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Animal Conservation. Print ISSN 1367-9430

There and back again? Combining habitat suitability

modelling and connectivity analyses to assess a potential
return of the otter to Switzerland
C. Cianfrani1, L. Maiorano1, A. Loy2, A. Kranz3, A. Lehmann4, R. Maggini5* & A. Guisan1
1 Department of Ecology and Evolution, University of Lausanne, Lausanne, Switzerland
2 Department of Science and Technology for the Environment, University of Molise, Pesche, Italy
3 Alka kranz, Graz, Austria
4 Institute of Environmental Sciences, University of Geneva, Carouge, Switzerland
5 Department of Ecology and Evolution, University of Lausanne, Lausanne, Switzerland.

Keywords
species recovery; habitat suitability model;
connectivity; multi-step approach; integrated
management framework; long-term
conservation strategies.
Correspondence
Email: carmen.cianfrani@gmail.com
Editor: Res Altwegg
Associate Editor: Madhu Madhusudan
*Current address: Centre of Excellence for
Environmental Decisions (CEED), University
of Queensland, Goddard Building 8, St
Lucia, Brisbane, Qld 4072, Australia.
Received 3 July 2012; accepted 29 January
2013
Abstract
After a steady decline in the early 20th century, several terrestrial carnivore species
have recently recovered in Western Europe, either through reintroductions or
natural recolonization. Because of the large space requirements of these species
and potential conflicts with human activities, ensuring their recovery requires the
implementation of conservation and management measures that address the envi-
ronmental, landscape and social dimensions of the problem. Few examples exist of
such integrated management.
Taking the case of the otter (Lutra lutra) in Switzerland, we propose a multi-
step approach that allows to (1) identify areas with potentially suitable habitat, (2)
evaluate their connectivity, (3) verify the potentiality of the species recolonization
from populations in neighbouring countries. We showed that even though suitable
habitat is available for the species and the level of structural connectivity within
Switzerland is satisfactory, the level of connectivity with neighbouring popula-
tions is crucial to prioritize strategies that favour the species recovery in the field.
This research is the first example integrating habitat suitability and connectivity
assessment at different scales with other factors in a multi-step assessment for
species recovery.

doi:10.1111/acv.12033

Introduction tion and management measures because these species

Following a steady decline since the beginning of the 20th
century, several species of carnivores such as the wolf, the
lynx, the bear and the otter have disappeared from most
Western European countries (Enserink & Voge, 2010).
Their decline is generally attributed to a combination of
human persecution, habitat destruction and losses of prey
species (Ceballos et al., 2005). However, in recent decades,
carnivore populations have recovered in most of Western
Europe, partly supported by reintroduction projects but
largely as a result of natural population expansions. For
instance, wolves returned in the Alps coming from the
Italian Apennines (Enserink & Voge, 2010), while otters are
recovering in many Central European countries (Kranz,
2000; Loy et al., 2010; Panzacchi, Genovesi & Loy, 2011).
But some exceptions are remarkable, such as the case of the
Iberian lynx (Palomares et al., 2011).
The recovery of any medium to large carnivore species
should be closely monitored to carefully design conserva-
usually require large areas and can potentially create con-
flicts with human activities. This need for management is
particularly true in human-dominated landscapes, such as
central-western Europe, where the habitat suitable for the
presence of many species may be highly fragmented (Temple
& Terry, 2007) and freshwater habitats are highly impacted
by introduced species, pollution, water captation and loss of
riparian vegetation.
Before planning the reintroduction of a species (Morrel,
2008) or establishing actions to favour the natural expan-
sion of threatened and rare species, at least three main ana-
lytical steps should be undertaken: (1) identify the areas
potentially suitable (or partially suitable, as here, depend-
ing on environmental data availability) for the species
(Martinez-Meyer et al., 2006; Olsson & Rogers, 2009; Cook,
Morgan & Marshall, 2009; Rodrigues-Soto et al., 2011); (2)
evaluate whether the suitable areas in a region are suffi-
ciently well connected to guarantee gene flow and survival of
a viable population (Kramer-Schadt et al., 2004; Singleton,

