Accepted Article e-mail: [email protected] e-mail: 33 74 62 93 (+47) Phone: Norway His, N-4817 Flødevigen, Research, Marine of Institute Sørdalen Knutsen Tonje Correspondence Australia 4811, QLD Townsville Kristiansand, Norway Norway Oslo, N-0316 Oslo, PO Box Blindern, 1050 2 This by is protectedarticle reserved. Allrights copyright. doi: 10.1111/eva.12611 lead to differences between this version and the throughbeen the copyediting, pagination typesetting, which process, may and proofreading hasThis article been accepted publicationfor andundergone peerfull but review not has 5 4 3 1 Sørdalen Knutsen Tonje Harvesting changesmating behaviorEuropean in lobster Article :Original type Article DR. HALVOR (OrcidKNUTSEN ID :0000-0002-7627-7634) MISS TONJE KNUTSEN SØRDALEN (Orcid ID :0000-0001-5836-9327)

Asbjørn Vøllestad Asbjørn National Lobster Hatchery, South Quay, Padstow, Cornwall, PL28 8BL, UK Cornwall, Quay, South Padstow, Lobster National Hatchery, University, Cook James Studies, Reef Coral for Excellence of Centre Council Research Australian Norway His, N-4817 Flødevigen, Research, Marine of Institute N-4604 Agder, of Sciences, University Natural of Department (CCR), Coastal for Centre Research of Biology, University of Department Centre for Ecological Synthesis (CEES), andEvolutionary 1 , Halvor Knutsen , Halvor 1,2,3 *, Kim Halvorsen Tallaksen 2,3 , Even Moland

Version of Record. Please Version as this cite ofRecord. article 2,3 , Esben Moland Olsen 3 , HugoB. Harrison 4 , Charlie Ellis 2,3 . 5 , Leif Accepted Article This by is protectedarticle reserved. Allrights copyright. gammarus Keywords: size. body smaller towards evolution fisheries-induced will likely accelerate selection sexual of weakening success, the higher reproductive relative with males) (particularly target large to continue individuals fisheries truncation of males by fishing thereduces oftraits variability thatsexual acts selection If upon. size that suggests strongly This areas. fished in was non-significant it while reserve, the males in body in onboth claw and size actedsize positively selection inthe larger reserve. Sexual significantly was own difference relative size the body but size, their than males larger matefemales with would showthat results Our reserve. inthe be larger to tend males whereas area, the size fished average in equal approximately of are females and males areas, two between the mortality fishing specific sex- sizeand in differences the to Due females. egg-bearing of harvest the on aban and size legal a by season, a minimum isregulated closed the fishing, fishery area Inthe to Norway. Southern open ( European lobster valued compare the highly to of overexploited and patterns mating assignment genetic parentage we In this study, used species. exploited of systems andthemating harvesting selective between onthe interplay knowledge empirical islimited yet, there sexual of selection strength dynamics and be dependent onthe strongly to is predicted the harvest-induced evolution of rate Consequently, selection. sexual and patterns effects onmating subsequent with outcome interactions, competitive of the and mates prospective of the availability affect can population wild a from Removing individuals Abstract , marine protected areas. Assortative mating, mating behavior, sexual selection, parentage analysis, analysis, parentage selection, sexual behavior, mating mating, Assortative

Homarus gammarus

) in a designated lobster reserve and nearby fished area in area fished nearby and reserve lobster a designated in ) Homarus Homarus Accepted Article This by is protectedarticle reserved. Allrights copyright. harvested populations. of rates andtrajectories evolutionary predicting al. (Kingsolver selection thanthat natural generated can bestronger by success fecundity and mating 2006;Kokko and Rankin (Kokko selection ratios (e.g. 2012), andQuinn this Kendall willlikely influence and strength the opportunity sexualof (Coltman malebody size in reduction sexually males dominant trophies for ledtoartif sheep bighorn ( this in effect demonstrated Swain 1989; mates (Wilber for access to competition are generallymate in and important that traits choice intraspecific body size; and claws) and antlers horns, (e.g. weaponry of size the as such characters, selected sexually against al. on selection of sexual intensity inthe variation underlying factors and success, reproductive of male populations’ contribution to phenotype potential Lane 2008; Roy and Fenberg 2003; Hutchings and (Rowe populations and mating systems exploited of betweenmortality human-induced been theinteraction dedicated to persistence (Jørgensen Enberg (Heino traits history of life evolution contemporary drive can harvesting size-selective (Berkeley limits minimum-size imposing regulations management or marked due preferences to individuals andlarge harvesting Typically, targets 2009). Hard Allendorf (Hutchings and Rowe 2008; population the from certain phenotypes removes always non-random and disproportionally virtually harvesting is For 2001). instance, (Palumbi nature selectivein forces be considered the strongest one of to also are activities human but goods valuableservices, and for onhealthy Humansecosystems depend Introduction 2001; Siepielski Siepielski 2001; select to tends harvesting First, reasons. several for surprising is research of paucity This 2015). et al. et al. et al. et 2012; Uusi-Heikkilä Uusi-Heikkilä 2012; 2004; Beamish 2004; et al et et al. et . 2011), illustrating the necessity of considering sexual when .of 2011), necessity illustrating selection the et al. et 2007, 2009; Kuparinen and Merilä 2007). However, far less attention has lessattention However, far 2007). Merilä and Kuparinen 2009; 2007, et al. et 2007; Woolmer et al. 2006). There is mounting empirical evidence showing that such that evidenceshowing empirical There is mounting 2006). et al. 2015) with consequences for population productivity and productivity population for consequences with 2015) Ovis canadensis Ovis 2003; Pigeon 2003; et al. et male remainstraits, largel icial evolution icial evolution towards smaller sizehorn and a 2012). Third, the strength of sexual selection on on selection sexual of the strength Third, 2012). et al. 2013). Coltman and colleagues (2003) Coltman andcolleagues 2013). (2003) et al. ) when the harvest of the larger and thelarger of harvest the ) when 2016). harvesting sex Second, alters 2016). if et al. et y ignored (Uusi-Heikkilä (Uusi-Heikkilä y ignored 2011).More so,the et al. 2015; et et et Accepted Article lobster fishery is regulated by closed season isregulated fishery lobster This by is protectedarticle reserved. Allrights copyright. 1999;MacDiarmid and Wolcott (Kendall species occurcrustacean many in (Karnofsky is seek to ready mate she a will out male and female when a that show studies Laboratory 2008). (since females egg-bearing of harvest the on ban Kleiven 1995; (Anonymous pressure fishing intense to subject therefore ( lobster American the and lobster European of consisting lobsters, clawed The years. multiple across fishing ( (Butler systems onmating effects fisheries- exploring when systems as valuable reference particularly are therefore 2005), and Dayton (Berkeley conditions baseline towards are restored be to expected composition size and sex ratios asdensity, such characteristics demographic population where reserves, No-take Fenberg marine and 2003; Roy 2008). dynamicsand Hutchings are(Rowe be intact to likely (Worm overexploited or Mobley2011; (Lane the wild in onmating behavior more research theneedfor studies underscore and field (Rowe species model and controlled environments to and matebeen have choice limited selection Mostsexual of studies task, species inmany especially aremarine noteasily which observedintheir natural environment. a of trivial is mating system no affect might thestability harvesting how alone. Disentangling fishing by thanexpected body forsize stronger selection of smaller is strength the overall mean variation, and morhua modelling the on work Atlantic cod ( (2008) and Rowe’s Hutchings marine environment. in the selection sexual of processes natural disrupt may potentially howharvesting directly address Homarus gammarus ) showed that if reproductive success increases with body size and harvesting decreases its its decreases body and size harvesting with increases success reproductive if that showed ) We investigated potential effects of harvesting on the mating system of the European lobster theEuropean of lobster mating the on system harvesting of effects potential We investigated to used been never have techniques assignment parentage knowledge, our of best the To Homarus americanus Homarus et al. et al. 1989; Bushmann and Atema 1997; Skog 2009a). Given that sperm limitation may sperm Givenlimitation Skog 1989;Bushmann 2009a). andAtema1997; that 2014). Considering that many commercially fished species are regarded as fully- regarded as fully- are species fished commercially many that 2014). Considering ) by comparing paternity data from a lobster reserve and and opento an adjacent area reserve a data paternity lobster from comparing ) by et al. et al et 2008; Uusi-Heikkilä, 2012), but discrepancy in results between laboratory between laboratory results discrepancy in but 2012), Uusi-Heikkilä, 2008; 2009), few locations for these species remain where natural mating natural where remain thesefor species 2009), few locations ), are long-lived iconic species with high commercial value and and value commercial high with species iconic long-lived are ), et al. 2015). 2015). , minimum legal size (> 250 mm total length, mmtotal 250 (> size legal minimum , preferentially choose a large individual as mate choose a individual large preferentially et al. et et al. 2012). The Norwegian et al. 1999; Kendall 1999; Kendall 2004; Birkeland Birkeland 2004; Gadus Gadus TL ) and a and ) et al. et al.

