BIRD-BANDING A JOURNAL OF ORNITHOLOGICAL INVESTIGATION

VoL XXXII January, 1961 No. 1

STUDIES OF THE BREEDING BIOLOGY OF HORNED LARK, WATER PIPIT, LAPLAND LONGSPUR, AND SNOW BUNTING ON BYLOT ISLAND, NORTHWEST TERRITORIES, CANADA By WILLIAM H. DRURY,Ja. INTRODUCTION This paperis dividedinto threeparts. The first part reportsdetailed observationson the breedingbiology of the four species.I haveincluded many detailsbecause they showecological factors deserving discussion, and behaviordifferences suggesting unexpected geographical variation. The secondpart discussesecological questions such as how the four speciessurvive together without competition, what adaptations allow them to breed successfullyin the Arctic; and what is the relation of large northernclutch-sizes to annualproduction and the shortbreeding season. The third part comparesthe courtshipbehavior of LaplandLongspurs with studiesof other buntings,specifically Andrew's (1957) aviary studies.These comparisons suggest either that thereare unusuallylarge geographicaland interspeciesdifferences among buntings, or that their evolutionaryrelations are not clear. Thereappear to be adaptivechanges in territorialbehavior during the early part of the longspur'sbreeding cycle. The 1954Bylot Island Expedition spent from 12 Juneto 29 Julyat the mouthof the AktineqRiver, southernBylot Island,approximately 73 ø North Latitude, 79ø West Longitude,District of Franklin, Northwest Territories,Canada. Bylot Islandis on the borderbetween the Low and High Arctic areas.and is just northof the north coastof BaitinIsland (see map in Miller, 1955). Shortdescriptions of the trip (Drurys, 1955) and of the area (Van Tyne and Drury, 1959) havebeen published. A popularaccount of certainaspects of the expeditionhas beenpu,blished as Springon an Arctic Island by KatharineScherman (1956). The observationsin this paper were madeby William and Mary Drury, and Dr. BenjaminFerris, who concentratedtheir time on the breedingbirds; and by JosselynVan Tyne,who contributed infor.mation gatheredon daily collecting trips. No collectingwas done in a studyarea of one squaremile. ContributionNo. 16 from the Hatheway Schoolof ConservationEducation, Massa- chusettsAudubon Society, South Lincoln, Massachusetts. 2 ] DI•uI•Y,Bylot Island Breeding Biology Bird-BandingJanuary

The study and preparationof this paper was made possibleby a sabbaticalhalf-year at HarvardUniversity in 1955. Sincethe ,first draft, timeand expenses associated with publicationhave been available through the ttatheway School,Massachusetts Audubon Society. JosselynVan Tyne's sicknessand untimelydeath preventedpreparation of a joint report and delayedpublication. The accompanyingmap (Figure I) showsthe vegetationof the study area and nests.A field map of the studyarea, showing elevations and namesused in the text,has .been published with the faunallist (Van Tyne and Drury, 1959), and in Drury (1960). Subspeciesidentifications and commentsare to be foundin the faunal list.

BARREN

DRY MAT PLANTS

WET MOSSY VEGETATION

• WATER PiPIT O LAPLANDLONGSPUR O NEST NOTFOUND ß LONG-TAiLED ,JAEGER PERCH •) RED-THROATED LOON • HORNEDLARK • SNOWBUNTING • NESTNOT FOUND

Figure 1. MAP OF THE STUDY AREA, ,showing•:he vegetation types and location of nests discussed in the text. Vol. 1961.XXXII DauaY,Bylot Island Breeding Biology [3

PART 1: SPECIES ACCOUNTS HORNED LARK Eremophilaalpestris (Linnaeus). (Eskimo:Kah-oh-r6d-lee-rah or Kah- 6h-dlee-rah). HABITAT Two nestterritories and the areasoccupied by 22 pairs were all in the mostexposed places of the uplands,almost free of vegetation.These areaswere on hilltops,ridges, old beaches,or fro•st-heavedareas where mostof the surfacewas tan, angulargravel. All were the first places free of snow,and the typical plantswere: black, crustoseli.chens and deadmosses, Gray Lichen (Stereocaalon paschale [L.] Ach.), andclumps of Purple Saxifrage(Saxi/raga oppositi/oliaL.), and Poppy (Papaver radicatumRottb.). Arctic Willow (Salix arctica Pall.) and Bell Heather (Cassiopetetragona [L.] D. Don), grassesand GrassRush (Luzula con/usaLindeb.) grew in mats in the hollowsand on slopes.

DISTRIBUTION AND DENSITY Hoyt'sHorned Larks (E. a. hoyti), althoughnot numerous in any spot, were the most widely distributedbird, exceptthat Snow Buntingsoc- curred in the mountainsabove the icecap.There were four pairs of larksin our studyarea. The centersof their territorieswere about 400 yardsapart. I havereported census data for the four passerinespecies dealtwith here in Van Tyne and Drury (1959).

TERRITORY Song. Larkswere singing when we arrived,and continued to singuntil 26 to 28 June; then frequencydropped and we heard noneafter 3 July. The birds sang over: the west end of Kungo Hill (Drury, 1960); the Upper PhalaropePonds; and Tui-Tui Tabletop. Each songpostwas constantand over the centerof a territory. We agree with Pickwell (1931, 1942) that onlyone bird wasin the air at once,,but we haveno indicationthat singinglarks flew overa neighbor'sterritory. We heard the "recitative"(Pickwell) rarely in the evening,and rarely heard any songgiven from the ground.These were short bursts, and seemedto be preliminaryto a flight song. Pickwellpoints out that humanintruders are greetedwith a flight song,and a disputeat GoldenPlover Creek descri.bedbelow may havebeen caused by our approach. Justbefore a bird flewup to sing,he stopped,standing unusually erect with hornsraised and featherssleeked (Figure 2) on a tuft of grassor a rock. He usedthe sametake-off perches regularly. The postureand movementcontrasts with the crouched,shuffling amble of a lark feeding undisturbed(Figure 2). The bird rose with undulatingflight, silent, sometimesas if in spiralsbut really"tacking" upwards. When he began to sing,he sailedwith spreadwings and tail (Figure2), or dropped with nearlyclosed wings. Betweensongs he climbedfor 10 to 20 beats. In this way he hung nearly in place,but movedup and downsinging short-lispingphrases, Pickwell's "intermittent song," more or lesscon- tinuously.Pickwell (1931) givesa, bout five minutes as the maximum time for a songflight. In our subjectiveimpression, the songis mellower thanthe rather dry songof a "Prairie"Horned Lark (E. a. praticola)and more like that of the northernsubspecies (E. a. alpestris).The bird 4] DauaY,Bylot Island Breeding Biology Bird-BandingJanuary performedthesame reckless drop to the ground after singing. Birds sang 150to 200 feet over Kungo Hill. Earlymorning, evening between 2000 and2200, and cloudy, windy days, were favored for singing. In a wind, thesinging bird soared with wings half-closed and tail widely spread, flyingalways into the wind. Under such conditions they often sang at about 50 feet.

AGGRESSIVE BEHAVIOR Wesaw one boundary quarrel on the west side of GoldenPlover Creek on21 June.A birdfrom overhead dove at thebird feeding on the river bankand drove it rapidlyabout 40 yardsin a zigzagcourse towards KungoHill, thenrose again to givea flightsong.

chwee-chwee. chwee tssoo.tssoo.ts$00 FLIGHT SONG

WATER PIPIT

BEGS AT SONG PERCH FLIGHT SONG FOR COURTSHIP FEEDING

SNOW BUNTING

....•• FEEDINGerillip DISPLACEMENT FEEDING BEFORE INTRUDER AT NEST

HORNED LARK Figure 2. WATERPIPIT, SNOWBUNTING, AND HORNEDLARK. Displays. Vol.1961XXXII DIeulaY,Bflot Islan'd. Breeding Biology [ 5

On 14 Junetwo larksfed freelyWith five SnowBuntings and five LaplandLongspurs onthe still wet, g•ssy'crest of the-Bluffsat the west- ernedge of theirterritory. This persistent flock behavior may havebeen correlatedwith the latenessof the 1954 sdason.Only 10 to 15 percent of theland surface was free of snowat thisperiod in 1954,-whereasD. V. Ellis tells us that at the time of his visit, 1 June, 1955: "There was very little snowon the groundby the (camp) site."

NEST AND EGGS Duringthe first week (12 to 20 June),the males were either constantly singingaloft or closelyfollowing the females.According to Pickwell, theseactions indicate nest building as muchas doesnervous activity of thefemale. We did not find any nestsuntil the younghad hatched. Nestsites and construction of hoyti agreewith thoseof Pickwellfor praticolaand DuBois (1935) for leucolaema,but differfrom those of alpestris.Townsend and Allen (1907), Sutton(1932), Soper(1946), Wynne-Edwards(1952), Suttonand Parmelee (1955) describethe nest of alpestrisas sunk in growthof lichenor moss,perhaps under a hum- mock,but definitelyin thevegetation. The two nests we foundwere on bareground, on a steepslope, and in a shallowexcavation with no evidentrelation to a clod. They wereon the upperedge of continuous matplant cover and had southern exposure. The lower side of thenest wasbuilt of small,flat stonesand pieces of dirt or clodheld together by dryingand by lichen thalli (compare DuBois, 1935). Thenest lining wastightly woven of gray,dead •asses and Grass Rush, with four to six feathers. The floor of the nest was dirt. •

Figure3. HORNEDLARK YOUNGIN •NESTNO. 2. Youngare four daysold and were not seen to be brooded during the day. 6] DRtmY,Bylot Island Breeding Biology Bird-BandinõJanuary

4''

Figure 4. HORNED LARK YOUNG IN NEST/NO. 1. Young is six days old.

Clutch-Size.One of our nestshad four and one five eggs; the four familiesfound out of the nesteach had five young.

HATCHING AND CARE OF NESTLINGS 1. Feeding. We did not see larks brood their young on the nest duringthe day. They did not fly directlyto the nest,but lit at a spot about20 yardsaway and ran, stoppingas if to feed, alongan irregular routeto the nest. The giving of food to the youngwas so rapid that it seemedto be a continuationof feedingunless one knew of the nestsite. After feeding,they ran 10 feetor more,stopped once or twice,then flew to a rock,dropped the fecalsac they usuallygot, and wipedtheir bill. Bothparents fed the young. 2. Developmento! theyoung. Pinfeathers were evident the third day amidthe yellow-buffdown. The nestlingskin was brown. The insideof themouth of thenestling for thefirst four days is yellowish,reddish below theto.ngue and ,bill rami. In this periodthe younggape a.t any disturb- ance. After their eyesare open(between the third and fourthday), the nestlings'press themselves flat and still into the bottomof the neston approachof a human.Fourth day youngare shownin Figure3. Sixth dayyoung are shown in Figure4. Theykeep this behavior until they leave the nest. For the first weekafter leavingthe nest,even when able to fly, the youngwill allow approachto within 15 to 20 feet. The parents' ]rrreeeeetalarm note makes them crouch until disturbedindividually.

'Det.ailed descriptionsof the nests, their constructi,on, topographiclocality and vegetationsurroundings, are on file at the HathewaySchool, Drumlin Farm, South Lincoln, Massachusetts.They are omitted here becauseof their bul'k and because they are of very specializedinterest. We have descriptionsfor all speciestreated in this paper. They can be consuhedon reauest. Vol. XXXII •061 DauaY,Bylot Island Breeding Biology [ 7

The youngwere five daysout of the nest (6-11 July) beforethey flew about30 yards,but five days coincides with the •ninimum for thespecies. Our observationsagree with Pickwell that the fledged young, barely able to fly, hop insteadof walk. 3. Nestlingperiod. The nestlingperiod was eight-nine days compared to 9-12 days in five nestsin leucolaema,DuBois (193'5).

REACTION TO PREDATORS Whenwe werenear their nests,the larks fed nervouslyand rapidly, coveringa large area and circling aroundthe nestand intruder. When feedingundisturbed, they fed creepingand walkinghere and thereover the barrendirt of "frostings"(Washburn, 1956) and wind-cleanedareas. Whenthere were eggs,or eggsand youngin the nest,the femaleleft the nestwhen we first appearedover the skylineat a distanceof 250 to 300 yards,"casual abandonment" mentioned by Pickwell(1942). Her flight was low and direct and followedthe sameroute each time. We foundno distressdisplay at the nest,but foundit frequentlywith parents with flightlessor weakly-flyingyoung. Whenwith their youngthe parents sat very close.then sprangfrom the groundwith noisy wingbeat. The male wasmuch less aggressive and under thesecircumstances often rose ten to fifteenfeet into the air and sanga few short,atypical phrases. On 30 June larks startedto give a different alarm note, a contralto shorebird-likejrrrreeeet; the regularalarm call is tseeeep.

WATER PIPIT Antht•s spi•oletta (Tuntall). i American Pipit). iEskimo: Engeo6k- dj•eyookl.

HABITAT This specieswas ahnostentirely restrictedto the sun-soakedravines cut into the Bluffswest of camp. Soper (1928, 1946); Suttonand Par- melee (1954a}: Wynne-Edwardsi19521 agree that this speciesis re- strictedto steepslopes, where the suncan havemaximum warming effect. We found pipits at this. the most northernrecorded nesting site, to be closely correlated with the .most mesophytyicvegetation, richest in speciesof plants.

DISTRIBUTION AND DENSITY Along the mile-and-a-halfstretch of the Bluffs west of camp, there were sevenareas where parents scolded. Each group was separatedby a gap of at leasttwo valleys.

TERRITORY Song. Pipits werein songon 12 June. Variousauthors (Sutton and Parmelee,1954a; Soper,1928) recorda songwhich changesin quality or accent. Others (Dixon, 1938; Pickwell, 1947) record songsthat changeonly in tempo,accelerating rapidly as the bird divesfrom the top of its flight. Townsendand Allen (1907) and Lewis (in Bent, 19501 recorda trilling song. We hcard uniformly two-partedsongs given on the ground and in the air. Just beforestarting to sing, the bir.d ran nervously.fitting onto rocks where he sangbriefly, and sleeked his body 8 ] DauaY,Bylot Island Breeding Biology Bird-Ban(lingJanuary feathers(Figure 2). As he flew up, usuallyto the level of the crestsof theBluffs, he calledtsoo-tsoo-tsoo-tsoo andbeat his wingsin a nervous, shallowbeat. Thenhe glideddown with primaries closed and secondaries open,tail furledor partlyopen, and immediately changed his songto an acceleratingchwee-chwee-chwee-chwee (Figure 2). He lit again on a conspicuousrock or ridge and stoodwith neck stretchedup and feathers pressedfiat, scoldingtseep or, as often,repeated his songfrom the ground. In the groundsongs, the chwee-chweewas more common. On 29 June,a pipit flew into a neighbor'sterritory calling the usual flightcall chip-chip,chip-chip-chip. As it trespassed,the territoryholder trilled loudly,dropped off its high perchat the crestof the ridge, and doveat the newcomerwho fled. Shortlyone bird returned,sat on its usual tussockperch, and called tseep-tseep,zzeeep-zzeeep-zzeeep. On anotheroccasion, we hearda parentgive the trilling songwhile carrying insects.This song seemedassociated with imminent attack. Song stoppedduring the first week of July. We heard song once 20 J.uly.

