Three New South American Species of Randia
(Rubiaceae, Gardenieae)
Claes G. R. Gustafsson
Botanical Institute, Systematic Botany, Box 461, 405 30 Goiteborg, Sweden.
claes.gustafsson@systbot.gu.se
ABSTRACT. Three new species of South American
Randia (Rubiaceae, Gardenieae) are described and
illustrated: Randia wigginsii Standley ex Gustafs-
son from montane tropical forests in Ecuador and
northern Peru is mainly recognized by its tomentose
to velutinous corolla tube and the lanate-velutinous
pedicels and fruits. The possible features of being
unarmed and having single male flowers are dis-
cussed. Randia pubistyla Gustafsson from lowland
tropical dry forests in western Ecuador and Colom-
bia is recognized by its puberulous style, the to-
mentose to puberulous fruits that usually are
crowned by a calyx with reflexed lobes, and by the
small thorns. Randia longifolia Gustafsson from
lowland tropical humid forests in western Ecuador
is mainly recognized by its relatively large, gla-
brous corolla, the linear calyx lobes, and by the
reticulate finer vasculature on the abaxial surface
of the leaves. The affinities for each species are
discussed.
Randia is a neotropical genus of approximately
90 species ranging from ca. 300N to 30�S. In South
America and the Caribbean there are approximate-
ly 45 species. The genus is represented by shrubs,
trees, and lianas in deciduous and evergreen veg-
etation from sea level to about 3300 m elevation.
Randia can be recognized from other members of
the Gardenieae by the following combination of
characters: dioecious (female flowers with nonfunc-
tional stamens, male flowers with a nonfunctional
stigma and rudimentary ovary), pollen in permanent
tetrads, a unilocular ovary with two parietal pla-
centas, fruits with many discoid seeds embedded
in a sweet pulp that turns dark when dry, thorns,
and conspicuous short-shoots with clustered stip-
ules and leaves. There are, however, exceptions.
Monoecious and hermaphroditic species have been
reported (Lorence & Dwyer, 1987; Burger & Taylor,
1993), pollen in dyads or monads have been re-
ported (Burger & Taylor, 1993), some species are
unarmed (Burger & Taylor, 1993; pers. obs.), and
sometimes the short-shoots are less conspicuous
(Burger & Taylor, 1993; pers. obs.). Within the ge-
nus there is also variation in the structure of the
inflorescence. The inflorescences are usually ter-
minal but sometimes axillary or cauliflorous (Burger
& Taylor, 1993). Female flowers are usually solitary
but sometimes in fascicles of 2 to 8 flowers (Burger
& Taylor, 1993). The male flowers are usually in
fascicles with a few to several flowers but some-
times form cymes with many flowers (Lorence &
Nee, 1987; Lorence & Dwyer, 1987) and are some-
times solitary. This large variation makes it difficult
to understand which genus has the closest affinities
to Randia.
Robbrecht and Puff (1986) discussed that all
genera in Gardenieae with pollen shed in perma-
nent tetrads may represent a natural group. This
gained support in a phylogenetic analysis by Pers-
son (1996) but was contradicted by Andreasen and
Bremer (1996, in press) and Persson (in press), who
discussed that pollen in tetrads may have arisen
several times in Gardenieae. In Persson's (1996)
analysis Casasia gained support as sister to Ran-
dia, but the later mentioned analyses did not point
out any strongly supported sister to Randia. Other
neotropical genera that in these analyses (Andreas-
en & Bremer, 1996, in press; Persson, in press)
grouped together with Randia and Casasia as pos-
sible relatives were Rosenbergiodendron, Sphinc-
tanthus, and Tocoyena. Sphinctanthus and Tocoyena
were not included in the analyses by Andreasen
and Bremer (1996, in press). In this group the ge-
nus with the morphologically nearest affinities to
Randia is Casasia by being dioecious and having
pollen in tetrads. Lorence (1986) and Lorence and
Dwyer (1987) discussed the morphological features
in Casasia and Randia and also expressed doubt
(Lorence & Dwyer, 1987) whether Casasia can be
maintained as a valid genus. More phylogenetic
studies in Gardenieae need to be done in order to
find the monophyletic group to which Randia be-
longs.
