A Succession of Paradigms in Ecology: Essentialism to Materialism and Probabilism
Author(s): Daniel Simberloff
Source: Synthese, Vol. 43, No. 1, Conceptual Issues in Ecology, Part I (Jan., 1980), pp. 3-39
Published by: Springer
Stable URL: http://www.jstor.org/stable/20111524
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SIMBERLOFF
DANIEL
A SUCCESSION
IN ECOLOGY:
ESSENTIALISM
AND
MATERIALISM
1. THE
IN
OF PARADIGMS
TO
PROBABILISM
MATERIALISTIC
REVOLUTION
AND
EVOLUTION
GENETICS
in his provocative
essay 'Darwin and Mendel-The
that by the time On the Origin of
Materialist
Revolution,'
suggests
was
the
in
notion of evolving
1859,
species was
Species
published
Embed
circles.
in
and
academic
both
established
lay
already firmly
Lewontin
(1974a),
in a matrix of new evolutionary
concept
and social
the arts, natural
sciences,
late eighteenth
and first half of the nineteenth
species
ding the evolutionary
areas of
in many
thought
sciences
centuries,
revolution.
during the
he points
to a different
the Darwinian
of
significance
worldview
was
The emerging
evolutionary
incompatible
the philosophical
tradition, stretching back to the Greeks, which,
still dominated
nineteenth
century
although
patently metaphysical,
with
viz. Platonic
thought:
views
1963). Idealism
imperfect embodiments
1961,
formal
structures.
Plato
essentialism
and Aristotelian
idealism
the material
objects
of
(Popper
as
the world
or ideal
essences
of fundamental,
unchanging
drew the analogy of shadows cast on a cave
the objects we can
reflections which constitute
for the imperfect
of
with our senses,
sought his version
although Aristotle
perceive
some
not
in
and
essential
forms within
matter,
spectral
particulate
wall
realm. Lovejoy
(1959),
(1949), Peckham
(1936), Wiener
worldview
evolutionary
(1969) suggest that the nascent
in
a reaction
to metaphysical
essentialism,
especially
engendered
the
the way
for
which
and economics,
eventually
paved
politics
transcendental
and Ghiselin
Darwinian
Lewontin
revolution.
metaphysical
philosophy
ing to the same type have
those
of differences
Synthese 43 (1980) 3-39.
Copyright
?
a necessary
is that differences
that
observes
a different
between
the types
of Greek
consequence
between
objects belong
and ontogeny
from
ontology
themselves.
The
former
are
0039-7857/80/0431-0003 $03.70.
1980 by D. Reidel
Publishing
Co., Dordrecht,
Holland,
and Boston,
U.S.A.
4
DANIEL
SIMBERLOFF
the latter, and can only confound
'noise' or disturbance
beclouding
or ideal structure of the universe.
our attempts
to see the essential
in this view, is to see through this
and science,
The goal of philosophy
the ideal forms. Mayr
and to try to understand
variation
(1963)
outlines
the
thought
typological
dramatic
of the most
event,
evolutionary
or
essences,
then
eternally
unchanging,
in the production
less result
by
systematics,
our understanding
For if species
speciation.
biology,
particularly
this outlook
retarded
and how
to types,
correspond
of
domination
how
of
ideals, which
could evolution
were
by definition
at all, much
(type)? The
'type'
occur
a new
species
of taxonomy
are, of course, a legacy of typological
specimens
in defining
the difficulty
species which
ing, and epitomize
on
us.
imposes
philosophy
think
a
such
were
tied to a typological
universe,
they
long as evolutionists
an
were confronted
could
with
insoluble problem: what mechanism
a type de novo
1969, Lewontin
1974a)? Two
(Ghiselin
produce
are possible:
either an individual must change
solutions
type or an
So
individual
different
Lamarck
of a
of one type must produce
(or individuals)
offspring
were
The Origin.
before
solutions
type. Both
proposed
that use
the former, with the specific mechanism
suggested
to change
of particular
organs could cause an individual
with
necks
is
well
his
known,
(and therefore
type;
giraffe example
the latter,
St.
Hilaire
Geoffroy
adopted
types) gradually
changing.
in type said to occur at
with dramatic
and discontinuous
change
or disuse
reproduction.
that
argues
observed
But
they
entities
neither
failed
or
solution
because
forces.
was
they
Lamarck's
accepted,
rested on
theory
on
founded
of
and
Lewontin
inferred
but
inheritance
un
of
was
from
not
characteristics
observations
acquired
of
could
saltational
nature, and though Geoffroy's
proposal
speciation
as evidence
the rare occurence
of unusual
variants
have adduced
it did not. Instead,
he posited
of normal parents,
among offspring
in type. Both
unobserved
ideas, therefore, were
easily and
changes
won a
who
the
dismissed
ant-evolutionist
Cuvier,
by
persuasively
The revolution
of Darwin
debate with Geoffroy.
widely
publicized
in examining
the individual variation
and Wallace
consisted
precisely
as "noise,"
rather than types,
which
been discarded
had previously
or ideals.
in
this variation
Instead
of viewing
essences,
among
A SUCCESSION
5
IN ECOLOGY
OF PARADIGMS
as a hindrance,
they took it to be the proper focus of study.
individuals
and variation between
between
Variation
species were no
In short, the
connected.
but rather were
causally
longer distinct,
a species were
to
differences
between
individuals within
converted
dividuals
between
differences
on a trivial
the mechanism
rested
of conversion
three
individuals.
of
And
species.
material
easily observed,
properties
of course, was that of Malthus
(1798),
syllogism
plus
The syllogism,
and that it independently
inspired Wallace
as
is a
it
did
Darwin
1972) precisely
(Ghiselin
reflection
1969, McKinney
of a widspread
in all areas of
of Greek metaphysics
by materialism
replacement
intertwined with
human thought (Barzun
1958) which was intricately
to above. The Dar
alluded
the burgeoning
of evolutionary
notions
winian
revolution
materialism,
materialistic
and
was
to become
it
is appropriate
formulation,
resting
should be at its root.1
and sex,
The three observations
Wallace
on
the
leading edge
that Malthus's
rise
of
consummately
for food
the requirement
in nature which
individuals
for the Malthusian
required
on
solely
this
of
Darwin
to produce
syllogism
and
evolution
were:
1.
Within
2.
siology,
Different
3.
The
a species,
individuals
and behavior.
vary
in morphology,
produce different numbers
is at least partly heritable.
variants
variation
are apparent
from even a casual
mechanism
The
Darwinian
breeding.
All
observation
of offspring.
of animal
no divine
phy
and plant
forces
or vital
requires
or essences,
nor the unobserved
as do orthogenetic
interpretations,
notes
events
Lewontin
of Lamarck's
and Geoffroy's
explanations.
that the Darwinian
than the
revolution was even more revolutionary
earlier
Newtonian
as Newton's
individual
one,
ideal
material
bodies
objects
for
it lacked
constructs
such
hypothetical
ideal trajectories
from which
following
varied somewhat.
in defending
his
difficulty
evolutionary
subsequent
nature
in
the
of
mechanism
of natural selection
lay
points (1) and (3)
it
It was required that, whatever
the mechanism
of heredity,
above.
Darwin's
must
of
allow both
parent
and
the production
offspring
(the
of variant
functional
offspring and the similarity
of
definition
'heritable').
6
DANIEL
the variation
Futhermore,
The problem which Darwin
but one never satisfactorily
of producing
to Lewontin,
capable
both
is
SIMBERLOFF
must be heritable.
itself, once produced,
and all other nineteenth
century biologists
resolved was to find a genetic mechanism
The
and difference.
similarity
can be
that Darwin's
failure
irony, ac
attributed
cording
the materialistic
outlook which
directly to his not applying to genetics
as did other nineteenth
served him so well for evolution. He believed,
that inheritance was generally
century naturalists,
blending, with the
characteristics
this belief
a blend of their parents'
of offspring
is readily
traced to Darwin's
attachment
(Ghiselin
1969). This mode
thought
typological
ill for Darwin's
notion
of natural
selection
for
it meant
of
of
that all new
characteristics;
to essentialist,
inheritance
variants
however
were
origin and significance
in the twentieth
century.
genetics
their
before
unrecognized
a
it arose,
the traits of
variation,
lost, or at least vitiated,
through blending with
the 'normal' mate. Mutants
(called
'sports') were not viewed
source
of
since
variation,
they were only recognized
important
were
abnormal
(and usually
they
poorly fit); in any
grossly
population,
would be
boded
within
the
as an
when
event,
rise of
was
with
this apparent
concerned
gap in his
increasingly
Whereas
the
first
the
of
stressed
edition
The
(1859)
theory.
Origin
evidence
for evolutionary
of natural selec
change and the operation
an overriding
tion, the fifth (1867) and sixth (1896) editions betrayed
Darwin
desire
to account
for the variation,
of inheritance
and did
so primarily
through
an
the
of
characteristics,
acceptance
explicit
acquired
no
the first edition discounted!
