Squamellaria apical `/ 2 stem apparently arising (fig. 3

advertisement
36
The
BLUMEA
53-61
(1991)
tuberous
epiphytes
A
of
revision
the
of
Institute, P.O.
4:
Squamellaria
M.H.P.
Christensen Research
Rubiaceae
Jebb
Box
305, Madang, Papua
New Guinea
Summary
Squamellaria Becc.
cies
is reduced.
Confined
The
the
to
is revised to include three
genus
on account
stigma,
of
Hydnophytum Jack,
fringed plates (squamellae)
of the tuber has revealed
Squamellaria species
a
has the
holes the tuber wall, of S. thekii in
ties. Each
cavity
in outline.
crenate
appears
cavity
is
and the base of the
a
Girdle Chambers interconnect
while
along
dorsally
and
their
and
basic
encircling
at
gives
rise
rise
apically
arise
only
at
'/3 of the tuber,
are
most
stems
are
In
apparently arising
appears
to
to
'Deep-lying
being
between each
of
be associated with the formationof
thickened and unbranched.
with
cylindrical,
are
similar cavi-
recognised (Jebb,
of
This chamber
entrance
and
edge,
blind-ended and
'Superficial
holes. The
to
Smooth Walled
From its inner
Smooth Walled Tunnels'
stems
to
entrance
a new
in that the
arise from the
the apex
holes
cavity.
one
(fig. 3);
In this
(fig. 1).
stems
apical '/ 2
(fig. 2). The
from
younger
densely waited,
previous cavity.
stems are more remote
ring
The
Hydnophytum.
it is
Hydnophytinae
clustered close
S. major the
capitate
structure, which fur-
(fig. 1).
their basal
numerous
the tuber
S. imberbis the
apex. In
slender and branched.
stem
rise
origin-
closely related,
essentially
ring
to a
along
also differs from the rest of the
Squamellaria
not
genus
structure:
'Girdle Chamber'
to
is the
species,
the tuber. Between these rings of
Chambers' which overlie the Girdle Chamber of the
gives
spe-
swells outwards, such that the tuber
apical edge. Ventrally they
wall the Girdle Chamber
existing
the tuber apex, between the last formed
older cavities
basally they give
three
most
and
Several distinct chamber types
to
one
bony endosperm.
tuber contains several,
superficial planar
all but encircles the tuber, and
cavities
same
particular,
A mature
stem.
just
which it is
to
embryos
produced sequentially
1985). Each cavity has
with
remarkably complex cavity
rimmed holes,
regularly spaced,
of which new, and
inside the corolla tube, the
the taxonomic 'moat' between this
strengthens
tuber of all
rings
this genus from
of the four
one
& Jebb, 1991). Beccari (1886)
Hydnophytinae (Huxley
and the curved pyrenes with short
Study
ther
species,
Fiji.
Fijian Islands, Squamellaria,
eastern
smallest genus of the
ally separated
is endemic to
to
stems are
another,
thus each
species
do
the
one
stem
stems
54
BLUMEA
1. Schematic
Fig.
Becc.
a.
cavity;
Tuber
e.
chamber;
The
f.
h.
deep lying
stigma
waited
and anthers
are
of
Scale bar
opening
or
sile inflorescences and
also
Squamellaria,
Squamellaria
10
stigma
shows
is
and
capitate
Wise.
finely fringed
and the
large capitate
No observations of
interesting
that
in the
Fijian species
of
noc-
solitary
relatively large (110-163 |im) four-colpate pollen,
occur
of later
presenter mechanism. The
pollination.
have been made. It is
(A. Gray)
smooth walled
by Rosemary
heterostyly,
a
imberbis
d. entrance hole
superficial
g.
Drawn
cm.
above the anthers, suggests
production
scent
of
preceding cavity;
=
exserted. The
species
1991
1,
girdle chamber;
often curved corolla tube suggests moth
turnal
in
extremities
None of the
edge.
hole of
entrance
c.
smooth walled tunnel.
stigma, lying immediately
long,
hole;
No.
36,
within a tuber
single cavity
b. entrance
girdle chamber;
the lower
along
of a
diagram
apex;
VOL.
as
ses-
found
80-130
Hydnophytum, (pollen
|xm across).
