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Biodiversity: Function and Assessment in Agricultural Areas: A Review

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Agron. Sustain. Dev. 25 (2005) 1–15 1
© INRA, EDP Sciences, 2005
DOI: 10.1051/agro:2004049
Review article

Biodiversity: function and assessment in agricultural areas. A review

Boris CLERGUEa*, Bernard AMIAUDa, Frank PERVANCHONb, Françoise LASSERRE-JOULINa, Sylvain PLANTUREUXa

a
UMR INRA-ENSAIA-INPL, Agronomie et Environnement, 2 avenue de la Forêt de Haye, 54505 Vandoeuvre-lès-Nancy, France
b Trame, 9 rue de La Baume, 75008 Paris, France

(Received 25 October 2004; accepted 25 October 2004)

Abstract – Biodiversity has become a central concept in agronomical research since the Rio de Janeiro summit in 1992. Agricultural areas
include a unique biological diversity which is the basis of human activities. Conservation of this biodiversity in agricultural and protected areas
is therefore fundamental and requires an operational approach. Biodiversity is a complex entity which can be spread over several levels (genes,
species, ecosystems and ecological processes) and can be related to three main functions: (i) patrimonial functions, (ii) agronomical functions
and (iii) ecological functions. The patrimonial function concerns conservation of the landscape aesthetic and threatened species. The
biodiversity function according to relationships with agricultural activities describes resistance to biotic and abiotic stress, and the production
of cultivated ecosystems. Biodiversity is also involved in ecological functioning through the existence of special habitats with particular species.
The relevance of assessment tools is required in order to understand and evaluate the impact of farm practices on the different compartments
of biodiversity on the patch scale to the landscape scale. Different methods, such as direct measurements with biodiversity indexes, biotic
indicators and models are described and their suitability and limits are discussed.

biodiversity / assessment / landscape / indicators / sustainable agriculture

1. INTRODUCTION scapes [12, 74, 144], and in natural areas [10, 25, 75, 97, 103],
but these specific cases will not be developed in this paper.
Biodiversity has become a central concept in agronomical Protection of biodiversity requires assessment methods in
research since the Rio de Janeiro summit in 1992 [24]. This order to understand disturbance effects on biodiversity, moni-
event indicated a world consciousness of the importance of bio- toring its state and the relevance of agri-environmental meas-
diversity protection for sustainable development [14]. Biodi- ures. However, biodiversity is a very complex entity with the
versity protection can be motivated by pragmatic reasons. For interaction of different scales (species, community, ecosystem
example, biodiversity represents a potential reserve of new and landscape). Biodiversity is not only a concept which
compounds for medicine, interesting genes for plant breeding expresses the “variety of life” but is also a socio-political con-
and services for agriculture [2, 37, 102, 105]. Biodiversity is struction and an ecological measurable entity [44]. Thus, oper-
also considered as mankind's heritage and human beings cannot ational definitions of biodiversity are necessary to determine
research directives, biological conservation measures and
decide on the existence or not of a species [20].
make environmental policies.
Considering the role of agriculture in the preservation of bio-
For instance, Noss [92] has described biodiversity by a hier-
diversity appears to be a key issue. For better biodiversity con- archic approach based on the distinction between “composi-
servation on the large scale of territories, knowledge and tion”, “structure” and “function” applied on different scales
creation of conservation tools are necessary not only in pro- (Fig. 1). The work of Noss has been a key reference in ecolog-
tected and restricted areas but also in agricultural areas. On the ical studies for monitoring biodiversity. Biodiversity “compo-
European Community scale, agricultural areas are more signif- sition” is an inventory of characteristics, such as biomass
icant (44%) than protected areas, which represent less than 5% production, species abundance, presence of threatened species
[114]. In addition, mosaic landscapes based on a mixture of or habitat proportions. Biodiversity “structure” is the organi-
agricultural and semi-natural areas represent a particular sation of biodiversity components and the relations between
reserve of biodiversity. Finally, biodiversity preservation in them. These components take into account structural data about
agricultural lands produces new challenges: to conciliate pro- population (sex, ratio, morphological variability, etc.), habitat
duction necessities with respect for the environment [2, 16, 105, (slope, foliage density, etc.) and landscape (connectivity, frag-
141]. Additional studies have been conducted in urban land- mentation, patch size, etc.).

* Corresponding author: [email protected]

2 B. Clergue et al.

Figure 1. Compositional, structural and functional biodiversity, shown as interconnected spheres ([92] modified).