Accepted Article This by is protectedarticle reserved. Allrights copyright. can a weakening lead success to reproductive high large males with targeting weargue fisheries that management aimed mitigating tools long-term at negative impact selectiveof harvesting. Specifically, developing for relevant selection, andsexual mating systems changes to fisheries-induced quantifying in variability the fished area. trait reduced the of because area fished the than reserve the in larger be to males, unsuccessful and successful values inthesebetween mean trait is, that difference the differentials, selection size-assort weaker of hypothesis our with Aligning body size male two atraits; on breeding within season, sexualselection, of strength estimateand compare the was second to Thus, our objective selection. be sexual strong under therefore clawsizeshould claws increases abilities a (Ate competitive male’s 1944) and larger Templeman, 1935; 2001; & Reynolds, Addison, (Debuse, than females larger claws Males Skog have relatively 2009b). 1986; (Atema contestdominance overand males shelters fight and intense is competition male-male clawedlobsters, In 2003). Cézilly Bertin and 1989; and Price mating. size-assortative of pattern aweaker creating thus reserve, the in compared tofemales area fished the sizesin smaller matemales with of should thatfemales predicted We therefore 1). (Figure tothe reserve relative in area fished the are smaller males and females between size mean differences the area), fished in the of females egg-bearing return mandatory to due sexes (and between the areas regulations conservation thedisparate of Probably because exists. of - matedbody individuals between size association -the nonrandom mating, thetwoareas size-assortative andwhether in pairs mated difference between al. (Wahle size female with increasing exponentially production increases as egg favour large females ensure passing of or largersizemalesto similar of Hines 2002; 2013). Our first objective was to determine to what extent there is a consistent relative size size relative there a consistent is whatextent to determine objective was Ourto first 2013). Our results contribute to a broader understanding of fisheries-induced evolution by evolution fisheries-induced of understanding a broader to contribute results Our Karnofsky (e.g., crustaceans many in selection be sexual under to shown been has size Body (carapace length, length, (carapace et al. et 2003; Jivoff 2003; Sato Jivoff 2003; CL ) and absolute and relative andrelative andabsolute ) et al. et 2006), females would expectedly prefer to mate with tomate with prefer 2006), expectedly would females sufficient sperm. In addition, malesalso should sperm. addition, In sufficient ma and Cobb 1980; Elner Cobb so ma and and Campbell 1981), ative inthe fishedmating area, we that predict claw size claw (width of crusher of crusher claws, (width CW ). et et Accepted Article from the tip of the foremost pleopod was stored in pure ethanol for later genetic profiling. All All lobsters genetic profiling. later for ethanol pure was in stored pleopod of the from foremost the tip pieceA small tissue of tonearest millimetre. measured was rear of carapace) the socketeyethe to and carapace Claw( width 2mm,length Hallprint). × 45 (TBA2, T-bar tags visible (Wiig in2011 anotherthe area in study fished with conjunction and in and females for 2012, in2011 fishing when sampled maleswere Additionally, above. described survey trapping research standardised of the assampled males the part of were Most intheanalysis. parentage ascandidates possible to 2010 include order from June as – - many paternal 2013 in December extensively fished throughout were Males two areas. the in size sample 2012to abalanced achieve - December inOctober season females were from obtained thefished area he with additional seven the reserve, in are caught more lobsters Because area). fished the reserve and 48from the from (60 2012 and 2011 in September - June from caught were 108 sampled, females bearing fishedwith the sameefforthauls (100 per year), seeMoland are area fished and The September. reserve days four August/early during 5-30 inlate meters depth at set traps parlour standard using lobsters samples survey annual this Briefly, 1. Figure in presented is survey atrapping and research from standardised data length incatch-per-unit-effort trends suggests very exchangelittle ofindividuals between areasthe two 2015). Temporal (Thorbjørnsen data meters ~800 andmark-recapture of a by distance separated area is fished and monitored the This by is protectedarticle reserved. Allrights copyright. Norway ( coastsouth-eastern in 2006,located at of Skagerrak the September in established reserve lobster adesignated in and fishing to open an area in conducted study was The sampling lobster systemStudy and Materials andmethods phenotypes. (smaller) productive less towards evolution fisheries-induced accelerate further could which sexual selection of et al. 2013). Captured lobsters were sexed, measured and individually tagged with externally with and tagged individually measured sexed, lobsters were Captured 2013). lp from local fishermen during the ordinary fishing lp local fishing duringfrom the ordinary fishermen et al. (2013) for details. Ofthe egg- Figure 2 Figure CL ). The reserve reserve The ). – rear of the the of rear – Accepted Article This by is protectedarticle reserved. Allrights copyright. MSATALLELE manually scored using CEQ sequencers capillary HGD wereThe one triplex into pooled loci (loci (Life Technologies). primers fluorescent- dyedforward MyCycleron amplified andPCR (Omega kit product Isolation Bio-Tec inc.) sampled650 males (n all area, thefished from area females 55 and reserve the 60females from from DNA extracted was genotyping microsatellite and DNA extraction males. sampled the with assignment parentage running when fathers actual the finding of likelihood the increase help should which combination, genotype offspring - mother each of based known the on offspring father the ustodeduceallowed genotype of the ethanol of (a total eggsten vials with fromeach with separate vials and in of sampledstored the tenpleopods the at the egg of top mass each near where oneegg offspring, was randomly of samples with along collected were samples tissue addition, In T-bar tags. tagged with individually and measured also were females egg-bearing Captured Genetic sampling of female and offspring quality. DNA highest the ensure to genotyped was sample tissue freshest the years, insuccessive Whererecaptured males were site. atthesampling were released SYSTEM SYSTEM and Knutsen 2009 for primer sequences). The DNA was extracted with E.Z.N.A. Tissue DNA DNA Tissue E.Z.N.A. with extracted DNA was The sequences). primer for 2009 Knutsen and All individuals tenmicrosatellitewere with genotyped and and HGB 106, 106, 6 v 8.0 respectively. As the length of the alleles slightly differed between the instruments, instruments, the between differed slightly thealleles of the length As respectively. 8.0 v HGD ) and one simplex (locus (locus andone ) simplex 111 (Alberto 2009), a script build on R, was used to bin the scored raw sizes from both both from raw on build 2009), scored sizes wasthe a R, used tobin (Alberto script and reserve reserve HGC GENEMAPPER ™ 118 = 331, n = 331, 8000 (Beckman Coulter) and ABI (Beckman Coulter) 8000 ), three duplexes (loci (loci duplexes three ), HGC fished v3.7 (Applied Biosystems) and and Biosystems) (Applied v3.7 120 = 319), and a total of 1,150 offspring from the 115 females. females. 115 the from offspring 1,150 of total a and 319), = ). Fragment analysis of PCR products was carried out on out carried was PCR Fragment). analysis of products female). Samplingfemale). fertilized ofeach eggs female HGC loci developedforEuropean lobster (see André 111 and and ™ 3130xl (Applied Biosystems), and and Biosystems), (Applied 3130xl HGC CEQ 131, 131, ™ 8000 8000 HGC ™ (Bio-Rad) using 129 GENETIC ANALYSIS and HGA 8, HGB 4 Accepted Article duplicates may be due to tag-loss and thus repeated tagging. tagging. repeated thus and tag-loss to due be may duplicates Such entries. genotype identical for checking samples by identifyus andremove duplicate to This by is protectedarticle reserved. Allrights copyright. in check function identity The analysis Genetic details. additional (S1.1) for analysis. information Seesupplementary and excludedquality further from poor DNA of lociweremore considered at five whichgenotypes or missing were for Individuals as missing. left were three repeat-runs after be solved not reliably could Genotypes that errors. scoring tominimize times three manually checked also were The assigned outerrors. genotypes rule candidate assignedmales with inthe initial parentage analysis re-amplified were re-extractedand to All PCR plateselectrophoreses. and used for each in 96-well was included control a samples, negative of cross-contamination To control the two. the results from calibrate correctly and analysers fragment there was no significant genetic differentiation between the two areas ( areas two the between differentiation genetic significant no was there 0.99), all samples were pooled in subsequent analysis. The fixation index (smallF, (smallF, index Thefixation analysis. subsequent in pooled were samples all 0.99), statistic of algorithm usingRaymond statistic Markov the in chain & Rousset (1995) a ratio likelihood of with each for loci was pair tested disequilibrium linkage Pairwise zero. from as from Hardy–Weinberg deviations (HWE, Equilibrium exactcritical significance All test). levels for F in sampled betweenlobster the genetic differentiation the candidate file. from removed subsequently duplicates were and these cases, cause all in probable asthemost confirmed was 38 tag-loss the data males, the for re-capture in compared were expectedannual growth When size and sampled the genotypes among males. 650 at each locus with with locus each at heterogeneity between localities (years pooled) was tested using using tested was pooled) (years localities between heterogeneity ST , with Weir and Cockerham’s (1984) estimator, estimator, (1984) and Cockerham’s Weir with , Genetic variation within samples Genetic variation within forthe estimated was adult samples We estimated the only. FSTAT . One sample t-test was used to assess if if assess to used was t-test . Onesample CERVUS v. 3.0.3 (Marshall (Marshall v. 3.0.3 θ , in , in the reserve and the fished area using Wright’s using area fished the and reserve the GENEPOP et al. CERVUS 1998; Kalinowski FSTAT F v. 4.5. Allele frequency Allele frequency v. 4.5. IS estimates differed significantly differed estimates significantly identified 38 duplicated duplicated 38 identified F v.2.9.3.2 (Goudet 1995). As 1995). (Goudet v.2.9.3.2 ST from 0.000 to 0.002, all to P > 0.000 0.002, from GENEPOP et al. F IS ) was measured v. as 4.5 well 2007) 2007) enabled Accepted Article This by is protectedarticle reserved. Allrights copyright. GERUD ( each locus at number allele including estimates diversity genetic multiple testing R-package wereadjusted after Benjamini with Finally,and Hochberg fdrtool, (1995). female and infers the most likely number of fathers contributing to each to Where batch. contributing fathers of number female the mostlikely andinfers probability given one parent is known for each locus and combined (EXC) were estimated with werewith estimated combined(EXC) each and for isknown locus given probability parent one paternity if offspring differed from the first male at five or more loci, did not show sign of scoring scoring of show sign not did loci, more or five at male first the from differed offspring if paternity equal Inferred probabilities). multip r DNA,multiple mother’s from example contamination based rates, ontheerror helped minimize an over batches. Altogether, a total of 612 (n males, 97females 612 of a total batches. Altogether, the of genotyping allow successful to were yield insufficient eggsfemalesDNA which had from missing to due ( data seven excluded maleswere from Genotypes Paternity and multiple mating analyses 4 ( information see supplementary error rates of andestimation approach and genetics, population (see methods of awere combination with estimated sampled, sampled, notall Although fathers were paternity inthe population. the multiple overestimating of level and avoided assignments negative and false minimize helped This higher. positive false confidence 95% or assigned with paternities accepted onlyWe both sexes.regard to paternities in multiple testing for a prerequisite bepolygamous, and males femalesWe both to individuals. allowed among and parentage assigning sib-ship in provides configuration that the mostprobable method) Monte Carlo program (Markov-chain a full-pedigree likelihood and(Wang Wang 2010), Jones 2004; inferred more than one sire in a batch, visual inspection of genotypes and changes made by genotypes andchanges of more ina than one sire inspection inferred batch, visual (and pooled) in the final parentage analysis ( parentage analysis thefinal in pooled) (and ). 2

