Fehérvári et al.: Modeling habitat selection of the red-footed falcon (Falco vespertinus)- 59 -

MODELING HABITAT SELECTION OF THE RED-FOOTEDFALCON (FALCO VESPERTINUS): A POSSIBLE EXPLANATIONOF RECENT CHANGES IN BREEDING RANGE WITHINHUNGARY
P. FEHÉRVÁRI 1,4* – A. HARNOS 1,2 – D. NEIDERT 3 – SZ. SOLT 4 – P. PALATITZ 41

2

Department of Biomathematics and Informatics, Faculty of Veterinary Science, Szent István
University

Adaptation to climate change” HAS Research Group,H-1078, István u. 2. Budapest, Hungary(phone: +36-1-478-4214; fax: +36-1-478-4217)
3

SZIE, KTI Department of Cartography, Geoinformatics and Remote Sensing H-2103 Páter
Károly út 1. Gödöllő, Hungary(phone: +36-28-522-000/2270; Fax:+36-28-415-383)

4

MME/BirdLife Hungary, Red-footed falcon Conservation Working Group H-1121 Költő utca
21. Budapest, Hungary(phone: +36-1-275-6247; Fax: +36-1-275-6247)*

Corresponding authore-mail: fehervari.peter@mme.hu(Received 21st March 2008; accepted 15th May 2008)

Abstract. Due to a severe population decline and shrinkage of distribution range in the past decades, thered-footed falcon has gained top priority in both worldwide and Hungarian nature conservation. As afacultative colonial breeder, in Hungary, this species predominantly nests in rookeries. The number ofrooks (Corvus frugilegus) has also dramatically fallen recently, but population decline did not affect thelarge scale breeding distribution of this species. In our study we analyzed the presence of red-footed
falcons at a colony in the case of current and historical breeding ranges based on landscape scaled habitat
variables. We used a potential colony home-range size, estimated from observed home-range sizes inorder to determine the scale of influential habitat variables. According to our results, the primary cause of
the observed range shift is the urbanization of rooks in definable regions of Hungary. The ratio of forestsand open water surfaces within the potential home-range had negative, while the ratio of grasslands had apositive effect on the probability of red-footed falcon presence. None of our models predicted red-footed
falcon presence at colonies outside the current breeding range, suggesting that a probable increase in redfooted falcon population numbers will not be accompanied by the expansion of the current breeding
range.
Keywords: red-footed falcon, habitat, modeling, breeding range, rook, spatial GLMM

Introduction
The analysis of habitat use and selection is important for adequate, well foundedplanning of species based conservation management (Pearce et al., 2008, Robles et al.
2007, Noss et al. 1997). Habitat selection and habitat use data are necessary for theprediction of a species’ distribution (Elith et al., 2006, Guisan 2005, Araújo 2007), forthe assessment of risk factors (Xuhezi et al. 2008, Groning et al. 2007), drafting of
habitat management regulations (Garcia et al. 2006, Franco et al. 2004) and for theconservation of a single species or a group of species. Habitat selection of birds of preyis a common research topic in conservation biology (Toschik 2006, Palomino &APPLIED ECOLOGY AND ENVIRONMENTAL RESEARCH 7(1): 59-69.
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Fehérvári et al.: Modeling habitat selection of the red-footed falcon (Falco vespertinus)- 60 -

Carrascal 2007, Lopez-Lopez et al. 2007, Bustamante 1997), since these birds are goodenvironmental bioindicators (Newton 1979, Roberge & Angelstram 2004) and are often
referred to as umbrella or flagship species (Sergio 2006, Ozaki 2006).
The red-footed falcon is an endangered colonial raptor species, with a continuallydiminishing population, classified as near threatened by the IUCN Red List
(http://www.iucnredlist.org/details/144562). The breeding range is in open, typical
steppe type habitats ranging from Eastern Europe to Lake Baikal in Central Asia (delHoyo et al. 1994). This species is a long distance migrant, with presumed wintering
grounds in South-Western Africa: from the Northern parts of the South African
Republic through Namibia, Botswana, Angola, Zimbabwe and Zambia (del Hoyo et al.,
1994, van Zyl pers. comm.).
The territory of Hungary – which is the westernmost edge of the species’ distribution
range (del Hoyo et al., 1994) – is almost negligible compared to its whole distributionarea, but it has considerable importance in the conservation of this small bird of prey
(Bagyura & Palatitz 2004). Hungarian nature conservation has always paid a great dealof attention to the protection of red-footed falcons, so the overall population estimates
for the country are probably the most accurate throughout the world (Bagyura & Palatitz
2004).
The first country-wide survey performed by Keve and Szíjj (1957) in the middle ofthe last century estimated the breeding population to be 2200-2500 pairs, while the sizeof the population in 2006 was estimated at 500-600 pairs (Palatitz et al. 2006). The
methods used in these surveys differed markedly, therefore direct comparison cannot be
made, but it is certainly true that the population had significantly decreased – possibly
by up to 50% – during the past decades (Fig. 1.).

