Ecological Indicators 34 (2013) 372–379

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Ecological Indicators
journal homepage: www.elsevier.com/locate/ecolind

Landscape heterogeneity metrics as indicators of bird diversity:

Determining the optimal spatial scales in different landscapes
Federico Morelli a,∗ , Fabio Pruscini a , Riccardo Santolini a , Paolo Perna b ,
Yanina Benedetti a , Davide Sisti a
a
University of Urbino, Campus Scientifico Enrico Mattei, 61029 Urbino, Italy
b
Terre.it, Spin-off of the University of Camerino, 62032 Camerino, Italy

a r t i c l e i n f o a b s t r a c t

Article history: Species distribution models are often used to study the biodiversity of ecosystems. The modelling process
Received 28 February 2013 uses a number of parameters to predict others, such as the occurrence of determinate species, population
Received in revised form 23 May 2013 size, habitat suitability or biodiversity. It is well known that the heterogeneity of landscapes can lead to
Accepted 24 May 2013
changes in species’ abundance and biodiversity. However, landscape metrics depend on maps and spatial
scales when it comes to undertaking a GIS analysis.
Keywords:
We explored the goodness of fit of several models using the metrics of landscape heterogeneity and
Spatial scale
altitude as predictors of bird diversity in different landscapes and spatial scales. Two variables were used
Land cover map
Landscape heterogeneity
to describe biodiversity: bird richness and trophic level diversity, both of which were obtained from a
Bird richness breeding bird survey by means of point counts. The relationships between biodiversity and landscape
Biodiversity indicators metrics were compared using multiple linear regressions. All of the analyses were repeated for 14 differ-
GIS ent spatial scales and for cultivated, forest and grassland environments to determine the optimal spatial
scale for each landscape typology.
Our results revealed that the relationships between species’ richness and landscape heterogeneity using
1:10,000 land cover maps were strongest when working on a spatial scale up to a radius of 125–250 m
around the sampled point (circa 4.9–19.6 ha). Furthermore, the correlation between measures of land-
scape heterogeneity and bird diversity was greater in grasslands than in cultivated or forested areas.
The multi-spatial scale approach is useful for (a) assessing the accuracy of surrogates of bird diversity in
different landscapes and (b) optimizing spatial model procedures for biodiversity mapping, mainly over
extensive areas.
© 2013 Elsevier Ltd. All rights reserved.

1. Introduction
In the face of continuous global changes, understanding the
way in which many factors affect biodiversity is becoming increas-
ingly relevant (Coblentz and Riitters, 2004). The role of biodiversity
is very important in the conservation of functioning ecosystems.
Conservation biologists now recognize that the biodiversity issue
involves more than just species’ diversity or endangered species
(Noss, 1990). Moreover, biodiversity can be studied in different
ways taking into account species’ richness (Alpha, Beta and Gamma
biodiversity; Whittaker, 1977) and taxonomic, phylogenetic and
functional diversity (Devictor et al., 2010). However, the study of
biodiversity is particularly complex, and requires a large amount of
time, a great deal of effort and significant funds. In many cases, the
data on biodiversity are hard to collect, and for this reason it is very
∗ Corresponding author. Tel.: +39 0722304251.
E-mail addresses: federico.morelli@uniurb.it, morellius@libero.it (F. Morelli).
useful to have a tool, or a set of tools, with which to estimate it indi-
rectly. One approach is to identify measurable attributes that are
used as surrogates or indicators, taking them from environmen-
tal inventories, monitoring work, existing maps, and assessment
programmes (Noss, 1990).
Indicators are measurable surrogates of environmental char-
acteristics such as biodiversity. Ideally, an indicator should be:
sufficiently sensitive to provide an early warning of environmental
change; widely applicable; able to provide a continuous assessment
over a wide range of stress factors; relatively independent of the
size of the sample; easy and cost-effective to measure or collect;
able to discriminate between natural cycles or trends and those
that are induced by anthropogenic stress; and relevant to ecologi-
cally significant phenomena (Caro and O’Doherty, 1999; Jeffrey and
Madden, 1991; Munn, 1988; Noss, 1990; Sheehan, 1984). The use
of indicators should be part of a comprehensive strategy to analyze
landscape quality by focusing on key habitat characteristics that
include corridors, mosaics and other landscape structures, as well
as species (Landres et al., 1988; Paoletti, 1999).

