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BLUMEA
29
(1984)
499-512
The genus Myxopyrum L.
(Oleaceae)
R. Kiew
Universiti
Pertanian
Malaysia, Serdang, Selangor, Malaysia
Summary
The
morphology
Of the
given.
nervosum
Bl.
(synonyms
M.
ellipticum),
(synonym
pinensis),
leaf
and
M.
with
thus
(Steen.)
enerve
horsfieldii,
one
M.
Hill
four
species
for the
Descriptions
M.
described
and
two
and
Kiew.
enerve
Bl.
M.
species,
M.
philip-
(synonym
Steen.
is
is
M.
Kiew
(Bl.)
cordatum,
smilacifolium
Myxopyrum
extra-Malesian
M.
species
recognised:
are
coriaceum
subspecies
one
to the
key
a
subspecies
macrolobum,
and M.
hainanense)
subspecies confertum (Kerr)
Kiew.
with
zippelii)
(synonyms
M.
(synonym
is
Myxopyrum
described
M. ovatum
Gagnep.
pierrei
serrulatum)
M.
of
anatomy
species previously
15
M.
Chionan-
smilacifolium,
is
given.
Introduction
Hill
reviewed the
(1910)
Kurz
ilicifolium
recorded
Java; M. coriaceum
Java
and M.
Five
the
(1938)
hainanense
Hill
by
Chai
since
(with
a
more
affinity
and M.
able.
the
those with
zippelii,
They
were
tory key
a
longer
could therefore
this flower character.
needs
to
Hill from
and M.
macrolo-
(1967)
(1912)
from
by
from Borneo and M.
He
and M. cordatum
and M.
in
nervosum,
lobes).
New
the
on
concluded M.
also
of the corolla tube
length
smilacifolium
long lobes)
(New
M.
than
corolla tube. Three
to
are
ellipticum
those of
species,
not
on
be
most
placed
in Hill's
specimens
are
key
serra-
closely related,
and M.
He considered the Indian
Guinea
and M.
Guinea species also with
M.
Malaya,
which
was
based
collected in fruit
leaf and inflorescent characters.
so
showed
Borneo and
M.
not
a
horsfieldii
species
cordatum,
imperfectly known, flowering specimens
In fact
be based
Elmer
Indochina, M. confertum
Van Steenis
short corolla tube and
long lobes)
then
from
groups based
two
ovatum
species
a
var.
Sumatra and
horsfieldii
(Admiralty I.)
philippinensis by
(1933)
by
enerve
corolla tube and short
to
smilacifolium
Malaya,
from Borneo; M.
described: M.
lobes).
with
corolla tube and
Hill
M. cordatum Hill
Gagnepain
genus into
species
from
nervosum
species. Myxopy-
ten
M.
(with
from Hainan.
(1974)
short
Java, i.e.,
been
by
Thailand, M.
M. macrolobum, M.
longer
ellipticum
M.
recognised
India
from New Guinea.
shorter than the
or
(Indian
tulum
are
from
Burma);
(Kei I.),
M. pierrei
divided the
(longer
as
zippelii
Philippines,
Kerr
Hill
have
species
from
Bl. and M.
ovatum
bum Hill and M.
of the genus and
history
Bl. and M. serratulum Hill from
smilacifolium
rum
coriaceum
being
avail-
primarily
that
a
on
satisfac-
500
BLUMEA
Most
on
of Hill's
single
a
or
clear that
come
species,
few
very
Myxopyrum
addition little
In
in
Oleoideae
of the tribes of
leaf,
simple
fruit
is
not
three
short
ovules
ascending
Johnson,
and
within
is
cork
and
as
and becomes
The
terete.
possess
terete
a
stem
(1980)
a
have identified
found
in
other
any
separate tribe, Myxo-
THE GENUS
triplinerved
are
leaves have
be confused.
usually
stem,
a
(Kiew, 1983).
grows older it
triplinerved
to
the
e.g.,
vascular strands in the
not
are
style,
differences,
distinct genus in the Oleaceae; it is
very
As the
Strychnos (Loganiaceae)
they
a
OF
young stems, the leaves
quadrangular
towards the apex.
serrate
are
which
Oleoideae
subfamily
wider range of
with tribe Oleeae the
also shows
This indicated it should be retained in
Vegetatively, Myxopyrum
plant
common
it
But
Myxopyrum
a
4 valvate lobes, short
triplinerved leaves, separate
stem,
MORPHOLOGY
with
has in
and in addition Harborne & Green
placed
when
allied with the tribe Oleeae than
with
present.)
are
of apigenin from
genus of the Oleaceae.
(It
ovule number and
on
from its traditional
position
However,
closely
corolla tube
diagnos-
be re-examined.
to
Over-emphasis
Jasminoideae.
is more
based
were
available it has be-
are
variable for many of the
Myxopyrum
remove
Jasminoideae.
subfamily
bilobed and flavones
glycosides
pyreae
to
Gagnepain
specimens
more
extremely
are
subfamily
comparatively
1984
2,
delimitation needs
Myxopyrum
climbing habit, quadrangular
petiole,
species
species
(1957)
to
characters is considered,
any
Now that
known about the genus.
was
caused Johnson
subfamily
No.
29,
those of Elmer, Kerr and
as
specimens.
tic characters used and that
position
well
as
VOL.
-
Species
develops
led
of
and in
a
thick
of
specimens
Strychnos
can
a
twining
some
layer
species
of soft
Myxopyrum
be told apart
have tendrils, the flowers have five
stamens
and
the fruit may have many seeds.
The
degree
Only
an
of serration of the leaf margin
and M.
vosum
one
smilacifolium,
species, M. ovatum, is
entire margin.
and is
serrate
given
not
This
as
spinose
yet known only
variation appears
such
as
M.
serrulatum and M.
so
from
that
species
taxa
smilacifolium
species
based
var.
as
to
M.
ner-
entire.
with leaves with
specimens
the
such
dentate
obscurely
to
randomly through
correlated with any other character
margins,
variable in
very
distribution
conspicuously
on
illicifolium
are
not
taxonomic rank.
