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This article isprotected rights by All copyright. reserved. 10.1111/ibi.12446 doi: lead tod been and throughtypesetting, proofreadingwhich may thecopyediting, process, pagination This article undergone has for and buthas accepted been not review publication fullpeer Runninghead: Reynolds Jim Editor: Article type Communication :Short Accepted Date :03 Received :17 Date 1 6

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This article isprotected rights by All copyright. reserved. in stored Kazakhstan. of Biodiversity commi The ConservationNature Western Cape Board. the underandconductedpermits issuedby SAFRING Tits Penduline three plastic colour rings ( studied from 2002 T Penduline Eurasian The (42 Kazakhstan Jabagly, near foothills Shan Tien the in 2008 in studied was (33 AfricaSouth WesternCape,Reserve, The METHODS are associated with genetic molecular provide understand Tit Tit Penduline 2008 suggesting consistent is strategy 2005 influence

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2008 approved by the Animal Ethics Committee, University of Cape Town Cape of University Committee, Ethics Animal the by approved

07 at 2008a, across were

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fishpond system is with a numbered metal ring and a unique combination of combination unique a and ring metal numbered a with ) care .

un Thus, (Seutin eeto behaviour desertion populations ° s

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a 19’N, 20 ei coronatus Remiz – et al. et 07 a Keeg Nature Koeberg at 2007) that - crowned PendulineTitcrowned ,

this unusual parental unusual this

greater levels o levels greater 1997 °

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This article isprotected rights by All copyright. reserved. 2014 score likelihood a had they if correctly assigned 2010) Wang & Jones allocat COLONY program and nest. care nestling nest) (12 provisioning recorded was nest the in spent parent each Tit Penduline crowned at recorded lat the during behaviour Tit Penduline noting blood samples obtained from 63 years subsequent in 1 blood Accepted Article 35 –

nestlings 5 as post days 15 White . )

- . sampled birds was 19 was birds sampled

es Thus

the Parentage was assigned usingassigned was Parentage was care Parental

aent ad aent va mxmm ieiod method likelihood maximum a via maternity and paternity

- identity rwe Pnuie Tit Penduline crowned , six

, rates , u c our

1 nss Ersa Pnuie i: 3 nests) 138 Tit: Penduline Eurasian nests, 18 : supported by supported

plus were genotyped

Cape Penduline Tit Penduline Cape birds (32and adults 31 nestlings) -

( hatc an additional eight birdsadditionalan eight were also calculated also were ontinual v 2.0.5 v and behaviour and ) . .

Due to the conservative assignment of COLONY, b COLONY, of assignment conservative the to Due ig a five at hing) o the For nes t e ) , which assesses the likelihood of sibships within the population the within sibships of likelihood the assesses which , recorded r ts ( ts

4 video footage video stages of incubation (8 incubation of stages bevtos f colour of observations

in the in 173

(211 adults and 443 White ±

of nests

ae edln Tit Penduline Cape populations Cape Pendu Cape hog o through

adult birds adult

- minutes of footage per nest) per footage of minutes 17 microsatellitemarkers 17 crowned Penduline Tit Tit Penduline crowned

from videos taken in the latter stages of nestling care nestling of stages latter the in taken videos from ( 299 ,

allowed us to identify the identify to us allowed that

s a as ± srain md every made bservations 44

line Tit line were sampled as nestlingswereasandadultssampled recruited ≥ ( Tbe 1). (Table

Cape Penduline Tit Penduline Cape 80 minutes of footage per nest per footage of minutes esr of measure

and in offspring; van Dijk – - % igd id truhu icbto and incubation throughout birds ringed 12 days after the last egg last the after days 12

(as in (as

population ( population nests

t he Eurasian P Eurasian he aent ws loae uig the using allocated was Paternity .

