Klimov: Numbers,reproductive success and geneticstructure of LapwingsVanellus vanellus

Numbers, reproductive successand genetic structure of Lapwings Vanellusvanellus in areasof varying pastoral regimes S.M. Klirnov

Klimov,S.M. 1998. Numbers,reproductive success and geneticstructure of LapwingsVanellus vanellusin areasof varyingpastoral regimes. International Wader Studies 10: 309-314.

Researchwas conductedin 1980-1989on constantsurvey squares in the greenbelt of (the CentralChemozyom Region). One hundred and fifty-four nests were observed regularly until hatching.The numberof LapwingsVanellus vanellus breeding in the studyarea decreased markedly,from 90-100 pairs to 10-15pairs. The causesof the declineare intensive cattle grazing, predationof clutchesby HoodedCrows Corvus corone cornix, Magpies Pica pica and RooksCorvus frugilegus,and recreational pressure. When moderate grazing by 100-150head of cattleoccured on a 400ha area,50-60 pairs of Lapwingsnested each year, but after the cattleherd increasedto 1,000 head,the number of breedingLapwings decreased to 10-15pairs. Averageclutch size in 1980-1983 was 3.9+0.1,and the reproductivesuccess was 46.1%. Over the next 12 years(1984-1996) the clutch sizedecreased to 3.3+0.3and the reproductivesuccess also decreased to 38.2%. In 1987-1989,clutch size(3.2+0.3) and reproductivesuccess (28.2%) was furtherreduced. We considernon-metrical characteristicsofeggs (colour and shellpattern) to be a reliablegenetic marker for differentbird groups.For Lapwing,there is a dependencebetween the scaleof intra-populationvariety (M) and thenumber of breedingpairs. Hencein 1982,when the number of breedingbirds was at its highest (100pairs), M = 2.4+0.1. In morerecent years, M hasnot exceeded2.26.

S.M. Klimov,Department of Zoology, Lipetsk State Pedagogical Institute, Lenina Str. 42, Lipetsk, 398020, .

KasMoB,C. M. 1998.qscaennocT•,, ycnex pa3m•omensn • reneTsqecKan cTpyKtypa q•i6scoB Vanellusvanellus • pafionaxc ncpcMem!•aM•4pC•MaMU sbmaca. International Wader Studies 10: 309-314.

H(;caeAonausnnponoAsa.cb n t980q989 rr. •a sna•paTaxnOCTOnUHblX sccaegonausfi n 3eae.o• noncer. AuneTcl•(qepno3e•nb•fi 8enTp). Hog peryanp.b• na6aioaeu.e• ao t54 rnc:•Aa.qncaennOCTb r.eaan½.xcn • paaones3yqcn.a qs6ucon Vanellus vanellus 3a•eT.o cnu3uaacb,OT90d00 napao i0q5 nap.Hpsqmib• co•pan•ensa qm;aeooocT• - nbmacCKOTa, pa3opea.e •aaAo• cepb•M•nopoaaM. Co•us coronecornix, (:opo•aM• Pica pica vpa.• Co•usfrugile•s, • pe•peal]•o..an•tarpya•a. Hp• y•epelt.o• .binaceot •00 Ao roao.cl•ota na nao•aA• 400 ra rne3A•a•cb•a•Ab•fi tog ot 50 Ao60 nap.6•co., a yneasqens.cTaga CKOTa Ao t.000 rOaOBKoAHqCCTBO vae3An•uxcn qsfscon G!tH3•AO('.b HaBceFo t0d5 nap.Cpea.ee soa. qecTno hug n ,aaasen 1980-t985rr. 6b•ao3,9&0.i. ycnex 6bla46A%. Bteqease caeay•o•.x 12 aCT (1984-t996) paa•ep •aaa•. y•enbmsacsua cuss.acs.ycnex pa3•no•ensn .a 382%.B nepsoac 1987no 1989 r. ,a, pa3Mep (5,2&0.3).Ta,. ycnexpaa•nome.nn (28,2%) 6blan e•e nnme.Mbl cq.tae• ueMeTp.qec,m• xapas•epncT.•.nm• (ospacsa • p•cyno, csopaynb0 •taae•ub• reneTuqec•.• .nAm•aTopoM Aaspa3anqnb•x rpynn nTn•. Aaa qu6ncaeCTb 3aBBC•MOCTb MemAy cTeneubD ,nytpsnonyasilsoanoropa3noo6pa3ss (M)s soasqecTso•rneaAm•sxcs nap. i982r., aoraa q.cao r.e3as•.xcs ntm! aoctsrao nn,a (i00 nap),M= 2,4k0,i.B nocaeaaseme roA•xscanquna M .e npesb•maaa2,26.