Leaf
shape
coriaceum,
M.
is
also
variable but
smilacifolium
date leaves of M.
ovatum
the
narrowly elliptic
subsp. confertum
enable them
nation with other characters
(table
to
opposed
lateral branches of
decreasing
terminal than in
produced (longer
depending
throughout
on
axillary inflorescences)
on
older
wood).
be identified
size
bear further branches which terminate in
great variation in length
and M.
leaves of M.
pierrei
by
nervosum
and the
ovate
subsp.
or
cor-
their leaf width in combi-
1).
The inflorescence is uniform in type
is
is
from almost
the genus. It is
towards the
cymules
the
and
of flowers.
position
on
a
lax
panicle
with
these lateral branches
apex,
Within
a
species there
of the inflorescence
(shorter
of the wood from which
age
In addition it appears that
they
the infructescence
in
are
un-
R.
The
L.
Myxopyrum
genus
501
Character combinations for the identification of
1.
Table
Kiew:
materialof
fruiting
Myxopyrum.
Leaf width
width
Inflorescence
Inflorescence
(cm)
M.
(mm)
3—1.5
3-7.5
8
8
12x10—20x15
12x10-20x15
(4 —)5 —8
—8(
( —11.5)
11.5)
(3.5—)5—10(—14)
(3.5 —)5 —10( —14)
17x12—20x20
17x12-20x20
nervosum
coriaceum
coriaceum
subsp.
subsp.
nervosum
nervosum
—
M.
ovatum
M. ovatum
M.
M.
M.
6-14.5
6-20(-32)
9x8-20x15
9x8-20x15
4-7.5
(6—9—)21 —35
(6—9—)21—35
13x16-19x17
pierrei
pierrei
smilacifolium
subsp. confertum
confertum
3 —4.5
3-4.5
subsp. smilaficolium
subsp.
4—8( —10)
4-8(-10)
dergoes
some
species,
it
(table
acute
ally
be
used
is
calyx
uniform for all
lobes about
between
forms
1
than
flower with
mm
1 and
3
long
mm
campanulate
tube and open, often
to
length overlaps
with other
conjunction
The
stamens
stamens
between
characters
encountered in M.
are
one
or
number,
suggested by
ovules have
In all
nor
seed
not
been
M. pierrei, all
pendulous
The
as
fruit
is
a
Blume
ovatum.
M.
seeds. The
that of M.
as
are
There
usu-
two
are
tube and rounded
linear lobes which
lobes,
nervosum);
and the
longer
than the
are
valvate, usually
4
but 5 lobes
and about
1
mm
scarcely
long. Rarely
three
smilacifolium.
stigma.
may
(1850)
Hill
The 2 locules each contain 2
ovules,
have between 4 and 1 seed. Neither
he
when
outer
meant
taxonomic character
a
1—3 ovules in the
gives drawings
and M.
ovules.
The
In
single
(1967)
it
as
contains
is
nervosum
record
a
ovary.
of
and
and
nervosum
of dissected flowers
addition Kerr
for M.
pericarp
as
recorded 1—3 ovules per locule but 3
I have observed this for M.
ellipticum
ascending.
berry; globose
two
few species.
fleshy
therefore be used
ovules is that of Van Steenis
when it contains
a
ovatum.
the fruit
can
showing ascending
confertum
thick
to
and M.
ascending.
horsfieldii,
in
mm
extrorse
reported since, perhaps
are
the base with 4 short
that after the corolla has fallen the
ovary may have
similarly
coriaceum and M.
ovules of M.
so
at
contained within the corolla tube and
are
oblong,
sessile 2-lobed
number,
the ovules
barium sheets of M.
of M.
ovatum
Gagnepain (1933).
species
subsp.
a
both may abort
and 1 ovule and
between 4
5
(such
open
tubular
outside. The corolla is very small;
a
oblong
specimens
anther is
The ovary is ovoid with
though
tube and
subsessile and
are
above it. The
protrude
shortly
reaching
but
long
it is
—
become recurved. The lobes
have been noted for several
trates
in
pubescent
tube, which scarcely
the
a
species
and is
the urceolate flower with
of corolla:
shorter
ovule
5x5-10x13
of inflorescence
range
recognise species
to
8x10-15x10
( —6.5)
(-6.5)
7-19
the
growth. Although
can
1 —5
1-5
(—6)
(-6)
1).
The
its
Fruit size
length
(cm)
(cm)
on
Gagnepain
(1938)
her-
illus-
describes the
Myxopyrum
possessing
nervosum.
single
fleshy
seed, broader than long
and either orange,
crimson
502
BLUMEA
when
purple
or
is
The inner
ripe.
The
papery.
seeds contain
(slimy
name
The
wheat).
that
5
mm),
most
have
species
nervosum
seeds
dry
Myxopyrum
of this
mata
3—4 of the
Rao &
Das
fresh
collected
2619,
TS of the
ovatum
Ramos &
Edano
The
petiole,
colleagues
the
Petiole
in
peltate,
25
pan
30-70 /jm
tate,
tissue
M.
M.
nervosum,
—
than
few
to
square
in
TS,
taller
1 and 7
flush with surface.
the abaxial
6
or
Kebun
Raya,
41711, Katik46864,
For each
specimen
made.
were
investigation by
in TS with
infrequent,
pierrei,
Baas and
1750—3330
slightly
in M.
convex
papillose only
sunken
sunken
long
brachysclereids
absent, except in
(absent
strands
major
glandular
eglandular pelpierrei.
Ground
associated with vascular
in all
2.5 pan thick
in M.
than
Cuticle
200—250 pan
abaxially.
surface smooth
Epidermis
thickened in M. pierrei.
or
plane
on
outer
Stomata
peltate
on
glandular
and
in M.
above
both
sur-
wall thinner
Epidermal
cells oval
straight, rectangular
abaxially (7—12 pan). Hypodermis
pierrei).
Trichomes
on
pierrei,
view, anticlinal walls sinuous
pan)
pierrei
ova-
(fig. 1).
thick in M.
species.
of M.
specimens
coalesce in M.
above in M. nervosum, M. pierrei,
8) cells, slightly protruding
surface, present
some
(these
of lateral traces
pairs
grooved
in surface
cytic,
(sometimes
many
moderately
adaxially (7-25
midrib
to
130—150 pun
adaxially,
anticlinal wall
above
as
cells.
