Download Contents - Naturalis repository

115-173
27(1981)
BLUMEA
Comparative leaf anatomy
of
the Asiatic
J. Koster
Rijksherbarium,
Myristicaceae
&
P.
Leiden,
Baas
The
Netherlands
Contents
Summary
116
Introduction
116
Materials
A
survey
117
and methods
of the leaf
anatomical characters
in the Asiatic
Myristicaceae
118
The indumentum
120
The cuticle
The
118
unspecialized epidermal
Some miscellaneous
121
cells
epidermal
features
122
122
The
stomatal
The
hypodermis
123
The
mesophyll
124
The
midrib
124
The
veins
125
The
petiole
126
complex
126
Crystals
Secretory
127
structures
128
Sclereids
Generic
128
descriptions
Explanatory
128
notes
Gymnacranthera
129
Horsfieldia
131
Knema
136
Hair
140
types of,Knema
141
Myristica
Value
of the characters
Taxonomic
Introductory
Below
146
151
implications
the
remarks
151
level
151
genus
Gymnacranthera
151
Horsfieldia
152
Knema.
153
A tentative
phylogeny
of the hair
types
in
Knema
155
156
Myristica
On the
On the
Remarks
genus
level
family
on
158
level
159
161
ecology
162
Keys
Synoptical key
to the
Malesian
Synoptical key
to the
groups
species
of Knema
and
varieties
species
of
Gymnacranthera
162
163
Acknowledgements
164
References
164
VOL.
BLUMEA
116
27,
I,
No.
1981
Summary
The
hairs,
of
leaf anatomy
Knema
Horsfieldia,
the cells
of which
characters
important
proper
on
Winteraceae. Guard
and
Knema
of
an
abaxial
free
The
cells
and
Myristicaceae
bundles
of the
Many
relation
in
the
family
(e.g.
within
can
oil
the
mesic
markedly
cells. The
subsidiary
alveolar material
hitherto
layer was
a
of the
overlying
complex in
The vascular system of the midrib is
Myristica.
latter is absent
from
into
only reported for
stomatal
paracytic
are
arms
be classified further
can
cutinaceous,
such
length
by typically arranged bordering cells, leaving a
(the
and
be
separated on
Leaf
paper).
anatomy
and
Canellaceae,
cells,
order
other
many
anatomical variation
easily
star-
composed
and there
Gymnacranthera).
are
varying
be
can
leaf anatomical
used
for
characters is
species grouping
leaf anatomical characters
(see
lends
close
little
the
although
leaf
paracytic stomata)
to
support
shows
family
anatomy
a
points
table
and
of
typically
isolated
fairly
a
and
affinity
several
to
V,
Magnoliales.
of the leaf anatomical characters
to the
in
value of these
of the leaf
Annonaceae and
with
ring
of
layer
of Knema
uniseriate
characteristic,
centre.
of this
end
a
bundle
the
the four genera
at
a
have
cell and the relative
angiosperms
overarched
forming
Much
Magnolialean characters
position
and
and taxonomic
identification, and
have
often embedded in the
are
is sunken
discussed.
synoptical keys
Myristicaceae
adaxial collateral
phloem
diagnostic
analysed
of
Myristicaceae (Gymnacranthera,
Myristicaceae
The hairs
genera.
the abaxial leaf surface. In
Knema and
and
the Asiatic
of the
genera
detail.
The number of arms per
arms.
to separate
Myristica
shaped opening in
always
have
in
is described
Myristica)
Many species
different types.
the cuticle
of the four Asiatic
species
60
c.
and
ecology
of
Myristicaceae
of the
highly xeromorphic.
are
This is discussed in
family.
Introduction
In
1897
his
published
Warburg
of the
monograph
another
recognized
15
attention
the remarkable hairs, present in this
revision
to
of
(Sinclair,
In
genera.
the
four
Asiatic
1958a and
b,
1968 his work
died in
and
has
1961,
started
the
as
species.
Complementary
explore
the leaf anatomical
can
often add
affinity,
in
Solereder
publication
recorded
fragrans.
(1895)
strands
on
Asiatic genera, with
an
hairs
species
the
studied. Of
one
species
a
of the
new
has
been
in
studied the
and
it
per genus
De
and
recognized
five
(1950).
patterns
some
to
of
The leaf
can
be
Warburg's
Worsdell
petiole
83
since leaf
specimens.
foliar sclereids in
Armstrong (1975)
two
been
Wilde
worthwhile
Myristicaceae,
Chalk
midrib
of
(1908)
Myristica
species
of the
included leaf anatomy in
Horsfieldia respectively.
c.
60
species, belonging
Knema. In order
species,
and
seemed
mentioned before.
the
of Knema and
on
material has
A short characterization
Metcalfe and
in
When he
Asiatic genera for Flora
the Asiatic
scarce.
species
6
years,
a
Malesiana
Horsfieldia.
1979),
work
Flora
of taxonomic boundaries and
is
and
he
paid special
recognized
several
Wilde,
revision
leaf anatomy of
emphasis
last
(De
within
phloem
(1975c)
anatomical variation within
were
and
in the identification of sterile
1908)
(1966)
and Wilson
study comprises
Wilde's
Myristicaceae
the
their studies of several
This
1975)
revision
genus
knowledge
medullary
Rao and Chin
family. Siddiqi
new
diversity
help
(1899,
on
of
De
to our
and be of
anatomical literature
found
to
a
first
for
and about 50 in
the
available
1895)
Myristicaceae
Myristica
he
Sinclair worked towards
family.
remained unfinished. Since much
Malesiana, with Knema
natural
the
in which
Myristicaceae,
(Warburg,
1974 and
1968,
become
(Rijksherbarium, Leiden)
anatomy
of
genera
37 in Knema, 72 in
Gymnacranthera,
collected
publication
to
specimens
or
ten
gain
of
some
most
specimens
to
the four
impression
species
were
or
of leaf
varieties
studied.
J. Koster
&
P.
Baas:
Leaf
MATERIALS
The
material
reduce the chance of
in the
or
material from
of Knema
the
Rijksherbarium,
from the Kew Herbarium. To
came
leaves
but
selected,
were
the
with flowers,
specimens
material identified
De Wilde
by
,
Fully developed
sometimes
case
in
specimens present
misidentification, only
identified by Sinclair,
117
Myristicaceae
METHODS
the material of Knema korthalsii
Leiden; only
taken.
AND
from dried
taken
was
of Asiatic
anatomy
of the
study
younger material, when the hairs had been shed from the older
required
mounted in Euparal,
leaves. Permanent slides,
were
made of transverse microtome
sections of the central part of the lamina and both distal and basal end of the
and of free hand
made
and
For
were
ether and
leaf
electron
hour in
an
warm
Measurementsand
at
their widest
linear eye
measurements were
were
measured,
veins
were
of
distributed over
counted;
made
at
(Warb.) Sinclair;
Borneo:
Rossum63.
van
--
Jacobs 5259;
G.
var.
van
(Houtt.) Warb.;
14970; Ashton
&
Java:
—
Subramanian
Warb.
var.
Borneo:
a
between the veins
areas
least 7
at
on areas
(0.063
2
mm
margin
),
was
made.
were
18969;
H.
1407. —
curtisii;
Wood SAN
var.
K.
S
var.
eugeniifolia;
Borneo:
H.
—
Sanusi b.
Tahir S
cult.
BW2796; Philippines: Merrill4031. —
Robinson
Koster
235;
Warb.;
Borneo:
BW
1123;
ex.
3944;
Rosli
Warb.
bb
23779.
Theunissen
West New Guinea:
—
BW
Kuswata &
Kochummen
Toroes
Borneo:
H.
macrocoma
H. parviflora (Roxb.)
Schram
4446;
Kostermans
Malaya:
Moluccas:
& Kadim bin
Sinclair
si
12325; Sumatra:
irya (Gaertn.) Warb.;
iryaghedhi (Gaertn.) Warb.;
Sinclair;
Rahmal
paniculatai (A. DC.)
zippeliana (Miq.) Sinclair;
var.
Guinea:
G.
Haviland
7241, 13696;
Rahmat si Roepa
(Warb.) Sinclair;
—
borneensis
var.
5326, 5363;
Sumatra:
Sumatra:
coll. 6591.
King's
44045 ;
Kostermans
17514;
crassinervis
var.
Tassim
1839\
Sinclair;
10437.
van
—
H.
.H.
Royen 4510;
Soepadmo 3; Pleyle
KEP80627;
van
(Miq.)
379.
superba (Hook. /:
&
SFN 40047.
ashtonii;
23314.
—
Borneo:
K.
Sibal
ak
attenuate
Luang
Ashton
A4139, Ashton
S
BRUN
18294.
5587; Malaya:
—
S
(Hook, /
conferta (King) Warb.; Singapore:
Borneo:
Sinclair
Borneo: Jacobs
et al. SAN
Sinclair
5749;
Junghuhn 230.
West New
Ashlon
<6
FRI
Th.) Warb.;
—
Warb.;
griffithii(Warb.) Sinclair;
var.
H. sabulosa
Sinclair
ashtonii Sinclair
Murlhv
cell
guard
measured with
were
magnification
bancana; Singapore:
var.
Galau S
Miranda
bb 3081 /; Moluccas:
Th.) Warb.; Singapore:
Knema
pole
to
400
x
G. contract
Superficially studied: H. fulva (King)
Borneo:
at
11044. —
3908;
Schram
Kostermans 767.
Gjellerup 407\
vision
Sinclair
Sleenis 5274.
West New Guinea:
sylvestris
of
forbesii; Malaya:
glabra (Bl.) Warb.;
Moluccas:
pole
per leaf. The number of basal parts of hairs
G. forbesii (King) Warb.
—
paniculata; Philippines:
Java:
or
The
wax.
any
follows: The width of the
out as
from
length
eugeniifolia (A. DC.)
Horsfieldia crassifolia (Hook./
Warb.;
to remove
per leaf for both midrib and areolae
9420;
coll. 6622.
Royen 5101;
ultrasonically
and the lamina between midrib and leaf
(Miq.)
Malaya; King's
H.
shaken
were
leaf. For the lamina thickness
out
Bujang S
Whitmore FRI
—
of leaves
studied.
bancana
8035; Malaya:
59.
acetic acid. The
glacial
glycerin-jelly.
1000 magnification in the free hand sections. Ten
x
field
a
counts
9437; Anderson
134; Zehnder
var.
per
carried
half
least 7
Gymnacranthera
3393;
at
with
were
chloroform or boiled in water, alcohol
or
carried
and the
the midrib
Specimens
Billiton:
and
out
specimens
gold.
counts were
part
of all
measured, the tips of the papillae excepted. For both characters
surface
a
ether, alcohol
carried
were
petiole
safranin-
a
for the cuticle thickness the thickest parts of the cuticle between the
measurements were
with
mounted in
microscopy fragments
micrometer
piece
tests
preparations
hydrogen peroxyde 30%
coated with
were
lignin
acid. Cuticular
stained with
were
rinsed in fresh dissolvent in order
subsequently
fragments
pairs
specimens
some
stained with Sudan IV and
scanning
for half
For
hydrochloric
volumes of
using equal
cuticles
sections. The sections
paradermal
mixture.
haematoxylin
phloroglucinol
was
indumentum
K. furfuracea
22841\
&
Sinclair
Whitmore
(Hook./.
Th.)
var.
cinnamomea
Warb.; India,
SFN 40046.
FRI
&
12326.
Th.)
K.
—
—
Warb.;
K.
de Wilde:
Peninsular:
curtisii
elmeri
Malaya:
(King)
Merr.;
Kadim &
BLUMEA
VOL. 27.
-
No.
1, 1981
118
—
Mahmud 22.
K.
2488.
K.
Kochummen
Malaya:
Borneo:
Chew
de
(Sinclair)
mandarahan
K.
-
oblongata; Borneo:
Java:
Ja.
Mikil
—
2502.
K.
S 18478. —
Sumatra:
Warb.;
24985;
K.
Java:
—
Dransfield
3137F
K.K. laurina
Witde-
de
Wilde;
linguiformis (Sinclair)
Plants
Warb.;
malayana
(Bl.)
Sumatra: de Wilde & de
Assam:
Steenis
van
—
Chai S 34822.
K.
K. hookeriana
—
Warb. ssp. kunstleri;
(King)
SAN
FRI 2431:
12308.
(Bl.) Warb.;
K. kunstleri
linifolia (Roxb.) Warb.; India,
(Miq.)
Bakar SA N
—
143; Kochummen
41273;
<£ Smitinand
K. intermedia
Borneo:
latifolia Warb.;
SAN
Hansen
12207. —
Cenabra FB 29973.
Borneo: Chai
Wilde;
Thailand:
Lorzing
Wee Lek
K.
—
K.
—
Ampuria
Paie S 27799.
llias
lunduensis
FRI 2252.
Borneo:
Duyfjes 14429;
Warb.;
Sumatra:
Philippines:
laurina; Malaya:
var.
40714.
Th.) Warb.;
korthalsii
i Warb.;
—
Warb.
globularia (Lamk.)
&
(Wall, ex Hook./.
10092.
of Assam
—
Merr.
oblongata
K.
SFN
Singapore: Sinclair
K.
—
6188.
ssp.
Chelliah
oblongifolia (King)Warb.; Malaya:
KEP 98224: Whitmore FRI0111.
K. pallens
;de Wilde; Borneo: Meijer 2533: Lantoh SAN 73312. —
K.
Sinclair
percoriacea
Sinclair;
Wilde;
Borneo:
Kostermans
-K.
forma sarawakensis
Malaya: Carrick
Anderson
S
5103; Malaya:
rubens(Sinclair) de
13963.
1510.
I.:
24236.
Houtt.
M. fatua
Koster
Guinea:
—
787.
Beguin
Mikil
Corner
Othman bin
Sinclair
12890.
BW 1016.
M.
M.
BRUN
Corner
32840).
M.
—
5365;
Warb.;
12547.
SURVEY
908.
M. subalulata
Kostermans
Borneo:
A
—
Miq.;
Sinclair
de
Borneo:
9908.
9907,
Unyong
S
New
Soepadmo 57;
—
Sulit
M.
lepidola
Bl.;
1289.
East
A.
New Guinea:
New
s.n
Guinea:
121
Lundquist
Sumatra:
Warb.;
West
Smith;
Floyd
Vriese
37177; Kostermans 4873;
Guinea:
M. maxima
C.
New
de
Sinanggul SAN 39995;
7; Borneo:
West New
—
SAN
Markgraf;
papuana
Moluccas:
SAN
Brass
15066; Singh
globosa Warb.; Papua
Kostermans UNESCO
M.
var.
15023;
SAN32924; Aganb. Ambulah
I.:
Guinea, Fergusson
Wood SAN
Borneo:
New
& Havel NGF
Brass
Guinea:
7446.
(=bb
Thorenaar
M.
—
&
villosa
Wood SAN A4747.
6438;
OF THE
s.n.
sphaerosperma
M.
—
plumulosa
Warb.; Borneo: Asah anak
Sinclair;
King;
Kusxvata &
Scortechini
-
5056.
rufa
K.
—.K. psilantha
pulchra (Miq.) Warb.;
(King) Warb.; Malaya:
K.
Houtt.; Philippines:
Java:
AbanGibot
1403.
4197.
Moluccas:
fragrans
FRI
Malaya:
—
MS.
Thailand:
& Geesink c.s. 4747.
van Beusekom
—K.
maingayi Hook./; Malaya: Maingay
(Endert) 55E1P624; Grashoff
Versteegh
Vogel
guatteriifoliaA. DC.;
SAN30298; Elmer 20850;
Wilde;
M. cinnamomea
fatua;
—
retusa
S 23645.
Luang
K.
S 29064.
entrecasteauxensis
var.
i
King; Malaya:
gigantea
M.
—
de
—K.
Sibat ak
Wilde; Malaya: Shah
MS. 1312.
setosa de
Brass 25893.
var.
Borneo:
Haron
SFN 29015.
Moluccas:
chrysophylla
25986; Normanby
—
Wilde;
Wilde; Malaya: Shah
(Miq.) Warb.;
Myristica
Carr
14373;
K. tenuinerviar de Wilde ssp.
tomentella
de
K. pseudolaurinai de
—
LEAF ANATOMICAL CHARACTERS
IN THE ASIATIC
MYRISTICACEAE
The
indumentum (figs.
Hairs
present,
are
Horsfieldia, Knema,
abaxial surface
frequent
on
Myristica
(in
at
and
on
frequent
most
in
both
1 —10,
19, 21, 25)
the adaxial and abaxial surface
on
examined of
and
sides in
the
on
leaves,
species
most
Gymnacranthera
the midrib
,
1—3, photos
least in young
some
or
Myristica species),
Horsfieldia
lamina in
Myristica)
and
only
usually
Knema and
Gymnacranthera
on
and
(in
the
most
adaxially
in
abaxially
in
Myristica.
The indumentum
hair type:
words,
each,
a
a
hair is
length
in
In
composed
two,
one or
excepted.
appears very
rarely
of
Gymnacranthera
The
appearance of
a
the
and
uniseriate
consisting
of
a
few cells
consisting
of
numerous
The
cells, having
the cells have
may
and the
length
tall
a
'body'
hair,
which
which will be discussed
are
occur.
sometimes
hairs
as
or two
of the
one arm
seen
and
is
basic
arms
from
same
(fig. 2,3;
above,
the
Knema, the walls
variable. Short
to
filiform,
In Knema the
separately.
a
In other
of each cell very small; this may
(most typical for Gymnacranthera)
cells,
one
two arms, not
Horsfieldia
of the
1895).
the so-called stalk cells,
and Knema have
give
In
recognize
one can
(Warburg,
epidermis,
Horsfieldia
arms
oblique
nature.
to
the
(e.g. photo 7).
250 /tm per cell in Knema, may
forms,
Myristica
the
arrangement of
stellate hair
of short
nearest
the cells in
between the cells
may be very
obscure
cells
per genus, but
uniseriate trichome
one row
more
Gymnacranthera ;
photo 4, 9).
different
sympodially branched,
with
species
hairs,
extremely long hairs,
a
show
length of
a
about
range of hair
In older leaves the hairs have often been
J. Koster & P. Baas:
Fig.
1.
deciduous
part
Diagram
and
of the
lucida
Fig.
hair);
drawing
3.
g
See
=
Myristicaceae
in
of the
text
the
hair. —
hair;
e
=
a
surface
view
different
for definitions
=
arm;
cutinized
unspecialized epidermal
of hair
Diagram
derivations.
of
persistent part of
cells.
b
=
ring;
—
body;
f
=
Fig.
types
of different
in
Knema.
types.
—
c
=
stalk
basal cells
2.
(e
cell;
of
arms
Arrows
d
=
line
and f form
Gymnacranthera
showing unequal length
hair
119
Leaf anatomy of Asiatic Myristicaceae
of hair
indicate
separating
together the
contracta.
cells,
x
—
the
basal
Camera
400.
tentative
phylogenetic
120
shed, but
of this
basal
a
small
cuticular wall
elevated.
to
cells' for
epidermal
The hair index
specimen
in
or even
but
variable,
studied. It is
at
of
fact
(in
it
is
essential
know
When necessary
not.
or
low power
with
magnification
cuticle
After
structured
11 —18,
(photos
in
boiling
this has
An identical structure, referred
Winteraceae
In the Asiatic
Bongers (1973).
species
examined of
material
usually
the
overlying
it
is
only
Knema
slightly present
and
stomatal
species
a
this
to
positively
with
acetolysis
it
of both
acid
IV.
the leaf
warm
material,
chloroform,
the
and the
cells
the
on
cells
Horsfieldia sylvestris
sunken
alveolar
by
of the
but in Knema
present
in
12);
cells. The
have
studied
most
When present the
guard cells,
margin, (photo 28),
(photo
a
has been recorded
thoroughly
was
only conspicuously
stomata
not
cells in
subsidiary
material
(see
under 'The
(photo
and
From
sulfuric acid.
(used
the
porous
16).
structured
species
14)
layer
is
or
in
Besides the
alveolar
not as
cuticle
(but
present
in
some
21)
and
a
erect
warts
present
Asiatic
(in
material,
most
Horsfieldia
and
15)
11),
in
13),
press)
for
of
Myristica
but
also present
the abaxial surface in
the
and
to
this
cutinaceous.
(photo
are
stained
moreover
solvents
organic
9:1
was
species
a
it is
dense
more
with rod-
in this genus. An
Horsfieldia iryaghedhi
a more elaborate
description
of the
Myristicaceae.
there may
layer overlying
species,
is
a
alveolar material
and Knema is composed
occur
(photo
on
to
plates (photo
plates
The
palynology)
resistance
composition
with
See Kosterand Baas
alveolar material in the
and Knema has been boiled in
Gymnacranthera
concluded that the
irregularly
structures
(photo
is
in
the abaxial side in many
on
Myristica species.
alveolar material in Gymnacranthera
irregular,
the
by studying
because the absence
this material is absent in
some
subsidiary
do
treatment
coarse
variable in appearance:
(photo
it
anhydride
Sudan
was
reticulum of
like
are
fully
complex').
mixture of acetic
The
the
on
alveolar
epidermis except
cells of the
usually
Myristica
One leafof
resistent
and in
plumulosa
subsidiary
in
have persisted
been checked
(1944), and
Myristicaceae
the whole
Knema
the
on
rough
a
leaves
fully developed
shaking
or
to as
Nast
midrib, the major veins and
intermedia and
bordering
and
Horsfieldia
covers
use.
per surface unit is
microtechnical artefacts.
to
ether,
or
for the
Bailey
slightly
patterns of
nevertheless
gives
that
the cuticle proper remains
species.
by
small
one
21, 28)
water, alcohol
layer, overlying
hairs)
whether hairs
of hairs from anatomical slides may be due
The
and 'oval
dissecting microscope
a
this
Myristica
these patterns with the hairs.
determine and
density, provided
to
and
oval, subtend
basal cells may be
basal parts of the
to
epidermis
and is therefore of little taxonomic
1965),
easy
the
may be very variable in leaves of one
density
remaining
more
course
leaves
developed
leaves
leaf(Stace,
one
not connect
on
(in Horsfieldia
or
and
described 'circular'
did
of hair
ring
to numerous
fig. 1). These
have
epidermis
cells. The upright walls
cutinized
a
Gymnacranthera
25 and
position (photo
of the difference in hair
impression
in
Usually
as
the
nearest
characteristic circle
a
Armstrong (1975)
The number of hairs
stalk cells
Two
(fig. 1).
in
cells, arranged
a measure
1981
1,
unspecialized epidermal
Horsfieldia, but
as
No.
of the hair remain
(photo 21).
circular cell takes this
27,
one or two
have the
as
of the hair has been shed
rest
Knema) mainly
sunken
left:
always
walls,
proximal part
most
after the
is
part
cutinized
strongly
also
VOL.
BLUMEA
be
the cuticle
abaxially
in
faintly adaxially
in
conspicuously
sylvestris,
other types
of
sculpturing
of the
proper). Striations (fine ridges)
Horsfieldia
some
are
crassifolia
Knema
species.
J.
Kostkr
Grooves
corresponding
present
in
& P.