1470-160X/$ – see front matter © 2013 Elsevier Ltd. All rights reserved.
http://dx.doi.org/10.1016/j.ecolind.2013.05.021
 F. Morelli et al. / Ecological Indicators 34 (2013) 372–379
Fig. 1. Distribution of sampled sites (black dots) with the breeding bird atlas data in the Marche region, Central Italy.
Land use or landscape parameters can be used as indicators of
landscape heterogeneity, and potentially as indicators of biodiver-
sity (Lindenmayer et al., 2002). One of the possible cost-effective
surrogates for obtaining appropriate ratings of spatial patterns for
species’ richness is provided by predictive modelling based on
remote sensing and topographic data (Luoto et al., 2004). In par-
ticular, land cover metrics could be regarded as good surrogates
of species’ diversity because, in ecology, habitat diversity is associ-
ated with an increase in niche availability for species (Kisel et al.,
2011). There is a growing body of literature demonstrating that
habitat fragmentation and the heterogeneity of landscapes can
lead to changes in species’ abundance and biodiversity (Andrén,
1994; Fuller et al., 1997; Morelli et al., 2012; Suarez-Rubio and
Thomlinson, 2009).
The opportunity to acquire tools that operate as indicators of
biodiversity is very important for management policies and the
conservation of habitats and species (Girardello and Morelli, 2012;
Morelli et al., 2007; Tobolka et al., 2012). In order to speed up the
decision-making process, and to estimate the areas with the great-
est species’ richness, we used environmental information that is
usually obtainable from current maps. However, the use of sev-
eral measures and tools is important because, as well documented
in recent literature on ecological models (Noss, 1990; Hawrot and
Niemi, 1996; Luoto et al., 2004; Thuiller et al., 2004), they could
have different levels of predictive accuracy depending on the type
of landscape, environment, thematic resolution and spatial extent
of the map from which the predictors were derived (Brambilla et al.,
2009; Chust et al., 2003; Debinski et al., 2001; Morris, 1987; Trani,
2002; Wiens, 1989; Wiens et al., 1993). For this reason, when mod-
elling bird species’ diversity and distribution, it is very important to
determine the best spatial scale for calculating the environmental
data (Cornell and Lawton, 1992; Kuczynski et al., 2010). Moreover,
the quality of regional or local environmental data is always con-
ditioned by the spatial resolution of the available maps (vegetation
maps, land use and land cover maps, etc.).
In order to analyze how bird species’ diversity in communities
could be an expression of landscape and land use characteristics,
and using bird data as response variables on different spatial scales,
we studied the reliability of several of these parameters, which are
often used as surrogates of biodiversity. A regional land use map
was utilized in the geographic information systems (GIS) project,
because it is considered to be the most common source of envi-
ronmental data on a territory, while the bird data were utilized
because they are largely used in the literature and are available
around the world (Brown et al., 1995; Gibbons et al., 1993; Mace
et al., 2010; Osborne and Tigar, 1992; Van Strien et al., 2001). Fur-
thermore, the accuracy of each parameter and the goodness of fit
373
of a set of modelled parameters were compared at different spatial
scales to select the best spatial scale for modelling purposes.
2. Methods
2.1. Study area
The study was carried out during 2011 with the study area being
the entire Marche region in Central Italy. This region covers a total
surface of 9.366 km2 (centroid: 43◦ 23 34.63 N, 13◦ 15 32.56 E)
(Fig. 1) and is at an altitude ranging from 0 to 2.476 m a.s.l. (Mt.
Vettore, Sibillini Mounts). The climate in Central Italy is tem-
perate (Tomaselli et al., 1972) and characterized by high spring
and summer temperatures and a marked summer drought. The
landscape in the study area consists mainly of cultivated fields
(52%), forest (20%), grasslands (10%) and other typologies (18%)
(AA.VV., 2008).
2.2. Bird data
The survey of birds was performed between mid-April and the
end of July 2011. Point counts (Bibby et al., 1997) were located