The third level, biodiversity “function”, is the whole of par- Noss [92] proposed a hierarchic approach involving the con-
ticular ecological processes, such as demographic processes or cept of the term “function” of biodiversity. He used it to define
population dynamics and genetics. The functional groups the- all the processes which occur on the different scales: gene, spe-
ory is another operational approach which links biodiversity to cies-population, community-ecosystem and regional land-
ecosystem processes. Each functional group is related to an scape (Fig. 1). Nutrient cycling and energy flow are especially
ecosystem process such as organic matter decomposition or taken into account. However, Noss has focused on the ecolog-
nitrogen mineralisation [71, 76, 78, 79, 137, 142]. An ecosys- ical functions of biodiversity.
tem process becomes an ecosystem service according to a On the contrary, Peeters et al. [106] have expressed biodi-
human point of view. For example, biomass production of versity functions essentially according to relationships with
grassland ecosystem represents forage production for cattle. agricultural activities. Biodiversity is split into three parts:
Ecosystem services therefore form a basis for human life [120]. (i) agricultural biodiversity, (ii) para-agricultural biodiversity, and
Agricultural areas contain a unique and useful biodiversity (iii) extra-agricultural biodiversity. “Agricultural biodiversity”
which results from farm management. In order to promote sus- represents the variety of life directly used for farming produc-
tainable agriculture, knowledge and conservation of biodiver- tion. It involves animal and plant species, races and varieties.
sity need clarification on two points: (i) the biodiversity “Para-agricultural biodiversity” (also called “functional biodi-
concept, especially the integration of the benefits of biodiver- versity”) is the variety of life indirectly used for farming produc-
sity, and (ii) assessment methods used to evaluate and monitor tion such as soil fauna, auxiliary fauna, pollinators, grassland
biodiversity. plant diversity and more generally ecosystem services. “Extra-
agricultural biodiversity” represents biodiversity in production
areas which does not contribute to production. These are mainly
2. BIODIVERSITY AS A MULTI-FUNCTION particular species, especially endangered species (orchids, but-
Biodiversity is a complex entity which can be spread over terflies, great mammals, etc.).
several levels. Authors have given, therefore, different ways to Gurr et al. [53] also reviewed benefits of biodiversity for
define biodiversity as a sum of several functions. agricultural production such as pest management which favours
Biodiversity: function and assessment in agricultural areas 3

Figure 2. The hierarchy of scale for potential benefits of multi-functional agricultural biodiversity ([53] modified).

enhancement of natural enemies. They also proposed a hierar- 2.1. Patrimonial functions
chy of biodiversity benefits based on the different scales of bio-
The biodiversity of a site is related to history, and thus con-
diversity. For instance, pest management is obtained on the
stitutes a patrimony. This patrimony is a common heritage with
patch scale by changing practices, and on the landscape scale
both a natural or biological and cultural patrimony. More often
by the integration of non-crop vegetation which increases
than not, these two patrimonies are inter-related. Patrimonial
diversity (Fig. 2).
functions are present on different scales: on a landscape scale,
The definitions of Peeters et al. [106] and Gurr et al. [53] biodiversity contributes to aesthetic, and on a smaller scale, to
show that agricultural activities are strongly linked to biodiversity particular habitat, species and a genetic patrimony.
components. Paoletti et al. [101, 102] previously highlighted,
by an inventory of biodiversity components, that agricultural pro- 2.1.1. Aesthetic function
duction is based on biodiversity.
Duelli and Obrist [37] have reviewed the different aspects Biodiversity contributes to the aesthetic value of the land-
of biodiversity with both an ecological approach and an agro- scape; this is also called visual or scenic quality. On the land-
nomical approach. They separated these aspects into three parts scape scale, patrimony therefore has an aesthetic function. The
which motivated preservation and studies on biodiversity: (i) aesthetic function creates a feeling of identity for residents, and
conservation (threatened species protection), (ii) biological a recreation object for tourists.
control (antagonist species diversity), and (iii) resilience (eco- For the European Landscape Convention [39], “landscape
system processes). The Duelli and Obrist [37] approach means an area as perceived by people, whose character is the
presents a biodiversity concept which manages several func- result of the action and interaction of natural and/or human fac-
tions or ecological services. The three parts may be, respec- tors”. Thus, the aesthetic value includes natural and cultural
tively, extended to three main functions: patrimonial functions, elements of the landscape. Steiner [133] stated that “usually, a
agronomical functions and ecological functions. The approach landscape is that portion of land or territory which the eye can
of Gurr et al. [53] highlights the necessity of taking into account comprehend in a single view, including all its natural charac-
the action of these functions on several scales. teristics”.
4 B. Clergue et al.

Figure 3. Aesthetic perception of landscape and levels of aesthetic cognition ([91] modified).

