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Article: Williams, R.C., Jackson, B.C., Duvaux, L. et al. (3 more authors) (2015) The genetic structure of Nautilus pompilius populations surrounding Australia and the Philippines. Molecular Ecology. Published Online 1 June 2015. https://doi.org/10.1111/mec.13255

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[email protected] https://eprints.whiterose.ac.uk/ Accepted Article

Burke Rachel C.Williams* † [email protected] UK. Email: forWildlifeConservati Centre Williams, * Rachel 3 2 1 of The structure genetic Article Original : type Article Accepted 27-May-2015 : Date Revised: 21-May-2015 Date Received 18-Feb-2015 : Date This isarticle by protected copyright.reserved. Allrights 10.1111/mec.13255 differences betweenthis version and theVersion Record.of Please cite this ar copyediting,through the paginationtypesetting, proofreadingand process, which lead may to This hasarticle been accepted publicationfor andundergone peerbutfull review not has

Population structure of structure Population title: Running structure microsatellite, population marine conservation, cephalopod, computation, Approximate Bayesian Key words: Centre for Wildlife Conservation, University of Cumbria, Ambleside LA22 9BJ, UK 9BJ, LA22 Ambleside Cumbria, of University Conservation, Wildlife for Centre These authors contributed equally equally contributed These authors Department of Animal and Plant Sciences, University of Sheffield, Department ofAnimal S10 Sheffield and 2TN,UK Sciences,Sheffield, of University Plant Sheffield, Sheffield S10 Sheffield 2TN, UK Sheffield, NERC Biomolecular Analysis Facility, Department of Animal and Plant Sciences, University of University Animal of andPlantSciences, Department Facility, Analysis Biomolecular NERC 2 , William Sinclair 1,2 Nautilus pompilius , Benjamin C. Jackson C. , Benjamin Nautilus pompilius 1

populations Australia and the surrounding Philippines 3† , Ludovic , Duvaux

on, University of Cumbria, Ambleside LA22 9BJ, 9BJ, LA22 AmblesideCumbria, of on, University 3† , Deborah A. Dawson A. Deborah , ticle as doi: 2 , Terry been Accepted Article Human access to the marine environment is no longer technologically inhibited, and defaunation of anddefaunation inhibited, technologically nolonger is environment marine the to Humanaccess of larg extinctions triggered the anthropogenically harvesting of marine animals began 42,000 years ago (O’Connor years(O’Connor agobegan 42,000 marine animals harvesting of declines (Hutchings 2000; Worm declines (Hutchings 2000; population significant are experiencing many species the world, commercial marine Throughout Introduction stocks. distinct genetically of diversity the preserving thereby can structure population Understanding classification. conservation ofinformation paucity rectifytheir inhibitingon biologythe the species’ currently facilitateif Philippine naturalrecolonization populati lack ofconnectivity thatimmigrants suggests not among west from would populations Australia size large effective population average vecan by beexplained present-day genetic similarity Instead, Australia. west and Philippines the between geneflow contemporary to due not divergence is this limited analysis that indicate computation Bayesian approximate using modelled scenarios Australian reefs, samples display a small degree of genetic structure ( structure genetic of degree asmall display samples reefs, Australian Philipp the between distances geographical the larger despite Conversely, respectively). design appropriate strategies to prevent the the prevent strategiesto appropriate design of degree andthe structure Genetic conservation. tosuccessful fundamental species is exploited in genetic of diversity distribution the Understanding Abstract This isarticle by protected copyright.reserved. Allrights microsatellite markers to evaluate the evaluate to markers microsatellite 14 we use Here, region. the in decline dramatic recent species’ the despite protection, legislative pompilius genetically isolated from Osprey Reef and Shark Reef in the Coral Sea ( Sea inthe Coral Reef and Shark Reef Osprey from isolated genetically relativePhilippines. Despite their geographical is fished unsustainably the Philippines for in andhas the shell ornamental trade limited et al.

s that persisted at both locations following their separation. The separation. their following locations at both persisted that s population structure of structure population 2006; Neubauer loss of distinct populations. populations. The losscephalopod ofdistinct e mammals seen on land (Koch & Barnosky 2006). 2006). Barnosky & (Koch land on seen emammals gene flow among populations must be assessed to to assessed be must populations among gene flow proximity, Great proximity, Barrier Reef individuals are ons These were toextinction. fished to data help ry genetic that due to has occurred limited drift allow us to facilitate sustainable harvesting, harvesting, sustainable facilitate us to allow et al.

N. pompilius et al. 2013; Watson 2011), up to 50,000 years after F around Australia and the the and Australia around ST =0.015). Demographic =0.015). F et al. et ST =0.312, 0.229, 0.229, =0.312, ines and west west and ines 2013). The Nautilus Nautilus Accepted Article times higher than extinction ratestimes extinction higher theabsencethan in (Pimm human of actions (McCauley occurring is now the oceans This isarticle by protected copyright.reserved. Allrights appreciated. isgenerally environments more than dispersal to barriers may contain marine that suggesting gene major represent to to barriers haveflow, shown 2010) alsobeen species with a high risk of extinction (Davidson extinction of ahigh risk species with rare ina marine environment (Palumbi 1994; Mirams barriers to geneflow sothat widely separatedar physical few present to considered are Oceans 1994). &Hauser stocks (Carvalho distinct genetically preserve of and the diversity extinctions local practices, prevent informsustainable harvesting used to canbe populations of structure genetic the of knowledge species, For vulnerable Sponaugle 2009). Cowen& 2006; (Levin variation genetic of through patterns shared of gene among populations flow detection the allow do moleculartechniques tracked, be cannot always individuals specific Although 200 (Pearse & genetic analyses Crandall among can population estimated populations be using have, however, been attributed to oceanic features, such as salinity (Rocha 2003; Lessios Lessios (Rocha 2003; as salinity such features, tooceanic attributed have, been however, depth and temperature (Zardi (Zardi andtemperature depth monitored using tracking devices, but success has been variable (see Semmens (seeSemmens variable been has success but devices, using monitored tracking been have populations among marine migration and movement Contemporary marineresources. strategies design effective conservation to canbe by CITES connectivityPopulation data used data for can prevent qualification CITES protection (De 2012). Angelis industrial biological information and global of absence the Unfortunately, implement (CITES 2015). is onbiological convention reliant toconstruc data CITES toensure aims the survivalspecies trade, of that not threatened and by is the international (CITES).and Flora Wild Faunaof Species Endangered in Trade on International the Convention is regulate to declines developed was that governments between agreement An international declines. et al. et 2007). Ocean fronts (Galarza Ocean(Galarza fronts 2007). et al. et 2015). Present extinction rates are probably a thousand athousand probably are rates extinction Present 2015). eas can remain connected, making absolute vicariance vicariance absolute making connected, remain can eas et al. t a specific participatingframework a for t countries to 2012) can facilitate policy changes and prevent can and policy2012) changes prevent facilitate et al. Population structure 2011). and speciation et al. et 2009) and currents (White (White currents and 2009) et al. et al. et 2014). Identifying 2014). 2007). Movement Movement 2007). et al. et 2003), 2003), et al. et for for 4).

