Widespread Gene Flow Between Oceans in a Pelagic Seabird Species Complex

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Article: Booth Jones, K.A. orcid.org/0000-0001-5228-8454, Nicoll, M.A.C., Raisin, C. et al. (10 more authors) (2017) Widespread gene flow between oceans in a pelagic seabird species complex. Molecular Ecology, 26 (20). pp. 5716-5728. ISSN 0962-1083 https://doi.org/10.1111/mec.14330

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[email protected] https://eprints.whiterose.ac.uk/ Sweet 7 6 U.S.A. York, Street,New West at79th 5 4 UK. 7NR, KentCT2 Canterbury, Building, Marlowe Kent, of 3 UK. Sheffield, Bank, Western Sheffield, 2 1 addresses: Author This article isThis article by protected copyright. All rights reserved. 10.1111/mec.14330 doi: differences between to lead the of version citethisarticle this and as Record.Please Version copyediting, pagination the been through andproofreadingtypesetting, may process, which This article acceptedhas been forpublication andundergone full peer review buthasnot 0000− Ι : (Ορχιδ ϑΟΝΕΣ ΒΟΟΤΗ ΑΛΙΧΕ Ρ. ΚΑΤΗΕΡΙΝΕ Dawson Authors: : 0000− Ι (Ορχιδ ΜΗ ΙΣΜΑΡ ΣΤΕΦΑΝΙΕ ΠΡΟΦ. Title: Αρτιχλε Οριγιναλ : τψπε Αρτιχλε Accepted U.K. Islands, Jersey, Channel Trinity, Les Manor, Trust, Augrès Wildlife Conservation Durrell Mauritius. Vacoas, Road, Grannum Foundation, Wildlife Mauritian Park Central Natural History, of Museum American Ornithology, - of Vertebrate Zoology Division NewZealand. Auckland, 1142 Auckland, of Sciences,University Biological of School University andConservation, Anthropology School of Ecology, and Conservation of Institute Durrell of University Sciences, Plant and Animal of Department Facility, Analysis NERC Biomolecular U.K. London, London, Society Zoological Zoology, of Institute Article 5 , Carl G. Jones , Carl Widespread gene in flow seabirdspecies oceans a pelagic between complex 2 , Helen Hipperson , Helen

Ka therine A. Booth Jones A.Booth therine

6.7 , Vikash Tatayah 2 , Gavin J. Horsburgh J. , Gavin 1,2,† , Nicoll A.C. Malcolm 6 , Kevin Ruhomaun , Kevin 0001 2 , JimJ. Groombridge −7437− 0001 3393) − 5228 8 and and Ken Norris 1

, Claire Raisin − 8454) 3

, Stefanie M.H. Ismar , Stefanie M.H. 2,3,‡ 1 . . , Deborah A. , Deborah

4, § , Paul Previous research assumed that Round Island represents a point of secondary contact between Atlantic Atlantic contactbetween of secondary apoint Island represents Round that assumed research Previous Ocean. intheIndian Island, Round on of petrels exists population hybridising amixed-species, of example only well studied The inseabirds. particularly isunderstudied, betweenspecies introgression flow forand gene potential the asinglespecies, of populations flowbetween different gene the on studies focus many Although populations. species and in diversity genetic decrease or increase either This article isThis article by protected copyright. All rights reserved. † Currentaddresses: 8 Abstract title: Running U.K. Email:[email protected] London, author: Corresponding words Key Germany. § ‡ British Trust for Ornithology, The Nunnery, Thetford, IP24 2PU, 2PU, IP24 Thetford, TheNunnery, Ornithology, Trust for British Mauritius. Reduit, Mauritius), of Service (Government Conservation and Parks National GEOMAR, Helmholtz Zentrum für Ozeanforschung Kiel, Düsternbrooker Weg 20, 24105 Kiel, 24105 20, Weg Düsternbrooker Kiel, Ozeanforschung Helmholtz für Zentrum GEOMAR, Accepted Article UK Chester, Westand Cheshire Chester, Chester Zoo, Global-scale gene flow is an important concern in conservation biology as it has the potential to has the asit potential biology conservation in concern important isan flow gene Global-scale

: dispersal, gene flow, hybridisation, petrel, Pterodroma, tracking. tracking. petrel, Pterodroma, hybridisation, flow, gene dispersal, : Gene flow in a seabird species complex species complex aseabird in Gene flow

Malcolm Nicoll. Address: Institute of Zoology, Zoological Society London, London, Society Zoological Zoology, of Institute Address: Nicoll. Malcolm

UK.

This article isThis article by protected copyright. All rights reserved. species between gene-flow revealed analysis STRUCTURE Conversely, Ocean. Indian the Pacificinto and Atlantic movement from the petrels of revealed unidirectional BAYESASS rates estimated migration using of Analysis complex. this in species between the demonstrated three oceanswas spanning flow gene and Dispersal Ocean. Indian outside the occurring hybridisation between-species of possibility the anddata address tracking to genotyping microsatellite uses study This heraldica). P. and neglecta Pacific species(P. and arminjoniana) (Pterodroma Introduction Introduction threatened seabirds. of the in conservation hybridisation migrationand widespread of consideration for the need demonstrating scale, a global on Procellariform species related ofclosely populations between flow to investigate gene kind its of isthe first study This assumed. was previously more are petrels common than Pterodroma in hybrids inter-specific that suggest results These Pacific. Atlantic and the to travelling individuals two Island petrels revealed Round of tracking geolocation Ocean.Additionally, Indian the outside hybrids occurring three-way potential with Oceans, important to the persistence of threatened populations (Bicknell et al. 2012; Blower et al. 2012). 2012). et al. Blower 2012; et al. (Bicknell populations threatened of the persistence to important tobe has shown been connectivity population 2003). Moreover, et al. (Grant depression inbreeding affectedby populations small of fitness genetic canboost the units between discrete flow gene of possibility the maintaining therefore sources biodiversity, unique of represent may populations Accepted 2006) (Seehausen reverse speciation through biodiversity current or reduce 1987) (Slatkin species new of evolution the and natural selection oppose 1996) Simberloff Rhymer & 2007; et al. Muñoz-Fuentes 1994; et al. (Gottelli rarespecies of pool the gene dilute may flow gene species, diverged relatedorrecently between closely occurring When Article Gene flow can be considered as a constraining or a creative force in evolution (Slatkin 1987) (Slatkin evolution acreative in or force asaconstraining be considered can Gene flow

. Conversely, genetically isolated isolated genetically Conversely,

of the Atlantic and Pacific and Atlantic the . , Dearborn et al. 2003; Gómez-Díaz et al. 2009; Milot et al. 2008; Morris-Pocock et al. 2010) et al. Morris-Pocock 2008; et al. Milot 2009; et al. Gómez-Díaz 2003; et al. Dearborn 2011; et al. (Ando betweenislands differentiation population in result always does not philopatry al et Gómez-Díaz 2015; Friesen in (reviewed populations seabird dividing animportant in role also play may that oceanregimes local to andadaptation 2010) et al. Morris-Pocock 2001; & Croxall (Burg distribution as non-breeding in such seabirds, flow gene barriers to potential other many arethere 2003; Austin et al. 1994; Avise et al. 2000; Burg & Croxall 2001; Welch et al. 2012) et al. Welch 2001; &Croxall Burg 2000; et al. Avise 1994; et al. Austin 2003; &Double (Abbott distinct becoming of seabirds populations island lead to can breed to natal island instinct for philopatry (Friesen et al. 2007; Steeves et al. 2005) et al. Steeves 2007; (Friesenet al. philopatry for instinct a strong by driven disperse, to reluctance their with coupled dispersal potential for huge between their dichotomy tothe due flow, gene and dispersal between relationship the and genetics evolutionary examine to which in model interesting an provide seabirds In particular, 2015). et al. (Grewe tuna and 2010) et al. Portnoy 2012; al. et (Blower sharks 1994), et al. (Baker whales in example for flow, gene to constraints behavioural show mobility incredible their to due populations form panmictic This article isThis article by protected copyright. All rights reserved. finches Darwin’s in e.g. environmental conditions changing to unsuitable may be phenotypes example, hybrid species; for bird of range awide in documented been have flow gene interspecific barriersto barriers. Biological biological oceans)or mountainranges, distance, (e.g. by physical prevented isusually but 2006), McCarthy 2005; Mallet & Grant1992; (Grant nature in common fairly species is between bird Introgression 2015). (Friesen understood remains poorly scales spatial atdifferent species between flow gene for potential colonies, the island at different seabird species is often restricted (Friesen 2015), therefore introgression of genes from one seabird genes fromof seabird one therefore introgression 2015), (Friesen restricted isoften species seabird asingle of populations between distinct Geneflow 2014). et al. (Neubauer gulls Larus in rate e.g. survival adult or areduced have 2005), et al. eagles (Helbig and Spotted 2008) et al. Svedin 1992; et al (Gelter Flycatchers sexes in e.g. both one or in fitness reproductive may suffer reduced hybrids

Accepted 2009). Schluter . 2009; Article

In marine ecosystems physical barriers to gene flow are few. Marine species that could potentially potentially could that species arefew.Marine flow gene to barriers physical marineecosystems In Although many studies focus on the gene flow and genetic structure within single seabird species single species seabird within structure genetic and the focusgene flow on many studies Although

. This preference for returning to their their to returning for This preference (Grant & Grant1992) (Grant . Natal Natal , but ,

. population. There is likely to be a range of possibilities that describe gene flow in petrels to and fromand to petrels in flow gene describe that possibilities arange of be to Thereislikely population. Pterodroma Island the Round flowinvolving gene of models betweenpotential two distinguish conditions. environmental underdifferent survive new areasand tocolonise for ability their implications important of many wide-ranging seabirds (Croxall (Croxall seabirds wide-ranging many of status conservation threatened separated. the Given and sympatrically allopatrically formerly are both that betweenspecies barriers of the breakdown and flow gene ofdispersal, the study role to which in ‘natura interesting particularly a represents therefore Island on Round population petrel The inthePacific. tohybridise known arenot they petrels, Herald and Kermadec of range Pacific This article isThis article by protected copyright. All rights reserved. (Brown here hybridise arminjoniana of colony 2015). et al. Brown 1982, Hunter giant petrels, 2012; 2010, et al. Taylor boobies, et al.1993; exist(murres, Friesen hybrids seabird occurring naturally of examples However, unlikely. very considered may be another species to several breeding locations (BirdLife International 2016b; Brooke 2004) Brooke 2016b; (BirdLife International locations several breeding 2005) Tove Imber2005; 2008; (Imber2004, evidence insufficient to due dismissed largely and debated been Atlantic has Ocean the in petrels Kermadec of presence thepossible although Herald petrels, or Kermadec with confirmed contact no petrelshave Trindade range Atlantic their in Ocean, Indian Accepted Article2004) (Brooke archipelago Vaz Martim and Trindade atthe Atlantic South the is in petrel theTrindade of location otherbreeding only The Ocean. Indian the species in Here microsatellite genotyping data, supplemented by geolocation tracking data, are used to to used are data, tracking geolocation by supplemented data, genotyping microsatellite Here On Round Island, off the coast of Mauritius in the south-western Indian Ocean there is an unusual isanunusual Oceanthere Indian south-western the in coast Mauritius the Island, of off Round On . In contrast, in the Pacific Ocean Kermadec and Herald petrels share a similar range, and and range, similar sharea Herald petrels Kermadec and Pacific Ocean the . contrast,in In Pterodroma , Kermadec petrel, Kermadec petrel, petrels. The population includes three species (Trindade petrel, (Trindade species includes three population The petrels. et al . 2011; Brown . 2011; P. neglecta P. et al. et al and Herald petrel Herald and 2012) the potential for inter-ocean gene flow has has flow gene for inter-ocean the potential 2012) . 2010), and is the only confirmed colony theseof colony confirmed only isthe . 2010), and , P. heraldica , P.

