BLUMEA
31
39-55
(1985)
The Winteraceae of
V.
Exospermum links
the Old World.
Bubbia to
W.
Zygogynum
Vink
Rijksherbarium, Leiden, The
Netherlands
Summary
Recent
collections
tween Bubbia
and
ly, Zygogynum.
new
of
The
combinations
The
oldest name
made; two
new
species
break-down
for the
these
characters, relating to
are
the
Exospermum completed
Zygogynum.
reductions,
and
of the
are
reviewed.
subspecies
one new
differences be-
generic
resulting enlarged
genus
One
unfortunate-
is,
name
new
and
30
described.
are
Introduction
In his
already
(=
treatise of the
winteraceous
new
existing
homoxylous
genera
Exospermum
nised
two
species,
these under the
To
tween
stipitatum
name
Drimys,
carpels
locules. In
and
firmly
are
they
Exospermum
carpels;
in
a
link':
the
relic of
carpel,
a
situation similar
ovules
on
the
they
Drimys
angle
are
I: Blumea
525;
18
to
that in
sealed
one
an
side,
to
the
Tieghem
v.
locule
abaxially
Van
or
(1942)
are
on
its
was
Zygogynum
are
or
Tieghem
recogunited
fully support.
and
the ovules
of the
inserted
ex
Guillaumin
Exospermum
carpels coherent,
c.s.
Forster
gynoecium.
already
on
united into
original separation
(1983)
Zygogynum,
(adaxial) carpel
(1970) 225-354;
IV: Blumea 28
on
in addition
position
be-
the other. In
an
ovary
with
in the walls between the
very
weakly
inserted in
the
a
so
in E. lecartii
position
(sealed) margins
where
of the
'on either side of the median
'reversal of placentation'. In E. stipitatum Van Tieghem observ-
median vein, and between these
*
but
Zygogynum they
he found the
internal
Zygogynum
vein' of the
ed
in
lecartii,
of his genus
free and in
a
Wintera
reduction which I
a
and Bubbia
in E. stipitatum. In
'belong',
and E.
importance
several locules and without
of the
characters
stipitatum,
Belliolum,
the
c.s.
E.
the
Tieghem
Van
Drimys
E.
by
erected the
Tieghem (1900)
Exospermum
Baillon.
Zygogynum
defined
was
and
Bubbia,
J.R. & G. Forster,
Drimys
Pseudowintera Dandy), and
Van
dicotyledons
genera Belliolum,
positions
on
II: Blumea 23
311-328.
but in E. lecartii he found the
margins,
on
either side of the
'missing
(abaxial)
the lateral walls.
(1977) 219-250; III:
Blumea
24
(1978)
521-
40
BLUMEA
Van
he
is
mentioned only the
Tieghem
indicated
never
of
occurrence
difference between
a
No.
31,
of ovules in each
two rows
and
simple
1985
1,
multiple
Burtt
than of
Exospermum
to
be included in
to
Drimys
tajania
are
comparison
a
endemic
as
outlined
as
by
Caledonia,
Tieghem.' However,
van
showed that
spermum, and
Madagascar respectively.
Caledonia thus harbours
Bubbia
we
to
appears
(though
in
leaves of
different
a
a.
an
B.
howeana,
in the
are
have united
axillary
structure
shares
a
carpels
normal
to
and
a
calyx
number of charac-
in this genus
states
comparison
and
also for
a
to
be
to
Bubbia
are
of the
twig
will
typical
turn
first
for Bubbia,
considerable part of the
have in
Zygogynum
con-
common
a
only
not
species.
terminal inflores-
in fruit.
as
following important
generally conceived,
with B. howeana
characters:
bear
inflorescences
partial
we
which is
Vink, 1983),
see
cataphylls
branched
in Exospermum
connate
carpels:
d. ovules:
(the
more
pollen:
to
free in
than
flowers in
solitary
Exospermum
in my earlier
('florescences'
and
publica-
exine
level:
species
of
most
one
are
ofZygogynum
species
always free),
Exospermum,
row
on
each
the
petals
in B. howeana all
free in B.
placenta
howeana,
in
of the
petals
connate in
Exospermum,
in
outer
are
series
free.
Zygogynum.
one row
in B.
Zygogynum.
structure
reticulate in B.
ploidy
and
inner petals
coherent
in
howeana and
f.
monopodial
Bubbia
Winteraceae,
in B. howeana.
tions)
mum,
a
Exospermum
and therefore the character
morphologically
inflorescence: the
b. petals:
e.
ZYGOGYNUM
Takhtajania
and
has
Drimys
differences of Exospermum,
principal
Zygogynum,
c.
and
early rupturing calyx persistent
Zygogynum
are
AND
Pseudowintera has its inflorescences
description
but
Exospermum,
The
BUBBIA
Zygogynum
very variable genus. For this
full
(for
nomenclaturally
and
relative.
ways),
WITH
well.
Bubbia is
and
in
to
Exo-
(including Belliolum),
with the other genera of the
Exospermum
closest
Zygogynum
as
howeana
cence
EXOSPERMUM
the bud until anthesis. However,
with
sidered
compare
be the
OF
monopodial branches,
enclosing
Bubbia
occurs
endemic
are
Zygogynum.
COMPARISON
When
Zygogynum
and South America. Pseudowintera and Takh-
Malesia, Australia, and New Caledonia. Exospermum and Zygogynum
New Caledonia. New
of Bubbia
former, although
well.
New Zealand and
to
it
retaining
in his intro-
wrote
suggestive
more
be combined with the
McPherson in New
by
Malesia, Australia,
in
occurs
in
justified
the genus is
effect,
might conceivably
made
collections,
'In
(1943):
and
Zygogynum
for the present I feel
recent
united Belliolum with Bubbia. A.C. Smith
(1936) already
duction
ters
locule;
this difference
so
rows,
amongst the original characters delimiting Exospermum.
not
has
VOL.
