Acta Bot. Neerl.
The
23(4), August 1974,
structure
primitive
a
and
function
Angiosperm
of
the
flower
–
*
discussion
Gerhard
Gottsberger
Departamento de Botanica,
de Sao
p. 461-471.
Faculdade
Medicas
de Ciencias
e
Biologicas
de
Estado
Botucatu,
Paulo, Brazil
SUMMARY
and
Morphological
discussed
Evidence
is
put
primitive
most
in
It is believed
dense
of
they
have
modes
of
of
or
to
on
large, solitary
and
still
come
seem to
to
of beetle
birds
numerical
suggested?
are
over
a
have
dominant
from the
mammals, since
and
with
pollination
regarding
changing
the
we
try
folded
carpels
(p. 335)?
Magnoliales,
directions within the
and
during the
during
saurochory
later
of
recent
as
in
Degeneria really
(1969)
dispersal types
He
has
is
were
convinced that
caused flower
dispersal syndromes
sperms. The purpose of the
present paper
Dedicated
the
Angiosperm
the
to Prof. Dr. L.
most
van
Is the
in
dispersal
more
modern
dominance
der
primitive
Pul,
flower
of
the
Den
are
states
Besides
in
this,
also have caused
structure
of
therefore is
living
to
is
stamens
generally
students of
perform
cantharophily,
specialization
flowering
“...that floral
primarily
ones to
in
leafy
as
so
studies
divergent
the
modes of
divergent adaptaprimitive Angio-
show what
in honour of his 70th
on
frequently
most
nowadays
flower structures, and what other
Haag,
struc-
floral axis
long
primitive
as
in mind when he
in the flower
*
flower
and their
archaic condi-
to the
Mesozoic
years.
beetle-pollination syndrome.
changing
as
the
whereas
group,
macrosporophylls prototypic
tions and correlated changes
regarded
carpels,
derive all other types from it? Are flat
to
mechanisms and
in
pollinated
were
the more effective
specialization,
maintained
insect
archaic
primitiveness
somewhat
What has Carlquist
anatomy”
observed
this
increase of stamens and
anatomy would have been served better if those who
be
not
flower
have occurred.
still
with its many micro- and
conduplicate,
can
are
lateral
end.
ideas
be
beetles
flowers
they gradually become adapted to
pollination. Together
inflorescences,
because
an
and should
dispersal
grouped into
unspecialized Angiosperm
did
phase
forced to switch
largely
dispersal by
Magnolia
floral
flowers
terminally-borne
ones,
are
differentiation.
early Angiosperm
pollination, many primitive Angiosperms
been
had
plants,
that
flowers
primitive entomophilous Angiosperm
aggregation and flattening might
prevailing
tures
of
conceptions
but rather middle-sized
primitive,
a later
of
cantharophily,
reptiles
The
most
Only in
reduction
regard
tion
to show
Angiosperms,
devastating type
more
enlargment,
In
that
beetles.
casuallyby
more
forward
the
features
with modem
or inflorescences.
aggregates
but
functional
and confronted
birthday,
struc-
462
still
although
tures,
considered
as
often believed
very
New
of the
Lhinfellner
from which
1956a,
a
ment
a
a
The
1969).
stamens
ventral blade
carpel primordia
as
Winteraceae,
families of this
Magnoliaceae
of
carpel ontogeny. Only
is
the
unifacial form
conduplicate folding
The
exhibited
by
the
carpels.
is
and besides
macrosporophylls.
it is
xylem)
and the
1968).
Based
have
now
broad and flattened
is
basically
wrong
Carlquist
reasons to
stamens
(For
textbooks where the
and laminar
Cronquist
of
and
1968 and
to
be
carpel
also very
are
are
(with
the
primi-
most
wood
primitive
very
(without
highest
poly-
Ehrendorfer
primitive Angiosperms:
Ehrendorfer et al.
conduplicate carpels
facts
new
flowers
still finds
are
believe
(1968)
ancient
bifurcation,
(p. 349-350).
