BLUMEA
25(1979)513-529
and fruits
Flowers
Hydnocarpus
Flacourtiaceae. IV.
Berberidopsis corallina Hook.
spp.,
Casearia
in
Kiggelaria africana L.,
spp.,
W.A. van
Rijksherbarium.
F.
Heel
Leiden.
The Netherlands
Summary
In
the hard
Hydnocarpus
where
tegmic.
Corner’s
pachychalazal
of the seed coat
layers
develop from
the
epidermides of
This does not conform with the division of the
they are contiguous.
suggestion
one, cannot
of two unallied
groups
be corroborated.
in the
Flacourtiaceae, namely
The seeds
of
both
Angiosperm seeds
an
or
integumental and
resemble
Kiggelaria closely
integuments
into testal
those
of
a
Hyd-
nocarpus.
development of the
In Casearia the
American
are
exotegmic,
The
studied. The three
species
as
berries
described
and
seeds
parenchymatic raphe.
inner
epidermis
beridopsis
by Corner;
of
The
fruit wall and its vascular bundles is different for the Asian and the
parietal placentae
the ovules
Berberidopsis
of the outer
are
integument
forms
a
for
the first
integument is
the base ofthe ovary. The seeds
time.
absorbed
The seeds
have
softly
a
during development.The
lignified crystalliferous layer.
The
of Ber-
affinity
is discussed.
Hydnocarpus
The flowers of
equal
over
atropous.
described
is small. The inner
embryo
confluent
are
are
number of
Hydnocarpus
petals,
ventral scale. The
and
pistil has mostly
ovules. The sessile
stigma
stamens
4 or 5
composed
or
Mostly they
as
or
5
sepals,
petals have
covered with
parietal placentae
of
have 4
staminodes. The
an
basal
a
anatropous
reflexed, widening, bifid lobes
many
with these. The fruits
indehiscent, large
as
there
are
hard,
and contain hard seeds. The reader may find detailed descriptions in Sleumer
(1954).
and
is
unisexual.
are
many
spp.
placentae,
The anatomical
recently
by
Vaughan
anatomical
alternating
structure
data, especially
of the seed
Corner
and
(1970)
as
to
the
was
are
and
described by Sleumer (1938) and
In
(1976).
relation of the
the
following
pistil, ovule,
some
and
more
seeds,
are
presented.
As
septa
not
a
regards
the
at most
just
form
stylar
result the
central
pistil
canals corresponding
cavity
split,
it is notable that the
touch in the centre, but
of the
which issues
series of cleared thick
The locules
are
pistil
on
mature
even
fruit is
sarcotestae
small lobes
not
continues
a
pistil
compared
matures, the endocarp forms
placentae,
formed
with the
a
can
by
soft
in
not
carpels
upwards
figs.
fuse
5
—
which constitute the
into
a
five-fluted,
of
endocarp
level the
or
and do
pistil.
This is illustrated
tissue, especially
pistil.
in the
As the
regions
However, the pulpous tissue
but is
of the seeds. At the very bottom of the
As
irregular,
by
a
8.
with the massive wall of the
mass
stylar
secondarily there,
stigma above.
be formed here.
the
is very short. At the
do
the surface of the
slices of
small as
style
they
entirely
pistil
due
to
pistil
of the
in the
the confluent
there is either no
placental
514
BLUMEA
Hydnocarpus polypetala. Fig. 1.
t.s., 7
x
.
Hydnocarpus
kurzii.
Ovule
Fig.
2.
l.s.,
-
60
VOL.
x
Vascular
25,
.Figs.
3
bundles
No.
—
2,
1979
8. Vascular
in
half of
bundle pattern of
young
pistil,
cleared
seed,
cleared, 7 x.
W. A.
tissue,
the
the
or
is
the latter
placentae
base
to
in
the
at
upwards. They
the ovules
split
sterile
the
bundles in
base and in
provided
are
with their
have their
of the
base of the
divide into
The ovules
are
ovule
branches
and
the direction
on
placentae.
on
forms
a
but has
Below,
they
of the
from the
lateral
vascular
bundles
carpellary
In the
stigma.
traces
blindly
of the basal lateral
origin
the median and
The
pistil.
run
from the
originate
is
connections
some
and many inward
6),
the
bundles which
have ramifications which end
level above the
a
Upwards,
branches
spread
fan-like in
into the
of the nucellar base,
developing
or
outer
outer
the
lobes
stigmatic
small seeds this is
definite endopyle.
In H.
of the
and
raphe,
anastomoses
Short
move
the
into
from the base of the recurved branches
coming
outer
distal part
integument.
does
integument
distal parts of the
prolonged
is notable that
tapering region
the outside (fig.
not
reach
unchanged,
However, in well developing ovules and seeds
the
it
8),
—
and fan-like.