Accepted Article associated with tracking them (Stark (Stark them tracking with associated difficulties bythe exacerbated this is and understood, however,poorly migration, Cephalopod is cuttlefish, squid), juveniles are often too small to be tagged, tag placement in adults can be difficult, canbedifficult, adults in placement betagged, tag to small too juveniles areoften squid), cuttlefish, ranges from panmixia (Lessios ranges (Lessios from panmixia that structure genetic in can result tomovement asabarrier act to potential the with features Oceanic This isarticle by protected copyright.reserved. Allrights appropriate protection measures for populations through measures populations for protection appropriate popula effect on and its migration Understanding anddecline. overexploitation the species’ toprevent mechanisms legislative appropriate al. cephalopod, cephalopod, the shelled data is connectivity population and biological lacking currently aspecies example of (Saleeffectiveconservation enable to must beresolved knowledge that scientific pr anthropogenic suggests that 2000) species (Hutchings (Neubauer species strategies and conservation thetechnology Despite (Griffiths consumer the to missold being are not catches illegal ensurethat (Grüss unit effort for per catch An in decline 80% 2012). (De Angelis are unknown recover, ability to their or populations, fishing on of effects Thelong-term trade. for shell overfished theSaunders) heavily ornamental are between, and distribution of genetic diversity among, populations (Semmens (Semmens of populations among, diversity genetic between, and distribution flowregarding gene long-term questions answer can timescales long over history demographic of beyond protected boundaries; Grüss beyondprotected boundaries; does no movement species that assess to (i.e. are areas appropriate protected marine whether evaluate to for example, conservation, areasof various in used canbe movements aspecies’ to infer 2010). The current deficiency of data on the species’ biology has inhibited the development of of the development inhibited has biology species’ onthe deficiency ofdata current 2010). The et al. et 2011). 2011). Nautilus pompilius. Nautilus pompilius. N. pompiliusN. et al. et 2013). The poor success of attempts to aid the recovery of commercial marine marine commercial of recovery the aid to attempts of success poor The 2013). during has1980–2010 beenreported from Philippine fisheries (Dunstan et al. et Nautiloids ( Nautiloids 2003) to (Baumscomplete 2003) to separation et al. et al. 2011), to trace the origins of a catch (Hobson 1999), or to or ofa catch 1999), 2011), tracethe origins (Hobson to Nautilus 2005; Semmens 2005; tion dynamics is important when deciding on on when deciding istion dynamics important available, population declines continue in most most in declines continue population available, spp. Linnaeus and spp. in-situ essure is too high and that there are gaps in gaps in are there that and high too essureis means, such as marine protected areas areas protected marine as such means, et al. et 2007). In coleoids (octopus, (octopus, In coleoids 2007). Allonautilus Allonautilus et al. et al. et et al. 2013). 2013). The inference2012). 2007). This ability 2007). This et al. et spp. Ward and spp. 2005). A key A 2005). t extend t extend et et

Accepted Article hypothesised differentiation of differentiation hypothesised and capture rates fluctuate andare often reliant on fishermen reporting This isarticle by protected copyright.reserved. Allrights Historical movement of movement Historical area. this aid progress will in and cost, devicesize decreasesin concomitant with technology, Improvements in al. be approximately 800 m (Saunders & Wehman 1977; Kanie Kanie 1977; Wehman & 800m(Saunders be approximately in which an at depth the maximum consequence, As a implosi of shell risk bylimited their is thewatercolumn in position deepest meansshell their that forprotection fr reef remainingclose to the New Guinean clade and a west Pacific clade Pacific and (Wray clade a west clade New Guinean Austra comprise a which clades, west monophyletic been about of connectivity known the current has yetand nothing management, genetic ofspecific design the andaid inform fishing sustainable (Saunders Unde & Landman 2010). relevant modern to also are constraints These temperature. andsea seas shelf adjacent between distance water depth, 1993): (Crick least barriers three biogeographical byat have distributionwill restricted nautiloid been (Wray in the Philippines population anancestral dispersal from (Sinclair (Sinclair differentiation hasoccurred genetic despite distances, and, small geographical dispersive barrier cytochrome c oxidase subunit I csubunit cytochrome oxidase morphology. Nautiloids are typically found between depths of 130 and 700 m (Dunstan m(Dunstan and700 130 of depths between are found typically Nautiloids morphology.

N. pompilius N. connectivitywill help to establish the impact that fishing is having acro 2011; Williams 2011; et al. population declines in the Philippines (Barord declines (Barord population inthe Philippines 2007), furthered by the absence of a juvenile larval stage that might aid might 2007), dispersal aid byabsence larval ajuvenile furthered stage of that the et al. Nautilus N. pompilius pompilius N. 2012). The observed population structur 2012). Theobservedpopulation and N. pompilius Allonautilus, Allonautilus, ( COI rstanding gene flow between current populations can between currentpopulations generstanding to help flow populations has been investigat populations ). Populations separated into three geographically distinct distinct threegeographically into Populations separated ). om predators. The internal arrangement of the the of arrangement internal The om predators. populations was confirmed by comparing variation at variation comparing by confirmed was populations whose movements are limited to some extent bytheir extent some to whose movements arelimited N. pompilius et al. et lian/Indonesian clade, an east Australian/Papua Australian/Papua an east clade, lian/Indonesian dividual would be encountered is considered to to considered is would be encountered dividual 1995; Sinclair 1995; et al. et al. et populations. Understanding population Understanding population populations. 2014). 2014). et al. et e was proposed to be the result of of result the tobe e wasproposed create These 1980). a limitations 1995). Historical expansions of of expansions Historical 1995). ed using molecular approaches; molecular ed using et al. et ss their distribution following catches(Sauer 2007, 2011; Bonacum 2011; 2007, et al. et et al. Nautilus Nautilus 2000). 2000). 2011), on. et et

