BLUMEA
27
175-201
(1981)
Sturcture and ontogeny of Stomata in
U.
Sen & E.
Polypodiaceae
Hennipman
Rijksherbarium,
Leiden
Summary
The
stomata
(Filicales)
their
occurring
as
investigated.
are
ontogeny investigated.
significance
for
on
the fronds
of the
sporophytes
A number of different stomatal
The
different types
of stomata
tracing phylogenetic relationships
in the
of
are
number of
large
a
types is
Polypodiaceae
discussed
in relation
Polypodiaceae following
to
their
cladistic
a
s.s.
and
recognised, (newly) described,
possible
analysis.
Introduction
The
ferns
basically
are
more
more
than
one
less
or
but
always
has
the
The
undergone
some
delimitationof
prevail
1973;
are
even
De la
many
of its
genera
cells,
its
due to
and Pichi
ignorance
our
with
has
of which
the
on
Sermolli,
They
are
composed
splitting
and
grammitoids
attained reasonable
not
on
and
a
and
species
the
of retention of
and
recognition
appear ill-defined. Disagreements
1977).
about the
sensu
al., 1975)
Many heterogenous
propriety
but also
family,
stalk
vicissitudes
loxogrammoids
family
differ
only
the
some
the
a
wealth of
and Christensen (1905).
it, yet
Crabbe et
and bilateral spores with
vast
about the position of the family in relation
Sota, 1973,
certainly
with
Polypodiaceae
by
distribution.
sporangia
taxonomic
many
not
members in
remaining
(i.e.
in
tropical
dipteroids, cheiropleuroids,
stability. Pteridologists
of the
family
(1902)
have meanwhile been removed from
taxonomic
wholly
less than four rows of
since the time of Diels
elements like
Polypodiaceae
transferred elsewhere
of exindusiate sori,
perispore.
distinct
the
taxa
almost
possession
generous diversities
especially
and
epiphytic
characterised by the
of
in
included
presently
1947, but excluding the
Copeland,
true
to
other ferns
These taxonomic
(Holttum,
disagreements
phylogenetic relationships
of this
family.
11
was,
therefore,
of different
available and
revision of the
structure
and
family.
*Permanent
address:
address:
Netherlands.
of the essential data on the
representatives
together
This paper
ontogeny
at
most
of the
brought
being investigated
**Present
felt that
plant parts
before
of
and ontogeny
the
way
of
serious
the
polypodioids
Leiden, Utrecht,
and
which
Kalyani.
Botany, Kalyani University, Kalyani, 741235,
Vakgroep Systematische
must
Plantkunde,
Heidelberglaan 2,
be
taxonomic
is the first of a series of reports
properties of
the Universities of
Department
opening
on stomata
of different
structure
of the genera and their allies
on
are
the
now
It includes
India.
3508 TC
Utrecht,
The
BLUMEA
176
apart from confirmation of
Claessens and Van
and ontogeny and
observations differ in many
1970a and
Cotthem,
1973),
of the
structure
stomatal types
of many
stomata
to
and
the
reconstruct
in the
structure
of
species. Further, parts
from those of previous authors
result, it appeared possible
a
1970a; Fryns-
the stomatal
and Van Cotthem,
1973; Fryns-Claessens
recognized
1981
I,
findings regarding
new
important respects
and Mickel and Lersten, 1967). As
stomatal types,
No.
previous reports (Van Cotthem,
some
Cotthem,
VOL. 27.
probable
1973; Pant,
describe
to
our
(Van
1965;
two new
of evolution of the
pathways
for the first time.
family,
ACKNOWLEDGEMENTS
It is with
him
to
Leiden
to
Rijksherbarium.
of Pure Research
Dr.
W. Jiilich
We would
with
that U.S. records his
pleasure
come
He would
for
(Z. W.O.)
like to express
and
of the
ontogenetic studies
his
financing
one
Except
a
few
plants
of taxa
Garden
under
where
cases
mainly
at
the
acetic acid:
glacial
overnight
peroxide
70%
the
at
and
alcohol and
cellulose acetic
For
AND
a
of
epidermal
hematoxylin
or
was
Leiden,
of
acetone
The
terminology
modifications. For
peels
were
Leitz
a
used
by
—
is
choosing Copeland's system
have,
Copeland
Drymoglossum
the
are
also due to
We
for
comments,
are
indebted
providing
help
the
to
material
for
herbarium specimens
to
and macerated
volumes of
were
stomatal
formalin: 5,
(40%
50% alcohol: 90),
equal
the
survey
fixed in F.A.P.A.
20% hydrogen
stained in Sudan IV in
Slides for references
surfaces
were
are
kept
in
also made with
1968).
development
(1947)
is used
alcohol.
were
Temporary
collected
acetocarmine
stained with Delafield's
Epidermal peels
also examined. Illustrations
were
but
followed
is due
(1970a, 1970b)
types of
to
by pteridologists
recognised
however,
though
made.
the
(1975)
used
used
various stages of
Van Cotthem
indicating
of classification —
being widely
of
were
drawing apparatus.
Crabbe
al.
facilities
climate chamber (phytotron) located in the Leiden Botanical
(1947) system
et
available,
epidermal
(Payne,
at
studies.
Mulder
were
2.5 and
Sudan IV in 70% alcohol
made with the aid of
the
for the Advancement
METHODS
Garden, fixed in F.A.P.A., and stored in 70%
mounts
disposal
phytotron.
glycerine-jelly.
Replicas
studies fronds
from plants grown in
the
these
acetic acid. These
glacial
Leiden.
sheets and
ontogenetic
in
Franklin's method in
mounted in
finally
in
boiled in water,
were
following
Rijksherbarium,
his
Organization
the work.
Mr. A.
to
material
preserved
and concentrated
also
cultivation
2.5, propionic acid:
58 °C
and interest in
investigated
Rijksherbarium,
types. Frond segments
at
year stay in the Netherlands. His thanks
MATERIAL
located
placing
sincere thanks to Mr. G. J. de Joncheere for his valuable
Botanical
from
for
like to thank the Netherlands
determination
Leiden
and
project
help, co-operation,
our
Professor Dr. C. Kalkman who made it possible for
gratitude to
this
on
particularly
for his various
nomenclature,
authorities
work
to
the
excluding
for
only
in
stomata
Phymatodes
indicated
as
that
a
as
genus
a
here with
taxa,
some
Copeland's
transferred elsewhere
purpose.
system
the world
over
adopted
various
taxa
practical
the fact that this
all
is
reason
of classification
despite
distinct
by
for
is still
many demerits. We
from
Microsorium
of
convenience.
same
level with the
purely
matter
The
instead of Pleropsis.
OBSERVATIONS
In
most
epidermis,
of the
species investigated
but in many
species
of
the
guard
Pyrrosia they
cells
are
are
in the
deeply
sunken in the
mesophyll
U.
Fig.
1.
Epidermis
in
Sen &
surface
E. Hennipman:
view
Stomata in
illustrating polocytic
Polypodiaceae
(a), copolocytic
cocyclocytic (d), cyclocytic (e), pericytic (f), copericytic (g), desmocytic (h)
Selligueafeei.
stomata. — a.
microphyllum.
—
—
h.
e.
b.
—
Colysis
membranacea.
Lemmaphyllum rostratum.
Platycerium quadridichotomum. —
i.
—
f.
—
c.
and
(b),
alcicorne.
—
—
g.
(All
anomocytic
codesmocytic (i)
Polypodiumfallax.
Pyrrosia flocculosa.
Platycerium
177
—
d.
(c),
types
of
Lemmaphyllum
Drymoglossum piloselloides.
x
750).
BLUMEA
VOL. 27,
No.
178
forming
of
the
external
an
with their
cells
The
epidermis.
found
are
fronds
in
stemaria and
The
superbum.
P.
oriented in the direction of the
discrete
areas,
become
become
they
the
of the
basis
number of
surrounding cells,
1.
Polocytic
surrounded
pore)
—
2.
cell
4.
—
the
Anomocytic type
cells
Desmocytic
region by
one or
distal
6.
7.
the
cells
9.
cells
not
—
of the
—
on
subsidiary
U-shaped
subsidiary
cell;
cell linked together towards
with
an
two
additional subsidiary
subsidiary
cells linked
differing
from other
1, b).
by
a
few cells
not
1, c).
encircled
completely
guard
cells
walls
by
(Fig.
subsidiary cell;
at
the
the distal
1, h).
desmocytic
two
a
both of them connected
or
of the
—
—
surrounded by
together by
of the
stoma
—
Cocyclocytic type
the
classified
cells and the encircled stomatal
polocytic type
(Fig.
stoma
stoma
linked
walls of the
at
guard
surrounded
stoma
—
apparently forming
least
are
subsidiary
type
with
an
cells linked
additional
together
at
the
1, i).
