FOUNDATION VAN EEDENFONDS

cf o Royal Tropical Institute, Amsterdam

FLORA OF SURINAME
EDITED BY

J. LANJOUW and A. L. STOFFERS

VOL. V. PART I

MORACEAE
BY

C.C. BERG and GORDON P. DEWOLF

URTICACEAE
BY

M.J.M. DE ROOIJ

LEIDEN
E. J. BRILL
1975
 MORACEAEt

BY

C. C. BERG2 AN D GORDON P. D EWO LF3

Dioecious or monoecious laticiferous trees (or herbs). Leaves alternate

(or opposite) with pinnate or palmate (or radiating) nerves, entire or
incised, stipules free or fused . I nflorescences uni- or bisexual, basically
cymose, mostly condensed into spikes, heads, or discoid to globose or
urceolate receptacles. Flowers small, unisexual, perianth basically with 2
whorls of 4 (often more or less) free or connate tepals, or sometimes
absent, basically 4 ( often less) epitepalous stamens, straight or inflexed
before anthesis, anthers 2-celled , dehiscing longitudinally, ovary one,
free or adnate to the perianth, style one, with 2( - 1 ) stigmatic arms, or one
capitate to peltate to sublingulate stigma (Conocephaloideae), ovule one,
anatropous, (sub)apical and pendulous or (sub)orthotropous and
(sub)basal (Conocephaloideae); fruit usually surrounded by the enlarged
perianth, seed surrounded by a solid endocarp, with or without
endosperm, embryo various.

D i s t r i b u t i o n : About 1 300 species, mainly tropical.

GENE RA L R E M A R K S

I n S urinam the Moraceae are represented b y 1 4 indigenous genera

with 56 species. Following the classification of Corner ( 1 962) the Moreae

1 The N etherlands Foundation for the Advancement of Tropical Research

(WOTRO) supplied C. C. Berg a grant to study material in some European herbaria.
Except for the contributions of G . P . DeWolf, the E nglish text has been corrected by
Dr. N . G . Bisset, Department of Pharmacy, University of London, Great Britain.
z I nstitute for Systematic Botany, U trecht.
J Arnold Arboretum, Jamaica Plain, Mass . , U . S . A . who contributed the treat­
ments of Ficus.
1 74 MORACEAI::
----- --��--�-------�

are represented by Clarisia and Sorocea (each with 2 species) ; the

Artocarpeae by Bagassa (2 species) ; the Bros imeae by Brosimum (5
species) and Trymatococcus (3 species); and the Olmed ieae by
Pseudo/media (one species), Perebea (2 species), Maquira (2 species),
He/icostylis (2 species), and Naucleopsis (2 species) . Ficus (Ficeae) is
represented by 18 species in S urinam. Dorstenia (Dorstenieae) has not
(yet?) been found in Surinam; a single specimen from French Guiana has
been seen.
Cecropia ( in Surinam 4 species), Coussapoa (in Surinam 4 species),
and Pourouma (in Surinam 7 species) were included in the subfamily
Conocephaloideae in Engler's system of the M oraceae ( 1 889), along with
some palaeotropical genera. This group differs from the other M oraceae
in several characters. One of the most important d ifferentiating features
is the (sub)orthotropous (sub)basally attached ovule, and probably in
connection with this the stigmatic portion of the style is not basically
bifid, as in other M oraceae. As regards these features the Conocephaloi­
deae are similar to the U rticaceae. In a study of the palaeotropical genus
Poiki/ospermum Chew Wee-Lek ( 1 963) proposed the transfer of
Cecropia and Coussapoa, both with orthotropous basally attached
ovules, to the U rticaceae and the retention of Pourouma, with subortho­
tropous subbasally attached ovules. in the M o raceae. Although
Pourourna d iffers distinctly from Cecropia and Coussapoa in several
characters of the flowers, placing these genera in separate families seems
unjustified in view of the similarities in morphological and anatomical
features. Corner ( 1 962) transferred the whole subfamily Conocephaloi­
deae to the U rticaceae on account of the position of the ovule, the shape
of the style, and some anatomical characters (cf. Renner, 1 907) . But it
appears to me that, by transferring this problematical group from the
M oraceae to the U rticaceae, only the problem is moved and that, to come
to a satisfactory positioning of this group within the system, it will be
necessary to make a thoro ugh comparative study of the U rticaceae and
the M o raceae. M o reover, it will be essential to consider the question,
whether it is correct to regard the two families as distinct . In spite of the
problematical position of Cecropia, Coussapoa, and Pourouma, I prefer
 MORACEAE 175

to follow the traditional concept and at this moment to deal with them as
part of the M oraceae.
From the fact that recent collections have revealed the occurrence of
species new for Surinam, it may be concluded that still more species
which are new for S urinam will be found on further botanical explora­
tion. A few species, expected on account of their assumed area of distri­
bution are also included in this paper.

PHYTOG EOG R A P H I C A L R E M A R K S

Some more or less roughly distinguished elements may b e recognized

in the Moraceae flora of Surinam. Firstly, a group of species which is
apparently (almost) confined to the Guiana shield region. To this group
belong, e.g. Naucleopsis guianensis, Ficus lanjouwii, F. erratica, Coussa­
poa microcephala, and Trymatococcus oligandrus. T. oligandrus has a
very small area; it is known only from French Guiana and eastern
Surinam; Ficus lanjouwii is known only from S urinam. A second group
comprises species which range from the L ower Amazon Basin ( Pani) to
the Guianas. Of this group may be mentioned Helicostylis pedunculata,
Maquira guianensis, Pourouma laevis, and Trymatococcus paraensis.
Some of them have not (yet?) been found in Guyana (British Guiana);
others seem to range only to the eastern part of Surinam. A third group
contains species which are more widespread than those belonging to the
other two groups. They occur at least in the U pper Amazon Basin and the
western part of the Guiana shield region, e . g. Sorocea muriculata, while
others are very widespread , like Brosimum guianense, B. lactescens,
Ficus citrifolia, and F. maxima. Considering also morphological resem­
blances, there seems in the M o raceae flora of Surinam to be a western
element (connected with the flora of the U pper Amazon Basin, and
Venezuela) and an eastern element (connected with the flora of the
Lower Amazon Basin) .
M ost Surinam M oraceae are true forest components. Cecropia
species, however, are distinct exceptions. The presence of Pourouma
1 76 MORACEAE

species is usually an indication of succession, as the seeds only germinate

in the light ( Lindeman, verbal information) . M ost species are restricted
to the interior of Surinam and the savanna belt, which comprises the
cover landscape and the old coastal plain and which is characterized by
the occurrence of several types of savannas ( see Cohen & van der Eij k,
1 953; van Donselaar, 1 965). Only a few species also occur in the young
coastal plain (the northernmost belt with young sediments), e.g.
Cecropia surinamensis, C. obtusa, Ficus maxima, F. pertusa en F.
trigona; Ficus ernestiana, however, seems to be confined to this region.

LITERATU R E CITED
BAILLON, H., 1 875. Stirpes exoticae novae. Adansonia 1 1 : 292-3 1 2.
BERG, C.C., 1 972. Flora Neotropica Monograph 7. Olmedieae and Brosimeae
( M oraceae) . New York.
B U RGER, W.C., 1 962. S tudies in New World Moraceae: Trophis, Clarisia,
Acanthinophylum. Ann. Missouri Bot. Gard . 49: 1 -34.
-, LANJOUW, J . , & J . G . WESSELS BOER, 1 962. The genus Sorocea St. Hi!.
( M orae.). Acta Bot. Neerl. 1 1 : 428-477.
CHEW WEE-LEK, 1 963. A revision of the genus Poikilospermum ( Urticaceae).
Gard. Bull. Singapore 20: 1 - 1 03.
COHEN, A. & J.J. van EIJK, 1 953. Klassificatie en ontstaan van savannen in
Suri name. Geologie en Mij nbouw l l . 1 5: 202-2 1 4.
CORNER, E.J . H . , 1 962. The classification of Moraceae. Gard. Bull. S ingapore
1 9 : 1 87-252.
DON S ELAAR, J. VAN, 1 965. An ecological and phytogeographic study of northern
Surinam savannas. The vegetation of Surinam 4.
DUCKE, A. , 1 922. Plantes nouvelles et peu connues de la region Amazonnienne.
Arch. Jard . Bot. R io de Janeiro 3 : 1 -28 1.
-, 1 939. Plantes nouvelles et peu connues de la region A mazonienne. Arq. Serv. Flor.
Rio de Janeiro 1: 1 -40.
MACBRIDE, J . F . , 19 37. Flora of Peru. M oraceae. Pub!. Field Mus. Bot. 1 3(2.2):
274-3 3 1 .
M IQU EL, F.A.W., 1 85 3. U rticineae i n Martius, Flora Brasiliensis 4( 1 ) : 76-2 1 8.
PITTIER, H . , 1 9 1 2. New and N oteworthy plants from Colombia and Central
America Ill. Moraceae. The American genera of Olmedieae. Contr. U . S .
Nat. Herb 1 3: 43 1 -443.
RENNER, 0., 1 907. Beitrage zur Anatomie und Systematik der Artocarpeen und
Conocephaleen, insondere der Gattung Ficus. Bot. Jahrb. 39: 3 1 9-448.
TRECUL, A., 1 847. Sur la famille des Artocarpees. Ann. Sci . Nat. Bot. I l l . 8: 38-
1 57.
WOODSON , R . E . , 1 960, Flora of Panama. Moraceac. Ann. M issouri Bot. Gard.
47(2): 1 1 4- 1 78.
 MORACEAE 1 77

KEY TO T H E GENERAl 2

I a. Leaves opposite . .. . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . I. Bagassa

b. Leaves alternate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
.

2 a. Leaves spirally arranged , stipules connate . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

b. Leaves distichous, stipules mostly free . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 6
3 a. Leaves peltate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 4. Cecropia
b. Leaves not peltate ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 a. Flowers in urceolate receptacles (figs); latex usually copious . . . . 1 1 . Ficus
b. Flowers in branched to capitate inflorescences; latex scarce to almost Jacking 5
5 a. Without conspicuous granular hairs on the j uvenile parts; staminate flowers
with I or 2 connate stamens; pistillate flowers in heads; often epiphytes
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 3. Coussapoa
b. Usually conspicuous granular hairs on the juvenile parts; staminate flowers
usually with 4 stamens; pistillate flowers not in heads . . . . . . 1 2. Pourouma
6 a. Inflorescences globose, turbinate, or discoid, mostly bisexual, with peltate
bracts; uncinate hairs common .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
b. Inflorescences otherwise; uncinate hairs absent (except in Clarisia ilicifolia) 8
7 a. Inflorescences (almost) entirely covered with peltate bracts . . . 4. Brosimum
b. I nflorescences with a few peltate bracts, the middle part of the inflorescences
almost ebracteate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. Trymatococcus
8 a. Scars of the stipules encircling less than half the twig; staminate inflorescences
spicate; pistillate inflorescences spicate, capitate, or uniflorous, without a dis-
tinc involucre . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
b. Scars of the stipules encircling more than half the twig; inflorescences discoid,
with a distinct involucre . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 0
9 a. Inflorescences usually o n special shoots; staminate inflorescences with a distinct
sterile groove, flowers close together; pistillate inflorescences uniflorous or
capitate . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. Clarisia
b. Inflorescences on normal twigs; staminate inflorescences without a distinct
sterile groove, rachis compressed, flowers not close together; pistillate inflo-
rescences spicate (to subcapitate) . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. Sorocea
10 a. Stipules fully amplexicaul . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 1
b. Stipules not fully amplexicaul . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 3
11 a. Leaves glabrous, without parallel tertiary veins . . . . . . . . . . . 1 0 . Naucleopsis
b. Leaves more or less hairy, parallel tertiary veins present . . . . . . . . . . . . . . 1 2
12 a. I nflorescences sessile; pistillate inflorescences uniflorous . . 6 . Pseudolmedia
b. Staminate inflorescences pedunculate, pistillate inflorescences pedunculate or
sessile, with several to many flowers . . . . . . . . . . . . . . . . . . . . . . . . 7. Perebea
13 a. Leaves almost glabrous or minutely puberulous, shining above, greenish when
dried . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8. Maquira
b. Leaves distinctly hairy beneath, dull above, usually brownish when dried
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9. Helicostylis

1 M ainly based on Surinam species.

1 For introduced M oraceae see p. 295.
178 M ORACEAE

1 . Bagassa Aublet, PI. Gui. Suppl. 15. 1775.

T y p e s p e c i e s : Bagassa guianensis Aublet.

Laurea Gaudichaud , Voy. Bonite t. 88. 1 844.

Laurea Kunth, Linnaea 1 3: 562. 1 839.
Ty p e s p e c i e s : La urea tiliifolia Gaudichaud ( Bagassa tiliifolia ( Hamilton)
�

Benoist).

Dioecious trees, with abundant white latex. Leaves usually opposite,

decussate, entire to 3-lobed to 3-parted, palmatinervate: stipules free,
lateral. Staminate inflorescences paired or solitary in the axils of the
leaves, spicate, pedunculate, bracteate; flowers with 4-parted perianths,
2 ( -4?) stamens, and a pistillode. Pistillate inflorescences paired or solitary
in the axils of the leaves, capitate, pedunculate, bracteate; flowers
with 4-lobed perianths; ovary free. Fruit an achene; seed with some en­
dosperm; embryo with equal plane cotyledons and a long curved ra­
dicle.

D i s t r i b u t i o n : Possibly 2 species, in the Guianas and Para (Brazil) .

The genus possibly comprises 2 species. I am inclined to regard B.

guianensis and B. ti/iifolia as conspecific. None of the twigs of specimens
which agree with Aublet's description of B. guianensis bears inflores­
cences. In only one collection have separate infructescences been found;
they are identical with those of B. tiliffolia. The main differentiating
character is found in the pubescence of the twigs and leaves, which is
scabrous in B. guianensis and consists of short, weak, straight or curved
hairs in B. ti/iifolia. The leaf tips are long-acuminate and the leaf margin
distinctly crenate to serrate in specimens with asperous leaves, but the
tips are acute to short-acuminate and the margin tends to be entire in the
other specimens. Leaves of some seedlings collected in Guyana and attri­
buted to B. tiliifolia (the only species known from that country) have much
the same characters as B. guianensis leaves. Sagot 522 from French
Guiarta is intermediate between the two species. The distribution of the
two species is about the same. These facts suggest that specimens with
 M ORACEAE 1 79

asperous leaves, etc., are j uvenile forms. H owever, trees from which such
leaves have been collected appear to reach the same height as those of B.
ti/iifolia. Evidence seems to indicate more and more that Bagassa has
only a single species: B. guianensis.

a. Twigs and leaves more or less scabrous, leaf tips long-acuminate I.B. guianensis
b. Twigs and leaves puberulous (white curved hairs), leaf tips short-acuminate to
acute . . . .. . . . .. .. .
. . . . . . ..
. . . . . . . .. . . .
. . . . ..
. . . . . 2. B. tiliifolia
. . . . .

I. Bagassa guianensis Aublet, PI. Gui. Suppl. 1 5, t. 376. 1 775.

T y p e : Aub1et s.n. , French Guiana (BM).

Trees up to 20 m tall. Leafy twigs 3-4 mm thick, scabrous. Leaves

deltoid to ovate (to elliptic), entire to 3-lobed or to 3-parted, 1 0-30 cm
long, 8-23 cm broad, chartaceous, tips long-acuminate, emarginate to
cordate at the base; margin crenate to serrate; surface more or less
scabrous; veins nearly plane above, prominent beneath; petioles 4- 1 4 cm
long; stipules 1 0- 1 5 mm long, puberulous.

D i s t r i b u t i o n : A few collections known from Surinam, French

Guiana, and Pani (Brazil) .

In Surinam collected in the Nassau M o untains, Brokopondo D istrict,

Jodensavanne area, and near M oengo, in forests.

Van Donselaar 1 097; Lanjouw & Lindeman 2433; LBB 1 3403; Lindeman 5958.

Only a single fertile collection is known; its separate fruiting pistillate

inflorescences are similar to those of B. ti/ iifo /ia.

2. Bagassa tiliifolia ( Hamilton) Benoist, A rch. Bot. Caen. 5

( M em. I ) : 3 1 . 1 93 3 . Piper tiliifolium Desvaux ex Hamilton,
Prodr. Plant. Ind. Occ. 4. 1 825. (Non Piper tiliifolium Schlechtendal
& Chamisso).
1 80 MORACEAE

T y p e: Martin s . n . , French Guiana (P, ex herb. Desvaux).

Laurea tiliifolia Gaudichaud, Voy. Bonite t. 88. 1844; d' Alleizette, Explication et
Description des Planches de !'Atlas de Voy. Bonite 1 57. 1866, deser.
T y p e : Gaudichaud, Voy. Bonite t. 88. 1 844; probably drawn from coil. Martin
s.n. fr om French Guiana.

Trees up to 45 m tall, (sometimes?) with buttresses. Leafy twigs 2-6 mm

thick, minutely puberulous. Leaves suborbicular to deltoid to ovate to
elliptic, entire to 3-lobed , 6-22 cm long, 4- 1 7 cm broad, chartaceous
to subcoriaceous, tips short-acuminate to acute, emarginate to cordate
at the base; margin entire to crenate; except for main veins almost gla­
brous above, densely minutely puberulous beneath; veins nearly plane
above, prominent beneath; petioles 3-9 cm long; stipules 5- 1 5 mm long,
minutely puberulous.
Staminate inflorescences 4- 1 0 cm long, 3-5 mm in diameter, with an
abaxial sterile strip; peduncles I 0- 1 5 mm long, minutely puberulous;
flowers close together; perianth ea. I mm high, 4-parted, minutely
puberulous; stamens 2, filaments 1 . 2- 1 . 5 mm long, purplish, anthers
0. 1 -0. 1 5 x 0. 1 -0. 1 5 mm, latrorse to extrorse, pistillode subulate, 0. 5-0 . 8
mm long; bracts oblong t o lanceolate t o spathulate, ea. I m m long.
Infructescences subglobose, 2. 5-3 . 5 cm in diameter; peduncle 1 . 3-
1 . 5 cm long, flowers close together; perianth tubular, (at fruit) ea. I cm
high, 4-lobed, minutely puberulous, fleshy, pulpy, yellow, soursweet;
fruits ea. 5 mm long, endocarp crustaceous, exocarp membranaceous;
seed ea. 3 . 5-mm long; bracts lanceolate to spathulate, about as long as
the perianths.

D i s t r i b u t i o n : In the Guianas and Para (Brazi!) ; in forests up to 650 m

altitude.

In Surinam known from forests of the interior and the savanna belt.

V e r n a c u l a r n a m e s : kauhoedoe ( Sur.) jawahedan (Arow.).

B W 1347; LBB 9466; Lindeman 3685, 37 1 2; Stahel Exp. Wilh. Geb. 1 5 2, 1 94 .

 MORACEAE 181

When food is lacking people i n the forest drink the milky latex of this
tree.

3 . Clarisia Ruiz & Pavon, Prodr. Fl. Peruv. et Chil. 1 28 . 1 794, nom.
conserv . ; Lanj ouw, Rec. Trav. Bot. Neerl. 3 3 : 254. 1 936; Burger,
Ann. Missouri Bot. Gard . 49: 1 . 1 962; Berg, Acta Bot. N eerl.
1 7: 309. 1 968.

T y p e s p e c i e s : Clarisia racemosa Ruiz & Pavan

Sahagunia Liebmann, Kgl. Danske Vidensk. Selsk. Skr. 5: 3 1 6. 1 8 5 1 .

T y p e s p e c i e s : Sahagunia mexicana Liebmann ( Clarisia biflora Ruiz &
=

Pavon).
Soaresia Fr . Allemao, Revista Braz. 1 : 2 1 0. 1 857 (non Soaresia Schultz Bip.).
T y p e s p e c i e s : Soaresia nitida Fr. Allenao ( Clarisia racemosa Ruiz &
=

Pavon).
Acanthinophyllum Fr. Allema o, Revista Braz. 1 : 368, 1 858.
T y p e s p e c i e s : Acanthinophyllum strepitans Fr. Allema o ( Clarisia
=

ilicifolia (Sprengel) Lanjouw & Rossberg).

Aliteria Benoist, Bull. Mus. H ist. Nat. Paris I I . 1 : 1 63. 1 929.
T y p e s p e c i e s : Aliteria sagoti Benoist ( Clarisia ilicifolia (Sprengel)
=

Lanj ouw & Rossberg).

D ioecious trees or shrubs. Leaves distichous, pinnatinervate;

stipules free, lateral, small, leaving small scars. U ncinate hairs present.
Inflorescences in pairs or solitary in the axils of the (fallen) leaves or al­
ternately on short shoots and subtended by "bracts". Staminate in­
florescences spicate, with a sterile strip, pedunculate, bracteate; flowers
sessile, close together in longitudinal rows; perianth (2-) 3-6-lobed to
(2-) 3-6-parted, or segments free; stamens 1 -5, straight in the bud.
Pistillate inflorescences capitate or uniflorous, pedunculate, bracteate;
flowers sessile; perianth tubular, subentire to 4-lobed; ovary adnate
to the perianth or nearly free, stigmas 2. Fruiting perianth fleshy, fruit
adnate to the perianth; seed without endosperm, testa rather thin,
vascularized, cotyledons thick, equal, free, radicle apical, straight, very
short.

D i s t r i b u t i o n : 3 species in tropical S outh and Central America.

1 82 MORACEAE

In 1 936 the genus was treated by Lanjouw, who distinguished 8 species.

Two of them were included in the re-established genus A canthinophy/lum
by Burger ( 1 962), who recognized only 2 species in Clarisia. The separation
of Acanthinophy/lum and Clarisia has proved to be untenable ( Berg, Acta
Bot. Neerl. 1 7 : 309. 1 968) .

a. Costa of the leaves prominent above; uncinate hairs present; pistil late inflo-
rescences capitate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. C. ilicifolia
b. Costa of the leaves plane to sli g htly impressed above; uncinate hairs lack ing;
pistillate inflorescences uniflorous . . . . . . . . . . . . . . . . . . . . . . 1. C. racemosa

I. Ciarisia racemosa Ruiz & Pavan, Syst. Veg. Fl. Peruv. et Chi!. 25 5 .
1 798.

T y p e : Ruiz & Pavon s . n . , Peru (MA, n o t seen).

Soaresia nitida Fr. Allemiio, Revista Braz. 1 : 2 1 0, cum tab. 1857.

C1arisia nitida (Fr. Allema o) Macbride, Pub!. Field M us. Bot. 1 1 : 15. 1 93 1 .
T y p e : Fr. Allemao, Revista Braz. 1 : tab. 1857.

Trees up to 40 m tall; latex white. Leafy twigs 0. 5-2 mm thick, minute­

ly puberulous to almost glabrous. Leaves elliptic to oblong to lanceolate,
5- 1 4 cm long, 2-6. 5 cm broad, often broadest below the middle and
somewhat inequilateral, subcoriaceous to coriaceous, acuminate, obtuse
(to emarginate) at the base; margin entire; glabrous and shining above,
sparsely puberulous, glabrescent beneath; costa plane to slightly im­
pressed above, for the rest the veins more or less prominent, 9- 1 8
pairs of secondary veins, without parallel tertiary veins; petioles 6- 1 5
mm long, canaliculate; stipules 1 -2 . 5 mm long, deltoid, puberulous.
Inflorescences usually in pairs subtended by reniform to ovate bracts
on leafless shoots which may be short and arise from older parts of the
twigs. Staminate ones 1 - 1 0 cm long; peduncle 1 -8 mm long, minutely
puberulous; perianth 0. 5-0. 8 mm high, consisting of 3-6 free or
sometimes basally connate, variously shaped , puberulous segments;
stamens 1 -2(-3), filaments 0.3- 1 .0 mm long, anthers ea. 0 . 3 x 0.4 mm;
bracts variously shaped , partly suborbicular and (sub)peltate, puberu-
 M ORACEAE 1 83

Fig. I Clarisia racemosa; leaf, staminate inflorescences (C. Blanco 509), pistillate
inflorescences (Ducke 1305 1 ) and infructescences (Prance et al. I 0488).
1 84 MORACEAE

lous. Pistillate inflorescences uniflorous; ped uncle 4 mm long or to 1 5

mm long at fruit, puberulous; perianth 2-3 mm high, subentire to
4-lobed, puberulous at the tip; ovary adnate to the perianth, style u p to
2 mm long, stigmas 5-7 mm long; bracts 3-7, suborbicular to deltoid,
often subpeltate, puberulous. Fruiting perianth ea. 1 . 5 cm high, ellipsoid.

D i s t r i b u t i o n : Apparently common in the Amazon Basin and the

coastal region of Brazil, but probably rare in north-eastern Co lombia,
northern Venezuela, and Surinam.

In Surinam a single sterile specimen collected m the Kaboeri Forest

Reserve near the Corantyne River.

BW 4968.

2. Clarisia ilicifolia (Sprengel) Lanj ouw & Rossberg, Rec. Trav. Bot.
Neerl. 33: 7 1 7. 1 936.
Excoecaria ilicifolia Sprengel, Neue Entdeckungen 2: 1 1 7. 1 82 1 .
Acanth\nophyllum ilicifo1ia (Sprenge1) W . Burger, Ann. Missouri
Bot. Gard . 49 : 27. 1 962.

T y p e : Without collector's name, Brazil (B, not seen).

Acanthinophyllum strepitans Fr. Allemao, Revista Braz. I: 369, cum tab.

1858.
Clarisia strepitans (Fr. Allemao) Lanjouw, Rec. Trav. Bot. Neerl. 33: 274.
1 936.
T y p e : Fr. Allemao, Revista Braz. 1: tab. 1858.
Clarisia strepitans var. guianensis Lanjouw, loc. cit. 274.
Clarisia ilicifolia var. guianensis Lanjouw & Rossberg, loc. cit. 7 18.
T y p e : Stahel Exp. Wilh. Geb. 1 20 9 , Surinam ( U ).
Clarisia strepitans var. paraensis Lanj ouw, loc. cit. 275.
Clarisia ilicifolia var. paraensis Lanjouw & Rossberg, loc. cit. 7 18.
T y p e Huber s.n. (HAMP 6965) 9, Brazil, Para (U).
Clarisia strepitans var. micranthera Lanjouw, loc. cit. 275.
Clarisia ilicifolia var. micranthera Lanjouw & Rossberg, loc. cit. 7 18.
T y p e : Glaziou 1 2 1 72 d, Brazil, Rio de Janeiro (K).
 MORACEAE 1 85

Clarisia strepitans var. cuyunensis Lanj ouw, loc. cit. 275.

Clarisia ilicifolia var. cuyunensis Lanjouw & Rossberg, loc. cit. 7 1 8.
T y p e : Davis s.n. (Forest Department 1 029) o , Guyana (U).
Sahagunia racemifera Huber, Bol. Mus. E milio Goeldi. 5 : 3 34. 1 908.
T y p e : Ducke s.n. (HAMP 2885) o , Brazil, Pani (MG, not seen).
Aliteria sagoti Benoist, Bull, Mus. H ist. Nat. Paris 11. 1 : 1 63. 1 929.
L e c t o t y p e : Sagot 530 9, French Guiana (P).
Sorocea stenophylla Standley, Tropical Woods 43: 1 8. 1 935.
T y p e : Capucho 447 9, Brazil, Para (F, not seen).
Clarisia spruceana Lanj ouw, Rec. Trav. Bot. Neerl. 33: 272. 1 936.
Acanthinophyllum spruceana ( Lanj ouw) W. Burger, Ann. M issouri Bot.
Gard . 49: 30. 1 962.
T y p e : Spruce 3279 o , Venezuela, Amazonas ( K).

Shrubs or trees up to 20 m tall, with white to cream ( resinous) latex.

Leafy twigs 1 -4 mm thick, usually with uncinate hairs; periderm often peel­
ing off. Leaves (elliptic to) oblong to lanceolate, ( 3-)8-33 cm long, ( 1 . 5-)
2 . 5- 1 2 cm broad, sometimes broadest below the middle, chartaceous,
sharply acuminate to caudate, acute to obtuse at the base; margin entire,
spinose to dentate, lobed or rarely deeply sinuate; glabrous and shining
above, glabrous or with a few uncinate hairs beneath; veins prominent,
( 5-)8-20 pairs of secondary veins, without parallel tertiary veins; petioles
3- 1 5 mm long, canaliculate, often with uncinate hairs; stipules 1 -3 mm
long, subulate, puberulous.
I nflorescences usually in pairs, subtended by ovate bracts, on (branch­
ed) leafless short shoots. Staminate ones 2-20 mm long; peduncle (0-) 1 -
5 mm long, with (rather) dense retrorse uncinate hairs; perianth 0. 5-0.6
mm high, (2-)3-4-lobed to ( 2-)3-4-parted, minutely puberulous; stamens
1 (-3), filaments 0 . 7- 1 .6 mm long, anthers 0. 2-0 . 3 x 0 . 2-0. 3 mm; bracts vari­
ously shaped, often subpeltate, minutely puberulous, sometimes also
with uncinate hairs. Pistillate inflorescences ea. 5 mm in diameter; pedun­
cle 3-20 mm long, with sparse to dense retrorse uncinate hairs; flowers 6-
20; perianth 1 .5-2.0 mm high, subentire, almost glabrous to rather den­
sely puberulous, sometimes with uncinate hairs intermixed; ovary free or
nearly so, stigmas 1 -2 mm long. Fruiting perianth 1 . 5-2.0 cm high, ellip­
soid to oblongoid , orange (to reddish or yellowish); fruit adnate to the pe­
rianth; some basally attached bracts below or sometimes among the flo­
wers.
186 MORACEAE

D is t r i b u t i o n : In the Amazo n Basin, the coastal region of Brazil, and

the Guianas.
In Surinam known mainly from the interior; also collected in the sa­
vanna belt; in the understory of rain forest; also in secondary forest and
seasonal swamp fo rest.

Van Donselaar 1 22 1 , 1 300, 208 2 , 33 1 8 ; Florschi.itz 1 633; Jonker & Dan iels 1 079, 1 08 1 ;
Lindeman 4778, 5053 , 5 1 78 , 6 1 6 1 , 6458, 6776; Stahel Exp. Wilh. Geb. 1 20; Wessels
Boer 1 080.

V e r n a c u l a r n a m e s : manletterh out, letterhout (Sur. D utch).

2. Sorocea St. H ilaire, M em. M us. H ist. Nat. Paris 7: 473 . 1821; Burger,
Lanj ouw & Wessels Boer, Acta Bot Neerl. 11: 428. 1962.

T y p e s p e c i e s : Sorocea bonplandii ( Baillon) W. Burger, Lan­

iouw & Wessels Boer

Balanostreblus Kurz, Jour. Roy. Asiat. Soc. Bengal. 42: 247. 1 873.
T y p e s p e c i e s : Balanostreblus ilicifolia Kurz ( Sorocea guilleminiana
=

Gaudichaud).
Pseudosorocea Baillon, Adan sonia 1 1 : 296. 1 875.
L e c t o t y p e s p e c i e s : Pseudosorocea bonplandii Baillon ( Sorocea bon­
=

plandii ( Baillon) W. Burger, Lanjouw & Wessels Boer).

Paraclarisia Ducke, Arq. Serv. Flor. Rio de Janeiro 1: I . 1 939.
T y p e s p e c i e s: Paraclarisia amazonica Ducke ( Sorocea duckei W.
=

Burger).

Dioecious trees or shrubs. Leaves distichous, pinnatinervate; stipules

free, lateral, leaving small scars. Inflorescences geminate or solitary in
the axils of the leaves, spicate (or subcapitate ), with suborbicular and usu­
ally peltate bracts and a more or less compressed rachis. Staminate flo­
wers sessile (in Surinam species); perianth 4-parted , segments imbricate;
stamens 4, straight in the bud . Pistillate flowers sessile or pedicellate, peri­
anth tubular, subentire to 4-lobed ; ovary free or basally adnate to the peri­
anth , stigmas 2, short. Fruiting perianth coloured and more or less
fleshy; seed without endosperm, testa thin, except a thickened suborbicu-
 MORACEAE 1 87

lar vascularized part, covering a concave unfused part of the equal cotyle­
dons, radicle short, deflexed .

Distribution: 22 to 25 species ranging from Guatemala to northern

Argentina.

Burger, Lanj ouw and Wessels Boer ( 1 962) revised the genus and divided
it into the subgenera Sorocea and Paraclarisia. The two closely related
species known from Surinam belong to the former subgenus. All Pseudo­
sorocea species described by Baillon (Adansonia 1 1 : 295 . 1 875) proved to
belong to Sorocea. P. poeppigii was mentioned among the synonyms of
Acanthinophyllum ilicifolia by Burger (Ann. M issouri Bot. Gard. 49: 27.
1 962). Baillon's description refers at least for the greater part to a speci­
men of S. muriculata. This specimen and a specimen of Clarisia ilicifolia
( Acan thinophyllum ilicifolia) were mounted on the same sheet. In 1 9 1 4
=

Rojas Acosta described Trophisomia edulis (in Bull . Geogr. Bot. 24:
2 1 1 ). The description is based on N iederlein 3359 from A rgentina
(Chaco). The vernacular name is yatita. This species is so poorly descri­
bed that it is not possible to be certain about its identity; it might be con­
specific with Sorocea saxicola Hassler, for which the same vernacular
name is known.

I a. Petioles 3-9 mm long, mostly with sparse, not distinctly patent, minute hairs
or almost glabrous; pistillate inflorescences 1 -4 mm long, usually subcapitate,
with 3-9 flowers; fruiting perianth ellipsoid to cylind rical 2. S. guayanensis
b. Petioles 1 .5-4(-7) mm long, usually with rather sparse, often distinctly patent,
minute hairs; pistillate inflorescences 5-25 mm long, distinctly spicate, with
6- 1 6 flowers; fruiting perianth (sub )globose, usually basally more or less
muriculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. S. muriculata

I. Sorocea muriculata Miquel, Fl. Bras. 4( 1 ) : 1 1 3, t. 34. 1 85 3 .

T y p e : Martius s.n. 9, B razil , Amazonas ( M ).

Sorocea amazonica Miquel, Fl. Bras. 4( 1 ) : Ill, t. 34. 1853.

T y p e : Martius s.n. <(, Brazil, Amazonas ( M ) .
Pseudosorocea uaupensis Baillon , Adansonia 1 1 : 297. 1875.
1 88 MORACE A E

E
u

Fig. 2 Sorocea, pistillate inflorescences; a . , S. muricu/ata (Prance et al. 76 1 6,

Schultes & Black 8547); b . , S. guayanensis (Rodrigues & D. Coelho 4744, Wessels
Boer 1 069) .
 MORACEAE 1 89

Sorocea uaupensis ( Baillon) Macbride, Candollea 5 : 349. 1 934.

T y p e : Spruce 27 1 5 o, Brazil, Amazonas ( P).
Pseudosorocea poeppigii Baillon, Adansonia 1 1 : 297. 1 875.
T y p e : Poeppig s.n. 9, Brazil, Amazonas (P).
Sorocea dentata Huber, Bol. M us. Emilio Goeldi 5: 3 3 3 . 1 909.
T y p e : Ducke s.n. (HA M P 8957) <;.>, Brazil, Pani ( MG , not seen; isotype
BM).

Shrubs or trees up to 12 m tall. Leafy twigs 0. 5-3(-4) mm thick, minute­

ly and usually rather densely puberulous, usually because of patent
hairs. Leaves (elliptic to) oblong to (narrowly) lanceolate, mostly broad­
est at or above the middle, usually more or less inequilateral, ( 2-) 5 . 5-
1 6.5( -22) cm long, 2-4.5( -7) cm broad , chartaceous to subcoriaceous, acu­
minate to caudate, acute to obtuse at the base; margin entire or more or
less distinctly dentate; glabrous or sparsely puberulous on the costa be­
neath; above the costa sometimes slightly impressed , the other veins
slight ly prominent to plane, the secondary veins sometimes slightly
impressed, the veins beneath prominent, 8- 1 2 pairs of secondary veins,
without parallel tertiary veins; petioles 1 . 5-4( -7) mm long, minutely and
usually rather densely puberulous, often because of patent hairs; stipules
2-4 mm lo ng, sparsely puberulous.
Staminate inflorescences l-4.5 cm long, including the l -3 mm long pu­
berulous penduncle, flowers rather distant at anthesis; tepals 4( -6), ea . 1
mm high, mostly unequal, almost glabrous to puberulous; stamens 4( -6),
filaments 0 . 3- 1 .2 mm long, often more or less incurved at anthesis,
anthers 0.4-0 . 5 x 0.4-0 . 5 .
Pistillate inflorescences 0.5- 1 . 5( -2.5) c m long; including t h e l-3 m m
long puberulous peduncle; flowers 6- 1 6, sessile or pedicellate; perianth
ea. 2 mm high, slightly 4-lobed, rather sparsely puberulous ; stigmas ea.
0 . 5 mm long, tongue-shaped .
I nfructescences up to 4 cm long, rachis red ; fruiting perianth (sub)glo­
bose, ea. 6 mm in diameter, red , the lower part usually more or less dis­
tinctly muriculate, the upper part (almost) glabrous , becoming earlier
red-coloured than the lower part, pedicels red, up to 6 m m long, normally
1 . 5-2 mm in diameter, sometimes swollen and as broad as or even
broader than the fruiting perianth and more or less muriculate.
1 90 MORACEAE

D i s t r i b u t i o n : The Amazon Basin ( Brazil, Peru, Ecuador, and

Venezuela) and Surinam; in no n-i nu ndated a nd river-flooded forests.

In Surinam known from the Left-Coppename River, the Upper Suri­

name River, and the Sipaliwini savanna area.

Florschiitz & Maas 2805; Oldenburger, Norde & Schulz I 1 27, 1 402; Stahel
Exp. Wilh. Geb. 1 4 3 .

Sorocea muricu/ata and S. amazonica, kept separated by Burger, Lan­

j ouw & Wessels Boer (1962), are without doubt conspecific. The strongly
swollen pedicels, mentioned as being unique to S. amazonica, have to
be regarded as an aberration . Intermediates between the rather slender
normal pedicels and the very broad ones are met with in some specimens.

2. Sorocea guayanensis W . Burger, Acta Bot. Neerl. 11: 439. 1962.

T y p e : Ll. Williams 14532 <;.>, Venezuela, Amazonas ( F. not seen;

isotype YEN).