Introduction sidesof the processof humanmodification of naturallandscapes such as occurs on farmland. More than60% of thetotal human population on the earthnow livesin largecities and towns,making Birds,being one of the mostnoticeable and well- the impactof humansone of the mostimportant studiedvertebrate groups, can serve as good models currentenvironmental issues. An improvementin for suchstudies. In many partsof EuropeanRussia the relationshipbetween man and natureis now the impoverishmentof bird populationsas a result extremelyurgent. Scientificknowledge of the reactions of natural communities to different kinds of humaninfluence has been slowed during the last two decades.Indeed, for severalspecies, including of anthropogenicinfluence will helpachieve this LapwingVanellus vanellus, an increasein numbers end. Long-termstudies enable a better hasbeen recorded. The biology and ecologyof this understandingof boththe positiveand negative specieshave beenrather well studied,although

3O9 International Wader Studies 10: 309-314

thereare generallyfew breedingdata from within mapped. Eachnest was examinedregularly; from urban landscapes. six to ten timesduring the nestingperiod. Periodic standard wader counts and searches for nests were Study Area and Methods also carried out. Our studies were conducted from 1973-1990 in the At the sametime as this work was taking place, town of Lipetsk,situated in the centralpart of similarstudies were carriedout on a fifth sample EuropeanRussia, and in the surroundingarea, a plot 40 km from Lipetsk,which was characterisedby total areaof 400ha (Figure1). Between1980 and only slightagricultural influence (sporadic 1989,studies were carriedout on four sampleplots pasturingand annualhay makingfrom the second eachof 5 ha, of varyinglevels of anthropogenic half of June onwards). influence.All the plotswere located on moistflood- plain meadowswith pools,ox-bow lakes, swamps Eggsuccess was estimated as the proportionof and shrub-thicketsalong the Voronezhand the hatchedchicks from the totalnumber of eggsin Matyrarivers. All themeadows were used for nestsbeing studied. pasture(mainly by cows)and hay making;on plots 2 and 3, moderatepasturing occurred, while plots1 As well as these traditional field methods, a new one and 4 sufferedintense grazing. Plot 4 waswithin an wasused: the pheneticapproach, based on the areaof activerecreation and thushay makingonly analysisof the colourof the egg-shell(Klimov et al. tookplace locally. 1989,1990). As hasbeen shown by Myand (1988), by our studies(Klimov 1987, 1990,1991; Klimov & Datesof arrival and the beginningof nest- Ovchinnikova 1988, 1992; Klimov et al. 1990, 1992a, construction,egg-laying and hatchingwere 1992b,1992c; Ovchinnikova & Klimov 1987),and by recordedannually for all waderspecies. Every nest was marked with a metallic stick and its location Vengerov(1989, 1990, 1991, 1992), eggs are convenientobjects for the analysisof morphological variationin bird populations. The colourof a Lapwingegg-shell is due to two components- diffuselyspread pigment, called the backgroundcolour, and variouslines, spots etc., calledthe tracery.Lapwing egg-shellcolour is

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5

ß; Elec Lipetsk

Figure1. Studyarea and location of controlplots. 1 - studyarea; 2 - controlplots with numbers;3 - administrative boundaryof Lipetskregion.

