Popul Ecol (2011) 53:351–359
DOI 10.1007/s10144-010-0229-2
ORIGINAL ARTICLE
A frequency distribution approach to hotspot identification
Valerio Bartolino • Luigi Maiorano •
Francesco Colloca
Received: 9 January 2010 / Accepted: 22 June 2010 / Published online: 24 July 2010
Ó The Society of Population Ecology and Springer 2010
Abstract We present a new global method for the iden-
tification of hotspots in conservation and ecology. The
method is based on the identification of spatial structure
properties through cumulative relative frequency distribu-
tions curves, and is tested with two case studies, the
identification of fish density hotspots and terrestrial verte-
brate species diversity hotspots. Results from the frequency
distribution method are compared with those from standard
techniques among local, partially local and global methods.
Our approach offers the main advantage to be independent
from the selection of any threshold, neighborhood, or other
parameter that affect most of the currently available
methods for hotspot analysis. The two case studies show
how such elements of arbitrariness of the traditional
methods influence both size and location of the identified
hotspots, and how this new global method can be used for a
more objective selection of hotspots.
Keywords Cumulative frequency distribution 
Density  Diversity  Spatial clustering  Tangent
V. Bartolino (&)  L. Maiorano  F. Colloca
Department of Animal and Human Biology,
Sapienza University of Rome, viale dell’Università 32,
00185 Rome, Italy
e-mail: valerio.bartolino@uniroma1.it
V. Bartolino
Marine Research Institute, Swedish Board of Fisheries,
PO Box 4, 45321 Lysekil, Sweden
L. Maiorano
Department of Ecology and Evolution, University of Lausanne,
Biophore Building, 1015 Lausanne, Switzerland
Introduction
The analysis of spatial data is a fundamental aspect of
ecological and conservation studies (Legendre and Fortin
1989; Haining 2003; Fortin and Dale 2005). This is par-
ticularly important for the identification of hotspots, i.e.,
spatial clusters where an out of the ordinary event or
phenomenon occurs (Nelson and Boots 2008). The defini-
tion of hotspots in ecology is typically related to species or
process abundance (Weathers et al. 2000; Houghton et al.
2006; Kissling et al. 2007) as a first step towards under-
standing mechanisms that generate the observed spatial
patterns. In conservation, hotspots are usually considered in
the context of biodiversity (i.e., Myers et al. 2000; Orme
et al. 2005) or abundance of species of particular concern
(i.e., Bonn et al. 2002; Houghton et al. 2006) in the attempt
to develop and maximize the effectiveness of both large
scale (Roberts et al. 2002; Worm et al. 2003; Kerswell
2006) and small-scale (Williams et al. 1996; Maiorano
et al. 2007) conservation and management strategies.
Although the definition of a hotspot could appear simple
and straightforward, it is generally not easy to define
objectively where the boundaries among hotspots and non-
hotspots occur (Fortin and Dale 2005). A wide array of
statistical techniques has been developed for this purpose,
and three main approaches can be distinguished: spatially
global, spatially local, and partially local methods (Nelson
and Boots 2008).
Spatially global methods compare the values measured
in single locations with those measured over the entire
study area. The most common application of this approach
has been the definition of biodiversity hotspots for con-
servation purposes (Myers et al. 2000). In most cases, a
subjective threshold is applied to a spatially explicit spe-
cies-richness dataset to identify the areas with the highest
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richness values that should be managed or conserved.
Usually the selected threshold ranges between 1 and 5%
(Myers et al. 2000; Gjerde et al. 2004; Grand et al. 2004;
Orme et al. 2005), but it has also been raised to 25% when
the identification of wider areas was required (Garcia
2006).
Spatially local methods compare the values measured in
particular locations with those measured in the immediate
surrounding areas. In this case, the definition of what
represents the neighborhood for the observation is usually
determined by subjective decision (Atkinson and Unwin
