modern telemetry: new possibilities in ornithology

ISSN 1062�3590, Biology Bulletin, 2011, Vol. 38, No. 9, pp. 885–904. © Pleiades Publishing, Inc., 2011.Original Russian Text © L.V. Sokolov, 2011, published in Zoologicheskii Zhurnal, 2011, No. 7, pp. 861–882.

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INTRODUCTION

The invention of new methods for tracking birdmovements has significantly broadened and, in somecases, radically changed our views on various aspectsof bird life. Attempts to mark individual birds areknown from the Middle Ages, when trapped heronswere fitted with tags or rings and then released (Stein�bacher, 1956). Bird marking for scientific purposes wasinitiated in 1890 by Hans Christian CorneliusMortensen, a Danish schoolteacher. He initiallymarked European starlings with zinc tags but in 1899began to use lightweight metal rings attached to birds'legs. Each ring had on it a unique number and theresearcher’s address. Three years later, German pro�fessor Johannes Thienemann defined bird banding asa major objective of research at the Rossitten BirdObservatory, which was established in 1901 on theCourish Spit, the Baltic Sea. Banding has become thebasic method of studies on bird movements. Bandingschemes implemented in different countries over morethan a century allowed specialists to compile a vastdatabase on retraps of banded birds, which is and willbe highly useful for bird studies. However, the methodof individual banding has certain drawbacks and limi�tations. In particular, birds should be banded en masseto achieve the number of retraps sufficient for analysis,but this does not provide the possibility of tracing theroutes of bird movements. This explains the necessityfor new methods of bird tracking.

Such methods began to appear in the 1960s, afterportable radio transmitters had been invented. Ini�tially, these were fairly cumbersome ultra�high�fre�quency (UHF) devices that could be attached only tolarge birds. To use them, special power sources, anten�nas, and receivers were necessary. Today, an assort�ment of tracking devices is available, from tiny short�range transmitters weighing less than 1 g to long�life

satellite transmitters weighing 5–20 g that allow notonly local movements but also extremely long migra�tions of birds to be tracked over several years.

After successful application in research on migra�tions of wild animals, telemetry has been used to studytheir local movements, dispersal, the size of individualhome ranges, location of habitats, etc. The availabilityof satellite transmitters has given rise to active studieson the routes, rates, and altitudes of bird migrations.Telemetry has also proved to be efficient in analyzingsocial behavior, intra� and interspecific relationships,mortality, and population dynamics of different ani�mal species (White and Garrott, 1990; Benvenuti,1993; Beekman et al., 1996; Seegar et al., 1996; Guanand Higuchi, 2000; Kenward, 2001; Fuller et al., 2005;Olival and Higuchi, 2006; Meyburg and Fuller, 2007;Whitworth et al., 2007; Hart and Hyrenbach, 2009).Today, this method is also increasingly used in studieson invertebrates, including insects.

ORNITHOLOGICAL PROBLEMS BEING SOLVED WITH THE AID OF TELEMETRY

Today, transmitters are used in studies on the fol�lowing aspects of bird biology:

(1) migration routes and directions (Higuchi et al.,1991, 2006; Meyburg et al., 1995, 2002, 2006; Johnsonet al., 1997; Fuller et al., 1998; Guan and Higuchi,2000a, 2000b; Kanai et al., 2000; Berthold et al., 2001,2004; Hake et al., 2001, 2003; Green et al., 2002; Foxet al., 2003; Haines et al., 2003; Kaatz, 2004; Gillet al., 2005; Judas et al., 2006; Trierweiler et al., 2007;Gschweng et al., 2008; Pütz et al., 2008; Stutchburyet al., 2009; López–López et al., 2009, 2010; Bächleret al., 2010; Egevang et al., 2010; etc.);

(2) migration speeds and altitudes (Bögel and Bur�chard, 1992; Javed et al., 2000; Hake et al., 2001,

Modern Telemetry: New Possibilities in OrnithologyL. V. Sokolov

Zoological Institute, Russian Academy of Sciences, St. Petersburg, 199034 Russia;e�mail: leonid�[email protected]

Received December 15, 2010

Abstract—Modern methods of bird telemetry and some research results obtained using these methods areconsidered. The development of high�technology methods for tracking bird movements has significantlybroadened and, in some cases, radically changed our views on the life of birds. After the invention of variousradio transmitters for birds (from simple, short�range to long�life satellite devices), new possibilities haveopened up for studies in many areas of ornithology.

Keywords: telemetry, ornithology, bird movements.

DOI: 10.1134/S1062359011090081

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2003; Kjellén et al., 2001; Meyburg et al., 2002, 2006;Miller et al., 2005; Soutullo et al., 2006; Strandberget al., 2009; Hedenström, 2010);

(3) duration of migration flights and stopovers(Aborn and Moore, 1997; James et al., 2000; Harriset al., 2000; Kanai et al., 2002; Shimazaki et al., 2004;Fransson et al., 2008; Galarza, Dennis, 2009; Strand�berg et al., 2009; Chernetsov, 2010);

(4) the timing of the start of nocturnal migrationand dispersal (Cochran et al., 1967; Åkesson et al.,1996, 2001; Moore and Aborn, 1996; Bolshakov andChernetsov, 2004; Bolshakov et al., 2007; Mukhinet al., 2009);

(5) the physiological and energetic state of birds(Sawby and Gessamen, 1974; Bögel and Burchard,1992; Weimerskirch et al., 1995; Pennycuick, 1996;Clausen et al., 2003; Pennycuick and Battley, 2003;Bowlin et al., 2005; Gill et al., 2005; Tsvey et al., 2007;Hedenström, 2010; Kosarev and Kobylkov, 2010;Stutchbery et al., 2011);

(6) orientation and navigation (Gudmundssonet al., 1995; Mouritsen et al., 2003; Chernetsov et al.,2004, 2005; Cochran et al., 2004; Thorup et al., 2007;Wikelski et al., 2007);

(7) homing and philopatry (Southern, 1970; Kosa�rev and Sokolov, 2007a; Mukhin et al., 2009; García�Ripollés et al., 2010; Oppel and Powell, 2010);

