Download species, and suggested it to affinity because peculiar finely (Fig. 2d

28
BLUMEA
(1982)
151
163
-
a new
Pentastemona,
5-merous
genus
of
Monocotyledons
from
North Sumatra (Stemonaceae)
C.G.G.J. van
Steenis
Summary
A new, short-stemmed
merous
and their characters
Attention is called
is
description
checked
were
the
the first
and
and
the four
seed
artificial
an
the Monocots.
key
the
to
the
Dr.
four
is
genera
Pentastemona sumatrana., there is
given by
J.
of the
that the
be
split
observations
Dr.
Muller.
regular,
5-
discussed
are
on
a
natural
new
family
family
A
up.
is
anatomical
features
P. Baas. A concise account
Besides
for the second
It has
family
for the first time described.
are
not
crystals, by
combination
one new
should
Some
genera.
Stemonaceae.
genera
It is concluded
and
structure,
established, especially concerning
the
in the
The four
genera.
of
or
palynology
aril in
peculiar
characters
given
is described
species
two
among
The fruit and seed of Stichoneuron
contrasted.
for the
vegetative
in
one,
with
genus
flowers, obviously
the
P.
species,
type
new
of
species,
egregia (basionym
Cryptocoryne egregia Schott).
In
the
1959,
of his work towards
course
Dr.
C.
H.
D.
Korthals collection
not
van
belong
den
in
dots
still
/.
the
West
to
the type of C.
a
family, possibly
some
feature
doubt
even
Araceae,
to
based
suggesting that
R.
Liliaceae. Dr.
but
in
Cryptocoryne
1968) because of the
(in
common
genus
egregia Schott (1863),
Sumatra, from this genus,
another
maintained
leaves,
revision of the Araceous
a
rejected
in Central
Araceae, but
to
Brink
raphide
de Wit
C.
on
a
it did
Bakhuizen
occurrence
otherwise
rare
of
in
Monocots.
The smallish
suggested
was
plant,
an
probably already
the type
with
in Schott's time,
so
submitted
was
tified Araceae
and
Dr.
to
among
by
species
As
I
W.
Dr.
would
Meijer
not
remained
examined the
was a new
1955,
Meijer and suggested it
connecting
to
the curved
he did
superficially
describe any floral parts.
not
collection of the
the
probably
interested in the
together
same
type locality.
m
In
1972
with other uniden-
species, hailing
600
at
altitude in
from the
primary
Also in Nicolson's view the
belong
nerves
to
Dr.
species,
1975) which according
thought of that
had also
rosulate leaves
Araceae.
to
distinctly
pollen (in
as
Nicolson, Washington,
Araceae-Philodendroideae. In 1978 Mr. J.
we
±
and
stem
Westcoast of Sumatra, collected
in
belong
Dan
them
well-known Anei canyon,
forest
short rhizomatous
a
habit, but the type (in L) is in bad and almost sterile condition. It
Araceous
to
Bogner
him
because
(Fig. 2d),
of
Muller,
examined the
to come to a
on
my request,
resemblance
specimen
As a
peculiar finely
the
but hesitated
a
Stichoneuron.
Stemonaceae, cf.
affinity
J.
suggested
with
collected
matter
of
by
fact
trabeculate veins
conclusion because of
the sheathing leaf-base and the manifest difference in habit with the other known three
genera
of the
herbs with
a
family, Stemona, mostly
leafy
stem, the leaves
rather insufficient for
accurate
a
climber,
having
no
description.
and
Croomia and Stichoneuron,
sheathing
base.
Meijer's
material
erect
was
also
BLUMEA
152
1978
In
Dr.
W.
and Mrs.
J.
J.
VOL.
O.
species.
canyon
material
flowers
that
the
to
it
same
distinct
but
stem
leafy
erect, rhizomatous
all
Tomlinson &
one
erect
rhizomatous
stem
they
Ridl. and
base.
Stemona
on
occur
(3
generality
most
a
Sav. the leaves
Franch.
Lour,
are
&
they
Gagnepain
one
peculiar
flimsy
the
strengthens
lower
cf.
opposite
are
a
in
scattered
lacking
on
a
a
short,
a
leaf-insertion,
narrow
in Stemonaceae
a
(Fig. lb).
or
or
pauciflora
(BKF32037),
in
stem
a matter
sessilifolia (Miq.)
S.
/. ( Wight 2821).
carrying only
scale-leaves!
1968).
are
the
material
In
Also
from
distichy
of
is in
its
of the
to make
is insufficient
same
(Bl.)
in habit Stemona is
Indochina four
in the
a
Stemona japonica (Bl.) Franch.
in whorls of 3-4-5, and the
moluccana
stem
statement
always distichous,
I feel doubt about the
in Stemona.
in S.
a true
in Stemonaceae. The
Stemonaceae
mentioned, but
Ottelia
a
rare
small,
holds for S.
C.
H.
more
erect
sessilifolia
Wright
variable
species
and in
than
and
S.
mostly
also
an
genera
sumatrana
is the
occurrence
feature in Monocots. It is also known, of
(Hydrocharitaceae),
other plants.
in Butomaceae
types for the four
occur on
Ayensu (1968:
with scale-leaves.
some
e.g.
present in
as
also
Torr. As
the absence of
largely
affinity
In
fine trabeculate venation between
occurs
leaves on the
stem
stem-base in Stichoneuron caudatum
absent, the
the mutual
wings (Fig. lc-d),
texture.
the
agreement for all three genera, and this sustains the
Furthermore,
Burmanniaceae,
Monocots, and
the normal
they
sheathing.
in many sheets in the Leiden
japonica Miq.
nodes
are
be
to
S. pierrei Gagn., and S. minor Hook.
not true
opposite.
of Croomia
because Tomlinson &
feature of the ovary and fruit of Pentastemona
much thicker in
venation
is
recorded
possibly
very
new
and
pairs)
two
viz.,
check whether scale-leaves
kerrii Craib
the leaves
just
certainly
and
(Liliaceae),
The
on
only provided
but distinct
in
course,
instead of
the stem-base of Croomia
satisfactory
Sav.
aphyllous
A
also
the stem-bases
assumed.
of
on
(corrected by Ayensu,
true
on
decision. However, it
tuberosa
in
genera,
(? spirally)
unknown
especially
the stem-base of C.
