Spatial Pattern and Ecological Analysis PDF

Spatial Pattern and Ecological Analysis
Author(s): Pierre Legendre and Marie-Jose Fortin

Source: Vegetatio, Vol. 80, No. 2 (Jun. 15, 1989), pp. 107-138
Published by: Springer
Stable URL: http://www.jstor.org/stable/20038425 .
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80: 107-138,
1989.
Vegetatio
1989 Kluwer Academic
?
Publishers.
Spatial
pattern
Printed
107
in Belgium.
and ecological
analysis
Pierre Legendre1
& Marie-Jos?e
Fortin2
1
sciences
de
C.P.
D?partement
biologiques, Universit? de Montr?al,
2
H3C
Canada
and
3J7;
Evolution,
Qu?bec,
Department
of Ecology
Brook, NY
6128, Succursale
State University
A, Montr?al,
of New
York, Stony
USA
11794-5245,
17.1.1989
Accepted
Keywords:
Ecological
Vegetation
map
theory, Mantel
test, Mapping,
Model,
analysis,
Spatial
Spatial
autocorrelation,
Abstract
The
spatial heterogeneity
for instance the theories
of populations
of succession,
adaptation,
interactions,
review how
competition,
predator-prey
and so on. This paper will
a central
and communities
parasitism,
the spatial
role inmany ecological

theories,
of species diversity, community
stability,
and other natural catastrophes,
epidemics
ergoclines,
structure of biological
and communities
populations
plays
maintenance
can be studied. We
first demonstrate
that many of the basic statistical methods
used in ecological
studies
are impaired by autocorrelated
if not all environmental
data. Most
data fall in this category. We will look
tests in the presence
valid statistical
of spatial autocorrelation.
Methods
briefly at ways of performing
now available for analysing
the spatial structure of biological
are
and
illustrated
described,
populations
data.
These
to
include
various
test
methods
for
the
of
in
by vegetation
presence
spatial autocorrelation
the data:
methods
and two-dimensional
and two
(all-directional
spatial correlograms,
and the multivariate
test and Mantel
Mantel
other
spectral analysis),
correlogram;
descrip
tive methods
of spatial structure: the univariate
and the multivariate
of clustering
methods
variogram,
with spatial contiguity
the partial Mantel
here as a way of studying causal
test, presented
constraint;
models
that include space as an explanatory
and
various
methods
for mapping
variable;
finally,
ecological
variables
and producing
either univariate maps
trend surface analysis, kriging) or maps
(interpolation,
univariate
dimensional
of truly multivariate
data (produced by constrained
clustering).
terms of the ecological questions
they allow to resolve. Reference
Introduction
In nature,
from
living beings
nor at random.
are distributed
role
neither
Rather,
they are aggre
or they form gradients
or other
structures.
of spatial heterogeneity
importance
its central role in ecological
theories
practical
table
ismade
shows
the methods
to available
computer
classified
in
programs.
several ecological
theories and models
of an ecosystem
that elements
that are
to one another in space or in time are more
Actually,
assume
uniformly
gated in patches,
kinds of spatial
The
in population
sampling
close
likely
to be
Such
process.
of epidemics
comes
theories
and
adaptations,
its
theory.
influenced
is the case,
or other
of competition,
maintenance
parasitism,
population
by the same
for instance,
generating
for models
for the
catastrophes,
evolution and
succession,
of
species
genetics,

diversity,
population
108
interactions,
growth,
predator-prey
assume
theories
behaviour.
Other
and
social
dis
that
zones
are
homogeneous
(succes
important for the structure of ecosystems
Allen
sion,
species-environment
relationships:
between
continuities
1911 \Allen & Starr 1982; Legendre

etal.
or
for ecosystem
1985)
dynamics
(ergoclines:
the impor
Lege?dre & Demers
1985). Moreover,
to eco
tant contribution
of spatial heterogeneity
etal.
1958; May
1984).
seems well
stability
logical
This
established
1974; Hassell & May

shows
clearly
that
(Huffaker
much
has been
the spatial variability

century
quantitative
a uniform
the
spatial
or
up to now about
of communities.
Most
19th
learned
studies
ecological
distribution
were
of living organ
assuming
area (Darwin
isms in their geographic
distribution
several
and
1881; Hensen
1884),
ecological
still assume, for simplicity,
that biological
are dis
variables
and
their
organisms
controlling
tributed in nature in a random or a uniform way
models
(e.g.,
simple
some models
models
etal.
(Legendre
1985). The spatial heterogeneity
of the physical
a
environment
thus generates
as
in
communities
of living beings,
well
diversity
as in the biological
and ecological
that
processes
can be observed
at various points
in space.
This
of
dynamics,
population
or
of forest or fisheries exploitation,
This assumption
is
productivity).
en
remote
from
since
the
reality
actually quite
is spatially structured by various energy
vironment
inputs, resulting in patchy structures or gradients.
Table
fied
1. Methods
1) Objective:
lation.
in
for instance
environments
(water,
and phytoplank
by aquatic macrophytes
ton, and air, inhabited
by terrestrial
plants),
of
thermal, mechanical,
energy
gravita
inputs
fluid
and even
radioactive
tional,
found, besides
light energy which
the
of most
chains;
trophic
heterogeneity
and advection
the formation
are
origins
lies at the basis
spatio-temporal
of energy inputs induces convection
in the fluid, leading to
movements
of spatial or temporal discontinui
between
ties (interfaces)
relatively homogeneous
zones.
and discontinuities
In soils, heterogeneity
From
are the result of g?omorphologie
processes.
the
there, then,
structuring of the
spatio-temporal
induces a similar organi
environment
physical
zation of living beings and of biological
processes,
as
as
well
Strong biological
temporally.
spatially
in interface zones
activity takes place particularly
surface
and
questions
statistical
Method
distance
ble
Method
2) Objective:
Method
Method
for
or
dimensional
single
c;
Geary's
and
autocorre
or
matrix)
Mantel
(or multi
space (geographical
a varia
test between
for multivariate
correlogram,
data.
structure.
of the spatial
Description
1 : Correlograms
(see above),
variograms.
2: Clustering
with
and ordination
spatial
poral constraint.
Test
3) Objective:
predictor.
Method:
causal
Partial
models
Mantel
A,
matrices,
test,
B
shape.
variable,
using
two-dimensional
a variable
and
matrix);
a model.
3 :Mantel
autocorre
spatial
is significant
spatial
the kind of pattern,
analysis.
test between
spectral
2: Mantel
of
auto
significant
spatial
to use parametric
in order
in the data,
tests.
1 : Correlograms
/
Moran's
classi
analysis,
is no
there
that
that there
1.2) Establish
lation and determine
Method
pattern
objectives.
for the presence
Testing
1.1) Establish
correlation
habited
chemical
aspects.
methodological
for spatial
by ecological
of ecosystem
In
includes
paper
1974; Levin
is an important
temporal structure of ecosystems
element of most
theories.
ecological
Not
(Legendre & Demers

1985). Within homogeneous
an aggre
biotic
often produce
zones,
processes
of organisms,
various
gation
following
spatio
and these can be measured
scales,
temporal
that
include
using
three
or tem
as
space
dissimilarity
et C.
4) Objectives: Estimation (interpolation) and mapping.

Method
Method
for a single variable:

trend
1: Interpolated
map
that provides
also the regres
surface analysis,
other
methods.
sion residuals;
interpolation
2:
into
taking
structure
Interpolation
autocorrelation
gram):
grams
function
that may
estimations,
sampling
3: Multidimensional
ordination
with
help
decide
spatial
(vario
for a single variable;

kriging map,
also
the
standard
deviations
give
add
Method
account
pro
of the
where
to
locations.
and
clustering
mapping:
constraint
(see above).
spatial

109
spatial autocorrelation
its influence on classical
statistical
is, and discuss
methods.
Then we shall describe
the univariate
methods
and multivariate
that we have had ex
anywhere near the first, this cannot be seen as a

random draw of elements
; the reason is that the
value of the variable
in (A) is now
observed
so that if the existence
and the shape of
known,
perience with for the analysis of the spatial struc

ture of ecological
communities
(list not neces
and illustrate this description
sarily exhaustive),
with actual plant community
data. Finally, recent
are also known,

one can
structure
the spatial
foresee approximately
the value of the variable in
the observation
is made.
This
(B), even before
in spatial analysis will be presented,

developments
to test simple interrelation
that make
it possible
are not
We
first what
shall define
models
varia
space as an explanatory
in this paper are also
described
that include
ble. The methods
to geology,
the
applicable
pedology,
geography,
earth sciences, or to the study of spatial aspects
of the genetic heterogeneity
of populations.
These
the study of observa
have in common
tions positioned
in geographic
space; such obser
are related to one another by their geo
vations
sciences
which
graphic distances,
that space. This
are the basic
relations
in
a
is organized
around
paper
series of questions,
of increasing refinement,
that
can ask when they suspect their data to
ecologists
be
structured
nomenon
by
(Table
some
underlying
spatial
phe
1).
shows
at neighbouring
points
from one another. Random
that observations
independent
or systematic
sampling designs
cated as a way of preventing
try to show that the methods

are not always
statistics
adequate
will
first
classical
of
to
study
space-structured
ecological
phenomena.
This will justify the use of other methods
(below)
when
the very
nature
of
the
spatial
structure
is of interest.
(autocorrelation)
In classical
one
inferential
statistical
analysis,
of the most fundamental
in
assumptions
hypoth
esis testing is the independence
of the observa
tions (objects, plots, cases, elements).
The very
existence
implies
of a spatial structure
that this fundamental
because
satisfied,
located at a given
influence
some
any
in the sample
assumption
space
is not
phenomenon
sampling point may have an
on other points located close by, or even
distance
ecological
away. The spatial structures we

find in nature are, most of the time, gradients
or
In
one
such
a
when
draws
first
cases,
patches.
sample (A), and then another sample (B) located
the
is suitable
to
systematic
sample of points
estimation
of the parameters,
achieve unbiased
since each point a priori has the same probability
of being
in the sample; we know of
included
course that the variance,
and consequently
also
the standard error of the mean, will be larger if the
cation
We
of
violations
or
now
structure
and spatial
advo
of inde
assumption
of errors. It is adequate, of course, when
pendence
one is trying for instance to estimate the parame
ters of a local population.
In such a case, a random
against
remains
statistics
been
this possibility
of
among observations
1977;
dependence
(Cochran
Green
then
1979; Scherrer
1982). This was
believed to be a necessary
and sufficient safeguard
is
distribution
Classical
have
unbiased.
that despite
of samples
but
their
estimation
patchy,
On the other hand, we know
the random or systematic
allo
space, observations
through
if
may retain some degree of spatial dependence
the average distance
between
is
smaller
samples
than the zone of spatial influence of the underlying
in the case of large-scale
ecological phenomenon;
spatial gradients, no sampling point is far enough
to lie outside
this zone of spatial influence.
A
is
to be autocorrelated
(or
to
it
is
the
regionalized)
possible
predict
values of this variable at some points of space [or
time], from the known values at other sampling
variable
said
when
points, whose
also known.
spatial
[or temporal]
[or temporal]
function,
by a mathematical
a spatial autocorrelo
structure function',
Spatial
tion can be described
called
gram and a semi-variogram

of such functions.
Autocorrelation
classes
are
positions
autocorrela
are examples
is not the same for all distance
sampling points (Table 2). It can

or negative. Most
often in ecology,
between
be positive
(below)

110
Table
2.
Examples
(non-exhaustive
Sign of spatial
autocorrelation
of spatial
list). Modified
short
often:
any
at short
(if the
step
sampling
small enough).
regularly
is
(e.g.,
spaced
plants);
sampling
step too wide.
or other
(e.g.,
furrows)
repeating
themselves
trough
positive
at short distance
is that positive
autocorrelation
distorts statistical tests such as correlation,
regres
and that this distor
sion, or analysis of variance,
tion is on the 'liberal' side (Bivand 1980; Cliff &

that when positive
1981); this means
spatial
in the small distance
is present
autocorrelation
tests determine
too
classical
statistical
classes,
Ord
space,
Spatial
(if also
is at least partially known

in
variable
new
but a
observation
each
contributes
observed
advance,
fraction of a degree of freedom. The size of this
so that
fraction cannot be determined,
however,
the proper
do not know
reference
statisticians
for the test. All we know for certain
distribution
for
Aggregates
is structures
that
phenomenon
Avoidance
ecology
(1979).
large
distances
distances
contagious
distance
Sokal
autocorrelation
Significant
-
+ Very
in
autocorrelation
from
gradient
significantly
at short
often
that correlations,
coefficients,
regression
differences
among groups are significant, when
are not.
fact
distance).
they
include randomization
or
in
to these
Solutions
tests,
problems
i-test
the corrected
proposed by Cliff & Ord (1981), the analysis of

autocorrelation
variable
takes
is positive
that the
(which means
similar values) for short distances
auto
In gradients,
this positive
is coupled with
short distances
as
for long distances,
autocorrelation
negative
points located far apart take very different values.
an aggregated
structure
at
Similarly,
recurring
among points.
at
correlation
for
autocorrelation
positive
to
the
between
gap
patch
corresponding
centers. Negative
for short dis
autocorrelation
tances can reflect either an avoidance
phenome
intervals
will
show
distances
non
(such as found
and solitary
among regularly spaced plants

animals), or the fact that the sampling
step (interval) is too large compared

so that any given patch does not
than one
to patch size,
contain more
the next
sample,
sample falling in the
interval between patches. Notice
finally that if no
a given scale of
at
is
found
autocorrelation
spatial
it
(i.e., a given intensity of sampling),
perception
mean
not
not
be
does
that autocorrelation
may
found at some other scale.
In classical
tests of hypotheses,
statisticians
count one degree of freedom for each independent
them to choose
which
the
allows
observation,
statistical
distribution
appropriate
for
testing.
This is why it is important to take the lack of

into account
independence
of autocorrelation)
presence
test of statistical hypothesis.
(that results from the

a
when performing
Since
the value of the
in the presence
of spatial autocorrelation
et al. (submitted),
etc. See
by Legendre
a
of
for
Edgington
(1987)
general presentation
see
also
&
randomization
tests;
Upton
Fingleton
as the other references
as well
in the
(1985)
variance
developed
to spatial analysis.
present paper, for applications
is
Another
the spatial structure
way out, when
a
to
extract
is
the
linear
gradient),
simple (e.g.,
first and conduct
the analysis
(see: trend surface
analysis,
after verifying that no spatial autocorrela
spatial component
on the residuals
below),
tion remains
The
in the data.
situation
classical
described
multivariate
been used
extensively
above
data
also
analysis,
by ecologists
applies to
which
has
for more
than
two decades (Orl?ci 1978;Gauch 1982; Legendre