(from
nervosum
and
in M. ovatum,
papillose
M. ovatum, M.
sclerenchyma
Isobilateral,
layers
4
under
Trichomes
below.
Vasculature of 3
polygonal
or
Malaysia
of the lamina
currently
sto-
type)
present in all species, associated with prismatic crystals in
crystals
M. ovatum. Midrib
cuticle,
clearings
peltate
epidermal
shallowly grooved, winged above,
or
nervosum,
in M. ovatum; prominent below
faces,
Reserve,
in M. ovatum, unicellular, 35—50 pan
2.5—5 pan thick
mm.
Myxopyrum.
pierrei {How 73504).
and M. ovatum,
with
described the
Myxopyrum
long
with between
Lamina
is
20
by
has the anomocytic
long
(Edano 41711).
ovatum)
(1908)
Epidermis rectangular
rectangular
Perivascular
ovatum.
tum
examined.
and M.
20
of
(5 by
head of 4 cells and that the
in M.
parenchymatous,
tissue. Ascicular
a
different from the other
Forest
subsp.
a remnant
Leiden.
deeply
nervosum
mm to
herbarium specimens: Edano
33666)
surface smooth.
in M. pierrei,
groove
(from
Myxopyrum
Outline TS
surface in M.
outer
Pasoh
Rijksherbarium,
Cuticle
across.
pun
—
of
family (Which
was
and TS, maceration and
wood anatomy of
at
species
generic
has smaller fruits
10
by
testa.
papery
THE GENUS
Solereder
slightly
were
from
M.
Bogor, Indonesia),
Lam
OF
a
hence the
oil, perhaps
generally
from 10
consisting
as
by
of M. nervosum, its
viscous
recorded the presence of foliar sclereids in
(1979)
specimens
covered
specimens
which
anatomically.
leaf anatomy of three
The
some
unusual for the
cells
neighbouring
and the septum between the locules
by copious
ANATOMY
smilacifolium
were
species
1984
2,
enveloped by mucilage,
as
fruits ranging in size
little known
is
hairs of M.
woody
smilacifolium
LEAF
glandular
of
surrounded
are
from M.
mucilage. Apart
dries
layer
No.
29,
copious horny endosperm,
Blume recorded the seeds of M.
coriaceum and M. pierrei
VOL.
-
abaxial surface
of 1
only,
or
2
anomo-
with stalk sunken and head of
above the
surface,
adaxial surface of M. nervosum, M.
usually present
on
pierrei; eglandular
R.
1.
Fig.
Petiole
vasculature
C. M.
(Lam 2619);
of
cells
layers,
The
A.
Myxopyrum.
L.
Myxopyrum
genus
M. nervosum
503
For.
(Pasoh
on
adaxial surface of M.
15—37.5 pm tall, in M.
Palisade
ovatum.
and M.
nervosum
the
pierrei
mesophyll
usually present
sade. Sclereids associated with veins,
Midrib
The
glandular
and
stomata are
the
specimens
they
are
found in the
terminal
(i.e.,
Oleaceae
The
uniformity
tionship
the
between
other
petiole,
species
the
in anatomy
far
so
single pair
clinal epidermal
are
&
M.
examined:
of lateral
family,
such
as
presence
neighbouring
cells
are
Che Su,
veins).
are
1982)
of the
but
but from exami-
different
scarcely
they
stomata
cells of the
enough
to
most common
be
type
unusual in that
are
In species of other genera, e.g. Olea and
diffuse.
that it is
means
though
that the
The foliar sclereids
associated with
species,
pali-
and 125-250 ptm
surrounding epidermal cells,
(Kiew
Chionanthus the filiform sclereids
of the
typical
noted
neighbouring
classed otherwise than anomocytic.
(filiform)
had
different from the
slightly
nation of these
2
moderately
trichomes, acicular crystals, anomocytic
(1908)
Solereder
long
to
specimens examined indicates that the anatomy of
show features
species
eglandular peltate
and foliar sclereids.
550—2000 pm
filiform,
to
sometimes in
mesophyll,
spongy
1
scarcely
TS, secondary veins arranged in row below palisade.
of these
the genus is uniform. All
of
in
crescent-shaped
foliar anatomy
in
is
palisade
differentiated from the spongy mesophyll. Spongy mesophyll compact
compact. Ascicular crystals
wide.
B. M. ovatum
Res.);
pierrei (How 73504).
trichomes present
peltate
Kiew:
pierrei
the
e.g.,
in the
traces
not
shows
helpful
in
determining
several characters
not
the
of unicellular hairs
presence
petiole (fig. 1)
rela-
shared
on
by
the
and the thickened anti-
walls of the adaxial surface of the lamina.
ECOLOGY
All
or
in
species
exposed
are
recorded
areas
been recorded from
such
from the lowlands below 1000
as
open
swampy
areas
poor acid soil of peat swamp and
Nothing
is known of the
thickets
and
more
its
rarely
pollination
or
or
seashores.
subspecies
m,
both in
Myxopyrum
coriaceum is
primary
forest
nervosum
confined
to
has
the
of heath forest.
fruit
dispersal
of
any
species of this genus.
504
BLUMEA
VOL.
-
TAXONOMY
The
genus is uniform with
in
the
of
tween
and
fruit,
(table
ters
Other
material
characters
and dichotomous character
discontinuous and there
is
using
a
of inflorescence
length
considerable
is
here
species recognised
be identified by
can
based
were
of
(degree
and
species
reduction in
cases
only qualitative
are
overlap
be-
nevertheless discrete
combination of these three charac-
1).
variable within
able
of these
none
for each character. The
species
fruiting
THE GENUS
there is variation in leaf shape and size,
Although
genus.
and size
1984
2,
habit, venation, size of flower, type of inflo-
to
regard
OF
and fruit. Form of corolla is the
rescence
No.
29,
the
are
serration
not
number of
useful
of
in
species,
flower
This has led
Hill's
particular
minute differences in flower
on
leaf texture,
margin,
taxonomically.
size and
to a
size)
which in
species,
are
considermost
of serration of the
degree
margin.
