Melnikov: Populationand rangefluctuations of AsianDowitcher Ltmnodromus semtpalmatus in the centralAsian arid zone

Population and range fluctuations of Asian Dowitcher Limnodromussemipalmatus in the central Asian arid zone Yu.I. Melnikov

Melnikov,Y.I. 1998.Population and range fluctuations of AsianDowitcher Limnodromus semipalmatusin the centralAsian arid zone. InternationalWader Studies 10: 351-357.

Themain breeding range of theAsian Dowitcher Limnodromus semipalmatus lies within the limitsof the arid zonein south-westernSiberia, Mongolia and China. Furthernorth, up to 56-57øN,it is patchilydistributed on isolatedareas of relictsteppe. At the end of the 19thcentury several hundredpairs were known to bebreeding at thenorthern limits of thespecies' breeding range. Theirpopulation during this period was probably stable. Since the mid-1970s the populationhas increasedto 6,000birds. Thisis mainlydue to extremedroughts in the southernparts of the breedingrange and an expansionof at least1,000 km in the extentof thenorthern boundary. This increaseoccured very rapidly overa periodof abouttwo years. The subsequentpopulation decreasewas slow, lasting more than ten years. Currently(1990-1991) the populationis similarto that at the beginningof the century.A slowdecrease in numbersin the northernpart of the breedingrange after the droughtended in thesouth indicates that the speciesis bothhighly site- faithfuland very mobile. The species' expansion is characterisedby markedirruptions during extremedroughts, their scale depending on the sizeof the areassuffering from drought.This behaviouris a characteristicadaptation to existencein arid regions.

Yu.I. Melnikov, East-SiberianSection ASRIF, SedovStr., 41, Irkutsk, 664003, Russia.

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Introduction zoneonly extendsinto Russiaat the border,the Publications from recent decades have shown that characteristicsof thisphenomenon could only be studied for the northern boundaries of the waders the spatialstructure of the rangesof waderssuch as AvocetRecurvirostra avosetta, Black-winged Stilt breedingin thesearid areas.The groupof waders Himantopushimantopus, Marsh Sandpiper Tringa that usethe narrow habitatspectrum in the arid stagnatitisand AsianDowitcher Limnodromus regionare of specialinterest, as most of them are semipalmatusbreeding near the water bodies of the includedin the RussianRed Data Book'sregional arid zoneof CentralAsia arevery variable listsof endangeredspecies and thusrequire special (Melnikov 1986a,1986b; Osipova 1986). As the arid measuresfor their protection.An important elementof suchwork is the analysisof those

351 International Wader Studies 10: 351-357

80OE 100OE 120øE 140øE 160øE Study

Lake Baikal

Figure1. AsianDowitcher breeding range: 1 - WestSiberian concentration; 2 - the LargeLakes depression (Mongolia); 3 - Selengariver basinconcentration (Mongolia); 4 - Russian-Mongolian-Chineseconcentration; 5 - Cicikarconcentration (China); ß - recordsof AsianDowitchers during invasionsto northernrange limits.