& Legendre
1983a, 1984a; Pielou
1984). The spa
of the data points
tial and temporal coordinates
are usually neglected
during the search for eco
out
which
aims at bringing
structures,
observations.
and relations
among
processes
time
of the space and/or
the importance
Given
as
in
the
in
theory,
component
argued
ecological
logical
Introduction,
study these
and
used
ecologists
important
clustering methods
are now
relationships.
in particular
to detect
and analyse
spatial
(e.g., Andersson
analysis
vegetation
though these techniques
to
beginning
Ordination
are often
structures
in
even
1988),
were not designed
specifi

Ill
are also being
for this purpose. Methods
that take spatial or temporal relation
developed
data
account
into
during multivariate
ships
con
of
the
methods
include
These
analysis.
cally
strained
methods
below,
clustering presented
ordination
of constrained
as the
as well
by
developed
Lee (1981),Wartenberg (1985a,b) and ter Braak

(1986, 1987) where one may use the geographical
of the data points as constraints.
coordinates
into
is divided by geographers
Spatial analysis
the distribu
point pattern analysis, which concerns
of
tion
(discontinuous
phe
points
physical
in space - for instance, individual plants
nomena)
a topological
and animals ; line pattern analysis,
of connections
to the study of networks
approach
points; and surface pattern analysis
of
phenomena,
study
spatially continuous
are attached
one or several variables
among
observation
to represent
Point pattern
and to
is any significant
spatial autocorrelation,
its type
establish
(gradient,
unambiguously
etc.). This can be done for two diametri
patches,
to
either (1) one wishes
purposes:
cally opposed
show
spatial autocorrelation,
to perform parametric
statisti
or
on
the
tests;
contrary one
(2)
that there is a spatial structure in
to study itmore thoroughly.

autocorrelation
spatial
study
order
Besides
of data points
the geographic
distribution
or not, and to describe
the type of
for inferring
this can then be used
pattern;
that might
have led to the observed
processes
structure. Graphs
of interconnections
among
is no
there
one wants
cal hypothesis
hopes to show
where
points, and each point is considered

of space.
its surrounding
portion
to
establish
is
intended
analysis
that
because
the
structure
Let us first study one variable at a time. If the map

and mapping,
of a variable (see Estimation
below)
eco
a
structure
is present,
that
suggests
spatial
there
logists will want to test statistically whether
for the
to
of a spatial
for the presence
Testing
In either case,
is conducted.
testing for the presence of a spatial struc

as well as
the
various types of correlograms,
ture,
a
periodograms,
provide
description of the spatial
structure, as will be seen.
whether
is random
that have been introduced by point pattern

analysis, are now widely used also in surface pat
tern analysis
they serve for in
(below), where
points,
stance
as basic
constrained
analysis,
of
for
relationships
autocorrelation
clustering,
spatial
etc. The methods
of point
pattern
in
and
the quadrat-density
particular
networks
analysis,
and the nearest-neighbour
widely used in vegetation
have been
methods,
science
(e.g., Galiano
1982;Carpenter & Chaney 1983) and need not be

have
any further here. These methods
expounded
been summarized by a number of authors, includ
ing Pielou (1977), Getis & Boots (1978), Cic?ri

etal. (1977) and Ripley (1981, 1987). The expos?
that follows will
for
surface
presently
then concentrate
pattern
analysis,
with.
experimenting
on the methods
that ecologists
are
autocorrelation
Spatial
coefficients
In the case of quantitative

variables,
spatial auto
can be measured
/
either
Moran's
by
correlation
(1950) orGeary's c (1954) spatial autocorrelation

coefficients.
Formulas
1. Moran's
Pearson's
are
in App.
presented
behaves
like
mainly
coefficient
since its numera
formula
correlation
of a sum of cross-products
of centered
a
is
covariance
in
(which
term), comparing
turn the values found at all pairs of points in the
tor consists
values
class. This coefficient

is sensitive
given distance
to extreme values,
or a
just like a covariance
Pearson's
correlation
coefficient. On the contrary,
c coefficient
is a distance-type
Geary's
function,
sums the squared differences
found at the various
pairs of
since the numerator

between
points
The
values
being compared.
statistical
significance
can be
tested,
that
and
confidence
of these coefficients
intervals can be
the distance
classes
highlight
or negative
autocor
significant positive
relation, as we shall see in the following examples.
More
detailed descriptions
of the ways of com
computed,
showing
puting
and testing
these coefficients

can be found
112
in Sokal & Oden (1978), Cliff & Ord (1981) or
& Legendre
Legendre
coefficients
also exist
Autocorrelation
(1984a).
for qualitative
(nominal)
variables (Cliff& Ord 1981); they have been used

to analyse
for instance
spatial patterns
of sexes
in
plants (Sakai & Oden 1983; Sokal & Thomson

types of
1987). Special
coefficients
have
been
specific
problems
& Gates
spatial
autocorrelation
to answer
developed
1983; Estabrook
(e.g., Galiano
the autocorrelation
puting
series analysis.
coefficients
of
time
1984).
A correlogram
is a graph where autocorrelation
are
in ordinate,
values
plotted
against distances
com
localities
(d) among
(in abscissa). When
one must be able to
puting a spatial correlogram,
assume
that a single 'dominant'
spatial structure
exists over the whole area under study, or in other
structure is the
that the main
words,
large-scale
same everywhere.
must actually
This assumption
to
be made for any structure function one wishes
other well-known
functions,
compute;
to characterize
include
spatial patterns,
gram (below), Goodall's
also used
the vario
(1974) paired-quadrat
the two-dimensional
correlo
function,
the
multivariate
and
gram
(below),
periodogram
Mantel
(below), and Ibanez' (1981)
correlogram
variance
auto-D2
already familiar with the use of cor

in
time series analysis will be reassured
relograms
to know that whenever
the problem
is reduced to
one physical dimension
or
a physical
(time,
only
or
a
instead
of
bitransect)
polydimensional
the coefficients
for different dis
space, calculating
tance classes
turns out to be equivalent
to com
Readers
All-directional
a single correlogram
over all
is computed
of the area under
one
investigation,
must make
the further assumption
that the phe
nomenon
is isotropic, which means
that the auto
When
directions
correlation
considered.
structure
functions
at a time;
direction
with two-dimensional
sional
which
by
is the
same whatever
the
In anisotropic
situations,
can be computed
in one
this
is the case
for instance
two-dimen
correlograms,
and
all of
analysis,
variograms,
spectral
are presented
Example
the result of a test of signifi
with
each autocorrelation
function
direction
function.
In correlograms,
cance
is associated
correlogram
below.
1 - Correlograms
at their shape,
are analysed mostly

since characteristic
of this test is that

the null hypothesis
coefficient;
is not significantly
different
from
the coefficient
zero. Before
value in
each significant
examining
looking
with types of spatial struc
shapes are associated
the
tures; determining
spatial structure can pro
about the underlying
vide information
generating
a number of
Sokal (1979) has generated
process.
the correlogram,
however, we must first perform
a global test, taking into account
the fact that
several tests (v) are done at the same time, for a
and published
the pictures of the
so
We
also done
have
resulting
correlograms.
a
structures
of
for
artificial-data
here,
variety
level a. The global test

significance
the correlogram
whether
checking
in ecology
encountered
similar to those commonly
a
of 9
surface made
(Fig. 1). Fig. la illustrates
at least one value which

is significant at
to the
the a' = a/v significance
level, according
tests
of correcting for multiple
Bonferroni method
fol
100 points were
sampled
bumps.
x
a
10
10
of
varia
The
regular grid
points.
lowing
ble 'height' was noted at each point and a correlo
gram of these values was computed,
taking into
given overall
is made
by
contains
(Cooper 1968; Miller

analogy
Q-test
1977; Oden
in time series analysis
1984). The
is the Portmanteau
(Box & Jenkins 1970). Simulations in
1984 paper show that the power of Oden's

for spatial series of
is an extension
Q-test, which
the Portmanteau
test, is not appreciably
greater
Oden's
than
which
of the Bonferroni
the power
procedure,
a lot simpler.
is computationally
spatial patterns,
bi-normal
of the sampled
the geographic
position
The
(Fig. lb) is globally
sig
correlogram
points.
at the a = 5% level since several
nificant
indi
are significant
at the Bonferroni
vidual values
= 0.00417.
corrected
level a' = 0.05/12
Examin
ing the individual
significant values, can we find
account
the structure's
main
elements

from
the c?rrelo
113
Correlogram
-
9 fat bumps
neighbouring points
- distance between
successive peaks
distance to the
I next peak
distance between peak and trough

7
8
9
3
A
6
10
5
Distance
Number
of point pairs in each distance class
Correlogram
9 thin bumps
Correlogram
Random
Correlogram
Single
numbers
Correlogram
thin bump
Correlogram
of artificial
correlograms
of distances
black
of
Single fat bump
Narrow wave
Correlogram
Wide
wave
Distance classes
the structure
text), (a) depicts

analysed
by the correlogram
in each distance
in this figure.
class, for all the correlograms
at the a = 5% level, before
the Bonferroni
values
represent
significant
applying
are non-significant
white
the correlograms;
values.
squares
(between
squares
significance
structures
Sharp step
Distance classes
Distance classes
Distance classes
Fig. 1. All-directional
spatial
in (b). (c) displays
the number
In the correlograms
(b, d-k),
to test the overall
correction
Distance classes
Distance classes
Distance classes
Correlogram
Gradient
Distance classes
Distance classes
Correlogram
classes
pairs
(see
of points)

11
12
114
since the alternation
of positive
gram? Indeed,
an
is precisely
of
and negative values
indication
encountered
(Table 2). The first value of spatial

patchiness
class 1), corresponding
autocorrelation
(distance
to pairs of neighbouring
points on the sampling
this means
and significant;
that
grid, is positive
the patch size is larger than the distance between
2 neighbouring
points. The next significant posi
at distance
is found
tive value
between
actual
distance
between
significant
and
peaks
Negative
between
found
values,
here to the
Notice
that
they could
if the bumps
unevenly
a
correlogram with
produce
significant
was
correlogram
the
classes,
equal distance
cannot be
coefficients
are based
upon
values
classes,
afterwards.
constructed
with
last autocorrelation
interpreted,
too few pairs of
histogram,
Fig. lc).
The other artificial
because
they
localities
(see
in Fig. 1
analysed
a 10 x 10 regular grid of
do fall in
an
has
in-between
5% level
that the 'zone of influence'
bump spreads
in (b) because
overall
into more
distance
significance.
of this single
classes
than
the phenomenon
here is not
limited by the rise of adjacent bumps.
Single fat bump (Fig. lh): a single bi-normal
curve occupying
the whole
surface.
sampling
The correlogram
has an overall 5% level signifi
cance. The 'zone of influence' of this very large
is not much
bump
-
larger on
the correlogram
than for the single thin bump (g).

drawn from a normal
numbers,
were
and used as the
distribution,
generated
on the same regular
to be analysed
variable
100 random
geographic grid of 100 points (Fig. li). None of

the individual
values
level of significance.
- Narrow
wave
(Fig.
so
between
crests,
across the sampling
are significant
at the 5%
lj): there are 4 steps

that there are 2.5 waves
has
(Fig. Id). The correlogram
level significance
(Bonferroni
surface. The correlogram

The distance
5% level significance.
crests of the wave show up
between
successive
in the significant value at d = 4, just as in (b).
- Wide wave
(Fig. Ik): a single wave across the
2 flat surfaces
(Fig. le).
step between
Sharp
has an overall 5% level signifi
The correlogram
cance. Comparing
with Fig. Id shows that cor
between
cannot distinguish
analysis
relogram
has overall
sampling surface. The correlogram
is the
The correlogram
5% level significance.
same as for the single fat bump (h). This shows
cannot be dis
that bumps,
holes and waves
are neces
;
maps
tinguished using correlograms
were
also
sampled
They are:
points.
- Linear
gradient
an overall
5%
structures
using
correction).
real data presenting