Bergner

population (317 nuain investment. incubation diinly vdo o parental of videos Additionally, for both for . ±38 The percentage of time that time of percentage The 38 : 21 nests, 21 :

et al. et al. care

(Wang & & (Wang

adults, 1 adults, iue o foae per footage of minutes

two the enduline Tit 654study irds were accepted as accepted were irds ,

provider 2010b 2014 od ape were samples lood Cape PendulineTit Cape as t ah nest each at days )

and

White was laid) were laid) was 3 , 2009 Santure

juveniles andjuveniles Gamero ). nine

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at each at White et al. and - , ,

This article isprotected rights by All copyright. reserved. Acceptedparity from differ not did analyse parentage the Tit Penduline crowned male adult of number nestlings 31 and total A Microsatellite genotyping RESULTS laboratory White ofpopulation entire the in ratio sex equalexpected the from deviations any for test to used was two A (visits/hour). nestlings of rate provisioning the or incubating spent time of proportion the either on sex of effect the for test to test ratio likelihood a using Articleal in term random White the both for approach model mixed linear a using tested were rates feeding and incubation problems the alleviate to model ne multiple term random a as included was (Eurasian, and species father) social the to unrelated offspring of percentage (the nest a in EPP of amount the ‘desert’ c paternal whether test to used was structure error - - crowned Penduline Tit Tit Penduline crowned Tit Penduline crowned of 1 of ) was explained by EPP in our model. The explanatory variables of the model included model the of variables explanatory The model. our in EPP by explained was ) A Bayesian generalized linear mixed mode mixed linear generalized Bayesian A

67 methodology can be found in sting attempts. A ‘normal’ A attempts. sting

( 34 )

l models and the null model (without sex) and the full model were compared were model full the and sex) (without model null the and models l adults, 1 adults, White White s are provided in Table 1. Table in provided are s s :

19 males 19 - - n f and

and sex crowned Penduline Tits Tits Penduline crowned crowned Penduline Tit Tit Penduline crowned in either in 3

juveniles and 12 and juveniles emale and and - typing

: 13 females). 13 : Cape PendulineTit Cape f opee eaain n h dataset. the in separation complete of because Cape Penduline Tit Penduline Cape ae edln Tit Penduline Cape s

- ( prior was applied to the fixed effects when running the running when effects fixed the to applied was prior ae edln Tit Penduline Cape

Supplementa

h sm ml ws cainly bevd across observed occasionally was male same the 0

nestlings)

Details of the microsatellite the of Details The or were genotyped were l (BGLMM, R Package ‘blme’ Package R (BGLMM, l Cape Penduline Tit Penduline Cape

overall ry Online

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females 10 day old day 10 .

details of field and field of details S loci

aibe ‘ae or ‘care’ variable, and 63 (32 adults(32 63 and x ifrne in differences ex

) with binomial with ) roughly genotyped , al exact test exact al and nestlings binomial similar White for -

This article isprotected rights by All copyright. reserved. caring female one and total a nests, a single in resulting contained the Of Parentage analysis Dijk ( Contr with provisioning ( male the (78%) nests 14 at provisioning nestling through continued Tit Penduline the of nest previous a from juvenile a observed provisioning the nest in addition to the focal pair nests observed the In Patterns ofparental care females, test exact Acceptednest 109/138 Article et al. a o vdne for evidence no tnl, n h Ersa Pnuie i nsln cr i dmntd y female by dominated is care nestling Tit Penduline Eurasian the in stingly, 13 ae edln Tit Penduline Cape

social parent was

binomial exact test, (1%)

2010b) , t extra Cape Penduline Tit Penduline Cape wo two three

(Figure 1) (Figure of .

population,

This low This level of s; s; nests h re the - - four until pair youngpair tailed; 79%) .

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nestlings). In these In nestlings). 11%) or the fema the or 11%) sult of sult fledging

with .

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genotyped both the male and female carried female and male the both 0. =

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the remaining nests remaining the pcfc ae roles care specific two nrseii bod parasitism brood intraspecific aetl ae was care parental

78 f offspring of nests where the social mother and father were father and mother social the where nests IBP is IBP is deo footage supported the field observations of observations field the supported footage deo - tailed; )

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nests containenests 11%) At i (29%) six low

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s includingwherenests -

pair young , and biparental care biparental and , were (29/138; 21%) (29/138; n female and across theacross majority

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genotyped White biparental care biparental before . caused by caused 1 mls 13 males: (18

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care two the ,

This article isprotected rights by All copyright. reserved. disentangle thedrivers of this relationship. and diverse with family by provided corre negatively with rates W D of collinearity or EPP from (24% of166nestlings) than that ob However, 2). (Table population Tit Penduline Eurasian the in prevalent more significantly was desertion nests 2 with parents extra of evidence offspring unrelated for caring be will female t unrelated are IBP) from result offspring species bird of ) ISCUSSION e provide evidenceprovide e .