2002; Haining 2003; Nelson and Boots 2008). The most
commonly used spatially local methods are kernel esti-
mation (Fortin and Dale 2005) and local measures of
spatial autocorrelation, like Moran’s Ii and Getis’ Gi
(Nelson and Boots 2008). Kernel estimators have been
widely used in ecological studies, mainly for home range
detection (e.g., Horne and Garton 2006), while Getis’ Gi*
and Moran’s Ii have been used especially in socio-eco-
nomic studies (e.g., Anselin et al. 2005; Moons et al. 2008).
Partially local methods use a combination of local and
global approaches, with a spatially global threshold that is
applied to a spatially local measure (Nelson and Boots
2008). For example, Grand et al. (2004) used a 5% global
threshold to identify bird and moth hotspots inside their
study area; however, in areas of particularly high species
richness, they raised the threshold value to capture in their
hotspots only the highest species counts, introducing a
local component.
Spatially local methods are usually more effective for
hotspot analyses compared to spatially global ones, espe-
cially when the area considered is large and the processes
being mapped are non-stationary (Nelson and Boots 2008).
Moreover, the smoothing effect of spatially local methods
can be important when the neighborhood structure has a
biological or practical relevance (e.g., structural connec-
tivity in a metapopulation context, known level of uncer-
tainty in the population data, etc.). Moran’s Ii and Getis’ Gi*,
in particular, are useful to identify clusters of values that are
high relative to a global mean; moreover, they are usually
more robust than kernel estimators, being less affected by
the choice of neighborhood and thus more appropriate when
only limited knowledge is available about the phenomenon
under consideration (Nelson and Boots 2008).
Although the most appropriate method should be
selected according to the specific objectives and applica-
tions considered, all the approaches briefly described above
suffer from one or more elements of subjectivity. Global
and partially local methods are based on the selection of a
pre-defined threshold that has little, if any, ecological jus-
tification, and that heavily influences the results. Local
methods, in general, work better with normally distributed
data although several corrections have been suggested for Ii
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Popul Ecol (2011) 53:351–359
and Gi* to handle non-normality (e.g., Tiefelsdorf 2002;
Griffith 2010). Simulation approaches are also commonly
used to assess significance in relation to a null hypothesis
of spatial randomization (Anselin 1995). Moreover, local
methods are affected by the identification of the appropri-
ate range of influence, i.e., by the smoothing factor in
kernel analyses (Horne and Garton 2006) and by the
neighborhood structure in local measures of spatial auto-
correlation (Fortin and Dale 2005; Nelson and Boots 2008).
Even though some statistical procedures can help in the
choice of an appropriate neighborhood/range of influence
(Haining 2003; Nelson and Boots 2008), an important
component of arbitrariness still remains that potentially
affects both size and location of the identified hotspots.
Materials and methods
A frequency distribution approach to hotspots
identification
We propose a new spatially global method which can be
used to identify hotspots for conservation or ecological
purposes. Our approach can be easily used for any phe-
nomenon that is distributed over spatially autocorrelated
surfaces, and is based on cumulative relative frequency
distribution (CRFD) curves. A CRFD curve f(x) is
approximated plotting the relative value of the variable
z (i.e., density, biodiversity, etc.), against the frequency
distribution of the same variable. The two axes of the plot
are given by:
z
x¼ ð1Þ
zm
where z is the variable of interest and zm is the maximum
value that z assumes in the study area, and
MðxÞ
y¼ ð2Þ
N
where M(x) is the number of cases (i.e., points, pixels,
areas) a value smaller than x is encountered, and N is the
total number of cases. The resulting f(x) curve is charac-
terized by having both axes ranging from 0 to 1.
The tangent to the curve f(x) at a point (x0, f(x0)) has
slope f 0 (x0), corresponding to the derivative of f(x) in x0,