Ehrendorfer
primitive Angiosperms
see
the
expression
described
folded
in
opinion
newer
conduplicate,
others).
of
believe that the still dominant idea of originally
and
primitive
stamens
30th edition of the
that
carpels
obviously represent a very
details of this
1969). Neglect
car-
manifested within this
from somewhat Winteraceae-like progenitors”
good
and
seems
variety
of Winteraceae
and
the
are
cytological findings
and Laurales “...
possibly emanating
We
their
on
Wintera)
sect.
stamens
therefore,
already
high paleopolyploidy
et
Magnoliales
early stages
pronounced flattening
much of the
stamens
stamens
chromosome numbers yet known in
al.
Drimys
other
some
the
pel-
primitive
great similarity between the micro- and the
only
not
The
ploid
that
is
flowering plants
tive in Winteraceae. Other characteristics
any vessels in the
less
Angiosperms,
beyond that,
this there is
But
in
structure
121-123).
1969:
such
and in
-
during
develop-
found
(1969)
including
fully developed
a more or
Similar-
before
stage
Degeneriaceae
in the
peltate primordium
Leinfellner
-
(Baum 1949;
be observed.
to
construction found in other
primitive,
and
a
be
members
most
congenitally.
peltate
Magnoliales
families
Winteraceae. But
family (Leinfellner
a
of
structure
with
unifacial tendencies
even
primitive
primitive carpel
most
terete)
rather
be confronted
to
connected.
are
carpels
in the Winteraceae (especially in
maintained,
whereas in all other
pels,
or
observed
order he
both
and fuse
through
pass
will have
and
start
develop
tate-utriculatecarpels in 14 families of the
ones
extent
stamens
unifacial (i.e.
normal form of utriculate
the
to
1956b,
dorsal and
the unifacial
basically
have
Magnoliales
should
primitive,
structures
what
to see to
have shown that the
investigations
be
to
less derived. Flower
more or
with functionalaspects in order
ly,
GOTTSBERGER
G.
having
as
discussion
in
most
basically
carpels (Takhtajan
(1971),
the
on
other
large
1959,1969;
in the
hand,
textbook takes into consideration the
“Strasburger”
of
modern
primi-
tive characters of the Winteraceae.
The
primitive
primitive
and
possibly
unidirectional
(1961).
structure
of the
to
Bailey and
case, Winteraceae
floral
characteristics
Winteraceae
was
considers the circular
than
encountered
others,
might
might
in
be
for
of
expected
to
hint
xylem
reasons
Magnoliaceae,
Degeneriaceae” (Carlquist 1969:338).
in
Winteraceae
correlated characters. “Trends
according
In that
primitive
wood
cited
have
to
other
more
numerous
Himantandraceae,
their
placentation:
in
Drimys (Sect.
are
by Carlquist
Another primitive feature to be
subapical placenta
more
evolution
Leinfellner
Wintera)
as
or
expected
(1966)
the
most
THE
AND
STRUCTURE
type found in
primitive
OF
FUNCTION
THE
PRIMITIVE ANGIOSPERM FLOWER
Further
Angiosperms.
primitive
A
-
463
DISCUSSION
features of Winteraceae
will be discussed later.
Since
(1916)
Diels
“Ranales”
as
attention has been
been made
Daumann
Heiser
by
demonstrated the
phylogenetical
a
paid
and Grant
for
(1962)
and
van
der
for
Pul
van
Pul
der
(1960,
are
studies
Recent
revealed
the
on
that this
winteraceous
pollination syndrome.
from the
ly
less
and there is
no
floral axis.
and
perianth
and others
were
-
any
flowers
From
Drimys
flowers
gentle
All
and do
other
species
are
time
same
visited and
Heiser 1962:
still
262).
lacking,
we
-
large
preted
an
“Flowers
of
in
the
during
has
beetle-
a
the
by
some
along
of
like
our
the
rather
for
to
us
learn that
Diptera, Thysanoptera
the flowers. The beetles
attack,
anthesis,
white colour of the
surprising
most
but also
whole
The unifacial micro- and
spiral position
of the
gnawed
their
were
we
the
eating
carpels.
petals,
observations
too, but that
attract
more
a
Hardly
stamens,
conclude
cantharophily
the
often
flowers
to
is
or
that
of
Drimys
cantharophily
by offering
in
certain
visitors.
specialized
considerable
elongated
are
damage
very
most
adaptation
are
and the venation of all
gigantism” (Carlquist
to
better
probably
parts
1969:
may
340).
