult.): Imperial College of Tropical Agriculture, River State
Diego Martinez, Field 19; primary branching ternate; growing pseudoapical;
branchlets and twigs ochraceous lanate-tomentose, leaves thin, flexible, very pale,
with costa and secondary nerves yellow green, with woolly, floccose, deciduous
tomentum beneath, green above, 31 VIII 1961, Cuatrecasas, Cope, & Bartley
25783T (US). Ibidem; trunk 15 cm., bark lenticellate-granulate; branching
ternate; terminal branchlets light brown or brownish ochraceous, lanate-tomen-
tose, crown very branched and leafy, the lower branches horizontal, the upper
ones ascending, young, tender, terminal leaves hanging, light yellowish green,
1 IX 1961, Cuatrecasas & Cope 25788 (US).

17. Theobroma subincanum Mart.
Ficurss 29, 33, 35, 39, 41; Map 11; Puarss 9, 10, 11

Theobroma subincanum Mart. in Buchner, Repert. Pharm. 35:23. 1830;
Linnaea Litt. Bericht, 32. 1831; Bernoulli (1869) 13; Schumann in Mart.
(1886) 77; Jumelle (1899) 27 (in part); Huber (1906a) 274; Ducke (1925)
132; (1940) 272, pl. 4, fig. 1; Addison & Tavares (1951) 25, pl. 3, fig. 1,

564 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

&0° 70° 60° 50° 40?

|_|,

{Noe rz

0°?

iT

Map 11.—Geographical distribution of Theobroma subincanum @ and of its vicariants at
the western side of the Andes, 7. hylaeum & and T. nemorale ©.

pl. 4, fig. A, pl. 11, fig. 8; Ducke (1953) 10; Cuatrecasas (1956) 699; Baker,
Cope, & al. (1954) 12, fig. 13.
Cacao sylvestris Aubl. Pl. Guian. 2:687, pl. 276. 1775.
Cacao guianensis Aubl. Pl. Guian. 2:684. 1775, pro parte (tantum folia).
Theobroma sylvestris (Aubl.) Don, Hist. Dichl. Pl. 1:622. 1831, non Mart.
1830; Chevalier (1946) 279; Lemée (1952) 380; Leén, (1960) 322, 321, fig.
Theobroma ferruginea Bernoulli, Uebers. Art. Theobroma 13. 1869.
Theobroma alba Ruiz & Pavén, Fl. Peruv. Chil. 6, pl. 68, ined.
Typres.—Amazonas, Brazil, Martius. French Guiana, Aublet (of
Cacao sylvestris). Peru, Ruiz & Pavoén (of T. ferruginea).
Medium-sized tree commonly 6-12 m. tall, at times up to 20 m. high,
the trunk 15-20 (-30) em. in diameter, with gray, almost smooth
bark, older bark rugose-rimose, reddish within, the wood whitish,
darker toward the center; growth pseudoapical; primary branches
ternate, grayish, spreading; juvenile branchlets covered by a dense
ferruginous tomentum of stellate hairs, when older glabrescent, pale
brownish or brown, somewhat rugose, rimose-reticulate; stipules
narrowly linear, densely ferruginous-tomentose, 5-7 mm. long, 1 mm.
wide, soon deciduous.
Leaves firmly coriaceous, rather thick and large; petiole robust,
subterete, densely and appressed ferruginous-tomentose, 8-15 mm.
long; lamina elliptic-oblong or subobovate-elliptic-oblong, very little

CUATRECASAS—CACAO AND ITS ALLIES 565

attenuate to the base, slightly unequal, rounded or very obtuse,
emarginate or rarely cordate at base, somewhat narrowed or rounded
and abruptly acuminate at apex, sometimes blunt, entire, or near the
apex dentate-sinuate, 16-40 cm. long, 5-20 cm. broad, the acumen
acute, 1-3 cm. long, when very young ferruginous-tomentose through-
out, but soon glabrescent above, when adult glabrous above, green,
somewhat brownish olivaceous when dry, the costa and the lateral
nerves depressed, filiform, the lesser veins obsolete, cinereous beneath,
the veins more or less tawny or ferruginous, the costa thick, very
prominent, the 9 or 10 pairs of secondary nerves very prominent,
subascending, thinner near the margin, decurrent, the superior arched,
anastomosing, the basal pair often straighter and forming a more acute
angle, the transverse tertiary nerves prominent, the minor ones and
small veins prominulous, minutely reticulate, the midrib, major nerves,
and reticulum more or less densely covered by mediocre, reddish or
tawny stellate hairs, the areoles between the veins with a dense
whitish indument of very small, delicate, intricate, stellate hairs.
Inflorescence small, few-flowered, axillary or extra-axillary on leafy
branches; cymes with 3-9 fasciculate branchlets, usually 1-3-flowered;
peduncles 2-8 mm. long, with 3 bracteoles at apex, the bracteoles
subulate, about 3 mm. long, deciduous; pedicels 3-6 mm. long,
thicker than the peduncle; buds ovoid-globose; sepals thick, carnose,
ovate, acute or subacute, densely stellate-tomentose outside, ferru-
ginous, the margin minutely whitish tomentose, shining inside,
purplish or red, subglabrous with minute, crowded, oblong-capitate,
glandular hairs at base, near the margin slightly pubescent, 8-9 mm.
long, 3-4 mm. broad, united at base for 2 mm., subpatulous.
Petal-hoods thick-membranaceous, pale yellow and red striate,
obovate, rounded-cucullate at apex, slightly emarginate, 7-nerved,
inside minutely hirtulous, glabrous outside except for the puberulous
margin, 3-3.5 mm. long, 2-2.4 mm. broad; petal-lamina pedicellate,
carnose, thick, rigid, red, suborbicular, 2-2.5 mm. long, 2.2-4 mm.
broad, with slightly retuse apex, slightly pilose at margin, the hairs
very slender, flexuous; pedicel 2 mm. long, compressed, pilose.
Androecium tube 1.5-1.7 mm. long, glabrous; staminodes laminar,
red, lanceolate-oblong, acute or subacute, with marked midrib, sub-
glabrous with sparse flexuous hairs at margin, 6-7.5 mm. long, 2mm.
broad; filaments rather thick, glabrous, about 1.5 mm. long, arched,
very shortly 3-furcate, 3-antheriferous, the anther cells ellipsoid,
about 0.5 mm. long; ovary ovoid-oblong, 1.3 mm. long, glabrous, with
very sparse, minute, granulate dots; styles 1.5 mm. long, connivent.
Fruit ellipsoid, light green, tawny or orange at maturity, smooth,
oblong-ellipsoid or obovate-ellipsoid, rounded at apex, often more
or less narrowed at base, 7.5-11.5 cm. long, 5-6.6 cm. broad; pericarp

566 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

coriaceous, rigid, hard, 3-4 mm. thick, the woody epicarp 1-2 mm.
thick, covered by a brown, thin, appressed, stellate-tomentose in-
dument; seeds ellipsoid-oblong or ovate-ellipsoid, 1.8-2.3 cm. long,
12-16 mm. broad, 8-11 mm. thick, the surrounding pulp rather
slightly sweet, scentless, white, becoming yellowish; cotyledons white;
fruiting peduncle 1-1.5 cm. long, 0.5-1.0 cm. thick; germination
hypogeous.

Common NAMES.—The commonest names in Brazilian Amazonia
are cupui and cupuai. Other names or other ways of spelling and
pronouncing the former are: copuf, copuaf, cupuhy, cupuahy,
cupuy do Igapé, cupuarana, cupti do matta, cupti-assuy, cupti-assti-
rana. In Colombia, Venezuela, and Peru this species is usually
called cacao de monte or cacao silvestre, and also cacao rana (Orinoco
valley), yurac-cacao (Yurimaguas), uchpa-cacao, cacao-ceniza (Peru),
cacao blanco (Peru, Ruiz & Pavén). Indian names recorded are:
abekaré (Makuna, Vaupés, Garcia Barriga), padama (Arekuna,
Venezuela, cumalaé (Peru). The Anglo-Colombian Cocoa Expedition
(Baker, 1952) recorded the following indigenous names: Win-cheék
(Puinave), Infrida-Guaviare; bawk (Maku), Piraparana-Taraira;
poo-hoo (Barasana), Upper Piraparand; a-ba-ka-ra (Makuna), Lower
Piraparand Popeyaca; mah-we-re (Yukuna), Miritiparand; no-tér-
ree-ka (Yanimuka), Guacaya; too-soo (Yauna), Lower Piraparand;
ma-oo-hee-reé (Aabuyart), Canarari; wa-k6 (Kubeo), Cuduyari;
wah-pek-la (Tukano), Papuri; a-sé-ya-ee (Piratapuya), Papurf;
wa-be-ga-ra (Vesano), Papurf; wa-be-ka-ra (Siriano), Paca; ma-wé-
roo-da (Kuripaka), Guainia.

Usres.—Although this species gives an acceptable chocolate it
is practically never used by the natives. The slightly sweet and
scentless pulp is occasionally eaten or sucked; it is very much sought
by animals, especially monkeys.

DistTrRIBUTION.—Widespread throughout Amazonia from Parf to
the most western tributaries of the Amazon River, the upper Orinoco
range, and the Venezuelan and French Guayanas; frequent in the
shade of the Hylaean rain forests, in noninundatable lowlands, often
on rich and humiferous soil but ascending small hills on sandy grounds,
along creeks and small rivers. Theobroma subincanum is the species
of most frequent and abundant occurrence and with the broadest
area of distribution, other than 7. cacao.

COLOMBIA: Mera: Acacias, Canaima, farm 350 m. alt., cultivated, 18 XI
1951, Patio 22 (F). Sierra de la Macarena, trail from Rfo Guéjar to Cafio
Guapayita, Cafio Yerli, 500-600 m. alt.; tree about 35 ft. tall, flowers deep red,
fruit ripening brown, leaves rusty beneath, 20-28 XII 1950, Idrobo & Schultes
776 (COL, IAN).

Putumayo: Rfo Caucayd, Laguna Primavera on Rfo Leguizamo; tree 18 m.,
symmetrical jorquette, 3 IV 1953, Holliday & Cope T/91 (COL, TRIN, US).

CUATRECASAS—CACAO AND ITS ALLIES 567

tA SS
‘ak PM EMSS)
SITE LIA)

7
=
=

“2

Mian ge eae HN
AIX ' is Kh / f (7 of ‘ <F, i A 3
AANA ATV Zs ACR
pew 4 jan At cen y* AY)
VGN ZN Ws ys <i) AN VAN NARA
SAAN wre
LASS

<i RAK Se .

fiey ANE Wesel
vA rf |

Ficure 41.—Detail of indument on the underside of leaf in: a, Theobroma subincanum
(Cuatr. 7277); B, T. sinuosum (Pavon, s.n.); c, T. nemorale (Cuatr. 21921); v, T. mam-
mosum (Leon 1363). a,c, and p X 25,8 XX 10.

Vicinity of Mocoa; tree in forest, 15 m. high, no fruits or flowers, 17 III 1953,
Holliday & Cope T/81 (COL, TRIN, US).
680-695—64——13

568 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Caqueti: Rio Cagudn, camp 4; tree 10 m., 26 LV 1953, Holliday & Cope
T/116 (COL, TRIN, US). Upper Rfo Cagudn; small trees, 10 IV 1953, Holliday
& Cope T/101 (COL, TRIN, US), 13 IV 1953, Holliday & Cope T/103 (COL,
TRIN, US). Solana, 8 km. SE. of Tres Esquinas, on Rfo Caquet4, below mouth
of Rfo Ortegiiaza, 200 m. alt., wet tropical forest; tree 15 m. high, 15 cm. thick,
bark gray, smooth, with lichen patches and bryophyte patches, long, ellipsoidal
orange-brown fruits 10 x 6-6.5 cm., edible, “cacao silvestre,’ 4 VI 1945, Little
& Little 9544 (F, US) (fruit US No. 1096). Municipality of Florencia, site San
Luis, right margin of Pescado River, ‘cacao silvestre’ 5 VI 1942, Ranghel 195
(in part) (COL).

Vaupés: Cafio Mayabo, near San Felipe, river level; only flowers and old
fruits, no young or ripe fruit at this season, 27 X 1952, Baker 33 (COL, F, TRIN,
US). Mitd: Cafio Parand-Midf, 200 m. alt.; tree 5 m., 19 X 1939, Cuatrecasas
7277 (F, US, COL). Cerro de Mitti, 380 m. alt.; small tree with bending, fruiting
branches, fruits 9.3 X 6 em., 17 IX 1939, Cuatrecasas 6890 (COL, F, US). Mitt
and vicinity, 280 m. alt.; tree, cultivated, 22-30 VI 1958, Garcia Barriga, Schultes
& Blohm 16064 (COL). Ibidem; tree 5 m. with spreading branches, flowers red,
10-13 XI 1952, Garcta Barriga 15139 (COL). Cerro de Circasia, 300 m. alt.,
base of hill 10 X 1989, fruiting tree, 10 X 1939, Cuatrecasas 7178A (COL, F, US).
Rio Barbas, river level, native in the forest, 28 X 1952, Holliday 43 (COL, F, TRIN,
US). Rfo Cuduyarf, tributary of Rio Vaupés, Yacara, middle and lower course,
highland, + 700-800 feet, 9 XI 1952, Schultes, Baker, & Cabrera 18552 (US). Rio
Cuduyarf, river level, showing jorquette of 3 branches, 16 X 1952, Baker & Cope 31
(F). Ibidem; young red flowers, Baker, Bartley, & Holliday 31 (COL, TRIN). Rio
Cuduyarf, Pakwativa Mission, river level, heavily infested with Marasmius
perniciosus, 19 X 1952, Baker & Cope 32 (COL, F, TRIN, US). Ibidem; flowers
distinctly paler in color, Baker, Bartley, & Holliday 32a (TRIN). Rfo Apaporis,
Soratama, above mouth of Rio Cananarf, Cafio Surruco, 900 feet alt., highland;
small tree, leaves rusty beneath, fruit rusty brown, ovoid, 30 I 1952, Schultes &
Cabrera 15116 (US). Rio Apaporis, Rio Jinogojé, river level; tree 40-50 ft.,
native in forest or river bank, 13 IX 1952, Baker & Cope 12 (TRIN). Jinogojé,
growing just above flood level with numerous ellipsoid pods, Baker & Cope 7
(COL, TRIN). Ibidem; river level, tree 16 m., 20 cm. diam. at base, a long, tall
thin pole, without obvious jorquettes, native in forest, 23 VIII 1952, Baker & Cope
3 (COL, F, TRIN, US). Ibidem, 250 m. alt.; tree 15 m., large, coriaceous leaves,
brownish fruits 7.5 X 4.5 cm., 25, 26 VIII 1952, Garcia-Barriga 14224 (COL, US).
Rio Piraparan4, cultivated tree, 21 VIII 1952, Schultes & I. Cabrera 17005 (AMES).
Rfo Piraparan4, 250 m. alt.; tree 3 m., red petals, sepals red above, 22-26 VIII
1952, Garcta-Barriga 14203 (COL, US), 14253 (COL). Rio Piraparand, near
confluence with Rfo Apaporis, river level; small tree 3 m., in Indian garden,
24 VIII 1952, Baker & Cope 4 (COL, F, TRIN, US). Rfo Infrida, near Morichal,
near the mouth of Rfo Papunand, 200 m. alt.; tree 10 m., 14 II 1953, Fernandez
2275 (COL, US). Rio Infrida, Raudal, 300 m. alt., 3 IT 1953, Bartley & Holliday
T/71 (COL, TRIN, U, US). Rio Infrida, Santa Rosa; tree 12 m., 25 I 1953,
Bartley & Holliday T/68 (COL, TRIN, U, US). Rfo Infrida, right bank below
Cafio Caribe (5 hours above Morichal); tree about 10 m., no flowers, one small
fruit, 22 11953, Bartley & Holliday T/65 (COL, E, TRIN). Rfo Infrida, Morichal,
30 m. alt., in forest, small fruits, no flowers now, 8 II 1953, Bartley & Holliday
T/72 (COL, TRIN, US). Rfo Infrida, affluent Papunand, 300 m., red leaf flush,
nearly mature pod 11.5 X 6 em., fruit pedicel 2 cm. long, fruit green with brown
pubescence, pericarp woody, pulp white, cotyledons white, convolute, 18 II 1953,
Bartley & Holliday T/75 (COL, TRIN, US).

