Anatomical contributions to
and
Floral
plant taxonomy
anatomy of Eliaea from
vegetative
from Indo-Malesia
P.
and
Madagascar
I.
Cratoxylum
(Guttiferae)
Baas
Contents
Summary
369
Introduction
370
Techniques
371
Materials
Results
371
Discussions
and
Anatomy
of the
flower
in
Cratoxylum
372
Anatomy
of the flower
in
Eliaea
373
Vegetative
anatomy
Vegetative
anatomy
Occurrence
Floral
and
Sectional
of
of
Cratoxylum
papillae
in
381
delimitation
and
and
Eliaea
383
Cratoxylum
and
xylem specialization
Cratoxylum
373
of Eliaea
as
anatomy
distinct
related
in
problems
the
genus
383
385
Cratoxylum
387
genera
Acknowledgements
387
References
387
Summary
The floral
Eliaea
and
vegetative
(Madagascar) is
stantial
enough
recognize
to
of the
anatomy
described.
Although
them
as
closely
the
two
distinct
allied
taxa
genera,
as
in
many features
common, differences
from
appears
the
following
with
Ovary
with
and
Styles
12
or
3(or
filaments
Secondary xylem
ovules
more
4)
incomplete
true
septa
glabrous
fibres
scarcely pitted
with
are
sub-
survey:
Eliaea
Cratoxylum
Ovary
and
guttiferous genera Cratoxylum (Indo-Malesia)
have
Ovary
with
6(—8)
Ovary
with
6(or 8)
being
true,
Styles
and
three
ovules
incomplete
filaments
Secondary xylem
three
septa,
false
mostly
with
villous
densely pitted
fibre-
tracheids
Vessels
Silica
The
In
some
species
absence
Floral
shows
be
may
and
of
This
are
369
in
on
ray
insertion
of the
of
features than
hypogynous
(families,
on
scales in
findings
the
in
genera, species)
leaves;
but
both
exclusively
solitary
absent
genera
favours
be
the,
of lower rank is
is related
are
are
the
view that
these
and
suggested by
geographical
The
seem
to
xylem
be
the
structure
more
presence
distribution.
of Eliaea
specialized.
The
discussed.
mainly vegetative,
described
to
discussed.
the flowers
plant geography
which
will
taxa
this
and Eliaea
Cratoxylum ,
for
almost
bodies
derivatives.
Cratoxylum
that of
is the first of a series
paper
Silica
further differentiation into
xylem specialization
of these
Vessels
of wood
staminodial
as
a
multiples
cells
epidermal papillae
primitive
different ranks
studies
radial
in
Cratoxylum
possible implications
*)
in
interpreted
of abaxial
and
more
and
present
vascularization
structures
or
solitary
bodies
used
to
anatomy
of
dicotyledonous
discuss their
affinities.
taxa
of
Anatomical
Carlemanniaceae,Dirachmaceae, Huaccae, Lophopyxis, Oncotheca, Phelline, and Sphenostemon
progress.
BLUMEA
370
In Cratoxylum the section
from
the
other
uous.
A
Isopterygium (C.
of
representatives
with
comparison
This
section
1970
very different in its wood anatomy
(concerning parenchyma distribution)
sections of
The other
Isopterygium.
about
2.
glaucum) is
difference
data from literature
some
No.
XVIII,
arborescens and C.
Cratoxylum.
the evergreen habit in the
correlated with
VOL.
other
groups with
Cratoxylum
deciduous
is
decid-
are
and evergreen
members is made.
Introduction
In his
the
monograph of
views
genus
described,
was
I doubt whether
it, but would certainly have given it
Tridesmos,
sect.
two
to
C.
of this
Cratoxywith the
genus
I
distinction
would have made
as
section
a
or
separate
a
genus
subgenus close
even
maingayi’. He listed the distinctive characters
of
to
of the
follows:
as
genera
closest
most
a
(Guttiferae, Hypericoideae,
the relationship
on
'If I would have studied material of Eliaea
monotypic Eliaea Camb. from Madagascar:
before that
Blume
Cratoxylum
genus
457) expressed his
Gogelein (1968:
leae)
Eliaea
Cratoxylum
Ovary with
at
least
Ovary incompletely
12
ovules
Ovary with
3-celled
ovules
6
6-celled
Ovary incompletely
Styles and fdaments glabrous
Styles and fdaments villous ('cottonny
hairy')
Fruit with three
valves, often partly
caducous
Gogelein expressed
ovary
doubt about the
some
true nature
to
ground of his limited
false, incomplete
a
undertaken in order
A
be able
to
important
to
Although
notes on
a
justify
discussing
Madagascar
of Eliaea
nature
the ovary
three
true
on
the
and three
of the
finds its response
ovary
in
of the
several
genera
was
certainty. The distribution
plant
relationships
(e.g.
genera
within such
considered
to
Nepenthes,
groups,
in
be sufficiently
elaborate investigation of this kind.
an
amount
genera
of anatomical information
would add
accounts
to our
on
Cratoxylum
comprehensive
a
knowledge of Guttiferae
of representatives of large families becomes
enigmatic
want
to
use
their
on
knowledge
can
more
and
in
summarized
on
structure
be found in Brazier and Franklin (1961), Chowdhury
1967).
Schofield
nodal and petiole anatomy. Chemotaxonomical data
et
of
Accu-
successfully
anatomy. Additional information
and Ghosh (1958), and Balan Menon, (1956,
by Hegnauer (1966). Jacquemin
some
small families.
Hypericoideae
of the timber
and
account
anatomy.
1908) and Metcalfe and Chalk (1950) comprehensively
the available information
Eliaea.
possesses
more
interpret this distribution pattern, is
the affinities of
apphcations
The former
(1951).
key 'loges biovulees',
original description of Cambesscdes (1830).
occurs
if comparative anatomists
Solereder (1899,
ofEliaea, because of
ovary
detaded study of the vegetative anatomy of both
better understanding
descriptive
urgent,
semipersistent
generic description
the secretory cavities in Eliaea have been published,
mulation of
and
in his
establish the mutual affinities with
substantial
the anatomy of these
more
a
Indo-Malesia and
to
ovary
further detaded study about the
to
Allanthospermum).
order
6
uniovulees'. Gogelein himself suggested
observations that the
Moreover,
paper.
pattern
condition, but
thus confirmed the
septa. He
Gogelein's plea for
in this
the three-celled
have '6(—8) loges incompletes
to
of the
Perrier de la Bathie
and fruit three-celled and the latter reported in his
which would refer
is said
finally
valves
conflicting evidence given by Engler (1925) and
called the
with
Fruit
al.
on
(1969) investigated
(1968)
Cratoxylum
some
gave
particulars
were
chemical
on
summarized
compounds
of
P.
The anatomical
Baas:
descriptions
Eliaea
Anatomy of
this
in
and
Cratoxyhm
made according
were
paper
371
to
a
more
or
less
standardized scheme largely following Metcalfe and Chalk's practice. Accounts of mature
secondary xylem
detailed than those of the secondary xylem of young twigs.
more
are
TECHNIQUES
For the
the
stalk
to
stigma
40° C in chloral-lactophenol
at
young buds and
paraffin
in
other flowers and buds
addition,
In
embedded
were
flowers, and
which
to
added
were
to
cleared according
were
in
serial sections from flower
wax;
stained with auramin, safranin and astrablue according
were
Vagas (1961).
(1949)
the floral anatomy, series of pickled
study of
of Eliaea also fruits,
case
drops
some
of
30
Maacz and
Vautier
to
% hydrogen
peroxide.
Leaf anatomy
was
studied in
sections
transverse
the distal and basal part of the petiole. Epidermal
cuticular preparations (using
concentrated glacial
mixture of
a
acid,
acetic
Twig and wood anatomy
Quantitative features such
as
60°
at
and
characters
and
scrapes
C.)
transverse,
vessel member length
macerated material. The macerations
studied in
were
equal volumes of 20 % hydrogen peroxide and
studied in
were
through the middle of the lamina
made in the
were
radial and tangential sections.
and fibre length
same
way
measured in
were
the cuticular
as
prep-
arations.
For wood elements the
diameter etc.)
Orthoplan
based
are
based
means are
on
3 —10
Other
measurements.
25
Illustrations
(stomatal
means
made using
were
a
Leitz
tube.
with drawing
microscope
on
measurements.
MATERIALS
In the
out
following list of specimens, particulars
them. All material studied
on
present
at
specimens
available
are
unless stated ortherwise. Institutional wood
Leiden,
were
at
given about the investigations
are
obtained
are
abbreviated according
the Rijksherbarium;
carried
derived from herbarium and wood specimens
was
duplicates of
Stern
to
collections from which
(1967). Slides for reference
of them have been deposited in
some
the Kew slide collection.