-
spaced,
the same
of 9 thin
the bumps.
thin
Single
bumps
(Fig. lg), about the same
size as one of the bumps in Fig. la. The correlo
gram
Notice
12
significant
but with no other
into
is 3.4 units).
the distance
were
this
the first of these

troughs;
at distance
class 2, corresponds
radius of the basis of the bumps.
structure
Since
neighbours
values
give
the flat area
succes
distances
class 4
includes
classes,
3.18 and 4.24, the unit being the distance
2 neighbouring
points of the grid; the
distance
between
points
a geometric
structure
unambiguously
most
since
of
the data points
bumps,
this one
class 4:
distance
between
gives the approximate
are
sive peaks. (Since the values
grouped
here
could
have
been
found
12 tests of a random
structure, for an
level a = 5%. 100 sampling
significance
are probably not sufficient
to bring out
among
overall
a sharp step and a gradient
respectively.
9 thin bumps (Fig. If) ;each is narrower than in
Fig. la. Even though 2 of the autocorrelation
are significant
at the a = 5% level,
coefficients
of the
since none
is not,
the correlogram
at the Bonferroni
is significant
coefficients
2
In other words,
level a' = 0.00417.
corrected
as
as
extreme
those
coefficients
autocorrelation
has overall
sary.
Ecologists
hypotheses
processes
nomenon
shape
are often capable

of formulating
or
as to the underlying mechanisms
the spatial phe
that may determine
the
study; they can then deduct
if these
spatial structure will display
are true. It is a simple matter
then to
under
the
hypotheses
construct
an artificial model-surface
correspond
as we have done in Fig.
ing to these hypotheses,

1,
115
to analyse
that surface with a correlogram.
of the difference
a test of significance
Although
is not easy to construct,
2 correlograms
between
and
of the values in
of the non-independence
at
the 2 correlo
each correlogram,
simply looking
- the one obtained from the real
data, and
grams
- suffices in
that from the model data
many cases
to find support for, or to reject the correspond
because
ence of the model-data

Material:
to the real data.
- These
data
Vegetation
a multidisciplinary
during
ecological
of the Munici
gathered
study of the terrestrial
palit? R?gionale
et al.
(Bouchard
ecosystem
de Comt? du Haut-Saint-Laurent
1985). An area of approximately

in a sector a few km north
sampled,
0.5 km2 was

of
the Canada-USA
o
o
o
o
o
o
o
o
o
o
o
o
"d o.io
o
'
700
200
400
500
cally
2.
Position
sampled
of
the 200
in Herdman
1983. From Fortin (1985).
vegetation
(Qu?bec),
quadrats,
the
during
systemati
summer
of
TV
/
i
?i/
1 2
iV i?i?i?iiji
Distance
10 11 12 13 14 15 16 17 18 19 20
classes
of
Fig. 3. All-directional
correlogram
spatial
distance
Abscissa:
densities
(Tsuga canadensis).
is 57 m. Ordinate:
of each distance
class
width
statistics.
Symbols
as
in Fig.
i?h-?i?i?i
m -*/
N^x0""^^
600
m ?tres
Fig.
-0.05
-0.10 1
0
300
two
second
/ \
o.oo ? ?i??t?i?
'
'
between
I 005 \
o
o
o
to
from 0 to 57 m), corresponding
on the sampling grid,
this means
that the
0.15
o
o
0.20j \
o
o
o
o
o
o
0.25
o
o
o
o
o
o
from this correlogram?

The first value of
class 1, includ
autocorrelation
(distance
o
o
to examining
in
significant
proceed
values: can we find the structure's main
o
o
then
o
o
o
o
o
o
2 - The correlogram
in Fig. 3 de
Example
scribes the spatial autocorrelation
(Moran's /) of
It is globally
the hemlock,
Tsuga canadensis.
sig
=
nificant
test, a
(Bonferroni-corrected
5%). We
groups 3,7 and 11 have high densities of hemlocks

at about
that
located
and have
their centers
o
o
o
o
o
o
o
o
o
o
o
o
o
o
o
o
examples
in this paper.
presented
peak
of this correlogram
(distance class 9, whose center
can be readily interpreted
is the 485 m distance)
as the distance among peak centers, in the spatial
see Fig. 10, where
of the hemlock;
distribution
o
o
for all the remaining
size is larger than the distance

sampling points. The
neighbouring
o
o
as the basis
used
the
about
the geographical
data set will be
patch
o
o
o
o
o
o
data
territory, plus geomorphological

200 sampling sites, and of course
This
of the quadrats.
locations
pairs of neighbouring
points
is positive
and significant;
o
o
o
o
o
o
height were noted and identified at species

level. The data to be analysed here consist of the
in this
of the 28 tree species present
abundance
spatial
ing distances
?
o
o
o
separated also by
at
5 cm diameter
breast
elements
rows
than
along staggered
Trees with more
dividual
intervals
50 m.
can
o
o
southwestern
o
o
in
border,
o
data were
Qu?bec. A systematic
sampling design was used
to survey 200 vegetation
(Fig. 2) each 10
quadrats
m
were
in size. The quadrats
placed at 50-m
by 20
1.

the
hemlock
classes;
Moran's
the
/
116
distance.
The
last few distance
because
interpreted,
all pairs of localities.
Two-dimensional
they each
classes
cannot
<
contain
be
1% of
Sokal (1986) have proposed

to compute
correlo
in pre-speci
grams only for object pairs oriented
fied directions,
and to represent either a single, or
several of these correlograms
together, as seems
fit for the problem at hand. Computing
structure
functions
in pre-specified
directions
is not new,
correlogram
and
All-directional
assume
the
has
been
traditionally
done
Fig. 4 displays
in variogram
a two-dimen
phe
as mentioned
above.
isotropic,
as
coefficients,
computed
Spatial autocorrelation
1 for all pairs of data points,
in App.
described
analysis
(below).
sional
spatial
correlogram,
sugar-maple Acer saccharum
a mean
the direction,
irrespective
produce
over all direc
smoothed
value of autocorrelation,
program used by Oden & Sokal (1986); the same
nomenon
correlograms
to be
of
tions. Indeed, a spatial autocorrelation

coefficient
class, which
gives a single value for each distance
is fine when
studying a transect, but may not be
several
for
phenomena
occupying
appropriate
geographic dimensions (typically 2). Anisotropy

is however
in ecological
field
are
often
generated
spatial patterns
Oden &
geophysical
phenomena.
often
data, because
by directional
encountered
computed
from our
were made
tation data. Calculations
for
the
test vege
with the very
information
could also have been represented
by
a set of standard correlograms,
each one corre
to one of the aiming directions.
In any
sponding
of aniso
case, Fig. 4 clearly shows the presence
in
the
not
which
could
structure,
tropy
possibly
have been detected
in an all-directional
correlo
the north-south
gram:
much
is
range of A. saccharum
than the east-west
range
larger (ca 500 m)
(200m).
Two-dimensional
spectral
analysis
This method, described by Priestly (1964), Rayner

(1971), Ford (1976), Ripley (1981) and Renshaw
Ford
(1984), differs from spatial autocorre
in the structure function it uses. As
lation analysis
in regular time-series
spectral analysis, the method
&
assumes
the data
to be
(no spatial
stationary
a combination
of sine
function rgh, as well
patterns. An autocorrelation
as a periodogram
with intensity I{p, q), are com
and made
gradient),
of
puted.
Just
as with Moran's
/, the autocorrelation
are a sum of cross products of lagged data;
in the present case, one computes
the values of the
for
of lags
function
all
combinations
possible
rgh
values
for the sugar-maple
correlogram
are geographic
and are the
directions
is sym
2. The
lower half of the correlogram
Fig. 4. Two-dimensional
Acer saccharum.
The
same
metric
tance
Moran's
dashed
as in Fig.
to the upper
half.
Each
ring represents
are as follows:
full boxes
class.
Symbols
/ coefficients,
half-boxes
on too
are based
boxes
100-m
dis
are
significant
are non-significant
values;
few pairs and are not con
the values
taken
represent
by
hachured
through
( + 0.5 to +0.2)
0.1), light dots ( 0.1
( + 0.2 to +0.1),
heavy dots ( + 0.1 to
to
( 0.2 to
0.5).
0.2), to white
sidered.
Moran's
Shades
of
/:
black
from
gray
(g, h) along the 2 geographic sampling directions

/ on the contrary,
the lag d
(App. 1); inMoran's
Besides
is the same in all geographic
directions.
one computes
a
the autocorrelation
function,
for
all
Schuster
two-dimensional
periodogram,
of spatial frequencies
(/?, q) (App.
as graphs
well
by
(first proposed
& Ford? 1983) called the ?-spectrum
combinations
as
1),
Renshaw

117
a
that summarize
and the 0-spectrum
respectively
of the dominant
the frequencies
and directions
1 for
waves that form the spatial pattern. See App.
details.
computational
Two-dimensional
spectral
been
used
to analyse
has recently
in crop
analysis
spatial
patterns
plants (McBratney & Webster 1981), in forest

canopies (Ford 1976; Renshaw & Ford 1983;
Newbery et al 1986) and in other plants (Ford&
of this technique
1984). The advantage
of
it allows
data,
analysis
anisotropic
are frequent
in ecology.
Its main
dis
Renshaw
is that
which
-6
for time
is that, like spectral analysis
advantage
a large data base;
this has
it requires
series,
from being applied to a
the technique
prevented
one
array of problems.
Finally,
the autocorrelogram
that although
wider
notice
35,-,-,-,-,-,-,-,-.-,-.-.-.-.
should
25-
can be
20.
as a
way
assumes
the periodogram
on the contrary the spatial pattern to result from
a combination
of repeatable patterns
; the periodo
are
R
and 0 spectra
very sensitive
gram and its
to repeatabilities
in the data, but they do not
essentially
interpreted
Moran's
correlogram,
detect
other
involve
in the
same
types of spatial patterns
which
R-spectrum b
30 T
1
1
15
10
do not
12
14
16
18
20
120
140
160
180
repeatabilities.
- -
saccharum.
For
.-.-.-.-._._,_,_,_,_L
12j-?-
? ,0
0-spectrum
the sake of this example, and

the data to form a
requires
since
this method
regular,
sugar
grid, we interpolated
rectangular
to
abundance
data by kriging (see below)
a rectangular data grid of 20 rows and 12
maple
obtain
10
R
3 - Fig. 5a shows
the two-dimen
Example
of our vegetation
data for
sional periodogram
Acer
an
The
periodogram
(Fig. 5a) has
5% significance,
since 4 values exceed the
critical Bonferroni-corrected
value of 6.78; these
4 values
72% of the spatial
together
explain
variance of our variable, which
is an appreciable
columns.
o-u?-_o=?o__c?--b-Hj^?^"t>-o?n?g?
overall
amount.
The most
-2\-,-1-,-,-,-,-,-,-,-,-,-,-,-,-,-,-,-,-,
-20
0
20
40
60
80
100
Angle
5.
The
ordinate
(a) Two-dimensional
periodogram.
the intensity
of the periodogram.
(b) ?-spectrum.
in
Bonferroni-corrected
values
(c) 0-spectrum.
significant
are represented
the spectra
for an overall
by dark squares,
Fig.
represents
prominent
values
are the tall blocks
located at (/?,q) = (0, 1) and (0, - 1); together,

they represent
62% of the spatial variance
indicate
that
the dominant phenomenon
they
and
significance level of 5%.
is an
wave with
a frequency
east-west
of 1 (which
means
occurs once in the
that the phenomenon
across the map). This struc
east-west
direction
"1
ture has
an angle
of 0 = tan
(0/[ 1 or
1])
= 0?
and
with
0-spectrum;
R =
is the dominant
+ l2) =1,
y/(02
Z?-spectrum.
This
feature
its
it also
east-west

wave,
of
the
frequency
dominates
with
the
its crest
118
in the north-south
direction,
elongated
visible on the map of Fig. 13 a.
The next
2 values,
that ought
is clearly
to be considered
together, are the blocks (1, 2) and (1, 1) in the

The
periodogram.
corresponding
angles
are
0 = 26.6? and 45? (they form the 4th and 5th

in the 0-spectrum),
for an average angle of
?
of the structure they
; the R frequencies
= 2.24 and
+
are
1.41, for an
represent
q2)
yf(p2
that the values of p and q
average of 1.8. Notice
values
about
35
as if the 2 geographic
axes
vertical
and
in
horizontal
directions
(the
Fig. 13)
were of equal lengths, as explained
1;
in App.
have been
standardized
these periodogram
values indicate very likely the
direction of the axis that crosses the centers of the
2 patches
of sugar-maple
of Fig. 13 a.
bottom
Two other periodogram
high
in
the middle
values
are
and
relatively
(5.91 and 5.54) but do not pass
the
test of significance,
proba
the
of blocks of data in our
because
number
bly
In
regular grid is on the low side for this method.
to
the
is
any case,
90?,
angle they correspond
which
is a significant
value in the 0-spectrum.
These periodogram
values indicate obviously
the
Bonferroni-corrected
north-south
the centers of the

crossing
in the upper and middle
parts of
direction
2 large patches
=
Fig. 13a (R 2).
These
dimensional
variograms
anisotropy
results
are
consistent
with
the
two
the
correlogram
(Fig. 4) and with
and
confirm
the
of
presence
(Fig. 9),
in the A. saccharum
data. They were
& Ford
using the program of Renshaw
Ford
of
vege
(1984).
(1976) presents
examples
tation data with clearer periodic
components.
computed
is the Mantel
ing out such analyses
This method
deals with 2 distance
2 similarity matrices,
obtained
independently,
and describing
the relationships
among the same
among the
sampling stations (or, more generally,
same objects). This type of analysis has two chief
of application
in community
Let us consider a set ofn sampling
we want
the first kind of application,
domains
a matrix
of ecological
ecology.
In
stations.
to compare
stations
among
distances
(X) with a matrix of geographic distances (Y)

same
the
among
tances
inmatrix
stations.
The
X can be obtained
dis
ecological
for instance by
to
with respect
all pairs of stations,

comparing
their faunistic or floristic composition,
of the numerous
association
coefficients
using one
available
in the literature; notice that qualitative

