Wentia
15
25-46
(1966)
relation
A
theoretical
and
pattern
Chr.G.
(RIVON
State Institute
,
approach to
in
process
vegetation
van Leeuwen
for Nature Conservation Research, Zeist)
(RIVON-communication
219)
nr.
(received July 15th, 1965)
CONTENTS
Abstract
25
Introduction
1.
1.1.
Cybernetics and ecology
1.2.
Elements
of
spatial patterns
27
1.3.
Elements
of
temporal patterns
28
The theory
2.
25
of basic
2.1.
Starting point
2.2.
Basic relation
2.3.
Spatial
2.4.
Temporal
2.5.
Basic relation
2.6.
Basic
28
of
equality (1st
basic
and
basic relation
relations
of
relation
Paradox
2.7.
the first
and
Concentration
4.
Some
5.
The
the first
between
and
29
relation)
basic
29
relation
31
inequality (2nd basic relation)
and
connection
separation
of the second
3.
28
relations
relations
connection
26
in
time
(3rd
32
separation
basic
in
and
space
33
relation)
34
basic relation
and dispersion in
35
space and time
36
properties of border areas
theory
of basic
relations applied
to
40
ecology
Acknowledgements
46
References
46
Abstract
A theoretical approach
and
their
based
on
Three
third
to the
problems concerning
environment, methodically allied
researches
made
fundamental
of which is
into
plant
relations
showing
the
character
connection
with
and
also provide an approach to the effects
time, which phenomena are considered as
regard
The
to
pattern and
relations
proved
to be
relations
an
described
Many examples
1.
with
concentration
expedient for getting
in border areas, which have
respectively
are
as
of
between
plants
and
cybernetics
distinguished,
been
pattern
and
of concentration
to be of
the
process. The three
and
dispersion
in
dominant importance
process.
associated
useful
have
between
basic relations
space
the relations
discipline
succession.
of universal
the basic
with
“limes
given
a
and dispersion
better
been divided
in two main
convergens” (ecotone)
with
regard
to the
on
understanding
and
“limes
application
their
turn
of the
ecological
types
of
have
boundaries,
divergens” ( ecocline).
of the
theory.
Introduction
During
has
the last
two
decades the botanical section
regularly investigated
over
the
annum.
whole country,
The results
more
with
of this
than 500 permanent
a
of
our
Institute
plots, distributed
frequency of about 200 samples per
yearly repeated
25
examination into
a
large
26
CHR.
of
number
vegetations
alteration of their
are
mainly used
In
to
pioneers in
the field of
ecology
and
is
attention
1.1.
LEEUWEN
regard
the
to
and
composition
alteration
other
directives for the control of nature
as
theory, founded
with
types
floristical
connection
other
G. VAN
the
on
given by
cybernetics
the
to
W.
Ross
reserves.
Ashby
able
we were
non-
patterns
to
(1956)
develop
and
a new
first mentioned and directed
investigations
conservation.
nature
paid
ideas
some
or
spatial
this
In
environmental
“Relation
conditions
Theory
”
to
much
in border
areas.
Cybernetics and ecology
Cybernetics has been started by N. Wiener during World War II as a
special scientific discipline with respect to the construction of machines
for fire-control.
also
Soon however he
be
successfully
applied
discovered that its
the
to
problems
and their internal and external vital functions. Since the
approach in biology and allied subjects, leaning
“
life
”
“
and
All
regulation
stitutes
ments
3)
for
research
central
ones
between
in
idea
stability
(
=
in
the
organisms
cybernetical
hypothesis
time.
are
These
change
(
=
basic
possibility
turned
primary elements,
four
con-
ele-
variety) in
inequality (difference,
time); 4)
the
to
The
mutual relations
cybernetics.
that
increased.
steadily
reduced
and
four
only
modern
equality
be
can
in space
inequality
1) equality in space; 2)
are:
space;
simple
more
the
or
has
allied,
by which complicated
reduction,
into the
closely
are
suitable
questions
problems of equality
of this
”
on
could
principles
concerning
inequality
in
time).
When
equality is seen as a special case of inequality (difference
zero) then we may agree with the assertion of Ross Ashby (1956) that
=
“
difference
two
”
is the
things
most
fundamental
in
concept
different
recognizably
are
or
that
cybernetics, either that
one
has
thing
changed
with time.
Viewed in
has
this
other
no
“
variety-in-space
”
communities. It
ed
with
the
even
the
such like
work of Nature
tracing
may
“
and
the
mutual
ecology
questions
same
regulation,
in
we meet
again
already
vegetation
on
between
within
the whole is
plant
concern-
does
as
selection and
(conservation)
process
connections
variety-in-time" (process)
be said that
Conservancy
itself
name
than
(pattern)
exactly
problems
light the study of pattern and
aim
cybernetics, studying
dynamic balance. In the
with all these
expressing
the
phenomena,
element
of
non-alteration.
Within
mation
the
It has been
better
cybernetics
developed by
Several
tried to
one
the
technical-mathematical Infor-
of its
best elaborated foundations.
chiefly in order
possibilities and difficulties linked
to
up
get
a
with
(carrying messages).
ecologists,
apply
as
C. E. Shannon a.o.,
of the
apprehension
communication
But
of
scope
Theory is considered
this
most
of them
working in U.S.A., have already
quantitative theory
nearly all publications
that have
on
their
appeared
study of ecosystems.
up
till
now
are
quite
PATTERN
in
the strain
or
“production",
of “biomass and energy
“
of
not
of
as
we
know
only
spatial patterns
of
degree
Margalef
two
of
types
the sake
of
(1957, 1961)
and
the size
of
this
we
to
suppose
ecology,
simply,
the
not
as
we
1.2.
example lime
gradual
is
Ecology
primarily
In the Relation
possible
not
are
able
in
we
theory
to
their
as
or
single place,
a
line
and
is
the
as
This
by
that
plants
and
temporal patterns,
the fact
that
an
organism
limitation or
spatial
restric-
of distribution
which
up
patterns,
patterns
and “no-no"
is
and
limits
are
equality
or
only present
within the
available. For,
the difference
same
way choice
thus with
measuring
non-pattern
is
two
found,
distribution
roundish
be
area
patch
large
or
gradual.
or
expressed
narrow
relatively
inequality
or
or
selection
using “no". One of the
the
amount
we
inequality between “yes"
is
connected with the
purposes
of information
or
of Shannon’s
selection
as
a
weighed quantity of such alternatives.
is
showing
3) border type.
1) form-, 2) size;
be
in
as
available!lime
the fact that life circum-
partly by
“no"
Every spatial pattern
A
on
time “lime
the
spatial
well
building
“choice",
“no”, expressing
statistically
can
cybernetical approach
a
couple giving “yes". Using the alternative “yes-no" however,
“no". In
sharp
taking
on
and pregnant
“yes-yes"
couple “yes-no",
may
suited
in environmental factors
and
space
are
react
equal everywhere
“inequality",
shall find
a
more
does
dealed
to trace.
combinations
viz.
System
than
Theory
addition still
of the soil
content
live under all circumstances.
are
and
an
that, applying
Now, generally spoken, spatial
either
General
of spatial patterns
tion offers the base
when
also
by the various environmental factors.