populationprocesses for speciesof which only we collecteddata on the changesin numbersof thrs limited data exist. The Asian Dowitcher is used as speciesin south-easternSiberia, as well as detailed an exampleof thisanalysis of fluctuationsin informationon the ecologyof Asian Dowitcherat numbersand changesin breedingrange the Selengariver delta (1973-1980)and the Irkut boundaries. river mouth(1983-1987). The mostprobable breedingareas were also surveyed (Melnikov 1985, The AsianDowitcher is a rare,poorly studied 1986a,1991), and all publisheddata were analysed. waderspecies, included in theIUCN RedData Book Totalcounts were made at the Selengadelta once and in the Red Data Books of several countries everytwo or threeyears using specially designed within the CIS.-The totalpopulation was estimated methods(Melnikov 1986c). Reproductive success not longago at 1,000birds. Surveysat the was estimatedaccording to Mayfield (1975), winteringgrounds in south-easternAsia made it modifiedby Payevsky(1985). The nest-historiesof possibleto updatethis figure to 5,000birds in 1984. a total of 625 nests were recorded: 531 nests at the After a furtherinventory of the winteringgrounds Selengadelta and 94 at the Irkut river mouth. in 1989,the populationwas estimatedat 15,000- 20,000birds (Howes& Parish1989). Surveysmade Results and Discussion within its breedingrange in Russiain yearswhen largenumbers of birdswere present estimated less The breedingrange occupies a narrowbelt in the than half this number (lessthan 6,000birds; forest-steppes,steppes and semi-desertsfrom Melnikov 1986a),whilst the datafrom otherparts of south-westernSiberia across the Baikalregions, the breedingrange are insufficient for evena Mongolia,and northernChina to the shoresof the preliminaryestimate of populationsize. PrimoryeTerritory. The main part of the breeding rangeis situatedwithin the arid zone,where birds Methods inhabitextremely variable wetland ecosystems. The characteristicdistribution is due to the fact that they Informationpresented in thispaper is basedon 20 chooseto breedin rather.flat, swampy meadows yearsof studiesfrom 1972-1991.During thisperiod with low grasscover, which often have numerous

