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Transcript
BLUMEA
45
(2000)
Revision
377- 396
the genus Cyathostemma (Annonaceae)
of
T.M.A. Utteridge
Department
of
University
of
Botany,
School
of Plant
Plant Science
Sciences,
RG6
Reading, Whiteknights, Reading, Berks.,
6AS,
Laboratories,
United
Kingdom
1
Summary
The
Cyathostemma Griff., occurring
genus
of ten
species
is
recognised.
is discussed
Tetrapetalum Miq.
to Uvaria
L. A
One
key
to taxa
is
new
and
from SW China to NE
reduced,
with
given,
C.
species,
in
new
siamensis,
part,
to
synonymy
and
descriptions
Australia,
is
with
status
of the
Cyathostemma,
distribution
and
revised,
is described. The
a
total
genus
and in
part
maps.
Key words: Annonaceae, Cyathostemma.
Introduction
Griff, is
Cyathostemma
from China
by
was
KeBler
genus of 10 species
a
(1993),
but this is
at
the
East Asian genera of the Annonaceae is
This is the first time that
a
Cyathostemma belongs
acterised
merous
by
valvate
The
ing
genus
at
on
to
sepals,
often latrorse
attached ovules
revision of
a
stamens
with
is characterised
currently
a
tribe of the
petals,
stellate
ligulate apices,
torus
by globose buds,
with
family
South
on
and it is estimated that
under
study (KeBler, 1990).
subfamily
or
many
a
work
has been undertaken.
Cyathostemma
conical
shallowly
are
of the
treatment
Monographic
currently neglected,
the Uvarieae,
imbricate
level.
generic
13% of the 35 South East Asian genera
only
of lianas in the Annonaceae, distributed
Malesia and into North Australia. The last
through
Annonoideaechar-
caespitose indumentum,
carpels
narrow
and
nu-
laterally
flat apex (Van Heusden, 1992).
with small
petals
not
expanding
or
reflex-
anthesis.
MORPHOLOGY
Members of
are
compound
branches
sensu
of
Cyathostemma
hairs
directly
are
of four
composed
from the
epidermis.
Cyathostemma
possess
caespitose
1)
consequently
Present
not
address:
Kingdom.
diagnostic
The
Royal
or
as
usually
long.
The hairs of
hairs, up
are
to
hairs
eight,
here termed
Cyathostemma
in
a
tuft which
caespitose (=
tufted
observed in Uvaria L. All members
rather than stellate hairs,
caespitose
of
m
more
These hairs
Steam, 1983), rather than stellate,
members of Uvaria also possess
is
climbers, 3-40
(KeBler,
pers.
although
comm.);
some
African
this character
Cyathostemma.
Botanic
Gardens, Kew, Richmond, Surrey,
TW9
3AB,
United
378
BLUMEA
The inflorescence of
the
on
it
is
can
for
upwards
grow
an
axillary, position
terminal
become
then
a
an
will
If this
extra-axillary
The
one.
small
and
globose flowers,
maturity.
the
will
growth
Four
petals
are
does
dominantly
has
uniform.
argenteum (Blume) J. Sinclair, which have
or near
of C.
i
micranthum is
for
occur
in C.
position
will
distinguished
expand
not
although
species,
not
seen.
is
Cyathostemma
which do
it is
petals.
or
not
reflex
at
known if
The genus pre-
C. micranthum and C.
and red flowers. Data
yellow-green
con-
longipes Craib, but
clearly
was not
two
then
growth
in C. micranthum does
whorl of four
or one
yellow-green flowers, except
same an
axillary,
terminal inflorescence
,
petals
the
exactly
here that
sometimes found in C. excelsum
two
not
position
hypothesised
initially
quite
if
not occur an
in C. excelsum (Hook.f. & Thomson)
seen
perianth segments
this is because of two whorls of
occurs
the internodeof the
same as
occur;
growth pattern
within the genus is
structure
is
internode of the
corresponding
extra-axillary position
in the herbarium material studied this
by
the
exactly
position
and the
manner
concaulescence. This
to
about by concaulescence, and this may also
come
Floral
is
side branches, and it is
sympodial
be due
Griff. The inflorescence
viridiflorum
on
position
J. Sinclair where it is terminal. The
same as
growth
is sometimes
as
The inflorescence
shoot.
DC.)
to
When this
occur.
is found,
J. Sinclair and C.
tinues in
distance the
inflorescence
opposite
extra-axillary position
here
1989): the
single prophyll
a
shoot is situated more towards the main axis, and
axillary
(Weberling, 1989).
main axis,
initially
a
2000
2,
rhipidium (sensu Weberling,
axillary
hypothesised
of the
primordium
main axis
a
for C. micranthum (A.
extra-axillary position
No.
45,
monochasial fashion from the axil of
a
adaxial side of the
extra-axillary, except
when the
is
Cyathostemma
inflorescence branches in
positioned
Vol.
gained
from herbarium labels of C. micranthum indicates that flower colour ranges from red
through
brown and orange
biology, however,
have
not
been made
C.
a
black
sap,
at
has
Cyathostemma
udes
and all
to
green. Petal colour may be
species
the
numerous
termed
may,
correct
a
time of year,
carpels
Floral characters,
in the past been used
in the
separation
of
stigma
notably
or
a
and (C.
day,
. In those
genteum.
taxa
not
longipes
characters for
species
known
or
only partially
where ripe fruits
are
nonaceae,
was
first named
Griffith
and
was
thought
by
Griffith
unsure
that
ring
ofhairs
at
ex-
King,
the junc-
of the
petals
and ovule number, have
distinctions. This has resulted
the
next
are
chapter).
and C.
longipes
glabrous, except
densely pubescent
(1854),
who described the
relationship
Cyathostemma
(see
known for C.
ar-
for
fruits.
HISTORY
was
of this
species,
single species
even
C. vi-
within the An-
intermediatebetween the Annonaceae
and the Schisandraceae, and that he knew of "no
that of this one."
a
known, the fruits
TAXONOMIC
ridiflorum.
have
which
hookeri
from Cyathostemma and the artificial distinction
C. argenteum and C. excelsum which both have
The genus
this character manifest.
to see
1982). Cyathostemma
flower colour, number of
Tetrapetalum Miq.
are
reproductive
glabrous U-shaped stigma
between C. vietnamense Ban and C. yunnanense Hu
Fruit characters
on
and the ovary.
diagnostic
as
with
dependent
time, have red petals, but collections
compitum (Endress,
glabrum (Span.) Jessup ex Utteridge
tion of the base of the
at some
anonaceous
plant
with
a
corolla like
T. M. A.
King (1893)
was
he described four
the
next
C. scortechinii King and
a
hookeri was described by
Thomson,
of
an
describe
scortechinii
Cyathostemma
and later to
a
been traced and
as
a
in the Annonaceae,
rare
such
plant
this would
as
that Uvaria
reduced
Craib
described C.
(1925)
distinct and
In the
natum.
viridi-
Sinclair(1955).
species
memberof
was
more common
Cyathostemma
described.
Gynodioecy
(Van Heusden, 1992). A
studied
intensively
an
from
longipes
a
collection by Kerr made in Thai-
single
at P
of
Sinclair
by
(1955)
Cyathostemma
common
species
description
of C.
yunnanense
to
to a
synonym
A continued
Cyathostemma.
known
now
These
by
to
species
two
seem to
2 &
(Map
C. yunnanense,
by Jessup, published
of the
here
volubile Miq.
as
following
in ovule number, and
limits of
thought
to
described
was
transferred to C.
being
gla-
combination.
be distinctfrom
by Miquel
in
Cyathostemma solely
1865.
Merrill (1929)
published by
Momose
(1998).
because
subsequently
recently
Within the Annona-
reports of dimerous flowers regularly occurring in species that normally
are
have trimerous flowers
two
here
is
has resulted in
Cyathostemma
described T. borneense Merr. from Elmer's collections of Bornean plants, and
of the
most
5).
extend into southern China,
differing only
generic
a new
T. lambirense K. Momose has been
there
make
be the
Hu (1940). Ban (1974) described C. vietnamense,
from northern Australia and Timor,
of its dimerous perianth,
ceae
acumi-
of C.
yunnanense.
understanding
glabra Span.,
Tetrapetalum
to
in the genus with the widest distributions
which is very similar
reduced
brum
Malaysia,
the dubious taxon C.
of the material in herbaria, and therefore
The distribution of the genus is
Uvaria
of the Annonaceae from Peninsular
recognised, including
were
excelsa Hook. f. & Thomson
large proportion
a
confirms that this
In this work Sinclair transferred Uvaria micrantha Hook.f. & Thomson and
Mitrephora
the
member
legitimate species.
treatment
species
seven
up
a
of C.
variety
dubious taxon, because the
has
type
a
land. Materialadditional to the type collection in the herbarium
a
Hook.f. &
was
like Peninsular Malaysia.
area
is
King. Cyathostemma
parviflora
to a
have been re-collected in
probably
wrayi King,
C.