Ayensu (1968) that
not
other
of Croomia with
that the difference with Pentastemona is
scale-leaves
Anei
this
liquid,
5-merous flowers with
regular,
tepals (mostly
their detailed expose
me to
This induced
Craib normal leaves
latter. The observation
(Miq.)
having
feature
a
scale-leaves
Siebold sheets),
aphylla
Tomlinson &
in
the horizontal rhizome in Croomia
the stem-base of Stemona
There is thus
opinion
4
of the Sumatran
which then lacks
genus,
occurrence
also
They
Franch. & Sav.
&
leaves
However, in
on
the
identify
stem
Stemona.
and
occur
on
having
differs from the
in Stichoneuron and in Stemona,
Herbarium.
In
to
tempted
fig. 35) depict their
of fact
it
stem, the
rooting
sheathing
Thus
the
the others
rather crowded leaves
having
described in full the scale-leaves
Stichoneuron
the
from
specifically
preserved
Ayensu (1968: 273) said that all three genera then known have distichous
without
is
was
three genera in the
leaf-base with wide insertion,
sheathing
leaves
from
(Fig. lc-d),
Also vegetatively
ovary.
different
Gajo Lands,
collections apparently
two
that in the one-celled ovary it would well fit into Stemonaceae, but
manifestly
inferior ovary and fruit
superior
in the
expedition
an
pleasure,
my
material
no
1982
1,
contain both flowers and fruit in excellent condition. In dissecting the
to
appeared
it
differs
to
though distinctly
genus,
Though unfortunately
proved
No.
de Wilde made
North Sumatra, and their material contained,
belonging
28,
Paris
delavayi
Dioscoreaceae there is also
the curved
(Hydrocleis).
of Stemonaceae.
In
nerves
is
not
a
(Franch.
wing,
but
uncommon
in
Fig. 2a-d I have pictured the
C. G.
Fig.
d.
E.
1. Pentaslemona
de
Wilde-Duyfjes
van
Steenis: Pentastemona
sumatrana Steen.
flower, tepals removed,
17010).
G. J.
x
6;
18695).
e.
—
a.
Habit,
ribbed seed with
Penlaslemona
x
4;
153
(Stemonaceae)
b. base
funicular aril,
egregia (Schott)
of leaf
x
25
sheath,
(W.
Steen.
f.
x
1½;
J. J. O.
Flower,
c.
flower,
de Wilde &
x
2
(W.
x
6;
B. E.
Meijer
BLUMEA
154
of
Root-system
273, fig. 35)
This has
gave
bud in the axil of
one
the
in
Descriptions
a
this
are,
are
rhizome
from
of
possible,
not
detailed
are
way
that the
course,
f.
less
of
(1965).
characteristic for
this
it remains
Concluding,
erect stem
be studied
to
in
Fruit
and
seed
incompletely
Small
(1933),
Ohwi
species;
surprising
who
is
wrote
the
finally
Croomia
is
thinner than
In
Stemona
cf.
kerrii Craib
(? storage)
of
roots
Pentastemona
5-10
c.
long,
cm
seem
egregia
without any
in
Croomia japonica,
in
The fruit of Stemonaceae
—
were
and
finally
Japanese species
very
of
plate
published
occur.
as
In
undescribed.
yet
as
stated and figured
2-valved in the American
of Croomia. This is rather
thin.
pericarp,
in that genus,
the
dehiscent remains unknown yet, but
as
many
field whether
superior
with about the size of thatin
seeds
a
thin
expected
to
it.
shimmering through
pericarp
shown in Croomia. In view of the 3-zoned
fruit of Pentastemona, if dehiscent, could be
is
obviously
parietal ovules,
no
the
be 3-valved, instead of 2-valved
in the other genera.
In Stemona the seed
'aril'
consisting of
(1964),
error
in
a
(1959),
dangles
account
studying only
hollow vesicular
also
obviously
mistaking
funicle and it is
studying
by
generally
Krause
described
(1930:
of Stemona, has shown that this is
dry
material in
body, bulging
and
the
irregular
dried material,
herbarium.
figured
a
seed
having
obviously
a
the
Actually,
in outline. In
as
an
fig. 30D). Swamy
passing,
serious
aril is
a
Hutchinson
of Stemona with the aril,
the latter for 'attached abortive ovules'.
Unfortunately
Tomlinson & Ayensu
Croomia, which they declare
Japanese species
which
long
on a
'tuft of hairs'. Thus it is pictured
accurate
very
caused by
complex
a
certainly
Fortunately
I
as
being
of Croomia Ohwi
reminds
had
one
of
the
(1968:
273)
'filamentous'
(1965: 279)
situation
collection with
a
in
ripe
did
not
against
calls it
a
describe
'tuft of
Stemona, by
or
figure
the aril in
vesicular in Stemona.
the
fleshy
use
fruit of C. japonica
setose
of the
For
the
appendages'
term
'fleshy
.
as
fleshy,
In
roots.
P. sumatrana.
the fruit is inferior and the
still
for dehiscence,
obstacle
in
the ones there
the excellent
on
2-valved capsule. So it is in Croomia,
that the fruit is
From
large number of thickened,
a
Stemonaceae.
confirmed this for the
plant
almost
Whether it is
superior,
pericarp
In the Sumatran
Croomia,
a
(1965)
the
as
the
rely
to
known; in Stichoneuron fruit and seed
Stemona the fruit is
by
of
genera.
in the other genera.
collected. The latter is anyway
much thickened
Stichoneuron and in Stemona subterranean stolons
was
have
is thick,
from
long, thick roots, but it is
many
not
complete plants
in
other
root-system, and
emitting
very
In the
is also found
situation
we
the
present
rhizome attached.
These
Stemona.
the
recently by Meijer,
stolons;
same.
for
rhizome with
erect
creeping
a
was
Torr.
