plants was the origin of plants. These organisms most algae. During

BLUMEA
37
(1992)239-261
Structure and ontogeny of stomata in ferns
U.
&
B.
De
741
Paleobotany Laboratory, Kalyani University, Kalyani
and
Pteridology
Sen
235,
Bengal,
West
India
Summary
On the basis of
into
ontogeny
twenty-four types
and
arrangement of
of which
seven
the
hemiparacytic, pluhemiparacytic, pluparacytic
ships
different stomatal
among
of
position
-
types
Psilotum, Tmesipteris
It is concluded
ties
one is
is, however,
form of the
accepted
and
codiacytic
Ophioglossum
and
are
-
The
new.
group
of
classified
are
polocytic
descendant of the
ontogenetic interrelation-
determining
the taxonomic
discussed.
are
genera
of stomata are not distinct
mesoperigenous types
other. The
as the direct
stomata in ferns
and the role of stomata in
traced
and the
that the mesogenous
the derived
not
are
cells the
surrounding
plupolocytic, pseudocopolocytic, sepcopolocytic, pseudo-
-
stoma which
is
mesoperigenous
in
enti-
origin
anomocytic (perigenous) type.
Introduction
One of the
major
the first land
in the
events
plants.
These
During their transmigration
adaptation
a new
tion of
A
in
two
are
a
waxy
stoma
cases
a
a
pore
single
1958;
cell
in
1962;
(Lele
& Van Cotthem,
man,
1973;
and
ontogeny
of
by
1961; Sen,
only, they
may
there have been
Sen
of
stomata
a
Lersten,
& Bhunia,
large
green
acquired
many
were
the forma-
cells and
guard
two
in the
occur
numerous
1881; Kondo & Toda,
&
of
algae.
1983). Although
1967;
1970a, 1970b, 1973; Probst,
1983;
these
origin
stomata.
surrounded
& Walton,
the
was
land they
on
Among
Viane & Van Cotthem, 1977; Bir
1981; Mehra & Soni,
structure
life
plants
from ancient
arose
to
tissue, and of
Mickel
1965;
Thurston, 1969; Van Cotthem,
habitat
most cases
1870; Prantl,
Pant,
probably
environment.
new
plants. Though
(Rauter,
stomata
Kondo,
aquatic
associated with the leaves
any of the aerial parts of
of fern
an
most
cuticle, of conducting
by
commonly
from
for survival in the
consists of
of the evolution of
history
organisms
et
epidermis
of
detailed studies
1956, 1959; Maroti,
Pant & Khare,
1969;
1971; Fryns-Claessens
Sen &
al., 1980;
1984; Sen,
number of fern
only
stomata
1986),
species
even
Hennipthen the
still remain
un-
explored. Moreover, the ontogenetic interrelationships among many major types of
of the
stomata
reports
filicopsids
about the
grettable.
of
not
been established
development
of
beyond
stomata
doubt. The
in many fern
contradictory
species
are
re-
A survey of relevant literature reveals that stomatal types have often been
identified
ignoring
have
pathways
on
the basis of their apparent structural
the fact that
development
may
stomata
following
occasionally
lution. These confusions,
as
show
also
a
different
apparently
comparative
properties
in
pathways during
similar
structure
dearth of data
mature
the
due
on
condition,
course
to
of their
parallel
stomatal
evo-
structure
240
BLUMEA
and ontogeny
largely prompted
better
this
of the
lead
to
ally
their structure,
and
ontogeny,
a
appreciation
VOL.
Pickled and dried
survey of
of the
mature
stomata
of
of this
fascinating
AND
in the
spores
mainly
the
at
Kalyani provided
borne
by
and foliar epidermal
these
cells.
sporophytes
Dried frond
were
segments
boiled in
acetic
acid, 2.5; propionic acid, 2.5;
equal
volumes of 20%
prepared
Spores
washing
finally
of
species
some
these
the
were
stained in
to
Fronds
in
a
in
fixing
These
and acetic acid
and treated with
water
prior
at
at
3:1
for
(formalin, 5;
58°C for 14 hours and
of dried frond
segments
were
solution of 2.5% NaOH. These
thorough
absolute ethanol: acetic
acid,
stages
of
development
the methods of Stebbins & Khush
(1961).
acetic acid, heated for 1 minute at
These
boiling point
were
in
a
treated according
were
fixed in
mixture. Peels of
fixed material
The
(1981),
a
was
terminology
and Sen
cations. Seven
time,
and
a
POM
was
solution of acetocarmine diluted
dehydrated
used
by
to
new terms
1958),
but the time in
and
half hour. The
to one
(1970a,
1970b, 1973),
the stomatal types is
have been coined for
new
a
Sen &
Hennipman
here after modifi-
adopted
first
types here described for the
Illustrations
were
made with the aid
(India) drawing apparatus.
Permanent slides of stomata, and herbarium of
University.
taxa
prepared.
wax
Occasion-
and mounted in clove oil.
Van Cotthem
designate
also
hematoxylin (Sass,
type has been redesignated.
reference have been
ani
were
reduced from four hours
was
then
(1986)
known
stained with acetocarmine
stained with iron
and mordant
stained material
of
epidermis
to
absolute alcohol;
3:1
1/2 strength with 50% acetic acid, and finally mounted whole in paraffin-bees
hematoxylin
re-
and
1% acetocarmine.
Pieces of fresh lamina at different
ally,
on-
Appendix.
then macerated in
1% NaCl solution. After
treated for 30 minutes in
ob-
various stages
FAPA
were
general
were
provided material, especially
Transparencies
pieces overnight
a
raised from these
Gametophytes
alcohol, 90).
hydrogen peroxide
by soaking
then rinsed in
water
50%
stained in Sudan IV in 70% alcohol.
were
also
Palaeobotany
of the material for
studies. The provenance of material investigated is indicated in the
togenetic
also
plants, especi-
and
Pteridology
most
laboratory ultimately produced sporophytes.
development
group of
METHODS
tained from the Botanic Gardens of Kew and Leiden.
of
that this will
hope
interrelationships.
housed
specimens
University
1992
1,
It is done in the
investigation.
stomata
MATERIALS
Laboratory
No.
37,
have been
For
deposited
recording
at
the
the data
pertaining
raised from spores for
specimens
and
to
the scheme of
arranged following
only for practical
Pteridology
Palaeobotany Laboratory, Kalystomata
of different
Crabbe, Jenny
species
& Mickel
the
(1975)
purposes.
OBSERVATIONS
General
properties
Stomata normally
thick
laminas, they
vestigation
are
of
stomata
occur
are
on
found
all parts of the lamina except
along
hypostomatic, only
a
the veins
few
as
over
the veins. But in
well. Most of the fronds under in-
amphistomatic.
In
amphistomatic
taxa
the
U. Sen
stomata
however,
are,
level of their
others
&
Structure
B. De:
more numerous
position
Guard cells differ from other
their anatomical
in
stomata
241
ferns
the abaxial surface. Stomata vary in the
Some
Guard cell
guard
layer
flush with the
are
fronds
a
cells have
small
practically
in the
all
epidermal cells,
morphology
cells of
epidermal
cells is
guard
but also in
feature.
significant
a
cells is
guard
fronds,
some
than those in the
numerous
mesophyll
more
prom-
cells.
unevenly
ledges
thickened walls and
cutinous
a
the adaxial surface of the
on
of lignin is also deposited
amount
in their
only
not
rich in starch. The nucleus of the
epidermal
often forms
occur
smaller and less
are
exceptionally
inent than that of other
tinous
in
chloroplasts
chloroplasts
are
cells
epidermal
properties. Chloroplasts
but the presence of
The
on
epidermis.
in the
of
ontogeny
sunken below the surface.
are
cells and
and
The
covering.
cu-
In very old
stomata.
the walls close
to
the stomatal
of the lamina are formed
by
the
on
poles.
of
Ontogeny
stomata
The abaxial and adaxial
initials of the
epidermal layers
meristems of the
marginal
anticlinal divisions and redivisions and
go
cells
initially
recognisably
are
uncommitted cells.
moids of
stomata
trichome
skipping
the
course
Stomatal meristemoids
and
cytoplasm
ser
a
are
of its
are
The
occurrence
adjacent
stomata
developing
organ
nucleus. In many
taxa
to
1.
the
subsidiary
Anomocytic
cell
or
two
Meriste-
specific
polygonal,
and have den-
contiguous protoderm
cells
stomata.
nullifies the view that stomatal meristemoids inhibit
also into stomatal initials (see Korn,
different types of
stomata are
1972).
recognised
on
the
surrounding cells, especi-
type:
guard
cells.
maturity surrounding
directly
acts
None of the
of the stomatal meristemoid
At
stoma or a
a
cells derived from the stomatal meristemoids.
The stomatal meristemoid
vides into
form
appendage.
into stomatal meristemoids and form twin
basis of their ontogeny and the mode of orientation of the
ally
or
distinct entities and each kind follows its
studied, twenty-four
taxa
an
transformed into meriste-
directly
can
uninucleate, isodiametric
conspicuous
of such
cells from
In the
cell
are
development
occasionally metamorphosed
are
protoderm
them. Some of these cells
produced
the meristemoid stage of such
moids of each of these kinds
during
protoderm
cells. The
of differentiationthey become
while others
epidermal cells,
and trichomes. No
protoderm
act as
processes
different from the cells which have
become the precursors of
pathway
By
marginal
frond axis. These initials under-
developing
(Fig.
cells
1:
are
as
the
mother cell
guard-cell
surrounding
cells is,
(gcmc)
therefore,
a
and di-
derivative
1-6).
indistinguishable
from other
epidermal
cells in
size and aspect.
2.
Polocytic
type:
The stomatal meristemoid divides
gcmc.
