Rguibi-Idrissi, H.

Revistes Catalanes amb Accés Obert

254 Spring migration in the western Mediterranean and NW Africa
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255Pied Flycatcher
Range
The Pied Flycatcher’s breeding range covers most of 
C and N Europe eastwards to W Siberia and scattered 
parts of NW Africa, Britain, France and Iberia (Glutz von 
Blotzheim & Baeur, 1991; Lundberg & Alatalo, 1992). 
Its wintering areas lie in western tropical Africa, most-
ly north of the Gulf of Guinea to c. 11ºN, and from 
Guinea and Gambia eastwards to the Central African 
Republic (Moreau, 1972; Urban et al., 1997; Cramp, 
1998). It does not breed in any of the study sites. Al-
though two rather distinct subspecies do breed in Iberia 
and Morocco (iberiae and speculigera, respectively; Sa-
etre et al., 2001), the vast majority of captures reported 
here refer to the nominate race. 
Migratory route
Birds migrating though the area in spring originate from 
a wide area ranging from Britain and C Europe, north 
to Scandinavia and eastwards to the Baltic and W Russia 
(fig. 1). Passage through the study area takes place fol-
lowing a main SW-NE direction, except for birds originat-
ing from Britain, which move due N. These results agree 
with the well-known migratory route of this species 
through Europe and N Africa (Zink, 1985; Cramp, 1998; 
Rguibi-Idrissi, 2002; Spina & Volponi, 2009). Spring pas-
sage takes place distinctly further east than in autumn in 
a text-book example of loop migration (Zink, 1985; New-
ton, 2008). Accordingly, in Iberia most autumn recover-
ies take place in the centre and west of the Peninsula 
(Telleria et al., 1999), while in Morocco the main autumn 
passage occurs along the western Atlantic coast (Rguibi-
Idrissi, 2002; Thévenot et al., 2003). In spring, on the 
contrary, the recovery and capture data presented here 
indicate that this species is very common along the Medi-
terranean Spanish coast and the Balearics (figs. 1, 2), and 
commoner along the Mediterranean coast of Morocco 
and inland than on the Atlantic coast (as also indicated by 
Rguibi-Idrissi, 2002; Thévenot et al., 2003). 
The species seems to stop relative more frequently in 
NW Africa in spring than in autumn, increasingly so to the 
east. In Morocco, most recoveries are from spring (243 
against 105 in autumn; Moroccan Bird Ringing Centre, 
unpubl. data), although observations are frequent in both 
seasons (Thévenot et al., 2003); in Algeria and Tunisia, 
however, the vast majority of recoveries take place in 
spring and the species is markedly more common then 
than in autumn (Dejonghe & Cournet, 1982; Isenmann & 
Moali, 2000; Isenmann et al., 2005). In SW Europe the re-
verse is observed, recoveries being much more common 
in autumn than in spring (Cramp, 1998; Telleria et al., 
1999). The notable passage and abundance of recover-
ies along the entire Mediterranean coast of N Africa indi-
cates that the species crosses the W and C Mediterranean 
along a broad front (cf. also Spina & Volponi, 2009).
Phenology
The first individuals migrate through the area in early 
April, with passage progressively reaching a peak in late 
April and early May, and then decreasing steadily after-
wards (fig. 2). The pattern is very similar in Catalonia 
and the Balearics / Els Columbrets, but with a peak in 
mid-April in N Morocco. Birds on passage can still be 
seen in the area in early June, outside the study period 
(Telleria et al., 1999; Thévenot et al., 2003). Overall, the 
phenological pattern in the C Mediterranean is very 
similar to that shown here, with the median date of pas-
sage on Capri (1 May) being nearly identical (Pettersson 
et al., 1990; Spina et al., 1993). At Gibraltar and N Mo-
rocco passage can start in mid- or late March, although 
not usually until April (Finlayson, 1992; Thévenot et al., 
2003), while in S Morocco it occasionally starts in early 
March, but mostly also occurs from early April onwards 
(Thévenot et al., 2003; Gargallo et al., unpubl.). 
Males pass distinctly earlier than females (differenc-
es in median dates 10 and 9 days in adults and second-
year birds, respectively), but adults only slightly earlier 
than second-year birds (3 and 2 days earlier in males 
and females, respectively). These sex and age-related 
phenological differences are well documented dur-
ing both migration and on arrival at breeding grounds 
(Dornbusch, 1981; Dejonghe & Cornuet, 1982; Cramp, 
1998; Messineo et al., 2001; Rguibi-Idrissi, 2002). 