Cyclocytic type
at
some
rarely
been observed.
never
These
in
stomata
mode of orientation of the
recognisable,
anticlinal cell walls of the
Copericytic type
periclinal
8.
two
region (Fig.
stoma
one
type
Pericytic type
cells
the
The
holttumii,
distributed and
more
family.
anticlinal cell wall of the
anticlinal cell
two
region (Fig.
subsidiary
of the
stoma
first; the
—
type
Codesmocytic
subsidiary cell;
the
easily
cells and the
in form and size
subsidiary cell and any
5.
guard
towards the distal
epidermal
or
in them has
and
P.
1, a).
(Fig.
partly enclosing
3.
single,
a
Copolocytic type
together
(i.e.
stoma
by
anticlinal cell walls of the
the distal end
cells
cells
level
Only occasionally,
oriented. Two
in the
guard
general
hypostomatic.
are
generally evenly
are
recognised
subsidiary
species
Platycerium grande,
of the frond.
polar contiguity
here
the
outwards above the
follows:
as
type
axis
irregularly
are
the
stomata
long
but
laterally contiguous
Nine basic stomatal types
of
most
1981
species investigated
no
protrude
to
been found in
amphistomatic condition has only
P.
In
respiratory cavity.
neighbouring
1.
two
a
cells linked
surrounded by 3
cycle
of cells
cyclocytic
proximal
subsidiary cell; guard
pericytic type
subsidiary
stoma
a
any anticlinal cell walls
end
if
(Fig.
stoma
not
or
with
at
a
(Fig.
second
the distal
more
cells and the
1, f).
subsidiary cell;
region (Fig. 1, g).
elongated
neighbouring
1, e).
with
an
additional layer of
throughout (Fig.
1, d).
encircling
U.
abundant
Sen &
E.
Hennipman:
Stomaia
in
Polypodiaceae
179
cocyclo
more
type
stomat l
cyclo
=
+
+
coperi
present;
STOMAT
types
peri
stomal
MATURE
OF
TYPES
=
codesmo
(+
below
table
desmo
the
in
DIFERNT indicated infrequent):
OF
DISTRBUION
are
studied
taxa
anomo
+
+
+ +
+ + +
+
+
+
+ +
+
+
+ +
+
+
+
type
stomat l copolo
+ +
+
+
+ +
+
+ +
+
+ +
+ +
+
+ +
+
+ + +
+
+ +
+
+ +
+
+
+ +
+
difer nt (+)
=
in
found
stomat
types;
polo
+
+
+ +
+ +
+
+ + +
or
type
mature as ociated
of
types
other
the
The
than
Aglaompha mey nia splendens Anap usia decurens Arthomeris himalyens lehman i walichan Belvisa an mensi califolia henryi longis ma mucronat novguine s platyrhncosrevoluta spicata validnervis Campyloneurm angustifolum leuconeuron lindigi phylitdes Christoperis sagita tricuspis varians Colysis eliptica hemiontdea macrophyla me brance
dura
BLUMEA
VOL.
27, No.
180
cocyclo
cyclo
I,
1981
+
+
+
+
coperi
+
peri
+
codesmo
+
desmo
+
anomo
copolo
polo
+
+ +
+
+
+ +
+
+
+ +
+
+
+
+
+ +
+
+
+
+
+ +
+
-1-
+ +
+
+
+
+ +
+ +
+
+ +
+ +
+
+
+
+
+
+
+
+
+
+
+
+ +
+ +
+ + +
pedunclat poilaneipothifolia wrighti Crypsinus albido-squamts ebenipes quin efidrhuymncophylus stenophylus taeniatus triquetrus veitchi Dendroc he an bela Dendroglsa minor Diblem a samrensi Dictymia browni mcke i Drymoglsum piloseloides Drymotaenium miyoshianumDrynaria bonii laurenti parishipleurido es propinqua quercifolarigidula sparsi ora Drynaiops heraclea
U.
Sen & E.
Hennipman:
Stomata in
Polypodiaceae
+
+
+
181
+
+ +
+
+
+
+ + +
+
+
+
+
+
+
+ +
+ +
+
+
+
+
+
+
+ + +
+
+ +
+ +
+
+ + +
+
+
+
+
+ + +
+
+ +
+
+
+
+ +
+
+
+
+ +
+
+ +
+
costula m beguini bro ksi
var.
var.
Eschatogrma desvauxi furcata Goniphlebium demrsumkorthalsi persicfolium subaricltum truncao-sgitrum verucosm Gram toperidumcostula m costula m costula m Holcsorus bisulcatus Holstachyumbuchan i Lecanopteris carnosa lomariodes sinuosaLemaphylum carnosum drymogls idemicrophylum microphylummicrophylum obvatum rostraum Leptochilus axilaris Margin opsiwiesbauri Merinthosru drynariodesMicrogam hetrophyalycopdio espricarfolia piloseloides
var. var. var.
cil ata
182
BLUMEA
VOL.
27,
No.
1,
1981
cocyclo
cyclo
+
+
coperi
peri
codesmo
desmo
anomo
+
copolo
polo
+
+
+ +
+ +
+ +
+ +
+
+ +
+ +
+
+
+ +
+
+
+ +
+ +
+ +
+ +
+
+ +
+
+
+ +
+ +
+
+
+
+
+
+
squamulosa vacinfolia Micros ium hanco ki mebranceum musifolum pteropus punctaum sablniaum spectrum tenuilore Neochirptes
fo.
ensata ensata
+
+
+
+
monstrifea normalis palmtopedat subhast a americanum lanceolatum balteiformis longifolia carinat crasifola decumanum speciosa com uta
Niphdium Oleandropsi fer ea Paltonium Par gam a Pes opteris Phlebodium aureum Photin peris Phymatodes
U.
Sen &
E.
Hennipman:
+
+
+
+
Stomala
in
Polypodiaceae
183
+
+
+
+
+ +
+
+ +
+
+ + +
+
+ +
+
+
+ +
+ +
+
+ +
+ +
+ +
+
+
+ + +
+ +
+
+ +
+
+ +
+ +
+ +
+ +
+ + +
+ +
+ +
+
+
+ +
+
+
+
+
+ + +
+
+ +
+ + +
+ +
+
)
+
+ +
+ +
+ +
+
+
+
(
bifurcatum
ssp.
bifurcatum
bifurcatum
ssp.
diversfolium nigrescens scolpendria Platycerium alciorne andinum bifurcatum cornarium elphantois grandebifurcatum
var.
laurenti
ellisi
veitchi
ssp.
var.
holtumi
ssp.
madgscarien quadrichotmu stemaria stemaria superbum bifurcatum walichi wandae bifurcatum Pleopeltis abreviat astrolepis bicolor elongat excavta loriforme macroap schraderi scolpendriasinensi thunbergianPodsorus angustausPolypdioters brachypoda colrat
var.
hillii
wil ncki
ridleyi
BLUMEA
184
VOL.
27,
No.
1,
1981
cocyclo
cyclo
coperi
peri
codesmo
desmo
anomo
)
+
+ +
+
+
+ +
+
+
+
+
+
+
(
copolo
polo
+
+ +
+ +
+ +
+ + +
+
-1- +
+ + +
+
(+) + (+)
+ +
+
+
+ +
+ +
+ + +
+
+
+ +
+ + +
+
+ +
+ +
+ +
+ +
+
+
(+)
+ +
+ + +
+ +
+
+
+
+
+ +
+ +
+ + -1-
aPolypodium m ianum amoenum atkinsoni atenuatm australe californcum cathrinae chnophrum crystaloneur falcaria falax fauriei formosan fraternum fraxinfolum friedchstalnum giliesilachnopus lepidopteris loriceum ma eins nip onicum pelucidum plumulapoly dio es praini pringlei pycnoarpum scouleri triseriale virgnia um vulgarePseudorynai cornas Pycnol ma metacoelum murdens
U. Sen & E.
+
+
+
Hennipman:
+ +
+ + +
+ +
+ +
+ +
Stomala
in
185
Polypodiaceae
+
+ +
+ + +
+
+ + +
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
num larifo
sphaerosticha
+
+ +
+
+ +
+
+
+
+
+
+
+
+
+
rigidum Pyr osia species ab reviat angusta confluens dimorpha floc igera floc ulosa foveolat gral ahet ractis lanceolat lingua longifolia mollis princepsrupestris serpens heare i stictica stigmosa Saxiglos um taeniodes Seligueafeei lima bifrons brunei Synam ia feuil ei cornucopia
Thayeria
Solanpteris
+
+
+
Thylacopteris diaphana papilosaWeatherbya acedns damunesi
186
BLUMEA
ONTOGENY
The
cytoplasm.
always
on
leaf
a
do
considerable
In
to
stomata
cells
unequal
proximal
stomata
The
guard
cells
a
a
in the
guard
mother cell,
cells.
The cell
b).
ordinary epidermal
e).
Thus in
necessary
Fig.
