Agriculture, Ecosystems and Environment 98 (2003) 8798

Biodiversity indicators: the choice of values and measures

Peter Duelli , Martin K. Obrist
Swiss Federal Research Institute WSL, Zrcherstrasse 111, CH-8903 Birmensdorf-Zrich, Switzerland

Abstract
Ideally, an indicator for biodiversity is a linear correlate to the entity or aspect of biodiversity under evaluation. Different
motivations for assessing entities or aspects of biodiversity lead to different value systems; their indicators may not correlate at
all. For biodiversity evaluation in agricultural landscapes, three indices are proposed, each consisting of a basket of concordant
indicators. They represent the three value systems conservation (protection and enhancement of rare and threatened species),
ecology (ecological resilience, ecosystem functioning, based on species diversity), and biological control (diversity of
antagonists of potential pest organisms). The quality and reliability of commonly used indicators could and should be tested
with a three-step approach. First, the motivations and value systems and their corresponding biodiversity aspects or entities
have to be defined. In a time consuming second step, a number of habitats have to be sampled as thoroughly as possible with
regard to one or several of the three value systems or motivations. The third step is to test the linear correlations of a choice
of easily measurable indicators with the entities quantified in the second step. Some examples of good and bad correlations
are discussed.
2003 Elsevier Science B.V. All rights reserved.
Keywords: Biodiversity; Indicator; Arthropods; Correlate

1. Who needs biodiversity indicators?

National and regional agencies for nature conservation, agriculture, and forestry have to monitor species
diversity or other aspects of biodiversity, both before
and after they spend tax money on subsidies or ecological compensation management, with the aim of
enhancing biodiversity (European Community, 1997;
Ovenden et al., 1998; Wascher, 2000; Kleijn et al.,
2001). Similarly, international, national or regional
non-governmental organisations (NGOs) may want
to monitor aspects of biodiversity at different levels
and scales (Reid et al., 1993; IUCN, 1994; Cohen
and Burgiel, 1997). In scientific research biodiversity
Corresponding author. Tel.: +41-1-739-2376;
fax: +41-1-739-2215.
E-mail address: peter.duelli@wsl.ch (P. Duelli).

indicators can be used as quantifiable environmental factors. Since the biodiversity of even a small
area is far too complex to be comprehensively measured and quantified, suitable indicators have to be
found.
Those who are responsible for comparing and evaluating biodiversity have a strong incentive to choose a
scientifically reliable and repeatable indicator, which
inevitably increases costs. The financing agencies usually opt for a financially reasonable approach, which
often results in programmes addressing only essential
work. The resulting compromises make optimisation
of the choice of biodiversity indicators and methods
of fundamental importance.
A recent international electronic conference on biodiversity indicators (http://www.gencat.es/mediamb/
bioind, 2000) has revealed widely differing views on
why and what to measure and quantify.

0167-8809/$ see front matter 2003 Elsevier Science B.V. All rights reserved.
doi:10.1016/S0167-8809(03)00072-0

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Fig. 1. Provisional domain tree of biodiversity based on the survey of 125 text documents in English (Kaennel, 1998). Concepts used by various authors to define biodiversity
are in square boxes, related concepts in rounded boxes. Type and direction of conceptual relationships are indicated by arrows. Synonyms and quasi-synonyms are in italics.

P. Duelli, M.K. Obrist / Agriculture, Ecosystems and Environment 98 (2003) 8798

2. Why is it so difficult to reach a consensus on

the use of biodiversity indicators?
The complexity of all the aspects of the term biodiversity is illustrated in Fig. 1. It is obvious that
no single indicator for biodiversity can be devised.
Each aspect of biodiversity requires its own indicator.
The difficulties for reaching a consensus on the use
of biodiversity indicators are manifold. They imply
differing choices for values and measures, which will
be discussed here more in detail.
Terms such as biodiversity, indicator or index are
not well defined and their use varies between different
countries and disciplines. Dismissing research findings
or scientific reports simply on the grounds of differing
views on the use of particular terms (semantic discrimination) would be counterproductive, but study reports
must clearly state what is meant by the terms used. A
helpful review on indicator categories for bioindication is given by McGeoch (1998).
In this paper, the term indicator is used in the sense
of any measurable correlate to the entity to be assessed: a particular aspect of biodiversity.
The most promising and convincing indicators of
biodiversity are measurable portions of the entity
that we consider to represent a target aspect of biodiversity. The term index is used here in the sense
of a scaled measure for one or several concordant
indicators.
3. Indicator FOR or FROM biodiversity?
A first major source of misunderstanding is, whether
biodiversity itself is to be indicated, or whether certain components of biodiversity are used as indicators for something else. Until 1990, the search for
bioindicators had focussed on indicators of environmental health or ecological processes such as
disturbance, human impact, environmental or global
change (Hellawell, 1986; Spellerberg, 1991; Meffe
and Carroll, 1994; Dufrene and Legendre, 1997).
After the world-wide launch of the term biodiversity
at the Rio Convention in 1992, there was a sudden
and drastic shift in the published literature towards
the search for indicators of biodiversity itself (Noss,
1990; Gaston and Williams, 1993; Gaston, 1996a;
Prendergast, 1997). Since then, however, the term