viridiflorum by
gynodioecious
androdioecy being
when
Cyathostemma,
parviflora A. Rich.,
was
have been made in the one hundred years since this
is
of
hookeri,
C. acuminatum
synonym of C.
further collections of
no
C.
namely:
when he recognised
379
Cyathostemma
species
new
the genus,
acuminatum is here treated
Cyathostemma
genus
later homonym of Uvaria
florum by Ridley (1922),
not
to
in
of the
gynodioecious species
King
illegitimate
Cyathostemma.
author
species
new
Revision
Utteridge:
genera
(Sinclair, 1955; Koek-Noorman et al.,
based
this
on
single
character has
1990),
recently
and the
been
separation
doubted (Van
Heusden, 1992; KeBler & Van Heusden, 1993). Apart from the difference in merosity,
there
are no
stemma,
other characters which support the
and
T. volubile has
KeBler & Van Heusden
now
remain
of these
better
to
petalum
be assessed.
species
placed
(1993).
as
The
outside of the
The
separation
been reduced
recently
relationships
petals
which
generic
to a
of T. borneense and T.
enlarge
limits of
and reflex
to
at
Cyathostemma,
members of Uvaria. Indeed, Momose
would be reduced
of T. volubile from
(1998)
Uvaria if the number of
Cyatho-
synonym of C. excelsum
tepals
anthesis
and
by
lambirense
place
they
are
both
more
commented that Tetrais
not
used
as
generic
character, but overlooked the reduction of T. volubile by KeBler & Van Heusden (1993).
Consequently,
the
new
here (see 'Excluded
combinations U.
taxa').
borneense and U.
lambirense
are
proposed
380
BLUMEA
Vol.
RELATIONSHIPS OF
The
of
position
No.
45,
CYATHOSTEMMA
within the tribeUvarieae has remained unchanged since
Cyathostemma
deae except for Walker's (1971) classification (see
is formed
the groups
by
Below the tribal level, Fries
of the inflorescence
account
in
sometimes have
axillary
an
group, Fries (1959)
position being axillary.
of
as
inflorescence
cauliflory
position.
the
genera
within the
Cyathostemma
placed
in the Malmea
Anomianthus Zoll.,
A
more
both
recent
natural
one
subfamily,
attempt
and
at
and
differs from
are
bilobed
into
(1998)
that
states
stigmas compared
petal
the flat,
sensu
Fries
be unnatural,
to
structure
of all genera of the
formed
natural group.
a
by
(1992)
also
be
or
that
not
Tetrapetalum
formally proposed.
Tetrapetalum
slightly
a more
including
it should be noted that
classification was
differs from
convex
seems to
notes
number. However,
a new
(1992) places
Van Heusden
that several of Fries' groups,
notes
Cyathostemma
to
who have
dissatisfied
classification. The Uvaria tribe
Uvaria group which
a
unnatural. Van Heusden
in
The Asimina
group
and
a
Cyathostemma
Tetrapetalum
Cyathostemma only
was
each other.
to
subtribal classification
Van Heusden's groups are informal, and
Momose
and he
and within the Uvaria tribe the genera Uvaria,
than that of Fries. Van Heusden
the Asimina group,
of the Asimina
authors
previous
This group appears
the basis for
as
and C. excelsum
description
African genus, one Australian genus,
pollen morphology
Tetrapetalum
Cyathostemma
position
taken into account, and it is doubtful that
are
within this group have any affinities
Annonaceae, and used these data
on
of the inflorescence
viridiflorum
In his
group.
one
Cyathostemma.
biogeographic relationships
Walker (1971) examined the
was
in the Asimina group
possessing axillary inflorescences,
and four Asian genera, including
if
C.
has confused
consists of eight genera restricted to America,
especially
The
Cyatho-
Scheff.
Rauwenhoffia
(1959) placed Cyathostemma
that
notes
quoted Cyathostemma
position
Uvaria and
comprising
Annonoi-
subfamily
The sister group of
below).
usually extra-axillary, although
is
Cyathostemma
with the
2000
moved the genus from the Unoneae, and within the
King (1893)
stemma
2,
concave
in
possessing
stigmas
found in
buds
maturing
Tetrapetalum.
forms
Cyathostemma
into small flowers, and
anthesis. However, it is
Uvaria when
stemma
Balonga
study
a
(including
a
Ellipeia
distinct flower
morphological
a
subacute apex which do
Tetrapetalum p.p.)
related
that
a
debatable whether
hoffia, closely
to
with
globose
not
Cyathostemma
to
there
are
variants"
or
reflex
at
Together
with
Cyatho-
six other genera, viz. Anomianthus,
Hook.f. & Thomson,
Uvaria which may
expand
will remain separate from
of Uvaria and related genera is undertaken.
Le Thomas,
Uvaria has still
group because of the
monophyletic
petals
Ellipeiopsis
in fact be
a
large
(Van Heusden, 1992).
be conducted, and this work is
presented
R.E. Fr. and Rauwengenus with "several
At present
here in the
a
study
of
knowledge
greater understanding of Uvaria and its related genera will possibly result in the
'loss' of several genera,
Cyathostemma
included.
DISTRIBUTION
The
genus
is
distributed from
Australia. The
Yunnan
most
province
southern
China
throughout
northern member of the genus is
of China and in Vietnam. Two
species
Malesia and into North
C. yunnanense, found in the
are
found in North Australia,
T. M. A.
C.
micranthum and C.
The
(Jessup, 1990).
Utteridge:
Revision
the
of
in
glabrum, particularly
distributed
widely
most
genus
and
mesophyll
is C.
species
vine forest
notophyll
micranthum, which
Malesia and extends north into Thailand and Vietnam
throughout
381
Cyathostemma
occurs
(Map 5).
TAXONOMIC TREATMENT
CYATHOSTEMMA
Cyathostemma Griff.,
Bengal
61
Not.
pi.
Asiat. 4
(1893) 8; Ann. Roy.
Bull.
(1922) 27; Hutch.,
Misc.
(1854) 707;
Gen. Indo-Chine
Suppl. (1938) 70;
Pflanzenfam.
2,
Flow.
PI.,
Blumea
Dicot.
17a II
(1992)
KeBler &
955; Engl.
257;
117.
&
Heusden,
—
et
al.
Dicot.
(eds.),
Fl.
1
3,
chasial cymes,
usually
green
abaxially.
extra-axillary
Petals
(4-)6,
in size, the innerwhorl
expanding
or
basally glabrous
with blunt
or
style
not
A
known.
Descriptions
branches reddish-
or
(1993) 111,
or
globose
Penins.
(1925) 29; Ast,
R.E.
(1979) 28,
f.
Gen.
11; Heusden,
Fam. Gen. Vase.
(eds.),
Fl. Australia
2
&
7
Suppl.
p.p.;
K.
Gen.
Hook.f.,
Bull.
(1992) 142,
1
(1867)
Kew
(1923)
pi.
Misc. Inform.
Blumea
Momose,
KeBler
p.p.;
43
grey;
at
on
elliptic,
in
obovate
pendulous
or
usually oblong,
clustered
or
terminal,
never
whorls of two, whorls
two
base and imbricate
mature
given
are
at
mono-
axillary,
flowers with
sometimes
over
and
densely
a
so
with
a
broadly
numerous,
anthers
taxon
some
species
if known.
CYATHOSTEMMA
sub-coriaceous, drying green
or
pale green-yellow;
inflorescencesubtended
by
a
mature
persistent
ovate-
2
drying green, brown, yellow,
brown, dark brown
ovate-ovate, coriaceous bract,
2a. Leaves sub-coriaceous;
b. Leaves chartaceous;
mature
petals
petals
red
never
5
basal when 2.
rows,
for each
c.
abaxially,
hippocrepiform stig-
above because the fruits of
given
small
concealing anthers,
pubescent
two
sub-equal
tips, coriaceous,
the adaxial surface. Stamens
arching
purplish-brown;
branches when
by
in
(1998)
Sepals (2 or) 3, valvate, densely pubescent
b. Leaves coriaceous, sometimes sub-coriaceous,
tended
2
pi.
(in press).
orbicular, chartaceous foliose bract
or
1
Fl.
Nat.
Fr„
(1964) 104; Hutch.,
3; Jessup,
from the old wood
lateral in
of fruits
J. Asiat. Soc.
Malay
Miq.
numerous,
not
KEY TO
la. Leaves chartaceous
Blumea
valvate
2—16(—18),
of fruits is
al.
Benth.
(1865) 1;
anthesis, resulting in
Carpels
Fl.
(1955) 219;
1
Sin. 30
et
f.
slightly clawed, pubescent
extrorse.
description
in Kubitzki
whorls of three or
connective apex
absent. Ovules
—
at
650; King,
Griff.
scarcely pubescent
very
ligulate
or
two
reflexing
latrorse, introrse or
Note
in
14
Fl. Java
Reipubl. Pop.
Flowers
opposite
t.
Fl. Siam.
Sing.
Bull.
hairs. Leaves alternate,
slightly smaller,
inner whorls
opening,
2
pi.
sometimes red.
yellow,
or
(1854)
(1893) 11; Ridl.,
3,2 (1888) 31; Hutch.,
volubile
emarginate.
or
1
Bakh.f.,
(1964) 86; Heusden,
caespitose
Asiat. 4
(1993) 60,
2
Lugd.-Bat.
