BLUMEA
22
(1975)
175—195
Comparative anatomy
of
the
1
van
G.J.C.M.
Crypteroniaceae
Vliet
Rijksherbarium, Leiden,
lato
sensu
and P. Baas
Netherlands
Summary
The
and nodal
leaf, twig,
is described in detail. This
be
to
heterogeneous,
very
The nodes
paper.
(or split lateral) plus
is
diversity
vascular
present
bundles
Anatomical
Axinandra
for
and
to
bers
of
as
and
Van Beusekom-Osinga and Van
the wood
trilacunar.
with
mechanical
only
Crypteronia.
of
diversity
Crypteroniaceae.
supports
These
not
Myrtalean
the
together
genera,
of the
which
family
with
its
character
with
appears
in
another
common
be present in addition.
may
the
veins,
wide
of all
gap
Further
of
arrangement
foliar
crystal complement,
anatomical
to some
Alzatea
known
shows
to
occur
Lythraceae, Oliniaceae,
of intimate
relationships
show
would
range
and
a
affinities with all families
Oliniaceae
,
mutual
features
also
affinity
cover
than
to
but remains
of
characteristic
most
of the
Rhynchocalyx. Rhynchocalyx
mentioned,
complex
close
several
and Melastomataceae
in the other families. The
between
and
genera,
Dactylocladus
Lythraceae, Sonneratiaceae, Oliniaceae, Alzatea,
Crypteroniaceae s.l., Melastomataceae, Lythraceae,
evidence
trace, of the
hypodermal development,
type,
support
diversity reported
complete girdling
Cortical bundles
stomatal
texture,
a
anatomical
Beusekom,
origin.
be closer in its
anatomy
of
supported by
midrib,
Rhynchocalyx
,
or
type,
cuticular
e.g.
petiole
evidence
its trilacunar node
ranges
trace
and
Alzatea, Axinandra, Crypteronia, Dactylocladus
newly delimited by
conclusion
a
median
in
as
simply unilacunar, unilacunar
Melastomataceae,
some
anatomical
pears
in
and cork
sclereids,
be
may
of
anatomy
family,
the
other
ap-
mem-
problematic
with
anatomical
pattern of overlapping
and Sonneratiaceae may be
interpreted
these families.
Introduction
In the present
family
issue of
These five
and Dactylocladus.
acters,
absent
or
Blumea,
uniting
Crypteroniaceae,
of
rare
Van Beusekom
Alzatea,
genera
occurrence
share
are
also characterized by
a
that of Melastomataceae and
however,
open
contribute
and
to
to
to some
our
compare
the
paper)
and
are not
necessarily
only
(presence
families
results
new
sense
appears
taxonomic
intra and
this
those
to
have
(Van
Data
on
anatomy
relic
char-
they
was
factual base
of the
comparable
family remains,
therefore undertaken
(Van
1975)
Vliet,
and
1975,
to
genera,
pollen
and this
(this paper).
five
genera
in the wood and
mentioned in
Sonneratiaceae) will
has been reviewed by Van
a
Myrtalean affinity for all
intraxylary phloem
the literature
as
are very
1965) briefly described the vegetative
in leaf and
putative
be considered in the
leafand twig anatomy of the Crypteroniaceae s.l.
and Stant (in Lourteig,
a
Beusekom,
of wood anatomy
history of all
pits
order,
Melastomataceae, Lythraceae,
Crypteronia
inter-family relationships of these
macromorphology
on
1975) with
of vestured
from
with
primitive
close. Furthermore,
very
Beusekom. Since the anatomical evidence clearly supports
genera
delimitation of the
Myrtalean families, implying that mutual
doubt. A detailed anatomical study
leaf, twig, and nodal
The complicated
number of supposedly
Sonneratiaceae. The naturalness
understanding of
morphology (Muller,
new
combination of more specialized features by which the delimi-
tation of the Crypteroniaceae in the
to
a
in other
affinities between the individual genera
a
proposes
Axinandra, and Rhynchocalyx
twig)
relatives
(e.g.
comparisons.
scanty.
anatomy
Hallier
(1911)
of Alzatea. Nie-
BLUMEA
176
recorded
denzu (1892)
Metcalfe
the
(1937) did
of the
these
to
Metcalfe
1975
for
information
some
Crypteronia,
Axinandra and
on
publications will only be made as far
a
Only herbarium specimens
Since the
for each species.
(Van Beusekom-Osinga
were
glycerin
available for this study. Collecting data will be given
in this
were
Beusekom,
prepared according
to
The anatomical terminology employed,
&
Baas,
as
studied
were
far
term
AND
node has been
employed: the node
is
of
as
at some
stem.
For
practical
use,
1973; Van Staveren
(Fig.
applied
1
are
1914) provides
Crypteroniaceae the
reasons
stem
In
the
even
traces
(1970)
conforming
to
complex
in the
same
97773,
a
single
traces
fail
On
to
one
one
gap
as
enter
course
of
deviation of the
not
be taken
on
as
a
devel-
Crypter-
the number of gaps and
1974). However,
that this classification
node differences
stem.
cannot
in
be
sections
per
species
descend from the leaf. The
species.
In C.
C.
and fuse with those from the
e.g.
or common
Rhizophoraceae.
In
off from each lateral
situation is
comparable
the stele after fusion
on
C.
trace
ib).
to
the
In
through
one
leaves with the
variability within
or
large
griffithii and
two
exist between the vascular
may
More evidence of
the median bundle (fig.
fig. ic) the
a
of the vascular bundles in
oppositely arranged. Serial
the split lateral
for several families
(lateral anastomosis) splits
gap.
are
of the
in all
girdle around the stele
mon gap,
men
within
opposite sides
Crypteronia three
ex-
Baas, 1973).
species.
the leaves
mediately with the stele
or
a
asterisk.
of vascular bundles will be
course
This should
ontogeny
found in the few instances that several nodes
traces
the
stem.
useful descriptive tool (Howard,
a
an
same as
in which the
therefore afford the study of vascular connections of
stele. It appeared that
at
data on the
such
pattern is
or
genera
Crypteroniaceae
nodal region
patterns
no
&
the
semi-
different ways. For this study the
in
available. The classification of nodal types based
(Sinnott,
employed for all
was
since
is
in
secondary
—4)
is that part of the
described as descending from the leaf into the
opmental interpretation,
stage of
marked with
are
uncommon
the vascular bundles from the leaf to the stele (or vice versa) produces
vascular pattern in the
1970;
DISCUSSIONS
Nodal anatomy
In the literature the
(Baas,
only bleached and studied
were
Bongers
1973;
RESULTS
following definition
origin will be cited.
standard techniques
thickening. Specimens used for the study of nodal anatomy
plained elsewhere (Jansen
also used for the revision
were
herbaria of
made of unembedded twigs of nodal and large parts
were
All nodal regions
mounts.
study
no
These serial sections
of internodal regions.
In all
to
METHODS
1975)
Van
Bongers, 1973). Serial sections
permanent
AND
specimens used
&
Sections and macerations
some
Crypteronia.
is relevant for the discussion
as
detailed histological study of leaf and twig, special attention is given
MATERIALS
traces
&
Sprague
of vascular bundles in the nodal region.
course
oniaceae
and
Chalk (1950) in compiling data from
&
results.
present
Apart from
the
included
only
No. 2,
22,
anatomical characters
some
for Rhynchocalyx.
so
older literature
Reference
VOL.
specimen
were
a
species
studied.
median bundle fuses im-
macrophylla (fig. la) lateral
opposite leaf
to
enter
condition described
gap
a
com-
by Howard
additional centric bundle
cumingii
an
to enter
the stele
C. paniculata var.
at a
level just below
paniculata (Kochum-
that in C. cumingii but the girdling lateral
place where
side of the node the lateral anastomosis
was
one
would expect
found
to
a
common
fuse with the median
G.
Fig.
1—4.
Idealized
Dactylocladus;
2a.
—
1b.
C.
A. beccariana
J.
C. M.
van
Vliet & P. Baas:
three-dimensional
cumingii;1c.
(similar
to
C.
diagrams
paniculata
var.
of nodal
types.
1.
Crypteronia
paniculata (Kochummen KEP.
Crypteronia paniculata p.p.);
Rhynchocalyx lawsonioides.
= cortical bundle; e.g. = common
Anatomy of Crypteroniaceae
2b. A.
alata;
2c.
A.
F.
1a.
177
C.
griffithii,
97773).
zeylanica. —
—
similar
to
Axinandra;
2.
3. Alzatea verticillata.
4.
c.b.
trace;
m = median
trace.
gap;
c.t. = complete girdling trace; l = lateral anastomosis;
l.t.
= lateral
BLUMEA
178
bundle above
C.
the level of its
the laterals
paniculata
as
the median
The fact that in Crypteronia
stele from
in
C.
paniculata
have their
makes it
two
at a
own gaps
to
the stele. In
the other three specimens
pictured
anastomoses
the median
cumingii)
C.
in
var.
value of these lateral
gap
a common
follows: Nodes with
traces
and
one
cumingii)
C.
gaps may
trace
be present
to
to assess
will be described as
gap
(to distinguish these from
reported by Howard, 1970).
as
a
as
it difficult
descriptions the different
a common
median
paniculata will be referred
in C.
as
pattern
to as
In the formal
anastomoses.
with split laterals plus
types where several lateral
The nodal
partly
anastomoses
paniculata and
var.
and
affinis,
situation where these
lower level (partly in C. paniculata
gaps
same
in their connections with the
vary
a
of
special
for Axinandra beccariana).