Figure 1. The estimated maximum number of breeding red-footed falcon pairs in the past 60
years. The red and green columns reflect the number of breeding pairs of Figure2.
and Figure 3., respectively.
Source: 1Keve & Szijj 1957, 2Haraszthy 1981, 3Haraszthy 1998, 4Tóth & Marik 1999,
5,6
Bagyura & Palatitz 2004, 7Palatitz et al., 2005, 8Palatitz et al. 2006

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Beyond the overall population decline, the spatial distribution of breeding birds wasfound to be radically different during the 2006 survey (Fig. 2.) compared to 1949 (Fig.
3.). It is obvious that by 2006 red-footed falcons occupied almost exclusively the Great
Plain region, practically deserting Transdanubia and Northern Hungary.

Figure 2. The distribution of red-footed falcons in Hungary according to the 2006 survey. Red
polygons mark municipipaity borders where breeding pairs were recorded in colonies or as
solitary pairs.

Figure 3. Distribution of red-footed falcons in Hungary according to the 1949 survey. Green
polygons mark municipality borders where red-footed falcon breeding was recorded inrookeries.

The conditions for the colonial nesting of red-footed falcons are primarily provided
by rookeries (Horváth 1964). The rook used to be a wide-spread, common species fromthe 1940’s (Vertse 1943) to the 1980’s (Kalotás & Nikodémusz 1981, Kalotás 1984),
but as a consequence of an intensive eradication campaign in the mid 80’s, thepopulation rapidly decreased throughout the country (Kalotás 1987, Solt 2008) fromAPPLIED ECOLOGY AND ENVIRONMENTAL RESEARCH 7(1): 59-69.
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about 260.000 to almost 23.000 pairs in approx. 30 years. In order to halt this trend, therook was declared protected by the Hungarian Nature Conservation Authorities in 2001.

Figure 4. Rook colonies in 2006. Despite the approx. 90% decline in the rook population, thelarge scale distribution of rookeries was not affected.