The
subsidiary
from the distal
end;
the
independently of the
subsidiary
cell becomes
by
a
curved wall
U-shaped
cell(s) abutting
on
guard
cells
form
are
1:
7-12).
linked
a
subsidiary
parts
the distal parts of the
stomatal meristemoid (Fig.
cell and that of the
to
and encircles
of the
guard
cell and
guard
cells
cells
originate
The anticlinal wall of the
together
towards the distal end.
BLUMEA
242
Fig.
1.
types
Stages of development of
of stomata.
—
podioides, polocytic
polocytic
stomata;
240;
12, 17,
26 &
anomo-,
x
1,
1992
polo-, copolo-, seppolo-, pseudopolo-,
13—17:
Pteris
Ibid.
, copolocytic
255.
stomata;
biaurita, pseudopolocytic
— 1—5, 7—11, 13—16 &
31
No.
Botrychium daucifolium, anomocytic
stomata;
22—26:
staurocytic stomata.
x
1—6:
VOL. 37,
22—25
x
275;
stomata;
and
18—21: Hypolepis
stomata;
18—20
staurocytic
7—12: Diplazium polypunctata,
27—31: Davallodes
x
1020; 27—30
x
sep-
hirsutum,
610;
6 &
21
U. Sen
3.
Copolocytic
&
B. De:
Structure
subsidiary
cells and the gcmc. It is
subsidiary
cell
or
ontogeny of
in
stomata
243
ferns
type:
The stomatal meristemoid divides
The cell
and
encircling
cells
abutting
most
by
of the
parts
with
stoma
from the
stoma
the distal
on
curved walls
superimposed
two
polocytic
a
of the
proximal
guard
cells
form
to
two
additional U-shaped
an
end
1:
(Fig.
are not
13-17).
the derivatives
of the meristemoid.
4.
Seppolocytic
As in the
type:
the stomatal meristemoid divides
polocytic type,
cell and gcmc. But instead of
one or more
parts of the
from
undergoing enlargement,
anticlinal walls. The
in such
stoma
epidermal
a
(Fig.
cells
the distal parts of the
formed
newly
that
way
they
the
stoma are not
produce
to
subsidiary
become almost
ontogenetic
a
subsidiary
cell divides
cells surround the
daughter
or
cells
by
proximal
in aspect
indistinguishable
In this type, also, the cell
18-21).
1:
the
abutting
on
derivatives of the stomatal meri-
stemoid.
5.
Pseudopolocytic
As
in the
type:
cells and gcmc. The
U-shaped subsidiary
is that
divisions
ly
after its
soon
6.
differentiation,
anticlinal walls. The
by
resemble the
stoma
the stomatal meristemoid forms
copolocytic type,
neighbouring
1:
(Fig.
the first
or
formed after
one
expansion
type
or
more
near-
very
copolocytic
type:
cells and gcmc. It is
three subsidiary
cell
undergoes
cells and the stomatal complex simulates the
The stomatal meristemoid divides
The cell
so
superimposed
copolocytic
22-26).
Plupolocytic
subsidiary
cell
subsidiary
cells
daughter
two
deviation from the
only
encircling
cells
of the
most
abutting
a
on
by
three
superimposed
copolocytic
stoma
with
stoma
from the
the distal parts of the
curved walls
an
proximal
guard
additional
end
cells
2:
(Fig.
are not
form
to
U-shaped
1-6).
derivatives of
the meristemoid.
7.
Pseudocopolytic
type:
The sequence of divisions in
stoma
up
to
is similar to that
the
stage
of
forming
stomatal meristemoid
a
in the meristemoids
occurring
three superimposed
imal parts of the gcmc. The distal
clinal septa
other
cytic
8.
to
produce
epidermal
cells
two
or
2:
(Fig.
subsidiary
more
7-12),
U-shaped subsidiary
cell there divides
daughter
and the
forming pseudocopolocytic
forming plupolocytic
cells which
stoma
so
at
by
cells
one
at
stomata
the
prox-
or more
anti-
simulate the
maturity
formed resembles
a
copolo-
stoma.
Sepcopolocytic
type:
The stomatal meristemoid is
ary cells and gcmc. The gcmc
sions form
ary
cells
the
newly
two
guard
undergo
formed
anomocytic
stoma
one
cells.
or
at
by
more
2:
wall
Following
the
13-18).
at
by
two
right angles
curved walls into
to
the
by
developing
anticlinal walls. After the
mature
stoma
very
two
previous plane
such divisions, both the
divisions
daughter cells,
(Fig.
first divided
a
subsidiof divisubsidi-
expansion
superficially simulates
of
an
BLUMEA
244
2.
Fig.
Stages
1—6: Pteris
tomium
x
of
VOL.
37,
No.
development of plupolo-, pseudocopolo-,
stomata; 7—12:
parviloba, plupolocytic
1992
1,
and
sepcopolocytic types
Ibid., pseudocopolocytic
falcatum, sepcopolocytic stomata. — 1—5, 7—11
&
13—16
x
of stomata.
13—18:
stomata;
12
275; 6,
&
17
x
—
Cyr-
120;
18
350.
9.
Hemiparacytic
type:
The stomatal meristemoid divides
tandy
with the
metaphase spindle
in the gcmc
and the gcmc is divided
during
ary cell
one or more
(Fig.
3:
a
changes
its
a
cell and gcmc. Concomi-
subsidiary
there is
polarity
septum which is
the gcmc
two
changes
more or
less
ary cell. The
cells
ordinary epidermal
a
from
more or
division of the gcmc. The
transverse
less parallel
to
cells and the
proximal
to
longitudinal,
the wall formed
stoma
side
by
is thus
sur-
the subsidi-
type:
hemiparacytic
ristemoid forms
posed
a
form
9-13).
Cohemiparacytic
This is
lies
by
to
subsidiary cell,
the first division of the meristemoid. The distal side of the
rounded by
10.
of the
enlargement
stoma
with
an
additional subsidiary
superimposed subsidiary
its
polarity
parallel
to
surrounding
and therefore the wall
the wall
cells
extending along
are
one
by
of
separating
which the gcmc is
the stomatal meristemoid (Fig. 3: 14-18).
guard
cells
the
separated
heterogeneous origin
side of the
cell. The stomatal
cells and gcmc. The nuclear
are
since
two
me-
spindle
guard
in
cells
from the subsidi-
only
two
ontogenetically
superimrelated
to
U. Sen
3.
Fig.
and
Stages
of
stomata;
aurita, hemiparacytic
dohemiparacytic
x
30
255.
of
Structure
para-,
and
copara-,
7 &
8:
ontogeny of
Ibid.
Cibotium
Pteris
stomata;
stomata;
stomata;
&
25—29
x
275;
19
&
20
1—3,
5 &
6:
9—13: Pseudophegopteris
19—24:
P.
biaurita, pluhemiparacytic stomata. —
10—12, 14, 16,17, 21—23
245
ferns
glaucum, paracytic
, pluparacytic
14—18: Ibid., cohemiparacytic
25—30:
in
stomata
plupara-, hemipara-, cohemipara-, pseudohemipara-,
stomata. — 1—4:
1—3, 5,
stomata;
stomata;
1020; 4, 6, 8,
x
B. De:
development of
pluhemiparacytic types
Ibid., coparacytic
9
&
x
hirtirachis,
1—3,
345; 13,
18
5 & 7
&
24
pseu-
x
x
360;
240;
BLUMEA
246
Fig.
4.
types
No.
1, 1992
stomata. — 1—5: Nephrolepisfalcata, diacytic; 6—10: Ibid., codiacytic
evecta,
ceolata, desmocytic
Ibid., copericytic
240.
37,
Stages of development of dia-, codia-, cyclo-, cocylo-, desmo-, codesmo-, periof
Angiopteris
x
VOL.
cyclocytic
stomata;
stomata; 25:
stomata.
—
17—20: Ibid., cocyclocytic
Ibid., pericytic
1—4, 6—9, 11—23
stomata;
&
stomata;
and
x
275; 5,
10
&
11—16:
21—24: Pyrrosia
26—29: Ibid., codesmocytic
26—28
copericytic
stomata;
24
x
lan-
stomata;
255; 25, 29 &
30:
30
U. Sen
11.
B. De:
Pseudohemiparacytic
This is
more
3:
(Fig.
with its
stoma
of
in
stomata
subsidiary
outer
their distinctive
Despite
expansion only
247
ferns
superficially
very
cell divided
the
nature
by
one
or
cells of the
daughter
resemble the other
epi-
type:
with
stoma
cohemiparacytic
a
ontogeny
19-24).
Pluhemiparacytic
This is
and
type:
cell after
subsidiary
dermal cell
Structure
cohemiparacytic
a
anticlinal radial walls.
outer
12.
&
an
additional subsidiary
cell. The stomatal
meristemoid forms three superimposed subsidiary cells and gcmc. The gcmc divides
by
an
anticlinal wall which is
less
more or
parallel
13.
Paracytic
type:
Meristemoids divide
The semilunar cell
tioning directly
central cell
by
as
by
curved wall into
a
enlarges
become
to
parallel
to
the
one
gcmc, the lenticular cell
a
long
axis of the
now
stoma.
Coparacytic
This is
a
abutting
guard
ent
from that of the
gcmc
15.
cells. The
at
the central
Pluparacytic
This is
a
cuts
16.
sides,
but
additional
an
development
more
guard
stoma
with
cells. Before
a
subsidi-
by
are
a
wall
there-
1-4).
cell
subsidiary
of this
type
of
on one or
often
on
both
sides,
is
stoma
cuts
before
both sides of
differ-
slightly
off
two
segments
differentiating
the
6).
segments
an
additional subsidiary
differentiating
either
on
cell
both lateral
at one or
the gcmc, the stomatal meristemoid
both the sides
or at
least
on one
(Fig.