CUATRECASAS—CACAO AND ITS ALLIES 569

Amazonas: Rfo Caquetd, La Pedrera, river level; tree 45-50 ft., native in
forest, 29 IX 1952, Baker & Cope 25 (COL, F, TRIN, US). La Pedrera, river
level, native tree in forest, 26 IX 1952, Baker & Cope 21 (TRIN). Rfo Caquetd,
Remolino, leaves from young tree, 2 V 1953, Holliday & Cope T/124 (COL,
TRIN, US). Rfo Apaporis, near mouth of Rio Cananarf; tree 45 feet tall in
forest, fruit rust colored, ‘‘cacao de monte,” III 1951, Schultes 12104 (COL, US).
Rfo Apaporis, between Rio Pacoa and Rfo Cananarf, Soratama, 250 m. alt.; weak
tree, leaves rusty beneath, flowers red, 26 IX 1951, Schultes & Cabrera 14140
(US). Rfo Miritiparand, near varadero to Rio Apaporis, river level; tree 50 ft.,
native in forest, 15 IX 1952, Baker & Cope 13 (COL, TRIN). Trapecio Ama-
zonico, Loretoyacu River, 100 m. alt., [X 1946, Schultes 8385 (AMES, F). Leticia,
100 m. alt.; bark rough, cracked, light gray, flowers red, 20 IX 1945, Schultes
6536 (F). Leticia, forest; flowers pale red, tree 5 m., 22 VIII 1946, Black &
Schultes 46-61 (USDA).

VENEZUELA: Boutvar: Mount Duida, 500 m. alt.; small tree, VIII 1928-IV
1929, Tate 944 (NY, US). Caronf, rain forests of the Icaburu valley, 440 m.
alt.; tree 15 m., fruit large and tasty, ‘‘Spadamd”’ (Arekuna), XI 1947, Cardona
2379 (US, VEN).

Amazonas: Alto Rfo Orinoco, Taman4, 121 m. alt.; medium-sized tree (10-12
m.) with few branches, flowers purplish or red on the branchlets, trunk bent, up
to 40 cm. diam. with no branches in 2-5 m., bark gray, inside light brown, wood
pale brownish; “‘cacao-rana,” Ll. Williams 15204 (A, F, G, US, VEN). Rfo
Guainfa, Maroa, river level, with one ripe pod (wall mealy, smooth, thin and
brittle), seeds extracted and sent to Trinidad, Baker 38 (COL, F, TRIN, US).

FRENCH GUIANA: Aublet s.n. (part of syntype of Cacao guianensis Aublet,
BM, Photo, Mo. Bot. Gard. no. 4028). “Guyane Francaise,’ Potteau, s.n. 1819-
1821 (G). French Guiana, “Cacao sylvestris Aublet,”’ Aublet (syntype, BM, Photo
Mo. Bot. Gard. no. 4029).

BRAZIL: Amazonas: Rio Negro, Barcellos, ‘‘cupuhy,’”’ 25 VI 1905, Ducke 7200
(MG). Rio Negro, Rio Caure, Igarapé Miritf; tree 5 m., 15 em. diam., red
flowers, in rather lowland high forest, VII 1948, Frées 23343 (IAN, P). Rio
Negro, Porto Cabary, “‘cupuhy,” 4 XII 1945, Frées 21482 (IAN, NY, USDA).
“Prov. Rio Negro Martius Iter. Brasilienses 325,’ Martius [898] (M) (Photo
F. M. 19645). ‘“‘Prov. Rio Negro, Martius Iter. Bras. 325,” Martius [872, 873 p. p.,
894, 895, 896, 897, 899, 900] (M). “Rio Negro Dr. Martius Iter Brasil. 325,
Theobroma subincanum,” Martius [893] (M) (Photo F. M. 40704). Rio Vaupés:
Jauraté, inundatable forest, 17 X 1945, Frées 21162 (IAN, NY, USDA). San
Antonio de Iga, forest; tree 10 m. “differt a T. ferrugineo Bern. foliis majoribus
latioribus floribus majoribus,’”’ Ducke 7679 (MG, F) (Photo F. M. 7679). Mu-
nicipality of Saé Paulo de Olivenca, near Palmares; tree 40 ft. high, trunk 5
inches diam., high land, “‘cupuarana,” 11 IX-26 X 1936, Krukoff 8226 (A, BM,
F, G, GB, K, LE, MO, NY, P, U, US). Basin of Rio Madeira, Municipality
Humayta, near Livramento, on Rio Livramento, terra firma; tree 60 ft. high,
“cupuarana,” 12 X-6 XI 1934, Krukoff 7016 (A, F, G, K, MICH, MO, 8, U,
US, WU). Rio Madeira, Varadouro do Morcego, 31 VIII 23, Kuhlman 18110
(U). Manaos, mata; arvore, tree 30 m. alt., 15 cm. diam., “cupuhy,” 17 II 1945,
Frées 20518 (IAN, K, NY, USDA). Manaos, Mata do Aleixo, “cupuhy,’” 16
III 1945, Frédes 20555 (F, IAN, USDA). Manaos Aurora Fazenda, 15 m.,
Urwald, “‘cupi do Matto,” 28 VIII 1921, von Luetzelburg 22079 (M). Without
locality Spruce 97 (K).

Amapd: Rio Amaparf, Serra do Navio, slopes of Curuca Ore Body, down
to Igarape Sentinela; occasional tree 6 m. tall; fruit brown, 9 XI 1954, Cowan
38186 (NY, US). Lower slopes of Observatorio Ore Body, heavily forested hills,

570 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

70-300 m. alt.; tree 10 m. tall, fruit brown, 8 XI 1954, Cowan 38164 (NY). Missao
do Servico Florestal no T. Amaps, IX 1955, Miranda Bastos 68 (IAN).

Pari: Belém, ‘‘cupuahy,” VI 1896, Huber 162 (BM, G, MG, P, (US). Belém,
Bosques Rodriguez Alves, 1 VIII 1944, A. Silva 317 (TAN, USDA). Belém,
Horto do Museo Goeldi; tree 481, small tree, flowers brown yellow, 22 XII 1958,
Cavalcante 9388 (MG, US). Belém, Bosque municipal; tree 30 feet high, sparsely
branched, 4 VIII 1942, Archer 7517 (IAN, USDA). Belém, south forest of the
Instituto Agronémico do Norte; large tree, contents of fruit eaten by birds,
“cupu-assuy,”’? 16 XI 1942, Archer 7820 (IAN, USDA). Belém, along roads on
lands of Inst. Agr. do Norte; large tree, flowers rose color, ‘‘cupu-assu-rana,’”’ A.
Silva 237 (IAN). E. F. Braganga, Jofio Coelho; tree 8 m., 14 III 1947, Pires
& Black 1414 (IAN). Region of Igarapé Pitoré; tree 10 m., flowers red, 19 IX
1958, Frées 34663 (IAN). Taperinha, near Santarém, bushed river margins of
Igarapé Asst, “cupuy do Igapé,” 23 VIII 1927, Ginzberger & Hagmann 801
(F, WU). Rio Tapajéz, Cachoeira do Mangabal, “beira de assahyzal,” “cupuy,”
6 IX 1916, Ducke 16464 (BM, MG). Matta do Alto Ariramba, “cupuy” 7 X
1913, Ducke 14925 (MG). Rio Purtis, Monte Verde, “cupuahy,” II 1904,
Goeldt 4225 (MG). Paranary, upper Amazon, “cupua-i,’’ 20 X 1874, Trazll 59
(K, P).

Guaporé: Porto Velho, km. 8, in high forest on firm land; tree about 30
m., ‘‘cupuf,”’ 17 VI 1952, J. F. Silva 143 (IAN).

Rio pe JANEIRO: Quinta de S. Christovao; small tree planted by Riedel
probably originally from Amazon basin, “cupuaf,’’ 16 II 1876, Glaziou 9633a (P).

PERU: Loreto: Mishuyaco, near Iquitos, 100 m. alt., forest; tree 6 m.
high, flowers dark red, “wild cacao,” X-IX 1929, Klug 87 (F, NY, US). Ibidem,
II-III 1930; tree 8 m. high, flowers wine red on branches, Klug 857 (F, US).
Alto Itaya, 145 m., in forest of Parafso, ‘‘cumald,”’ 30 IX 1929, Ll. Williams
3254 (A, F, G, 8, US). Upper Rfo Nanay, Santa Ana, “uchpa-cacao,”’ 7 VII
1929, Ll. Williams 1233 (F, 8, US). Rfo Nanay, Tierra Doble, deep forest,
“Campamento balatero Lira Dabu,” 8 VI 1929, Lil. Williams 1076 (US, F, S).
“Peru,” Ruiz & Pavén, “Theobroma alba R. & P.” (ined.), ‘Theobroma ferruginea
Bern.” (K, BM). Ibidem, Herb. Pavén 201 (G). “Theobroma alba,’ Rivero
1836 (P). Tessmann, s.n., NY-3717; probably the same as the Tessmann 4115,
cited by Mildbraed (Notizbl. 11:139. 1931) as 7. ferrugineum and collected near
Pongo de Manseriche, 160 m. alt., tree 19 m. tall, 21 em. diam., ‘“pako-kakao”’
(Dunkel Kakao), 23 IX 1924.

HuAnuco: Prov. Hudnuco, Tingo-Marfa, forest, tree about 25 m. high,
sepals olivaceous yellow without, more or less red within, petals yellowish gray,
31 VIII 1940 Asplund 13410 (S). Tingo Marfa, forest; tree 15 m., flowers dark
red, 8 VIII 1940, Asplund 12911 (S). Maynas, Yurimaguas, ‘‘yurac-cacao”’
ie., “Cacao album Peruvianum,” March, 1831, Poeppig 2352 p.p. (GH, P, WU).

TRINIDAD (cultivated): On grounds of the I.C.T.A., from seeds from
Miti Vaupés region, Colombia; one branch damaged by Marasmius pernictosus.
Cuatrecasas, Cope, & Bartley 25785 T (U.S.)

18. Theobroma hylaeum Cuatr., sp. nov. Figures 25, 39; Map 11

Arbor circa 10 m. alta, apicaliter crescens, ramis primariis ternatis,
ramulis brunneis juvenilibus minute ferrugineo-tomentosis denique
glabratis, cortice rugoso-fissurato; stipulae lineari-subulatae acutae
vel subacutae ferrugineo-tomentosae, circa 6 mm. longae, 1 mm.
latae.

CUATRECASAS—CACAO AND ITS ALLIES 571

Folia mediocriter coriacea, petiolo crassiusculo subtereti minute
ferrugineo-tomentoso, 6-10 mm. longo; lamina elliptico-oblonga vel
obovato-oblonga, basi symmetrica attenuata obtusa vel subobtusa,
apice leviter attenuata interdum rotundata subite acuteque acumi-
nata, margine integra vel apicem versus leviter undulata, 12-20 cm.
longa, 4-9 cm. lata, apiculo 6-15 mm. longo, supra in sicco viridi-
brunnescens glabra, costa et nervis principalibus filiformi-impressis
reliquis vix conspicuis, subtus cinereo-ochracea vel nervatione fer-
ruginea, areolis venulisque cinereo-tomentosis pilis stellatis minutis
albidis intricatis dense tectis et pilis stellatis crassioribus ferrugineis
sparsis vel in nervis copiosis, costa crassa elevata, nervis secundariis
7 vel 8 utroque latere eminentibus arcuato-ascendentibus, prope
marginem tenuioribus anastomosantibus, nervis tertiaris transversis
prominentibus, nervulis venulisque tenuioribus sed prominulis
reticulatis.

Inflorescentiae breves axillares cymis 1-3 floribus instructae,
ramulis brevissimis tomentosis, pedunculis tenuibus erectis vel sub-
flexuosis ad 12 mm. longis ferrugineo-tomentosis apice 3-bracetolatis
et cum pedicello articulatis; bracteolae anguste lineares subacutae,
3-6 mm. longae, 1-2 mm. latae, extus tomentosae; pedicelli erecti
tomentosi pedunculis leviter crassiores, 5-6 mm. longi.

Sepala crassiuscula ovato-oblonga acuta, basi ad 2 mm. coalita,
patulo-reflexa, extus ochraceo-tomentosa, intus margine minutissime
albido-tomentella excepta glabra, aurantiaca, basi ad insertionem
pilis crassiusculis oblongis glandulosis annulum formantibus, 8-9
mm. longa, 4-5 mm. lata; petala cucullo aurantiaco obovoideo, basi
angustato, apice rotundato, saccato-cucullato, 3 mm. longo, 2 mm.
lato, extus glabro ruguloso, intus minute hispidulo, 7-nervato, pediculo
2.5 mm. longo, parce puberulo, lamina obovato-subrhomboidea
crassa. brunneo-rubra, faciebus sparsis minutissimis pilis margine
pilis flexuosis tenuibus praedita, 4-5 mm. longa, circa 4 mm. lata;
androecium tubo circa 1.5 mm. longo; staminodia petaloidea aestiva-
tione reflexa in anthesin curvato-patula, brunneo-rubra obovato-
oblonga apice rotundata vel leviter retusa, 5-5.5 mm. longa, 2.2 mm.
lata, minute pilosula; filamenta fertilia 1 mm. longa; antherae 3.
brevissime pedicellatae; ovarium obovatum 1.2 mm. longum, 1 mm,
latum, crasse tomentosum; styli circa 1 mm. longi coaliti.

Fructus coriaceus circa 7 x 4 cm., viridi-brunnescens, adpresse
tomentosus laevis ellipsoideus, utrinque rotundatus, epicarpio rigido
lignoso in sicco 1 mm. crasso; pedunculus fructifer 1.5—-1.8 cm. longus,
0.8 cm. crassus.

Type in the U.S. National Herbarium, No. 2028683, collected in
the heavy rain forest around Villa Arteaga, northern region of An-

572 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

tioquia, Colombia, at 200 meters altitude, August 14, 1948, by J.
Araque and F. A. Barkley no. 18C745.

ComMMoN NAMES.—Chocolate de monte, cacao silvestre.

DistriBuTION.—Presently known only from the Chocé region of
northern Antioquia, Colombia, and probably also in Panama.

COLOMBIA: Anrioqura: Villa Arteaga, heavy wet forests at 200 m. alt.;
tree 10 m., flowers orange, corona cream orange, stigma black brown, 14 VIII
1948, Araque & Barkley 18C745 (holotype, US 2,028,683; isotype, US 2,028,684,
COL). Ibidem; seedlings showing symmetrical branching in 3’s, growth continu-
ing from above the jorquette, 24 VII 1953, Bartley & Holliday T 166 (K, TRIN,
US).

PANAMA: Co.tén: Along Rio Faté, in forests or thickets 10-100 m. alt.,
VII 1911, Pittier 4194 (BM, GH, US) (probably; specimen sterile; identification
needs confirmation by fertile specimens).

Theobroma hylaeum is closely related to T. nemorale from which it
differs essentially by the narrow-linear bracts and stipules, by the
subrhomboid lamina, and by the narrower obovate-oblong, rounded
and notched staminodia; furthermore, the peduncles and the pedicels
are short, the pedicels shorter than the peduncles, and the fruit is
harder, smaller and not constricted above the base. From 7’. subin-
canum it differs by the shape of the petal-laminae and staminodes,
by the tomentose ovary, by the smaller fruit and leaves, and by the
venation, in which the minor veins are less conspicuous.

19. Theobroma nemorale Cuatr. Ficurss 3, 41, 42; Map 11
Theobroma nemorale Cuatr., Rev. Acad. Colomb. Cienc. 8:487, fig. 4. 1952;
Baker, Cope, & al. (1954) 13, fig. 20; Leén (1960) 323, fig. in 321.

Typr.—Pacific coast, Colombia, Cuatrecasas 21291 (fruiting speci-
men), Patiio 24 (flowering specimen, paratype).

Small or medium-sized tree up to 15 m. high; growth pseudoapical ;
trunk up to 20 cm. in diameter; primary branches ternate; leafy
branches ochraceous or ochraceous-ferruginous, or brownish, minutely
and appressed tomentose, the older glabrate, dark grayish, rather
smooth or granulate-lenticellate, nitidous, the hornotinous greenish
ferruginous, tomentose; stipules subcoriaceous, oblong, obtuse,
striolate, tomentose, about 8-11 mm. long, 2-3 mm. broad.

