Internahonal Wader Studies 10. 152-169

Breeding distribution of Dunlin Calidris alpina in Russia E.G. Lappo & P.S.Tomkovich

Lappo,E.G. & Tomkovich,P.S. 1998. Breedingdistribution of Dunlin Calidrisalpina in Russia. International Wader Studies 10: 152-169.

Breedingrecords and breeding densities available from the literature, museum collections and unpublisheddata are summarised for six subspeciesof Dunlin Calidrisalpina in Russia.Differences in distributionand habitatsfound in subspeciesand populationsare usedto outlinetheir breeding ranges.Each subspecies or populationhas core areas where demities are high and habitats most diverse,but it was not possibleto discoversuch core areas for all subspecies.Maximum densities arenot thesame in differentsubspecies, thus centralis never reaches demities that are as high as foundin alpina,sakhalina and kistchinski. Coastal areas are often the most densely populated in Europeantundra and in theFar East. Dunlinavoid dense stands of dwarfbirch possibly because largehydrophilous Enchytraeidae worms, their main food, are scarce there. Thisis probablythe reasonfor Dunlin havinga patchydistribution in southerntundra; only isolatedbreeding localities on bogsare known in areaswhere large dwarf-birch tundra belt is present.

E.G.Lappo, Department of Biogeography, Institute of Geography,Russian Academy of Sciences, StaromonethnyPer. 29, Moscow109017, Russia. P.S.Tomkovich, Zoological Museum of Moscow State University, Bolshaya Nikitskaya Street 6, Moscow 103009, Russia.

Aanno,E. F., TOMKOBW•, Fl.C. t998. FHe3Aosoe pacnpocTpaHeHue qepHo•6uKa Calidris alpina • Po½cuu. International Wader Studies 10:152-169

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Introduction Abouthalf the totalbreeding range of Dunlin is in Russia,with six subspeciesdistributed from Beinga circumpolarspecies, Dunlin Calidrisalpina temperateto high arcticareas (Stepanyan 1990), so hasunprecedented geographic variability among onecan expect that generalphenomena can be arcticand subarcticbirds (Cramp & Simmons1983) tracedin thispart of the range.Available published with up to tensubspecies recognised. Knowledge descriptionsof breedingdistribution of Dunlin in of itsbreeding distribution is a basicrequirement thisvast area are incomplete either because they for otherstudies and conservationpurposes: it is were basedon now partly obsoletedata (Gladkov surprising,therefore that althoughshort 1951;Kozlova 1961; Uspenski 1969) or becauseof descriptionsof thisimportant feature can be found brevity(Stepanyan 1990). Uspenski(1969) was the mainlyin regionalpublications or generalised brieflyin handbooks,there have been no detailed firstauthor to outlinethe breeding range of Dunlin in Russiawith the help of vegetationmaps, but gave analyses. no evidencefor his generalisationfrom recordsnor did he outline the methods used. A different

152 Lappo& Tomkovlch.Breeding distribution of DunlinCahdns alptna in Russia analysisof Dunlin distributionfor north-eastern densities than when other methods are used. Asia was givenby Kistchinski(1988), based on data collectedbefore the 1980s.He distinguishedthe Transectcensusing methods with fixed or variable corearea and more local areas with high densitiesin distances to recorded birds are in wide use in sakhalinasubspecies and paid attentionto the Russia.Data obtainedthis way shouldbe more differences in distribution of birds of centralis and representativeof populationsin widespread,not sakhalinasubspecies. Nevertheless, there is no local,areas. However, not only are suchdata detailedaccount of thebreeding distribution of influencedby the experienceand methodsof the Dunlin in Russia,and this paper aims to fill the gap, observer, but also we do not know how data aswell assummarising more general patterns of the obtainedin thisway relatesto realfigures. Recently speciesdistribution. an attemptwas madeto checkthe latterissue (Soloviev1995; Soloviev et al. 1996). Table 1 shows Methods how widelybreeding densities may vary between observersusing similar transect methods (Ravkin Data aboutbreeding records, densities and habitats 1967) and between transectresults and those were takenfrom personalobservations from 1972- 1992 in different Siberian tundra areas,but the main obtainedby mapping.It canbe seenthat, on bulk of data was obtained from numerous VrangelIsland, Dunlin densitieswere found to be five timeshigher by Stishovet al. (1991)than those publications,mainly in Russian.Eggs and reportedby Dorogoi(1982) with both observers unfledgedchicks from the collectionsof several usingthe sametransect method. Data from the EuropeanMuseums were usedadditionally as ChukotskiPeninsula show that mappingproduces breedingrecords. The mostvaluable data were figuresseven or eighttimes higher than those foundin the ZoologicalMuseum of Moscow resultingfrom transects(Table 1). University (ZMMU), the BritishMuseum of Natural History (BMN-H),and the ZoologicalInstitute of Dunlinsshow strong site-fidelity, at leastat Yamal, RussianAcademy of Sciencesin St.Petersburg (ZIR). Taimyrand Chukoski Peninsulas (Ryabitsev 1993; Tomkovich1994; Soloviev et al. 1996;Pyabitsev & Alekseeva1998) and, as a result,there are no The precision,and sometimesquality, of the data indicationsof largeyear-to-year variations in differsconsiderably among publications. Therefore, numbersin differentlocalities: any changes are recordsof nests,unfledged chicks and fledglings,as probablydue to variationsin breedingsuccess and well asfemales containing eggs when they were differencesin departuretimes. The largest variation collected,were considered as cases of proved of densitiesrecorded was only three times (0.67 - breedingin a locality.Records of singingmales, 2.0pairs/km 2) during a tenyear study on a plot of territorialpairs, birds with distractiondisplays or 4.5 km2 in typicaltundra on theYamal Peninsula alarmingconstantly (characteristic behaviour near (Ryabitsev1993). Thus,the between-observer broods)and signs of familieswith fledgedyoung differences on Dunlin densities for the same areas constituteprobable breeding. In somereports can,in mostcases, be explainedmainly by different breedingcriteria are not stated;although in some methods of censusand calculation. Therefore, we casesexact records were used, in others this was not correcteddata for thepurpose of thispaper and the caseand sopossible breeding was assumed reducedStishov's figures by 80%. (Figures1 and 2). Thereare other sources of errorsin breedingdensity Apart from rangecontraction in the subspecies estimates. For instance the transect census method schinziidue to human activities(Malchevski & does not differentiate between local breeders and Pukinski 1983),no reliable data are availableabout migrantsin a summerbird population(Soloviev long term-changesin the breedingdistribution of 1995,pers. comm.), and usuallypopulations of Dunlin in Russia.Changes during this century approximatelythe firstand the secondhalves of a foundby comparingdistribution maps in summerseason are consideredseparately by handbooksare the resultof improvedknowledge, not in bird distribution. Hence we have combined observers.Authors of publicationsdealing with the data collected in different decades and even censusresults often do not give detailsof the status centuries. of the birdsunder study and thisleads to further uncertainty.We feel that some of theunusually high breedingdensities reported result from censusing Many differentmethods have been used for migrantsas well asbreeders. In particular,at least censusingand calculatingdensities and thismakes sometransect census data of Lobkov (1986)for comparisonsbetween studies very difficult. KamchatkaPeninsula and of Estafiev(1991) for the Densitiesobtained by territorymapping on defined north-eastEuropean tundra seem too high. studyplots should give the mostreliable data when As realdensities of breedingDunlins cannot usually carriedout accuratelyon quitelarge plots and at the be established,we decidedto simplifythe databy appropriatetime. Nevertheless,these results are assigningthem to low,moderate or high density probablyunder-estimates (Soloviev et al. 1996)and categories.This reduces errors resulting from showonly local densities unsuitable for large-scale different census methods and allows the inclusion of extrapolations.It is, likely that thereis a lower faunisticdata, where densities were evaluated only marginof errorbetween mapping data and actual

153 International Wader Studies 10: 152-169

Ira]Calidris a. schinzii 71Proved breeding ß C.a alpina 71Probable breeding 0 C.a. alpina xC.a. centralis 71 Possiblebreeding 0 C.a. centralis • C.a. sakhalina

:.....:.% "':'" ' •:••'t•7•.'• '"•- .••./.• -,• • • '•. .:: :..• '•.•"•:• '•• ' •.• ':6•:o; •' •":.:, :- ß ' ...., '" '.:',. . '• ,, -• ,. ,...... , ,,...... ,. ..

Figurela. Breedingrecords for Dunlin in Russia(Finland Gulf, European Russia, West and Central Siberia). Numbersrelate to siteslisted in theAppendix. asrare, common or abundant.By using all available someregions. These were Gribova et al. (1980)for data,it ispossible to determinetheir approximate northEuropean Russia and Sochava (1976) for west rangesof eachrelative density category (Table 2). Siberia.

