Systematic Botany (2019), 44(1): pp.

146–183
© Copyright 2019 by the American Society of Plant Taxonomists
DOI 10.1600/036364419X697994
Date of publication February 5, 2019

A Revision of Cuphea section Amazoniana s. s. (Lythraceae)

Shirley A. Graham
Missouri Botanical Garden, 4344 Shaw Blvd., St. Louis, Missouri 63110, USA
(shirley.graham@mobot.org)

Communicating Editor: Susana Freire

Abstract—Cuphea section Amazoniana, one of 13 sections in Cuphea, comprises 20 species, mostly narrow endemics in Colombia, Ecuador, and the
Guiana Shield region of Venezuela, Guyana, French Guiana, and Brazil. The species inhabit river margins, tepuis, scrubland, savanna, and low
montane forests in tropical to cold temperate climates. They are subshrubs defined by the common presence of a strigose indumentum of bifid
(malpighiaceous) trichomes; small elliptic to narrowly lanceolate or linear coriaceous leaves; and tubular flowers to 9 mm long with 11 deeply
inserted stamens included in the floral tube. In morphology, they are similar to or partially overlap species currently classified in Cuphea sections
Trispermum and Heteranthus. The revision provides a new key, comparative morphological descriptions, first descriptions and illustrations of pollen
and seed morphology, and distribution maps. Cuphea saxatilis S. A. Graham, sp. nov. from Colombia and Cuphea trisperma S. A. Graham, sp. nov.
from Venezuela are newly described. Cuphea killipii Lourteig is placed in synonomy of C. philombria Lourteig. Presence or absence of a floral spur,
position and shape of a nectary-like floral organ, the “disc”, and presence of two major pollen types suggest the section is not monophyletic, but
consists of a minimum of two phylogenetic lineages, one in the Guiana Shield region in Venezuela and eastward, the other in low montane forests
and on outliers of the Guiana Shield in Colombia and northern Ecuador. Fossilized Cuphea pollen grains resembling two types of pollen known in
sect. Amazoniana, establish the presence of the genus in the Mio-Pliocene to Holocene of northern South America in areas occupied today by members
of the section.

Keywords—Endemic species, fossils Guiana Shield, morphology, pollen, seeds.

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Cuphea sect. Amazoniana (Lourteig) Lourteig is one of 13
taxonomic sections of the neotropical genus Cuphea, the largest
genus of the Lythraceae with ca. 250 species. The present study
contributes to a revision-in-progress of the only monograph of
Cuphea (Graham 1988, 1989, 1998, 2017, 2019 in press; Graham
and Cavalcanti 2013). The monograph was initiated by
E. Koehne (1881a, b) and finalized in a monograph of the
family (Koehne 1903). The genus is immediately recognizable
in the Lythraceae by its unusual seed dispersal mechanism.
The elongate seed-bearing placenta is enclosed within a thin-
walled capsule surrounded by a persistent, 12-veined floral
tube. Unlike all other lythracean genera, the placenta with
attached immature seeds rises prior to full seed maturation
through complementary slits in the capsule wall and floral
tube, carrying seeds up and out of the tube for final maturation
and dispersal (Graham and Graham 2014: Fig. 18).
Section Amazoniana in subg. Bracteolatae S. A. Graham, the
larger of the two subgenera of Cuphea, is the most recently
described section, and one of the most homogeneous. It
consists of subshrubs with: small, elliptic or lanceolate to
linear, membranaceous to coriaceous leaves; plant surfaces
covered by a strigose indumentum of bifid (malpighiaceous)
trichomes which are usually mixed with a hirsute and/or
setose indumentum; flowers small for the genus at 2.5–9 mm
long; 11 included stamens; and conspicuous epicalyx seg-
ments in the form of flattened or cylindric lobes alternating
with the sepals. A globose to elongated fleshy organ, the
“disc,” stands external to the base of the ovary in the floral
tube.
The number of collections available for study is small due to
the relative inaccessibility of the plants and their narrow
distributions. More than half of the species are isolated along
river systems in savannas and on high mountain slopes of
tepuis surrounded by dense forests in the Guiana Shield region
and outliers of the Shield in Venezuela, Guyana, French
Guiana, northern Brazil, and sub-Andean Colombia and
Ecuador (Fig. 1). Some new collections are available as a result
of recent field work for floras (Steyermark et al. 1995–2005) and
checklists of floras of selected mountain chains and tepuis of
the Guiana Shield (e.g. Cortés-B. et al. 1998; Berry and Riina
2005; Cleef and Velasquez 2005; Funk et al. 2007; Rodrı́guez R.
et al. 2008; Grande et al. 2012).
The revision builds on the initial delineation of the section
and subsequent additions by Lourteig (1959, 1986, 1996). It
provides: a key to 20 species, two of them new; historical
taxonomic background; comparative morphological de-
scriptions, including the previously unstudied pollen and seed
morphology; distribution maps; a brief ecological and evo-
lutionary perspective; and notes on relationships for each
species suggested by morphology. The revision benefitted by a
generous loan of more recently collected specimens and types
from the Herbario Nacional Colombiano (COL).
Materials and Methods
Herbarium specimens and/or digital images from the following her-
baria were studied: BR, CAY, COL, F, GH, GOET, K, LD, MA, MG, MPU,
MO, NY, OXF, P, RB, TCD, U, UC, US, VEN, and W. Morphological
measurements of larger floral structures were taken from dried, rehydrated
specimens using a light microscope micrometer. Trichome, floral, pollen,
and seed samples were prepared for scanning electron microscope (SEM)
study and photography from herbarium material. Floral organs, trichomes,
and seeds were dry-mounted and sputter-coated at 35 A for 120 sec using a
gold-palladium target. Observations, measurements, and photographs
employed a JEOL Neoscope JCM-5000 scanning electron microscope.
Pollen grains were first heated in a drop of lactic acid on a glass slide to clear
pollen contents and then surveyed using light microscopy for shape and
colpi patterns. For SEM, pollen from mature flower buds was transferred
from anthers in 50% ethyl alcohol to stubs, evaporated, and sputter-coated
under the same settings as the other morphological material. Scanning
electron microscopy (SEM) provided details of the pollen exine and ac-
curate measurements of size.
Selected morphological characters are illustrated in Figs. 1–6. The
species and their disiributions are illustrated in Figs. 7–21. Illustrations of
the species use silhouettes from type specimens or a typical representative
(Figs. 7, 10, 11 12, 14, 15, 18–21) to portray their distinctive habits. Lateral
views of the flowers allow comparison of external details. Distribution
maps were prepared from geographical degree coordinates taken from
specimen labels and converted to decimal degree data. In the absence of
coordinates on specimen labels, coordinates in decimal degrees (degree
symbols omitted for brevity) were determined for the nearest locality cited
on the collections and added in brackets immediately following the lo-
cality. Maps were produced in ArcGIS (ESRI, New York) from files of the
decimal coordinates. A risk of extinction was estimated for each species
utilizing the categories and criteria established by the IUCN Red List
(IUCN 2012). Estimates of the extent of occurrence (EEO) and area of

146
 2019] GRAHAM: CUPHEA SECTION AMAZONIANA
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Fig. 1. Distribution of Cuphea sect. Amazoniana in northern South America. Dotted line 5 approximate limits of the Guiana Shield (shaded) plus outliers
(adapted from Hollowell and Reynolds 2005).
occupancy (AOO) were generated, and the number of localities analyzed
that were within protected areas was determined employing GeoCat
Bridge in ArcMap 1.5 on ArcGIS (ESRI, New York).
Taxonomic History
A complete nomenclatural history and classification of
Cuphea is available in Graham (1968) and subsequent sectional
revisions (Graham 1988, 1989, 1998, 2017, 2019 in press;
Graham and Cavalcanti 2013). Section Amazoniana (Lourteig)
Lourteig (1986) is the newest section of the genus, the other
twelve sections dating mainly to studies by Koehne (1877) for
Flora Brasiliensis. The one exception is sect. Ornithocuphea
(Koehne) Bullock, now a synonym of sect. Diploptychia Koehne
(Graham 1998).
Lourteig (1959) initially established Cuphea section Euandra
Koehne subsection Amazonianae Lourteig, based on collections
made chiefly by José Cuatrecasas, Richard Schultes, and
Bassett Maguire in Colombia, Venezuela, and Guyana in the
1950s (Lourteig 1954, 1955, 1957; Maguire et al. 1953). The
subsection comprised 11 species previously described and
classified in sect. Euandra subsect. Hyssopocuphea Koehne, two
by Koehne (1877) and nine by Lourteig (1954, 1957, 1959).
Subsequently, Lourteig (1964) added two new taxa to subsect.
Amazonianae, C. blackii Lourteig and C. rhodocalyx forma setosa
147
Lourteig. In 1986, Lourteig, without comment on the change of
status, elevated subsect. Amazonianae to the rank of section and
expanded it to include three new species, C. bolivariensis
Lourteig, C. curiosa Lourteig, and C. killipii Lourteig. Cuphea
rhodocalyx forma setosa Lourteig (1964) and C. rhodocalyx var.
setosa Lourteig (1986) became replaced synonyms for the new
name, Cuphea cardonae Lourteig, and Cuphea rigidula Benth.
was transferred to sect. Amazoniana from sect. Heteranthus
Koehne (Lourteig 1986). In the final study by Lourteig (1996),
the section grew by addition of C. chiribiquetea Lourteig, C.
maigualidensis Lourteig, C. insolita Lourteig, and C. curiosa var.
oresbia Lourteig.
At initiation of the present revision, the section comprised 20
species. With completion, 20 species are recognized, of which
two, Cuphea saxatilis S. A. Graham and Cuphea trisperma S. A.
Graham, are new. Cuphea odonellii Lourteig is transferred to sect.
Trispermum Koehne (See “Species Excluded from Sect. Ama-
zoniana”). Cuphea killipii Lourteig is placed in synonymy of C.
philombria, and C. curiosa var. oresbia is subsumed in C. curiosa.
Morphology And Other Comparative Aspects
Habit—Species of sect. Amazoniana range in habit from
decumbent or creeping to mainly erect, woody subshrubs
10 cm to 2 m in height. The vegetative structure is strongly
 148 SYSTEMATIC BOTANY
influenced by the challenging habitats the plants occupy: on
rocks in and along the margins of fast-flowing rivers; in open,
sunny, wet or dry savannas, on thin soils of granite outcrops;
on nutrient-poor white quartzite sandstone; and in acid peats.
The most common habit is a much-branched, woody subshrub
about 50 cm tall, with small, linear to lanceolate, coriaceous
leaves. Leaves and tiny flowers are often clustered toward the
apex of the stems and stems become leafless proximally with
age. The same traits are common in other endemic genera of
the open shrubland communities of the Guiana Shield region,
where the section is well represented (Grande et al. 2012).
Cuphea kubeorum Lourteig, with prostrate stems 10–45 cm long,
is one of the smallest species of the section, and the only one
described as a “low creeping herb” (Madri~ nán et al. 1121, MO).
Underground Structures—All species of the section are
perennial, and as far as known, all produce woody rhizomes
that generate a mass of fibrous roots. The extensive root system
forms a strong, effective anchor for stems against rushing river
waters and shifting sands, allowing many of the species to
thrive in the cracks and crevices of granitic river boulders and
on soft white quartzite sands in shrublands and savannas.
Stems—Stems are circular in transverse section and covered
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primarily with bifid, appressed trichomes, which are usually
accompanied by one or more other trichome types. Exceptions
occur in Cuphea rhodocalyx Lourteig, and to a lesser degree in
C. cardonae Lourteig, where youngest stems are quadrangular
in transverse section. Dichotomous branching occurs in two
species, C. rigidula and C. blackii.
Leaves—Leaves are decussate, thickly membranaceous to
coriaceous, elliptic to lanceolate, linear-lanceolate, or linear.
They spread outward horizontally from the stem, or are
partially to fully ascending on the stem, becoming closely
imbricate as internodes become progressively shorter distally.
The leaves of C. beneradicata and C. kubeorum are ericoid in
aspect, and in C. kubeorum, particularly, there is a tendency
toward production of short branchlets or fascicles. The ma-
jority of species have linear leaves less than 30 mm long that
appear uni-nerved on the distal portions of stems. Larger,
more proximal leaves display two to five sets of secondary
veins. Secondary venation is less visible on narrow, coriaceous
leaves. Three species from Venezuela, C. dactylophora Koehne,
C. pleiantha Lourteig, and C. rigidula have exceptionally large
elliptic to lanceolate leaves, to 45 mm long. Indumentum,
present on both leaf surfaces, varies among the species in type
and abundance; it can be the same or different on opposing
surfaces. Cuphea rigidula is unique in the section for an
abundantly brown-setose adaxial leaf surface; leaves of C.
philombria are setose to a lesser extent. Leaves in the majority of
species are exclusively bifid-strigose on both surfaces.
Trichomes—In all species of Cuphea there is at least one type
of trichome present, but often two or more types occur
intermixed on stems, leaves, and flowers (e.g. Fig. 2A, B, G). In
sect. Amazoniana, bifid (malpighiaceous), cystolithic trichomes
compose the principal indumentum. The trichomes are lon-
gitudinally oriented and closely appressed to the plant surface
(Fig. 2G–J) or have two semi-erect arms held at approximately
45° angles to the surface (e.g. Fig. 2D). Microcrystals are visible
on the trichome under magnification (Fig. 2G, H, I). The
common indumentum is referred to in this treatment for clarity
and brevity as “bifid-strigose.” Lourteig (1986) referred to it as
“malpighiaceous-pubescent” or “pubescent.” The bifid tri-
chomes differ subtly from species to species in length, thick-
ness, and abundance. Some are so small as to be nearly
[Volume 44
invisible without magnification (e.g. 0.2 mm long in C. boli-
variensis). Others are longer and very thin (e.g. 0.5 mm long in
C. curiosa). Opposing arms of the trichome typically differ in
length, with the distally-oriented arm longest.
Other trichome types are often intermixed with the typical
bifid type, forming a secondary hirsute or, rarely, a setose
indumentum. Setae are more substantial, multi-celled tri-
chomes (Fig. 2B, G). Several species in the section produce a
mix of coarse setae and bifid trichomes on the stems: C. cardonae,
C. cataractarum, C. philombria, C. rigidula, and C. sucumbiensis
Koehne. Cuphea philombria also mixes bifid trichomes and setae
with a sparser indumentum of long, thin, erect to lax, glandular
or non-productive trichomes (Fig. 2G). The combinations of
trichome types vary among the species and are not necessarily
uniform on all organs. For example, the stems of C. insolita are
setose and hirsute (Fig. 2B, E), whereas the leaves have only a
few thin trichomes or are glabrous.
Bifid trichomes have evolved in parallel several times in the
genus and are a conspicuous feature in two other sections
besides sect. Amazoniana, in the Central and South American
section Heteranthus and in the primarily Mexican section
Leptocalyx Koehne (Amarasinghe et al. 1991). The orientation
of the trichome arms on stems in those sections is either
transverse or longitudinal depending on the species, in con-
trast to the consistently longitudinal arrangement in sect.
Amazoniana (Lourteig 1988; Graham 1989).
Three species in the section have a unique indumentum;
they lack the normal bifid-strigose type characteristic of the
section. In Cuphea stygialis Lourteig, one arm of the cystolithic
trichome is so reduced as to be nearly or completely absent
(Fig. 2I, J). In C. insolita, the primary indumentum is hirsute,
consisting of simple, semi-erect trichomes without a second
arm or visible microcrystals (Fig. 2B, E). In C. chiribiquetea
Lourteig, the indumentum consists of smooth oblong to ovoid
glands filled with amorphous contents (Fig. 2C, F). An ab-
breviated arm occasionally develops at one end of the gland.
Inflorescence—The basic inflorescence in the section is a
terminal, simple or compound raceme. It can be indistinct, the
leaves gradually diminishing in size toward the stem apex, or
distinct, the vegetative leaves more or less abruptly reduced
distally on the stem to small bracts. Indistinct vs. distinct
racemes are best distinguished at full development.The di-
chotomous branching pattern of stems in C. blackii and
C. rigidula also appears in their inflorescences.This character is
also diagnostic for the morphologically similar sect. Heteranthus,
but the bracts of the inflorescence in sect. Heteranthus are more
distinctly hypsophylloid than in sect. Amazoniana, and the
flowers differ, arranged as offset pairs with one flower maturing
before the other.
Floral Morphology—The tubular flowers of the section are
among the smallest in the genus, 2.5–5 (rarely –9) mm long,
including in some of the species an incurved, “descending”
spur ca. 0.5–1.5(–2.0) mm long. The outer surface of the floral
tube is green, rose, or purple. Bifid trichomes are usually
present. Hirsute and/or setose indumenta occur infrequently.
Spurs—The extent to which the floral tube develops a
nectar-holding spur at the base varies among species. It is well
developed in the species endemic to Colombia/Ecuador. In
the Colombian Cuphea beneradicata (Fig. 7A) the spur is dis-
tinctly enlarged dorsally and can reach 2 mm in length. In
species endemic to Venezuela, Guyana, and French Guiana
(e.g. in C. bolivariensis and C. cardonae (Fig. 10A, B), the spur is
poorly developed or the floral tube is merely rounded at the
 2019] GRAHAM: CUPHEA SECTION AMAZONIANA 149
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Fig. 2. Stem trichomes in selected species of Cuphea sect. Amazoniana, SEM photos. A. Cuphea beneradicata: bifid cystolithic trichomes of strigose
indumentum mixed with longer, thin trichomes of hirsute indumentum; Idrobo & Zarucchi 9.431 (COL, MO), scale bar 5 500 mm. B. C. insolita: multicellular,
glandular setae mixed with underlying hirsute indumentum of simple, semi-erect, non-glandular trichomes; Hoffman & Marco 2260 (MO), scale bar 5
200 mm. C. C. chiribiquetea: indumentum of sessile ovoid to oblong glands; Franco 3282 (NY), scale bar 5 200 mm. D. C. beneradicata: semi-erect, bifid,
cystolithic trichomes; Idrobo & Zarucchi 9.431 (COL, MO), scale bar 5 250 mm. E. C. insolita: simple, semi-erect, non-glandular trichomes; Hoffman & Marco
2260 (MO), scale bar 5 200 mm. F. C. chiribiquetea: close-up of sessile ovoid to oblong glands; Franco 3282 (NY), scale bar 5 50 mm. G. C. philombria: cystolithic
bifid trichomes of strigose indumentum mixed with longer, thin trichomes of hirsute indumentum and coarser, glandular setae; Idrobo & Schultes 1062 (MO),
scale bar 5 200 mm. H. C. philombria: close-up of bifid, cystolithic, appressed trichomes; Idrobo & Schultes 1062 (MO), scale bar 5 200 mm. I. C. stygialis: one-
armed, cystolithic, appressed trichomes; Schultes & Cabrera 15330 (COL), scale bar 5 100 mm. J. C. stygialis: one-armed, cystolithic, appressed trichomes;
Schultes & Cabrera 15330 (COL), scale bar 5 200 mm.
 150 SYSTEMATIC BOTANY [Volume 44

base and not, or only slightly, extended beyond the pedicel
attachment. The well-developed descending spur of Cuphea
dactylophora (Fig. 12B) varies in length from 0.4 to 1 mm long,
on a 6–9 mm long floral tube.
Petals—Six equal to subequal petals typically are produced
in the section. The two dorsal petals are slightly smaller to
conspicuously smaller than the four ventral ones. All tend
to fall away easily and early. The full complement is reduced to
two or three in C. insolita. Petal color ranges from deep to pale
purple, pink, white, white with a purple base; or petals are bi-
colored, the two dorsals darker purple than the four ventrals.
Knowledge of petal color, size, and shape is incomplete due to
lack of recorded color data in the field and to loss of true color
on heat-dried and alcohol-preserved specimens. Variation,
especially in tone, differs among populations in some species,
judging by conflicting reports on collection labels.
Sepals and Epicalyx—The sepals, i.e. the distal lobes of the
floral tubes, are equal to subequal in size. In species from
Venezuela, and in some species from Colombia, the dorsal-
most sepal tends to be slightly larger or wider than the other
five sepals. In Cuphea rigidula, from Venezuela/Brazil, the
dorsalmost sepal is distinctly enlarged and sepals flanking it
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sepals at the apices of the floral tube veins. Most extend to half
the length of adjacent sepals. They are especially noticeable in
sect. Amazoniana because they tend to be darker purple or
green than the sepals.
Androecium and Gynoecium—The reproductive organs in
sect. Amazoniana are typical for the genus. The androecium in
Cuphea consists of eleven stamens of which the two dorsalmost
are more deeply inserted in the tube than the other nine. The
insertion level is apparent externally as a narrow contraction at
the “neck” of the tube. In sect. Amazoniana, stamens are in-
cluded, most extending no farther than the sinuses of the
sepals and no stamens are well-exserted beyond sepal apices.
The gynoecium consists of a superior, bilocular ovary. The
single style typically remains included or ultimately ex-
tends 1–2 mm past sepal apices (see Reproduction). Cuphea
bolivariensis, C. curiosa and C. saxatilis have a longer style.
Protandry (pers. obs.) suggests self-fertilization is the primary
system of reproduction in the section. The majority of species
produce 4–8 ovules. Cuphea bolivariensis, C. distichophylla
Lourteig, and C. trisperma produce just 3 ovules. Cuphea
philombria and C. sucumbiensis produce the highest number,
8–15 ovules.

Discs—The “disc” (Koehne 1903), is a fleshy, globose or

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are intermediate in size between the dorsalmost and the three

ventral sepals. elongate organ in the floral tube external to the base of the
Epicalyx segments are cylindric or slightly flattened oblong ovary (Fig. 3). It is sometimes described as a nectary and the
lobes, arising external to and slightly below the sinuses of the tissues composing the organ react to staining in the same way

Fig. 3. Discs in the floral tubes of selected species of Cuphea, sect. Amazoniana. A. Cuphea philombria: sub-triangular disc, semi-deflexed, positioned
outside and at base of pilose ovary, partially enclosed by part of the opened hirsute/setose floral tube; Idobro & Schultes 1062 (MO), scale bar 5 0.5 mm. B. C.
rhodocalyx: oblong disc, erect, outside and at base of opened, thin capsule wall and base of the split floral tube, digitate placenta to the right, an immature,
detached seed visible under the placenta; Leisner 24871(MO), scale bar 5 1 mm. C. C. curiosa: cylindric disc, reflexed over pilose ovary, covered above by part
of the opened floral tube; Wurdach & Monachino 41073 (MO), scale bar 5 0.5 mm. D. C. chiribiquetea: globose disc, semi-deflexed to erect, outside and at base of
the flattened ovary to the right; Franco 3282 (NY), scale bar 5 200 mm.
 2019] GRAHAM: CUPHEA SECTION AMAZONIANA 151

that proven nectaries in other genera of the family react (Tobe
et al. 1998). Nectar, however, exudes from the fleshy base
underlying the disc, rather than from the surface of the disc. In
sect. Amazoniana, the disc is usually dark red or purple-red in
color; it is most often pale cream or light green elsewhere in the
genus. Discs vary in shape and position in the floral tube.
Neither the shapes nor positions are narrowly defined, but
merge to some degree. For descriptive purposes there are
approximately four basic disc shapes: 1) sub-triangular in
outline, sub-deltoid, wide at base, tending toward sub-globose
in some species (Fig. 3A); 2) oblong in outline, slightly flat-
tened in transverse section, essentially a short, flattened cyl-
inder with obtuse apex (Fig. 3B); 3) cylindric, elongate,
merging to oblong when shorter and slightly flattened (Fig.
3C); and 4) sub-circular in outline, globose to sub-globose,
merging to subtriangular in outline or subdeltoid (Fig. 3D).
The position of the disc has been regarded as an important
taxonomic character. Lourteig (1986) grouped species of the
section based on three disc positions: deflexed, sub-horizontal,
and erect. I recognize three positions: 1) horizontal to deflexed,
the disc oriented in the longitudinal plane of the tube to de-
flexed downward toward the pedicel (Fig. 3A); 2) erect or
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C. distichophylla from Venezuela and C. insolita and C.blackii
from the Guianas (Table 1). Discs erect and reflexed are correlated
with cylindric to oblong discs in species endemic to Venezuela
(i.e. C. cardonae, C. cataractarum, C. curiosa (although variable),
C. maigualidensis, C. pleiantha, C. rhodocalyx, and C. trisperma),
and with C. dactylophora and C. rigidula in Venezuela/Guyana.
The correlation of disc position and shape with geographic
distribution suggests evolutionary divergence of the disc
characters along two distinct lineages, one in Colombia, the
other primarily in Venezuela and adjacent areas (Table 1).
Pollen Morphology—The pollen morphology of Cuphea is
unique in the family Lythraceae for exceptional diversity at the
species level, and the diversity has proven informative with
respect to taxonomic classification and evolutionary re-
lationships (Graham et al. 1968, 1985; A. Graham and S. A.
Graham 1971). Generically, the pollen is oblate, triangular to
oval-triangular in polar view and tricolporate; colpi are straight,
meridionally elongated, equatorially arranged, and equidistant;
pores are protruding or non-protruding; and the exine is psilate
or variously sculptured.
In sect. Amazoniana, there are at least two major pollen types.
Type 1 pollen (Fig. 4A–D, I, J) is: oval-triangular in polar view,

suberect, free-standing at the base of the ovary, approximately
perpendicular to the longitudinal plane of the floral tube (Fig.
3B, D); and 3) reflexed, erect to curving over the dorsal side of
the ovary (Fig. 3C). The position of the disc is described here
from pressed, dried flowers in early stages of flowering. The
original position in the fresh flower may be slightly altered
during pressing or changed during capsule dehiscence by the
erection of the placenta.
The disc positions are closely correlated with disc shapes
and the correlates sort geographically, although not exclu-
sively so. Horizontal to deflexed discs are correlated with
globose to sub-globose or sub-triangular discs. The combi-
nation occurs in all species from Colombia, i.e. Cuphea ben-
eradicata, C. chiribiquetea, C. kubeorum, C. philombria, C. saxatilis,
C. stygialis, and C. sucumbiensis, plus C. bolivariensis and
tricolporate, non-syncolpate; with exine low rugulate, less
often scabrate, the rugulae tending to parallel the grain
equator; pores slightly protruding; and grain length 13–22 mm,
average widest diameter 16.5 mm. Type 1 occurs in eight
species: C. beneradicata, C. chiribiquetea, C. kubeorum (Fig. 4A),
C. philombria (Fig. 4B), C. saxatilis (Fig. 4C), and C. sucumbiensis
(Fig. 4D) from Colombia; C. distichophylla from Venezuela (Fig.
4I; and C. insolita from Guyana (Fig. 4J). The shape and exine
sculpture of pollen of C. bolivariensis from Colombia (not il-
lustrated), suggests the grain is also Type I, however, a thick,
irregularly rugulate exine obscures the termination of the colpi
at the poles.
Type 2 pollen (Fig. 4E–H) is: triangular to oval-triangular in
polar view, tricolporate, syncolpate, colpi forming a triangular
opening at the poles enclosing a central exine remnant; exine

Table 1. Morphological variation in selected characters of Cuphea sect. Amazoniana arranged by geographical region. Abbreviations: EC 5 Ecuador;
VEN/GUAY/BR 5 Venezuela, Guayanas, Brazil; hor/defl 5 horizontal/deflexed; gl/s-del 5 globose/sub-deltoid; cyl/obl5 cylindric/oblong; n-syn 5
non-syncolpate; short 5 shorter than subtending leaves; dist 5 distinct; indist 5 indistinct.

