Posted on Authorea 9 Feb 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158121185.53650361 | This a preprint and has not been peer reviewed. Data may be preliminary. alu:tenwcs fbodparasitism Honza Marcel brood of cost new the Fallout: ie uvilnei a eicretycnue ihpeain hrfr,w togyrcmeduigvdorcrigfor words: video-recording without Key using Moreover, recommend strongly parasites. we brood Therefore, of predation. birds. cost with in underestimated confused costs of three and incorrectly predation nests. terms the substantial host be estimating in during in can care a restless parents parental it represents more foster insufficient are surveillance rate chicks also by chicks. of video hungry that falling-out cuckoo fed suitability because suggests big high less probably the This chicks, relatively significantly relatively in cuckoo nests. the were of role The the falling-out to nests important in facilitate risk the an stayed may intensity mortality that of plays feeding chicks high out size lower than a nest fell event have remained host that falling-out nests those chicks that before warbler with warbler cuckoo hours suggests reed compared that reed result 29) Eurasian found This great of that we bigger (4 32). and Moreover, rate much of events species falling-out (12 for these host these lower nests warbler of whether nests significantly warbler fate reed establish warbler a reed the To recorded reed video-recorded real we drown. 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Since in Nests 2000), made variation (Hansell builders. 2018). are birds explaining their Kempenaers they of for factors & adaptive success (Santema identify them breeding produced to make the young is on of ecology number effect evolutionary the crucial of in studies i.e. in success, goal major A INTRODUCTION [email protected] E-mail: Correspondence: Republic 1 coevolution nttt fVrert ilg CAS Biology Vertebrate of Institute nttt fVrert ilg,CehAaeyo cecs vta8 0 5 ro Czech Brno, 65, 603 8, Kvetna Sciences, of Academy Czech Biology, Vertebrate of Institute coehlsscirpaceus Acrocephalus 1 iolvCapek Miroslav , 1 alvJelinek Vaclav , , uuu canorus, Cuculus uuu canorus Cuculus 1 1 n ihlSulc Michal and , ro aaiim hc uvvl nest, survival, chick parasitism, brood ,oeo h otsuidbodparasite brood studied most the of one ), 1 ø ta.21) About 2012). al. et e ø tal. et e Posted on Authorea 9 Feb 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158121185.53650361 | This a preprint and has not been peer reviewed. Data may be preliminary. iitr iia iercre iiDRC-D05(hnhnCuTc o t,Cia lcdi a in placed China) and Ltd, illumination IR Co. without Chu-Tech recorders (Shenzhen inbuilt CH-HD0065 We with DVR cameras 2019). 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Czech Moravia, South in 17deg02’E) 54’N, 2 ø ta.21) hs otseisdiffer species host These 2012). al. et e yh angustifolia Typha .Ec etwas nest Each ). Acrocephalus Phoenicurus Phragmites ø e Posted on Authorea 9 Feb 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158121185.53650361 | This a preprint and has not been peer reviewed. Data may be preliminary. fpriua oe em eebsdo yeIIsmo squares. of sum the a III from as Type out visits P-values on fell feeding covariates. chick based parental R as were of = (categorical) in number terms (0 year variable, function fate model and binomial (continual) link chick particular response date cuckoo logit of a (continual), the and as daytime of fledged) and distribution consisted of predictor successfully error continuous chick out model binomial = fall initial 1 a will The or chick with 2018). nest cuckoo analysis. Team models that main Core linear probability this (R the generalized for 3.4.4 and used between used effort we were differed fledged feeding nest, nest successfully parental chicks the the between cuckoos cuckoo of relationship which of the out from test fell success nests To prematurely fledging Only or nests. the nest GRW the whether experimental left determine larger and and to nests test RW exact smaller Fisher’s and used We age count to nestling sufficient both not was on analysis recording Statistical of depend quality could the One unsuccessful parents because nests. assigned visits. analysis unsuccessful of in feeding the 15 those parental behaviour from and excluded in successful the was 15 as nest because of age unsuccessful comprised time same dataset the final