Six
taxa
relatively
here, four species
recognised
are
which stretches from India
Sundaland but
extending
ranging
wide
ranges
pines
over
ern
to
Guinea
M.
were
nervosum
origin with its
into Malesia
Java
with
along
entire
(the
Guinea).
on
species, although
few collections. M. pierrei
ovatum
centre
of
diversity
a
of
ranges
from the
Philip-
longer
corolla
diversity
in
here
account
Malesian
changes
M.
species,
for
the
provides
has
Philippines
a
key
smilacifolium
Malesian
in the Flora Malesiana
species.
short
corolla
had
coriaceum with
subspecies
probably ecologically adapted
New
to
identify
to
with
its
all
subsp.
Descriptions
spread
Sumatra and
spread through Malaya,
the
thick
to
the
other track from the
Guinea and M.
ovatum
has
re-
species.
species,
descriptions
confertum,
of the
tube)
north-
a
in the Thai-Indochinese region and has
nervosum
swamp and heath forest. The
the
through
a
tube). Myxopyrum probably
Borneo which is
of peat
that the species of India
statement
(both possessing
Celebes, producing
margins
is
related
closely
tracks: M.
two
Hill's
explains
species
more
tained the short corolla tube of the Indian and Indochinese
The
species
(based
nervosum
The other
are
species
northern
most
and M.
relatively
Borneo and M.
in
Two
subspecies.
two
Indochina)
known from
are
Sabah
(with
acid soils
impoverished
centre
(fig. 2)
to
New Guinea and
to
leaves
Burma to
into Celebes and New
of the
geography
New
than
area
Indochina
through
and
smilacifolium
New Guinea and New Britain.
to
The
and
a
from
M.
widespread:
and
common
Malesian
and
of the
extra-
nomenclatural
species
will
appear
account.
REFERENCES
HARBORNE,
J.B. &
the Oleaceae. J.
HILL,
A.W.
JOHNSON,
KIEW,
R.
P.S. GREEN.
Linn.
1910. The
L.A.
1984.
Myxopyrum
genus
A
chemotaxonomic
Two unusual
of the
family
Chionanthus
family.
survey
of flavonoids
in
leaves
of
155-167.
Myxopyrum (Oleaceae).
1957. A review
in the
1980.
Soc. Bot. 81:
Kew Bull.:
(Oleaceae) species
J. Arn. Arbor,
37-44.
Oleaceae. Contr. N. S. W. Natl. Herb.
(in press).
from
Borneo
and
2:
the
395-418.
position
of
R.
KIEW,
R.
&
I. CHE
nanthus
and
SU.
with
of leaf
anatomy
reference to foliar
special
505
L.
Myxopyrum
genus
Comparative study
1982.
(Oleaceae)
Olea
The
Kiew:
of
of Chio-
Malaysian species
sclereids.
Soc. Bot. 84:
J. Linn.
79-101.
RAO,
T.A. &
SOLEREDER,
Frisch,
1979.
S.DAS.
Bot. Notis.
Leaf
sclereids
H.
1908.
Oleaceae:
Systematic
TO THE
distribution
SPECIES
Transl.
dicotyledons.
the
angiosperms.
elliptic;
OF
MYXOPYRUM
2
3
if
lb. M.
narrowly elliptic
then chartaceous
Leaves
ovate to
b.
Leaves
elliptic
Inflorescence
cordate, less than twice
to
ovate, twice
large,
6—35
as
long
long,
cm
as
long
as
or
coriaceum
subsp.
nervosum
..
la. M.
3a.
subcoriaceous
subsp.
nervosum
wide
nervosum
2. M.
ovatum
wide
as
with
4
large fruits,
13
x
16—19
x
17
mm
3. M.
b.
Inflorescence 1—19
cm
long,
5a.
Inflorescence 7—19
cm
long
b.
*
Inflorescence 1—6.5
Fruiting material
1.
Myxopyrum
a.
subsp.
M.
Fl.
Gamble,
3
x
13
pierrei
5
mm
4b. M. smilacifolium subsp.
using
Blume
5 —10
x
4a. M. smilacifolium subsp. smilacifolium
long
cm
be identified
nervosum
with fruits 5
of characters
the combination
given
confertum
in table
1.
Fig. 1,2.
—
nervosum
BL,
Bijdr. (1826) 683;
Ind.
Bat.
nervosum
Miq.,
can
& F.E.
L.A. Boodle
by
Oxford.
Press,
coriaceous
narrowly elliptic, thickly
Leaf ovate,
4a.
in
campanulate
b*Corolla thin and
b.
and
fleshy
1 a*Corolla urceolate, thick and
Leaf
of the
anatomy
982. Clarendon
521-526,
KEY
2a.
occurrence
-
132: 319-324.
Mat.
Fl.
2
Mai.
(1912) 51; Ridley,
Backer & Bakh.
M.
horsfleldii Hill,
M.
zippelii Hill,
Pen.
Kew Bull.
Kew
4
Bull.
2
in
1
(1850) 320,
Hook,
(1910)
44.
44.
-
215.
-
-Type:
Ty
p
e:
f.,
Kew
(1923) 320; Kerr
(1965)
(1910)
Bot.
Clarke
(1905) 272; Hill,
Fl. Mai. Pen.
Fl. Java 2
f.,
Mus.
(1857) 549;
Bull.
in
herb.
T.
51; DC.,
India
s.n.
Prod.
8
(L!), Salak,
Lugd.
Bat.
(1844) 290;
(1882) 618; King
Fl. Siam. Enum.
Horsfield (K!),
Herb.
3
(1910) 42; Koord.,
Craib,
Blume
Type: Zippel
t.
Fl. Brit.
Exk.
2
Fl.
&
Java
(1939) 420;
Java.
Java.
908,161-792 (L!),
New
Guinea.
b.
subsp.
coriaceum (Bl.) Kiew,
M. coriaceum
Bl.,
Mus. Bot.
1
(= Strychnos nux-vomica?)
Ind.
M.
Bat. 2
3039
(K!),
Kew
Borneo.
Bull.
nov.
(1850) 320, non Chondrospermum
in
(1857) 550; Merr.,
ellipticum Hill,
stat.
Hill,
Kew
Bull.
Enum. Born.
(1921)
(1910) 42; Merr.,
?coriaceum
(1910) 44; Walp.,
489.