Darwin produced
very doctrine which
new material
to support the Lamarckian
evidence
view, and perhaps
the most poignant manifestation
the Darwinian
of the crisis in which
paradigm
found
itself
was
Darwin's
of Pan
Theory
under Domestication
'Provisional
in The Variation
and Plants
of Animals
at
to Nathaniel
back
least
(1651),
(1868). Pangenesis,
Highmore
dating
to Hippocrates
and in outline
entities
(400 B.C.),
tiny
postulates
that arise in particular
(Darwin called them 'gemmules')
parts of the
genesis'
the characteristics
of the part in which
body, and confer
the gemmules
that at reproduction
Darwin
hypothesized
from the different body parts to the reproductive
organs,
they arise.
are carried
there
to be
A
IN
PARADIGMS
OF
SUCCESSION
ECOLOGY
7
in sperm and eggs. This mechanism
allows the inheritance of
for it provides a means by which environment
acquired characteristics
to gametic cells.
ally induced changes in somatic cells can be transmitted
packed
came
Darwin
Though
his
to view
as
pangenesis
salvation
and
hypothesis
it as a reasonable
defends
century
Lewontin
both
and
his
favorite
Ghiselin
(Irvine
1955),
(1969)
in the light of nineteenth
hypothesis,
to explain
certain
observations,
empirical
observes
that it was a retreat to idealism or
knowledge,
perceptively
the unseen
ideal
gemmules
constituting
egregiously
entities. More
remarkably, Darwin made a number
essence-conferring
to a solution
of material
which had already
led Mendel
observations
essentialism,
to Darwin's
recorded
dilemma,
yet failed to recognize
that inheritance
need not be blending
but
He
significance.
rather that traits
sharply, that one
to manifest
the second
over the other,
trait may be dominant
itself only after two generations
of
:
even
an
1
3
two
and
ratio
of
(Irvine 1955),
approximate
two
in the second generation
of interbreeding
between
contrast
may
their
causing
interbreeding
flower types
varieties
of snapdragons
(Lewontin
1974a). His failure, and those of
to a
others doing similar breeding
Lewontin
attributes
experiments,2
on
as
Platonic
of
wholes
rather
idealism, focussing
groups
offspring
than on the physical
variation among the component
individuals.
Such
proceeded
logically from a world view of types (ideals) and
these types. Further, a concomitant
individuals
reflecting
imperfectly
a focus
of
this
view
between
was
the types
to seek the causes
of
attempt
causes
from the
of variation
separately
the
statistics,
(species).
Group
as
reported
type descriptions,
types
traspecific
Mendel
particularly
precluding
differences
within
the
were
means,
routinely
an examination
of in
variation.
alone
realized
that
both
and variation
similarity
on
and by concentrating
are
the
by the same mechanism,
individual offspring
of the same cross he deduced
among
not blending. Key to this deduction was
that inheritance
is particulate,
on individual differences.
his emphasis
Lewontin
observes
that Men
produced
variation
del
nowhere
offspring
cohort,
and
characterized
their variation.
had no place
in Mendel's
the
of
average
appearance
as a summary
statistic of a
scheme. Further, he kept offspring
separately
The mean,
8
DANIEL
crosses
of different
Both
of
appearance.
those of biometricians
terization
two papers
led him
SIMBERLOFF
even when
the parents
had identical
separate,
were
in contradistinction
to
these techniques
use of group charac
such as Galton, whose
to a blending
theory
of
inheritance.
until
1870) lay utterly unrecognized
nature of his materialistic
focus
truly revolutionary
or
as
Just
differences
discontinuity.
surely, his
(1866,
That Mendel's
1900 reflects
the
on individuals
and
simultaneous
and
in 1900 by DeVries,
and Tschermak
Correns,
discovery
in the largest sense the progress of the ongoing material
istic revolution
in all disciplines,
and more directly
that the sort of
independent
demonstrates
led to the recognition
that
inevitably
being done on heredity
is particulate
and to the basic mathematics
of genetic
in the Reports
In their papers
segregation.
of the German Botanical
in this
Mendel's
three
discovers
credited
him with primacy
all
Society
research
inheritance
but it is transparent
that he really had no effect at all on
recognition,
his retreat
the revolution
that bears his name. Merton
(1973) discusses
from research after failure to receive credit for his discovery.
II.
The
AND
PROBABILISM
materialist
THE
in evolution
revolution
with Mendel's
completed
ther developments
were
within
SYNTHESIS
NEO-DARWINIAN
and
genetics
I would
two of
required,
a permeation
of biology
which
(called
I have
insofar
as
they
could
only
of
'genes' by Johannsen
infer
quite
fur
advances
by an aspect of the
not yet discussed.
shown by geneticists
although
subsequently
(primarily
to be material
and Bridges)
bits of observable
Morgan,
were
or essences
still ideal constructs
for Mendel
adherents,
entities by
not
that three
argue
them technical
rediscovery.
the other
genetics,
based
rise
of materialism
broadly
the
of Mendel
'factors'
First,
was
Boveri,
in 1903),
Sutton,
chromosomes,
and his three
the existence
of
such
at morphologies
Only with
living organisms.
were
the
of
the
that
chromosomes
(1903)
cytologists
insight
was
the
the
in
of
the
materialist
revolution
(or bore)
genes
geneticists
in physical
to be firmly grounded
observation.
genetics
beginning
on a materialistic
Even
this insight did not completely
put genetics
looking
Sutton's
basis,
for starting
with
Johannsen's
distinction
(1903,
1909) between
A
SUCCESSION
OF
IN
PARADIGMS
9
ECOLOGY
came a growing
'phenotype'
recognition
we
and
behavior
which
observe
physiology,
and
'genotype'
morphology,
direct, one-to-one
that
the
are not
a
the product of a
and fundamentally
interaction
between
genes
complex
inseparable
and environment.
(1957, 1974, 1975) did most to elucidate
Waddington
this interaction,
and his and other work,
summarized
by Lewontin
translation
of the genes,
but rather
there
(1974b, particularly
chapter 1), indicates that in every generation
are four transformations
the genotype
and phenotype.
determining
even more
Two of these are steps between
and phenotype;
genotype
I will return
(a point to which
important
involves a completely
specified, deterministic
The
second
advance
technical
required
to be both
and unified
complete
and ultimately material
revolution
the particulate
continuum
of physical
how
which
Darwin
claimed
that natural
research,
are translated
and repressors,
effects,
operators
environment
during development,
was
by Morgan
genetics
thirty years of this century.
Even more fundamental
than these
four
the Darwin-Mendel
an understanding
genes could produce
of
the
in nature,
and on
an
area
acts. This is still
observed
selection
and
the
was
the
discrete
position
of genes with
are a few. But the
interaction
just discussed,
that continuous
variation
first clear demonstration
Mendelian
for
of
are known by which
and many means
into continuous
variation:
polygenicity,
of active
genes
that was
traits
soon), none
outcome.
and his
is compatible with
in the first
colleagues
technical
advances
to the com
was
revolution
in genetics
and evolution
pletion of the materialistic
the recognition
that evolution,
and its underlying
population
genetics,
are stochastic
a mathema
and not deterministic
processes.
Statistics,
tics of variation
comparison
was, until
view
of
with
variation?
study,
other
calculus.
to accord
seemed
indeterminism
branches
a scientific
recently,
success
the apparent
of
deterministic
alistic
and
It should
When
or probability,
late in
developed
of mathematics
(Robbins
1974) and
in
This
is not surprising,
stepchild.
the Newtonian
Newton's
so well
with
be very
clear
ideal
nature, why
that statistics
revolution
trajectories
focus on
with
its
and forces
individual
is inherently materi
'noise' as its object of
and anti-typological,
since it takes the
and not the type. Its two sixteenth century
beginnings
(Robbins
10
DANIEL
SIMBERLOFF
and the collection
of
success,
1974)-an
attempt to increase gambling
and industry of a state - are as materialis
data on population,
wealth,
it was an explicit
denial of typological
tic as can be. That
thought
success
in permeating
the various
for its meager
intellectual
until this century. An ideal or essential
universe
is ipso
disciplines
one.
a
deterministic
facto
accounts
mecha
century was dominated
by a deterministic
twin
ideal
Robbins
nics (Kac 1974,
1974), hypostatized
by
hypothetical
Demon. The former could pre
and Maxwell's
beings, Laplace's Demon
The
nineteenth
action-reaction
fashion the com
cause-and-effect,
the
and veloci
state
of
of
the
universe,
given knowledge
positions
plete
a
for
ties of all its particles
single instant. The latter could violate the
dict
in Newtonian,
second
motion
a perpetual
and in so doing, construct
the beginning
of the revolution
law of thermodynamics,
Kac
(1974)
machine.
in
determinism
against
velocities
of gas particles
It is almost
superfluous
Origin,
materialist
and
observation
that the
1859, with Maxwell's
are distributed
to a statistical
law.
according
to observe
that Barzun
of
Maxwell's
idealism
1859 was
that
views
the
and natural
in the arts
and
social
extends
the
revolution
have
been
observation
the year
it as
of The
signal year in the
Political
and
Economy
Critique of
to signify the
combining with The Origin
(1958)
with Marx's
revolution,
Tristan and Isolde
Wagner's
overthrow
dates
to
sciences.3
the
physical
in the physical
and it was
sciences
that an explicit
sciences,
recog
nature of the universe
nition of the probabilistic
led to the exorcism
of the demonic
reifications
of typology,
and idealism.
essentialism,
of
Aspects
this
exorcism
treated
(1967), and Klein
(1970). The key
Ehrenberg
Einstein's
(1)
Special Theory of Relativity
its consequence
that a measurement
affect the object
demons) would necessarily
with
(2) Heisenberg's
inherent property
taneously
particle,
(3) An
Uncertainty
of matter
both
Principle
precludes
and
velocity
measuring
much
less all of them.
modynamics
information
enunciated
theoretic
first
by Brillouin
tools were:
(1962),
(1905, cf. Lanczos
(such as those
1974),
the
by
measured.