The taxonomical work in Randia is complicated
by the intraspecific variation in size and shape of
leaves, calyx lobes, degree of pubescence, persis-
tence of stipules and calyx, and number of thorns.
NovoN 10: 201-208. 2000.


Novon
Extensive collecting in the Neotropics in the last
50 years has dramatically increased the number of
herbarium specimens available, facilitating our un-
derstanding of the genus. Even so, several species
are still unsatisfactorily collected with only a single
or a few flowering specimens collected or none at
all. During the course of a revisionary study of
South American and Caribbean representatives of
Randia (Gustafsson, in progress) collections rep-
resenting a number of undescribed species have
been encountered, three of which are described be-
low.
Randia wigginsii Standley ex Gustafsson, sp. nov.
TYPE: Ecuador. Loja: along Rfo Juntas, about
14-15 km S of San Lucas, ca. 2000 m, 10 Oct.
1944, 1. L. Wiggins 1099 (holotype, F; isotype,
NY). Figure 1.
Species insignis habitu arbore 4-10(-16) m alta, flori-
bus solitariis pedicellatis, pedicellis et ovariis et calycum
tubis lanato-velutinis. corollarum tubis 3-4 mm diam. to-
mentoso-velutinis, corollanrm lobis ciliatis, fructibus 4-5
x 3-4 cm longis. velutinis, a congeneris distincta.
Trees 4-10(-16) m high, dioecious. Leaf branch-
lets velutinous when young, glabrous when old, un-
armed. Stipules fused at base, on long shoots not
seen, persistent on brachyblasts, broadly to narrow-
ly triangular, 5-12 X 3-6 mm, acute at apex, outer
surface minutely verruculose, glabrate, inner sur-
face with hairs and colleters at base, margin ciliate
to glabrous. Petioles 5-17 X 1-2 mm, velutinous.
Leaf blades subcoriaceous, broadly elliptic to ob-
ovate, 8-13(-17) X 4-7(-8) cm, shortly acuminate
at apex, cuneate to shortly attenuate at base, ad-
axial midrib and secondary nerves slightly to dis-
tinctly impressed, tertiary veins slightly impressed
to plane, veinlets plane, abaxial midrib and sec-
ondary nerves prominent, tertiary veins slightly el-
evated to level, veinlets level, adaxial leaf surface
puberulous on midrib, otherwise glabrous, abaxial
leaf surface tomentose to velutinous all over or
mainly on midrib and secondary veins, margins gla-
brous. Flowers unisexual. Male flowers solitary,
pedicels 10-30 X 1-1.5 mm, lanate-velutinous.
Calyx tube 1-2 mm long, lanate-velutinous exter-
nally, with a ring of hairs at base internally, lobes
5, narrowly triangular to narrowly elliptic, 9-14 X
1-2 mm, acute at apex, pilose externally, glabrous
or with a few scattered hairs internally, margins
membranaceous, ciliate. Corolla white, salverform;
tube 15-18 X 3-4 mm medially, tomentose to ve-
lutinous externally, pilose in upper half internally,
glabrous below; lobes 5, broadly elliptic to subor-
bicular, ca. 12-14 X 11-12 mm, obtuse at apex,
glabrous externally, pilose toward base internally,
margins ciliate; anthers sessile, ca. 10 mm long,
dorsifixed in lower third, attached distally in the
widened part of the corolla tube, connective pro-
cess minute or absent; style glabrous, nonfunctional
stigma lobes ca. 4 mm long; ovary rudimentary. Fe-
male flowers not seen. Pedicels of fruit 10-25 X
1.5-3 mm, glabrate; fruits broadly ellipsoid to sub-
spherical, 4-5 X 3-4 cm, surface with slightly el-
evated veins, densely velutinous when young, fruit
wall ca. 1 mm thick, apex umbonate, persistent ca-
lyx lobes similar to those of male flowers in shape,
size, and pubescence, seeds irregularly discoidal,
10-14 mm diam., imbedded in pulp.
