Problems
and
use
of
classification
The
diagnostic
structure
of
evaluation
of
Braun-Blanquet
V.
(State
Institute
in
the
Westhoff
( RIVON-communication
(received November
228)
nr.
11th, 1965)
Introduction
It is
of
well-known fact
a
that
quite different criteria;
satisfying
critical
way
consider
the
which
a
and
This
classification,
area
for
such
of
in the
of
systems
The
the
to
point
and
in its
Poore,
first
deals
to
sight
such
be
on
fertility,
a
as
stratification,
floristic
of
of
the
rate,
appear
and
abundance,
in which
(see
495
defined
only.
et
quite
al,
a
1951,
one,
this
a
a
1965).
since
synthetic
characters
matter
complex
to
1957;
it
viewpoint
sociability,
As
has
appear
Becking,
Against
synthetical
starting
etc.)
composition
taxa
also
artificial
an
and presence.
on
this
1964
of the floristic
dominance,
based
(1910)
From
of faithfulness is
concept
is
vegetation
differences
structure.
1951,
1959; Westhoff
may
periodicity
and
Brussels
1932,
communities
analytical
classification
an
basis
less deductive formation
homogeneous.
vegetation
that the
number
such
of
habitats,
a
vegetation which is floristically,
1928,
Castro,
system
feature
argued,
a
unit of
i.e.
several
Cain and
vegetation
this
or
or
diagnostic value
faithfulness,
one
one
a
over
higher units of this inductive
more
Congress
(1913,
the
1955;
with
may
based
of
aspect
bound
as
is,
vegetation
presenting
geographical
physiognomy
on
of
communities
of
synit
fact
taxa.
physiognomically
Braun-Blanquet
especially emphasised
At
based
of association
of
whether the
to see
International Botanical
concept
comparison
analysis
In
one.
also
a
to
involved
are
Braun-Blanquet
system
plant
is
composition,
criteria
best
and, therefore,
the
or
less
give
to
classification,
floristical
uniform
compare
the units of the
the world,
ecologically
be
to
vegetation
on
the
even
study
author’s purpose
whether
and
behaviour of
autecological
correspond
used
us
areas,
or
dynamic
scientifically
a
of this
structural
consider
not
of
the basis
on
structure,
-
The
based
reality,
Europe,
It may thus be useful
system
be
geographical
as
large
in
widely
(e.g.)
as
even
the aim
system
to
valuable
only
most
items
mapping
does
enabling
large
as
claiming
how far,
the
the
however,
and
not
starting-points.
Braun-Blanquet
article
is
taxonomy
these
system
analyse
to
in it.
of
be classified
can
by physiognomy,
composition,
by habitat. It is
-
review
is
vegetation
e.g.
floristical
processes,
it
structure
system
of Nature Conservation Research ( RIVON ), feist, the
Netherlands )
1.
the
in
vegetation.
of
one,
of
the
vitality,
of
fact,
features
496
V. WESTHOFF
and
not
on
lated
all
1913,
as
ecological
own
objection,
hypothesis
of
of the
its
more
formu-
composition represents
since each
vegetation,
amplitude,
is
however,
Braun-Blanquet,
that the floristic
states
the other characters
its
Another
one.
fundamental
early
as
single
a
The
essential.
specific
taxon
and
genetic
presents
historical
background and its characteristic dynamic behaviour. Therefore,
floristic
and
relations
be
stated
Since
be
1913,
in which
the
munities
correlated
of
the
viz.
the
be
taken into
into
these
also
(see
Cain and
The
Castro
a
hierarchical
contain
high
to
com-
mosaic
complex
a
the level
to
in
plant
floristically closely
problem of relative
a
as
more
syn-
structural
at
which
classification system
a
and
Cain
Westhoff et
1963;
and
of coordination
possibility
used here
applied
in its
al.,
has
1959;
The
1965).
original
Beard
by
a.o.
system
Castro,
sense
as
and
(1944)
(1959).
Braun-Blanquet
into
offer
to
Braun-Blanquet
1948;
“formation” is
term
the
and the
Lam,
by Grisebach (1872)
given
or
with
account
topics
system
1962; Van Donselaar,
often abused
low
be
(2)
question
here
be
to
its
cases
composition.
considered
Doing,
built
to
formation
any
two
appear
vegetations
the
posing
structure,
entering
between
of
case
has
are
independently
less
or
exceptions
ourselves
when
units
There
biology.
more
will restrict
arising
proved this hypothesis
Such
different in structure;
quite
floristic
on
Before
We
system
one.
correspondance
in
vegetation.
problem
complexity has
based
of
laws
varies
composition
artificial
have
the
chorological
Braun-Blanquet
an
claimed
most
higher vegetation
or
homogeneity,
pattern
with
case
problems.
(1)
the
ecological,
the
reason
investigations
Nevertheless
the
with
natural than
a
number of
characters
but
related,
is
more
the floristic
other
problems:
to
a
as
taxonomical
this
be
to
valuable.
exceptions,
of
be
dynamical relations. For this
may
to
would
method
is
system.
It
more
lower
an
inductive
units and
“micro-method”,
with units
operates
are
which
time
every
from
charac-
by species with larger ecological amplitudes; at the same
time, these larger units are related to more comprehensive ecosystems,
terised
structural
greater
years
(1964)
has
of vicarious
soils in
species:
in such
system.
a
Even
can
be
if
that
possibility
may lead
to more
terranean
class
stoechidis
and
shrubs in
an
A class
by
recently
group
of
taxa
Until
some
Braun-Blanquet
has few or no species
higher rank consisting
of deciduous forests on rich
group
The class
the
groups represent
in a
having a floristic basis.
zone.
one
a
system
circumscribe
in well
floristically
since
than
of
to
they fit
purely
fulfilled,
unit;
areas.
the class
temperate
desirable now,
way,
highest
groups.
e.g.
classificatory
It would be
larger distribution
it is characterised
the northern
ultimate
the
were
added class
in common, but
wish
variance and
ago classes
the
diagnostic
method based
or
units of
criteria
on
class
a
are
floristic
groups
formation
used,
this
composition
possible system. An example is the medi-
Cisto-Lavanduletea
Helianthemetalia
open habitat,
the classes
with
guttati.
with
the
The first
the second ofopen
order
order
Lavanduletalia
consists
of
low
grassland and herbaceous
PROBLEMS
AND
USE
OF
STRUCTURE
IN
THE
communities. The latter however has
common
with the order
mediterranean
T.