Beccari's
new
The
present
and flower
original
and
species
species
two
here reduced
are
that all
reported
had
species
author collected all the
species
a
to one.
in 1985 and
All material is cited, marked
structure.
brackets afterFlower in the
descriptions
Smith (1967) described
chambered tuber inhabited by
was
if
n.v.
able
not
to
study
seen.
The
a
ants.
the tuber
figure
in
is the number of flowers dissected.
SQUAMELLARIA
Squamellaria Becc.,
4
(1891) 123;
A.C.
Smith,
Malesia 2
A.C.
Smith,
Fl. Vit. Nova
mecodia imberbis A.
Epiphytes.
J.
4
Arn.
(1988)
cylindrical,
blackish, occasionally with
prominent,
grey
to numerous,
or
36
—
Engler & Prantl, Nat.
(1955) 289;
S.
Darwin,
Type species:
S.
imberbis
numerous
rings
to
oblong.
numerous
Pflanzenfam.
Allertonia
(A. Gray)
2
IV,
(1979) 35;
Becc.
[Myr-
(1858) 42],
Spines simple, slender, mostly
around entrance holes. Surface smooth,
white tubercles
around the tuber,
to
4
to
mm
2
mm
in diameter. Entrance
in diameter, conical,
or
with
white, spine- fringed rims. Pores and ridges absent. Stems few
scattered
woody, branching
245.
ovoid
flexible, scattered, sometimes more
holes in discrete regular
K. Schumann in
Arbor.
Proc. Am. Acad. Arts & Sci. 4
Gray,
Tuber
(1886) 228;
or
or
arising
not,
to
50
from
cm
one or
long,
more
1.5
cm
basal
stock(s);
subherbaceous
thick, rounded, smooth,
to
mostly
M.H.P. Jebb:
Leaves
lacking spines.
clustered
A
revision
mm,
the
splitting opposite
petiole
at
to
with 4
Bracts
lobes. Corolla
towards middle,
markings
quadrangular
the corolla lobes,
on
wards base of tube,
pate, 110-163
occluded
in diameter,
pm
pm; vesicles absent.
Ecology
species
a
ridge
High
-
on
2a.
at
grain,
Petiole 0.4-3.5
Iridomyrmex
TO
THE
SPECIES
to
margin irregularly
undulate. Stems
0.5
cm,
Tuber apex blunt. Stems
depressed
at
Tuber apex
Leaves
apices.
6-30
apex,
tetra-
dense forest from 200-2000
to
OF
Stems
to
rhomboid, thin, with
laminaobovate
flat
a
1. S.
or ovate
markedly
to
margin.
imberbis
cordate, thick and fleshy,
thickened towards their apices
in middle, with
1.5
to
cm
in diameter
cordate, apex
ovate to
acute.
0.8
to
cm
2
their
near
acuminate.
minutely
red and green
in clusters,
arising
m.
SQUAMELLARIA
arising individually,
parallel-sided
tapered.
to-
less hooked,
more or
markings
2. S.
b.
or
red
partially
coarse,
siccate, markedly
or
Leaves obovate, the apex rounded and sometimes
Corolla lobes
open,
thickened towards their apex
Petiole very short,
apices.
or
green
ants.
lamina lanceolate
cm,
not
with
or
base
at
sometimes
reticulation
fleshy
entire
mm,
plates (squamules)
indistinct lobes,
Fruit
margin.
In-
petioles.
the abaxial surface.
inhabited by
are
Stems slender,
b.
fringed
±
with 4
the
as
3.5
to
exserted. Pollen 3-5-col-
mm,
pm;
low in host tree, in open
or
KEY
la.
4 barbate
long
as
cordate,
cm,
sharply
or
ovoid, orange-red; calyx remains prominent. Pyrenes 4,
or
sometimes with
All
almost
Stigma capitate,
recurved and sometimes
gonal
colpi
wall extensions; walls 9-14
by pollen
2-6
distorted, curving
pubescent without;
6.5
first, becoming pedunculate
at
inflated, white throughout,
or
x
thin, papery,
homostylous. Calyx
commonly
lumen. Anthers 4-8
occluding
17
-
between the
tearing
soon
in axil, sessile
Flowers
inconspicuous.
triangular
1.5
x
thick, fleshy. Stipules
first,
florescence solitary, laterally displaced
with age.
Lamina 3.5
or remote.
rhomboid-lanceolate or obovate, thin
55
of Squamellaria
in diameter
Corolla lobes obovate,
major
near
pale
their
green
3. S. thekii
1.
imberbis (A.
