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Migrating eagles and water barrier  545

             2011). Wind data at 1500 m. above sea level were  Table II. Reverse migration (flock and solitary bird sampling
             obtained  from    the  National   Centers   of    units) reported in the study area.
             Environmental  Prediction/National  Center  for   Species                 Inland zone  Coastal zone
             Atmospheric Research (NCEP/NCAR) reanalysis
             project (NOAA/OAR/ESRL PSD, Boulder, CO,          Accipiter nisus             2            1
                                                               Buteo buteo                 1            0
             USA,    http://www.esrl.noaa.gov/psd/;  National
                                                               Circus aeruginosus         15           13
             Oceanic and Atmospheric Administration/Oceanic    Circus pygargus             1            0
             and Atmospheric Research/Earth System Research    Circaetus gallicus          4           15
             Laboratory, Physical Sciences Division) and down-  Falco tinnunculus          1            0
             loaded using RNCEP package in R software (Kemp    Falco tinnunculus/naumanni  3            0
                                                               Hieraaetus pennatus         2            2
             et al. 2012). Variables in the model were chosen
                                                               Milvus migrans             11            0
             following a backward stepwise approach using an
                                                               Pandion haliaetus           1            0
             information  theoretic  approach  (Burnham  &     Pernis apivorus            85            1
             Anderson 2002), selecting the variables by the AIC
             (Akaike 1973).
                                                               Table III. Results of binary logistic regression analysis; asterisk
                                                               flags significance.
             Results
                                                               Explanatory terms        Wald     df       P
             A total of 12,147 directions of migration were
             recorded, involving 67,235 raptors belonging to 23  Watch site             10.5      1     0.001*
             species; of these only 158 (1.3%) resulted in reverse  Wind speed           0.03     2     0.9
             migration (Tables I, II). The short-toed snake eagle  Wind direction        1.9      3     0.8
                                                               Wind direction*Wind speed  0.01    3     1
             Circaetus gallicus (Gmelin, 1788) was the only species
             showing a significant proportion of reverse migration
             (19 vs. 64, 23%; Table II). As a result, we focused
                                                               observations involved migrating birds. The median
             the analysis on this species. Since this eagle does not
                                                               date of the passage of this species was on 26
             breed in the study area (Panuccio et al. 2015), our
                                                               September, and among aged eagles (N = 40) 33
                                                               (82.5%) were first calendar year (cy) birds. Finally,
             Table I. Numbers of observed directions of migration (flock and  we reported 69 solitary individuals and 14 flocks
             solitary bird sampling units) reported during autumn migration  containing on average 2.7 ± 0.7 (standard error,
             2011, 2012 and 2013 in southern continental Italy.  SE) birds. Among birds and flocks showing reverse
                                                               migration (N = 83), we directly observed the change
             Species                Inland zone   Coastal zone
                                                               in the direction (from expected to reversed) in six
             Accipiter nisus            42            26       cases (two flocks and four solitary birds), and always
             Aquila clanga               1             0       in the coastal zone; all birds of the two flocks
             Aquila pomarina             1             0       behaved in the same way, showing a strong social
             Buteo buteo                139           47
             Buteo rufinus                2             0       attraction. As regards statistical analysis, variables
             Circus pygargus/macrourus  39             9       were  not  auto  correlated  (P  >  0.05).  The
             Circus aeruginosus        3329         1085       ANCOVA run on the most parsimonious BLRA
             Circus cyaneus              2             1       model (Table S1) shows that only the observation
             Circaetus gallicus         47            36       zone significantly affected the behavior of short-toed
             Circus macrourus           19             2
             Circus pygargus            62            17       snake eagles (Table III). In particular along the
             Falco subbuteo/eleonorae   24             5       mountain chain eagles were more likely to migrate
             Falco tinnunculus/naumanni  1075        138       in the expected direction of migration (heading
             Falco eleonorae            13             5
                                                               south) than along the coastal zone. The AUC of
             Falco naumanni             106            4
                                                               this model was 0.96 (P < 0.001), so the accuracy of
             Falco peregrinus           14             9
                                                               the model was good.
             Falco subbuteo             116           31
             Falco tinnunculus          25             5
             Falco vespertinus          14             2
                                                               Discussion
             Hieraaetus pennatus        138           73
             Milvus migrans             427          137
                                                               The low proportion of raptors showing reverse
             Milvus milvus              10             1
                                                               migration was probably caused by the location of
             Neophron percnopterus       4             0
             Pandion haliaetus          99            29       watch points, far from the shoreline, but also because
             Pernis apivorus           4081          656       the Strait of Messina is not an insurmountable water
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