However, sexual differences in passage may be less ac-
centuated in the C Mediterranean, since medians differ 
only by 4 and 6 days in adults and second-year birds, 
respectively (Petterson et al., 1990). Recoveries show 
that the further north birds are ringed / recovered, the 
later they pass through the study area, indicating that 
northern populations tend to delay their passage.
Biometry and physical condition
Mean values for third primary lengths range from 60.4 
in N Morocco to 61.5 in the wet Balearics and S Mo-
rocco, figures that are very similar to those reported in 
the C Mediterranean (Messineo et al., 2001; table 1). 
Mean values for wing length vary from 78.2 in N Mo-
rocco to 79.7 in the wet Balearics, also similar to those 
obtained in other sites in the C Mediterranean (Rubo-
lini et al., 2004, Waldenström et al. 2004) and C and 
N Europe (Cramp, 1998). Wing length clearly decreases 
over time, which probably reflects the later passage 
of shorter-winged females (Cramp, 1998) (fig. 6). The 
same trend can be seen on the Tyrrhenian islands (Spina 
et al., 1993). Interestingly, birds trapped in the wet Bal-
earics have significantly longer third primaries than in 
the dry Balearics (also when considering only males). 
Fat reserves are significantly lowest in the dry Balear-
ics and, especially, Els Columbrets, and highest in Cata-
lonia, S Morocco and the wet Balearics (table 1, fig. 8). 
256 Spring migration in the western Mediterranean and NW Africa
Physical condition, on the other hand, is distinctly higher 
in N Morocco and the wet Balearics; in the dry Balear-
ics and Catalonia figures are similar and are clearly bet-
ter than in Els Columbrets. Fat tends to increase with 
time in the Balearics and N Morocco and physical con-
dition in the Balearics and Catalonia (figs. 7, 9). These 
increasing trends suggest that later arriving birds (more 
females) migrate in slightly better condition, possibly 
due to being in less of a hurry to migrate faster or to 
the better environmental conditions encountered as the 
season progresses en route and / or at wintering / fat-
tening grounds. Mean body mass varies from 10.6 in 
Els Columbrets to 12.3 in S Morocco, but with no clear 
overall trend (fig. 9). It decreases significantly with time 
in S Morocco, but in the dry Balearics it increases, albeit 
only slightly. Averages are highest in S and N Morocco, 
and lowest in the dry Balearics and, above all, on Els Co-
lumbrets. Birds from the wet Balearics are significantly 
heavier than on the other islands. Body mass on Els Co-
lumbrets and the dry Balearics is similar to that reported 
from the Tyrrhenian islands (10.8, n = 1,172; Spina et 
al., 1993), while the average from Catalonia is very simi-
lar to that obtained at Gibraltar (11.5, n = 28; Finlayson, 
1981) and further north in Switzerland (11.9, n = 42; 
Cramp, 1992). Birds from N Morocco have a similar av-
erage to N Tunisia (mean 12.3, n = 67; Waldenström et 
al., 2004), but those from S Morocco are slightly heavier 
than those reported at the nearby sites of Defilia (11.6, 
n = 60; Ash, 1969) and Merzouga (11.6, n = 101; Gar-
gallo et al., unpubl.). 
As birds trapped in N Morocco are at most only 
slightly heavier than in the south, mass gain in the 
area seems to be globally rather limited. In Catalonia 
birds are only in slightly poorer body condition than 
in N Morocco and show similar mass to those from S 
Iberia and C Europe, suggesting that once in conti-
nental Europe the species migrates in short bouts that 
do not require long stopovers or significant new gains 
in mass. Only birds passing through Els Columbrets 
and the Balearics show markedly poorer body condi-
tion, a sign of the efforts undertaken while crossing 
the sea. In the wet Balearics, however, birds are sig-
nificantly heavier, have larger fat reserves and longer 
wings than on the other islands, where apparently a 
higher proportion of birds in poor body condition are 
attracted to land. The fact that these birds also have 
on average shorter wings suggests that smaller size 
may make them more prone to suffer from unfavour-
able meteorological circumstances (particularly strong 
head winds; cf. Newton, 2008; Saino et al., 2010) 
and thus be more inclined to stop at any available 
site. Birds stopping at wetlands on larger islands may 
also gain mass faster and include a larger propor-
tion of birds that have been at the site for a few days 
(either at the site itself or other surrounding areas), 
which may also contribute to the overall better aver-
age body condition in these places. 