Accepted Article buffer (1 M Tris-HCl pH M7.5; 0.5 EDTA pH 8.0; Sambrook the information deficiency that currently inhibits the legal classification of deficiency currently thelegalthe information inhibits that classification contribute towardsrectifying Philippines. These data will andthe Australia surrounding populations Here, we use toassess techniques analyses of and statistical connectivity molecular the This isarticle by protected copyright.reserved. Allrights N. pompilius pompilius N. of Agriculture in the Republic of the Philippines. Departm the from Permit aGratuitous under made were Collections Palawan. and Roxas Tinitian, three loc werefrom collected (PH) the Samples from Philippines Reef. Barrier Great Far North Se the Coral in Reef Shark and Reef Osprey of consisted samples (EA) Australian East Reef. andScott Reef Ashmore Reef, Reef, Imperieuse Clerke reefs: four from taken were (WA) samples West (number1002548). Scientific Authority Management Australian Permit Australian Fisheries an under (Fig. 1) Ocean theIndo-Pacific in sevenreefs from collected were samples Australian Sample collection andMethods Materials currentl overfishing implies that This populations. shrinking contemporary into gene flow limited there is large populations, with associated usually the despite that, apparent it made and diversity genetic ha analysis (ABC) computation Approximate Bayesian endangered species, absolute ethanol in the laboratory for storage at room temperature (Sinclair storage (Sinclair at temperature for the room laboratory in ethanol absolute baited with ~1 ~1 with baited kg ofuncooked chicken ( EDTAsaturated pH 8, NaCl placed were immediately Tentacles was released. collection was non-lethal: a 1 wasnon-lethal: collection was caught using traps positioned onthe using waspositioned traps caught

and ourfindings can CITESfor be the species’qualification assess used listing. to 2 solution and stored at 4°C in the field. Samples were later washed in TE in later washed were Samples field. the in at 4°C stored and solution – 2 cm-long labial tentacle sample was collected before each individual eachindividual before collected sample was labial tentacle cm-long 2 Gallus gallus Gallus y threatens some populations with extinction. with y some populations threatens into a 20% DMSO (dimethylsulfoxide), mM DMSO (dimethylsulfoxide), 100 a 20% into reefs at a depth of ~200 m(Sinclair ~200 of adepth at reefs s enabled a powerful assessment of the species’ thespecies’ of assessment as enabled powerful persistence of high levels of variability that are that variability of levels high of persistence ), set at dusk and collected at dawn. Tissue set atdawn.Tissue ), and collected atdusk et al. 1989) et al.

2011). 2011). and placed into 1 and placed mlinto N. pompilius N. pompilius N. pompiliusN. et al. a, and the and a, ations: 2011) as an an as ent Accepted Article Error loci. rates toHoffman&according per were Amos(2005). reaction calculated acros and re-genotyped were re-extracted 60%ofsamples rate, assessgenotypingTo error 2005). Relatedness between individuals was estimated withS estimated was between individuals Relatedness Hardy–Weinberg equilibrium (HWE; m (1989) Goodnight’s Queller and population and population Oosterhout Oosterhout were assigned using the G the using assigned were sizes Allele Biosystems Inc.). (Applied standard LIZusing size Inc.) (Applied Biosystems GS500 analys were PCR products min. 10 for finally 72°C and s for 90 72°C 90 s, for 56°C s, 30 for of94°C cycles by 44 followed min, 15 for 95°C profile: Qiagen Multiplex Master Multiplexesamplified mix. under werethe following labelled) and1µl (6FAM primer fluorescent forward µM 0.2 primer, reverse µM 0.2 DNA, air-dried wasperf amplification PCR sampled. 215 individuals al. et GENEPOP onaelectrophoresis Fourteen 1%agarose gel. polymorphic was concentration quantified a using fluorometer Optima)(Fluostar and quality assessedits with DNA UK). Manchester, Ltd, (QIAGEN kits tissue DNeasy Qiagen using extracted was Genomic DNA Genotyping Validity and Microsatellite This isarticle by protected copyright.reserved. Allrights frequency alleles andidentify ofnull the estimate to was assessed locus Each microsatellite wasrequired. correction the which of in tests number the overinflating avoid basisto weremade apopulation-by-population Corrections on 2005). 2015) were selected based on satisfactory results from quality checks, and used to genotype all genotype used to and checks, quality from results satisfactory on based were selected 2015) (Raymond & Rousset 1995; Rousset 2008). LD was assessed using 1000 iterations per iterations 1000 using LDassessed was 2008). Rousset 1995; &Rousset (Raymond et al. et P 2004). Null allele frequency per locus was estimated using C -values corrected using the False Discovery Rate adjustment (FDR; Verhoeven Verhoeven (FDR; adjustment Rate Discovery the False using corrected -values ENEMAPPER easure of relatedness. Relatives were removed and departure from and departure removed were Relatives easure ofrelatedness. P v3.7 software (Applie v3.7 software <0.05) and linkage disequilibrium (LD) were calculated using using calculated (LD)were disequilibrium andlinkage <0.05) scoring errors due to stutter errors scoring stutter using due to ed on an ABI 3730 48-well capillary DNA analyser analyser DNA capillary 48-well 3730 ABI an ed on ormed in 2-µl PCR reactions, including 10ng PCRreactions, ormed in2-µl PAGEDI N. pompilius pompilius N. d Biosystems Inc.). Inc.). d Biosystems (Hardy & Vekemans 2002) using using 2002) & (Hardy Vekemans microsatellite loci (Williams loci(Williams microsatellite ERVUS MICRO , HEX, VIC or PET PET or VIC HEX, , v3.0 (Kalinowski v3.0 (Kalinowski - CHECKER (Van et al. s all s all