I found in the Leiden Herbarium
japonica Miq.
short rhizome
a
produced sympodially
mode is also
a
(1968:
pauciflora
continue the rhizome.
to
short rhizome with
short
a
is
available
not
growth
In Stichoneuron
pictured
observes
one
a
Tomlinson & Ayensu
—
stem
the
horizontal part
long
the situation is the
(BKF32037)
collected
with
roots, without
spindle-shaped
more or
he
(1888);
Stemona pictures
trace
In Croomia
provided
all
mentioned by Ohwi
Hooker
which
few specimens in the herbarium with
specimens,
1982
1,
scale-leaf, leaving another bud
mostly incomplete.
are
three
rather
No.
of the root-system in Croomia
herbarium material I doubt whether this
There
28,
Stemonaceae.
ample description
horizontal
long
a
an
VOL.
Miq. ( Hatusima 19946,
C. G.
from the
Ryukyu
matching
arillar
G. J.
and
Islands)
structures
Steenis:
van
to
Pentastemona
satisfaction
my
In the Sumatran plant the seed is of the
and Croomia,
that
matching
exactly
with
ellipsoid,
contained
and
shape
same
41
(1964: fig.
&
is
occurrence
excellent argument for
an
Of Stichoneuron the fruit and seed
the herbarium (as well
flowers),
as
seed
in
present
some
are
diagnosis
when
runs
follows: Fruit
as
1-seeded
11
c.
articulation of the
Funicle
flimsy
to
up
3 mm,
c.
made
(21-1-1947)
5-6
by
very
the
the apex caudate for
and wide-celled arillar, free,
halfway (c. 3-4
ridges appearing almost
This
confirms
Stemonaceae,
the
as
and
uniformity
the
a
seed
c.
16
stalk
4
mm,
the
very thin.
Pericarp
the base of the seed
long. Fig.
mm
in
structure
all
four
2e-f.
genera of
of their insertion, basal in Stemona and Croomia,
irrespective
by
above
sharply ridged lengthwise, the
in the herbarium, 5-7
of
number
5-6 narrow-lanceolate, very
c.
broad-ellipsoid,
wings
narrow
distinct
the seed with
to
in
rare
Leiden. In
second
when 2-seeded
remains
few millimeters.
a
at
boiled material the
equal appendages surrounding
Seed
long).
mm
±
From
description.
supported by tepal
apex close
at
winged,
detailed
glabrous, spindle-shaped,
mm,
at
pedicel,
a
extremely
fruits, respectively Ridley
14582, Corner SF 20242 (in spirit, unlocalized), and SF 30503. Of
Holttum has
and
This
(Fig. le).
the eight sheets
on
the Singapore Herbarium there were three of the 15 with
size
a structure
bulging
Croomia.
and
undescribed, and they
being
none
with
in Stemona,
with Stemona
affinity
were
with exactly
that in Stemona
provided
42)
the
hollow, but less coralloid and almost exceeding
inflated,
seeds
two
structure as
At its base it is
lengthwise ridges.
pictured by Swamy
this
for Stemona.
figured by Swamy
as
155
(Stemonaceae)
in
parietal
Pentastemona, and apical in Stichoneuron.
On
of the above
account
I have
Sumatran plant,
of the
the
family
Climbers
short,
or
erect,
curved
with
leafy,
or
lateral
stems
nerves
a
tepals
sessile,
a
raphide
and in Stichoneuron
or
5-merous with 5
whorl, free
free
or
enlarged apically
sulcate with
with
with
a
on
or
or
a
connate
shallow
stem
and
connected
an
by
dots, occurring
on
basal, anatropous,
with
or
2
new
genus
circumscription
with scale-leaves,
with
numerous
also
or
Tepals persistent,
inside
a narrow,
staminal tube,
surface.
many, either
sheathing
intramarginal vein,
of
in
not
at
on
pedicelled,
or
epitepalous,
a
the
on
a
connective
filament
apex or
2-valved
in 3
or
or
mono-
inferior,
parietal
(? always).
4
or
sometimes
Pollen inaperturate
from the
capsule,
2-merous
imbricate, 4-5. Stamens
shallow ring. Anthers
pendent
In-
the leaves
base. Inflorescences
Ovary superior, semi-inferior,
integuments. Fruit
parenchy-
(probably cincinnate) pauci-
valvate
latrorse.
a
petiole.
other organs.
Pentastemona
connate
in
in
or
ascending
and
crossbar veins;
sometimes
nodes and internodes. Flowers
tepals, whether
reticulate
or
features of the
distinct
(sometimes horizontal) scaly
a
subrosulate leaves
widened between the loculi,
granular
a
of the
provided
rarely solitary flowers, mostly
1-celled. Ovules either few
or
or
mostly distichous, with
florous bracteate inflorescences.
5 in
developed stem,
trabeculately
and/or
peculiar
represents
widening
a
stem, which is in the basal part
rhizomatous
the
axillary, peduncled,
or
causing
mostly absent, but present (as uniseriate hairs)
fruit ribs,
with 4
on
styloids
dumentum
leafy
of
description
that the latter
follows:
as
herbs, eitherwith
erect
Leaves entire,
ma
discussion and
concluded
Stemonaceae. Pentastemona,
family (cf. Krause, 1930)
rhizome and
or
given
definitely
zones,
Seeds
on
a
156
BLUMEA
2. Venation
Fig.
Franch.
&
egregia (Schott)
Fruit
seed
genera
genera
of
1,
Stemonaceae,
all
japonica Miq.;
of Stichoneuron
funicular
arillar
of Stemonaceae:
and two
the four
Croomia
No.
c.
28,
1982
x
Stichoneuron
2/3. — a.
Stemona
caudatum
sessilifolia (Miq.)
d.
Ridl.;
Pentastemona
Steen.
and seed
with
types in
b.
Sav.;
VOL.
common
with
lobes,
Ste
=
caudatum
8
x
Stemona,
Pentastemona
;
Ridl.
( Corner
with
—
SF
five
e.
Two
30503)-,
fruits,
g.
1- and
Scheme
unique characters,
similar ratios for Cr
Croomia,
=
2-seeded,
of affinities
common
two
Sti
=
f. ribbed
4;
x
the
among
with
four
Croomia,
Stichoneuron,
and Pe
=
Pentastemona.
short
or
long
funicle, broad-ellipsoid,
inflated, bulging
Stemonaceae,
274),
who knew
assemblage.
with
this
grooved
coralloid or lobed hollow aril;
only
a
natural
or
family.
three genera, stated that 'It is
Similarities
between
limited evidence
it
is
the
genera
clear
ribbed,
that
are
we
at
the
base with
endosperm copious; embryo
few
are
—
not
Tomlinson
evident that
but
not
&
dealing
form
many',
with
very
fleshy,
small.