3: 1-3,
8).
Cyclocytic
type:
the differentiationof the gcmc
two
a
U-shaped subsidiary
or more
the meristemoid orient themselves in such
subsidiary
cell which in its
anticlinal walls
(Fig.
Cocyclocytic
During
the
4:
subsidiary
undergo
division
which is
may also
epidermal
way
that
undergo
a
one
cell and
gcmc. Soon after
flanking
the distal end of
circle is formed
or more
along
divisions
by
with the
vertical
type:
of
cells and the
encircling
turn
a
cells
11-16).
early stages
velopmental pathway
of
cells
cells
a
type:
The stomatal meristemoid forms
17.
the
The stomatal meristemoid
3: 1-3, 5 &
region (Fig.
off three lateral
5, 7 &
of
paracytic type.
coparacytic
flanks of the
with
stoma
pathway
of the
at one
guard
guard
two
on
into
develops
type:
paracytic
the
least
lenticular one.
lateral semilunar segment and
a
gcmc forms
acts as
Both cells
a
cells. Instead of func-
subsidiary
off
cuts
fore the derivatives of the meristemoid (Fig. 3:
14.
semilunar cell and
of the
curved wall. The second semilunar segment
a
which
ary cell and the central cell
at
the subsidi-
separating
the walls
to
cell from gcmc (Fig. 3: 25-30).
ary
by
of
development
copolocytic
gcmc,
either both
one or more
at
least
at
Soon after the formation of the
subsidiary cells,
or
radial anticlinal walls, thus
cells. The distal cells and the
two-layered
the stomatal meristemoids follow the de-
stomata.
the
subsidiary
only
one
of
increasing
them,
the number
cells orient themselves in
proximal part (Fig.
4:
17-20).
two
may
a
ring
BLUMEA
248
18.
Desmocytic
VOL.
37,
No.
type:
The stomatal meristemoid follows the pathway of
A lenticular space
separation
of the
at one
by
subsidiary
a
cells. The lenticular space
guard
cells from the
guard
cell. The
19.
4:
the
Codesmocytic
one
with
stoma
same
an
of
pathway
lenticular space separates the
of the cell walls
Pericytic
This is
additional subsidiary
21.
a
ma,
and the
lost due
4:
(Fig.
cell.
stoma.
short radial
Later
a
the anticlinal
periclinal
seg-
cell
4:
(Fig.
of cell wall
segment/segments
the
Following
without the radial
stoma
subsidiary
to
cell.
developmental path-
connecting
wall segments be-
25).
connecting
the
Following
difference in the
guard
of
a
cells and the inner
rate
growth
link between the
development
of the
guard
copolocytic
subsidiary
sto-
cell is
cells and the subsidi-
guard
30).
During
maturationthe
the distal side of the
cells
guard
not a
at
stoma
their widest
by
subsidiary
a
curved wall
cell
on
orient in such
points
almost
a
the
similar to that
type the first
an
producing
two
a
diacytic
stoma.
acts as
gcmc
(Fig.
common
walls
The distal cell
abutting
meet
the
incidently
is
1-5).
The
only
forming
this type of
difference is that in the
4:
6-10).
This type is,
therefore,
stomata
is
codiacytic
subsidiary
a
diacytic
cell.
type:
The stomatal meristemoid divides into
sidiary
cell and
subsidiary
derivatives of the stomatal meristemoid form U-shaped
additional subsidiary
Staurocytic
a
side and the cell
type:
cells and the third cell
type with
form
way that their
right angles.
at
derivative of the stomatal meristemoid (Fig. 4:
Codiacytic
to
proximal
The sequence of cell divisions in the meristemoids
24.
cell. The stomatal meri-
copolocytic
Diacytic type:
gcmc.
23.
two or one
loses the radial
subsidiary
The stomatal meristemoid divides
on
a
type:
codesmocytic
a
by
subsidiary
finally
the radial anticlinal link between the
gradually
22.
and the
stoma, it
guard cell(s)
and the inner
ary cell
both remain
26-29).
minus the radial link
stoma
guard cell(s)
Copericytic
This is
cell(s)
encircled
type:
desmocytic
the
4:
in
as
cells from the inner subsidiary cell;
guard
(Fig.
desmocytic
a
between the
tween
or
and
points
at two
completely
cells
guard
the
about the
short anticlinal segments of cell wall
development
link between them is then maintained only
way of
of the
one
two
or
bringing
links
cells become
stoma.
connecting
type:
desmocytic
a
stemoid follows the
20.
polocytic
21-24).
This is
ments)
guard
cell and
subsidiary
connected in the distal region by
(Fig.
of a
expands,
subsidiary cell, leaving
point. Therefore, at maturity
later
development
appears in the intercellular material of the wall
soon
cell and the
subsidiary
1992
1,
cell initial
soon
divides
by
a
a
subsidiary
cell initial and
vertical anticlinal wall
to
form
gcmc. The subtwo
subsidiary
U. Sen
cells. The
&
cells in the
subsidiary
two
four walls of the
in different taxa
occurring
stomata in
and the
a
cells in
adjacent
two
cells. The
guard
cross
1:
(Fig.
27-31).
in Table 1.
given
are
of
shape
249
ferns
cells of the
surrounding
cells radiate into the
surrounding
The stomatal types
ontogeny of
proximal position
constitute the
position together
the distal
and
Structure
B. De:
DISCUSSION
the
Among
the
anomocytic
stomatal meristemoid
cell
different types of
twenty-four
vestigation,
soon
Table
study.
It is
The
signifcant
of
giatae (except
that in
course
the
sporangiatae, anomocytic
even
if
only
In the
from
ments
by
none
form
of the
(1983),
belonging
taxa
to
Ophio-
Marattiaceae,
Kondo & Toda
be confirmed in this
not
the so-called
Eusporanand
Lepto-
in association with other stomatal
types,
stomata occur
as
many
as
four
stomatal types the stomatal meristemoids
cells either
subsidiary
before
primary segments
directly
or
after
acting
division
undergoing
cut
off
gcmc. These seg-
as
redivision
or
radial anticlinal walls.
Among
and
these
twenty-three types,
in all fern families
occur
(1975). Seppolocytic
of Sen &
Hennipman,
Pseudohemiparacytic,
cytic, codiacytic
Not
only
glossaceae),
and
is the
the
polocytic
stomata are most
(i.e., mesoperigenous
stomata
stomata
have
of polocytic
a
rather restricted distribution.
stomata
it is also the basic form from which
universal in ferns
practically
sepcopolocytic, hemiparacytic, cohemiparacytic,
cyclocytic, cocyclocytic,
diacytic, codiacytic,
through
and
subsidiary cell(s)
growth
the
during
metaphase spindle
expansion
in
a
and other
of the
developing polocytic
types
of stomata, and also the factors that
types.
The
pericytic
the initial
wall links
As
stomata
and
stage
are
already
5
indicates
the
copericytic types
technically
of
remain
pathways
have all
of
about the
paragraphs
are
division(s),
re-
disharmonious
on
brought
forth these
of various
development
development
of different
the
of stomata,
mesoperigenous
stomata
been
cell and gcmc, and reorientation
stoma
bring
indicated in the
and all their derivatives
and
not
Figure
pericytic,
ontogenetically
surrounding cells,
subsidiary
evolved forms.
stomatal
have
staurocytic types
more
polocytic
Ophio-
pseudohemiparacytic,
desmocytic, codesmocytic,
certain elaboration or modification. Additional cell
orientation of the
rate
in
seppolocytic, pseudopolocytic, plupolocytic, pseudoco-
the
copolocytic,
(except
all other stomatal types,
polocytic,
derived
stomata
pluhemiparacytic, desmocytic, codesmocytic, pericytic, coperistaurocytic
especially
copericytic,
anomocytic
have the second-widest distribution.
1986)
1981, and Sen,
occurrence
pluhemiparacytic,
distributed
widely
(except Ophioglossaceae) recognised by Crabbe, Jenny
& Mickel
of
the
mother
in the
only
in the
reported by Hildebrand (1866),
could
type
freaks.
as
to
occurs
stomata
Ophioglossaceae), Protoleptosporangiatae,
remaining twenty-three
one
as
under in-
guard-cell
acts as
organisation
plants
since in this
simple,
of this type of
occurrence
and Mehra & Soni
(1969),
Thurston
(1959),
1).
in the
recognized
most
after its differentiation directly
and Dicksoniaceae,
Osmundaceae,
be the
this type of stomatal
(gcmc). Incidentally,
glossaceae (see
stomata
to
type appears
ontogeny
in
origin (Figs. 1-4).
originate mesoperigenously
mesoperigenous
as
lost. After the detachment of the link
long
walls,
as
only
in
the radial anticlinal
the
stomata
secondar-
BLUMEA
250
VOL.
37,
No.
1,
1992
Table 1. Distribution of stomatal
types.
Explanation:
1
=
=
hemiparacytic
17
2
=
polocytic
10
=
cohemiparacytic
18
=
copolocytic
11
=
pseudohemiparacytic
4
=
seppolocytic
12
=
pluhemiparacytic
5
=
pseudopolocytic
13
=
6
=
plupolocytic
14
=
7
=
pseudocopolocytic
15
=
3
8
*
9
anomocytic
*
*
sepcopolocytic
=
newly
*
16
*
*
=
desmocytic
19
=
codesmocytic
20
=
pericytic
paracytic
21
=
copericytic
coparacytic
22
=
diacytic
23
=
codiacytic
24
=
pluparacytic
*
cyclocytic
=
cocyclocytic
=
*
staurocytic
discovered
+
=
stomatal
type present
++
=
stomatal
type
(+)
=
stomatal
type infrequent
more
abundant than
the other associated
Adiantaceae
Ophioglossaceae
1
Botrychium daucifolium
Adiantoideae
+
1 +
lanuginosum
Pellaea
1
Helminthostachys zeylanica
1
evecta
2
sp.
