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V104 N01 P002-015 A
V104 N01 P002-015 A
migration cycles
of an adult Hobby
tracked by satellite
Bernd-Ulrich Meyburg, Paul W. Howey,
Christiane Meyburg and Klaus Dietrich Fiuczynski
Abstract A prototype of the smallest satellite transmitter produced so far,
weighing just 5 g, was fitted to an adult Hobby Falco subbuteo in Germany in August
2008. Two autumn and two spring migrations were recorded successfully from the
unit carried by this particular Hobby. All four migration routes were to the west
of a direct line between breeding site and wintering area. The migration route in
spring 2010 was up to 2,150 km farther west than that in autumn 2009, effectively
forming a ‘migration loop’. This was much less pronounced in 2008/09. The direct
distance from the breeding site to the southernmost point reached in Zimbabwe
in 2008/09 was 10,065 km. The fastest flight speeds on migration were recorded in
spring 2010 in Mali and Morocco, when 1,243 km was covered in two days. Also in
spring 2010, during migration from Morocco to southern France (1,032 km in two
days), the falcon migrated at night, when a fix was made over the Mediterranean in
the vicinity of Gibraltar. Migration across the Sahara took 4–4.5 days on each
migration. The West African equatorial rainforest appears to be a significant
ecological barrier, and the significance of this is discussed. Migration was noticeably
rapid there with distances of up to 580 km flown per day – also partly at night.
During both wintering periods the falcon spent the majority of its time in the
Angolan Miombo woodlands, but in winter it also travelled large distances: from
16th October 2008 to 7th April 2009, the bird covered a total distance of at least
9,025 km between identified night roosts. In 2009, the bird spent half the year in
the wintering area, a third on the breeding grounds and the remaining 18% of the
time on migration; i.e. 65% of its time in Africa and 35% in Europe.
he Hobby Falco subbuteo breeds across the species’ main wintering area is believed to
This paper is dedicated to Robin Chancellor, who contributed so much to raptor research and conservation by
editing the Proceedings of all Birds of Prey World Conferences, and who died in October 2010 (see pp. 42–43).
lands from southern areas north across the These data were decoded and processed using
central plateau to Bié. software produced by Microwave Telemetry,
Very little is known about the Hobby’s Inc. (USA) and displayed via Google Earth
migration. Only a small number of birds are Pro, ArcView GIS 3.2b (ESRI GIS and
observed at the well-known bottlenecks such Mapping Software, USA), and Animal Move-
as Gibraltar. A total of 5,720 Hobbies were ments Extension to ArcView GIS. To display
ringed in ten European countries between migration routes, only location classes (LCs)
1909 and 1998 (including over 1,000 in 3, 2, 1 and 0 were used. LCs of inferior and
Berlin and Brandenburg, Germany), of which poor quality (LC A, B and Z) were mostly
203 (3.5%) were subsequently recovered, excluded. The deviations indicated by Argos
although none south of the Sahara Desert for class LC 3 (best quality) are 150 m, LC 2
(Chapman 2002). Since then, we know of up to 350 m and LC 1 up to 1 km. Unless
only two ring-recoveries from Africa south of stated otherwise, transmitter data are given in
the Sahara. At present, mapping the win- co-ordinated universal time. Only where
tering areas of long-distance migrants is important data points were missing were LCs
based mainly on ring-recoveries, satellite of lesser quality used in our results.
telemetry, stable-isotope analysis of feathers We could not detect any effect of the
or other tissues and through the use of transmitter on the Hobby’s behaviour. The
genetic markers. Satellite tracking using the transmitter was programmed to operate on a
Argos system (Argos 1996) is now an duty cycle of 10 hours on/48 hours off. Eco-
accepted technique. logical digital Google Earth maps were used
to evaluate the data. They are based on orig-
Study site and methods inal publications such as Olson et al. (2001)
The Hobby has been studied in Berlin and its and DeFries et al. (2000).
environs for decades (Fiuczynski 1978, 1987, Where possible, the falcon’s night roosts
2007; Fiuczynski & Nethersole-Thompson were established in order to record habitat
1980). A nest-site in a disused Common use and changes in location in the wintering
Raven Corvus corax nest in pine trees north- area. As a rule, because of the transmitter’s
west of Berlin (52°50’N/13°0’E) was in a ter- programming, it was possible to localise only
ritory occupied each year between 2005 and every second roost (but most of them
2009, although young fledged from the site through several good fixes: LC 1–3). Other
only in 2005 and 2008. On 9th August 2008, fixes made in daylight hours were disre-
an adult female (weighing 265 g) was caught garded. The distances between the individual
at this nest-site, using an adult White-tailed night roosts were measured and the respec-
Eagle Haliaeetus albicilla as a decoy. The tive habitat determined by overlaying with
Hobby (which had raised two offspring) was digital maps.