Disc Position Disc Shape Pollen Colpi Pollen Exine Spur Internodes Raceme Ovules
COLOMBIA/EC
C.beneradicata hor/defl gl/s-del n-syn rugulate 1 short dist 4–6
C. saxatilis hor/defl s-del n-syn rugulate 1 sshort dist 5–6
C. chiribiquetea hor gl/s-del n-syn rugulate 1 short indist 4–5
C. stygialis hor/defl s-gl ? ? 1 short indist 4–6
C. kubeorum hor/defl gl/s-del n-syn rugulate 1 short indist 4–6
C. philombria hor/defl s-del n-syn rugulate 1 short indist 8–15
C. sucumbiensis hor/defl gl/s-del n-syn rugulate/scabrate 1 short dist 8–14
VEN/GUAY/BR
C. blackii hor/defl s-gl n-syn rugulate – long dist 3–6
C. insolita hor/defl s-del n-syn scabrate – short dist 3–5
C. distochophylla hor/defl s-gl n-syn rugulate/scabrate – short indist 2–3
C. bolivariensis hor/defl s-gl n-syn rugulate – short indist 3
VENEZUELA
C. cardonae refl cyl/obl syn rugulate – short dist 5
C. rigidula erect cyl/obl syn scabrate – short dist 9–10
C. cataractarum erect cyl/obl syn psilate – short indist 4–7
C. curiosa refl cyl syn psilate – short dist 5–9
C. pleiantha refl cyl/obl syn psilate – short dist 7–9
C. rhodocalyx erect obl syn psilate/rugulate – short dist 5–9
C. dactylophora refl cyl/obl syn psilate/scabrate 1 short indist 6–7
C. maigualidensis refl cyl ? ? – long indist 5
C. trisperma refl cyl syn psilate – short indist 3
 152 SYSTEMATIC BOTANY [Volume 44
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Fig. 4. Pollen of selected species of Cuphea, sect. Amazoniana; measurement is the greatest diameter of the grain illustrated. A. Cuphea kubeorum, Schultes &
Cabrera 19913 (P); 17 mm. B. C. philombria, Idrobo & Schultes 1062 (MO); 17 mm. C. C. saxatilis, Echeverry 2089 (COL); 15 mm. D. C. sucumbiensis, Cerón 20763
(MO); 18 mm. E. C. curiosa, Wurdack & Adderly 43590 (NY), with permission of A. Graham; 20 mm. F. C. pleiantha, Maguire & Maguire 35376 (MO); 19 mm. G. C.
trisperma, Steyermark 105451(MO); 20 mm. H. C. rigidula, Cardona 3013 (NY); 19 mm. I. C. distichophylla, Huber 13512 (P); 16 mm. J. C. insolita, Hoffman & Marco
2260 (MO); 17 mm. K. C. dactylophora, Maguire et al. 46715 (NY); 18 mm. L. C. blackii, Croat & Ferry 102998 (MO);15 mm.

typically psilate; pores not or slightly protruding; and grain
length 15–24 mm, average widest diameter 19 mm. Type 2
pollen occurs in seven species: C. cardonae, C. cataractarum,
C. curiosa (Fig. 4E), C. pleiantha (Fig. 4F), C. rhodocalyx, and
C. trisperma (Fig. 4G) from Venezuela, and C. rigidula (Fig. 4H)
from Venezuela/ Guyana. The pollen of C. trisperma (Fig. 4G)
and C. rigidula (Fig. 4H) differ in having raised colpi margins,
and in C. rigidula, a coarser exine.
Pollen of C. dactylophora and C. blackii (Fig. 4K, L) from
Venezuela/Guyana and French Guiana, respectively, possibly
represent a third pollen type or are Type I pollen modified by a
thicker exine.The grains are triangular in polar view, non-
syncolpate, with wide, non-protruding pores, and a thickened,
enlarged polar exine. The pollen types of two species,
C. maigualidensis, Venezuela, and C. stygialis, Colombia, are
unknown.
The Type I non-syncolpate, generally smaller pollen form is
known elsewhere in Cuphea in both subgenera. In subgenus
Bracteolatae, in addition to sect. Amazoniana, it occurs in section
Heteranthus and parts of sections Brachyandra and Euandra (S.
A. Graham and A. Graham 1971; Graham et al. 1985, 2006;
Graham 2017). Externally, it also resembles pollen of species
in sect. Trispermum. Internally, however, the pollen of sect.
Trispermum has distinctive interaperturate thickenings
(S. A. Graham and A. Graham 1971). Phylogenetically, Type I
pollen appears in several clades of subg. Bracteolatae (Graham
et al. 2006; Graham 2017).
Pollen type and geography are correlated in species of sect.
Amazoniana, as they are for disc position, disc shape, spur
development, and geography. Total evidence from these
differing sources supports, at minimum, two major evolu-
tionary lineages within the section, one of which has radiated
in Colombia and the other in Venezuela and regions to the east.
In a wider perspective, in a review of Myrtalean pollen, Patel
et al. (1984), illustrated syncolpate pollen grossly similar to the
Type 2 pollen described above (e.g. Patel et al. 1984: Fig. 41A,
42A). Within the order, only the Myrtaceae and Cuphea are
known to produce this form. The similarity does not infer that
Cuphea is closely related to those members of the Myrtaceae
with similar pollen morphology, but is likely the product of de
novo evolution out of the ancestral, ordinal-level genetic pool.
The similarity is mentioned here as a cautionary note when
identifying fossil pollen of these two families from tropical
America.
 2019] GRAHAM: CUPHEA SECTION AMAZONIANA 153

Seed Morphology—The seeds produced by species of the
section (Fig. 5A–L) are 0.9–1.7 mm long, 0.8–1.5 mm wide,
among the smallest in the genus, and mostly bilateral, with a
convex antiraphal face and flattened raphal face; in C. cardonae
they are sub-deltoid. Seeds vary in outline from cordiform to
obovate, circular, or oblong. All have finely reticulate or
scalariform seed coats. The seed margin is typically thick and
rounded. Cuphea blackii (Fig. 5L) and C. trisperma have a
thinned outer margin, most apparent under light microscopy.
There is some natural variability in shape within a species;
differences in shape also occur due to tight packing within the
capsule, especially if seeds are not fully mature when collected
and pressed. All seeds of the section appear to lack the
prominently enlarged chalazal area and strong raphe typical of
the genus (compare Graham and Graham 2014: Figs. 6, 7). The
total morphological variation in the section is not extensive
compared to the rest of the genus, and seeds do not sort into
discrete groups, as the pollen does. The embryos, which fully
fill the interior of Cuphea seeds, produce quantities of oil
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dominated by saturated short to medium chain fatty acids. The
oil composition of species in sect. Amazoniana is unknown
(Graham et al. 2016).
Reproduction—The stamens and styles of all species in sect.
Amazoniana are included at anthesis. Based on extensive
personal observations of living collections of Cuphea (none
from sect. Amazoniana), small flowers, such as occur in sect.
Amazoniana, are primarily self-pollinated. The versatile an-
thers encircling the distal portion of the floral tube face in-
wards toward the centrally-placed style and stigma. At anther
dehiscence, the style elongates passing through the circle of
anthers and the stigma, now receptive, is dusted with self-
pollen. The stigma typically remains within the floral tube or it
may become minimally exserted as seeds mature, but it is then
no longer receptive (pers. obs.). In some other sections, es-
pecially where the flowers are larger and more brightly col-
ored, the style elongates several millimeters beyond the
margin of the floral tube prior to stigma receptivity, and cross
pollination is the primary means to fertilization. No living

Fig. 5. Seeds of selected species of Cuphea, sect. Amazoniana, anti-raphal faces; measurement is the length of the seed illustrated. A. Cuphea beneradicata,
Cárdenas et al. 6514 (MO); 1.2 mm. B. C. kubeorum, Schultes & Cabrera 14267 (MO); 1.2 mm. C. C. philombria, Idrobo & Schultes 1062 (COL); 1.3 mm. D. C. saxatilis,
Echeverry 2089 (COL); 1.7 mm. E. C. cataractarum, Maguire et al. 30438 (MO); 1.2 mm. F. C. curiosa Steyermark et al. 107107 (P); 0.9 mm. G. C. pleiantha, Maguire &
Maguire 35102 (NY); 1.1 mm. H. C. rhodocalyx, Leisner 24871(MO); 1.0 mm. I. C. rigidula, Cardona 3013 (NY); 1.5 mm. J. C. insolita, Hoffman & Marco 2260 (MO);
1.5 mm. K. C. dactylophora, Berry et al. 6950 (MO); 1.3 mm. L. C. blackii, Croat & Ferry 102998 (MO); 1.5 mm.
 154 SYSTEMATIC BOTANY
collections from sect. Amazoniana were available for study.
Flowers of most Cuphea species appear to be promiscuously
pollinated by diverse small bees, butterflies, and day-flying
moths. Larger-flowered species attract larger Bombus bees and
hummingbirds; they are also common victims of small nectar-
robbing bees that puncture the sides of the nectar-holding
spur.
Distribution and Ecology—The species of sect. Amazoniana
are a common component of the floras in the Guiana Shield
and outliers of the Shield in Colombia, occurring along the
extensive river systems, on tepuis, in savannas, and less often
in forests of the regions. Only Cuphea sucumbiensis and
C. philombria lie to the west and south of the Shield in the sub-
Andean premontane to montane forests of southwestern
Colombia and northern Ecuador (Fig. 16).
In Venezuela and western Guyana, the species occur on
high, insolated, wind-swept plateaus and slopes of tepuis, e.g.
on Sierra de la Neblina, Mt. Roraima, Sierra Maigualida,
Serranı́a Yutajé, and Cerros Sipápo, Marahuaka, and Duida; in
La Gran Sabana; and, most likely, in other under-explored
mountains in the west-central and southern part of the Guiana
Shield (Maguire et al. 1953; Huber et al. 2018). Species of
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Amazoniana also occur in cool temperate and tropical montane
forests on mountain slopes, and in the lowlands of Amazonia.
Cuphea blackii is the easternmost representative of the section,
and the only one in French Guiana and adjacent Amapá, Brazil
(Fig. 9). It is unique in the section for occupying primary forest
habitats at low elevations from nearly sea level to 400 m. No
species in the section has been reported for the flora of Sur-
iname (Funk et al. 2007).
In Colombia, outliers of the Guiana Shield, such as the
Sierra de la Macarena, Serranı́a La Lindosa, and Sierra de
Chiribiquete, support Cuphea beneradicata, C. chiribiquetea,
C. kubeorum, and C. saxatilis on savannas and along numerous
tributaries of Guayabero and Guaviare rivers at elevations
from about 300 to 650 m. Cuphea philombria and C. sucum-
biensis inhabit the eastern sub-Andean forests. Cuphea sty-
gialis lies at low elevations (100–300 m) in tropical Amazonian
Vaupés.
The total range of elevation above sea level for sect.
Amazoniana is 10–2100 m. In Venezuela, nine species grow at
elevations from 100 to 2100 m, seven occurring above 1500 m,
with C. maigualidensis highest at 2100 m. In Colombia, seven
species occur from 200–1900 m; all seven are present below
700 m; and two reach 1500 m and above. Cuphea philombria and
C. sucumbiensis exhibit the widest tolerance in elevation,
C. philombria occurring along rivers and in montane forests
from 300–1900 m. It is found from 300–625 m and again at
1000–1900 on Sierra de la Macarena. Cuphea sucumbiensis oc-
curs from 50–1770 m in Colombia and 1500–1700 m in sub-
Andean Ecuador. Cuphea curiosa, in Venezuela, extends from
110–2100 m with a gap in collections between 830 and 1400 m.
The distribution of species in sect. Amazoniana may well be
underestimated because many parts of the extensive area they
occupy are unexplored or underexplored. The majority of
species in the section are narrow endemics, especially in
Venezuela where Cuphea insolita, C. maigualidensis, and
C. trisperma are known only from the type and C. bolivariensis,
C. rigidula, and C. stygialis are known from the type and one
additional collection.
The high diversity and endemicity of species in the section is
the result of the combination of at least three factors: the ability
to disperse seeds widely by water; access of seeds on dispersal
[Volume 44
to diverse habitats, thus enhancing the chance of plant es-
tablishment; and reproduction following self-pollination.
Plants are advantageously situated for seed dispersal along
rivers, by erosional outwash from mountain slopes, and by
seasonal flooding of the savannas bordering rivers. The to-
pology and physiography of the Guiana Shield in Venezuela
and the outlying fragments in Colombia, such as the sierras of
La Macarena, La Lindosa, and Chiribiquete, provide a variety
of micro-topographic habitats varying in temperature, expo-
sure, depth of organic accumulated organic matter, and water
levels (Huber 1988, 1995, 2005). The species are established on
both dry grassy and seasonally inundated savannas; in open
shrublands; in white-sands and herbaceous meadows; in
cracks and crevices of granitic rock outcrops; and along rocky
or sandy river margins.
Risk of Extinction—A risk of extinction is estimated for
each species according to the categories and criteria estab-
lished by the IUCN Red List (IUCN 2012). Exsiccatae from
distinctly separated localities provided estimates in km2 of
extent of occurrence (EOO) and area of occupancy (AOO). Of
20 species, nine are estimated as critically endangered (CR),
five as endangered (EN), three as vulnerable (VU), and three
are of least concern (LC), being geographically widely dis-
tributed, some in protected areas, and relatively common and
well-collected. Cuphea sucumbiensis (LC), has the greatest EOO
and AOO in the section. It differs from the two other species of
least concern in that only one of the 15 collections studied is
within a protected area. As a group, the species are perhaps
most vulnerable to changes in climate toward drier, warmer
environments due to their association with abundant seasonal
waters.
Morphological Relationships—Insights into evolutionary
relationships among members of the section and with other
sections or clades of the genus are limited to morphological
comparisons. The sect. Amazoniana was based on presence of
bifid (malpighiaceous) trichomes, leafy inflorescences, and
gibbous or spurred floral tubes (Lourteig 1959, 1964). Char-
acters that further define the section are the exceptionally small
flowers, 9 mm long or less, mostly 3–5 mm long, and included
stamens deep set in the floral tube. This character suite also
partially defines sections Heteranthus and sect. Trispermum,
with the important exception that in sect. Trispermum bifid
trichomes are recorded in only one speces, C. odonellii. Geo-
graphically, sect. Heteranthus in South America is distributed
along Andean slopes to the west of sect. Amazoniana, and sect.
Trispermum occurs widely in South America, being especially
species-rich in Brazil. Bifid trichomes are characteristic also of
the Mexican sect. Leptocalyx Koehne, and appear in one to few
species of five other sections that are not otherwise morpho-
logically close to sect. Amazoniana (Amarasinghe et al. 1991).
At present, the absence of an independent molecular data set to
test hypotheses of relationship from morphological evidence
leaves speculative the wider relationships of a clearly para- or
polyphyletic sect. Amazoniana.
Within sect. Amazoniana, position and shape of the disc,
pollen morphology, and morphology of the floral spur cor-
relate closely with two geographic areas, suggesting the sec-
tion consists of a minimum of two distinct lineages, one in the
Guiana Shield region in Venezuela and the other in Colombia
and southern sub-Andean Colombia/Ecuador. Each lineage
has produced a number of relatively homogeneous endemic
species that occupy comparable riverside, savanna, and forest
habitats in their region (Table 1).
 2019] GRAHAM: CUPHEA SECTION AMAZONIANA
In Colombia, seven species of the section (C. beneradicata,
C. chiribiquetea, C. kubeorum, C. philombria, C. saxatilis, C. stygialis
and C. sucumbiensis) are united morphologically by: distinct
floral spur development; dorsalmost sepal not conspicuously
larger than the others; discs globose to sub-triangular, hori-
zontal to deflexed; and non-syncolpate pollen (pollen mor-
phology unknown in C. stygialis).
In Venezuela, nine species of the section (Cuphea cardonae,
C. cataractarum, C. curiosa, C. dactylophora, C. maigualidensis
(pollen unknown), C. pleiantha, C. rhodocalyx, C. rigidula and
C. trisperma) are united by: spurs poorly developed to none
(except in C. dactylophora), the base of the floral tube horizontal
and rounded to very slightly extended (gibbous) past the
attachment of the pedicel; dorsalmost sepal largest; discs cy-
lindric to oblong, erect to reflexed over the ovary; and syn-
colpate pollen. The floral tubes are, overall, longer than those
of the Colombian species and appear more distended and
ampullaceous in fruit, although the ranges in length of the
floral tube overlap in Colombian and endemic Venezuelan
species, as do ovule numbers.
The relationships of the remaining species, viz., C.
distichophylla and C. bolivariensis in Venezuela and Brazil, C.
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insolita in Guyana, and C. blackii in French Guiana and Brazil,
are less clear. They are morphologically part of the Colom-
bian lineage based on equal-sized sepals, disc shape and
position, and non-syncolpate pollen, whereas the short to
non-spurred floral tubes are like those in the Venezuelan
species group.
Total morphological and geographic information suggests
that a lineage in Colombia\Ecuador and a second in Ven-
ezuela and adjacent regions independently speciated in the
recent history of the genus, forming two separate, relatively
homogenous groups of species. Each encompasses some
variation in branching habit, type and abundance of stem and/
or floral indumentum, shape and size of leaves, and degree to
which the inflorescence is distinct from the vegetative stem.
Additional lineages, possibly from Brazilian sources, such as
sect. Trispermum, should also be considered in future in-
vestigations of ancestral relationships.
Fossil History—The fossil record of Cuphea is based on
several reliably identified fossil pollen grains from North and
South America. The earliest known date from the early to
middle Miocene of Chiapas, Mexico and the late middle
Miocene of Trinidad (Graham 2013). Two grains from younger
deposits in northwestern South America bear a general re-
semblance to Types 1 and 2 in sect. Amazoniana. Type 1 (Fig.
6A) is from Meta, Colombia, at mid to low elevation sites of
Mio-Pliocene and Holocene ages. Type 2 (Fig. 6B) is from an
unpublished locality in northern South America of Mio-
Pliocene age (Wymstra and van der Hammen 1966; A.
Graham and S. A. Graham 1971: Figs. 1 and 8, copied here as
Fig. 6A, B; Graham 2013: Fig. 2a–e). Although not strictly
identical to pollen in sect. Amazoniana, the grains bear some
similarities, one being small, non-syncolpate, and psilate, the
other syncolpate with a polar triangular opening. They
demonstrate presence and diversity of the genus from at least
14 MYA in northwestern and north-central South America in
regions that harbor species of sect. Amazoniana today.
Speciation and Radiation—The expansion of the lineages
composing sect. Amazoniana is hypothesized to have occurred
beginning in the late Miocene, based on the presence of multiple
fossil pollen records (Berrio et al. 2002; Graham 2013). Today, the
lineages are adapted to diverse habitats within the four floristic
155
Fig. 6. Fossil pollen grains of Cuphea from northwestern South
America. A. Pollen of Pleistocene-Holocene age, Laguna de Agua-Sucia,
Meta, Colombia; actual size ca. 16 mm in greatest diameter. B. Pollen of
Mio-Pliocene age, northern South America; actual size ca. 18–20 mm in
greatest diameter. (From A. Graham and S. A. Graham 1971).
regions of the Guiana Shield: the Andean, Amazon, Pantepui,
and Caribbean regions (Huber et al. 2018; only C. blackii occurs in
the Caribbean biota). The lineages may have been present in
northern South America at the global Miocene Climatic Opti-
mum (MCO), and were present later in the Middle Pliocene
Climatic Optimum (MPCI, 3.3–3 MYA), when seasonality was
increasing and climates were becoming cooler and drier.
Diversification and radiation of the lineages in sect. Ama-
zoniana involved both morphological and physiological ad-
aptations to the mosaic of habitats composing the Guiana
Shield. The shrubby habit, small, thick coriaceous leaves, and
diverse kinds and densities of indumentum, common to
Cuphea and other shrubland genera in that region, whether
already in place in Cuphea or developing in response to the
changes, would have enhanced the ability of the plants to
withstand alternating wet and dry seasons and more extreme
fluctuations in temperature (Huber 2005), and favored their
diversification into the enlarging areas of drier shrublands and
savannas they commonly occupy today. In the period fol-
lowing the MPCI, dynamic cycling of 15–20 interglacial-glacial
fluctuations furthered topographic, geologic, and climatic
changes (Graham 2010) and must have influenced the rate of
species diversification and increased the ecological tolerances
to those now observed in the extensive altitudinal ranges of
many plant species (Huber 1988), including several in Cuphea
sect. Amazoniana, as noted earlier. Physiological adaptations to
nutrient-poor habitats, particularly for species now endemic to
granite outcrops and white sand savannas in the Guiana
Shield, were necessarily implemented as the lineages came to
occupy these specialized sites throughout the Quaternary to
present time. Distribution of the several species of the section
adapted to river margin habitats would have been fostered by
the extensive river systems present throughout the large
Orinoco River basin (León and Dı́az 1976).
Taxonomic Treatment
CUPHEA P. Browne, Civ. Nat. Hist. Jamaica 216. 1756. TYPE:
Cuphea decandra Aiton, Hort. Kew, ed. 2. 3: 3,151. 1811
(Graham 1968, 2001; Molero and Zijlstra 1999).
Annual or perennial herbs or subshrubs; stems commonly
with one or more types of indumentum. Leaves opposite and
decussate. Inflorescences leafy to bracteate racemes, one
flower at a node positioned between opposing petioles, ad-
ditional flowers at a node produced on axillary branchlets.
Flowers bilateral, tubular, 6-merous, conspicuously 12-veined;
 156 SYSTEMATIC BOTANY [Volume 44

floral tube perigynous, persistent, base rounded or short-
spurred (gibbous) to long-spurred, sepals 6, at apex of tube,
alternating with 6 epicalyx segments. Petals 0–6, crinkled,
with a midvein. Stamens 11, the 2 dorsalmost shortest and
most deeply inserted in tube. Ovary incompletely bilocular,
dorsal locule reduced; style one, stigma small, punctiform or
capitate; disc external to and subtending the ovary at base of
floral tube. Fruits dry, thin-walled capsules, enclosed in the
floral tube; dehiscence by complementary dorsal longitudinal
slits in capsule and floral tube, on dehiscence placenta and
attached seeds exserted through the dorsal slits. Seeds 2 to
many, small.
CUPHEA P. Browne subgenus CUPHEA
Pedicels lacking bracteoles.
CUPHEA subgenus BRACTEOLATAE S. A. Graham, Syst. Bot.
Monogr. 20: 42. 1988. Lectotype, designated by Graham
(1988, 2001): Cuphea viscosissima Jacquin.
Pedicels bibracteolate, rarely bracteoles reduced to tufts of
trichomes.
CUPHEA subg. BRACTEOLATAE sect. AMAZONIANA (Lourteig)
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Lectotype designated (as holotype) by Lourteig,
1959: Cuphea cataractarum Koehne.
Subshrubs 25–100 cm, mostly less than 50 cm; stems one to
several from a woody primary root or rhizome with abun-
dant fibrous roots, much-branched; indumentum basically
strigose, trichomes bifid, appressed, often mixed with hirsute
or setose indumentum, rarely stems, leaves, and/or flowers
glabrous. Leaves decussate, sessile or subsessile, spreading
or crowded and ascending, often imbricate distally on the
stem, elliptic-lanceolate to linear-lanceolate, linear, or oblong,
thickly membranaceous or coriaceous, frequently appearing
uni-nerved. Inflorescences racemes, leafy or bracteate,
simple or compound. Flowers with floral tube 2.5–7(–9) mm
long, margin flaring, base rounded, non-spurred or extended
as a spur to 2 mm long, spur horizontal or incurved,
descending; sepals equal to subequal, or the dorsalmost sepal
larger than the other 5, or the ventral 3 sepals slightly larger;
epicalyx segments thick lobes or cylinders at sinuses of se-
pals; internal surface of floral tube glabrous, villous, or pilose.
Petals 6, various shades of purple, rose, or white, dorsal 2
petals generally smaller than ventral 4. Stamens 11, the 2

Lourteig, Phytologia 60:17. 1986.
Cuphea sect. Euandra subsect. Hilairea Koehne, in
Mart. fl. bras. 13(2): 226. 1877, p.p. excluding
type of subsect. Hilairea.
Cuphea sect. Euandra subsect. Hyssopocuphea Koehne,
Bot. Jahrb. Syst. 1: 441, 152. 1881, p. p., ex-
cluding type of subsect. Hyssopocuphea.
Cuphea section Euandra subsect. Amazonianae Lourteig,
Notas del Museo, Univ. Nac. La Plata 19: 280. 1959.
dorsalmost shortest and inserted most deeply, included;
pollen tricolporate, colpi not united or colpi united in an open
triangle at the poles, exine psilate, rugulate or scabrate.
Ovules 2–15, mostly 4–8. Disc cylindric, or slightly flattened,
sub-triangular or oblong in outline, or sub-globose to glo-
bose, sub-circular to circular in outline; disc position reflexed
over the ovary, or erect at right angles to the longitudinal
plane of the floral tube, or horizontal to deflexed into the
spur. Seeds 0.9–1.7 mm long, bilaterally compressed, obovate
to circular or oblong in outline, finely to coarsely reticulate or
scalariform.