(+),
2 +, 16 ++,
hypoleuca
2
fraxinea
16 +, 17
Taenitis
(+)
2 +, 16 ++,
regalis
17
(+)
Todea
2
2
(+),
barbara
4
4 +
3 (+),
4
(+),
villosa
Mohria
20
(+),
2
2 +,
(+),
caffrorum
+,
3
21
2 ++, 3 +, 4
3 (+),
(+),
4
4
(+)
(+)
raddianum
(+),
18
(+),
9
(+)
+, 4 +
2
++,
(+)
++, 9
+,
2
3 (+),
++,
elongata
3
(+)
5
++,
4
3 (+),
+, 4
(+)
+,
(+),
5
(+),
3 (+),
4
4
5
(+)
+,
5
(+),
thalictroides
(+)
2
2
(+),
4
(+)
++,
++,
3
+,
4
(+),
3
+,
3
+, 5
(+)
9
(+)
(+),
4 +, 9
(+)
Pteridoideae
9
+
2 ++,
2 ++, 3 +, 4
Pteris aff. biaurita
(+)
2 ++, 3
Viitaria
(+).
biaurita
Ceratopteris
+,
2
3
2 +,
ensiformis
Parkeriaceae
5
4 +, 9
2 ++, 3 +,
2 +, 3
Antrophyum parvulum
circinnatum
microphyllum
19
(+),
(+),
Vittarioideae
(+)
Anemia
3
2 ++,
philippense
2 +.
Schizaeaceae
Lygodium
(+)
9 (+)
Plagiogyriaceae
22
5
5(+)
+
Plagiogyria glaucescens
+, 4 +, 5
2 ++, 3
caudatum
hispidulum
(+),
3
2 ++, 3 +, 4 +, 5
Coniogramme japonica
4 +
(+),
++,
blechnoides
Adiantum
claytoniana
+,
(+)
(+)
arifolia
palmata
Osmundaceae
Osmunda
+, 9
Hemionites
(+)
17 +
16 +, 17
(+),
4
++,
calomelanos
Pityrogramma
+
4
Angiopteris
2 ++,
induta 2
Trachypteris
+
1
3
+,
2 ++, 3 +, 4 +, 5
7(+)
Marattiaceae
Marattia
viridis
Doryopteris pedata
+
1 +
Ophioglossum bergianum
reticulatum
2
Cheilanthes farinosa
1 +
virginianum
nudicaule
type(s)
9
(+),
10
2
3
++,
10
catoptera
2 ++, 3 +, 4
2
(+), 5 (+),
(+)
(+),
+,
3
+,
12
4 +, 5 +, 6
(+)
++, 4 +, 5
(+)
(+),
U.
&
Sen
Pteridoideae
(Adiantaceae /
B. De:
Structure
and
ontogeny of
Pteris
251
ferns
Microsorioideae
(Polypodiaceae /
continued)
in
stomata
continued)
2
Phymatosorus scolopendria
dalhousiae
2
7
++,
2
ensiformis
+,
4
5 (+), 9 (+)
+,
4
5
+,
++, 4 +, 5
(+)
10
(+),
(+),
2
4
+,
5
+,
3
(+),
+,
4
5
+,
2 ++,
+, 4
3
9
(+),
2 +, 3
(+), 10 (+)
4
(+),
(+)
(+)
2 ++,
9
4 +,
9
+,
2 ++, 3 +, 4 +,
linearis
13
2 ++, 3 +, 4
10
(+),
2 +, 3
rigidula
2 +, 3
4
5
++,
+,
2
3
++,
+,
(+)
4
+,
++, 3 +, 4
5
(+),
coronans2
9
(+),
18 (+),
(+)
+, 3 ++, 4 (+), 5
++, 3
+,
19 (+), 20 (+),
21
(+)
(+),
9
heteractis
18
(+),
lanceolata
2
2
3
(+),
+
3
9
(+),
20
2
3
19 +,
+,
+, 21
+,
+,
18
+,
2
7
+,
2
punctalum
stigmosum
16
Phymatosorus
lucidus
10 (+)
5
2
2
2
3
+,
4
++,
(+),
(+), 9 (+)
3
4
++,
++,
sarawakense
(+),
2 ++, 3
3
++,
(+),
3
++,
++,
3
+,
3
4
4
(+),
4 +,
+,
5
9
(+)
(+)
5 (+),
commutatus
+,
9
++,
9 (+),
++,
3
+,
10
4
2
+,
3
++,
5
+,
(+)
(+)
4
+,
(+),
4
+
2 ++, 3 +,
9 (+),
2
++,
4
+,
+,
3
(+),
5
(+)
4 +
2
3
(+).
15
+,
9 +, 10
(+),
(+)
(+)
14
++,
+, 15
4
9 +,
(+),
(+)
2
roraimensis
9
(+),
+,
3
++, 4
(+),
(+)
canalicular
2
2 +, 3 ++, 5
3
++,
9
(+),
(+)
(+), 9 (+),
+, 4 +, 5
(+)
(+),
2
3 +, 4
(+),
5
2 ++,
3 +, 4
(+),
5
2
3 +, 4
+,
(+),
10 (+)
tussacii
(+),
(+)
++,
5
(+),
(+),
(+)
2
3
+,
+,
4
5
(+),
+,
6
(+),
9 (+)
Sphaeropteris
(+),
Dicksonia
9
(+)
(+),
14
metteniana
5
+
3
++,
Trichopteris oblonga
9
(+)
5
(+),
(+),
2
Alsophila gigantea
5
(+),
+,
lanceolata
rostrata
Nephelea
6 (+),
pteropus
4
5
(+),
2 ++, 3 +, 4
(+)
latebrosa
9
+
2
+
+,
Cyatheaceae
10
musifolium
Microsorum
(+),
3 +, 4
(+),
Lophosoria quadripinnata 2 (+),
Cyathea
++
18
Microsorioideae
8
2 ++, 3
Lophosoriaceae
5
21 +
++,
+,
Metaxya
Cnemidaria
(+),
(+),
Drymoglossum piloselloides
+,
16+, 17+,
19 +, 20 +, 21
+,
(+),
5
(+),
2 +, 3 ++, 4
Metaxyaceae
13
Platycerium grande 2
5
(+),
13
Platycerioideae
5
2
subecostata
(+)
9 (+)
splendens
19
(+)
+
aureum
Loxogramme
5
(+),
Aglaomorpha meyeniana
Pseudodrynaria
(+)
Grammitidaceae
(+),
(+)
Photinopteris speciosa
20
5
(+),
lepidopteris
4
9
+,
8
(+),
Goniophlebium subauriculatum
(+),
(+)
3
5(+)
(+)
Drynaria quercifolia
10
2 +, 3 ++, 4
Polypodium formosanum 2
(+).
Pyrrosia
3 +, 4 +, 5
++,
nudus
Phlebodium
(+),
(+)
+,
2
phyllitidis
4
Drynarioideae
5
+, 4
2 ++,
(+), 5 (+)
Lepisorus
4
Polypodiaceae
5
2 ++, 3
Niphidium crassifolium
3 (+),
(+),
2 ++, 3
Dicranopteris
++,
Polypodioideae
(+)
laevissima
3
(+),
(+)
Campyloneurum angustifolium
Gleichenia gigantea
22
+,
Selliguea feei
Gleicheniaceae
9
10
9 (+),
Pleopeltoideae
aureum
speciosum
3
++,
(+), 5
Crypsinus albido-squamatus
4
(+)
vittata 2 ++,
Acrostichum
4
Christiopteris tricuspis
6
(+),
(+), 9 (+)
semipinnata
9
+,
3
++,
2 +, 3
parviloba
3
9
elmeri
antarctica
(+),
2
10 (+),
2
13
++,
(+),
9
3
+, 4
(+),
(+)
3
++,
4
+,
5 (+),
(+)
Cystodium sorbifolium
5
++,
(+)
2 ++,
3
+,
4
(+),
BLUMEA
252
(Table
1
VOL.
