Acta Bot.
Need.
Facts
21(2), April 1972,
and
fiction
morphology
Hugo
in
with
floral
special
Polycarpicae.
to the
A.D.J.
113-127.
p.
1.
A
reference
general
survey
Meeuse
de Vries-laboratorium
Amsterdam
Botanicus,
Hortus
en
SUMMARY
The
of
possibility
demonstrated.
ceae, and
anthocorm
an
of ancient
unisexual
ambisexual
to
with
and
anemophily
some
of the
acting
upon
to
one
of
from the
groups of the
of
primary types
coaxial
associated
the
change-over
independent
and
of anthocormoid
and
structures
in
characterised
entomophily
in
from
parallel
gynoclads,
types originated
condition
with
is
concept
Polycarpicae) represent
androclads
The two ambisexual
gymnospermous
monocliny
anthocorm
primitive Piperales, Euptelea-
parts thereof,
or
separate,
the
on
of the
of a number
of the three
gonocladial androgyny.
forms
principal
the result
with
ones
partial switch-over
based
region
structures
anthocormoid
ambisexual
with
tion
of
floral
reproductive region
‘flower’ is shown to be
ones,
ones
the
a
tendencies
evolutionary
viz.
that the
Cercidiphyllaceae (and
surviving stages
of the
interpretation
an
By assuming
by dicliny
and
pre-
and
connec-
proto-
angiospermous groups.
A
‘flower’
true
tions
variously
phylls, and/or
sources
is
compounded of
always
associated
with the
with
other
supporting anthocorm
is not at variance
with
this
‘carpels’
this paper
groups and
derivatives
of several
In the third
the
are
the taxonomic
a
part
floral
of ovuliferous
other taxa
are
is
survey
phyllo-and stachyotaxis
development of
pistils
parts,
in the
and
of the
axis
of coaxial
true floral
(=
which
or the
on
of the
the basis
meaning
of the
for
In the second
in
new
of
part
ranalean
principal
anthocorm
of its connection
the
that
things,
gynoeciurn traditionally
of the
consequences
Evidence from various
of the
morphology
few excep-
subtending stego-
among other
constituents of the
reproductive region,
of its
their
axis).
implies,
with
gonoclads,
with
pteridospermous cupules.
discussed
given
kind,
same
and the floral
relationships
brief
number
interpretation
numerous taxa of the Polycarpicae the
called
a
gonoclads
theory.
interpretation for
with
the
comparative topology
ontogenetic
of fertile floral
organs.
1.
PREAMBLE
The
most
emphatic plea
neomorphological
tion has
met
not
an
(1915)
based
Thorne
was
published
1968;
region
most
(Meeuse
1965)
disappointing
since
of the
first
1966
Hutchinson
upon
Rohweder
to
essentially
new
so
often
already
a
notice that
(compare,
not
the
e.g.,
(1905,
present
Soo
1969)
1912)
century.
fare very much better.
1970) discussed
Caryophyllaceae.
observations,
relate
1969; Takhtajan
decades of
the whole did
(1967,
of Ranunculaceae and
groups has been studied
two
to
with taxonomic classifica-
scope and in method from e.g., Hallier’s
floral morphology
region
on
works
papers
example, recently
the floral
not
1968;
differ much in
Bessey’s
Interpretative
As
of the floral
interpretation
systematic
1967; Cronquist
do
the present author
by
with very little response. It
comprehensive
and
made
the
His
morphology
argumentation
of
is
because the floral anatomy of these
that
startling
discoveries
can
hardly
114
be
A.
It
expected.
decides
the
(compare
is
the
theoretical basis,
of such
outcome
also Moeliono
treatments,
about the obvious limitations
floral theories.
variance
within,
with
and the
tions in
question
can
of the
evaluation of the floral characters
synonymous
the various
1972),
attempt
different
a
It is my intention
to
still
and the
also
links
the
cycadopsid
gymnosperms,
There is,
or’
to
gymnospermous
there
Anyone
must
gy of the
there is
on
be
to
taxa
in
the basis offrappez
based
on
explained
I
relative
not
unbiassed,
the
in which
phylogenetic
progress of
inconsistencies
such
some
toujours
far
sources
(com-
awkward flaws is
these
and
into the consistent
cal patterns. The anthocorm
as
back and
wish
people
seems to
ovules,
failed
utterly
some
those
to
in the fossil record,
of the
Flowering
and does
of
accom-
no
Plants
from the
reproductive
admit that
not
is
to
not
in
for
the
wood.
However,
better times,
be convinced
at
all
-
so
that
-
soliloquy
a
1965, 1966), my floral theory,
quite original.
attempted
such
cupules
occurrence
babe
a
be indicated.
ecological
theory
with
directly
paleo-
of
Higher Cycadopsids
hoping
(Meeuse
information,
semophyleses
and
region
conventional, conceptual phytomorpholo-
but
palynological
origin
hard facts
Neumayer,
anatomical,
compatible
phytochemistry
reproductive
link all
is
of
only
of advancement of
much better
level of evolution of the
anthocorm,
of
the
as
to
in detail elsewhere
as
degree
Morphology
only regard
can
the
inquiries
out
inexplicable ‘gap’
as
face
not
just sitting
from other workers such
cerning
iron
Angiosperms
Old
no
amiss in the
the concept of the
from all available
of
evolutionary
in several
of the
early
be shown
can
1 doubt whether
As I have
the
feat the
a
does
region
point
compila-
‘derived’, which is practically
or
palynology,
angiospermous
the
who
something
floral
little
although
of
reckoning,
my
living
affinities
Plants. The
essence
level of
to
way
morphology
abominable’ mystery
is concerned if
region.
the
general
only
decidedly
systematic
show that indeed an alternative floraltheory, which leads
representatives
baffling
phytochemical
is also
approach.
to
living
plish.
is
on
detailed
classificatory hand-books,
‘primitive’
as
the
Angio-
based
1969)
of
b)
a,
Flowering
(and hence, of taxonomic groups), is
but
(1963,
recent
of ‘classical’
tenets
outcome
concerning
‘flower’
with all other relevant data from, e.g.,
botany,
and the
different evaluation of the
altogether
an
characters
of the basic
that
ponder
to
reason
point, phytomorphologically
a
of the
appraisal
sufficient
Huber’s
as
concerned. This has resulted
taxa
pare, e.g., Meeuse
to
ideas
to
in the
when and where of
how,
MEEUSE
anatomical data
morphological relationships
such
J.
assumptions,
satisfactory reply,
no
‘derivation’ of the
interpretation
with the
much the
1969, and Meeuse 1970
current
least up
at
primary
so
mildly)
be said of the above-cited
not
classical
still
are,
it
startling results,
phylogenetic
but this
the
the
of
opinion,
and
outlook,
in Kubitzki
(summarised
set
not
of the
(to put
systematic
rather
analyses, produced
studies
of
Surveys
holotaxonomic
broader,
at
There is still
1970).
descent, and this alone is, in my
sperm
a
the
and
pertinent question
the
to
studies,
D.
as
and
to
phylogenetic
other
data, and of
of certain
certainly
I borrowed
incorporate
morphological
based
evidence
sporangiate
course,
on more
heavily
indications
the
con-
organs,
results
of
and anatomi-
factual data and
FACTS
AND
FICTION IN
much
on
varied circumstantial evidence
more
monaxial flower,
such
gically
and it is
phytochemical
as
It also
provides
shall
of
I
see.