Aesthetic values result from the relationship between the Analysis of these perceptions showed that humans have “a
landscape and an observer. Observation provokes in the natural attraction for diversity which is a source of pleasure, sat-
observer a visual perception that is associated with thoughts isfaction, or happiness” [143]. A preserved natural landscape
and feelings. The NGO European Academy for the Culture of provoked the same feelings [4, 99, 100].
Landscape [112] attributed the first landscape description in Biodiversity gives origin to an aesthetic function on the land-
Europe to the Italian poet Francesco Petrarca (1304–1374). At scape scale but Nohl [91] showed another complexity level: “If
the “Mont Ventoux” summit (Vaucluse, France), Petrarca one compares the appearance of today’s landscape with that
related an observation experiment. He observed a panorama, of premodern and early modern time, one recognizes that the
the nature which he perceived as a totality: a landscape. This landscape did not only lose its wealth of elements but also its
observation is considered as an aesthetic perception. Nohl [91] sense of unity which gave form to that variety”. Landscape aes-
explained precisely the aesthetic perception process (Fig. 3). thetic is thus a result of the diversity of elements and their cohe-
He differentiated several levels of perception; especially sion or organisation.
between, on the one hand, results of observation and interpre- An agricultural landscape is a complex assemblage of agri-
tation, and on the other hand, objective (narrative aspect) and cultural, semi-natural and rural areas [114] and constitutes a
subjective approaches (poetic aspect). However, Weinstoerffer mosaic of many elements. Heterogeneity is a parameter that
and Girardin (2000) underlined that the first landscape studies helps with understanding the organisation of mosaic land-
which began in the seventies had used only an “objective pole” scapes. Heterogeneity is the diversity of landscape elements
with a descriptive science. This point of view includes natural- (patches) and the complexity of their spatial relationships.
istic approaches and agro-ecological approaches: the first Fragmentation and connectivity are measures that characterise
approach studies landscape structure by inventories of the char- landscape heterogeneity. Fragmentation gives information on
acteristics (vegetation, relief, soil, geology and climate), the the spatial organisation of a habitat by patch size, while con-
second approach is a taking into account of agricultural and nectivity describes the spatial relationships between patches
semi-natural elements [143]. Since the earlier work of Shafer [18]. These spatial parameters permit the understanding of the
et al. [122], more recent landscape studies have included both preferences of the observers.
objective and subjective approaches (See, for example: [5, 11, A landscape can offer some aesthetic qualities to the inhab-
13, 99, 100, 141]). Colquhun [27] and Bosshard [11] pointed itant or tourist but landscapes contain other elements which the
out that subjective approaches have the same scientific rigour public prefer, and ecological characteristics. These elements
as the objective approaches. This conviction is based on the also have a patrimonial value.
works of the German poet and scientist Goethe (1749–1832)
2.1.2. Patrimonial function on other scales
in botany (Plant metamorphosis, 1789) and optics (Theory of
colours, 1810). The American philosopher Emerson (1803– Biodiversity can also have a patrimonial interest that is more
1882) also sustained this point of view, especially in his essay due to its historical and socio-cultural context than its visual
Nature (1836). quality. On small scales, patrimony includes habitats, species
In addition, Schüpbach [121] underlined the fact that the and genetic patrimony.
tourist industry and landscape protection organisations (see, for On the European scale, the Directive 92/43/EEC (Habitats Direc-
example, [130]) use aesthetic perceptions in order to raise the tive) on the conservation of natural habitats, wild fauna and flora
public conscience of the landscape. has established the European ecological network Natura 2000.
Biodiversity: function and assessment in agricultural areas 5

Natura 2000 aims to maintain vital elements of the natural pat- in host species (bottom-up effect), while on the other hand, an
rimony. These natural areas are also related to economic activities increased diversity of predators controls pest populations ( top-
(agro-forest production and rural tourism), hobbies (hunting, down effect) [53].
fishing, outdoor hobbies, etc.), and contribute to maintaining Arthropods and birds are the main auxiliaries. The presence
the quality of rural life. of these useful fauna is strong correlated with semi-natural
Following the Convention on Biological Diversity the sig- areas [60, 62].
natory states must contribute to species conservation: this is In the case of the vole, their outbreaks are strongly correlated
especially the case for threatened species [23] that belong to with land cover. High values of the meadow/crop area ratio
natural patrimony. Based on the Red list concept of the World indicate a high risk of outbreaks [48]. Millan de la Peña et al.
Conservation Union (IUCN), threatened species are registered [84] showed that habitat diversity (connectivity vs openness)
for particular areas. allowed a diversity of rodents and thus decreased the generalist
So-called flagship species are used to increase public interest species.
and attract funding for ecological matters [21]. These species A high species diversity within a community enhances its
are often threatened species. Flagship species can be a plant resistance to invasion of alien species. The works of Levine
(orchids, etc.) or an animal (butterflies, eagle, bear, wolf, etc.) et al. [73] and Shea and Chesson [124] reviewed the different
with sometimes a cynegetic value (partridge, hare, etc.). Flag- studies examining this theory. The majority of studies were car-
ship species belong, therefore, to cultural and natural patri- ried out on plants in grasslands. In addition, they indicated that
mony. the most diverse natural communities were the most frequently
Pervanchon [109], owing to a request from French Regional invaded.
Natural Park managers, found that rarity characterises patrimo-
nial value in permanent meadows. The rarity criteria of a spe- 2.2.1.2. Disease and nematode control
cies is based on the rarity index of Janssens [58]. Pervanchon
[109] proposed a definition of a patrimonial species which cov- Crop protection against diseases is an important part of the
ers the concepts of both flagship and threatened species. A pat- farming budget. The diversity of plant and soil organisms may
rimonial species is “a rare or threatened species which needs help to control pathogenic microorganisms, especially fungi [1,
local management and which may be a flagship species and 117] and plant-parasitic nematodes [147]. In addition, disease
may have cultural importance” [109]. The Patrimoniality con- control by biodiversity helps to reduce pesticide inputs. Crop
cept is used in ecological studies in this sense (see, for example rotation (diversity in time) and the diversity of organisms in
[43, 70, 104]). organic amendments are management practices which increase
On the genetic scale, natural and agronomic species have a soil biological activity. For example, wheat diseases can be
genetic patrimonial value. Genetic diversity allows species per- reduced by cultivar blending [57], while compost amendment
enniality and species adaptation to environment changes. In increases soil biological activity and controls turfgrass diseases
addition, knowledge of genetic diversity gives measures for the [90].
breeding and conservation of plants [8] and animals [31]. This The presence of hedgerows limits propagation of some dis-
may also help in conservation of wild species and forest man- eases (e.g. Oidium) by reducing wind, but can induces other
agement [45]. Conservation of genetic resources has been com- disease in shaded and wet areas.
mitted internationally especially via the Global Programme for
the Management of Farm Animal Genetic Resources (FAO) 2.2.2. Abiotic stress resistance
and the Global Plan of Action for the Conservation and Sus-
tainable Utilisation of Plant Genetic Resources for Food and 2.2.2.1. Biodiversity benefits for soil properties
Agriculture (FAO, Leipzig, June 1996).
Soil biota regulates many ecological processes: litter decom-
position, nutrient cycling, pathogen control, mineral weather-
2.2. Agronomical functions ing, etc. From an agronomical point of view, the processes of
decomposition, immobilisation and mineralisation liberate
Agricultural production can be considered as linked to dif- nutrient elements according to plant growth [102]. Thus, losses
ferent biodiversity functions. This biodiversity may control by leaching are limited as plants absorb necessary elements.
crop and meadow stresses (pests, diseases, dryness, deficien- Moreover, symbiotic associations with mycorhizal fungi increase
cies, etc.) and support essential plant functions such as repro- nutrient availability, e.g. of phosphorus, and increase plant
duction via pollinators. Biodiversity acts on agronomic water uptake. Mycorhizal symbiosis is therefore important for
parameters on different scales: on the patch scale, on the matrix plant growth. It is present in all plant species except in the
scale, which includes semi-natural boundaries (bark, ditches Brassicaceae family [136]. Soil biota can also weather minerals by
and hedgerow), and on the landscape scale with hedgerow webs production of chelating agents and catalyses redox reactions [2].
(connectivity and fragmentation) or forest areas.
The diversity of soil organisms and their abundance are
2.2.1. Biotic stress resistance involved with processes that affect soil structure. Crossley et al.
[30] defined the influence of each organism category (micro-
2.2.1.1. Pest control flora, microfauna, mesofauna and macrofauna) on each soil
structure. These organisms act as much on particle aggregation
Biodiversity can control pest population by two mecha- and humification as porosity creation and organic-mineral
nisms: on the one hand, floristic diversity implies a decrease phase melting. Soil structuring increases the growth of plant
6 B. Clergue et al.