In
a
are
ways
against
or
of the fact
and other
otherwise
all
from
and
large
1904: 303,
closely arranged
thorough
studies
Magnoliaceae species
specialized
damage
the
Magnolia
beetles.
number of sexual
organs could be
escape
be
(Knuth
with many and
with
nutritious
some
The flowers of
the flowers
spite
Magnolia
attracting fruit-eating
large
to
large
floral axis.
believe that
Magnolia
specialized
more
beetles,
protecting
flower dimensions and
as
open
type. The small beetles attracted
The flowers
the
function by deceit
the
Magnoliales,
Brazil
progress)
principally pollinated by small, medium-sized
cause
may
sporophylls along
are
most
stigmatic portion
studied have
harmful side effects of the
beetles which
from
in
harm the flowers.
not
Magnoliales
the
at
was
result
cantharophilous
specialized
(1970)
(1950a,
Drimys brasiliensis differs, however,
and
to
of insect
signs
(1940),
Pervukhina
Ehrendorfer,
be attracted
It
the
across
significant
are
of
beetles,
all harmful
mechanisms which
particular
tissues,
this
less
“open”,
at
passed
showed
carpels.
more
not
and
pollen-tetrads
small
mostly
-
(1962),
brasiliensis
Drimys
remain
to
seem
flower odour.
a sweet
the flower visitors
Heiser
and
(1930)
Corner
and Gottsberger
of the interior flower
parts.
The insects
Eupomatiaceae,
Daumann
Magnoliales investigated by being apparent-
The flowers
grouped loosely
are
(1960),
and
like
species,
cantharophily
protection
for the
Uphof (1933),
(1960),
the
increasing
textbooks.
of
biology
of all other
specialized.
macrosporophylls
short
The
cantharophily
much
flower
most
Silberbauer-Gottsberger
(Gottsberger,
group,
found in the
papers of Grant
Leppik
1961),
in
cantharophily
this
studies. Contributions have
(1958)
(1910),
Kral
(1953),
(1967), Carlquist (1969), etc., and in
of
in
Calycanthaceae,
Wester
for Annonaceae. General discussions
1950b),
pollination
Hotchkiss
(1950a)
Magnoliaceae,
Zimmerman (1941),
importance
primitiveness
the results of
to
Hamilton (1897)
(1930)
of
sign
by
crude
examples
of increase in
suspected
of
having
size
Here,
inter-
pollinators.
phyletically,
increased
with this
464
Another example
of
gigantic
“
family Nymphaeaceae.
food
its
bodies)
Pul
large
flowers and
large pollinators
amazonica
Victoria
dynastid
of which
feeding-cycle
not
understood”
yet
The
another
Annonaceae,
(Gottsberger
way
heads. When the flower
flower
organs and
that
about
bring
which
petals
are
carrion,
odours
by
highly
The
borne
are
ovule
in
to
the
beetles still
gentle
The
Annonaceae possess
by
deceit: The beetles
they
seems to
possesses
same
as
the
carpels
and ovules
of
the
be another
and
of
the
special
petals.
feeding
This
on
Calycanthaceae,
food bodies
A nitidulid
on
1904:
(Knuth
1897:
51)
287), probably
attract
studied the
eaten
is
the
to
accom-
where the
successful
carpels
method of
studied
thoroughly
the
beetle, Colopterus
are
tips
truncatus,
pollinating
by
of the
was
the flowers
are
The
mostly
flower. Because of this
selective pressure
sporophylls
stamens
structure
up
and
and
to
to
very
more
to
we
by
flatten
a
sweet
pollination
to
be of
primitive
maintained
as
an
pleasant
odour and
sexual organs of the
we
increase
have therefore maintained their
fully developed stage.
a
stu-
cases.
because of its unspe-
harm the
or
to
are
Victoria
Eupomatia
more
think that there
the
aggregate them densely along the floral axis.