. Unlike the population in the the in theUnlike population . Despite the overlapping overlapping Despite the ) , known to extensively extensively knownto P. P. l experiment’ experiment’ l This article isThis article by protected copyright. All rights reserved. as referred to hereafter scenario, this scale. In ataglobal occur may species flowbetween gene and introgression that possible it is petrels, Pterodroma of dispersal potential thehuge given However, Island. therefore model’ 2010) etal. Brown 2011; et al. (Brown Pacificspecies and contactbetweenAtlantic secondary of is apoint Island Round that presumed been it has past, the extreme cases.two In represent thisstudy in models the tested Island,however Round Sample Collection and DNASample and Collection extraction Information. theand Supporting (2016) et al. Nicoll see tagging, geolocation and ringing on details For trips. inter-ocean making these of wererecorded two and migration, on recorded were geolocators with petrels fitted 330 the 116of and 2012, Between 2009 1B). Figure east, 57.78° ofPetrels Round tracking Island and Monitoring Methods gene-flow. two modelsbasic of our for theevidence investigate to we aimPacific Oceans and Indian Atlantic, the across populations island genotyping Atlantic.the By into Pacific genotypes and the into PacificOcean, genotypes Trindade of introgression of thepossibility investigate we oceans, between petrels disperse Pterodroma that co petrels from the Trindade Islands and Kermadec petrels from the Kermadec Islands (Table 1 (Table Islands Kermadec the from petrels Kermadec Islands and Trindade the petrels from in seen be can morphs plumage petrel Island Images Round of petrels’. Island

Accepted Article evidence historical the Given Ocean. Indian outsidethe Island, Round than islands other on -occur Blood samples were collected from Round Island Pterodroma Pterodroma Island from samples Round collected were Blood south; Reserve (19.85° Nature Island onRound ringed routinely petrels been 1994, have Since , individuals disperse from the Atlantic and Pacific to Round Island in the Indian Ocean, and and the Ocean, in Island Indian Round to Pacific and Atlantic from the disperse , individuals

co -occurrence of Atlantic and Pacific species and their hybrids should only arise on Round Round ariseon only should hybrids their and Pacificspecies and Atlantic of -occurrence the ‘ the widespread gene flow model’ flow gene widespread . In this scenario, referred to here as the ‘ the as here to referred scenario, . Inthis , Atlantic and Pacific species and their hybrids may may hybrids their and Pacific species and , Atlantic

(hereafter referred to as ‘Round ‘Round as to (hereafter referred Figure S1), Trindade Trindade S1), Figure secondary contact contact secondary , Figure Figure This article isThis article by protected copyright. All rights reserved. from the wassampled tissue species footpad study the of represent Pacificranges To the Pterodroma. tropical due petrels Murphy’s and Herald for and ofKermadec petrel populations Pacific other for samplesBlood wereunavailable 1B). (Table 1 (Table genotyping for out-group asan included species), were other the than less similar (phenotypically petrel Pacificgadfly anothertropical ultima), Murphy's of (P. petrel populations island samples from two Island.Additionally, at S2C) Round Figure underwing, a uniform (having brown plumage petrel-like Phoenix with of petrels sightings been have there since collected, were also petrels alba) (P. fromPhoenix samples this, investigate To fromIsland Pacific. the Round reaching species Pterodroma be additional there could that proposed (2011) al. et Brown Kermadec petrels, 002 TG03- included; also species were study in our utility cross species showedloci also that additional following the final marker set, Inthe 1995). et al. (Ellegren species study the two of three only from wereisolated markers that the of the selection caused by bias for ascertainment evidence of look species to study three the between compared was markers these of diversity The genetic heraldica. and neglecta P. arminjoniana, P. focalspecies: three the in amplified that were chosen heraldica) and from six arminjoniana P. from (five loci microsatellite validated 11 From libraries, the from Gambier Islands. the heraldica P. one and from Islands Trindade the arminjoniana P. from one development genotyping marker and Microsatellite procedures. standard used methods, which DNA extraction the to details of addition in methods, genotyping error rates and tests of Hardy-Weinberg equilibrium were carried out for each locus within within eachlocus for carriedout were equilibrium Hardy-Weinberg error tests of ratesand genotyping allelenull frequency, of Estimates PCR cycle. atouchdown and DNA polymerase Taq HotStar al

Accepted with PCR kits QIAGEN Multiplex using DNAwasamplified loci. of 16 atotal giving . 2007), Article , Two di- and tetra-nucleotide repeat enriched genomic libraries were created from blood samples blood from created were libraries genomic repeatenriched tetra-nucleotide and di- Two TG13-009 , American Museum of Natural History’s collection (Figure S2). (Figure collection History’s Natural of Museum American Figure 1B). The Supporting Information provides details of sample collection and storage storage and sample collection detailsof Information provides The Supporting Figure 1B). , TG13-017 (Dawson et al. 2010) TG13-017et al. (Dawson to the inaccessible nature of many of the islands in the Pacific range of of the thePacific range in of islands many of nature inaccessible the to , Tgu06 (Slate et al. 2007) and Calex- 2007) Tgu06et (Slate al.

In addition to Herald and to Herald and addition In 01 (Küpper et (Küpper P. P. P. P. locus, H locus, per alleles numberof The S1). Table and Results markers, potential see of 16 set(out final the in ra native its in petrel) Phoenix petrel and Murphy’s petrel, Kermadec petrel, ( developed from petre developed Trindade This article isThis article by protected copyright. All rights reserved. (H heterozygosity expected of diversity genetic Calculation Information. in theSupporting be found can marker selection and PCR conditions loci, microsatellite newof method, testing development library details the Further of eachpopulation. Estimation of Estimation differentiation genetic Team 2013). Core (R Development v.3.1.2. R environment programming the in ANOVA using performed test was This species. different the between markers these eight in andallelicrichness heterozygosity observed comparing wasby tested 1995) et al. (Ellegren bias ascertainment for DNA,evidence heraldica) (P. differentiation narrowed to K= to narrowed 3 was clusters specified numberof the therefore five, and between three lay clusters of number likely 2000) et al Pritchard 2003; et al. (Falush v.2.3.4. softwarethe STRUCTURE clustering using was estimated samples structure acrossall samples. Population contemporary and historical between the drift genetic of evidence was there whether investigate to location, and samesamples species the of contemporary the and Islands Kermadec from the petrelsamples Kermadec historical the between difference Accepted ArticleAr ) was calculated in FSTAT v.2.9.3 (Goudet et al. 2002). Since three of the 12 markers used were 12markersused the three of Since 2002). et al. (Goudet FSTAT v.2.9.3 in calculated ) was

Genetic diversity was measured by calculating the number of alleles per locus, observed and and observed locus, per numberthe calculating alleles of by wasmeasured Genetic diversity F ST . Preliminary analyses using STRUCTURE (Supporting Information) showed that the most the Information) showed that (Supporting STRUCTURE analyses using . Preliminary calculations were performed using FSTAT v.2.9.3.2 (Goudet 2001) to describe genetic to describe genetic 2001) (Goudet v.2.9.3.2 FSTAT using wereperformed calculations o and H between island populations. Additionally, F Additionally, populations. between island e were calculated in CERVUS v.3.0.7 (Kalinowski et al. 2007), and allelic richness allelicrichness and 2007), et al. (Kalinowski CERVUS v.3.0.7 in were calculated – 5 based on this evidence. Fifteen independent models were run for each number for of run models were independent evidence.Fifteen this on based 5 o and H and l ( P. arminjoniana) DNA, and five were developed from Herald petrel petrel from Herald were developed five DNA, and arminjoniana) P. e ), and allelic richness for each species (Trindade petrel, Herald Herald petrel, (Trindade eachspecies for and allelicrichness ST was calculated to quantify the genetic the genetic quantify was calculated to

nge, using the 12 markers 12markers the using nge, . This article isThis article by protected copyright. All rights reserved. 10 with clusters, specified probabilities for each value of K (Evanno et al. 2005). The output for K= for output The 2005). K for of eachvalue probabilities 3 et al. (Evanno K of value most the likely identify to HARVESTER (K the STRUCTURE mean Ln using into re-entered then were models These (Earl 2012). v.0.6.94 HARVESTER service STRUCTURE K of eachvalue for onlinelikelihood meanthe Ln highest using 15 total the of out were chosen the with models 10 admixture. The assuming and information, population prior any without were run Hybrid classification classification Hybrid the in Information. found Supporting can be these of outcomes settings and the Details on S7). (Figure populations petrel Phoenix and populations Herald petrel S6),and (Figure Island Trindade and Island Round between compare clustering to werealsoperformed analyses STRUCTURE Separate Information). identification(Supporting hybrid intervals on credibility of examine effect the to 1), = (ANCESTDIST turned on option ANCESTDIST the with out carried was separately analysis S5. This Figure as shown in 2004), (Rosenberg v.1.1 the softwareusing DISTRUCT visualised and 2007) & Rosenberg (Jakobsson software CLUMPP the using summarised wasthen HARVESTER ‘Trindade type’ (PureT), ‘Kermadec type’ (PureK), ‘Herald (PureK), ‘Kermadec type’ (PureT), type’ ‘Trindade were: These classes 1). (Figure analysis STRUCTURE using identified clusters to (Q) membership Herald- (PureM), type’ Accepted for >= 0.1 Q if hybridswereidentified Two-way forclusters. allother <0.1 and one cluster for >=0.9 was (Q) membership estimated clustertheir if a single to as belonging wereidentified individuals Forexample, (2011). et al. Marie and (2006) Primmer and Vähä on based (Table S2), groups more two or of ahybrid or to group species toa particular solely for belonging thresholds membership cluster using distinguished Classes were (TxHPxK). hybrid Kermadec and Herald-Phoenix Trindade, Article Petrels were assigned to one of nine possible pure o possible pure nine oneof to assigned Petrels were Phoenix and Kermadec hybrid (HPxK), Murphy’s and Herald and (HPxK), Murphy’s hybrid Kermadec and Phoenix Trindade and Kermadec hybrid (TxK), Trindade and Herald-Phoenix hybrid (TxHP), hybrid Herald-Phoenix and Trindade (TxK), Kermadec hybrid and Trindade 6 MCMC iterations, and a burn-in period of10 period aburn-in and iterations, MCMC r admixed ‘classes’ based on their estimated estimated their on based r admixed ‘classes’ - Phoenix type’ (PureHP), ‘Murphy’s (PureHP),type’ ‘Murphy’s Phoenix