Distribution and habitat. This new species is
found in montane tropical forest from 1500 to 2250
m in Ecuador and northern Peru. The only two flow-
ering collections examined (Wiggins 10996, Espi-
nosa 1919) were collected in August and October.
Randia wigginsii is easily recognized by the sol-
itary, pedicellate, male and female flowers; by the
flowering pedicel, ovary, and calyx tube being la-
nate-velutinous; by the tomentose to velutinous co-
rolla tube 3-4 mm in width; by the ciliate corolla
lobes, and by the quite large velutinous fruit (4-5
x 3-4 cm). Useful vegetative characters are the
petiolate, subcoriaceous leaves with glabrous ad-
axial sides (apart from the midrib) and slightly to
distinctly impressed midrib and secondary veins.
The abaxial sides are tomentose to velutinous (at
least on vasculature) and have prominent midrib
and secondary veins.
Randia wigginsii is here described as unarmed
by assumption, as no thorns have been found on
the studied material, and no mention of being
armed is mentioned on the specimen labels. The
only two male flowering specimens examined had
solitary flowers, which is unusual in Randia. Fur-
ther collecting will reveal if this is the true state
for R. wigginsii. The closest relatives are probably
found among a small group of undescribed montane
species (Gustafsson, in prep.) with large fruits, pu-
bescent leaves, and small pubescent flowers.
The name Randia wigginsii was written on the
here designated holotype by Paul C. Standley but
was never published by him.
Paratypes. ECUADOR. Chimborazo: canyon of Rfo
ChanchAn, ca. 5 km N of Huigra, 1500-2(XX) m, Camp
3316 (MO, NY, S, US). El Oro: Guayquichuma, 03�49'S,
79�34'W, 1600 m, Cornejo & Bonifaz 6022 (GB, GL(AY);
San Antonio, E of Zaruma, 2100-2200 m, Espinosa El919
(F. NY). Loja: 25 km Catamayo-Catocha, turnoff at Las
Chinchas 2.3 km toward Piftas, 03�57'21"S. 79�29'07"W,
2250 m, Jorgensen et al. 1464 (GB); Cerro de Celica. 2.7
km on Celica-Guachanama road. 04�05'46"S.
79�56'45"W, 2250 m, J0rgensen et al. 96 (GB). PERU.
Piura: Huancabamba. Canchaque. Cerro Chorro Blanco.
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New South American Randia
203
F
Figure 1. Randia wigginsii Standley ex Gustafsson. -A. Flowering branch, male specimen. -B. Adaxial side of
leaf. -C. Stipules, small stipules (first and second from left) from short-shoot, large stipule from long-shoot. -I). Male
flower bud. -E. Male flower in anthesis. -F. Calyx of male flower with two lobes removed. -G. Male corolla, opened.
-H. Fruit. A, Wiggins 10996 (F); B, Diaz & Balde6n 2455 (GB); C-E, Wiggins 10996 (F): F, Espinosa 1919 (F): G,
Wiggins 10996 (F): H, Diaz & Balde6n 2455 (GB).


Novon
1500-19(X) n, Diaz S. & Baldedn 2455 (GB), 1600 nm,
Diaz S. et al. 2779 (GB).
Randia pubistyla Gustafsson, sp. nov. TYPE: Ec-
uador. Guayas: 15 km E of Guayaquil, Cerro
Mirador de los Monos, 02�10'S, 79058'W, 200
m, 26 Feb. 1992, D. Rubio & W. Palacios 2439
(holotype, GB; isotypes, MO not seen, QCNE
not seen). Figure 2.
