Ada
Bot. Neerl.
22(4), August 1973,
Epidermal
p. 329-359.
leaf
characters
Malesian
M.G.C. van
Staveren
Rijksherbarium,
Leiden
of
the
*
Icacinaceae
and P.
Baas
SUMMARY
The cuticular
and
Continental
Asia have
different hair
slender
roids
those
types
in the
of Pennantia
been
hairs
cuticular
of
and
of
two
and
described
of the
part
as
to 25
Malesian species belonging
Australia
are
for this
Gonocaryum
flanges
all
of
from
studied
recorded
are
papilla-like
the leaves
of
characters
Icacinaceae
in
Zealand
detail.
family.
hydathodes
Stemonurus
New
is
species
Five stomatal
types
challenged. Penetrating
reported
for
this
the
from
and nine
of the
interpretation
Haberlandt’s
are
of
genera
some taxa
and
asteroscle-
family
for the
first time.
synoptical key
A
nostic
caryum,
The
and the
Bailey
that the
and
1.
and
lodeae
of
with the levels
and
of
of
and Howard. The
paracytic
this
that
characters
anisocytic types
on
the
basis
of the characters
study
distribution
more
the indication
isolated
indumentum
specialization
and
found
of
position
in
a
only
close
of the
be treated
stomatal
is
given.
The
diag-
as a
type
genus
do
types
primitive
over
separate
not
show
anatomy
the genera,
for Icacinaceae
between
relationship
genera Platea
wood- and nodal
of stomatal
are
characters
is discussed.
Mappianthusshould
anomocytic types
are
of cuticular
used
are
the
(all climbers),
impression
combined
correlation
value
implications
Taxonomic
Phytocreneae
by
to the genera
and taxonomic
and Gonofrom Iodes.
an
family
however, suggests
that the
and
absolute
for this
cyclocytic
derived.
INTRODUCTION
The
family
bution is
Icacinaceae
includes
c.
56 genera with
Sleumer
predominantly tropical.
(1971)
c.
300
revised
species.
Their
distri-
for
part of the family
Flora Malesiana.
All
and
of the Malesian Icacinaceae and those of Pennantia from Australia
species
Zealand
New
Such
preparations.
were
a
studied
of indumentumrecorded for the
and
because
of the
for
leaf
family by
suspected
range
the results
of this
characters
epidermal
study seemed rewarding
in
cuticular
because of the great range of type
Heintzelmann & Howard
in stomatal types
(cf.
Baas
(1948)
in Sleumer,
1971).
It is
hoped
that
study
will be of
some
fication of sterile material and of fossil leafremains belonging
A further aim of this
between
xylem,
leaf
as
study
epidermal
suggested by
was
to
*
This paper
on
reports
of the second.
their
initial
steps
the
specialization
and the
Bailey & Howard
results
of
It is dedicated
in the field
of
a
to
for the identi-
the Icacinaceae.
establish whether
relation could be found
any
The anatomical literature has been
vision
use
specialization
of the
node and
(1941).
comprehensively
post-graduate study by
summarized
the first
author under
to Prof. Dr. W. K. H. Karstens who
general plant anatomy.
guided
Sole-
by
the
both
super-
authors
330
M.
reder
&
(1899
Metcalfe & Chalk
1908),
For the indumentum the
for the
starting point
2.
MATERIALS
For
our
study
within
variability
an
area
60 °C in
mixture of
a
glacial
mounted in
The
acetic
used
are
In the Icacinaceae
major
and
eye
the
10
Apodytes
a
at
at
occur
the
1927)
be
X
or
is
on
was
types.
concen-
and
%,
areolae.
Malesia,
KEY
p.p.).
surface; only
in
Pyrenacantha
The
stomata are
They rarely
occur
in groups.
an
of the genera
species
for the
measured,
stomata were
near
always
measured with
stomata were
For each
and
on
almost
remaining
stomata, present in many species,
of which
21.
dimidiata,
In A.
were
not
the range and
descriptions
were
a
Continental Asia
a
and
of the lamina
rather variable
and
Winteraceae
Africa,
species
the
(Jansen
&
with
stomatal
species
remaining
This invalidates claims
diagnostic
Cotthem
Icacinaceae
areas
110
Baas,
it did
species.
in
the
also indicates that the stomatal index may
species.
a
the stomatal
examined,
Because of the great range in this
preparation)
van
THE
have been found
stomata
In the anatomical
Celastraceae
of
Malesian
description
genera. Herbarium
calculated. For that purpose 25
variable within
terminology
In
and
given.
specimen.
and Bongers, in
extremely
the abaxial
magnification.
water
AND
II and III
adaxial surface.
1000
index is
past that the stomatal
The
margin,
macerated
was
specific
larger
determine the stomatal indices for the
to
Current research
press,
5
(1969).
and width of the
indices varied between 7 and
useful
on
some
dimidiata, 11 specimens
distribution in
not seem
midrib and leaf
of each
for the
DESCRIPTIONS
least in the
Giant
examined in each
wide
THE
measurements.
index (Salisbury
were
IN
on
species
average for each
For
and
studied.
were
% hydrogen peroxide
beginning
Sleumer
and Citronella25
stomata.
included in these
by
as
length
micrometer
Apodytes, Cantleya
species
6
solution of Sudan IV in alcohol 70
the
at
reticulata
Miquelia
species
piece
dimidiata and
it
of
impression
some
Subsequently
water.
generic description
stomata
distributed,
For all
a
gain
Apodytes
middle, including
complex (plate I,
and minor veins
randomly
of
volumes of 20
in
for each of the
herbarium-specimen
latifolia, respectively,
boiled in
listed
are
USED
The stomatal
repanda
was
equal
abbreviated
CHARACTERS
3.1.
an
specimens
acid, stained
for small genera, after the
3.
(1971).
essential
an
glycerin-jelly.
specimens
numbers
11
from the
taken. All the material
trated
BAAS
study.
the 26 genera examined. To
species,
a
From each leaf
was
taken from
was
to
of Iodes cirrhosa and Platea
specimens
at
leaf
one
is
P.
METHODS
species belonging
111
and Baas in Sleumer
(1950),
of the present
interpretation
AND
STAVEREN AND
Heintzelmann and Howard
by
paper
VAN
G. C.
(1971)
the
character
was
stomatal
used
at
the
by
authors in the
species level.
for the various
complex
may
be
stomatal
anisocytic.
anomocytic,
cyclocytic, helicocytic,
Cotthem
van
paracytic
deviate from Stage
we
intermediate between
same
size
treated
are
the
as
mediate types the
which
stains
and
and
cell
guard
unspecialized
that the genera in
27 reference is
fig.
stomatal types; this may
or
deviating
cells,
cuticular
that
mean
question
made
actually
cuticle
a
striated
a
with intermediate
stomata
one
and
stomata
occur
in this respect
are
defined stomatal
clearly
or
unspecialized cells.
intermediate stomatal types
show both types of
the genera with each
intermediate between
to
such inter-
have
have
as
cells
characters,
In
may
or
here
subsidiary
cyclocytic.
pairs
cyclocytic
regarded
are
contrasted with the smooth cuticle of the
as
discussion
In the
than the
size
Stomata with
but with
the
intermediate type. Like
an
equal
anomocytic
surrounding
deeply
of
of
the definition of the
cyclocytic.
unspecialized cells,
cells
more
cuticle
granular
and
anisocytic
intermediate between
as
cells
or
in
(1965)
type. Stomata with three subsidiary
of the
331
CHARACTERS OF THE MALESIAN ICACINACEAE
LEAF
EPIDERMAL
type
only.
In all
the
species
inner stomatal
1971)
ledges
stomatal
and
only clearly
are
remains
chamber)
synoptical key
possess
In cuticular
present.
are
stomatal
stomata
ledges {fig. 10)
attached
species
ledges
where
The presence of
a
in the
very
common
only
recorded when
The
cross-piece
of
they
were
cells
distinctness.
The
Hairs
were
mostly
are
and
the
implies
the
descriptions
in fact
more
that inner
obvious than
ledges
of
the
a.
the
a
peristomal
on
to
9).
at
normal
hairs,
and i.
Heintzelmann and
They
b.
hairs,
the
These
that
is,
cells
or
was
present
are
absence
stomatal
poles
polar T-pieces
when both the
counted. Often the
radiate cuticular
rim
of
is
were
upright
III and
the abaxial
f.
enough
in
surface,
some
may
species
scarce.
mainly unicellular
hairs,
peltate-stellate hairs,
Howard,
When
walls
varying
intergrade
IV)
malpighiaceous
multicellular,
of
striations
of Heintzelmann & Howard
are
not.
or
periclinal
{fig. 9).
that surface and may be very
terminology
clustered
hairs.
T-piece
cells may be submersed
neighbouring
subsidiary
confined
papilla-like hairs,
ceous
leaves these inner
present together.
followed. Hairs in the Icacinaceae
e.
or
8 and
distinctly present,
indumentum(plate
hair types
hairs,
(Wilkjnson
therefore also
Concentric cuticular striations if massive
the midriband veins
types:
the
are
possible
(see fig.
concentric
possess
with the presence of
to
In
unnoticed because
T-shaped thickening
Icacinaceae
the number of
these
3.2.
remained
rim
also
species
some
cutinized part of the
flap (a
ledges
which
remains,
themselves. It is
they
cells and
subsidiary
possible
of Icacinaceae
stomatal
them.
to
In
ledges.
peristomal
a
cuticular material.
adhering
and the
a
the presence of inner cuticular
the inner stomatal
some
stomatal
preparations
visible if
presence of these other cuticular
in
outer
occasionally
defined a thin-base hair
“a
restricted
was
glandular hairs,
(see plate
largely
following
d. uncinate
globular hairs,
g.
distinguished
as
(1948)
and of the
c.
uniseriate hairs
moreover,
even
For the different
h.
III and
slender
IV).
thin-base and icacina-
long, tapering
hair with
a
thick
332\I
M. O. C.
Plate I.
Fig.
1.
Cuticular
Apodytes brachystylis,
stomata
with
surrounded
fig.
5.
preparations,
by
partly
x
radiate cuticular
arrows,
6.
Stemonurus
stomata
stomata;
subsidiary cells; fig.
striations; fig.
Rhyticaryum longifolium.
(arrow); fig.
P.
BAAS
200.
anomocytic
submersed
VAN STAVEREN AND
adaxial
grandifolius,
predominantly cyclocytic.
3.
2.
Rhyticaryum elegans,
Gomphandra
4. Medusanthera
surface
sites
fig.
of
showing
anticlinal
cyclocytic
water
laxiflora, anisocytic
pitted
penetration by
comosa,
cuticular
asterosclereids
stoma
stomata;
flanges
indicated
with
EPIDERMAL LEAF CHARACTERS OF
Plate II. Sections
and cuticular
Fig.
cunninghamii, paracytic stoma; fig.
7.
Pennantia
stomata with
surrounded
latifolia,
stoma at
verse
section
proper
by
low
through
and outer
glandular
preparations,
conspicuous T-pieces; fig.
T-pieces
hair.
concentric
focus
9.
X
900.
8.
Gonocaryum gracile, fragments
Gomphandra coriacea,
cuticular
showing
upper
333\II
THE MALESIAN ICACINACEAE
striations,
inner stomatal
cuticle
ledges; fig.
epidermis, showing granular
epidermal cell wall; fig.
12. Platea
also
excelsa,
stoma with
very
granular; fig.
11.
of two
faint
10.
Hartleya inopinata,
cuticular
transverse
layer
in
between
section with
polar
Citronella
trans-
cuticle
unicellular
334\III
M.
Plate
Fig.
III.
13.
fig.
with
not
hair with
15.
body
bulging
abaxial
and
basal
hardly
latifolia,
a
with
base
fig.
one
18.
and
minute
well
C.
VAN
STAVEREN
AND
P.
BAAS
200.
submersed
much restricted
part;
surface.
or
thin-walled
Citronella
malpighiaceous hair
walled
x
Gonocaryum macrophyllum, papilla-like
cyclocytic
normal
preparations,
Cuticular
G.
a
at low
(a)
and
subsidiary cells;
fig.
14.
second
hair
arm
stub. Hair
malpighiaceous
developed
and
lumen; fig.
one
17.
Miquelia celebica,
hair;
less
high (b) focus,
insertion
fig.
16.
developed
and
indicated
by
Gomphandra
arm.