Enum.
-
Ann.
Type :
Born.
3
Wall.,
Korthals
(1921)
489.
Cat.
no
2838
(1852) 18; Miq.,
-
s.n.
(L!),
Fl.
Borneo.
Type: Haviland
506
BLUMEA
Notes.
larly
Myxopyrum
Java
in
and
flower form
it
—
(the
the
noted that the corolla form
its
ler leaves
(4.5
6
by
shows
the flowers
can
There is also
1
in M.
as
a
i.e.
leaf size
and
mm
in M.
horsfieldii
that M.
and smal-
in M.
cm
of M.
nervosum
nervo-
horsfieldii
where flower
e.g. Koorders 31195.
single inflorescence,
a
6
by
specimen
type
nervosum
nervosum)
than 13
larger
different from M.
no
so
2.5
cer-
similar corolla.
a
species, distinct from M.
to
of the
length
within
its
open. Blume
tube) scarcely
resembles that of Noronhia and
new
horsfieldii
long,
mm
in
than the
closely
opposed
as
corolla
mm
3
to
difference
no
long
by
urceolate corolla tube which is thick and
an
shorter
or
most
horsfieldii
cm
be 3
to
from
vary
of the genus, particu-
species
genus of the Oleaceae of Malesia has
mm
Remeasurement of the
sum.
size
13
to
up
in any
described M.
corolla
longer
length
same
species
(1910)
Hill
by
other
1984
2,
most common
with
only species
fleshy,
no
No.
29,
It stands apart from all other species of the genus
(1826)
tainly
lobes
is the
nervosum
Malaya.
is the
VOL.
-
cannot
be maintained as
a
separate species.
Serration
rum
of the leaf
towards the
serrate
Hill
zippelii
and
margins
so
is
based
on
distinct
species
variable in M.
chartaceous
coriaceum Bl.
with
from
nervosum
to
size
not
300—650
to
response
to
a.s.l.
swamps
(Clemens
(higher
elliptic
leaf form of
30037)
from
species
also
to
fruits
4
also
narrowly elliptic,
and
1929).
It is
or
M.
been
inflorescence
In Borneo, M.
from
length
nervo-
confined
to
the acid
therefore that coriaceousness is
nutrient poor acid
collected.
ranges
ovate, coriaceous leaves from hill
likely
be
texture
chartaceous leaves from
coriaceum is
coriaceous
can
as
(table 1).
nervosum
long. However,
subsp.
a
is
Since M.
common.
as
a
coriaceum is
of M.
subspecies
leaves but
they
are
nervosum.
on
a
narrowly
single plant (Anderson
Myxopyrum ellipticum
coriaceum because it
and it
ovate
both this leaf form and the
found
a
and in such thick
soil)
typical
is
S
of sub-
therefore
re-
has coriaceous leaves and is
peat swamp and heath forest.
of
subsp.
coriaceum
subsp.
described the seeds
genus.
other characters, such
has
coriaceum
subsp.
shape, however,
coriaceousness of M. coriaceum that
it is considered here
mm
ex-
At first it appears that it is
Forest Reserve, Sarawak, where other specimens,
Occasional fruits of
the
margins.
lanceolate. Leaf
altitude
Hill also
synonymous with
confined
(1826)
to
Stapok
coriaceum, have
as
Dried
oil.
other extreme of leaf form. It has
margin and obscured venation is
specific habitat,
slightly larger flowers,
garded
s.n.,
serrate
extreme
with
and heath forest.
Myxopyrum ellipticum
has
plants
with
narrowly
(.Kostermans 21687)
environment
a
nervosum,
and includes
m
recurved
a
confined
It is the
differ from that of M.
lowland forest
of peat
leaves
do
thin leaves
finely
Myxopy-
nervosum.
very coriaceous leaves. Leaf
ovate to
subcoriaceous.
uncommon
primary
soil
the
often
are
toothed.
obscurely
are
which has
specimen
represents
narrowly elliptic,
falls outside the range of M.
and fruit
forest
sterile
a
coriaceous leaves, with entire recurved
tremely
is
variable. Chartaceous leaves
falls within the conspectus of M.
Myxopyrum
sum
is also
margin
margin, whereas thicker leaves
These
are
presence may be related
as
the
to
frequently
nervosum
mucilaginous
only
bird
pierrei
and it is
oil fruits of
dispersal.
contain copious
and M.
also
amounts
have
oily
this feature that
any oleaceous
species
in
of viscous
seeds.
gives
Blume
the
name
Malesia and
its
R.
Fig.
2.
2.
Myxopyrum
M. ovatum
M. macrolobum
(1927)
M. cordatum
(1927)
M.
Hill,
329.
Kew
Kew
of
Fig. 1,
Bull.
Bull.
Myxopyrum
genus
L.
507
Myxopyrum.
2.
(1910) 41; Lingelsh.,
-Type:
Hill,
329.
The
Bot.
Beccari s.n., VIII-1873
Type:
-
species
Hill —
ovatum
Kew Bull.
Hill,
(1927)329.
14
of Malesian
Distribution
Kiew:
61
(FI, drawing Kl),
(1910) 44; Lingelsh.,
Kei
philippinensis Elmer,
Leafl.
Philip.
Bot. 4
Bot. Jahrb. 61
(1912)
1483.
Guinea,
Bot.
14
New Guinea.
I.,
(1927) 3;
Nova
Bot.
Guinea,
New Guinea.
(1927) 2;
Type: Moseley Challenger Exp. (Kl), Admiralty I.,
—
Nova
(1927) 3;
Bot. Jahrb. 61
(1910)42; Lingelsh.,
Beccari PP 942
Jahrb.
(FI, drawing K!),
Nova
Guinea,
Elmer 12333
-Type:
Bot.
14
New Guinea.
(K!), Sibuyan
I., Philippines.
M.
smilacifolium
Challenger 1,
auct.
3
non
Bl.: Clarke
geb. (1900) 498; Lingelsh.,
Notes.
lium),
the
it
most
did
three
resembled M.
not
from
to
M,
cordatum
cordate leaf base.
New
was
from
based
Myxopyrum
differences in corolla
length:
and
(1910)
New
a
ovatum
M.