(1927), which
either demon
location
of
that an
states
from
even
simul
a
single
law of ther
analog of the second
(1929), and elaborated
by Szilard
by
A
Shannon
which
sort
SUCCESSION
OF
(1948, cf. Tribus
states that Maxwell's
fast
and
and Mclrvine
Demon
slowly moving
law), and that the minumum
get the information
enabling
can possibly
derive
from
PARADIGMS
must
IN
11
ECOLOGY
1971) and Brillouin
(1962),
in order to
have information
the second
(and so to violate
particles
amount of energy which
it will need to
it to do
the ordered
its sorting will be more
than it
state which
the sorting would
produce.
from the physical
sciences,
generalizations
plus two counter
intuitive theories
Planck's Quantum
(1900, cf. Wilson
1944)
Theory
and Bohr's
the
Victorian
(1928)-destroyed
Complementarity
opti
These
mism, engendered
by the success of the Newtonian
an improved
would
allow an infinitely
technology
and readily understood
true, precise,
knowledge
world.
One
by
wrought
Theorem
might
this
(1931),
thetico-deductive
to emphasize
be
tempted
overthrow
of Newton,
that any logical system,
that
Revolution,
to
close approach
of
the
physical
the essential
pessimism
to point
to G?del's
the Greek hypo
including
as the crush
assertions,
and
one, contains
unprovable
But
is better served by observing
the
my purpose
ing, crowning
statements
of
the
allowed
three
tools.
In
each
form
by
exorcising
statement which can be made about the
the most complete
instance,
blow.
is a probabilistic
universe
of the physical
distribution
of possible
state of the world
of
states
statistical
(1977)
ministic
sketches
This
the
replacement
by a probabilistic
and pessimistic
probabilistic
physics
one:
a distribution
of probabilities
some
of
(or
it), or a specified
part
outcomes
of some event. Dirac
in this century
of an
one based on observable
ideal,
deter
quantities.
rev
the materialistic
aspect of
the
Robbins
sciences.
(1974) argues
spread beyond
physical
that it has finally caused us to think statistically
about all aspects of
our day-to-day
existence.
The key figure in transferring
probabilistic
was
to the biological
sciences
sciences
thinking from the physical
olution
1974) and though his attempt began in
(Singer 1959, Robbins
in 1853,1 would argue that its culmination
did not occur until
the Neo-Darwinian
1971, Allen
(Provine
1976)
synthesis of 1917-1930
which wedded Mendelian
to
The
Darwinian
main
evolution.
genetics
Qu?telet
earnest
architects
of this union,
independently
produced
and Wright,
Fisher, Haldane,
working
a probabilistic
conclusion
very much
quite
in the
12
DANIEL
spirit of the
sciences. That
concurrent
in the physical
of probabilism
victory
not surprising,
be so is, in retrospect,
since
themselves
in nature. His ratios of
statistical
this should
results
Mendel's
SIMBERLOFF
were
cross were
never
of a varietal
progeny
generation
3 : 1 ; rather for all seven traits they deviated
from
exactly
slightly
on
exact
3:1. Mendel was not misled by this deviation;
the
focussing
traits
in second
numbers
kind
each
of
of offspring,
as the outcome
as
stated
of events
his results
interpreted
the variation was not 'noise,'
fixed probability;
an ideal ratio, and the probabilistic
not postulate
of the materialistic
necessary
consequence
he clearly
above,
which
happen with
but expected.
He did
was a
interpretation
The Neo
outlook.4
that the best we can do is to
in brief, concluded
synthesis,
some distribution
of probabilities
Darwinian
specify
certain
of gene
themselves
and
genotype
(and
given
underlying
probabilistic)
on production
and survival of gametes
and zygotes,
and
of gametes.
Thus ended forever
the notion
that evolution
frequencies,
constraints
on unions
can be orthogenetic,
at least with much precision
at the level of genes.
one of the major
to
forces which
the synthesis
demonstrated
affect
the gene and genotype
drift-even
the
frequencies
genetic
direction
of frequency
and the firmest
change cannot be predicted,
statement we can make
is that the magnitude
of the change after a
For
a specified
fall within
probably
stated above,
is to determine
the
rules
the
also
which
and
(doubtless
by
genotype
probabilistic)
a phenotype.
environment
interact to produce
Whatever
is learned
about
these rules, we can say that the Neo-Darwinian
synthesis
specified number of generations
thrust now,
range. The major
the death-knell
sounded
in biology
for Newtonian
as
as
will
as
did
cause-and-effect
determinism
in the
relativity
complementarity
the same time. Further,
the synthesis was
in the ongoing materialist
Thomas
revolution.
Pyn
as
two
Rainbow
Mex
has
(1973)
antagonists
Roger
surely
at almost
sciences,
physical
a development
chon's
and
Gravity's
ico, apostle of the new physics,
and Effect.'
only with Cause
Poisson
distribution
normal
curve.
representing
and Pointsman,
the 'Antimexico,'
'left
the utility of the
Mexico
propounds
and characterizes
himself by the equation
for the
The
the
metaphoric
shift in human
of these
significance
I have
thought which
characters,
outlined,
is
A SUCCESSION
in Mexico's
transparent
OF PARADIGMS
IN ECOLOGY
13
manifesto:
.. .there's
a feeling about that cause-and-effect
may have been taken as far as it will go.
a less..
to carry on at all, it must
science
look for a less narrow,
.sterile set of
we have
come when
The next great breakthrough
the courage
to
may
assumptions.
and strike off at some other angle.
entirely,
junk cause-and-effect
That
for
III.
THE
RISE
OF
PROBABILISM
AND
MATERIALISM
IN
ECOLOGY
about half a century
later than genetics
and
has undergone,
so strikingly
a transformation
similar in both outline and
evolution,
detail that one can scarcely doubt its debt to the same materialistic
events
in this transfor
revolutions.
and probabilistic
Many major
Ecology
described
(1961) and Mclntosh
by Ponyatovskaya
to
both inside and
the
(1975,
relationship
developments
seems not to have been noticed. An initial emphasis
outside biology
on similarity of isolated communities,
replaced by concern about their
mation
have
been
1976), but
examination
of groups of populations,
largely superseded
belief
in
deterministic
succession
of
individual
by study
populations;
to
into
with
the
introduction
of
statistics
shifting,
ecology,
widespread
realization
is probabilistic;
and
that temporal community
development
differences;
a continuing
entities
rather
observable
struggle to focus on material,
I have described
for
than ideal constructs;
trends which
all parallel
and
evolution.
genetics
as a
first paradigm was the idea of the plant community
Ecology's
in the first American
by Clements
super organism,
propounded
ecology book (1905) and elaborated
by him in numerous
subsequent
crux
The
this
of
publications.
concept was that single species popu
lations in nature are integrated
into well-defined,
organic entities, and
that temporal
in a sere is
succession
key subsidiary
aspects were
an
to
of
and
utterly deterministic,
individual,
analogous
development
leads inevitably
to one of a few climax communities.
The relationship
between
the stylized,
and the deterministic
integrated
superorganism
as
it is organic and fundamental,
successional
development
producing
(1920): "When we have admitted the necessity
pointed out by Tansley
our natural units, we have to find
of first determining
empirically
them. This way we can only find in the concept
of
ways of grouping
development.