AGGRESSIVE BEHAVIOR On 15 June we saw many chasesin severalravines •est of camp. Two birdsmet and had a brief fight; thenone chased the otherat break- neck speedin and out of the ravines,up and down from the Bluffs to the beach,round and round,then they just vanishedinto a ravine. In some cases three birds were involved.

TERRITORIES STUDIED Each pair of birds fed in four or five valle}s along a distanceof about 100 yards and 200 feet high. Sometimeswhen feeding, silent birds intruded into neighboringterritories without fights. At other times, seeminglywithout relation to the progressof the breedingcycle, intruders were vigorouslyattacked. NEST ANr• E•;•;S see Table I The nest was built entirely of grasses.

REACTIONS TO PREDATORS We found parentswith young in late July. A zzeeepscold-note and nervouswalking around in the low vegetation,accompanied by vigorous tail-bobbingwhile they seemedto feed in the mattedgrowth was asso- ciated with alarm at the nest. Such alarmed .birds did not run and dart, however,as they do whenreally feeding. On returningto the nest, the disturbedparent characteristically "fed" alongthe edgeof the slump terracesheltering the nest,then hoppeddown and enteredthe nest. When we first saw parentscarrying food to youngin late July, we heard a new alarm note, zing-zing.

FEEDING Occasionallythese birds fed on the frost-heavedbarrens on the slopes of KungoHill or on the meadowsat the crestsof the Bluffs,but they usuallyfed in the edgeof openareas, walking in and out of the tuftsof grass,through mats of Bell Heather,patches of Locoweed(Oxytropis MaydellianaTrautv.), andVetch •Astragalus alpinus L.•. Theyseemed Vol. XXXII •96• Daua¾.Bylot Island Breeding Biology' [9 10] DRURY,Bylot Island Breeding Biology Bird-BandingJanuary to like the .tallergrowth, where they fed threadingtheir way through, duckingunder branches,stepping over low growth. At other times, they fed by searchingover the surfaceand catchinginsects by short ru,nsand flights. Occasionally,when flying aroundin pairs feeding,they gavea call similar to that of a CommonRedpoll (Acanthis flammea)---dik-dik, dik-dik, dik-dik-dik.

LAPLAND LONGSPUR Calcariuslapponicus i Linnaeus). (Eskimo: Ktingnuktah). Accordingto Frazer Rowell (1957), our breedingrecords for this speciesare the farthestnorth; but the A.O.U. Check-list(1957) lists breedingat DundasHarbor, Devon Island, and at Thule, Greenland, in Latitudenearly 77 ø North. Nestsreported from Taimyr Peninsulaand New Siberian Islands (Tugarimov and Tolmatschev,1934: and Birula, 1907; in Grote, 1943) are as far if not farther north in Siberia.

MIGRATION AND ARRIVAL Longspurswere presentwhen we arrived. For the next three days their numbersincreased and the numberof femalesincreased rapidly. Theywere not paired on arrival,as suggestedin Russianarticles (Michel, 1935; Tugarimovand Tolmatschev,1934; and Birula, 1907; in Grote, 1943) for populationsin the far north of Siberia. Duringthe first week,groups fed togetherand seemedto revertreadily to flock behavior and the flock call. On 14 June, four male and one femalelongspurs fed in a flock with Snow Buntingsand Horned Larks on a grassyarea at thecrest of the Bluffson the southend of WestRidge. Grotesays that in Siberia,flocks are maintainedwhile the tundraremains mostlysnow-covered.

HABITAT Longspursoccupied the thick, moss-flooredvegetation of dry places in the uplands,primarily on the east-and south-facingslopes, not the bogs. Their distributionwas similar to that of Bell Heather,but Bell Heathergrew over large areaswhere we foundno longspurs.

DISTRIBUTION AND DENSITY This wasthe mostnumerous nesting species around our camp. It was, however,nearly equalledin numbersby Baird's Sandpiper(Calidris bairdii), and was less widely distributedover the island than Horned Larks. Where longspursoccurred, they were crowdedtogether. A surveyof territoriesin mid-July--nests,parental alarm, or flightless young--indicatedat least27 nestson our studyarea, and at least60 in the sevensquare miles betweenEclipse Sound and the snoutof the AktineqGlacier on the westbank of the river. As othershave found, longspursbecame inconspicuous at the end of July, 'but we saw no obviouslymolting birds, nor any flocking or migration. Vol. 1961XXXII DRURY,Bylot Island Breeding Biology [ ] ]

TERRITORY Song'. When we first arrived, full songand territorial disputeswere goingon overthe radio mastsat Pond Inlet, but were sporadicon Bylot Island. Song increasedin intensityand frequencyat camp 15 June, and was general, and territorial disputes vigorous, by 17 June. Males took up territories between15 and 25 June. The malessang: (1) a partial, abbreviatedsong on a raisedhummock near the nestsite; (2) a full but not rapidly repeatedsong on two or threesong perches on rocksor someconspicuous break in slope; (3) a whisperedsong on the groundnear the female;and (4) a flight song'. When maleswere singingon the ground,the primarieswere lowered, bill raisedobliquely, tail barelycocked, cro.wn feathers sometimes raised, back feathers sleeked,and belly and flank feathers slightly fluflexi

SONG ON GROUND

INCITING

spat-drrrr • zeep

RUNS SLDWLY •-'•-•• THREAT

'•• beggingnote [ •. PRECOPULATORY chatters ' - • ACTIONSOF PRECOPULATORY ACTIONS • FEMALE OF MALE

POST-

ACTIONS OF FEMALE

• ALARM AT , NEST COPULATORYTERRITORIAL SONG

IN FEEDINGWIND

LAPLAND LONGSPUR

Figure 5. LAPLAND LONGSPUR. Displays. 12 ] D•,],•¾.Bylot Island Breeding Biology Bird-BandingJanuary

(Figure 5). The whisperedsong had the sameelements as the full song, but was usuallyshorter, given only in closecompany of the femaleand before nest building started. The usual (90 percent) full song was a flight song,in which the male rosewith regular wingbeatsto a height of 20 to 40 feet, then floatedto the ground (usuallyto a rock or other conspicuousspot), singingseveral songs in rapid succession.His course on the way down was usuallya semicircleand as often on set wings as on quiveringwings ("moth flight" of Hinde, 195.3). When the wind blew 10 to 1.4knots, males hung suspended,occasionally beating their partly-foldedwings, spread their tails wide, and sangfive or six songs in oneflight (Figure 5). Many time malessang as they rosefrom their songperches, especially in a songduel and whensuddenly responding to borderaggression. When doing this, their wingbeatwas rapid and shallow. As I interpretthe descriptionsin Bailey and Niedrach (1938), the Lapland.Longspur's flight songis like that of the Chestnut-collared Longspur(Calcarius ornatus), but differsin severaldetails from that of 'McCown'sLongspur (Rhynchophanesmccownii). The flight song is an intense,vibrant display,in contrastto the undulantflight and lispingsong of Snow Buntings. We found little individual variation in song, which our field notes describeas liquid, wheezy,and like a Bobolink(Dolichonyx) heard in the distance,Soper (1928), and Sutton (1932). Songsstarted with a ringing and metalliczing, followedby a rolling and rapidly descending zizeleeaw;then a rolling, sustainedzizelee-ee (ending with the highest noteof the songee), and closedwith anotherrapid rolling and descend- ing zizeleeaw.We found no variation in phrasingwith the progress of the season. Our birds had two or severalsong percheson their territories; in contrast Salomonsen(1950-1951) reported single song perches in Greenland.They choseany rock or conspicuousplace as a songperch, and appropriatedpiles of dirt we madein excavatingfrost forms on their territories. The •nale at nest No. 5 sang from the dirt pile before I finishedthe hole, but he had no suitablerock or ridge, and was under pressurefrom Nos. 1. 6, 10, and 12 (Figure 7), and his femalewas building. Notes preliminary to ground and flight songsincluded: the alarm note, dzeeu, and tsuk (perhapsthe "twuu" whistle of Frazer Rowell, 1957). We cannotbe sure whetherthey are preliminarynotes to the song,or expressionof alarm at our intrusion,because longspurs were stimulatedto give their flight songby our approach. All observersagree that singing decreasesrapidly after the female startsto incubate:two _maleswere singing28 Junefrom 2000 to 2200: at midnight on 3 July one longspurwas singing from the late nest (No. 6). He sang wheneverwe approached,for four days after the clutchwas complete,and then rapidly decreasedhis singing. After the clutch was complete,nearly all singing was in flight. Grote (1943) reportsthat late seasonsong in Siberiawas on the ground. Singingwas sporadicuntil one to two days after the young had hatched,by which time it had ceasedeven as an expressionof aggressionto a human intruder. We heardno songsafter 10 July. Grotereports that in Siberia the songperiod lastsabout three weeks. Vol. !961XXXII DRURY,Bylot Island Breeding Biology [ 13

AGGRESSIVE BEHAVIOR As with McCown'sLongspur (Mickey, 1943), oncea maleLapland Longspurstarted to sing,he stayedon territoryexcept that some birds foraged'on the beachesor the camparea while most of the tundrawas underihe snow.The Russianpapers reviewed by Grotereport the same. We saw many territorialdisputes between 16' June and 20 June in the area of the territories of nestsNos. 6, 7, 5, and 12 (Figure 7). Disputesconsisted of vigorousanswering song flights or a dashing pursuit,ending in a short,very fast flight, and such a rushat an intrud- ing, singingmale often ended in a songflight or rapid returnto the gr-oundwithin the territory. On 14 Junewhile the first Bell Heatherareas were emergingfrom the snow, three malesdisputed an area below the 35-foot beach and 25 yardseast of the site of nestNo. 6, whereterritories Nos. 6, 5, and 12 cametogether (Figure 7). One flew up from .the ground and dove at the others,or ran low, as if creeping,with his headlowered and thrust forward,"wiists" barely protruding, feathers sleeked (or ruffledonly on upperrump), andbill partly open,as he calledsput dirrrrr. He held his chin up, so that the black throat and light bill were conspicuouswhen facingus (Figure5). Onebird did nearlyall the rushing.He attacked the other two males when their side was toward him. The victim flew away with a shallow,quivering wingbeat ("moth flight"). A female was presentduring this posturing. We saw no actual fighting, ,but Michejev (1939, in Grote, 1943) describesfighting on the ground andin the air, bitingand pulling out feathers.It wasdifficult to seethe posturetaken by the bird rushedat, but thosewe sawheld horizontal

Figure 6. LAPLAND LONGSPUR FEMALE OF NEST N'O. 6. Showsthe close associationof the nest with overhanginggrowth of Heather. ]_4] Daui•¾,Bylot Island Breeding Biology Bird-BandingJanuary posturecreeping thrm[gh the grass. These rushes are typical expressions both of supplantingattacks and head-forwardthreat as Hinde (1953, 1954) andAndrew (1957) havedescribed them. We did not seean)- raisingor wavingof the wings,or bill snapping.These have been recordedfrom McCown'sLongspurs (Mickey, 1943; DuBDis.1937b),

• •"• PLOV,E,.,R PLATEAU

TERRITORIESESTABLISHD JUNE13-16 • PLOVERPLATEAU •,,

AFTERJUNE20 • 7 Figure 7. TERRITORIES OF LAPLAND LONGSPURS. A. Clutches completed June 22-25. B. C]utches completed July 1-4. Vol. •06•XXXII DauRY,Bylot Island Breeding Biology [ 15

Chestnut-collaredLongspurs, and other Emberizines. The note we recordedas accompanyingthe head-forwardposture seems to differ from the "chaa" reportedbv Andrew (1957). We have recordeda note"chreep" (below}, which must be thesame. We saw no territorial disputesafter 1 July, which may explain the disagreementsummarized in Frazer Rowell (1957) aboutterritoriality. Territory was expressed,but disappearedas incubationprogressed. Toleranceof Non-aggressiveTrespassing. Some birds did not show conspicuoushostility and in severalcases silent .males were allowedto trespassinto territory No. 6, our camp. On 26 Junethe male of territoryNo. 12, chasingthe femalefrom territoryNo. 12, cameover territoryNo. 6 and the chasepassed over the male of No. 6. Male No. 6 crouchedin a sleekedhorizontal position, loweredhis head, thrust it forward, bill up (Figure 5), and gave a vibrantchreep that soundedlike a beggingnote; thenhe roseat once into the air and sang.The pursuingpair wasalready on its way back into territoryNo. 12, and maleNo. 12 sangas soonas he crossedthe boundary.In contrast,on 27 Junethe pair from territoryNo. 12 moved throughthe territory of pair No. 6 to the vicinity of the cook tent, whichwas a favoritefeeding area. As theyflew over the owner, they gave the winterflock call, a rattlingchi-chi-chi-chi-chi, and this wasanswered at onceby theresident male, who did not chaseor singuntil theyhad movedon. Wynne-Edwards(1952) reportslack of hostilityon common ground.W•ithout further testing under favorable conditions in thefield, I cannotsay whetherthis is simplynon-attack when the trespasserdid not singor act aggressively(Tin'bergen, 1939), or whetherthe wi.nter call is a formalizedact of non-hostility.Kluijver (1951) and Hinde (1952) have describedfor the Great Tit (Parus major) a large area in that residen•tspecies in whichproperty interest exists but whichis not territoryin the limited sense.Trespassing is toleratedin the larger "domain."This principlelnay applyto longspurs,but I doubtit. In the periodof 22 to 28 Junethe pairs fro.mterritories Nos. 5, 6, 7, and 12 all visitedthe centerof camp,although pair No. 6 spentmost of theday there and the female of pair No. 6 becameso tame she would feed on cornmeal spilled between Van Tyne'sfeet while he preparedbird skins.Mickey • 1943) reportsthat McCown'sLongspur did n.ot defend itsterritory a•ainst other longspurs feeding or goingthrough.

