Download (Asclepiadaceae) (Asclepiadaceae: to 300

BLUMEA
46
457-483
(2001)
Notes on the taxonomy and ecology of the genus
Hoya (Asclepiadaceae)
David
Kleijn
&
in
Ruurd
Central Sulawesi
van
Donkelaar
Summary
The
and
taxonomy
distribution:
mation
the
on
ecology
alistic
a
of
ecology
relationships
single species
with ants
that
of the
Hoya
(Roxb.)
(H. Karst.)
Warb. and
species
described.
Key
are
words'.
and
in
occur
the
species
Merr.
H.
is
unclear, especially
are
Jaya.
This
describes
Hoya species
4) species
described
eight
sussuela
and Irian
paper
genus:
eight species
1) species
with
in
grew
placed
in
pseudomaxima
association
synonymy
Koord.
Hoya, Marsdenieae, Sulawesi,
are
H.
reduced
ants. Four
of its
Sulawesi.
types
An
of mutu-
leaves to house
ants;
leaves; 3) species growing
on
and branches.
(types 2-4). Taxonomically,
ants
coronaria
to
area
available infor-
cavities in tree trunks
with
with
core
from central
specialised
imbricate
that root in ant inhabited
in the
summarises the
is their association with
for ants under
provides housing
the carton of ant nests; and
Four
many
R.Br,
Hoya
genus
Sulawesi
Hoya species,
trait of
important ecological
2)
of the
Sumatra, Borneo,
Blume,
imbricata
H.
and H.
Decne.
maxima
Three
new
gardens, ant plants, epiphytes.
ant
Introduction
The
to
genus
300,
R.Br.
Hoya
(Asclepiadaceae:
predominantly epiphytically
Marsdenieae)
growing species.
ranges from S China in the north, the
western
tralian rain forests in the south and the
rather
Most
at
vines with slender
are
each node. All
variety
species
abounds in
the size and
The taxonomic
description
contributions were made
den Brink f.
(1990,1992,
other authors
split
in
up in
many
by
stems
and
archipelago
of the
genus
Schlechter
and
Hoya
(1913)
is
SE Asia
typically
2)
diversity
can
areas.
fragmentary. Significant
for New Guinea, Backer & Bakhuizen
(1965) for Java, Rintz (1978) for Malaysia
are
of
one or a
few
species.
countries thatused
species being
described
of Environmental
e-mail:
Laantje 1,
1996).
leaves
Other than that,
highest species
The
to
and Forster & Liddle
more
Sciences,
geographical
range of Hoya is
4251
than
once.
Nature
[email protected]
EL
Werkendam,
large
and
be rather isolated scientifically. This resulted
Furthermore, Rintz (1978) noted
Conservation
WageningenUniversity, Bornsesteeg 69,6708 PD Wageningen, The
author,
opposite
two
margins.
best
corresponds
(Richards,
directly bordering
at
distribution
1993, 1995) for Australia and Papua New Guinea. Most contributions of
numerous
1) Department
(India)
of organs. The
shape
geographical
in the west, the northAus-
leaves with entire
petioled
be found in the Indonesian
probably
van
have
twining
estimated 200
an
Islands in the east, which
with the distribution of rain forests in
accurately
species
Fiji
Ghats
consists of
Its
The Netherlands.
and
Plant
Ecology Group,
Netherlands.
Corresponding
BLUMEA
458
that
numerous
that
species
Presently,
probably
which
ditionally,
nomical
than 400
more
the
not
does
the
making
2001
3,
identification of
correct
have been described in
species
synonyms may
be caused
and subdivision of the genus into well-defined sections
Kloppenburg,
1994)
are
still
(e.g.
This confusion may be due
geographical
well
as
range
(e.g.
Information on the
where
they
were
Irian
of all but
haphazard
size and
or
a
few
data
with
in which the
large
areas
the sections
have
species
in the heart of the
factors
absent.
completely
are
our
in the
Hoya
the habitat
to
pollinators, flowering
increasing
at
of
knowledge
core
of their
area
characters relevant
morphological
It summarises the available
some
is limited
for instance,
growth limiting
concise summary of the
a
species.
family
relatively unambiguous.
Hoya species
on,
series of papers aimed
a
Ad-
taxo-
attempts by Burton,
see
are
way
of
of members of the genus
ecology
supplemented
genus
lises
of
large
1997)
species.
Jaya, Borneo, Sulawesi, Sumatra).
ecological
gives
description
the
found. Basic
the taxonomy and
distribution. It
to
under-representation
ecology
This paper is the first of
the
partly
the
as
seed set, pests, diseases,
rhythm,
(Omlor,
unsatisfactory (Omlor, 1997); only
Eriostemma Schltr. and Acanthostemma (Blume) Koord.
been described,
genus
the
by
difficult.
specimens
of the genus. Both delimitation of the genus within the
complexity
1985 and
this
contain more than 300, many yet undiscovered,
number of
large
No.
46,
mentioned in the literature could not be found in the
were
various herbaria he visited,
Vol.
to
of the
ecological knowledge
novel observations made in central Sulawesi and finafrom central Sulawesi.
by describing eight species
MORPHOLOGY
Vegetative growth
Most
grow
species
of Hoya
adventitious
spaced
as
are
roots
arboreal shrubs
vines that twine counter-clockwise and
with which
down from their substrate (H.
leaves
oppositely positioned
that
develops only
species
display
densely
all
hirsute
adhere
cummingiana Decne.).
on
each node, the
adult imbricate leave per node
H.
extremely
and 30
(H. macrophylla Blume), however,
leaf size is
enormously
3-5-nerved (H.
ness
with
more
ex
Trail
two
H. imbricata Decne.
Leaf
shape
of different
linear (H. linearis Wall.)
leaf size of different
within
species
or
usually pinnate (H.
5-7-nerved (H.
individual
lacunosa
is
purpureo-fusca
to
P.I. Forst. &
extremely
plants
Blume),
Hook.),
diversifolia Blume).
(H. multiflora)
subsp. rupicola (Hill)
species
(H. microphylla Schltr.)
cm
succulent leaves (H.
of the leaves varies between herbaceous
australis R.Br,
to
species produce
one
(Fig. 8e).
species
just hang
glabrous (H. camphorifolia Warb.)
even
variable. Leaf venation is
species
All but
somewhere between 0.5
cinnamomifolia Hook.),
may be obscure in
(H.
usually
have branches that
Gamble)
parvifolia Schltr.). Average
variable but
cm
&
King
or
produce randomly
their substrate. Some
exception being
levels of hairiness between
i
to
multiflora Blume)
range from orbicular (H. curtisii
and may
and
one
H.
(e.g.
they
or
Thick-
succulent
Liddle) but,
like leaf
size, this may vary considerably with the environmental conditions a plant experiences.
Flowers
are
arranged
in umbels
nodes (a small number of
shape
across
of the
peduncle
on
species
persisting peduncles
have
peduncles
is variable between
environments making it
a
character
species
by
that
that
drop
are
positioned
off after
but rather
on
flowering).
constant
within
the
The
species
which otherwise similar species may be
when
separated
geotropic,
this is
an
& R.
Kleijn
D.
they
positively geotropic,
sections from the
two
coronal extensions
(see Fig. 1)
number of flowers
number of flowers
flower
Hoya species
rarely
a
while
characters,
vegetative
shape,
reliable. Note however that
some
upon the
and
species
conditions
to
herbarium sheets
based
on
be the
leaf size and thickness
vegetative
most
reliable
generally
are
simply indistinguishable
are
but the
species
growing
numerous
distinguish species
leaf venation and leaf shape
pubescence,
con-
trait for taxonomic purposes.
confusing
lack flowers, many attempts have been made to
characters. We find peduncle
between
considerably
infrequently,
or
also with
Miq.,
coronal lobes). The maximum
outer
single plant depends
a
umbels and bilobed
positively geotropic
concave,
per inflorescence varies
produced by
and
1978) that is particularly helpful in
and section Otostemma (Blume)
(Forster & Liddle, 1991) which makes it
As many
(Rintz,
positively
and
concave
negatively geotropic,
and
convex
umbels but with solid
positively geotropic
cave,
or
459
Hoya
genus
of the genus (section Acanthostemma (Blume)
rest
by
the
ecology of
The umbels may be
flowering.
Koord. which is characterised
and
Taxonomy
taxonomic character
important
separating
not
are
and
convex
Donkelaar:
van
until
they
un-
flower.