In the ovules the
less well
3
nucelli (fig. 1). The vascular
anatropous, bitegmic, and have large
bundle of the
curve
region.
anastomosing
number of smaller bundles toward the
a
they ramify profusely
(fig. 2).
upward
may touch each other in
they
vascular bundle strands which
Above, they
pistil,
vascular bundles.
carpellary
the
own
xylem
off from them.
regions
the
of
515
IV
Flacourtiaceae
pistil (figs.
'cortical' system
a
in
from the main vascular bundle system,
largely separate
with
fruits
their base and
at
of the
supply
supplied by
and
sterile locules in that
pistil, forming
the vascular
regards
As
massive wall
Flowers
may be sterile
placentae
of the
centre
Heel:
van
an
The inner
the inner
polypetala
is formed
ectopyle
integument.
the inner
beyond
that there is
so
as
a
In
one.
ectopyle.
no
short slit
by
integument always
integument
has
terminal
two
and lateral lobes.
The
development of
integument
in its
and
proximal
cative,
half. Thus the
cuticle between the
in
adjacent
On the
places.
outer
integument composing
integument
is
the
seeds
of
Towards seed
consisting of
Hydnocarpus
formation,
are
elongate
and
ment
to
differentiates into
the Soft
pulpous
inner
the
both
multipli-
in the
two
fruits,
layer originates by
integument, starting
cells of these radial
integuments. However,
in my
inner
middle
later
and
namely
layer
that the
on, so
inner
longitudinal
elongate
also
as
outer
to
divisions
opinion
the
testa
and
no
an
outer
layer
regions
is
one
as
a
integu-
giving
rise
which forms the
elongate
rows on
outer
an
cells.
separated
The
Sleumer)
of anticlinous
sclereids. The
the inside of the
outer
of the young seed. Later the innermost
radial cells which
sclerified,
axis of the seed,
parenchymatic
to
tegmen
parenchymatic
pigment layer
acc.
a
of periclinous
layer
body (' Kernhaut ').
the formation of radial cell
rows are
of cell
integuments,
an outer
of undifferentiated
Van der Pijl, aril
an
the
According
differentiates into
integument
of cells,
layers, namely
sensu
or
first
outer
occurs.
layers
from the distal
cells of these radial rows
that level.
because
at
that the
so
distinguished,
on
pachychalazal,
endosperm-embryo
layer (sarcotesta
mass
be
testa can
in the direction of the
a
layer desintegrates
dark sheath around the
are
of tissue
growth
pigmented layer,
The middle
are
persist,
the loss of the cuticle between the
three different thick
sclereids which
inner
the young
below the insertion of the
pachychalazal
level either the cuticles
remain partly free from each other
entirely integumental by
such
integument
inner
outer
especially
and inner integument vanishes gradually,
endopyle
and inner
occurring
and the
outer
and inner
Corner
of the cells of the
by enlargement
division of small cells of the inner integument
Corner, and may be distinguished by the dimensions of their cells. The
sensu
persisting
the ovule is marked
by profuse
but do
not
are
sclerified. The
first
outermost
elongate radially, and
in
BLUMEA
516
-
VOL.
25,
No.
2,
1979
1
2
Hydnocarpus glaucescens. 1.
t.s.,
slightly
younger.
Differentiation
of the
integumentsinto
seed coat, l.s.
(see text);
2.
Ibidem,
W. A.
this way
derived from both
outer
of the radial cell
integument.
The
outer
layer
formed
the
outer
integument
sensu
Sleumer is
layer
middle
Around the
caused
mainly
of
layer
which
and
this
forms
outgrowths
many
protruding
prior
outer
of the
layer
into
peak-like
is
rest
is absent, the
layer
dominant. In the subdistal part of the seed the sclereid
integument
the
periclinous
derived from the
are
endopyle
the
by
by the
without
sclerify
formed
are
by
which
Sleumer,
Sleumer is formed
sensu
heavily lignified
less
are
of the inner integument.
pigment layer
inner
by
inner
the three
Consequently,
The hardness of the seed is
the
The
517
IV
have been described by Sleumer (1938),
they
sclereids,
longitudinal sclereids, which
layer
in Flacourtiaceae
andfruils
anticlinous
rows
elongation.
dermal
as
integuments.
layer of
formed by the
radial
Flowers
of the seed coat,
layers
dominant
Heel:
differentiate into small isodiametric sclereids.
they
sclerified
van
the
outer
integument.