89

biodiversity has sometimes been used to allude to or

indicate some aspect of environmental quality.
If a species or a group of species is a good indicator
for lead contamination, it may not indicate biodiversity, i.e. there may not be a linear correlate to biodiversity. It is fundamentally a contamination indicator,
or an environmental indicator (McGeoch, 1998) rather
than a biodiversity indicator.
However, real biodiversity indicators may be
needed to measure the impact of e.g. lead contamination on biodiversity itself (indicator FOR biodiversity). Such an assessment is different from measuring
the impact of lead on a selected taxonomic group,
which had been chosen because it is especially sensitive to lead poisoning (indicator FROM biodiversity).

4. Alpha-diversity, or contribution to higher

scale biodiversity?
A second major dichotomy in the value system for
biodiversity indicators is the question of whether the
species (or allele, or higher taxon unit) diversity of a
given area is to be indicated (local, regional or national
level), or if the contribution of the biodiversity of that
area to a higher scale surface area (regional, national,
global) is important.
In the first case (alpha-diversity, e.g. species richness of an ecological compensation area), an indicator
ideally has to be a linear correlate to the biodiversity
aspect or entity of the surface area in question. Each
species has the same value.
In the second case, the value of the measurable units
of biodiversity (alleles, species, ecosystems) depends
on their rarity or uniqueness with regard to a higher
level area. A nationally rare or threatened species in
a local assessment has a higher conservation value
than a common species, because it contributes more
to regional or national biodiversity than the ubiquitous species. Thus a biodiversity indicator in the latter
case not only has to count the units (alleles, species,
ecosystems), but it has to value them differently and
add the values.
The best known examples are red list species. For
measuring alpha-diversity, they are not given a value
that is greater than any other species in a plot or trap
sample, but for measuring the conservation value of a
plot, their higher contribution to regional, national, or

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even global biodiversity has to be recognised. Raised

bogs are notorious for their poor species richness,
but if only a few raised bogs are left within a country, the few characteristic species present in a good
bog are of very high national importance. The problems of estimating complementarity or distinctness
are addressed e.g. by Colwell and Coddington (1994)
and Vane-Wright et al. (1991), endemism and spatial
turnover by Harte and Kinzig (1997).
This dichotomy between species richness and
conservation value is the most fervently debated
issue among applied biologists concerned with biodiversity indicators, and a recurrent source of misunderstandings. It will be elaborated further in the chapter
on value systems.

5. Indicator for what aspect of biodiversity?

After agreement on indicators FOR biodiversity,
and a decision between alpha-diversity and contribution to higher scale biodiversity, there is still
potential for disagreement on what is biodiversity?
(Gaston, 1996c). In practice, in a majority of cases,
species are the units of biodiversity (Claridge et al.,
1997). However, species diversity can be measured as
simple number of species, usually of selected groups
of organisms, or species richness may be combined
with the evenness of the abundance distribution of the
species. The best known indices are the Shannon in-

