9
 j. RaptorRes'. 6(4):256-264 ¸ 2002 The Raptor Research Foundation, Inc. VERNAL MIGRATION OF BALD EAGLES FROM A SOUTHERN COLORADO WINTERING AREA ALAN R. HARMATA 1 Fish & WildliJ•b rogram, epartment f Ecology, ontana St ate University, ozeman, T 59717 U.S.A. ABSTRACT.--Adult ald Eagles Haliaeetus eucocephalus) N = 15) wintering n the San Luis Valley (SLV), Colorado were radio-tagged with conventional ail-mounted transmitters between 1 January-18 March 1980 and 1981 to determine migration patterns and breeding areas. Migrating eagles were fbllowed primarily in a single vehicle with two trackers. n 1980, radio-tagged eagles N = 4) left the wintering grounds within a 15-d span n March but departures n 1981 (N = 7) ranged i:om mid-February o early April. Eagles nitiated migration on dayswith higher temperature anges,more clouds,and higher winds than other days during winter or spring. Subsequent ravel paralleled the northward movement of the 2øC sothermboth temporallyand spatially.Locations and pathways f migrating eagles were similar in both 1980 and 1981. All four eagles ocated on their summer range were within 102 km of each other in northeastern Saskatchewan and northwestern Manitoba. Mean distance fi:om the SLV wintering area to breeding or summer areas of Bald Eagles was2019 km. Adult Bald Eaglesapparently migrated alone in spring with mated males eaving first. Migration flights began between 1015-1045 H MST and ended between 1715-1745 H. Mean daily movement was 180 km. Migration flight speeds averaged about 50 km/hr. Altitude of flight ranged fi :om 30-4572 m above ground level (AGL), but most often was between 1500-3050 m. KEY WORDS: Bald Eagle;, aliaeetus eucocephalus; Colorado;adio-tracking;askatchewan; ernalmigration; wintering. MIGRACION PRIMAVERAL DE LAS XGUILAS CALVAS DESDE UN 3.REA DE INVERNACION AL SUR DE COLORADO RESUMEN.--Individuosdultos de figuila calva (Haliaeetus eucocephalus) N = 15) invernando en el valle de San Luis (VSL) Colorado, fheron provistos con r adios transmisores onvencionales montadosen la cola, entre el 1 de enero-18 de marzo de 1980 y 1981 para determinar os patrones de migracitn y las fireas de reproduccitn. Las •tguilas n migracitn fheron seguidas n primera instancia en un vehiculo sencillo con dos rastreadores. En ]980, las figuilas con radios (N = 4) abandonaron los campos de invernacitn en un intervalo de 15 dias en marzo, pero las partidas en 1981 (N = 7) fueron entre mediados de fEbrero y principios de abril. Las •tguilas niciaron la migracitn en dias con rangos de temperaturasmas altos,mas nubes,y vientos mas altos que otros dias durante el invierno o primavera. E1 subsiguiente uelo fue paralelo al movimiento hacia el norte de la isoterma 2øC anto temporal como espacialmente. Las localizaciones vias de paso de las figuilas migratorias heron similares en 1980 y 1981. Las cuatro •tguilas ocalizadas n su rango de verano estuvieron dentro de 102 km una de otra en el nororiente de Saskatchewan noroccidente de Manitoba. La distancia media desde el firea de invernacitn del VSL alas fireas de reproduccitn o de verano de las figuilas calvas he de 2019 km. Las figuilascalvasadultasaparentementemigraron solasen pri•navera mientras que sus machospareja sal- ieron primero.Los vuelos e migraci6n omenzaron ntre 1015-1045 H MST y terminaron entre 1715- 1745 H . E1 movimiento medio diario rue 180 kin. Las velocidades e los vuelos durante a migraci6n tuvieronun promedio de 50 km/h. La altitud de vuelo tuvo un rango entre 30-4572 m.s.n.m. pero la mayoria a menudo estuvo entre 1500-3050 m. [Traducci6n de C6sar Marquez] Relatively stable wintering populations of Bald Eagles (Haliaeetus eucocephalus ) xisted in the San Luis Valley (SLV) of southern Colorado through- E-mail address: [email protected] out the DDT era (Ryder 1965, Alamosa National Wildlif• Refuge reports 1954-83). In the 1970s, a mean of 185 (SD = 66.6) Bald Eagles was counted in the SLV annually (N = 5; Craig 19 81). However, origins of eagleswintering in the SLV were un- known. 256

Harmata 2002 Vernal Migration of Bald Eagles From Southern CO Wintering Area

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j. RaptorRes'. 36(4):256-264 2002 The Raptor ResearchFoundation, Inc.