Cratoxylum
and
Borneo,
C.
S.
of
H.
and
of
53506
of
flowers);
formosum (Jack)
formosum.
and
Java,
Cult.
Dyer. Borneo,
Philip.
and
Boschb.
Proefst.
sample
C.
ex
Malaya,
C. sumatranum
For.
C.
Serv.
bb
sumatranum
Eliaea
Arboretum
(tree
Hallier
Pickled
5) (clearing
and
twigs);
of
flowers);
Sumatra,
Alston
14359
sample).
material
C,
line
(leaves
samples).
(clearing
s.n.
(wood
VI.
Bog.
(wood
253,
33096
from
52a;
Botanic Gardens
Singapore
Buwalda
Borneo,
7979;
(clearing
Sumatra,
Forbes
Gardens
(clearing
and
3298;
Pickled
Hort.
SAN
Bog.
19.6.61
Nat. Herb.
Bogor
material
XV.
J.
A.
Bot.
Singapore
3
(leaves
and
and
twigs).
(wood sample).
87634 (clearing
Number
from
XII.
offlowers);
wood coll.
6468 (=
Borneo, Boschb.
Proefst. bb
bb 6770 of herbarium
14411
series) (wood
RTIw).
maingayi Dyer. Singapore,
twigs);
Kepong
twigs).
Borneo,
twigs)
21346
25128
pruniflorum.
ssp.
C. glaucum Kcrth.
(leaves
SAN
from
Noor 155; Sumatra, For. Serv. bb
and
21290
Cult. Hort.
(leaves
material
&
Hongkong,
Meyer
ssp.
formosum (Jack) Dyer
sectioning
C.
Dyer
flowers); Java,
Borneo, Meyer
C.
Blume.
Borneo,
Pickled
Kadim
Fuchs
25136;
(Lour.)
twigs);
formosum (Jack)
sectioning
Blume.
(Vahl)
flowers); Malaya,
SAN
cochinchinense
(leaves
C.
arborescens
sectioning
WT
6732
Burkiii
(Jack)
Blume
ssp.
3939
(leaves
and
(Jack)
articulata Camb.
HMB 2607
(wood sample
Blume.
Pickled
ex
sumatranum.
(clearing
of
flowers); Sumatra,
Grashoff 991
(leaves
and
KEPw).
Borneo,
Darnton
577
(clearing
of
flowers); Java,
Neth. Ind.
twigs).
Philippines, F.P.R.I.,
material
of flowers
S.H.
and
321
(wood sample
fruits collected
by
Dr.
ex
CLPw).
Capuron
at Fort
Dauphin,
BLUMEA
372
Madagascar,
was
used in
1967;
TEFw
gascar,
d'Alleizette
Madagascar,
clearing flowers.
SF
14577
VOL. XVIII, No.
(wood sample
of the
Anatomy
The following
account
of floral
parts
as
in
seen
is based
on
and Nov.
sectioning.
1970
1906. Material
The latter two for
from these 3 collections
vegetative
Mada-
anatomy.
TEFw).
ex
RESULTS
and of C. formosum and
Aug. 1906
s.n.
The first for serial
2,
AND
flower
on
DISCUSSIONS
in
Cratoxylum
(fig.
2)
1a,
serial sections of buds and flowers of C. arborescens
cleared material of all species of Cratoxylum. The arrangement
is illustrated in
section
transverse
fig.
Additional data
ia.
are
given below.
The
scales
hypogynous
are
inserted
the
at
same
several vestigial vascular bundles each from the
phalanges. This
tures
septa
(cf. Engler,
are
fused
cell
in
seems
(in
fused
C.
to
1925:
at
the
favour the view that they
The
183).
very
arborescens)
below the
ascendant anatropous
shown in fig.
runs
are
way.
styles.
Placentation is
attached
to
along
one
over
a
very
are
Fig.
1.
the attachment of
one
Transverse
indicated
there
axile;
at
ovary
3
(or 4) celled. The
(in
are
sections
o:
where
through flower
ovule;
p:
present.
petal;
struc-
C.
formosum)
buds.
p.s.:
or
3 —18 ovules per
the basal part of the septa
of Cratoxylum formosum is
distance (fig.
2a,
a
fragmented
b)
into
three
two
of the septa ('lateral' bundles). Minor vascular
supplied with vascular bundles
hypogenous scale;
bundles
coherent
the
as
staminodial
forms the dorsal bundle and each of the other
which
with the stelar cylinder. Some vascular tissue extends
h.s.:
as
incompletely
are
short
bundles connected with these three carpellary bundles
placentae
be regarded
the placentae
Vascularization in the
Each bundle branches
bundles of which the median
bundles
is
and receive
of the stelar system
Below the loculi three major vascular bundles depart from
2.
stelar cylinder.
Cratoxylum
the phalanges
as
region
may
base of the gynoecium and
(cf. Gogelein, 1968), wich
an
of
ovary
level
same
a.
are
to
occur
more
in
or
varying numbers. The
less linearly continuous
above the level of ovule insertion.
Cratoxylum formosum,
petal scale; ph: phalange;
X
s:
12.
b. Eliaea
sepal.
Loculi
articulata,
shaded;
X
12.
vascular
P.
Baas:
Anatomy of
The orientation of vascular bundles
was
Eliaea
and
Cratoxylum
difficult
very
determine because of the
to
low number of differentiated xylem elements before and
where orientation could
cases
lateral bundles
poles facing the loculi.
In the
found
were
distinct
no
vascular bundles. In C. arborescens the bundles
above the level of ovule attachment.
with their xylem poles facing
line through
not
material due
bundles
observed only in C. maingayi, but
the
enter
were
to
from
not
styles. The dorsal bundles
in the other
continue
are
as
a
median
derived from the
hardly observable
was
elements.
The
the septa could be
carpels opposite
species. The dorsal and lateral bundles
the stigmas where they branch pro-
to
up
(amphicribral)
observed running
of vascular bundles
different
com-
minor vascular bundles
found in the other species,
course
more
the individual
be centric
some
the scanty presence of differentiated xylem
to
fusion of lateral vascular
species
ovary
situation is much
boundary between
found
were
same
the axis of the
the septa. These bundles,
vascular system of the placentae. The
in cleared
the
In
and
vascular bundles (dorsal
have normal arrangement with xylem
to
placental vascular supply the
and is obscured because there is
plicated
very
the flowering stage. In those
at
be observed all peripheral
derivatives)
plus
373
fusely.
Secretory
cavities and canals
also in the style. They
in the
connective
abundant in the periphery of the
1968), but
petals (cf. Gogelein
from special
are
also abundantly present in the sepals,
are
glands
in
of the
Anatomy
absent
are
C.
formosum
flower
or
ssp.
in Eliaea
details in the gynoecium (see fig. ib) Fig.
comparison with those of Cratoxylum. The
the distance
although
uous
false
which the septa
over
composed
septa,
of
a
are
a
attached
to
the placentae
anatropous way. The vascularization of the
of Cratoxylum. There
broken
up
some
minor
to
are
be double
or
centric
do
tissue
ovary
in Eliaea is
of
ovary.
continue into
Spherical glands (i for each stamen)
stamens.
These
formosum
are
similar in
ssp.
occur
structure to
pruniflorum
are
among
the
In
ssp.
an
ascendant
are
is
that
not
Cratoxylum
arbo-
fused
some
over
The vascular strands in the
the styles.
to
placentae
fig. 3e). The
The dorsal bundles branch
is similar
to
that in Cratoxylum.
two
connective
1968, fig.
glands found in each
stamen
5k).
scales and the presence of
petal scales
in the
the other characters shared with Cratoxylum.
Vegetative
Leaf (fig.
carpels
two
in
As in
one ovary.
stelar cylinder
as
the
in the distal end of the connectives of the
(cf. Gogelein,
The vascularization of the hypogynous
corolla of Eliaea
in
fundamentally similar
The
to
of 6(—8)
presence
above the level of ovule attachment (see
not
for
(or 4) celled,
situated abaxially
6(—8) ovules
just below the stigmas. The distribution of secretory canals
of C.
3
sections
short. There are three conspic-
the basal part of the septa in
The lateral vascular bundles
lateral carpellary bundles
transverse
incompletely
differences, however.
distance (see fig. 3d—f) in die distal part of the
appear
that of Cratoxylum apart from
series of
just below the level of insertion of carpellary bundles
and formosum.
rescens
are
at
(apart
1b, 3)
superficial impression of the
loculi instead of the 3 (or 4) actually present. There
are
very
bulky parenchymatous
dorsal vascular bundle. These septa give
Cratoxylum they
a
also
is
free is
(fig.
to
3 gives
ovary
occur
pruniflorum).
