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Breeding scheduleand primary moult in Dunlins 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 migrate southward from thebeginning of August,when the moult is almostcomplete. On Kamchatka(kistchinski) and (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, .

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 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() 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 theranges of 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 Kitschinski et al. 1983). There are also a few recordsof even wadersand are usedin Figure 1 to illustratethis in Dunlinsin earlierhatching dates from the arcticcoast of Chukotka:27 four areas of the Far East. It can be seen that the latest and the Junein Enurmino (67 ø) (Portenko1972) and even 24 Junein 31

I I datesindicated for KamchatkaDunlins in Figure 1. Although far from beingcomplete, the datalisted above suggest that the datesin Figure 1 reflect the real situationin populations,at least

40 for peaksof hatchingdates.

Primary moult Chukotka •0 The largestset of dataon primarymoult camefrom Dunlinsof the coast of Chukotka, and these allowed the most detailedanalysis. It beginsin mid-June:no birdswere found 2O moultingbefore 13 June,while they were recordedon 14 June bothin Uelen andon BelyakaSpit. Almost all Dunlinswere in moult by 21 June. Theseresults are identicalto thosereported

10 by Holmes (1966b) from Barrow,north Alaska. Only two non- moultingfemales were caughton later dates,on 29 Juneand 5 July. These,and two more outlierswith delayedmoult, (female on 20 July and male on 11 August)were all found in Uelen. Thesefour birdswere excludedfrom furtheranalysis, and their I I I origin is discussedbelow.

Analysisof covarianceperformed on samplesfrom Uelen and Belyakarevealed no differencesbetween sexes in moult score 20 (p=0.163). This allowedpooling of the data from malesand femalesand addingof unsexedbirds to the samplefor further analysis.Dunlins in Uelen have a close,but statisticallyhigher, averagevalue of moult score(p=0.013) comparedwith thoseon BelyakaSpit, indicatingearlier primary moult there(Figure 2a). Most birdschecked after late July might be on their post- ß 4A ß ß 4A breedingdispersal and of uncertainorigin, as both local birds andmigrants from northAlaskan arcticola subspecies 1 o •0 20 9 DATE (MacLean & Holmes 1971; Norton 1971; Stiefel & Steufler JLRIE JULY AUGUr2 1989) could be encountered. The moult score of the birds Figure 2. Moult scorefor primariesof knownlocal Dunlinson a) Chukotka knownto be local andthose of unknownorigin differed andb) northSakhalin. Dots refer to Uelen,crosses to BelyakaSpit, triangles to north Sakhalin. substantially(p=0.01). This differencecan be inferredfrom the samplefrom BelyakaSpit, which includedmostly August birds Chaun Lowland, northeastChukotka (69 ø) (Kretchmar et al. of unknownorigin. We believethat thesebirds originated from 1991). Datesfor the startof breedingin high arctictundra on non-localpopulation(s). All checkedDunlins with completed WrangelIsland (71 ø) vary from yearto yearby up to two primarymoult belongedto birdsof unknownorigin. The first weeksdepending on the time of snow-melt.As a result,the Dunlin with completedprimary moult was caughton 5 August first songmay sometimesnot be hearduntil mid-June, on BelyakaSpit andon 6 Augustin Uelen, the last moulting althoughthe chicksstill hatchbefore mid-July (Stishov et al. of unknownorigin was recorded on 17 Auguston Belyaka. 1991), i.e. no later than on the mainland. There are almostno comparabledata for the Kamchatka The startingand finishingdates of primary moult on Chukotka Dunlin. In GekaBay (60ø),the northernpart of the kistchinski showthe earliestprimary moult amongall Dunlin subspecies, subspecies'breeding range, birds probably bred a little later in althoughprobably coming from birdsof differentpopulations. 1976-77, thanthe datesshown in Figure 1, because"mass Thus,it is possibleto usethe datesto determinethe durationof hatchingof chicksstarted in late June"(Firsova & Levada primary moult in birdsfrom Chukotka. It lastsfor 54-58 days, 1982). The few recordsof nestsand chicks, listed by Lobkhov a shortertime than in northAlaska, whereDunlins changetheir (1986) for Kamchatka,also fell into the perioddepicted in primariesin 60-70 days(Holmes 1966b). Figure 1, andhatching only occurredunexpectedly late, on 15 July,in a singlenest on the easterncoast of Kamchatka.Of Regressioncoefficients of primarymoult scoreson datedo not five local femalescollected on 30 and31 May 1983 by differ (p>0.1) betweenDunlins from Uelen andBelyaka, so the Gerasimovin the areaof the Moroshechnayaestuary, two had regressioncoefficient for the pooledsample of birdsknown to an egg in the oviduct,two were alsoegg laying, and only one be local is 0.904 _+0.056 (s.e.). As moult scorefor Dunlins of had not startedher clutch (collectionof ZMMU). Thus, Alaska was calculateddifferently by Holmes (1971), we cannot breedingin thesebirds was in completeagreement with the comparehis regressioncoefficient with ours. 32

