Posted on Authorea 7 Jun 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.162308964.47410185/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. n nlsdtedt;S n Kldtewiigo h aucit l uhr otiue rtclyto critically contributed authors All manuscript. the of writing collected publication. SI the for approval methodology; led final designed JK gave SC and and and SI drafts TH the data; Aoba-ku, SI, ideas; Kawauchi, the the 41 analysed conceived and authors Japan. All Miyagi, contributions: University, Author’s Tohoku Science, Life Japan 980-8578, of Miyagi School Sendai, Graduate address: ing address: E-mail author. *Corresponding 3 2 of 1 source the population source (S.I.) in a ITO colour Shun in shell selection snails’ Natural on title: act running Short selection does how population? origin: diversified the a in selection Natural predation. to due which not migration, Research was by that changed Original radiation was adaptive selection in natural diversification that phenotypic suggest of results were cause Our them ultimate colour. of the shell some explain mouse on and field could acted snails, Japanese predation 1,700 large an shape the than around and is the marked other predator mainland and compare factors the captured the that and showed we both selection traps total, trail-camera on natural In The speciosus detect selection island. predator. to an natural unknown an in mainland of by study the on cause previous preyed on the to experiment it land identify studies. of mark-recapture Japanese to phylogenetic strength the a traps and in using conducted trail-camera detected population then used diversified We been We of island. has population islands. it source to a although mainland in population, the known trait often diversified a not a on is has acts it of It snail However, selection source mechanisms. how radiation. these the adaptive demonstrates to becoming in study on contribute trait Our been that acts certain factors have a selection primary on speciation the natural acts of adaptive selection how one natural further is divergent selection and that Natural demonstrated diversification studies. been phenotypic of number with a radiation through clearer adaptive of mechanisms The Abstract 2021 7, June 2 1 population? diversified snails’ Ito a Shun on of act source selection the does in how colour shell origin: the in selection Natural ooUniversity Toho University Tohoku eateto ilg,Fclyo cec,Th nvriy hb,Japan Chiba, University, Toho Science, of Faculty Biology, Japan of Miyagi, University, Department Tohoku Studies, Asian Northeast Japan Miyagi, for University, Center Tohoku Science, Life of School Graduate uar eimhl simodae peliomphala Euhadra u hspeainddntcreaewt hl oor aeinapoc hwdta h tblsn eeto from selection stabilising the that showed approach Bayesian A colour. shell with correlate not did predation this but , 1 aaioHirano Takahiro , 1* aaioHRN (T.H.) HIRANO Takahiro , hssalsselclu a iesfiddet irpieslcinatrmgainfrom migration after selection disruptive to due diversified has colour shell snail’s This . 1 aoh Chiba Satoshi , [email protected] 1,2 aoh HB (S.C.) CHIBA Satoshi , 1 n uj Konuma Junji and , 1 eehn ubr 8-27576.Mail- +81-22-795-7560. number: Telephone . 1,2 n uj OUA(J.K.) KONUMA Junji and 2 Apodemus 3 Posted on Authorea 7 Jun 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.162308964.47410185/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. obndeet,adsmtms h rprin feet utaeacrigt h ere fimpacts of degrees the to according fluctuate effects of proportions divergence the traits trait snails’ sometimes, of regulate cause and a could be them effects, to many of combined shown have some been that snails also shown as have land conditions been Furthermore, such Environmental (Kraemer has 2010). it 2007). radiation, Chiba, Chiba, and competition 2004; adaptive & birds, 2009). interspecific (Barker, and Kimura 2006, preventing to mammals 2007; Crespi, in as Davison, due & such role & Parent predators, divergence a Chiba 2007; plays trait 1999b; Davison, displacement 1996, & of character of (Chiba, Chiba that cases process 1999a; demonstrated presented the (Chiba, have changes insight migrated, Studies also colour could has have shell that it and islands population determining morphology on a but (Yoder in selection acts, snails acts stabilising selection Land selection from how release natural unclear to divergent been due if often radiation population that adaptive not known source well has a (Gavrilets is it in selection Gillespie it it however, natural trait push case, 2013; as a Wainwright, then a trait such could on & certain In selection a act Martin natural on 2012). 