Download concepts and perspectives

Acta
Bot.
Neerl. 42(4), December 1993,
p. 413-433
Review
Mechanisms
vegetation succession:
of
review
a
of
concepts and perspectives
J.
Andel J.P.
van
Bakker
Department of Plant Biology,
and
A.P.
University of Groningen,
Grootjans
PO
Box
14,
9750
AA
Haren,
The Netherlands
CONTENTS
Introduction
413
Concepts
414
414
Vegetation
Succession
415
Kinds of succession:
416
phenomenology
mechanisms and functions
Causes,
Clarification of the
Vegetation
417
framework
418
mechanisms of succession
419
Seed bank
419
Establishment
419
Facilitation
420
421
Competition
Biogenic
causes
of
vegetation
succession
422
422
Herbivory
Vertebrate herbivores
422
Insect herbivores
423
Parasitism
424
Food webs
424
425
Ecosystem mechanisms of succession
Dune
Moist
425
valleys
426
grasslands
Salt-marshes
426
Mire systems
426
427
References
Key-words:
causes,
mechanisms, succession,
vegetation.
INTRODUCTION
succession
Vegetation
Nevertheless,
vegetation
there
has
been
much
change
suggest knowledge
&
Slatyer
Masing 1987;
(Burrows
the
gradually
1990).
Burrows
in
the
many
field
of
and
papers
terminology.
abandoned in favour of the
Phenomenological
of mechanisms, and
1977;
of
subject
confusion
succession has been
vegetation
(Connell
is
these
are
terms
labelled with
1990), ‘scenarios’ (Peet
Glenn-Lewin & Van der Maarel
413
1992).
et
The
more
been
terms
such
or
al.
reviews.
concept of
general
have
1992),
Pickett
several
term
created
as
‘classes’
to
‘models’
(Zobel
&
(1987) distinguished
414
J.
between
pathways,
to
attempt
autogenic
and
confusing
if
In
to
we
only
can
and
has for
long
succession.
The
towards
mechanistic
Grace
a
&
mechanisms
Connell &
applied
are
dynamics
competition
ANDEL,
J.
P.
of
BARKER
vegetation
AND
Tilman
in
guess
transient
been
recognized
in
approach
has
1990),
vegetation change (Tilman
also
in
as
lit
1988),
and
made
of these
use
an
between
is
terms
from
the research
a
as
mechanism of
vegetation
approach
plant
on
such
terms,
these notions. Plant
to
phenomenological
important
an
decennium,
competition
influenced the
positively
GROOTJANS
distinguishing
mechanistic
recent
way
Berendse & Elberse
1988;
tition, however, might result
which
P.
organization (vegetation, ecosystem).
dynamics (Tilman
the last
during
move,
different levels of
at
A,
succession,
models. When
Slatyer’s
allogenic mechanisms, following Tansley (1935), the
they
addition,
soil-driven
and
causes
relate these
VAN
view
The
1990).
(reviewed
by
mechanisms
on
emphasis
on
of
compe-
underestimation of other mechanisms of vegetation
an
change.
Overall,
enlarge
first
to create
and
conceptual
the
on
of the
understanding
we
review
to
particular
in
enables
knowledge
attempt
challenged
focusing
our
purpose,
felt
we
succession,
concepts
order in the
of
with
concepts
of
regard
perspectives
to
vegetation
they provide
succession.
vegetation
useful
hypotheses
and
predictions.
vegetation
and
succession,
of terms, thus
use
providing
a
For
and
this
then
framework for further
repeat earlier reviews,
not
to
such
Only
concerning
work. We will
experimental
their conclusions for the
several
evaluation of the
mechanisms
formulation
review
some
an
but rather
use
present purpose.
CONCEPTS
Vegetation
successional
In
composition,
floristic
terms,
least
at
to
another)
and
vegetation change
According
which in the
then been
if
changes
characterize the
there
is
history
two
association
is
not
communities,
Gleason
after
(1939)
aggregate of plant species
plant species,
of
textbooks,
rather
has created
community
while
a
than
also
species
basis
‘species’
one
of its
within
another,
to
when
implies
we
phytocoenon
a
labelling
together opportunistically
an
acknowledged
vegetation
being
unit with
by its
composed
does
For
not
‘a
a
plant
(1926,
justified
opinion,
but
unit,
coincidental
that
that
merely
aspect
of
a
plant
association
definite successional
in
an
is
an
status
floristic composition’.
is considered
problems.
population
this
Whittaker
not
we
prevailing
vegetation
species,
and have
Gleason
recognized by
from the
a
also
(cf.
of the coin: ‘Are
even
ecological
change
to
also
were
side
of
particular assemblages
are
come
emphasized
recognizable
number of
the
one
scarcely
having
forming
physiognomy,
majority
approach
have
far removed
organism,
an
However,
of uniform
emphasized
conclusion,
coincidence?’.
In the
communities
cover
aspects, individuality and order,
general
(from
phytocoenosis
one
the notion of
climate conditions continued
as
the
on
‘species replacement’.
plant
global plant
1926 paper he
the
to
(from
what
on
term
(1991),
of
in
change
a
community
present paper, dealing with mechanisms of succession,
focus
with the
Tallis
to
In his
coming
we
re-ordered
1975). The
a
we
of succession
phases
Here
phytocoenon).
of
community
as
worry about the distinction between stages of succession
not
and
plant
long
In the
composition.
do
1939).
a
as
of
be
to
plant
example,
become
composed
individuals.
a
species
extinct,
of
The
may
populations
population
disappear
either
from
because
the
MECHANISMS
boundaries
OF
of
a
the
Moreover,
using
vegetation science,
problems
be
can
population
term
characteristics
while the
(Grubb
of
species
such
the
about
generalizations
al.
et
life-history
vegetation
component of
dual
only
not
an
ecosystem (Odum
hierarchy, represented
histories and
and
energy
maintain
by
and
evolution)
much
of
individualistic
This draws
that
(Gleasonian)
‘it is
al.
with succession
disturbance
of
(e.g.
vegetation
White
(e.g.
Pickett
et
al.
the
has
(1992)
but
also
approach’
the
both aspects:
acknowledges
plant populations
[in
his
1951
in coherence with their sites and
growing
forming part
their site factors and all other forms of life
still far from
getting
to
the bottom of
into
recognize
to
with
of
cycling
and
a
life
to
holistic
a
in several studies
WesthofTs
1992)
which
of the ecosystem
or
growing
we
prefer]
together
between,
of
‘emergent properties’
with
definition
(1979)
largely spontaneously
individuals,
with
reasonable
& Pickett
in these sites’. In
occurring
insight
which result from interactions between
of
plant
wrote
functional
a
even
community
1991).
‘A system
he
paper
al.
et
considering
to
(dealing
(dealing
that
successional
as
(1986) proposed
1987),
species
shown
to
attention
our
1985; Falinska 1991; Myster
1989; Van Andel
plants.