Baas:
with the anticlinal walls
Horsfieldia,
Knema
sculpturings
of the cuticle
are
that
only clearly distinguish ridges,
one can
electron
scanning
adaxially.
way
The
surface
In this
because of the
The cuticle
although
a
The
1
in
or
been studied.
the
Only
of the vein is
course
In
epidermal cells).
are
the
on
is
very
major
midrib and the
areas
veins
indeed
thickness all
less
in
striking
straight,
proved
over.
in
nearly
all
are
are
perpendicular
rather than
perceptible
to
square
the
are
thin
some
the
cell
(photos
in
fairly
study,
view.
of the veins.
course
and
straight
to
they
on
Adaxially they
have
not
is when
(that
the minor veins
epidermis only;
are
in Knema
adaxially.
curved
are
usually
lower focus
at
less sinuous. Thin
are
cells
long
the overlying
pattern of
Myristica
the adaxial surface
slightly
The
with the
Venation patterns
of the abaxial
visible
faintly
constant,
13, 16, 20, 22, 30, 34)
surface
rectangular,
to
rectangular.
pattern
flanges
areas
are
areas
of
species
where
In
present.
of the cuticle
areas
areas
of
more or
(photo 22);
of cuticle of
of the cuticular flanges
frequencies
veins and
the
and
leaf
the midrib and the
transverse
the
only:
the
even on
some
and
Gymnacranthera
thin
but in low
major
this
(>Gymnacranthera
species
of minor veins has been recorded
veins
be thin
Pitting
in
present in the loops of the undulations. This character
species
so.
is
species
in the macerations in the
and
major
light
occur.
On the abaxial surface the cuticular
specimens,
the midrib,
on
to
these
inconspicuously
to
Gymnacranthera (photos 22,30),
prominent
In
sinuous.
found
veins
major
prominence
varying conspicuousness
the
sections.
considered
one
cells
focus and
high
at
of
specimens
polygonal
prominent abaxially
in
obscures this feature in
transverse
been
not
cells
epidermal
the abaxial surface,
granular
appear
present,
epidermal
In the macerations the cuticular
less sinuous
has
Gymnacranthera, Horsfieldia
visible in the
slightly
cells
may
when
The thickness per
two
epidermal
polygonal
on
of the alveolar material, varies between 2—18
/rm
of 9 /im has been
some
be
to
with the
sculpturings
of this material.
abaxially.
yum
1965)
material,
feature,
nature
the midrib and the
tend
18). It should be noted
have been examined this
specimens
be examined in the
must
difference between
parallel
they
11
—
unspecialized
usually
the
it
important
ephemeral
abaxially)
overlying
axis
case
an
unspecialized
The
Stace,
thickness, exclusive
and
contracta,
(sensu
convex
of the cuticle.
sculpturings
layer
although
adaxially
all
rarely
are
Adaxial
17).
and other
grooves
not
cells
epidermal
(photo
present in Knema rufa (photo
smooth. The alveolar
or
view.
Wax,
of the
Myristica
Moreover, alveolar material, when present
cuticular
microscope
and
microscope. However,
may obscure other
121
Leaf anatomy of Asiatic Myristicaceae
outer
flanges
are
mostly
Horsfieldia they
cuticle
be
may
(cf. Bongers,
where the
same
more or
are
slightly
present,
1973)
least
inconspicuous,
margin,
sections these
wall has the
but
is present in
inconspicuous
flanges
are
most
straight.
In
to
transverse
rectangular,
The abaxial
(photo 34).
the
sections the adaxial
on
unspecialized
conferta
they
unspecialized epidermal
are
epidermal
The cuticle of these cells is
measurements
Abaxial
the midrib
often
cells
are
slightly
usually
thickened
more
on
and Knema
occur
in
linifolia),
many
in
species
of
Knema
Horsfieldia iryaghedhi
in
are
less
or
usually
square
Gymnacranthera).
dome-shaped
the top of the dome. For
of the cuticle thickness these thickened
papillae
cells
dome-shaped (not
parts
have been taken.
in Knema
(most
striking
and in
Myristica gigantea
122
and
Myristica maingayi (photos
and Knema, where the
papillae
the
Myristica
cuticle
may
Often
and
in
and surrounded
occur
on
veins
the adaxial
and
is
or
properties,
In
the
less
are
and
are
However,
wall has
whereas in
strongly
thickened
the
(photos 28—31)
hole in
or a
sections
to
on
the
cells.
They
the
appear
may
major
large
as
situated in
extent
(in
are
of the
centre
midrib,
the cells
large
a
the
about circular in
epidermal
mostly
cells
epidermal
only
relatively small,
restricted surface
a
this
case
the
the cell is
cells. In view of
surrounding epidermal
present of several cells,
are
thin cuticular
relatively
narrow
to
are
they
epidermal cells,
and short cuticular
are
sharply delimited,
not
flanges (photo
30).
From
connected with this feature. The
in
Myristica adaxially,
which
are
large
flattened, having
flanges (photo 31).
and
Gymnacranthera
The
areas
Horsfieldia
absent in Knema.
each specimen examined, but usually in small numbers.
present in
distributed and
irregularly
more or
distribution,
in the Knema
Wilde refers
to
these cork
Knema
sabulosa,
less circular structures, identical
frequent
most
are
their origin is
shaped,
Horsfieldia
Horsfieldia glabra,
structures
Horsfieldia
dome-shaped
strongly,
a
four genera
oriented
radially
transverse
adjacent
and
abaxially,
kunstleri abound in
These
with
in
present
warts are
they
areas
idioblasts are
vary
have
features
all the
sunken and
by
forms of
probably secretory.
are
thin cuticle and
relatively
mentioned
Since
outer
covered
partly
the cells
deeply and
crystalliferous
Cork
In
not
cells
the abaxial surface,
margin.
The
cell does
usually
are
mentioned of
20).
sections it appears that druses
transverse
adaxially
as
species
thin cuticle
a
1981
as extreme
epidermal
less
epidermis,
macerations
many
staining
more or
well
as
mesophyll (photo 29).
pear-shaped)
by
the leaf
along
empty idioblasts in
these
single
over
with
These cells
(photo 28).
shape
or
I,
In the
epidermal
distributed
species
thin cuticle
present with a very
cell surface
one
top of the papillae (photo
miscellaneous
Some
No.
be considered
mentioned the
species
the
on
VOL. 27.
13, 16, 20).
cells, the thickness of the cuticle of
a
-
BLUMEA
the
on
species mostly
1979)
and
and Knema
regular
to
surface,
cork
having
a
warts.
regular
the midrib and the veins. De
along
and
on
(De Wilde,
warts
abaxial
traumatic.
probably
conferta
uses
them
as a
lens character
of basal cells
separate a group of species from others. In Horsfieldia glabra groups
to
of hairs
hairs
on
the
probably
are
in the leaf
early
of
origin
ontogeny,
of the cork
most
but basal
parts
the adaxial surface the groups of basal cells
small
sometimes
cells;
surface
relatively
abound.
Perhaps
development
frequently
some
and
occurs
The
bancana
veins
a
are
are
var.
basal
finally
on
a
cells
cork
wart
cell
is
become
a
to
the
the leaf
shows abnormalities.
a
is
is
not
from
13,
species
the
whereas cork
course
Why
they
are
leaf
this suberization
understood. Moreover there is
a
are
stomatal
Sometimes
complex.
recognizable
in
a
cork
wart.
23—27)
epidermis;
abundant on the
If
warts
of the
however,
in
Gymnacranthera
very small number is also present in the adaxial
margin.
leaves
numerous
On the abaxial
recognizable.
in
looses its
fully developed
and consist of
suberized.
are
part of it)
(photos
This
rarely present,
subdivided
only
abaxial
large
are
wart
are
developed
pair (or
complex
confined
borneensis
along
cork
the walls of these cells
randomly distributed,
and
to a
the abaxial surface
guard
stomatal
Stomata
They
the
evidence that
the outlines of
transition
a
small basal parts of hairs
warts.
remain. In the
areolae, scanty
on
the
epidermis.
midrib,
the
present here, the stomatal complex
J. Koster & P.
Several
authors
variable within
Baas,
1973).
The
39 f.im for the
but
the
cell
guard
and
a
of
type
in
little variation in
some
not
are
range from 8
and
a
is
paracytic
Knema
Therefore, the
genus.
specimens of Gymnacranthera
study.
27).
short
single
one
leaf
stomata
be up
can
to
are
to
8
the average values show
data for
the
stomatal sizes
per
have been recorded for
and Gymnacranthera forbesii
contracta
The
(in Myristica relatively
relatively
stomata
water
or
(photo
21 fim for the width and from 15
to
length respectively. However,
in tables. Giant
given
extremely
been determined in this
not
The variation in
occurs.
be
may
1973; Van Staveren &
different size range
a
Gymnacranthera
large overlap
stomatal index
examined
species
pairs
and 12 /<m for the width and
specimen
the
123
Myristicaceae
1973; Jansen & Baas,
The genera have
length.
stomata
fairly
that
out
Asiatic
oj
anatomy
Stomatal indices have therefore
stomatal
present),
pointed
Lea]
species (Bongers,
a
dimensionsof the
long
have
Baas:
var.
forbesii only.
The
guard
cells
cells
subsidiary
fully
Gymnacranthera
often embedded in the
are
and
Gymnacranthera
in
Knema; here,
enclose the
guard cells,
such sections also show
subsidiary cells,
section
a
the
Outer
rectangular.
stomatal
ledges
are
the
to
the
pore,
In
protruding subsidiary
lowly dome-shaped
well
in
conspicuously
excepted (photo 24).
sometimes
are
fairly
sides
poral
pointed,
cells, whereas the unspecialized epidermal cells
most
perpendicular
in
developed
or
square
to
species
of
complex
in
some
Horsfieldia only.
A
peculiar
Knema and
Knema
character is formed
Myristica.
species
the
stomatal
level than the other
cells
transverse
cells
characteristic
cells. A
sections show the
cuticle, overarched by
epidermal
more or
(photo
form.
structure' is formed
24);
Siddiqi
11
to
varying
in
stated
As
less
height
per
species,
the
elsewhere,
directed
(1975c)
cells. This
papillae
papillae
to
on
a
by
(photo 26).
less
In surface
above the stomatal
unspecialized epidermal
of the
(photo
cells in
a
thin
bordering
the
star
its
'star-shaped
the
be incorrect. In
to
more or
lower
epidermal
6
stoma
give
that
proposed
appears
overarched
ring
4
by
of the
position
nipple-shaped
cells
form a
left
sunken stomatal complex with
horizontally
partly
bordering epidermal
indicates the
Wilson
by hypodermal
observe, lying
to
star-shaped opening,
strongly
these
and
the sunken stomatal complex is
the 4
the stomatal
surrounding
itself is difficult
complex
epidermal
the stomatal complex,
overarching
The
23).
the cells
by
In the cuticular macerations and the free hand sections of the
Myristica
upright papillae
view these
of
papillae,
complex (photo 13,25).
most
species
are
without
papillae.
The
hypodermis
An adaxial
the
epidermis,
some
only
Knema
In
has
and
developed,
with the
Horsfieldia
as
a
continuous
been recorded for
species
locally
intergrades
(photo 32)
hypodermis
specimens
especially
are
cells of the
adjacent
to
tissue
Myristica
also
of
and
Myristica species.
Myristica guatteriifolia
near
the
midrib
the
occur,
hypodermis
occur
in the
and
have
the
periclinal
hypodermis,
the adaxial and sometimes
elevations of the leaf surface
ground
of
of translucent cells, distinct from
Horsfieldia
the
hypodermis
major
In
is
veins, and
palisade parenchyma.
some
genus swollen cells with thin walls
these cells
layer
some
(photo 32), and/or
midrib and
sometimes
the
petiole,
probably
are
to
or
the abaxial
present
where
in the
these
of traumatic
division walls. In this
few-layered
periphery
cells
origin.
groups of
epidermis,
in
in
of the
Knema and
BLUMEA
124
An
abaxial
inconspicuous
27, No.
VOL.
1.
1981
is present in
hypodermis
Knema
some
and in
species
Myristica sphaerosperma.
The
mesophyll
The
cells of the
virtually
four adaxial
to
with the
intergrades
in
range from
a
have
i.e.
lignified;
one
sclerified
Either the cells
possibilities.
walls
birefringent
spongy tissue is
(in
some
of
Knema and
Gymnacranthera,
(secondary
in
the
only
of
walls in
the
adjacent
to
N
o
t
Myristica)
the cell
sectioning
thickness
because
epidermis,
This
characters
thickness of
robust.
The
Horsfieldia,
there is
species
midrib
a
with
is
abaxially
Myristica
The vascular system is
phloem
often
an
sections
(in
the whole
or
species
some
In
of
cells have
palisade parenchyma
Knema mandarahan
large pits (in
strongly birefringent
the leaf
margin
the
cells,
the
some
in
soon, and for the transverse
domestic bleach
of bad
to be
lamina
quality
thickness
reservation
domestic
are
damaged
made
and the thickness
not in
(and
or
and
the
in the
study
of midrib
of
to
Horsfieldia specimens only,
bleach).
of lamina 3 and less:
quite
remarkable.
abaxial
arc-shaped
by
(in
some
in the
and
adaxially
and
there is
a
two
are:
more:
raised
in
Myristica species
and
some
the adaxial surface.
on
sometimes
in
more or
less straight
joined together.
One
layers.
to
The
numerous
tissue between the main bundles,
ground
elements
xylem
Generally
bundle,
ratio 6
Gymnacranthera
to concave
often
strands,
Horsfieldia
but in
less flat
more or
interspersed
are
accompanied
well
fine;
and
raised,
prominently
as
ridge adaxially),
in separate
arranged
bundles
range of
studied is fine to robust; criteria for these characters
the midrib is
adaxial bundle and
is
whole
a
and Knema)
the dried leaves break
leaves;
behaviour
raised central
Horsfieldia species
phloem
in
without cells with
expand completely in
midrib/thickness
Knema and
only
and
(figs. 4—7)
The midrib of the
ratio
the
Horsfieldia).
consequence,
swelling
layer
few leaves demonstrated to
a
mesophyll
with such cells
or even
in
have to be taken with
in
layer
isobilateral leaf. The
an
width ratio per
to
three,
to
abaxial
have sclerified walls.
the transverse
difficult. In
of lamina ratio
midrib
caused
to
walls in
the
extents
species
have brittle
walls cannot be made
of the variation
The
often
Horsfieldia species
process.
anatomical
also
are
mesophyll (particularly
the
Most
e:
sections
There
pallens).
two
most
abaxial layers of the spongy tissue have
thick, strongly birefringent walls, supplied
and Knema
length
Gymnacranthera
variable
to
sabulosa has
different
a
occurs
species
interspersed
the
five.
to
cells)
mostly
palisade parenchyma,
Horsfieldia
of cell walls
Strong birefringence
be
tissue.
spongy
of
with
dorsiventral,
always
layers
palisade parenchyma
species,
per
is
mesophyll
sometimes up
(fig. 4);
these
situated
centrally
phloem
bundles have been mentioned before by Worsdell (1908) for Myristica fragrans, by
Siddiqi
and
Wilson
Horsfieldia species.
and
In
Myristica.
pith (fig. 5);
in
(1975c)
for
some
some
strongly interrupted,
sclerenchyma
(figs.
often
4
—
7);
even
Knema
or
there
(fig. 7).
fibres,
there is
species
are
a
and
by
Armstrong
is present in
complete
which
centre
phloem
often
occur
of the
(1975)
Horsjieldia,
for
Knema
collateral bundle in the
the
adaxial bundle is
several small bundles (fig. 6). In Gymnacranthera
The whole
fibre groups also
in the
species
Horsfieldia and Myristica specimens
the adaxial bundle is absent, but
described above
Knema
This type of vascular system
bundles
occur
vascular system
are
associated
in the
phloem
centre
bundles
in the
centre
of the midrib
is surrounded
with
peripheral
by
groups
phloem
as
of
groups
of the midrib, associated with and
(figs.
4
—
7).
J. Koster &
Figs.
8:
7.
4
Gymnacranthera
basal
part
The
Baas: Leal
9. Vascular patterns in midrib and
—
— 4. Knema
16.
x
P.
of
laurina, midrib. —
Lines:
—
is
often
black:
of Asiatic Myristicaceae
Camera lucida
midrib.
lunduensis,
8. Knema
xylem;
tissue
ground
collenchymatous,
petiole.
5. Knema
contracta, midrib.
petiole.
anatomy
from
centre
interspersed
to
white:
4, 6, 7, and 9: x 40; 5 and
Horsfieldia glabra, midrib. —
drawings.
6.
—
linifolia, distal part
sclerenchyma;
of
petiole.
—
9.
Myristica lepidota,
phloem.
periphery
to
various
125
extent
parenchymatous
cells with
with
walls, especially in Gymnacranthera and Knema, in the former mostly abaxially;
these
genera there
midrib. Often the
one- to
of
collenchyma
Gymnacranthera
sclerified
The
The
flat
often several
or
cells,
veins
major
or
the
if
and
midrib,
layers
of these cells
at
the sclerified cells appear
several-layered hypodermis
continuous in
or
are
on
periphery
the
as
a
in
rarely
situated
abaxially
Knema
of the
in
of the
locally developed
the adaxial side of the midrib. In
Myristica,
to
sclerified
adaxial
specimens
chlorenchyma
,
collenchyma
or
is
the layers of
present.
(photos 33, 34)
veins
are
adaxially
raised; abaxially they
either
are
slightly depressed (not
always
raised.
in
Gymnacranthera),
126
The veins
veins
and
a
caps
by
constituting
one
number of Knema
sheath,
which
abaxial
more
less
or
well-developed
two
of
row
bundle sheath.
be continuous
and
A
sclerenchyma
fibres
often have
vertically
either the adaxial
birefrigent
or
walls and
the
veins,
transcurrent
in
are
caps
each
at
thus
side,
differentiated
usually poorly
sclerenchyma.
the abaxial
the cells of these bundle sheath extensions
epidermis
have
Myristica
(little developed
In
a
fairly well-developed parenchymatous bundle
a
to
major
intermediate between that of
Horsfieldia
continuous
there is
species
may
(photo 33);
from these
the
Knema
the
sclerenchyma fibres;
bundle sheath surrounds the bundle and the
parenchymatous
both
1. 1981
system,
Gymnacranthera,
slerenchymatous
a
No.
the adaxial and abaxial side of the bundle
In
Gymnacranthera).
connected
veins.
at
vascular
complex
more
minor
sclerenchyma
VOL. 27.
with collateral bundles and
supplied
are
have
may
midrib
or
-
BLUMEA
veins
are
are
epidermis
adjacent
or to
the adaxial
to
probably lignified. Apart
embedded in
usually
the
mesophyll.
Another character recorded for Knema
outline,
fibres
sclerenchyma
these
and
position,
in the
places
distribution
sometimes
only;
transverse
section
homologous with
minor veins.
The
(figs. 8, 9)
petiole
The
vascular
system
vascular system of the
Myristica)
the
of the
midrib,
adaxial
the
petiole
phloem
consists of three
bundles
is
sclerenchyma
Brachy-
more or
Crystals
idioblasts,
in
the
a
have
and
Knema
fig.
second
and the
are
crystal
tissue
10g),
of
found
of
consisting
is
may
with
comparable
mostly
the
Knema and
three
to
up
at
be considered
(in Horsfieldia,
bundles,
large
vascular system of the basal end of
collateral
bundles,
also with free
9);
the
the abaxial sides. Sometimes
to
in
(not
these
close and
Gymnacranthera),
transverse
interspersed
in the
mentioned
sections
ground
were not
from
tissue of the
petiole.
occur
of the
in the
by
midrib
(i.e.
'Some
mesophyll
restricted
in
enlarged
miscellaneous
of
species,
most
the
to
palisade
ordinary parenchyma
midrib and in
is
occurrence
(fig.
10a).
Metcalfe and Chalk
exists
spindle-shaped
are
druses
epidermal
cells of
cells;
these
Gymnacranthera.
have been found in
the
is
occurrence
may
tissue
type of frequent
the
Large
under
been found in the
their
(fig. lOh)
ground
before
They
particles
also present in the
(1950).
epidermal
are
cells
represented
in cells of the
bundle sheath of the veins. Most
mentioned
that
more
in the leaves studied.
specimens they
when druses
structures
petiole
usually minute, particles, grouped
parenchymatous
crystals
of
probably
often
been found in
not
spindle-shaped,
ground
of the
bundles,
few fibres may be present
petiole.
Myristica
Small druses
mesophyll
A
These
less arc-shaped
been
large diversity
31,
photo
druses have
or a
(photo 31, fig. 10)
show
parenchyma.
vein
the appearance of bicollateralbundles (fig.
epidermal features', have frequently
but
be mentioned: structures, in
to
confined for the greater part
usually
astrosclereids
Crystals
(photo 34).
(fig. 8). Generally the
basal end of the
to
one row
distal end
adaxially, giving
small collateral bundles
extreme
has
but often the abaxial and
adaxial to the main bundles. Most
the
only
consists
part
bundles for the abaxial part
only
unlike the
not
are
most
probably
In
as
some
by
more
mesophyll
abundant
these
are
or
less
and the
in
the
the acicular
Knema and
Myristica
well, but here the possibillity
fragments of
epidermis. Relatively
small
druses, especially
large spindle-
to
needle-
J. Koster & P.
Fig.
10.
Crystal types.
cell of bundle
sheath
(Myristica fatua
fragrans).
fragrans).
in
crystals
—
—
e.
h. small
Flattened,
specimens
more
in
species
of
to
a.
spindle-shaped crystals
needle-shaped crystals
in
prismatic crystal
(corroded?) edges
epidermal
cells
cells of
abaxial
tissue
some
druse in
big
in
parenchyma
in basal cells of a hair
cell
epidermal
in abaxial
(Myristica
epidermal cells (Myristica
(Myristicafragrans).
g.
—
127
Myrislkaceae
—
f.
mesophyll
irregular,
idioblast
small
(Knema
(Knema hookeriana).
Myristica specimens, especially
in
10b).
prismatic
crystals
bancana and
var.
the cells
crystals,
in abaxial
occur
Myristica have
crystals
with four
to
smaller
than the
solitary
makes, together
(fig. lOd).
630.
cell of spongy
ordinary
less
or
x
of Asiatic
of the
in
occur
several
parenchymatous
cells
epidermal
Myristica species
in
or,
bundle sheath,
in
some
mostly
10c).
small
Irregular
abaxial
epidermal
bancana
all
spindle-
(Knema hookeriana).
(fig.
of Knema, in
solitary (fig.
clustered,
cell
druse in
occur
Gymnacranthera
in
crystals
the basal cells of hairs
b.
flattened
c.
anatomy
drawings,
with rounded
crystals
epidermal
shaped crystals
—
fatua).
var.
clustered
—
lucida
Leaf
—
( Knema hookeriana).
d. flattened
abaxial
hookeriana).
Camera
Baas:
six rounded
study
if
edges;
Often
ones.
edges,
rounded
grouped,
cells in Knema hookeriana
peculiar crystals
also
with the rounded
Further
with
usually
epidermal
more
they
edges,
in
the epidermal
than
have
a
one
in
a
sometimes
lOf).
cells:
cell, they
Some
flattened
usually
are
corroded appearance, which
composition
a
(fig.
of calcium oxalate doubtful
would be necessary to determine the
exact nature
of these
crystals.
Druse-like
composed
clustered
of
crystals
relatively
few,
Intermediate forms have
been
edges
and
have been
prismatic crystals.
reported
Secretory
The
large,
to
enable
or
have been found in the
less
in
Myristica fragrans,
irregularly
found of clustered
shaped
sex
distinction of
but
crystals
unlike
particles
crystals, crystals
The different types of
structures
more
occur
large,
in
(fig.
druses
lOe).
with rounded
Myristica fragrans
seedlings by Nayar
et
al.
(1977).