TERRITORIES STUDIED On the warm,south-facing Bell Heather-coveredslopes north of camp, there were ten nestsand territoriesin an area of four acres (100 yards from nestNo. 7 to nestNo. 13, and 170 yards from nestNo. 3 to nest No. 4 {lCigure71. They were muchmore crowdedtogether than those recordedby Wynne-Edwardsf1952) at the head of Clyde Inlet, where nestswere 250 yards apart and territorieswere of five to fifteen acres. Grinnell (1944} foundnests at Churchillto be 200 yardsapart. Mickey 119431found no nestsof McCown'sLongspur less than 85 yardsapart. Territoriesin our camparea werea half-acreor less,probably a quarter- acre in the case of nests Nos. 5 and 6. Our territories are thus com- parableto thosestudied by Frazer Rowell t19571, or thosereported from Russianarticles reviewed by Grote (1943'l--Michejev reported 16] DRURY,Bylot Island Breeding Biology Bird-BandingJanuary

100 nestsin a squarekilometer of hummockytundra (their preferred habitat). The territoryof nestNo. 6 is an exampleof size,and locationof song perches.The male markedhis territorymost clearly in the few days after the femalestarted to incubate(4 July). It extendedup KungoHill about30 yardsfrom the nest (under the crestof the 35-foot,beach) to a songperch on a rock outcropat the 70-foot beach,then 10 yardsto the bank of SnowBunting Creek. From there his territory extendedalong the 70-footbeach 30 yardsto a barepatch on a shoulderof the hill above the nest. He had a secondsong perch on the bare groundthere and a third on the bare beachsurface 20 yards east of the nest. Thus his territory was a trapezoid,all sides with natural borders, 30 yards ,by 40 yardsby 30 yardsby 20 yards. He camein flight to sing over the westernmosttents of our camp,but did not perchthere and, (as above), allowednon-aggressive trespassing in that area. His territoryseemed to consistof an areaof ],000 squareyards. Evenif the areais extendedto includethe camparea, it covered1,800 to 2,000 squareyards--still less than half an acre. Territoriesof nestsNos. 1, 5, 7, ]0, and ]2, although lessclearly mapped because there was no pressureon at leastone border, were closeto the same size. The undisputed.borders had been along large snowpatches during territory estblishment.The twelveterritories which we studiedmost carefullyeach coveredless than half an acre.

COURTSHIP Pair Formation. The males' actions are similar {o those of McCown's Longspurs.When a male was associatingsteadily with a female,his singing decreasedin frequencythough not in vigor. This was not a cessationof song,but ratherdiffusion of actionsby additionalactivities. Pursuitflights were obviouslymutual affairs and, I believe,closely associatedwith pair formation.They were longer, slower, and with less zigzaggingthan teritorialskirmishes. If the male caughtup, therewas a burstof rapid zigzagging,but if he fell far behind,the femaleslowed downuntil he overtookher. We saw theseflights before any displays on the groundand throughoutthe periodof grounddisplaying. Male No. 6, displayingto his female,ran acrossin front of her or up to her side, standingat a'bouta 60ø angleto the horizontal,with his breast feathersfluffed out, headheld high and bill pointedslightly up, wings abouthalf spread,drooped and quivering.While he ran in this•way he wassinging his regular flight song. The female ran slowlyahead of him, crouchedin a horizontalposition with her headpartly lowered,wings partly spread,tail cockedjust abovehorizontal, calling zeep, zeep (Fig- ure 5). Sheand the maleoften pecked stiffly at the ground. Thesedis- playscoincide with thosedescribed by Andrew(1957), but he pointsout that in most Emberizinesthe male takesa horizontalposition in this situation. The chestnutnape "emphasized" the stiff bow by the male. The female ran around and ahead of him; then flew; if he did not fol- low, shecame back and repeatedher actionsuntil he did chase.During the chasesshe landedseveral times, ran alongthe ground,then flew again,fast and darting,or slowlyand on quiveringwings. Duringthe period of pursuits,because their attentionwas on the females,males sangless often and chiefly on the ground.Flight songswere conspicuous in the morning (0600 to 1000) and evening(1700 to 2200). Vol. 1961XXXlI DabmY,Bylot Island Breeding Biology [ ] 7

On 17 Junewe saw a femaleon three differentoccasions (or perhaps differentfemales), after beingchased by maleNo. 6, crossinto territory No. 12 or fly out of No. 6 into territory 7. The male did not pursue, and returnedto the groundbefore he sang. The male whoseterritory shenext visited usually took a shorttime to noticeher arrival, but when he did, he flew to her at once,singing (in the caseof maleNo. 12), and a pursuitfollowed. This pursuitmay havebeen territorial ratherthan sexual,or this may havebeen a vacillatingfemale, or we may havemissed some of the de- tails. Tinbergen (1939) was convincedthat suchsexual flights in the SnowBunting took placeonly with matedbirds, althoughhe observed onefemale to be pursuedby two males. I have no details on the changesin the male's aggressivepostures towardthe femaleafter pair formationand beforecopulation. We saw no signof courtshipfeeding. Pre-copulatoryActions. Abovenest No. 3 on 21 June (1800 to 1900), I saw the male rise with flutteringflight straightup into the air, flying with difficuhybecause of a wad of dark materialin his bill (Figure 5), a collection of leaves of Grass Rush, Black Mane Lichen (Alectoria nigricans[Ach.] Nyl.), and deadBell Heatherbranches. Before flying, he stoodvertically, bill horizontalor pointedup, wingsdrooped at his sides,and tail spreadand lowered. He sangwith thismaterial in his bill and succeededin gettingabout 20 feet off the ground,then sangas he fluttereddo;vn again. He pickedup evenmore material,and hopped onto a stone.He had so muchin his bill now that he couldnot get off the ground,but he couldsing. He stooda few momentsand readjustedthe materialin his bill, put it do;vnand picked it up again; thenhe pointed his bill up and flutteredhis wingswithout getting off the ground. This may 'be the wing wavingof Mickey (1943) or wing vibratingof An- drew t1957). Michel in Grote describesboth male and female flutter- ing up together.After severaltries he hoppedoff the stone,and walked and hoppedbetween clumps of Bell Heatherand Arctic Willow, picking up the darkestsprigs. The femalecrept hesitantly in a horizontalposi- tion throughthe grassas if feeding,but her tail wasslightly cocked up and her wingsoccasionally spread and quivering. Andrew (1957) sug- geststwo possibleorigins for this: (a) ritualizedfeeding action (out of context); or (b) ritualizedpicking up nestingmaterial (not out of context). When approachingthe femalethe male held his headhigh, neckex- tended and bill horizontal, with all the accumulation of dark material in his bill. He droppedthe load and pointedhis bill straightup, show- ing his blackthroat, while he draggedhis wings as he walkedup to her. Histail washalf spread and tilted down. Then he lowered his head, ruffled his scapular,back and rump feathers,and widelyspread his tail (An- drew's"fluffed run," 1957). She crouchedwith head low, tail cocked up, wingsquivering and partly spread. He flutteredover and, without hovering,mounted on her back in thesame position he hadassumed in frontof her,and copulated. After copulationhe hoppedoff andwalked in frontof herin thesame crouched position, while she stood up, raised herbill straightup and cocked her tail ashigh as possi'ble, chattering. 18] D}•uaY,Bylot Island Breeding Biology Bird-BandingJanuary

NEST AND EGGS Nesting;Sites. Although longsparswere among the early arriva]• their nestingsites in the great majority of caseswere under a patchof creepingBell Heather (Figure 6, nestNo. 6). Our observationssuggest as do Wynne Edwards' (1952) that one year'snesting is often near a previousyear's nestsite. Our nestsNos. 3, 4, 7, and3 wereplaced within 20 to 50 yardsof previousnests. Nests Nos. 3 and • were placedin Bell Heather, as were their associatednests. NestsNos. 4 and 7 wereplaced under Arctic Willow, an unusualsite at the Aktineq, and both associatedwith a previous year's nest under Arctic Willow. Male'sactions during nest building and egglaying. Songwas vigor- ous though ]essfrequent than before pairing. He answeredhis neigh- bors' songs,but usuallyhe directedhis songstoward her; while he followedthe female,who was gatheringnesting material and [eeding •eeming]y without attention to him. When she went to the nest while building or during the irregular periodsof incubationbefore the clutch was complete,he followed. When he lost sight of her during the egg- layingperiod, he beganto sing (as describedby Tinbergen(1939) for SnowBunting and Nice (1937) for SongSparrow). While shesat on the nestbefore the clutchwas complete,he wasseldom more than five feet away; but as soonas the clutchwas complete he stoppedfollowing her and madesong flights again over his wholeterritory, returning froin his flightsto a songperch rather than to her. He fed aimlesslywhile followingher, but not in the stiff way as when alarmed. Nest-Building.Females did all nest-building.The femaleat nestNo. 5, on 25 June,built the lining of her nestx¾ith white featherswhich she collectedwalking and hoppingalong amongthe GrassRush tufts and BellHeather on theedge of thebarren ground on the35-foot beach level. She,like other females,gathered nesting material inside her territory: deadgrasses for the'bulk of the nest,and white feathersor willow cotton for the lining, as Mickey (1943) observedof McCown's Longspur. Frazer Rowell pointsout that in somenests, the feather or cottonlin- ing may be absent.The femalehopped into a crack in the Bell Heather- coveredridge on the downslopeof a frost-crackon the edgeof the beach and into her nest,where she pushed the featheror clumpof grassesinto the side of the nest. Then she sat in the nest and wriggledback and forth with semicircularmovements similar to thoseused in settlingon the nest,kicking her feet and quiveringall over. I sawno forwardand backward movementof the bill ("tremble shove") to work the material in. After brieflywriggling in this way, sheflew awayto the upperedge of thebarren beach ridge. Onthe average,she returned to thenest every two minutes from 1400 to 1530. When she came from the nest she often stretchedher neck,spread the featherson her belly, exposingthe bare skin, spreadher wings,and cockedand partly spreadher tail. This agreeswith observedcorrelation of nest-buildingwith solicitingin Reed Bunting, Emberiza schoeniclus(Howard, 1929), and Snow Bunting (Tinbergen,1939). While shefed and gatheredmaterial for the nest (occasionallycalling cheeeplike beggingyoung), the malewalked around with headup and tail partly cocked,occasionally picking up browngrass only to drop it. Vol. XXXII 1961 D.[,l•. BylotIsland Breedit•g Biology [ 19 20] Daua¾,Bylot Island Breeding Biology Bird-BandingJanuary

TABLE III

LENGTH OF TIME IN NEST OF LAPLAND LONGSPUR EGGS

No. of days in nest before Nest No. 1 Nest No. 2 Nest No. 3 Nest No. 6 hatchin.g 5 (10%) 10 Individual 11 egg's number 6* 4 (12%) 12 (Numbered in 5 & 6 4 2 &3 13 sequence of 4 2 & 3 14 laying) 1 15 1 16 Table of the number of days that individually numbered eggs were in the nest before hatching. * Estimated by age of young; the day they hatched was missed.

They both gave low calls,inaudible beyond 20 yards: a reedy ,chatter, the winter rattlingcall, and the chip-chipflock calls. The male immedi- ately appropriatedas a perchthe pile of dirt from a test pit and sang every one to three minutes,or wheneverthe female hoppedinto the crackand ontothe nest. When the nestwas destroyed(presumably by an Eskimodogl on 2 July, the male stoppedsinging and the pair dis- appeared. Our observationson nestmaterial agree with Blair (1936), Grinnell (1944), Sutton and Parmelee t1955bl, and Wynne-Edwards(1952). Most nestswere tightly wovenof dead grass-likeleaves and stemsand were lined with a few feathers (two to about twelve) and ;viilow cotton (Salix sp.). Nestson the uplandswere made of grassesand GrassRush, and thoseon the edgesof marsheswere of sedgeparts. Early nestswere lined with white feathers;late nests(Nos. 6, 10, 12, and 13) were lined with both feathers and willow cotton. Whenthe femaleleft the nestduring the incubationperiod, the male joined her, or stayednear the nestwhile she was gone and calledin alarm whenan intruder enteredhis territory. Clutch-Size.Early nests(hatching 3-9 July) containedconsistently largerclutches (6, 6, 6, 6, 5, 4) than later nestshatching 10-15 July (5,5,4,4,4,4,4,3,3). Table I! summarizes the observed data on fifteen nests studied.

TABLE IV

NESTLING PERIOD OF LAPLAND LONGSPURS Nest Numbers No. Days 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 Young in Nest Total No. young Number of young 8 days 1 1 8 days in nest 2 No. young 9 days 3 4 3 2 4 2 1 4 2 2 1 9 days in nest 28 No. young 10 days 1 3 1 1 1 1 2 1 10 days in nest 11

Tableof numberof daysas nestlings. The youngwere not individually marked. Vol. XXXII 1961 Din,•Y. Bylot Isla;•d Breeding Biology [ 2 ]

Our egg-la3ingdates l first egg 17 Juneand clutchescompleted 22 Juneto 4 July) areclearly later than the dates of thepeak of firstegg datesin Fin•ark (56 p½•rcentbetween 11-20 June). as reportedby Shaaningin L0venskiold•1947), and in Clyde Inlet, Ballin Island (clutchescompleted by 6 June• by Wynne-Edwards(1952). They fall withinthe spread summarized in Frazer Howell t1957); andthey'come at the endof theperiod reported by Michejev(in Grote,1943): Timan Tundra,10-20 June: Murman Coast, 14-20 June; Kolyma region. 25-26 June; mouth of the Lena, 25 June to 4 July for full clutches.

ACTIVITIES DURING INCUBATION We haveno recordof malesincubating, although the stronglyde- velopedchestnut collar and blackish throat of several femalesmisled us. I)uBois(19371 shows that femalesof Chestnut-collaredLongspurs also may havepartially male plumages. The femaleat nestNo. 6 spentthree or four periodsof 15 minuteson the nestthree days before the clutchwas completed. She was on the nestabout half of the day on whichthe secondfrom last egg was laid, nearlyall of theday on whichthe next to lastegg was laid, andall day when'thelast egg was laid. FrazerRowell commented onthe start of incubation(sitting) with the layingof thefirst egg. Michejev(1939) reportsthe beginning of sitting(by thefemale alone) after the laying of the lastegg. DuBois(1937b) reportedthat incubationstarts when the last egg is laid in McCown'sLongspur. Mickey •1943) foundthat in onecase the femaleincubated all day ;•henthree eggs were in the nest althoughthe clutch of 4 wasnot completed until two days later. He found females"erratic" in respectto startingincubation. It wouldbe very interestingto keepa recordof the temperatureof the eggsduring the whole period. Malesand femalesstayed in their territoriesduring incubation. On 80 visits(after the clutchwas complete l to 14 nestswe foundthe female off 18 timesor two absencesper sevento nine visitsfor each nest (a consistentpercentage and not skewedby exceptional"absenteeism"l. FrazerRowell reviews the typicalbehavior of a femaleand male during the incubationperiod. Recognitiono! Damageto Eggs. While markingthe eggsat nest No. 13, we knockeda pieceof dirt into the nest. The femalereturned whilewe watched from 30 yards.picked the dirt out,flew 20 yardswith it, landed,and droppedit. Two eggswere punctured in marki.ngthem in nest No. 1, and these were both discarded by the same afternoon. At nestNo. 4 sixeggs formed the complete clutch ]from 25 June(when it wasfound) until 30 June. On 1 July onlythree eggs were present, and oneof thesewas damaged in marking.On 2 Julythe damagedegg had beendiscarded and the nestwas deserted 5 July. In contrastto these nestsin whichdamaged eggs were discarded(and none of the remains everfound), infertileeggs I oneeach in nestsNos. 2, 7, and 13, and three in nestNo. 10) wereleft in the nestuntil the youngleft or we removed theeggs t nest10). Removalof eggsduring the laying of theclutch and afterthe clutchwas complete had no effecton the numberof eggslaid. Resistanceto Cold. During the height of the thaw, 26-27 June, the bottomof nest No. 1 was soakingwet, the eggscold, and the female not seen. This wasthe fourth to sixth day after the clutchwas complete. 22] DRURY,Bylot Island Breeding Biology' Bird-BandingJanuary

The femalewas back on the neston 28 Juneand the four eggsthen pres- ent all hatched.Perhaps the youngbird with one foot undeveloped, which was founddead outsidethis neston 7 July, had its development damagedduring the periodof wetting,but otherwiseno complications resulted. IncubationPeriod. Incubationwas measured from the laying of the lastegg to thehatching of thelast egg (marked): 13 daysin nestNo. 1; 12 or 13 daysin nestNo. 2; 13 daysin nestNo. 3; and 101/2days in nestNo. 6. (SeeTable I for a summaryof incubationperiods.) The recordsin thistable support (by the incubationperiods indicated) our observationsthat the female spent more and more time on the nest as the clutch increased. DuBois (1935) found the incubation period in Chestnut-collaredLongspur to be 12•/5 days; Mickey (1943) found it to be 12 daysin McCown'sLongspur. Michejev reports incubation period of 10 days,but Grote in his review suggestsscepticism. Fertility of Eggs. We found 10.9 percentof the eggs(6 of 55) to be infertile. This is larger than the 5.3 percentreported by Sutton(1932) from SouthamptonIsland (53 nests,all with 6 eggs;in 17 oneeach did not hatch). It is muchsmaller than the 25 percentreported by Sutton and Parmelee(1955) from southernBaliin Island (of 97 eggs,22 did nothatch). Wynne-Edwards (1952) reportedthe hatching of all 29 eggs.