VERNAL

MIGRATION OF BALD EAGLES COLORADO WINTERINGALAN R. HARMATA 1

FROM AREA

A SOUTHERN

Fish & WildliJb Program, Department Ecology, of Montana StateUniversity, Bozeman, 59717 U.S.A. MTABSTRACT.--Adult Bald Eagles(Haliaeetus leucocephalus) = 15) winteringin the San Luis Valley (SLV), (N Colorado were radio-taggedwith conventionaltail-mounted transmitters between 1 January-18 March 1980 and 1981 to determine migration patterns and breeding areas. Migrating eagleswere fbllowed primarily in a singlevehicle with two trackers.In 1980, radio-taggedeagles(N = 4) left the wintering

groundswithin a 15-d spanin March but departures 1981 (N = 7) rangedfi:ommid-February in to earlyApril. Eaglesinitiated migrationon dayswith higher temperatureranges, more clouds,and higherwinds than other daysduring winter or spring. Subsequenttravel paralleled the northward movement of the 2Cisothermboth temporallyand spatially. Locationsand pathways migratingeagleswere of similar in both 1980 and 1981. All four eagleslocated on their summer range were within 102 km ofeach other in northeastern Saskatchewan and northwestern Manitoba. Mean distance fi:om the SLV

wintering area to breeding or summerareasof Bald Eagleswas2019 km. Adult Bald Eaglesapparently migrated alone in springwith mated males leavingfirst. Migration flightsbegan between 1015-1045 H MST and ended between1715-1745 H. Mean daily movementwas 180 km. Migration flight speeds averagedabout 50 km/hr. Altitude of flight ranged fi:om 30-4572 m aboveground level (AGL), butmost often was between 1500-3050 m.

KEYWORDS: BaldEagle;, Haliaeetus leucocephalus;Colorado; radio-tracking; Saskatchewan; vernalmigration; wintering. MIGRACION PRIMAVERAL DE LAS XGUILAS CALVAS DESDE UN 3.REA DE INVERNACION ALSUR DE COLORADO

RESUMEN.--Individuos adultosde figuila calva (Haliaeetus leucocephalus) = 15) invernando en el valle (N de San Luis (VSL) Colorado, fheron provistos con radios transmisores convencionales montadosen la cola, entre el 1 de enero-18 de marzo de 1980 y 1981 para determinarlos patronesde migracitn y las fireasde reproduccitn. Las tguilas migracitn fheron seguidas primera instanciaen un vehiculo en en sencillo con dos rastreadores. En ]980, las figuilascon radios (N = 4) abandonaron los camposde

invernacitn en un intervalode 15 dias en marzo, pero las partidasen 1981 (N = 7) fueron entremediados de fEbrero y principios de abril. Las tguilasiniciaron la migracitn en dias con rangosde temperaturasmas altos,mas nubes,y vientosmas altos que otros dias durante el invierno o primavera. E1subsiguiente vuelofue paraleloal movimientohaciael norte de la isoterma2Ctanto temporalcomo espacialmente.Las localizaciones vias de paso de las figuilasmigratoriasfheron similaresen 1980 y y 1981. Las cuatro tguilaslocalizadas su rango de verano estuvierondentro de 102 km una de otra cn en el nororiente de Saskatchewan noroccidente de Manitoba. La distanciamedia desde el firea de y invernacitn del VSL alas fireasde reproduccitn o de verano de las figuilascalvas fhe de 2019 km. Las figuilascalvasadultasaparentementemigraron solasen prinaveramientrasque susmachospareja sal-

ieron primero.Losvuelos migraci6n de comenzaron entre 1015-1045H MST y terminaronentre 17151745 H. E1 movimientomedio diario rue 180 kin. Las velocidades losvuelosdurante la migraci6n de tuvieronun promediode 50 km/h. La altitud de vuelo tuvoun rango entre 30-4572 m.s.n.m.pero lamayoria a menudo estuvoentre 1500-3050 m.