To reconstructthe breeding range of Dunlin, we Werecognised a corearea (optimal part according usedthe landscapeextrapolation method (Brunov to Brunov1982) of the species'breeding range, 1982)or extrapolationwith the help of vegetation whichis characterisedby severalfeatures (Brunov maps.For this purpose we useddata about both the 1982).In thisstudy, only two featuresof thesewere presenceand absence of breedingrecords and also usedto discoverthe coreareas: the highestdensities informationabout breeding habitats. We assumed of birdsand thelargest diversity of habitatsused for that thespedes inhabits the entirearea with the breeding.The maximum level of densityfor sametype of vegetation(broad habitat category) recognitionof the corearea was lower in centralis, shownon the vegetationmap. Theborders of the thanin othersubspecies (see Discussion). breedingrange were determinedin mostcases Preliminarydata about varying needs of different accordingto tundrasubzones and their type of subspeciesof Dunlin (Kistchinski1988) alerted us to vegetation.Different approaches to tundrazonation this difference. Therefore distribution, densities and existin Russia,but we followedthe terminologyof habitatswill be consideredaccording to subspecies Gorodkov (1935)included in Chemov (1985):three and evengeographic populations within a subzonesare recognisedwithin tundra - southern, subspecieswhen possible. Subspecific status of typicaland arctic.In the Europeanpart of Russia, Dunlinsfrom the Taimyr Peninsulais not yet quite southern tundra is often divided into two belts: clearso the Taimyrpopulation is considered southern(large dwarf-birch) and northern(low separately. dwarf-birch) (Gribova et al. 1980). Results Thevegetation map of theUSSR (Belov et al. 1990) C. a. schinzii was usedfor mostextrapolations. However, several othermaps turned out to be moredescriptive for Distributionis sporadicand associatedwith the

154 Lappo& Tomkovich:Breeding distribution of Dunlin Calidrisalpina in Russia

180

174 182 158 1 H 83 159

•86 165 164 •192

162 p 160,

199

• ,::153 156. .• 137 147,149 •;7 132133 •_'1,'• 145 201 220 202

......

= .,. 2•)1 ...... 206 ' '""' 217 ' [ • •• 2•0229 224 II ' II 216 • 209 .• •. •• 231 :• 25 ß.) ..... '.•-.'. :..'.d '• ,10

211 =' ß. ,,.'=.., :'=... 213••212 Figurelb. Breedingrecords for Dunlin in Russia(East Siberia, Far Eastand SakhalinIsland). Numbersrelate to siteslisted in theAppendix. Balticregion (Figure 1). Breedingis possiblefurther 11in Figurela), they are evenlocally numerous south-eastthan was previouslythought, more wherebogs with peat hillocksare available. Moss- preciselyat the floodplainof the Ilmen Lake, sedgelake-side marshes and tussocktundra are NovgorodRegion (site 8 in Figurela) with unique amongother favoured habitats. water table fluctuations (Mistchenko & Sukhanova 1998). No densitiesare available. The few recent In the regionbetween the White Seaand the recordswere all made at grazedlowland meadows, PechoraRiver, Dunlins are widely distributedin but birdsbred alsoon lightlyand moderately tundralandscapes but in varyingnumbers. They vegetatedmarshes in tussocksin the late 19th are rare in the southernmost coastal tundra areas century(Zarudny 1910). (Figurela) and were not foundbreeding in the southernpart of the Kanin Peninsula,but were C. a. alpina commonfurther north. The only mainlandarea Only a few breedingrecords are knownfrom the whereDunlins were reportedas commonwas Kola Peninsula.The speciesbreeds in tundraareas Shoyna,NW Kanin (site 18 in Figure lb; Avdanin, which are spreadover northernand easternparts of Filchagov,Leonovich, Vinogradov & Glukhovsky the region. Durdinsalso penetrate the south-eastern pers.comm.), but the areanorth and north-west part of the peninsulafollowing patches of damp from the Pechora River delta also seems to hold tundra alongthe seacoast. Distribution is not goodnumbers. The speciesis especiallyabundant uniformwithin the rangebecause of the on KolguevIsland (Morozov& SyroechkovskyJr., mountainousarea where Dunlins may be too sparse pers.comm.). Damp tundra of differentkinds forms to be found easilyor do not breedat all. No breedinghabitats, but the concentrationnear $hoina breedingis knownin theforest tundra. The tundra is on the fringe of the coastalmarshes. lowlands with lakes in north-central and eastern Kola Peninsulaare the only known areaswhere Furthereast in Europe,the southernlimit of the Dunlins are common(Mikhailov 1993;Filchagov Dunlin breedingrange according to Morozov (in pers.comm. - seeFigure 2a). Within the former press)passes within the largedwarf-birch tundra area,in the upper reachesof the YokangaRiver (site belt recognisedas the southerntundra subzoneof

155 International Wader Studies 10: 152-169

ß Low density ß Moderate density •1I Highdensity

ßLow density ""• Moderatedensity •'•'•.:

Figure2a. Densitiesof breedingDunlins in European Russia and West Siberia. •11Highdensity Figure2c. Densitiesof breedingDunlins in the Russian Far East.

ß Low density grasses,with higherdensflies on uplands.Further ß Moderatedensity easton the Tazovskiand GydanskiPeninsulas, the •11High density speciesdistribution is similarto thatoutlined for Yamal,but the densitiesreported are lower (Figure 2a). Dunlinswere not foundbreeding on Tazovski withinthe large dwarf-birch tundra belt, they were rare in the low dwarf-birch tundra belt of the southerntundra subzone of Gydanski,but were common to the north. In the arctic tundra of SibiryakovIsland (site 84 in Figurela), theywere quiterare (Figure 2a). Moss-sedgetussock tundra on slopes,and damp tundra with poolsin depressionsor lakesare among the best breeding habitats(Chernichko et al. 1994).

A separatebreeding population of Dunlin,probably belongingto thenominate subspecies, was found recentlyon extensive palsa bogs sometimes with •c•.:•.•'•'•.c•..•.,•.•,;,•.'.-•'v"'•• • ...... • • **•.• ...... •.'-.•.:..: :.:d• ...... •-•,-,•-..• scattereddepressed pine treesPinus sibirica within Figure2b. Densitiesof breedingDunlins in Centraland the northerntaiga natural zone of WestSiberia East Siberia. (Vinogradovet al. 1991).Densities vary, locally theregion. However, only isolated breeding records reachingmoderate values. No areasare known are known within this belt. The spedesis common wherethe palsa bog and the tundra populations of onlyin the typicaltundra subzone of boththe Dunlin meet. mainlandand VaigachIsland (Figure 3). Further south,it is commononly in coastalareas and is only C. a. alpina>< C. a. centralis a rarebreeder inland. Theprefered habitat is Dunlinsare widely distributedthrough the Taimyr Sphagnumbog with sparsecotton grass (Eriophorum Peninsulathrough the southernand typical tundra spp.)and sedge(Carex spp.) vegetation, and with subzonespenetrating along large rivers, western flat peathillocks. It seemsthat Dunlins are locally and easterncoasts into southernparts of the arctic numerousat coastalmoss-sedge tundra with lakes, tundrasubzone and evenon SverdrupIsland (site but it is difficult to be certain from the data 85 in Figurela) within thehigh arctic. No Dunlins presentedby Estafiev(1991). werefound in openbogs within forest-tundra, northerntaiga, or in mountaintundra on Putorany Dunlins are known to breed in almost the entire Plateau.There was only a singlebreeding record of YamalPeninsula. They arevery rarein southern a singlebrood at thenorthern limit of forest-tundra parts,but commonand locallynumerous in the (site105 in Figurela). centraland northern parts of thepeninsula (Danilov et al. 1984). The birdsare mostabundant in plain High densitieswere found in suitablehabitats in moderatelyhumid areasof tundrawith sedgesand mostplaces studied in westernand central Taimyr