Trifolium striatum,
Filago
F.
gallica,
be
therefore,
class.
one
and
Central
and
possible
the
join
to
the
split
to
Vicia
former
with
the
in
1931 he
the rank of an order
It
1940 the
the
of
homogeneous and logical
more
point. The objection
and
ranean
such
but
on
a
floristic
the author does
paper
made, that
and
given
was
Wagner, 1940).
basis
themetalia and
Festuco-Sedetalia;
be
The
necessary.
how
ally
to
just
meant
to
to
little
as
a
do with
to
Quite another approach
classification
Westhoff
who
(1964),
communities
in
has
coordinating
been
tried
established
primarily
a
present
Van
an
example
floristical
Maarel
local
secondarily
of
structur-
and
der
of
system
structural,
on
this
argumentation would
structural
by
In
join the orders Helian-
certain
vegetation
a
mediter-
has.
of communities can be classified in two
group
both on the basis of a floristic method.
different
ways,
a
in
view-
undesirable,
argument
elaborate
more
only
was
structural
a
class is
one
structural
a
intend, however,
not
case
as
from
combination of
a
has
consideration
chorological
a
classifying
be
may
non-mediterranean order
a
in
Helianthemion
unit
Joining the Helianthemetalia and Festuco-Sedetalia would result
classification
the
importance of this
former
(Braun-Blanquet, Molinier
would,
from
Festuco-Sedetalia
joined the alliance
in
but
Lavanduletalia,
as
gallica,
Vulpia bromoides,
lathyroides.
the
extra-
such
Silene
Helianthemetalia
Braun-Blanquet too, has recognised
to
Europe,
subterraneum, Hypochoeris glabra,
structuraTdifference;
guttati
Western
Jasione montana, Aira caryophyllea,
in
species
dry grasslands of the
minima, Poa bulbosa and
quite
Lavanduletalia
number of faithful
a
Festuco-Sedetalia,
of
region
Helianthemum guttatum,
497
CLASSIFICATION OF VEGETATION
dune
and
plant
floristical
on
criteria.
2.
Concepts
of
structure
One of the clearest
has
vegetation
between
structure
vegetation,
(1930)
and
(1958)
are
(l.c.)
The
desert.
also
concept
limit
Dansereau
characters
concept
the
of
it
is
to
to
be
course
and
“formation
assigns
On
order
of
has
include
(1958),
exact
or
such
Schimper
types”
in
vegetational
wood,
as
sense
and
shrub,
Rubel
of Dansereau
according
various
Fosberg
to
components
been defined in
the
feature than
(1898)
the
the contrary,
Physiognomy,
objective
categories
of
classifying
distinguished
in
ways.
space.
Many
and horizontal pattern; others, e.g.
periodicity, life forms, growth forms, leaf
dispersal.
Fosberg
(l.c.)
proposes
to
distribution of
the
limit
the latter
should be
of function. The present
to structure also:
(1)
author
structure
the
biomass
spatial
vegetation
adaptative phenomena which are functional
to
functional
under the concept
to
systems
used in
who
function.
and
much less
time
stratification
structure
and to exclude all
by
and leads
structure
(1958)
on
and
the criteria
Fosberg
structure
a
The
the
and seed
of
presumed
is
physiognomical.
structure
authors
given
and function
and
swamp
space
expositions
physiognomy,
of
aspect
been
in
brought
prefers
in time
-
or
together
a
middle
(see below);
498
V.
(2)
spatial
based
pattern
WESTHOFF
the distinction of life
on
easily distinguished morphologically,
Iversen
by
or
implicit
way
practical
symbol
similar
been
used
of
Dansereau
and
Arros
of
concept
Cain
by
concept
a
of
diagrams
Dansereau
by
83
also
of such
use
A
(1936).
-
which
their
to
has
Castro
has been
structure
which
are
Raunkiaer
by
structure
and
(1958),
(1959)
forms
those defined
e.g.
in
-
an
The
(1959).
proved by the
have been
structural
applied
of
diagrams
European associations.
It has
be stressed
to
and studied
within
distinguished,
static
the
dynamic
his
this
opinion,
of
not
static
and
a
result
on
years
1964,
between
of
the
1966a,
spatial
cation
and
in
time.
in
On
opposed
A
by
the
viewpoint,
the
Ross
1964;
is
furthered
in
other
species
the basis
time
hand,
are
of these
with
to
in
more
experiences
1965;
Westhoff
to
egalisation.
time;
the
the
spatial
Differis
spatial
pattern
discontinuities
borderline
convergent
induce
reverse
differentiated
in
two
or
1963;
Maarel,
der
disturbances,
the unstable
This
ecological
on
Isolation appears
a
theory,
(1958).
isolation and communi-
homogeneity
linked
between
Westhoff,
Westhoff,
by continuity
effects
i.e.