Elements
stances
we
succession and
This author
starting-point
better
a
have found
contrasts
Consequently
organisms
cannot
the
out
importan-
spatial patterns, particularly
much interested
so
or
available".
build up
not
are
for
sharp
other
the
purpose.
It should be borne in mind
to
1959;
“grains".
highly specialized Information Theory.
with hereafter
pointed
to
Ashby’s many-sided
give
to
seems
regard
ecosystem.
an
of its
Ross
has
approach, showing
in
ecology
Theory provisionally
for
’
of “matter and energy
(compare Patten,
differentiation with
or
or
“granulation"
For
thus
,
process"
stability
regulation
gives fruitful ideas about some qualities
about
27
VEGETATION
flow"
and
pattern
difference between these
ce
IN
PROCESS
1963).
Galoux,
So far
AND
of
or
For
mutually
example
related
1)
the
plant species (a spatial pattern)
look
small;
The
like
3)
a
its
narrow
external
ribbon;
and
2)
may represent
its
internal
properties,
geographical
extensiveness
limits
mutual relations between these three
by using
ribbon
large
a
or
three
means
the
alternative
that in the
quantity of “yes"
is
be
“yes-no".
longitudinal
available,
selectivity. Following this axis
may
qualities
we
so
axis
a
shall
of the
small
pattern
amount
continually
a
of
meet
28
CHR.
”
“the
large
ribbon, thus
a
tion pattern
looking
ecological
This
of
amount
like
a
grassland
however is
be
in
explained
surroundings,
species
not,
relation
what is
by
the
This
stream.
the
the
of
presence
localities
ecological
is
also
and
contrast
Dutch
place
has
river/dry
wet
the line
to
trans-
central
a
of
com-
the Netherlands formed
demonstrated in
clearly
plant bound
a
first
more
patch.
the
within
in relation
Germany
line-shaped distribution pattern of
sea
roundish
a
mainly connected with the
usually done,
principle
distribu-
a
contain
always
the river Rhine in the
to
munication between southern
selectivity. So
example,
several
on
less influenced
more or
or
ribbon will
pattern resembling
a
direction of the ribbon. For
region
by
inequality
narrow
than
significance
european
to
high
greater significance
versal
LEEUWEN
In
herewith
shall
find
contrast
we
a
equality'
quantity of “no”, in the transversal direction of the
or
same
relatively
G. VAN
“
tidal drift
to
the
along
coasts.
In the
“fine
same
value than
relations.
a
small
a
way
grained”
will
large, continuous
The scientific
of the main
scattered pattern, which
or
comparatively
objects
for
be
of
[“coarse grained”),
one
at
conservation,
indication
lation is
playing
a
on.
It
one
is foundedhere upon.
will become evident that in this
dominant
spatial
rarities,
qualities of patterns in respect of various border types
The
dealed with later
calling
are
least in
of endemics and local
importance
nature
we
ecological
more
also Van
part again (compare
will be
case
granu-
Leeuwen,
1965).
1.3.
Elements
With
regard
of
elements
is
to
the
showing
be established.
manence
process
and
appears
is
a
In the first
The
be
part
theory
c
contrary
patterns
now
should
we
the
the element
importance,
more
meant
inequality-m-
system
studied
of
which
is
that
suppose
or
instability
equality
means
showing
a
or
per-
that
the
more
or
of our
of
theory special attention will be given
spatial
basic
and
to
the
temporal patterns.
relations
Starting point
of Whitehead
nd
of
is
of behaviour”.
We have started
a
the
discern
may
equality
instability.
or
spatial
Nevertheless
to
On
also
we
By
inequality.
temporal pattern only when change
basic relation between
2.1.
of
stability.
of
case
determined and
less fixed “line
2.
or
(process)
patterns
those
indicating alteration, change
process
can
temporal
permanence
Analogous
a
to
equality
non-alteration,
time is
of temporal patterns
aspects
onnection,
of
with the idea taken from the
that every
(1926),
separation. Further
philosophical
relation shows aspects
we
but closed when there is
call
a
relation
separation.
open
Next
work
of connection
when there
we
place
in
is
one
PATTERN
line with
with
“
2.2.
“open” all
With the
up
the
basic
Connection
1.
can
relation)
closed
will
we
be written in the
the
following
two
“
basic
to set
relation
of
relation chains’''
:
a»unequal.
place
aim
to
express
that
difference between the three concepts
hardly any
able
we now are
call
equal
Both chains in the first
than between all other
more
equal (in space and time) and
unequal (in space and time).
position given under 2.1
which
29
VEGETATION
as
(1st basic
relation,
«aopen
Separations
spoken of
concerned with
of equality
of the
help
first
equality and which
2.
that is
closed” all that is
Basic relation
AND PROCESS IN
terms
which
are
there is
significally
linked
together, no
closely related to each of
them.
Spatial relations
2.3.
Moreover chain
and
equality-in-space
with
less
a common
obvious
that
we
life
must
living in
of
to
us
that
This
small,
of
one
two
environments
much easier
possibility
of
reach
will
the
reverse,
“
rather scattered
conduction
localities,
nearly showing the
the other
by equality
as
is
end
spatial inequality
in
must
spatial
and
inequality
bring separation
place
”
even
than
holds
demonstrated
by
Europe.
On the other hand chain number 2 indicates that
will
on
connection. The first conclusion agrees
the famous flora of zinciferous soils in Central
separation
connection
spatial
and that,
equality
fact, but the second result is a considerable
ecological relations it means among other things
In
expect.
case
tells
directly
spatial
to
lead
circumstances
should
in the
lead
well known
one.
species
a
same
1
number
must
openness
and the
first basic relation
spatial closure and
that,
with it.
on
the
reverse,
Both conclusions
fit
well the modern ideas about the relation between isolation and
very
genetic difference
The
following
as
given by
concepts
Dobzhansky
(1951).
representing
are
some
of
aspects
spatial
connection:
hole,
gap,
touching,
breach,
known,
public,
yes”,
affirming, permission,
neity, continuity,
monotony,
evenness,
Concepts expressing spatial
barrier, unknown, secret, locking,
“
veto,
no
”,
unevenness,
A
negative,
restraint,
“
on”,
joining,
“
positive,
contact,
communication,
unlimited, similarity,
homoge-
general {common), everywhere.
separation
are,
for
example:
wall, dam,
“off ”, distance, isolation, protection,
difference,
heterogeneity,
discontinuity,
negation,
variety,
differentiation, particular {uncommon), unique.
large number ofrelation chains of direct connection can be compounded
from
aspects
type number
of
1
spatial connection
we
have chosen:
and
separation.
Thus for
a
openpaplain•mherdzaaggregation.
chain of
We
not
only know the expressions “open plain” and “herd aggregation”, but
also
these
two
phenomena
belong
to each
other. For
an
example of
30
CHR.
number 2
type
The
give: island
can
significance of isolation
often
is
we
over-,
organisms
from
as
island in the
an
factor in
a
sea
operating
water
apart
it is
as
a
biogeography and ecology
In
under-rated.
respectively
salt
surrounding
G. VAN LEEUWEN
the
The
least in the
at
or
chain
of
certain
ascribe this
gives
direct
a
organisms
some
adjacent environment,
above
compounded
to
the
to
geographical barrier. But why
it that such islands often accommodate precisely
elsewhere,
absence
quite usual
found
are
is
which
wanting?
this
of
explanation
problem.
