Download (Euphorbiaceae) species (Esser, sufficiently well thyrses, large

BLUMEA
44
(1999)
149-215
A partial revisionof
the Hippomaneae
in Malesia
(Euphorbiaceae)
Hans-Joachim
Rijksherbarium/Hortus Botanicus,
P.O. Box
Esser
9514,
2300 RA
Leiden,
The Netherlands
1
Summary
The
are
Hippomaneae (Euphorbiaceae)
excluded. A
from
to all Malesian
key
treatments for the
previous
revised
are
genera
is
for Malesia.
provided.
region with
the
The
in
biogeography
Among these
sect.
are
described
genera
as new
having
genera;
Sebastiania
ly being
Sapium
sect.
and the African
as
as
on
and Triadica. All of these
used.
traditionally
Sapium
sect.
and Triadica
Falconeria, Sebastiania
Duvigneaudia is
13
species
are
accepted
Balakata
in
the
uncommon
for
names are
new
two
eight
show
sect.
but
(based
re-established
Microstachys
Malesia based
its
Sapium
as
and
on
Sapium
on
excluding
type)
distinct
Sapium
two former
and Sebastiania
new
as
previously
combinations
are
No
used
and
new
species
described,
are
but
(Sapium glandulosum, Stillingia
proposed
in the
present
revision: Balakata
luzonica, Gymnanthes borneensis, Gymnanthes inopinata, Gymnanthes remota,
Triadica cochinchinensis
neensis,
are
united with it.
revision.
present
accepted
namesare
subsp. pacifica). Numerous
Finally,
genera
indigenousto Malesia, with Sapium (= Sapium sect. Americana)former-
Shirakiopsis indica, Shirakiopsis sanchezii, Shirakiopsis virgata.
but
Balakata
Parasapium
is cited for the first time in
genus
differ markedly
cultivated.
two of the well-known
lineata
as
,
Altogether
baccata,
kept
and Homalanthus
discussed.
are
Falconeria, Microstachys
long time. Gymnanthes
longer considered
are no
only
a
is
Shirakiopsis (based
been treated
sect. Triadica for
species of
and
which
ecology,
only Stillingia
genera
Pleurostachya)
and
Excoecaria
following genera accepted: Balakata, Falconeria,
Gymnanthes,Microstachys, Sapium, Shirakiopsis, Stillingia
peculiarities
Only
generic concepts applied
re-introduced, namely
(the
synonyms
correct
are
name
Falconeria
for the former Sapium
proposed by uniting
Sapium plumerioides
with
Several
Stillingia
Sebastiania
lineata
other, previously
available
insignis, Microstachys chamaelea,
discolor),
lancifolia
and Triadica
with
sebifera.
Gymnanthes bor-
subsp. pacifica. Lectotypes
are
selected
names.
Key words: Euphorbiaceae, Hippomaneae,
Malesia.
Introduction
The tribe Hippomaneae
with
c.
300
monoecious
A. Juss.
ex
1)
thyrses,
biglandular
Neotropics,
Current
D-22609
Euphorbiaceae
1994). It is sufficiently well
large,
without petals and discs and with
in the
Bartl. of the
species (Esser,
address:
with fewer
Institut fUr
floral bracts,
comparatively
taxa
c.
30 genera
by elongate,
inclinate floral buds,
sepals.
Its
diversity
flowers
is centered
in Malesia.
Allgemeine Botanik,
Hamburg, Germany.
small
consists of
characterized
Universitat
Hamburg,
Ohnhorststrasse
18,
150
BLUMEA
For many parts of Malesia, the
Shaw
mann
of the
(e.g., 1975),
These
(1912).
based
were
Hippomaneae, however,
and fruits. For this
spectus (Esser,
in many
recent
who followed
of the
mostly
are
generic
in
Garden
Stillingia
(e.g., by
Pax
&
contrast to
applied
These
Webster
by
(Esser
(1994), although
HomalanthusA. Juss.,
1912),
but
con-
a recent
resulting
have been corroborated by
changes
following
et
effected.
not
Sebastiania
Sapium Jacq.,
to
the
Hip-
Spreng.,
Hippomaneae
into the tribe Hureae Dumort.
separated
a
al., 1998). Some
genera of Malesian
L. The genus Hura L. has often been allied
ex
reduced flowers
highly
the very diverse leaves
the Malesian taxa,
to
now
by Airy
of Pax & Hoff-
limits in the tribe were revised in
of the Malesian Hippomaneae
Hoffmann,
revised
quite recently
generic concepts
floralcharacters. The
on
The various revisions of Airy Shaw included the
pomaneae: Excoecaria L„
1999
1,
the
quite uniform,
been discussed
already
No.
have been
species
generic concepts.
phylogenetic analysis
of these had
44,
predominantly
This conspectus is
1994).
changes
the
reason,
Vol.
by
Webster (1994).
From these genera, Excoecaria is
prep.),
lished
Homalanthus (the
separately (Esser, 1997),
Homalanthuswill
was
still used
certain
the
probably
affinity,
Excoecaria
but
can
infructescence (Esser
A
taxa.
key
to
Trees
pistillate
rarely
to
with
of Malesian
flowers may be
spines. Stipules quite
of side veins
blade
not or
lacking
many-flowered,
1-20
elliptic
to
usually
few basal
usually
many distal
and with
of
un-
any genus
of
because of its
Stillingia
remains
the 'true'
Sapium
Malesian
indigenous
provided.
MALESIAN
can
HIPPOMANEAE
be summarized
sometimes and then
as
follows:
dioecy
small
to
large,
one
caducous. Leaves
in Excoecaria;
assumed),
to
simple, alternate,
petiole distinct,
symmetric, margin
or
entire
abaxially
or
triangular
to
on
or
serrate; venation
the blade close
bisexual. Bracts
ovate,
(0.4-)1.5
subtending single pistillate
ones
subtending cymules
(rarely divided) pair
of glands,
mm
or
sometimes
canaliculate, glandless
pinnate,
on
or
from the
compound,
spirally arranged,
long, covering
or
basal
the adaxial
to or remote
axillary, elongated thyrses, simple
diam., uni-
very
variously fissured, rarely
slightly different; glands nearly always present,
mm
ones
may be
Latex present, white. Indumentum absent
agallocha).
terminal
margin. Inflorescences
and
species
fit into
genera;
from the
separated
are
Hippomaneae
abaxial base of the blade, and/or
ceous
Omalanthus that
Hippomaneae
into several
of multicellular uniseriate hairs. Bark smooth
apically glandular;
or
not
pub[N.B.
herbs, often (always?) with sympodial branching. Monoecious (although
crowded, rarely opposite
pair
split
are
DESCRIPTION OF
dioecious (Excoecaria
consisting
was
1998; flowers still unknown).
Neotropical types,
general description
spelling
One isolated
treated in the present revision. Whereas
all Malesian genera will be
GENERAL
A
be excluded from the
and Sebastiania
and Sebastiania, with
the older
against
(?) myrioneura Airy Shaw, does
probably
are
by Djarwaningsih (in
the author and
by
and Hura will remain excluded from the tribe.
be conserved
al.,
et
The remaining genera
unchanged, Sapium
revised
being
has been revised
1997 (Brummitt, pers. comm.)]
by Esser,
Hippomaneae
currently
largest genus)
squama-
floral buds,
flowers and sometimes caducous,
of several staminate flowers,
rarely glandless.
persistent
Staminatepart caducous
H.-J. Esser:
after
sessile
nearly
bilateral
to
Staminate
flowering.
in bud,
to
connate
with
revision
partial
of
(.Homalanthus); petals absent; sepals free
disc and pistillode absent;
glandless;
sepals 2-6, triangular, entire;
with
excrescences,
2-3,
undivided
long,
indehiscent
with
1-3 seeds. Seeds
stamens
2-4(-20),
to
small
to
without
staminate flowers;
by
solitary
petals absent;
disc and staminodes absent; ovary 2-3-locular, smooth
descending epitropous
THE
MALESIAN
rows
90
to
of
ovule per locule;
or
style 1, stigmata
with
cm
GENERA
OF
mm
excrescences,
tall. Leaves
HIPPOMANEAE
minutely
serrate,
4.
spiny appendages
Leaves entire
of
a
extrorsely
bifurcate in Homalanthus), recurved. Fruits 3-31
shrubs up
trees.
rows
to
to
slightly
or
pendant, anatropous.
wide. Fruits with 6
b. Shrubs
fused
or
free
latrorsely
dehiscent in various ways, outside smooth
KEY TO
la. Herbs
one
(sometimes
or
inclinate
connective present. Pistillateflowers inclinate in bud,
longitudinal slits,
with
Staminate flowers
base, filaments present, anthers basifixed, opening
at
151
Hippomaneae
cymules (l-)3-18-flowered.
in the axil of each bract, sometimes surrounded
or
the
distinctly pedicellate, pedicel articulate; radially symmetric
to
zygomorphic
different degree,
A
or
distinctly
at
serrate,
smooth
spiny appendages,
or
least
rarely
15
with
mm
wide. Fruits
mm
6
12
to
up
Microstachys
of hooks
pairs
2
('muricate')
2a.
Leaves
shallowly
distinctly
to
b. Leaves entire, beneath whitish
3a.
Young
Fruits with
b. Plants
mm
Fruits with
Fruits
less than 2
of less
twigs.
than 5
present), flowering
at
seeds with
shortly (less
5a. Leaves without
on
glands
one
elongate thyrses,
often
surface
pale fleshy
separating
petiole
from seed,
on
surface
or
dry
6a. Plants with succulent
the
junction
a
on
leafy twigs.
sometimes
Pistillate flowers/fruits less
regularly dehiscent;
only
than 20
seeds with
dry
or
apex or laminabase visible from above. Staminate
sepals
remaining
and 3
at
stamens.
Seeds
dry, carunculate;
central columellaand
leaving
a
tiny
car-
scar
Excoecaria
glands
petiole
flowers 4-18 per bract,
fleshy
flowering
2. Falconeria
elongate thyrse (although
seed top
on
mm)
5
on
flowers 1-3 per bract, with free
b. Leaves with
than 1
(sometimes fleshy)
surface
reddish-fleshy
as
to
with thin
at
Shirakiopsis
least 20 per inflorescence/infructescence.
per inflorescence/infructescence. Fruits
uncle
mm,
distinctly
pericarp
4
Pistillate flowers/fruits
irregularly dehiscent;
sex
with massive
thick
mm
b. Pistillate and staminate flowers in
one
mm,
Staminate flowers sessile
pedicel
Pistillate and staminate flowers in separate,
leafless
hairs. Staminate flowers
of 8-30
6.
completely glabrous.
pericarp
yellowish
pedicel
thick
pedicellate.
4a.
3
7
not
leaves and inflorescences with
(1-3 mm) pedicellate.
least 2
or
whitish beneath
not
serrate,
apex
or
with fused
lamina base visible from above. Staminate
and 2
sepals
and carunculate, caruncle
stems.
Petiole with
a
structure
Seeds with reddish-
separating
from seed
quite inconspicuous
with the laminabase. Fruits with
tricornute
stamens.
not
woody,
after dehiscence; seeds
6
....
pair
of glands
on
indehiscent base that remains
dry. Indigenous
. .
7.
Stillingia
152
BLUMEA —Vol.
b. Plants
of
pair
succulent.
not
an
arilloid.
in cultivation
Only
gland(s)
b. Leaves without
Stipules
flowers
up
gland(s)
mm
or
only
in
with
(4-)6-20
with 2
tricornute base; seeds with reddish
laminabase
stamens.
Sapium
3
visible from above
8
...
or
apex
or
laminabase visible from above
10
venation reticulate. Staminate
tertiary
stamens.
Fruits 3-locular;
seeds
dry
and
9
Leaf blades with
Fruits
a
indehiscent
woody
tertiary
venation
usually percurrent,
leaves rarely reticulate. Staminateflowers bilateral
narrow
apex with
petiole
sarcortesta
long.
mm
a
Leaf blades with
long.
whitish with
5-200
Stipules
or
apex
petiole
on
but
5.
petiole
radially symmetric
brownish
b.
2
to
on
1999
adaxially glandless
alate columella without
base, leaving
1,
from the lamina.Fruits without
glands usually separated
7a. Leaves with
8a.
Lamina base
No.
44,
2(-3)-locular;
or
zygomorphic
seeds with reddish arilloid
....
Homalanthus
9a. Leafblades with lowermost pair of veins
Staminate flowers
(3-)5-8
b. Leaf blades with lowermost
cious. Staminate flowers
originating
at
laminabase. Monoecious.
per bract. Seeds with whitish
pair
of
veins
originating
1-3 per bract. Seeds
dry
8. Triadica
sarcotesta
above lamina base. Dioe-
and brownish
Excoecaria
10a.
5-200
Stipules
in
only
narrow
Leaf blades with
long.
mm
leaves
rarely
tertiary
venation
agallocha
usually percurrent,
reticulate. Inflorescences and infructescences
ter-
minal, simple. Fruits tardily dehiscent; seeds with reddish arilloid
Homalanthus
b.
Stipules
up
2
to
mm
long.
Leaf blades with
tertiary
venation reticulate
percurrent. Inflorescences and infructescences terminaland compound
and
simple
with
dry
to
compound.
Fruits
regularly
to
or
rarely
axillary
dehiscent or indehiscent; if dehiscent,
seeds
11
1 la. Staminate flowers 5-9 per bract. Fruits
1-
2-seeded, fleshy-indehiscent
or
..
1. Balakata
b. Staminate flowers 1-3 per bract. Fruits 3-seeded,
dehiscent
dry,
3.
1.
Gymnanthes
BALAKATA
INTRODUCTION
The
history
never
of the
classified as
two
relevant
being closely
described
as
a
species
and of the Icacinaceae ( Urandra
recognized
by
was
formally
studying
specific epithet,
transferred
by
the first available
teriflorum.
globose
of the
The latter
berries
elliptica, Merrill,
was
Merrill
supposed
(1920).
however,
very
to
untypical
was
1910), but these
(1920), respectively.
now
for
they
were
Euphorbia-
to
be
a
errors
Sapium by
described the
illegitimate, being
a
were
soon
Myrica luzonica,
Rolfe
A few years earlier, Merrill
flowering specimens,
name,
and up
Myricaceae ( Myrica luzonica, Vidal, 1883)
Rolfe (1886) and Merrill
oldest available
quite different,
related.
Balakata luzonica, with its 1-seeded
ceae, was
is
species
same
later
(1886)
(1906),
species
as
homonym
the
and
when
S.
la-
of the
H.-J. Esser:
Neotropical
S.
A
revision
partial
and
lateriflorum Hemsl.,
was
153
of the Hippomaneae
Pax & Hoffmann
replaced by
(1912)
with
S. merrillianum.
Pax & Hoffmann (1912)
of this
in
species
and erected
the first (and
were
detail.
section
because of the
Royle
Balakata baccata differs
globose, fleshy
by Roxburgh (1832)
tions under the
based
(1847;
the
even on
not
the
on
was
Sapium
S.
populifolium,
different collections)
on
pages of the
text
specimen
as
prepared
related
its bilobed 2-seeded
It
already
were
described
was
been used
he could
identical, although
description
of Excoecaria
many years earlier and
Miquel (1861)
Wallich
by
Wight recognized
was
based
described Stillingia
was
established
Hooker (1888).
by
Sapium baccatum, together
lingia by
Baillon
Excoecaria
species.
any
by
with the whole
genus
(1858,
Miiller Argoviensis
as
more;
only Kruijt (1996)
moved it
Triadica (Lour.)
to
the
which is, however,
until
transferred
was
(1877)
to
Stil-
listed it under Carumbium
not
within Sapium
of
Sapium.
was
To fit their
"semina in columella centrali
was
very
closely
not
Excoecaria. Pax & Hoffmann
sentence
In this way, Balakata baccata
position
Mull.Arg.
section, they added the
species description,
Kurz
(1866).
he also did with several other,
Since Hooker (1888), however, its
it in the section
Sapium,
but without publishing the combination under Stillingia) and
(= Homalanthus),
Reinw.
so
by
from another collection. The synonymy of all of these names
paniculata
to
that had
illustration. Finally,
Sapium
closely
contributed the first illustra-
One year later, Griffith's
Wight's
most
distinct from S. baccatum.
it had been
published (1854);
same
as
for it,
Euphorbiacea.
Wight (1853)
a name
isolated within
fruits.
an
book that both
same
figure captions.
the
change
affinis
name
baccatum.
as
the relationships
discussing
quite
from B. luzonica
fruits; therefore it has always been recognized
as
be
to
(Pleurostachya)
obviously
most
authors
only)
considered it
They
separate, monotypic
a
Falconeria
to
more
related
questioned
(1912) placed
of the
diagnosis
longe
adhaerentia" to
for its indehiscent fruits.
hardly applicable
from Balakata luzonica, and remained
separated
now.
CHARACTERS
Vegetative
characters
The genus is characterized
absent, but abaxially
visible without
absent, but
the
by
the base
a
magnification (Fig. lb).
the leaves of both
(much longer in
abaxial surface
quite
pair
always conspicuously
are
Otherwise,
Additionally,
near
B.
of
remarkable leaf
quite large,
Additional
(often
adaxial
less circular
marginal glands
ones
are
glands
is
may be present
or
smaller.
species
show
baccata), shape (ovate in
whitish in B. baccata,
the basal
glands:
more or
B.
several differences,
baccata, elliptic in
always
glands, although present
green and
B.
shining
and distinct in both
in
e.g.,
petiole
luzonica), and
in B.
luzonica).
species,
are
larger
in B. baccata.
Inflorescences and flowers
The
easy
regularly
recognition
branched
of the inflorescences
absent in related
thyrses
of the
genus in
are
genera),
are
unique
flowering
the sterile region
the
large
and
and
at
in Malesian
Hippomaneae
fruiting stage.
the base of each
conspicuous,
and allow
Other notable characters
thyrsal
branch
(peduncle,
undividedbracteoles of the stami-
154
BLUMEA
cymules,
nate
and the bracteal
Vol.
glands,
44,
which
quite irregularly shaped
are
the main axis. The individual flowers
along
1999
1,
No.
and
hardly diagnostic
are
in many characters. The bilocular ovaries and the
and decurrent
plesiomorphic
bistaminate male flowers,
constantly
however, may be noted.
the differences between the
Again,
B.
luzonica is the sterile peduncle
with
species
of each
sterile bracts
distichously arranged
are
remarkable.
(in
the
contrast to
characteristic for
Very
which is
thyrse branch,
densely
fractioned and
bracteal
multiple
fertile
spirally disposited
bracts; Fig. Id). Balakata baccata lacks this, but is characterized by the,
ficially,
covered
at
least super-
glands.
Fruits and seeds
Like in
many other
Balakata, they
fruits of the
In B.
nearly
are
luzonica, only
the
seed
one
terminal style result
case
are
(Fig. le, f).
nearly
Malesia
to
(Map 1).
AND
elaborated sufficiently yet.
and
stachys Baill.)
arisen
agree
a
with those of Falconeria,
veins and
to
Balakata
submarginal
The genus is
(B. luzonica).
to
be restricted
by
Balakata Esser, gen.
Genus
liter
novus
carnoso,
distincte
aliter
sect.
Harms,
similar
are
masculinis
pedicellatis
and
based
[=
style.
Asia, and the only
some
one
African ( Anomo-
genera, may have
within the Malesian
1998).
Hippoma-
The bistaminate flowers
but leaves and fruits
stipules,
one
in
are
quite
entire leaves with few side
(Map 1), one
E Malesia,
markedly zoochorous,
instructis
not
in W Malesia,
reaching
Australia
and their distribution
seems
of the inhabitedforest types.
earundem
5-9-floris
on
tribus
Pax & K. Hoffm. in
2,19c (1931)
Balakata
basalibus
membranaceis
indehiscentibus
sed
abaxia-
2,
indivisis
ovariis
bacciformibus
—
Type:
ornatis,
2-locularibus,
pericarpio
Balakata luzonica
S. Vidal.
Engl.,
202
eglandulosis
insigniter auctis, thyrsis
staminibus
Hippomanearum.
Myrica luzonica
ed.
adaxialiter
bracteolis
1-2-spermis
characteribus
Pleurostachya
Hoffm.
to
also in
into two vicariant species
ecological requirements
Nat. Pflanzenfam.
Pax & K.
al.,
Stillingia,
pedicellatis, sepalis connatis,
cum
(S. Vidal) Esser,
Sapium
et
glaber foliis distincte petiolatis integris
masculinis
seed aborts; in
lateral
nov.
compositis, cymulis
fructibus
and
a
and
vegetative characters, Triadica shows the closest
baccata),
glandulis submarginalibus
floribus
one
distinctly
with
develop,
glands).
diaspores
the
the
within the whole tribe cannot be
relationships
(long petioles,
leaf
India (B.
Their
Its
Sapium,
some
clearly separated
hardly reaching
a
restricted
relationships
separate paper (Esser
different in these genera. In
similarities
both seeds
rarely, however,
Compound thyrses, present
parallelisms.
as
elaborated in
genus. In
among
1-seeded berries
usually
Neotropical (Mabea Aubl., Senefeldera Mart.)
independently
neae are
unique
RELATIONSHIPS
Hippomaneae
Its
for the
typical
globular
1-seeded berries have
of the few genera of
one
that
so
In B. baccata,
flattenedand sulcate. Not
resulting globular,
endemic
most
are
bilocular berries, and
Hippomaneae.
develops,
BIOGEOGRAPHY
Balakata is
or
30 genera that constitute the
c.
the 2-seeded berries
this
the fruits
Hippomaneae,
indehiscent, fleshy, uni-
Pflanzenr.
IV.147.V
(1912) 243;
('Pleurostachys' ). —Type: Sapium
luzonica (S. Vidal) Esser].
in
Engl.
&
merrillianum
H.-J. Esser:
Trees.
Monoecious.
short
to
blade
long (1-9.5
mm
3.5-11
cordate, margin entire, apex acuminate
or
and with
pale-papillate
conspicuously enlarged
dary
veins distinct,
differing, intersecondary
not to
with
distinct sterile basal
a
tillate and staminate flowers in
gular, apically
sometimes
at
acute,
same
base with
fragmented glands
decurrent. Staminate
a
marginal
tertiary
pair
long
blade, glandless;
as
attenuate
to
acute to
towards the
of irregularly
the
margin,
veins percurrent
yellowish,
to
axis
basal
once to
ones not
twice branched,
2-7
by
mm;
of staminatecymules
pillow-shaped
ones
secon-
reticulate, smaller
staminate part 20-70
region,
basal
submarginal glands,
to
slightly
below smooth
magnification, rarely absent,
inflorescence. Bracts
touching
regularly alternate; petiole
half as
cuspidate, glandless above,
to
veins reticulate. Inflorescences terminal and axillary,
each branch with
leaves. Indumentum absent.
Leaves
wide, base
hardly joined
veins present,
155
Hippomaneae
nearly
visible without
usually
but
arching
to
cm
of several
a row
and
the
fruiting twigs
much shorter
elliptic,
to
of
revision
long, entire, glandless.
long),
cm
oblong
ovate to
and
Flowering
1.5-2
Stipules triangular,
partial
A
to
pis-
trian-
slightly flattened,
of the inflorescence and
slightly
bracteoles present, membranous, undi-
cymules 5-9-flowered;
vided, entire. Staminate flowers with pedicel elongating when flowering, but present
also in bud;
fused with
calyx basally
usually
2
absent; pedicel quite short but distinct (0.5-5
to
irregular,
than anthers. Pistillateflowers
slightly longer
fused
elliptic, slightly
at
acute
(1—)3—13
mm
at
tips;
with 2
long); calyx
a
stony seed
Distribution
mm
long);
a
thin
1-2-
sarco-
coat.
Two vicariant
—
or
base, entire, glandless; ovary 2-locular, smooth; style short,
seeded, smooth, fleshy berries, indehiscent. Seeds without caruncle, with
and
thyrse
sepals, triangular
stigmata 2, undivided, glandless. Fruits with distinct pedicel (1.5-27
testa
2, filaments
stamens
base of staminate
species,
distributed from NE India
to
Vietnam and
but unknown from Java, the Lesser Sunda Islands, and
of
parts
throughout Malesia,
New Guinea.
Note
balakat
—
The
gubat.
name
This
refers
means
to
the official
Philippine
'shoulder tree'. Thanks
are
for
name
one
of the
due to C. Ridsdale for
species,
providing
this translation.
KEY TO THE
la. Leaves
ovate to
elliptic,
2-seeded and sulcate,
SPECIES
below often whitish,
rarely
1-seeded and
petiole (2.5-)3.5-9
cm
long.
but then with lateral
globose
Fruits
style
..
1. B. baccata
b. Leaves
to
oblong
globose,
with
elliptic,
below
not
whitish, petiole 1-2.2
baccatum
Roxb.
5,
(1863) 121; Hook.f.,
Fl. Brit. India 5
Handl.
2
(1940) 276;
III-7
K.
—
Map
Fl. Ned. Ind.
(1964) 14; Malay.
(1888) 470;
3,1 (1900) 295;
Pen.
3
Diet. Econ. Prod.
Heyne,
17
G.
Watt,
ed.
Malay
ed.
Pen.
3,
1
Diet.
1832,3 (1832)694; Wight,
Econ. Prod.
Pax & K. Hoffm. in
(1924) 315; Gagnep.
Nutt. PI. Indon.
For. Rec.
1 -seeded,
1
(1853) 6; Baill., Etude Euphorb. (1858) 513; Mull.Arg.,
(1912) 240; Ridl., Fl. Malay
395, 400; Burkill,
nov.
[Hort. Bengal. (1814) 69, nomen] Fl.Indica
Icon. PI. Ind. Orient.
471; Boerl.,
long. Fruits
2. B. luzonica
1. Balakata baccata (Roxb.) Esser, comb.
Sapium
cm
apical style
2
in
Engl.,
Lecomte, Fl.
India
2
32
(1893)
Pflanzenr. IV. 147.v
Trees
(1950) 960; Wyatt-Sm„ Malay.
Biol.
6,
Indo-Chine
(1935) 1960; Corner, Ways.
(1965) 51, 113, 345; Medway,
Linnaea
J. Linn.
5
(1926)
Malaya
For. Rec.
Soc. 4
1
23,
(1972)
156
BLUMEA
Shaw, Kew
131, 138, 142; Airy
Kew Bull.
129; Airy Shaw,
Rec.
34
412.
(1877)
—
146
Not. PI. Asiat. 4
in
83]; MUll.Arg.
704
Griffith
or
Icon.
Prodr.
DC.,
706
HB
Tree, up
26
to
2
(A, P;
Fl. Ind.
stem
Bat.
iso
(holo U;
high,
when
dirty yellow
sweet
rarely
seen.
9.5
cm
long;
to
60
upper surface
hardly shining,
side, 0.4-0.8(-l.5)
mm
mm
SE Asia
I.M.
4
to
Endl.
2.
mm
Fl. Brit. India 5
461.
bole
high,
m
Sapwood
Stipules
lower surface
10-16
mm
staminate
2-3
elliptic,
with
cymules
mm
no to
0.6
c.
long; calyx
c.
long, stigmata
circular in
1
(-3)
mm
long,
0.75-2
1.5
densely
by
connate
mm
8-9
by
long.
9-11
nearly
white, soft, with
0.5
long
10-22
4-11
by
branch
at
—
E
Himalaya,
1-1.75
with
(1 —)3—12 glands
margin,
basal
circular
by
0.5-1
pedicel
c.
1.5
1.5-4
mm
India
style.
(Sikkim)
Seeds
and
c.
5
pedicel
mm
Habitat &
Ecology
forest, secondary
slopes;
in
—
Found in
MUll.Arg.,
a
0.1-0.5
over
mm
mm
by
4.5
Bangladesh
mm,
to
mm,
if
blackish.
Indochina and SW
Malaysia
(Kalimantan).
primary
it is very
and disturbed Dipterocarp
common
distinctive forest
Dipterocarps (Wyatt-Smith, 1964).
long.
long; (l-)2-seeded, nearly
forest, mixed deciduous forest, also along
Selangor (Malaya)
malaccense
Borneo
long;
0.6-1
China (only known from Yunnan), Andamans, and in W Malesia: Peninsular
(excl. Singapore), Sumatra,
of
cymules
mm
flattened with smallest diameter of 4-5
mm,
region
least super-
0.5-1
long; style
mm
covering
Bracts
mm.
anthers 0.25-0.5
sometimes absent;
or
initial
Staminate
c.
soon
in terminal
mm, at
long; calyx
flowering,
an
2-4
fragments.
mm
touching
sterile basal
a
per
glands
but
initially 75-85°,
by
—
acute to
revolute, apex acuminate,
not
midrib
base; ovary
Fruits:
cm,
base
percurrent. Inflorescences
when
sour to
a
caducous and
early
mm,
distant from
with
0.5-1.5
pedicel
thyrse
Evergreen.
deep longitudinal
withering yellow; petiole (2.5—)3.5
mm
glands
2-seeded sulcate, if 1-seeded with lateral
Distribution
—Type:
with many
above base of blade and sometimes
their
long,
numerous
10-13 per
shape,
affinis
(1833)
et Labat.
leaved.
caducous bracts, later with
with filaments 0.4-0.6 mm
Pistillate flowers
470.
(1888)
twisting
cream to
c.
conspicuous,
long,
5-flowered. Staminate flowers:
stamens
Fl. Norfolk.
Lectotype (proposed here):
few bracts, staminate part 20-70
mm
into
ficially disintegrating
c.
1211;
Forest Fl. Burma
and
bending
crown
pale-papillate,
pairs, angle
veins
—
inter Kebus
auriculate, margin
not
diam., usually 4-10
long
Gard. Bull.
(1866)
Excoecaria
—
[Prodr.
whorls and in the axils of few uppermost leaves, each branch with
8-12
2
15,
(Roxb.) Kurz,
dark grey when dead, with
to
diam. and 0-1
becoming 45-70°, tertiary
of numerous
Turner,
For.
2. Edible
(PROSEA)
Prodr.
DC.,
1950, f.
t.
non
183,
rarely elliptic, (8-)
cordate,
veins
(1861)
diam.,
cm
brown
ovate,
attenuate or
1.5-2.25
(1853)
illeg.,
brown when young,
pinkish
mostly
1
Suppl.
in whorls.
obtuse, rarely
midrib, secondary
&
(1973) 128,
Roxburghianae 2397), Bangladesh, Silhet.
2
5,
K), Sumatra, Palembang,
up
living,
Twigs usually
Leaves
blade
Icones
Orient.
cracks and fissures; inner bark fibrous.
smell.
in
2
Malaya
(1981) 341; Ng, Malay.
(1992) 355;
(1866) 1223; Hook.f.,
knobs, with irregular buttresses up
Bark
1
2,
Carumbium baccatum
PI. Ind.
15,
Tree Fl.
36
Res.
(Roxb.) MUll.Arg.
—
s.n.
PI.
(eds.),
Vietnam
Bull.
(GH, K, TCD), Burma, Mergue.
3677
m
Coronel
(1854) 486, nom. superfl.
Stillingia paniculata Miq.,
Teijsmann
83.
Type: Roxburgh
Sapiumpopulifolium Wight,
Griff.,
&
baccata
(1996)
1999
1,
(1975) 191; Kew
Ho, Cayco
P.H.