dex, the Simpson index and Fishers alpha (Magurran,

1988). Recent observations (Duelli, unpubl.) have
shown that when undergraduate biodiversity students
in entomology lectures have to choose which of the
two communities shown in Fig. 2 (without seeing the
text below them) they consider to be more diverse,
more than half of them decide for the left population, because they consider evenness to be of greater
importance than species numbers. When individuals
from other disciplines were asked during lectures and
seminars, particularly conservationists and extension
workers in agriculture and forestry, species numbers
are decisive. In recent years, indices involving evenness have essentially fallen out of favour, mostly
because they are difficult to interpret (Gaston, 1996c).
Particularly in agriculture or forestry, single species
are often collected in huge numbers with standardised
methods, which results in a drastic drop of evenness
and hence yields low diversity values, in spite of
comparatively high species richness.
The definition of biodiversity given in the international Convention on Biological Diversity (Johnson,
1993) encompasses the genetic diversity within
species, between species, and of ecosystems. Furthermore, Noss (1990) distinguished three sets of
attributes: compositional, structural and functional
biodiversity (see also Fig. 1). The most common approach is to measure compositional biodiversity. Presumably, both structural and functional biodiversity
are either based on or lead to higher compositional

Fig. 2. Which of the two populations do you consider to have a higher biodiversity? A choice test for biodiversity evaluation regularly
offered by the first author to students and at public lectures. For the vote, only the upper part without text is shown.

P. Duelli, M.K. Obrist / Agriculture, Ecosystems and Environment 98 (2003) 8798

diversity. We are convinced that ecosystem diversity, as well as structural and functional diversity, is
somehow reflected in the number of species present.
If they are not correlated with species richness, they
must be special cases and not representative as biodiversity indicators. More trophic levels will normally
include more species, and a higher structural diversity
will harbour more ecological niches. In fact, there is
increasing evidence that at least for some taxonomic
groups, species numbers are correlated with habitat
heterogeneity (Moser et al., 2002), but not in others
(Rykken and Capen, 1997).
For all these hierarchical separations or entities
within the huge concept of biodiversity, separate
comprehensible indicators can be researched and developed. In many cases, however, a rigorous scientific
test may show that the conceptual entities are difficult
to quantify (Prendergast, 1997; Lindenmayer, 1999;
Noss, 1999), or they are basically reflected in other,

91

better quantifiable measures of biodiversity, such

as species richness (Gaston, 1996b; Claridge et al.,
1997).
The aspect of intraspecific diversity is a different
case. To our knowledge there is no published example
of a tested correlation between inter- and intraspecific
diversity.

6. Value systems
People involved in developing or using biodiversity
indicators are influenced by their personal and/or professional goals. They all may want to measure or monitor biodiversity, but they address different aspects of
it. Their focus depends on their motivation for dealing with biodiversity. In an agricultural context, and
in an industrialised country in Europe, the three most
important motivations to enhance biodiversity are

Fig. 3. Illustration of the hypothesis that abundant species usually are of higher ecological but lower conservation value, in contrast to
rare and threatened species. Stars indicate red list species collected with pitfall traps, yellow water pans and window interception traps in
a semidry meadow (Duelli and Obrist, 1998). Number of individuals (N Ind(log)) are plotted versus number of species (N species).

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1. Species conservation (focus on rare and endangered

species).
2. Ecological resilience (focus on genetic or species
diversity).
3. Biological control of potential pest organisms (focus on predatory and parasitoid arthropods).
There are additional motivations, of course, but
either they are closely related to the ones mentioned
here, or their causal link to biodiversity is less clear
(e.g. sustainability, landscape protection, cultural
heritage).
Each of these three aspects of biodiversity requires
its own indicators. They often do not correlate with
each other or even show a negative correlation. Consequently, simply adding up different indicators may
lead to misinterpretations, as long as they do not address the same aspect of biodiversity. Species conservation focusses on rare and threatened species and
often regards more common species in a derogatory
way as ubiquists of little interest. Ecologists, on the
other hand, focus more on abundant species, because a
species on the verge of extinction is likely to have less
significant ecological influence. The hypothesis of an
almost vicarious relationship between the motivations

of species conservation and ecological resilience

is illustrated in Fig. 3.
Prendergast et al. (1993) found low coincidence of
species-rich areas and areas harbouring rare species
for either plants, birds, butterflies or dragonflies. An
investigation of carabid beetles in Scotland (Foster
et al., 1997) showed that neither the number of red
list species nor the number of stenotopic (faunistically
interesting) species are correlated with the mean total
number of carabid species in a variety of habitats such
as moorland, grassland, heathland, peat, saltmarsh,
bracken and swamps (Fig. 4). In an intense investigation with 51 trap stations and standardised sampling
methods in field and forest habitats in Switzerland,
the number of red list species of all identified arthropod groups was not significantly correlated to overall
species richness per trap station (Fig. 5), while e.g.
the numbers of aculeate Hymenoptera species correlated well (R2 = 0.88; Fig. 6). In an assessment of
the effects of ecological compensation measures in
Swiss crop fields and grassland, the number of butterfly species did not show any correlation with the
species numbers of spiders (Jeanneret, pers. comm.).
In an effort to test the suitability of Collembola as
indicators of the conservation value of Australian

Fig. 4. Neither red list carabid species nor stenotopic carabid species are correlated significantly with the average number of carabid species
collected in 18 types of habitats using pitfall traps. Data from Foster et al. (1997).

