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Integrating spatial multi-criteria evaluation and expert knowledge

for GIS-based habitat suitability modelling


Ron Store
*
, Jyrki Kangas
Finnish Forest Research Institute, Kannus Research Station, P.O. Box 44, Fin-69101 Kannus, Finland
Received 4 April 2000; received in revised form 23 October 2000; accepted 31 January 2001
Abstract
GIS data processing and spatial analysis, together with modern decision analysis techniques, were used in this study to
improve habitat suitability evaluation over large areas. Both empirical evaluation models and models based on expert
knowledge can be applied in this approach. The habitat requirements of species were described as map layers within GIS so
that each map layer represented one criterion. GIS was used as the platform in managing, combining and displaying the
criterion data and also as a tool for producing new data, especially by utilising spatial analysis functions.
Criterion standardisation, weighting and combining were accomplished by means of multi-criteria evaluation (MCE)
methods, the theoretical background being based on the multi-attribute utility theory (MAUT). By using continuous priority
and sub-priority functions in the evaluation, no classication of continuous attributes was needed and also non-linear
relationships between habitat suitability and the attributes could be considered. Sensitivity analysis was applied to consider the
temporal factor in the analysis and to nd out the effect of different criteria weights on the spatial pattern of the suitability
index. Changing the weights of permanent and time-changeable habitat factors shifted the location of optimal habitats for the
species. In the long run, permanent factors such as soil properties dene the habitat potential, which is important to take into
consideration; e.g. in forest management planning and species conservation. The method is illustrated by a case study in which
habitat suitability maps were produced for an old-forest polypore, Skeletocutis odora. #2001 Elsevier Science B.V. All rights
reserved.
Keywords: Habitat suitability modelling; Multi-criteria evaluation; Geographical information systems; Expert knowledge; Sensitivity
analysis; Skeletocutis odora
1. Introduction
Nowadays, the objectives set for forest use are more
diverse than in the past. Values such as recreation,
landscape beauty and conservation of biodiversity
have gained prominence alongside traditional wood-
production values. In wildlife management, protection
of critical habitats and conservation of endangered
species are the foremost tasks from the point of view
of biodiversity preservation. When performing these
tasks, it is important to know the factors affecting
habitat suitability and habitat selection, and also to
have methods for determining the suitability of an area
for certain species.
Habitat suitability modelling is often used to
produce probability maps depicting the likelihood
of occurrence of certain species and to nd out
Landscape and Urban Planning 55 (2001) 7993
*
Corresponding author. Tel.: 358-6-8743211;
fax: 358-6-8743201.
E-mail address: ron.store@metla.fi (R. Store).
0169-2046/01/$20.00 # 2001 Elsevier Science B.V. All rights reserved.
PII: S0 1 6 9 - 2 0 4 6 ( 0 1 ) 0 0 1 2 0 - 7
the landscape properties of preferred habitats (e.g.
Johnson and Temple, 1986; Pausas et al., 1995).
These models are typically made by exploring statis-
tically the relationship between existing occurrences
of the species and the site properties. However, sta-
tistical evaluation models must be based on adequate
empirical study material, which is usually expensive
and time-consuming to collect, especially in the case
of rare species. Furthermore, the number of species we
are interested in is continuously increasing. Thus,
empirical evaluation models for all the species of
interest cannot be expected to be available.
One possibility for dealing with this problem is to
use expert knowledge when models based on objective
information are not available. In this approach, habitat
suitability models, for example, are estimated on the
basis of expert judgements instead of empirical mea-
surement data. In order to enable the use of ecological
expert knowledge in habitat suitability modelling,
methods for transforming expert knowledge into a
numerical form, as well as appropriate tools for hand-
ling and producing data about environmental condi-
tions, are needed.
GIS applications have frequently been used in
producing new information by both combining infor-
mation from different sources and by spatial analysis
of existing data. Spatial modelling, especially carto-
graphic modelling, has been applied when looking
for areas suitable for a specic forest use or for
some species of interest (e.g. Shaw and Atkinson,
1988; Reisinger and Kennedy, 1990; Wadge et al.,
1993). Usually the objective in applications involving
cartographic modelling is to locate the area or areas
where the given criteria apply. In a classication
based on Boolean logic, an area is either accepted
or rejected based on a given threshold value. The
nal outcome of these applications is a map depict-
ing areas simultaneously fullling all the conditions
set.
However, problems have been noted with methods
for site selection and resource evaluation that rely on
classical Boolean logic (Carver, 1991). In situations
where the threshold value is not precise, loss of
information or error propagation may occur. Further-
more, the method does not offer any analytical pos-
sibility for examining which of the areas fullling the
criteria are the most appropriate for the purpose in
question or which areas are the best beyond the
feasible areas. Because of problems with Boolean
overlay, multi-criteria evaluation (MCE) methods
have been applied instead of Boolean logic (Carver,
1991; Pereira and Duckstein, 1993; Jankowski, 1995).
The main aim in using MCE methods is to provide a
basis for evaluating a number of alternative choice
possibilities on the basis of multiple criteria (Nijkamp
et al., 1990).