Shrubs or trees up to 8 m tall. Leafy twigs 0. 5-3( -4) mm thick, sparsely mi­
nutely puberulous to almost glabrous. Leaves oblong to (narrowly) lance­
olate, mostly broadest above the middle and slightly inequilateral, (2. 5-)
4.5- 1 8 . 5 cm long, ( 1 -) 1 .5-7 cm broad , chartaceous to subcoriaceous, acu­
minate to caudate, acute to subobtuse at the base; margin entire or more
or less slightly dentate toward s the apex; glabrovs or with sparse hairs on
the costa beneath; above the costa slightly impressed, the other veins
slightly prominent to plane, the veins beneath more or less prominent, ( 7-)
9- 1 3 pairs of secondary veins, without distinct parallel tertiary veins; pe­
tioles 3-9 mm long, very sparsely and minutely puberulous to almost gla­
brous; stipules 2-4 mm long, usually with sparse minute hairs.
Staminate inflorescences 2-2.5 cm long, including the 1 -2 mm long pu­
berulous peduncle; flowers rather distant at anthesis, sessile; tepals 4,
 MORACEAE 191

mostly unequal, ea. I m m long, almost glabrous; stamens 4, filaments 0 . 3-

1 .0 mm long, more or less incurved at anthesis, anthers 0.4-0 . 5 x 0.4-0 . 5
mm.
Pistillate inflorescences spicate to subcapitate, 1 -4( - 1 2) mm long, inclu­
ding the up to I mm long puberulous peduncle; flowers crowded, 3-9, sub­
sessile or pedicellate; perianth 1 - 1 . 5 mm high, slightly 4-lobed, puberu­
lous; stigmas 0.5- 1 . 5 mm long, usually recurved .
lnfructescences up to 1 . 5? cm long, (rachis red?), fruiting perianth
ellipsoid to cylind rical, ea. 8 mm high, sometimes curved, red, puberu­
lous, rather densely so near the apex, pedicels red, up to 4 mm long,
1 - 1 . 5 mm in diameter.

D i s t r i b u t i o n : S outhern Venezuela, the adj acent region m Brazi�

and in Surinam; in forests, mostly along streams?

In Surinam once collected along the Oelemari River.

Wessels Boer 1 069.

Neither the vegetative parts nor the staminate inflorescences and flowers
afford satisfactory characters for separating Sorocea guyanensis from S.
muricu/ata. H owever, the pistillate i nflorescences, especially when bear­
ing fruiting perianths, do give some good differentiating characters.
Both the resemblances and the nature of the differences j ustify the ques­
tion as to whether the two taxa should be regarded as distinct at the speci­
fic level. Burger, Lanj ouw & Wessels Boer ( l 962) noted that S. guayanen­
sis has blackish fruiting perianths; this was based on the field notes given
for some collections from Guyana. However, these collections differ
from the type collection in several characters and belong to another (still
undescribed?) species.
* The fruiting perianth turns black when fully mature at least in

Sorocea muricu /ata, but probably also in S. guayanensis.

192 MORACEAE

4. Brosimum Swartz, Prod r. Veg.lnd . Occ. 12.1788, nom. conserv . ; Pittier,

Contr. U. S. Nat. H erb. 20: 96. 1918; Berg, Flora Neotropica
M onogr. 7: 161. 1972.
Alicastrum P. Browne, H ist . Jam. 372. 1756, nom. rej ic.

T ype s p e c i e s : Brosimum alicast rum Swartz.

Piratinera Aublet, PI. Gui 2: 888. 1 775, nom. rejic.

T y p e s p e ci e s : Piratinera guianensis Aublet ( Brosimum guianensis
=

(Aublet) Huber).
Ferolia Aublet, PI. Gui . Suppl. 7. 1 77 5 . nom. rejic:1)
T y p e s p e ci e s : Ferolia guianensis Aublet ( Brosimum rubescens Taubert).
=

Galactodendrum Humboldt, Bonpland & Kunth, Relat. Hist. 2: 1 08. 1 8 1 9 .

T y p e s p e ci e s : Galactodendrum utile Humboldt, Bonpland & Kunth
( Brosi mum utile (H. B . K . ) Pittier).
=

Brosimopsi s S. Moore, Trans. Linn. Soc. 11. 4: 473 . 1 895.

T y p e s p e ci e s : Brosimopsis lactescens S . M oore ( Brosimum lactescens
=

(S. Moore) C.C. Berg).

M onoecious or dioecious trees. Leaves di stichous, pinnatinervate ; stipu­

les connate or free, fully amplexicaul or not so. Uncinate hairs usually
present. Inflorescences geminate or solitary in the axils of the leaves, bi­
sexual or unisexual, globose, turbinate or convexly discoid, receptacle
covered with peltate bracts. Staminate flowers several to many, perianth
distinct and with 1-5 tepals, vestigial or lacking, stamens 4- 1 , straight in
the bud, anthers latrorse to extrorse. Pistillate flowers one to several,
embedded in the receptacle, ovary ad nate to the embedded perianth,
stigmas vittiform to filiform. Fruit ad nate to the enlarged , often coloured,
receptacle; seed big and with out endosperm, testa with a thickened
(vascularized) part, embryo transverse or oblique in the fruit, cotyledons
thick and equal or unequal, radicle short.

D i s t r i b u t i o n: 13 species from M exico and the Antilles to

southern Brazil.

The genus is fairly well differentiated and shows a number of interesting

variations and morphological series. Some differences have given rise to

1 See Taxon 1 7 : 73 1 . 1 968, and 1 9: 273 . 1 970.

 MORACEAE 1 93

recognition of the genus Piratinera in addition to Brosimum s. str . , but

Piratinera has also been rega rded as congeneric with Brosimum. This and
the fact that Piratinera was established earlier than Brosim um has led to
the transfer of many epithets. The number of flowers in the inflorescence,
used by Pittier ( 1 9 1 8) as a criterion for separating Brosimum and Pirati­
nera, varies in most species from one to several and has little taxonomic
value. Brosimopsis was first inserted in the Euartocarpeae and then pla­
ced by D ucke (Arch. Jard . Bot. Rio de Janeiro 3: 30. 1 922) in the Olme­
dieae. Although he observed resemblances with Brosim um, he cons idered
the genus to be distinct and referable to the Olmedieae mainly on account
of the dioecism of the unisexual inflorescences. In 1 939 D ucke (Arq.
Serv. Flor. Rio de Janeiro I : 8) added to this character differences in the
habit, the wood, and the taste of the latex. However, considering the na­
ture of the differences with the Olmed ieae and the variation within Brosi­
mum, it is obvious that Brosimopsis has to be included into Brosimum.
Ferolia guianensis, referred to Parinari(um) by de Jussieu (Gen. PI. 342.
1 789) has proved to be identical with Brosimum rubescens ( Berg, Taxon
1 7 : 73 1 . 1 968).
In most species uncinate hairs occur on several parts of the plants.
This may help to distinguish sterile Brosimum specimens from members
of the Olmedieae. These hairs are also found in the related genus
Trymatococcus.
a. Stipules fully amplexicaul and connate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
b. Stipules not fully amplexicaul and free . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 a. Leaves with 20- 32 pairs of secondary veins, more or less densely hairy
beneath . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. B. parinarioides
b. Leaves with 1 2- 1 8 pairs of secondary veins, almost
glabrous beneath . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. B. rubescens
3 a. Costa of the leaves impressed above . . . . . . . . . . . . . . . . . . . . 2. B. lactescens
b. Costa of the leaves prominent to nearly plane above . . . . . . . . . . . . . . . . . . 4
4 a. Leaves almost glabrous above, papillose beneath . . . . . . . . . . 3 . B. guianense
b. Leaves scabrous to scabridulous above, without papillae
beneath . .......................................... I. B. acutifolium

1 . Brosimum acutifolium H uber ssp, acutifolium.

Brosimuin acutifolium Huber, Bol. M us. Emilio Goeldi 6: 66.
1 9 10.
194 MORACEAE

Piratinera acutifolia ( H uber) Pittier, Contr. U . S . Nat. H erb. 20:

100. 19 18.
Brosimopsis acutifo lia ( H uber) Ducke, Arch. Jard . Bot. Rio de
·

Janeiro 3: 30. 1922.

T y p e : Goeld i s.n. ( H A M P 8231), Brazil, Para ( M G, not seen;

isotypes B M , U).

Dioecious trees up to 35 m tall, with white latex. Leafy twigs 2-5 mm

thick, white puberulous. Leaves (elliptic to) oblong to lanceolate, 5- 1 8
cm long, 2 . 5-7 cm broad , chartaceous, acuminate to mucronate, acute to
obtuse at the base; margin entire; above scabrous to scabridulous, bene­
ath with rather dense minute to rather long, straight or curved hairs, usu­
ally slightly scabridulous, costa often with unci nate hairs ; veins nearly
plane above, prominent beneath, 8- 1 5 pairs of secondary veins, without
parallel tertiary veins; petioles 3-6 mm long, often with uncinate hairs; sti­
pules free, not fully amplexicaul, 2-8 mm long, white appressed-pubes­
cent.
Staminate inflorescences globose, 4-8 mm in diameter; peduncle 5-8
mm long, white puberulous; receptacle white puberulous; staminate flo­
wers many; perianth lacking; stamens 1 -3, filaments 0. 2-0. 5 mm long, ant­
hers 0. 1 5-0. 2 x 0. 1 5-0.2 mm, connective rather broad; many white puberu­
lous bracts.
Pistillate inflorescences (sub)globose, 8- 1 2 mm in diameter; peduncle
ea. 1 cm long, white puberulous; receptacle white puberulous; flowers 1 -
5 ; many white puberulous bracts. Fruiting receptacle subglobose t o sub­
discoid, up to ea. 2.5 cm in diameter, yellow to orange at maturity; seeds
ea. 1 cm long.

D i s t r i b u t i o n : From Para ( B razil) to Surinam.

I n Surinam in forests of the interior and the savanna belt; apparently

restricted to the eastern part of the country.
 M ORACEAE 195

Fig. 3 Brosimum /actescences; a., staminate inflorescences (N .T. Silva 1 1 72); b . ,

pistillate inflorescences ( Maas et a l . P. 13306).
1 96 MORACEA E

Geij skes 5; Lanjouw & Lindeman 2945; Lindeman 446 1. s . n . (inflorescences only);
Schmidt s.n.

V e r n a c u l a r n a m e s : takini (Car.) , iari (Arow. ).

The leaves are often markedly convex and therefore dry irregularly
folded in the press. B. acutifolium ssp. interjectum occurs in western
Para and eastern Amazonas; B. acutifolium ssp. obovatum occurs in
Brazil (Amazon Basin, west from M anaus), Peru, and Guyana.

2. Brosimum lactescens (Sp. M oore) C. C. Berg, Acta Bot. Neer1. 1 9 : 326.

1 970.
Brosimopsis lactescens Sp. Moore, Trans. Linn. S oc. 11. 4: 473 ,
t. 30, 3 1 . 1 895.

L e c t o t y p e : Sp. M oore 677, Brazil, Mato Grosso ( BM).

Brosimopsis oblongifolia Ducke, Arch. Jard . Bot. Rio de Janeiro 3: 3 1 .

1 922.
L e c t o t y p e : Ducke s.n. (HJ BR 1 2720), Brazil, Para ( R B, not seen;
isotypes B, K, U).
Brosimopsis d iandra S . F. Blake, Proc. Bol. Soc. Wash. 35: 1 80. 1 9 22.
T y p e : Curran 25, Brazil, Bahia ( U S).
Brosimopsis amplifolia Ducke, Arch. Jard . Bot. Rio de Janeiro 4: 3. 1 925.
T y p e : Ducke s.n. (HJB R 1 8 260), Brazil, Para ( R B , not seen; isotypes
B, K, P, U , US).
Brosimum ojoche Woodson, Ann. M issouri Bot. Gard. 47: 1 26. 1 960.
T y p e : Alien 5809, Costa Rica ( U S ) .
Brosimum belizense Lundell, Wrightia 3 : 1 66. 1 966.
T y p e : Gentle 1 737, British Hond uras ( L L , not seen; isotypes K, MO, NY).

Dioecious trees up to 35 m tall, with white latex. Leafy twigs 1 . 2 mm

thick, sparsely to densely puberulous. Leaves (elliptic to) oblong to
lanceolate, (2. 5-)4- 1 6(-23) cm long, ( l -) 1 . 5-5. 5(-6) cm broad, sometimes
broadest at the lower end, ( sub)coriaceous, acuminate, acute to obtuse
to subcordate at the base; above glabrous; beneath appressed
puberulous to almost glabrous, glabrescent; costa impressed , the other
 MORACEAE 1 97

veins more or less prominent above, the veins beneath prominent, 1 0-20
pairs of secondary veins, a few parallel tertiary veins; petioles .5- l 0 mm
long, often blackish; stipules free, not fully amplexicaul, 6- 1 0 mm long,
sparsely puberulous, often with prominent veins, often subpersistent,
often leaving prominent scars.
Staminate inflorescences globose, 3- 1 0 mm in diameter; peduncle
1 -6 mm long, puberulous, usually a bract at the base; flowers numerous,
close together; perianth (3-)4-lobed to ( 3-)4-fid to ( 3-)4-parted, 1 . 5-2.0
mm high; stamens 2-4, mostly unequal, filaments 2-4 mm long, anthers
0.6- 1 .2 x 0. 2-0 . 5 mm, connective narrow; numerous small bracts,
long-stipitate, ea. I mm high, puberulous.
Pistillate inflorescences (sub)globose to obovoid, 3-4 mm in diameter,
sometimes lobed; peduncle 2-5 mm long, puberulous, usually a bract at
the base; receptacle almost glabrous to densely puberulous and with
uncinate hairs; flowers 1 -4; style exceeding the receptacle by up to
2 mm, stigmas 5-7 mm long. Fruiting receptacles 1 . 2- 1 .6 cm in diameter,
at maturity yellowish; fruits ea. I cm long.

D i s t r i b u t i o n : From southern Brazil ( Santa Catarina) to M exico.

In Surinam only known from the Brokopondo District and the Voltz­
berg.

van Donse1aar 1449, 3 364; LBB 1 3147 .

V e r n a c u l a r n a m e : manletterhout (Sur. Dutch).

This widespread species varies in the number of stamens and in the pubes­
cence of the leaves and inflorescences. It may prove possible to distin­
guish infraspecific taxa, but it is not clear from the material seen hitherto
whether the variation is gradual or d iscontinuous. The staminate flowers
of specimens from the eastern coastal region of Brazil, Para, and north­
ern Central America have 2 stamens; those of specimens from the M ato
Grosso, Colombia, and Central America have 4 stamens; while those of a
specimen from Guyana have 3-4 stamens. The two sterile specimens from
Surinam most nearly resemble those from Para.
1 98 MORACEAE

3 . Brosimum guianense (Aublet) H uber, Bol. M us. Goeldi 5: 337. 1 909.

Piratinera guianensis Aublet, PI. Gui. 2: 888, t. 340. 1 77 5 .
Alicastrum guianense (Aublet) 0 . Kuntze, Rev. G e n . PI. 2 :
623 . 1 89 1 .
Brosimum aubletii Poeppig & Endlicher, N ov. Gen. et Spec. 2:
34, t. 1 48, 1 838, as synonym.

T y p e : Aublet s . n . , French Guiana ( B M ) .

Brosimum discolor Schott in Sprengel Syst. 4 (Cur. Post. App.): 403 . 1 827.
Piratinera discolor (Schott) Pittier, Contr. U . S . Nat. Herb. 20: 1 00. 1 9 1 8.
T y p e : Schott s.n . , Brazil (W. destroyed , replaced by F).
Piratinera panamensis Pittier, Contr. U. S . Nat. Herb. 20: 1 00, t.7. 1 9 1 8.
Brosimum panamense (Pittier) Standley & Steyermark, Pub!. Field Mus.
Bot. 23 : 40. 1 944.
T y p e : Pittier 4336, Panama ( U S ) .
Brosimum lecointei Ducke, Arch. Jard . B o t . R i o de Janeiro 3 : 28. 1 922.
L e c t o t y p e : Ducke s.n. ( HJ B R 1 2506, H A M P 1 5694), Brazil, Para
( RB, not seen; isotypes B, BM, F, NY, P, U, US).
Piratinera scabridula S . F . Blake, Jour. Wash. Acad . 1 2: 397. 1 922.
T y p e : Anderson s.n. (Forest Department 406), Guyana (US).
Piratinera velutina S . F. Blake, Jour. Wash. Acad . 1 2: 398. 1 922.
Brosimum velutinum ( S . F . B lake) Ducke, Arch. Jard . Bot. Rio de Janeiro
4: I. 1 925.
T y p e : B W 1 647, Surinam (US).
Brosimum tessmannii M ildbraed, Notizbl. Bot. Gart. Berlin 1 0 : 1 90. 1 927.
T y p e : Tessmann 3 268, Peru, Loreto (B).
Piratinera lemeei Benoist, Bull. Mus. H ist. Nat. 3 1 : 468. 1 93 1 .
Brosimum lemeei ( Beno ist) Lemee, Fl. Guyane Fran�. I : 5 1 0. 1 955.
T y p e : Lemee s.n . , French Guiana (P).
Piratinera mollis Killip, Jour. Wash. Acad . 26: 358. 1 936.
T y p e : Mutis 365, Colombia (US).
Brosimum palmarum Standley, Pub!. Field Mus. Bot. 1 7: 1 58. 1 937.
T y p e : Krukoff 8546, B razil, Amazonas (NY).
Brosimum rotundatum Standley, Bull. Torrey Club 75: 293. 1 948.
T y p e : Fanshawe 688 (Forest Department 3324), Guyana ( F).

Monoecious trees up to 45 m tall; with yellowish to white latex. Leafy

twigs l -3 mm thick, sparsely to rather densely white to brown puberu­
lous; periderm usually peeling off. Leaves elliptic to lanceolate, ( 2-)4- 1 3
cm long, ( l -)2-6 cm broad, often broadest below or above the middle,
 MORACEAE 1 99

more or less inequilateral, chartaceous to coriaceous, acuminate, acute,

obtuse or emarginate, at the base acute to obtuse to subcordate; margin
entire, often more or less revolute; above glabrous and shining; beneath
more or less densely papillose, mainly in the areoles, nearly glabrous to
appressed-puberulous to (sub)velutinous to scabridulous on the veins;
veins more or less prominent, 6- 1 4 pairs of secondary veins, without pa­
rallel tertiary veins; petioles 2-6 mm long, not canaliculate, often with un­
cinate hairs; stipules free, not fully amplexicaul, 2-5 mm long, puberu­
lous, caducous.
Inflorescences usually bisexual, broadly turbinate, convexly discoid or
hemispherical, sometimes lobed, 3- 1 2 mm in d iameter, subsessile or pe­
dunculate; peduncle up to 10 mm long, puberulous, often bracteate and
with uncinate hairs; receptacle almost glabrous to densely puberulous, of­
ten with uncinate hairs; staminate flowers many to several; perianth 0.2-
0.4 mm high, 3-4-lobed, puberulous; stamen I, filament 0. 3-0 . 8 mm long,
anther 0. 1 -0.25 x 0. 1 -0.25 mm, connective broad and swollen; one to se­
veral pistillate flowers; style exceeding the receptacle by 0. 1 -0 . 3 mm, stig­
mas ea. 0. 1 mm long; bracts sparsely to densely puberulous, often with un­
cinate hairs. Fruiting receptacle subglobose, ea. I cm in diameter, yellow
at maturity; seeds ea. I cm long.

D i s t r i b u t i o n: From eastern Brazil ( Rio de J aneiro) to Panama,

and in northern Central America.

In Surinam common in forests of the interior and the savanna belt.

van Asbeck 1 9, 1 9a; BBS 55; BW 56, 57, 58, 58a, 402, 1 1 58, 1 37 1 , 1 378, 1 634, 1 647,
1 85 1 , 29 1 2, 5095, 5429, 5495, 5497, 5498, 5499, 5500, 550 1 , 55 10, 5 536, 5828, 6247,
688 1 ; van Donse1aar 266, 1 686; Essed s. n . ; Gonggrijp s. n . ; 1rwin, Prance, Soderstrom
& Ho1mgren 54583; Lanj ouw & Lindeman 696, 1745; LBB 1 08 1 0 , 1 0992, 1 3260;
Lindeman 3676, 5 360, 5 376, 5473, 5 54 1 , 5 547, 6285, 6647: Maguire 24358, 40774;
01denburger & Norde 427, 5 1 0; Schu1z 7807, 7980, 8 1 38, 8 1 39; S plitgerber 90 1 (L);
Stahel Woodherb. 3, 3a, 2 1 9.

V e r n a c u l a r n a m e s: letterhout, manletterhout, kapiteinhout (Sur.

Dutch), sokone ibiberoe (Arow.), paida ( Car.) .
200 MORACEAE

Blake (Jour. Wash. Acad . Sci. 1 2: 39 1 . 1 922) d istinguished three separate

Piratinera species in Surinam, on account of differences in the indumen­
tum of the lower leaf surface. He recognized P. guianensis with appressed
short hairs (found throughout the distribution area), P. scabridula with
very short stiff patent hairs (occurring in Surinam, Guyana, and Trini­
dad), and P. velutina with longer and weaker, more or less patent, hairs
(found from Colombia through Surinam to Pani) . M ost specimens from
Surinam can be assigned to one of the groups, characterized by the indu­
mentum of the lower leaf surface. These differences are not correlated
with other differences and the taxonomic value appears to be slight.
These groups could be regarded as forms. Brosimum guianense varies
considerably in size, shape, texture, and indumentum of the leaves, and
in the characters of the inflorescences. The species is well characterized
by the occurrence of papillae on the lower surface of the leaf.

4. Brosimum rubescens Taubert, Bot. Jahrb. 1 2 ( Beibl. 27): 4. 1 890.

Alicastrum rubescens (Taubert) Taubert, Bot. Jahrb. 1 5 ( Beibl.
34): 1 6. 1 892.
Piratinera rubescens (Taubert) Pittier, Contr. U . S . Nat. Herb.
20: l OO. 1 9 1 8.

T y p e: G1aziou 1 2 1 69, Brazil, Rio de Janeiro ( B).

Ferolia guianensis Aublet, PI. Gui. Suppl. 7, t. 372. 1 775.

T y p e : Aublet s.n., French Guiana (BM).
Brosimum paraense Huber, Bol. Mus. Emilio Goeldi 6: 67. 1 9 1 0 .
Piratinera paraensis (Huber) Benoist, Arch. B o t . Caen 5(M e m. 1 ) : 4 5 . 1 93 1 .
T y p e : Goeldi s.n. (HAM P 8320), Brazil, Para ( M G, not seen; isotypes
BM, K , P . U).
Brosimum angustifolium Ducke, Arch. Jard . Bot. Rio de Janeiro 3 : 25 . 1 922.
T y p e : Ducke s.n. (HJB R 1 25 36, HA M P 1 6594), Brazil, Para (RB, not seen;
isotypes B, B M , K, P, U, US).
Brosimum lanciferum Ducke, Arch. Jard . Bot, Rio de Janeiro 3 : 24. 1 922.
Piratinera lancifera (Ducke) Benoist, Arch. Bot. Caen 5( M e m. 1): 46. 1 93 1 .
L e c t o t y p e : Ducke s.n. (HA M P 1 6646), Brazil, Para ( RB , not seen;
isotypes BM, F, P, US).
Brosimum caloxylon Standley, Tropical Woods 1 6: 1 9. 1 928 (nomen);
 MORACEAE 20 1

Tropical Woods 1 7: 1 1 . 1 929 (descr.).

T y p e : G . P . Cooper 535, Panama (F).
Brosimum platyneurum Ducke, Arq. Serv. Flor. Rio de Janeiro 1 : 4. 1 939.
T y p e : Ducke s.n. ( H J B R 23622), Brazil, Amazonas ( R B).
Brosimum longistipulatum Ducke, Tropical Woods 90: 7. 1 947.
T y p e : Ducke 1 9 1 6, Brazil, Amazonas ( R B) .
Brosimum brevipedunculatum Ducke, Tropical Woods 90: 8. 1 947.
T y p e : Ducke 1 9 1 8, Colombia, Amazonas ( R B) .

Monoecious trees up to 30 m tall, with white latex. Leafy twigs 1 -2(-3)

mm thick, angulate, sparsely to rather densely puberulous . Leaves ellip­
tic to lanceolate, 3- 1 3 cm long, 1 -6 . 5 cm broad, often broadest at the lo­
wer end, (sub)coriaceous, blunty acuminate to caudate, acute to obtuse
at the base; above glabrous, beneath sparsely appressed puberulous, usu­
ally with oblongoid-capitate pluricellular hairs, glabrescent; costa prom­
inent, other veins (almost) plane, 1 2- 1 8 pairs of secondary veins, without
parallel tertiary veins; petioles 2- 1 3 mm long; stipules connate, fully am­
plexicaul, 8-25 (-3 5) mm long, almost glabrous to appressed puberulous.
Inflorescences usually bisexual, (sub)globose, 3-4 mm in diameter; pe­
duncle 2- 1 2 mm long, puberulous; receptacle puberulous; a few stami­
nate flowers; perianth minute, puberulous; stamens 1 -2, filaments 0 . 2-0. 8
m m long, anthers 0.2-0. 3 x 0. 25-0.4 m m , connective broad and swollen;
pistillate flowers 1 -3 ; style ea. 1 mm long, puberulous, stigmas 0. 5-0 . 7
mm long; some tens o f bracts, 0.4- 1 . 2 mm in diameter, yellowish to white
puberulous. Fruiting receptacle ea. 1 cm in diameter, at maturity red­
dish.

D i s t r i b u t i o n : Amazon Basin, the Guianas, northern Panama, and

near Rio de Janeiro (Brazil) .

In Surinam in forests of the interior and the savanna belt.

BBS 62; BW 1 1 65, 5386, 5600, 5873, 6203, 6703 , 67 1 6, 6890; J unker s.n. or
5 1 2 (herb. Pfeiffer); Lanj ouw 1 004; Lanj ouw & Lindeman 24 1 1 ; L B B 1 1 0 1 6,
1 3427; Oldenburger, Norde & Schulz 1 2 1 9; Schulz 7387; Stahel Woodherb. 94, 1 70,
170a.
202 MORACEAE

V e r n a c u l a r n a m e s : satij nhout (Dutch), satine rubane (French),

doekaliballi ibiberoebaria (Arow.) .

The species is variable i n the indumentum, t h e length o f the stipules, and

the characters of the inflorescences (dimensions, number of flowers, and
bracts). The material from the Guianas is rather uniform; it agrees with
that of the Lower Amazon Basin, but differs more or less from many of
the specimens collected in the Upper Amazon Basin.

5. Brosimum parinarioides Ducke ssp. parinarioides.

Brosimum parinarioides Ducke, Arch. Jard. Bot. Rio de
Janeiro 3 : 27. 1 922.

L e c t o t y p e: Ducke s.n. ( HJ B R 1 25 1 9, H A M P 1 6972), Brazil,

Para ( R B , not seen; isotypes F, P, U) .

Trees up to 40 m tall, with abundant white latex. Leafy twigs 2-6 mm

thick, more or less angulate, minutely white to brown puberulous. Lea­
ves elliptic to oblong to subobovate (to lanceolate), 8-24(-50) cm long, 3-
8(- 1 4) cm broad, coriaceous, acuminate, acute to obtuse to emarginate
at the base; margin entire (to weakly undulate); above glabrous, beneath
with dense, white to yellow to brown, short, straight or curved hairs and
on the costa often some long weak appressed hairs, or not very densely
white to brown hirtellous with dark brown uncinate hairs on the costa
and secondary veins when juvenile; veins plane above, prominent be­
neath, 20-34 pairs of secondary veins, most tertiary veins parallel;
petioles 6- 1 4(-20) mm long; stipules connate, fully amplexicaul, 7-25 mm
long, whitish to brownish puberulous to pubescent to sericeous.
Inflorescences bisexual or pistillate, subglobose, 4-6 mm in diameter,
often foveate; peduncle 1 0-20 mm long, minutely brown puberulous; re­
ceptacle yellow puberulous; staminate flowers many; perianth 0. 2-0. 3
m m high, 4-3-lobed, puberulous; stamen 1 , filament ea. 1 m m long,
anther 0. 2-0.25 x 0. 2-0 . 25 mm, connective broad and swollen; pistillate
 MORACEAE 203

flowers one to several; style exceeding the receptacle by ea. I mm,

stigmas 0.8- 1 . 3 mm long. Fruiting receptacle ea . 1 . 2 cm in diameter; seed
ea. 0. 5 cm lo ng.

D i s t r i b u t i o n : Brazil (north of the Amazon River), Surinam, and

Guyana.

In Surinam in forests of the interior and the savanna belt.

van Donselaar 2337, 3427; van Emden s.n. ( B rownsberg Reserve, tree number
1 3 1 4, 1 8-IX- 1 93 1 , 2-X- 1 93 1 ); Geijskes 1 020; I rwin, P rance, S oderstrom &
Holmgren 55574; Lanjouw & Lindeman 247 1 ; LBB 1 0952, 1 0 1 2 1 ; Lems 5252 (NA);
Lindeman 446 1 ; Oldenburger, Norde & Schulz 1 1 1 8; Pulle 3 1 6; Stahel W oodherb.
1 29.

V e r n a c u l a r n a m e s : doekali (Sur.), doekaliballi (Arow.).

The material shows considerable differences in the indumentum of the ,J ow­

er leaf surface and in the size of the leaves. These d ifferences are proba­
bly due to different ages of the trees. Specimens apparently collected
from young trees have large leaves (up to 50 cm long) with rather sparse
hairs on the veins beneath. M o reover, the costa usually bears dark brown
uncinate hairs. Other, often fertile, specimens have smaller leaves with
dense, curved to straight hairs on the veins beneath, while uncinate hairs
are usually absent. Intermediate forms have been found. The other sub­
species, ssp . amplicoma, is known only from the Upper Amazon Basin.

5 . Trymatococcus Poeppig & Endlicher, Nov. Gen. et Spec. 2: 30. 1 838;

Lanj ouw, Rec. Trav. Bot. Neerl. 32: 262. 1 93 5 ; Berg, Flora
Neotropica M onogr. 7: 208. 1 972.

T y p e s p e c i e s : Trymatococcus amazonicus Poeppig & End­

licher
Lanessania Baillon, Adansonia 1 1 : 298 . 1 875.
T y p e s p e c i e s : Lanessania turbinata Baillon ( Trymatococcus amazoni­
=

cus Poeppig & Endlicher).

204 MORACEAE

M onoecious trees. Leaves distichous, pinnatinervate; stipules free, not

fully amplexicaul. Uncinate hairs common . Inflorescences in pairs or soli­
tary in the axils of the leaves, bisexual; receptacles cylindrical to turbi­
nate, with variously shaped, basally attached to peltate bracts; a few to
many staminate flowers on the upper part of the receptacle, perianth 3-
lobed (to 3-fid), stamens 3 , straight in the bud, anthers introrse, pistillode
present; one pistillate flower embedded in the centre of the receptacle,
ovary adnate to the perianth, stigmas vittiform. Fruit adnate to the enlar­
ged receptacle; seed big, testa with a thickened part, not vascularized, em­
bryo transverse in the fruit, cotyledons thick, usually more or less un­
equal, radicle minute.

D i s t r i b u t i o n: 3 species in the Amazon Basin and the Guianas.

Lanj ouw treated the genus in 1 935. At that time, only T o/igrandrus was
known from Surinam. The other species, T amazonicus and T para­
ensis, although closely related, can always be separated satisfactorily.

a. Leaves almost glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 . T . oligandrus

b. Leaves hairy on the costa above and on the veins beneath . . . . . . . . . . . . 2 .

2 a. Leaves coriaceous (to subcoriaceous), often more or less bullate above,

usually shorter than 10 cm . . . . . . . . . . . . . . . . . . . . . . . . . 2. T. paraensis
. . .

b. Leaves chartaceous (to subcoriaceous), never bullate above, usually

longer than 10 cm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I . T . amazonicus

I . Trymatococcus amazonicus Poeppig & Endlicher, Nov. Gen. et Spec.

2: 30, t. 1 42. 1 838.

T y p e : Poeppig s.n. (or 2 1 67), Peru, Loreto ( B).

Lanessania turbinata Spruce ex Baillon, Adansonia 1 1: 298 . 1 875.

Trymatococcus turbinatus (Spruce ex Baillon) Ducke, Arch. Jard . Bot.
Rio de Janeiro 3: 23 . 1 922.
T y p e : Spruce 1 825, Brazil, Amazonas ( P).

Trees up to 15 m tall, with yellowish to colourless latex. Leafy twigs 1 -5

mm thick, brown hirtellous to subvelutinous, often with uncinate hairs in-
 MORACEAE 205

termixed . Leaves oblong to lanceolate, 7-22(-32) cm long, 3-8(- 1 1 ) cm

broad, chartaceous, acute to acuminate to caudate, acute or obtuse at the
base; margin entire; above glabrous except on the costa, beneath puberu­
lous to hirtellous, often uncinate hairs on the costa; veins slightly
prominent to slightly impressed above, prominent beneath, 5- 1 3 pairs of
secondary veins, most tertiary parallel; petioles 6- 1 8(-25) mm long,
brown (sub)velutinous, often with uncinate hairs intermixed; stipules
3-7 mm long, appressed pubescent.
Inflorescences: peduncle 3-7 mm long, shortly brown (sub)velutinous,
with uncinate hairs intermixed ; receptacle narrowly turbinate to cylin­
drical, ea. 6 mm high, shortly brown velutinous, often also with uncinate
hairs, with variously shaped basally attached or peltate bracts; soms tens
of staminate flowers; perianth 3-lobed to 3-fid, puberulous, often with
short uncinate hairs; stamens 3( -2), ea. 1 mm long, filaments broad to nar­
row, pistillode small; style 3-3 . 5 mm long, stigmas 3-4 mm long. Fruiting
receptacle globose, ea. 20 mm in d iameter, coronate because of the stami­
nate flowers; seed ea. 1 2 mm in diameter, cotyledons slightly to very un­
equal.

D i s t r i b u t i o n : Upper Amazon Basin (Brazil, Peru, and Venezuela),

Surinam, and French Guiana.

In Surinam in forests of the interior and the savanna belt.

van Donselaar 1 272; Lindeman 377 3, 4735, 4928, 5042, 6442, 6762; LBB 1 0832,
1 33 1 0.

V e r n a c u l a r n a m e s: manletterhout, letterhout (Sur. Dutch).

The specimens cited under Trymatococcus amazonicus are sterile . Be­

cause of their leaf characters they are included in this species. H owever,
there are indications that they may have been collected from j uvenile
specimens of T. paraensis.
206 MORACEAE

2. Trymatococcus paraensis Ducke, Arch. J ard. Bot. Rio de J aneiro

3 : 22. 1 922.

T y p e : Ducke s . n . ( H J B R 1 2500; H A M P 1 6560), Brazil, Para

( R B , not seen; isotypes B, B M , K, P, U ) .

Trees u p t o 3 0 m tall. Leafy twigs l -5 m m thick, brown hirtellous to sub­

velutinous, often with uncinate hairs. Leaves (subrotundate to) elliptic to
oblong, 2- 1 2 cm long, 1 . 5-5 .5 cm broad, often broadest above the middle,
coriaceous to subcoriaceous, acuminate to mucronate, acute to obtuse at
the base; margin entire; above hirtellous on the costa, for the rest gla­
brous or with a few uncinate hairs, beneath hirtellous on the veins, someti­
mes uncinate hairs on the costa; the costa impressed, the other veins
plane to imp ressed above, veins prominent beneath, 5- l l pairs of second­
ary veins, tertiary veins usually partly parallel; petioles 3- 1 6 mm long,
subvelutinous to hirtellous, often with uncinate hairs; stipules 3-4 mm
long, appressed pubescent.
Inflorescences: peduncle 3-5 mm long, shortly brown (sub)velutinous,
often with uncinate hairs, receptacles narrowly turbinate to cylindrical, 4-
6 mm high, shortly brown velutinous, with variously shaped basally at­
tached or peltate bracts; staminate flowers some tens; perianth 3-lobed,
puberulous; stamens 3, 1 -2 mm long, filaments broad to narrow; pistil­
lode subulate, very small; style 4-5 mm long, stigmas 3-7 mm long.
Fruiting receptacle globose, ea. 1 5 mm in diameter, coronate because of
the staminate flowers, subvelutinous; seed ea. 9 mm in diameter, cotyle­
dons very unequal.

D i s t r i b u t i o n : Brazil ( Para), Surinam, and Guyana.

In Surinam only known from the Nassau M ountains, the Wayombo

River, and the Jodensavanne - M apane Creek area.

Lanjouw & Lindeman 2 177; LBB 1 3 365; Schu1z 7887; Stahe1 Woodherb. 366.
 MORACEAE 207

3. Trymatococcus oligandrus ( Benoist) Lanj ouw, Rec. Trav. Bot. Neerl.

32. 27 1 . 1 93 5 .
Lanessania oligandra Benoist, Bull. M us . H ist. Nat. Paris 27:
1 99. 1 92 1 .

T y p e : Benoist 1 573, French Guiana ( P) .

Trees u p to 2 0 m tall, with cream-coloured latex, turning red . Leafy twigs

0. 5-2 mm thick, appressed puberulous, often with uncinate hairs. Leaves
elliptic to lanceolate, 2- 1 2 mm long, 1 -4 cm broad, subcoriaceous, acumi­
nate to caudate, acute (to obtuse) at the base; margin entire; above at fi rst
with uncinate hairs, glabrescent, beneath sparsely appressed puberulous,
glabrescent; veins more or less prominent, 5- l 0 pairs of secondary veins,
tertiary veins not parallel; petioles 4-8 mm long, puberulous, often with
uncinate hairs; stipules ea. 2 mm long, appressed puberulous.
Inflorescences: peduncle l -6 mm long, densely puberulous to shortly
velutinous, often with intermixed uncinate hairs; receptacle cylindrical
to narrowly turbinate, 5-7 mm high, shortly velutinous, sometimes with
uncinate hairs intermixed , with variously shaped basally attached or pel­
tate bracts; staminate flowers 2-5 ; perianth 3-lobed, shortly velutinous to
puberulous; stamens 3 , filaments ea. I mm long, broad, pistillode su­
bulate; style ea. I mm long, stigmas ea. 1 . 5 mm long, hardly exceedi ng
the perianth. Fruiting receptacle 1 . 5 cm in diameter, yellow at maturity;
seed globose ea. 8 mm in diameter, cotyledons slightly to very unequal.