AcceptedWhite Article

a previous studyprevious a of male compared to female care female to compared male of (Table 2). (Table In our our In - crowned Penduline Tit Pendulinecrowned as as

the Eurasian Penduline Tit exhibits substantially higher levels of young resulting young of levels higher substantially exhibits Tit Penduline Eurasian the most ≥ complicates interpretation. B

o the social male per brood per male social the o 97% likelihood at the at likelihood 97%

GLMM lated with EPP with lated T Anl & wn 2002) Owens & (Arnold - he pair offspring was found was offspring pair

c 8% pseie species passerine 84%) (c. most important most

across three closelythreeacross

that demonstrated that the level of male care during chick provisioningis chickduring care male of level the that demonstrated that , EPP atypical parental care strategies care parental atypical . In the In was a was ( (7.1% of 42 and 0% of 29 of 0% and 42 of (7.1% M

Cape Penduline Tit Penduline Cape weak negative weak ø ller & Birkh & ller

variable

f ive

exhibit lower rates of EPP of rates lower exhibit -

nests

; including

w all nestlings (n nestlings all was was served in the mostly biparental as explaining

Ccbr 2006) (Cockburn at which at ead 5.4% predictor of paternal care paternal of predictor species 0.9%

1993) , Tit Penduline Eurasian the in

(across 28 nests) 28 (across the .

In the In

paternal care paternal of penduline tit tit pendulineof

both social parents were known were parents social both

may may , average .

respectively; Figure 2) Figurerespectively; Considering =

29) White provide , . were ihihig hs result this highlighting These results are consistent are results These percent - crowned Penduline Tit Tit Penduline crowned was . future opportunit future

assigned to both social both to assigned

that The that those those that age

across penduline tit penduline across species, as paternal as species, Cape Cape Penduline Tit

b probability

iparental care is care iparental of offspring that offspring ,

the problemthe with higherwith

( .% of 6.6% that a that ies

(Fig n a in

no to .

This article isprotected rights by All copyright. reserved. example Acceptedexception species same the of populations multiple in care parental and EPP assessed have studies few lab, and field the in both research, this of nature intensive the addition Tit Penduline that support populations distinct geographically across single the in system care sampl increased species and populations associated providingwith paternal care. caused Tit Penduline Eurasian observed been Article has behaviour care parental and ratio sex between association an ratio, sex population 2007) (Donald species, bird Dijk van pari showing see (but ratios sex adult unbiased with consistent be would species two these in found care biparental with combined EPP of rates low the adults, amongst ratios Penduline Tit ( species biparental two the in nestlings of ratios sex the in bias a of evidence sex each of opportunities mating betwe and within both system care parental and mating - e cnweg that acknowledge We the influence to predicted variable second the was ratio sex operational or adult The wide , ,

we acknowledge the need for increased sample si sample increased for need the acknowledge we if we were to sample five nests in the Eurasian Penduline Tit population (29/55 nests(29/55 population Tit Penduline Eurasian the in nests five sample to were we if ) (Liker . Nonetheless, our Nonetheless, ty of offspring sex ratio also in the polygamous Eurasian Penduline Tit). Penduline Eurasian polygamous the in also ratio sex offspring of ty and

and inferences aetl ae bevd n u single our in observed care parental populations Hwvr i a ao o brs that birds of taxon a in However, . s f nests of es et al et

Cape Penduline Tit nests seily ml pseie birds passerine small especially .