2009; acts so Nosil, sometimes selection, Losos, population, 2000; might disruptive & island (Schluter, population as an peak such source in adaptive selection, a into surface natural an in divergent migration to fitness trait valley the adaptive after (Yoder the the of competition from occurs, “valley” especially away from this a release changed, When in a located 2016). dramatically from Losos, be resulting adaptive radiation. be & habitat an adaptive Stroud of to can 2010; expansion according including , it influence the heightens occurs, to This and due diversification trait example, certain phenotypic islandsfor a 2000). how favours that (Schluter, study influence optimum primary to a model is selection ideal Natural an are Islands recap- Mark INTRODUCTION predator, Mammal camera Trail snail, selection, Land Stabilising selection, Disruptive ture, diversification, Colour radiation, due Adaptive not was migration, that by radiation changed adaptive was KEYWORDS in selection diversification natural phenotypic other that of factors suggest cause predation. from results ultimate to selection Our trail-camera the stabilising The explain the colour. could that shell predator. showed which and on unknown approach captured Bayesian acted mouse an we A field predation by total, Japanese colour. than on In large shell preyed island. the with were the an is correlate them in predator not both study of the on natural previous some that selection detect to showed and to natural it traps snails, mainland of of the 1,700 strength cause on and around experiment the shape marked mark-recapture identify the a to compare conducted and traps then a We selection trail-camera of island. used source an We snail and the land mainland Japanese how islands. on the demonstrates to using acts study mainland population selection diversified Our the of natural that population studies. how source factors phylogenetic a simodae. known in primary in peliomphala often trait certain detected the a a not been on of on acts is has acts one selection it it selection is although natural However, divergent selection population, that Natural diversified radiation. demonstrated adaptive been studies. in has have of It speciation trait number adaptive mechanisms. these further a was to and through work contribute diversification clearer this phenotypic with becoming of radiation been part adaptive The of mechanisms Peninsula. declare. The to Izu thank interest the of also in We conflict survey no ABSTRACT have Niijima. our authors in support The survey 18H02506. to interest: our and Ito of 17K07573 support Conflict Y. Number to and Grant Maechu KAKENHI Ito of JSPS H. by Maeda Ito, supported C. Yamamoto T. and E. and Hasegawa Ito, Maeda Yamamoto M. S. T. M. Yamamoto, and S. Tatani and Yamamoto, M. T. Sentaku-ya Uchida, thank We of S. Yamazaki, comments. and D. discussions Fukuda, useful S. for Kagawa, O. thank We Acknowledgements: tal. et 09.Oeal hs atr as irpieslcinadpooedvricto ytheir by diversification promote and selection disruptive cause factors these Overall, 2019). , hssalsselclu a iesfiddet irpieslcinatrmgainfrom migration after selection disruptive to due diversified has colour shell snail’s This 2 tal. et 00.Atog tblsn eeto should selection stabilising Although 2020). , pdmsspeciosus, Apodemus tal. et 2010). , u hspeaindid predation this but Euhadra tal. et Posted on Authorea 7 Jun 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.162308964.47410185/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. hm(ireYe era&Jre 07.Te eete eesda h oain hr hywr caught. were they on where numbers locations scarifying the and at beads released plastic then coloured were on They gluing 2007). by A Jarne, shell & the standardized. Henrya (Pierre-Yves on were them marked they were way. then snails same and captured the conditions, The in light measured by also caused was differences together minor higher shell correct the to with calculated photographed was USA) NY, pixels Rochester, those value Co., of luminance values The a RGB 1995). of the (Schlick, area and value code. Cupertino, square Inc., chosen, 3 luminance (Apple a Python randomly The X monitor, using were OS averaged computer Mac images and for a cropped measured Photos On software within automatically the were box. pixels using USM light-blocking 1000 cropped f2.8 was a USA). 