(8) nesting and postnesting dispersal (Bahat, 1992;Griesinger et al., 1992; Meyburg and Lobkov, 1994;Walls and Kenward, 1994; Hyrenbach and Dotson,2001; Rafanomezantsoa, 2002; McGrady et al., 2003;Mukhin et al., 2005; Glahder et al., 2006; Soutulloet al., 2006; Weimerskirch et al., 2006; Cadahía et al.,2008; Anker�Nilssen and Aarvak, 2009; Pearce andPetersen, 2009);

(9) individual home range size (Naef�Paenzer,1993; Kralovec, 1994; Jiguet and Villarubias, 2004;Judas et al., 2006; Meyburg et al., 2006);

(10) the timing of seasonal phenomena (Martellet al., 2001; Fox et al., 2003; Meyburg et al., 2005;Battley, 2006; Glahder et al., 2006; Kosarev andSokolov, 2007b);

(11) territorial and social behavior (Walls and Ken�ward, 1994; Tyack et al., 1998; Driscoll and Ueta,2002; Javed et al., 2006; Pütz et al., 2007; Therrienet al., 2008);

(12) territorial and biotopic distribution (Jouventinand Weimerskirch, 1990; Weimerskirch, 1990; Ancelet al., 1992; Grubb et al., 1994; Kanai et al., 1994; Falkand Møller, 1995; Davis et al., 1996; Higuchi et al.,2000; Berthold et al., 2001; Tamura et al., 2001;Kenow et al., 2002; Ueta et al., 2002; Aarvak andØien, 2003; Haines et al., 2003; Morimoto et al.,2005; Glahder et al., 2006; Meyburg et al., 2006; Mos�bech et al, 2006; Peterson et al., 2006; Therrien et al.,2008; Ktitorov et al., 2010);

(13) demography (Snyder et al., 1989; Bunck et al.,1995; Esler et al., 2000; Cadahía et al., 2005; Murray,

2006; Strandberg et al., 2010; Oppel and Powell,2010);

(14) ecology (Weimerskirch et al., 1993; Hamer et al.,2000; Glahder et al., 2006; Mosbech et al., 2006;Therrien et al., 2008);

(15) species conservation, acclimation, and rein�troduction (Higuchi et al., 2004; Galarza and Dennis,2009);

(16) transmission of pathogenic viruses (Whitworthet al., 2007; Prosser et al., 2009; Gaidet et al., 2010);

(17) locations and redistribution of game species; etc.Transmitters attached to birds, penguins and sea�

birds in particular, can be used to collect informationon hydrological and aerological parameters, includingwater temperature and transparency at differentdepths, air temperature and flow velocity, etc. (Bögeland Burchard, 1992; Wilson et al., 1994).

Since telemetry methods are fairly labor�intensiveand expensive, compared to simple banding, theresearcher has to explicitly determine the purposesand plan of the study, including (1) the size and type ofthe transmitter, (2) the most convenient and safe wayof its attachment to the bird, (3) the optimal mode ofbird tracking, (4) the possibility to retrap the bird andreplace or remove the transmitter, and (5) the optimalmethods of telemetry data processing and analysis.

Various aspects of planning and conducting radio�telemetry studies are addressed in many publications,which can be used as a basis for developing an originalresearch project (Kenward, 2001; Coyne and Godley,2005; Fuller et al., 2005; etc.). Strict regulations con�cerning wild bird trapping, treatment, and transmittertagging are in effect in many countries, including Rus�sia, and all necessary authorization documents andlicenses for such studies should be obtained inadvance.

Ultra�high�frequency (UHF) transmitters

The first transmitters used for bird tracking weresimple but cumbersome UHF devices attached to thebird’s back with special fittings. The equipment kitalso included a power source, antenna, and receiver.Today, the assortment of such transmitters ranges fromrelatively large ones, which remain in service for sev�eral months, to tiny (0.2 g) devices with a service life ofonly 5 days.

The basic service parameters of UHF transmittersare weight, power output (transmission range), andbattery service life (Table 1); the higher the power out�put, the shorter the battery life, and vice versa (Whit�worth et al., 2007). The optimal choice of transmitterpower depends on the purposes of research. If the birdsare expected to move over a large area, it is necessaryto increase the transmission range at the expense ofbattery life; conversely, if the birds are to be monitoredwithin a relatively small area, it is expedient to reducethe transmitter power output to save battery life and,


MODERN TELEMETRY 887

therefore, prolong the observation period. Since theservice parameters of transmitters depend on theirsize, they are better applicable to telemetry of rela�tively large birds, whereas the spatial extent and dura�tion of such studies on small birds are limited.Research involving UFH telemetry is more labor�intensive than satellite tracking, since it requires alarge amount of manual work and more manpower.This adds importance to the transmitter parameterssuch as power output (transmission range) and batterylife.

UFH telemetry requires a considerable amount ofsurvey effort aimed at locating transmitter�taggedbirds and fixing their coordinates. Signals from thetransmitter are detected with an UHF receiver con�nected to an antenna with a special cable. Modernreceivers can be programmed for scanning certain fre�quencies at specified intervals until the signal isdetected. During route surveys (on foot or on a vehi�cle), a triangulation method is used for accurate signallocalization. Scanning with a directional antenna isperformed from a fixed position with known coordi�nates to take the bearing to the signal source (thedirection wherefrom the strongest signal is received),and then this procedure is repeated from a differentpoint. The bearings are then plotted on a map, andtheir intersection point indicates the approximatelocation of the signal source. Simultaneous surveyingfrom an aircraft and on the ground (or on board of awatercraft) is usually the most efficient and cost�effec�tive approach. Aerial tracking is characterized by agreater spatial coverage and a longer range of signalreception (Whitworth et al., 2007), but the location ofthe signal source is determined with relatively lowaccuracy; in addition, aircraft time is expensive. On�ground tracking, in contrast, may be accurate to 5 m,and transmitter�tagged birds can sometimes beobserved directly if they are within a visibility range. In