[Traducci6n de C6sarMarquez]

Relatively stable wintering populations of Bald Eagles (Haliaeetusleucocephalus) existed in the San Luis Valley (SLV) of southern Colorado throughE-mail address: [email protected]

out the DDT era (Ryder 1965, AlamosaNational Wildlif Refuge reports 1954-83). In the 1970s,a mean of 185 (SD = 66.6) Bald Eagleswascountedin the SLV annually (N = 5; Craig 1981). However, origins of eagles wintering in the SLV were unknown.

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BALr> EAGLE VERNAL MIGRATION

257

Between 7January-18 March 1980 and 1981, 15 adult Between December-April 1977 and 1978, winBald Eagleswere captured and radio-taggedin the SLV tering Bald Eagles (N = 36) in the SLV were All were captured by a modified "Lockhart" method marked with yellow patagial wing markers to de- (Miner 1975) with and without live Bald Eagle and Goldtermine their geographicorigins or breeding areas en Eagle (Aquila chrysaetos) birds. Trap siteswere lure (Harmata and Stahlecker1993). Colormarking re- chosen on the basis of frequency and duration of the vealed fidelity of individual Bald Eaglesto this win- presenceof two adult Bald Eaglesof distinctlydissimilar size, presumablymated, within 1.6 km. In 1981, I estitering area was high, but by January 1981 only mated the SI.V winter population at 170 Bald Eagles three sightingsof yellow-markedeaglesoccurred (Harmata 1984).outside the SLV and none were in a documented

Mated status of radio-tagged Bald Eagles was determined by frequency and duration of time spent in the 1978, after being tracked locally for nearly six presence of another adult eagle of distinctly dissimilar weeks,an adult Bald Eagle wearing a conventional, size,observedcopulation (N = 3) in the SLV, or association with a nest site on the breeding grounds (N = 2). tail-mounted transmitter left her SLV wintering Eagles were considered unmated if diurnal movements area. She was tracked for two days over 300 km in the SLV were clearly independent of other eaglesand through the most rugged portion of the Rocky they were not observed to roost away from communal Mountains, being lost only due to lack of logistical roostswith just one other eagle, asmated birds often did Climatologicaldata associated with daysthat Bald Eaplanning, not our ability to maintain contact. This gles left the SLV were analyzedby stepwise discriminate serendipitousevent revealed the potential of de- analysis(Dixon 1981) to investigatemeteorologicalcontermining migration routes and breeding areas of ditions associated with initiation of vernal migration. ChBald Eagles usingconventional radio-tracking. This matologicaldata recorded near the geographical center

nesting area or during summer (Harmata and Stahlecker 1993). The primary objective of the marking program had not been realized. Clearly, other methodswere required to achieveobjectives in a timely and cost-efficient manner. Prior to the 1980s, technology for remotely tracking long-range movementsof individual birds (i.e., satellite platform transmitter terminals or PTTs) was not available. However, on 28 February

Gender of radio-tagged eagles assigned methods was by presented by Garcelon et al. (1985). Three were confirmed by behavior during copulation. All Bald Eagles were radio-tagged with two-stage radio transmitters mounted proximo-ventrallyon the tail. Transmitter frequencieswere between 148.500 and 148.950 mHz. Unit life expectancy was -->5 mo. Transmitter, antenna, and mounting tab weighed 50-57 g. Telemetry receiving equipment included fixed channel and programmablereceivers.

paper discusses results of subsequentlong-range tracking of Bald Eagleswith conventional telemetry from one seasonalrange to another prior to widespread use of PTTs on eagles (e.g., Grubb et al. 1994, Brodeur et al. 1996, Meyburg et al. 2001). Objectivesof this study were: (1) to determine breeding areas of adult Bald Eagles wintering in southern Colorado and (2) to gather information regarding factors associated with initiation of vernal migration, routes, duration, stopover habitats used, and other factors affecting the successful completion of migration.