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93

Fig. 5. No significant correlation exists between the number of red list species (from numerous arthropod taxa) and the overall number of
arthropods collected with flight traps, pitfall traps and yellow water pans at the same 51 locations (Araneae, Coleoptera, Diplopoda, Diptera
(Syrphidae only), Heteroptera, Hymenoptera (Aculeata only), Isopoda, Mecoptera, Megaloptera, Neuroptera, Raphidioptera, Thysanoptera).
Data from agricultural areas (Duelli and Obrist, 1998) and forest edges (Flckiger, 1999).

grasslands, Greenslade (1997) found no correlation

with species numbers of ants and carabid beetles.
The optimal approach is to select a basket of
indicators for each motivation, similar to the Dow

Jones index for the stock exchange. The measured

indicators within one basket have to be fairly concordant and are pooled to form an index. The result is a set of three separate indices for the three

Fig. 6. Species numbers of aculeate Hymenoptera (bees, wasps and ants) show excellent correlation with the overall number of arthropod
species at 51 locations (for details of data sources see Fig. 5).

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basic motivations conservation, ecology and pest

control.
7. How to select indicators for the three main
motivations
7.1. Several steps are necessary
The most accurate indicators of biodiversity are
proven linear correlates of the entity or aspect of biodiversity being evaluated. McGeoch (1998) proposed a
nine-step approach for selecting bioindicators among
terrestrial insects. Basically, the whole procedure can
be separated into three steps. The first step is to define the aspect or entity in as quantifiable a way as
possible. The second step is to actually quantify that
aspect or entity in a statistically reliable number of
cases. The third step is a rigorous test for linear correlation in a set of proposed indicators. The urgent
need to perform a scientifically solid test has been advocated repeatedly (Balmford et al., 1996; McGeoch,
1998; Niemel, 2000).
Starting with the first step, the three mayor motivations for protecting or enhancing biodiversity in agricultural landscapes are differentiated.
7.2. Conservation (an index based on the motivation
to protect or enhance threatened species)
For assessing the value of a given habitat, e.g.
an ecological compensation area, for species conservation, the entity to indicate is the accumulated
conservation values (e.g. red list status) of all species
present in that area. The highest values are contributed
by species of national or even global importance,
while the so-called ubiquists are of little value. The
second step thus is a comprehensive measurement of
the conservation values in a number of ecosystems or
habitat types.
The third step would be to find and test the best
linear correlate to that otherwise elusive entity conservation value. The standard indicators for the
conservation basket are numbers of red list species of
selected taxa, weighed according to their category of
threat. However, only very few of the tens of thousands of species present in a country are listed; in
Switzerland they are a mere 7% of all known animal

species (Duelli, 1994). Inevitably, the choice of the

groups of organisms used for an inventory depends
strongly on the red lists available, and on the availability of specialists to identify the listed organisms.
Lacking the information on the second step (full
account of the conservation value of an area), it is
not currently possible to come up with a scientifically
tested indicator for that value. Nevertheless, a correlation between the cumulated conservation values of all
presently available red listed species per habitat with
the conservation values of single taxonomic groups,
such as birds, butterflies or carabids, would at least
give greater credibility to the red list species approach.
In addition to red list status (degree of threat of extinction), species values have been calculated on the
bases of national or global rarity (Mossakowski and
Paje, 1985) or endemism. The rationale in the context
of habitat evaluation is that the presence of a nationally or globally rare species increases the biodiversity
value of that habitat, because it contributes more to
the conservation of national or global biodiversity than
the presence of a ubiquitous species.
Only after a reliable basket of indicators for conservation value has been established, are further steps
possible to test the correlative power of potential indicators such as length of hedgerows, amount of dead
wood, or the surface of ecological compensation areas per unit area. Environmental diversity (ED) as a
surrogate measure of the conservation value was proposed by Faith and Walker (1996), but so far there are
no empirical data to test their proposal.
7.3. An index for the motivation pest control
For the biodiversity aspect of biological control of
potential pest organisms, the first step may be to define the measurable entity as the species diversity of
all predators or parasites of potential pest organisms.
For short-term interests, the number of individuals of
beneficial organisms may appear more important than
species richness, because prey and hosts are reduced
by the number of antagonistic individuals rather than
by species numbers (Kromp et al., 1995; Wratten
and Van Emden, 1995). However, with a longer-term
perspective on maintaining a high diversity of antagonist species of potential pest organisms is certainly
more important. While the species richness of predators in a small area can be assessed with reasonable