Key To The Species Of Cuphea Sect. Amazoniana

1. Disc at base of ovary erect at right angles to longitudinal plane of floral tube or reflexed over the ovary, cylindric or oblong; pollen colpi united, forming
an open triangle at the poles (syncolpate).
2. Branching dichotomous; ovules 9–10; leaf adaxial surface bifid-strigose and abundantly brown-setose. 16. C. rigidula Benth.
2. Branching monopodial; ovules 3–9; leaf adaxial surface exclusively bifid-strigose, or glabrous, or a mix of bifid and simple trichomes.
3. Younger distal portions of stems quadrangular from decurrent leaf bases; leaf adaxial surface sparsely bifid-strigose or glabrous.
4. Stems glabrous or very sparsely bifid-strigose; disc short, erect; petals bi-colored, the dorsal 2 darker purple than the 4 ventrals or all
uniformly pale purple. 15. C. rhodocalyx Lourteig
4. Stems glandular hirsute to setose; disc elongated, reflexed over ovary; petals white with short dark-purple claw. 4. C. cardonae Lourteig
3. Younger distal portions of stems cylindric; leaf adaxial surface bifid-strigose or bifid-strigose mixed with longer, erect trichomes.
5. Ovules 3; leaves linear, 3–11 mm long; stem trichomes exclusively bifid-strigose. 20. C. trisperma S. A.Graham, sp. nov.
5. Ovules 4–9; leaves elliptic, lanceolate, or lanceolate-linear, (3.5–)10–43 mm long; stem trichomes bifid-strigose mixed with longer, simple, 6
erect trichomes.
6. Racemes distinct, vegetative leaves abruptly reduced to small bracts of the racemes.
7. Leaves ascending, imbricate, 15–60 mm long; floral tubes non-spurred, base not extended past attachment to pedicel; plants sturdy,
mostly 30–50 cm tall. 14. C. pleiantha Lourteig
7. Leaves mostly spreading, 5–40 mm long; floral tubes with spur poorly developed, extending ca. 0.3 mm past attachment to pedicel;
plants slender, mostly 10–25 cm tall. 7. C. curiosa Lourteig
6. Racemes indistinct, vegetative leaves gradually transitioning to bracts of the racemes.
8. Internodes long, mostly equal to or longer than subtending leaves; leaves spreading; disc reflexed. 12. C. maigualidensis Lourteig
8. Internodes less than half as long as subtending leaves; leaves imbricate or spreading; disc erect.
9. Leaves closely imbricate on sparsely branched stems, elliptic, 5–22 mm long, base rounded to obtuse; floral tube non-spurred, or
spur developed to 0.5 mm long. 5. C. cataractarum Spruce ex Koehne
9. Leaves loosely spreading, distanced along branched stems, elliptic-lanceolate, 10–45 mm long, base cuneate; floral tube with
well-developed spur 0.4–1 mm long. 8. C. dactylophora Koehne
1. Disc at base of ovary horizontal in the longitudinal plane of floral tube or deflexed into base of tube, globose, sub-globose, or sub-triangular; pollen colpi
short, not united at the poles (non-syncolpate).
10. Ovules 8–15.
11. Racemes distinct; stems 15–35 cm; seeds 1.0–1.1 mm long. 19. C. sucumbiensis Lourteig
11. Racemes indistinct; stems 30–50 cm; seeds 1.2–1.3 mm long. 13. C. philombria Lourteig
10. Ovules 2–6.
12. Stems bearing ovoid glands or other non-bifid trichome types.
 2019] GRAHAM: CUPHEA SECTION AMAZONIANA 157

13. Stems bearing ovoid, sessile, smooth-surfaced glands with amorphous contents (Fig. 2C, F); older stems glabrous or nearly so; racemes
indistinct. 6. C. chiribiquetea Lourteig
13. Stems bearing other simple or multicellular, non-bifid trichome types; racemes indistinct or distinct.
14. Stems exclusively bearing one-armed, cystolithic, appressed trichomes (Fig. 2I, J); racemes indistinct; floral spur well-developed. 18.
C. stygialis Lourteig
14. Stems bearing a mix of multicellular setae to 1 mm long and simple, single-celled, non-glandular, semi-erect trichomes to 0.2 mm
long (Fig. 2B, E); racemes distinct; floral spur poorly developed. 10. C. insolita Lourteig
12. Stems strigose, bearing exclusively bifid trichomes, or bifid trichomes mixed with simple, semi-erect, longer ones.
15. Floral spur well-developed, subdescending.
16. Racemes indistinct; stems and flowers exclusively bifid-strigose; leaves 2–6(–10) mm long, irregularly crowded to fasciculate
distally on stems. 11. C. kubeorum Lourteig
16. Racemes distinct; stems and flowers bifid-strigose and hirsute with trichomes simple, semi-erect, longer, fragile.
17. Internodes equal to mostly longer than subtending leaves; leaves elliptic- lanceolate, mostly 15–20 mm long; midvein and
secondary veins visible abaxially. 17. C. saxatilis S. A. Graham, sp. nov.
17. Internodes mostly shorter than subtending leaves; leaves linear, 5–11(–20) mm long; appearing uni-nerved. 1. C. beneradicata
Lourteig
15. Floral spur poorly developed or none, base rounded, scarcely extended past pedicel attachment.
18. Leaves half as long as to longer than the subtending leaves, membranaceous; racemes distinct; flowers paired. 2. C. blackii Lourteig
18. Leaves less than half as long as the subtending leaves, coriaceous; racemes indistinct; flowers alternate to subalternate, rarely
paired.
19. Leaves ascending, rigid, strongly distichous distally on the stem, 10–18 mm long, 1.5–3.0 wide, apex acute; stems bifid-
strigose, sometimes mixed with simple, longer trichomes. 9. C. distichophylla Lourteig
19. Leaves spreading and ascending, decussate to irregularly arranged, not strictly distichous distally on the stem, 6.5–12.0 mm
long, 1–2 mm wide, apex obtuse; stems exclusively bifid-strigose. 3. C. bolivariensis Lourteig