Besides the problems associated with the use of
Boolean logic, the accuracy of results depends on the
quality of the source data and on the quality of the
spatial data analysis used in the process. According to
Burrough and McDonnell (1998), the most important
factors affecting the quality of spatial data are cur-
rency, completeness, consistency, accessibility, accu-
racy and precision, and various error sources. Errors in
the nal results may originate from any stage of the
process, from the collection of the source data to the
interpretation of the nal results. Data quality is often
described by thematic accuracy, positional accuracy
and temporal accuracy. Also error propagation, mean-
ing the accumulation of errors from various sources,
affects the results of analyses.
One approach to MCE in a GIS environment is the
additive technique whereby the criterion scores are
standardised and the total score for each alternative is
calculated by multiplying each criterion score by its
weight factor and then adding the results. Weighted
linear summation is probably the best-known example
in this category (e.g. Berry, 1993). Additive techni-
ques are based on the multi-attribute utility theory
(MAUT), where the weights are interpreted in the
framework of a linear priority (utility) function.
Siddiqui et al. (1996) presented an additive
approach to a spatial problem based on the analytic
hierarchy process (AHP). AHP is one method of
producing the criteria weights and also of making
criteria of different kinds commensurable (Saaty,
1980). Siddiqui et al. (1996) called their method
spatial-AHP and applied it to supporting landll sit-
ing. It provides a way to exclude areas which are not
suitable for a certain purpose and to rank the remain-
ing areas based on area attributes. However, in this
method, attribute values have to be classied using a
limited number of classes. The problem in doing so is
that the classication may lead to loss of information
and, on the other hand, dening the threshold values of
classes may increase uncertainties in situations where
80 R. Store, J. Kangas / Landscape and Urban Planning 55 (2001) 7993
there is no exact threshold value or the value is not
known with certainty.
Decision criteria, used in habitat suitability eva-
luation, are mainly continuous ones. This is why
classication of the data describing the alternatives
may lead to increasing uncertainty and loss of infor-
mation. This problem can be avoided by utilising
continuous priority functions in the evaluation. Then,
no classication is needed in the evaluation of habi-
tats. In the present study, the evaluation technique
utilised in the HERO heuristic optimisation (Pukkala
and Kangas, 1993) is applied. It enables the evalua-
tion of habitats with respect to continuous attributes
by applying so-called sub-priority functions without
any classication of attribute values. Also, non-linear
relationships between habitat suitability and the
attributes can be considered using the sub-priority
functions.
Suitability indices have been utilised, for example,
in various aspects of natural resources planning (e.g.
Pukkala et al., 1995). In forest management planning,
decision alternatives are considered in a given area on
the basis of their consequences within the planning
horizon. For evaluating the alternatives, they are
assessed with respect to each objective set for the
forest and its use. Planning always concerns the future.
The objectives, in turn, can be related to the near
future or to longer time horizons. When a suitability
index is used in planning, it is, therefore, useful to
incorporate the temporal factor in the estimation of the
index values. In this study, a technique based on
sensitivity analysis is presented for taking the tem-
poral factor into consideration when producing habitat
suitability indices.
The aim of the present study is to improve habitat
suitability evaluation by using GIS-based data proces-
sing and spatial analysis along with state-of-the-art
decision-analysis techniques. Important qualities of
the habitat-evaluation method aimed at in this study
are the abilities to utilise expert knowledge when
evaluation models based on empirical studies are
not available, and to apply continuous attributes in
the evaluation in a reasonable way within a GIS
framework. The method is illustrated by a case study
in which a habitat-suitability map is produced for an
old-forest polypore, Skeletocutis odora. The approach
presented is appropriate for use with other species as
well.
2. Methods
2.1. The general outline
All species have specic habitat requirements,
which can be described by habitat factors. These
factors are connected to the critical characteristics
of the habitat, e.g. to those of vegetation or soil, but
also areas surrounding the habitat can inuence the
habitat quality (e.g., spatial structure of landscape
elements). Habitat factors can also be classied
according to the deterministity of the factor: a deter-
ministic habitat factor has to be present in a high-
quality habitat, but a non-deterministic factor has a
trade-off with some other factor. So, a deterministic
factor can be taken as a non-compensatory habitat
characteristic whereas a decrease in a non-determi-
nistic factor can be compensated by an increase in the
value of another non-deterministic factor, as expressed
in the habitat-evaluation model.
Habitat suitability is determined by habitat
factors. Habitat suitability can be measured by a
habitat suitability index, which is a unitless variable
describing the priority of the habitat with respect to
the needs of the species (or group of species) under
consideration. Typically, it can get values between
0 and 1, and is estimated on grounds of the measur-
able habitat characteristics. For producing habitat
suitability indices for large areas, methods enabling
the management and analysis of large amounts of
data are needed as well as the calculation para-
meters describing the most essential habitat character-
istics.
The habitat suitability modelling method applied in
this study consists of two basic phases. Firstly, deter-
ministic habitat factors are used to nd the potential
habitats fullling the absolute prerequisites for the
species in concern within the area. The result of this is
a feasible area meeting the absolutely necessary con-
ditions for the occurrence of the species. Secondly,
non-deterministic factors are used to evaluate the
potential habitats.
The evaluation procedure consists of the following
steps:
1. The assessment of a suitability structure: choosing
the habitat factors and determining their impor-
tance and how they affect the habitat priority.