(Jones 2005). Locus-specific genotyping error rates, allelic drop-out ( drop-out allelic rates, error genotyping Locus-specific 2005). (Jones COLONY can infer their genotypes from the pedigree analysis to the number of mates to mates of each to number the to pedigree analysis the genotypes their from can infer le paternities were only accepted as true cases of multiple of cases true as accepted only werele paternities Table 2 estimation ofmultipleestimation paternity cases (due to for Table 1 ). We assigned parentage using using parentage We assigned ). reserve reserve econstructions of alleles suggested with almost almost with suggested alleles of econstructions = 51, 51, n = for errorrates). For detailsdescriptive on ≥ N 5 missing loci) and eighteen and loci) missing 5 A fished ) and the theoretical exclusion exclusion andthe theoretical ) = 46) and = eggs967 were used ε 1 ) and false allele ( allele false and ) COLONY COLONY COLONY v 2.0 v 2.0

S1.2- ε 2

), Accepted Article This by is protectedarticle reserved. Allrights copyright. COLONY bycalculated been not had question in loci the if and genotype) mothers (miss-matching error the mated pairs (70%).For these itwas pairs, necessa from year the inmostof the of female sampling differed fathers inferred of The year sampling 2011. and 2010 in time some mated likely most 2012 and 2011 in sampled eggs (black) extruded newly contamination/amplification issue or had loci altered by altered loci had issue or contamination/amplification not not male be show resolved by did ancould at by fiveof theevidence first loci, offspring least the genotype if of paternity multiple be cases only of as resolved batcheswould cases. offspring The paternity multiple overestimate could and that missing, are genotypes where cases alleles propose in they moult and re-mate (Aiken (Aiken re-mate and moult they which after duration, a for to similar incubation eggs prior and fertilise months, usedexternally to 9-11 for mating stored are during received spermatophores whereby cycle, reproductive biennial a have females of majority The seasons. mating several across captured were lobsters that fact t-test.two-tailed For thesize relationship analysing and females of size overall the compared Wefirst mating Size-assortative original genotype data. This is because supplementary information supplementary information ( onthe settings see For used more details machine. Linux 6.7 M820, PowerEdge CentOS ona with and very runtime long full-likelihood, with settings thehighest precision with analysed inspected the results that were flagged as cases of multiple paternity by paternity multiple of cases as flagged were that results inspected the batches, we sired thefrequency multiple When determining of allele frequencies. population various scenarios ofmultiple contributions skew theof fertilisation between malesbased on the software P The probabilities ofdetecting multiple paternalcontribution (P due to missing alleles. missing due to R DM (Neff et al. et S1.3