2013)

because Lt 1991) (Lott

population of population is

(Griffith

. We therefore cannot rule out that bias that out rule cannot therefore We .

typical of typical results highlight a key difference key a highlight results after

hs td would study this xrpltn osrain fo a ige ouain limits population single a from observations extrapolating (Liker ). Although we cannot rule out thepossibility ofskewed sex et al. et ,

ldig ih b rsosbe for responsible be might fledging n because and each

the et al. et

2002 (van Dijk (van

E species

urasian Penduline T Penduline urasian 2014) ) .

oee, e ol age ht h pare the that argue would we However,

facili

benefit . In line with our prediction, we found no found we prediction, our with line In . et al. P etmts nrae n cuay with accuracy in increase estimates EPP t s ifcl t etmt aut e ratios sex adult estimate to difficult is it we ae edln Tit Penduline Cape tate require en species, as it potentially skews the skews potentially it as species, en

2010

zes in zes

from oe cuae siain o the of estimations accurate more ( worth further investigation. further worth a see

) definitive evidence definitive EPP studies EPP .

it is typical of that observed that of typical is it A increase Brommer lthough this lthough ed

adult sex ratios in the in ratios sex adult differing

and d

in general in et al. sample sizes sizes sample White White provides

ins costs fitness t al. et 2010 for - - crowned crowned

In most In

. this

Due to Due 2008b for an for

some ntal For . In of This article isprotected rights by All copyright. reserved. support logistic for Kazakhstan Kazakhstan the conducting support Koeberg at population thank to like would We that could explain therefore may obs care paternal reduced and EPP of levels and p care parental of assortment an confirmed has study Our traits. history species, related d Griffith approach multifaceted a require likely will it tackle to and biology to the evolution of EPP in variation intraspecific rates χ (Pea Tit Penduline Eurasian the in EPP from resulting young contain that the in EPP containing nests with y nests only gaining of likelihood the EPP) from resulting chicks contained 2 oung enduline iffe

Accepted Article1, = df 4.5, = Cape Penduline Tit Penduline Cape rences in mating and parental care among populations within species and among close among and species within populations among care parental and mating in rences between a ws h cs i the in case the was as , D t al. et iversity in iversity t ,

it family that family it as well as well as and to Sergey Sklyarenko at the Association for the Conservation of Biodiversity of Biodiversity of Conservation the for Association the at Sklyarenko Sergey to and aue Reserve Nature

2002

rvd a provide P populations

are

= 0.03). Thus, even low sample sizes can reveal a significant difference significant a reveal can sizes sample low even Thus, 0.03). =

diversity parental behaviour care

, promiscuity likely to generate novel insights into the into insights novel generate to likely ok & enos 2008) Jennions & Kokko

Gert Greef, ESKOM Greef, Gert

exhibit biparental care and high levels of fidelity, in contrast to the high the to contrast in fidelity, of levels high and care biparental exhibit the many field many the n

are

ilwr o the on fieldwork informative . in Or hns o o adr o ad ea oooa o hl and help for Voronova Vera and Bot Sander to go thanks Our . Cape Penduline Tit Penduline Cape oee, e oe future hope we However,

, associated with levels of levels with associated breeding systems the underl the White .

rates Thanks -

rwe Pnuie Tit Penduline crowned remains one of the most puzzling topics in evolutionary in topics puzzling most the of one remains assistants oe sse t ts seii eouinr hypotheses evolutionary specific test to system model ying reasons for reasons ying are and .

erved

also extended also CapeNature .

White more generally (18%) is also significantly lower than the 53% the than lower significantly also is (18%) e ol age ae o or idns that findings our on based argue would We

who collected data on the on data collected who i n the Eurasian Penduline Tit. Tit. Penduline Eurasian the n - rwe Pnuie Tit Penduline crowned

EPP research this variation and how this this how and variation this

for permission for

. The The . to Douglas Ross Douglas to i ol 34. h pooto of proportion The 3.4%. only is , co .