of the SLV (Alamosa, Colorado) were obtained from Na-

tional Oceanographicand AtmosphericAdministration, Asheville,North Carolina Monthly SummarySheets. Data for daysthat eaglesleft the SLV were compared to data for daysrandomly selectedbetween I January-15 April 1980 and 1981 that they did not. Variables selected for comparisons among days were maximum, mean, and range of temperature, percent of clear sky and mean wind speed. Migrating eagleswere followed primarily from a single 4 X 4 vehicle with two human trackers and a dog. An omnidirectional antenna and two element "H" yagi receiving antenna were mounted on the roof of the chase vehicle.The yagiwasattached a 360traversing to mount, allowing for directional tracking while the vehicle was moving.Manpowerand logistic limitationsplusvariability STUDY AREA AND METHODS in departure dates,routes, and travel speedsof Bald EaThe SLV is the largestand most southernof four large gles prevented ground tracking of more than one eagle intermountain basinsin Colorado. Encompassing 6475 at a time. One tracker drove while the other operated km9, the SLV is approximately the size of the state of the receiving equipment. Both shared navigational duDelaware.Mean elevationof the nearly-level valleyfloor ties. Due to often high chasespeeds(up to 150 km/hr) is 2286 masl. High (>3050 m) mountain ranges border and off-road "adventures,"visualcontactwith migrating the valley on eastand west,merging at the northern end. eagles could not be maintained, so migration behavior The Rio Grande and Conejosriversflow through the SLV often could not be recordedcontinuously, accurately, or and numerousnatural warm springsand wetlandsthat safely.Route, direction, and speed of the chasevehicle, seldom freeze have made the SLV attractive to waterfowl therefore, often was selectedprimarily to maintain maxand Bald Eagles,probablyfor millennia. Water develop- imum audio signal strength. When contact with a mentsand agriculturein the 20th centuryhaveprobably grating eagle waslost, an aerial searchwasimplemented improved the attractiveness. using local air services. two or three element yagi anA

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FEMALES

O

O

o

MALES

O

O

i

....

I

....

i

....

I

....

i

....

I

....

i

....

I

....

I

....

I

....

i

....

-40

-35

-30

-25

-20

-15

-10

-5

0

5

10

15

DAYS FROM VERNAL EQUINOX

Figure 1. Daysfrom vernal equinox that radio-tagged Bald Eaglesdeparted (N = 12) their San LuisValley,Colorado, winter rangesto initiate vernal migration,1978 (N = 1), 1980 (N = 4), 1981 (N = 7). Open circlesindicateeagles determined to be unmated, closedcirclesindicate eaglesthat were mated (see text), and squareindicateseagle ofundetermined mated status.

tenna was taped to the wing strut (high wing) or step (low wing) of light aircraft and the area surveyedin transectstylewith intermittent, "lazy" circlesat high altitude. Frequencieswere scanned to searchfor the target eagle and other potential migrating birds. When contactwas reestablishedand eagle's statusdiscerned as stationary, vehicle trackingresumed.Flight altitudeswere estimated based on eagle's position relative to the search aircraft and trees, geograph)5and structuressuch as buildings and radio towers while ground tracking. Eagles were monitored in their summering groundswith Beaverair'craft, snow machines, and snow shoes.

RESULTS AND DISCUSSION

Initiation of Migration. Over half (58%) of radio-taggedBald Eagleswith known departure dates (N = 12), initiated vernal migration within one week after the vernal equinox (Fig. 1). In 1980, radio-tagged eagles (N = 4) all left the wintering grounds within a 15-d span in March (Table 1). Range of known departure dates (N = 8) spanned 50 d in 1981,with the first confirmeddepartureof

Table 1. Residencyand mated statusof radio-taggedadult Bald Eagleswith known departure dates from winter rangesin the San Luis Valley (SLV), Colorado.MINIMUMRESIDENCY

EAGLEDESIGNATION SEX CAPTURED

DATEDEPARTED SLV

MATEDS[XI'US '

INSLV(DAYS)

378 180 280 380 480 181281 381

F F M M M FM F

21Jan. 1978 18Jan. 1980 23Jan. 1980 23Jan. 1980 27 Jan. 1980 7Jan. 19819Jan. 1981 9 Jan. 1981

28 Feb. 1978 21Man 1980 24Man 1980b 21Man 1980 8Man 1980 11 Feb. 198120 Feb. 1981 27Man 1981

U M M M U UU M M U?'

39 63 61 58 41 3643 78

481 581881

M FM

11 Jan. 1981 15 Jan. 198115Man 1981

27Man 1981 25Man 198121 Man 1981

76 707

981

F

18Man 1981

1Apn 1981

M

15

M = mated, U = unmated (see text).

First departed on 22 March 1980, but eagle encountereda winter slorm and returned to winter range.Undetermined.