Excoecaria
—
Biblioth. Bot.
Kruijt,
2
231.
(1995)
No.
44,
26(1972) 329; Whitmore,
Ser. 4
65A; Verheij
(1991) 382;
fruits and nuts
47
f.
(1991) 83,
Bull.
Add.
Vol.
and forms,
community
Soil: brown and
streams
together
bamboo
forest,
and
with
on
hills and
Endospermum
of the late succession,
yellow clay
and loam,
poor in
sandy loam,
limestone, granitic and volcanic bedrock. Altitude 15-1800 m. Flowers collected
H.-J. Esser: A
Map
1. Distribution
in Dec-Sept.;
not
flower
exude
of Balakata
baccata
(Roxb.)
and fruit
smell
a sweet
set
is
rare
of
Esser
the
in the
Uses
—
1935; Heyne,
Note
—
as a
mealy
branched with
—
Sumatra: bedi, damar kulihap,
timber tree and
and
types
long
of the
Myrica luzonica
Trees
S.
Vidal,
J. Sci.
16
Fl. Males.
Enum.
Vidal 610
577.
(1920)
243,
(1901)
nom.
J. Sci.
nov.;
Lamao
Blumea
Prov.
2680]; Elmer,
Sapium
Kruijt,
River,
17
=
The wood is
not
very durable.
flavouring (Burkill,
Biblioth. Bot.
263.
purely
staminate
Filip.
long),
and bisexual
cm
long);
in the last
Atlas
2
Pax
146
2565
&
(1983)
Mateo, Manila;
(1996)
90.
—
(holo PNHt;
B; Rolfe,
J. Linn.
Soc.,
—
Lex.
in
Philipp.
Lectotype (proposed
note.
see
nom.
44.
illeg.,
non
Hemsl.
(1911) 1303; Merr., Philipp.
K. Hoffm.
hardly
the basal
1
(1923) 461; Salvosa,
Is.
I, Suppl. (1906) 83,
Bot. 4
case,
Sapium luzonicum (S. Vidal) Merr.,
—
Flow. PL
Philipp.
t. 90
(1883) 40,
279.
regularly
ones,
length.
Fig. 1, Map
—
Euph. Philipp.
San
ones,
cm
nov.
(1951)
Philipp.
Leafl.
Engl.,
Pflanzenr.
[Hooker's
J. Sci.
IV.147.V
16
(1912)
Lectotype [proposed by Merr., Philipp.
iso
K, P, US), Philippines, Luzon,
Prov.
Mt Mariveles.
J.
Sci.,
Stemonurus ammui
(1969)
Bataan.
4
Enum.
merrillianum
elliptica Merr., Philipp.
17, nom. illeg.
I,
J. Sci.
16(1920)577]: FB (Borden)
Bataan,
Urandra
t.
—
comb.
(K), Philippines, Luzon,
Sapium lateriflorum Merr., Philipp.
Icon. PI.
do
doelpak dollong, handoelpak.
exceeds the staminate part in
Sin. Gen. PI. Lenos.
(1920) 577;
5-7
2-3
(each
Vidal) Esser,
(1963) 123; Airy Shaw, Alph.
here):
be found:
can
(each usually
thyrse usually
(1886) 294; Backer,
Philipp.
trees
etem.
wayside plant.
as a
of inflorescences
branches
2. Balakata luzonica (S.
21
the
studied. The flowers
and in Sumatra sometimes used for
sweet
branched and with shorter branches
Bot.
Medway (1972)
Malayan population
1950).
Two
pistillate part
to
(S. Vidal) Esser (■).
(Griffith, 1854).
names
Used
are
and B. luzonica
(•),
ludai, ludai kantijl. Simaloer Is.: banai delok, banai
The fruits
157
Hippomaneae
According
fruits collected in Jan.-Oct.
annually,
Vernacular
revision
partial
—
Type:
FB
Bot. 5
(1910) 195, non
Schellenb.
(Kaneh.) SleumerJ; Philipp. J.
(Topacio)
20003
(holo PNHt;
Sci.
iso
[Bot.
16
Jahrb. 59
(1924)
(1920) 577; Sleumer,
US), Philippines, Luzon,
158
BLUMEA
—Vol.
44,
No.
1,
1999
H.-J. Esser:
Tree, up
36
to
high,
m
white, 10
mm
10.5
Leaves:
base
cm,
yellow;
petiole
attenuate
papillate
not
cm,
elliptic,
by
and
oblong
to
obtuse
0.25-0.6
mm,
1.5
c.
1
mm
mm
region
long;
stamens
long.
Pistillateflowers
0.6-0.7 mm
4-5
long; calyx
mm
style
0.25
c.
mm
yellowish
1.25-2
c.
fleshy
1.5
mm
12-17 by
layer
outer
Distribution
—
Endemic
Ecology
deciduous forests, also
to
Malesia:
Vernacular
J. For.
listed
c.
0.4
5
man
—
Enum.
c.
1
Bracts
mm.
of
not
fragmented.
mm
long; calyx
anthers
sometimesabsent;
pedicel
sepals;
ovary 3-4
pedicel
8-27
flattenedand
mm
mm
not
long;
long;
sulcate,
thick, woody part of pericarp
mm
Philippines (Luzon, Mindoro,
in the
(Mangoli),
canopy of
Locally
m.
very
common
Flowers
on
Palawan,
Ticao,
New Guinea.
primary
steep hillsides and slopes and
Philippines:
Philipp.
or
secondary,
dry
semi-
the foot of a limestone hill
and dominant. Soil;
collected in Oct.-Dec.,
clay, sand,
Mar.; fruits col-
balakat-gubat [official
Flow. PI. 2
(1923)
common name
several additional
461.
fide Phiones
are
New Guinea: seganamuung,
(Kemtoek).
—
The
locality
original type
citation is
1 Balakata
seen
and annotated by Rolfe,
[Lomibao, Philipp.
Lumber-
a
a.
can
with
and
stony
No
specimen
with
be considered
as
authentic ma-
lectotype.
Habit of
closely packed
section, showing fleshy pericarp
(Manila)".
the collection Vidal 610, from
flowering plant, only
blade, lower surface, showing glands; c.
infructescence,
17601, L],
as
luzonica (S. Vidal) Esser.
b. detail of base of leaf
young
"Montalvan?
information could be traced. However,
terial, and is therefore proposed
d.
6-7
flowering,
mm, not
The wood is of potentially commercial value
San Mateo, Manila,
Fig.
veins
numerous
19 (1973) 22-29],
Note
this
or
free
(1974) 75, 135], mogalmod (Tagbanua);
sogonam, sogonamung
—
in
Sept.
names
by Merr.,
Uses
Found
on
Altitude 10-120
lected in Feb.-June,
lipp.
—
mangrove swamp.
volcanic rock.
by
when
long
Fruits:
11-15
tertiary
to
not
tightly packed, distichously
35-40
branch
long.
Bucas Grande, Mindanao), Celebes, Moluccas
bordering
mm
per
circular
glands
Inflorescences
with
c.
lower
shining,
2(-8) glands
base of blade and
flowers: pedicel
nearly
1.5-2.5
by
thick, style apical.
mm
Habitat &
or
1.5-2 by 0.6-0.7 mm,
thyrse
with
1
basal
above
reticulate.
0.75
long,
c.
with
margin,
mm
mmlong
c.
1-3 per
mm
long, stigmata
green, with
to
5-10
with filaments
1-seeded, nearly circular in shape,
0.15-0.25
to
7-9-flowered. Staminate
cymules
to
conspicuously auriculate,
upper surface
with midrib 55-65°,
pairs, angle
their glands
long,
1.25
mm
0.5-1
staminate part
long,
brown
light
1.5-2
Stipules
often
distant from
usually
veins 7-9
shallowly fissured,
elliptic, (7—)8—18 by (3.5-)4.5-
cuspidate,
to
indistinctly percurrent
c.
to
only slightly paler,
mm
Bark
inner bark
brown.
slightly cordate,
to
acuminate
apex
high.
m
red, moderately soft, odour- and tasteless;
to
turning
Staminate
c.
17
blade
but smooth and
sterile bracts
staminate cymules
to
159
Hippomaneae
dark brown;
to
long;
cm
leaf axils,each branch with basal
arranged,
bole up
cambium white,
touching midrib, secondary
inconspicuous
the
mid
side, 0.2-0.3 mm diam. and 0.2-0.5
0.5-1.25
of
light yellow
1-2.2
margin usually revolute,
surface
90
to
revision
partial
small scales,
thick. Slash
heartwood dark
mm.
dbh up
with
strongly peeling
A
distichous
seed
bracts at
coat; g. seed
staminate flowers
detail of staminate
base;
[a—c:
e.
Ramos
fruit;
thyrse,
f. fruit in
2013, L; d—g:
present;
late
bud;
longitudinal
FB
(Curran)
160
BLUMEA
—Vol.
No.
44,
1999
1,
2. FALCONERIA
INTRODUCTION
Falconeria Royle (1839)
the Botanic Garden
F.
at
later authors with F.
lectotype
third
of 2
insignis
They
the
as
F. malabarica.
Since then,
noted that the
arated
F.
taxonomically.
insignis,,
under F.
as
containing
by
insignis
are
Sometimes F. malabarica
but the
more,
distinguishing
species,
and
was
considered
ex
L.
(Royle, 1839). Tulasne( 1851)
s.
str., close
Falconeria
ment
to
up
was
to
accepted
Miiller
Miiller Argoviensis
classified
or
is still
Falconeria as
has
a
formal section of
often been sunk into
Shaw
(1972a)
view of
(1877),
by
to
Trimen
of
of Falconeria
Shaw
(1972a).
Antidesma Burm.
with.
by
its
irregularly,
Sapium
separate
as
Sapium
may
(1885).
sections
part
Ex-
kept
finally (1880) retained Falconeria
Hooker
Within the
quite
to
either be
(1888)
most recent
insigne, however,
was
the first
strange
of Carumbium (=
with
is the
byway
to
cite
decades, Falconeria
without further distinction, for instance
as
be sep-
variety
Falconeria in the Euphor-
(1994). Only Kruijt (1996) returned,
who classified Falconeria
about
malabarica,
generally agreed
that Falconeria and
distinct genus Falconeria. A
a
might
var.
taxa
distinguished particularly
Sapium.
Sapium
and Webster
a
unisexual inflorescences. A few years later,
sections of one genus, and he
contributed by him but
not
opinions
a mere
Airy
with uncertain position within Sapium. The combination of Sapium
was
insignis
described
for several decades after its establish-
moved Falconeria and
(1878) argued
as
(1853) placed
opinion
separate genus
strictly
and the
(1866)
coecaria. Bentham
separate
a
Argoviensis (1863),
dehiscent fruits
hardly
as
as
Other authors united all
any further taxa, e.g.,
and Wight
and this
Sapium,
most
Gagnepain (1926)
combination of
Falconeria had first been classified in the Antidesmeae, close
biaceae
by
F.
flowers and fruits with 3 instead
any
two
respective
Pax & Hoffmann (1912).
without
designated
(1975). Wight (1853)
distinct from those of India and
and Hooker (1888) made the
also
accepted
of
species changed repeatedly.
from Indochina
plants
superintendent
species simultaneously,
Pfeiffer (1874)
described
were
considered Falconeria
(1890)
name.
distinguished by pistillate
no new taxa
that time
two
distinct and therefore united
Wheeler
by
at
described
hardly
accepted
the distinctiveness of the described
Pax
Royle
were
of the genus, confirmed
species,
carpels.
India.
Serampore,
and F. wallichiana.
insignia
as
named after H. Falconer,
was
by Airy
doubt,
opinion
to
the
of Kurz
Homalanthus).
CHARACTERS
Vegetative
characters
Falconeria is remarkable
Contrary
to
branches
are
always
The leaves
pair
of
and
by
but
not
are
leafless
serrate
in
margins.
on
succulent
slightly
and deciduous leaves.
stems
other succulent genus of the
tribe),
the
flower-bearing
(Fig. 2c).
characterized
disc-shaped glands
its
by
Stillingia (the only
by petioles
the
petiole
and blades
apex
on
very
variable in
length, by
This combination of features
also be found in
may
any other genus within the Malesian
a
the junction with the blade (Fig. 2b),
Hippomaneae.
Sapium,
H.-J. Esser:
partial
A
the
of
revision
161
Hippomaneae
Inflorescences and flowers
The
inflorescences
branches very
This
is
in the
unique
unisexual and
strictly
are
simple
and
different in diameter;
Hippomaneae.
terminal
strictly
do
they
Likewise
terminate
is the
unique
of
many-flowered thyrses
the top of older, leafless
at
simply
not
annual shoot.
an
of
separation
still
equal size, although
sexes
the
on
in
same
plant.
The bract
the
glands
the staminate part of
The
same
pistillate
bistaminate and sessile
The staminate
flowering (Fig. 2d).
and the staminate
disc-shaped,
are
individual flowers
of
thyrses
flowers
and
sessile. The
are
taxonomic
plant. Therefore, a
is
thyrse, therefore,
Sapium
with
cymules many-flowered
before but
shortly
when
pedicellate
from
hardly distinguishable
Stillingia.
number is 2
carpel
3, but may vary
or
does
separation (var. malabarica!)
not
on
the
seem to
be
justifiable.
Fruits and seeds
The fruits
as
in
bers
are
subsessile like the
Sapium and Stillingia.
in
opens
In
irregularly.
red). Therefore,
The seeds
are
those taxa,
and the
elongated infructescences,
flowers and of the
pistillate
contrast to
covered
the fruits of Falconeria
the
and
monotypic
most
by
element
unique
common
and size
large
num-
membranous and
whitish (never
completely
in the tribe.
RELATIONSHIPS
in the
Himalayan region;
not a
it reaches
typical
of Malesian Hippomaneae (Esseret al.,
analysis
the view that Falconeria is very
closely
staminate flowers of the three genera are
arillate seeds with
sexes,
the leaves
ularly
the
lacking
has
Stillingia
only
Malesian
most
of its
,
Falconeria
1998)
Stillingia.
confirmed
Leaves and
and Falconeria shares
But the
spatial separation
the variable carpel number, and
woody
base
for
(characteristic
of
partic-
Stillingia)
species
are
in the New World, and
Hippomane
1994). All of these genera
unique,
Sapium
L., Senefelderopsis
several
showing
are
very different in its distribution,
as
well
Steyerm.)
not
are
some
are
characterized by
autapomorphic
being
as
re-
sim-
characters.
yet achievable.
restricted
to
Asia.
Royle
Royle,
1376; Wight,
Mull.Arg.,
24
Stillingia.
about the relations within this clade
precise hypotheses
Falconeria however, is
Falconeria
fruits without a
Neotropics (Esser,
but their fruits
ilar flowers,
and
of Falconeria.
related genera (like
the
Sapium
hardly distinguishable,
flowering branches,
irregularly opening
other, closely
to
on
related to
succulence with
Sapium,
support separation
Taxon
thinly
very
are
shape
(Map 2).
The recent cladistic
More
aril and
an
are
same
found in very
are
part of Malesia (Peninsular Malaysia) and is certainly
NW
stricted
is
pericarp
BIOGEOGRAPHY AND
Falconeria is
they
111.
Bot. Himal.
Mts
Icon. PI. Ind. Orient.
Linnaea
32
in
(1839) 354,
2
DC.,
Biblioth.
Prodr.
t.
(1853) 20,
(1863) 83; Benth.,
(1975) 535; Kruijt,
(Royle) Mull.Arg.
5,
J.
Bot.
15,2 (1866)
Linn.
146
1211.
Gen. PI.
84; Endl.,
t.
Soc.,
(1996)
—
Bot.
90.
Sapium
17
—
sect.
Suppl.
Etude
1866; Baill.,
1
(1836-1840)
Euphorb. (1858) 526;
(1878) 242; L.C. Wheeler,
Excoecaria
Falconeria
sect. Falconeria
(Royle) Hook.f.,
162
BLUMEA
Fl. Brit. India 5
Lex.
Kuntze,
241;
in
&
Engl.
Pfeiffer,
Pax in
(1888) 471;
Gen.
Phan.
Nomencl.
Bot.
in
Engl.
(1903) 498;
&
2
1,
(1874) 1334]:
Etude
Baill.,
No.
44,
1,
Prantl, Nat.
Pax &
K.
ed.
Nat. Pflanzenfam.
Harms,
Gymnobothrys Wall,
Vol.
Pflanzenfam.
Hoffm.
2,
19c
1999
in
(1931)
Falconeria
5
3,
202.
—
T. Post &
(1890) 98;
Pflanzenr.
Engl.,
IV.147.V
(1912)
Lectotype [proposed by
insignis Royle.
nomen in
Euphorb. (1858) 526,
synon.
Trees, with whorled distal branches, slightly succulent and shrinking when dry. Monoecious. Deciduous,
2
or
3 ciliae,
Leaves alternate but
disc-
cup-shaped
to
(1—)2—3.5
farinose
glands
attenuate to
different in
slightly
and with
papillate
veins distinct,
secondary
on
with
angle
with
pound, basally
no or
ovate,
staminate
long,
present, completely
mm
base with
in
sepals;
separate thyrse,
visible, stigmata 2
2-
or
thin (c. 0.1
marginal
or
or
than 1 mm)
long,
2-11
of
elliptic,
cm
teeth
(less than 10
diam. Bracts
pair
a
of
glands
margin,
not
basal
di ffering,
ones not
to
veins present but indistinct,
terminal, yellowish,
not
sterile basal
long)
mm
thyrses
of
com-
region,
90-
equal size,
ofstaminate cymules transversely
oblong-flattened
disc-shaped glands
to
cymules 9-15-flowered;
3 ciliae. Staminateflowers
nearly
bracteoles
sessile in bud,
during flowering, pedicel apically articulate; calyx
2, filaments slightly longer than anthers. Pistillate
stamens
c.
30-60
ovary 2-
basally fused, glandless;
thin
towards the
flowers in separate
pistillate
5-7
basal
marginal glands,
joined
midrib, intersecondary
divided into 2
pedicel (less
with 2 largely fused
long);
not
the axis of the inflorescence. Staminate
with short
cm
asymmetrical pair
or
mostly persistent glandular
reticulate. Inflorescences
thyrses
apically rounded, at
touching
flowers
but
inconspicuous
unisexual with staminate and
170 mm
with
serrate
of few
a row
arching
densely distinct,
smaller veins
symmetrical
with the blade; blade
junction
margin
acute,
0.6-6
apart, apex acuminate, glandless above, lower surface neither whitish-
mm
nor
the
a
divided into
Stipules
apically crowded; petiole
above with
apically
much shorter than blade,
wide, base
when leafless. Indumentum absent.
fruiting
glandless.
per
thyrse; pedicel
very short;
with 3
calyx
sepals,
3-locular, smooth; style short, sometimes hardly
or
3, undivided, glandless. Fruits with
short
a
pedicel (0.5-2
mm
3-seeded, smooth, pericarp partly fleshy in young fruits, later on dry, very
mm)
and
fragile, tardily
and
irregularly dehiscing;
columella alate with
vascular bundle, central part membranous and caducous. Seeds
fleshy arillus,
Distribution
—
pale,
with
a
carunculate.
not
One
distributed from India and Sri Lanka
species,
to
Vietnam and
China, in Malesia only known from Peninsular Malaysia (excl. Singapore).
Note
—
Obviously,
The characters used
a
1. Falconeria
Falconeria
146
90.
Carumbium
—
J. &
6,
2
above
inval.
—
Mts
Ho, Cayco
Pax & K.
Vietnam
12
Bull.
(1839) 354,
Mull.
2,
var.
1
see
note
2
1.
84a
in
genus is
or
98,
DC.,
(1877)
Hoffm. in
412.
—
Brit. India 5
Engl.,
(1992) 355;
genuinum
Pax
I.M.
in
s.n.
f.
reliable.
are not
not
justified.
2; Kruijt,
Prodr.
in
Turner,
Biblioth.
(1866)
IV.147.V
Lecomte,
Tree Fl.
Gard. Bull.
7022
2
Bot.
1212.
(1888) 471; G. Watt,
Engl., Pflanzenr.
(LIV
15,
—
Sapium insigne (Royle)
Pflanzenr.
(1972) 330; Whitmore,
Lectotype (proposed here): Royle
Raj pore';
t.
Arg.
(1916) 263; Gagnep.
26
genus
2
Forest Fl. Burma
Bengal
Kew
Sapium insigne (Royle) Trimen
nom.
Fig. 2, Map
Ceylon (1885) 83; Hook.f., Fl.
Proc. Asiat. Soc.
P.H.
within this
taxa
apparently monotypic
insignis (Royle)
(1893) 471;
(1926) 394, 395; Airy Shaw,
128, 129;
distinguish
111. Bot. Himal.
Excoecaria
Cat. Fl. PL
Prod. India
Burkill,
—
insigne (Royle) Kurz,
Trimen, Syst.
Econ.
insignis Royle
insignis Royle,
(1996)
to
further division of this
Fl.
Indo-Chine
Malaya
47
Diet.
(1912) 241;
2
(1995)
IV.147.V
5
(1973)
231.
—
(1912) 242,
i), India, 'Deyra
Doon
and
H.-J. Esser:
Falconeria
wallichiana
Bot. Himal.
Mts
parte), 'Burupa
pro
Gymnobothrys
Tree, up
40
to
extremely
lucida
in
t. 84a
petiole
0.6-6
note
surface
1-7
glands
cm
to
leaf
hardly shining,
auricles of the blade,
uously
not
0.25-0.5
mm
0.75
50
Slash
base
long,
inner bark
their
wallichii
pale
anthers 0.4-0.6
c.
long; schizocarp
Distribution
China
(Yunnan,
0.5
7
—
by
mm
mm
long.
attenuate to
0.75
mm
and
glandless
often
even
in both
1.75-3.5 by
1
mm
stamens
Pistillate
flowers: pedicel
mm.
0.5-1
Seeds 5
Sri Lanka and S India
Szechuan: Lee,
long;
by
to
mm
4.5
long.
sexes,
mm
and
axis
on
0-1
2. Distribution
with
conspic-
mm
Fruits:
mm
mostly completely
pedicel
with filaments 0.75-1
long; calyx
pedicel
up
mm
1.5-2
0.5-2
mm
mm.
and SW
its southern limit in Malesia: Penin-
Malaysia (excl. Singapore).
Map
or
in lateral
1.5-2.5
Nepal, Bhutan, Laos, Vietnam,
1956), reaching
mm.
with midrib 65-75°,
angle. Inflorescences
long
1
by
diam.; blade
margin hardly revolute,
acute,
brighter
strictly marginal,
mm
Bark
Branchlets light
c.
0.75-1.5
(13—)16—23 pairs, angle
long, stigmata
4.5-6
111.
7022 iii
spreading.
crown
yellow. Stipules
with smaller
glands
0.75-1
Royle,
synon.
pale yellow.
petiolar glands
diam. and
veins
nomen in
bole unbuttressed,
cm,
cm,
mm
Falconeria
—
Euphorb. (1858) 527,
lower surface somewhat
diverging
long; calyx
mm
long; style
sular
mm
354.
the axis, therefore often
separate from the bract. Staminateflowers:
along
long,
c.
(1839)
163
Hippomaneae
1.
thicker branches, fertile part 90-170
diam. Bracts
to
or
the
of
Lectotype (proposed here): Royle s. n. (LIV
with distal
secondary
differing
—
scars.
long,
per side, 0.25-0.5
basal veins
revision
rough, light brown;
elliptic, (4.5—)8—33 by (2—)4—11
upper
f. 3.
Etude
Baill.,
dbh up
thick and very
brown, with conspicuous
Leaves:
98,
or
Nepal'; see
Wall, in
high,
m
partial
111. Bot. Himal. Mts
Royle,
(1839)
A
of Falconeria
insignis Royle.
164
Fig.
leaf
BLUMEA —Vol.
2. Falconeria
blade,
d. detail
f.
part
upper
insignis Royle.
a.
Habit of sterile
surface, showing glands; c.
of staminate
thyrses, flowering;
of infructescence with sessile
showing
thin
Maxwell
92-6, L].
pericarp;
h.
seed
[a,
e.
No.
44,
habit of
plant
starainate
KEP FRI
1999
with crowded
flowering plant
flower with
bicarpellate fruits;
b:
1,
g.
with
short
bicarpellate
(Ng) 1626, L;
c,
leaves;
pedicel
fruit in
f—h:
b. detail of base of
pistillate thyrses, leafless;
and fused
sepals;
longitudinal section,
Maxwell
90—201, L; d,
e:
H.-J. Esser:
Habitat &
sun.
900
are
Ecology
Soil: limestone,
Vernacular
Uses
name
The
—
species
of
the
recently
was
slopes,
on
to
locally
ta
Notes
Harrison
collected in Feb.,
the
contains
recorded
as
a
7022 iii, has
one,
of the
one
immature and
flowers
does
mature
show
Oct. The flowers
12 (1916) 263].
skin and eyes.
cited in the
Only
with
(the
the first
label and
fragments
selected
as
and "Khira"; it
short filaments), the other
name
species,
as
sepals
lectotype
based
was
are
for Fal-
by Royle, although
which
on
the
male
No label data for Falconeria
"Kheera"
lectotype
The
long filaments).
longioribus").
the native
at
label,
of female and male flowers;
(with
latter with
a
of
female flowers.
only
The sheet 7022 i is selected
filamentis
(7022 i) has
one
"Chhiria", "Chirun",
no
publication
could be traced
they
for F.
The first
(= ?"Khira").
wallichiana
("sepalis
filamentis brevioribus").
(1866) cited
an
isotype
number of Falconeria is
carpel
number is
However,
at
constant
least
in the
species
but will
to
not
were
characters for the
Royle's
Apr.,
Bengal
at
K, which could
not
be found
during
K.
2. The
together
cases.
present except
Miiller Argoviensis
on
male flowers
Altitude 100-
fish-stupefying plant [Kulkarni
because it has the label with all details given
not
are
integerrimis,
to
envelopes
two
of sheet 7022 iii is
envelope
carpel
LIV (7022 i-iii).
("sepalis denticulatis,
wallichiana
LIV,
contains immature male flowers
envelopes
insignis
specimen
visit
at
entire and similar in all
coneria
at
female flowers. The sheet 7022 ii has
one
nearly
preserved
"Doon" and the native names
locality
only
The third
three sheets
are
in full
rocky places,
bawl.
the efforts of A. Gunn, however,
(1978). Through
LIV. There
citing
The Royle types,
1.
—
in
common.
J. & Proc. Asiat. Soc.
|Burkill,
165
Hippomaneae
al., Indian Forester 116 (1990) 333], The latex is toxic
et
a
Mar.-Apr.; fruits
Thailand: tang
—
revision
bedrock. Occasional
bees
by Melipona
partial
Found in deciduous forest,
—
granitic
Flowers collected in
m.
visited
A
level is
not
be
and KEP FRI
two
same
in
a
much discussed
single plant,
a
In fact,
are
usually
certainly
not
justified.
Maxwell 90-201). Therefore,
a
or
fruits
separation
The recognition of varieties may be reasonable,
here. The
only
available Malesian
contain
only
staminate flowers and leaves, thus the
(Ng) 1626,
the
most common.
collections studied showed 2- and 3-locular ovaries
thyrse (Beddome 7443,
applied
matter.
and 2-locular ovaries
specimens, Congdon
747
carpel
number is unknown.
3. GYMNANTHES
INTRODUCTION
Gymnanthes
tion
sepals
described by Swartz in 1788. In its
was
(following
Pax & Hoffmann,
of the staminate flowers.
Oe (1988), it is
actually
1912)
Although
Zijlstra, 1989).
delimitation,
peltis
as
Bertero
The reduced
homoplasies
ex
it has
recently
only by
its
been revised
greatly
sepals
are to
be
(compare,
are not
e.g., Webster,
sufficient and
expected (compare,
1983
probably
reduced
superficially by
still full of nomenclaturaland taxonomic difficulties,
its distinction from Ateramnus P. Browne
&
traditionally accepted circumscrip-
it is characterized
not
including
against Kruijt
necessary for
e.g., South American Adeno-
A. Juss., also with naked flowers but otherwise very
distinct).
Other
166
BLUMEA
defining
characters
currently assigned
is still
scription
upheld
is
a
to
Mtiller Argoviensis
Bentham
African
next two
ex
inclusion of other
in
Gymnanthes
(1863),
a new
lancifolia
and S.
sect.
is retained
Sarothrostachys
resulting
other
axillary
of the
taxa
Gymnanthes,
genus,
to
taxa
like
circumscrip-
is
as
as
distinct section
a
Sebastiania,
a
position
a
distinct
distinguished by
doubted but
Leonard (1959) removed
J. Leonard.
was
correctly
Van Steenis (1948), who described the
and S. remota),
borneensis,
or
MiillerArgoviensis himself
today. Only
Duvigneaudia
Sarothrostachys.
was not
sure
as was
about their sectional
In the present revision,
already suspected by Airy
Shaw
(1975),
species.
its filiform,
axillary, basally branching thyrses,
bunches of inflorescences, but is otherwise very similar
Hippomaneae.
It is, however,
with Sebastiania. The main
not
Sarothrostachys
in
broad
considered as identical
was
assigned
including
until
upheld
species (S. lancifolia
is united with S.
remota
in
a
Sebastiania borneensis Pax & K. Hoffm.,
species,
Sebastiania
related
to
and
but
classification. Yet, all three
species fit well into Sarothrostachys.
S.
1912, whose circum-
1994, who included another
Klotzsch),
he moved it
e.g.,
(1878)
species
The first Malesian
to
Pax & Hoffmann,
by Webster,
other genera by differentauthors,
changed by
assigned
1999
described by Klotzsch in 1841. It
was
Gymnanthes by
single,
extended
A critical revision of
in later publications. In 1866,
not
1,
needed.
badly
subsection
even
plausible.
Sarothrostachys
with
or
genus, Actinostemon Mart,
Neotropical
Sarothrostachys
tion is
Gymnanthes (compare
to
No.
44,
yet available. But considering the whole range of species
not
are
—Vol.
considered
now
difference between
is the inflorescence
being branched in
to
some
be identical with
to
Gymnanthes
and
and unbranched
Sarothrostachys,
Gymnanthes.
Sarothrostachys
bastiania: leaves
nate
shares several characters with
abaxially
often whitish, with
Gymnanthes
marginal
to
s.str.
but
not
with Se-
submarginal glands;
stami-
flowers, when flowering, shortly but distinctly pedicellate; fruits often (in Malesia
always) remarkably long pedicellate; pericarp comparatively
carp thickness
10/1); septa
c.
with separate basal
thick
with
triangle,
(fruit length/peri-
only
1 (-2)
vascular
strands; seeds with smooth surface.
The characters shared with Sebastiania
the tribe: staminate flowers with
are
also found in several other genera of
well-developed sepals,
CHARACTERS
OF
MALESIAN
and 3
stamens.
TAXA
Indumentum
The Malesian
species
are,
in
contrast
to
some
of the
Neotropical
ones,
totally
glabrous.