2002). P 2002). et al. The files input in ). 2004; Agnalt Agnalt 2004; COLONY R DM uses Monte Carlo simulations to calculate P DM simulations Carlo uses Monte can alter loci in accordance with error estimates and estimates error with in accordance loci canalter COLONY males inthe whole data set in both areas with a et al. et between mated pairs, we had to account for the account for to we had between mated pairs, ry to estimate male size for the time at which 2007). Thus, the egg-bearing females with with females egg-bearing the Thus, 2007). COLONY were set up with two two upwith and runs set were replicate . R COLONY DM) were quantifiedusing along along the with R DM under DM under Accepted Article following formula: the using males success mating with for adjusted length carapace of calculation werethe in included these twomodels values The predicted from using regression. whohadmoulted, a linear individuals capture of first for at the asa of increment carapace time estimatedlength growth yearly function the (f), and This by is protectedarticle reserved. Allrights copyright. (Agnalt errors be measurement to assumed were differences smaller year; previous from the was 5mm higher or had if moulting size We that the occurred difference inferred a capture at aswith carapace regression. logistic moulting first of the probability of length afunction we First, the estimated period. this within at any intwo point that hadbeenyears consecutive captured males and extracted 2004-2016 area from andfished thereserve from data we used mark-recapture Tothis end, patterns. mating to analysing size-assortative sampled were prior females corresponding respectively, as predicted from respectively, from predicted as Where Where model with only an additive area effect with the likelihood ratio test. test. ratio the likelihood onlyeffect with model anadditive with area against a2) (Eq. interaction x– size Area female models with comparing between areas, differed such patterns and testedwhether sizepairs) inmated andmalebetween female body correlation supplementary information mm 113 (> classes tobelow0.75 in sizes larger moulting decreases of probability thatthe and annually, below90mm males all showedthatalmost predictions the data file if samemale the if file the data sample mates between putative We six matings excluded 1) 2) alinear we used sizes, male adjusted the With CL

CL ). The overall probability that males of all sizes would moult once every year was > 0.5 (see 0.5 > was year once every moult would sizes all malesof that probability Theoverall ). and is the male carapace length (mm), (mm), length malecarapace the is ̂ = = is increment yearlyestimated growth the mm)(in andprobability of moulting,

S2, S2, ++ has mated with multiple females. multiple females. with mated has Figure S1 Figure + ). using linear and logisticregression. The model − Year × is the year of sampling for males (m) and females females and males (m) sampling for of istheyear model to test for model totestassortative for (a mating positive CL × d in different areas. Males were duplicated in in areas. duplicated Males were different d in

(the minimum legal moulted minimum (the size) × ̂

et al. 2007). We 2007). then Accepted Article Scotland has been estimated to 80 mm 80 to has estimated Scotland been This by is protectedarticle reserved. Allrights copyright. had not. males that to offspring, successfully sire size ( body the compare used to model was linear a Also, t-tests. two-tailed with assessed were differentials selection of Significance 1983). (Lande andArnold oneeach in area-year combination of deviation scaled toacentred and standard values trait mean- were calculations, studyto Prior our system. for andappropriate conservative 80mm of threshold father thus we a potential study, consider the Scottish mmwas sampling female73 our berried in However, smallest the 2003). CL males only mm 80 with fathers, larger or potential males among immature probability for including but to reduce the malesthis population, in known The isnot for sizeWade maturity of 1984). and (Arnold area each in fathers successful of mean the from fathers potential of value trait mean onmale si body differentials selection Standardised Selection onmale traits we maintain that sexual selection is likely to be the primary mechanism underlying these selection selection these be the primaryto underlying is likely mechanism selection sexual that we maintain Although years. the different in mating success obtaining of prospects the determine evidently would Because malessampled were over three seasonsand females in two, the aforementioned andnatural) fishing (both as mortality mayselection, sexual reflect pure not differentials selection and variable where each (2010-2012), modelsfor year linear generalized we used areas, univariate between differed malesof assigned proportion the whether years. To test amongsampling distribution variable inthe trait modeltoaccount for effect included as anadditive Year was assignment). and effect between area interaction inthe reservebe larger would (a significant males unsuccessful and successful between value trait mean thedifference in waswhether Of interest were included in the selection differential calcu differential selection the in included were 3)

Area + + × + + + = the predictor. CL for males and 79 CL ) of males that were successfully assigned and, thus and, thus assigned successfully maleswere of that ) lations. Size at maturity at maturity Sizelations. ze and claw size were calculated, subtracting the subtracting calculated, ze claw sizeand were CL assigned for females (Lizárraga-Cubedo (Lizárraga-Cubedo females for

(0, 1) was the binomial response response was binomial 1) the (0, CL CL for in for European lobster , compared to 82 mm in 82 to compared , and above to be and above to et al. et

Accepted Article the same model due to high collinearity (Lande and Arnold 1983; Zuur 1983; (Lande and Arnold high the model due collinearity same to = in notbe therefore (r and could included 0.90) traits correlated and sizebody width strongly were ( body size onmale 1998) and Stern (Janzen regressions logistic estimated werefrom gradients Selection 2012). This by is protectedarticle reserved. Allrights copyright. clawsize ( relative of a measure isthen clawsize, which theresidual length and used carapace and width claw between regression linear the from residuals the extracted we traits, both (Kingsolver alone differentials selection than selection of shape and represent strength the better andtherefore values trait when change, inthefunction fitness sensitivity whichcapture gradients, the selection of We 5out 272). out of 2011: 245; standardised estimated 24 (2010: analyses further warrant to year) (in following paternities assigned number of a sufficient at and alsohad the time reproduction of becausemalespopulation the represent they in2010 sampled an specific more a conducted wealso differentials, term for absolute size (body or claw), exemplified for exemplified size claw), (body absolute or term for an interaction effect with area for each of the traits ( traits the of each for area with effect interaction an differed relationships trait-fitness evaluate whether with together covariate structures: structures: for intermediatesized males(Uusi-Heikkilä might highest be thatmale mating success shown have studies as recent onmale selection size, linear) (i.e. rather than non-linear), directional (i.e. stabilizing the supported We whether alsodata explored 7) 6) 5) 4)

CL ) and claw size ( size claw and ) =+ =++× =++× =+ CL . We also fitted a ( claw . size modelonlyrelative including Wealso fitted CW

++ ), with mating success (s) as the response variable (0 or Claw 1). or (0 variable response asthe mating(s) success with ), et al. et

2012). For this, we ran models including Fora squared including this, we models 2012). ran × alysis of sexual selection, where we included only we included where selection, sexual alysis of between areas, we also included models testing for for modelstesting included we also between areas, CL, CW and CW CWand CL, CL below: below:

et al et res ) using the following model) using the following et al. et . 2012; Matsumura 2012; . 2010). Thus, to include to Thus, 2010). CW CW res res ). To ), as et al.

Accepted Article to 561 to 561 ( males of number the analysis, reducing further males so from removed year this 2013, all were from in the estimation, equation from Dodds equation from in the estimation, for accounted not are rates error of the (though effect males candidate selected randomly two between distinguish to power sufficient given indicating probabilitybe to 0.9998 genotype, a maternal known exclusion combined the estimated We 0.844. to 0.606 from ranged heterozygosity observed This by is protectedarticle reserved. Allrights copyright. loci across( all S2, see respectively, and 0.983 0.934 andoffspring; (adults was of over 0.946 typed individuals loci all The proportion Lobster samples genetics andpopulation Results ( gradients selection approximate estimated We wa which size, sample small for corrected criterion, All selection gradient (full and models reduced) body size were calculated by multiplying multiplying by calculated were body size claw and on gradients selection Mean and 1998). standardized Stern approach (Janzen regression Development inR 3.2.4 (R Core Team performed 2016). were analyses All statistical which has value and minimum maximum natural no (Hereford relative claw size, suchas for trait applicable across studiesis not but selection of strength comparing (Matsumura deviation Table 2