1979 and
concept of ecosystem function’. The latter aspect has been neglected
dealing
case.
of visible
subordinate
feasible and
of
be
can
the
not
composed
be
(Grime
approach’
quite
concept
quality
which is
in relation
plants,
‘process-functional
nutrients). They suggested
an
Brown
et
the
and Muller-Dombois &
to
plant species
O’Neill
of
determinants of ecosystem
related
1987).
of all
dormant seed bank.
a
association of individual
main
succession
individual
1971).
to
(1951)
‘population-community
a
a
an
the
more
of
strategies
as
population
thought
and
oversimplified.
composed
as
are
Grime
1982;
age of the habitat may have been
is
being
as
stability
is
that
Westhoff
plant community
species-richness
Grubb
1977;
community
boundaries
quantitative aspects,
to
community
productivity,
as
of
aspect
a
the
survives
suggest
may
populations. Only
Ellenberg (1974) recognized
Dominant
a
with
population
in addition
avoided if
overlapping
coincide
not
because the
or
involved in
individuals of
do
phytocoenosis
component populations,
Such
415
SUCCESSION
with
we
are
communities,
organisms (see below).
Succession
Until
now we
in
rather
a
According
that
change,
but the
if succession
into
a
succession.
considering
useful
to
‘changes
does
not
Tansley
of
an
escape
phases’ (Tansley
1935),
is
important part
the
sere.
which
reversible
Miles
(1979),
markedly
to
changes,
have
that
there is
in
no
whereas
as
having
into
a
successions
Similarly, Finegan (1984)
the
at
mechanisms of
of
use
the
all
historical
involves
not
which
can
be
abandon the
a
different
patch
type’,
directional
defined succession
as
of
merely
phases
or
separated
concept
stages.
it is
vegetation change,
vegetation
mech-
value of the
ascertainable laws. Even
to
of
term
non-committal
species composition (or abundance)
defined
are
to
vegetation
need
fluctuations.
the
of the
concept
notion of ‘a series’
alter the appearance of
changed
this
phases subject
of change
in
‘vegetation change’
include
useful when reviewing
seem
may
stated
sequence of
a
confusion
sufficiently acknowledge
(1935)
mechanisms of succession
considered
state.
much
differentiate between fluctuation in
successional
be
to
‘a continuous process
The
so
is
would
term
abandon ‘succession’ in favour
recognition
is
series of
latter
best
it
succession.
the
there
Though such
succession,
of
concept
succession’ and
‘vegetation
terms
Actually,
(1990),
seems
change’.
‘vegetation
of
it
the
using
way.
Burrows
to
succession
anisms
have been
arbitrary
dynamics
vegetation
added
changes
that
away
in
of
When
quite
and
general
a
as
such that it
can
fluctuations
are
from
‘the directional change
an
initial
with time
416
J.
of the
progressive,
mechanisms
hierarchical
al'
(1993) proposal
s
from
year
one
succession.
For
McIntosh
(1980),
We
start
Miles
from Pickett
succession:
calculate the
to
of
of the
et
al.'
or
particular
mechanisms.
action
pathways
are
mechanisms, specifying
well
as
Kinds
We
one
another
been
(cf.
the
al.
et
which
of
presented
species
of
‘phases’
succession,
between
pathways,
predict
or
vegetation change;
see
particular
successions
(ii)
of
is
cause
successional
of
sets
an
sere
an
is
and/or
causes
successional
explain
by
causes,
aspects
mechanism is
(iii)
a
describe and
to
of
a
Prach
to
(1992).
explain
on
rate
or
vegetation
change. Consequently,
particular
various
combining
between the mechanisms and the
pathway
as
1987).
between different types of
Burrows
some
from Muller-Dombois &
While
pathway.
attention
information
paid
we
succession.
‘classes’ of
vs.
prefer
clearly separate
Glenn-Lewin
vegetation
‘to
kinds
all
encompass
&
Van
dynamics,
We do
kinds
der
thus
scale of time and space: fluctuations,
Masing (1987) had proposed
community
and ecosystem
of jargon.
Overall,
McIntosh
(1980)
simplicity,
which
as
far
that
is
it
of
the basis
not
take this
of succession
Maarel
(1992)
providing
fine-scale
as
is
a
dynamics’,
open
on
not.
vegetation
causes
proposed
dynamics,
gap
vs.
The
change
areas
already
and mechanisms
into consideration
and mechanisms of
distinguish
to
on
an
seven
increasing
patch dynamics, cyclic
and secular succession.
Similarly,
‘spatio-temporal classification of several kinds
but these authors
provided
terminology
illusion
that
traditional in
areas
causes
proposal
vs.
phenomenological
an
of
framework based
a
succession, secondary succession, primary succession,
Zobel &
does
vegetation’, distinguished five ‘models’ of vegetation change,
allogenic influences).
to
have
we
terminology,
mechanisms, but actually
on
by vegetation), for another part
(e.g. autogenic
Ellenberg (1974)
literature,
phenomenological
to
the basis of the initial condition (non-vegetated
on
the
studying
statement.
attempt
an
in all kinds of
covered
because
in
(1990),
of
amount
may illustrate this
phenomena
partly
the
by
implicitly suggests
following
of
turnover
‘stages’
in
change
a
who
(1993),
of
knowledge
our
of
distinguish
to
construct
relationships
Fresco
al.
et
from
change
detect
the
concept of
might depend
of
to
to
for successional patterns;
successional
course
as
dilemma
describe,
to
the notion of‘kinds of succession’
surprised
fully
the
of the
change
conceptual
the
consider the notions
not
how
al.
et
of succession
(1987) proposal
which
is
GROOTJANS
site where climate
only
not
P.
of succession: phenomenology
adopt
distinguish
to
to
a
the
predict
or
rate
avoiding
responsible
pathway,
Models
Huisman
to
is the temporal pattern
interaction that contributes
type of
s
left
A.
single
a
is
contributing
as
problem
and Glenn-Lewin
(1987a)
AND
between trends and fluctuations, and
history
vegetation
(i) pathway
agent, circumstance
a
thus
of
succession
succession
only
refer
to
BAKKER
direction. We do
some
distinguish
to
review
a
mechanisms
and
suffices
another,
to
The
1989).
of models
set
degressive
or
Bakker
type. Here it
vegetation
et
regressive
(cf.
P.
physiognomy
that
implies
but should have
climax,
to
J.
ANDEL,
vegetation
constant’. This
effectively
pioneer stage
of
and
species composition
remains
VAN
to
science,
think
can
be
is
the
us
with another
concerned,
search
satisfied
by
for
we
regularity
replacing
set
agree with
and
terms
or
introducing ‘new-speak’.
The distinction of
understanding
two
largely depends
past.
