Three new species of Celastraceae from Costa Rica, one disjunct from Mexico
Barry E. Hammel
Missouri Botanical Garden
P. O. Box 299
St. Louis, Missouri, 63166
This paper was reviewed and accepted in November of 1996 for publication in Novon, Vol. 7 (2). In preparation for and in keeping with the theme of electronic distribution of the
Manual to the Plants of Costa Rica, the mss. was made available on two Web sites (INBio and MO) by 14 April 1997.
Abstract. Three species of Celastraceae, each represented by many collections from Costa Rica, are compared to near relatives and found to be new: Gymnosporia haberiana Hammel, disjunct from Mexico and Costa Rica, Maytenus recondita Hammel, of Costa Rica and Panama, and Crossopetalum enervium Hammel, of Costa Rica . The South American Gymnosporia magnifolia (Loesener) Lundell is here confirmed as a synonym of G. urbaniana (Loesener) Liesner, and Crossopetalum eucymosum (Loesener & Pitter) Lundell is placed in synonymy under C. parviflorum (Hemsley) Lundell. Keys are provided for all accepted neotropical species of Gymnosporia, and for all Costa Rican species of Crossopetalum and Maytenus.
For nearly twenty years collections of two large, cloud forest trees
in the family Celastraceae have been accumulating, primarily from the
Monteverde region of Costa Rica. Much of this material has been
distributed to various herbaria under tentative or approximate names.
More recent exploration along the central cordillera has extended the
range, within Costa Rica, of these two species and yielded many
collections of a third, shrubby species in the genus
Crossopetalum, also known from the Monteverde region. The two
trees are known almost exclusively from mid-elevations on the Pacific
slopes of the northern mountains, while the shrub extends nearly the
entire length of the country and is known from numerous collections
on both slopes.
Although one of the trees has always been known to be a Maytenus presumably related to M. schippii Lundell, generic placement of the other has been more problematic. This species, while having alternate leaves and capsulate fruits with arillate seeds, as in Maytenus, has greener drying, entire leaves, more open and branched inflorescences with 4-merous flowers,
and several seeded, 2--4 lobed fruits. In these respects the Costa Rican (and Mexican) tree can be distinguished from other alternate-leaved Celastraceae and coincides perfectly with two South American species recently described in or transferred to the genus Gymnosporia (Lundell, 1985; Liesner, 1993).
Gymnosporia haberiana Hammel,
sp. nov. TYPE: COSTA RICA. Guanacaste: Cordillera de
Tilarán, La Cruz de Abangares, 1400 m, 15 Jul 1985 (fl.)
Haber & Bello 2034
(holotype INB; isotypes, BM,
C, CAS, COL, CR, DUKE, F, FLAS, GH, HNMN, K, MEXU, MICH, MO, NY,
P, PMA, TEX, US, USJ, W, WIS). Figure 1.
A G. gentryi foliorum nervis lateralibus minus prominentis
et latiore divergentibus, ovario et capsula tantum lobato non
anguloso, a G. urbaniana floribus majoribus, disco manifesto, ab
ambobus sepalis latioribus quam longioribus.
Figure 1. Gymnosporia haberiana. A. fruiting shoot; B.
staminate flowers; C. pistillate flowers; D. fruit. (A, D from
Haber & Zuchowski 11048; B from Hammel 17661; C from Haber 9303).
Dioecious trees (3--) 5--30 m, the inner
bark bright yellow; twigs terete; leaves
and twigs drying pale to yellowish green, glabrous. Leaves alternate,
7-12 (--14) x 3--6 (--8) cm, elliptic, the apex acute to acuminate,
the base acute to rounded, the margin entire and often curled under,
all leaves often folded and curved along the midrib; main lateral
veins mostly 4 or 5; petioles 5-10 mm; stipules small, deciduous.
Inflorescences 2 or 3-branched, ca. 10--20-flowered axillary cymes,
the rachis minutely and sparsely farinose-puberulent, the pedicels
2-4 mm. Flowers 4-merous, ca 4.5--7 mm wide, greenish; sepals ca. 0.4
x 1 mm, broadly rounded; petals imbricate, ca. 0.4 x 1 mm, spatulate
or ligulate ca. 0.4 x 1 mm; stamens exceeding the style in staminate
(pollen-bearing) plants, shorter than the style and without pollen in
pistillate plants, anthers ± cordate, versatile; ovary ±
globose, confluent with the conically raised disk, 2-celled with 2
ovules/cell. Fruits ca. 1 x 1--1.5 cm, globose or often wider than
long and 2--4 lobed, (1) 2--4-seeded; seeds dark brown with white
Additional specimens examined. COSTA RICA. Alajuela:
Cordillera de Guanacaste, P. N. Rincón de la Vieja,
10°47'50"N 85°18'19"W, 1500 m, 6 Jul 1991 (fl.), Rivera
1423 (CR, F, INB, MICH, MO, USJ, W). Guanacaste:
Cordillera de Guanacaste, P. N. Guanacaste, Volcán Cacao, ca.