H ). See supplementary information (S1.3-4) for additional details. additional for (S1.3-4) information See). supplementary E = 0.738; = 0.738; TableS1 et al . 2011). The mean standardized selection gradient is recommended for Table 1 Table for females and eggs analysed) and the genetic diversity was high was high diversity thegenetic and analysed) andeggs females for ). The number of alleles per locus ranged from 9 to 19, and the allelesto 9 of The number from locusranged ). per et al. β avggrad 1996). No parentage was assigned to males sampled in in sampled males to assigned was parentage No 1996). β were compared with the Akaike information information Akaike the with were compared by the trait value´s mean and dividing by its standard standard its by dividing and mean value´s trait bythe avggrad s used to determine the most parsimonious model. parsimonious themost s determine to used ) for each trait with the Janzen-Stern logistic logistic eachwith Janzen-Stern trait ) for the et al 2004 and Matsumura andMatsumura 2004 et al. 2011). 2011). Accepted Article This by is protectedarticle reserved. Allrights copyright. and to males6.4% inthe fished area (GLM: the matings reserve males in the (totalto of of) assigning number 19.2% 2010, thebetween areas in probability differed thereserve. Assignment from involved males (66.7%) 27 matings,36 known those 54 Of high candidate confidence. with males) of 561 femalesmales 43 the (7.7% of one to ( father aknown were of assigned A 511 offspring total (52.8%) Mating patterns (GLM: females. Polygamous weremales la significantly not three matedand three with had two females five which with had mated more of one female, than multiply mated. may infact be matedalso single females our some of multiply cases matedfemales, of confirmed results (9:1) could only be detected with a 65% confidence. This suggests that, in addition to the two a the in of primary favour the skewin observed male however, moreusing 95% than nine offspring, a of Weconfidence (three 70:20:30 with sires) a skew alsodetect could to down equal contribution. multiple paternity only with ten offspring genotyped both information, cases supplementary (see S2 in ratio) these in a favour of amongskewed were primary highlymale matedin contribution the multiply (9:1 females The matedbe to multiply paternal females. concluded male beinga second bytherefore and sired of the broods evidence two (2.0%) of showed However, a of support becausesire. second lack of in only one male by sired were probably the broods of most that concluded we results, the assignment mm tohave ~140 been estimated reserve, the involved alarge pairs malefrom area. inter-area these Two of malesopposite mated the from with exchange of individuals across area boundaries, though five females (reserve females (reserve five though boundaries, area across exchange individuals of inferred genotypes from 41 un-sampled males that si β Of the inferred maleswith known identity, eight (reserve matings,after inspecting but multiple tobe of broods the cases flagged 24of Colony initially = 5.956, t 5.956,

= 1.333, df df 1.333, = 53, p = CL 0.188). On average, the level of polygamy was higher for for On polygamy higher of 0.188). thelevel was average, β at the time of mating. = 1.376, t t 1.376, red red offspring with females.42 There was little rger than males with only one recorded paternity paternity onerecorded only males with than rger at ten loci exceeded >99% confidence assuming confidenceexceeded >99% loci ten at = 2.384, df df 2.384, Table S1 Table Table 2 Table = 6, fished fished 6, = 154, p 154, ). ). We eggs 54 assigned from ). The power to detect = = = 2) had 2) mated with 3, fished 3, fished 0.017). 0.017). COLONY = 2) had

Accepted Article effect on mating success ( on mating effect success anarea support not did gradients selection sexual estimating used for mating dataof season) (2010 This by is protectedarticle reserved. Allrights copyright. area thefished ( males in than larger were females years, all sampling Across Size-assortative mating females. mating and 1.16 unknown) with (known 1.01 males and with females mating males, with interaction effectinteraction (LRT; Across all sampling years, selection differentials on body size ( size body on differentials selection years, sampling all Across Sexual selection Area: (GLM: area fished the to compared reserve the in size own 46, P = 0.0009, Multiple R Multiple 0.0009, = P 46, pattern (GLM: mating size-assortative positive There strong was a 0.051). difference smaller %) (6.4 and marginally (t-test: statistically t significant size averagethe female,the than with larger a male had pairs 15 three the but of all largerasof sizes, t (t-test, % 22.5 of difference size an average with female, a a male out of of smaller 34) with larger mating consisted mating(see S2insupplementary pattern information, (n pairs Inter-area 0.57). df Area; x parentage (GLM: bothbodyclaw size and for reserve inthe size larger males was unsuccessful ( area significantly positive in the reserve, while they were = 68.6, p-value p-value 68.6, Table 2 Table CL : β

= ). Correspondingly, the standardized trait difference between successful and successful between difference trait standardized the Correspondingly, ). 0.76, t 0.76, =

6.1799, df df 6.1799, = 0.037), but not not the but reserve in ( 0.037), = 2.44, P χ = 2 Table 3 5) were removed prior to analysing the area-specific size-assortative size-assortative to thearea-specific analysing were prior 5) removed = 2

= 1.479, P = 0.50, 0.50, 48.27, P <0.0001). Females in the fished area also paired with males with paired also Femalesarea P fished the in 48.27, <0.0001). = 0.02 and 0.02 ). Instead, only a additivemodel containing ofbody effects sizeand Figure = 0.224), females with mating males with larger to their relative CW

3) . An additive area effect was supported over an over supported effect was An . area additive : β

= Table 2 Table 0.88, t 0.88, more variable and non-significant in the fished the fished in andnon-significant variable more Figure S2 Figure ; t-test: t = 2.85, 2.85, P CL ). In the reserve,butpairs all two (2 β ) and claw size ) ( and clawsize

= = 17.65, t 17.65, = -0.57, df df -0.57, 0.005, 0.005, = β 2.034, df df 2.034, = 0.838, =0.838, Table 2 = Figure 4 Figure 3.722, 3.722, P = 98.5, p-value 98.5, CW ; t-test: t t = ) were = ). Thesubset 28.35, P 3.560, df 3.560, = 0.0005). = = 2.12, 2.12, =

=

Accepted Article This by is protectedarticle reserved. Allrights copyright. the marine reserve relative to the fished area. Alth in differentials selection the higher for positive isthelikely explanation male on traits sexual selection for scope theincreased Therefore, male diversity of phenotypes. a to wider access females have given that behavior, mating ‘natural’ more considered can be what of depiction good a display might reserve inthe almost a for lobsters decade, fishing from protection Having been afforded patterns onmating fishing of The effect evolution. fisheries-induced rate the of and trajectory and output reproductive for consequences potential with patterns, mating affect cangreatly fishing that findings suggest our size. Overall, claw andbody absolute on than size, rather claw relative on wasstronger selection Sexual area. fished inside the reserve, differentialsselection onthese werewhile traits weakerand the significant not in mating success their influenced strongly (body and claws) male size Wethat area. also documented size differe within-pair this that weshow Moreover, dispos have astrong wheremating females pattern, genetic assignment Our parentage sexual selection. toestablish reserve marine a and coastal outside environment natural inside its in lobster European exploited the system of We themating investigated Discussion AIC lowest the had success mating male on size claw residual term representing stabilizing and suppor traits onallthree positive significantly ( clawsize residual on positive strongly and body size on was positive selection sexual inference, modelfor parsimonious Using most the (

Table 3 Table ). A simpler model excluding the effect of body size also received some support ( support some received also size body of effect the modelexcluding simpler A ). or ( disruptiveor sexual selection Table 4 Table ). For comparison, univariate selection were gradients univariate selection For). comparison, whether canwhether harvesting affect and mating patterns ough females inthe fished areatendedto mate with ted over more complex including a squared models clearly demonstrates a positive size-assortative size-assortative a positive demonstrates clearly ition mate ition to with comparatively larger males. nce was larger in the reserve than in the fished fished inthe than thereserve in nce was larger Table 4, Figure 5 Figure 4, Table c score and therefore the most support themost support and therefore score ).