2.
the
to
wall
a
stomata in
third
early
ellisii.
meristemoid
at
stage
of
maturation; f,
stoma in
a
subsidiary
and
stomata
during
divisions
preceding
two
to
It
the
crescent
a
the
the formation
proximal pole
the distal
to
cell
a
curved wall and form
the
nearer to
cell. The smaller cell has
becomes
soon
first, forming
a
more
a
less
or
pair of guard
and the
pole
divisions in
d,
two
of the
lateral sides of the
mature
stomata
are
(Fig. 2,
the stomatal meristemoid
arcuate
in
to
stages
of maturation.
(All
x
are
the distal wall of the
of development. a, stomatal
meristemoids after second
stomatal meristemoid after second
early stage
lenticular
the stomatal meristemoid
The first is
and
and form the
which is the sister cell of the
copolocytic stomata
divison; c
by
develop
Those destined
ones.
cell which is
shape
subsidiary cell,
stoma.
and
in
d).
independently of
Polocytic
after first
division; h, copolocytic
succession
copolocytic
larger daughter
right angles
c
adjacent
mature
—
cells
of
mothercell).
cells. All
cell
of
staining
period
species investigated
guard
in
appear
anticlinal division
characteristic
polocytic type
Platycerium
meristemoid;b,
cells
a
no
the main
(guard-cell
two
sequence
staining properties.
cells derived
produce
They
neighbouring
The
surrounds the
or
and denser
long before
In
from the
distinguished
nucleus,
cell precursor
of them form
space between them. Meanwhile the
guard-cell
be
easily
can
observed in different taxa is noted below.
undergo
and
(Fig. 2,
assume
STOMATA
completed.
The
growth.
stomata
dense
size
is
guard
differentiates as
leaf,
rounded, and divides by
equal
epidermis
majority
a
nucleus and
cells of almost
OF
considerable number of stomatal meristemoids
a
(Fig. 2,
end of the
prominent
leaf
mature
while
produce polocytic
two
TYPES
segments before forming
developments
ellisii
Platycerium
1981
appear
simultaneously.
of
period
of different types of
polocytic
two
arise
not
to
functions as the
one or
meristemoidsand
1.
size, prominent
begin
in the foliar
directly
off
cuts
No.
leaf are isodiametric, tetrahedral,
polygonal,
developing
meristemoids
activity
meristemoid
It
DIFFERENT
their smaller
by
These
meristematic
a
VOL. 27.
and uninucleate. These cells
cells
surrounding
a
cells in
protoderm
slightly elongate
or
OF
-
division; e, polocytic
division;
750).
g, meristemoid
after
U.
E.
Sen &
Hennipman:
stomatal meristemoid and the second is
Slomata in
Polypodiaceae
perpendicular
the
to
187
same
wall of the
guard-
cell mother cell.
The sequence of cell divisions in the stomatal meristemoids
types
of
stomata
polocytic
is
similar to that
basically
The
stomata.
difference is that in the
only
divisions in the stomatal meristemoids there
type the meristemoids of the copolocytic
by
copolocytic type
producing
instead of
three divisions. As in the
are
first divide into
stomata
two
guard-cell
mother cell
The first
f).
divisions
guard
two
cells
towards
cells
the
by
is very similar
to
the
Drynariopsis
to
As
h).
of
the
terms
cells
(Fig.
two
the first and second
in the
the
polocytic type,
cells
guard
of cell
the
as
initiate
produced
right angles
g and
region
unequal
two
cells
unequal
developed
are
this pattern
alignment
polocytic type.
heraclea and
Belvisia
follow the sequence of cell divisions as
developing polocytic
from the stomatal meristemoids
validinervis
the
stomatal meristemoids
followed by the stomatal meristemoids in
are
and
Platycerium ellisii, and form poloBut in many
at
wall
(Fig. 2,
distal
produces
meristemoids thus
a
of the stomatal meristemoids. In
independently
In
wall similar to the first,
while the third cell
produce
surrounding
a
derivatives of the
two
subsidiary cells,
by
two
polocytic
curved wall. The smaller daughter cell, instead of functioning directly
a
2,
forming copolocytic
in the meristemoids
occurring
copolocytic
undergoes
(Fig. 3,
stomata
of these taxa, the
stomata
one or more
a-e;
b, f-h).
cell derived
subsidiary
anticlinal divisions
(Fig. 3,
d, i, j; d, i, k). Following such divisions the newly formed cells adjust themselves in
such
a
that these
way
their
recognise
copolocytic
more
the
radial anticlinal walls
stomata no
longer
the First
taxa
(Fig. 3,
m
g,
it is
anomocytic and
and
(Fig. 3, j
nature
of these
stomata
become
stomata
polocytic
in
more
few
a
possible
to new
nature
to
developing
cell becomes divided by
subsidiary
—o). Due
retain their copolocytic
no
1). Interestingly
one or
orientation of these cells
and become
polocytic (Fig. 3,
o).
In
Platycerium
meristemoids
stoma
in
a
during
alcicorne
recapitulate
P.
quadridichotomum
of
the
its ontogeny, while others follow the stages of cell divisions
occurring
one
both
to
(Fig. 4,
the
categories
lenticular space appears in
distal wall of the
e
—g).
Some of these
exhibit
soon
developing
further elaborations. A
the inter-cellular material
cells and
developing guard
and
a —c
by
stomatal
polocytic
belonging
conspicuous
some
segmentations
followed
a
developing copolocytic
stomata
and
all the stages of
surrounding subsidiary
the
connecting
cell.
the
Finally
lenticular slit expands bringing about the separation of the wall of the guard cells
from that of the
point (Fig.
desmocytic
subsidiary
4d and h
(with
codesmocytic
—i).
proximal end; Fig. 4)
leaving
adjacent
subsidiary
one
stomata
cell
The
(with
a
link
cell
cell
thus formed,
are
at two
cells situated
proximal
the
at
almost
two
first
at
or
superimposed
derived
rate
failure in
The
end;
and
wall into
subsidiary cell,
guard-cell
by
Pyrrosia
a
sp.
large
type of
while the small
a
mother cell as usual divides
which
it is
stomata
small cell
daughter
separated
by
from the
cell
a
at one
from
4,
at
the
the stomatal
probably
due to the
cells and
subsidiary
and
d)
cells
their
readjustments.
(Hennipman 5122).
and
Fig.
subsidiary
independently
cells and the
guard
of mutual
ontogeny of the pericytic
curved
wall
of the
enlargement
maintaining harmony
piloselloides
a
of
later
the distal end of the
meristemoid. The cause of the appearance of the lenticular slit is
unequal
and
points
at
was
studied in
Drymoglossum
A stomatal meristemoid divides
(Fig. 4, j).
acts as a
straight
The
large
guard-cell
wall almost
subsidiary
cell
by
cell becomes
a
mother cell. The
perpendicular
(Fig. 4, k).
to
the
The stomatal
188
BLUMEA
Fig.
3.
Drynariopsis heraclea.
anomocytic (c —d, i—j,
n,
x
750; e,
h.
j, l,
o,
and
x
i,
240).
k
—
—l),
Pathways
and
VOL.
of
27. No.
I,
1981
development of polocytic (a —e), copolocytic (b,
‘pseudo’ polocytic (g, m—o) types
of stomata.
(a
—
d, f,
g,
f—h),
i,
k,
m,
U.
Fig.
4.
Pathways
copericytic (n —q)
5122).
—
n—q,
of
Sen &
E.
development
types
of stomata.
Hennipman: Stomata
of
—
desmocytic
(a
—
(a —c,
Pulypodiaceae
—d), codesmocytic
a—i, Platycerium
Drymoglossum piloselloides.
in
e,
alcicorne.
f, j, k, l,
—
n, o,
(e
189
—i), pericytic (j—m),
j—m, Pyrrosia
x
750;
d,
g,
h, i,
sp.
and
(Hennipman
m, p,
q,
x
240).
BLUMEA
190
meristemoids
structure
of
the
of these
VOL. 27, No.
-
1,
1981
upto this stage of development,
plants
and function the stomatal meristemoidsof the
developing
further
stomata
intercellular substance
elaboration is
the walls
connecting
towards the distal end of the leaf swells
of
arrested
stoma, and the
cell
subsidiary
development
and the
just by
short
a
this link
even
guard
appreciably and
cells became
guard
cells maintain their
a
while in
instead of
These meristemoids divide
n).
by
The distal cell follows the
into
The
cells
the
two
material
intercellular
stomata
their distal
periclinal
stomatal
the
During
smaller
cell
the
flanking
position together
surrounding
subsidiary
cells
Sometimes the
fail
to
cells
as
region
so
end is
are
of
of the
a
to
subsidiary
and
ontogeny,
ultimately
stoma are
cells
a
ones.