and is calculated as:
y  f ðx0 Þ
f 0 ðx0 Þ ¼ ð3Þ
x  x0
A tangent with \45° slope (f 0 (x0) \ 1) will indicate
areas where the relative increase along the y-axis (the
cumulative relative frequency) is smaller than the relative
increase along the x-axis (the variable of interest), i.e.,
Popul Ecol (2011) 53:351–359
those areas where the variable considered can be found at
high densities. A tangent with [45° slope (f 0 (x0) [ 1) will
indicate areas where the relative increase along the y-axis
is greater than the relative increase along the x-axis, i.e.,
those areas where the variable considered can be found at
low densities. A tangent with 45° slope (f 0 (x0) = 1) will
indicate areas where the relative increase along the y-axis
is equal to the relative increase along the x-axis, i.e., those
areas where the variable considered can be found at
densities proportional to the frequency.
In practice, the tangent to the curve f(x) at each point can
be seen as the accumulation rate for the phenomenon of
interest. The CRFD curve generated by a random process
will approximate a 45° line, but almost any biological
process will produce a complex CRFD curve approaching
an upper asymptote. We suggest that the highest x0 corre-
sponding to a 45° slope tangent to the curve can be used as
a global threshold to identify hotspots. In fact, the 45° slope
tangent, and no other arbitrary threshold, will discriminate
the areas characterized by the highest values of the variable
z and at the same time by a rate of accumulation along the
x-axis higher than that along the y-axis. Other spatially
global methods, and also the frequency distribution
approach, compare local values with the distribution of
values across the whole study area, regardless of the local
neighborhood structure. However, under the assumption of
spatial autocorrelation, the high values selected as hotspots
will not occur in isolation, and the frequency distribution
method will thus identify all the areas with the highest
values of the phenomenon of interest (i.e., species diversity
and/or species density).
We show in the remainder of the paper two applications
of the frequency distribution approach presented, one to
detect density hotspots for European hake (Merluccius
merluccius) along the western coast of Italy, and another to
Fig. 1 Map of the two study
areas for the hake recruit density
(dark gray) and the terrestrial
vertebrate biodiversity (light
gray) case studies
353
detect biodiversity hotspots for terrestrial vertebrates on the
Italian peninsula.
Case studies
We obtained a dataset on fish density and one on terrestrial
vertebrate species richness, respectively, from the pro-
ject BECAUSE (http://www1.uni-hamburg.de/BECAUSE,
accessed on 2 March 2009; Colloca et al. 2009) and
Maiorano et al. (2007). The project BECAUSE provided a
Bayesian kriging estimate of European hake recruit log-
densities (n/km2) off the central-western coast of Italy (FAO
geographical sub-area 9, covering 42,419 km2; Fig. 1) over
a 2 km by 2 km estimation grid (7,290 cells). Maiorano et al.
(2007) provided validated distribution models for 468
species of terrestrial vertebrates occurring in the Italian
peninsula (Fig. 1). The original models have a spatial reso-
lution of 100 m by 100 m (more than 30 million cells); for
computational reasons, we re-sampled the entire dataset with
a resolution of 10 km, obtaining a total of 3,115 cells. We
calculated species richness as the number of species occur-
ring in each cell.
We used the two datasets to compare hotspots identified
by the frequency distribution approach that we proposed,
with hotspots identified through some of the most widely
used techniques in ecology and conservation: Moran’s Ii,
used to calculate spatially local hotspots, Getis’ Gi*, for
partially local hotspots, and a set of predefined thresholds
for global hotspots.
To implement the spatially local and partially local
methods, we set for both datasets three neighborhood lev-
els: 8, 24, and 48 neighboring cells. For each of the three
neighborhood levels in the spatially local method we
defined as hotspots all those cells with a positive local
Moran’s Ii and P \ 0.01. In the partially local method, and
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354 Popul Ecol (2011) 53:351–359