fitted with a new type of satellite transmitter The home ranges were determined with
(Platform Transmitter Terminal, or PTT), the help of fixed kernel techniques (Worton
which was at that time still in a development 1989) and the ‘Minimum Convex Polygon’
phase 1. The device was solar-powered and (MCP) method (Kenward 2001; Selkirk &
weighed 5 g (1.9% of the Hobby’s body Bishop 2002; Laver & Kelly 2008). The evalu-
mass); it was attached as a ‘backpack’ using a ation of the fixes was made using the home-
harness with neck and body loops. The range extension for ArcView from Rodgers &
overall weight of the harness was approxi- Carr (1998) as well as the home-range tools
mately 1 g. for ArcGis from Rodgers et al. (2007). Among
Satellite data were received through the other features, this computer program
Argos satellite-based positioning system. permits calculation of the MCP. The Floating
Amean method (‘proximity to recalculated month), the lowest number being recorded in
mean of selected fixes’) was used to calculate the Mediterranean region, which is explained
the 95% and 80% MCPs. By this method, the by interference in this area (Gros & Malardé
arithmetical mean of all fixes is calculated, 2006). The percentage of high-quality fixes in
the point farthest away determined and Europe (22%) was much lower than that in
deleted, and the arithmetical mean of all fixes Africa (72%). Signal transmission lasted for
recalculated. This procedure is repeated until 21.5 months.
only 95% or 80% of the fixes remain. Winter Altogether, two spring and two autumn
home ranges are calculated in surface area migrations, as well as two wintering periods,
and presented here over a background of the were recorded (fig. 1). In 2009 the bird spent
global tree cover derived from remote 49.3% of the year in the wintering area, 7.2%
sensing. Climate data from a large number of on spring migration, 32.3% in the breeding
meteorological stations along the migration area and 11.2% on autumn migration (fig. 2).
routes was obtained from the internet.
Behaviour at the breeding site
Results before departure on migration
The satellite transmitter fitted to the Hobby, After being fitted with the transmitter, the
the smallest and lightest unit available at the female behaved completely normally; it kept
time, delivered astoundingly high numbers of to its favoured lookout tree and remained
good Argos Doppler fixes. The complete close to the young birds. On 28th August
dataset comprised over 2,000 positions, of 2008 the bird was located in the vicinity of its
which 49% were high-quality locations (LC nest for the last time and on 30th August a fix
1–3). There were fewer fixes in Europe (on was made in southern Germany, near
average 65 per month) than in Africa (80 per Munich. The male parent remained in the
breeding area and fed the young
until 18th September. On 22nd
September the male and the
young had also departed on
migration.
Behaviour on migration
All migration routes were clearly
west of a direct line between the
breeding site and the wintering
area. The fixes recorded farthest
from this line on migration (and
the respective distances from it)
were as follows: 10°30’N 4°40’E
during the 2008 autumn migra-
tion in Nigeria (1,100 km);
14°20’N 10°5’E during the 2009
autumn migration, also in Nigeria
(510 km); 29°30’N 1°33’E during
the 2009 spring migration in
Bernd-Ulrich Meyburg
border between Nigeria and Cameroon (7°N forest region and 395 km over the Sahara. At
12°E), the four routes traced were fairly wide the beginning of May, after some 35 days of
apart but were quite similar between 7°N and migration, the bird reached its breeding site
the wintering area (fig. 1). In Africa, the (where a male had been observed during the
spring 2010 route was noticeably farther west last few days of April). In total, 7,640 km
than the other routes, being up to 2,150 km were covered at an average of 218 km/day.
west of the autumn 2009 route and up to Some 5,169 km in 21 days (246 km/day) were
1,500 km west of the spring 2009 route. This covered as far as Algeria. During the Sahara
resulted in a ‘migration loop’ for 2009/10, crossing, 1,184 km were covered in just three
which was not the case in 2008/09 (fig. 1). days towards the end of April.
