Nest Prospecting By Common Goldeneyes - University Of New Mexico

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The Condor 91:807-8120 The Cooper OrnithologicalSociety 1989NEST PROSPECTINGBY COMMONGOLDENEYESMICHAEL C. ZICUS AND STEVEN K. HENIVES*Minnesota Department of Natural Resources,Wetland WildrifePopulationsand ResearchGroup,102 23rd St., Bern@, MN 56601Abstract. We studied nest prospecting by Common Goldeneye (Bucephala clangula)females in north-central Minnesota. Adults unsuccessfulin nesting,those with broods, andnonnesting yearlings were captured in nests while prospecting.Prospectingbegan in lateMay and continued into early July. Active nests received up to 25 prospectingvisits perday with most visits occurringbetween 06:OOand 09:OOCDT. Adults appearedto prospectmore (P 0.05) in nest boxesthat had contained successfulnestsduring the current seasonthan in those where nests were abandoned or destroyed or those that were unused. Nestbox statushad no apparent effect on prospectingby yearlings. Body mass of prospectingadults that were unsuccessfulnestersand yearling nonnesterswas similar and was significantly less(P 0.05) than that of females still incubating nestsor those with broods. Ourobservationssupport the claim that prospectingfemales are preparingfor the next breedingseason,and we suggestthat prospectingis a meansof confirming information alreadygainedduring the current season.Prior knowledge of successfulnest sites could explain the preferential use of previously successfulnest boxes observedin a Swedish study.Kev words: Common Goldeneve:Bucevhala clangula; nestprospecting;nest boxes;postbrieding activity.INTRODUCTIONSelection of a good nest site is obviously important to successful reproduction, and nest-sitesearching behavior prior to egg laying has beendescribed for many ducks (e.g., Bennett 1938,Sowls 1955, Mendall 1958, Bellrose 1976).Another form of nest-site examination occursnear the end of the nesting period in hole-nestingBucephalu and Tadorna species(Bellrose 1976,Patterson and Makepeace 1979, Eadie and Gauthier 1985) and has been labelled “nest prospecting.” Eadie and Gauthier (1985) discussedpossibleinformation gatheredwhile prospecting,how it might be obtained, and the evolutionarysignificanceof the behavior. They proposedthatprospecting helped Bucephala females preparefor the next nesting season. By doing so, henscould determine availability and suitability ofscarcenest sites and possibly past history of thesite. They suggestedthat hens could obtain information by visits to active nests, by followingincubating females to their nests, or perhaps bydetecting the presence of shell fragments andmembranes in hatched nests.Exceptfor the workof Eadie and Gauthier (1985) and Patterson andI Received 2 1 December 1988. Final acceptance12June 1989.2Present address:5 174 N.E. Willamette Ave., Corvallis, OR 97330.Makepeace (1979) virtually no data have beenpublished concerning prospecting behavior incavity-nesting ducks.After reviewing 22 years of nesting data, Dowand Fredga (1985) concluded that CommonGoldeneye (B. clangulu) females tended to nestin sites that were occupied in previous years,especially successfulsites.The tendency was apparent even after subsequent reuse of a site bythe same female was eliminated from their analysis. It was not clear how females were able tochoose nests with a past history of success.Wehypothesized that females might learn about asite’s successduring the current nesting seasonand that prospecting might be a means of confirming this information. If so, then we predictedthat more prospectingwould occur in successfulnests than unsuccessfulor unused ones. We report data related to this prediction as well asadditional information about the timing of andparticipation in nest prospecting by CommonGoldeneyes.METHODSOur study was conducted in north-central Minnesotanear the southernedgeof the species’range(Bellrose 1976).From 1982-1985,weworkedonIsland Lake, a 1,250-ha lake having moderate toheavy year-round and summer residential development. From 77 to 90 nest boxeswere present along the shoreline, and goldeneyeslaid eggsPO71

