Landscape Ecology 18: 303314, 2003.

2003 Kluwer Academic Publishers. Printed in the Netherlands.

303
Research article
Temporal variability of connectivity in agricultural landscapes: do
farming activities help?
Jacques Baudry
1,
, Franoise Burel
2
, St ephanie Aviron
2
, Manuel Martin
2
, Annie Ouin
2,3
,
Guillaume Pain
1,4
& Claudine Thenail
1
1
INRA SAD Armorique, CS 84215, 65 Rue de Saint Brieuc, 35042 Rennes C edex, France
2
UMR 6553 Ecobio, CNRS Universit e de Rennes 1, Campus de Beaulieu, 35042 Rennes Cedex, France
3
INP-ENSAT Avenue de lAgrobiopole, B.P. 107, Auzeville Tolosane 31326, Castanet, Tolosan Cedex, France
4
Ecole Sup erieure dAgriculture, 55 rue Rabelais, BP 748, 49007 Angers Cedex 01, France

Corresponding author: (Tel.: 00 33 (0)223485621; Fax: 00 33 (0)223485620; E-mail: jbaudry@roazhon.inra.fr)

Key words: connectivity, farming system, landscape structure, simulations, temporal variability
Abstract
In landscapes where natural habitats have been severely fragmented by intensive farming, survival of many species
depends on connectivity among habitat patches. Spatio-temporal structure of agricultural landscapes depends
on interactions between the physical environment and farming systems, within a socio-economic and historical
background. The question is how incentives in agricultural policies may inuence connectivity? May they be used
to manage the land for biodiversity conservation? We used simulations based on property eld maps to compare
connectivity on the same landscape during seven years of crop succession for two dairy farming systems, one being
representative of conventional systems of western France, the second one representative of systems undergoing
intensication of production. Connectivity is a measure of landscape structure and species characteristics based
on individual area requirements and dispersal distance. Models used are based on weighed distances, considering
differential viscosity for different land uses. The results show that, for a given farming system, physical and eld
patterns constraints are such that landscape connectivity remains the same over years, while it is signicantly
different between the two farming systems. This is consistent with the recent input of policies to promote environ-
mentally friendly farming systems, and conrms that policies must encounter the landscape level. The analysis also
demonstrates that the localisation of forest patches, resulting from long term land cover changes, plays a central
role in connectivity and overrides changes in agricultural land uses.
Introduction
The main characteristic of temperate agricultural land-
scapes is the expansion of farmland at the expense
of forest. In western Europe this process has been
taking place for centuries (Duby and Wallon 1975).
For plants and animal living in these fragmented and
heterogeneous landscapes, movement is a key process
for survival (Wiens et al. 1993). For mobile species,
daily movements are for food search and predator
avoidance. For all species, individuals or propagules
disperse among local populations or to colonise new
habitats, at a time scale of a year or more. Before
the period of land use intensication grew from the
1950s on, patches and linear elements of semi-natural
habitats facilitated the movements of some species.
Since then, these habitats have decreased dramatically
in intensively farmed regions (Leonard and Cobham
1977; Agger and Brandt 1988; Meeus 1990). Nev-
ertheless, numerous forest species continue to thrive
in these environments (Burel 1996). Landscape con-
nectivity, dened in this paper as the degree to which
the landscape facilitates or impedes movement among
304
resource patches (Taylor et al. 1993) is an important
control of the populations of these species (Petit and
Burel 1998). Thence, we consider connectivity as an
estimate of population survival in landscapes. Land-
scape managers, conservation and integrated pest con-
trol planners tend to enhance connectivity to maintain
or restore biodiversity at the landscape level.
The measure of connectivity is a current debate in
ecological literature, mainly between landscape ecol-
ogists and metapopulation biologists (Moilanen and
Hanski 2001; Tischendorf and Fahrig 2001). The de-
bate mainly involves the patch centred approach of
the population biologists versus a landscape oriented
one for landscape ecologists, and a neutral view of
non habitat areas versus a view of a heterogeneous
matrix. Even if those ideas simply reect differences
in the scale of the studies and the precision of ex-
pected results, several authors have developed the idea