The autumn migration in 2008 (8,525
km) took 49 days, that in 2009 (7,793 km) Autumn migration 2009
took 42 days. Spring migration in 2009 Departure from the breeding area took place
(7,640 km) took 35 days, while the much on 29th or 30th August, the bird initially
longer spring 2010 route (8,678 km as far as heading almost due south. After resting on
the German border) took 38 days. In spring the Mediterranean coast in the vicinity of the
2010 the bird’s migration speed slowed dra- Tunisia–Algeria border from at least 7th to
matically in southern Germany and no fixes 14th September, no other rest stop was
were received after this time. recorded. The bird’s speed was quickest when
crossing the Sahara and the equatorial rain-
Autumn migration 2008 forest, with 850 km being covered in Niger on
After departure on 28th/29th August, the 21st–22nd September and 913 km on 26th–
route initially took a SSW direction and, after 27th September in the rainforest region. In
a short rest from 6th–13th September on the contrast, daily flight distances of only 70–220
island of Elba, off the west coast of
Italy, continued in a SSE direction
as far as North Africa. From Libya
(20th September, 29°50’N
13°38’E) migration resumed again
in a SSW direction, reaching its
most westerly point (10°30’N
4°40’E) on 30th September in
western Nigeria, near the border
with Benin. After an initial 90°
switch in direction to the south-
east, from 8th October on the
Nigeria–Cameroon border (7°N
12°E) the falcon took an almost
southerly course. It held this
course until arrival at its main
wintering area in southern Angola
on 17th October, after a total flight
of some 8,525 km.
Kinshasa
Fig. 3. The marked convergence of the migration routes against the background of the global tree
cover (derived from remote sensing) during the crossing of the West African evergreen rainforest.
The use of the narrowest part of the forest north of Kinshasa is clearly distinguishable.
area but then chose another site (which is then until evening it flew only a further 87
certainly possible since no male occupied the km.
original nest-site northwest of Berlin in
2010). Crossing the Sahara
Crossing the Sahara took some 4–4.5 days on
Migration speed each occasion. In spring 2009, the bird
The transmitter was programmed so that it crossed some 2,100 km of the Sahara in Niger
was generally possible to record the migra- and Algeria between the morning of 22nd
tion distance for a total of two or more con- April and midday on 26th. On 23rd–25th
secutive days only. However, on ten occasions April the average daily distance covered in
on spring migration the flight performance Algeria was 400 km. In spring 2010, the
on a single day was recorded. The minimum Sahara was crossed during 25th–29th April,
daily flight distance was 95 km, the mainly in Mauritania (and to a lesser extent
maximum 481 km and the average 248 in Mali and Morocco), a total distance of
km/day. However, the values of two or more 1,670 km. On five days the bird covered an
days taken together indicate that on some average distance of 440 km, including the
days the falcon flew considerably further than desert fringes. The exact daily flight distance
481 km, and must have covered at least 620 was recorded on three days – 25th April (481
km. On days with short flight distances the km), 26th April (212 km), and 27th April
bird presumably adopted a fly-and-forage (326 km) – followed by the remaining 1,240
migration strategy, especially between the km on 28th–29th. At 08.18 hrs on 25th, the
southern fringe of the Sahara and the bird had already flown 123 km from its night
northern edge of the rainforest. This was also roost, while between 10.00 and 11.00 hrs it
the case south of the rainforest, where the flew only a few kilometres. On 27th, the bird
bird covered only about half the daily dis- started migrating as early as 04.00 hrs (some
tance flown over the key barriers of the two hours before sunrise) but rested from at
Sahara and the rainforest. On 7th October least 06.30 to 11.00 hrs. In three cases, average
2008 in Cameroon, for example, the bird had flight speeds of 25, 30 and 42.5 kph over
covered only 40 km by midday, and from several hours of flight were calculated.
Migration across the rainforest bird flew an additional 260 km after sunset
The West African equatorial rainforest on 14th April 2009, and arrived at its night
appears to be a significant ecological barrier. roost, clear of the rainforest, after midnight.
All four routes across the rainforest (from On one part of the route, 57 km in length, it
approx. 5°45’N 12°33’E to 0°15’S 14°20’E) flew at an average speed of 34 km/h. In
were very close to one another, and from the spring 2010 the Hobby also covered part of
Nigeria–Cameroon border to Angola there the route in darkness until reaching the
was hardly any deviation. The routes passed northern fringe of the rainforest. During the
through a large, roughly north–south cor- early morning of 9th April, before sunrise, it
ridor (or ‘peninsula’) of more open habitats covered over 250 km; thereafter, until
dominated by farmland, degraded woodland midday, it flew only 90 km. The fixes did not
and moist savannah (fig. 3). This corridor is permit us to establish whether it migrated
some 370 km long and 200 km wide and throughout the night.