D i s t r i b u t i o n : French Guiana and S urinam.

In Surinam in forests of the interior and the savanna belt; apparently

confined to the eastern part of the country.

Boschbeheer s . n . ; BW 1 365, 34 17, 6098, 6645, 6823, 7236; van Donselaar 1 857,
2049, 2 1 74, 2 1 97, 222 1 , 3323; Essed s.n.; Lanjouw & Lindeman 2297; Lindeman
5934; Stahel Wilhelm. Geb. Exp. 50.
208 MORACEAE

6. Pseudolmedia Trecul, Ann. Sci. Nat. Bot. 1 1 I. 8 : 1 29 . 1 847 ; Berg,

Flora N eotropica Mo nogr. 7: 1 7 . 1 972.

L e c t o t y p e s p e c i e s : Pseudolmed i a ferrugi nea ( Poeppig &

Endlicher) Trecul ( = Pseudolmedia laevis ( R uiz & Pavan)
M acbride).

Olmediopsis Karsten, Fl. Columb. 2: 1 7 . 1 862.

T y p e s p e c i e s : Olmediopsis obliqua Karsten ( Pseudolmedia laevis ( Ruiz
=

& Pavon) Macbride).

D ioecious trees . Leaves distichous, pinnatinervate; stipules free, fully am­

plexicaul . Inflorescences on short shoots in the axils of the leaves, unisex­
ual, sessile, involucrate. Staminate inflorescences with discoid recepta­
cle, free stamens, intermixed with concentrically arranged bracts; sta­
mens straight in the bud . Pistillate inflorescences uniflorous; perianth tu­
bular; ovary adnate to the perianth, stigmas filiform; fruiting perianth
fleshy and coloured; seed big, without endosperm, testa vascularized, co­
tyledons thick, equal, radicle short and apical.

D i s t r i b u t i o n: 9 species in tropical America.

At present only the widespread P. /aevis IS known from Surinam,

but P. /aevigata is expected to occur there.

I. Pseudolmedia laevis ( Ruiz & Pavan) M acbride, Publ. Field M us. Bot.
1 1 : 1 6. 1 93 1 .
Olmedia laevis Ruiz & Pavan, Syst. Veg. FI. Peruv. et Chi!. I :
258. 1 798.

L e c t o t y p e : Ruiz & Pavon s.n. o, Peru ( H A L) .

Olmedia ferruginea Poeppig & Endlicher, Nov. Gen. et Spec. 2 : 3 1 , t . 1 43 .

1 838.
Pseudolmedia ferruginea (Poeppig & Endlicher) Trecul, An n . Sci. Nat.
Bot. I l l . 8 : 1 3 1 . 1 847 .
 MORACEAE 209
---- - ------- · -------- ····--- - - - - --··----- -------

T y p e : Poeppig s.n. o, Brazil, Amazonas (W, destroyed , replaced by P).

Olmediopsis obliq ua Karsten. Fl. Columb. 2: 1 7 , t. 1 09 . I S62.
Pseudolmed ia obliqua ( K arsten) Renner, Bot. Jahrb. 39: 375. 1 907.
T y p e : K arsten s .n . o, Colombia, M eta ( L E).
Olmed iopsis lanceo late Karsten, Fl. Columb. 2: 1 7 . 1 8 62.
T y p e : K arsten s . n . d . Colombia, Meta ( L E).
Pseudolmed ia hi rsuta Ba illon, Andansonia 1 1 : 295. 1 8 75.
T y p e : Triana 855 0 . Colombia, Meta ( P).
Pseudolmed ia sagot i Benoist , Bull. H ist. Nat. Paris 3 1 : 468 . 1 925.
L e c t o t y p e : Benoist 1 204 9 , French G u iana ( P).
Pseudolmedia multinerv is M ildbread, Notizbl. Bot. Gart. Berlin 1 0: 1 89. 1 872.
T y p e : Tessmann 4697 o , Peru , Amazonas ( B).
Pseudolmed ia hirtellaefolia R usby, Mem. N.Y. Bot. Gard. 7 : 228. 1 927.
T y p e : White 1 495 o , Bolivia, Beni (NY).
Pseudolmedia alnifolia R usby, Mem. N . Y . Bot . Gard. 7 : 229. 1 927.
T y p e : White 1 492 9, Bolivia, Beni (NY).
Olmedia caurensis Pittier, Bol. Soc. Venez. Ci. Nat. 8 : 306. 1 942.
T y p e : Williams 1 20 1 8 9, Venezuela, Bolivar (VFN).

Trees up to 40 m tall, with white latex. Leafy twigs 1 -3 mm thick,

sparsely puberulous, with longer yellow hairs intermixed . Leaves oblong
to lanceolate, 8-2 1 cm long, 3-8 cm broad , somewhat inequilateral,
chartaceous to subcoriaceous, acuminate to caudate, acute to obtuse
or sometimes emarginate at the base; margin entire or dentate towards
the apex; nearly glabrous above, sparsely pubescent on the veins
beneath; veins slightly prominent to plane above, prominent beneath,
1 2- 1 7 pairs of secondary veins, most tertiary veins parallel; petioles
3- 1 0 mm long, not or slightly canaliculate, (sparsely) puberulous, with
longer yellow hairs intermixed ; stipules 3-8 mm lo ng, with yellow,
appressed or patent hairs.
Staminate inflorescences usually up to 4 together; involucre with
4-5 series of deltoid to ovate to spathulate, obtuse to acute, subsericeous
bracts; filaments 0 . 5- 1 .0 mm long, anthers 1 . 2-2.0 x 0 . 3-0 . 5 mm, usually
inequilateral and provided with apical hairs; interstaminal bracts
narrowly spathulate, 2-4 mm long.
Pistillate inflorescences usually in pairs; involucre with 2-4 series
of reniform to ovate, obtuse to acute, subsericeous bracts; perianth ea.
2 mm high, subsericeous; style 1 - 1 . 5 mm long. stigmas 7- 1 1 mm long.
210 M ORACEAE

Fruiting perianth ellipsoid to obl ongoid , 1 0- 1 3 mm h ig h , yellow

hirtellous to hirsute; seed 7-9 m m l o ng.

Distribution: N o rt h e r n S o u t h A me r ica , east o f t h e A nd e s .

In S u r i n a m i n fo rests o f t h e i n t e r i o r a nd t h e sava n na be l t .

B W 4%5, 4966. 4990; L A B 1 1 1 90. 1 3 363. 1 4302; Lems 5235( N A ) ; Lindeman 5034;
S..:hulz 7X48. X 1 63 ; S t a h e l W oodherb. 96a .

V e r n a c u l a r n a m e s : letterh o u t , m a n lette rh ou t ( S ur. Dutch).

The Surinam specimens d iffer from the more typical specimens of this
species from the Upper Amazon Basin in the relatively large chartaceous
and almost glabrous leaves and in the slender sparsely hairy twigs. They
agree well with specimens collected in north-eastern Venezuela. Speci­
mens from French Guiana resemble those of the Lower Amazon Basin
more than the Surinam ones.

7. Perebea Aublet, PI. Gui. 2: 953. 1 775; Berg, Flora Neotropica

Monogr. 7: 3 8 . 1 972.
M ikania Necker, Elem. Bot. 2: 2 1 7. 1 790, as synonym.

T y p e s p e c i e s : Perebea guianensis Aublet

Noyera Trecu1, Ann. Sci. Nat . Bot . I l l . 8: 1 3 5. 1 847 .

T y p e s p e c i e s : Noyera rubra Trecu1 ( Perebea mollis ( Poeppig &
=

End 1icher) Huber ssp. rubra (Trecu1) C. C. Be rg).

Dioecious or monoecious trees or shrubs . Leaves distichous, pinnatiner­

vate; stipules free, fully amplexicaul . Inflorescences on short shoots in
the axils of the leaves, unisexual, discoid, pedunculate or sessile, involuc­
rate. Staminate inflorescences many-flowered (in Surinam); perianth
with 2-4 free or connate tepals; stamens 2-4, straight or slightly·incurved
in the bud . Pistillate inflorescences several to many-flowered (in
 M O RACEAE 21 1
-------

Surinam); perianth tubular, entire to 4-lobed or 4-fid; ovary free or

basally adnate to the perianth, stigmas ea. semidisciform to filiform.
Fruiting perianth fleshy, red(dish); fruit free or more or less adnate to
perianth; seed big, without endosperm, testa vascularized , cotyledons
thick, equal, free, rad icle short and apical.

D is t r i b u t i o n : 8 species in northern South America, extending to

Costa Rica.

I a. Twigs sericeo us; stipules 2-4 cm long; pluricellular hairs oblongoid-capitate;

stigmas ea. semidisciform; staminate flowers with 4 free tepals and 4
stamens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I . P. guianensis ssp. guianensis
b. Twigs not sericeous; stipules usually shorter than 2 cm; pluricellular
hairs globose-capitate; stigmas filiform; staminate flowers with 2-3 connate
tepals and 2-3 stamens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. P. mollis

l . Perebea guianensis Aublet ssp. guianensis

Perebea guianensis Aublet, PI. Gui. 2: 953, t. 3 6 1 . 1 775.

T y p e : Aublet s.n. <;? , French Guiana (not seen, obscure) .

Olmedia? grandifolia Trecul, Ann. Sci. Nat. Bot. I I I . 8: 1 28 . 1 847.

T y p e : Martin s.n. o , French Guiana ( P) .
Castilloa australis Hemsley, H o o k . l e . PI. I V . t. 2696. 1 90 1 .
Perebea australis ( Hemsley) Macbride, Pub!. Field M us. Bot. 1 1 : 1 7. 1 93 1 .
T y p e : Pearce s.n. <;? , Peru (K).
Olmed ia habas Pax, Repert. Sp. Nov. 7: 1 08 . 1 909.
T y p e : Buchtien 1 567 0 , Bolivia, La Paz (B).

Trees usually small, up to 20 m tall. Leafy twigs yellow to greyish seri­

ceous. Leaves oblong to lanceolate, 22-48 cm long, 7- 1 7. 5 cm broad,
chartaceous to coriaceous, acuminate, (obtuse to) emarginate to cordate
at the base; margin often dentate; sparsely hairy above, sericeous on the
veins beneath; veins almost plane above, prominent beneath, 1 7-22 pairs
of secondary veins, most tertiary veins parallel; petioles 5- 1 6 mm long;
stipules 2-3 mm long, sericeous; pluricellular hairs oblongoid-capitate .
Staminate inflorescences 6- 1 0 mm in diameter; peduncle 4-8 mm long;
212 M ORAC EAE
--- -- ---·-- -· --- -------- ·- -

involucre with 4-7 series of deltoid to ovate, acuminate to acute yellowish

subsericeous bracts; flowers numerous; perianth 0.8- 1 .0 mm high, with 4
free hirtellous tepals; stamens 4, filaments 0.8- 1 .0 mm long, often broad­
ened at the base; anthers 0.25-0. 3 x 0 . 2-0. 3 mm.
Pistillate inflorescences solitary, 8-30 mm in diameter, subsessile or
ped unculate; ped uncle up to lO mm long; involucre with 7- 1 5 series of
deltoid to ovate, acuminate to acute, yellowish subsericeous bracts; flo­
wers 1 0 or more; perianth 3-6 mm high, 4-dentate to 4-fid, yellowish
hirtellous to velutinous; style 2-2.5 mm long, stigmas ea. semidisciform,
0 . 8- 1 . 3 mm long. Infructescences 2-5 cm in diameter, fruiting perianth
1 0-22 mm high, yellowish hirtellous to velutinous.

D is t r i b u t i o n : E xtending from the Amazon Basin through French

Guiana to Surinam.

In Surinam probably only in forests near the eastern boundary.

L B B 1 1 976; Lindeman 4075, 5949.

2. Perebea mollis ( P oeppig & Endlicher) H uber, Bol. M us. Emilio Goeldi
5: 334. 1 909 .
Olmedia mollis Poeppig & End licher, N ov. Gen. et S pec. 2:
3 1 , t. 1 49 . 1 83 8 .
N oyera mollis ( P oeppig & Endlicher) Ducke, Arch. J ard . Bot.
Rio de Janeiro 3: 37. 1 922.

T y p e : Poeppig s.n. (or 3070) o, Brazil, Amazonas (W,

destroyed, replaced by P).

Trees usually dioecious, up to 30 m tall, with yellow latex. Leafy twigs

1 . 5-7 mm thick, with very short patent hairs and distinctly longer,
appressed to patent, whitish to yellow hairs. Leaves elliptic to oblong, 4-
36 cm long, 4- 1 4 cm broad , often broadest below the middle, · slightly
inequilateral, coriaceous to chartaceous, acuminate to mucronate,
 M O RACEAE 213

obtuse to subcordate at the base; margin entire or dentate; above hirsute

to strigose, except for the costa glabrescent, more or less bullate and
scabrous or not so, beneath hirsute to puberulous ; veins slightly promi ­
nent to somewhat impressed above, prominent beneath, 1 2-22 pairs o f
secondary veins, most tertiary veins parallel; petioles 1 -9 mm long,
stipules 5-25 mm long, yellow subsericeous to hirsute, sometimes subper­
sistent; pluricellular hairs globose-capitate .
Staminate inflorescences 8- 1 6 mm in diameter; peduncle 1 0-30 mm
long; involucre with 6-9 series of deltoid to ovate to linear, caudate to
acute, appressed puberulous to hirtellous bracts, the inner ones up to 8
mm long and incurved before anthesis; flowers many; perianth 1 . 7-2. 5
mm high, 2-3-lobed or 2-3-fid, puberulous; stamens 2-3 , filaments 2-3
mm long, anthers 0 . 8- 1 . 3 x 0.2-0. 3 mm.
Pistillate inflorescences solitary, occasionally accompanied by 1 -2
staminate ones, 8- 1 2 mm in diameter, subsessile to pedunculate; pedun­
cle up to 5 mm long; involucre with 4-7 series of deltoid to ovate to lanceo­
late, caudate to acute, appressed pubescent to hirtellous bracts, the inner
ones longest and incurved before anthesis; flowers 6-30, free or basally
connate; perianth 2-3 mm high, tubular, entire to 4-dentate, hirtellous ,
setose near the base or not s o ; style ea. 1 . 5 m m long, stigmas filiform and
2-3 mm long. Infructescences 1 . 5 -2.5 cm in diameter; fruiting perianth
hirtellous to hirsute .

There are two (almost?) allopatric subspecies; both have been collected
in Surinam.

2a. Perebea mollis ( Poeppig & Endlicher) H uber ssp. mollis.

Perebea paraensis Huber, Bol. M us. Emilio Goeldi 5: 3 34. 1 909.

T y p e : Ducke s.n. ( H A M P 4905) o', B razil, Para ( M G not seen; isotypes
BM, G, RB).
Perebea lecointei Huber, Bol. M us. Emilio Goeldi 5 : 334. 1 909.
T y p e : Ducke s. n . ( H A M P 6942) <(, Brazil, Para ( M G, not seen).
214 MORACEAE

Twigs with long ascending to patent hairs. Leaves chartaceous to

subcoriaceous, usually more or less distinctly bullate and scabrous
above, more or less densely hirsute, occasionally rugose, beneath.
P istillate flowers free or slightly connate at the base; perianth with
distinctly longer hairs towards the base; fruiting perianth not or slightly
connate at the base, hirsute.

D is t r i b u t i o n : The Amazon Basin ( B razil, Peru and B olivia), also

in southern Surinam; in non-inundated forests.

In S urinam known from a single collection collected in the Sipaliwini

savanna area near the B razilian frontier.

Oldenburger, Norde & Schulz 1 1 37.

2b. Perebea mollis ( Poeppig & Endlicher) H uber ssp. rubra (Trecul)
C. C. Berg, Acta Bot. Neerl. 1 8: 463 . 1 969.
Noyera rubra Trecul, Ann. Sci. Nat. Bot. I l l . 8 : 1 36. 1 847.

T y p e : Martin s.n. o, French Guiana (P).

Castilla ulei Warburg f. lecithogalacta R.E. Schultes, Bot. Mus. Leafl. 1 2 :

1 28 . 1 946.
Perebea 1ecithoga1acta ( R . E. Schultes) R . E . Schultes, Bot. Mus. Leafl. 1 3 :
298. 1 949.
T y p e : Schultes 5798 <j>, Colombia, Vaupes (ECON, not seen; isotypes
K, US).

Twigs with appressed to more or less patent hairs. Leaves coriaceous to

subcoriaceous, above slightly or not bullate and slightly or not scabro us,
beneath rather sparsely puberulous to hirtellous and mostly rugose, on
the costa longer, more or less apressed hairs.
P istillate flowers basally connate; perianth with short, dense, equally
long hairs; fruiting perianths connate, hirtellous.
 MORACEAE 215

D i s t r i b u t i o n : French G uiana, Surinam, and the U pper Amazon

Basin north of the Amazon R iver.

In Surinam in forests of the interior and the savanna belt.

BW 1 244, 1 786, 1 957. 5493, 6432; LBB 1 3292; Stahel Woodherb. 2 1 1 .

Vernacular names: letterhout (Sur. Dutch), sokone ( Arow.).

8 . Maquira Aublet, PI. Gui. Suppl. 36. I 775; Berg, Flora Neotropica
M o nogr. 7: 62. 1 972.

T y p e s p e c i e s : Maquira guianensis Aublet

Olmediophaena Karsten, Bot. Jahrb. 8: 397. 1 887.

T y p e s p e c i e s : Olmediophaena coriacea (Karsten) Karsten (= M aquira cori­
acea ( Karsten) C. C. Berg).
Olmedioperebea Ducke, Arch. Jard . Bot. Rio de Janeiro 3: 33. 1 922.
T y p e s p e c i e s : Olmedioperebea sclerophylla Ducke (= M aquira sclerophylla
( Ducke) C. C. Berg).

Trees, mostly dioecious. Periderm of the twigs peeling off easily. Leaves
distichous, pinnatinervate entire, greenish when dried; stipules free, not
fully amplexicaul. Inflorescences one to several on short shoots in the
axils of the leaves, unisexual, discoid, involucrate. Staminate flowers
several to many, perianth 4-lobed to 4-parted , stamens 4(-2), straight or
slightly incurved in the bud . Pistillate flowers one to many, free or basally
connate; perianth 4(-2)-lobed or 4(-2)-fid ; ovary almost entirely adnate
to the perianth, stigmas ea. semidisciform or linguiform. Fruit adnate to
the coloured perianth; seed big, without endosperm, testa vascularized,
cotyledons free, thick, equal, radicle short and straight.

D i s t r i b u t i o n : 5 species in northern South America, extending to

Nicaragua.
216 MORACEAE
���- � �� --- -- - ----- ------

a. Tertiary veins of the leaves parallel; pi stillate inflorescences with one or

a few connate flowers . . . . . . . ....
. . . .. .... . .
. . . . . 2. M . sclerophylla
. . .

b. Tertiary vei ns of the leaves not parallel; inflorescences with several to many
free flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I . M. guianensis
. . .

I . Maquira guianensis Aublet, PI. Gui. Suppl. 36, t. 389. 1 775.

Olmedia maquira Steude1, Nomencl. (ed. 2) 2 1 0. 1 84 1 .
01media guianensis (Aublet) Trecu1, Ann. Sci. Nat. Bot. I l l . 8 :
1 29. 1 847.

T y p e : Aublet s.n. d, French Guiana ( B M ).

Perebea laurifolia Trecul, Ann. Sci . Nat. Bot. I l l . 8: 1 33 . 1 847.

T y p e : Martin s.n., French Guiana (P).

Trees up to 25 m tall, with yellowish to white latex. Leafy twigs 1 -4 mm

thick, puberulous. Leaves elliptic to lanceolate, 4- 1 8 cm long, 1 . 5-6 cm
broad , broadest above the middle, coriaceous to subcoriaceous, acumi­
nate, acute to obtuse at the base; margin entire; glabrescent; veins more
or less prominent, 1 0- 1 4( - 1 8) pairs of secondary veins, tertiary veins not
parallel; petioles 5- 1 5(-20) mm long; stipules l -3 . 5 mm long, puberulous .
Staminate inflorescences up to 3 together, 8- 1 5 mm in diameter; pedun­
cle 5-20 mm long; involucre with (3-)5-6 series of deltoid to ovate, acute
to obtuse, puberulous bracts; flowers many; perianth ea. 0.5 mm high, (3-
4-lobed, puberulous to tomentellous; stamens 4, filaments 0. 5-0 . 6 mm
long, anthers ea. 0 . 3 x 0 . 3 mm, a tuft of hairs in the centre.
Pistillate inflorescences solitary or sometimes accompanied by l -2
staminate ones, l -2 cm in diameter; peduncle 5- 1 2 mm long; involucre
with ea. 5 series of deltoid to ovate to subrotundate, acute to acuminate,
densely puberulous bracts; ea. 1 5-25 free flowers; perianth 1 . 3-3.0 mm
high, 4-fid, brown velutinous; style ea. 1 . 5 mm long, swollen, densely
puberulous, stigmas ea. semidisciform, ea. 1 mm long. Infrutescences 2.5-
4.5 cm in diameter; fruiting perianth oblongoid, ea. 1 . 5 cm high, brown
velutinous.
 MORACEAE 217

D i s t r i b u t i o n : Brazil ( Para and Roraima Territory), French Guiana,

and Surinam.

In Surinam in forests of the interior and the savanna belt; apparently

common.

B W 1 6 1 , 1 39 1 , 4984, 5897, 6326; Daniels & Jonker 1 083; van Donselaar 1 009,
1 054, 1 378, 1 875, 22 1 9, 2354, 3347; Lanjouw & Lindeman 2 1 87, 228 1 , 2446,
28 1 1 ; LBB 8980, 1 0795, 1 3394; Lems 5263 (NA); Lindeman 3574, 36 1 2, 3650,
3670, 4757, 4775, 4825, 6233, 6700; Oldenburger, Norde & Schulz 1 1 99; Schulz
7342, 7982; Stahel Exp. Wilh. Geb. 46, Woodherb. 222.

V e r n a c u l a r n a m e s : manletterhout (Sur. Dutch), sokone (Arow).

This taxon, which is well-known under the name Perebea laurifo/ia, has
had to be removed from Perebea and must now bear Aublet's name
Maquira guianensis.

2. Maquira sclerophylla (Ducke) C. C. Berg, Acta Bot. Neerl. l 8 : 463 . 1 969.

Olmedioperebea sclerophylla D ucke, Arch. Jard. Bot. Rio de
Janeiro 3: 34. 1 922.

L e c t o t y p e : D ucke s.n. ( H J B R 8297; H A M P 1 7 1 1 9) o, B razil,

Para (RB, not seen; isotypes B, B M , NY, P, U, US).

Perebea xinguana Standley, Publ. Field M us. B o t . 1 7: 1 80. 1 937.

T y p e : K rukoff 1 1 33 c) , Brazil, Pani (NY).

Trees medium-sized to tall. Leafy twigs 3-6 . 5 mm thick, puberulous .

Leaves elliptic to lanceolate, 1 3-30(-38) cm long, 5- 1 5 cm broad, often
broadest above the middle, coriaceous (to chartaceous), acuminate,
obtuse (to acute) at the base; margin entire; glabrescent above, often
scabridulous beneath; veins prominent, 1 3-20 pairs of secondary veins,
most tertiary veins parallel; petioles 8-25 mm long; stipules 5- l 0(- 1 7) mm
long, puberulous.
218 MORACEAE

Fig. 4 Maquira. leaves and pistillate inflorescences; a., M. sclerophylla (N.T. Silva
1 4 1 8) ; b. , M. guianensis ( Froes 20785).

Staminate inflorescences up to 4 together, 3- 1 2 mm in diameter; pedun­

cle 5 1 3 mm long, sometimes bracteate; involucre with 3 4 series of
- -

broadly ovate, acute, puberulous bracts; flowers several to many, free or

basally connate; perianth 1 . 5-2.2 mm high, 4-lobed to 4-parted, puberu­
lous, outer tepals more or less cucullate; stamens 4( -3), filaments 2. 2-2. 7
mm long, anthers 0.8- 1 . 3 x 0.3-0 . 5 mm, usually inequilateral.
 MORACEAE 219

Pistillate inflorescences solitary o r accompanied by l -2 staminate

ones, subsessile or pedunculate, peduncle up to 8 mm long; involure
with 3 series of reniform to ovate, acute to abtuse, puberulous bracts; one
or 2-4 connate flowers; perianth 5-8 mm h igh, 4-lobed , puberulous
style short and broad , stigmas linguiform, 2.5-3 . 5 mm long. acute .
Fruiting perianth ea. 20 mm high, sparsely puberulous.

Distribution: Lower Amazon Basin (Brazil) and Surinam.

I n Surinam two collections known from Jodensavanne-Mapane

Creek area.

LBB 1 3386; Lindeman 5040.

9. Helicostylis Trecul, Ann. Sci. Nat. Bot . I I I . 8 : 1 34. 1 847; Berg, Flora
Neotropica M o nogr. 7: 75. 1 972.
Greeneina 0. Kuntze, Rev. Gen. Pl. l : 628 . 1 89 1 .

T y p e s p e c i e s : Helicostylis tomentosa ( Poeppig & Endlicher) R ushy.

Dioecious or monoecious trees. Leaves distichous, pinnatinervate;

stipules free, not fully amplexicaul. I nflorescences on short shoots in the
axils of the leaves, unisexual, pedunculate or sessile, involucrate. Stami­
nate inflorescences with a discoid receptacle; perianth 4-lobed to 4-
parted; stamens 4, often distinctly 2-merous, straight in the bud. Pistil­
late inflorescences with a discoid to hemispherical receptacle, several to
many-flowered; perianth 4-lobed to 4-parted, inner tepals cohering by
weak entangled thin hairs on the inner surface; ovary almost free, stigmas
vittiform to filiform, twisted or not so. Fruit free or almost so; seed
big, without endosperm, testa vascularized, cotyledons thick, equal,
free, radicle straight and apical.

D i s t r i b u t i o n : 7 species in tropical South America, exte nding to

Costa Rica.
220 MORACEAE

I a. Indumentum of the twigs pale yellow, leaves often dentate to denticulate;

pistillate inflorescences with 1 5-30 flowers, long pedunculate; stamens
narrow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. H. pedunculata
b. lndumentum of the twigs brownish, leaves rarely denticulate; pistillate
inflorescences with 5- 1 2 flowers, short pedunculate or sessile; stamens
relatively broad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I . H. tomentosa

1 . Helicostylis tomentosa ( Poeppig & Endlier) R ushy, Mem. Torrey Club

6: 1 20. 1 896.
Olmedia tomentosa Poeppig & Endlicher, N ov. Gen. et Spec. 2:
32, t. 1 45 . 1 83 8 .

T y p e : Poeppig s . n . o · , Brazil , Amazonas (W, destroyed , re­

placed by P).

Olmedia poeppigiana Martius, Flora (or Bot. Zeit.) 24(Beibl. 2): 93. 1 84 1 .
Helicostylis poeppigiana ( Martius) Trecul, Ann. Sci. Nat. Bot. I l l . 8 :
1 34, 1 847.
Greeneina poeppigiana ( Martius) 0. Kuntze, Rev. Gen. PI. 1 : 628. 1 89 1 .
T y p e : Martius 629 o , Brazil, Bahia ( M ).
Helicostylis affinis Steudel ex M iquel, Fl. Bras . 4( 1 ): 1 1 8 . 1 85 3 .
Greeneina affinis ( M i q . ) 0 . Kuntze, Rev. G e n . P I . 1 : 6 2 8 . 1 89 1 .
T y p e : H ostmann & Kappler 1 280 6 , Surinam ( U ) .
Helicostylis obtusifolia Standley, Bull. Torrey Club 58: 3 5 6 . 1 93 1 .
T y p e : Tate 389 o , Venezuela (NY).
Olmed ia polycephala Pittier, Bot. Soc. Venez. Ci. Nat. 7: 306. 1 942.
T y p e : Cardona 4 1 5 o . Venezuela (YEN).
Helicostylis podogyne Ducke, Bot. Tee. I nst. Agron. Belem 4: 3 . 1 945.
L e c t o t y p e : Ducke 1 202 9, Brazil, Amazonas ( R B).
H elicostylis duckei H awkes, Phytologia 3: 3 1 . 1 948 .
·
T y p e : Ducke 1 202 9, Brazil, Amazonas ( N Y).
Olmedia asperula Standley, Bull . Torrey Club 75: 299. 1 948.
T y p e : Fanshave 1 48 (FD 2757) 6 , Guyana ( F).
Trymatococcus guanabarinus A. P . Duarte, Rodriguesia 23-24 (35-36):
55. 1 964.
T y p e : Duarte 5658, Brazil, Guanabara ( R B, not seen; type fragm. U ) .

Dioecious or monoecious trees up to 30 m tall, with yellowish latex.

Leafy twigs 1 -7 mm thick, brownish puberulous to velutinous to tomen­
tose to tomentetlous. Leaves oblong to lanceolate, 5-32 cm long, 2- 1 5 cm
broad, sometimes broadest above the middle, more or less inequilateral,
 MORACEAE 22 1

chartaceous to subcoriaceous, acuminate to mucronate, acute to obtuse

at the base; margin entire, weakly undulate or rarely denticulate towards
the apex; above tomentose to puberulous on the costa, otherwise puberu­
lous, glabrescent to scabridulous, beneath puberulous to hirtellous to
tomentose to tomentellous; veins almost plane above, prominent
beneath, 8-1 7 pairs of secondary veins, often many parallel tertiary veins;
petioles 4- 1 7 mm long; stipules 3- 1 0 mm long, subsericeous .
Staminate inflorescences up to 1 5 together, 3 - 8 m m i n diameter;
peduncle 3- 1 4 mm long, puberulous to tomentellous; involucre with ea.
4 series of deltoid to ovate, acute, velutinous bracts; perianth 1 -2 mm
high, 4-lobed to 4-parted, densely puberulous; stamens 4, rarely 2
staminodial or wanting, two by two, more or less different, filaments
1 . 2-2.0 mm long, anthers 0.2-0.9 x 0.2-0. 5 mm, connective broad or
narrow, apiculate or not so.
Pistillate inflorescences solitary or sometimes a ccompanied by 1 -2
staminate ones, hemispherical to subglobose, 5-8 mm in diameter, subses­
sile or pedunculate; peduncle up to 5 mm long, puberulous to tomentel­
lous; involucre with 4-8 series of reniform to ovate to subulate, acute,
brownish velutinous bracts; flowers 5- 1 2; perianth 2.0-2. 5 mm high, 4-
parted , brownish velutinous; style lateral, 0. 7- 1 .7 mm long, hairy at the
upper end, stigmas vittiform, 2.5-5 .0 mm long, often twisted . I nfructes­
cences subglobose to globose, 2 . 5-5 .0 cm in diameter; fruiting perianth
yellowish, fruit ellipsoid, 6-7 mm long, seed ea. 5 mm long.

D i s t r i b u t i o n : Northern South America and eastern Brazil.

In S urinam in forests of the interior and the savanna belt; apparently

rather common.

BVV 1 352, 1 393, 1 422, 1 603, 1 6 1 4, 1 662, 3833, 4063, 4086, 4204, 4444, 5079,
555 1 : van Donselaar 1 920, 1 958a, 2077, 2346; H ostmann & Kappler 1 280;
LBB 8 1 37 , 836 1 ; Lindeman 4076, 4756, 5 299, 5356, 5357, 5469, 67 1 2; Oldenburger.
Norde & Schulz 1 228; Schulz 7 1 78; S tahel VVoodherb. 96, s.n.

Vernacular names: manletterhout, letterhout (Sur. Dutch),

222 MORACEAE

kaptinhoedoe (Sur.), sokone (Arow.), oempatapoe (Car.), poerienga

( Saram).

This widespread species is very variable. The specimens from the Guia­
nas agree most closely with those from Pani and the coastal region of
Brazil. Juvenile specimens differ from adult ones in having a sparser indu­
mentum.

2. Helicostylis pedunculata Benoist, Bull. Mus. H ist. Nat. Paris 25 : 298 .

1 9 1 9.

L e c t o t y p e : Benoist 667 9, French Guiana (P).

Dioecious or monoecious trees up to 25 m tall, with pale yellow latex.

Leafy twigs 2-7 mm thick, pale yellow tomentose to sublanate. Leaves
elliptic to lanceolate, 1 0-28 cm long, 5- 1 2 cm broad , sometimes broadest
above the middle, slightly inequilateral, coriaceous to chartaceous,
acuminate to mucronate, obtuse at the base; margin often denticulate to
dentate, mostly towards the apex; above tomentose on the costa, other­
wise puberulous, glabrescent to scabridulous, beneath tomentose; veins
almost plane above, p rominent beneath, 1 0- 1 7 pairs of secondary veins,
most tertiary veins parallel; petioles 6- 1 8 mm long; stipules 5-9 mm
long, subsericeous to tomentose.
Staminate inflorescences up to 6 together, 8-9 mm in d iameter; pedun­
cle 5- 1 5 mm long, tomentellous; involucre with 5-6 series of ovate, acute,
tomentellous bracts; perianth 1 - 1 .3 mm high, 4-lobed, tomentellous to
puberulous at the upper end ; stamens 4, filaments ea. 1 - 1 . 5 mm long,
anthers 0.6-0. 7 x ea. 0.2 mm.
Pistillate inflorescences solitary or usually accompanied by 1 -2 stami­
nate ones, discoid , 8- 1 1 mm in diameter; peduncle 6-25 mm long, tomen­
tellous, often bracteate; flowers ea. 1 5-30; perianth 1 . 5-2. 0 mm high, 4(-6)­
parted , tomentellous; ovary hairy at the tip, style subterminal, 0.2-2 . 1
mm long, hairy, stigmas vittiform, 2-3 mm long, not or somewhat
 MORACEAE 223

Fig. 5 Helicostylis; a., H. tomentosa, pistillate inflorescences (BW 4086);

b . , H. pedunculata, pistillate inflorescences and complemental staminate inflorescen­
ces ( BA FOG 7084) .

twisted. lnfructescences convexly discoid to hemisperical, 2-3 cm in

diameter; fruiting perianth pale yellow tomentellous; fruit ellipsoid, 7-8
mm long, seed ea. 6 mm long.

D i s t r i b u t i o n : From Surinam to Para (Brazil).

224 MORACEAE

In S urinam in forests of the interior and the savanna belt; apparently

confined to the eastern part of the country.

van Donselaar 1 727, 2003, 2992, 3 2 1 6; Lanj ouw & Lindeman 42 1 ; Lindeman
3544; Stahel s.n.

V e r n a c u l a r n a m e : Letterhout (Sur. Dutch).

l O . Naucleopsis M iquel, Fl. Bras. 4( 1 ): 1 20. 1 85 3 ; Berg, Flora

Neotropica Monogr. 7: l 04. 1 972.

T y p e s p e c i e s : Naucleopsis macrophylla M iquel

Ogcodeia Bureau in De Candolle, Prod. 1 7: 286. 1 873.

Oncodeia Bentham in Bentham & H ooker, Gen. PI. 3( 1 ): 373. 1 880.
L e c t o t y p e s p e c i e s : Naucleopsis glabra Spruce ex Baillon.
Acanthospaera Warburg, Verh. Bot. Ver. Brandenburg 48: 1 50, t.l. 1 907.
T y p e s s p e c ie s : Acanthospaera ulei Warburg (= Naucleopsis ulei (War­
burg) Ducke)).
Tramoia Schwacke & Taubert ex Giaziou, Bull. Soc. Bot. Fr. 59(Mem.3G):
674. 1 9 1 3, nom. nud.
Palmolmedia Ducke, Arq. Serv. Flor. Rio de Janeiro 1 : 20. 1 939.
T y p e s p e c i e s : Palmolmedia stipularis (Ducke) Ducke (= Naucleopsis
stipularis Ducke).

Dioecious trees. Leaves distichous, pinnatinervate; stipules free, fully

amplexicaul. Inflorescences on short shoots in the axils of the leaves,
unisexual, pedunculate or sessile, involucrate. Staminate inflorescences:
inner involucral bracts more or less petaloid and covering the flowers
before anthesis; flowers many to several; perianth with 0-8, usually 4, free
segments; stamens 4- 1 , straight in the bud. Pistillate inflorescences
with several to many flowers; perianth with 3- 1 0, mostly 4-6, free or
connate cushion-shaped to subulate tepals, more or less similar to the
pseudobracts in the periphery and among the flowers; ovary immersed
in the receptacle, stigmas linguiform to filiform. In fruit, tepals and
p seudobracts enlarged and hardened; seed big, without endosperm, testa
vascularized, cotyledons thick, equal and free, radicle short and apical.
 MORACEAE 225

D i s t r i b u t i o n : 1 8 species in tropical South and Central A merica.

The pistillate inflorescences and infructescences are very characteristic

for Naucleopsis. In most species the receptacle is covered with more
.
or less similarly shaped processes, called pseudobracts. Those
surrounding the styles can be regarded as tepals, those among the
flowers, i.e. not around the styles, are probably derived from the
perianths; the processes in the periphery of the receptacle are
inner involucra! bracts.

I a. Twigs and stipules usually distinctly hairy; petioles narrowly canaliculate;

secondary veins straight and arched 2-4 mm from the margin; stigmas ea.
3 mm long . . . ..
. . . . . . . . . . . . . . . . . . . . . . . . . . . 2. N . guianensis
. . . . . . . . .

b. Twigs and stipules (almost) glabrous; petioles broadly canaliculate, at

least at the upper end; secondary veins usually curved, arched 1 -2 mm from
margin; stigmas 7- 1 3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . I. N. glabra
.

l . Naucleopsis glabra Spruce ex Baillon, H ist. PI. 6: 206. 1 877.

Naucleopsis glabra Spruce ex Pittier, Contr. U. S . Nat. Herb. 1 3 :
440. 1 9 1 2, nom. nud .
Ogcodeia glabra (Spruce ex Pittier) M ildbraed, Notizbl. Bot.
Gart. Berlin 1 1 : 4 1 8. 1 932.

T y p e : Spruce 2793 C( , B razil, Amazonas ( P).