Accepted Article

clusters was selected from the highest mean log mean thehighest from selected was clusters sugges as log, density posterior the appearance of produced in produced in D To test for an association between between anassociation test for To between lo sampling 1984) &values (Weir Cockerham (Guillot distance against against distance correlated allele frequency model was used in G in used was model frequency allele correlated (Jost 2008)across (Jost lociwerecalculated using admixture and of the posterior probabilitythe posterior ( of removed from Plotting before &Goodnight 1989). dataset the the (Queller analysis natural logarithm were 0.5 a withrelatedness of apopulation within allindividuals on structure, kinship inferred information on (Supplementarythe sampling locations Material ToavoidTable 1). theinfluence of algorithm with 10,000 repeats to develop a consensus value for value for a consensus develop to repeats 10,000 with algorithm distance (IBD) was performed in S in performed was (IBD) distance for all datasets were averaged in C within the data set: S de to wereused methods clustering Three Bayesian Population Structure This isarticle by protected copyright.reserved. Allrights M 2005) as determined in S asdetermined in 2005) number of clusters (Supplementary Material (Supplementary number ofclusters ICROSATELLITE ICROSATELLITE et al. 2005). The software S The software 2005). K ISTRUCT F was inferred from the modal value of the replicate with the highest likelihood. of the likelihood. replicateAhighest from the inferred themodal value with was ST A /(1- NALYSER TRUCTURE F v.1.1 (Rosenberg 2003). Bayesian clustering ofT clustering Bayesian 2003). (Rosenberg v.1.1 ST TRUCTURE TRUCTURE ) was performed (Rousset 1997). 1997). (Rousset ) wasperformed P (Dieringer & Schlötterer 2003) was used to calculate pairwise calculate wasusedto 2003) &Schlötterer (Dieringer P ) of ) of (Pritchard LUMPP PAGEDI H K F ARVESTER given the data over multiple runs determined the predicted predicted the determined runs over multiple data given the ST TRUCTURE and geographical distance, Mantel’s test for isolation by for isolation Mantel’s test distance, geographical and v.1.1.2 (Jakobsson & Rosenberg 2007) using the using 2007) &Rosenberg (Jakobsson v.1.1.2 with 10,000 randomisations. A regression of the spatial ofthe spatial Aregression randomisations. 10,000 with et al. et DEMETICS DEMETICS Fig. 2), and this was compared with with wascompared this and Fig. 2), P v.0.6.93 v.0.6.93 &(Earl 2011). VonHoldt Independent runs ted by the software manual. The number of proposed proposed of number The software manual. bythe ted P 2000), T 2000), ENELAND was run with an admixture model and no prioradmixture model and withan wasrun (Guillot (Guillot termine the most likelynumbe most termine the (Gerlach (Gerlach cations, with Bonferroni corrections applied. Bonferroni with cations, et al. et and the burn-in length was based on the wasbasedon burn-in andthe length ESS Jost’s differentiation index differentiation ( Jost’s (Durand (Durand 2009). 2009). et al. et K 2010). 2010). . Graphical representation was was representation Graphical . et al. 2009) andG 2009) ESS r of genetic clusters r of was run without without run was K (Evanno D est ENELAND ) values Greedy et al. F ST

Accepted Article assumes that PH and of that effectiveassumes ofWA population the are part same size panmictic population of size of population an ancestral that IWOM assumes Model 3). (Fig. scenarios three evolutionary compared We populations), wepopulations), setModel different two from data (i.e. observed the to comparable statistics summary compute that to sizes sizes distribution as parameterized in asparameterized distribution extracted from a normalpriordistribution, and si stepwise-mutati a evolvedunder microsatellites that assumed we Foralldemographic models, 2). models (Table common between parameters The prior distributions were for uniform all demograp history. their whole throughout population apanmictic the samplesas – effectively generation modelling similar sample size, Ashmore Reef was chosen to represent the WA clade. theWAclade. chosenrepresent to was Reef Ashmore size, samplesimilar effects Toavoid distribution. parametersof the two models of interest (Beaumont 2010; Csilléry comparingfrom summary statistics by to is rationale Its 2006). Tavaré (Marjoram & the beingflexibility investigated in scenarios agreat allowing solve analytically, to impossible complex forwhic distribution models of posterior (Beaumont ABCanalysis an we conducted Results), (see samples betweenPHandWA similarity the genetic explain could that hypotheses test alternative To Methods Computation Bayesian Approximate This isarticle by protected copyright.reserved. Allrights their split at rates since migrants exchanged constantly and WAhave PH populations that the adjustment IWOM with N 1 and

N 2 , respectively (Fig. 3b). Model IM (Nielsen & Wakeley 2001) is equivalent to Model to equivalent is Wakeley2001) & ModelIM 3b). (Nielsen (Fig. respectively , m N 12 A and split split m t 21 generations ago into two daughter two into ago generations PAN , respectively (Fig. 3c). A null model was also tested as Model as tested also was model null A 3c). (Fig. respectively , ABC TOOLBOX ABC using Model observed data to the summary statistics obtained by simulating the bysimulating obtained statistics summary the data to observed et al. IWOM IWOM (Wegmann ngle-locus mutation rates were drawn from a ratesfrom ngle-locusGamma drawn were mutation 2010). 2010). on model. Mean mutation rates across loci were lociwere across rates mutation Mean on model. pass the need of an exact likelihood function function by likelihood an exact needof the pass parameters but fixed the divergence time to 1 1 divergence time fixed to parameters but the of substructure within the WA clade, and due to and dueto the WA clade, within of substructure h the likelihood function or be can difficult h the likelihood hic parameters and the same range was used for for used was samerange and the hic parameters et al. et al. et populations, PH and WA, of effective effective WA, of PHand populations, 2002). ABC aims thejoint ABC 2002). obtain to 2010), using uniform priorsforthe using uniform 2010), N P

(Fig. 3a). Note

PAN; this

Accepted Article summary statistics, we used a partial least-squares (PLS) transformation (Wegmann a we summary transformation statistics, least-squares partial (PLS) used perform the post-sampling adjustment step using the ABC-GLM (General Linear Model) in in LinearModel) (General the ABC-GLM step using adjustment thepost-sampling perform rejection step forthe statistics transformed summary comparab densities marginal comparison, For model estimation Model choice andparameter (Wegmann variables response and statistics) (modelparameters) byvariables maximising th identifies PLS orthogonal components. their extract We used used We range. prior the final within wascontained posterior the whole andensurethat distribution posterior on the t of assessment an allow to priors of sets varied with performed were therefore simulations TOOLBOX as an indication of each model’s ability to explain the toexplain ofeachthe asability data. an indication model’s parameter values and the weight assigned to them by the priors (Sousa priors them bythe to assigned weight and the values parameter the rangethe on of dependent less likely or canappearof models more that ABC is limitation were derived from the model choice procedure (Supplementary Material Table 3). wereMaterial Table procedure fromthe derived choice 3). (Supplementary model (Palero (Palero th on based chosen 2), SupplementaryMaterial Table populations; between and (within statistics summary 30 of set a tocalculate was used 2010) fell within the distribution of summary statisti summary of the distribution within fell (for statistics observed summary our checked that ABC analyses were conducted using the package thepackage using conducted were ABC analyses andsimulations Summary statistics This isarticle by protected copyright.reserved. Allrights simulations of our dataset of 14 of our dataset simulations et al. et FASTSIMCOAL (Wegmann 2009; Sousa 2009; et al. et (Excoffier & Foll 2011; Excoffier (Excoffier &Foll 2009). We retained the 5% of simulations closest to the observed data. We the data. observed to closest simulations of 5% We the 2009). retained et al. et microsatellites under each model. model. under each microsatellites 2012; Butlin Butlin 2012; et al. et cs from the simulations retained. Bayes Factors and Factors Bayes retained. simulations the cs from et al. 2009). both PLSboth components andraw summary statistics) e covariance matrix of predictor (raw summary of predictor matrix e covariance 2013). To ofthe reduce 2013). dimensionality the high le between models were produced using the PLS- the using produced were models between le components to explain variability ofresponse variability explain components to , while all raw summary statistics were used to to used were statistics summary allraw while , ose shown to be informative in previous studies studies previous in beinformative to oseshown ABC TOOLBOX ABC et al. et 2013) to run one million coalescent one to million 2013) run ARLSUMSTAT (Wegmann et al. (Excoffier (Excoffier & Lischer 2012). 2012). Exploratory P et al. et -values were taken et al. 2010). One 2010). heir effect heir 2009) to ABC P P