Flowers
The flowers of the
Asclepiadaceae
are
the
all the dicots (Endress,
1994). Organisation
been described in detail
by
from
the
an
elaborate
most
(e.g.
of the
Schlechter
to
(1913)
and the
separates Hoya
elaborate, complicated
flower. An
given
in
related
genera
and Micholitzia N.E.Br, from other related
genera within the
shape
of the
fields. However,
trait
(e.g.
all
a
related genera. The
tation within the
wings,
of the
shape
pollinaria
appendages
are
extremely variable,
and width (Forster & Liddle,
be used
in
rigidly
too
which may vary
notably
separating
Absolmsia Kuntze
as
lateral
darwinii
clearly separate
corolla lobe
length
have this
not
Thus
a
this genus from
taxon
delimi-
of this organ is
shape
corolla, anther
corona,
taxonomic
important
is the
germination
Loher).
character for
important
1978). The shape of the
further
length
not
an
H.
the additional benefit that the
size is
most
is
to
flower with
Asclepiadaceae
within the genus do
species
probably required
are
in dried flowers (Rintz,
even
that function
of the section Eriostemma Schltr.,
genus. It has
and anther
wings
considerable number of
species
number of different traits
preserved
have
pollinia. Hoya pollinia
therefore refrain
Hoya
distinct floral character
most
monospecific
a
1.
Fig.
and Omlor (1997) the
closely
we
of
example
flowers of
flowers have
Asclepiad
1990, 1991, 1993), and
Hoya
features is
important morphological
According
that
Kunze
description
most
and function of
characters. Flower
and width and
pollinium
1991). Size of the flower organs may therefore
different
even
between the successive
have
two
taxa.
The
flowers
same
on
the
applies
same
flower colour
to
peduncle.
Fruits and seeds
Hoya species
into
a
common.
Hoya
tightly packed,
Schltr. form
pericarp.
long,
an
On
thin and
follicles
comose
slightly
development
are
generally
of
two
to
the
rule, by
the follicle
splits
flattened seeds,
being
open
apically
usually
one
follicles per flower is
long, slim,
seeds. The follicles of the
exception
maturity,
ovaries. However, after fertilisation
the
follicle, although
have
species
thick
along
a
thin
pericarp
very
and
out
un-
contain
from the section Eriostemma
courgette-like
a
grows
not
single
with the still
with
suture,
damp
a
spongy
revealing
comas
the
(or plumes).
BLUMEA
460
Fig. 1.
the
1.
A flower
typical
Pedicel;
tensions;
18.
As
2.
calyx;
7. anther
area of
ceptive
of the
characters.
3.
corolla;
wing;
8.
style head;
b.
Top view;
a.
side
view;
4. inner coronal
9.
guide rail;
13.
14.
ovary;
46,
No.
3,
2001
Acanthostemma
Hoya (section
genus
-
Vol.
anther
c.
lobe;
sectional
5. outer coronal
appendage;
retinaculum;
(Blume) Koord.) showing
median
15.
10.
view;
lobe; 6.
16. caudicle
most
of
pollinarium.
bilobal
11.
pollinarium;
caudicle;
d.
coronal
style head;
17.
wing;
-
ex-
12.
re-
pollinium;
pollinium wing.
the
comas
caught by
helps
dry, they expand
and
push
the seeds outward and
the wind and float off (Rintz, 1978). The
the seed
the bark of
to
adhere
a tree.
(Rintz, 1978).
to
Seeds
Most
of follicle and seed
any
damp
surface with which it
germinate immediately
species
set
seed
morphology
for
plume
rarely.
taxon
upon
upward
is very
comes
not
they
are
which
into contact, such
encountering
It is therefore
until
hygroscopic
a
as
moist substrate
clear what the value is
delimitationwithin the genus.
ECOLOGY
Distribution in
Most
found
and
species
habitats
grow between 0 and 800
primarily along rivers,
branches of
secondary
found
The
Hoya
different
growing
species
be difficult
on,
usually,
lakes and
trees.
More
sea
m
altitude. In this
shores, usually
rarely, specimens
calcareous but
of
on
notice
more common
as
they
Hoya species
climb into and crawl
also
fruit and ornamental trees, in old
occur
trees
regularly
on
over
nearby
in rural
graveyards
and
areas,
and in
can
be
primary
can
be
of rock
types
outcrops.
of the section Eriostemma Schltr. all grow terrestrial,
to
they
zone,
stem
epiphytic species
other
occasionally
the
at
times
shrubs and
which
trees.
growing
on
plantations.
may
The
roadside,
Of
course,
D.
&
Kleijn
these observations
canopy of
canopies
cut
solitary
The
important
of
range
altitudes. The upper limit is
at
Seed
cracks
on
was
the
do
species
is restricted
Hoya species
1913),
±1500
m:
1700
±
m
m
so
from
tree
branches
because
Hoya species
the entire forest and many
throughout
Malaysia
in Java (Backer
open than lowland forests,
more
higher
to
in
in Sulawesi. Presumably
m
Krakatau Islands
heavily
species
can
covered
et
new
that is,
Once
collected
on
dispersal.
they
by
in
have been
and
two
eruption.
primary
Hoya
In 1979
and all three of the
of many
dispersal
modes of
potential
a
Java and Sumatra
common on
respectively
two
ants
Their
of 1883. The first
eruption
are
The mode of
1995).
plant.
actively
van
al.,
that
established
diplochorous,
as
the establishment of a
volcanic
species
two
firmly
were
may be characterised
runways (Docters
devastating
wind
to
the Krakatau Islands where
on
the islands between 51 and 68 years after the
(Bush
are
illustrated
diversifolia,
(some 30 km distant),
species
suggest adaptation
was
started after the
H. lacunosa and H.
dispersal
Hoya species
are
involved in
dispersed by wind, the seeds of
in
placed
their arboreal
Leeuwen, 1929; Kiew & Anthonysamy,
1987).
nests
and
On Java, Docters
Leeuwen (1928) found that the seedcoat of, amongst others, H. lacunosa, contained
high
concentrations of oils. He
collect seeds
their
nest
with
ants are
Hoya
although
and
Seed
dispersal
of the
of
Hoya
should
plant
them in the
ants to
carton
of
that associate
Hoya species
myrmecochorous.
grow in association with
varies between
et
species
al.,
but four
species
that have
1999). Hoya
pair
are
pressed
to
ants.
to
the host
multichambered structure. This
genera, many
root
were
The
tree
tip
and
margin
surface, forming
found
pair, covering
structure
to
specialisation
material in the chambers.
to
the
housing
adapted
nature
be
is
nesting
next
a
Hoya
cavity
pairs
them partly,
usually
to
house
domatia
pressed
ants
et
ants were
spherical
in
a
sim-
ultimately resulting
(Weissflog
darwinii also has
of ants. The domatia are
ants
between the bark
are
inhabited by
there
of the
recognised.
of the inner-most leaves of
systems develop only inside the domatia when
iting organic
be
can
mitrata Kerr forms, besides normal leaves,
the leaves of each older
of which
particular
leaves
or
and the underside of the vaulted leaf. The leaves of the
ilar fashion
The
general categories
myrmecodomatia,
that consists of about five leaf pairs.
tensive
the main stimulant for
was
numerous
therefore both anemochorous and
Hoya
(Weissflog
the first
that this
why they
ants
relationship
First,
suggested
it remains unclear
and runways.
Many species
a
the
forest had been
well. What
as
between 1200 and 2000
1965) and ± 1200
Hoya
dispersal
species (re-)colonised
in
growing
to
in
rock.
granite
seeds of
distance
succession
van
specimens
surrounding
in this niche
range of substrates,
a
in bare
plumed
long
these
Hoya
in New Guinea (Schlechter,
m
den Brink f.,
growing
any observer has
access
dispersal
The
for
occur
smaller number of
a
usually
altitudes grow scattered
higher
be found
to
van
of
Sightings
of mountainforests is much
stmcture
moss
± 2200
1978),
& Bakhuizen
the poor
461
Hoya
genus
this niche is in their habitat remains unclear.
geographical
(Rintz,
by
the
ecology of
trees, left standing after the
down, suggest that this genus does
and how
the
biased
probably
and
Taxonomy
lowland forests.
tall
m
Donkelaar:
van
are
primary
of 30
R.
of a
al.,
range of
1999).
Ex-
present depos-
dimorphic
leaves
as
a
and made up of several
BLUMEA
462
Vol.
leaves, tightly pressed together forming
with
Second, Hoya species
Decne.
that
fully
in
Hoya species
developing only
placed
over
the
underneaththe leaves, which allows the
(Karsten,
of
species
The
1895).
ants.