(nominal)
data can be handled as easily as quantitative
data,
since a number of coefficients
of association
exist
for this
type of data,
quantitative,
data. These
instance by Orl?ci
Legendre
others;
and
even
semi-quantitative
coefficients
have
for mixtures
and
been
of
qualitative
for
reviewed
(1978), by Legendre &
and
(1983a
see also Gower
of
comparison
and by several
1984a),
& Legendre
(1986) for a
Y contains
coefficients.
Matrix
distances
of
among
geographic
pairs
or
that
in
their
distances
other
m, km,
stations,
is,
units of measurement.
The scope of the study is
only
to determine
whether
the ecological
distance
as the samples get to be geographically
farther apart, i.e., if there is a spatial gradient
in
the multivariate
In
to
data.
order
do
ecological
increases
statistic is computed
and
this, the Mantel
as described
in App. 2. Examples
of Mantel
in the context
Ex.
The Mantel
test (1967).
or
matrices,
in this
of
spatial analysis
as well
as
paper,
tested
tests
are found
in Upton
in
&
Fingleton's book (1985).
test
test can be used not only in spatial

but also to check the goodness-of-fit
of
The Mantel
one of the scopes of community
is
ecology
a number of
between
of relationships
- on the one
- the
variables
species
biological
the
hand, and many abiotic variables
describing
Since
the study
on the other, it is often necessary

to
inmultivariate
these problems
terms, to
abundance
for instance
the simultaneous
environment
deal with
study
fluctuations
of several
species. A method
of carry
analysis,
data to a model.
course, this test is valid only

Y is obtained
independently
measures
from the similarity
in matrix X - either
or else if it derives from
by ecological
hypothesis,
if the model
Of
inmatrix
an analysis
of a different data set than the one
in elaborating
matrix X. The Mantel
test
used
cannot
be used
to check
the

conformity
to a
119
of a model
matrix
Goodness-of-fit
recently
derived
Mantel
from
tests
have
data.
used
been
used
coefficient
to investigate
very
to
of impor
related
questions
hypotheses
of climax
&
like the concept
(McCune
in vegetation
to compute
the X
studies
precise
tance,
control model
Allen
1985) and the environmental
can be
Another
1987).
(Burgman
application
among
Estabrook
& Rogers
similarity coefficient makes
to assemble mixtures
of quantitative,
and qualitative
into an
data
semi-quantitative
it possible
overall measure
of directional
studied
similarities
between
lobsters
and crabs.
were
area under study,
4 - In the vegetation
Example
2 tree species are dominant,
the sugar-maple Acer
saccharum
and the red-maple A. rubrum. One of
or both,
these species,
the 200 vegetation
of niche
hypothesis
are present
in almost all of
In such a case, the
quadrats.
comes to mind.
It
segregation
the
null
that
by stating
hypothesis
of the 2 species is the same, and the
that there is a difference.
hypothesis
that we
how
as ecologists,
to represent
established,
similar are the various pairs of semi-ordered
or unordered
from the point of
classes, considered
view of tree growth. The environmental
similarity
as X in Fig. 6.
is represented
matrix
The
of niche segregation
ecological
hypothesis
A. saccharum
and A. rubrum can be
between
the habitat
translated
alternative
are going to test this hypothesis

the environmental
data to a model
to
the
alternative
test. The
Mantel
by comparing
(Fig. 6), using

data were chosen
hypothesis
environmental
to represent factors likely to influence the growth

are: quality of
of these species. The 6 descriptors
stoniness
(7 semi-quantitative
drainage
classes),
of
the
soil
(7 semi-quantitative
classes),
topo
unordered
(11
graphy
qualitative
classes),
directional
exposure (the 8 sectors of the compass
= flat
card, plus class 9
land), texture of horizon
1 of the soil (8 unordered
and
classes),
qualitative
classes,
(6 unordered
geomorphology
qualitative
in Example
8 below). These data were
described
Y: Dominancemodelmatrix
of the alternative
as follows : each of the 200 quadrats

hypothesis
was
as having
or
coded
either A. saccharum
a model
Then,
similarity
was
contain
constructed,
among quadrats
were
that
dominant
ing l's for pairs of quadrats
for the same species
and
(maximum
similarity),
dominant.
matrix
differing as to the domi

is
species (null similarity). This model matrix
as
Y
as
in
is
where
it
shown
if
represented
Fig. 6,
all the A. saccharum-dominated
came
quadrats
0's for pairs of quadrats

nant
first, and all the A. rubrum-dominated

quadrats
came last; in practice,
the order of the quadrats
does not make any difference,
insofar as it is the
same in matrices
X and Y.
One
Mantel
matrix
X: Environmentalsimilarity
into a model-matrix
A. rubrum
corresponding
of similarity; for the descriptors

exposure and soil texture, the partial
to the overall coefficient

contributing
drawn from a set of partial similarity values
can be tested
We
(Estabrook
quadrats
similarity
& Rogers
1966; Legendre & Legendre 1983a, 1984a). The
found in Hudon & Lamarche (in press) who

competition
an Estabrook-Rogers
can obtain
the sampling
of the
distribution
statistic
realiza
by repeatedly
simulating
tions of the null hypothesis,
through permutations
to the lines and
of the quadrats
(corresponding
columns)
Mantel
in the Y matrix,
and
recomputing
the
statistic between X and Y (App. 2). If
there is no relationship
between matrices
and Y, we can expect the Mantel
statistic to
have a value located near the centre of this sam
indeed
if such a relation does

while
distribution,
statistic to be more
exist, we expect the Mantel
extreme than most
of the values obtained
after
pling
Sugar-maple
Fig.
model
6. Comparison
between
Red-maple
of environmental
(matrix Y), to test

the sugar-maple
Sugar-maple
Red-maple
(matrix X) to the
of niche
segregation
data
the hypothesis
and the red-maple.
random
Mantel
significant
permutation
statistic was
at p<
of
the model
computed
0.00001,

using
matrix.
and found
The
to be
in the present
120
case Mantel's
App.
must
that
t test, mentioned
in the remarks of
test. So, we
instead
of
the
2,
permutation
and accept the idea
reject the null hypothesis
there is some measurable
niche differentia
^4. saccharum
tion between
and A
rubrum. Notice
of this analysis
is the same as
that the objective
in classical discriminant
analysis. With a Mantel
one does not have to comply with
test, however,
of discriminant
the restrictive assumptions
analy
The Mantel
correlogram
on
Relying
a Mantel
test between
data
and
model, Sokal (1986) and Oden & Sokal (1986)

an ingenious way of
a correlo
computing
for
are often
multivariate
such
data
gram
data;
encountered
in ecology
in
and
population
found
The
that are rarely met by ecological

sis, assumptions
one can model
at will the rela
data; furthermore,
among
plants
(or animals)
tionships
by com
is to express
principle
ecological
stations
among
relationships
sampling
by means
of an X matrix of multivariate
and then
distances,
to compare X to a Y model matrix,
different for
each distance
for distance
class 1, for
class;
appro
puting matrix X with a similarity measure
as
as
to
to the
the
well
data,
priate
ecological
nature of the problem,
instead of being imposed
or a
a Mahalanobis
the use of an Euclidean,
station pairs (that belong

instance, neighbouring
to the first class of geographic
are
distances)
linked by l's, while
the remainder
of the matrix
zeros only. A first normalized
contains
Mantel
as it is the case
chi-square distance,
classical
multivariate
methods.
test made
case, the Mantel
mixture of semi-quantitative
inmost
of the
In
the present
to use a
it possible
and qualitative
varia
bles, in a rather elegant analysis.

do these
To what
environmental
variable(s)
tree species react? This was tested by a series of
a posteriori
each of the 6 environ
tests, where
was
the
mental
variables
tested in turn against
an Estabrook
&
model-matrix
Y, after computing
Rogers
variable
could
for that environmental

tests
that these a posteriori
matrix
similarity
only. Notice
have
table
by contingency
a
since they involve
single semi-quantita
analysis,
at a time; they were
tive or qualitative
variable
the
done by Mantel
testing here to illustrate
In any case,
of the method.
domain of application
tests show that 3 of the environ
these a posteriori
mental
been
are
significantly
stoniness
(p<
variables
model-matrix:
(p
phy
(p<
conducted
and
0.00028)
0.00001);
the other
related
0.00001),
topogra
were
not
related to Y. So the three first varia

significantly
either for studies of the
bles are likely candidates,
tor.
other
differences
adaptive
or for further
2 maple
species,
is presented
such
One
analysis
analyses.
8 below,
for the geomorphology
descrip
physiological
these
between
spatial
as Ex.
or
statistic
for this distance

class.
(r) is calculated
is repeated for each distance
process
class,
each time a new model-matrix
Y, and
building
the normalized Mantel
statistic. The
recomputing
The
of the normalized
Mantel
graph of the values
statistic
classes
against distance
gives a multi
variate correlogram;
each value is tested for sig
in the usual way, either by permutation,
nificance
or using Mantel's
normal approximation
(remark
in App. 2). [Notice that if the values in the X

or else
are similarities
instead of distances,
inmatrix Y,
if the l's and the O's are interchanged
then the sign of each Mantel
statistic is changed.]
matrix
Just as with
a univariate
(above), one
correlogram
a
is advised to carry out
global test of significance
of the Mantel
correlogram
using the Bonferroni
before trying to interpret the response of
method,
the Mantel
statistic for specific distance
classes.
to the
geomorphology
3 variables
genetics.
5
Example
pling stations
species
coefficient
the Bray
Legendre
gram was
- A
similarity
sam
among
from the 28 tree
matrix
was
computed
the Steinhaus
data, using
of similarity (also called the Odum, or
and Curtis
coefficient:
&
Legendre
abundance
1983 a, 1984a),
and the Mantel
correlo
computed
(Fig. 7). There is overall sig
in this correlogram,
nificance
since many
of the
individual values exceed the Bonferroni-corrected
= 0.0025. Since there is
level a' = 0.05/20
signifi
cant positive autocorrelation
classes
and significant
negative

autocorrelation
in
121
0.14
j
0.120.10-y\
statistical
univariate
0.08
0.06 .L
0.04
0.02
?
S
-0.06 1
0
does not lend itself to any

(below),
test of hypothesis.
is a
The variogram
limited to quantitative
varia
method,
is based
\
*.
Nl
that occur in
bles, allowing to analyse phenomena
or
2
3
dimensions.
one,
Burrough
geographic
to variogram analysis
(1987) gives an introduction
*-*^
1 2
5 6
7 8
Distance
10 11 12 13 14 15 16 17 18 19 20
classes
tree com
for the 28-species
Fig. 7. Mantel
correlogram
structure.
classes
distance
See text. Abscissa:
(one
munity
Mantel
standardized
is 57 m);
ordinate:
unit of distance
of the
values
Dark
statistic.
squares
represent
significant
<
Mantel
statistic
0.05).
(p
for ecologists.
Before using
sure
that
one must make

the variogram,
are stationary, which means
the data
that the statistical properties

(mean and variance)
of the data are the same in the various parts of the
area under study, or at least that they follow the
which means
that the incre
'intrinsic hypothesis',
a given
of
located
between
all pairs
points
zero and a finite
distance
d apart have a man
the same in the various
variance
that remains
ments
the large distances,
the overall shape of this cor
could
attributed
either to a vegetation
be
relogram
or
a
to
structure
with
steps
(Fig. Id)
gradient
auto
(Fig. le). In any case, the zone of positive
correlation
lasts up to distance class 4, so that the
size of the 'zone of influence' of multi
average
size of asso
(the mean
or (4
4
is
about
distance
classes,
ciations)
?
x
m.
57 m)
This estimation
classes
230
is con
in Fig. 10, where many of the
firmed by the maps
associations
delimited
have about
by clustering
variate
autocorrelation
that size.
of
parts
the
area
for distance
variance,
the semi-variance
this value of
study;
class d, is twice the value of
function
relaxed
y(d). This
under
form of the stationarity

it pos
assumption makes
or for that matter any
sible to use the variogram,
structure
other
correlograms),
(for instance spatial auto

data. Of course, a
ecological
function
with
large-scale
spatial structure, if present, will neces
be
sarily
picked up by the structure function and
may
mask
finer
structures;
large-scale
spatial
should be removed by regres
surface analysis) or some other form
in particular,
trends,
sion (trend
and description
Detection
of spatial
structures
above, the different

types of cor
relograms, outlined in the section entitled Testing
for the presence of a spatial structure', do provide
As mentioned
a description
that are more
of spatial
structures.
Other methods,
can also be
descriptive,
exclusively
for this purpose. They
used
are presented
in this
There
mental
of other,
are two types of variograms

the theoretical.
The
finer
: the experi
and
experimental
from the
is computed
variogram
(semi-variogram)
1. It is presented
data using the formula in App.
as a plot of y(d) (ordinate)
as a function of dis
a correlogram.
tance classes
As
like
(d), just
in App.
is a distance-type
1, y(d)
so
c
to Geary's
that it is related
function,
can be
coefficient.
The experimental
variogram
noticed
section.
The
of modelling
before
the presence
structures can be investigated.
The
often
(Matheron
semi-variogram
1962),
for simplicity,
is related to spa
variogram
tial correlograms.
It is another structure function,
called
allowing
non as
method,
as a description
of the structure function of
the spatial phenomenon
and in this way it is of
in
the spatial structure.
help
understanding
used
variogram
to study the autocorrelation

phenome
a function
of distance;
this
however
on which
the kriging contouring method
The variogram was originally designed

ing engineers, as a basis for the contouring
by min
method
known as kriging (below). This is how it became

known
to ecologists,
among
whom
its use
is
spreading (Burrough 1987). To be useful for