808MICHAEL C. ZICUSANDSTEVEN K. HENNESin approximately 70% of the boxes each year.Beginning in April, we inspected nest boxesweekly and recordednesting activity. When present, females were caught in the box, leg-bandedwith U.S. Fish and Wildlife Service bands,weighed to the nearest 5 g, and released.We collected nest prospecting data from 47nest boxes to which we had easy access.All buttwo of thesenest boxes contained eggsin at least1 year during 1982-1985. From 13 to 24 June1984 and 18 to 28 June 1985, we set nest traps(Zicus 1989) in boxesnot containing active nests.Traps were set in individual boxesfor 3 to 6 dayseach year and were checked twice daily between09:00-l 2:00 and 18:00-2 1:OOCDT. Nest boxeshaving traps were classifiedas (1) successfulif abrood had departed, (2) abandoned/destroyed,or (3) unused if no eggshad been laid in the boxthat season.Ducks captured while prospecting wereweighed, leg-banded if not already banded, andreleasedat the site. Common Goldeneyes do notnest as yearlings, and at least the Juvenal wingis retained into June or even July while the BasicI and Alternate I plumages are being acquired(Palmer 1976, p. 377). Because of these traits,we found Camey’s (1983) descriptions contrasting adult and immature wing plumage in falluseful to separate adult and yearling females inspring. Adult females were distinguished fromyearlings by the presenceof a distinct black bandacrossthe end of greater secondarycoverts alongwith several rows of white lesser secondary coverts. These characteristicsappearedas two whitewing bars, separatedby a dark bar, ahead of thewhite secondaries.In contrast, wing patterns ofyearling females appeared as a single grayishwhite patch ahead of the white secondaries.Adults and yearlings could also be distinguishedin flight by these patterns. Reproductive statusof most females we caught was known from ourweekly checksof all boxes on the lake. Femaleswere classifiedas (1) unsuccessfulif they had losta nest through abandonment or predation, (2)with a brood if they had hatched a nest we hadbeen observing or if we observed them rejoin abrood after having been trapped, or (3) yearlingnonnesters.Daily and hourly visitation rates to boxeshaving active nests were estimated during June andearly July for prospecting females using remoterecording sensor systems (Cooper and Afton198 1). These devices allowed us to record activ-ity in the nests at all hours. Becausethe systemssimultaneously used an event recorder at the boxentrance and a thermistor in the nest bowl, wecould distinguish departure and return by incubating females from “visits” by other birds ormammals based on consistent patterns of temperature change recorded on the strip chart. Although we could not be certain that all visitsrecordedwere by prospectinggoldeneyehens, wenever observed any speciesother than goldeneyes entering nest boxes at this time of year, andwe only once caught a speciesother than a goldeneye with our traps. Thus, we believe virtuallyall recorded visits were by prospecting goldeneyes.Daily capture rates for the three different nestbox classifications were examined using a oneway analysis of variance (ANOVA) after testingfor equal variances and normality. Individualmeans were compared using Fisher’s LSD procedure (Milliken and Johnson 1984). The firstcapture of a bird was used in the analysis, andwe performed separateANOVAs for adults andyearlings. Body mass of all females weighed between 13 and 28 June and having a differentreproductive status was also compared using aone-way ANOVA and Fisher’s LSD procedure.In addition to thosefemales captured while prospecting, we compared body mass of incubatinghens from Island Lake and those accompanyingbroods and captured by entanglement netting(Johnson 1972) on all area lakes during 19821985.RESULTSSeventy-four female Common Goldeneyes andone female Hooded Merganser (Lo&&escucullatus)were caughtprospectingduring 3 10 trapdays. Individual females were captured as manyas five times in the same season(Table 1). Thirtyof the 44 unsuccessfulnesters had been bandedas adults prior to the year during which they werecaught prospectingand were at least 3 years old.The remainder was captured for the first timeduring the current nesting season. Five previously banded goldeneye hens had successfullyhatched broods in nests we had been observingearlier in the season. We were not certain thatthese hens still had broods when they weretrapped while prospecting. The sixth brood henwas unbanded when caught, but called ducklingsfrom the shoreline vegetation and led them awayafter being released. Two yearlings had been