10°55'45"N 85°28'15"W, 1100-1400 m, 25 Nov 1989 (fr.),
Chávez 7 (CR, F, MO); Dec 1990 (fr.), 478 (CR,
F, INB, MO); Jul 1989 (fl.), Hammel 17661 (BM, CAS,
COL, CR, F, INB, MEXU, MO, TEX, US); 9 Feb 1995 (fr.),
Quirós 48 (CR, INB, MO); P. N. Rincón de
La Vieja, sendero al Volcán, 10°45'50"N 85°18'50"W,
820 m, Jan 1991 (fr.), Rivera 974 (CAS, CR, F, INB, MO).
Puntarenas: Cordillera de Tilarán, Monteverde Reserve
and vicinity, ca. 10°20'N 84°50'W, 1200--1550 m, 4 Mar 1990
(fr.), Bello 2110 (F, INB, MO, MV); 28 Jul 1977 (fl.),
Dryer 1589 (CR, MO); 4 Apr 1985 (fr.), Haber & Bello
1458 (BM, CAS, CR, F, LL, MEXU, MO, US); 12 Jul 1985 (fl.),
Haber & Bello 1984 (BM, COL, CR, F, LL, MEXU, MICH,
MO, P, UPS, W); 29 Nov 1985 (fr.), Haber ex Bello 3591 (BM,
CR, LL, MO); 17 Dec 1985 (fr.), Haber ex Bello 3935 (CAS, CR,
DUKE, F, LL, MEXU, MO); 22 Jul 1986 (fl.), Haber ex Clagget
5738 (CR, F, GH, LL, MO); 27 Feb 1987 (fr.), Haber & Bello
6728 (INB, LL, MO); Feb 1988 (fr.), Haber & Bello 8213
(F, INB, LL, MO, US); 20 Jul 1989 (fl.), Haber 9303 (CR, F,
INB, MEXU, MO, TEX), 26 Mar 1990 (fr.), Haber 9815 (CR, INB,
MO, US); 30 Jul 1991 (fl.), Haber & Zuchowski 10789 (CR,
INB, MO), 12 Mar 1992 (fr.), Haber & Zuchowski 11048 (CR,
F, INB, MO); 27 Jul 1977 (fl.), Hartshorn 1901 (CR, F);
25 Jan 1984 (fr.), Pennington 11432 (CR, MO). MEXICO.
Veracruz: Estación Biológica Los Tuxtlas,
18°35'N 95°05'W, 300 m; no date (st.), Calzada
79 (F); 29 Nov 1974 (fr.), Cedillo 436 (MO); 4
Feb 1986 (fl.), Cedillo 3518 (MO); 24 Jun 1984 (fl.),
Ibarra & Sinaca 1787 (MO); 15 Nov 1984 (fr.), Ibarra et
al 2124 (MO); 20 Jan 1985 (fr.?) Ibarra & Sinaca 2240,
2241 (MO); 4 Jul 1985 (fl.), Sinaca 122 (MO).
Distribution and biogeography. Gymnosporia haberiana is disjunct between one site at about 300 m elevation in Veracruz, México, and several wet-forest, mid-elevation sites in NW Costa Rica (Fig. 2). However, its nearest
relatives, as discussed below, are from low- and mid-elevation South America, a common pattern for tropical Mexican and Central American taxa (eg., Hammel, 1986 and references therein). Nevertheless, G. haberiana is not known from southern Costa Rica or from Panama. This disjunct distribution
between Mexico (or northern Central America) and Costa Rica, then skipping Panama to nearest relatives in South America, is not unique (eg. Ziziphus chloroxylon (L.) Oliv.), and if not an artifact of collecting, may itself reflect the complex geological history of the isthmian region.
