Two new species of Clusiella (Clusiaceae) with a
synopsis of the genus
Barry E. Hammel
Missouri Botanical Garden
P.O. Box 299
St. Louis, Missouri 63166-0299
Novon 9: 349-359. 1999. Published on 15 September 1999. ©
Missouri Botanical Garden 1999.
Abstract. Two new species of Clusiella, one from Costa Rica and Panama (C. isthmensis) and one from Brazil (C.
impressinervis), are described, one old species from Colombia
(C. cordifolia Cuatrecasas) is newly placed in synonymy, and
all eight species in the genus are brought together in a key. A
description of the genus and brief descriptions are given, specimens
are cited and distributions are mapped for all species. In view of
its stipuliform structures, contorted petals, psilate pollen exine,
baccate fruits, exarillate seeds, and embryos with well-developed
cotyledons, Clusiella's presumed close relationship to
Clusia is challenged.
When Planchon and Triana (1860) published Clusiella (based
on C. elegans) the genus was considered monotypic and known
only from pistillate flowers. It remained that way for nearly 100
years. However, Planchon and Triana appended to their treatment of
Clusiella a full description of staminate material of a
species now known as C. axillaris (Engler) Cuatrecasas.
Planchon and Triana implied that the collections on which that
description was based (Spruce 2159 and 2854)
might belong to a second species of Clusiella, but they gave
it no species name, and presented the description under the heading
"Asthotheca, Miers" with the prophetic injuction to see it
"without prejudice as to placement among the other Clusioids"
(Planchon & Triana, 1860, ser. 4, vol. 14: 254). Although all
genera, except this one, are specifically designated as such in their
large publication Planchon and Triana did effectively publish the
genus. The spelling "Asthotheca," however, must certainly have
been a typographical error for Astrotheca; on p. 240 of the
same publication they refer to the "genre manuscrit
Astrotheca, Miers." Bentham and Hooker (1862) repeated that
error when they reduced "Asthotheca" to a section of the genus
Clusia. Meanwhile, Engler (1888) published Clusia
axillaris Engler based on the same two Spruce collections. Vesque
(1892, 1893), using the correct spelling, and aware that "the
discovery of its female flowers and of the male flowers of
Clusiella might one day permit the combination of these two
genera" (Vesque 1892: 16), assigned two species to Astrotheca,
A. cuspidata Vesque and A. sulphurea (Poeppig) Vesque. The former was clearly a superfluous epithet
because Vesque cited the earlier Clusia axillaris Engler and
both Spruce collections. Astrotheca sulfurea is not a
Clusiella (Cuatrecasas, 1950). Cuatrecasas (1949, 1950)
finally brought Astrotheca (as to A. cuspidata Vesque) to rest in synonymy under Clusiella and described
five new species.
Key and Descriptions
Clusiella Planchon & Triana, Ann. Sci. Nat. Bot.
Sér. 4. 14: 253. 1860. TYPE: Clusiella elegans Planchon
Astrotheca Miers ex Planchon & Triana, Ann. Sci. Nat.
Bot. Sér. 4. 14: 254. 1860 (as "Asthotheca", a
typographical error). No species indicated, but based on
Spruce 2159 & 2854, from Brazil.
LECTOTYPE, here designated: Astrotheca cuspidata Vesque
(=Clusiella axillaris (Engler) Cuatrecasas).
Clusia subg. Criuva sect. Astrotheca (Miers
ex Planchon & Triana) Bentham & Hooker, Gen. Pl. 1: 170. 1862
Dioecious, epiphytic shrubs; resin (latex) clear; internodes
lenticellate, bearing several pairs of caducous or rarely persistent
bud scales usually clustered just above the node (one pair more
distant). Minute, interpetiolar stipuliform structures present, these
broadly triangular, scarious, drying dark, caducous. Petioles short,
somewhat channeled by virtue of the narrowly decurrent leaf blade.
Leaves opposite, the blades elliptic, sometimes cordate, ±
coriaceous, occasionally glaucous in dried material; major lateral
veins often very indistinct until dried, merging to a submarginal
vein, the intersecondaries often almost as prominent as the
secondaries; translucid (sometimes only with intense light) resin
dots and dashes present, occasionally drying dark and surficially
visible but then usually only on the lower surface, resin canals
sometimes also visible on thin, new leaves. Inflorescences terminal
or evicted by the growth of an axillary shoot and then appearing
axillary, at only one side of the node, short, 1-several-flowered,
± dichotomously branching cymes with numerous small, decussate
bracts dispersed on the rachis. Flowers with 5 small imbricate sepals
and 5 larger contorted petals, the petals white (maroon or wine-red),
often with yellow or pink markings at the base within. Staminate
flowers with the filaments connate for most of their length into a
narrow column but free for a short distance apically and forming a
± globose capitulum, the column surrounded by a collar of
clavate, resiniferous staminodia or resin globules; anthers ±
globose and only slightly wider than the filaments (Figs. 1, 2);
pollen spheroidal-oblate, tricolporate, psilate (Fig. 3). Pistillate
flowers with the ovary surrounded by a resiniferous collar of
staminodes that grade acropetally from resin glands to more clearly
stamenlike structures; stigmas sessile, 5 to ca. 20, circular and
distinct (when few) or packed into a disk and indistinctly triangular
to rectangular (when many); ovules sometimes with a long funicle.