Development
of vegation
is a concrete
fact equal with
14
DANIEL
SIMBERLOFF
to which
the extent
Mclntosh
(1975, 1976) illustrates
a series of
dominated
until recently
with
ecology
paradigm
well-chosen
texts.5 Suffice it here to
quotes and an analysis of ecology
one
cite Shelford
of
the
animal
of the first
(1913),
ecologists
leading
its structure."
this
half of this century:
is the
Ecology
the environment
of communities.
science
to the natural
the field
without
conceived
of
phenomena
of ecology.
of a single species
to
study of the relations
to communities
and, in the end, unrelated
.is not properly
in
and community...
included
A
reference
its habitat
the superorganismic
community
Although
arose
in plant ecology
ministic
succession
to de
sequence
Candolle's
won
it quickly
mations,
pioneering
acolytes
(e.g., Naumann).
limnologists
on animal
work
symbioses
and
among zoologists
(e.g., Shelford)
as early as 1931 had aligned his
Allee
with
the superorganismic
paradigm,
or less characteristic
set of animals
which
are
but are integrated
communities."
assemblages
focus also led him to the notion of an 'integrated
a lake com
described
(1939). Thienemann
community'
accidental
Emerson's
similar
ecological
munity
more
"_a
observing
not mere
its deter
concept with
almost as a logical con
of plant
for
descriptions
so closed
"a unity
of the highest order"
as
in itself
that
it must
be
called
an
even
1975). Probably
(fide Mclntosh
organism
more
in
than this zoological
and limnological
support
important
was
as
a
the
Elton's
solidifying
concept
paradigm
superorganism
of the food chain as a conduit for community
energy flow.
description
of the superorganism
Elton himself was not an adherent
Although
to the
such a diagrammatic
view, his discovery
provided
analogy
an
was
it
of
individual
that
readily incorporated
organism
physiology
as an
in fact, one of the forces
integral part of the superorganism,
it organismic
and
cohesion.
(1927), Clements
Citing Elton
giving
Shelford
(1939) stress that trophic structure studies "can be utilized to
of each process
in the working
of the com
significance
as
a whole."
role of coaction
Further, "the universal
munity
[including
is to be seen in the integration of plant and animal
trophic interaction]
an organic complex, which
to constitute
relations
is characterized
by
reveal
a certain
the
degree
of dynamic
balance
in number
and effects."
A
OF
SUCCESSION
a formulation
IN ECOLOGY
PARADIGMS
15
in Russia
appeared
the
1961) at almost
it achieved
comparable
status (Mclntosh
that its sway is not to
1975), also suggests
paradigm
or the
to the force of Clements'
be attributed
personality
simply
volume of his publications.
prose and prodigious
Similarly,
persuasive
That
almost
identical
to Clements'
1910, 1915, 1931, in Ponyatovskaya
(Sukachev
same time as in the United
and that
States,
the appeal of the superorganism
(1926) states that his philosophy
that larger philosophical
argues
concept outside ecology
(e.g., Smuts
of holism was inspired by Clements)
forces militated
for its acceptance.
to this end is that the superorganism
the strongest
evidence
from
its
critics - notably Gleason
had,
eloquent
inception,
paradigm
(1910, 1917, 1926, 1939) in this country and Ramensky
(1910, 1925, in
and Negri
1961) in Russia, but also including Lenoble
Ponyatovskaya
Perhaps
in Europe-whose
1947 (Kormondy
the
Surely
was uniformly
or ignored until
dismissed
Mclntosh
1965,
1975).
ultimate
basis
for the superorganism
philosophical
work
is Greek metaphysics,
and this explains
its strong appeal in
and others. For the
of data-based
by Gleason
objections
one of a small number of distinct climax communities,
superorganism,
paradigm
the face
is
an
explicitly
classification
of
typological
an observed
construct
which
allows
Differences
among
category.
viewed as less important than the similarities which
different
classified
together, and are ontogenetically
between
categories.
organizing
(Sutherland
The
as
the
popular
cause
them
to be
from differences
as a reflection
latter are viewed
(such
relationships
1974) in a currently
immediate
into an already described
are
within
that category
community
individuals
of different
stable
'multiple
equilibria'
incarnation
of the superor
former, as in pre-Mendelian
1973)). The
concept
(cf. Holling
are
as
the result of minor
rather viewed
'noise,' probably
genetics,
in physical
like soil chemistry,
differences
environment,
during
in
And
the
deterministic
of
succession
the
strictest
development.
path
ganism
Clementsian
monoclimax
as is a Newtonian
formulation
is as much
an ideal abstraction
conceptualization,
is as appealing,
is a tidiness, an ease of
trajectory. There
particle
on perfact paths that
to well-defined
ideals moving
as it
in ecology
both aesthetically
and functionally,
was
and evolution.
in genetics
Unfortunately,
it is as poor
a descrip
16
DANIEL
tion of ecological
paradigm did not
and
structure
references
SIMBERLOFF
as of evolutionary
lead to mechanistic
of nature
is not
reality.
That
the superorganism
of the operation
understanding
(1975, and
surprising. As MacFadyen
in many sciences
it has proven possible
points out,
features
of a higher
level deterministically
while
emergent
are stochas
that the underlying,
recognizing
component
phenomena
held out the
tic; it is in this spirit that the deterministic
superorganism
therein)
to treat
of an adequate
description
But
of natural communities.
com
of the almost unimaginable
the
of
rich
underlying
supposition
plexity
an
and omnipotent
connectedness
and interaction
under
precluded
a danger
in treating
mechanisms,
standing of low-level,
probabilistic
"Those who are obsessed
of
any field holistically:
by the interactions
promise
everything
conclusions
with
everything
are not drawn
else..
from
.are of necessity
diffuse. Practical
the holistic contemplation
of totality"
(Pirie 1969).
The watershed
in ecology was
Cain,
revolution
and probabilistic
year for the materialistic
in
which
three
1947,
(Egler,
respected
plant ecologists
in Ecological
all published
and Mason)
papers
Monographs
the Clementsian
and citing Gleason's
attacking
forcefully
paradigm
as
of
the first articulation
association'
'individualistic
concept
plant
The
formal
to Mendel's
of their view
(Mclntosh
1975).
analogy
but even more
is an examination
of
enlightening
the specific reasons given for this dramatic change. Egler cites Raup
(1942) and Cain (1944) to the effect that the Clementsian
assumption
in community
is an a priori explana
of cause-and-effect
development
resurrection
is patent,
derived mechanism,
and he claims
tion, rather than an empirically
on a series of Hawaiian
com
that his own extended
observations
are completely
in accord with the individualistic
of
concept
avers
his
in
interest
'all but forgotten
Cain
"actual,
paper.'
on the ground," and scorns the 'hypo
specific communities
that the mono
Clementsian
Later he suggests
community.
munities
Gleason's
concrete,
thetical'
as originally
theory
a panchreston,
and
climax
become
strate
Reality?',
section
he bases
was
on
wrong,
local
had
studies
subsequently
will demon
In a
individualistic
of Gleason's
hypothesis.
'Does the Association
Have
entitled,
Objective
are
answer on materialism
his negative
("Species
the correctness
'heretical'
stated
that focus
A
are
.Environments
facts,..
OF
SUCCESSION
IN
PARADIGMS
facts.")
while
17
ECOLOGY
lambasting
the
"pre
in the abstract." He is at
of the reality of the association
conceptions
a
to
stress
material
is real,
that
existence,
pains
specific stand, having
a
not.
In
is
the
while
final
the association,
ideal,
only
hypothetical
a
construct
is fictitious
because
for Cain, the superorganism
analysis,
1) unlike
objective
enough
a species,
criterion
to belong
the
to the same
association.
Mason
also
stresses
that
a real entity, and
species population
a fortuitous
limited only
the community
abstraction,
of environmental
factors by its
'coincidence
of tolerance'
continuity
genetic
lack of it renders
by
it has no continuity
and 2) there is no
by descent,
are similar
two stands
when
for determining
component
this notion.
species.
renders
He,
the
too, credits
Gleason
as the first proponent
of
texts early in this decade
1973,
(Colinvaux
spate of ecology
et al. 1973, Krebs
all
Poole
1972,
1974, etc.)
agree
that,
the Gleasonian
years after its rehabilitation,
twenty-five
paradigm had
one (Mclntosh
overthrown
the Clementsian
1975). If one asks why
it did, two convergent
lines of research
the revolution
occurred when
The
Collier
appear
to have
necessitated
it. First,
the
facts
that
real
stands
and when such boundaries
do
lack well-defined
boundaries,
generally
with
in the
exist
associated
they are frequently
abrupt changes
were often noted by Gleason
and Ramensky,
environment,
physical
an intolerable
of dogma by
but by 1947 constituted
contradiction
observed
all attack the superorganism
fact. Egler, Cain, and Mason
on this basis using data from field studies. This type of
paradigm
was
in the next decade
extended
observation
by two in
greatly
dependent
Mclntosh
Curtis'
groups;
Curtis
1955,
1951,
and
continuum'
(Curtis
'vegetational
and
Whittaker's
1959)
'gradient analysis'
as a
of plants
the spatial distribution
(1956,
1967) both describe
of
of the individual,
responses
consequence
relatively uncoordinated
to
in
the
individual
without
environment,
species
gradients
physical
or dying as a unit. It may
need to invoke groups of species' persisting
that discrete
that this work
also demonstrates
observed
popu
continuum
lations acting individually produce a community
(whatever
as the Morgan
is used to characterize
much
statistic
communities)
etc. resolved
of polygenicity,
effects,
position
group's understanding
be
18
DANIEL
SIMBERLOFF
that phenotypes
the apparent Mendelian
are, for the most
paradox
while
the al?eles of a gene are discrete.
part, continuous
I would
to
line of research which,
The second
argue, contributed
was a shift in emphasis within
the demise of the superorganism
the
to
which
led, inevitably,
study of animal and microbial
populations
on individual populations
focussing
most two or three of them together,
as proper objects
of study, or at
rather than the entire community
in 1940 to attract
failure of Haskell
The
they are embedded.
niche
in a hypervolume
of the species'
interest
(Mclntosh
concept
per
1976) is a reflection of the lack of interest in studying populations
se. That Hutchinson's
an
in
1957
identical
has
suggestion
generated
in which
is an
and relationships
parameters
to which
the plant population
research
indication
of the extent
by
and
Whittaker
and
animal
Mason,
Cain,
Curtis,
Egler,
population
or
the population
and Park (La.) had legitimized
studies by Nicholson
enormous
literature
on
as an object
of the
of study,
independent
were key to this shift of
on real populations
to population.
interaction
few-species
community.