310 Klimov: Numbers,reproductive success and geneticstructure of LapwingsVanellus vanellus thereforevery complicated,as in the majorityof Results and Discussion open-nestingbirds. Variationsin background colourcan be separatedinto threegroups: sandy Regularobservations were made on 228Lapwing (varyingfrom grey to olivetints, including those nests,the majority of whichwere observed until with light-bluebackground colour), green-brown hatching.During the studies,the breeding (backgroundcolour from greento tobacco-brown) populationof Lapwingwithin the studyarea and intermediatesandy-green-brown. The tracery declinedfrom 90-100pairs in the early 1980s,to 10- canalso vary, depending on the form and 15 pairsin the late 1980s(Table 1). Thiswas caused combination of the elements, and their distribution mainly by intensivegrazing, destruction of nestsby and densityover the egg-shell.The following corvids(Hooded Crow Corvuscorone cornix, Magpie tracerypatterns can be distinguished: Picapica, Rook C.frugilegus), and recreational pressure.When moderategrazing of 100-150head of cattleoccured on a 400 ha area,50 to 60 Lapwing 1. Form and combinationof traceryelements pairsbred annually;when the herd increasedto Althoughthese elements differ in shapeand size, 700-1,000head of cattle, numbersof breeding they canbe includedinto oneof two geometric Lapwingdeclined to 10-15pairs (Table 1). At the groups:spots or lines. Accordingto their sametime, changesin the distributionof Lapwing combinationon the egg-shell,four typesof tracery colonies and in the behaviour of birds were were distinguished: recorded.At the beginningof the study about95% 1.1. Spottytracery - only the spotsare present; of Lapwing colonieswere situatedon flooded 1.2. Spotty-linear- bothlines and spotsare present, meadows,while the restof the birdsnested on bogs althoughspots prevail; and hillockymarshes. In the later yearsof the study 1.3. Linear-spotty- bothlines and spotsare present, Lapwingssettled mostly on the floatingmats of linear traceryprevails; or bogsand, more rarely, on hillockymarshes, in 1.4. Linear- only linesare present. groupsof threeto five pairs. This situationwas observedat all the plotswithin the areasof 2. Distributionof traceryelements over the farmlandexcept plot 3, from whichLapwings whole egg-shell disappearedin 1984after it becameovergrown with In general,the traceryelements are circularly scrub. concentratedat the obtuseend of the eggs,rarely at the acuteone, and still morerarely at both of them. We believethat the declinein eggsuccess was due In addition,the traceryelements can be to the activitiesof avianpredators, whose numbers concentratedaround the equatorialpart of the egg, in the study areaare ratherhigh: 15 pairs of Hooded or spreaddiffusely over the whole egg. Thus,five Crows,70 pairsof Magpies,and sevenpairs of typesof distributionof traceryelements are Marsh Harrier Circusaeruginosus (Klimov & recognised. Aleksandrov1990). Low averageclutch size, recordedin the last four years(Table 1) was 3. Density of tracery explainedby the increasein the numberof Threetypes of traceryaccording to its densityon the replacementclutches, which were usuallysmaller. egg-shellwere distinguished: 3.1. Scarce- lessthan 30%of the egg-shellcovered On the areaswith moderategrazing, Lapwings with traceryelements; oftenformed mixed breeding colonies with other 3.2. Dense- traceryoccupies 30% to 70%of the egg- bird species,where the birdsdefended their nests shell; or moresuccessfully against corvids and Marsh 3.3. Solid- morethan 70%of the egg-shellis Harriers. The differentdefence strategies of the coveredwith tracery. differentspecies in mixedcolonies probably increasesthe generalresistance of suchcolonies to For the quantitativeestimation of this character predationpressure (Aleksandrov 1985, 1986). The under laboratoryconditions, we used a formation of mixed colonies within farmland can be polyethylenestrip 10 x 10 mm in sizewith a 1 mm consideredto be oneof the adaptationsof birdsto grid. The densityof traceryelements was calculated the presenceof humans(Klimov 1988).At the same accordingto the schemedescribed for two partsof time, if the pressureof humanactivities, particularly the egg:close to the obtuseand to the acuteends. In agriculturalones, is evenmore intense, this kind of the field, theseestimations were madevisually adaptationis not effective.The increasedsize of the withoutthe grid, as any under-estimationwas not cowherd in our studyled to the tramplingof places largeand at the sametime muchmore material which were suitablefor breeding,to the destruction could be collected. and separationof mixedcolonies into very small and mostlymonospecific ones, and finally,to the Statisticalanalysis was carriedout accordingto declinein breedingsuccess of Lapwing (Table1) as Zhivotovsky(1982). Usingthe threeegg well asother breeding bird species. characteristicsdescribed, an averageindex of within-populationcolour diversity was calculated. Studiesof Lapwingecology in plot 5 with slight Usingthis index,one can describe the differences agriculturalinfluence revealed stable breeding betweenlocal populations and any relationships numbersand a largerclutch size (mean = 3.8+0.2SE with differinglevels of humaninfluence. for five years)in comparisonwith moreintensively

311 International Wader Studies 10: 309-314

Table 1. Numbers,clutch size, breeding success and geneticstructure of Lapwingpopulation in the pastoralareas (plots1-4) of Lipetsk. 1980 1981" '1982 1983 1984 1985 1986 19•7 1988"' I•89"

......

