Flora of Panama. Part VIII. Family 155.

Ebenaceae
Author(s): Robert E. Woodson, Jr., Robert W. Schery, F. White
Source: Annals of the Missouri Botanical Garden, Vol. 65, No. 1 (1978), pp. 145-154
Published by: Missouri Botanical Garden Press
Stable URL: http://www.jstor.org/stable/2395355 .
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 tJLUIiA UtJ FAIN AIVLA-

BY ROBERT E. WOODSON, JR. AND ROBERT W. SCHERY

AND COLLABORATORS

Part VIII

FAMILY 155. EBENACEAE

F. WHITE2

froma woodyrhizome,withoutmilkysap;
Trees,shrubsor rarelysuffruticose
heartwoodsometimesblack;indumentumusuallyof simple,unicellular,setose,
strigose,or sericeoushairs,but multicellularglandular,unicellularbibrachiate,
and multicellularpeltate hairs sometimespresent. Leaves usually alternate,rarely
opposite, exstipulate,entire, the epidermalcells papillose in some species, mostly
in those with a glaucous lower epidermis. Flowers actinomorphic,hypogynous,
unisexual, but often with well-developed staminodes or pistillodes, solitary,
fasciculate, cymose or rarely in false racemes or panicles; calyx gamosepalous,
entire to deeply lobed, sometimesclosed in bud, otherwisewith open or imbricate
aestivation,always persistentin fruit and often accrescent;corolla gamopetalous
with contorted aestivation, shortly to deeply lobed, rotate, urceolate, salver-
shaped or sausage shaped, often constricted at the throat; stamens 2-100+,
epipetalousor borne on the receptacle, the filaments usually short and flattened,
the antherslanceolateor sagittate,often 2 or more arisingfrom a single filament,
usually dehiscing by lateral slits, rarely by apical pores, the connective well
developed, often larger than the anthers, often apiculate or rostrate; ovary
syncarpous,2-5 or more carpellary,the carpels 2-ovulate, usually completely or
partly divided by a false septum, rarely unilocular,the ovules pendulous from
the apex of the locule, the styles distinct or basallyconnate,less often more or less
completely united, the stigmas usually large, fleshy and irregularlylobed. Fruit
a berry, rarely tardily dehiscent; seeds large; endospermabundant,hard, some-
times ruminate;embryo relatively small, straight or slightly curved, the radicle
well developed, the cotyledons flat, oval, foliaceous, usually emergent, either
persistent and photosynthetic or fugaceous and non photosynthetic.
The Ebenaceae is a family with 2 genera; Euclea Murr.with a dozen species
is confinedto Africa,and the pantropicalDiospyroshas ca. 80 species in America,
94 on the African mainland,ca. 100 in Madagascar,and ca. 200 in tropical Asia
and the Pacific. It shows only feeble penetrationsinto temperatelatitudes and is
poorly represented on tropical mountains.

1 Assisted by National Science Foundation Grant DEB 77-04300 (W. G. D'Arcy,

principal investigator).
2Departments of Botany and Forestry, University of Oxford, South Parks Road, Oxford
OXI 3RB, Great Britain.
ANN. Missoum BOT. GARI. 65: 145-154. 1978.

0026-6493/78/0145-0154/$01. 15/0
146 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 65

Among Panamanian families the Ebenaceae appears to be most closely re-

lated to the Sapotaceae, from which it differs inter alia in the articulated, uni-
sexual flowers, biovulate carpels with apical placentation, bitegmic ovules, the
small hilum of the seed, and the absence of latex.
The number of floral parts in Ebenaceae is very variable, both between and
within species. The unripe fruit is very astringent but the ripe fruit of most
species is edible, though in a few species it is used as a fish poison. The whole
fruit is usually the unit of dispersal but it seems that in D. salicifolia individual
seeds with adherent pericarp are removed by birds or small mammals while the
fruit is still attached to the tree.
The Central American species D. digyna has been planted in many parts
of the tropics for its edible fruit. Diospyros blancoi A. DC., a native of the Philip-
pines which also has an edible fruit has been planted in a few places in Panama.
It is much better known under the illegitimate name D. discolor Willd.
Our knowledge of Panamanian Ebenaceae is very incomplete and more col-
lecting is needed. No species is represented in herbaria by complete material.
In this account 4 species are provisionally given specific names. All four are
widespread and variable, and the Panamanian representatives are somewhat
different from their relatives elsewhere. Further work may lead to the recognition
of distinct subspecies. The descriptions of the flowers of D. dig yna, D. salicifolia,
and D. artanthifolia (in part) are based on non-Panamanian material.