60-mm aperture in EF-S CA, shell an light taken the with were ring to Japan) shells Tokyo, close a snail Canon, shell under the X7; of (Canon) Kiss Photographs (EOS lens camera photographs. macro we reflex of single-lens where values island). digital RGB locations shell a the using using The on quantified and Schaffhausen, was 10J lip. Ltd., locations, colour eTrex shell shell Garmin Garmin geographic The HCx, the a conducted stage, LEGEND on and was (eTrex a growth mainland based devices survey in the identified their GPS Each on mobile was described recorded using Switzerland, adult) here. are we recorded overview or approach were snails, (juvenile an snails mark-recapture captured stage captured only the growth we include of The When we details colour. so The days. 2020), and 2–5 Konuma, 2020) 2018). km & for February October 1 - (Ito 2019 - about report (February 2017 were previous mainland the (March that on island areas month forest the every Peninsula conducted narrow were Izu were surveys was the sites Mark-recapture Island on Both Niijima Minamiizu-cho on Miyatsuka population. in Mount Niijima descendent on population including population island long. snail Islands A an Izu A population. the as ancestral to diversified. examined mainland migrated colour a Peninsula as shell Izu examined 2004), the their was Chiba, of & then (Hayashi part study and southern previous the Island, a in study to distributed to According was sites 1). that as (Figure Island respectively Niijima island, an and and Peninsula Izu the chose We experiments Mark-recapture METHODS Nijima with island, AND peripheral MATERIALS experiment a and mark-recapture 2020). mainland Konuma, a the & between conducted (Ito pressures selection Island then estimated We the predation compared of we differences Finally, islands. the and and traps trail-camera mainland using simodae snails among the the are between on radiation prey effects adaptive species the the what what of examined population, study this details This source in the the is what selection on and natural acted islands, snails. the has & and land of pressure (Hayashi mainland these cause selection the selection the what between what disruptive unknown are unknown also to is differences is due it to However, it monomorphic occurred migrated Furthermore, 2020). have have rapidly Konuma, case. mainland populations & has sympatric the Ito islands’ mainland diversification on 2004; peripheral The shell-colour distributed Chiba, the populations where in Islands. Ancestral observed islands, Izu is 2004). the dark the Chiba, to of & bright (Hayashi several from populations diversification as colour well while as shells, snail bright Peninsula, land Japanese Izu the the in on on studied studies been no also has almost snails simodae are population. of there source diversification colour but the shell in radiation, The adaptive determined are in traits focus Kraemer diversified a 2004; the Chiba, been how & has (Hayashi diversification island phenotypic the cases, of age the as such Y/Y Hysi&Cia 00 04 t oua 00.Ti ni naistemiln fJapan of mainland the inhabits snail This 2020). Konuma, & Ito 2004; 2000, Chiba, & (Hayashi ntemiln n siae h aua eeto rsue rmpeainadohrfactors. other and predation from pressures selection natural the estimated and mainland the on 0 au mle rgtrselclu,weesalower a whereas colour, shell brighter a implies value Y fteselclu a acltda follows: as calculated was colour shell the of Y 0 ftege cl mg KdkClrGe cl;EsmnKodak Eastman Scale; Grey Color (Kodak image scale grey the of 3 tal. et .p simodae p. E. Y/Y 09 t oua 00.I many In 2020). Konuma, & Ito 2019; , Y 0 au mle akrselcolour. shell darker a implies value 0 = . 299 ouain ntemainland the on populations R 0 + uar peliomphala Euhadra . 587 .p simodae p. E. G 0 + . Y/Y 114 .p. E. B 0 Posted on Authorea 7 Jun 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.162308964.47410185/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. 02.W sue httecoefficient the that assumed We 2002). calculated (Gimenez 2 Arnold, Equation second- & First, with (Lande used fitness. 