Fruit a ± globose, many-seeded berry with about as many locules
as stigmas; seeds elongate-ovoid, small (ca. 1 X 0.5 mm) with a
clear, gelatinous outer integument attached only at the hilum, the
exotegmen pitted (Figs. 4, 5), with sinuous anticlinal cell walls,
lignified; endosperm white, embryo with cotyledons ca. 2/5 the length
of the seed.
Figures 1--5. Clusiella (1--3, C. elegans
Planchon & Triana; Hammel & Kress 11287; 4, 5, C. isthmensis Hammel, Hammel & Trainer 12721). ---1. Staminate bud with petals removed, showing undehisced anthers and staminal column subtended by whorl of resinous staminodia. ---2. Mature staminate flower. ---3. Pollen grain. ---4. Seed with integument laid back to show pitted exotegmen. ---5. Seed with the clear integument partially removed. Scale bar: for Figs. 1, 4 & 5 = 3 mm; for Fig. 2 = 6 mm; for Fig. 3 = 10 µm.
Distribution. Clusiella is restricted to the
Neotropics, where species occur in Costa Rica, Panama, Colombia,
Venezuela, Ecuador, northern Peru, and northern Brazil (Fig. 6).
Figure 6. Distribution of Clusiella. (Occurrence of
C. axilaris with __> pertains to off-map
collection in Pará state of Brazil).
Key to the species of Clusiella
- 1a. Midrib impressed above; resin dots conspicuous (with hand lens) and prominent on both leaf surfaces....................C. impressinervis sp. nov.
- 1b. Midrib prominent above; resin dots obscure and visible mostly only on lower leaf surface.
- 2a. Leaves amplexicaul; petioles 2 mm or shorter.................................................C. amplexicaulis
- 2b. Leaves not amplexicaul; petioles 2 mm or longer.
- 3a. Petioles 8--10 mm; lateral veins mostly ca. 1 mm
- 3b. Petioles mostly 5(--8) mm or less; lateral veins mostly
ca. 2 mm or more distant.
- 4a. Fruits ± narrowly ovoid (ca. 5 mm diam. or less) with 5--7 distinct stigmas; leaves small, 4.5--5.5(--10) x 11.5(--4)
- 4b. Fruits globose, larger and with 10 or more connivent stigmas; leaves larger.
- 5a. Flowers wine red [buds ca. 5 mm diam.; fruits ca.
1.5--2 cm diam.; twigs tan or white; leaf acumen ca. 2
cm; Valle del Cauca, Colombia & Esmeraldas,
- 5b. Flowers white.
- 6a. Leaves drying dark brownish gray, never
glaucous; twigs black; flower buds ca. 10--15 mm
diam.; fruits, ca. 4 cm diam.; Amazonia (Brazil,
- 6b. Leaves drying lighter, usually glaucous; twigs
mostly brown to white; flower buds less than 10 mm
diam.; fruits ca. 10 mm diam.
- 7a. Fruit wall thick (ca. 0.5--1 mm) and
appearing woody in dried material; leaves strongly
coriaceous, drying glaucous above and below, the
apex usually abruptly acuminate, the petiole ca. 5
- 7b. Fruit wall thin (ca. 0.2 mm), leathery; leaves membranaceous, rarely drying glaucous below, the apex gradually acuminate, the petiole 2--3 mm; Costa Rica and Panama..................C. isthmensis sp. nov.
Clusiella albiflora Cuatrecasas, Ann. Inst. Biol. Mex. 20: 111. 1949. TYPE: Colombia. Valle del Cauca: Río Yurumanguí, Peña de Candelario, Costa Pacífica, 10 m, Feb. 1944 (stam. fl), Cuatrecasas 16123 (holotype, F; isotype, US).
Bud scales sometimes persistent, to ca. 4 cm long, their scars up
to 3.5 cm above the node. Petioles 8--10 mm; leaf blades elliptic,
8--12 x 4--6 cm, the base acute, the apex acuminate to a 1.5--2.5 cm
acumen, drying glaucous above, dark tan above and below, lateral
veins ca. 1--2(--4) mm apart, merging to a distinct submarginal vein
ca. 1--1.5 mm from the margin, the intersecondaries prominent and
easily confused with the secondaries. Inflorescences branched several
times from above a peduncle ca. 5 mm long, 1-several flowered; flower
buds ca. 8 mm diam.; petals ca. 10 x 5 mm. Staminate flowers with the staminal column, including
capitulum, very short, ca. 3 mm long. Fruits globose, ca. 10 mm
diam., the stigmatic disk with ca. 6 closely packed, ±
triangular stigmas, fruit wall rather thick, ca. 0.5 mm.
Distribution. Colombia, from near sea level.
This species is still known only from the type and one paratype.
It has longer petioles and closer lateral veins than the other
species as well as a very long acuminate leaf tip and a very short
staminal column. In leaf shape and color, though not in size, it
appears most similar to C. elegans.
Additional specimen examined. COLOMBIA. Cauca:
Río Micay, Guayabal, Costa Pacífica, 520 m, Feb. 1943
(fr), Cuatrecasas 14132 (F, US).