2a
(C. paniculata
gap
to
the stele in the
enter
fig.
no
fit them into the nodal classification of Sinnott. Their absence in
conditions will be referred
one
1975
Crypteronia species and the above reported variability makes
the systematic
having
at
paniculata and
var.
impossible
other
these lateral
three-traced condition
a
2,
from both leaves
the situation
to
No.
22,
complete horizontal girdling path showing
a
anastomoses
(similar
traces
into
entrance
form
locus of fusion; the lateral
VOL.
complete girdling
plus
trace
median bundle. Lateral anastomoses, ifpresent, will be mentioned in addition.
In Axinandra
may
the nodal patterns
—c). Cortical bundles
affinis (fig.
2a
be collateral
if sufficiently large,
or,
like extensions, and occasionally
periphery.
arranged
the four
at
in radial
rows
traces
at
the points of insertion of the lateral
cortical region, passing through the
bundles end in the
internodes
outer cortex
is
attached,
several nodes (fig. 2b). In A.
sectioned and
and part of
an
traces
a
coriacea two
no
as
cortical bundles
part of cortical systems
author
as
additional
girdling
in this
to
way
Dactylocladus
macrophylla
and C.
In Alzatea
conforming
the
fig.
2c
complete
trace
a
was
twig ( Meijer
SAN
found to be present in only
(the
over
37494)
one
node
were
found
(Howard,
a
all. As far
at
paniculata
1974;
or
common
since
they
are
are
gap
of Axinandra and Crypteronia paniculata
aware,
complete
re-
girdling
cannot,
trace.
The
however, be
also present in species without cortical bundles. The
system in Axinandra limits its
nodal anatomy conforms
to
use
a
as
the type
systematic
character.
described for Crypteronia
three bundles from each leaf
enter
the stele in separate gaps, thus
the well known trilacunar condition with three
the internodes
present in
continuous system.
we
affinis have only been
1886, 1887) and by the latter
Lignier,
split lateral
as
var.
griffithii (fig. ia).
(fig. 3)
to
successive nodes of one
in
situations with
traces
variability of the cortical
In
as
in Axinandra and Crypteronia
as
interpreted
no
cortical bundle system continuous
suggestive ofa
cortical system
ported
complete
is
outer
the cortical
internode, the other node and internode being devoid of cortical bundles.
In A. beccariana
girdling
and bend into the
various distances from the node. In A. alata
at
in
bundles descend from the
anastomoses
tissue. In Axinandra
zeylanica
layer of cork
wing-
with each
always fused
are
sections
decreasing
sectioned but the fact that both above and below the lateral
were
same?) cortical bundle
were
transverse
sometimes in
with the bundles
other just below the node. In the nodal region the cortical
lateral
In
corners,
In the latter instance the bundles
var.
in addition. These bundles
(amphicribal).
centric
situated
are
that in Crypteronia paniculata
to
however, be present
may,
through the internode these bundles
size towards the
all similar
are
are
Scarcity
attached
to
the lateral
of material made it
traces
traces.
Cortical bundles,
but apparently do
impossible
to
trace
not
the endings
form
a
of the
cortical bundles.
In
Rhynchocalyx
insertion:
a
only
one
leaf
trace
simple unilacunar node
is present and
with
one trace
enters
the stele
at
The unusual diversity of nodal anatomy in this small family calls for
Although
it would be
entirely unjustified
to
the site of leaf
(fig. 4).
attempt
some comment.
phylogenetic derivations of one nodal
G.
C. M.
J.
from the other, it is
type
reported here.
We
van
Vliet
possible
to
& P. Baas:
Anatomy of Crypteroniaceae
regard the simple unilacunar condition of Rhynchocalyx
condition within this
but
family,
the other
in
plus
plants from
are,
median
one
native
trace
to
and the unilacunar nodes with
Kochummen
of C. paniculata
97773
var.
versa
paniculata
Dactylocladus
1974) succeeded
common
or
by merely transferring
plus median
trace
situation
of C. paniculata and Axinandra:
trace
(fig.
isolated
again. Within Crypteronia there
common gap
complete girdling
a
diversity
an
similarities. Differences
from split lateral
vice
environment and back
greenhouse
to
or
as
gap situation in
common
trilacunar condition
a
transitions between the normal
moreover,
some
great, ifone considers that Howard (1970,
not so
are
the nodal pattern of Alloplectus (Gesneriaceae)
changing
gap
p.p.
show
genera
between the trilacunar nodes of Alzatea and the
and Crypteronia
series of part of the
morphological
construct
179
ic)
this
represents
more
or
less
intermediate condition.
Howard (1974) recently stressed the importance
vascular
arrangement
Applying this
to
the Crypteroniaceae there is
fundamentally
a
the entire pattern of changes in
use
taxonomic
common
gap
plus median
a
the petiole and usually also in the midrib. In Axinandra and Rhynchocalyx
connected
it is with
as
a
traces
closed ring
or
is found in the distal end of
plate plus latero-dorsal bundles
bundle is arc-shaped throughout,
purposes.
good correlation between petiole vasculature
Alzatea, Dactylocladus, and Crypteronia
similar types. In
separate adaxial and abaxial
a
the trilacunar node and
and nodal anatomy, accepting
as
to
the stem—node—leaf continuum for
in
complex
the main
petiole
simple unilacunar
or
petiole, and nodal anatomy present in the Crypteronionly few of the possibilities classified by Howard (1963), though he did not
node. The combinations of midrib,
aceae
even
represent
include the
common gap
plus median
Generic
Leaf, node, and twig
of
xylem
anatomy
twigs and trunks
Quantitative values
are
given
descriptions
of the individual
the subject
is
as
the full
of
range
a
studied.
Leaf (Fig.
In
5,
A.
Plate I,
surface
developed,
(subsidiary)
27—30
or
2;
to
Lamina
11 —15
—33 /urn
are
long,
than the
thick.
to
Midrib
an
of 2
range
1975).
of means.
6750*; Woytkowski 6196*.
Stomata confined
to
below the
abaxial epidermis,
warts
infrequently
/im
not
present in
prominent.
I
adaxially flattened
or
Guard cell pairs
ledges well developed.
Ule 6730 and of irregular
In
transverse
out-
section.
present
on
slightly larger
square,
same
level
as
unspecialized
either side of midrib.
tissue and rather compact mostly
slightly raised,
abaxial shallow arc-shaped
mainly
neighbouring
thick. Abaxial cuticle 7—11 fim thick, adaxial cuticle
cell-layer locally
layers of palissade
3 —7
guard cells.
abaxial epidermal cells. Stomata in the
cells. Adaxial hypodermis of
composed
Ule
Unspecialized cells of adaxial epidermis mainly
erect
(Van Vliet,
and
& Pavon
PERU,
hardly submersed
of traumatic
origin. Veins
square
range
26—28—31 fim wide. Outer stomatal
dorsiventral, 380—440
/im
full
intermediate between anomocytic and cyclocytic, with
Polar T-pieces absent. Cork
line, probably
or as
Indumentum absent. Cuticle granular, anticlinal flanges well
slightly curved.
cells, which
32
& Pavon:
separate
described below. Secondary
publication
II, 7)
view.
straight
anomocytic
&
verticillata Ruiz
are
genera
only,
ALZATEA Ruiz
Material
condition in his scheme.
trace
lignified
Mesophyll
spongy
abaxially prominently raised, supplied
bicollateral vascular
bundle and
one
to
adaxial bicollateral bundles. Whole system embedded in sclerenchyma,
tissue.
with
several smaller
abaxially
inter-
BLUMEA
180
VOL.
spersed with lignified parenchyma. Ground
phery but interspersed with sclereids.
No.
22,
tissue
Major
2,
1975
lignified
veins with
in
collateral vascular bundles, embedded in mesophyll and with
bundle sheath, enclosing adaxial and abaxial sclerenchyma
veins. Vascular system ofpetiole
Fig.
x 25;
5.
7.
5
—
11
12.
&
Alzatea
lucida
Camera
12,
x
verticillata
distal
part.
—
(de Joncheere s.n.),
calyx
lawsonioides
Caps absent from smallest
composed of a strongly incurved,
drawings
of vascular
systems in
petiole (5
—
peri-
hgnified parenchymatous
a
caps.
in
veins with
10)
almost
closed, bicollate-
and midrib
(11
&
12).
5
—10,
68.
(Ule 6750),
Crypteronia cumingii (Edaño
255),
'pith', unlignified
bicollateral, smaller
9.
PNH
basal
part.
37179),
—
distal
6.
Axinandra
Dactylocladus stenostachys (Teysmann s.n.),
distal
part.
—
(Strey 7750),
11.
zeylanica (Thwaites 2668),
part. — 8. Crypteronia paniculata
Crypterottia paniculata
var.
central part.
var.
distal
— 10. Rhynchocalyx
affinis(Kostermans 255)
midrib.
midrib.
xylem hatched; phloem dotted; sclerenchyma black; c=collenchyma; h=hypodermis.
part.