Leaves
Compared
characters.
and the
to
other genera of the tribe, the leaves have
Gymnanthes
submarginal
row
enlarged. Gymnanthes
longitudinally
G.
remota
borneensis is
linear
of abaxial
remota
glands
distinguished by
glands,
strictly
on
a
only
mostly
the lowermost ones
shows leaves
exhibits lateral veins with
its
not
the
few
distinguishing
whitish lower surface
usually conspicuously
whitish below and with characteristic,
abaxial
larger angle
margin (Fig. 3c). Furthermore,
and with
more
conspicuous loops.
H.-J. Esser:
A
partial revision
the
of
167
Hippomaneae
Inflorescences
The
mota
thyrses
of G. borneensis
do
rarely
(but
so
mostly
often
are
branch several times, whereas those of G.
difference. It is remarkable, however, that
were
most
bisexual, but in G. borneensis pistillate
staminate. Intermixed flowers of both
for African
Duvigneaudia,
but the floral bracts
axis in G. remota,
distinctive features.
give
G. borneensis,
resulting
in
They
are
and elevatedfrom the axis
correlated with this,
Perhaps
within
sexes
a
are
rare
and
biglandular
by
width of each
comparable
described
as
quite uniform,
and touch the
thyrsal
in G. borneensis.
much shorter in
are
branch in both
thyrsal
studied
remota
are
'peduncle'
a
re-
statistical
a
thyrses usually
inflorescence,
one
of staminate flowers
pedicels
the
only
inflorescences of G.
flowers
detected. Individual flowers
not
were
glandless
this is
simple: Fig. 3a), although
species.
Fruits
There
are no
differences between the fruits of the Malesian
significant
elongated pedicels (Fig. 3a)
absent in
some
from other
are
distinctive within the
very
of
Neotropical species
genera of the
Neotropical
Gymnanthes,
taxa
of
have
Gymnanthes
quite restricted
a
long fruiting pedicels,
remota.
borneensis
It
phyletic species
and G.
More inclusive
(i.e., all species
sufficient
2
we
species
with
large
only
exclude
without any
dispersal
a
inopinata,
that their
so
This
for the very
dispersal
for
thyrses),
of dubious
occur
are
in
but
(except
particular
to
to
of
still
distributed
in
always
a
Duvigneaudia
not
affinity (like
diversity.
of
taxon.
are
for
Neotropical
a
para-
remarkable
separation
(Kruijt
are
‘D.
This is the
obviously
a
1996),
this may
trees
point
the
known
but
are
or
very
3
distribution
c.
30
with limited
to
autapomorphies
obviously
then
and 2
Sarothro-
relationships
leonardii-crispi (J. Leonard) Kruijt
we
genus,
among the
case
Sarothrostachys,
own
an own
pantropical
only
If
J. Leonard'
hexaptera Urb.),
rain forest
inopinata’),,
Within
into
are
taxon
geography.
& Roebers,
has
to, e.g.,
taxa,
inclusive
inopinata (Prain)
Sebastiania
delimited by their
poorly
contrast
'
species
drupes
old
species
are
this shows
In
assess.
some
s.str. as next
Sarothrostachys certainly
As all
the
widely
more
different elevations, but
difficult
in Central Africa
centre
considerably
species.
quite
delimitation but
species
species
ones
W Malesia,
peripheral offspring.
red flower colours in D.
Neotropical
to
generic limits,
Hippomaneae.
abilities
a
compound
uncertain: The African
Malesian
(Map 3).
Therefore, it is possible that G. borneensis is
indehiscent fruits of
conspicuous
stachys being
the
obvious,
are
study. Regarding Sarothrostachys
Malesian, 2
are
genera of the
D.
remota
known from E and S Brazil.
are
zoochory
phylogenetic hypotheses
still in need of further
consider the
restricted
is
remota.
A. Juss., the
Microstachys
as
to
in rain forests from
occurs
lower altitudes than G.
and if
adaptations
distribution
forests). Moreover, except
may be restricted.
Gymnanthes
than G.
no
are
time also known
same
RELATIONSHIPS
may be related with their habitat (primary rain
capabilities
the
they
Hippomaneae.
BIOGEOGRAPHY AND
The Malesian
but
like Actinostemon. Mericarps and
Hippomaneae
seeds agree with those of many genera in the
at
The
species.
Hippomaneae,
&
[drupes
are
in
Roebers],
similar
to
the
168
BLUMEA
If Gymnanthes proves
picture
may
diversity
is restricted
K.
Prodr.
(1931)
Missouri
Ind.
G.L.
in
122.
ed.
38
32
—
in
Engl.
Arbor. 48
Britton &
&
Harms,
(1967) 387;
Excoecaria sect.
5
3,
Hook.f.,
Nat. Pflanzenfam.
Taxon
Kuntze,
Amer. Trees
Pax &
(1890) 100;
32
ed.
2,
Ann.
(1983) 304;
Gymnanthes (Sw.) Griseb., Fl.
T. Post &
Shafer, N.
Brit. W.
Lex. Gen. Phan.
(1903)
(1908) 600]: Gymnanthes
15,2(1866)
PI.
3
(1931)
G.L.
Arch.
Taxon
24
537.
Adansonia
Pax
in
7
1
—
sect.
147.v
&
Prantl,
L.C.
in
Pax
Harms,
Nat.
Regni Veg.
81
53
53
(1944)
(1988) 2; Kruijt
(1994)
122.
(1944) 5]:
&
Lecto-
—
Ateramnus
Etude
Atlas
Euphorb.
(1858)
Sarothrostachys (Klotzsch) Baill.,
sect.
Adansonia 6
Nat.
(1918) 4;
&
note 2.
see
&
Harms,
Taxon
Wheeler,
Mull.
5
3,
Nat.
24
in
Prodr.
DC.,
Hook.f.,
(1890) 94;
Pax
Pflanzenfam.
(1975) 537]:
multiramea
[= Gymnanthes
Sebastiania
—
Arg.
Benth. in Benth. &
Pflanzenfam.
Engl.
325.
(1867)
Sarothrostachys (Klotzsch)
(1912) 118;
Wawra
Nov.
Regni Veg.
36
Engl.
Ateramnus
Sarothrostachys (Klotzsch)
Engl.
ex
van
(1841) 185; Baill.,
Stillingia
(1861) 351;
subsect.
Lectotype [proposed by
Klotzsch
Nov.
Repert. Spec.
Naturgesch.
Engl., Pflanzenr. IV.
multiramea
Een revisie
Leiden
in
(1924) 204;
Ann. Missouri Bot. Gard.
Webster,
(1975)
1175.— Sebastiania
—
Vll
Gymnanthes lucida Sw;
on
(Spreng.)Miill. Arg.
193.
add.
IV.147.xvii,
Feddes
based
(1880) 336;
K. Hoffm. in
(1756) 339; Hallier,Meded. Rijksherb.
(1983) 304; Oe,
(1989) 322;
Euphorb. (1858) 524;
stachys
a centre
Benth. in Benth. &
(1931) 207; Rothm., Feddes Repert. Spec.
Taxon
Wheeler,
sect. Gussonia
19c
s.l. therefore has
Nat. Pflanzenfam.
Prantl,
(1912) 81;
Arnold
Pflanzenr.
19c
2,
(Sw.) Rothm.,
L.C.
Gen.
&
Gymnanthes (Sw.)
sect.
Hist. Jam.
Sarothrostachys Klotzsch,
Etude
IV.147.V
(1994)
81
type [proposed by Rothm.,
13;
the
proposed here,
as
Pax & Hoffmann (1912), without
Neotropics. Gymnanthes
Engl.
J.
Engl.,
Webster,
Zijlstra,Taxon
lucidus
in
Webster,
Sapium
Browne,
Hoffm.
Pflanzenfam.
5;
Sarothrostachys,
sensu
Sw.
elliptica
K.
—
Pax
Lectotype [proposed by
—
?Ateramnus P.
&
G.L.
191;
50.
337;
Pflanzenr.
Engl.,
Bot. Gard.
(1859)
498.
the
to
1999
(1788) 95; Baill., Etude Euphorb. (1858) 530;
255,
(1880)
Hoffm. in
19c
Gymnanthes
1,
Sw.
Gymnanthes Sw.,
3
with
congeneric
as
No.
44,
in the New World.
Gymnanthes
Gen. PI.
be
change somewhat,
Sarothrostachys,
of
to
—Vol.
(Klotzsch
ed.
&
2,
Sarothro-
Wawra)
ex
Miill.Arg.].
Duvigneaudia J. Leonard,
Bot.
146
neaudia
Bull. Jard. Bot. Etat 29
(1996) 12; G.L. Webster,
inopinata (Prain)
inopinata (Prain) Esser,
Shrubs
absent
2
mm
long),
to trees.
J.
Leonard,
comb.
Monoecious.
(rarely present
in
based
Flowering
Leaves
apex
and joined
rescences
acute to
terminal and
with short
but
basal
or not
several times branched
thyrse
inopinata
in
123.
(1994)
Prain
Kruijt,
—
pistillate
or
acuminate
ones
joined
fruiting twigs
[= Gymnanthes
with leaves. Indumentum
regularly alternate; petiole
blade
(rarely
or
ovate to
elliptic,
towards the
near
base and
sterile basal
flowers often
short
3-13
(up
cm
margin, tertiary
simple
giving
or
to
0.52
cm
wide, base
Neo-
some
Neotropical species), glanda row
of
strictly marginal
veins 10-16
enlarged, secondary
venationreticulate.
compound,
by
2-5
case
axillary
and staminate flowers in
staminate thyrse 6-60
pairs,
Inflo-
in the latter
theresemblance of crowded
region, pistillate
lacking,
in
retuse
smooth and with
sometimes
axillary, yellowish,
only
no
Biblioth.
Type: Duvig-
subcordate, sometimes slightly attenuate, margin entire (serrate in
submarginal glands,
thyrses,
and
glandless;
less above, lowersurface whitish-farinose
arching
Sebastiania
& Roebers
81
Neotropical species). Stipules broadly ovate-triangular,
much shorter than blade,
tropical species),
or
on
Bot. Gard.
nov.].
long, undivided, glandless.
acute to
(1959) 15; Kruijt
Ann. Missouri
mm.
same
Bracts
H.-J. Esser:
A
revision
partial
ofstaminate cymules broadly triangular
glandless
or
removed
or
with
by
a
a
Staminate
peduncle.
Staminateflowers with short
when
flowering; calyx
and
(larger
to c.
up
thyrse; pedicel
20
cm
3
(up
2
to
slightly longer
with 3
very
and
to
20 cm)
regularly opening along
a
separate basal triangle and
columellaalate except
free
sepals, triangular,
Distribution
c.
Zaire),
Notes
of
one,
a
broadly
and
—
to
the Antilles and
in Asia
two
of
1. Circumscriptions
doubt that the
have
to
monograph,
2..Ateramnus,
no
a
are
Malesian
two
be examined
tion and
again
still
species
extant
latter. The
(Sw.)
choice
was
a.
uously enlarged;
at
to
of multiple
rows
in
drupaceous
=
often
of appendages,
c.
an African
dry
spe-
10/1), septa with
remaining
without caruncle (in
Duvigneaudia,
some
Gymnanthes
com-
in the New World from the
occur
(Congo
of Pax
sense
Basin from Cameroun
Neotropical
prefer
a
as
well
& Hoffmann (1912), the
and in need of revision. There is
closely related,
are
but their
taxa are
being
generic affinity
over
no
may
revised.
genus with very insufficient
Gymnanthes,
if
descrip-
accepted
as
convincing.
not
still doubtful position ofAteramnus instead
Gymnanthes
of Ateramnus proposed
and
by Kruijt
probably
a
changing
&
species
the
Zijlstra,
A.
name
of the
glandulosus
of Sebastiania,
and the
unnecessary.
Glands of lower leaf surface
joined
pairs
thickness
in Africa
problematical
KEY TO THE
1
two
in the
C.D. Adams, is refused here; it is
proposed
with
Fruit with distinct,
Sarothrostachys (with compound thyrses)
definite synonymy with
neo-lectotype
1-3
after fertilization
given by Kruijt & Zijlstra (1989) are
In agreement with Webster (1994) I
a
and
type, would have priority
identical. However, the
arguments
some
Malesia).
to
when the
base
very
12 in
caruncle).
poorly understood, Neotropical
known
accepting
never
with 3
or
of them
Paraguay,
(endemic
Gymnanthes (with simple thyrses)
latest available
of
conical
thyrse
basally united, glandless;
to
elliptic, dry, smooth,
Including Sarothrostachys
—
at
to
sometimes divided, vascular strand each,
twenty-five species. Twenty-one
USA (Florida)
to
smooth
(fruit length/pericarp
base. Seeds
with
Neotropical species
prises
at
only
(up
than anthers. Pistillate flowers
the septa (indehiscent and
thick
cies); pericarp moderately
3
stamens
considerably elongating
pedicel; 3-seeded,
peduncle,
bracteoles present.
small and fused
style present, stigmata undivided, glandless.
long (up
a
long) pedicel, hardly elongating
mm
ovary 3-locular, sometimes with 3 pairs of appendages but
appendages;
by
the axis of the
glands touching
sepals, quite
distinct and often
length; calyx
sometimes removed
elliptic,
Neotropical species);
some
169
Hippomaneae
cymules l-3(-5)-flowered;
distinct
to
usually
filaments
Neotropical species),
base of
with
fused in
mostly
to
of globose-cylindrical
pair
the
of
MALESIAN
slightly
SPECIES
distant from
margin,
leaves often whitish beneath, with
towards the
margin;
basal
ones
secondary
floral bracts elevated from the axis,
often
veins
conspic-
indistinctly
glandless
....
1. G. borneensis
b. Glands of lower leaf surface
whitish beneath, with
bracts
touching
strictly marginal,
secondary
the axis,
veins
biglandular
basal
ones not
distinctly joined
enlarged;
towards the
leaves
margin;
2. G.
not
floral
remota
170
1.
BLUMEA
borneensis
PI.
Enum. Bornean
Bull.
Shaw, Kew
Icon. PI. 38
Ser. 7
Bull.
Shrub
iso
brown
2.5
Hoffm. in
Kew
Bull.
iso
(holo G;
FI,
n.v.,
Bull.
thick.
scars
Bull.
15
to
m
c.
Sapwood
0.4
shiny,
from
margin,
mm
veins
10-16
acute
basal
Bracts
(1995)
345.
P.
to
up
—
32
glandless.
long; calyx
flowering, hardly present
nearly
mm
absent in bud,
long
Map 3.
Esser
with free
Distribution
(■).
mm
Tanda.
sepals;
Syn.
with
1.2-2 cm
obtuse
mm
and
diam.
compound,
up
to
5
mm
0.4-1.2
to
stamens
in bud, anthers
to c.
10
c.
mm
mm
length
ovary smooth and without
of Gymnanthes borneensis
(Pax
cream.
& K.
Stipules
blade
ovate to
el-
distant
mm
rarely absent, secondary
and
not
or
staminate part
0.4-0.8
indistinctly
except for
mm,
(6-)15-30
apex rounded
peduncle
0.4-1
mm
pedicel only
mm
up
Pistillate flowers:
when
flowering; calyx
appendages;
Esser
(•)
style
1-2
0.2
when
pedicel
c.
0.8-1
mm
and G. remota
to
long,
to
long
long.
Hoffm.)
not
rounded, often slightly
with filaments 0.3-0.5
0.25
greyish-
orange-yellowish,
entirely yellowish
a
stilt
whitish-glaucous, rarely
or
long,
by
to
to
diam. and 0.5-3
mm
Kew
(holo BO,
nov.
yellowish
long;
to
usually
6925
irregularly shaped
creamish
1-flowered. Staminate flowers:
elongating
Hooker's
Sidiyasa, Tropenbos
(1948) 410; Airy Shaw,
55-65(-75)°, arching
region
long;
&
Bibl.
(1948) 410; Airy
Lectotype (proposed here):
yellowish
to
acute to
and 0.75-3
with midrib
cymules
0.4-0.6
—
17
cm,
to
per side, 0.2-0.4
of staminate cymules
Staminate
17
Type: Bunnemeijer
subacute, removed from the axis of the inflorescence by
mm
3
(1912) 122; Merr.,
KeGler
soft, pale
brown
pale
base
usually axillary
sterile basal
Map
(1973) 131; Airy Shaw,
232.
Lingga, Sg.
petiole (0.8-)
cm,
glands
2
Buitenzorg III,
(1981)
acuminate, lower surface
pairs, angle
a
47
medium hard, whitish
glands enlarged
reddish bracts, with
mm.
to
IV.147.V
—
Buitenzorg III,
(1975) 195;
smooth, hooped,
wide. Leaves:
with 2-8
joined. Inflorescences
2-3
36
high, girth
Outer bark
long.
attenuate, apex
green and
Bull.
nov.
K).
dark green; inner bark hard,
to
Bot. Gard.
Ser. 4
Bot. Gard.
(1975) 195; Kew
liptic, (10-) 14-23 by (3—)5—13
by
Add.
comb.
Pflanzenr.
Engl.,
Bull.
K, L), Sumatra, Lingga Archipelago,
cm
1999
1,
396; Whitmore, TreeFl. Malaya
3723;
lancifolia Steenis,
50
mm
seen,
t.
small tree, up
to
c.
roots
K.
(1994) 135, f. 103; I.M. Turner, Gard.
Add. Ser. 4
n.v.;
Pax &
(1921) 347; Steenis,
14 (1960)
(1974)
Beccari PB 3127
Sebastiania
No.
44,
borneensis (Pax & K. Hoffm.) Esser,
Gymnanthes
Sebastiania
—Vol.
long,
(Steenis)
H.-J. Esser: A
Fig.
3.
Gymnanthes
b. detail of
gland; d.
detail of staminate
nate flower with free
ticidal
dehiscence
(‘triangle’);
j.
i.
ecarunculate
glandless leaf
half of
Duyfjes 18260, L].
thyrse
sepals;
with
seed
(Steenis)
remota
pedicels;
the
f.
mericarp
[a—f:
De
Esser.
base
with
fruit;
two
partial
g.
with
revision
a.
Habit
stipules;
with
c.
showing apical
De
staminate
detail
thyrses
of leaf
and
with
and fruits with
margin,
lower
surface,
pedicellate flowers;
columella;
vascular
dehiscence
171
Hippomaneae
single-flowered cymules
after loculicidal
&
the
remnant of fruit and alate
septa,
Wilde
of
strand
h.
and
pericarp
Wilde-Duyfjes 15911, L; g—j:
mericarp
basal
Wilde
stami-
after
sep-
separate split
of medium
De
e.
long
with
&
thickness;
De
Wilde-
172
BLUMEA
stigmata
3-4
c.
circular
nearly
mm
Fruits
long.
Vol.
pendant
1999
1,
No.
pedicel (5—)8—17
on
in outline,
transversely elliptic
to
44,
9—11
sulcate, without appendages, reddish-pinkish; pericarp
5-6.5
by
mm,
without caruncle, brown, sometimes
Distribution
Borneo
pelago,
Habitat &
forest
old
or
Peninsular
—
—
periodically flooded,
collected in Mar.-June,
Vernacular
Note
name
Sebastiania
36
in this
344.
Type:
—
Poetjoek Angasan,
Shrub
cm
long;
and
Bull.
Bot. Gard.
comb.
6.5
m
elliptic,
(holo BO,
high. Stipules
8-12
brighter
elliptic,
enlarged, secondary
long
when
when
mm,
by
but
ovary
flowering, hardly
with 3
1—1.5
mm
pairs
long.
by
cm,
0.2-0.3
veins 10-15 pairs,
Fruits
Habitat &
common.
in June,
Note
3
Kew Bull
L), Sumatra, Atjeh, Gajolanden,
by
0.75
and
mm
mm.
Leaves:
long;
length; calyx
mm
2-5
and with
(0-)4 glands
with midrib 75-80°,
angle
and
ofstaminate cymules
by
Endemic
—
mm
with free
on
pedicel
to
mericarps
0.55-0.65
to
up
mm,
9
cm
long.
sepals
c.
c.
0.5-1
1-1.7
mm
mm
mm
long
Pistillate flowers:
0.5-0.7
mm
mm
long;
long, stigmata
circular
long; schizocarp
10-11
0.4-
glands
pedicel
with filaments 0.5
stamens
basal
arching
without a
rarely compound,
mm. Bracts
appendages ('muricate'); style
pendant
petiole
strictly marginal,
long; pericarp
without caruncle, brown,
to
1.2-1.5
spotted.
N Sumatra.
Found in
montane
rain forest,
also
on
Altitude 1600-2000m. Flowers collected in Jan., Mar.,
—
at
accepted
this
mm
in outline, sulcate,
Ecology
but is
ultrabasic
limestone; locally
Aug.;
fruits collected
July.
tion list
digits
typical
—
on
base obtuse, apex subacuminate,
visible in bud, anthers 0.3-0.4
thick. Seeds 0.7-0.75
Distribution
waxes,
(1948) 410; Airy Shaw,
iso
1-flowered. Staminate flowers:
0.5-0.75
10-90
of small
transversely elliptic
of
found
short (8 mm).
whitish-glaucous
never
0.7
c.
cymules
flowering; calyx
to
was
removed from the axis, apex subacuminate, their
diam. Staminate
pedicel elongating
the
17
n.v.;
0.5-1
c.
3.5-4
not
0.5
clay, sand,
June-Nov.; fruits
Fig. 3, Map
—
Buitenzorg III,
8273
0.6-1
by
Dipterocarp
and red
covering
specimen
nov.
staminatepart (10-)40-65 by
mm
a
distinctly looped. Inflorescences usually axillary
mm
yellow
remarkably
are
sterile basal region,
0.6
Soil:
collected in Feb., Mar.,
petioles
Van Steenis
to
blade
per side, longitudinally
not
mixed
primary
lacks the whitish
rarely
(Steenis) Esser,
lower surface smooth and
glands
Archi-
Lingga
note.
see
small tree, up
to
0.4-0.9
the
case,
Steenis,
remota
(1981)
C Sumatra,
hillsides or in semi-riverine, swampy,
or
( Vermeulen 883)
one case
remota
Gymnanthes
6-6.5
c.
Aug.-Nov.
The lower leaf surface
—
deeply
mm,
thick. Seeds
W Kalimantan: kelampah.
—
instead. At least in
(serpentine) soil;
2.
Flowers
m.
of
understorey
ridges
on
mm
Kalimantan).
and submontane heath forest.
Altitude 15-1000
serpentine.
shiny
Found in the
secondary forest, also
C
13-19
spotted.
Malaysia (incl. Singapore),
(Sabah, Sarawak, Brunei,
Ecology
1
c.
long; schizocarp
cm
(—13) by
The type cited
by
L, this specimen
name:
Van Steenis is Van Steenis 8237.
Murdannia japonica
(Thunb.) Faden],
have been confused, and the
specimens
species.
of
According
indeed is Aneilema herbacea Wall,
Gymnanthes
two
a
ex
his collec-
to
Kunth
Commelinacea.
[currently
Obviously,
sheets of Van Steenis 8273 present
remota, therefore
are
supposed
to
be
at
isotypes
L,
of
H.-J. Esser:
A
revision
partial
4.
the
of
173
Hippomaneae
MICROSTACHYS
INTRODUCTION
Microstachys
was
De Jussieu in
published by
Martius & Zuccarini (1824a) described the
based his genus
and
on
one
a
on
three species
Malesian type
Microstachys
But Miiller Argoviensis
distinct section with Sebastiania,
all later authors
Microstachys
up
Webster
to
tiania. This
even
(1866) changed
one
of Sebastiania
species
admittedthat it
should be
probably
Esser (1998b) and Esser
minor character: its floral bracts
tichously
and
most
as
in
Microstachys.
subsequent authors,
chocroton and
the
thyrses
and fruits
Microstachys
is the
are
Both
but
as
foundedand is therefore
are
et
arranged spirally
genera were
not
united
no
whereas several
Microstachys
quite variable,
separated
stem, the
is
as
are
and the
of the
multiple appendages
cylindrical
they
(1866)
classified Ela-
some
only
of
species
characters
to
not
seem to
be well
CHARACTERS
sessile
always
OF
widespread species,
one
Microstachys,
but it poses
unique
in the
Hippomaneae,
leaves, the pistillate flowers often
thyrse, i.e.,
large stipitate
a
of
characters of leaves, flowers
unique
situated
pistillate
of the central columellaof the fruit with
seeds with
less dis-
be subdivided in any way.
mostly opposite
from the staminate part of the
peculiar morphology
most
not
easily recognizable by
the inflorescences
Microstachys only
here.
taxonomic difficulties and it should
particularly
from Sebas-
not more or
and
discussed below. This distinction does
ignored
as
by
al. (1998).
Muller Argoviensis
by
It is well established that in all of Australasia there is
M. chamaelea. It is
followed
(e.g., by Airy Shaw,
separated
Pax & Hoffmann (1912);
by
was
different sections of Sebastiania. The symmetry
as
only difference,
shared,
ones
based
his mind and united it
Elachocroton was described by Mueller in 1857. It differs from
in
Neotropical
it has often been noted that section
(1994), although
corroborated by
was
two
(and only) species
he did with other genera. This
as
is very different from other
Webster (1994)
1975).
De Jussieu
authors up toGrisebach (1859) and Miiller
accepted by subsequent
Argoviensis (1863).
a
Tragia L.,
Indian type. The first
an
few months before Von
a
Cnemidostachys.
Cnemidostachys linearifolia Miq.
was
was
as
genus
included in
formerly
( Tragia chamaeleaL.) with
1824, only
same
at
different parts of the
flowers and fruits, the
regularly parallel margins,
caruncle.
MALESIAN
PLANTS
Indumentum
The often dense, colourless and
However, in M. chamaelea they
bearing
if
not
hairy,
them
are
are
quite
very characteristic:
totally hairy (glabrous
and the leaves
in
stipules
most
initially pilose
stiff hairs
shorter than in
other
but
widespread
are
most
and all
other
sepals
species);
in
species,
are
always
Microstachys.
and the
at
organs
least ciliate
the ovaries instead
are
never
early glabrescent.
Leaves
Extraordinary plasticity
name
of the
Neotropical
M.
in leaf shape and size is
polymorpha
typical
Mull. Arg.!).
for the
Especially
the
genus (compare the
peculiar
serrulation
174
BLUMEA
the
(Fig. 4b), i.e.,
Vol.
No.
44,
1999
1,
densely packed, enlarged, glandular
other, is useful for distinguishing
leaves with short but distinct
petioles,
strictly marginal glands
together
are,
the
teeth
from other
Microstachys
each
usually touching
genera. The
appressed (not erect) marginal
never
ovate
teeth and the
with the indumentum, sufficient for
recognition
of M. chamaelea within the genus.
Inflorescences
The
inflorescences
Microstachys,
the insertion of the
of the
elongation
of the
thyrses
the
useful characters
provide
not
quantitative
axillary
along
sexes
do
not even
very
ones.
often
for
the leaves. It
opposite
bud of the uppermost leaf.
delimitation in
species
for the whole
is, however,
genus
Noteworthy
originates by sympodial
is also the
Typical
spatial separation
stem.
Flowers
Most floral characters
of M. chamaelea
the
are
the very
Hippomaneae
remarkably
are
filiform,
not
narrow
fleshy
the
thyrses,
of the staminate flowers and the sessile
sepals
make
cences
within
constant
thick and
stipitate
pistillate
Microstachys.
in
as
to
other
some
falcate bract
The
stigmata
species.
flowers with the
Within
the free
glands,
typical
excres-
Microstachys easily recognizable.
Fruits
The fruits of Microstachys
oblong-cylindrical mericarps
(Fig. 4f;
absent in few extra-Malesian
stipitate caruncle,
alate
along
wings
unique
are
are
within the tribe. Notable
and seeds, the
multiple
its whole
obovate). Otherwise,
length
with
absent
or
to
is
a
there
are no
Microstachys
is
(Esser
and (when
badly
narrower at
other genera the
most
base, the columella shape then
differences between
has
to
the other
species
of
Microstachys.
RELATIONSHIPS
species formerly
in need of revision.
by
from
Microstachys
ships
of the Malesian
species
included in the collective
In
S and E Brazil,
many
De Oliveira
probably always
cannot
species
Only
the South American
(1983),
been
quite
be discussed
of
but the
separation
taxa
species,
artificial. Therefore, the relation-
now.
Microstachys
three of them local endemics and
as
well
as
and variable
to
a
occur,
and
M.
some
of them
only
a
quite
chamaelea is
small seeds,
a
shore
and therefore
are
as
local
probably
chamaelea, more widespread
corniculata (Vahl)
[M.
single species,
high degree. Therefore, Microstachys
only by dispersal. Microstachys
currence
one,
In the New World north of the Amazon
in Australasia (
( M. chamaelea) there is
of Elacho-
of Elachocroton
endemics, and often hardly variable in their characters. In Africa there
four
dry)
al., 1998).
et
treated recently
(Esser, 1998b).
the
somewhat isolated genus without any very close relative. In Malesia,
Excoecaria than
genus Sebastiania
croton were
particular
of the fruit surface
large disc-shaped
margins (in
nearly parallel
distinctly
BIOGEOGRAPHY AND
Microstachys
in
are
and the central columellaremaining after fruit dehiscence which is
completely
it is closer
the
species),
excrescences
may
plant
probably
each
Griseb.]
widespread
have reached Malesia
with often
easily
gregarious
dispersed.
oc-
This is
H.-J. Esser:
the fact that it is
supported by
border of floristic
markable
from the
Microstachys
A.
Benth. &
Ann.
fam.
Hook.f.,
Gen. PI. 3
(1931)
19c
Microstachys (A. Juss.)
by
G.L.
Webster,
Cnemidostachys
—
Adansonia
Baill.,
Webster,
192.
Pflanzenr.
—
6
F.
Pax
in
Engl.,
3.
—
Herbs
Type:
Etude
2,
323.
350.
than
elliptic,
1
ovate
19c
long;
or
to
few
marginal
and
usually looped
reticulate.
simple,
falcate
of
absent.
absent
to
—
5
(1890)
& Kuntze
subsect.
Lectotype [proposed
—
bicornis A. Juss.
[=
&
Gen.
PL
Sp.
Etude
Zucc.) Baill.,
&
1
&
Zucc.)
Zucc.)
Kew Bull.
(Oct.
Euphorb.
53
G.L.
(1998)
[= Microstachys daphnoides (Mart. & Zucc.)
Inflorescences
in
ones
the axis
0.5-2
of the
(L.) Miill.Arg.].
and
fruiting twigs
long,
mm
undivided
mm
and often
apart
and
to
some
glabrous,
pilose,
at
veins
often
not
opposite
to
acute, with
a
1-2
pair
Staminate
mm,
nearly
Neotropical
lower surface
veins
visible but
arching
presumably
leaves, yellowish,
and staminate flowers in
by
blade
base often with
slightly larger, secondary
5-60
or
subcordate, margin with very
glandless
thyrse.
Nat.
(1983)
regularly alternate; petiole short
0.3-0.6
axillary,
thyrse
(F. Muell.)
Harms,
de Janeiro 27
chamaelea
Flowering
white, glabrous
region, pistillate
&
Engl.
Jard. Bot. Rio
to
acute
hardly visible, tertiary
staminate
in
entire, glandulous margin in
basal
Elachocroton
Neotropical species), glandless;
some
never
sect.
Elachocroton
uniseriate, colourless hairs, mostly
Leaves
wide, base
an
Stillingia
sect.
Hoffm.