Bauplan
serve
owing
of certain
derived from
to
Another floral concept,
not
had such
hypothesis
and several
but it is
least
ductive
at
regions
strated the
basis
of
of the
regular
flower
in
ultimately
Theory
symbolised
floral
explains
relations between
‘cyclic’
or
Melville’s
coaxial
of
contrast to
Melville’s
all
all
by
does
and
not
sufficiently
viz.,
him;
the
flower
or
by
so
that
all this
postulation
Meeuse
certain
of
a
basic
from the
(i.e.
floral parts
(into
means
Gonophyll
between
(e.g.,
‘horizontal’
helices,
and corollae,
of vertical units
axis
sporangiate
the
quincun-
each
one
in
organs).
comprising
is rather similar
a
to
British-Australian worker
this
ovules, cupules,
microsporangia
interpretation
homologies
and
the
and
of the
carpels,
anthers
pistil
inhomologies
basis of his
highly conjectural
morphological unit,
and
proximal-distal)
instead of the
calyces
stamens
whorls
or
The
diagram.
better
a
supposedly
a
‘whorled’ assemblies of floral
on
gynoecial
quite apart
in helices
‘vertical’
megasporangia,
1971,
provide
(‘megasporophylls’),
pistils)
between
or
instigator,
of the repro-
studies have demon-
may
dissection of
floral elements
associated
structures,
(compare also
Winteraceae),
escape
a
one
discern between
other ovuliferous
stamens
of
least
polyaxial
a
the
morphology
which
floral
anthocorm concept, but unfortunately
own
my
at
of the
number of flaws in this
a
application by
heterogeneous
stamens
corresponding
interpretation
are
arranged
seeking
morphologically different,
leaf-like organ and
of
we
essential
no
as
as
application
treatments
‘flower’
painstaking
carpels
whorls, whole androecia and gynoecia,
xes,
divergent
which trends
1963, 1969) GonophyllTheory,
patterns
conventional
structures
groups
grouping
(1962,
traditional mental
of
morpholo-
of
details, but there is
reconsider the
to
morphologically
lobes and
perianth
of the
disciplines
classification
earlier
my
either. There
of certain
a
lines
pattern,
angiospermous
perianth lobes,
in
from
several
of the
of
terms
and
(‘microsporophylls’)
115
anthocormoid prototype.
Angiosperms.
than the
of the
the
taxonomic
a
inconsistencies in its
occurrence
comparison
monaxial
in
reception
glaring
of
diverge
meritory attempt
a
organisation
Melville’s
viz.,
cordial
a
theory
with other
the difficulties associated with the
taxa
basic,
a
that reveals
the frame-work
as
difference in the interpretation
has
POLYCARPICAE
than the ‘current’
compatible
reproductive regions
groups
present ideas
my
morphology
structure
of the
fundamental
same
add that
must
theory
a new
floral
can
better
rate
any
angiospermous
evolution from that
specialisation
at
THE
taxonomy.
basic
a
diverse
so
WITH SPECIAL REFERENCE TO
FLORAL MORPHOLOGY
hypothetical
of
of the
often
theory,
ambisexual
gonophyll.
A
small number of contemporary
developmental
and
anatomical
from the conventional
of the monaxial
cal
exhaustive
1961;
interpretative
reproductive region
interpretations
Barnard
based
Pankov
survey
by
on
the
1962;
workers
studies,
to
floral
or
Moeliono
Heel
(1970)
chiefly
of
1966;
a
from the
alternative
polyaxial
Moeliono
explains
by
means
inconsistencies
morphology (i.e.,
‘flower’) by
assumption
van
attempted,
explain
the
of
resulting
postulate
neomorphologi-
floral
region (e.g.,
1966, 1970).
historical
A rather
background
116
A.
and
also
the several
emphasis
ovules
variations
much laid
too
was
(stachyospory
of
interpretations
on
structures
gynoecial
A
borne
not
on
neglected
rather
leaves
but
at
and
ships,
pointed
be
enumerate
a.
it
must
Another way of
essential
universal
taxonomic
sense
recognized
in both
which
in the
entity
it
in
of
must
must
thus
that
provide
clear-cut
cycadopsid
it
taxa
must
floral
as
be
on
it
moot
must
must
the
a
a
and
barely
(because
in
cycadopsid
is
fulfil:
to
and
morphological entity ( Bauplan)
basis for
it
question
morphological
a
I
plausible
theory ought
in
as
small
no
Plants,
provide
point
and gymnospermous
can
groups,
comparisons (homologizations)
the gymnospermous with the
angiospermous
between, and indicate the topo-
homologies
organs
particular
and/or
the
intervening
and
the
the
the
complexes
hypothetical pre-angiosper-
connections between certain advanthus
primitive angiosperms,
‘missing
links’
with conditions and
postulate
in
floral elements of
organ
fertile
gymnosperms;
semophyletic
centrifugally
maturing
pinpointing
;
in the
phenomena
angiospermous
complications
of the monaxial flower and
in
its
taxonomic
stamens,
consequences be
anatomy, palynology,
not
on
in floral theo-
the conventional rules
laminal,
dedoublement,
parietal
and
‘free
compatible
phytochemistry
and
with evidence
other
from
taxonomically
and
disagree
more
with
theory does, and, conversely,
of
in
have,
irrelevant
floral
well
as
which have caused difficulties and
applicable criteria;
it
relation-
contended
placentation, etc.);
ontogeny,
f.
tenets
Flowering
aestivation, insertion and alternation of floral parts (apetaly,
central’
(OCU,
cupule-
the connection
emphatically
seems to
the
general
a
of Mesozoic
gymnosperms
the
obdiplostemony,
e.
is
phylogenetic
proved
with discrete rather than
organs
compatible
region
ries based
of
which
formulating
specific
in
relating
it must, therefore, show the
d.
most
of the
let alone
fundamental
a
of various
gynoecia
reproductive
certain
theory
angiospermous
provide
region,
androecia and
ced
are
all
evolutionary advancement;
reproductive
c.