(photos 34, 35)
spherical
mesophyll
oil
cells,
and the
as
mentioned in the
ground
older literature,
tissue of the midrib in
most
of the
128
BLUMEA
in great
specimens studied, usually
There
Warburg's
remark
indications
(colour
to
contrary
are
of
contents
these
West
mucilaginous.
large
cells
idioblasts of
the
1981
1,
that
and
they
also in
staining
tannin-like
that
properties)
(in
or
in
occurring
and for
mutually
the
irregular
confirmed in this
in
strictly
that
fact,
some
and
not
somewhat
study.
studied
leaves
peculiar
(photo
sometimes
be
Metcalfe
in the
protruding
even
the so-called
and
tanniniferous
Chalk
In the
not
S
been
c
le
r
e
id
Sclereids
in Knema and
they
often present in
Myristica
as
the
to
which
Astrosclereids
varying
they
occur
is in
amounts as
clustered
are
the
The
traces...'
for
study,
been
of the vein
epidermis,
structures must
according
latter
some
these
found
to
describes
Horsfieldia
have
structures
for
astrosclereids
(rarely
tissue of the midrib,
(photo 37).
recorded
tissue of the
findings
and
for
Gymnacranthera,
midrib,
most
in
two
Horsfieldia
Horsfieldia superba
frequent
in the spongy tissue
of Rao and Chin
abundantly
fulva
to
ground
high frequency
with the
Horsfieldia
finding
a
brachy-
in the
have been
ground
with
contrast
have
accordance with the
in
mesophyll,
examined:
superficially
vascular
caps
the veins
(1975).
examined in this
However,
Myristica:
adaxial
these
be
cell
followed and
specimens.
the
situated in
are
brachysclereids only)
sometimes
frequently
(photo 38),
towards
not
identify
of Knema and
specimens
Armstrong
rarely branched sclereids
and in the
adaxially,
staining technique
herbarium
could
to
observed.
mostly abaxially;
where
literature,
(photos 36—38)
s
are
Filiform,
some
which
mucilage
epidermal features';
impossible
epidermal layer (photo 35);
Horsfieldia specimens
clearly
even
species.
per
abaxially of the sclerenchyma
and
(1950)
dried
'branching'
sacs,
'...tanniferous tubules... adaxial
species.
in
and
sometimes
34),
On the whole it is
were
structures occur
fairly thick-walled cells, adaxially
bundles
the
cases
Annonaceae he noted
constant
like this, because of the
study
a
all
some
iryaghedhi)
related species. The empty idioblasts in
The presence of tanniniferous sacs, also mentioned in the
contents
in
Horsfieldia
also recorded tannin-like substances instead of oil in the
Myristicaceae,
their distribution is
unambiguously
Gymnacranthera,
absent in this genus.
probably
are
have been described under 'Some miscellaneous
epidermis
probably
No.
27.
frequencies (photo 34);
(1895),
are
(1969)
some
cells instead of oil cells
VOL.
-
as
well
(1966).
Horsfieldia
superba.
of Rao and Chin
species,
This
is
(1966)
in
and
Armstrong (1975).
Infrequent,
they
thick
filiform, branched sclereids
abundant in Knema
are
bundle sheaths of the veins
mostly
to
GENERIC
Explanatory
The
generic
expedient
in
individual
the
diagnostic
species
in
some
from the
Knema
species;
slerenchymatous
epidermis (photo 36).
DESCRIPTIONS
value only
or
descriptions
specimens
are
infrageneric variation
descriptions.
adopted procedure
present
notes
leaf anatomical
several characters the
such
the
are
extending
percoriacea,
the
For
some
generic
very
detailed. If
mentioned in the
presented
given
relevant and
descriptions.
For
in tables and referred to
variable characters
range is
makes individual
is
are
of limited
without further
or
specification.
specific descriptions superfluous.
as
without
The
J. Koster &
P. Baas:
GYMNACRANTHERA WARB.
Asiatic
Leaf anatomy of
Photos
figs. 2,7;
—
129
Myristicaceae
22,27,30,31,37,38;
figs. 2,7;
table I
6
5
species;
studied.
species
surface
In
frequent
on
the stalk
view: Hairs
present
the midrib,
cells
(up
length, spread
out
decussate
arrangement (fig.
sometimes up
oriented
horizontally;
epidermal
cells and
reticulum of
cells
Unspecialized epidermal
focus and
high
thin
conspicuous
Abaxial
straight
but
walls,
cuticle
on
but
rows
sinuous
in
abaxial
to
the
tending
on
near
be
in
rarely present (in
G.
6591).
Cork
origin.
Small cells
surrounded
In
1).
pattern
of abaxial
bancana
23
—
x
on
12—16 /<m,
absent
length
with
a
thin
sometimes
cuticle in
in
present
oriented
the
the
epidermal
section:
to
rectangular.
to
layer,
G. bancana
of about the
cells
overlying
same
size
midrib
usually
small and erect, sometimes
adaxial
epidermal
thin
cells
areas
of cuticle
epidermal
cells flat
lumen side.
as
to
cells;
protruding
above
subsidiary
stomatal
the
in
ledges
centre
of the
cell
and/or
adaxial
absent
x
10—19
2.3,
water
average
stomata
coll.
s
of traumatic
surface, probably
abaxial
130
excepted.
/un
410
—
thick
epidermis,
on
the
outer
Outer
abaxially
on
periclinal
periclinal
sometimes
cells.
thick (table
often
cells
slightly
cells, sometimes lower;
lumen side,
Subsidiary
surrounding epidermal
nm
(table I), usually
Unspecialized epidermal
wall
complex
complex
slightly
cells
abaxial surface
wall
corresponding
slightly sunken; guard
absent.
—
or
26
—
to
adaxial
cells.
convex,
cells
1.0
or
probably
the adaxial epidermal
in surface view;
slightly
G. forbesii var. forbesii, King
dome-shaped.
mostly slightly
Stomata and
subsidiary
as
15
pairs
Giant
sometimes erect,
with indentations
seen
of cuticular
present
width ratio
infrequent,
or
arranged
Stomata confined
adaxial surface often erect,
on
major
parallel
Pitting
scarcely
of
without
midrib and
axis
and
areas
abaxially
as
long
Guard cell
at
anticlinal
adaxial cells also
Lamina dorsiventral,
rectangular, usually flattened,
dome-shaped,
epidermal
cuticular
the
inconspicuous.
or
contracta, Zehnder 9437 and
by radially
granular
square
focus
abaxial surface. Minor veins
paracytic.
a
(photo 22).
high
at
abundant in areolae,
scanty and aberrant
T-pieces
transverse
a
undulations
overlying
with
borneensis
var.
as
margin.
lower focus and with
at
well
as
Adaxial cuticle 5—18 /<m thick, abaxial cuticle 2—11
with
veins and leaf
epidermis only.
variously present, usually
warts
cells,
rather than
conspicuous
stomata
18
adaxially
square
1,
radially
of the hair.
shedding
slightly curved
to
veins; corresponding
polygonal
G.
1.2— 1.9. Polar
secretory
straight
in
to
mostly
less
more or
anticlinal walls
Abaxial cells
to
veins and leaf margin,
/<m, average values
values
of the
most
cell
midrib;
midrib, major
G. bancana
rectangular
course
to
epidermal
epidermis,
epidermis
with
anticlinal walls.
rows,
flanges usually present,
visible in
or
cells
midrib, major veins and leaf margin with prominent thin
arranged
perpendicular
in
of cuticle,
areas
sinuous
same
with sinuous anticlinal walls
curved anticlinal walls
faintly
anticlinal walls.
along
conspicuously
veins
cells with
thin
inconspicuous
adaxially
of the
each other
with alveolar material
midrib, major
on
least
2 arms,
mostly
not
to
Basal
after
ring
of cuticle between anticlinal wall
areas
epidermal
cutinized
a
polygonal,
usually
parallel
oval, surrounded by
or
is absent
slightly
to
abaxial surface,
to
walls.
granular, abaxially
to
plates, which
erect
thin
with
subtending
smooth
adaxially nearly
Cuticle
circle
a
cell
same
of the whole hair
Arms
2).
6, arranged in
to
arms
confined
of flattened cells with
Arms of the
4) excepted.
to
(table I),
uniseriate, composed
of
with
of abaxial
indented
on
the
cells embedded in
pointed,
Hypodermis
as
a
sometimes
continuous
BLUMEA
130
Table
Sizes
in
++
µm;
of
because
abundant;
=
+
or
=
present;
27,
No.
characters
I. Some leaf anatomical
preparation
poor
VOL.
±
=
other reasons; ad.
±
scarce;
=
of
1981
1,
Gymnacranthera species
=
very
adaxial; ab.
=
?
scarce;
=
character
not
observed
abaxial.
t/1
"a>
1
o
o
CO
c
(J
"3
W
1
(/3
1
E
W
£
C3
c
>>
U
>N
o
1
o
c
—
o
1)
<D
1-
c*_
o
x>
CT3
E
O
o
T3
tA
C
■s
■J
u
t/i
55
S
c
C/3
cL'B
a
E
GA
1
3
g
•3
"3
3
'C
E
IE
Species
and
f—
studied
specimens
ad.
G.
bancana
var.
bancana
G.
bancana
var.
borneensis
al
—
"o
Z
ad.
8
30
18
10
*3
c
C/5 .3
C/5
SI
ab.
14
260
410
£
3
—
ab.
_
+
50
+
+
+
+
+
±
G. contracta
Jacobs
SAN
9437
Kostermans
Haviland
13696
9420
134
Kostermans
7241
11
7
5
±
+
220
5
2
15
±
+
260
7
6
5
±
+
300
8
8
15
-
15
-
-
14970
350
11
11
15
Jacobs 5326
240
5
5
10
210
6
2
15
Rossum
63
-
+
-
-
?
-
-
+
-
+
±
+
-
+
±
±
+
±
-
eugeniifolia var. eugeniifolia
250
8
5
5
±
+
-H-
260
8
6
5
±
+
++
Rahmat si Toroes 3944
200
8
4
10
±
++
++
coll. 6622
240
9
5
5
±
++
±
Rahmat si Boeea 8035
eugeniifolia var. griffithii
King's
forbesii
Jacobs
var.
crassinervis
5259
Rosli 3393
G.
270
8
FRI 3908
G.
15
10
van
G.
15
8
260
Galau
G.
6
310
17514
Anderson
5
10
200
5363
Zehnder
forbesii
var.
5
5
20
5
2
15
coll. 6591
210
5
2
20
220
6
2
20
G.
paniculata var. paniculata
130
5
2
15
G.
paniculata
210
7
5
15
layer
var.
absent.
cells, which
loose
zippeliana
Mesophyll composed of
are
spongy
conspicuously
Rosli 3393,
square
so
rarely
with
to
3
times,
tissue. Abaxial
in
G.
some
either with sclerified
supplied
2
—
3 adaxial
sometimes up
layers
to
7
+
++
++
±
7
of spongy
cells in the
centre
in
layers
an
abaxial
flat
to
shallowly
arc-shaped
and with several
+
-
-
+
-
+
as
long
as
wide, and fairly
G. forbesii
var.
to
the
phloem
var.
or
bancana
abaxially prominently
bundle with the
to numerous
margin
epidermis (table I),
in G. bancana
concave,
most
crassinervis,
of the spongy tissue sclerified. Leaf
parenchyma cells, often adjacent
with
7
+
-
tissue sometimes sclerified,
collenchyma. Midrib fine to intermediate,
raised, supplied
-
layers of palisade parenchyma
4 times
eugeniifolia,, locally present
robust, adaxially approximately
groups in 2
+
-
forbesii
FRI 5749
King's
180
190
phloem
in separate
bundles situated
adaxially
J. Koster
of the
collateral bundle,
of
Groups
with
the
parenchymatous
which
(table
phloem
hypodermis
and
on
the
with,
bundles. Ground tissue from
phloem
of the
scarcely present
collenchyma
the
or
elements
xylem
some
131
(fig.
7).
of the collateral bundle, associated
sclerified
the adaxial side. Ground tissue also
and
as
in
epidermis
locally
a
even
periphery
between the
the adaxial
near
cells
to
parenchyma cells,
vascular system
absent
to
and often
centre
with sclerified
collenchymatous, interspersed
groups,
often
I);
by
abaxially
abundantly present abaxially
are
central
to
accompanied
Myristicaceae
groups, and in the centre, associated
the adaxial
of,
rarely
Asiatic
Leaf anatomy of
fibres present
sclerenchyma
peripheral phloem
centre
P. Baas:
&
developed
with
interspersed
brachy-
to
astrosclereids; adaxial chlorenchyma continuous in the midrib (not in G. bancana),
situated abaxially of the
flat
adaxially
raised. Veins
abaxially
with
small
adaxially
at
in G.
(veins
eugeniifolia
with collateral bundles, embedded in
and
abaxially
caps
with very
paniculata
with 3
bundles
usually
little
less
more or
adaxially.
brachy-
astrosclereids.
to
idioblasts,
of several
Crystals
which
then
less
deeply
parenchymatous
tissue
var.
and/or
in
of midrib.
usually
be
in cells
most
Minute,
grouped
sometimes
50
species;
In
8
usually
species studied,
view;
in older leaves
some
hairs
usually
excepted.
on
2
in
the basal end
at
the
mostly
Arms extended
abaxial
walls between cells with
one
hairs
upwards
arms
very
robust hairs with thick walls also
thin
a
to
especially
present
more
Large,
usually
a
thin
Sclereids
the
ground
abaxially;
in
G.
less
or
tissue
spongy
cuticle, probably
usually
tissue
filiform,
ground
in
in
sometimes in
cells.
scarce,
as
spindle-shaped
sometimes
tissue,
and the
— Photos
Hairs present
the
less
or
in
var.
present
of the
as
midrib
rarely branched
tissue of the
midrib,
38).
of cells with
midrib
eugeniifolia
prismatic crystals
epidermis.
veins,
G.
flattened and have
idioblasts with
mesophyll
druses
large
as
(in
more
epidermal
on
16, 21, 32; fig. 6; table II
examined
adaxial and abaxial surface of
mainly shed, especially
on
cells
palisade parenchyma,
less
or
species superficially
persistent
uniseriate, composed
hair,
in spongy
Empty
major
in the
adaxially (photo
surface
leaves,
Minute,
abaxial
HORSFIELDIA WIILD.
C.
a
xylem,
visible in cuticular macerations
astrosclereids in
to
of the
frequently present
numerous
more
in
in
present
brachy-
sometimes
37),
cells.
sheaths, rarely
tissue of midrib.
sometimes clustered
sclereids
by
also with free
bundles,
frequent
extremely
invariably present, frequent
ground
secretory,
(photo
bundle
bancana,
oil cells
spherical
fibres
epidermal
to
may
staining
crystals usually present, grouped
ground
in the
interruptions
collateral
cuticle and thin and short cuticular flanges, also
bancana
surrounded
sclerenchyma),
sclerenchyma
mostly adjacent
griffithii only abaxially)
the
fibres
sclerenchyma
bundle sheath. Petiole with vascular
arc-shaped
of
Groups
veins
raised),
mesophyll, supplied
few
a
Major
never
the abaxial side of the bundles. Ground tissue abundantly interspersed
to
mesophyll
areas
eugeniifolia
var.
the distal end similar to that of midrib, but with
at
confined
with
the sclerified parenchyma cells.
or
G.
(in
differentiatedparenchymatous
the basal end
phloem
raised
sclerenchyma
mostly poorly
system
collenchyma
slightly
to
in
surface,
arm
longer
(cells
oblique,
in
glabra
and
particular
with
arms
directions,
arms
on
H.
as
the
adaxially
but
the midrib. Hairs
1
—
3
—
6 per
stalk
cells
in stellate hairs. Cell
with thin walls but in H.
Basal cells
young
sabulosa,
2—11, mostly 3
than elsewhere),
in different
present.
H.
1
—
sylvestris
13, mostly 4
or
8,
BLUMEA
132
VOL.
27,
No.
1,
1981
epidermis
curved;
adjacent cells,
to
the
ground
midrib of tissue
slightly
in
to
astrosclereids
straight
+
+
+
+
+
++ ++
+
++
±
+
++
+
++
+
Brachy-
crystals Spindle-shaped
to
+
Swollen
±
±
++
+
+
+
+
+
+
flattened
extremely idioblasts,
druse to
adjacent cells, epidermal
Some
=
st
midrib
of
pith
in
Xylem
+
+
+
+
+
±
s
+
+
focus;
species
<1
areas
cuticle of
Thin
±
±
+
s
st
s
st
8
high
at
ab.
st
st
s(h)
ad.
s(h)
c.st
s(h) s(h)
ab.
2
2
ad.
8
8
s
s
st
s(h) s(h)
st
St
St
s(h) s(h)
4
2
2
4
2
=
Horsfielda
Anticlinal
sinuous;
of
.
*
*
?
2
2
?
3
2,4 4,6
?
1.5 1.5
=
s
char cters
walls
(h)
1.5;
only)
hair
numbers
per
cells
common
basal
(most
16-43* 18-31*
of Number
and
4,6,8
1
antomical
leaf
betwe n
areolae
of
the in
parts
2
mm
hairs/0.1
basal of Number
II.
ad.
1.5 1.5 1.5
1
3
2.5
<1 <1
6
9
1
1
8
9
2
3
1
?
1
<
<
1.5
1
>
1
>
1
=
1
>
Some
<1* <1*
10 10
ab.
ab.
3
1
5
3
3
3
10
3
6
4
3
5
13
1;
cuticle of Thickness
ad.
and
4
0
Table
betwe n
=
table very
at
also
see
200 490
250 280
59
230
220 210 310
110 170 250 460
focus.
high
I.
lamina of Thickness
sinuous
legend only
=
For c.st
studied
specimens
and
Species
5101 5274
12325
en
Ste nis
cras ifola Theunisen SAR glabra Junghu nIbid.
irya van van
H.
H.
H.
Roy
4031
2796
iryaghedhi macro ma Mer il parviflora sabulosa
BW
H. H.
H. H.
J. Koster
_
+
+
+
+
+
_
+
+
+
_
_
_
_
_
+
± ±
± ± +
+
+
+
+
+
+
7
+
+
—
—
—
—
±
+
7
& P. Baas:
+
+
— _ _— — —_ _ _ — warts.
s
s
s
s
s
s
s
s
s
cork
C.St
->
St
2
C.St s(h) C.St
1
2
1
St
2
s(h) s(h) C.St
2
1,2
2
into
parts
basal
4
4,8
4
4
4
4
5,6,7 4,8
4
many
of
5
<1
3
1
15
1
3
10
3
<1 <1 <1
4
2
4
2
5
4
5
2
5
3
5
10
5
1.5 1.5 1.5
5
4
5
4
7
transfomatin
of
6
6
7
because
240 160 240 200 270 180 280 270 310
species,
other
3
1123
4510
en
sylvestri
H.
Soepadmo
235
407
4446
&
1
Roy 3081
BW van bb
with
379
Gjel rup Robinson Kostermans Kuswat Pleyte
1839
BW
compar ble
range.
Hardly
*
* Full
Leaf
anatomy
of Asiatic
Myristicaceae
133
BLUMEA
134
1
abaxially
most
—
1
15, mostly
2
or
VOL. 27.
in
arranged
a
circle
or
subtending
cells and
cutinized
a
probably
surrounded
oval,
in H.
(table II),
of the groups of basal cells
by
after
ring
No.
1981
1.
glabra adaxially 16-43, abaxially
into cork
changed
less
more or
of the hair.
shedding
Basal cells
warts.
oriented epidermal
radially
Usually
basal
most
parts of hairs per surface unit (table II) on the midrib. In H. crassifolia abaxially 2 or
more
but
basal
in
H.
often
parts
joined together.
smooth
glabra
then absent
iryaghedhi only,
and
at
sinuous,
least
at
cells
high
some
thickened. Abaxial
midrib and
overlying
axis
long
parallel
cells also
major
in
arranged
rows
but
the
to
epidermal
cell
epidermis,
abundant in
and
striations
of abaxial
pattern
leaf
sometimes
(photo 21),
the
to
faintly
—
1.2
35 x8—19
—
pairs
/OTI,
average
average values
3.3,
Polar
inconspicuous.
1.6
wide:
relatively
are
Subsidiary
in H.
values 25
2.8;
in H.
15—19 nm.
frequent
probably
only,
Smalt cells with
probably
glabra
In
and H.
grooves
—
110
—
section:
490 pm thick
10 /im thick
Cork
(table II),
cells
of cuticle
iryaghedhi;
complex
or
not;
overlying
in H.
circular,
from
stomatal
of the cell surface
adaxial
abaxial
and/or
in
frequent
most
to
midrib often
wall
cells.
flattened,
H.
than
of
sunken
(in
cells
in
absent
H.
surface
on
iryaghedhi
partly
view.
midrib,
and
slightly
embedded in
the
adaxial
epidermal
Abaxial
major
epidermis
and
on
abaxial side
more or
crassifolia
subsidiary cells,
but
with
leaf
to
epidermal cells;
erect,
cells
cells
slightly
side, corresponding
veins
in H.
sabulosa,
epidermal
sometimes
dome-shaped and/or
complex
H.
sylvestris with
in H.
Unspecialized
abaxially
lower
in
or,
crassifolia and usually
epidermal
higher
periclinal
seen
as
papillae
guard
variously
extent
cuticle 2—13 /im thick, abaxial
sometimes with shallow indentations on the lumen
areas
the
dorsiventral
Lamina
in H.
of
usually
dome-shaped,
small. Outer
usually
in
warts
and
centre
cell
guard
sometimes
distributed and
hairs
pairs
width ratio
to
less
to a
epidermal cells,
(table II). Adaxial
anticlinal walls
rectangular,
conspicuously
epidermal
of
parts
present
oriented
relatively
sabulosa.
above
to
square
sometimes
sylvestris
cells
ledges present,
regularly
basal
cuticular
H.
and H. sabulosa the
thin cuticle, sometimes in the
by radially
transverse
1
surface,
modified
a
secretory,
surrounded
isobilateral,
cuticle
abaxial
from
developed
complexes.
epidermis,
the
on
and
length
inconspicuous.
in
abaxial
midrib, major
on
prominent
of traumatic origin, but in H.
glabra and
present, probably
sabulosa
visible
to
then elsewhere. Guard cell
10—17 /im,
parviflora
or
adaxial
rectangular.
slightly
subsidiary
on
Outer stomatal
absent
T-pieces
32x
absent
with
then
Abaxial cells
to square with the
crassifolia
H.
granular
—
or
some
thin, but
iryaghedhi.
Minor veins
cuticle
glabra;
very
Stomata confined
cells
stomatal pore in
so
—
only.
areas
sinuous,
rather then
polygonal
curved
thin
veins; corresponding
inconspicuous.
thin and, when striations are absent, less
20
be
epidermis
paracytic.
perpendicular
to
H.
crassifolia
slightly
to
when walls
in H.
of the
in H.
inconspicuous
specimens
scanty and aberrant
areolae,
margin,
straight
rectangular
rows,
course
tending
few
papillae
in
arranged
of cuticular flanges present, but
Pitting
veins
veins
a
in
margin, irregularly
sylvestris faintly abaxially.
(table II);
in
cells with
epidermal
perpendicular
or
leaf
sometimes with
straight. Cuticular flanges
are
H.
granular,
obviously present
and
anticlinal walls
(table II),
of cuticle between anticlinal wall undulations
division walls
conspicuously
to
16). Cuticular striations present
polygonal;
focus
veins
and sometimes in
faintly adaxially,
Unspecialized epidermal
or
faintly
midrib, major
on
structured and distributed (photo
abaxially
Cuticle
Alveolar material
abaxially.
papillae
margin.
and H.
hardly
less
flat,
with thin
in
H.