stressed that the differentiation of the testa,
It should be
frequently
after the
starts
after the cuticles between the
young seed
tiations
integument
in this
each
before
start
the
entire
species,
which has
the distal parts
of the
with
is
testa
that
loss
the
of
as
new
growth
Flacourtiaceae
with
seeds.
an
as
even
ordinal level,
less
seed
a
a
the share of
in
so
far
from both
recognizes
Flacourtia group with
a
natural
possibly
similar way
thinks the
exotegmic integumen-
coat
is endotestal
in-
are
group
the Flacourtiaceae as
the seed
Hydnocarpus
relation of the
unity of
the
him the
to
Hydnocarpus
a
however, the seeds of the Hydnocarpus
a
a
Corner
there is
as
at
region.
also
consequence,
each
by comparison
originates
coat
Corner
of
multiplicative
Anyway,
remain free in that
As
assemblage
difference between testal and
important
and
are
differen-
share
and the tegmen has diversed in
testa
well, which pleads for
may indicate
the
be ascertained
boundary.
pachychalaza.
this character. The fact that in
exotegmic,
can
of the
part
seeds and
opinion,
my
integuments
small fruits and seeds.
cuticular
unnatural
pachychalazal
In
tegumental
are
that
so
described above (cf. his fig. 279) but according
of the
only
it is difficult
found in which the
were
cuticles,
integuments
hard
the
observed differentiationof the
tal
3)
differentiation
seed if the
the seed
composition of
comparatively
into the
The conclusion
integuments,
group
—
could be ascertained. Moreover, the
integument
into the
1
of the
loss
cases
above,
and
has in the differentiations. However, in H.
integument
stages (photos
described
as
integuments,
have vanished. In those
integuments
ascertain which share each
to
glaucescens
of the cells of the
multiplication
regards
as
well
as
Flacourtiaceae with Dilleniidae. The
tegmic seeds,
has its border
which
plays
a
role
at
family
cases.
Kiggelaria africana
The flowers and fruits differ from those of
such
as
the anthers
fruits could
of
Kiggelaria
Hydnocarpus
ovules.
strands.
carpel
is African.
the
Inverting ovule
traces
in
placentae
are
contrast to
Young pistils
(figs. 9—18), except
Correspondingly,
in
some
minor
aspects,
that the
do
not
are
not
the
genus Hydnocarpus,
very different from those
placentae
bear
have their
only
own
given off directly by the
two
rows
of
strong placental
commissural lateral
bundles.
The ovules
porus,
Hydnocarpus
pores and the dehiscent fruits. The dehiscence of the
be studied from my material. In
not
which is Asian,
opening by
the
are
anatropous, bitegmic, and have large nucelli. The endopyle
ectopyle
a
median slit.
Mauritzon
(1936)
reports that, when the
is
a
en-
518
BLUMEA
VOL.
-
25,
No.
2,
1979
3
4
Hydnocarpus glaucescens.
text).
—
Casearia
3.
tuberculoid.
Differentiation
of the
4. Differentiation
integuments into
of sclereid mantle
on
seed coat, l.s. at
inside of
micropyle (see
pistil wall,
t.s.
W.
is
dosperm
the
formed,
four cell
integument
with
Young fruits,
between the
cells of the
The
integument
start
and
reported by Mauritzon),
—
7
cell
seven
are
show the
places;
at
cells
a
consist of
inner
one
logical
to
of cells
the
of the
part
is
is
(which
rows
integument
of
two
as
difference here
no
in
the axis
to
the radial
lignification,
but here this
Hydnocarpus,
Also
chalaza.
of the
forming
a
tracheidal cup. There is
integument,
diameter)
which
the
raphe
there is in
as
curves
no
in
does
chalazal
in
ramify mainly
and divides
into the
antiraphe
direction of the chalaza, that is
The flowers of Casearia have
with
atropous ovules,
placentae.
a
a
style,
for instance
by
In the
Sleumer
following
(1954).
pistil
remains meristematic for
thickness. It forms
with
a
adaxial
spp.
distinct
a
layers give
differentiates
sclerified
some
the outside the
epithelium.
subepidermal
studied such
dermal
to
cell
Finally,
layers,
to
a
centripetally
its
fruit and
young
plays
sylvestris
6
—
7
and
outer
in
mm
bundle
grow
into
23).
about
stamens,
a
in line with the
in taxonomic
medianly
described
and seed
on
contributions,
are
responsible
for the
with many secretory
meristematic
activity
are
by
Corner
given.