HATCHING AND CARE OF NESTLINGS HatchingPeriod. Eggshatched in the same24 hour periodin nests 3, 6, 7, and 13 and over a 48-hour period in nests1, 11 and 14, and possiblyover 64 hoursin nest2. Wynne-Edwards(1952) reported that hatching occupiedfrom two to four days, while Sutton and Parmelee(1955b) reportedthree nests hatching within a day (up to 5 eggs),and threenests (4 to 5 eggs)hatching over two days. Frazer Rowellreports one nesthatching within twenty-fourhours, and three nests over a period of ninety-six hours. DuBois (1935) found the hatchingperiod to be two daysin Chestnut-collaredLongspur. Feeding o/ Nestling& When the young hatched,the male shared equallywith the femalein feedingthe young. In nestsnumber 3, 6, 8 and 12 the male was more active than the female in feedingduring evening and morning. Frazer Rowell shows that the female was the steadierfeeder on the last day of the nestlingperiod. She actuallyfed more, althoughw-hen he was feeding,the male made as •nany visitsas she did. Frazer Rowellpoints out that in the first few days after the young hatch, the male does most of the feeding. Such changesin behaviormay explain the variation in male and female concernand feedingdescribed by DuBois and Mickes. Grote (1943) reviews15 Russianarticles, one of which reportsthat both parentsfeed the young 20-22 hoursa day, stoppingonly 2-5 hoursat midnight. At nest No. 12 the male fed the young as often as did the female duringthe day,and he seemedto be alonein feedingthem in the evening of 19 July whenthe youngwere 6-8 daysold. Between1915 and 2010 that evening,he fed them at the rate of onceevery four to six minutes. At nestNo. 6 the male alsofed the youngas regularly as did the female. Over a two-hourperiod (2000 to 2200}, when the young were 3-5 days old, the mate only was feedingthem and came on an averageof Vol. XXXII 1901 DauaY,Bylot Island Breeding Biology [23 once in 6 to 10 minutes. In each case the male foraged in his own territoryin the Bell Heatheron the edgeof the 35-footbeach. Michejev reportagrees with oursthat parentsgathered food usuallyabout 20-30 metersfrom the nest,seldom beyond 40 meters(yards]. Develomento[ Young. The younghad somedark brown down on the dorsaltracts when they hatched.They gapedat any disturbanceon the secondand third day. Their eyesfirst openedon thethird or fourth da.•,and the tips of their first backfeathers began to appear.By the fifth or sixthday, they lookedat an intruderand oftengaped. They crouchedin thenest during the last day or two. Our observationsagree with Suttonand Parmelee(1955b) that the nestlingsare essentiallysilent in the nest. We occasionallyheard a quaveringcheep as if a food-beggingcry from youngthat had left the nest. The loud chee-chee(or, as we wrote it--pseep) reportedby Suttonand Parmelee was given when we picked up the nestlingsin their last daysin the nest,or whenwe caughtthem after they had left the nest.•hen theyoung were leaving the nest, the parents were very concernedabout the first one to leave,but usuallyshowed only mild alarm if intrudersfrightened the last young from the nest and it hopped"on all fours,"stumbling and tumbling over the ground,pseep- ing in alarm. FrazerRowell offers the interestinginformation that all youngwere ready to leave the nestat the sametime, eventhough in one nest one eggmay havehatched four daysahead of the last egg to hatch. More thanenough food may providea partialexplanation of this. NestlingPeriod. TableIV summarizesthe numberof daysthe young spentin the nest. The great majority of our birds left after nine days in the nest. These nestlingperiods are consistentlyshorter than the between10 and 11 days recordedby Grinnell (1944• from four nests at Churchill,Manitoba. They fall within the limits •8-10 days) reported by Grote (1943) from Michejev. In contrastto thesereports of longer nestlingperiods from more southernareas, our observationsagree closelywith thoseof Wynne- Edwards t19521 and Sutton and Parmelee I1955b) that the nestling period is 9 to 10 days. All these observersfound, as we did, that the young left the nest 3 to 5 days before they could fly. Grinnell (1944) and Nicholson (1930) indicated the same without comment. Frazer Rowell'schart doesnot allow me to analyzethe historyof all nestlings individually,but indicatesnestling period 8-11 days,chiefly 9 days, if I read it correctly. DuBois (1937a) found the nestlingperiod in Chestnut-collaredLongspur to be about 10 days; and Mickey (1943) and DuBois (1937bt found it to be 10 days in McCown'sLongspur. Leavingthe Nest. Wynne-Edwards(1952) reportedthat the young left the nest within 48 hours of each other. Our information includes four nestsin which the young left the nest within 24 hours of each other, and fi•e nestsin which the youngleft within 48 hours of each other. Our observationsagree with Frazer Roswellthat the nestswere left clean,not imariablv foul as HaGland (1916) reported. Frazer Rowell reportsin detail the behaviorof the femaleand youngat the time of leaving. 24] Dnu•Y,B•'lot lsla•,d Breeding Biology

TABLE V

LAPLAND LONGSPUR NEST SUCCESS Eggs for whose Young lost Nest Total damage we were Addled Young through presumed Young No. eggs responsible Eggs we took natural causes produced 1 6 2 1 3 2 6 1 1 4 3 4 4 4 6 1 (6) Deserted, presumably after disturbeelby dog. 5 4 4 Destroyed by dog. 6 5 5 7 5 1 3 •clisturbed ?) 1 8 6 1 5 9 4 4 10 5 3 2 11 3 3 12 4 4 13 4 1 3 14 4 4 15 3 1 2

Total 69 13 6 2 4 44

We have no accuratedata on how soonafter leaving the nest the fledgedyoung can fly, but it wasabout 3-5 days. Youngstayed in their parents'territory for thesedays and then drifted throughother terri- toriesto gather,as Frazer Rowellsays, in dampareas. Grotereports that they gather in hummockyareas while they can only fly weakly and are somewhatprotected from heavypredation bv harriers,merlins and jaeaers in the hollows. Once strong fliers, the young are less exposedto predationand moveto brushyplaces by pondsand streams. They are completelyindependent at 10 days. Nesting Success.Table V summarizesthe nesting successof the 15 nestswe studied. We found a total of 69 eggs,of which we or a dog (there becauseof us) destroyed10. Of the remaining 59 eggs, 6 were infertile, and 5 abandonedbecause of disturbanceby the same dog; one youngwas deformedand pushedout of the nest; and 3 young disappearedfrom a nest (whichwas six feet from a well-travelledpath at camp). In total, 42 youngleft the nest; we took two specimens.Eight •perhaps9) of the 15 nestssuccessfully fledged all youngfrom the full clutch. Thirteennests, or 86 percent,had one or more eggshatch, and four was the averagenumber of young that hatchedper successfulnest. Seventy-fivepercent of the eggsproduced fledged young. Grinnell (1944) reportedthat of 12 eggs,6 nestlingsleft the nest. Suttonand Parmelee(1955b) reportedthat of 75 chicks,62 left the nest successfully.Wynne-Edwards (1952) reportedthat of 29 eggs, all hatched--5 young were abandonedwhen wetted, 1 died, and 1 was killed accidentallyafter it left the nest. Presumably22' youngleft the nest.

REACTIONS TO PREDATORS To Humans. The first indicationof the presenceof a nest,and a con- stantaccompaniment to the nest-checkingrounds. was the male'salarm Vol. 1965.XXXiI D•t•. By/orlslat•d Breeding Biology [25 note--a wheezydyew or dzeeu. It wasgiven in the presenceof a human intruderand was givenby severallongspurs when a Peregrine(Falco peregrinus}flew over. Suttonand Parmelee(1955b) recordedthis note as whee-yeeor ear. Grinnell (1944) recordedthis note at Churchillas bee or a metallic kittyoo. In GreenlandNicholson (1930) reported a shrill "pipe" l usuallydouble). This seemsto be •norevariation than expectedfor different ears hearing the same alarm call, but in each area the alarm note is reportedto be consistent.We have heard the same note on the wintering grounds. and the bird takes the .same head-upposture when giving the call there. The male's alarm was soundedfor his territory rather than specificallyfor the nest, since no increaseof concernwas shownas we approachedthe nest, but he hoppedto an exposedperch or fed in a formalized,nervous way within 15 to 20 yards of the nest. Males often flew hesitantly,low over the tundrafrc;m place to placein thissituation. If the femalewas on the nest. his concern was much more marked and increasedif one ap- proached the nest directly. These actions agree with what DuBois (1935) found in Chestnut-collaredLongspurs. and in McCown'sLong- spur (1937b). Femalesvaried in tmnenessat the nest,but on the averagethey would flush from it at about 20 yards when we came to the nest directly. They usuallyflushed at aboutfive yardswhen the nestwas first found. Then they flew about20 yards low over the ground, as often in straightaway flightas in a quivering.hesitant flight. When they lit, theyusually shook themselvesand started to feed nervously,occasionally stretching their headshigh to look at us. Irregularly under these circmnstances.the femalewas heard to call a rapid pitze-pitze,which is probablythe note recordedby Nicholson 11930}. On a very few occasions,the female hoppedand fluttered over the ground. jerking her wingsand spreading her tail as if "injury-feigning." Grote'sreview (19•3) agreesthat the malestands watch while the femaleincubates. but he suggestsmuch more conspicuousdistraction display for both male and femalethan we saw. The female at nest No. 2 was very tame from the first, and allowed us to approachwithin six feet. The femalesat nestsNos. 5 and 6 were equallytame, especiallyafter severalslow approacheswere made to photographthem. After this,they, too. allowedapproach to within six feet. Nest No. 5 wasdestroyed. but No. 6 went on to successfulcomple- tion. and toward the end of' incubationthe female sat until we stretched a hand out over her. To Birdsand Mammals. Tinbergen (1939) suggestedthat the noisiness of SnowBuntings while in the nestin Greenlandmight reflecta lack of weaselsthere. Longspurs,like SnowBuntings, were essentially silent in the neston BylotIsland. where weasels were present. On 25 July we watcheda fernalelongspur, presumably from nestNo. 10,being pu•'sued by a Long-tailedJaeger (Stercorarius longicaudus). As describedin our accountof the jaeger (Drury, 1960), one made a seriesof passesat her whileshe fluttered upslope and downslope,back and forth acrossthe river. She had increasedtrouble with the steady wind and finally dashedinto the large stonesin the bed of the river. Afterhovering a minuteor two overthese rocks, the jaeger turned and glidedoff. 26 ] Dl•U•Y,Bylot Island Breeding Biology' Bird-BandingJanuary

FEEDING Longspursfed primarily on the edgesof densevegetation or in the areasof sparseclumps of plantson themargins of barrenspots. They fed almostcompletely on the ground,creeping, walking and hopping,and pickingtheir food off the dirt. DuringJune their food was chiefly plant (probablygrass and sedge)seeds. When they startedcollecting food after the younghatched, they searchedthicker vegetation, and many col- lectedfood along the frost-heavedridges in the boggyareas or in the thick Bell Heather and Net-leaved Willow (Salix reticulata L.) areas in sheltered,moist areas under solifluctionlobes on raised beach edges. Grote (1943) describesthe shift from grassseeds to insectsas insects becomenumerous and activethrough the season.He quotesfrom Miche- jev whostudied food brought to young:chiefly Diptera and Hymenoptera and only a few spiders. SNOW BUNTING Plectrophenaxnivalis (Linnaeus). (Eskimo: Kah-6h-dluk-tah). HABITAT SnowBuntings nested in steepravine sides,overhanging creek banks, and barren ridgesnearly free of plants. In contrastto the reportsfrom Greenland,and central and southernBari:in Island, this specieswas not the mostcommon bird, and it wasless widely dispersed than the Horned Lark in the plateauarea on the southwestpart of Bylot. Our surveys showed31 pairs (Van Tyne and Drury, 1959).

DISTRIBUTION AND DENSITY We found four nestsand four more territoriesin our studyarea: One territory was on the steepface of the West Ridge at the head of Golden PloverCreek; one eastof the mouthof the Little River; and two along the cut ba,nksof the Aktineq River at PtarmiganFlat (perhapsnest No. 5).

TERRITORY Many SnowBuntings were singing at Pond Inlet on 11 June,and we saw one territorialsquabble. On Bylot 13 June,Snow Buntingswere silentand we did not seeterritorial activity for severaldays. Tinbergen(1939) showedthat oncepairs are formed,song drops to a minimum,unless the femaledisappears. We arrivedin the later part of hisfourth period--paired but not yet nest.building. Song. The singingof two malesat campwas at its maximum20 to 27 June,and indicatedcontinuing territorial competition as late as the eve- ning of 2 July. This was,we assume,the recurrenceof songwhile the femaleis laying and whenshe starts to incubate.Two malessang while accompanyingfemales carrying nesting material, on 23 June. After singinga while from a perch,the malessang as they left or arrived back on their songperch (Figure 2). They sang in the air if they met a femaleor anothermale. Therewas marked individuality of songs.Of these,the male from nest No. 4 and that from nest No. 2--immediate neighbors--werethe mostsimilar. The male from nestNo. 3 sanga consistentsong of prrr-tsa-tsu-tsee,prrr-tsa-tsu-tsa, prr-tsa. Songsbecame abbreviated in late Juneand were shorterthan we ex- pectedat all times. Presumablywe did not hear the full songsof the earlypart of thecycle. Male No. 3 sanguntil 8 July, but disappeared whenthat nestwas destroyed. Vol. XXXII 1961. Dau•tY,B3/ot ls/a•d Breedb•g Biology [27

AGGRESSIVE BEHAVIOR We saw territorialfights on the Bluffsnear nestNo. 4, and between nestsNos. 2 and 3. The malefrom nestNo. 3 avoidedgoing westof the right bank of SnowBunting Creek where his femalebuilt her nest. He had a songperch on a largerock three-quarters of the way up the east (or left) side of SnowBunting Creek. Themale at nestNo. 4 ignoreda singingmale pipit and a HoaryRed- poll (Acanthishornemanni) which perchedand twittered within ten yardsof him on the stonyridge, and later took his perchand chattered whenhe flew downto chaseaway a malebunting from the west. Other territorialmales ignored singing longspurs and shorebirds.