hatching egglaying C. a. s•halina ,, 111111t . ( Chuko tka) moult J , primary

C. a. kist chin ski ( Kamohatk a )

C.a.actites I, I (North Sakhalin)

, ß !" '' ' ' I 0 20 1 0 20 1 o 2o June July August

Figure3 Overlappingof breedingevents and primary moult in different datesof startof moultin birdsof thesetwo populations are subspeciesof Dunlin in the Far East. closeto eachother and fall in thefirst half of July. The It is interestingto seethe similarityof theprimary moult in presenceof somenon-moulting birds in mid-Julyin Sakhalin Dunlins from different areas of Chukotka. The moult score of leadsto theconclusion that birds in thispopulation start 12 specimensfrom the ChaunGulf (collectionof ZMMU) does primarymoult over a moreprolonged period than in Chukotka. not differfrom that of Uelen andBelyaka (p>0.05). Moult scorevalues from severalspecimens available in the collection DISCUSSION from theAnadyr Lowland agree well with thosein Figure2. Holmes(1971) consideredthat the schedulingof themajor Thesedata indicate the similarityof primarymoult dates with energy-requiringevents of reproduction,moult and migration sakhalinasubspecies. is amongthe strategiesused in responseto a harsh environment. He was able to show that at Barrow, the further Kamchatka and Sakhalin northof two Alaskanareas compared, breeding and moult are There are not enoughdata from Kamchatkaand Sakhalinto compressedinto a shorterperiod and necessarily overlap. analyseprimary moult of Dunlinsin the way describedabove. Resultsfrom the Far Easthave shown some similarity with the This is partly becauseof the later startof moult in birdsof situation in Alaska. more southerlyregions. In particular,no Dunlinschecked in Junein Kamchatka(n=19) and Sakhalin (n=3) had startedtheir In Dunlinsfrom Chukotka, unlike more southerly populations, primarymoult. Only two malesof eightkistchinskii birds from primarymoult broadly overlaps with breedingand is Kamchatkawhich were checked on 1-21July (mainly on 1-9 compressedin time (Figure3). However,in spiteof thegreater July),had startedtheir primary moult, both having a moult latitudinalrange, 13ø-15 ø in theFar Eastcompared with 10ø in scoreof 10 on 2 and9 July respectively.Six of 10 malesof Alaska,the differences in breedingdates in differentFar Sakhalinsubspecies actites, collected from neartheir chicks in EasternDunlin populationsturned out to be the sameor several July,were in primarymoult (Figure2b). The latestdate on daysearlier than those in Alaska. In southernpopulations of whicha bird from thissmall sample had not startedmoulting theFar East,in contrastto Alaska,the breeding season is more was 19 July. Thesedata on SakhalinDunlins differ markedly compressedthan in the north, on Chukotka. This effectcan be from thosegiven in Figure 37 in Stiefel & Scheufler(1989), partlyexplained by shortageof data,although Blokhin (in becausemigrants of otherpopulations (e.g. all birdsfrom press)considers that Sakhalin Dunlins use spring water Augustto October)were undoubtedly included by these resourcesto raisechicks before the periodof summerheat, authors. droughtand fires. So,in the southernmosttemperate part of thebreeding range, Dunlins probably suffer sometimes from Althoughextremely scarce, moult dataon Dunlinsfrom evengreater time limitationson breedingschedule depending Kamchatka and Sakhalin are sufficient to conclude that the on the environment. 33