The arrangement of floral parts in Eliaea is identical
some
wall and
in various numbers
in the androecium
rare
very
ovary
occur
Anatomy of Cratoxylum
4d—k)
surface view: Adaxial epidermis glabrous in all material apart from C. formosum
pruniflorum.
Cells polygonal
in outline with curved
to
straight anticlinal walls.
BLUMEA
374
Fig.
2.
Some
mosum, from
transverse
bottom
and
through
ovary
vascular
bundle;
(a)
to
styles.
x:
sections
top;
through
all
x
d.b.: dorsal
differentiated
VOL.
29.
a.
the
XVIII, No.
gynoecium
Vascular
bundle; l.b.:
of
supply
lateral
a
2,
1970
flower
of central
bundle;
o:
xylem element(s) (solid black).
of
Cratoxylum formosum
part
ovule;
Loculi
st:
of
receptacle.
style;
shaded.
st.c.:
b—i.
ssp.
for-
Sections
style canal;
v.b.:
P.
Fig.
to
top;
3.
Some
all
X
transverse
29.
For
Baas:
sections
legends
Anatomy of EHaea
and
through the gynoecium of
see
Fig.
2.
ƒ.s.:
false
septum.
Cratoxylum
a
flower
of Eliaea articulata
375
from bottom
(a)
BLUMEA
376
Fig.
4.
Leaf
12.
X
200. e.
C.
200. i.
,
abaxial
abaxial
and
X
x
C. formosum ssp. formosum,
Bog., lamina,
tory cavity
or
canal;
v.b.:
Cratoxylum.
12. c.
epidermis,
epidermis,
Cult. Hort.
black.
of Eliaea
through midrib,
maingayi,
ssp. pruniflorum
X
anatomy
b. lamina
X
transverse
VOL.
abaxial
200. f.
X
200. h.
a—c.
Eliaea
epidermis,
C.
No.
X
arborescens,
2,
section,
7979, abaxial
X
200. k.
distal
petiole through
a.
d. C. sumatranum, abaxial
abaxial
epidermis,
ibid.,
1970
articulata
200.
C. formosum ssp. formosum,
Buwalda
vascular bundle.
XVIII,
epidermis,
Cult. Hort.
X
j.
200.
Buwalda 7979. h:
x
200.
Bog.,
g.
end,
epidermis,
C. formosum
abaxial
epidermis,
C. formosum ssp. formosum,
hair;
p:
papillae;
Xylem shaded; phloem dotted; sclerenchyma
in
a
s.c. :
and
secre-
b solid
P.
Abaxial epidermis
quently
present
surface.
Tall
on
to
Cratoxylum
pruniflorum (fig. 4g).
ssp.
cylindrical papillae
±
material of C. cochinchinense,
and C.
and
(see fig. 4h, j, Table
sumatranum
abaxial
on
formosum,
ssp.
III and discussion
and absent from
on
Abaxial epidermis
outline with almost
on
to
of papillae.
presence
ameter
Stomata confined
adaxial surface overlying the midrib in
Outer anticlinal walls of
shapes.
of
tending
stomata
perpendicular
subsidiary
to
the
on
section:
Cuticle
cells
cells curved
sinous
to
(mean
14//);
in
some
of C. formosum ssp.
specimen
fj,
veins.
polygonal
Cell outline often
guard cells and subsidiary
1 —3
for-
cells of various
(see figs. 4d —i).
Di-
orientation random or
thick. Epidermal cells
p,
in
present
absent from other material.
Mesophyll composed
cells; abaxial chlorenchyma
cells
of
I
to
2
or
of
margins composed of small
broadly V-shaped
to
groove
amount
multiple epidermis) of
layers of tall adaxial palisade
tending
be palisade-like;
to
downwards to various
to
extents.
Midrib with shallow
of collenchyma.
and prominently raised abaxial surface,
supplied with single
vascular strand with sclerenchymatous
crescentiform
I —3
glaucum,
fairly loose, composed of cubical
rounded chlorenchyma cells. Leaf margins pointed
shaped adaxial
flattened,
to
arborescens and C.
C.
of C. cochinchinense also
in remainder of material abaxial tissue compact
square
Hypodermis (derived from
inner layer(s) of the
irregularly arranged cell layers, adaxially
Mesophyll
maingayi,
abaxial surface apart from
to
I
pore 10 —20
protoderm and therefore representing the
V—U
of the
C.
epidermis) major
(see fig. 4d—i).
those of abaxial surface smaller than those of adaxial surface.
or
abaxial
coincide with direction of major veins.
to
transverse
lobed
on
in part
383). Papillae confined
p.
Unspecialized
sinuous anticlinal walls
paracytic with low dome-shaped
mosum;
In
straight
surface
guard cells and subsidiary cells of stomata
in remainder of material.
glabrous
cells
hairs also infre-
glaucum,
C.
and from elongated cells overlying midrib and (if conspicuous
found
with broad basal
sub-species
In this
present
C. formosum
unspecialized epidermal cells
obscured by
377
adaxial leaf surface and low bluntly conical papillae present
short
to
of Eliaea
Anatomy
with indumentum of mainly uniseriate hairs
C. formosum
present in
Baas:
of thick
caps
walled fibres interspersed with thin-walled cells. Ground tissue of midrib collenchymatous
at
periphery
abaxial
in
parenchymatous
to
schlerenchyma
caps.
vascular bundles and also
including
central parts.
more
Sheath of
Veins with
translucent parenchyma
if present. Vertical
caps,
without ad- and
or
cells
present
around
bundle sheath extensions
associated with major veins present in all species apart from C. maingayi. Petiole ± semicircular
and
bean-shaped
to
abaxial
convex
strand,
in T. S.
surface.
without incurved
through distal end, with
Vascular
margin
glaucum). Ground
parenchyma.
Crystals
mesophyll (often
in
ground
Secretory
or spongy
clinal
tissues of
part of
cell walls
remainder
of
midrib and
Epithelium
Axis
(twigs
Epidermis,
present in
which
in cells
cavities of
tissue of
of the
of secretory
c.
the section Isopterygium
present in
(C. arborescens
petiole composed of collenchyma
amounts; druses
3 —5
in
mesophyll.
present in both. In
thinner in
canals
canals present
cavities and canals
mm
predominant
and thick walled
in abaxial part of
larger than surrounding cells); clusters predominan.
Secretory
petiole. Broader
of
crescentiform
and Tridesmos.
varying
are
epidermis
epidermis.
cortex
±
petiole and midrib and
or
flat adaxial surface
Cratoxylum
system
(sub) spherical shape, varying greatly
mesophyll
to
of single
in material from the sections
2( —4) additional latero-dorsal bundles
and C.
concave
composed
diameter;
a
few
places
of
in
small
Prismatic
in
cases
also
secretory
diameter
tissue
badly preserved
see
the anticlinal
opposite
ground
crystals less
frequency confined
in
present
of
most
commont
to
palisade
and/or peri-
cavities than in
in
phloem
of
petiole and midrib.
herbariumspecimens.
table I)
and perivascular fibres caducous in all twigs which
are
3
mm
or
more
BLUMEA
378
Fig.
5.
Transverse
b. C. cochinchinense.
f:
tracheid;
v:
c.
vessel.
sections
of
C. formosum.
Parenchyma
wood
d. C.
VOL.
from
Cratoxylum
arborescens,
shaded.
XVIII,
SAN
No.
and
26136.
2,
1970
Eliaea;
e.
C.
all
X
glaucum.
30.
—
a.
C.
sumatranum.
f. Eliaea articulala.
r:
ray;
P.
thick. These tissues
Baas:
only
were
Anatomy of
in
seen
a
very
Eliaea
and
Cratoxylum
of C. arborescens. Cork
twig
young
379
ting in pericycle between perivascular fibre sheath and phloem; composed
approximately
wall
flattened cells with adaxially
to
square
thickenings; these layers
are
herbarium specimens; composed
with
rays
poorly defined
oblique
35 —31
to
end
reticulate
wall
in
in
thickness
contents
arranged,
radial
p.)
sparse
and
unilaterally
irregular,
in
maingavi and C.
glaucum. Rays
seriate
mainly
15
25/mm
—
biseriate, (1 —)
or
cells. Primary
upright
some
to
rather
in L.S.,
types in
proportion of different crystal
irregular shapes, noted in
the canals
in
ray
a
cells which
varying
a
types
I
minutely
to
varying
xylem
of
2
trace.
of secondary
than
those
Fibres
C.
in
pits.
predomin-
formosum.
c.
arborescens and c.
twigs investigated,
of
composed
uni-
square
to
conspicuous radial
Pith forming
rounded in T.S. and
a
Crystals
phloem.