I have
phylogenetic
a
course
logical equivalence
mous
cupule
primarily
important subject
ancestry
requirements
in
reproductive region linking
level of
b.
all
be
the
publications,
recent
denied).
to
be
ovules
of
point
serious consideration. This fundamental issue is
given
in
upon
not
can
equally
morphology.
for
quest
starting
the
leaf-borne
stems.
or
that any other floral
out
should be
touched
the
impeded
measure,
answer
the
MEEUSE
Regrettably,
versus
ovuliferous
1965, 1966) that the ‘classical’ morphological
(Meeuse
have
be the
(systematic classifications),
floral
interpretative
themes
because
must
1971): cycadopsid
least
between taxonomic affinities
main
two
the controversy of axis-borne
phyllospory),
versus
compare Meeuse 1963b, 1964a,
borne and
these
on
D. ].
teratological
provide
cases
better and
than
more
any
classical floral
consistent explanations
abnormalities.
It follows
that if
morphological
preferred
to
an
tenets
any
interpretation
meets
other
one
of the
all these
floral
region
based
requirements reasonably
which fails in
some,
or
in
on
certain
well, it is
neo-
to
be
most, of these respects.
FACTS
FLORAL
FICTION IN
AND
There is
in
point
no
WITH
MORPHOLOGY
the classical
discussing
REFERENCE
SPECIAL
floral
defence of the traditional interpretation of the Old
are
it
available (Eames
to
does
not
only require
ties mentioned sub
the
leaf
1972).
which
Grégoire)
of fertile floral
evolution of the
in thefloral
A
Meeuse
of
the
occurrence
floral part has
are
example
An
carpel
postulated
been
the
a,
1970),
the
Trollean
1937-1939,
b).
adduced and
reasoning
1967,
1970,
whilst
The
school
1939;
There is
alternative explanation
little
as
additional
2.
AN
and
or
the
stachyotaxis
is
straightforward
publications (see
of
floral
all
region supposed
and
called
as
a
parts
to
be
‘flower’.
if each fertile
some
to
way
even
ascidiform)
phytomorphologists
using
Leinfellner
the
against
paper
same
(compare,
1950,
Moeliono
or
1969;
arguments and
times before
fundamental issues
even
(compare,
1966,
with
disregarding
called ‘eclectic’
by
convince other workers of the merits of
an
by adducing
bit
the fundamental
changing
peltate (or
has been done many
attitude has been
by showing,
by bit,
manner
a
euphemistically
variety
that
can
a
be
of arguments
from
different
number of absurdities and in-
explained
without the
aid
in
a
logical, plausible
of sometimes far-fetched,
premises.
APPRAISAL
OF
The
Gonophyll Theory
of
blade
a
in
theory proposed by
interpreted
1949;
in
point
the second
avoiding
of
Baum
consequences of the conventional theory
often
interpretation
nature, and various inconsistencies and other diffi-
of the
only
and
Angiosperms
without
an
and
the
among
theory
certain evidence. Such
(1969).
monaxial floral
theory
Sterling
disciplines
and
Thomp-
continuity
phyllo-
interpretation
the
same
Rohweder
Moeliono
worst
phylogenetic
in my earlier
signalised
of this kind is the
by
1932,
Klopfer 1969
following
its
McLean
any
A floral
of
such
Piperales (Meeu-
The
reasons.
lacks
in
Gymnosperms
semophyletic
homologues.
subsequently
phyllomic
a
tenets.
Troll
of
assumption
is, I think,
(Saunders,
1964)
preconceived
explained by modifying
are
idea
already postulated,
an
Anatomical features
e.g.,
the
and their
already
1966), viz.,
ubiquitous
e.g.,
violates
(1961,
the
genitalia
and the
this
will be discussed in connection with
attitude has
peculiar
culties
Croizat
as
between
‘gap’
elements
region.
(as phyllomes)
of
by
sporangia
Plantefol (1948)
traditional
Suffice
for the difficul-
gynoecial
female
and
-
obsolete for various
now
support and
anatomical
fails
discuss older theories
not
are
also
theory
the
structures
and
1970).
normal, angiospermous
inconsistencies
“flowers”
Cercidiphyllum, Euptelea
floral
We need
unlikely
derived
as
taxa
bridge
to
-
Angiosperms.
e.g.,
such
to
unduly
The classical
shortcoming
son,
conformable to that of a
in
surveys
of the monaxial flower
to account
-
117
partly
-
good
-
Moeliono
principle
ancillary hypotheses
readily
but also leads
of
because
theory,
also
see
of the
POLYCARPICAE
Morphology
and for the anatomical structure of the
not
manifestly primitive
se
application
-,
occurrence
1967;
Rohweder
various
(d)
which is
(‘carpels’)
foliage
1961;
that the consequent
say
THE
TO
(to
MELVILLE’S
is based
on a
GONOPHYLL
hypothetical,
all intents and purposes
a
THEORY
ambisexual ‘unit’
leafy organ)
from which
a
consisting
fertile
struc-
118
A.
ture,
sporangial
a
supposed
are
the
truss
is
in
gonophylls’,
second
have
to
occurrence
order
several such
or
and
(or androphylls)
ants
their
of
which
relation
a
of the latter
(each
phyll
blade and the
cipal
gonophyll
associate
blade and
complex
more
between
an
of the
entities,
axis
of
coaxial blades
kind of
a
their
plus
of
of
axes
a
ultimate gono-
simple,
a
In addition,
‘gonophylls
first and
is assumed with
sporangial truss)
subordinate,
vari-
gynophylls)
sexes.
viz.,
of the
of
stipe
common
=
one
MEEUSE
J.
Microsporangiate
counterparts (or
reduction of
originated by
postulated
arise(s).
trusses
megasporangiate
D.
prin-
sporangial
trusses.
Melville based his concept of
stead.