Stopover
The percentage of retraps is low in all areas except N 
Morocco, where totals attain c. 18% of birds (fig. 5, ta-
ble 2). Mean stopover lengths are in the range 2-4 days 
and are not markedly different in any area in particular. 
Birds staying in the dry Balearics are in poorer condition 
when first captured than those not trapped again, indi-
cating that a higher proportion of birds in poor condi-
tion end up staying at these sites. When considering all 
the retraps, bird tend to show negative fuel deposition 
rates (although only significantly so on Els Columbrets) 
and nearly so in the dry Balearics. If one-day retraps 
are excluded, rates become positive at all sites except 
Els Columbrets, although never significantly so. The 
high number of retraps in N Morocco coincides with 
its expected role as a relevant stopover area; however, 
as shown by biometrical data no clear pattern of mass 
gain is observed. 
257Pied Flycatcher
Figure 1. Map of recoveries 
of birds captured in the 
study area during the study 
period (March to May). 
Table 1. Mean (± SD), range and sample size of main biometric parameters according to area.
 n Wing Third primary Body mass Fat score
Catalonia 1,729 79.6 ± 1.8 (72.0-87.0)  61.4 ± 1.6 (54.5-66.5)  11.9 ± 1.1 (8.8-16.4)  2.0 ± 1.1 (0-5) 
Columbrets 468 79.8 ± 2.3 (72.0-85.0)  60.9 ± 1.7 (54.5-68.0)  10.6 ± 1.1 (7.3-15.3)  0.7 ± 0.8 (0-5) 
Balearics (dry) 4,356 78.9 ± 2.0 (72.0-87.0)  60.7 ± 1.8 (54.5-68.0)  11.5 ± 1.2 (6.2-16.5)  1.6 ± 1.1 (0-6) 
Balearics (wet) 100 79.7 ± 1.6 (76.0-83.0)  61.5 ± 1.5 (57.0-65.0)  12.2 ± 1.5 (8.0-16.0)  2.4 ± 1.1 (0-5) 
Chafarinas 9  60.5 ± 1.5 (58.5-62.5)  11.1 ± 1.1 (10.0-13.5)  1.2 ± 1.0 (0-3) 
N Morocco 90 78.2 ± 1.9 (74.0-82.5)  60.4 ± 1.6 (56.0-64.5)  12.2 ± 1.3 (8.5-16.5)  2.0 ± 1.3 (0-5) 
S Morocco 88 79.6 ± 1.9 (76.0-84.0)  61.5 ± 1.5 (58.5-64.5)  12.3 ± 1.1 (10.0-15.1)  2.2 ± 1.0 (1-4) 
Table 2. Variation in fuel deposition rate (g / day) according to area and type of retraps involved (mean ± 95% CI and sample size are given).
 Catalonia Columbrets Balearics (dry) Balearics (wet) Chafarinas N Morocco
All retraps -0.08 ± 0.10 (97) -0.29 ± 0.18 (18) -0.06 ± 0.06 (277)     -0.07 ± 0.33 (16)
Retraps > 1 day 0.05 ± 0.10 (46) -0.03 ± 0.21 (8) 0.04 ± 0.06 (165)     0.14 ± 0.26 (8)
258 Spring migration in the western Mediterranean and NW Africa
                                    March                            April                              May
                       13  14  15  16  17  18  19  20  21  22  23  24  25  26  27  28  29  30
Pentad
                                 Total (median date = 30/4; n = 8,051)                            Adult male (median date = 23/4; n = 595)
                                            2nd-year male (median date = 26/4; n = 2,198)           Adult female (median date = 3/5; n = 866)
                                            2nd-year female (median date = 5/5; n = 2,158)
25
20
15
10
5
0
%
 C
ap
tu
re
s
Figure 2. Relative 
frequency of captures and 
mean daily numbers 
according to site during 
the standard period 
(16 April to 15 May). 
Figure 3. Frequency of 
captures during the  
study period. 
Aiguamolls
Llobregat
Canal Vell
Alfacada
Punta de la Banya
Columbrets
Illa de Colom
Albufera d’Es Grau
Illa de l’Aire
Albufera d’Alcudia
Dragonera
Cabrera
Conillera
Formentera
Illa Grosa
Chafarines
Kerbacha
Larache
Sidi Bou Rhaba
Fguig
Jorf
Mean daily number of captures
 0          1           2          3          4           5          6           7    
0              20             40              60             80             100
Frequency
     Relative frequency                           Mean daily captures Relative frequency                     Mean daily captures
259Pied Flycatcher
Figure 4. Variation in body 
mass and fat score 
according to site during
 the standard period 
(16 April to 15 May).
