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 adaptationsallow 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