Stomatal
sylvestris p.p.)
so
in H.
glabra
J. Koster & P.
H.
and
as
and
outer
macrocoma;
Hypodermis
continuous
a
Baas:
Leaf
inner
layer
translucent cells,
sometimes with
epidermal cells,
2
—
of
Theunissen
crassifolia,
of square
layer
flattened
to
sometimes broader than
division walls.
periclinal
often present.
probably
in H.
adaxial
one
higher,
to
ledges
(absent
135
Myristicaceae
adjacent
Mesophyll composed
of
3, in H. crassifolia, H. irya, Van Royen 5101 and H. sabulosa 3—4 adaxial layers
cells
palisade parenchyma
which
present),
loose
fairly
raised
to
square
ridge
cells,
present)
in
or,
in H.
abaxial
shaped
glabra,
bundle,
H.
and
parviflora
a
less
more or
concave,
adaxial
straight
in 2
usually only abaxially
often
pith,
or
(sometimes only
sabulosa,
H.
margin
epidermis,
adaxial bundle;
with the
rarely joined together
bundles in the
phloem
to numerous
raised
robust, adaxially
with
the
to
also
are
wide, and
as
tissue. Leaf
compact spongy
adjacent
long
as
bundle,
with 2 small adaxial bundles in H. parviflora, and
or
in small separate
groups,
phloem
palisade layers
5 times
to
up
usually
Midrib fine to
abaxially prominently raised, supplied
interrupted
less
more or
abaxial
some
times, sometimes
4
parenchyma
collenchyma.
central
H. sabulosa
(in
sylvestris)
H.
sclerified
with
supplied
are
(in
or
either with
a
lower
Asiatic
stomatal
often present
59, H. glabra and H. sabulosa), consisting of
rectangular
of
anatomy
an arc-
with
the
and with several
layers,
accompanied by xylem
groups
(table
II; fig. 6). Groups of sclerenchyma fibres present, surrounding
the vascular system,
associated with
fibres present in the
pith,
tissue
from
with the
tissue
groups, and
peripheral phloem
associated with and often
centre
to
collenchyma
in the
even
periphery parenchymatous
as a
are
usually present
in the
pith,
Veins
with
collateral
differentiatedparenchymatous
poorly
the
distal end
similar
that of
to
collateral bundles, also with free
fibres
the
at
basal end
Ground tissue
druses in
often
and
to
macerations
as areas
often present in the
crystals frequent
spongy
the
tissue,
the
to scarce,
ground tissue
sometimes
most
abundantly,
peripheral ground
the
H.
a
frequent
as
and H.
major veins, abaxially
sclereids
to
astrosclereids.
close
to
and the
glabra
sometimes
abundantly present
sylvestris,
(in
as
H.
to
with
supplied
by
a
usually
with
3
arc-shaped
sometimes with small
Groups
of
sclerenchyma
main bundles.
Crystals frequent
the adaxial and abaxial
also
II),
few, mainly
a
mesophyll,
parenchymatous
ground
visible in
less
more or
more
or
most
a
thin
cuticular
small druses
spindle-shaped
frequently
in the
bundle sheath, and in
less
tissue of the
probably secretory,
spherical
midrib,
oil cells
often in the
sometimes present in
and H. sabulosa. Sclereids often present,
clustered
brachy-
to
astrosclereids
(table II), usually abaxially, rarely
bb
large
as
epidermis,
flattened and have
extremely
flanges (table
in the
end
abaxial side of the
in cells in the
tissue of the midrib
parviflora
basal
the main bundles.
thin cuticle,
in H.
mesophyll,
adaxially,
(table II). Large,
mesophyll
brachy-
with
slightly raised, abaxially
to
deeply staining cells;
grouped
of the midrib
in the
the
at
bundles
which may be
especially
ground
parenchyma
and surrounded
tissue of the midrib. Minute,
idioblasts with
epidermis,
pith (in
to
cuticular
of several less
ground
sometimes
usually abundant
pith. Empty
short
sclerified
bundle sheath. Petiole with vascular system
brachy-
epidermal cells,
thin and
sometimes
the adaxial side;
interspersed
in
fibres,
confined to the
with
interspersed
on
some
depressed
few
midrib,
mesophyll idioblasts, mostly
adjacent
cuticle
mostly
a
phloem
collateral bundles close and adaxial
with
embedded
bundles,
caps, sometimes only with
sclerenchyma
bundles. Ground
phloem
collenchymatous,
to
and often
astrosclereids. Major veins adaxially slightly
raised.
of the
locally developed hypodermis
sometimes, mostly abaxially, interspersed
cells which
at
sclerenchyma
centre
30811),
crassifolia,
sometimes in the
H.
astrosclereids in the
glabra
and
superficially
H.
in
also in the
ground
tissue of
sabulosa),
examined
and
H. fulva
BLUMEA
VOL.
27. No.
and H.
superba,
solitary
in
or
at
least in the
groups
of the
periphery
LOUR.
KNEMA
Swollen cells with thin walls often present,
mesophyll.
if present,
hypodermis,
rarely adjacent
or
sometimes in elevations of the leaf surface
epidermis (table II),
the
in the
1981
1,
136
to
the abaxial
in
(photo 32), and/or
tissue of the midrib.
ground
1—8, 11,12,18, 23, 24, 28, 29, 33—36; figs. 3—5, 8, 10;
— Photos
table III
83
31
species;
studied
species
view:
surface
In
leaves often shed
midrib and
on
with
the
arms
1—3
to
many;
with
upwards
arms
short;
1—8,
very
fig. 3,
1
mostly
surrounded
by
ring
(table III)
or
midrib, major
is
plumulosa
mainly
adaxially;
as
with
rectangular
to
cells
or
not
in
epidermal
horizontally
papillae
Guard cell
to
usually
on
a
17
cell
pairs
are
inconspicuous.
traumatic
to
1
adaxially
in
circle
a
14,
—
or
oval,
subtending
a
granular, abaxially
plates,
which is absent
long
axis
and
parallel
or
arranged
rarely
rufa (photo
in K.
at
sometimes
18).
at
lower focus and
often
in
arranged
perpendicular
in rows but
Abaxial
straight.
walls, and
veins
major
epidermal
to
tending
the
to
cell pattern of abaxial epidermis,
epidermis.
with
rows,
course
be
of the
polygonal
Stomatal
nipple-shaped papillae
x
8
—
complex
of the 4
10—16
—
6
26
—
30
/rm;
Polar
variously
conferta and
absent
—
T-pieces
more
or
less
complex (photo 23).
27x9— 14
some
sometimes
usually
K. kunstleri
abaxial
midrib, major
by
guard
the
cell
present,
infrequent,
length
im,
oblongifolia.FRI0111,
in K. lunduensis
present,
on
to
bordering epidermal cells;
above the stomatal
16 /im, average values 20
pm.
warts
but in K.
or
overarched
visible
slightly
Stomata confined
in areolae, scanty and aberrant
30
cells
epidermal
of cuticle between anticlinal wall
areas
midrib
Cuticular
complexes.
with sinuous anticlinal walls
slightly curved anticlinal
star-shaped opening
Cork
origin,
relatively
Pitting of cuticular flanges present, but inconspicuous.
in
relatively long:
wide:
to
curved anticlinal walls
cell pattern of adaxial
—
sculpturings
convex
width ratio 1.3 —2.8, average values 1.7 —2.4; in K.
relatively
lobed
stomatal
midrib less sinuous,
adaxial cells also
directed
leave
bordering
slightly
margin, paracytic.
pairs
are
Arms
(table III, photos
cells and
almost smooth
polygonal, adaxially
overlying
rectangular.
veins and leaf
epidermal
with anticlinal walls of
inconspicuous
very
square with the
epidermis, abundant
the
straight
prominent
Minor veins
cells
to
corresponding
cells
veins; corresponding
rather than
hairs
arms.
may be
arms
structure
Basal cells
below).
reticulum of erect,
anticlinal walls
Abaxial
papillae.
in
(table III), arranged
adaxially
Cuticle
cells with straight
epidermal
have
not
(cells
elsewhere),
when
excepted;
where the
varying
oriented
adaxial cuticle with
focus and
undulations;
cells
than
longer
persistent
one arm
of the hair. Most basal parts of hairs per surface unit
restricted
sometimes
thin,
radially
a
Unspecialized epidermal
(very) high
less
hairs
of cells with
in stellate hairs. Cell walls between cells
midrib,
1 —4
surface, in older
some
veins and leaf margin; the alveolar material in K. intermediaand K.
striations and grooves
present
mostly
shedding
with alveolar material
on
1 —8,
the midrib.
on
as
the
classification of types
more
after
usually
distal cells do
most
Hairs
thick walls.
to
also
on
stalk
4)
to
the
in different directions,
oblique, especially
see
but
the midrib the hairs
on
hookeriana)
11, abaxially
—
cutinized
sometimes
in K.
with thin
arms
adaxial and abaxial
on
uniseriate, composed
midrib sometimes up
(on
extremely long (as
extended
veins. Hairs
major
one
Hairs present
(table III), especially adaxially,
to
guard
pairs
are
usually
probably
of
abundant, especially abaxially,
J. Koster &
and
and/or
abaxial
to
usually
Outer
of
leaves, especially
most
cuticular
lower
erect,
K.
absent
linifolia papillae
often with
midrib, major
on
papillae. Subsidiary
translucent cells,
continuous
major veins,
composed
high
as
layer
but
as
parenchyma
2
—
cells,
3,
are
up
to
square
(or
than wide (in K. curtisii and K.
K.
pallens
and K.
B RUN 5587, K.
tissue;
cells of
abaxial
layers
sclerified
palisade
margin
and
K.
laurina,
spongy
spongy
(table III);
supplied
with
a
straight
ashtonii and K. kunstleri, and
with the adaxial
bundle;
especially abaxially, and
accompanied by xylem
straight
present
to
arc-shaped
in
the
surrounding
an
3
lignified
cells
at
the
periphery,
side;
ground
or
often abundant
the sclerified cells
tissue
also
often
as
a
or
the main
of
2
in the
large,
var.
peripheral phloem
to
often
less
even
parenchyma
brachy-
or
are
present
groups, and
in the
centre
periphery parenchymatous
and also present
with
pith,
more or
fibres
to
cells, especially
in the
locally developed hypodermis
interspersed
the
(very)
to
abaxial bundle,
sclerenchyma
with sclerified
(table III)
Leaf
to
in K. ashtonii
associated with and often
bundles. Ground tissue from centre
collenchyma
as
Groups
of the
usually interspersed
large pits.
adjacent
rarely joined together
phloem bundles
sometimes 1
fibres present in the
collenchymatous,
with
the
only
many cells of
separate groups, usually in 2 layers,
associated with
pith,
pallens
(in
curtisii,
less compact
sometimes
interrupted
abaxial bundle,
sclerenchyma
phloem
more or
5 times
K.
abaxially prominently raised,
so,
oriented
5).
to
(in
to
Midrib of intermediate size
in
palisade
scarcely longer
to
sometimes up
supplied
never
Mesophyll
of
layers
sclerified,
adaxial bundle,
phloem
fig.
the vascular system,
adaxial
times,
walls
prominently
xylem bundles,
layer
rectangular,
the midrib and the
(table III), usually
one to numerous
III;
4
pseudolaurina)
groups (table III; fig. 4);
pith (table
near
slightly flattened)
or to
arc-shaped
with the
with
only
cells
present,
adaxial
one
wide, and fairly loose
collenchyma.
arc-shaped
directed
material; guard
of
in
and
Stomatal
margin.
horizontally
in K. mandarahan and K.
sometimes
to
as
parenchyma
with
robust, adaxially raised,
even
occasionally
tissue have thick
supplied
or
tissue
seen
as
conferta
palisade parenchyma.
3 —
or
and K.
oblongifolia
spongy
often with sclerified
epidermis,
long
as
in K.
flattened
of
sometimes
4,
linguiformis)
percoriacea)
of cuticle
leaf
less
consisting
layer)
with the
intergrading
sometimes
which
or
less shallow
adjacent epidermal cells, adaxially
locally developed,
and sometimes
of
lower than
or
areas
striking
small.
usually
stomatal ledges sometimes
outer
abaxial
inconspicuous
one
more
sometimes present,
Hypodermis
of
most
midrib
overlying
more or
thin cuticle, without alveolar
a
inner and/or
subsidiary cells;
inconspicuous.
a
cells with
cells with
veins and
cells with
complex sunken; bordering epidermal
cells
to
square
dome-shaped,
abaxial side
on
often
(table III),
adaxially
cells
epidermal
papillae,
thick
conspicuously
with thin
side, corresponding
epidermis
(sometimes
cells;
veins and
cells.
300 jum thick (table
—
9//m
—
to
partly
in adaxial
along midrib,
epidermal
of adaxial epidermal
complex
surface view. Abaxial
embedded in
slightly
adaxial side often dome-shaped,
wall
periclinal
abaxially
than adaxial epidermal
on
indentations on the lumen
as
oriented
by radially
and
on
layer. Unspecialized epidermal
sometimes erect,
rectangular,
and circular,
midrib,
thin cuticle, sometimes in the
a
cuticle 2—17 /im thick, abaxial cuticle 1
granular
higher
the
along
Small cells with
section: Lamina dorsiventral, 80
transverse
III). Adaxial
a
veins and
on
137
Myristicaceae
only, probably secretory, occasionally present
epidermis
often surrounded
margin,
In
Leaf anatomy of Asiatic
from stomatal complexes.
of the cell surface
centre
with
Baas:
regularly distributed, mostly
developed
leaf
P.
pith;
on
(mostly)
often the
the adaxial
brachy-
to
138
BLUMEA
midrib of tissue
-
VOL.
27,
No.
astrosclereids
to
1981
_
—
±
±
—
—
—
—
+
+
±
+
+
+
++
±
+
-
—
—
—
—
+
-
ground
+
in
1,
—
±
+
+
+
+
+
—
-
Brachy-
midrib of
tissue
cells
ground peripheral
parenchyma
+
in
+
1
midrib of
pith
in
margin
cells parenchyma
Xylem
species
Knema
bundles.
parenchyma
leaf
+
±
1
+
—
-
+
in
±
± ±
±
±
—
—
—
—
±
±
—
—
—
—
+
±
a
in
—
—
—
—
—
-
a
+
Sclerified
text) (see type
large
+
+
-
Sclerified
tissue spongy
cells
++
Sclerified
8a
Hair
2
7+8b 8a
8b
+
3
7
3
6
3+7+8c
7
++
—
±
—
—
±
±
1
2
2
2
2
2
2
4
2
2
4
3
3
3
5
1.5 1.5
3
8b 8b
7
a
+
7
=
1
of
layers;
char cters
abaxial
abaxially
with
only)
hair
covered
numbers
per
cells
leaves
common
basal
hairs
++
+
+
9
—
9
±
Adult
ab.
1
1
1
2
1
2
ad.
2
2,4
2
4
2
4
2
ab.
15
40
5
3
5
3
20 30
ad.
3
5
3
3
3
1.5
5
5
ab.
2
2
3
4
3
4
2
2
9
2
ad.
6
6
6
9
4
6
4
4
12
5
90
120
1
1
1
(most
of Number
=
2
2
8,9
a
antomical
II.
and
areolae
I
of
leaf
tables
Some
also
III.
see
Table
legend
the
parts
in
10 10
3
3
3
3
<1
8
3
2
6
3
12
9
7
13
4
mm2 hairs/0.1
basal
of Number
cuticle of Thickness
lamina of Thickness
170 140 140 230
120 110 230 160 140 180 160 220 120
For
studied
specimens
and
Species
ashtoni cin amo ea
var. var.
kunstleri
curtisi
var.
5587
A4139
ashtoni ashtoni atenuat conferta curtisi BRUN FRI123 6 elmeri SAN
K. K. K. K. K.
K.
ssp.
18294
S
fur ace globulari hokerian interm dia korthalsi kunstleri latifolia
K. K. K. K. K. K. K.
J. Roster &
++
+
+
+
+
+
-
+
—
—
+
+
-
+
-
±
-
++
P. Baas:
++
Lea)
++ ++
+
+
+
+
++
++
+
+
+
-
+
+
+
—
+
±
anatomy
±
-
++
1
-
+
-
—
-
-
-
+
-
-
—
-
-
-
++ ++
1
1
1
1
?
+
+
+
+
2
2
2
4
2
8,9
8
4,6
11
4
5
3
3
3
3
1
1
3
3
5
5
1.5 1.5 1.5
3
5
3
4
100 110 120 140
laurina Chew
2
-
2
2
2
4
1.5 1.5 10
-
a
+
7+8b
+
4
-
—
—
-
+
-
+
±
8c
8a 8a
•
++ ++
+
8b 8b
++
—
±
3 ?
-
-
2
2
1
2
2
2
2
4
6,7
8
2
2
4
4
5
20
10
3
3
1
3
3
1.5
15
1.5
1.5
2
3
3
2
3
5
3
3
2
2
6
3
7
8
5
4
14
7
8
2
2
8
100 180 210 150 300
170 210
3
1.5 1.5
5
80
140 120
SAN
—
++ ++
++
-
-
+
+
1
-
+(1)
+
—
-
—
-
+
+
7
-
+
-
—
++ ++
-
+
+
+
+
±
+
+
+
++
+
+
-
-
-
±
+
-
-
1
+
+
-
-
a
3
-
2
1,2
3
6
-
8a
5
-
8a 8a
7
—
-
2
1,2
2,3
2
1
4
4
5
4
4,5
2
2,4
3 3
3
15
3
1.5
1.5
15
2
2
1
3
4
3
4
4
1.5 10
5
<1
5
9
2
9
17
4
1.5
—
150 160
-
SAN
3482
S
K.
982 4
2533 73 12
pal ens Meijer SAN
percoiace plum losa pseudolarin psilantha
K.
K. K. K. K.
11
01
+
+
1
5
KEP FRI
S
S
K.
-
-
+
4
8b
-
+
2
2
2
2,3,4
4
2
4
35 20
3
3
1.5
3
10
5
3
3
4
3
2
3
3
7
9
3
8
12
6
1
2
1.5 1.5
1
180 250 150 150 200 150
150 180
pulchra Kostermans
++ ++
2
1
3
+
setosa
5103
14373 29064
+
8b 8b 8b 8c
+
forma
linguiformis linfolia lundensi mal yan mandran oblongat oblngifola
K. K. K. K. K. K.
+
percoiace
24985
Ja
-
—
-
230 300 260 160 130
ssp.
2502
+
—
a
++
±
1
—
oblongat
Wilde(-Duyfjes) 41273 FRI2431
de
1
—
5
±
+
5
143
var. Wee
-
4
+
++ ++
±
—
1.5 1.5
14 29
laurina Lek
8a
+
+
+
1
+
+
1
K.
-
-
139
of Asiatic Myristicaceae
9908 9907
29015
SFN
retusa Sinclair
K.
ssp.
Sinclair rubens rufa tenui ervia toment la
K. K. K. K.
140
BLUMEA
astrosclereids
then situated
adaxial
of the
No.
chlorenchyma
collenchyma.
bundle sheath,
sclerenchymatous
layers
the adaxial and
on
abaxial
1981
I,
continuous in
rarely
Major
veins
the
midrib,
adaxially slightly depressed
transcurrent
often
entirely
bundles
K.
with
fibres
sclerenchyma
often with
epidermis
adjacent
latter condition
sometimes
a
in
in the
interruptions
Minor veins
few-
many-celled
to
in K. laurina, K.
xylem,
also
bundles,
with
at
more
less
or
small
of
Groups
brachy-
to
in
(when large
collateral bundles close
fibres
sclerenchyma
astrosclereids.
in the
idioblasts)
the midrib, in the
and
to
the basal
at
the basal end with
at
free
epidermal
crystals frequent
to
parenchymatous
bundle
scarce,
present in epidermal
cells.
from
grouped
cells.
in cells in the
Flattened,
cells
epidermal
usually
tissue of the
ground
also in the
in
present
the
tanniniferous
idioblasts with
epidermis
sacs,
in the
rarely
also
sometimes
prismatic crystals
usually solitary. Irregular
sometimes clustered, present in
(fig. lOf). Large,
less
more or
mesophyll
in
small druses
spherical
in
and/or
oil
peripheral
in K. elmeri, K. kunstleri and K. retusa, Sinclair 9908,
midrib,
pith. Empty
grouped,
abundant,
sometimes
tissue of
less spindle-shaped
tissue of the midrib,
less
or
or
the
druses
large
to
ground
large; (very)
more
bundle sheath,
cells of K. hookeriana
present,
small
to
interspersed
mesophyll (most frequent
ground
more
with rounded edges,
crystals, usually
and
frequent
most
and in the
sheath),
as
the main
confined
mostly
often also present in the
Usually minute,
present in cells of the parenchymatous
abaxial
bundles
phloem
adaxially
end
Crystals usually frequent
mesophyll,
palisade parenchyma
often also
in
present
small
oblongata,
the distal end
abaxial side of the main bundles. Ground tissue usually abundantly
with
sheath
epidermis.
small,
especially
poorly
vertically
to
bundle
through
the
to
Petiole with vascular system
collateral
arc-shaped
by
sometimes with small, collateral bundles between the main bundles, and
adaxially,
bundles.
of
(the
with
the lateral sides, of several cell
on
surrounded
abaxial
and/or
pseudolaurina).
midrib,
less
or
rows
and K.
and
sides,
sometimes sclerified
composed
that of
to
more
adaxial
are
vertical
or
percoriacea
similar
the
to
which
extensions,
one-layered
bundle sheath, embeddedin mesophyll
differentiated parenchymatous
3
27.
raised, abaxially prominently raised. Veins with collateralbundles, supplied
to
a
(table III);
abaxially
VOL.
(photo
often present
a
29).
thin
cuticle, probably secretory,
More
and
adaxially
less
or
thick-walled
abaxially
of the
sometimes
cells, probably
sclerenchymatous
bundle sheaths of the veins, sometimes 'branching' towards the adaxial epidermis,
sometimes
even
protruding
in the
present, sometimes abundant,
12326)
or
as
tissue of the
pith,
in K.
Thick
sometimes clustered
oblongata
filiform,
veins
Hair
For
to
the
chlorenchyma
of
adaxial
of the
preparations
damaged during
studied
at
in
to
(in
K.
curtisii
astrosclereids in
some
ground
specimens
curtisii, FRI
var.
peripheral
from the
tissue of the
major
veins
morphology
were
percoriacea,
36).
of the hairs,
of individual hairs for
left for the
study
magnification
so
transverse
light microscopy,
have been used.
the microtechnical process,
low power
in the
(table III).
bundle sheaths of
sclerenchymatous
epidermis (photo
ground
of K. laurina also
Knema
study
views obtained with the S.E.M.
photographs
sclereids
brachy-
S. 34822 also in the
the, mostly
detailed
sections and
brachy
branched sclereids sometimes
abundant in K.
present,
types
a
Sclereids often
epidermal layer (photos 34, 35).
midrib, mostly abaxially,
extending through
the
as
Unfortunately
a
paradermal
well
as
the hairs
surface
are
often
that for certain leaves only S.E.M.
of the hairs. The entire
with
and
as
tomentum
dissecting microscope.
In
has also been
fully developed
J. K.OSTER &
leaves the hairs
cell
size,
Hairs
with
diversity
wall thickness
arms
short
very
other
using
such
arms,
total
etc.
of the leaf.
have
on
young leaves have been studied.
(very)
case
141
Myristicaceae
of the hairs of Knema have been classified
the midrib and the
on
Asiatic
Leaf anatomy of
type of stalk cells, relative size of the body of the cells with
as
remaining part
present
Baas:
often shed, in that
are
The morphological
characters
P.
veins
major
They
bodies)
and have
of the
leaf,
places
compact (which
more
short
relatively
have
different from those
usually
are
longer,
are
been
not
into
account
the
on
the cells
These hairs,
arms.
taken
means:
rarely
in
this
classification.
The
and
photos
indispensable part
also
a
If
key.
CLASSIFICATION
A.
One
or
Type 1.
more
stalk
Long
OF
2.
Long
one or
two;
some
1, then try type 2
AREOLAE AND
distinctly longer
cells
MINOR
considered
as
an
way the classification is
etc.