most cases, however, visual observations are impossi�ble, and the location of the birds is determined more orless approximately (depending on the trackingmethod). Thus, aerial tracking may be used forapproximate signal location in large areas, and on�ground tracking, for determining the accurate coordi�nates of tagged birds. The range of signal reception onthe ground increases when scanning is performed froman elevation. Transmitters can be fitted with program�mable data loggers (some transmitters have them builtin), which allow bird tracking from fixed receiver sta�tions. Data loggers are instrumental in confirming thepresence or absence of transmitter�tagged birds withina relatively small area. Like transmitters, they are pow�ered by a built�in battery. External power sources (e.g.,solar cells) may be used to prolong their service life.Loggers can be programmed to operate continuouslyor to switch on and off at certain intervals in order tosave power. The logged data can be directly uploadedto a laptop computer and processed in the field. Theprocedure of marking the locations of telemetry�tracked objects on a map has practically become obso�lete due to the development of the Global PositioningSystem (GPS) and commercial availability of reliable,precise, and affordable GPS receivers. These devicesare especially instrumental in determining the coordi�nates of transmitter�tagged birds or receiver stationsand delimiting the zones detected in the course oftelemetry tracking. GPS receivers are portable, simpleto operate, and compatible with the majority of com�puter programs for spatial data analysis, which makesthem an indispensable tool for any kind of radiotelem�etry research (Whitworth et al., 2007). The range ofUHF transmitters available today includes tinydevices weighing only 0.2–0.3 g, and they can be usedfor the study of any bird species.

Table 1. Parameters of modern transmitters used in bird studies

Parameter

Transmitter

Local, UHFsatellite

Geolocator loggerPTT GPS

Weight, g 0.2–12 5–18 20–60 0.4–12

Service life 5 days to several months

Several months to several years



Receiving range, km 0.1–100 Unlimited Unlimited –

Positioning accuracy 5–500 m 100–200 m 10–20 m 50–100 km

Tracking interval Continuous tracking

4 hours Continuous tracking Continuous tracking

Type of tracking Antennal Satellite Satellite Logging

Power source Power source Rechargeable battery, solar cell

Rechargeable battery, solar cell

Power source

Price $100–300 $2000–3000 $2500–3500 $100–200

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Satellite transmitters

There are two basic methods for determining thecoordinates of a transmitter�tagged bird using satellitesystems. In the first (simple) case, the coordinates ofthe transmitter are calculated from the Doppler shiftin the frequency of its signal received by the satellite.This method is simple and employs relatively light�weight transmitters, but its accuracy is low (Table 1).In the second case, the bird is fitted with a GPS devicethat receives signals from several satellites and calcu�lates its own position by comparing the relative powerof these signals. In most cases, the accuracy of thismethod is several tens of meters. The coordinatesdetermined in this way are transmitted to theresearcher immediately or with a short delay via thesecond communications system, which is not neces�sarily a satellite system: in most cases, a GSM cellphone or a short�range transceiver can be used to pickup these data. In particular, Alexander Minaev(http://moosefarm.newmail.ru) considers the use ofradio tags that collect the results of measurements andtransmit them over a public communications system(i.e., a system commercially accessible to any user).

Recent technological advancements have made itpossible to develop relatively small platform terminaltransmitters (PTTs) and GPS transmitters. Specificfeatures and parameters of PPT, UHF, and GPSdevices account for their application to different tasksof research on different bird species (Table 1). Thechoice of a PTT or GPS transmitter should be madewith regard to its size: the transmitter weight shouldnot exceed 2–3% of the bird body weight, although forsmall birds (<50 g) this proportion may be increased to3–5%.

Until recently, the smallest PTTs weighed 12–18 g,which interfered with their use in studies on bird spe�cies with a body weight of less than 500 g, such as smallducks or gulls. In contrast, some modern PTTs weighno more than 5 g and can be attached to cuckoo�sizedbirds (Kristensen, 2010). They are powered by solarcells or special batteries and remain in service for sev�eral months to several years, depending on the fre�quency of signal transmission to satellites. Currently,the most widespread is the ARGOS system, which hasfour satellites circling the earth on polar orbits at analtitude of 850 km. The satellites receive UHF signals(401 MHz) from more than 8000 ARGOS PTTs thatare currently in operation. Each satellite receives sig�nals from all transmitters within a radius of 5000 km,in all time zones, and sends them in real time tonumerous ground stations. This information isrecorded and transmitted to computer centers, wherethe coordinates of PTT�tagged birds are calculated withan accuracy of up to 150–300 m using Doppler shiftanalysis (Table 1). Such messages are sent every 65 s.

GPS transmitters weigh 20–60 g and can be usedmainly for large birds weighing about 1 kg or more,such as geese and swans. They send signals at two dif�

ferent frequencies to 24 satellites circling on circularorbits at an altitude of 20 000 km. The times of signalarrival at the two frequencies are compared to calcu�late the delay arising during the passage of radio wavesthrough the ionosphere, which allows the coordinates ofthe transmitter to be determined with an accuracy of upto 10–20 m. Thus, this tracking method is more accuratebut also more expensive than ARGOS tracking.

Geolocator loggers. Along with the above transmit�ter types, increasing use is made of devices referred toas geolocator loggers. They were invented by Russianscientist Vsevolod Afanasyev, who now lives inEngland. A logger is a programmable chip with anelectronic chronometer coupled to a light sensor,which is designed to record the level and timing of nat�ural illumination at certain intervals. Data on theduration of daylight and the time of sunrise and sunsetallow the coordinates of the tagged bird to be calcu�lated with an accuracy of 50–100 km (Table 1). How�ever, to retrieve these data from the logger’s memoryand upload them to a computer for analysis, it is nec�essary to retrap the bird and remove the logger.Another drawback of this method is that the illumina�tion level and the time of sunrise and sunset can beaccurately measured only if the logger is attached to abody part that is open to daylight. For example, itsattachment to the leg is ineffective in the case of sea�birds, which spend considerable time floating on thewater. Hence, loggers are often fitted with a small fiberoptic antenna protruding above the plumage surface,which allows the logger to measure illumination level.Regardless of these drawbacks, geolocator loggers arewidely used in bird studies. They are relatively inex�pensive (in Switzerland and Germany, for example,certain types can be purchased for $100) and havebecome increasingly lightweight in recent years. Thesmallest loggers weigh less than 1 g and can be used onsmall passerine birds. The battery life ranges from 2 to5 years, depending on logger size and type.