156 Lappo& Tomkovich:Breeding distribution of Dunlin Cahdrtsalptna in Russia

(Figure2b) within the typicaltundra subzone and subspeciesstarts further east (Tomkovich 1986), near the Dudypta Rivermouth, western Taimyr, occupyingAion Island,Vrangel Island, northern southerntundra subzone(site 106 in Figurela). coastal mainland, Chukotski Peninsula and the Within typicaltundra at easternTaimyr, Dunlins are Anadyr lowland. Chaun lowland in the north-west commonnear the TonskoyeLake (site126 in Figure of theregion supports a high-densitypopulation in la) in southerntundra. In bordersites of the range, a rangeof tundrahabitats (Kondratiev 1982; densitiesare low, and an averagedensity (Figure 2b) Kretchmaret al. 1991). In a part of the northern was only found in the Pyasinadelta area(site 96 in coast of the mainland with arctic tundra, Dunlins Figurela). Polygonalbogs and areaswhere dry are commonin severalareas and they are even tundrais interspersedwith bogsand marshes have locallynumerous in arctictundra on VrangelIsland. high densitiesof Dunlin. Only patchesof such Similarly,they are locallyabundant in different complexhabitats are occupied by Dunlinsat the coastal areas of the Chukotski Peninsula and in northernborder of thebreeding range (pers. obs. at coastalparts of theAnadyr lowland (Figure 2c), sites85, 89 & 126in Figurela). Bothuplands with which belongto the typicaland southerntundra drierhabitats and floodplains in areaswith subzones.Birds are mostabundant in placeswhere generallyhigh densitiesare less populated. tussocktundra is formedmainly by Eriophorum vaginatumand is interspersedwith marshy C. a. centralis depressionsand lakes.They readily use polygonal Thissubspecies, not oftenrecognised outside Russia, bogsand bogswith peathillocks which are usually coversan areafrom at leastthe KhatangaRiver in not extensivein the region. Our personal observations have shown that dwarf willow Salix the west to the Kolymadelta in the east. Breeding distributionis not fully known. Nevertheless,it is fuscescensis characteristic of preferredhabitats on the Chukotski Peninsula. clearthat in thisregion, as in the Taimyr,Dunlins are distributedfrom the tree-lineformed by Larix dahuricanorthward through the southernand Dunlins are rare, or only locallycommon, in typicaltundra subzones,and were found in mountainareas with predominantlydry habitats, southernparts of the arctictundra close to the suchas in theAmguema River valley (sites171-173 IndigirkaRiver deltaand nearthe AriabarRiver in Figurelb), (Portenko1972; Dorogoi 1993a). This mouth (Figurela). However,the specieswas not partlyexplains the absenceor low densitiesof recorded in the southern tundra subzone near the Dunlins in valleysless than 100km inland from the Ariabar River (Gladkov & Zaletaev 1965) nor in the northernmainland coast:Ekaenmyvaam River (69ø Yanadelta (Kistchinski1988; Syroechkovski Jr., pers. 73'N; 177ø 10'E),lower and middle EkvyvatapRiver comm.).Spring records on BolshoiLyakhovsky (sites168 & 170in Figurelb), Kychakvaamand Island (Rutilevsky1958) do not necessarilyindicate PegtymelRivers (65 ø 35'N 177ø 50'E), and lower possiblebreeding on theNew SiberianArchipelago. KaveemRiver (69ø 40'N 174ø 30'E) (Stishov pers. Birds do not breed within the forest-tundra even on comm.). Dunlins,do howeverbreed, although at extensivetundra-like bogs (Kistchinski 1988; low density,in mountaintundra at elevationsof Sorokinpers. comm.). 450-500m a.s.1.in the ElgygytgynLake depression (site198 in Figurelb; Dorogoi1993b) in the central Unlikeboth the Taimyr population and sakhalina part of the mountainregion. The species was also subspecies,no areasare known where centralis foundbreeding at low densityin non-floodedparts reacheshigh breeding densities, even locally. The of the plain of the middleAnadyr River (sites196 & only sitewith a locallyhigh breedingdensity (12.7- 197in Figurelb), whereBeringian forest-tundra is 16.9pairs/kin 2on a plotin differentyears) was characteristic.Birds occupy extensive tundra-like foundnear the mouthof the KhatangaRiver (site polygonalbogs and lake-side marshes there, among 128in Figurela) (Soloviev1995; Soloviev et al. 1996) areascovered by sparsedwarf pine treesPinus probablybelonging to theTaimyr population. Birds pumilaand tall shrubsof Alnusfruticosa,Betula were foundto be commononly in a few placesin Middendorffii(Kretchrnar et al. 1991). the westernhalf of thesubspecies range (Figure 2b). Theyoccupy plain tundra with a mixtureof damp, C. a. kistchinski wet and dry areaswith dwarf shrubs;these are Thisrecently recognised subspecies (Tomkovich usuallypolygonal marshes or marsheswith peat 1986;Browning 1991) is knownfrom the Kamchatka hillocksor ridges. WhenDunlins breed in dry Peninsula,the southernKoryak Highland, the tussockor moss-lichentundra, this is alwaysclose to northern coast of the Sea of Okhotsk and the marsheswhere they feedand to whichthey lead northernKuril Islands(Figure lb). Lack of data their young(Kondratiev 1982; Leonovich pers. precludesdetails of distributionbut it is clearfrom comm.;pers. obs.). Even in the southernmostareas the countsof Lobkov(1986) and brief personal centralisavoids habitats with tall shrubs (Kistchinski experience,that the lowlandcoastal belt with tundra 1988;pers. obs.). landscapeand numerous bogs and lakes along the westernside of Kamchatkais quitedensely C. a. sakhalina populatedby Dunlins.These waders have the same No Dunlins are known to breed for at least 150 km highbreeding density in lowlandswith similar eastfrom the KolymaDelta (Tundra Ecology -94 habitatson the Kamchatkaisthmus and probably expeditiondata). Thebreeding range of sakhalina nearGizhiga (site 220 in Figurelb), north-eastof the

157 International Wader Studies 10: 152-169

Seaof Okhotsk(Figure 2c). Mountainouslandscape placeswhere such habitats are in forests(mainly with woodlands in eastern Kamchatka and on the Betulaermanii) or are coveredwith scatteredtall northern coast of the Sea of Okhotsk, and the shrubsof Alnuskamchatica and Salixspp. Lobkov mountainsof KoryakHighland do not leavemuch (1986)states that the densityof Dunlinson spacefor Dunlinhabitat. Only small patches of Kamchatkais directlyrelated to the wetnessof the coastallowlands, usually near the mouthsof rivers tundra habitats and the number of lakes and near lagoons,are inhabited by Dunlinsin these areasresulting in a discontinuousbreeding range. C. a. actites Nevertheless,Lobkov (1986) considersDunlin as a Thebreeding grounds of Dunlin on SakhalinIsland breedingspecies of EasternKamchatka also at were only foundin 1975(Nechaev 1979, 1991) and altitudes of 400 to 650 m a.s.1. near lakes and in thebirds were describedas a separatesubspecies, mountaindepressions (site 208 in Fugurelb). very distinctfrom sakhalina and kistchinski (Nechaev & Tomkovich 1987, 1988). There are no indications In theKoryak Highland Dunlins are known only sofar of a possibleconnection between the breeding from coastalareas and arecompletely absent from rangesof actitesand kistchinski;breeding Dunlins inland mountaintundra valleys even in quite were not found near tidal meadows north from the suitablehabitats (Kistchinski 1980,1988; pers. obs.). Amur Rivermouth on the mainland(Babenko pers. Thereforethe breeding ranges of kistchinskiand comm.). On Sakhalin,Dunlins occupy tundra-like sakhalinaseem connected only by a chainof narrow boggyhabitats with numerouslakes spread on flat coastallowlands inhabited by Dunlins,although islets, and in narrow coastal areas of the north-east onlya few breedingrecords are known from there. and probablyin a few placesin the northernand north-westernparts of the islandwhere numerous A wide rangeof breedinghabitats has been lagoonsand shallowbays occur. Crowberry describedfor birdsof this subspecies.They breed Empetrumsibiricum and dwarf willow Salix on dry placesin areasflooded irregularly by sea fuscescensare characteristicfor elevationson bogs water (Leonovichpers. comm.), sometimes even on whereDunlins place their nests(Nechaev 1979, gravelwith sandspits, on coastalmeadows with 1991;Nechaev & Tomkovich1987). Blokhin (1998) dwarf shrubs(Gerasimov & Vyatkin 1973)and on statesthat Dunlin beginto breedwell beforethe differentkinds of bogs. The main habitat,however, vegetationof helophytesstarts, and, asa result,their is the tussockytundra with dwarf shrubsand with populationis ratherstable even in dry seasonsand lakesinterspersed with pools,as well asbogs with birdsare ableto raisechicks before the periodof poolsand ridges.Dwarf willow Salixuscescens was summer heat. foundin preferredhabitats in WesternKamchatka (pers.obs.). Frequentl• Dunlins can be foundin A locallymoderate density in ChaivoBay (site 228 Table1. Comparisonsof breedingdensities of Dunlinson thesame or nearbyareas obtained by differentobservers and/or methods.

Sibiryakov Island, Yenisei Bay Coastal lowland tundra plot c. 2.0 Chylarecki& Sikorapers. comm. transect 1.5 Koshelevpers. comm. LogataMouth, Central Taimyr Polygonaltundra transect 68.0 Gavrilov pers.comm transect 16.1 Chupin pers.comm. Malaya LogataMouth, C. Taimyr Polygonaltundra transect 44.9 Chupin pers.comm. transect 17.8 Gavrilovpers. comm Novorybnaya, SE Taimyr Mosswatershed near plot 16.9 Soloviev (1995) polygonaltundra transect 4.6 Soloviev (1995)

Vrangel Island Moss-sedgetussock tundra transect 15.0 Dorogoi(1982) transect 80-150 Stishovet al. (1991) Northern Chukotski Peninsula Dry patchycoastal tundra and plot 2-5 Tornkovich,unpubl. dump tussocktundra transect 10-50 Kondratyev(1982)

Uelen, Chukotski Peninsula Tussock tundra transect 2-3 Kuzyakin in Portenko(1972) plot 17.8-23.4 Tomkovich & S9rokin (1983)

158 Lappo& Tomkovich:Breeding distribution of Dunlin Calidrisalpina in Russia

Table 2. Rankingof breedingdensities of Dunlins,characterised by differentsurvey methods.

Categoriesof density Plot and transect Transect Transect Faunisticdata findividuals/km•) (individuals/m) ...... • .• ;•. .. ':• ,•.... ;.:.. .• . ..•. •.• ß ..c,•. :, .,...,•.

