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Origin and migration of woodcock Scolopax rusticola wintering in Spain

Article  in  European Journal of Wildlife Research · June 2011

DOI: 10.1007/s10344-010-0475-9

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Eur J Wildl Res (2011) 57:647–655
DOI 10.1007/s10344-010-0475-9

ORIGINAL PAPER

Origin and migration of woodcock Scolopax rusticola

wintering in Spain
Jose Luis Guzmán & Yves Ferrand & Beatriz Arroyo

Received: 11 November 2009 / Revised: 18 October 2010 / Accepted: 22 November 2010 / Published online: 29 December 2010
# Springer-Verlag 2010

Abstract Woodcock Scolopax rusticola is an important through hunting). We discuss the management implications
game species in Northern Spain, where it is mainly a of these results.
wintering species. Knowledge about the migration and
origin of the woodcock wintering in Spain is relatively Keywords Management . Migration . Circum-Baltic
sparse, existing to date only qualitative analyses dating Region . Ringing . Spain
more than a decade. From the analyses of ringing
recoveries of woodcock wintering in Spain, we evaluate
the relative importance of various countries or regions as Introduction
sources of the woodcock wintering in Spain, an estimate of
their migratory route. Our analyses show the Circum-Baltic Migrant game species are usually subject to hunting
Region to be the most important breeding area of the pressure throughout its migratory route, which may make
woodcock wintering in Spain. Within that area, both their management a challenge. Game management is
Sweden and Western Russia appear to be particularly usually regulated at a local or at most national level, but
important. Analyses of the ringing locations of woodcock in the case of migratory species, efficient management is
ringed during migration and recovered in Spain in winter only obtained if there is coordination among the countries
suggest that woodcock wintering in Spain migrate primarily or regions through which the species passes along the
through the South of the Baltic Sea, Germany and France. annual cycle. It is thus important to know the connectivity
The proportion of woodcock ringed in different French between breeding and wintering areas, as well as migratory
regions during the postnuptial migration months (October routes, to determine which countries should have stronger
and November) that was subsequently recovered in Spain coherence in the management of these species.
(mainly through hunting) declined with the proportion of Woodcock Scolopax rusticola is a forest wader with a
those birds that was recovered in France (also mainly wide Palaearctic distribution, present throughout temperate
areas in Asia and Europe, from the British Isles to
Kamtchatka, and from the Arctic Polar Circle to the North
Communicated by C. Gortázar of Africa and the Middle East (Cramp and Simmons 1983).
J. L. Guzmán (*) : B. Arroyo The European breeding population spreads from Fennoscandia
Instituto de Investigación en Recursos Cinegéticos, to the Mediterranean Region (in mountains) and from Western
IREC (CSIC-UCLM-JCCM), Europe to Eastern Russia (Hagemeijer and Blair 1997). It is a
Ronda de Toledo s/n,
pre-Saharan migrant with a leapfrog migration (Alerstam and
Ciudad Real 13005, Spain
e-mail: Joseluis.guzman@uclm.es Hendenstrom 1998): Populations from Russia and North
Europe are mainly migrant, spending the winter from Atlantic
Y. Ferrand to Mediterranean regions and North Africa, while the
Direction des études et de la recherche,
Southern and Western populations are mainly sedentary, with
Office National de la Chasse et de la Faune Sauvage (ONCFS),
5 rue de Saint Thibaud, short-distance migrations (Henderson et al. 1993). Through
St Benoit 78910 Auffargis, France ring recoveries, it has been shown that populations wintering
648 Eur J Wildl Res (2011) 57:647–655