Ant
that
more
favourable
that
species
in the
germination
In the Malesian
1978).
described
Ule
In his
by
him
D.
sinuosa
which
are
Table
1
were H.
closely
on
Java
was
spec.
larly. Interestingly,
(1987)
ant
van
ants
cies with
no
Hook.)
ex
one
nov.
have
never
carton
other
ings
area
of
poorly
observed
rarely
Huxley (1978)
a
a
set
were
of
first
that the
Iridomyrmex
of the
equivalent
In Sulawesi
by
to
be
Dischidia
same
phe-
major (Vahl)
Docters
of the
when
found the
we
the substrate of
van
nests
of
ant
gardens
Leeuwen (1928)
(Fig. 2).
gardens
ant
in central Sulawesi
ex
Lindl. while the ferns
major,
a
classical
house ants, such
to
by
ant
as
Kiew &
plant
that
800
construct
m
this
by
ant
pitcher
to
on
the
altitude. We
type
of nests
not
in
of
ant nests
speculate
that this
carton
as
the
conspicuous
Sulawesi
carton
nests
this
species
nor
are
the altitudinal distribution of the
building
leaves
nests, while spe-
the authors. In support of this view
examining
cordatus. In addition
Anthonysamy
with
H. brevialata and Dischidia
Hoya species growing
approximately
ants
Pyrrosia
sinuosa occurred somewhat less regu-
observed above this altitude;
in Indonesia visited
Iridomyrmex
baduvi Forel and
correlated with the location of
correlated.
theabsence of
ant nests are
of
one
gardens'
(1928) suggested
Leeuwen
Smith
Lecanopteris
adaptations
strongly
by
'ant
Such
nests.
rapidly (Kleinfeldt,
usually just
Table 1 confirms earlier observations
been observed above
may be caused
more
1910).
encounter
(see below), Dischidia imbricata, Dischidia nummularia
Morton and
obvious
therefore
ant carton
are
type of
most common
debris (Wheeler,
organic
the Asian
Copel.
primary components
may nest, is
nummularia are
far the
usually accompanied by
that the distributionof Dischidia
in which
imbricata is
Thus far, H.
may grow
species
R.Br, and the orchid Dendrobium bicaudatum Reinw.
longifolia (Burm.f.)
material that is
organic
of materials ranging from plant
nutritive
were
for the ants, while
nectar
ants.
ex
by
Crematogaster
cordatus Fr.
related to the
shows that the
brevialata
Scort.
longipeduncu-
punctata (Blume) Decne., and the ferns Lecanopteris curtisii
(Wall,
species Iridomyrmex
ant
observed
astephana
variety
soil and
arboreal
on
species
H. lacunosa
case
Merr., D. imbricata,
Baker and L.
grow
ant
observed
Mayr species
nomenon.
commonly
specialisation.
plants
in Brazil. Doctors
(1901)
of the
nests
conditions and
the
of several
one
This is
a
potentially
archipelago, Hoya
systematically
by
overgrown
on ant nests.
grow
decaying leaves, sand,
epiphytes planted
by
with this kind of
constructed from
nests are
fibres and thin bark to
Seeds of
D.
substrates
exposed
usually occupied by
more
only develop
roots
very
nutrients obtained from the
extra
Hoya species
Third, Hoya species
relationship.
grow
albiflora Griff.,
underneath the leaves
deposited
known
only
on
shelter and food in the form of floral
benefits from the
collected and
and
(Fig. 8e)
to
is
species
leaf per internode.
cochleata Blume, D. imbricata (Blume) Stcud.. D.
Ridl.), provides
plant
plants
D.
(e.g.
imbricata
Hoya
branches. This
or
one mature
this type of mutualism, which is
Supposedly,
& Gamble, D.
King
stem
space below each leaf is
in the related genus Dischidia R.Br.
the
(Kloppen-
below] has large imbricate leaves with
see
the bark of stems
tightly pressed against
are
amongst all
unique
convex-orbicular leaves.
specialised
A succession of such leaves is
the
with 4-6 cavities
globose pouch
H. maxima (H. Karst.) Warb.,
[syn.
margins
lata
2001
3,
1993).
burg,
the
a
No.
46,
in any
the find-
ant
species
inhabits the
D.
Fig.
2. A
&
Kleijn
R.
well-developed
Iridomyrmex
cordatus
van
ant
Donkelaar:
in
garden
overgrown
with
a
central
Sulawesi
number of
cavities in
most
to
Jack and
tane
forests where it
nophytum
for
was
and
ants
H.
Bada
ant
are
were
& Walker
at
commonly
pubera Blume,
at
2400
m
to
especially
Hydnophytum
be absent in upper
species, suggesting
for
instance
in New Guinea
on ant nests are:
picta Miq.,
However, the ecological characteristics of
expect that many other species,
of the ants
while she found
of the genera
1950 m in central Sulawesi
found
H.
A nest
Valley).
463
Hoya
garden epiphytes, epiphytes
high altitudes,
found
genus
1974). However,
plants
another Iridomyrmex
at
the
Pyrrosia piloselloides.
Janzen,
contrast to ant
common
Myrmecodia
parvifolia,
and
occupant of
replaced by
are
1975). Hoya species that
H.
(Gintu,
Jack in lowland forests, she found it
Lecanopteris spinosa Jermy
lata,
plants (e.g.
lower altitudes. In
to
housing
ant
most common
Myrmecodia
is restricted
vide
SE Asian
be the
ecology of
typical epiphytes: Asplenium nidus, Dendrobium
bicaudatum, Hoya brevialata, Lecanopteris sinuosa
I. cordatus
and
Taxonomy
most
H.
species
(Huxley,
(Jermy
mon-
that it
that proof
Hyd-
1978)
or
& Walker,
H. lacunosa, H. brevia-
spec. IPPS 4562 from Biak.
Hoya species
are
unknown and
we
from the section Acanthostemma (Blume)
Koord., demonstrate this type of relationship with
ants.
BLUMEA
464
Table
Plant
1.
(no. 1-11)
*
substrate
=
found
species
and
the
on
arboreal
Gimpu (Bada Valley,
bamboo;
**
=
Vol.
nests of
12-21)
no.
substrate bare
in
T
rock;
No.
46,
2001
3,
cordatus
Iridomyrmex
around Tentena
ants
central Sulawesi.
Total.
=
*
11
Antnestno.:
Ant nest no.:
2
2
3
3
5
5
4
4
6
6
7
7
8
8
9
9
10
10
12
12
11
13
13
14*15
14*15
16M7
16*17
18
18
19
19
20
20
T
T
21**
21*
species
species
Hoya
brevialata
brevialala
18
18
Dischidia nummularia
Dischidia
•
••
•
16
••
Dendrobium bicaudatum
•
••
.
.
Dischidia imbricata
Dischidia
imbricata
•
•••••
Pyrrosia longifolia
Pyrrosia
•
•
•
•
••
•
••
.
.
Myrmecodia
•
*
•
•
8
•
••••
...
•
7
4
•
spec.
spec.
3
.
rides L.
Asplenium
Asplenium tiides
1
•
Dischidia major
major
Dischidia
•
•
.
|3
13
]3
13
•••
.
.
•
.
•
•
•
•
.
•
•
Lecanopteris sinuosa
Lecanopteris
Pyrrosia piloselloides
••••••
I1
•
I1
•
spec.
Drynaria
Drynaria sparsisora
•
11
■
1
Moore
(Desv.)
(Desv.) T. Moore
Phymatosorus
scolopendrius
Phymalosorus scohpendrius
Pic.Serm.
(Burm.f.)
(Burm.f.) Pic.Serm.
Fourth, species
little holes in
Closer
1
(Fig.
a
ant
similar
a
epiphytes
ants
were
that tmnks and branches of the
of the
by
the
where
ants.
Little holes
roots
of both
of
stems
due to its
root
to
cm
under
by
this
ants
scavengers
and H.
Kuntze
root
plants
the
(Fig.
in
a
to
number
similar fashion from
1 le). The hole
of the genus
ants
parvifolia
ants to
was
closed
Ridl. in
Crematogaster
Lund
Malaysia. They
found
Sm. contained intera
single colony
reach the surface of the branches and
penetrate the central cavities in the branches
growing
was
Of both
supporting
ant-tree
pallilimba are
this type of relationship
to
out
centre
of trunks that
may be
quite
the
from Sulawesi (almost) the entire
the cavities,
nests
general
known
suggesting
that collectionof the
is the basic mechanism that
Hoya species
only
ants.
similar fashion from holes in the
ant-epiphyte relationship
were
also observed
view that the
relationship (Weir
the
with
a
Hoya species
confined
both
in
but may often have been overlooked in the past
and transport into their
on
growing
system within the rotted
epiphytic Asclepiads
relationship. However,
mecopa
the
a
the time of observation. Weir & Kiew
in diameter. Thus, this type of
ant-inhabited structures,
the
by
in
species.
of the genus Tetramorium Mayr.
Leptospermum flavescens
ofDischidia
inconspicuousness.
the
and D.
growing
was
from
growing
habit of parasitic
growth
substantial volume ofthe available space. Green (1993) observed
with their main
system of the
seeds
a
tree
at
found
was
of the presence of the ants) that housed
used
spartioides (Benth.)
trees
7.5-10
common
were
species
they occupied
Absolmsia
were
(irrespective
ants
root
spec,
between
astephana
the
trunks and branches. In
tree
below),
found
was
absent
relationship
Dischidia
connected cavities
site of
hollow Pandanus
carton, however,
(1986) reported
and the
a
(see
system of the plant
root
as a nest
(see below)
nov.
spec.
little circular hole in
with
nov.