Transmitters of the above types can also bedesigned to perform other useful functions, but thisincreases their weight, energy consumption, and cost.Transmitters can be equipped with sensors controllingambient and body temperatures, pressure, heart rate,wing stroke rate, etc. (Bowlin et al., 2005) and with atimer switching the devise of and off at certain inter�vals, which helps to prolong the battery life.

The effect of transmitter tagging on birds markedlydepends on the design of the transmitter and themethod of its attachment. It is usually recommendedto use smaller transmitters, as large and cumbersomedevices are more likely to produce an adverse effect. Inparticular, external transmitters increase aerodynamicdrag during flight and hydrodynamic drag in divingbirds. It has been shown that transmitter�tagged birdshave higher mortality, lower reproductive success andchick feeding rate, and are more susceptible to otherdisturbances (Whitworth et al., 2007). Ideally, thetransmitter should remain attached to the bird



throughout the study period and fall off as soon as thestudy is completed, but such a situation is very rare.There is no guarantee that the transmitter will remainattached to the bird, whatever the method used for thispurpose. For example, we used short�range transmit�ters to study homing in the pied flycatcher (Ficedulahypoleuca) on the Courish Spit and failed to removethem from five birds after the end of the experiment. Inthe next year, one of them returned from wintering inAfrica, still with the transmitter, and began nestingactivities, despite that the thin antenna was bent andinterfered with the bird’s entry into the nest box. Notethat the manufacturer guaranteed that the rubber bandused for attaching the transmitter would break aftertwo months. The presence of a transmitter can affectthe bird’s behavior, at least until the bird gets used toit. Some bird species show an extremely adverseresponse to transmitter attachment and can be effec�tively tagged only by devices implanted intraperito�neally or subcutaneously. Such implantation is a fairlycomplex surgical procedure that should be performedby a skilled specialist. An analysis of relevant publica�tions and advice from experienced researchers willhelp in choosing the most effective method of trans�mitter attachment. It is also advisable to conduct afield test on a small number of birds to reveal the prob�able adverse consequences of transmitter tagging.

Analysis of the movements and spatial distributionof animals by means of telemetry has proved to be avery complex process (Kenward, 2001; Whitworth etal., 2007). In some cases, such as studies on localmovements, the size of home ranges, or biotopic dis�tribution of birds, the analytical procedure can bereduced to connecting localization points so as to con�struct the smallest possible convex polygon coveringthe entire area used by the tagged bird (the minimumconvex polygon method). More complex probabilisticmodels allowing the analysis of differences in the pat�tern of home range use make necessary recourse togeographic information systems (GIS). Some knowl�edge of GIS is essential for anyone dealing with dataon animal movements. These systems provide the pos�sibility of complex mapping aimed at visual or statisti�cal analysis of relationships between the locations oftagged birds and habitat conditions, including climaticvariables (Whitworth et al., 2007). Satellite images ofthe earth’s surface accessible from Internet sites, suchas Google Earth (http://www.earth.google.com), canbe supplemented with user data, e.g., the coordinatesof points determined by GPS survey or the tracks ofbird movements. In particular, the program ArcviewGIS (Whitworth et al., 2007) offers a broad range ofoptions allowing the user to mark the locations of atagged bird on the map and quickly calculate the dis�tances and speed of its movement, evaluate the pat�terns of movements and home range and habitat use,and to solve a number of other tasks by means of spa�tial analysis. Detailed information on the methods oftelemetry data analysis can be found in special review

articles (White and Garrott, 1990; Fuller et al., 2005;Coyne and Godley, 2005; Meyburg and Fuller, 2007;Hart and Hyrenbach, 2009)

Migration studies. Traditional views on the routesand directions of bird migrations, based mainly onvisual observations and the results obtained over a cen�tury of bird banding, are changing in the context ofnew data provided by satellite telemetry technology.Important novel information on various aspects ofmigration in the nesting and wintering parts of therange has been obtained by means of satellite trackingeven for well�studied species such as the white stork(Ciconia ciconia), which has been banded en masseover more than 100 years and numerous ring recoverieshave been received along its migration routes. Duringthe past 20 years, German ornithologists headed byProf. Peter Berthold have satellite tagged approxi�mately a hundred white storks in Germany, Poland,Israel, and the Kaliningrad Region of Russia (Ber�thold et al., 2001, 2004; Kaatz, 2004). As a result, theyhave obtained valuable information on the directionsand distances of migration of birds from eastern andwestern populations, the timing (dates) of flights andstopovers (Fig. 1), flight altitudes and speeds at differ�ent stages of autumn and spring migrations, mortalityamong migrating birds, etc. Data on the migrationroutes of one pair of storks nesting in Germany are ofspecial interest. As shown by satellite tracking, thefemale migrated in autumn to the wintering grounds inSouth Africa, while the male overwintered in Spain.Regardless of the great distance between the winteringareas, both birds started the return migration on thesame day, February 22, but arrived to their nest at dif�ferent times: the male on March 9, and the female onApril 19.

No less interesting new data on migration routeshave been obtained for diurnal birds of prey (Table 2;Figs. 2, 3). For example, on the basis of visual observa�tions performed in the past century, specialists consid�ered that the small Eleonora’s falcon (Falco eleonorae)nesting on the Mediterranean islands migrates to itswintering grounds on Madagascar along the Mediter�ranean and Red Sea coasts and them along the easterncoast of Africa (Stresemann, 1954). However, recentstudies on its migration routes by means of satellitetracking have revealed remarkable facts (Gschwenget al., 2008; López�López et al., 2009). First, thesebirds do not migrate along sea coasts but fly directlyacross the Sahara Desert and the subequatorial belt ofAfrica, changing the flight direction several times, andapproach the Mozambique Channel (separatingMadagascar and Africa) from the mainland side (Fig. 3).Second, adult and young Eleonora’s falcons followdifferent migration routes. Adult birds fly to the win�tering grounds more or less directly, as describedabove, while the route taken by young birds is muchmore complex. They first fly south�southwest, towardthe Gulf of Guinea, but turn east or southeast at a lat�itude of central Nigeria (about 10° N) and follow to

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50°

40°

30°

20°

10°

0°

10°

20°

30°

Fig. 1. Autumn migration routes of white storks (individual birds) from different regions of Europe according to satellite telemetrydata (from Kaatz, 2004).