Figure 2. Distribution of four species of Celastraceae in Costa Rica
(hollow symbols mark type localities; 500 m contour is
Relationships. The closest relatives of Gymnosporia
haberiana are its two South American congeners. In 1985
Lundell transferred Maytenus magnifolia Loesener to
Gymnosporia and described a second neotropical species,
G. gentryi Lundell, thus reinstating for the New World
an essentially African genus that had come to be considered
congeneric with Maytenus. In contrast to species of
Maytenus, most of which (at least in the neotropics) have
small fasciculate inflorescences, 5-merous flowers, 1 or rarely
2-seeded fruits, and often toothed, gray- or tan-drying leaves, the
New World Gymnosporia have relatively large, open cymose
inflorescences, 4-merous flowers, usually 2--4-seeded fruits, and
entire, greenish-drying leaves (see also Hou,
1955). Study of specimens and bibliographic
material at MO supports Liesner's conjecture
(1993) that G. urbaniana (the
older name) and G. magnifolia are conspecific:
Gymnosporia urbaniana (Loesener) Liesner Monogr. Syst. Bot. Missouri Bot. Gard. 45: 1254. 1993: Rhacoma urbaniana Loesener Repert. Spec. Nov. Regni Veg.Vol. 1(11): 162. 1905. TYPE: Perú. Weberbauer 1875 (photo MO).
Syn. nov. Gymnosporia magnifolia (Loesener) Lundell Phytologia 57: 314. 1985: Maytenus magnifolia Loesener, Vernh. Bot. Vereins Prov. Brandenburg 48: 176. 1906 (1907). TYPE: Brazil. Amazonas: Flusse Juruá Miry, Ule 5721 (photo MO).
The new species differs from G. gentryi of Colombia
and Ecuador and G. urbaniana of Amazonian Brazil, Peru
and Bolivia by its generally smaller leaves with less conspicuous
lateral veins, usually fewer-flowered inflorescences, and sepals that
are much broader than long. Gymnosporia urbaniana has
smaller flowers with almost no nectar disk and is found mostly in
lowland Amazonia. Gymnosporia gentryi, from 1200--2800
m in the northern Andes, has flowers nearly as large as those of
G. haberiana, but the ovary (and fruit) is distinctly
4-angled and its leaves have very prominent (below) lateral veins
that depart from the midrib at a much narrower angle. By these
observations, G. haberiana and G. gentryi
appear to be sister species. The three neotropical species of
Gymnosporia can be separated according to the following
- 1 Flowers mostly < 4 mm wide, sepals ca. 0.5 mm wide and equally
long; filaments widening toward base, but nectar disk inconspicuous;
smallest twigs green; plants mostly Amazonian, from below 800
- 1' Flowers > 5 mm wide, sepals ca. 1 mm wide, much shorter than
wide or equally long; filaments scarcely widening toward base, nectar
disk conspicuous; smallest twigs red or green; plants Andean or north
of Panama, from above 1200 m.
- 2 Leaves mostly 18--20 cm long, the lateral veins very
prominent, arching strongly forward at < 45 degrees; smallest
twigs with reddish exfoliating epidermis; ovary (and fruits)
distinctly 4-angled; sepals as wide as long or slightly wider;
Ecuador, Peru, Bolivia...G. gentryi
- 2' Leaves < 14 cm long, the lateral veins not prominent,
arching outward mostly at 50 degrees or more; smallest twigs
green; ovary globose, the fruits often 4-lobed but not angled;
sepals much wider than long; Mexico and Costa Rica...G.
Mexican material of this species has sometimes been tentatively
identified as Maytenus grisea Lundell, but examination
of the Guatemalan type of that species (Contreras 6944,
LL!) shows G. haberiana to be quite distinct. Although
the two species are superficially similar because of their somewhat
inflated and relatively thin-walled fruits, in contrast to the open
cymes and lobed fruits of G. haberiana, the
inflorescences of M. grisea are only pedicellate
clusters on a very short or obsolete peduncle, the fruits are not
lobed, and the leaves are toothed and dry gray rather than green.