rate of bumblebees [7]. These pollinators are more efficient

than the honeybee in unfavourable climatic conditions (cold
and overcast sky). Pollination allows fecundation of ento-
mophile plants, especially dicotyledonous plants [85]. For crop
production such as rape or sunflower, pollination directly
affects a yield component, the seed number. In grasslands, pol-
lination allows reproduction of entomophile species. The con-
Figure 4. Duality effect in a bocage patch ([26] modified). With H: sequences of pollination are that sexual reproduction maintains
total height of bank and vegetation, about 10 m in the Western France genetic diversity that vegetative reproduction cannot. This
bocage. genetic diversity increases adaptation to environmental stress.

2.2.4. Crop and animal production

roots, anchorage and fluid circulation (air and soil solution). Regulation of biotic and abiotic stress and pollination are
Soil structuring also increases penetration of rain water. one of several agronomical functions of biodiversity. Aggre-
On the landscape scale, wind erosion is a soil quality matter gation of these functions gives biodiversity effects on crop pro-
often neglected. In open-field landscape without plant cover, a duction. However, other factors are linked to crop production.
low-speed wind (4 m·s–1) may provoke soil erosion of small For example, the species forage value and species richness of
particles. Humus is mainly present in the soil upper layers and a grassland is correlated to forage production. In addition, spe-
can be taken away by wind. The presence of hedgerow limits cies diversity influences crop and forage production and also
wind speed and thus soil erosion. The carbon value of hedgerow quality of dairy products (milk and cheese).
board soils is the highest [26]. The whole hedgerow/bank/ditch In the case of hedgerow effects on production components,
creates lateral discontinuity which limits water and particle lat- effects must be seen on the patch and landscape scales. In a
eral transfers. This process reduces soil erosion by hydric trans- bocage grid, the culture response of hedgerow effects shows a
fer due to, for example, superficial runoff and hypodermic flow. spatial heterogeneity. For the climatic effect, the centre of the
In the soils affected by this erosion, especially for sloped patch is controlled by regional context and the boundaries by
patches, soil is always deeper in bank uphill slopes than in local context (Fig. 2).
downhill slopes [118]. Erosion modifies the quantity of the dif-
The presence of hedgerow also influences growth rhythms
ferent soil elements (sand, silt and clay) which have conse-
and yield. A spatial heterogeneity is observed for precocity and
quences both on soil structure and fertility. Moreover, the
presence of the whole hedgerow/bank/ditch allows a better yield in strips which are perpendicular to dominant winds.
infiltration and thus a greater water stock than in open-field Favourable areas spread on 2 to 6 times the hedgerow height.
landscape [118]. Diversity and organisation of landscape ele- On the landscape scale, bocage increases crop precocity in
ments influence soil water availability, and thus plant growth comparison with open areas. However, effects on yield are
and yield. often contradictory because there is also an interaction between
the plant cultivar and its area.
2.2.2.2. Microclimate Protected Designation of Origin cheeses are characterised
by typical sensory properties (taste, odour and texture). In order
Microclimates are strongly connected to regional climate
but they are also linked to local geomorphology (slope, aspect to understand links between cheese properties and a geograph-
and relief which reduces winds) and human activities. The diver- ical area, a “Terroir”, several chemical studies were carried out.
sity of landscape elements, such as hedgerows, acts on climatic These studies have demonstrated clearly that some odour-
parameters (wind speed and Potential Evapo-Transpiration). active compounds (aldehyde, ester and terpenoid compounds)
found in grassland species can be transferred to the milk and
On the patch and regional scales, bocage structure decreases
cheese [15, 22, 29, 59, 139]. Dairy product quality is therefore
wind speed by about 30 to 50 per cent [26]. The effect of bocage
related to floristic diversity. A diversity of these compounds is
on the patch microclimate is due to landscape structure both on
produced by plant species adapted to a particular habitat (high
the patch and regional scales (Fig. 4). This wind speed diminu-
tion decreases Potential Evapo-Transpiration by 4 to 6 per cent. mountain pasture, extensive practices), especially dicotyledo-
Although several effects of bocage are known, effects on farm nous species such as Achillea sp., Meum sp., Thymus sp. or
production are difficult to measure. In a bocage, the day air tem- Geranium sp. [35, 82]. Odour-active compounds form a fin-
peratures are higher and the night temperatures are lower in gerprint of dairy products and may be used for traceability
comparison with open areas. [139].
In addition, pluviometry is higher in the presence of a hedge-
row web than in forest areas [129]. 2.3. Ecological functions