1 was of Equation (P-spline) index with an regression gradients as spline Gimenez selection rate penalized 2003; survival the Brooks, the estimate & used to Blows we used 1983; 2020), was Konuma, regression & polynomial (Ito mainland order study in previous a colour videos in these shell As camera from of The identified selection were system. natural predators trigger were of The traps motion Modelling respectively. camera a these sites, 5 on study and At 4 island operated laboratory. and and range. our 2018. mainland sensor in tripod camera’s February the a the at to entered USA). on up animal 0.2 Kansas, 2017 or set an diameter Ltd., tree when April Japan; Bushnell videos a Hiroshima, from ESSENTIAL, recorded to Ltd., automatically HD3 site attached (Daiso (TROPHYCAM was study strings cameras camera 5-cm-long trail island Each with from using the trees site monitored to at study and secured mainland mm) and were the 2020 snails at sites, conducted July study were to experiments a 2019 camera-trap as predators, December used and snails’ was 0), the recaptured identify juvenile were To snails 1; dead (adults 2020). Team, the stage non-predation Core when growth and (R month colour, (Bates (1) R The shell “lme4” Predation effects. the (GLMM). fixed effect. while as model random variables, stage, used mixed growth response were and linear interaction binary colour their generalised as shell rather with a used Fisher’s predators associated using with were was by island (0) them predations the attacks of and analysed from mainland number also were the the died between whether we that snails they examine predated snails To that of boring, recaptured number test. considered shell the Some exact we compared crushing, We so shell 2000). age. entry, shell-crushing, old Robinson, shell than of & as evidence classified DeWitt showed are 1979; dead gastropods (Vermeij, attack swallowing to whole-shell use and predators that predators modes and of The University identification Tohoku and of pressures regulations predation experimentation Measuring animal the to according University. Toho conducted was study This scale. equations, these In ( t variance and 0 mean with when distribution selection stabilizing the of that γ h ubr fknots of numbers The . col stennierslcingain,wihrpeet h tegho h irpieslcinwhen selection disruptive the of strength the represents which gradient, selection nonlinear the is P i − β stedge ffedmi h -pie hc a e to set was which P-spline, the in freedom of degree the is κ stelna eeto rdet hc ersnstesrnt n ieto fdrcinlselection. directional of direction and strength the represents which gradient, selection linear the is k ) P k/ hn(co when ( K 1) + logit unie yuigalvle fselclu,with colour, shell of values all using by quantiles f l K i ( − ( co sdcddacrigto according decided is t x logit κ 1 = ) l i k = ) means ) tal. et , ( 2 ≥ S µ , . . . , i,t 0 + logit = ) log r0otherwise. 0 or 07 a sdwt oi saln ucini h ttsia package statistical the in function link a as logit with used was 2007) , β < γ 12; 1 σ tal. et [ ( · x/ ε µ 2 S col 0 b Rye 2008). (Royle, i,t + i (1 k . 1 = i = )  of 09) ovsaietesria aeacrigt hl colour, shell to according rate survival the visualise To 2009b). , − β i + ersnsteidvda eeoeet,wihflosanormal a follows which heterogeneity, individual the represents · , x ( 2 · · · col K co )] f , . . . , . ( l µ i = + i co κ + − k steoealaeaefrtesria aeo h logit the on rate survival the for average overall the is l β min 4 i represents 1 2 κ p + ) 74( 1714 k · · B ) col γ ( + P t  I 4 · i olw omldsrbto ihma and 0 mean with distribution normal a follows stemnhyeet ntelgtsaefrmonth for scale logit the on effects monthly the is , i P col + nteIuPeninsula Izu the in 35) + i 2 B + ow e to set we so , P k X k K t =1 sfie nt,with knots, fixed ’s 3 = (2) #  i b + k k ( ayn ewe n 5(Ruppert, 35 and 1 between varying nti study. this in co B l t tal. et i (1) # − K κ k 06;b). 2006a; , 35 = ) ) + P κ ( Rpet 02.We 2002). (Ruppert, co 1 l κ < i − κ 2 k < ) + P · · · > γ seither is κ < 0 or k . Posted on Authorea 7 Jun 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.162308964.47410185/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. er-nonee 3 ftesal eesdi h ed(5 iigad8 edsal) erecaptured We snails). dead 80 and living (257 field total. in the snails in 1714 released released and snails marked the we of mainland, the 337 on re-encountered conducted We experiment mark-recapture pystan the used In we simulations, MCMC the natural For visualise RESULTS to summary. by distribution model 3. confirmed the posterior Python was In the in simulations (https://github.com/stan-dev/pystan) for all iterations. used (Gelman with 1,000 was each convergence statistics of parameters were of The burn-in Rubin that distribution initial required. simulations and posterior were the MCMC Gelman and the simulations 2,000 following instability obtain using MCMC five To numerical chains 2,500 of Markov avoid selection, 1996). rate three to Roberts, a generated MCMC & then used at the we colour (Gilks thinned using