Unexpectedly, this study has revealed that a rarely collected Costa
Rican plant from hills near Palmar Norte (Allen 6327, F
[as "Maytenus pallidifolius Standl. & L. O. Williams ined."];
Bohlke 20, F; Hammel et al. 20296, INB; Poveda & Hoet 3050, CR) is certainly
conspecific with M. grisea. Thus, M.
grisea, known only from the type in Guatemala and now from
southern Costa Rica (Fig. 2), is similar to
G. haberiana in being disjunct within the region.
Etymology and history. This new species is dedicated to my
friend and colleague, Dr. William Haber, long-time student and
professor of the rich flora of Monteverde, Costa Rica, locality of
the type and most other collections cited here. Bill's keen
observation and field notes brought to my attention the dioecious
nature of this species. The earliest known collection of this species
is from Mexico (Cedillo 436, Nov 1974). In Costa Rica
it was apparently first collected in Jul 1977 (Dryer
Maytenus recondita Hammel, sp. nov. TYPE: COSTA RICA. Puntarenas: Cordillera de Tilarán,
Monteverde community, 10°18'N 84°48'W, 1350 m elev., 11 Jul
1989 (fl.), Haber & Zuchowski 9286 (holotype, INB; isotypes, CR, F, MICH, MO, US). Figure 3.
A M. schippii atque M. guyanense inflorescentiis
cymoso-paniculatis, foliis in sicco viridi-griseis aut fuscis,
praesentia altitudinis majoris differet.
Figure 3. Maytenus recondita. A. flowering shoot; B.
pistillate flowers; C. fruits. (A, B from Hartshorn 1895; C from Haber & Zuchowski 8737).
Dioecious trees 5--25 m; twigs usually zig-zag, terete. Leaves
alternate, (5--) 7--11 x 2--5 cm, elliptic, the apex acuminate, the
base decurrent, shallowly toothed in the distal 1/2 or without teeth,
with 4--7 indistinct lateral veins, these looping and not reaching
the margin, glabrous, drying dark, gray-brown above, gray-green
below; petioles 2--5 mm; stipules small, triangular, ±
persistent. Inflorescences small, usually 1--4 (--8)-flowered,
axillary cymose, panicles, the rachis 1--7 mm, the pedicels 1--2 mm,
occasionally proliferating on ± leafless branchlets up to 4 cm
long. Flowers 5-merous, ca. 3--4 mm across, pale green; sepals ca.
1/2 or less the length of the petals, rounded, ± erose-margined;
petals 1 x 1 mm at base, more or less triangular; stamens yellow,
born on the margin of a flat disc ca. 1.5 mm wide, the filaments ca.
0.5--0.6 mm. Capsules bivalvate mostly 1 or 2 (to 6) per axis,
obovoid, 10--15 mm, bright orange, usually 1- or occasionally
2-seeded; seeds with a white aril.
Additional specimens examined. COSTA RICA. Alajuela:
Cordillera de Guanacaste, entre Río Celeste y cabeceras del
Río Chimurria 10°43'15"N 85°00'20"W, 700--800 m,
Jul. (fl. & fr.), Herrera 2008 (F, INB, MO); Cordillera de
Tilarán, altos del Río Caño Negro, 10°21'N
84°48'W, 1300 m, 17 Sep 1989 (fl.), Bello 1376 (INB, MO);
(Puntarenas, by error, on labels) Bosque Eterno De Los Niños,
10°23'N 84°42'W, 1100 m, fl. Apr., Bello et al 2156
(CAS, CR, F, INB, MO, W); 10°24'N 84°39'W, 800--900 m,
17 Jul 1993 (fr.), Haber et al 11544 (CAS, CR, F, INB, MICH,
MO, W); Parque Nacional Juan Castro Blanco, 10°15'30"N
84°15'30", 1200 m, 26 Jun 1993 (fl., fr.), Jiménez
1327 (COL, CR, F, INB, MO); Parque Nacional Arenal, 10°25'N
84°43'W, 1200 m, 18 Sep 1990 (fl.), Bello 2403 (INB);
Peñas Blancas River Valley, 10°20'N 84°50'W, 1250 m,
12 Oct 1985 (fr.), Haber ex Bello 3100 (CR, F, MO); Reserva
Biológica de San Ramón, 1017'N 8436'W, 1000 m, 16 Feb
1994 (fl), Herrera 6894 (CR, USJ). Guanacaste:
