Synopsis of Chrysochlamys (Clusiaceae: Clusioideae:
Clusieae) in Mesoamerica
Barry E. Hammel
Missouri Botanical Garden
P. O. Box 299
St. Louis, Missouri, 63166
Novon 9: 360--374. 1999.
Abstract. Delimitation of genera in the Tovomita group of
tribe Clusieae has fluctuated, in Central America, with almost every
new floristic work. Nevertheless, with some aberrations, the theme of
a unified Chrysochlamys Tovomitopsis has persisted, and
is here supported by concurring with Bentham and Hooker's placement
of Tovomitopsis in synonymy of Chrysochlamys. The
strictly South American Balboa is also placed in synonymy of
Chrysochlamys, which action necessitates a new name (here
provided) for its only species. A key to the 13 species of
Chrysochlamys recognized for Mexico and Central America is
provided. An annotated list of all, including brief descriptions,
taxonomic and nomenclatural notes, five new combinations, and two new
species, brings forth that concept and rectifies the nomenclature to
the practicalities of numerous regional floras. Tovomita croatii Maguire, a somewhat problematic species, transferred
to Tovomitopsis by D'Arcy, is included in the key to
Chrysochlamys and fully discussed.
The Tovomita group of the tribe Clusieae (Clusiaceae,
subfamily Clusioideae) comprises primarily arborescent, terrestrial
genera with succulent-capsuled fruits and 1(rarely 2) large arillate
seeds per locule. Three genera therein, Tovomita,
Chrysochlamys, and Tovomitopsis have suffered a
particularly tortuous taxonomic history. Planchon and Triana (1860)
made the genus Tovomitopis to accommodate Sprengel's
Bertolonia, a later homonym; Bertolonia Raddi
(Melastomataceae) is now conserved over Bertolonia Spin
(Myoporaceae) (Farr et al., 1979). Planchon and Triana distinguished
Tovomitopsis from Chrysochlamys claiming it had an
arilloid (originating from the micropyle) rather than the true aril
(originating from the funicule) they claimed for
Chrysochlamys. According to them, the so-called aril of
Tovomita is neither, but rather the outer integument of the
seed. These differences were further supported by floral structure:
Tovomitopis, calyx 4-merous; Chrysochlamys, calyx
5-merous; Tovomita, calyx 2- or 4-parted, the outer two sepals
overtopping the bud. Planchon and Triana also described Balboa
in the same publication with Tovomitopsis, but they placed it
in their subtribe "Euclusiae" rather than Tovomiteae. Bentham and
Hooker (1862: 167--176) recognized Balboa,
Chrysochlamys and Tovomita (as related) but treated
Tovomitopsis as a synonym of Chrysochlamys. However,
they transferred no species. Subsequently, Balboa was
apparently forgotten in discussions relating to the Tovomita
group. Hemsley (1879: 87) transferred all Central American species of
Tovomitopsis to Chrysochlamys. Cuatrecasas (1950) also
declared unjustified the separation between Chrysochlamys and
Tovomitopsis and transferred two (South American) species to
Chrysochlamys. Williams (1959) subsumed Tovomitopsis
within Tovomita, but kept Chrysochlamys separate.
Maguire (1977) tentatively recognized all three genera. D'Arcy (1980)
expressed the same taxonomic opinion begun by Bentham and Hooker and
put into practice by Hemsley and Cuatrecasas, i.e., a unified
Chrysochlamys/Tovomitopsis, but used the later name,
Tovomitopsis. D'Arcy also elevated Engler's section
Dystovomita of Tovomita to the genus
Most recently, an attempt has been made to circumscribe
Chrysochlamys as comprising species with succulent petals,
echinate pollen exine, and black seeds with white arils, rather than
membranaceous petals, psilate-foveolate pollen, and green seeds with
orange arils (cf. Hammel, 1984, 1986). Ignoring whether or not these
characters support a monophyletic group or even if they coincide,
typologically, with Chrysochlamys, neither such a separation,
nor one based on the considerations of Planchon and Triana is
practical. Dystovomita, with axillary inflorescences (i.e.,
mostly lateral and below the leaves), large, adaxial petiolar pits
and relatively small seeds, is easily recognized and may even lie
closer to Clusia (P. Stevens, in prep.). Tovomita, with
terminal inflorescences, leaves clustered at the ends of side shoots,
leaves sometimes with petiolar pits, outer two sepals fused in bud or
at least overtopping the bud, styles often evident, fruits usually
with the inner wall and placenta dark reddish wine, can also be
recognized, even with most herbarium material. The older name,
Chrysochlamys, takes precedence over Tovomitopsis for
the remaining species in the group, i.e., those with terminal
inflorescences, leaves not clustered on side shoots, lacking adaxial
petiolar pits, sepals (usually) not overtopping the bud, styles
lacking, and the inner fruit wall and placenta mostly white or pink.
Balboa is a synonym of Chrysochlamys.
The above paragraph can be summarized by the following key:
- 1a. Inflorescences apparently axillary; leaves with large,
adaxial, margined petiolar pits; seeds ca. 6
- 1b. Inflorescences usually terminal; leaves mostly lacking
petiolar pits; seeds ca. 1 cm or more.
- 2a. Leaves clustered at the ends of side shoots; outer two
sepals fused in bud or at least overtopping the bud; styles often
evident; inner fruit wall and placenta usually dark reddish
- 2b. Leaves ± evenly dispersed along the branches; outer
two sepals never fused in bud, nor both overtopping the bud; inner
fruit wall and placenta mostly white or
Chrysochlamys Poeppig, Nov. Gen. Sp. Pl.. 3:13, t. 211.
1842. TYPE: Chrysochlamys multiflora Poeppig.
Balboa Planchon & Triana, Ann. Sci. Nat. Bot., ser. 4,
13: 315, 14: 252. 1860. TYPE: Balboa membranacea
Planchon & Triana.
Tovomitopsis Planchon & Triana, Ann. Sci. Nat. Bot.,
ser 4, 14: 261. 1860. TYPE: T. paniculata (Sprengel)
Planchon & Triana. Basionym: Bertolonia paniculata
Dioecious, terrestrial trees or shrubs, sometimes epipetric
rheophytes, the branching decussate and the leaves dispersed more or
less evenly along the branches; resin mostly clear, occasionally
white, rarely yellow; internodes mostly without bud-scale scars.
Petiolar pits and interpetiolar stipuliform structures absent. Leaves
opposite, the blades elliptic; major lateral veins mostly prominent,
resin canals sometimes visible in dried material. Inflorescences
terminal (rarely left pseudo-axillary by the dominant growth of an
axillary, vegetative bud), many-flowered, panicles. Flowers 4- or
5-merous, corolla aestivation imbricate, petals thin, membranaceous,
to thick and succulent, pink, or yellow to white. Staminate flowers
with numerous (rarely few) ± separate stamens, or the filaments
fused basally into a short column, or rarely with fertile stamens
surrounding a central sterile capitulum, or very rarely the anthers
all sessile on a resiniferous capitulum. Pistillate flowers with
numerous separate staminodia, or staminodia mounted on a ring or
rarely just a ring surrounding the ovary; ovary with mostly 4 or 5
stigmas, rarely 3. Fruits globose to ovoid, usually terete, rarely
winged or star-shaped in cross section, succulent capsules mostly
with 1 seed maturing per locule, sometimes only 1 seed per fruit;
seeds green to nearly black with an orange or white aril.
Distribution and ecology. Chrysochlamys is known
from Mexico to Bolivia and Brazil, where it occurs in wet to very wet
primary forest from near sea level to at least 2300 m.
Chrysochlamys, in total, contains perhaps as many as 50
species, most of them South American, and is in much need of
revision. At least two Central American species, C.
glauca (Oersted, Planchon & Triana) Hemsley and C.
nicaraguensis (Oersted, Planchon & Triana) Hemsley, have
been erroneously reported from South America, and numerous South
American names (e.g., C. membranacea Planchon &
Triana, C. micrantha Engler, C. myrcioides Planchon & Triana) have been loosely (and erroneously) applied to Central American species, as well as within South America. The following key, distinguishing 13 species of Chrysochlamys for Mexico and Central America, and the subsequent annotated list, including citation of representative specimens and descriptions of two new species from Costa Rica and Panama, aim to clarify the species and names for the region. One or two possible new species, from among Panamanian material at hand, await resolution and are not included here.
Key to the species of Chrysochlamys (and one
Tovomita) in Mesoamerica
- 1 Internodes with a pair of bud scales (or their scars),
usually just above the subtending node; primary lateral veins
close (ca. 3 mm apart), ± prominent but difficult to
distinguish from the intersecondaries; inflorescences often
appearing axillary; anthers sessile on a capitulum; stigmas 3; far
eastern Panama to
- 1' Internodes without bud-scales; primary lateral veins
distant (usually 1 cm or more), they and the intersecondaries
obscure or prominent; inflorescences unambiguously terminal;
anthers with filaments; stigmas (4)5 or 6; Guatemala to northern
- 2 Leaves mostly < 2.5 cm wide, often drying tan; lateral
veins very obscure in dry material.
- 3 Flower buds 6--9 mm long; stamens 30--40; inflorescence
and infructescence stout, fruits pyriform, but not rostrate,
mostly 5-seeded; epipetric rheophytes................C.
- 3' Flower buds 3--4 mm long; stamens ca. 20; inflorescence
and infructescence delicate, usually nodding or pendulous;
fruits narrowly obovoid or pyriform and usually rostrate,
mostly 1 or 2 (3) -seeded; small shrubs in forest and sometimes
tenuis sp. nov.
- 2' Leaves mostly over 3 cm wide, drying green, gray-green,
reddish tan, to tan; lateral veins easily seen in dry material.