The aim of nature-protection,
ed
the
by
chain:
an
aspect of spatial
isolationsprotections hands
separation, is
offs
veto
s
express-
differentiations
particular.
It
is
(=
principally
regulation
ancestors’ taboo
tance
question of saying
a
like
mechanism)
(noli
of islands
as
a
and
(from Thermodynamics)
and the first
1.
One
which
“noise”
growing equality
Information
“
information
will be
For
tition
”
show
we
“
information
in
lead
nature
the
to
species,
sur-
“
~
Order and
”
disorder
follows:
as
known.
is identical
information
to
”
can
preservation
The
and “iunknown” is
that
demonstrates
of
of infor-
be communicated,
and receiver.
amount
an
the
”
giver
which
Theory
signifies
“
between
looking relation between
the
must
Information and noise” (from Information
noise
mation. On the other hand
means
for
islands)
border
isolate
can
great impor-
*
“
basic relation
Separation
remote
our
modern culture.
pairs of concepts
Connection ssequal
2.
the
rich
reserves
landscape impoverished by
The connection between the
Theory)
of
mechanism
of
analogue
protection
nature
development of island situations, viz.
by
isolation
Wadden and Delta
our
conservation. And in the reverse,
rounded
an
tangere). From this follows
me
(such
no,
modern
a
a
which
paradoxical
explained by
quantity
missing knowledge
”
of
which
the higher the more complex or isolated the system studied is.
ecological relations it is very important to consider that compe-
belongs
under 2.
to
the aspects of chain number 1 whereas
in
Expressed
terms
of Mather
(1961)
we
co-operation
therefore
can
fall 8
write;
like
unlike
Competition,
must
based
always lead to
while
The
of tasks and
setting
up
*=»
of spatial isolation necessary for observation and research
upon
Herein
we
finding
an excess
fore the
run
spatial equality, ultimately
variety and increasing monoto ny,
based on various
difference
types of spatial
interests), will result in growing variety-in-space.
(both aspects of chain number 1)
of disorder
co-operation
various types of
the loss of
spatial
co-operation,
(division
on
the
great
eco-
risks
or
of
must
automatically involve addition
bio-system
getting
a
which
distorted
want to study.
we
picture of reality by
of
equality aspects, competition for example. Therecompetition concept still plays a dominant part in the thought
PATTERN
of
the
their
leading
Combinations of 1 and 2
ficial
such
regulators,
“
The
apparatuses”,
yes-no
mentioned before in the
finds
connection
competition
same
tends
way
be
to
the
research in
ecological
decreasing spatial variety.
to
found in all types of natural and arti-
are
thresholds, canals, sieves, membranes, diaphragmas,
as
punch cards and valves.
these
In the
31
VEGETATION
result in serious disturbance of relation networks in
can
concerned,
area
PROCESS IN
realize that
not
activities.
own
reserves
nature
who do
ecologists
most
result of
AND
phenomenon “life” is closely
thus
case
on
of
the
nature
its
associated
application of “no”,
This fundamental
protection.
such
in
expression
cybernetical
concepts
vetoing, blocking and negative feed back.
restraint, prohibition,
to
already
as
as
Temporal relations and the first basic relation
2.4.
For relations in time
also
we
can
3.
Connectionsopen sequal
4.
Separation sclosedsunequal
Among
the
as
concepts
aspects
past
(change).
of connection in time in the first
place
we
short all that has
consider
do with
to
same.
In the second
side the
one
(remaining the same).
in
permanence, constancy, stability,
remaining the
the
write:
place
we
past and
goes among
other
through time with
the other the future.
on
Connection with the
things through:
tombstone,
remembrance,
have connection
can
on
experience,
inscription,
relic,
fossil,
historical
continuity.
Similarly, connection with
the future for
example
can
go
through:
instruction, programme, design, blueprint, agenda, testament, warrant, prediction.
The
tying-up of the past with the future
maintaining
vancy,
In
are
the status-quo,
archives,
addition
storage,
memory,
to
continuation,
these
provision,
conser-
saving, faith.
logistical aspects belonging
chain number 3
to
likewise:
low
long duration, durability,
speed, always
and less
Concepts expressing separation-in-time
instability,
ness,
to
through:
runs
conservation, preservation,
disturbance,
change, transformation,
new, news,
forget,
are,
ending,
production,
example:
interruption, strike, precarioustransmutation, mutation, loss,
start, find, invention, revolution,
destruction,
uncertainty.
for
adventure,
improvisation,
surprise,
risk, unexpected.
In addition
are
to
these heuristical
short
duration, shock, high speed,
Here
we
can
can
write:
times,
to
chain number 4
as
well
as
once
does
our
more
uncertainty.
between those of 4. So for
this
chain form
have started the winter with
continued existence of
and
of direct connection between the
conservations fossils storages
uncertainty. Not only
we
some
again link up chains
various aspects of 3
we
aspects belonging
likewise:
a
a
reassuring
satisfactory
technical
an
example of 4
provisions continuations less
fuel
thought when
supply, but also the
society is dependent
on
this.
32
CHR.
As
annual
manuring
productivity
of
VAN LEEUWEN
craft
agricultural
plants reproduction.
and environmental
productivity
the
of
example
an
G.
and
stability
of
given:
connection
ploughing&
between
and between
instability
an
be
can
The
a
low
which represents
ecosystem
high
degree
of
one
important scientific results within the scope of the study on
“biomass-and-energy flow” (Odum, 1963) fits very well the chains
most
given above.
Combinations of 3 and 4
joins
with
motion,
the
remaining
inheritance,
renovation,
process.
dynamic
Together
(compare
also
are
rhime,
repetition,
balance,
the
the
progressions where change
this
to
are
vibration,
succession,
form
they
found in all
Belonging
same.
among
line
evolution,
characteristic
combinations
1
of
other
adaptation,
and
of behaviour,
cyclic
of
elements
2
things:
restoration,
‘"life”
mentioned
under
§ 2.3).
At
‘ ‘
the
outset
Open-and-closed
have
we
’ 5
introduced
but
Theory,
.,
in
using the
“
terms
”
and
open
“
”
closed
abolished it in consultation with Van
this
issue). Therefore
do
we
theorems under the
our
not
for
temporal relations,
der
here
use
Maarel
we
and
the abbreviations o(s],
a
stands for
Basic relation
The
-
c(t), but:
in which
2.5.
have
(see elsewhere in
-V
o{t]
name
because many students have difficulties
basic
variety,
a
d(s)
(1)
a(s)
(2)
d(t)
(3)
a{t)
(4)
for
no
variety,
s
for
space and
t
for time.
of inequality (2nd basic relation)
relation
of inequality
we
see
in
the
difference
or
division
„
between connection and
(a)
in
definite
a
separation
In order
get from
As
to
of 90°
a
mentary
J/-1 (=
definition
develops
d
turn
at
the
of
a
there
of
through
connection
thread,
at
then
that
a
point.
an
an
angle
90°). One can
provides the key relation to all
say
also that the
other relations of
Expressed in symbols:
stands
closedseparation
but
means
transversely
a
give; The basic relation of inequality is the compleequal and unequal, that which is found in the expression
a
Here
introduce
and vice
we
second basic relation
inequality.
