Distichous and Pseudodistichous Mosses

6
Acta Botanica Neerlandica
Distichous
386—391
(1957)
and
Pseudodistichous
R.
van
der
Mosses
Wijk
( Groningen)
March
(;received
In
the
revising
family
found that in many
the leaves are,
that
of
type
of the
noticed
was
of the
regarded
Naturl.
Eucatagonium
comprises,
far
as
5a). The leaves
the
distances
The
equal.
I
between
check,
the
The last-mentioned genus,
and further in Africa
to
genus
as
a
the
the
175,
(1925)
first
not
the
to
the
on
family
Reichdt.
Phyllogonium,
genus,
species
only (Fig.
complanate branches,
in
leaves
both
S. America and Africa
of
subgenus
5b shows
At first
two
sight
and
by
also
occurs
rows
(Congo),
the
being
and
is
strongly ampli-
branches of
the
11
3
undoubtedly
one
the
(1925)
consecutive
5b
Fig.
they
a
a
leaves
not
on
fig.
side
and
et
Wils.)
arranged
in
of the
even
the
two
Fig.
Broth.
rows.
matter
a
of
Naturl. Pflanzf.
comparing figures
Fig.
the
to
mosses.
different from that
565 of Brotherus,
show them. In
each
unequal
are
f.
the
regarded
was
easily overlooked. As
the difference: in
sees
totally
America,
belonging
genus
regularly
not
are
of
copy
does
176,
is
phyllotaxis
are
irregularities
ed. 2,
Muell.,
in S.
transferred
change it
Eucatagonium politum (Hook.
leaves
3, which is
fact, Fig.
1922 Fleischer
family includes several complanate
it is clear that the
these
In
Before this
C.
Catagonium
this
Phyllogonium:
However,
Eucatagonium,
and Australia.
family Phyllogoniaceae.
Plagiotheciaceae;
in
The
consecutive
to
had
leaf bases.
ate
in
bryologists
distichophyllous
the well-marked alar cells
by
diagnostic
Phyllogoniaceae.
11
2,
alternately
is restricted
genus
characterized
Fleisch.
could
inserted
are
ed.
As
character
paid special attention
the
to
family,
disposed.
Phyllogonium Brid., Orthorrhynchium
(Broth.)
as
main
I
Malesiana,
distichously
not
have
Pflanzf.
Flora
included in this
that other
me
I
the
were
the
as
belonging
includes three genera, viz.
and
to
reason
species
Brotherus,
for
far
so
expectation
surprise
a
this fact. For this
phyllotaxis
In
it
which
to
is
phyllotaxis
family,
1957)
Phyllogoniaceae
species
contrary
11th,
5b and
3 the distances between
stem
are
whereas
equal,
alternation is
not
always
regular.
If in
is
the
Fig.
5b
the leaves
To
phyllotaxis?
are
answer
not
distichously
this
determine what the result would be, if
a
phyllotaxis
belonging
The main series
1,
2,
3,
5,
preceding
chies and
8,
in
etc.,
Each
simultaneously
is
term
the
main
formed
in which
any
have
by
term
series
386
is
stem
is
the
to
with
compressed.
the Fibonacci
indicates the number of
possible
attempted
radially symmetric
the so-called
phyllotaxis
13, 21
terms.
to
a
what then
disposed,
I
question
sum
numbers
of the
contact
two
parasti-
numberof orthostichies Schoute’s
R.
VAN
Fig.
serra.
e.
h.
5.
b.
DER
WIJK:
Stems and
Distichous
in h.
and
Eucatagonium politum.
Homaliodendronbeccarianum.
Capsule
of
Acta Botanica
O.
elegans.
f.
i.
Neerlandica
i.
and
capsules
c.
of
different
Mosses
mosses'(
Cyathophorella hookeriana.
Orthorrhynchium elegans.
Capsule
6
Pseudodistichous
(1957)
of
O.
g.
d.
X
6).
a.
Phyllogonium
Thamnium ellipticum.
Orthorrhynchium philippinense.
philippinense.
k.
Neckeropsis
lepineana.
DISTICHOUS
investigations
and
no
Braun,
Fig.