1. CUPHEA BENERADICATA Lourteig, Bot. Mus. Leafl. (Harvard shorter than sepals, some with simple, thin, red trichomes at
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University) 16: 226, Fig. 37. 1954. TYPE: COLOMBIA. Gua-
viare: Mesa La Lindosa [2.474, -72.696], 15–20 km de
San Jos é Guaviare, 400–600 m, 13–15 Dec 1950, J. M.
Idrobo & R. E. Schultes 640 (holotype: GH-00068434-
image!; isotypes: COL!, F-image!, MO!, P!, NY!, US32-
images!).
Subshrubs to 30 cm; stems one to several, erect to de-
cumbent from a woody rhizome, much-branched, branch
length and position on the stem irregular; internodes of the
branches very short, compressed, ca. 2–5 mm long, mostly
shorter than subtending leaves; stem indumentum of 2 types:
abundantly bifid-strigose, trichomes thin, short, longitudi-
nally or irregularly oriented on the stems, appressed or some
with arms semi-erect; and hirsute on youngest growth, tri-
chomes simple, thin, glandular, erect to lax, to 0.5 mm long.
Leaves decussate to irregularly crowded distally on the
branches, ericoid in aspect, abruptly reduced in size in the
inflorescence, appearing uni-nerved; petioles 0–1 mm long;
blades linear, 5–11(–20) mm long, 1–3(–5) mm wide, thickly
membranaceous, base cuneate, apex subacute, margin plane to
incurved; surfaces adaxially green, glabrous or sparsely
bifid-strigose with exceptionally minute, thin trichomes,
abaxially paler, bifid-strigose, midvein prominent, brown,
secondary veins visible only on largest proximal leaves.
Racemes distinct, sometimes tending to indistinct, terminal,
simple, bracteate to leafy, bracts lanceolate-elliptic, 2.5–4.0 mm
long, margin sparsely ciliate; flowers alternate, sub-alternate or
opposite, inter- or infra-petiolar; pedicels 1.5–2.5(–5.0) mm long;
bracteoles ovate-acuminate, 0.2 mm long, obscured by abun-
dant strigose indumentum. Floral tube, including a well-
developed, prominently dorsally enlarged, subdescending
spur 0.5–2.0 mm long, 4.0–5.5(–7.0) mm long, 1.0–1.5 mm wide,
neck slightly constricted at 2/3 length of floral tube, mouth
flaring, ca. 2 mm wide; outer surface green, abundantly
bifid-strigose with arms of trichomes sometimes semi-erect,
and sparsely hirsute with simple, thin, reddish trichomes to
0.5 mm long; inner surface white villous; sepals subequal,
0.7–1.0 mm long, 0.3–0.5 mm wide, deltate, apiculate; epi-
calyx segments thickened lobes, 6 cylindric, 0.5 mm long,
apex. Petals 6, pink or light purple, subequal, 2–4 mm long,
1.0–1.5 mm wide, oblong to narrowly obovate. Stamens: 2
short dorsalmost slightly deeper, other 9 inserted in a
nearly horizontal line at 1/2–2/3 length of floral tube,
included, antesepalous stamens extended nearly to sinuses
of sepals, antepetalous ones shorter; filaments white
villous, filament and anther color unknown; pollen oval-
triangular, tricolporate, non-syncolpate, pores not pro-
truding, exine shallowly rugulate to psilate, poles psilate, grain
diam 15 mm. Ovary villous; style villous, included at anthesis;
ovules (4–)6, at distal end of a flattened placenta. Disc hori-
zontal to deflexed, globose to sub-deltoid, ca. 0.5–0.7 mm long,
0.5 mm wide, thick, obtuse, dark red. Seeds 1.0–1.4 mm long,
1.1–1.5 mm wide, circular to slightly obovate, margin raised,
rounded, 0.2 mm wide. Figures 2A, D, 5A, 7A.
Phenology—Collected in flower March to December; in
abundant flower and fruit in August.
Distribution and Habitats—Colombia, Dept. Guaviare, in
the area of San José del Guaviare and the Serranı́a La Lindosa;
on rocks and in wet or well-drained sands on natural savanna
and in stunted forests (achaparrados); 200–600 m (Fig. 8).
Extinction Risk—The estimated risk if all IUCN guides
were to be followed is CR; EOO 5 28.79 km2 and AOO 5
16 km2. Three of the four collection localities analyzed occur in
protected areas.
Additional Specimens Examined—Colombia.—GUAVIARE: San José del
Guaviare, Serranı́a La Lindosa, carr. al Nuevo Tolima, ca. 1.5 km después
entrada a la Ciudad de Piedra, 2°270 39.6”N, 72°420 36.40W [2.461, -72.710],
15 Nov 2014, Betancur et al. 18989 (COL), entrada a la quebrada la Maria, 30
Apr 2015, Betancur et al. 19237 (COL); San José del Guaviare, Serranı́a La
Lindosa (Bh-T), 2°28’N, 72°41’W [2.474, -72.696], 22 Aug 1995, Cárdenas
et al. 6514 (COL, MO); Serranı́a La Lindosa, sector Los Pozos, 2°43”N,
72°390 040W [2.716, -72.651], 16 May 2014, González et al. 1607 (COL); Mesa
La Lindosa, orilla de un arroyo, 12 Aug 1979, Idrobo & Zarucchi 9.431(COL,
P); 12 km al sur de San José del Guaviare, Mesa La Lindosa, 18 Aug 1979,
Idrobo & Zarucchi 9.436 (COL, P); por la trocha Nuevo Tolima, Ciudad e
Piedra (Zona Turistica), sabanas naturales y bosques, desarrollados en
afloramientos rocosos, arenas blancas, 2°290 12.1”N, 72°410 12.90W [2.486,
-72.686], 9 Jul 1997, López & Betancur 3114 (COL); Serranı́a La Lindosa,
Ciudad de Piedra, bosques dominadas por Andropogon spp., Syagrus aff.
inajai, Bysonima crassifolia, 2°280 280N, 72°410 480W [2.474, -72.696], 13 Nov
1995, López & Giraldo-C. 651 (MO); San José del Guaviare, antiqua represa,
27 Mar 1993, Sastre et al. 9180 (COL, P); Ciudad de Perdida, 28 Mar 1993,
Sastre et al. 9225 (COL), 9230 (COL).
 158 SYSTEMATIC BOTANY
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Fig. 7. A. Habit of Cuphea beneradicata, Sastre et al. 9180 (P), and flower, Cardenas 6514 (MO). B. Habit of C. blackii, Granville 11749 (MO), and flower,
Croat & Ferry 102998 (MO).
Cuphea beneradicata is a species with abundant, small,
spreading to congested leaves, and flowers with a dorsally
enlarged, prominent spur. The species resembles the more
widespread C. kubeorum, which differs by smaller leaves
mostly 2–6 mm long, an indistinct inflorescence, stems and
flowers exclusively strigose, and a smaller spur. Both species
occur to the south of San José del Guaviare, Colombia, in the
sandstone savannas of the Serranı́a La Lindosa.
2. CUPHEA BLACKII Lourteig, Sellowia 16: 137. 1964. TYPE:
BRAZIL. Amapá: Oiapoque [3.844, -51.832], beira de
estrada que vai para o campo de aviaç~
ao, 30 Sep 1949, G.
A. Black 49–8243 (holotype: P-P01901552-image!; isotype:
IAN).
Subshrubs 30–100 cm; stems erect, slender, from an erect to
decumbent base with abundant fibrous roots, dichotomously
branched; internodes half as long to longer than subtending
leaves, much shorter on small branches; stem indumentum
exclusively bifid-strigose, trichomes abundant, some ap-
pressed to the stem, others with semi-erect arms. Leaves
spreading, ascending, abruptly reduced in size in the in-
florescence; petioles 0.5–2.0 mm long; blades elliptic to oblong,
13–30 mm long, 2.0–8.5 mm wide, mostly ca. 20–25 mm long,
4–5 mm wide, membranaceous, base cuneate, apex acute,
margin plane or narrowly incurved, ciliate in the inflorescence;
surfaces dark green, paler abaxially, finely bifid-strigose
adaxially, midvein and 3–5 sets of secondary veins visible,
brown-red. Racemes distinct, terminal, bracteate, dichotomously
[Volume 44
branched or simple, 5–10 cm long; flowers paired at nodes,
partially obscured by bracts, pedicels to 1 mm long, broadened at
base, one slightly more distal than the other, abundantly bifid-
strigose; bracteoles much reduced, obscured by indumentum.
Floral tube, including a horizontal to subdescending, poorly
developed spur, to 0.3 mm long, 2.5–4.0 mm long, 1.5 mm wide,
ampullaceous in fruit, neck at 1/2 the length of floral tube,
remaining wide in fruit, mouth flaring, 1.7–2.0 mm wide, slightly
oblique; outer surface deep red-purple, abundantly bifid-
strigose, sometimes also with thin erect trichomes to 0.5 mm
long; inner surface lightly villous distal to stamen insertion,
glabrous proximally; sepals equal, 0.5 mm long, 0.6 mm
wide, shallowly deltate, apex apiculate; epicalyx segments
thickened cylindric lobes, shorter than sepals, frequently
with few long, thin trichomes. Petals 6, medium purple to
pink with darker midvein, subequal, the 2 dorsals ca. 2 mm
long, 0.8 mm wide, obovate, smaller than the ventrals, ca.
2.3 mm long, 0.9 mm wide, narrowly obovate. Stamens: 2
short dorsalmost inserted slightly deeper, other 9 inserted
in a horizontal line at ca. 1/2 the length of tube, deeply
included, none reaching sinuses of sepals; filaments very
short, ca. 0.5 mm long, densely white villous, filaments and
anthers pale; pollen oval-triangular, tricolporate, non-
syncolpate, pores slightly protruding, exine low parallel
rugulate, thickened at poles, grain diam 15 mm. Ovary
glabrous; style glabrous, included; ovules 3–6. Disc hori-
zontal to deflexed, sub-globose, 0.3–0.4 mm long, 0.3 mm
wide, dark-red. Seeds 1.5–1.7 mm long, 1.4–1.5 mm wide,
 2019] GRAHAM: CUPHEA SECTION AMAZONIANA
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Fig. 8. Distribution of Cuphea beneradicata, C. cardonae, C. cataractarum, C. chiribiquetea, and C. distichophylla.
sub-circular to obovate, apex rounded, base obtuse to retuse,
margin thinned, narrow, sub-marginal ridge present. Figures
4L, 5L, 7B.
Phenology—Collected in flower and fruit throughout the
year.
Distribution and Habitats—In French Guiana and Amapá,
Brazil; primary forest, among rocks on wet sandy soil, on clay
creek banks, borders of ponds in savanna, in white quartz
sand, and on thin soil over granite boulders, in partial shade on
south facing slopes; 6–400 m (Fig. 9).
Extinction Risk—The estimated risk if all IUCN guides
were to be followed is VU; EOO 516,129.54 km2 and AOO 5
64 km2. Six of the 16 collection localities analyzed occur in
protected areas.
Representative Specimens Examined—Brazil.—AMAPÁ: Rı́o Oiapoque
savanna immediately S of Oiapoque airfield [3.522, -52.796], 30 Jul 1960,
Irwin & Westra 47244 (K-image, NY); Oiapoque airfield, 25 Apr 1960, Egler
1435 (NY); Rı́o Oiapoque, Cachoeira Cacheri, 3°43’N, 51°57’W [3.716,
-51.95], immediately west of mouth of Cricu River, 14 Aug 1960, Pires 47534
(NY); on large granitic outcrop surrounded by forest near Mt. Carupina,
3°33’N, 51°37’W [3.55, -51.616],16 Oct 1960, Pires &Westra 48869 (NY, also
K, P, RB,W images); Oiapoque, campina de areia branca, aeroporto, 20 Mar
1982, Rabelo 1546 (MG). French Guiana. Savane-roche du Quatorze Juillet-
Bassin du Bas-Oyapock, 3°58’N, 51°52’W [3.966, -51.866], 15 Apr 1991,
Cremers 12126 (CAY, NY); Savane-roche, Anabelle, Bassin de l’ap-
prouagues, 4°0’N, 52°16W [4.0, -52.266], 10 May 1997, Cremers 15252 (NY);
Cayenne, along road from St. Georges to Saut Maripa, 18.5 km S of turnoff
from N2 Hwy, 0.8 km N of Saut Maripa, 3°48’N, 51°520 460W [3.813,
159
-51.879], 13 Mar 2011, Croat & Ferry 102998 (CAY, MO); Mts. de Kaw,
Trésor Reserve, Kaw, near main trail, third tree after big Ficus, part of
main savanne, [4.35, -52.16], 30 Jan 2003, Ek et al. 1394, 1509 (NY), 29 Sep
2006, Ek et al. 1759 (NY); Haute Crique Armontabo [3.7, -51.966], in-
selberg granitique, 23 Feb 1981, Granville de 4373 (NY); Haute Crique
Montagnes de la Trinité, Inselberg N-Quest, savane roche exposée au
Sud, border of forest,16 Jan 1984, Granville et al. 6023 (NY); Pic Matécho
rocky savanna, 3°N, 53°10 400W [3.0, -53.017],16 Sep 2000, Granville et al.
14242 (NY); Pic du Grand Croissant, Bassin de l’Armontabo, 3°31’N,
52°22’W [3.516, -52.366], savane roche, à proximité de la lisière; Sur le
sommet do Piton Baron [3.25, -53.066],14 Apr 1993, Granville 11749
(MO, NY); Nouragues Field Station, 4°05’N, 52°41’W [4.083, -52.733],
and vicinity, south facing slope of inselberg on Chemin Terrasse Trail 1,
about one-half distance to summit, 16 Feb 2002, Mori et al. 25376 (NY);
Oyapock, St. Georges de l’Oyapock, airfield [3.901, -51.802], 12 Apr
1976, Sastre 4758 (NY); Commune de Roura-Savanne de la Crique
Marguerite, 4°380 44.28”N, 52°200 330W [4.645, -52.342], savane, tige
rouge, fleurs violettes, 17 May 2015, P. Silland & J. Sutra 932 (CAY, MO);
Inselberg Roche-Dachine, 3°28’N, 53°13’W [3.466, -53.216], Apr, Villiers &
Sarthou 6023 (NY).
Cuphea blackii is distinguished by elliptic to oblong leaves
separated by long internodes, and by stems dichotomously
branched distally. The terminal racemes are distinctive, having
very short internodes with flowers paired at the nodes, and
highly reduced bracteoles obscured by the indumentum.
Floral tubes are ampullaceous in fruit, abundantly bifid-
strigose, sometimes also sparsely hirsute, and the anthers
are deep in the floral tube on short filaments. Just 3 or 4 ovules
are produced. Cuphea blackii shares the characteristic
 160
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Fig. 9. Distribution of Cuphea blackii, C. bolivariensis, C. dactylophora, and C. insolita.
dichotomous branching pattern with C. rigidula, also occuring
in French Guiana, a species with alternate flowers, vegetative
parts abundantly setose, and 9 or 10 ovules.
3. CUPHEA BOLIVARIENSIS Lourteig, Phytologia 60: 30. 1986.
TYPE: VENEZUELA. Bolı́var: Urimán [5.350, -62.666], Rı́o
Apacará, 14 Aug 1954, A. L. Bernardi 1415 (holotype: NY-
00386500!; isotype: MER).
Subshrubs to 30 cm; stems several from a woody primary
root with many secondary fibrous roots, semi-decumbent,
distally ascending, unbranched or sparsely branched; in-
ternodes less than half as long as subtending leaves; stem
indumentum exclusively bifid-strigose, trichomes thin, ap-
pressed, very short, ca. 0.2 mm long. Leaves spreading and
ascending, decussate proximally on the stem, irregularly
arranged distally, not strictly distichous, similar in length
along the vegetative stem, very gradually reduced in size in the
inflorescence, crowded and imbricate; petioles 0.1–1.5 mm
long, dark when dry; blades linear, appearing uni-nerved,
6.5–12.0 mm long, 1–2 mm wide, coriaceous, not rigid, base
cuneate, apex obtuse, margin slightly incurved; surfaces
adaxially green, glabrous, abaxially paler, finely bifid-strigose,
midvein brown, prominent abaxially, secondary veins not
clearly visible. Racemes indistinct, leafy, simple; flowers al-
ternate, interpetiolar, crowded distally on the stem, partially
obscured by leaves; pedicels 2.5 mm long; bracteoles linear, to
0.3 mm long. Floral tube, including a poorly developed spur to
SYSTEMATIC BOTANY [Volume 44
0.2 mm long or non-spurred, 4 mm long, 1.9 mm wide, neck
contracted at 1/2 length of floral tube, mouth flaring,
2.5–3.5 mm wide; outer surface color unknown, indumentum
finely bifid-strigose as on stems, appearing nearly glabrous;
inner surface densely villous distal to stamen insertion, the
trichomes white or light purple, lightly pilose proximally;
sepals equal, 1.2 mm long, 0.8–1.0 mm wide, deltate, with a
central dark stripe, apex acute; epicalyx segments thickened
lobes, 0.3 mm long, much shorter than the sepals, without
setae. Petals 6, pink or light purple, oval to oblong, subequal, 2
dorsals 4.0 mm long, 2.4 mm wide, 4 ventrals 3.8–4.0 mm long,
2.0–2.2 mm wide. Stamens: 2 short dorsalmost deepest, other 9
inserted at ca. 1/2 length of the tube, all included, position with
respect to floral tube margin unknown; filaments and anther color
unknown; pollen oval-triangular, tricolporate, non-syncolpate,
pores slightly protruding, exine rugulate, rugulae low, parallel,
poles thickened and coarsely rugulate, grain diam 15–17 mm. Ovary
lightly pilose; style lightly pilose at base, included to exserted and
well surpassing sepals; ovules 3. Disc horizontal to deflexed, sub-
globose, ca. 0.3 mm long and wide. Seeds unknown. Figure 10A.
Phenology—Collected in flower and immature fruit in
August.
Distribution and Habitats—Venezuela in Bolı́var state and
in Roraima, Brazil; wet savannas and margins of streams; 500
and 850 m (Fig. 9).
Extinction Risk—The estimated risk if all IUCN guides
were to be followed is CR; EOO not available and
 2019] GRAHAM: CUPHEA SECTION AMAZONIANA
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Fig. 10. A. Habit and flower of Cuphea bolivariensis, holotype, Bernardi 1415 (NY). B. Habit and flower of C. cardonae,isotype, Cardona 2972 (MO).
AOO 5 8 km 2 . Two collection localities were analyzed, both in
protected areas.
Additional Specimen Examined—Brazil.—RORAIMA: Potaro-Siparuni
region, Pakaraima Mountains, Upper Ireng River watershed, savanna at
eastern base of Malakwalai-Tipu, 4°48’N, 60°16’W [4.8, -60.266], upland sa-
vanna, sand and claystone, 18 Jul 1994, T. W. Henkel 5694 et al. (NY, P). This
collection, labeled from Guyana, was made in Brazil (Henkel et al. 2016: 52).
Cuphea bolivariensis produces several sparsely-branched
primary stems closely arising from an enlarged woody root-
stock. Leaves are relatively uniform in length, closely arranged,
mostly 10–15 mm long (proximal leaves not seen), and separated
by very short internodes. Floral tubes, according to the proto-
logue, are spur-less, 4 mm long with pale pink or purple petals,
and three ovules. Leaves of C. bolivariensis resemble those of C.
distichophylla, but are not rigid, are less distinctly distichous, and
have an obtuse, rather than acute, apex. Stems and flowers of C.
bolivariensis are exclusively strigose, with the bifid trichomes
notably thin and closely appressed to the surface; in C. dis-
tichophylla, they are mixed bifid-strigose and hirsute. Cuphea
trisperma in eastern Venezuela, could be confused with C. boli-
variensis and C. distichophylla. Like them, it has indistinct racemes;
small non-spurred floral tubes, and three ovules. The leaves,
however, are more narrowly linear, and the disc is cylindric and
reflexed, rather than sub-globose and horizontal to deflexed. The
pollen of C. bolivariensis is unusual, having thickened, coarsely
rugulate poles that obscure the termination of the colpi.
161
4. CUPHEA CARDONAE Lourteig, Phytologia 60: 42.1986.
Replaced synonyms: C. rhodocalyx Lourteig forma setosa
Lourteig, Sellowia 16:136.1964; C. rhodocalyx Lourteig var.
setosa Lourteig, Phytologia 60: 42. 1986, non C. setosa
Koehne. TYPE: VENEZUELA. Bolı́var: suelo rocoso del Rı́o
Pauo, afluente del Rı́o Caura [ca. 4.8, -64.15], 390 m, Jan
1963, F. Cardona 2972 (holotype: P-05134203, image is an
unopened envelope; isotypes: MO-2983144!,VEN).
Subshrubs to 60 cm; stems erect, much-branched, young
stems slightly quadrangular from decurrent leaf bases; roots
undescribed; internodes one-half or one-third as long as
subtending leaves; stem indumentum of two types: bifid-
strigose, trichomes sparse to abundant, thin, silky, ap-
pressed; and glandular setose/hirsute, trichomes brown,
resin-producing, enlarged at base, less coarse distally on
stems. Leaves decussate, larger lower leaves loosely spread-
ing, distal leaves and bracts of inflorescence sub-ascending to
ascending, gradually reduced in size in the inflorescence;
petioles 0–1 mm long; blades narrowly elliptic, 12–17 mm long,
2–4 mm wide, thickly membranaceous, base cuneate, apex
subacute, margin plane; surfaces green and glabrous or
sparsely bifid-strigose, paler abaxially, midvein red-brown, 2
or 3 sets of thin secondary veins visible. Racemes 6 distinct,
terminal, transitioning from leafy to bracteate, simple; flowers
 162 SYSTEMATIC BOTANY
solitary, alternate, crowded distally, interpetiolar, pedicels
0.5–1.0 mm long; bracteoles ca. 1 mm long, 2 mm wide. Floral
tube, including a poorly developed spur to 0.5 mm long or
non-spurred, 4–5 mm long, 1.5–2.0 mm wide, ampullaceous in
fruit, neck slightly contracted at 2/3 length of floral tube,
mouth flaring, 2.0–2.5 mm wide; outer surface rose-purple,
sparsely finely bifid-strigose and sparsely hirsute; inner sur-
face villous distal to stamen insertion, lightly pilose proxi-
mally; sepals subequal, 0.5–0.7 mm long, 0.4–1.0 mm wide,
dorsal sepal largest, widest, deltate to broadly deltate, apex
acute; epicalyx segments thickened, 0.3–0.5 mm long, ovate,
about as long as sepals, glabrous, dark-colored. Petals 6, white,
obovate, 1.5–2.0 mm long, 0.8–1.0 mm wide including narrow
dark purple claw 0.5 mm long. Stamens: 2 short dorsalmost
inserted at ca. same level as other 9; filaments villous, filament
and anther color unknown; pollen triangular, syncolpate,
colpi convergent at the poles in a triangle enclosing a remnant
of exine, pores not protruding, exine coarsely, irregularly
rugulate, grain diam ca. 19 mm. Ovary and style lightly pilose,
style ultimately extended to apices of sepals; ovules 5. Disc
reflexed over ovary, cylindric to slightly flattened and nar-
rowly oblong, 0.9 mm long, obtuse, dark-colored. Seeds
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0.9–1.1 mm long, 0.8–0.9 mm wide, sub-deltoid. Figure 10B.
Phenology—Collected in flower and fruit in January and
February.
Distribution and Habitats—Venezuela, in Amazonas and
Bolı́var states; on rocky beaches of rivers; 200–390 m (Fig. 8).
Extinction Risk—The estimated risk if all IUCN guides
were to be followed is EN; EOO 5 735.36 km2 and AOO 5
12 km2. Three collection localities were analyzed, all occurring
in protected areas.
Specimens Examined—Venezuela.—AMAZONAS: Rı́o Metacuni, orillas
del rı́o Metacuni entre el tepuy-altiplanicie Sedukerawa [3.25, -64.933] y el
Salto Wânitı̂’ma, 30 Jan 1990, Stergios & Velazco 14480 (MO, NY); Rı́o
Metacuni, selvas ribere~nas y de banco, 3°2’N, 65°12’W [3.033, -65.12], 2 Feb
1990, Stergios & Velazco 14573 (MO). —BOLÍVAR: al lado Rı́o Nichare
(afluente de Rı́o Caura), arriba de la desembocadura con el Rı́o Cicuta
(Icuta), acercandose a las cabeceras del Rı́o Nichare en la dirección de la
Sierra Maigualida y Sierra Cervatana, 6°15’N, 65°50 W [6.25, -65.083], 25
Apr 1966, Steyermark & Gibson 95670 (NY, P); Rı́o Diamante Negro, trib-
utary of Rı́o Gua~na (4.2, -63.766), 27 Apr 1973, Steyermark et al. 107107 (P).
In Cuphea cardonae, young stems are slightly quadrangular
and abundantly glandular hirsute to setose, and distal leaves
transition to a raceme that is intermediate between leafy and
truly bracteate. The species was first described as a setose
variety of C. rhodocalyx, then later elevated to species rank
based primarily on the presence of setae, which do not occur in
C. rhodocalyx. Cuphea rhodocalyx, a narrow endemic from the
Marahuaka-Duida region southwest of the type locality of C.
cardonae, is the only other species of sect. Amazoniana with
distal portions of stems quadrangular. Cuphea rhodocalyx differs
from C. cardonae by stems glabrous or rarely sparsely strigose,
stems more distinctly quadrangular, and discs short and erect,
as opposed to reflexed in C. cardonae. Petal color is white with
purple claws in C. cardonae. In C. rhodocalyx it is pale purple or
white or bi-colored with the dorsal petals deeper purple than
the ventrals. I recognize C. cardonae as questionably distinct
from C. rhodocalyx and in need of further collections and study.
The two species share several characters with two other Ven-
ezuelan species, C. curiosa and C. dactylophora; all have spur-less
flowers with an enlarged dorsal sepal, an ampullaceous form in
fruit, and syncolpate, psilate to rugulate pollen.
5. CUPHEA CATARACTARUM Spruce ex Koehne in Martius, Fl.
Brasil. 13(2): 226.1877. TYPE: VENEZUELA. Amazonas: ad fl.
[Volume 44
Orinoco, pr. Esmeralda [3.164, -65.55] et ad fl. Rı́o
Cunucunuma, Dec 1853, R. Spruce 3261 (holotype: B†;
lectotype designated [as holotype] by Lourteig, 1986: P-
01901529-image!; isotypes: GOET!, NY! OXF!; isotype
images: BR! F 3 4! GH! LD! MPU! P 3 2! S! TCD!).
Subshrubs 30–80 cm; stems one to several, long, lax, as-
cending from a woody primary root with fibrous secondary
roots, sparsely branched, secondary branches long; internodes
less than half as long as subtending leaves, much shorter
distally; stem indumentum of two types: predominantly bifid-
strigose, trichomes very thin, closely appressed to the stem;
sometimes also hirsute/setose, trichomes erect to slightly
appressed, red or without color, longer, 0.5–1 mm long. Leaves
closely imbricate distally on the stem, gradually reduced in
size in the inflorescence; petioles 0–0.5 mm; blades elliptic,
5–22 mm long, 2–7 mm wide, appearing uni-nerved, thickly
membranaceous, base and apex rounded to obtuse, rarely
acute, margin plane; surfaces green, finely bifid-strigose, 1–3
pairs of secondary veins usually visible abaxially on the larger
proximal leaves. Racemes indistinct, terminal, leafy, simple;
flowers alternate, inter-petiolar, crowded distally on the stem,
partially obscured by leaves and bracts; pedicels 1–3 mm long;
bracteoles narrowly ovate, 0.3–0.4 mm long, 0.1 mm wide.
Floral tube, including a poorly developed spur to 0.5 mm long
or non-spurred, 5–7 mm long, 1.5 mm wide, neck contracted at
2/3 length of floral tube, mouth widely flaring, 2.5–3.0 mm
wide; outer surface color unknown, finely bifid-strigose as on
stems, appearing nearly glabrous, occasionally with red,
longer, thin trichomes; inner surface abundantly villous distal
to stamen insertion, glabrous to lightly pilose proximally;
sepals unequal, dorsalmost sepal largest, ca. 1.2 mm long,
1.2 mm wide, others ca. 0.7 mm long, 0.7 mm wide, all deltate,
apex acute; epicalyx segments cylindric lobes, 0.3 mm long,
much shorter than sepals, without terminal setae. Petals 6,
white, unequal, 2 dorsals smaller than 4 ventrals, spatulate,
1.4 mm long, 0.7 mm wide, ventrals narrowly spatulate,
1.9–2.2 mm long, 1.0–1.4 mm wide. Stamens: 2 short dor-
salmost deepest, other 9 nearly in a horizontal line at 2/3
length of floral tube, antesepalous stamens reaching apices of
sepals, antepetalous ones reaching sinuses of sepals; filaments
white villous, filament and anther color unknown; pollen
oval-triangular, tricolporate, syncolpate, colpi convergent at
poles in a triangle enclosing a remnant of exine, pores non-
protruding, exine psilate, grain diam 18–19 mm. Ovary gla-
brous; style glabrous, ultimately extended to apices of the
sepals or exserted 1 mm; ovules 4–7, terminal on a flattened,
narrow, placenta. Disc erect, sub-cylindric, obtuse, 0.7–0.9 mm
long, 0.5 mm wide, thick, dark-colored. Seeds 1.0–1.2 mm
long, 0.8–0.9 mm wide, scarcely compressed, obovate, apex
rounded, margin rounded. Figures 5E, 11A.
Phenology—Collected in flower and fruit November to
August.
Distribution and Habitats—Venezuela, a narrow endemic
in Amazonas state; in tall forest bordering rivers, on rocky
riverbanks, in crevices; 170–350 m (Fig. 8).
Extinction Risk—The estimated risk if all IUCN guides
were to be followed is EN; EOO 5 654.33 km2 and AOO 5
20 km2. Five collection localities were analyzed, all in protected
areas.
Specimens Examined—Venezuela.—AMAZONAS: Rı́o Cunucunuma, near
Cliff House, in marsh, 4 Jun 1950, Curran 176 (NY); 4 Aug 1957, Curran 241
(NY); Rı́o Cunucunuma a lo largo del Orinoco, Jan-Feb 1969, Fari~
nas et al. 462
(NY); Atabapo, Rı́o Cunucunuma, raudal Báquiro (Dukari-Shodo en lengua
 2019] GRAHAM: CUPHEA SECTION AMAZONIANA
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Fig. 11. A. Habit of Cuphea cataractarum, Maguire et al. 30348 (MO), and flower, Steyermark 125678 (MO). B. Habit of C. chiribiquetea, Barbosa 7963 (P), and
flower, Franco 3282 (NY).
Ye’kuana), 3°47’N, 65°54’W [3.783, -65.90], Mar 1990, A. Fernandez 7400 (MO,
NY); along Rı́o Cunucunuma near Culebra, 3°44’N, 65°44’W [3.733, -65.733],
15–16 Feb 1985, Liesner 17524 (MO, P 3 2); Rı́o Cunucunuma, sandbars along
river above Playa Alta, 14–15 Nov 1950, Maguire et al. 29497 (COL, NY); Rı́o
Cunucunuma, common along river in rock crevices at Culebra Rapids (north
base of Cerro Duida [ca. 3.513, -65.626]), 23 Dec 1950, Maguire et al. 30348 (MO,
NY, P, RB); entre las comunidades de Culebra y Huachamacari, entre el Cerro
Duida y Huachamacari, 3°40’N, 65°45’W [3.666, -65.75], 28–30 Jan and 6–8 Feb
1982, Steyermark et al. 125678 (MO, NY, P).
Cuphea cataractarum is recognized by the unique combina-
tion of: stems, long, sparsely-branched, proximally decumbent
or prostrate and distally ascending, with bifid and sometimes
also simple trichomes; leaves imbricate, elliptic; racemes in-
distinct; floral tubes 5–7 mm long, the spur poorly developed
or none; and ovules 6 or 7.
Determination of the type of C. cataractarum was based on
inspection of three successive publications. In the protologue
Koehne (1877) initially cited, “hb. Spruce” and “Venezuela.” In
1881, he expanded the citation to “Venezuela dec.: ad fl.
Orinoco pr. Esmeralda! et ad fl. Cunucunuma!” Full citation of
the type collection (Koehne 1903: 128–129) appeared as
“Venezuela: am Orinoco bei Esmeralda und am Cunucunuma
(Spruce n. 3261).” Although the material studied by Emil
Koehne at Berlin was subsequently destroyed, Spruce 3261 is
well represented in other herbaria by at least 14 isotypes. Three
specimens of Spruce 3261 with original labels were available to
163
Lourteig at P. She designated the one labeled “ad fl en
Cunucunuma,” and initialed by Koehne, as the holotype
(Lourteig 1986: 32). The specimen, now assigned identification
number P-01901529, is correctly considered the lectotype. The
other two Spruce 3261 specimens at P are from “prope
Esmeralda, ad flumen Orinoco, Dec 1853”, and were not seen
by Koehne.
6. CUPHEA CHIRIBIQUETEA Lourteig, Bradea 8: 27. 1996. TYPE:
COLOMBIA. Caquetá: Sierra de Chiribiquete, bajada hacia
refugio Bernardo, Quebrada Espa~ nolas [0.933, -72.701], al
pie de la cascada en rocas rezumantes, 635 m, 21 Nov
1992, M. Velayos, J. M. Cardiel, J. Pedrol, and M. T. Telleria
6329 (holotype: P-P01901526, image!; isotypes: COL!,
MA-image!).
Subshrubs to 40 cm; stems erect to decumbent from a woody
base with fibrous roots, branched; stem indumentum com-
posed of sessile ovoid or oblong glands, multi-celled at the
base, colorless with amorphous contents, ca. 100 mm long,
50 mm wide, often with a short emergent arm, older stems
glabrous; internodes to 5 mm long, much shorter than the
subtending leaves. Leaves spreading to ascending, crowded
distally on the stem, very gradually reduced in size in the
inflorescence; petioles 0–0.5 mm long; blades narrowly linear,
7–30 mm long, 0.5–1.5 mm wide, appearing uni-nerved,
 164 SYSTEMATIC BOTANY
coriaceous, base cuneate, apex subacute to obtuse, margin
plane to slightly incurved; surfaces glabrous, adaxially green,
abaxially paler, midvein brown, prominent abaxially, second-
ary veins not visible. Racemes 6 indistinct, terminal, leafy,
simple and compound; flowers solitary, alternate to sub-
opposite, inter-petiolar, crowded distally on the stem, par-
tially obscured by bracts; pedicels 1.5–2.0 mm long, slender,
persistent; bracteoles lanceolate, 0.5 mm long. Floral tube,
including a horizontal to descending, well-developed spur to
0.7 mm long, 3.5–4.5 mm long, 1.0–1.5 mm wide, neck slightly
contracted at 2/3 length of floral tube, mouth flaring, ca. 2.0 mm
wide, oblique, extended ventrally; outer surface green, glabrous
or nearly so, sometimes with sparse glands as on stems; inner
surface densely white villous; sepals subequal, the dorsalmost
reflexed, 0.8 mm long, 0.8 mm wide, ventralmost wider than the
others, 0.8 mm long, 1.3 mm wide, all deltate, apiculate; epi-
calyx segments cylindric lobes, 0.5 mm long, dark, shorter than
sepals, without terminal setae. Petals 6, light purple, subequal, 2
dorsals smaller than the 4 ventrals, obovate, 1.7 mm long,
0.7 mm wide, 4 ventrals narrowly obovate, 2 mm long, 1.3 mm
wide. Stamens: 2 short dorsalmost slightly deeper, other 9
inserted in a horizontal line at 2/3 length of floral tube, ante-
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sepalous stamens reaching apices of sepals, antepetalous ones
deeper; filaments densely white villous, filament and anther
color unknown; pollen oval-triangular, tricolporate, non-
syncolpate, pores not or slightly protruding, exine low,
coarsely rugulate, rugulae parallel, grain diam 15 mm. Ovary
and style pilose, style reaching apex of large ventral sepal;