same The of (Poˇzgayov´a the 2015). daytime were al. from chicks et cuckoo nest difference and same the mean nests which the (the had in season season, video-recordings breeding the from cuckoo same during nest the the where one timing from nest selected we similar ± each were we for nests nest, and counterpart falling-out, successful falling-out. control treatment the of These a chick of size they frequency as cuckoo (unsuccessful). affect falling-out (successful) where out the nest may of nests the fell parents preceding time in chick with foster span stayed the chick of out time cuckoo to effort the three-hour fell feeding closely where compared the chicks the we state during cuckoo whether hunger, cuckoo visits cuckoo test which on To of feeding from focused level parental nests the were all by in we counted affected parents As is foster nest successfully. the of fledged of behaviour hosts. falling-out feeding of of skills probability the building the above). the how (see not test nests To RW and non-manipulated nests feeding 42 of reed parental as features to Natural frequency of spatial attachment same their the Analysis site. the experimental fixed on nest we with the focused the checked or of were left were fastened effect we we nests loosely negative as Experimental after those any string avoided immediately notice jute we were almost by not the but chicks cuckoos stems did manipulated, cuckoo as feed We not and place to were installed same ones. started nests was the new RW parents video-camera at these foster the stems to when as reed nest day procedure ( the original the to ones the on often attached new strings GRWs from were build moved jute Since nests then with site. These nests study and RW same experiment. parasitism original the our to from for response obtained nests a were abandoned more as experiment are nests this they their for nests. and abandon used GRW 2011) nests GRW diameter with Schulze-Hagen 51.6 data). nest & nests cup: own RW (mean nest (Leisler of (Koˇzen´a-Touˇskov´a nests habitat 33 diameter RW composition same mean replaced GRW, material than the we and larger in survival, design one-third occur chick similar 1973; than 2010), a cuckoo Pearson have affects & they size Kennerley because nest nests GRW host used how We out to nest find the cm To check 20 card). to days from memory three varied experiment the every (change replacement nest visited data Nest and were stems the nests cameras reed download cloths Filmed between cut and masked density. distance by equipment stem by camouflaged The adjust reed covered poles content, the m and aluminium string. on 10 reeds thin jute depending ca on on by meters bank placed two hung together were record the were fastened Cameras to on recorders leaves capacity nests. placed with the and from were boxes with metres batteries resistant batteries five camouflaged; water to gel was two Ah and equipment 100 nests All / the days. 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Data may be preliminary. ta.93 ikn ta 08.I utb oe,ta h ukociki asdaoeadde o have not does and alone raised is chick cuckoo the that noted, (McRae movement be active has must Price more it 1991; to It related Indeed, Montgomerie 2008). positively events. & al is fall-out (Smith begging et accidental hungrier their Dickens the are and restless al.1993; 2002) to they are Leonard et when lead they & intensively Wright can thus 1996; more which and al. beg nests hungrier et nestlings are their chicks that chicks in well-fed satiated shown vigorously less less been more that these i.e. beg that hunger, may with speculate and We increases frequently. positively however, more falling-out hypothesis, of out probability This fall the chicks. nests that parasitic bigger found for also raising We preference for Molina- the better suitability 2004; that are physical suggest al. nests testing. better we bigger further et However, their larger built requires better. (Kleven by that in chick parasites males) explained parasitic eggs and/or be brood the their (females may feed of pairs lay also host choice to can that prefer nest and suggested quality parasites the 2016) 2013, why observed al. for that et explanation Morales studies also alternative Previous but of an nests nests. moment provide RW age). host in the also of than may till days frequently result with even 15 less solid This only often (after more not - period