( a)
we
a to a,
versa, one has to turn through
(compare the working of doors, bridges and bars).
relation between
to
When
direction, for example by
relation of
angle
separation.
same
for:
and
time
equal
#
for:
#
a.
openconnection,
“in
complementary
the
to
a
direction 90°
and
for:
unequal
different
from,
indissolubly allied with”.
AND
PATTERN
Basic relation between connection and
2.6.
and
The
in
separation
most
time
(3rd
a(t)
on
basic
important question
relation may exist between
we
and a, is
(Ad),
put in this chapter is: “Which
can
a{s)
on
hand and
one
dip) and
the other?”
Given that the second basic
a
separation in space and connection
relation)
and
(j)
a
33
PROCESS IN VEGETATION
the
just
same
should be
we
as
able
Now the
connections
of
we
to
and from
a(t)
(also
a{s)
to
should get:
d{s)
a{t)
(5)
a[s)
d{t)
(6)
and
(5)
This
reality.
d(s)
lead from
to
d(t). Thus expressing this in symbols
agreement with
thus the connection between
relation,
the relation between space and time
(6)
in
evidently
are
be demonstrated
can
by
good
very
different
many
examples:
In the field of
a)
time
long
protection
nature
the
over
problem
preference: “nature protection”
priority felt
by
second
one.
has
negative
has
a
a
many
The argument
sound,
on
nature
we
protection
The second law
change spatial
of
for
example
poor
rich
term
hand,
is
in
in
“conservation”
under the
positive
d{t).
write:
conservation.
states
that with
to
living conditions
communities.
species
precariousness.
for
protection”
every
(5):
change. J
precarious
on
the
“
standing
Salicornia
species,
a
for
spite of the
favour
is that
under the
thermodynamics
Stable environments
d)
munities
name
used
be
more
equality has increased. According
Environments with
species,
can
still
other
comes
nature
equality
c)
the
the first
conservation”
(6), therefore
to
name
against the first
while,
ranged for
should
term
“
a{s), but
According
b)
first
positive connotation. Now
negative
discussion has
“nature conservation” ? In
or
the
for
a
of what
are
According
poor
to
the other hand accommodate
example
coral
reefs.
in
(5);
com-
According
to
(6):
rich in
The territorial
e)
nous
to
in time.
By
species
singing of male grashoppers
means
of this other males
are
is
very
monoto-
repelled. According
(6):
repulsion
f)
stability.
Their
attracted
courtship
to
them
song
by
is
this
attraction
permanence.
strongly
varied.
variety-in-time.
variation.
Females
According
become
to
(5):
34
CHR.
G.
VAN LEEUWEN
The most
important conclusion that we can draw from the foregoing is that equality-in-space and equality-in-time cannot be treated alike,
than
more
any
Ross Ashby
Only variety
follows
that,
destroy variety.
can
the reverse,
on
variety-in-time.
this in his
assumes
variety-in-space
that
and
variety-in-space
(1956)
variety-in-time
can
be
requisite variety”:
formulas
our
counteracted
be
can
“Law of
From
—
only
(5)
and
by
change
(6)
it
and
destroyed on'y by spatial differ-
entiation.
The
mutually allied phenomena
aimed
at
nature
conservation has
formula
(6),
thus
other
no
In accordance with what is
also
Paradox
Paradoxical
is
relations
A
to
and
§1.2
spatial
between
easily
come
obviously
The work of
formula
§1.3
and
of this
from
out
the indissoluble bond
paradoxes
numerous
an
are
stability.
temporal
(6)
is
patterns.
of the second basic relation
confusion is due
are
and
purpose.
said in
expressing the connection
2.7.
life and regulation
variety-in-space
at
(
=
a)
scientific
important
very
key relation. This
type. To mention just
aspect of a although “to close” belongs
relations between noise and
the
between
and
a.
There
“Close
by”
to a.
of this
paradox
information
a
one:
is
type
found
in
the
and between communication and
which
the idea that noise and information
may bring us to
(or communication and information) are equal in fact. Inkspots blurring
information,
out
the letters of
written
a
message
deed carry the information that
they have disturbed
the
original
and
a
geneticist
presumed
to
information)
surprise,
be
of
an
electric
The
of
will
award
to
structure
of
aspect
to
show
Connection in
value
a
the
to
blueprint, according
certain
to
formula
direction is
to
or
These differences appear
When,
on
the other
then it
action of
is
mimicry
observe
(
differentiations
to us as
hand,
difficult
and
perma-
or
a
=
(
=
a)
same
connection
a) in
is
across
impossible
tied
up
to
in the
for
always
spatial patterns and
even
camouflage
(6):
possible where
being able
found,
of
blueprint.
The
separations
aspect
interested in isolation and the
more
separation
of
heuristical
within the ovum, will put in the forefront the
is present, and the
of
the difference
surprise.
mentary direction
presence
on
dealing with “information”
(5):
isolation ;<±
possibility
in-
by change.
engineer
great
a
formula
geneticist however,
logistical
patterns
doing the same. The first one, from whom may be
especially interested in communication (the carrying of
according
spatial
spatial
has messed with his
pen but
importance
communication
nence
permanent
context
Within this scope it is also of
between the ideas
as
someone
our
rests
reverse.
with the
surroundings.
structures.
our
line of
observe.
with the
comple-
the
key
The
sight is
hiding
relation. The
PATTERN
=
d) with
It is
the
surroundings
important
most
and vertical situations
to
in
or
horizontal
a
By
means
direction
the
account
(=
case
for
a),
=
and
in the
other. So
in
example
a
high
a
soil
a
amount
of
a) of the vegetation in question.
dispersion in
space
of the three basic relations it is
essential effects
’
when horizontal
with each
brought into relation
differentiation
one.
“basic paradox'
the
especially
connection
through
complementary
strongly layered forest, is coupled with
uniformity (
Concentration
very
35
in vegetation
our
take into
are
degree of vertical
profile
3.
in
in the
of ecological relations. This is
study
high
process
(= a)
object becomes invisible
(
and
belonging
to
and
time
possible
concentration and
the
approach
to
dispersion in space and
.
*
time.
For
concentration
densation,
(compression,
and
etc.)
convergence,
dispersion
(expansion,
accumulation,
con-
dilution,
divergence,
evaporation, etc.) may be considered as the two fundamental antagonistic processes which are completely dominating the events in the
world around
When
will
be
us.
concentration is
spatial
introduced
will
equality
the
to
increase
to
the
third basic relation in this
bility
combined with
more
case
of
hand,
dispersion and
the
external
true
to
stability
of Pask
and
According
or
to
inthe
internal insta-
more
spatial variety will be amplified in the
stability,
while,
the outside will increase
(1961)
when the external relations
Spatial
that
are
Fig.
also
establish
quantity
convergence
1, spatial dispersion
that
spatial
in
taken into
the situation is
concentration is attended
shown in
can
equality
stability.
divergence with instability
the internal aspects
lower
or
separation
on
the other
together
with the
instability.
The assertion
with
but
the result will be
external
to
connection
part
(externally).
thus the internal
spatial equality
a
(internally),
outside
On the contrary the internal
case
playing
centre
quite
by
a
by
Fig.
1.
With
regard
reverse.
a
concentration is
of border lines than has
going together
opinion is only
account.
the
coarse
however
is
our
grained pattern,
fine
as
granulation. We
characterized
spatial dispersion.
by
a
36
G.
CHR.