Accepted Article equal to or or equal larger thanthereal to other reef populations. The estimated frequency of null alleles was low for all loci ( loci all for low was alleles null of frequency Theestimated reefpopulations. other in quality due toits analyses retainedin was locus This stutter. dueto errors scoring from resulting verified by ensuring was Parameterestimation adjustment. forpost-sampling wasused 2010) Wegmann (Leuenberger & genotypes was found for wasfound genotypes locus heterozygous of A shortage locations. sampling any in loci any across drop-out allelic frequent for found was evidence replicates. No between determined zero was rateof error A genotyping Genotyping Validation Results underestimated. or over- parameter is a uniformity suggests departure from tochec used werealso data The pseudo-observed werewithin distributions posterior that and checking choice was calculated by estimating the FDR (Verhoeven FDR (Verhoeven the estimating by calculated was choice wi compared were probabilities Posterior compared. the pseudo-dat compared with original data were model’s A performed. modelswas two the data for observed and simulated of comparison pairwise To betweenturn. models each modelchoiceof our theusing robustness procedure, discrimination test in sets eachdata ofthe 1000 pseudo-observed using were to perform our used procedure model choice model each for sets data simulated onemillion (of files results original The priors. original using the each model for sets data pseudo-observed 1000 simulated we choice model procedure, our validate To This isarticle by protected copyright.reserved. Allrights components are extracted foreach are model. The implemented in extracted components GLM method PLS different because eachmodel for performedindependently was This PLScomponents. using carried out both were adjustment and post-sampling step the Fordistance estimation, parameter P- values were reasonably large (>0.05 as suggested in the the in assuggested (>0.05 large reasonably were values P P of the best model. model. ofthe best Npom08 in the Osprey Reef and Scott Reef populations, possibly possibly populations, Reef and Scott Osprey Reef in the a of the same model and those of the model being modelbeing ofthe and those same model the a of the prior Fig.Material 3). ranges (Supplementary k for uniformity of the posterior quantiles; a quantiles; foruniformity the posterior of k th a logistic regression. a Confidence model regression. logistic in th et al. et 2005): the frequency of the frequency of the 2005): ABC TOOLBOX ABC ≤ 0.05). Departure 0.05). ABC TOOLBOX P manual) manual) P being )

Accepted Article ranged from-0.04 to0.35, incorporating spatial data in datain incorporating spatial Results Reef. Osprey as referredto hereafter be will they 1); (Table relatedness of high degree and location geographicalclose their on based analyses Osprey for Reef with grouped Reef Shark was Population Structure Plots ofPlots Scott Reef, (iv) Great Barrier Reef and (v) Osprey Reef. divided five (i)PHsamples clusters into genetic of: The model comparison gave strong support to model to support gave strong comparison The model analysis computation Bayesian Approximate (Fig. 2cand2d). validating results clusters returned consisted ofCoral Sea’s Osprey Reef (Fig. Sub- 2b). consiste cluster the third WA, PHand populations (F dataset thefull in present clusters genetic of association between association reefs, the second cluster of the Great Barri Great the of cluster second the reefs, three and five genetic clusters, respectively. respectively. clusters, genetic three and five its best score was not a result of a bad fit by a fit of abad result not was score its best that summary indicating simulated statistics), the untransformedand post-rejection PLS-transformed range the within lay statistics summary observed (the well data the model fitted Moreover, this WA and betweenthe Philippines sustained migration of theresult were not data the highconfidence that with indicating (0.2%), FDR low was and the consistently showed LDconsistently showed in all of loci any at Nopairs loci. locations sampling across all detectedfrom HWE consistently not was This isarticle by protected copyright.reserved. Allrights K and LnP(K) from K and generated F ST and geographical distance and was not significant ( was significant distance and and geographical not D TESS est populations, suggesting that no loci were physically linked. no linked. lociwerethat physically suggesting populations, ranged from-0.04 to (Table 0.75 1). IBD analysisrevealed overall no (Fig. 2a)and STRUCTURE TESS er Reef, and third cluster of Osprey Reef. Reef. Osprey of cluster third and Reef, er GENELAND all models to the data. Pa data. the to models all returned the first cluster, including PH and all WA and all cluster,PHfirst returned the including ig. 2). The first two genetic clusters consisted of clusters of genetic consisted two 2). Thefirst ig. , (ii) Ashmore, Imperieuse and Clerke Reefs, (iii) (iii) Reefs, Clerke and Imperieuse , (ii)Ashmore, setting the data tolook further for division within IWOM d of EA Great Barrier Reef, and cluster four and Reef, Barrier cluster of EAGreat d results indicated four as the most likely number likely fourasthe most indicated results (see also Material (see Fig. Supplementary 4). ( P (Supplementary Material Fig. 1) returned 1) returned Fig.Material (Supplementary P = 1.0, Supplementary Material Table 3) 3) Table SupplementaryMaterial = 1.0, r 2 rameter estimation under rameter estimation = 0.139; 0.139; = P = 0.095). = 0.095). GENELAND F ST values of bothof