Literature:
Hiern, W. P. 1873. A monograph of Ebenaceae. Trans. Cambridge Philos.
Soc. 12: 27-300.
Howard, R. A. 1961. The correct names for "Diospyros ebenaster." J. Arnold
Arb. 42: 430-435.

1. DIOSPYROS
Diospyros L., Sp. P1. 1057. 1753. TYPE: D. lotus L.
Characters as given for family.

KEY TO FRUITING SPECIMENS

a. Fruitsbornein a lax, richlybranchedinfructescence; calyx 0.5 cm long with (4-)

5(-6) acute, deltateteeth-1. D. artanthifolia
aa. Fruitssolitaryor in 2-3-floweredfascicles.
b. Calyxcup-shaped;leavesto 9 cm long and 4 cm wide-4. D. salicifolia
bb. Calyxsaucer-shaped; leavesmuch larger.
c. Fruitsdistinctlypedicellate,at least 2.5 cm in diameter.
d. Fruitsca. 2.5 cm in diameter;calyxlobed to the middle-- 3. D. inconstans
dd. Fruitsat least 4 cm in diameter;calyx shallowlylobed- 2. D. digyna
cc. Fruitssubsessile,to 1.5 cm in diameter -5. D. sp. I

KEY TO FLOWERING SPECIMENS'

a. Corollalobes 4-5.
b. Corollalobes much longerthan the tube-1. D. artanthifolia

'The flowersof D. sp. I. are unknown.

1978] WHITE-FLORA OF PANAMA (Family 155. Ebenaceae) 147

FIGURE 1. Diospyros artanthifolia Mart-A. Leaves and fruits X7/10). [After Nee
7862 (MO).]-B. Male inflorescence (X21/;,). -C. Male flower (X3). [After Croat 14657
(MO).]

bb. Corolla lobes much shorter than the tube- 2. D. digyna

aa. Corolla lobes 3.
c. Leaves to 9 cm long and 4 cm wide, the tertiary nerves and veins indistinct
----------------------------------------------------------------------- 4. D. salicifolia
cc. Leaves much larger, the tertiary nerves and veins forming a close and conspic-
uous reticulum -3. D. inconstans

1. Diospyros artanthifolia Mart., Fl. Bras. 7: 7. 1856. TYPE: Maynas, Peru,

Poeppig 2266 (BR, holotype; L, OXF, NV, isotypes; FHO, US, photos).-
FIG. 1.

Tree 4-15 m tall; branches long, slender, subverticillate; branchlets setulose-

tomentellous. Leaves chartaceous, drying brownish black above, paler brown
beneath; lamina to 11 cm long and 4 cm wide, lanceolate, broadest just above
148 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 65

FIGURE 2. Diospyros digyna Jacq. Leaf and fruit (X). [After Gentry 6227 (MO).]

the base, the apex acuminate, the base cuneate, the margin revolute, the lower
surface strigose-puberulous and rough to the touch, the lateral nerves in 7-10
pairs, slightly impressed above, prominent beneath, the intercostal and other
nerves forming a subprominent lax reticulum beneath. Male flowers in 7-25-
flowered axillary cymes; pedicels 0.1-0.4 cm long; calyx 0.3-0.4 cm long, densely
strigillose outside, sparsely strigillose inside, the lobes (4-)5(-6), deltate, about
as long as the tube; corolla ca. 0.6 cm long, fusiform in bud, subrotate at anthesis,
glabrous outside except for the densely strigose mid-petaline lines, the tube
0.2 cm long, obconic, the lobes 5, lingulate; stamens ca. 30, 0.25-0.35 cm long,
attached to the base of the corolla tube, exserted, the connective with long setae
to 0.3 cm long; pistillode absent. Female flowers in ca. 5-flowered axillary
cymes, pedicels to 0.5 cm long; calyx and corolla similar to those in male flowers
but larger; staminodes ca. 6; ovary 0.4 cm long, 0.4 cm wide, ovoid-conoidal, the
styles 4-5, 0.4 cm long, divergent, united only at the base, locules 8 or 10, uni-
ovular. Fruit ca. 4 cm in diameter, subglobose, hispidulous; seeds 8 or 10 or
fewer by abortion, 2 cm long, 0.9 cm wide, 0.6 cm thick, black; endosperm
smooth; fruiting calyx scarcely accrescent.
This species ranges from Panama southwards to Brasil.
CANAL ZONE: Barro Colorado Island, Croat 14657 (MO). Pifia, Gentry 8724, 8731 (both
FHO). Ten km NW of Gamboa, Nee 7862 (FHO). DARIEN: Santa Fe, Duke 8403 (MO).
1978] WHITE-FLORA OF PANAMA (Family 155. Ebenaceae) 149