- 4 Leaf blades nearly sessile, 20--60 cm long; small shrubs
< 2 m.............Tovomita croatii
- 4' Leaf blades distinctly petiolate, shorter; shrubs to
trees, often over 3 m.
- 5 Lateral leaf veins mostly < 9 pairs; calyx
deciduous in fruit.
- 6 Flower buds < 4 mm long; stamens ca. 20--30;
inflorescence delicate, often pendulous; lateral leaf
veins mostly ca. 1 cm apart; leaves usually dull, drying
gray-green, often with obvious resin canals; Costa Rica
and Panama..................C. glauca
- 6' Flower buds >5 mm long; stamens 35 or more;
inflorescence stout, erect; lateral leaf veins >1 cm
apart; leaves shiny.
- 7 Leaves without obvious intersecondary veins or
resin canals, the primary lateral veins usually 2--3
cm apart, the leaves drying gray-green; flower buds
ca. 5--7 mm long; stamens 35--40; Nicaragua, Costa
Rica, Panama.........C. nicaraguensis
- 7' Leaves with obvious intersecondary veins and
often the resin canals also obvious, the primary
lateral veins mostly 1--2 cm apart, the leaves drying
reddish tan; flower buds ca. 5 mm long; stamens ca.
50; Mexico, Guatemala, Belize,
- 5' Lateral leaf veins mostly > 10 pairs; calyx
persistent or not.
- 8 Submarginal leaf vein usually distinct, at least
above mid-leaf; calyx persistent in fruit.
- 9 Petals succulent; filaments fused basally, the
free portion shorter than the fused portion;
staminodia connate, forming a thin ring around the
base of the ovary; fruits terete; central to eastern
- 9' Petals membranaceous; filaments free to the
base; staminodia distinct; Costa Rica to central
- 10 Fruits star-shaped in cross section; anthers
usually 2--3 times longer than wide; outer two
sepals ± equal, shorter than the bud; flower
buds ca. 7 mm long; inflorescence glabrous or
- 10' Fruits terete; anthers about as wide as
long; outer sepals very unequal, the larger ±
equal to the bud; flower buds ca. 5 mm long;
inflorescence glabrous...C. skutchii
- 8' Submarginal leaf vein lacking or indistinct; calyx
deciduous or persistent.
- 11 Calyx deciduous in fruit; plants from the
mountains (mostly > 1000 m) of Costa Rica and
western Panama; lateral leaf veins 9--15 distinct
pairs; leaves drying tan to pink or green.
- 12 Fruits ± pyriform, ca. 2--2.5 cm long,
often greenish yellow at maturity and the stigmas
set to one side; stamens >50; relatively large
(ca. 15 m) trees with ± coriacious, tan- to
pink-drying leaves; 700--2000 m.........C.
- 12' Fruits usually obovoid-mammillate, often 4
cm or more long, mostly red at maturity, the
stigmas centered; stamens mostly ca. 15--20; shrubs
or small trees (ca. 7 m or less) with
membranaceous, mostly green-drying leaves;
- 11' Calyx persistent in fruit; plants from lowland
Costa Rica (30--900 m) or from Central and eastern
Panama; lateral leaf veins 10--17 pairs, sometimes
nearly indistinguishable from the intersecondaries;
leaves usually dark, reddish brown to tan-drying.
- 13 Fruits strongly alate; leaves narrowly
elliptic, often falcate, ± coriaceous, the
apex gradually acuminate, the resin canals very
obvious on dry material; outer pair of sepals
± equal and nearly as long as the bud; resin
canals of petals obvious; resin of stems white;
lateral leaf veins 10--13; far eastern
- 13' Fruits not alate; leaves oblong to
elliptic, not falcate, membranaceous, the apex
abruptly acute to acuminate, the resin canals
obscure; outer pair of sepals very unequal, the
smaller much shorter than the bud; resin canals of
petals obscure or lacking; resin of stem clear or
- 14 Stamens ca. 50; leaves broadly elliptic,
mostly 8--10 cm wide; lateral leaf veins mostly
11--17 distinct pairs; resin of stems clear;
- 14' Stamens 20--30; leaves elliptic to
oblong, mostly 5--6(--9) cm wide; lateral leaf
veins mostly 8--10 somewhat indistinct pairs,
often difficult to distinguish from the
intersecondaries; resin of stems white; Costa
Chrysochlamys allenii (Maguire) Hammel, comb. nov. Basionym: Tovomitopsis allenii Maguire, Phytologia 38: 214. 1978. TYPE: Panama. Bocas del Toro: Robalo Trail, northern slopes of Cerro Horqueta, 2000--2300 m, Allen 5014 (holotype, NY; isotype, MO).
Tovomitopsis centistaminibus D'Arcy, Ann. Missouri
Bot. Gard. 67: 1030. 1980. Syn. nov. TYPE: Panama. Chiriquí: 8
km N of Camp El Volcán, 2000 m, Little 6070
(holotype, MO; isotype, US).
Trees 5--25 m, sometimes with stilt roots, occasionally reported
as epiphytic; resin clear to creamy. Leaf blades elliptic to slightly
obovate, widest just above the middle, 11--16 X 4--6 cm, the apex
rounded to acuminate, the base acuminate, coriaceous; lateral veins
ca. 0.8--1 cm apart, ca. 9--12 pairs, reaching to the margin; petiole
1--2 cm. Inflorescence 8--12 cm, erect, puberulent; flower buds ca.
5--7 mm; stamens 50 or more, anthers of both stamens and staminodia
about as wide as long, < 1 mm, filaments free; stigmas large (ca.
2 mm), distinctly spreading. Fruits pyriform, 2--2.5 cm,
yellow-green, the stigmas often mounted slightly to one side; calyx
deciduous in fruit.
Distribution. Very wet cloud forest, (700--)1450--2300 m;
continuous from the Cordillera de Guanacaste in northwestern Costa
Rica to Chiriquí province of western Panama, disjunct from
Darién province of far eastern Panama.
A larger tree than any other Chrysochlamys in Central
America and growing at higher elevation, C. allenii is
further distinguished by its coriaceous leaves, numerous stamens
(more than 50), often very large stigmas, and pyriform, often
greenish yellow fruits.
Although D'Arcy (1980) compared Tovomitopsis centistaminibus with T. psychotriifolia, the type specimen's high-elevation habitat, large stature, coriaceous leaves, and numerous stamens conspire to alienate it from the latter species, and instead, to bring it into synonymy with C. allenii. This species, however, can at times be confused with C. psychotriifolia, which usually grows at somewhat
lower elevations, has fewer (and often longer) stamens, and larger
fruits. Material at hand, especially that from far eastern Panama,
suggests that C. allenii may be closely related to,
perhaps not distinct from, C. colombiana (Cuatrecasas)
Representative specimens. Costa Rica. Guanacaste:
Volcán Cacao, 1100 m, Nov. 1990 (fr), Espinoza 24 (INB). Puntarenas: Monteverde Reserve, 1500 m, June 1985 (fl, fr), Hammel 13856 (MO); cloud forest above Wilson's finca S of San Vito de Java, 1600 m, Aug. 1967 (fr), Raven 21845 (MO). Heredia: 12 km above San Rafael, 1800 m, July 1967 (fl), Lent 1141 (MO). San José: La Palma, 1550 m, Aug 1898 (fl), Tondúz 7378 (MO). Panama. Chiriquí: 8 km N camp El Volcán, 2000 m, Mar. 1943 (fr), Little 6068 (MO); about 2 km NE of Guadelupe, 2 km N of Cerro Punta, Dec. 1971 (fr), Wilbur et al 15331 (DUKE, MO); Bajo Mono - Robalo trail, 1600--2300 m, July 1947 (fl),Allen 4839 (MO). Bocas del Toro: vicinity of Cerro Colorado, 1700 m, Apr. 1986 (fr), McPherson 8925 (MO); vicinity of Fortuna Dam, 1200 m, Apr. 1986 (fl), McPherson9049 (MO). Darién: Cerro Sapo, 800--1000 m, Mar. 1986 (fl, fr),
Hammel et al 14877 (MO).
Chrysochlamys angustifolia (Maguire) Hammel, comb. nov. Basionym: Tovomitopsis angustifolia Maguire, Phytologia 36: 399. 1977. TYPE: Panama. Veraguas, Mori & Kallunki 5357 (holotype, MO; isotype, NY).
Small shrubs to 2 m, often on rocks along streams; resin milky.
Leaf blades narrowly elliptic 7--13 X 1--2(2.5) cm, the apex acute to
acuminate, base acute, membranaceous, drying light gray- or tannish
green; lateral veins obscure, ca. 0.5--1 cm apart, ca. 5 or 6 pairs,
reaching to the margin; petiole ca. 0.5--1 cm. Inflorescence ca. 6
cm, erect, glabrous; flower buds 6--9 mm; stamens 30--40, anthers of
stamens and staminodia ± triangular, slightly longer than wide,
ca. 1 mm, filaments free; stigmas spreading to often erect. Fruits
pyriform, 1.5--2 cm, red or greenish red; calyx deciduous in fruit.
Distribution. Very wet, mostly lowland forest, usually
along streams, 100--1000 m; Bocas del Toro to Veraguas and
Coclé provinces of central Panama.
This distinctive, small, narrow-leaved, epipetric rheophyte with
flower buds over 6 mm long is endemic to a few lowland or mid-elevation sites in west central Panama. Its habitat, larger flowers, and stouter fruiting rachis easily distinguish it from the similarly narrow-leaved new species, C. tenuis, described below. Among Central American species, C. angustifolia is one of the few with copious milky resin rather than the more common clear to tardily creamy resin.
