The emendation
of the fern
genus
transference of
the
including
Christiopteris,
two taxa
to
the microsorioid Polypodiaceae
By
E.
and W.L.A. Hetterscheid
Hennipman
With
1
in
figure
the
text
Abstract
Hennipman,
E.
&
Jahrb. Syst.
the
In
sagitta
both
105:
present
are
Hetterscheid,
1—10. 1984.
the
paper
retained
in
one
remains
transferred to the
genus
Christiopteris
genus.
C.
variant
formas.
as
the
as
A.:
The emendation of the fern
two taxa to
the microsorioid
Christio-
genus
Polypodiaceae.
—
Bot.
ISSN 0006-8152.
—
genus
the
originally published
other
W. L.
the transference of
pteris, including
To
typical form
Microsorium
is
one
to
emended. The species
proved
to
of
consist
of these the rank
C.
tricuspis and
C.
two
of species is
variant. Furthermore,
C.
different species,
given,
both species
the
are
(Polypodiaceae).
1. Introduction
The
genus
Polypodium
its
Christiopteris
was
founded by COPELAND (1905)
systematic position
within the
Polypodiaceae
has
been
to
the
sagittum Christ. Subsequent authors all accepted
accommodate
genus
ambiguous
creation onwards. In its modern construction it should consist of three
i.e.
C.
tii;
mainland
sagitta (syn.:
Loyalty
Our
the
This
and
Asia
latter
study
varians
tricuspis (syn.:
(New Caledonia);
revealed
also
Christiopteris
much different
revealed
taxon
species,
C. eberhard-
reported
from the
to
from that
consist of
two
to
be
heterogeneous,
from New Caledonia.
different elements which
rank in Microsorium s.l.
is executed
(1984).
C.
1969).
being
given specific
Hennipman
C.
Polypodiaceae
on
from
the
have been
copelandii; Philippines),
Brownlie
studies
species
Besides,
Asia),
by
Is.
C.
though
from its
We
as
are
part of the
thankful
to
Polypodiaceae
co-workers
Project.
for
help
For details
0006-8152/84/0105-0001$
©
1
Botanische
Jahrbiicher, Bd.
105
1984 E. Schweizerbart’sche
see
in various
ways.
Verlagsbuchhandlung, D-7000
02.50
Stuttgart 1
E.
2
especially
Mr.
to
Hennipman &
R. P.
Baayen
the herbaria of Paris
present in
(U).
The
tial
thanks
due
are
information about the
Dr. F. Badre
his
(P) for
and Mr.
M. C. Roos.
London
(P),
Dr. B.
to
help
Zimmer
the
In
original
listed
(1906)
described
sagitta
Pleopeltis,.
sect.
and
later
He
could
cantoniensis
(Baker) Ching)
cantoniensis
cies
C.
not
be
Hooker)
even
varians
The
replaced
“Christopteris”).
LAND
be related
late
hairs,
and
Cheiropleuria
Dipteris,
the
Christiopteris
included
later
species
of
to
which
Myuropteris
genus
Ching
Colysis.
study
(= (C.
with
tri-
Leptochilus
spe-
which
Leptochilus
he
time corrected
ferns,
apart from the
the
e.g.
polypods
(1928) related
because of
shared
a
COPE-
with
stel-
Dipteris (Dipteridaceae)
genera
Bower
(Christiopteris
regarded by
was
(1915)
varians
included in Chris-
subsequently
same
thus construed
as
expressed
He
Copel.
(1933)
was
Ching
cantoniense
Campium
dilatatum in
he
the
doubts
(1930)
the
regarded
of
as
the
Christiopteris
occurrence
to
of the
Campium
by
nature
with both
(Christ)
Ching
of
to
monotypic
new
(1933), followed
Campium
to
Campium
Ching.
CHRISTENSEN
(= Drymoglossum
Ching
cantoniense
inclusion
with COPELAND and
Ching.
cordata
drymoglossoid.
the
to
The
be related
re-
(1929)
corda-
genus
Chris-
refuting
and
thought
dilatatum and
genus
im-
though
Leptochilus
to
of
resemblances of this
who emended Baker’s
Christensen’s
polyphyletic
cordatus
(Baker)
Myuropteris
type
regarded
transferred
Myuropteris)
strong
Christiopteris mentioning
species possibly conspecific
(syn.:
=
SCHUMANN
ferns by
to
Christiopteris cantoniensis, agreed
recognizing
Myuropteris.
this
(1928)
dilatatum
Campium
in
(1917)
the
at
was
Platycerium
cantoniensis in
Christ) Christensen,
tensen
Polypodi-
(“Christopteris”)
whether the latter
Cheiropleuria
the acrostichoid
number of different
and
and
better united.