3
(1882) 618,
(1891) 403; K.
I.
was
Bot.
Hemsl.,
Laut., FI.
nervosum,
in flower form and
so
Schutz-
(1910)
both based
a
as
a new
two new
on
and
and M. macrolobum
having
smilacifo-
(1882)
gave
Moseley's specimen
described it
described another
Guinea,
M.
Clarke
New Guinea. With the addition of
it became obvious that
Hill
p.p.;
the first from the New Guinea
coriaceum, M.
fruiting specimen
ovatum
Sch. &
2.
including
as
on
13
Admiralty
Guinea,
time Hill
and M. macrolobum,
Myxopyrum
FI. Brit. India
(1927)
smilacifolium
smilacifolium
same
f.,
Bot. Jahrb.
then known (M.
smilacifolium
collections from
belong
cordatum. At the
tum
species
distribution of M.
Beccari's
Hook,
Jahrb. 61
Bot.
Moseley's specimen
Of the
region.
in
(1885) 241; Warburg,
Beccari's
was
were
species,
species,
M.
collections.
distinguished by
separated by
corolla tube 1.5—1.75
M.
ova-
mm
its
minute
long
and
BLUMEA
508
corolla lobes 3
in
long
long
mm
opposed
as
No.
29,
tube 2—2.5
to a
mm
that there is
It is nowhere
to
stant
within
of the
mm
an
entire
(e.g.
old
wood.
appropriate
as
all
the
M.
(1910) (M. cordatum,
not
has inflorescences 6
46864)
Of
as
three
for
names
macrolobum,
M.
cm
New
specimens
long
M.
ovatum)
on
it
be
cannot
which
to
is
con-
cordate.
Spe-
veins, in fact they have
marginal loop
is
young
ovatum
in the basal
variable
twigs
—
chosen
is
single
a
and 20
described
species
have
or
ovate
length
Guinea
have cordate leaves
5
having
Inflorescence
prominent.
urceolate)
or
veins but in cordate leaves the
thick and
NGF
Hill
by
species,
is variable and
length
also varies from
species
described
were
longitudinal
leaf is
in this
shape
cordate leaves
of
single pair
on
Leaf
species.
cimens with
specimen
with
species
and unlike the other Myxopyrum
common,
unlike corolla form (campanulate
separate taxa,
used
long
and lobes 3.5-4.5
broad-leaved
single
a
has been collected from rivers and beaches. Corolla
region
long
in New Guinea and that its distributionextends northwards into the Phi-
margin
lippines (fig. 2).
a
1984
2,
M. macrolobum.
Additional material shows
leaf
VOL.
-
as
the
cm
Hill
by
most
corol-
exceptionally long
la lobes.
Elmer
15
by
described M.
(1912)
7—9
apparently
cm,
cordate leaves
(e.g.
Edano 41711, have
leaves
(14 by
6.5
very
shape
leaves
as
of both
cm) tending
synonymous with M.
Myxopyrum
ovatum
most
easterly
Hill
recorded
of M.
most
nervosum
a
M.
pierrei Gagnep.,
Ciaib,
Bull.
Fl.
sizes
Soc.
Siam.
on
cm). However,
and
the
than the
some
larger
specimens,
plant,
same
there is
leaves
the
e.g.
smaller
(17.5 by
differ in
other character and is
any
a
10.5
regard-
closely
related
to
in New Britain
M.
pierrei (see there)
Philippines
(Frodin
M.
NGF
zippelii,
to
New
and has
appa-
Guinea with its
26565).
from
New Guinea. This
is
a
(see there).
Myxopyrum pierrei Gagnep.
in
ovate
not
fourth species,
3.
Kerr
more
does
14.5
by
leaves,
which have
ovatum.
locality
described
(1910)
specimen
is
and
ovate
Philippine specimens
extremes
shapes
be
to
two
the island chain from the
rently spread through
specimen which has
a
leaf 23.5
a
between these
cm). Myxopyrum philippinensis
ed
from
different from other
Ramos 43178 with
range of leaf
complete
phiilippinensis
Bot.
—
Fig. 1,2.
Fr.
Enum.
2
80
(1933) 78;
(1939)
421.
Fl. Gen. Indo-Chine
Lectotype:
-
3
(1933) 1084,
Pierre
2804 (P!),
t.
123;
Cochin-
china.
M. hainanense
73504
Chia,
Fl.
Hainanica
3
(1974)
209
&
Addenda
(1974)
577.
Lectotype:
-
How
(A).
Extra-Malesian specimens examined.
THAILAND. Kerr
17705
VIETNAM. Pierre
2804
LAOS. Poilane
2399
CHINA. Hainan:
(BM, K),
(A, P),
18086
Poilane
2 &
3).
How 73504
It is
(P),
8050
(P); Tsang
27015
(A, K, P).
(P).
(A, BO),
Notes. The distribution of M.
(fig.
(K).
7426
73889
(A); Wang
pierrei overlaps
readily distinguished
from M.
34636
(NY),
with M.
34305
(NY).
smilacifolium
smilacifolium
by
its
in Indochina
large
fruit
(13
x
R.
16-16
is
19
x
distinct
a
long
and
mm)
resembles M.
closely
elliptic
with
species
each
ovate
young fruit and
it is
of veins
Hainan specimens
as
9.5
cence
lium
but
(How
has
short
Chai describes the corolla tube
the tube and the lobes
4.
Myxopyrum
a.
subsp.
M.
not
Mus.
Bl.,
(1910) 40; Gagnep.,
(1939)
421.
ii
Bot.
Bull.
Wallich,
Kew
e:
Kurz
2309
Hill,
Kew
Bourdillon
555
smilacifolium
Liana,
c.
(K!),
auct.
1
plinerved
cm
with
thickened.
Cat.
of
long
and
of the
one
an
infructes-
6.5
cm
narrower
in
the infructescence
leaves and smaller fruits
the lobes
are
equal
in
with the urceolate corolla
than the lobes which in the open
3.
Fig.
—
Hook.f.,
in
Gen. Indo-Chine
no.
2837
3
For.
Fl. Brit.
Fl. Brit. India
(1933) 1083;
Cat.
Burma
3
Kerr in
no.
2
(1877) 160;
(1882) 618,
Craib,
p.p.;
J.