Size of cow-herd 100-150 100-150 100-150 100-150 200-500 200-500 200-500 700-1000 700-1000 700-1000 on the 400 ha area

Number of nests studied 14 22 28 38 10 10 8 9 7 7

Estimated number of Lapwingnests on 400ha 40-50 40-50 90-100 70-80 50-60 40-50 20-25 20-25 10-15 10-15

Clutch size rn + SE 3.81+0.2 3.9+0.1 3.9+0.1 3.9+01 3.2+0.3 3.7+0.2 3.1+0.3 3.2+0.3 3.3+0.3 3.1+0.3

Coefficient of variation for the clutch size (%) 15.3 7.5 10.7 9.8 28.7 18.2 32.7 25.9 23.0 28.6

Egg success(%) 42.7 46.5 51.0 44.4 38.7 39.6 36.2 30.5 28.3 25.8

Indexof intra-population diversity + SE 2.00.2 2.10.2 2.40.1 2.30.0 2.20.2 - 2.00.2 -

Table2. Numbers,clutch size, breeding success and feneticstructure of Lapwingpopulations in areassurrounding Lipetsk town and subjectto only slightagricultural management.

-i983 1984 1985:. 1986 1987

Size of cow-herd on the 400 ha area 150-200 150-200 150-200 250-400 250-400

Number of nests studied 18 10 12 11 15

Estimated number of Lapwingnests on 400 ha 60-70 40-50 50-60 40-50 50-60

Clutch size +SE 3.9+0.1 3.8+0.2 3.9+0.2 3.8+0.2 3.9+0.2

Coefficient of variation in clutch size (%) 12.3 16.3 15.2 16.1 14.3

Eggsuccess (%) 50.5 46.2 48.6 43.5 48.8

Index of intra-population diversity(m) + $E 2.30.2 2.20.2 2.20.1 2.20.1 2.30.2

managedareas (mean=3.6+0.2 SE for ten years); It is clearthat Lapwingswhich breedin chronologicalvariation of clutchsize and phenetic agriculturallymodified areas differ greatlyin a diversityof egg-shellcolour in Lapwingsfrom plot numberof ecologicalcharacteristics from those 5 werenot large (Table2). Changesin plot 5 similar which breed in less modified habitats: dates of to thosein the moreintensively managed area were particularstages of reproduction,length of breeding alsoobserved during the lastyears of the study. period,clutch size, variation of phenetic Due to an increasein cattle-grazing,Lapwings parameters,egg-success etc. Birdsmay be startedto breedon the floatingparts of bogsand reasonablysuccessful in thoseareas managed by total numbers declined. men when the human influence on them remains moderate. Lapwingsbreeding in agriculturallymodified areas startnesting 7-10 daysearlier than birds in areas with little human influence (Table3). In addition, the breedingseason in agriculturalareas is longer due to high ratesof nest-failuresand clutch replacement.

312 Klimov: Numbers,reproductive success and geneticstructure of LapwingsVanellus vanellus

Table 3. The earliestdates of arrival and startof egg-layingin Lapwingsbreeding in the farmland areascompared with the slightlytransformed habitats.

Farmland areas: transfor areas Year Arrivaldate Layingdate Arrivaldate Layingdate

...... ß

1980 2 April 19 April no data no data

1981 25 March 10 April no data no data

1982 31 March 17April 1 April 24 April

1983 23 March 15 April 25 March 23 April

1984 24 March 10April 24 March 20 April

1985 28 March 15Apr"l no data 22 April

1986 27 March 15April 26 March 23 April

1987 31 March 18April 31 March 26April

1988 29 March 17April 30 March 25 April

1989 30 March 17April 1 April 26 April

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Vengerov,ED. 1989. Oomorphologicalcharacteristics of Vengerov,ED. 1992. Variationof eggswithin clutchesas Magpieand SongThrush in the urbanpark and in oneof the typesof individualvariation of birds. the nature reserve. In: A.P. Tsherbakov & O.P. Bull.Moscow Soc. of Naturalists, Biol. Branch, 5: 3-8. Negrobov(eds.), Monitoring and conservation ofthe Vengerov,ED. 1992. Comparisonof oomorphological environmentin CentralRussia, pp. 87-88.Voronezh parametersof birdsfrom natural and urban Univ., Voronezh. habitats.Ecology 1: 21-26. Vengerov,ED. 1990. Levelof intra-clutchvariation as the Zhivotovsky,L.A. 1982. Indexesof populationvariation morpho-physiologicalindicator of individualand accordingto polymorphicsigns. In: A.V. Yablokov populationstate. In: O.P.Negrobov (ed.), Problems (ed.),Fenetics of populations, pp. 38-44.Nauka, ofcadastre and conservation ofthe animal world of Moscow. Russia,pp. 82-83. VoronezhUniv., Voronezh. In Russian.

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