Download (Loranthaceae) Amyema and World, especially

BLUMEA
26
403-410
(1980)
A note on Heterophylly and Branching patterns
in
the Amyema complex
Job
of
University
(Loranthaceae)
Kuijt
Lethbridge,
Canada
Summary
in
Attention is drawn to the unusual distribution of flowers and inflorescences
to certain
far has
of branching and
peculiarities
escaped notice, involving
cataphylls.
A
similar
The genus
branching
Amyema
Ignoring
the
eight
or
dealing
with the
Danser
regarded
Amyema’.
several of these
complex
the
to
to
genera
in
Van
Sogerianthe
both
the system
the
but
(1931,
1933)
as
An
by Barlow).
been
context
encircling
in
wood)
between
peculiar
a
the stem and
words, inflorescences
feature,
1974;
a
of
including
which
internodes
Valeton
(
stem
to
also
are
for
more
very
Rubiaceae; Winkler
prolific
Amyema
creator
(1966,
1974).
Tetradyas,
Pa-
polymorphous
detailed investigations
a
was
in
the
has followed
not
nomenclaturally
for Amyema has
narrow
showing
of
more
and Rhizomonanthes are
even
to
regard.
workers
important
and Barlow
the latter genus
is
sense
an
several types which
at
a
inflorescence survey of Loranthaceae generally.
features of
Amyema
the
axillary region of
found
may be
a
an
found in those
1931).
lies in the fact that, in the
merely
in the axils of
zone
the
often
foliage
leaves. In other
(often profusely, especially
anatomical
The
like
the
curiosity
extreme
are
flowers
found
on
paid
in
expressions
species (Dicymanthes
leafy
completely
leaf pair. Little attention has been
constitutes
much
not
floriferous
transverse
(Merr.) Danser)where inflorescences
themselves,
the
most
regard
reliable generic limits and
develop endogenously.
inflorescence proliferation
and D. seriata
with
whole. A discussion of these types will follow
an
narrow,
on a
most
infrageneric organization
proposed.
the members of
inflorescences appear
constitute the
'extreme forms of the
more
so
intercalary
Helicanthes.
of Loranthaceae. Barlow's work
striking morphological
rather
of minute
Diplatia, Distrianthes,
great majority of species, the inflorescences emerge
organs
in
mostly by
two
diverse genus in its inflorescence structure,
One of the
heterophylly which
World, especially
need, however,
achieve
separate genera (Barlow
later date within the
number of species, and
a
pair
a
undoubtedly
Old
Tieghem,
Dicymanthes,
in Loranthaceae as
unique
is evident
generally although Dicymanthes
knowledge,
my
exceedingly
are
as
the
have been Danser
genera
overlooked
apparently
not,
names
He stressed the
Danser in these views
maintained
early disappearance of
relatives
nearest
a
explained by
nine additional genera proposed,
generic
placement
accurate
are
Much differenceof opinion has existed in this latter
Rhizomonanthes and
puanthes,
genus
its
and
The latter
flower distribution
and
pattern
and
generic circumscription.
of Loranthaceous
the formation
Loranthaceae in
complex of
difficult
phyllotaxy.
older
to
that
this
the
of such
suluana Danser
irregularly along
of Stichianthus
the
minutiflorus
404
BLUMEA
Plate
Sogerianthe versicolor,
I.
between
intercalary cataphylls
enlarged.
The
3.
Fig.
—
of young
prominence
lower
next
flower
Brass
3227,
two successive
zone.
node
Fig.
—
26,
L.
Fig.
—
of
pairs
from
4.
No.
VOL.
2,
1.
Young
leaves.
foliage
1,
Fig.
Emergence
1980
branch
from
Cataphyll
of
showing disappearance
of four flowers
of
tip showing position
2.
Fig.
—
Fig. 1,
and
cataphylls
from the flower
zone, all
in
approximately prophyllar positions.
In
number of
a
from
epicortical
endogenous
inflorescences
Amyema species
1
roots
In these
).
of flowers
origin
or
inflorescences,
number of quite
unrelated angiospermic
species
strictly
showing
endogenous origin,
is
nature
taken
referred
are
numerous
nor
both
genera
show the
species
buds
axillary
covered
the
covering
what
also,
all.