Squamellaria
imberbis
Squamellaria
Am.
Arbor.
Smith, Fl.
& Sci. 4
Hook.
Benth.
f.
Vit.
in
&
Squamellaria
Benth.
62266
wilsonii
J. Am.
nud.;
200.
—
247.
ex
—
Seemann,
f.,
Hook,
Malesia
Vit.
PI.
2
111. Fl. Ins.
Drake,
2.
Fig.
(1886) 228,
Fiji
Myrmecodia
Fl.
Gen.
2
PI.
Parham,
—
(1866)
Isl.
imberbis
138.
—
Pac.
46, figs. 1-12;
ed. 2
A.
132.
(1890)
—
200.
A.C.
Am. Acad.
inermis A.
Hydnophytum
—
Smith,
J.
(1972) 293; A.C.
Proc.
Gray,
Myrmecodia
sphalm. (1873)
Mar.
t.
(1964) 209,
Type: Fiji,
Arts
Gray
ex
imberbe
U.S.
Expl.
(holotype US; isotype K).
(Home
Arbor.
Home
J.W.
(1988)
B.C.
&
Hook.f.
(1988) 247, figs.
(1890)
4
Nov.
sphalm. wilsoni;
nom.
(A. Gray) Becc.,
(1955) 289;
(1858) 42;
Exped. U.S.
Smith,
36
Becc.
Gray)
S.
36
Darwin,
9A-D &
ex
J.W.
Allertonia
13a.
Baker,
Type: Fiji,
Baker) Becc.,Malesia
ex
(1955) 289;
—
Soc.
2
(1886) 229,
PI. Fiji
(1979) 35, fig.
Hydnophytum
J. Linn.
Home
2
Parham,
Bot.
wilsonii
20
1139 (holotype K).
Isl.
3D;
A.C.
Home,
(1883) 365;
t.
46, figs.
(1964) 209,
ed.
Smith,
A Year
Drake,
in
13-21;
2
Fl.
A.C.
(1972) 293,
Vit.
Nov.
4
Fiji (1881) 263,
111. Fl.
Ins.
Mar. Pac.
56
BLUMEA
VOL.
36,
No.
1,
1991
Wise.
Rosemary
by
Drawn
467).
Jebb
(all
pyrene
d.
fruit;
c.
flower,
b.
plant;
Whole
a.
Becc.
Gray)
(A.
imberbis
Squamelari
2.
Fig.
M. H. P. Jebb:
Tuber
in
liptic
glossy
longitudinal section,
conical, 0.6
to
green
4
tapering
veins
c.
to
5-7.
apiculate,
a
to
pedunculate,
branching,
1.5
x
8 5
-
3.5
to
long
cm
and 0.3
mm,
mm,
pubescent without, especially
Corolla tube
mm
2
a
long,
finely
more or
acute,
pm
the foot
across,
diately
3
to
pm,
layer
the
Ecology
given
Notes
unarmed
late this
in fruit,
and
-
The
holes is
and
from
the
wilsonii
to
was
the tube,
limb
470
vagaries
of
Smith
to
Navitho
between
J.W.P, P.B.T.
8326
1787
Lambasa,
to
pinkish;
long,
mm
sessile
pm
32
to
rarely glabrous.
so,
6.5
triangular,
in diameter;
of the
to
[11], Calyx
quadrangular,
base. Lobes
grain;
to
with
colpi
walls 9-10 pm
these lumina 6-16
perforations. Stigma
imme-
of short hairs below. Fruit
fringe
mm
3.5
long, blue-black; filaments
height
including calyx. Pyrenes
several
solitary,
5
mm
specimens usually
present
16935
longer
more acute
16°
481
50'
—
Vanua
gradually
light
more
attenuate
globose,
(Beccari,
of further collections and
proved insignificant,
SE
16°
Jebb 467
471
(BISH, K),
180°
179°
468
178°
or
due
to
py-
1886),
study
of
developmental
Wairiki,
00',
on
Smith 564
(n.v.).
Smith
750
of Mt
Manuka,
road
to
19', Thakaundrove,
25'
Korotini
(LAE, SUVA),
(BISH, FHO, K, SUVA),
Ndelaikoro,
and
30'
18° 04
472
—
Devoeux
Peak, Jebb
Freehold
uncertain.