Cordillera de Guanacaste, Parque Nacional Rincón de La Vieja,
cabeceras de Quebrada Rancho Grande, 10°46'N 85°49'W,
1350--1400 m, 2 Dec 1987 (fr.), Herrera 1474 (MO); Cordillera
de Tilarán, 3 km N Santa Elena, ca. 1020'N 8450'W, 1500 m,
near 20 Dec 1985 (fr.), Haber ex Bello 3824, (CR, F, MO),
3831, (CR, F, MO), 3863 (CR, MO); 5 km N Santa Elena on
road to Las Nubes, 10°22'N 84°49'W, 1400 m, 10 Nov 1988
(fr.), Haber & Zuchowski 8737 (INB, MO); Río
Negro de Tilarán, 1500 m, 4 Oct 1985 (fl., fr.), Haber
& Bello 2989 (CR, F, MO); 10°21'N 84°49'W, 1400 m,
27 May 1987 (fl.), Haber & Bello 7125 (F, MO).
Heredia: Cordillera Central, vicinity of Vara Blanca, 1710 m,
Apr 1938 (fl.), Skutch 3765 (MO, US); Volcán
Barva, 1700--2000 m, 24 May 1972 (fl., fr.), Stone 3261 (CR,
F, MO). Limón: Cordillera de Talamanca, Z. P. Barbilla,
frente a confluencia entre Río Caño Seco y Río
Dantas, 10N 8326'W, 150--350 m, 3 Nov 1988 (fl.), Herrera 2278
(F, MO). Puntarenas: Cordillera de Tilarán, Monteverde
Reserve and vicinity, ca. 10°20'N 84°50'W, 1000--1700 m
elev., 25 May 1989 (fl.), Bello 922 (MICH, INB, MO,
TEX, US); 21 Nov 1991 (fl.), Bello 4164 (INB); 17 to 20 Mar
1973 (fr.), Burger & J. Gentry 8601 (CR, F); 30 Oct 1976
(fl., fr.), Dryer 933 (CR, F, MO); 15 Dec 1976 (fr.), Dryer
1082 (CR); 13 Jan 1977 (fl., fr.), Dryer 1125 (CR,
F, MO); Gentry & Haber 48761 (F, MO); 20 Mar 1976 (fl.,
fr.), Gómez-Laurito et al. 1402 (USJ); 5 Aug 1978 (fl.,
fr.), Haber 162 (CR); Haber 446 (F, MO), 692 (F,
MO), 693 (F, MO); 20 Oct 1985 (fl.), Haber ex Bello
3108 (CR, MO); 20 Oct 1985 (fr.), Haber ex Bello 3117 (CR,
F, MO); 10 Dec 1985 (fr.), Haber ex Bello 3673 (CR, MO); 16
Jan 1986 (fl., fr.), Haber ex Bello 4266 (CR, F, MO); 2
May 1974 (fl.), Hartshorn 1470 (CR 4 sheets, F, MO); 19 Feb
1976, (fr.), Hartshorn 1820 (F); 26 Jul 1977 (fl.),
Hartshorn 1895 (CR 3 sheets, F, MO); 19 Aug 1995 (fl.,
fr.), Penneys & Zuchowski 683 (CR, F, INB, MO, TEX); 8 Aug
1975 (fl., fr.), Poveda 1110 (CR, F, MO, USJ); Cordillera de
Tilarán, Ojo de Agua, Río Aranjuez, 10°17'N
84°46'W, 1550 m, 14 Nov 1987 Haber & Bello 7738 (F,
MO); San Luis, Cerro Amapola, 10°16'33"N 84°47'45", 1100 m,
23 Nov 1993 (fr.), Fuentes 583 (INB, MO);
Península de Osa, Quebrada Agua Buena, 0842'40"N 8331'00"W,
500 m, 13 Mar 1996 (fl.), Aguilar 4524 (CR, INB, F,
MO). PANAMA: Chiriquí: Bajo Mono-Robalo trail ca.
1500--2100 m, 27 Jul 1947 (st.), Allen 4844 (F).
Distribution. Except for one sterile collection from western
Panama, Maytenus recondita is restricted to wet forests
of central Costa Rica (Fig. 2), primarily at
700--1700 m elevation on the Pacific slope from the Cordillera de
Guanacaste to the Cordillera Central, with one low-elevation,
Atlantic-slope collection from the Río Barbilla region,
Cordillera de Talamanca, (Herrera 2278) and another
isolated collection from the Osa Peninsula (Aguilar
Characterization. This species is recognized by the acuminate
and decurrent, gray-green drying leaves and the paniculate
inflorescence of reduced or aborted cymes. It is generally a larger
tree as compared to M. segoviarum Standl. &
Steyerm., with which it is sympatric in Costa Rica. It has been
identified in the past as M. schippii Lundell "vel. sp.
aff.," but differs from M. schippii by its shorter
petioles, branched, pedunculate (rather than fasciculate)
inflorescences, and higher-elevation habitat. Another similar
species, currently identified in Costa Rica as M.
guyanensis Klotzsch ex Reissek, differs by its much larger,
usually darker drying leaves, fasciculate inflorescences, and
lower-elevation habitat. The five species of Maytenus
recognized for Costa Rica can be distinguished by the following
1 Leaves distinctly serrulate with gland-tipped teeth; base of
peduncle densely covered with ferrugineous, laciniate bracteoles;
plants from > 2000 m...M. woodsonii Lundell
- 1' Leaves entire or indistinctly toothed; ferrugineous bracteoles
sparse or lacking; plants from 0--1700 m.
