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Breeding Schedule and Primary Moult in Dunlins <I>Calidris Alpina</I> Of

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Breeding Schedule and Primary Moult in Dunlins <I>Calidris Alpina</I> Of 29 Breeding scheduleand primary moult in Dunlins Calidris alpina of the Far East Panel S. Tomkoich Tomkovich,P.S. 1998. Breedingschedule and primarymoult in Dunlins Calidris alpina of the Far East. WaderStudy Group Bull. 85:29-34 On Chukotka,the breedinggrounds of the subspeciessakhalina, the primarymoult of Dunlin Calidrisalpina startsin mid-June,at the beginningof theincubation period, and birds migrate southward from thebeginning of August,when the moultis almostcomplete. On Kamchatka(kistchinski) and Sakhalin(actites), where breeding takes place earlier, primary moult startsin July,in the chick rearing period,or evenlater. Thesedifferences between northern and southern populations are similarto thosefound previously in Alaska (Holmes1971). The timing of theseevents is not strictlyrelated to latitude; it is thoughtthat, with localvariations, latitudinal differencesin datesof breedingand primary moult are brought about through subspecific characteristics. Knowledge of subspecific breedingschedules and primary moult periods can be usedtogether with biometricsto assignFar EasternDunlins to different populationsin the post-breedingperiod. P S. Tomkovich,Zoological Museum, Moscow Lomonosov State University,Bolshaya Nikitskaya Street 6, Moscow103009, Russia. INTRODUCTION Far East are comparedin this study. Two new subspeciesof Dunlin from the Far East, Calidris alpina kistchinski(Tornkovich 1986) and C.a. actites(Nechaev STUDY AREAS AND METHODS & Tornkovich1987, 1988), were describedrecently, in The main data were collectedduring expeditions in 1978-1980 additionto the well-recognisedC.a. sakhalina,which breeds to the vicinity of Uelen (66ø04'N, 17ø042'W), andin 1986- on Chukotka. Calidris a. kistchinski inhabits Kamchatka 1988 to BelyakaSpit, KolyuchinskayaGulf (67ø05'N, Peninsula,the easternpart of the northerncoast of the Seaof 17ø442'W), both on the northerncoast of ChukotskyPeninsula Okhotsk and the northernmost islands of the Kuriles and, in 1989, to the area of the MoroshechnayaRiver estuary, (henceforthreferred to as KamchatkaDunlin for brevity); the central West Kamchatka (54ø30'N, 15ø630'E). In addition, distribution of C.a. actites is restricted to coastal areas of informationwas obtainedfrom the literatureand from personal northernSakhalin Island (Lappo & Tornkovich,in press). The communicationson the breedingdates of Dunlinsin different validity of thesetaxonomic innovations is confirmedby the areas,especially the northof Sakhalin(52ø-53ø). The studiesof Browning(1991). collectionof the ZoologicalMuseum of Moscow University (ZMMU) was an additionalsource of data; skinscollected There is muchpublished work in Europeand North America recentlyfrom Sakhalinwere of particularinterest. concerningDunlins, but little is known aboutthe biology, moult, and migrationof birdsin the Far East. Only a few Descriptionsof the threemain studyareas are givenin regional papersdeal specificallywith Dunlinsin this region(Norton faunisticpapers (Tornkovich & Sorokin1983, Kretchmaret al. 1971; Gromadzka 1985; Stiefel & Scheufler 1989; Tornkovich 1978 and Gerasimovet al. 1992 respectively).However, it is 1994; Kim et al. 1994). Apart from these,data on breeding importantto realisethat Chukotkais an arcticarea covered datesand biology of the speciesare scatteredthrough Russian primarilyby tundra,while Kamchatkaand Sakhalin belong to faunisticpublications (the mostimportant are: Portenko1972; the temperatezone, where open, usually marshy or boggy Kondratyev1982; Lobkhov 1986; Kistchinski1988; Nechaev tundra-likelandscape predominates on coastalplains due to the 1991). Phenologyand numbersof migratingDunlins in the coolingeffect of the sea. It shouldalso be mentionedthat the Far East are also considered to some extent in faunistic studyareas at the ChukchiSea coasthave a severearctic sea publications.However, the dataare difficult to interpretas climateand deep snow, especially on the flat BelyakaSpit, therehave been various attempts to assignmigrants to whichis surroundedby icy sea. So, delayedbreeding of populations(subspecies) and, in any case,subspecific Dunlinscan be expectedthere compared with inlandareas of differencesin migrationschedule, migration routes and even Chukotka. winteringgrounds are to be expected.The main characteristics of migrationare linkedwith breedingand the moult schedule Breedingdates were obtainedfor Dunlinsfrom 1) nestsfound of birds' flight feathers. Knowledgeof the two latter features with still incompleteclutches, 2) recordsof hatchingin known canbe helpfulin distinguishingpopulations, and thus may lead nests,and 3) retrospectivecalculation for broodsof small to furtherunderstanding of Dunlin