NEST PROSPECTING809TABLE 1. Same-seasoncapture frequency of femaleCommon Goldeneyestrapped prospectingin nest boxes on Island Lake, Minnesota, between 13 June and28 June, 1984-1985.Female statusIFrequency2UnsuccessfulnesterYearling nonnesterWith broodUnknown (adult plumage)28166611200b35:0banded as flightlessyoung in the previous year.Unsuccessful nesters tended to be recapturedmore frequently than yearlings or brood hens.Adult females were trapped at different ratesin boxeshaving different classifications(F 9.26;df 2, 68; P -c 0.001). The daily capture rate ofadult females was higher in successfulnest boxesthan either those having abandoned/destroyednestsor thosethat were unused (Table 2). Clutchsize in the 14 successfulnests ranged from 9 to19 (X 13.5) and was not correlated with adultdaily capture rates (r -0.01, df 12, P 0.974). In contrast to adults, no differenceswereapparent among nest-classificationcapture rates(F 0.82; df 2, 68; P 0.45) for yearlings,and the rate averaged 0.06 birds/day.Field observations indicated that visits to nestboxes by prospectingfemales began in late May,when loose flocks comprised primarily of adultfemales began forming. These flocks of femalesflew around and vocalized while visiting differentnest boxes in a manner similar to that reportedand discussedby Eadie and Gauthier (1985, p.53 1). Eight nests being incubated by goldeneyeswere monitored with recording systems from 2June to 8 July. Each day, from one to five nestswere monitored and a total of 109 complete nestdays of data was obtained. From zero to 25 prospecting visits per day (median 2) occurred ina monitored nest (Fig. 1). Visits to these nests5015102520Visits per Nest-dayFIGURE 1. Prospectingvisits by female CommonGoldeneyeson Island Lake, Minnesota, to eight activeCommon Goldeneye nestsmonitored for a total of 109complete nest days, 1984-1985.were frequent throughout June, but rates appeared to decline in July (Fig. 2) as unsuccessfulfemales and yearlings left the study area. Incubating females often were present during prospecting visits (Zicus, unpubl. data), and mostvisits occurred between 06:OO and 09:OO CDTwith a smaller peak in the evening (Fig. 3). Onenest received 15 prospecting visits between 06:00 and 07:OOduring one morning. Sunrise andsunset occurred at about 05:30 and 21:30, respectively, at this time of year. Except for onenest in which the entire clutch of five eggswasinfertile, all monitored nestshatched successfully.TABLE 2. Capture rates (birds/day) of prospectingadult female Common Goldeneyes on Island Lake,Minnesota, 1984-1985.RateNest nR”SE1439180.41 A0.17 B0.12 B0.060.030.04aMeans having differentlettersare different(Fisher’s LSD procedure,P 0.05).712June17222727JulyFIGURE 2. Average daily visitation by prospectingfemale Common Goldeneyeson Island Lake, Minnesota, to eight active Common Goldeneye nests.Fromone to five nestswere monitored daily 2 Juneto 8 July,1984-1985.

810MICHAEL C. ZICUSANDSTEVEN K. HENNESTABLE 3. Body mass(g) of female Common Goldeneyes trapped on Island Lake, Minnesota, between 13and 28 June, 1984-1985.MassFemale statusIncubatingaUnsuccessfulnesterYearling nonnesterWith broodbWith brood400800120016002000Hour of Day (CDT)FIGURE 3. Averagehourly visitation by prospectingfemale Common Goldeneyes on Island Lake, Minnesota, to eight active Common Goldeneye nests monitored for a total of 109 complete nest days, 19841985.Body mass of females trapped between 13 and28 June varied with their reproductive status (F 37.26; df 4, 94; P 0.001). Prospectingunsuccessfulnesters and yearlings were lightest,whereas females trapped late in incubation wereheaviest (Table 3). Females that we believed hadbroods and that we retrapped while prospectingdid not differ in weight from those captured withbroods by entanglement netting on several nearby area lakes.DISCUSSIONResults of our study should be interpreted cautiously. Females often, but not always, prospected in flocks, and we frequently observed different individuals inspecting the same nest boxone after another. As evidenced by the activenestswe monitored, some boxes were visited byprospecting hens many times in one day. Occasionally, we observed two females occupy a boxsimultaneously. With the trapping method weused,a maximum of two birds per nest box couldbe caught each day. Once a hen was trapped ina particular nest, the box was no longer availablefor inspection by other females until we removedthe bird and reset the trap. Becauseof this limitation, capture of adult and yearling females orthose having a different reproductive statuscannot be considered independent events. We alsodo not know how unavailability of certain nestsites for part of the day might have changedtheattractiveness of those remaining available forprospecting. Certainly, the temporary elimination of the nest box prevented other females fromnPSE194418612659.7 A577.4 B580.8 B613.3 C612.5 C6.63.15.513.310.8- Trapped 1 to 22 daysbeforehatching(median 4 days).bTrappedwhile prospecting.CAlso includesfemales trapped by entanglementnetting on severalother nearbyarea lakes, 1982-1985.dMeanshaving differentlettersarc different(Fisher’s LSD procedureP 0.05).inspectingwhat might otherwisehave been a veryattractive site.Adult Common Goldeneye hens appeared toprospect more in nest boxes that had successfulnests during the current year than in those thatdid not. We believe this