- 2 Largest leaves 12-20 cm; plants from below 500 m elev.
- 3 Leaves and fruits drying dark gray to brownish-black;
infl. ± racemose or fasciculate from a short peduncle;
frs. obovoid, not inflated; plants mostly from P. N. Manuel
Antonio and Osa Peninsula, near sea level...M.
- 3' Leaves and frs. drying pallid gray-green, mottled; infl.
fasciculate, mostly without peduncle; frs. globose inflated;
Palmar Norte, 450 m...M. grisea.
- 2' Largest leaves 11 cm or <; plants from 40--1700 m
- 4 Leaf apex narrowly acuminate, sometimes mucronate, the
base markedly decurrent, the leaves drying dark gray-green to
greenish tan; infl. paniculate-cymose; frs. short-pedicellate
from an obvious rachis; plants mostly of cloud forest, (150--)
700--1700 m...M. recondita
- 4' Leaf apex acute to rounded, often retuse, the base
acute, the leaves drying light gray to tan; infls. fasciculate;
frs.arising from a knob; plants mostly of drier slopes below
cloud forest, 40--1100 m...M. segoviarum
Anthers on flowering material of M. recondita also
bearing young fruits appear (on close examination) not to bear
pollen. Although the relative length of stamens and style does not
differ strikingly in staminate and pistillate flowers, as in the
above-discussed species of Gymnosporia, M.
recondita must also be dioecious.
A series of collections from Costa Rica (U.
Chavarría 179, INB; Chávez
248, INB; Ezpinoza 1186, INB; G.
Herrera 850, INB; A. Rodríguez
188, INB) is here specifically excluded from the concept of
M. recondita. This entity shares the elevational range,
inflorescence type, and leaf color of M. segoviarum,
but has a leaf shape more similar to that of M.
recondita. It is not yet decided whether these collections
represent a sixth species for Costa Rica, hybrids between M.
recondita and M. segoviarum, or simply variation
within M. segoviarum.
Etymology and history. The epithet recondita ("hidden,
unpretentious") was chosen for this species because its primary
distinction is its reduced but nevertheless branched inflorescence.
In addition, although the species has been well-collected and
distributed to various herbaria for many years, it is unremarkable
and has remained undescribed. This species was first collected in Apr
1938 (Skutch 3765).
Crossopetalum enervium Hammel, sp. nov. TYPE: COSTA RICA.
Heredia: Llanura de San Carlos, S base of Cerros Sardinal,
Chilamate de Sarapiquí, 10°27'N 84°04'W, 70--100 m,
2 Jun 1985 (fl., fr.), Grayum & Jacobs 5351
(holotype CR; isotypes F, MO,
LL + 3, dist. from MO as C. eucymosum). Figure 4.
Differt ab species affinibus foliorum nervis lateralibus
inconspicuis, apprime a C. standleyi foliis latioribus sed
brevioribus, inflorescentiis ut videtur simplicioribus, a C. gomezii
foliis multo minoribus.
Figure 4. Crossopetalum enervium. A. shoot with flowers
and immature fruit; B. flowers; C. fruits. (A, B from Gamboa
48; C from Bello 2232).
Shrubs or small trees 1--5 (--8) m; twigs sharply quadrate-ribbed,
glabrous. Leaves opposite, 6--10 x 1.8--4 cm, elliptic, the apex
acute to acuminate, apiculate, the base acute and decurrent,
distantly but sharply serrate, glabrous; main lateral veins 4--6,
loop-connected well below the margin, very indistinct above and
below; petioles ca. 3--5 mm; stipules minute, caducous extensions of
the twig angles. Inflorescences ca. 1--1.5 cm long, mostly 3--7
(--25) flowered, usually only 1-branched, axillary to extra-axillary
cymes, glabrous or rarely puberulent; bracts and bracteoles small but
± foliaceous, gland-tipped; peduncles 7--13 mm, the pedicels
1.5--3 mm. Flowers small, 4-merous; sepals ca. 1/2 the size of the
petals, rounded; petals ca. 1--1.5 mm, ± orbicular, the usually
wine-red margins slightly crenate; stamens yellow, borne from just
inside the margin of a flat, circular or quadrate disc; filaments ca.