Silica bodies,
flattened
to rectan-
square
present
as
Amount and
globose
or
of
xylem of part of the material
layered epithelium;
narrower
±
Solid
bordered
amounts,
with
cylinder
varying infraspecifically
or
tracheids').
cochinchinense,
single leaf
are
in
Tracheids
specimens.
number of secretory canals.
parenchyma cells
the phloem being
in
Prismatic types abundant in
pith.
canals with
most
3 —4[i).
resulting
investigated.
confluent in C.
continuous
a
to
alternate arrangement,
to
high, heterogeneous,
Node unilacunar with
interspersed with
clusters and prismatic
investigated. Secretory
cells
many
short
secondary xylem. Walls varying
C.
or
periphery of secondary
of parenchyma
cylinder, composed
in
tangential diameter
('vascular
simple
present
bands in
apotracheal,
at
mean
specimens
of
tissue of
mainly paratracheal
—
xylem: Growth rings
compound,
frequent
with
provided
xylem forming
arrangement of vessels.
gular
3
vessels
in
constituting ground
sumatranum ;
of the
some
Average fibre length 420—740 p.. Parenchyma
antly
tangential cell-elon-
to
Perforations simple in the oblique
transitional
in
secretory
larger and slightly polygonal (diameter
in
present
oval in T.S.,
p..
in
pits
or
3
—
sparsely
very
(due
parts
wall). Secondary
to
255 —363
flattened
present
very
rows,
and
2
or
Tyloses
pattern.
paratracheally
arranged
polygonal
brown
amorphous
(—6); round
3
Lateral wall
walls.
cells, parenchyma,
intraspecifically). Vessels diffuse, solitary and
or
small and round-oval (diameter
pits
radial
extra
vessel member length
slightly oblique
Vessel—ray
an
absent (varying
or
radial multiples of 2
mean
[x,
slightly dilatating peripheral
very
of
and pitted
U-shaped
Secondary phloem often badly preserved
view.
of sieve-tubes, companion
the formation of
gation and/or
±
originalayers
occasionally interspersed with layers of thin-walled cells.
Cork cells ± hexagonal in tangential
canals and
lignified
of
present in
phloem and pith,
the pith.
Bark
Material:
sample
from bole
General features.
brown
than
For
2
tones.
cm
4 —5
apart.
mm
thick,
Shedding
field characters
Microscopic
see
of C.
arborescens,
Fuchs
Colour greyish brown
with
in thin
coarse
layers.
or
shallow longitudinal fissures, which
In T.S. with
ground
tissue.
Ground
Tangential bands (2 —3
and densely pitted
varying from
a
stone
tissue
composed
cells
few microns
mm
('phelloids');
to 0.7
mm
1—4
occurring
Bands
their inner sides.
cork
cambia
on
at
various
due to the rather
on
by
of sieve
both sides by
tubes,
more
companion
seriate. Secretory canals
thick) composed of
discontinuous bands accompanied
formed
are
yellow tangential bands.
numerous
Foxworthy (1927).
features:
cells and (crystalliferous) parenchyma strands. Rays
in
21290
dark brown with yellow and reddish
wavy
numerous
± cubical tliick-walled
intervals (their
course
of the
distance
bands). The
layers of flattened suberized cells.
BLUMEA
380
also Table II,
Wood (See
Cratoxylum and
A. Section
fig.
VOL.
Wood
General features:
Tridesmos (fig.
heavy
sumatranum, C. formosum and C.
yellowish
in
pinkish brown
to
other three species.
2
solitary and
Growth
in radial
oblique
pattern in C. cochinchinense and C.
in
T.S.,
3
4(
the oblique
predominantly
sometimes oval
oblique.
to
arranged
and granular brown
Parenchyma
10—32[i,
or
scanty.
very
species, tending
to
homogeneous,
very
in
B.
Section
apotracheal
1.1
two
shapes
in
mm, I —2
jd,
(fig.
of
—5/mm
elongated),
walls
tions
3 —4.(—6)
simple
pits round
slits
mean
[i
in the
to
,
solitary and
tangential,
tangential
6—8
oblique
of the
to
of
bands
maingayi,
wide.
cells
I —many
seriate in other
—3(or 4)
heterogeneous
tending
or
in
rough surface
cells.
arborescens
wood of C.
Growth rings
in radial
not
seen
multiples of
125—200fx,
two
and
C.
Texture
sections.
in material
examined.
in T.S.
—4( —5), oval
radial
glaucum.
brick red.
to
2
300—780 fx
diameter
(mean
up
to
—660
520
Vessels
(radially
—280/1,
210
/x). Perfora-
horizontal end walls. Lateral wall pits alternate. Inter-vessel
3 —6/u)
with oblique slit-like apertures.
neighbouring pits.
and then
tyloses and solid amorphous brown
narrow
more
cells
—4(—6)
Silica bodies with
than in the other
diameter
Vascular tracheids accompanying
few simple pits. Fibres
I
in
6)
unilaterally compound
together with vessels of
composed
1080—1320/t).
I
Pits
chyma cells (rays and axial parenchyma) half-bordered, similar
Thin-walled
by
sometimes
hexagonal
rows,
cells. Rays io—13/mm,
Crytals absent.
sometimes confluent with others from
occasionnally
vessels,
procumbent cells, but squarish cells occurring
thick, vessel member length
oval (diameter
and C. formosum
sumatranum
some
tyloses
heartwood vessels of C.
some
length 780—1690[i (mean
seriatein C.
mature
coarser
features:
2
C.
Apertures
sometimes
and minute. Thin-walled
noted in
Wood rather light. Colour light pink
fairly fine, but conspicuously
2
ray
e,
of
samples
General features:
diffuse,
Perforations
half-bordered;
distinct sizes in C. cochinchinense,
present in
Isopterygium
Material investigated:
walls
140—1909.,
Paratracheal parenchyma scanty. Diffuse parenchyma
mainly composed
Microscopic
but
variously disturbed radial
thick;
low uniseriates and in tails of multiseriates.
and of irregular
sparse
contents
Parenchyma strands of
be of
to
to
460—6oo;x).
bordered pits with strongly reduced borders and slit-
2—8/u
100 —400//.
exceeding
not
or
arranged
walls
predominantly
Distance between bands
high, height
an
(radially elongated)
with vessels of narrow diameter. Ground tissue
together
like ± vertical apertures. Fibres
diameter
pits
(also recorded for
few simple pits
very
similar
1906). Vascular tracheids accompanying
in radial 'tails'
offibres with
oval
to
(mean
—4(—6), forming
diameter 3 —5(—6) fx.
polygonal,
to
Vessel-ray
cochinchinense and C. maingayi
Moll and Janssonius,
in C.
Vessels
conspicuous,
2
diameter up
290—710(2
Vessel—fibre pits
unilaterally compound.
and solid amorphous
round
1
maingayi, light
C.
horizontal end walls. Lateral wall pits alternate. Inter-vessel pits
to
round,
horizontal
slit-like,
radial
90—iiOfx,
maingayi.
fine.
fairly
to
oblique multiples of
maingayi,
vessel member length
thick,
6)(j.
diameter
tangetial
mean
—
in
,
to
and IC.
C. formosum
Texture
rings absent
7 —18/mm
absent
Material investigated: samples of
heavy (specific gravity exceeding
to very
diffuse,
T.S.,
5a —c)
maingayi). Colour dark brown in
features:
Microscopic
simple
1970
2,
6)
5,
section
No.
wood from boles of C. sumatranum, C. cochinchinense,
mature
—
XVIII,
or
some
two
pits
contents
on
In
to
to
C.
glaucum
contiguous
inter-vessel pits
the parenchyma
noted in
some
the
parenor
cell wall.
heartwood vessels.
of the vessels, sometimes arranged in radial 'tails'
diameter. Ground tissue composed
less arranged
in radial
rows,
of fibres with
rounded
to
very
rectangular
in
P.
diameter
T.S.,
—38(2
20
Baas:
in C.
arborescens,
thick. Length 780—1560
[2 (mean
in
i—3
layers around the vessels
fluent in
some
places.
