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Home , Haines Junction, Kenai Peninsula, Manchuria, Sheenjek River

July]19611 327

THE MIGRATION OF LAPLAND LONGSPURS TO

LAURENCE IRVING

E^RL¾in May Lapland Longspurs,Calcarius lapponicus• arrive at Anaktuvuk Passin the centerof the BrooksRange of Alaska in large flocksthat fly swiftly northward in compactformations or stop to feed among the hummocksof sedgethat are exposedthrough the snow (Irving, 1960a). Judgingfrom my observations,verbal reports, and compilationsby Gabrielsonand Lincoln (1959), theselongspurs are generallythe commonestnesting species of the Alaskanarctic tundra. They also nest in westernAlaska over the grassyland beyondthe forestsas far southas the AlaskaPeninsula, on St. Lawrence(Fay and Cade, 1959) and other islandsin the Bering (Preble and Atee, 1923), and alongthe AleutianIslands to Attu (Murie, 1959). Publishedreports (Friedmmm,1935) showthe occurrenceof LaplandLong- spurson Kodiak. Unpublishedreports (Fish and Wildlife Service) indicatethat they are commonsummer residentsin the Kodiak , which comprisesa large part of . The extent of their presenceon Kenai Peninsulais not clearly shownin publications.Comments of residentson the northwesternshore indicatethat longspurspass that part of Kenai in migra- tion. Nestingprobably occurs on Kodiak and possiblyon Kenai, but I have not been able to verify this. On Middleton Island in the (Rausch, 1958), the nestingof longspurs is detached from their arctic and western Alaskan nesting grounds. They have also been reported nesting in the interior of Alaska at Mt. McKinley National Park (Dixon, 1938); and, accordingto Gabrielsonand Lincoln (1959), George Schaller found them nesting at Black Lake in the Talkeetna Mountains. Except for the reported nesting of longspurs in these mountainous areas of the interior, the nesting populationof the mainland of Alaska is settled in a peripheral band about its arctic and western perimeter (Figure 1). South of Barrow this heavily populated band extends from the coast about 320 kilometers (200 miles) inland to the forest. Elsewhere on the malnland the densely occupiednesting area is no wider, but it extends to the islands in the , southwestwardalong the and for some 2,200 kilometers (1,400 miles) into the Pacific over the . Since longspurs are the commonest birds nesting over this great area, they are one of the most numerousarian populatlons in northwestern America.

The breedingLapland Longspursof Alaska are ascribedto the sub- speciesC. 1.alascensis Ridgway (A.O.U. Check-list,1957) and are said to winter in interior western states and as as southern British Columbia(Munro and Cowan,1947). Taking latitude 50ø and longi- tude 134ø as referencepoints for the northwesternpart of this wintering area, their migrationwould extend 20 ø of latitudenorthward and 14ø of longitudewestward to the eastern nestinglimits of the race in 328 IRw•,'(;,Migration of LaplandLongspurs [[Vol. Auk78

160ø 150ø 140e 130e 120• 70 e

60*

160• 150• 140 ß

Figure l. Northwesternrange of Calcariuslapponicus (diagonal hatched) and location of well-recorded directional migration in spring (heavy arrows).

3,lackenzie(A.O.U. Check-list,1957). In the westernmostAleutian Islandsthe migrationwould extend their summerrange only 4 ø north- ward but 67ø westward. During severalyears I have been gathering evidencefor the courseand programof this extendedand large migra- tion in Alaska.

MIGRATION THROUGH THE BROOKS RANGE

Froin 1948 to 1960 Simon Paneak, a resident of Anaktuvuk Pass (latitude 68ø 19' N., longitude 151ø 26' W. in the central part of the Brooks Range), recordedfor me his observationsof birds and thoseof his family and neighbors. In 1949 ThomasBrower kept recordsthere during April, May, and June. Theseobservations are supportedby many specimens.I have beenat Anaktuvukduring all or part of five spring migrations. In 10 years of suitablerecords the first Lapland Longspurswere recordedat Anaktuvuk on 1 May 1949, 29 April 1951, 9 May 1952,4 May 1953,13 May 1954,5 May 1956,8 May 1957,3 May 19'58,11 May 1959, and 29 April 1960; the averagedate of these ar- rivals being 5 May. Julyl1961] IRWN½,Migration of LaphindLongspurs 329