2.2.3. Pollination According to Duelli and Obrist [37] biodiversity implied in

ecological functioning is involved in some services for agricul-
In addition to the domestic honeybee (Apis mellifera), pol- tural activities but diversity is also related to some ecological
lination is done by a diversity of insects (bumblebees, wild aspects. Biodiversity (i) creates typical habitats, (ii) includes
bees, etc.). A high diversity of habitat increases the occupancy particular species, and (iii) is related to ecosystem functioning.
Biodiversity: function and assessment in agricultural areas 7

2.3.1. Habitats (microbes, protozoa, vascular plants, nematodes, arthropods,

annelids and vertebrates). In the face of biodiversity losses,
Habitat is the place where an organism or population occurs ecologists have begun investigating this damage to ecosystem
naturally. The habitat of a (plant or animal) species is its “place functioning. The relationship between diversity and ecosystem
of residence” [93], which means the area to which it is adapted stability has been the most studied and debated since the 50s [78].
and which it is able to occupy. A habitat type includes specific
factors (ecological conditions) which allow the species to sur- According to several authors [32, 33, 78], biodiversity is
vive and to reproduce successfully. If the habitat quality linked to ecosystem processes: matter, energy and nutrient cycles.
changes (e.g. due to anthropogenic impact) or the ecological Although these relationships are known, especially through the
requirements of the species change, it is forced to retreat from food web concept, their understanding is limited. The majority
its place of residence [17]. of studies and models are based on the relationship between
diversity and stability (community and ecosystem process sta-
In addition, in agricultural areas, the presence of extensive
practices allows formation of habitats with a specific biodiver- bility) [71, 76, 79, 137], but are often led on a single trophic
sity [19]. level and on small scales.
Abandonment of these practices causes species impoverish- Loreau et al. [78] supposed that the first theoretical studies
ment. In order to prevent this phenomenon, Environmental applied the conventional wisdom (don’t put all your eggs into
Sensitive Areas (ESAs) have been established by the CAP (CE one basket). In this vision, diversity of pathways provides sta-
797/85). European Directives Habitats (92/43/EEC) and Birds bility. Below a threshold of biodiversity loss, stability is there-
(79/409/EEC) allowed establishment of the Natura 2000 net- fore broken and involves a cascade reaction of species loss, and
work. Natura 2000 areas are specific habitats or landscapes that the ecosystem is endangered. However, many results have
are selected for their biological diversity, and the presence of shown that diversity is related to different stability properties.
specific or threatened habitats and species. According to a plant A greater diversity is not always favourable to community sta-
diversity point of view, these habitats are also characterised by bility and process stability. One of the hypotheses about eco-
a particular phyto-sociological community [87]. system functioning involves idiosyncratic process. Species
make different contributions to ecosystems depending on cer-
2.3.2. Specific species tain conditions (e.g. community composition, etc.) [88].
Griffiths et al. [50, 51] illustrated another complexity of eco-
Biodiversity includes particular species in relationships with system functioning: the redundancy of functional groups by a
ecosystem processes. The literature has given several names to
stress-on-stress experiment. The first stress is a disturbance
groups of species that are related to certain ecological func-
applied to soil samples. It decreases the biodiversity but not all
tions. These different names may be cross-checked.
ecosystem processes. For example, organic matter decomposi-
Indicator species are species which are used for many rea- tion may be greater than before the disturbance. At the second
sons such as an indirect measurement of the health of the eco- stress, the ecosystem processes decrease greatly. These exper-
system (condition indicator), identification of an area of high iments showed that the first stress has affected biodiversity sta-
species richness (biodiversity indicator) or as markers of pop- bility while the second stress has affected process stability.
ulation size for other species (population/guild indicator) [68]. Hence, ecosystem processes are not linked directly to biodiver-
Keystone species have an important ecological function sity. As there is a functional redundancy in the soil community,
either in sustaining ecosystem functions or in sustaining pop- a distinction must be observed between community and proc-
ulations of other species. For example, barrage building by bea- esses.
vers creates a wetland, while a cavity dug by a woodpecker may
Loreau et al. [78] reviewed the different stability properties
be used for nesting of other bird species. Keystone species are
such as resilience or resistance. Resilience is “a measure of
therefore precious tools for ecosystem conservation [125].
speed at which a system returns to [a stable] state after a per-
However, they are not the panacea. Not all ecosystems have
keystone species. According to Bengtsson [9], ecosystem turbation”, while resistance is “a measure of ability of a system
engineers (earthworm in the soil, Daphnia in the aquatic food to maintain its original state in the face of an external disruptive
web) are like keystone species. force”. Resistance is the stability propriety of the ecosystem,
for example, against invasion by non-native species.
Umbrella species are used to locate the edges of a conser-
vation area. These are species that need a large area to survive. Raffaelli et al. [116] suggested there was an urgent need to
Conservation of this area provides protection for co-existing orient modelling on biogeochemical cycling and therefore
species. research on a larger scale. “There have been very few attempts
to explore the effect of biodiversity on the functioning of full
Using terms such as focal species and surrogate species
ecosystems comprising higher trophic levels, decomposers and
provoked a semantic and scientific polemic [3, 21]. However,
nutrient cycling and none as yet have considered stability
focal species and surrogate species are sometimes used to des-
ignate indicator species or other particular species. explicitly” [78]. Ecological studies consider two ecosystem
divisions: above-ground/below-ground, either plant-herbiv-
2.3.3. Ecosystem processes and nutrient cycling ore-predator or soil community. However, studies are often
limited to small-scale soil- and plant-associated ecosystems
Many studies [32, 33, 55, 67, 127, 128] have shown clearly (see, for example [33, 127]) and do not include mammals or
that farming practices (fertilisation, pesticides and tillage) birds. A multi-trophic approach is proposed to examine the eco-
affect the population size and the dynamics of several groups system process holistically.
8 B. Clergue et al.