As the result of the rook population decline, the number of rookeries suitable for thecolonial nesting of red-footed falcons also decreased. In order to compensate for thisloss, artificial nest box colonies are being established by the Life-Nature Red-footed
falcon conservation project personnel (Solt et al. 2005, www.falcoproject.hu). Despite
this large scale decrease in the number of breeding rook pairs, rookeries are stillavailable for red-footed falcon nesting throughout Hungary (Fig. 4.). Red-footed falcon
population decrease therefore can, to some degree, be explained by the crash of the rook
population, but the recent changes in breeding range cannot be directly linked. In thisstudy we analyzed the relationship between landscape scale habitat variables and thespatial distribution of colonies used by red-footed falcons in order to understand how
these variables affect the current distribution pattern.
Materials and methods
As an initial step, we collected the coordinates of all colonies suitable for red-footed
falcon breeding and defined the current breeding range in 2006. In the second stage weassessed the potential home-range of a red-footed falcon colony. This was carried out
using habitat use analysis data deriving from a three-year radio-telemetry study. The
following step was to draw the potential home-ranges around the coordinates of everycolony and to intersect these with a GIS database containing habitat describing
variables. The variables within the potential home-ranges of every colony were later
used for statistical modelling.
Data
The geographical coordinates of colonies derived from two separate databases: theintegrated population monitoring database of the red-footed falcon LIFE program, andthe database of Rare and Colonial Bird species of the Hungarian Ministry of
Enviorment and Water (Solt 2008).
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The analyses were carried out on two spatial scales. To be able to assess thedifferences between the 1949 (Fig. 3.) and 2006 breeding distribution (Fig. 2.) and tounderstand the pattern of colony occupation within the current breeding range, we
spatially defined the “historic” and “current” breeding ranges.
We considered the whole area of Hungary as the „historic” breeding range, because
the 1949 distribution (Fig. 3.) shows that there was at least one colony in every large
region (apart from high altitude closed forests), therefore we did not a priori exclude
any colonies based solely on its location. While the databases hold 198 potentiallysuitable colonies, we had to exclude the ones where the potential home-ranges
protruded Hungary’s border. Therefore 162 colonies: 41 colonies with red-footed falconpresence and 121 without red-footed falcon presence were used in the analysis
The current breeding range was defined based on the municipality borders of Figure 2.
by applying a 500 meter buffer to the outline of the polygons and connecting the outeredges (Fig. 4.). Two municipality borders (on the Northwestern and Northeastern partof the country) were excluded from the current breeding range because they hold only
four solitary pairs altogether. Therefore, the defined current breeding range holds allcolonies occupied by red-footed falcons and over 99% of the solitary pairs.
Defining the potential home-range of a colony
The true home-ranges of individual red-footed falcons were estimated from the dataof the LIFE program’s ongoing habitat preference analysis. We radio-tagged 24 birds inthe 2006 and 2007 breeding seasons with 3.5g “Biotrack TW-4” radio-tags. The birds
were directly followed during their hunting, and the exact location of hunting eventsand other habitat use variables were recorded (see methods in: Franco et al. 2007, Tella
et al.1998).
Based on the individual home-ranges, we estimated the potential home-range of acolony to be a 3000 meter radius circle, which covers at least 95 % of the localization
points of the studied colonies (unpublished data).

Figure 5. Defining the „current” breeding range of red-footed falcons. The red polygons shownon the map mark the municipality borders where red-footed falcon breeding occurred in 2006
according to the monitoring data. The ruby red large polygon shows the area defined as the„current” breeding range in our analyses.

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Habitat variables
The habitat variables were extracted from the CORINE land cover frameworks GIS
database. In Hungary, this database was created by SPOT4 Xi+M type satellite images
shot in 1998 and 1999. The CORINE has 79 land cover categories. The minimum areaof uniform polygons is 4 ha. Although the database is relatively out of date, it is still
usable to analyse certain variables due to its coarse resolution. The 79 variables weresummed and transformed into 12 biologically relevant variables (Table 3.).
The potential home-range size (3000 meter radius circles) was used to obtain thespatial explanatory variables surrounding each colony. We used the size, length andnumber of the habitat categories within the potential home-ranges for the analyses.
Table 1. Spatial variables used to model landscape scale breeding colony occupancy of
red-footed falcons
Variable
Water canals
ForestRiver
Road
Grassland
Small parcel arable land
Large parcel arable land
Farms
Settlements
Railroad
Water surface
Wetlands

Unit
Meter
Ha
Ha
meter
Ha
Ha
Ha
pcs
Ha
meter
Ha
Ha

Statitstics
We used Spatial Generalized Linear Mixed Effects Models (Spatial GLMM) withlogit link and binomial response to analyse the presence/absence of red-footed falcons
in relation to landscape scaled habitat variables on both historic and current breeding
range scales. Model selection was carried out by applying a decision tree (CART
model) to all variables. The variables grouping the observations were used as
explanatory variables in the Spatial GLMMs. Decision tree pruning was carried out by
optimizing the complexity parameter (Faraway 2005). We categorized the variables into
5 ascending categories to aid better understanding of decision tree outputs, while weused the relative size of the same variables in the GLMMs. We fitted a spatial GLMM
on the observations when the explanatory power of the decision tree was sufficient andthe hierachical structure of the grouping variables was biologically interpretable. The
explanatory power of the GLMMs was more or less smaller than that of the decision
trees, possibly due to the spatial autocorrelation of the expalanatory variables. The
CART models were unable to differentiate between artificial and natural colonies(decision tree, Cohen’s κ = 0.23, 95% Confidence Interval:-0.02, 0.47), therefore we did
not distinguish colony types in the analyses.
We used the QGIS sofware (www.qgis.org) to handle and map GIS variables, andthe R 2.8 sofware for data analysis (R Development Core Team 2007). The mostimportant R packages and their role in the analysis are presented in Table 2.