Very
of
i.i
mm,
(or 3)
scanty
to
few
in
present
6.
cells. Rays
10
tending
—
to
i/mm,
be of
cells, squarish
cells present
to
I
Tangential
(fig.
sections
through
view:
clinal walls. Cells
wood
most
rays;
x
77.
a.
ones.
composed
to
the
absent. Parennot
exceeding
arborescens,
rays
I
and in
the
or
tails
and of irregular
C.
glaucum.
b.
C.
formosum.
2
of
shapes
c.
Eliaea
articulata.
of Eliaea
pore.
epidermis
glabrous.
Diameter
widi
slightly elongated.
dome-shaped
cells.
subsidiary
perpendicular
to
cells
Epidermal
polygonal
the pore
of
2—5
p
straight
of
anti-
Stomata confined
Subsidiary
10—18
p.
cells
(mean
thick. Adaxial epidermal cells larger than
1
August
or
2
1906, absent in specimen
layers
of
palisade
cells and
wall
D'Alleizette,
tissue of
spongy
a
single
shallowly
strand
abaxial periphery
convex
or
sclerenchymatous
with thin-walled cells.
to
and
vascular strand with
with abaxial and adaxial
walled fibres interspersed
at
flat) adaxial surface
V-shaped
parenchymatous
in
thickenings
Nov.
1906.
at
abaxial
margins.
surface,
caps,
composed
lobed
Midrib
supplied
without incurved
Ground tissue of midrib
more
present
Mesophyll
inconspicuously
Leaf margins pointed downwards. Mesophyll parenchymatous
Vascular
tous
uniseriate
rough surface
and major veins
section: Cuticle
with sligh ly raised (almost
with
or
homogeneous, mainly composed
Multiple epidermis with irregular collenchyma-like
D'Alleizette,
cells.
scanty
orientation random.
transverse
abaxial
in
to
cells of abaxial surface
paracytic with high
often divided parallel
15p);
Adaxial and abaxial
unspecialized
overlying midrib
abaxial surface,
In
very
con-
21346. In C.
4a —c)
adaxial surface and
to
(2
cells.
ray
surface
In
—4( —6)
2
be aliform
to
in Fucbs
cells high, height
many
tending
in
tending
seen
distinct sizes in C.
Vegetative Anatomy
Leaf
only
parenchyma
bodies with
Crystals absent. Silica
rays.
bands,
two
or
381
glaucum; walls
(J, in C.
in C. arborescens
,
Diffuse
I
21
—
Cratoxylum
(2). Parenchyma predominantly paratracheal,
continuous
glaucum, heterogeneous
seriate in C.
multiseriate
—8
seriate and
i—4
of procumbent
Fig.
5
16
1070 —1210
glaucum parenchyma mainly aliform.
chyma strands
and
Anatomy of Eiiaea
margin.
of thick-
collenchyma-
central and adaxial parts.
Veins
BLUMEA
382
with sclerenchyma
provided
veins embedded in
vascular bundles
VOL.
which
caps,
XVIII, No.
are
surface.
and sclerenchyma
Vascular
Cratoxylum. Secretory
of
to
crescentiform
present in both
cavities
secretory
absent. Petiole
strand,
collenchymatous.
tissue
minor
present around
flat adaxial surface and
single
(sub)spherical shape
cells opposite
cells
sheath extensions
concave
Ground
margins.
cavities of
Adaxial epidermal
tissue.
prominent opposite xylem,
most
Bundle
caps.
composed
system
without additional incurved
1970
Sheath of translucent parenchyma
mesophyll.
semi-circular in T.S. through distal end, with
abaxial
2,
with
convex
with
Crystals
palisade and
as
or
in
spongy
thinner
conspicuously
anticlinal and periclinal walls than remainder of epidermal cells. Secretory canals abundant
and with
(twigs of
Axis
Peripheral
of
two-layered epithelium
a
in
Secretory canals
c.
tissues
diameter;
mm
3
as
in
also
see
diameter 35—36
fx,
mean
vessel
not
contents
seen.
sometimes flattened and
tracheids
Vascular
tissue of
secondary xylem.
length 670—780
bands apotracheal
periphery
as
in
of
or
oblique end walls. Lateral wall
Wood
further
present
also
Table II,
Material investigated:
fig.
outer
features:
General
dark brown.
—4
Wood
—1200
(j.
apertures
fx,
minute
with
brown-yellow
in
as
types in
tissue
(mean
i—3
to
composed
hexagonal
stem
of
5
heavy
very
cm
and
phloem
cells
wide
Parenchyma
exceeding
to
mean
or
clusters in
gravity
1). Colour
exceeding
Vessels diffuse,
c.
12/mm
in the
or
slit-like.
Vessel-ray
0.8
at
Tyloses
contents.
absent.
in
arranged
20—30
various
distances.
5 —8 cells.
p,
walls
oblique
and
Vascular
radial
4—11
apotracheal
to
horizontal end
vessel—parenchyma
tracheids
with
rows,
in
p
thick; length
c.
types
I
i/mm,
i
to
many
seen.
numerous
3 p..
Fibres
900 —1700
parenchyma
cells
p
axial
present in
scanty.
high (height
heterogeneous, composed
infrequently
c.
not
irregular tangential bands of
Diffuse and paratracheal
Rays
mm), uni—biseriate, markedly
bodies absent.
almost
,
Vessels in heartwood with
unilaterally compound.
in T.S., diameter
strands of
2
diameter 95 ft; walls 4—7 fx thick; vessel member
of fibre-tracheids,
upright cells. Crystals of prismatic
Silica
as
at
pith
diameter.
(specific
Growth rings absent.
p). Parenchyma predominantly
1290
node and
Cratoxylum.
bordered pits with slit-like, almost vertical apertures; diameter of borders
round
numerous
Parenchyma mainly irregularly banded;
6e)
and round
solid amorphous
Ground
provided
and
rows,
contiguous fibre-tracheids in alternate arrangement, diameter
to
half-bordered
similar
Walls thick and
u.
(mean 780 fx). Perforations simple
but
pits
Tyloses
in radial
elongated primatic
of
features:
walls. Lateral wall pits
3
Vessel-paren-
arranged
Cratoxylum. Primary xylem,
in
as
canals
5f,
part
exclusively solitary, roundin T.S.,
450
3 p..
Texture fine.
Microscopic
length
as
bodies absent. Secretory
(see
c.
unilaterally compound.
aliform confluent. Paratracheal parenchyma scanty. Raysc. 22/mm
secondary xylem,
Cratoxylum. Crystals
pith. Silica
diffuse,
Fibre-tracheids
rare.
bordered pits. Average fibre
constituting ground
Vessels
vessel member
mean
pits in alternate arrangement. Vessel-fibre pits round-oval, diameter
chyma and vessel-ray pits
petiole.
inconspicuous.
very
table I)
Perforations simple in the slightly
390—450 (X.
and
narrow
Cratoxylum. Secondary xylem: Growth rings absent.
almost exclusively solitary; round in T.S.,
length
tissue of midrib and
ground
cells in
large
of midrib and petiole
phloem
of
not
procumbent
parenchyma cells.
P.
Baas:
Occurrence
In
two
of
specimens
Cratoxylum
were
this
Results
where
either;
conspicuous
that in
papillae. Such
from
a
this
led
a
very
conspicuous,
trace
aim
My
very
striking
(magn.
was
to
of infraspecific
taxa
some
species
pattern is
rank.
and
not a
the
taxon
occurrence
papillae
formosum
ssp.
here
formosum;
reflects
this character
definitely deserves further
specimens of
C. cochinchinense listed under the
has
a
were
inter-
occurrence
we
find
of
high
a
be investigated
more
all
here that
differentiation into
further
to
whilst
lower,
tentatively suggested
It is
attention. The
not
remarkably
is
proportion
of papillae
taxa
an
in material from Java and Sumatra; in the material
this
variation patterns
differentiation of subspecific
exhibit
to
reliable diagnostic character for species of Cratoxy-
obvious in C.
the Philippines
infraspecific
found
kind of geographical pattern exists for the
a
from Celebes and S.E. Asia lack papillae.
lower rank. If
were
cases
smooth
not
was
From table III it becomes evident that the
low papillae.
very
but where the surface
seen,
in
60)
X
know whether
distribution of the specimens
geographical
to
The
study all material of
me to
stereomicroscope
section such leaves
transverse
very
of specimens with
Borneo
specimens
in
formosum sectioned
subspecies
with
to
could be
papillae
of epidermal papillae is
proportion
ssp.
absence of papillae.
or
relationship
any
when examined in
but
lum,
one
whether it could be used
mediate condition with
presence
Cratoxylum
listed in table III. Specimens hsted under the question mark represent
are
no
of
Rijksherbarium
presence
character showed
concerned and
383
Cratoxylum
completely devoid of papillae (fig. 4i, k).
the
present in
and
noted on the abaxial leaf surface (fig. 4h, j). The leaves of the
were
establish the
to
in
papillae
differences in the abaxial epidermides
order
Eliaea
of four leaf samples of C. formosum
two out
tall epidermal papillae
other
Anatomy of
of
taxa
extensively
comparatively high proportion of
mark could be
question
progressed
as
far
as
in
C.
an
indication that
formosum.