Accepted Article is feasible, it leaves individuals vulnerable to is itleaves vulnerable feasible, to individuals Results from software structure population for Mechanisms genetic drift. limited exhibiting consequently and 2014), al. et Barord pressure; fishing under areas in shown been have declines recentlylarge (population wereuntil sizesthat ancestral population to attributable ofmigration, butmay be theresult similarity wasnot t revealed scenarios demographic different modelling investigation Further unexpected. foun similarity genetic The reefs. Australian population structure but with significant pairwise significant with but structure population limited revealed samples West Australian Reef. Barrier Great the from Reef Shark and Reef Osprey Australian among east structure We population detected Discussion the Great Barrier Reef exceed 1700 the exceed BarrierReef m (Dunstan 1700 Great Reef SeaOsprey betweenthe Coral in Ocean depths in this differentiation. havebeen influential to appears floor) ocean the of geography physical (the Ocean physiography Philippine populations. the in populations Coral Sea be genetically to distin shown to be distinct, not not onlyshown to be distinct, Osprey from Reef Reef TheBarrier was (Saunders Great & Landman 2010). response escape demonstratedor nodefence pompilius model This isarticle by protected copyright.reserved. Allrights bias in parameter estimation (Wegmannparameter estimation bias in strong fr departures show not quantiles did posterior 3,080,000) PH (median: sizesof population current posterior density (HPD95 low, high): 62,816.9, IWOM IWOM individuals to attacks by teleosts showed that that they retreated into their shells and shellstheir they retreatedinto that that showed teleosts by attacks to individuals enabled estimation of theancestral effective estimation enabled STRUCTURE , TESS et al. and and 2009). predation (Yomogida &predation Wani The response2013). of d between the Philippines and west Australia was was Australia west and Philippines d between the et al. but also from the western populations, which supports supports which populations, western the also from but GENELAND 4,508,320), which was smaller than the estimated than the estimated which smaller 4,508,320), was F ct from the Great Barrier Great ct from the ST and WA (median: 2,610,000). The distribution ofThe distribution 2,610,000). and WA(median: 2011), and while movement through open water through open and whilemovement 2011), and and om uniformity, which is indicative of a lack of a lack of isindicative of which om uniformity, sampling sites, indicating genetic isolation of of isolation genetic indicating sites, sampling population size (median: 2,035,120; highest 2,035,120; (median: size population D showed Osprey Reef and Shark Reef Shark and Reef Osprey showed est between Scott Reef and the other west west other andthe Reef Scott between Reef, west Australia, and Reef, west Australia, hat this this hat and N. N. Accepted Article Australian reefs, despite shallower surrounding sea depths. depths. sea surrounding shallower reefs, despite Australian west surrounding and Reef Scott between isseen differentiation andthe Reef, Clerke and Ashmore Philippines as genetic Reefand Scott separate clus measures deviations from total differentiation (Whitlockmeasuresfrom total differentiation both deviations 2011); has been shown in other cephalopods (Pérez-Losada (Pérez-Losada cephalopods in other shown hasbeen 2008) but, but, like 2008) Due to Due depth, residing data their at the to surfacecurrents on explain cannot Reef Scott cluster. as genetic a separate Philippines and west Australia to the same genetic population, whereas population, genetic thesame to Australia west Philippines and the conclusions of previous evolutionary studies using partial partial using studies evolutionary ofprevious the conclusions This isarticle by protected copyright.reserved. Allrights an ability to utilise them to obtain food (O’Dor cephalopods (including (including cephalopods dedicated memory), lobes forlearning and esculenta esculenta knowledge of knowledge occurthisinnautiloidsbut behaviour &Basil does 2013),that (Crook gaps not our in remain each case due to natal philopatry (Kassahn (Kassahn natalphilopatry to caseeach due thiswasin that hypothesised hasbeen It distances. togeographical disproportionate distances genetic (Dunstan (Dunstan et al. et (Biuw patterns No significant correlation was found between between found was correlation No significant distribution is poorlyspecies’ documented. population West Australian results were not consistent across software; software; across wereconsistent not results West Australian 2011; Bonacum 2011; 1993), with recorded movements of up to 6 km that may have been facilitated by cur by facilitated been may that have 6 km to movementsup of with recorded 1993), et al. et (Zheng N. pompilius pompilius N. 2011). 2011). However, et al. et al. Nautilus, Nautilus, et al. 2007). Currents have 2007). Currents to onapositions been individual impact shown reef (O’Dor 2011; Williams 2011; Williams 2009) andthe squid the octopus the octopus ecology. Despite lacking the lensed eye and vertebrate-like brain of other other brain of vertebrate-like and eye lensed the lacking Despite ecology. N. pompilius N. et al. Octopus vulgaris vulgaris Octopus 2012). Loligo pealeii pealeii Loligo et al. et has also demonstrated strong resistance to currents and currents to resistance strong demonstrated hasalso F ST 2003; Buresch 2003; et al. et andlinear geographic distance(Rousset 1997). ters. Geographically, Scott Reef is located islocated Scott Reef Geographically, between ters. et al. 1990). impact 1990). The overall of currents on the (Moreira 2002; Kassahn 2002; (Buresch (Buresch F ST measures deviation frompanmixia,measures deviation COI et al. et N. pompilius N. STRUCTURE et al. et al. sequences (Sinclair (Sinclair sequences F 2006). It has been speculated speculated has been It 2006). ST 2006) 2006) have demonstrated all 2011), cuttlefish the and and et al. GENELAND GENELAND and and N. pompilius N. D 2003; Cabranes Cabranes 2003; has been shown to hasbe shownto been est values distinguish values distinguish TESS designates the the designates assigns the assigns et al. dispersal dispersal 2007, 2007, Sepia Sepia et al.