ovules 4 or 5. Disc horizontal, globose to sub-deltoid, obtuse at
apex, 0.3 mm long, 0.5 mm wide at base, thick, dark red. Seeds
1.0–1.1 mm long, 1.0 mm wide, obovate, base rounded to ob-
tuse, apex obtuse, margin thick, rounded. Figures 2C, F, 3D,
11B.
Phenology—Collected in flower and fruit in November and
December.
Distribution and Habitats—Colombia, in Departments
Guaviare and Caquetá, on Sierra de Chiribiquete; in mon-
tane forest along rivers and near falls, in rock crevices;
300–650 m.
Extinction Risk—The estimated risk if all IUCN guides
were to be followed is EN; EOO 5 480.38 km2 and AOO 5
16 km2. Three of the four collection localities analyzed occur in
protected areas (Fig. 8).
Specimens Examined—Colombia.—CAQUETÁ: Campamento Sur, en el
rı́o entre la cascada grande y la cascada de arriba, 0°55’N, 72°45’W, [0. 916,
-72.75], 8 Mar 1990, Franco et al. 3282 (COL, NY); Sierra de Chiribiquete, en
la via del Campamento a la Cueva de Pinturas, 1°05’N, 72°40’W [1.08,
-72.666], 17 Aug 1922, Franco et al. 3760 (COL).—GUAVIARE: extreme Norte
de parque, cerca de Dos Rı́os, 1°48’N, 72°44’W[1.8, -72.733], 19 Nov 1992,
Barbosa 7963 (COL, NY, P).
Cuphea chiribiquetea is a narrow endemic from the epony-
mous Sierra de Chiribiquete, Colombia. Uniquely, young
stems and tiny flowers bear microscopic sessile glands,
replacing the normal bifid trichome type of sect. Amazoniana.
Normal bifid trichomes occasionally are present along the
adaxial leaf midvein and scattered elsewhere on the abun-
dant, very narrow, linear leaves. The flowers at 3.5–4.5 mm
long are second smallest in the section, next to those of C.
blackii at 2.5–4.0 mm long. Three other species in sect.
Amazoniana occur in the Chiribiquete region: the similar,
poorly known C. stygialis; and C. beneradicata and C. kubeo-
rum, which differ by leaves typically much shorter (mostly
2–11 mm) than the mostly 15–20 mm long leaves of C.
chiribiquetea.
[Volume 44
7. CUPHEA CURIOSA Lourteig, Phytologia 60: 38, Fig. 3. 1986.
TYPE: VENEZUELA. Bolı́var: selva de galerı́a y sabana a lo
largo del Rı́o Kanarakuni [Canaracuni], N y NW de la
misión de Campamento de Sanidad del Rı́o Kanarakuni,
[ca 4.75, -64.433], 400 m, 17–29 Mar 1967, J. Steyermark 97835
(holotype: P-01901696-image!; isotype: VEN-image!).
Cuphea curiosa var. oresbia Lourteig, Bradea 8: 28–29. 1996.
TYPE: VENEZUELA. Amazonas: Atures, Sierra de Maigualida,
NW sector, small valley along an upper tributary of Ca~ no
Iguano [ca. 5.5, -65.537], 20 Feb-3 Mar 1991, P. Berry, O.
Huber, & J. Rosales 4904 (holotype: P-01901695-image!; iso-
type: MO!).
Subshrubs to 50 cm; stems erect, slender, ascending from a
woody, decumbent, rhizomatous base with abundant sec-
ondary fibrous roots, much branched distally; internodes less
than half as long as subtending leaves; stem indumentum of
two types: exclusively bifid-strigose, trichomes relatively
abundant, thin, silky, to 0.5 mm long on young growth; or
bifid-strigose mixed with simple thin to coarse, reddish-brown
trichomes, erect or lax, 0.5–1 mm long. Leaves spreading,
distal leaves ascending, imbricate, abruptly reduced in size in
the inflorescence; petioles to 0.5 mm long; blades narrowly
elliptic to lanceolate-linear, 5–40 mm long, 1–8 mm wide,
thinly membranaceous, base cuneate, apex acute or subacute,
margin plane; surfaces green, adaxially glabrous or with some
bifid trichomes, abaxially moderately to abundantly bifid-
strigose, midvein and 1–6 sets of thin, red-brown secondary
veins, visible on both surfaces. Racemes distinct, terminal,
bracteate, simple or compound; bracts elliptic to oval, mem-
branaceous, 2–4 cm long; flowers alternate, crowded distally
on the stem, not obscured by bracts; pedicels 0.5–1.5 mm long;
bracteoles narrowly ovate, 0.3–0.5 mm long. Floral tube,
including a poorly developed spur to 0.3 mm long, 4.0–5.5(–7.0) mm
long, 1.5 mm wide, neck at 2/3 length of floral tube, mouth widely
flaring, ca. 2 mm wide, neck and mouth open in fruit; outer surface
violet or green, finely bifid-strigose, almost glabrous, or mixed with
thinner, longer trichomes; inner surface villous distal to stamen
insertion, pilose proximally; sepals subequal, dorsalmost sepal
largest, widest, 0.4–0.7 mm long, 0.6–0.8 mm wide, the others ca.
0.6 mm long, 0.6 mm wide, deltate, apex acute; epicalyx segments
thickened lobes, half as long to as long as sepals, green or dark-
colored, erect to recurved, glabrous or with some bifid trichomes.
Petals 6, white, pink, pale purple, white with purple midvein or
purple base, or dorsals darker than pink ventrals (fide Lourteig
1986), subequal, 2 dorsals larger and wider than the ventrals,
1.7–1.8 mm long, 0.5–0.8 mm wide, 4 ventrals 1.7 mm long,
0.4–0.6 mm wide, all spatulate to obovate. Stamens: 2 short dor-
salmost deepest, other 9 inserted in a horizontal line at 2/3 length of
floral tube, antesepalous stamens extending nearly to sepal apices,
antepetalous ones shorter, not reaching sinuses of sepals; filaments
sparsely villous, filament and anther color unknown; pollen tri-
angular, tricolporate, syncolpate, colpi convergent at poles in a
triangle enclosing a remnant of exine, pores non-protruding, exine
psilate, grain diam 20–23 mm. Ovary and style sparsely pilose, style
extending to apices of the sepals or exserted ca. 1 mm; ovules 5–9, at
distal end of a flattened placenta. Disc erect to strongly reflexed over
ovary, cylindric with obtuse apex, 0.7–1.0 mm long, 0.4 mm wide,
thick, dark red. Seeds 0.9–1.1 mm long, 0.8–0.9 mm wide, ob-
ovate, apex rounded, margin rounded. Figures 3C, 4E, 5F, 12A.
Phenology—Collected in flower and fruit from October to
March; in sparse flower without distinct racemes in April and
May.
 2019] GRAHAM: CUPHEA SECTION AMAZONIANA
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Fig. 12. A. Habit of Cuphea curiosa, Boom & Marin 10546 (P), and flower, Wurdack & Monachino 41073 (MO). B. Habit of Cuphea dactylophora, Steyermark
et al. 104483 (MO), and flower, Berry et al. 6590 (MO).
Distribution and Habitats—Venezuela, in Bolı́var and
Amazonas states, and in Amazonas, Brazil; evergreen forest,
margins of rivers in gallery forest among rocks and in
neighboring savanna; at 110–830 m and 1400–2100 m (Fig. 13).
Extinction Risk—The estimated risk if all IUCN guides
were to be followed is LC; EOO 5 148,245.48 km2 and AOO 5
52 km2. Nine of the 14 collection localities analyzed occur in
protected areas.
Specimens and Images Examined—Brazil.—AMAZONAS: slopes of Pico
da Neblina [0.804, -66.006], slopes above Igarapé, Anta, rocks in creek, in
high forest, 15 Dec 1965, Silva & Braz~
ao 60641(MO, NY). Venezuela. —AMAZONAS:
Atabapo, Salto Yureba, Ca~ no Yureba, Bajo Ventuari, 4°30 N, 66°1’W [4.05, -66.016],
24 Oct–4 Nov 1981, aff., Delascio Ch. & Guánchez 10717 (MO, P), 10811 (MO); Alto
Yureba, Cerro Yureba, lower Ventuari, 4°03’N, 66°01’W [4.05, -66.01], 14 Mar
1985, aff., Liesner 18650 (MO, P); Rı́o Siapa [ca. 5.05, -67.79], between
raudal Gallineta and Salto Gallineta, 23 Jul 1959, Wurdack & Adderley
43590 (MO, NY, P). BOLÍVAR: Foráneo Aripao, Alto Rı́o Caura, 15 km aguas
arriba de la comunidad Y’dkuana “Aragua~ na”, ca. 4°74”(erroneous data)
N, 64°12’W [ca. 5.25, -64.2], 4–5 May 1988, Aymard & Delgado 6888 (MO,
NY, P); Cede~ no, along Rı́o Mawela, tributary of the Erebato, 5°8’N,
64°56’W [5.13, -64.933], 16 Mar 1992, Boom & Marin 10546 (MO, NY, P);
Cede~ no, cuneca alta rı́o Nichare, 5°53’N, 65°02’W [5.883, -65.033], May
1988, aff., Elcoro 342 (MO); Cede~ no, cuenca alta del Rı́o Mato, extreme
Norte de la Serrania de Maigualida, 6°35’N, 65°13’W [6.583, -65.216], Apr
1989, aff.,Fernández, A. 5231 (MO); Raul Leoni en orilla de quebrada, parte
N del Macizo Guanacoco, 4°57”N, 63°53’W [4.95, -63.883], Nov 1988, aff.,
Fernández, A. & Aymard 4845, 4870 (MO); Cede~ no, Sierra de Maigualida,
sector nor-oriental, altiplanicie tepuyana ubicada en las cabeceras del Rı́o
Chajura, afluente occidental del Rı́o, aprox. 100 km en linea recta al SW
del Campamento Entrerı́os, 5°33’N, 65°13’W [5.55, -65.216], 28 Feb 1988,
165
aff., Huber 12735 (P, paratype of C. curiosa var. oresbia); Rı́o Erebato,
Raudal Dacu, 5°08’N, 69°48’W [5.133, -64.8], Feb 1989, Sanoja 2619 (MO);
Riberas del Rı́o Canaracuni, (aguas negras), 4°27’N, 64°7’W [4.45,
-64.116], 13–26 Apr 1988, Stergios 11877 (MO, NY); Canaracuni, Rı́o
Canaracuni, entre Canaracuni y La Boca, 4°30’N, 63°42’W [4.5, -63.7],
13–16 Apr 1988, Stergios 11931 (MO, NY); along Rı́o Parguaza [6.444,
-67.162] between El Carmen and Raudal Maraca, 50–100 km from river
mouth, 1 Jan 1956, Wurdack & Monachino 41073 (COL, MO, NY, P, RB,
VEN-image).
Among the Venezuelan species in the section, delimitation
of C. curiosa remains the most problematic. Field studies
within this complex are still required for a fully satisfactory
taxonomy. Collections annotated as C. curiosa by Lourteig,
author of the species, exhibit considerable variation in habit,
indumentum, and leaf size and shape across an extended
geographical range in Venezuela. The type has: bifid-
strigose stems; elliptic leaves, broad at the middle, dis-
tinct from the bracts of the racemes; short pedicels
0.5–1.5 mm long, floral tubes strigose to nearly glabrous,
4.0–5.5 mm long; petals white or pale purple; and a dark red
disc reflexed over the ovary. Cuphea curiosa var. oresbia,
growing on Sierra de Maigualida at 2000–2100 m, was
distinguished from var. curiosa, at 400 m, by stems and
leaves bifid-strigose with some setae, and floral tubes in-
frequently with some short setae (Lourteig 2001). Because
var. oresbia is not accompanied by at least one distinctive,
stable diagnostic feature, I consider it synonymous with the
nominal variety.
 166 SYSTEMATIC BOTANY
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Fig. 13. Distribution of Cuphea curiosa.
Tentatively included in this species is an enigmatic group of
four Venezuelan collections that diverge from the type of C.
curiosa, varying in amount of branching, in the trichome mix on
the stems, and in size and spacing of leaves. The collections are:
from Amazonas, Venezuela: Delascio & Guanchez 10717 and
Liesner 18650; and from Bolı́var, Venezuela: Elcoro 342, and
Fernández 5231. I have annotated them as having affinity (aff.)
to C. curiosa.
Cuphea curiosa is similar to C. pleiantha, but the plant habit is
more delicate, plants are smaller with thinner leaves, racemes
are slender with fewer branches, and stems are predominantly
bifid-strigose.
8. CUPHEA DACTYLOPHORA Koehne in Mart., Fl. bras. 13(2): 226.
1877. TYPE: GUYANA: Rich. Schomburgk 1556, (holotype: B†;
lectotype (designated as holotype by Lourteig 1986: 36):,
Schomburgk 1556a, 5 K-000533053, image!).
Subshrubs to 45 cm; stems erect, ascending, one or more
arising from a small woody rhizome bearing abundant fibrous
roots, branched along the length of the stem; internodes one-
third or less as long as subtending leaves, leaves and in-
ternodes very gradually diminishing in length toward the
stem apex; stem indumentum bifid-strigose, trichomes short,
closely appressed. Leaves loosely spreading; petioles 0–2 mm
long; blades narrowly elliptic-lanceolate to lanceolate-linear,
[Volume 44
10–45 mm long, 2–8 mm wide, thinly membranaceous, base
cuneate, apex subacute, margin plane; surfaces adaxially deep
green, abaxially pale green drying yellow- to red-brown, both
surfaces strigose, the bifid trichomes very thin, short,
0.2–0.3 mm long, arms equal, midvein and 3 pairs of secondary
veins visible on proximal leaves, dark red-brown. Racemes
indistinct, terminal, leafy, simple; flowers solitary, sparse,
alternate, obscured among leaves; pedicels slender, increasing
in length with age of flower, 1–7 mm long; bracteoles orbicular,
0.5 mm long. Floral tube, including a well-developed, hori-
zontal to descending spur 0.4–1 mm long, 6–9 mm long,
1.3–1.5 mm wide, neck strongly contracted at 2/3 length of
floral tube, mouth widely flaring, 2.5–3.0 mm wide; outer
surface bifid-strigose, dark purple, at least distally, inner
surface light purple to white, villous distal to stamen insertion,
lightly pilose proximally; sepals subequal, dorsalmost sepal
scarcely larger, ca. 1.2 mm long, 1.2 mm wide, the others
0.7–1.0 mm long, 0.7–1.3 mm wide, or 3 dorsal sepals slightly
larger than 3 ventrals, all dark purple, deltate, apex acute;
epicalyx segments thickened lobes, 0.3 mm long, shorter than
sepals, dark purple. Petals 6, pale purple with dark purple
base, or white with dark purple midvein, obovate to ovate,
unequal, 2 dorsals 2.7–3.7 mm long, 0.8–2.0 mm wide, 4
ventrals 3.5–5.0 mm long, 2.0–3.0 mm wide. Stamens: 2
dorsalmost very short, inserted at ca. same level as the other
 2019] GRAHAM: CUPHEA SECTION AMAZONIANA
9 in a horizontal line, all included, antesepalous stamens
extending nearly to apices of sepals, antepetalous ones
proximal to or at sinuses of sepals; filaments white villous,
filament and anthers pale; pollen triangular in polar view,
tricolporate, syncolpate, colpi convergent at the poles in a
triangle enclosing a remnant of exine, pores non-protruding,
exine psilate or finely scabrate, grain diam 18–20 mm. Ovary
lightly pilose; style lightly pilose, included at anthesis, ulti-
mately exserted ca. 1 mm; ovules 6 or 7 at distal end of a
flattened placenta. Disc reflexed over dorsal side of ovary,
1.0–1.7 mm long, 0.2–0.3 wide, cylindric to oblong, thick, dark
red. Seeds 1.3 mm long, 1.5 mm wide, sub-circular, apex
rounded, margin rounded. Figures 4K, 5K, 12B.
Phenology—Collected in flower in April and in flower and
fruit in August, November, and December.
Distribution and Habitats—Venezuela, in Bolı́var state and
closely adjacent in Guyana; among rocks and in brown sand
along streams and rivers, in open wet sunny places, on slopes
in montane woodlands, in dense cloud forests, and in wet
gallery forests below waterfalls; 100–1450 m (Fig. 9).
Extinction Risk—The estimated risk if all IUCN guides
were to be followed is VU; EOO 5 9,702.67 km2 and AOO 5
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24 km2. One of the eight collection localities analyzed occurs
in a protected area.
Specimens Examined—Guyana. Cuyuni-Mazaruni, Paruima, 9 km W,
0.2–0.5 km E of Ararata scrub area, 5°490 36”N, 61°080 400W [5.82, -61.144], 5
Jul 1997, Clarke 5399 et al. (CAY, US-image); Potaro-Siparuni, Pakaraima
Mts., Mt. Wokomung, Suruwabaru DID Creek, E side riverine slopes
1–2 km from juncture with Yuarba River, 5°02’N, 59°54’W [5.033, -59.9], 9
Nov 1993, Henkel 4253 (NY). Venezuela.—BOLÍVAR: primeras faldas del
Paujil, 105 km sur de El Dorado, 26 Apr 1957, Bernardi 6802 (NY); slopes
along and above Salto El Danto, km 121–122 of Hwy 10, La Escalera, cloud
forest, 5°570 47”N, 61°230 300W [5.963, -61.391], 14 Nov 1997, Berry et al. 6590
(MO); 102 km de El Dorado hacia Sta. Elena, 29 Dec 1956, Foldats 2694 (NY);
Alto Rı́o Cuyuni, Rı́o Uiri-yuk, 10–15 km below Cerro Escalera [ca. 5.963,
-61.391], 17 Aug 1962, Maguire et al. 46715 (NY); 20 Aug 1962, Maguire et al.
46815 (MO, NY); near km marker 98, S of El Dorado along edge of rain
forest, 18 Dec 1979, Pruski & Steyermark 1369 (NY); Danto waterfall, in low
altitude cloud forest at km marker 118–119 S of El Dorado, 18 Dec 1979,
Pruski & Steyermark 1382 (NY); carretera a La Gran Sabana, Salto El Danto
[5.96, -61.38], 9 Apr 1988, Sastre et al. 8541 (MO); Cerro La Danta, NW of
Cerro Venamo [5.95, -61.38], 13 Apr 1960, Steyermark & Nilsson 10 (NY); vic.
road campamento at km 150 in valley of savanna of Rı́o Uarama below
Uarama-tepui, northeast of Luepa, 24–25 Apr 1960, Steyermark & Nilsson
634 (NY); Cerro Uananapan, S of Uei-tepui [5.016, -60.615] between Luepa
and Cerro Venamo, 25 Apr 1960, Steyermark & Nilsson 756 (NY); en el
drenaje del Rı́o Cuyunı́ al sur de El Dorado [6.715,-61.637], 22–28 Dec 1970,
Steyermark et al.104483 (MO); Sierra de Lema, sector La Escalera, al E de la
carretera, a unos 200 m del salto El Danto, 5°57.9’N, 61°23.7’W [5.95,
-61.383], 23 Apr 2006, Vivas et al. 1563 (MO).
In Cuphea dactylophora, all parts of the plant are densely
bifid-strigose, with very thin, short trichomes. The floral tube
has a well-developed spur, showy petals nearly half the length
of the floral tube, and an elongate, reflexed disc. Of importance
to identification of the species is the indistinct raceme, com-
posed of a few flowers on slender pedicels obscured by long
leaves. Cuphea curiosa, C. pleiantha, and C. rhodocalyx are closely
similar, but racemes are distinct. The four species form a
complex in Venezuela that will benefit from additional col-
lections and further study.
The only original material of Cuphea dactylophora is the
lectotype, which is a Schomburgk duplicate at Kew, sheet
000533053; the holotype at Berlin is no longer extant. There has
been great confusion about collection numbers and collection
localities of Robert and Richard Schomburgk (see van Dam
2002 for the most recent and thorough information on their
travels). The lectotype, cited as Schomburgk 1556a by Lourteig
167
(1986) does not bear that number. It retains, however, an
original label marked, 29, Cuphea, British Guiana, Schom-
burgk. That number is most likely a package number, not a
collection number (Van Dam 2002). The British botanist N. E.
Brown annotated the Kew specimen from the Bentham Her-
barium after seeing the holotype of C. dactylophora in Berlin in
1879. He noted that the holotype was numbered “1556” and
that Koehne said he had not seen this plant in any other
collection. The number 1556 appears to be a misreading by
Koehne for 155 S (1903: 128; Van Dam 2002: index). The “S”
was assigned to a separate set of collections made by Robert
Schomburgk and is referred to as “Rob. add. series” by Van
Dam (2002: 49). The type of C. dactylophora should not be
confused with the type of C. rigidula, which is also from the
Bentham Herbarium and bears the number number 155 S, or
from a second collection of C. ridigula, from the Hooker
Herbarium at K numbered 155 (see Cuphea rigidula for addi-
tional notes). The collection locality of the type of C. dactylophora
is unknown.
9. CUPHEA DISTICHOPHYLLA Lourteig, Mem. New York Bot.
Gard. 9: 357, Fig. 68. 1957. TYPE: VENEZUELA. Amazonas:
Cerro Sipapó (Paraqué) [4.957, -67.33], frequent in north
savanna, 17 Dec 1948, B. Maguire & L. Politi 27765 (ho-
lotype: NY-00386508!; isotypes: K, RB, US, as images!).
Subshrubs to 50 cm; stems erect to weakly decumbent,
from a woody base with fibrous roots, irregularly-branched;
internodes compressed distally, 1–4 mm long, much shorter
than subtending leaves; stem indumentum of two types:
predominantly bifid-strigose, trichomes thin, longitudinally to
irregularly oriented, some with arms semi-erect at a 45° angle;
sometimes also sparsely hirsute, trichomes bulbous-based,
purple, 0.5 mm long. Leaves ascending, rigid, closely imbri-
cate and stiffly distichous distally, gradually reduced in size in
the inflorescence; petioles 0–1.0 mm, thick, purple; blades
linear to lanceolate-linear, 10–18 mm long, 1.5–3.0 mm wide,
appearing uni-nerved, coriaceous, base cuneate, apex acute,
margin plane; surfaces green, sparsely bifid-strigose, adaxially
sometimes also sparsely hirsute, abaxially veins brown, not
visible on distal leaves. Racemes indistinct, terminal, compact,
short, leafy, simple; flowers alternate or sub-alternate,
crowded, partially obscured by leaves; pedicels ca. 2 mm
long; bracteoles ovate, 0.5 mm long, 0.5 mm wide. Floral tube,
including a horizontal, poorly developed spur 0.2 mm long
(0.8 mm long in Huber 13512) or non-spurred, 4.5–5.0 mm long,
1.5–2.0 mm wide, neck remaining wide in fruit, mouth flaring,
2.5–3.0 mm wide; outer surface dark purple-red, finely strigose
and sparsely purple hirsute as on stems; inner surface densely
villous distal to stamen insertion, lightly pilose proximally;
sepals equal, 0.5 mm long, 0.5 mm wide, deltate, apex acute;
epicalyx segments thickened lobes, 0.4 mm long, setose at
apex, shorter than sepals. Petals 6, fuchsia (Maguire & Politi
27646), pale purple (Huber 13.512), or white with purple base
and midveins (Maguire & Politi 27765), narrowly to broadly
obovate, 3.5–5.0 mm long, 2.0–3.0 mm wide. Stamens: 2 short
dorsalmost inserted slightly below other 9 in a horizontal line
at ca. 2/3 length of floral tube, none reaching sinuses of sepals;
filaments densely white villous except three ventral ones
glabrous, filament and anther color unknown; pollen oval-
triangular, tricolporate, non-syncolpate, pores not or scarcely
protruding, exine irregularly sinuously rugulate and partially
scabrate, grain diam 13–16 mm. Ovary pilose; style lightly
pilose, included; ovules 2 or 3. Disc horizontal to slightly
 168 SYSTEMATIC BOTANY
deflexed, sub-globose,1mm long, 0.6 mm wide, thick. Seeds
1.5–1.6 mm long, 1.5 mm wide, sub-circular, apex rounded,
margin rounded. Figures 4I, 14A.
Phenology—Collected in flower and fruit in December.
Distribution and Habitats—Venezuela, in Amazonas state,
Cuao-Sipapó Massif; in savanna and shrublands, on slopes, in
sand; 1400–1900 m (Fig. 8).
Extinction Risk—The estimated risk if all IUCN guides
were to be followed is CR; EOO 5 16.14 km2 and AOO 5
12 km2. Three collection localities were analyzed, all in pro-
tected areas.
Specimens Examined—Venezuela.—AMAZONAS: Atures, areniscas del
Cerro Cuao, Ca~ no Cabeza de Manteco, 73 km SE de Pto. Ayacucho, 5°06’N,
67°24’W [5.1, -67.4], Sep 1989, Fernandez et al. 6219 (NY); cumbre centro-
oriental del Macizo del Cuao, arbustales tepuyanos densos sobre vertiente
de arenisca inclinada hacia el SW, 4°58’ N, 67°19’W [4.966, -67.316], 10 Feb
1993, Huber 13.512 (P); Cerro Sipapó [4.957, -67.330], frequent in savanna on
southeast slopes of Peak I,12 Dec 1948, Maguire & Politi 27646 (NY, P, U-
image); Maguire & Politi 27764 (NY); Cerro Sipapó, savanna, N side of
middle Ca~ no Negro, 8 Jan 1949, Maguire & Politi 28217 (NY).
This Venezuelan endemic is distinguished by thick, as-
cending, rigid, strongly distichous, linear to linear-lanceolate
leaves that cluster on the distal few centimeters of the stems.
The flowers are bifid-strigose mixed with sparse, longer, thin
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trichomes, and produce only 2 or 3 ovules. The floral tube is
Fig. 14. A. Habit of Cuphea distichophylla, Maguire & Politi 27765 (RB), and flower, Huber 13512 (P). B. Habit and flower of C. insolita, isotype, Hoffman &
Marco 2260 (MO).
[Volume 44
minimally spurred and the disc horizontal within. The char-
acter suite indicates a closer relationship to the Colombian
endemics of the section than to the primarily Venezuelan
lineage (Table 1).
10. CUPHEA INSOLITA Lourteig, Bradea 8: 29. 1996. TYPE:
GUYANA. Cuyuni-Mazaruni region, Pakaraima Mountains,
Meanum River headwaters, 1.6 km W of Marali Falls, riv-
erside trail, 6°15’N, 60°27’W [6.25, -60.45], 4 Aug 1992, B.
Hoffmann & Marco 2260 (holotype: P-01901627-image!; iso-
types: MO!, US-image!).
Subshrubs to 50 cm; stems erect, one to several closely
arising from a woody rhizome with abundant secondary fi-
brous roots, much branched distally; internodes less than half
as long as subtending leaves, much shorter distally; stem
indumentum not bifid-strigose, of two types: hirsute, tri-
chomes simple, short, semi-erect, non-glandular; and sparsely
setose, trichomes multicellular, glandular, ca. 1.0 mm long.
Leaves spreading; petioles to 0.5 mm long; blades linear,
10–23 mm long, 1–2(–3) mm wide, appearing uni-nerved,
thickly membranaceous or coriaceous, base cuneate, apex
obtuse, margin plane, thickened, red-brown when dry; sur-
faces glabrous or nearly so, green, abaxially distinctly paler,
 2019] GRAHAM: CUPHEA SECTION AMAZONIANA
occasionally with sparse long thin trichomes, midvein slender,
red-brown; secondary veins not visible. Racemes distinct,
terminal, to 30 cm long, bracts reduced, almost hypsophylloid,
oblong to sub-obovate, 2.5 mm long, 1.0 mm wide; flowers
alternate to paired, crowded distally on the stem; pedicels
internodal, mid-way between nodes, 2.5–3.0 mm long; brac-
teoles flattened lobes ca. 0.1 mm long. Floral tube, including a
poorly developed spur 0.3–0.5 mm long, 3–4 mm long, 1 mm
wide, neck at 1/2–2/3 length of floral tube, scarcely contracted
in fruit, mouth flaring, 1.5–2.0 mm wide; outer surface color
unknown, indumentum as on the stems; inner surface villous
distal to stamen insertion, lightly pilose proximally; sepals
subequal, 0.5 mm long, 0.7–0.8 mm wide, deltate, apiculate;
epicalyx segments thickened lobes, dark-colored, 0.3 mm
long, much shorter than sepals. Petals 2 or 3, pink, 2 dorsals
oblong, 2 mm long, 1 mm wide, ventral petals 1 or 0, oblong,
2 mm long, 0.7 mm wide. Stamens: 2 dorsalmost very short,
glabrous or pilose, inserted slightly deeper than the others, 9
inserted at ca. 1/2–2/3 length of floral tube, lightly villous,
antesepalous stamens scarcely exceeding apices of sepals,
antepetalous ones reaching sinuses of sepals; filament and
anther color unknown; pollen oval-triangular, tricolporate,
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non-syncolpate, pores not or slightly protruding, exine
finely scabrate, grain diam 16–17 mm. Ovary lightly pilose;
style lightly pilose, ultimately exserted ca. 0.5 mm, sur-
passing the sepals; ovules 3–5. Disc horizontal to deflexed,
sub-deltoid, obtuse, thick. Seeds 1.5 mm long, 1.5 mm wide,
sub-circular, apex obtuse, margin rounded. Figures 2B, E, 4J,
5J, 14B.
Phenology—Collected in flower and fruit in August.
Distribution and Habitats—Western Guyana; steep slopes,
mixed forest, on riverside trails, and on sandstone boulders
along the Meanum river; 150–200 m (Fig. 9).
Extinction Risk—The estimated risk if all IUCN guides
were to be followed is CR; EOO not available and AOO 5
4 km2. One collection locality was analyzed, the type area,
which is outside protected areas.
Cuphea insolita, together with C. distichophylla, and C. bolivariensis
in Venezuela, have poorly developed spurs, horizontal to de-
flexed discs, low ovule numbers, and non-syncolpate pollen.
Unlike them, and uniquely in the section, C. insolita has only
simple trichomes. Flowers are alternate, subalternate, or paired
on internodal, not interpetiolar, pedicels. Reduction of petal
number from the normal six petals in Cuphea to two is also unique
in the section, although random reduction in petal number is
common elsewhere in the genus.
11. CUPHEA KUBEORUM Lourteig, Bot. Mus. Leafl. (Harvard
University) 16: 221, Fig. 35. 1954. TYPE: COLOMBIA. VAUPÉS:
Rı́o Kubiyú (tributary of Rı́o Vaupés), Cerro Ka~
nenda [ca.
1.0, -70.25], savannas about 15 mi upstream from mouth,
quarzite base, 800–900 ft, 10 Nov 1952, R. E. Schultes & I.
Cabrera 18306 (holotype: GH-0068449-image!; isotype:
COL!).
Subshrubs 10–45(–70) cm; stems one to several, erect, de-
cumbent, or prostrate and creeping from a woody rhizome,
much-branched, branch lengths and positions along the stem
irregular; internodes proximally longer than subtending
leaves, distally compressed on stems and branches, variable,
ca. 2–5 mm long; stem indumentum bifid-strigose, trichomes
appressed or arms partially erect at ca. 45° angle to stem.
Leaves decussate, gradually or not at all reduced in size in the
inflorescence, becoming irregularly crowded or fasciculate
169
distally, ericoid in aspect; petioles 0–1 mm; blades linear, 2–6
(–10) mm long, 0.5–1.0(–2.0) mm wide, appearing uni-nerved,
coriaceous, base cuneate, apex subacute, margin plane; sur-
faces adaxially green, glabrous or sparsely bifid-strigose, often
shining, abaxially paler and sparsely bifid-strigose, midvein
brown, secondary veins visible on widest proximal leaves.
Racemes indistinct, terminal, leafy, simple; flowers clustered
near stem apex, interpetiolar or internodal, opposite to sub-
alternate; pedicels 0.5–2.5 mm long; bracteoles ovate, 0.2 mm
long. Floral tube, including a well-developed, horizontal to
sub-descending, dorsally slightly enlarged spur 0.5–1.1 mm
long, 4–6 mm long, ca. 1.5–1.7 mm wide, neck slightly con-
stricted at 2/3 length of floral tube, mouth flaring, 2.0–2.5 mm
wide; outer surface green or purple, bifid-strigose; inner
surface white villous distal to stamen insertion, lightly villous
proximally; sepals sub-equal, 0.5–0.7 mm long, 0.3–0.5 mm
wide, deltate, apiculate; epicalyx segments thickened lobes, 6
cylindric, 0.3–0.5 mm long, shorter than sepals, without setae.
Petals 6, purple to pink, subequal, 2.0–3.5 mm long,
0.7–1.5 mm wide, oblong, apex obtuse. Stamens: 2 short
dorsalmost slightly deeper, other 9 inserted in nearly hori-
zontal line at ca. 2/3 length of floral tube, included, ante-
sepalous stamens reaching apices of sepals, antepetalous ones
proximal to sinuses of sepals; filaments white villous, filament
and anther color unknown; pollen oval-triangular, tricol-
porate, non-syncolpate, pores protruding, exine rugulate,
grain diam 17–18 mm. Ovary and style villous; style included
at anthesis, reaching apex of dorsalmost sepal; ovules 4(–6).
Disc slightly deflexed, 0.5–0.7 mm long, 0.4–0.5 mm wide,
globose or sub-deltoid, obtuse at apex, thick, dark red. Seeds
1.2–1.3 mm long, 1.5 mm wide, circular, base rounded, apex
rounded to obtuse, margin thinned. Figures 4A, 5B, 15A.
Phenology—Collected in flower and fruit from August to
December.
Distribution and Habitats—Endemic to Colombia, in De-
partments Amazonas, Caquetá, and Vaupés; savannas on
sandstone, and on wooded hills firmly rooted among sand-
stone rocks; 200–650 m (Fig. 16).
Extinction Risk—The estimated risk if all IUCN guides
were to be followed is LC; EOO 5 39,359.05 km2 and AOO 5
48 km2. Twelve of 13 collection localities analyzed were in
protected areas.
Additional Specimens Examined—Colombia.—AMAZONAS: Rı́o Caquetá,
nor-occidente de Araracuara, corregimiento de Pto. Santander, trocha a
comunidad Monochoa, 0°50’–55’S, 72°20’–25’W [-0.833, -72.333], 17 Nov
1993, Cárdenas et al. 4108 (MO); Rı́o Caquetá, Araracuara, camino a la
Sabana, 1° sabana de Tibeyes, 4 Jan 1977, Sastre & Raichel D. 5115 (P).
CAQUETÁ: Solano, Rı́o Mesay, Chorro Jacameya, extremo sur de la serranı́a
de Chiribiquete [0.916, -72.75], 25 Nov 1995, Cárdenas et al. 6866 (MO);
Campamento Norte, proximidades del campamento, 1°7’N, 72°50’W
[1.116, -72.833], 6 Dec 1990, Cardiel et al. 1014 (COL); Campamento Sur,
margen derecho del rio aguas arriba de la cascada grande, 0°55’N, 72°W
[0.916, -72.75], 6 Mar 1990, Estrada et al. 637 (COL); Sierra de Chiribiquete,
entre al Rı́o de los Musgos y el arco del triumfo, cerca a la pared con
pinturas, 1°05’N, 72°0’W[1.083, -72.666], 18–19 Aug 1992, Franco 3808
(COL, MO, P); Sierra de Chiribiquete, mesa encima de la Cueva de las
Pinturas, mesa facturada con profundas diaclasas, 1°05’N, 72°40’W
[1.083, -72.666], 21 Aug 1992, Palacios et al. 2400 (NY, P). VAUPÉS: Mitú,
[1.252, -70.233], Rı́o Vaupés, comunidad de Mandi [1.017, -70.383], for-
maciones de roca dura, vegetación de sabana, 29 Jul 1993, Martı́nez 522
(COL); Yapobodá [1.373, -70.908], 10 Dec 1943, Allen 3235 (MO); Rı́o
Kuduyari,Yapoboda, quartzite savanna near headwater, 4 Sep 1956,
Barclay et al. 532 (COL, NY); Rı́o Cubiyu, affluent du Rı́o Vaupés, 9–10
Nov 1952, Humbert & Schultes 27339 (NY, P); Ca~ no Cubiyú, comunidad
Indı́gena La Sabana, 1°15’N, 70°51’W [1.25, -70.85], 26 Apr 1993, Madri~ nán
et al. 1121 (MO); Rı́o Kuduyarı́, Yapoboda, 4–6 Oct 1951, Schultes & Cabrera
14267 (MO); 5–6 Oct 1950, Schultes & Cabrera 14389 (COL, P), 14 Aug 1960,
 170 SYSTEMATIC BOTANY [Volume 44
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Fig. 15. A. Habit of Cuphea kubeorum, Humbert & Schultes 27339 (P), and flower, Franco 3808 (MO). B. Habit and flower of C. maigualidensis, holotype,
Huber 12741 (P).