R. Store, J. Kangas / Landscape and Urban Planning 55 (2001) 7993 81
2. Producing map layers: raw data acquisition and
transforming to appropriate GIS layers.
3. Cartographic modelling: dening the feasible area
and combining the habitat factors.
4. Sensitivity analysis: demonstrating the effect of
different criterion weights on the spatial pattern of
the suitability index.
Step (1) is carried out using an external MCE
program which works outside a GIS while steps
(2)(4) are executed under a raster GIS.
2.2. Assessment of the suitability structure
The rst step in assessing the suitability structure
is to determine the habitat factors on the basis of
an analysis of existing studies and knowledge. Here,
judgements made by experts on ecology can be
applied. The next step is to classify the decision
factors into deterministic and non-deterministic ones
according to their exclusive character. Deterministic
habitat factors are Boolean type of variables, which
are used like geographical constrains to eliminate
areas from further consideration on the basis of certain
attribute values. In practice, very few habitat factors
can be considered so straightforwardly that a simple
Boolean variable is appropriate for describing it.
Next, the mutual importance of non-deterministic
habitat factors have to be evaluated. This can be
done by a number of methods. In this study, the
HERO method (Pukkala and Kangas, 1993) was
adopted. By its very nature, HERO is a heuristic
optimisation method including both the estimation
of the priority model and the procedure for maximis-
ing the value of the model among the decision alter-
natives. It was originally developed for the purposes of
tactical forest planning when dealing with a huge
number of alternative forest plans, each consisting
of a combination of treatment schedules for the forest
stands within the area under planning. In this study,
only the priority model estimation option of HERO is
applied.
Instead of forestry objectives, the priority model
here is estimated for habitat-evaluation purposes only.
Thus, it includes habitat factors as variables. The
resulting model can be utilised directly in forestry
planning calculations. Combining the habitat factors is
based on the multi-attribute utility theory (MAUT)
and accomplished by an additive priority function P:
P
X
m
i1
a
i
p
i
q
i
(1)
where P is the global priority, i.e. the habitat suitability
index, m the number of factors, a
i
the relative impor-
tance of factor i (
P
a
i
1), p
i
the sub-priority func-
tion of factor i, the maximum value for each factor is
one, and q
i
the amount of factor i.
The relative importances of the habitat factors are
used as coefcients a
i
in the priority function. The
relative priority of the habitats with respect to each
factor is estimated on a ratio scale with a sub-priority
function. The sub-priority function depicts the change
in habitat suitability as a function of the factor. The
sub-priority functions scale all factors between 0 and
1, thereby making the objectives commensurable (the
best priority value is always 1, other attribute values
getting sub-priorities relative to it). Sub-priority func-
tions are developed separately for each habitat factor
and the total suitability index is the weighted sum of
the sub-priorities.
There are several alternative methods for estimating
the coefcients a
i
. In this study, the coefcients were
solved applying pairwise comparisons carried out by
an expert on ecology. The relative importances of the
factors are computed using the eigenvalue method of
ratio scale estimation (Saaty, 1977). The importances
of the factors are compared pairwise using a graphical
interface instead of the verbal scale as proposed by
Saaty (1980). The relative importances of two factors
at a time are dened by adjusting the lengths of the
horizontal bars on the computer screen.
The method enables the presentation of a habitat
factor in a hierarchical manner. A factor may be
described by means of a model whose variables are
the components describing the factor in more detail
and whose coefcients are the components' relative
importance. This being the case, the sub-priority
functions are dened to depict the impact of the
detailed components on the priority obtained through
the habitat factors they explain.
When estimating the sub-priority function, the
maximum and minimum values to be considered in
the evaluation are determined rst and displayed by
the user interface. In addition, a few intermediate
values are selected. The desirability of these values
82 R. Store, J. Kangas / Landscape and Urban Planning 55 (2001) 7993
is then estimated by means of pairwise comparisons
and the values are given relative priorities dening the
sub-priority function (Fig. 1). The sub-priorities in-
between the compared values are solved by linear
interpolation. Although a piece-wise linear function is
applied, a sub-priority function can show a non-linear
relationships between habitat suitability and the envir-
onmental variables. The sub-priority function is esti-
mated separately for each habitat factor, or for its
components. Generally taken, the estimation can be
based equally on expertise or subjective value infor-
mation. The ratios of the values of objectively mea-
surable variables can also be used.
2.3. Producing map layers
A GIS application is used for managing, producing,
analysing and combining spatial data. Some of the
attribute data needed in the suitability-evaluation pro-
cess are collected by means of eld inventories and the
rest are produced from collected or existing data by
using different kinds of spatial functions and analysis.
For example, soil moisture is described by the topo-
graphic wetness index T
i
. It is a function of the
upstream contributing area and the slope of the land-
scape. It is calculated by means of formula (2) (used,
e.g. in the TopModel program, Beven et al., 1995). The
index can be interpreted as the spatial distribution of
soil moisture or as the relative depth of water table.
When the catchment area increases and gradient
decreases, the topographic index and soil moisture
content increase.
T
i
ln
a
tan B
(2)
where a is the area draining through a grid square, and
tan B the average outow gradient from the square.
The data describing the habitat factors are rasterised
into 25 m 25 m grid cells and every habitat factor is
stored in its own map layer.