rents IBD IBD D est

Accepted Article capable of both spatial learning and navigational strategy spatialcapable learning andnavigational (Crook ofboth This isarticle by protected copyright.reserved. Allrights flow was previously established between the sampled west Australian reefs (Williams (Williams reefs Australian west sampled the between established waspreviously flow from the were samples West Australian confirmed. not but was assumed which between connectivity the locations, several in conducted Philippines was Sampling the areas. from sampled the within substructure to inpart, due, areestimates potentially population current Suchlarge impact. has asignificant have yetto drift genetic suggest that 2,562,800) = Ashmore Reef (median and 3,190,920) = (median Philippines for the sizeestimates population Current & 1988). becoming fixed Nei beforedivergence (Pamilo a ofalleles probability lower of individuals between the two sites was unknown, but depth limitations of of limitations butdepth was unknown, between of individuals thesites two genetic the for admias an explanation Australia west and the Philippines between sustained migration of possibility not the modeldoes support ABC Our migration without Divergence our study. unfeasibl completely not is scale asmall on Migration et al. et 13.6 and 0.03 individuals perkm individuals 0.03 and 13.6 remo baited using made estimates density Population size oftheNWpopulation shelfpopulations. the have inflated may geneflow local This drift. Model drift. genetic limited due to populations both in then persist can Alleles 2006). &Knowles Maddison split; large relative sizeis effective population the likely if sorti and stochastic to theretention due populations among relationships about misleading can be data Marker sorting. lineage from incomplete resulted it distance; geographical a oversuch of isolation 2014). Using different methods to generate these results produced predictably dissimilar dissimilar produced predictably these Using togenerate 2014). results different methods IWOM indicated extremely large ancestral and current effective population sizes, resulting in in resulting sizes, population effective current and ancestral large extremely indicated 2 for Osprey Reef and Bohol Sea (Philippines), respectively (Barord (Barord respectively Sea(Philippines), Bohol and Osprey Reef for population estimatepopulation forAshmore reflectingReef, the is possible that the genetic similarity observed has has observed similarity genetic thatthe possible is most northern of these reefs, Ashmore Reef; gene gene Reef; Ashmore these reefs, of northern most ng of ancestral polymorphisms. This is especially especially is This polymorphisms. ngof ancestral xture shown in the structural analyses. Movement analyses. structural in the shown xture to lineage length (the time since the populations sincethe populations time (the lineagelength to te underwater video systems were calculated as as were calculated videosystems te underwater e, but seems unlikely seems e, at thescalebut in unlikely investigated et al. 2013). 2009; Crook&Basil N. pompilius are indicative are indicative et al. et 2012). 2012). Accepted Article protection. legislative enable to makers policy reach now must results Informative extinction. to continue area inthis fishing shoul berepopulated, would the Philippines unlikely itis that that indicates contemporary migration for thePhilippines thelong-term cluster. survival to is genetic imperative this of of The absence adequate protection adaptation, of local thepossibility with population aunique as but has occurred, selection divergent ifresulting unknown is it so and beenmeasured not have sites between differences mechanisms toprotect theseas populations discre theneed for highlights Australia west and Philippines between the migration of The absence implications Management modern modern of accepted species the forasimilar as time havebeen independently evolving populations current based on a generation time of 5.6 years (Saunders & Landman 2010)) bymodel 2010)) & Landman years (Saunders of5.6 time generation a based on years generations/1,660,366 (median 296,495 = estimation The time split Seaof the Coral (Fig. 4). Reefs Shark Osprey and distinct the more genetically in decline a with further Reef, Barrier the Great abundance data. Additionally, Barord Barord Additionally, abundance data. This isarticle by protected copyright.reserved. Allrights progenitor population located in the Philippines (Wray Philippines the locatedin population progenitor from the of colonisation direction proposed the supports also samples The Philippine richness allelic in markers. ourcurrent set of with detectable al. et nocultural orhistorical with new, is relatively time long generation in a results nautiloids 2010) of yet totake effect; lowfecundity reductionhave ofpopulation consequences genetic the that ispossible It 4). (Fig. sampled locations at other in thethan Philippines wasnolower butourdatashowed allelic richness reduction, that 2010), and so it is possible that we will see a genetic response to exploitation that is not yet not thatis exploitation to response genetic a see will thatwe ispossible and so it 2010), N. pompilius (Kröger et al. and long developmental (Carlson time and long 2011). et al. et al. (2014) documents evidence of a sudden population size population of asudden documents evidence (2014) significance in studied areas such as Palawan (Dunstan (Dunstan as Palawan areassuch studied in significance te conservation units (Moritz 1994). Environmental (Moritz 1994). units te conservation compared to other cephalopods. Fishing for shells shells cephalopods. for Fishing other compared to et al. 1995). There is lower allelic richness in in richness allelic is lower There 1995). et al. 1984; Landman 1984; IWOM IWOM indicates that et al. d

Accepted Article should be utilised to develop efficient species-specific management plans for declining populations. populations. for declining management plans species-specific efficient develop beutilised to should dynamics. Such genetic canan into migration andstudies genetic population data provide insight species, relativelyinaccessible In continued exploitation. for incentive decrease the CITESwould Allonautilus Allonautilus increased fishing pressure (Dillane (Dillane pressure fishing increased experience cephalopod islikely stocks targetsthat will fishingand it resources, industry the novel fo extrapolated are results when especially studies, range species-specific for need the demonstrates cephalopods within IBD seenin variation The This isarticle by protected copyright.reserved. Allrights 2012; De Angelis 2012; Barord Barord 2012; De Angelis 2012; wide (Hutchings 2000; Worm 2000; wide (Hutchings the local extinction ofthe local and pr reduce over-exploitation to the need emphasises migration of result the wasnot similarity this that conclusion Our considered. had been previously than Australia and west the Philippines Williams2011; A range of molecular studies has been conducted on coleoids (Allcock (Allcock on coleoids conducted been has studies of molecular A range Conclusions distance. account for by unitsand more should discrete geographic separation incorporate management than should plans management showhow Ourresults substructure. data presented reveals here resolution Australian reefs (Williams west the on DNAdata mitochondrial example, For protection. adequate establish and patterns connectivity determine to markers finer-scale or for multiple need the highlights study andour 2013) al. on previous evolutionary studies on these populations (Sinclair (Sinclair these populations on studies previous evolutionary on based Australia andwest between east division genetic hypothesised Wehad anautiloid. markers in population structure analysis using minisatellites using analysis structure population 2005; Zheng 2005; protection is ever increasing (Dunstan (Dunstan increasing isever protection et al. et al. 2012), but we found a greater degree of genetic similarity between samples from from samples between similarity genetic of degree greater a wefound but 2012), N. pompilius 2009; Moreira et al. et al. et 2012) suggested that the population was panmictic, but the higher- the but was panmictic, the population that suggested 2012) et al. 2006; Doukakis 2006; Doukakis inthe Philippines. Protection for et al. 2014). Overexploitation is threatening marine species world- species marine isthreatening Overexploitation 2014). et al. et 2005). The data supporting the need for The the need for 2005). data supporting 2011), but this is the first study to use microsatellite microsatellite use to study first the is this but 2011), , microsatellites and mitochondrial DNA (Dillane DNA (Dillane mitochondrial and microsatellites , r fisheries management. As fin fish stocks decline decline stocks Asfinfish management. fisheries r et al. et al. et 2010; Bonacum 2009; Neubauer 2009; et al. Nautilus 2007, 2011; Bonacum Bonacum 2011; 2007, et al. et et al. et al. and 2011; Williams Williams 2011; 2013; Watson 2013; Allonautilus 2015), including 2015), Nautilus