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Table 2. Most important R packages and their role used in the analyses
Package
Adehabitat
aspacespdep
mvpartMASS

Role
Home range estimation
Measuring spatial distance
Measuring spatial autocorrelation
CART models
Spatial GLMM

Authors
Calange
Remmel & Buliung
BivandTherneau & Atkinson
Venables & Ripley

Results„Current” breeding range
The grouping variables of the decision tree applied on the current breeding range
scale were: „Forest”, ”Large parcel arable lands” and ”Grasslands”. The spatial GLMM
fitted with these variables correctly classified 74 % of the observations (Cohen’sκ = 0.48, 95% Confidence Interval: 0.3, 0.67). The “Large parcel arable land” variable
had no siginifcant effect, the “Grassland” variable had marginally significant positiveeffect, while the “Forest” variable had significantly negative effect, according to themodel (Table 3.).
Table 3. The output of the spatial GLMM fitted on the presence/absence of red-footed
falcons at a colony within the „current” breeding range.
Intercept
Grassland
ForestLarge parcel arable lands

estimate
-1.37
0.03
-0.52
0.013

SE
1.53
0.019
0.20
0.019

t-value
-0.89
1.68
-2.58
1.01

p-value
0.3746
0.05990.01640.4951

Although the „Large parcel arable land” had a significant grouping effect in thedecision tree, it turned out to be non-significant in the spatial GLMM. This is probably
due to the strong spatial autocorrelation of this variable. Presumably, decision trees
overestimate the grouping effects of highly autocorrelated variables, hence thecontradiction of the role of this variable in the different statistical analyses.
„Historic” breeding range.
The decision tree applied on the „historic” breeding range scale (i.e. the wholecountry) classified 90% of the observations correctly. The grouping variables were:„Roads”, „Water surface”, „Forest” and „Grassland”. The spatial GLMM fitted withthese variables classified 87% of observations correctly (Table 6.). The classification ofthis model is also significantly deviating from random classification (Cohen’s κ = 0.52,
95% Confidence Interval: 0.51, 0.78). All grouping variables of the decision tree stayedsignificant in the spatial GLMM.

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Table 4. The output of the spatial GLMM fitted on the presence/absence of red-footed
falcons at a colony within the „historic” breeding range.
Intercept
ForestGrassland
Road
Water surface

estimate
-3.33
-1.5
0.042
-0.19
-1.07

SE
1.16
0.33
0.01
0.085
0.4

t-value
-2.87
-4.52
3.53
2.21
-2.63

p-value0.005<0.001<0.001
0.023
0.008

The significant „Road” variable is highly correlated with the ratio of „Settlement”
variable (Spearman’s rank correlation coefficient = 0.75, p<0.0001) within the potential
home range, therefore it can be used as an indicator of human presence. If the„Settlement” variable occupied more than 25% percent of the potential home-range of agiven colony, we classified it as urban colonies. Out of the 76 rookeries outside thecurrent breeding range, 58 are classified as urban colonies. On the other hand, only 3
out of the 86 colonies within the current breeding range are classified as urban colonies.
The „Forest” variable had a negative effect on the probability of red-footed falconpresence at a given colony on both spatial scales, although there is a large difference inthe ratio of forests between the two breeding ranges. The „historic” model does notpredict red-footed falcon presence at any of the colonies outside the current breeding
range (Fig. 6.).

Figure 6. The predictons of the historic breeding range model. According to our model
predictions, there are no colonies suitable for red-footed falcon breeding outside the current
breeding range.