(Cheiropleuriaceae).
Campium
ferred the
The
drynarioid ferns,
perfect description
tum
a
genus
de
eberhardtii Christ
questioned
genus
Christ
sagitta),
venation in the fertile parts of these ferns.
Copeland
to
to
Cheiropleuria,
diplodesmic
fern
of the
name
C.
cantoniense Baker
Fournier which he
Mett.)
tiopteris.
to
on
the
only.
derive
Christiopteris
C.
Christ,
Copel.
’erratique’
genre
stimulated Copeland
(1917)
to
Copeland
(“Christopteris”)
copelandii (=
added
Christ
in
placed
Detailed anatomical work
(Acrostichum
petit
(1908)
Christ.
better
by
Christ.)
sagittum
(Gymnogramme
thought Christiopteris possibly
and Bower
Christiopteris
represent “un
.
essen-
Mettenius, and
history
(“Christopteris”)
tricuspis (Acrostichum tricuspe
cuspis),
of
(Polypodium
Christiopteris
to
and Utrecht
in various ways.
description
the genus
thought
um
C.
(L),
gratefully acknowl-
who communicated
(B)
collections studied
holotype
2. Taxonomic
(1905)
The material studied is
Leiden
(BM, K),
of the Directors of these institutes is
cooperation
edged. Special
Hetterscheid, Christiopteris
W. L. A.
Leptochilus
Leptochilus
the polypodioid
sensu
genus
E.
Copeland
of
when
(1947)
and C.
tricuspis,
i
C.
gitta,
Hennipman &
W. L. A.
discussing
the Microsorium-Phymatodes
likely
more
to
in the
belong
The report of the
CoTTHEM
(1968)
the interest in the
strated
posed
relationship
Wilson did
induced PiCHl
s.str.
confirm
of the
much
is
in another
paracytic
stomata
also found
position
by
not
study
the
sa-
suggestive
whole
seems
Christiopteris by
in
of
stomata
(1977)
Dipteris
Wilson
Christiopteris
the
give
to
Cheiropleuria).
in the
that found
to
in
most
Christiopteris
within the
sup-
(1981)
position
failed
to
found that the shape
we
other
and
The
Polypodiaceae
isolated
an
genus
demon-
(1959)
(and
VAN
revived
—
sporangia of Christiopteris.
observations. Furthermore,
similar
of
a
as
genus
him in
Christiopteris. Earlier,
Polypodiaceae
being
The
Cheiropleuriaceae.
will be discussed
Polypodiaceae
paper.
3.
I.
The
from that found in Dipteridaceae
different
systematic
genera.
In the meantime Sen & Hennipman
COTTHEM’s
sporangia
of
of the sporangia of Dipteris
Sermolli
Polypodiaceae.
VAN
of
of the paracytic
occurrence
within the
of
stomatal type
several characteristics
Unfortunately
group
Christiopteris
“more
Crypsinus group”.
occurrence
a
—
the differences between
thought the latter species
varians,
3
Hetterscheid, Christiopteris
Taxonomy
Christiopteris Copel.
in
Christiopteris Copel.
Perkins, Fragm. FI. Philipp. (1905) 188; Philipp. J.
C, Bot. 12 (1917) 311, p.p.; Bower, The Ferns
Fil.
Fl.
(1947)
178, p.p.; Holttum, Ferns of Malaya (1954)
Philipp.
3
(1960) 456,
Caled. 3 (1969) 289,
p.p.
The
(1928) 213,
3
genus
is
p.p.;
—
Brownlie, Pteridophyta
Type:
here defined
Sci.
Gen.
210, p.p.; Copel., Fern
Aubrev.,
in
FI. Nouv.-
Christiopteris sagitta (Christ) Copel.
the characters
by
p.p.;
Copel.,
as
given by
COPELAND in the
original publication.
Key
the
species
Sterile fronds sagittate
la.
b.