Ayensu (1968:
they
differences
a
a
a
natural
and 'Even
homogeneous
C.
This
assemblage.'
also
I
always
genera have in
based
mostly
can
the
van Steenis:
on
only partly
Pentastemona
the
characters and
weighing
The conclusion
anatomy and
vegetative
endorse their
of
matter
a
common.
Croomia and Stichoneuron
opinion:
,
while all four genera share
characteristic aril.
that
taxonomy,
not
genetically,
To
me
taxonomic
well
apical,
a
superior
ovary
in
There is
a
at
the
of the
weight,
or
unique
ovaries
several
duced
roots
connective
basal
flowers;
Shares:
with
to
parietal
Moraceae
distinguish
do not
hold, e.g.
inferior ovary
an
is,
is, for all four
in the
Leguminous genus Archidendron.
course,
spindle-shaped thickened; flagellate-pro-
appendage;
long funicles;
very
ratio
pro
large
placentation.
Croomia
basal
often
stemona
this
a
in
example:
mostly scandent;
Unique:
though
even
characters sometimes
in the
with
essential thesis
structure
to
example
that of
to
structure
between the four genera, each of which, of
affinity
features. For
was
very similar in
varies from basal
placentation
level is used for
family important
one
distinct reticulate
in addition
Stemona:
illustrative
Whereas the peculiar seed
family
what
Ayensu
similar
similar seed
absolute
no
carry
&
are
extremely
the Rubiaceous genus Mastixiodendron and
Theaceous genus Anneslea,
possesses
characters
exactly matching,
feature which
and Urticaceae. Even within
a
situation appears
comprehensible.
genera of Stemonaceae,
and
the
are
surprisingly
a
considering
of Tomlinson
morphology.
habit and the venation and gross anatomy of the leaves
Pentastemona
157
(Stemonaceae)
be said of many families, however,
Olacaceae. And what does
e.g.
can
mean? It is
'homogeneous'
character-set
G. G. J.
curved
nerves
ovules
numerous
and
and
dehiscent
the
seeds;
with Penta-
fruit;
shallow
staminal
tube.
Unique:
Croomia:
a
horizontal
not yet
Shares:
rhizome
reported
with Stichoneuron
the
inflorescence,
and the
basal
Stichoneuron:
Shares:
reticulate
curved
valvate
Unique:
the
sheathing
genera
habit;
few,
exine
and
thick
ovules
of the
structure
dehiscent
dense, apically
unlikely
not
the short
pendent
(subterranean
bracts
but
pollen;
crowded
Pentastemona
lateral
broad
in
nerves
separated by
a
Croomia the
simple inflorescence;
ovules
ture of the
5-merous
sile;
the
and
connective;
the
with
simple
Stemona
the
fruit.
raceme
lobes, appressed
lanceolate
stolons
occur).
apical placentation,
5-6
free,
to
filaments;
with
pedicels spirally arranged;
with
Unique:
the
nerves
tepals;
with
other
setaceous, articulate
few
Pentastemona:
in
to lower
part
presence
the
with
aril
stiff,
of
c.
half of seed.
above
the
base;
of uniseriate
habit;short thick
seeds; apical placentation;
erect,
consisting
reticulate
thecae
hairs;
with
filaments;
exine
struc-
pollen.
flowers; mostly
leaf-base
sheathing;
fruit, winged in
compound inflorescences;
no
leafy
stem
proper; ovary
P. sumatrana; ovules
in 3
anthers
inferior,
parietal zones.
ses-
as
is
Pollen
inaperturate.
Shares:
with
Stichoneuron
by connective;
and
seeds;
the
suprabasal
uniseriate
hairs;
stamens inserted
on
lateral
with
a
nerves;
Stemona
shallow
the
thecae
staminal
Counting the characters mentioned the diagram (Fig. 2g) illustrates the
these characters,
unique
separated
numerous
ring-like
ovules
tube.
occurrence
and shared. From this it appears that Stemona, and
even more
of
so
Pentastemona, stand apart from the Croomia/Stichoneuron pair. And furthermore, that
Stemona and
especially
by pollen morphology.
Pentastemona have the
most
unique
characters. This is confirmed
158
BLUMEA —VOL.
Nakai
with
(1937)
inflate
to
tendency
Pentastemona, I have
no
characters would
hold.
I
feel
not
doubt that this
anatomical
celled ovary, and the
all four genera, whatever
least
of
Affinity
and
sense.
this is
have
the
with
Liliaceae
but
then
3-
regular
merous
small
family
(1960),
but
wise
who
the
course
in
an
propose
exaggeration,
conclusion.
the
is
accepted,
though
affinity
of
the
the
origin
of the most
a
of Stemonaceae
a
'to
of Dioscoreaceae',
features of the
striking
paragraph
in
in
possibly
with Dioscoreaceae
neighbourhood
One
affinities
This remains still obscure. As
suggested
general
of
subject
favours also
grounds Ayensu
of
5-merous
cq. 6-merous
regular 3-merous,
represented by
to
affinity
Proto-Liliales in
vague,
The
—
by Ayensu (1968: 164).
of Burkill
occurrence
Monocots
occur,
a
one-
exactly matching
should be devoted
to
of 5-merousness in Monocots.
occurrence
in
view
from the
seems a rather
The
viz., the
the
anatropous ovules,
seed structure,
peculiar
genus Pentastemona are the 5-merous flowers and
new
the
the
On anatomical
also
been
which
the
still find this in such
Stemonaceae.
traditional systems,
remote
I
the above
as
useful.
not
Stemonaceae has been treated
the
family rank,
to
may differ in placentation. One could of
they
three tribes for the four genera, but
at
in the
has small
the four genera together,
vegetative characters,
striking similarity
Croomial/
pair
If he had known
apical placentation.
hold
the characters which
impressed by
more
morphological and
the
raise
that the
ground
then also raised
was
to
proposed
the
on
anthers and
flowers, short filaments, unappendaged
1982
1,
ranks
family rank, Croomiaceae,
to
pair
Stichoneuron
his
No.