Type species: Tetrapetalum
Climbers with rufous
are
Li,
KeBler
Rheedea
Fam. Gen. Vase.
base cuneate, rounded or
mas,
&P.T.
Nat. Pflanzenfam.
PL,
Gard.
Backer &
37c, f;
f.
Ann. Mus. Bot.
Prantl,
Gen. Flow.
Kubitzki
mm
144,
pi.
(1923) 256; Craib,
Sinclair,
Type species: Cyathostemma viridiflorum
Tetrapetalum Miq.,
not
J.
(1959) 73;
(1964) 82;Tsiang
7
Suppl.
(1993) 114;
—
ed.
Kew
Inform.,
Ic.
(Calcutta) 4,
Bot. Gard.
to
red
green-yellow
or
or
black
black; inflorescence sub-
if foliose then
vestigial
3
1. C. argenteum
10. C. yunnanense
BLUMEA
382
3a. Leaves
No.
45,
2.5-8 (-12)
oblong-elliptic, small,
adaxial midrib densely rufous
inflorescence terminal,
Vol.
long, drying rusty-brown
cm
very short,
pubescent; petiole
opposite
2000
2,
flowers red, brown,
extra-axillary;
or
less than 3
b. Leaves
oblong-lanceolate
or
axially; petiole greater than 3
green,
so
b.
sometimes black and then
mm
and
long;
yellow
never
ad-
terminal; flowers yellow
4
the leaf surface;
obscuring
leaves
sometimes
abaxially pubescent,
densely
flowers sometimes dimerous;
peduncle absent;
2. C. excelsum
tomentose
grey, reddish brown
cent or
long, drying
cm
completely glabrous
or
dark brown-black; leaves
flowers
glabrous; peduncle present;
abaxially sparsely pubes-
always trimerous; ripe
monocarps
gla-
brous
5
5a. Leaves
than 5
drying light blue-green, glabrous; petiole
cm
b. Leaves
inflorescence of up
long;
dark green
drying
less
long; peduncle
or
than 3
to
mm
long; peduncle greater
25 flowers
brown, sparsely
8. C. viridiflorum
pubescent;
1-6.5(-8)
petiole
inflorescence of 1-10 flowers,
long;
cm
6-12
always
mm
less
than 20 flowers
6a.
Petiole 6.5-8
ripe
7a.
6
long; petals abaxially furrowed, less than 3
mm
monocarps greater than 30
b. Petiole
of
ripe
1—5(—7)
less
monocarps
Leaf apex
acute or
greater than 18
than 20
Adaxial midrib
by
2.5-6
never
wide; stipe of
4. C. hookeri
furrowed, greater than 4.5
mm
wide; stipe
long
mm
7
somewhat acuminate; inflorescence of 1-3 flowers;
mm
long; carpels pubescent
less
flowers, pedicels
mm
long
mm
long; petals
mm
b. Leaf apex acuminate, with the
8a.
acumen
than 12 mm
glabrous; pedicel
to
up
the base of the
at
3
cm
long;
8
stigma
inflorescence of 3
1.8-2.2
long,
cm
leaves
9a.
Twigs
pubescent; pedicel
flowers
cm; mature
striate with
9
5
mm
long,
up
Twigs slightly
1.
3-9.5
mm
to
2
cm
long; carpels
c.
cm
2
21
(1955) 220;
Java,
Backer &
Bakh.f.,
Bakh.f.,
Fl. Java
Ann. Mus. Bot.
Sarawak Mus. J.
?Bantam.
cm;
c.
20
longipes
inflorescence
and somewhat tuberculate, greater than
mm
9. C.
long
leaves 6-14.5
but smooth, less
than 5
by
wrayi
3-4.5 cm; in-
mm;
carpels
4-5
7. C. siamensis
(1830) 24; Miq.,
nitidum
5. C.
brown
leaves 16-29 by 6-10
J.
Sinclair
Cyathostemma argenteum (Blume) J. Sinclair, Sarawak
Ridl.,
light
glabrum
oblong-obovate,
long
Cyathostemma argenteum (Blume)
14
bark
conspicuous lenticels;
peduncles woody
leaves
long,
diam.;
becoming woody
striate with
florescence branched;
mm
15
6-17
diam.; bark dark brown-black
mm
inconspicuous lenticels;
unbranched; peduncles
b.
c.
or more
elliptic-oblong,
3. C.
6-6.5
pedicels
long; carpels pubescent throughout
flowers 8-10
cm; mature
b. Adaxial midrib
by
or
red
rarely
monocarps
Twigs
inflorescence
long;
Twigs black, conspicuously striate;
ripe
large, (6-) 12-27
obovate-oblong,
or
blue-green, yellow,
green,
or
black,
6. C. micranthum
green
4a.
or
mm
1
(1964)
Lugd.-Bat.
1,3 (1913)73; Merr., Bibl.
Blumea
12
(1963) 61,
nom.
illeg.
2
—
Map
Mus. J.
5,
104.
Uvaria
—
3
1
(1951) 599;
(1865) 8; Boerl.,
Enum. Born. pi.
—
Type:
Gard.
argentea
Icon.
(1921)
Bogor.
253.
VanHasselts.n.
Bull.
Blume,
—
1
Sing.
Fl. Javae
(1899) 95;
Cyathostemma
(holoL), Indonesia,
T. M. A. Utteridge:
Uvaria
bracteata
Fl. Ind.
Roxb.,
ed.
Revision
1832
of
(1832)
the
660.
—
383
Cyathostemma
genus
Roxb.
Type:
Icon
2290
(lecto K,
here
Type: Gomez s.n. {Wall.
Cat.
designated).
Uvaria
Climber to 5
drying
iso
auct.
non
forming
a
1
cm
ceous
a
bracts
broadly
axially.
mm
along
long,
the
c.
on
cuneate
both sides,
—
No
glabrous,
2
by
3
mm,
ovate, red
connective apices
edges,
c.
2
mm
and
the
mm;
3
c.
pubescent
5
on
mm
slit
and inter-
both surfaces
densely pubescent,
'midrib', membranabracteoles similar
mm
long. Sepals 3,
long,
to
cori-
both ad- and ab-
apex acute-obtuse,
the adaxial surface. Stamens
anthers latrorse.
long, stigma deeply
on
1-4-flowered, opposite
densely pubescent
green-yellow,
ligulate;
c.
apex acute;
ascending
pedicels
along
along the pedicel,
apex obtuse,
or
pubescent midrib,
rounded base,
or
long. Inflorescences
so
reddish brown
(4—) 10— 15(—21) by
reticulation visible
densely
type designated.
sub-coriaceous; minutely
to
curving
Peduncles absent;
the abaxial surface,
on
Carpels 4-angled, pubes-
the adaxial side, ovules
16.
minutely pubescent, oblong, slightly apiculate, slightly constricted,
somewhat tuberculate,
Map
10.
(1941)
oblong-lanceolate,
to a
margin,
mm
third of the
way
broadly
monocarps
rows,
the
extra-axillary.
ovate,
sparsely pubescent
2
on
abaxially pubescent,
c. one
Petals 6,
Ripe
—
1827.
sub-chartaceous, foliose, orbicular, 7-9 by 7-9
aceous,
cent
near
close-network; petiole 3-5
bracts, attached
2-2.5
narrowed
slightly prominent
faint broken line
long;
to
203.
when young, later
lamina
green-yellow;
the leaves, sometimes
c.
Bekn. Fl. Java 3
Backer,
gradually
cm,
12-14 pairs,
in
(1832)
September
lenticels indistinct. Leaves chartaceous
pale
or
green
arching
Blume:
8
when very young, later glabrous except for the adaxial
(1.5—)4.5—6(—7)
nerves
Geneve 5
Phys.
K-W) Burma, Tavoy,
Twigs densely pubescent
m.
finely striate;
puberulous
Mem. Soc.
DC.,
G n.v.;
Uvaria argentea
and
A.
gomeziana
6459) (holo
c.
4
by
2-2.5
cm,
stipes
stout,
c.
4
cm
long.
Seeds several in
smooth.
1. Distribution
of
Cyathostemma argenteum (Blume)
J. Sinclair
(•),
C.
yunnanense
Hu
(�).
384
BLUMEA
Distribution
most
—
in Peninsular
Cyathostemma
—
No.
45,
coriaceous leaves,
drying
or
with the thicker leaves,
is
petals
distinguishes
be
can
green-yellow
Cyathostemma
KeBler
Fl.
&
Ind.
excelsum
1
(1855)
Porters.n.
(Wall.
Cat.
excelsa
Wall., Cat.,
Born.
pi. (1921) 253; Ridl.,
6477), [holo K-W;
Tetrapetalum
volubile
1
nom.
(Calcutta) 4,
Type:
Uvaria
De
f.
Fl.
Ann. Mus. Bot.
Sarawak
s.n.
Mus. J.
m
brown
brown due
to
cm,
Univ. Calif.
