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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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