-Phoenix hybrid (MxHP) (MxHP) and hybrid -Phoenix 5 . All STRUCTURE analyses analyses STRUCTURE . All – 5 from STRUCTURE from5 STRUCTURE ) 10 al et Davy Following 2001). Beerli& Felsenstein (Beerli 2009; was sampled space parameter adequate that and 60% and 20% fellbetween acceptance rate the to ensurethat runs inpreliminary respectively, coefficients, andinbreeding rate, allelefrequency migration for 0.40 0.60 0.15, and to were adjusted parameters Mixing 1). distinct (Figure weregenetically asthese populations geneflow, of analyses 2003) Rannala of Estimation migration run with 10 with run longer afinal, in were used this run seedfor starting parameters mixing and andthe Meirmans (2014), in developed an R-script calculated using Bayesian deviance using the wasidentified of ten run allele frequency (>0.2) in three or more of the five species, and these were therefore discarded from discarded therefore were species,these five and the more of three or (>0.2) in allele frequency null estimated had ahigh loci the 16 of out four that each revealed species within separately markers microsatellite ofautosomal suite the of Analysis eachspecies. males for females and known in the o presence on based petrel) Murphy’s and Phoenix Herald, Kermadec, (Trindade, Results (2015). et al. Davy in asseen samples, Island Round the without but as above samethe parameters using also run was analysis the Pacific Oceans, and Atlantic the of betweenpopulations contact secondary point of or introgression stone for stepping This article isThis article by protected copyright. All rights reserved. clusters, and two 0.1 for three clusters, and three clusters, and for 0.1 . (2015), ten separate analyses were run using different random starting seeds. Each run had 2.5 x had 2.5 run seeds. Each differentrandom starting using were run tenseparate analyses . (2015),

Accepted 7 Article iterations and a 1.5 10 x a1.5 and iterations Gene flow was estimated using the software BAYESASS v.3.0.3. (Rannala 2012; Wilson & Wilson 2012; (Rannala v.3.0.3. softwarethe BAYESASS using estimated was Gene flow The microsatellite loci used in the analyses were autosomal in all of the five petrel species tested tested species petrel five the allof in analyses autosomal the were in used loci microsatellite The

8 iterations and a burn-in 10 of aburn-in iterations and . Based on preliminary STRUCTURE results, Murphy’s petrel was not included in in included was not petrel Murphy’s results, STRUCTURE preliminary on . Based Q < 0.1for the remaining two clusters. Thr clusters. two theremaining < 0.1for Q 7 < 0.1for the remaining cluster. cluster. theremaining < 0.1for burn-in, and the default sampling interval of 2000 iterations. The optimal iterations. The 2000 of samplinginterval default andthe burn-in, 7 . To investigate the role of Round Island as a possible asapossible Island Round of role the . Toinvestigate ee -way hybrids were identified if Q if identified were hybrids -way

f heterozygotes f heterozygotes >= This article isThis article by protected copyright. All rights reserved. (Parm20 analysis further H ANOVA, (one-way marker the set across allelicrichness and heterozygosity observed the museum) on or (blood type sample effect ±0.01). of S.E. significant ±0.07) was S.E. no There =0.02, (mean samples than blood =0.15, (mean for museum samples washigher allelic dropout Per-genotype (Table S5). negligible be to estimated datasetwas the in alleles ±0.01) (S.E. was0.02 false occurrence of the and genotype rateper- dropout Average allelic individuals 1). (Table 885 sample size a total of leaving analyses, from further 12 markers wereremoved remaining the of (eight) atleast 75% for amplify to failed that Data from samples together. clustered were petrels which Phoenix Herald and of exception the with analysis, the in speciesincluded the with thatwas consistent structuring agenetic reflecting S5), S4and 1A, (Figure runs between consistent were results the and markers different numbers of using performed were analyses STRUCTURE However, 2007). et al. (Kaeuffer analyses STRUCTURE in clustering of overestimate an to can lead LD dataset. the Strong leftin were these loci Phoeni (Trindade, Kermadec, groups species other the in was LD detected no and LD loci displayed pairs of few Since petrels. Herald andtwo in population, Island Round the in five S4), Table (LD, disequilibrium linkage displayed loci pairs of seven Island), at Round population mixed species the speciesand groups(five allsix in disequilibriumall loci between linkage of calculations paired 396 of Out orlinked selection. study, this to whichwere interest of groups, between genetic structure and migration indicate can equilibrium Hardy-Weinberg from the Deviation 2). species(Table five the or of more three for <0.05) equilibrium (P-value from Hardy-Weinberg the (Parm34, Parm29 (Parm34, Calex01 Weinberg equilibrium equilibrium Weinberg Accepted Article eightmarkers without (Parm34 and with both was conducted analysis , TG13-017 , Parm22, Parm20, Calex01, TG13- Calex01, Parm20, Parm22, , Phel12 O (P

: F , -value <0.05) for three or more of the five species (Table 2). STRUCTURE 2). (Table five species the more of or three for <0.05) -value Parm34 1, 1, 11 , Phel30, see Figure 1A and S4) that due to the Wahlund effect deviated effect deviated Wahlund the to that due S4) and see 1A Figure Phel30, = 0.09, P = 0.77; P = 0.09, = 0.77; , Phel30 and TG13- and Phel30 Ar : F 1, 11 017 017 = 1.09, P = 0.32). P = 1.09, = 0.32). ). Of the remaining 12 markers, eight eight remaining12 markers, Of the ). , Phel12, Phel30 Phel12, ,

, Parm29 ) deviated from the Hardy- deviatedfrom the x and Murphy’s petrels), Murphy’s and x , Parm22 , Parm20 , Phoenix- or Herald- petrels, Kermadec-type petrels, Trindade-type these describe to appeared and four, Ocean, whereas Kermadec-type, Herald/Phoenix- Kermadec-type, Ocean,whereas Atlantic the of were characteristic individuals Trindade-type distinctions, species adhered to largely that clusters STRUCTURE the from might expected be As groups. species more two or of hybrid four the on species, based ‘pure’ a either to individuals assign was to used clusters these eachof to (Q) membership Estimated same tothe assigned cluster. wereranges both Pacific. The proportion of individuals in each class for island populations is shown in in isshown populations island in for eachclass individuals of proportion Pacific. The between these temporally separated populations (F separated populations temporally between these genetic differentiation significant wasno there demonstrated 2011) et (Brown al. recently collected thoseand (1920s) Islands Kermadec the petrelsfrom Kermadec museum sampled between the species-specific markers between the different species we sampled. wesampled. species markers different the between species-specific the in 1995) et al. (Ellegren bias of ascertainment evidence wasno there revealed diversity genetic of from Ducie Atoll (F Ducie Atoll from petrels Herald of Island-populations Pitcairn two from the differentiated significantly not were Islands from Pitcairn samples collected the petrel Phoenix same the of However, species. populations island (five) were between majority the comparisons, Of these seven S6). (Table different significantly non- being pairs population island of 105 seven out only with differentiated, significantly most were between (H differ loci did species: F Pterodroma sample DNA did not differ significantly between the different species (H species different between the not differ significantly did sample DNA Pterodroma This article isThis article by protected copyright. All rights reserved. Accepted Article The most likely number of genetic clusters in the dataset using STRUCTURE was found tobe found was STRUCTURE datasetusing the in clusters genetic of number mostThe likely Analysis of genetic structure between the island populations of petrels using F petrelsof using populations island the between geneticstructure of Analysis The observed heterozygosity and allelic richness in the eight markers developed using using developed markers eight the in allelicrichness and heterozygosity observed The type petrels and Murphy’s type and petrels 4, 4, 28 = 0.47, P = 0.76; P=0.76; = 0.47, ST

= -0.01, P = 0.56), and from Oeno Island (F Island from and Oeno =0.56), P -0.01, O : F 7, 7, 28 Ar = 11.48, P <0.0001; P = 11.48, between species: F between species: -type petrels (Figure 1A). 1A). -type petrels(Figure type Murphy’s and ST = 0.004, P = 0.10). P=0.10). = 0.004, 4, 4, 28 Ar = 0.38, P = 0.82), although genetic diversity although geneticdiversity P = 0.38, = 0.82), : F 7, 7, 28 He

= 17.97, P < 0.0001). This comparison This P <0.0001). = 17.97, rald and Phoenix petrels across their their petrelsacross Phoenix and rald ST = 0.01, P = 0.24). Comparison = P 0.24). = 0.01, -type were characteristic of the the of werecharacteristic -type possible clusters, or a a or clusters, possible ST suggested that that suggested O between between ± 0.051). However, no reciprocal gene flow was observed from the sampled Phoenix petrel petrel sampled Phoenix from the wasobserved flow gene ±reciprocal However, no 0.051). 0.063 ± (Pitcairn 0.048 Islands) andPhoenix (Phoenix Islands) 0.046 ± 0.076 Island) Phoenix 0.052, ± 0.04 (Marquesas (Christmas Islands) ± Phoenix to Phoenix 0.04, 0.132 Atoll) (Ducie 0.071; Herald Islands) (Pitcairn Phoenix to Islands) (Herald(Marquesas islands different of atanumber populations petrel Phoenix to Atoll Ducie and Marquesasthe Islands on populations petrel Herald calculated from migrationrates were ± Round 0.048 Significant Island. to 0.01) Islands, Kermadec (Kermadec ± 0.014 and Islands, 0.01) (Marquesas Herald generation), per individuals Island Round of 0.036 ± (0.183 from movement Trindade was significant there whereas PacificOceans, or Atlantic the either islands, in to other Island from Round analysis the BAYESASS wasdetectedby migration significant No Figure 2. by rates are migration illustrated Significant Information. Supporting S8in the and S7 in Tables fully are described population Island Round mixed the petrels and Phoenix Herald petrels, Island. fromRound originated 39 of which cluster, Herald-Phoenix and Kermadec Trindade, between the assignment split three-way had 48 a globally, petrelsgenotyped 885 Of the asaKermadec-type. was classified Ocean Atlantic the petrels of Trindade from sampled the one individual and cluster, species Trindade-type the to as belonging were Kermadec classified the Islands of population petrel theKermadec sampled from Twoindividuals island. that the of ocean to different froma originating with acluster hybridised be to appeared respectively, % of22.2 eachpopulation, and 18.9% 24.1%, these In groups, hybrids. as classified were Islands petrelsfrom Trindade Trindade the of 22.2% and Islands Kermadec the from petrels of Kermadec 20.8% Rapa Island, petrelsfrom Kermadec of sampledislands 31.0% other the of however; Island toRound unique wasnot clusters Admixture between hybrids. as classified ofindividuals 43.2% with population, admixed mostIsland wasthe Round S3. Table in found This article isThis article by protected copyright. All rights reserved. Accepted Article Estimates of migration rates between island populations of Trindade petrels, Kermadec petrels, petrels, Kermadec petrels, Trindade of populations island rates between migration of Estimates canbe population island per eachclass in be calculated to individuals andnumbers of 1B, Figure