Ashby
1962,
relations
systems
1962; Van
theory,
each other.
various
(1960,
mutual
in research
place
twenty
the
behaviour,
also Tuxen and
this
about
in
ecological
an
Adriani,
In
to
types have been discerned:
stable
between
Leeuwen
whereas communication leads
the
On
has
of form
matter
for
out
out
theory of
(see
areas
1965).
continuity
poverty
Margalef
a
quadrats
dynamic
proposed
Van Leeuwen,
have been
pattern.
flow in
viewpoint;
from another
Van
be useful in the first
to
entiation in space
and
and
and has
Leeuwen,
true:
is
connection
carried
worked
has
Van Leeuwen and
differentiation,
also
the
structure
Netherlands,
1966b)
and borderline
Westhoff
of
concept
Structure
the
and
far,
so
this
1961)
viz.
the energy
with
agree
permanent
general systems
theory has proved
and Van
of
(1958,
aspects
discussed
deals with
not
analyses
of
structure
based upon the
Westhoff,
will
elucidate
succession
pattern and process,
gradients
latter
have
structural
shall
hundreds
types
1965,
I
two
theory.
of
some
vegetation
structure
structure.
dynamic aspect
that of systems
As
The
author
with
I
energy.
the
of
only be considered
not
Margalef
aspect.
which
aspect,
dynamic
do
to
should
structure
spatial
concept
energetic.
or
hardly anything
and
the
The present
ecosystems.
in
and
compositorian
or
that
now
in its static
ecosystem
limit and the
divergent limit.
or
limit
convergent
is
easily recognisable,
characterised
sharp
by
concentration
coarse-grained
contours,
effects,
thus
patterns
and
instability. A divergent limit is characterised by effects of spreading,
thus
only
patterns
with
and
transitional
less
difficulty
stability.
disturbance
horizontal
between
zones
synonymous
recognisable,
Convergent
concepts
zone,
noise
convergence
is
ecologically
are:
are
to
be
extreme
ecotone,
environment,
often
vague contours,
limits
stress
shuttle
combined
with
fine-grained
found in unstable
situations.
More
or
tension
belt,
environment.
The
zone,
a
vertical
one
(soil
PROBLEMS
hardening).
USE
OF
STRUCTURE
limits
Divergent
with
zones,
AND
complex
I
shall
the
return
to
two
fication
3.
found in stable,
More
synonymous
Westhoff, 1964).
in
of border situations
in
structure
the
transitional
less
or
Maarel and
der
types
increasing importance of
be
structures.
(see Van
these
499
CLASSIFICATION OF VEGETATION
THE
to
are
gradient
is the concept of ecocline
IN
discussing
Braun-Blanquet classi-
system.
Relative
independence
of
layers
as
a
problem
in
classification
Different
stands
the first
was
He
polymorpha :
vegetation
the
botanist
communities.
layer
of
whereas
common,
make
to
have
may
remaining
layers
of this
use
described sixteen
and
stand),
one
formation,
place
(alternating
spp.
taken
was
stands).
in
formation
one
1929;
Katz,
Alechin,
Katz
1932).
taken into
Dutch
the
1926). In
In
account.
dunes
this
method
1927; Sukachev,
et
al.
students;
the latter
independent
and
(1934)
ones,
by
named
Du
former named them
“unions”,
Botanical
of
special
shown
Congress
of
cases
synusia
the
(e.g. by Cain
later
a
and
Castro,
in
dominants
decreasing
Cain
and
be
can
where
Castro
useful
origin.
(1952),
Certainly
habitats is of
formations
are
so
to
only
in
These
Gams
of
be
are
not
found
are
A
for
but
the
more
classification
a
regions
whose
are
studied
difference
a
a
as
the
It
with
able
complicated
however.
floras
on
are
extreme
relatively
a
dominate
to
and
structure
in
area
and
to
basis
synusial
species and
limiting;
mountainous
been
According
a
poor
critically
present
has
classifying method
are
system
between
concepts
(1918).
areas
species
practical
not
them
by
whereas
refer
they
of
vulcanic
non-extreme
it will be shown below that such twin
great importance;
bound
even
much whether it is
synusiae,
is
many
with
conditions
who
the
areas
it
(l.c.)
environmental
Roach
to
warmer
by
his
approved by the International
1959) that such
simple vegetation structure, where
developed
1936) and
Konsozionen,
is functional for northern Eurasia, viz. in boreal
alternatively;
1932,
on
(Bijhouwer,
considered
units,
(1936).
concept
vegetation
applied
(1921,
on
term
Amsterdam
of
the idea has been
Rietz
the stratal
Sozietaten,
1928;
homo-
ecology
and
structure
analyses
similarly
was
somewhat different way
a
Lippmaa
Cladina
plant sociolo-
produced the concept of
of the first
one
tree
in the other
vulgaris
logous series of twin communities, in which both
are
the
Vaccinium
and
The older school of Russian
gists further developed the method (Keller,
the Pineta
e.g.
dominant in
Calluna
by
(1881)
Hult
myrtillus and Hylocomium splendens formed the ground layer,
their
in
layers
more
formations”,
sylvestris
whereas
or
different.
phenomenon in classifying
“twin
these formations had Pinus
(combining
are
in
boreal
to
the
tropics.
possible
to
diagnostic
circumstances,
of relative
however,
on
is
but
not
the basis
of
layer independence
system.
simple example showing that in
been the main
alpine
classify vegetation
what the consequences
Braun-Blanquet
and
My viewpoint,
some
cases
structure
has
always
criterion in this otherwise floristical system,
500
is
V.
by deciduous swamp
presented
of tall
WESTHOFF
herbs derived from
woods
them after
compared with
The floristical
resemblance between certain
herb
is
one
swamps
alliance,
a
that
great,
so
drawn: these communities
that
the
Alnetum
is
still
floristical
system
floristical
the
e.g.
—
alder
and it is
area,
marshwoods
only
a
on
assigning these associations
and
canopy
which
especially
to
the
grounds that it
different classes,
to
Sissingh
as
it
Vlieger
and
subalpine
can
with
in
zone
Pinus
has
been
dealt
The
(1939).
with
viz.
by
climatic
large parts of the Alps is
cembretosum,
mugo ;
extrasilvaticum,
of
communities
association
on
in the
justified
Querco-Fagetea
i.e.
without
different
indeed.
It
with
is
Pinus
climax of the
upper
the Rhodoro-Vaccinietum.
cembra;
tree
layer.
formations
have
a
that
true
of the
case
Braun-Blanquet,
In
mugetosum,
calamagrostidetosum
this
been
therefore,
case,
into
joined
the Rhodoro-Vaccinietum
one
pinetosum
cembretosum, open mosaic communities near the timber line,
hardly be considered to be woodland; especially as the succession
tendancy
man,
of
that
Ifsuch
an
for
alliance
be
This association has been divided into four subassociations:
can
an
Molinio-Juncetea.