(Fig. 4,
mother cell
subsidiary
lenticular slit in
cells
lost
and the inner
in
some
of the
maintain
always
cell
cells
division
to
are
by
a
and
a
cell
large
a
longitudinal
these
e). Occasionally
cells
the distal
at
form the
proximal position
other
the
adjoining epidermal
undergo
adjoining
the
to
follow
the
surrounding cells,
radial wall
5, g and
(Fig.
h).
the distal end of the stomatal meristemoid
at
or
after the initiation of the
undergo division, they elongate
surround the distal
cases
of
d and
These
occupying
preceding, during,
previous
pair
cell
stomata.
divide into
—c). They
mother
(Fig. 5,
(Fig. 5, f).
undergoes
their failure
a
sinuosa,
cyclocytic
meristemoids
Meanwhile the
region
of the
thus of dual
guard
origin.
cells. The
The
one
guard-cell
and
readjust
surrounding
adjacent
derivative of the stomatal meristemoid while those
at
the
to
the distal
independent origin.
Some of the stomatal meristemoids in
sinuosa
Drymoglossum
Lecanopteris
form
stomatal meristemoid
cells. Similar
guard
two
these
guard-cell
curved walls
neighbouring
as to
them,
(Fig. 4, m).
become
cells
carnosa,
(Fig. 5,
stomata
cell.
radial walls
with the
despite
in all the
proximal
by
division
undergo
guard
gradually
microphyllum
as
of the
by straight
two
mother cell. But
considerably
functioning
itself often
cell
cell
guard-cell
a
other,
subsidiary
two
development,
subsidiary
a
cells become divided
as
the
of the
Lecanopteris
polocytic
of
distal end
anticlinal division
in
of
stages
into
differentiating
walls
But the
Lemmaphyllum
pathway
daughter
stage of
attachment.
and
early
developmental
behaves
between them is
—q).
meristemoids
brownii
Dictymia
o
Final
unequal
and increase in size of
the
connecting
(Fig. 4,
developing
The
appearance
the anticlinal link
subsidiary cell,
cells
the
(Fig. 4, o). Following
the
sp. and
two
in
occurs
type of development and forms another
produced
partly enclosing
one
that
produce copericytic
stomata
the
cell
with the wall of the
subsidiary
a
(i.e. Pyrrosia
taxa
pericytic
copolocytic
larger cells,
others
subsidiary
events
In
1).
curved anticlinal wall into
a
cell. The smaller of the three cells thus
(Fig. 3, n).
contact
by
in
some
lenticular space appears
a
(Fig. 4,
surrounded
completely
developing
In
short anticlinal wall is lost in many of
A few stomatal meristemoids in these
piloselloides)
wall
connecting
through
stomata.
cells and the
guard
within the intercellularmaterial. Then follows the sequence of
desmocytic
resemble both
polocytic
Lemmaphyllum
adopt
form
copolocytic
cocyclocytic
shown in
Fig. 6,
a
development,
stomata.
—
f.
subsidiary
cell
increasing
the number of cells
(Fig. 6)
may
Dictymia brownii, Lecanopteris
microphyllum,
produce
The stages of
Occasionally
undergo
instead
division
encircling
the
of
following
two
subsidiary
development
either the inner (Fig.
by
one
guard
or more
cells
of
carnosa,
the
a
L.
polocytic
and
cells,
cocyclocytic
6, g)
or
the
outer
radial periclinal walls
(Fig. 6,
g —i,
and j
—
1).
U. Sen
Fig.
5.
&
E.
Hennipman:
Lemmaphyllum microphyllum. —
meristemoid;
b, meristemoid
encircled
subsidiary
by
subsidiary
cell
with
cell
after first
at
the
septation. (All
Though
the adult
of them pass
stomata
through
development.
The
development
in
been
cell
side
and
three
to
division
stages
of
development. a,
after second
surrounding
in the
the
AND
cells
division;
at
the
d
Stomatal
—f, guard
distal
region;
cells
g—h.
CONCLUSIONS
polypodioids
polocytic
exhibit nine sharply defined forms,
conditions
polocytic type quite curiously
development, leading
has
c, meristemoid
191
Polypodiaceae
750).
DISCUSSION
all
in
stomata in
Cyclocytic
division;
proximal
x
Slomata
different
directions
(Fig.
during
follows
7).
the formation of cyclocytic and
and/or
change
in
In
an
the
course
identical
one
of
cocyclocytic
orientation in
the
of
their
pattern of
these
lines
of
stomata, there
neighbouring
cells
192
BLUMEA
VOL.
27,
No.
I.
1981
U. Sen &
Fig.
7.
Different
types
of
E.
Hennipman: Stomata in
polypodioid
stomata
in
stages
193
Poiypodiaceae
of
development showing ontogenetic
interrelationships.
Fig.
x
240).
6.
Dictymia
brownii.
—Cocyclocytic
stomata in
stages
of
development, (a —g, j, k,
x
750;
h,
i, l,
194
VOL.
BLUMEA
occupying
the
by
the distal region of the
formation
desmo-,
developing
guard
the
cells
and
capacity
The second line is characterised
and
peri-
cell.
This
forms.
copericytic
breakdown the
to
subsidiary
the
1981
1,
stoma.
codesmocytic,
feature of this line is the
conspicuous
between
of
No.
27,
is
due
probably
A
wall link
periclinal
the
to
differential rate of expansion
of the guard cells and the associated subsidiary cell
cells
development. Polocytic
the late stage of their
at
fronds either arise
further
such
of
directly
differentiate,
juvenile
a
fail
eventually
or
retain
to
type of
developing
polocytic
ones
as
cell
their
Juvenile
stoma.
the
by
from
cells
copolocytic
resulting cells
cells.
epidermal
division of the
An
cell of
subsidiary
transformed
are
(i.e. due
phenomenon
same
other
the
not
cell
subsidiary
outer
and
division,
distinctiveness
result of the
a
undergoes
is formed
stoma
cell. The
or
mature
meristemoid cell which does
polocytic
'copolocytic'juvenile
copolocytic
anomocytic
polocytic
from a juvenile
from a
in
occurring
stomata
a
into
the division of the
to
subsidiary cell).
the cell
Though
are
of dual
adjacent
the
to
proximal
the
and
mesoperigenously
guard
peri-
becomes
distal
until the
so
that in all these forms they
Pant, 1965). The surrounding cells
stoma
the
at
significant
copericytic types
appears
cell is lost. It is
cells
and
see
of nine different forms vary
stomata
it is
of the
region
while those situated
Even in
subsidiary
mature
orientation,
or
origin (i.e. mesoperigenous,
meristemoid,
from it.
the
surrounding
in aspect and
considerably,
region
the
last
enclosed
by
the
derived
are
stomatal
meristemoid. The mesogenous condition in the adult stage is,
This
one.
thus fails
study
codesmocytic,
to
and
pericytic
copericytic
Cotthem
and Lersten
that in
the
(1967)
cell
surrounding
of the
developmental pathways
different from the
and
Van
the
be
not
guard-cell
and
cyclo-
'cyclomesoperigenous'
are
indeed
unique
and
The results of
contribution
to
studies
our
establish
on stomata
phylogenetic
are
Dr.
J. Th.
idea
set
Wiebes,
forth
observations
Platycerium (Hennipman
applied
to
In
by Hennig (1966).
establish
and
Roos,
phylogenetic
on
a
in
by Fryns-
and Mickel
attached
we
found
to
the
be very
of stomata in
and have
within the
never
the
been
approach
of his
to trace
colleagues
will be
their
to
possible
Polypodiaceae.
in
preparation) Hennig's
within the
Leiden,
genus,
are
Prof.
to
of the fern
ideas
As
phylogenetic
analysed according
forthcoming monograph
relationships
desmocytic,
plants.
to one
stomata
secundary
a
never
discussed in relation
relationships
several years
due
ago,
our
of
stomatal
stomata to
types
development
the second author became interested in the cladistic
relationships already
of
the
pair
diagrammed by Fryns-Claessen
cocyclocytic
recorded earlier in any other group of vascular
(1965)
Moreover,
cocyclocytic types
stomata
in their mode of
of
been claimed
mother cell is
confirmed either.
Cotthem (1973). The cyclo-
polypodioids
has
as
The observations of Pant
(1973).
Pyrrosia
could
stomata
origin
begins
with
that the
therefore,
confirm the view of mesogenous
Claessens and Van
link
cell derived from the
subsidiary
cell
independently
wall
contact
or
stomatal
development
anticlinal
after the loss of anticlinal wall
only
of the
derivatives
are
the
genus
succesfully
using
a
large
number of characters.
According
to
the
apomorphic (derived)
cladists,
monophyletic
character
states
(primitive) character states
mainly
applied
interested in his
to
are
in
zoological
theory (e.g.
The establishment of the
groups
only. Groups
can
based
on
be
based
shared
on
principle heterogeneous. Hennig's ideas
groups.