0.1--0.2 mm. Fruits drupaceous, 0.7--1.4 x 0.6--0.7 cm, obovoid, red
to black; seeds tuberculate.
Additional specimens examined. COSTA RICA. Alajuela:
Cordillera de Tilarán, Monteverde Reserve, Peñas
Blancas river valley, ca. 10°20'N 84°45'W, 800-1100 m, 22
Feb 1989 (fl.), Bello & Cruz 724 (INB, MV);
29 Nov 1986 (fl., fr.), Haber ex Bello 6480
(CR); Reserva Forestal de San Ramón, 800--1000 m, 4 May 1985
(fr.), E. Rojas s. n. (USJ); Llanura de San Carlos,
Pital, Yucatán 10°34'40"N 84°11'00"W, 100 m, 4 Oct
1994 (fr.), Estrada 246 (INB); Llanura de Guatuso, 3 km
NW of Florencia 10°23'N 84°28'W, 250 m, 28 Dec 1993 (fl.,
fr.), Haber & Guindon 11759 (CR, F, INB, LL,
MO, US); 8 km NE of Villa Quesada, near Artezalea, 550 m, 17 Feb 1966
(fr.), Molina et al. 17271 (CR, F); Pueblo Nuevo, 1100
m, 15 Apr 1939 (fl., fr.), A. Smith 1902 (F).
Guanacaste: Cordillera de Guanacaste, P. N. Guanacaste,
Volcán Cacao, ca. 10°55'45"N 85°28'15", 1100 m, 11
Feb 1995 (fl.), Alfaro 114 (INB); 2 Jun 1990 (fr.),
Bello 2232 (INB); 11 Apr 1991 (fl., fr.),
Chávez 543 (CR, INB); 14 Jul 1991 (fr.),
Chávez 585 (CR, INB); 8 Feb 1995 (fl., fr.),
B. Gamboa 48 (CR, INB, MO); 10 Feb 1995 (fl.,
fr.), Lobo 50 (INB); 3 Jun 1990 (fr.), Maass
14 (INB); 2 Jun 1990 (fr.), Obando 15 (CR,); P.
N. Rincón de La Vieja, 800 m, 27 Jan 1983 (fl.),
Garwood et al 726 (F). Heredia:
Llanura de San Carlos, near Tirimbina E of the Río
Sarapiquí, 10°24'N 84°07'W, 150--250 m, 12--15 Aug
1971 (fr.), Burger & Burger 8053 (CR, F).
Puntarenas: Cordillera de Tilarán, Monteverde area,
Hoge Middle, 13 Mar. 1979 (fl.), Koptur SK-102 (CR);
Cordillera de Talamanca, P. I. La Amistad, Estación Altamira,
09°02'10"N 83°01'20"W, 1350 m, 14 Apr 1995 (fl.),
Angulo 164 (INB); Las Alturas and vicinity, 1700 m, 26
Aug 1974 (fr.), Maas 1484 (CR, F).
Distribution. Crossopetalum enervium is
apparently endemic to Costa Rica, where it is known from both the
northern and southern parts of the country in wet to very wet forest
from ca. 100 to 1700 m elevation (Fig. 2). A
study of shrub and treelet species at Monteverde (Koptur et al.,
1988), includes phenological data about this
species (as C. eucymosum), vouchered by the above cited
Koptur collection (Haber, pers. comm); fruits take about 3 months to
mature and then may remain on the plant up to 5 months. Most
collections are from the northern half of the country in the
Cordilleras de Guanacaste and Tilarán and from the Caribbean
lowlands to the E. A few collections with slightly more prominent
venation, approaching that of C. standleyi, come from
an outlying population in the extreme S of the country on the Pacific
slopes of the Cordillera de Talamanca very close to Panama. Although
not yet known from Panama, this new species must certainly occur
Characterization and relationships. This species is
distinctive for the very faint (when dry) lateral veins and the
delicate, subumbellate inflorescences. In leaf size and certain
aspects of the inflorescence and fruit, this species is most like
C. standleyi (Lundell) Lundell (basionym Myginda
standleyi; [Standley 68938, istotype F!]), from
which it differs by its wider and slightly shorter leaves, markedly
obscure venation, and the subumbellate cymes with relatively long
primary peduncles. Both C. standleyi and C.
enervium have obovate fruits and short anther filaments.