Rhodoro-Vaccinion,
and
of
only show small
Just the opposite solution however has been given in the
and
used
be
may
and
comparable Valeriano-Filipenduletum
structural
true
Braun-Blanquet
belonging
—,
has
yet
It is
deal with
we
stands
also,
are
Macrophorbio-Alnetum
differences with
as
Filipendulo-Geranietum
of the
developed
there
joined into
developed
or
syntaxonomists
However,
soils,
peat
Alno-Padion
but all
only optimally
purposes.
mineralised
in
that
agree
classifying
same
well
a
considerable. The difference decreases when
Alnetum-coppice;
open
between
be
author
no
different classes.
to
Valeriano-Filipenduletum
a
well
very
which
assigned
are
difference
glutinosae and
they
woods and certain
swamp
could
however,
consequence,
the stands
cutting (“Hochstaudenfluren”).
a
the subassociation
Pinus
a
cembra
well-developed
association
of its
cembretosum is
is
canopy
canopy
but
own,
hardly
be
disturbed
found
by
anywhere.
described, it might be considered
were
still
easily
so
to
as
a
of the alliance Rhodoro-
part
Vaccinion.
Similar
situations
Russia
noscandia,
timber line
sisting
and
of
mainly
Diapensia
woods
and
occur
and
are
and
the
birch-pine
Katz.
in
subartic
In
communities
of
tortuosi,
The latter
occur,
In
circumboreal
habitats
as
formations”
joined
to
the
con-
procumbens
birch
zone
Cladonio-Betuletum
e.g.
been
Loiseleuria
Fen-
near
Loiseleurio-Vaccinion,
the next lower
“twin
the
have
the
such
and of lichens.
woods
and
wind-exposed
“Spalierstraucher”
lapponica
Hylocomio-Betuletum
Hult and
found
Siberia.
tortuosi
described
the alliance
by
Phyl-
lodoco-Vaccinion, which together with the Loiseleurio-Vaccinion have been
assigned
to
the
alliances differ
vegetations
approach
to
Vaccinio-Piceetalia,
considerably.
personally
these little
nanophanerophyte
A
comparable
will
agree
floristical
carpets
although
However,
any
that
the
structures
of these
botanist who knows these
any
other
syntaxonomical
differences between woodland and
would be
difficult.
situation in Ireland has been described
by
Braun-
PROBLEMS AND
mountains
of Luzula
munity
degradation
scribed
author
community
A
less
that
agrees
similar
Their
scattered
the
logical
problem is
area.
tree
should
Kerry)
be
of
been
slopes
a
com-
considered
and
Tuxen
extrasilvaticum,
be
can
found.
a
de-
although
This
solution
Braun-Blanquet
system
other
no
practical.
or
offered
the
by
nanophanerophyte
wooded raised
as
bogs in
with
communities
in them have been described
trees
has
having visited the classical spot the present
within
would be
Co.
Braun-Blanquet
single
odd one, but after
description
which
association
and nubulous
wet
reeks,
Blechno-Quercetum
as
no
Irish
roboris-petraeae
On very
of woodland.
surroundings
an
baltic
Quercion
occurs,
501
CLASSIFICATION OF VEGETATION
(McGillycuddy
maxima
stage
this
these
seems
alliance
by them Blechno-Quercetum.
maritime
in
THE
The eu-atlantic
(1952).
woodland
deciduous
named
of
OF STRUCTURE IN
and Tuxen
Blanquet
the
USE
class
a
the
more
or
Vaccinetea
uliginosi with the order Vaccinietalia uliginosi and the alliances Betulion
Piceo-Pinion
pubescentis,
al.
et
have
(1962)
uncinatae
alliances, but
woodland
trend of
syntaxonomic
the
the
France,
studied
in
detail
subassociations,
dine
degradation
pinetosum
The
are
different
feature,
attack
in
more
In
a
a
etc.
the
the
the
In
the
Eucalyptus
joined into
(l.c.)
are
layers is
one
(Scandinavian
of
a
R.-L.
R.-L.
ericetosum
community;
R.-L.
tree
Eucalyptus
not
as
Dansereau
a
(1955).
Meyer Drees
it
of various
on
another
particular
as
the herb
was
necessary,
into
this
method). Similar patterns
one
solution
on
were
described
of Timor.
occur,
leucophloea,
layer
In
under
spite of
therefore,
association.
and
bond
consist
densities
Acacia
in the open stands.
savanna
like
(1951)
the Indonesian island
groups
same
structure
considerably
association.
found where the vertical
alba,
in
savanna
where dwarf shrubs
major diagnostic
example of classification, their
distinguish locally dominante communities
mangrove.
been
constituting
only loose and the stands
of
savannas
did
has
formation:
pine;
shrub
Guinochet
situations
community and
however
presents
structure
this
to
nearly the
difference in
great
grassland
of
number
also communities with
consider
scattered
substrate
appeared
the
Drees
the
garigue
a
mentioned,
Aleppo
independent synusiae.
savannas
certain
trees
tropics
from
the
to
case
opposed
vegetation
less
or
case
these
on
strongly
the
association
finally R.-L. schoenetosum,
In this
who
is
complex of dwarf shrub stands and open
have been
(1959),
It
dwarf
not
are
West, but equally
dwarf shrub
of
to
the
especially its association
of this
be
to
closed
being directed against
between
such
and
structures
are
Even
of
mosaic
components.
Arros
and
“savanna”
a
the above kind
example
and
(1958b).
may
from woodland
a
vegetations;
minor
good
them
joined
variance with
at
atlantic
wet
structure
four
homogeneous
a
,
helianthemetosum,
herb
which
halepensis,
multiflorae
A
region.