LATE FLOCKING On 14 June,a flockof two maleand threefemale Snow Buntings, four male and one female Lapland Longspurs,and two Horned Larks fed in the grassymeadow between the generalsnow cover and the top of the Bluffsjust westof camp. The vegetationwas chieflyGrass Rush, Sweet- grass,Arctic Bluegrass(Poa rigensHartre.), and Water Sedge •,Carex equatilisWahlenb. var. stans [Drej.] Boott), with patchesof mosses, Averts,and Arctic Willow.

COURTSHIP While snowcover was general,Snow Buntingsfed togetherin pairs, and inspectedsheltered holes wherever they couldbe found. On 30 June while the femalewas off the nest, male No. 3 sang sporadicallyand preenedon his songperch. Shefed alongthe banksand whenshe came intosight, he glidedfrom his perch down to her andcalled a weakzzeeeep. He followeda while,then she hopped in front of him and cried wheezily, openedand raisedher bill, and waved (rather than quivered) her wings (Figure 2). Presumablythis was food begging.There were 4 eggsin her nest.

NEST AND EGGS All the nestswe found were in hollows, cracks, or tunnels excavatedin silt or sandybanks with one exception--undera crackedboulder. Our previousreading led us to expectnests in pilesof stones.All siteswere free of snowby 15 June. The lack of suitablesites may be responsiblefor the relativescarcity of SnowBuntings at the Aktineqas compared with the publishedreports from otherareas. Very few suitablerock pileswere presenteven in the Bluffs which were of friable and poorly consolidatedshales. and sand- stones.Many authorshave suggestedthat SnowBuntings are attracted to a permanentEskimo camp. On 3 July we foundtwo nestswithin five yards of each other in the stonewall of the nearestpermanent Eskimo house,on the shoreat Sermilikseven miles east of camp. We saw a femalecarrying nestingmaterial on 13 and 23 June, and founda partlybuilt neston 20 June. We foundthe first completedclutch on 23 June. Anotherclutch was completed 3 July. Of the full clutches we found one contained5 and two contained6 eggs. 28] D.•:,Y,By/or lsla.d Breedi;k. o'B/ology Bird-Ban•lingJanuary

ACTIVITIES DURING INCUBATION Attentivehess. We saw the female of nest No. 3 leave the nest to feed during the morning t'10-20minutes between 1000 and 1200), afternoon 15-12 minutestwice between1630 and 1800•, and one about midnight. Parmelee (Sutton and Parmelee, 1954,b) observedthat the male fed the female on the nest. We did not see the male ever go into the nest. When the female returned from a feeding excursion,she flew fast and directly, lit a few feet from the hole, and went in--quite in contrastto her aimlessmovements while feeding.

HATCHING AND CARE OF NESTLINGS In the only successfulnest we studied.all young hatchedon 6 July. They stayedin the nest12-13 days. B•)thparents fed the young, which were usually silent in thenest. We heard them chatter only when the parent actually was in the nest. Tin- bergent1939• saysthat in EastGreenland. young chattered loudly while the parentswere away. Oncethey had left the nest.the youngchattered hoarsely and loudIx' whenfed, and had a characteristicpit or sw#-switnote. We foundtwo youngwith a male at the head of Lark Gully. when the femalewas with the other young at the nest. This indicatesdispersal and parentalco- operationin care of the young.as Tinbergenmentions. The sameday. and for threedays afterward, 18-21 June.there were two pairsof parents and eightyoung in the stonycreek-bed on the westside of West Ridge.

REACTIONS TO PREDATORS Snow Buntings flew endlessIx'around and around with hesitating flight,low over the ground. They settled briefly and fed stifflyand ner- vously;then flew on again. perch{ng on exposed rocks. The female called cheepoccasionalIx'. Usually the female was more demonstrative,and the malefollowed her or remainedperched on a rock and occasionallycalled pirrit. The male actedas concernedas the femaleif shewere absent. On 28 Junethe male and femalefluttered doggedly after a weaselas it wentup the westbank of SnowBunting Creek and movedbetween their nest and the snowbank. The female fluttered so close when the weasel was near the nestthat it rushedher twice. The nestwas destroyedby a weasel.

FEEDING Snow Buntingsfed on the edgesof barren beaches.ridges. or steep slopeskept openby frost action. When feedingthey creptalong through the sparsevegetation, especially among clumps of grasssedge and rush. Most of their food was taken from the bare surface of the soil. but they alsopecked repeatedly at last vear'sgrass heads.

PART II: DISCUSSION OF ECOLOGY

/. SPECIES SEGREGATION AND DISTRIBUTION RELATIVE TO SUITABLE HABITAT. What elementsof habitatsegregation allow thesespecies to exist to- getherwithout competition for territory. cover,or food? Vol. XXXlI 1961 Daut•¾,Bylot Island Breeding Biology [29

1. In plants,there are seldomnumerous representatives of the same genusin onehabitat which is subjectto environmentalextremes; and we foundthe sameto be true for the birdsof BylotIsland. SnowBunting-- LaplandLongspur, and Black-belliedPlover--Golden Plover, seem to con- tradictthis. But alsoit is true in plantsthat whentwo membersof a genusdo occurtogether in the high north,they are closelyrelated--as in Saxijrga, Ranunculus,Salix, Draba, Antennaria, and Potentilia. When organismswidely separated from eachother converge evolutionarily on a commonhabitat, their manydifferences of structureand behaviorwill tendto keepthem from competition,as Darwinpointed out. 2. We sawno overlapor competitionfor food amongthe Passerines. In general,longspurs and pipitsfed in thicker,moister vegetation; and larksand buntings in drier sites.Pipits fed mostlyon insects,on or fly- ing abovethe vegetation.The othersfed on materialin or under the vegetation,longspurs in or belowmossy, heather-grown areas, and bunt- ings and larks in sparse,grassy areas. Our few observationson food broughtto the youngindicate that they werefed manytwo-winged flies and spiders.Presumably, this reflectsan abundanceof supply. All exceptlarks readily acceptedtrespassing by birdswho showedno territorialbehavior. This actionand the over-ridingeffect of flockcalls are valuableadjuncts to allowa sectionof the populationto crowdto- getherin hard times. 3. The four speciesdiffer most conspicuously in selection of nestsites: warm,sunny ravines for the pipit; exposedbarrens for lark; well-vege- tatedslopes for longspur;and holesin exposedridges or rock pilesfor bunting. Theseindicate complete ecological segregation even though therewas no shortageof thelast three types. 4. Larks had large territoriesand tendedto spreadout over wide areasof theuplands--not concentrating in favorablevalleys as did pipits and longspurs.This differencein dispersalreduces competition. The sunny,steep valleys occupied by pipitsdid not overlapthe lower sunny slopeterritories of longspurs.Snow Buntingswere widespread,like larks,and in our areawere influenced by restrictednesting sites so that thetwo seldomeverlapped. 5. Noneof the speciesoccupied the totalarea of habitatsuitable for it. LaplandLongspurs were most conspicuous in this. They werecrowded into certain places (mosaicdistribution), and occupiedonly a small percentageof the total suitablearea. There are at leastthree reasons for this:First--The birds settled on areasfree of snowwhen they first ar- rived (Figure2 in Van Tyne and Drury, 1959). Theseplaces were on south-facingslopes at low altitudesand werenot widespread.Second-- Mickey(1943) commented(McCown's Longspur) on the aggregation of nestsin seeminglyuniform grasslandhabitat. Svardson(1949) dis- cussedthe aggregationof Wood Warblers(Phylloscopus sibilatrix) at thelimits of theirrange in Sweden.He andAndrew (1957) agreethat the songof a malenot only attractsfemales, but alsoattracts other males to set up neighboringterritories. The natural behavior then leads to aggregationsof territorial malesover and above that forced on them--in our caseby generalsnow cover. Third--We found severalnests asso- ciatedwith a previousyear's nest (especially those in whicha peculiarity of constructionindicated that they were madebv the samebird•. This 30] Dinroy,Bylot Island Breeding Biology BirdJanuary -Banding iuggeststhat femalestend to returnyear after year to the samesmall territoryin thisspecies as in manyothers. H. COMPARISON OF ARCTIC ADAt•ATIONS OF PASSERINES. -What are the featuresof their biology which allow thesepasserines aloneto occupythis area? A. The behaviorof the LaplandLongspur which enables it to occupy the tundra habitat is sharedin detail, as far as I can discover,with the other longspurs.Indeed, the lundra at Bylot Island resemblesthe topography(rolling uplandsand deep-sided,sharply-cut river valleys) andvegetation aspect of thegrasslands of the highplains. Territorytype, songdisplay, restriction to territory,female incubating before the clutch is complete,short nestlingperiods, and young leaving the nest before theycan fly, are sharedwith McCown'sLongspur, and manyare shared with Chestnut-collaredLongspur; yet they are featureswhich would be assignedas adaptationsto the arctic habitat were it not for the excellent studiesby Mickey (1943), and DuBois (1935, 1937a, 1937b). 1. Toieranceof crowdingis of advantageto the species in a year suchas 1954,because thereby many pairs were able to breedsuccessfully whenonly a smallarea was available at thephysiologically proper time for nest-building. 2. With Lapland Longspurs,one adaptationis physiological.The toleranceof the eggsin nestNo. 1 to soakingfor two days (26-27 June) in ice melt-water is remarkable,allowing successfulreproduction in areaswith a "late" spring. DuBois (1937b) tellsthat beingburied two days under the snowkilled the eggsof a McCown'sLongspur, which extendedincubation nine daysbeyond the normal period. Birula (1907• describesthe destructionof LaplandLongspur nests by melt-waterfloods andby latesnowstorms in Siberia. 3. The ability to arrive on the breedingground and then feed for a week or two until suitablenesting sites becomes available, even though thisis afterthe peak laying periods in moresouthern parts of thespecies' range, contributesalso to local success.This feature of arctic nesters wasdiscussed by Lack (1933), and treatedin detailby Marshall (1952). Grote (1943) mentions it in his review of the Siberian .breeding of Lapland Longspurs. 4. The short period taken for the nestingcycle is anotherimportant quality,using longspurs as an example:eggs hatched 3-15 July, and the youngstayed in the nest8 to 11 days, and left still unableto fly. This concentratesthe breeding cycle (a) by shorteningat the beginning in responseto a late thaw. and (b) by shorteningtoward the end by successfulindependence of fledgedyoung. Presumablythe ecological pressuresdiscussed by Pitelka, Tonrich. and Treichel t1955) are im- portantin hurryingtile later end of thecycle. Those authors discussed the need for the predators'young to becomeindependent while food is abundant.The prey of longspursis soft insects,rather than lemmings. Nice i1937) showedthe influenceof temperatureon the start of laying in Song Sparrows.and since her studies,others have shown similar effectsin other species.most recently Snow (1958) in the European Blackbird (Turdus merula). This, coupledwith effectsof stimulationb3 the male and nestingsite. prestonably controls the start of femaleslaying. and decreasesthe chancesof her starting "too soon." Vol. 1961XXXI! Dm-aY,Bylot Island Breeding Biology'

5. The differencein clutch-sizefrom thesouthern portion of the range, Churchill, Manitoba---4-5eggs (Grinnell, 1944), to the northern part --6-7 eggs (our records,1954; and thoseof Wynne-Edwards,1952), conformswith the usual tendency;but we notice that in McCown's Longspurthe total annualproduction will be greatersince, as Mickey (1943) says,they tend to raisemore than onebrood a year of 3-4 eggs. The averagefor McCown'swould be sevenyoung per year which is the unusual•naximmn for LaplandLongspur. At the southernpart of the breedingrange of LaplandLongspur, there are nearly 20 hoursof daylightwhich is as much as our longspursactually used (Palmgren, 1935; Grote, 1943; Franz, 1949; Karplus, 1952; Hoffmann, 1959). Becauseof the adequacyof time "to feedmore young" where the clutch- sizeis smaller,and the greaterannual production in two-broodedmore southernspecies, it is hard to seethat t'he"reason" for LaplandLongspurs nestingso far north is that they are able to producemore young. The period during which the youngwere in the nest (8-10 days) is not decreasedin the Arctic as comparedwith McCown'sand Chestnut- collaredLongspurs. Interestingly, we found the sametypes of birds I larksand plovers ) sharingthe tundra habitat that Mickey (1943) found sharingthe grasslandswith McCown's•Longspur. The sameideas apply to larksand SnowBuntings, but it is hard to see how a bird so dependenton insectlife as the pipit surviveslate snows at this place. Our observationsindicate that pipits are at the extreme of their tolerance,but suggestthat larks are capable of nestingmuch farther north. The nestsites, behavior, and feedingof larks suit them at leastas well as longspursand Snow Buntings;yet they do not go further north while longspursgo north to EllesmereLand and Snow Buntingsare reportedto range over all land surfacesfree of snow, farther north than any otherland bird. The differencemay be related to the degree of dependenceupon insect food when they arrive on thebreeding ground. I find no obviouscharacteristics of productivitywhich make it ad- vantageousfor longspursto nest in the Arctic. Their various habitat- correlatedbehavior patterns are equallysuited to grasslandareas to the south. Certainlythe ability to maintainphysiological preparedness until thefinal stinmlus of suitablenest sites appear, together with the con- centrationof the breedingcycle into one brood, are factorswhich allow this speciesto nestfarther north where it lackscompetition. Probably the explanationis that by nestingin the Arctic the speciesescapes com- petition-the tool of selection--andany specieswhich can live there automaticallyhas an advantageby lackingcompetition. We shouldnot look. then, for the reasonswhy the specieshas an advantagein living in the Arctic. but look for the features which allow it to live there. Most importantof all is the fact that they do occur there. Historical accidentsand sourceof populationto occupythe area haveled to their presence.Once an organismoccupies an area, we may show why it can. but we have no hints on why, for instance,the closely related other longspursor pipits or buntingsdo not occupythe area. To say they are excludedby competitionis no answerunless we have details. B. All four specieshave well-developed flight songs.But they readily sing from elevatedperches, Horned Larks from telephonepoles, and 32 ] DRURY,Bylot Island Breeding Biology Bird-BandinLJanuary