Representative specimens. PANAMA. Bocas del Toro:
Upper Río San Pedro, 500 m, July 1979 (fr), Gordon 59Da (MO). Veraguas: vicinity of Santa Fe, dirt road past Escuela Alto de Piedra, 490 m, June 1987 (fl, fr), Croat 66871 (MO). Coclé: along Río San Juan near fork with Río Tife, 400 m, June 1978 (fl, fr), Hammel 3329 (MO).
Chrysochlamys eclipes L. O. Williams, Trop. Woods 111: 15. 1959. TYPE: Panama. Canal Zone: Barro Colorado Island, Standley 31459 (holotype, US).
Shrubs or trees 2--10 m; resin clear. Leaf blades elliptic to
oblong, 12--26(--29) X 5--12 cm, the apex acute, base acute,
membranaceous, drying dark reddish tan or rarely greenish tan;
lateral veins obvious, 1--1.5 cm apart, 10--12 pairs, loop-connected
to a submarginal vein ca. 1--3 mm from the margin; petiole ca. 1--3
cm. Inflorescence ca. 7--21 cm, erect or pendulous, minutely
puberulous; flower buds ca. 4--5 mm; stamens ca. 25--30, anthers of
stamens and staminodia wider than long, <1 mm, filaments basally
fused into a succulent torus, the free part of filaments <1 mm,
much shorter than the fused part; stigmas rounded, pressed against
the ovary. Fruits mostly globose, 2--3 cm, green to greenish red or
red; seeds black with a white aril; calyx persistent in fruit.
Distribution. Wet to very wet forest, from near sea level
to 1600 m; central to eastern Panama.
A distinctive, usually dark-reddish brown-drying, dull-leaved,
Panamanian endemic (from the Canal to San Blas and Darién)
with succulent and yellow rather than membranaceous white or pink
petals, stamens with short, free filaments mounted on a succulent,
fused basal portion, white rather than orange arils, and black rather
than green seeds, not at all closely related to C.
nicaraguensis, under which it was synonymized by D'Arcy
(1980). This is the only species, described from the region, with
succulent, rather than membranaceous petals. Even in fruit it can be
identified by the combination of large, dark-drying leaves and fused
staminodia. The petals dry black or very dark burgundy.
Some of this material with particularly large inflorescences from
far eastern Panama has been identified as C. floribunda
Cuatrecasas and C. weberbaueri Engler However, the
abundant material at hand suggests that probably only one species is
involved. Pending study of type material of the older, South American
names, I here use C. eclipes for all Panamanian
Representative specimens. PANAMA. Colon: end of Río Boquerón road, 500--650 m, Apr. 1978 (fr), Hammel 2445 (MO); Río Guanche, Feb. 1974 (fl), Foster 2780 (MO); Santa Rita Ridge Road, 450 m, Mar. 1975 (fr), Mori & Kallunki 5049 (MO). Canal Zone: Barro Colorado Island, Mar. 1969 (fr), Croat 8320 (MO); Pipeline Road, Sep. 1971, (fl), Gentry 1946 (MO). Panamá: Cerro Jefe region, 750 m, Jan. 1986 (fr), McPherson & Merello 8134 (MO); Gorgas Memorial Labs yellow fever research camp, 25 km NE of Cerro Azul on Río Piedras, 550 m, Nov. 1974 (fr), Mori & Kallunki 3371 (MO); Llano-Carti road, 350 m, Aug. 1986 (fl), McPherson 9962 (MO). San Blas: Cangandi, 30 m, Feb. 1986 (fr),
de Nevers & Herrera 7186 (MO).
Darién: Cana-Cuasi trail, Chepijana, 1600 m, Mar. 1940
(fr), Terry & Terry 1603 (MO); Parque
Nacional del Darién, ridge between Río Topalisa and
Río Pucuro, ca. 17 km E of Pucuro, 850 m, Oct. 1987 (fl),
de Nevers et al.8380 (MO); slopes
of Cerro Mali, 1300--1400 m, Oct. 1987 (fl & fr), Cuadros
et al. 3927 (MO); S of El Real on trail up Cerro
Pirre, 550--1030 m, Mar. 1985 (fr), McPherson 7045
(MO); trail to Cerro Pirre, 20--600 m, Aug. 1986 (fl), J.
F. McDonagh et al. 483 (MO).
Chrysochlamys glauca (Oersted, Planchon & Triana)
Hemsley, Biol. Cent.-Amer., Bot. 1: 87. 1879. Tovomitopsis
glauca Oersted, Planchon & Triana, in Planchon &
Triana, Ann. Sci. Nat. Bot., Ser. 4, 14: 264. 1860. Tovomita
glauca (Oersted, Planchon & Triana) L. O. Williams,
Fieldiana Bot. 29: 357. 1961. TYPE: Costa Rica. [San José]:
sur le mont Jaris, [ca. 850 m], Oersted 3 
Tovomita gracilis L. O. Williams, Fieldiana Bot. 29:
357. 1961. TYPE: Costa Rica. Alajuela: San Pedro de San Ramón,
1050 m, Brenes 4217 (lectotype, designated by Hammel
Tovomitopsis multiflora Standley, Field Mus. Nat.
Hist. Bot. Ser. 4: 234. 1929. TYPE: Panama. Chiriquí:
Progreso, Cooper & Slater 171
(holotype, F--573,086; isotype, NY).
Shrubs or rarely trees 1--6(--14?) m; resin clear to slightly
milky. Leaf blades elliptic, 7--20 X 2--7 cm, the apex and the base
acute to acuminate, drying gray-green; lateral veins 0.5--1.5 cm
apart, 6--10 pairs, fading towards the margin; petiole 1--2.5 cm.
Inflorescence 2--12 cm, often pendulous, glabrous; flower buds ca.
3--4 mm, outer 2 sepals equal to markedly unequal, shorter than the
inner ones; stamens 20--30, anthers of stamens and staminodia ovoid
to ± globose, < 1 mm. Fruits pyriform to globose, 1--2 cm,
greenish cream to nearly red, often two-toned; calyx deciduous in
Distribution. Wet to very wet forest, from near sea level
to 1400 m; Costa Rica and Panama.
Even segregating the material described below as
Chrysochlamys tenuis, C. glauca remains a
rather variable species, characterized by its usually dull,
few-veined leaves and small flower buds. Some of that variation may
be worthy of taxonomic recognition. Many of the collections from the
Osa peninsula in Costa Rica (cf. Aguilar 431,
Kernan 713) have leaves with especially obvious and
numerous latex canals and flower buds near the lower end of the size
range for the species, giving such material an aspect predictive of
its origin. Allen (1956: 345), in fact, distinguished two species
from the Osa region within the C. glauca complex. For
one (vouchered by Skutch 5286, F, not seen), he used
the name C. costaricana, stating that it was "nearly
identical" to C. glauca, "differing principally in the
smaller flowers." Although C. costaricana (discussed
below under C. psychotriifolia) is not its correct
name, the species distinguished by Allen may deserve recognition, not
since given. Populations at the other end of Costa Rica, in the
Cordillera de Guanacaste, have almost shiny leaves with relatively
obscure lateral veins. The plants also appear to flower more on
lateral shoots than material from elsewhere. The anthers are slightly
longer and the stigmas are larger than those of C.
glauca from elsewhere, usually at least 1/2 (vs. no more than
1/3--1/4) the length of the ovary in newly opened flowers. Several
collections from Panama, Panama province and San Blas (de
Nevers 7223, Herrera et al.
1137, Herrera et al. 1145),
tentatively placed here, have unusually dense and much-branched resin
lines, and fruits with persistent sepals. These may represent a
distinct species, but flowering material is needed.
Representative specimens. Costa Rica. Guanacaste:
Volcán Cacao, Rancho Harold, 700--1200 m, July 1986 (fr),
I. & A. Chacón 2154
(CR); Parque Nacional Guanacaste, Estación Cacao, 1100 m, Nov.
1990 (fr), Chávez 368 (INB). Alajuela:
Río Cataratas, San Ramón, 600 m, Jan. 1984 (fl),
Gómez-Laurito 9700 (CR). Limón:
Cantón de Pococí, Barra del Colorado, 15--20 m, Jan.
1995 (fl), Araya 713 (CR, INB, MO) Puntarenas:
airport 4 mi. W of Rincón de Osa, 40 m, Aug. 1967 (fr),
Raven 21564 (CR); wooded slopes above Golfito, 0--80 m,
Jan. 1967 (fl), Burger & Mata 4775
(CR, F); Las Cruces, Jardín Botánico Wilson, 1100 m,
Sep. 1993 (fr), G. Vargas et al.
1369 (CR). PANAMA. Bocas del Toro: Río Teribe,
alrededores de la estación Teribe II del IRHE, Oct. 1984,
Carrasquilla 2026 (MO). Chiriquí: E of
Cañas Gordas near Costa Rican border on road to Volcán,
800--1200 m, Feb. 1973 (fl, fr), Liesner 268 (MO); road
from Nueva California to Río Serano, 1400 m, Apr. 1979 (fl),
D'Arcy et al. 13057 (MO).
Panamá: Llano-Cartí road, 400 m, Jan. 1974 (fl),
Nee & Dressler 9371 (MO). San
Blas: Cordillera de San Blas, cabecera del Río Piriati,
460--799 m, Feb. 1992 (fr), Herrera et al.