migrationalong the East chicks. Three more caseswere addedwhen egg-layingfemales Asian-AustralasianFlyway. Data on breedingdates and were collectedon Chukotka. The following characteristics primarymoult schedulein severalDunlin populationsof the 30 were usedin calculations:the intervalbetween laying of mostprolonged period of hatchingin a population(27 days) successiveeggs, incubation period (between laying and was recordedat the northernmostsite, Belyaka Spit, while the hatchingof the lastegg in a clutch),and duration of hatching shortestperiod (7 days)was found in the southernmost period(from the first signsof hatching- 'starring'-to the breedingarea, Sakhalin. It couldbe thoughtthat, in the latter hatchingof the last chick). Dunlinsusually lay eggsdaily case,this resultsfrom too small samplesize; however,data (Holmes 1966a;Kondratyev 1982; pers. obs.). An incubation obtainedby threedifferent observers in differentyears agree periodof 21 dayshas been taken, because in five nestsat closelyon hatchingdates at Sakhalin. Hatchingdate records northernChukotka it variedfrom 20.5 to 21.2 days,with an from the secondhalf of July on Chukotkaprobably belong to averageof 20.8 + 0.28 (s.d.) days. At Barrow,North Alaska, replacementclutches, as was shownfor Alaska (Holmes 1966a, Dunlinsincubate for 21-22 days(Holmes 1966a). The 1971), but thereis no goodevidence for this. The absenceof a durationof hatchingwas 2-4 days,with 3 daysbeing most largehatching peak anddelayed median hatching date on usual(Kondratyev 1982; pers. obs.). Chick agewas calculated BelyakaSpit (Figure1) areundoubtedly due to a localeffect of from personal,unpublished data on the developmentof marked the levelled area with gradualand late snowmelting (see chickswith known hatchingdates. above); severalwader species had anomalouslaying dates there (Whitfield & Tomkovich 1996). Moult of primaryfeathers was studiedin Dunlinscaught for ringingon theirnests, near broods or in feedingareas on the The following calculatedranges of datesof clutchcompletion banks of water bodies and in collected birds. A detailed correspondto the hatchingdates shown in Figure 1:9 Juneto 5 descriptionof moult stateof eachprimary feather was recorded July on Belyaka Spit, 7 to 27 Junenear Uelen, 30 May to 12 and this was usedlater for calculationof primary moult score Junein Moroshechnayaand 3 to 9 Juneon northSakhalin. accordingto Ginn andMelville (1983). In total 115, 104 and 23 moult scorevalues are availablefor Uelen, Belyaka and The medianhatching date on Sakhalinis later thanthat for Moroshechnayarespectively. Every effort wasmade to west-centralKamchatka (U=I 1, p<0.05, Mann-WhitneyU test) eliminatedata relating to migrantsfrom non-localpopulations in spiteof the moresoutherly position of the formersite. This which might havebeen present among birds caught in feeding areasor collectedduring the secondhalf of the summer.In particular,only thoseDunlins at Kamchatkawhich were caught asbreeders or shotwhile showingbreeding behaviour were 1986-88 Belyaka, Chukotka:67ø • considered.At Sakhalin,only Dunlinswithin the sizerange typicalfor actites(Nechaev & Tomkovich1987) were considered,to excludemigrants and possible over-summering birdsfrom otherpopulations. 1978-80 Uelen, Chukotka:660 N As the smallsize of SakhalinDunlins is diagnosticand biometricsfor only a few malesof the subspeciesactites have beenpublished, it is usefulto updatethe biometricdata on the 1989 •oroshecnnaya,Kamohatka: 55øN basisof currentlyavailable museum specimens. Mean bill lengthof malesis 29.0 mm (s.d + 0.99, n=12, range26.8-31.2), of females34.0 mm (s.d. + 0.40, n=3, range33.5-34.2), wing lengthof males108.5 mm (s.d.+ 2.03, n=12, range105.5- 1975, 1983, 1988 NE Sakhalin I.: 520- 53ø N 111.5),of females112.3 (s.d. + 0.99, n=12, range111-113). 5 10 15 20 25 Analysisof variance(ANOVA) waschosen as the principal June J u • y approachto studythe dependenceof moult scoreon the sex Figure 1. Hatchingdates in nestsof Dunlin in differentareas of the Far East. and origin of birds. To allow for the influenceof calendardate, Arrow indicates the median date for an area.. it was included in the model as a covariate. The moult score, thoughnot continuous,was normallydistributed and justified raisesthe questionof how representativethe dataare, the useof parametricstatistical methods. All analyseswere especiallyfor Kamchatkawhere the few recordsthat existwere carriedout usingSYSTAT 5.0 (Wilkinson1990). collectedduring a singleseason. Breeding dates available for Dunlin from faunisticpapers for the Far East mostlyfall into RESULTS the rangesof datesgiven in Figure1. Moreover,the breeding Breedingschedule dates of Dunlins in warmer inland areas of Chukotka fall Hatchingdates are usuallythe mostabundant data that canbe mainly within the sameperiod as in the Uelen area(e.g. obtainedin studiesof the breedingschedules of northern
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