Table I. Indices used for biodiversity description.

Indices Formula Abbreviations

1 - Species richness (S) S = ni ni = species i
2 - Alpha diversity (Hα’) Hα’ = pi log2 pi pi = frequency of the species i
[123]
3 - Beta diversity (Hβ’) Hβ’ = S/m – 1 S = species number (all samples)
[145] m = average number of species per sample
4 - Evenness (J) E = H’/log2 S
[113]
5 - Dominance (D) D = Σ pi²
[126]
6 - Rarity index (IR) IR = Σ Ci/S Ci = rarity coefficient of the species i (from 1 to 13)
[58, 109]

Many authors [71, 131, 137] have given information on the viduals as the other, the Shannon-Weaver Index gives the same
relationship between plant diversity and above-ground biomass value [115]. The Shannon-Weaver index is also used on a land-
of grassland. Biomass production is greater with species-rich scape scale to evaluate diversity of landscape elements [41,
communities than the most productive monoculture. 89].The Shannon-Weaver index is used as an alpha-diversity
According to de Ruiter et al. [32], future research must focus index, because it gives information at species level.
on these links between biodiversity stability and process sta- Beta diversity could be defined as the difference in species
bility. Knowledge of these key properties will allow an under- composition between different communities. Beta diversity is
standing of the risks and effects of human disturbances. larger when there are fewer common species between different
communities [40]. The Whittaker index [145] could be the most
suitable among the beta diversity indices available. This is
3. BIODIVERSITY ASSESSMENT partly because it is easy to calculate and interpret [148]. It can
vary from 0 to 2.
Assessment tools are required to quantify and evaluate the Gamma or regional diversity is the total number of species
impact of agricultural activities on biodiversity. Many methods occurring in a system [81, 146].
have been proposed either by direct measurements on the site,
or by indirect measurements. Biodiversity studies are generally The evenness (J) [113] is a measure of abundance heteroge-
focused on one scale: either on the habitat, patch, or landscape neity between species in a community. This parameter can vary
scale. from 0 to 1. The maximum is obtained when all species have
the same abundance in the study site. Evenness is calculated
from Shannon-Weaver Index H.
3.1. Direct measurements of biodiversity
Touzard and Clément [138] used another parameter to
3.1.1. Simple indexes describe the diversity of the plant community: dominance. The
dominance (D) is measured from the inverse of the Simpson
The biodiversity definition provided by the Convention on diversity index [126]. When the dominance value is high, the
Biological Diversity takes composition (species, ecosystem, study site contains species with high abundance.
etc.) and structure (ecological process) into account. The tax- Janssens [58] used another parameter: the rarity index,
onomic richness is the first biodiversity measurement which which is an important parameter for biodiversity conservation.
gives the number of taxa (family, genus, species, variety and The rarity index may be used to give the patrimonial value of
ecotype) per unit area. This method is the most commonly used a study site [109]. Peeters et al. [106] proposed vulnerability
and represents the simplest expression of the diversity. Never- as a parameter which gives sensitivity of a taxon to extinction.
theless, the value of this criterion used alone is limited as spe-
cies number must be compared with a reference number for a These different methods show that there are many diversity
particular habitat. measures, but their suitability for use in different domains (soil
microflora, arthropods, plants and landscape elements) is not
Diversity indices are another method that uses the number always clear [40, 54, 148].
of taxa and their abundance (Tab. I). For example, communities
which have the same number of species may differ in the abun- 3.1.2. Biotic indicators
dance of each species.
The Shannon-Weaver index [123] (H) is the most commonly Direct measurement of biological diversity is frequently
used index. It gives information on community complexity and used for biodiversity studies. However, this measurement is
can vary from 0 (one species alone) to Log2S (where all species inconvenient due to high cost in terms of time and money, and
have the same abundance). However, this index is not sensitive the necessity for competence in species determination of very
to strength variation. In the case of two ecosystems which have diverse organisms (soil arthropods, plants, birds, etc.). In addition,
the same number of species, but one has twice as many indi- a sample represents a picture of biodiversity which changes all
Biodiversity: function and assessment in agricultural areas 9