Experiments
focus from community
Population
progenitor
in ideal abstractions.
The
had its own origin
ecology
the
deterministic
of
logistic equation, which has dominated
right into this decade, was a paper by Verhulst
ecology
population
(1838) communicated,
figure in introducing
that the
growth
niche
ironically
by Qu?telet,
enough,
The irony,
statistics
into biology.
is as much
curve
logistic
as a Newtonian
trajectory
the
seminal
of course,
is
an
ideal description
of population
an
is but
idealization
of particle
movement.
The
logistic model
ignored until the early
exhumed
and exploited
was
of single population
growth
virtually
twentieth century, when
it was simultaneously
(1907, 1925), Pearl and Reed (1920),
by Lotka
(1926, 1928); the first and last authors even
the
for
into a pair
eponyms
equation, which
they elaborated
or
to
describe
of equations
predator-prey
parasite-host
relationships.
on
Gause
(1934) performed
interspecific
competition
experiments
Pearl
(1925),
and Volterra
became
his basis for Gause's
Law, or the Com
(ultimately
microorganisms
were
which
he
Exclusion
believed
petitive
Principle)
satisfactorily
system based on the logistic. The logical
by a two-equation
explained
A
SUCCESSION
OF
IN ECOLOGY
PARADIGMS
to this proliferation
of logistic
and complete
conclusion
did not come until 1968, when Levins'
lation descriptors
to describe
and predict
the workings
matrix
attempted
19
based
popu
community
of an entire
on the assumption
based
that all species
community,
populations
to
the
ideal logistic
law modified
adhered
only by equally
ideally
characterized
interactions.
pairwise
But
long before
on material
based
abstract,
doubter
this conclusion,
properties
particles.
reproducing
version
of the pr?dation
discrepancies
to a failure
of the logistic had set in,
as opposed
to
living organisms,
an
was
Gause
himself
(1934)
early
criticism
of
of
between
the
results
experimental
the mathematical
model
his
logistic, and attributed
and equation predictions
to describe
the
adequately
a
(1954b) makes a
biology of
particular pair of species. But Nicholson
more
of the underlying
shift in
and, from the standpoint
cogent
am
I
which
the
of
dis
thought
describing,
analysis
suggestive
a
can
He
demonstrates
that
the
crepancy.
only represent
equations
mean
of
two large populations
of an interaction between
("these
the statistical
effects of the interaction
of large
represent
in adequate
of individuals
and that
operating
space")
were
to
too
Gause's
with
small
experiments
display
high frequency
the statistical
Observe
that variation
among populations
expectation.
result
equations
numbers
or replicates
had
among individuals
as
little place
in logistic formulations
had for the pre-Mendelians.
own work
Nicholson's
(1933, 1947, 1954a, b, Nicholson
was
more
even
1935)
severely critical of Lotka-Volterra
His
tems,
models
early two and three-species
and more
his
later
importantly
of
as variation
and Bailey
type models.
insect
host-parasite
and models
experiments
sys
of
of Lucilia
all
(the sheep blowfly),
cuprina
populations
not
that
did
not
formulations
and
could
(because
emphasized
logistic
too few parameters)
any popu
they contain
realistically
represent
lations but those of the simplest microorganisms.
In particular,
the
single
species
invariant
while
carrying
capacity,
as
occur
the fact
discrete
organisms
and the lack of lag factors
of many
ideals, cleansed
asymptote,
of
aseptic
'noisiest')
biological
properties.
that the equation
the
individuals,
is continuous
non-decaying
the logistic the most
the most
(and
interesting
all render
Nicholson
of
noted
that, aside
from
the
20
DANIEL
of
inappropriateness
the
SIMBERLOFF
as a model
logistic
account
realism
certain
for a laboratory
it took
environment,
in a controlled
(such as his blowflies)
of climatic,
edaphic,
own mechanistic
models,
population
no account
In his
even
biotic, and evolutionary
Nicholson
attempted
influences.
partially
to
for these more
for
factors, sacrificing generality
idiosyncratic
an impressive
to population
data in
match
achieving
in light of his discussion
of Gause's
instances.
Interestingly,
and
were always deterministic;
variance was
models
results, Nicholson's
nor confidence
never calculated,
limits given for goodness-of-fit
of
any real data. Nevertheless,
by his focus on single or few populations,
in modelling
could only damage
and partial success
them, Nicholson
the
claim
that
earlier,
supporters,
quoted
superorganism
is the science of communities."
By the nature of his models,
the
of
"ecology
he also began
to break
the hold
of Greek
on population
metaphysics
ecology.
flour
Park's
beetle
research
ing how
physical
between
and
1954, 1962, Park, Leslie,
was
impact. The primary emphasis
but rather on examin
trajectories,
(1948,
Mertz
1964) had an even greater
not on modelling
ideal population
environmental
factors
two
and
affect
on
populations
(such as cannibalism,
poisoning
The most
the outcome
is achieved.
petition
mechanisms
the outcome
of com
the
physical
precise
of the medium,
etc.) by
of
revolutionary
aspect
was
certain
the discovery
that under
Park's
work,
however,
a specific outcome
environmental
could not be predicted;
conditions
one could possibly
the process was stochastic,
and the best prediction
was
a probability
that a
in the neo-Darwinan
mold,
make, very much
which
on this
that Park published
species would win. It is significant
with
statistician
of
his
work
the
(Neyman, Park, and
Neyman,
aspect
one
into
was
this
the
first
inroads
of statistics
for
of
Scott
1956),
given
an explicit
that pro
and constituted
recognition
ecology,
population
cesses on the population
Park's stochas
level are not deterministic.6
on the population
level
and his success
ticity was a breakthrough,
to popu
communities
only aid the shift from superorganismic
lations as an object of study.
outcome
is the
indeterminate
A measure
of the impact of Park's
could
disproportionate
emphasis
placed
on
this
aspect
of
the Tribolium
A
work
this
SUCCESSION
OF
IN
PARADIGMS
21
ECOLOGY
text cited
every
ecology
subsequent
of the nature of biological
causality
an example
of indeterminacy
caused by the
(Mertz
1972). Virtually
review
result, and Mayr's
it as
(1961) adduced
extreme complexity
of ecological
that this two-species
laboratory
(it is notable, however,
must
be among
the
competition
was
a
effort
mounted
Further,
major
interactions
simplest ecological
phenomena).
and Dempster
1960, Lerner
(e.g., Hardin
that the indeterminacy
1971) to demonstrate
and Dawson
1962, King
was illusory and could be
con
in Park's experimental
variation
for example, were at pains to show that
for contrasting
among the cultures could account
genetic differences
outcomes
here.
in different
'replicates.' Two points require comment
accounted
First,
for by uncontrolled
and Dempster,
Lerner
ditions.
increased
pletely
genetic
eliminate,
unpredictability
uniform
strains
(cf. Table
2 of Lerner
and Dempster
that demographic
(1976) show
from the standpoint
of popu
Second,
must
of genetic
be
degree
heterogeneity
supposedly
and
Cawthon,
is also important.