Her reconstruction
that of
bivalved
a
microsporangiate
but also the
alliance;
any
on
the
event
the
the
was
of
question
seek for
be
can
ascertained,
pteridospermous
phylogenetic
of the
gnetalean
truss
of
of
the
(ovules)
are
coaxial ovules
(which
ovule
to
or
compelling
the
in
here.
among
have been
rate
opinion,
my
aril
a
and
organ
The
the
only
not
but also
1964a,
(Meeuse
the fertile organs adnate
single pistil
far
as
rare).
angiospermous
some
Plants
microsporangiate
represented by
Plum-
by
cycadopsids,
of
in
con-
reason
every
unisexual
rise
ultimately only give
to
Cupules
pteridospermous
always
Mesozoic
true
were
reason
glossopterida-
that there is
so
Flowering
single synangial
a
the ultimate level of evolution
no
was
side and
one
been confirmed,
never
point
any
is,
words,
could
the
on
Plum-
by
blade
leafy
a
described. However,
were
must at
In other
been
age,
of the
pistils. Any
or
erroneous
fertile organs described
of the
forms,
Glossopteris
angiospermous
or
cupules
1966).
to
organ among
common
peculiar
of other groups of
also Long
has this
crucial
megasporangiate,
gnetalean
pistil
likewise, only give rise
at
is
exceptions
of the
blades of
reproductive
if
cupule,
groups and of the
see
that
surely
elements remains obscure:
of fertile
homology
organs
so
archetype
chlamys
1971,1972;
only
other types have
of other seed ferns
cupules
attached
ovules? There is also
connection between the
a
stead and the
as
Not
and their derivatives of Mesozoic
cycadopsids
almost
an
megasporangiate
sporangial
or
only type
the contrary:
on
glossopterid pteridosperms
portion
was
one.
ultimate
synangia
taining megasporangiate
to
which
opposite
of the
thecate
that this
assume
lean
gonophyll
of the fertile
structure
on
nature
they sporangia,
to
the
of fertile organs of certain
interpretation
not
to
to
a
a
the
to
single
‘naked’
counterpart would,
(compare
van
the anther of the
Heel
1969),
angiospermous
stamen.
In Mesozoic
the
cycadopsids
megasporangiate
almost
bears
several
one must
(or
the
Meeuse
search for forms
subtending
represent
ovules
cupules:
a
an
consider the
exceptional
number
of
showing
co-axial
offshoot of the
ambisexuality
If
occur
in
Angiosperms
whole
1971).
gynoecium
one
a
is
a
fora
combination of
cupules.
glossopterid
majority
of their
it
is
a
of coaxial
ferns
groups
of
prospective prototype,
single
foliaceous organ
that
Assuming
seed
and
primitive placental region
consists
looking
(see fig. /),
trusses
(Meeuse
of their so-called sporocarps
in respect of the
and Nilssoniales
Caytoniales, Pentoxylales
as
homologues
that in the
universally agreed
modified
such
cupule
as
progenitors,
the
1961),
atypical
as
a
Marsileales
we
and
kind of
may
highly
speciali-
FACTS AND FICTION
Fig.
1.
Mesozoic
cupules
bly
or
with
of
MORPHOLOGY WITH
a
of
microsporangia (microsynangia)
bract which
the remains
may
of the fertile
left:
reproductive structures). Top
bottom
spermaceae,
zation which
water
and
most
which
ovules
based
the
one
giate
the
species
strand
(or
from various
truss
soon
trace
was
not
adnate
to
genus
to
the
resulting
combination of
Caytoniales {fig. 1)
From this stage
are
is
very
plausible,
a
Corysto-
most
primitive,
the
ones
independent
an
lowermost
(i.e.,
in
(as
system
off from
cupule (=
the
cupule
in the
sporo-
=
sporo-
majority
instead of
of the
ambisporanand
its
that of certain fossils such
as
(frond
development leading
a
associa-
coaxially superimposed
foliar organ
and such
in
bottom left:
undoubtedly
ovules is
resembles
a
found
the male and female
do with the survival of this
the frond petiole
cupules
Ovuliferous
axes, presuma-
bifurcating strand) branching
below the
119
text-books).
and bore several unisexual cupules
of ovuliferous
Hydropteridangium.
to
on
genera, Marsilea is the
of this
supply
a
forms.
vegetative phyllomes
in the centre, between
something
recent
POLYCARPICAE
borne
coaxially
that in the ancestral forms the fertile
axis
the
truss
angiospermous
frond is associated with several
frond-cupule
Marsilea)
ones,
axillary
(shown
have
the
The vascular
a common
carp-bearing)
of
Among
long-stalked
common
of
were
Nilssoniales, top right: Caytoniales,
even
conservative
carp). Assuming
species
perhaps,
ferns.
(see fig. 2).
on
have resembled
organs
right: Pentoxylales (adapted
may,
group of
the
SPECIAL REFERENCE TO THE
Cycadopsida, putative progenitors
groups
in the axil
tion
IN FLORAL
to
or
leaf)
the condition in
combination of
a
foliar organ
120
A.
2.
Fig.
At the
(adapted
leaf
(with
Various
vascular
prototype
of
is
sporocarps
fossil
a
as
could
easily
and
a
be the
of the
one
anthers),
such
an
arrives
at a
stalked
arillate
have
a
ovule),
a
also
fused
more
between
case c
with
original
the vascular
the
petiolar
condition.
basal
trunk
as
far less
Caytoniales
cycads,
recent
with
trusses
cycadopsids
origin.
Melville’s
vascular
ultimately
bract
,
an
organ
(androclad,
of
consisting
organ,
(=
stalked
(= stegophyll),
entity
male
in that their
are
may be
two
cupules (=
and it also
synangia
explains
why
and
a
respective
and
a
gonoclad (polygonon)-bract
now
was
originally
obsolete suggestion
by
vascu-
but
cycadopsid
it is
groups
and
Corystospermaceae,
chlamys=aril
more
or
unit is
ambisexual
Plumstead
as
we
that
by
of seed
in
fern
an
primitive higher
a
pistil
of cupular
postulated, primitive
Melville believes
(principally
have
or
gonoclad-
ways,
glossopterid type
the seeds of the
a
a
components
Marsileales) associated with
sporocarps in
nearly always enclosed by
primitive gonophyll
truss
by
principle emerging
several
in Mesozoic
occurs
derived from
in
Another fundamental difference between Melville’s
gonophyll
a
‘gonophyll’
it
a
represented
the
vascular trunk in
Nilssoniales, Pentoxylales
and
of coaxial
-
-
enigmatic
uni-ovulateand in certain groups
monandra
single
hypothetical
(not
system
its male counterpart
sometimes
cupules,
simple
(=
of the coaxial
type.
sporangiate complex
constituting
(= cupules).
sporocarps
gynoclad (or polygy-
a
microsporangiate complex
of
of
plant
compound
a
The somewhat
1972);
Meeuse
Antholithusarberi
or
sporocarps
MEEUSE
of a
piece
bears
the truss
then, is
cupules,
truss:
subtending
fertile
J.
related.
possibly
ovuliferous
a common,
unit resembles
axillant frond
of
of
caytonealean
(=
phylogenetically
the
is
In
a
stelar leaf gap. The monogyna and monandra are ebracteate. The
bract
the
the
coaxial
archetype
associated with
supplies
a
left)
phyllome.