that connectivity of a landscape depends not only
on the distance between habitat patches, but also on
the presence of corridors and stepping stones and on
the resistance of the surrounding matrix (Fahrig and
Merriam 1994; Moilanen and Hanski 1998; Pain et al.
2000; Ricketts 2001). In agricultural landscapes the
mosaic of crops and uncultivated patches, woodlots,
heath land, wetlands and hedgerows, inuences indi-
vidual movements for carabid beetles (Martin et al.
2001), butteries (Ricketts 2001), badgers (Schip-
pers et al. 1996), and damselies (Pither and Taylor
1998). Interactions among individual movements of
a species and a given land use depend on its pheno-
logical state (Ouin et al. 2000), available resources
and/or refuge effect (Henein et al., 1998). Milan de
la Pea et al. (2003) nd in Brittany, France, that
in landscapes showing similar organization of wood-
lands and hedgerows, communities of carabids differ
according to farming systems. In landscapes with few
woods remaining, carabid communities in dairy farms,
with a high proportion of maize, differ markedly from
the ones in landscapes where pig production, charac-
terized by a high proportion of cereals, is dominant.
The rst ones have communities characteristic of still
dense hedgerow network landscapes. A crop such
as maize, may, in some instance be a substitute for
woodland.
Nevertheless connectivity is a measure of land-
scape structure and determines animal or plant move-
ment or dispersal at the landscape level. Connectivity
involves spatial heterogeneity of the land as well as
individual species area requirement and dispersal dis-
tance, this is what Vos and colleagues develop as
Ecological Scaled Landscape Indices (Vos et al. 2001).
Agricultural landscapes are dynamic at several tempo-
ral levels. Within years, growth and harvest of crops
change the mosaic of resources. Among years, crop
succession in a given farming system changes the spa-
tial arrangement of the mosaic, with no or few effects
on the shape and size of the elds (Baudry and Papy
2001). On longer temporal scales, changes in farming
systems induce more durable changes that affect the
size and the shape of cropping areas and of natural or
extensively farmed areas (Baudry et al. 2000; Thenail
2002).
Due to the growing interest in sustaining biodi-
versity in agricultural areas, and for conservation and
integrated pest management (Altieri 1980), it is now
important to do more than consider habitats in a bi-
nary world that reduces a landscape to two basic
categories, suitable habitats and uninhabitable ma-
trix (Levins 1970; Gilpin and Hanski 1991; Hanski
and Gilpin 1997). Rather, the heterogeneity of the
whole matrix and its variability through time must be
included.
In this paper we considered the effects of short time
crop successions and long time changes in farming
systems on the measure of connectivity. The impor-
tance of eld patterns and the spatial differentiation of
landscapes due to farming activities have been demon-
strated in many instances. At a regional level, Simpson
et al. (1994) show the differentiation of Ohio land-
scapes according to the physical environment. Fernan-
dez Ales et al. (1992) describe a similar process in
southern Spain. This can also be shown at a global
scale (Turner II and Meyer 1994) or at a landscape
scale (Thenail 2002). Our simulations were based on
real landscapes and farming systems, and we took
into consideration the constraints of the physical en-
vironment, and of topology and farm property on the
organisation of the mosaic. Our hypothesis was that in
different crop mosaics, connectivity of the landscape
varies, and that variability is higher between farming
systems than within farming systems. Then, farming
systems can be one of the utensils of the tool box used
for the ecological management of the land.
We tested the effects of two dairy farm types, con-
sidering seven years of crop succession for each. The
two farm types differ in land use: the rst has more
permanent grassland and hedgerows, the second more
maize. They can be considered as two levels of intensi-
cation (Green 1989). Flamm and Turner (1994) have
tested the importance of using elds versus indepen-
dent pixels in land use simulations. They conclude that
305
Figure 1. Field pattern.
Table 1. Rules of land use allocation in farms of Farm type 1. For maize, cereals, sown grassland
and permanent grassland, the table gives the percent of each crop allocated to the group of elds.
Distance Hydromorphy Surface Maize Cereals Sown grass. Perm. grass.
Low Low <1 ha 0 0 100 0
(<0.5 Km) 1 ha 20 10 70 0
Medium < 1 ha 0 0 100 0