2.4. Cartographic modelling
Cartographic modelling is applied in producing and
combining spatial data describing the habitat factors.
Cartographic modelling is used as a means to deter-
mine the feasible area, standardisation and transform-
ing raw scores for priority measurements, and nally
to calculate a suitability index as a combination of
single habitat factors. In the rst phase, the feasible
area is produced by numerically overlaying a map
layer describing the study area and all the map layers
considering the deterministic habitat factors. This
overlay is carried out as a Boolean overlay, where
areas not included in the area determined as being
suitable by all the deterministic habitat factors are
excluded from further consideration. The remaining
areas establish the feasible area.
The map layers considering the attribute data used
in the MCE process may differ from each other as
regards the measurement units. To make the raw
scores commensurable, some kind of standardisation
is needed. Voogd (1983) presented various kinds of
standardisation methods to transform raw scores into
scores with ranges of variation 0 to 1. The problem
with the most commonly used standardisation meth-
ods is that they merely perform a linear transformation
between the raw score and the standardised score. The
linearity approach does not enable meaningful com-
bination of standardised criteria scores in situations
where the best value is either not the greatest value or
the smallest value, or where the relationship between
the raw score and priority is not linear.
To avoid those problems, the priority model as
applied in HERO is used as a standardisation tool
and for transforming the standardised scores into
Fig. 1. An example of sub-priority functions indicating a non-
linear relationship between habitat suitability and a habitat factor.
R. Store, J. Kangas / Landscape and Urban Planning 55 (2001) 7993 83
priorities. In the priority model, the differences in
measurement scales and units are not a problem,
because the method is based on direct comparisons
between signicance or priority of the elements in the
model. Non-linearities between raw scores and prior-
ity are handled by piece-wisely linear sub-priority
functions.
In order to technically enable the calculation of sub-
priorities and global priority (i.e. the habitat suitability
index) within the GIS application, where each habitat
factor is its own map layer and the data is rasterised
into grids, the following procedure is performed. First,
the raw scores in the map layers, describing the non-
deterministic habitat factors, are divided into groups
according to the ``corner points'' of the sub-priority
functions (see Fig. 1). The scores between two ``corner
points'', i.e. between the ones compared in the esti-
mation of the corresponding sub-priority function,
belong to the same group. Thus, the sub-priority
values for them can be calculated by the same inter-
polation formula. A separate interpolation formula is
constructed for each `piece' of a piece-wisely linear
function. Then the scores are dealt with group by
group to compute the sub-priority produced by a
certain value of a habitat factor. The sub-priority value
is estimated for each cell this way, resulting in so-
called sub-priority grids. Finally, the sub-priority grids
are combined by multiplying each sub-priority grid by
its weight coefcient a
i
and by summing the weighted
sub-priority grids by the means of arithmetical overlay
analysis. The end result is the habitat suitability index
for each 25 m 25 m cell.
2.5. Sensitivity analysis
Various weighting schemes are applied to the main
criteria when using sensitivity analysis. The main
purpose here is to examine how sensitive the choices
are to the changes in criteria weights. This is useful in
situations such as where uncertainties exist in the
denition of the importance of different habitat fac-
tors. In many cases, it is important also to know how
the results will change if the weights are changed.
If there are requirements in existence for a certain
kind of sensitivity analysis, these can be taken into
consideration already at the stage of building the
decision hierarchy. For example, criteria based on
facts can be classied under one main criterion
and those based on changeable preferences under
another. Then it is possible to change the weights
between the facts and the preference criteria, or one
can simply change the weights of different preference
criteria. An other example is to classify criteria under
permanent factors (e.g. soil characteristics) and time-
changeable factors (e.g. vegetation). Permanent
factors dene the habitat potential in the long run.
Time-changeable factors, together with the permanent
ones, dene momentary habitat suitability.
The general purpose of sensitivity analysis in this
study was to nd out the inuence of different criteria
weights on the spatial pattern of the suitability index.
In order to investigate the effect of the time horizon in
examining the location of the most suitable areas,
sensitivity analysis was connected to the temporal
factor. By means of this connection, it was possible
to investigate how the most suitable area shifts, if
suitability is related not only to the situation present at
the moment, but more and more to those character-
istics, which would not change in the long run. With
this purpose in mind in the hierarchy-producing phase,
the rst level factors were arranged according to time-
permanent and time-changeable factors.
3. Case study
3.1. Study area and general course of case study
The case study area, Kivalo forest estate, covers
about 6870 ha and is located in Finnish Lapland
(66823
0
N, 26823
0
E), about 20 km east of the town
of Rovaniemi (Fig. 2). It is state-owned land adminis-
tered by the Finnish Forest Research Institute. The
difference in elevation within the area is 278 m, the
highest point being 360 m above sea level and
the lowest point 82 m above sea level.
The study area was divided into 1833 compartments
composed of relatively homogenous tree stands and
having uniform soil characteristics. The dominant tree
species is Scots pine (Pinus sylvestris), accounting for
44% of the standing volume. Norway spruce (Picea
abies) and Silver Birch (Betula pendula) are the next
common tree species, both accounting for 24% of the
standing volume. The mean volume of growing stock
in the area was only 50 m
3
/ha due to the harsh climate;
the maximum volume recorded in the area was
84 R. Store, J. Kangas / Landscape and Urban Planning 55 (2001) 7993
262 m
3
/ha. The age class distribution was 43% <50
years, 18% 51100 years, and 39% >100 years. The
soil fertility of the area is mainly medium, with some
highly fertile (13%) and poor (15%) parts.
