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Ecology of a Breeding Population of Arctic Terns

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Ecology of a Breeding Population of Arctic Terns BIRD-BANDING A JOURNAL OF ORNITHOLOGICAL INVESTIGATION Vol. XXVIII April, 1957 No. 2 ECOLOGY OF A BREEDING POPULATION OF ARCTIC TERNS By OSCARHAWKSLEY INTRODUCTION Biologiststoday are •nuch concernedwith the ecologicalfactors which controlnatural populations. Avian populations,particularly of colonialspecies, lend the•nselveswell to investigationsof the survival and mortalityrates of offspringup to the ageof attainingflight. Factors whichcontrol the successof the socialunit or colonyas a wholeduring the reproductiveperiod may alsobe readilystudied. The presentstudy wasmade during the breedingseasons of 1947 and 1948 as a part of a stud)- of the behavior and ecology of the Arctic Terns •Steraa paradisaeayof MachiasSeal Island, Bay of Fundy, New Brunswick. Canada. Although a greater portion of time had to be allotted to behaviorstudies, sufficient data have been acquired to allowan analysis of eggand chickmortality of the ArcticTern and to providematerial for discussionsof the factorsaffecting group success in thisbreeding colony. Thisis fortunatesince Dr. RaymondA. Payntermade detailed studiesof the eggand chickmortality of HerringGu,11s (La. rus argea- tatussmithsonianus) and or.herspecies on Kent Island,only fifteen milesaway, during the summersof 1947 and 1948 t Paynter1949). Thus,by coincidence,so•ne comparison of data obtainedin the same locality,during the sameyears and for a relatedspecies, is possible. Due to the exceptionallywide distributionof the Arctic Tern, in- formationon the speciesis scatteredthroughout the ornithological ,literatureof manycountries and of manyyears. In this paper,an attempthas been made to bringtogether much of the moreimportant informationdealing with the ecologyof the species. I wishto expressmy appreciat.ionto Dr. RalphS. Palmerfor many helpfulsuggestions based on his experiencewith terns. Drs. RobertT. Clausen,Richard H. Backusand CharlesL. Remingtonhave assisted withidentification of plant, fish and invertebrate specimens. My wife assistedconstantly with field work which could hardly have been completedwithout her. SPRING ARRIVAL For manyyears, the lightkeeperand his wife, Mr. and Mrs. Ottawa Benson,kept recordsof the arrival datesof the Arctic Terns at Machia• SealIsland. Arrivaldates and comments from their log for fivecon- secutiveseasons are presentedbelow. 57 Bird-Bandi•ng 58] H•WKSLEY,Breeding Population o.fArctic Terns April In 1945,the terns arrived on May 17 and startedlaying on May 29. In 1946,they also arrived on May 17. In 1947,the terns arrived on May12 but did not alight on the island until May 20. Ternswere first seenon May 12 againin 1948but did not alight until May 17. In 1949, thebirds were heard on May 11 andfirst seen on May 12. Theylanded brieflyon May 14, but the presence of a DuckHawk (Falco peregrinus) anatum)kept them off for a coupleof days.After the hawk was killed by the keeper,the ternsreturned and eggs were found by May 29. Althoughthe terns arrived on May 12, 1948,there were only a few eggslaid whenI arrivedon June 3. We receivedthe impression that the numberof birds on t,he islandincreased for severaldays after that, but sawno arrivingflocks. Accordingto the Bensons,the ternsarrive at MacbiasSeal Island in large,high-flying flocks. The first time they appear,they circle high above the island, then leave. Afterthat they fly overthe island each morning in socialflight but do not alightuntil theyhave done this for severaldays. Pahner11941:401 has described thislast type of behaviorsimilarly for the CommonTern (Ster•mh. hirundo). THE ENVIRONMENT FOOD AND FOOD-GETTING The rich waterssurrounding Macbias Seal. Island provide an abundant foodsupply for thatternery. ArcticTerns, like otherspecies of terns, probablycolonize areas close to goodfood supplies.This doesnot necessarilymean that the ternery must be locatedclose to the sea. Colonies are known to exist even in the interior of the Scandinavian countriest Kullenberg 1946: 12t wherethe birds fish in lakesand rivers. In Bail:inLand, Dalgety (1936• notedthat Arctic Terns "appearedto preferfishing in freshwater." In contrastingterns with gull,s,Tinbergen (1932: 11-12) remarks itrans. 