the time (day/night, weather, season, years). Thus, scientists 3.2.2. Models predicting the threatening level
have tried to find indirect or surrogate measurements to deter- of natural resources
mine biodiversity. Instead of measure all the biodiversity, many
scientists hold the view that the dynamics of taxa gives a picture Potential impact models issued from German works of the
of the dynamics of biodiversity [34, 36, 37, 64, 65, 68, 80, 107, 1970s on ecological risks [42]. Impact means the level from
108]. An important contribution on using biotic indicators was which resources and/or ecological functions are threatened by
given in the special issue of Buchs [16]. Work of Duelli and harmful use to the ecosystem's health. Potential means that not
Obrist [36] suggested arthropod higher taxa were better biotic only are impact models in part based on field measurements,
indicators in terms of their ease of sampling and relationship but they are limited by the available data and approximation
inherent in modelling [42]. The model of Freyer et al. [42] pre-
with biodiversity. Assessment tools must be easily usable in
dicts the level of natural resources threatened due to human
order to be generalised for other case studies and to help deci-
activities such as pesticides and nitrogen inputs or mechanical
sion-makers involved with land-use management.
action (e.g. ploughing). This model can be applied on various
scales (see Tab. II for details).
3.2. Evaluation of biodiversity functions by models 3.2.3. Models based on life traits
Expert models are a novel modelling approach: they are
3.2.1. Modelling approaches considering live based only on the knowledge of some traits or biological char-
beings as dynamic systems acteristics of animal or plant species. There is no need for sta-
tistical analyses or empirical relations to elaborate such models,
Most of the models in ecology are based on a physical but only field observation and biometric measurements to build
approach of individual organisms, populations or ecosystems. a database. Once the database is built, expert models can predict
Live beings are not considered in all their complexity, but as very efficiently the species present in any ecosystem. These
dynamic systems which are determined by their state, as stated modelling approaches are the first concrete applications of
by physics [135]. For instance, it is the case of plant species functional groups theory based on life traits of plant species to
competition models [49, 119]. Gounot’s model is one of the predict animal or plant presence according to human activities
first theoretical ecosystem models. It is based on compartments and environmental factors [109].
which correspond to elements of the grassland such as the cat-
For instance, Pervanchon [109] developed an expert model
tle, the soil nutrients, the micro-organisms and the plant bio-
which predicts plant species' presence in any herbaceous eco-
mass. Matter and energy flows circulate between these system. This model is based on a database of 17 life traits or
compartments. Independent variables of the model are climate biological characteristics already identified in the literature for
and grassland management. VEGPOP 2 is a recent model based 2912 plant species. In order to predict the presence probability
on compartments, but it is operational thanks to the great of grassland plant species, the information of the traits and char-
improvement in scientific knowledge since the 1970s [119]. acteristics are aggregated with data on farming practices and
This model needs field experiments for plant species parame- environment factors by fuzzy logic associated with an expert
ters concerning physiology, resource allocation, nitrogen flow, system. With such a model, it is possible to predict a list of plant
flowering or population dynamics (see Tab. II for details). species with their patrimonial value, without realising floristic
VEGPOP 2 predicts the Shannon index, the plant biomass and sampling. The validation results of this model highlighted that
the vegetation spatial dynamics [119]. Numerous other models it is only necessary to improve the knowledge on live plant traits
are based on statistical analyses to quantify flows and compart- to improve the expert models [109].
ments (for instance, see [6]). Three kinds of analyses can be dis- The use of species traits to predict the presence of species
tinguished: the classical linear regression models, the linear according to human activities and environment factors was also
generalised relations, among which are the Gaussian, the bino- developed successfully to predict the presence of Syrphidae in
mial and the Poisson’s distributions [150], and the generalised any ecosystems [132]. If for plants, the scientific knowledge on
additive models. These models have up until now largely been biological traits still has big gaps, for Syrphidae, the traits are
used in ecology and they are well described elsewhere [52]. well detailed, and the lists of predicted species by the model
Besides these models, several models were inspired by the and observed species in ecosystems are very similar.
application of physics concepts. For instance, thermodynamics
[151] or automatics [83] can help to predict structure, dynamics
3.3. Surrogate measures of biodiversity:
and functioning of ecosystems.
landscape metrics
From the 1990s, the concomitance of the chaos theory, the
account of interactions between ecosystems and the improve- Ecologists have suspected for a long time that landscape
ment of computer performance was at the origin of numerous composition and landscape pattern are highly significant for
individual-based models in ecology [63]. Now, numerous dif- species diversity. However, the way in which species diversity
ferent models are available to explain or predict vegetation behaves in landscapes with different spatial arrangements is
structure and dynamics of ecosystems [38, 66, 69, 77, 98, 111, largely unexplained [134].
149]. None of these models evaluate the impact of farming One solution is to measure the elements that are related to bio-
practices on biodiversity, they only explain or predict vegeta- diversity. Landscape parameters may be correlated with species
tion structure or dynamics. diversity of many groups [60–62]. As a first step, a biodiversity
10 B. Clergue et al.