Pyrenacantha repanda.
uncinate
stomata
Gomphandra fusiformis,
globular
Hairs
with
normal
hairs
arrow;
papuana,
a
hair
on
thickwith
midrib
a
of
OF THE MALESIAN
EPIDERMAL LEAF CHARACTERS
wall and restricted lumenand
by
us
found
we
thickness and
and
we
without
a
very
restricted
a
thin-walled base and with
a
do
or
a
normal hairs
lumen and hairs
lumen which
without
a
thick-walled
and
body
suggests. This hair type differs from
of
second
a
14 and
(see fig.
stub
arm
sitions between normal hairs and
with
in
second
a
with
real
well
one
papuana
developed
small,
very
either thin-walled
the
in
presence of
the
a
the
lodeae
useful
found
we
less
one
here called
15).
presence
gradual
tran-
normal hairs
developed
developed
arm
are
hairs
as
-
hairs
the
arms,
hairs
Malpighiaceous
the
as
some-
be
may
restricted lumen.
very
a
which
recognize
an
only
can
be
separated
icacinaceous hair type. In
this type is called
synoptical key
the
malpighiaceous
well
two
a.
malpighiaceous
transitions,
to
normal hairs
Phytocreneae
normal hair with
a
layer
hairs,
the lodeae and
of the
hairs with
exhibit and
may
in
a
numerous
pits
which
is
very
is
the
in the wall of that part
(see
(except
normal hairs also
cuticular
possess
outer
Uncinate
the
and
basal part and
bulging
fig. 17). Many
on
(see fig.
mentioned
seem
character which
Another
ness
not
only by
stub.
arm
common
thick-walled with
and the
descriptions
second
in
or
it does
-
The real
as
of the above
Because
arbitrarily,
24)
hairs with
c.
(see fig. 23).
Citronella
well and
one
16 and
and
arm,
hairs
malpighiaceous
times
(see fig.
not so common
are
malpighiaceous hairs, namely
b. hairs with
stub,
arm
Gomphandra
but between normal
hairs,
Furthermore
25).
body
Therefore
not.
or
thin-walled base.
a
normal hair
a
normal wall
a
thick-walled
a
restricted
was
The icacinaceous hairs found in 9 of the 26 genera
name
with
with
between normal and thin-base
distinguish
not
hairs with
thin-walledbase”. In the material examined
transitions between
many
335
ICACINACEAE
Mappianthus),
of
markings
distinct-
varying
body (see fig. 25).
recurved
a
tip (see fig.
18 and
26), only
Uncinate hairs may also possess
Phytocreneae.
a
occur
bulging
in
basal
part.
Globular hairs
They
were
are
always
also very
common
in the lianoid genera (see
Clustered hairs consist of
groups
of hairs fused
central hair is sometimes several times
the
cluster and sometimes has
of the
fig.
a
as
long
at
the base
and thick
cuticular
verrucose
only
occur
considerable part of their
19 and
26).
(see fig. 19).
The
the other hairs of
as
markings
the
on
outer
layer
in Platea and consist of
body. They
we
also found
(1899) (see fig.
cellular hairs.
glandular hairs,
12 and
He
19).
to
spherical
group of cells fused
to
the
surface
(see
Heintzelmann & Howard
noted before for Platea excelsa
called this
(1948)
hairs in their material. We agree with them when
authors
a
appressed
are
20).
In Platea
Solereder
consider the
hairs
suggestive
globular type
of Platea have
between the wall and
very
18 and
body.
Peltate-stellate hairs
over
fig.
found in combination with other hair types.
of
a
cuticle,
a
as
thick
filled with
glandular
nature.
did
they
glandular
cuticle and
granular
hair type
not
find
truly glandular
the idea of
say that
is incorrect.
a
small
material
by
uni-
spherical,
some
However,
the
the
top
space
at
(see fig. 12),
which is
336
M.
The slender
to
a
found
papilla-like hairs
hydathodes
as
by
excentric
fig. 13
a
the
on
and
in
position
b, 21
and
water
guttation (cf.
also Esau
1964).
lower leaf surface noticed
stomata
water
are
The
by
The
The anticlinal walls of the
clinal
walls with thin
cuticular
in
as
the anticlinal and
lying
The
flanges,
pitted,
granular
cuticle,
In
in
as
nature
cells
to
the
1955)
of
cells
this
flanges
rounding
the
19.
Fig.
X
mis,
x
Platea
hair,
arms.
hair
arms
the
with
bullata,
1000;fig.
one
well
hairs
Miquelia celebica,
apex,
midrib
smooth,
and
veins,
The cuticle
granular,
are
over-
striated.
or
underneath the
epidermal
It is
x
cells
asterosclereids
are
are
of clustered
of
hair, peltate-stellate
peltate-stellate
radiate cuticular
hair.
Note
striations,
Gomphandra montana,
indicated
developed
with
a
by
arrow,
and
second
uncinate
pitted,
modified
of the cuticle
growth.
the
by
(fig. 6).
Foster
(1947,
(Rutaceae)
and Mouriria
grandifolius
the anticlinal
but
that the
flanges
sur-
not.
Gonocaryum gracile, papilla-like
23
intrusive
that in S.
noteworthy
is
pattern
flanges
photomicrographs.
detail
detail
21.
insertion
cum, normal
of curved
of the
by conspicuous
x
Hair
cells
The anti-
loops.
of the cuticular layer
for Boronia serrulata
unspecialized
electron
20. ibid,
500; fig.
surrounded
like
structures
of the
Scanning
250; fig.
because
straight,
may be
in the
overlying
penetration develops through
(Melastomataceae).
cuticular
Plate IV.
from the
as
species unspecialized
of cuticle
may be
structure
Stemonurus
recorded similar
huberi
well
that
for the
p.p.)
some
intrusion of asterosclereids between the cuticular
Presumably
likely
Gonocaryum.
longifolium (see fig. 5).
periclinal walls
is due
most
seems
pathways
as
both surfaces
on
In
areas
of
especially
Rhyticaryum
it
us
seems more
serve
droplets
mornings
unexplained, however,
I and II
sinuous.
periclinal
the
been found in Pennantia corymbosa.
only
(plate
or
in
from the upper as
guttation
protrudes
water
Hartleya inopinata (see fig. II).
species
two
Gonocaryum
unspecialized
slightly curved, slightly sinuous,
sometimes
It
water.
Haberlandtremains
cells
of
To
guttation.
guttate
absent from the upper leaf surface in
unspecialized
have sinuous
for the
can
uniseriate hairs have
Multicellular,
3.3.
structures
cells. From
structure
presence
Gonocaryum pyriforme
also present in
stomata
papilla-like
BAAS
with
structures
unspecialized epidermal
slender
a
responsible
as
that these slender
giant
cell
P.
have been referred
unicellular
are
Haberlandt noted the
22).
these hairs
regarded
this
lower leaf surface in
upper and
unlikely
the
&
VAN STAVEREN AND
C.
Gonocaryum
They
broad basal cell part inserted between the
an
in
only
Haberlandt (1894).
G.
and
250. Some
one
arm
x
less
stub
globular
of the leaf
hair
x
(p)
24.
developed
and
arm,
surfaces
vein.
with
glandular
of
water
22.
ibid.,
hair with
hairs
(g),
underlying epiderstoma
detail
two well
(w),
of
both
papilla-
developed
Gomphandrapapuana, malpighiaceous
verrucose
on
and
and
250; fig.
malpighiaceous
250; fig.
hairs
hairs
striated cuticle
x
250; fig.
cuticular
Uncinate
25.
Natsiatum
markings,
hair
fungal hyphae.
viewed
x
herpeti-
85; fig.
on convex
26.
side
EPIDERMAL
LEAF
CHARACTERS
OF
THE MALESIAN ICACINACEAE
337\IV
338
M.
3.4. The
patterns
veins in cuticular
that the
(usually
4.
I.
in
ANATOMICAL
APODYTES
Stomata
less
a
abaxial
A.
with
studied.
F.
the
2.
one
and
289 ;
B.S.
S.
24941',
Sudan
with
IV).
2.
CANTLEYA
1.
C. corniculata
Material
Stomata
and
a
thin-walled
surface
3.
cells
with
polar
s.n.',
Wylie
40872',
cells
Indumentum:
markings
sinuous
more
or
septate
without
walls, mostly
with
mostly
lumen
cells of adaxial and
pitted
cuticular
hairs
the
on
confined to abaxial
body.
surface,
Unspecialized
cells
of
walls.
the
with
cuticular
R.
F.
wide;
ixrn
guard
on
to
Java: Den
Lam 6048 ;
Thailand:
s.n.
hairs
less
cells
here
and
markings
on
also
the
on
Larsen
504.
length
27-30 [xm,
with
different
surface
with
sometimes
the cells
c.s.
786 ;
UNESCO
Philippines:
average
width 22-
of the
stomata
for
staining properties
one
body. Unspecialized
walls,
the stomata and
227 and
conspicuous grouping
adaxial
sinuous
mans
Rudatis
3616,
average
more or
abaxial
slightly
D.
Koster
Berger 605,
Natal:
with
or
more
and
cells of adaxial and
cuticular
overlying
septa
striations
on
the midrib.
Howard
Sumatra:
[xm
with
base.
curved
paracytic;
Grashoff F.R.I. 36E,
21-24-28
long,
cells with
striations
to abaxial
C1TRONELLA
Stomata
wide.
cuticular
slightly
20-24-29
surrounding
(Becc.)
27-31-35
confined
sinuous
slightly
abaxial
with
1380.
p.m
Madagascar:
straight
subsidiary cells; guard
hairs,
with
major veins,
body. Unspecialized
with
and
and
cells
distinctness. Indumentum
varying
Ridl.
studied.
subsidiary
of adaxial
with
Clemens
Wylie
verrucose
with
cells
epidermal
walls
of
Arn.
ex
long,
Indumentum:
surface
of the
curved, rarely
or
surface
surrounding
or without
abaxial
midrib
to
layer
outer
verrucose
Pierre 583 ;
jxm
polar T-pieces
confined
23-27-30
and
Meyer
786
cells
mostly
long,
Africa;
Rudatis
(xm;
and
(fig. I)
Borneo:
27-29-35
(epidermal
the
E.
Cambodja :
Stomata
26
(xm
studied:
genera.
prominent.
M.
abaxial
A. dimidiata
Material
v.
septum each,
adaxial
arranged differently
1)
(fig.
Queensland: Kajewski
Stomata 27-29-32
with
lianoid
ledges
verrucose
straight
veins not
brachystylis
Material
of veins
prominence
are
irregularly grouped, anomocytic; guard
or
anticlinal
striations;
Minor
flanges.
Am.
ex
hairs
slightly
or
surface
cuticular
Meyer
distributed
of normal
smooth
of
prominence
DESCRIPTIONS
outer stomatal
conspicuous
and
1.
E.
randomly
consisting
P. BAAS
cells in the areolae. Prominent minor veins
epidermal
in the
common
the minor veins
overlying
the
study. Only
recorded. In this
paper
was
cells
from the
rows)
examined in this
not
were
preparations
epidermal
particularly
are
C. VAN STAVEREN AND
veins
Venation
means
G.
over
D.
T-pieces, mostly
cyclocytic,
outer stomatal
locally
thin
cuticle.
surface and midrib of adaxial
and
Don
guard
the
664.
wide,
conspicuous
Unspecialized
walls
|xm
cells
with
(fig.
10
cells with
with
a
on
adaxial
faint cuticular
and
to
Indumentum
surface,
surface
3
with
striations.
5
with
a
polar T-pieces;
consisting
thick-walled
straight walls,
Minor
submersed
partly
and small
veins
not
on
of normal
body
the
and
abaxial
prominent.
15)
conspicuous
strongly
with
ledges
outer and
cutinized
inner stomatal
stomatal
ledges
chamber; subsidiary
and without
cells
partly
LEAF
EPIDERMAL
submersed,
cuticular
CHARACTERS
of
one
them
striations.
the whole
lamina
consisting
of normal
and
adaxial
striated
to
due
1.
C. latifolia
32-39-42
striations.
cells
of adaxial
granular
of cells
2.
C.
over
on both
lucidula
32-38-40
walls,
these cells
with
faint cuticular
striations.
adaxial
cular
with
striations
granular
4.
Indumentum
surface
over
on
C. suavolens
Stomata
28-32-37
of
consisting
adaxial
side,
4.
guard
peristomal
rings
of
rim
the whole
Unspecialized
to curved
on cells
1.