is
growth
in
spaces lined
confined
to
the
formed. In the Asian
in thickness is absent. In all
species subepi-
sclereids. This meristematic mantle
stretched
this sclereid
role in the
ten
the adaxial tissue of the wall of
time and is
tangentially
a
of the
raphe
arillate seeds
pistil, ovule,
mantle of initial
into
—
stigmatic parts
where radial cell families
(fig. 30, photo 4). Presumably,
wall of the
on
mesophyll
specialised growth
rise
petals,
no
descriptions
data
19
tegmic.
chalaza,
pistil has three parietal placentae
The anatomy of the seed is
more
part
(fruits
integument
(figs.
capsule, exposing
In the American spp. C. arborea and C.
the
calyx,
and three short
three-valved
some
outer
usual
as
into the
Hyd-
spp.
five-lobed
the valves. The reader may find detailed
(1976).
to
outer
of the
coat
the
short
a
with the
from the base of the
trimerous pistil. The
short
a
The fruit is
a
outer
in later stages
side of the
opposite
Casearia
number of staminodes, and
into the
prolongation
developed
irregularly
over
region
line
A
place.
also stretches
layer
the
the inside
take
not
that the seed
opinion
Hydnocarpus. However,
lateral bundles have
two
an
seed. It is
nocarpus group is integumental with loss of the cuticle, and testal as well
The vascular bundle of the
found
was
elongation of
differentiation of sclereids is present,
my
coat
described above. Therefore the
as
that
their
These results corroborate
with
the result of the differentiationof the
are
except
to
also
of isodiametric cells, and
outer one
respectively,
nucellus and
of the
an
sublayers
prior
rows,
are
acropetal
an
diameterof 17 mm, which contained seeds
longitudinally according
Hydnocarpus,
present
boundary
prolongation
integument.
to
sublayers,
integument
radial cell
pigment layer
basal
stretched
inner
is similar
pattern
thick
equally
presume that these
and
outer
two
inner
of the cuticle
disappearance
with complete embryos and endosperm, the stony layer of the seed
to
the
thick,
in this stage the inside surface
There
begins.
In fruits with
519
layers
divide into radial cell
to
IV
multiplicative (sensu Corner).
the outside of the inner
on
longitudinal
Hydnocarpus glaucescens.
is
mm,
integument
in Flacourtiaceae
fruits
integuments
diameter of 6
a
differentiation into
Flowers and
integument
outer
layers.
and inner
outer
outer
van Heel:
A.
cells
which later become
layer gives toughness
opening mechanism
to
the
of the commis-
sural valves. The dehiscence will be facilitated by the fact that in the radii of the main
vascular
bundles,
where
the
valves
become sclerified, but remains
separate, the
composed
of small
meristematic mantle does
parenchymatic
not
cells. The bulk of
BLUMEA
520
Kiggelariaafricana. Figs.
t.s., 7.5
x
Vascular
(fig.
24:
bundle
15
x).
pattern
9—18.
Distance
of
-
VOL.
Young pistil,
between
pistil. Base,
25,
No.
series of t.s., 15
sections indicated
cleared
t.s., 15
x
.
1979
2,
Fig.
x
in
.
Figs.
µm.
28.
-
19
—
26.
Young seed,
Casearia
Ibidem.
series of
flavovirens. Fig.
Half, cleared.,
15
x
.
27.
W.
A.
van
the fruit wall is formed
by
Heel:
extended
are
described above
creating
In
C.
because cells
prior
the
to
of the
arborea and
the
on
It is
three small locules
by
a
is less evident,
pericarp
layer
vascular bundle
becomes
boundary
become stretched
placentae
species
or
irregularly
radialexpansion
a
mesophyll
the
thicker,
much
radially
growing by
mainly.
feature of the Casearia
placental ridges
this
sylvestris
this
placentae (a 'short-cut'),
and the
pericarp
of their walls. Moreover, in these
cells of the
of cells
special
a
C.
side of the
lignification
mesophyll
multiplication
three
between the
continuous
not
inserted: Instead,
are
of the
placentae with their independent
separate position
a
strands.
secretory
the whole fruit the differentiation
to
where the ovules
placentae
boundary running
a
mesophyll
the
expansion
the top and pursues downwards. It is remarkable that the
starts at
the surface of the
forms
result of this
radial slits. In relation
as
the cells of the
expansion of
a
521
IV
of the meristematic mantle and its differentiation is
development
over
mantle. As
in Flacourtiaceae
fruits
marked radial
a
tissue outside the scarifying
spaces
Flowers and
(figs.
spp. studied
confluent and
are
30
rise
5 and
32, photos
—
give
in the base of the
that,
As this
6).
the
pistil,
three-rayed ridge separating
to a
is
structure
not
developed
secondary approximation of the placental ridges, the pistillary apex possibly
may
into this
develop
pistillary
C. arborea this
recognized
ovular
as
stalks
narrowing
space
on
is
phenomenon
the
ridges along
over
of the
the
wall.
pistil
pistil,
fuse
level of the
study
to
ridges
ridges
in the middle,
tuberculata,
stigma.
the
this
the three
the
Upwards,
Distally
secondarily
after the initiation of the
at once,
permit
not
prominent. Epibasally,
their whole surface.
small locules in C.