8-11
to
pubescent
cent on
narrowed
on
both
to
the
15
Fl.
Thomson,
pedicels
0-5
mm
degree
long;
the bracts, attached about
obscured due
to
both surfaces but less
the
Thomson,
Brit.
Ind.
263.
pi. (1921)
pubescent
1
—
long, pubescent;
sub-globose
(1929)
to
Type:
Porter
—
Nuovo
the
Bot.
Enum.
(Wall.
s.n.
Cat.
1832.
Giorn.
Enum. Bom.
Bot. Ital.
pi. (1921)
254.
61.
—
Type:
Elmer 21081
(lecto L,
Province.
later
glabrous,
very dark brown
with
pubescence varying
to
adaxially gla-
sometimes
abaxially pubescent,
of
Roy.
74; Merr.. Bibl.
Kalimantan.
or
age,
densely
drying
(9—) 13—22(—37) by (4 —)6
—
cordatebase, apex acute-acuminate;
reticulation
crookedly,
on
mm
the older
the
3-5.5
mm,
or
sometimes red, 5-6
rows.
Pe-
ovate; bracteoles
both bracts and brac-
Sepals
2
or
3,
connate at
6, broadly ovate, the inner
by
5-6
mm,
sepals, basally glabrous abaxially.
anthers latrorse
close
apex sub-acute, densely pubes-
the adaxial. Petals 4
12-16 lateral in 2
a
Inflores-
twigs, extra-axillary.
pedicel;
clustered flowers.
by
forming
long, pubescent.
pubescent, coriaceous,
halfway along
apices ligulate;
ovules
bracts
2-4
mucronate,
connective
Ann.
(1893) 22;
(1913)
Bibl.
oblong-obovate,
densely
so on
as
61
3
(1865) 1; Becc.,
Merr..
Tawau
and ascending rather
rarely
cent,
Bot.
emarginate
slightly clawed, green-yellow
mm
(1955) 226;
(herb. Benth.), K-W], Malaysia,
1,
34.
surfaces; petiole 7-10(-12)
coriaceous, semi-orbicular,
long;
iso K
2
darker below sometimes
green above,
orange-
pale
whorl
mm
2
(1913) 73;
sunken midrib,
sometimes
or
pairs, curving
usually
the base,
&
Enum. Born.
Bengal
(1922)
clustered cymes of 4-20 flowers, sessile
similar
Sing. 14,
lenticels indistinct. Leaves coriaceous;
the dense hairs; lamina
duncles absent;
teoles
1
Lugd.-Bat.
3
Publ.
the leaf surface, the
slightly
network, visible
cences
Sealy
2
Map
—
(herb. Benth.)], Malaysia. Penang,
long. Young twigs pubescent,
brous except for the
obscuring
1,
K
K, NY, P), Malaysia, Sabah,
iso
conspicuously striate;
nerves
Bibl.
Sarawak Mus. J.
(holo L), Indonesia, Borneo,
10-30
12(—16)
is unnecessary.
excelsa Hook.f. &
Hook.f.
(1858) 31;
Penins.
Miq.,
black and
very
Gard. Bull.
Mitrephora
—
J. Asiat. Soc.
18; Ridl.,
Malay
Climber,
pale
Sinclair,
(herb. Hook.f.);
K
nud.; King,
(1893) 26,
confertiflora Merr.,
and
1
J.
f. 3.
(1913) 86; Merr.,
(herb. Hook.f.),
Vriese
designated here;
so
3
iso K
(1871) 179; Ridl.,
—
Bat.
6477) [holo
Gard.
3
Thomson)
1,
different from
not
was
& Thomson) J. Sinclair
(1993) 60,
Fl. Ind.
114; Miq..
(1941)]
new name
1832.
Penang,
Uvaria
1
3,
Sarawak Mus. J.
(1872) 77; Ridl.,
Type:
&
(Hook.f.
Rheedea
Heusden,
together
Flora Indica icons.
Roxburgh's
excelsum (Hook.f.
Cyathostemma
red which,
C. argenteum from the similar C. yunnanense.
and so Bakhuizen's intended
discusses and lists
(1956)
2.
original concept,
or
sub-
to
subtending
and the foliose bract
Backer's usage of U. argentea [Bekn. Fl. Java 3: 10
Blume's
the chartaceous
distinguished by
pale green-yellow,
the 1-3-flowered inflorescence. The
and Borneo. Al-
(type locality)
Malaysia.
argenteum
green
2000
2,
Java
Bangladesh, Burma, Thailand,
certainly present
Notes
Vol.
slightly
to
Ripe
globose, tuberculate, yellow-orange
apex
introrse.
monocarps
sometimes
obtuse,
Stamens 2-2.5
Carpels
3
densely pubes-
brown,
2-2.6
by
T. M. A.
Map
2. Distribution
2-2.5
cm, on a
3
long.
c.
cm
—
(P), but with
Note
This
—
of
Revision
Cyathostemma
of
12, in 2
c.
the
(Hook.f.
2-2.5
385
Cyathostemma
genus
excelsum
pubescent receptacle
Seeds several,
Distribution
s.n.
swollen
Utteridge:
&
Thomson)
J. Sinclair.
diam.; stipes densely pubescent,
cm
rows.
Sumatra, Peninsular Malaysia, Borneo. A single collection by Pierre
no
locality data,
is
species
extends the distribution north into Indochina.
easily recognised by
the
abaxial leaf
densely pubescent
surface, with clusters of tomentose flowers with short pedicels.
3.
Cyathostemma glabrum (Span.) Jessup
Uvaria
glabra Span.,
Climber.
black
Linnaea
and buds
Young twigs
tinged
with red
elliptic, ovate-elliptic
acute or
shortly
sometimes
and
comb.
nov.
Map
—
3
—Type: Spanoghe s.n. (holo L), Indonesia, Timor.
when young,
soon
brown-
becoming glabrous,
distinctly striate; lenticels indistinct.
glabrous above, glabrescent below, drying rusty brown;
or
lamina
rarely obovate, (3.5-)6-17 by 2.5-6 cm, base rounded, apex
bluntly acuminate;
bifurcating
Utteridge,
densely pubescent,
especially
Leaves sub-coriaceous,
162.
(1841)
15
ex
before
nerves
8-12
pairs, interarching
joining together; petiole 2—4(—5)
mm
near
long,
the
margin,
channelled
above, sparsely pubescent. Inflorescence opposite the leaves, usually single-flowered.
Peduncle absent;
pedicels
c.
the
halfway along
acute or
ovate,
anthers latrorse.
dark
the
purple,
Carpels
10-16
from the
—
5-7
1.5
c.
junction
Distribution
seen
pedicel. Sepals
margins incurved,
at
mm
long;
mm
long;
bracts
3, broadly
stigma
10-15
Found
4.5-7 mm,
ovate, 3.5-4
on
Philippines.
mm;
puberulous.
connective
long;
numerous,
of the
by
by
mm
1.5-2
apices
by
3-3.5
mm
12-20
apex
depressed
Stamens somewhat
long, glabrous except
mm
mm,
or
dorsally-
somewhat blunt and rounded;
and ovary; ovules 5.
stipes
pubescent,
bracteoles similar to the bracts, attached
obtuse, pubescent abaxially, glabrous adaxially. Petals 6, broadly
ventrally flattened,
hairs
18-22
sparsely pubescent,
sub-coriaceous, ovate-obovate, 2-3
Ripe
long.
for
monocarps
a
ring
of
glabrous,
Seeds 1-3.
Timor and in northern Australia;
one
collection has been
386
BLUMEA
Note
—
Cyathostemma glabrum
the leaves, the
a
2-3
non-clasping
mm
which it differs in the
opposite
4.
Cyathostemma
1
Type:
f.
Curtis
Uvaria
parviflora
Uvaria
glabra
King,
Fl.
1213
oblong
never
to
non
Icon.
mm
Penins.
(lecto K,
1
by
C. micranthum from
to
long pedicels
here
(1899) 125,
and the inflorescence
Fl. Ind.
1
Bot. Jahrb.
lamina drying
light
Sinclair,
(1855) 103,
of 3
or
c.
the
Distribution
2.5
mm
49
Syst.
(1912)
cm,
apex
of
acute or
base
nerves
near
(Calcutta)
Sing.
14
(1955)
pi.
223.
Govt. Hill
124.
glabrous,
reddish brown
thinly coriaceous, sparsely
the
1
ovate, 2.5-3
or
very
distinct
pairs,
slightly rounded,
on
both surfaces,
margin; petiole 6.5—8(—10)
the leaves,
5-10
c.
cuneate
12-17
opposite
bracteoles
pedicel. Sepals 3,
Gard.
Enum. Born.
illeg.
later
Leaves
numerous.
4 flowers
long;
Gard. Bull.
nom.
Bot.
Bibl.
brown with the venation reddish brown; lamina
long; pedicels sparsely pubescent,
falling,
J.
42; Merr.,
t.
designated), Malaysia, Penang,
Thomson,
Span.: Diels,
1
long. Young twigs sparsely pubescent,
m
adaxially glabrous,
3.
adaxial midrib of
3
(1922) 28;
anastomosing indistinctly
along
King (�).
glabrous
Bengal 61 (1893) 10; Ann. Roy.