Clusiella amplexicaulis Cuatrecasas, Rev. Acad. Colomb. Cien. Exact. 8: 61. 1950. TYPE: Colombia. Valle del Cauca: Río Cajambre, Barco, 5--80 m, Apr. 1944 (stam. fl), Cuatrecasas 17188 (holotype, F).
Bud scales up to 7 mm above the node. Petioles indistinct, ca.
1--2 mm; leaf blades ovate, 13--15 x 7--9 cm, the base cordate and
± amplexicaul, the apex abruptly acujminate, drying slightly
glaucous above, light tan on both surfaces; lateral veins (3--)5--10
mm apart, merging to a distinct submarginal vein ca. 2 mm from
margin, the intersecondaries faint but occasionally reaching to the
submarginal vein; translucid resin dashes faint. Inflorescence with
ca. 1--3 flowers (in bud). Fruits unknown.
Distribution. Colombia, from near sea level.
This species is still known only from the type collection,
material with immature staminate flowers only. The large, amplexicaul
leaves with short petioles and distant primary lateral veins
distinguish it from all other species. In leaf color and texture
C. amplexicaulis is similar to C. elegans.
Clusiella axillaris (Engler) Cuatrecasas, Rev. Acad. Colomb. Cien. Exact. 8: 61. 1950. Clusia axillaris Engler in Mart., Fl. Brasil. 12: 413. 1888. Astrotheca cuspidata Vesque, Epharm. Genitalia Foliaque Clusiearum et Moronobearum. 3: 16, Tab. 80 & 81. 1892. LECTOTYPE here designated: Brazil. Amazonas: São Gabriel de Cachoeira: Rio Negro, Jan.--Aug. 1852 (stam. fl), Spruce 2159 (syntypes, GH, GOET;
photo A, MO).
Bud scales sometimes persistent, 5--20 mm long, the scars up to 20
mm above the node. Petioles 49 mm, deeply channeled; leaf blades
elliptic, 10--13(--14) x 4--6(--7) cm, the base acute to rounded, the
apex acute to acuminate to a 5--20 mm acumen, drying somewhat shiny,
dark olive gray above, tan below, lateral veins 2.5--5 mm apart,
merging to an indistinct submarginal vein 1 mm or less from the
margin, intersecondaries ca. 1--2 mm apart; translucid resin dots
visible only with intense light or on thin, new leaves. Inflorescence
usually 1-flowered; flower buds to 1.5 cm diam.; petals to 2 x 1.3
cm. Staminate flowers with the staminal column, including capitulum,
ca. 5--8 mm tall. Fruit to ca. 4 cm diam., the stigmatic disk with
ca. 20 closely packed rectangular stigmas, fruit wall rather thick,
ca. 0.5 mm.
Distribution. Amazonian Brazil, Colombia and Venezuela,
from ca. 100 to 750 m.
This species is easily recognized by virtue of its black twigs,
dark drying leaves, large flowers and large fruits. It is distinctive
also for its Amazonian distribution. It is probably related to C.
impressinervis and C. pendula.
Additional specimens examined. BRAZIL. Acre: E slope
of Serra da Moa, Apr. 1971 (stam. fl), Prance et
al. 12584 (MO); Rio Negro, Rio Cauaburí, 750 m,
Nov. 1965 (stam. fl), Maguire et al. 60422 (F,
GH). Amazonas: Maraa, Rio Japurá, Dec. 1982 (pist. fl),
Plowman et al. 12252 (F); São Paulo de
Olivença, Apr. 1944 (fl, fr), Ducke 1626 (A, F,
MO); basin of Rio Solimões, Dec. 1936 (stam. fl),
Krukoff 8626 (A, F, MO); basin of creek Belem, (stam.
fl), Krukoff 8933 (A, F, MO), Rio Solimões,
Igarapé Jandiaruba, Jan. 1969 (fl), Fróes
23906 (GH). Pará: Road BR 22, Camparema to
Maranhão, km 66, vicinity of Piriroro, Nov. 1965 (pist. fl),
Prance & Pennington 1971 (GH). COLOMBIA.
Amazonas: Leticia, Tarapacá, Parque Nacional Natual
Amacayacu, 200 m, Nov. 1991, Pipoly et al. 16055
(MO), 100 m, July 1992, Rudas et al. 5471 (MO).
Guainía: San Felipe Neri, on Río Negro, across
from San Carlos de Río Negro, Venezuela, 120 m, May 1979
(stam. fl), Clark & Clark 7161 (MO).
Vaupés: Río Apaporis, Sorajama (above mouth of
Río Kananarí) and vicinity, 275 m, (pist. fl),
Schultes & Cabrera 16089 (GH); Mitrú &
vicinity, along Río Vaupes about 6 km below Mirú at
Tukanaré, Aug. 1976 (fr), Zarucchi et al.
1883 (GH). PERU. Loreto: Maynas, Mishana, Río
Nanay between Iquitos and Santa María de Nanay, 150 m, Mar.
1979 (stam. fl), Gentry & Aronson
25305 (F, MO), 130 m, Nov. 1978 Diaz et al.
1984 (MO); Alipahuayo (Estación IIAP), Nov. 1984 (pist.
fl), Vásquez et al. 5887 (F, MO);
Requena, Jenaro Herrera, Río Ucayali, May 1982 (fr),
Encarnación 26132 (MO); Sapuena, Jenaro Herrera,
170 m, Nov. 1987 (stam. fl), Vásquez et al.