1.
to
Sterile fronds
1.
deeply palmately lobed,
lobes 3-5
Christiopteris sagitta (Christ) Copel.
188;
457.
Philipp. J.
—
Type;
1, Suppl.
Polypodium
sagittum
2
in
Bull.
Christiopteris sagitta
Christiopteris tricuspis
Perkins, Fragm.
(1906) 157, pi.
Christ,
2.
Fl.
Philipp. (1905)
13; Fern Fl. Philipp.
Herb.
Boissier
6
(1898)
3
(1960)
199.
—
Loher.s.n. (P).
Christiopteris
Bower,
Sci.
..
The
Syntypes:
Whitford.
copelandii
Ferns
small
3
Christ, Bull. Herb. Boissier,
(1928) 218;
specimens
Copel.
from the
Fern
Fl.
Ser.
Philipp.
collections made
3
by
2,
6
(1906) 990;
(1960)
457.
COPELAND
—
and
4
E.
Hennipman &
The
Description:
by
Christ
cit.
(op.
Christiopteris
2.
Philipp.
Sci.
J.
722—726, pi.
7, 2 (1941)
chum
Ferns
Brit. India
Ferns Brit. India
(1875)
mann,
139.
Flora
108
Christiopteris
ipp.
J.
Gen.
Sci.
Fil.
hardt 84
(P,
of
263.
—
Malaya
Bot.
(1954) 211.
Acrosti-
—
Gymnopteris tricuspis
—
Handb.
s.n.
Filic.
The
Schu-
(1906) 388;
(K, n.v.).
Phil(1908) 272; Copel.,
21
(Morot)
Hist.
tricuspe J. Smith.,
Chr., Index
(1917) 332; Bower,
Indo-Chine
descr.; Suppl. (1876) 27;
c.
Atkinson
Type:
Christ, J.
304.
FI.
fig.
214,
(1928)
in Lecomte,
Cheiropleuria
C.
3
Copel.,
273;
(1908)
21
Ferns
Ferns
(1928) 218; Copel.,
3
179; Holttum, Ferns of Malaya (1954) 211.
(1947)
2
12
The
C. Chr.
Ferns
tricuspis
(1915) 250.—
eberhardtii
Bot.
J.
(1864) 272, pi.
5
fig.
Leptochilus
Bot.
C,
Tard. &
(1866) pi. 53,
(1883) 434,
—
Christ,
Holttum,
Sp.
1905,1906).
(1917) 332; Bower,
Fil.
described and illustrated
excellently
Luzon and Mindanao.
(Hooker)
12
52: 2;
tricuspe Hooker,
Beddome,
Fil.
Bot.
fig.
450,
Philippines;
(frontispiece);
s.n.
is
species
present
tricuspis
C,
Hetterscheid, Christiopteris
and Copeland (op. cit.
1898)
Distribution;
W. L. A.
Eber-
Type:
—
sh.).
Description
See Bower
:
(op. cit.)
Distribution: Mainland Asia
and Holttum (op.
India
(northeastern
to
cit.).
Indo-China).
Excluded
3.
Christiopteris
cantoniensis
Baker in
nogramme cantoniense
cantoniense C.
Polypodium
—
toniense
Sinensia
Ching,
Fan Mem. Inst. Biol.
4
1
(Baker) Christ, J.
21
Bot.
Icon. PI. 3, 7
Hooker,
515.
Chr., Index Filic. (1906)
(1930)
(1933) 343.
53.
—
Leptochilus
Type:
—
272.
—
descr.
c.
Ching,
can-
Bull.
North
(Kwantung,
s.n.
Gym-
—
Campium
cantoniensis
Ford
C.
(1908)
(1887) pi. 1685,
River), (K, n.v.).
=
Leptochilus
cantoniensis
(Baker) Ching
II. Microsorium Link in Hort. Berol.
Full
ently
of this genus is
synonymy
under
investigation. Pending
here defined in the
ly
sense
accommodated in
Key
to
the
(1833)
given
110.
its
as
generic delimitation
the results of this
of COPELAND
Phymatodes
species
2
not
Presl
(1947)
i.e.
investigation
including
(= Phymatosorus
of Microsorium
those
Pichi
formerly referred
to
is pres-
the genus is
species
usual-
Serm.).
Christiopteris
varians
l.a.
Scales of the rhizome
of sterile fronds
b.