carpels
the
mode of
to
when
detailed
reveal many
carpels
by
not
294),
and
deceit. More
consider it
and
attracted
small and do
primitive
stamens
the beetles
1950a;
(Grant
will
probably
stamens
enlarge,
have flat
Magnoliaceae,
many
of Drimys
beetles,
by
eaten
(Hamilton 1897).
their flower visitors
Magnoliales
pollination
cantharophily.
by deceit,
that the
learn
to
decaying fruits,
staminodes,
archaic feature in the flowers of this winteraceous genus
no
-
being
Eupomatia laurina, Eupomatiaceae,
unifacial
type. We believe that the
cialized
and thick
characteristic of certain
these nutrient organs and
the floral chamber
to
flower odours in
Having
the sexual
attracted
are
Eupomatia,
highly significant
in Annonaceae, Calycanthus occidentalis
(Hamilton
from
floral axis.
flatter inner petaloid staminodes which
amazonica
on
to
time.
force their way
Just
thereby
essential
was
those of their normal substrate
excavation
the flowers
even
It
pollinators.
the
soft,
are
large
proceed
can
stigmatic
become
harmful beetles
carpels
protected
hard
having
function
protect
The curious flowers of
and
and
fascinating
and
further
stigmas
the
and the innermost
found in
sometimes
by
occidentalis
(1950a),
stamens
cial
der
(Grant 1950b).
Calycanthus
the
van
cantharophily,
stamens
are
devoured
imitating
an
protection
beetle flowers
Grant
stamens
and the
receptive
petals. Large
more
specialized
have flowers with
plished by Xylopia (Annonaceae), Calycanthus
was
(with
normal
dung.
or
Another way
not
&
(Faegri
beetles in another
visiting
carpels by
be
pollination.
Annonaceae flowers
flowers
the
by
while smaller and
excluded,
nutrient
starts to
closed
is
entrance
from
floral axis.
the
on
connective shields, and the
by large
with
family
damage
1970). Many species
densely aggregated
most
dies
and feeds
castanea, the
pollinator, Cyclocephala
is, however,
be found in the
can
imprisons,
attracts,
1971:116).
protect their flowers against
at
GOTTSBERGER
G.
number
of
It may be
primitive
unifa-
THE
STRUCTURE
AND
FUNCTION
OF
within the
secondary
beetle
laminar stamens
massive,
The
of
flattening
and
aggregation
as
sporophylls
phylls
from the
the
most
sporophylls
because
many
(1952) adopts
have
flattened
derived
structures
for
by specialization
caused
was
the
interprets
in
chance
to
save
dense
by
part
against gnawing.
Flowers
of the
some
beetles and this may have caused the
gnawing
465
DISCUSSION
view and
opposite
protection
better
a
A
-
primitive.
presumably
of better
should
the
FLOWER
caused
Magnoliales, probably
Canright
pollination.
ANGIOSPERM
who considers
(1969)
We agree with Carlquist
and
PRIMITIVE
THE
with
sporo-
sporophylls
to
increase in number and in size.
Attracting
primitive
fruit beetles
just
flowers,
compensate the
to
It is
in this
deceit
by
side
negative
direction that
we
have been
must
nutritious tissues
offering
as
effects of
have
Annonaceae, Calycanthaceae,
noliaceae,
large
secondary
of
structures
precise pollinators.
more
and
acquisition
other special
or
and
larger
the
to see
a
solitary
flowers of
Himantandraceae,
Mag-
Nymphaeaceae,
and others.
the
most
primitive
the flattened
stamens
pollination,
to
laminar
radial
some
found that
(1969)
organs
three-dimensial
in view of the
type of
open
questions
of
to
structure
the basic type in
(1969),
Heel
in
stamens
stamens
we
do
that the
and
like
to
somewhat
Takhtajan
stamen
with
trends in
that led
to
propose
(1969:
some
are
the
time
schemes
with
we
on
a
of
of
structures.
some
In the
sta-
of Heinsbroek &
evolutionary
considering
van
pathways
the data
on
the
a
opinion
Magnoliaceae-like
of
given
functional
as-
Magnoliaceae
occupied
explain
related
stamen
evolution in
the
Cronquist(1968: 89)
truncate
unifacial
was
Degeneriaceae.
the
Drimys- like
beetle
cases
Degeneria.