28,
the
flowers
staminode,
a
flowers
bisymmetrical
are
as is found
6-merous flowers.
or
flowers
in
and
in
anther
is
either reduced
Zingiberaceae, Lowiaceae, etc.,
Besides,
quite
some
—
In
Flowers with 5 anthers
prevail.
one
Monocots.
Monocots have
flowers. Therefore, Pentastemona with its 5-merous flower is
or
but this occurs
regular
an
quite
2-
or
4-
exception
the rule.
Mr.
with 4
J.
Bogner
or
6
tepals,
to me
wrote
tepals. This genus belongs
632)
that
on
the
Another instance
3-
mentioned
to
the
Urospatha (Araceae)I
flowers
Araceae tribe Lasieae, for which Hutchinson
or
to
is
found in
less
6-partite,
have 5
Sciaphila
often
6 segments.
to
females
tepals
are
also
usually
rarely
4-
or
occur.
of the Triuridaceae,
4-5-partite.
Bth.: the
6-merous, but he
The
plate
The
occur
(1959:
especially
in
was
the
Franch., where flowers may vary
pictured such
615, fig. 381).
a
5-merous
In
flower in
the perianth
of S.
wrote
copied by
section
from
Paris
4-
is
are
3: 313,
t.
that 5-merous
female flowers with 5
occurs
i
Euthyria
to
(3,
Bl.
6-merous, and also the
Hutchinson
(1959: 547, fig. 348).
in the genus Paris
Franch.
subsection
5-merous. Krause
polyphylla
nana
are 4- or
males
similar instance of occasional 5-merous flowers
Parideae),
where
female flowers
Martius described in Flora Brasiliensis
116, ii) the flowers of Sciaphila albescens
tribe
of
also mentions that the ovary is 4—5-locular.
usually
A
spadix
but that between these he had also observed occasional flowers with 5
J. Sm.
and
the Leiden Herbarium I found the flowers
to
so
(1930:
did
vary
(Liliaceae,
Submuticae
374, fig.
Hutchinson
in this
152F)
(1959:
species
from
G.
C.
G. J.
van
3-4-5-6-7-8-merous,without
genus.
Even
in
but
geographic
a
collection
one
Steenis:
specimens
Also in the genus Trillium of the
of
merousness
monographic
the
of the
study
North American
of variations and noted
occurrence
cases
commented, however, that all of them were single
merousness
and
The situation
that
they
were
similar
seems
In
and
all
only
the
occasional
flowers
in
Anatomy.
the
—
Dr.
egregia (Meijer3214):
crystals
cell
epidermis
abundant
walls
related
concluded
Stemonaceae
to
and
Meijer
that
the
also
that
crystals
in the leaves in
styloids
and
were
raphides
conglomerates
±
undefined
In
crystals,
Pollen
examined
densely
S.
by
J.
covered
by
in
Ridl.
the
5-merous. He
3-merous
where
populations.
sometimes
a
,
a
of
range
single
extremely rarely,
greater variability.
I
5-merous,
can
only
As
conclude
which is normally regular 5-merous.
on
the anatomy of
straight;
stomatal
minute
Also
de
to
irregularly
Pentastemona
complex tetracytic;
druses
Wilde-Duyfjes
tetracytic
minute
of
styloids
stomata
conjunction
Stemona.
are
throughout
cyclocytic;
the
anticlinal
18532):
crystals
shaped, solitary
and
is
raphides
different. He
with those
He
related
to
as
crystals.
suggestive
checked
also
of
the
in Pentastemona and the
confirmed
raphide
raphide
Muller
minute
and
Baas
needle-shaped
+
that in
Croomia the leaf
found
f.
in
the
leaves
diamond-shaped
bundles but certainly
membranaceum Hk.
but also
10082)
(Kerr
('clustered
morphology.
Dr.
in his
to
(no styloids).
Stichoneuron caudatum
absent.
in
in 1980
complex
as
the
of
shaped crystals')
to me
combination
but
of
crystalline
in the
3214.
occurrence
only contains raphides
not
specimens which deviated in
Tulipa
are
(de Wilde &
occurrence
In
species
raphides.
stomatal
raphides,
Dioscorea,
or
or
of Monocots
reported
sumatrana
straight;
occurs
69-83)
were
found in Monocots, but
are
specimens,
genus
styloids
and
styloids
Doubtlessly
Baas
P. Baas
Pentastemona
mesophyll.
odd
anticlinal epidermis cell walls
abundant
as
flowers
to
first
is
occurs.
Pentastemona
two
seems the
that it
tentatively
confined
or
in the
Japan.
large appendix
a
large, normally
to, for example, that in
5-merous
conclusion, regular
variability
to
(1917:
where flowers
individual
often found amidst
individual with 2-merous flowers
Himalaya
devoted
species,
number of
a
the
3-merous. Gates
usually
are
merousness.
from
range
This
tribe of the Liliaceae variation
same
which
flowers,
vary in
may
wide
its
159
(Stemonaceae)
of the 4-merousness, which is usual
preponderance
throughout
occurs
Pentastemona
( Koelz 5835)
not
identical,
he
found the
+
irregular
irregularly
while
styloids
same
rather
bundles.
—
found
granules.
The
pollen
to be
of both
inaperturate
Similar
pollen
species
with
grains
a
of Pentastemona
was
very thin exine which is
occur
in Araceae-Philoden-
droideae.
In
contrast,
structure
and S.
J.
the
pollen of Stemona, Croomia and Stichoneuron is monosulcate. The fine
of the exine of Stemona australiana
tuberosa Lour,
J. Smith
Croomia
it is
is
comparable
characterized
by
to
Wright,
S. japonica
(Bl.)
Franch. & Sav.,
that of Pentastemona, but of Stemona
the presence of
a
thin
layer
gloriosa
of columellae.
and Stichoneuron clearly differ from both Stemona and Pentastemona in their
reticulate exine
structure.
BLUMEA
VOL.
28,
1982
1,
No.
160
ARTIFICIAL KEY TO THE
absent, leaves crowded
stem
Proper leafy
la.
ciliate
sheathing petioles
Proper leafy
b.