Scales of the
appressed,
8 —20,
rhizome
apex
±
round
to
slightly elongate;
1.
acute
spreading,
triangular,
dex of lobes of sterile fronds 3-6,
obtuse
apex
apex
2.
...
index of lobes
Microsorium varians
acute
to
acuminate; in-
Microsorium latilobatum
E.
1.
Hennipman &
Microsorium varians
Fig.
W. L. A.
Acrostichum varians, Mett., Ann.
varians
varians Fount., Bull.
Leptochilus
Bot.
Nat.
18
(1873) 273;
nov.
—
J.
Philipp.
Nat.
Sci.
J.
Ser.
Nouv.-Caled. 3
See
note
12
Bot.
8
Bot.
B,
39
Jahrb. Syst.
C,
289.
7.
57.
(1861)
Ann.
Sci.
varians Diels
Gymnopteris
—
varians
Christiopteris
—
Brownlie,
Pteridophyta
Copel.,
Natl.
Mem. Mus.
Vieillard 1526, Poila
Type:
15
Roux, Nova-Caled. Bot. (1914)
Guillaumin,
(1917) 333;
—
Bot., Ser. 4,
France 15 (1868) 394;
442.
(1922)
(1906)
(1962) 202;
(3)
(1969)
45
Acrostichum varians Mett. f.
—
Nat.
Soc. Bot.
(1861) 56; Baker
15
Bot., Set. 4,
524.
in Sarasin &
Bonap.
Linn. Soc. Bot.
Bot.
Schlechter,
Nat.
Sci.
(1874)
ICBN, “normalis”), Ann. Sci.
24
(art.
ed. 2
Baker, Syn. Fil.,
38; Compton,
in
Hetterscheid, comb.
&
(Mett.) Hennipman
la.
in Hooker &
—
5
Hetterscheid, Christiopteris
Hist.
iso in P,
(B,
FI.
Aubrev.,
in
p.p.).
1.
Acrostichum
15, (1861) 57.
varians
Mett. f.
Mett.,
contracta,
Vieillard 1525, Balade
Type:
—
Ann.
Sci.
iso
(B,
in
Nat.
P,
Bot.
p.p.).
Ser.
See
4,
note
2.
varians
[Leptochilus
Bull.
Fourn.
P
var.
(1873) 273,
varians
Bonaparte,
terete
scales
scales;
3.5x0.7—2.0
terminated
marginal
cent
articulated
petiole
to
2—15
middle,
dex 8—20,
apex
wide;
the
the
cm
branched,
ex-
35x12—50
0.7
cm;
85
x
or
x
varians
(Mett.)
Ser.
5,
18
at
of 2—4
rows
cm
middle,
secondary
recurrent
terminated by
45—60
index
two
veins
fronds
to
up
to
pm, exospore
cell.
cm,
cm
long;
straight,
see
in-
margin entire,
cm,
other veins
also Fig.
la.
cm
Veins
immersed,
containing simple
or
Fertile
index 0.8—1.5,
15—
index 20—100, 8—24x0.2—
10 —20x0.3—0.8
glandular
translu-
widely spaced,
somewhat below
4
cm;
intermixed with 2—4-celled
a
rounded,
Venation:
cm.
long; lamina,
cm,
with
lowermost lobes 0.5—2
prominent,
6
stout,
set
a
20—55
usually
of smaller areoles
cm
apex
Fronds
up
non
1.0—2.5, 1.0—
clathrate with
—
spaced
3—30x0.7—4
5—17
2—14, spaced
polypodiaceous,
0, densely
cm
5—25x0.4—3.5
free included veinlets;
long; petiole
lobes
cells.
Sterile
lobes 4—14,
the
polygonal, consisting
and
are
of
the sinuses above the
and
Rhizome
epilithic.
0.1—1.0
auct.
nud.].
margin irregularly denticulate,
cm;
at
latioribus”
slightly triangular,
terminal segment, index 20—40,
45—60
angustioribus”).
Nat. Bot.
nom.
lamina index 0.6—1.3, widest
usually
rhachis
cm;
which
Sci.
pinnis
p.p.,
or
flattened,
segment, index 6—14,
tinct; sporangia
physes
rounded,
“a;
var.
rhizome, pinnatipartite.
the lamina
fronds 20—40
Ann.