This article isThis article by protected copyright. All rights reserved. on12 Round Island on geolocator the with was recovered The petrel breed. to areknown where Heraldpetrels 3), (Figure Island Raine close to also and Island Ho Lord on petrels known Kermadec of nearestcolony the close to travelled Hereit Pacific Ocean. the into eastwards andtravelled Island Round left 5H41524 petrel 2010 In February 3). (Figure Pacificpopulations. and Atlantic wasseenbetween the migration significant no analysis, from the wasremoved Island Round Pacific.When the petrel in species other with individuals exchange to observed were not petrels Kermadec populations. Herald petrel the into populations petrel. The petrel was subsequently recaptured on Round Island Island on 1 on Round recaptured was petrel subsequently The petrel. Trindade the site of breeding Atlantic the archipelago, Vaz Martim Trindade and the to Ocean,close Atlantic the into travelled Africaand tip of the southerly passed it around Oceanuntil Indian the higher on Round Island than in the other island populations sampled (Figure 1). Notably, admixture admixture 1). Notably, (Figure sampled populations otherisland than the Island in onRound higher Accepted were clusters Herald-type and Kermadec admixture between levels of however cluster, Trindade-type the to belonging individuals of mainly consist wasto Island shown Round on of petrels population The oceans. across three speciessampled five the between structure population the represented best clusters that four found STRUCTURE using data microsatellite of genotyping Analysis seabirds. Discussion (1.7%). Herald/ the to assignment alow had and cluster (28.7%), Trindade the alsoto but cluster(61.9%) Kermadec tothe predominantly wasassigned 5H41919 2 Island on Round departed 5H41919 petrel contrast, In shafts. Article paleprimary characteristic with plumage, pale in large and comparatively asitwas Kermadec petrel resembled a bird this phenotypically although 5H41524, the for available was information genotyping Only two individuals (~1% of petrels successfully tracked with geolocators) left the Indian Ocean Indian left the geolocators) with tracked successfully petrels (~1%of individuals two Only This study presents the first evidence of inter-oceanic gene flow between tropical Procellariform tropical between flow gene inter-oceanic of evidence presents first the study This

Phoenix cluster (7.7%) and Murphy’s cluster cluster Murphy’s and cluster (7.7%) Phoenix th November 2012. Unfortunately no Unfortunately 2012. November

nd October 2012 and remained within remained within and October 2012 st June 2013 (Figure 3). Petrel 3). 2013(Figure June we gene flow models that might best represent gene flow between the tropical tropical the between flow representgene best might that models flow gene more of specific are awiderange there Therefore PacificOceans. and Atlantic the between flow gene migrationand may facilitate population Island the Round reality admixed in that is plausible it and This article isThis article by protected copyright. All rights reserved. contact secondary contrastingThe from analysis. Islandwas the removed Round when persisted aresult that 2), (Figure reciprocally Pacific,or the to Ocean from Atlantic the rateswere detected migration generation per- significant no However, Ocean. Indian the outside flow gene and dispersal for evidence strong Pacific, the providing of petrel populations Phoenix Herald and Kermadec, and Ocean Atlantic the of population petrel the Trindade in Ocean, the Indian outside also seen was between clusters the findings of Brown et al. (2011), who found that one sampled Ducie Island Herald petrel possessed petrel Herald Island onethat sampled found Ducie who (2011), et al. Brown of findings the Accepted corroborates mixing population of This evidence Figure 1B). 11.1%, theIslands Pitcairn and 6.3%; Islands Phoenix 3.4%; Islands Marquesas 6.6%; (Christmas Island petrelpopulations Phoenix and 4.8%) Island Oeno 13.0%; Islands Marquesas 7.1%; (Ducie Atoll populations Heraldpetrel 24.1%), RapaIti 18.9; KermadecIslands 17.9%; Islands Fernández Juan 4%; Atoll (Ducie populations petrel in PacificKermadec were found genotypes hybrid Atlantic (Trindade-type) Similarly, 1B). (Figure migrant Kermadec-type apure wasclassified as individual one and respectively), the sample, 4.4% of and genotypes (17.8% hybrid Herald/Phoenix-type or Kermadec-type either having as wereclassified population Trindade Atlantic the in petrels some However, 2006) 2002, Randler 2011; & Wilson (McCracken related species of a population species in for rarer options mating inconspecific adeficiency of arearesult hybrids whereby 1955), (Hubbs 1B). (Figure populations Pacific and Atlantic than (43.2%) admixed to bemore appeared Island population Round the that 1A) (Figure analysis STRUCTURE the seenin was it clearly and Ocean, Indian the from outside Article species. between the links clarify to needed is research phylogenetic more detailed and this in study, included Historically speaking, the population of petrels on Round Island consists entirely of immigrants immigrants of consistsentirely Island Round on of petrels the population speaking, Historically and and widespread gene flow gene widespread This may relate to the Hubbs’ principle, or the “desperation hypothesis” “desperation orthe principle, Hubbs’ the to relate This may models presented in this study are scenarios, extreme study this in presented models

Pterodroma populations populations . This article isThis article by protected copyright. All rights reserved. cytochrome mitochondrial a Trindade-type genetic divergence between populations is low (F islow populations between divergence genetic if or 2016), et al. (Samarasin recentgenerations in has reduced connectivity ifpopulation example wasobserved. flow gene reciprocal no 2). However, (Figure Island Round to ± 4.8 (Kermadec Islands, population petrel 1%) ± Kermadec 1.4 Islands, the closest sampled 1%)and (Marquesas population petrel ± sampled Herald (18.3 fromclosest 4%), migration likewise the and population Island Round the the into Atlantic of population petrel flow from Trindade gene of the level high a that wasthere ratesuggested migration generation Estimates per of analysis. BAYESASS the in found as Oceans Atlantic and Pacific between the directly or Oceans, Atlantic Pacific or Pacificpopulations. and Atlantic between the flow forgene population intermediate actingasan and istherefore likely highesthad proportion the Islandpopulation Round flow gene widespread model. the but on Round Island, found exclusively werenot genotypes Hybrid the of variant the stepping-stone supports analysis STRUCTURE here, common asidentified be as not may hybrids although Therefore Information). (Supporting study this in criteria used the using hybrids as classified correctly individuals of reject88.4% would option ANCESTDIST S interval, credibility 90% a at showed that population hybrid simulated a using investigations However, status. hybrid true their on doubt casting hybrids, clusters all for potential to assignments for one to from zero Intervalsranged study. inthis caution must with be assigned hybrids Islands. Kermadec from the Kermadec petrels with haplotypes petrels sharing threshold (F threshold Accepted this be below to estimated Atlantic was the petrelsof Trindade the petrelsIsland and between Round Article These findings are apparently contradictory to the lack of migration from the Indian Ocean to the the Ocean to from lack migration Indian of the the to contradictory areapparently These findings putative suggested that analysis STRUCTURE during calculated intervals credibility 90% The BAYESASS has been shown to overestimate migration rates in certain circumstances, for for circumstances, certain ratesin migration overestimate to shown has been BAYESASS ST = 0.02, P < 0.01), and therefore the calculated migration rate of 18.3 ±18.3 rate of migration per 4%migrants andtherefore calculated the P <0.01), = 0.02, b haplotype, in addition to some Ducie Island Herald Herald some Island to addition Ducie in haplotype, ST

≤ 0.05, Faubet Faubet 0.05, ≤

et al. 2007). Genetic divergence Genetic divergence 2007). et al. TRUCTURE’s TRUCTURE’s 2004) et al (O'Reilly locus polymorphismand 2013) et al. (Jakobsson allelefrequency by influenced can be and Atlantic Oceans, the number of genotyped individuals from each of the of from individuals each genotyped numberof the Oceans, Atlantic and Wilson & Rannala 200 & Rannala Wilson 2007; et al. Faubet 2010; &Gibbs (Chiucchi asfewer five taken than usually afewgenerations, only the potential connectivity of their isolated populations (Figure 3). It is interesting that these two these two that Itisinteresting 3). (Figure populations isolated their of connectivity potential the demonstrating Oceans, Atlantic Pacificand the in colonies other close to coming tracked were (~2%) petrels. Phoenix and Herald petrels Pacific on conducted been have studies phylogenetic detailed asno well resolved, are not species these that two be may it Island.However, Round on genotypes petrel Phoenix of detection the enabled may and have (Figure S7), petrelsparticularly Phoenix and Herald between structure detect genetic to analyses of the power the haveincreased would This species. fivestudy our to all of specificity primer complete with markers marker homologous of microsatellite set alarger have to advantageous researchers. by visited infrequently and remote arevery arefound petrels wherethese the islands but studies; further to useful be would modernsamples of Thecollection populations. the all be same for would difference lack of this that it wasassumed and separatedgroups, temporally between two the variation differencegenetic in significant was there no Islands, Kermadec the from samples Kermadec petrel very low (Meirmans 2014) low very migrationrates are rate ifthe aper-generation bedetectableat potential to migrants enough include to are unlikely populations other ±sample for (S.E. sizes Therefore, 1.7). small,average 26.7 on very This article isThis article by protected copyright. All rights reserved. Accepted Article F on based Islands Trindade the Islandand Round between migrationrate calculated the dismissing anoverestimate. be However, may generation Although the number of tracked petrels from Round Island was limited (N = 116) two individuals individuals two (N =116) waslimited Island petrels from Round the number tracked of Although Although there was no evidence of ascertainment bias (Ellegren et al. 1995) , it would have been havebeen , it would 1995) et al. (Ellegren bias ascertainment of thereevidence was Although no . Additionally, due to the difficulty of gaining samples from the remote islands of the Pacific the of remote islands samples from the gaining of difficulty the to due Additionally, 3) . However, when F . However, when . Finally, BAYESASS analysis assumes that populations are separated by by areseparated populations that assumes analysis BAYESASS Finally, ST was calculated between contemporary andhistorical contemporary between was calculated ST must be viewed with caution, as maximum F asmaximum caution, with viewed must be