-
n
=
43,
Zygogynum;
with
tectum
howeana,
known
perforatum
to
microreticulate in
and with both types in
for three
unknown for
species
of Bubbia ;
Exospermum.
Exosper-
Zygogynum.
n
=
86,
known for
one
W. Vink:
Exospermum
Review
Inflorescence.
distal bracts often have
minal
which
partial
In
—
the
supplies
On this theme
In
a
and
and
of the
nodes
bract
buds
of
an
tative buds enclosed
at
above
the
tera
Sampson,
In
the
Drimys
only
are
one
flowerparts
is
confined
All
the
cled. The
bracts
sect.
have
such
but
only
*
(Vink,
In earlier
cataphylls
only
flowering
and short intera
from terminal vege-
a
enclose
situation
a
given
secondary
(Vink,
inflorescence is
base,
are
sect.
In
of
publications
I
twig
I found
after
to
pattern
leaf axil in consecutive
replaced by
a
vegetative bud,
a
they
can
f.
is
buds
on
or
clues in the herba-
monopodially
resulting
from
new
a
vegeta-
shoot has
non-flowering twigs
was
incorrect
as
no
do have
sometimes
the axils of bracts
partial inflorescences
terminology is
the
fasci-
well-developed (at
are
references
15)
are
so
from
range
(Vink, 1970,
flowerparts
no
1944;
vegetative
reduced and the flowers
inflorescences in
to these
that this
replaced by
buds in the axils of bracts
vegetative
me
is
the
on
(Nast,
that in Drimys.
to
flowering
vegetative
Secondary
have referred
Weberling kindly pointed out
In Pseudowin-
of the
inflorescence
Tasmannia
completely
Drimys
but terminal
1970, p. 251).
1963).
axils of bracts and
the flowerbuds until all
of the
1970)
comparable
flowers.
sect.
a
flowers
solitary
integral part
an
in
produced
are
sometimes
partial
solitary
is
of
elevates the flowerbud
elongating pedicel
inflorescences in the
to
leafless lateral short shoots in the axils
initiated (Sampson,
are
terminal flowerbud
arises
cataphylls. Development
observed
brac-
by
the flowerbud
flushlike, without cataphylls, and terminal
is
but
Tasmannia ; for
its
subtended
are
bracts.
flowers.
solitary
been initiated. After
tive bud in the terminal position in the inflorescence. The
at
to
some
inflorescence is formed in the axil of
inflorescences
secondary
material). Elongation
cataphylls
dichasia
inflores-
partial
axils of
inflorescences consist
partial
flowerparts
terminal
partial
to
is present. The
leaves,
case
dichasia
terminal
Non-flowering twigs elongate
and bracteoles of the dichasia
least in
rium
or
a
and enclose
fugacious
secondary
a
way
scars.
before all
normal
1963)
compound
axes
or
in the latter
least
compound
develop leaves,
not
Vegetative elongation
seasons.
there
their
formation of
twig bearing
shoot;
all
to
1970, f. 3) and hence flowers
(Vink,
at
and lateral flowers
axes
usually
are
flowering
least the terminal
the bracts
bud
number
by cataphylls.
In Pseudowintera
and
a
a ter-
a
described above, from buds in the axils of lower bracts.
as
do
all have
range from rich
old inflorescence.
of normal leaves
Often
flowering.
inflorescence is present in the
lateral
In this
(short shoots).
(scar)
can
some
twig is,
Sometimes these
after
twig
more
there is
axils.
Zygogynum
partial
bracteoles
till about the time that
elongation
usually
few of the lower bracts subtend
a
of the
elongation
dichasia the
(compound)
teoles. Bracts
or
seemingly empty
inflorescences
partial
The
also
usually
one
Usually
bears
flowering twig
of the leaves. The
2/5 spiral
number of variations is realised.
a
Exospermum, Bubbia,
and
of the
apex
the
inflorescences* in their axils and
sympodial
of the lowest bracts have
cence
the Winteraceae the
partial
inflorescence.
41
Zygogynum
-
characters
of
cataphylls (bracts) continuing
crowded
densely
Bubbia
-
'florescences',
were
but Dr.
for closed systems.
42
Fig.
BLUMEA
1. Petals.
connate;
15584).
s.n.).
—
b.
—
e.
&
—
c.
a-c.
two
d.
Z.
&
f. Z.
-
VOL.
31,
No.
Zygogynum (Bubbia) pauciflorum.
views
(Bubbia)
of
an
flower,
open
howeanum.
Open
in
a
regular
way
e.
and
1985
a.
corolla
flower
(Exospermum) stipitatum.
flower, petals rupturing apart
the
1,
Bud, calyx minute, petals largely
irregularly ruptured (x 6; Carlquist
bud, petals
free
Bud, petals halfway
(x 2½;
MacKee
25819).
(x 5;
connate;
R.B.
f.
Chinnock
just opening
W. Vink:
but
found,
not
sometimes the
shoot
flowering
the bract like
to
tary flower is
In
consist of
The
acter
and
Bubbia,
Although
mentioning
a
Petals.
nea)
the
petals
petal
blades and
few
a
side, providing
a
resulting
'free'
also
a
anthesis the
to
very
but
the
original
reason
one
petals
is
in
and
also
free,
but
secondary
The union of the
21204)
and B.
completely
the
e-g-
same
to
this
1
some
of the
in
the
thin
ring
less
in
petals
most
petals
of
Gui-
(New
thickening
from the adaxial
petals rupture apart
ruptured
tissue
the
petals
apices
is almost of
line of
so
of the
equal
of the
the ab-
along
has been overlooked for
long.
petals
thickness
rupture is absent and upon
irregularly in B.
spec.
(MacKee 21204)
pauciflora.
other
are
species
free.