Animal Conservation (2013) 2013 The Zoological Society of London 1

Can the otter return to Switzerland?
Gaines & Lehmkuhl, 2004; LaRue & Nielsen, 2008; Huck
et al., 2010; Carranza et al., 2012); (3) evaluate the level of
connectivity with neighbouring areas from were the popu-
lation can expand (Kramer-Schadt et al., 2004).
In this study, we illustrate these three steps using the
European otter (Lutra lutra) as a case study. The otter is a
flagship species for freshwater ecosystems, and it suffered a
severe decline during the 20th century in most European
countries (Foster-Turley, Macdonald & Mason, 1990;
Robitaille & Laurence, 2002) because of the reduction of
food supplies, increases in water pollutants, persecution by
humans and the destruction of riparian habitat (Foster-
Turley et al., 1990; Kruuk, 2006). In Switzerland, the otter
went extinct in the 1980s (Foster-Turley et al., 1990) and it is
still absent, even though the species is really close to the Swiss
border on the French side and is moving closer on the
Austrian side (Kranz et al., 2008; Fig. 1). The challenges and
opportunities for the comeback of the otter in Switzerland
have been discussed considering two alternative options:
active reintroduction projects and support to natural recolo-
nization (Weber, 1990; Weber, 2004). Building on previous
work (Cianfrani et al., 2010, 2011), we assess here the feasi-
bility of these two options using a framework that relies on
the most advanced spatial techniques and most robust data
available, and incorporating the three steps previously iden-
tified. We aim particularly at using this framework to answer
the following questions: (1) are there environmentally suit-
able areas for the otter in Switzerland?; here, we mainly assess
partial environmental suitability due to the lack of prey
availability and water pollution maps; (2) are the partially
suitable areas connected with each other within Switzerland?;
(3) would it be possible for the otter to naturally recolonize
Switzerland from neighbouring countries?
2 Animal Conservation (2013)
C. Cianfrani et al.
Figure 1 Otter distributions in neighbouring
countries of Switzerland and map of the
Swiss river catchments.
Material and methods
Our multi-step approach (Fig. 2) consists of three steps. We
first identify potentially suitable areas using habitat suitabil-
ity models (HSM; sometime referred to as species distribu-
tion models; see Habitat Suitability section) and then
evaluate the connectivity between the suitable patches (see
Internal Connectivity section). Considering that there are
populations of the species living in close proximity to the
Swiss border, we also evaluate the potential for these popu-
lations to recolonize suitable areas in Switzerland (see
External Connectivity section).
Habitat suitability
Given that no recent observations were available to cali-
brate a species distribution model in Switzerland, we used a
combined approach to identify potentially suitable habitats
for the otter in Switzerland. First of all, we fitted a purely
climatic model at the scale of the European Alps (see
Cianfrani et al., 2011), considering the largest possible cli-
matic niche for the species (see Pearson, Dawson & Liu,
2004), and then we combined this model with one based on
local environmental variables (i.e. mainly topography and
land use; climate excluded), calibrated at the regional scale
in Austria and projected to Switzerland (using the same
variables) (see Fig. 2). Although water quality and prey
availability are important environmental determinants of
the otter presence, these information are not available in a
spatially explicit way along rivers in both study areas (see
discussion). Therefore, none of these two variables could be
integrated in the local HSM. To overcome as much as
possible these limitations, we used surrogate variables in the
2013 The Zoological Society of London
C. Cianfrani et al.
Figure 2 Flow chart of the integrative approach.
model (e.g. elevation for fish density) and combined the
existing non-spatially explicit information later on with
the spatial predictions made with the other local predictors,
as a way to improve the interpretation of the the results. The
modelling approach, species data, environmental predic-
tors, model calibration and predictions combination are
described in detail in Supporting Information Appendix S1.
Given the particular ecology of the species, the projection of
the final model was restricted to a buffer of 150 m around
main rivers in Switzerland.
Environmental similarity between Austria
and Switzerland
Before calibrating the environmental model in Austria and
projecting it into Switzerland, we assessed the environmen-
tal similarity between Austria and Switzerland to ensure
that no bias would occur when a regional model fitted in
Animal Conservation (2013) 2013 The Zoological Society of London
Can the otter return to Switzerland?
Austria is projected in Switzerland. To perform this assess-
ment, we used the Mahalanobis distance to calculate an
index of environmental similarity. In particular, we calcu-
lated the Mahalanobis distance between the values of the
environmental predictors in each pixel in Austria and the
average of the same predictors calculated for the whole
Austrian environment (hereafter d_AUAU). Next, we cal-
culated the Mahalanobis distance between the values of the
same environmental predictors for each pixel in Switzerland
and the average values for Austria (hereafter d_AUCH).
The comparison between d_AUAU and d_AUCH allowed
us to test for the similarity between the Austrian and Swiss
environments. Using ArcGis, we mapped d_AUAU and
d_AUCH. We divided d_AUAU into 20 quantile classes
and we calculated the percentage of d_AUAU pixels
contained in each of the 20 quantile classes. Following the
same procedure, we calculated the percentage of pixels of
d_AUCH contained in each quantile class. At the end of this
process, we obtained the percentage of pixels of d_AUAU
and d_AUCH contained in each quantile class. We tested
whether these two distributions differed significantly using a
KolmogorovSmirnov test.
Internal connectivity
To assess the landscape permeability for an otter moving
between potential suitable patches within Switzerland, we
used the Circuitscape software (version 3.5.4; McRae,
2006). Following Carranza et al. (2012), the permeability
was evaluated within the whole suitable catchments, that
is, not restricting to the 150 m buffer around the river
stretches, as we were interested in exploring the potential
movements of otters both within (daily movements) and
between (dispersal process) catchments. The software
draws on circuit theory to evaluate the total landscape con-
ductance between sites based on multiple paths (McRae,
2006). The inputs for Circuitscape are a raster in which
each cell has a conductance value that corresponds to the
relative probability of the otter moving through the differ-
ent habitat types and a raster of focal nodes (patches suit-
able for the presence of the otter according to the habitat
suitability analysis).
To generate the conductance map, we considered the land
cover (Land use 25 categories; Maggini, 2011), slope (calcu-
lated from ASTER GD, see Table 1) and human density
(Swiss Federal Statistical Office) of the region. The conduct-
ance values of the different land-use classes were assigned
following Loy et al. (2009) and Carranza et al. (2012;
Table S3 in Supporting Information Appendix S3).
External connectivity
To assess the landscape permeability for an otter dispersing
from populations closer to the Swiss boundaries (external
permeability; Fig. 1), we used the same approach explained
in the previous paragraph.
3
Can the otter return to Switzerland? C. Cianfrani et al.