122
kriging, a theoretical variogram has to be fitted to
the experimental
the adjustment
of a
one;
to
theoretical
the
variogram
experimental
function
the parameters
used by the
provides
method.
The most
of these
important
kriging
are (l)the
structure, which
range of influence of the

is the distance where the
parameters
spatial
is
stops increasing;
(2) the sill, which
variogram
the ordinate value of the flat portion of the vario
the semi-variance
is no longer a
gram, where
function
of direction
and distance,
and cor
responds
to the variance
of
the
samples;
and
eventually (3) the nugget effect (see below). As in

curve fitting, the user must
any type of nonlinear
decide what type of nonlinear
is wanted
function
to adjust to his experimental
this step
variogram;
and
into the
both
experience,
insight
requires
ecological
theoretic
4 of
ment.
described
ferences
under
study. Several types of

are often used for this adjust
process
functions
the most
them,
in App.
between
common
1 and
illustrated
these
theoretic
are
ones,
in Fig. 8. Dif
functions
lie
in the shape of the left-hand part of the

mostly
curves, near the origin. A linear variogram indi
cates a linear spatial gradient;
this model has no
sill. Gaussian, exponential and spherical variograms
of the size of the spatial influence
give a measure
is
of the process
(patch size, if the phenomenon
as well as the shape of the drop of this
patchy),
as one gets farther away from the center
of the phenomenon;
the exponential model
does
not necessarily
have a sill. A flat variogram,
also
influence
called
the absence
'pure nugget effect', indicates
of a spatial structure
in the data, at least at the
scale
nugget
were made.
the observations
effect
refers
to variograms
The
so-called
that do not go
some
the origin of the graph, but display
even at distance
of variance
zero; this
effect may be caused by some intrinsic random
through
amount
or it
variance),
may suggest that the sampling has not been per
formed at the right spatial scale. Variograms
have
measure
to
been
used
the
fractal
dimen
recently
variability
in the data
sion of environmental
(sampling
gradients
(Phillips
1985).
separate variograms
Mining
engineers compute
to determine
for different
if the
spatial directions,
or
seen
structure
not.
is
We
have
spatial
isotropic
that this procedure
has now been extended
as well. The spatial structure
to correlograms
is
to
said
be isotropic when
are the
the variograms
same regardless of the direction
of measurement.
above
2 different
geometric
Geometric
can be detected:
of anisotropy
and
stratified
anisotropy
anisotropy.
(same sill, different ranges)
anisotropy
kinds
ismeasured
is equal
by the anisotropy ratio, which
to the range of the variogram
in the direction
the longest range, divided by the range
producing
in the direction with the smallest range. Stratified
(or zonal) anisotropy
(different sills, same range)
refers to the fact that the sills of the variograms
In the
may not be the same in different directions.
one
or
or
of
the
other
of
presence
type
anisotropy,
to obtain accepta
both, there are three solutions
ble interpolated maps
by kriging: one can com
pute
compromise
variogram
the formulas in David

Huijbregts
program
parameters,
using
(1977) or in Journel &
(1978); secondly, one can use a kriging

use of the parameters
that makes
of
in different
computed
variograms
separately
directions
of the physical
space (2 or 3, depending
on the problem);
or finally, one can use 'general
ized
intrinsic
functions
of order k?
random
1973) that allow for linear or quadratic
(Matheron
trends in the data.
6 -
were
Experimental
variograms
A.
for
Fortin
in
saccharum,
computed
by
(1985),
the 45? and 90? directions
(window: 22?), and in
?
?
all directions
the 45 and 90
9).
(Fig.
Comparing
as
shows the presence
of anisotropy,
variograms
Example
was
Fig.
8. Four
models.
of
the
most
common
theoretic
variogram
in Fig. 4. The range in the 45?

(dashed line) is about 445 m, while the
variogram
?
is about 685 m, so that
range in the 90 variogram
observed

123
methods
Clustering
constraint
with spatial contiguity
structures can be done by

multivariate
Describing
are classical
of clustering,
which
the methods
of multivariate
methods
600
Distance
analysis,
in
and
spatial contiguity
results are represented
on a map, the multivariate
structure of the data for instance - will be clearly
plant associations
described
by the map.
by clustering
If the clustering
particular
constraint.
400
data
with
800
with
Clustering
has been suggested
(m)
constraint
spatial contiguity
authors
since 1966
by many
& Burrough
1966; Webster
(e.g., Ray & Berry
1972; Lefkovitch 1978,1980; and others), in such

different
as
fields
science,
pedology,
political
and ecology.
economy,
psychometry
Starting
from multivariate
data, the common need of these
was
to establish
geographical
regions
of adjacent sites (i.e., a choropleth map: see
'Estimation and mapping'
below) which would be
In
with respect to certain variables.
homogeneous
authors
made
order
200
400
800
600
Distance
matrix
1000
1200
(m)
a
to do this, it is necessary
(1) to compute
of similarity among sites from the variables
on which
these homogeneous
regions have to be
based (of course, this step applies only to cluster
that are similarity-based),
then
ing methods
with
(2) proceed
with
methods,
the algorithm
any of the usual

the difference
that one
to cluster
only
clustering
constrains
sites or
these
site
that are geographically

The
groups
contiguous.
to the program
is provided
in the form
constraint
or spatial links, among
of a list of connections,
localities.
be
may
see App.
1.
of ways:
to existing programs
which we will not
problems
neighbouring
established
Connections
in a variety
such constraints
400
600
Distance
Fig. 9. Three
experimental
See
Acer
saccharum
data.
Ordinate:
values
800
(m)
On
variograms
text. Abscissa:
of the semi-variance
Adding
raises algorithmic
discuss here. Clustering
computed
distance
function
for
the
classes.
y(d). Dashed
lines: ranges.Modified from Fortin (1985).
as
can
be
ratio
computed
anisotropy
685/445 ? 1.5. The all-directions
variogram does
not clearly render this information.
the
esting properties.
set of mathematically
that are geographically
the well-known
where
different
with
constraint
has
inter
the one hand, it reduces the

to those
solutions
possible
this avoids
meaningful;
of
problem
clustering methods,
solutions may be obtained
after
to the
different
applying
clustering
algorithms
same data set; constraining
to
all these algorithms
are
results
that
consistent
produce
geographically
forces
them
solutions.
On
to converge
the other
towards
hand,

very similar
the partitions
124
in this way reproduce a larger fraction of
the structure's spatial information
than equivalent
obtained
without
constraint
partitions
(Legendre
ecological
cluster
1987). Finally, constrained
agglomerative
uncon
is
faster
sets
with
data
than
the
ing
large
clustering
conducted
strained equivalent,
because
the search for 'the
next pair to join' is limited to adjacent groups only
contiguity
from the
(Openshaw 1974; Lebart 1978).
described
obtained
Example
from our
Steinhaus
7 - A vegetation map was constructed

as follows.
same
test data,
(l)The
was
matrix
stations
among
similarity
as
in Ex. 5; it is based upon the 28 tree

data. (2) The spatial relation
abundance
species
ships among sampling quadrats were represented
used
by a list of connections
the list was established
Delaunay
;
among close neighbours
in the present case by the
method
(App. 1). The
triangulation
of a connection
2 quadrats
between
presence
tells the clustering
that these 2 locali
programs
ties are located
close
be included
eventually
to one another and thus may

in the same cluster, if their
allows.
(3) Agglomerative
similarity
with
constraint was
spatial contiguity
on the similarity matrix.
The spatial
was
constraint
read by the program
or 'link edges',
connections,
We
used a proportional-link
agglomerative
algorithm
(with 50% con
list of
above.
linkage
nectedness:
Sneath
a series
1966), that produced
one for each clustering
of maps,
level (Legendre
& Legendre
1984b). The map with 13 groups was
retained as being ecologically
the most meaningful
remain unclustered
at that
(Fig. 10a); 5 quadrats
level. Recognizing
13 groups
the
that
implies
=
mean
area per association
is 740000 m2/13
56 923 m2/association,
to an aver
corresponding
=
area
diameter
of (56923)1/2
238.6 m; this
age
size of the
compares
very well with the average
zone of influence
of our species
associations
found
in the Mantel
Agglomerative
small distortions
the hierarchical
results
from
correlogram,
230 m
(Ex. 5).
clustering may have produced
of the resulting map, because of
nature of the classification
that
such
tried to render our
So, we
algorithms.
as homogeneous
as
sequential
13 groups
in terms of vegetation
possible
composition,
using
a fc-means algorithm
(MacQueen
1967) with spa
tial contiguity
constraint.
A A;-means algorithm
uses an iterative procedure
of object reallocation
the sum of within-group
dispersions.
tends to produce
compact
to minimize
type of algorithm
clusters in the variable
This
space (here, the vegetation

is exactly what we are looking for;
to expect
is no reason however
this phe
nomenon
to affect the shape of the clusters
in
data),
there
which
our program with

space. We provided
geographic
the list of constraining
connections
in
computed
the 13-group
classification
step 2 above, with
in step 3 to be used as the starting con
obtained
figuration
clustered
10. Map
of
obtained
species),
Fig.
strained
where
by
vegetation
by constrained
clustering,
agglomerative
13 groups were
are materialized
map
the multivariate
by
structure
(a) Space-con
at the level
proportional-link
linkage,
the five unclustered
obtained;
dots,
space-constrained
(28
of
(b) Optimization
fc-means
clustering.
quadrats
the previous
(temporarily
allocating
to the group
quadrats
them geographically),
coordinates
computed
larity matrix
(since
the
that
un
enclosed
a set of principal
the Steinhaus
simi
and with
from
our fc-means
com
program
raw
from
variables,
variances
putes within-group
and not from a similarity or distance

matrix).
The
125
of the optimized
groups is shown in Fig. 10b.
of groups
remained
the same, of
number
map
The
19 objects out of 200 changed group

4
unmodified:
remained
groups
groups
(10%).
number
1, 6, 10 and 13 in Fig. 10.
were compared
The 2 13-group classifications
but
course,
to the raw species abundance

tables. This work
contingency
data
in a series of
was
facilitated by
range into
dividing first each species' abundance
a few classes, following
described
the method
by
Legendre & Legendre (1983b). Comparing the

of the 2 classifications,
the groups
were
classification
by the ?>means
to
than
those
easier
characterize
produced
slightly
Their main
classification.
by the agglomerative
interpretations
the A.
cated between
rubrum and^4.
and
3,7
1.
of communities
the development
by
et
structure
al
1985).
abrupt
(Legendre
jumps
has been used to segment
Since then, this method
transects
of ecological
data
(Galzin &
spatial
series
1988), as well as paleontological
Legendre
methods
for segmenting
the spatial or
account
lo
saccharum
such
taking
into
contiguity
of
series,
temporal
samples, have been proposed by Fisher (1958) for

univariate
economic
soil data,
by Hawkins
data, by Webster
(1973) for
& Merriam
for univariate
(1973) and for multivariate (1974) geologic data,

by Gordon & Birks (1972, 1974) and by Gordon
data. This work
(1973) for pollen
stratigraphie
has been summarized
by Legendre
(1987).
11.
in the three follow

is highest
diversity
of
dominated
stands,
by black ash
ing groups
Fraxinus
birch
and
Betula
alle
nigra
yellow
Causal
ghaniensis:
- In the
bottom
Although
and have
Species
5.
Group
- With
red
basswood
Fence-shaped
characterized
and American
to
ash Fraxinus
Tilia americana:
region
by white
and
pennsylvanica
8 and 9.
Groups
cleared
land)
(formerly
cedar Thuja occidentalis
elm Ulmus
with
but, con
F. nigra
and
americana
few
group 5,
trary
B. alleghaniensis:
13.
Group
or multivariate
Univariate
data
modelling
are used by ecologists

empirical models
their usefulness, modelers
often prefer to
include only the specific (ecological)
hypotheses
they may have about the factors and mechanisms
the process under study. The purpose
determining
of a kettle, with aspen Populus

cedar Thuja occidentalis
tremuloides, white
americana:
elm
and
American
Ulmus
of
model
of climatic
progress,
including the reconstruction
tree
fluctuations
and
the seg
by studying
rings,
of pollen
data.
Other
menting
stratigraphie
areas: Group
10.
- A. saccharum
4 and 12.
stands: Groups
- Stands dominated
white
by
pine Pinus strobus
6.
tremuloides: Group
and aspen Populus
Hemlock
canadensis:
stands, Tsuga
Groups
-
a
each group fusion,
before
structure;
munity
test
statistical
is
that
permutation
performed,
terms the ecological
into statistical
translates
(Bell& Legendre 1987).Other applications are in
produced
are the following:

biotic characteristics
Open area, with rare A. saccharum: Group
- A. rubrum
2.
stands, Group
Betula
Oldfield-birch
stands,
populifolia,
in P.L.'s
time). One such program was developed
lab to analyse
with
the
successions,
ecological
of locating the abrupt changes
explicit purpose
that may occur along successional
series of com
or
of modelling
is then to verify that experimental
field data do support these hypotheses
('causes'),
and to confirm the relational way in which
they
are assembled
into the model. Given
the impor
tance
of
in our
space
of spatial
complete without
review
analysis
ecological
methods
this
theories,
would not be
often need
how space can be

mentioning
in the calculation
of relationships
among
2 variables
variables.
if
related
may
appear
both of them are linked to a common
third one;
tiguity constraint
correlations,
seem to be
that
form
a surface,
in space, instead of covering
to be summarized
by identifying break
ing points along the series. Several authors have
con
to use clustering methods
with
proposed
transect
in a single dimension
(space
or
included
space
is a good
candidate
for creating such false

variables
may
actually
linked because
they are driven by a
since

126
spatial gradient. Even if correlation does
of correlation,
the absence
not mean
causality,
common
or linear, is sufficient to abandon

the
2 variables.
of a causal link between
monotonie
hypothesis
It is thus
in
interested
for ecologists
important
the
to
whether
check
causal relationships
spatial
at least in part,
gradient of A could be explained,
variable
structured
B, or if an
by a spatially
is not
2
variables
between
correlation
apparent
to a common
to be ascribed
unmeasured
or untested
spatial
structure
(an
varia
space-structured
ble causing A and B independently). There is still

to go before space can be included as
but we
in complex
ecological models,
can
in
at
be
included
least
show how it
simple
some way
a variable
will
C, as in Pearson
partial correlations).
(4) Test as
or
or
usual, either by permuting A'
B',
by Mantel's
to what
normal approximation.
This is equivalent
all 3 matrices.
by permuting
are not easy to interpret;
& Troussellier
(1988) have shown the
would
be obtained
Partial
Mantel
Legendre
in terms of significant Mantel

and
of all the possible
three
consequences,
partial Mantel
matrices
tests
statistics,
implying space. As in the case of

test (App. 2), the restrictive
influence
has not been fully
linearity assumption
models
the Mantel
of
the
tests.
yet for partial Mantel
investigated
numerous
of
This
has
type
analysis
applica
in space.
tions for studying variables
distributed
3 other forms of test of partial asso
Actually,
3 distance matrices
have been
involving
are
2
of
based
the
Mantel
these
upon
proposed.
one
&
Cheverud
test,
(Dow
by anthropologists
1985), the second one in the field of psychometry
ciation
models.
Partial Mantel
test
(Hubert 1985); the third one involves multiple

can a partial
bles be calculated,
How
correlation
controlling
between
two varia
for a space
effect?
Smouse et al (1986) dealt with this problem and

of each of the
the variations
suggested expressing
and
two variables by matrices
B) that contain
(A
between
all sampling
in values
the differences
station pairs. On the other hand, as in the Mantel
is expressed
test, the 'space' variable
by a matrix
stations
distances
of
among
geographic
(matrix C). Actually,
well be multivariate
matrices
A and B could
as
A partial
A
and B,
between
is
calculated
statistic
Mantel
for the effect of matrix C. The Smouse
controlling
et al partial Mantel
statistic has the same formula
distance
matrices.
correlation
product-moment
Mantel
from
standardized
coefficient,
computed
are
done as
the
statistics. Actually,
computations
statistic
follows in order to test the partial Mantel
as
partial
for the effect of

A and B, controlling
the
C: (1) compute matrix A' that contains
of
values
of
the
of
the
linear
residuals
regression
over
of C; (2) likewise,
A
the values
compute
matrix B' of the residuals of the linear regression
of the values of B over the values of C; (3) com
between
matrix
pute
(which
Mantel
the Mantel
statistic
between
A'
and
B'
the partial
is just another way of obtaining
B
for
statistic between A and
controlling
on distance
regressions
matrices
1986;
(Manly
Krackhardt 1988).
data to
8 - We will use our vegetation
Example
of
the
environ
much
debated
the
study
question
structures. We will
mental
control of vegetation
between vege
the relationship
study in particular
of the
structure
the
and
tation
geomorphology
structures
sites. Of course, vegetation
sampling
and this can be due
often autocorrelated,
is a
to the fact that biological
reproduction
some
or
to
between
process,
linkage
contagious
are most
either
and
vegetation
composition,
autocorrelated.
substrate
conditions,
and
geomorphology,
so
since
soil
on,
are
if we find a relationship
we will
and
geomorphology,
vegetation
do
the
data
ask the following additional
question:
a
link
causal
between
of
the
support
hypothesis
or is the
structure and geomorphology,
vegetation
So,
between
spurious, resulting from the

fol
and geomorphology
vegetation
some
a
common
low
through
spatial structure,
factor that could affect both?
unstudied
observed
correlation
fact that both
our vegetation
data are multivariate
(28
com
in
the
will
be
they
represented
Steinhaus
of multivariate
by a matrix
putations
as
Ex.
5.
in
similarities,
Space is repre
ecological
Since
tree species),

127
3. Above
Table
statistics
and
partial Mantel
of significance
N.
the diagonal:
simple
associated
probabilities.
and associated
statistics
the diagonal:
Tests
probabilities.
Below
are one-tailed.
Mantel
Geomor-
Vegetation
structure
tests
Mantel
standardized
for
the
of
effect
Vegetation),
controlling
not to be significantly
different
Geomorphology,
from zero; this condition
is not met in Table 3.
Space
phology
N.
Partial
Mantel
states that there is a spatial

in
the
is inde
data, which
component
vegetation
from the spatial
structure
in geomor
pendent
<->
phology
[Geomorphology
Space
Vegetation
If this model were
structure].
supported
by the
we
would
the
Mantel
statistic
data,
expect
partial
(2) The
Nv
tests
If this
Vegetation
structure].
Geomorphology->
model were supported by the data, then we would
the partial Mantel
statistic
expect
(Space
^v
-
Vegetation
structure
Geomorphology
0.38073
= 0.000
= 0.000
for the
(Geomorphology
Vegetation),
controlling
from
effect of Space, not to differ significantly
in Table 3.
that is not met
zero, a condition
0.36449
= 0.000
p
0.12384
= 0.000
p
Space
A-
0.09397
p
0.17053
= 0.000
0.15054
= 0.000
second model
(3) The third possible model (Fig. 11) claims that

structure
spatial
determined
by
partly
is
in the vegetation
data
in the
the spatial gradient
and partly by other factors not
the
sented by a matrix of geographic

The geomorphology
quadrats.
ordered
fied
classes:
qualitative
till ridge, reworked
sea deposits)
distances
variable
moraine
till, kettle,
was used
among
(6 un
ridge, strati
relict channel,
a
to compute
Similarities
simple matching
similarity coefficient.
= 1were
transformed
into distances
S)
(D
tests.
before computing
the Mantel
The results of the simple and partial Mantel
Champlain
are presented
tests (above
Mantel
tests
in Table 3. The
3 simple
the diagonal)
show that both
the vegetation
structure and the geomorphology
are autocorrelated,
as expected,
and also that
a
there exists
relation between
vege
significant
tation
Mantel
and
Notice
geomorphology.
statistic values do not behave
duct-moment
correlation
that
the
like pro
and do not
coefficients,
to be large in absolute value to be significant.
All 3 partial Mantel
tests (Smouse et al 1986) are
at
the Bonferroni-corrected
level
significant
a' = 0.05/3 = 0.01667. Of special interest to us is
on the
the unique
of geomorphology
influence
have
geomorphology,
identified
explicitly
to
According
3
this model,
all 3 simple and all
partial Mantel
tests should be significantly
different from zero.
in the model.
This is indeed what we find in Table 3.

of the correlation
this decomposition
Although
would best be accomplished
stand
by computing
coefficients
ard partial regression-type
(as in path
we can draw
some conclusions
by
at
show
the
Mantel
statistics.
They
partial
looking
the influence
that the Mantel
statistic describing
analysis),
on vegetation
structure
is
geomorphology
for the
reduced from 0.15 to 0.09 when controlling
of
influence
effect
of
The
space.
proper
of
geomorphology
the difference
on vegetation
is then 0.09, while
is
the
part of the influence of
(0.06)
on vegetation
to
that corresponds
geomorphology
the
of
component
spatial
geomorphology
=
(0.15 x 0.38
0.06). On the other hand, the par
the spatial determi
tialMantel
statistic describing
to the influence of
structure, compared
vegetation
To
decide
the
various
space.
among
possible
of interrelations
models
these 3 groups
among
of variables, we have to consider
in turn all 3
possible
competing
as follows.
nation,
the vegetation
spatial
and proceed by elimi

models,
The
states that
first model
(1)
is caused by the
spatial structure
->
structure
of geomorphology
[Space
Fig.
11. Diagram
structure,
of
interrelationships
and space.
geomorphology

between
vegetation
128
of the vegetation
structure not accounted
for by geomorphology
is still large (0.12) and sig
this shows that other space-related
fac
nificant;
tors do influence the vegetation
structure, which
is then not
determined
entirely
spatially
by
nation
Work
is in progress
geomorphology.
to
fill
the
gap.
hypotheses
on
other
to map
practicable
each variable
separately,
spondence
Several
analysis,
methods
ping. These
include
structured
quantitative
study of spatially
the varia
phenomena
usually starts with mapping
bles. Ecologists,
like geographers,
usually
satisfy
themselves
with
rather unsophisticated
kinds
Any
of map
common
The 2 most
representations.
kinds are (1) divisions
of the study area into non
since
overlapping
regions,
'many areal phe
nomena
studied by geographers
[and ecologists]
can be represented
as a set of
in 2 dimensions
contiguous,
used
to represent
the nested
of some
space-exhaustive
nonoverlapping,
1977), and
(Boots
polygons'
contoured
maps,
altitudes
isolines
continuous
or
(2)isoline maps,
for instance by geographers
on topographic maps, where

intensities
represent different
can be
Both
variable.
types
software. Before attempt
by computer
produced
a map,
ing to produce
by computer,
especially
sure that they satisfy the
must make
ecologists
all parts of the 'active'
following
assumption:
area
a
must
have
non-null
of
study
probability
found
being
be mapped.
in one of the states of the variable to

For instance,
in a study of terrestrial
plants, the 'active' area of the map must be defined

in such a way as to exclude water masses,
roads,
the
like.
and
large rocky outcrops,
Since
the map
obtained
samples
values have
derives
in most
cases
from
a surface,
intermediate
to be estimated by interpolation;
or,
then
for instance.
for
exist
map
interpolated
surface analysis,
local
Fourier
series modelling,
trend
weighted
and mapping
Estimation
or each
species
to map,
possible
environmental
variables
such
instead,
synthetic
as species diversity,
or else the first few principal
axes from a principal
or a corre
components
it remains
averaging,
average, kriging, kernel estimators,
spline, moving
and
boundaries
interpolation
by drawing
(in
case the resulting maps may
which
be called
or 'tessellations').
'choropleth maps'
They have
been
reviewed
by several
authors,
including Tapia
& Thompson (1978), Ripley (1981, ch. 4), Lam

(1983), Bennett etal (1984), Burrough (1986,
ch. 8), Davis (1986) and Silverman (1986). Com
can provide
an estimate
of the
puter programs
at all points of the surface considered;
variable
the density
of reconstructed
is either
points
user
or
selected
set
the
the
by
program.
by
are used to draw maps
algorithms
Contouring
from the fine grid of interpolated
points.
Besides
linear interpolation
between
simple
closest
haps
trend surface analysis

is per
neighbours,
the oldest form of spatial interpolation
used
by ecologists
(Gittins 1968; Curtis & Bignal
It consists
in fitting to the data, by regres

1985).
a
sion,
polynomial
equation of the x and y coordi
nates of the sampling
localities. The order of the
is determined
by the user; increasing
increases
the number of parameters
to
a better-fitting
be fitted and so it produces
map,
with
the inconvenient
that these parameters
become more and more difficult to interpret eco
polynomial
the order
For
the commonly
instance,
of degree one is written:
logically.
equation
used
from
grid, one can map

of regular tiles
are
whose values
given by the points in the center
of the tiles. One should notice
that interpolated
in the case of a regular sampling
as a juxtaposition
surface
the
can only represent one variable

maps
are not multivariate,
thus these methods
at a time;
although
two or
in some cases to superpose
it is possible
seem
or
three maps. When
it does not
desirable
f = b0 + bxx + b2y (1)

where
variable
mapped),
z is the estimated
value of the response

z (the one that was measured
and is to be
the b's are the three regression
while
parameters.
is:
second-degree
polynomial
f = b0 + bxx + b2y + b3x2 + 64xy + b5y2

model
(2)
129
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polynomial.
of sugar-maples,
the estimated
frequencies
of
the fitted
(z), trend
provide also a
values
analysis programs usually

the varia
of residuals
(z
z), representing
left undescribed
map.
by the interpolated
of
the
order
the
6th
illustrates
12a
map
poly
Fig.
to the A. saccharum data. Com
nomial adjusted
map
tion
to Fig. 13 (kriged map)

pared
is still crude, although
obtained
the
contouring
28 parameters
have been adjusted. Fig. 12b is the map of regres
in A. sac
sion residuals,
showing the variations
not expressed
charum frequencies
by the trend
trend
: :
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45
l>
il
?!
j
?!
"
-?*
*"
"
an exam
(1987) presents
Burrough
of soil data.
surface analysis
a single
surface analysis
computes
Since
poly
surface,
regression
equation for the whole
the resulting map cannot have the precision
that,
more
local criteria can provide. For that reason,
to compute
in ecology mostly
it is used
and
remove
the
first
trends,
large-scale
using
degree
nomial
equation
in most
cases, prior to further spatial
on the residual
that can be conducted
analyses
values. Trends
can also be detected
methods
by autoregressive
Coull
Another
valid
1987).
and modelled
&
(e.g., Edwards
use of trend surface
is the predictive modelling

of spatial dis
analysis
tributions of organisms,
coordi
using geographic
use
one
can
nates alone as predictors;
other
or,
to build such a model,
variables
alone
predictive