In
of
Bubbia,
that in
this
and in
Drimys,
Zygogynum pomiferum
buds have been found with their
length, suggesting
all
Zygogynum (Vink,
between the
of
anthesis the
strip
of
in B.
all
(almost)
is thinner than the blades of very
compressed
predetermined
d),
deviate from
Bubbia pachyantha
pauciflora (fig. 1, a-c)
more or
even
and have
shape; only the margins (except
ring
in
in Z. vinkii.
one,
worthwhile
genera, it is
described for
as
way
phenomenon
the
(2—)3—5
inflorescences:
B. comptonii,
rupture. Upon
a
char-
specific
species
petals
the
connate
occurrence
the
for up
of free
state.
petals
pauciflora
free
a
a
the latter genus the reduc-
time.
and
f)
1-, 2-, and
pauciflora.
the terminal
one,
same
&
e
incompletely
Takhtajania
partial
con-
species again they always
and B.
in
they
queenslandi-
reduced and
centrifugal flowering
'normal'
and B.
a
But
connecting part
the lower part
and often
fifth of their
a
of
in e.g. B.
flower is thus
single
comparison
union by
why
However, in B. howeana (fig.
to
only
Bubbia,
at
line of
petaloid ring ruptures
irregularly
are
of
in the
cross-section. Here
Pseudowintera,
petals
the
(fig. 1,
have
(MacKee 21204)
free,
throughout
and
epidermis penetrates
petals
of the
side, probably
In B. spec.
are
to a
1);
distinctly
in other
(MacKee 21204)'
picture
There the
3).
predetermined
apical part) testify
axial
species
general
double
a
f.
p. 313,
are
in later stages this thinner part is
petals;
and the
for
almost perpen-
variable. In B. howeana
are
character in Zygogynum.
are connate
f.
they
inflorescence
In Exospermum
—
1977, p. 229-230,
young
partial
pedicel,
indication that the soli-
an
1977, f. 1 & 2) the partial inflorescences
they
inflorescence;
e.g. 'B. spec.
inflorescence open
given
outer
whiteana
same
e.g..Z. baillonii,
the for the Winteraceae
flowers of
short leafless
inflorescence.
partial
(Vink, 1983,
triads
importance
here that
branched
reduction in the number of
a
of
not
a
var.
generic
1—3 in
pomiferum,
the insertion of the
at
but in Bubbia
to
solitary flowers,
a
of
in the
ones occur
reduction of the
in
a
regularly ( Pseudowintera), occasionally (Zygogynum),
Zygogynum (Vink,
dichasia
tion continues with
Z.
state
semecarpoides
B.
3-flowered
produces
known.
sufficiently
not
the bracteoles of dichasia. This is
solitary flowers,
compound
and
ana
reduced
a
Exospermum
consist of
sist of
is
rarely (Drimys) (a) bracteole(s)
dicular
43
Zygogynum
-
terminal inflorescence bud
Takhtajania
flowers have rather
Solitary
very
Bubbia
-
(Vink, 1970, Drimys insipida).
The inflorescence of
or
Exospermum
can
be considered
than in
completely
the other
connate
as
taxa.
outer
more
complete
in B. spec.
Bubbia then shows
petals.
(MacKee
the full
range
of
BLUMEA
Fig.
2.
Pistils
of
(patches
The
of torn
31,
VOL.
No.
Zygogynum (Exospermum) stipitatum.
coherent
cally strongly
-
(x 3;
epidermis
form of the
on
MacKee
two
calyx,
the
apices
free
—
and also
—
d.
&
(x
b.
6
(MacKee
calyptra
a
a.
connate,
Carpels basally
Carpels completely
coherent
of B. spec.
which is also
&
c.
carpels) strongly
petals
outer
minds of that of the
overlapping
15614).
1,1985
McPherson
;
44
or
api-
locally
very
6129).
and B.
21204)
with the
free
pauciflora
apical opening
rupturing irregularly, although
at
a
re-
closed
by
very
early
well
as
stage.
Carpels.
—
In
Pseudowintera and
the
carpels
brings
are,
us to
Van
at
B. howeana and
Drimys,
carpels
are
on
E.
connected
contact,
point.
two
By
are
its
to
intimately
each other
distinct
light
over
by
material,
other Bubbia
(or solitary).
coherent but
species,
In
some
as
Bubbia
essentially
pistil
thus
composed
shows
their lateral faces of which the
double
contrasts
the cells filled with
on
in
species
also free. This
yellow
the outside
united with their lateral faces into
their whole
colour this
contiguous locules,
which
free
stipitatum: 'Being separated
A series of cross-sections of the
in
many
are
Exospermum.
....
only
in
carpels
least in herbarium
Tieghem reported
grooves, the
the
surface;
vividly
essential oil
no
two
give
that the
neighbouring
that
colour
carpels
are
which
are
concrescence
every
to
by deep
single body;
epidermes,
which divides
epidermis,
with the
there is
a
at
any
separation
cortical
layers
the entire
of
of
carpel.'
W. Vink:
3.
Fig.
tissues
(all
flowering;
bud;
of the
6).
x
6063).
2975).
13832).
section) supplying
double
epidermis;
carpels
are
adherent. A
In
free
fact,
(fig. 2,
firmly
ing
c
coherent
from the
completely
not
quite
&
d).
reported:
only
free
a
not
is
In
one
Bubbia
fruitlets
also
at
lines
almost
not
line
carpels, branching
d. indicate
variations
base
of the
middle
locules
close
(just
past
(submature
(just past flowering;
below
to the
'A series of cross-sections of the
concrescent,
Van
but
sufficient
Tieghem's
are
to
even
the
surface
termination
of
of the
carpels
the
wide
Ill &
are
two
not
the
carpels.
species,
trace
of
the
In
ripe
rupturing
principally
shows also that
completely.'
carpels
I observed
variation of
VI).
very
specimens
pistil
faintly united, hardly
them
separate
without any
1943, pi.
in
locules
of locules
slightly
in
differences in
right (just opening bud; Carlquist 15590).