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BALD EAGLE VERNAI. MIGRATION

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Table 2. Mean climatological values(+99% confidence interval) which discriminated (P-< 0.01) between days radio-taggedBald Eaglesinitiated northward migration from their San Luis Valley, Colorado wintering area (Depart) and daysthey did not (Remain).TEMPERATURE PERCENT WIND

southerly winds; similar to conditions noted by Bagg et al. (1950) for other birds. Although Bald Eaglesare apparentlysensitive to local conditions when migration is initiated, they appear to be cognizant of little beyond their mmediate environs.Eagle 280 departed the SLV on22 March 1980 and traveled 145 km north before

RANGE

SKY

SPEED

(C)

CLE

(km/hr)

Depart (N = 11)Remain (N = 21) a' Randomlyselected.

24 (5)19 (4)

50 (26)56 (20)

22 (5)16 (5)

being stoppedby severesnowsqualls.He spentthe night on a mountain passand as the storm persistedto the north the next day, he returned to his winter range in the valley.He spent the remainder of that day and part of the next in closeassociation with a female (distinctlylarger eagle) and was observed to copulate during this period. He initiated migration again on 24 March, leaving the SLV forthe season.

a radio-tagged eagle occurring in mid-February and the last in early April (Table 1). Number of daysfrom the equinox that eaglesdepartedwasnot different between genders (Mann-Whitney U = 16.50, P = 0.81). However, mated eaglesdeparted the SLV closer to the equinox and later than unmated eagles (Mann-Whitney U = 3.00, P = 0.03; Fig. 1). Three climatological variablesdiscriminated between days that radio-tagged Bald Eagles left the SLV on migration and those they did not (Table 2). Eaglesinitiated migration on dayswith much larger range of temperatures, more clouds, and with higher winds than other daysover the winterspring period. Migration tended to be initiated about 5 hr after sunrise regardlessof the immediate meteorological condition. Photoperiodismis consideredto be the ultimate stimulusfor the onset of migration in birds (King and Farner 1963). All radio-taggedBald Eaglesdetermined to be mated left the SLV within 12 days of the vernal equinox, indicating breeding adults may be sensitive equal periodsof light and dark. to Unmated, nonbreeding, or immature eagles may be equally sensitivebut may not be driven by pressure to procreate. In fact, first observations subof adult eagles in northern ranges both years were not until at least two weeks subsequentto the arrival of the first radio-tagged adults. Migrational movement also paralleled the northward movement of the 2C isotherm (Lincoln 1979) both temporallyand spatially, hinting that thermal cues may also be involved in the initiation of migration. Once the urge is kindled, proximate affectors of Bald Eaglemigration appear to be coincidentwith incoming low pressure systems,associatedwide range of temperatures, cyclonic air flows, and

Migration Routes, Destination, and Navigation. In 1980, a relatively narrow migration corridor through Colorado, Wyoming, and Montana was used by three radio-tagged eagles (Fig. 2). Eagle 380 became sedentary in north-central Saskatchewan after 15 d of migration, six of which were spent sitting out bad weather. After four days of almost continual solitary soaring, he was fbund perched close to a larger eagle where he remained fbr severalhours. The next day the pair began construction of a new nest on Chachukew Lake, north

of Pelican Narrows, Saskatchewan. They eventually fledged one young in August 1980. In 1981, eagle 281 left the SLV on 20 February, and wasfollowed for two daysover 302 km through the mountains of western Colorado (Fig. 3). Direction was primarily northwest and 281 was the only eagle tracked that crossedthe Continental Divide. Subsequent tracking of 281 was interrupted becausea radio of the samefrequencyand pulserate as 281 was tracked for a day before I discoveredit was a transmitter on a collar of a bighorn sheep( Ovis canadensis).

Eagle 981 left the SLV on 1 April and was followed to about 65 km north of Casper,Wyoming,where she was lost in a snow storm because roads

became impassable. Eagle 481 was detected approaching Fort Peck Reservoir in east-centralMontana on 5 April. Afier leaving Fort Peck Reservoir, 481 flew due north about 35 km then graduallyshifted to a northwest course. This route took him

directly to the Missouri Coteau, a ridge line running longitudinally for over 150 km in southwestern Saskatchewan. continued north along the He Coteau for about 100 kin. Approaching the Canadian Shield, 481 made a sudden change in course

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Figure 2. Partial migrationroutesof radio-tagged adult Bald Eaglesfrom winter rangesin the San Luis Valley, Colorado in 1978 and 1980. Numbers indicate eagle designation and year (Table 1). Summeringarea of eagle 380 is indicated by () (confirmed nest location).