Species Randiae hebecarpae Bentham affinis. sed stylis
pubenlis, fructibus majoribus ad 2.5 x 2 cm diam., spin-
is parvis ad 0.5 cm longas, foliis majoribus 7-16 X 4.5-
10 cm differt.
Shrubs to small trees to 6 m high, deciduous,
dioecious. Leaf branchlets tomentose when young,
later glabrate, thorns 1 or 2 at base of brachyblast,
the thorns short, only to 5 mm, straight to slightly
deflexed. Stipules fused at base, persistent on long
shoots, narrowly to broadly triangular, 4-6 X 2-3
mm, aristate to mucronate at apex, outer surface
tomentose, inner surface with hairs and colleters at
base, stipules on brachyblasts persistent, broadly
triangular to suboblong, 3-8 X 2-4 mm, obtuse to
mucronate at apex, outer surface minutely verru-
culose, with many parallel veins, glabrous, inner
surface with hairs and colleters at base, margin mi-
nutely ciliate. Petioles 2-8 X 0.5-1 mm, tomentose
to puberulous. Leaf blades papery, broadly elliptic,
7-16 X 4.5-10 cm, shortly acuminate to acumi-
nate, sometimes apiculate at apex, attenuate at
base, all veins on adaxial side � plane, abaxial
midrib and secondary veins prominent, tertiary
veins less prominent to plane, veinlets plane, retic-
ulate, adaxial leaf surface glabrous or with scat-
tered hairs on midrib, abaxial leaf surface tomen-
tose to puberulous when young, glabrescent, the
hairs on midrib and secondary veins more persis-
tent, blade margins � ciliate. Flowers unisexual.
Male inflorescence in fascicles of 2 to 8 flowers,
pedicels 4-8 x 0.3-0.5 mm, tomentose. Calyx tube
to 0.5 mm long, pilose externally, glabrous inter-
nally, lobes 5, irregularly linear-oblong, 3-7 X 0.3-
1.5 mm, acute to shortly acuminate at apex, gla-
brous or pilose toward base externally, glabrous
internally, margin fimbriate to ciliate. Corolla white,
salverform, tube 9-13 X ca. 1.0 mm medially, dis-
tally widened, pilose externally, pilose internally
but glabrous toward base, lobes 5, oblong to broad-
ly obovate, 5-9 x 4-7 mm, obtuse to rounded at
apex, � pilose externally, glabrous internally but
basally pilose, margin glabrous; anthers sessile, 2-
3 mm long, dorsifixed in lower third, attached dis-
tally in the widened part of the corolla tube, con-
nective process absent; style puberulous, nonfunc-
tional stigma lobes ca. 1 mm long; ovary
rudimentary. Female flowers not seen. Pedicels of
fruit 5-8 x 1.0-2.0 mm, glabrate; fruits subspher-
ical to ellipsoid, ca. 2-3 X 1.8-2.5 cm, tomentose
when young, later becoming puberulous, yellow
when mature, fruit wall ca. 0.5 mm thick, calyx
reflexed and persistent; seeds irregularly discoidal,
5.0-9.0 mm diam, imbedded in pulp.
Distribution and habitat. Randia pubistyla is
found in lowland tropical dry forests in western Co-
lombia and western Ecuador. The only three flow-
ering collections examined (Brand 1082, Haught
4792, Rubio & Palacios 2439) were collected in
February (Ecuador) and April (Colombia).
Randia pubistyla is characterized by the puber-
ulous style and the tomentose to puberulous fruit,
which is often characteristically crowned by the al-
most free, reflexed calyx lobes. The small (only to
0.5 cm long) thorns on the branchlets, together with
the usually reticulate venation of the broadly ellip-
tic leaf blades, are useful vegetative characters. It
is similar to Randia hebecarpa Bentham, a species
found in northeastern South America, in size and
pubescence of the flowers and leaf shape, but that
species has a smooth style, distinctly smaller fruit
(to 1.5 cm diam.), larger thorns (0.5-1.6 cm long),
and smaller leaves (3.0-6.5 X 1.2-2.7 cm).