models, shell chain our The before each in prior z-score of 2020). parameter non-informative Konuma, the mixing a & by the used (Ito improve standardised also study we to was previous selection, models a natural in our visualise and described in parameters estimate parameters estimate to all to model with framework Bayesian the distribution the In in model. simulations each (MCMC) in Carlo Monte chain Markov 2012). used model We Schaub, Bayesian & the Kery S3-S4; in (Equations parameters addition, distributions Estimating In categorical the function. with variance modelled logit defined and We a were was 0 matrices function 2020). observation mean link parameters In Konuma, with the These & adults. distributed which in and heterogeneity (Ito normally framework, juveniles individuals. individual colour GLMM for dead shell same a same using marked to the recapture the colour the according for were shell vary matrix, they was of could this effect although rate months, parameters In recovery survey died these (3) the between snail. that addition, snail predation, differing marked and (5) adult unrecovered as than individuals, an snail, defined or of other adult live recovery were snail living causes (6) for a undiscovered predation, from of was an than recapture died other rate (7) or causes that capture and from snail first died predation, (4) that juvenile from predation, snail a from adult recapture died an of or of that recovery capture recovery snail first (2) juvenile (1) a snail, S2): of was juvenile recovery (Equation. rate living states observations modelled fidelity a (see seven was site defined of rate snails we the transition matrix, marked observation The constant and the among 2020). be fluctuation, In Konuma, mortality to monthly model. & have of assumed effects (Ito and to was study random causes previous juveniles a rate assumed the a with was predation of applied to comparing rate was The rate according from rate constant transition predation survival purpose. considered month The The the our and adults. to results). adults, colour to according the and juveniles shell separately from juveniles of 2 rate of or regardless transition an rate the 1 as and survival Equation site rate, to the study fidelity adult. the predation defined site an outside than the as we predation other adults, site than survival matrix, study factor (10) other juvenile, the factor the a a outside a from In as predation from site dead from dead study (8) (11) dead site the adult, juvenile, (12) outside study a predation an the and a from from as within adult, as dead dead site predation (9) site (5) study from juvenile, adult, the dead study dead from a an outside (6) as the (2) dead as site adult, juvenile, (3) outside site study an juvenile, survival a the study as a outside (7) as the site as within adult, site study site survival the an study study (4) within as the juvenile, the predation a within within than as survival temperature, other defined site factor as (1) we study such model, the S1): predation, this within than In predation (Equation. other 2020). states factor Konuma, 12 a & “dead Natural Marescot (Ito from with state: 2004; model matrix 2. 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No reuse without permission. | https://doi.org/10.22541/au.162308964.47410185/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. niseitdsmarcly h eei eainhpo h niswscoeo h sad n yrd of hybrids and island, the on close was brighter snails and (Wade, why darker the occurred which reason of in isolation The juveniles. bimodal, relationship post-zygotic was low. genetic than that colour is The shell adults be of rate might sympatrically. for distribution survival island The existed higher the the snails 2020). was when mainland on Matute, island the colour only & on Coughlan juvenile shell the adults rate 2002; in on on survival on acted act The that acted selection only mainland. whereas selection natural might the natural juveniles, of selection no that than natural to and adults Therefore, opposite island, among is island. the pattern pressures lower the on This selection and was juveniles the mainland 2020). on Konuma, of the & only difference between (Ito diversity The acted colour selection island. shell disruptive the of which Strong on difference snails, dimorphic the intermediate-coloured stabilising is in over variation The result snails colour then bright shell mainland. predation. would and the both the than on dark Hence, why monomorphic favours other is be predation. variation selection