DA
11901
Seemann
s.n.
(n.v.).
Forest
(K, P).
Mathuata
River
road
near new
179° 58
Range,
Western
Qacavulo Estate, A.C.S,
Smith 8191
479
(GH, L, UC),
(BISH, K, SUVA),
Above
Creek,
Yanawai
(BULOLO, K, SUVA),
16°48
Above
Wairiki,
Range,
Crest of Korotini
(SUVA).
(BO, H, UC).
east of
469
Taveuni.
(BISH, K, SUVA), 482 (LAE, SUVA, UPNG).
Locality
its shorter corolla, lobes
than filaments, fruit
and
In the
Levu.
(BRI). Slopes
(n.v.). Nggarawalu,
(type).
tuber, with fine-rimmed
The thin, lanceolate leaf and flowers also iso-
(BO, GH, K, UC).
and Mt
Somosomo
(K, L, UC).
wilsonii).
62266
Pass
D.K.
(BISH, SUVA),
16935
7
to
drying.
(A, BRI, CANB, SUVA),
slopes
to
green to
Flowers
densely
formerly distinguished by
stamens
(Smith, 1988).
FIJI ISLANDS.
-
Kasi,
Savusavu
between
cm.
slightly compressed
diagnostic.
hooked, and leaves
Collections
Mt
0.1
on
curved.
Not
m.
regular, oblong,
longer calyx
region,
to
mm
a
most-
elsewhere
rare
Inflorescence
Corolla white,
the
high,
cm
rhomboid, apex acute,
layer regularly reticulate,
fleshy,
fresh material, these differences have
or
caducous.
110-152 (134.5)
as
0.4
from the remainder of the genus.
species
relative
stages,
cm,
base, tapering
outer
not
tuber,
to
pink. Stipules
thick. Bracts
strongly
Altitude 300-700
-
entrance
less
renes
0.15
to
upcurving;
tree.
Squamellaria
longer
to
green
long
to
prominent below, pale
corolla lobes, bud
at
to
towards the apex. Internodes up
within the lumina with minute
turbinate, strongly quadrangular,
a
pendent
lanceolate
cm,
Entrance holes
protrusions.
close
very
Anthers 4-5
as
cm,
remote
pubescent.
on
in bud.
0.6
x
wide; surface
cm
surface; rims
upper
above anthers, with 4 indistinct lobes and
long, tighdy appressed
in
cm
less recurved
thickened border, about
slightly
to
(3-)4(-5)-colpate,
thick, the inner wall
3.5
pale
mucronate,
tetragonal
wide. Pollen
mm
x
cm,
2-3
or
50
to
small
hori-
nearly
from the side, el-
and 12
high,
cm
on
more numerous on
small central process
0.6
entire
mounted
mid green. Midrib
petiole,
18
long,
cm
cm
Spines only present
Petiole 0.4
with
30
2
subovoid
to
bluntly hemispherical
of the tuber, less
apical part
when sterile.
Leaves. Lamina 4.5
stem.
base
the
on
long
cm
Spines
in diameter,
cm
to
57
of Squamellaria
regularly oblong
less
more or
to
grey. Stems several,
ly arising
to
black.
to matt
revision
from host,
growing horizontally
zontal below, swollen above,
A
Horne
Prov.,
480
Nggathavula Estate,
1139
U.S.
(K) (type
DA
of S.
Expl. Exped.
US
58
BLUMEA
VOL.
36,
No.
1,
1991
Wise.
Rosemary
by
Drawn
483). Jeb
b—d:
475;
Jebb
(a:
fruit
d.
bud;
c.
flower,
b.
plant;
Whole
a.
Smith,
A.C.
major
Squamelari
3.
Fig.
M.H.P. Jebb: A
2.
Squamellaria
major
Squamellaria major
ed.
A.C.
2191043
Smith,
A.C.
(1972) 293;
2
A.C.
Smith
Fl.
Vit. Nov.
cm
cylindrical, obovate,
in diameterwith
Stems several,
nodes 0.5-1.5
cm,
Lamina obovate
near
from
to
20
narrowed
7
to
inflated towards middle,
to
2.5
at
to
0.5
at
the
pollen grain wall,
to
colpi being
with 4
or
3
x
-
pale
a
red
pm,
the foot
17
x
to
pm
6.5
0.3
or
on
trunks
The leaves of this
Collections
-
8323
483
5
to
at
mm
Tuber
fusiforme,
incrassati
orientes.
the centre,
paralled
when
Folia
16°
tumidum
apicem
to
mm.