Paratypes. COLOMBIA. Antioquia: La Playona, 15
km from Turbo, 10 Mar. 1958, Bernal 10 (COL 2 sheets);
Mun. de Arboletes, 5 km S of Arboletes on rd. to Turbo,
08�49'N, 76�27'W, 0-50 m, Zaruchi et al. 4940 (F, K,
MO, NY): Turbo, hills near Rfo Mulatas, 300 m, Haught
4792 (COL, US): rd. Tap6n del Darien, sector Rfo Le6n-
Lomas Aisladas, km 37, 0-50 m, Brand 1212 (COL. MO),
Brand 1082 (COL, MO); Corrto. Currulao, new Antioquia
rd. E-NE of Turbo, 11 km from Currulao, 0-50 m, Callejas
et al. 4966 (MO 2 sheets, NY). Choco: Mun. de Rfosucio,
vicinity of El Salto de Tilupo, corrto. Tilupo, Romero-Cas-
taneda 6246 (COL); Parque Natural Nacional, Los KAtyos,
Rfo Peye rd. to Tilupo, 0-100 m, Le6n 470 (COL); Los
K6tyos, Tilupo, 250-350 m, Ledn 492 (COL); ascent to El
Alto de Lim6n, 250-300 m, Le6n 430 (COL, MO); Peye,
rd. to El Alto del Iim6n, 50-200 m, Ledn & Forero P
1795 (COL, MO). ECUADOR. Esmeraldas: ca. 7 km SW
of Sia. 00�51'N, 78�58'W, 50 m, Webster 22822 (MO);
Cerro Mutiles, 00�54'S, 79�37'W, 200-300 m, Cornejo &
Bonifaz 5262 (GB, GUAY). Manabi: 5 km from Jama on
rd. to Pedernales, 00�10'N, 80014'W, 50 m. Cornejo &
Ronifaz 5836 (GUAY). Guayas: Bosque protector Parafso,
02�12'S, 79�57'W, 200 m, Cornejo & Bonifaz 1364 (GB,
GUAY), 150 m, 5598 (GB, GUAY); Cerro Azul, W of Gua-
yaquil, Asplund 15402 (S); Cerro Azul, 200 m, Harling
4839 (S); Cerro Azul. 02�08'S, 79059'W, 400-500 m, Cor-
nejo & Bonifaz 2926 (GB, GUAY); Cerro Azul, 10 km NW
of Guayaquil on rd. to Salinas, 50-1X) m, Dodson & Thien
527 (MO); Cerro Azul, Cordillera Chong6n-Colonche, be-
hind Cemento Nacional, 12 km W of Guayaquil. Canyon
of Canoa, 02�15'S, 80�00'W, Gentry & Josse 72348 (GB);
Chong6n, Hacienda Cerro Azul, 3()-500 m, Harling et
al. 9509 (GB); Cant6n Guayaquil, Bosque Protector Cerro
Blanco. 15 km on rd. to Salinas, 02�10'S, 79�58'W, 350
204


Volume 10, Number 3
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Gustafsson
New South American Randia
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1 cm
4 I
il ^:
A $ %
^r tj /^^ V:
:~- ^ / 5^t: \ r'4
S. 1 mm7
F GH E
9-f^LdC ^
Figure 2. Randia pubistyla Gustafsson. -A. Fruiting branch, female specimen. -B. Thorns. -C. Stipules from
short-shoot. -I). Short-shoot with male inflorescence. -E. Male flower bud. -F. Calyx of male flower, opened. -G.
Male corolla, opened. -H. Style. -1. Detail of style. -J. Detail of persistent calyx on fruit. A-B, Harling 4822 (S);
C-), Rubio & Palacios 2439 (GB); E-I, Haught 4792 (COL); J, Cornejo & Bonifaz 5598 (GB).