10051 (F, MO). VENEZUELA. Territorio Federal Amazonas:
Dept. Atures, Río Autana, 90--110 m, Nov. 1984 (fr),
Guanchez 3249 (MO); Dept. Río Negro, middle part
of Río Baría, 80 m, July 1984 (fr), Davidse
27549 (MO); 3 km NE of San Carlos de Río Negro, 120 m,
Apr. 1979 (stam. fl), Liesner
6777 (MO); Ríos Pacimoni-Yasya, Casiquiare,
100--140 m, Jan. 1954 (stam. fl), Maguire, Wurdack
& Bunting 37439 (GH); Río Negro,
Picora de Cocuí, Dec. 1947 (stam.? fl), Schultes
& López 9436 (GH).
Clusiella elegans Planchon & Triana, Ann. Sci. Nat.
Bot. 13 (ser 4): 254. 1860. TYPE: Colombia. Chocó: 2000 m,
1866 (fr), Triana s.n. (holotype, P; isotypes,
F-fragment, G [photo F, MO], W [photo
Bud scales inconspicuous, their scars clustered at or just above
the nodes. Petioles 1--2(--3) mm; leaf blades elliptic, 4--4.5(--10)
x 1--1.5(--4.5) cm, the base acute (rarely rounded), the apex
acuminate to a 1--1.5(--2) cm acumen, drying glaucous and olive green
above, shiny and tan below, lateral veins 2.5--5 mm apart, merging to
an indistinct submarginal vein ca. 1 mm from the margin,
intersecondaries faint, ca. 1--2 mm apart; resin dots and dashes
translucid or sometimes dark. Inflorescences unbranched or often with
up to 6 branches; flower buds 2--4(--6) mm diam.; petals ca. 4 x 3
mm. Staminate flowers poorly known, the staminal column, including
capitulum, ca. 2(6?) mm tall. Fruits ovoid, 4--8(--12) x 3--5(--10)
mm, somewhat apiculate; stigmas 5--7, ± distinct, circular.
Distribution. Panama and Colombia from near sea level to
Originally described from pistillate material, this species is
still known from very few staminate collections. Only three such
collections are known: one from the disjunct population in Panama
(Hammel & Kress 11287) and two from
Colombia (Garcia-Barriga 11130 and Luteyn
et al. 10682). In general, the plants are much more
delicate than the other species, but C. elegans is
particularly distinguished by its small, ovoid fruits with 5--7, more
or less distinct stigmas. As discussed below, the large-leaved forms
of C. elegans look much like C. isthmensis and also
like the small-leaved forms of C. macropetala. Indeed, the
glaucous upper surface of the leaves, as well as their color and
venation, suggest that the three species are closely related.
Betancur 802 (Cited below) has leaves that are large
for the species and somewhat glaucous below as in C.
macropetala. The fruits of that collection are also large for the
species, but in concordance with placement in C. elegans they
have a thin and leathery, rather than thick and bony fruit wall, and
the stigmas are few and ± distinct.
Additional specimens examined. PANAMA. Coclé:
between Llano Grande and Coclecito, 600 m, July 1979 (fr),
Antonio 1396 (MO), Dec. 1979 (fr), Antonio
3050 (MO); Mar. 1982 (stam. fl), Hammel &
Kress 11287 (DUKE). COLOMBIA. Antioquia: Amalfi
a Rumazón, 1550 m, Sep. 1988 (fr), Betancur et
al. 802 (MO). Chocó: between La Oveja and
Quibdó, Apr. 1931 (fr), Archer 1753 (US); 15 km
E of Quibdó, 75 m, Apr. 1931 (fr), Archer 2214
(US); Quibdó, Carretera Yute--Lloró, 70 m, Sep. 1976
(fr), Forero & Jaramillo 2721 (MO),
80 m, June 1983 (fr), Forero et al. 9700 (MO),
100 m, Aug. 1976 (fr), Gentry & Fallen
17817 (pro parte, mixed collection with C. macropetala,
MO), Nov. 1988 (fr), Ramirez 1 (MO); highway
Bolivar--Quibdó, km 52--70, 500--600 m, July 1944
Garcia-Barriga 11130 (US); km 175--176, 456 m, Mar.
1984 (fr), Juncosa 2578 (MO); 37--40 km W of El Carmen,
671--1360 m, May 1984 (stam. fl), Luteyn et al.
10682 (MO); road to Cabí SW of Tutuendo, 100 m, 19 Jan.
1979 (fr), Gentry & Renteria 24450 (MO);
Río Serrano, 4--6 km arriba de Guayabal, 50 m, Apr. 1975 (fr),
Forero et al. 1362 (MO). Valle del Cauca:
Buenaventura, Bajo Calima, 50 m, July 1988 (fr), Croat
69479, 69515 (MO), Dec. 1981 (fr), Gentry
35589 (MO), ca. 100 m, July 1984 (fr), Gentry
& Monsalve 48413 (MO), Feb. 1989 (fl),
Gentry et al. 65553 (MO), June 1987 (fr),
Monsalve 1509 (MO), Oct. 1987 (fr), Monsalve
1982 (MO), 50--100 m, Feb 1984 (fr), Juncosa 2125 (MO),
Buenaventura to Cali, 100 m, June 1944 (fr), Killip
& Cuatrecasas 38926 (F, GH, US); Punta
Arenas, Buenaventura Bay, near sea level, June 1944 (fr),
Killip & Cuatrecasas 38647 (US);
Quebrada de Aguadulce, 0--10 m, Feb 1946 (pist.? fl),
Cuatrecasas 19996 (F, US).