VEINS OF KNEMA
LEAVES.
than wide:
part
with
short and compact;
arms
arms
with
thick walls.
See
3.
photo 1, fig.
Type
HAIRS ON
cells
stalk
fit type
not
should be
3)
fig.
of hair types. In
descriptions
hair does
a
1—8,
drawings (photos
of the
stalk cells
up to
with
three; part
short and compact;
arms
arms
with thin walls. See
fig.
3.
Type
3.
Long
stalk cells
B.
Stalk cells
Type
Type
4.
5.
long;
walls.
See
Hairs
long;
thin
Type
6.
Hairs
with
Hairs
short to
thick
walls:
Type
Type
7.
8a.
Hairs
short;
are
thick
See
Hairs
b.
Hairs with
arms
Note:
in
the
case
MYRISTICA
C.
72
species;
In
(table
13
L.
2
to
obviously
—
surface
of about equal
distance
more
in
the
3.
proximal regions;
arms
with
thick
pointed tip,
a
and with
fairly
the
is
arms
considerable); arms thin,
3.
at the distal end
100
surface view
pm
relatively short,
and
less);
with thin to
with
arms
thin
to
3.
view 65
—
hairs with
the
250 pm. When
a
diameter of
hairs
more
with
than
a
diameter of 100 pm
also
100 pm
occur.
Arms
3.
photo
lengths
8,
3.
fig.
of hairs of type 8a and
c;
arms
with
fairly
thin to
3.
difficult; some species show
short
with
body,
between
or
long
a
gradual
transition from
Still this subdivision
may
help
one
to
subtype
identify
a
hairs.
10, 13—15,
17, 19, 20, 25, 26; figs. 9,
10; table IV
studied
view:
Hairs present,
on
sometimes
adaxial surface
almost
(table IV), uniseriate, composed
midrib and
the
the
near
give precise lengths.
Photos 9,
IV), in older leaves
(on
long (in particular
arms
photo 2, fig.
photo 4, fig.
walls. See
between
8 is
species
sometimes shed
or
with thick
type
of
in
(fairly) long,
photo 6, fig.
photo 7, fig.
of
broad
(diameter in
surface
it is useless
The subdivision
another. Therefore
specimen
See
in
See
(see type 7),
length
a
thick walls.
to
diameter
very
arms
and small
present
long;
longer than
long (the
photo 5, fig.
walls. See
c.
See
3.
walls.
of cells small;
short
very
and less
of cells
thin to thick
walls.
with
photo 3, fig.
long; body
long; body
Hairs
fairly short,
arms
thick
(very) long;
arms
3.
See
fairly
with
part
thin to thick walls.
at the distal end
arms
fig.
walls.
with
than wide:
longer
not
Hairs
two;
one or
proximal region),
major
veins often
length, spread
out
more)
to
always,
on
abaxial
and
at
excepted.
angles
Arms of
with
surface
abaxial surface
of flattened cells with 2
stalk cells
horizontally,
confined
arms
arms,
a
of
the 1
single
cell
adjoining
142
BLUMEA
VOL.
27,
No.
1,
1981
mesophyll
crystals; ocurenc
++
cells
epidermal
abaxial
in
Crystals
epidc
adaxial
in
Crystals
cells
=
of
•a s
+
++
+
(p)-c r-p-c r-p-c
p,s,n
r-p
P.s p,s
s,n s,n
s,n,h s,n,h
(s)
b-a
sc-b-a
r
spindle-shaped infrequent
+
+
crystals Spindle-shaped
c
s
druses; Types
ground
midrib of tissue
OJ
periph
in
midrib
of
cells
sc,b-a
sc-b
Sclerified
=
hypenatd.
crystals;
d
species
Myristca
astrocleids.
=
a
cluster d
hypodermis
forms
X<D
"a
i
o CJ
stomatal
=
c
char cters
brachyslerids;
crystals;
in
-
Xylem
•3
the
bordering
Intermdiate
X
epidermalof
ct
f,
—
"a. E oai stomatal
the
cells,
only.
-
-
Continuous
00
1
■,
of
pith
Number
1
SO
OS
+
+
bordering
on
++ ++
Papillae
abaxially
+
1
+
6-10 5-9 6-10 4-6
1
t-rt
1
1
6-10 6-9
+
+
+
1
1
6-9
6-10
+
+
antomical cells; prismatic hairs
wi
covered
leaves
6-8
+
hairs
++ ++ ++ ++
.c
+
+
Adult
=
b
=
lamina
of
c
p
leaf
Some
parenchyma edges;
rounded
IV.
Table
sclerifed with
=
sc
I—I I.
crystals
=
tables
r
only:
types
parts
mm2 hairs/0.1
ab.
5
15
40 30
15
basal of Number
ad.
in
5
1
1
+
+
ab.
6
6
6
ad.
9
12
8
material Alveolar
basal
1
1
4
+
2
15 15
50
5
3
25
1.5
15
+
+
+
+
+
3
3
3
3
5
3
5
6
7
8
8
1
cuticle of Thickness
=
h
crystals;
to
=
n
8 4
4
260 190 220 230 240
140 230 200 210
230 310
lamina of Thickness
studied
nedl-shaped
legends crystal spindleFor For
on
cells
also
see
the
brackets.
specimens
and
betwe n
Species
fatua
25986 25893
chrysophyla Brass cinamo ea
Brass
M.
M.
1506 24236
pa uan
s.n.
15023
var. var.
SAN SAN fatua fatua
787
Vriese
fragrans Sulit Beguin gi antea globosa
M. M. M.
de
M. M.
J. Koster
-
(P),d,(s)
—
-
+
-
-
-
-
d,(s)
-
—
—
-
—
-
+
+
+
s*
-
(P),d (s) d,(s)
—
—
-
-
-
-
+/±
-
-
+
+
-
+
-
+
++
-
-
-
+
-
-
-
-
s
(p),s
-
-
-
s,h
-
-
-
—
5
9
—
10
—
5
-
3
-
d
r,P
-
(P),d
s,n
-
b-a b-a b-a
sc**
sc
sc
+
+
-
-
-
sc
-
-
—
-
-
-
-
-
-
+
—
-
-
-
-
-
7
6-11
6-9
+
+
—
—
-
—
45 40 50 50 45 50 50 50 40 20
5
-
(P),d (s)
++ ++ ++ ++ ++ ++ ++ ++ ++
3
++
d,(s)
(s)
3
-
5
—
5
-
Leaf anatomy of
+
4-7 4-7 5-8 5-8 4-7 5-8 4-8 4-7 4-8
—
& P. Baas:
3
1.5
3
1.5
—
+
+
3
4
5
5
6
8
11
9
8
9
10
9
11 10
9
11 10
5
4-8 5-8
+
+
+
+
7-11 6-10
+
+
+
+
•
+
+
+
10 10 15 15
+
-
-
-
-
4-6 4-6
-
-
++ ++
30 35
1.5
5
5
10
+
+
+
+
+
+
5
4
4
6
3
5
7
7
7
11
7
9
8
270 190 220 190 260 290 310 260 310 180 200
1
7
170 200 320 220
Asiatic
5
280 350
origin.
traumatic
of
7
UNESCO
624
39 95
guaterifola Kostermans Sina g ul
M.
77
30298 20850 5365 32924
371
Mihil Elmer BRUN SAN Ambulah
55
**
908
Kostermans FRI50 6 lepidota maingayi maxima Thorenar Grashof sphaerosp ma subalu ta vilosa Kostermans
M. M. M.
probably
6438
E1P
4873
M. M. M.
A4747
SAN
minute;
very
*
Myristicaceae
143
144
cells. Arms with thin walls
In M. villosa
with
arranged
in
a
circle
less
very
frequent
smooth
material absent
on
cuticle between
in M.
wall
midrib and
or
major
veins
in
rows
cuticular flanges
but
epidermis,
abundant in
veins and leaf
25).
to
of
Guard
to
guard
cell
cell
areolae,
pairs
are
1.3
surface
origin.
be
—
—
Cork
average
and
on
or
Adaxial
overlying
long
absent
on
axis
cells also
of
Pitting
visible
slightly
Stomata confined
slightly
to
rectangular.
walls
in
abaxial
to
midrib, major
overarched
by
a
ring
of
papillae (table IV, photos 13, 15,
values 26
average
values 1.6
—
2.8;
27—39 gm. Polar
in
M.
—
34x11
—
18
m,
sphaerosperma
the
T-pieces sometimes present,
variously present, usually infrequent, probably
warts
Small cells with
especially
21
straight
of
areas
maingayi.
square with the
complex partly
without
or
thin
division
straight
Minor veins
M.
with
curved anticlinal
slightly
rather than
only.
in
adaxially
veins; corresponding adaxial
epidermis
Stomatal
shallow grooves
Very
and M.
to
inconspicuous.
ceils with
3.4,
of the
of variable
cells. Abaxial cells
epidermal
most
Cuticle almost
plates; alveolar
erect
cells with
by
usually
polygonal, adaxially
to
4,
after
ring
cell
one
(table IV)
present
with
rectangular
—
M. fatua,
if present,
inconspicuous,
scanty and aberrant
a
thin
cuticle,
sometimes in the
centre
sometimes present in the adaxial
only, probably secretory,
epidermis,
straight
gigantea
polygonal
but
relatively long:
usually inconspicuous.
of traumatic
to
21—39x8
pairs
rows,
course
margin, paracytic.
width ratio
in
abaxial
bordering epidermal
length
the
tending
cell pattern
4—11
in M.
papillae
usually present,
epidermal
cells
fragrans. Abaxial epidermal
arranged
perpendicular
cells
cells
undulations;
cells often smaller than abaxial
epidermal
margin.
sometimes very
usually with,
anticlinal
common
focus and
(very) high
at
leaf
epidermal
Unspecialized epidermal
curved anticlinal walls and with
arranged
veins and
of
the areolae.
reticulum of
a
in
cutinized
adaxially,
often with alveolar material
as
a
composed
in
frequent
most
midrib, major
lower focus and
if
surface,
or,
basal cells surrounded
subtending
Basal parts of hairs
sometimes
13—15),
sinuous anticlinal walls
parallel
abaxial
midrib, abaxially
gigantea and M. maingayi.
especially
on
1
mostly
row;
also
are
leaves. Basal cells 1
not on
a
walls and
oblique
these hairs
10);
mostly 2;
cells and
with anticlinal walls of
corresponding
at
adaxial surface
in M. villosa.
(photos
4 in
-
taller than elsewhere.
usually
(photo
bud scales etc.,
on
epidermal
granular, abaxially
to
appearance
walls
on
1981
of cells with
body
arms
sometimes with 3
oval,
basal parts
hair;
the
on
but
species,
I.
midrib
on
with
thick-walled
oriented
small, especially
27, No.
Hairs
(photo 9).
M. maxima
radially
of the
shedding
or
VOL.
extremely long,
parallel,
other
some
lepidotaand
more or
hairs
some
almost
short,
present in
M.
-
BLUMEA
along
the
midrib,
the
veins
and
of the cell
and/or
the
leaf
abaxial
margin,
surrounded by radially oriented epidermal cells.
In
transverse
IV). Adaxial cuticle 4
with
a
wall
cells
complex
the lumen
Abaxial
erect
to
overlying
midrib
higher
on
epidermal
side, corresponding
in M.
higher
in
M.
than
abaxial surface
with thin
gigantea
lower
to
cells with
—
350
thick
cells
thick
(table IV),
adaxially
on
same
with
as
as
the
seen
papillae
midrib, major
height
often
square
to
square
to
Outer
cells;
periclinal
less shallow indentations on
of cuticle
maingayi
to
(table
adaxial epidermal
often small.
more or
areas
and M.
Basal cells of hairs abaxially of about the
but
140
rarely dome-shaped, abaxially dome-shaped
thickened cuticle on the tips; papillae absent
cells,
dorsiventral,
layer. Unspecialized epidermal
flattened,
of adaxial
epidermis
Lamina
1 2 pm thick, abaxial cuticle 2—11
often erect,
rectangular,
epidermal
section:
cuticular
granular
rectangular,
—
in surface
with
a
view.
strongly
veins and leaf margin.
unspecialized epidermal
cinnamomea, M. guatteriifolia and M.
villosa
(photo
19).
P.
J. Koster &
of
stomatal
complex
guard
stomatal
cells
sometimes
ledges
continuous layer sometimes present
sphaerosperma
translucent
in
cells,
the
also
some
specimens
and
parenchyma.
4
—
occasionally
usually supplied
with
adjacent
the
to
fine, adaxially
adaxial bundle and
pith,
sometimes
with
of
the
developed
sometimes
IV);
to
with
supplied
through bundle
the
at
at
tissue
with
often
which
adaxially
veins
bundles
that of
to
adaxially,
tending
in M.
less
mostly
confined
to
druses
often
adjacent
(in
with
M. villosa
to
an
as
to
be
in the
situated
to
pith,
are
and
(table
abaxially
mesophyll,
differentiated
vertically
transcurrent
Petiole with vascular
to
fibres
sclerenchyma
astrosclereids.
at
to
the basal
usually
Crystals usually frequent
in idioblasts in the
small)
in
interruptions
collateral bundles, also
sometimes with small collateralbundles close
brachy-
they
locally
slightly raised,
poorly
sometimes with
arc-shaped
a
interspersed
embedded in
lepidota.
periphery
astrosclereids
midrib,
often
in the
even
to
collenchyma
depressed
with
associated
centre
to
of
fig. 4). Groups
the abaxial side of the main bundles. Ground tissue
abundantly interspersed
large
in
usually present
by
and adaxially from the main bundles. Groups of
end
as
separate
bundles in the
with and often
mostly
midrib,
more or
abaxially
in
phloem
tissue sometimes
slightly
surrounded
caps and
the basal end with 3
robust, rarely
to
system,
from
the
are
with collateral bundles,
extensions, especially
margin
parenchyma
present,
phloem
(IV,
vascular
associated
bundle sheath, sometimes
sheath
phloem
the
side; ground
(table IV),
ridge
with the
sometimes continuous in the
Major
Veins
central
groups table
pith,
sclerified
brachysclereids and/or brachy-
the distal end similar
xylem,
with free
cells
with
sclerenchyma
parenchymatous
system
the adaxial
chlorenchyma
raised.
abaxially
on
to
square
M. villosa. Leaf
some
and with 2 —9
Ground
collenchymatous,
parenchyma
collenchyma.
the
bundles.
are
less straight, sometimes interrupted
or
surrounding
in
groups, and
interspersed
adaxial
of the
more
xylem
near
palisade
sphaerosperma
which
of intermediate size
raised
a
in
so
abaxial bundle,
with
present,
hypodermis
with sclerified
arc-shaped
phloem
parenchymatous
a
M.
the
wide, and fairly loose spongy tissue; cells
sometimes with
only
a
M.
rectangular,
with
guatteriifolia,
sclerified, frequently
accompanied
peripheral phloem
centre
as
layers, especially abaxially,
fibres
sclerenchyma
long
as
sometimes
an
as
layer (in
adjacent epidermal
intergrading
in M.
—3,
epidermis. Midrib usually
prominently raised, supplied
than
palisade parenchyma cells,
collenchyma,
raised,
groups, often in 2
sometimes
of 2
of
layers
of spongy tissue
cells
and
5 times
to
inner and/or
Hypodermis
flattened
to
square
rarely higher
the
cells often
guatteriifolia only locally developed, especially
veins,
4 adaxial
sometimes up
times,
of M.
major
Mesophyll composed
and M. villosa 3
of
layer)
upright
thin cuticle, without
of one adaxial
(table IV), consisting
broader and lower,
a
subsidiary cells;
inconspicuous.
present,
abaxial
one
cells, usually
midrib
less embedded in
more or
less
cell, separating
bordering epidermal
sometimes slender, with
26). Subsidiary cells,
145
more or
small sunken
a
both lateral sides of the
on one or
alveolar material;
outer
cells often with
(table IV);
varying conspicuousness
present (photo
of Asiatic Myristicaceae
anatomy
sunken; bordering epidermal
Stomatal complex
papillae
Baas: Leal
as
palisade parenchyma,
adaxial epidermal cells, which then may be flattened and have
a
thin cuticle and thin and short cuticular flanges, also visible in cuticularmacerations
as
areas
of several,
epidermal
ground
in
cells
cells
less
deeply staining
(table IV) and/or
tissue of the midrib.
in
the
mesophyll,
parenchymatous
in
Minute,
most
cells;
small druses
more or
less
in
the
frequent
bundle sheath, and in the
sometimes
cells of the
parenchyma
present in
mesophyll
and the
spindle-shaped crystals, grouped
spongy
ground
tissue,
especially
in
the
tissue of the midrib, sometimes
BLUMEA
146
also present in
VOL.
-
where
epidermal cells,
they
shaped, especially
in basal cells of hairs
prismatic crystals
sometimes present in
Flattened
1
crystals with
sometimes in
Od)
Clustered
of
occur
Large,
more
in abaxial
prismatic
less
or
and in
mesophyll
maxima and
M.
epidermis,
few
oil cells
in
the
even
In the
pith,
generic descriptions and
been recorded. It is essential
species
(or
variety)
characters have
as
in
M.
More
in
thin
a
M.
cuticle,
less thick-walled
abaxially
of
the
towards the adaxial
Sclereids present in
epidermal layer.
brachysclereids and/or
tissue of the
sphaerosperma
edges.
abundant,
guatteriifolia,
or
and
'branching'
peripheral ground
in M.
with rounded
idioblasts with
most
Gymnacranthera,
a
THE
also in
sometimes
as
midrib, usually
ground
tissue of the
CHARACTERS
in the tables a very great number of characters has
analyse
a
which of these characters
and
genus
valuefor
which
is
list
are
in
not;
are
species (or variety),
or even
relationship
for
on
for
constant
are
other words:
a
possible
are
listed
to
a
limited
represented
following
those characters
by
their
are
extent
because
many
genus. This is
the
species
for
mentioned,
species (and,
specimens
two or more
constancy
species
which
or
not
are
a
which
and
of the characters the determination of the constancy for
varieties)
The characters
first
for
diagnostic
level. For
the
or
to
OF
for the determinationof patterns of
important
species (or variety)
In
(fig.
smaller in size.
sometimes
adaxially
into the
abundant,
VALUE
genus
less
Sometimes intermediate
midrib;
epidermis.
sometimes
protruding
astrosclereids in
to
grouped
crystals
present,
Empty
occur
sacs,
bundles,
sometimes
needle-
veins (table IV).
major
also
usually
and
pith.
sometimes
to
more or
corroded appearance
a
when
M. fragrans.
crystals
sometimes present in the
in M. maxima in the
abaxially,
cells of
villosa also
specimens only,
clustered brachy-
often with
tissue of the
caps of the vein
spindle-
epidermal cells, mostly solitary (table IV).
and clustered
spherical
and
larger
relatively few, large, irregularly shaped particles
epidermal
tanniniferous
probably
sclerenchyma
a
of
1981
be
peripheral ground
probably secretory,
cells,
may
1,
adaxially (table IV). Flattened,
edges,
6 rounded
—
No.
epidermal cells, mostly solitary,
crystals, composed
(fig. lOe) present
forms
4
27.
in this
a
for
study.
genera.
constant
for
a
species.
List
Pitting
of cuticular
Presence of
Shape
of
abaxial
of
cells
in
(papillae excepted)
ledges
prominence
of
interruptions
bundle of the
*)
Does
not
apply
midrib
in the adaxial
midrib*)
for
Number of
of
phloem
groups in
Gymnacranthera.
phloem
bundles
in the
pith
of the midrib
Presence
the
adaxially*)
Presence of
layers
the collateral bundles of the midrib
epidermal
section
Presence of stomatal
Degree
Number of
flanges
polar T-pieces
transverse
1
of
locally
developed
dermis in the midrib
Frequency
of druses
Frequency
of oil cells
Presence
of tanniniferous
sacs
hypo-
J. K.OSTER &
The characters,
P.
Baas:
Leaf
mentioned in list
anatomy
I
do
147
of Asiatic Myristicaceae
not
have
diagnostic
any
or
taxonomic
value.
The
next
for the
characters
applicable
are
to one or two
and
genera only,
not constant
are
species (or varieties).
List 2
Basal cells of hairs in
a
row
Minor veins
(Myristica)
Difference in size of adaxial and abaxial
cells
epidermal
Presence
of
adaxial
Presence of
(Myristica)
in
stomata
adaxial
the
of
giant
bundles
stomata
(Gymnacranthera)
The characters of list 2 do
the
value, especially
the
frequent
one
or
two
but for
genera
a
diagnostic
they
walls in
adaxial
the
and
midrib
abaxial
(Horsjleldia,
occurrence
or
are
few
within the
applicable
to
taxonomic value on the
these
'prominence
genera
are
species
of the minor veins',
concerned.
each genus and
genera
of the
indicative circumstantial
some
species (or varieties) they
especially,
transcurrence
(Knema, Myristica)
may have
latter features and the
two
The characters of list 3
species,
have
not
genus level
because of their
of the
of
in
of vertically
veins
on
the
( Knema)
Knema)
Degree
level either, but
not
periclinal division
Connection
water
or
in
prominent
or
hypodermis (Horsfieldia)
the
epidermis (Gymnacranthera)
Presence
slightly
epidermis
are
not constant
for
most
When the latter applies
are.
to
mentioned.
List 3
Cuticular
layer granular
or
Presence
( Gym-
not
midrib
nacranthera, Horsfieldia)
Sizes
of
guard
cell
pairs (Gymnacran-
of
in
the
of
pith
the
of
in
chlorenchyma
the
midrib
cells
secretory
(Hors-
Shape
fieldia)
of veins
adaxially
(Gymnacran-
thera)
Length/width
chyma
xylem
(Myristica)
Continuity
thera, Horsfieldia)
Frequency
of
cells
ratio of
palisade
Size of druses
paren-
Position of
(Knema).
Relative size of midrib
the
(Myristica)
brachy-
midrib
and
to
astrosclereids in
major
veins
(My-
ristica)
These characters have
species
or
In the
to
variety
following
species.
are
The latter
characters, without
either
acquisition
(very little) diagnostic
some
particular
list those characters
each genus, and which
for all
some
level in
them
or
constant
species
having
are
for
may be
on
the
genera.
mentioned,
most
of the
thought
become fixed
gradual loss).
and taxonomic value
to
in the
which
are
usually applicable
species (or varieties),
show
a
'tendency'
speciation
process
but
not
for these
(through
BLUMEA
148
VOL. 27,
No.
1.
1981
List 4
Number
of
basal
of
parts
hairs
Degree
of covering of the adult leaves
Leaf
by
walls
Anticlinal
Thin
grooves
of
the cuticle
cells
epidermal
or
Thickness of the lamina
of
of
layers
of
palisade
Presence
paren-
The characters of list 4 have
the characters
are
by ecological
factors,
Myristicaceae
not
of oil
thought
to
cells in
the
of the
pith
of
diagnostic and taxonomic
very strong, however:
and
some
most
of them
of them may be
brachy-
value
can
astrosclereids
to
on
the
possibly
age-dependent
species level;
be influenced
in the
persistent
leaves.
In list 5 characters
for
and
spindle-shaped
midrib
Frequency
constant
epidermis,
to
not
prismatic crystals
cells
chyma
flattened, when druse
adjacent
are
Frequency
Thickness of the cuticle
Number
cells
Epidermal
not
or
peripheral
tissue of the midrib
idioblasts
of cuticle present
not
or
of sclerification of the
ground
not
or
areas
in
sclerified
margin
Degree
Presence of
central
spongy tissue
the hairs
sinuous
of sclerification of the
Degree
per
surface unit
most
show
are
of the
a
mentioned, applicable
The
species.
tendency
species,
to one or two
for which
genera only, which
character is
a
inconstant,
are
are
for that character.