205
1 cm


Novon
m, Rubio et al. 1770 (GB, GUAY). 1810 (GB. GUAY), 300
m, Rubio et al. 1993 (MO, GUAY), 02�10'S, 79�50'W, 100
m, Tipaz et al. 839 (GB, GUAY); 13 km on rd. to Salinas,
02�10'S, 79�58'W, 350 m, Rubio & Tipaz 2383 (GB);
along Sendero Buenavista above visitor center. 02�10'S,
79�58'W, 250 m, Neill & Ndiiez 10512 (GB), 02�10'S.
80�10'W. 300-400 m, Neill & Niirlez 10662 (GUAY); Re-
serva Ecologfa Churute, Cerro Cimaldn. 02�25'S,
79�38'W, 50-1(X) m, Cornejo & Bonifaz 5126 (GB,
GUAY); Rfo I)aule below Pichincha, Hacienda Santa Bar-
barita, Harling 4822 (S).
Randia longifolia Gustafsson, sp. nov. TYPE: Ec-
uador. Los Rfos: Hacienda Clementina, 200 m,
27 Jan. 1947, G. Harling 186 (holotype, S).
Figure 3.
Species Randiae carlosianae K. Krause affinis, sed cor-
ollis majoribus, corollarum tubis 30-40 mm longis, cor-
ollarum lobis 20-25 mm longis, calycis lobis anguste
triangularibus ad linearibus 1.0-2.0 mm latis, venis par-
allelis numeris differt.
Shrubs to small trees to 6 m high, dioecious. Leaf
branchlets puberulent when young, hairs � erect,
unarmed. Stipules fused at base, persistent on long
shoots, narrowly triangular, 12-15 X 2-4 mm, acu-
minate to acute at apex, with many parallel veins,
glabrous externally, with hairs and colleters at base
internally, margin ciliate, stipules persistent on
brachyblasts, broadly triangular to narrowly trian-
gular, ca. 2-10 X 2-4 mm, acute to mucronate at
apex, with many parallel veins, verruculose, gla-
brous externally, with hairs and colleters at base
internally, margin ciliate. Petioles 3-6 X 0.8-1.2
mm, puberulent. Leaf blades papery, obovate, (11-
)13-21 X 4-8(-9) cm, shortly acuminate at apex,
shortly attenuate at base, adaxial midrib impressed
at base, otherwise all veins � plane, abaxial midrib
and secondary veins prominent, smaller veins less
prominent, reticulation evident, adaxial leaf surface
glabrous, minutely tuberculate, abaxial leaf surface
puberulous on midrib, secondary and tertiary veins,
margin � ciliate. Flowers unisexual. Male inflores-
cence in fascicles of 2 to 6 flowers, pedicels 5-10
X ca. 1 mm, puberulent. Calyx tube to 0.5 mm
long, glabrate externally, glabrate with a few basal
hairs internally, lobes 5, narrowly triangular to lin-
ear, 10-15 X 1-2 mm, acute at apex, with parallel
venation, glabrous to puberulent throughout, mar-
gins � ciliate; corolla white, salverform, tube 30-
40 X 2.0-4.0 mm medially, distal part widened,
glabrous externally, puberulent in upper part, gla-
brous below internally, lobes 5, ovate to broadly
elliptic, ca. 20-25 X 10 mm, subacute at apex,
glabrous or with a few small hairs externally, gla-
brous or with a few basal hairs internally, margin
glabrous; anthers sessile, ca. 6.0 mm long, dorsifix-
ed in lower third, attached distally in the widened
part of the corolla tube, connective process absent;
style glabrous, nonfunctional stigma lobes ca. 5.0
mm long; ovary rudimentary. Female flowers not
seen. Pedicels of fruit 2.0-10 mm X ca. 2.0 mm.
Fruits ellipsoid, 3-4 X 2.5-3 cm, puberulous when
young, glabrescent, yellow when mature, fruit wall
ca. 1 mm thick, persistent calyx erect; seeds irreg-
ularly discoidal, 7-10 mm diam., imbedded in
pulp.