of ovuliferous
single
together anatomically
in
shows
a
the
truss
a
less
or
case
of
the rhizome
bunch of coaxial
the relations
completely
more
but
a
Meeuse 1971,1972,sometimes
becoming
lar
associated
often
monogyna
pistil:
an
and
In this way
of
has
personal observations):
primitive gonoclad)
strictly
polyandron)
and
to indicate
(a, b, c)
Hydropteridangium (at
when
non
which
bundle)
drawn),
(stegophyll)
semi-diagrammaticrepresentations
,
text-books,
shown
cases are
separate
Marsilea
right;
from several
D.
seen),
on
the
that
ground
whereas
in all
FACTS AND FICTION
fossil and
gonoclads
have
to
were
pointed
exhibit
of
vice
in that
polliniferous
that
so
gonoclads
‘mixed’
seems
entities
this
lineages
survived
terminal groups with
the
as
Until
had
only
ture
of
limited
hardly
primitive
‘mixed’
(or,
gonoclads
not
as
ous
‘modified
familiar
Monocotyledons
which
gynoclads)
clads
in the
there
is
flowers
a
and
either
which
his
elements,
and in my
to
the
ville’s
in the
interpretation
often rather
which
and,
is,
to
in
my
doubly polyaxial)
logenetically
I have
and
already
is
(as
sex
axis
in
ap-
struc-
see),
and
the conventional
but
only
is
all
modified
unisexual)
why
flowers’,
must
one
‘mono-’ and
functional
in the
floral
male
of
in the
but
a
an
case
of
ancillary
upon
by
always
unisexual in
a
of
occurred
gonoclad,
theory
probable
the lack of
to
in
from
part
and
in
or
gynoecial
a
number
were
strictly
Thus
progenitors.
an
of the
flower,
same
only
again
of certain
starts
gonoclads
Mel-
complex androgonophylls
Cercidiphyllum,
‘gonophyll
or
Here
one
both androecial
means
of
unisexual
part).
different part of the
a
supply
female,
bear their andro-
interpretation
necessity
become
or
numer-
andro- and
(separate
distal
more
of the
of
presumably
consistently
androgynoclad
structures
morphologically
touched
in
and
gonoclads
exclusively
version
to
the
postulation
more
shall
we
that
to
of their gymnospermous
of floral
the
and
whereas in all other cases the
majority
fact, nothing
mind,
other
explanation
devious
as
occasionally
‘bisexual
almost
seems,
either first
floral
angiosperm progenitors,
as
it
interpretation
must
ultimate flower and of the
adnate
Mono-
androgynoclad
of the floral
anthocorms,
Dicotyledons
arranged
as
between my
Melville’s:
becomes
unisexual
and
lower, and their gynoclads
androgynophyll
advance of
some
groups. In several of the
Piperales,
homosporangiate
(which,
discrepancy
a
groups.
bore
were
‘mixed’ anthocorms
in
be emended
must
Cercidiphyllum),
indiscriminately
etc.
of other groups of
progenitors
few
a
rosalean
whole
not
‘mono-’
‘apetaly’,
angiospermous
idea
Polycarpicae
androgynophylls’);
terms
that the
interpretation
some
also in
are
and
Cercidiphyllum
‘perianth’,
units of all
The
in
the
but this
being.
of which
some
also
presumably
‘mixed’
into
evolution
Angiosperms,
and
the
to
also
came
floral
partial
Winteraceae and the monoclinousLaurales
(and
Morphology
(Melville:
apply
‘dicliny’,
mentioned
Old
of
I
clearly
the
additional phylogenetic
any
of several other groups of
taxa
‘flowers’ of the
evolution
quite erroneously,
the clue
provides
of
archetypes
also of the Hamamelidales and
probably
more
I believed,
least the Lactoridaceae, the
at
(and presumably
is
the
evolutionary potential,
because it
level
the
As
ovuliferous
equivalent
androgynoclads)
principally dicotyledonous
fairly recently
a
preciably,
of
(=
reproductive region: Piperales, Euptelea
cots.
polliniferous.
or
Chlamydospermae
the
by topologically
prefloral
clearly
number of
initially
incipient monocliny originated by
polygona
or
are
of
IX),
of the
usually
that
to assume
i.e., either ovuliferous
organs
at
distribution is
sex
plausible
more
1965, Chapter
(Meeuse
versa,
(ambisexual)
It
homosporangiate,
transitory phase
a
or
Such
dicliny.
as
out
‘replacement’
ones,
groups of gymnosperms the
recent
type referred
121
IN FLORAL MORPHOLOGY WITH SPECIAL REFERENCE TO THE POLYCARPICAE
the male ‘flower’ of
and of the
of
fit the
Magnoliaceae)
gonophylls’
hypothesis
(which
than
a
is
phy-
structure.
precision
of the
Gonophyll Theory
122
A.
when it
ferous
which led to,
1972),
logy
of
Meeuse
The
quality.
must
One
(we
the floral axis
to
there is
is found in
precision
a
bundles
or
an
late’ the
phylogenetic divergence
vascular
van
such
explanations
stegophyll,
tively
for
inevitably
by
structures
in
and
little
Melville
at
reader
is
ideas
We
rate
bract
to
anatomical
uses
bud
of agreement between
floral
decide,
but
it will
be
clear
why
I
to
and
of
interpretations
theory
nodal
so
and
need but rela-
which
think that
that
and mine. One
closely parallel,
question
at
stegophyll
characteristics,
run
has
‘trans-
‘leaves’
or
with the available data and indications from other
to
ai),
fertile axis
interpretations
coincide with mine. The
fertile axis,
(axis) complex,
his
morphology
a
et
have
a
that bract. For his
(leaf)-axillary
of
of the orien-
of
terms
be
cause
requiring
(‘inversion’)
foliar bract
only
can
may
even
explanation
in
a
e.g.,
of ‘invert-
which
shall, therefore,
of Melville’s arguments and
to
(compare,
Heinsbroek
of Melville
adnate
concerning
clue
turning
or
axial
Higher
syntelomes
occurrence
sometimes
(see
remainder which
any
a
points
readjustment
compatible
Theory
3.
our
several
sense
be
may
and
of
the arrangement and
a
the
The
gonophyll
with), because
parts of the stelic system of
case
(partially)
in
reported,
the combination of
yields
and the
indeed several
that
the
or
and done
us
again
ones.
foliar
polytelomes
give
twisting
straightforward simplicity.
simple gonophyll
why
the
of
is
a
organ systems
Morphology,
the
as
the
as
between
or
over
of floral parts
as
‘inverted’ bundles
such
subtended
may say
and
parts of organs in respect of other
anatomy, leaf gaps and all, of
best
(compare
tangible morphological
a
mainly
may
now
in the Old
interpretation
adnate
are
this
1969). Every
in the vascular system
which
of floral
and
tissues,
Heel
(the ‘gonophyll blade’)
that is
morpho-
adnation of
an
of
description
distinction
a
in which the
least the merit of
there
after
concerned with organs
clumsy
tation of
a sort
the vague
difference in the anatomical pattern,
difficulties of
‘stems’,
of
far off the mark
so
all,
Heinsbroek &
organs
not
(see Meeuse,
the floral
principle
rather than
pictorial
a
seek
always
after
are,
orientation of
rather
is
types of vegetative organs had long been
two
of
complex
lack of
Cormophyta
or
the basic
MEEUSE
1971).
same
organs
ed’
and monandraetc.
concerning
J.
of ovules, ovuli-
(interpretation)
rather confused idea
a
e.g.,
ancient structure’ with
‘an
into
definition
morphological
Drimys (Melville 1969), although
androgynous
an
as
the
to
comes
cupules (= monogyna), polygyna, poly-
D.
theory
is
disciplines
is
the
Gonophyll
inadequate.