(carpels)
& Nast,
&
c.
the
where
specimen (MacKee 13732)
by
in
to the
separating
broken
(as
of three
already
their form fits
(see Bailey
broken
and another flower with 4 free
covered
45
Zygogynum
-
epidermis extending to
epidermis
bundle
On the basis of the above the
Bubbia;
curved
number of Exospermum
carpels
area
Double
parts
carpel
slight boiling
in
b.
Double
c.
Bubbia
epidermis extending
—
different
abortive
On E. lecartii he
the
—
carpels;
d. One vascular
—
-
Zygogynum (Exospermum) stipitatum, showing
of the
Double
a.
—
of
pistils
continuity
McPherson
McPherson
MacKee
the
of
Longisections
the extent
Exospermum
are
one
completely
flower with 7
material, originatfruits
always
of the
show
epidermis.
different from those of
carpel
forms
encountered in
46
BLUMEA
Fig.
4.
Fruit
2739).
the
—
a.
Apical view;
continuity
of the
Takhtajania
There is
(Vink, 1977).
has
no
of
carpels
b. lateral
VOL.
31,
No.
1,
1985
Zygogynum (Exospermum) stipitatum (x 1¾;
view,
from
D;
c-d.
longitudinal sections showing
McPherson
variation
in
carpels.
has
Zygogynum
les.
connate
with
-
a
bicarpellate
2—°°
trace
This is
unilocular ovary
carpels completely
of
original
an
clearly
(Leroy, 1978; Vink, 1978).
united into
in
separation
the
an
ovary with
walls
different from the situation in
many locu-
as
the
separating
Exospermum
as
locules
described
above.
However, in Exospermum
Massif
(New
Caledonia)
the
collections
ripe
recently
fruits have
made
connate
by
that this fusion is also visible in flowers. There the double
down
tions
to
the base of the
carpels,
of fertile material from
various
adjacent
degree,
to
the
a
fusion of the
stamens
are
that is
to a
The
always free,
epidermis
does
level below the locules. In
the Panie Massif
carpels.
McPherson in the Panie
fruitlets (fig. 4). It turned
are
apices
but the
available
to
me
not
all,
out
extend
11 collec-
and all
show,
to
and the parts of the lateral walls
contiguous carpel
walls
are
connate
W. Vink:
5.
Fig.
more
c.
Pistil
of
0.2
over
to
0.8
of their
to
In this way
At
and
this
the
tween
epidermis
32379).
branches
to
the Mt Panie
(and
—
a.
(arrows) slightly
Lateral
view;
b.
(b)
or
longisection;
an
that in
Bubbia
with
Exospermum
is the
pistil
neighbouring carpels (fig. 3d);
'missing
with free
species)
fused
carpels
connate
link' between the remainder
—
whether
or
coherent
not
—
carpels.
is whether
question
with
axial bundles extend into
Zygogynum.
completely
important
cases
below the terminus of the double
different parts of the
to
Exospermum
some
Zygogynum
stage
In several
length (figs. 2,3).
comparable
Exospermum
carpels
double
MacKee
locules, dividing only slightly
two
provide
this situation is
of
pistil (x 6;
the
47
Zygogynum
-
d. cross-section.
the wall between
epidermis
Bubbia
-
Zygogynum (s.s.) baillonii, showing
distinctly (d) penetrating
apical view;
Exospermum
morphological
a
and the
difference be-
Zygogynum
pistil
can
be
defined.
In
with
(in
Exospermum
a
double
walls. In fruit these
ual
carpels
shown
in
still
are
Carlquist,
In e.g. Z. bicolor
pistil only
vidual
carpels
Z. baillonii
this
species
til
(fig. 5)
is
no
mum
are
by
their
even
epidermis
and this situation
morphological
and that in
up
on
till
there is
now)
and abaxial
as
a
double
possible
zone
contiguous carpel
material,
pers.
apices
but
(The
fruit
originated
comm.).
are
pomiferum (I.e.,
epidermis
recently
completely
distinction
Zygogynum.
In Z.
delineated and their
was
a narrow
c.q. fruit.
pistil
illustrative
Caledonia; Carlquist,
stigmas.
least
of
Both in flower and in fruit the individ-
the outside of the
added
at
margins
1977, f. 7c) the individual carpels
distinctly
8a-c)
double
was
of New
centre
(Vink,
more
f.
apical
outlined
1982, f. 12,
indicated
(I.e.,
a
the
epidermes neatly separate.
distinctly
from Katrikoin in the
the
all material known
epidermis along
are
between the
on
the outside of
f. 6 &
7b)
more or
apices
the indi-
less free. In
was
found. In
also found in the lateral walls of the
links up with that in
between the
Exospermum,
fusion of
carpels
in
so
pis-
there
Exosper-
48
BLUMEA
Ovules.
ovules in
that there
E.
mentioned
differences in
but
the
he
of E.
described the insertions of the
unlucky chance,
some
the insertion of the
type specimen
1985
1,
Tieghem
by
was,
No.
31,
Van
above,
Apparently
rows.
were
lecartii,
As
—
two
VOL.
-
also
stipitatum
shows
believe
to
made
ovules between E.
and
stipitatum
what he de-
exactly
scribed for E. lecartii.
A further
spermum
that is
development
'scattered
as
(1943,
477). Curiously
p.
discussion. A median
covering
that there
son
in
are
& Tucker
ovules
tion in
B. spec.
of ovules
21204)
them
fall
single
a
or
three
monads have
a
B. spec.
could
equally
level.