Hill,
As.
Kew
Fl. Siam.
Enum.
Prod.
(1844)
2837; DC.,
8
(Linn. Soc., London), Chittagong.
For. Fl. Brit. Burma 2
(1910) 40; Kerr
Bull.
S.
in
Craib,
Travancor.
Bl.:
recorded
thick,
x
apex,
(1910) 41; Gamble,
India,
as
bark
of old
4.5—8(—10)
rounded
or
(1877) 160;
Fl. Siam. Enum.
J. As.
2
Soc.
(1939)
Beng. 46,
421.
-
ii
Lec-
tertiary
or
pairs
veins
Inflorescence
for
Madras
Admiralty
stem
cm,
Fl.
I.
(see
white and
2
(1921)
under M.
papery.
membranous to
798.
-
Lectotype :
ovatum).
Leaf elliptic
to
ovate,
subcoriaceous, slightly glossy
cuneate, apex acuminate. Margin usually obscurely
to
entire
additional 3
prominent below,
cm,
long, though
cm
typical
are
describes the inflo-
(K!), Pegu.
above, glabrous, base
towards
Fl.
Bull.
non
(9—) 12—17(—24.5)
serrate
cm
6
compared
longer
(1850) 320; Kurz,
Clarke
ilicifolium Kurz,
var.
M. serratulum
t y
p
because
longitudinal
pierrei. Myxopyrum smilacifo-
to
Chondrospermum ?smilacifolium Wall.,
(1877) 245; Hill,
o
1
(1877) 245;
-
Type:
smilacifolium
t
is
as
illustration
smilacifolium
Beng. 46,
-
corolla tube
to
urceolate; however,
as
of
is in
it
recurved.
Soc.
301.
M.
the
having
recurved
are
smilacifolium Blume
smilacifolium
2
M.
where
but
in
pair
ovule in
that
pinninerved
(1974)
6
c.
of M.
range
specimens
to
erect
as
inflorescences up
1).
are
single
Myxopyrum species.
shorter. Chai
are
length
nervosum
a
2804) shows
main
as
has
pierrei
wide.
as
leaves and fruits that
73504) has inflorescences 5
differs from the Hainan
flower
have
Myxopyrum
(table
of M.
the
in all other
as
which fall within the
confertum
long
the leaf as
to
size
and the infructescence
long
cm
long,
cm
subsp.
it
4.5
c.
he cited
specimens
specimen (Pierre
equal
triplinerved
of
however their inflorescences
pierrei,
rescence
drawn of
are
as
M.
mm):
corolla and with
a
more
ovatum
less than twice
usually
are
It
1).
Myxopyrum
(9x8—15x15
with
as
long) (table
cm
three times
two or
are
509
that unfertilised ovules had aborted. The
probable
veins. The venation is in fact
The
35
to
L.
inflorescences.
in Flore Generate de l'Indo-Chine shows
pierrei
the lateral pairs
M.
smaller
locule but examination of the type
for M.
(up
large
cordate leaves which
to
generally
are
described the flower
Gagnepain (1933)
Myxopyrum
genus
which also has
leaves which
ovate-oblong
to
The
inflorescences
large
ovatum
wide and its fruits
as
Kiew:
deeply
serrate to
of lateral veins,
conspicuous
terminal,
above
sometimes
involute. Venation tri-
spinose,
impressed above, secondary
in
dried
axillary,
state.
veins
Petiole 0.5-1.5
paniculate,
7.5-19
cm,
510
BLUMEA
3.
Fig.
Distribution
of
VOL.
-
No.
29,
Myxopyrum smilacifolium
1984
2,
Blume;
subsp. smilacifolium ;o
•
subsp.
confertum.
hanging
like
bescent.
a
bunch of grapes when
Bracts
apiculate,
lobes ovate-acute,
long,
linear, opening
anthers
5x
globose,
Ovary
5 —lO
0.5—2.5
mm
less
globose,
long.
than
subsessile.
Calyx
campanulate,
1
Stamens less than 1
1
mm
fruit, finely
1.5
mm
mm
pumm
lobes
long,
long, subsessile,
bilobed. Fruit
Imm
dry, woody
1
less than
long, stigma sessile,
rugose when
xl 3 mm. Pericarp
thick in
becoming
yellow,
Corolla tube
pubescent.
recurved,
elliptic.
1
rachis
ripe,
Flowers
mm.
thick.
Seeds
1-2.
Distribution. N.
China
INDIA.
(A, P),
Beddome
Herb.
Thomson
(L, P),
&
E. India,
Burma,
Bangladesh,
Thailand,
Cambodia,
Laos,
(Canton).
s.n.,
Prazer
Mt
s.n.
1890
Bourdillon
555
Gamble
1563a
(K),
1859
(P),
(A,
Wallich
L, P),
2837
Hasal et al. 4060
Keenan
THAILAND.
368
Khasia
BANGLADESH.
BURMA.
(K),
s.n.
Gamble
s.n.
Kerr
1874
6040
(K),
(K), Chatterjee
Hooker
(K),
s.n.,
Sikkim
s.n.
s.n.,
1859
(P),
1902
564,
(A,
2197
Jenkins
P),
Griffith
(all K),
s.n.
(K),
3696
Hooker
Masters
&
s.n.
(K).
(L).
Keenan
(BM, K, P),
et
10516
al. 3868
(BM),
(A, K),
18086
Kurz
(P),
2309
(K),
Kostermans
Parker
811
(L),
2612
(K).
Put 4123
(BM), K, P).
CAMBODIA. Pierre
LAOS.
Spire
207
CHINA. Canton:
s.n.
Poilane
Ecology. Evergreen
Notes.
and
is
(P).
963
(A, P).
forests 700-1000
Myxopyrum smilacifolium
distinguished
however,
western
1970
(P).
from all other
exhibits variation
localities in India
on a
var.
m
(Gamble, 1921)
smilacifolium
species by
geographical
is
variable and
widespread
its smaller fruits. Even this character,
basis
produce inflorescences
with
thus
plants
of the coastal and
many tiny fruits,
5x5
mm
in
R.
with
compared
size,
Cambodia which
these
from the
fruits that
511
localities of Laos and
eastern
more
considerably
are
larger,
up
to
10
mm
in
larger fruit,
small-fruited
with the
compared
13
x
from the
Assam.) Apart
other differences when
no
L.