- evolution of these critical life critical these of evolution effort White

Anl & wn 2002 Owens & (Arnold - il focus will crowned Penduline Tit Penduline crowned s rson chi rson

and ytm ars the across systems , Jim Reynolds and Reynolds Jim , Cape Penduline Tit Penduline Cape

permits to work to permits

The ouain in population

- squared test, squared n assessing on within

may relatemay Remizidae in -

pair EPP ly - - ,

This article isprotected rights by All copyright. reserved. Brommer, J.E., Alho,J.S.,Bia Bleeker, M., Kingma, S.A., Szentirmai, I., Székely, T. & Komdeur, J. Birkhead, T.R. & Moller, A.P. Bergner, L.M., Jamieson, I.G. & Robertson, Bateman, A.J. Arnqvist, G. & Nilsson, T. Arnqvist, G., & Kirkpatrick, M. Arnold, K.E. & Owens, I.P.F. R (NBAF361) and (NERC) REvD and ADB. [SBP/JK/2007 Sciences [K109337 PL, and AP, TS, to 00564/2006] support financial further P has results these t hree orme ( rogramme AcceptedEFEREN Article Lubjuhn, T., Patrick, S.C., Rosivall, B., Tinbergen, J.M., der van Velde, M., Oers, van K., Hjernquist, M.B., Kempenaers, B., Komdeur, J., Laaksonen, T., Lehtonen, P.K., clutch desertion in consequences habroptilus relatedness among founders in a genetically depauperate parrot, theKakapo ( in insects. females. passerines: the strength of direct and indirect selection on extrapair copulation behavior in parental care and the risk of retaliation.

ano nymous reviewers CES ] fo te Schure the from , .

performed at the NERC Biomolecular AnalysisBiomolecular NERC the at performed Am. Nat. GEBACO;

1948. Intra Anim. Behav. The m The ). Conserv. Genet. . Academic

eevd udn from funding received olecular

165 - FP6/2002 P 53] to REvD, and REvD, to 53] enduline -

sexual selection in Drosophila. rm h Hungarian the from 2000. The evolution of polyandry: multiple mating and female fitness : S26

for 60

2002. Extra rd, C., Chapman, J.R.,Charmantier, A., Dreiss, A., - Press

ejrnk opnfns f h Ryl Ne Royal the of Poppingfonds Beijerinck 1992. work wasfunded bythe Natural Environmental Research Council : 145 –

providing 2005 S37.

15 - T , London

06 udr otat ubr 28696 number contract under 2006) it, it, : 1013 rm h Hnain National Hungarian the from – Sperm competition in Birds: Evolutionary

164. . The evolution ofinfidelity in socially monogamous Remiz pendulinus -

pair paternity and egg dumping in birds: life history,

from two University of Bath studentships awarded to awarded studentships Bath of University two from comments on the manuscript – , UK 1020.

B.C. Proc. the - .

ot Arcn negvrmna TT [OMFB TÉT Intergovernmental African South

2014. Combining genetic data to identify

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(2nd T it it

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viour – 11. – 250. P

1966.

147 . enduline

J. Ornithol.J. 2008b. Offspring sex ratioin the sequentially polygamous Penduline Tit, . 4

2010

: 66 : 1551 Adaptation and natural selection – Genetics b 77. . Nest desertion is not predicted by cuckoldry in the Eurasian T

– 2009. Parentage and sibship inference from multilocus genotype data it it are not related to breeding densities and mating opportunities.

Szentirmai, I., Székely,T. & Komdeur, J.

1565. Remiz pendulinus 149

P.W.

64 181 : 521

: 1425 1992. Parentage and the evolution of parental behavior. : 1579 - 527. – 1435. –

1594. : the process of clutch desertion. Székely, T. Behav. Ecol.

. Princeton University Press, Princeton

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: 20 2010 – 27.

a Remiz pendulinus

. Parental care strategies in 2008 Á - Constant, R. ., Szabad, J., a . The . The influence of Ibis Trends Evol.Ecol. 149 . : 530

P & & Ardea enduline tal Remiz - 534. , NJ,

This article isprotected rights by All copyright. reserved. Accepted Article Penduline Tit 1 Table Remiz Locus Remiz Remiz Remiz Remiz .