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accuracy and effort, the diversities of parasitoids are

much harder to quantify.
The second step is therefore to test inventory methods, and selected taxa for their correlation with the
above biodiversity aspect of biological control. At
present species numbers of carabid and staphylinid
beetles, as well as spiders, are often used as indicators because of established standardised collecting
methods (Duffey, 1974; Desender and Pollet, 1988;
Halsall and Wratten, 1988) and readily available keys
for identification and interpretation. Specialised aphidophaga among the syrphid flies, coccinellids and
Neuroptera are another option, but so far the methods are not fully standardised. Parasitoid wasps and
flies are promising, but so far there is no easy way to
identify them to the species level. Other possibilities
for indicators to test are ratios between herbivores
and predators, or parasitoids and a range of other
arthropods (see e.g. Denys and Tscharntke, 2002).
7.4. An index for ecological resilience
For the basket of indicators for the motivation ecological resilience (Balance of Nature, Pimm, 1991),
the entire genetic and taxonomic spectrum of biodiversity is the entity to be indicated. The assumption is
that the higher the number of alleles and species, the
higher is the ecological potential of an ecosystem to
react adequately to environmental change.
Here again, a first step requires quantification of
a measurable proportion of local organismic diversity, which can be trusted to represent total species
richness of animals and plants (alpha-diversity). Realistically, only few and small areas will ever be
fully assessed. For the second and third steps, approximations with large, measurable proportions
of alpha-diversity have to be used to test potential
indicators.
These ecological indicators can be seen as indicators for ecosystem functioning (Schlpfer et al., 1999)
and are representing a very basic notion of wholesale
biodiversity. Most studies claiming to measure or indicate biodiversity assume that the group of organisms
they investigate is somehow representative of biodiversity. However, in only very few cases has the correlation between a group or several groups of species
with a more or less representative sample of all organisms been measured and published (Abensperg-Traun

95

et al., 1996; Balmford et al., 1996; Cranston and

Trueman, 1997; Duelli and Obrist, 1998).
8. Effort and costs, the limiting factors for the
choice of measures
8.1. The dilemma of indicating complexity with
simple measures
Large environmental monitoring programmes usually avoid using invertebrates for their indicators,
although these constitute by far the largest portion of
measurable biodiversity. To cut down on effort and
costs, measurement of the immense richness and quantity of invertebrates has to be reduced to an optimised
selection of taxa. The proposed three-step approach
allows for testing all kinds of indicators for their correlation with aspects of biodiversity. The search for
linear correlates of quantified entities or aspects of
biodiversity is not limited to taxonomic units. Instead
of choosing birds or grasshoppers as indicators, the
spectrum of taxa considered can be determined by an
inventory method such as Berlese soil samples or flight
interception traps. The broader the taxonomic spectrum of the samples, the higher the chance of obtaining a good correlation with the entity to be assessed.
Furthermore, indicators, which are not part of the organismic spectrum, can also be tested in the three-step
approach: habitat diversity and heterogeneity, disturbance by traffic, neighbourhood or percentage of protected areas, etc. At present, various indicators are in
use, but few of them have been tested for their correlation with aspects of biodiversity. At least in Neotropical butterflies, a positive correlation of species richness
was found with composite environmental indices of
heterogeneity and natural disturbance (Brown, 1997).
8.2. Plots and transects
Plots (for plants) and transects (for birds and insects such as butterflies, dragonflies and grasshoppers)
are widely used relative assessment methods for the
species richness of a selected group of organisms (e.g.
Pollard and Yates, 1993; Wagner et al., 2000). The
main advantages are that the specimens survive the inventory (important for indicating conservation value),
and that large areas can be searched in a relatively