SnowBuntings and LaplandLongspurs from radio mastsat Pond Inlet. C. All four specieswalk and have a well-developedhind toe with an exaggeratedlylong claw. All four live on treelessareas where the wind ,blowsmost of the time. Having no shelterfrom sucha wind, smallbirds will needfirm footing,and the additionallong hind toe shouldsupply this. Anotherpurpose might be to supplya largerfoot for walkingon the snow.This correlateswith theirwalking rather than hopping--anadvantage on a soft surface.In my experience,tracks of larks and SnowBuntings on the snowdo not sink as deeplyas do those of Tree Sparrowsor SongSparrows. Walking or creepingalso avoids bufferingby thewind. HI. CLUTCH-SIZE /IND L/ITITUDE Lack'swork on clutch-size(1947, 1948a,1948b, 1954) is the founda- tion of our knowledgeof this subject,but I am unsatisfiedwith his explanationsof the large clutch-sizein the northernpart of a species' range. It is clearthat if the phenomenonis presentboth within popula- tionsand betweenseparate species, there must be a selectiveadvantage. On the other hand, our observationssuggest doubt that the larger clutch-sizeresults in larger annualproduction. How do the clutchsizes and total annualproductivity of thesespecies vary with latitude? For example:Information available in Bent(19}2) for HornedLarks r•rovidescomparison of the clutch-sizeof this speciesover its North American range--from the hot desertsof northern Mexico north to our Bylotnests, which are the farthestnorth recorded.The subspecies from thehottest part of the rangehave an averageclutch of threeeggs: (act& (2-3-5), adusta (3), enertera (3), merrillii (2-3, rarely 41, occidentalis(3}; the subspeciesof the westernand coastalgrasslands have a clutchof 3-4: insularis(3-4), giraudi (3-4}, leucolaema(3-4}, strigata (3-4); wetter prairies and the northeast usually have four; alpestris (3-4-5), tiara, Old World t2-4-5), praticola t2-4-5); the northernbirds, of whichonly hoyti is adequatelyrepresented, have 4-5. The annual productionof individual femalesis actually not larger in the north than in the two- or three-,broodedpopulations farther south. The clutch-sizeseems instead to be a telescopingof the breedingseason. The followingare clutch-sizesfor LaplandLongspurs: Hatched be- tween3 and 9 July--4 (6), 1 (5), '1 (4). Hatchedbetween 10 and 15 July--2 (5), 5 (4). 2 (3}. We found no nestswith clutch-sizeof 7, whichWynne-Edwards reports from the headof ClydeInlet, but he found completedclutches 6 June. Perhapsthis is because1954 was a conspicuouslylate season (we found the first completedclutch on 22 June). If the rule appliesthat later nestshave smallerclutches. our nestsfall into the patternhe found,because our nestsshould be com- pared with his later nests(completed 19 June to 2 July and containing three clutchesof 5 and two of 4). On this bas;•s,our earlier nests have larger clutchesthan the contemporaneousnests at Clyde Inlet. but the latitudinal distances are too small to be as effective as seasonal differences in influencingclutch-size; and the fact that early clutchesare laid by adultsand thus are larger than later clutches presumably laid by first- year females,must be taken into account. The clutch-sizein theset;•'o placesis larger than that reportedfrom FrobisherBay by Suttonand Parmelee (1955b): 3 (3): 9 (4}: 9 (5): 1 (6). Sutton 11932) found Vol. 1961XXXII DauaY,Bylot Island Breeding Biology' [33 mostnests tall but 7 of 78 nests)on SouthamptonIsland (64 ø N. Lat) to contain6 eggs. Grinnell's (1944) four nestsat Churchill,Manitoba {about 59 ø N. Lat.), contained4 to 5 eggs. Blair (1936) reported3 (7) and 1 (6) froin East Finmark (70 ø N. Lat.). Frazer Rowell (1957) reported1 (4), 7 (5), 3 (6), and 3 (5+). The clutchesof Snow Buntings1 (5) and 2 (6) were smallerthan thosereported from the northern part of their range and they were later than usuallyrecorded. Pleske (1938) reported5 to 7 eggsfrom Taimyr Peninsula,Siberia. Nicholson(1930) reported1 (7), 2' (6), 1 (5), and 3 (4) from southernGreenland. Manniche (1910) reported that the mostfrequent clutch-size in northeastGreenland was 5 or 6, seldom4, and only onenest of 3. Suttonand Parmelee(1954b) report: 1 (4), 11 (5), 1 (6). and 3 (7) froin Frobisher Bay, southernBattln Island. Tinbergen(1939) reported:1 (3), 2 (5), and 4 (6) from east Greenland. Tinbergenreported most clutchescomplete by mid-June;the nests Suttonand Parmeleefound after 19 Junewere alreadycompleted; and Wynne-Edwards(1952) reportsyoung after 2'5 June. Why larger clutch sizes in the north? Recoveryfrom Disasters. For a speciesto occupya hugearea of productivehabitat, its breeding biology.must allow it to meetthe vicissitudesof that habitat,and it is obviousto thosewho visit the far north, that heavy mortality comes from periodic disasterson migration or accidentson the breeding groundssuch as late snowstorms.Natural selection can be expectedto have detected a factor obvious to a human. For natural selection to act, however,it must act .on the advantageswhich allow the populationto occupythe favorablehabitat, not to •balancemortality, as Lack (1954) pointed out. In the north, speciesmust have a breedingpotential to allow re- coveryfrom periodiccatastrophes, and thereis selectiveadvantage of largeclutch-size to allowrapid recovery of the population.In contrast, in stableand uniform habitats,and especiallywhere there are resident populations,too many young in eachnest will ,beselected against, since it attractsspecialized predations and parasitism, and leads to competition amongthe membersof each large brood. The studiesavailable on breedingpotential suggest (viz., Lack's (1948b) work on Starlings, Sturnusvulgaris), that a specieswill producejust as many youngas it possiblycan bring through to ,naturity.Absence of a secondbrood, and lack of competitionfor abundantfood when the youngare 'becoming independentand the parentsare molting,allow arctic species to bring moreyoung through. Larger broods will be successfulas longas they producemore young to migrate.Large clutch-size has the advantage of allowingrecovery from disasters--butthat is not enough.How is the clutch-sizeenlarged, and why are clutchessmaller farther south? Is productivitygreater? Longer daylight alone is notthe explanationfor the largerbroods in the north,because of the daily cyclesof activity weand many others have o,bserved (Pahngren, 1935, 1949; Franz, 1949; Grote,1943; Karplus,1952; Hoffmann,1959). Birds at 74ø latitude useno moredaylight than those at 55ø latitude(20 hours),and across this latitudinal range. the longspurs'clutch-size varies "accordingto the rules." 34] DRURY,Bylot Island Breeding Biology Bird-BandingJanuary

GeographicalVariation in Clutch-Sizeand Daylength. In the geo- graphicalvariation of clutch-size,it is important to separatereleasers from selectiveadvantages. There is abundantevidence of larger clutches in the north and an obvious environmentalco-variant is daylength. Our longspurnests and thoseof Wynn-Edwardst1952) showthat within limits, the later t.he start of the clutch,'the larger the clutchuntil a peak is reached,and then the clutch decreases,and this is known for many species.In the caseof Kent's Island Tree Swallows(Iridoprocne bi- color)--(Paynter, 1954), later arrivals find a longer length of day than birds nestingto the south. If birds arrive later in an area slightly farther north, they find •nuch greater differencein daylengththan ex- pected,on a latitudebasis. I am convincedthat this is the "releaser." The length of day is one of the stimuli which startsthe mechanismof egg production,but it doesnot necessarilycontrol the upper limit of clutch-size. Nice (1937), Snow (1958), and others have discussed temperatureas a stimulus. I doubt that the reasonthe clutch is larger is ability to feed all the young. Lack's data on the clutch-sizeof the EuropeanRobin, Erithacus rubecula(1953) offers evidenceenough itself to throw doubt on this. My data on HornedLarks agreethat it is misleadingto apply only this hypot.hesis. The variations approximatelyfollow latitude or daylength, but many.other factors vary with latitudein the sameway. Large brood size is not involved in the most .characteristic of northern breeders-- the shorebirds.Long day is just one factor obviousto man. I suggestthat large clutch-sizeis a secondaryfactor; that in the north selectionoperates on concentratingthe brood into one annual nest. Ray (1913) suggestedthe sameeffect for Tree Swallowsat higher altitudesin California. In the south,as with larks,it is an advantageto spreadbroods over a long period--two or three broodsof two or three young. The young would not survive a breeding seasonof March- Septemberon Baffin Island, and thus larks there can have only one brood. Selectionis not simplyof large broods,but of a singleannual brood, which must be large enoughto maintainthe population.I sug- gestthat there are inherentlimitations to total annualproduction within the speciesand that theselimit the sizeof the singlebrood, becausethe total annual productionin the desertsof Mexico is the same as that in BaffinIsland. In studiesof the selectionof broodsize, annual production must be consideredas a factor; i.e., average clutch-sizeX average nmnberof broods. In the far north, productionis suitedto the short season,but productionis not necessarilygreater. Date of arrival, relative to length of day on arrival. still may be environmentalclues or time-giversin the annualcycle, but the)' are not the factorsleading to a largerclutch-size. This is supportedby Baker's 11938) evidencethat there is no generaltendency for birds to breed everywhereat the same daylength,nor for birds to breed when the daysare lengtheningparticularly quickly. If we speakof telescopedbroods or clutchesbeing selected in higher latitudes,it may help answersome of the questionsraised by the theory of daylengthcause: such as a largermean clutch-size in Connecticutthan on CapeCod at the samelatitude for Tree Swallows(Paynter, 1954): or a larger clutch-sizein Switzerlandthan in Englandfor Swifts fApus apus), (Lack and Lack, 1951). Vol. XXXII 1061 DaullY,Bylot Island Breeding Biology [ 35

The contradictoryclutches in Lack'sstudy of the EuropeanRobin are smallerwhere the populationtends to be resident. In approachingthis and otherproblems of variationwithin a species,we shou,ldstart at the centerof the rangeand ask not only why it is larger to the north, but whyit is smallerand subdividedto the south. A bundanceo] Foodand thePeriod o/Maximum Dernan&To examine Lack'shypothesis that the successfulfeeding of the youngin the nestis a selectionforce on clutch-size of passerines,we need to followthe weights of individualnestlings to seehow they compareas they leave the nest, and then follow color-banded birds that have left the nest to see whether and howlong the "runt" survives.T•his can be appliedto certaingroups only, becausein some--herons,hawks, and owls--the late or weakling youngdie and are eatenor trampledinto the nest floor. Snow (1958) has donethis for EuropeanBlackbirds and has shown that it was really impossibleto correlatethe breedingof this species with any particularly abundantprey species.The stimulusof rising temperatureled to the maximumnumber of eggclutches being actually presentwhen the chief food---earthworms(Lumbricus)--was abundant. Youngin the nestcame later, at a time whenworms often were harder to obtainand parentshad to seekother food. The artificial situationof the Botanical Garden, where Snow's studieswere made, may have a different annual supply of food than the more natural situation of woodlandBlackbirds which, as Snowpointed out, werenot so decimated by May droughts. The May and mid-summerdroughts hardened the soilsurface and largelyremoved the sourceof worms. The critical periodof the populationrelative to its food supplyshould be consideredto extend from hatchingof young until the young are independentand the parentshave completelymolted. No one period in this time is the critical one. The wholepopulation makes the maximum demandson the food supplywhen all younghave been produced and the parentsare molting(Pitelka, 1958). Furthermore,in the life of the individualfledgling, a criticaltime is whenthe parent has stopped feeding. In early broods,the female.may stop feeding fledglings to lay her next clutch(Tinbergen, 1939). If the youngshe has beenfeeding are not self-sufficientthey die. During thisperiod after leavingthe nestand while the parentsfeed, the youngwho haveinherited behavioral mechanisms for finding,killing, and eatingfood haveto learn (and very rapidly) what food is and where to find it. Ruiter's (1952) studieswith Jays (Garrulusglandarius) show that the bird blundersonto the caterpillar first, and whenit has a "picturein mind" of what to look for, it will seekout the food. But for youngto learn their food, the prey hasto be "superabundant"for the trials and errorsof the youngto happenon the prey oftenenough for themto learn,by "reinforcement."Carrick's (personalcommunication) work on food of birds in Australiashows extremeindividual differenceswhich must depend upon this sort of conditioning. The pointis that:larger broods in higherlatitudes are correlated with daylengthand dependupon ability to feedthe youngin the nestmust be extendedto feedingthe youngout of the nest--indeed,must include theperiod when the youngare independentand the parentsare molting beforemigrating. Pitelka (1958) madethis point in discussingthe molt of LaplandLongspurs. 36] D...¾,Bylot Island Breeding Biology Bird-BandingJanuary