2a.
by
stem
Roots
stem, with
inserted
in
3 parietal
Pentastemona
zones
not
winged
usually spindle-shaped thickened. Anthers with
connective. Ovules
produced
rhizomatous
erect
present. Flowers 2-4-merous. Fruit superior,
scandent.
Usually
short,
a
uniseriate hairs. Flowers 5-merous. Fruit inferior,
coarse
Ovules and seeds many,
winged.
on
GENERA
numerous,
basal.
Seeds
funicle.
long
on
Leaves
vinerved
b.
Small
cur-
Stemona
herbs with
erect
without
2
....
flagellate
a
leafy
Roots
stem.
short
on
appendage,
thick
thickened.
spindle-shaped
not
filaments.
Ovules
and
seeds
Anthers
few,
apically
inserted
3a.
3
Leaves
curvinerved. Flowers if
articulated
Anther-cells
pedicels.
than
more
spaced,
one
not
parallel,
separated
not
nodding,
few,
by
thick
a
on
stiff,
connective
Croomia
b.
Leaves
with
lateral
suprabasal
stiff, articulated pedicels
erect,
rated
by
Flowers
nerves.
at
apex of
Caulis
brevis,
rectus,
robustis
foliis
rhizomatosus,
ciliata;
lamina
ovalis,
marginem apicemque
parallelis
trabeculatis
ornatum.
in
3
5
confluentia.
Hypanthium
5
locum
embryo
insertionis
minutus.
Note:
though
a
—
It is
T
y
p
u
s:
ellipsoidea,
a
racemo
the
It
is, of
specimens
course,
are
not
small and
not
in
5-meri,
subglobosae,
arcuatis
densis
ad
racemos
specie typica
breviter
thecis
2
parallelis
zonis
hippocrepiformi formatis,
tribus
alis
3
brevia
in
eorum
10
(vel
foliis
pedicellati
(et fructus) inferum, 1-loculare,
vesiculari
basi
ovula
in
zonis
cum
conicum
stylum
transgredientibus
longitudinalibus
instructo.
ipsis
membranaceis
funiculus
brevis,
Endospermium
arillo
magnum;
Steen
new
that
omnibus
annularis, sub-5-lobatus,
longitudinalibus ornata;
remarkable
impossible
2-3
ultimo
breviter
sumatrana
pilisque
imbricata, persistentia, punctis
genus is found in Sumatra,
large island, is manifestly poorer in endemic genera
and Borneo.
Flores
antherae
specie typica
lobato
petiolata,
numerosis
(vel venis)
bracteato.
Ovarium
costulis
Pentastemona
that
noteworthy
sepa-
pinnatis 3-4-paribus
staminum
modo
Folia
instructa
3-5-paribus,
nervis
connata,
primariis instructum,
in
nervis
ultro
alternantibus; stigmata
irregulariter
magno,
Tubus
obtectus.
margine hyalina
bisexuales, regulariter
numerosis
inferus, pericarpio membranaceo,
late
se
dispositi,
instructus;
visibilibus
vasculis
on
liberis; mesophyllum punctis rhaphidum
libera vel basi
conspicuo separatis.
ovulis
inter
haud
densos
obsita.
sessilibus
bene
numerosa,
ultro
elongato,
sat
Flores
minus
numerosis
lata
spatha
intramarginali,
venularum
ternos
longitudinalibus
brevibus
instructo. Semina
areas;
3
epitepalis
longitudinalibus,
stigmatosis
Fructus
ad
prolifero.
et
latrorsis connectivo
parietalibus
usque
membranacea, plus
staminibus
lateralibus
lobis
solitarios
instructo
sicco
rhaphidum nervisque
intus
divergent,
1.
Fig.
—
ad laminae mediam
usque
ascendentibus,
conjunctis, apicibus
terminales
reductis
Tepala 5,
basi
nervo
Inflorescentia axillaris, pedunculo
cincinnos?)
paucis
laminae
nov.
gen.
radicibusque
probabiliter spiraliter aggregata,
uniseriatis
raceme
a
Stichoneuron
paullo suprabasalibus aliisque superioribus
ad
in
thick connective
a
sparsa,
Anther-cells
peduncle.
PENTASTEMONA Steen.,
integra,
crowded
numerous,
as
compared
which,
with Malaya
Pentastemona may be found in the latter
conspicuous,
and
may have
escaped
the attention
of collectors.
The beautiful
new
the seed, viz. that it
collection of
seems
in dried condition escape
to
Meijer of
possess
a
P.
egregia
showed
a
remarkable character of
distinct, proportionally thick sarcotesta, which will
observation. It
prevails in the family; field observations
would be
must
give
interesting
the solution.
to
A
know
whether
peculiarity
is
this
further
C. G.
the
of 5 rather
occurrence
staminal
G. J.
This
ring.
Stef.nis: Pentastemona
van
in the
deep pouches
observed in both
was
KEY TO THE
Flowers
in branched
at
racemes,
6-9
fresh,
(in
5-5.5
emarginate
cm,
when dried,
sicco
in
connate
a
cm
lata,
interdum
paribus;
petioli
subtus
10-30
Inflorescentiae
multinervosis
2-5
2-3
articulati,
longitudinalibus
in
mm
tenuiter
8-10
southern
SUMATRA.
lowland
rain
sicco
Alas
diam.
small stream,
a
dorso
1
green,
basalt
young
bracts
Semina
18532
t
genus,
e
s:
of the
J.
J.
m
Alas
(holo
Lands,
c.
late
on
rich
soil
ad
cm
ovata vel
longa, 6-9
Pedunculi
10-30
nervis
2.5
c.
haud
20, sessilia,
B.
1
c.
iso in
A, BO, K).
3°
N,
97°50'
E,
E. de
in
a
very
3
c.
longa,
G. Leuser Nature
rock,
vivo,
altum, alls
mm
mm
50 km south of Kota
basalt
in
Pedicelli
10
mm
costulis
irregulariter
E.
L,
c.
obsiti.
dehiscens,
magno
de Wilde &
1-nerviae,
longitudinalibus, disperse
ut videtur
arillo
foliis
apice
longi,
cm
uniseriatis
cylindricum,
c.
interdum
oblongo-lanceolatae,
in
over
4-5-
densius
marginem spathae
robustis
basi
O.
tributary Bengkong R.,
alt.,
membranacea, late
praebentes,
pilis
ornata,
J.
fringed
egregia
lobato
Wilde-Duyfjes
Reserve,
Alas
Tjane, in primary
shaded
damp place
~
O.
de Wilde &
white. Same
B.
place,
E.