C.
surface
walls
cells
and
the
and
Stomata 24-27-28
to
the
obscured
hairs
on
walls;
adaxial
concentric
by
only. Unspecialized
cuticle striated
side;
anticlinal
and
walls
of the cuticle.
coarsely granular structure
9550.
with
on
and
cells
without
few normal
a
the
overlying
with
abaxial
midrib;
of cuticle
structure
cuticular
hairs
of
straight walls,
cells
granular
wide; subsidiary
and
both
on
cells with
malpighiaceous
of abaxial
side
cells
with
hairs
slightly
overlying
anticlinal
sides;
striations.
a second
side
with
anticlinal
overlying
arm
slightly
walls
of
these walls.
the
walls
faint concentric cuticular
only. Unspecialized
sinuous
midrib
not
walls;
with
adaxial
on
cells
of
faint cuticuticle
side;
interrupted.
NGF17332.
23-27-30
midrib,
only,
absent
to curved
straight
with
without
wide,
ixm
hairs
walls,
granular
a
cuticular
from
with
cuticle
striations.
midrib.
Indumentum
Unspecialized
cells
faint cuticular striations
over
periclinal
walls
on
on
of
the
abaxial
walls.
present,
on
5
hardly
cells
rarely
not
on
submersed
or
sides,
often
the cells
or
of the
on
divided
for Sudan
consisting
with
subsidiary cells;
polar T-pieces
staining properties
straight walls,
striations
veins
to
rather distinct
Indumentum
of both
with
of small
without
abaxial
the whole
a
lamina
that
IV from
normal
second
surface
and with
so
arm
with
or
a
two
un-
hairs
stub.
straight
especially
prominent.
Howard
Andaman: King's
[xm
different
striations.
the midrib
Minor
3
ledges,
subsidiary
with
surface
cuticular
hairs.
with
cyclocytic,
cuticular
and
with
to
outer stomatal
guard cells;
are
(Kurz)
S.
prominent.
veins not
1983.
of anticlinal
of adaxial
surrounding
studied.
on
with
with
lamina
cells
often
Howard
around
andamanicus
Material
long,
conspicuous
subsidiary
granular
wide; subsidiary
of small
surface
Guinea:
overlying
specialized cells,
over
walls,
walls
interruption
with
cells
curved
slightly
to
and
nomine
24-27-30 (xm
grouped, anisocytic
cells
cuticle
seemingly interrupted
Minor
malpighiaceous
due to
striations
malpighiaceous
CODIOCARPUS
Stomata
cells of abaxial
walls;
Howard
|xm
small
cells
without
abaxial
New
and abaxial
epidermal
curved
(BÍ.)
studied.
side
surface
due
consisting
periclinal
Material
straight
(xm
Isl.: BSIP
long,
[xm
of
also
rarely
Howard
Solomon
Stomata 33-36-40
sine
of adaxial
(A. Gray)
studied.
sinuous
sometimes
subsidiary
of small
malpighiaceous
seemingly interrupted
C. samoensis
Material
wide;
(xm
abaxial
25-27-30
of small
cells
Unspecialized
indumentum
IS)
consisting
Caledonia;
long,
(xm
surfaces
Howard
New
consisting
Indumentum
24-28-31
on
walls
both
Unspecialized
slightly
with
species
some
on
24557.
surface with
walls
midrib;
these walls.
overlying
10 and
(fig.
B.S.
in
pore,
surface.
with
anticlinal
seemingly interrupted
(Sleum.)
Stomata
3.
abaxial
sides
studied.
curved
long,
periclinal
Material
stub.
Howard
Indumentum
and
of cuticle
structure
(xm
from the
curved, rarely
or
periclinal walls;
Philippines:
cuticular
absent
midrib, rarely
the
to
malpighiaceous hairs
of small
consisting
straight
over
(Merr.)
Stomata
subdivided perpendicular
confined to midrib of abaxial
hairs
with
granular
studied.
the
on
granular
coarse
Material
and
surface
and
often
Indumentum
339
ICACINACEAE
MALESIAN
OF THE
long,
20-24-27
coll.
|xm
s.n.
wide; subsidiary
cells often subdivided.
Indumentum:
340
M. G. C.
hairs
with
or without
the cells
on
2.
C. merrittii
with
of the whole
second
a
abaxial
studied.
Stomata
41-44-47
sides
of the
adaxial
6.
cells
of
in
in
stub,
tum
with
cells
of
montana.
with
these
cells
these
walls.
rostrata.
Material
1. G.
G.
to
711;
G.
and
G.
with
hardly
corners;
cells
submersed
polar T-pieces.
noted
arm,
on
both
on
sides
both
those
straight walls;
with
one
24,
with
3
see
(4)
also
not
in
a
smooth
with
a
two well
11
base;
surface
thin
and G.
to
Cork
straight
concentric
with
concentric
with
species
in 4
arms.
simalurensis)
0;
a
Hairs
to
in G.
mostly
walls, rarely
with
on
straight
both
walls
sides
anticlinal
coarsely granular
abundant
warts
species
most
second
G. papuana.
of sinuous
loops
periclinal walls;
over
finely
in
rarely
a
indumen-
species
curved
to
walls,
of cuticle
areas
cuticle
due
with
small
less
or
finely granular cuticle
developed
to sinuous
submersed
without
or
species
or
I)
hardly
more
in
hairs,
thin-walled
table
or
with
ledges,
cells of some
with
a
or
curved
striated
prominent.
and
subrostrata. G. tomentella,G. velutina,
with
mappioides,
not
veins
with
of normal
and
body
of adaxial
species
sometimes
cuticle
quadrifida
to
(i
(G.
with
walls
of
overlying
and G.
sub-
remaining species.
Merr.
Andamans:
Poilane
Merr.
13. G.
18354;
Merr.
papuana
Sleum.
some
absent in
(BÍ.)
G. javanica
Korthals
cells
developed
striations
cuticular
rims; subsidiary
in G.
surface
7
to
and faint
surface
stomatal
outer
varying distinctness,
abundant
23
14, 16,
helicocytic,
species consisting
23467 ;
coll.
Borneo:
C.
I
15. G. montana
G.
Borneo:
Guinea:
22.
New
australiana
: Loerzing
Wight.
G.
Merr.
Sleum.
14. G.
29394;
New
oligantha
Guinea:
Sleum.
19.
G.
Simalur
Isl.:
Sleum. New
Merr.
PNH
13617;
24.
mappioides
Carr
Achmad
G.
Sumatra: Loerzing
;
21.
64;
Guinea:
26.
comosa
433
9.
G.
4066 ;
Celebes:
16. G.
Valet.
oblongi-
12508;
18.
Sumatra:
quadrifida (Bl.)
Gjellerup 608;
Sleum.
G.
6.
Philippines:
Elmer
G.
;
Sleum.
13511;
Valet.
14501;
simalurensis
6685
K.D.
Sumatra: Meyer
parviflora (BÍ.)
(Birnie)
Guinea:
(Gagnep.)
Philippines:
Sleum.
subrostrata
Wight
12. G. luzoniensis
r(Merr.) Merr.
NGF31689;
Carr
India:
Philippines:
G. sawiensis
Guinea;
F.v.M. New
7009 p.p. ; 4. G.
donnaniensis
G. fusiformis■
(Schellenb.) Sleum.
pseudojavanica
561;
10.
6205;
7.
2814;
Clemens
17. G
.
(
2. G.
coriacea
flavicarpa (Elm.)
38543;
Sleum. New
Sleum.
25.
PNH
G.
Hallier
Valet. Java: Kostermans
20, G.
5.
1890;
13584;
Becc. Sumatra
Philippines: Elmer 8620;
Sumatra: Iboet
Achmad 1534;
8.
PNH
Vriese)
& de
lysipetala Stapf.
(Becc.)
schoepfiifolia
King's
F.-Vill.
Philippines:
882b;
Philippines:
I (Jungh.
capilulata(
fuliginea (Elm.)
folia
cells:
onthe
studied
Indo-China:
G.
in
lysipetala.
Minor
South
Hoorders
be
distinct
thick-walled
of abaxial
cumingiana (Miers.)
F.B.
24
a
Unspecialized
walls;
Lam 554 ; 3. G.
11.
surface,
apoensis (Elm.)
King.
in
with
cell
4
ledges
well
one
abaxial
at the
conspicuous
often seemingly interrupted
rare
striations
prominent.
of cuticular
of
with
(fig. 3, 9,
rather
to
with
cyclocytic,
and
rounded
malpighiaceous hairs
granular
cuticular
outer stomatal
adaxial
on
Lindl.
with
striations
walls;
curved
luzoniensis,
G.
faint
species
to abaxial
sinuous
slightly
cells
16
consisting
and G.
with
cells
veins not
ex
giving the impression
confined
wide,
conspicuous
thick walls
Wall,
others. Indumentum
the
arm
rarely subdivided. Indumentum:
7207.
am
anisocytic, occasionally tending to
radiate cuticular
or
with
Minor
subsidiary cells; guard
circles
Lam
malpighiaceous hairs
with
cuticle.
polar T-pieces,
cells
cells with
Tieghem ex Engl.
v.
30-35-39
Unspecialized
GOMPHANDRA
Stomata
Unspecialized
Tiegh.
long,
jjm
surface
finely granular
wide; subsidiary
ij.ni
stub.
arm
Caledonia:
consisting
of lamina.
striations,especially
91272.
PNH
(Baill.)
New
subsidiary cells; guard
Indumentum
with cuticular
surface.
surface.
G. austro-caledonica
1.
Material
Unspecialized cells
of abaxial
21-24-25
long,
GASTROLEPIS Van
5.
hairs
Philippines:
Stomata 24-27-29 p.m
mostly
the
BAAS
P.
Howard
(Merr.)
Material studied.
hairs
second arm stub.
a
surrounding
AND
VAN STAVEREN
Simalur
23.
Isl.:
tetrandra (Wall,
in
EPIDERMAL
LEAF
CHARACTERS OF
THE
341
ICACINACEAE
MALESIAN
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3
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rt
±
rb
+
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±
+
,
walls
—
of loops
u
cuticle
g
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4- ±
±±
±+
± —
±± ±±
±± ±+
|
-=
anticlinal
.
—
21-
22- 29- 20- 21- 23- 21- 20- 30- 24- 23- 22- 21- 24-
21- 31- 22-
=
tables. cone.
in
abaxial;
and
other
in
abreviatons curved;
=
Quantiave
species
slightly
apoensi australi na capitulat a comosa coriacea cumingxa don a iensflavicar ffuullmiginneeafusiformis avjanicaluzonie s lysipetal map iodes monta naohbl ngifola oligantha pa uan a pseudoiavnc quadrifdas wiensi schoepfi olasimrnlauensi subrosta tertandra toment lavelutina adaxial: resp ctively.
pa
pa
a
a
vicar
a
fl
I.
(7
G.
Table
flor
flora
vi
parparvi
ab
is
wiens
sa
G. G.
G.
G.
G. G. G. G.
G. G. G.
G. G. G.
G. G.
G. G.
G. G.
G. G.
G. G.
G. G.
G. G.
G.G.
G. G.
G. G.
G. G.
G. G.
G. G.
G. G.
G. G.
G. G.
G. G.
G. G.
1. 2.
2.
1.
3. 4.
6.7.
7. 8,
8. 9.
9.
4. 5. 6.
10.
10. 11. 12. 13.
13.14. 15. 16. 17.
17. 18. 19. 20. 21. 22.
22. 23. 24.
24. 25.
25. 26.
26. 27.
27. 28.
of
Explantion
to
=
ad
straight arms,
342
M.
Sleum.
Roxb.)
Kurz
1.
813;
28.
Indo-China:
GONOCARYUM
HCPM
Miq. (fig. 8,
Stomata intermediate between
faint
distinct
to
varying distinctness,
in
macrophyllum
G.
deeply
sides
in
with
Sudan
consisting
13, 21,
and G.
walls
more
and
G.
of cells
hairs.
striations
of
and
with
with
anticlinal
cells
cells
on
walls
in G.
hairs
of adaxial
adaxial
with
one
also
with
or
striations
of
ledges,
cuticular
granular cuticle, staining
more
Indumentum
calleryanum.
in G.
with
walls
both
faintly
calleryanum,
straight
with
surface
or
submersed
hardly
rims,
noted
abaxial
of
or
of cuticular
surface
sinuous
or
of cells
Hook. Burma:
outer stomatal
giving the impression
except
curved to
3 to 7 not
mostly
irregularly
an
cells
in
II)
table
with
massive
very
distinctness, occasionally
varying
also
see
A few normal
slightly
the midrib
overlying
flanges; periclinal
cyclocytic,
with
lobbianum
unspecialized
papilla-like
surface
22,
and
less concentric circles
or
impressinervium. Unspecialized
of abaxial
walls,
and
cells
Mast,
(Kurz.)
BAAS
P.