from north to northeast, paralleling the direction of many elongatedlakesand riversin the Canadian Shield country. Eagle 481 was subsequentlyfollowed to Reindeer Lake, Saskatchewan, arriving on malfunction, a dead eagle, or a bird in incubation 10 April. On 22 April, 481 was found by air asso- posture.Lack of contactwith other SLV eaglesmayciated with a nest site on Kamuchawie Lake in west-

Figure 3. Partial migration routesof radio-taggedadult Bald Eagles from winter ranges in the San Luis Valley (SLV), Colorado in 1981. Numbers indicate eagle designation (Table 1). Summeringareasare indicated by () (nest locations). Respective1981 and 1982 recovery locations BaldEagles of bandedin the SLV (January1977) are indicatedby (). The October 1981 recoverylocation of a Bald Eagle radio-tagged the SLV in March 1981 in is indicated by 781.

have been a function of transmitter failure (one

ern Manitoba near the Saskatchewanborder (Fig.3).

wasknown to fail in the SLV), premature sheddingof the transmitter (one transmitter was found still

Aerial surveys covering 210000 km2 of north- attached to broken tail feathersbelow a perch tree ca.ern Saskatchewan were conducted to detect signals of other SLV Bald Eagles in mid- to late April in both 1980 and 1981. During aerial surveys, eagle 181 waslocated on the Pagato River, south of Reindeer Lake, Saskatchewan, and eagle 581 was detected briefly in the area of Trade Lake, south of the Churchill River (Fig. 3). The signal was weakand intermittent, which indicated either a radio

in the SLV), mortality, incomplete surveycoverage, or summeringareaswere located outsideof northern Saskatchewan. However, the fact that 31% of

potentially-detectable eagles(N = 13) were located within an area roughly the size of Yellowstone National Park (ca. 9300 km2) suggests mostadult that Bald Eagleswintering in the SLV originated fromthis area of Canada.

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Mean distance (as the eagle flies) from the SLV wintering area to breeding or summer areas of Bald Eagleswas 2019 km (N = 4, SD = 50). All eagleslocated on their summer rangeswere within102 km of each other in northeastern Saskatche-

the remainder of migration clearly compensated for the one day blown off course (Fig. 2). Unless the eagle had been exposed to the area on previous migrations, true navigation is indicated. However, it is not unreasonable to assume that an adult

eagle over five years old, may indeed be familiar with a great portion of western North America as a result of vagariesof previousmigrations. Regardless,redundancy in navigational systems has been illustrated for homing pigeons (Columba livza) (Able 1980) and in all probability, severalbackup navigational systems are available to Bald Eagles, especiallyexperienced adults. ed the journey in only about 40% of the time. Migration Behavior.Mated adult Bald EaglesapHowever,the Arizona eaglewasa subadult (3rd yr) parently migrated alone in the spring. Eagle 380 and presumablywas not driven by the impetus to was seen roosting solitarilyat all but one stopover nest, as were adult SLV eaglestracked. location (six other eagleson the Yellowstone River Geography,celestialcues, and weather may all near Hysham, Montana). Eagle 481 did not miplay a role during migration of Bald Eagles.Prom- grate with a mate, but often moved northward with inent physiographic featuressuchasdeep canyons, other adult eagles.He roosted with other eagles rivers, and north-south oriented topography three times, but two sitesobviously were not comseemed to assistin visual navigation during flight munal or traditional because of lack of similarity (Griffin 1943). These features could have been im- to typical Bald Eagle roost sites (Kiester and Anprinted in the memory of eaglesduring their ini- thony 1983). Paired roostingwas probably a result tial migrations and experience dictated direction of facilitatory behaviorinfluenced by poor weather during subsequent flights (kinesis theory; Mat- and lack of daylightremaining. Subsequent obserthews 1963). Imprinting of migration routes from vations of migration flight confirmed he moved wintering areas,which compliment survivalduring alone. Once in the boreal forest of the Canadian the first year, would be more adaptive than im- Shield country, local eagles appeared to "meet" him in flight and escort him through their terriprinting during first southwardmigration. Eaglesdid not migrate on daysof total overcast, tories, but no overt agonistic encounters were obwith other eagles a phenomenonalsonoted by Gerrard and Gerrard served. Occasional associations (1982). The altitude of the cloud layer may have appeared incidental for all radio-taggedeaglesand an effect, but overcastlayers over 90 m were not were of short duration. Solitary migration behavior experienced during tracking. Sun compassorien- in established pairswould reduce the possibility of tation (Kramer 1952, 1957) with time compensa- both members being lost in a local catastrophe. tion, commonly referred to as sun-azimuthorienDaily migration flights consistentlybegan beH MST and ended between tation (Welty 1982) may, therefore, be important tween 1015-1045 for orientation of adult Bald Eaglesduring migra- 1715-1745 H. Mean daily movementwas 180 km, tion. but ranged from 144-435 km (N = 5) in 1980 and True navigation (selection of a compassdirec- 33-248 km (N = 5) in 1981. Speed of migration tion toward a known goal in unfamiliar territory; flights recorded averaged50 km/hr (N = 7, 22Able 1980) may alsobe a component of Bald Eagle 144 km/hr in 1980; N = 9, 20-105 km/hr in 1981). migration. Migrating eaglesappear to avoid strong Altitude of flights recorded ranged from 30-4572 winds during migration because strong winds ap- m AGL, but most often was between 1500-3050 m. parently influence direction of flight. Eagles did Total distances,speed, and daily duration of minot move, except locally, during dayswhen winds gratory flights indicate that under optimal weather in excessof 35 km/hr occurred prior to 0900 H. conditions, Bald Eagles can reach their breeding Eastwindsof 60-80 km/hr began about two hours grounds within 6 d after leaving the SLV. Pennyafter initiation of eagle 380's flight on one migra- cuick (1975) indicated a 2000 km migration for a tion day and his flight path deviatedwell west of bird the size of a Bald Eagle would be near maxidirect line to the eventualgoal. Flight direction for mum attainable without eating, assuminga 25%