Authorship of the four Central American Tovomitopsis names
published in Planchon and Triana (1860) has been variously obfuscated
in numerous publications, including my own. Oersted alone never
published these names, as sometimes implied (cf. Williams, 1959),
nor is there any basis for the even more common "Oersted ex Planchon
& Triana," (cf. Standley, 1937; Croat, 1978; D'Arcy, 1980), let
alone the terse permutation of the latter to just "Planchon &
Triana" (e.g., Hammel, 1986). Since Anders Örsted (1812--1872)
was professor of botany at the University of Copenhagen until 1862
during the time Planchon and Triana were working in Paris on their
Guttiferae memoir, nothing suggests other than the three, "Oersted,
Planchon & Triana," as co-authors, as originally indicated for
all four species in Planchon and Triana (1860).
The citation of two collection (?) numbers for the types of these
same four species results from examination of the actual specimens.
The single-digit numbers are cited in the original publication (e.g.,
Oersted 4), and are also written on or in packets with
the specimens. The four-digit numbers printed on the labels (e.g.,
Plantae Centroamericanae Oersted No. 3585) are not mentioned in the
publication. Neither series appears to come from normal consecutive
field-note numbers; earlier dates do not correspond, consistently, to
lower numbers within the series.
Chrysochlamys grandifolia (L. O. Williams) Hammel, comb.
nov. Basionym: Tovomita grandifolia L. O. Williams,
Trop. Woods 112: 96. 1960, new name for T. macrophylla
L. O. Williams, Trop. Woods 111: 16. 1959, non T.
macrophylla (Poeppig) Walpers 1842. TYPE: Costa Rica.
Puntarenas: Esquinas forest, between Río Esquinas and Palmar
Sur, 30 m, Allen 5968 (holotype, US; isotypes, EAP, F).
Shrubs or trees 2--10 m; resin clear to yellowish. Leaf blades
elliptic, 18--35 X 10--18 cm, the apex acute to rarely rounded, the
base acute to acuminate, drying reddish; lateral veins 1--2 cm apart,
14--18 pairs, looping to form a ± distinct submarginal vein;
petiole (2--)3--4 cm. Inflorescence ca. 6--10 cm, erect, glabrous;
flower buds ca. 7 mm, outer 2 sepals ± equal to each other and
shorter than the inner ones; stamens ca. 50, anthers of stamens and
staminodia longer than wide, ca. 2 mm. Fruits ovoid, strongly 5- or
6-ribbed (star-shaped in cross section), greenish cream to nearly
red, often two-colored; calyx persistent in fruit.
Distribution. Very wet forest, 10--600(--1300) m; Costa
Rica to central Panama.
This distinctive Central American species, clearly delimited by
its protologue, is characterized by strongly ribbed fruits and
monomorphic stamens with anthers up to 2--3 times longer than wide.
Costa Rican material from the Pacific slope has glabrous
inflorescences and anthers nearly 2 mm long, whereas that of the
Caribbean slope (also occurring at higher elevation than that of the
Pacific) and the Panamanian collections have markedly puberulent
inflorescences and smaller anthers.
This species was originally published as Tovomita
macrophylla L. O. Williams, a homonym of the earlier T.
macrophylla (Poeppig) Walpers Williams, therefore, made a new
name, T. grandifolia, for the same species.
Coincidentally, an earlier Chrysochlamys macrophylla
Pax would prevent the use of that combination for the present
species, in any case. Nothing, however, prevents the use of the
combination C. grandifolia (L. O. Williams) Hammel.
Although D'Arcy (1980) created the combination Tovomitopsis
myrcioides (Planchon & Triana) D'Arcy for this material
(with Tovomita grandifolia as synonym), the South
American C. myrcioides Planchon & Triana is a quite
different and poorly known species. It has very short petioles and
the dimorphic stamens of section Heterandra "externa fertilia,
libera, interna sterilia in corpus centrale concreta" (Planchon &
Triana, vol. 14: 260. 1860) with nearly globose anthers. Furthermore,
I know of no such rib-fruited Chrysochlamys species from South
America. Chrysochlamys membrillensis (see below) of far
eastern Panama also has somewhat ribbed fruits, but is otherwise
quite different from the present species.
Representative specimens. Costa Rica. Cartago:
Turrialba; 2 km al oeste de Grano de Oro, 1200 m, July 1995 (fl),
Cascante 639 (CR). Limón: Cord.
Talamanca, entre Cerro Chimú y Cerro Matama, 1200 m, Abr. 1985
(fl, fr), Gómez & Herrera
23536 (CR); fila entre cuenca superior del Río Xichiari
y cuenca superior del Río Boyei, 1300 m, Ago. 1995 (fl),
Herrera 8427 (CR). San José: ca. 20 km S
of San Isidro de El General, El Pilar de Cajón, 620 m, Nov.
1988 (fr), Zamora et al. 1521 (CR, MO).
Puntarenas: Cantón de Osa, Fila al Oeste de Rancho
Quemado, 300 m, Aug. 1991 (fr), J. Marín
127 (CR, MO); Sierpe, cabeceras de Quebrada Guerra, 200 m,
June 1990 (fl), Herrera 4240 (CR, INB, MO); Corcovado
National Park, trail from Los Chiles to base of hills, 20--400 m,
July 1977 (fl), Liesner 3175 (CR, MO); Los Planes (La
Gloria), 100 m, Feb. 1991 (fr), R. González
11 (CR, F, INB, MO); Cantón de Golfito, Refugio
Nacional Golfito, S end of Fila Gamba, 160--200 m, Dec. 1988 (fr),
Grayum et al. 9180 (CR, MO). PANAMA.
Bocas del Toro: road from Fortuna Dam to Chiriquí
Grande, 400 m, Apr. 1986 (fl), McPherson 9020 (MO).
Chiriquí: Camino hacia la finca Landau, NE del
campamento de Fortuna (Hornito), sitio de presa, 1100 m, Aug. 1976
(fr), Correa et al. 2339 (MO).
Veraguas: 5 mi. W of Santa Fe on road past Escuela
Agrícola Alto Piedra, 800--1200 m, Mar. 1973 (fr),
Croat 23151 (MO). Coclé: El Copé,
Atlantic slope E of sawmill, 900 m, Feb. 1979 (fr), Hammel
6271 (MO). Colón: Santa Rita Ridge Road, 600 m,
Feb. 1980 (fl), Antonio 3756 (MO). Panamá:
between peaks on Cerro Trinidad, May 1968 (fl), Kirkbride
& Duke 1647 (MO); Cerro Campana, near tower,
Mar. 1977 (fr), Folsom et al. 2311 (MO).
Chrysochlamys guatemaltecana Donnell-Smith, Bot Gaz. 13:
26. 1888. Tovomitopsis guatemaltecana (Donnell-Smith)
Standley & Steyermark, Publ. Field Mus. Nat. Hist., Bot. Ser. 23:
66. 1944. TYPE: Guatemala. Alta Verapaz: von Tuerckheim
989 (holotype, US; isotype, NY).
Trees or shrubs 3.5--15 m; resin clear to pinkish. Leaf blades
elliptic 12--25 X 5--9(--10) cm, the apex acute, the base ±
acuminate; drying tan to grayish tan and usually with rather obvious
resin lines; lateral veins 1--2 cm apart, ca. 8--12 pairs, fading to
the margin; petiole 1.5--2 cm. Inflorescence 7--15 cm, erect,
glabrous; flower buds ca. 5 mm, the outer 2 sepals ± equal, <
1/2 the length of the bud; stamens ca. 50, anthers of stamens and
staminodia about as long as wide, < 0.5 mm. Fruits pyriform,
2.5--4 cm, red tinged with green; calyx deciduous in fruit.
Distribution. Wet forest; 100--1450 m. Mexico to Belize.
This name has been considered a synonym of Chrysochlamys
nicaraguensis (cf. Standley & Williams, 1961) and
C. psychotriifolia (cf. Hammel, 1984). However, a
clearer picture of variation within the genus throughout the region
now supports its reinstatement. The species is characterized by more
or less shiny, usually reddish-brown-drying leaves with rather few
(6--9) and distant (1--2 cm) major lateral veins and usually quite
prominent intersecondaries and obvious resin lines. The rather large
flower buds (5--7 mm) have visible resin lines. It has more stamens
(ca. 50) than either C. nicaraguensis (35--40) or
C. psychotriifolia (ca. 20). The fruits are often quite
elongate, up to 4 cm, as in C. psychrotiifolia, and the
sepals are early deciduous.
Representative specimens. MEXICO. Oaxaca: 18 km al
este de Cuahutemoc, por la carretera Cuahtemoc-Sarabia, Dec. 1974
(fr), Vazquez et al. 1482 (MO); Mpio.
Matías Romero, 8.3 km al S de Esmeralda, luego 2.3 km al E,
110 m, May 1981 (fl, fr), Wendt et al.
3267 (MO); Mpio. Sta. María Chimalpa, Río Verde,
360--430 m, July 1986 (fl), Caletti et al.
210 (MO); Mpio. Minatitlán, lomas al S del Poblado 11,
ca. 30 km al E del Campamento La Laguna, 180 m, July 1980 (fl, fr),
Wendt et al. 2642 (MO). GUATEMALA.
Alta Verapaz: 7 mi. up road to Oxec, 700 m, July 1977 (fl),
Croat 41631 (MO). Izabal: Puerto Mendez, km 7,
Sep. 1970 (fl), Contreras 10237 (MO). BELIZE.
Toledo: Central camp, Edwards Road beyond Columbia, June 1951
(fl), Gentle 7354 (MO); SW Maya Mountains, Columbia
River Forest Reserve, 750 m, Apr. 1992 (fl), Holst 4407
(MO); vicinity of San José Mayan Indian village, June 1973
(fl, fr), Croat 24450 (MO). HONDURAS.
Copán: Parque Nacional Cerro Azul, 1440 m, Feb. 1992
(fr), Hawkins & Mejía 184
(MO). Olancho: Refugio de Vida Silvestre La Muralla, 1400 m,
June 1992 (fl), Mejía 164 (MO).