1 ha 10 0 90 0
High < 1 ha 0 0 50 50
1 ha 10 0 90 0
Medium Low < 1 ha 10 10 50 30
(0,5; 1 Km) 1 ha 25 25 50 0
Medium < 1 ha 0 0 50 50
1 ha 45 20 35 0
High < 1 ha 0 0 0 100
1 ha 50 10 40 0
High Low < 1 ha 0 0 0 100
(1 Km) 1 ha 50 50 0 0
Medium < 1 ha 0 0 0 100
1 ha 60 40 0 0
High < 1 ha 0 0 0 0
1 ha 35 5 25 35
306
Table 2. Rules of land use allocation in farms of Farm type 2. For maize, cereals, sown grassland and permanent
grassland, the table gives the percent of each crop allocated to the group of elds.
Distance Hydromorphy Surface Maize Cereals Sown grass. Perm. grass. Wood. fallow
Low Low < 1 ha 0 0 100 0 0
(<0,5 Km) 1 ha 20 20 60 0 0
Medium < 1 ha 0 0 100 0 0
1 ha 20 20 60 0 0
High < 1 ha 0 0 30 50 20
1 ha 20 0 80 0 0
Medium Low < 1 ha 10 10 50 30 0
(0,5; 1 Km) 1 ha 50 50 0 0 0
Medium < 1 ha 0 0 50 50 0
1 ha 60 30 0 0 0
High < 1 ha 0 0 0 75 25
1 ha 75 25 0 0 0
High Low < 1 ha 50 50 0 0 0
( 1 Km) 1 ha 50 50 0 0 0
Medium < 1 ha 0 0 0 50 50
1 ha 60 30 0 0 0
High < 1 ha 0 0 0 0 100
1 ha 75 0 0 25 0
only patch-based model with ownership boundaries
captured the complexity of the spatial pattern of the
landscape.
Landscape simulations
To simulate a realistic landscape we used the map of
a real landscape (a second stream order watershed)
from which we retained eld limits, built up areas,
roads and streams (Figure 1). This watershed is part
of a long-term landscape research project (Baudry
et al. 2000). The total area is 1060 ha. The current
eld pattern is the result of environmental constraints
and of technical and socio-economical history. It is
the result of physical heterogeneity and its diversity
of shape and size is the result of interactions be-
tween nature and society (Meynier 1966; Rackham
1986). Taking an actual eld pattern for simulations
permitted to incorporate both what forms a basis of
landscape structure and the spatial constraints within
which farming systems operate.
The actual land use of this area is a mosaic of
crops, grasslands, woods and hedgerows resulting
from the activities of 25 farms, all of which are not
included in the area, as in this part of France, the
territories of farms are scattered. Most of the farms
are specialized dairy farms, with more long duration
than short duration sown grassland, silage maize, and
a small proportion of grain maize. Beside maize silage,
the contribution of grass fodder (hay and silage) is
important as livestock food. Size and economic inten-
sity of the farms vary within a gradient going toward
more cash crops in the farming system, as farm area
increases (Thenail 2002).
For simulation we used seventeen farms, totally
included in the watershed, with the same farming sys-
tems for a given simulation. Both types of farming
simulated in this work are dairy farms: type 1 uti-
lizes more grassland than type 2 where maize has an
important role. We simulated land cover, categorized
as woodland, permanent grassland and other crops
(rotational grassland, maize and cereals), as well as
hedgerows over periods of seven years. Each agricul-
tural eld belongs to a farm, and all the farms have
all their elds in the watershed. This permitted us to
utilize within-farm rules of land allocation to simu-
late the landscapes. These rules (Tables 1 and 2) are
derived from empirical work in the region (Thenail,
unpublished). Distance to farmstead, soil hydromor-
phy and eld size are the main driving factors of land
use. We dene as hydromorphic, a eld with visible
307
patches of hydromorphic conditions (redoximorphic
features at the surface, on over 80% of its area). Loca-
tion of farm buildings are according to the distribution
of current villages. Fields were allocated to each farm
with simple rules of distance from buildings and size
of elds. We considered that farm type 2 is a change
toward agricultural intensication of farm type 1.
The simulations went as follows:
(1) We randomly assigned woods as to cover about
12% of the land. This simulated the fact that the pres-
ence of woods is due to a wide variety of causes,
outside the farming activities. Among the causes are
inheritance, leisure etc. For all subsequent simula-
tions, this woodland pattern was retained. We ran the
simulations with three different initial wood patterns
(A, B, C), so to test the effect of this wood pattern. In
each simulation, the same area of wood is kept, but as
wood is assigned randomly to polygons of the map,
we obtained a different spatial distribution.
(2) We ran simulations for farm type 1. We rst
simulated permanent grassland, we considered that
all elds with permanent grassland are delimited by