—
affinis (Kostermans
—
lawsonioides
12.
Rhyncho-
G.
ral vascular
C.
J.
Vliet & P. Baas:
van
in the distal part
bundle,
forming
sclerenchyma ring, and with
rupted
Ground tissue collenchymatous
side.
parallel
axis of
to
petiole). Crystals
Sclereids in lamina of
tissue
palissade
Node
two
with
present
a
closed ring, sheathed by
latero-dorsal
(material studied
Young twigs
cuticle
12—14
matous
large, little branched,
1—6
square,
mm
older twigs rounded in
thick, with conspicuous
fi m
with
stone
section.
transverse
anticlinal
flanges.
with in each
cells,
pericycle adjacent
in
sclerified in older material.
cells, chambered parenchyma,
corner
Secondary phloem
and (in
elements. Pith
infrequent sclerenchymatous
Crystals
of the twig x—3
Sclereids,
as
druses in
cortex
Wee Lek
1393;
Leaf (Fig. 6,
13,
surface
In
sparsely
A.
alata
Baill.:
Beccari
BORNEO,
Jacobs 5116*; Meijer
d'Alleizette
lamina in A.
SAN
2485; Meijer 492, 555*;
14;
in
on
(
s.tt.
3651*.
=
HB
Usually
p.p.
(Meijer 492) and
slightly undulated
with thin
areas
of
on
cells with thin,
periclinal wall
in adaxial than in abaxial
of them occasionally
13—17-—20fj.m
p.p.
infrequently
and
some
long,
(Chew
a
BORNEO,
BORNEO,
S
15661.
A.
—
zeylanica
Thw.:
beccariana,
in
curved
loops
major
[im
Giant
to
hairs
petiole and
undulated in adaxial
of undulations
wide.
stomata
or
cell
in part of the
Stomata confined
part
to
corners.
material,
abaxial epider-
submersed below the guard cells
subdivided perpendicular
1393).
on
absent. Anticlinal walls of unspecialized
epidermis.
minor or
12—14—17—20
Wee Lek
present and of
secondary
Baill.:
beccariana
abaxial side of lamina in A. alata. Cuticle
to
the
rarely
only distinct
present. Cork
traumatic origin.
veins prominent in A. beccariana and A.
cell pairs
ledges fairly well
present,
infrequently
irregular outline, probably of
Guard
pore.
Outer stomatal
developed. Polar T-pieces absent; cuticular 'cross-pieces'
coriacea
—
straight anticlinal division walls
mis, paracytic, subsidiary cells for
one
cells.
A. coriacea Baill.:
A.
in A.
epidermis;
xi—
section,
indumentum of short unicellular
glabrous, but
in abaxial
and
—
37494*, 49843; Smythies
cells straight
frequently
transverse
strongly sclerified
some
com-
scattered
Thwaites C.P. 2668.
to
more
tubes,
ring; phloem with
oval in
to
8299).
petiole and abaxial side of midrib
zeylanica
Unspecialized
of sieve
infrequently
Thw.
P.B.
indistinct
epidermis,
inter-
first formed
cells with unilateral
continuous
a
coarsely granular, anticlinal flanges
to
more
Plate I, 3, 4; II, 8)
view.
present
an
and pith, abundant in chambered phloem parenchyma.
3423*, 3458*; Jaheri s.n. (1893); Teysmann
CEYLON,
scanty, in
and pith.
see cortex
studied.
P.B.
material)
quadrangular
AXINANDRA
Material
square
composed
older
in older material with
lignified parenchyma,
present
square;
parenchy-
6—11
perivascular sclerencbyma,
to
phloem fibres. Primary xylem and internal phloem
of
prob-
with flattened thin-walled cells. Greater part of primary
alternating
panion
composed
cells
Epidermal
Cortex of
and thin-walled, in later stages layers of
square
thickenings
and
traces
diameter; Plate II, 6)
in
phloem
A.
tissue.
spongy
in the internode below.
cell-layers interspersed
phellem cells
Chew
fusiform
asterosclereids in
± isodiametric
cells of
Cortical bundles departing from lateral
traces.
rupted ring. Cork arising
Beccari
inter-
either
on
druses in ground tissue of laminaand petiole.
less centric cortical bundles (Plate II, 6). Perivascular sclerenchyma
wall
strongly
large sclereids (probably
numerous
as
a
wing bundles
centric
(Fig. 3)
ably ending somewhere
Axis
types:
2—3
181
Anatomy of Crypieroniaceae
(Plate II, 7), and idioblastic sclerified
Trilacunar with three
or
M.
zeylanica,
not
Primary
in
warts
veins
prominent
in
BLUMEA
182
Fig.
&
13
Axitiandra
14.
section of lamina
indicated
other
with
zeylanica (Meyer
dots. — 14.
material.
In
square
to
section
Paradermal
No.
of
1975
lucida
drawings,
and
fairly thin and slightly
13).
Stomata in the
(A. alata)
wavy
Hypodermis
composed
(fig.
thick.
3 —6 /urn
grooved,
with
without
or
chyma which
is
polar
extra
unlignified parenchyma
with ±
Major
cells
to
with
at
basal part of
except in
without sclerenchyma
Crystals abundant
styloids
in
to
petiole of
in A.
tissue
irregular
wall
periclinal
spongy
tissue.
a
flat
without incurved
or
unlignified, parenchy-
sheathed by sclerenchyma
usually bicollateral,
collateral, also usually sheathed by scleren-
the poles except in smallest veins. Outer bundle sheath
variously distinct. All
similar vascular pattern
very
present
A. alata.
prominently raised, supplied with
sclerenchyma, ground
veins
view)
clearly dif-
(Meyer 555), only
subdivided by thin anticlinal and
caps. Minor veins
abundant
more
in surface
layers of palissade cells and unlignified
I —3
sheathed with
system
to
areas
specimens, and absent from
collenchyma
abaxially slightly
collenchymatous.
matous
as
thick-walled
p.p.
arc-shaped, rarely interrupted, bicollateral vascular bundle, with
edges; whole
of
of
130—340/tm
Unspecialized epider-
cells. Adaxial hypodermis
and A. zeylanica
14), cells occasionally
Mesophyll composed of
Midrib adaxially
to
13,
layer
irregularly thickened anticlinal walls (fig.
midrib and major veins of other A. zeylanica
thickenings
walls.
to
the unspecialized
as
ferentiated in A. coriacea, A. beccariana,
over
cuticular
of irregular outline, adaxially larger than abaxially and the
or
level
same
Granular
dorsiventral,
Lamina
former with irregularly thickened outer periclinal walls (cf. thin wall
and
Transverse
x 970. — 13.
hypodermis.
hypodermis.
adaxial cuticle
fim thick,
flattened
2,
epidermis
section.
transverse
1 —4
of
22,
Camera
555).
showing irregular wall thickenings
thick. Abaxial cuticle
mal cells
VOL.
as
A.
midrib, but
veins embedded in
with
alata, sheathed by
extra
various
amounts
alata. Ground tissue parenchymatous
rather infrequent;
as
mesophyll. Petiole
latero-dorsal
to
wing bundles
of sclerenchyma,
collenchymatous.
druses in ground tissue of lamina and petiole
phloem of major vascular bundles. Idioblastic sclereids absent.
G.
Note
J.
C. M.
Vliet & P. Baas:
van
individual
on
in A.
in which
alata,
zeylanica
shows
the
epidermis
beccariana
somewhat
being
are
found
least developed. A.
are
the strongest development of irregular wall thickenings
A. coriacea and A.
epidermis;
species-to-species
generic description. The thinnest leaves
thickenings of
wall
irregular
183
of the restricted
Most
species.
variation in Axinandra is recorded in the
Anatomy of Crypteroniaceae
in
the adaxial
intermediate. A. coriacea has
the thickest leaves.
Node
(Fig. 2)
Complete girdling
plus
trace
one
with the basal part of the median
beccariana
ending
Beccari 3423 and
p.p.
in
of internode
cortex
(material studied
Axis
Twigs rectangular
1—4
one
fusing
in A.
of the successive nodes of A. coriacea SAN 37494),
over
at
least
2—4
sometimes
cells
very
to
square
parenchymatous cell layers,
Perivascular
corners.
infrequent. Cork arising
in
in T.S. with
rectangular
with thin-walled layers. Pericycle
c.
4
thick-walled sclereids
restricted
to
These layers
to
and internal
phloem
partly lignified, with
tous,
bordering
on
inner
Sclereids,
see
a
cortex
and
Material
Anderson
studied.
Poilane
KEP.
11924;
F.
cumingii (Planch.)
37179*.
—
—
C.
28337*;
INDIA,
97773*.
—
7,
8,
surface
to 20 /tm
Endl.:
griffithii
as
stone
long thin styloids
&
3180;
Thomson
paniculata
var.
CELEBES,
Clarke:
of
an outer
with
a
paren-
U-shaped
massive
ring of
Secondary
parenchyma. Primary
to
cells;
oval, parenchyma-
in A.
Crystals
in
outer
zeylanic mainly
present
as
sparse
and internal
to
phloem.
Bl.
b.b. 9704;
MOLUCCAS, Pleyte 382]
BORNEO, Ampuria
Beus.—Osinga:
macrophylla
Hallier
Hooker
C.