Oe), resembling
learned that the
inspection
of connected cavities that served
Hoya pallilimba
inhabited cavities in
ant
myrmecopa spec.
branches
tree
in
growing
central Sulawesi, H.
& Kiew,
species
epiphytes
1986).
within the
gives
growing
are
rise
outside
essentially
Thus far, H. myr-
genus that demonstrate
D.
& R.
Kleijn
Donkelaar:
van
and
Taxonomy
the
ecology of
465
Hoya
genus
Pollination
The
of all but
pollinators
that H. australis R.Br,
sothis Waterhouse
most
dusk however,
at
open
Hesperiidae),
dominant flower colours
are
insects
role in the
play
an
important
did not exclude the
observed
at
fragrance
night
& Matile
Altenburger
is emitted in
fall in
dark
a
circadian
a
Hoya species
of Hoya
species, feeding
the
ants
whether
to
copious
Ants
the
on
nectar
play
a
be
can
species
organisms (Beattie,
considered
are
stages of
barrier
to
pollen
in the
tible
ant
to
It is
1985).
with antibiotics that
genus
to
pollinated
notes
R.Br, that
may be
a
of their
species,
as
was
protected by
the flowers
strong incentive for
contrast to
of
nests
to
It is
doubtful
originally suggested
wax
attacks
juvenile
or
from
This
may have
1985).
presented
that
species
a
On the other hand,
which may make them less
of the
paper
micro-
themselves and their broods
ants cover
pollen growth.
some
relationship
on
nests.
be poor pollinators. In
are
emission would
the oils in the cuticular cells of
to
carton
pollinaria,
that
next
role in the
suscep-
reported
were
that, like Hoya species, deposit pollen
orchids,
were
that the
so
important
pollination systems (Beattie,
(1985)
of
scent
pollinators.
carnosa
scurrying incessantly
seen
that
inhibits
for the
attractants
relatively exposed
suggested
is released in
Hoya
Beattie
lie
ants
incidentally
the evolution of ant
damage.
an
of these
stages of the related bees and wasps, that
respectively, juvenile
period,
play
in the
pollination
for the
and the strong
determined for H.
In addition
nectar.
(1992)
partly responsible
nectar
produced by Hoya species
role in the
Schlechter (1916). Ants
by
ants.
seeds of these
plant
ants
and
of Forster
study
that is, flowers timed their emission of fra-
Nectar may also
period.
between
the seeds, the
rhythm,
were
respectively
(1988) experimentally
12 hours after the last dark
to occur c.
grance
richly produced
the main incentives
are
The
process.
insects
night flying
that nocturnal
pale yellow, suggests
or
pollination
The
pollination efficiency.
the flowers
again
that
The flowers of
day flying butterfly.
a
which, combined with the fact that the
white, greenish
possibility
walkeri
effectively pollinated by Ocybadistes
was
(Lepidoptera:
species
Hoya
remain unknown. Forster (1992) found
Hoya species
one
Traill
ex
to
be
pollinia.
as
Conservation
The main threat to
they
are even more
show
species. Many species
moss-covered branches
ants.
It may take
a
of the genus
species
susceptible
or
long
a
is habitat destruction.
Hoya
logging
to
and
clear-cutting
particular preference
time before
are
extinction threat and
many
to
easy
grow in situ; however,
orchids which
we
have
seen
only
once a
trees
or
even
other
urban
they
man on
placed
Such
their habitat for
a
appeal
on
at
a
minimum size,
branches
developed
all. Such
to
garden
the local
trees
by
a
Most
local
was
major
face
a
species
Unlike
population.
in E Java) that
species depends
occupied by
after
species
most
for
people,
used
as
an
therefore on the conservation
grow.
species
areas.
tree
disappeared already.
Hoya species (H. diversifolia
are
more
robust and may be encountered
For instance,
that adorn the main
squares
Malang (E Java).
may have
have low
Conservation of these
of the forests in which
Fortunately,
they
collected and
being
are
ornamentalplant.
rural and
species
proceed
to
of
trees
suitable substrate has
a
disturbance, if natural regeneration is allowed
high
for
co-inhabitantsof hollows in
are
Being epiphytes,
than terrestrial rain forest
('alun-alun')
species
long
H.
will
time.
diversifolia
grows
of large cities such
probably
survive
most
on
as
the
regularly
large
Medan
alterations
in
Ficus
(Sumatra)
imposed by
BLUMEA
466
Fig.
c.
in
3.
Hoya
flower
the
in
brevialata
median
Netherlands;
Kleijn
section;
IPPS
d.
& Van
Donkelaar.
pollinaria;
7718).
Vol.
e.
46,
a.
No.
3,
Flower
2001
in
growth habit (drawn
top view;
b.
flower in
from live material
side
grown
view;
indoors
D.
Kleijn
& R.
van
Donkelaar:
HOYA SPECIES
of both
as not
are
1.
and
vegetative
all
species
were
seen
brevialata
Caules tenues
characters and
flowering.
in Sulawesi will be between 15 and
species
This number is based
The eight
partly
partly
that
examination
on
vegetative
on
species
& Van Donkelaar, spec.
Kleijn
glabrescentes
viridia vel flavida ad clare
longi.
Umbellulae
albi ad
lobis
caudiculis
characters
observed
were
only
flowering
Umbels
than corolla,
corolla
to
bright
floribus
manifeste
proprie
Sulawesi
breves
alatis.
elliptica pallide
Pedunculi
longi.
cm
15-25. Florum
sursum
curvatis,
gynostegii
Tentena,
exposed.
positively geotropic,
deep red,
to
strongly recurved, glabrous
down
on
cm
fere
a
rubriores
corolla valde
re-
attingentes
&
Donkelaar
in
Van
plantation.
clove
Petioles 0.5-1
to
elliptic, light
long.
cm
Peduncles
with 15-25 flowers. Flowers: colour
corona
usually
ridged,
one to
several shades darker
bilobal extensions
both sides. Pollinaria anther
clearly upcurved,
characteris-
wings
the
gynostegium (Fig. 3b, c),
only
recorded from Sulawesi.
only halfway
3-6
colores
ad dimidio
—Typus: Kleijn
Province:
side of the coronal lobes
tically short, coming
with
C
suborbicularia ad
plerumque corolla aliquantum gradus
red when very
concave,
upper
alae
3
Fig.
—
with short internodes. Leaves suborbicular
glabrescent,
from almost white
ranging
corona
nov.
Petioli 0.5-1
angustis pyriformibus
(holo L), Indonesia,
green, yellowish
cm.
variantes,
cristatis extensionibus bilobatis
(Fig. 3b, c),
Stems thin,
brevibus. Folia
exposita.
positive geotropicae
utrinque glabra. Antherae
IPPS 8836
3-6
internodiis
rubra ubi
concavae
rubri
profunde
supra
curvata
caudicles
narrow
pear-shaped wings.
Distribution
Ecology
—
—
brevialata is
Hoya
From 0
to
600
in central Sulawesi
species
m
altitude, H. brevialata is by far the
due
probably
rural and semi-natural habitats.
Hoya
ant nests
but
subsequently colonising
attached with adventitious
trees.
cially along
shapes
as
Note
being
Table 1),
shores of rivers,
most
especially
conspicuous
abundant in
the entire
apparently starting growth
tree.
in
trees
lakes and
gardens,
using
seas
a
tree
but also
variety
simply
literally draped through
or
in clove
conspicuous
ants
from the
It also grows without ants,
the bark of tree trunks,
In natural habitats it is less
of
plantations
quite
trees
and in
common,
espe-
of all sizes and
substrate.
—
The
name
'brevis' and
H.
Hoya camphorifolia Warb.,
Type: Warburg
oblong
s.n.
to
brevialata refers
'wing' being
Hoya camphorifolia
Leaves
to
of ornamental and fruit
crown
roadside
roots
the fact that it is
to
brevialata often grows in association with
(usually Iridomyrmex cordatus, Fig. 2,
2.
467
Hoya
genus
described in detail below.
Hoya
the
a
species.
generative
the
ecology of
IN CENTRAL SULAWESI
We estimate that the total number of Hoy
20. We have observed 13 distinct
and
Taxonomy
(B,
Warb.
in
now
to
the
typically
short anther
wings,
'short'
'ala' in Latin.
—
Schltr.
Fig.
4
& Warb. in
Perkins, Fragm.
Fl.
Philipp.
1
(1904)
129.