Barkman
by
of
the
or
Rosmarino-Ericion,
Rosmarino-Lithospermetum.
clearly
Lohmeyer
palustris.
units and
Situations of
today.
mediterranean
southern
Pino-Ledion
higher
this solution is
only found in the cool North
in
and
these
rejected
would
to
join
Meyer
prefer
the basis of constancy
recorded from Timor
502
V. WESTHOFF
Van
Donselaar
Surinam
soils
on
he
develop
to
too
a
differed
little
so
that both
layer,
tree
from
had
Meyer Drees
Contrary
to
Van
Donselaar
principle
must
preponderate
unlogical
to
prefers
try
same
respects,
is
or
biotic
even
In
humidity;
burning
the
by
of
would be
formation
a
each other.
are
In
indeed,
in
pheno-
I
So
it
for
layer
should
with
be
not
high
extremely
the
frequent
ecological
overestimated
Van Donselaar
(l.c.)
which
problem,
the present
winters,
of nutrients and
minor
a
its
on
climate the
garigue
has
(master
cold
of Timor and Surinam the
agree
as
as
does
a
to
not
principal objections against the coordinating
structural
increasing
areas
structure
abiotic
here
tree
a
bogs the deficiency
cases
that
so
of
the mediterranean
these
that I agree with his
The
influence
subalpine
all-important; in oceanic
solution
syntaxonomic
and
the
Certain
preponderate
boreal and
savannas
levels,
man.
of floristical
other words
conditions.
factors
wooded raised
on
criterion.
diagnostic
in
habitat
ecological
subarctic,
water
importance
mean
from
separate
independent
synusiae,
extreme
in the soil; in the
alternating
4.
to
and wind
cover
layer
environmental
outdo
undergrowth.
his
it
case,
(l.c.)
floristical
in different classes.
grassland
quite
and
this
Arros
the
differing little in floristical but strongly in structural
bound
and
factors)
oxygen
in
that,
system
vegetations
independent
of stands
menon
air
and
stand and
tree
association.
same
that
convinced
quite
without
vegetation
the
as
and Dansereau and
(l.c.)
is
reckoned
examples mentioned show that the problem of twin formations
The
with rather
snow
place
to
association
an
of the
system
types
of South America
part
the open herb
be
to
however,
He
vegetation
the
system of associations, alliances, order and classes. He
problem that, in some cases, the undergrowth of savanna
stands
tree
these
to
found that
a
the
met
appropriate
was
He
of
vegetation
savanna
levels.
water
first botanist in the northern
the
was
the
investigated
alternating
method
Braun-Blanquet
and
(1963)
with
systems.
weight
of
structure
in
Braun-Blanquet
the
method
Hitherto I nearly took for granted, that all layers of a stratified
community should be studied in their totality as a single phytocenosis;
the
unistratal
mentioned
also
Lippmaa and
of
concept
(see
Cain
and
Du
Castro,
Rietz
has
only
Without
1959).
been
entering
deeply into this matter, it should be remarked that this assumption
does
is
are
possible
and
by
pellier
all
cases.
This
separate
workers
Epiphyte
be
epiphyte
limit
of
(1965).
generally
it
However,
the
on
communities
exception,
an
the
communities
problem
Barkman
to
of the soil.
moreover,
cryptogamic
draw.
with
to
stand;
rogam
in
generally held
independent
rather
even
to
hold
not
1958b)
the
is
not
community
between
soil
microcenoses
always easy
epiphytic
been
is
Barkman,
their habitat
from
surface
has
(see
because
the
or
phane-
communities
often
difficult
thoroughly
dealt
Apart from such problems, Ziirich-Mont-
agree
that
synusiae rooting
together
in
the
IN
PROBLEMS AND USE OF STRUCTURE
substrate
same
There
described
have
the
be
to
however,
are,
THE
considered
also
exceptions
We
have here
which
structure
the
is
associations
they
because
mended,
much
the
of
claimed
not
are
do
to
better
be
developed
sepii
(releves)
from
are
the other
be
species which
would
Yet
special
a
the
layer,
suspended
nymphaeid
have
and
layer
been
generally
elodeid
or
the
joined
with terrestrial
vegetations is
strata
to
appear
ones; for
be
the greater part
into
and the
class Lemnetea
and Segal
of
quences
much
so
former
this
on
which
strata
only in the
communities
are
Often such
soft
of
a
factors,
etc.
A
is bound
leads
to
described
by
W.
undoubtedly such
unit like
There
Blanquet
Europe,
the
the
are,
a
in
often
mono-
of
1926,
many
more
helophyte
Montio-Cardaminetalia
clones)
in which
are
large
alternating.
as
prove
is
years,
insisting
by
Braun-Blanquet
primarily abiotic
associations
not
been
an
showing
on
within
split
be desirable in
mostly
on
wind
in the
becoming
examples
to
Scirpo-Phragmitetum,
has
to
exposed
most
about
several
yet
dominates
angustifolia
zone
association
large
recent
and
communities,
with
patterns,
Typha
brought
splitting will
indeed,
more
and
(often
deepest
Scirpo-Phragmitetum
method
is
the
distinction
Koch in
the
primary abiotic habitat differences:
to
pattern,
the
However,
pattern.
not
up
Ceratophylletea,
s.s.,
but also in that of
general analytic procedure
a
splitting
hydrophyte
mosaic
coarse
individuals
Scirpo-Phragmitetum,
that such
conse-
based
system
a
Hartog
the
important argument in its
Phragmitetalia
examples of
pattern
the lemnid
Den
1960).
structures
of the Potametea,
of
hydrophyte
the terrestrial
micro-habitat.
same
Potametea
most
changing
orders
mud, Scirpus lacustris in
and waves,
a
case
than
thus
synusiae,
of
the
fact,
followed up
composed
classes
layer,
problem which will be dealt with below.
a
masses;
clusters
instance in
method is
the
into
rapidly
the
many
monospecific
for
form
life
character
pioneer
about
specific vegetation
there
on
lemnid
or
the order Lemnetalia
Tuxen,
have
Stratiotetea and Lemnetea. An
the
brings
Not
as
In
Lemnion,
and
isoetid
or
although the analogy
live in the
(1965)
They
Previously
firstly in subdividing
alliance
Segal
development.