The dates of first recordedarrivals of longspursare more variable than for many arctic species.The daily recordsof migrationsduring eight yearsare completeenough to showthat after their first arrival on the averagedate of 5 May only a few longspurs,usually identified as males,were seenbefore the averagedate of 17 May (11 to 20 May). On that averagedate flocksof 40 or "many" first appeared.Thereafter, for 10 days,numbers recorded by Tom Browerwere as largeas 2,000 a day in flocksthat commonlyincluded from 50 to 150 longspurs. Until about 20 May the flocks included mainly males, but thereafter the proportionof femalesincreased. The less conspicuousappearance and behaviorof femalelongspurs may have biasedreports, but we have not noticeda late excessof femalesthat would offset the obvionsearly preponderanceof males. Like pipits, ,4nthus spinoletta (Irving, 1960b), and several other species,the first male longspursarrive at Anaktuvuk fat and with partially developedtestes (Irving, 1960a, p. 123). After 15 May the testeshad reachedfull breedingsize, and flight songsbegan to be noticed among coherent flocks that were apparently still migrating northward.Pairing was recordedas early as 20 May, nest buildinga few days later, and first eggsjust before1 June in most years. In the confusionof so many migrating flocksI did not discernwhether entire flocks separated and settled at Anaktuvuk or whether individuals droppedout as they paired and nestedthere while the remainderof the flocks continued northward. The great northwardmigration of longspursthrough Anaktuvuk con- tinues for about 10 days after the averagedate of 17 May and inter- mittently for a few more days. The large numbersthat remain to nest are an insignificantpart of the many flocksthat passswiftly northward or occasionallyalight to feedand drink busilyas if in hasteto continue. Occasionallya flock has beenwatched while groundedfor severalhours, but I have not searchedto find if the migrants roost overnight in the valley. Sometimesa flock circlesor makes a short flight, and on a few occasionsflights have been seen to pass southwardout of view. In most of the flocksthe movementis well coordinated,closely organized, and pursuesits coursenorthward. I have gained the impressionthat the northward movementof longspursat Anaktuvuk is as pronounced a directionalmigration as is the tremendousmovement of eider ducks flying eastwardin springalong the arctic coastpast Barrow. The long- spur is the most numerousmigrant to passAnaktuvuk in spring, an(1 many thousandsare transferredtoward the arctic slope. Althoughthe NunamiutEskimos now are basedat AnaktuvukPass, 330 IRVING,Migration of LaplandLongspurx [Vol.[ Auk78 their olderpeople have lived in manyvalleys of the BrooksRang• betweenlongitude 150ø and 158ø , which their ancestorsoccupied as nomadic caribou hunters. These named six other valleys through which longspursmigrate northward in numberscomparable with or exceedingthe migration at Anaktuvuk and indicatedthat all valleyswere utilized accordingto the extent of their grassyland. I have seentheir migration through the Killik Valley and Howard Pass. Recordsfrom Anaktuvuk show that the earliestreports of longspurs in springdiffer from year to year morethan is the casefor other arctic migrants,each species of which commonlyarrives within'a few days of its regular date. Further, as a result of close observationby resi- dent people,it is apparentthat after the earliestlongspur was noticed, on the average 12 days elapsedwhile only a few individuals,mostly males,were seenuntil numericallysignificant migration in flocksbegan suddenlyand continuedfor 10 or 15 days. This rather unusualprogram of migration may be peculiarto the longspursat Anaktuvuk; or it may be that the observationof migration by many Eskimos resident in open tundra, where visibility is especiallygood and migration is con- strictedin a narrow valley, detectsthe few early migrantsthat escape the view of singleobservers at lessfavorable positions.