Duguetia? glabra Britton, Bull. Torrey Club 1 6· 1 4. 1 889.

Ogcodeia sandwithiana Mildbread, Notizbl. Bot. Gart. Berlin 1 1 : 422. 1 927.
T y p e : Rushy 1 378 C(, Bolivia, Beni (NY).
Ogcodeia tessmannii Mildbread, Notizbl. Bot. Gart. Berlin 10: 1 89. 1 927.
T y p e : Tessmann 4 1 09 o, Peru, Loreto ( B).
Ogcodeia tamamuri Macbride, Publ. Field Mus. Bot. 1 1 : 64, 1 9 3 1 .
T y p e : Williams 992 o, Peru, Loreto (F).
Ogcodeia acreana M ildbread, Notizbl. Bot. Gart. Berlin 1 1 : 42 1 . 1 932.
L e c t o t y p e : Ule 9320 o·, collected in J une 1 9 1 1 , Brazil, Acre Territory (B).
Ogcodeia pallescens Ducke, Arq. Serv. Flor. Rio de Janeiro 1: 1 8. 1 939.
T y p e : Kuhlmann 485 (HJBR 1 9807) c; , Brazil, Rondona (RB).
226 MORACEAE

Trees up to 20 m tall. Leafy twigs 1 -4 mm thick, glabrous or nearly so.

Leaves lanceolate to oblong, 7-3 1 cm long, 2 . 5- 1 0 cm broad, slightly in­
equilateral, often broadest above the middle, subcoriaceous to coria­
ceous, acuminate, attenuate to acute (to obtuse) at the base; glabrous or
nearly so; veins more or less prominent to slightly impressed above, 1 2-23
pairs of secondary veins, mostly curved and arched 1 -2( -3) mm from the
margin, without parallel tertiary veins; petioles 5-20 mm long, broad ly
canaliculate at least at the upper end ; stipules 8-20 mm long, glabrous or
sparsely appressed puberulous.
Staminate inflorescences up to 4 together, 5- 1 2 mm in diameter; pedun­
cle 1 . 5-7 mm long; involucre with 9-25 reniform to deltoid to oblong to
ovate, minutely puberulous bracts in 4-6 series; perianth 1 .0-2. 7 mm
high , with 3-7 free tepals; stamens 2-4, filaments 1 .0- 1 . 7 mm long, anthers
0 . 5- 1 .2 x 0. 3-0.6 mm.
Pistillate inflorescences solitary, 7- 1 0 mm in diameter, sessile or sub­
sessile; i nvolucre with ea. 1 5-25 renifo rm to deltoid to ovate to lanceolate,
obtuse to acute, minutely puberulous bracts in 4-8 series; 1 0 to ea. 30
flowers; perianth with 4-7 free or basally connate, 1 -4 mm long, obtuse,
puberulous tepals; style 2-4 mm long, stigmas 7- 1 3 mm long; pseudo­
bracts peripheral and among the flowers, similar to the tepals, subulate
to spine-like. Infructescences hemispherical or subglobose, 1 . 5-4,5 cm in
diameter; tepals and pseudobracts up to 8 mm long, more or less spine­
like and narrowed at the base.

D i s t r i b u t i o n : Northern South America.

In Surinam collected in the Brokopondo District and the Sipaliwini

savanna.
van Donselaar 3342; Oldenburger, Norde & Schulz 1 1 45.

The species is variable in the indumentum and characters of the pistillate

inflorescences . Most collections have come from the Upper Amazon Ba­
sin; a: few are known from northern Colombia, two from Surinam, and
one from French Guiana.
 MORACEAE 227

]�
a

Fig. 6 Naucleopsis, infructescences; a . , N. guianensis (BW 6688); b . , N. glabra

( Krukoff 1 300).

2. Naucleopsis guianensis ( M ildbraed) C.C. Berg, Acta Bot. Neerl. 1 8 : 465 .

1 969.
Ogcodeia guianensis M ildbraed, Notizbl. Bot. Gart. Berlin 1 1 :
422. 1 932.

T y p e : Sandwith 698 9, Guyana (B).

Trees up to 20 m tall, with yellowish to white latex. Leafy twigs 1 -3 mm

thick, sparsely whitish to brownish puberulous to more or less densely pu­
bescent to subtomentose. Leaves oblong to lanceolate, 5- 1 7 cm long, 1 . 5-
5 . 5 cm broad ; more or less inequilateral, often broadest above the mid­
dle, abruptly and bluntly acuminate, obtuse to acute at the base; gla­
brous; veins more or less prominent, 1 0- 1 9 pairs of secondary veins,
mostly straight and arched 2-4 mm from the margin, without parallel ter­
tiary veins; petioles 3- 1 0 mm long, narrowly canaliculate; stipules 7- 1 7
(-25) mm long, yellow to greyish appressed puberulous to pubescent.
Staminate inflorescences up to 4 together, 6-8 mm in d iameter; pedun­
cle 5- 1 2 mm long; involucre with 7- 1 0 subrotundate to deltoid to oblong
to ovate, minutely and sparsely puberulous, bracts in 3-4 series; perianth
1 . 5- 1 . 8 mm high , with 3-7 free · segments; stamens 2-4, filaments 0 . 8- 1 .2
mm long, anthers 0.6-0.9 x 0.4-0.6 mm.
228 MORACEAE

Pistillate inflorescences solitary, 7- 1 5 mm in diameter, sessile or subses­

sile; involucre with 1 0-25 subrotundate to deltoid to ovate, obtuse to
acute, minutely puberulous bracts in 3-5 series; flowers 5-20; perianth
with 3 -6 free, cushion-shaped to pyramidate, puberulous tepals; style 1 -
1 . 5 m m long; stigmas linguiform, 2.5-3 . 3 m m long; pseudobracts mainly
peripheral, similar to the tepals. I nfructescences 2-4 cm in diameter;
tepals and pseudobracts cushion-shaped, angulate and apiculate , up to
3 mm high; fruit ellipsoid , 8-9 mm long.

D i s t r i b u t i o n : From Guyana through Surinam to Amapa ( HraZil) .

In Surinam in forests of the interior and the savanna belt .

BW 6299, 6688; van Donselaar 336 1 , 379 1 ; Lanjouw & Lindeman 2544; Lindeman
3790, 4700, 6750, 6774; Schulz 798 1 .

V e r n a c u l a r n a m e : letterhout (Sur. Dutch).

This is the only species of the Olmedieae which is confined to the Guiana
Shield region. The pistillate inflorescences of the Surinam specimens dif­
fer from those of Guyana in the shorter and broader tepals and pseudo­
bracts and in the sho rter stigmas.

1 1 . Ficus Tournefort ex Linnaeus, Genera Plantarum ( ed. 5) 482. 1 754.

Oluntos Rafinesque, Sylva Telluriana 58. 1 838.

Urostigma Gasparrini, Nova Genera, Fici 7. 1 844.
Pharmacosycea Miquel, London J our. Bot. 7: 64. 1 848.

A genus (at least so far as the American species are concerned) of soft­
wooded , generally smooth-barked , trees and shrubs with milky juice.
Leaves alternate, in spiral phyllotaxy, entire. Stipules long or short, gen­
erally quickly deciduous, enfolding the buds. Flowers unisexual, haplo­
chlamydeous, borne inside a much enlarged and deeply invaginated
torus (the receptacle, or fig), the apical pore ( ostiole or" orifice) of
 MORACEAE 229

which is closed by a series of horizontal, inte rlocking bracts. The

female flowers are of two kinds: functional females, generally sessile
and maturing (after fertilization) into viable achenes, and ste rile
females (gall flowers), generally stalked and funct ioning as incubators
for the larvae of a wasp which polli nates the fertile flowers. Male,
female, and gall flowers are completely i ntermixed in the receptacles
of the American species. The female flowers receptive long before the
male flowers mature in the same receptacle.

I a. Figs borne in pairs in the axils of the foliage leaves; stamens in the male
flowers I , a red spot at the base of the style on the ovary of female and gall
flowers; 2-celled glandular hairs frequently present on the lower surface of
the lamina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . subg. Urostigma
b. Figs borne singly in the axils of the foliage leaves; stamens in the male flowers
2, ovary pale, unspotted; massive multicellular hairs present on the lower
surface of the lamina . . . . . . . . . . . . . . . . . . . . . . . . . . . . subg. Pharmacosycea

l l A. Ficus subg. Pharmacosycea ( Miquel) Miquel, Ann. Mus. Lugd.

� Bat. 3 : 299. 1 867.

Generally large, free-growing trees of the forest, forest margins, or

second growth. Figs generally large and thick-walled.

I a. Leaves at least weakly scabrid below, generally scabrid on both surfaces.

Epidermis of the petioles frequently exfoliating in dried material.
Lateral veins 5- 1 6 pairs. Leaf colour is characteristically very
dark green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I . F. maxima
b. Leaves smooth on both surfaces. Epidermis of the petioles not exfoliating
in dried material, but sometimes wrinkled . Lateral veins 1 0-25 pairs. Leaf
colour a bright green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. F. insipida

l . Ficus maxima P. M iller, Gard . D iet. (ed. 8 . ) 1 768.

L e c t o t y p e : S loane, V oyage to . . . . . . . Jamaica 2: 1 40. t.

223. 1 725.
230 MORACEAE

Ficus radula Humboldt & Bonpland ex Willdenow. Sp. PI. 4(2): 1 1 44.
1 806.
Pharmacosycea radula (Willdenow) Miquel, Versl . Kon. Akad. Wetensch .
Amsterdam 1 3 : 4 1 4. 1 862.
Ty p e : Humboldt & Bonpland s.n., Venezuela, Terr. Fed . Amazonas ( B).
Pharmacosycea grandaeva Martius ex Miquel, London Jour. Bot. 7: 70. 1 848.
Ty p e : Martius 7 1 , Brazil, Amazonas (U ).
Pharmacosycea guyanensis Miquel, London Jour. Bot. 7: 67. 1 848.
Ficus parkeri M iquel, Ann. M us. Lugd. -Bat. 3: 300. 1 867.
T y p e : Parker s.n., Guyana (K).
Pharmacosycea glaucescens Liebmann, Kgl. Danske Vidensk. Selsk. Skr.
V. 9: 332. 1 85 1 .
Ficus glaucescens (Liebmann) Miquel, Ann. Mus. Lugd . -Bat. 3 : 300.
1 867.
Ty p e : Liebmann 1 43 1 4, Mexico (C).
Pharmacosycea hernandezii Liebmann, Kgl. Danske Vidensk. Selsk. Skr.
V. 2: 332. 1 85 1 .
Ficus hernandezii ( Liebmann) Miquel, Ann. Mus. Lugd. -Bat. 3 : 300. 1 867.
T y p e : Liebmann 1 43 1 6, Mexico (C).
Pharmacosycea rigida M iquel in Seeman, Bot. Voy. Herald 1 95 . 1 854.
Ficus coybana M iquel, Ann. M us. Lugd . -Bat. 3: 300. 1 867.
T y p e : Seemann 638, Panama (BM).
Pharmacosycea mexicana M iquel, Versl. Kon. Akad. Wetensch. Amsterdam
1 3 . 4 1 5. 1 862.
Ficus mexicana ( Miquel) Miquel, Ann. Mus. Lugd. -Bat. 3 : 299-300. 1 867.
T y p e : ? Schiede 43, M exico ( U ) .
Pharmacosycea pseudo-radula Miquel, Versl. Kon. Akad. Wetensch.
Amsterdam 1 3 : 4 1 4. 1 862.
Ficus pseudo-radula ( Miquel) Miquel, Ann. M us. Lugd. -Bat. 3 : 299. 1 867.
T y p e : Schiede s.n., M exico ( U ) .
Ficus suffocans Banks ex Grisebach, F J . B rit. W . l n d . 1 50. 1 864.
S y n t y p e s : March s.n. (or 582), Jamaica ( K , mixed gathering);
Prior s.n. (or 297), Jamaica (K).
Ficus guadalajarana S. Watson, Proc. Am. Acad. 26: 1 5 1 . 1 89 1 .
Type: Pringle 2947A, Mexico (GH).
Ficus finlayana Warburg, Symbol. Antill. 3: 487. 1 903, pro parte.
T y p e : Finlay s.n., Trinidad (B, a mixed sheet: a single leaf = F. insipida,
a single fig = F. maxima).
Ficus picardae Warburg, Symbol. Antill. 3: 484. 1 903.
Ty p e : Picarda 983, Haiti (B).
Ficus rubricosta Warburg, Symbol. Antill. 3 : 486. 1 903.
T y p e : Eggers, 2625, Sto. Domingo (B).
Ficus subscabrida Warburg, Symbol. Antill. 3 : 485. 1 903 .
. Ty p e : Wright 543, Cuba (isotype P).
Ficus plumierii Urban, Repert. Sp. Nov. 1 5: 1 58 . 1 9 1 8.
L e c t o t y p e : Plumier, PI. Am. 1 23, t. 1 3 1 , f. 3. 1 757.
 MORACEAE 23 1

Ficus bopiana Rushy, Mem. N.Y. Bot. Gard . 7: 230. 1 927.

T y p e : Rushy 59 1 , Bolivia (isotype K).
Ficus ulei Rossberg, Repert. Sp. Nov. 42: 60. 1 937.
T y p e : Ule 57 1 6, Brazil, Amazonas ( B).
Ficus vicencionis Dugand, Caldasia 2: 385. 1 944.
T y p e : Jaramillo-Mejia 2 1 9, Colombia, Meta (isotype I AN).

S mall to large tree, 5-30 m tall, Twigs 3-5 mm in diameter, glabrous or pu­
bescent, the epidermis generally exfoliating in dfied material. Stipules to
25 mm long, narrowly delto id , glabrous, puberulent, or pubescent, at the
base. Lamina 2. 5-9 cm wide, 6-23 cm long, elliptic, broadly elliptic, lan­
ceolate, oblanceolate or obovate; apex blunt, acute, acuminate or long
acuminate; base cuneate; lateral veins 5- 1 6, departing from the midrib at
an angle from oo -30° ; basal veins 1 -2, departing from the mid rib at an
angle of 40° -60° ; intercostals slightly raised . Petiole 5-40 mm long, 1 I 5-
1 1 9 the length of the lamina, the epidermis generally exfoliating in dried
material.
Figs 1 0-20 mm in d iameter, globose, glabrous or puberulent, some­
times with a pedicel 2-7 mm long, borne among the leaves; colour green
or yellow, sometimes mottled darker; peduncle 2-25 mm long, glabrous
or pubescent; basal bracts 1 -2 mm long, deltoid, glabrous or pubescent;
orifice 1 -2 mm in d iameter, flat, or the bracts slightly outflexed . Female
flowers sessile or stalked, 1 . 5-3 mm long, tepals 4-5 , linear-oblanceolate,
linear-lanceolate, or linear-oblong, glabrous or ciliolate. Male flowers
long or short stalked , 0. 5-3 mm long, tepals 3-4, oblong, hooded, gla­
brous. S cales p resent among the flowers.
Leaf structure: cystoliths p resent on both surfaces, generally relatively
fewer and smaller on the upper surface; sclereids absent; lower epidermal
cells plane or somewhat sinuate and plicate; stomates superficial , or
sunken in sinuate-mouthed p its; d ruse cells abundant in the lower epider­
mis; tabular crystal cells absent; pubescence of massive, multi-cellular
hairs; to 0.2 mm long, acicular, unicellular hairs sometimes present. Scler­
eids absent from the fig wall.

D i s t r i b u t i o n : M exico, through Central America and the West Indies,

throughout South America to Paraguay.
232 MORACEAE

In Surinam from the young coastal plain to the interior; apparently

common in swamp forests and along streams.

BW 2028, 2405, 2434, 3760, 6 1 99; van Donselaar 1 1 54, 1 1 73; Gonggrijp 9; Lanj ouw &
Lindeman 1 468; Lindeman 5 1 37, 5407, 5497, 5594, 5640, 5693, 570 1 , 6347, 6635;
Wullschliigel 1 084 (BR), s.n. (BR) . .

2. Ficus insipida Willdenow, Sp. PI. 4(2): 1 1 43 . 1 806.

T y p e : Bredemeyer s . n . , Venezuela, D . F. ( B) .

Ficus glabrata H . B. K . , Nov. Gen. e t S p . 2 : 47. (or 38). 1 8 1 7.

T y p e : Humboldt & Bonpland s.n. , Colombia ( P).
Ficus adhatodaefolie Schott in Sprengel, Syst. Veg. 4(Cur. Post. App.):
409. 1 827.
Pharmacosycea adhatodaefolia (Schott in Sprengel) M iquel, London Jour.
Bot. 7: 70. 1 848.
T y p e : Schott s.n., Brazil ( B).
Ficus anthelminthica Martius in Spix & M artius, Reise in Brasilien 3: 1 1 5 8.
1 83 1 .
Pharmacosycea anthelminthica ( M artius) M iquel, London J our. Bot. 7:
66. 1 848.
T y p e : Martius s.n., Brazil, M inas Gerais (M).
Pharmacosycea vermifuga M iquel, London J our. Bot. 7: 70. 1 848.
Ficus vermifuga ( M iquel) Miquel, Ann. M us. Lugd. -Bat. 3: 300. 1 867.
T y p e : Martius s.n. , Brazil (U).
Pharmacosycea angustifolia Liebmann, Kgl. Danske Vidensk. Selsk.
Skr. V. 2: 333. 1 85 1 .
Ficus segoviae Miquel, Ann. M us. Lugd. -Bat. 3 : 300. 1 867.
T y p e : Oersted 1 4334, Nicaragua (C).
Ficus radulina S . Watson, Proc. Am. Acad. 26: 1 5 1 . 1 89 1 .
T y p e : Palmer "L", Mexico (GH).
Ficus krugiana Warburg, Symbol. Antill. 3 : 487. 1 903.
S y n t y p e s : Isert 87, Martinique (C) = F. insipida; Eggers 6670, St. Vincent
(P) = F. maxima.
Ficus crassiuscula Warburg ex Standley, Contrib. U. S. Nat. Herb. 20:
12. 1 9 1 7 .
T y p e : Pittier 1 6 1 50, Costa Rica ( B).
Ficus werckleana Rossberg, Repert. Sp. Nov. 42: 60. 1 937.
T y p e : Werckle 1 7436, Costa Rica ( B).
Ficus boyacensis Dugand, Caldasia 1 (4): 3 1 . 1 942.
T y p e : Daniel 2275, Colombia (isotype US).
Ficus crassa Klotzsch & Karsten ex Dugand, Caldasia 1 (4): 35. 1 942.
T y p e : Karsten s.n., Colombia ( B).
 MORACEAE 233

Large or small tree, 8-40 m tall. Twigs 2-6 mm in d iameter, generally gla­
brous, occasionally pubescent. S tipules 30- 1 25 mm long, narrowly del­
toid, generally glabrous, occasionally pubescent. Lamina I . 75- 1 1 cm
wide, 5-25 cm long, lanceolate to broadly elliptic, sometimes slightly in­
equalateral; apex blunt or acute, to acuminate; base cuneate, round cu­
neate, rounded, or emarginate; lateral veins 1 0-30, departing from the
midrib at an angle from 30° -60° ; intercostals slightly prominent. Petiole
1 0-65 mm long, 1 / 3- 1 / 8 the length of the lamina, epidermis not exfo­
liating, though frequently wrinkled in d ried m � terial.
Figs 1 5-25 mm (to 30 mm fide Little ex sched .) in diameter, globose,
with or without a pedicel, 1 -6 mm long, glabrous or pubescent, borne
among the leaves; colour green or yellowish green; peduncle 3-22 mm
long, thin or stout; basal bracts 2-3( -5) mm long, deltoid or semi-circular;
orifice flat, or somewhat crateriform or mammillate, 2-4 mm in diameter,
1 -2 mm high. Female flowers about 2 mm long, with 4-6 linear, lanceo­
late, linear-oblong, or narrowly linear tepals, glabrous or ciliolate, some­
times somewhat hooded . Male flowers 1 . 5-2 mm long, with 4-6 oblong,
hooded, glabrous or ciliolate tepals. Scales present between the flowers.
Leaf structure: cystoliths p resents on both upper and lower surfaces;
sclereids absent; lower epidermal cells plane; stomates generahy sunken
in small-mouthed pits, in B rasilian material sometimes only slightly
sunken; druse cells present in the lower epidermis; tabular crystal cells ab­
sent; pubescence of massive, multicellular hairs; simple or multiseptate,
acicular hairs sometimes p resent. Sclereids generally present in the fig
wall.

D i s t r i b u t i o n : Mexico to Paraguay in moist evergreen forest, from

0-2000 m altitude.

I n Surinam known from two localities near the Suriname River.

BW 2 1 5 1 ; Tresling 228.

1 1 B. Ficus subg. U rostigma (Gasparrini) M iquel, Ann. Mus . Lugd .

-Bat. 3: 260. 1 867.
234 MORACEAE

Small or large trees, or shrubs. These are the banyans, or strangler figs.
So far as is known the species of this sub genus normally begin life as epi­
phytes (or epiliths) and frequently finish life as (apparently) normal
trees, in the meantime having built up a false trunk through the coales­
cence of their aerial roots, and frequently having strangled their original
host.

I a. Figs borne both on wood of the current season (among the leaves) and on
specialized short shoots on wood of previous seasons (behind the leaves);
figs less than 10 mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
b. Figs borne only on wood of the current season (among the leaves); fig size
variable . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
2 a. Leaves generally more than 1 5 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
b. Leaves generally less than 1 5 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3 a. Leaves narrowly oblong or oblanceolate, glabrous; sclereids present in the
leaves; twigs 7- 1 0 mm in diameter . . . . . . . . . . . . . . . . . . . . . 9. F. longifolia .

b. Leaves elliptic-oblong to obovate, densely and minutely dark brown lanate

below; sclereids absent from the leaves; twigs 3-8 mm in diameter . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. F. albert-smithii
4 a. Leaves usually obovate or elhpt1c, 2-� . ) cm w1ae; stomates deeply sunken in
small-mouthed pits; tabular crystal cells usually absent . . . 4. F. clusiaefolia
b. Leaves usually oblanceolate or elliptic, 1 -4 cm wide; stomates only slightly
sunken; tabular crystal cells usually present on the lower surface of the
veins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
. . .

5 a. Leaves glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
b. Leaves densely and minutely dark brown lanate below . I . F. albert-smithii
6 a. Lateral veins departing from the mid rib at an angle from oo - 1 0° . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6. F. erratica
b . Lateral veins departing from the midrib at an angle from 1 0° -20°
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 0. F. mathewsii
7 a. Orifice of the fig level with the surface of the fig wall, or the bracts raised
into a mammillate structure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 2
b. Orifice of the fig surrounded by a raised rim of receptacular tissue, o r the
apex of the receptacle drawn up into a rim or crateriform structure, or the
orifice sunken below the surface of the fig wall . . . . . . . . . . . . . . . . . . . . . . 8 .

8 a. The orifice surrounded by a thin, narrow, more or less erect rim of recept-
acular tissue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 5. F. trigona
b. The orifice otherwise . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9 a. Orifice of the fig slightly sunken into the apex of the receptacle, or surround­
ed by a slightly raised rim or ring of receptacular tissue . . . . . . . . . . . . . . . 1 0
b . Orifice of the fig sunken into a prominent crateriform structure . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 3 . F. pertusa
10 a. Figs 1 0-25 mm in diameter; leaves relatively thick, generally more than 5 cm
wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 1
. . .
 MORACEAE 235

b. Figs less than 1 0 mm in diameter; leaves thin, generally less than 5 cm

wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 3. F . pertusa
11 a. Leaves generally more or less spreading pubescent, at least along the veins
below . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. F . gomelleira
b. Leaves totally glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 6. F. trigonata
12 a. Orifice of the fig level with the surface of the fig wall . . . . . . . . . . . . . . . . 1 4
b . Orifice of the fig o r the bracts raised into a mammillate structure . . . . . . 1 3
13 a . Leaves oblong, oblong-elliptic, oblong-oblanceolate o r oblanceolate, acumi­
nate; lateral veins 9- 1 8, departing from the midrib at an angle from
1 0° -30° . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 2. F . paraensis
b. Leaves elliptic, or somewhat obovate, rounded, acute or weakly acuminate;
lateral veins 6- 1 1 , departing from the mid rib at an angle from 20° -
40° . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 6. F. trigonata
14 a. Figs 1 5 mm or more in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 5
b . Figs less than 1 5 mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 7
15 a . Petiole 2 5 m m long o r less; leaves oblong o r oval . . . . . . . . 1 6. F . trigonata
b. Petiole more than 25 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 6
16 a . Leaves elliptic o r elliptic-oblong . . . . . . . . . . . . . . . . . . . . . . . 2 . F. broadwayi
b. Leaves ovate or ovate-rotund . . . . . . . . . . . . . . . . . . . . 1 1 . F . nymphaeaefolia
17 a. Figs less than 6 mm in diameter; twigs about 2 mm in diameter, minutely
pubescent or glabrate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 4. F. prinoides
b. Figs more than 6 mm in d iameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 8
18 a. Figs 8-9 mm in diameter; twigs and stipules densely appressed pubes-
cent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8. F . Ianjouwii
.

b. Figs 6-20 mm in d iameter; twigs and stipules glabrous or glabrescent . . . 1 9

19 a . Sclereid s present i n the leaves . . . . . . . . . . . . . . . . . . . . . . . . . 2. F . broadwayi
b. Sclereids absent from the leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20 a. Peduncle of fig 0-2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1
b. Peduncle of fig 2 m m o r more long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
21 a. Leaves oblong or oval; druse cells occasionally present; stomates super­
ficial or sunken . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 6. F. trigonata
b. Leaves lanceolate, ovate, oblong-lanceolate, or oblanceolate; druse
cells present; stomates superficial . . . . . . . . . . . . . . . . . . . . . . . 5. F. ernestiana
22 a. Leaves oblong or oval; tabular crystal cells absent from the leaves . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 6. F. trigonata
b. Leaves lanceolate, ovate, elliptic, elliptic-lanceolate, elliptic-ovate, oblong or
obovate; tabular crystal cells present on the lower surface of the
veins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. F . citrifolia

1 . Ficus albert-smithii Standley, Lloydia 2: 1 74. 1 939.

T y p e : A . C. Smith 365 1 , Guyana ( F).

Small tree, 2-4 m tall. Twigs 3-8 mm in diameter, densely and minutely
236 MORACEAE

Fig. 7 Ficus albert-smithii; a., twig ( 1 / 2 x); b., lower epidermis of leaf (c. 300 x ) ,
showing cellular detail, sclereids indicated by dashed lines; c., upper epidermis of
leaf (c. 300 x ) , showing cellular detail (Steyermark 9040 1 ) .
 MORACEAE 237

dark brown lanate, much shrunken and longitudinally ridged in dried

material. Stipules 7- 1 5 mm long, acuminate-deltoid. Stipules, petioles,
and lower surface of the lamina of the leaves densely and minutely dark
brown lanata. Lamina 8 . 5- 1 9 cm long, elliptic-oblong to obovate ; apex
bluntly acute to acuminate; base rounded or slightly emarginate; lateral
vei ns 5-20, departing from the mid rib at an angle of 30° -40° ; basal veins
I pair, departing from the midrib at an angle of 50° -60° ; intercostals
slightly raised . Petiole 1 0-20 mm long, I / 7- I I 9 the length of the lamina.
Figs 5-7 mm in diameter, depressed globose, pink, borne in pairs
among the leaves on growth of the current season and behind the leaves
on abbreviated short-shoots; peduncle l -2 mm long; basal bracts semi­
circular up to about 3x5 mm; orifice flat, about 2 mm diameter. Female
flowers sessile and gall flowers stalked , 1 -2 mm long, with 3 hooded
tepals. Scales present between the flowers.
Leaf structure: cystoliths few or none on the upper surface, few below;
sclereids absent; lower epidermal cells plane; stomates sunken in small­
mouthed crypts; druse cells absent; tabular crystal cells absent; pubes­
cence of clusters of multiseptate hairs .

D i s t r i b u t i o n : Apparently confined to the mountainous savanna

region of Guyana and Surinam, westward to Bolivar and Delta Amacuro
in Venezuela .

In S urinam a single collection from the Brownsberg.

BW 6497.

2. Ficus broadwayi U rban, Repert Sp. Nov. 1 5 : 1 1 0. I 9 I 7.

T y p e : B roadway 4382, Tobago (B).

Ficus mendelsonii Britton, Bull. Torrey Club 48: 330. 1 92 1 .

T y p e : Britton, Hazen & Mendelson 879, Trinidad (isotype G H).
U rostigma leucostictum Miquel, London Jour. Bot. 6: 535. 1 847.
238 MORACEAE

c
b

Fig. 8 Ficus broadwayi; a., twig (c. 14 x ) ; b., lower epidermis of leaf (c. 300 x ) ,
showing sclereids as dashed lines, druses of calcium oxalate in epidermal cells,
and tabular crystal cells on surface of the veins; c., upper epidermis of leaf (c. 300 x ) ,
cystoliths indicated by dashed circle, and druses of calcium oxalate in epidermal
cells (Steyermark 6 1 496).
 MORACEAE 239

Ficus leucosticta ( M iquel) M iquel, Ann. M us. Lugd . -Bot. 3: 297 .

1 867; n o n Sprengel, Syst. Veg. 3 : 7 7 9 . 1 826.
T y p e : Parker s.n., Guyana ( K).
Ficus manicariarum Stand ley, Bull. Torrey Club 7 5 : 297. 1 948.
T y p e : Fanshawe 2434, G uyana (isotype K).
Ficus savannarum Standley, Bull. Torrey Club 7 5 : 298. 1 948.
T y p e : Maguire & Fanshawe 23292, G uyana (NY).

Tree to 20 m tall. Twigs 3-7 mm in diameter, glabrous or minutely puberu­

lent. Stipules 8 mm or more long, deltoid , glabrous or puberulent with
the margins ciliate. Lamina 5- 1 2 cm wide, 1 2-23 cm long, elliptic or ellip­
tic-oblong; apex rounded or acuminate; base rounded, rounded-emargi­
nate, or subcordate; lateral veins 7- 1 4 , departing from the midrib at an
angle from 1 0° -30° ; basal veins 1 -2, departing from the midrib at a simi­
lar angle, or at an angle from 30° -40° ; intercostals only slightly raised. Pe­
tiole 25-80 mm long, 1 I 2- 1 I 8 the length of the lamina.
Figs 1 0-20 mm in diameter, globose, minutely puberulent, borne
among the leaves; colour yellowish; peduncle obsolete or to 5 mm long;
basal bracts more or less rotund, 1 -2 mm long; orifice 2-3 mm in diame­
ter, more or less plane, within a more or less prominent ring of recepta­
cular tissue. Male and female flowers about 1 mm long, with 2-3 hooded
tepals. The male flowers short stalked . Bracts present between the flo­
wers.
Leaf structure: cystoliths present on both surfaces; sclereids present;
lower epidermal cells sinuate and plicate or nearly plane and only a little
striate; stomates superficial; d ru se cells absent; tabular crystal cells gener­
ally absent; dark brown cells abundant along the lower surfaces of the
veins and in lower mesophyll; pubescence of 2-cellular glandular hairs.

D i s t r i b u t i o n : Trinidad and Tobago, Venezuela, Colombia and Brazil.

In Surinam known from near the Lower Corantine River, the U pper
Lucie River and the Brownsberg.

BW 4759, 6556; I rwin, Prance, Soderstrom & H olmgren 55452.

240 M ORACEAE

b c

Fig. 9 Ficus citrifo/ia; a., twig (c. 14 x); b., upper epidermis of leaf (c. 300 x),
cystolith indicated by dashed circle, druses of calcium oxalate in epidermal
cells; c., lower epidermis of leaf (c. 300 x), sclereids indicated by dashed lines, druses
in epidermal cells, tabular crystals in superficial cells of the veins (Steyermark 95437).
 MORACEAE 24 1

3. Ficus citrifolia P. M iller, Gard . Diet. ( ed. 8). 1 768 .

Ficus catesbaei Steudel, Nomenclator (ed . 2) 1 : 636. 1 840, as
synonym.

T y p e : sheet so la be led, ex herb. M iller ( BM).

Ficus pedunculata Dryander in Aiton, H ort. Kew (ed. I) 3: 450. 1 789.

lJrostigma pedunculatum ( Dryander) M iquel, London Jour. Bot. 6: 540. 1 847.
L e c t o t y p e : Plukenet, Phytographia t. 1 78, f. 4. 1 692.
Ficus populnea Willdenow, Sp. PI. 4(2): 1 1 4 1 . 1 806.
U rostigma populneum (Willdenow) M iquel, London J our. Bot. 6: 537. 1 847.
T y p e : sheet so labeled (ex hort. Paris) in herb. Willdenow ( B).
Ficus laevigata Vahl, Enum. 2: 1 83. 1 805.
U rostigma laevigatum (Vahl) M iquel, London J our. Bot. 6: 539. 1 847.
T y p e : West s.n., St. Croix (C).
Ficus lentiginosa Vahl. Enum. 2: 1 83. 1 806.
U rostigma lentiginosum (Vahl) Liebmann, Kg!. Danske Vidensk. Selsk. S kr.
V. 2: 323. 1 85 1 .
T y p e : Ryan s.n., M ontserrat (C).
Ficus gigantea H. B. K., Nov. Gen. et Sp. 2: 48. (or 39) 1 8 1 7, non Ficus
gigantea N oronha, 1 790.
U rostigma giganteum ( H . B.K.)Miquel, London J our. Bot. 6: 530. 1 847.
Ty p e : H umboldt & Bonpland s.n. ( P , Herb. H umboldt MN 722).
Ficus rubrinervis Link, Enum. PI. H o rt. Berol. 2: 448. 1 822.
T y p e : Link s . n . , cult. H ort . Berol., sheet so labeled (B).
Ficus eximia Schott in Sprengel, Syst. Veg. 4( Cur. Post. App.): 4 1 0. 1 827.
U rostigma eximium (Schott in Sprengel) M iquel, London Jour. Bot. 6:
527. 1 847.
Ty p e : Schott s.n., cultivated from a plant collected in the vicinity of Rio
de Janeiro ( Brazil), sheet so labeled (B).
Ficus pyrifolia Desfontaines, Cat. H ort. Par. (ed . 3) 4 1 3. 1 829.
T y p e : sheet so labeled in herb. Desfontaines ( FI).
Ficus botryapioides Kunth & Bouche, I nd . Sem. H ort. Berol. 1 5. 1 846.
T y p e : K unth & Bouche s.n., sheet so labeled ( B) .
Ficus syringaefolia K unth & Bouche, I nd. S e m . H ort. Berol. 1 5. 1 846.
T y p e : K unth & Bouche s.n., sheet so labeled ( B).
U rostigma angustifolium M iquel, London Jour. Bot. 6: 539. 1 847.
Ficus angustifolia ( M iquel) M iquel, Ann. M us. Lugd . -Bat. 3: 298 . 1 867 , nee
Blume, Bij dr. 1 826; nee Roxburg, H ort. Beng. 1 8 14; Fl. lnd. 1 820.
Ty p e : Parker s.n., Guyana (K).
U rostigma amazonicum M iquel, London Jour. Bot. 6: 54 1 . 1 847.
Ficus amazonica ( M iquel) M iquel, Ann. M us. Lugd . -Bat. 3: 298. 1 867.
S y n t y p e s : Martius 88, 89, Brazil, Amazonas ( M ).
Ficus brevifolia Nuttall, Sylv. 2: 3 . 1 854.
242 MORACEAE

T y p e : Blodgett s.n., Key West ( B M ) .

Ficus surinamensis M iquel, Ann. M us. Lugd. -Bat. 3: 299 . 1 867.
T y p e : Kegel 1 86, Surinam ( U ) .
Ficus thomaea Miquel, Ann. M us. Lugd . -Bat. 3:299. 1 867.
T y p e : Ehrenberg 236, St. Thomas ( B).
Ficus populoides Warburg, Symbol. Anti!!. 3 : 479. 1 903.
S y n t y p e s : Wright 1 686, Cuba ( K , P); J aeger 1 89, Haiti (C, K , P).
Ficus brittonii Bo1dingh, Fl. Dutch West. Ind. Islands 2: 20. 1 9 1 4;
non Pharmacosycea brittonii, Rusby, Bull. Torrey Club 28: 309. 1 90 1 .
L e c t o t y p e : Boldingh 5342, Cura�ao ( U ) .
Ficus guarantica Chodat, Bull. Soc. Bot. Geneve 1 1 . 1 1 : 254. 1 920.
T y p e : Balansa 1 986, Paraguay (isotype K).
Ficus turbinata Pittier, Bol. Soc. Venez. Ci. Nat. 4: 6 1 . 1 937, non Willdenow.
Sp. PI. 4: 1 1 4 1 . 1 806. I
Ty p e : Pittier 1 2277, Venezuela, D. F. (isotypes N Y , US).
Ficus guanarensis Pittier, Bol. Soc. Venez. Ci. Nat. 4: 7 1 . 1 937.
T y p e : Pittier 1 2077, Venezuela, Portuguesa ( Y EN).
Ficus subandina Dugand, Caldasia I ( 4): 66. 1 942.
T y p e : Dugand 2954, Colombia, Cundinamarca (COL, not seen; paratype
Gutierrez 1 42, U S).

Tree to 16 m tall, or, in exposed places, a shrub 0. 3-2 m tall. Twigs 2-6
mm in diameter, glabrous. Stipules 5-30 mm long, narrowly deltoid, gla­
brous, sometimes somewhat glaucous. Lamina 1 . 5- 1 2 cm wide, 2 . 5-
1 9 cm long, lanceolate, ovate, elliptic, elliptic-lanceolate, elliptic-ovate ,
oblong or obovate; apex acute to acuminate; base rounded, rounded­
cuneate, truncate, freq uen tly emargina te or su bcordate; lateral veins 4- 1 4,
departing from the mid rid at an angle from I 0° -40° ; basal veins l -2, de­
parting from the midrib at a similar angle, or to 60° ; intercostals not or
scarcely prominent. Petiole 7-70 mm long, l / 2- l / 7( - l / 8) the length of the
lamina.
Figs 6- 1 2(- 1 5) mm in diameter, globose, glabrous, borne among the
leaves; colour reddish or yellowish when ripe; peduncle 2- 1 8 mm long,
glabrous or puberulent; basal bracts 2-3 mm long, broadly deltoid to se­
micircular, with a hyaline margin, glabrous or puberulent; orifice 2-3 mm
in diameter, flat or very slightly raised . Female flowers with 2-3 hooded
tepals, l - 1 . 5 mm long. Male flowers with 2 hooded tepals, sessile or
short-stalked , l - 1 . 5 mm long. Scales present between the flowers.
Leaf structure: cystoliths present on upper and lower surface; sclereids
 MORACEAE 243

absent; lower epidermis plane; stomates superficial; druse cells

absent; tabular crystal cells present on the lower surface of the veins; pu­
bescence absent or of 2-5-celled glandular hairs on the lower surface.
The mesophyll makes a distinctive, reticulate pattern immediately below
the epidermis.