(As
axillary
endogeny
Old
it is very
as
appears
be
Worlds.
from internodes
there is
of these
plants.
and
but Hooker
as
In
a
It needs
The
implication
at
the
be
can
of all this
find
an
of
in
flaps
be
cannot
of it in
traces
of the
species
some
to
but
that
able
was
as
well
which the
cushion
hard, protective
that in
in fact
may
endogeny).
in the young
Amyema,
be this
not
understand how the
true
of
matting together
a
proper anatomical
a
show that
to
through
is, therefore,
nodes
flowering
anything
emphasis
we
tissue
to a
conspicuous hairy covering
no
such
for inflorescences
(1846)
caplike
a
axillary region, drying
endogeny
roots
usually
of
a
those
In Misodendrum (Misodendraceae), , several
mentioned above, however, it is difficult
enough,
in
by irregular flaps
phenomenon
a
general,
pronounced
secondarily by
over
in the
area.
to
Even
emergence. Such
bud eventually emerges. In Cladocolea there is evidence of
perhaps glandular hairs
from
known
evidence
some
ruptured during
and
New
phenomenon,
same
were
the
example,
(Kuijt 1975).
innovations
vegetative
been
in Loranthaceae in
rare
in
is
doubt of the
no
1969).
involved may be invested
This apparent
for
be
'chaffy scales' by Barlow (1974), but the implied leafy
to as
neotropical Cladocolea,
as
apparently
is it
there
(Kuijt
directly
may emerge
can
phenomenon
a
parasites
flowering
structures
appropriate.
not
face value,
at
nodal
the
which have
corky tissue,
Sogerianthe
as
flowers
or
latter instances there
at
flowering
Interestingly
flowering
study
zones
is needed
to
elucidate these features.
In the
course
of
general
evident that the overall
inadequately
explored;
relationships
to
understanding
complex,
present
studies of the inflorescence of Loranthaceaeit has become
structure
I
of the members of the
referring especially
am
the position of inflorescences.
of this
there is
no
doubt that it will
on
be
an
herbarium
key
My original attention
present
was
focussed
neither of which
seem
on
to
successive
than
')
at
See,
pairs
of
foliage
perpendicular planes
for
example,
A.
leaves
as
My
are
been
Rhizomonanthes)
also the inflorescences
of the above three
leaves.
and A.
can
a
Danser
seen
and
be rather
where two
misleading.
puzzling
the
we
features
literature. The
each
percurrent stem, however,
A.
in the
from
on
the
same
plants
hastifolium
Barlow. Flowers
both
when
which,
recorded in
expected
dilatipes
species) are
only
always aligned
would be
curvifolium (Krause)
Danser's
constituting
are
an
of the
material
branches of these plants frequently bifurcate. On
two
that
pretend
essential tool for this purpose.
Sogerianthe,
have
cannot
and their
systematic analysis
to
concerned with the position of branches particularly,
are
has been
Amyema complex
branching patterns
While I
is the
morphological aspect
contribution is based
to
in
plane
(Ridl.)
Danser
Distrianthes
epicortical
rather
with decussate
(together
(and perhaps
roots and in the axils of
Job
Kuijt:
Heterophylly
and
branching
patterns in
Amyema
405\I
406
BLUMEA
phyllotaxy (Plate I). Secondly,
from the
(often apparently
the
foliage leaves,
nearest
alternate with the
a
VOL.
often than
more
flowering zone)
in their axils
not
if
especially
somewhat
only
one
of the
2,
1980
not
the innovations
placed
are
In other
innovations
two
with
the
at a
node
obvious in
to
such branches
the members of
pressed material;
its orientation is often altered
develops,
internode
words,
between
always
not
arising
perpendicularly
foliage leaves, arising
nearest
aligned
be
to
No.
(Plate II).
As mentionedearlier, this feature is
pair.
26,
and
below,
it
may
be
to
appear
percurrent.
accounting for both
A solution
of
particularly good specimen
which has young
Islands, L)
that,
an
on a
of these aberrant features
in the process of
twigs
percurrent branch,
successive
two
with
presented itself
The fact is
elongating (Plate I).
of
pairs
foliage
leaves
separated by
are
extremely inconspicuous, caducous pair of intercalary cataphylls which
for
only
seen
a
very brief period. Each
and about 0.5
dorsally,
the
expanded
material in the
flowering
node before
cited above there
all in the
It
be
can
in
does,
leave
these
that
these
fact, exist but successive pairs
Innovations
The observed
exists in several
1959),
of
pairs
the
intercalary
novations, although
been
of the
first
material. I
at
or
a
few
P.
with the
calls
these
and
correct.
cataphyll
in
in
flowering
assuming
prophyllar,
all above the
I have
flowering
as
not
a
to
axils of
and
scarce
I
axillary positions
but in
some
observed any
In
seem
P.
as
Phoradendron,
to
subtend
in-
crassifolium (DC.)
intercalary cataphylls.