Clusiella impressinervis Hammel, sp. nov. TYPE: Brazil. Amazonas: Tonantins, Mar. 1944 (stam. fl), Ducke 1625 (holotype, US; isotypes, A, F). Figure 7.
Figure 7. Clusiella impressinervis Hammel; Ducke 1625. ---A. Habit. ---B. Bud scales. ---C. Base of leaf,
abaxial view. ---D. Base of leaf, adaxial view.
A speciebus aliis Clusiellae in costa supra impressa et punctis resiniferis utrinque foliorum manifestis, differt. Clusiella axillaris fortasse affinis, sed foliis et floribus minoribus.
Bud scales often persistent, several sets borne just above the
node and one pair borne further above. Petioles 2--3 mm, deeply
channeled; leaf blades elliptic to obovate, widest at or above the
middle, (7--)8--9 x (2.5--)3--3.5 cm, the base acute, the apex
abruptly acuminate to a 1.5-cm acumen, drying shiny, light green both
surfaces; lateral veins mostly 1.5(--3) mm apart, merging to a very
indistinct submarginal vein mostly hidden along the recurved margin;
resin dots very conspicuous, both surfaces. Inflorescences apparently
1-flowered; flower buds 7--8 mm diam.; petals ca. 10 mm long x 7 mm
wide, white. Staminate flowers with the staminal column, including
capitulum, ca. 4 mm tall, filaments ± free apically but
congested. Pistillate flowers and fruits unknown.
Distribution. Brazil and Venezuela, from ca. 50 to 1220 m
Since Clusiella impressinervis is so poorly known it is
difficult to relate to the other species. However, by virtue of its
non-glaucous leaves, its deeply channeled petioles, its persistent
bud scales, and its Amazonian distribution, the species may be
related to C. axillaris. Leaves of that species also
occasionally have resin dots visible on the upper surface. The new
species differs from C. axillaris by its smaller leaves and
flowers and from all species not only by its impressed midrib but
also because its leaves dry light green on both surfaces. More
material, especially fruiting, is needed to clarify its
Additional specimens examined. VENEZUELA.
Bolívar: Cerro Venamo, 1220 m, Jan. 1964 (fl. bud),
Steyermark et al. 92790 (F, US).
Steyermark et al. 92790 will key to C.
impressinervis by virtue of its impressed midrib and resin dots
on both leaf surfaces. It also has deeply grooved petioles and
somewhat persistent bud scales. However, it differs from the type by
its ovate leaves, ca. 6--7 x 3 cm, which are widest below the middle,
have a rounded base, and dry dull on both surfaces, the upper olive
gray and the lower reddish tan, and by its shorter petioles only ca.
1--2 mm long. It is also found at a much higher elevation and is the
easternmost collection known of the genus. By virtue of sharing
unusual vegetative characters with C. impressinervis this
specimen is tentatively included here. The description of the species
is based on the type collection alone.
Clusiella isthmensis Hammel, sp. nov. TYPE: Costa Rica. Limón: Cerro Coronel, 5 m, Sep. 1986 (fl, fr), Davidse & Herrera 31421 (holotype,
CR; isotypes, F, MO) Figure 8.
Figure 8. Clusiella isthmensis Hammel; Davidse
& Herrera 31421. ---A. Habit. ---B. Pist. flower.
---C. Base of leaf, abaxial view. ---D. Base of leaf, adaxial view.
Clusiella macropetala Cuatrecasas affinis sed
in exocarpio coriaceo (non osseo) folia membranaciore, et petiolio
Bud scales mostly deciduous, their scars congested at or near the
nodes, rarely persistent (Kennedy et al. 2394)
with a pair up to 3.5 cm above the node. Petiole 2--3 mm, somewhat
channeled; leaf blades elliptic-lanceolate, widest below the middle
(6--)8--14 x (2--)3--4.5 cm, the base acute to rounded, the apex
gradually acuminate to a 1--2 cm-acumen, in dried material brown to
glaucous and gray above, tan and sometimes glaucous below, lateral
veins 2--3(--5) mm apart, merging to an indistinct submarginal vein
mostly less than 1 mm from and hidden along the recurved margin;
translucid resin dots and dashes sometimes also surficially visible
on lower surface. Inflorescences mostly 1(--3)-flowered; petals ca.
10 x 6 mm, white. Staminate flowers with the staminal column,
including capitulum, ca. 6 mm tall, the filaments free for a short
distance apically. Pistillate flowers with apparently 10 stigmas but
these forming a disk and difficult to distinguish; ovary 10-celled.
Fruits ca. 1 cm diam. at maturity, translucent white, soft.
Distribution. Costa Rica and Panama, from near sea level to
Although this species was previously identified as Clusiella
elegans (D'Arcy, 1980; Hammel, 1986) it is more closely related
to C. macropetala. As here understood, C. elegans is
generally a much more delicate plant with smaller leaves, flowers,
and fruits than both C. isthmensis and C. macropetala.