GRW also nestling longer in are the much happened nests accommodated events in GRW damage falling-out comfortably later Moreover, fundamental result which nests. without a RW cups As endured the nest fledging. thus than on spacious time and standing more longer easily walls When have and a thicker remnant. moving nests for nest or period. GRW chicks stretching nestling trampled during cuckoo larger their a balance growing contrast, of losing on thirds In to only two susceptible down. sat very or were fell and half chicks first nest cuckoo platform the the flat during are researchers. damaged this in nests the frequently RW fitted by chicks happened design. only check events the nest chicks nest insufficient Later cuckoo falling-out by last which researchers all caused the cups was by that after deep falling-out induced small found hours that several we prematurely show and video-recordings as was fledging our unlikely Interestingly, it of events is and/or time falling-out explanation expected fledging the nests this before that of the However, days speculate under time several 2012). can found with al. One chicks reasons events. associated dead et unknown falling-out that to (Pietz be of believe due might consequences we dead video-recordings, recorded are found “In our (2004) we (2004): were of al. al. 10 light et et depredated, success nests In Kleven Kleven were RW breeding by nest“. by the in nestlings outlined the on cuckoo mystery raised under chicks effect clarify the or negative cuckoo to of inside apparently helps of 12 has also 32) nests that finding of warbler This cost (12 RW. reed serious by third a raised one represents chicks than This cuckoo more of nest. their that the found than of have out event we fell falling-out study the present the before In hours 2). three (Fig. the fledged which during finally chicks intensively they Cuckoos DISCUSSION which 1). less (Tab. from fed nest nests were the in of nest counterparts falling-out cuckoo the cuckoo cup. of of nest probability (N=2) out the the behaviour cleaned fell and comfort parent visits to foster feeding parental a due (estimate In of relationship when cup moving. negative the side or deep significant stretching the when a a during found to with (N=7) We rim moved chicks nest nest excessively undamaged the the it an from of because slipped from behaviour once or out comfort and stem fell down, reed caused cuckoos chick the fell was the occasions around falling-out and fed three wing typically the backwards by its parents cases, trampled or when hooked these was forwards (N=5) e.g. In nest chick feeding = leaned side. the parental P one it cases by: test, to 16 then triggered (Wilcoxon tilted chicks of nest nests and sometimes GRW cuckoo 13 the platform in Moreover, in on flat movement raised nest, 0.06–1.08). excessive those less the = by than or from CI age more falling-out younger 0.27, to cuckoos. were a chick chicks = adult at the cuckoo by ratio nests When ejected the odds 1). larger were of 0.044, Fig. from out %) than 0.024, = fell 32) (3 P nests of chicks RW 29, (12 two in nests of %) and RW raised natural (4 (14 nests smaller chicks nests the from GRW 10 frequently of successfully, experimental more out fledged significantly fell out %) fell %) (61 chicks (22 chicks Cuckoo chicks 45 chicks, 16 cuckoo depredated, video-recorded were continuously 73 the Of RESULTS 4 ± E=–5.62 = SE ± 18,P=007 ewe h number the between 0.047) = P 11.84, Posted on Authorea 9 Feb 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158121185.53650361 | This a preprint and has not been peer reviewed. Data may be preliminary. otdb h zc cec onain(rjc1-26S n yteIsiuinlRsac ln(RVO: Plan Research Institutional the AUTHORSHIP sup- by was and study (project17-12262S) JMK20189/2010, Foundation numbers The Science permit conserva- 68081766). 