352 Melnikov. Populationand rangefluctuations of AsianDowitcher L•mnodromus semipalmatus in the centralAsian arid zone shallowtemporary water bodies, as well as the Onelarge concentration where breeding occurs shallowwaters and mudflats of temporarylakes in regularlyis situatedin the Nonni and the the steppeareas. Most of the recordsfrom the Sunkhuaztsyanriver basins and probably at the northernboundary of thebreeding range have been Sungaririver basin, from where birds spread to in areasof steppeor similarhabitats. Judging from KhankaLake and the Zeya water reservoir.This publisheddata (Pevtsov1883; Kozlova 1930) the breedingconcentration is probablyconnected with sameis true alsofor the southernboundary of the the previousone as the ratherlow Kinganmountain species'breeding range. rangeprobably does not act as a barrier. Available data (Taczanowski1873; Velizhanin & The most characteristic feature of such arid areas is Velizhanin1929; Zalessky 1934) suggest that several the sharpcyclic fluctuations of waterlevel in lakes hundred birds bred at the northern limits of the and otherwater bodies (Shnitnikov 1957;Maksimov species'range (in Russia)in the late 19th- early 20th 1984;Melnikov 1986b;Krivenko 1991). Suchcyclic centuries.The specieswas consideredto be changesin the watertableaffect breeding conditions extremelyrare and probablygoing extinct. for shorebirdsand waterfowl. Consequently the Sometimes,according to theseobservations, the existenceof locationswhere breeding is always numbers of Asian Dowitcher would fluctuate, possible,only occursin vastswampy areas which althoughthe data for thatperiod are insufficient haveoptimal habitat diversity (Melnikov, 1986b). evenfor a generalanalysis. The appearanceof the The distributionof suchareas is determinedby speciesin ratherlarge numbers there during the landscapeconditions and, as a result,they are 1970swas rather unexpected (Melnikov 1979; situatedwell apartfrom eachother. In betweenare Gluschenko& Shibnev1979; Yurlov 1981). rathersmall swampy areas which cansupport only Accordingto maximumestimates made in different a few breedingAsian Dowitchers, often only in very areasin thenorth of thebreeding range, numbers of favourableyears (Figure 1). Asian Dowitchers at that time reached 6,000 birds (Melnikov 1986a). Sincethen the number of One of thesebreeding concentrations is situatedin breedingbirds has decreased and currently the south-west Siberian lowland. This area is well fluctuates from 700 to 1,000 - 2,000 individuals. The isolatedfrom the easternparts of the species' largestnumber, about 4,500 birds, was recordedat breedingrange, a factreflected both in the migration the SelengaDelta (Melnikov1979, 1985). routesof thesebirds and the positionof their winteringgrounds in north-eastAfrica and, The changesin numbersand in the structureof the probably,the Near East(Liedel 1982; Melnikov AsianDowitcher's breeding range are rather 1990).Although we do not rejectthe possibilitythat complicatedand aredetermined by geographicand birdsfrom Mongolia can penetrate to Western climaticconditions. As the latterchange cyclically, Siberia,especially in big expansionyears, we it is possiblethat theyinfluence breeding success. considerthis breeding population to be separateand In thissituation we canassume that thebiggest well-isolatedfrom the main part of thebreeding differencesin breedingsuccess would be recognised rangeand restrictedto Russia.The relativelysmall at thebeginning and end of a climatichydrological population,which comprises less than 1.0%of the cycle. Our data showsthe tendencytowards world populationof AsianDowitchers, means that increasingbreeding success during a hydrological specialconservation measures are extremely cycleat the Selengariver delta,and a simultaneous importantin Russia(Melnikov 1990). decreaseat theIrkut river mouth(Figure 2). Unfortunately,the regressionswere not significant Two largebreeding concentrations are situated in dueto a highdegree of variationduring the northernMongolia: at the LargeLakes depressions, intermediateyears of the cycle.Additional fromwhere birds penetrate to TuvaRepublic and estimatesof thedistribution of breedingsuccess theMinusinsk depression, and at the Selengariver usingthe iterationcriterion (Zaks 1976)did not basin.The last one is themost significant breeding reveala temporaltrend and provedthe random concentration,from which Dowitchers spread in distribution of the extreme values, which were unfavourableyears to areasclose to the southern recordedthree times at the SelengaDelta and once and south-westernBaikal Lake areas(Melnikov at the Irkut river mouth(Figure 2). Thus,we 1991).At HubsugulLake the birds appear believethat breeding success is not dearlyrelated to sporadically,probably penetrating from the Large fluctuationsin the hydrologicalregime. Lakesdepression or fromthe Selengariver basin. Anotherbreeding concentration is locatedat the Numbers of Asian Dowitcher are limited over the bordersof north-easternMongolia, southern Russia wholebreeding range by similarfactors, the most (Dauria)and north-westernChina (Barga), and importantof whichare high rates of predation, includesthe Dalai-Nor,Byi-Nor and the Toreiskiye fluctuationsin the watersupply, nest-trampling by Lakesand the basins of dischargingrivers, as well cattleand othertypes of human influence asthe middlereaches of the Argunriver. The (Gluschenko& Shibnev 1979;Yurlov 1981;Melnikov groupsof AsianDowitchers which sometimes 1985,1991). Althoughthe intensityof thesefactors appearat the EravnaLakes, in the Barguzinand the differsbetween years, the average breeding success VerkhnyayaAngara river basinsprobably originate remainsrather stable (Figure 2). Thisappears to be from thisMongolian-Russian-Chinese breeding a resultof compensatoryadaptations to the area. changingenvironment with largedifferences in

353 International Wader Studies 10: 351-357

a) Selenga Delta hatchingsuccess % 7O theSelenga Delta was probably connected with comparativelyhigh numbers during immigration. 60- Theslow decline in numbersobserved during the

50- nexthydrological cycle indicates that two opposite trendsexist within the population: high mobility but 40- alsosome site fidelity. In spiteof the restorationof

30- breeding suitablebreeding conditions in themain breeding Success area,site fidelity was possibly the main reason for 20- ...... •..•averagethe slow decline in the numbers of Asian

10- ß Dowitchersat thetemporary, northern, breeding grounds. 0 I I I I I I 1972 1973 1974 1975 1976 1977 1978 1979 1980 1981 year Anotherpeculiarity of the AsianDowitcher is the b) Irkutriver mouth hatching success % formationof largelocal breeding concentrations, 50 whichcan include up to 50%of the totalpopulation. Such concentrations were recorded at the Airik-Nor 40- .. average breeding Lake(Pevtsov 1883), the Malyi ChanLake in 1974 success (Yurlov1981), at the SelengaDelta in 1977-1978 30- (Melnikov1979), at the Khanka Lake in 1975 (Gluschenko& Shibnev 1979), and at theToreiskiye 20- Lakesin 1971(Shkatulova 1973) and in 1985 (Osipova1986). 10- Thebreeding success of AsianDowitchers even in