hedgerows, which is the current trend in hedgerow
network landscapes (Barr and Gillespie 2000) and par-
ticularly in this study site (Baudry et al. 2000). A
restricted number of crop eld boundaries (one third)
were also assigned as hedgerows.
(3) We simulated the crops for farm type 1 (7 runs)
with the empirical rules in Table 1. We maintained
the permanent grassland and hedgerow pattern con-
stant for the seven runs, which is coherent with current
agricultural use. The elds were divided into classes
according to their characteristics (distance to farm,
surface, hydromorphy) and within each class crops
were allocated to elds randomly in the proportion
given in Table 1.
(4) We also ran random simulations using only the
proportion of the different annual cover, assigned ran-
domly to elds, woodland and permanent grassland
remained identical. To keep the proportion of the crops
constant, as compared to rule simulations, we split
elds into two size categories: less or equal to 1 ha
and more than 1 ha. Thus hydromorphy and distance
to farmstead played no role.
We then ran the simulations with farm type 2.
We used the pattern of woodland, permanent grass-
land and hedgerows of farm type 1 with the following
changes:
(5) The decreasing role of permanent grassland led
to the abandonment of small, distant, hydromorphic
elds, therefore woodland may increase. This trend
Table 3. Summary of simulations.
Farm type Wood and Rules Random
hedgerow
pattern
1 A 1Arul (7 runs) 1Arand (7 runs)
1 B 1Brul (7 runs) 1Brand (7 runs)
1 C 1Crul (7 runs) 1Crand (7 runs)
2 A 2Arul (7 runs) 2Arand (7 runs)
2 B 2Brul (7 runs) 2Brand (7 runs)
2 C 2Crul (7 runs) 2Crand (7 runs)
1 A random 1Ahr (8 runs)
hedgerows
has been observed in several situations, when crops are
intensied in farming systems, use of wet meadows is
no longer part of cattle raising (Medley et al. 1995). In
contrast, some permanent grassland, in these hydro-
morphic zones, could be plowed and were considered
as elds. Subsequently, surrounding hedgerows were
removed, only one third are maintained.
(6) Other crops were allocated following empirical
rules for farmtype 2 (Table 2). Seven simulations were
run, each used as a year.
(7) Random allocation of crops were also simulated.
For one wood pattern (A) we ran eight alloca-
tions of permanent grassland at random to test the
effect of hedgerownetwork patterns largely associated
to permanent grassland. The different simulations are
summarized in Table 3.
Measure of connectivity
In this paper we considered a habitat-specic forest
species which moves between suitable forest patches
either for supplementation or complementation (Dun-
ning et al. 1992). Individuals move frompatch to patch
in continuously varying elements, within spatially ex-
plicit landscapes. Patches connected to woodlands can
be reached by a mobile individual moving out of a
patch and allowed to wander in the landscape on a
yearly basis. We used the concept of Minimal Cumu-
lative Resistance of landscapes (Knaapen et al. 1992).
Maximum distance from a patch was dened by the
energy cost of movement during this period, and is a
combination of distance and viscosity of encountered
patches.
308
Table 4. Coefcient of resistance affected to the dif-
ferent land cover types.
Land cover type Resistance
Hedgerows and wood 1
Roads and small streams 300
Built up areas 500
Rotational and permanent grassland 100
Maize 10
Cereals 100
We measured connectivity the following way:
We assigned a resistance value to each land use (Ta-
ble 4) to simulate the difculty of traversing the
landscape when moving out of woods or hedgerows.
These values were adapted fromprevious work on for-
est carabids using radio-tracing techniques (Charrier
et al. 1997) and dispersal estimated parameters (Pe-
tit and Burel 1998). A functional distance to wooded
elements was measured as the cost of moving in the
different agricultural elds, using the costpush module
of IDRISI
TM
.
The model species was a low mobility one, and the
maximum distance allowed for one run for movement
is 150 m along hedgerows, 75 m into maize elds and
7 m into grassland and cereals. The clusters of con-
nected pixels constitute the accessibility surface of Yu
(1996).
At the end of the simulations we had seven repli-
cates of maps of landscape connectivity for two types
of farming systems. These seven runs can represent
seven years of crop succession. As we considered that
the species cannot stay in elds after harvest, we run
the simulations independently for each year. By over-
laying the clusters of the different years we obtain
the frequency of connection for the different areas in
