The
descent
the
in
of
light
the
of
flowering
new
phytochemistry
evidence
and
from
plants
from
other
sources.
II
Suggestions
major
A.D.J.
Hugo
a
for
holotaxonomic
classification
Meeuse
de Vries-laboratorium
from Ada
(Continued
en Hortus
Bot.
Neerl.
19:
Amsterdam.
Botanicus,
61-72, 1970)
SUMMARY
In
this
second
cation
4.
of the
and
final
based
on a
PARALLELISMS
POSSIBLE
Parallelisms and
the
suggestions
part
Angiosperms
Angiosperms
may
mentioned sub. 2.1.
Analogies
in
extensive
In other instances
of
rence
such
as
and
with,
between
grains
a
the
also
the
in
by
a
reduction of
aril
(or
the
both Dicots and
Acta Bot.
Neerl.
of
characters
course
and
predominantly
some
fruit
doubt
structures),
dry
and of
are
or
Liliiflorae).
a
occur-
parallelism,
or
to
than is
taxa
clearly
fleshy,
methods of
“adap-
dehiscent
dispersal (asso-
of seed
taxa
appendages),
the
a
to
primitive
Some
trinucleate
pollen
(all Helobiae, Lemnaceae,
attributed
course
of
occurrence
general
zygomorphy,
than otherwise
of
so
account
suspect. The remarkable correlation
the
on
the
major
which
( e.g
fruits),
aquatic
be
may
number of stamens,
convergently
between
of edible parts of the fruit
are
and
not
are
into
of this condition and believe it
affinity
many-seeded
features
the other. The
outcome
.,
environment
cast
or
way
of
concensus
taken
are
of
phylogeny
a
phytochemical
Rosiflorae
one
origin
number of taxonomists.
of arilloid
developed
the
of seed and
Callitrichaceae)
would, in any event,
Helobiae
or
presence
special
characters
of the
morphological
hypogyny,
number of
Hippuridaceae,
of
decide
always
concerning
peculiarities
classifi-
major
characteristics
morphological
among
the
apparently
of the
occurrence
numbers
not
during
in several orders may be the
one-seeded
e.g.,
perhaps
ideas
of
but there is
compilations
can
Certain
may
indéhiscent,
ciated
holotaxonomic
a
CONVERGENCIES
indicative of taxonomic
assumed.
usually
or
own
my
clearly
more
tive’,
one
obdiplostemony
but I have
be
homology
if considerable
Huber’s
AND
anticipated,
be
a
(as
towards
convergent evolution of characters
opinion concerning
troublesome
made
are
pleiophyletic origin.
but
assigned
the
“trends”,
the
sympetaly,
(zygomorphy
convergence,
status
complete
are
and
more
such
loss
likely
sympetaly
to
as
the
of the
to
have
occur
in
Monocots).
19(2), April
1970
133
A.
The characters
fully
chosen
suspected
used
the
by
phanerogamists,
taxonomic
as
if
present author,
based
are
not
more
less
or
similarity”
5-7
paragraphs
generally
MEEUSE
are
care-
accepted
from the
apart
J.
between orders
difficulty. Relationships
“overall
on
in the
pointers
avoid this kind of
to
so as
D.
by
de-
more
monstrative (e.g., chemical) evidence.
5.
MAJOR
GROUPS
AND
LINEAGES
The conventional classification of the
and the
Monocotyledons
affinities of
groups and
monocotyledonous
groups,
(and
ranalean
all)
of the
respect
There
respectively.
Dioscoreales
of
(or
does
certain
Liliiflorae)
the
groups, whereas
orders
liliiflorous
dicotyledonous
the
(Huber
major angiospermous
groups is
1965, Chapter XIV)
(Meeuse
system
and
postulated,
of the
analysis
large
from
may
taxon
a
for
must
have existed side
and
classification,
several other
Kubitzki
is
clearly
see,
e.g.,
(such
concerning
and
on
‘Handles-branch’
true as
Hegnauer
(for
a
134
a
all
a
linking
and
the
(1959,
1969), have
the
on
186)
p.
least
at
but
groups
two
(1967)
the
is
as
often
so
of
systems
who
enumerates
Lanjouwc.j.