Bremer and
most
Only
recently
Wanntorp,
likely apomorphic
botanists
shared
plesiomorphic
havebeen
have
become
1978).
character
state
of stomatal
U.
is
types
looking
stomata, it
and
the
chart
be
can
the
apomorphic
conditions.
the
and
Cyclo-
the
and
desmo-
cladistic
stomata,
codesmocytic
differentiated from the
easily
pericytic
stomata
never
these
well
as
types,
and
polo-
types of
the
as
peritwo
represent
copolocytic
which
together
occur
When
methodology.
relations between
cocyclocytic
and
195
Polypodiaceae
the
ontogenetic
cyclo-
allied
states
of
application
that
supposed
Stomata in
Hennipman:
depicting
and
copericytic
different
for
prerequisite
a
at
E.
Sen &
in
itself
indicates their different evolution.
The
condition.
Polocytic
stomata
stomata
Grammitidaceae,
The second
of
possibility
family
retention. This
seems
since such
unlikely
It should be remarked that the
stoma
cell division and orientation makes it difficult
The
following
recognised,
stomata
or
These
has
with
synonymous
shown
requestioned
by
and
(1974)
that
the
The
Pyrrosia.
between
relationship
regarded
in different
stomatal
of
monophyletic
as
of
polocytic
polocytic.
occurrence
is
as
well
and
Pyrrosia
true
as
for
types
copolocytic
groups
unless
Saxiglossum
and
other features
(1940)
Ravensberg
(syn.:
Saxiglossum.
Polypodiaceae.
have in part
a
a
different
is
The stomatal types
relationship
accommodated
perispore (Hennipman
to
has been
Platycerium
On the other hand, the spores of
strikingly
(in
Pteropsis)
(e.g. stellate hairs) point
Such
Platycerium.
Ching
and
Hennipman
Drymoglossum
genus
same
several authors.
groups
is twofold.
stoma
separate family Platyceriaceae, while Pichi Sermolli (1977) placed these
Platycerium
or
through
polocytic again through
apomorphic
Platycerium, Pyrrosia,
Price
by
confirm this. The stomatal types
a
be
pericytic
stoma arose
place.
Drymoglossum,
research
preparation)
can
either the
'pseudo'
or
of the
one
Polypodiaceae
ascertain whether the
to
classified because of the
groups
to
polocytic
become
polocytic
share
genera
primarily
has taken
The genera
Recent
of
groups
are
only.
parallellism
1.
the adult stage is 'real'
found in
state as
in the
radiation.
of the
may
of
transformation series would
a
subsequent
interpretation
originally copolocytic
an
condition refers
appearance
plesiomorphic
a
likely sistergroup.
most
polo- and copolocytic
have involved retention combined with
The fact that
from
only
not
the
regard
we
plesiomorphic
however
the combined
expected
commonly
very
occur
a
be
could
as
stomata, from which the
cyclocytic
ferns shows
polypodiaceous
copolocytic
but also in
the
of the
plurality
and
polo-
two
in
a
genera
Pyrrosia
and
and Sen, in
preparation).
2.
The
genera
Dictymia,
Lemmaphyllum,
According
into
three
to
Dicranoglossum
Niphidium,
slight
different
Eschatogramma),
which
are
mutually
obviously
Lecanopteris,
(syn.: Neurodium)
differences in stomatal types this group
subgroups
is concerned,
present knowledge
(syn.:
and Paltonium
related
three
representing
be subdivided
can
closely
not
as
different
far
as
parallel
developments.
2a.
Dicranoglossum
The
from
stomata
those
cells
surrounding
suggests
2b.
The
It is
species
as
found in
found in
that
towards
cyclo-
genera
and
Dicranoglossum
other
the
groups
distal
cocyclocytic
stomata
and
the stomatal
types of
polypodioid
sori
as
listed
(syn.: Eschatogramma)
showing
region.
Dictymia, Lecanopteris
striking that
with
the
an
The
are
occurrence
liable
to
are
different
orientation
incomplete
of
this
parallel
of
condition
evolution.
Lemmaphyllum
Lemmaphyllum
by Copeland (1947),
and those of
have
a
Pleopeltis
derived
type
of
-
BLUMEA
VOL.
stoma.
Van Cotthem
Weatherbya,
In
have
stomata
a
1981
I,
found that
(1970a)
some
of the
polocytic
appearance because of an additionalpericlinal division of
cyclocytic
s.l. is
Lemmaphyllum
the cells.
No.
27,
196
under
presently
in
study
our
by
group
Mr.
J.
van
Scheepen.
The data further
the idea to
support
including
Myrmecopteris
genera separate
because
Sermolli,
Holttum
1977).
Pichi
Phymatodes.
of salient
Sermolli
in
recognize
Walker,
large
a
differences in
relates
rhizome
Solanopteris
other
on
Dictymia, Lecanopteris s.l.,
genus
Other
1975).
accommodated the
(1954)
confirmed. Further research
similarity
and
(Jermy
these
indument
(e.g.
Pichi
of
Myrmecopteris
whether the
is due
Lemmaphyllum
is
which
Lecanopteris,
features may show
and
keep
species
to
thus
Lecanopteris,
authors
to
in
not
stomatal
monophyly
or
parallelly.
2c.
and Paltonium
Niphidium
of the
spite
In
of exclusively
occurrence
cyclocytic
Paltonium, the relationship between these genera is
structural
similarity
3.
differences.
in stomatal feature is
case
stoma
or
Niphidium and
ascertain
to
necessary
homoplastic
both
some
other
whether
this
otherwise.
of
in
types
copolytic
the
stomata
remaining
is the rule in
derived. The
can
occur
singly.
stoma
polocytic
The
as
anomocytic
well
stoma
as
stomata
found in
are
two
we
but
and
Christiopteris,
bisulcatus, Holostachyum buchananii,
and
Synammia feuilli. Polocytic
anomocytic
stomata
occur
leuconeuron, Christiopteris
Drynaria,
Drynariopsis
heraclea,
Microgramma, Pleopeltis
and the Solanopteris
In the genera
in
tricuspis,
five
stomata
species
species
two
Goniophlebium
excavata, fourteen
species
Belvisia,
species
with
of
copolo-
Campyloneuron
three
Crypsinus,
demersum,
of
three
species
two
two
in association
of
of
of
type of
one.
Microgramma, Paragramma balteiformis, Pleopeltis scolopendria,
Pycnoloma,
type
one
in association
occurs
of
the
studied.
taxa
copolocytic
copolocytic
species
that
see
of the
having
the
always
and sometimes also with the
Holcosorus
Crypsinus,
polocytic
genera
most
ferns
plesiomorphic condition, polypodioid
polocytic
Exclusively
species
stomatal
and
necessarily
are
with the
A.
polo-
this is the
stomata
the
checking
combination of
of
is
in
obvious, due to
The microsorioid ferns
When
In
research
Future
stomata
not
Polypodium,
species
of
of
species
two
Selliguea feei,
species investigated.
Christiopteris
and
Solanopteris copolocytic
found
stomata were
to
be absent.
Species
B.
with
copolocytic
stomata
species investigated), Dendroglossa
Neocheiropteris subhastata,
Other
stomata.
the
of
species
Thus,
stomata
processes,
together
species
anomocytic
and
genera
are
found in
Colysis (six
(syn.: Leptochilus
Podosorus
have the
minus;
with
the formation of
with the
both
stomata,
genera
polo-
e.g.
and
closely
and
polo-
polocytic
related
copolocytic
Leptochilus,
out
of
eight
Colysis minor),
angustatus (syn.: Polypodium prainii).
combination of
the genera share the condition that with increased
forming
These genera,
having
these
only
minor
to
copolocytic
specialisation
stomata
is
them for other
stomata
and
Paraleptochilus,
of
suppressed.
reasons,
never
Diblemma,
having
and
Microsorium, should be treated together.
It
is
tempting
primitive
stomata
ferns,
to
discuss
and derived
associated
states.
with
these
The
copolo-
the genera related to
two
taxa
and
Crypsinus,
different types
with
polocytic
anomocytic
and
a
of
stomata
stomata
stomata
only
or
in relation
with
include the
number of genera whose
to
polocytic
drynarioid
relationship
is
U.
Sen &
E. Hennipman:
Stomala in
otherwise still obscure; the genera assembled in group A
a
natural group. This is in
the genera of group A
in
This
family.
Grammitidiaceae,
anomocytic
the
stomata
with
association
with copolocytic
that the
anomocytic
stomata.