Secondly,
primary
vs.
secondary
succession
only
of mechanisms of succession. First of all,
our
on
an
interpretation
mechanisms of
of what
succession
marginally
initially might
depend
much
contributes
to
the difference between the
more
have
on
happened
the
in the
initial abiotic
MECHANISMS
conditions
(e.g.
driven
whereas
successions
primary
debate
begin
that
it
is
succession
on
obviously
(such
kinds of succession and
primary
be
can
succession
primary
or
although
critical
being
as
(1990),
soils
relate
to
on
seres).
have
of
in
more
soils.
will
we
this
From
to
secondary
with
common
succession
Therefore,
mechanisms
on
that
recognized
rich
mechanisms
secondary
focusing
may
successions
secondary
mechanisms related
primarily by
however,
type of
kinds
to
of
discussing
stop
different hierarchical levels.
at
mechanisms and functions
Causes,
At
Tilman
and
start
is
(1992),
Peet
involving environmental change (soil, climate change,
confusing
primary
as
succession
a
view.
considers
changes,
nutrient-impoverished
than with
this
in environmental conditions
changes
Gleeson &
successions
whether
on
illustrate
succession would be driven
secondary
interactions.
species
small
may
successional
mechanisms
by
than
fertility)
examples
relatively
driving long-term
principally
etc.),
soil
following
that
recognizing
417
SUCCESSION
climate,
The
secondary.
factors
OF
which
level
According
to
of
community
Tilman
but idealized ends of
enological
to
a
‘It is
spectrum:
mechanistic
The
conceptual
occurred
In his
derived from comparative
determine whether correlations
related
problems
‘because
the
causal
used
from
for
phenom-
explaining
formulate testable
to
us
any
the usual
only through
manipulative experiments
Tilman
relationships.
short-term
mechanisms?
absolute entities
spectrum
optimal
allows
and
of
interpretation
have
the
for
not
are
will be found
view, theory
on
search
to
that is
point
a
observations,
based
are
to
ecologists
likely that, along
optimal point
trial-and-error process of science.’
have
we
and mechanism
there will be
theory,
given ecological pattern. (...)
predictions
do
organization
(1989), phenomenology
many of which have
experiments,
experiments
can
treated
(1989)
address
to
long-term
questions’.
Long-term
and
Beeftink
Only
and
observations
fluctuations
1987; Bakker
the basis of
on
of the relevant
mechanisms of
1992).
this
need
as
(see
ecosystem
to
be
Olff et al.
or
Roozen & Westhoff
succession
is
& Nisbet
attention has been
factors,
effects’
(Grubb
changes
in
paid
to
higher plants
salinity,
nutrient
on
we
&
autogenic
and
biotic and
abiotic
& Van der Maarel
vegetation
supply, quantity
farther than this and
allogenic
applied
rather than to the successions.
are
present in all successions’.
my
succession.
suggested
It is the
1991;
al.
not
‘relay
to
De Leeuw
can
fact,
(...)
new
I
Changes
composition
‘In 19261
I think
This classical view
that
has
now
in
terms
think,
of
the
a
succession’
to
species
allogenic
party
the influences of
in abiotic
factors,
groundwater,
proposed
to
e.g.
may also
distinguish
that I should have gone
first
autogenic
not
In
floristics’ and
So-called ‘third
1992) represent
succession.
and
find
represent
1992), ‘obligate
aspects.
sere
unravel
1993).
1980). With regard
Slatyer
allogenically induce succession. Tansley (1935) stated;
between
successional
a
does
et
may refer
al.
1993).
chronosequences.
frequently
et
1983;
al.
‘life histories’ and ‘vital attributes’ of
1976; Noble
1986; Glenn-Lewin
biota other than
1954;
i.e.
De Leeuw
Wilson
al.
et
be tested if we
can
succession,
1985;
concerned,
pre-emption’ (Egler
1973; Pickett
of
et
experimentally
to
Olff & Bakker
1990;
zonation
spatial
‘competitive hierarchy’ (Horn 1976),
(Drury
stages
Tooren
in
species
attempt
we
mechanisms of succession
that
aware
sequence of
can
Van
1982;
Turner 1990; Huisman
1990;
exact
processes,
situation,
e.g.
autogenic
‘initial floristics and
and
of the
knowledge
al.
et
between successions
distinguish
non to
Pickett
1987;
1969,
species replacement (e.g.
chronosequence
far
conditio sine qua
definitely represent phases
context we
As
a
1989; Andresen
In the actual field
sites which
are
Westhoff
(e.g.
been
place
and
to
the factors
allogenic
factors
sufficiently recognized
418
J.
until
tolerance
to
al.
et
Connell &
and
attempting
Peet
Pickett
recently.
proposed by
them in that
of
course
additional problem
a
successional
sequence.
causes
The
Pickett
(1987),
al.
et
study
anisms
of
view,
succession’
mechanisms and
and
it
long
and
autogenic
as
individuals
the
being
field
in
actors
which they
succession
vegetation
1935; Odum
the
O’Neill
1983;
analogy,
a
and
same
that
reason,
called ‘scenarios’
Farrell
may
An
1991).
between the
distinguish
that the
(1987) suggested
within
vary
successional
a
between
distinguishing
not
environment:
ability;
a
a
a
or
have
to
al.
et
fulfil
1986),
from
interest,
in
i.e.
a
allele in the
a
other
these
context
&
Slatkin
selection:
of the
mix of
Plant
is the
mechanisms of
discovered
(cf.
context
realized
a
in the land-
re-interpreted
that
the
aver-
of other groups and
competing
of
by
Tansley
of its function-
who
diseases
frequency
only
the ecosystem
context
context
competitors,
particular
context
‘Fisher
be
can
be held
functioning
(1992),
in the wider
of
the
However,
distinguished.
be
can
of its
can
a
‘mech-
as
mechanism
words,
wider
a
in
vegetation
Frank
context
of
clearly
be evaluated in
natural
the
In
to
an
delimited. From
this view
vegetation change,
functions.
of
be
according
processes
a
of
considered
succession’.
and
1992),
1987). Therefore,
quantity only
context
of
level
succession’ and
‘autogenic
terms
‘causes
can
be
can
study is clearly
as
the
on
1987). Indeed,
identify autogenic
to
ecosystem in the
of
of
influences
field
the
an
species
an
al.
depends
organization
Hobbie
e.g.
et
allogenic
useful
genotype in the
population;
(see
Allen
of the
use
influences
(Pickett
quote
we
group is
useful
seems
fundamental theorem
age fitness of
the
al.
et
from
1986;
if the system
but should also
ecosystem, and
For
Fisher’s
the
be
1987;
to
mechanism
of mechanisms of
being
reductionistic approach,
scape.
For the
to
facilitation,
seres
continuum of effects of earlier on later
level of
one
allogenic
considered ‘an efficient cause’
in
a
a
al.
et
that the
only
interact
causes
termed
on
implies
is useful
of
is
(O’Neill
mechanism
a
succession’
phytocentric point
ing
of
confusion results
that
addresses
another level. This
on
‘allogenic
action
unlikely
are
the three mechanisms
of each of the scenarios
Chapin
Connell
(Crawley 1986).
the main
interaction
specific
a
as
Slatyer (1977)
importance
&
GROOTJANS
‘models’ of
productive.
(cf. Walker
P.
and mechanisms.
organization
effect
be
to
A.
about entire
hypotheses
likely
extremes
AND
that
and directions of the interactions could
view,
our
sere
BARKER
emphasized
is that observational studies
strength
In
not
the relative
mechanisms represent the
the
species;
is
P.
mainly autogenic
alternative
way
Indeed,
different models of succession
proposed
i.e. the
the models of Connell &
(1992) suggested
in the
again
once
not
are
rather than mechanisms.
change
(1987)
J.