in the UK breed mainly in the Circum-Baltic Region, Norway Latvia, Estonia, and Western Russia (regions of Pskov and
and Denmark (Hoodless and Coulson 1994), whereas those St. Petersburg, Gossmann et al. 1998); Central Russia
wintering France breed mainly in Central Russia (Bauthian et (regions of Arkangel’sk, Vólogda, Tver, Vladimir, Moscow,
al. 2007, Gossmann et al. 1998). Smolensk; Gossmann et al. 1998); Central Europe, with
Knowledge about the migration and origin of the Hungary, Czech Republic, Switzerland, and Germany;
woodcock wintering in Spain is relatively sparse, existing Atlantic Coast (Netherlands, Belgium, France) and British
to date only qualitative analyses dating more than a decade, Isles (UK and Republic of Ireland).
which pointed to the Scandinavian countries as their main We calculated the percentage of woodcock recovered
breeding quarter (Onrubia et al. 1994). This study presents from each country/region out of total woodcock ringed
a comprehensive analysis of ringing recoveries made in there. For all countries except France, we did not have
Spain in winter, in order to obtain a quantitative estimate of information about the number of birds ringed in each
the importance of various countries or regions as sources period, so we used total number of birds ringed as an
of the woodcock wintering in Spain, as well as an estimate estimate of the effort of ringing during the breeding period.
of their migratory route, thus completing the pattern of This is probably correct for most of the northern countries,
migration of woodcock for Western Europe. We discuss our where ringing does not occur in winter. In the case of
results in the context of management of this species. France (where most ringing occurs during migration and
winter), we only considered birds ringed during the
breeding period. We tested whether the proportion of
Methods woodcock recovered in Spain varied among regions with
a Chi-square test. We used the standardized residuals (SR)
Data sources to look for the significance of each cell. SR higher or lower
than 1.95 indicates a significant deviation of the observed
This study is based on the analyses of woodcock ringed in from the expected values (Zar 1989). Subsequently, we
Europe and subsequently recovered in Spain between calculated an index of the importance of each region of
October and March. These data were obtained from the country as a source of woodcock wintering in Spain,
Central Bureau of Ringing in Europe (Euring), the Centro multiplying the percentage of recoveries by the breeding
de Migración de Aves (CMA, Spain), and the Sociedad de population estimate in each area and calculating the percent
Ciencias Aranzadi (Spain). Most of the recoveries (91.1%) value of the resulting figure.
occurred through hunting. Ringing effort (number of To analyze the variability in the winter distribution
woodcock ringed) of France and Russia was obtained within Spain of woodcock coming from different breeding
through the Office National de la Chasse et de la Faune areas, according to the leapfrog migration described for the
Sauvage (ONCFS). For other countries, it was obtained species (Alerstam and Hendenstrom 1998), we classed each
through the respective ringing centers. Ringing data ringing location during the breeding period in two groups:
correspond to the period 1911–2007 and recoveries to the “short-distance” for locations within the countries closer to
period 1911–2008. Spain (less than 2,000 km from Central Spain: Central
Both ringing and recovery data were grouped into three Europe countries, France and British Isles) and “long-
periods (breeding, migration, or wintering) according to date. distance” (more than 2,000 km from Central Spain:
We defined breeding period as April to September and Circum-Baltic countries and Russia). We tested with a
migration period as February–March and October–November general linear model whether latitude or longitude of the
(Lucio and Sáenz de Buruaga M 2000; Saari 2006). winter location varied according to the region of origin as
We analyzed a total of 244 ringing recoveries, of which described above.
57 corresponded to birds ringed in the breeding period
(Table 1) and 187 to birds ringed during migration (Fig. 1). Migratory route

Origin of the wintering population For the analysis of the migratory route, we considered
ringing locations of birds recovered in Spain in winter but
To determine the main breeding regions or countries of ringed during the migration period (n=187). Using the
woodcock wintering in Spain, we considered recoveries of geographical coordinates of all of those locations, we
birds ringed during the breeding period (n=57). Initially, we evaluated (through a regression analyses) the equation that
analyzed data at the country level (Table 1). Subsequently, best defined the dispersion of those locations using the
for statistical analyses (and given the low sample sizes), we program Tablecurve 2D v5.01. Of all possible curves
grouped countries according to geographic location as describing the dispersion, we chose the one with highest
follows: Circum-Baltic Region, with Sweden, Finland, percentage of explained deviance.
Eur J Wildl Res (2011) 57:647–655 649