MIGRATION ELSEWHERE IN THE WESTERN ARCTIC Longspursnest near Kittigazuitin the MackenzieDelta '(Porsild, 1943). Severalreports of migrationand nestingon the arctic coastof Territory have been compiledby Rand (1946), and Kesseland Schaller (1960) recentlyfound a few nestingnear the headwatersof the Sheenjek River in the northeasternAlaskan Brooks Range. We did not find them in the valley or on the tundra within 16 kilometers (10 miles) north of Old Crow, Yukon Territory (Irving, 1960a). While watching the spring arrivals at Demarcation Point and Humphrey Point, respectively,in 1914, Brooks (1915) and Dixon (1943) reportedthat a few longspursarrived there to nest. From their accountsand from the reportsof Eskimoswho have lived on the eastern arctic coast of Alaska, I have concludedthat there is no evidencefor a westward migration passing from Mackenzie Valley coastwiseinto Alaska (Irving, 1960a). Alfred Bailey (1948) reportedthat longspursseemed suddenly to arrive at Barrow and all alongthe westernarctic coastin the third week of May. During the 1st International Polar Year John Murdoch (1885)reported their arrival at Barrowon 20 May 1880. Sincemigra- tioncoastwise along the northeasternAlaskan coast has n6t beenre- July}1961J IRVING,Migration of LaplandLongspurs ported,I havetaken for grantedthat migrantsto the arcticslope and coastcame with the manyflights that havebeen seen passing through the valleys of the Brooks Range. Max C. Thompsoninformed me that during the springof 1960 he and severalother ornithologists(Williamson, et al., unpublished)who watchedthroughout the spring and summerat Cape Thompson(lati- tude 68ø 15', longitude 166ø) saw no group of longspursnear that northwesternAlaskan coastgreater than 10 and that during migration they noticedonly aboutas many individualsas they subsequentlycounted while nesting. Many longspursnest there, but it seemsthat the area is a terminus and not a through route for their migration. Leonard Peytonand I foundmany longspurs already nesting about Cape Prince of Wales and Lopp Lagoonon 15 June 1960,and Alfred Bailey (1948) reportedthat they were commonthere on 29 May 1922. His associate,Dwight Tevuk, who has made many importantcollections at the tip of ,recently informed me that many longspurs migrate at Wales early in the first part of May and that some have appearedon the coastfrom over the sea. Longspurshave been reported in the interiorKobuk Valley (Grinnell, 1900), where I have also found them nestingin summervisits in 1954 and 1957. Late in May 1951 I saw many longspursmigrating north- ward through Howard Pass, and just east of there in 1952 we found them the commonestbirds nestingalong the valley of the Ahlasuruk River (Irving and Paneak, 1954).

MIGRATION IN THE PORCUPINE VALLEY AND •N•ORTHERN YUKON

At Old Crow, Yukon Territory, Leonard Peyton and I first saw a longspuron 4 May 1957,and occasionallysaw a few malesuntil 12 May (Irving, 1960a). After that somefemales were also seenuntil 24 May, and groupsas large as a dozen were occasionallyseen flying east- ward up the PorcupineRiver as if coming from the Yukon Valley. We did not find them nestingin the PorcupineValley, but Kesseland Schaller(1960) founda few longspursnesting in the SheenjekValley 200 kilometers(125 miles) northwestof Old Crow. While watching for migrationat Fort Yukon in 1958, Peytonand Francis S. L. Wil- liamsonsaw a pair of longspurson 4 May, a male on 5 May, and a few on the edge of the airstrip on 6 and 7 May. Judgingfrom the small numbersseen at Fort Yukon and Old Crow, the PorcupineValley is not an importantmigratory route for longspurscoming to Alaska, and the northeasternpart of their nestingrange does not extendvery far south of the arctic coast. 332 IRw•c,Migration of LaplandLongspurs .Vol.[ Auk78