(1921) 252,
epigeous
rounded
consisting
long;
areoles
largest
13
somewhat
15—40x17—40
anastomosing,
Fourn.
sometimes
or
base
zone
terminal
lobis
(3:
Leptochilus
—
relatively large glandular cell, centrally
a
widest
acute;
(Mett.)
peltate,
mm,
by
(“var.
nud.
angustioribus”),
Notes Pterid.
epiphytic,
long-creeping,
(1
var.
nom.
nud.].
nom.
Usually
Fourn.
(1869) 394,
(“var. P; pinnis
[Leptochilus
Foum.:
the
(Mett.)
Soc. Bot. France 16
Spores
venation indis-
receptacular
microsorioid,
thin, perispore inconspicuous.
para70—
6
Hennipman &
E.
Habitat:
(up
to
5
In
mountain
wet
Distribution:
studied:
Specimens
got
P,
M.
see
(P,
Note
chosen
1
1),
collections made
and is without
tum, of which
a
by
a
2.
—
trunks
tree
Baudouin
(P);
8788
Viedlard
(P,
not
one
5646
6558
(P),
1526
98?) (P),
Lecard
(P);
(=
1526
Vieillard 1526f,
”)
the
other
2),
see note
from
Poila.
apart
from
from
only,
likely
are
to
one
the
represent
these represents Microsorium
varians
latilobarepresent Microsorium
two
viz.
1525
It
mixture.
specimens
Balade,
Three
Wagap.
Paris
at
latilobatum
constitutes
a
all from Balade
Microsorium varians,
Microsorium
a
from Poila (erroneously
one
“1526”
locality record,
of
from
Paris constitutes
at
Microsorium varians,
annotated with
a
s.
s.n.,
Maza-
(P);
B).
record. The other
has
Arembo,
Baumann-
(P);
s.n.
8892
(P),
Mackee
(P);
Aoui, Me
Me
Ou Hinna.
mixture.
It
from
apart
Vieillard
“No.
stat.
and
spec,
Bot.
Sen
(annotated:
Wagap.
latilohatum
Hennipman
Hetterscheid,
&
lb.
Fig.
57.
(P)
representing
Acrostichum
(1861)
1528
specimens
Microsorium
nov.
(=
Brownlie),
locality
1526—1528
=
958
The collection Vieillard
three
specimen
1525
(P),
Vieillard. One of
by
only
Note 2:
Ammeri,
(P),
Microsorium latilobatum from
herbarium of Pancher
comprises
(P),
98
specimens of
lectotype
of
1579
8777
The collection Viedlard
:
and
trees
on
1); Schlechter 15612 (B, P); Viedlard 1525 (P, p.p.;
see note
two
as
specimen
Mme Le Rat 63
note
p.p.;
comprises
Balansa
5441
Herbier Rancher
(P);
s.n.
p.p.;
1526
&
epiphytic
Dogny, Canala, Balade, Poila,
Bodenheim & Guillaumin
(P);
forests,
Caledonia: Me
New
Col d’Amieu, Plateau de
loc.
Hetterscheid, Christiopteris
Altitude 300—1100 m.
high).
m
W. L. A.
variant Mett. f.
Vieillard
Type:
—
Ann.
major Mett.,
1528
(“«Poila»
Sci.
Nat.
{1528)”) (B;
not
4,
P which is
15
M.
varians).
[Leptochilus
Soc.
Bot.
Foum.
273,
var.
nom.
a
16
(“van
(Mett.)
(1869)
a:
var.
(“var.
a
nud.
nom.
pinnis latioribus”),
varians
Bonaparte,
Usually
(Mett.)
epiphytic,
elongate,
Fourn.
Notes Pterid.
acute
13
sometimes
long-creeping, flattened,
tate,
Fourn.
394,
—
a:
lobis
Bull.
latioribus”),
Leptochilus
Ann. Sci. Nat. Bot.
varians
Ser. 5,
(Mett.)
18
(1873)
nud.].
[Leptochilus
Fount.:
varians
France
to
0.5 —1.5
(1921)
252
epigeous
cm
0,
“a;
var.
pinnis
p.p.,
or
nom.
latioribus”
epilithic.
densely
acuminate, index 1.3—5,
set
auct,
non
nud.].
with
2.0—8.0
Rhizome
scales;
x
stout,
scales
1.0—2.5
mm,
pelbase
E.