Low <1 <2 <1 rare Moderate 1-10 2-20 1-5 common High >10 >20 >5 abundant

in Figurelb) was the only estimationfor this rare alpinaand centralisis likely,Dunlins are found again subspecies(Blokhin 1998). The total numberof in southern tundra close to the northern tree-line in birdsin the populationis c. 300breeding pairs gooddensities, but not on bogswithin forested (Nechaev & Tomkovich 1987). areas.The populationshows a tendencyto lower densitiesin the easternpart of the peninsulawhich Discussion probablyreflects the generalfeature of centralisnot to havehigh densitiesthroughout its range. The breedingdistribution of Dunlin hasbeen comparativelywell studiedin Russiaconsidering Unlike centralis,high densitiesare characteristicof the vastareas involved (Figure 1), but of courseit is far from the detailed information available for the easternmostsubspecies sakhalina and kistchinski in favourable habitats. This difference between westernEurope (Cramp & Simmons1983; centralisand sakhalinawas stressed by former Gjershauget al. 1994;Hagemeijer & Blair 1997). It is observers(Vorobiev 1963;Kistchinski 1988). unlikelythat the situationwill improvegreatly in Dunlinsof both Far Easternsubspecies mentioned the near future. Nevertheless,the general canbreed in boggyhabitats with tall shrubsamong characteristicsof speciesdistribution and, to a lesser extent, of densities are more or less clear for most of forests,although in low densities.It is surprising that neithernorthern subspecies nor the southern the regionsdescribed. These, together with the onehave spreadinto tundraof the centralparts of understandingof the environmentalneeds of the theKoryak Highland situated between their ranges. species,provide a fairly accuratepicture of the Coastalareas support the highest densities and the distributionof Dunlin by a generalisationof widest distribution of these Dunlins. The actites availabledata. As statedabove (seeMethods) subspeciesof SakhalinIsland is exclusivelycoastal Dunlin is a conservativespecies with strongsite with adultsbreeding on tundra-likebogs but tenacityand smalldensity fluctuations. This leads probablyfeeding for the wholeseason mainly on to a relativestability of breedingrange limits and a tidal areas,unlike Dunlins of all othersubspecies rarity of siteswith irregularbreeding, which whichrely almostexclusively on tundraor boggy happenswith someother waders (Kistchinski 1988; habitats. Ryabitsev1993). An interestinghypothesis was advanced by The differencesfound in distributionpatterns and Andreeva(pers. comm.) to explainthe southern environmentdemands of subspecies,and limit in distributionof tundraDunlin populations. sometimespopulations, of Dunlin mustbe bornein Accordingto her unpublishedstudies (see also mind. The Balticpopulation of schinziidiffers most Kondratyev1982; Kondratyev & Kondratyeva1988) from othersubspecies, being mainly a temperate largehydrophilous Enchytraeidae worms are oneof meadowbird. Nominatealpina is the most the few basicfoods of Dunlinsbreeding in tundra, polytypicsubspecies regarding distribution among and these worms are common or numerous in thoseunder consideration.Thus, the populationof damp or wet mosseswith sedges,cotton-grass and Kola Peninsulainhabits damp tundra and bogsjust sometimes with willows, but are almost absent in near the forest-tundra, while further east from soilswhere dwarf birches(Betula nana, B. exilis) Kanin Peninsulato the Yeniseyonly a few sporadic form a densemat asthese plants cause drying out of recordsare known within the largedwarf-birch belt the soil. Therefore, Dunlins cannot live in areas of southern tundra close to forest-tundra. In the with densestands of dwarf birch. The breeding latterregion, the main populationsare concentrated distributionof Dunlinsagrees well with this in typicaltundra and the southernmostpart of arctic hypothesis:the tall dwarf-birchtundra belt of the tundra subzones, and in coastal areas within the southerntundra dominatedby Betulanana found Europeanpart of the range. The Yamalpopulation between the White Sea and the Yenisei is almost free hashigh breedingdensities on watershedswhilst on of Dunlins,the few isolatedrecords (Figures 1 and the Taimyrthe main concentrationsof Dunlin are on 3) comefrom bogswhere dwarf willowsand other lowlands. The Yamalpopulation, as expected,spans dwarf shrubspredominate. When Dunlins were almostthe whole arctictundra of the peninsula, found closeto the northern tree-line (Kola whereasat the Taimyrthe speciespenetrates to the Peninsula,Taimyr, Indigirka basin, upper Anadyr), subzoneonly alongvalleys of largerivers. largeboggy areas are present. Similarly, Stishov (pers.comm.) did not find breedingsakhalina It is almostcertain that the breedingranges of the Dunlins in several inland areas close to the northern palsabog and tundrapopulations are separated in mainland coast(see Results) where the tundra has WestSiberia. On Taimyr,where an intergradationof

159 International Wader Studies 10: 152-169

Limits of subspecies recognized Breeding boundary range Isolated breeding records outside the main breeding range Core area of breeding range C.a. sakhalina ',

Figure3. Reconstructedbreeding ranges of Dunlin subspecies.

dense B. exilis stands. morphologicaldifferences). It remainsuncertain if Thesefeatures and peculiaritiesof distribution,were the coreareas shown for Kamchatkabelong to usedto outlinethe breeding range of Dunlin in differentgeographic populations, or to a singleone Russia(Figure 3). This final map is not exhaustive stretchingalong the westernside of the peninsula. of course,and further revisionis necessary.Known areaswhere the specieshas high densitiesare also Definitionof suchcore areas is interestingnot only indicatedon the map. Howeverfor centralis, from a scientificpoint of view, indicatingthe density characterisedin generalby lower densities,areas centresof populations,but is alsoimportant for with moderate,not high densitiesare shown. Birds conservationpurposes. These areas with high in suchareas usually occupy the largestdiversity of densitiesprovide the main stockof populations,and habitats,and that is why thesecan be consideredas the generalreproductive success and well-beingof coreareas of populationsor subspecies. thepopulations is dependentprimarily on the situationin thesecore areas. Any necessary The corearea of schinziimost probably lies outside conservation measures should be concentrated in Russia.Lack of dataprevents the delineationof core theseareas of majorimportance for the species. areasfor actitesand the palsa bog population of alpinain west Siberia. Of course,not all areaswith Acknowledgements high densitiesof breeedingbirds have been found We thanksincerely all our colleagueswho made for othersubspecies and populations,but the their unpublisheddata on Dunlin breedingdensities distributionof alreadyknown core areas, in our and/or distributions available to us: V.O. Avdanin, opinion,reflects the generalsituation quite well. A.Yu. Blokhin, Yu.Yu. Blokhin, G.C. Boere, Thus,among several small areas with high densities I.I. Chupin,P. Chylarecki, I.V. Dorogoi, in the Europeanrange of alpina,the easternmostone A.V. Filchagov,A.A. Gavrolov,V.V. Grichik, probablybelongs to the Yamalpopulation where the V.N. Karpov,Ya.I. Kokorev,A.V. Kondratiev, corearea can be expectedto coverlarger parts of the N.B. Konyukhov,A.I. Koshelev,Yu.P. Kozhevnikov, peninsulabetween 69 ø and 72øN. Westernand V.V. Leonovich, K.E. Mikhailov, V.V. Morozov, centralTaimyr contain extensive concentrations of V.V.Nikolaev, I.V. Pokrovskaya,A.A. Romanov,G.T. Dunlins. For sakhalina,considerable differences can de Roos, Yu. Roschevski, V.Yu. Semashko, A. Sikora, be foundin the distributionof the areaswith high M.Yu. Soloviev, A.G. Sorokin, M.S. Stishov, densitiesshown by Kistchinski(1988) compared E.E. SyroechkovkiJr., T.V. Sviridova, A.V. Tikhonov, with Figure3. This is the resultof a different V.G. V•mogradov,A.E. Volkov,P. Yesou and classification: areas where Dunlins were either V.S.Zhukov. We particularlythank T.R. Andreeva abundantor commonwere consideredby for beingpermitted to useher ideasabout the Kistchinskito be thosewith high densities.It is reasonsfor Dunlin distributionpatterns. We thank clearthat the populationin the ChaunLowland is S.Pushkaryov for hishelp with mapsfor our separatefrom that of the ChukotskyPeninsula and figures. adjacentareas (see Tomkovich 1986 for