Figure 5. Variation in body 
mass by trapping status, 
minimum stopover length 
and frequency of retraps 
according to area.
Aiguamolls (n = 611)
Llobregat (n = 102)
Canal Vell (n = 752)
Alfacada (n = 35)
Punta de la Banya (n = 93)
Columbrets (n = 557)
Illa de Colom (n = 109)
Albufera d’Es Grau (n = 92) 
Illa de l’Aire (n = 786)
Albufera d’Alcudia (n = 7)
Dragonera (n = 365)
Cabrera (n = 2,457)
Conillera (n = 395)
Formentera (n = 510)
Illa Grosa
Chafarines (n = 9)
Kerbacha (n = 95)
Larache
Sidi Bou Rhaba (n = 9)
Fguig (n = 10)
Jorf (n = 78)
Fat score (± 95% CI)
0       0.5       1        1.5        2        2.5       3        3.5    
9             10           11            12           13           14
Body mass (±  95% CI)
    Body mass
14
13
12
11
10
9
8
7
6
    Catalonia    Columbrets  Balearics (d)  Balearics (w)   Chafarinas   N Morocco
   6% (1,840)       2.6% (724)      6.1% (4,758)       0% (102)             0% (9)         17.5% (120)
Area
% of retraps (sample size)  
B
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m
as
s 
(m
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 ±
 9
5 
C
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6
5
4
3
2
1
0
St
o
p
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ve
r 
(m
ea
n
 ±
 9
5 
C
I)
Fat score                     Body mass
 Not retrapped                    Retrapped (1st capture) 
  Stopover duration         Retrapped  (last capture)
260 Spring migration in the western Mediterranean and NW Africa
Figure 6. Temporal 
variation of third primary 
length according to area.
Figure 7. Temporal 
variation of physical 
condition according  
to area.
                                        March                                April                            May  
   13    14   15    16    17    18    19    20    21    22    23    24    25    26    27   28    29    30
Pentad  
      Catalonia          Columbrets            Balearic Is (dry)               Balearic Is (wet)
      Chafarinas         N Morocco            S Morocco
 T
h
ir
d
 p
ri
m
ar
y 
le
n
g
th
 (
m
m
) 
 
64
63
62
61
60
59
58
                                        March                          April                              May
   13    14   15    16    17    18    19    20    21    22    23    24    25    26    27   28    29    30
Pentad  
    Catalonia         Columbrets            Balearic Is (dry)               Balearic Is (wet)
    Chafarinas        N Morocco            S Morocco 
Ph
ys
ic
al
 c
o
n
d
it
io
n
0.220
0.215
0.210
0.205
0.200
0.195
0.190
0.185
0.180
0.175
0.170
Catalonia                      Columbrets     Balearic Is. (dry)        Balearic Is. (wet)
Ch farinas                     N Morocco     S Moro co
Catalonia                      Columbrets     Balearic Is. (dry)        Balearic Is. (wet)
Chafarinas                     N Morocco     S Morocco
261Pied Flycatcher
Fa
t 
sc
o
re
  
3.5
3
2.5
2
1.5
1
0.5
0
                                        March                             April                              May
  13    14   15    16    17    18    19    20    21    22    23    24    25    26    27   28    29    30
Pentad  
      Catalonia          Columbrets            Balearic Is (dry)              Balearic Is (wet)
      Chafarinas         N Morocco            S Morocco
Figure 8. Temporal 
variation in body mass 
according to area.
Figure 9. Temporal 
variation in fat score 
according to area.
B
o
d
y 
m
as
s 
(g
r)
13.5
13
12.5
12
12.5
12
11.5
11
10.5
10
                                        March                             April                            May
  13    14   15    16    17    18    19    20    21    22    23    24    25    26    27   28    29    30
Pentad  
    Catalonia         Columbrets           Balearic Is (dry)               Balearic Is (wet)
    Chafarinas        N Morocco           S Morocco 
C talonia               Columbrets  Balearic Is. (dry) . ( et)
Ch farinas              N Moro co  S orocco
                    l  Balearic Is. Balearic Is.
                      S orocco


Pied Flycatcher (Ficedula hypoleuca)

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Revistes Catalanes amb Accés Obert