Distribution and ecology. This species is known
only from the rapidly diminishing lowland tropical
humid forests in western Ecuador. The two sole
flowering specimens examined (Harling 186, Cor-
nejo & Bonifaz 880, unicates) were collected in late
November and January.
Randia longifolia is recognized by the relatively
large, glabrous corolla and by the long, almost free,
characteristically linear calyx lobes with parallel
venation. The fruits are usually crowned by a per-
sistent erect calyx. On the abaxial surface of the
leaves, finer vasculature makes the reticulation ev-
ident.
Randia longifolia might be confused with R. car-
losiana K. Krause, another species occurring in the
same area, by the similar fruits and large leaves,
but that species differs by being armed, by having
smaller corollas with tubes 20-26 mm long and
lobes 10-14 mm long, and by usually having foli-
aceous, obovate calyx lobes (broader than 2 mm)
with usually reticulate venation.
Randia longifolia is here described as unarmed
by assumption, as no thorns have been found on
the studied material and no mention of being armed
is mentioned on the specimen labels.
Paratypes. ECUADOR. Esmeraldas: Cerro Mutiles,
00�54'N, 79�37'W, 250 m, Cornejo & Bonifaz 5185 (GB,
(UAY). Manabi: Vuelta larga, 00�45'S, 80�50'W, 200 m,
Bonifaz 837 (GB, GUAY); rd. Portoviejo-Pichincha, 3 km
E of San Phlcido, 150-200 m, Harling & Andersson 24888
(GB). Los Rios: Hacienda Clementina, 200 m, Fagerlind
& Wibom 2650 (S); Hacienda Clementina, Cerro Samama,
above Rfo Mombe, ca. 38 km NE of Babahoyo, 01�39'S,
79022'W, 200-400 m, St&hl & Knudsen 1182 (GB); Jau-
neche, 70 km on rd. Quevedo-Palenque via Mocachi,
Cant6n Vinces, 100 m, Dodson et al. 7993 (MO), Dodson
& Gentry 9883 (MO, US); Jauneche, rd. Mocachi-Pal-
enque, 15 km E on rd. Empalme-Balzar, 50-100 m, Dod-
son et al. 14461 (MO, NY); near Pichilingue, 50-1(X) m,
Jdtiva & Epling 307 (NY, S). Guayas: Rfo Ayampe, Cerro
El Burro, 01�41'S, 80�40'W, Cornejo & Bonifaz 880
(GUAY); Bosque Protector Cerro Blanco, 02�10'S,
79�58'W, 400 m, Cornejo & Bonifaz 1783 (GB, GUAY).
El Oro: between Piedras and Moromoro, 300 m, Escobar
746 (MO).
Acknowledgments. I am grateful to the curators
of the following herbaria for making their collec-
tions available for study: COL, F, GB, GUAY, K,
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Volume 10, Number 3
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Figure 3. Ranndia longifolia Gustafsson. -A. Flowering branch, male specimen. -B. Fruiting branch, female spec-
imen. -C. Stipules. small stipule (left) from base of short-shoot, large stipule (right) from long-shoot. -I). Male flower
bud. -E. Male flower. -F. Calyx of male flower, opened. -G. Male corolla, opened. -H. Style. A. Harling 186 (5):
B. Bonifaz 837 ((;G): C-H, Harling 186 (S).
207


208
Novon
MO, NY, S, US. Special thanks are due to the cu-
rators of GUAY, MO, and QCNE who sent several
collections as gifts. I thank Pollyanna Lidmark for
preparing the excellent illustrations and Lennart
Andersson for providing useful comments on the
manuscript.
Literature Cited
Andreasen, K. & B. Bremer. 1996. Phylogeny of the sub-
family Ixoroideae (Rubiaceae). Opera Bot. Belg. 7: 119-
138.
& - . In press. Combined phylogenetic anal-
ysis in the Rubiaceae-lxoroideae: Morphology, nuclear.
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