160 Lappo& Tomkovich:Breeding distribution of Dunlin Cahdnsalptna m Russia

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Magadan,Institute for Biological Soloviev,M.Yu. & Syroechkovski,Jr., E.E. 1995. Wader Problems of the North. In Russian. MonitoringProject at the Taimyr:preliminary Pavlov,B.M., Beilmann,A.A. & Krashevsky,O.R. 1983. On resultsof comparisonsof censusingmethods. the avifaunaof the VerkhnayaTaimyra River basin. InformationMaterials of the Working Group on Waders In: B.M.Pavlov(ed.), Birds of Taimyr (ecology, 8: 11-12. conservationand practicaluse). Sci.-technicalBull., Sosin,V.E & Paskhaln)•S.P. 1995. Materialson faunaand SiberianSect. of the Acad. for Agric. Sci., ecologyof terrestrialvertebrates of BelyIsland. In: Novosibirsk.Issue 7, pp. 9-14. In Russian. V.S.Balakhonov (ed.), The current state of vegetation Podkovyrkin,B.A. 1956.Materials on breedingbiology of andanimal world of theYamal Peninsula, pp. 100-140. some birds at Northern Kuril Islands. Zool. Zhurnal Nauka, Ekaterinburg.In Russian. 35(12): 1892-1901. In Russian. Spangenberg,E.P. & Leonovich,V.V. 1960. Birdsof Portenko,L.A. 1939. Faunaof theAnadyr Land. Birds. Part northeastern coast of the White Sea. In: G.A. I. GlavsevmorputPress, Leningrad. In Russian. Novikov (ed.),Annals of Kandalaksha State Nature Portenko,L.A. 1972. Birdsof ChukotskyPeninsula and Reserve,2: 213-336. Murmansk Book Press, WrangelIsland. Part 1. Nauka, Leningrad. In Murmansk. In Russian. Russian. Stepanyan,L.S. 1990. Conspectusof the ornithological fauna Pozdnyakov,V.I., Solovieva,D.V. & Sofronov,Yu.N. 1996. of theUSSR. Nauka, Moscow. In Russian. Charadriiform birds in the Lena River delta. In: Stishov,M.S. 1990.In-landscape distribution of birdsin V.N. Vasilyev& V.I. Pozdnyakov(eds.), Soils, flora the typicaltundra subzone (based on the example andfauna in ArcticYakutia, pp. 54-65. RussianAcad. of the Aiyon island,West Chukotka). Zool. Zhurnal Sci., Yakut Sci. Centre, Yakutsk. In Russian. 69(9): 73-83. In Russian. Ravkin, Yu.S. 1967. On the censusmethods of birds in Stishov,M.S. 1993. Materialson faunaand bird population foresthabitats. In: A.A. Maksimov (ed.),Nature of on Aachimpeninsula (West Chukotka). Bull. of thetick encephalit hotbeds at theAltai, pp. 66-75. MoscowSoc. of Naturalists. Biol. section 98(4): 17-25. Nauka, Novosibirsk. In Russian. In Russian,with Englishsummary. Rutilevsky,G.P. 1958. Birdsof BolshoiLyakhovsky Island. Stishov,M.S., Pridatko,V.I. & Baranyuk,V.V. 1991. Birdsof Problemsof Arctic 4: 79-90. In Russian. WrangelIsland. Nauka, Novosibirsk.In Russian. Rogacheva,E.V. 1992. Thebirds of CentralSiberia. Husum SyroechkovskiJr., E.E. & Lappo,E.G. 1994.Data on fauna Druck und Verlagsges,Husum. and ecologyof birdsof the IzvestiyTSIK Islands Romanov,A.A. 1989. Birdpopulation of southeastern and SverdrupIsland. In: E.V. Rogacheva(ed.), coastof CheshskayaGulf. In: Yu.P.Yazan (ed.), Arctictundras of Taimyrand Kara Sea islands: nature, Organizationof theforms of nature-reserve fund objects,

163 International Wader Studies 10 ß 152-169

faunaand conservation problems 1: 111-150.Inst. Ecol. Uspenski,S.M. 1965. Materialson bird fauna of northern and Evol., Russian Acad. Sci., Moscow. In Russian. Anabartundra. In: N.A. Gladkov(ed.), Archives of Tatarnikova,I.P. 1983. Nestingof waderson Ainovy Zool.Mus. of Moscow Univ. 9: 63-97. MoscowUniv. Islands. Ornitologiya(Moscow) 18: 181. In Russian. Press, Moscow. In Russian. Tolmachev,A.I. 1928. To avifaunaof KolguevIsland. Year- Uspenski,S.M. 1969.Die StrandlauferEurasiens (Gattung bookof Zool. Mus. ofthe USSR Acad. of Sci.27(3): 355- Calidris).Die Neue Brem-Bucherei.A. Ziemsen 365. In Russian. verlag,Wittenberg Lutherstadt. Tomkovich,P.S. 1986. Geographicalvariability of the Uspenski,S.M., Boehme, R.L., Priklonski,S.G. & Vekhov, Dunlin in the Far East. Bull.of Moscow Soc. of V.N. 1962. Birds of North-Eastem Yakutia. Naturalists,Biol. Section91(6): 3-15. In Russian. Ornitologiya(Moscow) 4: 64-86. In Russian. Tomkovich, P.S. 1988. The birds of the southern coast of Vinikurov, A.A. 1971 Fauna of vertebrate animals in the the Buor-KhayaGull NorthernYakutia. In: V.E. areaof the Taimyrfield station(western Taimyr). Flint & P.S.Tomkovich (eds.), Archives ofZool. Mus. In: B.A.Tikhomirov (ed.). BiogeocenosesofTaimyr ofMoscow Univ. 26: 3-38. MoscowUniv. Press, andtheir productivity. Leningrad, Nauka: 212-230. Moscow. In Russian. In Russian. Tomkovich,ES. 1994. Sitefidelity and spatialstructure of Vinogradov,V.G., Krivenko, V.G. & Panfilov,A.D. 1991. populationsof RockSandpiper Calidris ptilocnemis Hotbedof tundraornithofauna in the upperpart of and Dunlin Calidrisalpina on ChukotskyPeninsula. the Pur Riverbasin. In: V.D. Ilychev(ed.), Materials Russ.J. Ornithol.3(1): 13-30. In Russianwith ofthe All-Union Ornithol. Conf. Part 1, pp. 52-53. Englishsummary. Nauka i tekhnika, Minsk. In Russian. Tomkovich, P.S.,Masterov, V.B. & Soloviev, M.Yu. 1991. Vronski, N.V. 1987. Materials to the avifauna of On the avifaunaof Mys Shmidta,the ChukchiSea. northwesternTaimyr. In: E.E.Syroechkovski (ed.), Ornitologiya(Moscow) 25: 175-176. In Russian. Faunaand ecology of birds and mammals of Central Tomkovich, P.S. & Sorokin, A.G. 1983. The bird fauna of Siberia,pp. 28-38. Nauka, Moscow.In Russian. Eastern Chukotka. In: V.E. Flint & ES. Tomkovich Vorobiev,K.A. 1963. BirdsofYakutia. USSR Acad. of Sci. (eds.),Archives of Zool. Mus. ofMoscow Univ. 21: 77- Press,Moscow. 336 p. In Russian. 159. Moscow Univ. Press, Moscow. In Russian. Voronin,A.Yu. & Korolyova,M.N. 1996. The centerof Tomkovich, P.S.& Vronski, N.V. 1988. The bird fauna of TaimyrPeninsula, area of SarytaturkuLake Diksonvicinity, northern Taimyr. In: V.E.Flint & (73ø40'N;96ø45'E). In: P.S.Tomkovich (ed.), P.S.Tomkovich (eds.). Archives of Zool. Mus. of Breedingconditions for wadersin Russiantundras Moscow Univ. 26: 39-77. Moscow, Moscow Univ. in 1994. WaderStudy Group Bull. 79: 76. Press. In Russian. Yakimenko, V.V. 1996. Lower reaches of Muksunikha and Tomkovich, P.S. & Vronski, N.V. 1994. Birds of lower Mungui Rivers(Lower Yenisei). In: P.S.Tomkovich reachesof the UboynayaRiver (North-Western (ed.),Breeding conditions for wadersin Russian Taimyr). In: E.V.Rogacheva (ed.), Arctic tundras of tundrasin 1994. WaderStudy Group Bull. 79: 76. Taimyrand Kara Sea islands: nature, fauna and Yurlov,A.K. 1982. Populationsand distributionin the conservationproblems 1: 161-206.Inst. Ecol.and summerbird communityin the lower reachesof Evol., Russian Acad. Sci., Moscow. In Russian. the PyasinaRiver (westTaimyr). In: V.I. Evsikov Trevor-Battye,A. 1895. &e-boundon Kolguev. Notes on the (ed.). Distributionand numbers of vertebrates in birdsrecorded, pp. 418-440.Archibald Constable Siberia. Novosibirsk, Nauka: 181-189. In Russian. and Company. Zarudny,N.A. 1910. Birdsof PskovProvince. Memoirs of Tugarinov,A.Ya. & Tolmachev,A.I. 1934. Materialsfor the RussianAcademy of Sci., Series 8, 25(2):1-181. In avifaunaof easternTaimyr. Annalsof thePolar Russian. Commissionof the USSR Acad. of Sci. 16: 5-47. In Zhukov,V.S. 1998. Seasonalchanges in distribution, Russian. abundance and numbers of waders in relation to Tulp,I., Bruinzeel,L., Jukema,J. & Stepanova,O. 1997. lemmingpopulation cycles in the westSiberian Breedingwaders at MedusaBay, Western Taimyr, in tundra. International Wader Studies 10: this volume. 1996. WIWO Report57, 90 pp. Zeist. Underhill,L.G., Prys-Jones,R.E, Syroechkovski,E.E. jnr., Groen,N.M., Karpov,V, van Roomen,M.W.J., Rybkin,A., Shekkerman,H., Spiekman,H. & Summers,R.W. 1993. Breedingof waders (Charadrii) and Brent GeeseBranta bernicla bernicla at PronchischevaLake, northeastern Taimyr, Russia,in a peakand decreasingletoming year. Ibis 135: 277-292.

164 Lappo& Tomkovich:Breeding distribution of DunlinCalidris alpina in Russia

APPENDIX Breeding recordsof Dunlin in Russia

Key No. Sitelocations on Figuresla & lb. # approximatelocation and co-ordinatesof sites.

Stat.Status of breedingrecords: 1 provedbreeding (nests, unfleged chicks, fledglings are found, female with egginside is collected); 2 probablebreeding (singing males, territorial pairs, characteristic behaviour near broods, and families with flying young); 3 possiblebreeding (indication of "breeding"without details given).