Fig.
a,
1.
Hennipman &
Projection photography
W. L. A.
of venation
Microsorium varians ( Balansa 1579, P).
rounded
to
tridentate,
cells, marginal
dular
cell,
centrally
of scale
clathrate with
pattern
—
b,
margin irregularly
teeth translucent
apex
Hetterscheid, Christiopteris
a
acute,
or
of the central
M.
part
7
of sterile lobes.
denticulate, provided
with
glandular
partly clathrate, each terminated by
terminated by
translucent
marginal
—
latilobatum (Vieillard 1526, P, p.p.).
a
relatively
zone
a
glan-
large glandular cell,
consisting
of
1—2
rows
of
8
E.
cells.
ile
Fronds
—
fronds
usually
est
the
at
margin entire,
long; petiole
cm
middle,
lowermost lobes 0.5—3.0
7
Venation:
cm.
veins
tertiary
ge-sized
taining
simple
lb.
Fig.
veins
which
index 0.8—1.5,
venation
15 —50
12—45
cm;
lobes 2—10,
the
crosorioid,
50—60
which
30—40
x
the
cm,
7—23x1.8—5.5
cm,
spaced
to
up
sinuses above
tertiary
the
—
long; petiole
lobes 2—8,
veins
2.5—15
spaced
of the
and part
cm
8
to
up
lar-
forming
smaller areoles
sized
two
15—25x3
free included veinlets;
recurrent
cm
wid5
secondary,
variously
Ster-
—1.6,
con-
also
see
long; lamina,
cm,
index 20—
terminal segment, index 15—50, 8—16x1.3 —1.0
cm;
paraphyses
the
at
rhachis,
indistinct; sporangia polypodiaceous,
ceptacular
middle,
1
index 3—5,
segment,
immersed,
and
ex-
fronds
pinnatipartite.
lamina index
long;
lamina
subdivided into
are
10—40x
3—20x0.3 —0.8
65,
the
anastomosing;
branched,
or
Fertile
—
cm;
terminal
other veins
prominent,
areoles,
wide;
cm
above
obtuse;
cm
the rhizome,
to
4 —25
usually
broadly
apex
Hetterscheid, Christiopteris
20—50x20—40
widest
3—6,
W. L. A.
articulated
widely spaced,
15—73
index
straight,
Hennipman &
terminated
are
x
30—40
pm,
intermixed with
by
a
cell.
glandular
exospore
cm;
4—5-celled
re-
Spores
mi-
thin, perispore
inconspicu-
ous.
Flabitat:
sometimes
In
epigeal
wet
or
Distribution:
de
d’Amieu, Plateau
Noir Forest,
mountain forests,
epilithic.
New
usually epiphytic (up
Altitude 300—800
Caledonia: Mt. Mi, Me
Dogny, Mepeou Forest,
Wagap,
Nimbayes,
high),
m
Aoui,
Col de Parari,
Col
Yahoue Forest, Tendia Forest,
Houailou.
Balade,
12
to
m.
—
Islands:
Loyalty
Pic
Mare
Island.
studied:
Specimens
Franc 656
(P),
686
(P),
P);
Germain
s.n.
M.
&
Le Rat
12141
Mme.
(P),
lard 1526*, P, 2
dat.
(P);
p.p.;
sh.;
see note
1
(P);
74
799
note
see
1525
(P,
1
to
p.p.;
(P);
(Ros.,
Baudouin
Fil.
(P),
(P);
7953
Herbier
Microsorium
see
note
2
to
1528 (B,
varians).
(P),
exsicc.
10247
8170
varians).
(P),
453
32) (B,
no.
(P),
10415
(P);
(P),
12043
(P),
Pancher “1526”
Anon,
Microsorium
not
Cribs
(P),
4
Nov. Cal.
Baumann-Bodenheim
Mackee 4777
Microsorium
to
s.n.
Montrouzier 250
(P);
Vieillard
Balansa
(P),
Guillaume &
(P);
14420
1081
s.n.,
Vieil-
(=
loc.,
s.
varians).
1526
s.
(P,
P).