Most importantly, the ca. 10 stigmas of the latter two species are
± fused into a stigmatic cap, whereas those of C. elegans
are fewer (ca. 5--7) and distinct. In addition to the characters
mentioned in the diagnosis, C. isthmensis can be distinguished
from C. macropetala by its generally smaller, more gradually
acuminate leaf blades and by its shorter, thinner petioles.
Nevertheless, in leaf size, shape, and color, both C. macropetala and C. elegans appear to overlap somewhat with C. isthmensis. The only other species that might be confused with C. isthmensis is C. pendula Cuatrecasas However, the
leaves of this species dry darker with more prominent lateral veins,
are shiny rather than glaucous, and the flowers are red or maroon
rather than white. One collection, Foster et al. 14661, from lowland Bocas del Toro province, Panama, is problematic. The leaf size, shape, and coloration, as well as flower color and fruit size, all correspond with C. isthmensis, but the fruits have fewer and nearly separate stigmas, more like those of C. elegans.
Original field notes for the type of this species indicate 11
duplicates. These were distributed as C. elegans, and their
whereabouts, except as indicated above, are uncertain.
Additional specimens examined. COSTA RICA. Heredia:
La Selva Biological Station just E of the juncture of the
Sarapiquí and Puerto Viejo Rivers, ca. 100 m, (all collections
from La Selva are pistillate), Apr. 1981, Folsom 9673
(DUKE), Apr. 1981, Folsom 9893 (DUKE, F, MO), Mar.
1980, Hammel 8252 (DUKE, F, MO), May 1980, Hammel 8625 (DUKE), June 1980, Hammel 8934 (DUKE), July 1980, Hammel 9286 (DUKE), June 1982, Hammel & Trainer 12721 (DUKE), Aug. 1981 Damon Smith 109 (DUKE, MO), Oct. 1981, Damon Smith 457 (DUKE, F, MO); Parque Nac. Braulio Carrillo, between Río Peje and Río Sardinal, Atlantic slope of Volcán Barva, 1200--1400 m, Nov. 1985 (fr), Grayum & Herrera 7867 (CR, MO); Estación el Ceibo, 520 m, Dec. 1992 (fr), Boyle 1448 (CR); along W fork of
Río Sardinal, Atlantic slope of Volcán Barva, 670 m,
Dec. 1987 (fr), Grayum 8501 (F, INB, MO). Limón: ca. 2 km S of Río Colorado along new road and ca. 1 km E, 5 m, Mar. 1987 (fl, fr), Stevens et al. 25074 (MO); Cantón de Talamanca, entre Sukút y Amubri, 700--900 m, July 1989 (stam. fl), Hammel et al. 17625 (CR); Alto Urén, 1190 m, July 1989 (stam. fl), Herrera 3368 (INB, MO); Alto Lari, 1300 m, Mar. 1992 (stam. fl), Herrera 5190 (INB, MO); 1500 m, Mar. 1992 (stam. fl) Herrera 5422 (INB, MO). PANAMA. Coclé:
Continental divide above El Copé, 750 m, Feb 1982 (stam. fl),
Knapp & Dressler 3410 (MO). Darién: Río Tuquesa headwaters, ca. 2 km from continental divide, ca 400 m, Aug. 1974 (fl, fr), Croat 27213 (MO). Panamá: El Llano--Cartí road, ca. 350 m, Mar. 1973 (stam. fl), Croat 22894 (MO); Feb 1973, Kennedy et al. 2394 (MO); Feb 1973, Kennedy et al. 2397a (MO); Mar. 1973 (fl, fr), Liesner 658 (MO); Mar. 1973 (stam. fl), Liesner 1208 (MO); Nov. 1985 (fl), McPherson 7585 (MO); Feb 1975 Mori et al. 4701 (MO); Feb 1975 (fl, fr), Mori et al. 4705 (MO). San Blas Comarca: El Llano--Cartí road, 13.8--15.8 km N of Interamerican Highway, 325 m, Aug. 1984 (stam. fl), de Nevers et al. 3747 (MO); 19.1 km N of Interamerican Highway, 350 m, Nov. 1984 (fr), de Nevers 4300 (MO). Veraguas: Atlantic slope NW of Santa Fe, 450550 m, Dec. 1974 (stam. fl), Mori et al. 3875 (MO).
Clusiella macropetala Cuatrecasas, Ann. Inst. Biol. Mex. 20: 110. 1949. TYPE: Colombia. Valle del Cauca: Río Cajambre, La Trojita, 5--50 m, 29 Feb 1944 (stam. fl), Cuatrecasas 16578 (holotype, F; isotype, US).
Bud scales deciduous, the scars clustered at or just above the
nodes. Petioles 3--10 mm, shallowly channeled; leaf blades elliptic,
(10--)13--19 x (4--)6--8 cm, the base acute to rounded, the apex
acute to acuminate to a 5--15 mm-acumen, in dried material glaucous
on both surfaces, pale gray-green above, tan below, distinctly
coriaceous; lateral veins 3--5(--7) mm apart, merging to a
submarginal vein ca. 1 mm from margin, the intersecondaries often
± prominent; translucid resin dots and dashes visible only by
intense light. Inflorescences unbranched or branched several times,
with ca. 1--6 flowers; flower buds ca. 7--10 mm diam.; petals 10--17
x 6--10 mm. Staminate flowers with a staminal column, including
capitulum, 5--10 mm tall. Fruits 8--12 mm diam., the stigmatic crest
with ca. 10 closely packed stigmas, fruit wall rather thick, ca. 0.7
Distribution. Colombia, from near sea level to 700 m.