* pers.comm. ** unpublisheddata collected by members ofthe "Tundra Ecology-94" Russian-Swedish expedition: C. Mintonand D. Rogers (Australia);T. Pierstoa(The Netherlands); E. SyroechkovskiJr.,I. Chupin,V. Karpov,E. Lappoand V. Yakovlev (Russia); EE. J6nsson,A. Lindstr6m,N. Holmgren,N. Kjellenand E. Isakson(Sweden); and R. Gill (USA).

R. river BMNH = BritishMuseum of NaturalHistory L. lake ZMMU = ZoologicalMuseum of MoscowUniversity P.peninsula ZIR = ZoologicalInstitute of RussianAcademy of Sciencesin St.Petersburg I. I-s (islands)

No. Star. Regions Co-ordinates Sources I I PskovskoyeL. 58ø00'N27ø30'E Zarudny (1910) 2 I PskovskoyeL. 57ø55'N28ø20'E Zarudny (1910) 3 2 ChudskoyeL., Samolva 58ø18'N28ø00'E Leonovich* 4 I ChudskoyeL., Gdov 58ø45'N27ø47'E Malcheuski & Pukinski (1983) 5 2 Ilmen L., Novgorod 58ø30'N31ø15'E Mischenko & Sukhanova(1998) 6 1 FinlandGulf, KurgakskyP. 59ø45'N28ø10'E Buzun & Merauskas (1993) 7 1 Finland Gulf, Kernovo 59ø50'N 29ø00'E Malchevski& Pukinski(1983) 8# 3 FinlandGulf, BerezovyeIs. 60ø17'N28ø40'E Khrabry(1984)

:C..a. a!pinla

No. Stat. Regions Co-ordinates Sources 9 1 Bolshoi Ainov I. 69ø50'N 31ø35'E Tatarinkova(1983); Kokhanov & Skokova(1967) 10 1 Kola E, between Kharlovka & 68ø38"N 37ø25'E Semashko* VostochnayaLitsa Rivers 11 1 KolaP., middle Yokanga R. 67ø55'N37ø50'E Mikhailov 1993 12 2 KolaE, KachkovskyBay 67ø24'N41ø01'E TundraEcology** 13 1 Kola P., lower Ponoi R. area 67ø18'N 41ø10'E Filchagov*,Tundra Ecology** 14 1 Kola Pen., lower Ponoi R. 67ø00'N 41ø10'E Biankekiet al. (1982) 15 1 KolaP., Pyalitsa 66ø20'N40ø10'E Malyshevski(1962, 1963), ZIR 16 1 White Sea coast Koida 66ø25'N 42ø15'E Spangenberg& Leonovich(1960) 17 1 Kanin P., Chizha 67ø05'N 44ø20'E Spangenberg& Leonovich(1960) 18 1 Kanin P., Shoina 67ø55'N 44ø20'E Vinogradov& Avdanin*;Leonovich* 19 1 Kanin P., Torna 68ø02'N 44ø12'E Spangenberg& Leonovich(1960) 20 1 KaninP., Vostochnaya Kambalnitsa 68ø32'N45ø16'E TundraEcology,** 21# 1 KolguevI., ArtelnayaR. 69ø25'N49ø15'E Tolmachev(1928); Trevor(1895) 22 I KolguevI., PeschankaR. 69ø08'N50ø00'E TundraEcology**; Syroechkovski & Morozov* 23 2 KolguevI., Paarchik_haR. 69ø00'N49ø55'E TundraEcology**; Syroechkovski & Morozov* 24 3 MalozemelskayaTundra, Pesha R. 66ø55'N47ø40'E Romanov (1989) 25 1 Timanskayatundra, Indiga 67ø52'N49ø30'E Gladkov (1951a) 26 1 Timanskayatundra, Ikcha R. 67ø40'N49ø40'E Gladkov (1951a) 27 2 Timanskayatundra, near 68ø19'N50ø47'E Morozov* SengeiskyStrait 28 1 RusskyZavorot E 68ø48'N53ø07'E TundraEcology** 29 1 RusskyZavorot P., Omulevka R. 68ø52'N53ø29'E TundraEcology** 30 1 RusskyZavorot P., KhabuikaL. 68ø32'N53ø53'E Schadilov& Bouler (1996) 31 1 near Pechora delta, Lovetski I. 68ø20'N 53ø55'E Schadilov& Bouler(1996), Mineyev & Mineyev (1997)

165 International Wader Studies 10: 152-169

32 1 near Pechora delta, 68ø20'N53ø30'E Bianki & Krasnov (1987) KorovinskayaGulf 33 1 near Pechora delta, former Alekseevka 68ø00'N 54ø00'E Seebohm& HarvieBMMU; Seebohm (1985) 34 2 Varandey 68ø50'N 58ø00'E Kalyakin(1994) 35 1 KhaipudyrskayaGulf (south) 68ø22'N 59ø20'E Tikhonov* 36 1 MoreR- mouth 68ø20'N 59ø45'E Estafiev(1983, 1991) 37 1 NovayaZemlya, Pukhovy Bay 72o40'N 52ø45'E Gavrilov (1996a) 38 1 NovayaZemlya, Pankova Zemlya 73ø10'N 53ø20'E Kuzyakin,ZMMU nearGribovaya Gulf 39 1 NovayaZemlya, Matochkin $har 73ø15'N 54ø20'E Schaanning(1916) 40 1 VaigachI., Vaigachsettlement 70ø23'N 58ø52'E Morozov (1998) 41 3 VaigachI., LyamchinaGulf 69ø55'N 59ø15'E Romanov* 42 1 VaigachI. 70ø17'N 59ø05'E Morozov (1998) 43 1 VaigachI. 69ø50'N 60ø15'E Morozov (1998) 44 1 VaigachI., 69ø43'N 60ø05'E Morozov (1998) 45 1 YugorskyE, BelyNos 69ø36'N 60ø13'E Gavrilov (1996b) 46 1 YugorskyE, YugorskyShar 69ø50'N 60ø40'E Grichik* 47 1 YugorskyP., Chaika Cape 69ø30'N 60ø35'E Estafiev (1991) 48 1 YugorskyE, TonkyCape 69ø50'N 60ø45'E Grichik (1995);Morozov (1998) 49 1 YugorskyE, BolshayaOyu R. 69ø27'N 61ø20'E Estafiev (1991) 50 1 YugorskyE, Amderma 69ø45'N 61ø45'E Estafiev (1991) 51 1 YugorskyE, KarskayaGulf 69ø17'N 64ø35'E Estafiev (1991) 52 1 Vorgashor 67ø40'N 63ø50'E Morozov (1987) 53 1 Khalmer 67ø50'N 64ø50'E Morozov (1987) 54 1 BaidaratskayaGulf (south) 68ø10'N 68ø23'E Kalyakin(1986) 55 1 Western Yamal P., Ust 68ø55'N 69ø30'E Flint, ZMMU 56 1 WesternYamal P., Nyabyyakha R. 69ø38'N 67ø36'E TundraEcology** 57 1 Western Yamal P., Marre 69ø45'N 66ø50'E Danilov et al. (1984) 58 1 WesternYamal P., Mordyyakha R- 70ø14'N 66ø17'E TundraEcology** 59 1 WesternYamal E, MordyyakhaR- 70ø20'N 67ø41'E TundraEcology** 60 1 WesternYamal E, MordyyakhaR- 70ø25'N 68ø02'E TundraEcology** 61 1 WesternYamal E, Charasavey 71ø10'N 66ø45'E Danilov et al. (1984); Morozov* 62 1 WesternYamal P., Syadoryakha R- 71ø40'N 68ø15'E Danilov et al. (1984); Morozov* 634 2 BelyI. 73ø02'N 70ø10'E Sosin& Paskhalnyi(1995) 64 1 EasternYamal E, Tambey 71ø28'N 71ø45'E Sosin& Paskhalnyi(1995) 65 1 Eastern Yamal E, Yaibari 71ø15'N 71ø45'E Ryabitsev(1993) 66 1 EasternYamal E, YasaveiyakhaR- 70ø08'N 71ø15'E Danilov et al. (1984) 67 1 EasternYamal E, SeyakhaR- 70ø10'N 72ø35'E Danilov et al. (1984) 68 1 EasternYamal E, Khanovey 68ø33'N 72ø45'E Ryabitsev(1993) 69 1 EasternYamal E, KamermyCape 68ø30'N 73ø35'E Danilov et al. (1984) 70# 1 SouthernYamal P., Porsyakha 67ø40'N 71ø20'E Danilov et al. (1984) 71# 2 Southern Yamal P., 67ø20'N 70ø00'E Danilov et al. (1984) middle KhadytayakhaR. 72# 1 SouthernYamal P., Schuchya R- 67ø35'N 69ø15'E Kozlova(1962); Andreeva*; Kalyakin* 73 3 Oblowland,Poluy R- 65o55'N 68ø45'E Pokrovskaya* 74 3 ObYeniseylowland, Nadym R- 65ø00'N 74ø10'E Pokrovskaya* 75 3 Oblowland,Pyakupur R. 64ø40'N 77ø00'E Vinogradovet al. (1991) 76 3 Oblowland, Taz R. 65ø35'N 82ø25'E Pokrovskaya* 77 1 GydanP., Antipayuta 69ø05'N 76ø55'E Zhukov (1998) 78 2 GydanP., Tadibeyakha 70ø22'N 74ø10'E Zhukov (1998) 79 1 Gydan E, Yuribey 71ø00'N 77ø00'E Zhukov (1998) 80 1 GydanE, Gyda R. 70ø55'N 79ø15'E Blokhin, ZMMU 81 1 Gydan E, KhoseinL. 70ø55'N 80ø05'E Naumov (1931) 82 1 GydanP., Yeniseiskoye L. 71ø25'N 79ø45'E Chernichko et al. (1994) 83 1 GydanP., Mamonta E 71ø55'N 76ø20'E Zhukov (1998) 84 1 YeniseiGull SibiryakovI. 72ø45'N 79ø05'E SyroechkovskiJr*; Chylarecki & $ikora*