(with
again
three-rayed configuration
wall. However, my material did
sterile, and,
a
ripe
this
stylar
space opens
fruits could
open
the
on
three-fluted
fig. 35). Apically,
see
Unfortunately,
In
be
can
fall apart into many
are
leaving
development.
placentae
be
not
studied.
The pattern of vascular bundles in the
A stele
pistil
of many small vascular bundles
bundles of the
upwards forming
adjoining perianth parts
the stele toward the
The
stamens
stele
pistil
three alternate
here),
which
of
ramify
as
later in the
departure
In C.
two
collateral
to
traces are
a
continuous
arborea and
C.
ontogeny of the pistil
stele which then falls apart
vascular bundles of the
tuberculata and
flavovirens)
the
confluent
sylvestris
wall
into three
or
as
the
off
There
a mass
placental ridges.
strands fade
out.
bundles
(C.
The
complex
(C. arborea),
(C. arborea),
calodendron),
or a more or
no
converge
connected
to
are
originate
Above
the
can
be
much
level of
bundles converge into the
the
complex
strands of
either cross-connections
arborea)
placental
(cf. figs. .28, 33)
(C.
in the base of
strands
placental ridges
concentric
up,
these
cross-section,
'median laterals'
bundles'.
Portions of the
less
main
the Asian spp. studied
(in
of bundles (C.
i
strands consist of
or a more or
37.
—
higher level, according
wall. On
remaining
off toward the ovules. In the
sterile
of the
parts
upper
array of bundles
or
stamens are
lateral bundles
'cortical
the
27
Five
pistil wall first; then, higher
pistil
the
figs.
inwards.
portions, forming
ridges.
there is
or
tangential series,
pistil wall,
placental
lower
given
the lateral bundles of the
as
of the bundles of the
three-rayed
on a
outwards
more
three main bundles
gives off
sets
lateral bundles appear
distinguished.
placed
are
the
remaining bundles
of the
traces
by
flower base.
five alternate commissural
by
each. The
The
pistil.
with the above mentioned vascular bundles
whether the
into the
followed
perianth parts separate off,
lateral traces of two
may be illustrated
diverges
either
are
split
the vascular
an
irregular
arrangement of collateral
less concentric arrangement of collateralbundles
especially below,
or
an
inverted
large
bundle
(C. tuberculata),
or
BLUMEA
522
Casearia tuberculoid.
t.s., 15
x
(fig.
41:
30
Figs.
x).
29
—
37.
Pistil,
-
VOL.
25,
No.
series of t.s., 15
x
2,
1979
(fig.
29: 30
x). Figs.
38
—
43.
Young
seed
W. A. van Heel:
again
series
tangential
a
vascular strands have
clear that
their
The ovules
the
dermal cell
two
layers
have
integuments
cells
only
38
(figs.
—
The
As
sclerified
a
embryosac
placental
pistil wall, they
in its
funicle,
the
The endo- and
plate
concave
outer
formed
at
cells
On
than the
inner
The
both
of
early stage, always
The
inner
outer
outer
in its base, but
seed, spreading cup-like
it,
epidermis
At
thin
walled
an
early stage of
seed
In its
more
the inner
integument
proximal region
or
less in
nucellar
is
planate embryo,
formed.
is
as
the seed is
forms
Newly
received
to an
(1977). Hitherto,
Possibly,
the
fruiting
fruits
fruits
have
are
and the
the
a
the
one
present
a
to
the
enclosing
some
are
no
fruits
are
which
protrudes
with
of the mesocarp.
almost
are
in
a
enlarged
small-celled
outer
small,
surrounds
on
some
a
Oncobeae
of the fruits and
mature.
length
and
torus or
The
and 9
stigma
are
mm
seeds.
fruits
are
in width.
persisting.
disc between the
that the wall of the fruit is
between the
bundle
of
The
lobed hard rim persisting between the stalk
indications of lignification,
berries,
the
in its natural habitat in
minor lobes may be present between the
pistil. Considering
that there
descriptions
style
the
large
layer
endosperm
in my paper
The
fruit, which represents
Also
no
material
a
the
integuments. Many
abscission
species, collected
alcohol).
enters
corallina
mm
fruit-stalks,
a
prolongation of
in Flacourtiaceae.
present measured 14
brown-red colour (in
attached
stamens.
and the
in
copious
descriptions given
taxonomic works gave
roundish, the largest
They
material of this
addition of the
A
form
remains
connected with this
are
with the
continuity
basal
into the
a transverse
common
Berberidopsis
Chile, leads
reaching
development
cells
parenchymatic
sized
moderately
not
a
become lined with the suberized
epidermides of
the
outer
its inner epidermis into
brachysclereids
vascular bundles of the aril
longitudinal
on.