The differentpatterns for the occurrence of papillae in different species growing under
similar conditions (cf. Gogelein,
1968)
that the
suggests
presence
or
absence of papillae is
the environment.
determined by
not
Floral and
xylem
specialization
and related
problems
In applying the general criteria for trends of phylogenetic flower specialization (see
Takhtajan 1959)
that of
one
has
to
regard
Cratoxylum because of
a
the gynoecium of Eliaea
seems
character. A
with other
not
comparison
of trends
available,
Detailed data
Frost
(1930),
purpose
1) long
it
far
as
floral
in
on
as
to
represent
known
to
more
specialization,
but
would be
rather than
necessary
comprehensive
e.g.
specialized than
for
presence
some
a
data from
of
vestigial
good underliterature
are
me.
xylem specialization
to
a
Hypericoideae
specialization,
Tippo (1938), Metcalfe
suffices
be
reduced number of ovules in the former. The
false septa in Eliaea also
standing
to
summarize the
are
and
to
be found in Bailey and Tupper (1918),
Chalk
(1950),
and
Carlquist (1961).
For
our
following trends:
fusiform elements (particularly
vessel members)
are
more
primitive than short
ones;
fibre
2)
tracheids
3) xylem
are
with diffuse
more
primitive than libriform
solitary vessels
Bailey (1957) and Carlquist (1961)
is
more
have
fibres;
primitive than xylem with vessel multiples.
pointed
out
the dangers of using these trends,
which have been established for the higher categories of land plants
of
specialization
within small
however, equally
groups
of
closely
allied
taxa.
In
my
to
judge the degree
opinion these trends
valuable for both closely and distantly related groups, perhaps
are,
even
BLUMEA
384
readily interpretable for small
more
VOL.
XVIII,
groups.
In the
No.
and members of different
anatomy. These factors
other
on
expect linear
The
species.
like
only
result
we
I
II),
&
may
the
to
find
some
characters
primitive
and
cochinchinense (in twigs
wood).
for xylem specialization
in
Eliaea,
mature
It is
though
C.
wood)
course
we
we
know
must
not
the other
to
which
representatives
are
more
Cratoxylum and Eliaea,
to
the longer
in
confusing
secondary xylem
group of which
a
and in exclusively
equalled
great, is
formosum (in twigs) and
that fibre length is
we
fusiform elements
bordered pits
comparatively
known, however,
since
families and
as
more
member had been ancestral
fibre tracheids with conspicuously
in
mature
is
in their
comparable
than others. Returning
more
solitary vessels. Fibre length
twigs and
hope for
much
a
closely related. Of
are
primitive
more
ancestor
common
find that Eliaea shows
(tables
if the
as
sequences,
possible
a
that its representatives
grounds
not
play
may
less important within
apparently
are
often
are
groups
1970
higher categories such
orders, parallelism and different speeds of speciahzation
part
2,
C.
by
c.
maingayi (in
not an accurate
index
considerable elongation of these cambial products
takes
place.
Here it should also be stressed that
in the
disposal
at our
was
since Eliaea is
a
the
Cratoxylum
case
only
a
wood sample from
of Eliaea. Bole-samples
in
as
shrub. It might well be that in branch wood of
Quantitative
xylem characters
because of
arborescens).
For
a
listed in tables I and II
as
high intraspecific
problem
our
occupies
the maximal
Eliaea resembles that of
extremes.
It is difficult
Cratoxylum.
to
a
are,
as
reconcile the
plant
in
might
however,
that the
taxa
expect from
gross
or
flower and fruit
(Cratoxylum
Eliaea is
a
or
at
shrub found in the
of Eliaea is
time
d'Asie
its
used
must
more
ct
'Ce
to
rare
phenomenon
(see
oxylum and
anatomy in Cra
that Cratoxylum and Eliaea
to
any
other
or
not
are
so
For the main
genus.
hypericoid
Vismia the reader is referred
For each of these
genre
d'Oceanie. C'
occurrence
along
primitive than
be explained
that Cratoxyleae
found in Eliaea.
explaining
of a mixture of obviously
no means a
there
genera
apart
strip of dunes along the east
narrow
(194.8: 114) stated:
tentatively ascribed
a
that the
closely than that of
morphology. Within the Hypericoideae
est
the
monotypique
done
the
An
occurrence
in
that of
un
east coast
seeds. The derivation of Eliaea from Cratoxylum
now
c.
of values Eliaea
are
to
either
from the Cratoxyleae
and Eliaea).
affine des Cratoxylon
Madagascar
range
more
occurrence
by
xylem characters which put them rather far
Perrier de la Bathie
xylem
table II:
formosum;
vegetative anatomy from the other
Engler (1925) and Metcalfe and Chalk (1950).
He
is
Ascyrcum, Haronga, Hypericum, Psorospermum
genera:
wide
tentatively conclude
can
that floral and xylem
morphology
gross
pronounced.
suitable for diagnostic
I: C.
common ancestor
closely related mutually than
more
differences in either
we
a
disposal;
at my
even more
contradictory conclusions from floral and wood
mentioned, however,
one
diameter
available
sinnlar diameter from
a
not
are
(e.g. table
know that in
differently diverging lines implies
closely related
they
possible
Cronquist 1968). The suggestion
Eliaea show
to
From the above
advanced and primitive characters
e.g.
variation
it is sufficient
xylem of
should be
anatomy. It
cm
are not
values would be lower than in the material
quantitative
this would make the differences between Cratoxylum and Eliaea
purposes,
of 5
stem
a
Cratoxylum
is,
to
by oriental origin it,
type
or
its
coast
most
If the
see
presence
understand the present
on
by
distribution pattern Madagascar—Indo-Malesia
goes
back
to
since
of Eliaea
the
on
have arrived
xylem characters
Danser
Madagascar,
distribution of Eliaea. This hypothesis
est
etc.
of the winged
unlikely
ancestor, must
already used
of primitive Nepenthes species
mais il
d'origine orientale,'
Indo-Malesia still showed the primitive
alternative hypothesis,
of Madagascar.
endcmique,
dispersal by
however,
Cratoxylum.
est
(1928)
can
implies
also
for
be
that the
the time that land connections
P.
between the
two areas
Anatomy of
Eliaea
and
still present and that
were
the whole
Eliaea occupied
Baas:
385
of Cratoxylum and
common ancestor
a
(see also Schuchert
range
Cratoxytum
1932;
Willis
1932; Van Steenis 1962
and Florin 1963). A later fragmenting of this Gondwana continent made an independant
evolution of both Cratoxylum and Eliaea possible. During this process Cratoxylum retained
primitive
flower characters
grandifolia (Oxal),
an
and Eliaea
retained primitive
endemic form from
,
also Sarcotheca, all from Malesia (Veldkamp
which the data known
at
before it is
pattern is needed
Great
care must
gantly established, but
1944)
and
possible
characters
together
a
general
two
list
70t
any
wood
(C.
sumatranum
and C.
the phylogenetic
to
trends
ele-
very
two
Tridesmos
the
grounds of
that
him
C.
arborescens
can
be
papers
distribution.
cited
distinguished
section
is
wood-
ligustrinum
own
obser-
Moll and
One group
Cratoxylum
comparatively
5a —c),
group
anatom-
confirm the fact
possess
parenchyma (see fig.
fairly thick-walled fibres (high specific gravity). The other
My
Cratoxylum.
The species of these sections
banded
C.