—
lost), Philippines, Luzon, Tayabas Province, Sampaloc.
elliptic, shortly accuminate, glabrous
whitish main veins. Petioles
glabrous,
characteristically long (5-11
1-2
cm
long,
with
usually
three
thicker than the
cm), very thin. Umbels
convex,
conspicuous
stems.
negatively
Peduncles
geotropic,
468
Fig.
BLUMEA
4.
Hoya camphorifolia
section;
IPPS
d.
pollinaria;
8845).
e.
Warb.
growth
a.
Vol.
Flower in
habit
(drawn
46,
No.
top view;
from live
3,
2001
b. flower in
material
side
grown
view;
c.
flower in
indoors in the
median
Netherlands;
D.
with
20-40 flowers.
corolla, open for
sides with broad
corolla
ly
& R.
Kleijn
van Donkelaar:
Flowers whitish
tapering
469
Hoya
genus
darker than the
corona
lower
concave,
than inner coronal lobe
tips higher
tips;
lobes recurved. Pollinaria retinaculum short and thick, broad-
faintly pubescent,
flat top
the
purplish,
to
coronal lobe
outer
grooves,
pink
the
ecology of
coronal lobes lanceolate, upper sides
day only;
one
and
Taxonomy
off about
exponentially
to
narrow
bottom
tip;
caudicles
narrowly
winged.
Distribution
and has
the
Ecology
undergrowth
shrubs.
or
of H.
Even
seashore
growing
is
forests in
common
in bushes
No association
ing plantations.
tion in the
was
that this
Philippines
Note —The Sulawesi
have
or
less flat corolla lobes. Otherwise the
3.
Hoya
coronaria Blume
coronaria
Hoya
Blume, Bijdr.
designated here),
Blume
—
16
[Coronaria ariadnes punicea Rumph.,
Roxb.,
Fl.
Ind. ed.
Herb. Amboin.
in
(possibly
velutina
(1917)
31.
—
—
Types:
ariadna
Hoya
Blume, Rumphia
438.
anthropogenic
more
in shrubs border-
growing
Elmer's
(1938) observa-
coronal lobes than the
Philippine counterparts
898.168-128),
no.
Herb. Amboin. 5
—
the
on
behind the
mangrove
areas
Philippine
have
more
similar.
are
4
—
in
172.]
t.
A.DC.,
Hoya
898.168-121,
no.
sussuela
Asclepias
—
Prodr.
sussuela
Herb. Amboin. 5
Syntypes: Rumph.,
(lecto L,
s.n.
(1750) 464,
Decne.
31.
(1849)
Blume
Java.
8
635.
(1844)
—
(Roxb.) Merr., Interpr.
(1750) t. 172, Roxburgh
s.n.
BM).
Wight,
Hoya grandiflora
no.
(1832)
corona-ariadnes
Hoya
Hoya
2
1063.
(1827)
green and
very abundant
just
despite
from both
plants
woody
6
Fig. 5,
(iso L,
s.n.
or
dark
an ant nest.
narrower
and have recurved corolla lobes while their
plants
become
succulent with age.
found in
regularly
ants
stems
highly
flood plains
grew from
species
stems
through
are
altitude. It is
m
observed with
specimens
of
leaves
Young
between sawahs
trees
masses
in the genus, old and young
or on
It is also
palms.
dangling
as
species
age.
from 0-600
environments, hanging from large
common
often these scramble
green and become
rock outcrop
on
Philippines
probably
equally
distinct appearance. The
a
pale
turn
and Pandanus
trees
have
in thickness with
coriaceous, but they usually
Hoya camphorifolia
than other
more
camphorifolia
considerably
and increase
growing
seen
branches of trees, but
primary
is
species
and Sulawesi.
may be
Floy a camphorifolia
—
plant parts
described from the northern
originally
was
Philippine archipelago
from the
hanging
species
before been recorded for Sulawesi. The
not
throughout
This
—
Contr. Bot. Ind.
Blume
ex
Decne.
in
35.
(1834)
A.DC.,
—
Type:
Prodr.
8
Contr. Bot. Ind.
(1844)
635.
—
(1834)
8150.
t.
Blume
Type:
s.n.
(lecto L,
898.168-123, designated here).
Hoya speciosa
Decne.
in
A.DC.,
Prodr. 8
(1844)
634.
—
Type:
Labillardiere
s.n.
(holo
P
n.v.),
Amboina.
Hoya
coronaria
F.M.
(=
Blume
Mt
(B,
now
Stems thick,
from
a
Feddes
lost), Papua
densely
broadly
midriband less
BRI
Bailey, Queensland Agric.
n.v.), Papua
large
Repert. Spec.
New
Guinea,
hirsute. Leaves
rounded
or
pronounced
short, thick, pubescent.
very
F.M.
papuana
New
Guinea,
N
J. 3
(1898)
156.
foot of Mt
Prov.,
—
Type:
Trafalgar
Iamiwara).
Hoya hollrungii Warb.,
661
var.
Bailey AQ360787 (holo
slightly
side
Umbels
flowers. Flowers:
W
Nov.
Regni Veg.
oblong,
(1907)
342.
base obtuse-rounded,
narrowed
nerves.
convex,
3
—
Type: Hollrung
Sepik (= Sundaun) Province, Augusta Station,
Petioles short,
hairy,
1-2
negatively geotropic,
coronal lobes
shortly
top, thickly coriaceous,
cm
with
a
1887.
accuminate
with
long.
a
strong
Peduncles
small number of
yellowish white, usually positioned
in
a
BLUMEA
470
Fig.
5.
Hoya
section;
Lake
d.
coronaria Blume.
pollinaria;
Poso;
horizontal
IPPS
e.
growth
a.
Flower in
habit
No.
46,
top view;
(drawn
from
a
b.
3,
2001
flower
in
side
plant growing
on
view;
c.
flower in
the north-eastern
median
shore
of
8891).
plane although
in
specimens
some
elevated, inner coronal lobes acute,
from
Vol.
slightly campanulate, patent
outer
to
two
segments of
transparent membrane attached laterally
pollinia wingless.
outer
coronal lobe
tips
are
somewhat
varying
stellar, usually yellowish white or reddish yellow
often violet dotted. Pollinaria retinaculum
side, caudicles made of
the
coronal lobes obtuse-rounded; corolla
pear-shaped
a
over
dark
with
tip
on
the
upper
broad
secreted material connected
the entire
length
by
a
of the dark segments,
D.
&
Kleijn
Distribution
Hoy
—
northern Australia,
Peninsular
R.
—
Hoya
Donkelaar:
throughout
It is unclear whether it also
coronaria grows
vegetation
Notes
in
—
in roadsides and
agricultural
wide
range and
in the
on
the
that
central Sulawesi
(in
the
drawn
plates
some
in
reddish
only
by
Rintz
trees
and
extremely
of disturbed natural
remnants
(e.g. Rintz,
1978;
from the
compared
Forster &
section
yellow,
(1978;
shape
only
of our
H. coronaria,
one,
were
com-
very
with the types of
of corona and corolla
stellar flowers have been
comparison
Liddle, 1992)
Eriostemma
specimen
our
differences could be found in
resembled both types. A
specimen
our
It is
altitude, especially along the shores
m
species
two
of this section, and therefore
species. Although
into
high
ground.
described for Sulawesi, H. coronaria and H. sussuela. We found
species
be found in
Philippines.
It may climb
vegetation
occasionally
Examination of the literature
mon
can
areas.
and herbarium sheets showed
both
471
Hoya
genus
the Indonesian archipelago and in
occurs
terrestrially.
in central Sulawesi between 0 and 800
of rivers and lakes,
ecology of the
extremely
an
New Guinea,
shrubs but may also grow amongst herbaceous
common
and
Taxonomy
coronaria has
a
Papua
Malaysia.
Ecology
van
observed),
overall
plate (Fig. 5, Sulawesi)
with
Forster & Liddle (1992; H.
Malaysia),
sussuela, Australia), those in table 182 and 184 by Blume (Blume, Rumphia4,
1849;
H. sussuela,
7023)
showed that,
and
shape:
Especially
the
shape
Fig.
almost
6. A
b. H.
comparison
different
of the
pollinaria
from
of the
Biak,
shape
geographical
from the
Irian
H.
of the
locations.
Province,
Philippines, top:
Jaya,
upper
is
very
constant
ciliata Elmer
side.
—
pollinaria
Hoya
Sumatra
lower
All
ex
of four
bottom:
1
specimens
Irian
specimens
coronaria
bars
in corolla colour
from
mm.
upper
significant.
not
over
specimen,
Burton from the
(pollinia lost), top:
side,
scale
in
of the Sulawesi
from Biak,
species
a.