The objective of the case study was to provide
support for the choice of the most suitable old-forest
areas for the polypore species S. odora. The process of
assessing the ecological value of an old-forest area is
often done by considering the threatened aphyllopor-
ales and other old forest indicator species persisting in
the area. S. odora is considered to be one of the most
important boreal old-growth forest indicator species,
e.g. in Finland and Sweden. In the examination of the
ecological potential of the area, it was chosen as one of
the key species to be considered in landscape ecolo-
gical planning.
According to a list approved by the Ministry of
Environment in Finland, S. odora is classied as a
threatened polypore falling into the threat class `rare'.
Also in Sweden and Norway it has been found to be
vulnerable and classied as a threatened species
(Kotiranta and Niemela, 1996). S. odora is an old-
growth forest species growing on Norway spruce.
Several records have also been made of it growing
on Aspen (Populus tremula). It requires a fertile mire-
like habitat with a humid micro-climate and downed
large-sized boles with the bark still attached (Kotiranta
and Niemela, 1996). Scarceness of shady, old-growth
forests with a humid micro-climate is the main reason
for its rareness.
3.2. Estimation of habitat suitability indices
3.2.1. Data procurement
The habitat quality required by S. odora was found
to depend on both vegetation and soil characteristics
(Fig. 3). Its vegetation requirements are related to
the occurrence of a suitable host tree and a humid
Fig. 2. Location of the case study area.
Fig. 3. Habitat factors of S. odora, as applied in the case study.
R. Store, J. Kangas / Landscape and Urban Planning 55 (2001) 7993 85
micro-climate. The following stand-specic mean
characteristics were used in the case study: age of
spruce, stem volume of spruce, diameter at breast
height (dbh) of spruce, and the density of the growing
stock. The habitat factors related to the soil's physical
properties and topography were: slope direction
(aspect), soil moisture and soil fertility. Suitable soil
characteristics enable a humid micro-climate and also
moist and fertile forest site types.
One expert on polypores and another on soil
sciences took part in the evaluation process. The
expert on polypores chose the most important habi-
tat-quality factors for S. odora and also assessed the
importance of these factors. Furthermore, in the pro-
cess of producing sub-priority functions, the experts'
task was to assess the relative priority produced by the
different criterion performance scores. The expert on
soil science determined the factors inuencing soil
moisture and the relative importance of these factors.
In addition, the soil expert evaluated the consequences
of different soil textures with respect to soil moisture
in the study area.
Most of the variables needed in the evaluation
process were collected by means of ocular compart-
ment inventory, and they were then stored into a
database of a GIS application. Some of the evaluation
criteria, such as the spatial distribution of soil moisture
and slope direction (aspect), were elaborated by spa-
tial analysis functions using GIS operations. In GIS,
the collected and produced data were managed so that
the data for every habitat factor were stored in separate
map layers. The GIS used in the case study was Arc/
Info workstation (UNIX) version 7.2. Arc/Info is a
commercial GIS software product developed by ESRI.
The priority functions for habitat suitability were
estimated on grounds of the expert judgements. The
general priority function for habitat suitability is pre-
sented in Table 1. The vegetation factors were con-
sidered to be more important than the factors related to
soil properties (0.7 versus 0.3). The factors directly
related to the properties of the spruces were consid-
ered to be clearly more important than general stand
properties such as the density of growing stock (0.59
versus 0.11). Regarding soil properties, soil fertility
was more important than the factors related to soil
moisture (0.17 versus 0.13).
The sub-priority functions (Fig. 4) of the habitat
factors were estimated using a minimum, maximum
and 23 intermediate values in-between them, and
then comparing the values pairwise with respect to
their sub-priorities. The sub-priority functions for
stem volume and mean age of spruce indicated
decreasing marginal sub-priority. However, the mean
diameter of spruce at breast height and the density of
the growing stock were assessed to have nearly linear
sub-priority functions. The sub-priority functions for
the mean diameter of spruce and the density of
growing stock could also show decreasing marginal
priority, if the maximum values recorded in the case
study area were higher. It is also worth noting that,
according to the judgements made by the expert, very
high soil moisture values decrease habitat suitability in
the case of S. odora.
3.2.2. Overlay analysis
Two deterministic habitat factors were used in the
combining phase to screen out unsuitable areas.
Firstly, the area has to be forest land with the condi-
tions sufcient also for tree growth as regards hydrol-
ogy. Secondly, the area's site class should be fertile
enough to enable a suitable micro-climate and appro-
priate host trees with regard to tree species and size.
Map layers describing these factors were overlaid
numerically to build up a feasible area fullling both
deterministic habitat factors simultaneously.