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Discussion
Although the classification of the model fitted on the current breeding range ismoderately accurate (74%), it reveals an interesting pattern in the case of the “Forest”
variable. This variable has significant negative effect on the probability of red-footed
falcon presence in models fitted on both spatial scales, even though the variable range isquite different. The CORINE can only differentiate forest patches larger than 4 ha,therefore the 3-4% of forests in the potential home ranges of colonies within the current
breeding range practically mean a few small patches or one larger patch. The Goshawk(Accipiter gentilis) is a regular, well distributed (Haraszthy 1988) predator of the redfooted falcons (pers. obs., Bagyura & Haraszthy 1994), occupying small to large forest
patches for breeding. Therefore, the observed colony occupying pattern of red-footed
falcons can be explained as a predator avoiding strategy (e.g. Brodie & Formanowicz
1991, Fontaine & Martin 2006).
The positive effect of grasslands was not as significant as expected in the current
breeding range model. The habitat use analyses revealed that this species uses
agricultural lands relatively often for hunting (Fehérvári et. al 2006), suggesting that thebirds can substitute grasslands to a certain degree, similar to lesser kestrels (Tella et. al1998), causing the “Grassland” variable’s lower explanatory power. The significant
difference between colonies outside the current breeding range and the ones currently
occupied by red-footed falcons can be explained by two reasons: 1) the previously
mentioned urbanization of rooks outside the current breeding range, and 2) that thelandscape has been significantly transformed over the past decades. Source: Central
Statistical Institute: http://portal.ksh.hu/pls/ksh/docs/hun/agrar/html/tabl1_3_1.html).
Presumably, landscape modification has been greater outside the current breeding
range, but most probably these two main causes acted synergistically to generate thecurrent red-footed falcon breeding distribution. This presumption may be confirmed by
the prediction map of the “historic” breeding range model, which does not predict redfooted falcon occurrence outside the current breeding range (Fig. 5.). Although ourmodel variables derive from a coarse GIS database, and we only considered the
distribution pattern of one year’s red-footed falcon breeding distribution, our model
predictions may aid the mid-term nature conservation strategy of this near threatened
species. It is quite clear that − in the current situation − without the local redistributionof rookeries (i.e. from urban to natural habitats outside the current breeding range of
red-footed falcons) there is a low chance of red-footed falcon breeding range reexpansion.
Acknowledgements. This work was partly supported by the ”Conservation of the Red-footed falcon in
the Pannon Region” LIFE NATURE program and OTKA T049157 grant and MME/BirdLife Hungary’s
Monitoring Centre. We are grateful for Anna Nagy and two anonymous reviewers who provided helpfulcomments on this manuscript and we thank every volunteer for their indispensable help in data collection.

REFERENCES[1][2]

Araújo M.B., New M. (2007): Ensemble forecasting of species distributions. −Trends in
Ecology & Evolution 22: 42-47
Bagyura, J., Haraszthy, J. (1994): Adatok a héja (Accipiter gentilis) ragadozómadár- ésbagolytáplálékához −Aquila, 101, pp. 89-92

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[3]
[4]
[5]
[6]
[7]
[8]

[9]
[10]
[11]
[12]
[13]
[14]
[15]
[16]
[17]
[18]
[19]
[20]
[21]
[22]
[23]

