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Reproduction and Survival in a Declining Population of The Reproductionand Survivalin a Declining Population of the Southern Dunlin Calidris alpina schinzii Paul Eric J6nsson J6nsson,P.E. 1991.Reproduction and Survival in a DecliningPopulation of theSouthern Dunlin Calidrisalpina schinzii. Wader Study Group Bulletin 61, Supplement: 56-68. Reproductivesuccess and survival was studied in a declining population of theSouthern Dunlin in SW Sk,Sne,SSweden, in 1981-1986.Hatching success was low; on average only 30% of clutches hatched, dueto intense nest-predation from mainly Crows, Foxes and mustelids. Some nests (8%) were also lostby tramplingfrom grazing animals. Within a season,there was no significantdifference in hatchingsuccess between early and late clutches, but the overall annual hatching success was higher in yearswith an early onset of breeding.On average,the Dunlins yearly produced 0.3 - 0.4 hatched young/adult.Fledging success was estimated at36% and survival from fledging tothe age of one year at56%. Adult survival, estimated from rerum rates, averaged 89% in males and 77% in females (83% of thesexes combined). The sex-relateddifference in return-ratesbelieved to reflecta loweractuai survivalrate in females, assuggested bythe male-skewed sex-ratio inthe population. The average life expectancyfor an adult Dunlin is estimated at5.4 years and the mean longevity at7.4 years. Survival tendedto decrease with increasing age (up to 5 yearsafter ringing). The oldest bird recorded in the studywas 17 yearsold. The observed reproductive rate was found to be insufficientto maintaina stablepopulation and unless nest-predation issignificantly reduced, the Dunlin will become extinct in SW Sl•ne within20 to 30 years. PaulE. J6nsson,Dept. of Ecology, Ecology Building, University of Lund, 223 62 Lund, Sweden. Introduction Study area and methods On the south-easternedge of its distribution, Thestudy was performed in theyears 1981-86, alongthe coast of theBaltic andthe North Sea, on the coastalwet meadowsof SW Skfine, the SouthernDunlin Calidrisalpina schinzii SouthSweden (Figure 1). Duringthe study hasdecreased markediy in numbersduring the period, these meadows held the bulk of the past50 years(Gromadzka 1983, Cramp& Sim- Swedishschinzii-population; 60-80 pairsout mons 1983, Piersma 1986). The main mason of anestimated total of about250 pairs (Tjern- for thisdecline is believedto bethe rapidly di- berg1985). Due to alteredmanagement (abo- minishingareas of suitablebreeding habitat, lishedgrazing and hay-cutting) and exploita- i.e. wet meadowsand pastures, grazed by do- tion (expansionof urbanand industrial areas) mesticanimals, or otherwiseharvested by man of wetmeadows, suitable breeding habitats for (Larsson1969, Dybbro & JOrgensen1971, Dunlins and other waders have become scarce Soikkeli&Salo 1979,Emanuelsson & Kjel16n andfragmented in the whole of SouthSweden 1981, Hansen 1985, Tjernberg 1985, Kr61 (Emanuelsson& Kjell•n 1981, Tjernberg 1986). 1985).The totalarea of typicalDunlin-habitat in SW Skfineis, at thetime of writing,less than In thispaper I presentthe results of a six-year 300 ha, most of which is situated around the studyon a decliningpopulation of theSouthern shallow Foteviken Bay (about 55ø17'N, Dunlin in SW Skfine, South Sweden. Data on 12ø58'E). reproductiverate and survivalare presented, andthe possible factors controlling the popula- Themain study area, Vellinge •ingar (South), is tion parametersand maintenance of thepopu- locatedon theeastern shore of FotevikenBay lation are discussed. (Figures1 and2) andis comprisedof approx. 60 haof wet,intensively grazed shore meadow. WSG Bulletin61, Suppl.: 56-68 (April 1991) J6nsson: Dunlln 57 Table 1. Mean population size and density of breedingDunlins in SW Sk•ne, 1981-1986. Number Maximum Area of breeding (ha) pairs density (pairs/ha) Main study area Vellingelingar (S) 60 14-24 0.40 Control areas VellingeItngar (N) 58 15-20 0.35 Eskilstorps1tngar 55 10-25 0.46 Kungstorp 9 1-2 0.22 Hammars Nlts 33 3-5 0.15 Ar16vs1tngar 44 3-4 0.09 Jtmgsniiset 10 1-2 0.20 the visits to the control areas were to estimate the numberof breedingbirds and to checkfor andidentify marked individuals, however, I al- so ringed a few adults and chicks. In 1983- 1985, the main studyarea was expandedto in- cludealso Vellinge gngar (North) and.•ngsng- set. i THESAœTI• SEA Nests were searchedfor by careful investiga- tions of suitableparts of meadows,and by Figure 1. Locationof the studyareas (in black) on the coast of SW SloPe, South Sweden. Fields N This locality still holdsone of the largestcon- centrationsof breedingDunlins in Sweden(15- 25 pairs during the studyperiod). The areais FOTEVIKEN BAY separatedfrom thenorthern part of Vellinge'•'a- gat by a ditchand a small(140 x 80 m) treeplan- 0 50 tO0 1SOrn I Fields tation. However, grazing animals can move freelyover the whole area, totalling almost 120 ha. The meadowsare grazed annually, from late May to well into October,by 150-250 cattle • FeslucaLollurn perenne-veg.rubraI and 50-80 riding horses.A furtherdescription !!i}::i[!:'Juncus gerardii-veg. of the areais givenin J6nsson(1985). .:'!