continued)
14
14
(+)
(+)
4
(+),
13
(+),
++,
2
bipinnata
2
3
4
(+),
3
++,
2
speluncae
4
+,
2
3
++,
+,
3
+,
2
luzonica
curroii 2
9
+,
4
+,
4
2
+,
+,
4
+,
9
+,
(+)
(+)
5
(+),
(+)
(+)
3
(+),4
4
(+),
3
4
++,
5
+,
(+)
+,
+,
5
(+)
5 +, 10
(+)
5
(+),
9
2 +, 3 ++,
9
2 ++,
3
+, 5
2 ++,
immersa
3 +, 4 +, 5 +, 9
3
2 ++,
2 ++,
uliginosa
Pseudophegopteris
+, 4
(+),
+, 4
3
aurita
10
5 (+), 9 (+),
hirtirachis
(+)
5
(+),
5
+
(+)
(+), 5 (+)
2 ++, 3
2
4
+,
(+)
3
4
+,
(+),
5
+,
2
9
(+)
3 +,
++,
4
5
+,
(+)
5 (+)
2 ++, 3
(+),
4 +,
(+)
2
3
++,
2
5
+, 4 +,
++,
3
(+),
2 ++,
2 ++,
(+)
4 +, 5
2 ++, 3
4
+,
jaculosa
+, 4
3
+, 4
3
(+)
5
(+),
(+)
5
(+),
(+)
(+), 5 (+)
Asplenioideae
2 ++,
3
2
formosum
2 ++,
++,
2
3
++,
2
glaucophyllum
(+),
hemitomum
lucidum
+, 4 +, 5
auriculata
10
2
(+), 9 (+),
2 ++, 3 +, 4
(+),
(+)
(+),
++,
Macrothelypteris
3
+,
4
ornata
(+),
5
(+)
2 ++, 3 +, 4
(+),
4 +
3
4
(+),
3
+, 4 +
+,
++,
Diplazium
2 ++,
3
+, 4
(+),
9
(+)
4 +, 5
5
(+)
+
(+)
2 ++,
3 +, 4 +,
2 ++, 3 +, 4
esculentum
2
2
++,
(+)
2++, 3+,4
++,
3
3
++,
3
++,
+,
4
4
+,
+,
5
5
+,
+
(+)
6
(+)
6
(+),
Tectarioideae
Ctenitis
9 (+)
4
(+),
(+),
4 +
(+)
5(+)
proliferum
(+),
(+),
4 +, 5
Athyrioideae
pectinatum
arthrotricha12 ++, 3 +, 4
3
++,
3 (+),
2 ++, 3
2 ++, 3
polypodioides
torresiana
++,
+,
+, 4 +,
3
Athyrium filix-femina
2
(+)
(+), 10 (+)
2 ++, 3 +, 4 +,
3 (+),
2 ++,
falcatum
+, 4 +,
(+)
viscosa
2 ++,
repens
dalhousiae
10 (+)
Cyclogramma
5
3
Asplenium africanum
(+)
2 ++, 3 ++, 4 +, 5 +, 9
rectangularisr
(+),
(+),
(+)
parasitica
10 (+).!!(+)
Coryphopteris
+,
Aspleniaceae
baramensis 2 +, 3 ++, 4
harveyi
10
(+),
9
(+),
ecallosa
++,
molliuscula
(+)
Thelypteridaceae
9
5
(+),
arida
2 ++, 3 +,
darjeelingensis
(+)
3
++,
(+)
2
dentata
Tapeinidium pinnatum
Thelypteris
2
Pseudocyclosorus esquirolii
5
(+)
(+)
yar.
(+),
+, 5
coriacea
ensifolia subsp.
9
3
++,
(+)
invisus 2 ++, 3 +, 4
Christella appendicular
2
chienii
odorata
9
(+),
tylodes
(+),
5
2 ++, 3 +, 4
2 ++, 3 +, 4
truncata
(+)
2 ++, 3 +,
subdigitatum
(+)
(+), 5 (+), 9 (+)
(+),
2 ++, 3 +, 4 +, 9
penniger
5
4
4 +, 5
(+)
10 (+)
Pneumatopteris
(+)
+,
(+)
5 (+)
Lindsaeoideae
5
+, 4
+,
5
5
(+),
(+)
(+)
11
Sphaerostephanos
4
Monachosorum
Lindsaea
3
nudatum
5
3
++,
(+),
Ampelopteris prolifera
Monachosoroideae
4
++,
5 (+),
unitus
5
+,
+,
9 (+),
(+), 5 (+)
++,
3
3
9 (+)
2 +, 3 +, 4
3
+,
incisa
Histiopteris
Blotiella
2
++,
++,
:
Hypolepis punctata
Paesia
2
+, 4
3
(+),
4
+,
2 +, 3 +, 4
striatus
4
Microlepia platyphylla
strigosa
2 ++,
++, 4
Mesophlebion crassifolium
++,
2
tenera
3
++,
Pronephrium
Dennstaedtioideae
scabra
3
extensus
2
gongylodes 2
9
(+),
Dennstaedtiaceae
Dennstaedtia
2 +,
flaccida
ferox
15 (+)
+,
1992
Cyclosorus
(+), 9 (+),
15 (+)
+,
2
hawaiiense
4 +, 9
(+),
4
glaucum 2 (+).
4 +, 9
3 (+),
2 ++, 3
Culcita macrocarpa
++,
1,
Metathelypteris dayi
2 ++,
Thyrsopteris elegans:
13
No.
(Thelypteridaceae continued)
Thyrsopteridaceae
Cibotium
37,
ampla
9 (+)
2
+,
4
+,
5
+,
U. Sen
&
B. De:
Structure
and
(Aspleniaceae / Tectarioideae continued)
(Ctenitis)
eriocaulis
Pleocnemia
2 ++, 3 +, 4 +, 5
irregularis
9
Tectaria
(+),
2 +, 3 ++, 4
2 ++,
2
+,
3 ++, 4
7
(+),
Cyclopeltis
3
+, 4
24
5
(+),
4
(+)
+, 4 +,
3
aculeatum
2
2 ++,
3 +, 5
4 +,
9
(+),
5
(+),
9
(+),
(+)
22
(+),
3 +, 4 +, 5 +,
(+)
Rumohra
(+),
2 ++,
3
4 +, 5
(+),
2 ++, 3
Dryopteris filix-mas
+, 4
(+),
22
heteroclita
3 +, 4 +,
2 ++,
5
(+),
2 +, 3 ++, 4 +, 5 +,
2
singaporensis
++,
2
3
++,
Lomariopsis
+,
3
4
+,
(+),
4
5 +
(+),
(+),
++, 4
(+),
22
23
5 +
2 ++,
3
+,
+
+,
2 ++, 3 ++, 4 +, 9
22
+,
2
3
2 ++,
3
+,
2 ++, 3
+
(+)
3
++,
+,
+, 4 +,
(+)
(+)
+, 5
(+)
2 ++, 3
+, 5
+
(+)
2 ++, 3
paucivenius
17
(+),
24
+,
24
+,
monocarpa
2
+,
3
5
+,
+
(+),
(+)
2 ++, 3 ++,
+, 5
16
(+),
17
+,
16
17
+
(+),
17
(+),
(+),
22
(+),
(+), 23 (+)
2
++,
3
16
+,
+,
(+)
2 ++, 3 +, 4
orientale
Sadleria
sp.
2 ++,
3 +, 4
(+),
5
+,
2 ++, 3
Woodwardia radicans ■ 2 ++,
tenuifolia
2
palustris
2 ++, 3
9
(+),
+, 4
(+),
2 +, 3 +, 4
pallida
Stenochlaena
Davallioideae
2 ++, 3 +, 4
brasiliense
(+),
9 (+)
punctatum
+
(+)
9
(+)
(+)
3
+, 4
+, 4
5
(+),
3 +, 4
++,
(+)
(+)
(+)
Marsileaceae
lanuginosa
ophioglossa
2 ++,
2 ++,
(+),
22
3
3 +, 4
Trogostolon falcinellus
+, 4
(+)
(+),
22
Marsilea
(+)
2 ++, 3 +, 4 +,
(+),
22
2
Regnellidium
++,
3
(+),
3
+,
4
(+),
4 +
+
2 +, 4 +
diphyllum
Salvinia
cucullata 2
++,
4
+,
9
Azollaceae
(+)
undulatus
2
drummondii
minuta
Salviniaceae
(+)
Scyphularia simplicifolia 2++,3++,4+,
Parasorus
2
2 +, 3
2 ++, 3
Blechnum
(+)
Davalliaceae
5
2 ++, 3
(+)
cordifolia
+,
2 +, 3 ++, 4 +, 5
cochinchinensis
5
Elaphoglossum
5
(+),
(+)
africana
falcata
5
Elaphoglossoideae
Humata
+, 22
5
Blechnaceae
2 +, 3
sp.
Teratophyllum ludens
9
9
Arthropteris
9 (+)
8(+)
4
2 ++, 3 ++,
4 +,
adiantiformis
exaltata 2 +, 3
(+)
semicordata:
Bolbitis
3 ++,
9
(+),
articulata
16
crispatula
+
Oleandroideae
Lomariopsidoideae
9
(+)
9
+
Nephrolepis acutifolia
(+)
+,
(+)
Psammiosorus
(+)
5
9
(+),
5 (+),
+,
2 ++, 3 ++, 4 +, 5
(+),
tenella
cervina
Bolbitis
5
neriiformis
obtusissimai 2 ++, 3 +, 4
9 (+)
Olfersia
2 ++,
(+),
Oleandra
+
2 ++,
Cyrtomium falcatum
8
22
(+),
4 +, 5
(+)
3 (+),
++,
+,
(+)
pallida
2 ++, 3 +, 5
4
++,
++, 4 +, 5
3
3 ++, 4
++,
Leucostegia immersa
5
(+),
3
++,
grammatosorum
4 +, 5
(+)
Arachniodes
2
Gymnogrammitis dareiformis
anomalum 2 ++, 3 +, 4
6
Davallodes
5 (+),
truncatula 2 ++,
cyatheoides
varium
2 ++,
2
hirsutum
(+)
(+)
Polystichum
5
divaricata
5 +
Dryopteridoideae
Peranema
denticulate
Araiostegia hymenophylloides
2 ++,
Didymochlaena
+,
(+)
(+)
sp.
9
5
(+),
2 ++, 3 +, 4 +,
subtriphylla
5
(+),
+, 4 +,
3
continued)
9(+)
fijiensis
fernandensis
incisa
5
+, 7
(+)
Pteridrys australis
6
5 (+)
+,
253
ferns
Davallioideae
Davallia
(+)
3
2 ++,
in
stomata
(Davalliaceae /
2 ++, 3 +, 4
hemiteliiformis
ontogeny of
2 ++, 3 ++, 4
(+)
Azolla
pinnata
2
+,
4
+,
9
(+)
(+), 13 (+)
VOL.
BLUMEA
254
No.
37,
1,
1992
Table 2. Distribution of stomatal types in different families and subfamilies
(sensu Crabbe
et
1975).
al.,
*
Newly
=
discovered.
*
*
*
O
"3
o
*
f
o
i
*
*
a.
Families /
Subfamilies
anom cytic pol cytic cop locytic sep ol cyti pseudoplcyti plu ol cyti pseudocplocj sepco lcyti hemipar cyti cohemiparcyti
*
e
o
-C
0
"3
O
v.
pluhemiparcyt par cytic coparcytic pluarcyti cylocytic coycloytic desmocytic codesmocyti pericytic copericytic diacytic codiacytic staurocytic
a.