OBSERVATIONS OF LONGSPURS MIGRATING WESTWARD TOWARD ALASKA

Since the early studiesof Alaskan birds (E. W. Nelson, 1887 and 1913), it appearsto have been commonlyconsidered that the Lapland Longspursnesting in Alaskawinter in the interior westernstates. They are reported in winter as far north as southern British Columbia (Munro and Cowan,1947) and migratenorthward through the western MackenzieValley (Cooke,1910 and Munro and Cowan,1947). In 1958, while Peyton and Williamsonwere proceedingto observe at Fort Yukon,I set out from Anchoragealong the GlennHighway to join the and travel eastwardthrough the into southernYukon Territory. I had compiledevidence (Irving, 1960a,p. 277) that observationalong this coursemight be expectedto find flightsof a numberof continentalland birds migratingwestward and northwardinto Alaska. I first identifiedlongspurs on 24 April 1958,in a flockof 30 alongthe GlennHighway, 265 kilometers(165 miles) northeastof Anchorage. The next flock of 50 was identified on 26 April, at mile 1,045on the AlaskaHighway at BoutilierSummit, YukonTerritory, between Lake and HainesJunction. On the fieldsof the WhitehorseExperimental Farm westof HainesJunction a compactflock of about 100 longspurswas millingabout in rapid, swervingflight and occasionallyalighting briefly on the stubblefields. Just beyondWhitehorse on 27 April a flockof 100 took off from the airfieldand headedwestward. In anotherflock of 100, whichwere drinkingand feeding so busily that they appeared reluctant to fly,I ex- amined50 individualswith binocularsat closerange and counted48 malesand two females. It had beenmy impressionthat in the six flocksnoticed so far, mostof the birds were males. The trend of their flights was westward. Ninety-fivekilmneters (60 miles)east of Whitehorseat Johnson's Crossingover Teslin River many flocks of longspurs were flying swiftly downthe river in thefirst days of May,occasionally swerving and some- timescircling before passing over the high bridge and embankment on thehighway. Only one flock was observed to alightbriefly on theflats alongthe river. In one periodof five hoursI notedthat I saw flocks estimatedat 2,500 birds flying northwestdown Teslin River. I doubt if I noticedhalf of those passing within a halfkilometer of myposition, for I waswalking about searching for otherbirds. The longspurs werenot restrictedto the flatsof the TeslinRiver Valley, for I saw flocksmoving westward over the high ground when I occasionally traveledon the highway. As I waspassing southward along the high•vayto Atlin, British July]1961] IRVING,Mi#ration of LaplandLon•tspurs 333

Columbia,on the east side of Lake Atlin on 2 May, I saw many flocks oi longspursin flight, a few alightedon the ground,and a few small groupsperched in trees. I studiedthree of theseflocks at closerange as theyfed on the groundand countedthe proportionof femalesas one, two, or at most three among 10 birds. An automobileahead of me had ki!led or wounded15 longspursfrom a flock on the Atlin Highway, and only three of them were females. At the village of Atlin there were severalflocks of longspurs,one of which includedabout 200 birds, but they were apparentlypausing in the village. During this day I esti- mated that I noticed about 40 flocks of from 20 to 150 longspurs, averagingabout 50 birds or some2,000 in all. The commondirection of their flight was xvestward. While with Charles Sheldon hunting for specimensof Dall's sheep on the Macmillan River, Osgood (1909) saw large flocks of longspurs early in September,and Rand (1946) saw them migrating in the same country in late summer.Many longspursappear suddenlyin spring at Dawson (Betts, 1940). The westwardcourse of the spring migration of longspursthat I observedextended over the country between the AlaskaHighway and Atlin for a width of 80 kilometers(50 miles) from north to south. Reports listedby Rand (1946) indicatethat the migra- tory path in Yukon may extend 300 kilometers(200 miles) northward to the southernwatershed of the Ogilvie Range. H.S. Swarth (1936) reported the first longspursat Atlin, B.C., on 25 April 1930, 4 May 1931,21 April 1934,and 11 April 1935; the averagedate for thesefour years being 22 April.

PROGRESS OF •IIGRATION IN ALASKA

Taking 49 recordsof arrival in variousyears at 30 localities,the dates at 27 localitiesfollow the average date of arrival in Atlin (Table 1). I have chosenAtlin as a reference point hecausea careful observer, H. S. Swarth (1936), observedthere during the spring and sumlnerof four years and becauseit is in the path of large migratory flightsthat I observedin progresswestward. Migration to Mackenzie Delta is not likely to pass near Atlin, and arrival at Mountain Village four days after Atlin wouldrequire flight at the improbablespeed of 413 kilometers per day. Omitting these records,the average advanceof the front of niigration from Atlin to 17 placesin northern and northwesternAlaska was 8• kilolnetersper day, a distancethat longspurscould travel in two hours. Migran's may not be c•served until some days after they arrive. Further, the arrival of longslmrshas been variable from year to year 334 IRvx•c,Mi#ration of LaplandLo•gspurs [VoL[ .Auk78 July1961 IRWSG,Migration ofLapland Longspurs 335