1.
stems.
on
the
AND
have shown
PSEUDODISTICHOUS
that it
the
observing
Phyllotaxis
The
right
figures
to
with
on
those
2
divergences
(A),
the left
with
3
(B),
relate
either
one
5
to
or
these numbers of
are
others, which nearly always
in
occur
in
(C),
or
8
(D)
without
rows
parastichies,
higher plants. Alexander
phyllotaxis
stems
two
387
MOSSES
at
a
much
orthostichies
median
of median
on
leaves,
leaves.
earlier
complanate
the
figures
388
R.
the
date, found
the
VAN
DER
WIJK
numbers of orthostichies in the greater
of
part
same
mosses.
1
Fig.
demonstrates what will
is
respectively
a
complanate
sides
of the
2, 3,
5
the
one;
median
if the number of orthostichies
happen
8, when the radially
or
leaves,
line,
far
so
then shifted
are
symmetric
they
as
either
becomes
stem
found
are
different
on
the left
to
the
1a). The leaves
are
to
or
right.
Leaves
a.
there
is
second
a
dorsal
and
in
direction,
one
b.
the
the
Leaves
twice
that
in several
hookeriana
in
in
would
d.
in
the
Id
(Fig.
on
then
that the
they
call
are
them
of the
this way
the
the
two
two
viz.
the
Cyathophorella
1c). Let
in
of
and
is
that if
is
are
on
the
is
on
are
in
the
side
one
2:3:2:3. Neither
did
nor
I
discover
moss.
In the transitional
dorsal
and
case
ventral
a
one
then leaf
ventrally inserted,
2:3:2:3.
If
median
leaves
If
the
side it is
one
we
measure
consecutive
indeed
near to
1:2:3:2:1
in
Fig.
leaves
each other,
side.
My
not
are
really
eight orthostichies. For
5b
on
there is
is
on
it is
etc.
of Eucata-
either
and
1:2:3:2:1 etc.,
opposite
are
leaves
characteristic ratio varies
very
on
etc.
is
leaves
between the consecutive
This
etc.
1
the
rows,
both sides the ratio of the distances
on
distances
a
in
two
orthostichies
Eucatagonium
in
arranged
leaves,
first
us
orthostichy
Eucatagonium,
complanate
leaves
leaves
leaves
leaves
type of
is realized
consecutive
five
eight orthostichies.
between
if
a
side,
on
one
large
gap
conclusion therefore
distichous,
this
but
that
we
will
reason
pseudodistichous.
That this
arrangement
in other genera of the
that
in
(Fig.
one
the other side
of
If leaf number
1:2:3:2:1
stem
between the
is
in
find that the ratio
side of the
of
on
of median
the distances
we
this
possibility
apparently disposed
between
5b
the other side 3:2:1:2:3
gonium
case
become
of lateral leaves and
(amphigastria)
that with
are
other
any
right).
in
the
this
may
find
not
the other
orthostichies.
none
Fig.
consecutive
sides
sides
two
If
dorsally placed,
the
both
the
on
in
the ratio
absent,
did
two rows
i.e.
1:2:1:2,
with
rows
the
number 5 is
between
five
distances
disposed
two
to
I
the
on
pointing
In
1b).
is found in the median line,
case,
etc.
ratio
agrees
Leaves
are
(Fig.
the distances between the consecutive leaves
arrangement
there
in
the
row
line,
be
leaves
in
novae-hollandiae Brid.
median line
However,
mosses.
inserted
then found
the
all
However,
Fleisch.
2:3:2:3
arrangement
this
the other side.
The lateral leaves
each
the ratio
median
on
transitional
median line.
of the
median leaves
disposed
the
side
5a).
are
are
Rhizogonium
one
5c. shows the
Fig.
(Griff.)
Leaves
consider
and
that
of ventral
with
case
on
complanate
mosses.
row
c.
This
of these orthostichies
one
single
leaves
as
in
the leaves
realized in
(Fig.
leaves
if the
stem
in three orthostichies
disposed
large
as
phyllotaxis
(Fig.