Table 1 Countries where there has been woodcock ringed during the breeding season and subsequent recoveries in Spain during winter

Country Number ringed Number recovered Percent recoveries Breeding populationa Breeding estimateb Index

Denmark 1,968 0 0.0 2,000–3,000 2,500 0.0

Norway ? 2 ? 30,000–50,000 40,000 0.4c
Finland 3,236 9 0.27 100,000–150,000 125,000 3.3
Sweden 1,783 17 0.95 250,000–1,000,000 625,000 58.9
Latvia 388 1 0.25 20,000–30,000 25,000 0.6
Estonia 360 1 0.27 20,000–50,000 35,000 0.9
W Russia 1,986 9 0.45 1,200,000–5,000,000d 465,000e 20.8
C Russia 1,112 3 0.26 1,200,000–5,000,000d 465,000e 12.0
Switzerland 282 0 0.0 1,130–1,630 1,380 0.0
Czech Republic ? 2 ? 2,000–4,000 3,000 0.1c
Hungary 61 1 1.63 10–20 15 0.0
Germany ? 0 0.0 12,000–24,000 18,000 0.0
Netherlands ? 1 ? 2,000–3,000 2,500 0.1c
Belgium 811 4 0.49 1,600–2,800 2,200 0.1
France 153 2 1.31 10,000–30,000 20,000 2.6
British Isles 9,862 5 0.05 7,900–23,700 15,800 0.1

Index represents the estimated proportion of woodcock wintering in Spain coming from each country or region
a
BirdLife International (2004)
b
Median of the estimate in the previous column
c
Calculated with estimates of 0.1% recoveries for Norway, 0.4% recoveries for Czech Republic, and 0.6 for the Netherlands (these figures represent the
average for the countries in each group, except for Norway, where we added a minimum of 0.1 as there exist some recoveries)
d
For the whole of Russia
e
Estimated as 15% of the median for the whole of Russia

Fig. 1 Ringing location of wood-

cock recovered in Spain during
winter, according to ringing time
(breeding period: filled circles;
migration: open circles)
650 Eur J Wildl Res (2011) 57:647–655

A more detailed analysis of spatial and temporal Atlantic Coast were recovered in Spain more frequently than
migration variation through France may be of interest, as expected, whereas the opposite was observed for woodcock
it has implications for management. For example, ringing coming from the British Isles (Table 2). Taking into account
data may give indications about timing of spring migration breeding population size, it would appear that more than
through France, which are relevant for regulating closure of 80% of the woodcock wintering in Spain would breed in the
hunting seasons there, and determining whether certain Circum-Baltic Region (including Western Russia) (Table 1).
French regions are particularly important as a migratory Figure 3 suggests a different distribution of birds of
flyway for woodcock wintering in Spain is also useful as different origin within Spain, with the most western and
hunting pressure varies strongly among regions there. To southern recoveries corresponding to long-distance migrants
analyze the spatial and temporal variation of movements and short distance migrants being restricted to northern and
within France, we analyzed data of birds ringed in France eastern areas. However, neither longitude nor latitude of the
during migration or wintering months, and subsequently winter location varied significantly between both groups
recovered in Spain, by month and region of ringing. (F1,55 =0.10, P=0.7 for latitude; F1,55 =0.47, P=0.5 for
Regions were defined as in Bauthian et al. (2007) longitude).
(Fig. 2). We used Chi-square tests, and SR to look for the
significance of each cell. Migratory route

General migratory route

Results
Figure 4a shows the modeled curve from the distribution of
Origin of the wintering population locations of woodcock ringed during migration and
subsequently recovered in Spain in winter (n=187). This
Overall, the proportion of woodcock ringed during the curve explained 58% of the variance and suggests that
breeding period and recovered in Spain during winter varied woodcock wintering in Spain probably pass primarily close
significantly with ringing region (X42 =53.78, P<0.001): to the South of the Baltic Sea and subsequently through
woodcock coming from the Circum-Baltic Region and the Germany and France.