Bogor.
obovate, (7.5-)12-20 by 5.5-8
and
soon
the way
Map
J. Asiat. Soc.
Malay
glabrous. Inflorescence
and
the
is similar
species
the very
Map
—
emarginate, apex acute-acuminate;
curving
2000
and the inflorescence which is subtended
This
finely striate; lenticels small and
pubescent below,
0-2
King
40; Boerl.,
Hook.f. &
auct.
Climber, 8-25
and
bract.
2,
distinguished by
glabrous midribs,
hookeri
hookeri
(1893) 12,
(1921) 263; Ridl.,
—
long
is
long,
mm
No.
45,
the leaves.
Cyathostemma
4,
18-22
pedicel
Vol.
mm
mm
by
never
long;
long,
bracts foliose,
ovate, attached
2.5-4.75 mm,
obtuse. Petals 6, furrowed
Cyathostemma glabrum (Span.) Jessup
axillary.
ex
mm
or
long,
Peduncles
vestigial
c. a
third of
abaxially pubescent,
somewhat tuberculate
Utteridge (•)
and C. hookeri
T. M. A.
the abaxial surface,
on
apex
cleft and
2.5
up
around seeds, 3-4
Distribution
This
—
of 3
cymes
be
or
c.
2
cm,
a
Ripe
at
monocarps
the
by
2-3
mm,
incurved and sometimes
Carpels 4-angled,
semi-introrse.
to
4-6
yellow,
truncate,
of hairs
ring
of the
junction
1.5-
stigma
and
glabrous, slightly tuberculate,
blue-green; stipes (2—)3.5—4.8
Peninsular Malaysia,
special
gave
as
there is
cm
Seeds
long.
species
is
the
to
(holo K;
the
green; lamina
unequal,
tinctly
petals vertically
not
select any
specimens
iso
Craib
—
Misc.
Map
to
furrowed
abaxially
particular specimen
margin, prominent
mm
long,
covered with indument
4. Distribution
as
of
(1925) 8;
Fl. Siam.
sub-coriaceous, pubescent
oblong-oblanceolate,
sides; petiole 3-5
Map
traced for the
as
4
the
long,
cm
when
dry.
It
holotype,
a
Curtis
6.5-19
on
(1925)
on
29.
—Type:
the upper midrib and
sparsely pubescent, drying
by 3.5—5.5(—6.5)
nerves
cm,
channelledabove, below
interarching
sub-prominent
indis-
on
both
transversely furrowed, similarly
Inflorescence
Cyathostemma longipes
dull olive-
base rounded, often
10-14 pairs,
both sides, reticulation
the branches.
1
Udawn.
abaxial midriband lateral nerves
the
was
4
apex acuminate or caudate-acuminate;
near
collections have
not
of Curtis, hence the selection of
Inform. Kew
BM, P), Thailand, Pu,
Climber. Leaves chartaceous
nerves,
outer
Bull.
Cyathostemma longipes Craib,
lateral
Single
locality
exact
lectotype.
Cyathostemma longipes
Kerr 8607
though
the
distinguished by glabrous fruits, stipe usually
King (1893) did
mention
Sarawak and Sabah.
cultivated record from Java.
a
4 flowers with
that
seen
specimen
5.
—
Additionally
Note
but
two rows.
from Ceram and Sumatra,
seen
latter.
can
for
apices
387
Cyathostemma
genus
8.
to
been
connective
anthers latrorse
by
the
of
ovate, inner whorl clawed,
glabrous except
mm,
ovary; ovules (4-) 10-12, in
bulging
to
long;
mm
appearing bi-lobed;
0.5-1
by
elliptic
Stamens 1-2
acute.
Revision
Utteridge:
Craib
two-flowered, extra-axillary.
(�)
and C.
wrayi King (•).
BLUMEA
388
Peduncles
long;
lanceolate-ovate, 1-2
Sepals 3, broadly
ovate,
5
c.
5
by
adaxial surface
ly pubescent,
projecting
stigma
and
stamens,
China
—
Note —This
mm
this
except for
Ripe
a
the
pedicel.
along
1-2
ovate,
by
sparse towards the
Stamens somewhat dor-
Carpels
of hairs
ring
a
broadly
1
c.
mm
long,
not
of the
junction
the
at
unknown.
Indochina and Thailand.
easily distinguished by
The inflorescence
long pedicels.
very
is
a
It is two-flowered; the youngest
Cyathostemma.
condensed nature of the inflores-
highly
dichasial cyme with
terminal
non-developing
a
of the bracts is examined the inflorescence is
position
cm
sparsely
more
glabrous.
monocarps
(Hainan province),
the appearance of
flower. When the
tips becoming
anthers latrorse.
and because of the
develop,
not
3-9.5
bracteoles
halfway
c.
is very distinct within the genus and is
species
gives
the
elsewhere
tip,
monochasial cyme like all members of
cence
long;
Petals 6, coriaceous,
at
the few-flowered inflorescence and the
flower does
mm
obtuse-acuminate abaxial, surface sparse-
apex
long;
glabrous
ovules 2, basal.
ovary;
Distribution
the
at
sally-ventrally flattened, 0.8-1
attached
long,
mm
mm,
glabrous.
base, abaxially pubescent only
above
2000
2,
long; pedicels sparsely pubescent,
mm
adaxially densely pubescent
cm,
No.
45,
coriaceous lanceolate-ovate, 3-6
pubescent,
pubescent,
1-1.7
1-3
densely pubescent,
bracts
Vol.
obviously
a
monochasial cyme.
6.
micranthum (A.DC.) J. Sinclair
Cyathostemma
Cyathostemma
Australia
micranthum
2
micrantha
(in press).
4
(Calcutta)
1
Enum.
Wall.
Type:
Cat.
6449
iso
Fl.
sumatrana
Ned.
Ind.,
Sumatra, Lampung,
Popowia
nitida
King,
nated), India,
J.
42.
—
(1832)
Brit. Ind.
1
Ann.
(1893) 21;
4
Bot. France
42.
Jessup,
Fl.
Fl.
Uvaria
—
(1872) 51; Kurz,
Roy.
Bot. Gard.
(1906) 70;
(1915) 230; Ridl.,
BM), Burma,
(1832)
near
3
bijv.
(1861)
Malay
Fl. Gen. Indo-Chine
Fl.
Gen.
Penins.
1
Suppl. (1938)
Amhearst.
Type: Gomez
s.n.
(Wall.
Cat. 6430
) (holo
minate or acute;
the
J. Asiat. Soc.
to
m.
Bengal
61
92.
(1893)
4
—
t. 58.
—
Anaxagorea sumatrana
4383
on
Type: King
July
Young twigs pubescent,
iso
(holo L;
GH, K),
s.n.
(lecto K,
here
desig-
1891.
later
glabrous,
the young twigs, later very
nerves
8-15
2
cm,
base
faint on both surfaces;
continued
long,
petiole
of 2 flowers,
sympodial growth,
0-3
pubescent, chartaceous,
mm
shortly
pale
brown-black and
and
quite
somewhat
attached
ovate-rounded, obtuse, 2-2.5 by 2.5
c.
mm
1.5-3
initially
up
to
mm
or
rounded, apex
dividing
long,
into
two
acu-
before
channelledabove,
sub-terminal then
appearing
three inflorescences per branch.
long; pedicels pubescent, 2-7(-10)
ovate, 4-5
by
halfway along
mm,
black; lamina oblong-
or
cuneate
often
pairs, curving irregularly
woody, sparsely pubescent,
c.
(1899) 126,
Type: Teijsmann
sparsely pubescent, drying rusty-brown
margin,
bracts
cent, ovate,
1
—
Islands, Hobdaypur,
sparsely pubescent. Inflorescence
due
382.
Tegineneng.
elliptic, (2.5—)3—8(—14) by (2-)5.5-8
long;
61
Soc.
10
(1955) 225;
thinly coriaceous, adaxially glabrous except for the densely pubescent
midrib, abaxially
mm
Bot.
(Miq.) Boerl., Icon. Bogor.
South Andaman
distinct. Leaves
Peduncle
(1855) 103; Fl.
Bengal
Bull.
2
14,
Sing.
Mem. Anon.
(1923) 155; Ast,
iso
(holo K-W;
finely striate; lenticels indistinct
opposite
Sc.,
Flow. PI. 2
Mem. Anon.
Eerste
Climber, (2—)8—10(—15)
reaching
1
5
BM), Burma, Tavoy.
Cyathostemma
Miq.,
Ind.
Gagnep.,
Bull.
A.DC.,
J. Asiat. Soc.
Finet &
18;
Philipp.
Polyalthia fruticans A.DC.,
K-W;
micrantha
(1907) 54; Merr., Philipp.
(1922) 33; Merr.,
—
f.
Gard.
Sinclair,
Thomson, Fl.
(1877) 29; King,
(1893) 26,
Indo-Chine
63.
1
J.
Guatteria
Hook.f. &
(A.DC.)