2683.
polar T-pieces; subsidiary
IV than
of
G. tomentella
Kjellberg
anomocytic
neighbouring or subsidiary cells; guard
without
21.
2727;
Sleum. Celebes:
G. velutina
G. C. VAN STAVEREN AND
granular.
with
cuticular
veins
Minor
curved
cuticular
pitted
often
both
gracile,
anticlinal
loops;
often with
surfaces
G.
slightly
to
shallow
on
mostly
prominent.
Material
1. G.
PNH
6252;
Endert
Siam:
1.
3.
2443 ;
Put
Borneo;
8.
studied
calleryanum (Baill.)
G.
5.
G. litorale
1. G.
894;
Sibal
Table
inopinata
Meijer
Sleum. Celebes:
(Bl.)
4664
2.
;
6765 ; 4. G.
Cel
Sleum.
G.
Elm.
cognatum
impressinervium
IE/184;
6. G.
Sumatra: sine
lobbianum
8.
nomine;
Philippines:
Sleum. Borneo:
(Miers.)
G.
minus
(fig. II)
New
[an
Quantitative
Guinea:
long,
and
Hartley
29-33-37
11836.
wide, cyclocytic,
(am
qualitative epidermal
with
leaf characters
4
of
7
to
submersed
partly
Gonocaryum.
(g)
2
§
(s)
3 .
£
stomata
S
o3
g
m
of
°
N8 —
3 pm
sizes
in
C/5
■p3
8
«
species
species
T-pieces
—
t
|
S
'5
length
—1. G.
3
o
r> O
U»
3.
Ci l C
iS
o 2S 522
calleryanum
£tQ
523
3
cognatum
G. CTQ
go S'
gracile
-o*.
4. G.
S'3
s■>1
1 Ia'
impressinervium
<-*
5. G.
2 S'
litorale
6. G. lobbianum
On
O' 2"
S'S522
7. G.
00
8.
F5
ps
macrophyllum
V
II
C. minus
*3
40-
a
width
polg.ar
3
g
und lation anticlnal
“
M
U»
31-34-36
rw
ON
±/-
u>
33-36-39
r w
LO
K-if
H
+
ÍS
41-
±
+
+
1
47-50-55
■^i LA ? LA LA
33-36-38
U» U) w
00
+
+
±
If
-
U>
31» u>
KJ<-*
25-26-28
S)
00
±/if
T
U»
36-40-42
hJ
N>
27-29-31
NJ vO
-
+
+
u>
32- U» fUJ "J
25-28-31
N> LA K) CC
-
u>
33U) Li
U» la
—
K)
28-29-31
00 N> VO
45-48-52
LA -L
LA N)
U>
32-35-39
N) U> LA
f? EE
L*J LA
1
ft
irrt
1
1
1
+
1
1
i
1
ab
a*
03
±
If
1
±
If
1
1
1
1
1
f
+
-2.
ad
CL
03
—
I
1
“■
s H±
v> TO*
s/g
±
H-
1
1
1
1
1
+
h
1
g
veins
'53
3
>
c_>
iges
miInor
ab
cr
-g flai
pit
,g
s±
If
(A
ab
ab
+
+
H±
ad
ad
H±
03 a*
ab
+
s
(/>
HC/l
s/g ±
1
A
s±
HC
l
Í
+/±
3
"O
±
H-
1
1
cuticular
“
ted
ad
CL
03
I
s
cuticle and/or
73
aj*0'.n
3
S
| I
rim
| ||
cuticular
promine t
.S
ca
striated granular
walls
§ t
5-U.30
K)
2. G.
Kurz.
Sleum.
Sleum.
31-36-41
II.
Kostermans
macrophyllum (Bl.)
Sleum.
Material studied.
Stomata
Borneo;
Sumatra:
S 24582.
HARTLEYA
H.
Becc.
gracile Miq.
1
03 o*
ab
g
+
+
+
Hg±
TO
If
T
03 CL + 03cr
g±ad+ab±/CTQ
KOQ
g±
H00
g±
C/5
s±
H-
CA +
s+
I
1
-
cr
03
ab
+
+
+
LEAF
EPIDERMAL
cells
subsidiary cells; guard
cells
with
surfaces
both
walls
periclinal
9.
with
IODES BI.
tions
often
stomatal
yatesii
often with
between
confined
a
finely
to
varying
Material
with
pits
in
and uncinate
and veins
surface
walls
and
with
Minor
I. velutina
bulging
hairs in
surface
pitted
with
cuticle
sizes,
wall.
verrucose
I. ovalis.
a
overlying
of
cells
cuticle
of
over
with
in
3
anticlinal
surface
and
and on small
hairs
sinuous
outer
and
type
in
3
in
5
hairs
species,
intermediate
species,
anticlinal
of both
midrib
markings
surface with
walls;
cells
walls and
A
present
cells of adaxial
on
normal
present
markings.
species;
prepara-
distinct
cuticular
verrucose
Uncinate
cuticular
in cuticular
rather
normal, uncinate,
to abaxial
slightly
to
with
Globular hairs
Unspecialized
flanges
cells
subvillosa
var.
basal
straight
cuticular
granular
consisting
confined
species,
without
Unspecialized
with
completely submersed,
of two distinct
in
polar T-pieces; subsidiary
absent.
prominent.
Indumentum
of both sides.
coarsely granular
distinctness.
in all
body (except
of abaxial
walls,
sides
also
straight
walls
of
often with
with cuticular
striations
Steenis
and
of
strikingly prominent.
veins
studied
1. I. cirrhosa
Turcz.
Celebes:
Malay Peninsula: Sinclair
Table
present
bulging basal wall,
normal
of adaxial
curved
to almost
partly
sometimes
surface,
yatesii).
in
pits
5
polar T-pieces.
hairs
of the
to midrib
to sinuous
cells
without
layer
small
3 to
veins not
Indumentum
distinguishable, subsidiary cells; guard
normal
var.
table
with
of adaxial
the outer
of I.
III)
ledges,
and veins
on
also
(see
slightly
to very
and without
ledges
walls.
periclinal
Minor
343
MALESIAN ICACINACEAE
outer stomatal
(see fig. II).
hardly
globular hairs;
THE
over
straight
cyclocytic,
Stomata
with
cuticle
granular
a
OF
CHARACTERS
111.
Teijsmann
9875 ;
H.
B.
12624 ; Java:
Sumatra: Maradjo
Quantitative and qualitative epidermal
Van
12717
12717a;
253 ; Thailand; Hansen & Smitinand
leaf characters
12985 ;
of lodes.
83
flanges
so
sizes
sizes
of
°
in
(xm
p.m
o
s
3
<D
dJ
jC
J-
■S
£
'S)
T3
c
S
B
1. I.
/. cirrhosa
M
cd
C
normal
C
C
E
'5
o
g
c
§
3
V-
"c
-S
globular
3
3
3
CJ
C
cs
•§
-a
33
g
ad
ab
T3
pi§t ed
■«
2020-21-23
15-17-18
15-17-18
++
+
+
+
+
1717-20-21
15-18-20
15-18-20
+
+
+
—
±
—
16-19-21
13-16-18
+
—
—
—
9875
9875
1919-22-26
18-20-21
+
+
253
2121-23-25
17-20-21
+
12985
12985
2020-22-26
16-18-20
+
—
1818-19-20
14-16-17
+
O
o
20-22-24
16-19-21
+
+
O
o
1616-21-25
17-19-21
+
+
+
4. I. reticulata
15-18-20
13-16-18
13-16-18
+
+
+
+
—
5. I.
1919-21-23
19-21-23
+
+
+
—
1717-20-21
16-18-20
16-18-20
+
+
1818-21-25
14-17-19
+
+-
17-18-20
13-16-18
13-16-18
+
Teijsmann
van
van
id.
Steenis
Maradjo
Hansen
Hansen
ƒ.
I. ovalis
ovalis
Bakh.
v.d.
Loerzing
3. I.
I.
8. I.
1.
O
Brink
Brink
5709
5533
philippinensis
seguinii
6. I.
I. velutina
7. I.
12717
12717
12717a
Sinclair
2.
2.
12624
I
%
"5
wal§ls
—
und lation anticlnal cuticular
species
species
53
§
32
hairs
stomata
var.
subvillosa
subviltosa
var.
vitiginea var.
vitiginea
vitiginea
vitiginea
yatesii
=
var.
yatesii
intermediate type
between
normal
and
uncinate hair.
—
—
—
—
-
—
+
+
+
±
±
?
+
+
±
±
±
±
—
±
±
±
+
+
+
+
+
+
+
—
-
—
-
±
±
—
—
-
±
±
+
—
+
-—
—
+
+
—
—
—
±
—
—
—
-
±
±
—
-
--
+
+
344
2.
M.
I.
ovalis
Bl. Java: Bakhuizen
Cuadra
Merr. Borneo:
I.
Sleum.
cuticle
of
and
veins
of
to sinuous
Stomata
24-26-30
surface
M. iodoides
VAN
STAVEREN AND
5533 ;
Loerzing
Malay
Peninsula;
6.
I. velutina
Tsang 29923;
Hemsl.
3.
King.
vitiginea.
var.
I.
King's
P.
BAAS
philippinensis
coll.
6687;
5.
subvillosa
var.
Thailand:
Kerr
6908.
studied.
Stomata
26-28-31
surface with
walls
11.
Minor
China:
stub
in
curved
with
walls,
in M.
Stomata
32-35-40
gracilis
wide.
walls;
cells
of adaxial
and
abaxial
thin
Unspecialized
cells
of adaxial
and
abaxial
areas
submersed
not
or
of cuticle
in
present
loops
of anticlinal
with
with
rarely
a
very
or
cells
mostly
radiate
minute
with
with
cuticular
second
to abaxial
Unspecialized
sinuous
surrounding
surface.
cells
to sinuous
hairs
or
Adaxial
of both
hair
surface
sides with
walls;
bases
cells
often
rather
a
finely
striations.
stub,
arm
base, possibly articulate; mostly leaving
confined
slightly
concentric
with
cells
subsidiary cells; guard
polar T-pieces; subsidiary
hairs
gracilis.
those
a
with
without
straight
to
a
hairs
slightly
of both surfaces
with
a
papilla-
often
striated cuticle.
cells
Sleum.
Malay
Peninsula:
adaxial
3428.
FRI
27-29-31
long,
pm
of
surface
pm
with
wide.
curved
Indumentum
to
sinuous
present
walls,
of
on
both
abaxial
sides.
surface
Unwith
walls.
M. laxiflora
(Miers.)
Material studied.
Stomata
[xm
long,
cuticular
faint cuticular
samoensis
Howard
Moluccas:
30-33-36
concentric
M.
Unspecialized
prominent.
studied.
with
to
with
1093.
pm
and a thin-walled
for M.
gracilis (King)
sinuous
Cheo
without
being shed, mostly
Material
specialized
surface
(fig. 4)
hardly
of normal
body
except
veins not
Minor
&
laxiflora occasionally
M.
granular cuticle,
a
M.
after
3
ledges,
consisting
bases,
wide.
pm
22-25-27
Seem.
with
stomatal
outer
very thick-walled
hair
abaxial
cuticle
confined
epidermis.
granular cuticle,
like
IV than
hairs,
652.
curved to sinuous
anisocytic,
Indumentum
and
with
granular
curved walls.
Steward
MEDUSANTHERA
distinct
adaxial
cells
finely
prominent.
21-24-26
long,
pm
slightly
of adaxial
Stomata
of
a
Sudan
of normal
consisting
cells
with
with
deeply
more
Indumentum
Unspecialized
Korthals
slightly
to
walls, staining
veins
cells
T-pieces; subsidiary
Sleum.
long,
pm
completely submersed subsidiary cells; guard
Hand.-Mazz.
Material
3.
King.
Sumatra: Buwalda
cells.
surfaces.
Sumatra:
straight
with
guard
both
(Bail!.)
studied.
or
T.
polar
anticlinal
and
walls.
M. hookerianus
without
and
cells
Material
2.
Sumatra:
5709;
vitiginea (Hance)
yatesii.
var.
and
ledges
periclinal
over
straight
1.
I.
W.
with 4 to 6 almost
cyclocytic,
unspecialized
midrib
or
Merr.
yatesii
stomatal
outer
2.
Brink
I. reticulata
C.
MAPPIANTHUS Hand.-Mazz.
10.
Stomata
1.
7.
Ajoeb 164;
Sumatra:
8. I.
4.
Rehd. Indo-China:
seguinii (Levi.)
21351;
v.d.
A 2342 ;
G.
studied.
Stomata
27-34-37
71.
27-30-31
striations.
jxm
wide; subsidiary
Unspecialized
striations.