In this connection
in
the
stomata
or
types of
should like
could
Polypodiaceae
from
Loxogrammaceae
and
polo-
stomata
the
are
that
the
in
the
polo-
and
the
The fact that
more
occur
in
stomata, but
copolocytic
second argument. It is further
a
never
in association with
occur
mention that the observation of Van Cotthem
to
anomocytic
used the stomatal features in
(1977),
that
Polypodiaceae,
1970a).
be confirmed. Van
not
fact
and group 2 sometimes
1
with
the
which
process
condition in
original
by
the
alone may represent
specialised
we
of
group
in group
stomata,
the
represent
(Van Cotthem,
listed
constitute
not
forming
stomata
supported
that the combination of polocytic and
Pichi Sermolli
and
sister
rule
as
polocytic
noteworthy
(1970a)
the
are
should
strongly
possible
stomatal types
specialised
never
is
hypothesis
true, the
being
do
obviously
with the genera of group B,
contrast
microsorioid. This
unmistakingly
occurs
sharp
197
Polypodiaceae
does
not occur
followed
(1970a)
by
distinguishing Grammitidaceae
which
Polypodiaceae,
stomata
Cotthem
could
be
not
confirmed
either.
In
the
conclusion,
Polypodiaceae
,
within the
If
suggested
Polypodiaceae
Pichi Sermolli
a
are
a
selection of
of
number
a
of
taxa
polypodioid
possible natural
in part congruent with the groups
recognized
authors
(1975)
with the subfamilies listed
three subfamilies
are
publication
including
Crabbe
et
al.
and
consisting
Of the
9 groups, the groups
(which
unambiguously
are
here,
out
et
al.
(1975),
our
Platycerioideae,
If
groups.
of 63 genera
four
our
groups
observation
(1977)
a
data
the
suggest
remaining
genus
Van Cotthem
(Van
by Pyrrosia, Drymoglossum,
and the
Pichi
by
Pichi
by
by
subdivided into
these is the
genus of doubtfulreliance.
compare the
recognized
recognized
as
opinion)
we
are
only. Amongst
represented
in
congeneric
Polypodiaceae
genus
Pichi Sermolli
monophyletic
which contain 46
one
an erroneous
is ranked
by
the
(1977)
of
Cotthem, 1970a)
remaining
and
markedly heterogeneous.
because of
Christiopteris which,
Crabbe
by
Drynarioideae
of Pichi Sermolli
14 groups, five of which
suggested
of
number of
large
a
(1977).
compared
Saxiglossum
in
types
recognition
only
recently by
that apart from the subfamilies
In the
stomata
study of the ontogeny of
evidence for the
species, gives strong
groups. The groups
of the
analysis
together with
and
drynarioid ferns,
remaining
Sermolli,
Sermolli
seven
are
groups
with the groups
as
likely
be
are
to
heterogeneous.
It is
indeed
gratifying
stomatal features in
the
perspective
towards
of the
solving
to
be able
determining
to
some
point
methodology applied.
many
a
phylogenetic
the
to
phylogenetic
Of
problem
undoubted usefulness of the
affinities in these
course,
in the
plants,
this study is only
a
and
first step
family.
REFERENCES
BREMER, K.,
CHING,
& H.
R. C.
CHRISTENSEN,
COPELAND,
E.
WANNTORP.
1978.
Phylogenetic systematics
1940. On the natural classification of the
C.
B.
1905. Index Filicum.
1947. Genera
in
Botany.
Taxon
27
317-329.
(4):
family ‘Polypodiaceae’.Sunyatsenia
5: 201
—
268.
Copenhagen, Hagerup.
Filicum,
the
genera
of ferns.
Waltham,
Mass.: Chronica
Botanica
Co
-
BLUMEA
198
COTTHEM, W.
Jard.
VAN.
Bot.
1970b.
classification
A
classification
CRABBE, J.
A.,
L.
1902.
139
Rev.
E. & W.
39
1966.
W.
HENNIG,
141
in the
Bull
Filicopsida.
Bot. J.
In
A.
Linn.
C.
Soc. 63:
Jermy
235
al.
et
—
246.
(Ed.)
The
—
and
phylogeny
71.
—
and J. T. MICKEL.
3):
&
stomata
of the
1975. A
new
generic sequence
for pteridophyte herbarium
162.
-
In A.
K.
&
Engler
Prantl
Die
(Eds.).
natiirlichen
Pflanzenfamilien
Leipzig, Engelmann.
FRYNS-CLAESSENS,
Bot.
(2
types.
systematics.
59
Polypodiaceae.
339.
—
of stomatal
and
types
JERMY,
11
!, 1981
81 -239.
(2):
of the ferns:
A. C.
Fern Gaz
DIELS,
40
Belg.
Stomatal
1973.
27, No.
Comparative morphological study
1970a.
Nat.
VOL.
(1):
VAN COTTHEM.
1973. A
new classification
of the
of stomata
ontogenetictypes
71 -138.
Phylogenetic Systematics. University
of Illinois
Press, Urbana, Chicago
&
London:
1-261.
E. & W. A. RAVENSBERG. The
HENNIPMAN,
species
of
Pyrrosia formerly
referred
to
Drymoglossum
and
Saxiglossum. (In preparation).
& M. C.
Roos.
& T. SEN. The
HOLTTUM,
-
A
monograph
of
spores
R. E. 1954. A revised
the
Posing
1973.
ferns:
1
flora of
problems.
(2
MICKEL,
J.
T. &
D. D.
N.
R.
1965. On the
1:
M. G.
R.
31
(2):
Lecanopterisspinosa
Bot.
1967.
54(9):
and classification
phylogeny
of the
—
A new ant fern from Indonesia.
Fern Gaz
ontogeny
Floating
(adetostomy)
stomates
in
ferns:
distribution
and
1181-1185.
of stomata and other
The
1977. Tentamen
homologous
production
Pteridophytorum
fern
new
names.
Plant Science
structures.
Series,
of epidermal casts. Ward's Bull. 7
in taxonomicum
genera
Kalikasan, Philipp.
1973. On the classification and
phylogeny
and classification
List
Aglaomorpha meyeniana Schott (Ramos
of
phylogeny
J.
species
BS13649,
Biol.
of the
of the ferns: 229
ordinem
redigendi
3:
173—178.
In A. C.
Polypodiaceae.
—
Jermy
etal
244.
studied
B.
L)
spicata (L.f.)
Mirbel
Copel.
ex
( Wild
6571,
M)
A. splendens;(J. Sm.)Copel. (Hoogland6257, L)
decurrens
Anapausia
Presl
(Bl.)
3836a, L; Phengkhtai 37, L,
Artromeris
700,
T. Shimizu
et al.
validinervis
5541,
T
himalayense (Hook.) Ching (Gammie
Bot.
(Mett.) Ching (Dickason 8104, L)
A. wallichiana
(Spring) Ching (Mann
s.n.,
L)
callifolia (Christ) Copel. (Poilane 12210,
L;
Kostermans
B.
dura
(Copel.) Copel. (St.
15621
, UC)
s.n.,
12.1.1969, L)
John
8979, L;
Maireau
B. mucronata
(
Hennipman
Edano PNH
Halschbach
Fee
(Pringle
10257, L)
Fee ( Buchtien
s.n.,
L
9137331)
lindigii (Mett.) Ching (Schlim 440,, L)
C. phyllitides
(L.)
Presl
(Van Lansbergen 65, L)
Christiopteris sagitta (Christ) Copel. (Copeland
C.
P)
tricuspis (Flook.)
Christ
(Hennipman 3678,
L)
(Hieron.
longissima
C.
s.n.,■
<£
ex
C.
Chr.)
Raymond
(Hennipman 3376, L)
Holttum
C. varians
Colysis
(Brass 30362, L)
(Mett.) Copel. (Franc 686, L)
elliptica
(Thunb.)
(Fee) Copel. (Kjellberg 1422, L)
C. hemionitidea
(Wall,
ex
novoguineensis (Ros.) Copel. (Brass 30462,
L)
C.
B.
platyrhynchos (Kunze) Copel. (McClure BS
C. membranacea
(Bl.)
C.
(Hook,
16058, C)
(Bl.) Copel. (Holslvoogd
226,
Ching
(Hennipman
3391, L)
B.
B. revoluta
Copel.
Garden;
860, L)
C. leuconeuron
L)
henryi
(Kunze)
cult. Leiden
Campyloneurum angustifolium (Sw.)
A. lehmanni
Belvisia
B.
(Hennipman
11386, L)
B.
The
313-512.
1974. Nine
DE LA.
(Eds.).
B.
Government Printer.
Malaya. Singapore:
6-7.
Webbia
E.
of
(Eds.).
1-24.
PICHI SERMOLLI, R. E. G.
SOTA,
et al.
W. W. 1968. The use of cellulose acetate film for the
(52):
PRICE,
LERSTEN.
Amer. J.
Allahabad
PAYNE,
II. Ferns
Jermy
165-176.
3)
ontogeny.
PANT,
Platycerium (Polypodiaceae). (In preparation).
10.
—
&
genus
Malaya
In A. C.
JERMY, A. C. & T. G. WALKER. 1975.
11
of the fern
Polypodiaceae. (In preparation).
L)
Mett.)
Presl
(Tagawa
1938, L)
macrophylla (Bl.)
pedunculata
Presl
(Lorzing 12370, L)
Presl
(Hennipman 3881, L)
(Merrill 10493, L)
et
Grev.)