Chrysochlamys membrillensis (D'Arcy) Hammel, comb. nov.
Basionym: Tovomitopsis membrillensis D'Arcy, Ann.
Missouri Bot. Gard. 67: 1036. 1980. TYPE: Panama. Darién:
Upper Río Membrillo, 100--800 m, Duke 10872
Shrubs or trees to ca. 8 m; resin clear. Leaf blades narrowly
elliptic, 16--25 X 4--6 cm, the apex acuminate, the base acute and
slightly decurrent, drying reddish to grayish tan and with numerous,
closely spaced, often-branched, resin lines; lateral veins 1--2.5 cm
apart, ca. 10 pairs, fading towards the margin and very indistinctly
loop-connected; petiole 1--3.5 cm. Inflorescence ca. 10 cm, erect,
glabrous; flower buds ca. 5--6 mm, the outer 2 sepals ± equal
and as long as or shorter than the others; stamens ca. 40, anthers of
stamens and staminodia slightly longer than wide, ca. 1 mm. Fruits
ovoid-pyriform, ca. 3 cm, strongly alate but the wings thin, pink;
calyx ± persistent in fruit.
Distribution. Very wet primary forest, 100--850 m. Known
for sure from just five collections, only from Panama, low to middle
elevations of Colón province and extreme eastern Darién
province. This species must certainly also occur in nearby Colombia,
but no such specimens have been seen.
This Panamanian endemic has narrowly elliptic and often falcate,
coriaceous leaves, with very visible resin canals. The lateral leaf
veins are without prominent intersecondaries, and the fruits are
basically pyriform and winged. Although the winged fruits are very
characteristic of this species, they are not obvious in young
pistillate material nor even more mature, dried material.
Specimens examined. PANAMA. Colón:
Santa Rita Ridge Road, 600--700 m, Apr. 1981 (fl), Sytsma
et al. 4176 (MO); 250--400 m, May 1982 (fl),
Knapp & Schmalzel 5264 (MO).
Darién: Parque Nacional del Darién, between
Río Topalisa y Río Pucuro, 750--850 m, Oct. 1987 (fl,
fr), Hammel et al. 16282 (MO); Upper
Río Membrillo, 100--800 m, Apr. 1967 (fr), Duke
10863 (MO); Apr. 1967 (fr), Duke 10901 (MO).
Chrysochlamys nicaraguensis (Oersted, Planchon &
Triana) Hemsley, Biol. Cent.-Amer., Bot. 1: 87. 1879.
Tovomitopsis nicaraguensis Oersted, Planchon &
Triana, in Planchon & Triana, Ann. Sci. Nat. Bot., ser. 4. 14:
266. 1860. Tovomita nicaraguensis (Oersted, Planchon
& Triana) L. O. Williams, Trop. Woods 111: 18. 1959. TYPE:
Nicaragua. [Río San Juan]: along the Río San Juan, [ca.
10--50 m], Oersted 2  (holotype, C).
Chrysochlamys standleyana L. O. Williams, Trop.
Woods 111: 16. 1959, new name for C. pauciflora
Standley, Ceiba 3: 214. 1953 non C. pauciflora
Steyermark 1952. Tovomitopsis standleyana (L. O.
Williams) D'Arcy, Ann. Missouri Bot. Gard. 67: 1040. 1980. TYPE:
Panama. Bocas del Toro: Chiriquí Lagoon, Wedel
1019 (holotype, F; isotype, MO).
Shrubs or trees, 3--8 m; resin clear. Leaf blades elliptic to
obovate-oblong, 14--25(--27) X 6--10(--12.5) cm, the apex acute to
rounded and apiculate, the base acute to acuminate, drying gray or
tannish green, more or less glossy; lateral veins 1.5--3 cm apart,
(5--)6--8(--10) pairs, fading to the margin; petiole 1.5--4 cm.
Inflorescence 3--19 cm, usually erect; flower buds 5--7 mm, outer
pair of sepals ± equal to each other and shorter than the inner;
stamens 35--40, anthers of stamens and staminodia about as wide as
long, < 1 mm. Fruits obovoid to globose, 1.5--3 cm, greenish pink
to red; calyx deciduous in fruit.
Distribution. Very wet forest of the Caribbean lowlands,
from near sea level to 600(--1200) m; Nicaragua, Costa Rica, Panama.
Chrysochlamys nicaraguensis, often erroneously
interpreted as a wide-ranging species (cf. discussion under C.
eclipes), is still only known for certain from the very wet
Caribbean lowlands of Nicaragua, Costa Rica, and adjacent Panama. It
is distinctive for its very glossy and distantly veined (2--3 cm)
leaves almost totally lacking visible intersecondaries and resin
canals. For discussion of authorship and type specimen citation of
this species, see under C. glauca.
Representative specimens. NICARAGUA. Río San
Juan: Campamento La Lupe, ca. 15.5 km al Noreste del poblado El
Castillo, 100 m, Dec. 1991 (fr), Grijalva 5633 (CR).
COSTA RICA. Alajuela: Cantón de San Ramón,
camino al Volcán Muerto, 1100--1200 m, May 1993 (fl),
F. Araya et al. 294 (CR, INB, MO).
Heredia: Cantón de Sarapiquí, La Selva Research
Station, 100 m, Feb. 1981 (fr), Folsom 8775 (CR, DUKE).
Limón: Cerro Coronel, 60--100 m, Mar. 1987 (fl),
Stevens et al. 24851 (CR, MO); Parque
Nacional Tortuguero, ca. 3 km al Sur del pueblo, 3 m, Nov. 1988 (fr),
Robles 2175 (CR, MO); Cantón de Talamanca, Aguas
arriba de Río Banano, Fila Asunción, 400 m, Feb. 1995
(fr), Rodríguez 580 (CR, INB, MO); hills between
headwaters of Quebrada Mata de Limón and upper branches of
Quebrada Tigre (Sixaola region), 25--30 m, Nov. 1984 (fr),
Grayum et al. 4468 (CR, MO). PANAMA.
Bocas del Toro: Fish Creek Mts., vicinity of Chiriquí
Lagoon, Apr. 1941 (fl), von Wedel 2300(MO);
Peninsula Valiente, on E side of Lagoon of Chiriquí, 25 m,
Aug. 1987 (fl), McPherson 11453 (MO). Veraguas:
valley of Río Dos Bocas along road between Escuela
Agrícola Alto Piedra and Calovebora, 450--550 m, Aug 31 (fl),
Croat 27581 (MO). Coclé: Alto Calvario, N
of El Copé, 700--900 m, May 1977 (fr), Folsom
3274 (MO); region N of El Valle, 1000 m, Sep. 1946 (fl),
Allen 3683 (MO). Panamá: Cerro Campana,
800 m, Aug. 1982 (fl), Hamilton & D'Arcy
662 (MO); Finca El Indio at headwaters of Río Indio,
slopes of Cerro Jefe, 750 m, Nov. 1979 (fl), Antonio
2425 (MO). Colón: ca. 2--3 m up Río
Guanche, 10--20 m, Jan. 1973 (fr), Kennedy &
Foster 2193 (MO). San Blas: Cerro Habú,
500--850 m, Dec. 1980 (fr), Sytsma et al.
Chrysochlamys psychotriifolia (Oersted, Planchon &
Triana) Hemsley, Biol. Cent.-Amer., Bot. 1: 87. 1879.
Tovomitopsis psychotriifolia Oersted, Planchon &
Triana, in Planchon & Triana, Ann. Sci. Nat. Bot., Ser. 4. 14:
263. 1860. TYPE: Costa Rica. [Cartago]: Turrialba, [ca. 650 m],
Oersted 4  (holotype, C).
Tovomitopsis costaricana Oersted, Planchon &
Triana, in Planchon & Triana, Ann. Sci. Nat. Bot., Ser. 4. 14:
265. 1860. Chrysochlamys costaricana (Oersted, Planchon
& Triana) Hemsley, Biol. Cent.-Amer., Bot. 1: 87. 1879. TYPE:
Costa Rica. [Cartago]: Turrialba [ca. 650 m], Oersted 1
 (holotype, C).
Tovomitopsis faucis D'Arcy, Ann. Missouri Bot. Gard.
67: 1031. 1980. Syn. nov. TYPE: Panama. Chiriquí: forests
around El Boquete, 1000--1300 m, Pittier 3039
Shrubs or trees, 3--10 m; resin clear or slightly creamy. Leaf
blades oblong-elliptic, 12--21 X 3--8(--9) cm, the apex acute, the
base acute to acuminate, dull, drying gray-green; lateral veins
mostly 1(--1.5) cm apart, 10--15 distinct pairs, fading to the margin
or somewhat loop-connected towards the leaf apex; petiole 1--2 cm.
Inflorescence 6--16 cm, erect, minutely pubescent or glabrous; flower
buds ca. 5--7 mm, outer 2 sepals ± equal to each other and
shorter than the inner; stamens 15--20, anthers of stamens and
staminodia about as long as wide, < 1 mm; stigmas ± reflexed.
Fruits oblong and often apiculate, 3--5.5 cm, greenish yellow-pink to
red; calyx deciduous in fruit.
Distribution. Very wet montane forest, 600--1500 m; Costa
Rica and Panama.
Chrysochlamys psychotriifolia, well known in Costa
Rica, is a species with numerous, rather close, lateral veins, few
(ca. 20) monomorphic stamens, and usually large (3--5 cm),
obovate-mammillate fruits. Tovomitopsis costaricana,
T. faucis, and what D'Arcy (1980) called T.
membranacea (Planchon & Triana) D'Arcy were based on very
scant material that is, nevertheless, now seen to fall easily within
the variation of C. psychotriifolia. For discussion of
authorship and type specimen citation of this species, see under
C. glauca. The South American Chrysochlamys
membranacea Planchon & Triana, by its type and numerous
more recent collections is clearly a different species, belonging to
Planchon and Triana's section Heterandra, with dimorphic
stamens in the staminate flowers and the staminodia connate into a
ring or collar in the pistillate.