the landscape. From these maps we extracted the area
of the different clusters formed by connected woods
and hedgerows and connected pieces of farmland. We
tested for differences between treatments for the total
area of connected areas of farmland.
Results
Land cover
Table 5 gives the proportion of wood, maize and
hedgerows in the simulated maps. Figure 2 gives
examples of land use maps.
There was no change in woodland from Farm 1 to
Farm2 in these simulations. Hedgerows decreased and
maize increased. Due to the fact that we used elds as
units for simulations, there has been no strict control
of area but within Farm 1 percent of maize stayed
below the one within Farm 2, and within a wood-
land/hedgerow pattern and Farm type, the order was
similar.
Connectivity
Table 6 gives the measure of connectivity, considering
only hedgerows as suitable habitat or also considering
maize as suitable, though not optimal, for the different
situations. Examples of connectivity are presented in
Figure 3.
By construction, connectivity due to hedgerows
decreases from Farm 1 to Farm 2, by about 25%.
When maize is considered as suitable habitat, con-
nectivity increases (by construction); a larger area of
maize, even with less hedgerows, increases connectiv-
ity.
Test of differences between simulations. Differences
between initial woodland/hedgerowpatterns and types
of farming systems were tested by analysis of variance
of the SYSTAT software.
Overall, connectivity of farmland differed accord-
ing to wood pattern (A, B, C) and rules of land
allocation (Figure 4 and Table 7). The differences are
signicant with the different measures (total number
of connected pixels, cluster of 0.25 ha or more or clus-
ter of 1.25 ha or more). Results are only given for
connectivity measured with clusters of all connected
pixels. Landscapes organized according to rules of
farm 2 were signicantly less connected than those
organized according to farm1 (Figure 4), but these dif-
ferences were less than between woodland/hedgerow
initial structure. Comparisons of simulations of farm 1
vs. farm 2 show that differences were signicant for
each woodland/hedgerow pattern (Table 7). We found
no difference between rule and random crops, nei-
ther within farm rules or within woodland/hedgerow
patterns. Using rules only generated differences in
connectivity measured for each run.
Within both farming system rules, differences be-
tween woodland/hedgerow patterns were signicant
in 5 simulations out of 6 (Table 7). This empha-
sized the importance of woodland/hedgerow patterns.
Differences in connectivity showed that, when sig-
nicant, differences between woodland/hedgerowpat-
309
Figure 2. Examples of land cover patterns.
Figure 3. Examples of connectivity patterns.
310
Table 5. % of the different land cover types that varies (when, by construc-
tion the value does not change, it is noted id.)
Farm type Rules/ Simulation Wood Hedgerows Maize
random
1 Rules A 12.3 6.1 19.3
id. Random A id. id. 18
id. Rules B 13.9 5.7 18.7
id. Random B id. id. 17.2
id. Rules C 11.4 6.1 20.6
id. Random C id. id. 18
2 Rules A 12.3 4.7 25.2
id. Random A id. id. 28.1
id. Rules B 13.9 4.3 25.4
id. Random B id. id. 24.5
id. Rules C 11.4 4.6 27.3
id. Random C id. id. 29.8
Table 6. Measure of connectivity in the different landscapes (made with
rules) comparing the role of hedgerows and hedgerows +maize (values in
km
2
)
Simulation Connectivity due to Connectivity due to Increase
hedgerows hedgerows + maize (%)
1A 1.02 1.79 0.75
1B 0.90 1.50 0.66
1C 0.95 1.68 0.77
2A 0.75 (26.3%) 1.67 1.21
2B 0.70 (22.9%) 1.41 1.02
2C 0.68 (28.7%) 1.52 1.24
terns were more important than differences between
farm types (Table 7).
A two ways analysis of variance (woodland/hedge-
row pattern and Farm type) showed that both variables
explained differences (for the type of farm F=21.929
and p <0.001 and for woodland/hedgerowF=34.819
and p <0.001) and had no interactions (F=0.577
and p =0.567). This analysis conrmed the higher
importance of woodland/hedgerowpatterns.
Effects of initial woodland/hedgerowpattern. Wood-
land initial patterns were randomly generated and
did not differ by either mean distance between wood
patches or fractal dimension. Thence, we tested for
the effect of hedgerow patterns, within one wood-
land pattern (A). Hedgerows, in our model are closely
related to permanent grassland. So we randomly as-
Figure 4. Mean connectivity in the various simulations.
311
Table 7. Results of the tests of differences between simulations: connected
farmland (average over 7 runs)
Simulation F P % connectivity difference
(average)
1ABCrul/2ABCrul 8.214 0.007