PART III: DISCUSSION OF BEHAVIOR A. COMPARISON OF BEHAVIOR OF LAPLAND LONGSPURS WITH OTHER BUNTING AND FINCHES. Do our observations on the behaviorof the two buntingshelp to clarify their taxonomousrela- tionship? Completedescriptions of the ,behaviorof Emberizinefinches are few but they suggestseveral comparisons. The displaypostures of Lapland Longspursseem closer to the Yellowhammer(Emberiza citrinella)than to the otherEtnberizines, including Snow Bunting, but are more similar to thoseof Snow Buntingand other EuropeanBuntings than to the AmericanEmberizines (Song Sparrow,Melospiza meIodia, andTree Sparrow,Spizella arborea). The)' lack the wing-wavingdescribed in McCown'sand Chestnut- collared Longspurs,but they include wing-vibrating (Andrew, 1957). They differ from the SnowBunting in the natureof the songflight, and in that the longspur"emphasizes" his breastwhile the Snow Bunting "emphasizes"his back (Tinbergen,1939) in displayingto the female. Thereare alsodifferences in songperiod. In SnowBunting, copulation stopsafter the first egg is laid, and the maleno longeraccompanies the female. The female Snow Bunting starts incubation one to three days a/ter the completionof the clutch,instead of one to three days be/ore completionof the clutch as in the Lapland Longspur. Food foraging in LaplandLongspurs was restrictedto the territory, and muchless so in SnowBunting. The aggressiveand courtshipactions agree with generalpatterns for Europeanbuntings described by Hinde (1955), and with specificpatterns describedby Andrew (1957) for Lapland Longspur."Supplanting attack" and "sleeked head-forward threats" were shown in the border disputes,and in the first encounterswith the female 0nale fluffed--of Andrew). Later, and with no transition that we observed, the male assumedan upright posturewith droopingwings and under-feathers fluffedout whenparading before the female (bill-raisedrun, courtship displays,of Andrewl. In our observationsthe wingswere net vibrated in this posture,but we must have missedthis detail. In contrastto Andrew we found this as much associatedwith the song period as the fluffed run. Accordingto Hinde (1953), droopingwings and parading showed sexualdrive, while the uprightposture and fluffingof feathersshowed drive to flee. More recent work has led Hinde and his students to believethe upright postureis associatedwith sexualdrive (Andrew), sincethis postureis assumedduring copulation.The two were combined in the pre-copulatorybehavior of longspurmales. The combinationof uprightposture and bill-up paradingshowed off the male'sblack throat and white belly (the posture must have precededthe plumage, as Lorenz (1941) has suggested). Exaggeratedupright posture,drooping and vibrating wings, and handlingof nestingmaterial by the male precededcopulation (wing- quiveringand nest-sitedisplay; bill-raised run, Andrew). My observa- tionssuggested that this wing-vibratingwas an attemptto fly up rather than a ritualizedpart of the display,and I saw it only immediately beforecopulation. The malecrouch, with headlowered and rump fluffed --just before, during, and after copulation--isdescribed by Andrew •ro•.1961XXXII Dt•t;t•Y,Bylot Island Breeding Biology [37 only under bill-raisedrun, and not in this context. Hinde (1952) stronglysupports the idea that the rise of sexualdrive in bothmale and femalesuppresses the driveto attack,and that sincethe male'ssexual tendencygrows faster than that of the femaleduring most of the court- shipand nesting period. he is subordinateto her. Hence,following the uprightposture by the fluffedand horizontalposture at the peakof "conflict"may showthe strengthof the male'sthwarting because he is subordinate. The differencesfound whennay notes are comparedwith Andrew's suggest(1) a varietyof specificpostures of particular"meaning" which can,be combined in variousways or (2) variationsin posturesbetween populationsor (3) greatersexual drive in the malesI studiedbecause of the latenessof the season,and the advanceof their conditionbefore the fernaleswere receptive.I prefer to avoid assigningmotivations until moreis knownof specificsituations in whichthe displaysappear. Pointsof interestin placingLapland Longspurs' displays properly in comparisonwith the displays of otherfinches are: (1) songflight (which is to be expectedin a tundraregion); (2) horizontalhead-forward postureof aggression;(3) lackof bill-snapping,wing-raising, pivoting or twisting,or courtshipfeeding; {4) symbolichandling of nesting material;and (5) unusualposture of the malejust beforecopulation. Theseare differences,while the rest of the posturesfit into the general pattern,sleeked head-forward posture, supplanting attacks, head-up postureand fluffed posture--the main elements found by Hinde(1953, 1954, and 1955). B. CHANGES IN M,4LE BEHAVIOR ,4ND TERRITORY ESTAB- LISHMENT OF LdTE dRRIFdLS. What explanationcan we offer for the two separateperiods of territoryestablishment in our areas?The maleLapland Longspur's hostility changes (1) whenhe has been joined by a female,(2) whenthe femalehas started building the nestand layingeggs, and (3t whenthe femalehas startedto incubate.The aggressivetendency of thenaale sinks as his sexualdrive rises; at the sametime his singingand responseto his neighbors'activities change. Eventuallyhe seemsto lose interestin territory borders,just as the pressure]s highest.if the"reason" for territoryis food. We recorded changesin territoriesof LaplandLongspurs indicating that this change allowsthe establishmentof territory by maleswhich arrived later or those which are unable to establishterritories early in the season i Figure7). Later arrivalstake up territoriesin undefendedareas as the aggressivenessof the territoryholders fades or is concentratedonto a smallerarea. Meverriecks(1959) has documentedthis changein territorial defense and its effect on territories in colonial Green Herons ( Butorides virescens). Of the sixteennests of LaplandLongspurs in half of our studyarea, sevenclutches (Nests Nos. 1, 2, 3, 4, 7, 8, and 1 unnumbered)were completedearly (22-25June)•(Figure 7a), andall of thesenests were about100 yardsapart. Nine clutcheswere completed late (1-4 July)-- i Figure 7b). Thesebirds were not banded,and we do not have detailedobserva- tions on the changein behaviorof individuals;but the evidenceis clearthat therewere two setsof territoryestablishment nine daysapart. 3• ] DRURY.Bylot Island Brecdb•g Biology •E•rd-•Eand•ng$anuary

The later nestswere placedinside the area includedin the five earlier territories. Smith (1950) describedthe late arrival of Yellow Wagtails (Mota- cilla tiara) in Englandat an area where three territorieshad already been established. The newcomer established himself in five acres t half of the total area) where there were two territorial males whose females had nestsand were incubating.At the sametime a singlemale, whose nestingactivities had not proceededas rapidly, held five acres and excluded the newcomer. The functionsof territory are complex,and authorshave differed on whetherthe mechanismsare simple or various (Symposium,Ibis, 1956). One feature often neglectedis that selectionacts on maximum successfulbreeding of the whole populationrather than on especially successfulbreeding of a few individuals.The late breedingof yearling birdsis souniversal that it suggestsselective advantage for the population as a whole. This advantagemay comefrom the increasedopportunities for the inexperiencedbirds to establishterritories when the experienced adultshave passed their peakof hostility. Young birds whichhave not establisheda territory in a previousyear are at a disadvantageunless theycan choosea time whenthe hardenedcampaigners will, for various reasons,be "willing" to let them insinuatetheir territoriesamong those of the maturesegment of the population.Usually it has been asstuned that oncea territory is established,it remainsconstant for that breed- ing season,and late arrivalscan only carve out a territory by dint of specialaggressiveness. Most theorieson the function of territory relative to food for nestlings(Lack, 1948b) dependon this premise. But what doesit meanto the biologicalfunction of territoryif the "any defended area" (Noble, 1939) is a great deal smallerwhen the young hatch than whenthe singlemale first establishedhis territorial boundaries?There is dangerof confusionof proximatevs. ultimatecauses, and dispersion mustbe carefullyseparated from territorialityif we considerthe main selectionto be concernedwith the food supply. Furthermore,there are adequate"needs" to lead to the phenomenonof territoryexpressed in courtshipbehavior. Tinbergen, Lorenz, Lack, and their studentshave shownthe "need" for a male to act aggressivelyin many casesin order .to effectsex recognition. This, combinedwith site tenacity (Ortstreu) leads to a courtshipexplanation of territory as Tinbergensuggested (1957). But I do not suggestthat food is not an evolutionaryadvantageous effect also. Territorycan be expectedto have differentexpressions and differentselective advantages in every group in which it •isexpressed. I presenteda report on LaplandLongspurs. including the idea just discussedat the meetingsof the American Ornithologists'Union at CapeMay in September1957, and in responseto questionsI saidthat I thoughtthe idea had alreadybeen published; but sincethen Meverriecks and I havenot beenable to find it in print.

C. SELECTIVE ADVANTAGE OF MIGRATION IN TERRITORY ESTABLISHMENT. Althoughthere are many selectiveadvantages in migration,our studiesof longspurssuggest ariother, the apportionment of availablebreeding territory. On the average,mortalit;' comes in large Vol. •9alXXXII DauaY,Bylot Island Breeding Biology [ 39 juvenilefailure and a steadydeath rate of adults[Nice, 1937: Kluijver, 1951; and summaryin Lack, 1954). Recentpopulation studies have shownthat the greatpeak of juvenilemortality in residentspecies comes betweenJuly and October,even though migratory dangers do not exist. As Kluijver (1951) has suggested,mortal dangersfor residentspecies result from the emigration of juveniles into strangehabitats, seeking to establishtheir ownterritories. Such young often occupy less desirable habitats,such as the pine woods,in Kluijver'sstudies. Snow (1958) has datawhich show that anotherimportant peak in mortalityoccurs during territory establishmentand the early part of the breedingcycle in the EuropeanBlackbird. In a wholly migratoryspecies, when the returningpopulation takes up territoryin spring,newly territorial birds do not face an entrenched winteringpopulation with inexperiencedyoung at a maximal disad- vantagerelative to the experiencedadults. This mustbe an advantageto the popula.tionas a whole,since it allowsmore adjustmentamong the arrivalsand lessabsolute exclusion. It may be that for the year-crop, migrationis an advantagein spreadingout the population--balancingthe migration hazardswhich becomethen no worsethan thosehazards met by juvenilesleaving their parents'territories when neighboring birds are in a phaseof temporaryascendancy of territorial behavior(Lack 1953, Nice 1937). D. LONGSPUR NEST SITE SELECTION. Longspursat the Aktineq built their nestsconsistently under the decumbentbranches of clumpsof Bell Heather (Figure 6). How can this observationbe rationalizedwith the nearly universaldescription of the nestingof this speciesin low, rolling, wet, hummockytundra, under a tuft, or in the sideof a mossor sedgehummock (Blair, 1936: Dalgety, 1936; Grinnell, 1944; Nicholson,1930; Soper, 1928 and 1946; Sutton, 1932; Suttonand Parmelee,1955b; and Wynne-Edwards,1952)? Havi- land (1916) found nestsfrom the wettestto the driest habitats on the YeneseiRiver, and A. Murie (1946) found them on gentleslopes cov- ered with Avens (Dryas integri[oliaM. Vahl). Grote (1943) reports longspursnesting on all sorts of tundra in Siberia--whereverthere is plant cover--but agreeswith Birula (1907) that the speciesis charac- teristicallya hummock-tundrabird. Only one nest at the Aktineq (No. 12) couldbe said to be placedin a low, wet place,and No. 11 was in a frost-crackon the edge of a wet area. Bell Heathergrew in shallowdepressions on slopeswhich (after the barren and exposedridges) were the first free of snow,and longspurs chosethe edgesof the Bell Heatherpatches first free of snowon slopes facing southeast,south, or southwest,from which the snow melted very early. Among26 nestsfound, there was only one exceptionconcerning exposure. Birula i1907), Michejev t19391, and Tolmatschev11934) report nestsfacing south, east. or southeastand shelteredfrom the wind by hummocks,grass or low heaths.In the Siberianareas studied a south- east wind is almost unknown in summer. Frazer Rowell { 19571 reviews the literature o.nnest sitesof this species.DuBois' (1935) descriptions indicate that the nests of Chestnut-collaredLongspurs are similarly sheltered.in contrastto the exposedsituations of McCown'sLongspurs' nests. The breedingsites we found are similar to thosefound bv Havi. 40] DRURY,By/ot Is/and Breeding Biology Bird-BandingJanuary land and Murie, presumablybecause the wet meadowsites were snow- coveredtoo late in the species'breeding cycle and •becausethe species, displacedfrom ideal nest sites,selected a form resemblingthe overhang of a tussock. At the Aktineq, 14 of 19 nestswere directly associatedwith Bell Heather,'but at Ooyarashukjooeet(Miller, 1955) only 2 of 6 nestswere so associated.Nests at Ooyarashukjooeetwere overhungby Avens, ArcticWillow or a clumpof GrassRush and placed in frost-cracks,under solifluctionlobes (Washburn, 1956), or on the sidesof a raised beach ridgewhere no Bell Heathergrew. In theAktineq region, steeper slopes, lessmarked with deep frost-cracksand mounds,provided fewer hum- mocks,and the best availableform was provided by the decumbent growthof BellHeather. The regularfrost patterns with steep-sidedcracks at Ooyarashukjooeetsupplied the proper forms, as did tussocksnear lemmingmounds and plant cushionson the sandbars. This seemsto indicatethat longspursnest in a shelteringoverhang facingtoward the southand that actuallyBell Heatheras suchdid not matter. It was a form requirementrather than a speciesrelation. Ob- servationsof other authorsfarther southand in Greenland(Salmnonsen, (1950-51) refers the speciesto the verge of high arctic areaswhere Crowberry(Empetrum nigrum L.) is replacedby Bell Hdather),indi- catethat the speciesdoes not belongto a particularvegetation associa- tion. Rather, it takes what is available and best fits its needs in those areasthat satisfythe physiologicalrequirements, and from which it is not driven by competingspecies. SUMMARY Horned Lark 1. This specieswas the mostwidespread on Bylot Island, and had the largestterritories among the passerines. 2. Nearlyall singingwas in the air and the flightsong was identical in form, althoughdifferent in quality,from that of praticola. Song flightstook place over the individual'sterritory. 3. We found a clutch-sizeof 4-5 eggsand recordeda specificalarm notegiven by parentswith young. 4. ta) "Casualabandonment," lb) very rapid feeding at the nest, and (c) lack of broodingof the youngduring the day, contributed to effectiveconceahnent of nestlings. FFater Pipit 1. Flight songwas regularly two-parted. 2. The trilling songcorrelated with immediateattack on an intruder. 3. Territoriesaveraged less than 100 yards long and about 200 feet high in the ravinesin south-facingbluffs. 4. The single nest found containedfive eggs and was excavatedinto a slightoverhang. It wasbuilt entirelyof grasses. 5. This speciesis the only clearlyLow Arctic representativeon Bylot Island. The nest is. we believe,the most northern recorded. LaplandLongspur 1. This wasthe mostabundant nesting species: 27 territoriesin the studyarea, and at least60 in the sevensquare miles betweenthe Vol. •9•!XXXII DRURY,Bylot Island Bree•ting Biology [41