I such short-term concentrations can influence the 1985 year sizeof the populationover a fairly longperiod. Comparativelylow averagereproductive success Figure2. Variationin AsianDowitcher breeding success a) (Figure2), aswell assome other characteristics of in theSelenga Delta and b) in theIrkut rivermouth during thespecies' ecology, suggest that thepopulation a completehydrological cycle (from maximum to remainsstable or increasesin periodsto sharp m'mimumwater supply). increasesin breedingsuccess in singleyears (Figure breedingsuccess occuring only in yearswith 2). As thespecies has a ratherlong life expectancy extremeconditions (Melnikov 1982). Statistically andtherefore a complicatedage structure within the significantdifferences in thebreeding success of population,high productivity seasons maintain AsianDowitchers (p<0.01) found at differentstudy ratherlarge and stablenumbers over a longperiod areasindicate geographic variations in population 10000_a)number ofbirds at Selenga Delta -- dynamics. -- --

-- A sharpincrease in the numbersof AsianDowitcher couldnot be theresult of changesin therelationship

betweenlocal productivity and mortality rates. The 1000_- breedingsuccess of birdsin theyears before peak numbersremained at the usualaverage level (Figures2 & 3). Althoughthe population peak of 1977coincided with veryhigh breeding success, no increasein numberswas recorded in the following years(Figure 3). In addition,there was no 7219'74 1976 1978 19801982 1984 1986 1988 1990 19•92 correlationbetween numbers and reproductive year success. The latter varied while numbers remained similar(Figure 4). Thereis no doubtthat 10o_b) number ofbirds at Irkut river mouth _ fluctuations in numbers at the northern limit of the species'range can be explainedonly by emigration andimmigration processes, following from sharp environmentalchanges in themain part of the breedingrange. lO_-

-- Our observationsat theSelenga river deltaand the -- Irkut rivermouth demonstrate that the change in numberswas similar (Figure 3) duringa mass -- appearanceof AsianDowitchers both in the casesof large-scaleimmigration and duringlocal 198019'81 19 2 19831984 1985 1986 1987 19 8 1989 resettlementof birds. A sharpincrease in numbers year wasobserved over a periodof oneto two years Figure3. Changesin numbersof AsianDowitchers a) in followedby a declineto theoriginal level. The the SelengaDelta andb) in the Irkut river mouthin south- prolongeddecline in numbersto the lower level at easternSiberia. (Note log scale).

354 Melnikov:Population and range fluctuahons of AsianDow•tcher Limnodromus semtpalmatus tn the central Asian arid zone

the neighbouringparts of Mongoliain 1977. As a 100. result, in 1975 Asian Dowitcher was recorded in largenumbers at KhankaLake (Gluschenko& Shibnev1979), and duringthe most severe droughts an influxwas recorded at the Selengariver delta (Melnikov 1979;1986a). So,the increasein numbers of Asian Dowitcher which was observed in Russia is connectedwith the largeimmigration of birdsfrom the main breedinggrounds which were suffering 10- from severedroughts.