Ophioglossaccae
+
Marattiaceae
+
Osmundaceae
+
Plagiogyriaceae
+
+
+
Schizaeaceae
+
+
+
Parkeriaceae
+
+
+
+
Adiantoideae
+
+
+
+
Vittarioideae
+
+
+
+
Pteridoideae
+
+
+
+
+
+
+
Drynarioideae
+
+
+
Platycerioideae
+
+
Microsorioideae
+
+
+
Pleopeltoideae
+
+
+
Polypodioideae
+
+
+
+
Grammitidaceae
+
+
+
+
Metaxyaceae
+
+
Lophosoriaceae
+
Cyatheaceae
+
+
+
Thyrsopteridaceae
+
+
+
+
+
+
+
+
+
+
+
+
Adiantaceae
Gleicheniaceae
+
+
+
+
+
+
+
+
+
+
+
Polypodiaceae
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
Dennstaedtiaceae
Dennstaedtioideae
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
Asplenioideae
+
+
+
+
Athyrioideae
+
+
+
+
+
Tectarioideae
+
+
+
+
+
Dryopteridoideae
+
+
+
+
+
Lomariopsidoideae
+
+
+
+
Elaphoglossoideae
+
+
+
+
Davallioideae
+
+
+
+
Oleandroideae
+
+
Blechnaceae
+
+
+
Marsileaceae
+
+
+
Salviniaceae
+
+
+
Azollaceae
+
+
+
Monachosoroideae
Lindsaeoideae
Thelypteridaceae
+
+
+
+
+
Aspleniaceae
+
+
+
+
+
+
+
+
+
Davalliaceae
+
+
+
+
+
+
+
+
+
+
+
+
+
U. Sen
Fig.
5.
tion
of
cell(s).
of the
ily
Evolutionary
metaphase spindle
become
Hennipman
precursors of the
& Lersten
gcmc.
F
The
that the
and
stomata and
=
of these types of
tween
the
ontogeny of
their
in
255
ferns
=
Additional
segmentation
subsidiary cell(s).
of additional
subsidiary
C
=
cells
in
the
=
Orienta-
subsidiary
Change in orientation
on
both sides.
thus confirms the observations of Sen &
present study
polocytic
and
and
copericytic
stomata
polocytic/copolocytic
short-lived. Moreover, it is
stomata
ontogenetic interrelationships. — O
A
cells and
Formation
Fryns-Claessens
origin
and
neighbouring cell(s).
pericytic
(1967),
Structure
growth of guard
in
mesogenous.
(1981)
fern
and
Disharmonious
=
B. De:
trends in
subsidiary cell(s)
D
&
not
copolocytic
stomata,
stomata
respectively.
& Van Cotthem
(1973)
are
the
Pant
did
ontogenetic
(1965),
not
Mickel
describe the
correctly, probably because the transitory stage
and the
at
all
pericytic/copericytic
possible
to
condition is
detect this stage unless
be-
extremely
one
fixes the
BLUMEA
256
material
of
at
the
developing
gory of
ting
right
into
stomata
moment.
or
The
distinct lines:
one
from the
in
not
in the
only
of this
acterised
association,
stomatal
two or more
we
view that the
sophyta,
and
collateral
in
condition, elliptic
stem
filicalean
stomata,
ceae are most
article Kato
should be
to
same
It
can
be
on two
hemiparacytic
one.
stomatal
two
types
established fact in ferns that
a
of
extant
In the
1).
genera
char-
member of the
treatise)
conspic-
are
(Sen,
distinct division, the
1967)
Ophioglos-
of collateral cauline bundles,
coiled leaves
stalk in the
group
other
any
occurrence
in the
pits
lamina (see Table
thus supports Sen's
tracheids,
even
in the
embryonic
and the lack of sclerotic
Ophioglossum
extant
group
a
are
extant
hairless
similar
upright simple
and
of
genera,
are
are
vascular
to
pro-
Ophioglossa-
of oval circular border-
towards the distal end of
cylinder
differ
'aerial
as
only
Unfortunately
from the progymno-
in the
are
ophioglossoids
not
in
of
the
a
possession
numerous stout
tap
radially
less pit membrane and mucilage ducts which all
diploxylic
roots
of
cylinder,
traces
perigecortical
are
char-
originating
often associated with apo-
seriate
seem to
of tori in
progymnosperms.
this, however, the cycads
number of
antero-
Bierhorst, 1971)
development
living
rounded stems, eustelic vascular
to
of
possession
complex'
also in the
ophioglossoids
the
not
cryptogams
storage root,
from the bases of the stems,
plants.
possession
living
are
the
non-arborescent, fleshy and devoid of any
stipular outgrowths. Contrary
cauline strands,
ophioglossaceous
plants,
of the progymnosperms. But unlike any of the
acterised by circinately coiled leaves with
roots
seed
indeed resemble the progymnosperms.
stomata
ophioglossoids
fleshy,
Ophioglossaceae
extant
In the
phyllo-sporangiophore (i.e.
cycads
stomata,
separate
that the
that among
of eustelic vascular
membrane. So it appears that
geotropic
in the
distinct entities. In
that these
same
in
placed
Moreover, the ophioglossoids
associated with
from
stomata
have evolved
of the present
sense
Cycadophyta.
to
development
divided
parenchyma
copara-
be considered as the direct de-
Ophioglossum
structure
(1988) suggested
sperms but also from all other
The
of
two
stomata
now an
non-circinately
presumed
ophioglossoids
sclerenchyma.
nous
subsidiary
mesoperigenous type.
reason
It is
related
the
(in
common
clearly related
progymnosperms, all
pit
not
circular bordered
is known about the
posteriorly
are
Stomatal
2).
stalk traces,
and in the
and in the
the stem, the
the
represent
not
but also in the
stomata
1 &
further
gymnosperms. He
nothing
off
properties.
a recent
pits
simple
Filicophyta. Certainly the
to
their cut-
form paracytic,
the other from the
conclude that the
Ophioglossaceae
not
for the
accidently.
taxon
anomocytic
common
cells in the
type,
even
same
absent (Tables
uously
cut
cate-
a new
over
them often
types having ontogenetic interrelationships among
by anomocytic
Filicales where
ed
not
derived form of the
a
definitely
can
stomata
ultimately
mesogenous types
forms do
two
desmocytic stock,
scendant of the anomocytic
never occur
In
and
of stoma, however, should
polocytic type
The
stomata, instead
simply by changing
mesogenous
gcmc
and
mesoperigenous
shows that the
taxa
produce
inferred that in ferns the mesogenous types of
safely
light
1992
stomata.
of
occurrence
to
differentiating
fact, the mesogenous type is
occur
1,
developing hemiparacytic
the mesogenous type, that is,
-
pluparacytic
frond in many
No.
37,
hemiparacytic forms, often initiate the development of
both sides before
on
cytic,
not
Some of the
face. The initials destined
cells
VOL.
secondary phloem,
exclude any
torus-
affinity with
the
U.
Taking
stomatal features
Bierhorst's
port
which
Structure
as
(Bierhorst,
characterised
are
and the
pteris strongly
a
and
ontogeny
parameter
1968,
of
presence
of
1973)
1971),
257
ferns
it is also very difficult
affinity,
anomocytic
all ferns.
at
are
terminal
plurilocular
stomata in
of
contention that Psilotum and
by having only perigenous
1963; Pant & Khare,
Mehra,
phylly
B. De:
Sen &
to
sup-
Tmesipteris,
stomata
(see
Pant &
Indeed the absence of megain Psilotum and Tmesi-
sporangia
indicate their non-filicalean nature.
ACKNOWLEDGEMENTS
This
paper is
friendship
sor
K.U.
to the
dedicated
and
of
memory
to the
guidance
senior
Kramer and Dr. H.P.
R.E. Holttum in
Professor
author
for over three
(U.S.)
for suggesting
Nooteboom
many
view
of his
scientific
support,
decades. The authors thank
Profes-
modifications.
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W. 1971.
Rauter,
1971.
Bernh. Ann.
Bharati
1881.
Farnen.
1992
1,
420-424.
morphologisch
Probst,
Epidermal
No.
37,
Bot. 33: 795-805.
sipteris tannensis
Sci.
1969.
VOL.
and
in
stomatogenesis
Azolla pinnata
Bot.
P. Brown.
46: 957-968.
Bot. 52:
Ann.
201-204.
Sen, U.,
&
Sen, U.,
1984. Stomatogenesis
S. Bhunia.
montana and
& E.
D.
Blumea
splendida.
30:
in
1981. Structure and
Hennipman.
Gleichenia
linearis
gigantea, Dicranopteris
var.
13-16.
ontogeny
of stomata
in
Polypodiaceae.
Blumea
27:
in
the
175-201.
& C.S.
Stebbins, C.L.,
leaf
of
epidermis
Thurston,
E.L.
Khush.
1961. Variation
and its
monocotyledons
1969. Taxonomic
in
the
organization
bearing on
significance of
their
stomatal
of the stomatal
phytogeny.
patterns
in
Amer. J.
complex
Bot. 48: 51-59.
ferns. Amer.
J.
59: 63-
of some
Kenyan
Fern
79.
Viane, R.,
& W.
van
Cotthem.
1977.
Ber. Deutsch.
Asplenium-species.
Spore morphology
Bot. Ges. 90:
and stomatal
characters
219-239.
APPENDIX
List of
studied.
species
All
material
housed at the
University
Ophioglossaceae
—
lanuginosum
U.
Sen
De
—
nudicaule
—
Plagiogyriaceae
Schizaeaceae
—
—
—
Adiantaceae
Ta3;
A.