diverse
Magnoliaceae
main
Mag-
pollination,
this resulted in
After the flatten-
positions,
difficulty why
We
two
trend within the
specialized
and
most
without
Angio-
shown in
most extreme
the adaxial surface in the
closely
It is
the telo-
the
may admit that there existed
evolution. One
abaxial. This could
situated
Contrasting
or
problem. Relying
related groups, connected with
flattening
the pollensacs
gia
same
Starting
stamen
stamens
processes
surface in
the
the
regards
explanation,
should have
stamen
findings
tentative scheme of
a
57).
ing
or
at
possibly
flattened,
so.
the blade-like
adaxial,
263)
apical pollensac-groups
the
with the
unsolvable
an
structure.
Heel
simple
harmful beetles.
132-135), defending
Degeneriaceae-
different from the
Angiosperm
noliales and
as
Leinfellner and
think
“typical”
The
functional
a
by large
primitive Angiosperm
flat
(1972:
Kubitzki
not
We would
sperms,
could find
respect
van
the
not
homology” (p. 443).
pollinated
Angiosperms
Angiosperms
Baum and
by
pects,
Victoria amazonica “...are
flower is
about the
(1952)
in
&
-
non-flattenedbut otherwise rather
of Canright
considering
but rather flattened three-dimensial
be,
of these
me-theory, already argued
and
specialized
as
be settled. Heinsbroek
structures, of unknown
fact that the
Angiosperms
in this respect that Zimmermann(1965;
interesting
mens as
can
and other Winteraceae
Drimys
in
primitive Angiosperms
stamens
appear
-
of
stamen
microsporophyll
of other
the
they
unifacial
or
unifacial
truncate
with the
Starting
as
the
but
have, however,
marginal,
microsporan-
on
to
the abaxial
admit that
466
an
is
G.
of
apical position
The
has
abaxial
1969:
what
question
sperms
caused
well
as
microsporangia
as
57-59) suggested
adaxial,
and
abaxial,
like
not a
Drimys
1971:
we
-
obtain
that another trend in
with
unifacial
insects
and anther. We may
more
advanced than
of the sterile basal tissue of the
truncate
producing
while the
remained
with
typical
a
more
or
filament,
less
that
also
(see
of
flattening
when
of the
Calycanthaceae,
principles
and
large
many
after
hand,
some
a
said,
and
Degeneriaceae,
Eupomatiaceae,
data make
solitary
cences
by
flowers
resent
many
were
in
the
in
Magnoliaceae, Degeneriaceae,
authors,
characterised
(p. 55). Why
too,
But he did
when
he
unwilling
solitary,
minds of
A
the
as
flower became
large
in
flower
at-
protected
mind,
one
was
Himantandraceae,
and others.
to
from
Starting
be
(1945),
regarded
on
solitary
the
as
other
flower found
to
be derived
family.
have
cases
occurred.
of
Ims
(1964,
Himantandraceae,
to
and
not
stated
flowers
appear derived
74, 78, 79)
Himantandraceae
fully recognize
that
fact, just
replike
Angiosperms
leafy
branches...”
Angiosperms
on
the
solitary
probably
this
“...the earliest
terminal
from inflores-
noted that the
accept the idea that the earliest
by large solitary
are we
the
Magnoliales
reduction. Also Takhtajan (1969:
other
large,
a
time better
Magnolia,
probably
assumes
pollination
Having this
within this
discussed
structured
and Annonaceae and maintained that reliable
flowers
reduced inflorescences.
Angiosperms
secondary phenomenon
primitive
165)
as
Winteraceae had
In other families similar reduction series
1967:
in accordance
the
radially
visits.
same
Bailey & Nast
a
already
occurred, finally
solitary
concluded that the terminal
members of the Winteraceae is
after Pervukhina,
and
Nymphaeaceae,
in many
as
of
argue
stamen
constriction
or
are
of beetle
the
at
flowers
Magnoliales.
thorough study
is
may
and later on.
flower
large
pollinators.
solitary
from the inflorescences otherwise
cit.