Microsatellite - - - - -

01 14 10 09 07

Penduline Penduline Penduline Penduline Penduline Penduline Penduline Penduline Penduline Penduline Penduline crowned crowned crowned crowned crowned crowned Species White White White White White Cape Cape Cape Cape Cape Tit Tit Tit Tit Tit Tit Tit Tit Tit Tit

- - - - - characterization

# individuals #

G enotyped 168 166 168 148 67 67 67 67 67 -

in the White range (bp) range Allele size Allele 206 169 107 206 187 177 108 113 159 – – – – – – – – – -

214 190 269 223 211 206 137 168 179

- crowned alleles

20 11 3 6 # 9 7 5 9 6 -

Penduline Tit 0.23 0.6 0.29 0.76 0.7 0.4 0.4 0.37 0.8 Ho -

3 8 6 5 1

0.2 0.7 0.75 0.8 0.77 0.48 0.7 0.74 0.7 He and -

3 3 0 9 9

Cape Estimated frequency n ull allele ullallele - - - 0.06 0.442 0.02 0.030 0.26 0.33 0.01 0.005 0.016 -

4 6 9 4 9

This article isprotected rights by All copyright. reserved. Accepted Article Remiz Locus CAM CAM CAM CAM Remiz - - - - 17 15 13 10 - -

17 18

Penduline Penduline Penduline Penduline Penduline Penduline Penduline Penduline Penduline Penduline Penduline Penduline crowned crowned crowned crowned crowned crowned Species White White White White White Cape Cape Cape Cape Cape Cape Tit Tit Tit Tit Tit Tit Tit Tit Tit Tit Tit

- - - - -

# individuals #

G enotyped 173 168 172 172 167 67 67 67 67 - -

range (bp) range Allele size Allele 100 142 203 283 215 213 179 183 83 – – – – – – – – – - - 114

394 164 211 385 221 218 187 194

alleles

24 11 9 6 # 5 7 5 4 7 - -

0.85 0.89 0.7 0.37 0.8 0.76 0.5 0.43 0.68 Ho - -

8 5 1

0. 0.89 0.71 0.3 0.85 0.77 0.50 0.42 0.72 He - - 80

Estimated frequency n ull allele ullallele ------0.045 0.004 0.03 0.034 0.00 0.042 0.00 0.00 0.041 - -

6 5 1 5

This article isprotected rights by All copyright. reserved. Accepted Article CAM Locus TG01 TG01 CAM CAM - - - - - 18 24 20 124 040

Penduline Penduline Penduline Penduline Penduline Penduline Penduline Penduline Penduline Penduline Penduline Penduline crowned crowned crowned crowned crowned crowned crowned Species White White White White White White Cape Cape Cape Cape Cape Tit Tit Tit Tit Tit Tit Tit Tit Tit Tit

------

# individuals #

G enotyped 149 171 167 64 67 66 67 - - - -

range (bp) range Allele size Allele 334 203 396 404 291 198 97 – – – – – – – - - - - 102

340 204 400 406 293 211

alleles

11 4 2 # 2 2 2 3 - - - -

0. 0.46 0.28 0.16 0.35 0.22 0.86 Ho - - - - 50

0.5 0.49 0.3 0.18 0.3 0.3 0.83 He - - - -

4 1 3 5

Estimated frequency n ull allele ullallele - - 0.04 0.02 0.04 0.05 0.208 0.037 0.026 - - - -

1 8 0 9

This article isprotected rights by All copyright. reserved. Accepted Article TG03 Locus TG05 TG05 TG04 TG04 TG04 ------053 046 061 041 012 098

- - - - -

# individuals #

G enotyped 173 171 173 173 171 169 67 67 67 - -

range (bp) range Allele size Allele 137 136 229 233 201 326 188 184 170 – – – – – – – – – - -

145 141 230 234 295 332 196 195 190

alleles

10 5 3 2 2 # 9 3 4 7 - -

0.64 0. 0.58 0.43 0.79 0.5 0.2 0.67 0.7 Ho - - 40

8 7 1

0.6 0.4 0.50 0.49 0.7 0.5 0.2 0.7 0.7 He - -

7 2 8 3 5 7 1

Estimated frequency n ull allele ullallele - - - - 0.010 0.01 0.06 0.06 0.009 0.076 0.013 0.04 0.06 - -

3 5 6 7 5

This article isprotected rights by All copyright. reserved. Acceptedand estimated null allele frequencies were bp, basepairs; Ho Article TG11 Locus TG13 TG12 - - - 017 015 011

Penduline Penduline Penduline Penduline Penduline Penduline Penduline Penduline and crowned crowned crowned crowned crowned Species White White White White Cape Cape Cape Tit Tit Tit Tit Tit Tit Tit Tit