Both
remains
integument
sheath downward into the chalazal tissue,
integumental sclerified
integument
integument.
its
integument,
integument.
top
layers
and consists
soft cell layers of the testa, consisting of
of sclerified tissue
layer.
which probably
subdermal
laciniate,
an
integument.
ectostome
by
outer
may be
sometimes
absorbs the central distal part of the
(Corner) longitudinal sclereids,
rows
integumental
later
as
proximal part, by
aril
an
or
distal part of the vascular bundle cup. A large complex vascular bundle
small
well
as
integuments
two
ninety degrees
ectostome
layers distally.
suberized plate of tissue. Moreover a rim of
the
to
integument
outer
larger
regards
the inner epidermis of the
protected by
the inner
the endo- and
and forms the
cells of small radial cell
by
the
case
be secured,
can
the distal part of the
dermalcell
44).
pigment layer (photo 7).
and
on
fewer but
two
into ribbon-like
develops
layer
into
and
entirely parenchymatic
large
of up
angle
an
the distal cells of the
remaining covered by
has fewer cell
placentae
distal part. The ovules have
layers only.
protrudes
nucellus and
of the
later. The distal half of the aril is lobed to
being incorporated
of
in its
from the dermal
originates
at
micropyle is formed by
integument
outer
supply
the nucellus with
atropous. However,
are
The
even more.
no
the sclerified mantle described above.
the end of the funicle
at
In
(C. cf. flavovirens).
523
IV
connection with the lateral bundles of the
a
fruit maturation the
at
protection by
attached
of bundles
in Flacourtiaceae
fruits
from the vascular bundles in the flower base. This construction makes
directly
come
Flowers and
rind
nor
scars
and
a
seems
softly parenchymatic
no
and
stamens
entirely parenchymatic,
of dehiscence lines, it
seeds. There is
perianth
of the
and
that the
endocarp
differentiationinto radial cells
524
BLUMEA
-
VOL.
25,
No.
2,
1979
5
6
Casearia
tuberculata.
5.
Three-rayed placental ridge
in
ovary
base,
t.s.
—
Casearia
arborea.
6. Ibidem.
W. A.
Casearia
clutiaefolia. Fig.
Twenty
and 2
to
thirty
44.
are
are
ovules,
at
.
x
present in
—
by
integument.
C. arborea.
one
are
bundle
A
integuments.
is the first
two
the
cells. The inner
epidermides
walls
integument
lignification
crystal
at
(figs.
of
which
at
are
to
layer
of tissue,
epidermis
the
at
the
pale, softly
the
the
at
signs
there is
a
lignified
ectopyle also mesophyll cells
for the
palissade layer,
which
is
present
nucellus and the
a
inner
partly
causing
Only
a
no
forms
layers,
and
remain
of many small
or
2
near
layers
of
mesophyll
two
not
of the
epidermis
outer
a
show
inner
the chalaza; it
and
also
of palissade
pitted.
Each cell contains
integuments
layer
can
the
outer
only
one
a
be
the
rhomboid
partly
with that
by comparison
as
cells,
Frequently,
free
region
integument.
This
that, in ovules
in the form of cell
activity.
at
At the
undulating lignification of their walls. Except
disrupted,
integument.
absorbed.
inside and radial walls,
the
red,
consists of 1
the inner epidermis of the
show the
the
by
chalaza,
raphe, however, they
endopyle
it could be verified
conspicuous layer
between
narrow
of the ovule consists of
conspicuous layer
beginning differentiation
of
a
their cell pattern, but do
regular by
the cell in the form of bars.
ectopyle,
the
staining
has four cell
of its ends. As the distal
of the
parts
shows
and
long
disappear.
photo 8)
palissade layer represents
disrupted
mm
bordered
ectopyle
little in
mesophyll
post-anthesis
is in accordance with the observation of this
anthesis,
of
integuments,
two
wide
a
at
integument
undulatingly
are
traverses
from each other
that the
cell
47 and 48,
which
one
3
.