se island populations was populations island ST

. This article isThis article by protected copyright. All rights reserved. and lessonii) Accepted (Pterodroma petrel White-headed a between asahybrid is posited Islands Antipodes the eastof Seas Expedition the South Whitney during collected asinglemuseum specimen of study phenotypic mitochondrial and although 2010), et al. Brown 2011; et al. (Brown Island from Round researchers. visited by rarely and isolated petrels are Pterodroma by inhabited islands unusual, asthe highly is evidence colony-switching of providing records ringing of occurrence However,the old. years at minimum 28 2012, a of age in May again and in2008 egg October with an Island Round on present as recorded wassubsequently Thisindividual & Reimer 1991). King 3; (Figure 1987 until partner was petrel The 2010). (Tatayah chick ayoung with wasdiscovered petrel pale-plumaged aringed, 2006, inApril Island onRound monitoring routine seabird during data.However, return would not colony change permanently that geolocators with tracked anybirds be obtained, to data for be recovered must since geolocators Unfortunately, are unknown. colonies natal their Island,so Round on as adults Article ringed and caught both wereinitially although Round Island, to returned Oceansubsequently Indian the outside petrels tracked Both locations. of breeding a switching in result colonies will breeding Atlantic. the presence in their support to evidence adds here presented data tracking and genotyping the butboth 2005), Tove Imber 2005; 2008; (Imber2004, the past in contested has been Ocean Atlantic the in breeding potentially and petrels visiting Kermadec of The possibility analysis STRUCTURE cluster in the Kermadec-type to assignment petrels(61.9% Trindade than PacificOcean the petrels of similar Kermadec to moregenotypically appeared Island Round from the Ocean Atlantic to travelling the petrel surprisingly, Most directions. migration toestimated counter travelling tracked petrels were To date the only well-studied instance of introgression between species of Pterodroma petrel is is petrel Pterodroma of betweenspecies introgression of instance well-studied theonly date To other visits all to not means that Procellariformof seabirds philopatry strong the Of course, originally captured as a Herald petrel on Raine Island in 1984, where it bred with the samethe with bred 1984, where it in Raine Island on petrel as aHerald captured originally

This article isThis article by protected copyright. All rights reserved. mollis (P. petrel Soft-plumaged Accepted presence this The of events. tostochastic itssusceptibility and therefore range limited breeding petrel Trindade the for status IUCN List Red ‘vulnerable’ the example, For someseabirds. of management status and conservation the of assessment the to relevant may be particularly colonies with other species and hybridisation and migration of species. Consideration wide-ranging related, closely of populations between dispersal and flow gene about isknown little how highlights study future. This their well for bodes new in environments thrive to potential the and islands, extirpated colonise new or them and help disperse may to Pterodroma tropical behaviour of wide-ranging Island. The Round to unique hybrids arenot and migrants ocean inter- However, PacificOceans. inthe and Atlantic species originating contactbetween secondary important of an is Island zone clearly Round populations, island and differentspecies of complex this Within species. related closely these between flow gene widespread and dispersal supporting unique. is currently therefore This finding 1B). Figure Article petrels:2.2%, 3.3%; Trindade Christmas Island 3.4%, Islands Marquesas 5.6%, Islands Pitcairn Phoenix petrels: 3.6%; Atoll petrels: Ducie Herald 6.9%; Rapa Iti 7.1%, Islands Fernández Juan petrels: (Kermadec Ocean Indian the outside and Island (8.1%) Round on are occurring hybrids way three- possible but Pterodroma, tropical of populations the widespreadbetween hybrids inter-species are only Here, not 2006). (McCarthy cage birds and falcons, aspheasants, such birds captive-bred in reported aremore commonly hybrids three-way Avian 2006). (McCarthy hummingbirds and 1965) & Harrison (Harrison ducks describedin anecdotally birds, although in documented are rarely hybrids (compound) three-way occurring Naturally 2013). et al. Tennyson 1997; et al. Moore 2001; et al Moore McCarthy 2006; 2001; et al. Holdaway 2015; et al. (Brown individuals single sizesor smallsample on based are these and species, Procellariform between other hybridisation two-way Our results provide evidence of gene flow between three oceans in Pterodroma petrels, Pterodroma oceansin three flowbetween gene evidence of results provide Our ) (Tennyson et al. 2013). Indeed there are few published examples of examples there published arefew Indeed 2013). et al. (Tennyson (Birdlife International 2016a) is based on its is based on 2016a) International (Birdlife

. This article isThis article by protected copyright. All rights reserved. Accepted declare. interest to of conflict no have authors The 1230649). reference: NERC,UK (Award by was supported Jones Booth Katherine NE/H5081500). (UK) (Grant Council Research NaturalEnvironmental the by supported wasprogramme tracking petrel The technicalsupport. provided RachelTucker and Krupa Andy NBAF844. and grants NBAF446 under Sheffield of University at the (NBAF) Facility Analysis Biomolecular (NERC) Council Research Environment Natural at the was performed genotyping and Microsatellite development manuscript. this improved which comments, helpful their for anonymous reviewers and thank subject to editor the also like We would genetic analyses. on advice invaluable their for Wang Jinliang and Burke Terry to also Thanks DNAextraction. museum sample with helping for Gottelli samples Dada petrel and of transport and with the organisation helping for Simon Island Tollington We thank petrel). Brown(Round Ruth by provided data werekindly associated and samples blood Additional geolocators.the of on use valuable insight provided Survey) Article Antarctic Ratcliffe (British Norman Island. Round on programme research petrel the facilitating HelenGath in and Rouse Lucy PatBanville, Tom Churchyard, Baxter, Richard Goder, Martine Zuel, Vimul Nicolas to thanks Nundlaul, particular, sampling. In blood and geolocators fitting assistance with data, ringing providing in assistance Service their for Conservation and Parks National Acknowledgements efforts. conservation for information valuable provide also Pacificmay the in studies concentrating tracking and genetic similar and Trindade petrel, foraccount the into taken be should Island, Round namely locations, other species in Mauritian Government the and and volunteers staff Foundation Wildlife the Mauritian We thank

This article isThis article by protected copyright. All rights reserved. by speciation of evidence and genetics conservation structure, Genetic MC(2003) Double CL, Abbott References Ando H, Kaneko S, Suzuki H, et al. (2011) Lack of genetic differentiation among subpopulations of of amongsubpopulations geneticof differentiation (2011) Lack al. et Suzuki H, S, H, Kaneko Ando Austin JJ, White RW, Ovenden JR (1994) Population-genetic structure of a philopatric, colonially colonially aphilopatric, of structure Population-genetic JR(1994) Ovenden RW, JJ,White Austin Baker C, Slade R, Bannister J, et al. (1994) Hierarchical structure of mitochondrial DNA gene flow flow DNAgene mitochondrial of structure (1994) Hierarchical al. et J, R,Bannister Slade Baker C, Avise JC, Nelson WS, Bowen BW, Walker D (2000) Phylogeography of colonially nesting seabirds, nesting colonially of Phylogeography D(2000) Walker BowenBW, WS, Nelson Avise JC, Bicknell A, Knight M, Bilton D, et al. (2012) Population genetic structure and long and structure genetic (2012) Population al. et D, M, Bilton Knight A, Bicknell Beerli P, Felsenstein J (2001) Maximum likelihood estimation of a migration matrix and effective and matrix amigration of estimation likelihood Maximum J(2001) Felsenstein BeerliP, Beerli P (2009) How to use MIGRATE or why are Markov chain Monte Carlo programs difficult to to programs Carlo difficult chain Monte areMarkov why or use MIGRATE to How BeerliP (2009) BirdLife International (2016b) Species factsheet: Pterodroma heraldica. http://www.birdlife.org heraldica. Pterodroma Species (2016b) International BirdLife factsheet: IUCN(2016a) International Birdlife Red http://www.birdlife.org International, Birdlife List birds. for The IUCN phaeopygia. RedList Speciesof Pterodroma (2015) International BirdLife Threatened Blower DC, Pandolfi JM, Bruce BD, Gomez-Cabrera MdC, Ovenden JR (2012) Population genetics Population JR(2012) Ovenden MdC, Bruce BD,Gomez-Cabrera JM, Pandolfi DC, Blower Brown RM, Techow NM, Wood AG, Phillips RA (2015) Hybridization and Back-Crossing in Giant Giant in Back-Crossing and Hybridization RA(2015) AG,Phillips Wood NM, Techow RM, Brown Brown RM, Jordan WC,Jordan Brown RM, CG,Jones Faulkes V,Palma BugoniL,Tatayah RL,Nichols Across of World the World) the Families (Bird Petrels Oxford Albatrosses and (2004) MD Brooke Burg T, Croxall J (2001) Global relationships amongst black amongst Globalrelationships Croxall J(2001) T, Burg Brown RM, Nichols RA,FaulkesNichols CG,Jones V,GottelliBrown RM, BugoniL,Tatayah D,Jordan Accepted Stattersfield B, Lascelles SHM, Butchart JP, Croxall, Article range expansion in shy and white-capped albatrosses. Molecular Ecology Molecular albatrosses. white-capped shyand in expansion range the black-footed albatross on the Bonin Islands. Journal of Zoology Journal Islands. Bonin the on albatross black-footed the nesting seabird, the Short-tailed Shearwater (Puffinus tenuirostris). The Auk (Puffinus Shearwater Short-tailed the seabird, nesting tenuirostris). among humpback whales Megaptera novaeangliae, world novaeangliae, Megaptera humpback whales among Molecular Ecology Molecular fuscata). tern(Sterna sooty the in matrilineal patterns global to special reference with National AcademyNational of Scien of the Proceedings acoalescent approach. byusing sizes subpopulations n population in use. Population genetics for animal conservation conservation for animal genetics Population use. 327. http://www.birdlife.org http://www.birdlife.org e.T22698020A79009412.2015: and low effective population sizes. Marine Ecology Progress Series Series Ecology Progress Marine sizes. population effective low and events dispersal transoceanic structure, spatial fine-scale reveals sharks white Australian of http://www.birdlife.org 21 Ecology Molecular persistence. for colony Implications populations: seabird among dispersal of Hybridization in Seabirds. Plos One One Plos in Seabirds. Hybridization of a Summary and Island,Georgia, South atBird M. and halli) giganteus Petrels (Macronectes Secondary Contact and Hybridization. Plos One One Plos Hybridization. Contact and Secondary RA Press, Oxford. University WC evidence from microsatellite genotypes. Molecular Ecology Molecular genotypes. microsatellite fromevidence Seabird Conservation Status, Threats and Priority Actions: a Global Assessment. Bird Bird Assessment. Actions: aGlobal Priority and Status, Threats Conservation Seabird Ecology Molecular microsatellites. DNA mitochondrial and structure using population of analysis , 2863-2876. , 2863-2876.