Myxopyrum
genus
have been collected from
exhibit
specimens
The
collected
plants
produce
fruit sizes
(Both
size.
Kiew:
specimens.
Another character
that
smilacifolium,
is the
leaf margin
frequently
is
The
spinose.
both
reasons
duced
to
Kurz
M.
synonymy
15
(K!),
Kew
7
c.
cm,
veins.
Inflorescence
Corolla tube
when
of the
particular
a
Stat.
(1938)
differs
THAILAND.
LAOS.
axillary,
1—5
133; in
Craib,
smilacifolium
var.
illicifolium
located specimens
not
Fl.
Siam.
Enum.
(1939)
2
characters from
following
base
as
of
420.
Type
-
subsp. smilacifolium:
long,
cm
to
up
mm
globose,
Margin entire,
cuneate.
6.5
8
x
10—15
sometimes
about
oblong,
10
x
coar-
additional pair of lateral
in fruit. Bracts
cm
lobes
long,
1
with
triplinerved
1—1.5
campanulate,
Thailand, Vietnam,
Collins
806
Spire
Ecology.
455
mm,
1.5—2
a
mm
third the
orange red
bright
447
s.n.
(L, P),
1909
Laos.
1712
(BM, K, P),
Smitinand
van
(L, P),
(BM),
Harmand
5099, 5099A, 12823,
Kerr
Beusekom
& Santisuk 2859
1291
(P),
Pierre s.n.
Evergreen
the
forest
(sometimes
eastern
mixed
dry
deciduous
range of distribution of M.
the
same
Laos
Thailand,
up
folium by
15
specimen (Kerr 5099),
locality
he determined
in its short inflorescence,
ceptional
to
single
a
x
a
to
cm
long
in
M.
mm
in size.
as
though
M.
1-1.5
and Vietnam also have
10
combination of the
10
1869
(all BM,
(P).
a
short inflorescences
narrowly
two
lanceolate
Kerr
leaf,
c.
m.
occurs
M.
(1938)
His
described
type specimen
other
(up
to
6
specimens
cm
also differ from
characters show
10—500
specimen {Kerr 5099A)
long. However,
smilacifolium). They
These last
similar
smilacifolium.
cm
forest),
smilacifolium
smilacifolium subsp. confertum (M. confertum Kerr) (fig. 3).
from
16448
(L, P).
(P).
Notes. Within
fruit,
re-
the Cocos Islands.
or
thinly coriaceous,
VIETNAM. D'Alleizette
posed
are
nov.
the
in
corolla tube. Fruit
K, P), Phengnaran &
from
Kurz
ripe.
Distribution.
this
For these
area.
ilicifolium
var.
thick, bark corky and light brown. Leaf narrowly lanceolate, 8.5—
long.
length
or
smilacifolium.
towards the apex. Venation
serrate
smilacifolium
the
be entire
Thailand.
mm
3—4.5(—6)
x
Bull.
the range of M.
smilacifolium,
correlate with other char-
Andaman and Cocos I. However, I have
confertum
stem
not
from
the distribution of his M.
from either the Andaman
5099
Subsp.
sely
with M.
reported
confertum Kerr,
Kerr
does
specimens
to
and M.
In M.
towards the apex, but may
serrate
confined
it
confertum (Kerr) Kiew,
var.
Old
is
nor
Martaban (Burma),
b.
obscurely
M. serratulum Hill
(1877)
Myxopyrum
randomly throughout
of serration of the leaf margin.
spinose margin, though conspicuous,
differences,
acter
apparently
varies,
degree
4
cm
some
wide,
long
subsp.
is
ex-
from
as
op-
smilaci-
and the
overlap (table
larger
1)
but
512
BLUMEA
if used
these
in
two
combination with
However,
9.5
the
even
length
(P).
cm
on
table
within the
The
flower
with lobes
in
fleshy
No.
length
1984
2,
one
can
in
most
range
can
of M.
be
the
on
distinguish
cases
from 0.3—0.5
which
Blumea
Steen.,
J. Arn. Arbor. 64
(Steen.) Kiew,
specimen)
to
that the great
4
cm
(K)
up
species
to
rank
of the
majority
the combination of characters
is confined
distribution of M.
mm
to
a
relatively compact
smilacifolium
size
in
(table
(0.3—1
15
mm
area
in
is matched
these
a
campanulate
subsp. confertum
M.
long)
nervosum
which
has
scarcely
by
species
two
to
a
that
also
corolla
0.5—2.5
thick
and
open.
SPECIES
(1967)
(1983)
1),
possesses
length
open recurved:
EXCLUDED
enerve
down.
smilacifolium.
Myxopyrum smilacifolium
(ranging
breaks
length,
wide range of inflores-
a
be maintained at
cannot
grounds
Besides the differences in fruit
form.
BM
distinguished using
urceolate corolla tube with lobes
Myxopyrum
the
(in
Kerr,
occurrence
widespread
smilacifolium)
subsp.
cm
confertum
level
inflorescence
in Thailand, shows
Kung
from 2.5
that their
and
variability
differ
mm
confertum
greater
nervosum.
in
Klawng
subspecific
to
1 and
of M.
tube
29,
reliable character,
most
In view of this, M.
specimens of M.
in
inflorescence
single plant
a
and is here reduced
given
VOL.
taxa.
Kerr 6040, collected from
cence
-
152,
fig.
3,
=
Chionanthus enerve
589—596.
ACKNOWLEDGEMENTS
The
abled
tors
author
of the
permission
ting
loans:
Paris,
is
this work
and
grateful
to be
Herbaria
to
work
Arnold
to
Universiti
completed,
at
British
in their
Pertanian
Professor
Museum
institutes
Arboretum,
Rijksherbarium
to
Leiden.
of
Malaysia
Botany
(Natural History)
and
National
to the
Curators
Herbarium
for
of
granting
sabbatical
leave which
Cambridge University
and
Royal
of the
Botanic
following
Washington,
New
Gardens,
herbaria
York
en-
and the CuraKew for
for facilita-
Botanic
Gardens,