7.6
1ABCrand/2ABCrand 1.655 0.296
12ABCrul/12ABCrand 0.069 0.793
1ABCrul/1ABCrand 0.747 0.392
2ABCrul/2ABCrand 0.129 0.721
1Arul/2Arul 5.572 0.033

7.08
1Brul/2Brul 6.374 0.027

5.96
1Crul/2Crul 10.614 0.007

9.50
1Arul/1Arand 0.637 0.440
1Brul/1Brand 0.936 0.352
1Crul/1Crand 2.338 0.152
2Arul/2Arand 0.050 0.827
2Brul/2Brand 0.170 0.687
2Crul/2Crand 0.530 0.481
1Arul/1Brul 37.065 0.000

16.3
1Arul/1Crul 3.677 0.079 6.20
1Brul/1Crul 24.055 0.000

12.07
2Arul/2Brul 37.222 0.000

15.29
2Arul/2Crul 11.268 0.006

8.64
2Brul/2Crul 5.337 0.039

7.85
signed permanent grassland, then crops according to
farm 1 rules in 7 simulations and tested the difference
in connectivity between considering all non wooded
elements as hostile (viscosity = 100) and connectiv-
ity as measure above; so the differences in hedgerow
network patterns and their relations to wood patches
can be assessed. We found a relationship between both
measures (Figure 5): the higher the connectivity with
hedgerows, the higher the connectivity considering
both hedgerows and crops.
Cumulative time effects
By overlaying annual cluster within each simulation,
we saw the effects of time. As shown in Table 8,
the farmland being in a cluster was six times larger
with farm type 2 than 1. For farm type 2, it was also
higher with simulations according to rules than with
random land allocation. This difference was negli-
gible for simulations with farm type 1. This result
demonstrated the importance of taking time into ac-
count. Some hedgerows stay isolated (functional isola-
tion) in similar proportions in the different simulations
(Figure 3).
Figure 5. Relationship between connectivity due to hedgerows and
overall connectivity.
Discussion
Our simulations demonstrated that rather slight
changes in farming systems (the same crops in dif-
ferent proportions and a decrease of hedgerows) can
lead to signicant differences in landscape ecologi-
312
Table 8. Test of the cumulative effects on connectivity.
Test Sum=1 Sum=6 Isolated hedgerows
Rules vs. random F=0.341, p =0.572 F=6.397, p =0.030

F=2.897, p =0.12
1rules vs. 1random F =0.602, p =0.481 F=6.149, p =0.068 F =0.315, p =0.604
2rules vs. 2random F =3.462, p =0.136 F=7.696, p =0.050