Representative specimens. COSTA RICA. Guanacaste:
Parque Nacional Rincón de La Vieja, 950--1100 m, Nov. 1987
(fr), Herrera 1446 (CR); Cantón de
Tilarán, 1--2 km W of Lago Cote, 800--900 m, Aug. 1993 (fl),
Haber & Zuchowski 11626 (CR, MO).
Puntarenas: Reserva Monteverde, Vert. Pacific, 1520--1580 m,
Sep. 1976 (fl), Dryer 757 (CR). Alajuela: La
Palma de San Ramón, 1250 m, Nov. 1925 (fl), Brenes
4600 (385) (CR). Heredia: Horquetas de
Sarapiquí, Reserva Rara Avis, 700 m, July 1995 (fr),
Martén 1008 (CR). San José:
Braulio Carrillo National Park, vicinity of Estación Carrillo,
600 m, July 1985 (fr), Hammel & Trainer
14276 (CR, MO). Cartago: Cantón de Turrialba,
Valle del Reventazón, Grano de Oro, 1100 m, Sep. 1993 (fl),
Campos & Campos 185 (CR, INB, MO).
PANAMA. Bocas del Toro: road to Chiriquí Grande N of
Fortuna Dam, 650--700 m, June 1987 (fl), McPherson
11130 (MO). Chiriquí: in forest along road to
La Fortuna, 1400 m, Feb. 1979 (fl), Hammel 6232 (MO);
ridge road at Cerro Colorado, 1450--1750 m, Aug. 1977 (fl, fr),
Folsom et al. 4832 (MO); Veraguas:
above Santa Fe beyond Escuela Agrícola Interamericana, Apr.
1976 (fr), Croat 34216 (MO).
Chrysochlamys silvicola (Hammel) Hammel, comb. nov.
Basionyym: Tovomitopsis silvicola Hammel, Selbyana 9:
113. 1986. TYPE: Costa Rica. Heredia: La Selva Biological Station
[ca. 100 m], Hammel 9404 (holotype, DUKE; isotypes, CR,
Shrubs or trees 3--6 m; resin white. Leaf blades oblong-elliptic
(13--)16--22 X 4--7(--8) cm, the apex acute to acuminate, the base
acute, dull both surfaces, drying reddish tan; major lateral veins
0.5--2 cm apart, ca. 10--16 pairs, difficult to distinguish from the
intersecondary veins, fading to the margin where indistinctly
loop-connected; petiole 1--2.5 cm. Inflorescence 5--10 cm, ±
pendent, minutely pubescent or glabrous; flower buds ca. 4--5 mm,
outer 2 sepals often markedly unequal with the larger one ±
overtopping the bud; stamens 20--30, anthers of stamens and
staminodia about as long as wide, < 1 mm; stigmas ± reflexed.
Fruits ovoid to globose, 1.5--2.5 cm, red throughout or whitish at
base; calyx ± persistent in fruit.
Distribution. Very wet, primary lowland forest, Caribbean
slope, from near sea level to 700(--900) m; Costa Rica and Panama.
This species, particularly as to its type, can be characterized by
its primary lateral veins and intersecondaries being almost equally
salient. Nevertheless, a few collections from Costa Rica included
here (Heredia, Cantón de Sarapiquí, Rara Avis, 400--700
m, Martén 948 (CR); G. Vargas
1650 (CR); O. Vargas 2 (CR); O.
Vargas 4 (CR)) have fewer and less prominent
intersecondaries. Material from Panama appears to grade imperceptibly
into an extreme with the intersecondaries very obscure. All the
material has dull, yellowish to reddish tan-drying leaves, the outer
pair of sepals often very unequal, the petals usually without obvious
resin canals, and stamens about 20--30. At least in Costa Rica, the
resin of the twigs is bright white, rather than clear or tardily
creamy as in most other species in Central America. For its reddish
tan-drying leaves, this species sometimes resembles the more northern
C. guatemaltecana, which has more stamens and shiny
leaves with fewer, more distant veins.
Representative specimens. COSTA RICA. Alajuela:
Cantón de San Carlos, (near) Boca Tapada, Finca La Monalisa,
100 m, May 1995 (fl), Rodríguez &
Estrada 702 (CR, INB, MO). Heredia: between
Río Peje & Río Sardinalito, Caribbean slope of
Volcán Barva, 700--750 m, Apr. 1986 (fr), Grayum
6695 (MO); Finca La Selva, Puerto Viejo de Sarapiquí
(ca. 100 m), July 1979 (fl, fr), Grayum 1902 (CR,
DUKE). San José: Parque Nacional Braulio Carrillo,
Carillo, 600 m, May 1982 (fl), Gómez-Laurito
8440 (CR). Limón: Ref. Nac. Barra del Colorado,
forests and pastures between Chirripocito and Río Sardina, 12
m, Apr. 1990 (fr), Grayum 9794 (CR, MO, F); La Aurora,
Guácima, ca. 5 km al E del pueblo, 100 m, Aug. 1988 (fl),
Robles et al. 2087 (CR, MO, F);
Cantón de Talamanca, Alto Urén, 700--900 m, Sep. 1994
(fr), Gallardo & Lépiz 313
(CR, INB, MO, F). PANAMA. Veraguas: NW of Santa Fe, 1 km from
Escuela Agrícola Alto de Piedra, Feb. 1975 (fl), Mori
& Kallunki 4794 (MO), 900--1200 m, May 1981
(fr), Sytsma & Andersson 4640 (MO).
Coclé: El Copé, 1200 m, June 1979 (fr),
Antonio 1189 (MO); N of Penonomé on road to
Coclesito, 100--170 m, Feb. 1978 (fl), Hammel 1681
(MO). Panamá: between Cerro Azul and Cerro Jefe, July
1969 (fl), Foster 1157 (MO); El Llano-Cartí
Road, 400--450 m, Jan. 1974 (fl), Nee &
Dressler 9352 (MO), 350--400 m, Feb. 1973 (fr),
Kennedy 2543 (MO). San Blas: Río Diablo y
vecindad de Duque Sui, 80--110 m, July 1992 (fl), Herrera
et al. 1174 (MO). Darién: vicinity
of upper gold mining camp of Tyler Kittredge on headwaters of
Río Tuquesa, Aug. 1974 (fr), Croat 27209 (MO).
Chrysochlamys skutchii Hammel, sp. nov. TYPE: Costa Rica.
San José: basin of El General, 675--900 m, Mar. 1940 (fl),
Alexander Skutch 4759
(holotype, CR; isotype, MO). Figure 1.
Figure 1. Chrysochlamys skutchii Hammel (A, B,
Skutch 4759; C, Aguilar 1771). --A.
Flowering branch. --B. Staminate flower. --C. Fruit.
Altitudo altior C. allenii affinis a qua differt foliis majoribus,
tenuioribus, par extimus sepala inaequalia, calyx ad maturiatem
fructus persistens et inflorescentia glabra.
Trees or shrubs 2--10 m; resin clear to creamy (red?). Leaf blades
elliptic, (17--)21--35 X 6--12 cm, membranaceous, the apex and base
acute to acuminate; lateral veins ca. 1--2 cm apart, ca. 10--17
pairs, mostly reaching to the margin or the apical veins looping into
a ± distinct, submarginal vein; petiole 2--4 cm. Inflorescence
7--15 cm, erect, glabrous; flower buds ca. 5 mm, outer pair of sepals
very unequal, one < 1/2 the length of the bud, the other ±
equal to the bud; petals membranaceous, lacking distinctly
contrasting resin canals; stamens ca. 50, varying in length
centripetally, 1--4 mm, the anthers nearly as wide as long, ca. 0.3 X
0.4 mm; stigmas sessile, spreading or slightly elevated. Fruits
± pyriform or obovate, 2.5--4 cm, pale green-pink to red; seeds
ca. 1 cm, the aril orange; calyx ± persistent in fruit.
Distribution. This species is known only from a few
collections from relatively low 100--750(--900) m elevations in wet
forest on the Pacific slope of southern (and eastern) Costa Rica,
from the regions of Acosta (Fila Bustamante), Valle del General, the
Osa Peninsula and vicinity.
Flowering or sterile collections of this species may be difficult
to separate from Chrysochlamys grandifolia, a species
most readily distinguished by its strongly ribbed fruits. The two
grow sypatrically and have been confused on Costa Rica's Osa
Peninsula. However, the leaves of C. grandifolia
usually have a more distinct submarginal vein, it has fewer stamens,
and the anthers are often 2--3 times longer than wide rather than
about as long as wide. In many details, e.g., fruit shape and color,
number and size of stamens, leaf coloration, C.
skutchii comes much closer to C. allenii.