Monoecious.
glandless.
cymules triangular,
glands touching
K.
—
[= Microstachys
glandular teeth,
but
&
Oliveira, Arq.
F. Muell.
terminal and
separated,
Pax
Stipules triangular,
long
cm
de
17.
(1857)
Sebastiania
—
mucronate, above
acute or
papillate
but often
staminate
498.
955.
Benth. in
3,
Cnemidostachys (Mart.
sect.
of multicellular,
submarginal glands,
flowers
(1863) 90;
Nat. Pflanzen-
Harms,
T. Post
(Spreng.)
(1903)
&
Cnemidostachys (Mart.
sect.
114;
high.
m
consisting
without sterile basal
pistillate
Bracts
to
32
(1998)
Nat. Pflanzenfam.
Engl.
(Apr-June 1824) 136; Nov.
517.
(1912)
each other (rarely fused into
and smooth
in
53
15,2 (1866) 1166;
(1994) 121]: Microstachys
Mart. & Zucc.
asperococcus
linear, 0.3-2
taxa), apex rounded,
paler
Prantl,
Linnaea
Bull.
Lectotype [proposed by Esser,
—
(1931) 192; A.S.
dense, minute and persistent
touching
81
Excoecaria
—
386.
(1967)
IV.147.V
Elachocroton
cm
&
(1912) 91;
Sebastiania
Sebastiania
—
divided into several ciliae,
apically
Kew
Prodr.
DC.,
Engl.
J. Bot. Kew Gard. Misc. 9
vegetative parts, rarely
(less
in
Cnemidostachys (Mart.
Euphorb. (1858)
with leaves. Indumen tum
on
50; Miill.Arg.,
Lex. Gen. Phan.
7.1 Beil. 4
sect.
subshrubs, less than 2
to
stressed
was
2.
Pflanzenr.
Pflanzenfam. ed.
Pax in
Bot. Gard.
(1861)
1
(1867)
Mueil., Hook.
(F. Muell.) Baill.,
f.
15,
t.
(1994) 122; Esser,
IV.147.V
Kuntze,
Flora
Zucc.,
Stillingia
note
see
81
sect.
Sapium
957]: Cnemidostachys myrtilloides
Elachocroton
a
The absence
established and
sufficiently
re-
be
(Vahl) Griseb.].
J. Arnold Arbor. 48
Miill.Arg.];
be
eventually
usually underrepresented.
are
1824) 48,
Gard.
(1880) 336;
T. Post &
Adansonia
(1858) 515;
(Feb.
Ann. Missouri
Mart. &
40.
t.
plants
seems to
Bot.
Engl.,
corniculata
Microstachys
1824) 66,
Australia and has crossed the
collection from New Guinea may
any
Microstachys (A. Juss.) Miill.Arg.
sect.
2,
ed.
Gen.
Missouri
Pax & K. Hoffm. in
94;
to
(1980).
Juss., Euph.
Webster,
Sebastiania
Shaw
175
Hippomaneae
A. Juss.
Microstachys
G.L.
The lack of
Philippines, however,
recently by Airy
the
of
fromAfrica
widespread
because seashore
collecting artifact,
revision
partial
On the other hand, its absence from
of Malesia is
parts
regions.
(Map 4).
A
same
glabrous
thyrse
to
or
pilose.
of flattened-orbicular to
cymules
bracteoles absent. Staminateflowers with very short (less than 0.5
1-3-flowered;
mm
long) pedicel,
176
BLUMEA
when
hardly elongating
staminate
thyrse
with 3
often
or
1999
1,
spathulate,
glabrous
with
usually
from staminate part and
separated
6
hirsute;
to
10/1), septa
to
longitudinal
style
spine-like
with
regular,
of
rows
absent,
usually
free
quite large,
a
thin
of minute basal
spine-like
excrescences
with very
glands;
each
on
regular shape,
dry, dehiscing regularly
the
along
(fruit length/pericarp thickness
exocarp
small separate basal
a
sepals;
base of
different height, sessile;
on
pair
a
at
stigmata 3, undivided, glandless.
excrescences,
hirsute; mericarps with
very
with
or
long) pedicel; 3-seeded, oblong
mm
of
rows
septa, glabrous
with 2
usually
Fruits with short (0-2
>
with 3
flowering; calyx
sepals, triangular, free, glandless
ovary 3-locular,
carpel,
No.
44,
3, filaments longer than anthers. Pistillateflowers 1 (very rarely 2)
stamens
calyx
Vol.
triangle
and 1 furcate vascular
strand; remaining central columella very regular, slightly alate with parallel margins
over
tent
its whole length. Seeds
caruncle
and lenticular when
stipitate
Distribution
—
of the Amazon,
Fourteen
one
W
Notes
species
based
posed problems,
distinctiveness of the
and
Philippines,
have
recently
N Australia,
to
not
primarily
only
Asian
and several
however, has
species,
1994). However, the plate illustrating
of the
genus
diagnosis
panying
the
also the
only
of the several
one
C.
by
1.
Esser
chamaelea
Microstachys
PI.
L., Sp.
Excoecaria
Miill.
5
DC.,
Bot.
Chine
5
Wild
Backer & Bakh.
Malaya
686;
1
2
2
(1992) 352;
Hoffm. in
Fl.
Fl.
Malay
(1959) 464,
Fl. Java
Turner,
Herb.
1
Miill.
was
a
The
shape.
questioned.
simul-
lectotype (e.g., Webster,
also cited
C.
reason
F.
Arg.
32
f.
Fig.
6
2
as
the
one accom-
(1824a),
was
and it
of
drawings
was
diagnostic
obviously
myrtilloides
meant as
was
proposed
Tragia
chamaelea
H.
Kew
Keng,
Gard. Bull.
(1867)
323.
vol.
Muell.,
var.
4:
3
IV.147.V
&
Handl.
4
chamaelea
Fl.
Ned.
Pen.
26
in
2
Lecomte,
(1935)
Bot. Gaz.
124
335
3,
Fl.
1
J.
IndoM.R.
(1962) 58;
(1972) 339; Whitmore, Tree
(1975) 195;
Bull.
—
(L.)
Fl. Brit.
Ind.
1988;
Kew
Bull.
Sing. (1990) 113; P.H. Ho, Cayco
Gard.
835.
(1826)
(1912) 116; Merr., Philipp.
Malay
Bull.
3
(1873) 151; Hook.f„
47
(1995)
232.
(1857)
17.
—
35
(1980)
Vietnam
2,
Lectotype
(BM), Ceylon.
J. Bot. Kew Gard. Misc. 9
asperococca
—
Sebastiania
Maitreyi,
Add. Ser.
(1993) 87;
no.
1.
(1924) 317; Gagnep.
N.C. Nair
Bull.
43,
Hook.
note
—
Fl. Austral. 6
Econ. Prod.
Cone. Fl.
45
see
(L.) Spreng., Syst. Veg.
(1891) 619; Boerl.,
Penins.
418;
4
Map
(1863) 95;
Pflanzenr.
Diet.
4,
(1964) 498; Airy Shaw, Kew
Hermann
asperococcus
(L.)
Engl.,
55.5-14; Burkill,
(1980) 244;
I.M.
(proposed here):
chamaelea
Gen. PI.
(1973) 131; Airy Shaw,
Muelleria4
Elachocroton
f.,
f.
—
(1866) 1175; Benth.,
Revis.
10(1915) 190; Ridl.,
Hend., Malay.
Fl.
15,
Pax & K.
(1926) 454,
been
of Cnemidostachys
and therefore it
chamaelea
Adansonia
(L.) Baill.,
Prodr.
Linnaea
Arg.,
Cnemidostachys
(1888) 475; Kuntze,
(1900) 268, 296;
Sci.,
—
chamaelea
in
Arg.
India
Mull.
(L.)
981.
(1753)
notoriously
(1998b).
chamaelea (L.) Müll. Arg.
Microstachys
has
species
never
select
myrtilloides
1824b)
in
Microstachys (Esser, 1998b).
species
to
one
Malesia in the
throughout
that included separate
plates
characters (Von Martius & Zuccarini,
lectotype by
Mexico, and
to
Von Martius & Zuccarini
illustration representative for the genus. For this
as
S America south
tropical
characters of indumentum and leaf
on
authors avoided
previous
large persis-
known from New Guinea. Three
yet
been attributed to
2. Von Martius & Zuccarini (1824a) described 17
taneously,
in
ten
In the New World, the distinction between the
1.
—
Asia
through
a
dry.
species currently accepted,
but absent from the
part,
ends, dry, with
truncate
in northern S America and C America up
the Old World from Africa
additionalAfrican
with
elliptic-oblong
(F. Muell.)
Pax in
Engl.,
—
Sebastiania
Pflanzenr. IV.147.V
(1912)
H.-J. Esser:
J.
117; Merr., Philipp.
add. VI
MEL,
Sci.,
iso
Bot.
Cnemidostachys linearifolia Miq.,
(holo U;
iso
K), Java, Banka,
Perennial herb
originating
mm
to
(-1.2)
cm
cm,
a
0.6-2
by
acute
mm,
completely
axis
by
sessile
of 5-6 each;
sometimes
to
olive brown
pericarp
c.
0.15
—
Widespread
India and Sri Lanka
Malesia: Peninsular
to
cm
15
F. Miiller
Type:
—
long.
(holo
9
to
to
and
ciliate. Leaves:
by
0.1
long,
thick. Seeds
0.5
with 0-6
13-35
c.
mm
long,
long
2.5-2.15
by
long, nearly
with
distinctly
long,
excrescences
by
2-2.25
distinctly
ciliate
Fruits 3.5-4.5 mm
mm
pairs, angle
1-flowered. Staminate
mm
with
0.4
to
strictly
part 5-15
staminate
cymules
mm.
petiole
teeth, apex rounded
glabrescent,
long); calyx
mm
in total diameter;
lanceolate, 1.6-6 by 0.3-0.8
Inflorescences:
Staminate
cm
Indumentum 0.25-0.4
sepals;
green,
in 6
mm,
rows
brown,
mottled.
in Africa
(e.g., Cameroun, Togo)
and in Asia from
Vietnam and S China, the Solomon Islands and N Australia;
Malaysia (Thailand, Malaya, Singapore), Sumatra, Java,
(Brunei, Sabah, Sarawak).
Map
147.xiv,
s.n.
Type: Kurz (as 'Amann')
tall and 30
cm
appressed
±
pilose
orange, with 0.6-1
brown-yellowish-whitish
Distribution
90
elliptic
0.6-1.2
0.5-1
mm
460.
of staminate cymules
glands.
calyx
to
Pflanzenr. IV.
Engl.,
—
diam., secondary veins
mm
visible.
c.
(1861)
c.
with
0.2-0.4 mm
glabrous; stigmata filiform,
turning pinkish
blade
Bracts
the bract
(pedicel
ciliate sepals. Pistillateflowers:
ovary
1
densely pilose
mm,
margin
hardly
usually glabrous.
covered
flowers nearly
root
but smooth,
per side, 0.2-0.25
347.
(1921)
ad flumen Victoriae'.
Suppl.
tap
0.2-0.5
bright
177
Hippomaneae
aridis ad rivulam Muntok'.
pratis
obtuse,
to
with midrib 45-75°, often
1-1.5
Fl. Ind. Bat.
yellowish-brown
mucronate, lower surface
marginal glands
PI.
locis sterilioribus
'in
the
of
Pax & K. Hoffm. in
Bornean
long, pilose, glabrescent;
base
revision
many-stemmed subshrub, up to
from
long. Stipules
0.1-0.25
Enum.
'in
K), Australia,
partial
11(1916)76;
Bibl.
(1919) 58; Merr.,
n.v.;
A
4. Distribution
of
Microstachys
chamaelea
(L.) Müll.Arg.
Borneo
178
BLUMEA
Vol.
44,
No.
1,
1999
H.-J. Esser:
Habitat &
also
on
Ecology
alluvial and lateritic
the whole year
Vernacular
Uses
and
in the
particularly
together
to
300
rainy
does
Flowers and fruits collected
m.
1935).
with
meat
and
(1824),
to
whole young plants
vegetables,
use
his
constitute
not
valid
a
the first
was
of the
plant
as
astringent, tonic,
genus. But because he did
definitely
chamaelea cannot
34.1
publication (Art.
and accept this
to use
mentioned
,
not
be attributed
under Sebastiania
(Microstachys)
ICBN). Milller Argoviensis
name.
Connaropsis monophylla
the herbaria of Bentham and
lea; the same is
true
recognized
pines, really
3.
Contrary
although
the
weaker
from
to
Hook.f.,
was
2329
at
specimens,
totally
Cuming
Hook,
ex
f.)
Oxalidacea. At K
can
be referred
2324 is
only Cuming
found
not
an
all. Thanks
labelled
2324
an
are
as
to
2324
Hallier
f.,
(including
M. chamae-
originally
due to J. F.
labelled
Veldkamp
collected in the
Philip-
Malaya.
data in the literature, leaves with entire
appressed
to
ex
Hooker), only Cuming
this confusion. Both
of the fruits
appendages
are
are
Planch,
for FR. At L, however,
M. chamaelea, and Cuming 2329
who
Cuming
chamaelea. Nevertheless,
monophylla (Planch,
is the type collection of Sarcotheca
on
to
2329. Miiller Argoviensis (1866) and Pax & Hoffmann (1912) cited both
Cuming
based
Tragia
combine
synonym, which
as a
2. There has been confusion in the labels and the literature between
(K)
are
[5 (Mamit) 35141], Dragendorff
describing Microstachys
when
new
the combination Microstachys
species,
specimens
Borneo:
in India.
syphilis
belonging
as
(1863), therefore,
and
Soil: sand,
season.
him (Art. 33.1 ICBN). Baillon (1858) cited this combination only
also
places,
waste
gregarious.
ambin-ambin (Burkill,
1935) mentions the
by Burkill,
1. De Jussieu
—
and open woodland, often
Peninsular Malaysia:
—
diarrhoea and
Notes
beaches, sandy sites, roadsides and
Altitude: sea level up
but
179
Hippomaneae
recovery for women after giving birth
speedy
chamaelea
on
the
of
(Lundu Malay).
(1898; repeated
against
revision
partial
grassfields
When cooked
—
used for
Found
ground.
through,
names
daun merayat
genus
—
lateritic outcrops,
A
teeth
are a
very
absent.
margins
were
never
found,
very small and may be overlooked. In Malesia, the
are
regular
Specimens
feature. Those ofAustralian
from Australia
may
also be
plants, however,
distinguished by
their often wider leaves.
4.
Airy
Microstachys
chamaelea is sometimes confused with
Shaw in the herbaria,
fruits with
petals
a
and carinate, reddish seeds. Thanks
the determination of the latter
Fig.
b.
4.
Microstachys
detail
of
leaf
inflorescence,
sepals;
e.
young
e—h:
split,
often
Forman
&
are
due to C. Barker for
(L.) Müll. Arg.
a.
Habit with
pistillate
densely packed glandular teeth;
fruit;
d. staminate
f. fruit with
as
a
with the
861, L].
c.
flower, nearly sessile,
typical
excrescences;
large septa,
typical triangle;
Blewett
providing
species.
with
columella; h. mericarp
and the basal
, L;
and
infructescence;
teristic alate
3709
chamaelea
margin
Sauropus bacciformis (L.)
herb with similar habit but entire leaves and smooth
i.
g.
the
seed with
flower and staminate
detail
with
of habit
large
bract
partly opened fruit
apical
large
vascular
caruncle
strand
[a—d:
with
thyrses;
stipules,
glands and
with
(left
free
the charac-
upper
Haviland
part)
& Hose
180
BLUMEA
—Vol.
No.
44,
1999
1,
5. SAPIUM
INTRODUCTION
in Malesia has
Sapium
been known for its difficult circumscription.
long
old Greek word of Plinius, used for Pinus but
an
described
by
Browne
the former
(1760),
but has
(1756),
probably being
a
referring
its sap.
to
been conserved in the
recently
sense
name
was
of
is
first
Jacquin
Zijlstra, 1989; Brummitt,
&
Gymnanthes (Kruijt
The
Sapium
1994).
The first Malesian
In the
1805).
scribed
Sapium
in
or
Sapium
following decades, several species
combined; but also several
newly
taxonomic
species
Pax
history.
upheld by
most
authors up
Webster
to
(1994).
were
sometimes
assigned
Excoecaria
to
e.g.,
to
indicum (Willdenow,
to
Sapium.
Stillingia (Baillon, 1858), resulting
& Hoffmann (1912)
This
a
sometimes
to
other genera,
Indeed, Sapium is characterized by thered aril of the seeds, accompanied by
elaborated
as
Eusapium
must
the
only
sepals,
two
Malesian
subg.
Sapium
Americana Pax & K. Hoffm.' of Pax & Hoffmann
the
to
Neotropics
and
comprises
described under
species
Sapium (the pollen
has
20
c.
species, mostly
not
regard. This, however,
the
species, Sapium being
ing
the delimitationof
allowed
one
is
a
lack the aril and, therefore,
Sapium
been
as
a
source
of
species
of rubber and
Sapium
are
have been
consequence of the difficult determination of
of the most difficult
genera
in the Euphorbiaceae
the
species (Webster, 1967). Fortunately,
clarification, although Kruijt did
a
Sometimes S Ameri-
studied).
have been introduced in Malesia
Sapium
Kruijt (1996)
'
It is therefore identical with
records of the genus for Malesia. At least six
cited in this
constant
stamens) and pollen ('equatorial
characters of their leaves.
indigenous
of
sect.
indigenous
be excluded from
plants
can
is
taxon
distinguished by
All
by Kruijt (1996).
Pax & K. Hoffm.
This
(1912).
blurred
species, e.g.,,S. indicum,
some
1926),
broad
a
circumscription, however,
result,
Sapium (Gagnepain,
applied
earlier authors; it is still
by
(Smith, 1910).
features of the staminate flowers (two fused
ring'),
de-
Sapium, newly
removed from
were
concept of Sapium, including several sections separated
the delimitation against other related genera. As
S.
was
assigned
were
united with other genera, i.a., with
was even
quite complicated
a
described under
species
recent
of
consider the
explicitly
not
regard-
monograph
Malesian cultivars.
Sapium Jacq.
Sapium Jacq.,
Nat.
[Civ.
Enum.
Hist.
J. Arnold Arbor.
Bot.
Gard.
7
(1841)
1
(1861)
Engl.
498.
&
—
81
187.
351.
Syst.
48
—
(1912) 200,
nom.
(1756) 338];
123.
Stillingia
Sapium
Nat.
Excoecaria
subg. Eusapium
(1760) 9;
2
(1967) 391; Kruijt,
(1994)
—
Prantl,
PI.
Jamaica
—
sect.
sect.
Biblioth. Bot.
43
(1994) 114],
Gen.
146(1996)27;
Eusapium
3,
5
Sapium (Jacq.)
Mull.
Arg.,
(1890) 98;
Mull.
Arg.
Linnaea
32
T. Post &
in
DC.,
inval.;
in
Engl.
&
Harms,
S.
G.L.
Webster,
Klotzsch,
Arch.
Naturgesch.
Adansonia
(1863) 115, nom. inval.;
Kuntze,
Prodr.
Nat. Pflanzenfam.
P. Browne
Sapium
G.L. Webster, Ann. Missouri
Lex. Gen. Phan.
15,2 (1866)
Pax & K. Hoffm. sect. Americana Pax & K. Hoffm. in
nom.
non
(1824) 49;
Sapium (Jacq.) Baill., Etude Euphorb. (1858) 513;
Sapium aucuparium Jacq., nom. illeg. [=
(1989).
[Taxon
Juss., Euphorb.
Stillingia subg. Sapium (Jacq.)
Pflanzenfam.
sect.
cons.
A.
ed.
2,
Engl.,
19c
1202.
—
Pax in
(1903)
Sapium
Pflanzenr. IV.147.V
(1931)
198.
glandulosum (L.) Morong]; compare Kruijt
—Type:
&
Zijlstra
H.-J. Esser:
A
partial
Trees. Monoecious. Leaves present
Stipules broadly
sent.
Leaves
ovate to
of
1.5-2.5
triangular,
blade
quite variable,
with teeth 3-5
serrate
mm
less
whitish-farinose and with
long, undivided, glandless.
than half
2.5-9
as
below
that often
papillate
nor
cilia-like
marginal lobes, basal glands hardly different, secondary
tinctly percurrent
Bracts
axis of the
of
reticulate,
to
axillary, yellowish,
staminate
with
largely
with
cymules
fused
a
pedicel
sepals;
absent
to
0.5
c.
3-seeded
with
10/1), septa
completely
covered
Distribution
the
(1-
the
a
the
mostly
flowering;
or
to
10
at
2-locular in
base of staminate
species;
some
species;
some
to
in Malesia),
not
not
shortly
smooth,
thin (fruit length/pericarp thickness
septa, pericarp
columella
remaining
slightly
alate. Seeds
red aril.
Approximately
—
in
vascular strand,
one
by
2-seeded
or
and restricted
twenty-five species, indigenous
to
Argentina
to
and Bolivia. Introduced
tropical regions.
Sapium glandulosum (L.) Morong
Ann. New York Acad. Sci. 7
Sapium glandulosum (L.) Morong,
146
(1996)
Enum.
44.
Syst.
aubletianum
Diet.
Econ.
Arbor.
Tree,
the
24
—
PI.
(L.) Miill.Arg.
Hippomane glandulosaL., Sp.Pl. (1753)
(1760) 31,
var.
Prod.
(1943)
Malay
Miill.Arg.,
2
Bull.
Herb.
1.5-2.2
with
a
1.2-2
by
of
pair
mm,
often
when young, later
but
nate
not
part
glands
serrate
per side
mm,
joined
towards
2-2.25
by
mm
to
—
162.
c.
long
the
6
margin,
mm.
and
basal
Bracts
mm
Sapium biglandulosum
nom.
(1906) 362,
superfl.
nom.
—
to
20
m
persistent.
t.
up
to
40
cm
irregular longitudinal
mm
petiole
diam.
2.5-6.5
by
J. Arnold
8.
tall and dbh
Leaves:
0.5-1
229 f.
Sapium
superfl.; Burkill,
Lectotype [proposed by Croizat,
smooth with
Bot.
0.8-2.4
in
fiscm
usually separate
cm,
base
acute
to
and in small lateral auricles of the blade 0.4-1
completely
with filaments 1.25
—
(1863) 117,
VI
Biblioth.
Sapium aucuparium Jacq.,
nearly entire, apex acuminate, below with
veins (12-) 14-23
7-8-flowered. Staminateflowers:
stamens
2,
(1855)
obovate, (5—)10—17
strictly marginal
secondary
60-140
1
32
distinctly stipitate glands
elliptic
glands
—
buttressed. Bark grey,
from the blade; blade
0.5-1
Boiss.
1960.
(1935)
obtuse, margin
0-3
Linnaea
known in Malesia but up
measurements not
Stipules
Pen.
Retzia
superfl.; Hassk.,
(1893) 227; Kruijt,
1191.
176]: Plukenet, Phytographia Almagestum (1694)
Neotropics, slightly
sures.
nom.
aubletianum
(Miill.Arg.) Huber,
long, apically
c.
when
long only present
mm
from Mexico and the Antilles
Neotropics
inflores-
disc-shaped glands touching
Pistillateflowers up
into the USA, sometimes cultivated in other
by
same
2, filaments longer than anthers. Pollen
stamens
short; ovary 3-locular (1-
dry, opening regularly along
1.
of
pair
a
terminal and
Inflorescences
Malesia), smooth; style short, stigmata undivided, glandless. Fruits sessile
pedicellate;
>
found in small
are
veins distinct, basal
and staminate flowers in
pistillate
equatorial ring (fide Kruijt, 1996).
thyrse; pedicel
in
in-
or
neither
usually
Staminate cymules (3-)7-18-flowered; bracteoles present,
thyrse.
with 2
smaller veins reticulate.
compound,
not
small. Staminate
flowers with
calyx
the blade,
as
different, intersecondary veins present, tertiary veins usually distinct, indis-
ones not
cence.
long
wide, margin entire
cm
marginal glands
ab-
Indumentum
fruiting twigs.
mm
glandless above,
apart,
181
Hippomaneae
and
flowering
on
the
spaced; petiole distinct, always
regularly
usually apically glandular;
distinctly
revision
of
on
not
with midrib 65-70°,
differing. Inflorescences:
staminate
cymules
c.
0.6
mm
arching
stami-
long,
their
the axis below the bract. Staminate
cymules
pedicel
long
pairs, angle
ones
up
when
to
0.5
mm
flowering,
long; calyx
1.25
anthers 0.5-0.75
mm
mm
long;
long.
182
BLUMEA
Pistillateflowers
5-10 per
c.
3-locular;
ovary
style
9
3-seeded, nearly circular,
Seeds
5
c.
4.5
by
Distribution
—
Habitat &
long,
mm
Indigenous
to
mm
long,
1.5
c.
green-brownish;
with 3
glandless sepals;
long.
mm
pericarp
Ecology
—
the
sessile;
Fruits
0.25
c.
thick.
mm
and
Singapore
In the
Neotropics
in the lowland
regions,
fromthe Caribbean Islands and Mexico
Neotropics
In the first half of this century
Argentina.
in the Botanical Gardens of
in coastal
1
long, stigmata
mm
1999
1,
mm.
the Guianas and
to
No.
44,
thyrse; sessile; calyx
0.5
c.
Vol.
up
400
to
in Malesia, e.g.,
planted
Buitenzorg.
found
m
in
especially
altitude. It is the
forests and
secondary
most common
species
of the genus.
Uses
Planted
—
and in
some
fore it
never
sonous
became
latex
Notes —
atical
one
as
a
properties
of rubber. The rubber is of
superior
to
It is also used
warts.
is the
species
most
of this difficult genus.
high quality, comparable
Hevea rubber, but is difficult
commercially important.
against
1. This
source
even
by
Burkill
Indians
variable and
(1935)
to
to
harvest; there-
mentions
use
of the poi-
catch birds.
taxonomically
Kruijt (1996) suggests
to
the
unite 57
problem-
most
(!) in this
names
species.
2.
Burkill
tried in
S.
(1935) cites
Singapore
the
Sw. [an
jamaicense
Both
(1996: 59)].
at
are
two
other American
beginning
illegitimate
name
distinct from S.
by
their elliptic,
by
its 1-seeded fruits. Because
hardly
for S.
whose cultivation had been
species
of this century,
namely
S.
may be
Malesian
specimens
S.
e.g.,
jenmanii
species
and
Kruijt
distinguished,
glands,
of these
Hemsl.
Griseb.:
laurifolium (A. Rich.)
glandulosum, they
obovate leaves with indistinct petiolar
no
jenmanii
also
were seen,
the
be confirmed.
names cannot
6. SHIRAKIOPSIS
INTRODUCTION
Hurusawa
name
the type
species,
the section
to
(1954)
applied by
Sapium
Shirakia
Parasapium,
as
illegitimate
was
described the genus Shirakia.
Sapium
because
replaced
a
japonica (Siebold
which
later
homonym
species
a
species
their leaves
of
Esser
species
a
and
Hook.f. The
S.
nudum)
was
based
to
on
allied
finally
name,
namely
japonica,
of Shirakia, added
provided
S.
however, is
cochinchinensis
the latter
two
being
the
type.
additional Malesian
the necessary combinations.
share similar inflorescences. Two of them, however,
and fruits:
the earliest
fossil fern
genus, Shirakia S. Kawas., and
in Shirakia Hurus.,
preliminary synopsis
ones
on
a nomen
(1998a).
indica (Willd.) Hurus., and S.
and several African
These
in
in
by
1843,
movedbetween different genera but
Parasapium (Miill.Arg.)
Hurusawa included three
(Lour.) Hurus., S.
was
is based
name
& Zucc.) Hurus. Shirakia
sect.
with Neoshirakia
Kruijt (1996),
The
Von Siebold & Zuccarini (Croton shiraki,
cochinchinensis, which is
now
are
quite
transferred
to
different
Triadica
(see there), and S. japonica. The latter differs by, e.g., being totally glabrous and by
often entire leaves,
fruit
different, laminar leaf glands, much larger stipules, and
wall. A cladistic
included
(Esser
et
al.,
study
1998)
of Malesian
showed
that
Hippomaneae,
in
Kruijt's concepts
which S.
cannot
a
japonica
thin
was
be maintained.
H.-J. Esser:
Either
this
A
broad concept of Shirakia
a
must
would have
Gymnanthes
case
most
includes three Malesian
Shirakiopsis
indica
discussed
Later
later authors.
it
on,
183
Hippomaneae
has
japonica
applied here,
new
was
from
separated
genus has
a new
This
Parasapium.
Gymnanthes (in
be
to
to
be
esta-
genus, Shirakiopsis,
species.
described by Willdenow (1805)
was
several
to
names
as
S.
bingyricum,
hurmais)
for S. indica, which
contributed the
Wight (1853)
indicum. It had
Sapium
authors,
pre-Linnean
Hamilton (1837) and Merrill (1917). Wallich
(1847) cited
by
nomina nuda (Sapium
some
the
S.
or
case,
of the section
been known under different
already
of
include additional species of
priority),
the remainder of the genus. In the latter
blished, congruent with
revision
partial
some
were
was
taken up
first illustrations of this
moved around between several genera,
as
additional
by
species.
including Excoecaria, Sapium,
Shirakia, and Stillingia.
Miquel (1861)
for
a
long
Excoecaria
(1981,
sect.
Triadica),
was
considered
Miquel (1859),
species, poorly
Argoviensis
identical with S.
Excoecaria
all later authors and is
recognized by
as
This
Miiller
by
indica
known
(1866,
by Airy
as
Shaw
indica).
of
species
[Sapium] diversifolia.
treated under Triadica
even
Excoecaria
as
Another
was
described Stillingia
time and
The third Malesian
species
described
was
by
here
Merrill
It remained poorly known for several decades, and
Zoll. & Moritzi
virgata
accepted
ex
Miq.,
too.
(1913)
Sapium sanchezii.
as
only Kruijt (1996)
gave additional
information.
CHARACTERS
Vegetative
characters
The indumentum of coloured hairs is
of the three
size and
species
are
Shirakiopsis indica,
their leaves. This is
are most
and remarkable. Also the leaves
glandless petioles,
of the blades, the serration, and the
proportion
usually enlarged (Fig. 5a-c),
ones
uncommon
characteristic. The short
are
unique
S. sanchezii and S.
very
uncommon
for
in Malesian
virgata
are
the
quite
marginal glands
constant
with the basal
Hippomaneae.
definitely indistinguishable by
where
Hippomaneae,
usually
leaf characters
useful for differentiation.
Inflorescences and flowers
The inflorescences of Shirakiopsis
The mentionedpubescence
of the bract
latter
are
two
glands
and the
characters
few and
agree
are
to
those of
higher
number of
with Triadica.
as
most
is the
long-pedicellate
Contrary
to
other
Hippomaneae.
regularly elliptical shape
staminate flowers. The
Triadica, the pistillate flowers
shortly pedicellate.
Like in characters of the leaves, S.
distinguishable
most
similar
is, however, remarkable,
indica, S. sanchezii and S. virgata have hardly
inflorescences and flowers.
Contrary
to
the leaves, this is
common
in
genera of the tribe.