Phyllogonium
viz.
side.
is often
in
seen
complanate
ventral
number of
orthostichies.
two
is
as
possibility,
in the
median line;
in
disposed
inserted,
distichously
Eucatagonium
is
is
also
present
Plagiotheciaceae,
closely
allied
in
the
may be
to
genus
taken
Catagonium,
Catagonium and
as
of
an
indication
which
it
was
DISTICHOUS
originally
ming
much
borne
short
the
shorter
on
a
linear
In
Eucatagonium
nacci
the
8
numbers of
of
was
Eucatagonium
is
the
is
seta
of the
aware
not
the
transferring
correct
and
walls,
pitted
Phyllogonium
in
assu-
whereas
Catagonium,
and
who
its
for
reasons
place is
(sub-)
the
near
Plagiotheciaceae.
I
wish
orthostichies,
the
in
mistake
a
Phyllogoniaceae;
completeness
precisely
accept
made
the
to
in
as
capsule
whereas
seta,
389
the leaf cells in Eucata-
have incrassate
opinion, Fleischer
my
in
Catagonium
For the sake of
to
other
are
genera:
rhombic. The
to
leaves,
pseudodistichous
genus
There
two
long-linear just
leaf cells
well-developed
absent.
or
genus
MOSSES
subgenus.
a
thin-walled and
are
Phyllogonium
are
as
PSEUDODISTICHOUS
close relation between the
a
gonium
in
regarded
AND
main
2.
Fig.
with
that it is
13
or
orthostichies
series)
without
Phyllotaxis
between
gence
add
to
for
the
not
(the
too
we
leaves
Fibonacci
necessary
next
Fibo-
arrive
at
is
a
diver-
the
orthostichies,
consecutive
the
21
137° 29',
angle.
disposition which agrees with the observation. The differences between
the dispositions
obtained in the case of 8 and 13 orthostichies are
small
very
indeed. To
these differences I
phyllotaxis
Fibonacci
stem
angle
135°
of
to
the
is
stem
the
mentioned
a
case
given
in
the
Fig. 2
limiting angle
short
the
leaves
slight shifting,
we
viz■
small
comparatively
the situation
of the
are
we
stem.
observe
1:2:3:2:1.
regular
disposed
a
as
main
the
orthostichies),
remains
have
same
of the
part
considered,
above,
the
nearly
with 8
[i.e.
arrangement
branch,
demonstrate
in the
series,
the
angle, i.e. 137° 29'. The resulting disposition in
is in that
attention
the
with
have
closer
at
slight
Our
along
we
least
Only if
a
in 8
one
a
a
we
much
the
at
orthostichies,
approximation
to
an
our
of
the ratio
is
length
but
the
with
longer part
conclusion
whole
a
so-called
confine
deviation of
final
of
of
complanate
arrive
if
size
case
this
of
:
if
the
if there is
a
limiting angle
390
R.
of the main series.
In this
VAN
it is
case
the number of orthostichies is
Further
that
study of
in several
occur
Plagiotheciaceae
Let
in
5e
Fig.
us
the
different
is
moss
is
also
philippinense
C.
the
(Fig.
s f)
referred
f.
C.
Muell.
now
phyllotaxis
3.
Eucatagonium politum
Natiirl.
Brotherus,
to
seems
The
is short
(Fig.
developed but
of difference
Now in
that O.
have
clearly
under this
name.
perfectly distichous,
C.
philippinense
Muell.;
Reichdt. which
C.
Muell.
was
from
ori-
New
ed.
11).
8
or
really belongs
moss
make
O.
pseu-
565
(fig.
Pflanzf.
arranged in
are
name
are
this
In
improbable.
attention
to
the
more
to
The
rows.
Orthorrhynchium.
search
for
other
capsule of this Orthorrhyn-
nearly absent, and the capsule itself is long and
or
5
The
i).
short
(the
was
unknown. However, the specimen
11944) in New Guinea proved to bear capsules.
seta
difference
of the
But
to
capsule
capsule)
what
Journ. of Bot.
philippinense C.
of O.
and is ovoid.
philippinense C. Muell. will
rhynchium.
be
occurs
which hitherto
by Carr (no
development
leaves
whether this
collected
seta
which
specimens
re-examination
beccari
the
Kindb.