Soundand the snoutof the AktineqGlacier on the westbank of the river. 2'. Whenwe arrivedwe foundmixed flocks containing .many male and a few femalelongspurs, as well as SnowBuntings and Horned Larks. Males took up territoriesbetween 15 June and 25 June. 3. Mostsongs were given in flightat a heightof about20 feet. Each malehad severalsong perches, and sangrepeatedly from the ground. The maleshad a whispered'songgiven in closecompany with the femaleafter pairing. 4. Territorial disputesinvolved flying chases,song duels, and, on the ground,forward postureswith a scoldnote and gapingbill. Rushes were directed at the side of an intruder. 5. Trespassingwas tolerated and territorialdefense seemed suppressed by the winter flock calls. Song stimulatedimmediate response in theform of singingand dereaseof territory. 6. Territories occupieda half acre or lessfor the twelvemost carefully studiedpairs. Once a maletook up territory,he did notleave it until the end of the breedingcycle except for very brief periods. 7. The first displaysto a femaleresembled hostile displays. In sexual pursuits,one female was pursuedby two or three malesin succes- sion. When she settledin one territory, she was still pursued'by her male, and the pursuit flights still sometimestrespassed. The femaleincited pursuit in flight and on the ground. At this time the males'displays assumed an upright posture. Song remained vigorousbut less frequentthan during the pre-pairingperiod. It occurredon appearanceof a neighborin songor whenthe female disappeared. 8. Males' pre-copulatorydisplay includedcarrying dark material. uprightposture, and song. This wasfollowed by a crouched,fluffed posture before copulation. The female crept and hopped in a crouchedposition. After copulationthe malereturned to the crouch- ing position. The female stood erect, raised her head high and cocked her tail. 9. Nestingsites included a numberin which Bell Heather overhung the nest. 10. The nestswere constructed entirely by the female,and were made of dead and dry grass,tightly knit and lined with white feathers and often willow cotton. The ,maleconstantly accompanied the female while she built, occasionallypicking up brown nesting material similar to that she used, only to drop it. The male ac- companiedher closelyuntil she completedher clutch. 11. Clutcheswere completed between 22 Juneand 4 July. Thosehatch- ing 3-9 July contained:4 (6), 1 (5), 1 (4) eggs;and thosehatch- ing 10-15 July contained:2 (5), 5 (4), 2 (3) eggs. 12. The femalestarted to "incubate"sporadically on the secondday be. fore the lastegg was laid. and incubatedmost of the day beforethe lastegg was laid in nestNo. 6. The day afterthe lastegg was laid, the male stoppedhis attendanceon her and sangactively for three days after that. Then his songdiminished until it had effectively 42 ] D.u.Y.Bylot Island Breeding Biology Bird-BandingJanuary

stoppedwhen the younghatched. The malestook no part in incu- bation, but were constantlyalert to trespassingon the territory. 13. During 80 visitsto 14 nests,the femalewas found on the neston all but 18 visits; two absencesfor 7-9 visits.The femalekept the nest clean of any dirt that fell in amongthe eggs,and removed brokeneggs to sucha distancethat we did not find any of four we damaged.Removal of eggsduring the layingof the clutchand after the clutchwas completed had no effecton the numberof eggslaid. 14. Hatchingwas delayedin nestNo. 1 whenthe eggswere soakedin ice melt-waterfrom the fourthto the sixthday after the clutchwas completed,but the embryoswere not killed. Onemalformed nestling (perhapsthe resultof the chilling) waslater ejectedby the parents. 15. Eggswere in thenest from 16 daysfor thefirst egglaid in nestNo. 6, to a minimumof probably101/.2 days for the fifth egg in the same nest. The usualperiod for eggsconstantly incubated was 11 to 12 days. One eggin eachof nestsNos. 2, 7, and 13, and threeeggs in nest No. 10 were infertile. Theseeggs were left in the nest by the parentsuntil the youngleft. 16. In some nests all eggs hatched during the same 24-hour period (Nos.3, 6, 7, and 13), andin othershatching extended over as much as 48 hours(Nos. 1, 11, and 14), or possibly64 hours (No. 2). 17. The male took at leastas large a part in the feeding of the young as did the female; and in nestsNos. 3, 6, 8, and 12 the male was more active than the femalein feedingduring the early morning and all evening,when the youngwere 3-5 and 6-8 days old. 18. Twenty-twonestlings left the nestafter 9 days,12 left after 10 days, and 3 left after 8 days in the nest. All left 2-3 days before they could fly. Nestlingswere essentiallysilent until the last day in the nest. They had dorsaldown on hatching,feathers on their backs at four days (whentheir eyesopened) and were fully leatheredby the seventhto eighthday. Youngleft the nestfrom 12'-24July. Our recordsindicate that the young left within 24 hours of each other in four nests, and within 48 hours of each other in five nests. 19. Sixty-nineeggs •¾ere found, of which 10 were destroyedby us or by a dog with us. Of the remaining59 eggs,6 were infertile and 5 abandoned;one young was deformedand pushedout, and three young disappeared.Forty-two young left the nest (we took two specimens).Eight l perhaps9) of the 15 nestssuccessfully fledged all of the young from the full clutch. Thirteen nests,or 86 percent of the total,hatched one or moreeggs. The averagenumber of eggs that hatchedin successfulnests was four; 75 percentof the eggs producedfledged young.

SJ,ou' Bunting 1. We arrived whenbirds werepaired, but nest-buildinghad not started. 2. Four nestswere in holesexcavated into loosesand on exposedridges or vertically-cut-banksof the river. One nestwas under a boulder. 3. We found 2 (6) and 1 (5) clutchescompleted 23 June to 3 July. 4. The female at nest No. 3 repeatedlyleft the nest to feed, but also Vol.1961xxx• D[tt'•¾.Bvlot Island Breeding Biology

beggedfor foodfrom themale. This samemale performed a pre- pairingdisplay to herwhen there were four eggs in thenest. 5. The youngin thenest were as silent as nestling larks and longspurs, butwere noisy after they had left the nest.Both parents fed the young. 6. Of five nestsfound, only two weresuccessful. We foundone group of eightyoung with two setsof parents. Ecology 1. Exceptfor plovers,only single representatives of a genuswere found in our district. 2. We sawno ecologicalcronpetition among the four passerinespecies. Territorywas largest in HornedLarks (1,,• sq. mile) andsmallest in LaplandLongspurs • 1/.2 acre). 3. We foundno specialadaptation in LaplandLongspurs (i.e., different fromclosely related longspurs) which would prepare them to occupy their northernrange. 4. All four specieswalk and havea long hind claw whichmust be regardedas functionaland thereforeof no systematicimportance. 5. Clutch-sizesshow no greaterannual productivity in the norththan do multiple-brooded,smaller-clutched populations in the south. Adaptationhas concentrated the breeding season. 6. No particulartime in the breedingcycle is criticalwith regardto the food supplythereby controlling clutch-size. Food is a .critical factorwhen the speciesarrives, when there are youngin the nest, andespecially when the youngfledge and the parentslnolt. 7. An importantforce selecting annual productivity in arcticbreeders is the dangerof climatic--non-densitydependent--disasters. 8. The advantageof single-broodednessis that recently-fledgedbut incompetentyoung are lesssubject to abandonmentthan when the female starts a second brood.

Behavior 1. LaplandLongspur behavior is closeto publ:sheddescriptiong of Yellowhammerin most of its features,and differs in severaldetails from otherlongspurs. 2. We foundbill-raised run (Andrew) associatedwith songperiod, and wing-vibrationassociated with precopulatoryflight. Bothdrooping wingsand uprightposture were associatedwith sexualsituations. The precopulatoryposture we saw is describedunder anothercon- text by Andrew. Behaviordoes not vet clarify the taxonomicrela- tionshipsof LaplandLongspurs within the buntings. 3. We founddecrease in aggressivenessof territorial lnales after pair formationto coincideMth intrusionon occupiedterritories by later males. This allows concentrationof the populationin the few suitableareas available,and may be an evolutionarymechanism permittingyearlings to establishterritories later than adults. 4. In migratorybirds, less well-motivated or inexperiencedyearlings settingup territoriesare lesssubject to discriminationthan are yearlingsof residentspecies. 44] •)RuRY,Bylot Island Breeding Biology Bird-BandingJanuary

5. Nest site selectionby longspursgave insight into the form of the environmentsituation which they chose;i.e., shelteringoverhang-- whether moss hummock,grass tussock,frost-crack, or decumbent growthof Bell Heather.

BIBLIOGRAPHY A.O.U. Check-list of North American Birds--Fifth Edition. 1957. 691 pp. Balti,more. ANDRew,R. J. 1957. T'he aggressiveand courtship behaviour of certain Em- berizines. Behaviour 10(3-4): 255-309. Ba•Lz¾, A.M., and R. J. NmDRacm 1938. The Chestnut-collared Longspur in Colorado. Wilson Bull. 50: 243-246. Ba•zR, J. R. 1938. The relation between latitude and ,breeding season in birds. Proc. Ecol. Soc. London :4108: 557-582. BENT, A. C. 1942. Life histories of North American flycatchers, larks, and swallows. U.S. Nat. Mus. Bull. No. 179. ß 1950. Life Histories of Nort,h American wagtails, shrikes, vireos, and their allies. U.S. Nat. ,Mus. Bull. 197: 1-411. BmuLa, A. 1907. Sketches of the Bird Life of the Arctic Coast of Siberia. Acad. Sci. St. Petersburg 1907. BLare, H. M.S. 1936. On the birds of East Finmark. Ibis 78: 280-308, 429-459, 651-674. DALCETY,C. T. 1936. Notes on birds observed in Greenland and Ballin Land, June-September 1934. Ibis 78: 580-591, pls. 9-11. DIXON, JosEPH S. 1938. Birds and mammals of Mount McKinley National Park, Alaska. Fauna Nat'l. Parks U.S., Fauna Series No. 3. 236 pp., 1 pl., 85 figs. DRuRY,W. H., JR., and M. C. DRuRY. 1955. The Bylot Island Expedition. Bull. Mass. :4ud. Soc. 39(6): 259-265. DRuRY, W. H. JR., 1960. Observations on the breeding cycle of Long-tailed Jaeger, Herring Gull, and Arctic Tern on Bylot Island, Northwest Territories, Canada. Bird-banding 31: 63-79. DuBo•s, A.D. 1935. Nests of Horned Larks and Longspurs on a M'ontana Prairie. Condor 37: 56-72. 1937a. Notes on Colorationand Habits of the Chestnut-collaredLong- spur. Condor 39: 104-107. 233.•38•937b.TheM'cCown's Longspurs ofa Montana Prairie. Condor 39: FRANZ,J. 1949. Jahres- und Tagesrhythmuseiniger VSgel in Nordfinnland. Z. Tierpsychol. 6: 309-329. FRaZ•.RROWELL, C. H. 1957. The breeding of the Lapland Bunting in Swedish Lapland. Bird Study 4(1): 33-49. GRINNELL,L. I. 1944. Notes on breeding Lapland Longspurs at Churchill, M'anitoba. Auk 61(4): 554-560. GROTE,H. 1943. Ueber das Leben der Spornammer (Calcarius lapponicus) in der Tundra. Beitrage zur Fort. de Viigel 19: 98-104. HaVlLAND,M.D. 1916. Notes on the Lapland Bunting on the Yenesei River. British Birds 9: 230-238. H•Nm;, R. A. 1952. The behaviour of the Great Tit (Parus ms/or) and some other related species. Behaviour, Supplement 2. ß 1953. The conflictbetween drives in the courtshipand copulationof the chaffinch (Fringilla coelebs). Behaviour 5: 1-31.

Beh•violu9r54.7: 207-232.Thecourtship andcopulation ofthe Greenfinch (Chlorischloris). . 1955. A comparativestudy of the courtship of certain finches (Frin- gillidac). Ibis 97(•1): 706-745. HOFF•aaNN,K. 1959. i)ber den Tagesrhythmusde Singv/igel in arktischen Sommer. Journ. /iir Orn. 100: 84-89. HOWARD,H. E. 1929. Introduction to the study of bird behaviour. Cambridge Univ. Press. KaRPLUS, M. 1952. Bird activity in the continuous daylight of the arctic summer. Ecology 33: 129-134. Vol. XXXII 1961 lh•vn¾.Br[ot Island Breeding Biology [45

KLUIJVER,H. N. 1951. The Population Ecology of the Great Tit (Parus m. ma/or L.). Arden 39: 1-135. L.ac<, D. 1933. Nesting conditions as a factor controlling breeding time in birds. Proc. Zool. Soc. London: 231-237. 1947. The Significance of Clutch-Size--Part I. Ibis 89: 302-352. 1948a. The Significance of Clutch-Size--Part II. Ibis 90: 25-45. 2:9•-1109. 48b'Natural seS-ction andfamily size in the Starling. Evolution 1953. The Life of the Robin. Pelican Books, London. 1954. The Natural Regulation of Animal Numbers. Oxford Univ. Pres•,London. ---, and E. Lack. 1951. The Breeding Biolo•-q;of the Swift, .4pus ap.,•s. Ibis 93: 501-546. LoaœNz, K. 1941. Vergleichende ,Bewegungsstudienan Anatinen. Jour. /iir Ornithologie, Festschrift Oskar Heinroth: 194-293. LOVr•NSK•OL•),H.L. 1947. Handbok over NorgesFugler. Os]o. Ma.•N•Cm;. A.L.¾. 1910. The terrestrial mammals and birds of north-east Greenland. .lieddel. om GrOnland 45: 1-200. M^asn.a•.L,A. J. 1952. Non-breedingamong Arctic Birds. Ibis 94: 310-333. Mz•qmamc

humans; with special reference to the role of the aggressive-,escape-, and brooding-drives.Behaviour 8: 130-173, pls. 1-10. S,'aXT•, S. 1950. The yellow wagtail. Collins, London. SNOW,D.N. 1958. A Study of Blackbirds. George Allen and Unwin, London. Sovza, J. 1928. A faunal investigation of southern Baitin Island. Nat. Mus. Canada Bull. No. 53 (Biol. Ser., No. 15). 143 pp. (incl. 7 pls.), ,map. ß 194.6. Ornithological results of the Baffin Island expeditions of 1928- 1929 and 1930-1931,together with more recent records. •4uk 63fl): 1-2k pls. 1-2, 2 maps., pp. 420-421. SUTTON,g. M'. 1932. The birds of Southampton Island. Mere. Carnegie Mus. 12 (Part II, sec. 2). , and D. F. Parmelee. 1954a. Survival problems of the water-pipit in Baitin Island. Arctic 7(2): 81-92, 3 figs. 1954b. Nesting of snow bunting on Baitin Island. g/ils. Bull. 66 (3): 159-•79. 1955a. Nesting of the horned lark on Baffin Island. Bird-Banding 26(i): 1-18. 1955b. Summer activities of the Lapland Longspur on Batfin Island. g/ils]Bull. 67: 110-127. Svaal)SON,G. 1949. Competition and habitat selection in birds. Oikos 1(2): 157-174. TINBEat;EN,N. 1939. The behavior of the snow bunting in spring. TransßLinn. Soc. New York, 5: 1-94. 1957. The Functions of Territory. Bird Study 4(1): 14-27. TOWNSEt'D,C. W., and'G. M. AzzEN.1907. Birds of Labrador.Proc. Boston Soc. Nat. Hist., 23 (7): 277-428, pl. 29. TuGaaXNOV,A, and A. Toz•aTSCl•Zv. 1934. Information on the Avifauna of Eastern Taimyr. Leningrad Acad. Sci. VAN TYNE, J., and W. H. DauaY, Ja. 1959. Faunal studies of the 1954 Bylot Island ExpeditionßOcc. Papers Univ. Mich. Mus. Zool. No. 615: 1-37. Was•auaN, A. L. 1956. Classificati.on of patterned ground and a review of suggestedorigins. Bull. Geol. Soc. ,4ruer. 67: 823-866. WyNN•;-EI)waal)S,V. C. 1952. Zoology of the Baird expedition (1950). 1. The birds observed in central and southeast Baitin Island. •4uk 69(4•: 353-392. Received January, 1959

MIST-NETTING BIRDS ON ANDROS ISLAND, BAHAMAS

By LAWRENCEH. and CLARAM. WALKINSHAW

From the 16th to the 26th of March, 1960, we were on Andros Island, Bahamas,making a studyof the birdsin the areawhere we stayed.We hopedto capturethe long elusiveKirtland's Warbler (Dendroicakirt- landii) by the useof mist-netsbut failed to evensee nor hear the species. However,we did have someluck in capturingother species. The weatherwas very goodbut at timeswind bothered.We operated from one to five nets for a total of 305 net-hours. Since the soil is chieflycoral and almostimpossible to dig into and the fact that we wereunable to getmetal poles, we wereforced to uselong wooden poles improvisedin positionby ropes,stones, etc. We had much trouble with localwild dogsand at timescats (someof which were alsowild). We capturedonly 105 birds.