(Reinecke)
Material
(fig. 4)
Atasrip
Samoa:
pm
Howard
Reinecke
long,
104.
22-26-28
pm
wide.
cells
cells
occasionally
overlying major
with
faint radiate
veins on adaxial
surface
EPIDERMAL LEAF
4.
M. vitiensis
CHARACTERS OF
Stomata
34-35-38
studied.
Stomata
24-26-29
and
stub.
veins
not
and minor
with
of
basal
of hairs
mostly
walls,
Minor
M. caudata
29-33-36
M.
outer
and
reticulata
stomatal
with
ledges
globular hairs;
straight
and veins
6
almost
of normal
consisting
also
4 to 5
and
to
with
completely
ledges
hairs
with
curved
slightly
a
walls;
pitted cuticular flanges.
Adaxial
on
hairs
with
flanges; cells
low
numbers
submersed
polar T-pieces.
layer
of the
with
some
mostly
curved to sinuous
of abaxial
major
on
subsidiary cells;
Indumentum
infrequent, with pits
outer
the
surface
in
present
partly
without
normal
markings
surface
and abaxial
anticlinal cuticular
in
mostly
all
types
hair bases
only.
body;
walls: in M. caudata
surface with
a
finely granular
prominent.
Peninsula:
long,
pm
anticlinal
Bl.
studied.
Celebes:
Stomata
29-32-37
confined
to abaxial
cuticular
surface;
jxm
wide.
1168.
Unspecialized
cells
adaxial
Indumentum
uncinate
of
surface
with
flanges.
26)
Forslen
long,
(xm
Shah & Sidek
22-26-29
18 and
(fig.
Material
s.n.
24-26-28
normal
|xm
hairs
wide.
infrequently
present
:
on
both
and
globular hairs
surfaces.
Meisn.
Material
studied.
Stomata
26-29-31
4.
in
to abaxial surface.
Malay
coarsely granular
India:
sine
long,
pm
nomine
1447.
22-24-28
pm
wide.
Indumentum
very
scanty.
Hair
bases
absent
surface.
M. reticulata
Material
midrib
4 to
faint inner stomatal
King
Stomata
from adaxial
surfaces
surfaces with
surface, cyclocytic,
of adaxial
veins
studied.
M. kleinii
outer and
26)
with very fine cuticular
confined
Material
M. celebica
both
overlying
surface,
uncinate
coarsely granular
cuticle.
18 and
(fig.
conspicuous
Unspecialized cells
with
conspicuous
of both
adaxial side
of adaxial
normal,
bulging
of
with
wide, cyclocytic,
(xm
with
Indumentum
to abaxial
veins
cells
consisting
on
Meisn.
Stomata confined
guard
NGF32614.
22-25-28
long,
prominent.
MIQUEL1A
13.
Sleum.
Unspecialized cells
walls of cells
anticlinal
3.
Kaneh.
Britain:
|j.m
wide.
pm
subsidiary cells; guard cells
arm
Minor
New
polar T-pieces.
with
second
2.
(Hemsl.)
M. megacarpum
Material
1.
30-31-34
long,
pm
MERRILLIODENDRON
submersed
345
ICACINACEAE
Smith 6159.
Fiji;
studied.
1.
MALESIAN
Seem.
Material
12.
THE
Merr.
studied.
Philippines:
Stomata 28-31-34
[o.m
long,
B.S.
43303.
22-24-26
|xm
wide,
a
few
present
on
and
near
the veins
on
adaxial
surface.
14.
I.
NATSIATUM Buch.-Ham.
N.
herpeticum
Ham.
Material
studied.
Stomata
19-21-24
cytic,
with
partly
and without
(fig. 25)
India:
jxm
Griffith
long,
submersed
polar T-pieces.
828.
16-18-20
fim
wide,
intermediate
neighbouringcells; guard
Indumentum
consisting
cells
between
with
anomocytic
and
distinct outer stomatal
of many normal
hairs,
with
or
cycloledges
without
a
346
M.
second
also
stub
arm
and with
of
consisting
both
surfaces.
present
on
abaxial
surface with
anticlinal
15.
intermediate
conspicuous
cuticular
surfaces.
curved
slightly
in
in N.
mostly prominent
N. foetida
Material
normal
many
cuticular
neighbouring
2.
25-27-30
3.
N.
over
Stomata
36-40-45
16.
cells
walls.
pits
in
veins
both
hairs
One
may
curved
also
(see
cells
foetida
and
cuticle
with
of
over
of
of both
surface
cuticle.
granular
with
confined
Minor
prominent in
faint
normal
some
cuticle
less
or
with
major veins
striated
most veins not
more
of abaxial
walls,
with
with
consisting
and
with
to
veins
N. montana.
faint cuticular rims. Indumen-
Unspecialized cells
stub.
anticlinal
Minor
veins
of adaxial
of abaxial
walls;
surface
surface
with
especially
mostly prominent.
walls.
stomatal
complex
normal
also
the
wide.
of
veins
both
not
normal
hairs
with
surfaces
with
finely
distinct
a
granular
prominent.
5
hairs
preparations
hairs;
with
normal
and
verrucose
hairs
present
midrib
to
without
cuticle
over
a
second
periclinal
species
with
3
has
been
present
on the
clustered
of abaxial
of the
cluster,
cells
foetida subsidiary
cuticular
classified
striation.
as
of
cells
intermediate
P.
normal,
species;
hairs
on
and with
present
central
in
hair
5
of
with verrucose
with
species
of abaxial
Because
and
all
body
hairs
surface;
sinuous
walls;
in
T-
polar
surface, globular
markings
species,
minute
bracteata,
consisting
surface in 4
of N.
stronger
P.
cells of adaxial
slightly
neighbouringcells
Nothapodytes
in
hairs
adaxial
cuticular
and veins
subsidiary cells; guard
without
Indumentum
globular
on
or
borneensis,
and thick as the other hairs
long
a
of P.
areolae).
over
submersed
and with
ledges
surface, occasionally
hairs
by
normal
finely granular
very
completely
stomatal
of hairs
confined
cells
Indumentum:
IV)
to almost
body. Unspecialized
in
epidermal
in
pm
table
Uncinate
times as
walls,
Minor
surfaces with
partly
abaxial
the
Indumentum',
cells
mostly prominent.
clustered
wall.
on
53.
wide.
pm
in cuticular
of hairs
call
other
cyclocytic.
sinuous
pittosporoides
arm
distinct outer
on
several
from the
and
N.
Unspecialized
of both
3 to 5
and
basal
types
markings
*
Wall,
only;
cuticular
slightly
granular
walls,
& Li 179.
veins
not visible
clustered
to
cells
Minor
mainly
bulging
species;
hairs
curved
polar T-pieces,
to midrib
wide; guard cells
pm
26-30-31
long,
with
uncinate, globular,
midrib and
sin
faint
Indumentum
with
& Kuswala
due to the abundance
hairs
wall,
of
Sleum.
Fan
faint to rather
pieces (stomata
normal
basal
types
3778.
21-22-25
anticlinal
and
pm
cyclocytic,
macrophylla
with
guard
very
confined
pittosporoides,
periclinal
stub.
arm
Unspecialized
with
foetida.
cell
N.
second
small
over
China;
PHYTOCRENE
Stomata
Edano
25-28-30
long,
pm
periclinal
anticlinal
N.
surface
striated cuticle.
second
a
studied.
and
a
pittosporoides (Oliv.)
stub.
in
walls;
Java: Kostermans
Material
arm
slightly
BAAS
BI.
Stomata
cuticle
with
cells with
studied.
of
with
bulging
both
arm;
surface
cyclocytic;
without
or
stub, mostly
and N.
foetida
long,
pm
Material
indication
pore
montana and
striations
montana
N.
of abaxial
and
with
of adaxial
Philippines:
hairs
surface
in
pits
developed
of adaxial
cells
with
body,
well
P.
AND
Sleum.
(Wight)
studied.
Stomata 36-41-45
tum:
N.
and
arm
sinuous
slightly
to
neighbouring* cells;
1.
second
cells
Unspecialized
walls;
anomocytic
ledges
to the
parallel
a
cells
of the
one
STAVEREN
Bl.
between
without
or
with
VAN
C.
prominent.
stomatal
outer
strations
with
hairs,
sinuous
veins
NOTHAPODYTES
Stomata
hairs
Unspecialized
slightly
Minor
walls.
layer
verrucose outer
a
malpighiaceous
G.
because
of this
straight
surface
they
deviate
deviation
between
with
the
anomocytic
curved to sinuous
walls
of cells
cells
of abaxial
of cells
in
overlying
straight
areolae
surface
midrib
a
to
curved
slightly
not visible
veins
overlying
surface often with
of abaxial
walls
cells
347
THE MALESIAN ICACINACEAE
with
walls, rarely
unspecialized
species
OF
CHARACTERS
LEAF
EPIDERMAL
due to
with
3 above mentioned
abundance
in
mainly straight
these
finely granular cuticle overlying
and veins
in the
walls;
the
of
hairs
(anticlinal
densely hairy species);
anticlinal
pitted cuticular flanges in
4
walls;
species.
anticlinal
Minor
veins
strikingly prominent.
Material
studied
1. P. anomala
Merr. Borneo:
905 ;
Rant
5.
macrophylla.
P.
8. P.
S 22325
1840;
Sleum.
interrupta
Sumatra:
Malaya 1759;
Borneo:
B.S.
2. P. borneensis
Malay Peninsula: Maingay
3. P. bracteata Wall.
palmata
Murthy
Guinea:
New
Teijsmann
s.n.;
Wall.
2715
1. P.
Malay
(
Carr
Becc. Borneo:
=
K.D.
12612;
377);
6.
P.
oblonga Wall. Malay
Peninsula:
King's
coll.
H.B.
Teijsmann
8459;
4. P. hirsuta Bl. Celebes:
macrophylla (Bl.)
Peninsula:
2226;
9. P.
Phyt.
BI.
var.
Surv.
Fed.
Sleum.
racemosa
& Ashton.
PITTOSPOROPSIS Craib
17.
kerrii
1. P.
Craib
Material
studied.
Stomata
20-23-25
Thailand:
[jim
subsidiary cells; guard cells
consisting
of both
adaxial
18.
of
surface
with
PLATEA Bl.
a
with
Quantitative
IV.
cells
19 and
and
with
of
with
wide, cyclocytic,
one
adaxial
cuticle.
4 to 5
cells
(xm
outer stomatal
granular
(fig. 12,
cyclocytic,
peristomal rim; guard
Table
with
Unspecialized
138.
17-20-21
malpighiaceous hairs
sides.
Stomata*
Garrett
long,
ledges
two well
or
and
Minor
and without
developed
abaxial
veins not
about
surface
5
partly
polar T-pieces.
arms,
with
submersed
Indumentum
confined
sinuous
to midrib
walls;
cells
20)
partly
with
subsidiary
submersed
conspicuous
outer
qualitative epidermal
stomatal
cells
and with
a
well
ledges, probably
leaf characters
of
in
developed
all
walls
~
of
®
am
3
T=
5
u
iS
O—
species
species
und lation anticlnal cuticular
-b
u
I
length
V-
e3
TD
<U
cluster d
—
C
•-<
,2
J3
•mal
O
no
3ÖOOC
3
S
CO
.tS
5
2
T3
Ills
uncinate globular
wiIdth
c
S
species
Phytocrene.
hairs
3* 3. 1A
Stomata
O3
p p
sizes
5’
C/5
n (A in
D
of
prominent.
.
3
CJ
§§
H
“
.ll
ed
ah
a Dp
aaaD
P a*
•
flanges
—
1. P.
k
anomala
Ö
S'
I
g
to
23-26-30
L^i to TL*J o
18-20-22
oo to?
to
to
2. P.
borneensis
<>
sw ns 1
?
•o
•o
?
1
U)
3.
bracteata
Öso s
C>
•o
?
•o
?
1
1800
to? toto
15-16-17
LA
T -O
to
T U)? LtJ L*J
26-30-33
fN) -J
24-27-30
6
+
+
•O
?
+
+
+
+
+
+
-
+
-
+
+
+
+
1
+
+
+
+
1
+
+
+
+
+
+
-
TÍ^)
P.
>
H
4. P.
k
LA
5.
hirsuta
5S52S
s3
P. S"
to
interrupta
T3^
6.
as P.
t
O
"■4
1. P.
k
l3
00
8. k
P.