Leaves coriaceous, moderately rigid; petiole 9-12 mm. long, thick,
subterete, densely and appressed ferruginous-tomentose; blades el-
liptic-oblong or obovate-elliptic-oblong, slightly attenuate and asym-
metrical, or equilateral, rounded at base, rounded or obtuse and
abruptly acuminate at apex, entire or upwardly sinuate or coarsely
dentate, 10-32 cm. long, 3-12 cm. broad, the acute acumen 1.5-2.5
em. long, above green, pale brown when dry, rather shining, glabrous,
the costa depressed, thin, the other nerves rather inconspicuous,
cinereous or greenish cinereous beneath, or pale tawny when dry, the
costa very prominent, the secondary nerves about 8 on each side very

CUATRECASAS—CACAO AND ITS ALLIES 573

KAR

Ficure 42.—Leaves of Theobroma, X%: a, sinuosum (Pavon, s.n.); B, mammosum (Leon
1363); c, nemorale (Cuatr. 21291).

prominent, subparallel, ascending, curving and anastomosing near
the margin, the transverse tertiary nerves prominent, parallel, the
nerves of the fourth rank prominulous, broadly reticulate, the minor
reticulate veins concealed, the whole nervation tomentose, greenish
ochraceous or subferruginous, by rather thick, tawny, densely dis-
tributed stellate hairs, the surface within the veins covered by a dense
and appressed cinereous tomentum of white, fine, minute stellate hairs.

Inflorescences axillary or on exfoliated branchlets very small, re-
duced to 1-3 (-5) flowers, the cyme-axis extremely short, the peduncles
5-10 mm. long, erect, ferruginous-tomentose, 3-bracteolate at apex, the
pedicels thicker, ochraceous-ferruginous tomentose, 2-5 mm. long, the
bracts ovate, obtuse, 2.5-3 mm. long, 1.5-2 mm. wide; bracteoles 3,
broadly cochlear, embracing the bud, orbicular or ovate-rounded,

574 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

ochraceous-greenish tomentose, 7-11 mm. long and broad, soon
caducous.

Sepals thick, ovate-oblong, subacute or subobtuse, 9-10 mm. long,
4-5 mm. broad, united in the lower third or fourth, first curved-
patulous, later reflexed, inside purplish or rose, shining, glabrous
except for the minutely white-tomentose margin, and for the minute
oblong-capitate, glandular hairs at the base, thick-tomentose outside,
ochraceous, the stellate hairs with long, thin rays.

Petal-hoods thick-membranaceous, ochraceous or orange, obovate
with rounded-cucullate apex, 7-nerved, glabrous and rugose outside,
minutely hispidulous-pubescent, inside with 7 prominent veins, 3-3.5
mm. long, 2-3 mm. broad; petal-lamina pedicellate, carnose, rigid,
red or crimson becoming brownish red, oblong-obovate-spatulate,
rounded or subtruncate-emarginate at apex, with very minute, sparse
hairs on the surface and copious longer, slender, weak, spreading hairs
on the margins, 5-7 mm. long, 3-5 mm. broad; pedicel narrowly
linear, folded, puberulous, at the broader base barbate, 3 mm. long;
androecium tube 1.5-2 mm. long; staminodes laminar, rather thick,
obovate-oblong, rounded or subspatulate at apex, purplish red or
crimson, covered with sparse, thin, minute hairs, 6-7.5 mm. long,
3.5-4.5 mm. broad, reflexed in bud, spreading in anthesis; filaments
moderately thick, 1-1.2 mm. long, pilose towards the base, 3-furcate
(the branchlets 0.4 mm. long), 3-antheriferous, the anther lobes
ellipsoid 0.5-0.6 mm. long, connivent; ovary 1.3-1.5 mm. long, oblong-
ovoid, subtruncate, 5-ridged, thickly ochraceous, hirsute-tomentose;
styles linear, glabrous, 1.5 mm. long, coherent.

Fruit ellipsoid, smooth, rounded at apex, slightly constricted above
the base, 8-10 cm. long, 4.5-6 cm. broad; pericarp coriaceous, rigid,
1-1.5 mm. thick, when mature fragile, its surface densely appressed
stellate-tomentose, greenish or bluish, at maturity yellowish, brown
or tawny; seeds compressed ovoid or subellipsoid, 17-19 mm. long,
9-12 mm. broad, the surrounding pulp white, yellowish when ripe;
cotyledons white; fruiting peduncle robust, 1.5-2 em. long, about 0.8
mm. thick; germination hypogeous.

ComMMon NAMES.—‘‘Bacao de monte,’”’ “chocolate de monte,”
“bacao,” ‘“bacafto,’”’ “cacao de monte,” “cacaito de monte.’’

Usrs.—It is said to produce a fairly good chocolate; not known to
be used by the natives.

DistrisutTion.—Restricted to the Colombian Pacific coast and
the Chocé area between the parallels 3° and 5°3’ N. latitude. It is
recorded from the Calima, San Juan, and Cajambre Rivers.

This very interesting species is closely related to 7. subincanum and
T. hylaeum, but it differs specially by its three broad, orbicular
bracteoles subtending the flowers, which before anthesis embrace

CUATRECASAS—CACAO AND ITS ALLIES 575

and cover the short-pedicellate bud. This feature is unique in the
genus Theobroma.

The leaves of T. nemorale, as well as those of T. hylaeum, are very
similar to those of T. angustifolium, but they are broader, more oblong-
elliptic, with a tendency to an obovate shape, and the ochraceous or
ferruginous hairs beneath are smaller, with shorter, somewhat
thicker rays than in T. angustifolium.

COLOMBIA: Cuocé: Rfo San Juan, Palestina; part of branch brought in by
an Indian, one immature fruit 7 x 4.5 em., 4 VIII 1953, Holliday T/149 (TRIN,
US). Ibidem, young tree 3 m., sterile, 2 VII 1953, Holliday T/147 (TRIN,
US). Ibidem, sucker 14 m. from leafless fallen trunk, 2 VII 1953, Holliday T 148
(TRIN). Ibidem, Quebrada de las Sierpes; tree 10 m. tall, leaves green yellowish
above, ashy beneath, 24 IX 1961, Cuatrecasas & Willard 26051 (COL, US).
Istmina, tree about 7 m., sterile, flowers apparently borne only on branches, 2
VIII 1953, Holliday & Bartley T/173 (TRIN, US).

Et Vaute: Pacific Coast, Rfo Calima, La Trojita, 5-50 m. alt.; small tree;
leaves coriaceous yellowish green, fruit ovoid-ellipsoid, 8 x 4.5 cm., light brown,
“bacao de monte,’”? 28 II 1944, Cuatrecasas 16544 (F, VALLE). Rfo Calima,
Quebrada de la Brea, 20-40 m. alt.; tree 8 m., leaves subcoriaceous, medium
green above, green cinereous beneath, nerves green ochraceous, fruits 10 x 6 cm.,
ellipsoid, rounded at apex, constricted at base, tawny, on the branchlets, ‘‘choco-
late de monte,” 24 V 1946, Cuatrecasas 21291 (holotype, F; isotypes, B, F,
VALLE). Ibidem, La Brea, flowering specimens, Patio 24 (paratype, F; type
of flowers; isoparatypes, F, US). Estacié6n Agroforestal del Calima, 30-50 m.
alt.; erect tree with abundant handing pods, 8 I 1953, Patifio 117 (US, Herb.
Cuatr.). Ibidem; small tree 8-10 m., ‘‘cacao,” “‘cacaito de monte,” “‘bacaito,”
8 I 1953, Patiwio 116 (US, Herb. Cuatr.). Ibidem; tree with primary ternate
branches from near the ground; leaves light green; twigs tomentose ochraceous
or ferruginous, sepals thick, rose or purplish rose inside, ochraceous outside,
petal-laminae and staminodes thick, rigid, purplish red or dark brown red, hoods
ochraceous with 7 red veins, “bacao,” 23 IX 61, Cuatrecasas & Willard 26007
(COL, US). Cafio La Brea; young tree 1.5 m., sterile, 29 VI 1953, Holliday
T/141 (TRIN, US). Estacién Agroforestal; tree 8-10 m., in land cleared from
forest, jorquette symmetrical, crimson and yellow flowers borne singly or in pairs
on small branches, pods 8.5-10 x 5-5.5 cm., fruit peduncle 2 cm. long, 0.75 cm.
thick, 29 VI 1953, Holliday T/146 (TRIN, US). Pacific coast, Rio Cajambre,
Silva, Loma de la Vigia, 5-80 m. alt.; small tree, leaves green above, gray beneath,
‘“bacao de monte,’”’ II-V 1944, Cuatrecasas 17503 (F, VALLE). Ibidem, Que-
brada del Corosal 0-5 m. alt.; tree 15 m. tall, trunk 20 cm. diameter, leaves
coriaceous, green above, ashy beneath, fruits ellipsoid rounded at apex, con-
tracted above the base, smooth, brownish, 10 x 5.5 cm., “chocolate de monte,”
17 V 1944, Cuatrecasas 17738 (F, VALLE).

TRINIDAD (cult.): Imperial College of Tropical Agriculture, River State
Diego Martinez, Field 19; 7-8 years old tree, 31 VIII 1961, Cuatrecasas, Cope, &
Bartley 25782 T (US).

20. Theobroma sinuosum Pavén ex Huber Fiaurss 41, 42; Map 9; Puate 12
Theobroma sinuosum Pavén ex Huber, Bull. Herb. Boiss., II, 6:274. 1906.
Theobroma Tessmannii Mildbr. Notizbl. Bot. Gard. Berlin 11:139. 1931.
Theobroma sinuata Ruiz & Pavén, Fl. Peruv. et Chil. Fol. E, Plate 417, ined.

576 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Typres.—Chicoplaya, Peru, Ruiz & Pavén (lectotype, G). Marafién
River, Peru, Tessmann 4928 (type of T. tessmannit).

Large tree with erect trunk; leafy terminal branchlets brownish
ochraceous when dry, densely hirsute-tomentose, covered by minute,
whitish, stellate hairs with intricate, slender rays (0.1-0.2 mm. long)
and by larger stellate, ferruginous hairs with straight, acute, long
rays (1 mm. long); stipules lanceolate, about 1 cm. long, deciduous.

Leaves subcoriaceous, firm; petiole short, thick, densely tomentose-
hirsute, about 1 cm. long; blades obovate-oblong, slightly attenuate,
rounded and slightly asymmetrical and emarginate at base, sub-
rounded and abruptly triangular-cuspidate at apex, sinuate-dentate
in upper third, otherwise entire, about 30 cm. long, 13 cm. broad,
pale green brownish above, when adult glabrous with abundant
reddish, punctiform scars, the midrib and secondary nerves filiform
impressed, the other nerves less visible, softly tomentose beneath,
subochraceous, covered by a dense, cinereous layer of minute, white,
fine, stellate hairs and, especially on nerves, by larger ochraceous,
stellate hairs, with 4-6, erect, acute, rays 1-1.5 mm. long; the midrib
thick, prominent, densely tomentose-hirsute, the secondary nerves
about 9 on each side, ascending, prominent, vanishing near the margin,
hirsute, the tertiary, transverse nerves prominent, parallel, 5-10 mm.
distant from each other, the minor veins reticulate and prominulous.

Sepals ovate-lanceolate united at base, reflexed ; petal-hoods obovate,
concave; petal-lamina obcordate-“trigonous”; staminodes lanceolate-
obovate; stamens five, 3-antheriferous; ovary hirsute; fruit sub-
rounded-pyriform, the epicarp smooth, hard, woody, ferruginous
tomentose.

The preserved original specimens from the Rufz and Pavén herbar-
ium surely collected by Tafalla have only leaves; they have been used
by me for the above description. The short description of flowering
characters has been taken from the manuscript of the unpublished
Flora of Peru and Chile of Ruiz and Pavén. This description was
written by Ruiz or Pavén based on data sent by Tafalla. Some of
the data were misinterpreted by the authors who describe the anthers
“quinque in singulo filamento,” and this is not the case. Tafalla
wrote in his “notas” that each filament was divided in six “lacinias”’
bearing one anther each, and that the number of stamens was five in
T. alba, sinuata, and cordata, whereas it was ten in 7. digitata. Pavén
described the inflorescences as being cauline; we have no basis either
to affirm or deny that assertion. The petal-lamina is described as
“trigona”; probably it was slightly 3-dentate, and this feature was
extremely exaggerated by Pulgar in his drawing.

Theobroma sinuosum, because of the little information available, was
disregarded by most of the authors who listed cacao species; Chevalier

CUATRECASAS—CACAO AND ITS ALLIES 577

who saw specimens of it considered it as a synonym of 7. ferrugineum,
which was not well defined by him in his revision. My study of the
sterile specimens of the type convinces me that T. sinuosum is a very
different species, unique in possessing an indument comparable to
that of the young plants of 7. chocoense and T. simiarum. On the
other hand, after careful study of the description and photograph of
T. tessmannu Mildbr. which I had formerly associated with 7’. subin-
canum, I arrived at the conclusion that the Tessmann specimen is
definitely distinct from T. subincanum and that it coincides with T.
sinuosum. Unfortunately the Tessmann specimens are not existing
any more, but the Mildbraed’s detailed description of the indument
permits us to differentiate his species from the closest allied species
growing in the same area, namely T. subincanum. Some doubts
may remain about the identity of 7. sinuosum and T. tessmannii.
The Tessmann plant definitely had axillary inflorescences, whereas
the Pavén plant was described as being cauliflorous, but the species
could well have both cauline and axillary inflorescences. The geo-
graphical range and the identical kind of indument are the reasons
that I consider the species synonymous.

I wish to supplement the description given above with the data
taken from Mildbraed’s description: Leaves subcordate at base,
25-35 cm. long, 8-12 cm. broad, with about 10 secondary nerves;
sepals lanceolate, acute, about 10 x 4 mm., connate at base for 2 mm.;
petal-hood whitish, lamina red, subquadrate-rounded and apiculate,
3.x 3 or 2 x 3 mm.; staminodes dark red, subspatulate-elliptic, 6 mm.
long, 3 mm. broad; stamens 3-antheriferous; pedicels 2 cm. long.

CoMMON NAMES.—Cacao de monte, Pako Kakao (Tessmann).

Usns: The pulp of the fruits is eaten by the natives.

DistrisutTion.—Upper river valleys of the Huallaga and Marajién
in Peru.

PERU: Chicoplaya, ‘‘Pavén,’’ collected by Tafalla (lectotype, G; BM). Rio

Marafién from Iquitos to the mouth of Santiago, near Pongo de Manseriche,
ca. 77°30’ W., Tessmann 4928 (type of T. tessmannii, photo F. M. 17942).

21. Theobroma canumanense Pires et Frées, sp. nov. Fiaure 43; Map 9

Arbor 18 m. alta pauciramosa ramulis dense ferrugineo-tomentosis,
pilis crassiusculis mediocribus stellatis 8-14 radiis acutis 0.3-0.7 mm.
longis instructis.

Folia rigide coriacea, petiolo crasso dense ferrugineo-tomentoso
8-10 mm. longo. Lamina oblongo-obovata vel obovato-elliptica basi
paulo angustata rotundataque leviter asymmetrica apice subrotundata
subite breviterque acuminata margine integra vel sursum leviter
grosseque dentata, 8-20 cm. longa, 3-8.5 cm. lata, supra in sicco
tabacina subnitida leviter rugosa juventute pilosa deinde glaberrima,
costa nervisque secundariis filiformibus impressis ceteris paulo con-

578 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

spicuis, subtus ferrugineo-tomentosa, costa valde elevata, nervis
secundariis 5 vel 6 utroque latere elevatissimis duobus basilaribus
ascendentibus ceteris patulo-ascendentibus arcuatisque ad marginem
decurrentibus anastomosantibus, nervis tertiariis transversis bene
elevatis 3-6 mm. inter se distantibus, nervulis quaternariis elevatis
transversis, minoribus prominenter minuteque reticulatis, pilis dimor-
phis: a) pilis stellatis albidis minutissimis dense intricatis areolas
tectis, b) pilis ferrugineis crassiusculis mediocribus stellatis radiis
patulis supra nervationem copiosissimis.

Flores in cymis axillaribus vel extra-axillaribus paucifloris congeste
glomerati; pedunculi ad 6 mm. longi tomentosi apice 3-bracteolati,
bracteolis 1-1.5 mm. longis triangularibus tomentosis; pedicelli
tomentosi 1 mm. longi. Alabastra globosa crasse ferrugineo-tomentosa
cirea 5 mm. diam. Calyx sepalis crassiusculis carnosis in anthesin
subpatulis paulo reflexis, circa 7 mm. longis, 4 mm. latis, basi 1.5 mm.
longitudinaliter coalitis, intus glabris basi excepto, pilis densis crassis
glandulosis ad marginem insertione praeditis extus dense stellato-
tomentosis margine dense minutissime pilosulis.

Cucullus petali obovoideo-ellipsoideus circa 3-3.5 mm. longus,
2-2.2 mm. latus, carnosulus extus glaber intus 7 costis elevatis stri-
gulosis instructus. Lamina petaloidea rubra crassa semirotundata
apice emarginata, circa 2 mm. longa, 3 mm. lata, sursum utrinque
minutissime pilis margine ciliata, basi subite in pediculum circa 2 mm.
longum longe ciliatum contracta.