the
apical
Ehrendorfer
somewhat fixed in the
about accidental
Annonaceae,
in
only partly
exceptions,
talking
with
not
accept
pollensac-groups,
Our ideas
general recognition
was
inflorescences
derived features
more
it
the
199). Ehrendorfer also
133,
Angiosperm flowers,
we
“typical”
has
the
believes that
primitive Angiosperm
with few
If
that in flowers
parts,
Pteridosperms
occurred
the crude side effect of the
thinking
in
is,
the beetles better,
against
such
in
requisite
necessary
most
botanists. After
primitive syndrome
tracts
probably
stamens
structure
contemporary
who
Zimmermann 1965:
The concept of the
and terminal
fertile
still
was
we
reduction
a
microsporophylls
664)
some
also
the
to
assume
beetles,
original position.
(1971:
from
originated directly
sporophylls
a
in their
those of Ehrendorfer
may have
evolution led
an
(1959: 84,
living members,
again
stamen
Angio-
89-90)
fourth,
a
in archaic
interpretation (see
truncate
stamen
Angiosperm
filament, connective,
pollinated by
but
marginal.
derived from
as
clearly present
in
(1968;
Takhtajan
from the
stamen
been derived.
primitive
most
opinion
primitive.
position
much clearer
a
with the
634). Starting
but
one,
truncate
have
marginal original position,
laminal
marginal positions
hypothetical
is the
may be
position
the
somewhat
positions
Cronquist’s
In
possibly
a
deriving
position
radial,
a
pollensacs
dispute.
adaxial
an
of
position
some
convinced about
-
on
condition from which the other
primitive
the
GOTTSBERGER
THE
AND
STRUCTURE
could have
had
FUNCTION
OF
THE
inflorescences
terminal flowers, and that the
be better
interpreted
beetle pollination
studied
genera
beetle
nant
has
pollination
of
have
its
least
flower
flowers of
Magnoliales
already
While the
feature, Drimys
ANGIOSPERM
FLOWER
with
less
flower in these
flowers
specialized
aggregated
flowers
Have
the
at
end
solitary
not
forgotten
and
less
of the
the
that all
specialized
groups is indeed
and
could
generalized
we
flowers
highly specialized
solitary
and
living Magnoliales
much and
too
467
A DISCUSSION
-
aggregates
most
derived from inflorescences?
as
far
so
at
or
solitary
syndrome
pollination?
PRIMITIVE
domi-
a
beetle
specialized
branches in
cymose
inflorescences.
What about the evidence for
flower?
lateral
primitive inflorescences.
have lateral
(in part)
matiaceae
lateral
between the
lateral
conclude that
or
have
primitive Angiosperms
flower, although
archaic
later
probably
Angiosperm
acquisition
of the
the
similarity
Magnolia?
it,
by
The size of
is also
at
all
significant.
noliaceae,
unspecialized
with the
comparable
of the
Degeneriaceae,
have robust and
most
with
cences
Such
or
solitary
solitary
flowers
or
in
respect
flowers which
flowers
and is also found in its
position
of
solitary
flowers is
only
beetle
pollination,
as
in
Thysanoptera) probably
sized
sweet-smelling
gentle (as
in
Drimys),
Drimys
were
not
a
realized
the
pollination
like
not
Mag-
Eupomatiaceae,
beetle
pollina-
Some
possess
Magnoliales,
which
their derivation from inflorescences.
size.
larger
in
The lateral
Magnoliales
by
some
started with
more
or
beetles
(later
less
were
very harmful. To
by
on
also
chance
by
one
The terminal
flowers.
unspecialized
Diptera
and
these middle-
anthophilous,
improve
of
position
than the terminal
large, specialized
The small beetles
These
following:
pollination
have somewhat reduced inflores-
Angiosperms
attracted
flowers.
and
:
a
con-
and the
Cycas
primitive representatives.
most
It is therefore to be assumed that
beetle
flower aggregates.
common
probably
without
families,
specialized
unspecialized
or
attain
more
the
as
perhaps
beetle
may conclude the
we
with
still show
normally
inflorescences is
less
seen
middle-sized and
are
Calycanthaceae,
more or
pollination,
to
specialized
living
Magnolia
not
obsessed,
to
reason
most
still
strobili of
Drimys
distin-
apparently
the
large
is
who
81)
flowers.
primitive Angiosperms
specialized
more
the
Eupo-
not
and the
flower. All other
Himantandraceae,
mostly larger
still
we
flowers of
did
terminal flower
terminal female
middle-sized flowers in inflorescences
are
are
(1969:
although
because
or
predominantly
What is
derived structure,
Or
all facts mentioned together
Bringing
of the
it
truly
large Magnolia
Annonaceae, and Nymphaeaceae,
tion,
Is
flowers with and without
primitive
The
Is the
Angiosperms?
terminal flower of
fessing
somewhat
type?