The deterministic habitat factors used in this study
were not very strict because the purpose was not to
rule out any areas where the circumstances are or
could, in the future, be suitable for S. odora. Of the
6870 ha of the case study area, 5444 ha were con-
sidered to be suitable. Most of the areas deemed to be
feasible were suitable according to their soil and
Table 1
Habitat suitability criteria and their relative importance
Criterion Relative importance
Vegetation 0.7
Density of growing stock 0.11
The dbh of spruce 0.25
Stem volume of spruce 0.14
Age of spruce 0.20
Soil characteristics 0.3
Soil fertility 0.17
Slope aspect 0.04
Soil moisture 0.09
86 R. Store, J. Kangas / Landscape and Urban Planning 55 (2001) 7993
vegetation characteristics, but also areas the vegeta-
tion of which could in the long run develop suitable
were chosen.
The value grids, describing the non-deterministic
habitat factors, were divided into groups according to
the ``corner points'' of the sub-priority functions. For
example, the habitat factor `stem volume of spruce'
was divided into the following groups: <60 m
3
/ha (I),
60120 m
3
/ha (II), >120 m
3
/ha (III). The correspond-
ing linear parts of the piece-wisely linear sub-priority
function could then be determined via formulas used
in the interpolation, respectively:
I y 0:0097x
II y 0:0045x 0:31
III y 0:0025x 0:55
where x is the value of habitat factor, and y the sub-
priority produced by value x.
The habitat suitability index was calculated for each
cell by multiplying each sub-priority grid by its weight
coefcient and then summing the weighted sub-prio-
rities by the means of an arithmetic overlay analysis.
The index map resulting in the habitat suitability
analysis for S. odora is presented in Fig. 5. In this
basic calculation, where weights for vegetation factors
and soil characteristics were 0.7 and 0.3, respectively,
the maximum suitability index value recorded in study
area was 0.83 and the minimum was 0.10 (scale 01).
The mean value was 0.44. The areas in the middle part
of the study area, located highest above the sea level
within the area, seemed to be best suited for S. odora.
Especially the north-facing hillsides in these areas got
high index values. Also, the biggest continuous high
index areas were located in this area while the high
index areas in northern and western part of the study
area were relatively small. The range of variation in
the index value in the study area was high and,
furthermore, high and poor index values were clearly
clustered. Steep gradients follow the forest stand
boundaries, because the coefcients of vegetation
factors were high.
3.3. Sensitivity analysis
Sensitivity analysis was used to present a technique
enabling the temporal factor to be taken into consid-
eration in the process of producing the suitability
index and, on the other hand, to investigate how
changing the weighting of various habitat factors
affected the determination of the preferred areas.
Special attention was paid to the weighting of time-
permanent factors. Giving more weight to them means
giving less weight to factors whose values change with
time. Correspondingly, emphasising the importance of
permanent factors means preferring the future habitat
Fig. 4. The sub-priority functions estimated in the case study.
R. Store, J. Kangas / Landscape and Urban Planning 55 (2001) 7993 87
potential over the present qualities of habitats. The
habitat factors were divided into time-permanent and
time-changeable factors for this analysis.
The habitat factors connected to soil characteristics,
such as fertility and moisture as well as topography,
can be considered to be relatively constant. However,
habitat factors connected to vegetation mostly change
slowly through time, but also rapid and radical
changes are possible. Usually the most extensive
changes are due to the consequences of human activ-
ities, e.g. clear cutting. The weighting scheme, when
approaching the problem this way, reects the impor-
tance of the present vegetation and the vegetation
potential in the future (determined by soil character-
istics).
In practise, sensitivity analysis was accomplished
by applying different weighting schemes for the two
main decision criteria. In the basic computation the
weight of vegetation (0.7) and the weight of soil
characteristics (0.3) demonstrate the importance of
the two main factors at the present moment according
to opinion of the expert on polyspores. In addition to
Fig. 5. The process of combining habitat factors to suitability index.
88 R. Store, J. Kangas / Landscape and Urban Planning 55 (2001) 7993
Fig. 6. Sensitivity analysis, where weights 0.5, 0.7 and 1.0 where applied for soil characteristics.
R. Store, J. Kangas / Landscape and Urban Planning 55 (2001) 7993 89
basic calculation, weighting schemes 0.5 and 0.5, 0.3
and 0.7, 0.9 and 0.1, and 0.0 and 1.0 were applied
for vegetation and soil characteristics, respectively.
The results illustrate how changes in weighting affect
the optimal choice of habitats to be conserved. Also,
the alternatives where suitability maps are based either
only on soil characteristics, or only on vegetation
characteristics, were useful in identifying areas where
vegetation preservation was less important because
soil factors limit the site's suitability.
For the purpose of sensitivity analysis, suitability
maps for every weighting scheme were created in GIS.
In suitability maps the pixels of the index maps were
classied into ten categories according to suitability.
The class ranges were determined so that the area
covered by pixels was equal in each category. Sensi-
tivity analysis was done by comparing these suitability
maps. In the basic calculation, the most suitable areas
were located in the middle parts of the study area.
When the importance of soil characteristics was
increased, the within-stand variation of the forest
compartments also increased. When the importance
of soil characteristics increased close to 1, then also
the location of the most suitable areas shifted from the
middle part's old high-elevation spruce forest to more
fertile and moist, but also younger, spruce forest in the
northern part of the study area (Fig. 6). This shows
that, in the case of S. odora, the temporal factor had a
major inuence on the results.