Fig. 2 Regions in France used

for analyses (modified from
Bauthian et al. 2007)
Eur J Wildl Res (2011) 57:647–655 651

Table 2 Proportion of woodcock ringed in various regions and lower than expected for the Western Region (Table 4). There
subsequently recovered in Spain during winter
was a negative correlation between the percentage of
Region Number Number Percent SR woodcock ringed in each French region during the postnuptial
ringed recovered recoveries migration months that is subsequently recovered in Spain and
the percentage of those woodcock subsequently recovered in
Circum-Baltic 5,767 37 0.64 4.87
France (Fig. 5, r=−0.906, P<0.0001).
Central Russia 1,112 3 0.26 −0.15
Central 343 1 0.29 −0.01
Europea
Atlantic Coastb 964 6 0.62 3.17 Discussion
British Isles 9,862 5 0.05 −4.46
Origin and distribution of woodcock wintering in Spain
SR standardized residuals of the Chi-square test
a
Data from Germany or Czech Republic not included since no ringing Our analyses indicate that the Circum-Baltic Region is the
effort was available most important breeding area of woodcock wintering in
b
Data from Netherlands not included since no ringing effort was available Spain. Within that area, both Sweden and Western Russia
appear to be particularly important (although it is important
to remark that maybe the difference between Central and
Temporal and spatial variation through France Western Russia is not as clear as we present here: as the
ringing effort in this area takes place in September, when
Not surprisingly, the probability of recovering in Spain postnuptial migration may have started, some of the birds
woodcock ringed in France varied significantly according ringed in Western Russia may have originated further east,
to ringing month (X52 =33.03, P<0.001), being significantly e.g., in Central Russia). Woodcock breeding along the
higher than expected for October and November and Atlantic Coast would also have a relatively high probability
significantly lower than expected for January and February of wintering in Spain, but given the lower importance of the
(Table 3). breeding populations in those countries, the total contribu-
The probability of recovering in Spain woodcock ringed in tion to the wintering population in Spain would be lower.
France during the postnuptial migration months (October and Finally, birds breeding in the British Isles and Central
November) also varied significantly according to ringing Europe would only occasionally winter in Spain.
region (X52 =19.4, P<0.005): Recoveries in Spain are higher These data fit among those described for the British Isles
than expected for birds ringed in the NE and CN regions and (Hoodless and Coulson 1994), which indicate a strong
Fig. 3 Recovery location of
woodcock coming from
short-distance countries
(filled circles) or long-distance
countries (white circles)
652 Eur J Wildl Res (2011) 57:647–655

Fig. 4 a Modeled migratory

route of the woodcock winter-
ing in Spain. Filled circles, best
fit curve; open circles, 95%
confidence limit; open triangles,
95% prediction limits.
b Best-quality locations of two
woodcock tagged with satellite
transmitters by the Spanish
Woodcock Hunting Club
(CCB). Data obtained from
March to May 2009 and kindly
made available to us by CCB

preponderance of the Circum-Baltic and Scandinavian On the other hand, we did not find any firm evidence
regions as origin for the woodcock wintering in that area, that woodcock coming from different breeding areas had
and those described for France (Gossmann et al. 1998; different wintering distribution within Spain, although lack
Bauthian et al. 2007), where ringing data showed a stronger of statistical differences may be due to the small sample
importance of the centre of Russia as origin of the French sizes, so this issue merits further attention.
wintering woodcock. Hence, our analyses complete the
general migratory pattern of woodcock in Western Europe, Migratory routes and timing
suggesting a strong longitudinal gradient among breeding
and wintering areas, with birds breeding further east Analyses of the ringing locations of woodcock ringed
wintering also further east. during migration and recovered in Spain in winter suggest
Eur J Wildl Res (2011) 57:647–655 653