SOME EVIDENCE IN
FAVOUR
OF
THE CONCEPT
OF
A
POLY AXIAL FLORAL
REGION
The evidence is
the
first
(1952)
of the
way
can
partly
of
circumstantial and
argumentation,
a
partly
paper
by
be cited in which the conclusion
angiosperms
makes it rather difficult
lution of the
Flowering
lae, and also
to
Plants from
Spadiciflorae (still
a
was
to
more
direct. As
Suessenguth &
drawn that the
believe that
Magnoliales
wide-spread
to
a
an
example
great diversity
monophyletic
Amentiferae
of
Merxmueller
evo-
and Composi-
belief apparently!)
could have
FACTS AND FICTION
taken
the
in the
place
Early
short
relatively
Tertiary (when
or
discernible in
clearly
In
of floral morphology, the
one-ovuled
such
in
in
of
consisting
and
least
at
this
of
purposes
flowers
of
Euptelea
the latter becomes
unilocular
than
primitive
the
of Chloranthus and
exhibits
either
ancestral
,
floral
the
of
more
than
‘
when
The
In such
or
can
conventional
its
to
Piperales,
not
unlikely
(it
a
‘flowers’ of these
of
magnolialean
descent of
piperalean
a
the
true
Schisandraceae and
important
as
tool
or
are
forms from
a
forms from
show
of
stigmatic
supposedly
derived
explanation
Euptelea,
primitive
struc-
in
Meeuse
on
respect
Among
the
the
of
1970a, b)
magnoliaceous prototypes
stelated floral axis
more
crests,
ranalean flower is thus
1969,
around).
Sar-
as
classical
least either
very
the
receptive
considerable doubt
(Kubitzki
other way
the
The derivation of the
most
histological
in the flowers of the Lac-
Winteraceae, which renders the interpretation
modified monaxial shoots untenable
to
of
ranunculalean Polycarpicae
etc.
ovule
signs of a
any
of Amborella, of
nature.
throw
single
complete phy-
inevitable)
say:
Urticales
‘carpellary’
data also
Helobiae,
would rather be
of these flowers
a
primitive
Phytochemical
arguments, the absence of
toridaceae,
be of
possibly
Centrospermae,
and render
even
at
for its
account
would
a
under the
suture, of
a
more
ranalean
disappearance
so-called ‘flower’
pistil,
(I
leaving
would,
one
uni-ovulate,
vesselless Dicot
primitive
a
the
the localization
clear indications of
plausible
without
of Sarcandra (and of other Piperales),
rendered absurd.
position
to
a
pistil morphology
to
the total
to
standardised
considerably
the
These relate
carpel
‘apetalous’
‘conduplicate
describing
Amborella)
find in the
to
ovule in
most
the
all intents
to
a
solitary carpel bearing
crests, and
to
be
may
conventional
connection with
face the fact that
style
histologically,
carpel'.
pistils),
one
the
pistils
which
‘basic’
to
from
and
Piperales,
peculiar
semophyletic
oli-
manifestly pri-
family
a
the
pluri-ovulate
also those of other Laurales and of the
tures
an
p.121),
surfaces of
applies
expect
more
Surely
is that the
and
same
hypothesis,
status.
has
proper
stated:
or
the
of the
far removed
so
basic,
stigmatic
the
of
apex
the
(and
‘carpels’ (i.e.,
and
the ventral
morphologically
external double
at
candra
(1950,
Sarcandra,
and
of their direct
a
gynoecium
modifications,
with
orders, especially
highly modified evolutionary changes.
logenetic fusion of
surface
theoretically
pseudo-mono-
theoretical derivation. The
simply
without
organ
Swamy & Bailey
carpel’.
suture
assumption
be
to
series of
compatible
Middle Cretaceous,
absurd. One
too
pistillar
considered
whole
a
morphologically
are
ranalean flower that the
in
Urticales,
number of Monocots
a
(dioecious) Cercidiphyllaceae,
the
as
in
Laurales,
1970).
one-chambered,
apocarpous ranalean
an
all
at
of that
out
of the
old
as
not
a
groups
Muller
basic, conduplicate (or peltate-ascidiate),
of these
some
bottom
‘flowers’
apetalous
supposedly
carpels through
least
at
the
and
from
cases)
and reductions, is
status
knocks
and
foliar
pluriovulate
and
Piperales
Helobiae,
many
number of
number of
a
gomerisation,
mitive
and
a
123
POLYCARPICAE
angiospermous
derivation of
semophyletic
many
THE
record: compare, e.g.,
Hamamelidales, in Euptelea, and in
some
Cyperaceae
as
merous
found in
pistil (as
Juglandales,
basic
perhaps all,
the fossil
TO
between the Middle Cretaceous and
time-span
some,
become
terms
SPECIAL REFERENCE
FLORAL MORPHOLOGY WITH
IN
the relative
complexity
(Meeuse
of the floral
1971),
region
is also
of the
124
A.
and their associated
Magnoliaceae
in another
of the
anthocorm
such
interpretation
Winteraceae, and
of
taxa
advanced
gynoecia,
rous’
theory,
and do
Euptelea,
be
to
prove
not
more
appear
as
‘overall’
groups with
(such
in
interpretation
if interpreted
demonstrate. In this
Cercidiphyllum,
truly
terms
Lactoris,
primitive types
some
incongruous,
‘monome-
dicliny, apetaly,
as
axial stele in the floral
an
plausible typological
to
shall attempt
features
morphological
the consequence of their
I
as
the much
primitive Angiosperms,
Piperales,
as
absence of
of
regions
Schisandraceae
Angiospermous plants
highly
provided by
between the floral
relationships
terms
MEEUSE
will be discussed
as
groups,
J.
paragraph.