D i s t r i b u t i o n : Widespread from Lake Okeechobee, Florida, U . S .

through the West Ind ies, i n Central America north t o Oaxaca ( M exico) ,
south to Paraguay.

In Surinam from the young coastal plain to th : interior.

BW 2085; I rwin, P rance, Soderstrom & H olmgren 57585; K egel 1 86; Lanj ouw 878,
895; Lanj ouw & Lindeman 1 075, 3 1 59; LBB 1 2490; Mennega 488; Pulle 254;
Schulz 7784; Wullschlagel 1 082 (BR).

4. Ficus clusiaefolia Schott in Sprengel, Syst. Veg. 4(Cur. Post . ; App. ) :

409. 1 827; n o n U ro stigma clusiaefolium (Schott in Sprengel)
M iquel, London Jour. Bot. 6: 543. 1 847; M iquel, Fl. B ras. 4( 1 ):
1 02. 1 8 5 3 .

T y p e : Schott s . n . , B razil (B).

Ficus guianensis Desvaux ex Hamilton, Prod. Fl. Ind. Occ. 62, 1 825.
Ty p e : Desvaux s.n., French Guiana (P).
Ficus anacardiifolia, Kunth & Bouche, Ind. Sem. H ort. Berol. 1 5. 1 846.
U rostigma anacardiifolium ( K unth & Bouche) M iquel, London Jour. Bot.
6: 587. 1 847 .
T y p e : K unth & Bouche s.n. (B).
Ficus splendens Kunth & Bouche, Ind . Sem. H ort. Berol. 1 4. 1 846.
U rostigma splendens ( Kunth & Bouche) Miquel, London Jour. Bot. 6: 587.
1 847.
Ty p e : Kunth & Bouche s.n. (B).
Ficus martinii Miquel, Ann. Mus. Lugd . -Bat. 3: 2 1 9. 1 867.
Ty p e : Martin s.n. (or 1 23), French Guyana (isotypes B, CGE, FI, L, P).
Ficus umbonigera Warburg, Symbol. Anti!!. 3 : 480. 1 903.
T y p e : Hart 2808, Trinidad (isotype T R IN).
244 MORACEAE

Ficus grenadensis Warburg, Symbol. Antill. 3 : 48 1 . 1 903.

Ty p e : Eggers 6 1 38, Gre nada ( B).
Ficus myriasycea Pittier, Bol. Soc. Venez. Ci. N at. 4: 75. 1 937.
T y p e : Pittier 1 1 924, Venezuela, D . F. ( Y E N , not seen; paratypes Pittier
1 255 1 , B; Pittier 8706, NY, US).

Tree to 8 m or more tall. Twigs 2-3(-6) mm in diameter, glabrous

or minutely puberulent. Stipules to 1 2 mm long, narrowly deltoid, gla­
brous or minutely white puberulent. Lamina 2-8 . 5 cm wide, 4- 1 5 . 5 cm
long, elliptic, oblong-elliptic, obovate, or oblanceolate; apex rounded ,
acute or abruptly short blunt acuminate; base cuneate or rounded cu­
neate; lateral veins 5- 1 5( - 1 7) , departing from the mid rib at an angle from
1 0° -30° ( -40° ); basal veins 1 -2, departing from the mid rib at a similar or
slightly steeper angle; intercostals generally slightly prominent. Petiole 4-
30 mm long, I I 3- 1 I 1 2 the length of the lamina .
Figs 4-8 mm in diameter, globose, glabrous, borne among the leaves
and on short shoots behind the leaves; colour yellowish or rose scarlet; pe­
duncle 1 -5 mm long, glab rous or minutely puberulent; basal bracts 0.5-
5.0 mm long, suborbicular, frequently apparently somewhat excentric,
sometimes irregularly splitting to give 3-4 lobes, glabrous or minutely pu­
berulent; ostiole 1 -3 mm in diameter, flat. Female flowers with 2-3
hooded tepals, 0.8- 1 .0 mm long, sessile or stalked. M ale flowers with 2-3
hooded tepals, 0 . 8- 1 .0 mm long, sessile or short-stalked.
Leaf structure : cystoliths present, but generally few, on upper and
lower surfaces; sclereids absent; lower epidermis plane, or somewhat sin­
uate and plicate; stomates sunken in small-mouthed pits, the mouth
either smooth ( if the lower epidermis is plane) or rugose (if the lower
epidermis is sinuate and plicate); tabular crystal cells absent; druse cells
absent; pubescence absent.

D i s t r i b u t i o n : Trinidad and Tobago, along the coast of S outh

America to Rio de Janeiro .

In Surinam known from the M iddle Saramacca River and the Perica
River.

Lindeman 5425; Maguire 23962.

 MORACEAE 245

5. Ficus ernestiana Pittier, Bol. Soc. Venez. C i . Nat. 4: 5 5 . 1 93 7 .

T y p e : Pittier 1 2522, Venezuela, Gmirico ( Y EN ) .

Ficus paludica Stand ley, Bull. Torrey Club 7 5 : 298. 1 948.
T y p e : Maguire & Stahel 22790, S urinam (F).

Tree 8-20 m tall. Twigs 2.5-4 mm in diameter, glabrous or minutely pubes­

cent. Stipules 1 0-25 mm long, acuminate, deltoid, glabrous or minutely
pubescent. Lamina 4.5- 1 0 cm wide, 7- 1 8 cm long, lanceolate, ovate, ob­
long-lanceolate or oblanceolate; apex bluntly acuminate; base rounded
to somewhat emarginate; lateral veins 7- l l , departing from the mid rib at
an angle from ( 1 0° -)20° -30° ; basal veins not or only slightly different; in­
tercostals not prominent. Petiole 20-40 mm long, 1 / 3- l / 5 the length of
t he lamina, minutely pubescent to glabrous.
Figs 9- 1 1 mm in diameter, ± globose or globose-pyriform, glabrous,
borne among or j ust behind the leaves; peduncles l -2 mm long, basal
bracts 2, semi circular, ± 2 m m long, pubescent, somewhat reflexed; ori­
fice about 2 mm in diameter, somewhat mammilate. M ale and female
flowers about I mm long, with 3 hooded tepals. Scales present between
the flowers .
Leaf structure: cystoliths on both surfaces of the leaf: sclereids absent;
lower epidermal cells plane; stomates plane with the surface of the leaf;
druse cells present in the mesophyll; tabular crystal cells absent.

D i s t ri b u t i o n : Venezuela and Surinam.

In Surinam in the young coastal plain; in swamp forests.

Lanj ouw 1 1 56; Lanjouw & Lindeman 477, I 1 05, 1 1 68, 1 395; LBB 1 2522;
Lindeman 5702; Maguire & Stahel 22790; Stahel 23.

6. Ficus erratica S tand ley, Bull. Torrey Club 75: 295 . 1 948.

T y p e : Fanshawe 2436, Guyana ( F) .

Ficus mensalis Standley, Bull. Torrey Club Club 75: 297. 1 948.
T y p e : Maguire 24584, Surinam (F).
246 MORACEAE

Weakly or diffusely branched glabrous tree to ± 1 5 m tall. Twigs 3-5 mm

in diameter, glabrous. Stipules not seen ( 1 2- 1 8 mm long fide Standley).
Lamina 2. 7-7 cm wide, 6- 1 5 cm long, elliptic-oblong or oblanceolate;
apex generally abruptly acuminate; base cuneate; lateral veins about I 5 ,
departing from the mid rib at an angle from 0° - 1 0° , basal veins departing
at a slight steeper angle; intercostals not prominent. Petiole I 0-25 mm
long, I I 6- I I 7 the length of the lamina.
Figs 5-7 mm in diameter, globose, glabrous, reddish in colour, borne
on minute short shoots among and behind the leaves; peduncle 3-4 mm
long; basal bracts 2, semi-circular, I mm long, glabrous; orifice about I
mm in diameter. Male flowers around the orifice, male and female flow­
ers about I mm long with 3 hooded tepals; scales present between the
flowers.
Leaf structure: cystoliths on the lower surface of the leaf, absent above;
sclereids absent; lower epidermal cells plain; stomates sunken into pits in
the cuticle; druse cells absent; tabular crystal cells present on the veins.

D i s t r i b u t i'o n : From Guyana to Amapa ( Brazil) .

In Surinam only known from the Tafelberg.

Maguire 24563, 24584.

7. Ficus gomelleira Kunth & Bouche, Ind. Sem. H ort. Berol. I 8 . I 846.
Urostigma gomelleira (Kunth & Bouche) M iquel, London Jour.
Bot. 6: 5 3 1 . I 847.

T y p e : specimen so named (B).

Urostigma doliarium Martius ex M iquel, London J our. Bot. 6: 527. 1 847.

Ficus doliaria Martius, Systema Materia Medicae Vegetabilis Brasiliensis
1 88 . 1 843; nomen.
Ficus doliaria ( M artius ex Miquel) Miquel, Ann. M us. Lugd . -Bat. 3: 297.
1 867.
Ty p e : H ort. Monac. s.n. ( M ) .
 MORACEAE 247

Ficus acaroianiensis Benoist. Arch. Bot. Caen 3: ( Bull.) : 1 70, 1 929.

T y p e : Sagot 526 ( isotypes BR, P ) .
Ficus pakkensis Stand ley, B u l l . Torrey C l u b 75: 297. 1 948.
T y p e : M aguire 23960, Surinam ( isotype U ) .

Tree to 30 m or more tall. Twigs 4-6 mm in diameter, pubescent. Stipules

to 1 5 mm long, deltoid , densely appressed brown pubescent. Lamina 6.5-
l 4(-20) cm wide, 9. 5-24(-38) cm long, broadly ovate or elliptic; apex
rounded or acute; base rounded , truncate and emarginate or weakly cor­
date; lateral veins 7- 1 4( - 1 6), departing fro m the midrib at an angle from
20° -40° ; basal veins 1 -2, departing at a similar angle; intercostals dis­
tinctly to extremely prominent. Petiole 1 5-55 mm long, 1 I 5- 1 I 8 the
length of the lamina, harshly pubescent.
Figs 1 7-20 mm in diameter, globose or subpyriform, pubescent, borne
among the leaves; colour "green"; peduncle 4-7 mm long, pubescent; bas­
al bracts 2-5 mm long, semicircular, pubescent; o stiole 1 -2 mm in diame­
ter, plane with the surface or slightly sunken. Female flowers with 2-3
h ooded tepa1s, 1 . 5-2 mm long. M ale flowers long stalked with 2-3 hooded
tepals, 1 . 5-2 mm long. Scales present between the flowers.
Leaf structure: cystoliths small and numerous on the upper surface,
few or lacking on the lower surface; sclereids abundant; lower epidermal
cells plane, somewhat striate or strongly plicate and sinuate; stomates su­
perficial; druse cells absent; tabular crystal cells absent; pubescence of
long, multiseptate hairs, to 1 mm long, and short conical hairs on the
lower surface, sparingly on the upper surface of the leaf, abundant on pe­
tioles, twigs and stipules.

D is t r i b u t i o n : Coastal mountains Venezuela to Brazil.

I n Surinam collected in the Jodensavanne - M apane Creek area, along

the Perica R iver, and along the M iddle Saramacca River.

Lindeman 52 1 5 , 5383; Maguire 23960.

248 M O R ACEAE

Fig. 10 Ficus /anjouwii; a . , twig (c. V:! x); b. , female flower (c. 7Y:! x); c. and d. ,
male flower (7Y:! x); e., lower surface of leaf (c. 300 x), without cellular detail,
showing tabular crystals of calcium oxalate along the veins, ( I ) and druses of calcium
oxalate (2); f. , lower surface of leaf (c. 300 x), showing cellular detail, at left,
view of epidermis showing stomates and (top) a cystolith, at right, subepidermal view
showing sclereids along the veins, tabular crystals of calcium oxalate along the
V
 M O RACEAE 249

8. Ficus lanjouwii DeWolf, Jour. Arnold Arb. 50: 478. 1 969. 1

T y p e : B W 4639, Surinam (A).

Tree. Twigs 3-4 mm in diameter, densely pubescent with ascending hairs

on new growth. Stipules 6-8 mm long, acuminate, deltoid, densely ap­
pressed pubescent. Lamina 2. 5-5 cm wide, 7.0- 1 4. 0 cm long, oblanceo­
late; apex acuminate; base cuneate; lateral veins 1 0 1 1 5 pairs, departing
from the midrib at an angle from 20° -30° ; basal veins l pair, departing
from the mid rib at an angle from 50° -60° ; intercostals very slightly
raised . Petiole )- 1 5 mm long, l I 1 3- 1 I 1 5 the length of the lamina, densely
appressed pubescent.
Figs 7-9 mm in d iameter, ± globose-pyriform, borne in pairs among the
leaves; peduncles l -2 mm long, densely appressed pubescent: basal bracts
semicircular, about l mm across densely appressed pubescent; ostiole
plane with the surface of the fig about l mm in diameter. Female and gall
flowers about 1 .0- 1 .2 mm long with 3 narrowly lanceolate tepals with hy­
aline margins. Scales present between the flowers.
Leaf structure: cystoliths few on the upper surface, more abundant on
t he lower surface; sclereids present generally along the veins; lower
epidermal cells plane; stomates superficial; d ruses present in the meso­
phyll cells; tabular crystal cells present.

D i s t r i b u t i o n : Only known from Surinam.

In Surinam collected in three localities in the savanna belt in eastern

Surinam.
BW 3639, 4639, 5859; H eyligers 70 1 ; Lindeman 6088.

This species appears to be allied to the widespread Ficus trigona L.f. It

differs from that species in a number of characters : 1 . the greate r

' I take pleasure in commemorating the name of the Senior Editor of the Flora of
Suriname who has been connected with studies of the M o raceae for more than thirty
years.
250 M O R ACEAE

number of lateral veins in the leaves ( 1 0- 1 5 pairs versus 3-9 pairs)

and their lower angle of departure from the midrib; 2. the orifice of
the fig is not surrounded by a raised rib of receptacular tissue; 3. the te­
pals of the flowers are narrowly lanceolate (not hooded).

9. Ficus longifolia S chott in S p rengel, Syst. Veg. 4( Cur. Post. App.):

409 . 1 8 27.
U rostigma longifolium (Schott in Sprengel) M iquel, FI. Bras.
4( l ) : 95. 1 8 53.

T y p e : Schott s.n., Brazil ( B).

Ficus caballina Standley, Pub!. Field M us. Bot. 1 3( 2): 30 1 . 1 937.

T y p e : Ll. Williams 2075 , Peru, Loreto ( F).
Ficus ramiflora Standley, Pub!. Field M us. Bot. 1 7 : 1 75. 1 937.
T y p e : K rukoff 6446, Brazil, Amazonas (NY).

Epiphytic shrub, "strangling vine" or small tree, to 10 m tall, glabrous .

Twigs 7- 1 0 mm in diameter, glabrous. Stipules 1 5-23 mm long, glabrous,
broadly deltoid . Lamina 5 - 1 1 . 5 cm wide, 1 5-47 cm long, narrowly ellip­
tic, spathulate or obovate; apex rounded, acute or abruptly short acumi­
nate; base cuneate; lateral veins (8-) 1 3-23 , departing from the midrib at
an angle from 20° -30° ( -40° ); basal veins I , departing from the mid rib at
an angle from 30° -40° ; intercostals not or only slightly raised . Petiole .7-
30 mm long, 1 I 9- 1 I 1 4( -I I 35) the length of the lamina.
Figs 5-8 mm in diameter, globose, glabrous or minutely puberulent,
borne on short shoots generally behind the leaves; colour red , reddish­
yellow or yellow, with or without darker spots; peduncle obsolete to 5
mm long, puberulent; basal bracts to 2 mm long, semicircular, puberu­
lent; ostiole 1 -2 mm in diameter, flat. Female flowers about I mm long,
with 2-3 hooded tepals. M ale flowers about I mm long, sessile, with 2-3
hooded tepa ls. Scales present between the flowers.
Leaf structure: cystoliths abundant on both surfaces; sclereids
abundant; lower epidermal cells plane, or more or less sinuate and
 MORACEAE 25 1

plicate; stomates more or less sunken into small mouthed pits; tabular
crystal cells absent; pubescence of a few 2-cellular, glandular hairs.

D i s t r i b u t i o n : The Amazon Basin and the Guianas.

In Surinam known from the U pper Suriname River and the Tapanahony
River.

BW 4 1 45; Stahel Exp. Wilh. Geb. 1 42.

1 0 . Ficus mathewsii ( M iquel) Miquel, Ann. N.t us. Lugd . -Bat. 3: 298 .
1 867.
U rostigma mathewsii Miquel, London J our. Bot. 6: 549. 1 847.

T y p e : Mathews 2059, Peru ( K) .

Ficus gleasonii Standley, Pub!. Field M us. Bot. 1 7: 1 70. 1 937, descr. ;
Kribs, Tropical Woods 1 3 : 34. 1 928, nomen.
T y p e : De La Cruz 2396. Guyana (isotype NY).
Ficus sprucei Standley, Pub!. Field M us. Bot. 1 7: 1 76. 1 937.
T y p e : Spruce 2804, Brazil, A mazonas (isotypes B R , C, CGE, E, K).

Tree 3- 1 6 m tall. Twigs 2-4 mm in diameter, generally minutely puberu­

lent, sometimes glabrous, the epidermis frequently exfoliating. Stipules
to 9 mm long, generally minutely puberulent, sometimes glau­
cous. Lamina 1 -4 cm wide, 4.5- 1 2 cm long, oblanceolate; apex rounded,
bluntly acute or acuminate; base cuneate; lateral veins 1 0- 1 7, departing
from the midrib at an angle from I oo -20° ; basal veins 1 -2, departing from
the mid rib at a similar angle; intercostals not or scarcely prominent. Pe­
tiole 9-26 mm long, I I 4- 1 I 1 6 the length of the lamina.
Figs 4-5 mm in diameter, globose, generally minutely puberulent,
borne among the leaves on new shoots or behind the leaves on specialized
short shoots on older twigs; colour yellowish or reddish; peduncle 1 -2
mm long puberulent; basal bracts about 2 mm in diameter, flat. Male
252 M O R ACEAE

and female flowers about I mm long, with 2-3 hooded tepals. Male
flowers sessile. Scales present between the flowers.
Leaf structure : cystoliths present on both upper and lower surfaces:
sclereids absent; lower epidermal cells plane; stomates somewhat
sunken; druse cells absent; tabular crystal cells usually present on the
lower surface of the veins; pubescence absent, or a few long, white multi­
septate hairs present.

D i s t r i b u t i o n : Surinam and Guyana to Colombia southward to east­

en Bolivia and the western Mato G rosso in Brasil.

In Surinam only known from a locality on the lower Corantine River.

BW 2 1 90.

1 1 . Ficus nymphaeaefolia P. M iller, Gard. Diet. (ed. 8). 1 768. Ficus

nymphaeifolia Linnaeus, Mantissa 305 . 1 77 1 .
Ficus nymphoides Thunberg, Ficus Genus, Dissertatione 7 . 1 786.
U rostigma nymphaeifolium ( Linnaeus) M iquel, London J our.
Bot. 6: 527. 1 847.

T y p e : Miller s.n. (BM).

Ficus ierensis Britton, Bull. Torrey Club 4 8 : 329. 1 9 1 1 .

T y p e : Britton, H azen & Mendelson 774, Trinidad (isotypes G H , TRIN).
Ficus anguina Benoist, Bull. M us. H ist. Nat. Paris 30: 1 04, 1 924.
S y n t y p e s : Sagot 1 1 54, French Guiana (B, BR), Benoist 343, French
Guiana (P).
F. duquei Dugand , Caldasia 1 (4): 42. 1 942.
T y p e : Duque 265, Colombia, Tolima (COL).

Tree 1 5-3 5 m tall, becoming massive with age, glabrous. Twigs 4- 1 3 mm

in diameter, glabrous or minutely puberulent. Stipules to 40 mm long,
lanceolate or deltoid, glabrous or minutely puberulent, or glabrous. La­
mina 6.5- 1 7 cm wide, 9-23 cm long, ovate or ovate-rotund ; apex acute,
 MORACEAE 253

Fig. 1 1 Ficus nymphaeaefolia; a . , twig (c. 1 / 4 x); b . , lower epidermis of leaf

(c. 300 x), showing cellular detail, sclereids indicated by dashed lines, clavate uni­
cellular hair at lower left, druses of calcium oxalata scattered in cells; c., upper epider­
mis (c. 300 x), showing cellular detail, sclereids indicated by dashed lines, cystolithes
indicated by dashed circles.
254 M O R ACEAE

broadly acute , rounded or sometimes shortly acuminate; base cordate; la­

teral veins (4-)5- 1 1 , departing from the mid rib at an angle from 1 0° -40° ;
basal veins 1 -3 , departing from the mid rib at an angle from 0° -20° ; inter­
costals not prominent. Petiole 30- 1 25 mm long, 1 / 2- 1 / 4 the length of the
lamina .
Figs 1 5-25 mm in diameter, globose, minutely puberulent or glaucous,
bo rne among the leaves; colour greenish, with purple spots; ped uncle ob­
solete to 5 mm long; bracts 4- 1 9 mm long, deltoid or ovate , connate , mi­
nutely puberulent or glaucous; ostiole 2-3 mm in diameter, probably pla­
ne with the surface, but in dried mate rial frequent ly raised, or surrounded
by a more or less distinct ring of receptacular tissue. Female flowers
about 2 mm long, with th ree hooded tepa ls. Male flowers stalked , about
2 mm long, with three hooded tepals. Bracts present between the flowers.
Leaf structure : cysto liths present on both surfaces; sclereids present
and abundant; lower epidermal cells nearly plane or with the cells va ri­
ously sinuate and plicate; stomates superficial; tabular crystal cells gener­
ally absent, but present on the veins of the lower surface in material from
Colombia; druse cells generally absent.

D i s t r i b u t i o n : Panama, Colombia and northern South America to

the State of Ceara in Brazil.

In S urinam in the young coastal plain and the savanna belt; known from
four localities from the Coppename River eastwards.

BW 1 502, 2494, 3370, 44 1 0, 5864; Lindeman 53 1 5, 56 1 2.

1 2. Ficus paraensis ( M iq uel) M iquel, Ann. M us. Lugd . - Bat. 3 : 298. 1 867.

Urostigma paraense M iquel, London J our. Bot. 6: 534. 1 847.

S y n t y p e s : M artius 1 2 1 7, 1 48, 1 49, Brazi l, Para ( M); Parker

s.n., Guyana ( K ).
 M O R ACEAE 255

Fig. 1 2 Ficus paraensis; a . , twig (c. 1 / 4 x); b . , upper epidermis of leaf (c. 300 x),
showing cystolith as dashed circle; c . , lower epidermis of leaf (c. 300 x), showing
tabular crystals cells in superficial cells of the veins ( Steyermark 99937).
256 M O RACEAE

Ficus panamensis Stand ley, Contrib. U . S . Nat. H erb. 20: 1 5 . 1 9 1 7 .

T y p e : Pittier 3908, Panama ( isotype G H ).
Ficus thelephora Benoist, Arch . Bot. Caen 3( Bull . ) : 1 7 1 . 1 929.
T y p e : Sa got 1 1 65, French Guiana ( P) .
Ficus haughtii Standley, Pub!. Field M us. Bot. 1 7 : 1 70. 1 937.
T y p e : Haught 1 466, Colombia, Santander (isotype K).
Ficus uberrima Standley, Pub!. Field . M us. Bot. 1 7 : 1 77. 1 937.
T y p e : K rukoff 5055, Brazil, Amazonas ( isotypes A, K , NY, US).
Ficus putumayonis Dugand , Caldasia 1 (4): 62. 1 942.
T y p e : Cuatrecasas 1 1 265, Colombia, Putumayo (COL, not seen; paratypes
Cuatrecasas 1 1 1 00, 1 1 2 1 3, U S ) .
Ficus arukensis Stand ley, Bull. Torrey Club 75: 295. 1 948.
T y p e : Fanshawe 2435 , Guyana ( isotypes A, K ) .

Small to large tree or shrub. Twigs 2-7 mm in diameter, glabrous or mi­

nutely puberulent. Stipules to 35 mm long, narrowly deltoid, glabrous or
minutely puberulent. Lamina 2. 5-9 cm wide, 6-27 cm long, oblong, ob­
long-elliptic, oblong-oblanceolate or oblanceolate; apex acuminate, the
acumen commonly 5- 1 0 mm long, but sometimes to 25 mm; base cor­
date, emarginate, rounded or cuneate; lateral veins 9- 1 8 , departing from
the midrib at an angle from I oo -30° ; basal veins 1 -3 , at least the strongest
departing from the mid rib at an angle from 40° -60° ; intercostals
scarcely, or somewhat prominent. Petiole 5-35( -80) mm long, I I 3- 1 I 9
( - 1 I 1 1 ) the length of the lamina.
Figs I 0- 1 8 mm in diameter, globose to subpyriform, glabrous or mi­
nutely puberulent, borne among the leaves; colour greenish or yellowish,
sometimes with red streaks and sometimes with crimson basal bracts; pe­
duncle obsolete to 3 mm long; basal bracts 2-5 mm long, ovate, connate,
puberulent; ostiole 2-4 mm in diameter, umbonate, to 3 mm high. Female
flowers 1 . 5-2 mm long, with 2-3 very membranous hooded tepals . Male
flowers to 1 mm long, sessile, with 2-3 membranous hooded tepals.
Scales present between the flowers.
Leaf structure: cystoliths present on the upper surface of the leaf,
seldom on the lower surface: sclereid s absent; some lower epidermal
cells papillate, frequently conspicuously so; stomates superficial; druse
cells absent; tabular crystal cells sometimes present on the lower surface
of the veins; pubescence of two-cellular, glandular hairs.
 M O R ACEAE 257

D i s t r i b u t i o n : British H onduras southward m lowland forest

to and through the Amazon Basin.

In Surinam collected in three localities in the savanna belt.

BW 2 1 26; van Donselaar 3497; Lindeman 6942; van der Vecht s . n .

1 3 . "Ficus pertusa Linnaeus fil. , Suppl. 442. 1 78 1 .

U rostigma pertusum (Linnaeus fil.) M iquel, London Jour.
Bot. 6: 548. 1 847.

T y p e : Herb. Linn. 1 240. 9 ( LINN).

Ficus padifolia H . B. K . , Nov. Gen. et Sp. 2: 47 (or 38). 1 8 1 7 .

U rostigma padifolium ( H . B. K . ) Liebmann, Kgl. Danske Vidensk. Selsk.
Skr. V. 2: 324. 1 85 1 .
T y p e : H umboldt & Bonpland s . n . , M exico (P).
Ficus complicata H . B . K . , Nov. Gen. et Sp. 2: 48 (or 39). 1 8 1 7.
U rostigma complicatum ( H . B. K . ) Liebman, Kgl. Danske Vidensk. Selsk.
Skr. V . 2: 3 24. 1 85 1 .
T y p e : H umboldt & Bonpland s . n . , M exico ( P) .
Ficus ciliolosa Link, Enum. P I . Berol. 2: 450. 1 822.
T y p e : Link s . n . ( B).
Ficus myrtifolia Link, Enum. PI. Berol. 2: 450. 1 822.
T y p e : Link s.n. ( B).
Ficus arbutifolia Link, Enum. PI. Berol. 2: 450. 1 822.
U rostigma arbutifolium ( Link) M iquel, Versl. Kon. Akad. Wetensch.
Amsterdam 1 3 : 4 1 2. 1 862.
T y p e : Link s . n . ( B).
Ficus cestrifolia Schott in Sprengel, Syst. Veg. 4(Cur. Post. App.): 409. 1 827.
U rostigma cestrifolium (Schott in Sprengel) M iquel, London J our. Bot. 6:
54 1 . 1 847.
T y p e : Schott s . n . , Brazil ( B).
Ficus lancifolia H ooker & Arnott, Bot. Beech. Voy. 3 1 0. 1 838.
U rostigma lancifolium ( H ooker & Arnott) M iquel, London J our. Bot.
6: 538. 1 847.
T y p e : a sheet so labeled in Herb. Hook. (K).
Ficus subtriplinervia M artius, Flora 24( Beibl. 2): 67. 1 84 1 .
U rostigma subtriplinervium ( Martius) M iquel, London Jour. Bot. 6 : 542.
1 847.
258 MORACEAE

T y p e : M a rt ius 1 79, Brazil, M inas Gerais ( M , not the specimen photographed

by Macbride).
Ficus arpazusa Casa retto, Nov. Stirp. B ras. Dec. 1 5 . 1 842.
T y p e : Casaretto 1 234, Brazil, Rio de Janeiro (TO).
Ficus cerasifolia Kunth & Bouche, l nd . Sem. H ort. Berol. 1 6. 1 846.
T y p e : Kunth & Bouche s . n . ( B).
Ficus consanguinea Kunth & Bouche, l nd. Sem. H ort. Berol. 1 7. 1 846.
U rostigma consangineum ( K unth & Bouche) M iquel, Versl. Kon. Akad.
Wetensch . Amsterdam 1 3 : 408. 1 862.
T y p e : Kunth & Bouche s.n. ( B).
Ficus ligustrina Kunth & Bouche, l nd . Sem. H o rt. Berol. 1 6. 1 846.
T y p e : Kunth & Bouche s . n . ( B).
Ficus planicostata Kunth & Bouche, l nd . Sem. H ort. Berol. 1 6. 1 846.
U rostigma plan icostatum ( K unth & Bouche) M iquel, London J our. Bot. 6:
586. 1 847.
T y p e Kunth & Bouche s.n. ( B) .
Ficus periplocaefolia Kunth & Bouche, l nd . S e m . H ort. Berol. 1 6. 1 846.
T y p e : Kunth & Bouche s . n . ( B).
U rostigma erythrostictum M iquel, London J our. Bot. 6: 540. 1 847.
Ficus eryth rosticta ( M iquel) M iquel, Ann. M us. Lugd. -Bat. 3 : 298. 1 867.
T y p e : Parker s . n . , Guyana ( K ) .
U rostigma geminum Ruiz e x Miquel, London J our. Bot. 6: 547. 1 847.
Ficus gemina ( R uiz ex M iquel) M iquel, Ann. M us. Lugd. - Bat. 3 : 298. 1 867.
T y p e : Ruiz s . n . , Peru ( isotypes B, Fl, P).
U rostigma schiedeanum M iquel, London Jour. Bot. 6: 539. 1 847.
Ficus schiediana ( M iquel, Ann. M us. Lugd . -Bat. 3 : 298. 1 867.
T y p e : Schiede & Deppe 1 1 6, Mexico ( U ) .
Pharmacosycea peruviana M iquel, London. J our. Bot. 7 : 7 2 . 1 848.
Ficus peruviana ( M iquel) Rossberg, Repert. Sp. Nov. 42: 6 1 . 1 937.
T y p e : M athews 206 1 , Peru ( K ).
U rostigma baccatum Liebmann, Kgl. Danske Vidensk. Selsk. Skr. V . 2: 327.
1 85 1 .
Ficus baccata ( L iebmann) M iquel, Ann. M us. Lugd. -Bat. 3 : 299. 1 867.
T y p e : Liebmann 1 4309, Mexico (C).
U rostigma sapidum Liebmann, Kgl. Danske. Vidensk. Selsk. S k r. V . 2: 327.
1 85 1 .
Ficus sapida ( Liebmann) M iquel, Ann. M us. Lugd . -Bat. 3 : 298. 1 867.
T y p e : Oersted 1 4327, Costa Rica (C).
U rostigma turbinatum Liebmann, Kgl. Danske Vidensk. Selsk. Skr. V . 2:
328. 1 85 1 .
Ficus turbinata ( L iebmann) M iquel, Ann. M us. Lugd. -Bat. 3 : 298. 1 867.
T y p e : Liebmann 1 4336, M exico (C).
U rostigma sulcipes M iquel, Versl. Kon. Akad. Wetensch. Amsterdam 1 3 :
4 1 3 . 1 862.
Ficus sulcipes ( M iquel) M iquel, Ann. M us. Lugd . - Bat. 3: 298. 1 867.
T y p e : Sch iede & Deppe 1 09 1 , M exico ( U ) .
 M O R ACEAE 259

Ficus ochroleuca G risebach, Fl. B rit. W . Ind. Isl. ! 5 1 . 1 864.

T y p e : Prior s . n . , Jamaica ( K) .
Ficus faydeni M iquel, Ann. M us. Lugd . -Bat. 3 : 2 1 9. 1 867.
T y p e : Me. Fadyen s . n . , Jamaica ( U ) .
Ficus palmicida Pittier, B o l . Soc. Venez. C i . Nat. 4: 69. 1 93 7 .
T y p e : Pittier 1 23 24, Venezuela, G uarico (isotype NY).
Ficus kanukuensis Standley, Lloydia 2: 1 74. 1 939.
T y p e : A . C . Smith 3 1 43, G uyana ( isotype ! A N ) .

Large spreading tree to 30 m or more tall. Twigs l -2 mm in diameter, gla­

brous. Stipules 5-7 mm long, narrowly deltoid, glabrous or minutely pu­
berulent. Lamina l -4 . 5 cm wide, 2.5- 1 1 cm long, elliptic or lanceolate;
apex bluntly acute, acute or acuminate, the acumen 5-20 mm long; base
commonly cuneate, sometimes rounded cuneate, seldom rounded and
slightly emarginate; lateral veins (4-)6- 1 2, departing from the midrib at
an angle from 20° -60° ; intercostals somewhat raised. Petiole 5- 1 3 mm
long, l I 5- l I 8 ( -1 I 1 2) as long as the lamina.
Figs 5- 1 8 mm in diameter, globose, glabrous, borne among the leaves;
colour pink, yellow or green with brown flecks; peduncle 2-5 mm long,
glabro us; basal bracts 1 -2 mm long, semicircular, glabrous; ostiole l -2
mm in diameter, crateriform, the bracts sunken into the receptacle or sur­
rounded by a raised ring or receptacular tissue. M ale and female flowers
about l mm long, with 2-3 hooded tepals. M ale flowers sessile. Scales
present between the flowers.
Leaf structure: cystoliths on both upper and lower surfaces; sclereids
absent or very weakly developed ; lower epidermal cells plane, striate
or slightly sinuate and plicate; stomates superficial; tabular crystal cells
usually abundant in the lower surface of the veins; druse cells usually
present but generally obscured by the presence of dark brown inclusions
in the lower epidermal cells; pubescence absent.

D i s t r i b u t i o n : W idespread in tropical America, from Mexico,

through Central America and the West l ndies to Brazil; in dry evergreen
or deciduous forest from sea level to l 000 m.

I n Surinam throughout the country; common along streams and m

swamps.
260 MORACEAE

BW 1 83 5 , 3892, 4902; van Donselaar 1 1 70; Focke 1 385; H ekk ing 1 1 45; Kegel 342,
356 ( P); K ramer & Hekking 27 1 4; K uyper 22; Lanj ouw 1 1 4, 978; Lanj ouw &. Lin­
deman 1 47 1 ; LBB I 0629; Lindeman 4994, 5580, 5583. 5597, 5703, 6626, 6638;
Maguire 23928a; Mennega 4 1 9; Pulle 448; Reij enga 4 1 8 ; Samuels 43 1 ( P); Stahel
24, Woodherb. 238; Went 266, 552; Wullschliigel 469 ( BR ) , 1 08 1 ( B R ) , s.n. ( BR).

14. Ficus prinoides H umboldt & Bonpland ex Willdenow, Sp. PI. 4(2) :
1 1 49. 1 806.
U rostigma prinoides (Willdenow) Miquel, London J our. Bot.
6: 540. 1 847.

T y p e : Humbo ldt & Bonpland s . n . , Colombia ( B) .

Ficus pallida Vahl, Enum. 2: 1 94. 1 806.

U rostigma pallidum ( Vahl) M iquel. London. J our. Bot. 6: 5 30. 1 847.
T y p e : von Rohr s . n . , Colombia, M agdalena (C).

Tree to 30 m tall. Twigs about 2 mm in diameter, minutely pubescent or

glabrate. Stipules to 1 0 mm long, glabrous or minutely pubescent. La­
mina 1 -3 . 5 cm wide, 3- 1 0 cm long, elliptic, elliptic-oblong, lanceolate, or
oblanceolate; apex rounded , blunt, acute or slightly acuminate; base cu­
neate or cuneate-truncate; lateral veins ( 5-)8- 1 2(- 1 6) , departing from the
mid rib at an angle from 20° -40° ; basal veins 1 -2, departing from the mid­
rib at an angle from 40° -50° ; intercostals scarcely raised . Petiole 4-20
mm long, I I 4- I I 8 ( - 1 I 1 6) the length of the lamina.
Figs 4-6 mm in diameter, globose, glabrous or glabrescent, borae
among the leaves; colour green with red spots or whitisch dotted with
brown or green; peduncle 2-4 mm long, minutely puberulent; basal bracts
0. 5- 1 (-2) mm long, minutely puberulent, semicircular; ostiole about 2
mm in diameter, flat or slightly sunken. Male and female flowers about
I mm long, with 2-3 hooded tepals. Male flowers sessile or short stalked.
Scales present between the flowers.
Leaf structure: cystoliths present on both upper and lower surfaces of
the leaf; sclereids absent; lower epidermal cells plane; stomates superfi­
cial; druse cells present; tabular crystal cells present on the lower surface
 MORACEAE 26 1

of the veins; pubescence absent. Lower mesophyll extremely dense and

dark coloured, obscuring the lower epidermis.

D i s t r i b u t i o n : Northern coast of S outh America from Colombia

to Brazil.