E. de
Wilde-Duyfjes 18695, phnt jiiicy, bright
about 1 km further on,
herb,
bracts green, flowers
from
the
inflorescences,
on
damp shaded slope,
5-merous, tepals white,
W.
J.
J.
O.
de
Wilde
&
anthers
B.
E.
among
yellow,
E.
de
often
Wilde-
(BO, L, US).
1 have chosen
Pentastemona
because the material,
specimens
2-3
longae.
mm
J.
petiole
1.
Fig.
cordata, 9—13(—16)
bracteae
ellipsoidea
W.
Typus :
the
long, halfway
mm
suprabasalibus) 3-4-, superioribus
seriiformis,
subgranulosis
by
papyraceous
10 strong ribs,
—
statu sicco
basi
7-9
2. P.
nov.
disperse,
distanter
Fructus
North Sumatra
50-125
corolla
green
plants developed
N o
when
green,
rocks, juicy
Duyfjes
with
oblong,
instructa, mesophyllo punctis rhaphidum
et
Ovarium
longa.
membranaceo.
Lands,
c.
6-9
angulos
latis.
connate
Leaves
sheath of
11-13
c.
cm.
glabrous,
the
only
sp.
racemulos
pilosae,
ad
mm
mm
oleosum.—
W.
Folia
instructae;
~
along
3-5
c.
apex,
Fruit
acuminata,
uniseriatis
robustis
plani,
ornata, 3
the mouth
forest at
altus.
cm
breviter
longae, apice
cm
North: southern
near
Steen.,
uniseriatis
obscurioribus
albido; endospermium
valley,
recurved.
mm,
(basalibus
cm
1-4
membranaceis
longitudinalibus
River
cystoliths;
longi. Tepala obovato-oblonga, albida,
albidis
altus, pericarpio
,
at
Tepals thick,
pilis
nervos
mediano
nervo
punctis rhaphidum
18695
acuminate
inferioribus
longi, pilis
cm
videtur, 3-4-angulati,
haud
well
sumatrana
caulis 5-10
ad
nervis
ornato,
6-20
membranaceae, in
ut
long; peduncle
mm
cm.
9
by
subinaequilateralibus,
glabra,
supra
dispersis
obsitis.
6
Pentastemona
aquifero-membranacea;
tenuibus
reductis
lobes
long, thin, oblong,
often antler-like branched, hair-like emergences
1.
Planta
Petiole 4-15
thick tube,
by
beneath and petioles
nerves
mm
in the axil of uninerved, sessile,
racemes,
dotted with punctate
hairs.
3
9-13
apex, very watery when
at
1. P. sumatrana
rather
base,
at
Tepals
cm.
pauciflorous
10-30 cm. Leaves
c.
wings
bracts 5-10
scarious)
densely
uniseriate
by short,
elliptica,
they represent
apical, spaced,
spaced oblong cystoliths;
condensed
pauciflorous
of 2-3
leaves; peduncle
base, very short-acuminate
at
entire
filmy,
foliaceous
with
and the
base
style
that
likely
seems
SPECIES
consisting
reduced
uniseriate hairs. Petiole 6-20
Fruit with
Flowers in
1.
cordate
2-5
when dried, with
filmy
spaced
at base.
inflorescences,
often with
apex
distinctly
cm,
with
and it
structures.
nectarial
1.
between the
receptacle
species
161
(Stemonaceae)
the
fruiting
though only
inflorescences
are
sumatrana as
crowned
inflorescences sag, their
the type of this
available in the dried
tips
by
state,
reduced leaves,
may form
new
is
new,
remarkable
complete.
In some
resembling proliferation;
plants
in
reaching
the soil.
A
BLUMEA
162
similar proliferation is shown in the
cf. Fl.
I, 5: 326, fig. 5. The
Males.
missed
possibly
in P.
sumatrana,
VOL.
1982
1,
Echinodorus
Malayan
of
occurrence
of which
No.
28,
I had
dried
only
(Alismataceae),
Steen.
ridleyi
observed in
a sarcotesta
material.
P.
egregia
This calls
I
have
for further
field observation.
2.
Cryptocoryne egregia Schott,
Stem
5-10
cm,
in
long
many
Ann.
Miq.,
terete, thick,
petioles, emitting
comb.
egregia (Schott) Steen.,
Pentastemona
fleshy,
roots;
Mus. Bot.
1
c.
glabrous, broad-ovate, the base shallowly
1
Lugd.-Bat.
0, with
cm
Fig.
—
122.
(1863)
Leaves
distinctly
or
acuminate, beneath densely dotted by punctate raphide dots, 7-17 by 5-10
the base 3-4
4-6(-15)
5
crowded,
c.
6
little
perianth,
fringed
c.
halfway in
5
a
the
in
appendage,
the
axil
Pedicel
c.
at
3-4
apex.
mm
less
separated.
sometimes antler-like branched,
thickish,
1.5
c.
seed. Seeds almost
globular,
transparent sarcotesta
Between the
high.
mm
Ovules
pouches (nectaries?).
very
c.
through
1.5
by
1.25 mm, 1
which the ribbed
13
whitish,
or
very
low
when
staminal
the
not
5
ring
distinct
all developing into
with
dry, glossy,
shimmers
are
Style short,
emergences.
and the staminal
endocarp
fleshy
shorter than
with 10 firm ribs which
mm,
hair-like
style
mm
yellow
Ovary
the three placentas,
on
many
lanceolate, 1-nerved
pale greenish
to c.
enlarging
a
lowest
scarious,
long, the lobes roundish, imbricate,
Stamens 5, inside a
or
filiform,
raphide-dotted,
in the axil of
fleshy,
uniseri-
one
below the
long
mm
at
coarse
flattened, with
1.5
c.
acute-
cm; nerves
(in life); petiole
fringed-hairy by
uninerved,
mm,
13
c.
more
in fruit
of
and
tube, in all
thecae
high, fleshy,
mm
and
somewhat
long,
setaceous
above
shallowly depressed
margin hyaline
cm
wide, recurved
mm
short,
by
cylindric,
2-3
4-6 mm.
by
receptacle,
the
on
sheath
3-4
c.
all
pairs,
Perianth rather thick
over
long, 7-8
mm
tube
5-12
long.
mm
connate a
the
glabrous,
downwards curved
Flowers
1-nerved bracts
bract
terete,
Peduncle
sometimes
bract.
long,
cm
hairs.
ate
up 3—4
pairs, higher
(in sicco),
papyraceous
apex short
emarginate,
2d.