X/:X

FIGURE 3. Diospyros inconstans Jacq.-A. Leaves and female flowers ( X) [After

Croat & Porter 15467 (MO).]-B. Fruit (X3/,-,). [After Gentry 13458 (MO).]

2. Diospyros digyna Jacq., Hort. Schoenb. 3: 35, tab. 313. 1798. TYPE: Hort.
Schoenb. 3: tab. 313. 1798.-FIG. 2.
Tree to 20 mntall; bark black, scaling in small pieces; branchlets minutely
strigillose. Leaves subcoriaceous, drying grey green or blackish; lamina to 18
cm long and 7 cm wide, elliptic, oblong-elliptic or lanceolate-elliptic, the apex
obtuse to subacuminate, the base cuneate, the lower surface glabrous except for
a few minute, in part forking, strigillose hairs, the lateral nerves in 8-12 pairs,
the tertiary and other veins forming a close but inconspicuous reticulum. Male
flowers subsessile in ca. 5-flowered cymules in the axils of leaves or bracts
towards the base of the current year's shoot; peduncle ca. 0.5 cm long; calyx to
1 cm long, minutely puberulous on both surfaces, the tube cup shaped, 0.4-0.5
cm long, the lobes 4-5, ca. 0.5 cm long, plicate, reduplicate; corolla to 1.8 cm
long, more or less conical in bud, urceolate at anthesis, the tube ca. 1.2 cm
long, strigillose-tomentellous outside, the lobes 4-5, suborbicular, 0.6 cm long,
0.6 cm wide, glabrous except for the sparsely strigillose part which is exposed
before the flower opens; stamens ca. 20, to 0.9 cm long, attached mostly in pairs
to the base of the corolla tube, included, the filament and connective densely
strigillose, the connective short apiculate; ovary rudimentary, minute, setulose.
Female flowers solitary or in fascicles of 2-3, borne otherwise as in the male
150 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 65

flowers; calyx and corolla similar to those in the male flowers but the calyx
longer and the corolla shorter and broader; staminodes ca. 12, 0.3-0.8 cm long,
attached to the base of the corolla tube; ovary 0.5 cm long, 0.4 cm wide, sub-
globose, tomentellous, the styles 4-5, ca. 0.5 cm long, united at the base, ending
on the inner face in fleshy, irregularly lobed stigmas, the locules 8 or 10, uni-
ovulate. Fruit (4-)6-8 cm in diameter, subglobose, black and glabrous when
ripe; seeds 8 or 10 or fewer by abortion, 2 cm long, 1 cm wide, 0.5 cm thick,
reddish brown; endosperm not ruminate; fruiting calyx 1.4-2 cm long, saucer
shaped, glabrous outside, tomentellous inside, the lobes hemiorbicular to ovate,
0.6-0.9 cm long, the margin undulate.
Diospyros digyna, which has edible fruit, extends southwards from Mexico
to Panama and Colombia. For centuries it has been extensively cultivated else-
where in the tropics and is widely known under the name Sapote negro. Until
recently its place of origin was uncertain. Because of its wide dissemination as
a cultivated plant its nomenclatural history is complex, and for a long time it
was known under erroneous or invalid names. Howard (1961) has recently
established the correct name.
VERAGUAS: above Santa Fe on slopes of Cerro Tute below Agricultural School, Gentry
6227 (FHO).