Chrysochlamys skutchii is distinguished from C.
allenii by its larger, thinner leaves, glabrous inflorescence,
unequal outer sepal pair, persistent sepals in fruit, and lower
elevation habitat. The epithet is in honor of the species' first
collector, one of Costa Rica's primier natural historians, Alexander
Additional specimens. COSTA RICA. Puntarenas:
Cantón de Golfito, La Gamba, "Bosque de los Austriacos," 300
m, Dec. 1993 (fl), Huber & Weissenhofer
68 (MO), (fl), Huber & Weissenhofer
90 (CR), (st.), Huber & Weissenhofer
94 (CR); Cantón de Osa, Parque Nacional Corcovado,
Cerro Brujo, 600 m, Jan. 1991 (fl), Castro 268 (MO);
Cerro Rincón, 745 m, Aug. 1990 (fr), Morales et
al. 20 (CR, MO); Cerro Mueller, 150--650 m, May 1988
(fr), Kernan & Phillips 474 (CR, MO);
Estación Los Patos, 250 m, May 1995 (fr), Moraga
191 (CR, MO), 500 m, July 1988 (fr), Kernan
& Phillips 662 (CR); Los Planes, La Gloria,
170 m, Feb. 1991 (fr), Induni 266 (CR, INB, MO), 100 m,
Feb. 1991 (fr), R. González 32 (CR);
Rincón, Fila Casa Loma, 50--400 m, July 1990 (fr),
Chacón 941 (CR, MO); Camino de Altura, 100--300
m, Aug. 1967 (fr), Raven 21512 (MO); Aguabuena, 3.5 km
W of Rincón, 450 m, Nov. 1992 (fl), Thomsen 188
(CR); Rancho Quemado, 500 m, Nov. 1991 (fl), Marín
et al. 256 (MO), 300--400 m, May 1988 (fr),
Hammel et al. 16898 (CR, MO), 200--350 m,
Apr. 1993 (fr), Aguilar 1771 (CR, INB, MO).
Chrysochlamys tenuifolia Cuatrecasas, Revista Acad. Colomb.
Ci. Exact. 29: 59. 1950. TYPE: Ecuador. Río San Miguel o
Sucumbíos, 250 m, Cuatrecasas 10880 (holotype,
F; isotype, COL).
Shrubs or small trees to 2.5--7 m, most twigs with paired
bud-scale scars just above the node; resin yellow to orange. Leaf
blades elliptic, 8--14(--24) X 3--4.5(--9) cm, the apex acuminate,
the base acute to obtuse, drying gray-green; major lateral veins ca.
3(--20) mm apart, ca. (12--)16 prominent pairs but with several
nearly equally prominent intersecondaries, fading to the margin or
looping to an obscure submarginal vein; petiole 0.6--1.2 cm.
Inflorescence appearing axillary or terminal, 1--3 cm, erect,
glabrous; flower buds 2--3 mm, the outer 2 sepals 1/3 the length of
the bud; stamens ca. 60, apparently without filaments, anthers of
stamens and staminodia broader than long, < 0.5 mm, surmounting a
resiniferous capitulum . Fruits ellipsoid-falcate, bean-shaped, red,
1.5--2.5 cm, 3-valved but usually just 1-seeded, the seed black or
brown with an orange aril; sepals persistent in fruit.
Distribution. Very wet forest; 300--1400 m. Eastern Panama,
Because of its apparently axillary inflorescences, bud-scale
scars, and resin-producing staminate flowers, this species's position
in Chrysochlamys is somewhat dubious. It has many close and
very prominent lateral veins, and small, essentially terminal
inflorescences that are often left in an apparently axillary position
(only one per node) by the dominant axillary vegetative shoot. It
also sometimes has unambiguously terminal inflorescences, and has an
androecium with the stamens more or less connivent into an apparently
resiniferous capitulum, much as in the resin-producing flowers of
some species of Clusia. The staminodia of the pistillate
flowers are connate into a ring or collar around the base of the
ovary as in Chrysochlamys membranacea. The small,
narrowly ellipsoid-falcate, capsulate fruits mostly have only three
sessile stigmas, and usually one seed with a markedly cellular aril.
In leaf venation and bud scales it is very similar to
Symphonia, but for numerous reasons would be even more out of
place there than in Chrysochlamys. The species must certainly
belong to the Tovomita group, but further study is in order.
Although D'Arcy (1980) created the combination Tovomitopsis
micrantha (Engler) D'Arcy for this material, the Peruvian
Chrysochlamys micrantha Engler, by its original
description and type photo, bears it no resemblance.
Representative specimens. PANAMA. San Blas: forest
SE of Puerto Obaldia, Aug. 1971 (fr), Croat 16754 (MO);
mountains above Puerto Obaldia, Aug. 1971 (fr), Gentry
1487 (MO). Darién: Cerro Mali on Colombian
border, 1400--1500 m, Jan. 1975 (fr), Gentry &
Mori 13806 (MO); Parque Nacional del Darién,
ridge between Río Topalisa and Río Pucuro, ca. 17 km E
of Pucuro, 600-850 m, Oct. 1987 (fr), de Nevers
et al. 8343 (MO), (fl), Hammel et
al. 16379 (MO), 700--1400 m, Oct. 1987 (fl, fr),
Hammel et al. 16357 (MO); Río Tacarcuna,
vicinity of old Tacarcuna village, 580 m, Jan. 1975 (fl),
Gentry & Mori 13586 (MO).
Chrysochlamys tenuis Hammel, sp. nov. TYPE: Costa Rica.
Limón: Cordillera de Talamanca, Fila Matama, 09°47'18"N,
83°08'45"W, 1350 m, Apr. 1989 (staminate fl), G.
Herrera & A. Chacón
2673 (holotype, CR; isotypes, MO +
2). Figure 2.
Figure 2. Chrysochlamys tenuis Hammel (A--C,
Herrera & Chacón 2673, D,
Berrocal et al. 74). A. --Flowering
branch. --B. Flower bud. --C. Staminate flower. --D. Fruit.
Chrysochlamys glauca affinis a qua differet foliis
angustioribus et plerumque in sicco ferruginis vice glaucorum,
venatione magis inconspicuo, inflorescentia tenuiore.
Shrubs or small trees 2--5 m; resin clear to tardily creamy.
Petiole 0.5--2 cm. Leaf blades narrowly elliptic, 5--13 X
1--2.5(--3.5) cm, the apex acuminate, the base narrowly acute and
shortly decurrent, both surfaces dull, drying reddish tan to
gray-green; lateral veins ca. 0.5 cm apart, 6--8 pairs, very
indistinct, often difficult to distinguish from intersecondaries,
fading to the margin or looping to form an obscure submarginal vein;
resin canals usually very obscure, ± parallel to the midrib or
at least somewhere crossing the lateral veins. Inflorescence
paniculate, very lax to slightly congested, ± globose, ca.
2--5(--8) cm long, 3- to ca. 30-flowered, the rachis very slender,
often deflexed or pendulous, glabrous; flower buds 3--4 mm long,
usually conspicuously striate with resin canals; outer 2 sepals
± equal and shorter than the bud, sometimes very narrowly
triangular, rarely (in the type and one nearby paratype,
Herrera 2495) the third and fourth sepals
long-acuminate with the filiform tips overtopping the rest of the bud
by ca. 2 mm; petals membranaceous; stamens ca. 20, free, 2--2.5 mm
long, the anthers ± globose, ca. 0.3 mm long; stigmas 5. Fruits
narrowly obvoid to pyriform, usually rostrate, 2--3 cm long reddish,
often tinged with white or pale yellow, 1--3-seeded; seeds with an
orange aril; calyx deciduous in fruit.
Distribution. This species is known from scattered
localities in very wet forest on the Atlantic slope of Costa Rica and
Panama, and from a few collections from wet forest on the Burica
Peninsula, shared by both countries along their Pacific border.
Although most material is from below 800 m, the type is from nearly
The type and all other Costa Rican, Atlantic slope collections, as
well as one Panamanian collection from Bocas del Toro
(Kirkbride & Duke 621), one from
Veraguas (McPherson 10724), and two from the Río
Blanco de Norte area of the Atlantic lowlands of Coclé, have
particularly long, narrow leaves without obvious intersecondaries and
relatively few-flowered, very lax inflorescences with the outer
sepals slightly to very narrowly triangular. The Burica Peninsula
outliers and all other collections from Panama tend to have somewhat
broader and shorter leaves with more obvious intersecondaries,
multi-flowered, more congested inflorescences, and broader outer
Chrysochlamys tenuis is similar to C.
glauca in its dull, few-veined leaves and small flower buds.
The Río Guanche and Cerro Campana populations cited here were
discussed by D'Arcy (1980, p. 1034) under that species as "suggestive
of a distinct variety or even species." Rather than further blurring
what then seemed only suggestive, more recent collections,
particularly those from Atlantic Costa Rica, strengthen the case for
recognizing this entity as a distinct species. Other than the
narrower and usually reddish tan vs. gray-green-drying leaves, more
obscure venation and smaller, more delicate inflorescences, C.
tenuis is distinguished from C. glauca by the
resin canals of the leaves, which, when visible, are more or less
parallel to the midrib or at least somewhere crossing the lateral
veins, rather than parallel to the lateral veins, as in C.
glauca and most other species. The narrowly obovoid fruits of
C. tenuis are usually only 1- or 2-seeded, whereas the
more or less globose or broadly obovoid fruits of C.
glauca are normally 5-seeded. Mori et al.
3950 recorded the seeds as "steel blue." The new species also
superficially resembles the Panamanian C. angustifolia,
which, however, is an epipetric rheophyte with milky resin, stouter
inflorescence axes, and much larger flower buds.
Original field notes for the type of this species indicated four
duplicates. These were earlier distributed as either
Tovomitopsis angustifolia or T. glauca,
and their whereabouts, except as indicated above, are uncertain.
Additional specimens. COSTA RICA.
Limón: Cordillera de Talamanca, Fila Matama, Cerro
Muchilla, 850 m, Apr. 1989 (fr), Robles &
Chacón 2678 (CR, MO); Reserva Biol. Hitoy
Cerere, Cerro Bitárkara, 800 m, Feb. 1989 (fl bud),
Herrera & Solís 2495 (CR, MO);
Río Tercero, 600 m, May 1985 (fr), Berrocal et
al. 74 (CR); Suretka, 200 m, July 1982 (fr),
Gómez-Laurito 8839 (CR). Puntarenas:
Burica Peninsula, Punta Burica, M. M. Chavarría
306 et al. (CR, INB). PANAMA.