4. Discussion
The possibilities of GIS were used in this study in
combination with state-of-the-art decision-analysis
methods to develop more exible habitat suitability
analyses utilising expert knowledge. The theoretical
background in the combining phase was provided by
the multi-attribute utility theory (MAUT) and conti-
nuos attributes and non-linearities were handled by
means of sub-priority functions. A GIS was used as a
platform enabling the management of the criterion
data, production of criterion layers, calculation of
attributes by means of spatial analysis, carrying out
the combining of decision criteria by means of carto-
graphic modelling, and nally conducting sensitivity
analyses and production of the maps needed in the
evaluation process.
The major advantage of the method presented in the
present study is in the possibility it offers for produ-
cing suitability indices for large areas within a reason-
able period of time, and also for species not having
statistical suitability models based on empirical data
available. This is enabled by using expert knowledge
instead of evaluation models based on empirical data
and on the other hand by utilising the data processing
and production capabilities of GIS. The use of sub-
priority functions is one way to handle the non-line-
arities between the raw scores and the relative priority
produced by these scores. With sub-priority functions
it was also possible to deal with continuous variables
without classifying them.
In the discipline of landscape ecology, discussion
has arisen about the nature of boundaries between
different features in the landscape. In many applica-
tions of landscape ecology, or more generally in the
management of natural resources, the boundary is
determined by means of crisp borders. However, in
real-life landscapes, the boundary between two fea-
tures is more often soft or fuzzy rather than crisp.
In the case study, the data used were collected for
the purpose of enabling forest management planning
where the basic spatial entity is a homogenous forest
compartment with crisp boundaries. This being the
case, spatial patterns of suitability variables based on
stand characteristics, such as stem volume and density
of the growing stock, follow crisp stand boundaries.
However, the data produced by the digital elevation
model, such as aspect and wetness index, can be
considered as being a continuous surface and they
can be discretised into regular grids, for instance. Then
each cell can have a different value for the attribute
without any unnatural crisp boundaries. Besides the
problem of crisp boundaries, the forest compartment
approach involves the problem that in reality there is
often within-compartment variation as regards stand
characteristics.
The method presented in this study is based on
analysing and overlaying the data in raster format.
This, and the technique whereby the evaluation of
continuous variables is done by means of partial
priority functions, provides a methodical capability
for handling both crisp and fuzzy boundaries, and also
the within-stand variation. The main factors limiting
the full use of the capability were the accuracy and
resolution of input data. The nature of the boundaries
90 R. Store, J. Kangas / Landscape and Urban Planning 55 (2001) 7993
and the needed accuracy depends on the requirements
of the application.
The suitability index calculated for S. odora was not
as such the most essential point of this study. The
index was produced mainly in order to demonstrate
the purposes and qualities of the method, and there-
fore, eld checks were not included in the case study.
The criteria used in the case study represented just one
expert view on the habitat factors of the species.
Instead of using only one expert it is possible to
use a group of experts and estimate the sub-priority
functions and criteria weights according to the opinion
of that group (Kangas et al., 1998). The Delphi
technique, for example, could be used in searching
for the coherence between the views of experts
involved in the process.
Determining the habitat factors and their weights
are two crucial phases in the process. It is possible that
even slight changes in weight coefcients have a
conclusive effect on the results of suitability analyses.
Also, the structure of the decision hierarchy can affect
the results. In practice, it is impossible to include all
the factors affecting the suitability to an evaluation
model. The choice of the model is always a compro-
mise between accuracy and costs.
Also, the capabilities embodied in the model for
handling spatial relationships have bearing on the
accuracy of the results in the case of S. odora. For
example, variables describing the size of the core area
and the distance between habitat patches could
increase the accuracy of suitability analysis. Some
further research and method development is needed to
get more spatial relationships included in the analysis.
In a situation where the decision criteria are not
independent of each other, the additive assumption as
applied in the case study is not valid. In this context,
the additive assumption means that a suitability index
value can be calculated as the weighted sum of sub-
priorities with respect to individual habitat factors.
One way to deal with non-additive variables in the
model is to estimate the interactions of habitat factors
and add the interaction terms to the additive priority
function (Keeney and Raiffa, 1976). Another way to
handle this independence problem is to use rules of
combination method as presented by Hopkins (1977).
Using the said method, suitabilities are assigned to sets
of combinations of factors and expressed in terms of
verbal logic. It is also possible to deal with non-
additivities by transforming and combining habitat
factors (Pukkala et al., 1997). When HERO-type
priority models are applied, also multiplicative parts
can be added into the model (Kangas and Kangas,
1998).
In sensitivity analysis, the effects of changes in the
weights of different habitat factors were investigated.
In addition to the uncertainties associated with the
weight coefcients, the quality of the nal results was
inuenced by the accuracy of the input data and error
propagation. The assumption in the approach pre-
sented in this study was that the input data are error
free. However, the information available in habitat
suitability analysis is often uncertain and imprecise
because of matters such as measurement errors and
errors arising from processing of the data.
In the case study, most of the data layers were
collected by means of eld inventory and the rest
were produced by using spatial analysis. The most
signicant factors affecting the quality of the results in
the case of S. odora were the uncertainty associated
with the data produced by spatial analysis and the
uncertainty associated with the modelling of expert
knowledge. The data measured in eld inventories,
e.g. tree age, stem volume, diameter at breast height,
were not error free, but the errors associated to them
were not crucial. It is possible to determine the
sensitivity of the model to errors and error propagation
by means of tools based on the analytical error pro-
pagation method and Monte Carlo simulation (Bur-
rough and McDonnell, 1998).