•?' ! Reedbeds :.'.?. Fields The control areas consistedof the remaining Dunlin localities in SW Skfine(Table 1), most L• Baresand of which are located within 4 km of the main studyarea; although two localities, •ngsngset and Ar16vs, are more distant (8 and 22 km, re- spectively)(Figure 1). The Danish island of Saltholm,which in 1980 held up to 15 pairsof Dunlin (Hansen1985), is situated25 km NW of Vellinge th'agar,but was for practicalreasons not includedin this study. In the main studyarea, my aim was to capture andmark asmany birds as possible, and to find Figure2. Map of the mainstudy area, Vellinge •tngar all nestsin everyseason. The main objectives of (S), showing the distributionof some important vegetationtypes. WSG Bulletin 61, Suppl.: 56-68 (April 1991) 58 J6nsson: Dunlln watchingthe behaviour of thebirds. During the tions (see Soikkeli 1973, Ferns & Green 1979 firstperiod of egg-laying,from mid-April until and J/Snsson1987). mid-May,nests were relatively easy to findjust by searchingon the dry spotswith low vegeta- On everyvisit to thestudy areas, colour-ringed tion;i.e. the only placeswhere it waspossible birdswere lookedfor and, if possible,identi- for the birdsto lay their eggsat that time. Later fied usingbinoculars (10x40) and a telescope in the season,with virtually the whole area (20-40 x 60). With favourablelight conditions, driedup andwith highervegetation, nests were colour-ringscould be read accuratelyat a di- more difficult to find (see JOnsson1985). On stanceof 150to 200meters, using the 40x mag- averageI estimatethat 75-95 % of all nestsin nif'lcationocular on the telescope. the main studyarea, were foundannually. All nestsand broodswere plottedon a field Results map,and their fate subsequentlyrecorded. The hatching date of incubated broods was Nesting success estimatedby immersingthe eggsin water(see Westerskov (1950) and van Raasen et al. The frequencyof clutchesthat surviveduntil (1984). Nest losseswere recordedas well asthe hatching,varied betweenyears from 16% in probable causesfor these losses (predation, 1983, to 38% in 1984, averaging30% for the tramplingor desertion).In predatednests, signs whole studyperiod (Table 3). of the predatorwere looked for. Thus,a largemajority of the clutcheslaid each Adult birds were caughton the nestduring in- year weredestroyed or deserted.The mostim- cubation,using walk-in-traps (see Bub 1974), portantfactor affecting nest survivalis preda- while chickswere caughtby hand,as soonas tion. In total, 60% of all clutches were taken, possibleafter hatching. completelyor partially,by predators.In most casesno signof the predatorwas found, but All birds caughtwere ringed with one metal- sometimesegg-shell fragments showed the ring and 2-5 colouredplastic rings. A unique characteristic marks of Crows Corvus corone combinationof colour-ringswas usedfor each or small Mustelids(most probablyStoat Mu- single individual. Table 2 gives the annual stelaerminea). Another important predator on numbersof ringedbirds in the main studyarea Dunlin eggswas probably the Red Fox Vulpes and the control areas. vulpes,which was seen regularly patrolling the meadows for waders' nests (see also J/Snsson Sex, in adultbirds, was determinedfrom plu- 1985). magecharacters and measurements, in combi- nation with subsequentbehavioural observa- Partial predation,i.e., where just one or two Table 2. The annual numbers of Dunlins trapped and ringed in the main study area and the control areas.Left figure = adults,right figure= young. 1981 1982 1983 1984 1985 1986 Vellingeartgat (S) 43/31 14/18 9/3 1/6 5/6 1/- Vellinge•tngar (N) -/- 1/- 15/7 23/28 10/25 3/3 Eskilstorps -/- -/- 2/7 -/- -/- -/- Kungstorp -/- -/- -/- -/- -/- 3/- Hammarsnlts -/- -/- -/- -/- -/- -/- ArlOvs•tngar -/- 1/3 1/- -/- 2/2 1/- •ngsn'aset -/- 2/4 1/4 -/- -/- -/- Total 43/31 18/25 28/21 24/34 17/33 8/3 WSG Bulletin61, Suppl.: 56-68 (April 1991) J6nsson: Dunlln 59 Table 3. Nestingsuccess in a Dunlin populationin SW Sk•ne in 1981-1986.Figures refer to clutches. Hatched Predated Trampled Deserted Total n (%) n (%) n (%) n (%) n (%) 1981 13 (36.1) 20 (55.5) 2 (5.6) 1 (2.8) 36 (100) 1982 8 (28.5) 18 (64.3) 1 (3.6) 1 (3.6) 28 (100) 1983 6 (16.2) 26 (70.3) 4 (10.8) 1 (2.7) 37 (100) 1984 12 (37.5) 18 (56.2) 2 (6.3) 0 (0) 32 (100) 1985 12 (29.3) 21 (51.2) 7 (17.1) 1 (2.4) 41 (100) 1986 9 (31.0) 19 (65.5) 0 (0) 1 (3.5) 29 (100) Total 60 (29.6) 122 (60.0) 16 (7.9) 5 (2.5) 203 (100) ,_ eggswere taken from the clutch,accounted for = 11.3,0.
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