Androecium rubrum; tubus circa 1.5-1.6 mm. altus glaber. Sta-
minodia crassa curvata subspathulata sursum dilatata apice leviter
emarginata, circa 6 mm. longa, 2.4-2.8 mm. lata, margine longe
flexuoso-ciliata cetera glabra. Stamina filamentis glabris crassis,
1.2-1.4 mm. longis triantheriferis, lobis antherae ellipsoideis circa
0.4mm. longis. Ovarium 1.2-1.5 mm. longum obovatum pentagonum
in angulis et subapicem hirsutulum. Styli circa 2 mm. longi glabri
versus apicem plus minusve liberi. Fructus ignotus.

Type in the U.S. National Herbarium, No. 2404642, collected in low,
firm land, Rio Canumao, tributary of Madeira River, municipality of
Borba, State of Amazonas, Brazil, October 5, 1957, by R. L. Frées
(No. 33783). Isotype in the herbarium of Instituto Agronomico
do Norte, Belém do Para.

Theobroma canumanense is closely related to T. sinuosum; its
leaves and indumentum conform well with those of the type of this
species showing only some differences due to the fact that the T.
sinuosum specimens came from a young plant. The vegetative
characters of the Frées plant also coincide with the description of
T. tessmannii given by Mildbraed, but I found a few differences
which indicate that the Frées specimens belong to a different species.

CUATRECASAS—CACAO AND ITS ALLIES 579

Ficure 43.—Theobroma canumanense (Froes 33783): a, petal, inside view; B, petal, lateral
view, X 5; c, androecium, X 5; p, stamen, X 10; £, sepals, inside view, X 2; F, pistil,
xX 5; 6, bud, X 2.

T. canumanense has smaller flowers, the sepals being only 7 mm.
long; they are 10 mm. in J. tessmannii which also has a longer
androecium (8.5 mm. high); the petals and staminodes are glabrous
in the Tessmann plant according to the Mildbraed description and
drawings, while they are long-ciliate in J. canumanense. The
inflorescences are more compact in this species than in T. tessmannii.
The photograph of the latter shows the adult leaves to be subcordate
and broader at base and provided with one or two additional pairs of
basal veins (not conspicuous in J’. canumanense); the secondary
nerves are more numerous (8-11 pairs) in Tessmann’s plants.
T. canumanense also differs from all related species by its extremely
short pedicels (up to 1 mm. long).

Common NAMES.—Names and uses not recorded.

DisTRIBUTION.—Limited to the region of Rio Canumao, a tribu-
tary of the Rio Madeira in Brazil.

BRAZIL: Amazonas: Regiéo do Rio Madeira, Rio Canuméo, municipio de
Borba, 5 XI 1957, R. L. Frées 33783 (US, holotype; IAN, isotype).

Section 6. Andropetalum

Theobroma sect. Andropetalum Cuatr., sect. nov. Figure 4

Lamina petalorum anguste spathulata longe attenuato-stipitata ;
cucullum 7-nervatum; staminodia crassa latissima petaloidea arcuato-
reflexa petala obtegentia; stamina 3-antherifera; fructus ellipsoideo-
oblongus laevis tomentosus supra basim constrictus ad apicem
angustato-mammillatus epicarpio duro lignoso; semina germinatione
hypogaea; folia subtus adpresse stellato-tomentosa ; inflorescentiae
axillares brevissimae; caulis incrementum pseudoapicale; rami

580 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

primarii ternati; calyx cymbiformis 3-lobatus, sepalis usque ad
tertiam partem vel ad medium connatis.

Type specius.— Theobroma mammosum Cuatr. & Leén.

This section comprises a single species. It is characterized by the
broad, obovate-spatulate staminodes (as broad as long), which are
reflexed even during anthesis, completely covering the petals, and
by the reduction of the petal-laminae, which are narrower and
smaller than in the closely related section Glossopetalum.

22. Theobroma mammosum Cuatr. & Leén
Fiaures 6, 36, 41, 42, 44; Map 9
Theobroma mammosum Cuatr. & Leén, in Leén, Inst. Interamer. Cienc.
Agr. Bol. Técn. 2:1-6, figs. 1949; Leén (1960) in 320, 317, fig.

Typr.—Siquerres, Limén, Costa Rica, Ledn 291.

Small tree, 6-7 m. high; trunk about 25 cm. in diameter with
rather smooth, brown bark 1 mm. thick and white, hard wood; growth
pseudoapical; branches ternate, from near the base, spreading or
more or less descending; terminal branchlets grayish or somewhat
ochraceous, appressed and minutely stellate-tomentose, later gla-
brous, gray, rugose; stipules subulate, acute, 4-5 (-10) mm. long, 0.6
mm. wide, deciduous.

Leaves subcoriaceous; petiole rather thick, subterete, straight or
somewhat flexuous, ochraceous or ferruginous, stellate-tomentose, 5—
12 mm. long; blade elliptic-oblong or oblanceolate, slightly attenuate
to the apex and suddenly acuminate, slightly narrowed toward the
asymmetrical base, rounded in one side, subrounded or subcuneate
at the other side, the margin entire or sinuate-dentate near the apex,
10-25 cm. long, 3.5-10.5 cm. broad, including the acumen, this acute,
8-20 mm. long, glabrous above, dark green, when dry pale brownish,
the costa filiform, depressed, the other nerves hardly noticeable,
cinereous beneath, except for the main nerves densely and appressed
stellate-tomentose, covered by thin and minute, white stellate hairs,
the costa very prominent, the prominent secondary nerves 9-12 on
each side, regularly parallel, subascending, curved and vanishing near
the margin, the transverse tertiary nerves filiform, prominent, the
lesser veins reticulate, thin, prominulous, but covered by the tomen-
tum, the midrib, secondary, and tertiary nerves with scattered or
copious, larger, thicker, spreading, stellate hairs.

Inflorescences very small, axillary, cymose with few (usually 2)
flowers (1-3), the axis extremely short, tuberculate, giving rise usually
to a single ferruginous-tomentose branch 8-12 mm. long, this 2- or 3-
furcate at apex into 2 or 3 peduncles; peduncles very short, about 1
mom. long, 3-bracteolate at apex, each articulate to a pedicel; bracteoles
linear, 2-4 mm. long, tomentose; pedicels rather thick, tomentose,

CUATRECASAS—CACAO AND ITS ALLIES 581

CH. REINZECH.

Ficure 44.—Theobroma mammosum (Le6n 1363): a, petal, inside view, X 5; B, petal,
laterally, X 5; c, part of androecium, X 5; p, flower in anthesis, X 2.5; E, gynoecium,

x 5.

8-12 mm. long; buds globose, densely ferruginous tomentose, about
1 cm. in diameter.

Sepals 11-12 mm. long, 5-6 mm. broad, thick, triangular-ovate,
united completely by pairs or by 2 and 3, and all five in the lower
third, forming a cupular, umbilicate, 2 or 3 short-lobate calyx, yellow-
ish green inside, red tinged, glabrous except for the glandular, thick,
erect, fasciculate, congested trichomes at the inner base, these 0.4—
0.7 mm. long, densely, thickly, stellate-tomentose outside, greenish
ochraceous or ferruginous.

Petal-hoods about 7-8 mm. long, 5-7 mm. broad, elliptic-obovate,
broadly rounded-cucullate at apex, dark red, rather carnose with
7 thick nerves, between the nerves veined with yellowish or rose by
transmitted light, minutely papillose inside, rugulose outside with
sparse, long, acute, straight or slightly flexuous hairs throughout,
these more copious at margin; petal-lamina erect in full anthesis,
narrow, thick, dark red or purplish red, truncate-spatulate or slightly
emarginate, usually plicate, with long, ferruginous, slightly flexuous
hairs, these scattered outside, abundant at margin and inside, 3 mm.
long, 2.5-3 mm. broad, gradually narrowed toward the base into a
plicate pedicel about 4 mm. long, 0.5 mm. wide.

582 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Androecium purplish red or dark red, the tube carnose, about
4 mm. high, 3-4 mm. broad, glabrous; staminodes large, petaloid,
reflexed, the lower part thick, carnose, toward the apex gradually
thinner, membranaceous, transparent-veined, obovate-spatulate, sub-
truncate and slightly sinuate at apex, laterally usually 1-dentate, the
lower half suddenly narrower, about 11-12 mm. long, 10-12 mm.
broad at top and 2.5 mm. wide at base, glabrous or with sparse hairs
near the top outside, with copious, ferruginous, simple or stellate,
rather long hairs inside; filaments thick, glabrous, about 1 mm. long,
shortly 3-furcate, 3-antheriferous, the lower branchlet extremely
short, the other two lateral about 1 mm. long; anther lobes ellipsoid,
suborbicular, about 1 mm. long after anthesis; ovary subglobose,
slightly 5-angulate, whitish, densely tomentose-hirsute, about 1.8
mm. high and thick; styles 2 mm. long, glabrous, erect, acute, con-
nivent, only united at base.

Fruit at maturity 16-22 cm. long, 6-8.5 cm. broad, cylindric-oblong,
terete or seldom slightly pentagonous, broad and umbilicate at base,
strongly contracted and slightly pentagonal above the base, suddenly
narrowed near the apex forming a mammiform, umbilicate end 2-3
cm. long and 2-2.5 cm. broad. Pericarp coriaceous, hard, smooth
or slightly verrucose downward, pale green, covered by dense, short,
ferruginous tomentum, the woody epicarp 1.5-2 mm. thick with a
tomentose epiderm outside, the mesoendocarp carnose, white, about
7 mm. thick; pulp enveloping each seed fleshy, fibrous, white; seeds
ovoid-amygdaloid, brownish, 22-27 mm. long, 14-17 mm. broad,
10-13 mm. thick; cotyledons white; germination hypogeous.

ComMOoN NAME.—Cacao silvestre.

DistripuTion.—Kastern coastal mountains of Costa Rica, where
it is extremely rare in primary forest. It has been found wild only
twice, at altitudes from 300 to 800 meters. Cultivated in few agri-
cultural experimental stations.

COSTA RICA: Lrmén: La Lola, 100 m. alt., 12 V 1948, Escamilla, s.n. (MO).
Siquirres, Finca La Lola, 300 m. alt., 11949, Leén 291 (holotype, TURRI).
La Lola, cerca de Madre de Dios, 100 m. alt., cultivated, 30 I 1949,Leén 1363
(paratype, F, TURRI). Ibidem, Experimental Station II.C.A., cultivated;
tree 6 m., stem 20-25 cm. diam., leaves chartaceous, firm, green above, pale
beneath, branches ternate, growth pseudoapical, flowers dark red and brown red,
fruit ellipsoid, contracted at apex, 6 XI 1961, Cuatrecasas & Paredes 26535 (US).

Herepia: Puerto Viejo, Sarapiqu{, 700-800 m. alt., IV 1959, Holdridge
146 (TURRI).

Cartago: Turrialba, grounds of I.I.C.A., cultivated, 600 m. alt., 4 XI 1961,
Cuatrecasas & Leén 26516 (US).

TRINIDAD: River State Diego Martinez, I.C.T.A., Field 19, cultivated from
seed received from Belém do Par4; tree 8 m. high, stem rugose-tuberculate, brown,

abundant dry fruits hanging from branchlets, 1 1X 1961, Cuatrecasas & Cope
25791 (US).

CUATRECASAS—CACAO AND ITS ALLIES 083
Hybrids

Theobroma angustifolium Moc. & Sessé o&’' X mammosum Cuatr. & Leén 9.

Trinidad, I.C.T.A., Cuatrecasas & Cope 25800. Well-developed
tree, with intermediate characters.

Theobroma grandiflorum (Spreng.) Schum. X obovatum Klotzsch.
Belém do Paré, Brazil, Museu Goeldi, arvore 480, 22 XII 1958,
Cavalcante 937 (MG, US).

Theobroma grandiflorum (Spreng.) Schum. ? X subincanum Mart. o.
Hybrid fertile obtained by Addison and Miranda (1951) 13. In-
termediate features of parents.

Theobroma grandiflorum (Spreng.) Schum. ? X obovatum KI. .

Brazil. Fertile hybrid obtained by Addison (1951) 10, pl. 1, fig. 8,
pl. 2, fig. C. F. cultivated at Belém do Paré, 6 XI 52, Pires 4343
(IAN). Ibidem, Pires 4344 (IAN).

Theobroma mammosum Cuatr. & Leén o& X simiarum Donn. Smith.?

Costa Rica, Turrialba IICA, robust young trees obtained by Soria.

Theobroma mammosum Cuatr. & Leé6n 9 X simiarum Donn. Smith o.

Costa Rica, Turrialba, IICA, robust young trees obtained by Soria.

Theobroma obovatum Klotzsch | X subincanum Mart. 9.

Fertile hybrid obtained by Addison and Miranda (1951) 14, pl. 13,
jig. 13, pl. 4, fig. C. Also found spontaneous by Cope «& Holliday
in Colombia: Rio Caqueté, Remolino, tree 11-12m., with some
characters of T. obovatum and others of T. subincanum, 2 V 1953;
it has almost smooth pods but bears flocose, woolly pubescence on
the young shoots and leaves, Holliday & Cope T 122 (COL, TRIN, US).
Caqueté, Rio Caguin, tree 8 m., 27 IV 1953, Cope & Holliday T 117
(COL, TRIN). Brazil, Amazonas, Fonte Béa; tree 15 m. alt. ‘cu-
purana,” Frées 20648 (IAN, US). Trinidad, ICTA, experimental
camps, Cuatrecasas & Cope 25797 (US).

Theobroma speciosum Willd. ex Spreng. @ X sylvestre Mart. ?.

Belém do Paré, Brazil, 6 XI 1952, Pires 4345 (IAN, COL). Ex-
perimental hybrid by Addison. It has intermediate characters as
described and illustrated by Addison & Miranda (1951) 14, pl. &

Jig. 8, pl. 6, fig. C.
Theobroma speciosum Willd. ex Spreng. ? X sylvestre Mart. <.

Belém do Pard, Brazil. Experimental hybrid obtained by Addison,
similar to the former. Addison & Miranda (1951) 15.

At the Instituto Interamericano, Costa Rica, Dr. Soria (1961)
tried to hybridize T. cacao with T. angustifolium, mammosum, simia-
rum, and bicolor. The cross 7. angustifolium o X cacao 9 gave small

680-695-6414

584 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

plants which did not grow more than about 10 cm. in two years.
Theobroma cacao 2 X mammosum of gave fruits, but the seeds were
very weak; the same happened with 7. cacao 9 X simiarum o. The
cross 7’. bicolor & XX cacao 9 was negative. See also Addison and
Miranda in the Historical Sketch above.

Rejected Names and Excluded Species

Theobroma Kalagua de Wildeman, Bull. Herb. Boiss. 7:957, pl. 11. 1899.

Typr.—Panama, Patin, s.n. (lectotype, BR, leaf) (Ff. M. Photo
40742).

This species was described by de Wildeman as very remarkable,
because of the extraordinary combination of flowers similar to those of
T. angustifolium and T. simiarum, fruits resembling T. simiarum,
and quite different leaves which although resembling those of T.
angustifolium in shape differed by the lack of pubescence. But
Patin gave assurance to Wildeman that these different parts came
from a single tree. De Wildeman wrote: ‘‘qui appartiendraient sans
le moindre doute 4 la méme plante. Une confusion aurait cependant
pu étre possible parce que le Th. simiarum existe également en
Colombie, ot M. Ch. Patin, qui s’adonne a la recherche des plantes
utiles, l’a découvert 4 Choco, province de Cauca” (Wild. p. 958).
In the U.S. National Herbarium, attached to a herbarium sheet
(1,382,338), is a letter sent to J. Donnell Smith by Ch. Patin, dated
Brussels 19 Oct. 1900. Here Patin states ‘“‘As concerns 7. Kalagua

. . there was a doubt about the leaves used to determine the specy
[sic]: the young plants which we have just got here from seeds have
proved that it occurred really a mistake in the description made of
the leaves brought to me by my collector.” This contradicts the
earlier assurances given by Patin about all his specimens having been
collected from a single tree and shows that he was not the collector
himself. Later on Patin writes, “I think that the 7. Kalagua is
just your 7. Simiarum.” Patin never explained where his specimens
were collected. Probably they came from the region of Panamé.
Regarding the specimens sent with the above cited letter, the seedling
leaves and small fragments of fruits might well be those of T. simiarum.