uniformly over the entire Marche region (Fig. 1) and were at least
1000 m apart. The points were visited between 06.00 and 10.00 h
for 10 min, and all of the birds detected both visually and acous-
tically within a radius of 100 m from the observer were recorded.
Only breeding species were considered in this paper. Bird species’
richness (BR) was quantified as the total number of bird species
recorded per site (Gaston, 1996), because all sites were surveyed
with the same sampling effort.
Other data were elaborated on by performing a calculation of
the diversity of the trophic level (DTL) to provide more information
about the composition of the bird communities. This parame-
ter was used as another indicator of bird community diversity,
because higher trophic levels should testify to unaltered food
chains and, therefore, communities, ensuring more ecological func-
tions (Mouysset et al., 2012; Sergio et al., 2005). Information about
the trophic level for each bird species was extracted from the Com-
plete Birds of the Western Palearctic CD-ROM v.1, 1998 (copyright
Oxford University Press) and the British Trust for Ornithology (BTO)
website (www.bto.org). The trophic level was classified into five
categories (1 = eat vegetal; 2 = eat invertebrate; 3 = eat vegetal and
invertebrate; 4 = eat vertebrate and invertebrate; and 5 = eat vege-
tal, invertebrate and vertebrate). Using these categories, the trophic
level diversity for each sampled site was calculated by utilizing
 the
Shannon–Weaver diversity index with the formula H = pi ln pi ,
where the different pi are the proportions of the different trophic
374 F. Morelli et al. / Ecological Indicators 34 (2013) 372–379
Table 1
Bird diversity, landscape heterogeneity metrics and the altitude of sites in the different environments studied. The values are expressed as the mean and SD in parentheses.
The values of the landscape heterogeneity metrics in the table were calculated for the spatial scales of a radius of 250 m around the sampled points. The variables’ codes are
as follows: BR: bird richness; DTL: diversity of trophic level; LCF: land cover features; LCR: land cover richness; LCD: land cover diversity; WEDG: weighted edge density;
EDG: edge density; and LPI: largest patch index.
Environment N Bird diversity Landscape heterogeneity metrics and altitude
BR DTL LCF
Cultivated 922 12.3 (4.05) 0.67 (0.28) 19.20 (9.41)
Forest 480 9.57 (3.09) 0.65 (0.30) 12.24 (8.22)
Grassland 286 5.40 (3.33) 0.37 (0.34) 8.53 (7.9)
Total 2091 10.5 (4.33) 0.62 (0.31) 16.17 (10.1)
level components of the bird community (Shannon, 1949). The
response variables (BR and DTL) were calculated using all of the
bird species recorded per site.
2.3. Landscape, environmental and ecological parameters
Land cover data for the study area were obtained from the land
cover map of the Marche region (1:10,000) (AA.VV., 2008). This
was chosen as it is a data set that has been classified and validated
and is readily available on a national level. The ArcGIS 9 software
(ESRI, 2009) was used to delineate the polygons with a variable
radius around the sampled site, and Fragstats 3.2 and ArcGIS 9 were
used to derive and process the landscape heterogeneity metrics
(Schindler et al., 2008).
In order to describe the landscape heterogeneity, a multi-scale
approach was used by taking the following steps: (a) the creation
of a series of increasing buffer zones around each survey point as
multiple spatial scales; (b) using overlap buffers with a land cover
regional map; and (c) using an intersect operator to calculate the
following landscape metrics:
• Land cover features (LCF): number of total polygons within the
buffer.
• Land cover richness (LCR): number of different typologies of land
use within the buffer (L).
• Land cover diversity (H ) (LCD): calculated using the Shannon-
Weaver diversity index (see formula above) on land use types.
• Edge density (EDG): sum of the perimeters of all polygons in the
buffer zone.
• Weighted edge density (WEDG): sum of the perimeters of all
polygons in the buffer zone per number of land use types/buffer
surface (Hargis et al., 1998).
• Largest patch index (LPI): percentage of total area occupied by
the largest patch.