< < < < < • < • < < • 336 IR¾•'G,•Iigration of LaplandLon.qxpurs [Vol.[ Auk78

in localities that have been under sustained observation. Erroneous reportsor erratic migrationcould causethese records to be late and so lead to calculations of slower than actual progress. Since it is a likely possibilitythat the swift-flyinglongspurs can travel 150 kilometers per day, a delay of five or even 10 days in any of theseannual local records would, if corrected, still lead to a plausible calculationof the progressof migration. Reports show that migration to the western coastsof the Gulf of Alaska evidentlypursues a different schedule. The recordedarrival of longspursat Dyea, Yakutat, and Homer precededthe recordsfor Atlin. Among the nine other recordsfrom the south-centraland southwestern coast and Aleutiau Islands, only two (Cold Bay and Unalaska) follow arrival in Atlin by an interval sufficientto make the calculatedprogress of the van of migration within the possiblespeed of flight. The other five records would require flight from Atlin at an impossiblespeed. In these cases erroneouslylate records would only increasethe im- probabilitythat flightsto southwesternAlaska and the Aleutian Islands proceedthrough the headwatervalleys of the .

MIGRATION FROM. TO ALASKA

Dr. Frank Pitelka informed me that from his observations at Point Barrow the Lapland Longspursmight arrive in northern Alaska from wintering in Asia. Accordingto his comment,thousands of longspurs have been seenat Barrow flying from the west late in May and in the first part of June. Further, of 1,400 bands applied to longspursat Barrow, none were recovered either from well-populated western America or from the interior of Asia that we know so poorly. If direction of migration can be inferred from flights observedat one point, Dwight Tevuk's remarks that longspursappeared over xvater near Wales might also suggestmigration froin Asia. Stejneger (1885) reported that longspursfirst reachedthe Com- 1nanderIslands on 21 April 1883, but the specimensfroin there were later assignedto the darker race, C. I. coIoratusRidgway (1898), which occurs in Kamchatka and adjacent (Vaurie, 19'59). Olaus Murie (1959) repeated Ridgway's (1901) opinion that over the Aleutian Islandsthe longspursdid not differ froin C. I. alascensisof the mainland. These reports indicate opinionsthat eastern Siberian long- spurs do not migrate to the Aleutians. Vaurie (1959, p. 706) observedthat "possiblyall or the majority of visitors to are coloratus." That distinction indicates the unlikeli- hoodthat many of the light-coloredC. I. alascensiswinter in China and July]1961] Im'•N(;,Mi9ration of LaplandLoncdspurs 337 suggeststhat the nearestAsiatic range in whichto suspectthat some Alaskannesting longspurs might winter unrecognizedamong Eurasian C. l. lapponicuswould be in . The distanceand directionof such a migration does not appear ilnpossiblefor the swift-flyingand sturdy longspurs,because populations of four smaller and weaker- appearingspecies of land birds, Oenantheoenanthe, Luscinia svecica, Motacillatiara, and Phylloscopusborealis, regularly migrate from win- teringin warmareas in Asiato nestin Alaska(Irving, 1960a,p. 270). Longspursnest on the ChukchiPeninsula adjacent to Alaska,but their migratory schedulesand taxonomiccharacters have not been reportedin detail that providesevidence for the extent and direction of exchangesof longspursbetween Alaska and Siberia.

•'IIGRATION IN SOUTHEASTERN ALASKA

Migrating longspurshave been frequentlyreported in southeastern Alaska (Gabrielsonand Lincoln, 1959), but the reports do not show that the coastwisemigration is a great contributorto their nestingin Alaska. Longspursnest on Middleton Island 120 kilometers(70 miles) southof the mainlandin the Gulf of Alaska (Rausch,1958). Consideringthat longspursreach west Greenlandfrom America early in May (Salomonsen,1951), it would be possiblefor them to make extensiveflights over the Gulf of Alaska to reach south-centraland southwesternAlaska at the early dates when they appear.