1f,
scarious bracts between the
some
of stolons.
trace
no
nov.
a
rather thick
than
through; larger
the vesicular arilloid.
SUMATRA.
canyon,
locality,
c.
600
W.
West
W.
alt.,
Korlhals
Central:
Meijer's
other
two
s.n.,
holotype
Meijer 3214, flowering 22 April 1955,
Meijer 17010, flowering and
Notes:
poor the
m
new
collection has
collections
material, realizing that in
fruiting
some
points
have
the
1981,
so
much that it will
base of the midrib and
its
own
was
P.
small roots,
loose leaves
obviously
a
means
escape
sent
of
ditto,
Anei
forest;
same
898-88.409;
in primary
slope
material
liquid).
in
the
description
with sizes and
occurrence
observation. The
had
produced
a
with
texture
of
a
apex
this
new
will exceed
sarcotesta; when
of the
pedicel
and
small adventive shoot with
vegetative reproduction;
in
P.
sumatrana
this
observed in the inflorescence.
egregia
flowers in
There is
viz.
some
easily
also
amplified
comparison
those of herbarium material. A marked character is the
dried it shrinks
sub
hill
clarified many points and has shown how
now
I
were.
8 Nov.
L
in
on
a
is
distinctly
more
different from P.
simple
a rather
Pentaphragma,
striking
of the
sumatrana,
being
inflorescence, the ribbed ovary,
resemblance
in habit with
Campanulaceae
or
a
much
coarser,
much
larger
plant
genus,
etc.
completely
Pentaphragmataceae,
different
and the
two
could be
C.
found
for P.
in the
G. G.
by the mostly asymmetric,
which
do
not
the
raphid dots,
can
easily
from the
pairs
midrib,
valid
especially
be
distinguished
the reticulate
sheathing petiole base, the absence of
not
the
inflorescence, the stalked anthers,
boragoid
the celled ovary with axile
the
Pentaphragma
leaf-base, the mostly toothed leaf margin, the lateral
in decussate
emerge
163
(Stemonaceae)
forest bottom in loose earth, and this is
cuneate
venation, the
trabeculate)
Pentastemona
because of the size of its flowers.
egregia
nerves
Steenis:
van
on the
habitat,
same
J.
the
(not
white crystal
thick, large, solid style,
and non-ribbed seeds without arilloid appendage
placentas,
at
apex of the funicle.
ACKNOWLEDGEMENTS
In the
effort to solve
de
Wilde
for
granting
Further I extend
the
examining
flowers
and
assistance
detail
fruit
Dr.
W.
the
Anei
identity
the
me
I
of
which
species
sincerely
normal binocular
from these
forest and
has
jungle
in
data
on
privileged
W.
the
extremely
the first
are
place
to thank
due to Dr.
in
discovery
W.
J.
van
delicate
acknowledge
the microtome
Heel,
the
examination
I could
which
in
material
visited
1981,
with
sympathetic
of the
hardly
flowers. Warm thanks
hunting Rafflesiaceae
O.
North Sumatra.
and to Dr. J. Muller for
anatomy
I
Mr.
J.
(Singapore) for providing me
with
A.
collecting complete
precise
new
descriptions
Dr.
by
in
Thanks
F. Maxwell
Latin
rambles
succeeded
defeated
long
so
his
J.
I feel
sumatrana
ovary
Meijer who, during
canyon
Most
providing
the
I have
suggestions.
his remarkable
to Dr.
of P.
of the
a
P. Baas in
For
and
study
to
indebted
Stichoneuron.
of Dr. H. Sleumer.
under
Cryplocoryne egregia
privilege
most
am
of
encouragement
thanks to Dr.
pollen.
structure
obtained
my
the
for his
Bogner (Munchcn)
J.
at
have
go to Prof.
my
of Pentastemona
request
egregia,
a
identification.
REFERENCES
AYENSU, E. S.
9
I. H.
BURKILL,
J.
R.
R.
relation
K.
15a:
NAKAI,
T.
56:
1917. A
D.
HUTCHINSON,
KRAUSE,
Bot.
to Paris
J.
HOOKER,
t.
Comparative vegetative anatomy
1960. The
Soc.
Linn.
GATES,
2,
1968.
of the Stemonaceae.
Bot. Gaz.
129:
160-165,
fig.
systematic study
and Medeola.
1888.
J.
The
evolution of
Dioscoreaceae,
of the North American
Bot.
membranaceum.
families
Stemonaceae.
224-227, fig.
the
Ann. Missouri
Stichoneuron
1959.
1930.
organography and
the
of the
family
yams.
319-412.
of
Gard.
In:
flowering plants.
In: A. ENGLER & K.
genus
4:
HOOKER,
2.
Trillium,
its
variability
Icon.
PI.
18: t.
1776.
Monocotyledons: 657, fig.
PRANTL,
and its
43-87.
Die
natiirlichen
E.
Oxford.
Pflanzenfamilien, ed
80.
1937. Croomia
kiusiuna
Makino.
Iconographia plantarum Asiae
Orientalis
2(3): 159-160,
60.
OHWI,
J.
SMALL,
J.
1965.
K.
Flora
SWAMY, B. G. L.
Lour.
of
Japan:
1933. Manual
P.
B.,
(Stemonaceae).
J.
&
E.
14:
S.
Washington,
Southeastern
1964. Observations
Phytomorphology
TOMLINSON,
279.
of the
on
the floral
458-468,
AYENSU.
Am. Arbor.
49:
44
D.C.
Flora:
309, fig.
morphology and embryology
of Stemona tuberosa
fig.
1968.
260-275,
Morphology
48
fig.
and
anatomy
of
Croomia
pauciflora