22600 (COL), 22605 (COL, MO), 22657 (COL); savanna off Rio Kubiyu [ca. Chajura, afluente occidental Rı́o Erebato, aprox. 100 km
1.033, -70.2], 25 Mar 1979, Soejarto & Lockwood 2417 (COL); Rı́o Karurú (en linea recta) al SW del Campamento Entrerı́os, 5°33’N,
(tributary of Rı́o Vaupés), mesa de Yambı́ [1.475, -71.349], savannah Goo-ran-
hoo-dá, 15–16 Apr 1953, Schultes & Cabrera 19177 (P); Rı́o Paraná- Pichuna
65°13’W [5.55, -65.216], 2100 m, 28 Mar 1988, O. Huber
(tributary of Rı́o Vaupés), general location 1°10’N, 70°30’W [1.166, -70.5], Jun 12741 (holotype: P-01901742-image!; isotype: VEN-
1953, Schultes & Cabrera 19913 (P); Mitú [1.252, -70.233] and vicinity, lower Rı́o image!).
Paraná-Pichuna, savanna at major rapids, 9 Sep 1976, Zarucchi 2011 (COL, P).
Subshrubs 30 cm; stems erect, branched from near the
Cuphea kubeorum stands out among the species of sect.
base, branches sparse, long; roots undescribed; internodes
Amazoniana in Colombia for the multitude of tiny, linear leaves
mostly equal to or longer than the subtending leaves; stem
that are crowded or fasciculate at the apices of short branches
indumentum a mix of two types: bifid-strigose, trichomes
along woody, decumbent or creeping stems. The indistinct,
closely appressed to the stem; and hirsute, trichomes erect,
few-flowered raceme is obscured by leaves at the stem apex.
bulbous-based, thin, dark red, 0.6–1.0 mm long. Leaves
The flowers are spurred, densely white villous within, and
spreading, not imbricate, very gradually reduced in size in
have a slightly deflexed disc. Closest morphologically are C.
the inflorescence; petioles 1–2 mm long; blades elliptic,
beneradicata, C. chiribiquetea, and C. stygialis. The species most
3.5–20.0 mm long, 1.5–5.0 mm wide, thickly membranaceous,
likely to be confused with C. kubeorum is C. beneradicata, which
base obtuse or cuneate, apex subacute to obtuse, margin plane;
is a more erect plant with longer, slender internodes, slightly
surfaces green, bifid-strigose, sometimes also with sparse long
longer leaves, more distinct racemes, and both bifid-strigose
trichomes, abaxially a prominent midvein and 2 pairs of
type and sparse longer trichomes on young growth. The two
secondary veins visible. Racemes indistinct, terminal, leafy,
species are sufficiently similar that they might prove con-
simple; flowers solitary, alternate, interpetiolar; pedicels 1 mm
specific once variation at the population level is better known.
long; bracteoles elliptic, 0.7 mm long. Floral tube, including a
12. CUPHEA MAIGUALIDENSIS Lourteig, Bradea 8:28. 1996. TYPE: rounded non-spurred base, 5.5–6.0 mm long, ca. 1.5 mm wide,
VENEZUELA. Bolı́var: Cede~
no, Sierra de Maigualida, sector neck slightly contracted at 2/3 length of floral tube, mouth
nor-oriental, altiplano tepuyano, cabeceras del Rı́o widely flaring, 2.5–3.0 mm wide; outer surface violet, strigose
 2019] GRAHAM: CUPHEA SECTION AMAZONIANA
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Fig. 16. Distribution of Cuphea kubeorum, C. philombria, C. saxatilis, C. stygialis, and C. sucumbiensis.
and hirsute as on stems; inner surface lightly pilose; sepals
equal, deltoid, 0.6 mm long, 0.6 mm wide; epicalyx segments
thickened lobes, half as long as sepals, with setae. Petals 6,
violet, 3.5–5.0 mm long, 2.5–3.5 mm wide, 2 dorsals slightly
smaller than 4 ventrals, all obovate-oblong or sub-orbicular.
Stamens: 2 short dorsalmost deepest, other 9 inserted in a
nearly horizontal line at 2/3 length of floral tube, all included;
filaments pilose, filament and anther color unknown; pollen
unknown. Ovary lightly pilose; style lightly pilose, included;
ovules 5. Disc reflexed, appressed to ovary wall (fide Lourteig
1996), cylindric, 1 mm long, 0.2 mm wide. Seeds 1.5 mm long,
1.3 mm wide, sub-circular, margin thick, rounded. Figure 15B.
Phenology—Collected in flower and fruit in March.
Distribution and Habitats—Venezuela, in Bolı́var state,
on a plateau in the Sierra de Maigualida; mossy river banks,
shady damp areas; 2100 m (Fig. 17).
Extinction Risk—The estimated risk if all IUCN guides
were to be followed is CR; EOO not available and AOO 5
4 km2. The species is known only from the type area, which is
not a protected area.
Cuphea maigualidensis is unusual in the section for the long,
slender internodes that are mostly equal to or longer than the
subtending leaves, even in the inflorescence. The description
above is based on a digital image of the holotype and on the
descriptions and line drawings by Lourteig (1996: 28; 2001: 75).
The type of Cuphea maigualidensis, Huber 12.741, is from the
same locality and habitat, and was collected on same day as a
paratype of C. curiosa var. oresbia, Huber 12.735 (P). The species
are not likely to be confused. Cuphea curiosa has short distal
171
nodes, imbricate distal leaves, and distinct inflorescences.
Cuphea maigualidensis, in addition to longer internodes with
typically well-spaced leaves, has an indistinct inflorescence
and petals more than half the length of the floral tube. Lourteig
(1996) considered C. maigualidensis most similar to C. distichophylla
and C. cataractarum and the three species to be part of a homo-
geneous group endemic to the Guayana region. The numerous
differences in general habit and morphology among the three
species, however, do not suggest a close relationship.
13. CUPHEA PHILOMBRIA Lourteig, Bot. Mus. Leafl. (Harvard
University)16: 223, Fig. 36. 1954. TYPE: COLOMBIA. META:
Cordillera de La Macarena, (extreme northeast), macizo
Renjifo [3.105, -73.934], summit and surroundings, on rocky
banks at river’s edge, roots embedded in moss, in shaded,
damp areas, 1300–1900 m, 6–20 Jan 1951, J. M. Idrobo & R. E.
Schultes 1062 (holotype GH-00068452-image!; isotypes: COL
(32)!, MO!, NY!; from images P!, US (33)!).
Cuphea killipii Lourteig, Phytologia 60: 24.1986. TYPE: COLOM-
BIA. Meta: north side of river near junction of Güejar and
Sanza (Zanza) rivers [3.278, -73.916], N end of Cordillera
de La Macarena, S. G. Smith and J. M. Idrobo 1480 (holo-
type: GH-00068448-image!; isotypes: MO!; from images,
COL!, P!, UC!).
Subshrubs 30–50 cm; stems erect to decumbent, one to
several from a woody rhizome with abundant fibrous roots,
much-branched; internodes less than half as long as the
subtending leaves; stem indumentum a mix of two or three
 172 SYSTEMATIC BOTANY
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Fig. 17. Distribution of Cuphea maigualidensis, C. pleiantha, C. rhodocalyx, C. rigidula, and C. trisperma.
types: bifid-strigose, trichomes thin, closely appressed to the
stem; setose, trichomes coarse, multicellular, slightly enlarged
at the base, ascending, red-brown, 0.3 mm long; and/or
sparsely hirsute, trichomes thin, multicellular, glandular, to
0.5 mm long. Leaves spreading, loosely imbricate distally on
the stem, gradually reduced in size in the inflorescence; pet-
ioles 0–0.5 mm long; blades linear to linear-lanceolate,
7–30 mm long, 0.5–4.0 mm wide, appearing uni-nerved, co-
riaceous, base cuneate, apex subacute, margin slightly incurved;
surfaces adaxially green, glabrous (strigose in Caquetá), abaxially
paler, bifid-strigose, midvein brown, 2–4 pairs of thin secondary
veins visible on widest, most proximal leaves. Racemes in-
distinct, terminal, leafy, simple; flowers interpetiolar, alter-
nate, crowded distally on the stem, partially obscured by the
leaves; pedicels 2.5–4.0(–7.0) mm long; bracteoles sub-
orbicular, thick, inconspicuous, 0.3 mm long. Floral tube,
including a well-developed, horizontal to slightly descend-
ing spur 0.5–1.0 mm long, 4.0–8.5 mm long, 1–1.5 mm wide,
neck slightly contracted at ca. 1/2–2/3 length of floral tube,
mouth widely flaring, 2.0–4.0 mm wide; outer surface green
or purple [fide Lourteig,1986], finely bifid-strigose with
sparse to abundant, coarse hirsute/setose trichomes on the
veins, inner surface white villous distal to stamen insertion,
[Volume 44
lightly pilose to nearly glabrous proximally; sepals subequal,
0.8–1.0 mm long, 0.7–1.2 mm wide, ventrals slightly wider, all
deltate, acute to apiculate; epicalyx segments thickened
lobes, 6 cylindric, 0.3–0.5 mm long, dark red-purple, shorter
than sepals, 2 dorsal segments with 1–few terminal setae, others
with 0–2 setae. Petals 6, purple to white with pinkish tint, or 2
dorsals purple, 4 ventrals pale purple to white, all approx.
equal, broadly elliptic to obovate, 2.5–4.5 mm long (–7.0 mm
fide Lourteig 1986), 1.0–3.5 mm wide. Stamens: 2 short dor-
salmost slightly deeper, other 9 inserted in nearly horizontal line
at ca. 2/3 length of floral tube, all included, none reaching si-
nuses of sepals; filaments white villous, except three ventral
antesepalous ones glabrous; filament and anther color un-
known; pollen oval-triangular, tricolporate, non-syncolpate,
colpi short, pores not or slightly protruding, exine coarsely ir-
regularly rugulate, thick at poles, grain diam 17–18 mm. Ovary
and style sparsely pilose to glabrous; style included; ovules 8–15.
Disc horizontal to slightly deflexed, 0.4–0.5 mm long, 0.5 wide,
sub-deltoid, obtuse at apex, dark red. Seeds 1.2–1.3 mm long,
1.0–1.2 mm wide, obovate, base and apex rounded, margin thick,
rounded. Figures 2G, H, 3A, 4B, 5C, 18A.
Phenology—Collected in flower and fruit from November
to March, and in July and August.
 2019] GRAHAM: CUPHEA SECTION AMAZONIANA
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Fig. 18. A. Habit and flower of Cuphea philombria, isotype, Idrobo & Schultes 1062 (MO). B. Habit of C. pleiantha, Maguire & Maguire 35376 (NY), and
flower, Steyermark et al.123626 (P).
Distribution and Habitats—Colombia, in Departments Caquetá
and Meta, Cordillera de La Macarena; on plateaus in permanently
wet low depressions, on sandstone, on rocky river banks and along
river margins, in shaded damp areas in moss, and in sand of river
beaches; 300–625 and 1000–1900 m (Fig. 16).
Extinction Risk—The estimated risk if all IUCN guides
were to be followed is VU; EOO 5 16,368.85 km2 and AOO 5
44 km2. Thirteen collection localities were analyzed, only one
was within a protected area.
Additional Specimens Examined—Colombia.—CAQUETÁ: San Vicente
del Caguán, carr. en construcción San Vicente-Neiva, tramo entre la
Campana (casa del paisa) al Rı́o Pato, Vereda Las Perlas [2.415, -74.744], 27
Jul1990, Betancur et al. 1898 (MO, NY); Belén de los Andaquies, vereda Las
Verdes, setor bocana quebrada Los Verdes,vegetación a corolla de que-
brada, 1°360 21.5”N, 75°520 46.30W [1.605, -75.879], 26 Jul 2011, Cardenas et al.
42119 (NY); Florencia [1.615, -75.5664], Quebrada de las Perdices,
matorrales, 29 Mar 1940, Cuatrecasas 8846 (COL); Rı́o Hacha inter Venecia
[1.563, -75.532] et Florencia, 5 Aug 1926, Woronow & Juzepczuk 6454 (MO). —META:
Carr. Villavicencio-Acacias [3.991, -73.766], márgenes de los Rı́os Sardinita y
Acacias, 13 Mar 1986, Fernández A. et al. 5629 (COL); Vista Hermosa [3.123,
-73.751], trail to falls of Rı́o Sardinata, Serranı́a de la Macarena, 2 Feb 1985,
Henderson 116 (COL, excluding NY specimen); Sierra de La Macarena, Vereda El
Tablazo, rivera alta del Rı́o Güejar, 29 May 1973, Garcı́a et al. 511 (COL); Mt.
Macarena, plateau [ca. 2.854, -73.944], 19 Jan 1942, Gilliard s.n. (NY, P); Reserva
Nacional de La Macarena, Rı́o Guapaya, 16 Mar 1957, Idrobo 2431 (COL, NY);
Sierra de La Macarena, region norte, vertiente oriental, union de los Rı́os Sansa y
Güejar [3.278, -73.916], 22 Feb 1956, Idrobo & Jaramillo 2047 (COL, MO); Sabanas de
173
San Juan de Arama [3.346, -73.889], margen izquierda del Rı́o Güejar,
alrededores “Los Micos” [3.281, -73.869], 5–20 Dec 1950, Idrobo & Schultes
564 (COL); Cordillera La Macarena, Idrobo & Schultes 1061 (COL, P);
Cordillera La Macarena, mesa del Rı́o Sansa, 23 Jan 1951, Idrobo & Schultes
1288 (COL); Sierra de la Macarena, Rı́o Guapaya, 29 Nov 1949, Philipson
et al. 1609 (COL); Reserva Macarena, San Juan de Arana [3.346, -73.889],
Cuchilla del Neme [3.278, -73.916, left side Rı́o Güejar, 21 Jan 1968, Thomas
et al. 1467 (COL, P 3 2).
Cuphea philombria is recognized by the combination of: in-
distinct racemes, stems and flowers with a mix of abundant,
appressed, thin bifid trichomes and coarse, ascending, red-
brown setae; and by the high number of ovules. The leaves are
generally glabrous adaxially and strigose abaxially except in
the Caquetá specimens, where they are strigose on both sur-
faces. The presence of one or two setae at the apices of the
epicalyx segments is a striking, but inconstant, feature.
Cuphea philombria, including the synonymous C. killipii, is
most similar to and separated with difficulty from C.
sucumbiensis. The stem indumentum, floral size and high ovule
number are the same or overlap quantitatively. Plants are taller
and more substantial in C. philombria, with branches spreading
more than closely ascending, and leaves are longer on average.
Seeds of C. philombria are slightly larger than in C. sucumbiensis
(1.2–1.3 mm vs. 1.0–1.1 mm long). The racemes are indistinct in
C. philombria and distinct in C. sucumbiensis. Cuphea philombria
 174 SYSTEMATIC BOTANY
ranges along the Sierra de la Macarena, Colombia, to the north
and east of C. sucumbiensis, whereas C. sucumbiensis occurs in
Andean pre-montane forests of Colombia and Ecuador, and
along rivers on both sides of the Colombia-Ecuador border.
The two species are allopatric to the extent known.
Cuphea philombria and C. killipii were taxonomically sepa-
rated primarily by the dorsal petals darker than the ventrals in
C. killipii, compared to either uniformly purple petals in C.
philombria (e.g. in Idrobo & Schultes 1062 and Betancur et al. 1898)
or petals white with pink tint (Idrobo & Jaramillo 2047). Petal
color is unreliable and inadequate as the sole basis for species
delimitation in Cuphea. Other differences attributed to C. killipii
(Lourteig 1986), e.g. size of petals and spur, shape of disc, and
degree of visibility of leaf trichomes, are not sufficiently dif-
ferent from C. philombria to support recognition of two taxa.
14. CUPHEA PLEIANTHA Lourteig, Mem. New York Bot. Gard.
9: 356, Fig. 67. 1957. TYPE: VENEZUELA. Amazonas: Serranı́a
Yutajé [5.680, -66.075], Rı́o Manapiare, common in right
branch of Ca~ no Yutajé, 9 Feb 1953, B. & C. K. Maguire
35126 (holotype: NY-00386521!; isotypes: images P! and
US!).
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Subshrubs, 30–50 cm; stems erect, sturdy, from a woody
rhizome with abundant fibrous roots, branched, virgate dis-
tally in the inflorescence; internodes mostly less than half as
long as subtending leaves; stem indumentum of two types:
primarily bifid-strigose, trichomes very thin, closely appressed
to the stem, 1 mm long; rarely also sparsely hirsute, trichomes
erect, thin, 0.5 mm long. Leaves ascending, imbricate, 6
abruptly reduced in size in the inflorescence; petioles
1.0–1.5 mm long; blades narrowly elliptic to lanceolate-linear,
15–45(–60) mm long, 1–7 mm wide, appearing uni-nerved,
coriaceous, base cuneate, apex subacute to obtuse, margin
plane, thickened, especially on larger leaves; surfaces green,
adaxially glabrous or finely strigose, abaxially sparsely bifid-
strigose, midvein only or rarely midvein and 3 pairs of thin
secondary veins visible. Racemes distinct, terminal, bracteate,
simple or compound, with 2–10 branches, to 8 cm long, erect;
bracts lanceolate, thickly membranaceous, 1–4 mm long,
0.7–1.0 mm wide; flowers solitary, alternate, crowded distally
on the raceme; pedicels 1–2 mm long; bracteoles ovate-oblong,
0.5 mm long. Floral tube, including a rounded, non-spurred
base, 5.5–7.0 mm long, 1.5–2.0 mm wide, neck slightly con-
tracted at ca. 2/3 length of floral tube, mouth widely flaring,
ca. 2.5–3.0 mm wide; outer surface purple, sparsely to mod-
erately bifid-strigose, appearing glabrous, trichomes very thin,
arms short; inner surface purple, densely villous distal to
stamen insertion, glabrous proximally; sepals subequal,
dorsalmost sepal lighter in color, distinctly larger, 1.2 mm
long, 1.2–1.4 mm wide, broadly deltate, the others 0.7 mm
long, 1.0 mm wide, deltate, apex acute; epicalyx segments
thickened lobes, glabrous, dark-purple, 0.7–1.2 mm long,
shorter than sepals. Petals 6, unequal, 2 dorsals 3 mm long,
2 mm wide, slightly smaller than 4 ventrals, all petals light
purple or white with purple base, oval to obovate. Stamens: 2
short dorsalmost inserted at about same level as the other 9,
in a horizontal line at ca. 2/3 length of floral tube, included,
none reaching sinuses of sepals; filaments white villous except
3 ventral antesepalous filaments glabrous, purple or colorless,
anthers dark purple; pollen triangular, tricolporate, syn-
colpate, colpi convergent at the poles in a triangle enclosing a
remnant of exine, pores not protruding, exine psilate, grain
diam 18–19 mm. Ovary villous distally, pilose proximally; style
[Volume 44
sparsely pilose, included; ovules 7–9, at the distal end of a
flattened placenta. Disc reflexed and appressed to ovary,
0.9–1.2 mm long, 0.3–0.4 mm wide, cylindric to slightly flat-
tened and oblong, thick, dark red. Seeds 1.1 mm long, 1.3 mm
wide, sub-circular in outline, base rounded, apex rounded,
margin rounded. Figures 4F, 5G, 18B.
Phenology—Collected in flower and immature fruit in
February.
Distribution and Habitats—Venezuela, a narrow endemic
in Amazonas state, in the Cerro Yutajé region; forests and river
margins with many boulders, among rocks in the river;
150–300 m, 830 m, and 1300–1500 m (Fig. 17).
Extinction Risk—The estimated risk if all IUCN guides
were to be followed is EN; EOO 5 336.39 km2 and AOO 5
16 km2. Of the four collection localities analyzed, one is in a
protected area.
Specimens Examined—Venezuela.— AMAZONAS: Atures: Ca~ no Yutajé at
S base of Serranı́a de Yutajé, 5°38’N, 66°06’W [5.633, -66.1], 17 Feb 1987,
Holst & Liesner 3069 (MO, NY); Serranı́a Yutajé, Rı́o Manapiare, common in
right branch of Ca~ no Yutajé, 9 Feb 1953, Maguire & Maguire 35102 (NY, P);
common among boulders and rocks in streambed of middle section of main
branch, left fork of Ca~ no Yutajé, 15 Feb 1953, Maguire & Maguire 35246
(NY32, P, RB); common among boulders, Ca~ no Yutajé, 23 Feb 1953,
Maguire & Maguire 35376 (NY, L-image); base del Cerro Yutajé, a lo largo
del rı́o Yutajé, vecindades de la Chorrera, rı́o arriba de la pista del aero-
puerto, 5°37’N, 66°8’W [5.616, -66.133], 8 Nov 1980, Steyermark et al. 123626
(P-image).
Cuphea pleiantha is notable for: compound, highly floriferous
inflorescences composed of up to 10 slender long branches; for
long, very thin trichomes accompanying the bifid-strigose
stem indumentum; and for relatively large, narrowly ellip-
tic, ascending leaves. The species is morphologically similar to
some collections of C. curiosa. Both can have distinct com-
pound racemes, poorly developed spurs, and reflexed discs.
Cuphea curiosa does not reach the size of C. pleiantha and is a
more fragile plant than C. pleiantha, with slender, delicate
inflorescences, and a bifid-strigose stem indumentum that is
sometimes also hirsute. In C. pleiantha, the ascending leaves on
the distal half of the stem are coriaceous, narrow, and generally
without visible secondary venation. In C. curiosa, the distal
leaves are more membranaceous and spreading, and sec-
ondary venation is visible. Collection labels record color of the
petals of C. pleiantha in the main branch of Ca~ no Yutajé as
lavender or purplish, and in the right branch as bi-colored
purple and white.
15. CUPHEA RHODOCALYX Lourteig, Mem. New York Bot.
Gard. 9: 355. 1957. TYPE: VENEZUELA. Amazonas: Cerro
Marahuaka [ca. 3.5, -65.5], along streams, montane slopes
at 1000 m alt., 9 May 1949, B. Maguire & B. Maguire, Jr.
29201 (holotype: NY-00386524!; isotypes: as images-F!,
P 3 2!, US!)
Subshrubs 15 to 60 cm; stems erect or decumbent, one or
more from a woody rhizome with abundant fibrous roots,
branched, young stems quadrangular proximal to the nodes
from decurrent leaf bases; internodes short, less than one-third
as long as subtending leaves; stems glabrous or very sparsely
bifid-strigose. Leaves decussate, ascending, most often
abruptly reduced in size in the inflorescence; petioles 0–1 mm
long; blades narrowly elliptic to lanceolate-linear, 10–25 mm
long, 1–4 mm wide, appearing uni-nerved except at the most
proximal nodes, thickly membranaceous, base cuneate, apex
subacute, margin plane; surfaces adaxially green, abaxially
green or light brown, both surfaces glabrous or sparsely
strigose, abaxial midvein prominent, dark brown. Racemes
 2019] GRAHAM: CUPHEA SECTION AMAZONIANA
distinct, sometimes weakly so, terminal, leafy to bracteate,
simple and compound; flowers solitary, alternate, crowded
among distal bracts, interpetiolar; pedicels 1–2 mm long,
sturdy; bracteoles 0.4–0.7 mm long, 0.2 mm wide. Floral tube,
including a poorly developed, horizontal spur to 0.3 mm long
or non-spurred, 4.5–6.0 mm long, 1.5–2.0 mm wide, in fruit
ampullaceous and neck slightly contracted at 2/3 length of
floral tube, mouth widely flaring, 2.5–3.5 mm wide; outer
surface light purple, glabrous or sparsely bifid-strigose, tri-
chomes very thin; inner surface villous distal to stamen in-
sertion, lightly pilose proximally; sepals subequal, dorsal
sepal slightly larger, all 0.7–1.0 mm long, 1 mm wide, deltate,
apex acute; epicalyx segments thickened lobes, 0.3–0.5
(–0.7) mm long, shorter than the sepals, glabrous, dark-
colored. Petals 6, subequal, bi-colored, 2 dorsals purple, ob-
ovate, 2.5–4.0 mm long, 1.0–1.5(–2.5) mm wide, 4 ventrals pale
purple or white, narrowly obovate, 3–4 mm long, 1.5(–3.5) mm
wide, or all uniformly pale purple (fide Lourteig 1986). Sta-
mens: 2 short dorsalmost inserted at ca. same level as other 9,
all in a horizontal line at 1/2–2/3 length of floral tube, in-
cluded, not reaching apices of sepals; filaments white villous,
except 3 ventral antesepalous filaments glabrous, filament and
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anther color unknown; pollen triangular, tricolporate, syn-
colpate, colpi convergent at poles in a triangle enclosing a
remnant of exine, pores not protruding, exine psilate to in-
distinctly shallowly rugulate, grain diam 18–19 mm. Ovary
lightly pilose; style lightly pilose, ultimately reaching apices of
the sepals or exserted 0.5 mm; ovules 5–9. Disc erect, short,
0.4–0.6 mm long, 0.2–0.3 mm wide, slightly flattened, oblong,
thick, dark-colored. Seeds 0.9–1.4 mm long, 1.1–1.5 mm wide,
circular or sub-circular in outline, slightly wider than long,
apex rounded, margin rounded. Figures 3B, 5H, 19A.
Phenology—Collected in flower and fruit in October and
January to May.
Distribution and Habitats—Venezuela, in Amazonas state;
along streams and riverbanks and on montane forest slopes;
130–1200 m and 1500–1600 m (Fig. 17).
Extinction Risk—The estimated risk if all IUCN guides
were to be followed is EN; EOO 5 208.86 km2 and AOO 5
24 km2. Six collection localities were analyzed, all from pro-
tected areas on two neighboring massifs.
Specimens Examined—Venezuela.—AMAZONAS: en la ribera del rı́o,
Cerro del Duida, Jan-Feb 1969, Fari~ nas et al. 408 (MO); Atabapo, slope of
Cerro de Marahuaka, Rı́o Yameduaka, arriba, 3°38’N, 65°28’W [3.633,
-65.466], 17–18 Feb 1985, Liesner 17617 (MO, P); Atabapo, below Salto Los
Monos on tributary of headwaters of Rı́o Iguapo, 3°35’N, 65°23’W [3.583,
-65.383],11 Mar 1985, Liesner 18510 (P); slopes Cerro Marahuaka, “Sima”
area, 3°43’N, 65°30’W [3.71, -65.5], 15 Oct 1988, Liesner 24871 (MO, P); in
saddle between Duida and Marahuaka near base of Duida, 3°34’N,
65°32’W [3.566, -65.533], 27 Oct 1988, Liesner 25502 (MO, P); Atabapo,
“Sima Camp” south-central portion of forested slopes along eastern branch
of Ca~
no Negro, 3°43’N, 65°31’W [3.716, -65.516], 21–22 Feb 1985, Steyermark &
Holst 130645 (MO, NY, P).
Cuphea rhodocalyx is a small shrub with a strong woody base,
glabrous or sparsely strigose stems that are noticeably qua-
drangular below the nodes on young growth due to decurrent
leaf bases. Abundant slender leaves are ascending, closely
imbricate, uni-nerved, and nearly glabrous. Flowers are also
nearly glabrous and inflated in appearance in fruit. The short
disc stands erect. I have not seen it as strongly reflexed as
originally illustrated in the protologue. The collection, Liesner
25502 (MO), differs by leaves sparsely strigose on both sur-
faces; Liesner 18510 has exceptionally long floral tubes, 7–9 mm
long, and larger petals.
175
Cuphea rhodocalyx forma setosa Lourteig (1964), later cited as
var. setosa (Lourteig 1986, 2001), was elevated in rank as C.
cardonae Lourteig (1986) based primarily on the occurrence of
setae on the stems, with the additional mention of smaller
bracteoles occurring lower on the pedicel, and visible sec-
ondary leaf veins. Cuphea rhodocalyx was restricted to plants
with glabrous or sparsely strigose stems.
Four species endemic to central and southern Venezuela,
Cuphea rhodocalyx, C. cardonae, C. curiosa, and C. dactylophora,
share many features and remain difficult to separate taxo-
nomically. Cuphea rhodocalyx differs from the others by leaves
that appear uni-nerved; in the other three species, secondary
venation is visible. Cuphea cardonae displays slightly qua-
drangular young stems as in C. rhodocalyx and differs from it
by more indumentum, a longer cylindrical disc and, possibly,
different petal color. It remains questionably distinct from C.
rhodocalyx. Cuphea dactylophora has spreading, wider leaves
and a long, reflexed spur. Cuphea curiosa is morphologically
similar to C. rhodocalyx, but has larger leaves, up to 40 mm
long, and 8 mm wide, and more distinctly bracteate racemes.
The most numerous collections of C. curiosa are from north of
the few C. rhodocalyx localities; the two species may be sym-
patric in the southernmost part of the range of C. curiosa.
Floral tube and petal color, according to collection data, vary
among the four species and appear to vary within the ex-
tensively collected C. curiosa. Colors are missing on most
exsiccatae in sect. Amazoniana and this is the case in C. rho-
docalyx. Label information describes the floral tube as lavender
and petals as white, lavender, or purple. The specific epithet
rhodocalyx refers to the “rosado-violáceo” floral tube. Petals are
described as “violáceos,” and in the key to the section as
“rosados” (Lourteig 1986). In C. curiosa, the floral tube is
described on specimen labels variously as violet or green, and
petals as white to pale purple, pink, rose-purple, white with
purple vein or base, or bi-colored with dorsal petals darker
than the ventrals. In C. cardonae petals are white with purple
claws, and in C. dactylophora they are pale purple with a darker
purple base or white with purple midvein.
Until more extensive morphological data is available, I
accept Cuphea rhodocalyx, C. cardonae, Cuphea curiosa, and C.
dactylophora as distinct species in Venezuela, but acknowledge
the need for additional studies to test species limits and the
relationships suggested by the current evidence.
16. CUPHEA RIGIDULA Bentham, J. of Bot. 2: 316.1840, non C.
rigidula Seemann, Bot. Voy. Herald 1(4):121.1854. LECTO-
TYPE (designated, as holotype, by Lourteig 1964): BRAZIL.
Roraima: Rı́o Parima, near Meretani Mts. (5 Rio Uraricoera
[Parima], Merewari or Merevari Mts.; aprox. 3.333, -63.0),
Robert Schomburgk 155 S (K-000532982-image!; isotype: K-
000532981-image!).
Subshrubs, to 30 cm, stems erect, brown-red, branched,
branches dichotomously branched; roots undescribed; in-
ternodes half or less than half as long as subtending leaves;
stem indumentum of two types: predominantly coarsely
setose, trichomes abundant, erect to slightly appressed,
bulbous-based, typically red-brown, 0.5–1.0 mm long; and
inconspicuously strigose, trichomes bifid, appressed to the
stem, longest arm oriented distally. Leaves loosely arranged,
ascending, becoming imbricate distally, sessile, abruptly re-
duced in size in the inflorescence; petioles 0; blades broadly
elliptic-lanceolate, 15–43 mm long, 5–13 mm wide, thickly
membranaceous, base cuneate, apex subacute to acute, margin
 176 SYSTEMATIC BOTANY
Copyright (c) American Society for Plant Taxonomists. All rights reserved.
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Fig. 19. A. Habit of Cuphea rhodocalyx, Steyermark & Holst 130645 (P), and inflorescence, Liesner 24871 (MO). B. Habit and flower of C. rigidula, Cardona
3013 (NY).
plane; surfaces adaxially dark green, abundantly setose, setae
1.0–1.7 mm long (–2.5 mm long fide Lourteig 2001: 70),
abaxially paler green, finely bifid-strigose, midvein and 2 pairs
of secondary veins visible on both surfaces. Racemes distinct,
terminal, bracteate, dichotomously compound; flowers soli-
tary, alternate, crowded distally on the stem, partially ob-
scured by leaves and bracts; pedicels 0.5–1.0 mm long;
bracteoles lanceolate, ca. 1 mm long, 0.3 mm wide. Floral tube,
including a poorly developed, horizontal spur to 0.2 mm long,
or non-spurred, 5.5–6.0 mm long, ca. 1.5 mm wide, dorsally
convex in fruit, neck slightly narrowed at 2/3 length of floral
tube, mouth widely flaring, ca. 2.5–3.0 mm wide; outer surface
purple, finely bifid-strigose, appearing nearly glabrous, with
setae or not; inner surface lightly villous distal to stamen in-
sertion, lightly pilose proximally except villous along the veins
of the 2 dorsalmost stamens; sepals subequal, the 3 dorsalmost
slightly larger, wider, 1.2 mm long, 1.2 mm wide, the others
0.7 mm long, 0.7 mm wide, all deltate, apex acute; epicalyx
segments prominent, thickened lobes, dark-colored, 0.3–1.0 mm
long, the dorsal ones longer than the sepals. Petals 6, pink,
obovate to sub-spatulate, size uncertain, 2 dorsals darker at the
narrowed base, 4 ventrals short-clawed. Stamens: 2 short dor-
salmost inserted at nearly same level as other 9 in a shallow V-
shape at ca. 2/3 length of floral tube, antesepalous stamens
not reaching apices of sepals, antepetalous ones not reaching
[Volume 44
sinuses of sepals; filaments of 2 antesepalous stamens
sparsely villous, all others glabrous, filament and anther
color unknown; pollen oval-triangular, tricolporate, syn-
colpate, colpi convergent at the poles in a triangle enclosing a
remnant of exine, pores slightly protruding, exine scabrate,
nearly psilate, grain diam 19–20 mm. Ovary lightly pilose;
style lightly pilose, ultimately extended to apices of the sepals
or exserted ca. 0.5 mm; ovules 9 or10. Disc erect, 0.5–0.6 mm
long, 0.2 mm wide, cylindric to slightly flattened, obtuse,
thick, dark-colored. Seeds 1.2–1.5 mm long, 1.2 mm wide,
oblong to sub-obovate, base rounded, apex rounded, margin
rounded. Figures 4H, 5I, 19B.
Phenology—Collected in flower and fruit in April.
Distribution and Habitats—Venezuela, in Bolı́var state,
and Brazil in Roraima state; rocky outcrops at the base of falls;
680 m (Fig. 17).
Extinction Risk—The estimated risk if all IUCN guides
were to be followed is CR; EOO not available and AOO 5
4 km2. One collection locality was analyzed; it occurs in a
protected area.
Specimens Examined—Venezuela.—BOLÍVAR: Alto Caura, frecuente
entre los pe~
nascos [rocky outcrops] situados al pie de Salto Mareveni, en la
falda del Cerro Mareveni [4.5, -64.166], Apr 1965, Cardona 3013 (NY, P-image).
Cuphea rigidula is one of two species in sect. Amazoniana with
dichotomous branching, C. blackii, an endemic species of
 2019] GRAHAM: CUPHEA SECTION AMAZONIANA
French Guiana and Brazil, being the other. Cuphea rigidula is
further distinguished by abundant, coarse, brown setae on the
stems and leaves; which also may occur in a less coarse form
among the finely bifid trichomes of the floral tube. Cuphea
rigidula was first questionably classified in sect. Heteranthus
(Koehne 1877) because, although it had the requisite dis-
tinct bracteate inflorescence, it lacked the other primary de-
fining feature of that section, unequal maturation of paired
flowers on the raceme. Cuphea rigidula was subsequently
transferred to sect. Amazoniana (Lourteig 1986; see Morpho-
logical Relationships).
The lectotype of Cuphea rigidula, Kew sheet 000532982, from
the Bentham Herbarium, is from a set of collections made by
Robert Schomburgk that was numbered apart from his pri-
mary series and identified by the letter “S” following the
collection number. It is preferentially cited as “Rob. add. Ser.
1” (Van Dam 2002). Mounted on the same sheet as the lec-
totype is a second Schomburgk specimen of C. rigidula, Kew-
000532981, annotated “155” and “Guiana 1840” that was part
of the Hooker Herbarium. It appears to be from the same plant
population, morphologically identical to and at the same
flowering stage as the lectotype, and can be reasonably con-
Copyright (c) American Society for Plant Taxonomists. All rights reserved.
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sidered an isotype.
At the time Schomburgk collected C. rigidula, the boundaries
between Venezuela, Brazil, and Guyana were not fully re-
solved. Van Dam, following extensive research on the plant
collections of Robert and Richard Schomburgk, supplied the
collection locality for the holotype of C. rigidula, Robert
Schomburgk 155 S, as “Rio Parima near Meretani Mts” (Van
Dam 2002: Index 2). That locality today is in the state of
Roraima, Brazil (Huber et al.1984).
The two Kew specimens of C. rigidula, the lectotype,
Schomburgk 155.S from the Bentham Herbarium, and Schom-
burgk 155 from the Hooker Herbarium, are not to be confused
with the lectotype of C. dactylophora, marked Schomburgk 1556,
which was almost certainly a misreading of 155 S, from an
unknown locality. Van Dam (2002: 97) makes note of the
duplicate numbers, listing C. dactylophora as 155 S and C.
rigidula as 155.S*.
17. CUPHEA SAXATILIS S.A.Graham, sp. nov. TYPE: COLOMBIA.
Meta: La Macarena, Zona de Preservación del AMEM,
vereda cachivera, Parque la Macarena, por el camino que
conduce del Rı́o Guayabero al Ca~ no Cristales, 2°200 20”N,
73°430 000 W [2.338, -73.716], 1 Aug 2000, R. López C., M
Gaitán, J. L. Ramı́rez, & M. Zubieta 6729 (holotype: COL!;
isotype: COAH).
Diagnosis: Subshrubs 30–45 cm; stems erect, suffrutescent,
internodes mostly shorter than subtending leaves, abundantly
bifid-strigose. Leaves lanceolate to elliptic, 5–22 mm long,
2–8 mm wide, midvein and secondary veins prominent
abaxially. Racemes distinct. Floral tube including 1–2 mm long
spur, 6–8 mm long; sepals subequal, shallowly deltate. Petals
6, pink or fuchsia, 2 dorsal petals smaller than the 4 ventrals.
Pollen tricolporate, non-syncolpate, exine shallowly rugulate.
Ovules 5 or 6. Disc horizontal to deflexed, globose to sub-
deltoid. Seeds 1.5–1.7 mm long.
Subshrubs 30–45 cm; stems one to several from a woody
rhizome, sturdy, erect, suffrutescent, branched, some branches
as long as primary stem; internodes mostly equal to or shorter
than subtending leaves, except most proximal and most distal
internodes sometimes longer than subtending leaves; stem
indumentum of 2 types: abundantly bifid-strigose, trichomes
177
both appressed or with arms semi-erect at 45° angle to the
stem, bristly; and sparsely hirsute on youngest growth, tri-
chomes simple, fragile, glandular, erect to lax, to 0.5 mm long.
Leaves decussate, spreading to loosely ascending, 6 abruptly
reduced in size in the inflorescence; petioles 0–0.5 mm long;
blades lanceolate-elliptic, 5–22 mm long, 2–8 mm wide, mostly
15–20 mm long, 5 mm wide, thickly membranaceous, base
cuneate, apex acute, margin plane; surfaces green, bifid-
strigose mixed with sparse thin trichomes, abaxial midvein
and 3 sets of secondary veins visible on nearly all leaves,
prominent, brown. Racemes distinct, terminal, simple, brac-
teate, bracts elliptic, 5–10 mm long, margin plane; flowers
opposite, interpetiolar; pedicels 2–4 mm long; bracteoles much
reduced, obscured by abundant strigose indumentum. Floral
tube, including a well-developed subdescending spur
1.0–2.0 mm long, 6–8 mm long, 1.5 mm wide, neck constricted
at 2/3 length of floral tube, mouth flaring, 2–3 mm wide; outer
surface green, abundantly bifid-strigose, bristly; and sparsely
hirsute as on stems; inner surface white villous; sepals sub-
equal, 0.4 mm long, 0.9–1.2 mm wide, shallowly deltate, apex
acute to apiculate; epicalyx segments thickened lobes, 6 cy-
lindric, 0.5 mm long, slightly shorter than sepals, some with 1
or 2 setae at apex. Petals 6, pink or fuchsia, subequal,
2.0–2.2 mm long, 0.5–1.0 mm wide, oblong to narrowly ob-
ovate, 2 dorsals slightly smaller than ventrals. Stamens: 2 short
dorsalmost slightly deeper, other 9 inserted in nearly hori-
zontal line at 2/3 length of floral tube, all included, none
reaching sinuses of sepals; filaments densely white villous,
filament and anther color unknown; pollen oval-triangular,
tricolporate, non-syncolpate, pores not protruding, exine
shallowly parallel-rugulate, grain diam 15–17 mm. Ovary
villous; style sparsely villous, ultimately well-exserted, sur-
passing the sepals; ovules 5 or 6. Disc horizontal to slightly
deflexed, 0.5–0.7 mm long, 0.4–0.5 mm wide, sub-deltoid,
obtuse, thick, dark red. Seeds 1.5–1.7 mm long, 1.4–1.5 mm
wide, obovate, margin round. Figures 4C, 5D, 20A.
Phenology—Collected in flower and fruit in June and
August.
Distribution and Habitats—Colombia, Dept. Meta, in La
Macarena region; savanna and disturbed forest on rocky
outcrops; 400–480 m (Fig. 16).
Extinction Risk—The estimated risk if all IUCN guides
were to be followed is CR; EOO not available and AOO 5
8 km2. Two collection localities were analyzed, both within
protected areas.
Additional Specimens Examined—Colombia.—META: La Macarena,
Reserva de La Macarena, 20 km NO vı́a Los Conejos, on rocks, dominated
by Vellozia, 2°10’N,73°48’W [2.166, -73.8], 7 Aug 1988, Callejas & Marulanda
6915 (MO); primeros cerros del sur de La Macarena, frente al Rı́o
Guayabero, al pie de los acantilados, 10 Jun 1970, Echeverry E. 2089 (COL).
Cuphea saxatilis is newly recognized in the richly endemic
flora and fauna of the Sierra de la Macarena. The Sierra is an
isolated, north-south oriented mountain range to the east of
the Colombian Andes, and is part of the older Guiana Shield
(Gilliard 1942). The new species has a unique combination of
stems with a mix of bifid trichomes and simple, long, thin
trichomes; lanceolate-elliptic leaves; distinct racemes; and 5 or
6 ovules. Cuphea philombria is also in the Sierra de la Macarena in
localities to the east of C. saxatilis; it differs from C. saxatilis by
linear-lanceolate leaves, indistinct racemes, and 8–15 ovules.
18. CUPHEA STYGIALIS Lourteig, Bot. Mus. Leafl. (Harvard
University)16: 222, Fig. 37. 1954. TYPE: COLOMBIA.Vaupés:
Rı́o Apaporis, Raudal de Jirijirimo [0.133, -70.649], (below
 178 SYSTEMATIC BOTANY
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Fig. 20. A. Habit and flower of Cuphea saxatilis, holotype, Lopez et al. 6729 (COL). B. Habit of C. stygialis, holotype, Schultes & Cabrera 15330 (GH), and
flower (adapted from type illustration, Lourteig 1954).
mouth of Rı́o Kananarı́), general location 0°50 N, 70°40’W
[0.083, -70.666], 14 Feb 1952 [1932 in error], R. E. Schultes &
I. Cabrera 15330 (holotype: GH-00068455, image!; iso-
types: COL!; as images–GH, K, S, US!)
Subshrubs to 50 cm; stems one to several, erect or slightly
decumbent, from a woody rhizome with many short fibrous
roots, abundantly branched; stem indumentum exclusively
strigose, the trichomes almost all simple, 1-armed, closely
attached to the stem by an elongated base; distal internodes
less than half as long as to longer than the subtending leaves.
Leaves spreading to ascending, imbricate distally, gradually
reduced in size in the inflorescence; petioles 0–0.5 mm long;
blades linear to linear-lanceolate, 8–20 mm long, 1.5–3.0 mm
wide, appearing uni-nerved, coriaceous, base subacute, apex
obtuse, margin plane, thickened; surfaces green, glabrous or
abaxially paler and sparsely strigose, with prominent brown-
red midvein, secondary veins not visible. Racemes indistinct,
terminal, leafy, simple; flowers solitary, alternate, inter-
petiolar, partially obscured by leaves; pedicels 2.0–4.5 mm
long, persistent; bracteoles ovate to oblong, thick, ca. 0.5 mm
long. Floral tube, including a well-developed, horizontal to
descending spur 0.5–1.0 mm long, 4.5–5.5 mm long,
1.5–2.0 mm wide, neck at 2/3 length of floral tube remaining
wide, mouth widely flaring, ca. 3 mm wide; outer surface color
unknown, surface strigose, trichomes 1-armed as on stems,
[Volume 44
also sparsely hirsute or not, trichomes thin, reddish, to ca.
0.5 mm long; inner surface densely villous distal to stamen
insertion, lightly villous proximally; sepals subequal, ca. 0.7
long, 1.2 mm wide, deltate, apex apiculate; epicalyx segments
thickened lobes, shorter than the sepals. Petals 6, subequal,
white with purple midvein, subequal, 2 dorsals, 2.5–2.6 mm
long, 1.5–1.7 mm wide, oval to obovate, smaller than 4 ven-
trals, ventrals narrowly spatulate, ca. 3 mm long, 1.7 mm wide.
Stamens: 2 short dorsalmost deepest, other 9 inserted in a
horizontal line at ca. 2/3 length of floral tube, included, none
reaching sinuses of sepals; filaments white villous, except 3
ventrals glabrous; filament and anther color unknown; pollen
unknown. Ovary pilose; style pilose, included, extended to
apices of the sepals; ovules 4–6. Disc deflexed, 0.6 mm long,
0.7 mm wide, sub-globose, obtuse, thick, dark-colored. Seeds
1.2–1.3 mm long, 1.2 mm wide, sub-circular, apex obtuse,
margin thick, rounded. Figures 2 I, J, 20B.
Phenology—In flower and fruit in February.
Distribution and Habitats—Colombia, in Department
Vaupés; on quartzite base; along river margins; ca. 100–300 m
(Fig. 16).
Extinction Risk—The estimated risk if all IUCN guides
were to be followed is CR; EOO not available and AOO 5
8 km2. Two collection localities were analyzed, both in a
protected area.
 2019] GRAHAM: CUPHEA SECTION AMAZONIANA
Additional Specimen Examined—Colombia.—AMAZONAS: corregi-
miento de la Victoria, margen derecho del Rı́o Apaporis, sector Cueva de
Guacamaya en la entrada al raudal Jirijirimo, 0°50 05”S, 70°550 300W [0.084,
-70.925], 18 Mar 2009, Cárdenas, D. et al. 22006 (COL).
Cuphea stygialis is one of four closely similar Colombian
species in sect. Amazoniana. It shares with C. chiribiquetea, C.
kubeorum, and C. philombria: indistinct racemes, well-
developed spurs, and deflexed discs. The species differ
distinctly in stem indumentum. Two lack the typical bifid
trichome form present in the rest of the section. In Cuphea
stygialis, the strigose indumentum is composed of cystolithic,
1-armed trichomes attached to the stem by an elongated base
(Fig. 2 I, J). In Cuphea chiribiquetea (Fig. 2C, F, the indumentum,
uniquely in the section, consists of ovoid glands. Cuphea
kubeorum and C. philombria have typical bifid-strigose indu-
mentum, with some trichomes in C. kubeorum having semi-
erect arms, rather than being fully appressed to the plant
surface. In C. philombria, they are all closely appressed to the
surface. Cuphea stygialis and C. philombria are similar vegeta-
tively, but the latter has normal bifid trichomes and a high
number of 8–15 ovules vs. 5 or 6 ovules in C. stygialis.
19. CUPHEA SUCUMBIENSIS Lourteig, Bot. Mus. Leafl. (Harvard
Copyright (c) American Society for Plant Taxonomists. All rights reserved.
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University) 16: 225, Fig. 36, 1954. TYPE: COLOMBIA. Putu-
mayo: frontera Colombo-Ecuatoriana, selva higrófila del
Rı́o Sucumbı́os (San Miguel)[ca. 0.133, -75.850], entre los
afluentes Conejo y Hormiga, 300 m, 15 Dec 1940,
J. Cuatrecasas 11069a (holotype: F-1365302-image!; iso-
types: COL!; P, US-images!).
Subshrubs 15 to 35 cm; stems erect to decumbent, one to
several from a woody rhizome with fibrous roots, abundantly
branched distally; stem indumentum a mix of 2 types: bifid-
strigose, the trichomes thin, silky on young growth, closely
appressed to the stem, some with semi-erect arms; and
abundantly setose, or setose/hirsute, the trichomes bent and
slightly ascending, brown-red, slightly enlarged at base,
0.5–1.0 mm long, hirsute on young growth, the trichomes
thinner, more fragile; internodes less than half as long as
subtending leaves, gradually shorter distally. Leaves
spreading, crowded distally on the stem, gradually reduced
in size in the inflorescence; petioles 0–1 mm long; blades
linear-lanceolate, 3–18 mm long, 0.5–3.0 mm wide, appearing
uni-nerved, thickly membranaceous, base cuneate to sub-
acute, apex obtuse, margin incurved; surfaces adaxially
green, glabrous, or sparsely bifid-strigose, abaxially paler,
bifid-strigose, midvein prominent, brown, 1–3 pairs sec-
ondary veins rarely visible on larger, more proximal leaves
(Aulestia et al. 492, Ecuador). Racemes distinct, terminal,
simple, bracteate, bracts elliptic to oblong; flowers alternate
or alternate and opposite, interpetiolar or internodal, pedicels
1.5–2.5 mm long; bracteoles oval to oblong, obtuse, thick,
0.5 mm long. Floral tube, including a well-developed hori-
zontal to descending spur 0.5–1.0 mm long, 3.5–6 mm long,
1.0 mm wide, neck at 2/3–3/5 length of floral tube, not or
only slightly contracted, mouth oblique, widely flaring, ca.
2 mm wide; outer surface green with red veins, strigose as on
stems, sometimes also sparsely hirsute with long, thin tri-
chomes; inner surface villous distal to stamen insertion,
glabrous to pilose proximally; sepals subequal, 0.4–0.8 mm
long, 0.5–1.3 mm wide, the dorsalmost widest, all deltate,
apex apiculate; epicalyx segments thickened lobes,
0.3–0.7 mm long, 2 dorsal segments larger than others, all
segments shorter than sepals, setae 0–4. Petals 6, white, pink
179
or pale purple, subequal, 1.6–2.5 mm long, 0.7–1.1 mm wide,
2 dorsals slightly wider than 4 ventrals, all obovate to oblong.
Stamens: 2 short dorsalmost deepest, other 9 inserted in a
nearly horizontal line at 2/3–3/5 length of floral tube, lightly
to densely villous on proximal half of the filaments, 3 ventral
antesepalous stamens sometimes glabrous, all included, none
extending to sinuses of sepals; filaments white villous, fila-
ment and anther color unknown; pollen oval-triangular,
tricolporate, non-syncolpate, colpi short, pores not pro-
truding, exine finely scabrate, grain diam 18–19 mm. Ovary
and style sparsely pilose; style included; ovules 8–14. Disc
horizontal to typically deflexed, 0.5–0.6 mm long, 0.3–0.5 mm
wide, shape variably sub-globose to sub-deltoid or short
cylindric to sub-globose thick, dark red. Seeds 1.0–1.1 mm
long, 0.8–0.9 mm wide, sub-circular, base and apex rounded,
margin rounded. Figures 4D, 21A.
Phenology—Collected in flower and fruit throughout the
year.
Distribution and Habitats—Colombia and Ecuador; among
rocks along river margins, in wet riverside forest, in tropical,
pre-montane, primary forest; 50–1750 m, mostly 200–1000 m, at
1500–1770 m in montane forest, Napo, Ecuador (Fig. 16).
Extinction Risk—The estimated risk if all IUCN guides
were to be followed is LC; EOO 5 52,217.76 km2 and AOO 5
56 km2. Of 14 collection localities analyzed, only one occurs
in a protected area.
Additional Specimens Examined—Colombia.—CAUCA: La Espriella, a
orillas del Rı́o Caunapı́ [1.469,-78.640], 7–19 Jun 1986, Léon et al. 1425
~
(NY).—NARINO : Barbacoas [1.673, -78.138], Rı́o Telembı́, entre Barbacoas y
ca. 15 km rı́o arriba, 20 Nov 1986, Bernal & Hammel 1322 (COL, MO).—
PUTUMAYO: Mocoa, carr. Mocoa-Villa Garzón, km 5, Balneario Fin del
Mundo, bosques alrededor del Rı́o Dantayaco, 1°6’11.2” N, 76°370 17.80W
[1.103, -76.621], 28 Oct 2010, Bernal & Manrique 4625 (COL). Ecuador. —ESMERALDAS:
San Lorenzo, Parroquı́a Mataje, Reserva Etnica Awá, Centro Mataje,
1°08’N, 78°33’W [1.133, -78.55], 21 Sep 1992, Aulestia et al. 492 (MO, NY);
San Lorenzo, Rı́o Negro, a 2 km de El Placer, 0°55’N, 78°35’W [0.916,
-78.583], 20 Feb 1991, Palacios 6944 (MO). NAPO: Lago Agrio, Rı́o Oyacachi,
23 km from Baaza, 0°18’S, 77°48’W [-0.3, -77.8], 31 Oct 1976, Balslev &
Madsen 10506 (COL, MO, NY, P); Archidona, carr. Hollı́n-Loreto, Rı́o
Huataraco, dos horas a pié por bosque primario desde la aldea de Guagua
Sumaco, 0°43’S, 77°32’W [-0.716, -77.533], 23–30 Aug 1989, Cerón & Factos
7424 (MO); Orellana, Parque Nacional Sumaco, Proyecto Amoco, Bloque
18, lı́nea sı́smica AM-96–07, margen occidental del Rı́o Payamino, 0°24’S,
77°21’W [-0.4, -77.35], 2–4 Nov 1996, Dik 1698 (MO); Cerro Antisana,
banks of Rı́o Napo at Puerto Napo [-1.05, -77.783], 14 Jul 1960, Grubb et al.
182 (NY); Pastara a Borja (Virgilio Dávila), 15–26 1959, Harling 3849 (P);
Union of Rı́o Borja and Rı́o Quijos, west bank, 0°25’S, 77°49’W[-0.416,
-77.816],19 Sep 1980, Holm-Nielsen et al. 26173 (MO, P); Union of Rı́o Borja
and Rı́o Quijos, east bank, 19 Sep 1980, Holm-Nielsen et al. 26274 (MO, P);
Archidona, carr. Hollı́n-Loreto, km 17, cerca del Rı́o Hollı́n, 0°41’S,
77°41W [-0.683, -77.683], 14–22 Feb 1989, Hurtado & Shiguango 1662 (MO,
NY); Archidona, s slopes of Volcán Sumaco, 4 km N of Huahua Sumaco
community, primary forest near Rı́o Huataraco, 0°43’S, 77°32’W [-0.716,
-77.533], 13 Dec 1989, Neill et al. 9167 (MO); El Chaco, Codo Sinclair,
bosque primario en el valle del Rı́o Quijos, 0°8’S, 77°27’W [-0.133, -77.45],
16–20 Sep 1990, Palacios 5650 (MO).—SUCUMBIOS: Gonzalo Pizarro, Campo
Bermejo 6 Norte, 2 km al norte del Campo, 30 km al NW de Lago Agrio,
0°14’N, 77°13’W [0.233, -77.216], 30 Mar 1990, Cerón et al. 9377 (MO);
Gonzalo Pizarro, Cofán de Sinangue, bosque primario entre la comuni-
dad de Sinangüe y el Rı́o Candué, 0°08’, 77°27’W, [0.133, -77.45], 2 Nov
1992, Cerón 20763 (MO).
Cuphea sucumbiensis is the only species of the section oc-
curring in Colombia and Ecuador. At its northern limit in
Colombia, it is separated with difficulty from C. philombria,
which occurs farther north along the Cordillera de la Mac-
arena. Cuphea sucumbiensis and C. philombria can be distin-
guished from the rest of the section by the combination of:
stems both bifi-strigose and setose; high ovule numbers of
8–15; and a horizontal to deflexed disc. Cuphea sucumbiensis is
 180 SYSTEMATIC BOTANY
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Fig. 21. A. Habit and flower of Cuphea sucumbiensis, Aulestia 492 (MO). B. Habit and flower of C. trisperma, holotype, Steyermark et al. 105451 (MO).
shorter in stature than C. philombria (35 cm vs. 6 50 cm), with
shorter leaves 3–18 mm long, mostly less than 15 mm long vs.
(7–)15–30 mm long in C. philombria; and with distinct brac-
teate inflorescences vs. indistinct leafy ones in C. philombria.
In both species, setae on the epicalyx segments are in-
consistently present. Leaves in both species generally have a
glabrous adaxial leaf surface. In C. sucumbiensis, however, in
Aulestia 492 (MO, NY), Bernal & Hammel 1322 (MO), and
Palacios 6944 (MO), the leaves are bifid-strigose on both
surfaces. Pollen is Type 1 in both species, but differs in exine
sculpture and colpi shape and size in the grains examined
with SEM.
Petal color varies in Cuphea sucumbiensis. Of two collections
from Esmeraldas, Ecuador, one has white petals, the other
pink (rosadas). In Napo, the petals are described as white,
pink, or cream-violet, in Sucumbios as lilac. Petals in C.
philombria are purple, white, or bi-colored with the dorsal
petals purple and ventral petals pale purple. Two habit forms
are indicated in C. sucumbiensis, although this needs to be
verified by field observations. The form seen in the type col-
lection has leaves spreading outward along the stem and
distinct inflorescences; it occurs throughout the range of the
species. A second form is noted in three collections from Napo,
Ecuador, inside the range of the typical form in Napo. The
[Volume 44
leaves are ascending and more closely imbricate on the stem
and inflorescences are possibly indistinct.
Boiled leaves of Cuphea sucumbiensis are used to treat sore
throats in northern Ecuador, where the name in Cofrán is
Quni’cco Suvu (Cerón 20763, MO).
20. CUPHEA TRISPERMA S.A.Graham, sp. nov. TYPE: VENEZUELA.
Bolı́var: On igneous outcrops along a stream at km.108
south of El Dorado [6.715, -61.637], carretera El Dorado
toward Santa Elena de Uairen, 1200–1400 m, 19–22 Feb
1972, J. A. Steyermark, L. & R. Smith, G. C. K. & E. Dun-
sterville 105451 (holotype: MO-2680818!; isotype: NY!)
Diagnosis: Subshrubs 30 cm tall, all surfaces bifid-strigose;
stems several from near the base, slender, erect. Leaves sessile
or subsessile, decussate, linear, 3–11 mm long, 1.0–1.5 mm
wide, abundant, crowded, coriaceous, apices often deflexed.
Racemes indistinct, leafy. Floral tube 5.5–6.0 mm long; spur
poorly developed or none; dorsalmost sepal larger than the
others. Petals 6, pale lavender, 2 dorsals smaller than the 4
ventrals. Pollen tricolporate, colpi convergent at the poles in a
triangle enclosing a remnant of exine. Ovules 3. Disc strongly
reflexed over ovary wall, cylindric. Seeds 1.2–1.3 mm long.
Subshrubs to 30 cm; stems erect, dark brown, several closely
arising from a short woody rhizome with woody secondary
 2019] GRAHAM: CUPHEA SECTION AMAZONIANA 181