Fruits and seeds
The fruits of each
support
with
a
species
of
their delimitation.The
thick
pericarp
and
Shirakiopsis
species
always
one
are
very
characteristic and, therefore,
have trimerous and
comparatively large
vascular strand per septum.
They
fruits
may dehisce
184
BLUMEA
immediately
and
with
hard capsules,
woody,
very indistinct
—Vol.
S.
with
virgata
absent in all
of
species
Shirakiopsis
and the Lesser Sunda Islands, S.
all
species
a
few localities of the
shore
are
is remarkable. This is
resemble each other very
adapted
to
in
floating
about one year
long time;
a
and
by
its fruits with
to
of S.
indica
Although
and endemic
species
syndromes,
the
well-
being obviously
after
germinate only
may be
its
by
serrate
thick fruit wall and
remarkably
pubescence,
easily recognized by
leaves without
dry
its indu-
petiolar glands,
seeds. The inflorescence
resemble Triadica, but this is based
Gymnanthes,
cladistic
a recent
as
study (Esser
et
al.,
some
on
Within Malesian Hippomaneae, the genus is however most
symplesiomorphies.
related
a
for the
except
widespread
the seeds of S. indica
Hippomaneae, Shirakiopsis
mentum, especially of the inflorescences,
characters,
Java, and S. sanchezii is
to
and the Lesser Sunda Islands.
capsules
a
Philip-
1991).
(Ng,
Within Malesian
for
and have
closely
but absent from Java, the
is restricted
virgata
Philippines
dehiscent
hardly
water
widespread
correlated with the different dispersal
probably
very thick-walled and
are
RELATIONSHIPS
the difference between
plants,
indica
The caruncles
species.
vicariant distribution (Map 5): S. indica is
pines
irregularly (S.
to
drupe-like fleshy ones).
BIOGEOGRAPHY AND
known from
1999
1,
regularly (S. sanchezii: Fig. 5h), orretardedly
to
The Malesian
No.
44,
closely
corroborated.
1998)
Excoecaria differs in several characters, e.g., the sessile staminate flowers with 3 free
sepals
and
(mostly 3)
3 stamens, the fruits with
constantly
vascular strands; it is
The three African
reddish (not
species
yellowish)
are
not
very
very
hairs and
similar
to
related.
the Malesian
ones,
differing mainly
often very small fruits
two-carpellate,
in Africa and Asia,
Shirakiopsis, therefore, occurs
very thin wall and septa with several
a
closely
on
(Esser,
both continents with
a
in
1994).
comparable
diversity.
Shirakiopsis Esser,
Genus
novustribu
gen.
Hippomanearum pilis
laminis serratis subtus
centiis
non
nov.
papillatis
nec
nec
masculinis
ramosis, cymulis
glandulas marginales ferentibus,
plurifloris,
floribus
masculinis
cellatis, sepalis connatis, fructibus septicidaliterdehiscentibus,
sime carunculatis
on
Sapium
Shirakia
Hurus.,
Biblioth.
Bot.
Linnaea
32
vel
J. Fac. Sci.
146
(1863)
in
123.
Stipules
to
yellowish
ovate
to
—
Mull.
Engl.
Trees. Monoecious.
pale
Univ.
Tokyo,
(1996) 7,91,
(Mull. Arg.) Hook.f.,
of
—
TyP e: Shirakiopsis
seminibus
indica
inflores-
distincte
pedi-
siccis minutis-
(Willd.) Esser,
based
indicum Willd.
sapium (MU11. Arg.)
(1912) 249;
ecarunculatis.
foliorum
obtectis, petiolis eglandulosis,
coloratis
albidis
Excoecaria
in
Arg.
DC.,
Fl. Brit. India
&
Flowering
alternate; petiole 0.6-2
cm
sect.
Prodr.
Bot. 6
—
15,
2
pro
sect.
(Hochst.)
(1866)
1216.
—
Pax & K. Hoffm. in
ed.
fruiting twigs
mm
(1954) 317,
Excoecaria
Sclerocroton
(1888) 471;
some
1-2
long,
3,
Pflanzenfam.
and
(reddish in
triangular,
5
Nat.
Harms,
Sect.
parte excl. type.
pro
2,
19c
(1931)
parte
excl.
type; Kruijt,
Parasapium MU1I. Arg.,
Mull.
Arg.
Sapium
Engl.,
subsect. Para-
sect.
Parasapium
Pflanzenr. IV. 147.v
202.
with leaves. Indumentum consisting
African taxa), multicellular,
long, undivided, glandless.
much shorter than blade,
uniseriate hairs.
Leaves
glandless;
blade
regularly
oblong
to
H.-J. Esser:
elliptic
to
ovate, base obtuse
apart, apex
but
acute
basal
ones
pairs)
and
slightly
to
papillate,
not
slightly larger
but
arching
revision
partial
the
of
attenuate,
185
Hippomaneae
margin
with teeth 3-5
serrate
mm
acuminate, glandless above, lower surface paler than upper
to
whitish and
not
A
but very similar,
strictly marginal glands
secondary
towards the
joined
not
with 0-10
margin,
veins
basal
quite
pair
numerous
similar.
one
each side,
on
(15-26
Inflorescences
terminal, yellowish, simple, without sterile basal region, pistillate and staminate flowers
in
thyrse,
same
each
triangular, pilose
ciliate,
(3-)5-7-flowered.
pedicel distinct; calyx
at
with 3
Pistillate
length.
base, glandless;
with 3
ovary
least 8 mm)
1-3
at
3
at
base;
stamens
thickness
fleshy
<
3, filament and anther of
sometimes absent;
thyrse,
in S.
species), undivided, glandless.
virgata), regularly
with
very thick
fused
species), smooth, usually
3-seeded (2-seeded in African
mericarps
cymules
in bud and
species) sepals, irregularly triangular,
in African
(2
elliptical
to
long) pedicel
mm
3-locular (2-locular in African
pedicellate;
of staminate cymules
elongate-spheroidal
base of staminate
in African
(2
smooth, dry and woody (partly
(fruit length/pericarp
fused
sepals,
flowers
septa (sometimes tardily so);
a
of
pair
Bracts
pilose.
mm,
a
and sometimes decurrent. Staminate
thyrse
glabrous; style present, stigmata
distinctly (at
with
Staminate flowers with distinct (1-3
flowering; calyx
similar
4-6
by
base
at
the axis of the
glands touching
when
40-120
thyrse
to
Fruits
species) mericarps,
dehiscent
along
in Malesian
pericarp
the
taxa
10/1; but much thinner in African taxa), septa with
separate basal triangle and 1 vascular strand, central columella alate. Seeds elliptic,
dry,
caruncle very
Distribution
inconspicuous
Six
—
Asia from India
to
to
absent.
from these three in
species,
Cambodia and
Africa and three in
tropical
Malesia up
throughout
to
tropical
the Caroline and the
Solomon Islands in the Pacific.
Note
—
for the
Except
the genus is characterized
still insufficient, but
Neotropical
taxa,
an
too.
unusually
thick
mostly by
elaborationwould
The
species
of the fruits and the indumentum,
pericarp
the absence of characters. Its
very similar
are
to
circumscription
is
studies of African and
require comparative
each other, but
at
least the fruits
allow clear distinctions.
KEY TO
la. Fruits
acute
at
both ends,
THE
longer
MALESIAN SPECIES
than wide,
always
with
a
fleshy
Java
3. S.
b. Fruits
ripe.
layer.
outer
basally
—
rounded
slightly
attenuate,
at
least
as
wide
as
virgata
long, dry
when
Not in Java
2a. Fruits
very hard,
spread
to
—
2
hardly
and
slowly
but
regularly opening, hardly
sulcate.
in Malesia, but absent from the Lesser Sunda Islands and the
—
Wide-
Philippines
1. S. indica
b. Fruits
regularly
and
easily
opening,
deeply
sulcate.
—
Lesser Sunda Islands,
2. S. sanchezii
Philippines
1.
Shirakiopsis
Sapium
indicum
indica
(Willd). Esser,
Willd., Sp.
PI. ed.
4,
1832,3 (1832) 692; Buch.-Ham.,
Wight,
Icon.
PI.
Ind.
Orient.
6
4
comb.
nov.
—
(1805) 572; Roxb.,
Trans. Linn.
(1853)
t.
Soc.
17
Map
Hort.
5
Bengal. (1814) 69;
(1837) 229; Hassk.,
1950; Baill.,
Etude
Retzia
Fl.
1
Indica
ed.
(1855) 158;
Euphorb. (1858) 513; Benth.,
186
BLUMEA —Vol.
J. Linn.
Soc.,
India 5
(1888) 471;
Pflanzenr.
Bot.
Fl.
Pen.
Malay
K.
Econ.
Fl.
Malaya
Ser.
8
PI. Res.
SE Asia
3,
17946
see
Bot.
Bot. 6
2,
14
1
Herb.
Roxb.
Roxburgh
in
Mtill.Arg.
DC.,
Bull.
353.
36
Biblioth.
37
(1940) 277;
(1972) 330; Whitmore,
f.
I.M.
65B;
Linnaea
J.J. Sm. in Koord. &
Goel
&
Bot.
(1996)
(PROSEA handb.)
93.
Herb.
Type:
—
P. H.
Sci. Univ.
J. Fac.
IDC microfiche
n.v.,
Bull.
Chakrab.,
Wulijarni-Soetjipto (eds.),
(Willd.) Hurus.,
146
in
Valeton,
(1975) 114; Kew
PI. Res. SE Asia
&
Turner,
(1863) 123;
32
(1982) 20;
Lemmens
1
tannin-producingplants (1991) 73,74;
indica
(holo B-WILLD,
s.n.
26
Add. Ser. 4
Bull.
(eds.),
in
Malaya
(1991) 83,
(1877)413;
Kew
& Coronel
and
34
Kew Bull.
(1981) 297;
Shirakia
Rec.
Burkill,
(1926) 394-396;
(Willd.) Mtill.Arg.,
Forest Fl. Burma
—
Baill.,
ex
Fl.
?iso
Ind. Bat.
Icones
BM, BR;
1
Suppl.
Ho,
Tokyo,
Willdenowfol.
1297/7), India, Tripura;
Handl. Fl. Ned. Ind.
A;
iso
G-DC,
3,
(1900)
Bot.
296.
IDC microfiche
n.v.,
146
10-11.
see note
—
1.
diversifolia (Miq.)
Excoecaria
—
Biblioth.
1
f.
(1858) pi. 6,
Roxburghianae 1296);
461.
(1861)
15, 2 (1866) 1211; Kruijt,
U sheet in
Atlas
Euphorb. (1858) 513,
Etude
(holo G;
s.n.
Prodr.
of
(holo U; drawing
(1996)
88.
Sapium
—
—Type: Teijsmann HB
4215
2619/9), Sumatra, Lampong,
Toelang bawang.
Tree, up
30
to
m
and buttresses up
high,
to
2
high, branching
to
to
40
peeling
in small
blade
sparsely pilose to glabrous;
numerous
oblong
to
to
olive,
rectangles;
outer
greyish
white
homogeneous, dirty
of moderate weight and hardness, with
with
twisting,
2-4
bark
to
pale yellow
pores. Leaves:
elliptic
to
to
diam., basal glands 0.5-0.9
mm
mm
at
base
thin; inner
diam. and often
smell.
sweet
1.1-1.4
petiole
ovate, 7-14
slightly
in
thick,
mm
straw-coloured,
base obtuse, apex subacuminate to acuminate, lower surface with 2-4
0.25-0.4
spines
Young twigs pilose, especially
fibrous. Slash with strong,
light brown, darkening rapidly,
and heartwood
bole
cm,
from base.
Bark brown
yellowish.
to
furrowed and fissured,
yellow
Sapwood
diameter up
stem
m
Indumentum pale
vertically
bark
indica
(1991) 376; Purwaningsih
diversifolium (Miq.) Boerl.,
axils.
For.
in
(1921) 348; Ridl.,
PI.
1.
note
fluv.
(1950)961; Airy Shaw, KewBull.
(1990) 738; Verheij
(1992)
5
Engl.,
J.J. Sm.
(1917) 328;
Trees
1961; Corner, Ways.
Excoecaria
Pax & K. Hoffm. in
Herb. Amboin.
Fl. Indo-Chine
Lecomte,
Fl. Brit.
(1880) 335; Hook.f.,
6,2 (1893) 471;
Bibl. Enum. Bornean
(1910) 615; Airy Shaw,
Kew
Stillingia diversifolia Miq.,
a
1
(1954) 317; Kruijt,
Sapium bingiricum
Type:
—
Buchanan Hamilton
=
3,
in
(1935)
(PROSEA handb.) 3, Dye
Vietnam
Cayco
12
fruits and nuts
Edible
Sect.
(1995) 231.
(1980) 89;
J. Econ. Tax.
2,
ed.
2
(1973) 128, 129; Ng, Malay.
Boomsoort. Java
Bijdr.
Pen.
Malay
2
Gen. PI. 3
Hook.f.,
(1917) 548; Merr.,
15,2 (1866) 1216; Kurz,
Prodr.
Add.
12
(1924) 317; Gagnep.
Prod.
Gard. Bull. 47
DC.,
3
Benth. &
Diet. Econ. Prod. India
1999
1,
(1912) 251; Merr., Interpr. Rumph.
Nutt. PI. Indon.
Heyne,
Tree
Watt,
Nova Guinea
Herderschee,
Diet.
G.
IV.147.V
in
(1878) 242;
17
No.
44,
3-4
by
glands
per
cm,
cm,
side,
the midrib,
touching
secondary veins 18-24 pairs, angle with midrib 60-66°, smaller veins distinct. Inflorescences
pilose
30-55 by 6-8
to
1-2
pedicel
mm
flowers:
1
rounded
long; calyx
at
mm
or
black when
or
mm
mm.
c.
1.5
attenuate
ripe, dry
mm
c.
at
stamens
5
mm
and without fleshy
sometimes
often with
less
very
outer
irregularly
mm
mm
4-6
long.
Pistillate
1.25-1.75 mm
mm
18-30
by
long.
Fruits:
20-32
slightly sulcate,
layer; hardly
broken
than three seeds,
trimerous; mericarp with pericarp wall (2—)3—4
sparsely pilose;
long; calyx
in outline,
not or
long,
mm
with filaments 0.5-0.6
long, stigmata
base,
1.25-2
Staminateflowers
long, ciliate;
nearly globose
unopened,
loculicidally;
ofstaminate cymules
absent; pedicel
long; style
slightly
dehiscent and often shed
septicidally
or
Bracts
0.5-0.9
absent in bud, anthers 0.4-0.5
long; schizocarp
both ends
becoming
mm
by
0.6-0.8
flowering, nearly
ovary 2.5
8-22
pilose.
1-1.75
(rarely 2) per thyrse
long, pilose;
and
mm
when
long
pedicel
axis
mm,
ciliate, their glands
but
or
and
mm,
green
tardily
opened partly
always regularly
thick, septum remaining complete-
H.-J. Esser:
ly
4
c.
mm
long
medium
the back,
partial
therefore without any
mericarp,
at
only
A
to
to
septal
nearly absent,
pale brown,
revision
not
the
187
Hippomaneae
gap or basal
alate. Seeds
7-8.5
by
mm,
Distribution —Widely distributed from Sri Lanka and India (Malabar
to
columella
triangle; remaining
11-13
keeled
on
without caruncle.
spotted,
not
of
coast,
Ganges)
Thailand, the Caroline Islands and Solomon Islands; in Malesia known from Penin-
sular
Malaysia (incl. Singapore),
Guinea
Habitat &
Ecology
grove forests,
inundated
up
to
the Bismarck
including
75
in
Soil:
places.
m.
—
primary
E Sumatra, Borneo, Celebes, Moluccas, and New
Archipelago.
Found
along
rivers and seashores, in
and advanced
clay, sand,
mud.
Very
tidal and
gallery,
forests of swampy and
secondary
common to
Flowers in Malesia collected in Dec-Jan.,
man-
seasonally
scattered. Altitude: sea level
June-July,
outside Malesia
however, the whole year through; fruits collected the whole year through. The seeds
germinate only
Uses
—
after 318-413 days
The timber is used for
Leaves have been
leaves
can
as
boiled
are
applied
together
be obtained from it
a
fish
poison,
Kinomeri
1935)
to cure
and
can
e.g.,
to
as
supposed
suitable for indoor work.
gonorrhoea (Burkill,
black
dye,
in lower Siam: Burkill,
the Dusun (Borneo). The
but also
1935).
'juice
be eaten, whereas the fruit wall
of uses, the
species
other crops
(Purwaningsih,
fish
is considered
names
apid, gurah (Bisaya
as
—
and
as
well
as
of the fruits' is
other
an
interesting
crop
species
colours
yellow-green
drying
parts
poison (Dragendorff, 1898).
as
Fruits and
1935).
The green fruits
toothache [NGF (Floyd) 8039]. The seeds contain a
tain toxic latex, also known
Vernacular
and is
fever and
cure
with clothes
(e.g.,
by
(Ng, 1991).
canoes
are
applied by
oil (Burkill,
of the
plant
In view of the
for
places
used
con-
variety
too wet
for
1991).
Thailand: kula. Peninsular
Malay),
Malaysia:
keboean. New Guinea: Irian
gurah.
Jaya:
fa
Borneo:
apid
(Asmat), farid,
sakottoebo; Papua New Guinea: dordi (Kinomeri).
Map
S.
5.
Distribution
of
Shirakiopsis
virgata (Merr.) Esser (�).
indica
(Willd.)
Esser
(•),S. sanchezii (Merr.)
Esser
(■),
and
188
BLUMEA
Notes
—
1.
Hamilton (1837)
tion, transmitted it
to
S.
s.
n.
explained
Willdenowunder the
that the collections Roxburgh
burgh
Vol.
s. n.
No.
44,
that Roxburgh,
name
Fig.
5.
S. indica,
Shirakiopsis
fruiting plant;
face,
with
thyrse
bud;
to
S.
a—c:
of leaf
was
right
virgata
as
Vegetative
base,
and
lower
that
with
thick
pericarp
&
with
of S.
and
by
septa
c.
Herb. Rox-
isotypes
detail
synonym.
ex
of leaf
must
vascular
partly
fruit;
be
re-
h.
g,
habit
lower
sepals,
mericarp
and basal
h:
of
sur-
d. staminate
fused
strand
Convocar) 10342, L; d—f: Zondag s.n., L;
a.
margin,
Miq.) Esser;
g. sulcate
single
of
earlier authors.
& Moritzi
sanchezii;
with
=
in fact
litorea Rumph.
e. staminate flower with
of S.
his collec-
sanchezii (Merr.) Esser;
glands;
virgata (Zoll.
are
homotypic
Ichthyoctonos
characters
surface,
pedicellate flowers;
(Mendoza
a
had been done
stamen; g & h: fruit characters
PNH
sent
It may be, therefore,
(BM, BR, G) and Wallich Cat. 7963 A
d—f: flower characters of S.
dehiscence
(‘triangle’) [a—c:
233, L].
not
indumentum
f. basifixed
septicidal
certainly
Esser.
b. detail
gland;
with
whom he
(K, K-WALL, P), both labelled as Sapium bingirium,
2. Merrill (1917)
to
to
Sapium bingirium.
indica, and Sapium bingiricum of Baillon (1858)
ferred
1999
1,
late
after
split
Kartawinata
H.-J. Esser: A
2.
Sapium sanchezii Merr., Philipp.
Trees
revision
sanchezii (Merr.) Esser,
Shirakiopsis
IV. 147.vii
partial
Sci.,
J.
Biblioth.
(Merr.) Kruijt,
146
Bot.
Flow.
Fig. 5a-c, g-h, Map
—
Pax & K. Hoffm. in
PI.
Is.
44.
(1983)
—
Lex.
Shirakia
Philipp.
sanchezii
Bolster 373
Lectotype (proposed here):
—
5
Pflanzenr.
Engl.,
(1923) 461; Salvosa,
2
Euph. Philipp.
93.
(1996)
nov.
(1913) 406;
Philipp.
Enum.
(1963) 123; Airy Shaw, Alph.
comb.
Bot. 7
Enum.
(1914) 423; Merr.,
189
of the Hippomaneae
(UC),
Philippines, Mindanao, Prov. Surigao, Surigao.
Tree, up
mm
1.5-2
to
15
to
1.5-2
by
entire
mm,
blade
glabrous;
diam. and
1-2
c.
ciliate. Leaves:
to
5-10
glands
side, 0.3-0.4
per
above base of blade,
mm
4
mm,
details
axis
pilose.
fruits:
only
Bracts
of
staminate
mm
in bud.
long
distinctly pedicellate; calyx
when young,
dry
to
mm
Seeds
long.
c.
1-1.5
when mature;
8
by
4
c.
5.5-6
veins 17-25
15-19
mm,
pairs, angle
studied in bud, these
0.75
c.
c.
long.
0.5
by
not
by
17-20
mm
2.5-3
at-
1
mm
with
c.
20
further
long
in
studied, but according
to
mm,
c.
mm
15-30
mm
long;
deeply sulcate, fleshy
3-6
long, stigmata
mm
pilose,
mm,
pedicel
Fruits:
2.5
rounded
slightly
midrib, lower surface almost
sparsely pilose; pedicel
mm
style
long, sparsely pilose
cm
diam., basal glands 0.5-0.7
mm
cymules
mericarps separating easily, glabrous; pericarp
7-8
on
Pistillate flowers
circular in outline,
schizocarp nearly
cm,
secondary
visible in bud. Staminate flowers
not
bud; calyx 0.6-0.7
Bark
yellowish.
to
base obtuse, often
Inflorescences only
midrib 60-75°, smaller veins distinct.
by
by
0.6-1.3
petiole
2-3.5
acuminate, upper surface pilose
and with 0-3
glabrous
pale
deeply fissured; inner bark brown, paler inside. Stipules
elliptic,
ovate to
acute to
tenuate, apex
Indumentum
high. Young twigs pilose.
m
thick, somewhat rough,
mm
long;
thick; remaining columella
the back, brown, caruncle absent
on
inconspicuous.
very
Distribution
Endemic
—
Malesia:
to
Philippines (Mindanao),
Lesser Sunda Islands
(Sumbawa).
Habitat &
forest
on
Ecology
ridges.
Vernacular
—
In thickets and forests
names
Note
only
monsoon
Sumbawa: k.
tanduk; Philippines:
Flow. PI. 2 (1923) 461; Salvosa, Lex.
Philipp.
The collections studied from Sumbawa differ from the
separation
being
can
18-19
Philipp.
mm
of 15-17 mm).
long (instead
Philippine plants
Obviously
no taxo-
be corroborated.
& Moritzi
virgata (Zoll.
Shirakiopsis
ex
comb.
Miq.) Esser,
nov.
—
Fig. 5d-f,
5
Map
Excoecaria
virgata
(1863) 123;
Java
the seashore, in
June.
123].
in their fruits,
nomic
3.
—
or near
May,
Lesser Sunda Islands,
—
bantiano [fide Merrill, Enum.
Trees (1963)
along
Flowers and fruits collected in
12
in
Zoll. & Moritzi
DC.,
15,
(1910) 613; Koord.,
499; Airy
Kew
Shaw,
Miq.) Hook.f., Fl.
K. Hoffm.
in
Miq.) Kruijt,
L, P; G-DC,
Tree up
Prodr.
to
40
Engl.,
m
Add.
5
dbh
up
Young twigs pilose, especially
Ser. 4
Fl. Ind. Bat.
(1996)
(1912) 506;
(1912)
93.
—
60
cm,
(1859) 416; Miill.Arg.,
114.
—
Handl.
249.
& Valeton.
Backer &
Bijdr.
Bakh.f.,
—
Fl. Ned.
Shirakia
Ind.
3,
1
1
(holo U;
32
(1964)
& Moritzi
(1900) 295;
virgata (Zoll.
3035
Linnaea
Boomsoort.
Fl. Java
Sapium virgatum (Zoll.
Type: Zollinger
2620/1), Java,
to
2
1,
J.J. Sm. in Koord.
(1975)
IV.147.V
146
IDC microfiche
high,
Miq.,
(1888) 471; Boerl.,
Pflanzenr.
Bot.
ex
(1866) 1216;
Exkurs.-Fl. Java 2
Brit. India
Biblioth.
n.v.,
Bull.
2
& Moritzi
iso A,
ex
Pax &
ex
BM, G, K,
Bantam.
branching
5
m
above base. Not deciduous.
in the axils. Indumentum yellowish
to
pale.
Bark brown.
190
BLUMEA
inner bark
vertically fissured;
0.8-2
petiole
cm,
cm
long, pilose
base obtuse, often
lower surface
sparsely pilose
midrib 60-65°, basal
pair
blade
glabrous;
pilose.
or
only
glabrous
to
mm
with filaments 0.3-0.5
0.4
c.
in
elliptic
with
4
long.
mm
1.5-1.75
9-10
Habitat &
whole year
Ecology
flowering, nearly
(or 2)
per
thyrse
mm
when
—
pericarp
brown
to
to
8-15
long; calyx
mm
long; schizocarp
mm
slightly sulcate,
with
fleshy
outer
wall 2-2.25
mm
style
2-4
layer
mm
thick. Seeds
without caruncle.
cream,
Grows in
primary
Java: kisereh;
—
stamens
Malesia: Java.
fmits collected in
name
Staminateflowers
blackish and wrinkled;
long,
axis
mm,
long, ciliate;
both ends,
acute at
mm
mm.
mm
inter-
absent in bud, anthers 0.25-
pedicel
becoming
18-21
fide Backer & Bakhuizen
July,
dry,
margin,
whole surface
on
absent; pedicel 4
or
studied. Fruits:
green but
leaf
to
40-70 by 5.5-6
0.5-0.7
long; calyx
when
16-18
of
2.5-4
nearly glabrous,
per side, 0.25-0.4 mm
long, entire, pilose
mm
and disturbed coastal forests
inundated. Soil: loam. Altitude:
through;
Vernacular
mm
layer,
Endemic
—
plains, periodically
collected in
pale
mm,
Distribution
glands
Inflorescences
tardily dehiscent; mericarp
dry, woody part
5-5.5
by
long
outer
and
long; hardly
thick when
up
mm
ciliate. Leaves:
nun,
elliptic, (5—)7—13 by
to
1.25-1.75 by 0.6-0.7
glands
1.5
to
23-31 by
outline,
1-1.5
by
and with 2-10
1.25-1.5
long, entire; style not
leathery-fleshy
mm
their
Pistillateflowers 1
mm
2
c.
oblong
diam., secondary veins (15—)19—26 pairs, angle with
ofstaminate cymules
sparsely pilose; pedicel
1999
with smaller angle, therefore parallel
margin (ciliate),
on
1,
attenuate,apex acuminate, upper surface
secondaries distinct, smaller veins distinct.
Bracts
No.
44,
pale yellow. Stipules
to
slightly
diam., basal glands 0.4-0.7
Vol.
sea
level up
den Brink Jr.
van
to
550
on
m.
alluvial
Flowers
the
(1964) flowering
Apr., July-Sept.
see
also
Smith
(1910)
and Pax & Hoffmann
(1912).
Uses
The wood is durable and
—
Afriastini 3421).
S.
indica and
Notes
uses
listed
1.
Smith (1910) and
two taxa
are
not
Airy
although they
sufficiently
2. The first citation of the
was
be used for tools and houses
can
by Dragendorff (1898)
for India
(McDonald
probably
&
refer
to
are erroneous.
of S. indica,
variety
of the
—
The
Shaw (1975)
formally
never
distinct
to
species by
recognize
Baillon
regarded
S.
them as distinct
(1858,
as
to
be
only
a
species.
Excoecaria
information and does
any additional
accompanied by
virgata
transferred it. However, the fruits
not
virgata Zoll.)
constitute
a
valid
publication.
3. Miiller Argoviensis
bly
was
erroneous
and the
specimen
flowering
cannot
one.
(1866)
cited S.
virgata
from Moulmein, Burma, which
(Hooker, 1888). The species has
by
seen
Miiller
Because the fruits
be identified
as
S.
never
Argoviensis (G-DC,
are
typical
for this
proba-
been collected outside Java,
IDC microfiche
species,
2620/3)
is
a
the mentioned specimen
virgata.
7. STILLINGIA
INTRODUCTION
The genus
Stillingia
was
Garden and named after
Because the
generic
established
by
Linnaeus
Benjamin Stillingfleet,
limits within the
an
(1767),
English
Hippomaneae
based
on
the
authority
of
botanist of the 18th
century.
have
always
been
a
matter
of
H.-J. Esser: A
dispute,
for
assigned
to
decades
some
many, very
Bentham
(1878,1880)
and
cing regularly
forming
a
the
of all parts of the
species
and
(1858
Bertero
Stillingia
of
species
S. thouarsiana
Stillingia by
the
wooden structure, the
peculiar fruits,
ready
never
Stillingia
Of the three
should
Baill.,
described it under the
accepting
The
as
was
proposed by
two
distinct
of
of the
geography,
not
of S.
apically crowded,
plasticity
in the
only
been
e.g.,
vary
on
had alwas
has been discussed
pacifica
distinction of
(1966)
and
the fruits,
Van Steenis,
only
1966). The
species
one
supported by
Coode
with
(1982),
to
two
seems
this view,
is without doubt. It is
similar
very
Malesian
subg. Stillingia,
as
least 3-flowered staminate
subgenus
to
noted above.
fits in
species
at
as
in its series
because of its succulence
OF
MALESIAN
lineata
it has welland
cymules
Dichotomae D.J.
Rogers
characters of
(the palynological
STILLINGIA
succulent (a
character in the
rare
of leaf
shape
not
and index, noted
unique
when
conspicuous
by
compared
Hippomaneae!)
leaf
Van Steenis
scars
(1966)
with
(Fig. 6a).
and Coode
with other members of the
Hippo-
Microstachys.
setae.
the
are
non-succulent leaves and very
plants
The blades bear
of blade and
otherwise
(1942)
conspecificity
studied).
The leaves of Malesian
junction
A
Hippomaneae
and within the
is remarkable but
glandular
two
pacifica,
characters
stems
maneae,
S.
species,
Croizat
but their
notedbelow, S.
as
status.
Van Steenis
CHARACTERS
The
genus,
one rare
(1963).
Shaw
Sapium plumerioides,
pistillate flowers,
Pachycladae Pax)
S. lineata have
(1982),
only
one,
the other
its distinction from the very similar S. lineata from Mauri-
Stillingia
carunculate seeds,
The
from the
species.
Within the genus, the
developed sepals
Vegetative
by Airy
They
and the Mascarenes,
Madagascar
separated
Most
(1912).
have been revised
it should be noted, however, that Smith (1978, 1981) objected
position
sect.
and
diagnostically important
Pax & Hoffmann
the New World.
In Asia, there is
Sapium, distinguished particularly by
(=
be
probably
tius, and therefore its taxonomic
preferable;
are
(Airy Shaw, 1963, 1972b; Smith, 1978, 1981;
of opinion
subspecies,
plant
and is established in the present revision for the first time. Because
recognized
several times
to
from
species
name
of its rareness and remarkable
only
the
on
The South American genus
carpophor.
presented by
was
restricted
are
from Malesia for the first time
matter
the upper part dehis-
A. Juss., however, has very similar fruits but naked staminate
ex
conspecific (Coode, 1982).
reported
were
name.
defined
complete monograph
by Rogers (1951).
may be
tropics
considerable part
a
too.