1 6*
macrophylla
S' I
oblonga
r
O
vo
9. P.
fc
*
racemosa
2£
o Co&
i
Length
the
17-19-21
«O
VO
•—
to
21- lO fto -O
19-20-21
VO to
*-*
1
K)
22K>
toLA 1 to 00
17-20-22
•O (O
to
1
and width of stomata
guard
cells
are
(
obscured
1
! toUJ
19VO tO
S'
palmata
-
l
•O
?
2ÖQ
+
+
by
were
a
only
1
??
measured
peristomal
rim.
1
-
in P.
+
1
+
1
+
1
+
+
+
+
+
•o
+-?
+
+
+
+
++
+
+
+
+
+
+
+
+
+
+
+
1
-
1
+
1
1
•o
?
1
1
±
H-
+
+
•O
+-?
+
+
1
sclerophylla. In
the
c3
pit
a?n
1
-
1
remaining species
348
M.
with faint
in
P.
in
bullata
cuticle
material.
the
minor
with
of
Unspecialized cells
with
mostly
with
conspicuous
bullata Sleum.
Material
studied.
Indumentum
both
2.
sides
with
P. excelsa
Material
Bl.
Java:
consisting
surface
with
a
abaxial
cells
with
P. latifolia
4.
P.
with a
glandular hairs.
scandens
of
almost
New
and clustered
hairs.
Unspecialized cells
of
glandular
hairs.
overlying
veins
cells
of adaxial
cuticular
striations;
Unspecialized
with
faint
striations.
China:
nomine ;
F.C. How
Sumatra: Kostermans
73310 ;
Peninsula;
Malay
Sinclair
267.
and clustered
striated
cuticle;
hairs.
abaxial
Unspecialized cells
cells
with
of
conspicuous
of the
types
body
celts
slightly
Guinea:
Docters
17-18-20
pits
confined
curved
in
stomatal
Unspecialized
without
ledges
with
cells
surface
finely granular
a
cuticular
of
peltate-stellate and
striations.
10613.
Minor
variable
a
with
besides also
Unspecialized
number
outer stomatal
of normal hairs with
surface.
veins
with
cells
a
coarsely
consisting
cells
of
partly
ledges
verrucose
of
and
outer
globular hairs.
of adaxial
and
abaxial
prominent.
Moeser
and
with
of adaxial and
cuticle.
Leeuwen
wide, cyclocytic,
consisting
abaxial
walls.
thick-walled
with
van
[xm
bulging basal walls,
to
predominantly paracytic,
of abaxial
Indumentum consisting
sides
subsidiary cells; guard
Indumentum
PSEUDOBOTRYS
hairs
wide.
pm
of both
Becc.
and with
of hairs
with
29-33-39
submersed
polar T-pieces.
Both
Chew 8026.
long,
(itn
completely
layer
normal
cells
prominent.
Becc.
19-21-23
outer
and
cells
peltate-stellate, glandular
Unspecialized
Stomata
Stomata
cuticular
sine
long,
pm
studied.
20.
walls;
abaxial
cuticle,
166.
21597 ;
Borneo:
Material
surface
curved
slightly
on
abaxial
to
striations.
coarsely granularand/or
POLYPORANDRA
without
also
usually
confined
thick
a
granular
Sleum.
sclerophylla
28-32-37
P.
with
with
striations.
Stomata
or
Sulit
consisting
studied.
1.
cells.
20)
peltate-stellate
Borneo:
Material
19
to
striated
a
hairs
and
present,
straight
with
veins
surface
if
guard
(fig. 12)
Dakkus
of
hairs,
BAAS
glandular hairs;
filled
cuticle,
abaxial
on
surface with
Minor
Glandular
and
the
overlying
unicellular
P.
Bl.
surface
cuticular
striations.
cuticular
conspicuous
40618; Philippines:
adaxial
abaxial
coarsely granular cuticle,
studied.
Indumentum
present
hairs.
wall
AND
STAVEREN
19638 PS. Ashton.
excelsa
var.
studied.
Material
S
the
peltate-stellate, glandular
conspicuous
Indumentum
3.
19 and
of
and
clustered
coarsely granular, rarely
a
rim
peristomal
clustered
surface;
of adaxial
Borneo:
consisting
hairs
glandular
cuticular
(fig.
of
between
space
adaxial
without
or
due to the
VAN
C.
peltate-stellate and sessile,
consisting
small
a
and
veins
cells
P.
also
latifolia
top
adaxial
1.
of
species consisting
Peltate-stellate
and
surface.
all
and P.
and at
major
often not visible
polar T-pieces,
Indumentum
G.
a
striated
Minor
sometimes
well
body
and
abaxial
and
also
developed
a
anomocytic; guard cells
polar T-pieces.
thin-walled
surface
with
base,
straight
finely granular cuticle,
veins not
prominent.
confined
to
with
Indumentum
to
slightly
those
conspicuous
consisting
abaxial
curved
of adaxial
of
surface.
walls;
surface
cells
with
a
EPIDERMAL LEAF CHARACTERS OF
1.
P. cauliflora
studied.
Stomata
24-27-29
2.
P.
dorae
New
studied.
Stomata
27-29-31
1.
Guinea:
New
Guinea:
studied.
Stomata
mainly
surface,
confined
markings
and
on
of
slightly
over
hair
or
areas
hairs,
bases.
cuticle
curved
in
to
periclinal
pm
cells
long,
with
confined
the broad
of uncinate
hairs
from
paracytic
to
anomocytic.
12760.
|xm
exclusively paracytic.
17359.
surface,
conspicuous
and with
body
the latter
and
sinuous
a
in
pits
cells
outer
bulging
on
present
and
basal
surface
on
coarsely granular
strongly
over
anticlinal
of
T-
cuticular
of
globular
surface without
sinuous
small.
polar
with
consisting
Adaxial
both surfaces with very
cuticle
hairs
besides
sinuses
veins
submersed
partly
and without
normal
veins.
and minor
3 to 6
ledges
wall,
with
major
with
of
major
amplitude of
cells
on
stomatal
midrib and
of adaxial
walls;
also
wide, cyclocytic,
surface, consisting
especially
loops; wave-length
slightly
few
a
13-15-16 pm
abaxial
to
Unspecialized
walls
almost
wide,
17)
Elmer
to abaxial
17-19-21
Indumentum
Carr
Wight (fig.
Philippines:
subsidiary cells; guard
pieces.
9307.
wide, ranging
Merr.
repanda (Merr.)
Material
adaxial
Leeuwen
van
ixm
26-28-30
long,
(xm
PYRENACANTHA
P.
Doctors
21-23-26
long,
(xm
Moeser
Material
21.
Sleum.
(Pulle)
Material
349
THE MALESIAN ICACINACEAE
walls
Abaxial
with thin
cells
with
finely granular cuticle
walls.
Minor veins
prom-
inent.
22.
RHYTICARYUM
Stomata
cyclocytic,
cells; guard
Table
with
Quantitative
2 and
in
partly,
also
see
table
V)
R. oxycarpum almost
outer stomatal
polar T-pieces
species only
and
5,
conspicuous
small
Indumentum: in 6
V.
(fig.
with 4 to 6
cells with rather
granular material,
cuticle.
Becc.
some
in
hair
6
species
and
bases noted
qualitative epidermal
completely
ledges,
in
R.
mostly
on
adaxial
leaf characters of
submersed
subsidiary
longifolium with coarsely
with
or
a
finely granular
abaxial
in R.
side;
Rhyticaryum.
cuticle
CL n
c
stomata
stomata
sizes
in
in
v.
of
®
(xm
[xm
walls
-
u
s
species
P ecies
T-pieces
3
lJength
a
1. R.
2.
R.
R.
3.
3. R.
4.
6. R.
*
a
polaar
—
und lation anticlnal
adaxial
g- 2.S.
abaxial
gL'
g 2.
p
’c'5
5
§
i
ab
ab
T3
S
2
«
3
=
§
a
S
=
elegans
27-29-32
24-26-28
24-26-28
+
+
+
22-24-28
22-24-28
21-22-24
21-22-24
+
+
+
+
±±±
±
—
—
longifolium
longifolium
37-40-45
31-35-39
+
+
+
+
±
±
±
26-28-29
26-28-29
23-25-28
23-25-28
+
+
—
26-29-31
23-27-28
23-27-28
+
—
—
+
±
—
-
novoguineense
novoguineense
oleraceum
—
+
+
+
+
—
-
—
—
-
±
—
—
—
22-24-26
+
25-30-31
25-27-28
25-27-28
+
±
±
+
8. R.
8.
R.
25-26-28
22-24-26
±
+
+
+
racemosum
racetnosum
+
+
+
±
±
26-29-32
—
-
+
7. R. oxycarpum
7.
oxycarpum
--
cuticular
13
granular striated granular striated
c
ad
25
3
11
fasciculatum
R. macrocarpum
macrocarpum
5. R.
5.
width
~
o
--
—
--
—
“>
■««
ad +ab
ad+ab
--
±±±
±
±
±
pit ed flanges
|
—
-
±±
—
ad
ad
ad+ab
—
ad + ab
ad+ab
—
ab
ad
350
M.
oleraceum and
R.
Unspecialized
cells
slightly
curved
surface
in
striated
Material
1.
R.
6
carpum
23.
Schellenb.
1085 ;
Becc.
3.
R.
New
with
sinuous
anticlinal
hairs
in
walls,
walls
of cells
Adaxial
flanges.
veins not
mainly
Guinea;
K.
BW
and
with
the
on
AND
second
a
P.
BAAS
stub.
arm
with
remaining species
adaxial
abaxial
and
cells
/or
with
abaxial
granular,
prominent.
Becc.
s.n.;
W. &
8.
with
stubs noted.
with
straight
&
5.
2.
13351;
Laut.
R.
R.
New
fasciculatum
Guinea:
7.
503;
Becc. New
racemosum
Becc.
Clemens
R. oxycarpum
Guinea:
New
Van
4.
1028;
Sleum.
novoguineense (Warb.)
Moluccas: Kornassi
R.
New
K.
Royen
Guinea:
R.
macro-
Guinea:
Sch.
& Laut.
3129.
A.
surface, cyclocytic,
conspicuous
Indumentum:
Unspecialized
slightly
BW
Sch.
11366 ;
confined to abaxial
cells
to
Guinea:
New
Ledermann
arm
curved
stomatal
outer
hair
only
cells
bases
of adaxial
cells
walls;
on
with 4 to 5
ledges
and
both sides
partly
with
few normal
a
surface
and
with
with
submersed
faint to
hairs with
distinct,
minute
second
of abaxial
straight walls,
distinctly
subsidiary
rather
surface
striated cuticle.
Minor
prominent.
S. kleinii
Material
W.
& A.
studied.
Stomata confined
S.
of normal
consisting
sides
walls;
Minor
longifolium
polar T-pieces.
veins
both
cuticular
pitted
6. R. oleraceum
cells; guard
2.
with
on
sinuous
cuticle.
SARCOSTIGMA
small
species
slightly
to
species
Guinea:
Stomata
1.
6
indumentum
STAVEREN
studied
NGF8386;
New
in
or smooth
elegans
Tuyama
racemosum
VAN
G. C.
paniculata
Java:
Zollinger
to abaxial
studied.
Stomata
mainly confined
Table
Borneo:
24-27-28
VI.
23-25-28
pm
long,
21-23-25
pm
wide.
Pierre
Material
surface,
228.
surface,
p,m
SAN
long,
Quantitative
39985.
to abaxial
and
surface,
24-25-26
a
few
also present
and
on
near
midrib
leaf characters
qualitative epidermal
of
stomata
O1 P
sizes
W5
tsi
in
5'
Stemonurus.
S
Cd
S
Cd
predominatly
predominatly
,S
urn
3
.9
Ë
_
E
j-
GO
Ä
c
•?
peristomal distinct
$■
D.T3
2q
.5
op
-5
•-
6
u.
D.
5
width
’S
fc*
cd
length
o
’S
—,
J3
9
o
rim
‘C
0' C/Ï
U>
species
f
-1. S. ammui
O 55J5
.a
O
stomat anisocytic
tá
5“
c
t
o
£
o
to
S
Ui
30-32-35
VUi
Ui Ui
Ki
29-31-33
NO
Ui Ui
+
+
1
2. S. ri
celebicus
O- r>’ s:05
Ki
34fUi l Ki
Ui
Ki
27-29-34
sNO Ui
+
+
+
Ui
3.
Ui
35UiTUi £Ki
Ui
33-34-35
Ui
fUi Ui
1
H±
3
a
&9
grandifolius
-N
4. S. oq
r>
c -J
40-45-47
[>
o
40-45-50
Ui
Ui
Ui
5.