No
Sogerianthe.
is
zone
The
next
flowers
difficult
they
seem to
and such
be
to
foliage
appear
tempting
too
designation
a
regular pattern
to
It is
petiole (fig. 4).
some
petiolar base,
to
leaves.
foliage
parallel
Trel.
never
such
species
single flowering
the
same
far
to
observe
leaves
designate
the side for this
may therefore
in the further
in
the
certainly
are
to
are
not
development of
be
the
zone.
Danser did
same
in
mind the
(H.B.K.)
persistent,
mature.
common
of caducous
pair
a
nearest
that the flowers in the axils of
to
a
which
It is of considerable interest that another
the
single
(Kuijt
elaborate
immediately flanking
positions
not at
a
one
piperoides
are
more
least among the first
those found
along
genus Phoradendron of Viscaceae
immediately
common
in
am
In the collection
Regular decussate phyllotaxy
frequently
most
position
neotropical
cataphylls
seen
The sequence of
pressed
of the
swelling
leaves
intercalation of
the
Eichl. inflorescences regularly stand in the axils of
flowers have
fully
corky
leaves find themselves in the
foliage
originate
seem to
phenomenon
species
especially
are
leaves below
foliage
scars.
foliage
keeled
of the
cataphylls provide
intercalary
of
and thus alternate in
cataphylls,
however,
discernible
no
successive
ten
leaves
foliage
production
because of the considerable
plane (on percurrent branches) through
cataphylls.
the
be
can
scale,
plane.
therefore,
seen,
before the
for both of the above mentioned features.
explanation
brownish
minute,
between it and the node of the
cataphylls
least
at
are
same
a
fallen, possibly through
zone
the
flowering,
Even
long (fig. 2).
have
is
cataphyll
Both because of the cork and
(fig. 3).
shoot,
mm
cataphylls
a
versicolor Danser (Brass 3227, Solomon
Sogerianthe
not
consider
closely
branching pattern
cannot
but, certainly
crescent-shaped
a
of it is
to
(monotypic)
Sogerianthe
described above.
be certain that all
most
the members of
as
related
branching
(Plate III). Young
leaf-pair.
The first flower
leaf scar) is in
a
is
as
or
genus, Helicanthes, which
Amyema,
precisely
described for Sogerianthe above,
innovations arise in
(only
shows
Since material of Helicanthes is
a
scar
in
positions
fig 10, directly
primary axillary position,
but
between
above the
subsequent
ones
JOB Kuijt:
Sogerianthesogerensis.
Plate II.
Flower
positions
indicated
pattern,
branching
as seen
petiole,
those of
Sogerianthe.
2
),
one
branching
5. Cruciate
Fig.
—
black
dots.
Amyema
patterns
branching
in
pattern.
luzonensis.
—
—
not
6
Fig.
8.
Figs. 7,
Since flowers
supposed,
in
affinity
even
a
series of flowers
Helicanthes,
between the
if the
two
geographical
or
again
as seen
from
Variations
been able
to
ascertain the
scars
in
as
genera
in
above.
cruciate
very much like
Sogerianthe
might
not
distributionof the
and Eastern New Guinea and island groups to the east,
distinct. I have
407\II
Amyema
from above.
wonders if the
closer than Danser
(India,
by
and
and older nodes show
flank the
solitary
Heterophylly
occurrence
of
,
are
be much
two
respectively)
genera
is
quite
intercalary cataphylls
in
Helicanthes.
2
)
The statement The inflorescence
(Johri, Agrawal,
probably
referred
and
to
Garg
a
leafy
1957,
p.
branch
is
a raceme
352)
with
is not
a
with
about fifteen sessile
only self-contradictory
nodal
whorl
of flowers.
flowers
but
arranged
also in
error;
in
the
a
whorl'
authors
408\III
BLUMEA
VOL.
26,
No.
2,
1980
Job Kuijt:
While
all
pattern,
numerous
species
of
lateral branches in
Australian
the
be
of
for
in
this
Amyema species.
lack
cataphylls
type
there
This is
A. maidenii. As would be
There
quite separate.