When
single
a
territory,
100
taken into
available.
nal
m
2
When,
on
possible,
contrary,
point
on
formation is
the
thousand
a
means
high
a
case
hundred
of
of
are
exter-
more taxono-
no
found
are
on
the
the
considering
dispersion less information is
a
same
external
with
met
to
high degree of internal selection. Nearly
in
territory
(1961)
question
has shown
important when
very
species
probable,
more
species
content
species being present.
one
the inside of the
tions.
we
is
studying
are
from
differing
this distinction in
pattern
ecological
rela-
•
Further
ought
we
taneity) has the
of
results
time. This
the
by
temporal concentration (simulby spatial concentration, just
be
can
amplified
brought
from all
by spatial
internal
type of
demonstrated
dispersion
flocking
be solved
to
are
that
caused
as
time. Difficulties
traffic
(for example:
consider
to
effects
same
spatial
distribution in
fore
a
combined with
each other. As Margalef
the
only
situation is
relations. Thus in
the outside
surroundings
information. To the inside however
the
the
then
every
that in the
concentration in fact
mical selection is
area,
species is dominating over a relative large
example, then this situation is rather improbable,
account
or
LEEUWEN
plant
for
Spatial
selectivity
VAN
by spatial
on
is
regulation
as
well
as
or
concentration
there-
cross-roads)
on
quarters
dispersion
by spreading
by spreading in
as
well known
in
nature
seasonal differences in the floristical
as
is
composition
of many woodland communities.
4.
Some
By
properties
dispersion
already
as
has
Leeuwen,
1965).
The
which
called
“limes
are
and “limes
areas
of the external and internal basic relations connected with
means
concentration and
discerned,
of border
main
two
most
to
by
(allied
dispersion)
these
spatial
to
may be
be
can
elsewhere
us
important qualities of
convergens”
diverges” (allied
of boundaries
types
been demonstrated
two
(Van
types,
concentration)
recapitulated
here in
short.
The
first-named
and
granulation
in
its
characterized
type,
patterns
by coarse
sharp, often straight border lines (“ all-or-nothing”)
is
,
marked
by unstable conditions
of internal
limes
variety-in-space.
convergens show
a
the
on
inside, coupled
Life circumstances
high degree
of
to
a
in border
low
amount
areas
of the
”
(“ now-and-then
instability
or
“
to-and-fro ”)
them
while
represented by
suggested
to
considering
A limes
is marked
the
concept
tension
as
an
of
in
poor
(production
“
aspect
of
”
ecotone
of
species,
to
mass).
this
most
of
We have
border
type,
instability.
divergens,
on the
contrary, which shows a fine granulation
together with faint lines of demarcation (“ more-or-less ”)
by stable conditions
variety-in-space.
between
vegetation
there is
many individuals
confine
stress or
in its patterns
of
the
mutually
The
on
the
vegetation
inside, coupled
of such
differing environments
gradual
is rich
in
to
a
high
amount
transition
species,
zones
many
of
PATTERN
them
it
as
is
represented by only
has been worked
quite appropriate
Both
Van
by
the limes
to
accommodate
types
Characteristic
VEGETATION
few individuals. The
a
out
IN
AND PROCESS
concept
“ecocline”,
Maarel and Westhoff
der
(1964),
divergens.
their
plant species for
37
of
organisms
own
the limes
and
convergens
communities.
for example:
are
Rumex
crispus, Ranunculus repens, R. flammula, R. sardous, Potentilla anserina,
P.
Trifolium repens, T. fragiferum, Hydrocotyle
replans.
vulgaris, Plantago
maior,
Inula britannica, Juncus
effusus, J. inflexus, J. maritimus, Carex hirta,
C.
C.
otrubae,
vulpina,
Agropyron
be
to
seem
common:
stances
and
and
things result
This
different but
very
in soil
“loss
often
which show
(manuring)
which
structure''
and
wet
heavy
disturbance,
“
'
rather
riding
for
as
looking like
of tanks
example
The
vegetation),
herds,
the
in all
marsh
be
agricultural
(treading,
when
to
it
for pastures
Potentilla
anserina
proved
be
to
influence of
Natural
the
more
and
or
banks,
on
areas
dominates
“
place
Secondly
regard
to
especially
Next
to
eminently
a
and
these
the
is
so
that
great
which
plants
of
Coast.
maior
permanent
“
So-called
often
of the
a
plots
on
to
several
like
tread-plants ”,
found
succession,
salt-
former
considered
connection has
This
overgrazed.
series
of
are
of
succession
a
along
roadsides,
there without any
are
plant communities of
to
with
met
along
inundated,
oligotrophic
sea
the
limes
and river
coasts
lines
boundary
on
environments
where
“rick”
”
poor
structure
the
in
vegetation
and
damaging of the
large animals living in
by
desalination
now-and-then
and
on
“environmental noise”, thus
bad
a
treading being present.
or
eutrophic
heaths
induce
to
found in the unstable situations
surroundings favourable
flat
reclaimed
on
done
less
Sea
Plantago
convergence in the first
between
from
measures.
those
important elements
grazing
the top
manuring
accelerated
North
in
types of heaviness
possible
similarity
clearly been demonstrated in
along
is
what is
This
artificially
localities
other
such as
Trifolium
Triglochin palustris and Festuca arundinacea,
accompanied by just
typical
and fresh
among
salt-marsh, with species
wrong
especially
an
it is
grazing
convergens.
on
is
of
salt
On soils sensitive for this
tractors.
found
are
brackish
a
closely allied
are
limes
based
of
means
effects
and
Netherlands,
fragiferum, Ranunculus sardous,
only by
in
’
ojf-the-road-locomotion”
of the
parts
dry,
changes
(increasing convergence)
.
to
the
to
important feature
one
_
eastern
canina,
the synsystematical
compactioning.
of spatial
also due
treading
A.
stolonifera,
to
irregular temporal variety in life circum-
by alterations between
rich
_
soil is
and
caused
as
bound
et
Prsg. 1950. The plant communiliving in environments which, mutually compared
are
strong
poor
all
Plantaginetea maioris Tx.
ot the
ties of this class
1
class
Agrostis
repens,
Festuca arundinacea and Folium
perenne,
a
which
water
natural
for
can
the
places
where
of
is found.
a{t),
grow
induce
management
in extensive flat
these
other
plant communities
of
suited
all
high degree
quick
from
on
disturbance
various
or
strong
bone-dry
to
soil
types,
or
the
changes
with
soaking
wet,
areas.
possibilities
limes
man-made
convergens
and
landscapes
its
are
vegetation
pre-
types,
38
CHR.
above
all in their
and
more
straight
present form.
LEEUWEN
For
the modern technical civilization
favours the limes convergens
more
demarcation,
G. VAN
vegetations,
monotonous
canals,
soil
by
and
compactioning
by making sharp lines of
barrages,
scale
by
metalled
roads,
enlarging
in
re-
allotments.
The
limes
anthropogeneous landscape represents
is
ever
showing
convergens
external
regulation
always
by
seeming
a
The
man.
Environments of the limes
of
which
among
plants
Silene nutans, Dianthus
Agrimonia
eupatoria,
is
The internal
ensure
the
presence
in
expected
the
this
types
or
natural
the
where
situation
dunes
distances
transition
lagg-zone
where
areas
an
covered
The
by
where
quite opposed
of
Instructive
bogs
on
to
oligotrophic
water,
a
stable
down from
the
”
it is
also
of the limes
on
divergens.
gradual differences in
to
this
such
as
subsp.