Table 1 Percentage area in the catchment, suitable area considering the catchment area, suitable area considering the entire area of Switzerland
(CH), permeable area considering the catchment, permeable area considering the entire area of Switzerland (CH) in each catchment
Catchment surface Suitable area in the Suitable area in the Permeable area in the Permeable area in the
River area/Swiss surface catchment/catchment catchment/total catchment/catchment catchment/total permeable
catchment area (%) surface area (%) suitable area in CH (%) surface area (%) area in CH (%)
Aare 26.6 33 30.7 44.0 38.5
Adige 0.5 0 0 0.0 0.0
Inn 6.2 0 0 12.5 1.7
Limmat 7.04 43.2 10.6 26.1 4.5
Mera 0.6 0 0 7.6 0.1
Posh 0.7 7 1 40.1 0.7
Reuss 9.12 36.9 11.8 23.1 5.8
Rhine 25.9 28.6 25.9 51.1 38.6
Rhone 13.15 25.6 11.8 10.1 5.2
Ticino 9.7 26.9 9.1 20.8 4.9

Figure 3 The map visualizes the Mahalano-

bis distances between the values of the
environmental predictors for each pixel in
Switzerland and the average values for
Austria. The graphic represents the distribu-
tion of the Mahalanobis distances between
the values of the environmental predictors
for each pixel in Austria and the average
values for Austria (hereafter d_AUAU) and
between the values of the same environ-
mental predictors for each pixel in Switzer-
land and the average values for Austria
(hereafter d_AUCH) (orange line). The x-axis
represents the 20 quantiles for which the
Mahalanobis distance between the mean
of the distribution of the Austrian environ-
ment and each point in Austria has been
classified, and the y-axis represents the per-
centage of pixels included in each quantile.
We performed a KolmogorovSmirnov test
to compare the two distributions.