Potametea
Utricularietea,
is
the
should
swimming-and-rooting
the
obvious.
not
account
(Miyawaki
and
(1964)
class
favour
units:
separate
in
substrate.
species
the bottom
interdependent
they do
This fact has been taken into
strata
viz.
association,
very
less
recom-
possible
same
which
just swimming
one
not
much
hardly
fact,
although
be
not
vegetations.
water
layer,
only
into
is
to
as
communities,
water
units,
should
It
In
tropics.
not.
presented by the
of
the
Calystegion
vegetation
of
type
rooting in the
choice
and which
is
case
strata
in
(1950)
communities)
separate the blanket Hanes
to
also
subjective
a
included in the list
the different
or
be
a
synusial
are
difficulties.
analytical
Tuxen
alliance
an
of
This solution
so.
plant community.
rule.
as
503
OF VEGETATION
blanket
(liane
outpost
making sample plot analyses
The result
this
ecotones
an
Calystegion
to
of
with
total
one
to
“Schleiergesellschaften”
of river banks and similar convergent
sepii.
CLASSIFICATION
in
up,
near
this
first
but
future;
anachronism.
that
Western
structure
as
the
and
a
BraunCentral
diagnostic
504
criterion.
well
flats
This
in
as
De
been made into
Oberdorfer,
and
1965). One
the stands
the
here
careful
stand
analysis
and
far
vegetation,
called
(1964)
essential
unstable
of
analysis
a
hierarchic
said that every
two
classes.
cases
is
of
poor
relatively
difficult;
to
units”.
with
exposed
Other
water,
It
is
patterns
ecotones
convergent
are
in
common
what Pignatti
do
to
single
relation
or
irregular
to
examples
are
the
the Cakiletea of
the
Plantaginetea maioris of the border
and
wet
situation,
A
Salicornia
no
floristical
more
have
we
(brackish)
of
is
vegetation
system
This
and
is
factor
species.
stand have
have any
changes.
and the
extremes
poor in
well-developed
of time
course
and soil rich and
1962,
therophyte,
a
higher vegetation
the
and
even
Beeftink,
important
very
Spartina
communities
shore,
sea
the
untrodden,
the
i.e.
poikilohalinic
the
between
areas
of
on
of
have
they
the former
more
a
not
more
in such
fluctuations
of the
tidal marks
inflation
which in the
ecosystem
Ruppietea
that
out
us
mud
(Braun-
Spartinetea (Tuxen
1962;
stands
pure
they do
urge
that
habitat,
areas
convergent
and
“the
realise
to
However,
a
concerned).
are
will
marine
the great difference in life forms:
developed
that
so
al.,
et
as
of Salicornia
association
recently
synusiae,
two
points
well
phanerogams
pioneer
major
always
equally
an
the stands
one
more
change is
do with
to
in common,
species
(as
have
we
1936);
one.
communities
pioneer
Thero-Salicornietea and
present
geophyte
a
in
1936,
compose
Lohmeyer
1958;
for this
reason
In
northwestern-atlantic
on
to
classes,
mentioned
latter
that
stricta
Leeuw,
two
obvious
ones.
considered
and
Blanquet
(climax)
Spartina
still
were
is
development
terminal
and
europaea
of
WESTHOFF
V.
as
dry, salt and fresh,
trodden
in nutrients. In all such
simple, but
the
it
overstatement
an
cases
development
might
be
associations from this
type of environment represent
two
(See also Van der Maarel,
Westhoff and Van
Leeuwen,
1964).
In
contrast
category
be
the
stable,
divergent
fine-coarsed
gradient
and vague
pattern
stable environment where fluctuations
a
is
problem
In
such
classification.
cases
where the
drawn and which level of
basis for
the
are
with
species,
rhythmic and regular.
difficult:
to
in
living in
contours,
are
this
to
rich
communities,
As
limits
between
more
the releves
homogeneity should be
have
taken
as
a
problem, the syntaxonomy
of the woodland margin communities in particular,
the “mantles”
a
result
(if any)
analysis is
vegetation
of this
(shrub) and “skirts” (borderlines of tall herbs),
up
at
to
about
least
refining
1950.
In
and
in
Western
of
analytical
and intricate
merging
each
years
Central
of
two
other
or
Europe,
of
more
within
was
however
methods and
contact
into
later
a
somewhat
neglected
situation
changed,
this
on
special
account
of
attention
gradual
a
to
the
close
adjacent communities gradually
complicated
divergent
border
the
Braun-
pattern.
Tuxen
(1952)
made
Blanquet system in
on structure.
with
He
comparable
a
an
important
classification
development
which is
to
a
on
larger
extend based
joined the mantle communities of woodland margins
communities of shrubs
and
hedges into
one
newly
PROBLEMS AND
described
the
to
This
and
least
the
its
as
in
species
result
with
in
all
rank of
It
The
next
the
class
the
vicarious
mountains
described
Doing Kraft
account
in
Cotinetalia
sense
the
(l.c.),
-
(1930).
in the
-
and
by designating
system
In
refining
has
(1962)
the
a
quite
i.e.
new
the tall
which
of
diagnostic
by
of
formed
communities
resolving
Trifolio-Geranietea
analysis
feature.
on
He
goes
In
a
fact,
the
the
“sibljak”.
Prunetalia
two
as
in
and
the
mediter-
of Rubel
sense
(l.c.)
“macchia”
higher altitudes,
(1955)
the
completed
old
pattern
this
Muller
so-called
described
and
“Step-
developed
“Saumgesellschaften”,
or
margins of woodland mantle,
sanguinei.
stronger
the
which
“pseudo-sibljak”.
class of “Hochstaudenhalden”
herb
in
phy-
of
by giving them
at somewhat
as
syntaxonomically.
1962);
formation into
by the
divergent vegetation
by
areas,
Aestifruticeta
Adamovig
the Prunetalia
analysis
he named
refining
is
(1961)
and
name
system
Doing Kraft
sibljak.
the
coggygriae”.