To clear up this question, in 1953 I asked Dr. Robyns, director of
the Jardin Botanique de l’Etat, Brussels, to send me the material
of T. kalagua preserved at that herbarium. Very kindly, Dr. Robyns
sent me the only existing herbarium sheet of the type material of
T. kalagua, which is represented only by one leaf, undoubtedly the
same used to illustrate the plate in the publication. To a further
request to Dr. Robyns the answer was given that no other material

CUATRECASAS—CACAO AND ITS ALLIES 585

of this species existed except for some fruits which “selon toute
vraisemblance appartiennent au matériel Patin s.n. recolté en 1899,
se trouvaient dans nos collections de fruits.’’

Examination of the Patin specimens proves that the leaf mounted
on this sheet at BR (photo F. M. No. 40742) is the original Patin leaf
used for de Wildeman’s illustration and that this leaf belongs to
Theobroma cacao. This leaf is the only unquestionable part of the
type apparently existing, and I select it as the lectotype. The three
fruits received on loan from Brussels labeled T. kalagua were: 1) A
half shell, to be discarded as belonging to Theobroma bicolor. 2) An
entire pod with a modern label from “J. Bot. Br.” reading “Th.
Kalagua Wild. Colombie, sans date Coll. Patin.” This fruit is
smooth, ellipsoid, oblong, slightly attenuate and rounded at apex and
broadly rounded, umbilicate at base, 18-20 cm. long, and 8.8 cm.
broad; the surface is tomentose, the woody epicarp about 1.5 mm.
thick, the dried, spongy mesoendocarp 2-4 mm. thick; a fragment of
this shell is lacking, and this could be the fragment of fruit sent by
Patin to Donnell Smith and preserved at the U.S. National Herbarium
(1,382,338). This fruit is not that used for the illustration, because
the original was sectioned; furthermore, this pod looks somewhat
more oblong than the one figured in the plate. It may well belong to
T. simiarum, as may also the seedling leaf sent to Donnell Smith.
3) The third fruit received from Brussels bears two labels; one reads
“Theobroma kalagua, Colombie, Chocéd, Prov. de Cauca, leg. Ch.
Patin 1899,” and the number 823; the other label reads ‘Theobroma
simiarum D. Sm. Colombie, leg. Ch. Patin’’; it consists of half a shell
whose section is ovate-ellipsoid, 19 cm. long and 14.2 cm. broad and
about 2 cm. thick; the woody epicarp is 1.5-2 mm. thick; the thick
mesoendocarp is compact. It is different from all species of Theo-
broma known to date; it might be an undescribed species, but it could
also belong to another genus.

In conclusion, we may infer that Patin gathered several specimens
of different Theobroma species collected by different persons and
coming from several places (Panama, Chocé ...). The flowers
described by de Wildeman could well be flowers of 7. angustifolium
or T. simiarum; the possibility of their growing in Panama, perhaps
planted, cannot be discarded. They also could have been flowers
from trees of 7. stipulatum, T. chocoense, or from some undescribed
species. An important character of the flower, its color, was
not mentioned by Patin. The fruit shown in the plate is thicker
than that of T. simiarum; it recalls very much that of 7. grandiflorum,
a species more or less widespread in gardens, and it is also similar to
that of 7. chocoense and T. stipulatum. It may also belong to an
undescribed species. ‘To ascertain to what species such a fruit belongs,

586 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

it would be necessary to see at least two associated organs from the
same tree, that is to say fruit and leaves, fruit and flowers, or, also,
flowers and leaves. The above comments on the available data
prove the disorderly way of Patin’s work and his unreliability.
There is no reason to believe that the flowers and fruits of Patin’s
collections were brought from the same tree or from the same species.
Patin never mentioned the locality of the specimens sent to de Wilde-
man or who collected them. Colombia is always cited but at that
time Panama was part of Colombia. At present the only identifiable
Patin specimen from his syntypes is the leaf (BR) and that is T.
cacao. Consequently, T. kalagua becomes a synonym of T. cacao.

Theobroma guianense (Aubl.) Gmel., Syst. Veg. ed 13, 2:1151. 1796.
Cacao guianensis Aubl. Pl. Guian. 2: 683, pl. 275. 1775.

Syntypres.—Aublet illustrations (l.c.) and specimens at British
Museum (Natural History).

Under Cacao guianensis, Aublet gave a detailed and illustrated
description made up of a mixture of three different species. There is
complete agreement between the description of each part and the
corresponding illustration. The flowers were described from specimens
of Theobroma cacao L.; the branches and leaves from 7. subincanum
Martius; the fruits cannot be identified with any other species included
in Bernoulli’s and Schumann’s treatments. The illustration of this
fruit (Pl. 275, figs. 16 & 17) agrees unmistakably with only one recent
collection of Theobroma fruits, namely that from French Guiana by
Benoist, which is the type of T. veluténum Benoist. Aublet’s short
diagnoses of the fruit, especially the French description ‘‘l’ovaire
devient une capsule ovoide 4 cing arrétes arrondies saillantes” also
agree perfectly with it. It seems that Aublet considered the most
typical part of his ‘‘species” the fruit, since he headed the description
of his Cacao quianensis with a short definition based only on the
fruit: “CACAO (G@uianensis) fructu ovato, quinquangulari, tomen-
toso, rufescente (Tabula 275)” (p. 683); furthermore, as Sandwith
pointed out, the French name given by Aublet [Le Cacaoier anguleux.
(Planche 275)] was taken from the same diagnosis. Thanks to Benoist
we know now that this kind of fruit belongs to a species with a kind
of leaves very different from those described by Aublet, the species
described by Benoist as 7. velutinum.

But the nomenclatural problems have to be solved on the basis of
types according to the International Code. At the British Museum
there are preserved Aublet specimens and among them type specimens
of Cacao guianensis which are syntypes. In 1954, I could identify
by close examination an herbarium sheet (with foliage, one flower,

CUATRECASAS—CACAO AND ITS ALLIES 587

and an immature fruit) as belonging to 7. subincanum Mart.; this
specimen agrees with the description and drawings given by Aublet
for the leaves and branchlets; this evidence could easily incline
us to use this specimen as lectotype for C. guianensis. But in the
carpological collection of the British Museum there is a fragmentary
fruit, also a syntype of Aublet’s species, which belongs to T. cacao.

Until now, there has been only confusion about the identity of
Theobroma guianense. Bernoulli (p. 7) wrote: ‘Cacao guianensis
bleibt somit eine vollstaendig ungewisse Art. Sie scheint auch von
keinem weitern Autor gesehen worden zu sein, sondern immer nach
Aublet citiert zu werden.” Schumann placed it as synonym of
Theobroma cacao, surely on account of its flowers. Chevalier con-
sidered the species synonymous with 7. speciosum Willd., erroneously
interpreting Aublet’s descriptions and drawings of the leaves and a
photograph of the Aublet herbarium specimen (syntype) at the
British Museum; these, as I have pointed out above, belong to T.
subincanum Mart. He also identified the Benoist collections of 7.
velutinum erroneously as T. speciosum.

Consequently, Cacao guianensis Aubl. is not a true species, but a
mixture of three species. Therefore the Aublet “species” and name
has to be rejected as “nomen confusum”’ (articles 63(3), 65, and 66
of the Code of Nomenclature). The name Theobroma guianense
has never been consistently used in monographs and general books
for any known species.

“Theobroma fossilium” Berry, Proc. U.S. Nat. Mus. 75(24):8, pl. 1, figs. 13, 14.
1929.

In regard to this unfortunate name, the following opinion of R. W.
Brown, former paleobotanist of the U.S. Geological Survey, is final:
“This specimen, considered by Berry to be the first fossil record of
Theobroma, is a section of the forepart of a reptilian jaw. Berry
mistook the bony structure for the pulp, and the teeth for the seeds
of a chocolate-bean pod. Although described among Tertiary
plants, the specimen, as stated by Berry, came from near Leiva,
Department of Boyacé, Colombia, where Cretaceous strata crop out.”
(Journ. Washington Acad. Sci. 36:353. 1946).

Theobroma alba Bernoulli, Uebers. Art. Theobroma 14. 1869; Jumelle (1899)
35; De Wildeman (1902) 98. 1902.

Typr.—Essequebo et Cuyaunic, British Guiana, C. F. Appun 1,
1860 (holotype, Herbarium Hookerianum, K).

The type of this species consists of leaves of a young, sterile plant
of the genus Licania. I have identified it by comparison from the
abundant material existing in the Royal Botanic Gardens, Kew, with

588 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

L. venosa Rusby, a widespread species of the Guianas. The following
new combination is necessary:

Licania alba (Bernoulli) Cuatr., comb. nov. Theobroma alba Bernoulli, Uebers.
Art. Theobroma 14. 1869. Licania venosa Rusby, Descr. New Sp. So.
Am. Pl. 26. 1920.

Theobroma albiflorum (Goudot) De Wildeman, PI. Trop. Gr. Cult. 90. 1902==
Herrania albiflora Goudot, Ann. Sci. Nat. III, 2:230, pl. & 1844.
Theobroma aspera (Karsten et Triana ex Triana) Van Hall, Cacao, ed. 2, p. 49.
1932. (Brotobroma aspera Karsten et Triana ex Triana, Nuev. jen. y esp.
fl. Neo-Granat. 12. 1854)=Herrania nitida (Poepp.) Schultes, Caldasia

2:16, 17, pl. 1948.

Theobroma augusta L. Syst. Nat. 3:233. 1776; Willd. Sp. Pl. 3:1424. 1803=
Abroma augusta (L.) L. f. Suppl. 341. 1781.

Theobroma balaénsis (Preuss) De Wildeman, Pl. Trop. Gr. Cult. 89. 1902=
Herrania balaensis Preuss, Exped. Centr. und Siid-Amerika 253, pl. 7.
1901.

Theobroma camargoanum (Schultes) Ducke, Bol. Técn. Inst. Agron. Norte
28:15. 1954=Herrania camargoana Schultes, Bot. Mus. Leafl. Harvard
Univ. 14:120, pl. 29, 32. 1950.

Theobroma celtifolia Salisburg, Prodr. 387. 1796=Guazuma ulmifolia Lam.
Encyel. Méth. 3:52. 1789.

Theobroma guazuma L. Sp. Pl. 782. 1753=Guazuma tomentosa H. B. K. Nov.
Gen. Sp. 5:320. 1823. Freytag (1951) 214.

Theobroma hastata Chevalier, Rev. Bot. Appl. 26:273. 1946, nomen nudum;
lapsus calami for T. sagittata Pavén ex Chevalier.

Theobroma laciniifolium (Goudot ex Triana et Planchon) De Wildeman, PI.
Trop. Gr. Cult. 90. 1902=Herrania lacintifolia Goudot ex Triana et
Planchon, Prodr. Fl. N. Granat. 209. 1862.

Theobroma mariae (Martius) Schumann in Mart. Fl. Bras. 123:71, pl. 16.
1886= Herrania mariae (Mart.) Decaisne ex Goudot Ann. Sci. Nat. III,
2:233. 1844.

Theobroma montana Goudot ex Bernoulli, Uebers. Art. Theobroma 15. 1869,
nomen nudum. No description.

Theobroma nitidum (Poepp. et Endl.) Schumann in Mart. Fl. Bras. 123:72.
1886. (Abroma nitida Poepp. et Endl., Nov. Gen. Sp. Pl. 3:73. 1845) =
Herrania nitida (Poepp. et Endl.) Schultes, Caldasia 2:16,17, pl. 1943.

Theobroma purpureum Pittier, Repert. Sp. Nov. Fedde 13:319. 1914=
Herrania purpurea(Pittier) Schultes, Caldasia 2:333. 1944.

Theobroma pulcherrimum (Goudot) De Wildeman, Pl. Trop. Gr. Cult. 89.
1902=Herrania pulcherrima Goudot, Ann. Sci. Nat. III, 2:232, pl. 6.
1844.

Theobroma sagittata Pavén ex Chevalier, Rev. Bot. Applig. 26:274. 1946,
nomen nudum= Herranta nitida (Poepp. et Endl.) Schultes, Caldasia 2:16,
17, pl. 1948.

Theobroma tomentosa (H. B. K.) Gémez, An. Hist. Nat. 19:217. 1890==
Guazuma tomentosa H. B. K. Nov. Gen. Sp. 5:320. 1823.

Theobroma undulata Pavén ex Chevalier. Rev. Bot. Appliq. 26:268. 1946,
nomen nudum: lapsus calami for T. sinuosum Pavén ex Huber.

Collections Cited

Acosta Sorfs, M., & Giuer, M. Baker, R. E. D., Bartiey, B. G., &
12392 gileri Hou.ipay, P. C.
12423 _ gileri 31 subincanum
Auten, P. H. 32a subincanum
3105 grandiflorum Baker, R. E. D., & Corr, F. W.

4593 bicolor 2 bicolor
6259 angustifolium 3 subincanum
6341 angustifolium 4 subincanum
Apron, C.F. 5 grandiflorum
1 Alba 6 grandiflorum
ARAQUE, J., & BARKLEY, F. A. 7 subincanum
18C745 hylaeum 11 glaucum
ArcHer, W. A. lla bicolor
7517 subincanum 12. subincanum
7537 obovatum 13. subincanum
7549 grandiflorum 21 subincanum
7551 microcarpum 25 subincanum
7619 speciosum 26 bicolor
7721 speciosum 27 obovatum
7734 grandiflorum 28 microcarpum
7820 subincanum 29 microcarpum
ASPLUND, E. 30 obovatum
10271 bicolor 31 subincanum
12911 subincanum 32 subincanum
13410 subincanum BarRTLEY, B. G., & Houuipay, P. C.
AvuBLET, J. B. C. F. T 46 grandiflorum
s.n. cacao (fruit) T 47 bicolor
s.n. subincanum T65 subincanum
BAFOG (Bureau AaR. ET FoRESTIER T 66 bicolor
GUYANAIS) T 68 subincanum
136M _ velutinum T69 glaucum
7386 velutinum T70 = glaucum
Baiuey L. H. T 71 subincanum
s.0. angustifolium T 72 subincanum
BakKER, C. F. T 74 glaucum
62 grandiflorum T 75 subincanum
421 grandiflorum T 166 hylaeum
2102 angustifolium BENoIsT, R.
Baker, R. E. D. 516 velutinum
16 grandiflorum BERNOULLI, G.
33 subincanum 94 bicolor

34 bicolor

37 glaucum

38 subincanum
s.n. angustifolium

95 angustifolium
BERNOULLI, G., & Carrio, R.
3188 angustifolium
3145 bicolor

589

590 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Brack, G. A.
47-1496 obovatum
47-1502 grandiflorum
47-1889 speciosum
Buack, G. A., CorpErro, E. &
FRANCISCO, J.
52-14649 microcarpum
52-14655 speciosum
Buack, G. A. & Lepoux, P.
50-10644 sylvestre
Buack, G. A. & Scuuutss, R. E.
46-61 grandiflorum
46-61 (USDA) subincanum
46-III_ grandiflorum
BoEHLMER, F. DE
12229 bicolor
BonpLanp, A.
s.n. bicolor
Brenss, A. M.
12333 bicolor

British Honpuras Forests Con-

SERVATION
H2192/29 bicolor
Broapway, W. E.
8935 angustifolium
BurcuE.u, W. J.
9375 grandiflorum
9467 grandiflorum
B. W. [BoscHwezEn]
1161 velvutinum
CaLpEROn, 8S.
627 bicolor
630 angustifolium
23610 bicolor
Camaroo, F. C.
8 microcarpum
2395 speciosum
Capucno, P.

397 speciosum
CarvENosa, B., MurGuEITIO,
P. R. & BarKkuey, F, A.

17C934 bicolor
Carpona, F.
2379 subincanum
CAVALCANTE, P.
310 bicolor
339 obovatum
937 grandiflorum X obo-
vatum
938 subincanum

Copr, F. W., & Howupay, P. C.

(see Holliday & Cope)

Cook, O. F.
4 bicolor
Cook, O. F., & Doyrtg, C. B.
50 bicolor
Cook, O. F., & Griaas, R. F.
756 bicolor
Coorer, J. J.
10244 simiarum
Cooper, G. P., & Suater, G. M.
242 = angustifolium
Cowan, R. S.
38164 subincanum
38186 subincanum
CUATRECASAS, J.
6890 subincanum
7277 + +subincanum
7178A subincanum
14897 cirmolinae
15336 cirmolinae
15700 _ cirmolinae
16160 bernouillii subsp. capil-
liferum
16526A bicolor
16544 nemorale
16896 chocoense
17034 bernouillii subsp. capil-
liferum
17034A _ bernouillii subsp. capil-
liferum
17350 bernouillii subsp. capil-
liferum
17350A bernouillii subsp. capil-
liferum
17503 nemorale
17738 nemorale
21291 nemorale
21339 = stipulatum
Cuatrecasas, J., & Corr, F. W.
25788 obovatum
25789 angustifolium
25790 angustifolium
25791 mammosum
25792 = simiarum
25794 simiarum
25795 bicolor
25797 obovatum X subincanum
25800 angustifolium X mammo-
sum
25801 grandiflorum

Cuatrecasas, J., Corr, F. W., &

CUATRECASAS—CACAO AND ITS ALLIES

Bart.ey, B. G.