To generate the conductance map, we considered the land Results

cover (Corine Land Cover 2000 categories), slope (calcu-
lated from ASTER GD, see Table S3 in Supporting
Information Appendix S3) and human density (from Euro-
pean Environmental Agency) of Austria, Alsace (Germany),
and Haute-Savoie (France), for which data on otter pres-
ence were available at a 1-km2 resolution. Following Loy
et al. (2009) and Carranza et al. (2012), we assigned differ-
ent conductance values to the different land-use classes,
(Table S3 in Supporting Information Appendix S3). In this
case, we considered the areas with stable otter presence in
the countries surrounding Switzerland as the focal nodes.
These areas include the Inn, Rhine and Rhone catchments.
Habitat suitability
The results showed that the distributions of Mahalanobis
distances calculated for the average environmental predic-
tors in the whole Austria (hereafter d_AUAU) and the same
predictors for each pixel in Austria (d_AUCH) and Switzer-
land (d_AUCH) were not significantly different (D = 0.37,
P-value = 0.15), providing support to the transferability of
the HSM from Austria to Switzerland (Fig. 3). The areas
with the highest difference in habitats between Austria and
Switzerland are located in the Swiss Alps sector (Fig. 3).

4 Animal Conservation (2013) 2013 The Zoological Society of London

C. Cianfrani et al.
Overall, all of the HSM calibrated with the different tech-
niques can be considered as potentially useful (sensu Swets,
1988) since they have area under curve (AUC) values > 0.7.
In particular, the bioclimatic models calibrated considering
the entire Alpine range had a minimum AUC of 0.835 [for
Generalized Linear Model (GLM)] and a maximum of
0.905 [for random forest (RF)]. Comparable results were
obtained for the models based on the environmental
variables and calibrated in Austria, with a minimum
and a maximum AUC of 0.711 (for GLM) and 0.807 (for
ANN), respectively (Table S2 in Supporting Information
Appendix S2).
According to our ensemble model derived from the inter-
section of the bioclimatic and environmental models, 31% of
Switzerland (considering only buffers of 150 m around
rivers) is suitable for the otter (Figs. 4ac; Table 1). The
30.7% of suitable areas is located in the Aare River catch-
ment and 25.9% in the Rhine catchment, were they are
mainly concentrated in the northern part of the catchment.
The remaining suitable patches occur in Rhone and Reuss
catchments (11.8% each), and in the Limmat (10.6%;
Fig. 4c; Table 1).
Internal connectivity
Almost 33% of Switzerland has a permeability value greater
than the median permeability of the country (Fig. 4d;
Table 1). The Aare and Rhine catchments together contain
almost 80% of the most permeable areas, while the Reuss
and Rhone catchments are much less permeable (5.8 and
5.2% of permeable areas, respectively; Table 1). Extended
permeable areas connect the Aare and the Reuss catch-
ments, the Rhone and Ticino catchments (in particular
along the Rhone Valley), the Ticino and the Reuss catch-
ments, and the Limmat and the northern part of the Rhine
catchment (Figs. 4df). In the Alpine sector, the connectiv-
ity is mostly very low because of the rugged topography
(Figs. 4df).
External connectivity
The external permeability map clearly shows that the otter
could easily colonize Switzerland from the surrounding
populations in the Haute-Savoie (Fig. 4f). The Rhone catch-
ment in Switzerland should be the first to be recolonized,
and from here, the otter could easily expand into the Aare
catchments. According to our model results colonization
from the Alsace to the Rhine catchment is less likely, and
colonization from Austria to the Inn catchment (where there
are permeable areas but not suitable habitats for the otter)
appears very difficult but this possibility can not be excluded
in the long term (Figs. 4e,f).