The best indirect evidence is
in
(and derivative)
D.
“axis”, etc.)
would be
as
of the classical floral
morpho-
logy.
4.
PRIMITIVE
CONDITIONS
IN
THE
REPRODUCTIVE
REGION
OF
SOME
LIVING ANGIOSPERMS
An
interpretation
tates
in
structures
ly
living
derived. The
living
of the floral region
documentation in the
some
taxa
taxa
obvious absurdities
more
useful
as
a
the
Probably
of these
subsequently
of the
relationships
from which
occurrence
may
a
of
clues
anthocormoid of the very old genus
by
Cercidiphyllum.
the
stamens
(monandra), traditionally
be the
of
reduction of
numerous,
only
has
axes
of
a
and
both
bearing
it, but is rather
a
polyandra,
brachyblast.
and of the axis of the whole
The male anthocorm bears in the
units each
consisting
of
few
a
of
a
at
conventionally
structure
derivative of
bracteate
a
pistils
gynoclad
is
the
called the
pistil
in
other
species
(androgynoclad).
indicative of
primitive
in
a
very
the
status
complex
primitive
presence
of this
form
to
a
fertile
reasons
the
bracteated ovuli-
a
carpel,
of the
a
few androclad-
topological
words,
not
Old
must
the
represent
a
of 2-8
complex
female anthocorm
The conventional ‘naked’ flower of
gonoclad
or
if
the
as
shortening
a
anthocorm,
recent
process
some,
primitive androclad,
being
monandra. For
(polygynon);
least
carpels,
female counterpart of the androclad in this dioecious genus,
ferous
unisexual
male flower devoid
a
and
stamens
perfect example
result in
long semophyletic
advance in respect of the ancestral condition
of the
bract
product
is,
bracteate but other-
bisexual ranalean ‘anthostrobilus’ with
perianth lobes,
Morphology
the
a
not
not
in any event,
primitive
Each
possibly
can
in
phylogenetic
one.
wise ‘naked’ cluster of
perianth,
the
working hypothesis
of
a
floral
logical-
morphological patterns
concerning
provided
be
can
If these deductions do
is
case
special, primitive
structures
than the conventional
illustrative
anthocorm necessi-
polyaxial
advanced
good
the
incongruities,
of
examples
complicated
more
question.
general theory
most
terms
more
provide
in
groups
or
in
form of
The
of
genus
a
simple
precluding
is in fact
Euptelea
simple
very
pattern
morphological
the
possibility
of
elements,
viz.,
stipitate
bracteate
mixed
vascularization
structure,
the
is
generally
of wholesale reductions
system of floral venation. The stalked pistils
gynoecial
a
are,
ovuliferous
consequently,
cupules
or
mo-
FACTS AND FICTION
The
nogyna.
IN
MORPHOLOGY WITH
FLORAL
patterns
same
ancient type is that of the
raceae
noclad with
a
few
few
a
and
uni-ovulate)
1971,
arises whether the
question
phylogenetic
number of
from OCUs,
increased,
secondarily
of ovules
quantities
in all other
supposed
be
to
more
the conclusion
in
e.g.,
less
or
(or ‘carpel’
taken
in the
least
at
marked
taxa
the
less
by
related
primitive
that
the
a
primitive
assembly, provide
a
a
a
all
ones
one
deny
not
can
have
must
enormous
the numbers found
orchids.
After
of the Rosaceae
a
groups
thorough
(published
1969), Sterling
each
in
to
came
element
gynoecial
corollary
is
increase of the number of ovules has
of
representatives
with
taxa
derived
pistils
monocotyledonous
pistil
the
clear that in
is biovulate. The inevitable
secondary
not
and
along only
produce
surpass
Sterling
rosaceous
with
pluri-ovulate pistils
because
only
primitive
a
possible example
think,
of
have
they
are
an
con-
group
be
can
ear-
pluri-ovulate
gy-
old offshoot of the
secondary polyspermy
among
Polycarpicae.
of the
most
In
the
excepted)
that
an
happen
other
gonoclads
which
anthocorms)
is
far
the
to
noecial elements. The Aristolochiaceae which, I
ranalean
It is
which
advanced)
summarized in
Rosaceae
most
that
ex-
Meeuse
(see
variable
number of ovules
Orchidaceae,
closely
with uni-ovulate and
the
as
is
other hand
the
taxa
terminology)
current
some
stamen
one
monogynon
gynoecial morphology
the
It follows
place.
taining
in
only
On the
place.
flower, quantities
of the
that
per
angiospermous
series of
cited and
papers
that in
took
an
androgy-
an
the evolution of one-ovuled
(and generally speaking,
comparative study
a
per
caused
is
most
Pipe-
monogynon (normally
one
progressive oligomerization.
oligomerisation
that in several advanced
only
( Chloranthaceae)
manifestly during
cases,
‘flower’
evolutionary development proceeded
trend of
The
four subterminal monogyna (each
or
of ovules
number
reduction has
saururaceous
families there is
two
sometimes
The
fig.la).
and three
stamens
alliance.
piperalean
and
oligomerization
conventional
in the other
ovules);
single
A
simplification.
the
125
POLYCARPICAE
because in the Chloranthaceae and
Saururaceae,
towards
tendency
a
cessive
found in
are
REFERENCE TO THE
SPECIAL
Dicotyledons
are
act as
functional flowers.
anthocorm evolved into
only
The basic
once.
a
flower,
partly
ial, separate
one
groups
semophyletic
trends of
distal
zone,
almost
but,
region
monogyna
follows
evolution into
specialisation.
gonoclads
male
(zones,
whorls)
with androgynoclads.
although
that these
flower
seem
the
to
one can
by
the
(found
ones
types,
the
and
can
occurrence
of
serve
the result of the
subtype
groups
with
androclads,
as
same,
or
taxa)
coax-
and
prototype
a
not
of three
in all diclinous
one
state
process did
semophyletic
gynoclads
as
Juglandales
pattern in several major
of these
have
gynoecial
androgynoclads,
(combined
of
Each
two
as
the
for
a
of different
Ambisexual anthocorms with
female
separate male and
always
of the
a
of
again
such
complexes (modified
Generally speaking
caused
types of primitive anthocorms, viz., unisexual
and ambisexual types, the latter
other
Amentiferae
but this
morphological
different. This is
fundamentally
(some
combined in various ways into
with
a
distal female
elements
the latter may
proximal
different prototypes,
or
distal
are
and
zone
usually
occasionally
monandra,
combined with
some
a
proximal
inserted in the
bear
proximal
respectively).
divergent
It
evolu-
126
A.
tendencies,
tionary
SOME
The
in
ADVANCEMENTS AND
early
phylogenetic
have started before
early Tertiary.
manifest and
lutionary
The
in floral
divergence
easily
of
the
advancements
be
can
the
recognised,
a.
progressive oligomerisation
of the number of monogyna
of the
of
aggregation
of
or
monandra
into
fascicles,
tufts,
gonoclad, and/or
cupule);
bundles,
or
be
evo-
per anthocorm,
gonoclads
monandra per
monandra into
and/or
in the
already
groups. A number of
number of
(OCUs) and/or
monogyna
lineages,
principal (and partly alternative)
of the number of ovules per monogynon (= per
an
culminated
various
must, therefore,
basic
more
of the
mainly completed
was
morphology
demonstrable in the
which
Angiosperms,
in the Cretaceous and
or
being:
b.
of floral
early divergence
types of floral organization
ones
a
MEEUSE
DEVELOPMENTS
divergence
the fixation of the basic
must
the manifest,
J.