25780T grandiflorum
25781T grandiflorum
25782T nemorale
25783T obovatum
25784T bicolor
25785T subincanum
25786T bicolor
25787T bicolor

Cuatrecasas, J., & Lrdén, J.

6823 obovatum

7200 subincanum

7202 bicolor
7202B bicolor

7216 sylvestre

7397 bicolor

7638 bicolor

7679 subincanum

7704 obovatum

7822 sylvestre

7884 speciosum

26515 simiarum
26515A simiarum
26516 mammosum

CuaATRECASAS, J., & PaREDEs, A.

26534 bicolor

26535 mammosum
26536 simiarum
26537 angustifolium
26538 microcarpum

Cuartrecasas, J., & WILLARD, L.

26007 nemorale
26051 nemorale
26074 chocoense
26167 gileri
Curopontis, G.
92 bicolor };
599 simiarum

DauucReN, B. E., & Sexua, E.

10 microcarpum
438 grandiflorum
634 grandiflorum
733 grandiflorum
739 grandiflorum

Davipson, G. W. R.

s.n. bicolor
Dawe, M. T.
83 bicolor

Dopesr, C. W., & Gorrcer, V. F.

9420 simiarum

Dopez, C. W., & NEVERMANN

7164 simiarum

Dopesz, C. W., & Tuomas, W. S.

6399 angustifolium
Duckg, A.
100 sylvestre
103 sylvestre
265 obovatum
281 speciosum
283 microcarpum
598 grandiflorum
4878 sylvestre
6773 microcarpum

7975 speciosum
10669 sylvestre
12187 sylvestre
14734 sylvestre
14925 subincanum
16458 grandiflorum
16464 subincanum
16466 microcarpum
21044 obovatum
21045 microcarpum

Duque JARAMILLO, J.

1205 bicolor

4403A bicolor

Kocrars, H. F. A.
14244 bicolor
ELLENBERG, H.
2551 obovatum
2565 speciosum
EscaMILLA, G.
s.0. mammosum

2371 bicolor
FERNANDEZ P, A.
2275 subincanum
Fossera, F. R.
21310 bicolor
Francisco, J.
1966 grandiflorum
Fréss, R. L.
20463 sylvestre
20518 subincanum
20555 subincanum
20556 = sylvestre
20625 bicolor
20645 glaucum
20646 obovatum
20648 obovatum X
canum
20655 sylvestre
20750 microcarpum
20885 glaucum
20942 glaucum

591

FacEerLinp, F., & Wrsom, G.

subin-

592 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

21162 subincanum
21482 subincanum
21556 grandiflorum
23343 subincanum
23926 obovatum
23963 microcarpum
25554 sylvestre
26526 speciosum
28382 sylvestre
29732 speciosum
30180 speciosum
30432 speciosum
31414 speciosum
33783 canumanense
33788 sylvestre
34663 subincanum
34949 sylvestre
Fr6égs, R. L., & Frixo, J. P.
29465 sylvestre
Garcia Barriaa, H.
11178 bicolor
14203 subincanum
14224 subincanum
14253 subincanum
14380 glaucum
14416 bicolor
15139 subincanum
Garcfa Barriga, H., Scuuxtss, R. E.,

& Buon, H.
16064 subincanum
GARDNER, C. A.
870 bicolor
GENTLE, P. H.
3464 bicolor
GEOFFROY,
8.0. speciosum
Giuer, M.
162 gileri
168 _ gileri
Giter, M., & Patifo, V. M.
164 gileri
165 _ gileri
166 gileri

GINZBERGER, A,

802 speciosum
GINnzBERGER, A., & Hacmann, M.
801 subincanum-.
GINZBERGER, A., & ZERNER, H.
800 grandiflorum

Guaziou, A. F. M.
9633 speciosum
9633a subincanum
9643 grandiflorum

GoELDI, ANDREAS
4225 subincanum
4226 obovatum
4228 microcarpum

Goneceryp, J. W.
4108 velutinum
4127 velutinum
4148 *;velutinum

GouDoT
s.n. bicolor
GUERRERO, H.
26074 chocoense
HANBURY
9471 grandiflorum
Hart, J. H.
158 bicolor
5381 augustifolium
HOrFMANNSEGG, W.
8.n. speciosum
Houprives, L. R.
146 mammosum
5133 _ gileri
s.n. bicolor
Houurpay, P. C.

T 43 subincanum

T 139 cirmolinae

T 140 cirmolinae

T 141 nemorale

T 142 bernouillii subsp. capil-
liferum

T 143 chocoense

T 144 chocoense

T 145 bernouillii subsp. capil-

liferum

T 146 nemorale
T 147 nemorale
T 148 nemorale
T 149 nemorale
Houupay, P. C., & Bartuey, B. G.
T 163 gileri
T 165 stipulatum
T 167 gileri
T

liferum
173 nemorale
175 stipulatum

HAHAH

liferum

4

liferum

172 bernouillii subsp. capil-

176 chocoense var. bullatum
177 _bernouillii subsp. capil-

178 bernouillii subsp. capil-

CUATRECASAS—CACAO AND ITS ALLIES

Houupay, P. C., & Corz, F. W.

T 77
T 79
T 79A
T 81
T 90
T 91
T 94
T 95
T 96

|
i=)
oe)

101
103
114
115
116
117
118
119
122
123
124
125

HHA RRA AAR a aA

162
1567
1748
4008
4295
7081

bicolor
glaucum
glaucum
subincanum
obovatum
subincanum
glaucum
obovatum
glaucum
obovatum
subincanum
subincanum
obovatum
glaucum
subincanum
obovatum Xsubincanum
glaucum
obovatum
obovatum Xsubincanum
obovatum
subincanum
microcarpum
Huser, H.
subincanum
speciosum
speciosum
grandiflorum
obovatum
microcarpum

HumsBo tpt, A., & BONPLAND, A.

5.n.

Iproso, J.

776

bicolor
M., & ScHu.tss, R. E.
subincanum

INPA (INstitTuTO NACIONAL DE
PrEsquisas DA AMAZONICA, MANAOs)

1966
2125

903
237

8.0.
KIL.uIpP,
30006
30011
30320

87
857
1523
2021

grandiflorum
sylvestre
JOBERT, DR.
speciosum
Jounson, H.
bicolor
Karsten, G.
glaucum
E. P., & Smita, A. C.
bicolor
grandiflorum
grandiflorum
Kuve, G.
subincanum
subincanum
bicolor
bicolor

2983

1080
1117
1274
1644
1668
5295
5388
5759
6203
6263
6592
7016
8226
8275
8280
9019

593

obovatum

Krovuxorr, B.

speciosum
speciosum
grandiflorum
microcarpum
obovatum
speciosum
obovatum
obovatum
microcarpum
obovatum
microcarpum
subincanum
subincanum
obovatum
microcarpum
bicolor

KuHLMANN, J. G.

18110

§.n.

12056

subincanum
LABROY
grandiflorum
LANGE
bicolor

Leymann, F. C.

7909
9021

291
937
1363
3189
4267
4832

bicolor
bicolor

Lroén, J.
mammosum
angustifolium
mammosum
simiarum
angustifolium
gileri

LInDEMAN, J. C.

5725

simiarum

Lirrte, E. L., & Lirriz, R. R.

9544
9598

8.n.

subincanum

bicolor
Luano, E.

bicolor

Léprz, J. R.

8.n.

1

angustifolium
Lucas, A.
bernouillii subsp. asclepi-
adiflorum

LUETZELBURG, PH. V.

22007
22079
23065
23287
23895

grandiflorum
subincanum
bicolor
grandiflorum
bicolor

594 CONTRIBUTIONS FROM THE NATIONAL

Martius, C. E. P.
862 bicolor
863 bicolor
864 bicolor
865 bicolor
871 sylvestre
872 subincanum
873 p.p. grandiflorum
873 p.p. subincanum
874 grandiflorum
875 grandiflorum
876 grandiflorum
884 microcarpum
885 microcarpum
886 microcarpum
887 sylvestre
888 sylvestre
889 sylvestre
890 sylvestre
891 sylvestre
893 subincanum
894 subincanum
895 subincanum
896 subincanum
897 subincanum
898 subincanum
899 subincanum
900 subincanum
Observ. 2832 sylvestre
Observ. 2890 microcarpum
Matupa, E.
16690 bicolor
16733 bicolor
16840 bicolor
Mexfa, Y.
7214 bicolor
MIRANDA BastTos
68 subincanum
Mocrifo, J. M., & Sesst, M.
3618 angustifolium
3620 bicolor
3621 bicolor
s.n. angustifolium
s.n. bicolor
MONTEIRO DA CosTa
121 grandiflorum
Mutter, J. V. S.
s.n. bicolor
Patin, C.
S.n. cacao
PaTiNo, V. M.
22 subincanum
24 nemorale

115
116
117
163
169
171
171A
171B
s.n.

617
201
8.0.
8.0.

HERBARIUM

chocoense
nemorale
nemorale
grandiflorum

chocoense var. bullatum?
chocoense var. bullatum
chocoense var. bullatum
chocoense var. bullatum

speciosum
Pavon, J.
bicolor
subincanum
bicolor
sinuosum

GENTLE, Percy

3464

Puitipson, W. R., Iproso, J. M.,

bicolor

& FERNANDEZ, A.

1552

136
4340
4343
4344

4345
6575

Pires, J.

695

740

742

743

744

746

1414

8.n.
Pires, J.

5886

Pires, J. M., Nixo, T., & Sriva, A.

4339

4105
4194
6883
11112
14016
16142
s.n.

Pirrrer, H., & DurRanp, T.

3925
8536

glaucum

Pires, J. M.

sylvestre
bicolor

grandiflorum * obova-

tum

grandiflorum X obova-

tum

speciosum X _ sylvestre

simiarum

M., & Buack, G. A.

speciosum
speciosum
microcarpum
obovatum
grandiflorum
bicolor
subincanum
obovatum

M., Frées, R. L., &
Sitva, N. T.

speciosum

obovatum
PiTtier, H.
bernouillii
hylaeum?
bicolor
angustifolium
simiarum
angustifolium
angustifolium

simiarum
angustifolium

CUATRECASAS—CACAO AND ITS ALLIES 595

Pirtier, H., & Tonpvuz, A.
4074 angustifolium
Porppia, E.
18 _ bicolor
1845 obovatum
2352 p.p. subincanum
2352 p.p. obovatum
2746 p.p. obovatum
2746 p.p. bicolor
s.n. bicolor
s.n. obovatum
PoiTEau, A.
s.n. subincanum
Preuss, P.
1381 angustifolium
RANGHEL, A.
195 subincanum
Rexo, B. P.
6068 bicolor
Ricwarp, L. C.
s.n. grandiflorum
RIEDEL, L.
1373 grandiflorum
s.n. grandiflorum
RIVERO
1836 subincanum
Romero CastaNepa, R.
5405 bernouillii subsp. capil-
liferum
5500 stipulatum
s.n. bicolor
Rvufz, H., & Pavén, J.
s.n. bicolor
8.0. sinuosum
s.n. subincanum
Russy, H. H.
647 speciosum
654 speciosum
Sacort, P.
1206 velutinum
SANDEMAN, C.
2233 ~=grandiflorum
ScHoMBuRGE, R.
870 p.p. bicolor
870 p.p. obovatum
8.0. grandiflorum
Scuuutss, R. E.
3471 bicolor
3922 bicolor
6536 subincanum
6921 obovatum
8065 grandiflorum
8178 grandiflorum

8385 subincanum
12104 subincanum
Scuu.tses, R. E., BAKER,
R. E. D., & Caprera, I.
18552 subincanum
Scuuttss, R. E., & Buack, G. A.
8146 grandiflorum
Scuuttses, R. E., & Casrera, I.
14140 subincanum
15116 subincanum
17005 subincanum
17775 =obovatum
17780 microcarpum
17781 grandiflorum
18695 gileri
Scuu.tes, R. E., & Corpetro, E.
6507 speciosum
Scuuutss, R. E., & Lépez, F.
9204 grandiflorum
Scuuttess, R. E., & Siva, A.
8066 speciosum
ScHULTZE-RHONHOF
2312 glaucum
Sressé, MociNo, Casti.1o, &
MALDONADO
3618 angustifolium
3620 bicolor
3621 bicolor
SIBER
4 grandiflorum
S.n. speciosum
Sitva, J. F.
143 subincanum
155 obovatum
Sitva, A.
237 =subincanum
317 subincanum
SIQUEIROS, R.
4008 grandiflorum
Smitu, J. Donn.
6457 simiarum
7313 simiarum
7731 simiarum
SNETHLAGE, E. H.
300 grandiflorum
10044b speciosum
Spruce, R.
97 subincanum
166 sylvestre
456 speciosum
1609 bicolor
1737 speciosum
1822 grandiflorum

596 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

s.n. bicolor

8.n. speciosum

s.n. sylvestre

STANDLEY, P. C.

22317 angustifolium
36822 simiarum
37374 bicolor
37377 simiarum
79068 angustifolium
79069 bicolor
82446 bicolor

STERN, W., & CHAMBERS, K.

140 angustifolium
STEYERMARK, J. A.
44941 bicolor
49317 bicolor
STOCKDALE, J. A.
s.n. bicolor
SuCHTELEN, N. J. v.
8.0. simiarum
Tate, G. H. H.
944 subincanum
TEssMANN, G.
3433 obovatum
4079 bicolor
4115 subincanum
4928 sinuosum
5398 speciosum
s.n. subincanum
Tonpvuz, A.
4074 angustifolium
6852 simiarum
8373 simiarum
113804 bicolor
12822 simiarum
13110 bicolor
18222 simiarum
TraILu, J. W. R.
59 subincanum

60 bicolor
61 obovatum
62 glaucum
s.n. bicolor
TRIANA, J. J.
5333(-3) bicolor
s.n. bicolor
TUERCKHEIM, II. v.
7824 bicolor
Utes, E. H. G.
5030 bicolor
5637 obovatum
9609 speciosum
14448 speciosum
WEBERBAUER, A.
6245 bicolor
WEDEL, H. v.
681 bernouillii subsp. ascle-
piadiflorum
1535 p.p. bernouillii subsp. ascle-
piadiflorum
1535 p.p. Lauraceae sp.
Wicxuaw, H. A.
S.n. grandiflorum
WI.uraMs, R. O.
12121 angustifolium
WI.LuIAMs, LLEWELYN
161 obovatum
230 obovatum
1076 subincanum
1233 subincanum
2149 bicolor
2401 grandiflorum
3254 subincanum
3346 bicolor
5268 obovatum
9345 bicolor
15204 subincanum
15614 grandiflorum

Collections of Theobroma cacao L. Seen

Acosta Solis, M., 6332, 10724a.

Allen, Cyril, 880, 881.

Asplund, E., 13408, 14464, 14583,
14788.

Baker, C. F., 61, 63, 125.

Baker, R. E. D. & Cope, F. W., 15.

Banks, s.n.

Barkley, Araque, & Gomez, 410.

Bartlett, H. H., 13108.

Bartley, B. G., & Holliday, P. C., 51.

Bernoulli, G., 96, 97, 98.

Bernoulli, G., & Cario, R., 3150, 3151,
3152.

Bertero, C., 35.

Blanchet, J. §., 3, 115, 5068, s.n.

Blanco, 579.

Bonpland, A., 1102, s.n.

Box, H. E., 1536.

Brenes, A. M., 12334.

Broadway, W. E., 787, 4827, s.n.

Buchtien, O., 187, s.n.

Burchell, W. J., 9276.

Calderén, S., 107.

Chevalier, A., s.n.

Clement, B., 1931.

Collins, J. H., 15, 16.

Converse, O., 74.

Cook, O. F., & Doyle, C. B., 53, 610,
621, 622, 625, 674, 726.

Cook, O., & Gilbert, G. B., 1668, 1685.

Cook, O. F., & Griggs, R. F., 320, 321.

Cope, F. W., & Holliday, P., 83, 99,
102, 104, 105, 107, 111, 127.

Cuatrecasas et al., 2555, 7756, 7770,
13329, 13377, 25802, 25803, 25804,
25805, 26004, 26005, 26006, 26224,
26225, 26492, 26493, 26539, 26540,
26561, 26562, 26563, 26564, 26565.

Curran, H. M., 122, 163.

Dahlgren, B. E., et al., 7, 610931.

Dawe, M. T., 227.

Doustan, Dr., s.n.