Integrated predictions
Our results indicate that natural recovery is most likely to
occur from populations in Haute-Savoie, potentially recolo-
nizing 12% of the suitable habitat and 5% of the permeable
Animal Conservation (2013) 2013 The Zoological Society of London
Can the otter return to Switzerland?
areas in the Rhone catchment in Switzerland (Table 1).
From these areas, colonization can spread to the Aare
catchment, which is well connected to the Rhone catchment.
These results suggest that, if repopulating Switzerland with
otters was to become a main goal, the permeable areas
between the Haute-Savoie region and the Rhone catchment,
as well as that between the Rhone catchment and the Aare
catchments (Fig. 4e,f) should be preserved to the greatest
possible extent, and eventually restored where needed.
Discussion
After a severe otters decline in the 20th century in most
European countries (Robitaille & Laurence, 2002), several
otters reintroduction actions started between the end of
1970s and the beginning of 2000, for example, in the Neth-
erlands, Alsace, Spain (Catalonia), Italy, Switzerland,
Czech Republic and Sweden. In Switzerland, eight otters
from Bulgaria were released in 1975 in the Schwarzwasser
River (Bern canton), but the reintroduction proved unsuc-
cessful (Weber, 1990). In others countries, the reintroduc-
tion proved successful in a first step, but current populations
seem not viable in the long term. This is the case, for
example, in the Netherlands, where the reintroduction
started in 2002, with 31 otters released along 6 years. The
reintroduced populations maintained themselves so far, but
are small and isolated, with dispersal and migration being
severely limited by heavy vehicular traffic. In 2008, an
important inbreeding rate was measured in these isolated
populations, urging for the creation of dispersal corridors to
connect them with the larger neighbouring German popu-
lations (http://www.otter.wur.nl). In such case, a more thor-
ough evaluation of habitat suitability and connectivity
would have certainly helped planning a reintroduction
program with a stronger emphasis on dispersal and migra-
tion of reintroduced individuals, and associated longer term
viability of reintroduced populations.
Our study does indeed illustrate that, before deciding if a
species recovery should be supported by reintroduction or
favoured through natural expansion from neighbouring
populations, several key aspects need to be investigated
(Morrel, 2008). This process requires the type of assessment
and tools that we have proposed.
Assessing habitat suitability
The first step consisted in identifying whether suitable habi-
tats were available that could be eventually repopulated and
where they were located (Morrel, 2008). This step can be
difficult when the target species is no longer present in the
area targeted for recovery because it requires fitting a suit-
ability model in a different area from which the model can
be safely transferred to the area of concern. Such transfer-
ability proved not to be an issue here, as shown by our
environmental similarity diagnostic test, but it could be
critical in other cases and should be considered carefully.
Several studies have similarly assessed model transferability
to evaluate how well a model calibrated in one situation can
5
Can the otter return to Switzerland? C. Cianfrani et al.

Figure 4 (a) Map of the bioclimatic suitable areas; (b) map of the environmental suitable areas; (c) map of the total suitable areas obtained by
overlapping the bioclimatic and the environmental suitability; (d) map of the connectivity between all suitable areas; (e) map of the integrated
results; the permeable areas were identified as those areas with a permeability value greater than the median permeability in Switzerland. The
points refer to PCBs measurements expressed in function of the effects on the otter.