List 5
Presence
of cuticular striations
Hors-
(
Presence of
fieldia, Knema)
Number
matal
of
pith
cells,
complex
the
bordering
the
(forming
Minor veins
sto-
of
'ring';
on
sunken
slightly
Presence
the level of the
of
a
{Horsfieldia,
Myristica-,
Myristica
is
Presence
of
Presence
locally
bundles
as
fibres
peculiar
cells
of
sclereids
hypodermis
the
as rows or
epidermal
or
( Horsfieldia)
continuous
developed hypodermis
Spongy
epidermis
xylem
sclerenchyma
Presence
Myristica)
Stomata
large
in the
of the midrib (Knema)
small bundles
(Knema)
crystals
in
the
(Myristica, Knema)
relatively
thick
filiform
cells
in
(Knema)*)
of
swollen
the
subepidermal layer (Horsfieldia)
in Knema and
highly inconstant)
tissue
compact
or
not
{Horsfieldia, Knema)
Abaxial
of
layers
rified
or
not
spongy
tissue
scle-
(Gymnacranthera,
Knema)
These characters also have
level,
*)
but
only
some
diagnostic
and taxonomic value
in certain genera. The value is low
The thin filiform
sclereids
in
Gymnacranthera
are
on
the
not meant here
on
genus level.
the
species
J. Koster &
The
characters
following
constant
for the
P.
species,
Baas:
anatomy
applicable
are
but
Leaf
of Asiatic
to
one
for the genera in
not
149
Myristicuceae
two
or
genera
and
only,
are
question.
List 6
Type
of hairs with cells with
one
Presence
arm
(Horsfieldia, Knema)
Basal parts of hairs often
joined
or
stomatal
Basal parts of hairs, when
cell, very small
one
cells
of
hairs
unspecialized
cork
regular
many
of
composed
or not
epidermal
not
warts
than
cells
or
or
chyma
not
or
(Knema)
with
veins
Major
not
(Myristica)
conspicuous
sclerified and with
parenchyma
large pits
the
the
cells, bordering
complex,
(Myristica)
Palisade
(Myristica)
higher
of
Upright papillae
not
(Horsfieldia)
Basal
of
(Horsfieldia, Knema)
little
very
with
or
scleren-
(Gymna-
caps
cranthera)
The characters of list 6 have
certain genera;
The
species
on
few characters
next
and for
diagnostic
and taxonomic value on the
level in
species
the genus level the value is low.
most
are
applicable
to
each genus and
are
for the
constant
of the genera. The genus for which the character is inconstant is
mentioned.
List 7
Basal
Alveolar
not
of hairs
parts
present
or
not
adaxial surface
on
material
cells
obviously present
and
(Horsfieldia
of
Degree
(Myristica)
being
in
embedded of the
the
guard
cells
subsidiary
(Horsfieldia)
or
Stomata sunken below
Myristica)
epidermis
Midrib
or
not
the level of the
(Horsfieldia)
raised
adaxially
not
or
(Horsfieldia)
These characters have
genera,
and taxonomic value on the
diagnostic
for which the characters
are
inconstant;
species
level for the
the characters also have
and taxonomic value on the genus level, but this value is
not
of the
same
diagnostic
degree
for
each genus.
In the
constant
following
list those characters
for each genus, but
are
are
which
mentioned,
inconstant for the
are
applicable
to
and
family.
List 8
Basal
of
parts
frequent
on
Arms of hairs
the latter
hairs
the
most
midrib
one or two
case,
of the
or
least
abaxially
size
or
of
bordering
per cell, and, in
same
Structure
not
Major
or
veins with
caps
(and
Position of
The characters of list 8 have
diagnostic
stomatal
complex,
the
cells included
big
sclerenchyma
the size of the
druses in the
and taxonomic value
on
sheaths
caps)
mesophyll
the genus level.
BLUMEA
150
In the last list
characters
some
and which
only,
are
No.
1,
1981
mentioned, which
applicable
are
to one
genus
for that genus.
constant
are
VOL. 27,
List 9
vascular bundle in
Adaxial
the midrib
Filiform,
absent ( Gymnacranthera)
Phloem
groups
midrib small
of collateral
The classification of
characters,
noted that
far
as
Dicotyledons
for
possibilities
Rijksherbarium
(Baas,
1975),
the
taxonomic
general
Icacinaceae
(Van
the genus
not
strict.
the
midrib
& Baas,
Baas,
and
1973)
Den
1973;
more
that
likely
the
throughout
confirms
proposed
Vliet &
is
the different levels. It may be
occurrence
& Baas,
level.
When
it
studied,
in
the
evergreen
limitations and
studies
earlier
of widely differentfamilies such
(Jansen
Staveren
on
analysis
as
purposes
representatives
Celastraceae
the
of
on
is
would have been
above
1978), Crypteroniaceae (Van
Baas,
tissue
outlined above,
be inconstant
to
concerned,
on
sclereids
and
the
and taxonomic value
as
characters of
as
are
branched
mesophyll
ground
species
more
characters would appear
more
rarely
in
(Gymnacranthera)
diagnostic
and
species
per
bundles in
( Horsfieldia)
These characters have
specimens
present
Hartog-Van
Ter Tholen &
1975), Guttiferae (Baas,
Winteraceae
the
in
Aquifoliaceae
as
1970),
1973; Vink,
(Bongers,
1977).
important
Some
Knema' and
have
Horsfieldia
studied,
papillae
in
Sinclair
examined have
reasons
are
to
at
most
moreover
papillae
papillae
species
in
dome-shaped
full range of
the
occur
occurrence
of
Horsfieldia,
not
unfortunately only
M. gigantea and
unspecialized
M.
one
in
cells.
species.
to
species
Myristica species
epidermal
cells
specimen
conspicuous
maingayi,
the other
for these
are constant
have such
astrosclereids
is unknown. The very
slightly dome-shaped
do
of
lists.
closely related, whereas
as
suppose, that the
different: the
genus;
(1968)
the
foregoing
of which
species
the constancy for the
so
Myristica, however,
regarded by
and
included in the
have been recorded in
Papillae
has been
epidermal papillae (in species
respectively),
been
not
as
the type and distribution of the alveolar material (variable in
Knema
and
Myristica
characters such
Myristica),
papillae
pronounced shapes
This
In Knema
as
in this
occur
in
gives
things
Myristica.
These facts make the constancy of this character in Knema doubtful. The constancy
for
Horsfieldia
papillae
reflecting
The
of the
alveolar
studied with
only
one
specimen
of this character for
over
the leafis
restricted
plumulosa,
placed
these
for
constant
C.
In
to
a
a
these
species
In
scanning
they
(sub)species
character
electron
species
per
(a
but
were
into
regarded
taxa
variable in
microscopy
Baas,
as
in
of lower rank.
Myristica)
only; however,
in this
has been examined. Therefore the constancy
the stomatal
study by
series. This
Winteraceae
alveolar material is
be
can
with the
is uncertain. The distribution of the alveolar material
bordering
one
1970)
possibly
character variable in Knema. The presence of the material is
the cells,
represented
species.
material
species
Cratoxylum (Guttiferae,
formosum,
further differentiation of the
a
type
thoroughly
S.E.M.
is unknown.
iryaghedhi
not
are
(Bongers,
variable in
some
one
complexes
specimen each;
suggests
1973)
species.
a
mainly
in K. intermedia and
De
Wilde
(1979)
K.
has
constancy of this character for
the distribution and the type of the
J.
Koster &
P. Baas:
Lea/
Astrosclereids have been recorded for
superba;
only
according
be
specimen
one
Sinclair
to
for these
constant
in Knema have either a
leaf are
on a
constant
as
far
and H.
species
are,
expected
some
The
and
of
Specimens
numbers,
cells,
one
whereas
frequently
most
Myristica species,
trend for Knema
inconstant
species.
of the cuticle in Knema
sculpturings
to
(not
and Knema
Horsfieldia
occur.
smallnumber of basal
a
Gymnacranthera
is
rufa
only specimen examined of this species, while
been observed in other
not
These
the constancy is concerned: The
as
Horsfieldia species.
for
convex
because it is the
unknown,
number or
high
and reflect
Horsfieldia species
The constancy of the
has
studied.
with small numbers also
species
this character is inconstant for
for
been
The character 'number of basal cells of the hairs'
species.
but in these genera
species,
found
has
numbers of basal cells per hair have been counted in
highest
151
Myristicaceae
related. Therefore this character is
(1958) closely
mentioned in the lists either) is difficult,
species
Asiatic
species only: Horsfieldia fulva
two
species
per
of
anatomy
simply
the feature
species.
The feature 'isobilateral leaf' (recorded for
Horsfieldia sabulosa)
will be discussed
elsewhere.
TAXONOMIC
Introductory
In the
on
an
remarks
following discussions, suggestions
evaluation of the
species,
IMPLICATIONS
genera
shared and
and families. With
our
anatomy, this phenetic approach is
cladistic
methods
matics in the
(cf.
sense
Bremer &
of
whether certain character
types in Knema
are
states
perhaps
will be further
significance
feasable than an attempt towards
plea,
Wanntorp's
mainly
based
because there is
and their
this,
to
applying
1978, for phylogenetic
should be considered derved
exception
an
only
leaf anatomical characters of
present stage of knowledge of systematic leaf
more
This is
Hennig).
about natural affinity will be
distinguishing
or
no
way
of
syste-
telling
ancestral. The hair
possible phylogenetic
explored.
In view of the limitationsof the research materials, the taxonomic suggestions
can
be tentative.
only
Below
the
genus
level
Gymnacranthera
This genus is
this
because of
of hairs
some,
per
the
fairly homogeneous:
G. bancana may be
context
mostly epidermal
surface
unit,
species
regarded
as
features:
cuticular
layer
differ
taking
a
only
relatively large
not
granular,
cuticle, anticlinal walls of unspecialized epidermal cells
and
the
chlorenchyma
recognized
the
by
of the
ground tissue
features
of G.
are
bancana
var.
findings
as
midrib,
sometimes also
forbesii
These
the
forbesii
are
most
other, especially
continuous
not
high degree
G.
and the
in
the
a
a
in
few
parenchyma
G. forbesii,
specimens
agreement
cells
but
of G.
and G.
forbesii.
thick
G.
less
adaxial
sinuous
conspicuously
eugeniifolia
can
be
tissue, the peripheral
margin.
extent.
contracta
are
(1958a, b);
distinct species, and the remaining species
eugeniifolia
position
number of basal parts
of the leaf
to a
with Sinclair
few characters. In
relatively
not
midrib.
a
less isolated
of sclerification of the spongy
present
and
partly
in
in
a more or
as
One
These
specimen
very similar.
he
regarded
very close
to
G.
each
152
BLUMEA
Where all Malesian
to
give
a
key
be
1.
1981
and their varieties have been examined, it
species
and varieites of
species
and
species
leaf
separated
No.
enable identification on leaf anatomical characters. A
to
the Malesian
to
paper. All
VOL. 27,
-
varieties, except
is
Gymnacranthera
G.
and
contracta
given
seems
useful
synoptical key
the end of this
at
G. forbesii var.forbesii,
can
anatomically.
Horsfieldia
It is
possible
surface unit
H.
to
distinguish
features
epidermal
H.
of hairs
smooth)
and
papillose
and stomatal
are
distributed
species, which
H.
the
grooves,
H.
a
many
more
unit
all
abaxially;
abaxial
sylveslris
mesophyll.
epidermis
has some, for
too
tissue
spongy
surface),
and abaxial cuticle
more
less
or
Other characters
less reticulate pattern, but there
or
characters in
important
are
common:
share the presence of cuticular striations and
sylvestris
sunken
slightly
in
species
and
has isobilateral
on
of hairs per
the abaxial
on
adaxially
H.
anatomically:
parts
clearly present,
often robust
stomatal
cuticular striations and the
prominent
surface
(hairs
relatively
and
crassifolia
material
only species
as
the
over
have
joined together
complex distinctly sunken);
features
unique
H. sabulosa
compact);
basal
numerous
of many cells
composed
iryaghedhi (alveolar
H.
examined of Horsfieldia
species
and basal parts often
adaxially
glabra (basal parts
Horsfieldia
most
crassifolia (relatively
with
complex
relatively
epidermal
the
characters,
parts
recorded
not
cells
subsidiary
basal
many
with
of hairs
for
the
per
other
Horsfieldia species examined.
H.
in
crassifolia
as
common,
and H. sabulosa also have
the
H. sabulosa and
(and unique
of adult
for
leaves,
some
relatively
thick
H.
share another
glabra
Horsfieldia)
many
are
regular
(in
view
our
unimportant) features
cuticle and lamina.
epidermal:
cork
warts
of
set
hairs
characters;
abaxially and
the
important
most
absent from the lamina
completely
many secretory cells in the
epidermis.
H.
and H.
fulva
the other
by
Sinclair
according
H.
not
examined
a
1975),
to
Sinclair,
are
placed
are
on
concerned. The
one
Bivalves
crassifolia,
Section
H.
H.
irya
H.
sylvestris
H.
iryaghedhi
H.
macrocoma
H.
sabulosa
special
relationship
relationship
and
H.
the
by
crassifolia
sabulosa,
species
Sinclair.
and H.
leaf anatomy. H.
between H. fulva and H.
between
as
pairs
Our results
by
this
mentioned
do
not
superba
and
crassifolia
suggested
of the
irya.
H.
H.
study,
were,
support
a
is
supported by
sylvestris,
has
not
however,
is
a
species
with
a
and
been
as
the
species,
placed
deviating
not
this
study.
between
recognized
special relationship
Also the division into the sections does
iryaghedhi
far
as
related
superba
H.
The close
closely
Trivalves
fulva,
glabra
section
mentioned
recorded for
line.
H.
Sinclair;
not
given here,
more
parviflora
glabra
characters
character
this classification is
H.
A
the
on
mesophyll,
results, outlined above, with the classification of Horsfieldia,
(1958a, 1974,
examined leaf anatomically
species
Section
(species
Horsfieldia species.
To compare the
made
superba
both have many astrosclereids in the
above)
in
the
H.
by
same
between
H.
appear from the
inflorescence
(Sinclair,
J. K.OSTER & P.
it has
1975). Although
the
other
the
species,
Asiatic
Leaf anatomy of
of
majority
the
153
Myristicaceae
leaf antomical characters,
important
some
Horsfieldia
Baas:
recorded for
not
characters is
in
range of
the
Horsfieldia.
key
No
species
the
to
species
of
is
Horsfieldia
small number of
because of the
given,
examined.
Knema
Only
a
few of the Knema
examined
species
K. laurina have
unique
has
peculiar crystals
the
mesophyll and
Some other
K.
point
(the
distribution of
kunstleri
to
a
with
Most other
the
these characters
with
diagram
the
alveolar
cork
less
(c. 20, largely
compartments
with many features in
formed:
Wilde's
Group
K.
K.
K.
K.
linifolia
has
percoriacea
for Knema
many
unique
and
hair
type 6);
mandarahan and
and
K.
(The
characters
constant
each character
gives
K.
K.
sclereids in
features
tenuinervia
and
conferta
and
type 4);
(hair
for each
roman
—
distributed in
6 of the
point
species.
to
a
are
K.
behind
the
close
species
listed in
there appear to be
relationship,
few features. In this
of
way groups
numbers
But when
refer
to
the
1
Group
2
Group
whereas
series
3
K.
ashtonii
II
K.
malayana
XI
latifolia
V
K.
attenuata
X
K.
oblongifolia
I
linguiformis
VI
K.
elmeri
II
K.
globularia
IV
K. laurina
I
K.
IV
oblongata
K.
pseudolaurina
I
K.
pulchra
IV
K. retusa
II
K. tomentella
X
Group
4
K. rubens
Group
5
a
species
species
XI
6
K. intermedia
IX
K.
conferta
XII
K.
furfuracea
III
K.
korlhalsii
IX
K.
kunstleri
XII
K. hookeriana
III
plumulosa
IX
K.
K.
reticulate pattern
a
previous chapter)
possible species pair
which may
none or
are
VI
Group
(which
plumulosa
classification).
curtisii
the
pallens (sclerified
another classification of the
from lists 3
common,
other combinations share
been
K.
couple of,
material
on
adaxial cuticle.
for instance: K. intermedia and K.
warts);
K.
recognized
be
cinnamomea, K. korthalsii and
combination of types, K. hookeriana
unique
sculptured,
a
var.
psilantha (hair type 5).
more or
species:
a
epidermis,
one, or a
large pits);
lunduensis and K.
over
or
special relationship),
(many regular
mesophyll
has
rufa
share
species
may
hair types
in the abaxial
individually
can
basis of leaf anatomical characters: K. ashtonii
K.
linifolia
I
K.
lunduensis
III
K.
mandarahan
IV
K.
pallens
III
K.
percoriacea
III
K.
psilantha
III
K.
rufa
IV
K.
tenuinervia
IV
have
of De
BLUMEA
154
The first group contains
lamina thin
leaf
margin
the
linguiformis
1.
conspicuously
not
1981
which have the characters: hair type 7, 8a
species
midrib
sclerified,
not
No.
adaxial cuticle less than
/an),
than 5 pm thick, adult leaves
and
VOL. 27,
weakly
cells
palisade parenchyma
10 pm
covered with
hairs,
sclerified. In
K.
spongy tissue
curtisii
than wide,
scarcely longer
are
8b,
or
abaxial cuticle less
thick,
and
K.
unique
a
feature in Knema.
The second group contains
This group exhibits the
lamina usually intermediate
<5
margin
adult
pm,
leaves
sclerified
weakly
important characters,
shared
<
pm,
200
each other in
midrib
not,
with
usually
cuticle
hairs,
are:
8b, thickness of
or
10 /im, abaxial
<
tissue
spongy
and
sclerified.
moderately
of this group
by species
reticulate pattern.
a
1, 2, 7, 8a
/on), adaxial
covered
usually
or
to
features: hair type
120
(>
cuticle
linked
species,
following
relatively
numerous
basal
parts of hairs per surface unit abaxially, spongy tissue compact, anastomosing
as rows
of the midrib. K. elmeri and K.
pith
similar. The second group shares
The third group contains
thickness of lamina
not
covered
such
The fourth group contains
and/or 8c,
<
characters:
is rather
group
hair type 7
abaxial cuticle 4
—
weakly sclerified,
6 /mi
leaf
group, K. kunstleri in
the
and K.
compact spongy
covered with
The sixth group contains
which have the characters:
adaxial cuticle
usually
(strongly) sclerified,
shares
>
leaf
species,
and
7
spongy tissue and
leaf
which
species
have
>5
/mi), adaxial cuticle
or
the
/mi,
not, spongy tissue
not, midrib moderately sclerified. This
or
linked
3, 6,
10 pm, abaxial cuticle
<
contains
(>200
characters with
some
hair types
have
tissue.
hairs,
the sclerification of the abaxial
as
characters
particular
sclerified.
thick
sclerified
adult
<5m,
some
laurina in
adult leaves covered with hairs
thick,
and/or 8b,
which exhibit the features: hair type
heterogeneous
particular,
the sixth group, such
Other
as
species,
margin
group.
sclerified. This group shares
8a, lamina
or
appear
sclerified, leaf margin
spongy tissue
oblongifolia
conspicuously
not
particular
of the first
adaxial and abaxial cuticle
hairs,
sclerified, midrib (moderately)
not
fifth
The
species
thickness of lamina intermediate, adaxial cuticle
5 pin, adult leaves
margin
with
moderately
features,
common
in
retusa
characters with
which have the characters: hair type 7
of the second group; K.
species
certain
species
some
usually intermediate,
conspicuously
not
sclerified and midrib
with
veins
of sclerenchyma fibres, large xylem bundles in the pith of the midrib and oil
cells present in the
leaves
leaf
Other
each other in
to
5
3, 4,
5 /mi, adult leaves
or
not
of the spongy tissue.
reticulate pattern, and
a
8c, lamina intermediate
covered with
margin usually sclerified,
important characters, shared by species
of the third and
species
layers
midrib
of this group
hairs,
(moderately)
are:
to
thick,
spongy tissue
sclerified.
sclerification of the
abaxial layers of the spongy tissue and large xylem bundles in the pith of the midrib.
K. furfuracea,
in K.
long
pallens
as
wide,
characters in
species
K.
a
K.
hookeriana,
and K.
percoriacea
Some
of the fourth group,
De Wilde
(1979),
as
of series VII and VIII
refers
to
the
and K.
far
were
as
species
especially
are
with
by
us.
in
particular
cells
and K.
group share
are
up
rufa
some
to
are
similar;
five times as
also have
many
characters with
K. korthalsii.
below with the classification
examined in this
studied
leaf anatomical group.
linifolia
of this
compared
species,
not
percoriacea
palisade parenchyma
feature in Knema. K.
unique
common.
The results outlined above
by
pallens
the
study
are
proposed
concerned.
The number behind each
Species
species
name
J. Koster &
1: Series
K.
K.
K.
P. Baas:
Asiatic
155
Myristicaceae
Series Sericeae
II:
Laurinae
Leaf anatomy of
III: Series
Lamellariae
linifolia
6
K. elmeri
2
K.
hookeriana
6
oblongifolia
3
K. retusa
2
K.
furfuracea
6
pseudolaurina
2
K. ashtonii
2
K.
pallens
K. laurina
2
K.
K.
IV:
Series
Obovoideae
V: Series
Latifoliae
2
1
K.
oblongata
2
K.
mandarahan
6
VI: Series
K. tenuinervia
6
K.
linguiformis
1
K.
rufa
6
K.
curtisii
1
K.
globularia
2
latifolia
XI:
percoriacea
6
IX: Series
pulchra
K.
6
lunduensis
K.
K.
6
Mamillatae
plumulosa
4
K. intermedia
4
K. korthalsii
4
K.
Curtisianae
6
psilantha
XII: Series
X:
Series Glomeratae
K.
tomentella
2
K.
rubens
3
K. kunstleri
5
K.
attenuata
2
K.
malayana
3
K.
conferta
5
The classification based
Wilde's
classification.
discussed,
(1958a,
this
Wilde describes
K.
a
by
to notes
linifolia
K.
and K.
oblongata
common
supported
oblongata) together
classification
with K.
will
be
The classification of Sinclair
(1979).
with K.
with K.
close
laurina,
study.
and
K.
K. retusa,
K.
resembling
(according
oblongifolia
leaf anatomical
common
globularia
K.
placed
and K.
rubens,
in the
species
of
close
vegetative parts,
attenuata
to
to
species
having
as
globularia
According
one.
with K.
these
tomentella and other
K. curtisii in the
has
De
a note
tenuinervia.
in series IV
placed
were
Sinclair
K.
laurina,
the
to
and K. tenuinervia. In
linifolia
pulchra
whereas leaf
isolated position,
an
rufa
De Wilde considered K.
this
by
has characters in
oblongifolia
resemblances
whereas in leaf anatomical aspects
series)
affinities with his series X and XI, and
is
De
supports
generally
particular
in series I, albeit in
resemblance between K.
leaf anatomical group 2.
which
and
De Wilde
together
goes
heterogeneous
have much in
the
K.
species
a
globularia,
De Wilde
differences
Punctatae
will also be cited, if this is relevant.
De Wilde placed
anatomically
leaf anatomical characters
on
Some
with reference
1961)
Series Glaucae
in
(and
one
De
K.
group,
Wilde,
same manner as
a
K.
in leaf
anatomy.