Ducke, A., 1095, 12148, 23970, 23976.

Duque Jaramillo, J., 4411, 4404A.

Duss, P., 2039, 2900.

Echevarria, 866.

Emrick, G. M., 14.

Engel, s.n.

Espiritu Santo, J., 94.

Ferreyra, R., 4905.

Fredholm, A., 3117.

Frées, R. L., 20573, 20882, 21484,
21524, 23925.

Galeotti, M., 7237.

Garcia Barriga, H., 8388.

Garganta, M. de, 717.

Gentle, P. H., 1740, 3292.

Glaziou, A. F. M., 9644, 12190.

Graham, E. H., 500.

Gregg, 1774.

Haenke, T., 1533, 2301.

Hahn, 112.

Harvey, D., 5215.

Heller, A. A. & Heller, 726.

Heyder, H. M., 35.

Hitchcock, A. E., 449.

Hinton, G. B., 7531.

Hodge, W. H., 6715.

Hohenacker, R. F., 39.

Holton, I. F., 765.

Hostmann, W. R., 1, 440.

Huber, H., 1392, 4392.

Idrobo, J. M., et al., 784, 940.

Isert, 87.

Jack, J. G., 4334.

Jovert, Dr., 542.

Jungner, J. R., 79.

Kappler, A., 1636, s.n.

Karsten, G., s.n.

Kellerman, W. A., 4842, 5565, 6045.

Kidder, N. T., s.n.

Killip, E. P. & Smith, A. C., 29434,
3022, 33603.

Klug, G., 926, 2938.

Krebs, s.n.

Krukoff, B., 4736, 10661.

Kuntze, O., s.n.

Lehmann, F, C., 5641.

597

598 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Lemée, A., s.n.

Leén, Jorge, 2001.

Leonard, E. C., 8550, 9249, 9252.

Levy, P., 1.

Liebmann, F., 586, 15078, 15079.

Little, E. L., 6483, 8624.

Little & Little, 9574.

Llave, P., s.n.

Lloyd, 1128.

Luetzelburg, Ph. v., 2585.

Lundell, C. L., 2799.

Macbride, J. F., 5278.

Martius, C. E. P., 866, 867, 868, 869.

Matthews, A., 18, s.n., 1653.

Mell, C. D., 29.

Mexia, Y., 6399.

Miranda, F., 6644, 9299.

Molina, A., 2346.

Mulford, 953.

Myers, J. G., 5829.

Nadeaud, J., s.n.

Nelson, E. W., 2490.

Oca, N. de, 47 bis.

Orcutt, C. H., 4250.

Pav6on, J., 623, s.n.

Pérez Arbeldez, E., 686.

Philipson, W. R., et al., 1565, 1569.

Pierre, 119.

Pittier, H., 3927, 6615, 11934, 11953,
8.n.

Poeppig, E., s.n.

Poiteau, A., 209, 211, 214, 217.

Proctor, G. R., 18348.

Raunkiaer, Ch., 2525, 2864.

Reko, B. P., 3393, 4720.

Richard, L. C., s.n.

Ricksecker, s.n.

Rodin, R. J., 594.

Rose, J. N., 21991.

Ruiz, H. & Pavén, J., s.n.

Rusby, H.H., 655.

Ryan, J., s.n.

Sagot, P., 52, s.n.

Sagra, R. de la, 118.

Salzmann, s.n.

Sandeman, C., 3382.

Sargent, F. H., 333.

Schipp, W. A., 178, 419, s.n.

Schomburgk, R., s.n.

Schott, A., s.n.

Schultes, R. E., et al., 3309, 5858b,
6117, 6667a, 8371, 8524, 8604.

Scolnick, R., et al., 19An526.

Sessé et al., 3619.

Shafer, J. A., 3428.

Shannon, W. C., 147.

Shuttleworth, 1250.

Sintenis, P., 315, 6370.

Sneidern, K., A1324.

Splitzberger, F. L., 1097.

Standley, P., et al., 19430, 21640,
22699, 25693, 27968, 29673, 30479,
31104, 31384, 44954, 45715, 48644,
52877, 54143, 54879, 55742, 79081,
82445, 91139,

Stern, W. & al., 171.

Stevenson, J. A., 116, 3631.

Steyermark, J. A., 45950, 49218,
54947.

Swartz, 900, s.n.

Tessmann, G., 3036.

Theresa, Prinz., v. Bayern, 8.n.

Thiebout, C., 501.

Thieme, C., 5156.

Tonduz, A., 6984, 9928.

Traill, J. W. R., 58, 63.

Triana, J. J., 5333, s.n.

Ule, E. H. G., 5032.

Urban, I., 315, 6370.

Williams, R. 8., 806.

Williams, Llewelyn, 148, 2105, 2349,
3510, 4160, 5278, 8457, 8981, 9021,
9022, 9346, 11718, 15869.

Wilson, P., 162.

Wright, C., et al., 23X, 2610.

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CONTR. NAT. HERB. VOL. 35, PT. 6 CUATRECASAS—PLATE

Theobroma sylvestre Mart., lectotype at Munich (photo FM 19644).

GS0-695—64 16

CONTR. NAT. HERB. VOL. 35, PT. 6 CUATRECASAS—PLATE 2

Theobroma speciosum Willd. ex Spreng., isotype formerly at Berlin-Dahlem; identical with
the holotype in the Willdenow Herbarium (photo FM 9640).

CONTR. NAT. HERB. VOL. 35, PT. 6 CUATRECASAS —-PLATE 3

Theobroma speciosa Willd. ex Spreng. isotype of 7. quinguenervia Bern., Spruce 1737 at

Berlin-Dahlem (photo FAT 9639)

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CONTR. NAT. HERB. VOL. 35, PT. 6 CUATRECASAS PLATE 8

Theobroma grandiflorum (Spreng.) Schum, lectotype of 7 macrantha Bern. and neotype of
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INDEX

Page references to descriptions or definitions in Boldface.

Synonyms in [talics.

Tribal origins of native names in parentheses

a-ba-ka-ra (Makuna), 566
abekard (Makuna), 566
Abroma, 388, 389, 390, 391, 392, 394,
395, 432, 433, 435, 435 (fig.), 437,
445
augusta, 435 (fig.), 443 (fig.), 445,
588
augustum, 390
fastuosum, 390
nitida, 588
Acaju, 387

aligator, 399

alligator, 399, 400, 402, 518, 516

Amazonian forastero, 409, 515

Ambroma, 388

amelonado, 397, 399, 400, 409, 413,
501 (fig.), 510, 515, 516, pl. 6

amelonado amarillo, 394, 402

amelonado colorado, 394, 402

Amygdalae similis Guatimalensis, 384

Amygdalis similis guatimalensis, 495

Amygdalus, 383

Andropetalum, sect., 421, 422 (fig.),
425, 431 (map), 451, 452 (key),
579

angoleta, 399, 501 (fig.), 508, 516

ad (Makuna), 463

Arriba, 400

Arvor cacavifera Americana, 384, 495

a-si-ya-ee (Piratapuya), 566

Assonia, 388, 432

Attalea, 476

Avellana Mexicana, 384, 495

Ayenia, 388, 389, 391, 394, 395, 396,
414, 432, 433

bacafto, 574, 575

bacafto de monte, 547

bacao, 400, 462, 463, 465, 541, 574, 575

bacao de monte, 545, 546, 574, 575

ba-dja-na-hoo (Makuna), 556

bahia, 400

balam (Kekchi), 462, 464

balamati, 462, 464

680-695 6418

balao, 400
bawk (Maku), 566
bawk-pom (Maku), 556
beira de assahyzal, 570
Bertholletia, 476
bicco, 488, 489
bik (Térraba), 550
bizoya, 512
boldo, 445
Bombacaceae, 391, 393
béo-e (Mirana), 524
Bouchi-cacao, 485, 486
Brotobroma aspera, 588
Bubroma, 388, 390, 396, 400
grandiflorum, 390, 396, 552,
556
Guazuma, 390
polybotryon, 390
tomentosum, 390
Bubroma, sect., 396, 400, 405, 409, 451,
467, 517, 526
Bubroma subsect. Glossopetalum, 405,
526
Bubroma subsect. Oreanthes, 405, 467
Bubroma subsect. Telmatocarpus, 405,
517
Buettneria, 392, 394
Buettneriaceae, 442
Buettnerieae, 392, 394
Buettneriées, 437
Buttneria, 388
Biittneria, 389, 395, 396
Biittneriaceae, 391
Biittnerieae, 391, 394, 395, 396
Biittnerinae, 396
Byttneria, 388, 432, 433, 435 (fig.), 437
arguta, 435 (fig.)
Byttneriaceae, 391, 432, 4338, 437
Byttnerieae, 391, 432, 433, 437
Byttneriées, 433
Byttnerinae, 433
eabeea de Umbi, 560, 563
cabeea de urubd, 524, 525, 560, 562, 563
607

555,

608

cabosse, 428
cacado de monte, 512
cacahoacentli, 383
cacahoacuahuitl (Nauhatl), 512
cacahoaquahuitl, 383
cacahoatl, 379, 383, 512 (Nauhatl)
cacahuatl (Nauhatl), 379, 512
cacafto de monte, 574, 575
Cacao (genus), 383, 384, 385, 386, 387,
388, 449, 495
album Peruvianum, 570
bicolor, 389
guianensis, 387, 388, 390, 396, 495,
564, 586, 587, pl. 11
guyanensis, 393
minor, 385
minus, 388, 495, 511, 513
sativa, 387, 388, 401, 495, 510
sylvestris, 387, 388, 391, 475, 562,
564, 569, pl. 10
Theobroma, 495, 511
Cacao, sect., 393, 406, 409, 495
Cacao, subsect., 405, 495
cacao (common name), 512
cacao (Kofdn), 466
cacao amelonado, 512
cacao azedo, 482, 483
cacao azul, 474, 475, 476
cacao biaro, 482
cacao blanco, 400, 462, 463, 466, 566
cacao bravo, 463, 466, 525
cacao calabacillo, 512
cacao ceniza, 566
cacao chuncho, 405, 512
cacao claro, 482
cacao complex, 406
cacao criollo, 394, 404, 512, 513
cacao d’Anta, 466
cacao de Castilla, 463
cacao del pafs, 400
cacao de la India, 533
cacao de macao, 560, 563
cacao de mico, 533, 534, 550, 551
cacao de mono, 550, 551
cacao de monte, 493, 524, 541, 543, 545,
546, 566, 569, 574, 577
cacao de monte bravo, 493
cacao do matta, 482
eacao do matto, 482
cacao dulce, 404, 512
cacao forastero, 394, 512
cacao grande de monte, 545
cacao indio, 541

INDEX

cacao lagarto, 3938, 400, 505, 512, 513

cacao malacayo, 462, 464

cacao marraco, 463, 466

cacao meco, 533, 534

cacao rana, 474, 483, 524, 525, 566

cacao sacha, 482, 483

cacao sauvage, 485, 486

cacao silvestre, 463, 465, 512, 533, 566,
568, 572, 582

cacao trinitario, 512

cacao-hu, 474, 476

cacaohy, 482, 525

cacao-i, 482

cacaofllo, 482

cacaorana, 483

cacao-rana, 482, 569

cacaotlquahuitl (Nauhatl), 512

cacao-u, 482

cacao-y, 482

cacau, 482, 562

cacau azul, 474, 475, 489

cacau baju, 463, 466

cacau bravo, 474, 476, 524, 525

cacau do Pert, 463, 467

cacau rana, 474

cacau silvestre, 516

cacauatl, 408

cacauf, 474, 482, 524

cacau-i, 482, 483

cacau-rana, 482

cacaurana, 475, 524

cacauu, 474, 482

cacava quahuitl, 384

cacavate, 384

cachu azul, 489

caco, 12

cahequa (Tarascin), 512

calabacillo, 394, 397, 398, 399, 400, 402,
403, 406, 407, 414, 502 (fig.), 510,
515, 516, 517

calabacillo amarillo, 394, 397, 401, 402

calabacillo colorado, 394, 397, 401, 402

caocauatzana (Zoque), 512

caracas, 399

carupano, 399

carupano grande, 399

carupano legitimo, 399

carupano mestizo, 399

carupano parcho, 399

carupano taparito, 399

carvu (Kabekara), 462

cauca, 400

chocoatl, 379

INDEX

chocolate, 543

chocolate de monte, 493, 522, 543, 547,
572, 574, 575

chocolatillo, 482, 483

chuct, 488

chudechu (Otomi), 512

coca mono, 533, 534

cocoa, 512

cojén de toro, 400

Commersonia, 389, 391, 392, 394, 395,
396, 432, 433, 437

copuaf, 566

copu-afi, 560, 563

copuassti, 556, 557, 558

creole, 398

Crescentia cujete, 463, 556

criollo, 383, 386, 394, 396, 397, 398, 399,
400, 401, 402, 403, 405, 406, 407,
408, 409, 414, 415, 506, 507, 508,
509, 510, 513, 516, 517

criollo amarillo, 396, 400, 401

criollo caldas, 498 (fig.)

criollo caracas, 397

criollo colorado, 396, 401

criollo legitimo, 400

criollo mestizo, 400

criollo Venezuela, 507

cucuh, 512

cuculat, 512

cu-lu-hu (Chok6), 462, 465

cumacaco, 393, 404

cumajé (Chok6), 547

cumald, 566, 570

cundeamor, 394, 397, 399, 516

cundeamor vars., 402

cundeamor legitimo, 400

cundeamor verugoso amarillo, 394, 396

cundeamor verugoso colorado, 394, 396

cundiamor, 498 (fig.), 425 (fig.), 427
(fig.), 508

cupai-acd, 556

cupasst, 556

cup do matta, 566

cupt do matto, 556, 569

cupt-asstrana, 566

cupt-assuy, 566

cupt-curta, 560, 563

cupuact, 466, 556, 557, 558

cupti-act, 557

cupuahy, 566, 570

cupua-f, 463, 466, 570

cupuarana, 566, 569

cupuasst, 463, 466

609

cupu-asst, 556, 557, 558
cupuasti, 556

cupul, 566, 570

cupuhy, 483, 566, 569
cupurana, 482, 560, 563, 583
cupu-uasst, 556, 557

cupuy, 482, 570

cupuy do igapé, 566, 570
cupuyh, 482

curupano grande, 400
cushta, 533

deghy (Otomi), 512
Dicarpidium, 395

Diosma, 387

dolé (Doraske), 512
Dombeya, 388, 432
Dombeyaceae, 391
Dombeyeae, 395
Dzug-mang-ud (Brunka), 550
erefa (Guatuso), 462
Eriolaenae, 391

Eriolaenées, 437
Eutheobroma, sect., 395, 396, 400, 405,

409, 458, 495
Eutheobroma, subsect. Cacao, 405, 495
Eutheobroma, subsect. Rhytidocarpus,
405, 458

farinha, 483

forastero, 394, 396, 397, 398, 399, 400,
402, 403, 405, 406, 407, 408, 409,
414, 506, 515, 516, 517

forastero amelonado, 401

forastero amelonado amarillo, 397

forastero amelonado colorado, 397

forastero cundeamor, 401

forastero ordinary amarillo, 396

forastero ordinary colorado, 396

forastero vars., 402

Forcipomyia, 432

Frankliniella parvula, 408, 432

Glossopetalum, sect., 393, 409, 417 (fig.),
421 (fig.), 422 (fig.), 425, 431
(map), 438, 451, 452 (key), 453
(key), 526, 580

Glossopetalum subsect., 405, 526

Glossostemon, 389, 391, 392, 394, 432,
433

bruguieri, 443, 445

guandbana, 556

guayaquil cacao, 399

Guazuma, 385, 387, 388, 389, 390, 391,
392, 394, 395, 396, 411, 432, 433,
435, 435 (fig.), 445, 555

610

Guazuma—Continued
grandiflora, 552
polybotrya, 443, 445
tomentosa, 435 (fig.), 588
ulmifolia, 389, 446, 555, 588
hé-ha (Tanimuka), 463
heé-a (Maku), 463
Helicteraceac, 591
Helictereae, 399
Heritiera, 442
Hermannia, 388, 395
Hermannieae, 395
Herrania, 392, 393, 394, 395, 396, 403,
405, 407, 408, 409, 411, 412, 413,
432, 433, 435 (key), 457, 444,
445
albiflora, 392, 588
balaénsis, 588
camargoana, 444, 588
cuatrecasana, 425 (fig.), 437 (fig.),
444
guianensis, 484, 485, 486
quyanensis, 484
laciniifolia, 392, 588
mariae, 401, 408, 411, 412, 413, 444,
588
nitida, 409, 511, 588
paraensis, 482
pulcherrima, 392, 588
pulcherrima v. pacifica, 437 (fig.),
443 (fig.), 444
purpurea, 588
Herrania, sect., 395, 396, 400
Hugonia, 387
judromajé (Choks), 547
kao-kra (Brunka), 512
kajo (Guatuso), 512
kako (Mixe), 512
kau (Tiribf), 512
kicob, 512
kicou, 512
Kleinhovia, 388, 432
kno (Penonomé), 512
ko (Térreba), 512
ké6o (Brunka), 512
kua (Guaimf), 512
ku-gin (Térraba), 550
kuk (Rama), 512
kréaku (Guatuso), 550
largarto, 400, 425 (fig.), 500 (fig.), 506,
514, 516
largarto amarillo, 514
largato rojo, 514