Durifruticeta
types,
advanced further
penheide-problem”
sere
similar
formation
take the latter
“pseudomacchia”
between macchia and
shrub
communities,
parallel
Tuxen,
priority Rhamno-Prunetea
under
Cotinetalia
distinguished
were
plains
A
the
to
Crataego-Prunetea ; Doing
Braun-Blanquet
name
shrub
evergreen
which
to
in
continental
“
order
deciduous shrub
are
thorn
southern
the first
was
Carbonell
to
structurally
by AdamoviS (1909)
(1955)
of Rubel
ranean
and
agreed
valuable
new
the Prunetalia
Borja
name
are
that
to
the frame of the
provisionally
and
(referred
the
proposed
the
the
superseded
independently by three authors;
Goday
The Prunetalia
name.
mediterranean
first
Rivas
far
forest
with
way,
have
raise
Sambucetea. For the sake of
name
togeographically
was
criteria
to
was
Rhamno-Prunetea
himself
(1962)
the valid
adjacent)
another
be added that this renovation has been
consequence
botanists
Spanish
(often
choose
the structural
case,
must
(l.c.)
at
which
possible, because by
corresponding
however,
that
so
alliance
of their faithful and differential
part
class. This has been effected
described
(1962)
the
is
(1961)
described
edaphically
solution,
Europe and that it has induced much
over
a
Tuxen
this
ones.
research.
is
Tuxen,
that,
floristical
with
common
a
same
This
is
Querco-Fagetea.
newly
the
companion.
by Jakucs
Prunetalia
associations,
in
placed
Braun-
by
their
near
woodland
be
woodland
order
if all
avoided
assigned
was
1937
as
of the
had been left
would
tried
early
as
subdivision of the
been
shrub associations have
most
This order
spinosae.
corresponding
comparable
already been
have
505
CLASSIFICATION OF VEGETATION
admission
associations
association
associations.
the
could
and mantle
shrub
THE
generally accepted
chorologically
each
has
the
to
IN
described
The
Vlieger.
consequence
shrub
STRUCTURE
Querco-Fagetea,
and
Blanquet
OF
named Prunetalia
order,
class
contrary
USE
Along with the further
accentuation of
the alliances
and
structure
associations
all
as
a
newly
described
been
even
by Muller (l.c.) within this class Trifolio-Geranietea have
faintly characterized
they hardly present any faithful or
that
good differential species with a comforting selectivity
so
—
—
nobody would have brought them into
arguments
were
available.
This
a
separate
weak
class if
floristic
no
structural
characterization,
506
V. WESTHOFF
nothing
has
however,
communities
terization in
within
the
this
the
opposite
most
is
a
case
of
rate
these
problem
of charac-
the
on
the
small.
always
limit
a
In
of
in
species
great stability
a
on
mile, and
quadrat
a
is
species
every
of
the
rhythmic
ecosystem
not
As
Blanquet
being
is
the
in
An
in
5.1.
Braun-Blanquet:
only good and bad ones”.
at
twenty
European
classifying
onomic
the
this
(1962);
has
of the
survey
alpine
structural
sake of
units
no
classes
and
right
occurring
units
himself here
class
names; references
to
1963),
(1963).
The
as
know,
most
a
have
may
completely
recent
by
syntax-
Lohmeyer et
Ellenberg
to
(1963)
al.
and
classify the units of this system into
is
attempt
are
Dutch
presented
given,
this
region.
are
(l.c.)
relating
to
to
of
The author
prefers
non-alpine
authors’
been
provided
these classes
Doing
(1964),
may
Kraft
Segal
is
not
W.
and
(l.c.).
with
should
names
avoid confusion,
and Oberdorfer
the
highest
also found in the surveys
have
(1962),
Segal
desirable
For
the
are
This restriction
vegetation.
the Netherlands. As
are
below.
they
as
the bulk of the classes
names
Doing
Hartog and
try
An
they
Ellenberg
deviating
hoff
to
in
have been omitted if
(1962,
I
as
by
over
described in
Europe occur
only be added when such
Den
published.
been taken
big disadvantage, since
(l.c.),
chapters.
only the classes
which have been
al.
changed,
As far
areas) has been published
classification.
surveyability
restrict
considerably
(1964).
Central
or
vegetation
We
pragmatism.
are
of
important
vegetation of Northwestern and central Europe
It seemed worth while
et
most
Braun-Blanquet system of classification
been
not
has
survey
Oberdorfer
the
has
preceding
survey
(except for
a
the
structural
into
years,
vegetation
in the
up-to-date
to
to
have
Braun-
Introduction
appeared
a
dogmatism
suitable
we
possibilities
more
The
types.
“There
Netherlands
In the last
of
from
shifting
and
flexibility
ecosystem
by very
the
provides
structure
with
attempt
the
of
use
greater
a
different
gradual
conclude
wrong systems,
5.
increasing
with
system
applied
point
may
the
species
tree
represented
is
As yet, the Braun-Blanquet
system
community type. This is, however, exceptional.
above,
just
divergent
on a
few individuals.
shown
for
but
for
this
the
too,
case,
of energy flow within internal
with
relative
pioneer communities.
in
case
this
possibility,
whole, is the tropical rain forest, where hundreds of
a
occur
contours
vague
i.e.
very
given community compared with
a
to
species:
optimum,
phytocenosis rich
coincides
dynamic processes
The
dependent
those of the
a
maximal
in
poverty
species.
presence
in
attains
from
reasons
extreme
habitat, where
as
the
species
system
in
directly
pattern:
any
with
rich
communities,
adjacent
Braun-Blanquet
The
is
case
mosaic
accumulation of
the
do
to
extremely
are
such
names
of Lohmeyer
More
their
recent
authors’
be found in Beeftink
and
(1965),
Westhoff
(1959),
Tüxen and West-
PROBLEMS AND
An
has
attempt
below in
USE OF STRUCTURE
been
of
terms
factors
(Edaphical
possible
A
It
quite
The scheme is
be
practical
Scheme
5.2.
1)
lump
to
others
some
up
meant
be
to
5
2,
(e.g.
and
an
Classes:
of
Lemnetea
(J.