MIGRATION OF I.ONGSPURS THROUGH WESTERN

In British Columbia1ongspnrs nilgrate northward in numbersthat passmainly east of the RockyMountains (Munro and Cowan,1947). These authors designatedthe longspursof British Columbia C. I. alaxcensis. At Fort SimpsonPreble (1908) considered24 out of 33 longspurscollected soon after 25 April 1904 to be C. I. a[ascensis.In assigningthe longspursof Banks Island to C. I. lapponicus,Manning, H6hn, and Macpherson(1956, p. 116) distinguishedthem from the longspursmigrating through western Canadianprovinces and Alaska. They related the birds from the latter localitiesto C. l. ala,vcensis. These recent opinionscontinue the older views that the longspursthat migrate northwardin springthrough the westernMackenzie Valley to Alaska resemble those from Alaska and can be differentiated from the longspursthat neston ,Canadian Arctic Islands,on the main- land east of Mackenzie Bay, and in northern . 338 INVlNG,Mi9ration o/ LaplandLon9spurs [Vol.I Auk 78

•¾[IGRATION OF OTHER SPECIES IN YUKON TERRITORY

At Old Crow in the Porcupine Valley, migration carries several speciesthat migrate along the Pacific Coast eastward, but no migration has been reported passing westward through the arctic part of Yukon Territory toward Alaska. I have reviewed evidence based upon dates of published reports from Alaska, Yukon, and British Columbia,which indicatedthat a number of American species of land birds migrate to Alaska through the watershed of the Yukon River in southwesternYukon Territory (Irving, 1960a). While I was observing the migration of longspursthrough southwesternYukon during late April and early May in 1958, birds of the following specieswere also observed moving north- westward but not eastward: Branta canadensis,.4nser albifrous, •qnas platyrhyn- chos,.4nas acuta, a few Buteo lagopus,a small number of Circus cyaneus,Irido- procne bicolor, Dendroica coronata, Euphagus catolinus, and Spizella arborea. Only two species, Bucephala islandica and Larus canus, were observed flying southeastwardalong Teslin River, as if they had entered western Yukon Territory from eastern Alaska.

DiscussIoN

A courseof migrationcan be inferredfrom observationsshowing a commondirection of flight over a significantportion of a migratory path. Directional flights have been watchedin southwesternYukon as they passedmany points along a coursewestward for 260 kilometers (160 miles) betweenJohnson's Crossing and HainesJunction. Great numbersof longspurswere observedproceeding uniformly westward over the 80 kilometers(50 miles) south of this line to Atlin, B.C. Directional flights have also been observedduring severalyears at Anaktuvuk. I have watchedflights moving northward at pointscover- ing about65 kilometers(40 miles) in the narrowupper valley of the Anaktuvuk River. Since 1947 residentEskimos have annuallyre- portednorthward flights of longspursin springover their campsand huntingparties that were distributedthrough 130 kilometers(80 miles) of the upper valleysof the John and Anaktuvuk rivers. This north- ward migrating stream is only a few miles in width, but it has been observed to transport many flocks of longspursnorthward. The Nunamiuthave seenflights of longspurspassing northward while they were earlier living as nomadicfamily groups encampedin spring in variousvalleys of the BrooksRange between Howard Pass (longitude 158ø W.) and the Itkillik Valley (longitude150 ø W.). The same northwardmigration of longspurswas describedto the presentNuna- miut by their parents.Over coursesextending from 80 to 160 kilometers (50 to 100 miles) northwardand across290 kilometers(175 miles) from east to west, a great migrationof longspurspasses through the valleys of the central Brooks Range. July19611 l IRvxx6,Migration of LaplandLongspurs 339