Forest fl. Burma
(A.DC.)
—
Map
—
3.5-4
the
mm;
pedicel.
abaxially pubescent.
bracteoles
pubes-
Sepals 3, broadly
Petals 6, ovate, green,
T. M. A.
5. Distribution
Map
orange, red
yellow,
Utteridge:
of
truncate; anthers introrse.
remaining
Distribution
below the
—
Note
10
mm
—
This
cm,
Carpels
to
c.
1
long,
mm
fruiting
torus.
Seeds
(A.DC.)
most
species
is
widely
distributed
distinguished by
and the terminal
or
petals
connective
ovules 4-6.
mm
J. Sinclair.
outer
mm,
long,
mm
black, stipes 3-10
small monocarps less than 20
long,
micranthum
Stamens 0.8-1
389
Cyathostemma
genus
Ripe
4
3
mm,
rounded
monocarps
gla-
some-
l-2(-4).
through
Malesia into North
species.
the small leaves 4-6
long, stipes
mm
cm
long, always
less
of the monocarps less than
inflorescence of red, orange,
extra-axillary
by
apices
long, perianth segments
From Burma and the Andaman Islands
Australia. This is the
than 12
Cyathostemma
pubescent.
brous, 12-20 by 10 mm, red
times
of the
brown, inner petals 4.5-5 by 3
or
whorls
apex acute, both
or
Revision
or
yellow-
green flowers.
7.
Cyathostemma
Differ!
a
iso
Noi
Large
dark
conspicuis
K), Thailand,
River
Basin
climber.
glabrous
spec.
nervis
pedunculis gracilis
South Western:
nov.
intra
et ramis.
Kanchanaburi
marginem
—
Typus:
[Tripagodas,
arcuatis
confluentibus,
Kostermans
428
Burmese border
(holo
(Kwae
Expedition)].
and
and buds
slightly striate;
pubescent,
lamina
becoming
lenticels distinct
above and below except for the
drying pale brown;
cuneate
et
Young twigs
brown-grey
ceous,
Utteridge,
Cyathostemma wrayi foliis parvis,
lenticellis
L;
siamensis
elliptic-obovate,
on
pubescent
6-14.5
by
glabrous
the older
twigs.
when mature,
Leaves coria-
midrib and
secondary veins,
3-4.5
base rounded
cm,
or
obtuse, apex attenuate; nerves 8-10 pairs, brochidodromus, interarching 3-4
BLUMEA
390
from the
mm
with age.
pedicels densely pubescent,
tached
c.
half
6-9
whitish-green,
mm
Note
and
8.
—
5-9
mm,
15-25
long;
mm
adaxially glabrous, abaxially pubes-
long;
mm
so
bracteoles similar
to
ovate,
the bracts,
4
5-6
by
at-
mm,
This
—
having
a
Malay.
Rheedea
species
is close
to
C.
their
3,
(1893)
Penins.
BM),
1
1
black and
1
61
62.
scortechinii
12,
f.
Not.
(1893) 8;
38.
—
length. Ripe
King,
—
slightly
sepals.
Stamens
extrorse.
Carpels
monocarps
unknown.
with
more
flowers per inflorescence,
Roy.
6
(1854) 707;
Bot. Gard.
Gard. Bull.
432
J. Asiat. Soc,
Bengal
61
collector
14
Sing.
(lecto K,
Cyathostemma viridiflorum
Ic.
pi. Asiat.
1
(Calcutta) 4,
here
4
(1955) 221;
scortechinii
(Scortechini)
KeBler
f.
5857
Bot. Gard.
(King) Rial.,
(lecto
K, here
37; Ridl.,
&
designated), Malaysia,
(1893)9; Ann. Roy.
var.
(1854) t. 650; King,
(1893) 12,
Heusden,
Malacca.
(Calcutta)
Fl.
Malay
designated; iso
Malaya, Perak, Gopeng.
m
long. Young twigs slightly pubescent,
finely striate;
pubescent
Map
—
Asiat. 4
Type: Griffith
(1922) 27.—Type: King's
15-25
pi.
Ann.
(1922) 27;J. Sinclair,
(1993)
Peninsular
Climber,
to
but differs in the smaller elliptic-obovate
wrayi,
viridiflorum Griff.
Bengal
Penins.
Cyathostemma
1
ligulate;
like the
pubescent
anthers latrorse
branched inflorescence.
J. Asiat. Soc.
Fl.
apex rounded,
Type locality.
Cyathostemma viridiflorum Griff.,
the
bracts
drying rusty brown, conspicuous lenticels,
Cyathostemma
4,
when young, less
cauliflorous, when terminal growth
pedicel. Sepals 3, broadly
long, angular, pubescent along
Distribution
leaves
by
connective apices
long,
mm
4-5
the
along
or
rounded, glabrous adaxially, sparsely pubescent abaxially. Petals 6, broadly ovate,
apex
5
thirds
to two
2000
woody, densely pubescent,
long;
mm
2,
long, densely pubescent
Peduncles
7-12
chartaceous-sub-coriaceous, 2-3
cent,
c.
mm
No.
45,
of 3-5 flowers terminal
continuing sympodially.
then
3-7
margin; petiole
Inflorescence
Vol.
adaxial midrib,
drying light blue-green;
lanceolate, (8—) 12—17(—27) by 5.5-8
Map
later
glabrous,
lenticels indistinct. Leaves coriaceous,
6. Distribution
of
cm,
base
dark brown-
glabrous
lamina oblong-elliptic
emarginate
or
except
or
for
oblong-
rounded, apex acute;
Griff.
Cyathostemma viridiflorum
T. M. A.
10-13
nerves
larly
but
the
to
pairs,
distinct above and
margin, interarching
prominent; petiole
not
Revision
Utteridge:
6-12
genus
prominent
the
near
the
of
on
in
margin
the lower surface,
a
curving irregu-
broken line, reticulation visible
channelled above with
mm,
391
Cyathostemma
transversely arranged
lenticels, pubescent. Inflorescence cauliflorous, each cyme producing up
of which
2.5-8
pendulous,
ceous,
2 will be
only
3-4(-6)
in size and
long, persisting
mm
3
at
any
3-5
apex obtuse
mm,
6-9
mm
long;
yellow
sometimes
ovules
4-5
by
mm,
apices bluntly ligulate,
16, lateral in
15-20
long,
5-6
orange,
connective
acute,
two
Ripe
rows.
apex
the
connate
monocarps
at
25 flowers
coria-
pubescent,
pedicel. Sepals 3,
3-4
Carpels
c.
2
long,
mm
the seeds,
apex
long, pubescent,
mm
2.5-4
glabrous, oblong-ovoid,
to
abaxially
the base,
rudimentary claw, greenish
a
Stamens
acute.
truncate.
wide, with irregular bulges due
mm
bracts
halfway along
c.
or
both whorls with
rusty-tomentose. Petals 6, broadly ovate,
papillate;
to
densely pubescent,
after the flowers have fallen; bracteoles similar
the bracts and attached
to
by
time. Peduncles
one
long; pedicels pubescent,
cm
shape
ovate,
broadly
anthesis
at
stipes
2
c.
cm
cm
long.
Seeds 7-10.
Distribution
which
432 (K). This
9.
can
Sumatra and Borneo.
its
elongated peduncle,
bracts
be considered
have been
to
material from which he
is therefore selected
specimen
and cymes of up
the inflorescence. The
on
examined
definitely
as
the
and
by Griffith,
published iC. viridiflorum,
is
to
only
Griffith
lectotype.
Cyathostemma wrayi King
wrayi
a. var.
4
Map
—
Cyathostemma wrayi King,
(1893) 12; Ridl.,
—
Type:
m
a
1316
or
10-14
about 5
the leaves
cent, up
to
2
the
4-6
cm
mm
long.
Roy.
Bot. Gard.
Bull.
c.
3
or
yellow-green,
8-10
on
later
glabrous,
by
long;
10
acumen
2-3
cm
(1955)
4,
1
224.
dark brown and
glabrous adaxially
6-10
cm,
lamina
narrowed
sometimes acute;
long,
reticulation distinct
on
sparsely pubescent,
long;
ovate,
c.
c.
1
cm
or
to
nerves
3
by
4
mm,
acute,
inner whorl
narrower
apex sub-acute,
outer
opposite
woody, sparsely pubes-
long;
bracts
sparsely
rufous-pubescent
slightly warty,
especially
2 flowers,
bracteoles similar to bracts, attached
coriaceous and
mm,
both surfaces,
of 1
the older wood. Peduncles
rudimentary claw,
1.5 mm
2
sparsely abaxially, drying olive-green;
margin,
mm
Petals 6,
incurved base
(Calcutta)
Sing. 14,
Perak.
long, pubescent. Inflorescence
Pedicels
pedicel. Sepals 3,
c.
Ann.
Sinclair, Gard.
evenly, impressed above, prominent beneath, interarching
not
from tubercles
glabrous adaxially.