C.
Kostermans
SAN
BORNEO,
13012.
—
s.n.*; PHILIPPINES,
affinis Beus.-Osinga;
11;
C.
ak
Banying
paniculata
Edaho PNH
PHILIP-
41449*] MALAYA, Wy art-
Bl.
Nyudong
var.
S
19193*,
paniculata: LAOS,
Kochummen
17897; MALAYA,
MALAYA KEP.
F.
98861*] THAILAND,
Plate I, 5)
view.
Usually glabrous, but indumentum of short unicellular hairs
long) and long uniseriate, multicellular hairs (up
rarely finely granular, adaxially striated
or
en ray
ring. Pith quadrangular
frequent thick-walled
adaxial and abaxial epidermis of C. paniculata
with
perivascular sclerenchyma,
255*.
Leaf(Fig.
In
C.
76336.
& Paie S
Kostermans
(up
F.
centric cortical
pith.
Edaho PNH
Smith KEP.
/urn
thick in
interrupted ring,
phloem, and interspersed with sclereids.
and
I—2
—7 fim
phloem of older twigs. Phloem fibres
outer
CRYPTERONIA
PINES,
an
if present, continuous with
are,
continuous
pith,
5
layers of sclereids
internal phloem. Pith caducous in older twigs.
druses in
frequent
in
rare to
in
cell layers wide and composed
phloem composed of sieve tubes, companion cells, and axialxylem
and
species with
fibres
pericycle adjacent
Cuticle
erect.
unilaterally thickened adaxial walls, alternating
differentiated from
the region of primary
in all
sclerenchyma
chymatous portion, and variously differentiated
adaxial wall thickenings.
to
square
thick in A. beccariana and A. zeylanica,
/im
A. coriacea. Cortex of 5 —9
bundles in the twig
nodes (A. alata, fig. 2b).
two
section. Indumentum of short unicellular
transverse
hairs present except in A. alata. Epidermal cells mainly
thick in A. alata,
anastomoses
diameter)
in
rounded in
to
present,
Cortical bundles mostly present (absent
continuous
or
mm
median bundle. Lateral
trace.
without thin
rather indistinct
to
areas
in
loops
conspicuous,
undulated. Stomata confined
to
or
var.
not,
to
160 /urn
long)
present
on
affinis. Cuticle mostly coarsely granular,
abaxially infrequently striated
of undulations in adaxial
adaxially curved
abaxial epidermis,
to
over
veins,
epidermis. Anticlinal flanges
undulated,
abaxially
straight
to
paracytic, subsidiary cells only for
a
BLUMEA
184
minor part submersed below the
11 —14—19—20 /im
areolae,
in
veins
griffithii,
C.
outer
22, No.
2,
1975
guard cells. Guard cell pairs
stomata
complex
to
anomocytic
distinct
infrequently
Polar
cyclocytic.
stomatal
12—14
—19 —22
faint
T-pieces
well developed.
ledges
prominent
In
on
veins,
over
present
jum
more
long,
rarely
in
absent, but fairly
to
Cork
and of irregular outline, probably of traumatic origin.
present
nent.
wide. Giant
VOL.
warts
infrequently
Primary and secondary
both surfaces; elaborate network of minor veins sometimes promisection. Lamina dorsiventral, no—300
thick. Adaxial
fim
transverse
cuticle 2—6 .Mm thick, abaxial cuticle 1—3
thick. Unspecialized cells mainly flattened,
//in
than
adaxially larger
of
hypodermis
in
from
paniculata
C.
Stomata in the
abaxially.
paniculata
var.
flattened
more
partly lignified,
or
spongy tissue,
square
hgnified
not
less
triangular bicollateral
differentiated
& Thomson
s.n.),
absent
flattened, larger than adaxial
cells, and rather
compact,
paniculata. Midrib adaxially
in C.
vascular system,
not
present,
of palissade
layers
I—3
to
supplied with
slightly raised, abaxially prominently raised,
to
or
cells
cells. Adaxial
unspecialized
as
paniculatap.p. (Hooker
affinis. Hypodermal
var.
epidermal cells. Mesophyll composed of
wholly
level
same
layers of usually thick-walled lignified cells
of lamina of C.
parts
some
I —2
frequently
usually closed,
a
of distinct
composed
vascular bundles separated by lignified 'pith'—'cortex' connections, with small additional
bundles in
medullary
wholly
tissue
or
phloem. Major
macrophylla; whole
C.
veins
sheathed by sclerenchyma.
system
sometimes sclerified in
partly lignified,
bicollateral,
'pith' region adjacent
minor veins collateral. Most veins with
an
many-layered sclerenchymatous bundle sheath and/or polar sclerenchyma
and
lignified
a
little
or
unlignified parenchymatous
or
sclerenchyma.
no
sclerenchymatous
to
Major
veins
in basal part, and
centric
to
wing bundles
midrib.
C.
parenchymatous
tissue of lamina and
as
unlignified
or
C.
two to
Large unbranched sclereids
petiole, and
infrequently
not
several latero-dorsal bicollateral
collenchymatous.
to
paniculata
midrib, often
styloids
as
present
in
phloem
ground
in
amounts
Crystals
resent
P
of
of
as
petiole and
tissue of
petiole of
cumingii, rarely clustered.
Note
on
Most of the variation from
individual species.
in Crypteronia is recorded in the
ally thin, and the lesser
amount
morphic
in
Crypteronia.
C.
of lignification in the supporting
paniculata
vascular tissue in the petiole with
a
not
var.
affinis has
a
genus
as
slightly aberrant
species
typic-
the
most
meso-
arrangement
of
entirely closed, deeply arc-shaped bicollateral bun-
(e.g. cuticular striations, conspicuousness
of anticlinal wall undulations etc.)
to
are
tissues around the veins
dle with strongly incurved edges (fig. 11). Most of the other characters
within the
species
generic description. The leaves of C. paniculata
and in the ground tissue of petiole and midrib characterizes this species
as
in
except
either side. Vascular system sheathed by various
tissue
sclerenchyma. Ground
druses in ground
on
one- to
girders,
circular in distal part, less frequently
to
connections; with
fragmented by 'pith'—'cortex'
caps or
by lignified
similar vascular pattern
kidney shaped
internal
inner
bundle sheath. Smallest veins with
transcurrent
bundle sheath extensions,
parenchymatous
which has embedded veins. Petiole with ±
entirely closed
outer
vertically
Ground
to
appear
to
of thin
be variable within
reported
to
vary
areas
of cuticle in loops
some
individual species
well.
Node (Fig.
Typically
without
fusion
or
1)
common
common
not.
gaps
gaps
Lateral
with
split laterals and
but with complete
anastomoses
C.
paniculatap.p., but entering
in
C.
present
one
girdling
p.p.,
traces,
between lateral
the stele in individual
cumingii and C. paniculata
median bundle. In C.
after having
gaps
showing
traces
paniculata
distinct places
of
and median bundle in
below the level of leaf insertion
traversed the
cortex.
G.
Axis
C. M.
J.
(material studied
Twigs rounded
in
2
—
4.
P. Baas:
of 5 —8
poorly developed (var. paniculata)
adjacent
affinis. Epidermal cells
var.
parenchymatous
(see under node). Perivascular fibres
species
glabrous, but with indumentum of
Mostly
paniculata
without cortical bundles except
griffithii,
or
in
a
with adaxial unilateral wall
in C.
thickenings;
continuous ring,
is
present). Cork
paniculata
with thin-walled
ally thick-walled cells alternating
cell-layers,
insertion in
but in
and C.
cells
paniculata
C.
pericycle
outer
square
some
flattened,
to
griffithii layers of unilater-
in other
cells;
with
interspersed
just below leaf
in-
flattened,
to
square
absent (var. affinis). Cork arising in
perivascular sclerenchyma (if the latter
to
185
Anatomy of Crypteroniaceae
diameter; Plate II, 9)
in
section.
Cortex
thick.
fim
sclereids in C.
in C.
&
Vliet
1.5 —4 mm
transverse
frequent unicellular hairs
cuticle
van
thin-walled
species
cells irregularly distributed between unilaterally thick-walled cork cells. Secondary phloem
of sieve
composed
tubes,
companion
Primary xylem and internal phloem
by hgnified
In
'Hartbastbiindel',
and pith,
griffithii whole
C.
31). Pith caducous
1911:
styloids
as
outer
in
studied.
D.
stenostachys
composed of
frequent
near
Crystals
present
druses in
as
and pith.
see cortex
Oliv.
Anderson
BORNEO,
most
strongly sclerified (Hallier's
in older material.
phloem. Sclereids,
Oliv.:
the latter
of pith
part
in T.S.,
rectangular
to
DACTYLOCLADUS
Material
fibres.
phloem
less continuous ring, sometimes traversed
parenchyma interspersed with sclereids;
internal phloem.
and frequent
cells, parenchyma,
a more or
connections. Pith oval
pith—xylem
mostly Hgnified
cortex
in
b.b.
8541;
32378*; Teysmann
s.n.
(
=
HB
7930).
Leaf(Fig.