ANDEL,
Slatyer (1977),
inhibition,
test
VAN
and
food
avail-
genotypes within
competing
alleles
at
a
locus’.
CLARIFICATION
After
to
having
propose
will
start
up with
a
THE
FRAMEWORK
reviewed several concepts and the associated
reduction of the
from the
current
mechanisms
OF
phytocentric point
views,
and
jargon along
of view
but will finish with
causes
at
with
a
on
terminology,
we
feel the need
clarification. In the present
vegetation
succession
considering ecosystem
different hierarchical
levels
are
review,
in order
succession.
concerned,
we
As
to
we
link
far
as
take the
following position.
(a)
Interactions
mechanisms
between
of
plants,
succession.
i.e.
autogenic
processes,
will
be
termed
vegetation
MECHANISMS
Effects of biota other than
(b)
the
heading ‘biogenic
If autogenic
(c)
VEGETATION
established.
such
we
first
framework
et
longer distinguishable and it is
al.
leave the
we
mechanisms of succession’.
SUCCESSION
and
competition
after
only
occur
the
interactions.
In
broad
other
emergent
have
plants
mechanisms of colonization and
species
proposed by Pickett
will be treated under
that governs succession,
facilitation,
can
treat
thereafter mechanisms of
only
OF
as
community,
Therefore,
are no
two
discussing ‘ecosystem
start
plants,
plant
a
causes
these
community
a
succession’.
vegetation
MECHANISMS
between
of
properties
of
interaction between
view and
phytocentric
within
higher plants
causes
mechanisms and allogenic
the
particularly
Interactions
419
OF SUCCESSION
become
regeneration,
outline,
and
follow the
we
(1987).
Seed bank
Plant
species
can
bank
or
result of
as
become established
immigration
These processes
1990).
Buried
viable
attention
seed
important
are
context
Seed banks of
whether
Grime
their
not
or
related
longevity,
the
to
Therefore,
al.
et
(i) transient
seeds
of
adequate
seeds that
grasslands
seed
long-term persistent
the seed bank.
problems
et
al.
Sykora
seed
bank.
1983;
related
to
McDonald
the
rate
in chalk
or
This
of
on
nature
are
useful in
of
rare
is
an
the
less; (ii)
in
use
as
or
to
&
far
as
&
they
can
Blom
short-term
be
1992).
proposed (Bakker
distinguished
1989;
three types:
persistent
seeds
seeds
studies indicate that the
(Willems
see
if we do
der Valk
(Bakker
the process
in the
1983; Poschlod
Van
grasslands
of
1991),
1992),
1989)
have
not
dry
no
of re-establishment of
propagules,
rather than
landscape increasingly
poses
introduce them
(Gartner
spatial dispersal
urgent research
so
according
(Thompson
(iii) long-term persistent
immigration
species
land
little information
too
Voesenek
(e.g.
1987; but
reserves
in
the year
provides
Preliminary
that
scientific
plant communities,
‘transient’
or
throughout
grasslands
means
condition.
1992).
5 years;
years.
Maas
Constraints
of succession
to
1982), and moist
availability
1993).
of 1
least 5
at
depends
Fragmentation
of lack of seed
responses
first
a
received
change
to
These authors
1991).
period
a
species
in disturbed ecosystems
species
on
&
have
They
‘persistent’
as
in the soil for 1
year
persist
& Grime
who termed
(1916),
being
bank classification has been
1986; Pfadenhauer &
(Van Dijk
plant
disturbances
cf. also Poschlod
of characteristic
(Vyvey
vegetation.
of the latter
habitat
that remain viable in the soil for
majority
by
(‘nudation’)
detected
were
because estimates
1993;
the
been classified
that remain viable in the soil for
litter fens
of
Clements
this classification of seed banks
frequency
a more
Thompson
have
availability
Jimenez & Armesto
germinable
1979). However,
about the
site
present, either in the seed
are
neighbouring populations (Hodgson
of potential
1989a;
species
species
of conservation and restoration of
predictors
management (e.g. Peart
if viable
been mentioned
part
are
only
from
already
‘ecesis’,
banks
in the
mainly
had
and
‘migration’,
them
they
a
of
plant
propagules
as
topic.
Establishment
Once
seed
plants have become established
availability
Hester
(Grubb
et
al.
may become
1991).
1977)
is
a
limiting
and
In many ecosystems,
condition
to
set
for the
maintain
a
fruits,
microsite
regeneration
availability
from seeds
(e.g.
rather than
Peart 1989c;
permanent existence of ‘regeneration
species richness,
as
has been
niches’
shown for chalk
420
J.
grasslands
(Grubb
al.
et
&
De
be
to
seems
Vries
1992).
indicated
requirements during the
resource
of
concept
‘safe
term
P.
BARKER
&
(Harper
entire process of
A.
P.
niche
1977),
GROOTJANS
Rusch
1984;
is
for
1993),
& Russell
(Hutchings
regeneration
site’
AND
Verkaar
for salt-marshes
and
1992)
The
the
by
J.
ANDEL,
Schenkeveld
1982;
& Blom
flood-plains (Voesenek
Bakker
VAN
much
1989;
wider
than
this niche includes all
as
from dissemination
regeneration,
to
establishment.
Here,
do
we
feel
not
because
concerned,
syndromes’ (cf. Rozijn
the
germination
emerged
stage is
seedling
life of
active
for
over
germinated quickly
for
competition
productive
to
thought
survive
to
to
(Voesenek
in
these
a
in
germinating
soils, germinated
less and
at
successional
thus
alternating temperatures,
the
perceive
pioneer species require
conditions for
1980; Van Baalen
Bouwmeester
et
a
al.
1993).
In
Fenner
species (such
colonizing ability
Rather
1989b).
gradients,
et
for
in
example
al.
1987).
On
Gleeson & Tilman
suggesting
in
this
a
is
this
for
studies
on
were
the
different
of
capable
escaping
species,
delay
less
on
more
of
the
been
rich soils,
relatively
they responded
a
on
having
for
hay
successional
Grime &
(e.g.
1988; Pons
discussed the
clearly
germination
successions
bare
on
not
shrubs and
as
trees),
to
be
the
the basis
of
hypothesis
this
detected
trade-offbetween colonization
a
In
relationship
1992;
seeds
to
of
between seed
species (Fenner
species
Werner
1978;
successional
along
(1979)
primary
angustifolius
proposed
succession
on
the
poor
reproductive allocation,
decreasing
and
ability
al.
related
general,
and in Rhinanthus
a
et
as
smaller sized than the seeds
are
phenomenon,
of succession’.
(1990) similarly
in
soils,
but the
within
case
1975; Silvertown
of seeds
among different
straightforward
seems
Jarvis
1991; Voesenek
properties
Solidago species (Werner 1979)
‘colonization-competition
soil,
way
Therefore,
succession:
suggesting
pioneer species (mainly annual and perennial ruderals)
of later successional
(Ernst
is
that the
stimulus and/or alternating temperatures
Pons
(1987)
secondary
light
survival
seedling
1982; Van Tooren &
succession.
vegetation
Peart
1992).
cut
that these
slower rates, and
a
early photosynthetically
species,
Later
autumn.