F=2.748, p =0.173
1rules vs. 2rules F =4.625, p =0.098 F=32.780, p =0.005

F=0.029 , p =0.873
cal characteristics, notably changes in connectivity,
an indicator of potential use of space by animals.
Three factors play a role in determining connectivity:
(1) land cover distribution, (2) the initial woodland and
hedgerow network patterns and (3) farm internal rules
of land allocation.
The explicit program of Landscape Ecology is to
consider the effects of landscape heterogeneity on eco-
logical processes as well as to understand the causes of
changes in this heterogeneity. Up to now, most stud-
ies have paid attention to important changes, mainly
changes affecting forest cover. As the time of consid-
ering the matrix as neutral or hostile comes to an end
(Ricketts 2001), it is necessary to increase the under-
standing of the dynamics of this matrix (farmland in
the case of forest as the main habitat). The organiza-
tion of landscape patterns may be driven by different
factors. For instance, forest growth results from social
or historical events (forest attached to castles, sub-
sidies for individual reafforestation), while farming
activities are constrained by eld pattern and soil con-
ditions, thence afforestation and agricultural land use
are, in our simulations, independent. Our simulations
showed that the way they are combined in space can
lead to different landscape ecological characteristics,
as the relationships between woodland, hedgerows and
maize eld can vary.
Our results showed that a relatively small change
in farming modied connectivity in a signicant man-
ner. Both farming systems considered here, can be
classied as dairy farm in the EU farm types (EU-
ROSTAT). It was the rules of within farm land use
allocation that made the differences, not the propor-
tions of the different crops. The importance of farm
functioning was emphasized by the cumulative ef-
fects. Farming systems induce landscape changes at
two time scales. Over a few years time, crop suc-
cessions produced year to year change, but overall
there was a stable pattern. In our simulations, for
any treatment, the variability between runs was rather
small. Over a longer period, landscape changes were
determined by changes in the farming systems, and
associated changes in cropping systems. The relative
stability of connectivity within the functioning of a
farming system can be interpreted as limits within the
landscape/farm systems with a narrow range for its
behaviour. Relatively small changes in farming sys-
tems created signicant changes in connectivity, thus
changes in landscape boundary conditions.
The co-operation between scholars of farming sys-
tems and landscape ecologists is crucial to decipher
landscape processes, as farming activities are often
the most important factor driving the dynamics of
rural landscapes. The concept of micro-regional crop-
ping system (the repetitive combination of crops in a
given area (Papy 2001)) may play an important role
in understanding agricultural landscapes. It is the ex-
pression of land use by farming systems integrating
both the type of farming systems and the physical and
eld pattern constraints. Types of crop successions and
crop management can be inferred from micro-regional
cropping systems. A stronger relationship with farm-
ing system scholars would help to model landscape
dynamics and look for thresholds in their trends. It
would also permit one to extrapolate landscape pat-
terns and processes at a regional scale. This integration
of causes of landscape changes in landscape models
has been advocated by Baker (1989) as a condition to
understand dynamics.
Our simulations demonstrated that the processes
that organize landscape patterns must be taken into
account in the analysis of connectivity models. These
ndings question the way we map landscapes and the
type of information put in the models. The impor-
tance of land covers between optimal habitat patches,
whether to impede or facilitate movement, is widely
recognized in empirical and modeling studies (Pe-
tit and Burel 1998). Land cover in the matrix is
usually more unstable than habitat patches of impor-
tance in the maintenance of biodiversity; this is why
either cumulative effects of land uses or land use dy-
namics over a number of years must be incorporated
313
in connectivity models. A measure of connectivity
based on dynamic structural patterns and assessment
of potential movement offers the possibility to closely
link biological processes (species behaviour) and land-
scape dynamics. It should be suitable to assess the
ecological outcomes of various landscape scenarios.
The cumulative effects of crop succession are con-
sistent with empirical ndings on the relationships
between crop successions and eld margin ora. They
are stronger than those between ora and the adja-
cent crop et the time of the survey (Le Coeur et al.
2002). The close relationships between hedgerows
(eld boundaries in general), structure and composi-
tion, and adjacent land use is established in many in-
stances (Barr and Petit 2001; Baudry et al. 2000). Our
results imply that land management can use a design of
partially suitable habitat to increase connectivity. This
also means that agri-environmental policies cannot be
restricted to semi-natural elements, and that normal
activities of farmers can help.
Acknowledgements
We thank the Programme Environnement Vie et So-
cit of the CNRS (Motive) for its nancial support.
Stimulating discussions with Nicolas Schermann were
helpful.
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