Plate I,
9;
surface
In
conspicuously
straight
to
—
19
21
warts
some
of irregular
7 —10 /um
erect,
unspecialized
cells, and
outline and
taller than the
spongy
tissue which is
mesophyll. Vascular
veins
of traumatic
to
square
erect
fx m
coarse
flanges
which
21 —24—29
ledges
distinct,
not
rarely
are
/urn
for
a
long,
well-developed.
origin frequent
Veins
or
thick.
on
abaxial
prominent, except for
cells
Stomata in the
of
2—3
in central part. Midrib
parenchymatous
same
level
as
adaxially flattened
a
flatter adaxial bicollateral bundle;
tissue
Adaxial
of adaxial
layers of palissade
prominently raised, supplied with
a
thick.
330—450/nm
Unspecialized
abaxial cells.
faintly lignified
to
shallowly
arc-
whole system
collenchymatous,
bicollateral, with unlignified parenchymatous bundle sheath
Minor veins with little
or no
sclerenchyma. All
distal end similar
system of petiole in
latero-dorsal wing bundles
as
s.n.
Mesophyll composed
on
to
crystal sand (fragmented
as
veins embedded
that of midrib, but with
either side; in basal part with
almost closed bicollateral bundle. Crystals present
petiole,
cells,
(subsidiary)
dorsiventral,
2—5
by interrupted fibre ring. Ground
enclosing polar fibre caps.
2
Anticlinal
Outer stomatal
Teysmann
Lamina
cells. Hypodermis absent.
partly lignified. Major
i or
in
abaxial cuticle
abaxial bicollateral bundle and
sheathed
faintly granular; abaxially
very
s.n.
section.
thick,
probably
infrequent
slightly raised, abaxially faintly
shaped
in
neighbouring
T-pieces indistinct.
primary veins in Teysmann
epidermis
to
smooth.
to
abaxial epidermis, typically anomocytic,
to
with 4—7
wide. Polar
transverse
cuticle
or
cyclocytic,
of Anderson 8541,
epidermis
In
/urn
—
Cork
to
Stomata confined
smooth
striated
only submersed below the guard cells. Guard cell pairs
minor part
16
Glabrous. Cuticle
striated, adaxially faintly
curved.
anomocytic
1)
view.
a
strongly incurved,
druses in ground tissue of lamina and
druses?)
in
sclereids present in ground tissue of lamina and petiole.
phloem. Thick-walled fusiform
BLUMEA
186
of the vascular bundles of the veins.
caps
than the sclereids
type
Node (cf. Fig.
are
They
2, 1975
connected with the polar sclerentherefore of an entirely different
are
of Alzatea.
1a).
Common gaps with
Axis
No.
22,
The fusiform sclereids in the mesophyll
Note.
chyma
VOL.
(material studied
Young twigs
laterals and
split
1.5 —3
more
Epidermal cells mainly
in
mm
less
or
square,
median
one
diameter)
twigs rounded
older
rectangular,
cuticle
trace.
8—10
fim
in
section.
transverse
thick. Cork arising in subepidermal layer.
in older
Cork cells flattened with lignified anticlinal and inner periclinal wall thickenings;
twigs
2—3
layers of thickened cells alternating with
I
Cortex
layer of thin-walled cells.
of 7 —9 parenchymatous cell-layers, interspersed with branched and unbranched sclereids.
Perivascular
closed and
sclerenchyma
cylinder
not
composed of thick-walled
poorly
or
of sieve tubes, companion cells, and parenchyma,
older material. Primary xylem in
less
or
discrete units opposite
and
fusiform sclereids,
material.
Crystals
present
infrequent druses. Sclereids,
Note.
The
and with
phloem
see cortex
and
their
twigs, almost
infrequent scattered fibres
to
rectangular
periphery. Pith caducous
at
in
know whether they
merely
other by phloem parenchyma,
separated by pith parenchyma
are
difficult
part of the internal
the
been traced because of difficulties in
correct
to
distinguish
in the
Leaf(Fig.
In
R. lawsonioides
from the internal
pith. Apparently they
view.
granular, and striated
occasionally
sidiary cells,
identifying
In
Oliv.: S.
Anticlinal
veins.
to
abaxial
seemingly
long,
13—15
T-pieces absent. Cork
probably of
AFRICA. De
course
possible
in the nodal region has
not
Oliv.
Joncheere s.n. (1973); Strey 7550*, 7565,
traumatic
transverse
7750.
—16—18
warts
(see
cells. Unspecialized
cells
square
to
abaxially. Adaxial hypodermis of
i
note),
for
/im
a
4—7
Outer
present
of primary and
2—4
with
to
to
finely
slightly
stomatal
ledges
well-
and of irregular outline,
secondary
dorsiventral,
and
neighbouring/sub-
only. Guard cell pairs
minor part
wide.
infrequently
Lamina
cuticle 3 —7 [I m thick, abaxial cuticle
straight
epidermis, intermediate between anomocytic
paracytic
origin. Network
section.
striated, abaxially smooth
flanges well-developed,
submersed below the guard cells
16—17—20—22 fim
developed.
not
separated from each
them in herbarium material.
Glabrous. Cuticle adaxially
over
Stomata confined
cyclocytic,
system,
12)
10,
surface
curved.
phloem
Their
one.
RHYNCHOCALYX
Material studied.
as
whether the interpretation of them as medullary strands
or
is
pith
pith.
predominantly peripheral position
are
in
in older
and
cortex
fuse occasionally with the internal phloem. Without ontogenetic studies it is
to
in
more
with thick-walled
interspersed
druses and crystal sand,
as
medullary phloem strands
phloem because of
young
major primary xylem poles. Pith oval
medullary phloem bundles
in
in
Secondary phloem composed
less closed cylinder. Internal phloem in
a more or
composed of unlignified parenchyma
section,
transverse
developed
fibres in older material.
veins
prominent.
200—270[im
thick.
Adaxial
thick. Stomata above level of unspecialized
/mi
flattened, adaxially lower, but slightly broader than
collenchymatous
cells larger'than those of adaxial epidermis.
to
parenchymatous
Mesophyll composed of
2
layer
present;
layers of palissade
G.
and
cells,
unlignified
in
with little
M.
with
most
adaxial sclerenchyma
an
tissue
minor veins
sheath
unlignified bundle
parenchymatous
a
sclerenchyma girder, abaxially
bicollateral, with
extending
girder; smallest
and
petiole,
as
solitary prismatic crystals
in
sclereids frequent in ground tissue of petiole,
to
a
veins embedded in
as
collenchymatous.
collenchymatous
and lower
to upper
slightly incurved single bicollateral vascular bundle,
a
187
abaxially prominently
parenchymatous
without latero-dorsal wing bundles. Crystals present
and
flattened,
adaxial hypodermis by
to
sclerenchyma (fig. 12). Ground
no
Anatomy of Crypteroniaceae
Midrib adaxially
tissue.
Vascular bundles of major and
enclosing
& P. Baas:
single, crescentiform, bicollateral vascular bundle (with incurved
a
Strey 7550), linked
or
Vliet
van
spongy
raised, supplied with
edges
C.
J.
to
epidermis, and
mesophyll. Petiole
sheathed by collenchyma,
druses in ground tissue of lamina
phloem. Thick-walled (probably fusiform)
infrequent
in
tissue of midrib
ground
and
lamina.
Note. In the
seemingly paracytic
cells
cytic subsidiary/neighbouring
Node
(material studied
/«m
cortex
1
to
cells of the ring of anomocytic
arranged parallel
to
thick.
layer
1—4
in
mm
Cortex of 8—12
from the
Plate II,
diameter;
section.
transverse
to
cyclo-
pore.
10)
Epidermal cells flattened, early caducous,
cell
parenchymatous
periphery, composed of layers
layers. Cork arising
of thin-walled cells,
3rd
in
cuticle
to
5th
alternating with
several layers of flattened cells widi unilateral adaxial wall-thickenings. Perivascular
sclerenchyma virtually absent
in
cells, chambered parenchyma
and internal
phloem
in
crystals
Systematic
and
as
present
cortex,
10). Sclereids,
cells,
and
infrequent thick-walled sclereids. Primary xylem
transverse
see
as
clusters
and
at
the
one wants
solitary prismatic crystals
diagnostic
in chambered
is often much
at
more
the
difficult
are
usually
of good
diagnostic
level within the Crypteroniaceae.
genus
evaluate and depends
to
on
the
Their
problems
solve. For instance, the stomatal type, in combination with several other
to
of a close
affinity between Axinandra
but its taxonomic value becomes of little help
Melastomataceae which both
indicate relationships
are
in
comparisons
in this respect.
heterogeneous
also show this
feature,
&
for
Chalk,
and
which is of
1950).
very
rare
occurrence
In view of this
the systematic anatomist is
to
some
Styloid crystals
list
as
many
judge mutual affinities and relationships with other families
with
also
very
serve
strong
representatives of this family
in the
variability
Crypteronia,
of Crypteroniaceae
between Axinandra and Crypteronia, but provide
evidence of Melastomataceous affinities too, because
possibility
phloem parenchyma
value of some of the anatomical characters
characters, provides good evidence
(cf. Metcalfe
in
pith.
species and often also
systematic value
of
(sometimes irregular) solitary prismatic
The anatomical characters recorded in the descriptive part
value
section, composed
interspersed widi infrequent thick-walled sclereids, caducous
Crystals
pith and
in
(Plate II,
thick-
very
Secondary phloem composed of sieve tubes, companion
continuous cylinder. Pith oval in
a
unlignified parenchyma
older material.
twigs, interrupted and composed of
young
walled sclereids in older material.
to
the
trace.
one
Twigs rounded in
i—2
stomata 2
(Fig. 4)
Unilacunar with
Axis
are
in
Dicotyledons
systematic
characters
on
as
as
value,
a
whole
the
only
possible and
to
the basis of a good overall
VOL.