1: In manyways the gullsappear remarkably adaptable, much more so than the terns. The gullslook for food in many ways,changing with the circumstances.The terns, on the other hand, show a strar•gepsychic unadaptability. Their behavior,especially on their searchfor food, is very stiff. We could call them, briefly, over- specializedplungers. Althoughthis conceptis fundamentallycorrect, one mustnot think of the Arctic Tern as a wholly unadaptablespecies. It is principallya deep-waterfeeder, yet it quicklyadapts to the useof shall.owowaterfood organismsor of insectsat timeswhen these are especiallyabundant. In a colonyfive milesfrom the sea,Wright (1909) observedthat the youngwere being fed not with fish,but with largecrane-flies and may- flies. These terns were caught for banding on hook and line baited with crane-fliesand "greendrake." Palmer (1941: 17) did not see pelletsfrom Commonor RoseateTerns (Sterna dougalli), but I fre- quentl.ynoticed pellets, containing insect parts, on the rocksin the roost- ing areas on MachiasSeal Island. The food habits of the Arctic Tern have been little studied. A com- pilationof specificfood organismswhich the speciesis knownto utilize the world over would be of litt.le value in this study and might even Vol.1957XXVIII H•WKSLEY,Breeding Population o[- Arctic Terns [59 give an incorrectimpression. Specific names will be usedonly in referringto food usedin the studyarea. A surveyof the literatureindicates that the three mostimportant food groupsinclude fish, crustaceansand insects. Fish from the families Gasterosteidae,Ammodytidae, Stromateidae, Scorpaenidae, Cyclopteridae,Blenniidae, and Gadidaeare much used. Crustacean groupsmuch used include branchiopods, copepods, mysids, amphipods, euphausids,and decapods.Insect orders represented are Ephemerop- tera. Odonata,Lepidoptera, Coleoptera, Hymenoptera and Diptera. A few pelagicmolluscs such as Clio, Limaci,a and .Loligoare eaten, as well as marine worms. Availabil.ity is obviouslya very importantfactor in whatfood is used. In Labrador, O. L. Austin, Jr. (1932: 128• did not seeArctic Terns eat anythingbut the lanceAmmodytes americana, but thesesha•llow water fish werenot usedat all at MachiasSeal Island. CaptainFielden (1877) madethe interestingobservation in polar regions,where continual ice preventsfish from comingto the surface,that the amphipodAnonyx nugaxworks its way up throughtidal ice cracksand is eatenin numbers by the Arctic Tern. The fishesmost used for food by the terns at Machias Seal Island werehake (Urophysissp., probably U. chuss),the 1.umpfish(Cyclopterus lumpus) and the dollar fish (Poronotustriacanthus). However, the supplyof skeletonshrimps (Meganyctiphanes norvegica) and of lump- fish appearedto be mostconstant. Thesewere used extensively during intervalswhen hake and dollar fish seemedto be lessplentiful and were usedto someextent throughout the entire season. There werenoticeable seasonal "runs" of mostof the food organisms. Notableamong these were the "runs" of cicadas(Oka,agana cana•4en- sisy an annelid (Nereis pelagica), small squids (Loligo pealei) and dollar fish. The cicadaswere used during the secondweek of July in both 1947 and 1948. In 1947, Nereiswas used from July 8 to July 11, squidscame into usefrom July 10 to July 17, and dollar fish were used in large numbersfrom July 24 until we left the islandin early August. Other fish, in the order of the frequencywith which they were brought to the colony, were four-'beardedrockling (Enchelyopus cimbrius),Clupea sp., haddock(Melanogrammus aegehfinus), rock eel (Pholis gunnellus) , three-spinedstickleback (Gasterosteus aculeatus) , rosefish (Sebastosmarinus), and barrelfish (Palinurichthysperei- ]ormis). The nine-spinedstickleback (Pu,gitius pungitius) was com- monly usedat coloniesnear Churchill,Manitoba, during studiesmade there in 1949. Unidentifiablemoths and a dragonfly(Tetragoneuria spinigera) were found in nestswith young. EFFECTS OF WEATHER AND LIGHT The importanceof weatherto ArcticTerns is indicatedin manyparts of this paper. Weatherconditions have a direct effectupon the ability of adults to seek food. Later it will be shown to what extent weather affectsthe survival of the young. Roostingor brooding adults may becomeso wet in rain or densefog that they are unableto fly out of tal,1 60] [la;;•sLEY,Breeding Poptdation o.iArctic Terns Bird-E•andingApril lOO • 70 • 60 • so 40 2 3 4 5 5 20 25 3031 1 July Aug. Fig. 1. Growth of chicks as related to •eather. vegetationand may be easilycaught. Weatherand light influencethe amountof socialflying to so•nedegree. Weather, especiallythat which affects visibility, and the need for brooding young, has a very decided effect upon the ability of Arctic Ternsto providefood for their young. Weightsof chicksfrom July 2 to August1, 1948, showeda definitecorrelation between certain weather conditionsand the averageweight of the group of chicks. Figure 1 is a graph showingthe averagedaily gains and lossesin weight and the visibility factors which seemedto contrt•l them. It is interestingto note that a singleday of fog or unfavorableweather may not resultin weight losses,but that several days of unfavorable weather cause a definite weight loss. It becomesunderstandable that a continuationof adverseweather of this sort might greatly decreasethe survival of young. Kullenberg (1946) is probably correct in thinking that the absenceof the Arctic
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