1962). Mertz,
stochasticity
some
in nature,
lations
a universal
considered
premise;
outcomes
different
reasons
greatly reduced, but did not com
in outcome
of competition
between
uniformity
for
on
a Poisson
these
Park
to explain away observed
attempting
to seeking
is analogous
the
in
buckets
in
the
raindrops
grounds
distribution
of
of each drop. Of course such
of the trajectory
specific determinants
exist and in principle
could be found (at least to the
determinants
But the information we
limits imposed by the uncertainty
principle).
are likely ever
of a stochastic
to have
the epitome
renders rainfall patterns
us well
serve
it
in
and
this
will
process,
viewing
light
I
and allow sufficient predictions.
would
argue that the genetic com
as the physical
will be as elusive
of natural populations
position
forces acting on precipitation,
and our understanding
of population
available
will require stochastic
treatment.
phenomana
in single species
interest
I do not wish
to imply that the new
was to the exclusion
of concern with community
proper
populations
as a collection
of the community
ties. Rather
the growing conception
of
species,
analytic
munity
that a superorganism,
of
than synthetic mode
rather
rather
properties
were
approached
led in some
community
through
assembly
quarters
research:
to an
com
of populations
22
DANIEL
and pairwise
population
died first. Mertz
(1972)
SIMBERLOFF
interaction
which
were
well stu
ostensibly
of this tack from the
instances
gives many
literature; two further examples will suffice here. First, the
determinants
of community
have
stability
long been
sought as a
matter
interest.
Clements
and theoretical
of both practical
(1936)
the increasing
ascribed
stages in a sere to an
stability of successional
Tribolium
com
and integration
of community
increasingly
tight organization
on
interest
has
focused
another
More
ponents.
community
recently,
as possibly
attribute,
stability. Goodman
conferring
species diversity,
in its simplest form is disproven
that this hypothesis
(1975) suggests
level
1977)), but even while other community
as potential
determinants
(1973)
(e.g. May's
a reductionist
of the community
matrix),
properties
(but cf. McNaughton
are sought
attributes
discussion
of
was
explanation
teristics
of one
charac
(1968): the population
by Frank
proposed
or a few critical
whether
the
determine
species
as a whole
to
is stable, by whatever
statistic one wishes
community
characterize
this notion,
(1975) strongly supports
stability. Goodman
is a fortuitous
of properties
that a community
consequence
property
of one or a few component
species.
Horn's
matrix
only
attempt (1975) to study
of transition probabilities
reduces
succession
between
as an exponentiation
tree species
individual
of a
not
to the level of populations,
but
phenomena
as a fundamental
successional
stochasticity
to a
succession
of deterministic
antithesis
leading
are
treatments
conceived
climax.
by Ste
Similarly
successional
introduces
explicitly
the
property,
superorganismic
and Waggoner
phens
(1970),
vations
and Stephens
(1970), Leak
(1970), Waggoner
obser
and Botkin,
Janak, and Wallis
(1972). Other detailed
is
at the population
level have also suggested
that succession
a population
Nisbet
and
1973), largely
(Drury
phenomenon
truly
determined
Needless
supports
by the effects
to say, the work
this view.
IV.
Finally,
one ought
THE
and
tolerances
of a few
of Curtis
and Whittaker
CURRENT
STRUGGLE
not to be left with
the feeling
critical
discussed
species.
earlier
that the materialistic,
A
PARADIGMS
OF
SUCCESSION
IN
23
ECOLOGY
is a fait accompli.
in ecology
revolution
Essentialism,
still rampant
and determinism
are, if not dominant,
idealism,
(Slo
is partly a reflection
of ecologists'
bodkin
1975). Their persistence
of their field compared
diffidence
because
of the apparent
sloppiness
probabilistic
to
is the curse of biology"
"physics-envy
scatters
of points
and jagged trajectories
cruves
for new island
the
colonization
(e.g.,
the physical
sciences;
The
1971).
large
(Cohen
which
typify
communities
ecology
and Wilson
(Simberloff
on different
of herbivores
the dispersion
of the
(Strong and Levin
species
is not quite so scientific as
1977)) seem to foster the view that ecology
a search
militate
for more
ideal
and
and
for
chemistry
physics,
a
sciences
themselves.
For example,
often from the physical
models,
number
of deter
during this decade has been the erection
as
some
if
into
rules for packing
communities,
clearly
species
bounded
'com
entity could be denoted
physical
permanently
topical
ministic
and
1970) and
plant
endeavor
In MacArthur's
in isolation.
and studied
summary work
an analogy of species-packing
to crystal packing,
(1972), he presents
Such an ideal, deterministic
suggested by Gordon Lark, a biochemist.
a
to
be
of
wide variety of proposals
appears
approach
symptomatic
on species-packing,
and May
(1974, fide Mclntosh
1976), a physicist
munity'
the leading figure in analytic
to an even
the metaphor
raises
1973),
(e.g., May
ecological modelling
the
eventual
of
establishment
many
higher level, envisioning
'perfect
in ecology
and the consequent
of ecology
emergence
crystal' models
as co-equal
to "the more conventional
(and more mature) branches of
who
turned-ecologist
science
and
has quickly
engineering...."
The
crystal-packing
do not appear
however,
a retreat
they constitute
action
coefficients
is still used
from
and
outcome
deterministic
point.
related
to have
models
for
species-packing,
successful;
strikingly
The community
matrix
certainly
of inter
the deterministic
to characterize
of
been
to idealism.
1968)
munity
properties
(e.g., Culver
and the community
in general
since it incorporates
statistical
borhood
become
(Levins
logistic equations
communities
and to explain com
1975). May's
treatment
(1973), appears
to produce
noise
of the matrix,
to be
revolutionary
a stochastic
neigh
rather than a single
community
dynamics,
But the underlying
equations
are
distressingly
24
DANIEL
SIMBERLOFF
is an ad hoc
ideal, and the noise distribution
he
form Q, which
quadratic
claims
suggestion. MacArthur's
minimizes
(1969,
competition
of logistic equations,
consequence
1970), is not only a deterministic
an entity
as Darwin's
but as metaphysical
Maxwell's
gemmules,
in statistical mechanics,
and Adam
hidden hand in
Demon
Smith's
it is condoned
economics.
(1969). Limiting
by Lewontin
Surprisingly,
from logistic equa
similarity L of coexisting
species was calculated
a data-based,
and Levins
tions by MacArthur
(1967);
despite
of its incorrectness
demonstration
mechanistic
(1973) and
by Dayton
a cogent, damning theoretical
as a possible
characteristic
treatment
of
nature
(1976) it is still cited
1976). Neill's
(e.g., Fraser
form of the logistic model
by Heck
showing that the multispecies
experiments
rests on untenable
(1974) seem not to have been heeded.
assumptions
the
from the
and independence
(1975) deplores
autonomy
MacFayden
to
the essentialist
world which
strain in ecology
appears
biological
have achieved,
but his explanation
truly
frequent failure to propose
- is
cause.
The
the proximate
ultimate
testable
only
hypotheses
the
is
the
of
Greek
worldview.
difficulty
tenacity
metaphysical
Another
of
the
manifestation
nascent
stochastic
of this tenacity
school
of
is the relative
independence
and
community
from
school
above,
ecology,
epitomized
text of ecology
A striking
differential
by logistic-based
equations.
in 1969,
from the probabilistic
by E.C. Pielou
viewpoint was produced
and though it attracted
favorable
reviews
and a few new
generally
population
the mathematical
discussed
to the stochastic
adherents
methods
dominate
are given
American
short
cause
shrift
ecology
1970), its ideas and
(e.g., Wangersky
texts which
the
by
logistic-oriented
in this decade;
earlier
stochastic
treat
ments by Chiang (1954, 1968), Skellam (1955), Bartlett (1957, 1960),
and Leslie
(1958), and Leslie
(1960), Leslie and Gower
Gower,
were
even
more
to
in these
oblivion
(1958, 1962)
summarily consigned
were
not more enthusiastic
than
quarters. Systems
analysis ecologists
Bartlett,
mathematical
cizing
systems
ones.
For
models
improve
appreciably
text states,
Pielou's
in mathematical
work
example, Watt
of communities
that stochasti
(1968) predicts
will not
and large populations
their performance.7
review
Cohen's
(1970) of
assess
"This book
should
liberate
those who
ecology
according
to its projection
along
an
A
axis
from
OF
SUCCESSION
to Davis
Princeton
IN
PARADIGMS
by
stantial
efforts,
accomplishments,
The liberation
is still
directions."8
25
ECOLOGY
them in the very sub
informing
in orthogonal
and opportunities
in the future, and will come only
in ecology.
of a materialist
revolution
Recent
the completion
differential
and Hobbie
papers by Tiwari
(1976a,b)
stochasticizing
a simple aquatic ecosystem
describe
may be in its
equations which
with
vanguard.
unease
vis-?-vis
and the zeal with which
of ecologists
physics
are misplaced.
science models
What
they seek deterministic
physical
as
to
is
music
the
the
view
noise
of
individuality
physicists
ecologist;
The
is their most
and
intrinsic,
striking,
the
of ecological
and
apparent
indeterminacy
inspiring characteristic,
their study a less valid pursuit. Mayr
does not make
systems
(1961)
the
of
entities
that
and
suggests
phenomana
biological
uniqueness
one of the major
differences
between
and the
constitutes
biology
and
populations
physical
difficult
There
it particularly
makes
and that this difference
sciences,
concerns.
to understand
for physical
scientists
biological
are three types of indeterminacy
at issue here. One is at least
as fundamental
in fact
is
and
communities
of
expression
to ecology
grounded
this sort of
as Heisenberg's
in the latter.
is to physics,
uncertainty
most
the
Perhaps
elegant
in ecology
is Lerner
and
indeterminacy
(1962) of the potential
"A molecular
accident
Dempster's
example
tion in a flour beetle:
effect
of a neuron,
in its wanderings
the stimulus
threshold
slightly
the path taken by a beetle
be modified,
and perhaps deflected
be
eaten."