Breedingand intensive primary moult are both energetically responsiblefor thetiming of thebreeding season and primary costlyprocesses: temporal overlap of suchfunctions is rare in moult of Dunlins. If this suppositionwere correct,then the birds. Holmes(1971) considersthat with the long snow-free southernDunlin populationsof the Far East, situatedfurther periodat the breedingrange of the southernAlaskan souththan those studied in Alaska,should have progressively populationof Dunlin, thereis sufficienttime for moult to be earlierbreeding, completely separated from primarymoult. completedbetween the endof the breedingeffort andthe Comparisonof Figure 3 with breedingdates of Dunlinsin west autumnmigration, thus explaining the absenceof overlap Alaska (61 øN), where most chickshatched before 20-29 June there. The samecan be saidabout southern Dunlin populations (Holmes 1971), showsthat in the southernmostareas, in theFar East;these processes are usually separated there Kamchatkaand Sakhalin,breeding takes place not earlierbut also. It is difficult,however, to agreewith Holmesthat evensomewhat later, with no reductionin the overlapof breedingand moult necessarily overlap at Barrow(and also on breedingand moultingperiods. Chukotka). Dunlinsof subspeciesarctica from breedat evenmore northerly latitudes (700-80 ø ) and do not The resultsof this studydo not contradictthe general undergoprimary moult on thebreeding grounds (Cramp & conclusionof Holmes(1971) that featuresof the breeding Simmons1983). This meansthat northern populations of the seasonand the periodof moult are relatedto the lengthof time Pacificsubspecies of Dunlin couldalso, by evolution,have favourablefor breeding.However, this relationship is not avoidedthe overlapof breedingand moult, delaying primary directlylatitude dependent. The similarityin breedingdates of moultuntil winter. Neverthelessthis does not happen,and is Dunlinsin differentparts of Chukotkasuggests a common probablydue to historicalevents in thePacific populations of breedingperiod for the wholeChukotka subspecies sakhalina. Dunlin,resulting in overlappingof thesetwo processes. This, in turn, may indicatethat latitudinaldifferences in breedingand moult dates arise from differinggeographical In Dunlinsof Chukotka,as in northAlaska, primary moult populations(subspecies in manycases). The qualityof the almostcompletely overlaps with the breedingseason. As a environmentin somelocalised areas may resultin breeding rule, it startssoon after the beginningof the incubationperiod takingplace on earlier(e.g. ChaunGulf) or later (BelyakaSpit, andreaches about half way at peakhatching in thefirst week WrangelIsland) dates than those which are typicalfor the of July. Post-breedingdispersal of Dunlinson Chukotkafirst geographicpopulation range. In the light of thesedata on becomesnoticeable about a week afterhatching, before mid- populationspecific ranges of breedingdates, a recordof a July. However,Chukotka Dunlins probably continue and Dunlin pulluson 7 May in China (Greenwood1980) seems almostcomplete their primary moult either on the breeding completelyunrealistic. groundsor closeto them,for instanceon lagoonsaround Chukotkaand in river mouths(Kistchinski 1988). Departure It was noted earlier that moult score outliers were found on andlong-distance migration of Dunlinsfrom Chukotkaand the Chukotkaonly in the Uelen area. This fact couldbe due to the northernKoryak Highlands start only at the very end of July local breedingpopulation not beinghomogeneous. In an andthe beginningof August(Kistchinski 1980, 1988), and earlierstudy (Tomkovich 1986), it was emphasisedthat mostprobably it is thesebirds which initiate the mass unusuallylong-billed Dunlins can be foundamong migrationof Dunlinson Kamchatkain thefirst half of August C.a. sakhalina on the easterncoast of Chukotka Peninsula, in (Lobkhov1986). Thusthe beginning of the southward particularin Uelen,which was thought to indicatean influx of migrationof ChukotkaDunlins precedes the primarymoult long-billedbirds of subspeciespacifica from westAlaska to the completion(growing of the outermostprimaries). area. Bill lengthof two females(41.3 and43.2 mm) among four moult scoreoutliers from Uelen exceededthe rangeof Dunlins of Kamchatka and Sakhalin do not moult in the variationof thisfeature in birdsof the subspeciessakhalina. nestingperiod, and some males start primary moult only As relativelylate primarymoult is characteristicof pacifica duringthe chick-rearingperiod (Figure 3). Femalesof these birds,the presenceof moultscore outliers also suggests that Dunlinsstay with thefamily for only a few daysafter hatching smallnumbers of Dunlinsof Americanorigin are present in the andthen disappear from breedinghabitats (Nechaev 1991; local populationin the Uelen area. Gerasimovet al. 1992)to unknownareas. They may concentratefor furthermoult in nearbyestuaries and lagoons, The subspecificdifferences in datesof breedingand moulting or may undertakelonger-distance migration to special in the Far EasternDunlins can be usedtogether with biometrics moultingareas. The latteridea is supportedby the to distinguishadult birds of thesesubspecies in the post- observationsof Nechaev(1991) andBlokhin (pers.comm.) in breedingperiod. In southernareas, for example,in the first northern Sakhalin, where hundreds and thousandsof Dunlins half of August,Dunlins with completedprimary moult should form flockson shallowwaters of baysfrom the lastweek of belongto northerntundra populations (sakhalina and/or Juneand stay there at leastthroughout July, when the Alaskan arcticola), while thosestill in active moult shouldbe migrationof northernpopulations starts. kistchinskibirds if theyare large, or actitesbirds if theyare small. The first attemptto usethese findings for understanding Holmes(1971) thoughtthat latitudinal differences were migrationdifferences of Dunlin populationsin the Far Easthas 34 beenmade above. Much more canbe donenow, paying special Kistchinski, A.A., Tomkovich, P.S. & Flint, V.E. 1983. The birds of the KanchalanRiver basin,Chukotsky Autonomous Area. In: Flint, V.E. & attentionto the migrationand population differences outlined Tomkovich,P.S. (Eds.)Archives of Zool. MuseumMoscow State Univ. 21: 3- here. 76. (In Russian).