Ching
U. Sen &
C.
poilanei C.
C.
pothifolia (Don)
C.
Chr.
& Tard.
Presl
E. Hennipman:
( Poilane 32103, P)
G.
al.
5451,
C.
612
(Meijer
,
L)
rhynchophyllus (Hook.) Copel. ( Brass 31635,
Chr.
C.
Holttum
stenophyllus (Bl.)
(S w.) Copel.
(
Meijer 2471,. L)
palmatus (Bl.)
var.
C.
(Forster 203, L)
Wilde & de
Lecanopteris
veitchii
(Maire
L.
206.
L
L.
(King
Diblemma
minor (Fee)
BS
Copel. (Brass 25074,
L;
L.
20400, L)
samarensis
J.
(Edano
BS
brownii
D. mckeei
Tindale
NSW
8009, L)
Presl
Drynaria
s.n.,
L
Hieron.
Bedd.
pleuridioides
D.
propinqua (Wall.)
Presl
Presl
obovatum
var.
(Harr.)
(Bedd.) Tagawa (Rosenstock
de
Fit.
172, MICH;
Mann
Wilde
Wilde-Duyfjes
&
de
s.n.,
L
quercifolia (L.)
Sm.
D.
rigidula (Sw.)
D.
sparsisora (Desv.)
Bedd.
(Larsen
Mett. J.
et al.
Smith
L; cult.
Leiden
Bot.
desvauxii
Eschatogramma
squamulosa
Smitinand
(Rulten
Garden)
C. Chr.
ined.
(Petersen
&
&
Fisch.)
Copel.
908289452, L)
(Baker) Ching (Hansen
&
12806, L)
(Don) Ching ( Tagawa 3721,
L)
M.
musifolium (Bl.) Copel. (Jacobs 5070, L)
(Bl.) Copel. (Hennipman 3955, L)
pteropus
M. punctatum
(Terpslra
6517, L)
(Langsd.
L
M. membranaceum
Ching
Ule
(Kaulf.)
vaccinifolia
(Anonymous,
(Hennipman 3949,
(K.I.)
(Broadway
Hjerting 659, L)
M.
1963,
Presl
piloselloides (L.) Copel. (Hatschbach 17178,
L; Herzog 2153, L;
M.
(Franc 33, L)
(Kunze)
3256,
persicariifolia (Schrader)
(Henry
L)
heraclea
al.
lycopodioides (L.) Copel. (Columb 152, L)
3025, L)
(Lam 3386, L)
Moore
(Burchell
M.
M.
13092, L)
Drynariopsis
(Willd.) Copel.
M.
(Leeuwenberg
(Lorzing 15268, L)
ex
ciliata
Microsorium hancockii
D.
(L.) Copel. (Hennipman 3065, L)
M. sablanianum
35, L)
(Christ) Copel. (Brooke
BAU
9869, L)
(
Wine s.n., L 936224372)
demersum
C. Chr.
(Brause) Copel. (Croft
E. furcata (L.)
Goniophlebium
LAE
60621, L;
korthalsii
(Mett.)
908294531;
Schodde
Bedd.
Rosenstock
persicifolium (Desv.)
G. subauriculatum
3449,
s.n.,
L
Presl
(Hennipman 3363,
Sm.) Copel. (Iwatsuki
N.
L
(Hook.)
& Lelean
J. Sm.
(Pulle
LAE 65608,
(Brass 7672, L;
L;
Grammatopteridum costulatum (Ces.)
(Lam 826, L)
Lam 708,
C. Chr.
et
al.
ensata
(Thunb.) Ching (Oldham,
ensata
(Thunb.)
Ching
fo.
monstrifera
Iwatsuki
3635,
L)
N.
L)
var.
M. F.
908300530)
(Tagawa) Tagawa ( Tagawa &
ined.
&
(fixed material, L))
N. normalis
verrucosum
(J.
Neocheiropteris
L)
1126, L)
costulatum
tenuilore
s.n.,
(Bl.)
L.
1169, L)
1200, L; Hennipman 5618 (fixed material, L);
5619
(Elmer 11631, L;
(Kaulf.) Copel. (H.
spectrum
Lyon
14722, L)
Bedd.
truncato-sagittatum (Brause)
Croft
M.
M.
1438, L)
(Korthals
Surbeck 720, L)
G.
Garden;
heterophylla (L.) Wherry (Smith et
M.
D.
G.
Bot.
2402, L)
(Larsen 10634, L)
(Christ)
parishii (Bedd.)
L;
Leiden
L)
961244621)
bonii Christ
laurentii
Makino
(Mak.)
miyoshianum
Microgramma
M.
3088, L)
G.
(Forrest
33, G; Tagawa 8612, UC)
exsicc.
90828829;
Leiden
(cult.
Garden)
(Umemura
et al.
Presl
908338568)
9779, L)
(Lam 7219, L)
piloselloides
Drymotaenium
G.
(Meijer 2511, L)
(Wall.)
17428, W)
rostratum
15133,
Leiden
(Wikstr.) Copel. (cult.
Constable
Garden;
Drymoglossum
D.
L
Baker
13702, L)
Bot.
D.
Leiden
(cult.
Warburg 10937, B)
formos.
Sm.
s.n.,
microphyllum
L.
BO)
Dictymia
Bot.
(Brause)
Mann
Maire
Presl (
Ramos
Bl.
(Reinw.)
microphyllum (cult.
BO)
Dendroglossa
M.
(Forbes) Copel.
(Brass
(Copel.) Ching (Floyd
carnosum
Gaudichaud
annabellae
Copel.
drymoglossoides (Bak.) Ching (Caxalerie627,
91842127)
Dendroconche
carnosa
L. lomarioides
Wilde-
s.n.,
(Hook.)
Garden)
L;
Copel.
(Bak.)
brooksii
var.
(Brooks 361, S., BO)
buchananii
Lemmuphyllum
Duyfjes 15281,
■ L)
C.
beguinii
var.
Chr.
C.
(Ces.)
C. Chr.
bisulcatus
18924, W;
iriquetrus (Bl.) Copel. (de
Chr.
12791, L)
Bot.
C. taeniatus
C.
(Brass 11699, L)
5241, L)
L)
C.
costulatum
Holostachyum
ined.
quinquefidum (Bak.)
(Cesati)
C. Chr.
(v.A.v.R.)
Holcosorus
L)
ebenipes (Hook.) Copel. (Polunin el
199
Polypodiaceue
costulatum
Copel. (Elmer
L)
C.
in
(v.A.v.R.)
wrightii (Hook.) Ching (Fosberg 37472, L)
22107, ,
G.
(Tagawa 7871, L)
Crypsinus albido-squamatus (Bl.)
C.
Stomata
Don) Tagawa ( Hennipman3393, L)
palmatopedata(Bak.)
Christ
(Kouy-tch 4162,
L)
N.
subhastata
Filices
(Baker)
Tagawa
chinenses exsicc.
US, L)
(Rosenstock,
200
BLUMEA
americanum
Niphidium
J.
(Hook.)
Sm.
VOL.
(Rose
27, No.
P.
P.
5334, L;
lanceolatum
Presl
(L.)
(Eggers 259, L)
P.
longifolia (Bl.)
s.n.,
Moore
ined.
(Buchtien
Underw.
& Maxon
P. decumanum
(L.)
J. Sm.
Sm.
commutata
diversifolium (Willd.) Pic.
J. Sm.
nigrescens
LAE
Serm.
(Varekamp
alcicorne
Baker
bifurcatum
Ching (Nedi & ldjan
9, L)
Leiden
Bot.
Leiden
(cult.
C.
Chr.
Bot.
Leiden
bifurcatum (cult.
Bot.
(Konig ex Muller)
Garden)
bifurcatum
ssp.
Garden)
Desv.
(cult.
Leiden
Bot.
Garden)
Bot.
elephantotis
(cult.
P.
grande
Leiden
(cult.
(Fee)
Bot.
Leiden
(cult.
Kunze
hillii
C.
bifurcatum (Cav.)
ssp.
(Moore)
De
ined.
Jonch.
(cult.
Chr.
Leiden
&
Hennipm.
P.
(cult.
Baker
(cult.
Bot.
Leiden
Tardieu
(Henderson 22617, C)
Desv.
(Beauv.)
(cult.
Leiden
(Beauv.)
'var.
Bot.
P.
delaurenlii
De
Garden)
Desv.
(cult.
Leiden
Bot.
Bot.
(cult.
P. wandae
P.
Jonch.
Pleopeltis
P. bicolor
L)
(cult.
(cult.
H.
Fee
L
(Und.)
Leiden
et al.
Chr.
Steenis
Bot.
Garden)
Joncheere
(De
willinckii
Kaulf.
MAD
( Heller 5011, L)
Langsd.
et Fisch.