Vigorous research in the eld of statistical metho-
dology of modelling expert knowledge and on analys-
ing and reducing uncertainties in expert judgements is
going on (Alho et al., 1996, 2001; Alho and Kangas,
1997; Kangas et al., 1998; Leskinen, 2000). In the
future, the results of these studies will be integrated
with GIS-based habitat suitability modelling. This
will, hopefully, alleviate the problems regarding the
quality of the analysis results.
If the suitability index is used to select a network of
areas where selection is not independent of the com-
position of previously selected sites, the problem is
that the method does not offer any analytical way to
take into consideration the effects of earlier selections
on the ranking of the remaining areas. In those cases,
e.g. nature reserve network selection, optimisation or
heuristics algorithms provide a better approach than
R. Store, J. Kangas / Landscape and Urban Planning 55 (2001) 7993 91
the method presented in the present study (e.g. Pressey
et al., 1997).
Sensitivity analyses examined the inuence of the
temporal factor by changing the weights given to
permanent and time-changeable habitat factors. The
spatial pattern of the suitability index changed along
with the weighting schema. This means that the
optimal values of the time-permanent factors did
not follow those of the time-changeable factors. Also,
the variety within the forest compartments was
increased when more emphasis was given to the
time-permanent decision factors.
Different weighting of time-permanent and time-
changeable factors leading to a different choice among
habitats has consequences both for management plan-
ning and species conservation. The most effective
alternatives in the long run can be found in areas
which are not among the best ones at the moment,
when a momentary evaluation is made emphasising
factors whose values change with time. Especially
when dealing with goals related to nature conserva-
tion, the time horizon has to be long enough. This is
important to take into consideration, e.g. in landscape
ecological planning.
Sensitivity analysis using the temporal factor is
most useful in situations where the objective is related
to a long time horizon and when one is not in such a
trade-off situation where one would have to give away
some areas if one were to want to preserve some
others. For example, the temporal factor plays an
important role if one is going to increase the size of
an existing nature conservation area and at the same
time one wants to nd out the optimal solution in the
long run. Then it might be better to choose an area with
optimal soil characteristics and adequate vegetation
characteristics rather than an area with optimal current
vegetation characteristics but only average suitability
with regard to soil characteristics.
On the other hand, species usually require a con-
tinuum of suitable habitats in time and place to survive
and, therefore, it is important to avoid changes in
species habitats that are too sudden and distinct.
Certainly it is not appropriate to give away all of
today's best habitats because vegetation development
takes a long time and involves many uncertainty
factors. One needs to nd an appropriate dynamic
`habitat slide' if the purpose is to shift habitats in the
course of time to the best areas from the viewpoint of
time-permanent habitat factors. The modelling
approach presented, especially the sensitivity analysis
as carried out in the case study, gives valuable support
to this kind of dynamic forest planning. However, the
method presented is at its best in tasks where the
purpose is to rank a set of alternative sites according to
the habitat requirements of a certain species or where
the aimis to nd sub-areas where it is most probable to
nd the species under examination.
Developing methods for managing dynamic ``habi-
tat slides'' in nature conservation problems, as well in
any natural resources planning, is an important topic
of future research. Other important topics include
the combined use of expert knowledge and existing
evaluation models in habitat suitability modelling,
including several species' concurrent suitability con-
siderations, and taking more spatial factors into suit-
ability considerations. Furthermore, dealing with the
interdependencies between habitat factors considered
in modelling processes is worth studying further. The
GIS-based framework presented in this study gives a
solid basis for this kind of methodological develop-
ment work.
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Ron Store is presently a Researcher at the Finnish Forest Research
Institute, Kannus Research Station. He received the MSc degree in
Forestry in 1995 from the University of Joensuu (Finland). His
field of research includes utilising the capabilities of geographical
information systems (GIS) in multi-objective forest management
planning. His other areas of professional interests are GIS,
cartographic modelling, multi-objective forest planning, multi-
criteria evaluation.
Jyrki Kangas is at present the Director of the Kannus Research
Station of the Finnish Forest Research Institute. He has been the
Professor of Forest Management Planning (acting) and Associate
Professor of Forest Economics (acting), at the University of
Joensuu. He received his MSc in 1986, and DSc in 1992 from the
University of Joensuu. He has been a Docent at the University of
Joensuu since 1994, an Editorial Board Member of Journal of
Environmental Management, Managing Board Member of Silva
Fennica, as well as a referee of a number of journals, supervisor of
many doctoral theses, opponent or peer-reviewer of many theses,
and involued in international evaluation tasks. His areas of interest
include forest planning, natural resource management, optimisation
methods, multicriteria decision support, ecological modelling,
expert judgments. Kangas has also received the Scientific
Achievement Award conferred upon him by the International
Union of Forest Research Organisations (IUFRO). His total number
of publications is about 150 of which about 90 are scientific
publications and about 45 are peer-reviewed articles or mono-
graphs.
R. Store, J. Kangas / Landscape and Urban Planning 55 (2001) 7993 93

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