factor could colour than disruptive shell a other the the by factor why contrast, be a it caused In could Thus, also which are snails, is island. bright selection the selection favours on disruptive selection disruptive predation low of and relatively exclude cause was selection not predation the previous did In stabilising from that a method shells 2020). suggested from Bayesian snail Konuma, been & dead differs the (Ito of has result from island proportion This estimated the the However, on selection mainland. colour disruptive the effects. shell selection the on the stabilizing study, predation on weak previous acts than that the selection other suggests disruptive 4A; highest factor that the (Figure a showed which being from island study, snails colour the bright-shell shell on of on rate than acts survival higher the is of result pressure (Figure The predation island bright-shell showed where the of colour mainland, on effects shell dimorphic the predation was the excluding 2). on Table rate it of highest survival while the distributions the mainland, was that frequency showed the snails The experiments on mark-recapture persistence island. monomorphic The the the was pressures clarify 2). variation and selection To and colour mainland variation 1). colour the the Table the that between compared 3; then snails (Figure we mouse, snails, the colour field in shell of colour Japanese shell with diversified large correlated the a of not processes only was frequency demonstrated predation first the we study, this In BCI 95% the and DISCUSSION 4A), (Figure monomodal not were S2-S4. snails juvenile mainland of of rates Bayesian survival the the on contrast, based of simulations (BCI) gradient MCMC interval selection the estimated was nonlinear in rate the survival colour snails of adult γ shell adult median The of 4A). the mainland (Figure function and of colour framework, polynomial effects shell second-order of predation terms a excluding in rates using monomodal field survival but Japanese the bimodal large not that were only showed the regression these, P-spline Of The and S1). birds, (Table mainland. four range the mammals, camera on eight snails’ ones filmed monitoring juvenile mouse and the than predators = snails within the z-value adult reptiles identify 1; more three to (Table attacked experiments than predators sites trail-camera mainland that study conducted implies We both the and result in on This 37 colour predated 1). were shell (Table were the There snails with 9.46, correlated = More 3). significantly 2.86, odds (Figure not respectively. test, was predators island, Fisher-exact snails by and predated S1; crushing mainland (Figure colour shell the island shell of on the the evidence snails while mainland had 2A), such the snails (Figure 8 on unimodal dead snails was 2B). the recaptured colour (Figure of Some shell Most bimodal was the times. island of four the distribution was of number the distribution maximum and the shells, and bright once, had than more snails some uprsta h -piergeso uv a ocv pad lhuhte9%Bysa confidence Bayesian 95% the Although upward. concave was curve regression P-spline the that supports γ nldd0(al ) h te aaeesetmtdfo h aeinmto r hw nTables in shown are method Bayesian the from estimated parameters other The 2). (Table 0 included P pdmsspeciosus Apodemus .5.Hwvr h ubro rdtdsal a orltdwt rwhsaeo h mainland the on stage growth with correlated was snails predated of number the However, 0.05). > γ a -.7 .3,9.%of 94.2% 0.33], [-2.47, was a takn h nis(iue3;Mve 1S nDryad). in S1-S3 Movies 3C; (Figure snails the attacking was γ ausetmtdi h iuainrn eengtv.In negative. were runs simulation the in estimated values 6 P .0) nteGM nlss h ubrof number the analysis, GLMM the In 0.001). < γ a 10 Tbe2.Tengtv au of value negative The 2). (Table -1.05 was .speciosus, A. rydo nis but snails, on preyed Posted on Authorea 7 Jun 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.162308964.47410185/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. uprigMveS-3 orecd n aaue ntesuyaeaalbefo h ra Digital Dryad the from available are study GitHub. the the from in available are used mark-recapture REFERENCES data in used and codes code stan stage. The source initial Repository. an S1-S3, in Movie ultimate especially the radiation, Supporting explain adaptive could of study process colour the this shell STATEMENT AVAILABILITY in in DATA then diversification presented colour and have as shell we occur, such such that can of predation, selection radiation factor Kraemer than