Table 3). 3). Table Accepted Article This by is protectedarticle reserved. Allrights copyright. al. et Debuse Cowan and Atema 1990; (Atema females 1986; chances with increase their shelters to ought better of occupancy males, whose be typically smaller to available morewould than shelters good free up should large of lobsters mortality fishing high Further, 1989;Arnqvist 1992). behavior(Conner and mates competitors between encounter rates lower tobe islikely through selection relaxed have more difficulties finding a larger mate. When and the fact thatfemales thesame about were size lobsters of density lower the area, fished the In fished. heavily more site a in pattern mating random a but pressure, area fishing caught femalesan moderate in larger in of pattern mating assortative size- positive a found who (2003), colleagues and Gosselin by obtained lobsters female American ( smaller much was sexes between size body in difference average the themselves, than larger males following year, relative to a similar sized male in the reserve. This implies that the selection mate the andsuccessfully surviving of as have chances 2010, lower should a area in fished male areas. Moreover, temporal replicates, tracking several selection episodes and ideally also including including also episodes and ideally selection several tracking replicates, temporal areas. Moreover, and fished of pairs reserves be multiple using tested should the approach pressure, that in fishing in situ regimes harvesting the effects of study to may theavailable bestoptions of beone continues as usual, fathers. and betweenunobserved observed the distributions trait different reason weno assume to find However, be inferred. canonly selection therefore true and fitness The consequences for sampled. (females) and withoutpaternitywith onthe broods males of we compareonly but distributions patterns, the trait affecting mating harvesting is indeed selective show that work this from results Arguably, large individuals. against selection fisheries of component may mayan unidentified reflect but also be purely not selection differentials sexual due to ♂ > ♀ 1999, 2003). 2003). 1999, ; 6.4%) . To test thegenerality findings,the spatialof in ecology variability attribute to and indisputably High fishing mortality could also mortality High the lo could fishing explain Contrasting no-take reserves and complementary monitored control areas where harvesting areasharvesting control where monitored and complementary no-take reserves Contrasting than in than in reservethe ( ♂ > ♀ ; 22.5%). ; 22.5%). These results are in line with those for wild-mated wild-mated for those with line in are results These as males in this area, implies that females would individuals are more sparsely distributed, sexual wer assignment probability for males in the assignment formaleswer probability in Accepted Article (Grafen 1990; Berglund Berglund (Grafen 1990; and choosy females tocompetitors male signals of likelycombatquality are be honest to anddisplay, of dualfunction like claws, with structures model. Sexually selected multivariate parsimonious the to sizebody according on most weaker selection with along inmaleselection lobsters, driving sexual trait the be to appears size claw relative et Interestingly, al 2004). (Hereford estimates published 140 calculated from above being were high, median well both (3.56) relatively (1.93) the round (Waddy season and isable a females round inseminate multiple to breeding within maleall-year- sperm can produce since an females, areintermoult malethan choosy less lobsters of themating female even time-out the to period due more fecund alarger, with opportunities mating lower cost of the at this comesfemalesif smaller matewith to reluctant be could males hand, other the On males. larger choose to females for reasons receptiveness tomate, forprotection during the need moulting andsperm are quantity plausible MacDiarmid 1997; males (Jivoff smaller than faster sperm load to depleted the sizeand replenish female, the of ejaculate tailoring of capable are reserves, sperm greater have decapods large male Secondly, 1990). This by is protectedarticle reserved. Allrights copyright. female (Atema soft-shelled the of guarding post-copulatory and pre- successful increase to believed time, a strategy some for cohabitate and mate after soon will she where ina shelter, males’ moult usually lobsters femaleFirst, clawed choosingmales. larger from a benefit canFemales have direct small males. someofthe largest males hadmated withsmall females, while the largest femalesnever mated with mating pattern; assortative driving role positive in the anplay appears to Female important choice inclawed lobsters selection sexual of Drivers sexualselection. in consistency stability and temporal individual be and time totest usedtoestimate fitness could life selection, and fisheries natural The univariate selection gradients (mean standardized) on both claw (3.04) and body claw size on both (3.04) standardized) (mean gradients selection The univariate et al. et al. 1999;Kendall 1979; Karnofsky 1979; Karnofsky et al. 1996). Relative claw size might therefore be a better male be of measure a clawRelative size 1996). therefore might better et al. et al. et 2001; Gosselin Gosselin 2001; 1989a,b; Karnofsky and Price 1989; KarnofskyCowan and Atema andPrice 1989; 1989a,b; t. Nevertheless, is reasonableit to assume that et al. 2003). Thus, the narrow time-window of of time-window thenarrow Thus, 2003). et al. et 2017). 2017). Accepted Article indicates that the likelihood of sperm limitation is present. As for males, we found eight individuals males, individuals eight Asfor is present. we found sperm of limitation thatthe likelihood indicates mate with relatively sm thatfemalesthe Europeanbut finding lobster our unknown, remain iscause for Whether concern it for linked to sperm-limitation due to fisheries-induced sex ratio imbalance (Gosselin (Gosselin imbalance ratio sex fisheries-induced dueto sperm-limitation to linked This by is protectedarticle reserved. Allrights copyright. undetected. went but broods single sired among our present were broods sired multiple additional that it is possible chance, by thatwere discovered beside cases two Thus, the high with probability. skew 9:1 of sire parental asecondary detect to power nothave we statistical did offspring, sample of al. paternal (Streiff skew of haveto shown level have often (e.g. high broods Multiply-sired crustacean theother deviated siblings. nine from each from brood offspring a asingle andonly male primary (Ellislobster in the of European no paternity the UK inaregion study multiple found a recent of contrast, incidence In analyzed). broods 97 of out (2 males different bytwo being sired of evidence area had the fished but females in caught pattern, two fertilization was theprevalent broods femaleon Single paternity Multiplematings and sperm limitation female attraction. individuals large very also implying that size, claw Meeren for andUksnøy Note that we didnotfind 2000). support stabilizingon body and selection VanDer Elner andCampbell 1981; of thanand Cobb body1980; victors predictor size (Atema American larger clawslobster, increa found to are InPratt 1998; both Oliveira and European and McLain and males 1983; Custódio 2002). (Christy thansmall-clawed more and attract females more males competitions win crabs, large-clawed where Fitness benefits accruing largeto rela males with age. toold survival be simply could chance to due size,body claw and absolute than which quality 2004;Yue Multiple paternal American and Multiple lobster fertilizations paternal in havepopulations, been documented et al. et al. 2015). Both cases found in our study had contributions highly skewed in favour of of favour in skewed highly contributions had study our in found cases Both 2015). 2010; Bailie aller males (presumably with lowersp with malesaller (presumably et al. 2011). However, withonly method because However, our 2011). a of limited

tivelylargeclaws Fiddler well documented are in se male competitive abilities tobea abilities se and better male competitive maintain high male-male competitiveness and/or and/or competitiveness male-male high maintain erm storages) in the fished area area fished the in storages) erm et al. 2005). et et Accepted Article 2007). Accounting for evolutionary processes in management can potentially increase long-term long-term increase can inmanagement potentially processes evolutionary for Accounting 2007). This by is protectedarticle reserved. Allrights copyright. (Jørgensen in and populations adaptive of potential loss evolution minimize harvest-induced (Urbach andCotton 2008). phenotypic effects thanonother evolutionarytraits Therefore,stronger we may1995). anticipate and Moller (Pomiankowski traits selected non-sexually may thanfor characters be much higher selected underlying sexually variation genetic evolution because the fisheries-induced studies of in component future a of sexualselection this, we inclusion of encourage In spite induced selection. obtaining data forestimating sexual selection is ofte be that could The reason 2008). Cotton and Urbach (Hutchings andRowe 2008; conclusions their in studies out as of tends tobeequations the left body size. and less towards smaller productive evolution likely fisheries-induced will accelerate sexual selection to target individuals (particularly males) with high continue fisheries If fishing. to due selection sexual weakened for support empirical first the provides lobster onEuropean study our best To our knowledge, Roweand 2008). random (Hutchings mating of under than the assumption evolution harvest-induced tofaster lead to isexpected fishing due to traits these in variability mean and in areduction fisheries), in targeted and females by preferred both are males larger when (e.g. selection harvest opposing under traits for non-random is Whenmating Implications andmanagementfor evolution fisheries-induced perspectives and Kokko 2006). WadeRankin 2003; and females (Shuster malesmonopolize to and mate multiple for with density increased opportunities suggests population where higher the reserve, the males came from Seven these of a single thosewith mating. size from with known identities thathadmated more with than