Uganda for a long stopover. Then, they fly southeast tothe Mozambique coast and make an attempt to crossthe Mozambique Channel, which is about 500 kmwide. They do not always succeed: many birds land onsmall islands in the channel or return to the continentand remain there for several weeks before repeating theattempt to reach their traditional wintering grounds.These data raise a number of principally importantquestions. The main question is how these youngbirds, which have not yet wintered on Madagascar andcould not get imprinted to migrate there (i.e., learn thecoordinates of their wintering grounds), manage tocover such a long and difficult route alone, without thecompany of adult birds. Specialists as yet have no def�inite answer to this question. Furthermore, the return(spring) migration route of young of the year and adultEleonora’s falcons has also proved to be largely unex�pected: instead of flying from Madagascar northwest�ward, directly to the nesting grounds in Italy and

Greece, they move westward, sometimes reaching thewestern coast of Africa in the regions of Senegal andMauritania (young birds) or Morocco (adult birds),turn northeast to northern Italy, and only then make asharp turn south to arrive to the Mediterranean islands(Fig. 3).

Birds migrating to their wintering or nestinggrounds often follow unexpected routes. At the 25thInternational Ornithological Congress in Campos doJordáo (Brazil), Higuchi presented satellite trackingdata on the return migration of crested honey buzzards(Pernis ptilorhynchus) to their nesting range in Japan(Yamasuchi et al., 2008). As in the study on Eleonora’sfalcons, it was found that these birds sometimesarrived from the north rather than from the south, asthey should have in the case of direct spring migration.

In telemetry studies on migrations of the osprey(Pandion haliaetus) from North America to Centraland South America and back, American researchers



Table 2. Bird species whose migration routes have been traced by means of telemetry

Species References

Emperor penguin Aptenodytes forsteri Ancel et al., 1992

Adelie penguin Pygoscelis adeliae Davis et al., 1996

Common loon Gavia immer Kenow et al., 2002

Wandering albatross Diomedea exulans Jouventin, Weimerskirch, 1990; Nicholls et al., 1996; Weimerskirch, et al. 2006

Black�footed albatross Phoebastria nigripes Hyrenbach, Dotson, 2001

Cory’s shearwater Puffinus diomedea Ristow et al., 2000

Dalmatian pelican Pelecanus crispus Morimoto et al., 2005

Great frigatebird Fregata minor Weimerskirch et al., 2006

Magnificent frigatebird Fregata magnificens Weimerskirch et al., 2006

White stork Ciconia ciconia Kaatz, 2004

Oriental white stork Ciconia boyciana Shimazaki et al., 2004; Higuchi et al., 2000

Black stork Ciconia nigra Jiguet, Villarubias, 2004

Trumpeter swan Cygnus buccinator Strikwerda et al., 1986

Whooper swan Cygnus cygnus Pennycuick et al., 1996; Kanai et al., 1997

Tundra swan Cygnus columbianus Higuchi et al., 1991

Ruddy shelduck Tadorna ferruginea Prosser et al., 2009

Greater white�fronted goose Anser albifrons Fox et al., 2003

Snow goose Anser caerulescens Blouin, 1999

Lesser white�fronted goose Anser erythropus Lorentsen et al., 1998; Aarvak, ien, 2003

Bar�headed goose Anser indicus Javed et al., 2003; Prosser et al., 2009

Pink�footed goose Anser brachyrhynchus Glahder et al., 2006

Brent goose Branta bernicla Green et al., 2002

Northern pintail Anas acuta Miller et al., 2005

Spectacled eider Somateria fischeri Petersen et al., 1999

Steller’s eider Polysticta stelleri Petersen et al., 2006

Egyptian vulture Neophron percnopterus García�Ripollés et al, 2010

Griffon vulture Gyps fulvus Griesinger et al., 1992

Osprey Pandion haliaetus Kjellén et al., 1997; Hake et al., 2001; Stout, Green, 2009

European honey buzzard Pernis apivorus Hake et al., 2003; Higuchi et al., 2005;

Montagu’s harrier Circus pygargus Limiñana et al., 2007

Marsh harrier Circus aeruginosus Strandberg et al., 2010

Common buzzard Buteo buteo Walls, Kenward, 1995; Strandberg et al., 2009

Broad�winged hawk Buteo platypterus Haines et al., 2003

Swainson’s hawk Buteo swainsoni Fuller et al., 1998

Golden eagle Aquila chrysaetus Brodeur et al., 1996

Eastern imperial eagle Aquila heliaca Meyburg, Meyburg, 1998

Steppe eagle Aquila nipalensis Meyburg, Meyburg, 1998; Meyburg et al., 2003

Lesser spotted eagle Aquila pomarina Meyburg et al., 2000, 2006

Greater spotted eagle Aquila clanga Meyburg et al., 2005

Short�toed snake eagle Circaetus gallicus Meyburg et al., 1998

Bonelli’s eagle Hieraaetus fasciatus Cadahia et al., 2005, 2008

White�tailed eagle Haliaeetus albicilla Ueta et al., 1998

Bald eagle Haliaeetus leucocephalus Grubb et al., 1994

Steller’s sea eagle Haliaeetus pelagicus Meyburg, Lobkov, 1994; McGrady et al., 2003

Ø′

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have found that young birds prefer moving along thecoast and over the Caribbean islands, whereas youngbirds make nonstop 2000�km flights over the open sea.Some adult birds fly the same route from year to year,but in other cases the same bird chooses a differentroute every autumn, apparently depending on weatherconditions of the season (Martell et al., 2001; Stoutet al., 2009).

Russian ornithologists have obtained interestingdata on autumn migration routes of poorly studiedtundra peregrine falcons from the Yamal Peninsula(Sokolov et al., 2010). Some female birds migrated toSpain or Greece, while others preferred wintering inSaudi Arabia or Sudan. It is noteworthy that these fal�cons nested in the desolate tundra but often winteredin densely populated cities, preying on pigeons.