34437/2014). Czech (Czech nature animals the MUHOCJ local protected by to JMK115874/2013, wild ported obliged on 41433/2012, are work We MUHOCJ to guidelines. us JMK23530/2011, ethical allowing with for compliance authorities in tion conducted for was can research advantageous research. ground The future become the for may above avenues vegetation fruitful nests Finally, are in in systems. areas Acknowledgements events concealment parasitic these falling-out that because brood accidental believe fitness different We whether survival. across parasite study nestling widespread brood to more affect interesting is be positively it also whether would or Americas it the species-specific rate in is those rejection cost like low systems or ( the brood-parasitic densities blackbird in different ( breeding winged be on host cowbirds can high needed insufficient falling-out predation, by are of the nest studies represented cost that Further low both This eggs. from costs that factors. parasitic gains serious contributing demonstrate of by represents be We counterbalanced may study parasite. time feeding this the parental same in insufficient low of and observed by success size chicks compensated breeding nest cuckoo be the of can reduce falling-out nests RW substantially frequent 1993). from Martin the 1969; fall-out conclude, (Ricklefs the To failure habitat. by nest this induced of 2005;, in cuckoos source (Weidinger Gosler rate for main habitats & predation the cost woodland (Neto nest is the or predation reedbeds survival that open nest in Nevertheless, in suggests where nest). breeding reason breeding This 2014) the species main species al. in other with the et died of comparison was Praus one in nests fall-out 2006; and high the or predated falling-out are nests 2016) were 2014) and video-monitored al. 12 al. (Jel´ınek overestimated Musilov´a 74 et et out, nests been our RW fell has in chicks of predation fact, cuckoo rates that In (16 well. findings failure as chick new nest cuckoo effect for our the strong of of RW a light disappearance the have considered in by might authors raised suspect these cuckoos study, we Although of this particular, predation, 2016). success to In al. Jel´ınekas breeding Similarly et 1998; 2000). and/or conclusions. fledgling Granfors al. and low & et relatively results (Honza factors (Pietz a false of common reported to have misclassification relatively lead studies the is potentially nest previous can that eggs/nestlings its rate revealed of defend predation method loss of not This or ( overestimation in failure does rate nests. nest high bird 2016), host to a of al. cuckoo leading surveillance (Jel´ınek at et frequent video eggs area advocate most cuckoo small also accepts the relatively We or a is 2004) on RW in al. Therefore, specialization densities the et cases. that high (Honza this, most suggesting aggressively in Despite in cost, breeds trap. high 16) it evolutionary a of because R have an (7 drowned & can out become territory (Moksnes water fell can the Europe that the from host chicks above cuckoo trace a nest the without such RW, that the disappeared species In prematurely host habitat. knowledge or parasitizing the in 16) on gap depending of apparent differs an (4 falling-out is events. of there these Interestingly, cost for 2009). The or triggers Bednarz 1996) as al. & act et (Bader that (Kleindorfer nests factors from their jump of of 2009), Bednarz out in & pushed observed Bader are been 1999; Morrison also 1955; has Vietinghoff-Riesch age (von fledging ( ( reached kite swallow they Mississippi barn before like nests species frequency. feed therefore their other feeding 2005), to leaving many Nowicki lower likely nestlings & with more of (Searcy associated are phenomenon need parents be The short-term where may signals events position reliably falling-out a begging that to Yet, believe moving 1993). we and al. jockeying et by (McRae siblings them other with compete to iud rustica Hirundo glisphoeniceus Agelaius ø kf 95 hc rsmbyipista hshs sohrievr utbe e.g. suitable, very otherwise is host this that implies presumably which 1995) skaft Molothrus n osahdwrlr( warbler moustached and ) p. n hi ot.Ahs reigoe ae uha h red- the as such water over breeding host A hosts. their and spp.) ) , cii mississippiensis Ictinia ol eapriual utbemdlt xmn hte this whether examine to model suitable particularly a be would 5 coehlsmelanopogon Acrocephalus ,nrhr aaaa( caracara northern ), uce l 2019). al. et Sulc ˇ aaaacheriway Caracara ,we hyfl out fall they when ), ), Posted on Authorea 9 Feb 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158121185.53650361 | This a preprint and has not been peer reviewed. Data may be preliminary. lidre,S,Hi .&FslB 19) lr al n hc ecin ntemutce warbler, moustached the in reactions chick and R calls ( A., Moksnes, Alarm O., (1996). Kleven, B. Fessl London. Black, & Helm/A&C H. melanopogon Christopher Acrocephalus Warblers. Hoi, Bush S., and Reed Kleindorfer, (2010). 