A general objective in an evolutionarily enlightened management framework should be to be to should management framework enlightened an A evolutionarily in objective general selection sexual evolution, ratesfisheries-induced of for thepotentialramifications Despite sessing this subject, with potential consequences for for consequences potential with this subject, sessing er relative theweakening reproductive success, of n more challenging than for natural and fisheries- natural n more for than challenging one sampled female, they not but differ in did et al . Accepted Article 1 (starting fishing season For 2017 the footprints. mayinduced evolution considerable left have already fisheries- Thus, aage large reaching size. or high individuals of the prospects diminished profoundly Gwinn Gwinn resilience (Matsumura and environmental productivity populations therefore be mandatory to consider in evolutionary enlightened management. management. enlightened evolutionary in consider to mandatory be therefore evolution is of and part anshould Sexual selection integral andbiased lowered sex ratio. density may through toact at selection but sexual berelaxed of selection the sameupon, strength time, the sexual for variability thephenotypic reduces wild European in behavior Selective lobster. fishing 2015). andBarnett Baskett also: in (see areas fished evolution and fisheries-induced selection buffer sexual strengthen possibly thecould reserves from males ‘attractive’ larger, more of and spill-over largewith males) (mated large females from output reproductive increased for the potential within, recovering selection sexual are with increased, thenumber reserves and size of If, however, trajectory. evolutionary the on haveanyeffects are strong to unlikely coastline the along established reserves small of a of management, spatial handful benefit the As for coast. caught lobster theSkagerrak along This by is protectedarticle reserved. Allrights copyright. Halvorsen Tiainen 2017; 2005; and Dayton (Birkeland sex ratios and balance ageandsize structure also restore can of largeprotection individuals managementin towards A shift 2017). and Zimmermann Jørgensen 2008, Fraser and slots (Hutchings through gear (e.g. modifications reducing entrance traps) diameter ormaximumin size limit/harvest large individuals harvest of such fishing, as of restricting selectivity the maychanging by beachieved Uusi-Heikkilä Roweand 2008; random of mating (Hutchings tothe scenario relative body size smaller towards toslow evolution are body predicted with size) correlate genetically (that Mollet Jørgensen2017; (Zimmermann and stability collapse andecosystem population yield, theresilience to st of October), a maximum size limit of 320 of mm a mm length total size maximum CL) (~116 for was implemented October), limit of et al. et et al. Long-term overfishing has left the European lobster in Norway at a historically low level and level low Norwayin ata lobster thehistorically left European has overfishing Long-term In conclusion, our paper presents novel empirical support for how fishing affects mating affects fishing how for support paper novelempirical presents our In conclusion, 2016). Fishing thatcan maintain increaseor variability ofsexuallythe selected traits 2015). 2015). et al. et 2016), which should have positive effects on have on effects positive should which 2016), et al. 2011, Arlinghaus 2011,Arlinghaus et al. al. et et al. 2016). This This 2016). 2010; 2010; Accepted Article This by is protectedarticle reserved. Allrights copyright. and cycle, reproductive Growth, K.E. (2007). and T.S. Jørstad, A.L., Kristiansen, Agnalt, cited Literature https://doi.org/10.5061/dryad.1b1f023 Repository: Digital Dryad the from available Data Data archivingstatement comments thathelpedto improve the manuscript. constructive for reviewer oneanonymous Arlinghaus and Robert to samples and lobster providing for Arendal in to fishermen the thelaboratory, assistance in Hanne Sannæs to for parentage data, the with and adviceWangassistance helpful for Jinliang to gratitude We our express Education. of Ministry Norwegian the from funding Agder through of by University the supported This was study Acknowledgements Council, Report to the Minister of Fisheries and Oceans. FRCC 95, 95, R FRCC 1. and Oceans. Fisheries of the Minister Report to Council, Conservation Resource Fishery Atlantic lobster. framework for A conservation Anonymous (1995). animals. wild of harvest through selection caused by unnatural evolution Human-induced (2009). J.J. F.W. andHard, Allendorf, alleles. microsatellite of MsatAllele F. An R binning the package tovisualize (2009). 1.0: Alberto, American Aiken, D.E., Mercer, S.M. and Confirmation Waddy, of External S.L. (2004). in Fertilization the Lobster,

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J.H., Xia, C.M., Le, Wang, Li, J. Yue, G.H., statistical statistical to Aprotocol exploration data for avoid C.S. (2010). and Elphick, common E.N. Ieno, A.F., Zuur, sciences biological of problems. Methods in Ecology and Evolution and Ecology in Methods 6, 107–15. 6, 107–15. Procambarus clarkii Procambarus 1, 3–14. 1, 3–14. . International journal journal International Accepted Article COLONY2, italic.shown in This by is protectedarticle reserved. Allrights copyright. 2 1 and EXP 1: Table (1-4) Tables mother is known, critical P-value for critical HWE P-value isknown, = for mother (a 0.05); test (EXP) of the second theprobability parent: excluding of whenthe a chosen randomly non-father Flødevigen area, Norway given southeast Also in2010-2013. areprobabilities expected exclusion the

denoted ‘*’ showing null-alleles at high frequency, frequency of null alleles; nullalleles; of athighfrequency, frequency null-alleles denoted ‘*’ showing ε and observed ( observed and Average Average C129 C111 C120 D106 C118 N Locus B6 B4 A8 C131 C131 D111 ε ε 2 2 1 , false allele rate. The samples are based on 727 (612 males and 115 female) lobsters. lobsters. female) males and (612 on727 115 are based samples The false rate. allele , =Allelic drop-out rate estimated from andPedant Micro-checker, the latter is denoted‘ = False allele rate estimated from Pedant. Where Pedant estimated 0.000, 0.010 was implemented in in implemented was 0.010 0.000, estimated Pedant Where Pedant. from estimated rate allele False = average

Description of loci used in the paternity analysis and error rates. error and analysis paternity the in used loci of Description 11.9 0.710 0.738 0.999 0.023 0.014 0.021 0.012 0.012 0.021 0.014 0.023 0.040* 0.096 0.040 0.999 0.738 0.710 11.9 0.000 0.582 0.779 0.706 14 0.001 0.001 0.021 0.013 0.023 0.008 0.009 0.735 0.725 0.529 9 0.001 0.745 0.870 0.844 0.017 19 9 0.703 0.709 0.494 0.431 0.013 0.012 0.012 0.012 0.013 -0.060 -0.031 0.013 0.431 0.494 0.709 0.703 9 0.073 0.370 0.587 0.619 9 11 0.738 0.818 0.646 0.000 0.001 0.044* 0.044* 0.001 0.044 0.010 -0.004 0.006 0.001 0.062 0.000 0.646 0.818 0.738 0.062* 11 0.000 0.399 0.606 0.606 0.116 9 0.818 0.661 14 0.712 0.000 13 0.806 0.830 0.669 0.231 0.025 0.012 0.012 0.012 0.025 -0.024 -0.012 0.000 0.231 0.669 0.830 0.806 13 0.909 0.405 0.631 0.645 12 a

H EXP EXP calculated byGERUD2 according to the equations inDodds O ) and expected ( expected and ) H O H E H EXP E ) microsatellite heterozygosity for the adult European lobster at lobster European adult the for heterozygosity microsatellite ) Uncorrected p-value F F IS IS

, inbreeding coefficient; F(null), loci loci F(null), coefficient; inbreeding , F (null) (null) ε 1 1

et al ε Number ofalleles N 1 m , allelic drop-out rate; rate; drop-out , allelic m m m m m