3. Diospyros inconstans Jacq., Enum. P1. Carib. 34. 1760; Sel. Stirp. Amer.
276, tab. 174, fig. 67. 1763. TYPE: New Granada, Carthagena, Jacquin (not
seen).-FIG. 3.
Small tree 5-8 m tall; branchlets sparsely and minutely strigillose. Leaves
chartaceous, drying dull dark brown, paler beneath; lamina to 16 cm long and 7
cm wide, oblong-elliptic, the apex rounded to shortly cuspidate, the base
rounded to obtuse, the lower surface sparsely and minutely setulose, the lateral
nerves in 7-10 pairs, prominent beneath, paler than the lamina, the nerves
and veins forming a close and conspicuous reticulum. Male flowers subsessile
or shortly pedicellate in mostly 3-flowered, shortly pedunculate cymules; calyx to
0.7 cm long, strigillose-puberulous outside and inside, the lobes 3(4), imbricate,
as long as the tube, suborbicular, apiculate; corolla ca. 1.5 cm long, subrotate,
the tube ca. 1 cm long, densely strigose, lobes 3, 0.5 cm long, 0.45 cm wide, sub-
orbicular-apiculate, glabrous except for the strigose mid-petaline lines; stamens
ca. 10, to 0.4 cm long, attached to the base of the corolla tube, included, gla-
brous, the connective long-apiculate; pistillode absent. Female flowers solitary
in the axils of leaves or of reduced leaves towards the base of the current
year's growth; pedicel to 0.4 cm long; calyx 0.4 cm long, 0.5 cm wide, shallowly
cup shaped, sparsely strigillose outside, tomentellous inside, the lobes 3, 0.2 cm
long, rounded, the margin reflexed; corolla similar to that in male flower but
shorter and broader; staminodes 3, attached to the base of the corolla tube;
ovary 0.25 cm long, 0.25 cm wide, subglobose, strigillose-tomentellous, the
style 0.15 cm long, stout, undivided, ending in 3, fleshy, bi-lobed stigmas, the
locules 6, uniovulate. Fruit ca. 2.5 cm long, 2.5 cm wide, subglobose, glabrous;
seeds 6, or fewer by abortion, 1.5 cm long, 0.8 cm wide, 0.5 cm thick, brownish
1978] WHITE-FLORA OF PANAMA (Family 155. Ebenaceae) 151

A
0?~~~~~~~~~~~~~~
f 2 C-

FIGURE 4. Diospyros salicifolia H. & B. ex Willd.- A. Leaves and young fruit (X%1)
[After Dwyer & Hayden 7525 (MO).]-B. Leaves and mature fruit (X3/m)). [After Tyson
1819 (MO).]

black; endosperm smooth; fruiting calyx ca. 1.2 cm long, saucer shaped, lobed
to the middle, sparsely puberulous outside, tomentellous inside.
Following Hiern (1873), this species is interpreted in a wide sense. It extends
southwards from Panama and the West Indies to Paraguay and Argentina.
Specimens from some parts of this enormous geographical range, especially from
the extremities, are so different that it is difficult to believe that they belong
to the same species. They are, however, connected by a complex series of inter-
mediates.
El Real, Croat & Porter 15467 (MO). El Real, Quebrada Trapiche, Duke 324
DARIEN:
(MO). El Real, Gentry 4508 (MO), 13458 (FHO). El Real, Rio Tuira, Stern et al. 455
(GH, UC, US), 763 (GH, MO, US).

4. Diospyros salicifolia Humb. & Bonpl. ex Willd. in L., Sp. PI., ed. 4. 4: 112.
1805. TYPE:? near Acapulco, Mexico, Humboldt & Bonpland (B, not seen;
photo FHO).-FIG. 4.
Shrub or tree to 15 m tall; branchlets tomentellous with short hairs and
longer sparser spreading hairs. Leaves chartaceous, drying reddish brown or
yellowish brown; lamina to 9 cm long and 4 cm wide, oblanceolate, the apex
152 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 65