Corol31
about
by
edges
6 pm, the
a
pm in diameter;
long
as
the
as
number of fine
of the
outer
mm
Tube
mature.
towards apex, and then
125-163 (140)
across,
or
wall
mm;
colpi
height
of
outgrowths
of
and
colpus
interdigitate;
irregularly reticulate,
small
numerous
large
limbs of trees in closed forest
species always
180°
out-
to
3
mm.
to
Pyrenes hooked,
at
300-400
m.
appear chlorotic and diseased. Exami-
00',
cover
the
out-
in them.
SE
on
16°
50'
road to
Jebb
Jebb, spec.
inter
versus,
fasciculata,
cotypus
50'
thekii
ovoideus, calyce
BISH, K;
35
above the disc. Anthers 5-6
prominent calyx
fungal hyphae
margine corrugata. Calyx
Fructus
to
long.
179°
59' Slopes
Devoeux
Peak,
of Mt
Jebb
Manuka,
475
east of Wari-
(FHO, SUVA),
476
(SUVA).
Squamellaria
paullo
apices.
at
Margin irregularly
with
FIJI ISLANDS. Taveuni.
(type).
(BISH, SUVA),
3.
entrance
few in number.
side of the tuber have
Smith
cm across.
apex rounded,
cm;
nation of fresh tuber material reveals that the small white tubercles that
ki,
Entrance
of
ring
sessile. Flowers [3]
within the lumina with
layer
Surface
cm.
long.
cm
each venation branch, often
at
narrowing
less occluded
to
x
pink patch
or
mm
5-colpate,
14
17
-
gentle undulations,
which arise from both
Growing
-
solitary
Note
US
=
mm.
Ecology
Often
Isl.
Fiji
smooth, grey; inter-
above and below.
in diameter,
to
more or
fleshy, quadrangular below,
5.5
PI.
8323
square in section with 4 mounds, exserted above anthers. Fruit
growths. Stigma
to
surface
rounded in bud, sides
12 pm thick, the inner wall
these lumina 9-30
cm,
Smith
cordate, drying thin, yellowish
or
Inflorescence
cm.
mm
Pollen 4-
mm.
grain,
1
7
to
4.5
x
blunt
mouth; barbate scales 2
the
walls
8
prominent
margin,
barely thickened, colpi
the
1.5
to
within each
fleshy,
cm,
narrowing abruptly
pubescent,
mm,
widening slightly
1.5
x
la recurved, white, lobes green with
lobes
24(-40)
to
flexible,
to
immediately
abruptly narrowed,
Lateral veins
apex.
Calyx cup-like,
apex blunt
broadly oblong-elliptic,
to
undulate, white. Petiole 0.3
edges
Parham,
Type: Fiji,
—
shortest and thickest towards apex. Leaves clustered
not, base
or
filaments
J.W.
(1967) 90;
247.
thickened, spine-topped, white rim
a
green above and below. Midrib
long;
37
(1988)
stiff
Spines
apparently arising
holes, unbranched, scattered,
bifurcate
4
(holotype US; isotype BISH, K, L, US).
Tuber subpendent,
apiculate
3.
Fig.
—
59
of Squamellaria
Cont. U.S. Natl. Herb.
Smith,
with small white tubercles, black.
holes 1
revision
annulos
ad
obovata
8
mm
usque
manifeste
—
Fig.
foraminum,
ovata,
coronatus.
apice
alba,
—
lobi
4.
qua foramina
diametro, ramosi,
4-dentatus. Corolla
478, SUVA).
nov.
acuto
ab tubere
usque
tumidi in
Typus: Fiji,
margine
uno
rotundo,
alabastro,
Jebb 477
incrassato.
fasciculo
basi
Caules
vel
pluribus
obtuso,
carnosa,
obovati
in
maturitate.
(holotypus SUVA; isotypi
60
BLUMEA
VOL.
36,
No.
1,
1991
Wise.
Rosemary
by
Drawn
477).
Jebb
(all
pyrene
f.
apex;
fruit
at
disc
and
calyx
e.
fruit;
d.
bud;
c.
flower;
b.
plant;
Whole
a.
thekiJeb .