wan or northwestern Manitoba (Figs.2 and 3). Duration of travel betweenwinter and summer range for SLV Bald Eagleswith known departure and arrival dates (N = 2) was 15 d. Although SLV Bald Eaglestraveled about 2,5 the distanceto their summer ranges compared to one Bald Eagle tracked by satellite from Arizona to the Northwest Territories (3032 km, Grubb et al. 1994), they complet-

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massloss.During this study,no radio-taggedeagles were observedfeeding during migration and mean migration distance was 2020 km. Captive eagles commonly fast more than two weekswith no apparent deleterious effects (Brown and Areadon 1968, pets. observ.) and wild raptors can lose up to 30% of body masswithout problems (Newton 1979). These compensatory capacities undoubtedly allowed adult Bald Eaglesto reach their breeding grounds with sufficient energy reserve for breeding. All radio-tagged eaglesarrived in their summer range at a time when lakesand most stretches of rivers were still frozen. The only areasof open water were rapids or narrows on rivers or between lakes.Radio-tagged eaglesspentmost of their time there, presumablyforaging for fish. Other eagles observedduring aerial surveys were associated with ubiquitousholesin lake ice and viscera piles of fish left by native commercial fishing operations. Foraging eagles were also seen on or near caribou (Rangifertarandus)and moose (Alces alces) carcasses killed by natives or wolves (Canis lupus).A few eagleswere seen with snowshoe hare (Lepusamercanus) remains.

Stopoverareas used during vernal migration of SLV Bald Eagleswere generallywidely distributed. A tree of adequate size, secure from human disturbance in any type habitat, was all that seemed necessary roosting. In late March 1982, Swenfor son (1983) counted 232 Bald Eaglesalong the YellowstoneRiver between the mouth of the Bighorn River and Miles City, Montana. A site where eagle380 roosted is in the middle of this stretch and

within an area where the river seldom freezes, con-

tained the most highly braided portion of channel, most heavilywooded islands,and highest Canada goose (Branta canadensis) populations of three sections of river studied by Hinz (1974). Use of this sectionof the Yellowstone River by adultsmay be dictated primarily by tradition and availabilityof water, becausemigrating adults were not known to feed during this study. Areas in eastern Montana may be equally important to asmany or more migrating eaglesasthe more highly-publicized areas, where ephemeral concentrations of eagles occur in western Montana.Leightonet al. (1979) estimated population a of 14000 Bald Eagles in Saskatchewan. Some eagles from north-central Saskatchewan were captured in autumn at Hauser Lake (Restani et al. 2000) and Glacier National Park (McClelland et al. 1982), while others passedthrough eastern Mon-

Regional Relationships. McClelland et al. (1994) noted that Bald Eaglesradio-taggedin Glacier National Park, Montana, in autumn wintered west ofthe Continental Divide and summered in the

tana during migration (Gerrard et al. 1978, Harmata et al. 1985). Both Hauser Lake and Glacier

MacKenzie River watershed of northern Alberta, northwest Saskatchewan, and Northwest Territo-

ries. The summer range of a Bald Eagle that wintered in Arizona (west of the Divide) also was in the MacKenzie River watershed (Grubb et al.