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P. Duelli, M.K. Obrist / Agriculture, Ecosystems and Environment 98 (2003) 8798

short time. Scientifically, the drawback is that usually

there are no voucher specimens kept for verifying the
identification. Also, these popular groups (except for
vascular plants) have only few species in agricultural
habitats, so their species richness, even if cumulated,
never reaches 1% of the local species diversity of all
organisms. Their correlation power with local species
diversity has never been tested. Vascular plants, on
the other hand, seem to correlate reasonably well with
overall organismic diversity (Duelli and Obrist, 1998).
Plots and transects are low budget measures and worth
testing for their correlation power in the conservation
and ecology baskets of indicators.
8.3. Standardised trapping methods for arthropods
Pitfall traps for surface dwelling arthropods and various kinds of flight traps for insects are often used
for biodiversity assessment in agricultural areas. Either one or a few taxonomic groups are collected over
longer periods, or a larger number of taxa are sampled
within a shorter collecting period. In both cases, suitable correlates have been found for the indicator basket of ecological resilience (Duelli and Obrist, 1998).
Bugs (Heteroptera), and wild bees and wasps (aculeate Hymenoptera; see also Fig. 6) collected during
an entire vegetation period, where highly correlated
with overall species richness, while carabids and spiders in pitfall traps were not. Reducing the collecting
time to five carefully selected weeks, but extending
the spectrum of identified taxa (Duelli et al., 1999),
yielded correlation values comparable to those of seasonal collections of bugs or bees. Tests are under way
to further reduce the effort required for collecting and
identifying through an adaptation of the Australian
method of Rapid Biodiversity Assessment (Cranston
and Hillman, 1992; Oliver and Beattie, 1996). With
that method, the whole taxonomic spectrum collected
within a few selected weeks in a standardised trap
combination is considered, but only at the level of
morphospecies, i.e. without identifying the catches
to the species level (Duelli et al., unpubl.). Obviously, the resulting indicator will not be useful for
the indicator baskets of conservation or pest control,
where identification of the species is essential. However, it is a promising monitoring device for the indication of alpha-diversityor the ecological resilience
basket.

9. Conclusions
There is no single indicator for biodiversity. The
choice of indicators depends on the aspect or entity of
biodiversity to be evaluated and is guided by a value
system based on personal and/or professional motivation. Each biodiversity index for a particular value
system should consist of a basket of methods with one
or several concordant indicators. In order to achieve
greater reliability and a broader acceptance, indicators
have to be tested for their linear correlation with a substantial and quantifiable portion of the entity to assess.
The challenge now is to assign all the presently used or
proposed indicators to a basket with a declared value
systemand to test them with empirical measures.
References
Abensperg-Traun, M., Arnold, G.W., Steven, D.E., Smith, G.T.,
Atkins, L., Viveen, J.J., Gutter, M., 1996. Biodiversity indicators
in semiarid, agricultural Western Australia. Pacific Conserv.
Biol. 2, 375389.
Balmford, A., Green, M.J.B., Murray, M.G., 1996. Using
higher-taxon richness as a surrogate for species richness. I.
Regional tests. Proc. R. Soc. Lond. B 263, 12671274.
Brown, K.S., 1997. Diversity, disturbance and sustainable use
of Neotropical forests: insects as indicators for conservation
monitoring. J. Insect Conserv. 1, 2542.
Claridge, M.F., Dawah, H.A., Wilson, M.R. (Eds.), 1997. Species:
The Units of Biodiversity. Chapman & Hall, London.
Cohen, S., Burgiel, S.W. (Eds.), 1997. Exploring Biodiversity
Indicators and Targets under the Convention on Biological
Diversity. BIONET and IUCN, Washington, DC and Gland.
Colwell, R.K., Coddington, J.A., 1994. Estimating terrestrial
biodiversity through extrapolation. Phil. Trans. R. Soc. Lond.
B 345, 101118.
Cranston, P.S., Hillman, T., 1992. Rapid assessment of biodiversity
using biological diversity technicians. Aust. Biol. 5, 144154.
Cranston, P.S., Trueman, J.W.H., 1997. Indicator taxa in
invertebrate biodiversity assessment. Mem. Mus. Victoria 56,
267274.
Denys, C., Tscharntke, T., 2002. Plantinsect communities and
predatorprey ratios in field margin strips, adjacent crop fields,
and fallows. Oecologia 130, 315324.
Desender, K., Pollet, M., 1988. Sampling pasture carabids with
pitfalls: evaluation of species richness and precision. Med. Fac.
Landbouww. Rijksuniv. Gent 53, 11091117.
Duelli, P., 1994. Rote Listen der gefhrdeten Tierarten der Schweiz.
Bundesamt fr Umwelt Wald und Landschaft. BUWAL-Reihe
Rote Listen. EDMZ, Bern.
Duelli, P., Obrist, M.K., 1998. In search of the best correlates
for local organismal biodiversity in cultivated areas. Biodivers.
Conserv. 7, 297309.