(2011) Phylogenetic Relationships in Pterodroma Petrels Are Obscured by Recent by Obscured Petrels Are Pterodroma in Relationships Phylogenetic (2011)

(2010) Range expansion and hybridization in Round Island petrels (Pterodroma spp.): spp.): petrels (Pterodroma Round Island in and hybridization expansion Range (2010) 10 , 2647-2660. , 2647-2660. 9 , 1783- ces 1792.

98 , 4563-4568. , 4563-4568.

10 , e0121688. , e0121688. AJ, 6 Sullivan B, Symes A, Taylor P (2012) P (2012) Taylor A, SymesB, Sullivan 17 , e20350. , e20350. , 42- ┽

browed and grey and browed 79. ┽ wide. Molecular Ecology Molecular wide.

19 , 3157-3170. , 3157-3170. 283 455 ┽ , 28-36. , 28-36. , 229-244. , 229-244. headed albatrosses: albatrosses: headed 12 ┽ , 2953-2962. , 2953-2962. 111 distance distance , 70- 79 3 , 313- . . Ellegren H, Pr Ellegren Grant PR, Grant BR, Keller LF, Markert JA, Petren K (2003) Inbreeding and interbreeding in in interbreeding and Inbreeding (2003) K Petren JA, Markert LF, BR, Keller Grant PR, Grant Grewe PM, Feutry P, Hill PL, et al. (2015) Evidence of discrete yellowfin tuna (Thunnus tuna discrete yellowfin of (2015) Evidence albacares al. et Hill PL, P, Feutry PM, Grewe Grant PR, Grant R (1992) Hybridization of bird species. Science species. bird of Hybridization R (1992) Grant PR, Grant Falush D, Stephens M, Pritchard J (2003) Inference of population structure using multilocus genotype multilocus genotype structure using population Inferenceof J(2003) Pritchard M, D, Stephens Falush movements Inter-island UG (2003) Mueller Adams RMM, EA, Schreiber AD, Anders DC, Dearborn Gelter H, Tegelström H, Gustafsson L (1992) Evidence from hatching success and DNA success and Evidence hatching from (1992) L Gustafsson H, Gelter H,Tegelström 2010. for Plan Action Birds Australian G(2011) J, Dutson Szabo Garnett S, Canberra: 2000.[Online]. Birds Australian for Plan Action The G(2000) Crowley Garnett S, Harrison JM, Harrison JG (1965) A presumed trigen duck involving mallard, pintail and gadwall. gadwall. and pintail mallard, involving duck A trigen presumed JG(1965) Harrison JM, Harrison Friesen VL, Burg TM, McCoy KD (2007) Mechanisms of population differentiation in seabirds. seabirds. in differentiation population of Mechanisms KD(2007) McCoy TM, Burg Friesen VL, Gómez-Díaz E, González-Solís J, Peinado M (2009) Population structure in a highly seabird, pelagic a highly in structure Population (2009) M Peinado J, González-Solís E, Gómez-Díaz Goudet J, Perrin N, Waser P (2002) Tests for Testsfor P (2002) N, Waser J, Perrin Goudet indices(version fixation and diversities testgene estimate and program to a FSTAT, J (2001) Goudet Friesen, VL, WS Molecular Friesen, RT, (1993) Barrett, Montevecchi,Davidson, WA and Bayesian multilocus a of performance the Evaluating OE (2007) Gaggiotti RS, Waples P, Faubet ID,Bacon B, Bird Hansson KL, AD,Durrant IR,Ball C,Stewart GJ, Küpper Horsburgh Dawson DA, Gottelli D, Sillero-Zubiri C, Applebaum GD, et al. (1994) Molecular genetics of the most endangered mostendangered the of genetics Molecular (1994) al. et GD, C, Applebaum Sillero-Zubiri D, Gottelli Friesen V (2015) Speciation in seabirds: why are there so many species…and w species…and so many there are seabirds: why in Speciation (2015) V Friesen bat among structure Spatialgenetic (2015) SV CKR,Good Willis F, Martinez-Nunez CM, Davy This article isThis article by protected copyright. All rights reserved. Accepted Article highly among flow andhistorical gene contemporary of Similarity H(2010) Gibbs JE, Chiucchi Darwin's finches. Evolution finches. Darwin's Reports Reports Scientific resource. important globally this management for of demands rethink populations data: Linked loci and correlated allele frequencies. Genetics Genetics allelefrequencies. correlated lociand data: Linked Ecology Molecular a seabird. in pelagic differentiation population and Ecology Resources utility cross-species primerhigh setsof develop and loci microsatellite identifyconserved to methods New (2010) AP A, Krupa Klein S, Genetics genetic markers. Molecular Ecology Molecular markers. genetic albicollis hypoleuca Ficedula flycatchers pied× collared hybrid fertility of the for fingerprinting Australia. Birds Australia and Environment Molecular Ecology Molecular ecology. Marine Ecology Progress Series Series Ecology Progress Marine ecology. shearwater Cory’s the 2.9. 3). 2.9. hybrid and a male Common Murre. Canadian journal of zoology journal andamaleCanadian hybrid Common Murre. Murre identification between afemale ofabackcross Thick-billed Common Murre× Ecology Molecular rates. migration of theestimation for method canid: the Ethiopian wolf Canis simensis. Molecular Ecology Molecular Caniswolf simensis. Ethiopian the canid: 16 Genetics Conservation pathogen. spreading arapidly of edge the leading along hibernacula Journal of Ornithology Journal Conservation International International Conservation fra , 1013-1024. , 1013-1024. gmented populations of an endangered rattlesnake. Molecular Ecology Molecular rattlesnake. of an endangered populations gmented immer C, Sheldon B (1995) Microsatellite ‘evolution’: directionality or bias? bias? or directionality ‘evolution’: Microsatellite B(1995) C,Sheldon immer 5 . 11 , 1 Ibis , 360-362. , 360-362. 6916. 6916. 134 , 62-68. 10 16 , 475- , 1765-1785. , 1765-1785. Calonectris diomedea: an examination of genetics, morphology and and morphology genetics, of examination an diomedea: Calonectris 156 , 27- 22 494. 57 , , 2911-2916. , 2911-2916. 1 39

– 34. . 11 sex , 1103-1114. , 1103-1114. ┽ 382 biased dispersal using bi using dispersal biased , 197-209. , 197-209. –

as demonstrated for birds. Molecular Molecular birds. for demonstrated

256 164 , 193. , 193. 3 , 301- , 1567-1587. , 1567-1587. 16 ┽ , 1149-1166. , 1149-1166. 312. parentally inherited parentally 12 71 hy aren’t there more? there aren’t hy

, 2835-2843. , 2835-2843. 19 ,1474-1477. ,1474-1477. , 5345-5358. , 5345-5358. Nature Nature ×

) Helbig AJ, Seibold I, Kocum A, et al. (2005) Genetic differentiation and betweenand hybridization Genetic differentiation (2005) et al. Kocum I, A, AJ,Seibold Helbig King B, Reimer D (1991) Breeding and Behavior of the Herald Petrel Pterodroma arminjoniana arminjoniana Pterodroma Herald Petrel of the Behavior and Breeding Reimer B, D(1991) King Kalinowski ST, Taper ML, Marshall TC (2007) Revising how the computer program CERVUS CERVUS computer program Revising howthe (2007) TC Marshall ML, Taper ST, Kalinowski This article isThis article by protected copyright. All rights reserved. Holdaway RN, Worthy TH, Tennyson AJ (2001) A working list of breeding bird species of the New the species of bird list of breeding A working AJ (2001) TH, Tennyson Worthy RN, Holdaway Hubbs CL (1955) Hybridization between fish species in nature. Systematic nature. in species fish between Hybridization CL (1955) Hubbs zoology Küpper C, Horsburgh GJ, Dawson DA, et al. (2007) Characterization of 36 polymorphic of 36 polymorphic Characterization (2007) et al. GJ, Dawson DA, C, Horsburgh Küpper Marie AD, Bernatchez L, Garant D (2011) Empirical assessment of software efficiency and accuracy accuracy and efficiency software of assessment Empirical Garant D(2011) L, Marie AD,Bernatchez Moore PJ, Taylor GA, Amey JM (1997) Interbreeding of black-browed albatross Diomedea m.Diomedea albatross black-browed of Interbreeding (1997) JM GA,Amey Taylor PJ, Moore of Black-browed ratio sex and Provenance CD(2001) GA,Millar Taylor Burg TM, PJ, Moore Kaeuffer R, Reale D, Coltman DW, Pontier D (2007) Detecting population structure using structureusing population Detecting D(2007) Pontier DW, D, Coltman Reale Kaeuffer R, McCracken KG, Wilson RE (2011) Gene Flow and Hybridization between Numerically Imbalanced Imbalanced between Numerically Hybridization Geneand Flow (2011) RE KG, Wilson McCracken of hybrids ofHandbook EM(2006) Press. McCarthy University Oxford world the avian Evol Ecol Trends the genome. of invasion asan Hybridization Mallet J(2005) Morris-Pocock JA, Steeves TE, Estela FA, Anderson DJ, Friesen VL (2010) Comparative (2010) VL DJ,Friesen FA,Anderson Estela TE, JA, Steeves Morris-Pocock and genetic structure dispersal, paradox: Theseabird (2008) Bernatchez L Weimerskirch E, H, Milot Imber M (2004) Kermadec petrels (Pterodroma neglecta) at Ilha da Trindade, South Atlantic Atlantic Ocean South Trindade, da atIlha neglecta) petrels(Pterodroma Kermadec (2004) ImberM S(1982)Emu breedingHunter Interspecific ingiant petrels at Georgia. South Meirmans PG (2014) Nonconvergence in Bayesian estimation of migration rates. Molecular Ecology Molecular rates. of migration estimation Bayesian in Nonconvergence (2014) Meirmans PG Jakobsson M, Edge MD, Rosenberg NA (2013) The Relationship Between F(ST) and the Frequency Frequency the and BetweenF(ST) The Relationship NA (2013) Rosenberg MD, M, Edge Jakobsson ocean. Atlantic North Azores, the off neglecta) Petrels(Pterodroma Kermadec ImberMJ (2008) Accepted Article Imber of (2004). rebuttal Tove’s M. A to response (2005) ImberM greater and lesser spotted eagles (Accipitriformes: Aquila (Accipitriformes: eagles lesserspotted and greater clanga Ecology Molecular assignment. paternity in success increases error genotyping accommodates Raine Island, Queensland. Emu Queensland. Raine Island, Zealand region at first human contact. New Zealand Journal of ZoologyNew contact. human at first region Zealand Journal Zealand Ornithology loci and their amplification in four other Charadrius species. Molecular Ecology NotesMolecular species. Charadrius other four in amplification their and loci sex-linked two including alexandrinus) (Charadrius Kentish plover the in loci microsatellite Bulletin of Club The Ornithologists' British Conservation Genetics Genetics Conservation charr(Salvelinus brook in scenarios stocking variable under introgression detect to fontinalis). Emu-Austral Ornithology D. m. albatross black-browed NewZealand and Island. on Campbell impavida melanophris 329 Emu Campbell NewZealand. on breeding Island, melanophrys, Thalassarche Albatross, STRUCTURE software: effect of background linkage disequilibrium. Heredity Heredity linkage disequilibrium. background of effect software: STRUCTURE Populations of Two Duck Species in the Falkland Islands. Plos One Plos Islands. Falkland the in Species Duck ofTwo Populations Phylogenetics and Evolution and Phylogenetics Molecular seabirds. pelagic in flow gene preferenceon habitat barriersand physical of The influence boobies (S.red-footed and sula): leucogaster) (Sula of brown phylogeography 17 Ecology Molecular species. philopatric albatross but mobile, dynamics a highly in population and in the North Atlantic. Notornis Notornis North Atlantic. the in and 39. Resources of the Most Frequent Allele. Genetics Allele. Frequent Most the of Notornis , 1658-1673. , 1658-1673.