(E.
el
Kze
chnoophorum
Midler
(Balansa 2878, L)
Ros.
crystalloneuron
Kze.
falcaria
(Buchtien
L
s.n.,
(Bourgeau 3187, L)
Schlecht.
fauriei Christ
& Cham.
(Bourgeau 1831, L)
(Tagawa 8500, L)
formosana (Bak.)
Nakai
fraternum Schlecht.
(Tagawa 7940, L)
et Cham.
Kunze
P.
lachnopus Wall,
P.
lepidopteris (Langsd.
(Pringle
s.n.,
L
C.
Chr.
( Stork 4794, L)
(Gerth 16, L)
ex
Hook.
et
(Bor 16265, L)
Fisch.)
Kze.
(Boom
Witte s.n.,
L
936224342)
loriceum
L.
(Kills
s.n.,
P. mammeiense Christ
P.
nipponicum
P.
Mett.
pellucidum
30703, L;
plumula
H.
al.
T1507, L)
(Tagawa 7538, L)
Kaulf.
Heller
L)
( Tagawa et
(Degener
&
Degener
2075, L)
B.
Willd.
(Arsene
s.n.,
L
913153591)
P.
polypodioides (L.)
no.
a!.
O.
90912309)
gilliesiii
(Heller 6300, L)
(Takeda) Sledge (Polunin
134, L;
2016, L)
friedrichsthalianum
1822, L)
(Fee) (Poilane 7939, P)
Liebm.
gladitum
var.
Sandwith
20, L;
P.
Garden)
ssp.
19615, L)
fraxinifolium Jacq. (Hatschbach 17495, L)
P.
Bot.
T
at.
(Splitgerber 374, L)
Willd.
B.
et
910211377)
Sinnema
californicum
L
(Ludlow
B. Willd.
s.n.,
(Iwatsuki
11465, L)
P.
(cult.
Garden)
Leiden
C.
(Van
s.n.,
P.
P.
Bot.
(Cav.)
ined.
abbreviata
astrolepis
Hennipm.
C. Chr. ssp. veitchii
Leiden
Hooker
bifurcatum
&
Garden)
Racib.
(Hooker)
P.
De
bifurcatum (Cav.)
P. wallichii
Chr.
attenuatum
australe
el al.
27412, L; Hennipman 4146, L;
superbum
&
90910270)
Garden)
Leiden
Tagawa ( Koidzumi
Mett.
ex
Shimizu
atkinsoniiC.
P. fallax
(cult.
Garden)
Bot.
Christ
P. stemaria
ined.
PNH
(Gutierrez
9137207)
P.
P. stemaria
P.
42600, L)
32467, L)
Wall,
amoenum
s.n.,
P.
quadridicholomum (Bonap.)
Leiden
P.
Moran
Copel.(Bell 2019, L)
Nijhoff &
Bot.
Garden)
Bot.
madagascariense
Wild.'
T
& Morse 674,
(Dorsett
amamianum
(Buchtien
P.
holttumii
ridleyi
P.
Bot.
Garden)
P.
colorata
P. catharinae
Garden)
Garden)
P.
Clemens
P. attenuatum H.
P.
bifurcatum
Leiden
P.
P.
Bak.
Garden)
P.
S.
Polypodium
P.
Schweinf.
Garden)
P.
al.
et
5.5.1923, L)
(cult.
Desv.
(Cav.)
coronarium
var.
(Iwatsuki
Holttum
angustatus
9581, L;
Leiden
P.
Bonner
et
Polypodiopteris brachypoda (Copel.) Copel. (J.
P.
(Brass 23887, L)
Schmutz F—
P. andinum
P.
(Maas
(Jacobs 8535, L)
Alston
(Don)
Kaulf.
thunbergiana
Podosorus
Garden)
P. ellisii
Tardieu
(Mett.)
L; Fosberg 37302, L;
M.
Platycerium
P.
Klf.
Willd.)
ex
(Christ) Copel.
sinensis
P.
L)
var.
Alston & Bonner
78332, L)
164, L;
P.
Meeuse
4916, L; 5781, L)
(Hennipman 3151a, L)
(Lam 6672, L)
(Bl.) Ching (Croft
&
(Lam
9586, L)
(Schlim 153, L)
Bl.
P. scolopendria (Burmann)
P.
s.n.,
6311, L)
scolopendria
(Boom 21075, L)
60581, L)
P.
schraderi
P.
aureum
Photinopteris speciosa
6,
P.
(Wright
1022, L)
Phymatodes
Parks,
16058, L)
(Bory
macrocarpa
P.
L)
crassifolia (L.)
et a!.
Geesteranus
(Meijer 1110, L)
(Lellinger)
carinata
Phlebodium
P.
(Ludlow
P.
12075, UC)
Moore
Maas Geesteranus
loriforme (Wall, ex Mett.)
Paragramma balteiformis (Brause) Copel. (Brass
Pessopleris
&
(Degener
(Bory)
excavata
L)
Paltonium
Kaulf.
23.6.1933, L)
Oleandropsis ferrea (Brause) Copel. (Brass 12795,
P.
1981
elongata
22816, US)
P.
1.
4894,
P.
prainii(Bedd.)
L)
Hitchock
(Rosenstock
det.
16336, L)
C. Chr.
(Schimizu
et al.
13254,
U.
P.
P.
pringlei (Maxon)
C. Chr.
C. Chr.
pycnocarpum
Sen & E.
P. scouleri Hook. &
Grev.
triseriale
P.
virginianum L. (Rolland
P.
vulgare
Sw.
L.
11855,
( Pringle
2332,
(Herzog
P.
Hennipman:
L)
L)
(Rose 66081, L)
(Smith 2564, L)
—
Germain
6013, L)
coronans
(Wall,
ex
Mett.) Ching
metacoelum
C.
Chr.
(Haltier 2942, L)
murudense
C.
Chr.
&
(J.
M.
S.
C.
Chr.
(Hirano
&
Leiden
Garden)
abbreviata
(Zoll.) Ching (Kostermans
22122,
P.
angustata
De
P.
Morton
(Van
Farwell
(Tagawa 2904, L)
(Sw.) Ching (Bunnemeijer6802, P;
Wilde &
De
Morton
( Brass 23703,
L;
rupestris (R. Br.) Ching (Evans 2478, L)
(Forsi. f.) Ching (Bernardi, L)
serpens
P.
sheareri
P.
sphaerosticha (Mett.) Ching (
(Bak.) Ching (Tagawa 109, L)
Waalkes
1668,
stigmosa
Waalkes
Van
Borssum
L)
(Sw.)
Ching
taenioides
5122,
Borssum
(Van
293, L; Hennipman
species (Hennipman
Saxiglossum
3015,
L)
L)
(C. Chr.) Ching (Fan
& Li
Selliguea feei Bory (Kostermans 293, L)
(v.A.v.R.)
confluens (R. Br.) Ching (Lam 7660, L)
dimorpha (Copel.)
floccigera (Bl.) Ching (Hennipman 3814, L)
Parris
(Hoogland4173, L)
P. flocculosa (Don) Ching (Hennipman 3352, L)
P. foveolata (Alston)
Morton
(Van Royen 4315,
heteractis
(Mett.
(Hennipman 3831, L)
ex
Kuhn)
Ching
(Iwalsuki el
al. S
1068,
brunei
(Werckle
ex
Christ) Wagner
(Liesner 662, L)
S.
bifrons (Hooker) Copel. (Cuatrecasas 11209,
US)
Synammiafeuillii (Bert.) Copel. (Lam 7885, L)
Thayeria cornucopia Copel.
(Copeland
s.n.,
uc
769060)
Thylacopteris
12838, L;
T.
L)
gralla (Gies.) Ching (Tagawa 3023, L)
Holtt.
L)
Wilde-Duyfjes 19544, L)
P.
P.
(Mett.)
Royen 5189, L)
Solanopteris
(L.)
P.
P.
P.
Royen
L)
lanceolata
L)
princeps
S. lima
longifolia (Burman)
4549,
P.
,
(Kunze) Ching (Surbeck 1012, L)
702, L)
L)
P.
Togashi 447, L)
3350
nummularifolia(Sw.) Ching (Jacobs 8407, L)
P.
cult.
Farw. (
(Hennipman
P.
Hotta
163, L)
Pyrrosia species (Hennipman 5122,
P.
mollis
P.
rigidum (Hook.)
Bot.
P.
201
P.
Clemens
33980, L)
P.
stictica
P.
(v.A.v.R.)
Polypodiaceae
lingua (Thunb.)
Van
(Hennipman 3995, L)
P.
P.
in
P.
(Pauca 1637a, L)
Pseudodrynaria
Pycnoloma
Stomata
diaphana (Brause)
Schlechter
papillosa (Bl.)
Copel. (Brass
18220, L)
Kze.
(Anderson
S.
29900, L)
Weatherbya accedens (Bl.) Copel. (Elmer 9721, L)
W. damunensis
G)
( Ros.) Copel. (Schlechter 14399,