‘germination
into
important aspect
from several
been
ecological
spring.
next
size and
al.
et
of the
had
grasslands
The
in such
secondary grassland
high percentage, suggesting
a
evolved
for
that,
arranged
1992).
report the results
(1994)
species
be
to
vulnerable part
most
25-years period. Early
light by
Seeds of many
to
al.
et
purposes,
a
stratification and
until
OUT
plant.
agricultural
of time
periods
considered the
characteristics of
germination
used
a
chance
a
1985;
(1979)
be
can
Van der Valk
is
germination physiology
as
Chabot
physiologies
are
far
as
&
Angevine
germination
physiologies
into detail
go
& Van Andel
have
seedlings
to
with
agree
species-specific
purposes,
that
the need
we
competitive ability (for nitrogen
case).
Facilitation
It
is well
two-way,
matous
1991),
known
plants
positive
phenomenon
water
and
Robinson
that plants of different species
effect. This
(Joenje
1978;
exemplified by
Schat
interactions between
of
hydraulic
nutrients
1991),
(De Hullu 1985)
‘Experiments
may be
in
and
or
1984; Laan
different
lift in semi-arid
through
interspecific
benefits from
can
et
al.
a
beneficial,
areas
(Caldwell
mycorrhizal
N-fixing plants,
soils
1989; Schat &
Vaccinium
after their death (Akkermans
dry grasslands
exhibit
the aeration of anoxic
species
et
al.
as
Van
1991),
or
Beckhoven
(Maillette
networks
either
one-way
by aerenchy-
1988),
the
the transport of
(Grime
et
al.
1987;
hosts for hemi-parasites
1971). We quote from Gigon (1990):
in Switzerland show that adult
plants
can
have
positive
MECHANISMS
OF
microclimatic and
SUCCESSION
effects
edaphic
Other experiments
species.
421
and
between adult individuals of many
not
direct but
via
other
of YAM
composition
to
abandoned fields
Within the scope of the
Connell &
if
Only
plant species,
know does
wood
not
plants,
concern
(Shigo
species
As
1975).
far
cite the
(1982)
N-fixation
which
succession.
on
europaeus
the
by
Bay (Crocker
et
al.
(Joenje
facilitating
of the
of
plants
are
et
initially
et
al.
(1980)
The
in
al.
majority
of
et
al.
fungal species
1993),
an
Miles
areas
and
(1987a)
from which
shrub
increase in the
examples
dead
on
Gerlach
1958;
and in reclaimed
are
in Glacier
e.g.
(Olson
of
forest,
of facilitation
1974),
we
Ulex
rate
Australian open
Major
in dune
1967),
of successional
examples,
an
to
(Horn
of facilitation we
example
showed
(see
is related
succession’
N-fixing legume
fire
a
poor soils
OUT
1991;
few
a
Facilitation
term
establishment
clear
the effects of the
on
Lawrence
‘obligate
concerned,
mentioned only
succession.
on
in
soil
indirect facilitation
indirect
Farrell
In
polder
this under the
leaving
the
but
(Miles
Tilman
1991), analogous
prefer
we
exhibit their beneficial effects
formation
(sensu
practice, however,
distinguished. Therefore,
term
plants
(direct facilitation),
soil
dynamics’
(see
competition.
will discuss
and
changes
‘soil-driven
of
processes
interaction
plant-to-plant
a
plant-induced
resulting
time
most
This
aspect.
and
between
distinguish
areas
1978).
Many
result
secondary
of
between soil
succession.
to
viz. the succession of
Grove
Westhoff & Van Oosten
1989;
The
should
we
leading
the latter
to
Macrozamia riedlei after
Major 1955;
&
al.
successions
primary
to
et
Similarly,
cycad
facilitate
may
related
of Bradshaw
study
those
adequate.
(1992)
are
species
positively
was
inter-relationship
facilitates the
fungi,
Glenn-Lewin & Van der Maarel
the
succession
during secondary
(Egler 1954)
higher
as
that
fungal community
succession,
on
and
directly
but
changed
showed
(1991)
density.
floristics’
facilitation
term
plant
the beneficial effects
most cases
al.
concomitant
a
YAM
coexistence
1916), ‘relay
plant
one
is the
indicating
1977) is confined
Slatyer
In
et
of the YAM
present paper
result in
‘reaction’ (Clements
1976).
diversity
and
properties, plant productivity
(sensu
Johnson
communities
fungal
forest. The
and establishment of other
germination
plant species.
organisms’.
related with soil C and N, thus
interactions which
the
on
literature survey show that positive interactions exist
a
The
the
heading ‘ecosystem
‘facilitation’ in
Hobbie
latter
favour
as
be
a
via
thus
1992),
might
called
distinction between direct and
direct and indirect facilitation
consider facilitation
to
not
indirect way,
an
1987b;
1988).
to
in
as
an
mechanisms of
of ecosystem
hardly
can
be
ecosystem process and
succession’,
terms
such
the
at
same
feedback
as
mechanisms.
Competition
for
Competition
succession
al.
1987;
Tilman
but
mechanism,
importance
1986).
of
Grace
the
have
they
1991)
(i)
resource
outcome
knowledge
long
Horn
1976;
None
competition
as:
has for
differ
in
in
of
and
been
their
or
the
not
claim
the
plant
an
1977;
communities
views of Grime
‘stress-tolerators’
The
1985) is
question
of the mechanism of
to
Grime
intensity
nature,
(cf.
and
(Grime
the main
mechanism of
important
competition
whether ‘maximum resource
requirement’ (Tilman
of succession.
of
as
Slatyer
authors
view
contrasting
(ii)
recognized
Connell &
these
discussed
whether
(Tilman 1987),
‘minimum
ing
1916;
1990).
(1990,
questions such
nutrients
resources
(Clements
and
be
Welden &
Tilman
to
the
the
et
only
relative
Slauson
by considering
1979) do compete for
capture’ (Grime 1987)
plant property
understand its role
as
or
in determin-
be discussed here is whether
competition
Pickett
1979;
to
a
we
should
mechanism
422
of
J.
succession.
vegetation
plant properties
competition
cession,
as
the
quantify
of
intensity
growth,
population
provides
as
(1993),
Elberse
feedback
Bobbink
in chalk
which
to
Berendse &
is
requirement’,
we
competition
for nutrients
establishment
gain
in
it
and
the
of
the
on
of
replacement
&
nutrient
increase in
is
abilities of
species
may
not
(Marrs
& Hicks
the
context
of
BIOGENIC
a
&
the
on
(1982,
Weissing
Slatyer
can
and
(1991)
tetralix by
Erica
species
as
Agnew (1992)
in
cycling
the ecosystem.
become
to
It is worth
for nutrients
concept of
1988)
more
much
as
result in
a
proposed by
model.
general
In
than is
complex
in
a
of
‘positive-feedback
which
succession,
change,
1988). The
But
be it
term
switches in
the
represents inhibition,
due
to
no
example,
for
competition
a
change
cyclic
(Watt
inhibition
at
there is
dominance. For
competitive
plant
competitive
community modifies
a
way, this
allelopathic competition.
of
prevention
particular aspect
treated
processes
successional
predomi-
mentioning
‘relative nutrient
more
the
meaning
may inhibit further
to
a
to
1993).