The connectionbetween changes in thenumbers of ß Permamentstudy area of thisspecies at the boundariesof its breedingrange 100km 2 with the distributionand intensityof droughtsat the 0 Extrapolatedfor the whole mainbreeding grounds is emphasizedby the mass SelengaRiver Delta appearenceof AsianDowitchers at the north of the breedingrange at the sametime asthe droughts occurin the south(Melnikov 1986a). Although the 100 1000 5000 winteringgrounds of AsianDowitchers from Figure4. Relationshipbetween the breeding success and differentbreeding concentrations are at leastpartly numbersof AsianDowitchers at the SelengaDelta. Note shared(Verheugt et al. 1990),the asynchronous log scales. populationfluctuations suggest that these of time. In otheryears reproduction only partly concentrationsare independentfrom each other. makesup for annualadult mortality and, as a result, the populationdeclines until thenext favourable An increasein rainfallin Russiausually coincides breedingseason. Obviously, when breeding success with droughtsin neighbouringsouthern areas is extremelylow, it cancause a significantdecline in leadingto the emigrationof birdsfrom the main the population. breedinggrounds. Although the optimalconditions for breedingAsian Dowitchers are found on early Our studiessuggest that favourable breeding stagesof ecologicalsuccessions (Melnikov 1985; conditionsoccur once every eight to tenyears 1991),its massappearence on thenorthern breeding (Figure2). Thisis comparableto the averagelife of areasdoes not alwayscoincide with theformation of onegeneration, calculated by regression(Payevsky optimalbreeding habitats. Thus we canconclude 1985),and alsoclose to thelength of one11-year that emigrationof AsianDowitchers is determined hydrologicalcycle. onlyby environmentalconditions at the main breedinggrounds. The latteris provedalso by the Changesin numbersat thelimits of the breeding factthat, even in the extremeyears, no morethan rangecan be explainedby cyclicchanges in climate. half of the totalpopulation appears in thenorthern Thus,the 20th century is characterisedby a general parts,while the restevidently stay at themain warmingof climatein thenorthern hemisphere breedinggrounds, where suitable conditions remain (Budyko1980), accompanied by increasingly locally. frequentdroughts, sometimes extreme ones. These droughtsare observed asynchronously in different Sinceit hasbeen shown that breedingsuccess in areas,passing gradually to the neighbouringregions large,local concentrations of AsianDowitchers (Pasechniuket al. 1977;Rauner 1979;Koshelenko formedduring drought periods influences 1983).For example, the severe drought of 1968-1973 populationlevel for a ratherlong period. We can started in Africa and then moved eastwards, suppose,judging by the resultsof winter counts coveringthe large areas of India, Nepal, Burma,Sri (Howes& Parish1989) that the mostrecent years Lankaand southern China. In 1972it passedto have been rather favourable for this wader. It is morenorthern areas and spread over Europe and quiteprobable though, that high population CentralAsia (Koshelenko1983). estimatesfor Asian Dowitcheron the wintering groundswere made in theyears of themost In 1973-1974the period of ratherhigh humidityin successfulbreeding after the immigrationof birdsto south-westernSiberia changed to a periodof northernbreeding areas. At the sametime we droughts,when large areas of thelake depressions consider that the coincidence of unfavourable within the rangeof AsianDowitcher dried out conditionsfor the speciesat differentbreeding (Maksimov1984; Krivenko 1991). It wasprobably concentrationsover several years can cause the main reason for the concentration of Asian catastrophicchanges in the stateof the Asian Dowitchersfrom this part of thebreeding range on Dowitcherpopulation. the easternshore of theMalyi ChanLake, the Chulymand the Kargat river valleys, where the The absenceof massemigration of Asian conditionsfor breeding had become optimal (Yurlov Dowitchersin theperiods before and after the 1970s 1981). Since1975, severe and prolongeddroughts is explainedby the absenceof severeextensive haveoccurred in China(Cheng 1978; Ofori-Sarpong droughtscovering the mainbreeding grounds of the 1980),and were at theirworst when they spread to