Satellite tracking provided evidence for the abilityof migrating birds to fly nonstop over great distances,which could not be revealed traditional tracking meth�ods. Thus, one of transmitter�tagged whimbrels(Numenius phaeopus) in spring made a 5150�km non�stop flight over the territories of the United States andCanada, covering this distance in 146 hours (6 days)(Watts, 2008). Another sandpiper species, the bar�tailed godwit (Limosa lapponica), makes even longernonstop flights over the Pacific. These birds migrate inautumn (August 30–September 7) from Alaska toNew Zealand (11 700 km) and return in spring by adifferent route, covering 10 300 km over the oceanfrom New Zealand to China (March 17–24) and then

6500 km from China to Alaska (May 2–8) (Gill et al.,2005; Hedenström, 2010). Seven female bar�tailedgodwits with implanted transmitters covered the dis�tances of 8117 to 11 680 km nonstop within no morethan 6–9 days! Observations have shown that thesebirds start migrating from Alaska during tailwinds. Thecapacity for long�distance nonstop flight is evidencefor outstanding physiological properties of the birds(Pennycuick, 2003; Gill et al., 2008).

Important data on long�distance bird migrationshave also been obtained with geolocator loggers. Inparticular, these relatively new devices have been usedin studies on the arctic tern (Sterna paradisae), whichconducts the longest known migrations (among birds)from the Arctic to Antarctica and back (Egevang et al.,2010). The results show that arctic terns migrate inautumn mainly along the western coast of Africa,while the routes of spring migration lie closer to Northand South Americas (Fig. 4). In autumn, the birdscover a total of about 34 600 km at an average speed of330 km/day; in spring, 25 700 km at 520 km/day; con�sequently, the period of migration averages 93 (69–103) days in autumn, compared to only 40 (36–46)days in spring. The majority of birds arrive to the win�tering region by November 24 (the total period of arrivalis from October 25 to November 30) and depart on April12–19, remaining in the region for 139–173 days. Thewintering birds actively move over the region, with thedistance of their movements during the season averag�ing 11 000 km. Thus, the total distance covered by the

Table 2. (Contd.)

Species References

Peregrine falcon Falco peregrinus Fuller et al., 1998, Sokolov et al., 2010

Eurasian hobby Falco subbuteo Strandberg et al., 2010

Eleonora’s falcon Falco eleonorae Gschweng et al., 2008; López�López et al., 2009, 2010

Common crane Grus grus Higuchi et al., 1992

Hooded crane Grus monacha Higuchi et al., 1992; Harris et al., 2000

Red�crowned crane Grus japonensis Higuchi et al., 1998

White�naped crane Grus vipio Higuchi et al., 1996; 2004; Harris et al., 2000

Siberian crane Grus leucogeranus Kanai et al., 2002

MacQueen’s bustard Chlamydotis macqueenii Judas et al., 2006

Far Eastern Curlew Numenius madagascariensis Driscoll, Ueta, 2002

Whimbrel Numenius phaeopus Watts et al., 2008

Bar�tailed godwit Limosa lapponica Green et al., 2002; Gill et al., 2005, 2008;

Lesser black�backed gull Larus fuscus Pütz et al., 2008

Arctic tern Sterna paradisae Egevang et al., 2010

Northern fulmar Fulmarus glacialis Falk, Møller, 1995

Common guillemot Uria aalge Hatch et al., 2000

Thick�billed guillemot Uria lomvia Hatch et al., 2000

Atlantic puffin Fratercula arctica Anker�Nilssen, Aarvak, 2009

Common cuckoo Cuculus canorus Kristensen, 2010



arctic terns over the year reaches 71 000 km. In addi�tion, it has been shown that these birds have a stopoversite in the North Atlantic, where they remain for about25 days (mainly in the second half of August).

Recently constructed lightweight loggers (<1 g)suitable for tagging small passerine birds allowedAmerican researchers to track the migration routes ofthe wood thrush (Hylocichla mustelina) and purplemartin (Progne subis) over the Gulf of Mexico. Analy�sis of data on several birds has shown that woodthrushes in autumn and spring prefer to migratedirectly across the gulf, whereas purple martins nest�

ing in Pennsylvania migrate to South America andback mainly over the land (Stutchbury et al., 2009).Both species migrated several times more rapidly inspring than in autumn. One female purple martindeparted from the sintering grounds in the AmazonRiver basin on April 12 and covered about 7500 km in13 days (9 days in flight and 4 days at stopover sites)with an average speed of 577 km/day. The springmigration speed of thrushes was approximately halflower, 233–271 km/day.

Bird tracking by means of satellite telemetry allowsresearchers to obtain valuable information not only on

Fig. 2. Autumn migration routes of Montagu’s harriers (individual birds) from western and eastern Europe according to satellitetelemetry data (from Limínana et al., 2007).

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SOKOLOV

the direction and range of migration but also on theflight speed in different segments of the route, flightaltitude at different times of day, etc. For example,

studies on the autumn migration of honey buzzards(Pernis apivorus) from Europe to East Africa showedthat the migration speed of both adult and young birds

(a) (b)

(c)

Fig. 3. Autumn and spring migration routes of young (n = 7) and adult (n = 6) Eleonora’s falcons according to satellite telemetrydata (from Gschweng et al., 2008): (a) autumn migration of young birds, (b) autumn migration of adult birds, and (c) springmigration of young (n = 2) and adult (n = 2) birds.



reached a peak of about 400 km/day) when theycrossed the Sahara Desert and decreased to a mini�mum in their nesting and wintering areas (Fig. 5). Thehighest flight speed (over 50 km/h) was recorded in thefirst half of the day (Hake et al., 2003).

An analysis of the spring migration of satellite�tagged bar�headed geese (Anser indicus) from India toNepal showed that they flew over the Himalayas at analtitude of more than 6000 m (Javed et al., 2003;Prosser et al., 2009) and successfully arrived to theirnesting grounds on the Tibetan Plateau (4450 m a.s.l.),750 km north of the wintering region.