130
.0
100.0
200.0
300.0
400.0
?00.0
?00.0
0.0
100.0
200.0
300.0
400.0
EO?.Q
600.0
r eOQ.O
0.0
Fig.
13.
100.0
(a) Map
200.0
of Acer
900-0
4OO-0
saccharum
500-0
obtained
600-0
0-0
by kriging,
and
(b) map
100 0
of the standard
200-0
deviations
900.0
400-0
500-0
of the estimations.
BOO.O
From
Fortin
(1985).
or
in conjunction
using multiple
with
geographic
or some
regression
coordinates,
other form of
modelling.
and
by mining
engineers
to
estimate
mineral
(1966)
a more detailed map
produces
Kriging,
developed
named
after Krige
resources,
usually
than ordinary
interpolation.
to trend
Contrary
analysis,
kriging uses a local estimator
located
that takes into account only data points
as well
in the vicinity of the point to be estimated,
surface
as the autocorrelation
structure
of the phenome
can be provided
either by
or
above),
by generalized
non; this information

the variogram
(see
intrinsic random functions
to make
of order k (Matheron
in
valid interpolation
1973) that allow

the case of non-stationary
variables
(Journel &
as fol
is
The
used
1978).
variogram
Huijbregts
lows during kriging: the kriging interpolation

method
estimates
other data points

the variogram
of
a point by considering
located in the observation
(given
by
the direction
all the
cone
and
window
them using
aperture angles), and weighs
the values read on the adjusted
theoretic vario
gram at the appropriate
distances;
furthermore,
this
among
kriging
splits
weight
neighbouring
so that the result does not depend upon
points,
the local density of points. Kriging programs pro
duce not only amap of resource estimates but also
one of the standard
deviations
of these esti
(David 1977; Journel & Huijbregts

this
map may help identify the regions
1978);
mations
where
should be intensified,
the map
sampling
being often obtained from amuch smaller number
of samples
than in Fig. 13.
The
nomena
seems
of mapping
multivariate
problem
phe
is all the more acute because cartography
an understanding
of the
to
for
instance
correlo
brought
by
light
could be done in the multi
analysis. What
essential
to reach
structures
gram
variate
case? How
could
one
combine
the varia
into a single,
bility of a large number of variables
and
understandable
Since
map?
simple

131
the
to compute
are available
programs
in this
of spatial
described
analysis
is not exhaustive.
list of programs
4. The
Table
following
methods
various
This
paper.
of
Package
Methods
BLUEPACK
Variogram, kriging.
CANOCO
spatial
Constained
analysis
GEOSTAT
Variogram,
kriging.
Kellogg's
Variogram,
kriging.
of constrained
developed by Lee (1981), by

Wartenberg (1985a, b) and by ter Braak (1986,
redundancy
this. They
1987) are other ways of accomplishing
of principal com
differ from the simple mapping
or correspondence
scores,
ponents
analysis
analysis.
Two-dimensional
data. The methods
ordination
canonical
analysis,
CORR2D
clustering,
of multivariate
ordinations:
correspondence
in some detail
explained
- and
can
be
that
above, produces groups
mapped
can be
indeed constrained
programs
clustering
to draw these maps directly (Fig. 10) - we
made
heuristic maps out
have here a way of producing
constrained
correlogram.
at the beginning of this section, in that

the spatial relationships
they take into account
resemble
them in that itmay
;
among samples
they
to draw several maps
be necessary
in order to
mentioned
NTSYS-PC
'R'
test.
Mantel
Simple
autocorrelation
(quantitative
Spatial
and nominal
data),
simple Mantel
cor
test, partial Mantel
tests, Mantel
con
with
spatial
relogram,
clustering
time
with
clustering
tiguity constraint,
A variety
of connecting
constraint.
networks.
SAAP
SASP Two-dimensional
special
variability
extracted
by all
axes. MacDonald
the
but orthogonal
&
important
Waters
of
(1988) give examples
palynological
Sur
maps obtained using Lee's Most Predictable
are
other
found in
face Analysis
(MPS);
examples
These methods
should
Wartenberg
(1985a,b).
find ample use among community
ecologists, who
multivariate
study essentially
(multi-species)
phe
(Moran's/
autocorrelograms
Spatial
and Geary's
c).
the
represent
analysis.
nomena.
SYMAPTrend
surface
inter
methods.
polation
UNIMAP
analysis;
other
Variogram,
methods.
kriging;
other
interpolation
Conclusion
- The BLUEPACK
package is available from: Centre de
et de morphologie
math?matique,
F-77305
Fontainebleau
Cedex,
g?ostatistique
Saint-Honor?,
- The
CANOCO
Braak,
for Applied
Wageningen,
The CORR2D
available
-
Road,
Country
The GEOSTAT
tems
The
New
B, Setauket,
Bldg.
is available
package
Qu?bec,
Kellogg's
M.
by Geoffrey
Biostatistics
Inc.,
Applied
4385
Inc.,
Canada
programs
H2J
rue
York
is
Jacquez
100 North
available
in PC version
Country
Road,
from: Geostat
Saint-Hubert,
from Applied
the Computer
Biostatistics
New
Inc.,
York
The
R package
for multivariate
Vaudor
(P. Legendre's
by Alain
data
lab.:
English
SAAP
and IBM
VAX,
microcomputers,
and French
versions.
speaking
is a set of FORTRAN
programs
package
from
Daniel
of
Wartenberg,
Department
and Community
Robert Wood
Medicine,
Medical
675 Hoes
School,
Lane,
Piscataway,
Johnson
New Jersey 08854, USA.

The
SASP
program
of
Statistics,
ment
100
11733,
analysis',
developed
see title page),
The
available
from
B, Setauket,
Sys
Suite
1,
from E. Renshaw,
of
University
Depart
Edinburgh,
King's
Kingdom.
- SYMAP is not distributed

any longer by Laboratory for
Computer
and Spatial Analysis,

Graphics
It is however
USA.
still available
centers.
puting
UNIMAP
is available
is
is available
Buildings, Mayfield Road, Edinburgh EH9 3JZ, United
versity,
USA.
-
for Macintosh
Environmental
2X1.
are available
Bldg.
available
11733, USA.
W.K.
Station,
Laboratory,
Kellogg
Biological
Michigan
State University,
Corners,
49060, USA.
Hickory
Michigan
The NTSYS
is
Rohlf,
package,
developed
by F.James
North
should
mainframes.
written
program
International
Montr?al,
-
from Cajo
J.F. ter
TNO
Institute
Group,
Box
AC
100, NL-6700
Mathematics
Science,
Computer
The Netherlands.
from
rue
is available
program
Agricultural
35
France.
stand? As we have seen,

ecologists
the physical
environment
is not homogeneous,
and most ecological
theories are based on precise
Where
tors A/S,
Norregade,
from: European
DK-2800
Lyngby,

Harvard
Uni
at many
com
Software
Denmark.
Contrac
132
about the spatial structure of popu
assumptions
lations and communities.
If we rely upon models
as many
that assume,
still do for simplicity,
that
biological
populations
or at random in space,
are distributed
uniformly
of obtaining valid
are small since the ecological
reality is
predictions
quite different.
chances
or hypothe
in the descriptive
a
of
research, ecologists who
sis-generating
phase
of organisms
should
sample spatial distributions
consider a priori that their data are structured
in
So,
space (i.e., are autocorrelated);

they should then
test for the presence
of spatial autocorrelation,
the spatial structure using maps and
and describe
In some cases, itmay
spatial structure functions.
to remove
be adequate
large-scale
spatial struc
or model-fitting
in order to
tures by regression
on residu
out
classical
statistical
carry
analyses
als, but in doing so, one must be careful
remove one of the important determinants
not
to
of the
is
since heterogeneity
In
the
ecosystems.
hypothesis-testing
functionalin
two
phase of a research, when
(model-testing)
linked by a causal
variables or groups of variables
one should
are
both
autocorrelated,
hypothesis
under
processes
study,
if significant,
test whether
could
their correlation,
be spurious and due to a similar spatial structure
in both. This in turn could give clues as
present
to the identity of some other spatially autocorre
lated causal
their
common
variable
that may
autocorrelated
have
given
structure.
them
a
In
is the rule rather

where
spatial structuring
can prevent
the
this
than
exception,
precaution
one from reaching unwarranted
conclusions.
world
of spatial analysis (descrip

Statistical methods
are currently under develop
tive or inferential)
ment, and already they offer a way of answering
about populations
and
relevant questions
many
of the
1): demonstration
or
de
structures,
spatial
temporal
or multi
of these structures, univariate
communities
existence
(Table
of
scription
with models,
variate mapping,
analy
comparison
on
structures
of spatial
sis of the influence
statisti
assumed
causal links between
variables,
the inde
which
do not assume
cal analyses
available
of the observations.
Programs
pendence
for spatial analysis are becoming widely available.
Some are listed in Table 4; this list is not exhaus
tive.
We can expect the spatial approach

to ecologi
to bring about a quantic
cal problems
jump for
and
who
had
ecologists
population
geneticists
learned a type of statistics where one had to hide
It is now possible
to
space or time structures.
use these structures and to integrate them into our
as fully-fledged
analyses
controlled
variables.
Acknowledgements
This
is publication
No. 339 from the Groupe
des Eaux
Universit?
de
douces,
d'?cologie
and contribution
No.
689 in Ecology
Montr?al,
and Evolution
from the State University

of New
at Stony Brook. We are indebted to Pierre
at Universit?
Ph.D.
student
de
Drapeau,
York
who
directed
the sampling
Montr?al,
program
ex
that produced
the data used for the various
this paper. Dr Michel
that illustrate
amples
de Montr?al,
?cole
David,
gave us
Polytechnique
to his GEOSTAT
for and access
instructions
that we used
for kriging.
package,
M.
State
of New
Geoffrey
Jacquez,
University
on
York
at Stony Brook,
revised
the sections
computer
two-dimensional
analysis. Alain Vaudor,

some of the pro
computer
analyst,
developed
for Multivariate
Data
grams of 'The R Package
are
and
We
for
the
present study.
Analysis'
during
to
Dr
R.
State
Robert
Sokal,
University
grateful
of New York at Stony Brook, who gave us access
to Dr Neal
spectral
L. Oden's
two-dimensional
correlo
and provided
funds to
computing
are
4
this
of
We
also
indebted
paper.
produce Fig.
to Dr E. David Ford and Dr Cajo J.F. ter Braak
gram program
for very helpful
comments.
ported by NSERC
Legendre,
Fortin.
This
grant No.
and by a NSERC
study was
A7738
scholarship
sup
to P.
to M.-J.
Appendix 1
Formulas
Spatial
and technical
autocorrelation
points
analysis
is no spatial
there
autocorrelation.
The values
of the
H0:
are spatially
variable
of the /
Each
value
independent.
l
is equal to E(7) =
coefficient
is
1)~ ? 0, where
(n
E(7)

133
of data
of I and n is the number
the expectation
= 1.
of the c coefficient
equals E(c)
spatial autocorrelation.
H}: there is significant
each
points;
of
are
the variable
=
E(7)
ferent
1(d)
E(c)
l)-1
=
[?I I
the /
y)(yj
?v<Jt
Z (y,
yjfVW
along
(y,
y)]/[W?
y)2]
(1)
y)2]
(2)
class d.
for each distance
computed
are for
are the /s. All summations
but
of data points,
from 1 to ?, the number
/ and j varying
1when
take the value
i = j. The
the cases where
exclude
w?Js
are
of the variable
values
the pair (ij) pertains to distance class d (the one for which
the coefficient
of
the
whole
Wy's,
square
is the sum
W
is computed),
and 0 otherwise.
of pairs
the number
in other words
(in the
into
taken
of distances
matrix
among
points)
or
for the given dis

the coefficients
computing
1 to
from
varies generally
coefficient
tance class. Moran's
or
+1
can
1
it
exceed
(Fig. Id, h, k);
1, but sometimes
auto
to positive
/ correspond
of Moran's
values
positive
coefficient
varies from 0 to some indeter
correlation.
Geary's
account
when
3 in real cases;
rarely exceeds
auto
to positive
1 correspond
can be tested for significance;
coefficients
These
correlation.
error of the estimated
the standard
for computing
formulas
minate
values
value
positive
of c smaller
which
than
for nominal
coefficient
autocorrelation
(qualitative)
data is described by Cliff & Ord (1981) and by Sokal & Oden
(1978).
Technical
-
points:
autocorrelation
Spatial
than ca. 30
with
fewer
not
should
because
be performed
of
the number
class then
in each distance
pairs of localities
results.
small to produce
significant
are two ways
of dividing
distances
There
or
either by forming
classes,
equal distance
makes
This
last
solution
equal frequencies.
valid
compute
the correlogram
analysis
localities,
even
coefficients
too
becomes
a data
classes:
absence
would
autocorrelation
to
it possible
right-hand
equal distance
reflect
the
fact
for that variable,
that
which
then
the
part of
classes
the
zeros,
because
be
overestimated
localities
is not what
share
the best way
between
of expressing
pairs
of localities
geographic
spectral
topological
of
network
networks,
(see: Connecting
in terms of number
of edges
distances
analysis
The
autocorrelation
spatial
values
autocorrelation
sample
matrix
contains
rgh, corresponding
of
pairs
to all possi
all
ble lags (g, h) where g is the lag along the x geographic axis
of sampling
is the ratio
and
of the
variance
sample
as
is computed
lag along the y axis. Each value

rgh
at lag (g, h) to the
autocovariance
sample
The
autocovariance
of the
sample
sgh
ytJs.
h is the
m-g
=
sgh (i/mn) X
!,&?,?-yKyu+8,J+H)-y)
(3)
- n <
0 < g < m and
h < n; m and n are respectively
rows
of the geographic
of
and columns
sampling
over j = 1,...,?The
second
summation
is
taken
h if
grid.
n if h < 0. There
h > 0 and over j = \h\ + 1,...,
is no need
to compute
the whole
autocorrelation
surface
( m < g < m)
since the surface
is a reverse
of
itself
round
either of
image
where
the number
The
lag axes.
Schuster
periodogram
can also be computed,

again
of lags (g, h). The periodogram
of the spatial pattern
than the
and power
Periodograms
correlogram.
for all possible

combinations
is a more
compact
description
full two-dimensional
are often
as functions
of frequencies
expressed
=
of periods
For
convenience,
(frequency
1/period).
are here multiplied
frequencies
by the size of the series
the whole
that occupies
(m or n) so that a wave
length
spectra
instead
area has a frequency