INDEX

la-na-pee-ta-ma-ca-la-chu-na-ni (Yaku-
na), 463
Lasiopetaleae, 395
Leptonychia, 394, 395, 452, 453, 437
Licania, 587
alba, 588
venosa, 588
liso amarillo, 394
liso colorado, 394
macambo, 463, 467
machala, 400
maga (Barasana), 556
mah-we-re (Yukuna), 566
majambo, 463
Malvaceae, 393, 395
Malvales, 392, 437
ma-oo-hee-rée (Kabuyarf), 560, 566
maraca, 463
marraco, 465
Marasmius perniciosus, 525, 570, 605
mate, 463, 556
ma-wé-roo-da (Kuripaka), 566
Maxwellia, 394
mazorcea, 428
meeacahoatl, 383
Melhania, 388, 432
Melochia, 395
me-tré-ree-moo-ee (Karihona), 524
Monilia roreri, 522
monkey cocoa, 534
mountain cacao, 464
nacional Ecuador, 508, 517
najambu, 463, 467
Nicaraguan criollo, 402
no-térree-ka (Tanimuka), 566
nunisup (Rama), 550
fiee-aw (Tanimuka), 556
Oreanthes, sect., 393, 409, 421 (fig.),
422 (fig.), 425, 431 (map), 438,
451, 452 (key), 467
Oreanthes, subsect., 405, 467
pacxoc, 512
padama (Arekuna), 566, 569
pako kakao, 577
patachtli, 383
pataiste, 462, 465
patas, 463, 466
patasht, 462, 464
patashte, 462, 463, 464
pataste, 462, 464
pataste de sapo, 462, 464
pataste simarr6én, 462, 464
patatle, 462

INDEX

pataxte, 462
pazoli, 464
pec (Pokonchi), 462
Pentapetes, 432
petaste, 462, 464
petaxte, 462, 464
Philippodendrae, 391
Pistachia, 384
Polyadelphia Pentandria, 432
porcelaine, 399
porcelaine criollo, 516
porcelaine Java criollo, 514
poo-hoo (Barasana), 566
quauheacahoatl, 383
ree-ka (Tanimuka), 566
Rhytidocarpus, sect., 393, 409, 421
(fig.), 422 (fig.), 425, 488, 451,
452 (key), 458
Rhytidocarpus, subsect., 405, 458
Rulingia, 391, 394, 395, 396, 432, 433,
437
— sambito, 399, 410
sangre de toro, 399
saparén (Estrella), 462
skar-ub (Bribrf), 462
Sapokaia brasiliensis, 476
Scaphopctalum, 394, 395, 422, 433, 437
searbo (Bribrf), 462
soré (Bribf), 533
Sterculiaceae, 391, 392, 393, 395, 396,
432, 433, 437, 442
Sterculieae, 395, 437, 442
Surinam, 399
Telmatocarpus, sect., 393, 409, 42], 422
(fig.), 427, 438, 452 (key), 453
(key), 517, 518 (map)
Telmatocarpus, subsect., 405, 517
teta negra, 550
Thalamiflorae, 392
Theobroma, 435 (key), 449, 452 (key),
455 (key)
alba, 393, 564, 570, 587, 588
albiflorum, 588
album, 395, 398, 405
angustifolia, 390, 391, 398, 409, 415
angustifolium, 395, 398, 404, 405,
407, 414, 427 (fig.), 431 (map),
440, 441, 443, 446, 454 (key), 456
(key), 502 (fig.), 526, 528 (fig.),
531, 533, 539 (fig.), 575, 583, 584,
585
angustifolium, * cacao, 583
angustifolilum, * mammosum, 583

611

Theobroma—Continued

asclepiadiflorum, 408, 415, 489, 490,
491, 492, 493

aspera, 403, 588

augusta, 588

balaénsis, 588

bernouillii, 403, 405, 408, 409, 414,
431 (map), 440, 443, 453 (key),
457 (key), 488, 489, 491, 492
(key), 493

bernouillii subsp. asclepiadiflorum,
431 (map), 469 (map), 472 (fig.),
473 (fig.), 477 (fig.), 492 (key),
493

bernouillii subsp. bernouillii, 431
(map), 469 (map), 472 (fig.), 477
(fig.), 492 (key)

bernouillii subsp. capilliferum, 419
(fig.), 427 (fig.), 429 (fig.), 431
(map), 469 (map), 472 (fig.), 473
(fig.), 477 (fig.), 481 (fig.), 492
(key), 493, pl. 5

bicolor, 383, 389, 390, 391, 392, 393,
395, 396, 398, 400, 401, 403, 404,
405, 407, 409, 411, 412, 413, 414,
418, 419 (fig.), 425 (fig.), 428,
437 (fig.), 488, 440, 443, 446, 452,
456 (key), 458, 459 (fig.), 460
(map), 479 (fig.), 522, 537 (fig.),
583, 585, 605

bicolor * cacao, 411

bicolor X cacao, 584

cacao, 417 (fig.), 419 (fig.), 421
(fig.), 425 (fig.), 427 (fig.), 455
(key), 481 (fig.), 495, 498 (fig.),
499 (fig.), 501 (fig.), 503 (fig.),
512 (key), 508

cacao subsp. cacao, 494 (map), 497
(fig.), 499 (fig.), 512 (key), 513

cacao subsp. cacao fma. cacao, 512
(key)

cacao subsp. cacao fma. lacando-
nense, 502 (fig.), 512 (key), 514

cacao subsp. cacao fma. leiocarpum,
502 (fig.), 506, 512 (key), 514, 516

cacao subsp. cacao fma. pentago-
num, 497 (fig.), 500 (fig.), 512
(key), 513, 516

cacao subsp. leiocarpum, 413, 496,
514, 515

cacao subsp. pentagona, 496, 513

cacao subsp. sativa, 496

612

Theobroma—Continued

cacao subsp. sphaerocarpum, 494
(map), 497 (fig.), 501 (fig.), 502
(fig.), 513 (key), 515, pl. 6

cacao fma. leiocarpum, 408, 496,
515

cacao fma. pentagonum, 413, 425
(fig.), 427 (fig.)

cacao var. leiocarpa, 407, 496, 515

cacao var. letocarpum, 514

cacao var. typica, 407, 496, 513

cacao var. typica X v. leiocarpa,
407, 496

cacao X mammosum, 584

cacao X microcarpum, 411

cacao X obovatum, 411

cacao X simiarum, 584

calodesmis, 408, 412, 413, 414, 415,
486, 488

Camargoanum, 413

camargoanum, 588

canumanense, 431 (map), 455 (key),
457 (key), 535 (map), 577, 578,
579 (fig.)

capillifera, 414

capilliferum, 410, 412, 446, 489, 490,
491, 492, 493

caribaea, 390, 495, 511

celtifolia, 389, 588

chocoense, 412, 421, 421 (fig.), 431
(map), 454 (key), 458 (key), 529
(fig.), 535 (map), 538 (fig.), 543,
545, 546, 549 (fig.), 550, 551, 585

chocoense var, bullatum, 546

cirmolinae, 408, 412, 414, 415, 421
(fig.), 422 (fig.), 423 (fig.), 431
(map), 446, 453 (key), 458 (key),
528 (fig.), 531 (fig.), 534, 535
(map), 536 (fig.), 538 (fig.), 539
(fig.), pl. 7

cordata, 460, 467

ferruginea, 393, 409, 414, 564, 569,
570, pl. 9

ferrugineum, 406, 570

foliis integerrimis, 495

fossilium, 587

gileri, 411, 412, 415, 425 (fig.), 427
(fig.), 430, 453 (key), 455 (key),
503 (fig.), 517, 518 (map), 519
(fig.), 520 (fig.), 521 (fig.), 522

glauca, 392, 393, 409, 414

INDEX

Theobroma—Continued

glaucum, 391, 395, 398, 405, 419
(fig.), 431 (map), 448, 443 (fig.),
444, 451, 453 (key), 457 (key),
471 (fig.), 473 (fig.), 475 (map),
477 (fig.), 481 (fig.), 486, 488, 493,
pl. 4

grandiflora, 409, 415

grandiflorum, 390, 395, 398, 405,
407, 408, 411, 412, 413, 414, 421
(fig.), 425 (fig.), 426 (fig.), 428,
429 (fig.), 431 (map), 438, 440,
443, 446, 454, (key), 457 (key),
474, 526, 533 (fig.), 538 (fig.),
552, 552 (map), 553 (fig.), 585,
pl. 8

grandiflorum X subincanum, 583

grandiflorum X obovatum, 583

Guazuma, 386

gquazuma, 588

guianense, 388, 389, 390, 480, 586,
587

guianensis, 390

hastata, 409, 511, 588

hylaeum, 430, 431, 431 (map), 455
(key), 458 (key), 502 (fig.), 553
(fig.), 564 (map), 570, 572, 574,
575

integerrima, 389, 495, 511

Kalagua, 398, 400, 414, 496, 584,
585, 586

laciniifoliwm, 588

laeve, 514

leiocarpa, 393, 410, 446, 495, 514

leiocarpum, 395, 404, 405, 406, 407,
408, 409, 411, 414, 504, 505, 506,
508, 510, 511, 514, 515

macrantha, 393, 552, pl. 8

macro nthum, 396, 555, 556

mammosa, 415

mammosum, 411, 414, 423, 426,
(fig.), 431 (map), 452, 456 (key),
535 (map), 538 (fig.), 567 (fig.),
573 (fig.), 580, 581, (fig.) 583

mammosum X simiarum, 583

mariae, 588

Martiana, 391

Martti, 395, 398, 405, 467, 474

microcarpum, 391, 393, 395, 398,
401, 405, 407, 408, 409, 411, 412,
413, 414, 415, 430, 438, 440, 441,
443, 444, 446, 453 (key), 456 (key),
503 (fig.), 517, 518 (map), 519

INDEX 613

Theobroma—Continued

microcarpum—Continued
(fig.), 521 (fig.), 522, 523, 537
(fig.)

montana, 588

nemorale, 411, 412, 414, 421 (fig.),
431 (map), 455( key), 457, 473,
(fig.), 528 (fig.), 529 (fig.), 531,
564 (map), 567 (figs.), 572, 573
(fig.), 575

nemoralis, 415

nitida, 393, 467, 469, 474, 475

nitidum, 474, 588

obovatum, 393, 396, 401, 408, 409,
411, 412, 413, 414, 415, 431
(map), 438, 440, 441, 446, 454
(key), 456 (key), 533 (fig.), 537
(fig.), 552 (map), 553 (fig),
559, 561, 562, 583

obovatum X subincanum, 583

ovatifolia, 390, 391, 393, 460

ovatifolium, 390, 396, 398, 400, 463

Patastle, 463, 464

pentagona, 393, 402, 403, 410, 414,
446, 495, 510, 513

pentagonum, 395, 398, 400, 405, 406,
409, 410, 411, 414, 504, 505, 506,
508, 509, 510, 511, 514

pulcherrimum, 588

purpureum, 403, 407, 588

quinquenervia, 393, 476, pl. 3

quinquenervium, 396, 480, 483

sagittata, 409, 410, 511, 588

saltzmaniana, 393, 410

Salizmanniana, 496

saltzmannianum, 504, 511

salzmannianum, 395

sapidum, 407, 496, 511, 513

sativa, 401, 410, 496

sativa var. leucosperma, 410, 496,
511, 513

sativa var. melanosperma, 410, 496,
511

sativum, 391, 510, 511

silvestre, 552

simiarum, 397, 398, 400, 405, 407,
409, 412, 414, 415, 425 (fig.), 426
(fig.), 431 (map), 446, 454 (key),
458 (key), 531 (fig.), 538 (fig.),
545, 547, 549 (fig.), 550, 561
(fig.), 583, 584, 585

sinuata, 575

Theobroma—Continued

sinuosum, 401, 409, 431 (map),
454 (key), 457 (key), 535 (map),
573 (fig), 575, 578, 588, pl. 12

speciosa, 393, 409, 482

speciosum, 390, 391, 395, 396, 398,
401, 405, 407, 408, 411, 412, 413,
414, 425, 427, 431, 425 (fig.),
427 (fig), 431 (map), 438, 446,
452 key, 457 key, 467, 468
(fig.), 471 (fig.), 473 fig., 474,
475 map, 476, 479 (fig.), 408,
481 (fig.), 482, 484, 485, 488,
552, 587, pl. 2, pl. 3.

speciosum var. coriaceum, 401, 443,
444, 459 (fig.), 476

speciosum var. quinquenervia, 476

speciosum var. Spruceana, 467

speciosum Xsylvestre, 583

sphaerocarpa, 401, 403, 410, 496,
515

sphaerocarpum, 404, 498 (fig.), 505,
510

spruceana, 393, 467, 469

spruceanum, 396, 405, 408, 412,
413, 474, 476

stipulata, 415

stipulatum, 411, 412, 431 (map),
453 key, 458 key, 528 (fig.),
537 fig., 546, 550, 551, 531 (fig.),
535 (map), 539 (fig.), 541, 546,
585

subincana, 393

subincanum, 387, 390, 395, 396,
398, 401, 405, 407, 408, 412,
413, 414, 430, 431 (map), 438,
440, 455 (key), 458 (key), 482,
521 (fig.), 533 (fig.), 537, 543,
553 (fig.), 562, 563, 564 (map),
566, 567 (fig.), 569, 572, 574, 583,
586, 587, pl. 9, pl. 10, pl. 11

sylvestre, 390, 395, 398, 401, 405,
412, 422 (fig.), 431 (map), 437
(fig.), 438, 440, 441, 452 (key),
457 (key), 467, 468 (fig.), 469
(map), 474, 479 (fig.), 559, 562,

pl. 1

sylvestris, 391, 393, 409, 415, 474,
475, 564

tessmannii, 406, 414, 577, 578, 579,
pl. 12

Tessmannii, 575

614

Theobroma—Continued
tomentosa, 396, 588
undulata, 409, 588
velutina, 409
velutinum, 404, 42! (fig.), 431
(map), 452 (key), 456 (key), 459
(fig.), 471 (fig.), 478 (fig.), 479
(fig.), 484, 485, 586, 587
Theobroma, sect., 421 (fig.), 422 (fig.),
427, 488, 451, 452 (key), 495
Theobroma sect. Andropetalum,
(key), 579
Theobroma sect. Cacao, 495
Theobroma sect. Glossopetalum, 452
(key), 453 (key), 526
Theobroma sect. Oreanthes, 452 (key),
467
Theobroma sect. Rhytidocarpus,
(key), 458
Theobroma sect. Telmatocarpus, 452
(key), 453 (key), 517
Theobroma sect. Theobroma, 452 (key),
495
Theobrominae, 396, 433
tiger, 463
Tilia, 388
tlalcacahoatl, 383
tlapal, 408
Tlapalcacauatl, 408
too-soo (Yauna), 566
Toxoptera aurantii, 408, 432

452

452

INDEX

Tribroma, 403, 449

Tribroma bicolor, 403, 460, 463
Trinidad criollo, 402

trinitario, 399, 400, 415, 508, 517
Trinitario amargo, 400
Triopteris, 387

tsiru, (Cabécara), 513

tsird, (Bribri) 512

tsird-kuru, (Cabécara) 512
tutuma, 463

uchpa-cacao, 566, 570

uerba (Térraba), 462

Uirub (Bribrf), 550

Uir-ub (Bribri), 550
Urubti-acaim, 560, 563
Venezuelan criollo, 402
wa-be-ga-ra (Desano), 566
wa-be-ka-ra (Siriano), 566
wah-pek-la (Tukano), 566
wa-k6é (Kubeo), 566

Waltheria, 395

wariba, 463

Wasmannia auropunctata, 408, 432
wild cacao, 512, 513, 570
win-cheek (Puinave), 560, 566
win-cheek-choo-ai (Puinave), 556
xochicacahoatl, 383

xocoatl, 379

yagabizoya (Reko), 512
vurac-cacao, 566, 570

U.S. GOVERNMENT PRINTING OFFICE:1964