(1968),
large
primitive”)
current
the
Soo
1969),
and
lineage.
affinity
(Huber
even
but
least “most
at
of
Thorne
Dilleniales,
who
had
Kubitzki
(1968).
1963,
also
the
as
is,
types being
out
Pittosporales
I believe,
Hallier
out
not
that
so
and
(1912)
there
are
Ranunculales and Rutaceae,
Hegnauer
Compositae.
their early
postulated
1966), pointed
pointed
expressed
evidence considered the
such
between
several
quoting
previously
However, this
workers,
Nooteboom
Rutales and
possibly
liliiflorous line
a
greater,
even
(or
majority
of certain alkaloid
Swain
be
mainly phytochemical
see
between this
affinity
of
was
Huber’s
principle.
findings,
angiospermous)
the
relationships.
taxonomic
lines
time.
long
Hegnauer)
dead-end
Several workers
in Harborne &
Araliaceae,
their
discussion,
the presence
significant.
relations
be
to
a
Polycarpicae
the basis of
Huber’s
may
Kubitzki suggests. Several
clear indications of
especially
of
close
a
in
two
rather fundamental differences between the
out some
Melchior and
as
reveal
“basic”
examples, Cronquist (1968),
views
divergence,
in
Takhtajan
morphology
authorities
lineages
even
maintain this
to
certain
even
forecast
previous
a
I assumed in 1965 (fig. 20
as
the
isolated
evolutionary
“early” separation
side for
not
reflected
(1969) pointed
androecial
is
essentially
dicotyledonous (or
group of all
assumed
one,
by
number of
the
assembly
not
An
conformable
sufficiently
be
Dicots
the
strictly
did
instance, palms.
number of
to
reason
every
liliiflorous assembly
and,
even
ranalean
similar
is
spadiciflorous-pandanalean
not
lineages
In
there
of
A dismemberment of the
unavoidable. In
a
between
relationship
very
closer
dicotyledonous
precursors
are
certain
be
groups may
the
Helobiae
ultimately
between
the various
and
1969).
of
a
or
number of indications of a
a
hence:
taxa, the
major
two
Relationships
monocotyledonous
are
in
agree well with the natural
not
of their subordinate groups.
some
than those within the various
early
Plants
Flowering
Dictotyledons,
considered
on
several
that there
with
the
The overall
to
be rather
occasions,
are
other
e.g..
likely
Umbelliferae
similarities do
Acta Bot. Neerl.
19(2), April
and
not
1970
DESCENT OF
Fig.
1.
FLOWERING PLANTS AND PHYTOCHEMISTRY II
Diagrammatic representation
sperms.
Acta
Bot. Need.
of the
putative major evolutionary lines
of the
Angio-
(Centrospermae omitted).
19(2), April
1970
135
A.
plead
against
these
points quite clearly
families
“Cor nales”)
groups
and
the
regard,
I
with
of coumarins in
positae,
knit
and the
these
taxa
Harborne &
a
ellagic
pointers
of
together
(compare
the
as
Hegnauer
postulated by,
positive
including
Umbelliferae
ones:
p.
the pres-
Rutaceae and Com-
in Araliales and
1964,
these
“negative”
the
544, fig.
Asterales,
and
29,
in
132, fig. 2).
are
rather isolated and this is reflected
Angiosperms.
Meeuse
e.g.,
of
and
other
between
Araliaceae and
some
acetylenic compounds
Centrospermae
the absence
are
well
Cornaceae and
convincing;
Rulaceae,
as
MEEUSE
Umbelliferae (which
plus
similarities
chemical
number of these families
Swain 1969, p.
is
group
Araliaceae
highly
J.
Huber’s detailed analysis
contrary;
iridoid-containing
acid in
substances,
several classifications of the
this
the
The
as
presence
In several respects
in
the
Hegnauer,
and the absence of iridoid
ence
with
Pittosporaceae.
the absence of
pointers, i.e.,
on
relation between
a
associated
not
are
assumptions,
to
D.
ellagic acid,
A
(1965)
iridoid
long independent
and
Kubitzki.
compounds
existence of
Phytochemical
and characteristic
alkaloids.
Fig.
2.
Putative
N.B.
relationships
“Woody
in
the
Polycarpicae
include
Polycarpicae”
and their
probable
Magnoliaceae s.l.,
derivatives.