1977),
be considered
1986; Gimingham
climax
process-
experimental
of the latter
N-supply.
competition
based
between inhibition and
and nutrients in addition
this
words,
opposite of facilitation (or negative feedback switches). However,
distinguishing
which its
to
other
In
(1986). Examples
that the
suitable for itself. In
more
stands of Pteridium aquilinum
light
al.
competitive ability
of Tilman’s
which indicate all
environment, making
to
suc-
important
most
the level
resources).
competition for light is
(Huisman
Wilson &
1989b).
plant communities’,
discussing
we want
vegetation
The
Brachypodium pinnatum
an
which
case
by existing plants,
Peart
shown
showed
that
Connell
(sensu
in
importance
be found in Tilman & Wedin
that the model of
aware
if
a
is that it relates species responses
soil
et
when
hand,
as
question
refer here.
increase in
1987),
particular
a
should be
‘Inhibition’
ability (cf.
(1984,
competition
species
On the other
GROOTJANS
avoid the
population-community
can
only
result of
a
can
P.
habitat, and that this replacement is irreversible due
(1988)
as
of
A.
treat
we
parameter (R*:
O’Neill
grasses
we
to
should be known.
limiting
by
AND
levels,
relative
ecosystem
Molinia caerulea
al.
shows
(1993)
Aerts
an
of the
of
et
an
have
(1990)
grasslands
that Huisman
least the
its
between the
successional
on
eutrophication
positive
between,
and
reduce
advocated
Molinia caerulea results from
Similarly,
to
can
interface
an
research
Berendse &
nant
insufficient
succession.
plant competition
allocation)
approaches,
Wedin & Tilman
a
it
BARKER
competitive ability
vegetation
competition
equilibrium
in
functional
result of
Is
P.
of Tilman’s (1988) mechanistic approach
(dynamics,
follow-up
words:
determine the
mechanism of
a
the mechanisms of
advantage
concept
other
J.
ANDEL,
in this context? In view of the hierarchical
phenomenon
of which
In
VAN
in
competitive
1947,
seems
1955)
useful
only
or
in
stage of succession.
CAUSES
OF
VEGETATION
SUCCESSION
Herbivory
Vertebrate herbivores.
vegetation
depends
productivity (due
to
herbivores.
is assumed
to
level. In such
to
According
on
the
to
Oksanen
limiting nutrients
or
At intermediate levels of
be close
a
1987). Probably,
to
the
still
levels of
the
impact
soil-plant
of herbivores
system.
At
low temperatures), vegetation is
herbivore is
higher
(1990),
the
primary productivity
carrying capacity,
situation the
at
of
productivity
so
that the
‘prisoner
the
primary productivity
not
crop
on
the
primary
attractive
consumption
standing
of its food
low
is kept
supply’ (Drent
pressure
at a
low
& Prins
the herbivore densities
are
MECHANISMS
the
longer regulated by
no
vegetation
This
abiotic
has been
(Joenje
1985;
Olff
and
1992)
by lemmings (Batzli
herbivores
1993). As
Bakker
the
soon as
&
of
(deflections
in
Verkaar
species-rich
way)
a
&
and
Brown
competition
a
Fresco
and Carex
savannas
grazed
Prins
of
rate
1989;
among
lesser
responsible
on
longer
a
herbivores. In
a
sheep
could
grazing
succession,
vulgaris
after
grasslands,
Trirhabda spp.
succession
was
area
while
snow
to
(Kerbes
the
grazing
al.
et
vulgaris
(1990)
acidic soil.
for
of
plant
and
increases.
supports
1988,
a
for
on
feed
Diptera
A
similar
but in
their
by
modifications
the
intensity
of
distinctive and
a
review).
individual
on
a
of
cover
al.
et
increase in relative
on
dominant
conclusion, foliar-feeding
by
recycling
and
the
case
its
and
light
of
a
the
forbs,
was
a
or
is
applied
prairie
even
a
cyclic
underground,
chewing
period
to
on
com-
detect the
succession
vegetation
(Brown
larvae of
mainly
bare sandy and
of 3 years first resulted
thereafter in
a
relative increase
of
by above-ground Hemiptera
annual forbs.
a
in
by chrysomelid beetles,
fluctuation
which resulted
insects caused
in
beetle
microfloral/microfaunal
enhanced
on
result
heather
observed
was
early successional plants
during
may
faeces,
quality.
irreversibly replacing
grasses
have been
on
via
food
geese
densities of
complex
plants
grasses,
for
‘optimization
1992).
canadensis
Insecticides
of annual
as
higher photosyn-
the
of nutrient
These authors showed that
roots
of studies
majority
of increased fitness
consequences
regrowth
high
this
preferential sap-feeding
Cicadellidae feed
In
the
phenomenon
Elimination of these soil herbivores
to
avoided
intended for
in
on
temporal
a
grubbing
1990; Hik
heathland,
1983),
Gange 1991, 1992).
and
due
were
result
showed that
speeding-up
and forbs. The latter effect
grasses
herbivory,
to a
1987).
al.
et
relative increase of the
richness.
large
Jeugd
external factor,
land,
induced. In grasslands, most of the primary production
of insect herbivores
Coleoptera
phases,
by
der
dominated by Puccinellia phryganodes
attack of the dominant Solidago
(McBrien
patches
examples
consider
not
swards
phenology
time-scale
(Berdowski
an
Stanton
Brown &
perennial
& Van
succession
In the latter case,
account
did
on
few
a
evidence.
Jefferies
geese
related
the
for
only
ratio may
Hik &
Calluna
rhizosphere
munity (see
a
dune
1990),
browsed
some
depending
Lochmaea suturalis may result in subordinate perennial
in
and
review of the literature that the
convincing
was
destruction of the habitat
1990;
This
waterfowl
by
al.
et
Prins
vegetation
while other
kept intact,
defoliation. He
subspathacea,
was
However,
impact
either
as
1977), coastal tundras
On calcareous former arable
defoliation,
species.
grazing intensity’ by
and the
that the
1992).
leaf
higher
following
rate
Calluna
so
long
as
grazed
1989; Andresen
&
1986;
the
direction of
concluded from
(1988)
rate
and
grasslands,
of the
result of defoliation have
Insect
and
1980),
succession.
indicate adverse effects of
which
salt-marshes
e.g.
1985,
Georgiadis
stable
relatively
1957; Zeevalking
carrying capacity,
enabling
grazing (Gibson
growth
is
density of herbivores becomes reduced by
of the
thus
sheep,
thetic
parasites,
al. (1983) showed that moderate sheep grazing intensity resulted in patches of
et
restoration
a
ecosystems,
al.
et
grassland being intensively grazed
the
and
least retard further succession.
at
or
cattle (Bakker
(see McNaughton
below the level
of
variety
a
by rabbits (Watt
grazed
by predators
vegetation
biotic factors inhibit
or
shown for
grasslands
grazed
indicates that the
general picture
factors
but rather
vegetation,
into woodland.
develop
can
stress
423
OF SUCCESSION
slowing
in
In later successional
increase of species
an
of the
rate
of succession
grass-dominated sward, and thus resemble the well-documented effect of vertebrate
herbivores,
whereas
root-feeding
insects effected
an
increase in the
rate
of succession.