Lac.)
specimens, included under the
totally different (Fig. s h). They
I turned my
discrepancies,
cylindrical
O.
more
species.
are
the
i.e.
dodistichous,
species
to
O.
to
or
Broth.).
several
a
Wils.)
et
et
Orthorrhynchium,
proved
8
the Neckeraceae and
(Hamp.)
but
shown
has
mosses
reality in
ellipticum (Bosch
Muell.,
2,
chium
in
herbaria
Australia.
same
from
The
in
are
Orthorrhynchium
Fig.
question is
higher.
species have been included
moss
from
determine whether
to
even
complanate
genus,
the
elegans (Hook.
described
The other
In
incorrectly
O.
reality
Guinea
Thamnium
region.
The leaves of the first
ginally
5d
philippinense
two
or
families, especially
consider the third
now
Malaysian
this
in
Homaliodendron beccarianum
showed that
it is in
possible
not
13, 21, 34
phyllotaxis
(see Fig.
been named O.
but
WIJK
leaves, i.e. leaves that
pseudodistichous
rows,
and
the
DER
elegans
my
phyllotaxis
sufficiently
have
other
43
in
In
to
and
prove
5 g)
has
these
that
that
in
this
two
a
shape
part
well
points
and
of O.
be removed from the genus Orthor-
genus should
(1905)
(Fig.
opinion
342 A.
Muell. from the
we
transfer
and E.S.
Philippine
the
Gepp
Islands
species?
recognized
and Neckera
DISTICHOUS
Sull.
phyllogonioides
that in Bartram’s
is
named O.
it is
better
part
Neckera
phyllogonioides
1.
2.
a
few remarks
rows
Homaliodendron,
8
rows
Neckera,
be
but
an
the
the 4
or
rows
and
Fig.
4.
between
1:2:1:3:1
Neckeropsis
the
same
this
(Fig.
case
in
8
only if
two
not
the
for
the
name
4
or
consecutive
but
few leaves
did
rows
not
not
are
agree
directed
with the
either there
accessory
leaves
7
(B)
the
(Fig.
is
series
Fig. 5b, which relates
is
also
case
to
on
complanate
arrangement
in
shows that
4
rows
I
11,
have
of the
stems
1:3:3:1
dispositions
Eucatagonium.
rows,
orthostichies
The ratio
or
theory
4
are
1, 3, 4, 7,
of 4
4b).
1:3:1:3
of these
figure clearly
in
that the
in
orthostichies
leaves
Neither
our
key
easily recognizable;
is
rows
a
orthostichies
(A)
a
Neckeropsis.
possibilities;
of the
of 7
with
5k),
Muell.
species
really exist. To solve this problem,
respectively.
etc.
as
do
that
Phyllotaxis
distances
are
disposition
in
species
opinion
Himantocladium.
phyllotaxis is of the first
calculated the
(Fig. 4a)
the leaves
arrangement in 4
There
phyllotaxis.
etc.,
of
difficulty
some
C.
the
reason
another dilemma. In
on
4
arrangement
this
232 this
finds:
in
and then
in
one
in
laterally,
of
philippinense
Leaves
The
is for
Bartr. To my
ex
accept for
to
Leaves
there may
It
(1938)
Sull.
wind up with
the genera of the Neckeraceae
to
of O.
the genus Neckera and
to
us
this
transfer
Philippines
(Sull.) E. G. Britt,
391
MOSSES
conspecific.
are
Mosses of the
phyllogonioides
to
PSEUDODISTICHOUS
from Luzon
present
Let
AND
and
in
occur
the ratio
is
The conclusion
is
a
fiction;
the
392
R.
moss
is
not
tetrastichous,
VAN
but
DER
WIJK
pseudo-tetrastichous;
in
reality
it
is
octostichous.
The
above
considerations
morphological study of
false
the
taxonomic
phyllotaxis
the
may
conclusions and
be
may
serve
phyllotaxis
that
of taxonomic
to
may
at
show
refrain
least
in
that
us
some
importance.
in
mosses
from
moss
a
drawing
families