x
large polygonal
integument consists
outer
becomes absorbed, except
soon
In the seed
of the
15
t.s.,
angularovoid,
are
spreads
outer
nuclei. The
containing large
differentiation. In ovules
integument
Young fruits,
contents
patch
safranine;
of the
epidermis
and
integument,
cells which stain red with
larger parenchymatic
and
45
525
IV
protuberant raphe consisting
a
vascular
raphe
the
entering
isodiametric cells
layers,
the
anatropous; they have
the inner
The
unstained. The inner
any
Fig.
berry, they
kind of hypostase. The nucellus is thick. The
large cuboid
in Flacourtiaceae
fruits
(in alcohol) by
marked
anthesis,
endopyle bordered by
ramifications
15
and
tissue.
parenchymatic
outer
Flowers
Seed, ± l.s.,
seeds
surface cells. The seeds
The
Heel:
reddish brown
wide,
mm
van
in the seed is formed
but
by
partly persisting,
All the cells of the inner
the inner epidermis
polygonal pattern
on
the thick cuticle
epidermis
integument
persists
the cuticle
of the
become
in the form of its
layer,
or
rather it
BLUMEA
526
-
VOL.
25,
No.
2,
1979
7
8
Casearia
Base
clutiaefolia. 7.
of young
Differentiation
seed, ± l.s.
of the
integuments into
seed
coat.
—
Berberidopsis corallina
8.
W.
A.
van
and
fruits
be the cuticle which penetrates between
may
have
a
small
seed
are
the endotestal
embryo; 2)
in
Flacourliaceae
the inner
of cells,
copious endosperm, consisting
Four characters of the
the
Heel: Flowers
with
cells. The seeds
epidermis
small
a
527
IV
embryo
unusual for Flacourtiaceae
very
development
as
the
Berberidopsis
reasons
beridopsideae,
and
be
to
similar
of the
strength
much like
very
that of
also
as
belonging
to
clearly
Magnoliidae
exotegmic
This
than
is of
small embryos.
In
Berberidopsis.
much
course
evident
Unfortunately, however,
parietal Magnoliid
and
does
(Miller, 1975) follows
that
1973),
purely
well
has
Berberidopsis
a
contact.
coat
especially
occur
partly
are
as
mixed
a
Flacourtiaceous,
affinity
an
structure as
From
of the
with
far
as
study
a
of
of the
xylem anatomy
within the
primitive position
very
described above, also
present in Dilleniaceae. Possibly,
endotestal seed in
em-
although
Berberidopsis (and
Strep-
in the Dilleniaceae, and also in the Theaceae. The seed
anatomy of Berberidopsis,
embryos
study
a
Flacourtiaceae. Miller thinks many wood characters
tothamnus)
large
Berberidaceae, Lardizabalaceae,
as
from
as
coat
endotestal seeds and the
the rather diverse seed
families
as
is
the indication of
present many points of
not
pollen morphology (Keating,
has
seed
group, viz.
Hydnocarpus
the
by
a
Erythrospermum,
Erythrospermum
Berberidopsis
more
known from such
Papaveraceae
in the
coat as
of
specimens
Herbarium, have
that
means
Evidently,
one.
Erythrospermum, which
genus
our
on
with three
gynoecium
Furthermore, Erythrospermum
the Oncobeae, has a seed
endotestal and
more
Kiggelaria.
Warburg reported.
from
judged
I found that various
Berberidopsis. However,
from New Guinea, present in
Erythrospermum
bryos,
with
Ber-
Warburg (1893),
authors considered
prior
dehiscent anthers and the
longitudinally
similarity
rank.
the genus in the Flacourtiaceae
Warburg placed
parietal placentae. Another argument concerned the
has much
separate tribe (cf.
a
given family
Berberidaceae and Lardizabalaceae,
to
external appearance. However,
the
forming
as
be
must
or
the genus in the Flacourtiaceae. All
Gilg (1925) placed
Berberidopsis
1967),
of the inner
in Flacourtiaceae as well. For these
rare
be considered
must
Hutchinson,
is
fleshy raphe
the
1)
crystalliferous palissade layer; 3)
a
conspicuous notched cuticular layer; 4) the lack of differentiation
integument. Presumably,
it.
in
namely
,
Hydnocarpus,
and the
ancient relation of the Flacourtiaceae with
the
perianth
in Theaceae. Minute
occurs
structure
of
the
Berberidopsis,
of Oncobeae, all indicate an
Dilleniidae.
ACKNOWLEDGEMENTS
My
sincere
thanks
(Stellenbosch),
(Leiden), Dr.
as
indicated
subsidized
Dr.
H. Sleumer
by *,
by
Maatschappij
was
P.
due
to
the
Directors
Ng (Kepong),
(Leiden), all
collected
for
during
voor
Rumphius Fonds,
during a
visit
subsidized
of
my
the
visit
mentioned,
Steenis
to the
Kebun
in
de
Mr.
part
of the
in
Bogor
material,
of
Drijfhout
van
in F. A. A. Part of the
1969.
of Tropical Research
University
P.