a b c

Cumulative frequency

Frequency

Latiitude (°N)
Relative density Density Longitude (°E)

Fig. 2 a Cumulative relative frequency distribution curve of hake with the proportion of area (shade area) above the identified density
recruit density showing the 45° slope tangent to the curve (dotted line) threshold (dotted line). c Spatial distribution of hake recruit density
at the highest density value. b Frequency distribution of fish density hotspots from the frequency distribution method

Fig. 3 Spatial distribution of hake recruit density hotspots from Moran’s Ii (with 8, 24 and 48 neighboring cells), Getis’ Gi* (with 8, 24 and 48
neighboring cells), and global predefined thresholds (5, 10 and 15%)

for each of the three neighborhood levels, we applied a approach, we applied three arbitrary commonly used global
global threshold to select as hotspots all cells in the upper thresholds, considering as hotspots, respectively, the 5, 10,
5% of the Gi*-statistics distribution. In the spatially global and 15% highest values.

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Popul Ecol (2011) 53:351–359 355

To calculate the hotspots according to the frequency Table 1 Total number of hotspots of hake recruit density, percent-
distribution approach, we built for both datasets a CRFD age of the Tyrrhenian and Ligurian Seas identified as hotspots, and
percentage of overlap with the hotspots identified by the frequency
curve and we calculated the related 45° slope tangent.
distribution method (FD)
Because the curves were approximated by discrete fre-
quencies, the tangents to the curves were calculated as the % Hotspot % Overlap with FD Number of hotspots
lines passing through pairs of adjacent points. G8 5.0 100.0 9
G24 5.0 100.0 5
G48 5.0 100.0 4
Results Glob.thr 5 5.0 100.0 10
Glob.thr 10 10.0 100.0 8
The density hotspots for European hake recruits calculated Glob.thr 15 15.0 100.0 10
with the frequency distribution approach (Fig. 2) occupy
FD 15.3 – 9
approximately the same areas as those identified by the
I8 14.6 100.0 8
other three methods (Fig. 3; Table 1). All the methods
I24 19.8 77.1 9
identified density hotspots that are consistent with previous
I48 23.5 64.7 8
knowledge on the bathymetric and spatial distribution of
hake recruits in the study area (Ardizzone and Corsi 1997; Results obtained with three different neighborhood levels (8, 24 and
48 cells) for the Getis’ Gi* and the Moran’s Ii, three global thresholds
Abella et al. 2005; Bartolino et al. 2008). Hake recruits
(Glob.thr 5, Glob.thr 10 and Glob.thr 15%), and the frequency dis-
concentrate at the limit of the continental shelf and the tribution approach
upper slope, where elevated local productivity supports the
rich demersal communities of the shelf-break (Colloca Table 2 Total number of hotspots of vertebrate species richness,
et al. 2004). However, the number and size of the hotspots percentage of the Italian peninsula and islands identified as hotspots,
being defined was different for different approaches and percentage of overlap with the hotspots identified by the fre-
quency distribution methods (FD)
(Tables 1 and 2). We identified a minimum of 4 (using
Getis’ Gi* with 48 neighboring cells) and a maximum of 10 % Hotspot % Overlap with FD Number of hotspots
hotspots (using the 5 and 15% global thresholds) for hake
G8 5.0 31.4 15
recruits. The high percentage of overlap between the hot-
G24 5.0 26.3 8
spots identified with our method and all the other local,
G48 5.0 22.4 4
partially local, and global approaches (Table 1) suggested
Glob.thr 5 4.6 21.6 27
that the same peaks of fish density were detected as hot-
Glob.thr 10 9.8 32.7 45
spots. Hotspots identified through the frequency distribu-
Glob.thr 15 14.8 69.4 44
tion approach occupied 15.3% of the study area. Similar
FD 3.2 – 22
estimates for the area occupied by the hotspots were
I8 6.2 42.7 14
obtained with Moran’s Ii with 8 neighborhood cells
I24 15.5 18.7 10
(14.6%), and with the 15% global threshold (Table 1).
I48 22.2 13.7 14
The frequency distribution biodiversity hotspots for
terrestrial vertebrates in Italy (Fig. 4) covered 3.2% of the Results obtained with three different neighborhood levels (8, 24 and
study area and clearly identified the middle elevation areas 48 cells) for the Getis’ Gi* and the Moran’s Ii, three global thresholds
(Glob.thr 5, Glob.thr 10 and Glob.thr 15%) and the frequency dis-
along the two main mountain ranges, the Alps and the tribution approach
Apennines, as already identified by Maiorano et al. (2006,
2007). All the other methods identify the same geograph- last 40 years (Falcucci et al. 2007), and some of them are
ical regions as important (Fig. 5), with generally larger under some level of protection (Maiorano et al. 2008).
areas (Table 2). In this case, the variability in area and As expected, the results obtained with the ‘‘traditional’’
number of hotspots as identified with the ‘‘traditional’’ methods (Getis’ Gi*, Moran Ii, and global threshold) were
methods was high (Table 2). Moran’s Ii with 8 neighbor- highly dependent on the particular setting that was chosen,
hood cells and the 5% global threshold identified as with hotspots appearing and disappearing (Figs. 3 and 5)
hotspots 6.2 and 4.6% of the study area, respectively. depending on the threshold or neighborhood level selected.
A maximum of 22.2% of the study area was selected as a In general, an increase in the neighborhood levels or
hotspot by the Moran’s Ii with 48 neighborhood cells thresholds corresponded to larger areas, while the number
(Fig. 5). All methods identified as important the areas of identified hotspots showed no clear relationship with the
along the lower mountain slopes in the Italian peninsula. particular settings adopted, with the exception of the Getis’
These areas, especially along the northern Apennine range, Gi* hotspots, whose number decreased with increasing
have been abandoned by agriculture and reforested in the neighborhood levels.