4. Discussion
In
an
reasons
of
extensive
that
Christiopteris,
species
and
by
also
Bower
Christiopteris
physes;
apex.
fronds
To
of
the
mentioning
tricuspis
Christiopteris,
The
can
two
COPELAND
the
tricuspis
results of the
mentioned included small,
(1917)
as
a
gave
Christiopteris
simple,
rhizome-scales
or
drawn
ample
second species
anatomical studies
similarities between both
venation; peltate
this
the
on
recognition of Leptochilus
(1917).
diplodesmic
hair-like
sterile
discussion
justified
on
branched
out
into
this
sagitta
a
para-
long
be added the resemblance in venation pattern of the
species,
being
a
complex
pattern
of
areole-layering.
E.
The
inclusion
Copeland
(rather
of
the
at
Hermipman
weak)
& W. L. A.
Leptochilus
varians
time seemed
same
diplodesmic
as
third
a
the
of
species
based
primarily
venation,
9
Hetterscheid, Christiopteris
on
“translucent”
Christiopteris
by
the resemblance of the
scale
and
margin,
the
acrostichoid condition.
Our results
as
Christiopteris sagitta
the inclusion of
sense
varians. The
C.
other genera of
to
The
sister
above confirm the
given
of
consisting
species
called
which both
species
occur
shape
to
prep.);
occurring
P.
cromwellii,
the
all
type
P.
(e.g.
but
nigrescens,
or
ned
a
a
suggest
atavistic
remains
relationship
fronds
raleptochilus
ty
in
to
a
M.
possibly
of
spe-
of
P.
sco-
Hennipman, in
e.g.
Phymatodes
sablanianum,
etc.
Hennipman 1984;
of the microso-
are
in the greater
of
species
has the
Phymatodes
lepisorioid
to
spore
one
spe-
the characters mentio-
alliance, using
specific
fronds of
these acrostichoid
element in Microsorium s.l.
do
polypods
decurrens
compared
associated with the
commutatum
M.
varians
condition which recalls
one
the atavistic
e.g.
to
& Hennipman
Very
soral
might
informative
of
for
fronds of Pa-
conditions
generally
soral instabili-
1984). Further,
development
ferns
which is essential
give information
phylogenetic relationships,
(Leptochilus)
fronds
to
states
similar in
are
occurring
type
which
species
of Microsorium latilobatum and M. varians
of other
Phymatodes
atavistic
&
that of
to
of the
part
have been
Phymatodes (viz.
Baayen
spore
in
scolopendria
P.
Colysis (Hetterscheid
which is
e.g.
in
not
and
They
species
bamlerianum,
a
inferred
two
ambiguous.
the reconstruction of
found in
latter
Hetterscheid &
1983),
wondered whether atavistic
We
the
resemblance
in the Microsorium
species-group
above,
ROOS
of Microsorium,
species
type). The relationship
cies
a
of
the character
by
Lecanopteris,
Microsorium
paraphyses), cf.
&
(HENNIPMAN
in this restricted
two
of both Microsorium varians and M. latilobatum
of the
part
commutatum,
e.g.
these
out
subgeminatum;
P.
refuting
time
investigation.
of
paraphyses
species of
and
have “normal”
spores
rioid
in
same
varians consists
bom
especially
Christiopteris
genus
the
in New Caledonia.
exclusively
venation pattern shows
P.
nigrescens,
(which
latilobatum. The
those
the
is under
Christiopteris
cies of the Microsorium alliance is
lopendria,
at
Microsorium. The resemblance of
accommodated in
Microsorium
tricuspis,
relationship of Christiopteris
Polypodiaceae
formerly
the
justification of
and Ch.
is found
acrostichoidy
and Microsorium bamlerianum. However, the
studied
( Vieillard 1526, B,
of the soral situations
do
P)
not
show
a
found in Micro-
generally
as
sorium s.l..
As
tum
regards
the
relationship
between Microsorium
it should be noticed that the
sized than
of M.
spores
those of M. latilobatum. The
this, that these differences
are
possibly
suggestion
due
to
varians
varians
usually
is
tern,
—,
etc.
Roos
supported by
when
1983).
applying
If
the
the
characterstates of the
the out-group rule
suggestion
of
made in
polyploidization
matically qualifies the differentiating characterstates of
phies
in
a
M.
—
varians
paraphyses,
general
(autoP)-polyploidy
latiloba-
and M.
significantly larger-
are
way
is
cases
which
as
like
auto-
automor-
venation pat-
(Hennipman &
true,
we
have
a
E.