Kaulf.:
L.:
U.
melanos (L.)
U. Sen
S75;
Grev.:
cult Kew;
U. Sen
virginianum (L.) Swartz:
Ophioglossum
-
L.: B. De
(Forst.) Hoffm.:
Sm.: U. Sen
Sen
thalictroides
L.: B.
Fee:
K28
Willd.: cult Kew
Link: U. Sen Kll
O.
the
s.n.; B.
B. De
bergianum
SB1
-
Schlecht.:
M6.
B. De
Dar9;
A.
hypoleuca
K85.
regalis
L.: B. De P13
-
Todea
barbara
-
-
Lygodium
Mohria
(L.) Brongn.:
B. De
De
raddianum
B6;
-
A.
Pellaea
K106
circinnatum
(Burm.)
Hemionitis
Kln4;
A.
arifolia (Burm.)
-
Taenitis
U.
-
-
Sw.: B.
De
Cheilanthes farinosa
U. Sen
Moore: B.
Sen K125
blechnoides (Willd.)
B.
hispidulum Swartz:
Presl: U. Sen K108
viridis (Forssk.) Prantl:
Sen
U.
K197.
B. De
Coniogramme japonica (Thunb.) Diels:
-
Sw.:
Desv.: U. SenK114.
caffrorum (L.)
caudatum L.:
U. Sen K65
Link: U. Sen
-
B. De SB11.
Adiantum
U. Sen Kill
opteris pedata (L.)
mata
villosa
Adiantoideae:
philippense
(Forssk.)
L.:
s.n.
A. evecta
fraxinea
claytoniana
B.
O. reticulatum
B5;
Plagiogyria glaucescensChing:
Anemia
Ceratopteris
—
U. Sen
K63;
Hook. &
ex
02;
Paleobotany Laboratory of
otherwise.
Sen K47.
K21;L. microphyllum
Parkeriaceae
Marattia
-
Osmunda
U. Sen
De
and
unless stated
Wall,
B.
Hook.:
L. f.: B. De
Angiopteris sp.:
Moore: U.
& Grev.:
zeylanica (L.)
Vriese: U. Sen K68
Osmundaceae
(L.)
O.
SA5;
Hook.
ex
Pteridology
Kalyani,
Botrychium daucifolium
Wall,
Helminthostachys
Marattiaceae
of
De
K50
Dory-
-
Ke4;
H.
Pityrogramma
De A2
-
pal-
calo-
Trachypteris
Sen
U.
induta
(Maxon)
A10
Vittaria
-
R. & A.
B.
Tryon:
B. De
P. biaurita L.: M. Mishra
De
P.
Klnl8;
semipinnata L.:
Gleicheniaceae
—
K97
Dar21;
U.
—
K74
Sen
Sen K28
Pichi
Sen
auriculatum
(L.)
C.
Metaxyaceae
Cyatheaceae
-
—
—
R.
&
U. Sen
G.
s.n.;
A.
K98;
P.
K35
-
laevissima
U. Sen K33
-
&
U.
Sen
(Mett.
AB3
K102;
Presl:
A3
-
A.
Microsorioideae:
—
—
M.
pteropus (Bl.) Copel.:
Phymatosorus
-
Sermolli:
U. Sen
U.
s.n.; P.
Sen
Sen K127
(Bl.)
K79
scolopendria
—
Pleoipeltoi-
Polypodioideae: Campyloneurum
Presl: U.
Holt-
(Baker)
commutatus
angus-
Goniophlebium sub-
-
U. Sen
Niphidium crassifolium (L.) Lellinger:
-
Mett.)
ex
B. De
Kuhn) Ching:
ex
M. sarawakense
Selliguea feei Bory:
s.n.
Bedd.: U. Sen
(Wall,
coronans
Crypsinus albido-squamatus (Bl.) Copel.:
Pichi
U. Sen K129
splendens (J. Sm.)
rigidula (Sw.)
Bhattacharyya
-
phyllitidis (L.)
K126
Nakai:
Polypodium formosanum (Baker)
U.
-
Sen
Kunze: U. Sen K123.
Fisch.)
lanceolata
(Sw.)
Presl:
A.
L.
Bhattacharyya 31;
Presl: K.U. Kramer
(H.B.K.)
Tryon:
R.
(Domin)
antarctica
(CAL)
—
D.
heteractis
U. Sen K87
(Roxb.)
J. Smith: U. Sen K15
rostrata
subecostata
s.n.
(CAL)
elmeri
H.H. Smith 2224
Culcita
-
149; A.
cult. Kew
-
metteniana
Hance:
canalicular
Cyathea
(Copel.)
Nephelea
tussacii
R.
CAL
Tryon:
(Desv.)
10585
67,
s.n.
J. Smith:
Cystodium sorbifolium (J.E. Smith)
-
AFS
Willd.:
R.
Tryon:
Trichopteris
-
(CAL).
glaucum (Sm.)
cult. Kew
-
s.n.
C. Chr.: cult. Kew.
N.C. Das
Tryon;
Labill.: cult. Kew
Sphaeropteris
-
Cibotium
Dennstaedtioideae:
—
bipinnata (Cav.)
Mett.: U.
Sen Kl
Hook. & Arnott: cult.
Kew;
Presl: cult.
macrocarpa
—
(Alderw.)
s.n.
Histiopteris
Sen:
—
U. Sen
B.P.S. s.n.
Kew
—
&
(Hook.)
(Thunb.)
T. Sen
Jermy:
Holttum:
B.
K69;
s.n.;
De
Holttum
tricha Holttum: Holttum
R.
(Wall.)
J.
M.
strigosa (Thunb.)
chienii
-
C. hawaii-
Thyrsopteris elegans
-
L. odorata Roxb.
C. Chr.: U. Sen
C.
var.
arida
(D.
s.n.;
C.
C.
Don)
Sen K131
J.
-
A9;
D75;
U.
-
Sen
Paesia
subdigitatum (Bl.)
L.
ensifolia
darjeelingensis Sen
-
Denn-
D. tenera
Hypolepis
Sm.:
Presl: G.B. Nair s.n.
(Baker)
B. De Kln84
Holttum:
parasitica (L.)
viscosa
U. Sen K100
Sw.
punc-
K17;
Kuhn:
subsp.
M.
luzonica
G.P.
coriacea
& Sen: T. Sen
s.n.
-
K122.
H84/2; C. jaculosa (Chr.)
s.n.;
Tryon:
Moore: U. Sen
Sm.: U.
B. De
Ching:
Ampelopteris prolifera (Retz.) Copel.:
Holttum:
Brownsey
K5;
incisa
curroii
D. scabra
Microlepia platyphylla (D. Don)
-
Lindsaea
Tapeinidiumpinnatum (Cav.)
Thelypteridaceae
K42
K70;
Monachosoroideae: Monachosorum
Lindsaeoideae:
Kramer:
Blotiella
Maxon: U. Sen
Moore: U. Sen
Christ: G.B. Nair
Nair s.n.
-
Mett.: U.
speluncae (L.)
(Kuhn)
SMP8X;
cult. Kew.
(Thunb.)
(Presl)
C.
K86;
(Wall.) Copel.:
Ogura:
Dennstaedtiaceae
tata
U. Sen
roraimensis
1015
oblonga (Kl.)
(Pr.)
Mishra
Sen
Hook.:
Sen K120.
Dicksonia
Thyrsopteridaceae
staedia
M.
& T. Sen
U. Sen
K73;
Bhattacharyya AB1;
Alsophila gigantea (Wall.) Holttum:
-
Nakai
A.
K58
Lophosoria quadripinnata (J.F. Gmelin)
C. latebrosa
H.H. Smith
Kunze:
(L.)
Metaxya
—
A.
lucidus
(Hook.) Ching:
U. Sen
Chr.: U.
cult. Kew
ense
P.
Loxogramme
—
Cnemidaria
(CAL);
K95;
Presl:
(Bl.)
Sen
musifolium (Bl.) Copel.:
lepidopteris (Langsd.
Lophosoriaceae
142
F6e:
Farwell:
U.
(L.) Copel.:
Sermolli:
nudus
aureum
Grammitidaceae
(Hook.)
U.
Pseudodrynaria
-
Pyrrosia
-
stigmosum (Sw.) Ching:
U. Sen
Pichi
tifolium (Sw.)
Phlebodium
M.
K90;
Sermolli:
P.
P. dalhousiae
montana Holttum:
var.
Sm.: U. Sen
U. Sen K84
P. lanceolata
M. punctatum
Lepisorus
K83;
Underw.
Blume: U. Sen K61
Kunze:
Microsorum
-
K22;
(N.L. Burm.)
deae:
Acro-
P15K.
(subg. Diplopterygium):
Christiopteris tricuspis (Hook.) Christ:
tum: U.
Bl.: B. De
Pteridoideae:
Platycerioideae: Drymoglossum piloselloides (L.)
—
Bhattacharyya AB21;
U. Sen
—
K39;
Christ:
parviloba
M. Mishra
Drynaria quercifolia (L.)
-
Platycerium grande (Fee)
U.
P.
H27;
U. Sen
Drynarioideae: Aglaomorpha meyeniana Schott:
Sen K99
U.
Ching:
Sen K81
Willd.: U. Sen K16- Pteris aff. biaurita L.: U.
(Burm. f.)
Wall, ex Hook.
Photinopteris speciosa
-
De
259
ferns
Antrophyum parvulum
Sw.: U.
catoptera Kunze:
P. vittata L.:
linearis
Vitarioideae:
—
in
stomata
DeSBlO.
Polypodiaceae
Copel.:
P.
ontogeny of
ensiformis
V.
speciosum
SMP6X;
Dicranopteris
Gleichenia gigantea
Christ: B.
A.