under et al. et event et al. et al. and and et et

Accepted Article Microsatellite primer sequences for for sequences primer Microsatellite accessibility Data and RCWproject. al draftedthemanuscript the TBoversaw DAD, WS, data. the and interpreted analysed LD and BCJ RCW, marker quality. the and RCWRCW genotyping. WSDAD assessed the RCW and performed samples. obtained Author Contributions supported by theLeverhulme numberRPG-2013-198). (grant trust at (NERC) the Council Research Environment by UKNatural the anddataanalysessupported The work were laboratory discussion. helpful throughout the project. We are gratefulto members Molecular oftheSheffield Ecology Laboratory for advice Mortega for Kim Tuckerand Rachel Steeves Buckland, We thank analyses. with support comm for Aaron Comeault Gumbsand We Rikki thank Acknowledgements (Neubauer marine populations of conservation the imperative for is protection, legislative with collaboration in these, Enforcing This isarticle by protected copyright.reserved. Allrights Mantel’s test are available in Dryad, doi:10.5061/dryad.j251f. Dryad, available areMantel’sin doi:10.5061/dryad.j251f. test theandinformation for geographic data genotyping the Microsatellite HG918068-HG918111. et al. 2013). 2013). N. pompilius N. pompilius l authors contributed edits and comments. and comments. edits authors contributed l NERC Biomolecular Analysis Facility. LD Facility. was Analysis NERC Biomolecular are available through NCBI: accession numbers accession NCBI: through available are ents on the manuscript, and Martin Hinsch for for Hinsch Martin and manuscript, the ents on Accepted Article Allcock A, Lindgren A, Strugnell J (2015) The Lindgren (2015) cont J A,Strugnell A, Allcock References This isarticle by protected copyright.reserved. Allrights De Angelis P (2012) Assessing the impact of the Assessing impact DeP (2012) Angelis CITES (2015) Convention on International and ofWild EndangeredTrade Species Fauna on International CITESFlora. 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Fisheries , 123 Nautilus Nautilus , 264–274. 264–274. , , 10 versity , 564– , ng , Accepted Article Koch PL, Barnosky AD (2006) Late Quaternary Extinctions: State of the Debate. of the State Extinctions: Late Quaternary AD (2006) Barnosky Koch PL, Kassahn KS, Donnellan SC, Fowler AJ Fowler SC, KS,Donnellan Kassahn Guillot G, Leblois Guillot A, StatisticalFrantz Coulon R, methods AC inspatial (2009) genetics. Guillot G, Mortier F,Guillot Estoup Geneland: package forlandscapeA (2005) a genetics. computer Hardy JO, Vekemans X (2002) SPAGeDiHardy X(2002) Vekemans JO, Gerlach Kraem A, G, Jueterbrock Kanie Y, Fukuda Y, Nakayama H, Seki K, Hattori M (1980) Implosion Implosion M of (1980) living H, Seki K, Y,Nakayama Hattori Kanie Fukuda Y, Kalinowski ST (2005) Hp-RareKalinowski a 1.0: pr computer Hobson K a. (1999) Hobson Ka. (1999) Tracing and migration origins Jost L GST Jost andits measure (2008) relatives not do differentiation. 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National Academy Sciences National America States ofthe of United of Ecology Notes Ecology juvenile movement formarine areas. protected and early-life-historytraits on connectivityamong littoral fis 1801–1806. 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t N N Parameters Prior range Mean Median Mode HPD95 low HPD95 high high HPD95 HPD95low Mode Median Mean range Prior Parameters MUT ARG Ashmore Reef ( Philippines ( Population Philippines Clerke Reef ( Scott Reef ( Great Barrier Reef ( 31) Imperieuse Reef ( Shark Reef( Osprey Reef ( 13) 2 1 A Uniform [0- 5×10 Uniform [0- 5×10 Uniform [0- 5×10 K U Uniform [0 - 6] 1.78785 1.62815 1.35678 0.105534 3.72541 3.72541 0.105534 1.35678 1.62815 1.78785 [0- Uniform 6] Uniform Uniform N N N N = 30) 0.263* 0.097* - 0.015* 0.015* 0.173* 0.354* 0.268* 0.268* 0.354* 0.173* 0.015* 0.015* - 0.097* 0.263* =30) N =8 061 062 076 068 071 043 -.1 - -0.014 0.483* 0.731* 0.678* 0.736* 0.612* 0.651* 8) = = 27) - 0.015* 0.044* 0.014* 0.024* 0.130* 0.330* 0.237* 0.237* 0.330* 0.130* 0.024* 0.014* 0.044* 0.015* - 27) = Uniform [0- 10 = 32) 0.114* -0.028 0.084* - 0.004 0.124* 0.322* 0.234* 0.234* 0.322* 0.124* 0.004 - 0.084* -0.028 0.114* =32) = 45) 0.693* 0.696* 0.740* 0.703* 0.745* 0.5* - 0.012 0.012 - 0.5* 0.745* 0.703* 0.740* 0.696* 0.693* =45) N = 29) 0.136* - 0.018* -0.004 0.006 0.121* 0.319* 0.228* 0.228* 0.319* 0.121* 0.006 -0.004 0.018* - 0.136* =29) N = N = [10 -6 0.158* 0.044 0.111* -0.042 - 0.144* 0.343* 0.343* 0.144* 0.044 0.158* 0.255* 0.111* - -0.042 0.527* 0.485* 0.586* 0.494* 0.528* - 0.312* 0.312* 0.229* 0.485* 0.527* - 0.586* 0.494* 0.528* - 5×10 - 6 ] 353845 6 6 6 ] 2.61E+06 2.56E+06 2.09E+06 665830 665830 4.77E+06 2.09E+06 1.05E+06 4.99E+06 2.56E+06 62816.9 4.51E+06 3.79E+06 ] 2.61E+06 3.19E+06 1.31E+06 ] 3.08E+06 2.04E+06 ] 2.17E+06 -4 ] 5.00E-05 4.11E-05 2.61E-05 -2.53E-07 0.000122236 0.000122236 -2.53E-07 5.00E-05 4.11E-05 2.61E-05 ] Ashmore Ashmore Reef Scott Reef 296495 296495 Clerke Reef 105529 105529 Imperieuse Reef -2511.56 -2511.56 839452 Great Barrier Reef Osprey Osprey Reef Shark Shark Reef Accepted Article

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Accepted Article

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