These parameters were calculated for different spatial scales
(radiuses of 50, 125, 250, 375, 500, 625, 750, 875, 1000, 1125,
1250, 1375, 1500 and 2000 m). The initial spatial scale (spatial
extent), which was fixed to start the analysis, was a 50 m radius.
This is because the environmental information extracted from the
regional maps at a lesser spatial scale becomes inaccurate (at the
lowest spatial resolution many land use typologies are poorly rep-
resented on a 1:10,000 map). A landscape description at a specified
scale also contains information about landscapes at lower scales,
because these parameters were calculated as spatial indices in spa-
tially nested landscapes surrounding each sampling site (Bergin
et al., 2000).
The sampling sites were classified as cultivated (arable fields),
forest or grassland when the main land cover was >50%. The altitude
was recorded for each sampling site as one measure in the centre
of the point count using a GPS (accuracy rate: ±15 m).
LCR LCD WEDG EDG LPI Altitude
6.52 (1.67) 1.05 (0.36) 0.31 (0.13) 0.045 (0.01) 0.15 (0.05) 247.26 (193.58)
4.23 (1.75) 0.66 (0.36) 0.17 (0.11) 0.035 (0.01) 0.11 (0.08) 757.11 (300.37)
3.24 (1.71) 0.50 (0.35) 0.11 (0.10) 0.031 (0.01) 0.21 (0.09) 1176.64 (379.34)
5.64 (2.28) 0.95 (0.46) 0.26 (0.16) 0.041 (0.01) 0.18 (0.12) 536.64 (435.13)
2.4. Statistical analysis
Two response variables were regarded as the surrogates of bio-
diversity: bird richness (alpha diversity) and trophic level diversity
in bird communities. The predictor variables considered, i.e. the
environmental parameters, were four scale-dependent variables:
land cover features, land cover richness, land cover diversity, and
edge density; and one non-scale dependent variable: altitude.
The spatial autocorrelation was checked using a Mantel test,
which compares two similarity of distance matrices computed for
the same sites (Legendre and Legendre, 1998). The test was per-
formed on paired matrices containing the correlation coefficients
between the response variables and the geographical distance
between each pair of points.
A first screening of the relationships between the bird data and
the environmental data was performed using the GIS tool “inverse
distance weighted” (IDW) technique that interpolates a raster sur-
face from several point values (bird species’ richness and landscape
heterogeneity measures) and distances between the points. How-
ever, because the output value for a cell using the IDW technique is
limited to the range values utilized, this procedure was only used
for preliminary descriptive purposes.
In order to explore the relationship between the response
variables (bird richness or trophic level diversity) and the environ-
mental parameters, multiple linear regressions were applied for
each spatial scale (ranging from a radius of 50 to 1500 m around
the buffer). To stabilize variance and ensure a more normal distri-
bution of the dependent variables, Box–Cox transformations were
used where appropriate (Box and Cox, 1964).
Linear regressions were also performed on subsets of the sam-
pled sites classified as cultivated areas, forest and grassland. To
determine the relative contribution of each independent variable
(landscape or altitude) when accounting for variation in bird rich-
ness, zero-order correlation coefficients of the response variables
were considered for each model. The goodness of fit of multiple
linear models (McCullagh and Nelder, 1989) was determined using
multiple R correlation coefficients (Chust et al., 2003). Further-
more, the best model was evaluated by plotting observed against
predicted species’ richness values. All tests and elaborations were
performed with the R program (R Development Core Team, 2011)
and SPSS v.19.0.
3. Results
3.1. Multi-scale performance of landscape heterogeneity metrics
as indicators of bird diversity
During this study, collected bird species’ presence data from
2091 sites were sampled (Fig. 1). The sampled sites were treated as
independent units, because the spatial autocorrelation values were
low (relationship between geographic distance and biodiversity
or the environmental dissimilarity of sites: Mantel test r = −0.018,
 F. Morelli et al. / Ecological Indicators 34 (2013) 372–379 375