(particularly
overall differences were
comparison, pollinaria
specimen,
coronaria from Aceh
H. ciliata
material from Sumatra H. coronaria, IPPS
(
campanulate),
unidentified Eriostemma Schltr.
from
c.
to
range. For
geographical
pickled
the flowers varied somewhat
although
from stellar
mentioned Sumatra
an
and
Moluccas),
Jaya
the whole
the above
Philippines,
are
given
in
Fig.
of section Eriostemma
central
lower
side;
Sulawesi,
side,
d. unidentified
6.
Schltr.
bottom
bottom:
and
upper
side;
side;
Hoya species
472
Fig.
BLUMEA
7.
Hoya dolichosparte
section;
d.
pollinaria;
IPPS 8831).
e.
Vol.
Schltr. a. Flower in
growth
habit
(drawn
46,
No.
top view;
3,
2001
b. flower in
from live material
side
grown
view;
c.
flower in
indoors in the
median
Netherlands;
D.
In
&
Kleijn
R.
Donkelaar:
van
the retinacula of the
particular
and the Sumatra specimens
the Sulawesi and Sumatra
as
specimens
H. coronaria and H. sussuela
from
as
are
one
nearly
genus
473
Hoya
differ from the Sulawesi
species
another.
contrast, the retinacula of
By
identical. We therefore conclude that
synonyms and
are
the
ecology of
and Biak
Philippine
well
and
Taxonomy
H. sussuela in synonymy with
place
H. coronaria.
4.
Schltr.
Hoya dolichosparte
Hoya dolichosparte Schltr.,
C
B), Indonesia,
Stems thin,
with
lignify quickly
Toii-Toli.
with age,
pocked
cm
Schlechter 20642
—Type:
with short adventitious
3-5 main veins connected
Peduncles
long.
negatively geotropic,
13.
34,2 (1916)
Province:
light venation,
Petioles thick, 1-2
vex,
Beih. Bot. Centralbl.
Sulawesi
conspicuous
7
Fig.
—
short
characteristically
15-40 flowers. Flowers:
by
roots. Leaves
numerous
2
(<
cm).
ovate
side veins.
Umbels
coronal lobes white,
outer
(holo
con-
acute-
acuminate; inner coronal lobes bright red, acute-acuminate; overall lobe shape elliptic;
coronal lobe
outer
tips slightly
dotted. Pollinariaretinaculum with
ties where the
wingless
Distribution
to
Ecology
sea
—
level
lemon
corolla
tips;
are
faintly
violet-
yellow, usually
laterally protruding margins surrounding
Note
the cavi-
attached.
is endemic
dolichosparte
cle
often found
sets
the
the
along
It
was
a
rather
common
altitude. It
m
sea
was
twining
shore, in large
also found growing
vine in central Sulawesi
observed
trees
growing epiphytically
rivers and lakes and
along
terrestrially
on
limestone
outcrop
the Poso river.
Warb. Furthermore,
are
trees
Vegetatively,
—
is
600
approximately
and
along plantations.
overlooking
5.
inner lobe
pale
available data suggest that H.
Currently
Hoya dolichosparte
to
undergrowth
in shrubs
H.
caudicles
a
Sulawesi.
from
in
—
elevated than
more
from pure white to
pubescent, recurved, ranging
H.
growth
habit and habitat preference
growing through
another. In this
one
species clearly apart.
two
bears close semblance with H.
dolichosparte
The
case
is
species
camphorifolia
similar, and the
are
the
only
shape
undoubtedly
a
two
of the
species
pedun-
close relative of
nicholsoniae F. Muell.
Hoya
Hoya
imbricata Decne.
imbricata Decne.
forma
s.n.
typica,
Koord.,
A.
maximum H.
Meded.
Repert. Spec.
but
in
inval.;
8
DC., Prodr.
8
DC. in
(P n.v.), Philippines, Luzon,
Conchophyllum
Hoya
nom.
Fig.
—
obviously
imbricata
Syntypes:
parently
Lands Plantentuin
Nov.
3
Regni Veg.
Karsten's
Decne.
own
from
from
Stems thin,
single
PNH,
leaf
(1898)
(Edaho), Samar,
PNH,
now
per
Sel. PI. 5
(1846) 37,
Manila; Deless.,
Buitenzorg
534.
non
—
Hoya
Teijsm.
12
90.
t.
Icon. Sel.
158.
(1895)
maxima
& Binn.
15
—
(1919) 263,
PI. 5
—
(1846)
Dischidia
t.
—
Type:
90.
maxima
(H. Karst.) Warb.,
(1863).
incl.
Syntypes: Callery
Feddes
not indicated
collection(s).
now
BS
Koord., Philipp.
18893
J. Sci.
15
(1919) 264,
(McGregor), Biliran, Copeland
t.
399
2,
3.
(all
—
ap-
lost), Mindanao, Davao district,
J. Sci.
15
(1919)
lost), Philippines, Luzon,
sparsely twining,
mature
19
(1907) 342,
Hoya pseudomaxima Koord., Philipp.
loaned
=
Jard. Bot.
forma basi-subcordata
BS 24910
loaned
'Calawan'
Karst., Ann.
J. Sci.
(1844) 637; Koord., Philipp.
Deless., Icon.
grow
out
internode. Leaves
to
265.
—
Rizal
TVpe:
BS 22089
Province,
considerable
(Ramos) (apparently
Bosoboso.
length
large, orbicular, peltate,
before
developing
with various
a
degrees
474
Fig.
BLUMEA
8.
Hoya
section;
d.
Netherlands;
plantation
imbricata
pollinaria;
c.
Decne.
e.
IPPS 8838.
30 km
Flower
a.
growth
habit
Growth
west of
habit
Vol.
in
top view;
(flowers
drawn
Kolonodale).
No.
46,
b.
drawn
from
a
3,
2001
flower
from
in
live
side
view;
material
plant growing on
a
c.
flower
grown
shade
in
median
indoors
tree
in
a
in the
cacao
D.
of incision of
darker spots
white
and
Taxonomy
petiole;
both
covering
the
ecology of
and
green
white with
yellowish
tically long forming
light
or
outer
a
shade of red
green and dark green; lower side
medium-long
of the
top
to
long (>
anther
corona,
characteris-
appendages
short
wings typically
8b, c), corolla inside densely hirsute, strongly recurved. Pollinariaretinaculum
caudicles with
ly elongated,
and attached almost
large wings
5 cm).
yellowish
the inner coronal lobes, inner coronal
on
coronal lobes horizontal, anther
a crown on
of
complex pattern
a
with 10-20 flowers. Flowers white,
positively geotropic,
lobes almost vertical,
red
purplish
Petioles (sub)peltate. Peduncles
purple.
475
Hoya
colour combinations greyish green
lighter background,
on a
genus
and adventitious roots,
stem
tightly against substrate; upper surface with
concave,
or
towards
purple,
dark
Donkelaar:
van
superimposed
with brownish
Umbels
R.
margin
margins pressed
uniformly
&
Kleijn
at
(Fig.
narrow-
the bottom end of the
retinaculum.
Distribution
imbricata
Hoya
—
Sulawesi from 0
it
in the
occurs
600
to
rest
m
from N Sulawesi
originally reported
was
(Karsten, 1895; Koorders, 1920). Nevertheless,
this
is
species
quite
altitude and further investigations
of Sulawesi
as
well. The
also
species
will
common
show that
probably
the
throughout
occurs
only
in central
Philip-
pine archipelago.
Ecology
found it
our
to
(1920) description
Just like Koorders'
in open rain forests and
occur
on
colour and
blend with the
manner
of
the shade
are
irrespective
of the diameterof the
factor
be the
may
run-off: the
as
species rarely
roots
stem
are
activity
species
during
the
produced
housing
ants.
early
to
belong
separation
to
a
plant growth
species
of
the
ants
scent
grows
on
is
species
stem
a
to-
of trees,
important
root
will pass. The
enhanced
at
to use
dusk, but
most
by
this
the leaves
are
scentless
pollinators.
Conchophyllum
maximum H. Karst.,
two
Dischidia flowers
imbricate leaves he collected. In the late
appeared
by
on
this
Karsten
form from the
previously
truly
(1895)
of the leaves and stems, these traits
and
the
water
probably
sweetly
very
number of papers
physiology
of two
on
branches. An
more
it
growing upwards,
has been observed
nocturnal
described
large
the
as
causes
material underneath the leaves. As
because the first paper in this series
and
spot
the space between the imbricate leaves
use
organic
(1895) originally
20th century
morphology
Decne.