Squamelari
4.
Fig.
M.H.P. Jebb:
Tuber
surface
5
cm
1.5
numerous
Stems several,
across.
36
0.8
x
a
Stipules rounded,
Bracts papery,
to
0.2
less
to
cm.
0.2
to
one
to
long,
9-15
rise
giving
wide;
cm
hair-like
to
in
numerous,
the inner edge
cm,
few
base and apex, ab-
to
cm
some
than 0.1
to
lobes,
31
to
Corolla tube
mm.
ments
8
to
mm,
unbranched,
to
3
Pollen 4
mm.
ened; walls
to
14
pm,
these lumina9-22
anthers,
2
to
mm
colpate,
colpi
4.5 pm, the
to
round the foot
deep
layer
in diameter,
on
groove
the
above the disc. Anthers
150-163 pm;
the inner wall
pm,
a
mm
19 pm
to
wall
outer
3.5
across,
to
6.5
x
3
fila-
mm,
thick-
edges
irregularly reticulate,
within the lumina smooth.
slightly recurved,
mm,
on
gently
surface,
outer
to
with
10
to
mm,
abaxial
prominent
a
above
Stigma
in diameter, rounded, margin reflexed and fimbriate. Fruit
ovoid, fleshy. Pyrenes
to
0.2
fleshy, quadrangu-
minutely pubescent
6
to
to
sessile.
Inflorescence
mm,
Corolla white,
with
mm
5
to
Calyx
towards middle,
less orbicular in bud, scales 2
more or
mm.
margin.
at
swelling
recurved. Lobes obovate, blunt,
36
to
0.8
apex. Lamina cordate
at
caducous.
long,
cm
to
1
rings
abrupt,
scattered clusters,
Leaves clustered
cm.
Flowers [2]
4-dentate, abruptly flattened
lar,
protuberances,
from
arising
27
to
black. Entrance holes
to
thickened rim
Internodes 0.3-3
cm.
holes,
rhomboid, apex acute, base blunt to cordate, margin crinkled. Petiole up
ovate
cm.
61
of Squamellaria
entrance
small
dark brown
long,
cm
of
rings
with
apart; prominent,
cm
to
with
rough,
to
spines
revision
subpendent, regular, cylindrical-fusiform, tapering
swollen between the
ruptly
A
keel.
Ecology
400
Notes
gest
an
low level
epiphyte, growing
species
status.
in S.
species;
clusters
The
two
species, only
trunks of
on
at
stems
and S. imberbis the
Its
stem
trees
in forest
at
300-
apparently
the tuber surface is
stems are
not
as com-
and leaf characters sug-
but flower characters indicate
species,
on
and
once,
sympatric.
between the other
The distribution of
major
collected
with which it is
intermediateposition
specific
peculiar
to
this
scattered, while here they arise in
the apex of the tuber and elsewhere.
specific epithet
derives from the
Fijian
name
'theke theke nkau' for
ant-plants,
'testicles of the trees'.
meaning
Collections.
477
A
A little known
-
the
mon as
its
-
m.
(type),
478
FIJI ISLANDS. Taveuni.
SE
16°
50'
180°
00',
Taveuni
road
to Devoeux
Peak,
Jebb
(cotype).
ACKNOWLEDGEMENTS
I
tion,
thank F. White and C.R.
and
Rosemary
Huxley
Wise for her
for their support,
meticulous
A.C.
Smith for nomenclatural
clarifica-
drawings.
REFERENCES
BECCARI,
O. 1886. Piante
HUXLEY, C.R.,
the
JEBB,
Hydnophytinae.
M.H.P.
Thesis,
SMITH,
SMITH,
1985.
Oxford
A.C.
from
Fiji.
A.C.
Ospitatrici.
Malesia 2
:
228-230.
& M.H.P. JEBB. 1991. The tuberous
Blumea
36:
Taxonomy
1-20
and
epiphytes
of the Rubiaceae
1: A
new
subtribe
-
(this issue).
tuber
morphology
of
the
rubiaceous
ant-plants.
D.Phil.
University.
1967.
Studies
Contr. U.S.
1988. Flora
of Pacific
Natl. Herb.
Island
37:
plants
XVIII. New
and
noteworthy flowering plants
69-106.
Vitiensis Nova 4. Pacific
Tropical
Botanic
Garden,
Hawaii.
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