Fig. 2. Distribution of bird diversity and landscape heterogeneity metrics in the Marche region using the inverse distance weighted (IDW) technique. The values are
represented in a coloured-scale from green (lowest values) to red (highest values). (For interpretation of the references to colour in this figure legend, the reader is referred
to the web version of the article.)

nbootstrap = 999, p = 0.974; Betts et al., 2006). A total of 79 bird species

were recorded in the study area. The mean bird richness was 10.5
species, with the range being two to 31 species (Table 1). The
variable bird richness was transformed using the square root trans-
formation, as suggested by the Box–Cox plot (value = 0.61, near to
0.5).
To demonstrate the graphic relationships between response
variables and predictors (Fig. 2), the inverse distance weighted
(IDW) measure was used in the GIS to interpolate a raster
surface from our sampled point values (bird diversity and land-
scape heterogeneity measures) and distances between these
points.
Separate tests on relationships between the response and
explanatory variables revealed that bird species’ richness and
trophic level diversity were significantly related to five of the six
variables (p < 0.05). The correlation coefficients (R2 ) between bird
richness and the predictors from the initial spatial scale analyzed
(radius of buffer 50 m) were: 0.35 for land cover features, 0.43 for
land cover richness, 0.38 for land cover diversity, 0.42 for edge den- Fig. 3. Comparison among the mean values of the coefficients of correlation (R)
sity and −0.59 for altitude. Meanwhile, the correlation coefficients between bird richness and landscape heterogeneity metrics.
between trophic level diversity and the predictors for the same
spatial scale were: 0.13 for land cover features, 0.22 for land cover
richness, 0.14 for land cover diversity, 0.21 for edge density and way, decreasing the spatial scale (e.g. 50 m radius) also decreased
−0.36 for altitude. Using all of the spatial scales, bird richness was the predictive power.
always more correlated with the predictors than with trophic level
diversity. For this reason, only bird richness was preferred for fur- 3.2. Performance of landscape heterogeneity metrics as indicators
ther analysis. of bird diversity in different landscapes
With respect to the mean values for all spatial scales, the land-
scape heterogeneity metrics that were most correlated to bird The mean bird richness was different in different landscapes.
richness were land cover richness, weighted edge density and edge The highest values of species’ richness were found in cultivated
density (Fig. 3). However, the predictive accuracy of the variables environments, followed by forest and grassland (Table 1). More-
and linear regression models varied among the spatial scales, and over, the partial contribution of the independent variable in each
also in relation to different landscapes. spatial scale model varied in the different landscapes. Altitude
The best linear regression model of bird richness using land- was more important when it comes to explaining bird richness in
scape heterogeneity metrics accounted for more than 39% of the a grassland environment when compared to cultivated land and
variance (r = 0.64) of the response variables (p < 0.001) (Fig. 4). The forest (determination coefficient values: −0.54 (p < 0.001); −0.24
maximum values of the determination coefficient were recorded (p < 0.001); and −0.14 (p = 0.002) respectively).
at a radius of 125 m that corresponded to the optimal spatial scale In summary, the models that relate bird richness to landscape
for modelling bird richness (Fig. 5). However, the determination heterogeneity metrics and altitude seem to be more accurate when
coefficients for all of the linear models were lowest as a predictive performed on grassland with a spatial scale radius of 125–250 m
power when increasing the spatial scale of analysis. In the same (Figs. 5 and 6). The predictive power of all of the predictors and
376 F. Morelli et al. / Ecological Indicators 34 (2013) 372–379
Fig. 4. Predicted values from multiple-regression models of bird richness and land-
scape heterogeneity against observed values at a spatial scale of 125 m (R2 = 0.39,
p < 0.001). The values in the axis are the square root transformations of the response
variables.
linear regression models was highest in grasslands. A minor accu-
racy was recorded in cultivated environments.
4. Discussion
Over the last few years, the diffusion of digital maps has
increased the opportunities to apply biodiversity models to broad
areas, enabling changes in landscapes to be related to poten-
tial changes in patterns of species’ richness (Coops et al., 2009;
Griffiths et al., 2000; Scott et al., 1993; Stoms and Estes, 1993).
The use of remote sensing approaches due to their multi-scale
capabilities is a helpful tool with which to fill the gap between
intensive ecological research and landscape management (Estes
et al., 2010; Gulinck et al., 2000). In fact, in recent years, several
studies have been developed to quantify biodiversity, with the
focus on the use of environmental data derived from existing maps
(Guisan and Zimmermann, 2000; McCain, 2005; Rushton et al.,
2004). For example, landscape heterogeneity metrics were used
Fig. 5. Performance of the best model using landscape heterogeneity metrics as indicators of bird richness at different spatial scales and on the basis of landscape type.
to study relationships with avian diversity in agricultural ecosys-
tems in Spain. In this study had demonstrated how the total bird
species richness is mainly related to landscape diversity (Pino et al.,
2000). Studies of this type are needed because it is very important to
understand functioning farmland, as well as the actions that can be
taken to reduce the decline of bird populations in agro-ecosystems
(Tryjanowski et al., 2011).
However, the insight drawn from such models depends on the
quality and quantity of the input data. It is also important to
consider the fact that in ecology, the scale of the habitat generally
varies with species and wider home ranges (Cornell and Lawton,
1992). Accordingly, it is important to determine the best spatial
scale for the environmental data recorded if we are to improve the
prediction of biodiversity.
On the basis of our results, the most useful landscape hetero-
geneity metrics for studying bird diversity are: land cover richness,
weighted edge density and edge density as scale-dependent vari-
ables; and altitude as a non-scale dependent variable. These
environmental parameters take into account an important quan-
tity of variation in species’ richness (more than 39% in some cases).
The dependence of bird species’ richness on these landscape het-
erogeneity metrics confirms that a high degree of species’ richness
can be indicated by a surrounding landscape that is characterized
by greater heterogeneity and high edge density. Such an outcome
is expected according to the habitat heterogeneity hypothesis (e.g.
McArthur and Wilson, 1967) and due to increases in niche avail-
ability (Kisel et al., 2011).
Fuller et al. (1994) suggested that a useful scale for using the
land cover map in ecological modelling is upwards of 1.5 ha or
200 m along a linear dimension. Our results also demonstrate
that for studies of bird richness by means of environmental data
obtained from land cover maps at a 1:10,000 resolution, spa-
tial scales under 50 m are neither useful nor precise enough
for building models. The optimal spatial scale around the sam-
pled sites for deriving the landscape metrics for studying bird
species’ richness was a radius of 125 m (about 4.9 ha). Further-
more, given that we worked with passerine birds, which also
have variable home-ranges during the breeding period, the opti-
mal spatial extent (making a compromise between accuracy and
the dimension of patches) ranged from a radius of 125 to 250 m
around the sampled site (4.9–19.6 ha). Similar work was also
 F. Morelli et al. / Ecological Indicators 34 (2013) 372–379 377