355 h•ternational Wader Studies 10. 351-357

species.It is known that suchdroughts last for two SteppenschlammlaufersLimnodromus semipalmatus or threeyears and recurno moreoften than oncein (Blyth).Mitt. Zool.Mus., Berlin. Bd 58. Ann. Orn. a century(Koshelenko 1983). We thereforesuppose 6: (S)147-162. that the observed fluctuations in the number of Maksimov,A.A. 1984.Long-term jquctuations ofanimal birdsalso depends on suchlong-term cyclic climatic numbers,their reasons and forecasting. Nauka, changes.Therefore we do not expectanother mass Novosibirsk.249 pp. In Russian. appearenceof the speciesat thenorthern breeding Mayfield,H.E 1975.Suggestions for calculatingnest areas in Russia in the near future. success.Willson Bull. 87(4): 456-466. Melnikov,Yu.I. 1979.Numbers and ecology of Asian Dowitcherat theSelenga river Delta. In: Yu.V. Conclusions Labutin(ed.), Migration and ecology ofbirds of Siberia Long-termobservations on AsianDowitcher during pp. 160-161.Yakutsk Section of SiberianBranch of a 20-yearstudy and analysisof the literature USSR Acad. Sci.,Yakutsk. In Russian. demonstratethat the latestsharp increase in Melnikov,Yu.I. 1982.Certain adaptations in coastalbirds. numbersof this speciesin Siberiawas connected TheSoviet Journal of Ecology 13(2): 134-139. with the displacementof the optimalpart of its Melnikov,Yu.I. 1985.On the ecology of Asian Dowitcher breedingrange to the northernareas due to at theSelenga Delta. Bull.of Moscow Soc. of emigrationof birdsfrom concentrations on themain Naturalists,Biology Branch 88 (5):21-28. breedinggrounds. The scaleof suchemigrations are In Russian. determinedby the sizeof thearea affected and the Melnikov,Yu.I. 1986a.Number dynamics, peculiarities of durationof the drought. Sucha dynamic distributionand protection of AsianDowitcher in distributionis consideredto be themain adaptation theUSSR. In: V.E. Sokolov, E.E. Syroechkovsky & of AsianDowitcher to dependanceon wetlandsin V.A.Kuzyakin (eds.), All-Union Meeting on the an arid zoneenvironment. This adaptation is problemsofcadastre and censuses ofanimals, part 2, realizedeither through local redistribution of birds pp. 341-342.Rosagropromizdat, Moscow. withinbreeding concentrations or throughlarge- In Russian. scaleemigration, caused by sharpchanges in the Melnikov,Yu.I. 1986b.Cyclic changes of wetland wetlandecosystems of thearid areas during long- ecosystemsand conservationof rare shorebird term cyclicfluctuations in humidity.Even in periods speciesin naturereserves. In: V.E. Sokolov,I.Yu. of large-scaleresettlement, only part of the Asian Koropachinsky& I.V. Taran(eds.), Recent state and Dowitcherpopulation appears in Russia.However, perspectivesofscientific studies in Siberiannature 6,000birds recorded during the pre-breeding period reserves,pp. 148-150.Nauka, Moscow. In Russian. in theRussian part of thebreeding range represents Melnikov,Yu.I. 1986c.The experience of number aboutone third of the totalworld populationof this estimationin AsianDowitcher. In: L.V.Sopin (ed.), species.The changesin numbersat the limitsof the Organizationand technology ofgame-economy breedingrange are completelydetermined by productionpp. 11-17.Inst. of AgriculturalResearch, emigrationand immigrationprocesses. Irkutsk. In Russian. Melnikov, Yu.I. 1990. Current status of Asian Dowitcher. References In:V.A. Krasilov (ed.), Ecological problems ofnature conservation,Part 1, pp. 102-103.Nauka, Moscow. Budyko,M.I. 1980. Climatein thepast and in future. In Russian. Gidrometeoizdat,Leningrad. In Russian.350 pp. Melnikov,Yu.I. 1991.Ecology of AsianDowitcher at the Cheng,S.S.M. 1978. Dry spellsduring mid-summer over limit ofbreeding range in EasternSiberia. Ecology, SouthChina. Techn.Nat. Roy. Observ. Hong Kong. Sverdlovsk 3: 52-58. In Russian. No 44. 35 pp. Ofori-Sarpong,E. 1980.The 1975-1977 drought in Ghana. Gluschenko, Yu.N. & Shibnev, Yu.B. 1979. New materials Hydro-meteorologicalaspects. Bull. Inst. fondam. on Asian DowitcherLimnodromus semipalmatus at afr.Noize 42(4): 649-661. the Khanka Lake. In: Yu.M. Korotkov & V.A. Osipova,M.A. 1986.On breeding of AsianDowitcher in Nechayev(eds.), Biology of birds at thesouthern Far theSoutheastern Transbaikalia. In: R.L.Potapov Eastof the USSR, pp. 67-74.Far-Eastern Branch of (ed.),Studies of birdsof the USSR, their protection and USSR Acad. Sci., Vladivostok. In Russian. rational use. Material of the 9th USSR Howes,J.R. & Parish,D. 1989.New information on Asian OrnithologicalConf., Zool. Inst. of the USSRAcad. shorebirds:a preliminary review of the Interwader Sci.,Leningrad. Part 2, pp. 121-122.In Russian. Programme1983-1989 and priorities for thefuture, 32 Pasechniuk,L.E., Zhukov, V.A., Zoidze, E.K. & Mamayev, pp. Univ. of Malaya,Kuala Lumpur. L.G. 1977. Characteristics and distribution of Koshelenko,I.V. 1983. Droughtsand drought-fighting. droughtson the territory of USSR.Transactions of The review. Obninsk,VNIIGMI-MCD. Vol. 4, 56 IEM 11: 3-18. In Russian. pp. In Russian. Payevsky,V.A. 1985.Demography ofbirds. Nauka, Kozlova,E.V. 1930. Birdsof south-western Transbaikalia, Leningrad.285 pp. In Russian. NorthernMongolia and Central Gobi. USSR Acad. Pevtsov,M.V. 1883.The sketch on the travel through Mongolia Sci.,Leningrad. In Russian.397 pp. andnorthern provinces ofinner China: West-Siberian Krivenko,V.G. 1991. Waterfowlbirds and their protection. Branchof ImperialRussian Geographical Soc., Agropromizdat,Moscow. 271 pp. In Russian. Omsk. 355pp. In Russian. Liedel,K. 1982. Verbreitungund Ocologiedes