No. Stat. Regions Co-ordinates Sources 85 1 Kara Sea,Sverdrup I. 74ø35'N79ø20'E SyroechkovskiJr. & Lappo (1994) 86 2 Kara Sea,Arcticheskogo Instituta I. 75ø05'N82ø01'E TundraEcology94** 87 1 WesternTaimyr, lower UboinayaR- 73ø38'N82ø25'E Mork et al. (1994) 88 1 WesternTaimyr, Dikson 73ø30'N80ø30'E Tomkovich & Vronski (1988); ZMMU 89 1 WesternTaimyr, Meduza Bay 73ø20'N 80ø30'E $viridova*; Hertzler & Gunther*

166 Lappo& Tomkovich:Breeding distribution of Dunlin Calidrisalpina in Russia

90 3 WesternTaimyr, Rogozinka R. 72ø48'N80ø50'E Vronski (1987) 91 I WesternTaimyr, Yenisey Gulf, 72ø17'N81ø00'E Popham,BMNH Korsakovski I. 92 I WesternTaimyr, Golchikha R. 71ø45'N83ø30'E Popham,BMNH; Chernikov* 93 I WesternTaimyr, Pustoye 71ø10'N83øI5'E Popham,BMNH; Chernikov* 94 I WesternTaimyr, Mungui R. 70ø25'N83ø30'E Yakimenko (1996) 95 I WesternTaimyr, 69ø48'N85øI0'E Seebohm,BMNH; Seebohm(1985b) SukhayaDudinka R. 96 I PyasinaR. basin,Lidiya mouth 74ø07'N 86ø50'E H6tker (1995) 97 I WesternTaimyr, Chadyrymota R. 74ø00'N 87ø00'E Kokorev* 98 3 WesternTaimyr, Binyuda R. 73ø40'N 89ø15'E Kokorev & Lisenko (1989) 99 3 WesternTaimyr, Koreulakhbigai R. 73ø48'N 90ø50'E Kokorev & Lisenko (1989) 100 I WesternTaimyr, lower TareyaR. 73ø17'N 91ø10'E Vinokurov (1971);ZMMU 101 1 WesternTaimyr, Ust 73ø15'N 90ø34'E Yurlov (1982) 102 1 WesternTaimyr, middle PuraR. 72ø23'N 85ø25'E Kokorev* 103 1 WesternTaimyr, Purinskiye L. 71ø50'N 88ø40'E Kretchmar (1966) 104 1 WesternTaimyr, Agapa mouth 71ø25'N 89ø20'E Leonovich*; Kozhevnikov* 105 1 WesternTaimyr, middle DudyptaR. 71ø11'N 92ø13'E Lappo& SyroechkovskiJr.* 106 1 WesternTaimyr, Dudypta R., Kresty 70ø50'N 89ø55'E Kretchrnar (1966) 107# 3 WesternTaimyr, upper PyasinaR. 70ø40'N 89ø30'E Kozhevnikov* 108# 2 NorthernTaimyr, 74ø25'N 91ø50'E H6tker (1995) middle LenivayaR. 109 1 NorthernTaimyr, Nizhnyaya 75ø32'N99ø10'E Kokorev* TaimyraR. 110 1 NorthernTaimyr, Trautfetter mouth 75ø25'N99ø52'E Chupin* 111 1 EasternTaimyr, Ledyanaya R. 75ø28'N107ø25'E Kozhevnikov(1982); Rogacheva (1992) 112 1 EasternTaimyr, Neizvestnaya R. 76ø11'Nlllø23'E TundraEcology 94** 113 1 EasternTaimyr, Pronchischeva L. 75ø16'Nl12ø28'E Underhill et al. (1993) 114 1 EasternTaimyr, 73ø36'N106ø40'E Yakovlev,ZMMU; Yesou& Chupin* BolshayaBalakhnya R. 115 2 CentralTaimyr, Rysyukov Cape 74ø22'N 100ø05'E HOtker (1995) 116 1 CentralTaimyr, Ozhidaniya Bay 74ø40'N 101ø00'E Sdobnikov (1959) 117 3 CentralTaimyr, Bikada R. 74ø45'N 106ø00'E Rogacheva(1992) 118# 1 CentralTaimyr, YamuTarida R. 74ø25'N 102ø50'E Tugarinov& Tolmachev(1934) 119g 3 CentralTaimyr, Luktakh R. 73ø16'N 93ø44'E Pavlov et al. (1983) 120 3 CentralTaimyr, Logata R. 73ø12'N 95ø55'E Chupin* 121 1 CentralTaimyr, lower LogataR. 73ø07'N 96ø10'E Kokorev* 122 2 CentralTaimyr, 73ø50'N 96ø55'E Gavrilov* BolshayaBootankaga R. 123 2 CentralTaimyr, Sarytaturku L. 73ø40'N 96ø45'E Voronin & Koroleva (1996) 124 1 CentralTaimyr, Malaya LogataR. 73ø23'N 98ø24'E Gavrilov (1989); H/Stker (1995) 125# 3 CentralTaimyr, Novaya R. 72ø38'N 100ø40'E Volkov* 126 1 CentralTaimyr, Tonskoye L. 72ø16'N 98ø50'E Karpovet al. in press. 127# 1 CentralTaimyr, Boganida 1• 71ø20'N 97ø00'E Middendorff, ZIR

No. Stat. Regions Co-ordinates Sources 128 1 KhatangaR,, Novorybnoye 72ø51'N106ø02'E Chupin*;Soloviev (1995), Soloviev et al. (1996) 129 2 Anabar tundra, Paksa P. 73ø40'N 113ø00'E Uspenski(1965) 130 3 Anabar tundra, Uele R. 73ø20'N 114ø10'E Uspe'nski(1965) 131 3 Anabartundra, Yuryung-Khaya 72ø50'N113ø12'E Gladkov& Zaletayev(1965) 132 2 Anabartundra, lower PeschanayaR. 73ø40'N115ø30'E Gladkov& Zaletayev(1965) 133 2 Anabartundra, Oyulakh-Yurgakh R. 73ø27'N117ø00'E Lappo* 134 1 Anabartundra, Chaidakh-Yurgakh R. 73ø18'N116ø55'E TundraEcology** 135 2 Anabartundra, Terpyai-Tumus P. 73ø30'N118ø35'E TundraEcology** 136# 2 Anabartundra, Kuogastakh R. 72ø25'N124ø20'E Pozdnyakovet al. (1996) 137# 2 LenaDelta,Khaas-Khaata-Aryta I. 73ø15'N125ø20'E Labutin et al. (1985) 138# 1 LenaDelta, Yuggyus-D'ie L. 72ø50'N126ø10'E Labutin et al. (1985);Blokhin & Blokhin (1986) 139g 1 LenaDelta, upper Bolshaya 72ø25'N126ø20'E Blokhin* Tumatskaya 140 1 Lena Delta, Nordenskiold station 72ø13'N 127ø55'E Boere* 141# 1 LenaDelta, Bulkurskaya 72ø10'N126ø03'E Posdnyakovet al. (1996) 142 2 Tiksi Bay 71ø35'N128ø50'E Gladkov(1958); Kapitonov (1962) 143 1 Sytygan-TalaBay, 70ø42'N131ø20'E Tomkovich (1988)

167 International Wader Studies 10: 152-169

144 1 Yana delta, Ilin Shar 71ø20'N 134ø50'E Syroechkovski;Roschevski* 145 1 Yanadelta, lower Maly Samandon 71ø30'N135ø20'E Kistchinski,ZMMU; Kistchinski(1988) 146 2 Yanadelta, Nizneyansk 7!ø30'N136ø10'E Syroechkovski 147 1 Yanadelta, Yukagir 71ø50'N139ø50'E Syroechkovski;ZMMU 148# 2 Khromalowland 72ø05'N 147ø55'E Uspenskiet al. (1962) 149 1 Khromalowland,Berelyakh 70ø50'N146ø55'E Tomkovich*; Leonovich*; Flint, ZMMU 150# 1 Khromalowland,Batyntai L. 70ø57'N147ø30'E Flint, ZMMU 151 1 Khromalowland,Gusinaya Bay 71ø35'N149ø14'E TundraEcology** 152# 1 Khromalowland,Indigirka delta, 71ø05'N150ø40'E Flint, ZMMU Keremesit 153# 1 Khromalowland,Indigirka delta, 71ø00'N151ø20'E Kistchinski(1988), ZMMU Kolesovo 154• 1 Kolymalowland, lower Chukochya 70ø15'N159ø25'E de Roos* 155# 3 Kolymalowland, lower 69ø45'N159ø37'E Kretchmar et al. (1991) KonkovayaR. 156• 1 Kolymalowland, Vankhotveem 69ø00'N158ø30'E Kretchmaret al. (1991);Kondratyev (1982) 157# 1 Kolymalowland, Khalerchinskaya 69ø00'N160ø00'E Kretchrnaret al. (1991);Kondratyev (1982) tundra

