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Reproductive Success and Nest-Site Selection in a Cooperative Breeder: Effect of Experience and a Direct Benefit of Helping

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Reproductive Success and Nest-Site Selection in a Cooperative Breeder: Effect of Experience and a Direct Benefit of Helping TheAuk 116(2):355-363, 1999 REPRODUCTIVE SUCCESS AND NEST-SITE SELECTION IN A COOPERATIVE BREEDER: EFFECT OF EXPERIENCE AND A DIRECT BENEFIT OF HELPING B. J. HATCHWELL,• A. E RUSSELL,M. K. FOWLIE,AND D. J. Ross Departmentof Animal and Plant Sciences, University of Sheffield,Sheffield S10 2TN, UnitedKingdom ABSTRACT.--Wedetermined whether nest-site characteristics influence reproductive suc- cessand whetherexperience influences nest-site selection in a populationof cooperatively breedingLong-tailed Tits (Aegithaloscaudatus). Nest predationwas high; only 17%of breed- ing attemptsresulted in fledgedyoung. The heightof nestswas an importantdeterminant of success;low nestswere significantlymore successfulthan high nests.A breeder'sage, natal nestsite, and breedingexperience had no significanteffect on nest-siteselection. How- ever,failed breeders who helped at thesuccessful nests of conspecificsbuilt subsequentnests lowerthan nestsbuilt prior to their helpingexperience. Failed breeders who did not help showedno reductionin tlseheight of subsequentnests. Moreover, the subsequentrepro- ductivesuccess of failed breederswho helped was significantlyhigher than that of failed breederswho did not help.We concludethat helpersgain informationon nest-sitequality throughtheir helping experience and thus gain a directfitness benefit from their cooperative behavior.We suggest that experience as a helperoffers a morereliable cue to nest-sitequality thanbreeding experience because helpers are associatedwith nestsonly during the nestling phasewhen few nestsare depredated.In contrast,although successful breeders may expe- riencesuccess with a low nest,they are evenmore likely to haveexperienced the failureof low nestsbecause of the high rate of nestpredation. Received 26 December1997,accepted 28 July1998. A MAJORDETERMINANT of reproductivesuc- acteristics,then no consistentselection may ex- cessfor manyorganisms is the abilityof breed- ist for choiceof particularnest sites. However, ers to protecttheir offspringfrom predation. if breeding successis consistentlyrelated to Thisis particularlytrue of manyopen-nesting certainnest traits, then high ratesof nestpre- passerineswhere the rate of nestpredation may dation will exert strong directional selection be extremelyhigh (Lack 1954,Ricklefs 1969, againstthe choiceof low-qualitysites by breed- Martin 1995). A variety of antipredatorstrate- ers, resultingin low variancein the critical gieshas evolvedamong birds to reducenest characteristics.Alternatively, in the absenceof predation, including colonial or dispersed consistentselection for particular nest sites, breeding,use of cavities,nest concealmentor site choicemay be a variabletrait, with adap- camouflage,elaborate nest design, and protec- tive nest-sitechoice occurring either through a tive nesting associations(see Collias and Col- learningprocess of trial and error, or as a con- lias 1984).Numerous studies have shown an in- ditionalbehavior depending on predationrisk traspecificrelationship between nesting suc- or habitattype. cess and various characteristics of nests or nest Individual breeding performance can in- sites, e.g. nest structure (Baeyens1981), nest creasewith age and experiencein many ver- density (Anderssonand Wiklund 1978,Potts et tebrates(see Clutton-Brock 1988), and this has al. 1980, Hatchwell 1991, Chamberlain et al. often been attributedto increasedforaging ef- 1995, Meilvang et al. 1997), and nest conspic- ficiency(e.g. Desrochers 1992a, b) or increased uousness(Picman et al. 1993, Hatchwell et al. reproductiveeffort; i.e. the "constraint"and 1996),although other studieshave shownno "restraint"hypotheses of Curio (1983).Indi- sucheffect (e.g. Holway 1991, Colwell 1992, Fil- vidualsmay acquireforaging skills at anytime, liater et al. 1994,Cresswell 1997a). If predation but more specializedparenting skills, suchas risk is randomwith respectto nest-sitechar- nest placementor incubation,may only be ac- quiredthrough breeding. Such experience may be gainedthrough personal reproduction (e.g. E-mail: [email protected] Marzluff 1988, Marzluff and Balda 1992) or 355 356 HATCHWELLET AL. [Auk, Vol. 116 throughobservation of others(e.g. Boulinier et we investigatewhether and how individuals al. 1996), but among cooperativebreeders it acquire information about nest-sitequality. In also may be acquiredby helping a breeding particular,we examinethe hypothesisthat ex- pair rear its offspring(Brown 1987). Few stud- periencegained through helping at the nestof ieshave shown a directbenefit to helpersof in- conspecificsinfluences subsequent selection of creasedexperience, although