These
and collect
one
stems
maxima (H. Karst.) Warb., after characteristics of
species. Probably
in the
ants.
to
its substrate)
and nutrients that passes the rather limited
for shelter. The flowers
Karsten
usually
we
of abandoned coffee (in
and downwards with leaf tips
downstream the substrate, the
More than
mistakenly thought
the
water
on
and, less regularly,
day (Koorders, 1920) indicating
—
Hoya
19th and
more
stem
also located under the leaves,
of the
Notes
of
grows without
for
of this
later
amount
both
pointing skywards,
wards the earth (Koorders, 1920). Hoya imbricata
and the
in Minahassa,
species
trees
growth (tightly pressed against
trunk. Leaves
tree
in leaves with their tips
resulting
system
of this
cacao) plantations (Fig. 8e). In this region it is usually difficult
case
pattern,
to
—
remarkable
focused
played
a
key
on
role
described H. imbricata
Hoya maxima (H. Karst.) Warb., H. pseudomaxima Koord. and H. imbricata
Decne. forma basi-subcordata Koord.
upon the
following
of the leaves; 2) presence of
side of the leaves;
were
distinguished
characters: 1) presence of
3)
the
humps
depth
on
stomata on
from H. imbricata based
the upper and/or lower side
the cuticular cells and/or hairs
of the incision
at
on
the base of the leaves
the upper
(Koorders,
1919,1920). Although there is the obvious (minor) variation in flower size and other
476
Fig.
BLUMEA —Vol.
9.
Hoya
section;
IPPS
d.
minahassae
pollinaria;
8816).
e.
Schltr.
growth
a.
Flower
habit
in
(drawn
46,
top view;
from live
No.
3,
2001
b. flower
material
in
side
grown
view;
c.
indoors in
flower in
the
median
Netherlands;
D.
&
Kleijn
R.
floral characters, flower
We do
constant.
the
H.
477
morphology
of plants with different types of leaves is
relatively
and
Taxonomy
ecology of
forms into separate
growth
Warb., H.
(H. Karst.)
forma basi-subcordata Koord.
that H. shallertiae Burton
synonymy with H.
described from
was
Koord.
pseudomaxima
in
in synonymy with H.
placed
and
and main branches of
minahassae Schltr.
Hoya
minahassae
Hoya
imbricata Decne. We suspect
Stems thick,
with
corolla
narrow
thick.
Umbels
upwards
Distribution
Ecology
elliptic,
almost
—
others; the wings
to
Hoya
—
m.
just
furthermore found
was
It
was
at
found
1100 m in
Type:
Schlechter
Peduncles
long.
20434
(holo
shape
characteristically
with 1-6 flowers.
ovate, inner coronal
coronal lobes acute, upper side
on
is
structure
furthermore
Pollinariaretinaculum
particularly
obvious in this
uncommon
in
covering
to
Sulawesi
the side of the Tineba mountain range (central
more
humid than the side
trees
the Bada
facing
Valley.
in the open mountain forests. It
mountainrange north of Bunku. Schlechter's
a
0.5-1.5
cm
Kleijn
longi.
& Van Donkelaar, spec.
Folia ovata ad
obscura. Pedunculi
corolla
sparse
exteriores
hirsuta
necessariis, apicibus
floribus
sub
extensionibus
corollam
& Van Donkelaar
on
IPPS 8840
limestone
minutis
anthesi
bilobatis
valde
loborum exteriorum multo
of Tentena
elliptica
tenues plerumque
convexae
the
other than that it is confined to mountain
species
from Tomohon, Minahassa (hence the name)
myrmecopa
apices
—
sides.]
covered branches of
negative geotropicae
lobi
waymaniae
growing
at an
2.5-7.5
sursum
6-16.
lobi
laterales
superantibus;
cm
altitude of
1-2
acuta ad
cm
corona
lobi interiores
Umbellae
viridiflavide
coronae
integrarum partibus
acuminati
—
Province:
shore
acuminata
paulo latiore;
alis minutis.
C Sulawesi
longi.
coronaque
lateralium
the north-western
10
Fig.
longa
Corolla
Caudiculae
(holo L), Indonesia,
—
curvati
valde recurvata
superantibus.
ridge forming
nov.
of Lake
apicibus
eis
Typus: Kleijn
c.
10 km west
Poso.
was
original
m.
nervatura
alba;
cm
outer
minahassae is endemic
moss
Petioli
the trunk
attached, caudicles with long wings made
are
the lateral
Here it grew
on
are
excretions. [This
Little is known of this
above 800
Hoya
morphology
against
acuminate, venation absent but
acute to
vertically,
Lake Poso, which is
7.
15.
(1916)
positively geotropic,
concave,
Sulawesi) facing
800
2
34,
densely hirsute, patent to slightly campanulate.
entire caudicles rather than
some
closely
top and broad, protruding 'hips', retinaculum groove widens significantly
species compared
specimen
species
flower
flowers
would also
Province, Minahassa, Tomohon.
honeycomb-structured
areas
containing
9
Fig.
downwards from where the caudicles
of
less
and corolla white, overall coronal lobe
corona
lobes acute, turned
ridged,
so,
similarities in
more or
wherethe midrib is. Petioles 1-2
short (0.5-1.5 cm),
Flowers:
—
Leaves succulent,
fleshy.
depression
a
this
and H. caudata Hook, and H.
species
Beih. Bot. Centralbl.
Schltr.,
N Sulawesi
B), Indonesia,
for
trees)
placed
suggest close kinship.
Kloppenb.
6.
of this
pressed
justifies
and H. imbricata Decne.
herbarium sheet
a
imbricata. The
habit (stems and leaves
growth
genus
We have therefore
taxa.
of H. imbricata and branches of Dischidia imbricata. If
be
the
think that the variation in above-mentioned vegetative traits
of these
separation
maxima
not
Hoya
Donkelaar:
van
478
Fig.
c.
BLUMEA
10.
Hoya
myrmecopa
flower in median
indoors
ants
on
IPPS
in the
d.
ridge
on
& Van
pollinaria;
Netherlands, growth
limestone
8840).
Kleijn
section;
habit
Vol.
46,
Donkelaar.
e.
a.
No.
3,
2001
Flower in
top view;
growth habit (flowers
drawn from the
the north-western shore
drawn
b. flower in
from live
plant growing from the
of Lake
Poso,
which
side
material
view;
grown
nest of Tetramorium
is also the
type specimen;
D.
Leaves 2.5-7.5
0.5-1.5
&
Kleijn
Donkelaar:
van
long,
cm
ecology of
and
Taxonomy
elliptic,
ovate to
Peduncles 1-2
long.
cm
R.
acute to
acuminate, venation obscure. Petioles
sions that
are
inner coronal lobe acuminate,
much elevated above
outer
lobe
tips,
la. Pollinaria caudicles with small
Distribution
Ecology
was
as
growing
had
was
the
found
Note
being
to
—
without any
The flowers
markedly
this
sign
in floral
at
in
that
originate
It may represent
are
an
in this
and
'ope',
found
8.
or
which
in the fork of
from
sections
knowledge
name
to
species
proved
to
be
also
was
a stem.
Hoya species
within the genus that differ
H. myrmecopa is the
only species
umbel and bilobed coronal extensions.
1,3, 11), which have
outer
coronal lobe,
ex-
the
of the entire lateral sides of the lobe.
link between the sections Acanthostemma
'hole' in Greek, and refers
of
myrmecopa consists
the niche where this
(e.g. Fig.
'myrmex'
species
was
'ant'
or
commonly
growing.
Hoya pallilimba Kleijn
Caules
cm
tenues
tropicae
—
sine
5
saepe
nervatione
cm
parum
fere
alis magne
margins
than 5
cm
glabrescentes.
valde
verticaliter
Folia ovata 3-8
alatis, retinaculo
pallide
Umbellae
recurvata;
sursum
in
viridis.
positive
geo-
coronae
erectis;
lobi
interiores
medio sulco latissimo.
(holo L), Indonesia, C
glabrescent.
Leaves 3-8
cm
basal half of the leaves thin,
long,
geotropic,
15-30 flowers. Flowers:
vertically
upwards,
outer
longi.
cm
dimidio basali
Sulawesi
Province:
mouth of Ranu River.
near
on
more
apicibus
in
11
Fig.
—
0.5-1
intus hirsuta
Van Donkelaar IPPS 8864
&
Stems thin, hirsute when young,
Peduncles
demum
15-30. Corolla
erectis. Caudiculae
Reserve,
obvious venation,
hirsuti
exteriores
bilobatis;
Typus: Kleijn
Morowali
Petioli
nov.
manifesta, marginibus
longiores
concavaefloribus
extensionibus
apicibus
& Van Donkelaar, spec.
juniores hirsute, glabrescentes.
longa acute
Pedunculi
cm.
Once the
area.