6 Animal Conservation (2013) 2013 The Zoological Society of London

C. Cianfrani et al.
be transferred to a different context (e.g. another time
period after a climatic change, or another region for an
invasive species; Araujo & Guisan, 2006; Randin et al.,
2006; Anderson & Raza, 2010; Maiorano et al., 2012), and
our study illustrates well why testing for environmental
similarity is an important step to perform prior to transfer
environmental niche models.
In our example, predictions of environmental suitability
were restricted to a 150 m buffer around rivers, as otters are
rarely found beyond 150 m from streams (Philcox, Grogan
& Macdonald, 1999; Kruuk, 2006). In this case, having
presence data of high locational accuracy for calibrating the
HSM becomes particularly important. In this respect, the
Austrian data were the only data with a sufficiently accuracy
but also from a similar enough environment to allow rea-
sonable transferability. To better capture the realized niche
of the species, we combined climatic data at large scale with
other non-climatic regional environmental factors. We com-
bined these data because it is known that climate mostly
controls the distributions of species at macroscales (e.g.
Cianfrani et al., 2011), whereas other abiotic or biotic
factors control species distributions on a finer spatial scale
(Pearson et al., 2004; Lomba et al., 2010).
Assessing internal connectivity
If there are suitable habitats throughout an area to be even-
tually repopulated, the second step consists in evaluating
whether the suitable habitats are sufficiently connected to
allow movements between populations to assure that suit-
able patches can be colonized (Kramer-Schadt et al., 2004;
Singleton et al., 2004; Morrel, 2008; LaRue & Nielsen, 2008;
Kuemmerle et al., 2011; Fig. 2). The analyses of the internal
connectivity among the suitable habitats, allows for the
identification of the land matrix available for species move-
ments within catchments and dispersal between river basins
(Carranza et al., 2012). This analysis provides a general
framework to assess the species possibilities to move across
the landscape and to identify the patches that could be
possibly colonized (in our case, river catchments; Loy et al.,
2009; Carranza et al., 2012). This type of analysis is essential
in Central Europe where landscapes are profoundly domi-
nated by humans (Kramer-Schadt et al., 2004; Kuemmerle
et al., 2011).
In response to the extensive habitat modification and
fragmentation, conservation planners have developed a
number of tools to increase connectivity between core
areas of habitat to facilitate the movement of species.
Among them, ecological networks, which aim to identify
core areas, species corridors and buffer zones, the integra-
tion of ecological concerns into spatial land-use planning
and broader approaches to increase landscape permeability
(Jongman & Pungetti, 2004; Crooks & Sanjayan, 2006).
Providing increased connectivity is a vitally important
aspect of mammal conservation in Europe and will provide
a key tool to allow species to adapt to current habitat frag-
mentation and projected future climate change (Temple &
Terry, 2007). The maintenance and restoration of land-
Animal Conservation (2013) 2013 The Zoological Society of London
Can the otter return to Switzerland?
scape permeability is a particularly important issue and a
main responsibility for Switzerland since it occupies a
central position in the middle of the three important Euro-
pean otter subpopulations (Cianfrani et al., 2011). We rec-
ognize that we only considered structural connectivity in
this analysis, while functional connectivity was only indi-
rectly evaluated (Boitani et al., 2007). This is due to the
fact that no data on otter movements during dispersal are
currently available, and thus our approach could not be
further improved. In fact, the only available data refer to
the radiotracking monitoring of reintroduced otters in the
Netherlands and Spain (Saavedra Benditi, 2004; Koelewijn
et al., 2010), but no details are provided there on the envi-
ronmental characteristics limiting or favouring otter move-
ment across the landscape.
Assessing external connectivity
Considering that in recent decades carnivore populations
have recovered in most of Western Europe mainly as a
results of natural population expansions, the evaluation of
the level of connectivity with the neighbouring populations
is an important step before deciding if a species recovery can
be supported by a reintroduction or by actions to favour the
natural expansion of the species from areas where it is cur-
rently present. Investigating the possibility of a natural rec-
olonization from neighbouring populations is crucial to
prioritize conservation strategies and to establish where to
act. As reintroductions have a number of limitations and
extra costs, if a recovery from natural populations is feasi-