Studies on orientation and navigation. Telemetrymethods provide the possibility of studying not onlymigrations themselves but also the capacity of migrat�ing birds for orientation and navigation. An illustrativeexample is the experiment with fledged white storksdetained on the Courish Spit in order to test thehypothesis of inherited migratory orientation in youngbirds (Chernetsov et al., 2004). On the basis of numer�

ous bird�banding data, it was hypothesized that whitestorks from eastern and western Europe follow differ�ent inherited programs of autumn migration fromtheir nesting areas to the wintering grounds in Africa:birds from eastern populations, including those ofeastern Germany, round the Mediterranean Sea onthe east, following the so�called eastern flywaythrough Turkey, Israel, and Egypt, while birds of west�ern populations (from western Germany to Spain)take the western flyway through the Strait of Gibraltar,Morocco, etc. To test whether the route of the firstautumn migration is indeed programmed genetically,it was expedient to detain young storks in the region ofbirth until the departure of all adult birds and thenrelease them and track their migration by means ofsatellite telemetry. Such an experiment was performedby Chernetsov et al. (2004) at the Rybachy BiologicalStation on the Courish Spit, the Baltic Sea. Threegroups of chicks taken from the nests in the Kalinin�grad Region were reared in an aviary, individually

60° W 0° 60° E

60° N

30° N

0°

30° S

60° S

1 2

Fig. 4. Routes of spring, (1) winter, and (2) autumn movements of arctic terns during migrations and in wintering areas accordingto logger data (from Egevang et al., 2010).

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tagged with satellite receivers, and released on Sep�tember 7, 11, and 21 (as a rule, storks depart from theregion by the beginning of September). Chicks of thecontrol group were satellite�tagged in the nest andcould migrate together with their parents. Asexpected, they flew southeast, following the route nat�ural for eastern stork populations: through Turkey,Israel, and Egypt, to Sudan (Fig. 6) The birds of exper�imental groups unexpectedly flew southwest, throughPoland and Germany. One of them reached the Med�iterranean coast of France, crossed the sea, andremained for wintering in Tunisia (Fig. 6). Storks usu�ally avoid flying over open waters, but this bird suc�cessfully covered 752 km over the Mediterranean in26 hours. The pattern of its migrations in subsequentyears is also of interest. The bird spent the second win�ter near Lake Chad in Nigeria, migrated in spring toSpain, and remained there until the next spring, whenit was recorded in Poland, 220 km away from its birth�place (on March 30, 2003). Its migration to Africa forwintering started on August 24 of the same year, butthis time the bird followed the southeastern routecommon to storks from eastern populations. Itapproached the last�year wintering site near LakeChad on September 28 but then flew east and

remained for the winter in western Sudan. In earlyMarch 2004, the bird started the return migration to itsbirth region via the traditional route (through Egypt,Israel, and Turkey) and completed it in northernPoland on April 14 (Chernetsov et al., 2005). Theauthors concluded that the choice of migration routein the first�year white storks is governed not so muchby an inherited program (which probably specifiesonly its general southward rather than northwarddirection) as by social interactions with adult bird,whom they follow during their first migration. This isconfirmed by observations on a satellite�tagged storkfrom the group detained on the Courish Spit, whichinitially migrated southwest but suddenly turnedsoutheast when flying over central Germany. Appar�ently, it met a late�migrating adult bird and followed iton the southeast route to Turkey it is due only totelemetry tracking that the experiments yielding suchvaluable data could be performed. Another experi�ment to study the magnetic orientation of birds duringmigration with the aid of short�range transmitters wassuccessfully performed on thrushes of the genusHylocichla in the United States territory (Cochranet al., 2004).

500

400

300

200

100

0

(a)M

igra

tio

n s

pee

d,

km/d

a

WestAfrica

Sahara EuropeMales

Females

1

2

3

4

5

6

500

400

300

200

100

0

(b)

Mig

rati

on

sp

eed

, km

/day

1

2

3

Youngbirds

0°N 20°N 40°N 60°N

Fig. 5. Autumn migration speed of (a) adult (n = 6) and (b) young (n = 3) European honey buzzards in Europe and Africa accord�ing to satellite telemetry data (from Hake et al., 2003).



Satellite telemetry can also be used in studies onnavigation capacities of birds. Some researchers cur�rently perform experiments on displacing birds fromthe summer migration route in order to find outwhether they can perceive this longitudinal displace�ment and correct the course of flight to their nesting(native) area (Chernetsov et al., 2008). Such experi�ments are so far conducted with passerine birds, usingthe Emlen funnel cages to reveal migratory orienta�tion. The next step will involve experiments in nature,with the birds displaced from the migration routebeing fitted with satellite transmitters to trace theentire course of their flight to the destination point.Before releasing the birds, it would be expedient toexpose them for a certain time in Helmholtz rings,which allow the direction of the magnetic field to bealtered, and to test the effect of this treatment on theirmigratory orientation. Such experiments can providea much deeper insight into the functioning of bird nav�igation mechanisms, compared to laboratory studies.In the latter case, there is always uncertainties as towhether the observed phenomena do indeed take placein nature or are artifacts resulting from bird keeping incaptivity.

Studies on dispersal and homing. Such studies datefrom the 1960s, when Godfrey and Marshall (1969)used ratio transmitters to study brood disintegrationand dispersal in the ruffed grouse (Bonasa umbellus).Two transmitter�tagged broods disintegrated on Sep�tember 7, but the young remained at the same placesfor 7 more days. Their subsequent dispersal occurredin two stages, on September 24 and October 7, in bothcases coinciding with the passage of a katafront.Despite the absence of connections between individ�ual birds, the dispersal began synchronously, and theirpreviously chaotic movements became oriented in the

same direction. The dispersing birds walked most ofthe way. The daily distance covered by a bird reached1740 m, averaging 874 m. They advanced 1.5–8.0 kmin 2–4 days, and then their movements became cha�otic again. Lance (1970) performed a similar study inthe blue grouse (Dendragapus obscurus). The resultsshowed that the tagged brood walked in the daytimestrictly in the same direction, crossing various obsta�cles without any attempts to circumvent them. Thus,the birds covered 2.5 km (on September 1–6) and kepttogether for 6 more days before dispersing. Such adetailed information on the movements of young birdsof the above species would be impossible to obtainwithout using ratio tracking.

The availability of transmitters weighting less than1 g has made it possible to study dispersal and homingin small passerine birds. Using these devices, special�ists have obtained novel information on the postnest�ing life of birds and their ability to find their individualterritories in the nesting period. For example, studiesperformed by Mukhin et al. (2005) on the Courish Spitallowed them to discover the phenomenon of noctur�nal postnesting dispersal in young reed warblers (Acro�cephalus scirpacius), blackcaps (Sylvia atricapilla),and garden warblers (S. borin). These birds (aged 30–50 days) in their native region proved to fly at night todistances of up to several kilometers. The authors con�sider that these flights are necessary for the young birdsto develop their capacity for orientation by stars beforethe autumn migration and, probably, to imprint navi�gation cues for the spring migration.