This
Linum
to
respect
mutabilis and
last
of the
shown
are
in
situated in
higher level,
a
G.
is
space,
Gentiana
germanica,
Carex
found
on
varying
an
”
places
from
a
could
unstable
by
a
to a
palustris,
Spiranthes spiralis,
be
instability
low
are
degree.
belonging
large quantityjaf species
baltica,
campestris subsp.
Parnassia
already
form of the
where
high
G.
Fritillaria
flacca, Briza media, Juncus capitatus,
orchids like
is found in
divergens
convergens
mark off
which is characterized
Scirpus planifolius,
J.
sub-type
catharticum,
uliginosa,
the
“rich'”,
be clear that several environments just mentioned
sub-type
and
in this article.
of the limes
possible
shows
may
wet
eutrophic and
between woodland and non-woodland
treated later
sub-type
limes
It
border
dry and
of limestone hills
slopes
“
distinguished,
on
sandy soils.
specially
“
as
dominates
on
gradual
of
off into
mentioned in
coming
biological culminating-point
which will be
shading
slopes
swimming marshland
eutrophic environment and
poor
on
examples of this situation
in
be
fundament
a
found
are
valleys,
“poor”
can
may
where
as
boundaries between
what is
limestone,
gradual transition-in-space
Just
on
hand
is found,
convergens
They
zones.
one or
single spatial
divergens
places
to
a
these
to
than
of years.
tens
available
are
pulicaris,
of orchids.
belonging
limes
inundated
never
environments
being
limes convergens.
reaches
of
another
C.
species
the other
on
the limes
confined
always
great
gradual
sand
oligotrophic
the
of
all
almost
to
tiny populations for
soils often inundated and
clayey
case
Trifolium medium,
dioica,
to no more
comes
between salt-marshes and fresh dune
rather poor
the
area
of the situation
localities
nearly
of
and
required being limited
of such
height
nissolia,
Carex
of individuals
certain
a
stability
same
case
differences in
for other
in
environments
Typical
L.
odoratum,
Eriophorum latifolium
present
point.
areas
to
for
call
total number
two, the circumstances
in
the
divergens accommodate quite
we
Polygonatum
the
cases
many
species which
but
related
demonstrated in
Lathyrus montanus,
C. hosteana, C. buxbaumii,
In
is
example: Botrychium lunaria,
superbus, D. armeria, Hypericum pulchrum, Pinguicula
Genista tinctoria,
vulgaris,
which
of now-and-then how-
and dikes.
barrages
set
is
as
special sub-type of the
”
uncertainty
here
threatening
a
stability
Orchis
amarella
meleagris,
J. tenageia,
morio, O. maculata,
Epipactis
even
palustris,
”
“
divergent
”
side of which
the limes
that
idea
the
limes
types
In
showing
water
a structure
with
thus
in
life
is
has
relations,
scale
is
and
on
of the
Isolation
influence
lead
dry sand
This
internal
and
agricultural
of
amplification
to
spatial
the
regulation
not
change for centuries. By
steadily increased.
distance
by
the
on
chapter.)
prefer loamy
layers.
top
the modern situation the former
circumstances.
variety-in-space
2.
to
auspicious micro-instability
The methods did
1.
the “Law of
which is intermediatebetween
an
to
one
the other side
the last
appear
the
on
expression in
dealed with in
providing
the
developed
conceivable,
standing
finds its
(We
conopsea.
Theory, started from
amplifying
result of three factors:
the
mainly
Gymnadenia
divergens also
automatically
systems
diversity
and
instability”
management
contrast
mining
concept
of the unstable limes
water-logged clay,
in the
convergens
39
VEGETATION
Relation
our
gradual
a
This
divergens.
Species
was
bifolia
environmental
requisite
soil
Platanthera
IN
framework of the three basic
convergent
and
PROCESS
thesis here that
put the
can
“
the
PATTERN AND
dwelling-places in the
the outside with all
a
gave
This
landscape.
of the action-radius and
centre
human
to
minimal
was
the
stability
restriction
gradual
restraint
this
the
near
maximal
possible degrees of influence between the
at
extre-
mes.
3.
Our ancestors were operating gradually and on a small scale.
By this dispersion-in-time, which, as we have already shown before,
has
the
Besides these
control of
our
the
by
means
It
still
three main factors
may
in
such
and
and roads
be obvious that the
vegetation
one
types
as
as
the
variety
rules
or
only
for
the
of diver-
of the
unevenness
soil
and the borders
watercourses
not
basic
smaller elements
of
metalled
primitively
of the
preservation and control
belonging
most
extensive
flat
areas
provided
to
the old
important and
tasks for the Dutch Nature
equalisation
at
country is missing
the
the shelter
by the differences of level
which
areas
time
same
The
Conservancy.
of
man-made land-
more
so
against
common
to
most
as
our
spatial
regions
accidented.
more
The
that
(now used
numerous
landscape,
of ways
footpaths
scape constitutes
over
spatial
of loam.
contain
difficult
reserves)
the
windings
old-fasbioned
the
dispersion-in-space,
as
more.
nature
occurred
gence
surface,
effects
same
increased still
relation
diking
in
theoretical
approach
to
makes
ecology
and reclamation of the Wadden Sea,
it also
clear
combined with
straightening of the coastal line along the Wadden-islands should rob
the
Low
based
on
from
Countries
nearly
all
types
their
of
last
contrasts
area
where
reaches
an
spatial
divergence
almost mountainous
height.
The continued research
pay
special attention
to
the
into the
properties of border
ecological relations bound to
areas
has
to
the steepness
40
CHR.
of environmental
gradients
between convergent and
observed
various
on
The
In
theory
the
been
pointed
by
as
zones
have
we
shaped
screen
like networks.
the
certain
applied to
ecology
connection
possible
ecological
other
important
between
phenomena
of the
facets
has
the
already
broaden this
to
relations between
and their environment.
postulate,
plexity
of
want
of
and
to
problems
the
to
and
pattern
we
border
and
the
basic
of
way
relations
the
thinking,
we
com-
(or structure and organisation) in the
Form, size and border type are considered
process
of
respect
stability for temporal
course
from
cybernetical
study.
in
elements
leading
degree
ecological
according
ecosystem
on
meeting
which,
often characterized
are
chapters
some
Approaching
size
spatial
many times. It may however be useful
out
to
the
to
as
relations
relations and
perspective
plants
basic
preceding
three basic
LEEUWEN
well
as
places,
of
VAN
divergent environments,
vegetation patterns looking
5.
G.
of
type
spatial
lake
a
will
we
of
are
have
character of the local
next
patterns,
So
ones.
type and
to
opinion that form,
determining
a
succession
influence
from open
water
to
marshland.
To
from
start
and
and
structure
that the famous
organization
Struggle for life”
regulation,
thus
for
threatening from all sides,
only by
of the
means
This
d{t).
thus
,
means
interpreted
0
a
among
can
w,
other
things
for survival
struggle
directed
fight,
against a[t),
as
against
change
be crowned with
success
_
various
of
aspects
a(s),
thus
by spatial
variety.