As noted above, the leaves of C. macropetala are glaucous
on both surfaces and the fruits have a thick, several-layered fruit
wall that is bony when dry. Smaller-leaved and smaller-flowered forms
may otherwise be confused with C. elegans and C.
isthmensis (q.v.), but C. macropetala is probably more
closely related to the latter.
Additional specimens examined. COLOMBIA. Antioquia:
Anorí, Valle de Anorí, entre Dos Bocas y Norí,
700 m, June 1971 (stam. fl), Soejarto 2869 (F, MO, GH).
Chocó: Quibdó--Istmina road to Lloró, ca.
100 m, June 1983 (fl, fr), Forero et al. 9562 (MO), June 1983 (stam. fl), Forero et al. 9591 (MO), June 1983 (fl, fr), Forero et al. 9596 (MO), June 1982 (stam. fl), Gentry & Brand 36855 (MO), June 1982 (fl, fr), Gentry & Brand 36958 (MO), June 1982 (fl, fr), Gentry & Brand 36963 (MO), Aug. 1976 (stam. fl), Gentry & Fallen 17817 (pro parte, mixed collection with C. elegans, MO); San José del Palmar, hoya del Río San Juan, alrededores de Docordó, 0 m, Mar. 1979 (stam.? fl), Forero et al. 4328a (MO); Tutuendo--Quibdó road to Tubadó, 90 m, Jan. 1979 (fr), Gentry & Renteria 24507 (MO). Valle del Cauca: Bajo Calima, Buenaventura, 100 m, Jan. 1988 (stam. fl), Monsalve 2074 (MO); Río Cajambre, 580 m, May 1944 (stam. fl), Cuatrecasas 17497 (F, US).
Clusiella pendula Cuatrecasas, Ann. Inst. Biol. Mex. 20: 109. 1949. TYPE: Colombia. Valle del Cauca: Bajo Calima, Buenaventura, Bahia de Buenaventura, 0--10 m, 20 Feb 1946 (stam. fl), Cuatrecasas 19877 (holotype,
F; isotype, US)
Clusiella cordifolia Cuatrecasas, Rev. Acad. Colombiana
Cien. 8: 61. 1950. Syn. nov. TYPE: Colombia. Valle del Cauca: Bajo
Calima, Buenaventura, coastal thickets, Killip 11731 (holotype US; isotype GH).
Bud scales deciduous, the scars clustered at or just above the
nodes. Petioles ca. 3--5 mm, shallowly channeled; leaf blades
elliptic to cordate, (9.5--)11.5--16(--19) x (3.5--)4--6.5(--8) cm,
the base acute or rounded to sometimes cordate, the tip acuminate to
a ca. 2 cm acumen, in dried material often shiny, (not glaucous) on
both surfaces and dark olive above, dark brown below; lateral veins
2.5--4 mm apart, merging to a submarginal vein ca. 1 mm from margin,
intersecondaries often also quite prominent; resin dots and dashes
translucid. Inflorescence usually much-branched and with several
flowers; flower buds ca. 5 mm diam.; petals ca. 10 x 5 mm,
purplish-brown (maroon) to red. Staminate flowers with a staminal
column, including capitulum ca. 3 mm tall. Fruits globose or slightly
ovoid, ca. 15--20 x 10--15 mm, green(?) at maturity; stigmas ca. 8 or
9, closely packed and difficult to distinguish on fruits.
Distribution. Coastal Colombia and Ecuador from near sea
level to ca. 800 m.
This species is easy to distinguish by virtue of its dark red
instead of white or pinkish flowers and especially by its
dark-drying, shiny leaves with prominent lateral veins. For the shape
and color of its leaves it is most similar to the amazonian C.
The many new collections from Colombia, Valle Dept., Buenaventura
in Bajo Calima, near the type locality of both C. pendula and
C. cordifolia, allow the conclusion that the latter should be
considered a synonym of the former. The leaf color, texture and
venation of the type of C. cordifolia coincide with those of
C. pendula. Although the type specimens differ in shape of the
leaf base, it is now obvious that this can vary from acute through
rounded to slightly cordate even on a single specimen.
A label or numbering mix-up may have occurred with the type
specimen of C. cordifolia. As noted by Cuatrecasas in the
original description of the species, the label reads "Tree, fruit
blue." However, not only are trees and blue fruits otherwise unknown
in the genus, the specimen is of a flowering staminate individual!
Additional specimens examined. Colombia. Valle del
Cauca: Bajo Calima, Buenaventura, Bahia de Buenaventura, Apr.
1939 (stam. fl), Killip 34966 (F, US); coastal
thickets, Oct. 1922 (stam. fl), Killip 11731 (GH, US);
el Forge, sea level, June 1944 (fl, fr), Killip &
Cuatrecasas 38962 (F, US); Carretera Hans, km 22, 50 m,
Dec. 1987 (fl), Gentry et al. 59554 (MO);
Concesión Cartón de Colombia, 50--100 m, July 1987
(stam.? fl), Faber-Langendoen & Renteria
1209 (MO), Dec. 1981 (sterile), Gentry 35301 (MO),
Feb 1983 (stam. fl), Gentry et al. 40394 (MO),
Mar. 1986 (stam. fl), Gentry et al. 53618 (MO),
Apr. 1987 (stam. fl), Gentry et al. 56701 (A,
MO), Nov. 1986 (stam. fl), Monsalve 1206 (MO), Dec.