Winteraceae,
Schizandraceae, Trochodendrales, Annonaceae, Myristicaceae , Laurales,
ber
of
small
Berberidaceae,
a
few
on
families;
Lardizabalaceae
associated
dubious
136
associated
,
taxa.
account
-
Menispermaceae, Ranunculaceae,
Position
of presence
“Ranunculean-Berberidalean
of
Nymphaeales
s.s.
(not
Iliciaceae,
and
assembly’
a
num-
includes
Nelumbonaceae
included)
more
or
and
less
of ellagitannins.
Acta Bot.
Need.
19(2), April
1970
DESCENT OF
A
FLOWERING PLANTS AND PHYTOCHEMISTRY II
considered
major lineage,
assembly
the
Rosiflorae,
sarily
is
related
closely
line which
one
the
phytochemical
includes
the
Cornales
iridoid and
in
Gentianales,
also
Rubiales,
here
belong
referred
The
or
same,
more
Jay
the
(Huber
may
one
related
be close
to
another
There
general
a
to
the
orders
(Ebenales),
Styracales
The
Celastrales
Aquifoliaceae, often
genus of the celastralean
relations
with
the
Primulales.
line led
saxifragaceous-rosaceous
to
neces-
out.
with
divergent, evolutionary
Rosaceae,
s.s.,
this
Plantaginales.
have
that the
not
sympetalous
includes the
least
s.s.; at
early
Amentiflorae,
Saxifragaceae
as
and
(phytochemically
and
1968)
in the Cornales
an
Ericales,
out
are
sense,
number of
a
Tubiflorae,
least related
probably
to
constituents:
Dipsacales,
iridoids)
restricted
a
this line is believed
have led
to
related
at
are
contains
Hamamelidales
such
or
Celastrales,
to
Icacinaceae
in
bear
patterns
ranalean
heterogeneous
pointed
several groups which
includes
and
and
of counterpart of the
includes the rather
it,
the Parietales. Huber has
assembly
guttiferalean-dillenialean
rich
kind
a
of iridoid substances and
representation
also
and
Guttiferales
conventional rosalean
be
to
derived from
directly
not
and
(or several)
to
the
groups:
noniridoid families
to
and
Chrysobalanaceae
perhaps
the
Leguminosae.
The
nexus,
remaining
and
a
major
number of
presumably
Capparidales, Passiflorales, etc.)
talous
one
the
assembly
upon the
and,
of the
characterised
whole, general
lines is
previous
the
TAXONOMIC POSITION OF
A
number of
taxa
render the
of
as
a
absence
complete
ellagic
Violales,
predominantly choripe-
acid.
of the
iridoids
A close relation with
the relation Actinidiaceae
glycosides
SOME
(Myrtales, Malvales,
regarded
almost
suggested by
6.
characters
be
must
by
dillenialean-guttiferalean-parietalean
derived
occurrence
of aucubin-like
occurrence
the
branch,
Ericales and
-
in Actinidia.
SMALLER
GROUPS
following
taxonomic
conclusions
highly
probable;
Araliaceae and
and
with
Pittosporaceae
Sapindales
the
and
are
relation is
origin
approximate
Compositae
Polemoniales)
as
previously
Rutales
->
discussed,
Araliales
->■
are
related
Asterales
Rutales
to
lineage, together
Campanulales.
independent
Sapindales
to
and
->
be associated with the Rutales and
may
possible
related
this
Umbelliferae,
in the Ranales
belong
least
at
accepted
developed
the
of
Leguminosae
according
according
to
to
of
the
in this
case,
Rosaceae.
Huber.
their
Boraginaceae (and
independently
would,
and
point
to
Comaraceae
Boraginaceae
phytochemical
be
may
patterns;
if
possibly
all
the
non-iridoid
majority
of
the
tubiflorous
Sympetalae.
Papaverales
the
are
close
old Rhoeadales
Ranmculales
to
(=
Capparidales
s.s.
( Berberidales),
sensu
Takhtajan)
but
the
is allied
remainder of
to
the
Cistales
(Parietales).
The order
the
family
Nymphaeales,
in the
Nelumbonaceae
Acta Bot. Need.