Ploidy
counts
three
(Ehrendorfer
species
all in
the
number
of
same
size of the
Up
a
'We
Exospermum:
irregular
retarded
num-
causes
prefer
rather than
row,
with almost
to
two
equal justification'
or
be
seen
Drimys,
and
Bubbia,
1968)
one
is
to
now no counts
differentiating
this is
a
as
an
to
be
all
be
in
Drimys
tetraploid:
sect.
n
=
by Bongers (1973)
for
Exospermum
have
Zygowith
species
have
specific
level
In
fact,
character
of
on
the
Exospermum.
with reticulate
reticulate
and
Counts
Australia,
this may
pollen
as
exine
published
to
same
found:
for
chromosome
n
be related
my
as
Caledonia,
=
43.
to
knowledge.
Zygo-
the
be characteristic for the
come
exine.
the number of
are
and New
1; here the
was
pollen
local) development.
be assessed
Drimys
to
in
whereas the other
purpose.
86;
However,
species
cannot
low for this
to
species investigated
and the
have
B. howeana in table
as
(tetrads)
(specific
a new
tetraploidy
too
in all
character.
Exospermum
Takhtajania
perforatum
tectum
as
for the single species
from each New Guinea,
group
appeared
occur.
defined
is
microreticulum,
perforatum
—The role of
c.s.,
shed in
tetrahedral
permanent
found: Z. acsmithii
true
can
are
monads
exine
Zygogynum
in
stomata, reported
till
the latter also
In Bubbia the exine is,
well be
tectum
species
single
pollengrains
good
are
in Pseudowintera and
as
gynum baillonii
genus.
a
perforatum
(MacKee 21204)
I consider the
a
description
of the
1979).
Therefore,
As Pseudowintera,
develop-
similar situa-
50—100, B. spec. (MacKee
c.
conclude for
in
Samp-
found. All have much higher
was
Exospermum
Zygogynum
structure
tectum
their
a
50—70, and the close packing of the ovules
either
structure
(Praglowski,
reticulate exine.
and that
well,
of
reticulate. This would be
far,
line
out
demonstrated that the
they
in B. megacarpa;
; in
one
& Tucker
adopt
can
of ovules.
during
being arranged
species
both types of
gynum
a
as
(1966) pointed
rows
photographs).
the
Exospermum
microreticulate
so
some
three
found
In the Winteraceae the
—
tetrads; only in
In
in
& Nast
deviation in their
but that
placenta,
placentation
by Bailey
the bodies of the ovules
but Leinfellner
two to
of
type
important
indeed shows
walls,
each
and
c.
Sampson
one
also their beautiful
Pollen.
aligned
B. megacarpa
of line.
rows, but
this
over
each with
than other Winteraceae:
out
modified
in different directions. I
on
primordia neatly
consider the ovule bases
(see
row
21204)
(MacKee
40—50,
c.
to
and
carpel
a
placentas,
two
ovules become
with ovule
carpel
bers
initiated in
are
of
of the lateral
a
Lardizabalaceae'
have made further observations and
(1978)
adjacent
ment
locule,
enough they passed
area
reality
of the
Nymphaeaceae
longisection
fairly large
a
of the insertion of the ovules in Exo-
description
the walls
of certain
suggestive
the
was
over
large
whole
W. Vink:
Anyhow,
polyploidy
Ehrendorfer
les for the
genera
Hedycarya
thus
not
decisive
a
Bubbia
-
argument in
and Liriodendron
Magnolia
Siparuna
drawing generic
in the
(Magnoliaceae),
Hernandia
(Monimiaceae),
49
-Zygogynum
(1968) report infrageneric polyploidy
c.s.
and
is
Exospermum
limits;
Magnoliales
e.g.
and Laura-
(Annonaceae),
Annona
(Hernandiaceae),
and
Chloran-
(Chloranthaceae).
Table 1.
Table
1.
*sT
5
x
c
(3
1
bicolor
«
■S
PE(xMaospnterimué) Exospermu
3
Uj
B.
oej
21204
•5
Z.
N
Nor1th South
g£
div.,
1
acsmithi
i
spec.
&
$j
Z.
N'
M1acKe
3
0
&3
B
b
g,
&
Z.
N'
s
pauciflora
05
B.
MscPherson
div.,
spec.
&
spec,
&
43125
g.
e.
u
NGF1
0
tj
1
B.
03
05
B.
B.
03
2.
2.
petals
petals
3.
carpels
carpels
1-flowered
1-flowered
connate
4. ovules
>
>
5.
pollen
microreticulate
microieticulate
s:
carpel solitary;
1
1
row
row
on
on
*
placenta
placenta
etc.
0**
o**
o
0
0
0
oolllooool
1
1
0
o
marily
**
carpels
carpels
is
left out,
connate
petals. Exospermum
carpels.
been
and the
surveyed
The
1
0
loo
1
0
coherent
or
or
New
is
as
as
the
split
states
currently
the
petals,
Guinea
representation
0
not.
comparison
and Bubbia
state, derived from
free
11
lolloooo?oooo
1
l
7
0
0
O
0
0
0
0
0
species-groups, representing
fusion of the
B.
03
0
0
0
0
0
oosoosoo*o
0*
s
0
s
species
secondary
B.
03
1
the distribution of the characters under discussion is
Zygogynum pomiferum
B.
03
"sj
1
l
1
lllllloooooll
1
1
1
0
0
0
0
0
carpels
carpels coherent;
Zygogynum, Exospermum,
B.
05
s
i
1
New
Guinea
In table 1
or
•5
y
£•
0
The
of
£
0
llllllloloooo
1
1
1
1
1
1
1
0
0
0
1
111
1
connate
Sj
—».
1
rNC W
l1
1. part, infloi.
1.
part, inflor.
01
5j
B.
«5
l^o.
New Caledonia
.3
megac rpa sylvestris pachyantha clem nsiae
s
spec.
&
<u
■s
1
s
tj
6
0
3
spec.
&
9
0
l
O
1
E
5
5
1
3
e.g.
howean
6197
into
free
two
for
a
number
of these characters in the genera
conceived.
petals
and thus
species
given
parts
thought
are
not
to
to
be
show
of Bubbia have
of their character
states
may
to
represent
comparable
to
a
pri-
the differences in
only
not
preliminarily
be exhaustive.