Angiosperms.
types among the
5.
for
responsible
are
D.
subterminal
of
or
whorls
monogyna
into
‘phalanges’;
c.
the
cyclisation
of
androclads and
d.
the
adnation of
e.
the adnation of
f
the
association
the
such
In
gonoclads
as
the
1971):
‘syncarpous’
some
the
genus Lactoris
the
case:
anthocorm
one
longitudinally
congenital).
etc.,
initially
appear
and
In
a
,
one
as
In
or
the
false
is
a
three
a
column
complex
gynoecia,
apex
for
,
and the
interpretation
subterminal
(in
two
(their
three
Dicots
Capparidales
is
instance),
or
their monandra with
for
,
a
Malvales
with
one
more
instance).
(Meeuse
sometimes
the universal trend
Among
1965).
an
became
large part
(Tiliales),
whorl of gyno-
whorls)
sometimes
they
became
apparently
comprising Dilleniales,
s.s.,
with
are
gonoclads
concrescence
each with
nearly
(in Flacourtiaceae, Hypericaceae,
anthocorms
vas-
(Meeuse
gonoclads
(occasionally
fascicles
single gonoclad,
Cyperaceae
into
subterminal monogynon and
dialypetalous
( Dilleniaceae
primitive
axis);
organisation
usually ambisexual, probably
axis, only
protruding
Centrolepis
floral
staminodial. The
be
a
to
single
a
Violales,
Euphorbiaceae-Ricinoideae
filaments
few whorls of
and
form
to
following
monandra. The androclads
tufted bundles
Pandanales have
the
reduced
of the
nexus
apex);
their bracts
give
to
may
Cistales,
(a
floral
(=
gonoclads
false
relatively simple
us
form
to
are
a
stegophylls;
subterminal
and
whorl of androclads
numerous
broadened floral
of
large
Theales
the anthocorms
clads and
of
each with
of which
connate
or
one
types.
became
androgynoclads)
monandra,
Clusiales,
subtending
the anthocorm axis
gonoclads
cularisation pattern enable
this
their
to
to
concrescence
of
only
gynoclads remaining;
continuation of the floral axis
forming
g.
of
gonoclads
longitudinal
into whorls, often
gonoclads
one
free and
Tiliaceae,
‘buried’
in
the
of their individual
the
Monocots, the
The conventional flower
androgynoclad.
apparently
was
the
reduction of the
FACTS
AND
gonoclads
ous,
single
to
is low,
formable
these
the
which
in the
are
and
assembly
of the older
of taxa,
most
have free
Saururaceae
deed there
and the
(and
even
perspicuity
of
fossil
tween
as
the
these
criteria
trends
Given the evidence of
essentially completed
this
period
would have survived.
shown
beyond
as
late
onward
They
reasonable
morphology (and concerning
ment
of
such
hindrance
a
general
and
to
of
as
All
morphology
can
putative evolutionary
a
and
number of
is
not
as
out,
degree
of
all
hopeless
a
that was
process
most
I believe
ideas
would
no
can
serve
to
etc.
perfectly
advanceform
by
way.
In the
a
numerous
the
the above-mentioned trends of
natural
have
Plants with
more
as
taxa
concerning
longer
qualify.
Saururaceae, but with
that
unlikely
phylogenetic
Ranmculales)
be-
living early
type of plant, in
at
a
as
the
to
relationships
these
preconceived
monogyna
traditionally
a
form of intermediate
pointed
of group I
taxa
of the
to
common
Winteraceae, Magnoliaceae, Ranunculaceae,
les, Dilleniales, Hamamelidales,
netic advancement in
be
the relative
inserted monandra and
Lactoridaceae,
and
evolution
in the
provided
Magnoliales
their recognition.
and floral
helically
archetype
groups
fossils’
exist and
doubt,
taxa,
In-
and double fertilisa-
Senonian, it would be
‘living
floral
the
obvious
Mystery’,
Angiosperm
the
as
no
the
closely.
traditionally angio-
hemi-angiospermous
‘Abominable
taxa
those of the
as
1964b). Owing
Meeuse
and
gnetalean
the quest for the ancestral,
the forms
These
rather
and
etc.,
morphology
absolute:
not
recent
groups,
such
them,
between such
sack
of the
A number
as
region.
1971a, fig. a)
Amborella,
specialisation
solution of Darwin’s
proposition.
from
are
of
emerge
The differences boil down
embryo
as
clearly
of
number
a
taxa
spicate inflorescence), approach
a
(Meeuse
Euptelea, Piperales,
early
some
true
1966, Chapter XVIII;
reproductive
the
differences
Chlamydospermae.
cycadopsid
Angiosperms,
fact
form
very few essential
almost technical details such
tion
of
con-
Some of
1968).
obsolete, however!).
are
considered to be
ancestral
putative
gymnospermous
there
a
other Dicots and of the
Chapter XIV;
1965,
group I of table ƒ,
to
there may be
the various
to
ovarial
per
3. In table I
fig.
of the
lineages
heterogene-
well be
may
1947, Kaul 1967a, b,
shown in
127
POLYCARPICAE
pistil (or
per
restriction that
anthocorms of
(conventionally
forms
spermous
Meeuse
primitive morphology
are
the
few main
a
THE
TO
organization
indicated in relation
are
belonging
gonoclads
those of the
(with
interpretations given
those
viz.,
with the
of their floral
diagramatically
to
Monocots (compare also
some
number of ovules
flower: compare Uhl
divergent specializations
ranalean
the
interpretation
trends
divergent
SPECIAL REFERENCE
WITH
monandra. The Helobiae may be
or
monogyna
the Lactoris pattem
to
floral axis
MORPHOLOGY
FLORAL
but in groups in
locule)
of
IN
FICTION
Laura-
phyloge-
following paragraphs
the
sequences will be discussed in greater detail.
(To
be
continued)