424
VAN
J.
J.
ANDEL,
BARKER
P.
AND
A.
P.
GROOTJANS
Parasitism
In
the
Dutch
cleared away
influence the
pathogens
they
die from
or
The
or
Alexander
mentioned
(1987)
parasitizing
plant
the
example,
and
&
the
soils.
result
Rosa
N-fixing
in
on
arenaria in coastal
and
soils,
appeared
also
thus
contrast
research
to
(semi)natural
culture,
on
dune
amount
on
have
might
of
rust
to
et
for
thus
This
1981).
nigra,
Ligustrum
on
reduced
Festuca rubra ssp.
root
system,
Otten
Empetrum nigrum
responsible
al.
reduce the
mycorrhizal
of
capable
are
Sambucus
to, e.g.
or
acidified
of
vitality
arenaria
(Van
1993).
interactions
almost
unfortunately
are
For
willows
stands
pure
This
They
areas.
related
favouring
plant-parasite
ecosystems
of the
Longidorus
genus
the
of succession
dunes,
selectively
succession.
the
succession,
1988;
plant-
well. Zadoks
as
seedlings
and other nutrients (Oremus &
Putten & Troelstra 1990; Van der Putten
In
the
in coastal
calcareous
nematodes
species,
fungi,
vegetation
if
became dominantand killed the willows
fungi
nodules,
few
formed nearly
the
or
interactions in
& Van Damme
a
the ice had receded.
During
proved (Den
develops differently
community
many
be
whether plants
not
pathogenic
which
once
spruce.
acceleration
rubiginosa
the
were
to
of whether pests
be,
only
by
that
killed
species
Nematodes of
1939).
of phosphate
Endoparasitic
Ammophila
der
the
to
Nooij
may accelerate
Hippophaë rhamnoides
uptake
may
and
vulgare
birch and
to
Wadsworth
including
reducing
showed
river Yukon in Alaska,
stands of
damaging
damage
pioneer
but various canker and heart rot
declined,
and
populations
which
De
dominated
community,
a
alexensis,
accelerated succession
system,
is
been
never
test
supposed
plant-pathogen
level (e.g.
Melampsora bigelowii
rust
S.
banks of the
(Baxter
in
species
willow
pulchra
gravel
both
studies
few
a
concerning
two-species
plant community
a
that ‘the crucial
plant communities has
of studies
the
relations may affect
pathogen
Salix
If
1990).
but this has
pests, but whether the community
on
is
Labyrinthula macrocystis
catastrophy,
of
submarine Zostera beds
evergreen
Harper (1990)
majority
natural communities focus
of
pathogen
composition
disease
excluded’.
are
1932. The
We agree with
1987).
Hartog
15 000 ha
some
for this almost irreversible
responsible
suffer
estuaries,
around
in
the
of
context
devoid of such
a
agri-
type of
investigations.
Food webs
McNaughton (1992)
has
that disturbance
proposed
at
level in
one
a
trophic
have effects far removed from the initiallocus of disturbance. Because of the
of food
webs, only
a
few
case
studies
are
to
be referred
in the
to
of
context
web
can
complexity
vegetation
succession.
Prins
Africa
&
ungulates
section,
the
such
(1991)
central
increased
an
effect
with
species
on
events
reconstructed
of
bush encroachment in
savanna
rinderpest pandemic
having
treated the
be
reported
aware
on
coast.
of the
Depending
symmetric
or
on
in
limpet
the
an
asymmetric,
on
Acacia tortilis
plants
intertidal
in the
former
community
activity
competition
but it is
East
among
effects, via animals.
foraging
limpets,
epidemics
even-aged
parasites
succession
decreased
of
of indirect
occurrence
of
study
Barnacles
abundance.
could be
a
direct effects of
anthrax
or
impala, which enabled the establishment
Oregon
algal
not
and
on
thereby
between
two
clear whether this has
algal abundance.
Robertson
herbivores
Jeugd (1993)
should also
we
barnacle
as
after
So,
Farrell
der
coinciding
as
stands.
Van
(1991)
studied mangrove forest ecosystems
consumed overall less
than 5%
of the
canopy
in
tropical
production,
Australia.
but the
Insect
rates
of
MECHANISMS
herbivory
OF
SUCCESSION
individual
on
species ranged
tree
Heritiera
marina,
(Avicennia
425
differences in chemical composition.
common
are
mangrove
species
tree
also contribute
messa)
playing
role in the
key
a
References
to
ECOSYSTEM
For
of
reasons
case
autogenic
latter,
confined ourselves
we
clearly
be
level
of
the
succession
to
of
entity
The
and
case
in
such
an
examples
succession and ecosystem
Dune
seres
in
meant
are
case
of abiotic
causes
two
thus
that the
recognize
to
the
of
context
complex
processes.
have been
we
succession.
distinguishing
With
regard
it is
allogenic factors,
and
interaction
(feedback
to
succession
make
a
can
longer
no
at
the
mechanisms)
is involved
choice for the
the
to
mechanisms
autogenic
types of process indistinguishable.
vegetation
illustrate the
to
with
succession,
primary
Westhoff & Van Oosten
and
causes
Cenlaurio
1991)
interface
increasing emphasis
This is
in ecosystem
‘ecosystem’
between
as
vegetation
the latter.
on
nigricantis
Saginetum
governing
the
rates
of
more
pathways
matter
(indirect facilitation).
blowing
The
saline
various
allogenic origin,
becomes
but
they
by
(Stal
1985;
(De
Wit
the
early
1989)
Visscher
1992).
establishment of
the main
plants,
the
autogenically
1988; Roelofs
et
al.
an
rate
Dieren
induced oxic environment around the
et
al.
from
1984) retard the succession
mollusc
shells,
and
cause, either
determinants of how
1991). High pH values,
to
secondary
a
the
bare and
(Pluis
1993),
on
beach
plains
1990).
After
hydrological regime
long
the
pioneer stage
low input of nutrients, and
plant
roots
(Ernst
higher productive
precipitation
are
organic
fixation of C and
Pluis & De Winder
of decalcification and the
of
seres
mats
important part by
1934;
i.e.