,
Therefore
“
a
strangler
for
light
a
”
and
plant which is able
water, but
temporal variety within
can
grow
only
could
and
disturbance
etc.
an
The
organ
where
caused
as
most
effective
“
homeostasis
”
we
do
which
observe
can
are
forming
the
by
mass
offshoots
is
determining when they
a.o.
abiotic
to
all
competition
a
wipe
plant
out
even
allowed
will be
Mentha
is offered
consider
not
and
concerning ecosystem. Such
internally resistance
genetic damage, uniform
called
form offshoots
experimentally demonstrate on
Externally
or
to
fellow-creatures in ruthless
the
places
on
,
we
as
local circumstances
besides,
as
its
tyrannizing
and,
formed,
aquatica.
of “noise”
types
environment,
by
virusses,
growth, quarrels within communities,
weapon
forged
for
this
many-sided
battle is
"
(Cannon,
il
1929)
or
the know-how
of equalisation-in-
time”,
During history
of life
quantity of regulation
with
aim
the
of
the various
a(s)
numerous
richness
stood
regulator itself with respect
as
a
belonging
closely
mental
available,
their
One of these
each
to
the
of which
status
aspects
can
a
large
quo ante”
be
is
the
under-
stability of the ecosystem
to.
The function of
be
species
maintaining
aspects.
enormous
it is
in
organisms have developed
mechanisms for
a
certain
associated with
a
plant species
special
as
a
regulator
we
suppose
to
trajectory and degree of environ~
instability.
In
other
words:
Every plant
species
would need
an
AND
PATTERN
and
trajectory
own
functioning
as
In
for
this
has been
in
rapidity of
a
Further,
six
of
and b
dry and
sake
regarded
plant
lying within
many
as
of
left
are
of all
of
possible
instability)
other environmental
the
simplicity,
out
same
single
or
and
frequency
of consideration.
a-f
species
useful
a
Fig. 2.
scale
(a type
wet
of
“Law
may be drawn into it, either
the
for
Active
are
course
this
regard
we
graduated
a
extreme
but of
top
the alteration
respect
Species
the
at
only
showing change
combination.
In
model
extreme
are
which
instability”
and
development
of
further research is represented in
our
drawn
factors which
41
simplification
_
hypothetical
changes between
A
regulator.
environmental
hypothesis
VEGETATION
degree of instability for its germination,
efficient
an
_
requisite
IN
PROCESS
the
is
figure
showing:
trajectory (a), but they
are
different in size.
and d
Species
c
Species
e
a
low
are
requires
equal in size but different in trajectory.
a
high degree
Next
a-e—f,
we
can
d-e,
c-e,
combinations
and e—f
possible, but
are
d-f.
Epipactis palustris — Potentilla
couple Epipactis palustris
that
assume
diminishing,
of
amount
are
low
a
higher
try
to
locality
anserina
explain
far
'
V.
a-c,
the
and Lolium
(c—e)
so
why
as
When
wc
we
know,
finally
2.
will
increase,
degree
one.
of
the
third basic
possible
instability being
The Relation
Theory
’
dissemination, accessibility (Heimans,
Dr.
may
relation,
numbers of
much
Westhoff
the
total
trajectories
larger
than of those
may also be
helpful when
explain the fundamental problems belonging
"
by
a-d, a-d-e, a-b-e-,
the combinations
found, but not,
inaccordance with the
species
indicating
like
f needs
species
species has its own trajectory and degree of requisite
Fig. 2, demonstrates that as environmental instability
marking
a
so
— Lolium perenne (c-d).
Fig.
we
while
every
then
instability,
not
This, for example,
perenne — Potentilla anserina (d-e)
is
instability
read that the combinations a-b,
c-f and
b-c, b-d, c-d,
the
of
one.
phenomena
1954), accession (term proposed
to
,,
meaning the integration of
and for the first time mentioned in Van
a
plant
Leeuwen,
with
a
1962a)
new
and
succession.
For briefness
sake
we
shall here confine ourselves
to
some
problems
42
CHR.
around
succession.
plant
succession is
showing
a
As
G.
VAN
to
and characterized
stability
It is
high
as
striking
allowed
as
has
a(t)
maritimus,
a
but
coast
into
plant
not
also
in
a
A
able
are
every
in
poor
tidal
and
degree of
a
settle
to
this
formed
waters
delta of
the
in the
claypits
as
newness,
is
along brackish
water
with
species
good example
found
sand-
around young waterfilled
in
how
matter
no
the fresh
process
complete series
a
situation.
the local
by
being.
only
In
convergent pioneer stage
pioneer plants
that many
come
regulate
high degree of environmental instability
by
community rich
a
environment has arisen,
new
of
terminal
divergent
a
and
orderly
an
marked line of behaviour.
vegetation develops gradually from
species
LEEUWEN
soon
an
as
by
Scirpus
the
near
Biesbosch
eastern
a
aspect
sea
and
of
parts
the Netherlands.
For such
usually
it is
appearing
be
can
a
an
bare
ecologists
that
fact
by
luck when
It
to
its
are
when
we
we
the
come across
that the
intervention
can
and
a
certain
inclined
to
has
species
that
to
may
be
of their first
of such
the
species
After
time.
right
established itself
once
this
more
the classic
species coming
point-of-view
relation-theoretical
favourable for the
change
being fulfilled
view in
into the
a
further
however
when the environment is still
area
”
as
a
ecology
compe-
species in question.
communis and Cladium
ground
or
change in
Carex
the
species
mariscus
lasiocarpa.
water
then
cannot
win
On
the
management of
other
as
even
for
hand
like
type
not
a
square
example
just
one
to
Phragmites
centi-
Plantago
abnormal
divergent swimming marshland
a
give Phragmites complete dominance within
may
a
This condition
expansion will be improbable. According
their weaker brethren
on
expansion
an
showing
experience so-called powerful and intolerant plants
coronopus
gradually
begin.
a
of
is
species
conquest of the empty space
quite in accordance with
of
rich in
even
“
consider this
only probable
metre
very
species
a
moment
settling
in
comes
often observe
individuals and
more
From
one
or
years.
The alteration which underlies
the environmental
( t)
shortness
extreme
means
like this is
two
for
surroundings, starting from the first individual. Many
of course,
tition will
our
aggregation of d(s) is
possibility
the
that
an
also
individual of
plain
extending
matter
the
establishing has become impossible.
When
on
good
anywhere.
prevented
that its
to
no more than one time.
Just in the case of perennials which
growing in closed formations (for example Juncus maritimus)
to
are
bit of
a
next
itself is marked
establish
confined
environments the herd
young
characteristic,
elements.
(1961)
has
instability,
This
pointed
increase
out,
a
succession consists
thus in
of
an
stability
by storage
of
in
augmentation
is
stimulated,
organic
decrease of
a
of the local d
Margalef
as
structure
or
“infor-
mation”.
This may
happen by formation of
increasing divergence
(refining
growing co-operation (division
at
the end of the
humus and peat in the
of the
spatial
of tasks and
series, often consisting in
a
topsoil, by
pattern grains)
interests).
and
The stable
cyclic change
as
by
state
clearliest
PATTERN
is shown in heaths and
AND
the greater
part
the
on
quantity of d (t)
helpful in destroying
large
it
landscape
forms
of the
to
common
“climax
a
the
In
community”
thus
the
reverse
be
may
within
variety-in-time
community
for
dependent
surroundings.
environmental
part of,
a
1947; Stoutjes-
1962) is always remaining
stability
the
43
VEGETATION
1923; Watt,
bogs (Oswald,
1959; Van Leeuwen,
dijk,
IN
PROCESS
functioning
the
as
a
regulator.