Geographicalsuccessiou of datesof first-arrivingmigrants is also an indication of the course and progress of the van of migration. Iu Alaska north of the Alaska Range reports from 19 localitiesfollow the averagedate of arrival of longspursat Atlin. At 18 localitiestheir ar- rival occurred at dates consistentwith reasonablerates of flight from Atlin. Records for the significant18 localitieshave been averaged in four series accordingto intervals of longitude in Table 1. The cal- culated speedto points betweenlongitudes 134 ø and 145ø is 62 kilo- metersper day; between148 ø and 156% 73 kilometersper day; 157ø and 164% 91 kilometersper day; and 1660 and 171ø , 124 kilometers per day. The speedof the van of migration increaseswestward. The advanceof migration over south-centralAlaska and the Aleutian Islandsexceeded the possibilityof flight from Atlin. When I informed Dr. Axel Hemmingsenof the earlinessof arrival of longspursin south- central and south•vesternAlaska, he brought to my attention the fact that migration of some specieseastward and westwardthrough Asia and proceedsmore rapidly than northward. He had indicated (1951) that the advanceof a numberof migrationsin Asia proceeded northwardabout as the movementin springof fixed daily amountsof incidentsolar radiation. This relation would be fulfilled by the simul- taneousarrival of the migratory front of a speciesall along a given high latitude. Recordedarrivals of longspursin south-centraland western Alaska are not simultaneous,but they do not show progressioneastward or westward.These recordsare not only fewer than those in the north, but I know that severalof them were madeby travelingnaturalists as they arrived at localitiesand first saw longspurs. The recordsfrom north of the Alaska Range were made by observersresident in those localitiesduring the spring migration, and usually in villages where resident people attentively watch and discussmigration. I suspect that carefulobservation will provideeven earlier recordsof longspurs arriving on south-centraland southwesternAlaskan coasts. Recordsfrom the Aleutian Islands are too early to indicate that migrationreaches there by skirtingthe coastof the Bering Sea after crossingBering Strait or Sea from the ChukchiPeninsula. Only the existenceof C. I. coloratuswest of the Aleutiansis evidenceagainst mi- grationfrom Siberiathrough the Aleutians. Althoughthe progression of arrivalsin northernand westernAlaska is consistentwith migration throughthe headwatervalleys of the Yukon, it is possiblethat some longspursmigrate from Siberia to Alaska. Four passerinespecies regularlymigrate in that direction,but a larger numberof speciesmi- 340 I}•V•NG,3iicjratio•z of Lapla•zdLo•tcjspurs [Vol.[ Auk 78 grate over the interior of Americato reacheastern Siberia. In recent yearswe havehad no view of the exchangebetween Asia and Alnerica at the northern point where they are so closegeographically. No doubt there are other localitiesthan in Yukon Territory and the Brooks Range where migrating flocks of longspurscan be watched along courseslong enough to define the direction of their migratory flights. By strategicselection of a few localities,a prearrangedtactical schemeof observationmay definethe physicalprogress of migrationin one year. in addition, the flocks can be characterizedby ratios of sexes,social behavior, plumage, fatness, and conditionof gonads. These characteristicscan be used to distinguishthe flocksand to demonstrate social and physiologicalfactors that are causally related to the or- ganizationand progressof migration.

SUMMARY

Lapland Longspurs,ascribed to Calcariuslapponicus alascensis Ridg- way, nest in great numbersover the arctic tundra and grassy land westof the forestsof Alaska and over the Aleutian Islands. During 10 yearsgreat numbersof longspurshave been observedin May migrating north through Anaktuvuk Pass. Similar migrations proceed through other interior valleys of the Brooks Range, taking vast numbers of longspurstoward the arctic tundra. There is no evidencethat this migration proceedswestward to Alaska through arctic Yukon Terri- tory or eastwardfrom Siberia. In the last week of April and first of May 19'58, great numbers of longspurs,in flocks at first containing mostly males, were seen mi- grating westwardover a distanceof 260 kilometers(160 miles) through the headwater valleys of the Yukon River in southwesternYukon Ter- ritory. Longspursreach northern and western localities north of the Alaska Range after their appearancein southwesternYukon Territory at dates indicatingplausible rates of lnigration from Yukon Territory through Alaska. South of the Alaska Range and on the Aleutian Islands, arrivals are so early that theseregions cannot be settledby the migrantsobserved in Yukon Territory. Unobservedflights coastwiseor overwater are sug- gested. The northward advanceof the front of longspursmigrating through .Alaskais slow, but it increasesas the northwardcomponent of migra- tion diminishesand as the westwardcomponent increases. July19611 l I[•VXNG,Migration of LaplandLongspurs 341

LITERATURE CITED

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