Bagyura, J., Haraszthy, L., Szitta, T. (1996): Gyakorlati ragadozómadár-védelem. −Magánkiadás. ISBN: 9630462842
Bagyura, J., Palatitz, P. (2004): Fajmegőrzési tervek, Kék vércse (Falco vespertinus). −KvVM TvH.
Brodie, E.D., Formanowicz, D.R. (1991): Predator avoidance and antipredator
mechanisms–distinct pathways to survival. − Ethol. Ecol. Evol. 3: 73–77.
Bustamante, J. (1997): Predictive models for Lesser kestrel Falco naumanni distribution
abundance and extinction in southern spain. − Biological Conservation 80: 153-160.
del Hoyo, J., Elliot, A., Sargatal, J. (eds) (1994): Handbook of the Birds of the World.
Vol. 2. New World Vultures to Guineafowl. − Lynx Edicions, Barcelona.
Elith, J., Graham, C. E., Anderson, R.P., Dudík, M., Ferrier, S., Guisan, A., Hijmans,
R.J., Huettmann, F., Leathwick, J.R., Lehmann, A., Jin Li,. Lohmann, L.G., Loiselle,
B.A., Manion, G., Moritz, C., Nakamura, M., Nakazawa, Y., Overton, J.M.M., Peterson,
T.A., Phillips, S.J., Richardson, K., Scachetti-Pereira, R.,. Schapire, R.E., Soberón, J.,
Williams, S., Wisz, M., Zimmermann, N. (2006): Novel methods improve prediction of
species’ distributions from occurrence data. − Ecography 29 (2): 129–151.
Faraway, J. (2005): Extending the Linear Model with R − CRC Press
Fehérvári, P., Neidert, D., Solt, Sz., Kotymán, L., Szövényi, G., Soltész, Z., Palatitz, P.
(2006): Kék vércse élőhely preferencia vizsgálat – egy tesztév eredményei. − Heliaca: 5159.
Fontaine, J. J., Martin, T. E. (2006): Parent birds assess nest predation risk and adjust
their reproductive strategies. − Ecology Letters 9 (4): 428-434.
Franco, A.M.A, Palmeirim, J.M., Sutherland, W.J. (2007): A method for comparing
effectiveness of research techniques in conservation and applied ecology. − BiologicalConservation (134) 1: 96-105.
Franco, A.M.A, Sutherland, W.J. (2004): Modelling the foraging habitat selection oflesser kestrels: conservation implications of European Agricultural Policies. − BiologicalConservation (120) 1: 63-74.
Garcia, J.T., Morales, M.B., Martinez, J. et al. (2006): Foraging activity and use of spaceby Lesser Kestrel Falco naumanni in relation to agrarian management in central Spain. −Bird Conservation International (16) 1: 83-95.
Groning, J., Krause, S., Hochkirch, A. (2007): Habitat preferences of an endangered
insect species, Cepero's ground-hopper (Tetrix ceperoi). − Ecological Research 22(55):
767-773.
Guisan, A., Thuiller, W. (2005): Predicting species distribution: offering more thansimple habitat models. − Ecology Letters 8 (9): 993-1009.
Haraszthy, L. (1981) Contributions to the quantity conditions and hatching biology of
red-foot falcons hatching in the Hortobagy 1973. − Aquila. 1981; 87: 117-122.
Haraszthy, L. (ed) (1998): Magyarország madarai − Mezőgazda Kiadó, Budapest.
Horváth, L. (1964): A kék vércse (Falco vespertinus L.) é s a kis őrgébics (Lanius minorGm.) élettörténetének összehasonlító vizsgálata II. A fiókák kikelésétől az őszi vonulásig− Vertebrata Hungarica VI. 1-2.
Kalotás, Zs. (1984): A vetési varjú (Corvus frugilegus) állományfelmérése
Magyarországon 1980 tavaszán. − Puszta (2)11: 109-121.
Kalotás, Zs. (1987): A vetési varjú (Corvus frugilegus L.) hazai állományának felmérése
1984 tavaszán. − Állattani közlemények 74 (1-4): 65-76.
Kalotás, Zs., Nikodémusz, E. (1981): Szelektív varjúirtás lehetősége a 3-klór-4metilanilin-hidroklorid anyaggal. 1. Etetési és szabadföldi vizsgálatok a vetési varjún(Corvus frugilegus L.). − Állattani közlemények 68(1-4): 89-96.
Keve, A. Szijj, J. (1957): Distribution, biologie et alimentation du Faucon kobez Falco
vespertinus L. en Hongrie − Alauda 25(1): 1-23.

APPLIED ECOLOGY AND ENVIRONMENTAL RESEARCH 7(1): 59-69.
http://www.ecology.uni-corvinus.hu ● ISSN 1589 1623
 2009, Penkala Bt., Budapest, Hungary