In Surinam known by a single collection from the Zuid River near the
confluence with the Lucie River.

l rwin, Prance, S oderstrom & H o lmgren 55800.

1 5 . Ficus trigona Linnaeus fil. , Suppl. 44 1 . 1 78 1 .

U rostigma trigonum (Linnaeus fil.) Miquel, London J our. Bot.
6: 548. 1 847.

T y p e : H erb . Linn . 1 240. 1 2 ( LI N N) .

U rostigma fagifolium M iquel, London Jour. Bot. 6: 544. 1 847.

Ficus fagifolia ( M iquel) M iquel, Ann. M us. Lugd . -Bat. 3 : 299. 1 867.
T y p e : Martius 25, B razil, Pani ( U ).
U rostigma hartwegii M iquel, London Jour. Bot. 6: 545. 1 847; pro parte.
T y p e : H artweg 1 384, Colombia ( K) .
U rostigma euomphalum Miquel in Martius. F l . B ras. 4( 1 ): 1 03. 1 853.
Ficus euomphalum ( M iquel) M iquel, Ann. M us. Lugd. -Bat. 3 : 299. 1 867.
T y p e : Gardner 3425, B razil ( K).
Ficus colubrinae Standley, Contrib. U . S . Nat. H erb. 20: 1 6. 1 9 1 7.
T y p e : von Tiirckheim l l . 1 56, G uatemala ( U . S, not seen) .
Ficus vulpina Benoist, Bull. M us. H ist. Nat. Paris 11 . 1 : 1 64. 1 929.
T y p e : Sagot 524, French Guiana ( P).
Ficus llewelyni Standley, Publ. Field M us. Bot. 1 3( 2) : 304. 1 937.
T y p e : Williams 2308, Peru, Loreto ( isotype B).
Ficus weberbaueri Standley, Publ. Field M us. 1 3(2): 308. 1 937; W illiams,
Publ. Field M us. 1 5 : 84. 1 936.
T y p e : Weberbauer 6954, Peru, Cuzco (isotypes B, U S M ) .
Ficus llanensis Dugand, Caldasia 1 (4) : 5 3 . 1 942.
S y n t y p e s : Cuatrecasas & Garcia-Barriga 4435 ( U S); Jaramillo 1 89 ( I A N ,
US).
Ficus fanshawei Standley, B u l l . Torrey Club 75: 295 . 1 948.
T y p e : Fanshawe 2363, Guyana ( isotype K).
262 MORACEAE

Fig. 1 3 Ficus trigona; a . , twig (c. l / 2 x); b . , upper epidermis of leaf (c. 300 x),
showing cystolith as dashed circle; c . , lower epidermis of leaf (c. 300 x), showing
sclereids as dashed lines, and tabular cells in the superficial cells of the veins
( M aguire, Schulz, Soderstrom & H o lmgren 54 1 59).
 MORACEAE 263

Ficus maguirei Standley, Bull. Torrey Club 75: 296. 1 948. ..;
T y p e : M aguire and Fanshawe 23528, Guyana ( i sotypes A, K).
Ficus arctocarpa Stand ley, Bull. Torrey Club 7 5 : 294. 1 948.
T y p e : M aguire 23865, Surinam ( isotypes N Y , U).

Tree or epiphytic shrub to 35 m tall. Twigs 2-7 mm in diameter, at least

the young growths appressed pubescent (pubescence to I mm long) or
rarely glabrous. Stipules 7-25 mm long, deltoid, appressed pubescent.
Lamina 2- 1 0 cm wide, 3. 5-20 cm long, narrowly or broadly elliptic, ovate
or obovate; apex blunt, acute or short and abrupt acuminate, the
acumen 6- 1 2 mm long; base cuneate or rounded , sometimes slightly
emarginate; lateral veins 3-9, departing from the midrib at an angle
from 30° -60° ; basal veins 1 -2 departing from the mid rib at a similar
angle; intercostals not prominent. Petiole 6-40 mm long, appressed
pubescent or glabrous, 1 / 4- 1 / 9( - 1 / 1 2) the length of the lamina.
Figs 6- 1 2 mm in diameter, globose, glabrous, minutely puberulent, or
more or less appressed pubescent, borne among the leaves; colour green
or olive green; peduncle obsolete, more commonly about 2 mm long, or
up to 4 mm long, pubescent; basal bracts 2-3 mm long, ovate, connate,
generally appressed pubescent; ostiole 1 -3 mm in diameter, surrounded
by a thin, e rect or incurved rim of receptacular tissue. Male and female
flowers with 2-3 hooded tepals, 1 - 1 . 5 mm long. Male flowers usually
sessile, sometimes short stalked . Scales and bristles present between the
flowers.
Leaf structure: cystoliths present on the upper surface, few or lacking
on the lower surface; sclereids absent in mate � ial from eastern South
America, present in material from the U pper Amazon and Rio Negro
d rainages; lower epidermal cells plane or slightly sinuate and plicate;
stomates superficial; druse cells absent; tabular crystal cells sometimes
present; pubescence of multiseptate hairs on veins, petiole, stipules and
twigs.

D i s t r i b u t i o n : Northern S outh America.

In Surinam apparently throughout the country; often in or near swamps,

or along streams.
264 MORACEAE

Lindeman 5 1 1 0, 5584, 56 1 3 , 63 1 8 ; Maguire 23865; Maguire, Schulz, Soderstrom &

H olmgren 54 1 59; Mennega 369; Pulle 438; WullschUigel 1 083 ( BR ) .

1 6. Ficus trigonata Linnaeus, PI. Surinam. 1 7. 1 775.

U rostigma trigonatum ( Linnaeus) Grisebach, G oett. Abh. 7 : 1 94.
1 857.

T y p e : P lumier, PI. Am. (ed . Burmann) t. 1 32, fig. I. 1 757.

Ficus crassinervia Desfontaines e x Willdenow, Sp. P I . 4(2): 1 1 3 8 . 1 806.

U rostigma crassinervium (Willdenow) M iquel, London J our. Bot. 6: 535.
1 847.
T y p e : A specimen so labelled , ex hort. Paris, in herb. Willd. ( B ) .
Ficus berteroi Warburg, Symbol. Antill. 3 : 468. 1 903.
T y p e : Bertero 2328, Jamaica ( B) .
Ficus eggersii Warburg, Symbol. Antill. 3 : 469. 1 903.
T y p e : Wright 1 444, Cuba ( isotype CGE).
Ficus mamillifera Warburg, Symbol. Antill. 3: 470. 1 903.
T y p e : H arris 5206, J amaica ( B) .
Ficus stahlii Warburg, Jamaica Symbol. Antill. 3 : 470. 1 903.
T y p e : Stahl 485, Puerto Rico ( B ) .
Ficus hartii Warburg, Symbol. AfJtill. 3 : 4 5 8 . 1 903.
T y p e : A lexander 5043, Trinidad (TR I N ) .
Ficus combsii Warburg, Symbol. Antill. 3 : 456. 1 903.
T y p e : Combs 500, Cuba (B).
Ficus mitrophora Warburg, Symbol. Antill. 3 : 457. 1 903.
T y p e : Ehrenberg 352, H aiti (B).
Ficus yucatanensis Standley, Contrib. U. S . Nat. H erb. 20: 3 3 . 1 9 1 7.
T y p e : Goldman 554, Mexico ( U S ) .
Ficus iturbei Pittier, B o l . S o c . Venz. C i . N a t . 4: 6 8 . 1 937.
T y p e : Pittier 1 23 1 9, Venezuela, G mirico ( Y E N ) .
Ficus interfector P ittier, Bol. S o c . Venez. Ci. N a t . 7 : 1 33 . 1 94 1 .
T y p e : Pittier 1 4746 Venezuela, Guarico (YEN) ) .
Ficus squamulosa Pittier, Bol. Soc. Venz. Ci. Nat. 7: 1 34. 1 94 1 .
T y p e : not cited .
Ficus perez-arbelaezii, Dugand , Caldasia 1 (4): 60, fig. 1 0. 1 942.
T y p e : Perez-Arbelaez 1 0072, Colombia, Cundinamarca (COL, not seen).
Ficus campbellii I . M. Johnston, Sargentia 8 : 1 1 4. 1 949.
T y p e : I . M . Johnston 72 1 , Panama (A).
Ficus leavensii I . M. J ohnston, Sargentia 8: 1 1 7. 1 949.
T y p e : I. M . J o h nston 1 1 25, Panama (A).
 MORACEAE 265

Large, spreading tree, to 25 m tall. Twigs 2-5 mm in diameter, glabrous or

minutely pubescent. Stipules 6-40 mm long, deltoid , glabrous or pubes­
cent. Lamina 1 . 5 - 1 0 . 5 cm wide, 4-32 cm long, oblong, elliptic or obovate;
apex generally obtuse, sometimes varying to slightly acuminate; base
cuneate, rounded or emarginate, sometimes slightly inequilateral;
lateral veins 5- 1 5 pairs, departing from the midrib at an angle from 20° -
400 ; basal veins I pair, departing from the mid rib at an angle from 30° -
50° . Petiole 5-25 mm long, I I 4- 1 I 9 the length of the lamina.
Figs 6-20(-30) mm in diameter, globose, minutely pubescent, borne
among the leaves, green when immature, reddish when mature; peduncle
0-9 mm long, glabrous or puberulent; basal bracts 2-6 mm long, broadly
deltoid or semicircular, sometimes variously split, glabrous, ciliolate or
pubescent; orifice 1 -3 mm in diameter, plane or slightly umbonate, or
with an erect rim of receptacular tissue.
Leaf structure: cystoliths present on the upper epidermis, few and
scattered on the lower surface; sclereid s absent; lower epidermal cells
p lane to somewhat sinuate and plicate; stomates superficial or somewhat
sunken; tabular crystal cells absent; druse cells occasionally present on
the lower surface; pubescence of 2-cellular glandular hairs on the lower
epidermis, short, conical, unicellular hairs may be present on either
surface, or may be entirely lacking.

D is t r i b u t i o n : Greater Antilles, Caribbean coast of Central America,

Venezuela and Surinam.

In S urinam a single collection from the U pper Saramacca River.

M aguire 24039

1 2. Pourouma Aublet, PI. Gui. 2: 89 1 . 1 775.

Puruma Jaume St. H ilaire, Expos. 2 : 3 1 3 . 1 80 5 .

T y p e s p e c i e s : Pourouma guianensis Aublet

266 MORACEAE

Dioecious trees, often with stilt roots. Leaves spirally arranged , entire to
palmately incised , pinnati- to palmatinervate; stipules fully amplexicaul,
connate. Inflorescences geminate or solitary in the axils of the leaves,
variously, mostly di- to trichotomously branched, sometimes un­
branched, often with scattered linear bracts and a basal bract. Stami­
nate flowers more or less densely glomerate at the ends of the inflores­
cence branches or partly solitary, sessile or pedicellate; perianth 4( -3)­
lobed to 4(-3)- parted ; stamens 4( -2), filaments straight in the bud,
connective small. Pistillate flowers solitary or clustered , usually pedicel­
late; perianth tubular, entire to 4( -3)-lobed to 4( -3)-dentate; ovary free,
ovule suborthotropous, subbasally attached, stigma usually discoid and
2-lobed; fruiting perianth fleshy (or dry?), fruit big, endocarp crusta­
ceous; seed without endosperm, cotyledons equal, thick, radicle apical,
short and surrounded by the cotyledons.

Distribution: About 30 species in tropical Central and South

America.

The genus comprises a number of clear-cut species, e.g. P. minor Benoist,

P. heterophylla M artius ex M iquel (syn . : P. myrmecophila Ducke),
and P. ovata Trecul (syn . : P. longipendulina Ducke), and also a number
of complexes in which the taxa (both specific and infraspecific) are
difficult to delimit owing to the present lack of knowledge about the
variability and the taxonomic value of the characters. Several characters,
such as leaf shape and indumentum of the inner surface of the stipules, to
which important diagnostic value has been attached (cf. Benoist, Bull.
Mus. H ist. Nat. Paris 28: 3 1 8 . 1 922), have to be used with caution. For
the reasons mentioned under Cecropia (p. 287) , I have not put names into
synonymy, with exception of a few in P. minor, one in P. maroniensis,
and one in P. guianensis.
As in Cecropia, the j uvenile parts of several species are more or less
densely covered with pluricellular granular hairs, which soon disintegrate
into a powdery layer. In general, if entire and divided leaves occur in
the same species, the leaves of fertile twigs are entire or are less divided
 MORACEAE 267

than those of sterile twigs. In such species, as they grow from seedling
to young sapling, the leaves soon become more divided and larger, and
it is only after the sapling stage that gradual reduction follows.

a. Leaves scabrous to scabridulous above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

b . Leaves smooth above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
.

2 a. Young twigs with dense granular hairs or (sub)sericeous intermixed with

sparse granular hairs; leaves often incised . . . . . . . . . . . . . . . I . P. guianensis
b. Young twigs with sparse granular hairs and pale yellow patent hairs; leaves
usually entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. P. velutina
3 a. Stipules, twigs and inflorescences with white a rachnoid indumentum . . . . . . .
. . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6. P. maroniensis
b . Stipules, twigs and inflorescences with white a rachnoid indumentum
. . .. . .. . . .. . .. . ... . . . . . .. . .. . . . .. . . . . .. .. .. . . . . . .. . . .. . . . . . . . . . . . 4
4 a. Stipules, twigs, and inflorescences pale yellow sericeous and with a few granu­
lar hairs; leaves entire and usually b roadest above the middle . . 7 . P . minor
b. Stipules, twigs, and inflorescences not yellow sericeous but with dense granu-
lar hairs; leaves if entire usually broadest below the middle . . . . . . . . . . . . 5
5 a . Lower leaf surface with dense weak patent hairs . . . . . . . . . . . . . 4. P. mollis
b . Lower leaf surface without dense weak patent hairs . . . . . . . . . . . . . . . . . . . 6
6 a . Petioles with rather long yellow hairs . . . . . . . . . . . . . . . . . . . . . 3. P. laevis
. .

b . Petioles with short appressed hairs . . . . . . . . . . . . . . . . . . . . . 5. P. melinonii

I . Pourouma guianensis Aublet, PI. Gui. 2: 892, t. 34 1 . 1 77 5 .

T y p e : Aublet s. n. <;.> , French Guiana (BM).

Pourouma aspera Trecul, A n n . Sci. N a t . B o t . I l l . 8: 1 02. 1 847.

T y p e : Poiteau s.n. <;.> , French Guiana( P).

Trees up to 25 m tall, often with stilt roots. Leafy twigs 3- 1 3 mm thick,

with dense to sparse, brown granular hairs intermixed with sparse to
dense, short, white or yellow, appressed (to patent) hairs. Leaves entire,
(oblong to) subovate to ovate to deltoid, 7- 1 6 cm long, 4. 5- 1 0 cm broad
or 3-lobed to 3-fid up to 5-7-fid to 5-7-parted, 1 3-30(-42) cm long, 1 3-30
(-40) cm broad , chartaceous to subcoriaceous, tip(s) acuminate (or
obtuse), (obtuse to) truncate to deeply cordate at the base; margin entire
to weakly repand ; above short yellow hairs on the main veins, otherwise
scabrous to scabridulous or rarely almost glabrous, beneath with short
268 MORACEAE

yellow appressed or partly more or less patent hairs, between the tertiary
veins arachnoid-tomentellous; veins almost plane above, prominent
beneath, 6-40 pairs of secondary veins; petioles (2-)4-30( -46) cm long
with brown granular hairs intermixed with white to yellowish, appressed
short hairs; stipules 3- 1 3 . 5 cm long, with brown granular hairs inter­
mixed with white to yellow, appressed hairs outside, inside with dense
appressed yellow hairs or almost glabrous.
Staminate inflorescences with dense brown granular hairs intermixed
with sparse to dense, yellow to whitish, appressed or patent hairs on
peduncle and branches; peduncle 2. 5-4 cm long; flowers solitary or
clustered at the end of the branches, usually sessile, sometimes short
pedicellate; perianth l - 1 . 5 mm high, 3-4 parted or segments free, lanceo­
late, acute, puberulous; stamens 4, filaments 0.4-0 . 6 mm long, anthers 0.3-
0.4 x 0 . 3-0.4 mm; a few hairs in the centre of the flower.
Pistillate inflorescences usually distinctly branched, with dense brown
granular hairs intermixed with sparse to dense, yellow to whitish,
appressed to patent hairs on peduncle and branches; peduncle 2-4 cm
long; flowers few to many, pedicel 2-4 mm long; perianth 3-4 mm high,
scabrous to puberulous to velutinous mainly on the lower part; stigma
d iscoid, 1 - 1 . 5 mm in diameter. Fruiting perianth ovoid, ea. 1 5 mm high,
scabrous to sparsely puberulous or subve1utinous, pedicel 4-8 mm long.

D i s t r i b u t i o n : Probably throughout northern South America

and tropical Central America.

In Surinam apparently common; also collected in the young coastal

plain .

BW 2030, 3376, 4033, 4866, 5036, 6292; van Donse1aar 1 262, 1 382, 3333;
F1orschtitz & Maas 3 1 32; Lanj ouw & Lindeman 2 1 1 3; Lindeman 5468, 6 1 23 ,
6 1 66, 6765 ; 01denburger, Norde & Schulz 1 1 20, 1 406; Schulz 83 1 6; Stahe1 Woodherb.
1 23, 1 66, 1 66b.

V e r n a c u l a r n a m e s : bospapaj a (Sur. Dutch), granboesipapaj a

(Sur.), boroma (Arow.) yarayara (Car.), poeroema (Car.).
 MORACEAE 269

The Surinam specimens determined as P. guianensis probably belong to

a widespread complex in which a great number of species have been
described (e.g. P. palmata Poeppig & Endlicher, P. digitata Trecul, P.
radula Benoist, P. scobina Benoist) . There are also distinct resemblances
with P. acutiflora Trecul and P. cecropiaefolia Martius. S ome sterile
specimens from Surinam can hardly be distinguished from the latter
species. P. guianensis is rather variable in Surinam, especially in the
indumentum of the twigs, stipules, inflorescences, and flowers.

2. Pourouma velutina Martius ex Miquel, Fl. Bras. 4( 1 ) : 1 30, t. 4 1 .

1 8 53.

T y p e : Martius s.n. o, Brazil, Para (M).

Trees up to 15 m tall. Leafy twigs 1 . 5-4 mm thick, with sparse brown

granular hairs intermixed with dense, pale yellow or whitish, patent to
appre,ssed hairs. Leaves usually entire, (lanceolate to) oblong to
subovate, 8-25 cm long, 3 . 5- 1 0 cm broad, sometimes 3-lobed to 3-fid, 1 0-
24 cm long, 1 0-25 cm broad , chartaceous, acuminate (to acute) at the
tip(s), obtuse to emarginate at the base; margin repand ; above scabrous,
beneath with dense pale yellow, appressed to patent hairs on the veins,
white arachnoid-tomentose between the tertiary veins; veins plane
above, prominent beneath, 8- 1 3 pairs of secondary veins; petioles 2-9 cm
long, with rather dense pale yellow, appressed to patent hairs; stipules 3 . 5-
7 cm long, outside with rather dense brown granular hairs intermixed
with yellow appressed to patent hairs, inside yellow subsericeous.
Staminate inflorescences patent pubescent to puberulous, with inter­
mixed brown granular hairs on peduncle and branches; peduncle 2. 5-4 . 5
c m long; flowers n o t densely clustered ; perianth e a . I m m high, 4-parted ,
segments elliptic to oblong to lanceolate, puberulous; filaments 0. 5-0. 7
mm long, anthers 0. 3-0.45 x 0.3-0.45 mm.
Pistillate inflorescences poorly to hardly branched, pale yellow to
whitish, appressed pubescent, with intermixed brown granular hairs on
270 MORACEAE

peduncle and branches; peduncle 3-7 long; flowers 4-7(- 1 2), pedicel 0. 5-2
mm long, pubescent to puberulous; perianth ea. 3 mm high, pale yellow
velutinous, with brown granular hairs intermixed , at the tip less densely
hairy; stigma discoid , ea. 2 mm in diameter, often 2-lobed . Fruiting
perianth ovoid, ea. 1 5 mm high, (sub)velutinous, pedicel 2-5 mm long.

D i s t r i b u t i o n : From Surinam to Para ( Brazil).

In Surinam in forests of the interior and the savanna belt.

BW 1 368; Lanj ouw & Lindeman 1 76 1 ; Lindeman 4057, 4794, 64 1 8 .

V e r n a c u l a r n a m e s : bospapaja (Sur. Dutch), granboesipapaj a

(Sur.), boroma (Arow.), yarayara (Car.), poeroema (Car.) .

This species seems to b e closely related t o P. fulginea M iquel (syn . : P.

mildbreadii Standley) from the Upper Amazon Basin.
Although there is no doubt that P. velutina is a good species, it is not
easy to indicate important differences with P. guianensis, to which it
shows most resemblance. As a rule, P. velutina may be distinguished
from P. guianensis by the relatively narrow usually entire leaves, which
have rather few secondary veins and by differences in the indumentum.

3. Pourouma laevis Benoist, Bull. M us. H ist. Nat. Paris 28 : 32 1 . . 1 922.

S y n t y p e s : Sagot 5 1 70, 972c;, Benoist 34 I c;, French Guiana

(P).

Trees up to 30 m tall. Leafy twigs 4- 1 5 mm thick, with dense brown granu­

lar hairs intermixed with sparse to dense, yellow to light brown, patent or
appressed hairs. Leaves 3-lobed to 3-parted, rarely 5-lobed to 5-parted ,
occasionally entire, 1 0-3 1 cm long, 8-28( -40) cm broad, subcoriaceous to
coriaceous, tip(s) acuminate, deeply cordate to obtuse at the base;
margin entire to repand; above with yellow hairs mainly on the main
 MORACEAE 27 1

veins, glabrescent; beneath with sparse to dense, very short and long,
yellow , more or less appressed (sometimes patent) hairs on the veins,
white arachnoid-tomentellous between the tertiary veins; veins nearly
plane to somewhat impressed above, prominent beneath, ( 5-) I 0- 1 6 pairs
of secondary veins; petioles 5-20 cm long, with brown granula r hai rs
intermixed with short and long, patent (to appressed) hairs; stipules 4- 1 4
c m long, outside with brown granular hairs intermixed with yellow,
appressed or patent hairs, inside with yellow appressed hairs.
Staminate inflorescences with brown granular hairs intermixed with
dense rather short, yellow, appressed to more or less patent hairs on
peduncle and branches; peduncle 2-6 cm long; flowers rather densely
clustered, sessile; perianth 1 - 1 . 5 mm high, 4-3-parted (to 4-3-fid),
segments triangular to lanceolate, acute, membranaceous, sparsely
puberulous; stamens 4(-2), filaments 0.6- 1 .0 mm long, anthers 0.4-0.5 x
0.4-0 . 5 mm; a hair tuft in the centre of the flower.
Pistillate inflorescences with brown to light brown, granular hairs
intermixed with white to yellow, more or less appressed hairs on pedun­
cle and branches; peduncle 3 . 5-7(- 1 1 ) cm long; flowers 1 5-30, sessile to
pedicellate, pedicel up to 6 mm long with dense brown granular hairs
intermixed with sparse patent yellowish hairs; perianth ea. 4 mm high,
with dense, brown to purplish, granular hairs intermixed with white
hairs; stigma discoid , ( 1 . 5-)2( -3) mm in diameter. Fruiting perianth
ovoid , ea. 20 mm high with dark brown to dark purplish hairs intermixed
with sparse white hairs, pedicel up to 1 2 mm long.

D i s t r i b u t i o n : From Para ( Brazil) to Surinam, or possibly to Guyana .

In Surinam collected in forests of the interior and the savanna belt.

BBS 297; B W 2095, 3255, 3302, 40 I 0, 5390, 6509; van Donselaar 1 888; lrwin,
Prance, S oderstrom & H o lmgren 55576, 55884; LBB 1 1 0 1 7, 1 4296; Lems
5240 (NA); Schulz 8000; Stahel Woodherb. 1 23b, 1 66a.

V e r n a c u l a r n a m e s : bospapaja (Sur. Dutch) granboesipapaja (Sur.),

boroma (Arow.), yarayara (Car.), poeroema (Car. ).
272 MORACEAE

Fig. 1 4 Pourouma, leaves and pistillate inflorescences; a., P. maroniensis ( Lanjouw

& Lindeman 432); b., P. minor ( P rance et al. 1 0089) .

4. Pourouma mollis Trecul, Ann. Sci. Nat. Bot. I l l . 8: 1 02. 1 847.

L e c t o t y p e : Leprieur l 4 l c;> , French Guiana ( P) .

Trees u p t o 2 5 m tall, with stilt roots. Leafy twigs 3- 1 0( - 1 5) mm thick,

with dark brown to blackish, granular hairs intermixed with more or less
patent, yellow hairs. Leaves 3-lobed to 7-parted, 1 7-50 cm long, 1 4-54 cm
broad , or entire and obovate, ea. 10 cm long, ea. 6 cm broad , subcoria­
ceous, tip(s) shortly acuminate, (acute to) obtuse to emarginate at the
base; margin repand; above with dense yellow hairs on the main veins,
 MORACEAE 273

for the rest with sparse hairs, glabrescent, and brown globose p luricellu­
lar hairs on the secondary and tertiary veins, beneath with weak and
yellow, mainly patent hairs on the veins, white arachnoid-tomentellous
.between the tertiary veins; veins nearly plane above, prominent beneath,
5-35 pairs of secondary veins; petioles 7-20(-37) cm long, with brown
granular hairs intermixed with more or less patent, yellow hairs; stipules
3 - 1 8(-25) cm long, outside with rather few brown granular hairs inter­
mixed with dense, more or less patent, weak and yellow hairs, inside with
· sparse to rather dense, yellow hairs.
Staminate inflorescences with brow n granular hairs intermixed with
patent yellow hairs on peduncle and branches, dense patent yellow hairs
on the terminal branches; peduncle 3-4. 5 cm long; flowers densely glomer­
ate in globose heads and a few solitary flowers, sessile or pedicellate,
pedicel up to 0 . 2 mm long; perianth ea. I mm high, 4-3-lobed, puberu­
lous; stamens 4( -3), filaments 1 . 3- 1 .8 mm long, free or slightly connate at
the base, anthers 0. 5-0 . 5 x 0 . 3 5-0.45 mm; a hair tuft in the centre of the
flower.
Pistillate inflorescences poorly branched, with brown granular hairs
intermixed with dense, appressed to patent, yellow hairs on peduncle and
branches; peduncle 4.5- 1 1 cm long; flowers 1 0-20, mostly clustered in
two groups and turned to one side, pedicel 2-4 mm long; perianth ea. 5
mm high, densely yellow velutinous on the lower part, sparser towards
the tip; stigma d iscoid, 2-3 mm in diameter. Fruiting perianth ovoid , ea.
20 mm high, yellow velutinous, pedicel 2-7 mm long.

D i s t r i b u t i o n : In the Guianas and in all probability in the U pper

Amazon Basin (Brazil and Peru) .

In Surinam in forests of the interior and the savanna belt.

BBS 1 1 28 ; BW 24 1 0 , 4494, 5048, 5440, 558 1 , 5992; van Donse1aar 1 373,

1 68 1 , 2924, 3788; H ostmann (& Kapp1er) 1 272; Lanj ouw & Lindeman 2 1 06;
LBB 9 1 99, 9843; Lems 5262 (NA); Lindeman 363 1 , 3989, 6923; Schu1z 9424.

V e r n a c u l a r n a m e s : bospapaj a (Sur. Dutch), granboesipapaj a (Sur.).

274 MORACEAE

Tn!cul based the description of P. mol/is on three collections : Leprieur

1 4 1 (French Guiana) , H ostmann 1 272 (S urinam) , and Blanchet 236 1
(Para, Brazil) . The latter belongs to P. melinonii, although it differs from
the typical specimens of the Guianas in the occurrence of weak patent
hairs on the lower leaf surface. In this character it agrees with P. mol/is,
to which it is very closely related . The two species can be separated easily
in Surinam, French Guiana, and Amapa (Brazil), owing to the difference
in the indumentum of the lower leaf surface. Moreover, the taxa differ in
the pistillate inflorescences: those of P. melinonii are distinctly branched
and the flowers are not clustered in two groups as in P. mollis. This also
applies to the specimens known from Para (Brazil).

5 . Pourouma melinonii Benoist, Bull. M us. H ist. Nat. Paris 28: 3 1 8. 1 922.

L e c t o t y p e : Sagot 990<;> , French Guiana ( P).

Trees up to 20 m tall, with stilt roots or buttresses. Leafy twigs 3- 1 0 mm

thick, with dark brown to dark purplish, granular hairs intermixed with
sparse white hairs (mainly at the nodes) . Leaves ovate to deltoid , entire to
3-lobed , 9-2 1 cm long, 6- 1 7 cm broad , coriaceous, tip(s) shortly acumi­
nate (to acute), (obtuse to) truncate to emarginate at the base; margin
entire to weakly repand; above sparsely puberulous on the main veins,
and with brown globose pluricellular hairs on the tertiary veins; beneath
with short yellow hairs on the veins, white arachnoid-tomentello u s
between the tertiary veins; veins plane above, prominent beneath, 1 0- 1 4
pairs of secondary veins; petioles 6- 1 7 c m long, with dark brown granular
hairs intermixed with sparse white appressed hairs and sometimes also
with sparse arachnoid hairs; stipules 8- 1 2 cm long, outside with dark
brown to dark purplish, granular hairs intermixed with short, white to
yellow hairs, in side glabrous.
Staminate inflorescences with dark brown to dark purplish granular
hairs intermixed with short, yellow, appressed to patent hairs on pedun­
cle and branches; peduncle 2. 5-3 cm long; flowers densely glomerate in
 MORACEAE 275

globose heads and a few solitary flowers, sessile (or pedicellate); perianth
ea. I mm high, 4( -3)-lobed ; stamens 4( -3), filaments ea. 1 . 5 mm long,
basally connate, anthers ea. 0.4 x 0.4 mm; a hair tuft in the centre of the
flower.
Pistillate inflorescences distinctly branched with dark brown to dark
purplish, granular hairs intermixed with short, yellow to whitish,
appressed to patent hairs on peduncle and branches; peduncle 3-7 cm
long; flowers (4-)8-25, pedicel 2-4 mm long; perianth 4-5 mm high, with
brown to purple, granular hairs intermixed with yellow hairs (dense on
the lower part, sparse near the tip); stigma discoid, 1 . 7-2. 3 mm in diame­
ter, slightly 2-lobed . Fruiting perianth subovoid , ea. 1 8 mm high, yellow
( sub)velutinous, pedicel 6- 1 2 mm long.

D i s t r i b u t i o n : From Bahia (Brazil) through Para and the Guianas

to southern Venezuela.

In S urinam known from the J odensavanne - M apane Creek region

(near the Suriname R iver).

( H ostmann &) Kappler 1 272 (L, P); Lanj ouw & Lindeman 399; LBB 8976.

V e r n a c u l a r n a m e s : bospapaj a (Sur. Dutch), granboesipapaj a

(Sur.), boroma (Arow.), yarayara (Car.), poeroema (Car.).

The original description of P. melinonii was based on the collections

Sagot 5 1 7 and 990, Melinon 457, and Kappler 1 272. Sagot 5 1 7 also
occurs among the syntypes of P. laevis and ( H o stmann &) Kappler 1 272
among those of P. mol/is. I n both cases specimens of different species
have the same number. Specimens from Para and Bahia ( Brazil) differ
from those from Amapa (Brazil) , French Guiana, and Surinam in the
occurrence of patent weak hairs on the lower surface of the leaves. P.
melinonii is closely related to P. mol/is, as already mentioned under the
latter. In Surinam the two species can be separated by the nature of the
indumentum on the lower leaf surface and on inner surface of the
276 MORACEAE

stipules. In the region in which the species occur side by side intermediate
forms have not been found.

Fig. 15 Pourourna, leaves and pistillate inflorescences; a., P . rnelinonii ( l rwin,

Pires & Westra 47832); b., P. mol/is (Elburg ( LBB) 9843).

6. Pourouma maroniensis Benoist, Bull. M us. Hist. Nat. Paris 28: 3 1 8.

1 922.

T y p e : Wachenheim ser. 2. 392o, French Guiana ( P).

Pourourna essequiboensis Standley, Lloydia 2: 1 75. 1 939.

T y p e : A.C. Smith 273 1 0, Guyana ( F, not seen; isotypes K, U).

Trees up to 20 m tall, often with stilt roots. Leafy twigs 2-7 mm thick,
younger parts white arachnoid-villous with intermixed (rather) sparse,
 MORACEAE 277

short yellow appressed hairs, mainly at the nodes. Leaves entire, ovate
to subovate to elliptic, 5- 1 8 cm long, 3- 1 3 cm broad, 3-lobed to 3-
parted, 1 8-22 cm long, 1 8 -23 cm broad , or 5-lobed to 5-parted , 2 1 -40 cm
long, 1 8-20 cm broad, subcoriaceous to coriaceous, tip(s) acute to acumi­
nate, obtuse to emarginate, or (in incised leaves) deeply cordate at the
base ; margin repand ; above short and long, yellow hairs on the main
veins and globose, pluricellular hairs on the secondary veins; beneath
white arachnoid-tomentellous to -villous on the main veins, with inter­
mixed sparse yellow hairs; veins plane to slightly impressed above, promi­
nent beneath, 1 0-26 pairs of secondary veins; petioles 4- 1 5 cm long;
stipules 3- 1 1 cm long, outside with long, yellow, appressed to patent hairs
intermixed with sparse to dense, white arachnoid hairs, inside glabrous.
Staminate inflorescences white arachnoid-tomentose to - villous, with
intermixed short yellow appressed hairs on peduncle and branches;
peduncle 2-3 cm long; flowers densely glomerate, sessile or mostly
pedicellate; perianth 0.7- 1 .0 mm high, 3-4-dentate to 3-4-fid , glabrous or
sparsely puberulous; stamens 4, filaments 1 - 1 . 5 mm long, often basally
connate, anthers 0.4-0 . 6 x 0. 3-0.4 mm; a hair tuft in the centre of the
flower.
Pistillate inflorescences white arachnoid-tomentose to -villous, with
intermixed yellow appressed hairs on peduncle and branches; peduncle 5-
7 cm long; flowers 5-20, pedicel 2-4 mm long; perianth 4-5 mm high�
white arachnoid-tomentose, with intermixed short yellow, more or less
patent hairs, dense on the lower part; stigma discoid, 1 . 5-2.0 mm in
diameter. Fruiting perianth ovoid, ea. 20 mm high, with sparse yellow
hairs and remains of the arachnoid indumentum, pedicel 6- 1 1 cm long.

Distribution : At least in the Guianas and Amapa (Brazil).

In Surinam known from the Brokopondo District and between M oengo

and Albina.

BBS 298; van Donselaar 1 226; Lanjouw & Lindeman 432.

V e r n a c u l a r n a m e s : bospapaja (Sur. Dutch), granboesipapaj a (Sur.),

boroma (Arow.), yarayara (Car.), poeroema (Car.).
278 MORACEAE

The species is closely related to P. acuminata Martius (syn . : P. populi­

folia Standley), a species from the Amazon Basin and the eastern coastal
region of Brazil, and to P. apiculata Benoist and P. tom(mtosa Martius
ex Miquel, both from the U pper Amazon Basin . The differences between
these taxa are found mainly in the indumentum.

f
7. Pourouma minor Benoist, Bull. Mus. H ist. Nat. Paris 30: 1 03 1 924.

T y p e : Benoist 960c;> , French Guiana ( P) .

Pourouma aurea Warburg ex M ildbread, N otizbl. B o t . Gart. Berlin

1 0: 4 1 8 . 1 928.
T y p e : Ule 57 1 8 6, B razil, Amazonas (B, not seen; isotype K).
P ourouma isophlebia S tandley, Pub!. Field M us. Bot. 1 7: 1 82. 1 937.
T y p e : Krukoff 80739, B razil, Amazonas (NY, not seen; isotypes K , U).
P ourouma subplicata Standley, Pub!. Field M us. Bot. 1 7: 1 84. 1 937.
T y p e : Krukoff 5282c;>, Brazil, Acre Territory ( F, not seen; isotypes K , U ) .
P ourouma umbellata Standley, Pub!. Field M us. B o t . 1 7: 1 85 . 1 937.
T y p e : Krukoff 707 1 '( , Brazil, Amazonas ( F, not seen; isotypes K , U ) .

Trees up to 25 m tall. Leafy twigs 1 . 5-3(-5) mm thick, light yellow

sericeous. Leaves entire, obovate to oblong to subobovate to lanceolate,
6-23 cm long, 2-7 . 5 cm broad, often subplicate, chartaceous to
subcoriaceous, acuminate (to obtuse to emarginate), acute at the base;
margin entire to repand ; above yellow sericeous on the costa, brown
globose pluricellular hairs on the secondary and tertiary veins, beneath
yellow sericeous on the veins, between the secondary veins white arach­
noid-tomentellous; veins slightly prominent to plane above, prominent
beneath, 7- 1 8 pairs of secondary veins; petioles 1 . 5-4 cm long, angulate to
broadly canaliculate, yellow sericeous; stipules 3-7(- 1 2) cm l ong, outside
yellow sericeous, inside glabrous.
Staminate inflorescences yellow (sub)sericeous on ped uncle and
branches; peduncle 2-2.5 cm long; flowers not densely clustered , sessile
or mostly pedicellate; perianth 1 . 5-2) mm high, 4-parted, segments
narrow, acute, glabrous o r nearly so; stamens 4( -2) , filaments 0. 6-0 . 8 mm
long, anthers ea. 0.4 x 0.3-0.4 mm; hairs in the centre of the flower.
 MORACEAE 279

Pistillate inflorescences unbranched or nearly so, subumbellate; pedun­

cle 2. 5-6 cm long, yellow (sub)sericeous; flowers 1 -4(- 1 0) , pedicellate,
pedicel 3-6 mm long, yellow sericeous; perianth 5-6 mm high, light yellow
sericeous; stigmas knob-like, 4-5 mm high, 4-5 mm in diameter. Fruiting
perianth ovoid, 2 1 -26 mm high, acuminate, pedicel 1 6-26 mm long.

D i s t r i b u t i o n : I n the Amazon Basin ( B razil, Peru, and nor thern

Bolivia), Amapa (Brazil) , and the G uianas.