These factors
autogenic build-up
primary allogenic
(Van
baltici-
hydrological regime,
blowing.
formation of microbial
allogenic (and partly autogenic)
persist (Grootjans
carbonates
This is the
in blown-out dunes
or
Junco
beginning of these successional
which green algae and cyanobacteria play
1991;
allogenic
communities,
the local
on
and sand
may be modified by
the
1988,
the Parnassio-Juncetum atri-
conditions,
depend mainly
At the very
stabilized
pioneer
al.
et
between
association
calciphilous
from
develop
acidification, desalination,
mainly
among
exemplify the complex interaction
under
or,
of
sand
valleys (Westhotf 1947; Grootjans
may
may
Further successional
capilli.
dune
mechanisms.
autogenic
Schoenetum
can
Sesarma
leaves,
valleys
Successional
are
of mangrove
study.
few
next
where
(e.g.
because community components
causes,
that it is advisable
extent
in
place
vegetation
continuous
soil renders the
wetlands,
of
allogenic
the
influence the
can
crabs
the need
stress
takes
existing literature,
causes
in the
between
and
SUCCESSION
biogenic
to
because
vegetation,
vegetation
generally
the
distinguish
to
the
between
allogenic
distinguished. However,
worthwhile
OF
in touch with
remaining
to
sesarmid crabs
addition,
These
decomposition
serve to
vegetation
MECHANISMS
In
zone.
due
propagules of eight of the
propagules
processes and ecosystem
community
mechanisms and
insects.
35-0%
to
probably
area,
function’ of mangrove habitats.
studies may
of
leaf
20-80% of the
intertidal
the process of
(Excoecaria agallocha)
expanded
mangrove
the
across
to
development
interactions among
on
‘nursery ground
these few
and
patterning
largely
the
damaged by
were
species
from 03%
Moreover,
important post-dispersal predators
distribution patterns of
N
of
littoralis)
an
& Van der Ham
communities. Both
of calcite
may
act
as
426
J.
in the
buffering agents
formation
Moist
P.
(F.
thus
zone,
J.
ANDEL,
P.
BAKKER
preventing organic
AND
matter
A.
P.
GROOTJANS
deposition
and soil
Sival, personal communication).
grasslands
When intensively used
abandonedand
in
changes
vegetation
to restore
be
to
first
the former semi-natural
decreased. The
system
largely governed by
nutrients
limiting
was
hay
plant-soil
peaty soils appeared
in the Drentse A brook
agricultural grasslands
for
cut
of the
productivity
to
root
VAN
limited
N
by
succession
secondary
and
such
and
K,
by
later
P
the
species
of the entire
addition.
in
Holcus lanatus and
as
were
these
on
the response of the successional
1987; Olff 1992). The productivity
(Pegtel
indicators of eutrophic conditions,
valley system
species-rich hayfields,
As
result,
a
Agrostis stolonifera,
were
replaced by Plantago lanceolata, Anthoxanthum odoratum and Rhinanthus angustifolius,
which in
had
turn
to
make
room
for
a
acutiflorus
Juncus
dominated community
1989). Here, the competitive abilities of the successional species
nism) depend
on
the nutrient
autogenic influences
the
status
of the soil.
regard
the
to
and thus demand
indistinguishable,
are
With
an
(Bakker
mecha-
(an autogenic
latter, allogenic
and
ecosystem approach
of
soil system.
plant
Salt-marshes
Olff
reconstructed 200 years of
(1992)
Here, plant species replacement
of the
parts
of
progress
of
gradient, probably
course
which
nitrogen accumulation
of succession
the
relative
supposed
was
be
could
to
short
in
succession
than
had
the
to
in
the
of
higher
observed
rate
for
the
replaced
were
of
study
derived from
studying
the lower
on
clay.
(an
the
the
(an
sea
autogenic
and
as
responses
In
result
influences
of
understanding
a
mechanism)
allogenic
develop-
geomorphological
species
The
allogenic
succession.
species,
taller
autogenic
case
evidence for
more
be faster
vegetation
by
competition
this
to
of sedimentation of
sediments from the
clay
light
in
retrospect,
been
was
appeared responsible
Though
provided
ecosystem
of coastal bar island salt-marshes.
development
succession
plant species
importance
increase.
distinguished
of the
ment
due
rather than the desalination
cause)
the
during
and
vegetation
competitive
abilities.
Mire systems
Zobel
the
reviewed
(1988)
‘autogenic
term
processes
bog
succession in boreal mires. He
succession’
or
‘self-development
rather than vegetation succession in itself. He
comparison
with mineral land
deterministic
(though
not
communities, bog
tiate between external and internal factors.
determine autogenic processes, for
peat formation
processes,
e.g.
(Wheeler 1992)
Hornets
(1984)
development
logical
and
over
a
wider
example
area,
it is
when the water table
or
when
a
showed
may
conditions
allogenically
occur
of
a
in
bog
bog
it
is
likely
rising
very
Sphagnum
it
difficult
sometimes
may
for
we
comparable
dealing
are
faster than the
However,
difficult
growth.
to
without
and
by
to
that,
in
a more
differen-
possible
to
sequences in
with
are
allogenic
changes
encountered.
autogenic
knowledge
distinguish
Small
of
peat could accumulate
drained and unusual sequences
allogenic
seem
use
ecosystem
is characterized
where it is
that
regards
the conclusion
came to
searching
development
simultaneously.
induced
also
was
bog has been
that
Though
when
mires’
succession
unidirectional) development,
showed that his
clearly
of
between
in
the
types
of the
of
hydro-
autogenically
hydrological
MECHANISMS
regime
al.
OF
SUCCESSION
427
may result in acidification of the top
Wirdum
layer (Van
Van
1991;
Diggelen
et
1991).
succession
Bog
and
drain
may
structures
perform
of
growing
contribute
to
respect
processes
flow,
water
of
random
a
regime,
the
The
the
terms
arise
communicate
which
functions
with
of
of
succession,
discussions
species
moss
autogenic
on
undirectional
in
met
Masing (1987) put it:
is it
e.g.
caused
changes
the
‘In
unidirectional
a
and
hydrologists
feedback mechanisms with
by
carried
be
the
bogs
autogenic
certain
a
should
literature,
of
case
or
allogenic
or
type of research, because it should be simultaneously
As Zobel &
these
the continuous
explain
to
autogenic
terms
structures
hydrological
hydrological
Phytocentric
frequently
are
mire.
easily
can
that
so
formation,
structural characteristics of individual
on
nature
complex
which
bog
hydrological
by identifying
spatial scales.
discussions about the
or
a
the
in
ecologist
species.
such
entangle
of
parts
based
moss
During
expressed
An
bog.
various
on
be
can
such research
to
carpets
even
out
the
other
development,
ecosystem
meaningless.
flood
or
is
actually
become
allogenic
process
disturbance
considered
as
artificial’.
In the above
of
ecosystem
succession
ecosystem
appeared
it
examples,
succession.
could only
another review
(see
that
of ecosystem
the
study
vegetation
‘In
choosing
that
understand the
to
1979).
mechanisms of
But these
does enable
an
analysis
of
simulate
a
interest,
is
a
one
may
focus
on
the
this
However,
unit and limits one’s
single
size
one
of
(1983):
functions. We sometimes
forcing
that
require
emphasize
understanding
functions.
forcing
details of
to
would
this
as
are meant to
much
as
context
vegetation succession,
We quote from Odum
of
system
a
on
system
least
at
influences outside
concentrates
examples
approach.
interplay of pathways
model and
must
one
al.
processes
is reductionistic and
procedure
on
receive additional attention,
et
all its
by making
succession in the
study vegetation
to
does the mechanistic
as
the boundaries for
unit of interest
ability
e.g. Gorham
succession
useful
In the present paper
than the
larger
say
of
one
interest.’
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