1986 (stam. fl), Monsalve 1405 (MO), Apr. 1987 (stam.
fl), Monsalve 1457 (MO), Sep. 1987 (fr),
Monsalve 1823 (MO), Oct. 1987 (stam. fl),
Monsalve 1892 (MO), Oct. 1987 (fr), Monsalve
1933 (MO), Oct. 1987 (stam. fl), Monsalve 1980
(MO), Jan. 1988 (stam. fl), Monsalve 2083 (MO), June
1982 (stam. fl), Murphy 558 (MO); Concesión
Pulpapel, 100 m, Aug. 1984 (fr), Monsalve 324 (GH);
Quebrada Algeria, Bahia Malaga, 50 m, Dec. 1985 (pist. fl),
Gentry et al. 53334 (MO); Río Calima, La
Esperanza, 5--10 m, Mar. 1944 (stam. fl), Cuatrecasas
16761 (F, US); road to Juanchaco Palmeras, 100 m, July 1984
(pist. fl), Gentry et al. 47819 (MO), July 1984
(fr), Gentry et al. 48291 (MO). ECUADOR.
Esmeraldas: San Lorenzo, 22 km from Lita on road to San
Lorenzo, 800 m, May 1990 (fl), Gentry et al.
69990 (MO); Alto Tambo, 15 km oeste de Lita, 400 m, Sep. 1990
(fr), Rubio 618 (MO); Carretera Lita--Alto Tambo--La
Punta, 400 m, Feb. 1991, Gudiño &
Moran 1263 (MO).
Relationships of the Genus
Discussing the placement of Clusiella within the family,
Planchon and Triana (1860) pointed out that the contorted aestivaton
of the petals was characteristic of the tribe Moronobeae (subfamily
Moronoboideae of Engler, 1925). However, because of the sessile,
largely cupuliform stigmas and nonascendent ovules of
Clusiella, they preferred to place it in their tribe Clusieae,
although they were not certain of their disposition because of the
lack of fruits and staminate flowers. Engler (1925) also felt that
these "incompletely known" plants "probably belong" next to Clusia
in subfamily Clusioideae. Subsequent published accounts have made
no explicit statements about possible relationships of
Clusiella, although most imply a close relationship with
Clusia (but see Hammel, 1984, unpublished thesis).
Numerous additional characters, including interpetiolar stipliform
structures, bud scales, resin gland-dots in the leaves, psilate
pollen exine, baccate fruits, small fovelate seeds that lack an aril,
and an embryo with unusually large cotyledons, are now known for
Clusiella and make this genus very unusual if not misplaced
among the clusioid genera. Stipuliform structures are uncommon in
Clusiaceae, but ones similar to those found in Clusiella also
occur in the Moronoboideae (e.g., Moronobea, Symphonia)
and in Garcinia s.l. Bud scales also occur in Moronobea
and Symphonia (and in some related genera), while some
species of Garcinia have one or two pairs of bud scales, and
bud scales occur in some of the Calophylloideae. Contorted petal
aestivation does not otherwise occur within the whole subfamily
Clusioideae (Engler, 1925), but is most likely the primitive state
for the family (P. Stevens, pers. comm., 1987). The psilate pollen
exine of Clusiella also appears to be otherwise unknown within
the family except for Symphonia and Moronbea (Cf.
Seetharam, 1983: It should be noted that Seetharam's study and light
micrographs of "Clusiella elegans Klug 1950,"
describing it as having "perreticulate" exine, pertain to a species
of Quapoya (i.e., Clusia), not Clusiella). The
small, foveolate seeds and embryo with well-developed cotyledons of
Clusiella are most like those in the Hypericoideae and basal
Calophylloideae, e.g. Marila. Were it not for its epiphytic
habit, dioecy, and resiniferous androecium (all reminiscent of
Clusia), but particularly because of its small foveolate seeds
and embryo structure, Clusiella would seem to be closely
related to Symphonia. More detailed, comparative studies of
pollen and the foliar resin system in particular are needed.
Characters such as those noted above in a sense pull Clusiella
strongly outward and downward from Clusia. Work in
progress towards an understanding of relationships among genera in
the whole family, in fact suggests that Clusiella may be
sister to a large clade including both the Clusioideae and
Moronoboideae (P. Stevens, pers. comm.). The diminutive
Clusiella, distinctively derived in many ways, and long
masquerading as Clusia's little cousin, seems to be an epiphytic
refugium for a number of relatively primitive characters in the
Clusioideae-Moronobiodeae, and could be called, more accurately,
Clusia's great aunt.
Acknowledgments. This work was supported, in part, by funds
from the National Science Foundation through grants (eg.,
DEB-9300814) to the author and co-PI M. H. Grayum for the Manual to
the Plants of Costa Rica. An early draft of the manuscript benefitted
greatly through correspondence with Peter Stevens. I thank John
Meyers for the line drawings of the two new species.
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