Table 3 Proportion of woodcock ringed in France and subsequently 0,6

% woodcock recovered in
recovered in Spain according to ringing month 0,5

Ringing month Number ringed Number Percent SR 0,4

Spain
recovered recoveries 0,3

October 1,715 9 0.52 2.72 0,2

November 16,144 51 0.32 2.67 0,1

December 14,796 30 0.20 −0.39 0
January 10,191 6 0.06 −3.44 0 5 10 15 20 25 30
February 6,763 7 0.10 −2.02 % woodcock recovered in France
March 5,003 16 0.32 1.54
Fig. 5 Relationship between the percentage of woodcock ringed in
SR standardized residuals of the Chi-square test each French region during the postnuptial migration months (October–
November) that are recovered subsequently (mainly through hunting)
in either Spain or France
that woodcock wintering in Spain migrate through a route
closer to the Baltic Sea than to Central Europe, in
accordance with the above-mentioned results. These results
also coincide with those obtained from various woodcock recoveries in Spain belonged to birds ringed in France in
tagged with satellite transmitters since 2007 (see Fig. 4b), migratory months. According to our results, the probability
arising from a study by the Spanish Woodcock Hunting of recovering woodcock ringed during migration in western
Club (CCB) (http://rtvs.ccbp.org/). regions is lower than elsewhere, suggesting that there may
A previous study determined the existence of two be a lower migratory flux through those regions (as also
migratory routes for woodcock wintering in France, one suggested from Fig. 4). Recoveries from north-central and
going through Fennoscandia and another one through north-eastern regions were more common than expected by
Eastern Europe (Bauthian et al. 2007). This study was chance, which is also in accordance with the general
based on birds ringed in winter in France and subsequently migratory flux. In contrast, it was surprising that south-
recovered (mainly through hunting) along the migratory western or central-southern regions were not significant.
route. Our study, in contrast, in based on the ringing Alternatively, this pattern may be related to the spatial
location of birds recovered in Spain. This implies that the variation in hunting effort within France. The negative
importance of certain countries in the migratory route may correlation found between the probability of recovery in
be hidden if they do not have woodcock ringing programs, France and Spain of woodcock ringed during the migrating
as is the case in many Eastern Europe countries. It is months in different French regions supports this hypothesis. It
therefore plausible that woodcock arrive to Spain also is thus possible that hunting pressure suffered by migrating
through two migratory routes, including another eastern woodcock in France affects the populations wintering in
one, although this will need to be confirmed in further Spain. However, it is important to interpret these results with
studies. caution, because they are just correlations, not causal relation-
France is a compulsory corridor for the woodcock ships. It would be important to do more detailed studies to
wintering in Spain, and more than 40% of all ringing evaluate the relative importance of the heterogeneous passage
through France and the influence of hunting pressure on
wintering populations in Spain.
Table 4 Percentage of woodcock ringed in France in October–
November and subsequently recovered in Spain according to ringing
The analysis of temporal variation in probability of
region (see Fig. 2) recovery indicates that the peak of post-breeding migration
through France occurs in October and November and that
Ringing Number ringed in Number Percent SR pre-breeding migration through France does not occur
region October–November recovered recoveries
before March. These results agree with the average
W 4,541 2 0.04 −2.75 departure dates (October 10) and arrival dates (9 May)
SW 2,969 6 0.20 −0.49 observed in Southeastern Finland (Saari 2006), as well as
NE 3,856 17 0.44 2.43 with migration data suggested for Spain based on ICAs
SE 3,562 11 0.31 0.75 (Lucio and Sáenz de Buruaga M 2000). Similarly, these
N 5,179 12 0.23 −0.22 data are coherent with observed departure dates (to
CN 1,303 7 0.54 2.11 throughout March) from Spain toward breeding grounds
CS 2,906 5 0.17 0.81 obtained from satellite transmitters and for radio-tracked
woodcock in Northern Spain (Guzmán and Arroyo,
SR standardized residuals of the Chi-square test unpublished data).
654 Eur J Wildl Res (2011) 57:647–655

Management implications be having an impact on the Spanish wintering population.