Fehérvári et al.: Modeling habitat selection of the red-footed falcon (Falco vespertinus)- 69 -[24] Lopez-Lopez, P., Garcia-Ripolles, C., Soutullo, A. (2007): Identifying potentially suitablenesting habitat for golden eagles applied to ‘important bird areas’ design. − Animal
Conservation 10 (2): 208–218.
[25] Negro, J.J., Hiraldo, F. (1993): Nest-site selection and breeding success in the lesser
kestrel Falco naumanni. − Bird Study 40 (22): 115-119.
[26] Newton, I. (1979): Population Ecology of Raptors. − Poyser, Berkhamsted, UK[27] Noss, R.F., O'Connell, M.A., Murphy, D. D. (1997): The science of conservation
planning: Habitat conservation under the endangered species act. − Island Press[28] Ozaki, K., Isono, M., Kawahara, T., Iida, S., Kudo, T., Fukuyama, K. (2006): A
Mechanistic approach to evaluation of umbrella species as conservation surrogates. −Conservation Biology 20 (5): 1507–1515.
[29] Palatitz P., Solt Sz., Fehérvári P. (2005): Kékvércse-védelmi Munkacsoport 2005. évi
beszámolója. − Heliaca: 12-17[30] Palatitz P., Solt Sz., Fehérvári P., Neidert D. és Bánfi P. (2006): Kékvércse-védelmi
Munkacsoport 2006. évi beszámolója. − Heliaca: 16-24[31] Palomino, D. Carrascal, L.M. (2007): Habitat associations of a raptor community in a
mosaic landscape of Central Spain under urban development − Landscape and Urban
Planning 83 (4): 268-274.
[32] Pearce, J. L., Kirk, D. A. P., Lane, C. P., Mahr, M. H., Walmsley, J., Casey, D., Muir,
J.E., Hannon, S., Hansen, A., Jones, K. (2008): Prioritizing avian conservation areas for
the Yellowstone to Yukon Region of North America. – Biological Conservation 141(4):908-924.
[33] R Development Core Team (2007): R: A language and environment for statisticalcomputing. − R Foundation for Statistical Computing, Vienna, Austria.
[34] Roberge, J.M., Angelstram, P. (2004): Usefulness of the umbrella species concept as aconservation tool. − Conservation Biology 18 (1): 76–85.
[35] Robles, H., Ciudad, C., Vera, R., Olea, P.P.,. Purroy, F.J., Matthysen, E. (2007):
Sylvopastoral management and conservation of the middle spotted woodpecker at thesouth-western edge of its distribution range. − Forest Ecology and Management. 242 (23): 343-352.
[36] Sergio, F., Newton, I., Marchesi, L., Pedrini, P. (2006): Ecologically justified charisma:preservation of top predators delivers biodiversity conservation. − Journal of Applied
Ecology 43 (6): 1049–1055.
[37] Solt Sz., Pigniczki Cs., Utassy T., Fehérvári P. és Palatitz P. (2005): Kiskunsági kék
vércse kitekintő – avagy egy ígéretes műfészek telep kialakításának tapasztalatai. −Heliaca: 67-71.
[38] Solt, Sz. (2008): Vetési varjú konfliktus kezelési javaslatok tervezet. MME gondozásban.
In Press.
[39] Solt, Sz., Pigniczki, Cs., Utassy, T., Fehérvári, P., Palatitz, P. (2005): Kiskunsági kék
vércse kitekintő – avagy egy ígéretes műfészek telep kialakításának tapasztalatai. −Heliaca: 67-71.
[40] Tella, J.L., Forereo, M.G., Hiraldo, F., Donazar, J.A. (1998): Conflicts between Lesser
Kestrel Conservation and European Agricultural Policies as Identified by Habitat Use
Analyses. − Conservation Biology 12 (3): 593–604[41] Toschik, P.C., Christman, M.C., Rattner, B.A., Ottinger, M.A. (2006): Evaluation ofosprey habitat suitability and interaction with contaminant exposure. − Journal of
Wildlife Management 70 (44): 977-988.
[42] Tóth, I. és Marik, P. (1999): Kék vércse felmérés. – Madártávlat 4: 4-5.
[43] Vertse, A. (1943): A vetési varju elterjedése, táplálkozása és mezőgazdasági jelentősége
Magyarországon. − Aquila (50): 142-248.
[44] Xuezhi, W., Weihua, X., Zhiyun, O., Jianguo, L., Yi, X., Youping, C., Lianjun, Z.,
Junzhong, H. (2008): Application of ecological-niche factor analysis in habitat
assessment of giant pandas −Acta Ecologica Sinica 28 (2): 821-828.
APPLIED ECOLOGY AND ENVIRONMENTAL RESEARCH 7(1): 59-69.
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