1994). McClelland et al. (1994) suggested that wintering areas may be related to the watershed of origin and Bald Eaglesbe managed by application of a "Migration FlywayConcept." Adult Bald Eaglesradio-taggedand banded in the SLV wintering area (east of the Divide) were tracked to breedingareas in Saskatchewan and Manitoba, all in the

National Park Bald Eagle concentrations now are defunct due to a collapsed, exotic food base (kokanee salmon, Oncorhynchus nerka), but eastern Montana habitatsstill support large numbersof native prey (lagomorphs,ungulates,waterfowl). However, lack of a concentrated food base, diffusion of

Churchill River watershed. Jenkins et al. (1982) followed two adult Bald Eaglesradio-taggedin Wyoming during winter. One trapped on the west sideof the the east Continental side of the Divide Divide was fbllowed to to the the

MacKenzie River watershed,while one trapped onwas followed

Churchill

River watershed,

similar to those from

the SLV. These data suggesta "Churchill-East Slope" Migration Flywayexists,distinct from the "Mackenzie-Intermountain" Flyway proposed byMcClelland et al. (1994).

roost sites,solitary habits of migrating eagles,plus dispersion of departure dates from winter (this study) and summer ranges (Harmata et al. 1985), prohibit any accurateestimateof numbersof Bald Eaglespassingthrough easternMontana. Relatively low numbers of eagles present at any particular time at some stopover areas in eastern Montana may belie the true importance of these areas to migrating eagles.Turnover of individualsappeared to be daily, over months. Therefore, western prairie statesmay provide important migratory habitat for a large proportion of the continental population of Bald Eaglesover long periods.ACKNOWLEDGMEN

J. Stoddart initially suggestedBald Eagle research in the SI,V. Dale Stahleckerservedas co-investigator during

DECEMBER 2002

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263

preliminary marking studies.P. Harmata (then age 5) Bald Eagles Haliaeetusleucocephalus fi'om Arizona and helped track the first migrating Bald Eagle in 1978. G. Michigau with satellite telemetry. Pages 347-358 zn Montopoli wasinstrumental for successthl completion of B.-U. Mcyburg and R.D. Chancellor fEDS.I, Raptor the first year's migration tracking. M. Lockhart provided conservationtoday. World Working Group on Birds details of eagle capture techniques and field assistance. of Prey. Pica Press,London, U.K. S. Werner and E. Spettigue participated in winter and A.R. 1984. Bald Eaglesof the San Luis Valley, migration tracking.L. Stevenson PelicanNarrows,Sas- It^RMATA, of Colorado; their winter ecologyand spring migrauon. katchewan, donated flight expertise and air time. FinanPh.D. dissertation, Montana State University, Bozecial assistance was provided by R. Koteen; Fred Jcnse, man, MT U.S.A. Dept. of Veteran'sAffairs;J. Lincer and W. Clark of the National Wildlifk Federation; R. Phmkctt of the National --, .i.E. TOEPFER, ANDJ.M. GERP, Am). 1985. Fall miAudubon Society;D. Flath of Montana Fish,Wildlilk, and gration of Bald Eagles produced in northern SasParks; I,. Jahn of the Wildlife Management Institute and katchewan. BlueJay43:56-62. American PetroleumInstitute;T. Ingram of EagleValley --ND D.W. STAHLECI4ER. 1993. Fidelity of Bald Environmentalists; Rainey and C. Merrit of American S. Eagles to wintering grounds in southern Colorado Wilderness Alliance; M. Malone, Montana State Univerand northern New Mexico. J. Field Ornithol. 64:129sity (MSU) Research CreativityProgram; R. Eng, MSU Agricultural Experiment Station;and R. Moore, MSU Biology Department. T. Grubb, M. Raune, M. Rcstani,and D. Stahlccker made helpful comments on earlier drafts of the manuscript."Sarge" wasthe dog, a great companion and a loyal friend.134.

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Received21 December 2001; accepted6 July 2002