10
unique
of
case
search
endemic
to
of
study
and
processes
rather small
a
differences in characterstates.
for the
object
Hetterscheid, Christiopteris
W. L. A.
sister-species
two
number of
large
Hennipman &
This
island, showing
should be
of
strategies
a
promising
speciation
a
re-
tropical
in
ferns.
References
F. O.
Bower,
1917:
“acrostichoid”
Bot.
nie
2
1—39,
The ferns (Filicales).
G.
Brownlie,
et
Ching,
31:
(London)
1928:
—
in
Studies
1969:
phylogeny of the Filicales
1930:
The
reference
VI.
showing
Ferns
derivatives.
dipteroid
to
the
Ann.
—
pi.
Vol.
3. The
Pteridophyta.
Dependances. Vol.
R. C.
the
condition, with special
3. 307
leptosporangiate
ferns.
—
Cambridge.
Aubreville, A., Flore de la Nouvelle-Caledo-
In:
—
pp.
studies of Chinese ferns
I.
Species
novae.
Sinensia
—
1: 43
—
56.
The studies of Chinese ferns
1933:
—
Mem. Inst. Biol.
H.
Christ,
Filices insularum
Philippinarum (2).
Fougeres d’Annam franyais recueillies
1908:
21:
Microsorium, Colysis,
Leptochilus.
Bull.
—
Fan
Bull. Herb. Boissier,
—
Ser.
6
2,
987—1011.
(12);
—
1906:
X.
4: 293—-354.
M.
par
Eberhardt.
—
J.
Bot.
(Morot)
261.
C.
Christensen,
with
of their
E. B.
butions
—
1917:
—
1928:
coenosori
phylogeny.
Copeland,
Taxonomic
1929:
longitudinal
1905:
Dansk
—
Ferns.
fern-studies
Ark.
Bot.
In:
—
The
Christiopteris.
genus
Leptochilus
and
—
6:
13
Ill:
with it.
—
with
a
genera
discussion
Florae
Philippinae.
Contri-
175—194.
12: 331 —336.
Sci., C. Bot.,
Philipp. J.
polypodioid
pi.
J.; Fragmenta
Islands, fasc.
confused
genera
of the
Revision
“Drymoglossinae”);
1 —93,
Perkins,
the flora of the Philippine
to
I.
and
(Cochlidiinae
Sci.
Philipp. J.
37:
333—416,
32
pi.
—
1947:
Genera
copsida.
—
icales).
1984:
Taxon
—
E.
Hennipman,
(Filicales).
&
—
Chronica
Botanica
Co.
studie
Vergelijkend-morfologische
A
33:
new
approach
de
van
M.C.
Roos,
Verb. Naturwiss.
the
to
systematics of
bij de Fili-
the Polypodiaceae
&
1983;
Phylogenetic
Vereins
Hamburg
(Fil-
Hennipman, E.
1984:
systematics
of the Polypodiaceae
26: 321—342.
Venation
patterns,
of the fronds of the microsorioid Polypodiaceae.
shape
stomata
Gent.
140—141.
W. L. A.
Hetterscheid,
tics, and
—
1968:
Proefschrift, Rijksuniversiteit
E.
Hennipman,
Filicum.
W.
Cotthem,
van
—
soral
characteris-
Bot.
Jahrb. Syst.
105: 11—47.
Pichi
Sermolli,
R.E. G.
dinem redigendi.
Schumann,
E.
1915:
—
1977;
Tentamen
Webbia
Die
Pteridophytorum
genera
in
taxonomicum
or-
31: 313 —512.
Acrosticheen und ihre Stellung
im
System
der Fame.
—
Flora
108:201—260.
Sen,
U.
—
&
Wilson,
—
Hennipman,
Blumea
K.
A.
Contr.
27:
E.
1981;
Structure
and
ontogeny
of
stomata in
Polypodiaceae.
175—201.
1959:
Sporangia of the fern
Gray Herb.
genera
allied with
Polypodium
and
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185: 97—127.
Accepted for publication May 7,
1984
Address of the authors:
E.
Hennipman
& W. L. A.
Hetterscheid, Institute of Systematic Botany,
sity of Utrecht, Heidelberglaan 2,
NL-3508
TC Utrecht, The Netherlands.
Univer-