2)
habitat and
aquatic
-
Tx.,
3)
species
(on
of rocks
and
14),
assigned
to
to
split
to
system.
them
amphiphytes
(grow-
hardly emerging from it).
or
1964)
Hartog
with
et
Potametea
-
Segal,
haloid soils
1964)
up
Ruppietea
-
Zosteretea
-
character,
from
consisting
single
a
rich in ammonium
and
to
many
nitrates,
(or)
habitats).
Cakiletea
-
Chenopodietea
—
pioneer
a
built
therophytes,
Thero-Salicornietea
in
“formations”.
and walls.
vegetations
open,
Pioneer
4)
been avoided.
might appear
and
and
not
way.
better-balanced
a
hydrophytes
yet, it is
as
as
It
are
only
rupestris.
summer
juncetea
poor
them
the 37 classes
(Krausch,
nearly always in convergent
Classes:
get
only.
The
ecological
here has
13
given
Littorelletea.
Ephemeral
of
units
spectra
classification
walls;
only.
12,
to
not
(Den
1961)
Asplenietea
mainly
Charetea
-
Vegetations
Class:
6),
of structural units with
Vegetations consisting
ing in
provisional
(e.g.
this
an
design
to
units
some
and
than in
presented
appropriate
form
understanding).
of rocks
units
for
better
otherwise
the structural
not
life
507
OF VEGETATION
the structural
and
essential
to
2, vegetations
group
for
term
seemed
aid
an
define this unit
to
define
to
not
are
as
THE CLASSIFICATION
arrangement
which
only given in brackets
exception is
made
spatial
IN
Bidentetea
-
Isoeto-Nano-
-
Secalinetea.
-
vegetations consisting mainly of rhizome geophytes
species
marine
(on
and
mudflats
coastal
the
ridge
and
of
sea
dunes).
Classes:
5)
of
Spartinelea
More
several
less
or
species,
hemicryptophyta
disturbed soils
Classes:
made
Epilobietea
in
et
nitrogen, always
in
maioris
Plantaginetea
-
of
up
scandentia
rosulata,
rich
instable
secondary,
open,
mainly
Communities
6)
Ammophiletea.
-
consisting mainly
of
vegetations,
perennial
scaposa
(on
more
Artemisietea
helophytes
chiefly
less
or
habitats).
convergent
—
consisting
plants:
(in
vulgaris.
and
swamps
springs).
Phragmitetea- Scheuchzerio-Caricetea nigrae (syn.: Parvo-caricetea)
Classes;
- Montio-Cardaminetea.
Closed
7)
and open
hemicryptophytes
tosa
—,
winter
(on
a
annuals
-,
communities
grassland
the bulk of them
considerable
dry soils and
Classes:
-
number
and
on
next,
the
of
if
ecotone
Sedo-Scleranthetea
-
Saginetea maritimae
therophytes
any,
-
geophytes
between
et
for
the
and
caespi-
larger
part
chamaephytes
dry and salt habitat).
Festuco-Brometea
(Tuxen
mainly of
consisting
being hemicryptophyta
-
Violetea
Westhoff,
calaminariae
1963).
-
508
v. westhoff
Closed
8)
of
caespitosa
the
damp
Classes:
group
to
wet,
Closed,
herbs,
mostly
fresh
with
ous
tripolium (Beeftink
shrubs
more
and
—
raised
sometimes
therophytes
open
1947)
communities
1960).
of
tall
intricate
an
perennial
and
chamaephytes
scaposa,
mosaic
habitat,
ecocline
divergent
Arrhenatheretea
-
Hanes;
pattern
mostly
on
sanguinei.
with
nanophanerophytes
predominating
mostly conspicu-
a
tussocks
bryophytes:
of
layer
—
of
heaths and moorlands.
bogs;
Ombro-Sphagnetea
Shrubs
11)
differing from
Westhoff,
et
forming
(in
Trifolio-Geranietea
Classes:
but
absence of
or
mainly
hemicryptophyta
soils).
Communities of
10)
content
(Braun-Blanquet,
shrub communities
Class:
mostly
soils).
hemicryptophyta
calcareous
dry
salt
or
sometimes
bordering
of them
mainly chamaephytes,
Molinio-Juncetea
9)
bulk
by the smaller
- Asteretea
often
the
—
exceptionally
previous
(on
grassland communities consisting
(rarely open)
hemicryptophytes
or
Nardo-Callunetea.
-
Fruticeta
(on dry
to
soils, mostly in divergent
wet
habitats).
Classes:
Rhamno-Prunetea
12)
stratified
Poorly
growth
in
poor
in the
often
chamaephytes
rather
(1936);
rich
(azonal
Borja-Carbonell,
et
Salicetea
-
especially
always with
of Iversen
sense
(though
1962)
deciduous forests
species,
but
phanerophytes,
(Rivas Goday
(Doing,
Franguletea
in
poor
certain
a
moss
with
(but
in
amount
swamp woodlands
of
and
not
and
nano-
telmatophytes
very
conspicuous
of
consisting
or
under-
Hanes);
therophytes
layer either
species)
1961);
purpureae.
“carr”
sphagnoid
on
wet
peat
soils).
Class:
Alnetea
Poorly stratified deciduous forests (but with Hanes); in boreal,
13)
baltic
in
and
species,
even
glutinosae.
without any
dominant
woodlands
Class:
Querco-Piceetea
on
roboris-petreae
14)
Richly
of
and
Class:
very
et
moss
acid,
dry
with abundant
layer
often
or
conspicuous
damp mineral soils).
to
Westhoff,
Vaccinio-Piceetea
poor
often
1959)
(syn.:
Quercetea
lumped together).
undergrowth rich in species and
forests;
forms, but rarely with a conspicuous amount
stratified
nanophanerophytes
or
poor,
(Doing
of many life
mineral soils
but
telmatophytes,
nanophanerophytes;
(azonal
consisting
Undergrowth
Europe also acicular forests.
montane
(zonal
woodlands
mineralised peat
soils
with
rich,
on
a
crumb
dry
to
damp,
structure).
Querco-Fagetea.
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