notable excep- nest sites. tionsexist. In White-wingedChoughs (Corcor- ax rnelanorharnphos),helpers acquire foraging METHODS skillsduring their long prebreeding association with cooperativegroups (Heinsohn et al. 1988), We studieda populationof 15 to 35 pairsof Long- and in SeychellesWarblers (Acrocephalus sechel- tailed Tits from 1994 to 1997 in the Rivelin Valley, lensis),both helpersand breedersgain experi- Sheffield,United Kingdom. The study site (ca. 3 km2) compriseda variety of habitatsincluding hedge- ence in nest building, nest guarding, and in- rows, scrub, mature deciduous woodland, and small cubation (Komdeur 1996). standsof coniferoustrees. The breedingattempts of In this study,we used the cooperativebreed- individually markedbirds were closelymonitored ing systemof the Long-tailedTit (Aegithalosthroughout each breeding season(March to June). caudatus)to investigatethe effects of nestsite on We found a total of 178 nestsbelonging to 67 males reproductivesuccess and the effectsof experi- and 68 females.Each individual wasrepresented by ence on nest-siteselection. Long-tailed Tits betweenone and eightnests in our dataset. In some build domednests in very diversesites and suf- analyses,this raises the potentialdifficulty of pseu- doreplication,but we consideredeach nest to be in- fer high ratesof failure,largely caused by pre- dependentin analysesof reproductivesuccess with dation (Lack and Lack 1958,Gaston 1973, Glen respectto site characteristics.This is justified be- and Perrins1988). A new nestis built for every causealthough some nestsbelonged to the same breedingattempt, and a pair builds between birds, each was located in a different site, and it is oneand four nests in a singleseason. Each nest nest location that is important for this question constitutesa large reproductiveinvestment be- (Hatchwell et al. 1996, Cresswell1997b). Further- causethey are amongthe mostelaborate struc- more, nests of the same individual often encom- tures of any Europeanbird speciesand require passedthe wholerange of possiblenest sites, and the a lengthybut variablebuilding period (the av- same individuals did not build successive nests at the sameheight (males, F = 1.33,df = 38 and 97, r eragebuilding period for firstnests of the sea- = 0.086; females, F = 2.28, df = 36 and 81, r = 0.294). sonwas 25 days,although later nests were built In analysesconcerning experience and nestsites, we in just 8 days;B. J.Hatchwell unpubl. data). The used individuals as independentdata. Divorce was nestis made of mossbound with spider'ssilk, frequent both within and between seasons(B. J. the exterior is covered by several thousand Hatchwellunpubl. data), so we treatedthe sexes sep- piecesof lichen,and the interioris lined with aratelyin analysesof nest-siteselection. an averageof about 1,500 feathers(Hansell The greatmajority of nestswas found during the 1993,B. J.Hatchwell unpubl. data). The sexra- early stagesof building.Although we madepartic- tio is 1:1in the cooperativebreeding system of ular efforts to find replacementnests following nest failure, a few nests were never found; this was usu- Long-tailed Tits, and all birds attempt to re- ally attributableto renestingof failed pairs outside producein pairs.If their ownbreeding attempt the studyarea and sowas unlikely to biasthe sam- fails,however, breeders may become helpers at pling of nests.Nests were visited every one to three the nestsof their relatives,assisting in the care days during building, laying, and incubation and of nestlingsand fledglings (Lack and Lack were observedevery two days during the nestling 1958, Gaston 1973, Glen and Perrins 1988, periodto recordthe presence and identity of helpers. Hatchwell and Russell 1996). Thus, helpers Frequentnest visits might elevatenest predation maygain information about what constitutes a rates if potential predatorsobserve such visits or if successfulnest site through their own breeding visits reducenest concealment(Mayfield 1975, Len- experienceand/or by helpinganother pair. ington 1979).We do not considerthat an observeref- fect on nest predationbiased our resultsin any way First, we investigatewhether the high pre- for three reasons.First, depredatednests were usu- dation rate of Long-tailedTit nestsis influ- ally torn apart (see below), creating a "drift" of encedby characteristicsof the nest site. We feathersfrom the lining, so the survivalof the great show that nestplacement plays an important majorityof nestscould be checkedfrom a distanceof role in determiningbreeding success. Second, several meters. Nest contents of accessible nests were April 1999] Experienceand Nest-site Selection 357 checkedonly to confirm the start of laying, clutch 50 size,hatching, and during bandingof nestlings.Sec- ond, mostof the nestsplaced high in treeswere rel- 40' H [] Failed atively inaccessibleand so were visited only if they survived to day 11 of the nestling period, when
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