19(2), April
(the
1970
usual
genus
circumscription,
Nelumbo)
is
heterogeneous
is rather
aberrant
and,
in that
if
not
137
A.
far
removed,
related
distantly
only
in
Nymphaeaceae. Nelumbo deviates
Cabombaceae,
to
and
phin-
aporphin-type
that the
non-alkaloid and
of
in the
place
Nymphaeales
“
name
the
(minus
s.s.
Cornales
and
from
This
characters
also
(the
the
is
in
Styraceae,
to
exclude
their
and
position
of the
related
in
with
Geraniales is
Ebenales
not
the
substances),
including
'Grossulariaceae
and
Dia-
chemodiagnostic
but it is
for this
ebenaceous families
and
include
to
heterosides
(iridoid
not
also
(and
Garryaceae
The
present).
in any event,
are,
to a
sense
Huber
Aquifoliaceae
conventionally
(some)
to
(but
Symplocaceae,
quite
close.
but the absence of alkaloids and
clear,
render
trihydroxylated phenolic compouds
GENERAL PICTURE
out
origin
restricted
a
according
Diapensiaceae ?)
Alangiaceae
with Ranales-Rutales rather
is
for the
place
independent
in
applied
and
s.s.
good agreement
the
ericaceous
with Ericales and
the presence of
7.
Araliales
reasonably
better
present)
relationships
The
s.s.!),
presumably
(aucubin
suggest
but
“better”
a
highly probable.
presence of aucubin and
perhaps
reason
is
believes
(1968 a)
family Nymphaeaceae
Cornaceae) Escalloniaceae, Philadelphaceae
Saxifragaceae
pensiaceae.
not
should henceforth be
families without the
group of
(apart
I dare
Nelumbonales),
assembly
”
Bate-Smith
ellagic acid-containing
Polycarpicae.
of the ranalean
respect
The
order
and
and chemical
the presence of proapor-
by
alkaloids.
isoquinoline
J. MEEUSE
Ceratophyllaceae
morphological, palynological
affinities with the
Polycarpicae
respects and shows
D.
direct
a
relationship
improbable.
AND
SOME DETAILS OF
THE
HOLOTAXONOMIC
CLASSIFICATION
The
to
major lineages
tentatively depicted
are
least
Monocotyledons (at
orders),
three,
and ranalean
Piperales
to
two or
here
one
and
orders,
(see figs.
of them
to
lean
(clusialean,
dent”,
s.s.,
or
at
relative
is of
sequencies
another
and
of
age
parietalean
lines of descent
and
of
origin
1).
of
these
course
Some
(cistalean)
leading
Scitamineae
perhaps
we must
(see fig.
and
quite conjectural
course
representation
one
dubious,
Centrospermae
The
tic
thealean)
least
are
lineages
and is
place
the
contrasting
close
2)
to
assemblies.
Helobiae,
of closest
lineages
leading
to
phytochemical
“Indepen-
Nymphaeales
here.
independent
considered. In
not
leading
dillenialean-guttifera-
disregarded
as
as
proranalean
to
group of lines
a
rosalean-hamamelidalean, cornalean-saxifragalean
1 and
fairly
phylogenetic
a
diagramma-
affinity
next to
characteristics
are
shown.
Of
in
the
fig. 1,
individual
but
derivatives
ingoff”
an
(see fig. 2;
of the
assessment
under
this
of their overall
5)
and
agree
sufficiently
are
be made for the
dendrogram
mere
is in
so
shown in essential
Polycarpicae
and their
far fictitious that the
guesswork
although
it
is
detail
putative
“branch-
based
on
an
similarities).
in several essential
of Kubitzki
dismemberment of the
138
must
various groups is
The conclusions
(see
lineages several
exception
Dicots,
(1969,
p.
but there
are
with those of
respects
366,
Abb.
also
4) concerning
some
Hegnauer
the
major
discrepancies, especially
Acta Bot.
Neerl.
19(2), April
1970
DESCENT OF
with
FLOWERING PLANTS AND PHYTOCHEMISTRY
regard
the
to
overall
picture
genetic
lines
in
ranalean branch.
the
details,
to me
seems
to
II
Although
be
evidence may
new
of
early divergence
least
at
three
the
change
four
or
phylo-
established.
firmly
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