In S urinam in forests of the interior.

Lanjouw & Lindeman 2 1 1 4; LBB I 1 027; Schulz 8 1 6 1 ; Stahel Exp. Wilh. Geb. 82.

Ve m a c u l a r n a m e s : bospapaja (Sur. Dutch), granboesipapaja (Sur.).

P. minor is a clear-cut species which can be easily distinguished by the

entire leaves, the yellow sericeous indumentum, the subumbellate few­
flowered pistillate inflorescences, and the knob-like stigma. The speci­
mens from the Amazon Basin d iffer more or less from those of the
Guianas and Amapa in the leaf shape and in the occurence of patent
hairs, mainly on the lower leaf surface. Pluricellular hairs are rare in this
species.

1 3 . Coussapoa Aublet, PI. Gui. 2: 955. 1 775.

L e c t o t y p e s p e c i e s : Coussapoa latifolia Aublet

Dioecious terrestrial or epiphytic trees or shrubs. Leaves spirally

arranged, entire, pinnatinervate (or subpalmatinervate ); stipules
copnate, fully amplexicaul. I nflorescences paired or solitary in the axils
of the leaves, more or less dichotomously branched or (by reduction)
unbranched, with terminal bracteate (sub) �lobose heads. Staminate
flowers with usually 3, more or less connate tepals and one or two
completely connate stamens. Pistillate flowers: perianth tubular, thick-
280 MORACEAE

ened at the upper end ; ova ry free, stigma subpeltate to subpenicillate,

ovule orthotropous, basally attached . Fruiting perianth coloured (at
least at the upper end); fruit small, endocarp crustaceous, exocarp
(often?) mucilaginous; seed with some endosperm, embryo with equal,
� rather flat cotyled ons and a straight apical radicle.

D i s t r i b u t i o n : About 35 species from M exico to southern Brazil.

The species of Coussapoa can be distinguished rather easily.

According to Lindeman (verbal information) in S urinam epiphytic
specimens send down roots which embrace the stems of the host tree
without j oining together and really strangling it, as the strangler figs do.
In specimens with mature fruiting head s examination showed the
outer of the fruitwall to be mucilaginous. It seems likely that the
mucilaginous (dark-coloured?) mass splits open the perianth and pushes
up the endocarp body which is covered by irregular projections and
the remains of the mucilaginous mass. It is not certain whether this
phenomenon is common in Coussapoa, as too few specimens have been
examined . Birds probably play an important role in the dispersal.

a. Leaves scabrous to scabridulous . . . . . . . . . . . . . . . . . . . . . . 4. C . asperifolia

.

b. Leaves smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 a. Leaves (at least the younger ones) arachnoid-villous beneath . . . . . . . . . . . 3
b. Leaves minutely puberulous or glabrous beneath . . . . . . . . . . . . . . . . . . . . 4
.

3 a. Arachnoid indumentum persistent . . . . . . . . . . . . . . . . . . . . 5. C. ferruginea

.

b. Arachnoid indumentum (almost) disappearing . . . . . . . . 2. C. microcephala

4 a. Leaves with 3-6 pairs of secondary veins; pistillate inflorescences repeatedly
branched ; staminate flowers with 2 connate stamens . . . . . . . . I . C . latifolia
b. Leaves with 2-4, mostly 3 pairs of secondary veins; pistillate inflorescences
(usually) unbranched ; staminate flowers with one stamen 3 . C . angustifolia

l . Coussapoa latifolia Aublet, Pl. Gui. 2: 955, t. 362. 1 77 5 .

T y p e : Aublet s . n . 9 , French Guiana ( B M ) .

Coussapoa obovata M iquel, H e t l nstituut 1 842: 200, t. 3. 1 843.

 MORACEAE 28 1

Fig. 1 6 Coussapoa /atifolia, leaves and pistillate inflorescences; from Het Instit-uut
1 842, t. 3.

Coussapoa latifolia Aubl. var. obovata ( M iquel) M iquel, Fl. Bras. 4( 1 ) :

1 35 . 1 853.
T y p e : Focke 99, Surinam (L).
Coussapoa froesii Standley, Pub!. Field M us. Bot. 1 7: 1 62. 1 937.
T y p e : Froes 1 907o , B razil, M aranhao ( N Y , not seen; isotype U).

Shrubs or trees up to 20 m tall. Leafy twigs 1 . 5-4. 5 mm thick, minutely

puberulous to almost glabrous. Leaves subrotundate, elliptic, oblong,
obovate, subobovate or subovate, 4. 5- 1 9. 5 cm long, 2. 5- 1 3 cm broad,
subcoriaceous to coriaceous, emarginate, obtuse or acuminate, at the
base acute to obtuse to subcordate; above glabrous, beneath with curved
pluricellular hairs on the veins; 3-6 pairs of secondary veins;
282 MORACEAE

petioles 0. 5-6 . 5 cm long; stipules 0 . 5- 1 . 7 mm long, minutely puberulous.

Staminate inflorescences repreatedly loosely branched with some tens
of heads each ea. I mm in diameter; peduncle minutely puberulous;
a few minute linear bracts on the branches; perianth ea. 0. 5 mm high,
3-parted , glabrous; stamens 2, completely connate, filament 0 . 6-0. 7 mm
long, anthers 0. 1 -0. 1 5 x 0. 1 -0. 1 5 mm; interfloral bracts subpeltate
to cucullate, shorter or longer than the flowers.
Pistillate inflorescences branched with 3-8 heads 2-3 mm in diameter;
peduncle 0 . 5- 1 . 5 cm long, minutely puberulous; perianth ea. l mm high,
glabrous; a few subpeltate to cucullate interfloral bracts. Fruiting
heads 4-8 mm in diameter; fruiting perianth ea. 2 mm high, colour
unknown .

D i s t r i b u t i o n : From Surinam to Maranhao (Brazil) .

In Surinam as terrestrial or epiphytic plants in forests of the interior.

BBS 3003; BW 732; van Donselaar 2569; van Emden s. n.d, s.n.c;> ; Focke
9 (L), 4 1 8?; Irwin, Prance, Soderstrom & H olmgren 55487; LBB 1 1 747 ; Lindeman
6938; Mennega 27 1 , 52 1 ; Schulz 7787; Stahel Exp. With. Geb. 1 96.

V e r n a c u l a r n a m e : abrasa (Sur.) 1

2. Coussapoa microcephala Trecul, Ann. Sci. Nat. Bot. I l l. 8: 96.

1 847 (August).

Type : Schomburgk 276 (published as 876)d , Guyana ( P).

Coussapoa fagifolia K lotzsch, Linnaea 20: 528 . 1 847 (October).

T y p e : Schomburgk 276 (or 1 366 in B) d , Guyana ( B).
Coussapoa cuneata M iquel, Fl. Bras. 4( 1 ): 1 38 . 1 85 3 .
T y p e : Schomburgk 287d, Guyana ( U ) .

Shrubs or trees u p t o 1 5 m tall. Leafy twigs 2-6 m m thick, brown (to

I The general name for epiphytic trees, including several Ficus and Clusia species.
 MORACEAE 28 3

whitish) arachnoid-villous, with intermixed brown to white stiff hairs ,

glabrescent. Leaves (subrotundate to) elliptic, deltoid, ovate, oblong,
obovate or subobovate, 3 . 5- 1 6 (-27) cm long, 2 . 5- 1 2. 5(- 1 6) cm broad,
coriaceous to subcoriaceous, weakly acuminate, obtuse or acute, at
the base acute to obtuse to cordate; above glabrous, beneath at first
whitish to brown arachnoid-villous, later the indumentum remaining
mainly on the main veins and margin, finally almost glabrous; 4-9 pairs
of secondary veins; petioles 1 -4( -6) cm long; stipules 1 -4.5 cm long,
brown shortly arachnoid-villous, with intermixed whitish to brown
stiff hairs.
Staminate inflorescences repeatedly loosely branched with more than
10 heads each ea. 2 mm in diameter and with ea. 1 0-20 flowers;
peduncle 2-6 cm long, brown (sub)tomentose; perianth ea. 0 . 5 mm
high, 3-parted , glabrous, stamens 2, completely connate, filament ea.
1 mm long, anthers ea. 0.2 x 0.2 mm; interfloral bracts spathulate,
shorter than the flowers.
Pistillate inflorescences repeatedly loosely branched with several
to many heads each ea. 3 mm in diameter and with ea. 1 0-30 flowers;
peduncle 1 .5-3. 5(-7) cm long, brownish (sub)tomentose; perianth ea.
I mm high, glabrous; interfloral bracts spathulate, shorter than the
flowers. Fruiting heads up to 7 mm in diameter; fruiting perianth ea.
2 mm high, scarlet to purple.

D i s t r i b u t i o n : Apparently common m Guyana, probably rare m

Surinam.

In Surinam once collected on the Tafelberg.

Maguire 24588.

3. Coussapoa angustifolia Aublet, PI. Gui. 2: 956, t. 363. 1 775.

T y p e : Aublet s . n . , French Guiana (BM).

284 MORAC EA E
� ----- � -- - ----------

Shrubs or trees up to 1 7 m tall; bark exuding brown sap. Leafy twigs

2-5 mm thick, minutely puberulous to nearly glabrous. Leaves obovate
to oblanceo late , 2.5- 1 1 ( - 1 8) cm long, 1 . 5-5 .5( -7) cm broad , coriaceous,
obtuse or emarginate, acute at the base; glabrous; 2-4, mostly 3 pairs
of secondary veins; petioles 1 -4.5 cm long; stipules 0.5- 1 .0 cm long,
minutely puberulous.
Staminate inflorescences shortly branched with 2-4 separate
conglomerate heads each 3-5 mm in diameter or heads more or less
fused; peduncle 0 . 5- 1 . 5 cm long, puberulous; perianth ea. 0.5 mm high,
3(-5)-lobed , glabrous; one stamen, filament as long as the perianth,
anther O. l -0 . 1 5 x O . l -0 . 1 5 mm; interfloral bracts subpeltate, as high
as the perianths.
Pistillate inflorescences unbranched; heads 3-6( -8?) mm in diameter,
globose, sometimes 2-lobed, occasionally 2 nearly separate heads; pe­
duncle 0.3-2.0 cm long, puberulous; perianth ea . l mm high, glabrous; in­
terfloral bracts subpeltate, as high as the perianth. Fruiting heads 0.5- 1 . 5
cm in diameter; perianth ea. 2; 5 mm high, orange (or yellow?).

D i s t r i b u t i o n : Known from Surinam, French Guiana, and Amapa

( Brazil).

In Surinam collected east of the Coppename River in the interior and in

the savanna belt, as terrestrial and ep iphytic plants.

BW 772; lrwin, Prance , Soderstrom & H olmgren 57574; LBB 1 0344a ; Lindeman
6362, 6545 , 6600; Maguire 23867, 23942; Tresling 55.

V e r n a c u l a r n a m e : abrasa (Sur. ) .

4. Coussapoa asperifolia Tn!cul, A n n . S e . N a t . Bot. I l l . 8: 96. 1 847.

S y n t y p e s : Hostmann l 1 89o , Surinam ( P); Perrotet s . n . ,

French Guiana ( P) .

Shrubs or trees up t o e a . 1 5 m tall. Leafy twigs 2- l O mm thick, minutely

puberulous to scabridulous. Leaves subrotundate, elliptic, obovate, or
 M O R ACEAE 285

subobovate, 6-22 cm long, 3- 1 5 cm broad, chartaceous to subcoriaceous,

emarginate to obtuse to acute or broadly acuminate, at the base
(acute to) obtuse to emarginate; above scabrous, beneath with short white
stiff hairs, weaker white curved hairs, and brown pluricellular hairs;
5-8 pairs of secondary veins, 2 pairs arising from the leaf base;
petioles 1 . 5-8 cm long; stipules 0. 7- 1 . 5 cm long, minutely puberulous to
scab rid ulous.
Staminate inflorescences repeatedly branched to unbranched, with
many globose heads each 2-3 mm in diameter or a few or one, often
I obed head 1 0- 1 5 mm in diameter, respectively; peduncle 2-3 cm long,
covered with short, white, straight or curved hairs and brown pluri­
cellular hairs; perianth l -2 . 5 mm high , 3-lobed , minutely puberulous at
the tip; stamen one, filament as long as the perianth, anther 0. 1 5-0.2 x
0 . 1 5-0.2 mm; iqterfloral bracts many, subpeltate, as high as the flowers.
Pistillate inflorescences (almost) unbranched with one subglobose to
subdiscoid , mostly lobed head 1 -2 . 5 cm in diameter; peduncle 2-3 . 5 cm
long, covered with short, white, straight and curved , hairs and brown
pluricellular hairs; perianth ea. 1 .5 mm high, minutely puberulous at
the tip; interfloral bracts subpeltate, as high as the perianths. Fruiting
heads 1 .5-3 .0 cm in diameter; perianth ea . 2 mm high, colour unknown.

D i s t r i b u t i o n : From Surinam to Amapa (Brazil).

In Surinam probably throughout the country; in forests as terrestrial

or epiphytic plants.

BW 1 322, 2070, 3430, 5262; Hostmann 1 1 89; Lindeman 5709; Rombouts 803;
Splitgerber s.n. (L, P).

V e r n a c u l a r n a m e : abrasa (Sur.).

C.t asper!folia is very closely related to C. magnifolia Trt!cul ( syn.

C. ficina Standley), which ranges from the Amazon Basin to Panama
(cf. Woodson, 1 960); they differ slightly in the pubescence and the
venation .
286 MORACEAE

5 . Coussapoa ferruginea Trecul, Ann. Sci. Nat. Bot. Ill. 8 : 93. 1 847.

T y p e : Leprieur s.n.d , Fr� nch Guiana ( P).

Tree. Leaves oblong to subovate, 4- 1 6 cm long, 2-8 cm broad , coriace­

ous, glabrous above, persistently densely brown arachnoid-villous
beneath; 5-8 pairs of secondary veins. Staminate inflorescences
repeatedly branched with many globose heads each ea. 3 mm in
diameter; flowers with 2 completely connate stamens, filament far ex­
ceeding the perianth.

D i s t r i b u t i o n : The few specimens seen came from Bolivia and

French Guiana, and possibly also from Guyana.

Apart from the type collection, a Schomburgk collection from Guyana

bearing the number 287 has been found in Paris. As a Schomburgk
,
collection of C. microcephala bears the same number and as Trecul did
not refer to this collection it is not quite sure that this species does
indeed occur in Guyana . The species is apparently widespread but rare;
further botanical exploration may lead to its discovery in Surinam.

1 4 . Cecropia Loefling, Iter Hisp. 272. 1 758, nom. conserv.

T y p e s p e c i e s : C. peltata L.

Coilotapalus P. Browne, Hist. Jam. I l l . 1 756, nom. rejic.

Ambaiba Barrere, Ess. hist. nat. Fr. equin. 1 0. 1 74 1 ; Adanson, Fam.
2: 377. 1 763. as synonym; 0. Kuntze, Rev. Gen. PI. 2 : 623, 1 89 1 .

Dioecious trees with hollow partitioned trunks and branches, often with
stilt roots. Leaves spirally arranged, excentrically peltate (to subpeltate),
lobed to divided to the petiole; nerves radiating, basically palmate;
stipules connate, fully amplexicaul. Inflorescences paired or solitary
in .the axils of the leaves, pedunculate, with a digitate cluster of spikes
 MORACEAE 287

in youth enclosed in a spathe. Staminate flowers with a tubular 2-lobed

perianth and 2 free stamens. Pistillate flowers: perianth tubular, with
a pore, thickened at the upper end ; ovary free, stigma sublingulate to
subpeltate, ovule orthotropous, basally attached . Fruiting perianth dry,
fruit small, more or less tuberculate, endocarp crustaceous, exocarp
membranaceous; seed with some endosperm, embryo with equal, rather
flat cotyledons and a straight apical radicle.

Distribution: Throughout tropical America, about 80 species.

The genus is very poorly known and poorly understood taxonomically.

In spite of its common occurence and conspicuous habit, Cecropia is not
well represented in most herbaria and the few specimens collected are
often incomplete and adjusted to the size of a single herbarium sheet.
,
The great number of aggressive ants inhabiting the trees of many species
a nd a uniformity in habit and in habitat have no doubt been the cause
of the slight interest of collectors for this genus.
Cecropia seems to be very succesful in Colombia (see Cuatrecasas,
Rev. Acad . Colomb. 6: 274. 1 945, 9: 325, 1 9 56), but elsewhere there
seems to be a great discrepancy between the number of species described
and the number of species which may prove to be "good".
A great number of names will have to be put into synonymy, but it is
undesirable to make a start with this in the present paper because of the
many problems still extant, the few types seen, and the fact that Dr.
Cuatrecasas is working on a revision of the genus.
The variability of the species, especially at different ages, appears
to be considerable; and in order to understand and interpret this
taxonomically, it will be necessary to have available a large number
of representative collections of many species from many localities and
to have extensive experience of the plants in the field.
In a manuscript treatment of Cecropia prepared by Dr. Snethlage
(about 1 9 20) for publication in the first part of the Flora of Surinam,
seven species were recognized ; two of them were expected to occur
only in Surinam. On the basis of the material seen for the present
288 M O R ACEAE

work the number of species accepted has been red uced to four; one of
these has been recently described ( Berg, Acta Bot. Neerl. 2 1 : 655. 1 972).
Young seedlings of Cecropia have pinnately veined, basally attached,
leaf blades; they soon become palmately veined and then peltate. Adult
specimens have peltate leaves with radiating nerves. Juvenile Cecropia
specimens can usually be recognized by the occurrence of uncinate hairs
- these are lacking in Pourouma.

a. Leaves divided to or to near (at most I cm from) the petiole . . . . . . . . . . . 2

b. Leaves not so deeply divided . . . . . .. .. . . .... ....
. . . . . ....... ..... 3
. . .

2. a. Leaves scabridulous above . . . .

. . . . . . . . . . . . . . . . . . . . . . . . 3. C. silvae
. . . .

b. Leaves smooth above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. C. sciad ophylla

3 a. Leaves arachnoid hairy on the main veins beneath and on the
petiole . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I . C . obtusa
b. Leaves not arachnoid hairy on the main veins beneath and on the
petiole . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. C. surinamensis
. .

l . Cecropia obtusa Trecul, Ann. Sci. Nat. I l l . 8: 79. 1 847.

Ambaiba obtusa (Trecul) 0. Kuntze, Rev. Gen. PI. 2: 624. 1 89 1 .

L e c t o ty p e : Leprieur s.n. (or 1 95) d, French Guiana ( P).

Trees up to (at least) 10 m tall. Leafy twigs 1 0-45 mm thick, scabrous

due to white, curved to uncinate hairs intermixed with brown pluri­
cellular hairs. Leaves subrotundate (occasionally more or less deltoid t9
ovate), 25-50 cm in diameter, 6-, mostly 8-, rarely 9-lobed to -parted,
segments obovate to subobovate to pandurate, always narrowed at the
lower end, usually <;>btuse, sometimes acute to acuminate at the tip,
chartaceous to subcoriaceous; above scabrous, scabridulous or sub­
strigose, brown pluricellular hairs on the main veins, young leaves
more or less densely white arachnoid-villous; beneath white arachnoid­
villous to -tomentellous in the areoles, this indumentum more or less
disappearing from the main veins of older leaves, often brown,
pluricellular hairs on the main veins and on their proximal parts some
curved to uncinate hairs; veins nearly plane above, more or less
 MORACEAE 289

prominent beneath, the costa ' 23-40 cm long with 1 0- 1 5 pairs of stout
secondary veins; petioles 1 6-37(-46) cm long, cylindrical, white
arachnoid-villous, with intermixed brown pluricellular hairs and at the
lower e !]d also more or less scabrous; pulvinus with brown pluricellular
hairs; stipules 7- 1 3 cm long, outside white (rather) sparsely hirsute to
strigose, with intermixed dense pluricellular hairs, inside with white to
yellow hairs.
Staminate inflorescences: peduncle 4-8 . 5 cm long, more or less com­
pressed , broadened at the upper end , scabrous due to white, curved to
uncinate hairs; spikes 1 5-2 1 together, 4- 1 3 cm long, ea. 3 mm thick,
stipitate (stipe 5-1 5 mm long), rachis with stiff, up to I mm long hairs ;
spathe similar to the stipules; perianth 1 . 5-2 mm high, muriculate at the
tip, puberulous below the tip .
Pistillate inflorescences: peduncle 5-7 c m long, more o r less com­
pressed , broadened at the upper end , scabrous due to curved to
uncinate hairs, sometimes also sparsely arachnoid-villous; spikes ( 2-)4
together, 3- 1 0 cm long, ea. 4 mm thick, sessile or shortly stipitate,
rachis with stiff hairs; spathe similar to the stipules; perianth
1 . 5-2 mm high, arachnoid-tomentose below the tip. Fruiting spikes up
to 1 4 cm long, ea . 8 mm thick; fruiting perianth ea. 3 mm high, fruit
2-2.5 mm long, often acute at one end and somewhat tuberculate.

D i s t r i b u t i o n : In the Guianas and in all probability also in the

Amazon Basin.

In Surinam common in open places such as clearings, forest margins,

roadsides, and river banks.

BW 1 56 1 , 1 6 1 3, 1 62 1 , 1 7 1 1 , 1 892, 1 943, 224 1 , 2967, 303 1 , 38 1 8, 3953, 4208, 4708,

5077; Focke 448 ( L); Lanj ouw 453; LBB 8628, 1 2742, 1 2743; Oldenburg, N orde &
Schulz 1 297; Samuels 506 (P); Splitgerber s.n. ( P); Went 1 30; Wullschlligel 1 557
( B R) .

' The midrib o f the midsegment o f the leaf.

290 M O R ACEAE

V e r n a c u l a r n a m e s : bospapaja (Sur. Dutch), boesipapaja (Sur. ),

soro-soro (Car. ), wana-soro (Arow.).

C. obtusa differs distinctly from C. surinamensis in the indumentum of

the lower leaf surface. Moreover, the leaves of C. obtuse usually have
8 segments, whereas those of C. surinamensis mostly have more than 8.
The original description of C. ob tusa was based on four collections
originating from quite different regions of tropical America. It is
convenient to designate the collection Leprieur 1 95 at Paris as the
lectotype.

2. Cecropia surinamensis Miquel, Fl. Bras. 4( 1 ) : 1 43 . 1 853.

Ambaiba surinamensis ( M iquel) 0. Kuntze, Rev. Gen. PI. 2:
624. 1 89 1 .

T y p e : Focke s.n.o , Surinam ( U ) .

Trees u p t o ( a t least) 1 5 m tall. Leafy twigs 0. 5-4 c m thick, scabrous

due to curved to uncinate hairs intermixed with brown pluricellular
hairs. Leaves subrotundate, 1 5-60 cm in diameter, 8- 1 4-, often 9-lobed
to 9-parted , segments (elliptic to oblong to) obovate to subobovate
to pandurate, always narrowed at the lower end , acuminate to obtuse
at the tip, chartaceous (to submembranaceous); above scabrous to
scabridulous, beneath arachnoid-tomentose in the areoles and some­
times also on the parallel veinlets, on the main veins brown pluricellular
hairs and sparse to dense, curved or uncinate hairs; veins plane above,
prominent beneath, the costa 1 3-45 cm long, with 1 0-20 pairs of stout
secondary veins in the upper part; petioles 1 8-67 cm long, cylindrical,
covered with sparse to dense, short curved hairs or longer uncinate
hairs and brown pluricellular hairs; pulvinus with brown pluricellular
hairs; stipules 5 . 5-9 . 5 cm long, outside with brown pluricellular
hairs, rather stiff long white hairs, and / or short curved hairs, not
or sparsely arachnoid-villous, inside with sparse hairs or glabrous.
 MORACEAE 29 1

Staminate inflorescences: peduncle 6- 1 2 cm long, compressed,

broadened at the upper end , slightly scabrous to minutely puberulous,
with intermixed brown pluricellular hairs and long weak hairs mainly on
the upper end; spikes 1 2-24 together, 2-9 cm long, ea. 2 mm thick, sessile
or stipitate (stipe up to 5 mm long), rachis with stiff up to 0 . 5 mm long,
hairs; spathe similar to the stipules or more densely arachnoid­
villous; perianth 1 - 1 .5 mm high, puberulous under the tip; anthers
0 . 3-0.4 x 0.2�.3 mm.
Pistillate inflo rescences: peduncle 8- 1 7 cm long, compressed, .
broadened at the upper end , indumentum like that of the staminate
inflorescences; spikes 4 together, 7- 1 2 cm long, ea. 3 mm thick,
sessile or shortly stipitate, rachis with stiff, up to ea. 0 . 5 mm long
hairs; spathe similar to the stipules or more densely arachnoid-villous;
perianth 1 - 1 . 5 mm high, arachnoid-tomentose below the tip. Fruiting
spikes up to 24 cm long, ea. 5 mm thick ; fruiting perianth 2-2 . 5 mm
high, fruit tuberculate and obtuse at both ends.

D is t r i b u t i o n : At least in the Guianas. This species is probably

widespread; specimens resembling the Surinam collections come from
Mexico to R io de Janeiro (Brazil) .

In Surinam common in wet or dry open places such as clearings, forest

margins, road sides, and river banks.

BBS 4, 223; Focke s . n . ; Kramer & Hekking 23 1 7; Lanj ouw 720, 922; Lanj ouw &
Lindeman 1 28 3 ; LBB 1 062 1 , 1 2739, 1 2750; Oldenburger, Norde & Schulz 1 362;
Pulle H 94; Rombouts 64; Schulz 76 1 6 , 8533, 8622; Splitgerber 1 80 ( L); Tresling I;
Versteeg 4 1 8 ; Wessels Boer 97 1 , 972, 973, 1 256; Wullschliigel 1 556 ( BR), s.n. ( BR).

V e r n a c u l a r n a m e s ; bospapaja (Sur. Dutch), boesipapaja (Sur. )

Regard ing the shape o f the closed spathes and the indumentum o f the
spathes, the nerves of the lower leaf surface, and the petioles, two
forms can be distinguished in the Surinam material: firstly, plants
with usually mucronate to acuminate, white arachnoid spathes and
sparse short curved or longer uncinate hairs on the veins of the lower
292 MORACEAE

leaf surface and on the petioles; secondly, plants with usually acute to
long-acuminate spathes without arachnoid hairs and ( rather) dense short
curved or longer uncinate hairs on the lower leaf surface and on the
petioles. The type of C. surinamensis belongs to the first group.
Intermediate forms exist, so that not all the specimens could be
placed in one or other of the two groups.

3. Cecropia silvae C. C . Berg, Acta Bot. Neerl. 2 1 : 655 . 1 972.

T y p e : S ilva 9569 , Brazil, Para ( U ) .

Tree u p t o 3 0 m tall. Leafy twigs hispid t o hispidulous t o scabrous due

to mainly uncinate hairs of different lengths. Leaves subrotundate to
broadly ovate, 15 (or more than 1 5?) -parted to within 0.5- 1 cm from
the petiole, segments oblanceolate, mostly obtuse, sometimes subacute
to subacuminate, chartaceous to subcoriaceous; above scabridulous
due to short rigid hairs of different length s, sparsely intermixed with longer
weaker hairs and brown pluricellular hairs, beneath arachnoid­
tomentose, this indumentum disappearing from the main veins with age,
on the main veins whitish puberulous to hirtellous, the hairs distinctly
different in length, the longer ones mostly uncinate; the veins
slightly prominent to plane above, prominent beneath, the costa
24-26 cm long, with ea. 35-45 pairs of secondary veins, at most 0 . 5 cm
from each other; petioles 35-47 cm long, cylindrical arachnoid­
tomentellous and sparsely to rather densely, hirtellous, but hirsute to
hispid at the upper and lower ends; pulvinus with brown pluricellular
hairs and longer white 'unicellular ones.
Pistillate inflorescence with a 6-7 cm long, hirtellous to scabrous
and towards the apex hirsute to hispid peduncle; spikes 4 together,
8-9 cm long, ea. 9 mm thick, sessile, rachis with up to 2 mm long
stiff white hairs; perianth ea. 3 mm high, towards the tip densely
arachnoid-tomentose.

D i s t r i b u t i o n : Only known from northern Pani ( Brazil) and Surinam.

 MORACEAE 293

In Surinam once collected in the Sipaliwini savanna area.

Oldenburger, Norde & Schulz 1 1 1 2.

4. Cecropia sciadophylla Martius, Flora ( od. Bot. Zeit.) 24 ( Beibl. 2) : 93.

1 84 1 .
Ambaiba sciadophylla ( Martius) 0 . Kuntze, Rev. Gen. PI. 2:
624. 1 89 1 .

T y p e : Martius 630 ( 1 )9 , Brazil ( M ) .

Trees u p to 30 m tall, often with stilt roots. Leafy twigs 1 . 5-5 cm thick,
covered with a few weak long white hairs, glabrescent. Leaves sub­
rotundate, 22-70 cm in diameter, 1 1 - 1 5 parted to or nearly to the petiole,
segments oblanceolate, undulate, sometimes short petiolulate, (acute to)
acuminate, often slightly plicate, subcoriaceous to coriaceous; above
glabrous and shining, beneath white arachnoid-tomentellous in the
areoles, sometimes also on the parallel veinlets, the main veins glabrous
or with a few weak white hairs and few or many, brown pluricellular
hairs; veins almost plane above, more or less prominent beneath, the
costa 1 4-45 cm long, with 22-38 pairs of stout secondary veins; petioles
35-70 cm long, cylindrical to subtrigonous, almost glabrous; pulvinus
lacking; stipules 30-36 cm long, outside sparsely puberulous with
many brown pluricellular hairs intermixed, inside with appressed
long white hairs, sometimes mainly on the 2 stout longitudinal veins.
S taminate inflorescences: peduncle 7- 1 5 cm long, compressed to
angulate, greatly broadened at the upper end , minutely puberulous with
many brown pluricellular hairs intermixed; spikes 8- 1 3 together,
8- 1 2 cm long, 4-8 mm thick, stipitate ( stipe 1 . 5-3 mm long), rachis with
stiff, up to ea . 0.5 mm long hairs; spathe similar to the stipules;
perianth ea. I mm high, muriculate at the tip, white arachnoid-tomen­
tose below the tip; anthers 0. 5-0.6 x ea. 0 . 2 mm.
Pistillate inflorescences: peduncle 3-8 cm long, compressed to
angulate, broadened at the upper end, minutely puberulous, with many
294 MORACEAE

brown pluricellular hairs intermixed; spikes 4-6 together, 8- 1 2 cm long,

7-8 mm thick, sessile or subsessile, rachis with stiff, up to 0. 5 mm long
hairs; spathe similar to the stipules; perianth ea. 2 mm high, muriculate
at the tip, white arachnoid-tomentose below the tip. Fruiting spikes up to
20 cm long, 1 0- 1 5 mm thick; fruiting perianth ea. 3-3 . 5 mm high, fruit
2.5-3 mm long, (sub)acute at one end , tuberculate.

D i s t r i b u t i o n : In the Amazon Basin and the Guianas.

In Surinam in forests; not collected in the young coastal plain.

BW 1 593, 3836, s.n.; lrwin, Prance, Soderstrom & Holmgren 55577; Lanjouw
824; Lanj ouw & Lindeman 2485 ; LBB 1 0 1 09, 1 2744, 1 2745, 1 2746, 1 2747, 1 279 1 ;
Lindeman 36 1 7; Stahel Woodherb. 1 79.

V e r n a c u l a r n a m e s : bospapaja (Sur. Dutch), manbospapaja (Sur.)

soro-soro (Car.), wana-soro (Arow.).

Several infraspecific taxa have been distinguished within C. sciadophylla

(see Snethlage, N otizbl. B ot. Gart. B erlin 8 : 358; 1 923; Cuatrecasas
Rev. Acad. Colomb. 6 : 299. 1 945) . S nethlage distinghuised var.
juruyiana (A. Richter) Snethlage, characterized by non-petiolulate
leaf segments, and var. decurrens Snethlage, characterized by shortly
petiolulate leaf segments. The Surinam specimens have leaves with short­
ly petiolulate and non-petiolulate segments. H owever, both forms appear
to pass into each other gradually, so that the distinction of two infra­
specific taxa proves not to be justified for Surinam. Specimens collected
in the Amazon Basin (including the type of C. sciadophyl/a) have leaves
with long petiolulate segments and in this leaf character are distinct from
collections made in the Guianas.
* A fifth species of Cecropia probably also occurs in Surinam: C. palmara

Willdenow, which is characterized by reddish and subpersistent stipules,

leaves with 7- 1 1 obtuse segments, staminate inflorescences with 4- 1 1
ea. l- 1 . 5 cm thick spikes, and pistillate inflore scences with 4 ea. 1 cm
thick spikes. The species is common along the Lower Amazon River.
 MORACEAE 295

MORACEAE INTRODUCED AND C U LTIVATED IN SU RINAM

Most of the species mentioned below have been discussed by Stahel

( 1 944) 1 and Ostendorf ( 1 962)2 in their works dealing with the useful
plants of Surinam.

Antiaris

Antiaris toxicaria Lesch . var. macrophylla ( R . Br.) Corner

Probably a single tree (still?) cultivated in the Experimental Garden at

Paramaribo .

Artocarpus

Artocarpus communis J. R. & G. Forster ( Breadfruit)

Artocarpus incisus L.

Monoecious trees up to 35 m tall. Leaves spirally arranged, elliptic, up

to ea. 50 cm long and up to ea. 20 cm broad , pinnatifid; stipules 1 0-25 cm
long, leaving annular scars. Inflorescences in the leaf axils, the staminate
ones cylindric to clavate, 1 0-30 cm long, pedunculate, the pistillate ones
ellipsoid to globose, pedunculate, the upper part of the perianths conical
and pointed. I nfructescences ellipsoid to globose, 1 5-30 cm in diameter,
covered by conical pointed processes, but without these in seedless
forms.

V e r n a c u l a r n a m e s : seeded form - kastanj ebroodboom (Sur. Dutch),

sinibredebon (Sur.); seedless form - njamsi-bredebon (Sur.).

1Stahel,. G . , 1944. De nuttige planten van Suriname. ed . 2.

2Ostendorf, F. W . , 1 962. Nuttige planten en sierplanten in Suriname. Bulletin
Landbouwproefstation in Suriname no. 79.
296 MORACEAE

Both the seeded and seed less form are cultivated .

Artocarpus heterophyllus Lam . (Jack, Jackfruit)

Artocarpus integrifolia and Artocarpus integer auct. mult.

M onoecious trees up to 10 mm tall. Leaves spirally arranged, elliptic to

obovate, up to ea. 25 cm long and up to ea. I 0 cm broad, entire; stipules
1 . 5-8 cm long, leaving annular scars. Inflorescences mostly on the stem
and branches, the staminate ones clavate, up to I 0 cm long, and peduncu­
late, the pistillate ones ellipsoid to globose, pedunculate. Infructescences
ellipsoid to cylindric, 30-60 ( -1 00) cm long, covered by conical tapering
processes.

Both the form with firm fruiting perianths and that with soft fruiting
perianths are cultivated .

Castilla

Castilla elastica Sesse ssp . elastica

Castilloa elastica auct.

M onoecious to dioecious trees up to 30 m tall, with more or less droop­

ing, partly self-pruning, branches. Leaves alternate and distichous on the
twigs, pendulous, oblong, 1 5-40 cm long, 7-20 cm broad, hairy, emargi­
nate to cordate at the base; stipules fused , 2- 1 0 cm long, leaving annular
scars. Staminate inflorescences flabellate and 2-valved, 1 0-25 mm broad,
involucrate, ped unculate, those accompanying pistillate ones often smal­
ler and cup-shaped to funnel-shaped; pistillate inflorescences discoid, 1 -2
cm in diameter, involucrate, sessile, with 1 5-30 flowers. I nfructescences
2.5-4.5 cm in diameter, fruiting perianth orange to red .

Introduced for rubber production . Nowadays occasionally found as

an ornamental tree or run wild at plantations.
 MORACEAE 297

Ficus1

A number of species of Ficus exist in a cultivated or quasi-cultivated

state. In regard to the latter cond ition, plants of various native species
of Ficus are frequently left standing when land is cleared for pastures,
apparently to afford shade for the cattle. They are also allowed to
survive along roadsides and in public squares to provide shade.
Finally, a few species, notably Ficus citrifolia, furnish "living fence
posts". Freshly cut stakes, with the bark intact, are driven into the
ground, soon root and put forth branches and grow to form good-sized
trees.
The following species of Ficus have been recorded as cultivated in
Surinam. It may be ' expected that other species will be recorded, as
interest turns to those species that accompany man.

Ficus benjamina L. ( Weeping Fig.)

Tree to 35 m tall, typically with a single trunk and several widely

spreading branches, with the branchlets d rooping (weeping) at the tips.
Aerial roots rare or absent. Leaves ovate to elliptic, to 1 0 . 5 cm long x 4.3
cm wide, the base rounded , tip acuminate. Figs sessile, depressed
globose to obovoid , to 1 1 mm tall x 8 mm in diameter, red to blackish
at maturity in the typical form, or to 2 1 mm tall x 1 8 mm in diameter,
yellow at maturity in var. nuda ( Miq.) Bartlett ( var. comosa ( R oxb.)
=

Kurz).

Ficus carica L. ( Edible Fig)

A large shrub or small tree with coarse twigs. Aerial roots absent.
Leaves orbicular to ovate, generally 3-5- lobed or -lobulate , to 3 3 cm

I by G.P. DeWolf
298 MORACEAE

long x 25 cm wide, the base cordate to cuneate, tip blunt. Figs with a
peduncle 2- 1 5 mm long, globose to obovate, to 8 cm long, yellowish
green to brownish purp le when ripe.

Ficus elastica Roxb. ex Hornem. ( Rubber Plant)

Tree to 30 m tall with a massive trunk and ascending branches. Aerial

roots produced . Leaves elliptic to oval, to l l cm long x 5 . 6 cm wide on
mature plants, to 30 cm long x 17 cm wide on sucker shoots or young
plants, base rounded , tip briefly acuminate. Elongating twigs with
large, quickly deciduous rosy pink stipules around the terminal bud .
Figs sessile or nearly so , oblong, 1 0 mm tall, greenish yellow when ripe.