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a b
Cumulative frequency
Frequency
Relative species richness
Fig. 4 a Cumulative relative frequency distribution curve of
terrestrial vertebrate species richness showing the 45° slope tangent
to the curve (dotted line) at the highest species richness value.
b Frequency distribution of species richness with the proportion of
Discussion
Given their importance for ecology, management, and
conservation, several approaches have been developed for
the detection of spatially explicit hotspots (see Nelson and
Boots 2008 for an updated review). All these approaches,
however, have different elements of strength and weakness,
and some level of subjectivity (Aldstadt and Getis 2006),
which may have important consequences for their practical
applications (Nelson and Boots 2008).
We developed a new spatially global method which uses
CRFD curves to identify hotspot areas. Cumulative fre-
quency plots have been previously used to describe species
spatial distribution (Marshall and Frank 1994) and to cal-
culate distributional indices mainly oriented to the study of
spatially density-dependent processes (Swain and Wade
1993; Swain and Sinclair 1994), but they have never been
used for the identification of hotspots.
We tested our new spatially global method using two
different examples for the definition of density and bio-
diversity hotspots, and compared the results with those
obtained using well-established hotspot identification
techniques. Although the main hotspots were detected by
all the techniques that we considered, we observed wide
variations in the number and size of the identified hotspots,
with important differences among the different methods
and parameters that were selected. Major discrepancies in
the results were observed in the terrestrial vertebrate spe-
cies richness case study due to the lower autocorrelation
that characterized this dataset (i.e., 48 neighboring cells,
global Moran’s I = 0.551, P \ 0.01) if compared to the
hake recruit density dataset (i.e., 48 neighboring cells,
global Moran’s I = 0.874, P \ 0.01). In fact, a large
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Popul Ecol (2011) 53:351–359
c
Latiitude (°N)
Species richness Longitude (°E)
area (shade area) above the identified species richness threshold
(dotted line). c Spatial distribution of terrestrial vertebrate biodiver-
sity hotspots in the Italian peninsula from the frequency distribution
method
variety of processes drive the distribution of the many
terrestrial vertebrate species in the Italian peninsula (Fal-
cucci et al. 2007; Maiorano et al. 2007), generating high
patchiness in the spatial distribution of species diversity.
As a result, global methods tend to identify more frag-
mented hotspots than local and partially local methods.
We have not been able to identify any objective criterion
that could be used to select one approach over the others,
and also, in the published literature, the selection of a
particular method is driven in many cases by the expected
and most desirable result rather than by scientifically sound
reasoning. Considering the important differences in the
results obtained applying different approaches, the use of a
variety of methods to the identification of hotspots in
ecology is often necessary and worthwhile for a more
complete understanding of the spatial process studied.
Our geometric approach offers an easy and practical
way to identify hotspots by using geometrical properties of
CRFD curves. This method offers the advantage that no
threshold, neighborhood of influence, or other parameter
need to be selected a priori by the observer, in contrast with
most of the commonly used global, partially local, and
local measures. The shape of the CRFD curve varies in
relation to how the intensity of the spatial process changes
across the area, revealing important aspects of the under-
lying spatial phenomenon being considered. Through the
detection of a geometric feature of the cumulative curve,
our method translates this spatial structure into an objective
way to identify those areas characterized by the highest
intensity of the phenomenon.
However, as with all the other global approaches to the
identification of hotspots, the frequency distribution method
is based on the definition of a single threshold to be applied
Popul Ecol (2011) 53:351–359 357

Fig. 5 Spatial distribution of terrestrial vertebrate biodiversity hotspots from Moran’s Ii (with 8, 24 and 48 neighboring cells), Getis’ Gi* (with 8,
24 and 48 neighboring cells), and global predefined thresholds (5, 10 and 15%)

across the whole spatial process, and consequently it does
not specifically search for local structures. This makes the
geometric approach presented here and all the other global
measures attractive for their low computational costs, and
particularly easy to be applied on high-resolution or large-
scale studies. At the same time, global approaches are not
able to account for the local context in which high values
occur. This represents an important limitation for the
application of all spatially global methods, including the
frequency distribution approach. Consequently, the method
can be effectively applied only in the case of spatially
autocorrelated processes, where high values have a low
probability to occur in isolation. Moreover, the spatially
global methods should be employed under the assumption
of homogeneous processes (i.e., hake recruits in our study
area are part of a single fish population). Because this

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condition rarely occurs, large areas characterized by non-
stationarity should be partitioned into smaller regions of
investigation before spatially global thresholds should be
applied without the risk of losing locally relevant hotspots.
The example on terrestrial vertebrates that we provided
is particular in this respect. In fact, species diversity in
the Italian peninsula cannot be assumed to represent an
homogeneous process (Italy hosts at least 6 different bio-
geographic regions; Falcucci et al. 2007), but the same
peninsula is a single management/conservation unit at the
political level, necessarily asking for analyses that do not
consider hotspots that are important only locally.
An interesting future development of our approach
would be represented by the implementation of a rational
framework to include local analyses based on the use of
CRFD curves, or some other related curve (e.g., geosta-
tistical aggregation curves; Petitgas 1998), and their asso-
ciated tangent with 45° slope. In this case, the method that
we propose could be specifically tailored towards a more
detailed definition of the boundaries of hotspots that have
been identified considering also the local spatial context.
Further testing is required using different datasets and
spatial processes; however, we are confident that the fre-
quency distribution approach can be successfully intro-
duced in ecology and conservation studies as an effective
contribute to the current set of methods available for the
identification of hotspots.
Acknowledgments We especially thank Nathan M. Bacheler for his
helpful comments on an earlier draft of the manuscript, and two
anonymous reviewers for a useful discussion.
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