The latest
of the
united
later)
flowers, and therefore the large, fused sepals of Stillingia
for
191
Hippomaneae
being caducous, the lower part instead remaining
(2-)3-horned,
Adenopeltis
of
different
Baillon
Stillingia. Especially
of the whole tribe under this
revision
partial
same
are
membranous and
petiole (Fig. 6c)
plant; they
comparable glands
serrate
marginal glands abaxially.
are
of
are
usually
Sapium.
often
not
The
not
with
very obvious
separated
texture
mostly persistent
The adaxial
or
glands
on
the
absent, and may
from the blade, unlike the
and distinct serrulation of the
192
BLUMEA —Vol.
leaves, together with the
the
are
1,
1999
and the
lacking marginal glands
never
characters
only
No.
44,
visible venation,
easily
the African
the Asian from
distinguishing
of
specimens
S. lineata.
Floral characters
The inflorescences
bracts bear
but
a
of
pair
are
unbranched, which is
quite large,
not
The flowers of both
uniform,
as
is
be
to
sexes
latter
large,
in
dry,
into three
sessile fruits
structure
are
typical
resembling
carpophor, carpidiophor
pistillate
is
Stillingia
very
(Esser,
or
a
woody
al.,
et
are
to
at
are
anthesis,
are
3-carpellate
and sessile.
base remains
the
on
(Fig. 6g)
regularly
plant, forming
and named
the
gynobase,
AND
RELATIONSHIPS
and Falconeria,
as
was
shown in
a recent
1998). In particular the staminate thyrses and leaves
aboutthe
exact
Rogers'
close
best
are
Neotropical
genera, like
are
their fruits. In
separated by
Senefelderopsis
or
with
only
one
7
(1951)
species
and may be
phylogenetic
namely
and is
a
to
the
stems
sufficient; because his revision concentrated
Dichotomae
on
only superficially,
diagnosis
and
a
never
Remarkably,
not
marginal
to
species
only
to
been studied
Rogers
not
treated
be established
cannot
most
the
or
of the Dichotomae is
phylogeny
in the New World is S. uleana. It differs from S. lineata
and the succulent leaves with distinct
has
pollen
two
peculiar
subsp. pacifica),
the North American taxa,
without re-evaluating his work. Within this series, the
marginal,
cited
succulence and
lineata
variable character. The
(1951). Therefore, his
species by Rogers
(S.
The
hypotheses.
monophyletic. Rogers
lateral pore. The succulence, however, may be restricted
trapezoidea Ule)
tropical
lacks any
characters for them,
leaves (S. uleana Pax & K. Hoffm.),
both (S.
of S. lineata within the genus are hindered
relationships
revision
putatively unique (autapomorphic)
stems
pedicels
various authors.
by
Sapium
comparably
some
Dichotomae consist of 6
the
quite
are
1994).
the fact that
in all
and
cymules usually
with short but distinct filaments; the
three-horned cupule
coccophore
related
closely
study (Esser
Hypotheses
pollen
staminate
flowers
similar in these three taxa, and the
genera
addition, there
by
The
for the genus. The distal part dehisces
BIOGEOGRAPHY
very
observed
glands
Stillingia subg. Stillingia
two stamens
but the thickened,
mericarps,
remarkable
cladistic
semi-cylindrical,
of these
shape
sessile when young, but have short
are
anthesis. The
at
for
typical
Hippomaneae.
2-lobed calyx and
visible
and
The
Hippomaneae.
to
characters
Fruiting
The
a
only
are
length
disc-shaped
taxonomically significant.
are
expected
5-7-flowered. Their flowers
each with
be
seem to
in the
common
least when dried
The variation in
cyathiform glands.
never
in S. lineata does
at
similar to S. lineata
in the
non-succulent
serration and basal
glands purely
adaxial-petiolar.
there is
a
clear distinctionin the distribution between series Dichotomae
and the remainderof the genus,
as
The other four series of the genus
recognized by Rogers (1951)
are
all
strictly American,
for the New World.
confined
to
temperate
or
H.-J. Esser:
and the
subtropical regions
but
they
however,
in succulents),
the
of the
margin
the
of
Mascarenes (Mauritius, La
elevations in the Andes,
higher
tropics.
dry
193
Hippomaneae
or
tropics
core
known from the
truly tropical, only
are
revision
partial
absent from the
completely
are
A
All
species
of the Dichotomae,
of NE Brazil (a
areas
Reunion),
some
rich
region
scattered islands
in
Malesia, and from Fiji.
Stillingia
lineata has
which
(Map 6),
several other
is
with
taxa
limit for Australasian
for the
scattered distribution in the Mascarenes, Malesia and
a
quite
remarkable but
a
is
because Malesian Stillingia is
the
dispersal causing
related genera
Stillingia
Garden
Garden
Stillingia
(1824) 49;
Pflanzenfam.
in
180;
Engl.
Bot. Gard. 3
Gard.
510,
ex
81
5
3,
&
122.
—
Mull.
to
Monoecious.
site in
some
than blade,
in
stipitate
usually
not
or
with
some
of
1
sepals;
hardly
long
a
to
visible.
ovate
mm
3
48
498.
to
2,
with
above
Juss., Euphorb.
in
Engl.
Pflanzenr.
D.J.
ex
—
Sapium
&
Missouri Bot.
Euphorb. (1858)
Stillingia (Garden
sect.
Garden
Type: Stillingia sylvatica
some
New World
a
Inflorescences
to
same
thyrse,
transversely
thyrse.
species);
(less
than
short
pair
(less
of
mm
than 2
conspicuous
on
serrate
the
long,
often divided into
long),
cm
to
much shorter
inconspicuous
or
with
a
0.5-3(-5)
pair
glands
towards the
terminal
or
not
Staminate
with
a
mm
of glands
nor
on
apart,
the
or
often
junction
papillate,
with
or
margin,
smaller veins reticulate
axillary, yellowish, simple, pistillate
pair
cymules
when
is
sometimes
different, secondary veins
very variable in size. Bracts
ovate,
(even
with the blade that
junction
with teeth
basal
slightly longer
15)
L.
species perennial
of
cup-
ofstaminate cymules
to
disc-shaped
glands
5-13-flowered (1-3-flowered in
bracteoles present, filiform. Staminateflowers
filaments
ex
with leaves. Indumentum absent.
less than 2
glandless
joined
not
Nat.
(1912)
Ann. Missouri
5; Ann.
Etude
Gen.
Prantl,
IV.147.V
Rogers,
f.
Baill.,
below smooth and neither whitish
prolate, tricolporate
several
closely
A.
(1967) 388,
1155.
—
Pax
Engl.,
Klotzsch
(1866)
species) glands
—
flowers
2
acumen,
long) pedicel only present
stamens
postulate
all
Leaves alternate, sometimes apically crowded (oppo-
species); petiole
truly entire,
the axis of the
spheroidal
because
(1767) 637;
(1931) 198;
fruiting twigs
strictly marginal glands,
New World
to
15,
(1903)
Dichotomae), margin
never
and staminate flowers in
(up
and
apically
distinct, arching but mostly
touching
19c
2,
,
petiole (see there),
triangular
landmass. But
could
one
separate from the blade; blade elliptic in various proportions,
a row
but often
ed.
in series Dichotomae
New World
some
shores,
(1880) 334;
Hoffm. in
Eustillingia
Prodr.
DC.,
Flowering
New World
indistinct but
without
in
& K.
2
12,
3
J. Arnold Arbor.
sect.
basally glandulous.
succulent (series
with the
Webster,
Lex. Gen. Phan.
(succulent
glandless
Pax
61, 62;
Stillingia
Stipules ovate-triangular
several ciliae,
f.
G.L.
Arg.
Nat. ed.
Gen. PI.
Hook.f.,
Nat. Pflanzenfam.
Kuntze,
herbs).
trees
&
(1890) 96,
Shrubs
geological explanation
distribution, especially
(1767) 19; Syst.
Benth.
Harms,
inval.;
T. Post &
eastern
restricted in their distribution.
more
Mant.
in
(1951) 207;
(1994)
nom.
L.)
L.,
ex
at
the
as
L.
ex
Benth.
wide
Fiji. Fiji
to a
Fiji
mentioned
(1966)
the old Australian
predominantly growing
remarkably
much
are
and this could point
not rare,
distribution correlated with
present-day
Steenis
between Mauritius and
disjunction
taxa
Van
unique.
not
flowering; calyx
than anthers. Pollen
with 1 lateral pore
with 2
with short
largely
fused
1951)
(fide Rogers,
(series Dichotomae).
Pistillate
-
at
base of staminate
sepals; sepals triangular, free, glandless (absent in
thyrse, nearly sessile; calyx
some
New World
species);
with
ovary
194
BLUMEA
3-locular (2-locular in
tinct, stigmata 3 (2 in
No.
44,
1999
1,
few New World
species), smooth; style usually
short but dis-
few New World
species), undivided, glandless.
Fruits
a
a
Vol.
nearly
sessile; 3-seeded (2-seeded in few New World species), smooth, dry, dehiscing regularly
the septa in the upper part, but with
along
the
plant
and 1
form
to
a
2 vascular strands visible
or
thickened, woody lower part remaining
a
at
(2- or) 3-lobed, woody coccophor; mericarps with persistent septae
each septum. Seeds
on
with
elliptic, dry,
without
or
caruncle.
Distribution
Series Dichotomae,
and three in
Brazil)
strictly American, ranging
instead, includes four species
Mauritius,
Sunda Islands, Moluccas,
Habitat &
Ecology
—
open woods, scrubland,
on
La Reunion,
Note
Argentina
in S America
and
Fiji
to
well
(as
Kansas, USA.
of NE
(dry regions
few islands in Malesia: Lesser
a
Philippines.
Preferably growing
under xerophytic conditions,
roadsides, semideserts,
The
—
vague,
boundaries in
species
hybridization
at
least the
rocky hills,
often
Paleotropical species
lineata (Lam.)
Stillingia
lineata
Stillingia
Pflanzenr.
2
Shrub
small tree, up
to
in
?iso
to
Prodr.
DC.,
(1912) 183; Coode,
597/1;
Linnaea
G, L, P, P-JU),
12
a
hardly
(3 —)5
apex
pair
same
diam.
junction
orbiculate
to
below with 0-17
16-26
115.
(1863)
to
are
very
Stipules
elliptic
a
pair
of
indistinct
to
Bracts
cylindrical,
Staminate
c.
of staminate cymules
1.5-3
by
1.25-2.5
1-2
mm,
to
indistinct,
mm
long; style
mm
long.
up
long;
stamens
1
mm
in diameter. Seeds 5
long, stigmata
—
by
4-5
mm,
long,
0.3-1.7
absent and
c.
0.5
mm,
long, glandless
mm
on
varying
or
the axis and
long.
by
cm
c.
5
partly
pedicel
long
1.5
when
mm
to
the
long,
mm
the
greenishflattened-
below the bract.
0.2-0.5
mm
flowering,
long;
Fruits 5-6
apart,
even on
mm,
glands disc-shaped
mm
mm
the junction
towards
indistinctly joined
thyrse; calyx
mm
Bark
mm.
0.3-0.75
side, 0.5-0.75
per
Staminateflowers:
1.5
cm
by
with teeth 2-3 (-4)
their
with filaments 1
Pistillateflowers 5-9 per
to
sulcate, apically retuse; pericarp
Distribution
mm
diameter c. 6
0.8-1.5
cup-shaped glands
glands
mostly along
cymules 5-9(-15)-flowered.
0.75
(holo
s.n.
slightly obovate, (4-)7-24 by
to
serrate
strictly marginal
distinct
Commerson
Type:
Engl.,
lineatum
Sapium
with the blade; blade membranous and
obtuse, margin
pairs,
—
K. Hoffm. in
17. —
Mauritius.
mm
retuse, above with
to
32
Pax &
(1982) 86, pi.
smaller veins distinct to indistinct. Inflorescences 4-6
margin,
0.4
the
on
that sometimes
veins
secondary
calyx
rounded
petiole
twig,
yellow.
above
cuneate
base
cm,
acute to
with the
glands
are
collected.
rarely
(1866) 1157;
160
apically crowded; petiole
slightly succulent,
to
—8
of
Stillingia
for N America.
by Rogers (1951)
high. Twigs succulent, apical
m
divided. Leaves alternate but
2
15,
Fl. Masc.
smooth, covered with leaf-scars 2.5-3
with
shown
known and
poorly
(1790) 734; Mull.Arg.,
microfiche
as
Müll.Arg.
(Lam.) Mull.Arg.
IV.147.V
Lam., Encycl.
P-LA,
still
are
of the New World series of
some
frequently,
occurs
The Dichotomae, however,
c.
from
seashores.
quite
1.
of which
Thirty species currently accepted, twenty-seven
—
four of the five series)
as
ovary
mm
c.
long,
thick; remaining carpophor up
long;
anthers
to
4
mm
deeply
11
mm
carunculate, brown, dotted.
Mauritius, La Reunion, Fiji and in Malesia
the Lesser Sunda Islands, the Moluccas and the
Philippines.
a
few localities from
H.-J. Esser:
Habitat &
Ecology
A
revision
partial
Often found
—
of
the
195
Hippomaneae
beaches, but also in forests up
on
300
to
m
elevation.
Note
—
The
(Smith, 1978; Coode, 1982).
and La Reunion has firm
and
no
distinction between the
morphological
well established
marginal glands,
to
Shaw
Airy
a.
Stillingia
302,
lineata
map
Engl.,
71; Airy Shaw,
Nov.
2
n.v.;
iso
DC.,
Kew
Bull.
f.
Shrub
3-8
—
Type:
to
small
cm,
index
Bartlett
to
(1.5-)2-4,
below with 5-17
Distribution
—
glands
27
2
15,
IV. 147.v
not
Fig. 6, Map
hardly
differ in
ecological
Type:
(1972) 93;
(1866)
A.C.
U.S.
Kew
Bull.
visible venation
generative
thin leaves
characters.
distinction between the
(1966).
37
Blumea
(1982)
Fl. Vitiens.
Occ. Papers
Allertonia
Expl. Exped. Capt.
1
—
f.
Mus.
20;
(1966)
Stillingia
(1867) 232;
Bishop
(1978) 397,
Wilkes
5
Suppl.
35.
18
Pax
&
(1944)
Fl. Vitiens.
(holo presumably
G
or
Ovalau.
J. Arnold Arbor.
23
(1942) 507; Salvosa,
Lex.
Philipp. Trees (1963)
(holo A), Philippines, Babuyan Group, Dalupiri
10
m
high,
trunk
15
cm
side, secondary
Fiji (several
islands ofthe
Island.
Syn.
nov.
diam. Leaves: blade (4-)7-24
membranousand non-succulent,
per
sufficiently
from Mauritius
6
1156; Seem.,
Sm.,
seems
distinguishable by
Van Steenis
(1912) 183;Croizat,
(1963) 372;
—
15167
tree, up
Bull.
Prodr.
GH, US), Fiji,
Sapium plumerioides Croizat,
123.
Kew
16
151.
—
are
Arg. subsp. pacifica (Miill.Arg.) Steenis,
Pflanzenr.
(1981) 567,
G-DC,
Shaw,
in
pacifica Miill.Arg.
K. Hoffm. in
Mull.
(Lam.)
169; Airy
do
proposed by
Steenis
subsp. pacifica (Mull. Arg.)
they
doubt the
(1978)
inland plants)
vs.
entire leaves with
slightly succulent,
and Smith
(1972b)
subspecies
typical subspecies
whereas the Malesian plants
with obvious serration, venation and glands;
subspecies (beach
The
margin distinctly
by
serrate,
and smaller veins distinct.
group)
and Malesia: Lesser Sunda Islands
(Sumbawa, Babar, Timor), Moluccas (Seram), Philippines (Babuyan Group).
Map
6. Distribution
of
Stillingia
lineata
(Lam.) Müll.Arg. subsp. pacifica (Müll.Arg.)
Steenis.
196
BLUMEA
—Vol.
44,
No.
1,
1999
H.-J. Esser:
Habitat &
plantations
Ecology
poorly
in coastal thickets and
edges,
collected in Feb.,
197
Hippomaneae
known from coastal
only
known. On
forest and
Fiji occurring along
limestone cliffs. Altitude 0-300
on
Philippines:
—
the
of
fruits collected in Jan., Feb.,
July, Dec.;
Vernacular name
revision
partial
In Malesia
—
coral soil, but
on
A
loi
coconut
on
Flowers
m.
forest
on
Fiji
Dec.
July,
fide Salvosa, Lex.
[Tag.,
streams,
Philipp.
(1963)
Trees
123].
Note
(1942) did
and
The type of
—
subsp. pacifica.
neata
Stillingia
was
consider this genus, but
not
and therefore could
Sapium
is
Sapium plumerioides
As
not
of S. li-
typical flowering specimen
a
known for Malesia
not
the
compared
this time, Croizat
at
with Excoecaria
specimen only
match it.
8. TRIADICA
INTRODUCTION
Triadica
described
was
both
species
had
a
Loureiro (1790) with
by
T. sinensis. This concept
different taxonomic
quite
The type of T. cochinchinensis
Argoviensis (1866),
and for
and Merrill (1935). Miiller
sapium,
of its
therefore
pistillate
separated
(1925)
and Merrill
(1912),
mann
publications
case
on
only by
the
in
species
Excoecaria
it
as
distinct
a
about T. cochinchinensis
(1954)
and
on
and
as a
T.
not
to
Sapium
about the
Bentham
first
to
same taxon
the
He
Sapium
as
species
Miiller
on
6.
plant;
Stillingia
b. habit
margin,
lower
flowers with
cited
was
Argoviensis,
insufficiently
as
Gagnepain (1926,
placed
discolor. Miiller Argoviensis
the
species
in
Sapium
fused
pericarp
1624, L].
(Lam.) Müll.Arg. subsp. pacifica (Müll.Arg.)
fruiting plant;
surface,
after dehiscence
with
lineata
of
with
sepals;
with
f.
c.
detail of leaf
serration
thickness;
h.
base,
gland;
fruit, unopened,
carpidiophor;
of medium
and
Para-
E. cochinchinensis,
under
sect.
Sapium
(1863)
e.
upper
detail
with distinct
seed
Steenis.
with
of staminate
woody base;
half of mericarp after
i. carunculate
surface,
[a—e:
septicidal
Van
to
Habit of
a.
glands;
thyrse
the
Excoe-
flowering
d. detail
with
g. central
nearly
columella
and loculicidal
Steenis
was
Triadica, effecting
the combination Sapium discolor. Later he transferred the section Triadica
Fig.
separated
sect.
species.
described Stillingia
(1854)
discuss its relationships.
the
cases
revisions except
ignored
Pax & Hoff-
separate genus named Shirakia. Pax & Hoffmann
cochinchinensis. However, in all
without further divisions)
as
as
species.
Sapium.
these authors
cochinchinense (Lour.) Kuntze which is based
known, and all authors of local
necessary,
repeated by
Kruijt (1996), although
Kruijt, although writing obviously
referred
glandular sepals
was
of
species
was
(1925)
sect. Para-
very different
a
the whole section from Excoecaria. Pax & Hoffmann treated it
sapium, and Hurusawa and Kruijt
the
E. loureiroana,
a nomen novum,
(1935) accepted
Miiller
monographer,
one
Loureiro's herbarium, Moore
described E. cochinchinensis.
opinion
Hurusawa
history.
re-examined
it from Triadica (see below), based
already
Miiller Argoviensis'
was
Argoviensis placed
flowers. In this
Loureiro (1790) had
Moore
two
T. cochinchinensis and
species,
two
unaltered by De Jussieu (1824). Later, however,
accepted
was
of leaf
sessile
of fruit
dehiscence
18184, L; f—i:
De
Voogd
198
BLUMEA —Vol.
caria (1866) and made the respective
ed
to
published,
never
this
was
Shaw (1972a, 1975,1981). Only
and transferred it
Sapium
to
No.
1,
1999
combinationof E. discolor. Hooker (1888)
the classification under Sapium and
cluding Airy
44,
Excoecaria
followed
Kruijt (1996) excluded
the
species
from
The combination under Triadica
again.
Hurusawa, who re-established Triadica (1954;
as
return-
all later authors, in-
by nearly
see
was
below), neglected
species.
described
Miquel (1861)
This
type.
Argoviensis (1866)
most
of
known for
poorly
dubious species of Excoecaria
was
separate from
time. It
long
a
Sapium.
Airy
Midler
by
(which included, in his opinion,
repeated by Boerlage (1900)
Shaw
Malesian
on a
cited
was
and Pax & Hoffmann (1912),
it
(1981) recognized
identical with
as
discolor.
Sapium
In the
T. cochinchinensis and Sapium discolor
present revision,
be identical, this
It is the
making
a name
only indigenous
Gangopadhyay
Ham.,
a
This
Sapium).
but then
as
based
Stillingia (sect. Sapium) lanceolaria,
remained
species
a
species
Malesian
united
(1990)
change
of the
Sapium
from India and the
discolor
'
are
considered to
well known S. discolor necessary.
of Triadica (see
species
'
quite
with
below). Chakrabarty
This effected
Himalayan region.
a
&
Buch.-
Sapium eugeniifolium
remarkable
extension of the distribution.
Triadica sinensis,
united the
Triadica
new
its
as
with T.
species
a
names
on
a
Croton
were
Webster
It is
1753).
indigenous
cultivated in different
names were
Seborium
(Stillingfleetia Bojer,
only
Argoviensis (1863)
followed by all later authors.
sebiferus (Linnaeus,
several
long time,
although
Baillon (1858) and Midler
commercially important species,
of the earth for
generic
remained in Triadica,
always
lectotype.
sebifera (L.) Small; they
is based
sebifera
E Asia. Because it is
regions
instead,
it
(1994) designated
applied
to
Raf.). Together
to
warmer
it, including
two
with the whole of
Triadica, it has been combined with different other genera.
After De Jussieu
(1824),
other, but different genera. It
1858,
1861), Sapium
Hoffmann, 1912),
with
Sapium
although
or
Triadica
was
usually
united as
(Midler Argoviensis,
Excoecaria (Midler
without
generically separate
from
considered
was
1863; Hooker,
investigations
Sapium”],
or
changing position,
columella of the fruit,
type collection of T.
was
(Hooker,
(e.g.,
S.
even
when the
also the
pericarp
cochinchinensis
Therefore, the species
cences
was
usually
are
Stillingia (Baillon,
1890; Pax &
was
united
1880; Webster,
1994,
may show that it (
Triadica)
with Carumbium (= Homalanthus;
a
separate genus, only followed
opinions
has
‘Sapium
separated.
attached
to
T.
not
are
visible.
was
sebifera (Hurusawa, 1954).
baccatum’ with its berries and
sometimes included in Triadica,
species
the central
Because the fruits of the
immature, these characters
monotypic, only containing
as
about the included
a sarcotesta
excluded from Triadica, and Triadica
fairly
although
some-
On the
similar leaves and inflores-
it does
not
develop
a sarcotesta
1888; Pax & Hoffmann, 1912). The position of some extra-Malesian species
rotundifolium Hemsl.), however,
includes
distinct part of
and in the present revision.
In addition to the
other hand,
a
Sometimes it
[Bentham,
varied. Characteristic for Triadica are the seeds with
times considered
be
1888; Pax,
Argoviensis, 1866).
Kurz, 1877). Hurusawa (1954) considered Triadica as
by Kruijt (1996)
to
separate section with
delimitation
infrageneric
Webster admitted that "further
should be
a
more
than
two
species.
still has
to
be evaluated, and Triadica probably
A
H.-J. Esser:
partial revision of
the
199
Hippomaneae
CHARACTERS
characters
Vegetative
The architecture of T. cochinchinensis
some
with
detail
by
one
metres
is
in
occurs
of the renewal
monopodial
(1958)
axes
The
account
'Koriba's model'
as
of Koriba
(Halle
two
sympodially
branches
et
honoured
was
al.,
All
1978).
but the apex of the vertical trunk dies away,
pseudoverticels,
trunk grows
attained,
has been described in
(as Sapium discolor)
(1971).
becomes vertical and continues the trunk;
in
a
zigzag
After
manner.
simultaneously,
grow
as a
and the
a
result the
apical
pseudo-
of several
height
ramification
This may be different from other genera, but data
pseudodichotomous.
appears
and Halle
this type of architecture
naming
ramification
Koriba
are
scarce.
The leaves
Triadica,
not
are
adaxial-basal and
most
useful for
very
the basal
of the leaf
part
nensis is the
also
are
may
only species
blades. The leaf base
in
not
with
'
be
the
leaf
remarkably long petioles,
glands (Fig.
the base of the blade and constitutes
at
Homalanthus,
distinguishing
always paired
are
the whitish
in the
which is
the
quite
(papillate
similar in habit.
other
sebifera (and
show orbicular
two
or
shared
Triadica cochinchi-
species.
whereas T.
ovate-elliptic blades,
Sapium rotundifolium’)
glands
the
7b) and the lower-
mentioned. The reticulate venation is
with
for
important
mentioned above, like
taxa
are
margin (Fig. 7a). Additionally,
with many related genera, but
The leaves
genera,
originates strictly
lower leaf surface
glaucous)
of the
Constant characters for
genus.
nearly globose (not disc-shaped)
of side veins that
pair
recognition
found in related Malesian
Malesian
to
broadly
species,
but
ovate
single
Sapium rotundifolium’.
Inflorescences and flowers
Remarkable characters of the
per
of
of
the
thyrse,
a
large
the
are
comparatively long pedicels
per
species
flower is 2
3
or
of Triadica show
(invariable
quite large
of the
number of staminate flowers per
stamens
The
thyrses
most
flowers and the combination
and distinct
thyrse
in
distinguishing
no
pistillate
numbers of pistillate flowers
pedicels.
The number
other Malesian genera of the
characters in the floral parts
tribe).
at
all.
Fruits and seeds
Besides
some
fruits. The seeds
leaf characters,
are
correlated with it
covered
as
part
by
of
a
an
Triadica is best characterized
pale
to
white sarcotesta,
endozoochorous
central columellaafter dehiscence of the
carps
are
Another
mature
unique
syndrome,
pericarp (Fig. 7f),
by
feature is the
fruit will be still
regularly
fact
that, if
not
all
trimerous but with
seeds of
a
on
the
fruit
plant,
too.
develop,
the
locules. In other related
empty
carpels
of
size.
The fruits of all
species
of Triadica are very similar,
Triadica cochinchinensis may be
testa
the
at
and the septa of the meri-
genera, the abortion of seeds will result in rudimentationof locules and
unequal
Probably
the seeds remain
membranous, very fragile and remain between the seeds
noteworthy
the very unusual
in the tribe.
that is
pale
but
never
distinguished by
white and often
a
even
in
quantitative
characters.
relatively weakly developed
quite inconspicuous.
sarco-
200
BLUMEA
Vol.
BIOGEOGRAPHY AND
Triadica is
most
respect
and
common
Triadica is
unique
Triadica cochinchinensis
the
,
C Malesia, but is
rarely,
of the
phylogeny
A
species
Triadica is
often
a
In
taxon.
long petioles
Malesian
as
separate branch in
a
vious sister
only
are
this
it is
most
similar
only
7).
It is
probable,
1998). It shows that
al.,
Sapium,
but without
(entire
Balakata
an
ob-
leaves with
be noted, however, that the
must
from other
taxa
tribe.
in W and,
The relationships
moment.
et
to
to
small part of the tribe,
a
with
comparison
a
at
group of genera close
a
In this
occur.
from the north.
paper (Esser
and often whitish beneath). It
Hippomaneae
whole, and
be established
a recent
respects,
some
occurs
the SE Asian mainland(Map
only by dispersal
cannot
of the genus have been treated in
species
predominantly Neotropical
a
Malesian taxon,
only indigenous
reached
was
RELATIONSHIPS
Hippomaneae,
most common on
therefore, that Malesia
1999
1,
diverse in E Asia, where all
most
in the
No.
44,
probably
not
is still
regions
monophyletic
lacking.
Triadica Lour.
Triadica
Gen.
Lour.,
Bot. 6
81
1
Fl. Cochinch.
ed.
123.
(1994)
351.
—
—
sect.
Brit. India
5
&
Lex. Gen. Phan.
Kuntze,
237;
48
in
Engl.
(1967)
—
(1888) 469;
&
392.
Harms,
—
sinensis
Lour.
[=
Seborium
Triadica
Pax in
Engl.
Triadica
sect.
G.L.
2
&
(1793) 735,748; A. Juss., Euphorb.
(1996) 7;
G.L.
J. Fac. Sci. Univ.
Prantl,
Nat. Pflanzenfam.
Pax & K.
Triadica
Linnaea
Hoffm. in
Engl.,
2,19c (1931) 201;
in
(Lour.) Miill.Arg.
Ann. Missouri
32
3,
Sect.
Tokyo,
3,
Ann. Missouri Bot. Gard.
Webster,
(Lour.) Baill., Etude Euphorb. (1858) 511;
Webster,
Adansonia
(1863) 121; Hook.f.,
5
Pflanzenr.
f.
Fl.
63;
T. Post
IV.147.V
(1912)
(1890) 98,
G.L. Webster, J. Arnold Arbor.
Prodr.
DC.,
Bot. Gard.
81
1210.
15,2 (1866)
(1994) 123]:
Triadica
sebifera (L.) Small].
(1837) 284,
Hortus Maurit.
Tellur.
2,
(Lour.) Miill.Arg.,
(1903) 498;
Excoecaria
Raf., Sylva
146
Nat. Pflanzenfam. ed.
Lectotype [proposed by
Stillingfleetia Bojer,
Bot.
ed.
(1828) 194; Hurus.,
veg.
Triadica
Stillingia sect.
Sapium
(1790) 598,610;
regn.
Biblioth.
(1954) 315; Kruijt,
(1861)
2
1,
(1824) 50; Rchb., Consp.
(1838)
63.
—Type:
nom.
nud.
Seborium
chinense Raf.
[=
Triadica
sebifera (L.)
Small],
Trees. Monoecious.
although
some
and
Flowering
organs
are
fruiting twigs
erose-ciliate.
Stipules
with leaves.
ovate to
Indumentum absent,
triangular,
0.5-2
mm
long,
undivided, glandless. Leaves alternate, but sometimes apically crowded; petiole 1-6
cm
long,
at
times
on
2.5-7
cm
least half as
the
petiole
long
apex,
wide, base cordate to
acuminate, above with
nate
to
base
(sometimes
more on
the
whitish-farinose, often with
secondary
the basal
blade, usually glandless (but basilaminar glands
as
see
below); blade orbiculate
acute to
a
pair
attenuate,
of
of few
veins distinct, lowermost pair
leaf
margin,
with
a
different
angle
not
staminate part 25-140
thyrse
or
with
slightly
a
some-
slightly
ovate,
to mucro-
pair
by
to
on
the
distinctly
the
margin,
forming
not
with the midrib,
touching
intersecondary
veins
terminal and in axils of uppermost
5-10
of
and pale
from the very leaf base and
without sterile basal
leaves, yellowish,
the axis of the
to
apex obtuse
papillate
submarginal glands
originating
of staminate part,
triangular, acuminate, basally
margin entire,
lower surface
present, smaller veins reticulate. Inflorescences
compound,
elliptic
large, nearly globose-spheroidal glands
petiole apex),
a row
to
region, pistillate
mm.