50
BLUMEA
The
the
characters
plesiomorphic
young
The table
A
compatibility
of
pairs
four
different
New
In
New
characters;
of the
there
as
the
are
no
The New Caledonia
1
—
2—3,
made and
1
—
of NGF 44125
5, 2
shows that there
(1981)
—
3, and 2
cladograms
pollen
by '1';
too
are
—
(genera).
only
are
5. With these four
based
on
these all have
pairs
paral-
combinationsof
(up
1—2—4
till
now) only
found in
form
of
set
a
as
compatible
character states, the branches
apomorphic
separately
—
have three
sets
of three
2 —5; with these three different
these all have
on
are
be ordered.
taken
and 1
exine
absent from New Guinea (and Australia
characters
remaining
based
be resolved into groups
cannot
—
3,1
states are
cannot
taxa
2—4,
cladograms
indicated in the table
flowers
branches.
higher
resulting cladogram
acters:
—
and microreticulate
Guinea
The
Meacham
be made and
can
these character
Caledonia;
well).
in their
by
to
characters: 1
bipartitions
carpels
Connate
'o\
complex
according
diagram
developments
1985
1,
exine.
reveals that this
compatible
No.
31,
apomorphic states,
indicated
are
pollen
clearly
four
lel
states
check the
to
listed for their
are
VOL.
-
compatible
tripartitions
parallel developments
charbe
can
in their
higher
branches.
The
remaining
that
have
we
whether
is
question
‘North’ and
Exospermum
Bubbia
consider Exospermum
to
Exospermum
If Bubbia,
contains the
most
and
Exospermum,
most
of the
on
the
separation
plus
means
level
combined into
are
This
possible.
level.
specific
represented by
as
Zygogynum
into
‘South’ is
specific
the
on
Zygogynum
primitive type
advanced types
artificial
an
plus Exospermum
B.
one
howeana,
this
Winteraceae, Zygogynum;
is
as
a
genus, this genus
well
as
highly
one
of the
undesirable
situation.
As
far
as
I
can
see
this situation could
could be made into
two
united with Bubbia, and
the
mum
could be united with
Both
E.
the
gether
southern end
table
are
a
North
Van
by
carpels
a
E.
seem
to
and in
to
be part
a
with free
division of
carpels)
(with partly
Exosper-
that could be
fused
E.
stipitatum
Col d'Amieu.
carpels)
that
differences in altitude,
as
(Panie Massif)
and when both parts
a
leaves.
in smaller
However,
are
as
taken
is
to-
clinal variation with decrease in size and
in the
not
and
to
the collections
overlap
reveals
as
specimens
only
differences
a
thicker
from the Panie Massif and
leaf size.
the
is documented by
Katrikoin.
lecartii, belong
Especially
flowers
larger
are
‘North’
and E.
smaller
of
epidermis
correlated with
near
to
South. Leaf anatomy
the altitudinal ranges
Mt Panie and
‘North’
Tieghem,
considerable
thicker adaxial
apparently
Although
on
there is
the differences
lamina and
by
2,
be avoided if
‘South’
of its
differ from E.
range
and
stamens
numbers from
these
one
which is known from Inedete
numbers of
shown in
other
only
E.
one
Zygogynum.
species recognised
‘South’,
from
species,
same, the differences
collections from
are
not
induced
comparable altitudes
W.
Vink:
Exospermum
Bubbia
-
Table
2.
Exospermum
Exospermwn
number of
petals
number of
of
stamens
number of carpels
in
in
pedicel length
pedicel
petal length
in
leaf
leaf in
longest
This
leaves
10—13
10-13
65—188
65-188
97—205
97-205
(incl.
(inch petiole)
7—19
7-19
12—47
12-47
3—8
3-8
14—20
14-20
19—28
19-28
10-22
10—22
1714—26
17&-26
350-800
m
with
only
4—10
4-10
the difference between
20—650
20-650
'carpels
for 0.2—0.8 connate'. Even if the distinction between
acceptable
is
us
10—15
10-15
(1—)2—5
(l-)2-5
mm
mm
altitudinal range in
'carpels
‘North’
'North'
5—52
5-52
mm
cm
'South'
‘South’
(3 or)
diameter in
and I
—
unacceptable
I think
to
51
Zygogynum
-
think it is
—,E.
not
‘North’ and E.
‘South’
completely
are
free'
and
and 0.2 would be
zero
much alike that it
so
them in different genera.
place
that the fusion of the
the other characters and that all
carpels
is
part
a
Exospermum
of the clinal variation observed in
collections
are
samples
of
one,
variable,
species.
CONCLUSION
Bubbia
The oldest
(including
name
The Winteraceae
-
Zygogynum:
rupturing
free
—
to
calyx
a
a
an
following
be united.
after
genera:
flowering;
free
or
after
early stage, probably
inflorescence
the
outer
terminal; calyx
series
flowering;
united; carpels
inflorescence termi-
persistent;
elongation
of
twig;
petals free; carpels
inflorescences
early stage, persistent; petals free; carpels
elongation
after
upon anthesis and
free.
to
pistil.
monopodial
at an
monopodial
free; carpels
at
unilocular
calyx rupturing
have
Zygogynum
plurilocular pistil.
rupturing
Pseudowintera:
Drimys:
consist of the
and
unfortunately, Zygogynum.
probably sympodial elongation
calyx rupturing
-
genus is,
early stage, persistent; petals
united into
united into
—
now
enlarged
sympodial elongation
at an
Takhtajania:
nal;
Belliolum), Exospermum,
for this
flowering;
axillary
inflorescence
usually dropping,
to
leaves;
free.
sometimes
pseudo-terminal;
persistent; petals
BLUMEA
52
VOL.
-
No.
31,
1,
1985
NEW TAXA
Three undescribed
complete.