Table II. Comparison between several models and indicators which evaluate the impact of agricultural practices on biodiversity and agronomic
value of grasslands.

Characteristics of the models Model of Freyer et al. (2000) VEGPOP2 Model of Pervanchon, 2004
(Schippers & Joenje, 2002)
Model type Potential impact model Vegetation dynamic model Expert based model associated
with fuzzy logic
Model objectives Evaluation of human activities Evaluation of human activities’ Evaluation of human activities’
impacts on ecological functions impacts on plant diversity impacts on ecological and
(biomass, Shannon Index) agronomic function of any kind
of herbaceous surfaces
Targeted users Unknown Unknown Agricultural development
managers
Model structure
Parameters Human activities Pesticide inputs, ploughing, Hay cutting, N fertilisation, Hay cutting, grazing, water
Environment drainage, proportion of the herbicides management (drainage,
characteristics different activities on a irrigation), N and P fertilisation,
Studied species landscape scale calcareous inputs
Groundwater, soil, climate, Field perimeter Corine habitat, N and P soil
species, biotopes, landscape fertility, soil depth, pH, soil
and amenities. moisture, temperature
Unknown: species and biotopes Parameterisation for 4 plant 2912 European plant species
are together in the frame of species (Poa annua, Holcus
“environment protection”. lanatus, Anthoxantum odoratum,
Festuca ovina tenuifolia)
Model outputs Potential impact of human Shannon index, plant biomass Plant species lists (names)
activities on resources according to agricultural
practices
Input data Measured data, maps, statistical Fertilisation and disturbance Data given by farmers and maps
data levels
Scale Landscape, field, biotope or Field boundary Herbaceous surface
species (e.g.: grassland)
Calculation methodology
Statistical analysis None Yes: relations for spatial None
representation of plant
competition
Fuzzy logic Yes: exponential, logarithmic, None Equivalent of sigmoid functions
multilinear and linear functions
Other methods Type of equation Ratio and average of weighted None Choice of the minimum: plant
parameters. species presence probability is
the minimum of the all
probabilities.
Justification None / Precaution principle and limiting
factor theory
Application area Various biotopes or landscapes Areas where the 4 species are 4 French Regional Natural Parks
parameterised
Model generalisation Unknown Unknown Any herbaceous surfaces where
some of the 2912 species are
potentially present
Validation Unknown Satisfying results for some parts Satisfying results for French
of the model permanent grasslands; not tested
for other surfaces
Computerised version yes yes yes

parameter is studied in relation to spatial information. For the abundance of the species and spatial structure, this link is
example, data are searched for on the presence of a target spe- modelled and then validated. In the end, landscape data are only
cies in different habitats. After determining the link between necessary for monitoring the target species. Currently, a higher
Biodiversity: function and assessment in agricultural areas 11

diversity level on the landscape scale is used to predict a lower ecological indicators are predictive tools and help with deci-
diversity level (species richness, etc.) [60], and even if biodi- sion-making. These indicators use easily accessible data that
versity is linked to landscape parameters, there are no general can be collected by non-specialists However, the building of
models. indicators is dependent on scientific knowledge and indicators
For this reason, very many indicators based on spatial infor- are only suitable if they are validated for sensitivity and usa-
mation have been built. Piorr [114] reviewed agri-environmen- bility value [47].
tal indicators and landscape indicators used in the European
Union.
The OECD produced agri-environmental indicators which 4. CONCLUSION
were adjusted to the driving force-state-response (DSR) frame-
work [94–96]. DSR indicators focus on the causes of change Functional vision is an operational approach which permits
in environmental conditions in an agriculture area, the effects clarification of the complex concept of biodiversity. Biodiver-
of agriculture on the environment and the efficiency of any sity is too large to be entirely assessed by a single criterion. Bio-
actions taken. logical diversity must therefore be evaluated according to precise
objectives: ecological, agronomical or patrimonial approaches.
The OECD Expert Meeting, May 1999, in Paris suggested Many tools have been built to assess biodiversity but they meas-
more concrete indicators [86]. One goal was to select relevant ure only some parts of biodiversity. For example, models are
landscape indicators for the data available. An example for EU often limited to simple systems, while validation of indicators
territory monitoring is the Corine Land Cover (CLC) [28]. This shows the complexity of these systems. Future studies ought
monitoring at the EU level allows determination of anthropo- to examine understanding of the relationships between biodi-
genic impacts on landscapes. Initiatives aiming to preserve the versity and agro-ecosystems with complementary approaches
quality of landscapes can be designated. However, at the EU (agronomy and ecology) and produce suitable tools that permit
level monitoring is limited. A specific level must be chosen and decision-making. Studies frequently examine only one scale
the data are of limited significance for specific analysis. whereas these relationships are relevant on different scales and
The European Community initiated a project proposal on are interconnected.
agri-environmental indicators called the PAIS project. This
project contained indicators within the domain of landscapes,
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