(Fig. lOe)
Once, it
genus Tetramorium.
integral part
3,11) and Hoya (e.g. Fig. 4, 7). The
on
found
material
ant nest
well apart from any other known
our
that grows
myrmecopa
usually
was
cavities
from the lower lateral sides of the
phylogenetic
a
the north-
forests
open rain
It
range.
most
branch
of the
negatively geotropic
extensions of H. myrmecopa
Hoya
trees.
relatively
However, unlike species from section Acanthostemma (Fig.
tensions
on
the time of observation which may have
tree
a
ants
species
To
morphology.
a convex,
ants
of
species
characteristics of
combining
that has both
set
tips
above corol-
rarely twining species
prolonged dry spell
a
small hole in
a
Although
abandoned
cavities inhabited by
growing
small leafed,
from the main branches of
branches.
tree
considerably
found in central Sulawesi
only
from the Tineba mountain
the end of
from
growing
entrance
was
a
dangling
generally
were
do with it
to
observed
in
they
elevated
the north-west of Lake Poso in the
on
from cavities in
seen,
tips
600 m).
(±
ridges extending
exten-
wings.
myrmecopa is
Hoya
found only
limestone
was
—
cluster of stems
a
lobe
outer
myrmecopa
shore of Lake Poso
western
usually
Floy a
—
greenish/
corona
coronal lobe with bilobed
outer
of the entire lateral sides of the lobe, inner lobe
integral part
an
convex,
sparsely hirsute, strongly
recurved, only marginally wider than the coronal lobes, corolla and
yellowish white,
479
Hoya
genus
long, thin, generally upturned. Umbels
cm
6-16 small flowers. Flowers: corolla
negatively geotropic,
the
hirsute
corona
to
glabrescent.
yellowish,
long,
pale
ovate, acute, without
green.
Umbels
Petioles 0.5-1
concave,
inner coronal lobe
coronal lobes have bilobed extensions with
positively
points
tips
almost
that
point
BLUMEA
480
Fig.
c.
11.
Hoya pallilimba Kleijn
flower in
indoors in
species
in
median
the
section;
d.
Vol.
& Van Donkelaar.
pollinaria;
e.
Reserve,
which
a.
No.
3,
Flower
2001
in
top view;
growth habit (flowers
Netherlands, growth habit depicts
the Morowali
46,
is the
the
plant growing
type specimen;
drawn
b. flower
from live
in
from the stilt-root of
IPPS
8864).
side
material
a
view;
grown
Pandanus
D.
&
Kleijn
R.
van
Distribution
Ecology
in
Pandanus
however,
plant
a
The
species.
ants
found
was
opening
level
of
by
ants on
log
a tree
alluvial soils
on
just
not
the inside,
on
been observed
recurved.
strongly
the
on
shores of
eastern
with
found
was
As
ant
only
without any obvious
Both
plant.
plants
grew
at
sea
far from the Ranu River in the Morowali Reserve. As indicated
not
soils (Soroako-Malili Road,
Van
Balgooy 3820, L),
on
species growing
coral-chalk
on
ultrabasic
Is-
(Tukang-Besi
lands, Elbert2537, L) and in Opa swamp (Polipolia 200-300 m, Prawiroadmodjo
Soewoko
Note
s.
n.,
&
L).
resembles H.
—Vegetatively, Hoya pallilimba
tinctive is,
a
carton;
The second, and
root.
growing
far away from the first
depicted
stilt-root of
a
something resembling
the hollow
opening
species
ants.
little circular hole in
a
the herbarium sheets, others have found this
notes on
481
Hoya
genus
grows in association with
from
closed
was
found when
not
were
hirsute
the
Besi Islands.
growing
other observed individual of this
signs
has
species
Tukang
Hoya pallilimba probably
—
1 le,
Fig.
the
on
ecology of
retinaculum with very wide groove in the middle.
large wings,
So far, this
—
nearby
and
Taxonomy
corolla dull whitish red,
slightly upwards,
Pollinariacaudicles with
Sulawesi and
Donkelaar:
however, the thin,
sides
of the
petiole
until
name
of the
species
is
Latin. The thin
pale
green
halfway
approximately
derived:
for
'palleus'
quite closely.
From this
and 'limba'
margin
the
meaning 'margin'
material is dried, e.g.
plant
Dis-
downwards from both
the leaves.
'pale'
is also distinctive when
margin
myrmecopa
margin extending
on
in
herbarium
sheets.
ACKNOWLEDGEMENTS
The authors
comments
Provision
of flowers
sions
with,
the translation of the
greatlyappreciate
insightful
Dale
on a
by
draft version
Ard
Vogel
is
by
English diagnoses
His
Veldkamp.
Green
were
a
great
in Latin
as
well
help greatlybenefited
gratefully acknowledged.
and Ted
Kloppenburg
J.F.
The
help of,
and
the
many
as the many
manuscript.
lively
discus-
stimulus.
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(Leptospermum and Dacrydium)
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Weissflog, A.,
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W.M.
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Ants,
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J. Linn.
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R. Hashim
30: 45-51
Malaysia
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Dischidia
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113-132.
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myrmecophytic epiphyte from
Maschwitz.
1999.
Southeast Asia with
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5: 221-225.
their structure,
development and
behavior.
Columbia
University
Press
D.
& R. van
Kleijn
Donkelaar:
and
Taxonomy
Identification
The numbers
Alphonso &
Blume
after the collectors
3;
Chai s.33565:
21:3
3
H. brevialata
5
=
H.
=
H.
camphorifolia
6
=
H. minahassae
3
=
H.
coronaria
7
=
H.
4
=
H.
dolichosparte
8
=
H.
Edano 9588:
—
3;
Fox
4192:
8975:
Hallier
Ismail
2;
2
3;
5;
5793: 4
34523:
Frake
36055:
Kerr
—
5;
3
Posthumus
Ramlanto
3
—
3.
Church & Ismail
—
5984: 3.
II:
III: 3
3;
Elbert 3401: 2
3;
3
J.C.
1551: 3
Kleijn
—
8845:
s.41868:
—
Dransfield
—
Elmer
26847:
3
8;
—
Leonardo
—
3;
Jensen
—
15244:
400: 3
& VanDonkelaar
8864:
2;
Maxwell 80-39:
—
3
—
series:
Toxopeus
7;
22126:
3;
Lee
RAM.168:
39912:
5228:
40154: 3
3;
2;
4164:
15802:
5
3.
—
Eyma
4081:
8891:
3
—
Kornassi
3531:
85-916:
2
Johansson
—
7718:
(all IPPS)
Koorders
688:
3
1;
6;
8831:
4;
16187:5; 16189:5; 16192:5;
Kostermans
—
105: 2.
8816:
6: 3.
5.
Meyer
—
9997:
3.
627: 3.
Rant 392:
Sandakan
De Vriese
Buwalda
—
3.
12447: 3
8840:
16237:
—
1125: 3
Maingay
—
—
Coode
3
s.38246:
3.
1;8838: 5;
16201:
5
—
Ching
—
3.
173: 5
Lam 2659:
3
—
Ramos
Ridsdale
SAN
17603:
1118: 5
73660:
3
6513A:
3
—
—
20500:
3;
Rutten
3;
41265:
5
—
Ramos & Edano
36729:
3
—
164: 3.
Schlechter 20434:
6;
20642:
4
—
Schmad
486: 3
—
Sinclair
3.
58: 3.
Balgooy
Weir
5728:
s.n., Fort A. de Kock:
8836:
Van
80526: 3
Buwaya
11015:
—
98903:
Kaudem
Chew Wee-Lek CWL 513:3
&
myrmecopa
pallilimba
3.
898.168-121-125, 898.168-128):
no
imbricata
3.
247:
Jacobson
A.236:
(L
Vogel 5661: 2;
De
to:
=
Conklin
—
refer
1
s.n.
483
Hoya
genus
list
numbers
collecting
the
2
Samsuri
483:
and their
ecology of
4867:
3;
1: 5 —Whitmore FRI
—
3707:
Van Niel
3745:
3.
3.
Index
Accepted
the
names
species
are
numbers
Asclepias
Coronaria
type,
given in
Roxb.
new
names
3
5
and
hollrungii
imbricata
Koord.
5
3
camphorifolia
Warb.
Blume
coronaria Blume
3
papuana
dolichosparte
F.M.
Donkelaar
2
corona-ariadnes
1
Blume
Decne.
refer to
ex
Decne.
Bailey
Kleijn
pseudomaxima
3
sussuela
velutina
Koord.
5
Warb. 5
Schltr. 6
pallilimba Kleijn
3
3
5
(H. Karst.)
myrmecopa
speciosa
Schltr. 4
Numbers
Warb. 3
minahassae
& Van
Kleijn
italic.
forma basi-subcordata
maxima
ariadna Decne.
in
synonymy
grandiflora
punicea
R.Br.
var.
bold
(Hoya)
Karst.
3
maxima
brevialata
in
the text.
maximum H.
ariadnes
Rumph.
Hoya
roman
as
sussuela
Conchophyllum
Dischidia
in
& Van Donkelaar
& Van Donkelaar
Koord.
5
Decne. 3
(Roxb.)
Wight
3
Merr.
3
8
7