Fig. 6. Performance of each variable of the landscape heterogeneity metrics: as indicators of bird richness; as multiple coefficients of correlation (R) at different spatial scales;
and on the basis of landscape type.

carried out in forest landscapes in Greece to evaluate the rela-
tionships between landscape metrics and species’ richness for
several taxa (vegetation, insects, amphibians, reptiles and small
terrestrial birds) at a number of spatial scales. There, the authors
made similar findings, highlighting how spatial scale affects the
performance of landscape metrics as indicators of biodiversity,
and finding also that spatial scale affect the performance of the
metrics for modelling small terrestrial birds, improving the predic-
tively at smaller extents of surrounding landscape (Schindler et al.,
2013).
In this study, we detected that landscape heterogeneity metrics
as indicators of species’ richness have performed better in some
landscapes than in others. Indeed, this fact is of special interest and
has implications for landscape conservation planning. Bird species’
richness was most sensitive to variations in landscape heterogene-
ity metrics in grasslands, where the highest model performances
in terms of predictivity were recorded. Grasslands may consti-
tute vulnerable ecosystems because they have the lowest values
of bird richness and, at the same time, the lowest habitat hetero-
geneity. In spatial terms, the nature of the habitat matrix in these
378 F. Morelli et al. / Ecological Indicators 34 (2013) 372–379
landscapes has a bearing on different ecological niches (e.g. edge
effects are less pronounced because there are fewer heterogeneous
landscapes; Kisel et al., 2011). However, the assessments of regional
grassland bird habitats must be conducted with multiple scales
because, according to other studies (Bakker et al., 2002; Morelli,
2012), some species only respond to local conditions within grass-
lands, whereas others respond to habitat structure at landscape
scales.
The fact that cultivated landscapes have a higher degree of bird
species’ richness could be explained by the presence of hedgerows,
uncultivated patches and point elements such as shrubs, all of
which can increase landscape complexity and, then, ecological
niches, making them available for a greater number of species (Kisel
et al., 2011; Norton et al., 2009). However, local studies are also
important because, as Tryjanowski et al. (2011) highlighted, farm-
lands are different in the various parts of Europe. This means that
to understand how bird populations are related to habitat struc-
ture, and later to real conservation, clear data with a good local
background are required.
In conclusion, we confirm that the use of a 1:10,000 scale land
cover map, which is often used by regional administrations, has
good potential for modelling and predicting bird species’ diver-
sity patterns, as demonstrated in the bibliography (Salem, 2003).
However, the selection of suitable landscape parameters for mod-
elling biodiversity must take into account all of the considerations
expressed earlier (the optimal spatial scale and mainly the land-
scape type that affect the predictive power). On the other hand,
the kind of limitations that are related to the use of bird data
from atlases is due to the fact that bird communities may fluctu-
ate from year-to-year. Furthermore, the availability of quantitative
data could be used to better describe the relationships between
birds and landscape metrics. Yet, an analysis that is even more
meaningful for conservation could be conducted on the number
of rare or declining species (conservation concern of species in the
sites), rather than on the total number of bird species.
Our findings clearly underline that landscape heterogeneity
metrics (relatively available and easy to obtain) can be a useful
tool for multidisciplinary frameworks when it comes to integrating
ecological approaches into conservation management plans. The
methodologies and constraints described in this and other recent
work (Schindler et al., 2013; Wiens et al., 2008) are useful for
modelling and have implications for landscape and environmental
management over extensive areas.
Acknowledgements
We are very grateful to Maria Balsamo, Federico Bastianelli,
Joseph McCulloch, Mario D’Atri, Raffaele Secchi and Roberto
Mazzeo for their important suggestions and contributions in the
work. We also thank ION Proofreading for the critical revision of
English version of the manuscript.
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