Telemetry tracking of reed warblers provided evi�dence not only for the aforementioned night flights ofthe young but also for the ability of adult birds to returnhome at night during the nesting period (Mukhin

(a) (b)

Fig. 6. Autumn migration routes of white storks from the Kaliningrad Region according to satellite telemetry data: (a) birds fromthe control group (n = 4), (b) birds from the experimental group (n = 5) (from Chernetsov et al., 2004).

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et al., 2009). Males transferred to different distancesfrom the nest (2–21 km) proved to return only atnight, after several days, since a sufficiently longperiod of time was required to develop the nocturnaltype of locomotor activity similar to that during migra�tion. The authors suppose that such a behavior hasadaptive significance for bird species inhabiting frag�mented biotopes.

Unlike reed warblers, pied flycatchers showed nocapacity for nocturnal homing in radio telemetryexperiments (Kosarev and Sokolov, 2007a). All of40 males transferred from the nest to distances of 2 to10 km returned only during the daylight period. More�over, they returned much more rapidly in sunnyweather (within a few hours) than in cloudy weather(after no less than 24 hours). Analyzing the trajectoriesof most these birds, two distinct phases of their move�ments in search of the nest were revealed. For at least3 hours after being released, the birds moved in differ�ent directions within a radius of about 0.5 km, but thenheaded straight to the nest, covering 10 km in 20–30 min. In our opinion, the birds at the first phase weretrying to find their nests near the site of release (someof them even examined empty nest boxes) and deter�mine their coordinates relative to the nesting area,which took several hours; having taken bearings totheir nests, the birds quickly returned there. It appearsthat seeing the sun is necessary for their correct orien�tation in space. Gavrilov et al. (2010) obtained similarresults in telemetry experiments on the pied flycatcherand European robin (Erithacus rubecula) at theZvenigorod Biological Station, Moscow State Univer�sity. In both species, adult birds carried away from thenest for 1–5 km proved to return home only during thedaylight period, although European robins are gener�ally more active at dusk, compared to flycatchers. As inthe previous case, they returned more rapidly in sunnythan in cloudy weather.

Studies on the timing of seasonal phenomena. It isoften necessary for research purposes to determineexact dates of the onset of migration, dispersal, orother kinds of bird movements. Conventional methodsare of little or no use in this case. Thus, still very littleis known about the dates of bird departure from win�tering grounds, although this information is veryimportant for understanding the role of recent climatechanges in the life of birds (Sokolov, 2010). The major�ity of ornithologists adhere to the traditional conceptthat migratory birds, particularly long�distancemigrants, start their spring migrations on approxi�mately the same dates from year to year (with a devia�tion of no more than 7 days from the long�term aver�age date) and that these dates are genetically specificfor each species and population (Dolnik, 1975). Thisconcept still prevails, despite that studies performed indifferent countries of Europe and North America inthe past three decades provide evidence for signifi�cantly earlier arrivals of birds to their nesting regions,compared to the previous period (Sokolov, 2010).Such a tendency has been observed in many speciesmigrating not only within a certain continent but alsobetween continents. The shift in the dates of springmigration is usually attributed to climate warming inthe Northern Hemisphere. However, some specialistsconsider that changes have occurred not so much inthe timing of bird departure from wintering grounds asin the speed of their movement along the migrationroute, although factual data in favor of this importanthypothesis are as yet scarce. Hopefully, satellite track�ing will help to resolve this issue.

We managed to analyze satellite telemetry data onthe timing of departure of white storks from their Afri�can wintering grounds, which were collected by Ger�man ornithologists over almost 20 years (Kosarev andSokolov, 2007b). In particular, 22 adult birds weremonitored during 6 years (beginning from 1992) todetermine the exact dates of the onset of their springmigration to Europe. The results showed that the aver�age date was February 20, but migration in 1997started a month later, on March 20 (Fig. 7). In thatyear, a cold spell in February was recorded in a vast ter�ritory, from the Black and Caspian seas to the Gulf ofGuinea coast, including the regions of Sahel and EastAfrica. As a result, foraging conditions for storksapparently deteriorated, and they postponed theirdeparture from wintering grounds. On this basis, weconcluded that the time of the onset of spring migra�tion in white storks largely depends on weather condi�tions in their African wintering grounds and maymarkedly vary between years (by up to 30 days), whichcontradicts the generally accepted concept of mecha�nisms regulating the development of spring migratorystate in birds. Unfortunately, there are practically noother examples of long�term direct observations onthe dates of bird departure from wintering grounds inAfrica or Central and South Americas.

Apr. 4

Mar. 29

Mar. 19

Mar. 9

Feb. 27

Feb. 17

Feb. 720031994 1997 2000

Dep

artu

re d

at

Years

Fig. 7. Dates of departure of white storks fitted with satel�lite transmitters from African wintering sites in differentyears. Each square corresponds to an individual bird (fromKosarev and Sokolov, 2007a).



Preparing this review, I have managed to findapproximately 200 information sources providing fac�tual data of bird studies by means of telemetry, includ�ing scientific journals, books, and websites. Obviously,some printed or electronic publications have escapedmy attention, but even those included in analysis pro�vide conclusive evidence for the importance and wideapplications of corresponding methods. Unfortu�nately, they are not yet widespread in Russia, which isexplained primarily by their relative expensiveness, butprogress in Russian field ornithology will be impossi�ble unless telemetry tracking is introduced on a largescale. However, telemetry is not the key to resolve allproblems but just a research method, and it is impor�tant to use it competently and have a clear�cut idea ofwhat is to be studied with its aid. When financialresources are insufficient to afford expensive satellitetransmitters, attention may be focused on short�rangemicrotransmitters, which are relatively low�priced butcan provide no less valuable data when used in studieson dispersal, homing, orientation, territorial behavior,physiology, and other aspects of bird life. Moderntelemetry offers wide opportunities for bird research.

ACKNOWLEDGMENTS

This study was supported by the Russian Founda�tion for Basic Research, project no. 10�04�00721�a.

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