disruptive 2006; adaptive other to Chiba, islands, factor selection & a on stabilising was (Davison was snails which diversified selection land population rapidly island, natural is some the without source changing changed on In of be the colour effects. could cause in selection shell environmental direct natural on persisted whether the unknown acted effects, colour is selection it predation shell disruptive Although population. contrast, monomorphic derived In in variety that diversified a selection. of suggested stabilising use to findings the due triggered our have diversification. could colour conclusion, predation shell In from occurs then release predation and a from microhabitats (Losos Islands, release or Izu predation a habitats the Constant when avoid of In diversified 2016). to is Losos, monomorphism. trait & use (Stroud restricted shell-colour microhabitat a in contrast, or In persistence Svanb¨ack, habitat (Ekl¨ov 2006). & of restricted driver in Lapiedra indirect result than & an (Inoue adults can ground among be the predation observed on could frequently eggs more predation lay to been However, trees have from would from down all mainland. predation risk come ground the of predation to the on constant evidence need on juveniles face the snails dwell survival. would adult so and the The snails 1981), trees reduces mainland Nakada, climbing ground. that the the at factor on a Thus, poor not only mice are 1981). be field do mice Nishikata, can pressure mouse field 1978; mice that predation field this (Imaizumi, from reported like so year predation studies (Rosin rodents 2007), that previous (Conway, colour suggested fact, addition, field types is shell In In it the cone of so two regardless so shells. idea, with this randomly bright experiments, with blind occur or the mark-recaptured colour could dark of the mice are number on in from they The based colour because snails Our snails. shell colour predate the mainland. the recognise to the of with on unlikely predators rates correlated main are survival the not mice the were was reducing mice in snails field factor predated primary that showed a be experiments would trail-camera pressure in the predation polymorphism than high of an lower The is case in possibility the that camouflage this snails in of Hence, the as advantage on experiments. preyed snail, an 2013). trail-camera birds this has our no for Although in colour hypothesis filmed effect. shell environmental were opposite brighter birds the have some a would studied, example, shell we darker for a alternatives scenario, while (Allen an habitat, to predation diversification is open lead birds a colour feature and from shell In geographical habitats predation promote this of contrast, 2019). Thus, predators of heterogeneity In bird altitude 1). as 2012). because The such (Figure of Schilthuizen, hypothesis a conditions mainland & (O˙zgo environments. function environmental while the diversification closed complex a environments, than colour in create islands has open shell might sunlight the in colour islands less on sunlight the shell to higher to of that is exposed exposed area when is when distributed advantageous selection the temperature be natural could body of optimum shell them. cause darker an between maintain the incompatibility on to for genetic snails only help Lellis a hypotheses of exists is (Di all the shells there regulation bright almost of since thermal and which rate one dark survival in lower between In adults, mainland, hybrid a In have 2020). the the they Konuma, that on that implied & colour and results Ito shell island These 2004; of the colour. Chiba, distribution brighter & unimodal a (Hayashi were a colour was intermediate there showed contrast, snails brighter and darker tal. et 08.I uhacs,atati te oteaatv piu codn otehabitat the to according optimum adaptive the to fitted is trait a case, a such In 2018). , tal. et 02 Schweizer 2012; , 7 tal. et tal. et 09.Freape h rgtrselcould shell brighter the example, For 2019). , 09.Tecag fntrlslcinfrom selection natural of change The 2019). , tal. et 01.Orrslsaeconsistent are results Our 2011). , eaanemoralis Cepaea tal. et 98 Kraemer 1988; , tal. et (Schilthuizen, 2004; , tal. et , Posted on Authorea 7 Jun 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.162308964.47410185/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. rs oaRtn FL. Raton, In: carlo Boca convergence. monitoring Press monte and simulations chain from markov Inference 2011. of Others. & data. K. 