De Wilde created
a
series for K.
in accordance with this
Not half the number of
examined in this
leaf anatomical
study.
latifolia, considering
of Knema, included
species
Therefore it is useless
groups,
however,
may
help
parallelism
of the classifications based on
characters,
there is
De Wilde
species
as
related
chance that the
to a
examined. For
De Wilde's series
of Knema is
A
a
tentative
at
the
to
to
give
a
by
key
as
restrict the
De Wilde
to
the
related which is
macromorphological
species,
not
it
may be
A
key
the
In view of the
study, regarded by
fits the group of the
that K. lamellariade Wilde
the leaf anatomical group 6. A
been
to
and leafanatomical
examined in this
hoped
(1979) has
species.
possibilities.
examined leaf anatomically,
species,
example,
III) fits
given
series VI
study.
synoptical key
to
the
(from
groups
end of this paper.
phylogeny
The indumentumof Knema
can
of
the
hair
types
be classified in types with
in
Knema
(very) long
hairs
(type
3, 4, 5, 6 and 8c) and types with shorter hairs (fig. 3). Leaves with long hairs loose
156
BLUMEA
these
hairs when
they
retain them
usually
least
Between type 8a and 7
27,
No.
the adult stage,
come to
(at
VOL.
1,
1981
whereas leaves with shorter hairs
partially).
there is
(short hairs)
hairs of type 8a small hairs also
distinction. On leaves with
sharp
no
The classification into these types is artificial
occur.
and made for identification purposes. Hair type 8b and
least
specialized
structures
types, having
(as long stalk cells, long
then
occurrence)
rare
hairs with
been
of the
indicated in
with
part
arms.
c) through
peculiar
through
The types
between
are
the 'forerunners' of
&
laurina
hookeriana,
3 and
are,
latter
and
1 and 2
ashtonii
K.
korthalsii,
K.
according
to
K.
var.
pallens
De Wilde
however, has
species,
and K.
hairs belonging
one
to
series
It is remarkable that the
mandarahan and K.
the
(type 8c,
species
groups
with
type 8c.
The
species
group
type 8b)
these
presumably
one
to
are
very
K.
The
6.
belonging
K.
types.
belonging
types
each
to
excepted.
K.
type
The
and
linifolia
K.
remaining
These three
a
hairs all
long
of
group
particular
just mentioned),
species
both have
with K. korthalsii.
species
but without
long hairs,
4).
type 5 and have
to
together
with the derived hair types with
and
Only
furfuracea,
intermedia and K. plumulosa
in De Wilde's series IX,
4
3.
fig.
relationships
derived hair
most
K. korthalsii
three
both have hairs
This supports the
of group
without
species (or variety)
have hairs
ancestral form of the hair types
groups and their mutual
within
are
these species have been discussed
type 4;
to
the hair types of K. korthalsii (of group
ones
The
belonging
also have
rufa,
putative,
are
is outlined in
here,
percoriacea
De Wilde. K.
by
type 6 and
leaf anatomical
7 and 8a
for the determination of
(1979), closely related,
hairs
psilantha
in
short hairs
cinnamomea, respectively).
before. K. lunduensis and K.
placed
these
(the types
c
represented by
are
belonging
the
proposed
as
tenuinervia both have hairs
been
and
types
cells.
types,
Wanntorp, 1978).
1, 2, 3, 4, 5 and 6. Type
(K.
no
and 2 may also have been derived from the types 7 and 8a
1
of the hair
(Bremer
taxa
that
8b and
type
derived characters should be used
uniquely
stalk cells)
(long
derivations of these
possible
are
may have been derived from the types 8b
(7 and 8a)
extension of the stalk
phylogeny
The
Other
relatively
have
may also
species each)
one
a
There
c.
5 and 6. The other
3, 4,
through specialization
reduction; another possibility is
structures are
common!).
(b and)
the
as
special
3.
fig.
The types with short hairs
(and
c)
regarded
hairs with
long
the distal end etc., types with
at
1 and 2, recorded for
(type
type 8b (and
be
may
have been derived from type 8
may
stalk cells
long
derived from
reduction
arms
c
The
structures.
propose evolution lines between the types
to
argumentes
peculiar
no
moreover
natural character of these
are
6,
in
K.
structure
like
are
one
the
of
only
leaf anatomical
relationship.
1,2,3
and 5 have shorter
hairs, belonging
2 the derived hair types 1 and 2. Hair type 7 and 8a
in different groups, which may be
an
to
different types,
occur
(together
argument for the primitive
with
nature
of
types.
Myristica
The
have
species
some,
leaves, basal
in
mesophyll
abaxial
examined of
for
Myristica
are
fairly
Myristica unique features,
cells of
not
hairs,
if
solitary,
large) and
hypodermis present,
M.
very
e.g.
distinct
small;
sphaerosperma
sclereids
in
anatomically.
M. villosa
major
Some
(remarkable
species
hairs
on
the
sclerified and druses
spongy tissue
(guard
veins
cell
and
pairs relatively long,
druses
in
the
abaxial
J. Koster &
M.
epidermis);
fragrans,
P. Baas:
Lea)
only
as
anatomy
has
species,
157
of Asiatic Myristicaceae
clustered
in
crystals
the
abaxial
epidermis.
M.
and M.
gigantea
maingayi
are
Most other
more or
less
characters
constant
question
reticulate pattern. The characters in
adult
on
cells
epidermal
on
etc.
sclerification of the
spindle-shaped
characters
diagram,
cells
As
in
the
done for Knema, it appears that
'character
own
none or
complement';
of
a
crystals
a
in the
the
hypodermis,
frequency
of the
mentioned. When
these
previous chapter)
some
in
thickness of the
unit,
presence
be
to
species
presence of hairs
epidermal:
per surface
7 of the
—
epidermis).
distributed over the
often
have
mesophyll
from lists 3
features in
rare
tissue of the midrib, and the
peripheral ground
crystals
some
abaxial
characters the presence of
non-epidermal
whereas others share
common,
are
are
bordering the stomatal complex,
(c. 25, largely
as
papillose
a
number of basal parts of hairs
leaves,
cuticle, papillae
has its
similar and have
very
(as grooves in the adaxial cuticle and
common
species
have
many
few features. Of the former
nevertheless groups of
listed
are
each
species
species
can
in
a
features in
pair
be formed.
Not each character involved will be mentioned in the discussion of the groups, but
those which
only
are
more
or
M. chrysophylla, M. fatua,
globosa,
M.
and M. subalulata have many
lepidota,
features, in particular: M. chrysophylla and M. lepidota,
common
lepidota; M. fatua
and M.
M. subalulata. The
crystals
in
crystals
play
important
an
and M.
lepidota;
important for
with
sphaerosperma
M.
(such
adult
M.
leaves)
fatua.
features in
It is
the
are
M.
as
and
M. villosa have many
basal cells
mm
abaxially
striking.
cinnamomea and
The
M.
two
higher
too
has
to
than
study
are
I:
series Maximae
or
sphaerosperma
M.
chrysophylla
series Fatuae
M.
M.
lepidota
fatua
villosa
far
as
the
series Cinnamomeae
series Littorales
M.
9:
maingayi
M.
as
Myristica
8:
Maingavae
gigantea
series Fuscae
M.
some,
M. cinnamomea
Section
II:
mainly
common
features;
and many hairs present
have also
guatteriifolia
series
M.
10:
more
given here,
7:
M. maxima
M.
30
concerned.
Section
series
of
on
mainly epidermal
(1958a;
with the reticulate distribution of the characters
in this
M.
M.
the
unspecialized epidermal
results with Sinclair's revision
compare these
struggled
and the series. His classification is
4.
couple
again
latter Matures have also been found in
maxima
species
1:
a
and
species.
common.
interesting
Sinclair
most
this group of
shares
2
cells, basal parts of hairs/0.1
globosa
of this group, and
species
and M.
M. fatua
and M.
chrysophylla.
guatteriifolia,
the hairs
concerning
are
common
role.
characters with M.
M. cinnamomea. M.
globosa
particular
has many features in
epidermal
M.
globosa;
fragrans
those
less characteristic for the groups.
M.
Fragrantes
fragrans
II: Fatua
14:
series Cimiciferae
M.
15:
globosa
series Subalulatae
M.
subalulata
species,
1968).
over
the
examined
BLUMEA
158
The presence of M.
this
by
also
study.
some
deviating
and M.
it is
distinguish
this
position,
notes;
he
so
globosa
M.
from the
Apart
sections is
common
anatomical
species
to
that the
series
while
case
9
as
he has
this
M.
to
and
fragrans
varieties examined of M.
to
the
species
of section
villosa)
alone will
species together
in
one
from the
differing
others in
those of
to
present results.
the division into
villosa,
the
two
to
series 8, series 10 is
globosa
and
M.
nearest
is
globosa
series 11, M.
to
also
similar
to
The
supported by this study.
are
M.
two
rather different, which is in agreement with the
are
characters.
macromorphological
key
fatua
M.
a
with M.
some extent.
to
Sinclair series 7 is close
has resemblances
M.
has leaves somewhat similar
in agreement with the
study,
common
within the sections and series
distinct series,
a
M.
M. fatua
but M. villosa takes
than with
these
placed
has
some
from
deviates
features in
I)
chrysophylla
nevertheless
some comment:
characters,
relationships
subalulata in many ways; all these remarks
No
has
species
more
of section
M. fragrans
of M.
has
strongly supported
and M.
vegetative features (of
fatua,
this is
lepidota;
supported by
According
lepidota
this
too
respect
his ideas about
regarded
and
sphaerosperma
while this
from M.
given
section I in
many ways,
M.
M.
guatteriifolia (species
species
series. Sinclair has
sphaerosperma
support.
Sinclair stated
Although
series is
one
of M.
ecological
have many
lepidota
in
maingayi
1981
1.
mountain species. The series 11 needs
a
cinnamomea and M.
in
in
characters;
somewhat distinct
II.
relationship
leaf anatomical
chrysophylla:
and M.
gigantea
The close
No.
VOL. 27,
species
of
is
Myristica
because of the small number of species
given,
examined.
On
the
The
level
genus
Asiatic
characters
genera of the
Myristicaceae, unique
This
applies
with cells with
midrib
in
table V.
two arms
to
Gymnacranthera.
of different
Horsfieldia
genus has
flat
absent, major veins
to
with
is the least distinctive;
for the Asiatic part of the
The
some,
important
length, subsidiary
shallowly
the
family
cells of
concave,
small
(very)
branched sclereids present in the
unique
distinct.
leaf anatomically
are
Each
for
the
The
Asiatic
features.
particular
adaxially approximately
in the midrib
rarely
Myristicaceae
summarized in
are
characters
stomata
are:
hairs
protruding,
adaxial vascular bundle
sclerenchyma
filiform,
caps and
mesophyll.
only character,
constant
is the small size of the
for the genus and
phloem
in the
groups
midrib.
Knema has
sclerenchyma
For
length
Myristica
and the
papillae)
characters the
unique
as
sheaths
the
unique
'ring'
above the
in the
of
major
characters
bordering
stomatal
Knema.
two
groups:
Regarding
grouping. According
to
with
supported by
the
most
this
characters,
Sinclair
(1958a)
a
primitive
more
or
arms
conspicuous,
less
close
leaf anatomy
and another
and
concerning
Myristica
there appear
the
to
and
of
equal
upright
be
four genera
the hairs separate
2) Horsfieldia
no reasons
are
close
to
as
to
it
provides
the
most
and
no
Knema
motives
advanced.
last.
to
This
indicate
and
for such
a
each other;
series of increasing advancement: first
Gymnacranthera
study;
hairs with cells with two
(with
each genus, those
to
furthermore he arranged them in
Horsfieldia
are:
cells
1) Gymnacranthera
the other
the
complex.
Of the characters, applicable
the genera in
stomatal structure' and
'star-shaped
veins.
Myristica,
view
one
is
not
genus
as
J. Koster & P.
Table V.
Baas:
Lea)anatomy of
Differential leaf anatomical characters
+
present; — = absent; —/+
=
of
in
=
Asiatic
Horsfieldia,
Gymnacranthera,
some
159
Myristicaceae
species absent,
in others
Knema and
Myristica
present
03
t—
<D
I
03
o
2
03
O
03
C3
c
characters
Important
Basal
of hairs
parts
Most basal
More than 6
Number of
basal
arms
than
more
one
cells
surface
per
basal
per
X
-
unit
abaxial side of the midrib
on
-
of hair
part
o
Stomata
characters
Important
Arms
sunken
spongy
in
of hair cells of different
Arms of hair
bordering
adaxially
+
+
complex
flat (to
-
shallowly
+
concave)
+
Phloem groups in midrib small
with small
Major
Thin,
veins with
filiform,
recorded
Characters,
branched
for
one
Large xylem
bundles
cells in the
Garrat
(1933)
characters for
number in
of
and
presented here,
do
be
the
the
can
is leaf
be used
family
more
mentioned
as
recent
stem
& Wilson
a
as
(Wilson,
and node
pollen (Siddiqi
-
+
-
+
-
+
-
-
-
-
-
-
-
-
-
-
-
-
+
-
-
-
-
-
-
-
-
-
recorded
(1980)
(1968)
found
and Wilson
±
-
±
±
some
-
+
-
-
-
wood
deviating
deviating
chromosome
(1978) regarded
the anatomy
a
somewhat different from those of the other genera.
not
support
a
distinct
position
somewhat isolated. A
for
Horsfieldia; only
grouping
of certain
with the present results.
synoptical key
to
the four
genera.
level
literature the
the closest allies of the
have been regarded
androecium
as
-
-
-
anatomically
supported
+
-
genus:
fibres
Ehrendorferet al.
as
cannot
Table V
In
Armstrong
Horsfieldia
Gymnacranthera
On
sclerenchyma
Horsfieldia species; Armstrong
The results,
for that
subepidermal layer
Horsfieldia;
of the flowers of
genera
constant
epidermis
of the midrib
pith
Anastomosing veins, composed
Swollen
-
-
+
and not
genus
in the
-
-
-
sclereids
of minor veins in the
Prominence
+
caps
sheaths
sclerenchyma
rarely
-
-
+
-
sclerenchyma
-/+
-
-
approximately
2
+
-
-
stomatal
Adaxial vascular bundle in midrib absent
Major veins
+
-
dome-shaped
cells above
-
1
only:
stomatal structure
Star-shaped
Midrib
genera
length
equal length
of stomata
cells
Subsidiary
Ring of
cells of
single
-
+
(+)-
+
-/+
"/+
+
(+)-
-
tissue
occurring
+
1
+
distinctly
druses in
+
+
2
Alveolar material
Large
•c
+
(-)
-
hair cell
per
£
genus:
adaxial surface
on
of hairs
parts
in
occurring
a
c
52
the
family
Annonaceae and
Myristieaceae.
most
closely related,
1966; Wilson & Maculans,
(Wilson,
1965; Siddiqi & Wilson,
& Wilson,
1975b).
Based
on
the
Canellaceae have
The Canellaceae in
based
1967),
on
been
particular
the anatomy of the
the anatomy of the young
1975a)
and the
pollen morphology
structure
the
of the
Annonaceae
BLUMEA
160
have also been mentioned as the
(Joshi,
Sinclair
1946).
To
short
a
family,
and
ochocoa,
Scyphocephalium
(Compsoneura
ulei,
platyspermum,
Virola
The anatomy does
to
allied with the
strongly
this
compare
Staudtia
is
family
V.
of the African
Pycnanthus
the
Iryanthera
deviate from that of the
on
indispensable.
specimens
and
gabonensis)
V. loretensis and
calophylla,
based
the leafanatomy of the three
cuneatum,
lehmanni,
Myristicaceae
relationship,
leaves of
Coelocaryon
Dialyanthera
not
1981
advocated
Myristicaceae,
acuminata,
species (Brochoneura
I.
survey of characters of each
general
the survey of the
complete
most
also
(1958a)
characters. In order
macromorphological
families,
VOL. 27, No.
angolensis,
American
ulei,
species
Osteophloeum
have been examined.
pavonis)
Asiatic genera in
the characters
mentioned here. The variation in the presence of the alveolar material and in the
vascular
of the midrib,
system
found in the African
described
recently
shows
a
similar
well
as
found in the
Bernardi and
by
variation
Asiatic
Myristicaceae
in the American genera. The
as
in
as
for
Spichinger (1980)
the
Asiatic
confidence that the leafanatomical characteristic of the
The
comprehensive.
have
genera
their
implications
closest
the
on
relatives
on
genus
other
i.e. the
level;
American genera,
give
below is
possibility
would
continents,
been
of the hairs
similarities
family given
of African and American material, and is
detailed study
some
These
genera.
has also
,
morphology
that
require
a
us
fairly
some
more
the scope of this
beyond
study.
The characteristics of the
Myristicaceae
when hairs have been shed,
present.
Stomata
embeddedin the
collateral
phloem
bundle,
cells.
with
often
caps
or
more
also with free
and
spindle-shaped particles
phloem
Sclereids
the
fragrans,
complement
in
the
to
mesophyll
dinisii,
the
less
shaped
guard
the
mesophyll
the
ground
as
tissue
epidermis. Tanniniferous
filiform, rarely
a
ground
of
cells
large
to
sacs
mannii)
epidermis.
absent. Sclereids
also
the basal
at
small druses
the
sacs
midrib and
often present
of
tissue
species
not
present
in the
in
midrib,
has
been
Cinnamosma
examined
are:
Hairs
to
not
Midrib with
an
to
arc-
in groups. Veins with collateral
as
at
spindle-shaped
Oil cells present in the
cells sometimes
usually
the
(Canella alba,
Petiole with three collateral bundles
small druses and sometimes
characters,
bundles, with
only abaxially present, paracytic
phloem apparently
caps.
the
bundles in the
to
macranthum,
embedded. Leaf dorsiventral.
not
less
branched elements.
number of
Warburgia
tissue of the midrib; secretory
Most of the
ground
targe
or
arc-shaped
the mesophyll and the ground tissue of
present in
and
and sometimes in the
Tanniniferous
the
mesophyll,
as
arms;
mostly
adaxially,
three collateralbundles
mostly
astrosclereids in the
as
phloem
more
an
with collateral
groups. Veins
leaf anatomy of
collateral bundle, the
Crystals
with free
scanty data in the literature. The characteristics
bundles, with sclerenchyma
end.
cells
collateralbundle
straight
recorded. Alveolar material absent. Stomata
bicyclocytic;
Alveolar material
dorsiventral. Midrib with
cinnamodendron corticosum. C.
madagascariensis,
the
visible.
adaxially. Crystals mainly
Oil cells
brachy-
Canellaceae,
C.
Hairs uniseriate, with cells with
layers; always
cells present in the
as
sometimes also in the
Capsicodendron
bundles
epidermis.
midrib; secretory
the veins.
or
two
sheaths. Petiole with
end,
Of
a
Leaf usually
accompanied by xylem
centre, sometimes
sometimes in the
are:
remain
usually only abaxially present, paracytic; guard
subsidiary
in separate groups, often in
sclerenchyma
the
basal parts
present
the basal
particles
mesophyll
in the
in
and
epidermis.
absent.
Myristicaceae (stomata only abaxially,
J. Koster &
bundles;
crystals
have
present)
Asiatic
Leaf anatomy oj
wide
a
druses
as
and
in the
1950). Important characters, present
alveolar material,
bundles in
the
Canellaceae.
guard
the
of the
pith
Therefore,
midrib and
close
a
particles;
Dicotyledons
the
the
between
&
Chalk,
the
peculiar hairs,
subsidiary cells,
tanniniferous
the
relationship
cells
secretory
(Metcalfe
Myristicaceae (as
embedded in
cells
with
sclerenchyma; petiole
spindle-shaped
distribution among the
161
Myristicaceae
leaf dorsiventral; collateral vein bundles with
paracytic;
three
P. Baas:
sacs)
phloem
the
absent
are
these families
in
the
be
hardly
can
supported.
The
characteristics
of
the
Annonaceae
Jovet-Ast
to
Blunden et al.
not
Stomata
collateral
vascular system
with free
several
Petiole with
druses and
mesophyll
sclereids
the
brachy-
mesophyll and/or
to
the
the
centre);
in the
at
with
the
mucilage
or
same
too
lack
are
as
absent.
sacs
astrosclereids
Sclereids
or as
resemblance,
sclereids.
close
Magnoliales
(taken
candidates for
to
it
between
other
in
close
the
to
large
sometimes
brachy-
as
broad
A
the
and
hairs
(the
superficial
a
the
filiform
indications of
of the flowers in
and
comparison
of
versa
only
groups
no
less
more or
close
a
Myristicaceae.
Myristicaceae
sense
with
are
are
The Annonaceae
and vice
show
phloem
there
morphology
the
Myristicaceae
Myristicaceae.
characters
that
say
families).
affinity
present
the
with
Cronquist,
Myristicaceae.
other
does
1968)
Yet
did
particular,
Annonaceae
or
families
not
the
of the
yield
Myristicaceae
not
other
clearly
this order through their leaf anatomical features like oil cells and paracytic
The
embedded
alveolar
material
cells in the
guard
of
the
subsidiary
Winteraceae
(Bongers,
cells of the Lauraceae
the
branched hairs and
sympodially
vascular system of
petiole
and midrib, that
the
take
an
the
additional
Magnoliales.
separate phloem
Myristicaceae
and
1973)
provide
circumstantial evidence of links with individual families of the
through
small
tissue of the midrib and
and cavities present in the
Myristicaceae
the
bundles,
to
the
(1972), studying
(among
stomata.
medullary
suffice
relationship
Canellaceae
belong
in the
Some
instance).
between the Annonaceae and the
Also Nair
a
the
as
Therefore
relationship
see
for
as
in
sheaths.
or
thin, branched, filiform sclereids in the
those between the Canellaceae and the
different,
or
(not
groups
caps
Crystals
ground
cells
bundles,
tissue of the midrib.
ground
important characters, present
very
large
sclerenchyma
the basal end.
mesophyll,
not
an arc-
medullary part (not
a
often in
phloem
The resemblances between the Annonaceaeand the
the
cells
dorsiventral. Midrib with
abaxial and
are,
and
(1969)
Alveolar material
peltate.
or
anatomy,
West
tissue of the midrib; oil cells sometimes in the abaxial
ground
Tanniniferous
or as
leaf
(1950),
consisted of several small
arc
adaxial,
an
Oil cells
epidermis.
and the
epidermis.
with
three vascular strands
mostly
prismatic particles
in
especially
complex,
Veins with collateral bundles,
layers).
Chalk
stellate
isobilateral, mostly
bundles in the
phloem
simple,
sometimes the
bundle,
more
and
usually only abaxially present, paracytic; guard
embedded. Leaf sometimes
shaped
if present,
(1973): Hairs,
recorded.
Metcalfe
(1942),
the
concerning
,
according
It
is
groups of the
isolated
position
within the order.
REMARKS
The
majority
forests
(Sinclair,
of the
1958a),
Myristicaceae
a
ON
are
ECOLOGY
confined
to
the
lowland
very humid habitat. Therefore it is
tropical
surprising
to
rain
find
so
162
BLUMEA
characters in
xeromorphic
many
and
adult
dense
a
(characters, usually
No.
198!