As
a
our
study
succession
permanent plots has shown,
on
and
course
direction of
of the basic
highly dominated by the rapidity
are
ation, the transition from a{t)Xod(f). This transition passing in
short time,
tens
of years
on
be the result. This
mental
we
the botanical
do
not
richness
circumstances will
the
the
of
surface,
future
the
during
their
controller
of such
horses
and
cattle
is
will
reserves
of the basic
slowing-down
to
a
at
the
one
which
a
buffer.
with
the
environmental
blow. Those
showing
are
will
areas
relatively
a
the localities
next to
and those which will remain salt for the
the
very
leading
regard to
along the Zealand
Delta-activities,
speak,
so to
as
high
too
marshlands
a nature reserve
as
working
expectations
our
put
change,
highest value
accidented
soon
the same
spot a much more divergent situation will
divergence will become the larger the more environ-
where,
aestuarium,
develops
vegetation. When however the transition lasts
divergence is already available
Therefore
get
heels succession”
over
monotonous
convergent
for
“head
a
altera
and
be
We
change.
best
already being less saltish now
longest time. The main task for
at
to
the
in behalf of the
regulate
have
found that
same
time
grazing by
practical
most
method for this purpose.
Many
means
cerning
other
aut-
and
of the Relation
synecological
Theory
with
example
phytogeographical
sense
regard
is
occurring of
Europe
Stebbins &
a
areas
Major,
At the end of this
ideas about the
ties
or
Beeftink
Prunetalia
chapter
spinosae
Th.
some
Tx.
Crataegus
sanguinea and
his
recent
western
attention may be
and
1962).
and
zone
In
monogyna,
Rosa
Outskirt
central
forming
canina,
(Prunetalia
paid
to
our
environment spatially
is the
nature
can
most
communities
Europe
the
spinosae)
here,
even
at
or
botamcal
spatial transition
characterized
Prunus
variegated and
produce
new
(Mantle communi-
non-woodland and
Rhamnus cathartica
one
of
coast
transition
a
or
spinosa,
like
Agrimonia eupatoria ( Trifolio-Geranietea ). This richness indicates
stable
a
thesis
the
more
Trifolium
and
that their
same
than
by
Cornus
medium, Polygonatum odoratum, Geranium sanguineum, Origanum vulgare
most
An
in
He could associate
along the
of forest borders
1952
Muller
between woodland
like
endemics
1965).
richness culminates in these communities
species
of
in
Europe.
plants
in each of which
plant communities
Trifolio-Geranietea,
zone
con-
is
gradual
pair of mutual different climatic types (compare also
with four
found between
approached by
mention those
problem
Western
endemic
numerous
the
to
given by
(1965) about the salt-marshes of
the
be
can
we
and endemics.
pests, relics, rarity of species
ecotypes, adventives,
interesting
questions
among which
time the
that of
a
44
CHR.
woodland.
pure
maximum
forest
of
is
VAN LEEUWEN
composed
realised,
botanical
a
gradient
already
of them. The
necessary
all
as
contrast
these
kinds
found where the
in
the other
are
the
on,
a
(first
stage
from
3.
Fig.
concerned with
later
of
species
these
of the
of
The
through
a
grassland.
angustifolium,
as
etc.
may
periods
zones
are
by the
to
be
not
also known
this
is
contrast
forest
not
edges
are
occurring.
exist in time and space
degradation
and
Rosa
a
spec.,
to
figure)
woodland
stage
we
roses
(at
degree
indicate
in view
a
change
the foot of the
glutinosa,
high
as
is
figure).
by quite another
Alnus
state
with,
met
needing
of
a
and,
(second
now scorn
are
we
through
canina should
low
in space
heath
idea, still often
characterized
Betula
or
transition)
A former
monogyna
spec.,
are
or
on an
combination
seems
where
periods of radical change
Salix
a
the top of the
that hawthorns and wild
transition
itself
grassland
spatial variety combined with
such
or
scarcely
are
(at
case
temporal
along gradual transition
two
plants
to
Crataegus
supposition
amount
found
like
etc.
formed
3.
the first
of
a
shaped
only develop
and
artificial
many
succession from
woodland
transition
two
In
state
temporal transition)
that
can
spot
landscape
hand,
woodland/non-woodland
contrast
pioneer
the
concerning communities
illustrated in
wedge
outskirt-communities
Fig.
The
gradient
on
woodland/non-woodland
mantle- and
on
outskirt
gradient “pur sang”.
present
of localities
while,
wanting,
as
vertically
wet/dry, salt/fresh, grazed/not grazed,
contrasts
and
mantle
by
each
that this botanical
must assume
ecological
G.
mosaic
a
divergence
edge giving
We
from
In
set
of
Epilobium
PATTERN
AND
PROCESS IN
Fig.
The
close
relation
between
important gradients
determined
cally
The
greater part
on
a
between
surface
the
water
contrasts
well
and
lime
on
are
occurring
frontier.
and
and
which
holocene,
slopes
as
specimina
along
as
for
acid boulder
the
and
spatial
east
the
of
most
geomorfologi-
or
as
shadings.
of the Prunetalia and
indicating border
and
saltish
Limburg table-land,
landscape,
slopes of
example
clay
lines
are
of the South
the whole dune
on
between fresh
gradients suitable for mantle
particular spots
stone
been drawn
these lines
to
the
as
edges of rivervalleys,
smaller scale such
munities
have
of the natural dutch
pleistocene
as
communities
of the Netherlands their
map
larger scale, mainly based
Trifolio-Geranietea is bound
areas
4.
Prunetalia
gradients is shown in Fig. 4. On this
45
VEGETATION
-
etc.
On
a
and outskirt
dikes
many
meeting
area
Winterswijk
and
-
much
com-
ditches
between basic
near
the German
46
CHR.
the lines drawn
By
country
nities have been found
has in
the
on
particular
by
characterized
or
still
time those
same
the
occurring. Beyond
limited
areas
of the
areas
and
plant species
rare
within the
case
concentration of
a
the
of its
most
are
been the
LEEUWEN
at
map
indicated where
are
G. VAN
commu-
this
seacoast
by
a
circle and
gradients.
ACKNOWLEDGEMENTS
With
regard
leagues in
all have
E.
van
him,
Maarel
der
RIVON,
and
Mr.
greatest recognition
fundament already
spatial
ten
differentiation
Dr. Ir.
M.
L.
J.
W.
most
Dr.
and
that
out
has
last
Drs.
brother.
laid
essential
the
conservation and
nature
A
expe-
The
Beeftink,
author’s
Westhoff, who
col-
They
practical
criticism.
Ir. W. G.
the
his
to
grateful
of nature reserves.
scientific data
Leeuwen,
pointing
hand
is
and regulating
Soest,
van
van
by
ago
going
are
interest
is directed to Dr. V.
years
author
of
stream
a
permanent
Prof.
the
to many controllers
next
only by
not
their
concerns
here
developed
ideas
of the
with
also
especially
His
the
to
out
sustained
but
riences,
facet
and
in hand.
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