Ficus kurzii King

Tree to 50 m tall, the major branches with conspicuous aerial roots

which may reach the ground and form false stems. Leaves elliptic or
ovate-oblong, to l l cm long x 5 cm wide, base rounded, apex short acu­
minate. Figs with the base narrowed to a stalk above the 3 bracts at the
base, globose, to 20 mm tall x 1 5 mm in diameter, dark purple to black at
maturity.

Ficus microcarpa L.f. (Indian Laurel)

Ficus retusa and Ficus nitida auct. mutt.

Tree to 30 m tall with spreading or ascending branches. Aerial roots

abundantly produced, some reaching the ground and forming false
stems. Leaves elliptic to oblong -obovate, to 10 cm long x 4. 7 cm wide,
the base rounded, tip acuminate. Figs sessile, globose or depressed glo­
bose, to 10 mm in diameter, bluish black at maturity .
 MORACEAE 299

Ficus pumila L. (Climbing Fig)

Creeping or climbing shrub, attaching its branches to the substrate by

clusters of very short adhesive roots. Leaves with a conspicuous dimor­
phism: on the lower (vegetative) portion of the plant ovate to ovate­
oblong, up to 3 . 5 cm long x 2.3 cm wide (frequently smaller) , the base cor­
date, tip blunt; on fruiting branches to 8 . 5 cm long x 4. 5 cm wide, the base
rounded to subcordate, tip blunt to acute. Figs with a peduncle to 1 8 mm
long, campanulate or conical, to 75 mm long x 65 mm in diameter, dark
blue or purple when ripe.

Ficus religiosa L. (Bo Tree, Peepul)

Tree to 25 mm tall. Aerial roots absent . Leaves broadly ovate, to 23 cm

long x 12 cm wide, base truncate to cordate, tip profoundly acuminate,
the acumen to l l cm long (about 12 the length of the lamina). Figs
sessile, depressed globose, to 1 5 mm in diameter, blackish at maturity.
 URTICACEAE

BY

M .J . M . DE ROOIJ

Herbs, shrubs or trees, monoecious or dioecious, sometimes armed

with stinging hairs or prickles, usually with cystoliths in the epidermal
cells. Leaves simple, alternate or opposite; stipules 2, free or connate .
Inflorescences uni- or bisexual, basically dichasial, by reduction of axes
often resembling spikes, cymes, head s, panicles or clusters; flowers
small, unisexual, rarely bisexual; perianth sometimes wanting. Staminate
flowers with a (2-)4-5-parted perianth; stamens (2-)4-5, epitepal, inflexed
before anthesis; pistillode usually present. Pistillate flowers mostly with
(2-)3-5 free or fused tepals, perianth sometimes wanting; ovary one,
superior, unilocular; style undivided ; stigma penicillate-capitate or fili­
form; ovule one, orthotropous, (sub)basal. Fruit an achene, sometimes
drupaceous, sometimes enveloped by an enlarged (fleshy) perianth.

D i s t r i b u t i o n : About 40 genera with more than 700 species in the tro­

pics and temperate zones of both hemispheres.

I n Urticaceae the caracters of the pistillate flowers are important for

recognation and identification, in contrast, the staminate flowers are
very similar throughout the family.

LITERATU R E CITED

CHEW WEE-LEK, 1 965. Laportea and allied genera ( U rticaceae). Gard . Bull.
Singapore 2 1 : 1 95-208.
-, 1 969. A monograph of Laportea ( U rticaceae). Gard . Bull. Singapore 25: 1 1 1 - 1 78.
KILLIP, E . P . , 1 9 60. Flora of Panama. U rticaceae. Ann. M issouri Bot. Gard . 47:
'
1 79- 1 98.
MIQUEL, F.A.W., 1 853. U rticineae. In Martius Flora Brasiliensis 4( 1 ) : 76-2 1 8.
 U RTICACEAE 30 1

W E D D E L L, H . A . , 1 856- 1 857. M onographie de la famille des U rticees. Arch.

Mus. H ist. Nat. Paris 9.

KEY TO T H E G E N E R A

a. P lants with stinging or irritant hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

b. P lants without these hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 a. S h rubs or trees, 1 - 1 0 m high; inflorescences unisexual; stigma penicillate-
capitate or linear and straight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I . U rera
.

b. Annual herbs up to 2 m high; inflorescences usually bisexual; stigma linear

and reflexed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. Laportea aestuans
3 a. Leaves opposite; pistillate perianth uneq ual 3-parted; staminate flowers
4-parted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. Pilea
b . Leaves alternate; pistillate perianth tubular, contracted at the throat; staminate
flo wers 3-parted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. Boehmeria ramiflora

I . Urera Gaudichaud in Freycinet Voy. Bot. 496. 1 826.

T y p e s p e c i e s : U rera baccifera (L.) Gaudichaud ex Weddell

Calostima Rafinesque, Fl. Tell. 3 : 47. 1 836.

T y p e s p e c i e s : Calostima aculeata Rafinesque (= U rera baccifera).

Shrubs or small trees, usually dioecious, with herbaceous to ligneous

stems and branches, usually with stinging hairs or prickles. Leaves sim­
ple, alternate, triplinerved with 5- 1 0 pairs of secondary veins; stipules
co nnate, more or less free at the top. I nflorescences unisexual, bracte­
ate, dichotomously to more or less irregularly branched; flowers sessile
to pedicellate, solitary or in glomerules. Staminate flowers with a 4-5-par­
ted perianth and with 4-5 stamens; pistillode membranaceous. Pistillate
flowers with a 4-fid to 4-parted perianth; tepals unequal or equal; stigma
pennicillate-capitate, sessile, persistent. Achene lens-shaped, oblique.

D i s t r i b u t i o n : About 30 species mainly in tropical America and

Africa.
The neotropical species of U rera are not well differentiated and are
therefore not easily separated. Revision of the genus is necessary.
302 U RTICACEAE

a. Leaves pinnati-lobed to -parted . . . . . . . . . . . . . . . . . . . . . . . . . . 3 . l l . laciniata

b. Leaves entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.
2 a. Leaf margin coarsely dentate to sinuate-dentat�:; branches usually with stout
prickles with an up to I cm broadened base; inflorescences sub-dichotomously
to irregularly branched. flowers usually solitary, sometimes 2-4 together;
fruits 3-5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I . U. baccifera
b. Leaf margin crenate to dentato-crenate; plant without prickles with a
broadened base; inflorescences repeatedly dichotomously branched , flowers
up to ea. 50 in compact glomerules; fruits 1 - 1 . 5 mm long . . . . 2. U. caracasana

I . Urera baccifera ( L. ) Gaudichaud ex Weddell, Ann. Sci. N at. Bot. I l l .

1 8: 1 99 . 1 852.
U rtica baccifera L. , Sp. PI. (ed. 2) 1 398. 1 763.
Calostima aculeata Rafinesque, Fl. Tell. 3 : 48. 1 836; as a
synonym.

T y p e : Plumier, Plant. Amer. 1 0: t. 260. 1 760.

U rtica horrida H . B . K . , Nov. Gen. et Sp. 2: 33. 1 8 1 7.

U rera horrida ( H . B. K . ) M iquel, Fl. Bras. 4( 1 ): 1 92. 1 853.
U rera baccifera var. horrida ( H . B. K . ) Weddell, in DC. Prod. 1 6( 1 ): 94. 1 869.
T y p e : Humboldt & B onpland 1 639, Colombia, Santander ( P) .
U rtica armigera Presl, B o t . Be m erk. 1 1 0. 1 844.
U rera armigera ( P resl) M iquel, Fl. Bras. 4( 1 ) : 1 92. 1 853.
T y p e : Lhotsky s . n . , Brazil, near R io de Janeiro ( PR ) .
U rtica nitida Vellozo, Fl. F l u m . 1 0 : t. 2 0 . 1 827.
T y p e : Fl. Flum. 10: t. 20. 1 827.
U rera denticulata M iquel, Fl. Bras. 4( 1 ): 1 92. 1 853.
T y p e : Martius s . n . , Brazil, M inas Geraes ( M ) .
U rtica grand identata Liebmann, Kg!. Danske Vidensk. Selsk. Skr. V . 2 :
296. 1 85 1 .
T y p e : O rsted 1 4283, Costa R ica (lrasu) (C).
U rera baccifera var. angustifolia Weddell, in DC. Prod . 1 6( 1 ) : 94. 1 869.
Based on and as a synonym of U rera denticulate M iq uel, U rera armigera
M iquel, U rtica armigera Presl and U rtica grandidentata L iebmann

Shrubs, usually dioecious, rarely monoecious, up to 3 m, sometimes up

to 6 m high, with erect subligneous to herbaceous pithy aculeate stems
and branches, young growths hirtellous. Leaves broadly to narrowly
 U RTICACEAE 303

0 b

(8
a

t3
c

e
,'[1
Fig. I Urticaceae, pistillate flowers; a . , Urera baccifera; b . , Urera caracasana;
c.,Urera laciniata; d . , Laportea aestuans; e., Boehmeria ramiflora; f. , Pi/ea
pubescens.

ovate to elliptic or oblong, or sometimes to suborbiculate, 1 2-3 1 cm long,

8-26 cm broad , chartaceous; apex acuminate to acute; base obtuse, trun­
cate or emarginate to subcordate; margin coarsely dentate to sinuate-den­
tate; lamina above almost glabrous, usually scabridulous, beneath ap-
304 U RTICACEAE

pressed pubescent to substrigillose, with ret rorse stinging prickles on the

costa and the secondary veins; triplinerved ; veins plane above, promin­
ent beneath, 6- 1 0 pairs of secondary veins, many parallel tertiary veins;
petioles 3-22 cm long, hirtellous, usually with stinging prickles; stipules 6-
1 2( -25) mm long, connate except for their upper parts, puberulous.
Inflorescences unisexual, up to 9 cm long, usually repeatedly subdi­
chotomously branched, tending to form a main axis, bracteate, with re­
trorse stinging hairs and numerous reddish globose-capitate pluricellular
hairs; interfloral bracts present . Staminate flowers sessile or subsessile,
usually solitary, sometimes 2-4 together; perianth 1 - 1 . 5 mm high, ( 4-)5-
parted , tepals subcucullate; stamens ( 4-)5; pistillode small, membranace­
ous. Pistillate flowers with an up to 0 . 2-2 mm long pedicel, usually soli­
tary; perianth 4-fid with unequel segments, not overlapping each other;
receptaculum flat. Achene lens-shaped , oblique and apiculate, ea. 3-5
mm long, the basal part enveloped by the up to 3 mm high fleshy peri­
anth.

D is t r i b u t i o n : Widely distributed in tropical America.

In Surinam in the interior; most collections from the Wilhelmina Range,

Emma Range, and Tafelberg, also one collection from the Kabalebo
River near the Bakhuis Mountains.

BW 6976; Geyskes 13; Florschtitz & Maas 2524; Gonggrijp & Stahel 1 78; I rwin,
Prance, Soderstrom & H olmgren 54666, 54856; Schulz ( L BB) 1 0266; Wessels Boer
1 573.

2. Urera caracasana (Jacquin) G risebach, Fl. Brit. Br. W. l nd . 1 54. 1 856.

Urtica caracasana Jacquin, H ort. Schoenbr. 3 : 7 1 , t. 386. 1 798.
U rera jacquinii Weddell, Ann. Sci. Nat. Bot. I l l . 1 8 : 200. 1 852.

T y p e : Jacquin, H ort. Schoenbr. 3 : t. 386. 1 798.

Urtica alceaefolia Poiret in Lamarck, Encycl. Suppl. 4: 2 2 7 . 1 8 1 6.

U rera alceaefolia Gaudichaud in Freycinet Voy. Bot. 497. 1 826, cited as a
synonym by Weddell, in DC. Prod. 1 6( 1 ): 89. 1 869.
T y p e : Martin s.n., French Guiana ( P).
 U RTICACEAE 305

Fig . 2 Vrera, leaves and pistillate inflorescences; a., V. baccifera (Geijskes 13 and
Schulz .(LBB) 1 0266 respectively); b . , V caracasana (A. C. Smith 3 505 and Fanshawe
( FD) 48 1 9 respectively); c., V. /aciniata ( Lindeman 5660 and M exia 8326 respectively).
306 U RTICACEAE

U rtica ti liaefolia H. B. K . , Nov. Gen. et S p . 2: 34. 1 8 1 7 .

T y p e : H umboldt & Bonpland 1 6 3 3 , Colombia ( P ) .
U rtica ulm ifolia H . B. K . , N ov . G e n . et S p . 2: 3 4 . 1 8 1 7 .
U rtica jacquinnii var. ulm ifolia ( H . B. K . ) Weddell. Arch. M us . H i st . Nat.
Paris 9: 145. 1 8 56.
T y p e : H u mboldt & Bonpland 1 427, Colombia, Bolivar ( P ) .
U rera subpeltata M iquel, Fl. Bras. 4( 1 ): 1 89 . 1 8 5 3 .
T y p e : Martius s . n . , Brazil, Bahia ( M ).
U rera jaquinii var. subpeltata ( M iquel) Weddell, Arch. M us . H i st. Nat.
Paris 9: 1 45 . 1 856.
U rera caracasana va r. subpeltata ( M iquel) Weddell, in D . C. Prod. 1 6( 1 ) : 90.
1 869.
Based on and as a synonym of U rera subpeltata M iquel and U rtica tiliaefolia
H . B. K.
Urera subpeltata var. mo rifolia M iquel, Fl. Bras. 4( 1 ) : 1 90. 1 8 53.
S y n t y p e s M a rt iu s s . n . ( o r 25?), Brazil, Amazonas ( M ); R . Spruce 633,
Brazil, Para ( M ) .
U rtica mitis Vellozo, Fl. F l u m . 1 0 : t . 1 9 . 1 827.
U rera mitis ( Vellozo)M iquel, Fl. Bras. 4( 1 ) : 1 9 1 . 1 853.
T y p e : Vellozo, F l. Flum. 1 0 : t . 19. 1 827.
•u rera caracasana var. mitis (Vellozo) Weddell, in DC. Prod. 1 6( 1 ): 90. 1 869.
Based on and as a synonym of U rera mitis Miquel, Urera jaquinii var.
ulmifolia Weddell, and U rtica ulmifolia H. B. K .
U rera acuminata Miquel, F 1 . Bras 4( 1 ): 1 90. 1 85 3 .
U rera j acquinii var. miquelii Wed cfell, Arch. M u s . Hist. N a t . Paris 9 : 1 45 .
1 856.
Urera caracasana var. miquelii ( Weddell) Weddell, in DC. Prod. 1 6( 1 ) : 90.
1 869 .
T y p e : Mikan s . n . , Brazil, Rio de Janeiro (not yet traced).
Urera acuminata Miquel, F1. Bras 4( 1 ): 1 90. 1 85 3 .
T y p e : P o h l s.n. Brazil, pro b . R i o de Janeiro ( n o t yet traced).
Urera jacquinii var. miquelii Weddell, Arch. Mus. H ist. Nat. Paris 9: 1 4 5 .
1 856.
Urera caracasana var. tomentosa ( Weddell) Weddell, in DC. Prod. 1 6( 1 ): 90.
1 869.
Based on and as a synonym of U rera densiflora M iquel and U rtica verrucosa
Liebmann.
U rtica verrucosa Liebmann, Kg!. Danske Videns k . Selsk. S kr. V. 2: 295.
1 85 1 .
T y p e : O rsted 1 4284, Costa �ica (lrasti) (C).
Urtica corallina Liebmann, Kg!. Danske Vidensk. Sels k . Skr. V . 2: 295. 1 85 1 .
T y p e : O rsted 1 4282, Costa Rica (Aguacate) (C).

Dioecious shrubs or small trees, up to 5 m, sometimes up to 10 m, sublig­

neous to ligneous, young growths hirtellous to strigillose. Leaves broad ly
 U RTICACEAE 307

to narrowly ovate, 1 2-36 cm long, 8-26 cm broad, chartaceous; apex acu­

minate; base cordate to emarginate to obtuse, sometimes with overlap­
ping lobes; margin crenate to dentato-crenate; lamina above sparsely stri­
gillose and usua lly scab rid ulous, beneath substrigillose to subhirtellous;
veins plane above, pro minent beneath ; 5-9 pairs of secondary veins; peti­
oles 2.5-22 cm long, hirtellous to strigillose; stipules 5- 1 4 mm long, pube­
ru lous.
Inflorescences unisexual, repeatedly dichotomously branched, bracte­
ate, with greatly shortened terminal axes, hence with up to 50 flowers in .
compact glomerules, axes puberulous to hirtellous, also with retrorse
stinging hairs. Staminate flowers sessile; perianth 4( -5)parted , ea . I mm
high and 1 . 5-2 mm broad; tepals subcucullate; pistillode membranace­
ous. Pistillate flowers (sub )sessile; perianth up to 0 . 5 mm high; tepals 4, al­
most free, the outer ones smaller than the inner ones. Achene lens­
shaped, ea. l - 1 . 5 mm high, the basal part enveloped by the up to I m m
high perianth .

D i s t r i b u t i o n : Widely distributed in tropical America.

In Surinam known from two collections, one obtained in the Bakhuis

M ountains and one from near Paramaribo .

Florschtitz & Maas 2824; Splitgerber 38 1 (L).

I n the glomerules of the male inflorescences ea. 1 . 5 mm long stipes with a

peltate apex may occur among the staminate flowers. According to
Miquel ( 1 853) these stipes are deflo rated pedicels. In some cases the
terminal axes of the female inflorescences are extremely swollen and
fleshy. These aberrant inflorescences may occur together with normal
inflorescences.
Urera caracasana appears to be highly variable in many of its charac­
ters; infraspecific taxa can probably be distinguished (cf. Killip, 1 960).
308 U RTICACEAE
------ · -------- - -- - - - ---------

3. Urera laciniata Weddell, Ann. Sci. Nat. I l l . 1 8 : 203 . 1 852.

Urtica laciniata Goudot mss . , cited as a synonym by Weddell,
Ann. Sci. Nat. Bot. I l l . 1 8 : 203 1 8 52.

L e c t o t y p e : Goudot s . n . , Colombia ( P).

U rea girardi nioides Seemann, Bot. Voy. Herald 1 94. 1 854.

T u p e : Seemann 494, Panama, Canal Zone (BM).

Perennial herbs or shrubs with up to 7 mm long stinging prickles on the

leaves and stems, probably dioecious, up to 4 m high. Leaves pinnati-lo­
bed to -parted, 20-23 cm long. 20-30 cm broad segments dentate to
lobed, base cordate to truncate; lamina above glabrous or with very
sparse appressed hairs at the base,_ beneath gla brous or with sparse
appressed hairs, the mid rib and the secundary ve ins usually wit h
prickles; 5-7 pairs of secondary veins; petiole ea. 1 5 cm long with very
short patent hairs. Stipules inconspicuous.
Inflorescence unisexual, more or less paniculate, ea . 1 0-20 cm long
and ea. 7- 1 2 cm broad , with hirtellous branches; peduncular and inter­
floral bracts present. Staminate flowers with a 0 . 8(- 1 ) mm long
pedicel; perianth I mm high and 2 mm broad , 5-parted; tepals sub­
cucullate, with reddish globose-capitate pluricellular hairs; pistillode
membranaceous. Pistillate flowers ( sessile to) short pedicellate, ea .
2 mm high and 0.7 mm broad ; perianth I mm high, hairy, 4-parted ,
tep'als acute, the outer ones much smaller than the inner ones, ciliate
with globose-capitate pluricellular hairs; stigma I mm long, subulate
with long and dense papillae; ovary compressed-ovoid to -ellipsoid,
at the base fused with the perianth . Achene sessile or very short stipitate,
2 mm high and 2 mm broad, compressed ; stigma persistent, oblique;
perianth persistent, the 2 larger tepals enlarged up to 1 . 6 mm high , the
smaller ones not or hardly enlarged .

D i s t r i b u t i o n : From Costa Rica to Bolivia and Venezuela, also in

Surinam.
 U RTICACEAE 309

In Surinam known from one locality near the mouth of the

Coppename River.
Lindeman 5660.

2. Laportea Gaud ichaud in Freycinet Voy. Bot. 498. 1 826, nom. conserv.

T y p e s p e c i e s 1 : Laportea canadensis ( L. ) Weddell

U rticastrum Heister ex Fabricius, En. PI. H elmst. 204, 1 759, nom. rei­
T y p e s p e c i e s : U . divaricatum ( L.) Kuntze ( L. canadensis (L.) Wedd.).
=

Oblixilis Rafinesque, Fl. Tellur. 3: 49. 1 837.

T y p e s p e c i e s : Oblixilis divaricata (L.) Rafinesque ( Laportea Canadensis
=

( L.) Weddell).
Fleurya Gaud ichaud in Freycinet Voy. Bot. 497. 1 826.
T y p e s p e c i e s : Fleurya spicata Gaud ichaud ( Laportea interrupta (L.)
=

Chew).
Schychowskia End licher, Ann. Wiener Mus. 1 : 1 87 . 1 836.
T y p e s p e c i e s : Schychowskia ruderalis ( Forst.f.) Endlicher ( Laportea
=

ruderalis ( Forst.f.) Chew).

Fleuryopsis Opiz, Lotos 3 : 240. 1 8 53.
T y p e s p e c i e s : Fleuryopsis petiolata ( Decne.) Opiz ( Laportea aestuans
=

(L.) Chew).
Sceptrocnide Maximowicz, Bull. Acad . Petersb. 22: 239. 1 877.
T y p e s p e c i e s : Sceptrocnide macrostachya Maximowicz (= Laportea
macrostachya (Maximowicz) Ohwi).

Monoecious herbs or shrubs with irritant hairs. Leaves alternate. Stipu­

les fused at the base, intrapetiolar. Inflorescences unisexual or bisexual,
usually paniculate, pedunculate, the flower in loose glomerules. Perianth
of the staminate flowers 4-5-parted. Perianth of the pistillate flowers
4-parted, the 2 lateral tepals equal-sized, the dorsal and ventral ones
very unequal-sized and smaller; ovary ovoid; stigma linear. Achene
usually compressed-ovoid to -semicircular, sessile to stipitate.

D i s t r i b u t i o n : According to Chew 22 species, mostly in Africa.

This genus has a few species, among them L. aestuans, which are almost
pantropical in distribution.

I Type species chosen b y Chew Wee-Lek in Gard . Bull. S ingapore 2 1 : 1 9 5-208. 1 965.
310 U RTICACEAE

l . Laportea aestuans ( L . ) Chew, Gard . Bull. S ingapore 2 1 : 200. 1 965.

U rtica aestuans L. , Sp. PI. (ed . 2) 1 397. 1 76 3 .
Fleurya aestuans (L.) Gaudichaud ex Miquel, F l . Bras. 4 ( 1 ) : 1 96.
1 85 3 .

T y p e : Herb. Linn. 1 1 1 1 - 1 4, Surinam.

For· a comp lete list of synonyms see Chew Wee-Lek, Gard . Bull. Singapore
25: 1 64. 1 969.

Monoecious herbs, up to 2 m high; stem subligneous with few branches,

with dense to sparse up to 0. 7 mm long irritant patent hairs often inter­
mixed with 0.8- 1 . 5 mm long glandular hairs. Leaves ovate to broad­
ovate, 7-2 1 cm long, 4- 1 6 cm broad , chartaceous; apex acuminate; base
obtuse to acute; margin dentate to crenato-dentate; up to 1 .5 mm long ir­
ritant hairs, above scattered , beneath mainly on the veins; veins plane to
prominent above, prominent beneath; petioles 6- 1 5 cm long with dense
to sparse up to 0.7 mm long patent hairs often intermixed with up to 0 . 8
m m long patent glandular hairs; stipules fused a t the base, u p to I 2 m m
long.
Inflorescences unisexual or bisexual, paniculate, up to 20 cm long and
IO cm broad, the flowers in loose glomerules, the main axes either with
up to I mm long patent glandular hairs or with up to 0.5 mm long patent
irritant hairs, but mostly both glandular and irritant hairs; peduncular
bracts absent. Staminate flowers with up to I mm long pedicels; perianth
I mm high , I .5 mm broad ; tepals 5, subcucullate, fused at the base with a
single vein almost reaching the apex and bearing either 4-5 long glandu­
lar hairs or 2-5 irritant hairs 0.2 mm from the apex; stamens 5; interfloral
bracts inconspicuous, minute. P istillate flowers with up to 0.5 mm long
pedicels; perianth zygomorphous; tepals 4, the 2 lateral ones enclosing
the ovary and ea. 0.5 mm long with minute reddish pluricellular hairs at
the margin, the dorsal one with 3-5 glandular hairs or 2-5 irritant hairs,
the ventral one minute and glabrous; ovary dorsi-ventrally compressed
ovoid, ea. 0.5 mm long; stigma linear, reflexed, ea. 0.2-0. 3 mm long; in­
terfloral bracts inconspicuous. Achene short stipitate, up to 2 mm long
 U RTICACEAE 31 1

and up to 2 mm broad, compressed-ovoid to -ellipsoid with oblique

apex and up to 2 mm long pedicel.

D i s t r i b u t i o n : An almost pantropical weed .

In Surinam apparently common near Paramaribo, probably also

occurring more southward s along the Saramacca and S urinam R ivers;
a single collection is known from the Tapanahuni R iver.

J . & P.A. Florschiitz 946; Focke 26, 5 1 and 1 22 (L); van Hall s.n. (L); Hekking
880, 1 237; Hostmann s.n. (L); Lanj ouw 1 6, 27; M olkenboer s.n. (L); Pulle 2 1 ;
Samuels 427 (L); S plitgerber 2 2 ( L), s.n. (L); Tresling 67, 1 58; Tulleken 1 0 ( L);
Versteeg 668; Went 1 9 ; Wullschlagel 870 ( BR); several collections without
collector's names.

In contrast with the data given by Chew ( 1 969) in the material of L. aestu­
ans studied for this flora three d istinct groups could be distinguished : (a)
plants with sparse spiny hairs, but without glandular hairs, and with their
flowers having only spiny hairs; (b) plants with both spiny and glandular
hairs, the glandular ones abundant, and with flowers having only glandu­
lar hairs; (c) plants with both spiny and glandular hairs, both being
sparse and the glandular ones more or less rudimentary, flowers having
only spiny hairs. A form having only glandular hairs was not encoun­
tered.

3 . Boehmeria Jacquin, Enum. PI. Carib. 9. 1 760.

T y p e s p e c i e s : Boehmeria ramiflora Jacquin

Splitgerbera M iquel, Comment. Phyt. 1 33. 1 840.

T y p e s p e c i e s : Splitgerbera japonica M iquel ( = Boehmeria biloba
Weddell).

Herbs, shrubs or trees, monoecious or d ioecious, without stinging hairs

or prickles. Leaves opposite or alternate-, 3-nerved or triplinerved , those
of the adjacent nodes sometimes unequal and dissimilar; stipules free or
312 U RTICACEAE

fused at the base. Inflorescences unisexual or bisexual clusters; flowers

(sub)sessile. Staminate flowers 3-4(-5)-parted . Pistillate flowers tubular,
contracted at the throat, 2-4-toothed , somtimes entire; stigma filiform.
Achene enclosed by the persistent perianth.

D i s t r i b u t i o n : Pantropical with about 50 species of which a few in

America.

I . Boehmeria ramiflora Jacquin, Enum. PI. Carib. 3 1 . 1 760, nomen; Stirp.

Amer. Hist. 246, t. 1 57. 1 763, descr.
Cat urus ramiflora (J acquin) L., Mantisse 1 27, 1 767.
Procris ramiflora (J acquin) Poiret in Lamarck Encycl. 5 : 630.
1 804.

T y p e : Jacquin, Stirp. Amer. H ist. t. 1 57. 1 763 .

Boehmeria cuspidata Weddell, Arch. M us. H ist. Nat. Paris 9: 345. 1 856.
Boehmeria ramiflora var. cuspidata ( Weddell) Weddell, in DC. Prod. l 6( 1 ) :
1 97. 1 869.
T y p e : Broteri 280, M exico, Veracruz ( P).
Boehmeria ramiflora var. subvelutina Blume, M us. Bot. Lugd . -Bat. 2. 1 99, t.
1 3 . 1 856.
T y p e : Blume, M us. Bot. Lugd . -Bat. 2: t. 19. 1 8 56.

Subligneous monoecious shrubs, branches with appressed to patent

short hairs. Leaves alternate, those of the adjacent nodes unequal, elliptic
to lanceolate or, especially the larger ones, to ovate , 4- 1 3 cm long, 2. 5-7
cm broad ; apex acuminate to caudate, base acute to obtuse; margin cren­
a to-serrate, ciliate; lamina with appressed hairs, above 0. 2-0 . 7 mm long,
beneath 0. 1 -0.2 mm long and on the veins up to 0 . 6 mm long; tri(pli)­
nerved; veins plane above, more or less prominent beneath; petiole strigil­
lose, up to (0.5-) 1 -5 ( -7) cm long; stipules free, up to 8 mm long, narrowly
triangular, sessile, puberulous on the midrib.
Inflorescences bisexual (or unisexual?); flowers in small sessile clusters
on leafless and leafy parts of the branches; interfloral bracts 0.6 mm long,
 U RTICACEAE 313

almost obtuse. Staminate flowers sessile; perianth I mm high, 0.6 mm

broad; tepals 3 , cucullate, distinctly corniculate; stamens 3 ; pi still ode cla­
vate, 0.6 mm high. Pistillate flowers sessile; perianth up to 1 . 5 mm high,
tubular, 2-toothed, contracted at the throat, hairy; ovary sessile, com­
pressed-globose to -ellipsoid , up to I mm high, up to 0.6 mm broad;
stigma filiform, reflexed . Achene up to 1 . 2 mm high, 0.8 mm broad, ses­
sile to short-stipitate, stigma persistent.

D is t r i b u t i o n : As far as could be ascertained from the material

studied in the West lnd ies, Central America and N orthern S outh­
America.

In Surinam known only from the Wilhelmina Range, near Julianatop.

lrwin, Prance, Soderstrom & H olmgren 54605; Schulz ( LBB) 1 0265.

It is not clear if Boehmeria ramiflora is represented in the neotropics by

a single species with several taxa at infra-specific levels or by a number
of closely related species. For this treatment the latter concept has been
provisionally adopted . The genus needs revision.

4. Pilea Lindley, Collect. Bot. t. 4. 1 82 1 , nom. conserv .

T y p e s p e c i e s : Pilea muscosa Lindley ( = P. microphylla ( L . )

Liebmann).

Adicea (or Adike) Rafinesque, Anal. Nat. 1 79. 1 8 1 5.

T y p e s p e c i e s : Adicea pumila (L.) Rafinesque ex Torrey ( Pilea pumila
=

(L.) Gray).
Dubreuilia Gaudichaud in Freycinet Voy. Bot. 495 . 1 826.
T y p e s p e c i e s : Dubreuilia peploides Gaudichaud ( Pilea peploides
=

Hook. & Arn.).

M onoecious or d ioecious herbs, often succulent, sometimes suffru�

tescent, without stinging hairs or prickles. Leaves opposite, those of a
3 14 U RTICACEAE

pair unequal (in all Surinam species) or equal, similar or dissimilar. I n­

florescences unisexual or bisexual, branched to unbranched, flowers in
clusters. Staminate flowers with 4( -2-3)-parted perianth. Pistillate flow­
ers with 3-parted perianth, the middle segment usually larger than the 2
lateral ones; stigma sessile, more or less capitulate to penicillate. Achene
compressed-globose to -ovoid , envelopped by the persistent perianth;
stigma persistent to almost disappearing.

D i s t r i b u t i o n : 1 50-200 species, most of them South-American, richly

represented in the Andean region from Colombia to Peru.

I a . Leaves orbicular to ovate, shorter than 10 mm, single-

nerved (or nerveless), succulent, entire, usually crowded throughout the length
of the branches . . . . ...
. . . .. . . . .. . . . . . I . P. microphy lla
. . . . . . . . . . . . . .

b. Leaves suborbicular to ovate, longer than 10 mm, trinerved, herbaceous,

crenato-serrate, usually massed at the end of the branches . 2. P. pubescens

I . Pilea microphylla (L.) Liebmann, Kgl. Danske Vidensk. Selsk. Skr.

. V. 2: 296. 1 85 1 .
Parietaria microphylla L . , Syst. Nat. (ed . 1 0) 1 308. 1 759.
Urtica microphylla (L.) Swartz, Act. H olm. 66. 1 787.
Dubreuilia microphylla (L.) Gaudichaud in Freycinet Voy. Bot.
495 . 1 826, cited as a synonym by Weddell, Ann. Sci. Nat. Bot.
Ill. 1 8; 206. 1 852.

T y p e : Herb. Linn. 1 220-8 .

U rtica callitrichioides H. B. K . , Nov. Gen. et Sp. 2: 33. 1 8 1 7.

Pilea callitrichioides ( H . B. K . ) Schlechtendal, Linnaea 27: 494. 1 854.
T y p e : Humboldt & B onpland 255, Venezuela, Sucre (P).
Pilea muscosa Lindley, Collect. Bot. t. 4. 1 8 2 1 .
T y p e : U nknown.

Monoecious much-branched herbs, usually up to 20 cm high; branches

with linear-fusiform up to 0 . 5 mm long cystoliths, glabrous. Leaves succu­
lent, crowded throughout the length of the branches, those of a pair un-
 U RTICACEAE 315

equal, the larger ovate to obovate, up to 5 mm long and 3 mm broad, pe­

tiolate, the smaller broad-ovate to suborbicular, rarely more than 3 m m
long, sessile to subsessile; apex subacute to obtuse; base tapering, more
or less oblique; margin entire; with linear cystoliths transversely arranged
across the lamina; single-nerved ; petioles up to 2.5 mm long; stipules in­
conspicuous.
I nflorescences small, sessile to short-pedunculate, usually few­
branched ; flowers usually in bisexual clusters. Staminate flowers with an
up to 1 mm long pedicel; perianth ea. 0.6 mm broad and 0 . 5 mm high; te­
pals 4, subcucullate; obtuse and slightly corniculate. Pistillate flowers
with an up to 0.3 mm long pedicel; perianth ea. 0.4 mm high; the larger te­
pal subcucullate, subherbaceous to herbaceous, the 2 smaller ones almost
plane, subtriangulate, membranaceous to subherbaceous, acute; ovary
short stipitate. Achene (ovoid to) broad elliptic, compressed , up to 0 . 5
m m long, usually 0.3 mm broad, apiculate, very short stipitate to subses­
sile, partly enveloped by the persistent fruiting, up to 0.6 mm high peri­
anth; stigma almost disappearing.

D is t r i b u t i o n : A weed occurring throughout the tropics.

In the cultivated area of north-eastern Surinam.

J. & P.A. Florschiitz 1 099; K ramer & Hekking 3 1 29; Soepratq 89, without
collector's name and number, collected at Paramaribo.

2. Pilea pubescens Liebmann, Kgl. Danske Vidensk. Selsk. Skr. V. 2:

302. 1 85 1 .

T y p e : Lund s.n., Brazil (C).

Pilea guyanensis Weddell, Ann. Sci. Nat. Bot. I l l . 1 8: 22 1 . 1 852.

Pilea pubescens var. guyanensis (Weddell) Weddell. in D C . Prod. 1 6( 1 ) : 1 53 .
1 869.
T y p e : Melinon 55, French Guiana (P).
Pilea gaudichaudiana Weddell, Ann. Sci. Nat. Bot. I l l . 1 8: 226. 1 852.
S y n t y p e s : Gaudichaud 93, 1 077, Guillemin 74 1 , Brazil (P).
316 U RTICACEAE

Fig. 3 Pi/ea p ubescens; from Arch. Mus. H ist. Nat. Paris 9: t. 6, C, I. 1 856- 1 85 7 .
 U RTICACEAE 317

Monoecious (or dioecious?) herbs with repent stem and erect o r ascend­
ing branches; stem and branches usually with dense 0. 2-0.5 mm long line­
ar-fusiform longitudinally arranged cystoliths in the epidermis, glabrous
to pubescent to strigillose, sometimes patent and appressed hairs inter­
mixed . Leaves usually massed at the end of the branches, the upper ones
usually larger and narrower than the lower ones, those of a pair unequal
but similar, ovate to broad-ovate to suborbicular, 1 0-52 mm long, 8-3 1
mm broad ; apex almost obtuse; base acute to truncate, more or less ob­
lique; margin crenato-serrate, usually ciliate; lamina with dense 0. 1 -0 . 3
m m long linear-fusiform cystoliths, above glabrous to strigillose, be­
neath strigillose and usually more d ensely so on the veins; trinerved, only
the costa reaching the apex, wj th (2�)3-5( -6) pairs of secondary veins;
veins plane above, prominent beneath; petiole pubescent to strigillose; sti­
pules up to 0.2 mm long.
I nflorescences unisexual or bisexual, dichotomously branched with
pedicellate flowers in loose clusters at the terminal axes, occasionally soli­
tary; peduncular bracts present, membranaceous, up to 0.5 mm long.
Staminate flowers with up to 3 mm long hairy pedicels; perianth up to 2
mm high; tepals 4, fused at the base, subcucullate; stamens 4; pistillode
0. 1 -0.2 mm. Pistillate flowers with up to 0 . 3 mm long glabrous pedicels;
perianth up to 0.5 mm high, 3-fid, the large tepal more or less cucullate
and herbaceous, the 2 smaller ones sharp triangulate and herbaceous to
membranaceous; ovary compressed-ovoid to -globose with readish glo­
bose-capitate pluricellular hairs; stigma sessile, more or less oblique, ca­
pitulate-penicillate. Achene compressed-ovoid to globose, oblique, short
stipitate, up to 0.6 mm high and 0 . 3 mm broad .

D i s t r i b u t i o n : Throughout the American lowland tropics.

In Surinam known from the right Kabellebo River, the Lucie River, the
S ipaliwini River and from near Paramaribo .

Florschiitz & Maas 2527; l rwin. Prance, Soderstrom & Holmgren 54667; Stahel
s.n. ; Wessels Boer 723.
318 U RTICACEAE

The Surinam collections of this species are not very similar.

Additi on:

Pi/ea now appears to be represented in Surinam by 3 species. The third

species is P. imparifolia Weddell, a dioecious epiphytic glabrous herb
with thin repent sterile stems bearing almost equal and similar leaves
on each node, and ascending o r erect flowering branches bearing unequal
and dissimilar leaves on each node; the leaves contain dense linear­
fusiform cystoliths.