Amyema complex,
the
thus
are
two
is,
In those
together.
occur
leaves.
are
My impression
with
normal, percurrent system,
a
decussating foliage
example
of 'cruciate'
Similarly,
cataphylls.
always
409
Amyema
have
to
appear
all
clearly
and
patterns
in
several
predicted
in such
seen
cushions from which inflorescences emerge
axillary
two
ramification in the
which
branching
the axils
confluent but
not
seen
for
true
Amyema
representatives,
species also,
are
not
branching
either we would thus expect
showing
not
Sogerianthe
this is
that cruciate
however,
species
of
species
branching
and
Heterophylly
at
node
a
distinct patterns of
very
of which needs
systematic significance
to
explored.
The
In
habit of
forking
the
of
majority
which forms
only
The entire
place.
aborts
we
which
are
many
this
cases
one
decussate
with the
is
effect,
area on
lateral innovations
only
a
single
and
leaves,
own
two
minute scales
in
placed
are
can
field
a
or
each
of
apex takes
internode long. Where
cushion that future flowers
hairy
develop,
before abortion of its
which sometimes
foliage
brownish hairs. It is from this
abortion of the shoot
apex.
early
on an
two
foliage leaves,
in
innovation,
very blunt
see a
that
means
of
pair
is based
species
an
be
apex
seen,
of short,
inflorescences
originate.
In
least
at
axils of the
of
species
one
leaves
foliage
the
fourinnovations is present above the
species,
such
even
on
the
same
inflorescences;
the
as
at
luzonensis,
cruciate
two
two
foliage
individual, is the
quartet of branches (fig.
a
these
Strictly speaking,
A.
Amyema,
may join
compound
an
apparent whorl of
Also
common
of four foliage
in this
leaves under
8).
any rate, the transition
with
normal innovations in the
Thus
(fig. 7).
occurrence
to one
is
not
already
be
considered
leafy
great. In several members of
inflorescences of this type have evolved,
Distrianthes, Diplatia,and Papuanthes.
Amyema species
leaves
lateral innovations may
Amyema complex, indeed, specialized
in
two
ones.
In turn, it is
possible
that the
umbels may have inflorescences of
derivation, involving the reduction of all leafy organs
on
such
a
numerous
comparable
branches.
ACKNOWLEDGEMENTS
I should like to
acknowledge working
from the National
Science and
facilities
at the
Research
Engineering
Rijksherbarium, Leiden,
Council,
and financial support
Canada.
REFERENCES
BARLOW,
B. A.
1966. A revision
of the Loranthaceae
of Australia and New Zealand. Austral.
J. Bot.
14:
421-499.
1974. A revision
Bot.
DANSER,
of the Loranthaceae
of New Guinea
and the South-western Pacific. Austral. J
22: 531-621.
B. H.
1931. The
Loranthaceae
of the Netherlands Indies. Bull. Jard. Bot.
Buitenzorg,
Ser.
3,
11
233-519.
Plate
III.
innovations
axillary
Helicanthes
(i);
flower
one
scar
elastica,
Wight 35,
leaf removed.
(s)
above
a
leaf
—
Fig.
scar,
L.
10.
and
—
Fig.
9.
Cruciate
Flowering
node
two flowers
(f).
branching, showing
showing
two
older
two
innovations
young
(i),
an
BLUMEA
410
.
1933. A
new
Old World
29:
Sect.,
HOOKER,
JOHRI, B.
J.
D.
M.,
system for the
species
JOB.
No.
genera of Loranthaceae
subfamily.
1946. Flora
Antarctica.
Verh. Kon.
Part II.
J. S. AGRAWAL, and S. GARG.
1959.
A
Phoradendron
.
26,
1980
2,
Loranthoideae, with
Akad. Wetens.
a
nomenclator for the
Amsterdam, Afd. Natuurk.,
2d
1-128.
Loranthaceae.-I.
KUIJT,
of this
VOL.
1969. The
Helicanthes
study
of
elastica
of
Loranthaceae,
pp.
Morphological
(Desr.)
heterophylly
(Loranthaceae).
biology
1957.
and
Danser.
8:
—
302.
embryological studies
Phytomorphology
inflorescence
Acta Bot. Neerl.
289
and
506
—
parasitic flowering plants. University
structure-
in
7: 336
—
in the
family
354.
Dendrophthora
and
546.
of California
Press, Berkeley
and Los
Angeles.
.
1975. The
WINKLER,
H.
genus Cladocolea
1931. Ueber die
Valeton.
Planta
13:
(Loranthaceae).
J. Arn. Arb.
eigenartigeStellungderBlUten
85—101.
56: 265
—
335.
bei der Rubiacee Stichianthus minutiflorus