Bracts
of
spheroidal-cylindrical
decurrent. Staminate
flowers
staminate
at
base
cymules
glands touching
cymules (3-)5-8-flowered;
H.-J. Esser:
bracteoles present,
nearly
fused for the
filaments
partial
long
as
most
with
part
revision
the
with 3, sometimes
spheroidal glands
apically
at
to
(3-6);
glandless
with
one
pale
a
to
with
hardly
to
septal
largely remaining
the central columella for
at
considerable time after
at
and loculicidally;
central columella,
columellaconspicuously
a
3,
long; 3-seeded,
mm
without separate
basally
remnants,
remaining
or
sometimes with
or
nearly simultaneously septicidally
thick pericarp, septa
vascular bundle;
apical
Seeds attached
with
moderately
therefore
mericarps
with
and
dry, opening regularly
mericarps
2
long; calyx
mm
style present, disarticulating
base, stigmata 3, undivided, glandless. Fruits with pedicel 2-15
smooth,
flowering;
stamens
2-10
pedicel
divided sepals,
completely
ovary 3-locular, smooth;
margin;
in bud and when
number of lobes
varying
ciliate.
undivided, margin slightly
than anthers. Pistillate flowers with
longer
201
Hippomaneae
long) pedicel
mm
a
of
the bracts,
as
with distinct (> 2
Staminate flowers
calyx
A
triangle,
alate, persistent.
covered
ripening,
whitish sarcotesta, without caruncle.
Distribution
three or four
Probably
—
E Asia (Cochinchina,
China); only
all in Asia and
species,
in W Malesia
species
one
mostly
distributedin
Celebes and Palawan
to
up
(Philippines).
Note
—
the genus is well-defined
Although
the relations of some
S.
Especially
Hemsl. will
rotundifolium
KEY TO
la. Leaves
b. Leaves
ovate
elliptic,
to
broadly
at
ovate, less
loureiroana
(Lour.)
Mull.
3,
24
in
least twice
as
long
than twice
as
long
in
—
Cochinch. ed.
15,
Shirakia
Boerl.,
Bot.
Map
IV.147.V
1. T. cochinchinensis
. .
wide
as
wide
as
2. T. sebifera
7
1,2 (1790) 610;
2,2 (1793)
ed.
nom. nov.
(1912) 252;
—
S.
(1926) 395, 401; Merr.,
cochinchinensis
Biblioth.
Champ,
Benth.)
ex
Handl.
ex
Benth.,
Mull.
Fl. Ned.
Arg.,
Ind.
(1912) 239; Ridl., Fl. Malay
146
(Lour.) Hurus.,
(1996)
93.
749.
—
—Type:
Excoecaria
Sapium cochinchinense
Moore,
J. Bot. 63
(1925)
Trans. Amer. Philos.
J. Fac. Sci. Univ.
3,
Hook. J. Bot. Kew Gard. Misc. 6
Linnaea
1
Pen.
2
3
(1935) 1960; Corner, Ways.
19, 70; Wyatt-Sm., Malay.
Whitmore,
Tree
C.F.
Fl. Taiwan 3
Hsieh,
Fl.
Euphorb. (1983) 44;
Jones
Vietnam
(eds.),
2,
Benth.)
1
2
PI. Res.
Arg.
in
(1863) 121; Hook.f.,
Pax
&
J. Sci. 30
Trees
17
(1973)
& M.G.
SE Asia
K.
Hoffm.
(1924) 316; Gagnep.
Loureiro
Malaya
in
(1854)
1
1.
Fl. Brit.
in
Engl.,
Lecomte,
(1926) 404; Burkill,
s.n.
Tokyo,
Soc.,
Sect.
(holo BM;
Shaw,
Gangop.,
Kew
Bull.
36
Bull.
Gard.
iso
Add.
14
Turner, Gard.
DC.,
Prodr.
Bull. 47
(1995)
231.
15,2 (1866) 1210; Kruijt,
Lectotype (proposed here): Champion
s.n.
(holo K;
iso
—
discolor
(1888) 469;
3
Bull.
IV.147.V
5
17
4
Malay
(1958)
PH.
't
GH, K), Hongkong.
146
5;
Philipp.
Mannetje
Ho, Cayco
Excoecaria discolor
Bot.
f.
(1975) 192;
Enum.
(1990) 183;
Biblioth.
(1926)
(1971) 57,59,
Ser.
(1981) 341; Alph.
J. Econ. Tax. Bot.
5
Pflanzenr.
(PROSEA handb.) 4, Forages (1992) 252;
I.M.
Sapium
Diet. Econ. Prod.
(1965) 113,344; Halle, Biotropica
129; Airy
—
India
Fl. Indo-Chine
(1940) 276; Koriba,
(1977) 494; Airy Shaw, Kew
Chakrab.
(1992) 355;
Miill.
For. Rec.
Malaya
32
(1900) 295;
394, 395,399, f. 46.2-8; Merr., Philipp.
ex
5
be examined.
Triadica.
to
SPECIES
(1866) 1217,
Pflanzenr.
Engl.,
Fl. Indo-Chine
(1954) 318; Kruijt,
discolor
(Champ,
&
2
7,
belong
to
prove
leaf characters,
to
Cochinchina.
Stillingia
Pen.
Fig.
—
Prodr.
DC.,
Lecomte,
(1935) 241.
Bot. 6
BM),
in
Pax & K. Hoffm.
288; Gagnep.
n.s.
Lour., Fl.
Arg.
probably
some
still have
Sapium
THE MALESIAN
1. Triadica cochinchinensis Lour.
Triadica cochinchinensis
its fruits and
by
included in
species currently
(Champ,
(1996)
88.
—
202
BLUMEA
Miq.,
? lanceolaria
Stillingia
(1981)
342.
Boerl.,
Handl.
Fl. Ind.
Excoecaria
—
Fl. Ned.
?
Ind.
to
tresses.
light
tree, up
grey
0.6-0.7
long;
blade
long,
base
to
high,
crown
a
distant from
pair
ovate to
to
1-1.5
mm
of basal
mm
40
&
iso
in
DC.,
K. Hoffm.
diam., bole up
cm
in
Kew
Shaw,
Prodr.
Engl.,
2
15,
Bull.
36
(1866) 1221;
Pflanzenr.
IV. 147.v
CAL, n.v.), Sumatra, Padang, Poeloe Pisang.
to
to
20
m
yellowish
glands
margin, secondary
long,
their
articulate
high,
to
without butnot
fissured,
orange. Sapwood
1-3
blood red;
cm,
glands
apex obtuse
per side,0.5-1
veins 9-17
pairs, angle
margin,
1-1.5
by
base; calyx
7. Distribution
by
to
acute
7-10
1
mm
of Triadica
mm.
long;
to
as
Bracts
cochinchinensis
as
whitish-papillate
angle
mm
arching
with midrib
ofstaminate cymules
Staminate flowers:
stamens
cm
acuminate,
with midrib 65-75°,
mm.
1-3
wide
diam. and (0—)0.5—1
lowermost pair with
0.5-0.75
c.
mm
petiole
less than half
1-1.5 by 0.5-0.75 mm, lower surface
towards the
glands
Map
rarely obtuse,
staminate part 25-50
near
withering
when young,
to
and with 0-3
Inflorescences:
long,
Pax
(holoU;
elliptic, (2.5-)3.5-7 by
attenuate
only indistinctly joining
30-45°.
stem
(1861) 183, 461; Airy
bare for 1-2 weeks. Bark rather smooth,
pinkish
Leaves
slightly
whitish-glaucous
but
m
549
1999
(Miq.) Miill.Arg.
(1900) 296;
HB
1
Suppl.
1,
white, soft; sapwood and heartwood little differentiated. Stipules 0.3-2
mm.
cuneate
above with
to
25
1
3,
No.
44,
brown; inner bark granular mottled, pale
to
pale yellow
by
to
Deciduous with
Bat.
lanceolaria
(1912) 170.—Type: Teijsmann
Shrub
Vol.
pedicel
2-3
with filaments 0.75-1
Lour.
H.-J. Esser:
Fig.
7.
base,
Triadica
upper
dicels and
partly
dehiscence;
seeds;
h.
(Amin
et
cochinchinensis
surface,
with
fused
f. central
seed
al.)
with
glands;
sepals;
,
L].
and
partial
Lour.
c.
part
a.
revision
Habit
of
of staminate
partly opened fruit;
columella
raphe
123379
d.
A
of fruit with
without
e.
plant
the
with
staminate
with
half of
mericarp
seeds;
[a—c:
SAN
203
Hippomaneae
thyrse
attached
caruncle
of
g.
thyrse;
b.
detail
of leaf
many-flowered cymules, long peafter septicidal and loculicidal
central
(Amin
et
columella
of fruit without
al.) 117787, L; d—h:
SAN
204
mm
BLUMEA
when
long
2-4
pedicel
1-3 mm
style
flowering,
anthers 0.4-0.5
long, stigmata
circular in outline, 7-9
mm,
marginally
sarcotesta
with
Distribution
mm,
long.
Fruits:
with
granitic
Found in
thickets,
sand.
sulcate, apex
not
or
glands;
long; schizocarp
mm
slightly
vein. Seeds 3.5-5
brownish when
hardly white,
—
perthyrse;
basimarginal
2.5-6
pedicel
base often clavate,
Fairly
to
2.5-3
by
Taiwan and E China
and Celebes.
Sarawak), Philippines (Palawan),
primary forest,
hillsides and
on
3.5-4.5
by
dry.
1956), and W Malesia: Peninsular Malaysia (incl.
N Borneo (Sabah,
Ecology
years, and
and
yellow
Pistillateflowers 0-13
NE India, Burma, Thailand, Indochina up
—
Singapore), Sumatra,
forest of 5
long.
thick; remaining columella distinctly alate by 1.5-2.5
mm
(known from eight provinces: Lee.
Habitat &
mm
1999
1,
long, sepals rarely
mm
mm
No.
44,
conspicuously arching
but
pale
2-3
7-9
by
rounded; pericarp 0.3-0.4
mm,
1-2
long; calyx
mm
—Vol.
common.
disturbed forest,
Altitude 10-1000
m.
secondary
young
also in
steep slopes,
dry places.
Soil:
Flowers collected in
Jan., Apr.-May, Aug.-Sept., Nov.; fruits collected the whole year through.
Vernacular
Borneo:
Uses
as
names
't
Mannetje
minor forage
seed
coat
(Aziz,
plant.
is poorer in
& Jones
Notes
—
(1898)
293 refers
Usually
all
from the
guished
to
(under
'
both variants.
(1940)
to
light
as
timber tree,
a
additionally
of great use (Burkill,
not
1935).
and is therefore
sebifera,
However,
statement
a
that it "is
not
The
used
for
(1940)
distinctly
taxonomic
a
collective
&
over
syntypes
separation
species
which had been
two
its whole
slight
not to
of
Revis. Gen.
T. cochinchinen-
with
also
may
be distin-
lectotype) by
the
structure
Stillingia
of the cuticular
discolor
represent
be corroborated. Croizat's
cannot
numerous
united the
local forms"
species
already suggested by
probably
with the Indian
Hooker
(1888);
ofdifferences ofthe leaves very similar
mentioned syntypes.
range,
(including
whitish. The
Gangopadhyay (1990)
the differences between the
Croizat
more
kept separate before because
T. cochinchinensis
(Lour.) Kuntze,
like Pax & Hoffmann (1912) and Merrill (1935)
different. The
eugeniifolium Buch.-Ham.,
had been
cochinchinense
from Malesia and southern Cochinchina
obviously
justified. Chakrabarty
taxa
Sapium
of the northern
of its range
part
plants
ornamentations is
S.
ludai.
Sapium’).
specimens
the lower surface of the leaves
both
species
and
Excoecaria cochinchinensis Lour.,
subsequent authors,
used it for Triadica
not
cite the
(1992)
acid than that of T.
1. The combination
sis Lour. Most
is
Sumatra:
pelandok.
(Ibang).
The wood is soft,
palmitic
mamah
Malaysia:
1987).
PI. 3 (2)
2.
Peninsular
tapang lalat
sengajang,
—
—
Sapium
Considering
the
of
variability
laui from Hainan, described
differences of leaves and measurements, may prove
to
by
be
synonymous with T. cochinchinensis.
2. Triadica sebifera
Triadica
Bot.
4,
Small
sebifera (L.) Small, Man.
6
(1954) 315,
(1753)
ed.
(L.)
4
1004.
—
f.
45; Kruijt,
f.
(1933) 789; Hums.,
Biblioth.
Bot.
146
Stillingia sebifera (L.) Michx.,
(1805) 588; Hassk.,
(1858) pi. 7,
S.E. Fl.
26-30; Miq.,
Cat. Hort.
Fl. Ind.
Bot.
Bat.
1,
(1996)
Fl.
J. Fac.
89.
—
Bor.-Amer.
2
Sci. Univ.
Croton
(1859)
413.
—
Sect.
(1803) 213; Willd., Sp.
Bogor. (1844) 234; Baill.,
2
Tokyo,
sebiferus L., Sp.
Etude
Euphorb.
3,
PI.
PI.
Atlas
Sapium sebiferum (L.) Roxb.,
H.-J. Esser: A
Fl. Indica
[Hort. Bengal. (1814) 69, noraen]
7972; Mull.Arg.,
Econ.
Prod.
Koord.
&
India
Chine
5
2
6,
32
Boomsoort.
Hoffra. in
(1926) 394,
Howes,
Kew
Bull.
Bakh.f.,
Fl. Java
1
Engl.,
Pflanzenr.
(1949) 573;
3
Stillingfleetia sebifera (L.) Bojer,
in
(L.) Mull.Arg.
Burma
2
Prodr.
DC.,
412.
(1877) 411,
15,
—
—
2
Linn.
No.
1140.9
(1858) 512; Merr.,
to
to
tree, up
8
apex acuminate, above with
and with
papillate
no to
and joined towards the
70-140
cences
thyrse.
by
Fruits:
6.5-7
with
mm,
Distribution
—
pair
Soc.,
under
very
a
of glands
mm
to
Uses
uses
in
There
—
is
glands
c.
Growing
—
2.5-4
shade
tree
provide
candle-making
drying
oil
in
1
and
0.75
by
from
to
waterproofing
—
especially
erroneous
black
is
cm
c.
mm.
12
as
mm
30.
—
Herb.
in
s.n.
Loureiro
s.n.
(holo
long;
blade
long,
base rounded,
broadly
pairs, arching
Pistillateflowers 2-6 per
sulcate. Seeds 6.5-8
long,
and
nearly
all
provinces: Lee, 1956) and
regions
to
and sometimes
natu-
in former times also
grown in
Japan,
hardy.
or
2).
note
near
waters, and
thriving
It flowers after three years with
four months for
kirendang (Smith,
ripening (Aziz, 1987).
1910).
especially China, Japan, India,
planted
plantations
to
dye.
ornamental and has
as
of other crops,
naturalize
easily.
The waxy seed
with
linseed oil,
properties
but
umbrellas (Lee,
It is also
coat
yields
similar
also used
as
especially
an
a
the
sebifera
as
the
effects when
species
grows
minor timber tree, and the
tallow' used for
'vegetable
endosperm
to
and the USA (e.g.,
positive
tung oil,
of the seeds
in its
provides
a
drying capacities
illuminant (Burkill,
1935)
and
for
1956).
populneus (Geiseler)
records of T.
Type:
veins 8-10
1. Triadica sebifera has often been confused with
H.
Osbeck
distinctly reticulate. Inflores-
warmer
and soaps in China, whereas the
(Stillingia oil)
comparable
Notes
a
Forest Fl.
(1948)
wealth of literature available concerned with cultivation and
very well and often tends
leaves
61
nomen
sebifera
diam., lower surface whitish-
well e.g. in moist soil
Java: kasoembi,
1950). The species
as
&
sarcotesta.
the fruits take three
—
a
(1838) 63,
Excoecaria
Mag. Tokyo
241.
mm
closely
often cultivated in
outside Malesia,
regions
Howes,
planted
names
F.N.
Backer
2,2 (1793) 749; Baill., Etude Euphorb.
petiole
long; schizocarp
China (known
uses
—
fragrant flowers;
—
(1986) 86]:
(1935)
0.75
c.
wide range of soil conditions. It is frost
Vernacular
Tellur.
than half as wide
cm, more
in the USA, N India,
Ecology
Indo-
1962;
(1950) 961;
1
284.
Indochina, and Malesia (Singapore, Bogor, Timor; compare
Habitat &
(1935)
in
(1912)
Lecomte, Fl.
2
sebiferum (L.) Kurz,
Bot.
172
ed.
24
n.s.
smaller veins
Bracts:
6-15
Native
nowadays
in
Pen.
3,
(1857)
Carumbium
Fl. Pakistan
Diet.
J.J. Sm.
Exkurs.-Fl. Java 2
Raf., Sylva
Hort. Maurit.
—
(1900) 295;
Ser.4(1975) 136, 192; C.F. Hsieh,
sebiferum Hurus.,
white, persistent
a
ed.
submarginal glands, secondary
margin,
Taiwan. For its different
ralizing,
a
few
long.
mm
pedicel
Bull. Add.
Deciduous. Leaves:
high.
m
1
3,
Malay
Indon.
(1847)
Cantonem Sinarum'.
orbicular, 3.5-6.5 by 2.5-7.5
ovate to
Prod.
PI.
1,2 (1790) 610;
Trans. Amer. Philos.
'circa
BM, n.v.), China,
Shrub
Cochinch. ed.
Ind.
List.
(1888) 470; G. Watt,
Canton.
(LINN, n.v.), China,
Lour., Fl.
Nutt.
1210.
(1866)
Lectotype [proposed by Radcliffe-Smith,
Triadica sinensis
Ned.
Seborium chinense
Seborium
India 5
(1912) 237; Gagnep.
Econ.
Kew
superfl.
(1977) 496, pi.
Fl.
IV.147.V
Heyne,
K.
Fl. Brit.
(1910) 613; Koord.,
Diet.
(1964) 500; Airy Shaw,
700.
12
Fl. Taiwan
—
1832,3 (1832) 693; Wall., Numer.
Handl.
Java
395, 398; Burkill,
4
ed.
205
of the Hippomaneae
(1863) 121; Hook.f.,
(1893) 472; Boerl.,
Valeton, Bijdr.
Pax & K.
507;
Linnaea
revision
partial
Pax.
Airy
Shaw
(1975)
species
cites
of
Homalanthus,
some
for Malesia. Homalanthus species may be
examples
of
distinguished
206
BLUMEA
from Triadica by the remarkable
flowers and
mostly
carpels,
two
2. Although
from this
a
numerous
thin and
quite often
region
1999
1,
leaves with percurrent (not reticulate)
3-flowered staminate
glands,
stamens, and
and seeds with
fleshy pericarp,
could be studied
[Horsfield
s.n.,
Timor
The other localities cited
s.n.,
Java (BM);
Java, Bogor Bot. Gard. (L sheet
were
no
with
reddish arilloid.
a
no.
four collections
only
Teijsmann
s.n.,
Java
R. Brown
904.117-515);
taken from the literature.
cultivation in Malesia has been abandoned, and
cymules
very different, smaller fruits with
by
cited in the literature regarding Malesia,
K); De Vriese
(BM)].
No.
44,
large stipules,
different adaxial leaf
tertiary venation,
compressed
Vol.
(P,
s.n.,
Obviously,
the
records from the 20th century
are
available.
SEBASTIANIA
Sebastiania
a
was
described
the first who, in 1866,
the genus,
in 1821 and named after Antonio Sebastiani,
by Sprengel
and botanist of Rome who died the
physician
namely
assigned
a
species
year. Midler
same
from the Old World
S. chamaelea. Later contributions
to
Argoviensis
the Malesian
area
were
Pax & Hoffmann (1912) described another species, S. borneensis, Van Steenis
two more,
notes
on
useful
It is
S.
lancifolia
S. borneensis
on
comments
becoming
is unnatural
and S.
caria L.
et
(Esser,
more
and
al.
(1998).
1994).
by
All other
accepted
e.g., Webster,
The
two
Shaw
as
and
Microstachys),
on
again
even
noted
Sebastiania
by Airy
including
Shaw
as
and Esser
are
ones,
assigned
Euse-
Excoe-
genus,
one
(1980)
all Malesian
formally
('sect.
Paleotropical
should be
which will be
and
circumscription
s.str.
be combined into
they
(1948)
additional
chamaelea(1980),
S.
to
few.
(1975).
and very similar to
different sections;
quite
1974) published
that Sebastiania in its actual
1994). Indeed,
of Sebastiania,
species
whole
a
genera could
(1878)
(1960,
illustrations),
strictly Neotropical
genera ( Gymnanthes and
taxa
more
Bentham
included in distinct and
sian
its first
(including
(compare,
discussed
Airy
Malesian Sebastiania
bastiania Mull. Arg.') is
was
remota.
was
(including Malesia)
to
still
different
effected for the Male-
in the present work.
NOMINA EXCLUDENDA
1.
Sapium
cochinchinense (Lour.) Kuntze, Revis. Gen. PI. 3
(2) (1898)
293
=
Excoe-
caria cochinchinensis Lour.
2.
(Benth.)
3.
Fl. Indica ed. 1832, 3 (1832) 693
Sapium cordifolium Roxb.,
=
Alchornea mollis
Mull. Arg.
Leafl.
Sapium crassifolium Elmer,
Philipp.
Bot. 2 (1908) 485
=
Antidesma bunius
Spreng.
4.
Sapium subrotundifolium Elmer,
subrotundifolium (Elmer)
5.
Stillingia populnea Geiseler,
(Geiseler)
Pax.
Leafl. Philipp.
Bot.
3(1910)
930
=
Blumeodendron
Merr.
Croton.
Monogr. (1807)
80
=
Homalanthus populneus
H.-J. Esser: A
partial
revision
the
of
207
Hippomaneae
ACKNOWLEDGEMENTS
study
HBG, K, K-WALL, L, LIV, NY, OXF, P, P-JU, P-LA, TCD, U, UC,
the
is based
directors
material
on
facilities. The Netherlands
grant (B 85-291),
C.
Jarvis
and the
(BM)
at
and A. Radcliffe-Smith
assistance
comments. The
and
thanks
and manifold
support.
were
(©
produced with
B.
are
due
J. H.
the
to P.
M.M.J,
van
Os is
van
the Latin
for
Name
Typification Project
diagnoses.
Two
responsible
is
anonymous
thanked
Ridsdale,
the
manuscript,
(©
P.
drawings.
Welzen, L)
van
and
a
help with
with
Linnaean
reviewers
for various
C.E.
for the beautiful
to
work with
invaluable
Balgooy,
of the COOR database
help
(NWO) supported the
(LIV) provided
reviewing
van
Welzen
indebted
am
of visits and use of their
Gunn
Rijksherbarium, Leiden,
staff of the
discussions, especially
Very special
Plant
Linnaean
(K) improved
and US. I
possibility
Research
gratefully acknowledged. A.
types,
useful
for loans or the
for Scientific
Organization
which is
types
very
herbaria
curators of these institutions
and
LIV,
of the
A, B, BKF, BM, BR, BRI, DBN, E, FHO, FR, G,
This
GH,
made
support,
J.F.
Veldkamp.
critical
comments
The distribution
maps
and the KORT program
Hansen, C).
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3.
Fleischer,
210
BLUMEA
0.1788.
Swartz,
Trimen, H.
wild
in
de
1885. A
species plantarum
systematic catalogue
83.
Ceylon:
L.-R.
Tulasne,
Skeen,
1999
1,
seu Prodromus:
95.
Swederi, Stockholm, Uppsala
Bot. Gard.
1883.
de las
15:
3,
note
17:
3,
1966. Pacific
Sinopsis
flowering plants
Stilaginellas
Bot. Ser.
Ser.
Buitenzorg,
C.G.G.J.
Steenis,
S.
Nat.,
C.G.G.J. 1948. Provisional
Steenis,
Vidal,
of the
and fems
indigenous to
growing
or
Colombo.
1851. Antidesmata et
eis affinibus. Ann. Sci.
Bull.
Van
No.
44,
Abo.
&
Van
Nova genera et
Vol.
on
novum
recenset
genus
nonnullaque
the
genus
Sebastiania
in
Malaysia (Euphorb.)
409-410.
Plant Areas
2. Blumea
lenosas de las
plantas
plantarum
255.
5:
Suppl.
Filipinas.
302,
Atlas:
169
map
90. Chofre
40, 41, pi.
y ca.,
Manila.
Von
Martius,
C.F.P. &
brasilianische
Von
J.G. Zuccarini.
Pflanzen. Flora 7.1
Martius, C.F.P.
1824a.
Beil.
& J.G. Zuccarini.
Ankfindigung
der
eines
Fortsetzung
Werkes fiber
4: 136-139.
1824b. Nova
et
genera
species plantarum
1:66-71.
Lindauer,
MUnchen.
Von
Siebold,
P.F. &
Akad.
Bayer.
Wallich, N.
J.G. Zuccarini.
Wiss., Math.-Phys.
1847. A numerical list of dried
collected under the
269, 274-275.
Webster,
G.L.
Arbor. 48:
Webster,
Wheeler,
Wight,
Japonicae
familiae
naturales
1. Abh.
Konigl.
145-146.
of plants, in
specimens
Dr. Wallich of the
the East India
company's
company's museum,
Botanic Garden at Calcutta:
London.
of
genera
Euphorbiaceae
in
the
Southeastern
United
States.
J. Arnold
392.
1983. A botanical
Taxon
G.L.
Missouri
superintendenceof
1967. The
G.L.
biaceae).
Webster,
1843. Florae
Kl. 4:
1994.
1975.
of the
Synopsis
Bot. Gard. 81:
L.C.
gordian knot:
the
case
of Ateramnus
and
Gymnanthes (Euphor-
32: 304-305.
genera
and
suprageneric
Taxon
Euphorbiaceae genera lectotypified.
W. 1853. Icones
taxa
of the
Euphorbiaceae.
Ann.
120-125.
plantarum
Indiae
Orientalis,
5, part
vol.
2:
24:
535,
537
5,6,20, pis.
1866,1950.Pharoah,
Madras.
Willdenow,
C.L.
1805. Caroli
Linne
a
Plantarum.
Species
J. 1964. An introduction to forest
Wyatt-Smith,
Identification
Material
of
=
1-2
=
2-1
=
3-1
3-2
numbered collections
baccata
6-1
Balakata
luzonica
6-2
Falconeria
8-1;
=
=
=
Shirakiopsis
indica
Shirakiopsis
sanchezii
Shirakiopsis virgata
bomeensis
7-la
=
Stillingia
lineata
subsp.
7-lb
=
Stillingia
lineata
subsp. pacifica
605:
1-1;
679:
3-1;
1-1;
W 495:
—
3213:
906:
4-1;
—
15334:
Baker
8-1;
15167:
7-lb —
=
8-2
1-1;
1053:
bin Shukor
W 791:
1203:
1-1;
19: 6-1
797:
=
3-1;
3-1—Amherst
—
Triadica
cochinchinensis
Triadica
sebifera
1-1
—
Adams
d'Alleizette
846:
6-1
1398
lineata
=
—
3045:
6567:
Ambriansyah
Wallich
4-1
7-la
7797 H:
—
—
& Arifin
4-1;
Aet
(exp.
Altmann
1718
546:
=
456:
6-1;
Wallich
183: 4-1.
4-1;
18988:
4109: 6-1
3214:
8-1
Ambriansyah
3-1;
1-1 —Andrew
3-1
chamaelea
31:6-1 -—Ahmad
Amann 9: 4-1
Bartlett
Shirakiopsis, Stillingia,
remota
4578:
3031:
Berlin.
Gymnanthes
Microstachys
7966:
Nauk,
14
Gymnanthes
Sapium glandulosum
Backer
III-7:
=
=
122:
572-573.
23,
=
=
—
4:
list
6-3
insignis
5-1
4-1
vol.
cited):
Balakata
Lundquist)
W
(only
4-1
Achmad
4,
For. Rec.
Balakata, Falconeria, Gymnanthes, Microstachys, Sapium,
and Triadica studied
1-1
Ed.
types. Malay.
8-1
—
—
4-1;
19299:
Balakrishnan
4-1;
888:
20681:
1-1
—
4-1
Bangham & Bangham-Masters
Bartlett & LaRue
176: 4-1
—
—
Bacon 212:
Balansa 702:
633:
8-1
bb series 5346:
—
8-1;
4-1
703:
Barber
1-1;
9089:
—
8-1;
Bakar
3212:
6752: 4-1
1-1;
—
11323:
H.-J. Esser:
11553:
6-1;
26437:
35728:
1-1;
1855:
6-1;
BeunwSe
8536:
XV
1-1
4353:
4-1
—
Beccari
—
1168:
6-1;
8331:
10573:
8-1;
40313:
1-1
Shah
168:
—
—
8-1
115:
1
—
1
series
36942:
-1;
1-1
Chevalier
—
Chow &
475:
—
—
1-1;
8-1;
3618:
B:
6-1;
8-1
Collins
—
Mermaid)
Dachlan
281:
HB
2096:
NEP
1-1
8-1
Elmer
4-1;
2382:
730:
2565:
1-2;
20120:
C:
4-1;
22718:
1487:
&
4709:
Haines
—
2040:
5672:
2-1
BS series
2755:
4-1
6-1; 6607:6-1; 7509:
Brooke
—
634
1-2;
BKF
=
Burkill
&
Buwalda
—
8124:
1-2;
—
Brass
—
—
1681: 3-1
8-1;
6-1;
8-1
35078:
Bunchuai
—
1-1;
1087:
8859:4-1
4-1;
12431:
1-1;
al.
et
Chin
et
—
Craib
836:
1-1;
2-1
1605:
—
—
—
4-1;
1-2;
9176:
1999: 6-1
419: 3-1
Clarkson
6786:
—
4-1;
—
Coode
K.S.
H.H.
13710:
8-1;
4-1
—
Coert477:
Congdon
—
—
—
5205:
4-1;
9756:4-1
—
4-1
Clarke
—
7-la
3-1
Craven 3842:
4-1
5541:
1484: 8-
1782: 8-1
3014:
M.S. Clemens
2329:
33: 4-1
12836:
Dickason 6652:
et al. 6722:
3-1;
—
20252:
8-1;
Chermsirivathana
Chuang
—
IV-160:
8-1;
18829:
747:
2-1
& Richardson
3844: 4-1
—
Craven &
Cunningham (2nd Voyage of
6-1.
Darnton
—
174:
1-1
IV-81:
1-1;
Chippendale
8-1
Nangkat
156: 8-1
Wilde-Duyfjes
1-1
&
2324:
—
—
Commerson 657:
—
Coode
—
A:
series
12456:
6-1
78428:
21665:3-1
2372: 6-1
4-1
Cel
—
1-1;
1240:
2152:
al.
38099
4-1;
Cuming
—
7238;
1-1;
6692:
—
6908:
1-1;
6-3
14343:
3-2;
150: 5-1
3-1
1584:
Vogel
13767:
Djumadi
—
3-1;
De
1-1;
6-1
De
—
Voogd
15911:
3-2;
Dobremez NEP 682:
—
Drummond
3-2;
—Diepenhorst
26356:
2-1
2-1;
Duthie
—
1-1.