BLUMEA
188
22,
No.
2, 1975
1
Melastomatoideae
Astronioideae
Memecyloideae
Melastomataceae
Lafoensia
0
0
—
X
0
—
0
X
0
0
—
X
0
0
X
—
0
0
0
—
X
X
—
x
X
—
—
0
0
0
0
0
0
0
0
0
—
—
—
—
—
X
X
—
—
—
X
—
—
0
0
—
—
0
0
—
—
+
—
—
+
—
—
—
X
X
—
—
—
0
)
7
relatives
Lagerstroemia
Lythraceae
puta ive
Sonneratiaceae
and
Oliniaceae
—
X
0
0
l
)
+
—
X
—
+
?
+
+
+
X
—
X
—
0
0
0
—
+
S
O
O
O
D
—
+ D
O
0 S
O
+
O
)
e
Crypteoniac e
of
charcters
Rhynchocalyx
s
r
|
oniaceae
>OG 11?
Cryptei
i
I
X
—
X
X
—
X
—
X
X
—
X
X
—
X
—
X
—
X
—
X
—
X
X
—
(X)
X
—
X
—
X
—
—
X
X
X
—
—
X
X
—
—
X
—
X
—
X
—
X
X
X
—
—
X
—
—
X
—
—
—
X
—
—
—
X
—
X
—
—
—
X
X
—
X
—
)
)
5
"
fr
—
8
)
8
Alzatea
X
—
—
X
X
X
antomical
Some
I.
Table
)
)
8
cy locytic)
cluster d
(to
granularsmo th par cytic anom cyti
Leaf
Cuticle Stomat
4
A B
presentabsent
petiol-midrbpetiol-midrb transcurent phloem
system
Hypodermis
Vascular
sand
prismatic
cluster d crystal solitary styloids
in
scler ids
outside
phloemphloem phloem phloemscler ids foliar
Veins Crystals
Unbrached Branched
verticaly
in in
in in
subepidrmalpericycle cortex
petiole
in in
origin,
cork
of
Axis
Site
G.
C. M.
J.
van
Vlibt
& P. Baas:
Anatomy of Crypteroniaceae
189
)
18
0
0
0
0
+
(0)
+
—
+
(0)
0
0
0
X
—
—
+
0
X
0
—
o
—
—
4-
—
0
—
0
0
—
0
0
0
—
0
(0)
0
—
(0)
0
—
0
0
0
0
—
0
0
0
X
—
—
—
X
—
—
—
—
—
—
—
—
+
—
midrib
and
(X)
—
fibres
in
)
16
x")
—
X
X
—
0
0
0
+ +
p>
D
X
—
-
X
or
cells
0
s
0
+
D
0
—
0
—
0
-16)
parenchyma
)
14
0
—
+ +
—
+
—
+
—
X
X
—
(X)
—
X
—
X
—
in
oc ur
)
10
X
—
—
X
—
—
X
—
—
X
X
—
—
X
—
—
—
X
—
X
—
—
X
X
—
X
—
—
—
(X)
X
—
—
X
X
—
—
X
—
present. only.
)
12
crystals
petiole
of
)=rarelyDuab nga
only.
chamber d
part
)
12
—
X
X
—
—
X
—
—
—
X
—
X
—
—
X
X
X
—
(X)
—
—
—
—
X
—
family
the
for
recorde . recorded
of
part
in
present
alternate
o=charcter
pits
1
2
Vevsl—ray
3
parenchyma
type type type
)
9
1975)pits,
Vliet,
interv s el
(Van
Wood Vesturing
larger
Unes
short
scal rifom,border d border d
to
and
reticulate
distinctly
or
to
pits
Fibre
present
)
minutely septate
simple
Fibres
crystals
vasicentric ag regates with
15
III
I
rays
in
aParenchyma liform-cnluet partacheal HetrogenousHetrogenous Intercluar
partly
distal
(
—
in
dif use
fibres
SeptateRays:
canals
—
or
species
and
not
or
absent
char cter
D=
only;
=char te
char cter. (sub)family.
genus
midrib
for
petiole
veins
largest
closed
whether
complet ly
descriptons
in
edges
or
Son erati incurved adaxial Crypteronia
.
for
midrib
over
recorde only
X
-f
*) *)
studied
by bv
and
without
or
with
and
in
abaxial
an
of
S
=
paniculat scler nchymacolenchvma
part
charcteisc =charcter hypodermisarc-shaped, composed
=
the
in
transcurent transcurent
transcurent adaxialyadaxialv
not
also also
veins midrib
4
5
8
material
Crypteronia
and
twig
present
in
only in
than
larger
wings
reticulate, narow
scanty-prachel
very
•)
8
Mujica
from
evident
to
Alzatea
)
homgenous
and
parenchyma
of
7
III.
to
than
Crypteronia Axina dra
in
vasicentric Cutler's
midrib only smal er alternate with
))
))
)
3
cumingi
was
Rhyncoalyx
not
is
(1950) are
rays
Kribs
to
the
taxa
ac ording these
in
it
)
))
)
)))
10
n
12
13
14
15
ie
BLUMEA
190
similarity
of consistent differences in
or
table I for the
5
Crypteroniaceous
anatomy have been included
Of the
characters
of
are
No.
22,
2,
1975
number of characters. This has been done in
a
and several putative relatives. Data
genera
well and
as
those
listed,
the cuticle
probably also
they
VOL.
are
referring
taken from Van
Vliet,
and
little value for comparison with other families, yet
very
included because within the Crypteroniaceae they have
are
1975.
unbranched sclereids,
hypodermis,
to
wood
on
good diagnostic value.
a
The systematic value of the wood anatomical characters is discussed elsewhere (Van Vliet,
1
975).The implications of the diversity
We have found
reasons
no
is due
systematic discussions
to
Within the
genus
within the
species),
a
there is
Crypteronia
into
subdivision also
some
adopted by
mesophyll and
not
studied.
taxa
and
paniculata)
Beusekom-Osinga
Van
192.
in the
anatomical diversity supporting Niedenzu's
Crypteronia paniculata is the only species with
veins embedded in the
on p.
Crypteroniaceae
sections, Crypteronia (C.
two
discussed separately
are
different ecologies of the
Relationships
(1892) subdivision
in nodal anatomy
believe that the anatomical diversity employed
to
a
&
complete girdling
Basisperma (the
other
Van Beusekom (1975).
trace
in the
node,
with
transcurrent, and abundant coalescent
vertically
apertures in the secondary xylem vessels of the twigs (Van Vliet, 1975).
The
subgeneric division of Axinandra
ca) and
section Naxiandra
Axinandra alata of the latter section is
of the
by
genus
rather
section Axinandra
monotypic
not
anatomical characters.
styloid crystals,
different from the remainder
is
descriptions
it
Particularly
the
petiole, total absence of a
epidermal cell walls.
appears
that Axinandra and Crypteronia share
common
occurrence
found elsewhere in the Crypteroniaceae,
not
Supporting evidence
of the
(A. zeylani-
correlated with anatomical differences.
anatomically slightly
regular outline
From table I and from the
and
is
its lack of wing bundles in the basal part of the
hypodermis, and
many
into the
(the other species)
in nodal
provided by the overlap
of paracytic
suggest
stomata
close affinities.
conditions, and similarities
in
cuticular texture, cork origin, and overall wood anatomy. Differences in petiole anatomy
and parenchyma distribution of the wood
thickening of the cell walls
throughout the
constant
of
are
minor
The peculiar collenchymatous
ones.
epidermis and/or hypodermis
in
species
and does therefore not provide
genus,
a
of Axinandra is
not
good distinctive char-
acter.
Dactylocladus
shares its
nodal type with Crypteronia, but differs from both Axinandra
and Crypteronia in stomatal type,
and
ray
type. These differences
with either
by
Van
Crypteronia
cladus into one
and
or
Beusekom-Osinga
tribe,
or
by
crystal complement,
are not
Axinandra
&
to
be able
Van Beusekom
Muller
vessel—ray pitting,
to
advocate
(1975)
who
(1975) who reported
a
a
strong
affinity
as
suggested
placed Crypteronia and Dactylo-
similar pollen type for Axinandra
Dactylocladus.
Alzatea differs strongly
Vhet,
1975)
With its
genus
from the above mentioned
wood anatomy (Van
genera in its
and also bears little
sclereids,
stands
out
clustered
resemblance
crystals
ence
from all other Crypteroniaceae,
these in its other anatomical characters.
phloem cells, and trilacunar
though
it is
differs, however,
genus
the characters restricted
with Alzatea, differs from it
to
in cork
as
or
to
vice
node the
derive the nodal
versa.
in Axinandra.