Ui
32- Ui
S. OQ
CoI
gitingensis
of adaxial
wide.
p.m
1
1
1
■
+
-
D.
o
stomat cy locytic
rtu
o
c
C
7-
o
y
too
+
+
1
H±
+
f
1
S. malaccensis
3 O o"
5ö a
Ui "J
Ki
29-31-33
NO
TUi Ui
+
H±
6. S. 5
On
monticolus
oa ri
- cS 1*
Ui
35-37-39
Ui Ui
Ui NO
Ui
33-36-38
Ui
fUi 00
+
+
7. S.
& 5g
O
perobtusus
Ui
33Ui Ui V'Ui
Ki
25-27-30
Ui
Ui O
+
+
+
+
1
00
8. S.
C«ïÖ o■g o' 1
scorpioides
Ni
25-28-30
Ui Ki 00 Ui O
Ki
26-29-30
NO Ui O
1
±
H-
±
if
NO
9.
S.
%r> s:
i
g.
secundiflorus
K>
28-31-32
00 Ui
Ui K)
Ni
26-27-29
-0 Ki NO
+
1
10.
o
S.
umbellatus
o'1
5£■
Ui
33-36-37
Ui Ui TU) --J
Ki
27-30-32
? u> Ni
1
1
1
1
1
1
1
1
1
TT
1
+
+
1
+
H±
1
1
STEMONURUS
24.
Stomata
anisocytic
rim
peristomal
thick
of
dumentum
or
cells
a
in
of
finely
both
and
grandifolius
S.
the
S.
with
abaxial
submersed
rather
straight
cells
slightly
to
with
curved to
with
pitted
rounded
of
cuticular
adaxial
on
curved
cuticle
distinct
flanges.
cells.
S. malaccensis and
on
abaxial
in
S.
S.
with
in-
celebicus,
often with
Anticlinal
S.
of both
cells
grandifolius and
not
and
umbellatus
abaxial surface.
faint
walls
in
S. umbel-
cells
(see fig. 6)
prominent.
Corkwarts
unspecialized
side in
S.
walls;
moreover
veins
Minor
a distinct
remaining species
striations.
asterosclereids between
epidermal
and
walls;
In S.
with
ledges
and/or
sinuous
slightly
abaxial
with
secundiflorus
var.
in the
surface,
noted
bases,
stomatal
secundiflorus
var.
abaxial
to
hair
subsidiary cells,
distinct outer
secundiflorus
confined
penetration
of
abundant on adaxial side in
in the
with
through
sides
scorpioides
impression
cells
VI)
partly
coarsely granular cuticle;
to
on abaxial side with
giving
table
3 to 4
in
hairs
evident
secundiflorus
S.
with
Indumentum
only
also
see
species; guard
and S. umbellatus
surfaces with
striations,
{fig. 6,
cyclocytic,
malpighiaceous
Unspecialized
latus
7
absent
perobtusus,
S.
in
Bl.
to
polar T-pieces.
consisting
351
CHARACTERS OF THE MALESIAN ICACINACEAE
LEAF
EPIDERMAL
umbellatus, rare
to absent
remaining species.
Material studied
(Kaneh.) Sleum.
1. S. ammui
Cel
3.
II/339 ;
SAN
Borneo:
Indo-China:
Pierre
secundiflorus.
W.
grandiflora
Material
S.
Sleum.
2838;
Sleum.
malaccensis
Philippines:
(Mast.)
Guinea:
New
8. S.
Sumatra:
studied.
Becc.
scorpioides
Thorenaar
T.
bb.
33;
Elmer
Sleum.
25059;
Borneo:
10. S.
12277
4.
;
Thailand:
7.
S.
grandifolius Becc.
292;
perobtusus (Gagnep.)
9. S.
SAN 24315 ;
umbellatus
S.
Swanakoses
Becc.
6.
S.
Sleum.
secundiflorus Bl.
Borneo:
S 8641.
Sleum.
Solomon
Stomata 40-41-45
submersed
without
a
5.
16782;
NGF2027 ; 2. S. celebicus Valet, in Koord. Celebes:
Guinea:
WHITMOREA Sleum.
25.
1.
New
gitingensis (Elm.)
(Schellenb.)
monticolus
var.
S.
walls;
Schodde
33-35-37
subsidiary cells; guard
cells
cells
base
abaxial
on
of adaxial
surface
with
a
3629.
wide, anisocytic
surface
Adaxia!
straight
cyclocytic,
to
with
3
to 4
distinct outer and inner stomatal
of normal
consisting
surface.
of both surfaces with
(& Craven)
(xm
cells with rather
Indumentum
polar T-pieces.
thin-walled
cialized
Isl.:
long,
[xm
walls,
hairs
without
of abaxial
finely granular cuticle
with
over
a
and
body
and
thick-walled
hairs
or
surface
periclinal
hair
with
walls.
hardly
ledges
bases.
Unspe-
slightly
curved
Minor veins
not
prominent.
26.
PENNANTIA
Stomata
polar T-pieces.
uniseriate
1.
P.
with
Baylis
surface.
cells
cuticle.
hairs;
in P.
types mainly
of adaxial
and
Minor veins
and with rather distinct
ledges
corymbosa
confined to
abaxial
surface
also
major
with
of
multicellular,
and minor
curved
to
veins
sinuous
prominent.
Forst.
studied.
25-26-28
multicellular,
P.
apex; both hair
Unspecialized
outer stomatal
of normal
Three
Kings
Isl.;
Baylis
22875
(=
Plectomirtha
wide.
Indumentum
baylisiana);
New
Zealand:
s.n.
Stomata
2.
conspicuous
consisting
coarsely granular
corymbosa
Material
cells with
Indumentum
hairs with rounded
of abaxial
walls,
Forst.
paracytic; guard
long,
(xm
uniseriate
cunninghamii
Material
studied.
Stomata
21-23-25
16-19-20
|xm
consisting
of normal
hairs
and
hairs.
Miers.
New
|xm
South
long,
Wales:
Nat.
16-18-19 jxm
Herb.
wide.
11151.
Indumentum
only consisting
of normal
hairs.
352
5.
M. O. C.
SYNOPTICAL
KEY
TO
THE
foregoing descriptive part
From the
has become apparent. This
epidermal
characters and
characters
only.
discussed the
the
fact that
Dr.
out
group of
the key.
In the
In
the
able
were
to
two
key
1.
Apodytes,
13.
19.
Hartleya,
14.
25.
Stemonurus,
1.
Cantleya
8.
Miquelia,
Stomatal
a.
20.
b.
3.
a.
b.
4.
one
is
certain of
leaf
aware
down
unable
were
to a
of the
only.
of herbarium material
we
cases
possibilities
to
to
to
try
the genus.
distinguish
structure
10.
4.
a
genus
in
21.
genus shows
A genus number between
intermediate stomatal type.
an
5.
Codiocarpus,
Mappianthus,
Nothapodytes,
that the
means
question.
16.
11.
Gastrolepis,
Medusanthera,
Phytocrene,
22.
Pyrenacantha,
17.
6.
12.
Gomphandra,
18
Pittosporopsis,
23.
Rhyticaryum,
7.
Merrilliodendron,
Platea,
Sarcostigma,
24.
Pennantia.
(4)
5
6
(7)
11
8
9
20
10
13
12
(14) (15)
16
17
18
19
21
22
23
10
11
13
14 15
16
17
18
21
22
25
(24) (25)
(24) (25)
cells:
inner
without
stomatal
inner
3
ledges
stomatal
ledges
12 25
1
2 4 5
6 7 8
9
19 20
21
22
23
26
cells:
polar T-pieces
without
polar T-pieces
2 4 5
6
7
6 7
9
10
1
3
8
12
11
15
16 20
13
14
22
23
15 16
24 26
17
18 19
cells:
distinct
a.
with
b.
without
a
24
Subsidiary
b.
26
6
Guard
a.
20
helicocytic
with
15.
(7) (14) (15)
2
23
5.
1
3
(4)
Guard
Leenhouts
types:
cyclocytic
24
down the
satisfactorily
Gomphandra
Citronella,
Iodes,
Pseudobotrys,
anisocytic
with
9.
Whitmorea, 26.
paracytic
Guard
3.
,
Natsiatum,
anomocytic
2.
when
key
on one
fragments
that the genus exhibits
2.
Polyporandra,
by
these
using
Icacinaceae. In several
be cautious and be
to
based
underlined number for
an
implies
Gonocaryum,
of them
genera.
several alternative types of the
brackets
structure
characterize genera
to
for this purpose.
the
use
8 of these
identify
ICACINACEAE
most
narrowing
are
10 leaf
BAAS
key.
family
has
one
descriptions
selected
to
the
in
helpful
of Medusanthera and
case
between the
to
species, though
kindly
We
belonging
P.
considerable range of variation
a
synoptical key
of this type of
will be
all the
virtually
Sleumer
possible
identify
to
only possible
course
with leaf material
or a
makes it
diversity
chose the
we
specific descriptions
species
MALESIAN
AND
STAVEREN
great diversity in leaf epidermal
a
consequently
advantages
At present it is of
dealing
OF
Since many genera also show
amongst their species,
(1966)
GENERA
VAN
not to
partly
a
peristomal
distinct
25
rim
peristomal
4
rim
1
18 24
2 3
5
6 7
4 5
6
2 3
9
8 9
10
11 12
13
14
15
16
17
19 20
21
22
26
cells:
hardly
submersed
to almost
completely
submersed
10
12
13
14
15
16
17
18
19
21
22
23
24
LEAF
EPIDERMAL
6.
a.
10-20
b.
20-30
c.
30-40 (xm
23678
11
40-50
5 7
25
Average
a.
b.
c.
d.
8.
[xm
14
or
14
16
4 6
7
17
9
sides
adaxial
on
22
23 24
18
19 20
22
23
24
26
19 21
10
11
26
12
13
15
16
18 20
22
23
24
25
7
8
24
abaxial
and/or
side
1
2345679
10
11
12
13
14
15
16
17
18
19
25
consisting of:
b.
malpighiaceous
hairs
hairs
1
2 3
4 6 7 9
3
5
14
6
globular
hairs
913
uncinate
hairs
9
e.
clustered
hairs
16
f.
peltate-stellate
g.
glandular
h.
papilla-like
i.
multicellular,
hairs
16
10
11
12
13
14 15
16 19
20 21
22 23
23
26
25
26
17 24
19
21
131621
1
18
18
hairs
18
hairs
7
hairs
uniseriate
26
hairs:
with
a
Normal
with
b.
without
a
with
second
a
a
13
1 2
part
stub
arm
14
3
16
19
4 6 7
21
10
11
12
346H12141522
stub
body
thick-walled
22 23
15 20
22
25
1
2 4
6 7 9
10
11
23
13
14 15
16
19 20
21
22
25
26
and
body
a
thin-walled
and
a
base
thin-walled
2 6
base
11
20
25
1 3 4 6 7
9
10
12
13
14 15
16
19
26
hairs:
a.
with
cuticular
b.
without
Anticlinal
a.
with
b.
without
24
markings
cuticular
cell
pitted
23
arm
second
thick-walled
without
Normal
9
part
basal
bulging
hairs:
a
21
a
hairs:
a.
Normal
basal
bulging
without
Minor
17
hair bases:
normal
Normal
16
15
1315182224
24
(xm
a.
b.
15.
12 13
40-50
Indumentum
a.
14.
10
257811182224
present
b.
13.
9
b.
a.
12.
9
15 24
1 3
absent on both
d.
11.
6
(xm
a.
c.
10.
4 6
width of stomata:
20 21
9.
1
|im
30-40 (xm
Hairs
353
ICACINACEAE
9 21
jxm
10-20 [xm
20-30
MALESIAN
of stomata:
Average length
d.
7.
OF THE
CHARACTERS
the outer
1
of their
layer
2 3 4 6
7
10
11
12
body
15
20
flanges
cuticular
at least
flanges
1
on
midrib
2 3 4
5 6 7 8
and veins
9
10
11
24
25.
26
veins:
a.
prominent
b.
not
191314161921
22
23 25
26
walls:
cuticular
pitted
25
on
markings
prominent
7
1
9
10
2 3
13
4 5
14
15
6 7
8
16
11
18
12
19
21 23
15 17
20
26
22
179
13
14 15
12
16
162224
17
18 19
20
21 22
354
6.
M.
O. C. VAN STAVEREN
AND
P.
BAAS
DISCUSSIONS
6.1.
The
value
of the
characters
The characters used in the
taxonomic and
same
type of granulation
Apodytes
e.g.,
walls varied from
exhibit
however,
and/or
specimens
rather
constant
of which
cuticle. In other genera such
are not
all of the
variable,
very
such
as
anticlinal walls and also the
(see
343 &
336,
pp.