As
they
Frutex vel
gae, 6-14.5
Pedicelli
3—5,
in
petala
nov.
(1.5-)2-4
30—58
alabastro
sub
connata,
leviter
seriata. Fructus
The
ignotis.
x
cruminatum
3.5-7
album abaxiale
16—19,
c.
Carpella
tis.
c.
u s:
yp
Vink,
nov.
longi.
cm
The
specific
carpels
Zygogynum
basi attenuata,
partiales
c.
uniflorae.
lobatus. Petala exteriora
non
Mt Me
(P),
long stigmatic
10-18
x
5-9
mm;
Pollen in
productis.
Ori,
uni- vel
pauci-
New Caledonia.
crest.
spec.
Laminae foliorum
apice
mm
obovato-oblongae
oblongae,
partiales
200—275 connectiviis
non
leviter lobatus. Petala
productis.
Pollen
carpellorum aequilonga.
Baime
Wau
Creek,
stratum
uniflorae. Flores alabastriis
longi. Calyx subcrassus,
(L),
vel
obtuso vel rotundato, basi acuta;
Inflorescentiae
Kairo NGF 44125
Subdist.,
ignotum.
Fructus
igno-
Morobe Dist.,
like
epithet ('filled
a
purse')
refers
to
the
large
number of
stamens
in the flowerbud.
tieghemii
Vink
(see
nov.
uniflorae,
simul florentes.
—
Typus:
McPherson 6197
N. of Koumac, New Caledonia.
NEW
New combinations in
survive the
53)
page
Vink, subsp.
Inflorescentiae partiales
(L), Tiebaghi Massif,
to
irregulariter
irregulariter separata,
discreta; stigmata apicibus
subsp. synchronanthum
ed
Inflorescentiae
MacKee 21204
(sub)coriaceae,
observatum.
more
avoid
to
Laminae foliorum obovato-oblon-
rotundato vel retuso,
obtectis.
the very
to
picture
B. spec,
Carpella 4—9, discreta; stigmata longissima, apices
u s:
Pedicelli usque 40
35-50,
the
New Guinea.
Papua
and
non
T
discreta. Stamina
T yp
—
cm,
cogniti.
tantum
—
refers
Arbor. Petioli 1.5-2
12—21
as
excendentia; ovula 43—50, in utraque placenta
specific epithet
Zygogynum
albo
anthesi
tetradis exutum, exinio reticulato.
carpellorum
indicated
discreta. Stamina 68—95 connectiviis
8—16,
interiora
strato
longi.
apice
longi. Calyx subcrassus,
mm
make
to
paper
were
Zygogynum.
cm
cm, coriaceae,
stomatibus
this
in
they
spec.
arbuscula. Petioli 0.6—1.5
x
nisi
non
described in
are
cristatum Vink,
Zygogynum
be used
to
of the Bubbiatype,
confusion, but below they
subtus
had
taxa
are
current
Zygogynum
revision.
COMBINATIONS
are
made for
only
those
names
that
are
expect-
W.
Vink:
Zygogynum amplexicaule
Vieillard
308,
Bubbia
—
Bubbia
-
France &
amplexicaulis
53
Zygogynum
-
Vink, comb.
(Parmentier)
Parmentier, Bull. Sc.
ex
10 f. 34.
t.
Exospermum
nov.
27
Belg.
—
Drimys
(1895)
amplexicaulis
231 in obs.,
(1896)
J. Bot. 72
(1934)
(Parmentier) Dandy,
40.
Zygogynum
comb.
amplexicaule
Smith,
Vink
(Parmentier)
nov.
(1942)
Zygogynum
nia
10
293.
bullatum
Zygogynum
54
(1916)
243.
balansae
nov.
294.
Bubbia archboldiana
Bubbia argentea
-
comb.
(Diels) Vink,
nov.
balansae
Drimys
-
(Baill.) v.Tieghem,
Tieghem,
v.
Bubbia bullata
-
comb.
et
nom.
Bubbia balansae
-
Zygogynum
non
—
Vink,
335.
-
A.C.Smith,
436.
(1942)
tieghemii
(1873)
nov.
Vink,
(1900)
433.
Zygogynum argenteum (A.C. Smith) Vink, comb.
J. Arn. Arbor. 23
(v. Tieghem)
J. de Bot. 14
Tieghem,
v.
Vink, comb.
(A.C.Smith)
J. Arn. Arbor. 23
isoneurum
var.
Bubbia isoneura
-
archboldianum
Zygogynum
A.C.
et stat.
nov.
J. de Bot. 14
nov.
—
de
Baill.,
Bot.
(1900)
bullata
Diels,
Arbor.
Arn.
Adanso-
14
(1900)
341.
J.
Drimys
(Diels) A.C.Smith,
J.
Jahrb.
Bot.
23
(1942)
426.
J.
Smith,
Arbor. 23
Arn.
(1942)
Jahrb. 54
(1916)
244.
Bubbia
—
clemensiae
J.
Smith,
Arn.
Zygogynum
J. Bot. 72
(1934)
sonia
sub
3315,
(1868)
Drimys calothyrsa
-
Diels,
A.C. Smith, J. Arn.
Bot.
Arbor. 23
7
nov.
-
Bubbia clemensiae
A.C.
267. —
nov.
Bubbia
-
Drimys comptonii
E.G
comptonii (E.G. Baker) Dandy
(Baill.) Vink,
comb.
nov.
Bubbia crassifolia
-
—
Drimys crassifolia Baill.,
(Baill.) Burtt,
Hook. Ic. PI.
34
Adan-
(1936)
p. 2.
(1869)
(F. Muell.) Vink,
17.
comb.
nov.
Bubbia glauca A.C.Smith
-
433.
(1942)
howeanum
comb.
Bubbia howeana
—
nov.
—
(F. Muell.)
Drimys
v.
howeana F.
Tieghem,
J.
de
Muell.,
Bot.
14
293.