0.023 . (1996). . (1996). m ’. 0.010 0.010 0.010 0.010 0.010 0.010 0.010 0.018 ε 0.010 0.010 0.010 2 2

a

Males Females claw width (P-value) (P-value) width claw diff* Selection St. (P-value) carapace diff* Selection St. mm (CV), Mean claw width mm length (CV), carapace Mean assigned) candidate males of No. (No. male candidate assigned No. offspring mm length (CV), carapace Mean offspring) No.females (No. of Accepted Area Article This by is protectedarticle reserved. Allrights copyright. counts. the in once than more appear hence and females multiple with mated have that males including males, byknown matings of number the counts and reported is confidence different selection Significant parentheses. value) in value (P- with body confidence size and clawwidth for (diff*) differentials selection standardized variation (CV), of corresponding coefficient in millimetres with claw width length and crusher carapace mean parentheses, males in andassigned candidates of number For males, candidate males. assigned the offspring number of variation (CV), percentage and of coefficient corresponding with mm in length carapace mean offspringparentheses, in of females andnumber of For females, number inyear inthe analysisseparated Table Summary andarea. 2: on European used results lobsters

Reserve Fished Fished Reserve Fished Reserve ReserveFished 0.94 0.80 0 01) 0(.3 9 01) 8(.5 11(.9 8 01) 99 (0.18) 88 (0.16) 101 (0.19) 88 (0.15) 95 (0.17) 90 (0.13) 104 (0.18) 8(.5 4 01) 1(.4 4 02) 5(.2 4 02) 4(.6 45 (0.20) 54 (0.26) 44 (0.22) 55 (0.32) 44 (0.21) 51 (0.24) 46 (0.18) 58 (0.25) 98 (20) (00) -0.37 (0.53) (<0.01) (<0.01) -0.25 (0.67) - 9 7 ) 9(3% 5 6 ) 9(6) 5 6 ) 158 (34 %) 353 (69 %) 92 (0.11) 69 (36%) 46 (457) 57 (63 %) 97 (0.12) 89 (33 51 (510) %) 94 (0.13) 296 (70 %) 105 (0.15) 19 (188) 91 (0.10) 9(90) - 96 (0.11) 27 (269) - 42 (420) - - - - 2010 2011 2011 2010 2012 Years pooled 80 (5) 80 (5) 0.67 0.45 148 (11) (00) 0.22(0.45) (<0.01) (.1 0.29 (0.33) (0.01) ials are in bold. Only paternity assigned at 95% Only95% paternityassigned at bold. arein ials 111 (8) 1.66 1.66 28 (5) (.8 0.7 (0.19) (0.08) (.6 0.43 (0.37) (0.06) 6() 7 3) 287 (18) 274 (36) 96 (5) 0.93 0.78 (00) 0.22 (0.34) (<0.01) (00) 0.16 (0.48) (<0.01) 88 (0.15) (0.15) 88

Accepted Article This by is protectedarticle reserved. Allrights copyright. reserve Area, fished; and width; variables (standardized): Explanatory score. criterion information AIC, parameters; of Akaikes P,number variable. response as the success reproductive 2010 using 3: Table The AIC model with lowest is indicated bold. in number number Model 2 CW 1 8 CL x Area 4 4 1 2 CLxArea 9 Null 2 8 7 CL 6 CW 4 5 CW x CW CL+ 4 CW Area 3

Model selection. CL + CW CL + Structure P AICc res res 2 x Area 4 4 x Area res res x Area 5 5 xArea

Logistic regression modeling on selection of male European lobster from of male modelingfrom lobster onselection European regression Logistic CW 3 re s, s, relative claw size (residuals from claw body size regression). size claw regression). body from size (residuals claw relative 134.06 132.26 129.98 128.60 151.04 147.65 143.97 138.18 135.24

CL , carapace length; length; carapace , CW , claw , Accepted Article This by is protectedarticle reserved. Allrights copyright. gradients selection approximate ( ( coefficients regression logistic the table For eachJanzen-Stern gives 2010. Flødevigen trait, in during sampled male Europeanlobster 4: Table interest are carapace length ( length are carapace interest analysis. to 1andmean-centred of prior deviation a were standard traits scaled to size. tobody relative All claw size for asa are andclaw used proxy width length carapace between regression the from linear Model no. Trait Trait Model no. 4 3 1 2

Sexual selection estimates. CW CW CW CL CL res res

1.544 0.835 0.609 0.424 1.320 CL β SE SE z-value p ), claw width claw ), ( width β β ) and their corresponding standard error (SE), z- p-value,(SE), error and standard and the corresponding ) their avggrad Sexual selection operating on body size and relative claw size in in relativesize claw size and body operating on selection Sexual 0.391 0.203 0.201 0.227 0.394 ) andthemean standardized selection gradient ( CW 3.350 3.949 4.111 3.033 1.868 ) and residual claw width ( claw width and residual ) <0.0001 0.965 - - 0.965 <0.0001 <0.0001 1.170 - - 1.170 <0.0001 3.039 0.657 <0.0001 0.002 0.512 0.512 0.002 3.555 0.06 0.310 - - 0.310 0.06 β avggrad CW

re s ), where residuals residuals where ), β μ

β μ ). Traits). of Accepted Article This by is protectedarticle reserved. Allrights copyright. group. each in lengths mean indicate lines Vertical years. sampling the in assignment grey) four (light (blue) andwithout with confirmed European lobsters parentage. Figure Males 4: with size. equal in males are and where females marks Y= X) (isometry, line stippled and Value 1.0 black and malecapture point female represents Area excluded. are pairs Inter-area period. four-year inthe grey) area and (dark reserve (red) fished (corrected sizes, seeMaterials and methods) and female European lobster that formed pairs (n = 51) in Figure 3: monitored fished area.Inserted image: European lobster ( line: stippled reserve line: boundary, Solid (B). area fished and reserve and lobster Flødevigen Figure Sampling 2: location. in Norway cm). (25 lobsters for size legal minimum the line denotes horizontal Thestippled line. stippled in females and line insolid males with the around separated Sex mean. is error standard bars The error depict colour. with inred grey area dark andfished in reserve line), stippled – (2006 vertical by2013, indicated after and 2006) - (2004 reserve the of establishment to prior survey trap research theannual from and totalbody panel) (mm) length (upper (CPUE) of effort legal catch per sizedunit European lobster area. reservefished population in and thelobster of distribution sizeCatch and Figure1: (All figures arestored asseparate pdf). (1-5) Figures

Size-assortative mating. Study area on the Norwegian Skagerrak coast (circular marking) (A) marking) (A) (circular coast areaStudy Skagerrak onthe Norwegian Length (carapace distributions length, The relationship between body size (carapace length) of male of length) between (carapace size body The relationship CL is adjusted according to the year of the mating event. event. mating the year of the to according adjusted is Homarus gammarus CL in mm) of male ). Mean Accepted Article This by is protectedarticle reserved. Allrights copyright. table5). 8, size among males(Model 2010 Tableamong males (Modeland 5, 5), 2010 clawsize on relative ( selection sexual estimate each for used were to and 2) 1 (0, residuals male. matings number These of showing are among coloured in 2010. Filled circles carapace lengthandmales clawsampled width inmale ontraits Figure lobsters selection Sexual 5: European Table 4 & 5 Table & 4 ). B : Probability of mating success as a function of body of size as a of matingfunction success Probability : C: C: Probability ofmating a ofsuccess as claw function . A : Correlation male: Correlation between Accepted Article This by is protectedarticle reserved. Allrights copyright.

Accepted Article This by is protectedarticle reserved. Allrights copyright.

Accepted Article This by is protectedarticle reserved. Allrights copyright.

Accepted Article This by is protectedarticle reserved. Allrights copyright.

Accepted Article This by is protectedarticle reserved. Allrights copyright.