ble, a reintroduction plan should be avoided and actions
to favour the natural expansion should be preferred
(IUCN/SSC Re-introduction Specialist Group, 1998). This
is particularly true for the Eurasian otter. In fact, otter
reintroduction projects did face such a series of problems
and extra costs (Weber, 1990; Ralls, 2002). In contrast, the
species seems to have high dispersal capabilities, as sug-
gested by its fast recovery in many European countries,
including Germany, Austria, Italy, France, UK and Spain
(Clavero et al., 2010; Marcelli & Fusillo, 2009; Loy et al.,
2010; Marcelli et al., 2012), following its overall decline in
the 1990s. In our example, considering that the species is at
present close to the Swiss border on the French side and is
moving closer on the Austrian side, the investigation of the
otter re-expansion dynamics in Austria and France would
be a crucial step.
In our study, we used a conductance approach to evalu-
ate the connectivity of neighbouring populations of France
and Austria with the locations of suitable habitats in Swit-
zerland in order to preliminarly assess if a natural recoloni-
zation (as opposed to a human reintroduction) would be
possible.
Our results, obtained from the external permeability
analyses, indicate that, in the short term, a recolonization
could most likely occur from populations in Haute-Savoie,
potentially colonizing 12% of the suitable habitat and 5% of
the permeable areas in the Rhone catchment in Switzerland
(Table 1). According to the model, from these areas,
7
Can the otter return to Switzerland?
colonization can spread to the Aare and Ticino catchments,
which are well connected to the Rhone catchment. This may
well be true for the Aare, but not for Ticino because of the
topography there. This highlights the risks of modelling
suitable habitats without considering connectivity, and
emphasizes the need for careful interpretation. However, the
results suggest that, if repopulating Switzerland with otters
was to become a main goal, the permeable areas between the
Haute-Savoie region and the Rhone catchment, as well as
between the Rhone catchment and the Aare catchment,
(Fig. 4e,f) should be preserved and monitored to the great-
est possible extent.
Our results show that, according to the environmental
variables considered, suitable habitats are available in
Switzerland and sufficiently connected between them and
with those outside the borders to allow for the return of
the otter. On the basis of these results, we can draw some
preliminary recommendations for the prioritization of
actions required for the otter recovery in Switzerland. The
main one would certainly be the preservation of the suit-
able riverine habitats in the Rhone catchment that are
closest to the Haute-Savoie populations. Also, according
to the recent IUCN recommendations (Vasilijevic &
Pezold, 2011), the development of a concerned trans-
boundary conservation plan would be crucial to ensure the
protection and establishment of source populations in
Austria and France.
Missing factors
Although we tried to be exhaustive in covering the main
factors affecting the distribution and the expansion across
the landscape of the otter, three factors previously reported
as important causes of the otters extinction in Switzerland
could not be included in the analyses. These are (1) pollu-
tion by polychlorinated biphenyl (PCB)s (Foster-Turley,
1992; Weber, 1990; Gutleb & Kranz, 1998; Weber, 2004),
(2) food availability (Kruuk, 2006) and (3) hunting and
social acceptance.
Data on PCBs, fish biomass and social acceptance are not
available in a spatially explicit way for all rivers in the
country, and available measurements or information are too
scattered to be spatialized in a reliable way and used in the
HSM. Furthermore, these data were not available for
Austria where the regional model was fitted. And last, envi-
ronmental data for freshwater bodies such as water tem-
perature, depth, water velocity or water chemistry do not
have a sufficient level of spatial accuracy to allow fish
density or PCB contamination to be satisfactorily modelled
across Switzerland and be used as predictors in the otter
models. In particular, fish distrtibution is affected by small,
local variations in the river network so that interpolation
along the river network is often inaccurate and not reliable
(White, McClean & Woodroffe, 2003). Furthermore, the
relationship between fish density and otter distribution is
difficult to assess because this information is unevenly avail-
able within an otter range and existing studies led to con-
8 Animal Conservation (2013)
C. Cianfrani et al.
trasting results (Thorn et al., 1997; McGarvey, Johnston &
Barber, 2010). As a rough surrogate for fish assemblages
and abundance we included altitude in the model (White
et al., 2003; Remonti, Balestrieri & Prigioni, 2009). For