Zygogynum
longjfolium (A.C. Smith) Vink,
J. Arn. Arbor. 23
Zygogynum
megacarpum
montanum
(1942)
(1942)
J. Arn. Arbor.
23(1942)
(1936)
t.
—
Bubbia
longifolia
A.C.
comb.
nov.
—
Bubbia
megacarpa
A.C.
319.
nov.
—
—
Tetrathalamus montanus
Bubbia
montana
(Laut.)
A.C.
Laut.
Smith,
426.
(1913) 71,
sub
comb.
(1905)
Zygogynum oligocarpum(Schlecht.)
Jahrb. 50
nov.
434.
(Laut.) Vink,
in K. Schum. & Laut., Nachtr.
comb.
429.
(A.C.Smith) Vink,
J. Arn. Arbor. 23
Zygogynum
PI. 34
A.C.
41.
199.
J. Arn. Arbor. 23
Zygogynum
comb.
Vink,
glaucum (A.C.Smith) Vink,
Zygogynum
Bot.
Bubbia calophylla
(1942) 431.
(E.G. Baker)
crassifolium
8
Smith,
nov.
comb.
(A.C. Smith) Vink,
Arbor. 23
comptonii
Zygogynum
Smith,
comb.
calothyrsa (Diels)
Baker, J. Linn. Soc. Bot. 45 (1921)
(1900)
-
427.
(1942)
Zygogynum
Fragm.
nov.
436.
calothyrsum (Diels) Vink,
Zygogynum
t.
comb.
calophyllum (A.C.Smith) Vink,
Zygogynum
3315,
f. 1.
p.
3.
Vink,
-
comb. nov.
Bubbia
—Drimys oligocarpa Schlecht.,
oligocarpa (Schlecht.) Burtt,
Hook. Ic.
54
BLUMEA
Zygogynum
A.C.
Smith,
J. Arn. Arbor. 23
(1942)
336.
nov.
pauciflorum (E.G. Baker) Vink,
Baker, J. Linn. Soc. Bot. 45 (1921) 268.
polyneurum (Diels) Vink,
Jahrb. 54
244.
(1916)
3315,
t.
Bubbia
-
pachyantha
Adansonia
Drimys pancheri Baill.,
-
Hook. Ic. PI. 34
sub
(1936)
t.
comb.
nov.
Bubbia
—
E.G.
Drimys pauciflora
—
pauciflora (E.G. Baker) Dandy,
72(1934)41.
Zygogynum
sub
nov.
1.
Zygogynum
J. Bot.
comb.
(Baill.) Burtt,
Bubbia pancheri
-
1,1985
428.
comb.
pancheri (Baill.) Vink,
(1873)
3315,p.
No.
31,
pachyanthum (A.C.Smith) Vink,
Zygogynum
10
VOL.
-
Bubbia
-
nov.
polyneura
—
Diels,
Drimys polyneura
Bot.
(Diels) Burtt, Hook. Ic. PI. 34(1936)
3.
p.
queenslandianum (Vink)
Zygogynum
comb.
comb.
Vink,
nov.
Bubbia
-
queenslandiana
Vink, Blumea 28 (1983) 318.
queenslandianum
Zygogynum
-Bubbia
Zygogynum
laumin,
queenslandiana
schlechteri
Ic.Pl.
34(1936)
Zygogynum
nov.
—
p.
Vink
A.C.Smith,
(F.Muell.)
comb.
(1983)
Bubbia schlechteri
—
nov.
322.
Guil-
nov.
Drimys semecarpoides
-
(F. Muell.) Burtt,
semecarpoides
Hook.
3.
(F. Muell.)
Bubbia whiteana
Burtt
var.
var.
J.
whiteanum (A.C.
Arn.
Arbor. 24
whiteana
Smith) Vink,
(1943)
145.
(A.C.Smith) Vink,
—
comb.
Bubbia
Blumea
28
325.
Zygogynum
54
3315,
t.
semecarpoides
semecarpoides
(1983)
sub
(1891)
nov.
comb.
Bubbia
—
(Vink) Vink,
Blumea 28
3.
(1942)
15.
Vink,
comb.
(F. Muell.) Vink,
semecarpoides
F. Muell., Vict. Nat. 8
subsp.
australe
subsp.
australis
(Guillaumin) Vink,
Bull. Soc. Bot. Fr. 89
Zygogynum
Vink
(Vink)
Vink
sororium
245.
(1916)
comb.
(Diels) Vink,
Bubbia sororia
-
nov.
(Diels)
—
Drimys
A.C.Smith,
sororia
J.
Arn.
Diels,
Bot.
Arbor.
23
Jahrb.
(1942)
427.
Zygogynum
sylvestre (A.C. Smith) Vink,
J. Arn. Arbor. 23
Zygogynum
comb.
nov.
—,Bubbia sylvestris
A.C.
Smith,
(1942) 430.
umbellatum
comb.
(Ridley) Vink,
Trans. Linn. Soc. Bot. 9
(1916)
11.
—
nov.
—
Drimys
Bubbia umbellata
umbellata
(Ridley) Dandy,
Ridley,
J. Bot.
72(1934)41.
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J.M.
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Epidermal
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CARLQUIST,
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1982.
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W.
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Bubbia
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C.A.
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Taxon
30:
591
—
600.
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C.G.
the
1944.
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SAMPSON,
The
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F. B.
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Pollen &
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Taxonomic
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W.
and
VI. Vascular
anatomy
of
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Pseudowintera,
8:
the
member
of
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New
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13: 403-423.
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J.
Arn.
Arbor.
119-164.
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1900.
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Homoxylees.
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Blumea
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1978.
Ibidem,
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1983.
Ibidem,
IV.
of the
Zygogynum
on
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I. Pseudowintera
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Drimys
-
morphology
225-354.
—
morphology
the ovary
The Australian
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of Bubbia.
Blumea
24: 521-525.
28:
311-328.