P. Duelli, M.K. Obrist / Agriculture, Ecosystems and Environment 98 (2003) 8798

Duelli, P., Obrist, M.K., Schmatz, D.R., 1999. Biodiversity
evaluation in agricultural landscapes: above-ground insects.
Agric. Ecosyst. Environ. 74, 3364.
Duffey, E., 1974. Comparative sampling methods for grassland
spiders. Bull. Br. Arach. Soc. 3, 3437.
Dufrene, M., Legendre, P., 1997. Species assemblages and indicator
species: the need for a flexible asymmetrical approach. Ecol.
Monogr. 67, 345366.
European Community, 1997. Agenda 2000, vol. L, For a Stronger
and Wider EU. Office for Official Publications of the European
Communities, Luxembourg.
Faith, D.P., Walker, P.A., 1996. Environmental diversity: on the
best-possible use of surrogate data for assessing the relative
biodiversity of sets of areas. Biodivers. Conserv. 5, 399415.
Flckiger, P.F., 1999. Der Beitrag von Waldrandstrukturen
zur regionalen Biodiversitt. Doctoral Thesis. PhilosophischNaturwissenschaftliche Fakultt, Universitt Basel, Basel.
Foster, G.N., Blake, S., Downie, I.S., McCracken, D.I., Ribera,
I., Eyere, M.D., Garside, A., 1997. Biodiversity in Agriculture.
Beetles in Adversity? BCPC Symposium Proceedings No. 69:
Biodiversity and Conservation in Agriculture, pp. 5363.
Gaston, K.J. (Ed.), 1996a. Biodiversity: A Biology of Numbers
and Difference. Blackwell Scientific Publications, London.
Gaston, K.J. (Ed.), 1996b. Species Richness: Measure and
Measurement. Biodiversity: A Biology of Numbers and
Difference. Blackwell Scientific Publications, London.
Gaston, K.J. (Ed.), 1996c. What is Biodiversity? Biodiversity:
A Biology of Numbers and Difference. Blackwell Scientific
Publications, London.
Gaston, K.J., Williams, P.H., 1993. Mapping the worlds species
the higher taxon approach. Biodiv. Lett. 1.
Greenslade, P., 1997. Are Collembola useful as indicators of the
conservation value of native grassland? Pedobiologia 41, 215
220.
Halsall, N.B., Wratten, S.D., 1988. The efficiency of pitfall trapping
for polyphagous predatory Carabidae. Ecol. Entomol. 13, 293
299.
Harte, J., Kinzig, A., 1997. On the implications of speciesarea
relationships for endemism, spatial turnover, and food web
patterns. Oikos 80, 417427.
Hellawell, J.M., 1986. Biological Indicators of Freshwater
Pollution and Environmental Management. Elsevier, London.
IUCN, 1994. IUCN Red List Categories. Prepared by IUCN
Species Survival Commission. IUCN, Gland.
Johnson, S.P., 1993. The Earth Summit: The United Nations
Conference on Environment and Development (UNCED).
Graham and Trotman, London.
Kaennel, M., 1998. Biodiversity: a diversity in definition. In:
Bachmann, P., Khl, M., Pivinen, R. (Eds.), Assessment of
Biodiversity for Improved Forest Planning. Kluwer Academic
Publishers, Dordrecht.
Kleijn, D., Berendse, F., Smit, R., Gilissen, N., 2001. Agrienvironment schemes do not effectively protect biodiversity in
Dutch agricultural landscapes. Nature 413, 723725.
Kromp, B., Pflgel, G., Hradetzky, R.I.J., 1995. Estimating
beneficial arthropod densities using emergence traps, pitfall
traps and the flooding method in organic fields (Vienna,