selection would be unlikely if prospectingwas the only means a femalehad to obtain information about a nest site. Morelikely, prospectingfemaleswere searchingfor potential future nest sites and at the same timeconfirming information about current nest useby other females already gained during the nesting season. Apparently, this information is notacquired early in the seasonbecauseadult capture rates were not higher in successfulneststhathad been parasitized during laying, and abandoned nests often had large clutches (i.e., 15eggs)but low capture rates. These nests, used byseveral females, would have received much activity early in the season.If this activity was thesourceof information for prospectingfemales, itshould have influenced capture rates.Our observations support the belief that prospecting females are preparing for the next nestingseason (Grenquist 1963, Eadie and Gauthier1985) and may explain why Dow and Fredga(1985) observed a tendency for females to nestin boxes that had been used the previous year.Subsequent use was evident even when reuse ofthe same nest by the same female was eliminatedfrom their analysis; use was especially high forboxes having successfulnests the previous year.Predation was a significant factor, and Dow andFredga (1985) believed that females nesting inpreviously successfulnest sitesbenefited becausethey were less likely to lose nests to predation.They also speculated that females might recog-

NEST PROSPECTINGnize these sites through “familiarity” with thearea.The status of a nest box apparently did notinfluence the daily capture rate of yearlings. Ifthis contrast with adults is real, and not an artifact of our trapping, it further supportsour suppositions about how information is obtained andabout the function of prospecting. We suspectthat prospectingadults become aware of the status of a particular site by observing the activitiesof other femalesat the site throughout the season.Most season-long activity would occur at siteshaving active or successfulnests,thus more prospectingwould be expectedat thesesitesif femaleswere able to differentiate boxes receiving muchuse from those receiving little or no use. We didnot observeyearling goldeneyesin our study areauntil the end of May when prospecting was beginning. Unlike adults that had been residing onthe lake all spring, yearlings would have no wayof knowing, except by associationwith adults, ofpast activity indicative of the status of differentnest boxes. However, some other mechanismmight be operating, and we cannot rule out thepossibility that adult females prospectmore thanyearlings and that capture of adults in successfulnest boxes precluded a higher yearling capturerate than we observed.Prospectingdatesand the actual rateswill likely vary among lakes and yearsdepending on geographic location, numbers of breeding females,potential nest sitesavailable, and current nestingsuccess.Perhapsbecauseof a more southerly latitude, prospectingbegan at an earlier date in ourstudy than in British Columbia (Eadie and Gauthier 1985). However, we observed similarmorning and evening peaks in activity. UnlikeEadie and Gauthier (1985) we caught at leastone prospectingfemale that had a brood. We alsoobservedgoldeneyefemalesleave broodsand enter nest boxes (Zicus, unpubl. data). Our datasuggestthat unsuccessfulhens may prospectthemost, but that prospectingby brood hens is morecommon than Eadie and Gauthier (1985) concluded. They reasoned that brood hens had noneed to prospectbecauseof the tendency for successful goldeneye females to subsequently nestin the same site (Johnson 1967, Eriksson 1979,Dow and Fredga 1983). We believe females withbroods would also profit from nest prospectingbecauseof the probable loss of some cavity treesfrom year to year. Prospecting by brood hensmay be less frequent becauseof the need to bal-811ante time spent prospectingwith that spent caring for broods. However, knowledge of the location and suitability of potential nest siteswouldbe an obvious advantage to all females nestingin an environment where the existence or condition of nest sites might change yearly.Unsuccessfulfemalescaughtprospectingin ourstudy had body massesnearly identical to thatof prospectingyearlings and to thosereported forall prospecting females by Eadie and Gauthier(1985). In contrast, incubating females and henswith broodsweighedduring the same period weresignificantly heavier than prospecting females.We do not know if body condition contributedin any way to lossof nestsor whether lower bodymass was the result of increased activity, particularly flight, while prospecting. We suspectthatprospecting may subject females to greater energydemandsthan thosefacedby incubating