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No reuse without permission. | https://doi.org/10.22541/au.162308964.47410185/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. )tesellf fe ri aeacpue cn that scene a captures camera trail a by after predated left shell the B) and that dark scene the value bimodal, the Peninsula, showed luminance Izu it 3. the the whereas Island. In Figure using existed Niijima quantified colours. colour in brighter existed shell was representing colour bright colour values shell mainly higher bright Shell and with unimodal snail, showed 2020). the distribution Konuma, of photograph and a from Ito using Island from Niijima and (modified Peninsula Island both Izu on the 2. conducted of peninsula, was cross-section Figure Izu survey The the The C) In B, Island. side. Island. Niijima west Niijima including the and Islands Peninsula Izu Izu the the and (mainland) Peninsula Izu 1. Figure Deviation Standard SD: heterogeneity Table2. and Peninsula Izu the in adult) and (juvenile stage growth and colour shell with Ecolo- Island. predated) Niijima 2010. 1: others, L.J. the Harmon, (0: & Jochimsen, S.F. radiations. J., 1. Table Hagey, Spear, adaptive W., J.J., of Godsoe, Schenk, origin epistasis. L., B.A.J., the Gentry, of https://doi.org/10.1111/j.1420-9101.2010.02029.x Sarver, and J.M., view J., opportunity Eastman, eye Robertson, S., gical gene’s B.P., Roches, A Oswald, Des speciation: D., E., and Clancey, selection J.B., epistasis, Yoder, of view biology. eye evolutionary gene’s of Journal A 2002. M.J. Wade, eut fgnrlzdlna ie oe nteascaino as fdahi akdln snails land marked in death of cause of association the on model mixed linear generalized of Results )Mp fsuystsi h z Peninsula. Izu the in sites study of Maps A) siae ierslcingradient selection linear Estimated mgso rdtdepyseladtelreJpns edmouse field Japanese large the and shell empty predated of Images rqec hl oordsrbto nmre nisi )teIuPnnuaadB Niijima B) and Peninsula Izu the A) in snails marked on distribution colour shell Frequency .speciosus A. σ 2 .speciosus A. sn h aeinmdlo ahgot stage. growth each on model Bayesian the using ijm sadItret-3.17 Intercept Island Niijima z eisl necp -1.24 Intercept Peninsula Izu tg aaee einS 5 C 5 BCI 95% BCI Adult 95% SD Median Juvenile Parameter Stage on nmr-eatr uvy thscaatrsistesltigo h pxor apex the of splitting the characteristics has It survey. mark-recapture in found rdtdalvn ni hc a ehrduigsigi rn fatalcamera. trail a of front in sting using tethered was which snail living a predated 15 neato .506 0.55 0.60 0.55 0.49 -0.15 Interaction Stage Growth 0.09 Colour Shell 0.28 1.61 Interaction Stage Growth Coefficient Colour Shell effect Fixed σ γ β σ γ β 2 2 3–4.http://doi.org/10.1046/j.1420-9101.2002.00413.x 337–346. : .p simodae p. E. .733 .114.8 0.33 1.47 3.11 12.9 3.32 -2.47 0.73 -1.17 0.70 0.73 0.21 0.66 9.77 3.63 -1.05 -1.35 0.01 -1.35 0.54 3.68 0.54 -0.34 -0.16 β sihbtdol nsuhr ato it. of part southern in only inhabited is 11 n olna eeto gradient selection nonlinear and uar eimhl simodae peliomphala Euhadra ± ± ± ora feouinr biology. evolutionary of Journal ± ± ± ± .speciosus A. .705 0.57 0.57 0.87 0.52 0.87 0.16 0.64 0.55 0.43 .8-.6<0.05 < -2.56 0.48 .3-.1<0.001 < -3.41 0.93 .3-.80.86 -0.18 0.83 .628 0.001 < 2.86 0.56 ± .%97.5% 2.5% EZvleP-value Z-value SE rydo ni.C mt shells empty C) snail. a on preyed pdmsspeciosus Apodemus sdsrbtdo the on distributed is γ 23 n individual and 1581–1596. : A) . Y Posted on Authorea 7 Jun 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.162308964.47410185/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. rdto ffcs h oi n ahdlnsidct h einadte9%Bysa ofiec interval, simulations. excluding confidence Carlo Bayesian after Monte 95% chain estimated the natural Markov was and from the selection median represents obtained natural the distribution shape indicate the This posterior lines Peninsula, the dashed adult. Izu in and is respectively, the solid II) In The juvenile, effects. is colour. predation I) shell 2020); on Konuma, worked and selection Ito from (modified Island 4. Figure shell. the of shattering the h eainhpbtensria aeadselclu nA h z eisl n )Niijima B) and Peninsula Izu the A) in colour shell and rate survival between relationship The 12 Posted on Authorea 7 Jun 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.162308964.47410185/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. 13 Posted on Authorea 7 Jun 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.162308964.47410185/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. 14