348). In,
examined,
the anticlinal
(B.S. 24941).
Some genera,
were
sinuous
to
be
may
striation of the cuticle
12
straight (Rudatis 786)
a
synoptical key
Some
undulation of the
flanges,
dimidiata,
used
and the
value.
diagnostic
of the cuticular
pitting
descriptions
kind of striation
in their
and/or granulation
characters may be restricted
part of the species
to
only.
The
flaps
This
genus.
hair
types
used when
easily
easily
by
stomatal
to
some
stomatal
synoptical key).
species
of
a
genus may
normal hairs with
(e.g.
be
to
and
types
with
ledges
found
usually
their
isolated
diagnostic
Platea and
adhering
constant
hair
for
a
though
types,
The absence of certain
be due
to
thick-walled
a
value of hair
the fact that
body
and
a
should
only
clearly
demon-
key
indumentum
positions
and
only,
within
the
of
material
more
on
the
These
taxa
Gonocaryum
because
family
be
can
and
of the
be
Platea
slender
indumentumof glandular,
complex
and clustered hairs in the latter. In
is
types
corymbosa.
Pennantia
hairs in the former genus and the
detailed study
large
species
genera like
level
might
Gomphandra
prove useful
I).
Taxonomic
Engler
applies
taxonomic and
rather
table
6.2.
also
Gonocaryum,
peltate-stellate
(cf.
peristomal
inner
was
positive.
papilla-like
a more
rim
Therefore, character 9 of the synoptical
recognized
occupy
of distinct
leaves of
shed
base).
The great
in
a
underlined figures in
(see
mature
are
thin-walled
of
mostly
in
these hairs
strated
and
exist
exceptions
absence
or
presence
cuticular
(1896)
implications
and
Sleumer
and
(1942
personal
information)
distinguished
four tribes within the Icacinaceae:
Icacineae
Citronella I,
(Apodytes I, Cantleya II,
Gomphandra II, Gonocaryum II, Hartleya II,
III,
III,
Nothapodyles
Rhyticaryum III,
lodeae
Pennantia I,
Codiocarpus II, Gastrolepis II,
MedusantheraII,
Stenonurus II and
Whitmorea
(lodes III, Mappianthus III,
Merrilliodendron
Pseudobotrys
Platea I,
I,
Pittosporopsis
I,
II).
Natsiatum III and
Polyporandra
III).
Sarcostigmateae (Sarcostigma III).
Phytocreneae
In
his
refrained
( Miquelia III, Phytocrene
recent
more
from
subdividing
correlation of the
The
roman
represent the
revision
of
the
the
family
macromorphological
figures
three
Bailey & Howard
following
levels
(1941),
of
each
III
and
Malesian
into
Pyrenacantha III).
Icacinaceae,
tribes
because
Sleumer
of
(1971)
unsatisfactory
characters with anatomical data.
genus
specialization
name
in
in
the above
Icacinaceae
who based themselves
on
as
enumeration
recognized
by
wood anatomy and nodal
EPIDERMAL LEAF
OF THE
CHARACTERS
is
I
anatomy.
Group
trilacunar
nodes;
characterized
II
Groups
and trilacunar
perforations
355
MALESIAN ICACINACEAE
by
scalariform
by
mixture
a
simple
and group III has
nodes,
vessel
perforations
of scalariform and
vessel
and
vessel
simple
perforations
and unilacunar nodes.
Callen-Lobreau
in
(1973,
for the
specialization
family
as
the
three levels coincide with those of
The present results
leaf
on
with the subdivisions of the
The lodeae and
hairs
have
usually
partly
almost
to
stomatal
a
bulging
inent. The leaf
phenomenon.
close
well
as
are
and
common,
prom-
isolated
not an
Phytocreneae
a
stomata
always
are
tribes is
in
points
many
quite
The
anomocytic.
two
with
stomata
Natsiatum has
Only
very unusual. The veins
have
be
to
appear
cyclocytic
cells.
and
epidermal
these tribes. The normal
of the lodeae and the
representatives
their wood anatomical features
morphologically they
subsidiary
cyclocytic
are
comparison
in leaf
strongly
to
resemblance between these
epidermal
All
restricted
are
submersed
usually small; polar T-pieces
are
suitable for
are
basal part. All genera show
completely
of
outlined above.
as
intermediate between
type
features
resemble each other
Phytocreneae
levels
characters. These
and Howard for wood and node.
Bailey
features. Globular and uncinate hairs
three
distinguishes
pollen morphological
on
epidermal
family
also
press)
based
lianas,
pollen-
(Callen-Lobreau, personal
communication).
Mappianthus, belonging
in
1969,
name
reduced
was
here because there is
features.
strikingly
bulging
Particularly
different in
18-23
the
x
16-20
two
prominence
taxa:
pollen
indumentum
in
,
a
and
bulging
Mappianthus
is
Sarcostigma (the only
genus of
are
Leaf
The
T-pieces
as
a
xylem, node,
epidermal
and
a
averages
and
kindly
striking
the lodeae and the
Mappianthus
informed
us
to
that the
are
for
is of
climbing
anatomically, however,
a
second
(except
for
Mappianthus),
present
therefore in
in
a
arm
not
Sarcostigma
stub. This is very
Natsiatum),
whereas in
Sarcostigma,
way
habit
there is
where the
Sarcostigma
absent from the
view of
supports Engler’s
rather isolated genus.
remaining Icacinaceae,
geneous in leaf
lodes
and the other lianas. In
Phytocreneae
Epidermal anatomy
regarding Sarcostigma
are
Mappianthus only
polar T-pieces
Engler’s Sarcostigmateae)
markings (except
smooth. Polar
other lianas.
of
also
press)
Sarcostigma
unusual for the lodeae and the
they
whilst
Stomatal size in
indumentum consists of normal hairs with
hairs show cuticular
dimensions
different from that of Iodes.
much resemblance between
the
generic
epidermal
24-25 [xm. The characters shared
x
absence
the
Sleumer
normal hairs with
support the reduction of
not
(in
in leaf
stomatal
globular hairs,
26-28
like the lodeae and the Phytocreneae.
so
the
affinity
ubiquitous throughout
are
therefore do
Mappianthus
and
basal part.
cyclocytic stomata,
Madame Callen-Lobreau
of
and
close
a
by
have retained the
lodes possesses
“genera”.
hairs,
from lodes
kept apart
1971. We
indication of
of the minor veins
Phytocreneae
lodes.
[im
lodes in
to
two
uncinate
basal part,
the lodeae and
no
the
the
has normal hairs without
by
to
him
by
all
belonging
characters. This
pollen features,
to
the
Icacineae,
apparently
because the
also
are
rather hetero-
applies
genera may be
on
to
secondary
all levels
of
356
M.
in
specialization
leaf
Using
of
pollen
Howard’s
characters
it is
only
and
is,
VAN
P.
BAAS
classification.
clear pattern
a
obvious correlation
no
We therefore restrict ourselves
genera in which the leafanatomical evidence
some
AND
STAVEREN
recognize
to
moreover,
wood anatomical characters.
or
the discussion of
C.
Callen-Lobreau’s
impossible
within the Icacineae. There
relationship
with
and
Bailey
epidermal
O.
to
seems more
straightforward.
Most
thera
species
of
Gomphandra
and
(see descriptions
between
two
the
anisocytic
As
both
Gonocaryum
for
the
other genera,
the stomatal type
no
absolute
rather
specialization
character
characters this
of the
mediate types
we
correlation of
see
and
331,
p.
is
genera
intermediate type of
however,
there is
the
specialized
a
in genera
anomocytic
Icacinaceae
(1941)
the
For
node and
level II may
In
have
genera and
of the
the intermediate
together
to
the
the
with
Callen-Lobreau
the lack
and
the
that
anisocytic
an
inter-
or
cyclocytic
or
an
absolute correlation
two
be restricted
paracytic
group I. It thus becomes
stomatal
and
anisocytic
and
restricted
wholly
anomocytic
to
intermediate types
level II. The
are
inter-
and intermediate
stomata to
and
intermediate,
cyclocytic
of
of absolute
with stomatal type.
pollen
of
specialization
their
primitive
most
paracytic
condition and
and
anisocytic, cyclocytic
of this lack
spite
pollen
here.
interpretation
the genera of level III have the
stomata.
at
speculation
shows
diagram
wood,
and
indeed,
node and
wood,
the genera and the level of
over
Howard
The
Without due
fig. 27).
very hazardous
of the cyclocytic
strong tendency
types,
belonging
primitive
346.
within the
positions
paracytic, anomocytic, cyclocytic
mediate stomatal types, and
the
with
ones
specialization
summarized.
in
also
epidermal,
level of
specialization level
stomatal types. The genera in
genera
relationship
only
about their mutual affinities
(see
refrain from further
Bailey &
to
Level I houses
with
genera
to occur
the
354).
p.
would be
with other
type and
according
press)
most
close
to a
are
isolated
might speculate
guiding
established,
The stomatal
the
they
(see
In fig. 27 the distribution of the stomatal types
(in
points
because
so
occupy
one
a
as
correlation
characters could be
6.3.
which
because of their unusual indumentum
consideration of other
since
more
even
from those of Medusan-
distinguished
stomata.
Platea and
family
using
genera,
be
cannot
synoptical key),
types
to
that in
likely
represent
stomata
are
a
more
derived.
For other
paracytic
assume
series.
plant
groups
stomata to
that
the
Guyot
be
several authors have also claimed the
primitive. Rohweder,
anomocytic
(1966)
and
type is
primitive
for the Umbelliferae and
however,
assume
angiosperms,
(Baranova
paracytic
that
the
1972).
For
are
starting point
(1971)
the
more
for
regard
many
occurs
stoma
in
a
Icacinaceae
primitive
is
basic
to
the
we
than
also
advance
anomocytic
stomata
Other
primitive
the
ones.
as
authors,
other types
number of so-called
or
(1971)
specialization
anomocytic
Saxifragaceae, respectively.
paracytic
because this type
stomata
a
Gorenflot
anomocytic
Schlumpf & Krattinger
in
the
families
hypothesis
that
It is obvious that
EPIDERMAL LEAF CHARACTERS OF
Fig.
27.
Stomatal
Howard. See also
species
with
types
and level
of
the text. The circle
helicocytic
357
THE MALESIAN ICACINACEAE
specialization
with
stomata. Generic
genera
names
in
wood
and node
having anisocytic
abbreviated.
according
stomata
to
Bailey
also includes
and
the
358
M. G.
studies should be undertaken
ontogenetic
trends of
7.
Epidermal
for
the
in
types
this part of the
for
surprising
and
in
diversity
the
Malesian Icacinaceae
most structures
but
affinities,
the
the
considered renders it
other
or
also show
histology
detailed study
equally
Icacinaceae from Africa
America
range of variation within the
huge
remaining
gain
a
of intersubdivide
to
This is in fact
taxa.
of vessel
not
perforation
less continuous variation. We
or
should be
to
of the ideas advanced here and
some
the
and
of the
employed
characters
occurrence
impossible
suprageneric
a more
profitably
epidermal
therefore support Sleumer’s view that any subdivision of the
artificial. An
of the
understanding
be
can
instances
some
where other characters like type
family
a
wood
general
into tribes
family
In
indications of
straightforward
mediate
better
a
BAAS
P.
CONCLUSIONS
identification of genera.
provide
get
AND
in the stomatal apparatus of the Icacinaceae.
specialization
GENERAL
to
C. VAN STAVEREN
would be
family
genera and
undertaken in order
more
of the
species
comprehensive
to
test
of
picture
family.
ACKNOWLEDGEMENTS
The
authors
indebted
greatly
are
correlation
of anatomical and
tion
disposal. Special
at
our
discussions
helpful
technical
assistance.
Mr. C. L.
Marks,
the illustrations.
read
on
and
thanks
Mr.
J.
H.
Finally
the
we
M.
and
Os
would
and
due
was
H.
of the
in
helpful
Mr. B.
for
kindly put
Dr.
to
palynology
Bongers
van
Callen-Lobreau
data. She
also
are
the taxonomy
Mr, J.
improved
Mrs. D.
to
palynological
lengthy
some
Sleumer
family.
and
Mr.
are
Mr.
W.
critically testing
N. Kieft
discussions
unpublished
the
acknowledged
like to thank Prof. Dr. C. G. G. J.
van
J.
Star
on
the
informa-
Muller
kindly
for
gave
synoptical key.
for their
Steenis who
share
in
critically
manuscript.
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