Crossopetalum enervium is also similar to the type of
C. riparium (Lundell 1476, F!) in its
smallish leaves and delicate, glabrous inflorescences. The
inflorescences of the latter species, however, are more distinctly
branched, and its leaves dry brownish (rather than gray-green), and
are less serrate and more prominently nerved than those of C.
enervium. The stamens are nearly sessile on the type of
C. riparium. Both C. riparium and
C. standleyi (originally described from Guatemala) are
restricted to northern Central American. That the close relatives of
this new species are from N of Costa Rica is to be expected; the
genus is basically Central American and West Indian, with only one
species reported from South America (Gentry,
1993). One collection of C.
enervium, Chávez 543, has slightly
Among Costa Rican Crossopetalum, the new species is most
similar to C. gomezii Lundell, which differs by having
much larger leaves with very prominent venation. It has been confused
with the most common Costa Rican species, formerly known as C.
eucymosum (Loesener & Pittier) Lundell, here considered a
synonym of the older C. parviflorum (Hemsley)
C. parviflorum (Hemsley) Lundell Wrightia3(1): 8.
1961: Euonymus parviflorus Hemsley, Diag. Pl. Nov. Mex.
p. 6. 1878. TYPE: Nicaragua, Chontales, R. Tate
292 (photo! US).
Syn. nov. Crossopetalum eucymosum
(Loesener & Pittier) Lundell Wrightia3(1): 7. 1961:
Myginda eucymosa Loesener & Pittier, Contr. US.
Natl. Herb. 12:175. 1909, Pl. 18. TYPE: Guatemala. Alta Verapaz:
Cahabon River, Pittier 239 (US!).
Crossopetalum parviflorum has much larger, more
open, pubescent inflorescences, larger flowers with longer stamens,
and leaves that dry brownish instead of gray and have more obvious
secondary venation. The five Costa Rican species of
Crossopetalum can be distinguished by the following key:
- 1 Leaves pubescent, mostly less than 1.5 cm wide, acute; plants
from near sea level on the dry Pacific coast...C.
uragoga (Jacquin) Kuntze
- 1' Leaves glabrous, mostly more than 2 cm wide, acuminate or acute;
plants from higher or at least wetter habitats.
- 2 Fruits cylindrical; leaves acute at apex, conspicuously
toothed; flowers nearly sessile at the end of a short
peduncle...C. tonduzii (Loesener) Lundell
- 2' Fruits obovoid; leaves acuminate at apex; flowers both
pedicellate and pedunculate.
- 3 Inflorescence pubescent, openly cymose-branched; largest
leaves mostly 10(--13) cm long, inconspicuously toothed, drying
brownish; filaments nearly 1/2 length of petals...C.
- 3' Inflorescence glabrous (very rarely pubescent in
C. enervium), little branched, appearing almost
umbellate; leaves larger or smaller, drying gray; filaments
much less than 1/2 length of petals.
- 4 Leaves inconspicuously toothed, the largest at least
(12--)15 cm long; lateral nerves conspicuous
- 4' Leaves conspicuously toothed, the largest mostly 6
(--7) cm long; lateral nerves very indistinct
Etymology and history. The epithet "enervium" is chosen in
reference to the inconspicuous lateral venation of the leaves in dry
specimens. Here again "reconditum" was a serious contender for the
honors; this rarely collected species has actually been in
collections for quite long, the earliest known gathering being that
of Austin Smith 1902, in 1939. That collection
and others, including the type, were distributed as
Crossopetalum eucymosum, wherein the new species has,
until now, remained hidden.
Acknowledgements. I thank Michael Grayum for his ever
careful reading of an early draft of the mss, two anonymous reviewers
for further refinements, Silvia Troyo for the line drawings, and Henk
van der Werff for fine tuning the latin descriptions. The photographs
of Gymnosporia haberiana and Maytenus recondita were
supplied by William Haber. Field work resulting in many of the
collections here cited was supported by various National Geographic
Society grants to William Haber and to Michael Grayum and by National
Science Foundation (NSF) grant BSR-8700068 to M. H. Grayum and the
author. Publication was supported by NSF grant DEB-9300814 to the
author and M. H. Grayum.
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