Kondratiev,A.Ya. 1982. Biologyof wadersin the tundrasof north-eastAsia. ACKNOWLEDGEMENTS Nauka, Moscow. (In Russian). Field studiesin Chukotkaand Kamchatkawere financedby the ZoologicalMuseum of MoscowState University, and State Kretchmar,A.V., Andreev,A.V. & Kondratyev,A.Ya. 1978. Ecologyand distributionof birds in north-eastUSSR. Nauka, Moscow. (In Russian). Nature Reserve"Wrangel Island" provided transport for studies on BelyakaSpit in 1986-88. Invaluablehelp in gatheringdata Kretchmar,A.V., Andreev,A.V. & Kondratyev,A.Ya. 1991. Birdsof northern in differentyears was received from V.V. Morosoy,A.I. plains.Nauka, Leningrad.(In Russian).

Oleksenko,M.Yu. Solovyevand A.M. Sokolov. Important Lappo,E.G. & Tomkovich,P.S. In press.Breeding distribution of Dunlin in informationand museum specimens were receivedfrom A.Yu. Russia. Blokhin, N.N. Gerasimov, V.A. Nechaev, E.G. Lobkhov and Lobkhov,E.G. 1986. Breedingbirds of Kamchatka.Far-eastern branch, V.A. Ostapenko.M.Yu. Solovievhelped greatly with statistics RussianAcad. of Sci., Vladivostok.(In Russian). and commentson the early draft of the paper. I am extremely gratefulto all thesepeople as well as to JeanPatterson for MacLean,S.F. & Holmes,R.T. 1971. Bill length,wintering areas, and taxonomyof NorthAmerican Dunlins Calidris alpina. Auk 88: 893-901. correctionof my English. Nechaev,V.A. 1991. Birdsof SakhalinIsland. Far EasternBranch, Russian REFERENCES Academyof Sci., Vladivostok.(In Russian). Blokhin,A.Yu. In press.Population of breedingwaders at the marinecoasts of Nechaev,V.A. & Tomkovich,P.S. 1987. A new subspeciesof the Dunlin north-easternSakhalin In: Hotker et al. (eds.) Migration and international Calidrisalpina litoralis ssp.n.(Charadriidae, Aves), from SakalinIsland. Zool. conservationof : researchand conservationon northAsian and Zhuma166:1110-1113. (In Russian). Europeanflyways.International Studies 10. Nechaev, V.A. & Tomkovich, P.S. 1988. A new name for Sakhalin Dunlin Browning,M.R. 1991. Taxonomiccomments on the Dunlin Calkkis alpina from northern Alaska and eastern . Bull. B.O.C 111: 140-145. (Aves,Charaddidae). Zool. Zhuma167: 1596. (In Russian)

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