Magnolia
had terminal flowers,
lateral ones?
a
flower
inflorescences.
Takhtajan
Eupomatia.
flowers
and Himantandra
Eupomatiaceae,
terminal flowers of
inflorescences of
Angiosperm
solitary
and Annonaceae have
inflorescences.
original Angiosperms
of the
lateral
few-flowered
or
16)
the flowers of
to
apparently
flowers
solitary
flowers
position
have
Drimys,
Magnoliaceae, Degeneria
1969, fig.
few-flowered
or
terminal
a
terminal position
original
Some
solitary
Takhtajan
(see
flowers
attributed
guish
an
Many Winteraceae, including
relatively
pollination,
lateral
468
G.
flowers
solitary
primitive
er
They
ways.
built
utilize
to
started
were
better
was
enlarged.
At
One flower
probably
adapted
The
good
the
or
less
same
few
a
gave
to
reasons
specialized,
by
Xylopia,
early
are
on
land and
beetles
did many
It
flowers
were
reduced.
more
precisely
middle-sized flowers
beetle
may
doubt
beetle
as
regularly
a
birds
cantharophily
on
birds
by
a
flowers.
are
mammals?
or
in
pollination
phylogenetic
eaten
from
while their seeds
as
groups
cast
with
interpreted
pollination
unspecialized
by beetles,
animal
as
an
Magnolia,
“reminiscence”
mammals and
or
have very much influence on the
not
der
Pul(1966: 603-614,
are
1969a:
was
discussed
which
extensively
following
situation.
reptiles
but the
some
we
by
often
tropical,
number of reptiles still occur, as, for example
humid
Most
still
van
some
saurochory
where
basin,
of
dispersed by
are
regions,
in the Amazon
primitive
dispersal
majority
other agents. The archaic condition of
in
the
88-89,1969b: 22-24).
them still show links with herbivorous
preserved
origin
played
fruit and seed
most common
beetles. In their fruit and seed
pollinated by
birds, mammals, and
reptiles
dispersal by reptiles (saurochory),
therefore confronted with the
Angiosperms
the
were
primitive Angiosperms,
find in modern
At that time
Angiosperms.
presumably
This archaic
dispersal agents.
can
commonly
inflorescences
specialized
Drimys
did
certainly
differentiation of the
are
primitive Angiosperm
these “modern” animals?
dominant role
We
seeds
be
to
casual visitors.
regard
we
and
Guatteria,
Birds and mammals
and
even
do
Why
when their fruticles and
dispersed by
modern
such
in number
and crude visitors.
flower therefore can be
through
contradiction which
phenomenon?
Talauma,
resulted
or
than
more
solitary
offered breed-
flattened in order
inflorescence with middle-sized and
or
dispersed
not a
archaic
results
primitive Angiosperms pollinated
are
commonly
Is that
that
Lat-
dung-beetles;
or
carpels increased
were
attracting specialized
small and
terminal
having
as
flower aggregate
Why
flowers
carrion-
specialized
more
specialization
pollination
or
stamens
time flower aggregates
large
better
lateral
large
and
forming
in the most diverse
precise attraction; they
more
by
branches,
Angiosperm history.
syndromes
the visitors. Stamens and
with this
probably parallel
of the
in
deceit, attracting fruit-,
destruction
protected against
end
their pollination
densely aggregated
more
the
at
Drimys) quite early
nutritious tissues for
up
ing places and/or trapped
and
aggregated
in
(as
Angiosperms specialized
one
they
have
might
inflorescences
GOTTSBERGER
the
is
a
more
great
Guianas,
Borneo, the Everglades of Florida, and the famous Galapagos Islands.