M. J.
Malaya
for the Advancement
as
Balgooy
material,
This visit
was
The
(WOTRO),
Tropen (Treub-Maatschappij),
Another
Organization
at
to
Dr.
(Leiden),
or
Raya
for the Advancement
Leersumfonds.
the Netherland's
van
material in alcohol
short visit to the Botanic Garden of the
by
Institutes
C. G. G. J.
Onderzoek
Wetenschappelijk
and the Van
Dr.
collecting
the Netherland's Foundation
collected
was
are
F. S.
the Greshoff's
indicated
at Kuala
by **,
Lumpur.
of Pure Research
was
This
(ZWO).
MATERIALS
Hydnocarpus alpina Wight:
Campus University
H.
kurzii
Garden
of Malaya, Kuala
(King) Warburg,
Garden
Sleum, Sarawak, Jacobs5427; ibid.,
Kepong**,
van
Heel
1137.
—
Economic
Lumpur**,
Economic
Kebun
Plants, Bogor*,
van
Plants, Bogor*,
Raya, Bogor*,
Kiggelaria africana L.,
van
Heel 433.
Balgooy2189,2253, van
Bot.
van
van
Heel
Heel
104.
—
432.
—
H.
—
Heel
glaucescens
1118,
H.
Bl.:
1146,1147.
-
polypetala (Sloot.)
H. woodii Merr.,
Arboretum,
Heel
1148; ibid.,
Garden, Stellenbosch,
van
528
BLUMEA
Berberidopsis
coat, t.s., 220
x
corallina.
.
Fig.
46.
-
VOL.
Ovule, l.s., 120
x
.
25,
Fig.
No.
47.
2,
1979
Seed,
transmedian
s.,
30
x
.
Fig.
48. Seed
W. A.
Bot.
num.
—
Casearia arborea
BW3655.
Sleumer
1140.
C.
—
s.
van
C.
(L.
cf. flavovirens
num.
Cameroon,
al.
Kirstenbosch,
Garden,
van
—
Voucher
—
specimens
For
Flowers
in Flacourtiaceae
fruits
23811;
Rich.) Urban,
Brasil, Steumer
Bl., Kebun
Bl.,
are
in
ibid.,
Raya, Bogor,
Kebun
Berberidopsis
the
and
Steenis 23810,
C. tuberculata
Breteler 1148.
Heel:
Bot. Garden, Pretoria.
Burck
s. num.
s. num.
Raya, Cibodas*,
corallina
529
IV
van
—
Lotsy
&
Goddijn
s.
clutiaefolia, New Guinea,
C.
—
C. sylvestris
Heel 373.
—
Swartz, Venezuela,
C. calodendron
Gilg,
Hook./, Quebrada Honda, Chile, Marticorena
el
L.
meaning of*
and**
see
under
Acknowledgements.
REFERENCES
CORNER,
E. J.
GILG, E.
1925.
HEEL, W.
A.
H.
VAN.
J.
HUTCHINSON,
KEATING, R. C.
Gard.
MILLER, R. B.
tiaceae.
H.
1954.
of
Dicotyledons.
Engler
und K.
Prantl,
Die natiirlichen
1977. Flowers and fruits in Flacourtiaceae
1967.
1973.
The
genera
of
Pflanzenfamilien.
III. SomeOncobeae.
flowering plants. Dicotyledons.
Pollenmorphology and relationships
1936. Zur
1975.
of the
Vol.
2. Aufl.
21.
Blumea 23: 349-369.
2.
Flacourtiaceae.
Arb.
J. G.
O.
6a:
anatomy of the
56:
Monographie
Flacourtiaceae.
VAUGHAN,
1970.
The
Ann. Missouri
Bot
xylem
and
Svensk
comments
on
the
Bot.
Tidskr. 30: 79—
relationships
113.
of Flacour-
20— 102.
der
Flora
Gattung Hydnocarpus
Malesiana,
structure
1 893. Flacourtiaceae.
15.
Parietales-Familien.
Embryologieeiniger
Systematic
J. Arnold
1938.
WARBURG,
Ill,
A.
60: 273-305.
MAURITZON, J.
SLEUMER,
1976. The seeds
Flacourtiaceae.
ser.
I,
and utilization
A.
Engler und
K.
5:
1
—
Gaertner.
Bot.
Jahrb. 69:
1 —94.
106.
of oil seeds.
Prantl,
Die natiirlichen
Pflanzenfamilien.
1
Aufl.,