roots and some thin fibrous roots; branches slender, numerous Species Excluded From Sect. Amazoniana
distally; internodes longest proximally, becoming very short
C. fluviatilis S.A.Graham, Novon 19: 45–48, 2009. In the
distally; stem indumentum exclusively bifid-strigose, tri-
protologue, the relationships of this species were unsettled.
chomes moderately thick, arms sub-equal, semi-erect at ca. 45°
Vegetative habit, floral morphology, and riverine habitat
angle to stem. Leaves decussate, spreading, abundant,
suggested a position in sect. Amazoniana, whereas absence of
crowded and irregularly positioned distally; petioles
bifid trichomes, and pollen morphology indicated ties to
0–0.5 mm; blades linear, 3–11 mm long, 1.0–1.5 mm wide,
clades comprising species of sections Brachyandra and Euandra
appearing uni-nerved, coriaceous, apices often deflexed, base
(Inglis et al. in progress). Following a detailed study of all
obtuse, apex acute, margin incurved; surfaces bifid-strigose,
members of sect. Amazoniana, I believe that morphological
adaxially dark green, abaxially paler, drying yellow-brown,
similarities of C. fluviatilis to members of sect. Amazoniana are
midvein prominent, 1 or 2 pairs of secondary veins occa-
the result of parallel evolution, especially in vegetative char-
sionally visible abaxially on largest leaves. Racemes indistinct,
acters like the subshrub habit, small linear leaves, and massive
terminal, leafy; flowers sparse, alternate; pedicels 1–2 mm
fibrous root system. These characters likely evolved in parallel
long; bracteoles ovate, ca. 0.2 mm long. Floral tube,
in C. fluviatilis and in species of section Amazoniana as they
including a poorly developed spur 0.1 mm long or non-
adjusted to the same adaptive pressures generated by wide
spurred, 5.5–6 mm long, 1 mm wide, neck contracted at 2/
seasonal changes in riverine and savanna habitats, especially
3 length of floral tube, mouth widely flaring, 2.5 mm wide;
in changing water levels and varying strengths of river flows.
outer surface dark purple, abundantly bifid-strigose as on
In addition, the strongly striate pollen of C. fluviatilis is unlike
stems, trichome arms bristly, semi-erect; inner surface villous
pollen of any member of sect. Amazoniana, but is characteristic
above stamen insertion, sparsely villous or pilose proximally;
of numerous species in the clades comprising species of sec-
sepals subequal, ca. 1.2 mm long, 0.8 mm wide, dorsal sepal
tions Brachyandra and Euandra.
largest, all narrowly deltate, apex sharply acute; epicalyx
Cuphea odonellii Lourteig, Notas del Museo, Universidad
Copyright (c) American Society for Plant Taxonomists. All rights reserved.
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segments thickened lobes, 0.3–0.4 mm long, shorter than