(m or n) of a side of the sampling
is then
considered
range of frequencies
(p or q) of 1. The
=
=
p
0,...,
(
/i/2),...,
1) where
(m/2) and q
((/i/2)
(ra/2)
is intended
may
relationships
in
not be
when
travel
the
of the sine wave
under
x and y
in directions
window)
the direction
of
sign of q gives
As
in
time
series
study.
analysis,
of the periodogram
/(/?, #), for each frequency
measures
the amount
of variance
of variable
y
intensity
combination,
that
is explained
by the given
to the data,
(p, q), after fitting
series (sum of sines and cosines)
combination
of frequencies
a Fourier
by least squares,
the given combination
with
of frequencies. See formulas inRenshaw & Ford (1984), for

sional
as a three-dimen
is presented
periodogram
with
the axes
of the
(/?, q) along
frequencies
and
the intensity
of the periodogram
plane,
the response
variable.
The
instance.
their
most
applications.
distances
Euclidean
localities
in the observation
frequency
of the sampling
surface. The
with
classes
in the
a lot of double
set that contains
some
instead
and (h/2) are respectively the Nyquist frequencies (highest

into
1983); with
(Sokal
too
on the contrary,
the number
of pairs of points becomes
classes
in the large distance
small for valid testing
(Fig. lc).
cannot be performed
with
autocorrelation
analysis
Spatial
of
degree
and would
use
between
Two-dimensional
the zero
statistics can be found inCliff & Ord (1981), Sokal & Oden
(1978) and Legendre & Legendre (1984a). A special form of
spatial
can
and compute
this network.
below)
wv(y,
coefficients
These
one
less,
connections
1.
c{d) =[(?-1)11
The
from
dif
one could use

\?d or
1917), or some other
& Gates
(Estabrook
1984).
is felt to be meaning
distance
Instead,
et at.
Jumars
1967;
(Mantel
transformation
appropriate
the Euclidean
In cases where
values
of
c is significantly
of
the value
0;
The
value
different
significantly
-(?
from
dependent.
spatially
is
coefficient
The
l/d2
data.
ecological
analysing
value
plot,
controlling
/(/?, q) as
The polar
frequencies
spectrum
and angular
of
aims at measuring
wave
of the dominant
the data
directions

the
pat
134
terns
in the data.
2 graphs,
present
are produced.
& Ford
(1983),
measures
the /?-spectrum,
the frequencies
Renshaw
first
The
by
proposed
first one, called
of the waves
The
pattern.
spatial
in the periodogram
that have
e = t2Xi-\plq)
same
the
approximately
values
R and 0
are first
of classes
along the abscissa

(that is, the various
into a manageable
divided
number
are drawn.
the graphs
The
value
have been
I(p, q) values
of 1, so that a data set with
an ?-spectrum
produce
to 1. Since the individual
are approximately
for
be
tested
and
no
a 0-spectrum
values
significance
In the same
mentioned
values
shows
that the spectra
analysis
ments
for interpreting
the spatial
of a descriptive
has more
value.
of two-dimensional
are
the most
useful
|d \/a)]
C0; a is the range.
C0 + C
a,
[1
exp(-d2/a2)].
as distance
increases
of a vario
two-thirds
when
the
describing
structure.
spatial
With
ecological
the data often
close
spectral
instru
is rare and
property
'drift' in
called
the stationarity
some overall
data,
contain
trend,
the kriging jargon; drift can affect significantly

the accuracy
of kriging.
Thus
in the presence
of non-stationarity
and
intrinsic
functions
of
random
drift, the use of 'generalized
order
mate
instead
the autocorrelation
structure.
of
Graphs
describe
interrelations
autocorrelation
clustering,
spatial
methods
that
information
require
other
connect
are used
to
among
points
for such data analysis methods
interconnections
spatial
constrained
as
to esti
networks
Connecting
and
of a variogram,
k' is recommended,
In the case
localities.
neighbouring
of sampling
the periodogram
structure;
y(d)
in the computation
decreases
the first
lc). Thus,
only about
should be taken
into account
(Fig.
way,
use
Actual
sill
are computed
in correlograms,
for distance
variograms
which
of pairs of points
that the number
implies
used
and 0-spectra
k individual
authors.
C0 + C [1 exp(
points:
gram
with
and C =
classes,
q) in the periodogram
of/(/?,
as
then they can
(\00/mn)x22),
a critical
value
of
against
distributed
model:
Technical
values)
before
values
in the
particular
to intervals
that correspond
of /, can be tested
values
for
say,
containing,
a critical value of / = [l/(2k)]x2a
significance
against
2ky As
one should
in all cases
of multiple
the usual
testing,
apply
Bonferroni
correction
level
and use the corrected
significance
v is the number
a' = ol/v where
of tests performed
simultane
this point
had not been
ously;
by the above
emphasized
(100/m?)xfa,2)of the Rgraphs
Gaussian
the
an average
structure
should
spatial
y(d)
effect
=
Sphericalmodel:y(</)
C0 + C [(3<//2a) (?3/2?3)]if?<
while y(</) = C0 + C if </> a.
to have
scaled
C0
- As
angle
In these graphs,
(O?<0<18O?).
is the nugget
form
of the
is a graph
^-spectrum
ing
in the periodo
average
response
I(p, q) of all the elements
the same frequency magnitude
gram that have approximately
+ q2). The
R =
second
called
the 0-spectrum,
one,
*J(p2
measures
the directions
It is presented
of the waves.
(angles)
as a graph of the average
of
the elements
all
response
I(p, q)
the
model:
Exponential
where
the
locations,
each
('rook's move'),
he so chooses.
analysis,
about
of a regular
square grid
is simple,
since one can
solution
to its neighbours
point
or else in all 8 directions
in all
4 directions
if
('queen's move')
design has the form
of
If the regular
sampling
connections
rows, as in Fig. 2 for instance,
(also
staggered
in all
'link edges') may be established
called
with neighbours
are irregular
If the sampling
tiles that
6 directions.
localities
Variogram
The
called
the vario
experimental
(often
semi-variogram
as a function
is a plot of the values
of semi-variance
gram)
of distance.
The estimator
of the semi-variance
is
function
touch
one
a natural
another
choice
and cover
the whole
is to connect
localities
surface
under
study,
a border
in
that have
common.
It often
y{i))2] (4)
y{d)=[\l{2nd)]YJ[y^d)-
not
at distance
located
of pairs of points
nd is the number
The
is for i varying
from
d from one another.
summation
c autocorrelation
ference
Some
are
the
is a distance-type
in the denominator
lies mainly
of the most
following
often
(Fig.
used
8). Other
theoretic
models
coefficient
function;
of
(above),
the dif
the function.
models
variogram
are proposed
by
Journel & Huijbregts (1978).

Linear model: y(d) = C0 + bdwhere b is the slope and C0 is
the
intercept
(nugget
effect).
happens,
a regular
however,
that
In such
pattern.
the
localities
do
sampling
one should wonder
cases,
not provide
if the ecological
under
study would
problem
are. If
the close neighbours
what
way of deciding
no such ecological
can be found,
then one can rely
criterion
on the more
criteria. The most
commonly
arbitrary
geometric
first
where
to nd. Just like Geary's

this structure
function
form
a natural
used
graph-theoretic
criteria
are the minimum
tree
spanning
(Gower & Ross 1969), the Gabriel graph (Gabriel & Sokal
1969),
or
the Delaunay
triangulation
which
is
simply
an
algorithmic method of dividing a plane into triangles that

obey some precise set of rules (Miles 1970; Ripley 1981;
to note
that
It is interesting
1981).
a
tree
Gabriel
is
of
the
subset
graph,
spanning
is a subset of the Delaunay
triangulation.
Watson

the minimum
which
in turn
135
Appendix 2
to the reference
test
of the Mantel
Theory
value
of
obtained
and Y),
sidered
Hypotheses
in matrix
X are
among
points
H0: Distances
to the corresponding
in matrix
related
distances
as
distances,
represents
geographic
H0 reads
(or the multivariate
variable
data)
in X
is not
the
follows:
Remarks
autocorrelated
z or
r statistics
the
in matrix
X
points
distances
in matrix
Y.
are correlated
to
to a table
referring
test gives
(1967) statistic:
Normalized
Mantel
ponent
of
distance
matrices.
the
is the number
column
and
of the standard
normal
indices.
into
another
distribution.
linearity,
tance matrices,
the
Strong
of
and
[(x? x)lsx] [(yg y)/sy]
between
relationship
non-linearity
themselves
expressing
to
(1983)
correlation
formula.
parametric
statistic:
r = [l/(n - 1)]
I iI
for i^ 7, where
row
j are
/ and
transformed
This
a good
from
relationships
this led Dietz
test;
/ 7? y, where
be
(1967), which can be tested by
of the probability
when
the
approximation
number
of objects
is large.
Like Pearson's
correlation
the Mantel
statistic
coefficient,
formula
is a linear model,
out the linear com
that brings
Statistics
can
statistic, called t by Mantel

among
under
is one
linearly
Y
Y. When
The
H ^Distances
the corresponding
for
obtained
statistic
not
as a gradient.
- Mantel
H0. If the actual

to have
been
likely
the null hypothesis
under
between
X
(no relation
to be con
then H0 is accepted;
if it is too extreme
a likely result under H0,
then H0 is rejected.
distribution
the Mantel
not
influence
or both
in one
transformations
has
through
the use
suggest
The
yet been
fully
two
in the
values
can probably
prevent
the Mantel
of
a non
of
lack
of
of the dis
investigated.
(6)
i and j are row and column

and n
indices,
of distances
in one of the matrices
(diagonal
excluded).
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Spatial Pattern and Ecological Analysis

Author(s): Pierre Legendre and Marie-Jose Fortin

This content downloaded from 148.206.159.132 on Thu, 14 Nov 2013 22:47:52 PM

role inmany ecological

This content downloaded from 148.206.159.132 on Thu, 14 Nov 2013 22:47:52 PM

1911 \Allen & Starr 1982; Legendre

1974; Hassell & May

the spatial variability

for the presence

(Legendre & Demers

4) Objectives: Estimation (interpolation) and mapping.

for a single variable:

for a single variable;

This content downloaded from 148.206.159.132 on Thu, 14 Nov 2013 22:47:52 PM

anywhere near the first, this cannot be seen as a

perience with for the analysis of the spatial struc

are also known,

in spatial analysis will be presented,

ble. The methods

are the basic

try to show that the methods

away. The spatial structures we

gram and a semi-variogram

is not the same for all distance

sampling points (Table 2). It can

This content downloaded from 148.206.159.132 on Thu, 14 Nov 2013 22:47:52 PM

tion is on the 'liberal' side (Bivand 1980; Cliff &

is at least partially known

proposed by Cliff & Ord (1981), the analysis of

among regularly spaced plants

step (interval) is too large compared

This is why it is important to take the lack of

(that results from the

the value of the

two decades (Orl?ci 1978;Gauch 1982; Legendre

This content downloaded from 148.206.159.132 on Thu, 14 Nov 2013 22:47:52 PM

Lee (1981),Wartenberg (1985a,b) and ter Braak

to study itmore thoroughly.

points, and each point is considered

Let us first study one variable at a time. If the map

for the presence

testing for the presence of a spatial struc

that have been introduced by point pattern

1982;Carpenter & Chaney 1983) and need not be

ing Pielou (1977), Getis & Boots (1978), Cic?ri

In the case of quantitative

(1950) orGeary's c (1954) spatial autocorrelation

class. This coefficient

since the numerator

This content downloaded from 148.206.159.132 on Thu, 14 Nov 2013 22:47:52 PM

variables (Cliff& Ord 1981); they have been used

plants (Sakai & Oden 1983; Sokal & Thomson

gram (below), Goodall's

already familiar with the use of cor

are analysed mostly

of this test is that

level a. The global test

at least one value which

(Cooper 1968; Miller

in time series analysis

(Box & Jenkins 1970). Simulations in

1984 paper show that the power of Oden's