Stamens
J.
designated), Malaysia,
here
the midrib,
from the
bescent, broadly ovate,
along
(lecto K,
Bengal 61 (1893) 9;
(1922) 27;
acuminate with
mm
petiole
arising
on
but
pairs, curving
the lower;
1
broadly oblanceolate, (10-) 16-25(-29) by
emarginate base,
distinctly
Penins.
lenticels indistinct. Leaves chartaceous-coriaceous,
few hairs
oblong-obovate
the
J. Asiat. Soc.
Malay
long. Young twigs minutely pubescent,
distinctly striate;
except for
Fl.
Scortechini
Climber, 40
on
Malaysia,
distinguished by
represented by persistent
are
original
therefore the
is
species
25 flowers which
specimen
Peninsular
—
Note —This
c.
pu-
halfway
abaxially,
ovate-orbicular with
with
whorl
a more
an
distinct claw,
minutely puberulous.
connective apices truncate; anthers latrorse-introrse.
Carpels
392
c.
BLUMEA
2
mm
stipes
1
Ripe
Seeds 1
long.
cm
to
wrayi
var.
Peninsular
is
1-1.5
long;
cm
Malaysia.
the
recognised by
which become
long peduncles
cm
10 mm
by
Cyathostemma
2000
2,
glabrous, ovoid, reddish,
monocarps
Note
—
No.
45,
2.
or
Distribution —Vietnam
with 2
b.
long, pubescent.
c.
Vol.
almost
large
glabrous leaves,
and tuberculate, and the
woody
large,
10
c.
petals.
indochinensis Ast
Cyathostemma wrayi
15387
var.
indochinensis.Ast, Notul. Syst. (Paris)
(P n.v.), Cambodia, Mimot;
Poilane
21936
9
86.
(1940)
—
Poilane
Syntypes:
(P), Vietnam, Annam; Poilane
22517
(P n.v.),
ibid.
Climber. Leaves acute-acuminate, 25-28 by 5-9
1-1.5
carps red,
Distribution
Note
acumens,
orbicular, and
acute at
the range of
leaves
10.
never
Sin.
30
to
c.
Hu,
Ban,
Yen
cm,
arching
drying
rounded
and
and the ovary is
11.
—
or
Peduncles absent;
tam.
halfway along
6-7
by
5-7
2.5-3.5
cent
mm,
mm
to
along
monocarps
their
prominent
green
or
Note
—
near
1157.
(1974)
oval-
are
the base. The
at
within
acumens
greater in number and the
(1940)
is distinct.
variety
121; Tsiang
(holo
PE n.v.;
—Type:
To Thuc Vat
at
the
becoming
&
iso
P.T.
Li,
F1
GH), China,
connective
pubescent
long;
at
mm
long;
lamina obovate,
12 or
nerves
long;
the
ovules
re-
pairs,
5—6(—8)
mm
cymes.
sparsely pubes-
Petals 6, coriaceous,
apex
13
extra-axillary
bracts
above,
7—17(—20) by
channelled above,
cm
(holo
veins and
secondary
bracteoles similar, attached
c. one
broadly
third
ovate,
both inside and outside. Stamens
apices bluntly ligulate;
3.5-4.5
veins and reticulate
few-flowered in
1-1.5
1823
ibid.
when mature,
glabrous
cuspidate;
margins; petioles
Thuc Vat
(LE),
chartaceous, glabrous except
secondary
the base, apex
To
1823A
midrib and elevated
pedicel. Sepals 3, pubescent.
length,
Tsiang
of ovules ( C.
59
Paratype:
pedicels sparsely pubescent,
apex obtuse,
—
10
74879
pale green-yellow;
sub-cordate
pubescent, ripe
Distribution
Inst. Biol.
Type: Wang
glabrescent. Inflorescence
the
long;
narrower
whorls
1
twigs pubescent,
cent, foliose, ovate-orbicular, 4.5-5 mm
to
which has
are
establish ifthis
to
Map
Mem.
Bot. Zhurn.
bai, Dong
the
forming loops
long, pubescent
Fan
—
petal
glabrous only
veins
secondary
the slender elevated midriband
ticulate beneath,
veins. The
and striate; lenticels indistinct. Leaves
sparsely pubescent along
5.5-9
Hu
t.
long. Young
m
purple-brown
puberulous along
Bull.
(1979) 28,
vietnamense
3
mono-
Fo-Hai
province,
LE), Vietnam, Tonkin,
dark
the
because of the
wrayi
var.
secondary
is Poilane 21936 (P),
seen
yunnanense
yunnanense
Reipubl. Pop.
Climber
tip,
the
wrayi, although
Cyathostemma
Yunnan
more
thinner. The author is unable
slightly
Cyathostemma
and
the author has
var.
Cyathostemma
differs from
variety apparently
leaves, much longer
only specimen
pairs. Ripe
Cambodia and Vietnam.
—
This
—
15-17
cm, nerves
long.
cm
anthers introrse.
(4—6)—14
in 2
Carpels pubes-
rows.
Half-grown
monocarps unknown.
China and Vietnam.
& P. T. Li (1979)
vietnamense 4-6,
note
that C. vietnamense differs only
C. yunnanense 14), but
they
were
in
the number
unable
to
see
the
T. M. A.
material.
type
the
same,
allied
C. argenteum,
to
two
taxa
the
type material of both
difference
only
red. These
differing only
may be
being
more
they
yunnanense
needs
two taxa
to
the
are
closely
chartaceous, petals
C. yunnanense may be better
or
is
never
placed
as
a
be collected,
from Vietnam, Thailand and South China, before any taxonomic decision
especially
be made.
DUBIOUS
acuminatum
Cyathostemma
Gard. (Calcutta) 4, 1
J. Asiat. Soc.
King,
(1893)
TAXA
13.
—Type:
Note —
type has
This
taxon
been
not
that from the
is described
although
seen,
description
Uvaria borneense
other material of
it
seems
very like C.
Bot.
15 (1929) 64.
iso BM, IBSC, L, NY
Uvaria lambirense
n.v.,
—
note
SAR n.v.),
Neither of these
and reflex
enlarge
that
T.
at
borneense
not
—
a
possesses
nov.
hirsute
designated;
—
Tetrapetalum
lambirense
Momose 5069 (holo KYO
n.v.;
Lambir Hills National Park, Miri.
Cyathostemma
stigma,
a
because of the
in Uvaria. It is
placed
better
specimen
borneense Merr.,
Tawau.
K.
Type:
are
Tetrapetalum
—
comb.
members of
taxa are
anthesis, and
of this taxon, and noted
Elmer 21211 (lecto K, here
Malaysia, Sarawak,
members of Cyathostemma. The type
material which is
—Type:
117.
integrity
TAXA
nov.
(K. Momose) Utteridge,
examined has
Cyathostemma
wrayi.
P), Malaysia, Sabah,
K. Momose, Blumea 43 (1998)
(photo!),
comb.
(Merr.) Utteridge,
Univ. Calif. Publ.
Note
Bot.
Roy.
traced), Malaysia,
not
and female flowers. The
having hermaphrodite
as
no
EXCLUDED
which
3468 (?CAL
flowers. Sinclair (1955) doubted the
gynodioecious
iso L
61 (1893) 10; Ann.
Bengal
Wray
Perak.
Upper
to
Cyathostemma
in leaves
conspecific,
yunnanense here, since
and concludes that
taxa
ovule number.
being
393
Cyathostemma
genus
of C. argenteum. More material of these
subspecies
can
the
seen
of
vietnamense is included in C.
Cyathostemma
the author has
Revision
Utteridge:
character
of T. borneense
at
not
petals
interesting
found within
BM includes
fruiting
Annonaceous.
Cyathostemma grandifolium
K. Schum. & Lauterb.
Haplostichanthus longirostris
=
Heusden.
ACKNOWLEDGEMENTS
I thank
the
material
keepers
of
their material
their
help
Baker
work
on
and
and Paul
was
of
loan.
and the
Thanks
Bygrave
for their
while I
K
was
and P for
herbarium
to Paul
supervision during
completed
studentship.
BM, C, IBSC,
Cyathostemma,
the
KeBler
allowing
keepers
(L),
undertaking
helpful
receiving
of
me
A,
Mark Coode
of this
Science and
(K)
project.
comments whilst
a
to visit
B, BM,
I
was
their herbaria
GH,
L
and David
I would
Moore
also like
preparing
Engineering Research
and
and LE for
this
examine
sending
(RNG)
to thank
manuscript.
Council
for
Bill
This
postgraduate
394
BLUMEA
Vol.
No.
45,
2000
2,
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apocarpy
117-120.
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1929. Plantae
A.G.
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E.D.
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1993. Annonaceae.
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P.J.A. & E.C.H.
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1-131.
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flowers
and
translated
inflorescences;
by
R.J. Pankhurst.
Cam-
bridge University Press, Cambridge.