Future studies should address the effect of different
As the main source of woodcock wintering in Spain management guidelines on populations.
appears to be the Circum-Baltic Region (particularly
Sweden and Western Russia), it would be particularly Acknowledgments This study has been funded by the Comité
Interautonómico de Caza y Pesca Continental in Spain and has
important to establish connections between those countries,
been carried out within the Framework of the Interregional Study
in order to include productivity in those regions in the on Woodcock Scolopax rusticola in Spain (project number
hunting management in Spain. For example, hunting bags 200430E471). We thank the staff in ringing offices that provided
could be cupped up in relation to the productivity in those woodcock ringing data: Chris du Feu (EURING Data Bank), Óscar
Frías (Oficina de Especies Migratorias, Dirección General para la
regions. In a recent study, spring temperature in Norway Biodiversidad, Ministerio de Medio Ambiente), Ainara Azkona
appeared to be a good predictor of woodcock hunted in that (Oficina de Anillamiento de Aranzadi), Kaarel Kaisel (Estonia),
country in early autumn (Selas 2006), which would point to Jacquie Clark (British Trust for Ornithology), Juris Kazubiernis
spring temperature as a potential predictor of woodcock (Letonia), Jari Valkama (Finlandia), Dr Thord Fransson (Bird
Ringing Centre, Swedish Museum of Natural History), and Didier
productivity. If verified, this could be an interesting
Vangeluwe (Institut Royal des Sciences Naturelles de Belgique,
management tool, allowing the determination of various Centre Belge de Baguage). Additionally, we are grateful to all the
scenarios (“good” or “poor” reproduction years) and thus Spanish ringers that contributed to the project: David Lizaso, Miguel
various degrees of hunting bag limits. San Martín, Alberto Artácoz, Felipe Oiarzabal, Iñaki Balda, Peio
Oiarzabal, Juan Jesús Picabea, Nestor Domeño, Iñaki Ripa, Angel
Looking at the countries that form part of the
María Sarratea, Eduardo Sarratea, Javier Oiaregui, Ion Aldunzin,
migratory route of woodcock wintering in Spain (accord- Fernando Belzunce, Rafael López de Aberasturi, Jesús Gutierrez,
ing to results in this study) and the duration of the Melchor Alzugara, Agustín Mendiburu, Mikel Olano, Esteban
hunting season in those countries (Lutz and Jensen 2006), Iriarte, Alejandro Onrubia, Ibón Telletxea, Javier Monge, Pedro
Zuleta, Marco Escudero, José Luis Rivas, Javier Lucientes, Carlos
woodcock wintering in Spain are apparently hunted through- Pérez, Javier Ferreres, Rubén Ibáñez, Marco Escudero, Marc Palles,
out the annual cycle, except in June, July, and March. For Celestino Suarez, Miguel González-Vélez, Justo José Robles, and
example, in countries like Russia, Lithuania, and Latvia, Pablo González. Enrique Castién provided useful comments on the
hunting is practiced in April and May (Ferrand and manuscript.
Gossmann 2001; Blokhin and Fokin 2005), which would
include part of the pre-breeding migration and the breeding
period, when hunting is prohibited in European Union
according to Annex II of the Birds Directive (European References
Commission 2004).
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sible about 25% of all woodcock hunted in the European migration theory. J Avian Biol 29:343–369
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unknown. In any case, France is a compulsory corridor for trends and conservation status. BirdLife conservation series 12.
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between both countries seems critical for the long-term Cramp S, Simmons KEL (1983) Handbook of the birds of Europe, the
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