Eberhardt
1071:
Falconer 837:
13232:
1-1
1438:
8-1;
3282:
1-1; 1438:6-1;
—
Fan & Li
17601:
1-2;
30157:
1-1;
1-1;
—Fosberg
4892;
8-1;
1488:
6-1;
8-1
89 E
—
Edgeworth
1 P 763:
222: 2-1
1-1—Evrard
1-2;
118: 8-1
18289:
Fung
2831
1-2
—
—
50224:
2-1
Fox
—
12716:
857:
1-1;
1-1
338:
19153:
9;
Fernandes
1-1—Forbes 2776:
20194:
FB series
18784:
32419:
1-1;
A:
9;
2-1;
23431:
Santisuk
2-1;
5502:
—
1713:
19808:
5714:
66990:
1328:
2-1;
3630:
1-1;
—
5054:
2-1
1-1
1-1
2-1;
4711:
—
6518:
2-1;
2-1 —Feuilletau
2788:
1-1
—
—Foxworthy
6817 A:
6-1;
—
—
Giesen
6-1;
7559:
2-1;
27537:
4-1
—
2-1
2-1
—
6-1;
Haniff
—
—
Guppy
947:
—
3709:4-1
6-1
2-1
—
& Santisuk
1991:
20003:
de
Bruyn
Forman & Blewett
9934: 6-1
1-1;
5927:
Griffith 704:
6-1
1076:
10:
1-1
—
6-1;
—
—Fried-
—
706:
6-1;
4-1;
8653:
20864:
308:
2-1
—
6002:
1-1
—
Goodenough
1161: 4-1
1-1;
—
1-1;
Grierson
4706:
Gwynne-Vaughan 297:
1108: 6-1
Harmand 4:
Henry 11569:2-1;
1-1;
4481: 7-lb
Gillespie
239:
7765:
14595:
Gaudichaud
8-1; 913:4-1; 1097:8-1;
4577:
2-1;
(Borneo)
5905:
463: 8-1;
51:
Gressitt339:
4712:
Hallier
27035:
2-1;
Geesink, Phanichapol
4299:
1-1;
Handel-Mazzetti
Haviland & Hose
—
1-2;
1-2;
8-1:20308:4-1.
24064:
6-1
1080:
1-2;
1-2;
1-1; 10114:6-1;11127:2-1; 11169:4-1; 13827:4-1; 14486:2-1;
6-l;4710: 1-1;
500:
—
8-1;
4-1
227:
1-1—Grashoff752:
10136:4-1
4-1
11:
—
& LaFrankie
Long
7659:
1-2; 20628:9; 22955:1-2; 29251:
9887:
Gentry
2-1
—
Dransfield
1101:4-1
&
4-1;
1-2 —Endert
1-2;
4-1;
Geesink
1623:
21303:
1-1;
A:
3111:
Djimat Tetong
—
—Fleury
Gamble 2830
4257:
—
13812: 6-1
3-1;
Bon
—
Blake
—
IX C 96:
5-1;
—
18954: 4-1
4-1;
443: 3-1
Burkill
—
9
6-1
Brandis 3044:
—
18953:
4-1;
series
Church, Mahyaretal.
Curtis
1179: 7-la —H.
mann
1-2;
12075:
8-1
4-1;
Dalziel
18: 2-1
—
1-2;
861:4-1;
1-1
IX C 80:
234:
Beguin
14788:7-la
7-la;
42017:
1-1;
373:
—
1-1;
35344:
8-1;
4-1.
4-1
244:6-1
—
25007
3921:
—DyPhon
12630:
3-1
K.S. Chow
—
al. 7325:
et
—
—
—
1833: 2-1
3774:
4-1
18845:
Eardley-Wilmot
37529:
De Wilde & De
—
18260:3-2;
series
1-1;
865: 6-1
6-1;
2322: 6-1
1624:7-lb
1096:
4-1; 7337:4-1;
6925:
2781:
1-2;
Bolster
—
Tukirin et al. 445:
609:
8-1
6-1;
Cooper
—
Schodde 787:
Fagg
516:
Coode, Ferguson
5064:7-la
4-1;
TB
BRUN
—
5-1;
IX C 79a:
7-la
18893:
4-1;
4-1
1002:
Burley,
1-1
4-1;
J. & M.S. Clemens 3006:
—
18415:
CCC
8-1;
80071:
20624
2716:
16920:
8-1;
Chantaranothai, Chayamarit
32270:
P.P Wan
Chung
T 843:
3556:
1558:
—
2780:
144: 8-1
8-1;
CF
1388:
40313:
6-1;
7443: 2-1
14588:
7-la;
25618:
6-1;
34183:
6-1;
2669:4-1.
—Byrnes
Cantley
1-1;
25012:
71: 6-1;
IX C
Beddome
—
24568:
1-1;
33883:
6-1;
Bernardi 5:
—
Boivin
BSIP series
1-1
1108:
6799:6-1—BW series
1-2
—
Brown
Bunnemeijer
—
175: 2-1
Garden
6-1; 18403:4-1;
3-1
PB 3127: 3-1
6-1;
22452:
6-1;
211
Hippomaneae
32653:
1-1;
BKF series
5-1
T & P 519:
83440: 9
4-1;
—
6-3
XX C 71:
6-3;
1199:
the
of
22099:
1-1;
32183:
1-2;
Bdjeaud
Bogor Botanical
4-1;
44473:
4490:
6-3;
Boschproefstation
8156:
PB
—
revision
partial
15348:
8-1;
29911:
1-1;
1948: 6-1
6-1;
B VI 8:
j
14946:
8-1;
28956:
1-1;
A
11946:
1 -1
—
Hance 878:
1-1;
—
307:
2-1;
8-1;
6-1.
8-1;
653:
Herb. Hermann
212
BLUMEA
vol.
3, 26,
4-1;
831:
TFB
1-1;
8-1;
1268:
2956: 6-3
70:
1-1
—
& Noor
1739: 4-1
515:
1-1
1172:
13305:
8-1;
7734:
16336:
1-1;
Khan
1-1
2673:
2675:
6-3;
28260:
4-1
Krukoff
330:
Kuschel
9141:
2331:
1-1
5175: 7-la
—
507:
1-1;
4-1;
26156:
—
4-1 —
7288:
12: 4-1
LU
4-1;
6-1;
86-571:
201:
2-1;
91-582:
7232:
Mooney
27 b:
Nelson
6-1;
4-1
Parker
—
10648:
8-1;
29851:
25532:
445:
1-1
8-1;
—
643:
Kurz
—
1-1;
—
1-1
532:
6-1;
8-1;
5915:
3168:
1-1—Koorders
6-3;
27363:
Kostermans
103:
—
1-1;
4-1;
6-3;
7128:
13320: 2-1
Koyama
4-1;
15567:
5836:
33708:
—
6907:
1-1;
6-1;
3083:
8-1;
T-30826: 4-1
Wongprasert
9:
(as 'Amann')
523:
1-1;
16771:
20778:
1-1;
27246:
6-3;
1166: 8-1
Terao &
4977:
series
915: 6-1
6-1;
8-1;
Koyama,
KL
30827:
26128:
809:
5800:
—
23688:
14245:
1-1;
series
13304:
8-1;
1-1;
6-1;
20776:
Kassim
Kermode
—
4-1;
423:
1-1;
1-1
1-1;
6-3;
(exp. Rutten.)
8-1
—
4-1;
2499:
1-1;
2-1
—
—
—
424:
41139:
—
—
—
—
Meebold
114:
6-1
8-1
6-1
—
Nooteboom
1-1
—
—
6100:
8-1;
Lawson
—
Lei 3:
4-1;
110:
929:
3329:
Lorzing
1-1;
7338:
85-625:
8-1
Monod
1-2
6-1
—
4-1;
1-1;
85-670:
1-1;
85-
1-1;
89-355:
1-1;
90-
8039:
Must
1280:
3400:
6-1
611:
4-1
—
6-1
—
—
—
Musser S
1530: 4-1.
4-1;
& Clarkson
Nowicke &
—
Melegrito
McKee
—
2969:
6-1; 11741: 6-1; 18444:6-1;
Niyomdham 772:
3-1
3421: 6-3
Murton 27: 4-1
Neldner
MADw
4-1 —Maxwell
de Froideville
—
—
—
& Islam
2264:
6-1;
88-1102:
48 A: 8-1
6-1;
4107:
Majumder
—
1957:
1-1;
—
1272:
1-2;
8023:
6-1;
8-1
4355:
4-1;
McDonald & Afriastini
Muchtar
729:
L 794:
6-1;
88-278:
4-1;
Collector 2441:
42940:
6-1;
4-1
4-1
NGF series 502:
10574: 6-1
3049:
4-1;
85-383:
1-2;
Native
Liou
1-1;
8551:
1-2;
—
85-378:
& Ruskandi
Smitinand & Warncke
3564:
6045:
8-1;
62214:
6-1;
—Lam & Meeuse
17088: 4-1.
599:
1-1
1-1
—Larsen,
4-1;
62169:
6-1;
657:
6-1;
Lecomte & Finet 7: 8-1
6-1;
86-914:
2168:
8-1
1369:
594:
4-1;
4-1;
—
4-1;
2983:
6-1;
387:
4-1;
273:
—
63097:
94-748:
4-1;
59113:
363:
3180:
8-1;
TFB
7890:4-1
2-1;
1641:
LAE series
—
1554:
4-1;
16018:
1-1;
Mirmanto
1-1
Laumonier
Liang
80-221:
3-1
483:
1414:
4-1;
8408:
4808:
8-1;
86-684:
4-1;
1032:
—
2034:
8-1
8-1;
3895:
—
Po Khant
8-1;
22392:
1042:
11141:
1-1;
1-1;
22695:
Parkinson
4453:
8-1;
Phengkhlai,
6-1—Phusomsaeng
1-2
6-3;
10646:
3926:
2-1;
6-1;
4900:
1-1;
8-1;
6-1
Kanis
—
—
KEP FRI
11836:
21882:
1-1;
9
Niyomdham et
Jayasuriya
4-1
—
27894:
al.
283:
278: 6-1.
8-1.
42017: 6-1
40:
6-1;
Noordin
519:
Petelot
325:
1189:4-1
Olsen 629:
—
1-2;
3-1;
27895:
—
4824:
1-1;
307:
1-1;
Jayasuriya
—
233:
—
8-1;
13731:
19986:
1-1;
1-1 —Lakshnakara
—
H.Y.
92-6:
2-1;
4-1;
202:
245:
—
TFB 634:
13054: 6-1
1-1
5756:
6-1;
1-1;
Larsen & Larsen 33122:6-1
7814:
8-1;
2-1;
6-1;
4-1
1591:
1-2;
Nangkat
6119:
6-1;
Meijer
—
357:
1-1;
—
8-1;
15717:
1402:
75-87:
980:
6093:
2679:
2-1;
4-1
4-1;
series
1-1 —Marcan
2-1;
8-1
13:
27559:
Lemaire
74-922:
29754:
2249:
-1;
1-1—Lau
5074:
24543:
1
Lamont 636:
Lazarides
1401:
Maingay
6-1
4-1.
37619:
3690:
1-1;
291:
6-1;
1-1;
T-48952:
3080:
6-1;
2099:
4-1;
—
1-1;
HFP 1072:4-1
98637:
1-1;
1-1;
13340:
4-1;
10364:
1-1;
6-3;
830:
2-1;
8813:
306:
Kartawinata
18168:
1-1;
19238:
1-1;
Kornassi
—
Hue RH
Hatusima
—
3-1;
37096:
3581:
1-1;
10725:
4151:
1-1;
406:
Hore
—
4-1.
794:
4-1
2596:
Kunstler 4394:
—
2960:
6-1;
6-1;
&
4-1
5754:
1-1;
17086:
Kostermans & Anta
—
6-1—LaFrankie
4-1
6-3
Konta & Nanakhon
Koyama,
Lace
27209:
2677:
6-3;
33712:
6-3;
25362:
6-1;
18987:4-1;
63:
94885:
1-1;
8-1;
17849:
1-1;
2-1
—
Johnson 4929:
Kanehira
—
35910:
1-1;
7787:
1-1;
15800: 3-1
Hohenacker
—
1212:
6-1;
Ramamoorthy
—
3723:
Collector 289:
2286:
8-1;
6-1
8-1;
3-1;
4-1;
1-1;
King's
7593:
1-1;
Koelz
—
16479:
—
1896:
1-1;
7574:
15140:
9973:
8-1
94439:
6-1;
1-1;
4-1;
Jarrett &
83:
2036:
1874:
2270:
2-1;
8102:
1-1;
5540:
1759:
8-1;
8-1
1031:
4-1;
—
8-1;
34218:
6-1;
Hotta
—
5060: 4-1
643:
Karayanaswami
—
80282:
13493:
8-1;
Kerr
—
8-1;
1098: 2-1
1819:
3;
27397:
1-1;
Kanehira
—
Ho
—
2-1;
1999
1,
4-1.
Jeng
—
No.
44,
4-1
584:
294: 6-1
465: 6-1
1616: 2-1
4-1
335:
no.
5844:
Hyland
—
Janowsky
Kanjilal
1626:
Hose
—
KEP series
—
1-1;
26571:
1-1
332:
—
4, 43,
& Thomson
4-1
& Sumithraarachchi
Kadim
vol.
4-1;
Hooker
—
Horsfield 28:
Ja series
335:
no.
4-1
Vol.
1-1;
5217:
Tamura et al. 4080:
197:
1-1
382:
8-1;
4-1;
1-1
—
1-1; 213: 1-1—Pleyte
—
14396:
8-1;
1912:
Poilane
4-1;
25525:
851:
6-1;
14914:
1-1;
2-1;
5760:
8-1
Phengkhlai
6-1
—
—
Perrottet 33:
8-1;
et al. 6915:
20292:
3782:
7545:
1-1;
1125
Phengkhlai
8-1
840: 6-1—PNH series
1401:
8-1 —Polunin
1959:
4-1;
20729:
7699:
1-1;
2-1 —Pullen
—
10342:
8-1;
21868:
1795:
—
BKF
Phromdej
9;
37874:
7855:
8-1;
6-1;
4-1
=
8-1;
22252:
7076:
6-1;
H.-J. Esser:
7081:
Rahil
6-1
355:
Rao
PBU
series
51052:
50211:
3-1;
76141:
82660:
3-1;
8-1;
3-1;
6-1
88892:
8-1;
109372:
118230:
8-1;
118552:
3-1;
119827:
130234:
6-1
Sandom
Schodde
25782:
24: 6-1
& Craven
29744:
4-1;
4478:
34564:
4-1;
Kadim
973:
4-1
Shaik Mokim
4-1;
6256:
983:
8-1;
Abbe 6588
7-lb
6886:
Stainton
5137:
Taam 867:
Y.
HB
&
4-1
—
—
Tsiang
624:
—
1221:
1869:
HB
1-1;
483
=
—
1603:
8273:
8-1;
—
G:
4-1;
B:
1-1;
3222:
U.S. North
Yapp
366:
8-1;
8-1;
121613:
8-1;
123379:
8-1;
128089:
3-1;
128322:
3-1;
7972 C:
2-1;
—
8-2;
8021
Wenzel
6078:
3-1;
1231:
series
35826:
1-1;
Shimizu
—
9877:
255:
11019: 4-1
9: 9
4-1
8-1
—
—
Smitinand
Soejarto
1296 A:
4-1;
859:
3-1
4-1
8-1
4-1
418:
7213:
6-1
1-2;
—
8-1
&
—
Sprensen,
St. John
Stone & Mahmud
—
bin Tassim
Smitinand
8-1;
—
SF 38363:
=
1-2;
—
632:
617:6-1;
6-1;
Shah &
Sidek bin
—
190:
7212:
1296 B:
4-1;
—
& Kadim
6111:4-1
& Madulid
1649: 6-1
HB
1-1;
=
KL
—Teijsmann
4215:
3083:
Thwaites
4-1;
654:
339
8-1;
6-1;
HB
568
=
2-1;
=
LU
549:
HB
4233:
KL
744:
8-1;
1-1
3168:
2-1;
17088:
Phengkhlai 373:
&
2644:
6-1;
9733:
1-1
4-1;
7797
H:
4-1;
C:
1-1;
D:
8-2;
C.
9;
7644:
6-1;
4-1;
7797
—
18057:
12696:
6-1
—
8-1
—
4-1
519
=
836:
—
6-1;
Tongseedum
—
LU
HB
6-1;
2743: 6-1
Thorel
—
2132:
4-1;
1872:
HB
Telado
16018:
4-1
4-1
1-2;
4456:
2-1;
5528:
& Mohamed
5252:
4-1;
—
—
7972 F:
8-2;
—
Wilson
2-1;
12748: 4-1
—
—Zollinger
3035: 6-3.
3180:
2674: 2-1
11463: 4-1
WKM
B:
8-1;
8-2;
3-1
—
3736:
4-1;
1-2.
7797 E:
7963
6-1;
7972
7973:
J.E.
A:
8-1;
C:
8-2;
8014:
1-1 —Webster & Hildreth
Kalima
Wight 2643:4-1;
P.G.
7963
7970:
7972 G:
8-1 —Ward 6625:
—
6-1;
D:
1570:
vanRoyen
1-2;
7797
4-1;
1-1;
883:
3726:
1-2;
C:
A:
7966:
—P.
Vermeulen
1325:
7963
1-1;
443: 8-1.
1-1
4-1;4559: 6-1
4-1; 7797
—WongWKM 518: 6-1;
Explor. Exped.
—Van Borssum Waalkes
Whitmore & Titi
6345:6-1
283:
I:
D:
8-2;
34252:
9
610:
7797 B:
7965
7972 E:
Wang
1-1
3440:
18184: 7-1 b
3-2;
S. Vidal
—
7965
6215:
62:
2-1;
4-1 —VanNiel
4-1
Zainudin, Salleh
Sinclair
SMHI series
—
Larsen & Hansen
Sprensen,
207:
—
40039:
4-1;
23873:
6-1;
33503:
8-1;
183: 7-la —Sinclair 5617
10575:
1-1;
Expedition
22891:
4-1;
& Fukuoka M-14738:
Fl. Maurit.
7-la;
Santisuk et al. 401:
—
32788:
1-1;
38363:
8-1;
1-1
3030:
6-1;
31583:
1-1;
37265:
1934:
1-1;
1848:
30572:
1-1;
3-2;
—
3455: 6-1
3418:
Pacif.
7972
B: 2-1
I.B. Wilson
2-1;
SF
—
Wallich 7797 A:
7965
2-1;
3-1;
117787:
15717:
5075:
7797
—
106873:
Beusekom
1-1;
1-2
3-1;
VanderMaesen4149:
4-1;
Winkler 3271:
6-1;
82397:
3-1;
8-1;
—
LU
F:
643:
68030:
6-1;
82375:
6-1;
3-1;
44428:
6-1;
83: 6-3
Van Steenis
& Subari
67202:
6-1;
82048:
6-1;
45468:
8-1;
35482:
6-1;
6-1;
38965:
3-1;
116200:
4055:
A:
14396: 7-lb
64375:
8-1;
45218:
3-1;
27641:
6-1;
26786:
3-1;
37563:
1-1;
105292:
Steward & Cheo
7797
Wally
—
43076:
80588:
3-1;
25465:
3-1;
36214:
4-1;
Soejarto
156: 6-3
—Van
2096: 6-1
8-2;
2937:
3-1;
Specht
7965
8021 A:
Ridsdale
—
Chand
111375:
1-1;
1 -2
—
4-1;
7972 B:
A.S.
26085
104336:
1109:4-1
—
6-1;
2-1;
4-1
Rup
—
—
4-1;
3-1;
L.S. Smith
—
Tsang218
8-1;
6-1;
14940:
4-1;
1-2
24182:
8-1.
3064: 7-lb
3094: 6-1
Waas
6-1
2-1
(Kostermans cs.)
Vidal
2013:
1-2;
3-1;
Guizhou
Pachiappan
Balgooy 2113: 6-1
8-1;
1407:
90324:
5-1;
—
1-1
Thorenaar
W.T.
2207:
4-1.
3677:
Leong Eng
UNESCO
Van
890:
1-1;
110905:
8-1;
1533: 8-1 —Takamatsu
8-1;
4-1;
1475: 4-1
36:
1-2
BKF 25012:
=
Swanborn ML
Teo
988:
8-1;
Put 692:
8-1;
6-1
6574:
6-1;
& Larsen
—
3393:
6300:
Gaerlan et al. 8506:
Soejarto,
Larsen
35523:
Sino-American
—
—
1334: 2-1
4-1;
25284:
63178:
6-1;
78223:
Sieber Fl. Maurit. 67:
8-1;
10196: 3-1
969:
—
4-1
RHT series
20158:
4-1;
SAN series
30160:
4-1;
6-1;
—
Ritchie
6-1;
Sangkhachand
—
34476:
Kiah 97: 4-1
—
14518:
8-1;
35141:
3-1;
41153:
3-1;
—
6-1;
3-1;
—
4613:
4-1.
200: 6-1
—
18635:
3-1;
60030:
6-1;
78150:
8-1;
108846:
1-1
4-1
213
Hippomaneae
8-1 —Ramos
4935:
4-1;
361:
33528:
40989:
3-1;
3-1;
1-1;
2334:
16576:
106905:
—
8094:
8-1;
Ripley
3-1;
57592:
3-1;
85825:
8-1;
—
4254:
4-1;
Reksodihardjo
32356:
51953:
8-1;
77487:
8-1;
—
the
& Shah
Purseglove
4206:
7364:
of
1-1.
40973:
3-1;
54725:
—
8-1;
6-1;
3-1
13684:
3-1;
3-1;
392:
6198:
26964:
8-1;
40592:
8-1;
PBU
1-1;
1615:
Ridley
—
10043:
26828:
3-1
3912:
revision
partial
Rastini 74: 6-1
—
3-1;
4295:
1-1;
S
1 -1
4-1
155:
1 -1;
—Rahmatsi Boeea
38887:
51373:
3;
3868:
2-1;
1-1
4867:
Purseglove
—
2252:
4-1;
A
—
1085: 3-1
Wray
Jr.
1-1
—
Winit
—
Widjaja,
R.S. Williams
1670:
1-1
—Worthington
201:
1-1
—
—
67:
C.Wright
214
BLUMEA
Index
References
accepted
or
made
only
are
Letters refer
species.
Alchomea
mollis
P.
lucidus
(Sw.)
Esser
in
E2
3
lucida
Esser
1-1
Esser
chamaelea
1-2
1-1
8-2
4-1
Duvigneaudia J.
4-1
F. Muell.
asperococcus
F.
&
(Royle) Miill.Arg.
Griseb.
sect.
Gymnanthes (Sw.)
sect.
Parasapium Miill.Arg.
sect.
Sapium (Jacq.) Miill.Arg.
sect.
Sclerocroton
Triadica
Griff.
(L.)
Baill.
ex
var.
2-1
(Miq.) Miill.Arg.
loureiroana
Miill.Arg.
Zoll.
& Moritzi
insignis Royle
wallichiana
wallichii
2
Royle
Royle
2-1
& K. Hoffm.
(Spreng.)
3
6
1
T. Post & Kuntze
T. Post &
4
T. Post &
7
(Lour.) Miill.Arg.
(Miill.Arg.)
Roxb.
8
Huber 5-1
5-1
1-1
aubletianum
bingiricum
(Lour.) Kuntze
cochinchinense
(Lour.) Pax
crassifolium
Miq.
Elmer
(Champ,
E3
ex
Benth.) Miill.Arg.
Boerl. 6-1
glandulosum (L.) Morong
var.
Willd.
6-1
Trimen 2-1
genuinum Pax
lateriflorum
El
& K. Hoffm. 8-1
Roxb. E2
diversifolium (Miq.)
6-3
5-1
6-1
cochinchinense
discolor
8-2
Miill.Arg.
Roxb. ex Baill.
insigne (Royle)
2-1
Pax
Hook.f.
Microstachys (A. Juss.)
cordifolium
8-1
8-1
ex
8-1
indicum
2-1
Royle
6-1
6-1
lanceolaria
sebifera (L.) Miill.Arg.
T. Post & Kuntze
biglandulosum (L.) Miill.Arg.
Benth.) Miill.Arg.
(Willd.) Miill.Arg.
5
Hook.f. 2
Stillingia (Garden ex L.)
baccatum
Lour. El
(Champ,
Hoffm.
Arg.
(Royle)
Pleurostachya
aubletianum
4-1
& K.
Mull.
aucuparium Jacq.
insignis (Royle) Miill.Arg.
virgata
Pax
Parasapium (Miill.Arg.)
Kuntze
1-1
4
1-2
Gymnanthes (Sw.)
sect.
1-1
diversifolia (Miq.) Miill.Arg.
Falconeria
S. Vidal
Eusapium
sect. Triadica
(Roxb.) Miill.Arg.
4
Zucc.) Miill.Arg.
5
Kuntze
8
&
Americana Pax & K. Hoffm. 5
subsect.
6
4-1
Griseb.
(Vahl)
sect. Sebastiania
(Hochst.) Miill.Arg.
E5
Juss. 4
(L.) Miill.Arg.
sect.
5
Pax
Juss. 4
sect.
sect.
6
(Lour.) Miill.Arg.
cochinchinensis
indica
2
3
Parasapium (Miill.Arg.)
Miill.Arg.
discolor
Zucc.)
4
chamaelea
A.
sect. Falconeria
sect. Falconeria
baccata
Microstachys
sect.
Cnemidostachys (Mart.
affinis
populneus (Geiseler)
sect.
4-1
L. 5-1
Homalanthus
subg. Eusapium
Excoecaria L.
sect.
Wall. 2
Wall. 2-1
Sapium Jacq.
3
4
Muell.
Esser 3-2
(Steenis)
Gymnobothrys
daphnoides (Mart.
3
Wawra)
ex
3
Myrica luzonica
J. Leonard
Esser 3-1
Hoffm.)
Esser 3
(Klotzsch
comiculata
Leonard
inopinata (Prain)
& K.
(Pax
chamaelea
L. 8-2
sebiferus
Sw. 3
bicornis A.
Mart. & Zucc. 4
number.
page
Hippomane glandulosa
Mart. & Zucc. 4
myrtilloides
Elachocroton
Kurz
bold,
genus
are
Sw. 3
remota
(Elmer)
2-1
(L.) Spreng.
linearifolia Miq.
by
in
accepted
names
to those of the
are
Miill.Arg.
(Roxb.)
Kurz
Kurz
New
parts.
refer
Other references
lucida
Cnemidostachys
subsect.
nomenclatural
borneensis
subrotundifolium
insigne (Royle)
Baill.
names
multiramea
(S. Vidal)
sebiferum (L.)
1999
Gymnanthes
Merr. E4
sect.
the
species (E).
1
Carumbium baccatum
Croton
1,
inopinata (Prain)
Rothm.
Blumeodendron
No.
italics. Numbers
E3
Spreng.
(Roxb.)
luzonica
in
Browne 3
Balakata
baccata
to excluded
44,
scientific
treated
taxa
(Benth.) Miill.Arg.
Antidesma bunius
Ateramnus
to
to
in roman, and
synonyms
names
Vol.
Merr.
2-1
1-2
5-1
8-1
H.-J. Esser:
A
partial
revision
lineatum
Lam. 7-1
Merr.
merrillianum
Pax & K.
plumerioides
Croizat 7-la
1-2
Hoffm. 1-2
Shirakiopsis
indica
Merr. 6-2
sanchezii
sebiferum (L.)
subrotundifolium
8-2
Moritzi
Stillingia
Sebastiania
Klotzsch
3
ex
sect.
Wawra 3
&
Zucc.)
G.L. Webster 4
Elachocroton
sect.
Eusebastiania
sect. Gussonia
Pax
(F. Muell.)
4
Miill.Arg. [p. 206]
Microstachys (A. Juss.) Miill.Arg.
sect.
Sarothrostachys (Klotzsch)
Pax & K.
chamaelea
(L.) Miill.Arg.
Prain
lancifolia
Steenis 3-1
4
Triadica
Triadica
8-2
(Willd.)
(Lour.)
Hurus.
1-1
Vahl
8-1
Hurus. 6-1
8
sebifera
(L.)
sinensis
Lour.
Urandra
8-2
4
L. 4-1
Lour.
cochinchinensis
6
Steenis 7-la
7-la
Michx.
bicornis
chamaelea
Raf. 8-2
cochinchinensis
7-1
7-la
Geiseler E5
sebifera (L.)
Seborium Raf. 8
3
8-1
paniculata Miq.
Tragia
Baill.
8
6-1
Miq.
lineata
4
7
5
subsp. pacifica (Miill.Arg.)
4-1
Baill.
Benth. 8-1
ex
(Lam.) Miill.Arg.
populnea
Hurus.
Baill.
ex
(Lour.) Baill.
Champ,
subsp.
Steenis 3-2
sebiferum (L.)
(F. Muell.) Baill.
Klotzsch
pacifica Miill.Arg.
Pax
Zucc.)
Sarothrostachys (Klotzsch)
Benth. 3
3
&
Sapium (Jacq.)
lineata
4-1
(F. Muell.)
inopinata
indica
Klotzsch 5
sect.
Hoffm. 3-1
borneensis
Shirakia Hurus.
8-2
L. 7
sect.
lanceolaria
sect.
asperococca
Eustillingia
diversifolia Miq.
3
Miill.Arg.
Elachocroton
sect.
discolor
(Spreng.) Miill.Arg.
Esser 6-3
4
sect.
sect.
Sarothrostachys (Klotzsch)
Miq.)
Cnemidostachys (Mart.
Baill.
Spreng. [p. 206]
sect.
chinense
ex
subg. Sapium (Jacq.)
Klotzsch
Cnemidostachys (Mart.
subsect.
Garden
ex
8
sebifera (L.) Bojer
Miq.)
ex
Hook. f. 6-3
Sarothrostachys
multiramea
Esser 6-2
& Moritzi
Stillingfleetia Bojer
Elmer E4
virgatum (Zoll. &
Esser 6-1
(Merr.)
virgata (Zoll.
Roxb.
6-3
Miq.) Kruijt
ex
Esser 6
(Willd.)
sanchezii
6-2
& Moritzi
virgata (Zoll.
1-1
populifolium Wight
remota
215
Hippomaneae
sanchezii (Merr.) Kruijt
luzonicum (S. Vidal)
var.
the
(Shirakia)
(Sapium)
sect.
of
8-1
Lour.
Small 8-2
8-2
elliptica
Merr.
1-2
Notes
V. Malecot
only
Now
p.
Y.
one
(Paris),
species
of
the African
who
currently
works
Stillingia occurs
representatives
on
the
in Africa
of the
Hippomaneae of Madagascar, confirmed
(i.e.,
genus
the
can
Mascarenes), namely
be
considered
as
S. lineata
sufficiently
that indeed
(pers. comm.).
well
known
(see
190-193).
Tseng [Fl. Reip. Pop.
synonym
note
2,
p.
of
Triadica
204).
Sin.
44,
3
(1997) 18-19] recently proposed Sapium
cochinchinensis
(as ‘Sapium discolor’),
laui Croizat
and confirmed
my
as
a new
suspicion (see