-334. 16 55 14 , 1099-1106. , 1099-1106. , 106-108. , 106-108. 146 , 726- , 226-234. , 226-234. 733. 12

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, 515-528. , 515-528. 40. 85

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Notornis Notornis , A. pomarina). Journal of Journal A. pomarina). 6 52 , e2317 20 , 58- 28 , 229- , 119- 4 59. , 1 3. 82

237. - 99 , 312-314. , 312-314. 20. 187. , 374-380. , 374-380.

7 , 35- 101 on

, R Development Core Team (2013) R: A language and environment for statistical computing. environmentstatistical for and R: A (2013) Team Core R Development language Randler C (2002) Avian hybridization, mixed pairing and female choice. Animal female and choice. mixed pairing hybridization, Avian C(2002) Randler Behaviour Svedin N, Wiley C, Veen T, Gustafsson L, Qvarnström A (2008) Natural and sexual selection against against selection sexual Natural and A (2008) Qvarnström L, Veen C, T,Gustafsson Wiley N, Svedin Portnoy DS, McDowell JR, Heist EJ, Musick JA, Graves JE (2010) World phylogeography and and phylogeography World Graves JE(2010) JA, Musick HeistEJ, JR, DS, McDowell Portnoy Nicoll MAC, Nevoux M, Jones CG, et al. (2016) Contrasting effects of tropical cyclones on th on cyclones tropical of effects Contrasting (2016) al. et Jones CG, M, Nevoux MAC, Nicoll Rhymer JM, Simberloff D (1996) Extinction by Hybridization and Introgression. Annual Introgression. and by Hybridization Extinction D(1996) ReviewSimberloff JM, Rhymer of Ibis birds. in hybridization natural of correlates ecological and Behavioural C(2006) Randler Seehausen O (2006) Conservation: Losing Biodiversity by Reverse Speciation. Current Biology Current ReverseSpeciation. by Biodiversity Losing Conservation: (2006) O Seehausen Pritchard J, Stephens M, Donnelly P (2000) Inference of population structure using multilocus multilocus using structure population of (2000) Inference P Donnelly M, Stephens J, Pritchard Schluter D (2009) Evidence for Ecological Speciation and Its Alternative. Science ItsAlternative. and Speciation Ecological for Evidence D(2009) Schluter a in connectivity recentgenetic estimating of problem The (2016) F Rodd B, Shuter Samarasin P, Rannala B (2012) BayesAss edition 3.0 manual. 3.0 user’s BayesAss edition B(2012) Rannala Taylor SA,Anderson DJandFriesenTaylor for strong Evidence assortative VL mating, (2012) (Sula SA,Zavalaga CBTaylor andFriesen Hybridization VL between Blue-Footed (2010) Mauritius. of University biology The RVV(2010) Tatayah of breeding Petrel, Round the Island Slate J, Hale MC, Birkhead TR (2007) Simple sequence repeats in zebra finch (Taeniopygia guttata (Taeniopygia zebrafinch repeats in sequence SimpleTR (2007) Birkhead MC, Hale Slate J, Steeves TE, Anderson DJ, Friesen VL (2005) A role for nonphysical barriers to gene flow in the in flow gene to barriers nonphysical Arolefor VL (2005) DJ, Friesen Anderson Steeves TE, Slatkin M (1987) Gene flow and the geographic structure of natural populations. Science natural populations. of structure geographic the (1987) Geneflow and M Slatkin This article isThis article by protected copyright. All rights reserved. Accepted Article females hybrid do revisited: Haldane's rule (2014) M Zagalska-Neubauer P, Nowicki G, Neubauer white- between Hybridization (2007) MD JJ, Sorenson GreenAJ, Negro V,VilàC, Muñoz-Fuentes Foundation for Statistical Computing, Vienna, Austria. http://www.R-project.org/ http://www.R-project.org/ Austria. Vienna, for Computing, Statistical Foundation hybrid flycatchers. Proceedings of the Royal SocietyProceedings flycatchers. hybrid SciencesBiological B: male Ecology Molecular structure. population weak resolving for implications chalcogramma): (Theragra pollock walleye fish, marine the polymorphism in microsatellite between and F Inverserelationship P (2004) KM, Bentzen Bailey MF, Canino PT, O'Reilly Ecology and SystematicsEcology and annual survival of a pelagic seabird in the Indian Ocean. Global Change Biology Change Global Ocean. Indian the in seabird apelagic of survival annual 119. genotype data. Genetics data. genotype changing world. Conservation Biology Conservation world. changing 459 hybrid zone in northern Peru. Journal of Avian zoneinnorthern Journal Biologyhybrid Peru. limited geneflow, differentiation and strong across theblue 251-257. Waterbirds Northern Peru. Boobiesin nebouxii)(Sula variegate) andPeruvian R334-R337. R334-R337. Ecology Molecular dactylatra). (Sula booby masked the seabird, vagile ahighly of diversification 792. Genomics BMCpasserines. studies of genetic evolutionary for new resource tags:a expressed sequence 19 species. Journal of Evolutionary Biology species. Journal large gull betweentwo zone arecentcontact in hybrids of Survival lifespan? ashorter have Ecology Molecular Spain. in ducks ruddy introduced and ducks headed , 1994-2010. , 1994-2010. -467.

┽ mediated gene flow in the sandbar shark, Carcharhinus plumbeus Carcharhinus sandbar shark, the in flow gene mediated 14

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43 , 311-324. 27 ┽ . footed/Peruvian booby 16 . 13 Molecular Ecology Molecular , 629- 3

23 , 1799-1814. , 1799-1814. 275 , 737-741. , 737-741. , 735. , 735. 638.

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, 787- 550-565 R 148 e 33 , 103- 16 , , , ) . Vähä JP, Primmer CR (2006) Efficiency of modelof Efficiency CR(2006) Primmer Vähä JP, Welch A, Fleischer R, James H, et al. (2012) Population divergence and gene flow in an endangered an in flow gene and divergence Population (2012) al. et H, James R, A, Fleischer Welch http://www.seabirdtracking.org/ database: OceanWanderers Tracking International BirdLife the via data areavailable Tracking - http://dx.doi.org/10.5061/dryad.fg86p Digital Repository: Dryad the from data areavailable genotyping Microsatellite - CK307697). DV948691, DV575298, LT608740, LT608738, LT608733, LT608720, LT608717, LT608694, LT608692, LT608685, AM072445-AM072447, numbers: (Accession - Accessibility Data between divergence genetic and segregation (2012) Foraging al. et P, Welch A, Ostrom A, Wiley (Pterodroma Petrel Hawaiian Genetics of Endangered the Conservation AJ (2011) Welch This article isThis article by protected copyright. All rights reserved. To soft- that suggests petrel gadfly A hybrid (2013) GA,MJ Imber HA,Taylor AJD,Lawrence Tennyson genotypes. Genetics genotypes. multilocus rates using migration recent of inference Bayesian B(2003) GA, Rannala Wilson table. pop-up from resulting the 1326 selecting ont clicking and then arminjoniana Pterodroma species the The microsatellite sequences isolated are available via the ENA-EBI EMBL sequence database sequence EMBL ENA-EBI the via available are isolated sequences microsatellite The

Accepted Article Ocean Atlantic the in neglecta) Petrels(Pterodroma Kermadec (2005) ve MH Molecular Ecology Molecular numbers loci. of different with and scenarios hybridization different under individuals and highly mobile seabird. Heredity mobile seabird. highly and geographically proximate colonies of a highly mobile seabird. Oecologia Oecologia mobile seabird. a highly of proximate colonies geographically Time, Across sandwichensis) and Space USA. Maryland, of University dissertation, Doctoral Notornis Notornis mollis) the 1920s. by Islands the Antipodes hadcolonised (Pterodroma petrels plumaged 52 , 56- 58. 60

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109 , 19-28. , 19-28. ┽ based Bayesian methods for detecting hybrid hybrid detecting for methods Bayesian based he datasets ‘view list’ button and button list’ datasetshe ‘view

– 168 a rebuttal. Notornis Notornis a rebuttal. , 119- 130.