The
From
origin
sharing
and
ray
the
same
pres-
Crypter-
in stomatal type and septate fibres
Alzatea. Dactylocladus,
moreover
possible
from that in Alzatea
of cortical bundles might indicate similar tendencies
onia and Axinandra the
in
to
in chambered
condition in Crypteronia and Dactylocladus
as
cork origin,
sufficiently outweighed by anatomical similarities
as
well
stomatal type
few similarities
type. The
G.
J.
C. M.
van
remaining (see table I) do not
Vliet
& P. Baas:
Anatomy of Crypteroniaceae
compensate for the above mentioned
that relationships of Alzatea with the other three
is also
Rhynchocalyx
aberrant in this
very
is
genera
191
differences, indicating
at most remote.
of genera; its wood anatomy shows
group
similarities with that of Alzatea (Van Vliet, 1975) but is very different from the other three
genera.
Within Crypteroniaceae it is
crystalliferous fibres
in the
If
phloem.
in the
the only
to
genus
wood, cork arising
with each of the other four
compared
Alzatea (stomatal type, and overall wood histology)
in cuticular texture, mechanical support of veins,
besides the important characters restricted
in
common
with each of these
Van Beusekom's
Accepting
heterogeneous
of
group
genera
petiole
anatomy,
of these
relationships
three
It is
possible
with Dactylocladus
genera
weighted
more
evidence
at
calyx and Alzatea
belong
appear
to
to
with
Differences
the number of char-
to
faced with
anatomist is
enough
advocate
only
supports
this is
not
some remote
very
a
close
supported
anatomical
if the anatomical characters shared
by the
Alzatea would
and anatomical characters
with Axinandra,
a
provide
Crypteronia, and Dactylo-
the Myrtales. The anatomical affinities between Rhyncho-
be of the
same
both Crypteronia and Axinandra, and
Alzatea and
are
vessel—ray pitting,
heavily than the differentiating characters.
all of affinities of Rhynchocalyx
except that all
cladus,
and
Rhynchocalyx.
in which the anatomical evidence
palynology.
similarities
being extremely limited (see table I).
be much further removed from this assemblage,
even
no
genera are
or
outspoken,
are most
family delimitation the
genera,
or
solitary prismatic crystals
most
genera,
affinity between Axinandra and Crypteronia. Remarkably
by macromorphology
simple unilacunar node,
a
but considerable differences remain
either Alzatea
to
with Crypteronia, Axinandra, and Dactylocladus
acters
have
in the cortex, and
may
remoteness
be taken
Rhynchocalyx have been raised by
as
to
those between Dactylocladus and
support
the tribe level
Beusekom-Osinga
Van
&
to
which
Van Beusekom
(i975)As
an
alternative one
Van Beusekom's concept of the
might reject
search for the closest relatives of each individual
will be
explored below.
Position in the
As stated
Vliet, 1975)
are
in the
and
Myrtales.
many more
in the past
in twig and
characters in each of the
families
(see
are not
table I).
as
leaf, characteristic for
genera
literature (see comprehensive
survey
to
those families of the
by
with that of Olinia
have been included in the Lythraceae
as
Chalk,
most
summarized in table I
1950;
relevant
Mujica
genera
&
are
Cutler,
at some
mainly based
stage
on
was
1975).
is
by
or
other
(Van
Myrtales,
one
or
families
genera
in the systematic
issue). These families
noted in
a
a
are
striking similarity
superficial
survey
Both Oliniaceae and Sonneratia-
of botanical history. The compari&
1974) but additional observations have been made of
using the Rijksherbarium
no means
most
data from the literature (Metcalfe
slide collection. It should be borne in
mind that the body of anatomical information, although
compared,
of the
Myrtales which have been
Van Beusekom in this
of the anatomy of all Myrtalean families (Van Vliet,
sons
Cunoniaceae
Melastomataceae. Oliniaceae have been added because
between the wood of Rhynchocalyx
ceae
genera
witnessing affinities with
Affinities with
close relatives of the individual Crypteroniaceous
Lythraceae and
all
in the wood
pits
supported by anatomical evidence.
The comparisons have been limited
suggested
supports inclusion of
Besides the presence of vestured
intraxylary phloem
several Myrtalean
suggested
and affinities with other families
Myrtales
before, the anatomical evidence fully
Crypteroniaceae
there
genus
Crypteroniaceae and
in other families. This alternative
very
substantial for the families
complete, rendering conclusions about the lack of affinities
BLUMEA
192
VOL.
No.
22,
2,
1975
with certain groups tentative. For Lythraceae separate additional columns for Lagerstroemia
(Lagerstroemieae) and Lafoensia (Diplusodontineae)
given because of suggestions of close
are
affinities with Rhynchocalyx and Alzatea respectively (see
The
two
of Sonneratiaceae,
genera
those characters in which
which have
they differ or
are
as
ed; for the other anatomical characters data in the literature
in table I, the
as
based
the overall
Beusekom, 1975).
in
one
of the
wood anatomy is
insufficient
were
vegetative
genera.
concern-
do this.
to
anatomy, summarized
anatomy will be dealt with.
implications of nodal
Comparative nodal
on
Van
presented individually for
only been studied
For Melastomataceae separate columns have been added as far
Before discussing the affinities
by
survey
Duabanga and Sonneratia,
anatomy
The nodal anatomy of the families
compared
only
is
incompletely known.
In
the
literature Sinnott (1914) andLignier (1886,1887) provide useful information for Lythraceae
and Melastomataceae,
provided
Lythraceae
with
far
so
one trace
show
or
for
these families Dr.
yet
are
unilacunar with
common
to
some
(Oliniaceae)
node
one
the nodes of
comparing
of these
range
Rhynchocalyx
—
indicate which
presence
common
than
affinity rather
In
Sonneratia caseolaris
a
Chalk, 1950).
All nodes
as
appeared
to our
—
very
families compared,
at
bf
to
report
o
components of the
those of the
with Melastomataceae, which
obviously
known
Oe
and Olinia
present.
unusual
and
nodes;
family
the unilacunar
the situation reported
for
Therefore, general nodal anatomy
two
genera.
However, the
median
condition would favour
trace
a
Melastomataceous
family of disputed Myrtalean affinity,
the nodes
are
also character-
(Howard, 1970). Evidence from other anatomical features (notably
the unvestured pits and
against
&
(Sonneratiaceae)
family of the Myrtales”.
have the closest affinities with these
plus
gap
common gaps
however,
far
a
species (fig. 2). Cortical and
relationships with the other families mentioned.
Rhizophoraceae,
ized by
'derived from cortical bundles',
bundles in Alzatea and the suggested similarity of its trilacunar node
of cortical
with the
in all
as
All
unilacunar
Howard furthermore found
trace.
"(the diversity of) the 'new'
are
genera.
are
Axinandra, Crypteronia, and Dactylocladus with
occurs
taxa
Melastomataceae
Axinandra
elsewhere in the Dicots
Alzatea is unique in Myrtales
cannot
median
our
(Arnold Arboretum)
cite Dr. Howard's letter in response
to
above mentioned families the affinities
node of
trace.
is broader than I have found in
any
Crypteroniaceae
the
a
Howard
sectioned for comparison.
was
be interesting
may
one
A.
substantial number of
a
number of genera (cf. Metcalfe
a
the nodal diversity in Crypteroniaceae:
covers
of
species Duabanga moluccana and
unilacunar. It
In
with
gap
R.
on
in Rhexia and Arthrostemma
present in
are
data
unpublished
comparable
medullary bundles
cymosa
a
traces
perhaps
each of the
as
studied
complete girdling
situation
and
with his
us
the absence of intraxylary
close affinities of Rhizophoraceae
phloem
in
Rhizophoraceae)
with those Crypteroniaceae
pleads,
showing
similar nodes.
Comparative
Alzatea
overall
differs
from
vegetative
anatomy
in
the Lythraceae
which it
sclereids, and
provides
ray
type. The
common occurrence
another distinctive
Lythraceae.
Stant
Alzatea, but
same
applies
acters
there
we
to
are
character
(in Lourteig,
1965)
have been unable
secretory
sacs
to
for
most
of
in
included previously,
was
of the vascular tissue in petiole and midrib,
anatomy, arrangement
mucilage cells
though
reported mucilage
not
all
in the leaf
nodal
of foliar
presence
epidermis
representatives of the
cells in the palissade
find them in material of the
same
tissue
of
collection. The
she mentioned for the petiole. In several anatomical char-
suggestions of affinity (wood
anatomy,
stomatal type, etc.,
see
table I).
G.
Diplusodontineae
C.
J.
applies
more
of about the
are
the basis of
on
would be
unjustified
particular
share
to
appear
193
features with
Crypteronia, and Dactylocladus with
Axinandra,
same
use
with
this
we
with Alzatea. The
different
over
those with these three families. As
as
in
range
as
same
features,
have been unable
to
find
except
a
very
one
affinity
the latter
of Alzatea with
in the Melastoma-
being haphazardly
of Melastomataceae. More detailed and
anatomical studies in Melastomataceae would be necessary
to
conse-
for its trilacunar node. It
close
representative
Alzatea characters,
most
genera
a
Alzatea would fit into this
Melastomataceae,
absolute evidence of
the combination of
distributed
order
of its anatomical
most
to
Melastomataceae, because
taceae
in
Anatomy ofCrypteroniaceae
substantial than those of Lythraceae
of the wide anatomical
quence
& P. Baas:
Oliniaceae and Sonneratiaceae. The affinities and differences between Alzatea and
Rhynchocalyx
family
Vlibt
the differences of
however,
are,
to
van
and Diplusodon)
(Lafoensia
Alzatea. Anatomically
Alzatea
M.
comprehensive
the anatomical affinities
judge
with Alzatea.