He represent observed and expected heterozygosity

- - - -

# individuals #

G enotyped 170 66 67 67 - - -

calculated in CERVUS v 3.0(Kalinowski range (bp) range Allele size Allele 279 210 231 308 – – – – - - -

281 216 240 320

alleles

2 4 9 # 4 - - -

0.09 0.64 0.88 0.42 Ho - - -

respectively 0.10 0.63 0.8 0.43 He - - -

et al . 2007). Estimated . Ho, frequency n ull allele ullallele - - 0.03 0.004 0.01 0.03 - - -

2 3 5

This article isprotected rights by All copyright. reserved. Accepted Dijk parentssocial ha parasitism nestlings found to be either Fig nest. identity and Fig Figure Article e Fixed included as a random paternal care ( Table 2. ure ure Species ( Species (Eurasian)Species EPP

et al. The Eurasian Penduline Tit data are sourced from

titles 2. 1.

ffect

Parameter estimatesParameter ( E Diversity

2010 (IBP) xtra

s White sex ofthe paren

‘c - b pair paternity (EPP) are

ve are shown for each species ) . -

crowned Penduline Tit ’ of been identified

term nestling

‘ abandon .

within t (s) (s)

provisioning behaviour

and fit statistic ’ provisioning the young post ) (nat nests -

pair young (WPY) and

variation in

t he Eurasian Penduline Tit data are sourced from )

. All data arebased on offspring in where nests both

= s 1.31 - - Estimate

8.19 1.01 74) ofthreep of the

three

, theresult of EPP or Bayesian in p van Dijk

enduline

three penduline tit

1. 2.01 0.92 s.e. - 94 enduline tit species. hatching were

GLMM

et al. t it it species.

(2010b) model 0.67 - - z 4.08 1.

intra 10

recorded at each species The used to predictused

- specific brood

proportion of Male IDwas .

The 0. <0.00 0. P

50 27

van

Percentage of young (%)

0 20 40 60 80 100 Percentage of nests (%)

Percentage of young (%) 0 20 40 60 80 100 Eu

This article isprotected rights by All copyright. reserved. 0 20 40 60 80 100 Figure 2. Figure 1. ra Percentage of nests (%)

Acceptedsi Article

a Percentage of nests (%) n Percentage of young (%) 0 20 40 60 80 100

(n Eu

=1 0 20 40 60 80 100

0 20 40 60 80 100 ra Eu 6

si 6 ra a ) n si

(n a =1 W n

(n h 3 Eu 8 i Eu t =1 ) e ra -cro Eu 6 ra si 6 a ra si ) n w si a

(n n a n W e n =1

W (n

d h (n 3 h

i =1 (n t =1 8 i e t ) e -cro =2 Sp 6 -cro 3 6 8 Pe 9 ) ) e ) w w c n n W n Pe e i e e d d h d W

s i u n (n

t (n h e C d l =1 i -cro i W t =2 u a e n 8 -cro p l h e Pe i ) 9 i e n t w

e )

T e (n -cro w n n

i n T e t =4 Sp d e d i w s t d u

C 2 (n

n e s p (n l ) a e i c p =2 e d n =1 p i e

c e e C (n e c 9 8 s

i a

(n i =1 ) e ) T e p s e =4 s i 8

) (n Sp 2 s =2 C ) p a e 1 Sp e p ) c c e i e e

i C (n e s c a s i =4 p e C

e s

a 2 (n p ) e =2 P

(n a 1 =2 ) r e 1 n I EPP W ) BP t a PY g e

I EPP W BP PY Pa re

Bi Bi U U n n n t p p a i i a a p p l re re a a

I EPP W ca re re BP n n t t PY n n re a a t t l l a a . w l l . . t ma f yp i e t h ma Bi Bi U U . e l h e n n p p

e l i i e a a l p p Bi Bi U U p re re a a e n n p p re re r n n i i a a p p t t n n re re a a a a t t l l re re a a . n n w l l . . t t n n ma f a a i

e t t t l l h a a . ma w . l l l . . h e ma f i e e t l h e ma l p . l h e e e r l e l p e r