Chiriquí: Burica Peninsula, 8--11 mi. W of Puerto
Armuelles, 200--500 m, Feb. 1973 (fl, fr), Croat 21995
(MO), Mar. 1973 (fl, fr), Croat 22472 (MO); La Fortuna
hydroelectric project, 1200 m, Mar. 1978 (fl, fr), Hammel
2181 (MO). Bocas del Toro: Quebrada Huron on Cerro
Bonyic, 170--400 m, Apr. 1968 (fr), Kirkbride &
Duke 621 (MO); Chiriquicito-Calderas trail, Apr. 1968
(fr), Kirkbride & Duke 765 (MO).
Veraguas: Escuela Agrícola Alto de Piedra, road to
Calovébora, 450--550 m, Aug. 1974 (fr), Croat
27665 (MO), Nov. 1974 (fr), Mori &
Kallunki 3224 (MO), Dec. 1974 (fr), Mori
et al. 3950 (MO); slopes of Cerro Tute, 600--750
m, Mar. 1987 (fl), McPherson 10724 (MO).
Coclé: above El Copé, ca. 700--900 m, Apr. 1977
(fr), Folsom & Robinson 2371 (MO),
Apr. 1977 (fr), Folsom & Kauke 2652
(MO), Oct. 1977 (fr), Folsom et al. 5707
(MO), Nov. 1977 (fr), Folsom & Collins
6516 (MO), Jan. 1978 (fr), Hammel 953 (MO), Apr.
1978 (fr), Hammel 2432 (MO), June 1978 (fr),
Hammel 3587 (MO), Jan. 1989 (fr), McPherson
13627 (MO); Atlantic slope from El Copé to Río
Blanco del Norte, ca. 150 m, Feb. 1980 (fl, fr), Antonio
3653 (MO), 350 m, Feb. 1982 (fl), Knapp 3675
(MO); near El Valle, ca. 700--1000 m, Aug. 1967 (fr), Duke
& Dwyer 13936 (MO), Jan. 1972 (fr),
Gentry & Dwyer 3654 (MO), Dec. 1972
(fr), Gentry 6848 (MO), Sep. 1978 (fr), Hammel
4794 (MO), May 1967 (fl), Lewis et al.
1769 (MO), 1785 (MO), Nov. 1974 (fr), Mori
& Kallunki 2960 (MO), June 1975 (fl),
Mori et al. 6565 (MO).
Panamá: Cerro Campana, ca. 900 m, May 1971 (fl),
Croat 14681 (MO), 14753 (MO), Sep. 1982 (fr),
D'Arcy 15071 (MO), Mar. 1969 (fl), Porter
et al. 4259 (MO); Cerro Trinidad, 800-1000 m,
Oct. 1946 (fr), Allen 3773 (MO); El Llano-Cartí
Road, Feb. 1977 (fl, fr), Foster 2770 (MO).
Colón: Santa Rita Ridge, Jan. 1968 (fl), Dwyer
8564 (MO); Río Escandaloso near Cerro Bruja, 10--200 m,
Apr. 1978 (fl), Hammel 2672 (MO), 2713 (MO),
Mar. 1982 (fl), Knapp & Kress 4283
(MO), (fr), Knapp et al. 4461 (MO);
Río Guanche near Portobelo, ca. 0--100 m, May 1984 (fl, fr),
Churchill 5119 (MO), Oct. 1978 (fr), Dec. 1973 (fl),
Gentry 8764 (MO), Oct. 1978 (fr), Hammel
et al. 4932 (MO), July 1971 (fl), Kennedy
1174 (MO), Jan. 1973 (fr), Kennedy &
Foster 2132 (MO), Aug. 1981 (fr), Knapp
1024 (MO), Mar. 1975 (fr), Mori &
Kallunki 5194 (MO), Mar. 1975 (fl), 5212 (MO);
June 1975 (fl), Mori et al. 6443 (MO),
Aug. 1975 (fr), Mori & Witherspoon
7955 (MO), Dec. 1973 (fr), Nee &
Gentry 8719 (MO), July 1976 (fr), Sullivan
155 (MO), Oct. 1980 (fr), Sytsma 1645 (MO).
Although the strictly South American Balboa does not occur
within the geographic focus of this paper, taxonomic clarity and
logical consistency prescribe its mention here as an additional
synonym of Chrysochlamys. Planchon and Triana described the
genus and its single species, B. membranaceae based
only on staminate material and placed it in their subtribe
"Euclusieae" rather than Tovomiteae. Examination of type material and
more recent collections, of both staminate and pistillate plants
(e.g., Boyle & Boyle 3303 (MO),
Croat 73004 (MO), Gentry et al.
14655 (MO), Hammel & Trainer
15831 (MO), Hoover 1287 (MO), Hoover et
al. 3136 (MO), Romero-Castañeda
2739 (COL), Rubio & Quelal
1339 (MO)) support the view that B. membranacea
is a species of Chrysochlamys:
Chrysochlamys balboa Hammel, nomen nov. Basionym:
Balboa membranacea Planchon & Triana, Ann. Sci.
Nat. Bot., ser. 4. 14:252. 1860, non C. membranacea
Planchon & Triana, 1860. TYPE: Colombia. Between Tuquerres and
Barbacoa, 1176 m, Apr. 1853, Triana s.n. (holotype, P; isotype
Chrysochlamys clusiifolia subsp. panamensis
Maguire, Phytologia 36: 395. 1977. TYPE: Panama. Darién: Cerro
Mai, Serranía del Darién, Gentry &
Mori 13769 (holotype, MO) = Dystovomita
paniculata (J. D. Smith) Hammel, Ann. Missouri Bot. Gard. 76:
D'Arcy (1980) already showed this to belong in his newly elevated
Tovomitopsis membranifolia (Standley) D'Arcy, Ann.
Missouri Bot. Gard. 67: 1035. 1980. Tovomita
membranifolia Standley, Field Mus. Nat. Hist. Bot. Ser. 22:
160. 1940. TYPE: Panama. Darién: Caná-Cuasí
trail, Chepigana District, Terry & Terry
1478 (holotype, F; isotype, MO) = Tovomita
stylosa Hemsley, Biol. Cent.-amer. 1: 88. 1879.
Hammel (1986) already showed this to be a synonym of
Tovomita croatii Maguire, Phytologia 36: 404. 1977.
Tovomitopsis croatii (Maguire) D'Arcy, Ann. Missouri
Bot. Gard. 67: 1031. 1980. TYPE: Panama. Veraguas: road to
Calovébora Croat 14268A (holotype, MO).
Shrubs 1--2 m, resin clear. Leaf blades obovate, widest above the
middle, 20--60 X 7--25 cm, the apex acuminate, the base narrowing to
an acute or cordulate base; lateral veins ca. 1--2 cm apart, ca.
16--22 pairs, looping to form a distinct submarginal vein 2--5 mm
from the margin; petiole lacking. Inflorescence 10--26 cm, usually
pendulous, glabrous or puberulent; flower buds ca. 6 mm, the outer
sepals often overtopping the inner; stamens 25--30, anthers of both
stamens and staminodia about twice as long as wide, ca. 1.5 mm;
stigmas concave and spreading. Fruits pyriform, green to pink, ca. 2
cm; seeds ca. 1 cm; calyx ± persistent in fruit.
Distribution. Very wet, mostly lowland forest,
30--350(--1500) m; Costa Rica, Panama, Ecuador.
This rare, somewhat problematic species might appear to walk the
generic borderline between Chrysochlamys and Tovomita.
The outer pair of sepals, although thinly membranaceous, exceed the
bud as in Tovomita, yet styles are lacking and the fruits
apparently do not have the dark maroon placenta and inner fruit wall
typical of Tovomita. Nevertheless, two Costa Rican collections
(Grayum 3374, Grayum & Schatz
5276) clearly indicate the presence of side shoots with
clustered leaves. With present knowledge, placement of this species
in Tovomita, particularly because of its branching pattern, is
more justifiable than placement in Chrysochlamys. As a
species, it is very distinctive, not only for the nearly sessile and
sometimes very large leaves, but also for the numerous (16--22)
lateral veins that are loop-connected to a distinct submarginal vein.
Probably because of the species' rarity, its short stature and large
leaves, the typical Tovomita branching pattern of T.
croatii has gone unobserved and unrecorded.
Representative specimens. COSTA RICA. Limón:
Reserva Biol. Hitoy Cerere, 90--200 m, July 1985 (fr),
Grayum & Hammel 5763 (CR, M0);
Cantón de Talamanca, Sukut, 350 m, July 1989 (fr),
Herrera 3168 (CR, MO); Finca Anai, W of Mata de
Limón (Sixaola region), 30 m, May 1985 (fl), Grayum
& Schatz 5276 (CR, MO). PANAMA.
Chiriquí: Fortuna Dam Area, 1100 m, May 1984 (fl),
Chuchill 5243 (MO). Coclé: El Valle,
Cerro Gaital, 900--1000 m, July 1968 (fl, fr), Dwyer
& Correa 8913 (MO). ECUADOR.
Pichincha: Quito a Sto. Domingo de los Colorados, km 94,
1300--1500 m, Dec. 1986 (fr), Zak 1529 (MO).
Acknowledgments. This work was supported, in part, by funds
from the National Science Foundation through a grant (DEB-9300814) to
the author and co-PI M. H. Grayum for the Manual to the Plants of
Costa Rica. I thank Mike Grayum for his always useful comments on an
early draft of the manuscript and Sylvia Troyo for the line drawings
of the two new species.
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