This article isThis article by protected copyright. All rights reserved. Accepted manuscript. the on comments provided samples and andmuseum blood collected geolocators, recovered and deployed experiment, the KNdevised and (NPCS) Service Conservation and National Parks the through Island petrels Round data on and Island Round access to KR facilitated MauritianFoundation; Wildlife the through Islandpetrels Round data on and Island Round to access facilitated CJ VT and History; Natural of Museum American at the skinpetrel collection SouthSeas Expedition Whitney the to facilitated access PS manuscript; Article the on comments provided and Kermadec Islands the petrelsof samples from Kermadec blood provided SMHI applications; NERCfunding of the development advisedon and work genetic and library microsatellite initial with assisted JJG devised experiment, ; the genotyping and development marker supported and sequences microsatellite the GJHisolated manuscript; the on comments and data analysis on guidance HHprovided manuscript, the comments on provided and analyses, and and genotyping validation marker development, on guidance DADprovided Information; Supporting the co-wrote and markers microsatellite validated the and CR characterised manuscript; the comments on provided samples and museum and blood collected geolocators, recovered and deployed experiment, the MACN devised manuscript; the wrote and all analyses genotyping, microsatellite the andmuseum performed samples, blood collected geolocators, KABJ deployed Contributions: Author

Accepted isThis article by protected copyright. All rights reserved. neglecta) (Pterodroma Kermadec petrel heraldica) (Pterodroma Herald petrel arminjoniana) (Pterodroma petrel Trindade Article Unknown species Putative 1 Table Figures and Tables : Number and origin of samples. N of origin and : Number

Kermadec Islands Islands Kermadec Kermadec Islands Islands Fernandez Juan Ducie Atoll Island Oeno Islands Marquesas Ducie Atoll Islands Trindade Island Round location Geographic S = Number of samples, N samples, of = Number

G = Number genotyped at >75% of 12 microsatellite markers. markers. 12 microsatellite of >75% at genotyped = Number Pacific Ocean Pacific Ocean Pacific Ocean Pacific Ocean Pacific Ocean Pacific Ocean Pa Pacific Ocean Ocean Atlantic Ocean Indian Region cific Ocean cific Ocean 41 29 30 30 21 23 30 52 561 N S

blood blood museum museum museum museum museum museum blood blood Type 24 29 28 25 21 23 28 45 484 N G

Islands Phoenix Islands Marquesas Islands Pitcairn (Kiritimati) Island Christmas (BassIslands) Island Oeno Island Marotiri (BassIslands) Rapa Island

Pacific Ocean Pacific Ocean Pacific Ocean Pacific Ocean Pacific Ocean Pacific Ocean Pacific Ocean 30 1001 16 29 21 30 30 28

museum museum museum museum museum museum museum 885 16 29 18 30 30 26 29

This article isThis article by protected copyright. All rights reserved. Mean Tgu06 TG13- TG03- Phel Phel33 Phel28 Phel15 Phel12 Parm31 Parm29 Parm22 Calex01 Locus 2 Table individuals from the native range of the species, and not from the Round Island population. Size: allele size range, N Size: range, allelesize population. Island from Round the not the species, and of range from native the individuals H O

Accepted Article heterozygosity, : observed 35

009 002

: Characteristics of the 12 microsatellite markers genotyped in each of the potential species found at Round Island. Each species group only includes includes only group species Island. Each atRound speciesfound thepotential eachof in genotyped markers microsatellite 12 the of Characteristics

152 185 123 124 116 230 157 199 229 130 91 86 Size – – – – – – – – – – – All spp. -146 107 169 199 129 182 164 254 177 207 244 90

H 7.83 7.83

N 13 10 11 11 8 8 3 9 6 6 8 5 5 4 E A : expected heterozygosity, P-value: P-value for Hardy-Weinberg test. for Hardy-Weinberg P-value P-value: heterozygosity, : expected

3.92 3.92 N 5 5 3 3 9 9 5 1 5 2 1 2 2 A

3.45 3.45 4.75 2.50 2.43 7.67 7.85 3.75 1.00 4.46 1.95 1.00 2.00 2.00 Ar Trindade petrel Trindade

0.36 0.36 0.51 0.27 0.15 0.83 0.81 0.67 0.00 0.40 0.13 0.00 0.21 0.35 H O

0.39 0.39 0.63 0.24 0.14 0.86 0.84 0.58 0.00 0.63 0.19 0.00 0.22 0.40 H

P-value <0.01 <0.01 0.04 0.04 1.00 1.00 0.58 0.59 0.65 0.06 0.52 0.02 - - - -

5.17 5.17 N 10 9 9 4 3 9 5 2 6 3 4 4 3 A

3.71 3.71 4.51 2.76 1.97 6.44 7.89 3.43 1.37 5.61 2.81 2.71 2.87 2.20 Ar

0.34 0.34 0.69 0.14 0.10 0.75 0.83 0.17 0.03 0.68 0.21 0.19 0.10 0.15 Herald petrel H O

0.36 0.36 0.61 0.16 0.09 0.77 0.86 0.28 0.03 0.76 0.26 0.22 0.17 0.17

H E

P-value

<0.01 <0.01 <0.01 <0.01 0.43 0.43 0.06 1.00 0.09 0.01 1.00 0.01 0.02 0.06 - -

A : number of alleles, Ar: allele richness, allele richness, Ar: alleles, of : number

5.58 5.58

N 11 7 7 4 3 8 5 3 6 7 3 5 5 A

3.82 3.82 4.23 1.94 2.47 6.98 8.14 3.63 2.66 3.78 3.65 1.62 4.14 2.64 Ar

Kermadec petrel Kermadec 0.35 0.35 0.47 0.05 0.39 0.82 0.71 0.30 0.22 0.28 0.29 0.05 0.49 0.12 H O

0.40 0.40 0.57 0.08 0.37 0.83 0.74 0.47 0.22 0.33 0.30 0.05 0.68 0.13 H E

P-value <0.01 <0.01 <0.01 <0.01 0.01 0.01 0.92 0.08 0.01 0.87 0.04 0.13 1.00 0.02

- -

009 002 continued. continued.

152 185 123 124 116 230 157 199 229 130 91 86 Size – – – – – – – – – – – - 107 – 1 199 129 182 164 254 177 207 244 90 All spp. 46 69

7.83 7.83 N 11 11 13 10 5 5 4 8 3 9 6 6 8 A

4.67 4.67 N 7 7 3 3 8 8 4 1 6 7 3 3 3 A

3.81 3.81 4.87 1.91 2.27 7.41 5.42 3.57 1.00 4.38 6.63 2.69 3.00 2.61 Ar

Murphy's petrel Murphy's 0.41 0.41 0.70 0.07 0.29 0.88 0.79 0.54 0.00 0.50 0.77 0.21 0.00 0.23 H O

0.49 0.49 0.66 0.07 0.30 0.84 0.72 0.63 0.00 0.64 0.81 0.20 0.61 0.40 H E

P-value <0.01 <0.01 <0.01 <0.01 0.84 0.84 1.00 0.70 0.02 0.69 0.06 0.12 1.00 - - -

4.83 4.83 N 11 5 5 3 4 8 3 3 7 4 6 2 2 A

3.58 3.58 3.74 2.26 3.01 5.93 7.49 2.65 1.82 5.65 3.39 4.39 1.43 1.19 Ar

0.38 0.38 0.45 0.10 0.28 0.76 0.83 0.19 0.05 0.79 0.52 0.53 0.03 0.01 H Phoenix petrel Phoenix O

0.39 0.39 0.53 0.12 0.25 0.76 0.83 0.22 0.06 0.78 0.46 0.57 0.03 0.01 H E

P-value <0.01 <0.01 0.25 0.25 0.01 1.00 0.09 0.34 0.05 0.07 0.89 0.18 0.04 1.00 - -

This article isThis article by protected copyright. All rights reserved. 1A Figure and island of origin (below the plot, separated by black vertical lines), with the exception of Round Island, where species is unknown. isunknown. where species Round Island, of exception the with lines), vertical black separatedby plot, the (below of origin island and a part to assigned ancestry anindividual’s of proportion the represent bar each in Accepted Article types. Kermadec and Herald/Phoenix Trindade, between hybrid three-way =two TxK type, PureM= Murphy’s type, =Herald/Phoenix PureHP type, Kermadec PureK = type, PureT e.g. =Trindade analysis, STRUCTURE using clusters identified potential four the eachof membership to estimated from derived chartis pie each in petrels of classThe assignment see S3). Table ateachlocation sample numbers (for from sample the originated source-island time and sampling at species recorded separatedby individuals represent (star). charts Island Pie Round and Martim Islands, and Vaz =Trindade TI Islands, Fernández =Juan Ducie, JFI Islands- = Pitcairn (D) PI Oeno, Islands- (O)=Pitcairn PI all, Islands, =Pitcairn PI Islands, =Marquesas MI Marotiri, = BassIslands- (M) BI Rapa Island, Islands- Bass = (R) BI Islands, Islands, CI =Christmas =Phoenix PhI Kermadec Islands, east, Westto = KI sampled. ofislands distribution : Plot showing STRUCTURE analysis results, where K (number of clusters) equalled 4. Each vertical bar represents a single individual and colours colours and K individual where results, asingle STRUCTURE analysis showing vertical represents Each Plot bar 4. equalled clusters) of (number

icular cluster. Individuals are grouped by grouped are Individuals cluster. icular -way hybrid between Trindade type and Kermadec type, TxHPxK= type, and Kermadec type between Trindade hybrid -way putative species (above the plot) plot) the species(above putative B: The global The Accepted isThis article by protected copyright. All rights reserved. Islands. =Pitcairn grey light = Islands, grey Marquesas group: dark from island same populations the ovals represent background Grey 1. Figure are thesameasin abbreviations Island label. island and species a a circle, with by species isrepresented a of population island Each betweenoceans. indicatelines divides of movement. Dotted direction the represent Black arrows per generation. population petrel Trindade originates the from population Island the Round ± (18%) example, For 0.183 of 0.036 (2012)). Rannala asin deviation, standard the x 2 Figure Article : Migration rates (proportion of migrants from population x in population y per generation) between island populations, ± confidence intervals (1.96 (1.96 intervals ±populations, island between confidence generation) per population in y x of frommigrants rates (proportion population : Migration

View publication stats Kermadec petrel colony to Round Island. Island. Round to colony Kermadec petrel Accepted isThis article by protected copyright. All rights reserved. 2006 IslandRound between on breeding wasfound it subsequently before 1984 in adult asabreeding was ringed 061-39302 petrel where Herald (Australia), Raine Island = Diamond petrel. Trindade of the colony known other only (Brazil), the =Trindade Islands square Black Island. Star =Round 18/08/2010. - (male) between19/02/2010 from 5H41524 petrel locations Crosses represent 11/03/2013. - 02/10/2012 between (female) 5H41919 from petrel represent locations points circular Black Pacific Oceans. theAtlanticand migrated into Ocean and 3 Figure Article : Recorded movement of individuals between oceans. Two individual petrels fitted with geolocators that departed from Round Island in thein Indian Island from departed Round that geolocators petrels with fitted individual Two oceans. between individuals of movement : Recorded

– 2012. Triangle = Lord Howe Island, the closest known Pacific closest known the Island, Howe = Lord 2012. Triangle