1899,
therein),
in
maingayi) and by
C.
by Englcr
differs
(C. formosum and
parenchyma
(C. formosum and
cochinchinense).
wood anatomy,
mention the decisive
from literature (Solereder
and the
1950
grouwps
on
Cratoxylum
genus
names or
emphasized
type of
vessels, abundant predominantly
narrow
specific
descriptions
Janssonius 1906, Metcalfe and Chalk
represented by section
and
generally
represented
by
C.
arborescens and C. glaucum constituting the section Isopterygium. Wood of these species
characterized by wide vessels, less
thin-walled fibres (low
specific
abundant
gross
not
be
interpreted
morphology. That the
In
fact, the
two
genus
other sections. It is
are
The
This indicates
distinction
a
two
be
distinguished instead
Cratoxylum
that they have
no
sections
are
group,
unlikely that,
C.
wood
cannot
be separated
two
It
groups
in
coincides
Isopterygium
would be
terms
very
of
a
are
to
differences between
with
to
energy
another
trees;
genus.
important
those of the
interpret the anatomical
functional adaptation
requirements
a
other members of the
to
the behaviour
increased storage facilities in the abundant parenchyma
useful for
on
flowering material,
evergreen
attractive
and
Isopterygium and that
of such
in the absence of
structure
of the
anatomy
sound morphological basis.
glaucum would be related
of the section
deciduous.
differences between the
the deciduous
or
surprising than the lack
wood anatomical
of the foliage. Initially,
[i
can
groups
disagrement between
a
rather isolated position of the section Isopterygium within the
in
feature: the representatives
other
mean
more
most
wood sample of C. arborescens
genus.
as
great differences between the wood anatomy of section
of the remainder of the
the
not
two
Tridesmos and
sections
the basis of wood anatomy does
is
sd, e), and
paratracheal parenchyma (fig.
gravity).
The fact that wood-anatomically only
three sections should
to
similar distribution
a
different from those recognized
species investigated by
that wood anatomically only
is
weight
the
that,
paper
sections,
having another
in
with
with
probability of the explanation offered.
anatomy in
Desch (1941)
the other
cochinchinense)
C.
=
vations,
in
into
he did
involved.
anatomically from
merely a model in
proved.
(1926) suggested
Cratoxylum should be split
ical
is
No factual evidence is available
groups
much
too
similar explanation.
a
hypothesis
absolute certainty of their irreversability has only been claimed
never
(1925). Unfortunately
plant
ascertain the
to
Sectional delimitationand
Den Berger
fairly well.
attaching
call for
might
Dapania
Dapania species but
two
The general trends of xylem specialization have been
an
was
fit
research into
more
also be observed in
of xylem specialization.
(Baily
seem to
present
much
Obviously,
support it.
and
1968)
here that the above mentioned
emphatically stressed
It is
characters.
xylem
Madagascar, resembles
during leafless periods,
were
cells of
consid-
ered. Some data from literature
out
is the
general relationship
habit
parenchyma
fagus and found wood parenchyma
New Guinea, whilst it
was
example
rule
a
in order
cases
or
find
to
whether there is
the evergreen
or
no
deciduous
the wood anatomy of the genus Notho-
abundant in the evergreen section from
very
small
a
evergreen
reversal, therefore, of what we found in Cratoxylum.
a
and Muller (1942) tried
(1939) and Tilson
of American Quercus.
for subgeneric segregation
of
the deciduous sections and in
sparse in
very
Zealand. This is
section from New
Williams
be
to
1970
distribution and
Ingle (1954) described
all. Dadswell and
at
2,
comparable
collected about
were
whether the situation in Cratoxylum
between
No.
XVIII,
VOL.
BLUMEA
386
to
find wood-anatomical evidence
found
They
no
substantial
from
difference
surprisingly
and of deciduous oaks, apart
between the wood of evergreen
of species. The above mentioned
in the former
more
pronounced ring-porousness
though
examples,
very
few,
group
remove
ground for accepting
any
functional explanation
a
for the differences in wood anatomy in Cratoxylum. Suggestions
more
of the
knowledge
knowledge
is
Returning
the xylem
differences found
the
to
diameter and frequency
and vessel
From this
glaucum
,
young
be banded
may
i
mature
xylem of
juvenile secondary
such topics require
trees
involved and such
secondary xylem
parallelled by
not
are
those
and
In young twigs of C. arborescens
twigs.
almost the
to
do
it should be noted that the great
Cratoxylum,
same
extent
as
in the other
m
C.
species
show deviations from the remainder ofthe
not
the
might conclude that
one
represents
often retains the
of
anatomy
exist in the
glaucum parenchyma
genus.
autecology
and cambial activity of the
on
obtainable in field studies.
only
to
not
a
xylem
mature
of C. arborescens and of C.
first formed secondary xylem
derived condition, considering that the
a
more
Such
(see Bailey 1944)-
characters
primitive
a
conclusion
seems to
where the vessel members of the
features listed in table II
disagree with the quantitative
and
C. glaucum belong to the longest in the
of C. arborescens
secondary
mature
xylem
This again would
genus.
imply
a
more
primitive condition
for young twigs
However, if we consider the data
arborescens
are
increase in
secondary
xylem
length, which
interfere with
186) could
specialization,
to
to
species
to
species (see
the picture
as
Isopterygium
in the
case
may
increase from the
as
related
to
to
phylogenetic
of Cratoxylum. Considering
in C. arborescens and C. glaucum
one
be derived rather than primitive
with the other sections of Cratoxylum (cf. Carlquist,
and C.
It has beenknown
Bailey and Tupper, 1918:
e.g.
theories of vessel member length
our
section
formosum.
two species.
glaucum
mature' secondary xylem. This
the much later formed
paratracheal parenchyma
regard the
for the former
that both C.
that fusiform element length
varies from
and complicate
wide vessels and
inclined
of C.
within the limits of individual variability
from Sanio's time onwards (1872)
first formed
note
we
as
the
would be
compared
1961, for trends of xylem
speciali-
zation).
The considerations offered above do
Eliaea
there
having
are
a more
consistent
primitive xylem
for both
young
Another anatomical character
of the
Isopterygium
species
not
and
mature
provided
within the
affect
the
conclusions
than Cratoxylum since
genus
secondary
argument
an
as
a
the section
other
Isopterygium
species.
The presence
or
absence of
a
often infraspecifically (e.g. in Eliaea articulata,
break
down in
a
Cratoxylum
as
a
xylem.
whole: in the distal end of the
occur
which
multiple epidermis
would offer yet another character
to
diagnostic feature
are
position
petiole
of
absent in the
in leaves of species of
distinguish them from
multiple epidermis
seepage
about
employed
for the rather isolated
latero-dorsal bundles
C. arborescens and C. glaucum 2 ( —4)
other species of Cratoxylum. The presence of
page 384
on
the characters
varies,
however,
the
quite
381), and this character could well
if
more
material
is
examined.
P.
Baas:
Anatomy of Eliaea
and Eliaea
Cratoxylum
From the
in
stamens
phalanges;
also essentially similar. However, quite
as
well. All characters
wood of Eliaea,
could be added from
The differences between
Isopterygium
mainly based
370) could be confirmed and
anatomy:
be
moreover,
two
are
sections of
I
Eliaea and
agree
axis is
present
three more
with
form
to
bundles in the
Eliaea.
to
one
differ in
genera
are
xylem. The latter character
grounds for
see no
com-
genus.
Cratoxylum,
genus
C.
maingayi
probably
is
by the fusion
supported
this view is
Anatomically
of
less important than vessel grouping and
taxonomically
with Gogelein that within the
closely related
two
whilst the differential characters within Cratoxylum
as
Cratoxylum
occurrence
Cratoxylum.
substantial than between the
more
distribution in the secondary
regarded
of
Cratoxylum. The
of fibres. Considering both floral and vegetative anatomy I
type
bining
are
fibre-tracheids
common
the representatives
in
rays
Cratoxylum and Eliaea
parenchyma
on
in
and the other
both floral and wood characters,
may,
p.
vegetative
contrasted with scarcely pitted fibres and the
as
multiples and silica bodies
section
general histology of leaf and
the
bordered pits, almost exclusively solitary vessels, and the absence of silica in the
numerous
vessel
characters both in
are many
number of distinguishing characters
a
listed by Gogelein (see
as
characters
diagnostic
genera
shared by Cratoxylum and Eliaea. For instance petal scales,
structure
glands, and
hypogynous
distinct
as
387
Cratoxylum
follows that there
part of this paper it
descriptive
floral and vegetative
and
most
of lateral
and the absence of vertical bundle sheath extensions in the veins of
ovary
the leaves in both Eliaea and C. maingayi. These
features
two
were
not
encountered in the
other species of Cratoxylum. Moreover, C. maingayi possesses xylem fibres with pits that
are most
and of which the borders
numerous
of the other Cratoxylum species. These fibres
fibre-tracheids
least reduced
yet
as
compared
different from the
very
with those
densely pitted
however.
Eliaea,
in
are
are
ACKNOWLEDGEMENTS
after
study became possible
This
Prof. Dr. C. G. G. J.
van
Steenis.
the revision
Through the
of Cratoxylum
and
courtesy of Dr. R.
suggested by
was
ofTananarive
Capuron
he obtained excellent pickled material of Eliaea; later Dr. Capuron also provided
sample.
Ir.