With the
zoic
the
rapid development
high
therefore
Angiosperms
“modern” modes of
brought
about
tion for
dispersal by
33),
but
known
In
as
a
should
the
of
time
were
dispersal
divergence
to
came
in order
to
between the
have
plastic
such
a
caused
and
too
end.
an
forced
largely
to
survive.
switch
This
(van
der
the end of the Mesooverall
over
Pul
was
not
1969a:
for
the
dominating
other,
to
more recent
great problems
adaptive higher plant
switch-over
at
The
reptile-dispersal syndrome
birds and mammals
not
most
pollination
of birds and mammals
reptiles
more
switch-over
and
88-89,
adapta1969b;
Angiosperms,
group.
necessary.
The
beetles
were
the
THE
STRUCTURE
AND
dominant insect
still the
condition of
whereas in
group
does
saurochory
find
one
maintained
becomes
ago
This is
is
maintained
still
or
evident
discussion
we
born
middle-sized,
and
few
relatively
axis;
and the
ting
its
of
the
unspecialized
We
pollinated by
in
major
any
has main-
genus
mode of
of birds
morphological
in
primitive
early
very
Angiosperms:
on
in
not
a
situated
laterally
Angiosperm
flowers
inflorescences;
with
acyclic
very
floral
long
apical
in
long
and mammals.
the
tentatively
most
have
dispersal
themselves
reptiles
groups
which
regions
archaic
that
microsporangia
are
totality. Only sporadically
branches;
lateral
both of them
carpels utriculate,
wide spectrum
a
in
carpels loosely arranged
with
truncate
in fact
the archaic
more
any
single
a
in undisturbed
possibly
of the
primitive
and
stamens
stamens
even
may summarize
are
and
why
Angiosperms.
many
dominant
not
modern
more
end
the
at
also formation of
history
so
reason
unprotected, hermaphrodite, protogynic;
open,
aggregated
by
469
DISCUSSION
nearly exclusively
the fact
considering
and functional characteristics which
Flowers
the
The relict character of this
the
our
not
species-groups
widely replaced by
Concluding
is
and
A
-
Angiosperms
probably
dispersal syndrome
species
more
of
origin
today.
primitive family
saurochory.
even
were
ANOIOSPERM FLOWER
primitive Angiosperms
its
as
some
PRIMITIVE
dispersal, saurochory
taxonomic group. No
tained
THE
the time of
cantharophily
find whole families of
beetles,
at
group
insect
largest
OF
FUNCTION
position,
unifacial; flowers unspecialized,
beetles; dispersal principally
attrac-
by
sauro-
chory.
After the first
protogynic,
sexual
the
er
acyclic
organs
lateral
carrion-beetles
dies
are
or
by
petals,
mechanisms
by
flowers offer
deceit;
like
by
by
the
closing
sometimes
carpels
rochory,
We
the
or
the
mostly utilizing
aminoid
more
side
effects of
which
help
to
or
production
by
exclude the
apical,
but
floral
to
axis;
conduplicate;
Prof. Dr. F.
over to
very
stamens
or
dispersal
less
still
dispersal by
Ehrendorfer, Vienna,
bo-
shield,
or
large beetles;
the
normal
densely grouped
flattened with
lateral
crude
food
imitation of the
more or
mostly adaxial, abaxial,
and
mostly switching
thankful
manuscript.
numerous stamens are
and
of
connective
a
upon
fruit-
attracting
specialized
more
the
borne in
acting directly
common,
and
number of
solitary,
or
odours
odours
larger flowers, by
elongated
numerous
but later
are
robust;
and
protection, alimentation, breeding places,
still somewhat
sacs;
hermaphrodite
enlarged
semi-inferior ovary,
a
substrate of the beetles; the
along
primitive entomophilous
but
flowers
fruity
compensated
thick
in
increased; flowers in reduced inflorescences
of visitors:
instincts
picture
somewhat: Flowers still
hemicyclic,
or
the
specialization
terminal position;
or
beetles
of
phase
presumably changed
Angiosperms
position
sometimes
of
principally
pollen-
by
sau-
birds and mammals.
for
a
critical revision of
470
GOTTSBERGER
G.
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