1,
A thick cuticle,
tomentum
sunken stomata, overarched
leaves,
alveolar material
27,
species.
most
tissue of the midrib,
ground
VOL.
of the
by papillae
for the
constant
the
These characters do
press).
lack these
particular
apparently
are
adaptations
different. Some
not
conditions
to
subjected, quite
semi-arid
a
vesselles
for
phase).
now
surface unit
cells
appear
Another
(if
leaf,
same
to
are
at
is
most
characters,
least
the
this
there is
the isobilateral leaf. The
any) for
important
(an
factor)
call for
appears
basal
many
an
of hairs
for
on
the
Myristica.
never
are
of them is.
isobilateral
leaf
(Esau,
1960),
(1974),
have the leaves in
a more
to
serve
as
explanation
an
of the leaves with
different
only
Sinclair
cannot
position
not
parts
high papillae
not constant
Myristicaceae
be
to
for
passing
similar relic
a
The three characters
one
less distichous arrangement. This fact, however,
sun
i.e.
were
presumably primitive,
species has, according
other
relics;
are
Myristicaceae
features.)
some extent.
but
character
whereas
rows,
and
in
species
(cf. Doyle, 1978,
might
(in agreement with
complexes
each other
the
perhaps
the
the alveolar material and the
Horsfieldia sabulosa;
leaves in several
or
on
xeromorphic
for
adult leaves
on
areolae),
replace
to
present together
recorded
the
mesic habitats
as
species
differentiationof angiosperms
markedly xeromorphic
occupying
the stomatal
bordering
They
on
of
habitat,
present
early ecological
The
characters
and
1963, Koster & Baas,
circumstances of the
ancestors
the leaf anatomical
explaining
The dense tomentum
per
the
on
Winteraceae,
hypothesis
the
cells
regarded
may be
species (Horsfieldia
ecological
xeromorphic
which
to
in all
occur
different from those of the
interesting hypothesis
through
not
whereas the
features),
mesophyll
bordering
species)
features, restricting the transpiration (Burstrom & Odhnoff,
in
sclerified
the abaxial surface of the
on
regard
from that of
the
to
the
other
Myristicaceae (De Wilde, personal communication).
In
the
conclusion,
seems
defy
to
features
all
the
to
Myristicaceae provide
generalizations
prevailing
situation in the genus
xeromorphic
interpretation
not
even
equally
a
is
if
in
to
they
to
of
an
(cf.
of a
Baas,
earlier
stretches
It leaves the
of the
example
plant
group which
the
question
for
1975,
a
similar
adaptive significance
limits
unanswered
of
of the
functional
why
at
present
grows in semi-arid habitats. However,
family
it is
visualize the development and retention of so many xeromorphic
were
research
needed
hypothesis
Myristicaceae
speculation.
portion
another
adaptability of various leaf anatomical
conditions
ecological
The
at
on
all
enable
stages
in evolution
totally meaningless
the actual role of the anatomical
householdand gas
water
badly
and
difficult
characters,
the
features
small
Physiological
Ilex).
the
on
exchange
more
in these and other
fruitful discussions
on
structures
tropical
ecological
for survival.
concerned in
rain forest
plants
and functional leaf
anatomy.
SYNOPTICAL KEY TO THE
MALESIAN
GYMNACRANTHERA
a:
Number of basal
parts of hairs
abaxially (average
G.
14-22:
—
55:
the
values of about five
2—11:
28
on
SPECIES AND
(p.p.
=
pro
VARIETIES OF
parte)
areolae and
minor veins
counts):
contracta
p.p.,
G.
eugeniifolia
G.
contracta
p.p., G.
forbesii, G.
G.
bancana
paniculata
/
0.1
2
mm
J.
b:
Koster
& P.
Baas:
Leaf anatomy of Asiatic
Thickness of lamina in /im (This character has
when other leaves than examined here
<
200:
S: 200,
G. forbesii
<400:
G.
forbesii
>400:
c:
G.
forbesii; G.
var.
bancana
G.
panicuiata
<
12:
>12:
paniculata
Abaxial
Abaxial
paniculata
eugeniifolia;
G.
zippeliana
var.
borneensis
var.
layers
of
contracta
p.p., G.
forbesii;
G.
contracta
p.p., G.
eugeniifolia
G.
bancana
layers
tissue sclerified,
spongy
of spongy tissue
at
G. forbesii
eugeniifolia;
G.
least
var.
paniculata
locally:
crassinervis p.p.
sclerified:
not
G. bancana, G. contracta, G. forbesii
forbesii
Major
veins with well
G.
G.
Druse idioblasts
developed
Druse idioblasts
G.
var.
crassinervis p.p., G.
paniculata
sclerenchyma
caps:
forbesii
sclerenchyma:
paniculata
bordering
G.
forbesii,
var.
bancana, G. contracta, G. eugeniifolia, G.
veins with little
Major
f:
reservation;
quite possible)
var.
contracta; G.
G.
G.
e:
some
are
Thickness of adaxial cuticle in /im:
>7,
d:
bancana;
var.
be used with
G.
crassinervis;
var.
bancana
to
studied, overlaps
are
163
Myristicaceae
the abaxial
on
eugeniifolia
bordering
var.
epidermis only:
griffithii
the abaxial and adaxial
on
G. bancana, G. contracta, G.
epidermis:
eugeniifolia
var.
eugeniifolia,
G.
eugeniifolia,
G.
forbesii, G. paniculata
g:
Minute, prismatic crystals present in the abaxial epidermis:
G. bancana
var.
bancana
Minute, prismatic crystals absent:
G.
bancana
G.
forbesii,
SYNOPTICAL
(the
a:
Adult leaves
groups
densely
2
Adult leaves
not
KEY
borneensis,
TO THE
specified
are
5
very
G.
contracta, G.
GROUPS OF KNEMA
on
p.
covered with hairs
p.p.,
(or
var.
paniculata
153) (p.p.
=
SPECIES
pro
parte)
abaxially:
p.p.
sparsely)
covered with hairs:
1, 2 p.p., 3, 4, 5 p.p., 6
b:
Hair type
(see
page
141):
1: 2 p.p.
2: 2
6: 4 p.p.
7:
1
p.p., 2 p.p.,
3: 4 p.p., 6 p.p.
8a:
1
p.p., 2 p.p., 5 p.p.
4:
8b:
1
p.p., 2 p.p., 3
p.p.
6 p.p.
5: 6 p.p.
c:
Number
of basal parts
abaxially (average
<
12:
>
12:
8c:
3 p.p., 4
p.p., 5 p.p.
4 p.p., 6 p.p.
of hairs
on
the
values of about five
counts):
1, 2 p.p., 3 p.p., 4, 5, 6
2 p.p., 3 p.p.
areolae and
minor
veins
/
0.1
2
mm
BLUMEA
164
d:
Thickness of lamina in /im
character has
(This
when other leaves than examined here
:g
120:
>
120,
No.
VOL. 27,
are
1981
1.
to
be used with
studied, overlaps
some
reservation;
quite possible):
are
1, 2 p.p., 3 p.p.
<200:
2 p.p., 3 p.p., 4, 6 p.p.
2
p.p., 5, 6 p.p.
Thickness of adaxial cuticle in /mi:
e:
>5,
10:
<
10:
1
p.p., 2 p.p., 3,
1
p.p., 2 p.p., 4 p.p., 5
4 p.p., 6 p.p.
p.p., 6 p.p.
5 p.p., 6 p.p.
Thickness of abaxial cuticle in /mi:
f:
<5:
1, 2, 3,
4, 5 p.p., 6 p.p.
5 p.p., 6 p.p.
tissue
Spongy
g:
sclerified:
obviously
2 p.p.,
tissue
Spongy
3,
5 p.p., 6
sclerified:
(obviously)
not
1, 2 p.p., 4, 5 p.p.
h:
Leaf
margin
sclerified:
Leaf
margin
not
2 p.p., 5 p.p., 6 p.p.
sclerified:
1, 2 p.p., 3, 4, 5 p.p., 6 p.p.
tissue of midrib
Peripheral ground
i:
2 p.p., 3,
tissue of midrib
Peripheral ground
bundles in
Xylem
sclerified:
weakly sclerified
not:
or
2 p.p.
1,
j:
strongly
6
4, 5,
pith
of midrib
large (see fig. 5):
2 p.p., 6 p.p.
bundles in
Xylem
pith
Regular cork
k:
warts
of midrib
2 p.p., 3,
1,
not
large
or
absent:
4, 5, 6 p.p.
abundantly present:
5
cork
Regular
warts
absent:
1, 2,
3, 4, 6
ACKNOWLEDGEMENTS
We wish to thank
on
Knema
Horsfieldia.
in
the text
Dr. W.
to their
prior
Mr.
M.
figures
J. J.
O. de Wilde
publication. Dr.
Groeneveld,
and
for
J. E.
Mr. B. N.
photoplates. Special
givingvaluable
taxonomic advice and
Armstrong kindly provided
Kieft and Mr. C. L. Marks
thanks are
due to Dr.
are
D. F.
with
us
access to
acknowledged for
Cutler
his results
his thesis
copy of
a
for
testing
on
their share
the
alveolar
material.
REFERENCES
ARMSTRONG,
J. E.
1975.
Thesis. Miami
& T.
K.
The
comparative
University, Oxford,
1978.
WILSON,
Floral
anatomy
and
morphology
of
Horsfieldia (Myristicaceae).
Ohio.
morphology
of
Horsfieldia (Myristicaceae).
Amer.
J.
Bot.
65:
441 -449.
BAAS,
&
—
P.
1970. Anatomical contributions to
from
1975.
1980. Wood
Madagascar
Anatomy
and
22:
Horsfieldia (Myristicaceae).
Cratoxylum
Vegetative anatomy
Oncotheca. Blumea
of
and
taxonomy
affinities
of
IAWA
I. Floral and
from Indo-Malesia
the
311 -407.
plant
Bull.
N.S.
I:
121
vegetative anatomy
(Guttiferae).
Blumea
—
129.
of Eliaea
18: 369 — 391.
Aquifoliaceae, Sphenoslemon,
Phelline
and
,
J. K.OSTER &
1. W. &C. G.
BAILEY,
and
&
L.,
133-
J. Am.
Asiatic
of
anatomy
comparative morphology of
Arb. 25:
1980.
SPICHIGER,
R.
Leaf
165
Myristicaceae
the Winteraceae
V. Foliar
epidermis
342-348.
Las Miristicaceas
del
Jenaro Herrera.
Arboretum
Candollea
35:
182.
BI.UNDEN, G.,
AYE KYI & K.
macrophyllus,
H. G.,
BURSTROM,
comparativeleaf
and G. velutinus.
leaf characters
1978.
WANNTORP,
& C.
1973. The
JEWERS,
malayanus
Epidermal
E.
& H.
BREMER, K.,
G.
1973.
M.
J.
BONGERS,
1944. The
NAST,
sclerenchyma.
BERNARDI,
P. Baas:
anatomy of Goniothamus
Linn.
Soc.
67:
Blumea
in
botany.
Vegetative
21:
G.
381 -411.
Taxon
27:
317
—
Alb.
plants. Stockholm,
of
anatomy
andersonii,
361 —376.
of the Winteraceae.
Phylogenetic systematics
1963.
ODHNOFF,
Bot. J.
329.
Bonniers
Boktryckeri.
A.
CRONQUIST,
1968.
The evolution
of
and classification
Mifflin
flowering plants. Boston, Houghton
Company.
DOYLE,
J. A.
1978. Fossil evidence
P.
systems (Ed.
EHRENDORFER,
F.,
F. KRENDL, E.
K.
ESAU,
1960.
G.
Anatomy
A.
HARTOG-VAN TER
lato.
Acta
W.
JANSEN,
Bot. Neerl.
P.
&
T.,
1946.
affinities
A
1942.
on
J., & P.
METCALFE,
C.
N. C.
(in
R.,
certain
S.
Murty
living
in
1978.
of
Woods
Myristicaceae. Trop.
Epidermal characters
35: 6
—
of the Celastraceae
48.
sensu
388.
—
leaf
Comparative
of
anatomy
Kokoona
and
Lophopetalum
153— 178.
the
development of pollen
J.
les
sur
national
of
Bot.
Indian
Myristica fragrans
Soc. 25:
d'histoire
van
Houtten
and
the
139- 143.
d'lndochine. -
Annonacees
du Museum
1981. Alveolar material
the
Anatomie
naturelle;
-
foliaire.
Repartition
fascicule
serie; XVI,
nouvelle
& L.
of nutmeg
sex
A. N., & W. Y.
Flora Jena
1975b.
Pollen
Leaf
J.
1958b.
Curr. Sc. 46:
malabarica
Myristica
In: Advances in
Press.
Lamk. with
a
discussion
Y.
plant morphology (Ed.
simple morphological technique for distinguishing
156—
157.
sclereids in certain
1975a. Stem
members of Annonaceae
anatomy and nodal
anatomy of the
and
Myristicaceae.
genus
Knema.
Pak. J.
The
whitmorei
The
Bull.
Bull.
genus
—
XX.
-
23:
XXXI.
18:
Sing.
precursores
Sing.
102
—
—
spp.
The
Gard.
genus
Bull.
21:
167—
Sing.
16:
175.
205-472.
Gymnacranthera (Myristicaceae)
in
nov.
The
genus
Knema
(Myristicaceae)
in
Malaysia and
327.
XLII.
Gard. Bull.
The
in
Sing.
Horsfieldia (Myristicaceae)
genus
197-200.
in
Myristica
genus
Malesia
and
outside
1—540.
Horsfieldia (Myristicaceae)
J. Sinclair
7:
17: 96—120.
precursores
Gard.
Pak. J. Bot.
(Myristicaceae). Biologia
Malayan Myristicaceae.
precursores
Sing.
Malesianae
Gard.
(Myristicaceae).
genus Knema
of the
Malesianae
Gard. Bull.
Florae
Knema
of the
revision
Malaysia.
Malesia.
1975.
genus
Florae Malesianae
1974.
of
Myristicaceae.
1977. A
VATSALA,
1966. Foliar
WILSON,
of the
Florae
outside
1968.
of
Clarendon
Dicotyledons. Oxford,
of the
Anatomy
systematics
seedlings.
anatomy
1958a. A
Malaysia.
1961.
Bot. J.
67-76.
1975c.
SINCLAIR,
Suppl.
156: 220-231.
M. R. & T. K.
Bot. 7:
Cuticle.
277.
—
CHIN,
B.
In: The Plant
Myristicaceae.
morphology and embryology
264
al.).
1950.
CHALK,
aspects of the
et
in
press).
B. K., R. RAJENDRA & P.
SIDDIQI,
as
trees
Univ. Press.
& Sons.
London, Wiley
BAAS,
1973.
Recherches
1972. Floral
on
the
RAO,
Tropical
Cambridge
1968. Chromosome numbers and evolution
SAUER,
den, &
family Myristicaceae.
BAAS,
Linn. Soc.
NAYAR,
3-30.
353.
New York,
P.
and forests. In:
trees
125-308.
KOSTER,
NAIR,
—
of the wood
21:
geographique.Memoires
3:
plants.
355
BAAS,
note
of the
S.
JOVET-AST,
27:
Blumea
& W.
337
tropical
Zimmermann)
anatomy
R. M.
THOLEN,
(Celastraceae).
JOSHI, A. C.
of seed
17:
of
origin
HABELER,
Systematic
1933.
the
& M. H.
Taxon
primitive angiosperms.
GARRAT,
on
B. Tomlinson
in
and
27:
and
outside
Malesia
I:
H.
sabulosa
and
H.
Sing.
28:
133-141.
outside
Malesia
II.
Gard.
Bull.
1-181.
SOLEREDER,
H.
1899 &
1908.
Systematische
Anatomie der
Dicotyledonen & Erganzungsband. Stuttgart,
Enke.
STACE,
C. A.
STAVEREN,
1965. Cuticular
M. G. C.
Neerl.
22:
VAN &
studies
P. BAAS,
as an
1973.
aid to
plant
VLIET,
Bull. Brit. Mus.
of the Malesian
(Bot.)
Vol.
Icacinaceae.
4
no.
1.
Acta Bot.
329-359.
VINK, W. 1977. The Winteraceae of the old world II.
23:
taxonomy.
Epidermal characters
Zygogvnum
—
Morphology and taxonomy.
Blumea
219-250.
G. J.
C.
173-195.
M.
VAN
&
P.
BAAS,
1975.
Comparative
anatomy
of
Crypteroniaceae
s.l.
Blumea
22:
BLUMEA
166
1897.
Monographie
WEST, W. C.
WILDE,
1969.
WILSON,
T. K.
1966. The
Amer.
WORSDELL,
1
Photos
170.
hair
type 5,
hair
type 7,
C.
52: 369
—
1967. The
Internal
4. K.
Photos 7—12.
types
8b,
x
350.
material
as a
hairs,
and
x
Scanning
140.
—
phloem
in
3,
material
note tall
on
warty
abaxial
rods
(heavily
restricted to
stomatal
a
alveolar
corresponding
surface with
complex,
warty layer.
complex (s),
(‘spongy’)
electron
infested
x
material,
with
convex
with
Note
800.
anticlinal
CI. 96:
Blumea
329 —
25:
of the
Anatomy
—
82.
344.
321 -478.
young stem
morphology and
conclusions.
Ann. Bot.
Lond.
types in Knema.
note tall bodies
cuticle,
alveolar
x
type 8c,
of hair
820. —
material,
tall
90.
x
—
hairs,
the
x
1.
—
K.
Myristica fragrans
x
9.
material
laurina,
x
—
80.
—
oblongata,
hair
type 1,
3. K.
lunduensis,
5. K.
globularia,
hair
type 8a,
note
170.
material.
—
(±
11. Knema
surface
as in
—
7. Knema
retusa,
cinnamomea. thin-walled
Myristica
150.
527.
85.
x
cells,
K.
entire abaxial
intermedia,alveolar
526
lying hairs,
alveolar
types and
of hair
plates covering
22:
—
alveolar
oblongata,
(exept margin,midrib
photo 11) restricted
to cells
x 410.
epidermis covered
above stomatal
bot.
I. Flowers of
Myristicaceae.
of hair
with
micrographs
12. Knema
—
II.
IV. Floral
stalk cells of free
abaxial
mandarahan, hair
of erect, lobed
13 — 18. Scanning
Photos
opening
electron
of the
Myristica.
plumulosa, hair type 6,
complex,
Torrey
1 —680.
54: 214-220.
micrographs
hair type
Knema
1730.
x
stomatal
Bull.
13: 78
68:
(Myristicaceae).
the Canellaceae.
the Canellaceae.
Myristica villosa, thick-walled,
reticulum
major veins),
overlying
papillose
10.
—
Deutsch. Bot. Ges.
Leop.-Carol.
Ranales.
genus Knema
morphology
J. Bot.
electron
also alveolar
note
woody
d.
Ber.
Acta Acad.
378.
dome-shaped unspecialized epidermal cells
hair
the
of the
1981
1.
343.
Amer.
percoriacea,
—
—
336
1908.
170.
x
in
comparative morphology of
— 6. Scanning
2. K.
—
Bot.
MACULANS,
W.
of oil cells
Nova
comparative morphology of
M. malabarica.
and
x
The
Amer. J.
J. Bot. 53:
& L. M.
Myristicaceen.
1979. New account
DE,
1965.
and node.
der
Ontogeny
W. J. J. O.
No.
dieHaarbildungderMyristicaceen.
O. 1895. Ueber
WARBURG,
VOL. 27.
x
fungal hyphae),
16.
270.
walls;
sculpturings
material of erect
features.
plates.
part
of hair
x
1660.
(arrow)
15.
—
and
—
Note
alveolar
820. — 14. Myristica globosa,
x
basal
—
of miscellaneous
micrographs
with alveolar
of cells
Myristica maingayi,
of cells
material
Myristica
ring
13.
ring
villosa,
overlying
as a
alveolar
and
Horsfteldia iryaghedhi, abaxial, papillate epidermis
—
17.
Myristica maingayi,
note basal
of cuticle
part
which
of hair
are
adaxial
in centre;
much smaller
x
surface with
400.
than
—
cells,
of
material
surrounding
with
grooves
18. Knema
individual
surrounding
reticulum
rufa,
x
porous
in cuticle
adaxial
1470.
J. Koster
& P.
Baas:
Leaf anatomy of
Asiatic
Myristicaceae
167
168
BLUMEA
VOL.
27,
No.
1.
1981
J.
Koster & P. Baas:
Leaf
anatomy
of Asiatic Myristicaceae
169
BLUMEA
170
Photos
with tall
19
—
basal
thick cuticle
22.
Light micrographs
cell,
on
x
tips
600.
of
x
20.
papillae;
prominent striations, and
cuticle with
—
VOL.
of various
27,
No.
I,
features.
epidermal
Myrislica maingayi, transverse
x
240.
basal part
undulating anticlinal
—
at
five cells,
high
focus
x
19.
—
Myristica guatteriifolia, hair
papillate epidermis.
Note
abaxial cuticular maceration
with
section
21. Horsfieldia crassifolia,
ofhair with
flanges
1981
with
380. — 22. Gymnacranihera forbesii,: adaxial
and thin
areas
of cuticle
in
loops
of undulations,
600.
Photos
23
of abaxial
stomatal
Knema
—
27.
complex
laurina,
complex
Light micrographs
of stomatal
complexes. —
23. Knema laurina.
epidermis showing star-shaped opening (arrows) overlying
and
in
focus;
transverse
leaving
a
inset:
section
star-shaped
structure
as
seen
showing ± horizontally
star-shaped opening
as
shown
in
stomatal
in cuticular
directed
photo 23,
paradermal section
complex;
maceration,
papillae overarching
x
600.
—
25.
x
at
bottom
600.
—
24.
the stomatal
Myristica.
—
a.
M.
J. Koster &
P.
Baas:
chrysophylla, paradermal section showing rings
—
x
b. M
380.
cinnamomea, ibid,
—
26.
Myrislica chrysophylla.
cells, forming ring
cells,
x
600.
in cuticular
—
as shown
27.
in
photo
Gymnacranthera
171
Leaf anatomy of Asiatic Myristicaceae
of cells
overlying
maceration. Note
transverse
25. Note
section of stomatal
also small cells
contracta,
and
bordering
also small basal
paradermal
complex
(arrows)
section
parts
overarched
between
with
the stomatal
of hairs
both
by ± papillate
subsidiary
paracytic
complex.
(arrows),
cells and
stomata,
x
ring
600.
172
BLUMEA
Photos
abaxial
absent
28
—
32.
cuticle
near
Light micrographs
near
leaf
leaf
margin.
margin (right),
x
cells
—
29.
with
flanges
x
380.
in
—
contracta,
centre,
31.
maceration
corresponding
Ibid.,
transverse
Horsfieldia sylvestris,
adaxial
with
section
location
hypodermis
I,
features.
198!
—
thin cuticle
very thin cuticle
with
of adaxial
with
very
No.
28.
Knema
pallens,
(some arrowed);
maceration
alveolar
Ibid., transverse section, showing globular cells
(or mesophyll cells intruding into epidermis?)
Gymnacranthera
27,
of miscellaneous
Note
380.
VOL.
shown in
cuticle, showing region
of
large crystal
subepidermal
with
as
locally
druse
swollen
idioblast
with
in
idioblast,
cells
relatively
in
x
epidermis
240.
—
30.
thin anticlinal
subepidermal mesophyll,
cf.
(arrow),
photo 28;
of
material
photo 30;
x
240.
x
380.
—
32.
J.
Photos
vertically
33
—
38.
Roster & P.
Light micrographs
transcurrent
bundle
phloem, entirely composed
(arrows),
palisade
x
240.
very
sclerenchyma
brachy-
to
—
35.
Knema
cap of vein
astrosclereid,
in
fibres;
cross
x
pallens,
380.
x
with
a.
380.
section
38.
240.
—
34.
tanniniferous
36.
Knema
epidermis,
x
features.
Knema
note oil cell in
—
adaxial
—
x
173
Leaf anatomy of Asiatic Myristicaceae
of miscellaneous
sheath,
of
thick-walled,
sclereids, mostly
Baas:
x
33.
tubule
running
percoriacea,
240.
150.
—
37.
Knema
minor
mesophyll (arrow);
Gymnacranthera
(some arrowed),
—
laurina,
veins
devoid
into
filiform
vein
with
xylem
or
epidermis (arrow);
sclereid
Gymnacranthera forbesii,
contracta,
of
b. note tanniniferous tubules
from vein
thick
malayana,
mesophyll
with
connecting
midrib
numerous
with
filiform