Th. Wassink
J.
of
wood samples
I have
to
(Amsterdam)
and Dr. Meng
van
Jodrell Laboratory, Royal
the english
wood
of the
Heel and Mr. A. J. F. Gogelein for their keen interest and
the initial stages of this study I received hospitality in the
During
Gardens,
Botanic
Kew. I wish
and the members of the anatomy staff for their
very
a
some
Cratoxylum.
thank Dr. W.
critical suggestions.
(Kew)
(Kepong) provided
kindly provided
with references
me
help
on
in
to
thank Dr. C. R.
many
ways.
Miss M.
Metcalfe
Gregory
anatomical literature, and polished
text.
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I.
1944. The
W.
Am.
research.
——
1957. The
of
&
BALAN
J.
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MENON,
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lingen
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timbers
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van
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in the
anatomy
study
of
phylogeny
and classification
243 —254.
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W. W. 1918. Size variation
P.
1967. Structure and identification
BBRGBR,
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421 —428.
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H. D.
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A.
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R.
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1941.
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FOXWORTHY,
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G.
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C.
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W.
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SANIO,
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8:
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Penninsula
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II.
Malayan
Evolution
For.
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Rec.
No.
3.
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Pflanzen,
R. R.
Sure
1969.
phytothcrapie
A
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new
Bl.
Cratoxyluin
(Guttiferae).
Blumea
15:
453 —475.
4.
Hypericacee
unc
de
Madagascar,
l'Eliaea
articulata
196 —203.
3:
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of cellulose
and
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Mikros-
L.
in
Mem.
Ober
1906.
1948. Notes
de
die
of the
Mikrographie
Dicotyledons.
des Holzes der
aufjava
vorkommenden Baumarten,
biogeographiques sur quetques plantes ayant
Sci.
l'lnst.
H.
Humbert,
Anatomy
1950.
H. H.
Flore
de
de
Madagascar,
Madagascar
Grosse der Holzzellen
der
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ser.
des
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I:
contribue
au
I.
peuplement
113—131.
Comorcs,
135c
Famille.
gemeinenKicfer (Pinus silvestris).
K.
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401 —420.
D. E. K.
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STEENIS,
C.
C. G.
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1932.
H.
Blumea
G.J.
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and related
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Mem.
N.Y. Bot. Gardn.
A. H.
O.
Quercus.
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Geol.
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1968. A revision
1939.
Bot.
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875 —915.
with
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to
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49.
Angiosp*rmen.
29:
of
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Moraceae
florale
chez
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their
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Polygonacees.
Bull. Geol.
Soc.
Bot.
12:
Dapania Korth. (Oxalidaccae).
tissues of the oaks
indigenous
delimiting Erythrobalanus
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Candollea
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43:
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353—365.
1932. Isthmian
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523 —529.
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Secondary vascular
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WILLIS, B.
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1949.
VELDKAMP, J.
Die
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1959.
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VAUTIER,
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1908.
235—542.
1967.
A.
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SCHUCHERT,
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18:
TILSON,
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CHALK,
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SCHOFIELD,
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&JANSSONIUS,
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Chemotaxonomie
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A
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90:
DURET, S.
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100:
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Blumea 15: 519 —543.
States,
Bull.
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219—343.
I. The
Torrey
importance
Bot.
Club
66:
P.
Baas:
Anatomy of
Eliaea
and
389
Cratoxylum
and
length
diamet r,
310—430
p
p.
Tracheid
Eliaea vessel
in
mean
340 340
320
—320
310
510510
280—
370
280
410—
620
720720 960960
for
and
%
only.
820820 850
35
Cratoxylum length, measur nts
of
xylem
secondary
member
first
of
char cters
on
vessel based
for are
%
25
formed
10
length
to
tracheid
amount
of
They Means
table. length.
this
in
Quantiave recorde
I:
TABLE
not
are
fibre
for
%
range
470— 650—8—50
mean
600
—620
350—850 490—9—0
30 — 40 — 420—
41100—5—2 72015— 68 1— 637 240—853 16
length
300
p
[A
in
in
Fibre
510510
670
420— 670—780
21—84 21—63
15—5 16— 26—
37—51
35—36
60 —730
740
560
84
84
diamet r
55
16— 31—68
räange
58
58
5252
4
0
40
35—36
3
p
Er-
in
in
Vessel
36
mean
Ves el
45
440
length
member
4
422
range
215— 25 —435
mean
310 315
42
36
485
185—325 2 0— 2 5—650
175—540 175—4 5
p.
F-
in
25 —365
270
265—
345
Cratoxylum:
Cratoxylum
Section
sumatrnum cohinc ens
C.
C.
C.
Tridesmos:
Section
Isopterygium:
(5
(5
formosum maingayi
C.
340
390—450
specimens) specimens)
specimens)
deviatons
Standard
900
C.
C.
Section
(2
(2
arboescens
glaucum
C.
C.
articulat a
Eliaea
VOL.
BLUMEA
390
wall
diamet r,
Fibre
thicknes
Fibre
diamet r
(A
}i
in
in
5
range
vessel
for
Eliaea
[A
fj.
in
in
and
of
length
length,
25— 13
length.
fibre
for
to
of
char cters
%
amount
this
Quantiave
II:
8
6
8
3-8 3-8
32
20
3
2222
10—15
16— 14—
15601560 16901690
1300130 1610
3
4
IXIX
4—
38
30 28
3030
1430
1430 15601560
21
20—38 20—30 20— 16—
20—
1—
Tangential
member
in
recorded
VeVess sel
1190 1260
1080 1320
1120 1070 1210 1120
1290
315 160160
140140
145145 160160
diamet r inji.
[A.
in
mean
90
90
90
90
60— 60— 50—315 50— 50—
90
90
180
155
155 180
no
no
550 710
600
290— 365—560
ranngege
230230 270270
20— 30—
200
200
125
95
95
—780
—1120020
290— 40 —
—790
400
30 —715 390—845 325—780 450—
325
450
470 600
620
620
520 660 620
780
780
[A
inji,
m
mean
470 460
470
25136
are
Cratoxylum:
deviatons
Standard
1700170
910—1430 780—1430 104 — 780—1430 90 —
125
length
1430
780— rio —
not
TABLE
4
2—3 2—4 2—3 2—4
910— 104 —
5 —140 30—125
range
17
They and
table.
mean
vessel
%
24
(i
JA
in
m
Fibre
member
for
mature
1970
921 36—50 14 780—3295 47016— 602—491783520—9847 10— 54962
range
xylem vessel
secondary
2,
%
35
Cratoxylum
No.
2—5 3—6
16—
16—
range
XVIII,
Cratoxylum
Section
sumatrnum cohinc ens
C.
C.
C.
Isopterygium:
Tridesmos
Section
formosum maingayi
C.
C.
Section
SAN
2212901290 21346
F
F
arboesc ns arboesc ns arboesc ns
glaucum
C.
C.
C.
C.
articulat
Eliaea
P,
BAAS:
Anatomy of Eliaea
and
391
Cratoxylum
(see
0
1
??
glaucum
steromicsope,
- — — 421
C.
magnifcto
Cratoxylum
C.
+
+
C.
present
III:
—
ssp.
—
??
—
+
+
when
papil ae
4
24
24
— —
0
0
0
0
210 180 070 500
i
2
8
I
epidermalpapilae
doubtfuly
0
o
040 050
4
5
0
0
9
5
? ?
cohinc heins
—
+
+
C.
7
0
0
5
669 305
6
0
6
3
i
6
6
81
80
8
37
8
30
30
19
4
4
0
19
42
3
3
0
—
—
one
—
—
Only
0
0
08 5190
8
0
—
19
5
—
—
C.
ssp.
does
specimens). pruniflorum
(7
blancoi
ssp.
ssp.
formosum
4
0
— — —
with without
in
specimens pecimens specimens
(mainland)
of of of
number number number
=
+
=
?
Locality
AsiAsiaa
S.E.
MMaal ylaya Sumatr a
Java
Java
Borneo o Cel eb ses
Philp nepsines
376).
p.
on
0
C.
sumatrnum papilae.
C.
and
specimens)
—
2
pos es
(43
arboescens
showed
specimens)
(170
—
which
+
+
conspicuo s
not
0420 040
sumatrnum ner ifolium — 21122 — —
specimen
—
are
conspicuo s epidermal papilae
the
—
??
TABLE
Phil pines papilae
from
6=—0x — — —
using
papil ae
Epidermal
maingayi
formosumformosum
in
0
+
+
? ?
of
00
2
in
sumatrnum
found
ssp.
C.
were
papil ae
sumatrnum
C.
No
of
descripton