356 Melnikov: Populationand rangefluctuations of AsianDowitcher Limnodromus semipalmatus in the centralAsian arid zone

Rauner,Yu.L. 1979. Synchronyof droughtsin the cereal Velizhanin, A.P. & Velizhanin, G.A. 1929. List of birds of cropsareas of NorthernHemisphere. News of the the Bamaulregion. Uragus1: 5-14. In Russian. USSRAcad. Sc•., Geography Branch, 1: 5-19. Verheugt,W.J.M., Danielsen, E, Skov,H., Purwoko,A., In Russian. Kadarisman, R. & Suwarman, U. 1990. Seasonal Shkatulova,A.E 1973.Species composition, numbers and variationsin thewader populationsof the migrationof wadersat theChita region. In: V.E. Banyuasindelta, South Sumatra, Indonesia. Wader Flint (ed.),Fauna and ecology of waders 2: 86-89. StudyGroup Bull. 58: 28-35. Moscow State Univ., Moscow. In Russian. Yurlov,A.K. 1981. Asian Dowitcher (Lirnnodrornus Shnitnikov,A.V. 1957. Variationin totalhumidity of semipalrnatusBlyth) at the areaof the ChanyLake NorthernHemisphere continents. Notes of the (WesternSiberia). In: K.T.Yurlov (ed.), Ecology and USSRGeographical Soc. Vol. 16,335 pp. USSRAcad. biocoenoticconnections ofbirds of Western Siberia, pp. Sci.,Moscow-Leningrad. In Russian. 102-109. Nauka, Novosibirsk. In Russian. Taczanowski,V. 1873. Criticalreview of ornithological fauna Zaks, Z. 1976. Statistic estimation. Statistica. Moscow. ofEastern Siberia. Proceedings of the3rd Congress Zalessky,E 1934.Mystery birds. SiberianHunter (Okhotmk of Naturalistsof Russia.St-Petersburg. In Russian. Sibiri) 1: 22-24. In Russian.

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