stretches north from Kinshasa, DR Congo, to The bird migrated in the dark in at least
about 0°15’S 14°20’E. This significantly the early morning hours while crossing the
shortens the distance to be flown over con- Sahara in 2010 (see above). Furthermore, on
tinuous rainforest, to some 450–600 km. 30th April 2010, the Hobby was located over
In autumn 2008, the lowland forest in the Mediterranean near Gibraltar at 02.54
Congo was crossed in a 680-km-long stretch. hrs, some 2.5 hours before sunrise. From its
Here the falcon travelled at a rate of about surmised night roost in Morocco it had at
580 km/day, one of the highest daily dis- this time already covered 122 km and so
tances recorded. In autumn 2009 the distance must have been on the move for at least three
flown over the rainforest was c. 695 km, hours.
covered at a rate of about 450 km/day. In
spring 2009, a stretch of 730 km was flown Wintering
over the rainforest between the afternoon of On both occasions, the Hobby wintered
12th April and c. 01.00 hrs on 15th April at mainly in the same regions (Cunene, Huila
an average of 380 km/day. In spring 2010, and Cuando Cubango) of south and central
daily distance covered over the rainforest and Angola and, to a lesser extent, in the far
adjacent areas was c. 400 km. northeast of Namibia, where it visited the
same areas in both winter periods. The ‘home
Migration during twilight and range’ in winter was very large with the
at night actual size dependent on the method of cal-
Migration at night was recorded repeatedly culation. In 2008/09 it comprised 406,000
while crossing the Sahara and rainforest km² and in 2009/10 it covered 461,000 km²
areas, but also in the Mediterranean. For (Kernel 80%). Calculations based on the
example, in order to reach the northern MCP method arrived at the following values.
perimeter of the rainforest in Cameroon, the In 2008/09, MCP 95% equalled 280,000 km²
Table 1. Habitat use in the wintering area calculated on the basis of identified night roosts, and
taking account of the excursion into Zambia 2008/09.
Discussion
With the advance of satellite-
telemetry technology, information
on the ecology and life history of
raptors has been greatly increased
compared with what could pos-
sibly have been gathered during
decades of conventional telemetry
and ringing. However, when using
telemetry as a study method, the
weight of the transmitter (and its
harness) should not exceed 3% of
the body mass of the study
subject, in order not to affect the
bird’s behaviour and thus bias the
study results (Meyburg & Fuller
2007). This limitation has not
been respected in several recent
falcon studies. The 3% rule still
excludes 85% of all bird species
from satellite telemetry studies
and, of nearly 10,000 species of
birds alive today, only about 1,500
are large enough to carry even the
smallest, 5-g transmitters. The
prototype of what is currently the
smallest transmitter, weighing just
Fig. 7. The Kernel home range in winter 2009/10, based on
5 g, was first used successfully by 77 night roosts (plotted against the background of the global
us on the adult female Hobby tree cover, derived from remote sensing).
described in this paper.
1987). The provision of small birds as food the satellite tracking of four Hobbies between
for the young is clearly an important precon- their breeding grounds in Sweden and their
dition for the long-distance migration flight wintering areas in southern Africa (although
that begins in September, and this, as well as they were not able to follow the birds in
territorial defence, is reliably carried out by winter or on their subsequent spring migra-
the male as the young improve their flying tion).
skills. Our satellite-tracked Hobby showed no
clear preference for the shortest sea crossing
Migration routes when traversing the Mediterranean. Frequent
For most long-distance migrants, ring-recov- sea crossings by the smaller falcons have been
eries from Africa are far too few to properly reported by earlier radar studies at the south
explain the patterns and strategies associated coast of Spain, in both spring and autumn
with migration. We are aware of just two (Meyer et al. 2000, 2003).
such ring-recoveries for the Hobby south of The four flight paths plotted were far
the Sahara. A Dutch bird, ringed as nestling apart across much of Africa, the maximum
on the Wadden Sea Island of Texel on 26th east–west separation being 2,150 km, a dis-
July 1982 (53°03’N 4°43’E), was found dead tance that far exceeds the expected normal
on 26th November 1992 near Kabwe, Zambia range of vision. This suggests that the Hobby
(14°26’S 28°27’E), a recovery distance of did not navigate by following familiar land-
7,820 km (Dowsett et al. 2008); while a marks but that it could find its target regions
nestling ringed in Pälkäne, southern Finland, after extensive deviation, presumably by
on 19th July 2006 was found electrocuted on ‘map-based’ navigation and possibly in com-
7th March 2007 in South Africa, a recovery bination with path integration. The four
distance of 9,650 km (Finnish Ringing routes showed several distinctly non-random
Centre, www.luomus.fi/english/zoology/ features, however. In particular, there was an
ringing/index.htm). In addition, Strandberg et overall westerly bias to the routes, compared
al. (2009a,b) published results obtained from with a direct line between the breeding and
wintering areas. The westerly deviation from spring and autumn of these two species (see
a straight-line route was particularly pro- www.hirondelle.oiseaux.net/migration.html),
nounced over and to the north of Nigeria but the exact migration routes of hirundines
during spring migration, especially in 2010. and swifts within Africa are unknown. In
Furthermore, there was a striking conver- spring, experienced Barn Swallows may be
gence of the four routes over an inland area able to return north in about five weeks at a
close to the equator, well north of the bird’s speed of about 300 km/day (Mead 1970); this
final winter destination in Angola, which again is similar to the performance of the
closely matches the findings of Strandberg et Hobby, which may exploit the swallows as a
al. (2009b). Such convergence of migration convenient prey species. This is also sup-
routes is generally thought to occur in ported by observations in Zambia (see
response to distinctive topography (for below).