PCBs contamination, only scattered point measurements
were available for Switzerland and, because of the complex-
ity of modelling pollutant fluxes in freshwater systems, these
data could not be included in the models.
Obtaining these information in a spatially explicit way
should constitute a priority field for future freshwater
research in Switzerland.
Nevertheless, available PCB data can provide a general
idea of the freshwater contamination levels in Switzerland.
Data indicate that contamination by PCBs may still be too
high (Weber, 1990; Michelot et al., 1998; Boscher et al.,
2010; Fig. S4 and Table S4 in Supporting Information
Appendix S4) to allow a proper return of the otter in the
country. Similarly, qualitative data on fish in Switzerland
reveal that fish biomass is alarmingly low in Switzerland
(Fischnetz, 2004) and may further prevent the establishment
of viable populations in many river basins. These issues
require further investigation to allow a definitive statement
for the otter. Finally, the social acceptance information was
difficult to include in the models because it was not available
for Austria (calibration area) and cannot be easily made
spatially explicit. In particular, this social study was prelimi-
nary and only aimed at testing the overall acceptance of an
otter comeback in Switzerland. A more exhaustive social
study should be conducted to make it more spatially explicit
and then possibly be incorporable in the models.
Would the otter be welcome
to Switzerland?
Concerning the hunting and social acceptance dimension,
the species is now under strict protection laws. Therefore,
only the question of social acceptance remains. Independent
results from a preliminary survey on the socio-economic
perception of a species return in Switzerland (Mateus, 2011;
see Supporting Information Appendix S5 for the details)
revealed quite high percentages of favourable perceptions,
even by fishermen and fish farmers, those stakeholders most
susceptible to experiencing damaging losses from an otter
recovery. In the same survey, most stakeholders stressed the
importance of revitalizing Swiss rivers as a first step, so as to
increase fish biomass and reduce PCB pollution. For these
stakeholders, under the present conditions of low levels of
fish biomass and high levels of PCB contamination, a
reintroduction of the otter would not be viable. The deter-
mination of stakeholder attitudes about a potential return
of a carnivorous species is essential to anticipate future con-
flicts (Treves, 2009).
Conclusion and perspectives
Our study proposed a three-step approach to assess the
recovery of a species in an area where it was formely present
2013 The Zoological Society of London
C. Cianfrani et al.
but now extirpated, based on the incorporation of connec-
tivity analyses into traditional habitat suitability assess-
ments. As few studies so far have considered all these facets
within a same conservation case (Morrel, 2008), our study
proved novel in this regard. By applying it to the case of the
otter in Switzerland, we reached the following conclusions:
(1) From a climatic and land-use perspective, suitable habi-
tats are present, reasonably well interconnected and con-
nected with existing populations outside Switzerland.
(2) However, some key factors like water contamination
and food resources could not be included in our analyses
and would need to be carefully considered in addition to our
predictions in any future otter recovery plan.
(3) Despite the fact that the otter would be socio-
economically welcome and that suitable and well-connected
habitats could support its return, current levels of PCB and
fish density seem still too critical for a full recovery of the
otter in Switzerland.
(4) Future studies should aim at improving existing spatial
data on fish denstity and water quality for freshwater bodies
in Switzerland, and use these to improve estimates of habitat
suitability and connectivity.
Acknowledgements
W. Hordik helped with the elaboration of the ensemble
models; S. Matteus performed the human dimension study;
H. Satizbal provided useful comments on the paper. This
study was supported by the Swiss Federal Office, MAVA
Foundation and ECOCHANGE project (EU 6th Frame-
work Programme; contract number FP6-036866).
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Supporting information
Additional Supporting Information may be found in the
online version of this article at the publishers web-site:
Appendix S1 Habitat Suitability
Appendix S2 Model evaluation
Appendix S3 Conductance matrix
Appendix S4 Water contamination (PCB)
Appendix S5 Social perception of the otters return
Appendix S6 (a) Otter occurrence for the Alps sector, at a
1-km resolution, which were used to develop the bioclimatic
suitability model. (b) Otter occurrence data in Austria
that were used to develop the environmental suitability
model.
11