No. Stat. Regions Co-ordinates Sources 158• 1 WrangelI. 71ø15'N179ø45W Portenko(1972); Dorogoi (1982); Stishov et al. (1991),ZIR 159 1 WrangelI., MamontovayaR. 70o54'N 179o33'E TundraEcology* 160# 1 ChaunGulf, Ayon I. 69o55'N 168o20'E Stishov (1990) 161 1 ChaunG .ulf, Ayon I. 69ø48'N 168o38'E TundraEcology* 162 1 Ust 68o45'N 170o40'E Ostapenko(1973); Kondratyev (1982); Kretchmar et al. (1991) 163 3 North Chukotka,Nolde Bay 69ø45'N 173ø20'E Dorogoi* 164 1 North Chukotka,Cape Aachim 69ø50'N 173o35'E Stishov (1993) 165 1 North Chukotka, ShalaurovaIzba 69o50'N 174o30'E Stishov (1993) 166 1 North Chukotka,Cape Yakan 69o35'N 177ø30'E Stishov (1993) 167 1 North Chukotka,Shmidta Cape 68ø55'N 179o25'W Tomkovichet al. 1991;Dorogoi,* 168 1 North Chukotka, lower 68ø43'N 178ø55'W Stishov (1993) EkvyvatapR. 169 3 North Chukotka, middle 68ø18'N 179ø50'W Stishov (1993) EkvyvatapR. 170 1 NorthChukotka, Ukouge lagoon 68o08'N 177o08'W Portenko (1972);Kitschinski (1988) 171 1 North Chukotka, 67o55'N 177o35'W Portenko (1972);Kitschinski (1988) lowerAmguema R. 172 1 North Chukotka, middle 67o40'N 178o45'W Dorogoi(1993a) AmguemaR. 173 1 North Chukotka, middle 67ø02'N 178o55'W Portenko (1972);ZIR AmguemaR. 174 1 North Chukotka,Belyaka spit 67o05'N 174o42'W Leonovich*;Tomkovich*; Kondratyev (1982) 175 2 North Chukotka,Rekokaurer Cape 66o20'N 173o35'W Flint* 176 1 North Chukotka,Cape Serdtse 66o55'N 171o45'W Leonovich*;Portenko (1972) 177 3 East Chukotka, Uteveem R. 66o15'N 170o45'W Portenko (1972) 178 1 East Chukotka, Uelen 66o10'N 169o50'W Portenko (1972);Tomkovich & Sorokin (1983); ZMMU 179 1 East Chukotka, Koolen L. 65ø55'N171ø10'W Dorogoi* 180 1 East Chukotka, Puotenveem mouth 65ø50'N170ø40'W Tomkovich& Sorokin(1983) 181 1 EastChukotka, Lavrentiya 65ø35'N171ø00'W Portenko(1972); Kiryuschenko (1973); Tomkovich& Sorokin(1983) 182 2 East Chukotka, Arakamchechen I. 64ø45'N 172o25'W Dorogoi* 183 1 EastChukotka, Chaplino 64o25'N 172o15'W Kuzyakin*;Tomkovich & Sorokin(1983) 184 1 EastChukotka, Provideniya Bay 64ø18'N 173o35'W Portenko (1972) 185 2 EastChukotka, Kurupkan delta 64ø40'N 174ø15'W Konyukhov* 186 2 KrestaBay, Notapelrnen 65ø35'N 178ø20'W Portenko (1972) 187 2 KrestaBay, Egvekinot 66ø20'N 179ø10'W Dorogoi*;Tomkovich* 188 1 KrestaBay, Uelkal 65ø30'N 179ø20'W Portenko (1972);Yakobi*; Leonovich* 189 1 upperKanchalan valley 66ø10'N179o10'E Kistchinskiet al. (1983);Kistchinski (1988); Flint, ZMMU

168 Lappo& Tomkovich:Breeding distribution of Dunlin Calidrisalpina in Russia

190 1 middle Kanchalanvalley 65ø35'N 178ø20'E Kistchinskiet al. (1983);Kistchinski (1988); Flint, ZMMU 191 1 lower Kanchalanvalley 65ø15'N 177ø25'E Kistchinskiet al. (1983);Kistchinski (1988); Flint, ZMMU 192 1 Anadyr estuary,Shakhtersky 64o45'N 177o35'E Dorogoi*;Portenko (1939); Kiryuschenko (1973) 193 1 Anadyr estuary,Anadyr 64ø45'N 177ø30'E Leonovich*;Portenko (1939);Kolonin (1980) 194 1 Anadyr estuary,Avtatkuul R. 64ø05'N 178ø20'E Kondratyev* 195 1 Anadyr estuary,Tumanskaya mouth 64ø00'N 178ø35'E Portenko(1939) 196 2 middle Anadyr R., between 65o10'N 171o30'E Kretchrnaret al. (1991) Anadyr R. & Main R. 197 middleAnadyr R., KrestovskayaR. 65ø20'N 171ø40'E Kretchmar et al. (1991) 198 ElgygytgynL. 67ø30'N 172o07'E Dorogoi(1993b)

a, 'kistchinSki

Stat. Regions Co-ordinates Sources 199 2 KoryakHighland, Khatyrka lagoon 62ø08'N 175ø28'E Kistchinski(1980) 200 2 KoryakHighland, Apuka mouth 60o25'N 169o45'E Kistchinski (1980, 1988) 201 I KoryakHighland, Tilichiki 60ø27'N 166ø03'E Kistchinski(1980) 202 I KoryakHighland, Geka Bay 60ø00'N 165ø10'E Kistchinski(1980); Firsova & Levada (1982) 203 1 Kamchatka P., Ossora 59ø55'N 163ø05'E Gerasimov* 204 1 KaraginskiI., Markelovskaya 58ø50'N 164ø15'E Gerasimov& Vyatkin(1973) 205 2 East Kamchatka, Ust 56o14'N 162o12'E Ryabushinski,ZIR 206# 1 EastKamchatka, Bolshaya 55ø04'N 161ø47'E Lobkov (1986) Chazhma R. 207 1 EastKamchatka, Kronotskaya R. 54ø30'N 160ø40'E Lobkov (1986) 208 I East Kamchatka, Uzon volcano 54ø30'N 160ø25'E Lobkov (1986) 209 1 EastKamchatka, Semyachiksky 54ø10'N 160ø00'E Lobkov (1986) lagoon 210 2 East Kamchatka, Avacha mouth 53ø05'N 158ø30'E Gerasimov (1968) 211 2 SouthKamchatka,Kambalnaya IL 51ø10'N 156ø50'E Gerasimov, ZMMU 212 1 Shumshu I. 50ø40'N 156ø20'E Podkovyrkin(1956) 213 1 Paramushir I. 50ø30'N 156ø00'E Golovushkin* 214 1 West Kamchatka, Plotnikova R. 52ø55'N 156ø35'E Gluschenko(1984) 215 3 WestKamchatka, Bryumka R. 54ø30'N 155ø50'E Lobkov (1986) 216 1 West Kamchatka, 56ø30'N 156ø30'E Gerasimovet al. (1992);ZMMU MoroshechnayaR. 217 3 West Kamchatka, 57ø10'N156ø47'E Lobkov (1986) lower KhairyuzovaR. 218 3 WestKamchatka, lower Tigil R. 58ø10'N 158ø20'E Lobkov (1986) 219 3 WestKamchatka, Parapolski 61ø20'N 164ø40'E Lobkov (1986) depression,Talovskoe L. 220 1 ShelikhovaBay, Gizhiga 62ø00'N 160ø30'E Allen (1905) 221 1 ShelikhovaBa]6 Yamsk 59ø35'N 154ø05'E Kistchinski(1968); Kondratyev* 222 1 ShelikhovaBa]6 Babushkina Bay 59ø12'N 153ø45'E Kistchinski(1968); Kondratyev* 223 1 Magadanarea, Ola delta 59ø30'N 151o05'E Dorogoi* 224 1 Magadanarea, Arman 59ø40'N 150ø10'E Dorogoi* 225 1 Magadanarea, Ojra 59o45'N 149o50'E Leonovich* i C;a...a:ites

No. Stat. Regions Co-ordinates Sources 226 1 North Sakhalin,Piltun Bay 53o10'N 143o15'E Leonovich*;Nechaev & Tomkovich(1987); Nechaev (1991) 227 2 North Sakhalin,Chaivo Bay (north) 52ø30'N 143ø15'E Blokhin* 1 North Sakhalin,Chaivo Bay (south) 52ø20'N 143ø10'E Nechaev (1979, 1991);Nechaev & Tomkovich(1987) 229 2 North Sakhalin,Dagi Bay 52ø15'N143ø07'E Nechaev& Tomkovich(1987); Nechaev (1991) 230 2 North Sakhalin,Nyisky Bay 52ø07'N143ø05'E Nechaev& Tomkovich(1987); Nechaev (1991) 231 2 North Sakhalin,Nabilsky Bay 51o35'N143ø23'E Nechaev& Tomkovich(1987); Nechaev (1991)

169