List
of
collections
7
Cyathostemma
I
2
3
Agama
AA
1081:6
284:
—
11;
60039:
4
BRUN
606:
—
=
excelsum
=
9
10
glabrum
=
hookeri
=
viridiflorum
=
wrayi
=
yunnanense
=
5
=
longipes
11
=
6
=
micranthum
12
=
Ambriansyah
Anderson
2
S
—
&
BRUN 5167:
Uvaria
Arbainsyah AA
Amin
16417:
2.
siamense
=
4
AA 429:
2;
8
argenteum
2
SAN
—
95310:
987:
2;
Arbainsyah
1;AA
SAN
et
borneense
lambirense
1660:2
105998:
al. AA
1945:
—
4
Ambriansyah
—
2;
Amin &
AA
& Arifin
Joseph
1987: 2
—
SAN
Ashton
T. M. A.
Bakar
SAN
25044:
Brand SAN
Carr
11088:
31146:
1582: 6
38:
2;
105:
2;
SAN
17397:
Geesink &
8
Haniff SFN
422:
2
1968:
11;
Ismawi S 37445:
Kadim
—
6
Kidh
6482:
mans
Lau 535:
428:
7;
Madani SAN
174:
FRI 5492:
KEP
27210:
6;
5
6
2
—
9
4
Van
—
9
—
11;
12
10;
195:
8
Curtis
—
21341:
2
50:
—
1213:
2;
26541:
1;
Collins
—
4;
4;
162:
6;
2808:
6;
19330:
19554:
8;
19917:
2;
8
Dewol
—
SAN
1551:
2234:
2;
9
& Santisuk
5662:
1
Gibot SAN
—
93980:
2
8.
14189: 9
Haron
—
Haniff &
S 21376:
711: 6
6;
&
Kenneally
2
—
SFN
Nur
Hashim
—
Hillard
10904:
Hyland
12761:
9;
collector
2
12966:
6;
307:
—
6;
4269:
5
KL
Hardial
—
K.F. No. 4936:
9
&
Haviland
—
1744: 2 —Hose
6
—
Meijer
Mujin
100032:
601:
11.
1103:
4635:
4207:
4;
2299:
8;
75-501:
11
2
—
—
6;
2;
3250:
75-564:
Merrill
Mtiller
8
182: 8
75-658:
6;
1180:
—
6
—
332:
2
Mikil
—
2.
8.
Marcan
76-540:
6;
2;
Koster-
8131:
2;
& Ashton
5981:
—
Lefevre
76-390:
8;
1
1557:
6;
PK 1190:
6;
5857:
6210:
9;
1;
9271:
Murthy
—
18767: 8
Mansus & Amin:
—
1;
3345:
2
5862:
2;
6 —Korthals
1665:
6;
8;
PK
2;
5451:
9;
4151:
15133:
6;
8;
2
44321:
S 22619:
6;
—
2.
2,
Pascual
1048:
12426:
2
2
—
Petelot 6.421:
—
15367:
6;
30612:
36:
32834:
33826:
6;
14876:
6;
PK
2;
4;
9 —Kooy 339:
110:
Kerr 2291:
—
13107:
Leeuwenberg & Rudjiman:
6;
75-396:
4047:
6;
Kunstler
11 —
S 52340:
—
6;
—
15387:
6;
Purseglove
4;
1667:
Ridsdale
2
—
Sanusi
5
2
Pierre
—
16375:
9;
P 4965:
4;
BS 41225:
CEROL
10: cf.
al. SAN
56772:
SAN 30015:
118670:
878
—
6449:
10
2
2
—
—
Put
209:
17488:
6;
1421:
6;
6
6;
Podzorski
—
21936:
2291:
9;
3137:
6;
SMHI
22517:
6;
9;
3173:
2;
6
Ramos &
—
SMHI 1619:
3;
Edano
SMHI
BS 44124: 6
1866:
6
—
—
Russell-
4383: 6
Tukirin
—
6459:
1548:
—
6;
1;
Jr.
8
Sinanggul
&
—
—
Scortechini
1316: 9
PBU 422: 2
54532:
Reynoso
6
880
4
—
—
Shah & Kadim MS
Sidiyasa
& Kochummen
Sinclair SFN
—
6242:
al. 2558:
et
1842:6;
—
(KL 3480): 9;
Teo & Bruno KL
1;
6
—
Soejarto
39246:
8;
al. 6407:
et
SFN
6
—
2
(KL 3543):
—
Thorel
2
8.
9
—
389: 6
Talip SAN
—
To Thuc
1.
—
Winkler 2548:
1166: 4.
Zollinger 961:
1043
3420:
1887A:6 —Van Steenis
6477:
23: 8.
al. AZ 5158: cf. 8
—
Sidiyasa
Soejarto
6468:
1837:
Wray
—
2.
VanBeusekom
6;
—
2
(KL 3478): 9;
Teijsmann
6
bin Tahir 9749:
SAN 76088:
Minjulu
1823A;
8 —Winit
Yuang Guang
1480:
—
SAN
6127:8
6430:
et
et
Singh
SAN A4632:
Zainudin
Endert
Maingay 29: 6;
(KL 3420): 2;
cf.
Balgooy
3128:
Clemens
—
2
1929: 3.
Sigin
1823:
Wallich
—
4733:
11;
Lee
4790: 4
Shea &
—
T. & P. 820
Vat
4
520:
2.
6.
Heifer 433:
King's
Ramos
—
Sumbing Jimpin
65449:
—
Kochummen FRI 26243:
8
Prance
Smith & Lucas
39341:
Bernstein
—
1034: 8.
2314: 2 —KeBleret al. PK 819: cf.
—
—
Saikeh Lanteh 67224:
2
4
& Saikeh SAN 70051:
Corner SFN 29460:
SFN
4;
9613:
Maxwell
KEP
Ridley 2112:8;
565:
1811:
Buwalda 6617:
—
8.
332:
1022: 6
—
—
—
Poilane
—
11
1837:6
5;
cf.
—
1744: 2
2;
—
4310:
Ramlanto
4222:
99276:
2171: 6
—
85-771:
8;
1
2;
8
92017:
Millard KL
Pagi
4
2708: 6
6;
80-211:
Ng
1282:
1751:
5;
Cockburn
De Wilde
—
Guinet
—
3678:
8607:
35304:
8526:
4;
551:
6;
KeBlerPK
—
3
etal. 821:
Church
—
Beccari
—
4922:
Geesink, Phanichapol
—
2250: 4
2;
2
Brown
395
Cyathostemma
genus
4.
7843:
6;
6
Harmand
& Noor KN
7419:
9
Dilmy
—
21211:
2;
SFN
6;
—
179:
9;
2
433: 6
8;
1068:
Sidek417:
107:
7848:
Hiepko
Griffith 432:
—
3810:
—
—
—
Fukuoka T-36219:
—
the
of
8.
99417:
21081:
6;
Falconer 534:
cf. 2
7661: 2
2;
3505:
9;
1077: 9
32081:
6;
Coode 6758:
—
80810:
Elmer
Brass 8557: 6
—
6 —Chin
31534:
2;
3015:6; 3018:6;
David
2
BNBFN
Balajadia
—
11509:
6;
cf.
2
30855:
Revision
Utteridge:
Wang 74547:
8
—
Wong
10;
308:
1106:4—Vidal4110:
74879:
8.
10 —Whitmore
2 —Wood
SAN
16166:
BLUMEA
396
Vol.
45,
2,
No.
2000
Index
The number after each
=
excluded
Anaxagorea
sumatrana
Cyathostemma
acuminatum
Griff,
names are
King
1
Thomson)
ex
Popowia
J. Sinclair
Utteridge
3
(excl.)
2
borneense
J. Sinclair
(excl.)
in
Heusden
bold.
(excl.)
2
6
(excl.)
Momose
(excl.)
I
L.
argentea
8
Blume
7
1
1
auct.
(Merr.) Utteridge (excl.)
Roxb.
1
Boerl. 6
(Miq.)
Ban
viridiflorum
Griff. 8
excelsa Wall.
(King)
Ridl.
8
glabra Span.
glabra auct.
9a
var.
indochinensis
var.
wrayi
Merr.
confertiflora
10
scortechinii
Ast 9b
9a
Hu
2
6
argentea
vietnamense
yunnanense
K.
Miq.
bracteata
Guatteria
taxa;
names
A. DC. 6
King
Merr.
lambirense
bomeense
var.
new
5
(A. DC.)
King
wrayi King
,
excelsa Hook.f. & Thomson
nitida
volubile
Utteridge
sumatrana
dubious
=
italics
4
Bakh.f.
siamensis
(dub.)
in
Tetrapetalum Miq. [p. 381]
Uvaria
scortechinii
the text;
Polyalthia fruticans
K. Schum. & Lauterb.
Craib
micranthum
nitidum
in
synonyms
Mitrephora
J. Sinclair
&
(Hook.f.
species
roman,
Haplostichanthus longirostris
King (dub.)
grandifolium
longipes
in
6
Miq.
glabrum (Span.) Jessup
hookeri
of the
printed
[p. 381]
argenteum (Blume)
excelsum
is the number
name
Accepted
taxa.
10
micrantha A. DC.
3
4
gomeziana
A.DC.
lambirense
(K. Momose) Utteridge (excl.)
micrantha
6
2
2
parviflora
(A.DC.)
Hook.f.
1
Hook.f.
& Thomson 6
& Thomson
4