FIGURE 5. Diospyros sp. I. Leaf and fruit (X45). [After Bristan 1229 (MO).]
1978] WHITE-FLORA OF PANAMA (Family 155. Ebenaceae) 153

acute to subacuminate, the base cuneate, the lower surface densely strigose on
the midrib, more sparsely so elsewhere, the lateral nerves in 4-6 pairs, indis-
tinct, the tertiary nerves and veins forming an indistinct reticulum. Male flowers
subsessile in (1-)3(-5)-flowered, shortly pedunculate cymules; calyx to 0.6 cm
long, tomentellous outside and inside, the lobes 3, with valvate or open deltate
aestivation, as long as the tube; corolla ca. 1.2 cm long, narrowly urceolate,
densely strigose outside except for the margins of the lobes, the lobes 3, deltate,
0.3 cm long, 0.15 cm wide; stamens ca. 9, to 0.5 cm long, attached to the base of
corolla tube, included, glabrous, the connective long-apiculate; pistillode 0.1
cm long, 0.1 cm wide, densely setulose. Female flowers solitary, in the axils of
leaves or of reduced leaves towards the base of current year's growth; pedicel
0.1-0.4 cm long; calyx similar to that in the male flower but to 0.9 cm long
and the lobes valvate-reduplicate in young flower; corolla ca. 1.5 cm long, other-
wise as in the male flower; staminodes 6, ca. 0.3 cm long, glabrous, attached
to the corolla tube, 3 opposite to and 3 alternating with the lobes; ovary 0.3 cm
long, 0.2 cm wide ovoid-conoidal, strigillose-tomentellous, the style 0.15 cm
long, stout, undivided, ending in 3, fleshy, bi-lobed stigmas, the locules 6,
uniovulate. Fruit ca. 3 cm long, 3 cm wide, orange yellow, glabrous, globose;
seeds 6, or fewer by abortion, 1.4 cm long, 0.7 cm wide, 0.5 cm thick, reddish
brown; endosperm smooth; fruiting calyx accrescent, to 1.5 cm long, cup shaped,
shallowly lobed, sparsely puberulous outside, tomentellous inside.
Diospyros salicifolia is widely distributed in Central America from Mexico
(Sinaloa) to Panama. It occurs in a wide range of forest types up to 1,000 m,
varying from swamp forest to scrub forest on dry rocky slopes. The fruit
frequently is not the unit of dispersal, but the seeds are often removed singly,
possibly by birds, while the fruit is still on the tree.
CANAL ZONE: Farfan Beach, Dwyer & Hayden 7525 (F, COL, MO, SCZ, UC); Tyson
1819 (MO). DARIEN: Cativo swamp, Rio Chucanuque, about half an hour below Morti,
Duke 11739a (MO).

5. Diospyros sp. I.-FIG. 5.

Tree; branchlets irregularly compressed, sparsely and minutely strigillose.
Leaves chartaceous, drying reddish brown; lamina to 20 cm long and 8 cm
wide, more or less elliptic, the apex shortly subacuminate, the base rounded,
slightly asymmetrical, the lower surface glabrous except for a few minute
strigillose hairs, the lateral nerves in 6-8 pairs, prominent beneath, the tertiary
nerves subscalariform. Male flowers unknown. Female flowers in axillary 2-3-
flowered fascicles, otherwise unknown; pedicels (in fruit) ca. 0.2 cm long. Fruit
subglobose, ca. 1.5 cm long and 1.5 cm wide, glabrescent with patches of per-
sistent sericeous hairs; fruiting calyx 0.6 cm long, saucer shaped, sparsely strigil-
lose outside, more densely so inside, 4-lobed, the lobes 0.3 cm long, hemi-
orbicular.
This species seems closely related to the Mexican and Guatamalan species
D. campechiana Lundell.
DARIEN: Cerro Pirre, Bristan 1229 (FHO, MO).
154 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 65

INDEX OF LATIN NAMES

Numbers in bold face type refer to descriptions; numbers in roman type refer to syno-
nyms; numbers with dagger (t) refer to names incidentally mentioned.
Diospyros 144t, 146 salicifolia 146t, 151
artanthifolia 146t, 147 sp. 1 153
blancoi 146t Ebenaceae 145, 146t
campechiana 153t Euclea 144t
digyna 146t, 149 Sapotaceae 146t
discolor 146t Sapote
inconstans 150 negro 150t
lotus 146t