example the bottlenecks of large soaring The convergence of the Hobby’s routes
birds at places such as Gibraltar and the near the equator fit neatly with the hypoth-
Bosporus) and/or reflect a species’ navigation esis that the bird aimed to take advantage of
to a particular target area, such as a well- the narrowest crossing of the equatorial rain-
defined winter range. forest, and this suggests that this habitat is
The most challenging aspects of our find- avoided by northern migrants not only for
ings to explain are the extensive migration wintering but also during migration. The
loop, in particular the extreme westerly rainforest may be a barrier not only because
course, of the spring 2010 migration route; the habitat structure of the forest is
the convergence near the equator; and the unfavourable for hunting, but also because of
vast distances that the bird covered in its the weather conditions, with high rainfall.
wintering area. In particular, we have no con- Our results thus support Strandberg et al.
vincing explanation for the 2010 spring (2009b) that the continuous evergreen rain-
migration route, especially given that most forest constitutes an important barrier that
species usually migrate more to the east in affects the major flyways and stopover strat-
spring than in autumn (Newton 2008). egies by which Hobbies, and perhaps other
Analysis of the data on wind direction and migratory landbirds, travel to and from their
precipitation from all meteorological stations winter quarters in southern Africa. It is most
along the migration routes provided us with important that we understand the role of the
no further clues, although Strandberg et al. rainforest as an ecological barrier of possibly
(2009b) suggested that the westward shift of continental-scale significance for the evolu-
routes they observed over the Sahara was in tion and future of the Palearctic–African bird
response to drift in easterly crosswinds. Inter- migration systems.
estingly, two adult Eleonora’s Falcons F.
eleonorae also migrated much farther to the Migration speed
west in spring than in autumn (Gschweng et The flight speed of the Hobby tracked in our
al. 2008). One contributory factor to the study during outward migration was signifi-
bird’s route could be the migration tracks of cantly faster (8,525 km in 49 days or 7,793
Barn Swallows Hirundo rustica, House km in 42 days, 174 and 185 km/day respec-
Martins Delichon urbicum, Common Swifts tively) than the somewhat longer migrations
Apus apus and other important prey species of the Swedish Hobbies tracked by Strand-
(hirundines and swifts make up 55% of the berg et al. (2009a), which needed on average
avian prey species of the Hobby around 61 days for a mean distance of 9,223 km (151
Berlin; Fiuczynski et al. 2009). Barn Swallows km/day). To date there are no other data for
in particular have similar migration routes us to compare with the spring migration
and wintering areas to the Hobby, and depar- routes described here. The differences that we
ture and arrival times in Europe are also very found in migration speed during the Sahara
similar to those of the Hobby (e.g. Sparks & and rainforest crossings, compared with
Braslavská 2001). In France in recent years other parts of the route, were also recorded
(2008–10), there has been a close similarity for the Swedish Hobbies. The daily rains
in terms of the passage periods in both associated with the rainforest may limit the
Bernd-Ulrich Meyburg is chairman of the World Working Group on Birds of Prey (WWGBP). He has carried
out research on no fewer than 16 raptor species since 1992 using satellite telemetry. Paul Howey is president
and owner of Microwave Telemetry, Inc. (USA), and a pioneer in producing satellite-tracking devices for birds.
He built the first 5 g satellite transmitters. Christiane Meyburg worked as a researcher at the National
Institute of Statistics and Economic Studies (INSEE) in Paris. Since 1992 she has helped her husband analyse
the satellite-tracking data from his projects. Klaus Dietrich Fiuczynski worked as a teacher, completed a PhD
on the Hobby and has written a major monograph on this species (a new edition of which is almost
complete).