With the evidence available it is impossible for the anatomist
systematic affinity of Alzatea. The
contribute much
to
support the isolated relic character of this genus, also deduced from its
type
(Muller,
shares
heterogeneous
the Lythraceae it is,
characters in
one
a
of Rhynchocalyx
however, impossible
representative.
than
is
veins
are
there
the other
to
identical
to
constitute the
genera
The
that of Rhynchocalyx.
origin, the differences
minor importance.
are
similarities
crystalliferous fibres
Affinities of
only
major difference
with Oliniaceae
in
in the wood.
differences
having large mucilage cells
Similarly
;
they
also
(see Metcalfe
&
characters. Punicaceae
have
may
a
p.p.,
Chalk, 1950). Summarizing, Rhynchocalyx
Melastomataceae
p.p.,
Oliniaceae,
those with the above mentioned families
traditional placement of Rhynchocalyx
in the
solution by the anatomical evidence
at
a
on
table I;
A
com-
Sonneratiaceae
or
it
here.
presented
seems
other characters
shows
p.p.,
even more
Lythraceae,
to
large portion of the
Punicaceae
than
same
remote.
In view of the
appropriate
Sprague
in
affinities
of about the
and
are
more
to
&
to
support
Metcalfe (1937)
similar conclusion.
Dactylocladus
and Oliniaceae,
distinctive
(see
mesophyll).
some
Axinandra, Crypteronia, and Dactylocladus. Affinities with Alzatea
arrived
based
also be mentioned in relation
crystalliferous fibres, and have
Lythraceae
this
as
however, impossible
with Alzatea it is,
to
as
in overall
differentiating anatomical character:
one
as
present
are
in the
to
order
to
in mechanical support of the leaf
Rhynchocalyx
but here substantial
too,
moreover
combination of Rhynchocalyx
Rhynchocalyx
be closer
to
appear
The wood anatomy of Olinia
individual member of the Melastomataceae showing
an
common
Rhynchocalyx
of the Crypteroniaceae.
with Melastomataceae results in only
indicate
and Lawsonia from the
Lagerstroemieae,
Lagerstroemia
therefore also closer than those with other Crypteroniaceae. With Sonneratia-
Sonneratiaceae differ
parison
only (gradual) difference. Within
find the complete combination of Rhynchocalyx
from the Diplusodontineae share a fair number of characters
cork
anatomy is in
being of
anatomy
ceae
less
or
more
Both
to
Thus the anatomical affinities of
with Rhynchocalyx.
vegetative
to
unspecialized pollen
considerable number of anatomical features with Lythraceae. The
rays
and Lafoensia and Physocalymma
Lythraceae
the
1975).
Rhynchocalyx
more
to
of links with several families may be used
presence
shows several anatomical features
not
present in
Lythraceae, Sonneratiaceae,
the nodal anatomy and wood anatomy providing
characters.
similarities (table I)
are
with Axinandra and
Affinities with
of the
same
Crypteronia
these
order
are
as
families,
suggested
by
the
significant
most
certain
anatomical
those with Alzatea and Rhynchocalyx.
Affinities
somewhat closer. Melastomataceae show all characters
BLUMEA
194
family
with this
both
is,
1975
in the
phloem.
moreover,
strands of Dactylocladus
relationship
An intimate
supported by the unusual nodal
The medullary phloem
taxa.
22, No. 2,
crystal sand
except the
Dactylocladus,
present in
VOL.
occurring
in
be suggestive
of
anatomy
would also
Melastomataceous affinities. Thus, the results from anatomy support the traditional place-
challanged
Axinandra is
cladus,
as
a
more
and has
an
its inclusion in
Melastomataceae, though
of Dactylocladus in the
ment
deae could be
on
subfamily Memecyloi-
wood and nodal anatomical grounds (cf Van Vliet,
different from Lythraceae, Sonneratiaceae, and Oliniaceae than Dactylo-
additional Melastomataceous character of rare
occurrence
whole: styloid crystals. Its anatomical affinities with Melastomataceae
close
1975).
are
in the Dicots
about equally
with Crypteronia, and much closer than its affinities with Alzatea and Rhynchocalyx,
as
Lythraceae, Sonneratiaceae, and Oliniaceae.
somewhat
more remote
than with
Its anatomical
affinities with Dactylocladus
Crypteronia. Vegetative
anatomy
on
are
the whole favours
the retention of Axinandra in the Melastomataceae.
Crypteronia shares
fore be
equally
most
of
out
of its anatomical characters with Axinandra, and would there-
place
in
affinities with Melastomataceae
could
mataceae
be
supported
Sonneratiaceae, and
Lythraceae,
are so
Oliniaceae. Anatomical
striking that the transfer of this
than with Alzatea and
anyway stronger
with
Similarities
positively.
and
Rhynchocalyx
to
a
to
genus
several
lesser
the Melasto-
Melastomataceae
extent
are
also than with
Dactylocladus.
CONCLUSIONS
In the
previous
the anatomical
sections of this
paper
two
alternative
considerable anatomical heterogeneity of related
and Sonneratiaceae (see
strongly weakens
e.g.
the
table I), which
anatomical
Dactylocladus and Axinandra
transfer Crypteronia
an
anatomical alternative
macro-
and
arguments
pattern of
A
There is
polyphyletic
no
general
in
the
family
to
independent
group.
retain
in the Lythraceae, and
to
to
offer
evolution of
If Crypteroniaceae would
primitive Myrtalean stock, the anatom-
families
would exemplify
Lythraceae,
a
very
Oliniaceae,
peculiar
or
Sonner-
origin would explain the anatomical diversity much better.
agreement of the anatomical data with those from
although
the palynological
concept
for Crypteroniaceae
evidence
as
or
may
be
floral, fruit, and
interpreted
as
either
supporting the inclusion of several
Lythraceae and/or Melastomataceae (Muller, 1975).
The only possible synthesis
fruit specializations
Lythraceae,
of
natural groups,
proposals have been made
with those in other families (Melastomataceae,
pollen morphology,
supporting
a common
given
divergent evolution within the family together with developments parallel
convergent
atiaceae).
very
which have been put forward
demonstrate the apparently
and resemblances with other
have been
of Van Beusekom. The
like the Melastomataceae, Lythraceae,
taxa
Melastomataceae, Rhynchocalyx
natural group derived from
a
ical diversity
a more
to
sense
generally considered as
and micromorphological characters in this plant
represent
genera
in the
are
the Melastomataceae. Yet these
to
interpretations
of the Crypteroniaceae in the
heterogeneity
have led
etc., but that the
of all the conflicting evidence is
to
between all the
morphology alone. This conclusion
macro-
agrees
taxa
more or
and micromorphological
ranges witnesses
discussed than aparent from external
less with Hallier's view from 1918,
characters in this intelligent
the systematic position of amongst others, Alzatea, Rhynchocalyx,
and Dactylocladus.
accept that floral and
overlapping of anatomical and palynological
intimate relationship
who used both
to
the recognition of distinct families like Melastomataceae,
account
of
Axinandra, Crypteronia,
G.
C. M.
J.
The wide anatomical
anatomist to support
patterns
P. Baas:
Anatomy of Crypteroniaceae
the putative
relatives
in the
conse-
for
the
delimitation
new
would be
anatomy
on
family, be
same
and
impossible
it
for the
(1975) proposals
of this family. The only straightforward suggestion based
Axinandra and Crypteronia together
195
of Crypteroniaceae
overlapping each other, make
Van Beusekom's
reject
or
in
ranges
quently the complicated
&
Viiet
van
to
keep
either Crypteroniaceae
it
or
Melastomataceae. More detailed knowledge of the anatomical characters in all members
of Melastomataceae,
and Sonneratiaceae would be useful
Lythraceae,
about which
suggestions
representatives
Crypteroniaceae. Comparative
similar
are most
cannot
anatomy
specific
more
decide the family delimitation in this part
of the Myrtales, but the anatomical character complexes
to
for
the individual genera of the
to
are
unusual
specific and
enough
indicate probable relationships and aid in future revisions of those taxa.
ACKNOWLEDGEMENTS
The
authors wish
to
and
(Leiden, systematics
share in
M.
van
record
nodal
(Arnold Arboretum,
for
appreciation
Mr.
relationships).
the micro-technical
Zoelen
their
anatomy),
the
typing
Mr.
W.
L.
Mr. C.
work;
and
help
and Miss A.
Star
discussions
are
Howard
R. A.
F.
van
Beusekom
for
acknowledged
the
preparing
read the
critically
Dr.
to
and Dr. C.
M. Kuenen
Mr. B. Kieft for
Kalkman
C.
valuable
(Leiden, palynology),
Marks and
Dr.
manuscript.
for
Muller
J.
their
and Miss
plates;
manuscript.
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