sepals, strigose, dark-colored. Petals 6, lavender with dark
midvein, spatulate to obovate, unequal, 2 dorsals 1.5 mm
long, 0.7 mm wide, 4 ventrals larger, 2.0 mm long, 0.7 mm
wide. Stamens: 2 dorsalmost slightly deeper, other 9 inserted
in a nearly horizontal line at 2/3 length of floral tube, all
included, antesepalous stamens nearly reaching sinuses of
sepals, antepetalous ones shorter; filaments sparsely white
villous, filaments and anthers cream-colored; pollen oval-
triangular to triangular, tricolporate, syncolpate, colpi con-
vergent at poles in a triangle enclosing a remnant of exine,
pores not protruding, exine psilate, grain diam 17–20 mm.
Ovary and style slightly villous to glabrous; style ultimately
extended to apex of dorsalmost sepal; ovules 3, at distal end
of placenta. Disc reflexed over the ovary, cylindric, 0.8 mm
long, 0.1 mm wide, obtuse, dark red. Seeds 1.2–1.3 mm long,
0.9–1.0 mm wide, obovate, margin narrow, thinned. Figures
4G, 21B.
Phenology—Flowering and fruiting in February.
Distribution and Habitat—Venezuela, Bolı́var state; on
igneous outcrops along a stream; 1200–1400 m (Fig. 17).
Extinction risk—The estimated risk if all IUCN guides were
to be followed is CR; EOO not available and AOO 5 4 km2. The
species is known only from the type, which is in a protected
area.
Cuphea trisperma is a member of the group of Venezuelan
species in the section characterized by spur-less floral tubes,
cylindric discs reflexed and appressed to the ovary wall, and
syncolpate, psilate pollen. The species is distinguished
vegetatively from the others by the several slender stems
arising from the base of the plant and bearing multitudes of
tiny, linear leaves. The flowers are exclusively strigose,
have a large dorsal sepal, 6 pale purple petals, and 3 ovules. Two
other Venezuelan species in the section, C. bolivariensis and C.
distichophylla, also produce just 3 ovules. They are morphologi-
cally and geographically distant from C. trisperma. Cuphea dac-
tylophora (Steyermark et al. 105451, MO) occurs in the same
geographical area as C. trisperma, to the south of El Dorado in
eastern Venezuela near the Guyana border. Unlike C. trisperma, it
has large, wide, lanceolate-linear leaves, flowers 6–8 mm long,
and seven ovules.
Nacional de La Plata 19: 279–284. This species, widespread in
savannas of western South America in Bolivia, Peru, Colombia,
Venezuela and Brazil, was originally described in sect. Euandra,
subsect. Amazoniana Lourteig. On the basis of pollen morphology,
however, supported by presence of a curled deflexed disc, it
belongs among species of sect. Trispermum. The pollen type, in
which the walls of the grain are thickened between the pores
(interaperturate thickenings), and the curled disc, are unique
apomorphies of sect. Trispermum. (S. A. Graham and A. Graham
1971; Graham et al. 2006, Clade 4). Cuphea odonellii is here
transferred to sect. Trispermum.
Acknowledgments
I am indebted to several individuals who generously contributed their
expertise or technical assistance in the course of this study: A. Graham for
guidance on the past history of the Amazon and Guiana Shield regions; A.
Arbelaez for plant illustrations, P. Inglis for sharing results of phylogenetic
studies, B. Bassüner for map preparation and IUCN risk estimations; and
M. Stiffler for library acquisitions. I thank the curators of the herbaria CAY,
COL, GOET, MO, NY, OXF, P, and RB, who supplied collections for study,
including valuable types. I also found indispensable the online images of
specimens now available from several herbaria and JStor.org. I thank the
reviewers for their meticulous reading of the manuscript and for changes
suggested. The Missouri Botanical Garden and President Peter Wyse-
Jackson are gratefully acknowledged for generous use of the research
facilities of the Garden.
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logische Mededelingen 38: 71–90.
Appendix 1. Index to numbered collections examined. The numbers in
parentheses in the Index refer to the corresponding species in the numerical
list presented below. Type collections are starred following the collection
number.
1. C. beneradicata Lourteig; 2. C. blackii Lourteig; 3. C. bolivariensis
Lourteig; 4. C. cardonae Lourteig; 5. C. cataractarum Spruce ex Koehne; 6. C.
chiribiquetea Lourteig; 7. C. curiosa Lourteig; 8. C. dactylophora Koehne; 9. C.
distichophylla Lourteig; 10. C. insolita Lourteig; 11. C. kubeorum Lourteig; 12.
 2019] GRAHAM: CUPHEA SECTION AMAZONIANA
C. maigualidensis Lourteig; 13. C. philombria Lourteig; 14. C. pleiantha
Lourteig; 15. C. rhodocalyx Lourteig; 16. C. rigidula Benth.; 17. C. saxatilis S.
A. Graham, sp. nov.; 18. C. stygialis Lourteig; 19. C. sucumbiensis Lourteig;
20. C. trisperma S. A. Graham sp. nov.
Allen, P. 3235 (11); Aulestia, C. et al. 492 (19); Aymard, G. & Delgado, L.
6888 (7).
Balslev, H. & Madsen, E. 10506 (19); Barbosa, C. 7963 (6); Barclay, A. S.
et al. 532 (11); Barriga, H. G. et al. 15817 (10), 15862 (10), 15871(10); Bernal,
R. & Hammel, B. 1322 (19); Bernal, R. & Manrique, H. F. 4625 (19); Bernardi,
A. L. 1415* (3), 6802 (8); Berry, P. et al. 4904* (7), 6590 (8); Betancur, J. et al.
1898 (13), 18989 (1), 19237 (1); Black, G. A. 49-8243* (2), 49-8315 (2); 49-8823
(2); Boom, B. & Marin, E. 10546 (7).
Callejas, R. & Marulanda, O. 6915 (17); Cardona, F. 2972 *(4), 3013 (16);
Cárdenas, D. & González, M. F. 45513 (10); Cárdenas, D. et al. 4108 (11),
6514 (1), 6866 (11), 22006 (18), 42119 (13); Cardiel 1014 (11); Cerón, C. E.
20763 (19); Cerón, C. E. & Factos, M. 7424 (19); Cerón, C. E. et al. 9377 (19);
Clarke (H.) D. 5399 et al. (8); Cremers, G. 12126 (2), 15252 (2); Croat, T. B. &
Ferry, G. 102998 (2); Cuatrecasas, J. 7691 (10), 8846 (12), 11069a* (19);
Curran, H. M. 176 (5), 241 (5).
Delascio Ch., F. & Guánchez, F. 10717 (aff. 7), 10811 (7); Dik, A.1698 (19).
Echeverry E., R. 2089 (17); Egler, M. A. 1435 (2), 24591(2); Ek, R. C. et al.
1394 (2), 1509 (2), 1759 (2); Elcoro, S. 342 (aff. 7); Estrada, J. et al. 637 (11).
Copyright (c) American Society for Plant Taxonomists. All rights reserved.
Delivered by Ingenta to IP: 79.110.25.129 on: Sat, 23 Mar 2019 19:39:05
Fari~
nas, M., et al. 408 (15), 462 (5); Fernández, A. 5231 (aff. 7), 7400 (5);
Fernández, A. & Aymard, G. 4845 (7), 4870 (7); Fernández, A. et al. 6219 (9);
Fernández, F., J. L. et al. 5629 (13); Foldats, E. 2694 (8); Franco, P. et al. 3282
(6), 3760 (6), 3808 (11); Froés 25941 (2), 25952 (2).
Garcı́a, M. C. et al. 511 (13); González, M. F. & Cárdenas, D. 2619 (10);
González, M. F. et al. 1607 (1); Granville, J. J. de 496 (2), 4373 (2), 6023 (2),
11749 (2), 14242 (2); Grubb, P. J. et al. 182 (19).
Harling. G. 3849 (19); Henderson, A. 116 (13); Henkel, T. W. 4253 (8), 5694
(3); Hernández, M. et al. CHI-1407 (10); Hoffman, B. & Marco 2260* (10);
Holm-Nielsen et al. 26173 (19), 26274 (19); Holst, B. K. & Liesner 3069 (14);
Holst, B. K., Steyermark & Manara 2137 (8); Huber, O. 12735 (aff. 7), 12741*
(12), 13512 (9); Humbert, H. & Schultes, R. E. 27339 (11); Hurtado, F. &
Shiguango, J. 1662 (19).
183
Idrobo, J. M. 2431 (13); Idrobo, J. M. & Jaramillo, B. 2047 (13); Idrobo &
Schultes, R. E. 564 (13), 640* (1),1061(13),1062* (13),1288 (13); Idrobo, J. M. &
Zarucchi, J. 9.431 (1), 9.436 (1); Irwin, H. S. & Westra, L. T. 47244 (2).
Léon et al. 1425 (19); Liesner, R. 17524 (5), 17617 (15), 18510 (15), 18650
(aff. 7), 24871 (15), 25502 (15); López, R. & J. Betancur B. 3064 (1), 3114 (1);
López, R. & D. Giraldo-C. 651 (1); López, R. et al. 6729* (17).
Madri~ nán, S. et al. 1121 (11); Maguire, B. & Maguire, B., Jr. 29201* (15);
Maguire, B. & Maguire, C. K. 35102 (14), 35126* (14), 35246 (14), 35376 (14);
Maguire, B. & Politi, L. 27646 (9), 27764 (9), 27765* (9), 28217 (9); Maguire
et al. 29497 (5), 30348 (5),46715 (8), 46815 (8), 60547 (8); Martı́nez, X. 522 (11);
Mori, S. A. et al. 25376 (2).
Neill, D. et al. 9167 (19).
Palacios, P. et al. 2400 (11); Palacios, W. 5650 (19); 6944 (19); Phillipson,
W. R. et al.1609 (13); Pires, J. M. 47534 (2); Pires, J. M. & Westra, L. 48869 (2);
Pruski, J. & J. Steyermark 1369 (8), 1382 (8).
Rabelo 1546 (2).
Sánchez, F. FAS 145 (1); Sanoja, E. 2619 (7); Sastre, C. 4758 (2); Sastre, C. &
Raichel D., H. 5115 (11); Sastre, C. et al. 8541 (8), 9180 (1), 9225 (1), 9230 (1);
Schomburgk, Robert 155 S* (16), 1556* (8); Schultes, R. E. & Cabrera, I.
14267 (11), 14389 (11), 15330* (18), 18306* (11), 19177 (11), 19913 (11), 22600
(11); 22605 (11), 22657 (11); Silland, P. & J. Sutra 932 (2); Silva, N. T. &
Braz~
ao, U. 60641 (7); Smith, S. G. & Idrobo, J. M. 1480* (13); Soejarto, D. &
Lockwood, T. E. 2417 (11); Spruce, R. 3261* (5); Stergios, B. 11877 (7), 11931
(7); Stergios, B. & Velazco, J. 14480 (4), 14573 (4); Steyermark, J. A. 92831(7),
104113 (8), 105451* (20); Steyermark & Gibson 95670 (4); Steyermark &
Holst, B. 130645 (15); Steyermark & Nilson, S. 10 (8), 634 (8), 756 (8);
Steyermark et al. 97835* (7), 104483 (8), 107107 (4), 123626 (14), 125678 (5).
Thomas, J. et al. 1467 (13).
Velayos, M. et al. 6329* (6); Villiers, J. F. & Sarthou, C. 6023 (2); Vivas et al.
1563 (8).
Woronow, G. & Juzepczuk, S. 6454 (13); Wurdack, J. J. & Adderly, L. S.
43590 (7); Wurdack, J. J. & Monachino, J. V. 41073 (7).
Zarucchi, J. L. 2011 (11).