1975–1987. D. 70, Pearson Sociobiol., & Ecol. P. Behav. Kennerley nestlings? host suffer London. than chicks Helm, cuckoo more common Christopher predation Do Icterids. nest (2016). The M. Blackbirds: Honza, Proch´azka, World & K., P. New Karasov´a, Weidinger, Jel´ınek, V., T., 1999. P. Burke 104–108. & 45: A. Study, Jaramillo, Bird position. nest to relation in M, WarblersHonza R Reed & parasitized A. experimentally 256–263. Moksnes, of 51, Jr, Study, behaviour M., inspection Cambridge. Capek, nest Press, T., and University Cambridge Grim, behaviour. M., construction 10.1111/j.1365-2656.2010.01798.x Honza, and doi: nests Ecol. Bird Anim. (2000). J. M. hosts? parasitic Hansell do potential why common choice: host some on exploit Constraints for (2011). rarely M. Costly birds Honza, is & Virulence O. Samaˇs, Mosk´at, Kleven, Constrained T., P., C., Experimentally Grim, 14–22. (2009). 115, M.E. Ethology, Hauber, Chicks. & Cuckoo P. Cassey, Common ( J., blackbird Rutila, red-winged T., Grim, of Selection (2002). : 1023–1029. ( S.G. cowbird 86, mocking- doi Sealy, by brown-headed Behav. mobbing at: Anim. & violence: Available cowbirds. N.D. of brood-parasitic wages (v10.1). Grant, The against List (2013). defence A. frontline Bird Kacelnik, a World & as J.C. IOC birds Reboreda, V.D., (2020). Fiorini, R., (eds.). Gloag, P. 2020. 02. Rasmussen, 04. & accessed Last D. 10.14344/IOC.ML.10.1., Donsker, F., Gill, London. hungry Plc, strategies: Publishing begging 167-174. Helm/Bloomsbury 75, offspring Behav., and Anim. care father. Erritz Biparental the (2008). towards move I.R. nests? tits Hartley, Blackbird blue & nestling Red-winged D. locate Berridge, to M., Dickens, use Cowbirds Brown-headed 1181–1189. do errors. 55, recognition Behav., cues make Anim. What to (1998). it Common induce E.D. not the Clotfelter, do of but Warbler presentations Reed 763–769. Eurasian Repeated 112, the kites: (2010). Condor, by Mississippi defence M. for nest Honza, failures increase Cuckoo nest & Poˇzgayov´a, 598–606. of Proch´azka, P. 29, M., causes M., Wetlands, and Capek, Arkansas? success eastern Reproductive in population (2009). sink J.C. A Bednarz, & T.J. Cuckoo Bader, common Chat nestling Bush the of Rufous gape for conspicuous stimulus The (2004). F. Alvarez, References manuscript. M and the MH wrote study, the designed MH uuu canorus Cuculus ø ,J,Mn,CF,Bamr .. ulr ..(02.Ccoso h ol.Christopher World. the of Cuckoos (2012). R.A. Fuller, F.P., Brammer, C.F., Mann, J., e, Ø e J ose .&RsatE 98 uvvlo edWrlrArcpau craesclutches scirpaceus Acrocephalus Warbler Reed of Survival 1998. E. Roskaft & A. Moksnes IJ, ien hcs ea.Eo.Scoil,4,41–46. 47, Sociobiol., Ecol. Behav. chicks. ) oohu ater Molothrus nm ea. 1 1199-1206. 51, Behav., Anim. . ø kf,E oz,M 19) otseisaet h rwhrt fcuckoo of rate growth the affects species Host (1999). M. Honza, & E. skaft, ectihsgalactotes Cercotrichas olce h aa CaV nlsdtedt,M,M,V n M and VJ MC, MH, data, the analysed VJ a MC data, the collected S ˇ .Br ea.1,21-30(10) 15, Behav. Bird ). 6 ot.Ada9,63–68. 92, Ardea hosts. ø kf,E 20) etdfne nm recognition enemy defence, Nest (2004). E. skaft, glisphoeniceus Agelaius uuu canorus Cuculus coehlsscirpaceus Acrocephalus sasupernormal a as ot ythe by hosts ) Bird . S ˇ Posted on Authorea 9 Feb 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158121185.53650361 | This a preprint and has not been peer reviewed. Data may be preliminary. adokn eae?Sxa oflc vrprna aei ro aaiehs n t osqecsfor consequences its and host parasite brood 1053–1061. a 69, in Sociobiol., care Ecol. parental Behav. and over growth. males chick conflict Lazy Sexual (2015). females? M. hardworking 34–38. Honza 81, & Process., M.M. Behav. Abraham, Poˇzgayov´a, Jel´ınek, incubation. Proch´azka, V., P., parasitism Beˇno, female-only R., M., brood with against host behaviour a defence in Sex-specific (2009). M. Poˇzgayov´a, Honza, & Proch´azka, P. M., cuckoos Pietz, common & hosts: 879–891. suitable III. Choosing Grassland canorus F.R., (2009). Thompson, Cuculus Video-Monitoring M. C.A., from Honza, Ribic, & 3-22. Gained (eds. Polaˇcikov´a, pp M.I. Berkeley, using Knowledge Cherry, Birds. Proch´azka, P., Press, Nesting L., nests (2012). California of passerine of C.A. Surveillance University grassland P.J.). Ribic, Video of In: & fates Nests. D.A. Passerine and Granfors, predators 71-87. 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