Such molecular
indicate,
the outcome
of
affect
could, conceivably,
in consequence
of which
through the medium may
from an egg that otherwise would
as Lerner
and Dempster
indeterminacy,
to ecological
is intrinsic
of a random muta
systems
a
to be critical to
but unlikely
between
like competition
event,
particular
of two species. A second form of apparent
indeterminacy
populations
Poole
has only recently been addressed
Oster
1975,
1977),
(May 1974,
and takes
certain
equations.
the form of
non-linear,
Despite
'chaotic'
behavior
of populations
by
governed
of difference
systems
quite realistic,
apparently
are com
the fact that the underlying
equations
as
the resulting trajectories may be so complex
pletely deterministic,
to appear random, and Oster
(1975) suggests
that itmay
be impossible
26
DANIEL
SIMBERLOFF
true stochasticity,
data to distinguish
among
biological
experi
flows of a deterministic
mental
model. However,
error, and complex
similar situations
arise in meteorology
(May 1974) and even in that
with
billiard
table
ideal, the Newtonian
(Oster
quintessential
physical
can
so
to the status of second-rate
1975),
ecology
hardly be relegated
science
on
these
grounds.
is probably most foreign
third type of ecological
indeterminacy
cause
to the physical
sciences
and is the primary
of ecologists'
defensiveness.
This
is apparent
by the
indeterminacy
engendered
them
enormous
number of entities even in simple ecological
systems
The
selves
rather
than
Further,
systems.
are often
actions
by
these
the
form
entities
subtle. Whether
(1970)
Wangersky
deterministic
pletely
that
this
of
may
we
the
describing
and the
interacting,
believe with Mayr
complexity
ecological
description
and better instrumentation
ally, that better
within
Heisenberg's
equations
be
will
(1961)
the
inter
and
com
forever
preclude
or feel, more optimistic
and effort will bring us to
of ecological
description
limits of a perfect
1966), we must agree that we will not, in the near
(Holling
or insight to produce
as
information
future, have sufficient
equations
as
most
those
of
and
the
other
On
physicists
engineers.
predictive
able to predict weather
hand, neither are meteorologists
patterns with
systems
of money
and the amount
and manpower
precision,
to the study of single ecological
systems pales compared
to the effort involved
rocks. With
in, say, scooping
up a few moon
resources
more
been
successful
and
sufficient
have
notably
ecologists
remarkable
committed
are given by DeBach
in their predictions;
many
examples
In any event,
the nature of genetic
for biological
control.
the
evolution
of
and
fact
that
systems
constantly
living organisms
a
occurs
amount
in the outcome
insures
certain
of variability
of
precise
(1974)
events
is among
the
1966), and this variability
(Pimentel
ecological
most
the
of
Further,
aspects
interesting
ecological
phenomena.
as in Schaffer and Elson's
amount of variation
itself can be predicted,
as discussed
(1975). Finally,
study of salmon life history phenomena
is evolutionarily
early in this paper, a certain amount of variability
adaptive;
manifests
se
per
variability
consequently
of ecological
itself in the workings
is
selected
systems.
for,
In sum,
and
that
A
OF
SUCCESSION
PARADIGMS
IN ECOLOGY
27
because
of their complexity
systems
variability
ecological
typifies
less
than
the
worthwhile
does not render their study
study of physical
different mathematical
and
rather it suggests
systems;
approaches
to force biological
than attempting
"rather
success,
a
created
into
mold
economics,
by hydrodynamics,
phy
phenomena
instituted by
sics, or what have you" (Slobodkin
1975). The policy
extensive
in 1963 of not publishing
tabular data symbolizes
Ecology
for
criteria
such an attempt, coming in the midst of new
and
of populations
ideal characterizations
the stochastic
of
recognition
nature
interest
in logistic-based,
communities;
of ecology
demands
increasing
a re-evalu
tation.
I end
this
ecology
on
section
the status
of
the materialistic
in
revolution
the observation
with
that the first ecological
ideal, Clements'
is not dead, but rather transmogrified
into a belief that
is the proper course for ecology
of ecosystems
(Watt
superorganism,
holistic
study
1966b, Levins
and Levins
1968, Lane, Lauff,
1975, Johnson
1977,
as
Mclntosh
views
the
Odum
1977; cf.
1976).
(1964)
ecosystem
as
to
to ecology
molecular
the same relation
the cell does
bearing
a clearly superorganismic
Patten
(1975) sees the
biology,
conception.
Odum
as
a
unit
of coevolution,"
and argues
that
is a
linear
behavior:
ecosystems
good
"Nonlinearity
an
no
not
mathematical
one, and
ecosystem
property,
ecological
a relation that describes
until someone writes
is nonlinear
it
process
a
from
Aside
the
is as much
so."
fact
that
linearity
ecosystem
"holistic
evolve
mathematical
of
artifact
existence
toward
property
Patten's
as is nonlinearity,
the well-behavedness
an ecosystem
of defining
convention
is an
out
of
it is egregiously
ill-behaved; Holling
(1973) views the
as evidence
domains
of attraction
in the
for multiple
of the validity of the holistic, well
The true measure
when
same behavior
same
system.
behaved
ecosystem
it provides
is whether
insight into com
concept
et al.
and its record here is equivocal
(Mitchell
munity mechanisms,
et al. 1977). Well known
instances of abrupt fluctua
1976, Auerbach
are
(1969) gives
yet Preston
ignored,
one
claim that poor
that
might reasonably
sufficiently many examples
increases
nature. The widely
characterizes
behavior
cited, dramatic
and
in
the
Pacific
of Acanthaster
1973,
1974)
(Branham
Glynn
planci
tions
in single
populations
28
DANIEL
SIMBERLOFF
terebrans on Florida mangroves
Sphaeroma
(Rehm and Humm
1973)
are two more recent examples
of ill-mannered
ecosystems.
One may
ask why
so many
focus on ecosystems
has seduced
in
the face of its failure
to add substantially
to our
ecologists
of the workings
understanding
failure
is noted,
it is ignored
of
nature.
even
Indeed,
or
when
this
as Kuhn
away,
(1970)
explained
is typical for a paradigm before
it is finally overthrown
by a
revolution.
the predominant
failure of the U.S.
Admitting
International
the most massively
(I.B.P.),
Biological
Program
sup
one
effort
and
in a
conceived
ported American
ecological
wholly
suggests
scientific
vein, Odum
ecosystem
(1977), a leading ecosystem
that
it
is
not the paradigm which
is faulty, but
suggests
with which
the paradigm was used in organizing
consistency
holistic,
adherent,
rather the
research.
In short, the LB.P. effort was not holistic enough!
One suggestion
for the apparent paradigmatic
status of the ecosys
tem concept
in the face of conflicting
data is that it provides
support
for the notion of self-regulatory
inherent in unfettered
powers
capi
a
talism
For
if
of
1971).
(Leigh
community
organisms,
naturally
each to maximize
the representation
to a single organism
to be analogous
common
so ought a competitive
purpose,
selected
shown
unified
whole
metaphor
activities
was
which
benefits
that of a hidden
all. This
hand
of its own genes, can be
whose
parts all work to a
a
to produce
capitalism
is an old notion; Adam Smith's
the profit-maximizing
converting
into the good of the whole. That this should
be true for ecological
on both evolutionary
is questionable
systems
of observed
irre
(Levins
1974) and the grounds
grounds
ecological
were
described
true
Even
it
I
above.
for
gularity
suspect that
ecology,
it is not the primary attraction
of the ecosystem
but one
paradigm;
of
individuals
the strength with which
ought always to recognize
even an economic
our perception
one, structures
related
main
a basic
philosophy,
un
of apparently
that
the
argument
the most
phenomena.
Perhaps
convincing
attraction of holism is not as a subtle justification
of capitalism
is
credentials
long-standing,
impeccable Marxist
and Levins
1968, Lewontin
(e.g., Levins
1976).