hensor those rearing broods because many unsuccessfulfemales had body massessimilar to thoseof successfulhens when weighed earlier in theyear (Zicus, unpubl. data).Eadie and Gauthier (1985, p. 533) suggestedthat nest prospectingand delayed maturity weretraits that evolved in relation to the scarcity ofnest sitesusedby North American cavity-nestingducks. Finding suitable cavities is likely a function of: (1) cavity abundance, (2) easewith whichcavitiesare found, and (3) time available to searchfor nests. Besides the Bucephalu species, nestprospecting has been reported in Hooded Mergansers(L. Fredrickson in Bellrose 1976, p. 444,this study), but prevalence of the behavior is unknown. Prospecting by Common Mergansers isalso problematic; Bellrose (1976, p. 444) mentioned it as being “noticed once.” The behaviorhas not been described in Wood Ducks (Aixsponsa)or Black-bellied Whistling-Ducks (Dendrocygnaautumnalis),which are speciesthat nestas yearlings. Prior knowledge of nesting siteswould be an obvious benefit in short, northernnesting seasons.Thus, prospectingmight be morecommon in specieshaving a northern distribution. Common Mergansers (Mergus merganser)have delayed maturity, a more northerly nestingdistribution than Wood Ducks and whistlingducks, and nest both on the ground and in cavities. BecauseCommon Mergansers are not obligate cavity nesters, prospecting may be lessprevalent in this species.In comparison, HoodedMergansers have delayed maturity but a widedistribution. Determination of the extent of

812MICHAEL C. ZICUS AND STEVEN K. HENNESsites by a hole-nesting duck, the Goldeneye Bucephalaclang&a. Ibis 127116-30.EADIE,J. McA., AND G. GAUTHIER. 1985. Prospecting for nest sites by cavity-nesting ducks of thegenusBucephala.Condor 87:528-534.ERIKSSON,M.O.G. 1979. Aspectsof the breedingbiology of the goldeneyeBucephalaclangula. HolACKNOWLEDGMENTSarct. Ecol. 2: 186-l 94.GRENQUIST,P. 1963. Hatching lossesof CommonWe thank personnelof the Chippewa National ForestGoldeneyesin the Finnish archipelago.Proc. XIIIBlackduckDistrict and many private land owners onInt. Omithol. Congr. (1962):685-689.Island Lake for permission towork with their water- JOHNSON,L. L. 1967. The common goldeneyeduckfowl nestboxes.We acknowledgethe help of numerousand the role of nesting boxes in its managementindividuals employed as summer duck bandersby thein North-Central Minnesota. J. Minn. Acad. Sci.Wetland Wildlife Populationsand ResearchGroup for34:110-l 13.weighing brood hens while entanglementtrapping. R. JOHNSON,L. L. 1972. An improved capturetechniqueM. Pace, III, S. J. Maxson, R. E. Lake, J. W. Barrett,for flightlessyoung goldeneyes.J. Wildl. Manage.and J. McA. Eadie made helpful comments on the36 1277-1279.study and manuscript. L. N. Hamilton typed several MENDALL,H. L. 1958. The ring-necked duck in therevisions of the manuscript.northeast. Univ. of Maine Bull. 60. No. 16.MILLIKEN,G. A., AND D. E. JOHNSON. 1984. AnalysisLITERATURE CITEDof messy data. Volume 1: Designed experiments.Lifetime Learning Publications,Wadsworth, LonBELLROSE,F. C. 1976. Ducks, geese, and swans ofdon.North America. StackpoleBooks,Harrisburg,PA.BENNETT,L. J. 1938. The blue-wingedteal, its ecoloav PALMER,R. S. 1976. Handbook of North Americanbirds. Waterfowl. Vol. 3. Yale Univ. Press. Newand management.Collegiate Press,Ames, IA. -IHaven, CT.CARNEY,S. M. 1983. Snecies.aae. and sex identifiPATTERSON,I. J., ANDM. MAKEPEACE.1979. Mutualcation of nearcticgoldeneyesf;o’m wings.J. Wildl.interference during nest prospectingin the ShelManage. 47 754-761.duck, Tadorna tadorna. Anim. Behav. 27:522COOPER,J. A., AND A. D. AFTON. 1981. A multiple535.sensor system for monitoring avian nestina beSowrs, L. K. 1955. Prairie ducks.The StackpoleCo.,havior. Wilson Bull. 93:325-533.Harrisburg, PA and Wildlife Management InstiDow. H., AND S. FREDGA. 1983. Breedine and nataltute, Washington, DC.dispersalof the goldeneye,Bucephala lalangula.J.ZICUS, M. C. 1989. Automatic trap for waterfowlAnim. Ecol. 52:681-695.usingnest boxes. J. Field Omithol. 60: 109-I 11.Dow, H., AND S. FREDGA. 1985. Selection of nestprospecting by both mergansers as well as investigations into the energy costs of all postbreeding activities in prospecting speciesmightfurther advance our understanding of the evolutionary significance of the behavior.

Abstract. We studied nest prospecting by Common Goldeneye (Bucephala clangula) females in north-central Minnesota. Adults unsuccessful in nesting, those with broods, and nonnesting yearlings were captured in nests while prospecting. Prospecting began in late May and continued into early July. Active nests received up to 25 prospecting visits per

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