Al;
Burm.: B.
ensiformis
B. De
U. Sen K119
B. De
A4;
and
Structure
B. De:
elongata Sw.:
stichum aureum L.:
K8;
&
B.
De
Holttum:
Lev.:
T. Sen
Christella
-
S84/1; C.
U.
s.n.
Holttum: Holttum
Sen
-
appendiculata
dentata
K23;
C.
(Forssk.)
molliuscula
Coryphopteris arthro-
s.n.
-
Cyclogramma
auri-
BLUMEA
260
U. Sen
culata (J. Sm.) Ching:
U. Sen K25
Ching:
s.n.; M.
Holttum
s.n.
Holttum
Das
(Gaud.) Ching: N.C.
T76
Sen
-
U.
(Poir.) Holttum:
s.n.
P. hirtirachis
DarB14;
Holttum: B.
(Zoll.)
De
immersa
ecallosa
—
deae:
-
K56;
Bedd.: B.
esculentum
U. Sen
Ching:
C. Chr.
&
K117;
Sen
DarB32
U. Sen K124
Sen
U.
P.
U. Sen K72
T. Sen
Sen
1178
D.
-
fijiensis
D.
Bot.
637-69;
Price
(L);
cult. Kew
cinellus
-
(Forst. f.)
(L.)
Genus
1035
ex
(L);
Hook.)
T. Sen
s.n.
B.
Fee:
Alderw.:
1074
N.
Olfersia
-
U.
B.
Cav.: Price
Presl var.
cult.
L.
Pleyte 363 (L);
& Hernaez
monocarpa
-
1478
Sll
B. De
aculeatum
(L);
T46
cult.
Humata
cult. Bot.
ex
Christ:
—
cordifolia (L.)
Joncheere SEU39
(L);
U.
Sen
Kew
Presl: T. Sen
(L);
recognised by Crabbe, Jermy &
T. Sen
Mickel
-
D19;
A19;
Schott:
Sen
U.
Sen
K115;
ludens
B.
(Fee)
divaricata
207-71;
Bl.:
Price
in
Alderw.:
Dept. Kalyani
Bot.
Rumohra
7358
Univ.
under
T. Sen
14393
M. Hotta
adiantiformis (G. Forst.)
cult.
s.n.
-
cult.
Dept. Kalyani Univ.
(L)
Kew
-
Trogosiolonfal-
362-76
T.
—
Olean-
Sen T53;
Nephrolepis acutifolia (Desv.)
U. Sen
(1975).
(L.)
Lomariopsi-
lanuginosa
Kuhn) Copel.:
s.n. under
K89
Kunze:
grammatosorum
(Cordem.) C.Chr.: Pichi Sermolli 6763 (L);
268
Presl:
(L.)
Ching:
D.
Jermy
Hirano & Hotta 172
299 (L); Craig
T. in-
Murata et al. T16067
A8;
Davallodes
397-68.39702; cult.
T. Sen
Ching
K62;
dareiformis (Hook.) Ching
-
-
Dryopteridoi-
cervina
Holttum: U.
under
amplissima
pallida (Mett.
Kew
K118
(Racib.)
U. Sen K112.
sp.:
Univ.
2759
—
Teratophyllum
-
Kuhn:
T. Sen
prolif-
australis
U. Sen
heteroclita (Pr.)
Sen K9
Price
DarB3
D.
sp.: U. Sen
Pr.: U. Sen K40
singaporensis
Mett. ex
(L);
B. De
Bonpl. ex Willd.)
U. Sen K64
Gymnogrammitis
-
D23;
under
J. Smith: Dockrill
(L);
(L.)
SB25;
De
Scyphularia simplicifolia Copel.:
Schott de
not
B.
(J. Smith) Copel.:
& Sleumer
&
Alderw.:
Athyrioi-
—
Cyrtomium falcatum (L. f.)
-
s.n.;
Hook.: B. De
SB6;
Pteridrys
-
Polystichum
-
Dept. Kalyani
Univ.
Presl:
B. De
Cyclopeltis
K104
Sen
varium
Ca2;
(Burm.)
Bot.
ophioglossa
(Presl) Copel.: Price
Surbeck
tala
H.
(Wall,
Walker
droideae: Arthropteris
tenella
hirsutum
(BO);
Parasorus undulatus
Ching:
denticulata
Hook.: cult.
Elmer 9900
1888
P.
cochinchinensis
Dept. Kalyani
immersa
U.
B. De DarB19
De
dalhousiae
Araiostegia hymenophylloides (C. Chr.) Ching:
Davallia
-
3881
Biinnemeijer
De DarB21
Elaphoglossoideae:Elaphoglossum
—
& C.Chr.: Smitinand
Leucostegia
B.
Holttum
glaucophyllum
ampla (Humb.
Sen K13
s.n.;
Thelypteris
-
Forst.: U. Sen K49
-
-
rectangularis
Pleocnemia hemitelliformis
-
s.n.
Holttum
K38.
pectinatum (Wall.)
K71
s.n.; P.
Holttum
SB4
fernandensis (Baker) C. Chr.:
SB2;
B.
(Moore) Ching:
Lomariopsis
s.n.;
cult.
Don:
B. De
A.
A.
T2;
subtriphylla (Hook. & Arn.) Copel.:
cyatheoides
B. De
U. Sen
Sen K53;
Ctenitis
P.
DarJ;
Holttum:
polypodioides Bl.:
Holttum: U.
Tectaria
Das
(Hook.) Ching: U. Sen
B. De
Proctor ex Iwats.:
B. De
Alston: U. Sen K66
crispatula (Wall.) C. Chr.:
-
T56
Price
(L);
Tardieu
(L);
T.
A.
Holttum
(Hope) Ching:
(Forst. f.)
lucidum
D.
K26;
Tectarioideae:
(F6e)
-
A.
DarS;
B. De
(Schum.)
N.C.
L56;
Holttum: U. Sen
aurita
L33;
Holttum:
(L.)
Desv.: U.
K7;
irregularis (Presl)
Davallioideae:
K57;
Copel.:
Kew
U. Sen
Holttum:
repens
harveyi (Mett.)
J. Sm.: U. Sen
anomalum Bedd.:
semicordata
—
unitus
Dryopteris filix-mas (L.) Schott:
-
Holttum: U. Sen K77
U.
P.
(Sw.)
Sen K125
K27;
Bolbitis
B.
Davalliaceae
(L);
—
P.
invisus
formosum Willd.:
Roth:
eriocaulis
Peranema
-
K105;
doideae:
sp.:
A.
U. Sen
(Roxb.)
uliginosa (Kunze) Ching:
Arachniodes obtusissima (Mett.) Ching:
B. De
K88;
s.n.; T.
Sw.: U. Sen
(Retz.)
truncatula
Ching:
cisa Cav.: U.
C.
K32;
Holttum: U. Sen
U.
A15;
Thouars: U. Sen K34
(Lam.)
Didymochlaena
deae:
De
Athyrium filix-femina (L.)
erum
s.n.; S.
T.
hemitomum Hieron.:
A.
Diplazium
in
K29;
nudatum
Holttum: U. Sen
Asplenioideae: Asplenium africanum
D6; A.falcatum
U. Sen
Holttum
K12; C.ferox
striatus
M. flaccida (Bl.) Ching:
s.n.;
Pseudophegopteris
-
Kel;
(Holttum)
Sen L5;
U.
Sphaerostephanos
-
U. Sen
(Bl.) Ching:
Sen L32
(C. Chr.)
Holttum
(C. Chr.) Ching:
baramensis
Aspleniaceae
DarK
Holttum:
S. penniger (Hook.)
T.
U.
tylodes (Kunze) Ching:
s.n.; B. De
C.
De
Holttum: Holttum
Pseudocyclosorus esquirolii (C. Chr.) Ching:
s.n.; P.
U. Sen
B.
Bedd.) Ching:
ex
Pronephrium
-
(Bl.) Ching:
Link:
-.Mesophlebion crassifolium (Bl.) Holttum:
s.n.
Pneumatopteris
Sen
1992
1,
extensus
(Wall,
ornata
Metathelypteris dayi (Bedd.)
-
s.n.; U.
truncata
Cyclosorus¹
-
No.
37,
gongylodes (Schkuhr)
Macrothelypteris
-
torresiana
T77.
s.n.; C.
Holttum
(Bl.) Ching:
VOL.
under cult.
Kew;
N.
A.
Chr.:
exal-
K59; N. falcata (Cav.) C.Chr.:
U. Sen
Lee
O.
UL6
(L)
-
neriiformis
Blechnaceae
latum
lidium
Salviniaceae
Azollaceae
—
—
Sadleria
B. De
ontogeny of
U. Sen
K24;
Psammiosorus
Desv.: U. Sen
S.
stomata in
O. articulata
paucivenius
K51;
pallida (Hook.
S05;
drummondii
Lindm.:
Salvinia
Azolla
-
and
&
tenuifolia (Desv.)
(Sw.)
Presl: U. Sen
C. Chr.: Last
B. orientale
Arn.)
Hill:
261
ferns
s.n.
L.: B. De
U. Sen
Moore: U.'Sen
K44
K31
K103;
(BM).
Ca4;
B.
punctu-
Stenochlaena
-
-
Woodwardia
Sen K43.
Marsilea
diphyllum
-
Bedd.:
U.
Dar37
brasiliense
Sen K45
(L.) Sm.:
—
Structure
Oleandra africana Bonap.:
palustris (Burm.)
Marsileaceae
B. De:
Cav.: B. De
Blechnum
—
Sw.: U.
radicans
&
A. Br.: U.
Sen
K91;
M. minuta L.:
U. Sen K92.
cucullata Roxb.: B. De Kln22.
pinnata
R. Br.: cult Bot.
Dept. Kalyani
Univ.
B. De Kln21
-
Regnel-

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