Mclntosh
documents
the transformation
in the 1950's and
(1976)
a
1960's of American
into
the era of
ecology
big-money
operation,
that it has adherents
with
A
OF
SUCCESSION
IN
PARADIGMS
ECOLOGY
29
that the big money
is primarily
in
'Grant Swinger,' without
observing
the area of ecosystems.
the I.B.P. was followed
For example,
by the
of a new program,
in the National
creation
Studies,
Ecosystem
over
this
Science Foundation.
has
twice the annual
program
Currently
of
budget
the General
Ecology
awards
Consequently
proposals.
mon
in Ecosystem
Studies
Program,
in excess
and
fewer
than half
the
are com
of $100,000./yr.
in General
To the
Ecology.
of
determinant
academic
important
and unusual
that grant funding
is an
and economic
advancement,
well-being
Merton
1973), one might
reasonably
extent
a general
argue
1973,
goal (Storer
that the ecosystem
of
alone,
independent
on economic
is seductive
grounds
paradigm
or biological
either philosophical
considerations.
Yet another attraction
is that
of the ecosystem
it lends
itself
to
a vogue vocation
via systems
in
interpretation
analysis,
States for about two decades
(Mclntosh
1976). Indeed, the
has been systems analysis
(Patten
primary thrust of ecosystematists
into a
1959, 1971, Watt
1966a) and the glamor of turning ecology
cybernetic
the United
the terminology
and
of engineering
science,
replete with
a
must
itself
been
the
have
inducement
of
ecosys
physics,
powerful
tem approach,
with
the economic
fitting hand-in-glove
appeal. The
space-age
concurrent
the appeal
rise of computer
further augmented
technology
a systems
and though Mclntosh
analytic
study of ecosystems,
to have crested,
that the tide appears
nevertheless
(1976) observes
a powerful
this aspect
remains
force
in
of ecosystem
research
=
Odum's
cell:molecular
(1964) analogy,
today.
ecology
biology
may be rel?vent not only for the light it throws on
ecosystem:ecology,
of
the relationship
to metaphysical
of the ecosystem
concept
thought
as an
its intellectual
debt to the superorganism
but
also
concept,
to
the
of
the
of
achieve
desire
expression
ecologists
respectability,
and
even
in its heyday, when Odum wrote.
glamor, of molecular
biology
reason
I suggest
that the chief
for the persistence
of the
with
is
that
it
accords
Greek
The
ecosystem
paradigm
metaphysics.
But
the notion that 'everything affects everything
of holism,
else' (Watt 1966a), includes not only its tidiness but its determinism,
are included
in the system and linked to all
for if all components
attractiveness
others
by deterministic
equations,
then no exogenous,
random
input is
30
DANIEL
The myth
and
consciousness,
of
possible.9
moner's
SIMBERLOFF
the balance-of-nature
(1971); Colwell
paradigm and the balance-of-nature.
thing'
by
in the popular
Com
in Barry
systems
ecological
of all ecology
into 'You can't change
just one
(1970) also notes the identity of the ecosystem
condensation
That
attracts
the
persists
form
takes
an idea so readily accepted
as well
adherents
is not
lay public
professional
when
the idea has 2000-year
old roots. Even
surprising, particularly
a founder of stochastic
Albert Einstein,
viewed
mechanics,
quantum
as an instrument
for dealing with atomic systems, not as a true
"I am absolutely
of the universe:
convinced
that one
representation
will eventually
arrive at a theory in which
the objects
connected
by
laws are not probabilities,
but conceived
facts...
."(Born 1949). That
it only
to a 'dice-playing
God' was
irrational, however,
objection
"I
cannot
he readily admitted:
for my
provide
logical arguments
but can only call on my little finger as a witness,
which
conviction,
cannot
to be respected
claim any authority
outside my own skin"
Einstein's
(Born
1949). Small
influence ecologists!
wonder
that Greek
metaphysics
to
continues
there is something
disturb
Certainly
profoundly
random elements play a large role. Just as
ing about a nature in which
to Darwinian
much of the opposition
evolution
by natural
powered
so the
the
to
selection was engendered
role
chance,
by
large
assigned
idea of an unbalanced,
spires distrust.
The
ecosystem
stochastically
paradigm
purports
driven
to have
in
natural
community
corrected
the superor
that succession
ganism's
shortcomings,
primarily by explicitly
noting
on certain
need not always
lead to the same climax and by focussing
individual
characteristics
But its most
of ecosystems.
fundamental
are determinism
in a high-level
in
and interest
ideal entity;
it is squarely
in the camp of the essentialists.
The success of the
in other disciplines,
revolution
materialist
evolution
and
particularly
for ecology,
but Greek metaphysics
will not
augurs well
genetics,
features
these
vanish
easily.
A
OF
SUCCESSION
IN ECOLOGY
PARADIGMS
31
ACKNOWLEDGMENTS
R. Strong,
to Donald
Jr. for numerous
I am grateful
sug
specific
as well as for a long
on
this
of
all aspects
manuscript,
gestions
interest
in the history
and philosophy
of
freely discussed
standing,
F. Connor,
Frances
C. James, Eugene F. Kaelin,
science.
Edward
B. Mertz,
discussion
David
both
Alexandra
and several
Department
of Biological
Florida State University,
and Edward
Radbil,
key references.
O. Wilson
provided
Science
Tallahassee
NOTES
1
Malthus
stated his syllogism
thus:
to the existence
is necessary
"First, That food
the sexes
between
and will remain
is necessary
"Assuming
then, my
indefinitely
greater
of man.
That
Second,
in the present
state.
the passion
as granted,
I say, that the power
of population
postulata
in the earth to provide
than the power
subsistence
for man.
is
check
on population
a strong
"This
and constantly
the
from
implies
operating
This difficulty must
of subsistence.
fall somewhere
and must
difficulty
necessarily
be severely
felt by a large portion
of mankind."
2
Seton and Goss, Knight,
Haacke
Verlot,
Gaertner,
Vilmorin,
Naudin,
(Olby 1966)
3
Wiener
the related
idea that the increasing
of the role of
(1949) develops
recognition
use of statistical
in a number of disciplines,
the consequent
approaches
plus
on evolutionism
in The Origin
and the individual
and Mill's
On Liberty
in 1859!), led directly
to twentieth
(also published
century American
pragmatism.
4
It is ironic but appropriate
that the integrity of Mendel's
into doubt
reports be thrown
chance
and
the emphasis
to the Neo
foremost
and one contributor
statistician
century's
as a group,
Fisher
too
that Mendel's
ratios were,
(1936) noted
ones
even
to the hypothetical
to be caused
for the deviations
alone,
by chance
are as Mendel
can
that the underlying
Mendel
probabilities
suggested.
Perhaps
by R.A.
Darwinian
close
this
Fisher,
synthesis.
given
be excused
this disturbing
on
anomaly
the grounds
of
the novelty
of
statistics
in his
field!
5
The power of the paradigm
individual
species. Margaret
changes,
has
been
frustrated
of the pioneer
vegetation
for a non-existent
climax
(Colinvaux
1973).
was
such
B. Davis,
by
days was
that they
as to preclude
gathering
in her long-term
studies
this
lacuna:
"We
do
not
of data
of
oriented
long-term
what
know
were
all the ecologists
to record what was actually
toward
vegetation
the virgin
like because
so busy
forgot
growing
looking
there"
DANIEL
32
6
adumbrated
Gleason
this
'fortuitous
immigration'
and
laws of probability
SIMBERLOFF
as he did so many
he wrote
when
others,
discovery,
and plants'
"in accordance
with
distributed
being
was
in its infancy
in biology,
(1929). But statistics
of
the
(1926)
chance"
as
on the spread of
A literature
and Gleason's
above,
insight was
imprecise.
was
as a stochastic
at the time of Park's
also extant
work
process
(e.g.,
epidemics
a formal
had published
statistical
1956,
1957, Bartlett
1960), and Kendall
Bailey
treatment
into ecological
of population
(1949), but this work was not integrated
growth
described
thought.
7
He quotes
as a stochastic
(1957): "Considering
population
change
those obtained
with
different
from
deter
appreciably
of critical
such as total
for the prediction
models...,
phenomena
especially
the group.
It is our
in particular
of
extinction
however,
belief,
that,
except
of effort both in deriving
the stochastic
and in
instances...
,the multiplication
equations
Beverton
.often
process..
ministic
and Holt
results
gives
not have been
them would
the standard
of accuracy
of our
justified when
computing
we are dealing,
of the biotic
and the order of
the complexity
system with which
are all taken into account."
of the expected
discrepancies
magnitude
8
was widely
as a center
of Robert
viewed
the institution
of
Princeton,
MacArthur,
data,
at Davis
K.E.F.
while
the University
of California
housed
mathematical
ecology,
Watt's
group.
systems
analysis
leading ecosystem
9
A witty, widely
but unpublished
'On the Tendencies
of Motion'
circulated
manuscript
lambastes
the
'Isadore Nabi')
and R. Levins
the pseudonym
(under
by R. Lewontin
on grounds
of both messiness
and tunnel vision. A bogus
stochastic
experi
approach
on motion
ment
and
the
variance
with
of apples,
cannonballs,etc.
of the data to multiple
subjection
in a series of nonsensical
results
massive
regression,
conclusions
amounts
of data is described,
and analysis
of
analysis,
to Newton's
without
leading
factor
laws.
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