97

Austria). In: Toft, S., Riedel, W. (Eds.), Arthropod Natural

Enemies in Arable Land, vol. 70. Acta Jutlandica, Aarhus
University Press, Denmark, Aarhus, Denmark, pp. 87100.
Lindenmayer, D.B., 1999. Future directions for biodiversity
conservation in managed forests: indicator species, impact
studies and monitoring programs. For. Ecol. Manage. 115, 277
287.
Magurran, A.E., 1988. Ecological Diversity and its Measurement.
Croom Helm Limited, London.
McGeoch, M.A., 1998. The selection, testing and application of
terrestrial insects as bioindicators. Biol. Rev. 73, 181201.
Meffe, G.K., Carroll, C.R., 1994. Principles of Conservation
Biology. Sinauer, Sunderland.
Moser, D., Zechmeister, H.G., Plutzar, C., Sauberer, N., Wrbka, T.,
Grabherr, G., 2002. Landscape patch shape complexity as an
effective measure for plant species richness in rural landscapes.
Landscape Ecol. 17, 657669.
Mossakowski, D., Paje, F., 1985. Ein Bewertungsverfahren von
Raumeinheiten an Hand der Carabidenbestnde. Verh. Ges.
kol. Bremen 13, 747750.
Niemel, J., 2000. Biodiversity monitoring for decision-making.
Ann. Zool. Fenn. 37, 307317.
Noss, R.F., 1990. Indicators for monitoring biodiversity: a
hierarchical approach. Conserv. Biol. 4, 355364.
Noss, R.F., 1999. Assessing and monitoring forest biodiversity: a
suggested framework and indicators. For. Ecol. Manage. 115,
135146.
Oliver, I., Beattie, A.J., 1996. Invertebrate morphospecies as
surrogates for species: a case study. Conserv. Biol. 10, 99109.
Ovenden, G.N., Swash, A.R.H., Smallshire, D., 1998. Agrienvironment schemes and their contribution to the conservation
of biodiversity in England. J. Appl. Ecol. 35, 955960.
Pimm, S.L., 1991. The balance of nature? Ecological Issues in
the Conservation of Species and Communities. University of
Chicago Press, Chicago.
Pollard, E., Yates, T.J., 1993. Monitoring Butterflies for Ecology
and Conservation. Chapman & Hall, London.
Prendergast, J.R., 1997. Species richness covariance in higher taxa:
empirical tests of the biodiversity indicator concept. Ecography
20, 210216.
Prendergast, J.R., Quinn, R.M., Lawton, J.H., Eversham, B.C.,
Gibbons, D.W., 1993. Rare species, the coincidence of diversity
hotspots and conservation strategies. Nature 365, 335337.
Reid, W.V., McNeely, J.A., Tunstall, D.B., Bryant, D.A., Winograd,
M., 1993. Biodiversity Indicators for Policy-makers. WRI and
IUCN, Washington, DC and Gland.
Rykken, J.J., Capen, D.E., Mahabir, S.P., 1997. Ground beetles
as indicators of land type diversity in the Green Mountains of
Vermont. Conserv. Biol. 11, 522530.
Schlpfer, F., Schmid, B., Seidl, I., 1999. Expert estimates about
effects of biodiversity on ecosystem processes and services.
Oikos 84, 346352.
Spellerberg, I.F., 1991. Monitor Ecological Change. Cambridge
University Press, Cambridge.
Vane-Wright, R.I., Humphries, C.J., Williams, P.H., 1991. What to
protect?systematics and the agony of choice. Biol. Conserv.
55, 235254.

98

P. Duelli, M.K. Obrist / Agriculture, Ecosystems and Environment 98 (2003) 8798

Wagner, H.H., Wildi, O., Ewald, K.C., 2000. Additive partitioning

of plant species diversity in an agricultural mosaic landscape.
Landscape Ecol. 15, 219227.
Wascher, D.W., 2000. Agri-environmental indicators for
sustainable agriculture in Europe. ECNC Technical Report
Series, Tilburg.

Wratten, S.D., Van Emden, H.F., 1995. Habitat management for

enhanced activity of natural enemies of insect pests. In: Glen,
D.M., Greaves, M.P., Anderson, H.M. (Eds.), Ecology and
Integrated Farming Systems. Wiley, Bristol.