Posted on Authorea 22 Jun 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159285541.11256769 — This a preprint and has not been peer reviewed. Data may be preliminary. aze Liu Wanzhen southern of regions montane China F.P. the chuniana to endemic in () cycles Metcalf glacial to linked dynamics Population atAi umrmnonadcniuu aia vial o cuain ope hsorpyplus physiography Complex occupation. for available intensified habitat with continuous China, simula- in coalescent and and period monsoon interglacial modeling Lushan-Tali summer niche geographical warm was Asia ecological the as by isolation East during function inferred Geographical occurred to As contact thought east. Pleis- physiography. secondary ranges, the complex middle tions, mountain to their the adjacent those to 11 in in from due among located separated barriers occurred variation populations Mts. divergence genetic the Nanling of between population the detected pattern that in spatial revealed populations dynamics. the when estimation population investigate tocene, time its to elucidate Bayesian dis- (GBS) on to narrowly sequencing China, focused populations. of subtropical we by evolution in climate Here genotyping ranges the Pleistocene used species. mountain affected We southern widespread geo- to have the more the response China to than driving endemic southern in studied species in processes less populations ranges the is mountain investigating species of key tributed for radiation the system How and excellent biodiversity fluctuations. an of are distribution China graphic southern of mountains The how ecosystems. understanding montane are our China approaches advances southern Abstract genomic study in population This species plant that of species. shows history distributed work evolutionary narrowly Our the occupation. of affected for have dynamics expansion began. available cycles demographic population migration glacial habitat the the contemporary continuous facilitated detecting which cycles and Lushan-Tali As in from glacial effective warm monsoon Mts., across physiography. the summer Nanling conditions complex during Asia the ecological their occurred stable in East to long-term contact intensified due plus secondary with physiography barriers simulations, Complex China, geographical coalescent in populations as and the variation period function between modeling detected interglacial genetic to niche was of isolation thought ecological Geographical pattern by when ranges, east. Pleistocene, spatial inferred mountain the middle elucidate to the adjacent the to those in investigate in from China, occurred separated to located divergence subtropical Mts. (GBS) population in Nanling that ranges sequencing the revealed Here in mountain by estimation populations southern time species. genotyping Bayesian the widespread used populations. to more We 11 than endemic among studied species dynamics. less woody in is ranges population a species mountain chuniana, its key distributed of Cercis the narrowly distribution How on of geographic fluctuations. focused evolution the climate we the driving Pleistocene affected processes to have the response China investigating in for southern populations plant system of excellent radiation an and are biodiversity China southern of mountains The Abstract 2020 22, June China 510650, Sciences, of Academy Chinese 3 Garden, Botanical China 2 1 Gong Wei and oaia eerhIsiueo ea,10 nvriyDie otWrh X717 USA 76107, TX Worth, Fort Drive, University 1700 Texas, of Institute South Research Utilization, Botanical Sustainable and Conservation Resources Plant of Laboratory Key University Agricultural China South 1 inun Xie Jianguang , 1 1 u Zhou Hui , 1 agu Kong Hanghui , 1 2 agHao Gang , ecschuniana Cercis 1 ee Fritsch Peter , woody a , 3 , Posted on Authorea 22 Jun 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159285541.11256769 — This a preprint and has not been peer reviewed. Data may be preliminary. nulse aa 00.A ihiscneesi hn,i xiisa dpaint ei environments mesic to adaptation an exhibits al the it et (Fritsch near China, apex resolved in blade congeners is leaf its acuminate species with its The through As morphologically. 2020). identifiable data, easily unpublished it the and Wuyi makes of the which and base from sizes 1940), extending population Metcalf, larger distribution, increasingly 2010; geographical with narrow Mountains among a Nanling Unique the has densities. to It westwards China. Mountains species. widespread China southern et Eastern distributed on of Liu narrowly focused STEBF of 2016; has the evolution al., research to the with this et affect involved of might (Gong chuniana most scenarios ranges China Cercis However, mountain the southern 2020). key elucidated Wang, how in has & ignored species Bao, research and Wang, plant species Previous Ye, have of 2015). Tian, expansions history al., 2012; range evolutionary al., et rapid postglacial Tian or and 2011; localized glacial Comes, subsequent & which diversity Fu, of genetic al Qiu, of out et genetic centres China, (Chen with and correlated inferred southern isolation refugia been in glacial geographical elevation Multiple identified both in 2006). habitat for been Su, lowering of & opportunity have belts Cui, degree vegetation the (Shi, the providing with occur China, to range, periods, southern divergence geographic glacial 105 In during in 1980). to contracting decreased Wu, (east have & and to China Pu, now thought (Duan, eastern are is Glacials data in Tali connectivity considerable and periods debate, Lushan under glacial Dagu, still Poyang, four populations, Although support al of 2009). Skov, to history et & available demographic Li Normand, the 2004; Svenningm, shaping 2018; (Hewitt, important and Suding, an regions biodiversity as montane considered of in also pattern are particularly distribution and Pleistocene the the biodiversity during driving al of cycles fluctuations. factor glacial et distribution climate with (Li geographic Pleistocene associated cycles oscillations to the Climatic glacial response driving in Pleistocene processes populations and the plant events investigating of orogenic radiation for both arising system presumably with excellent 2011) line 2011; Ge, an in of al & Comes, rates years, well et Ying, high & Liu million Sun, and exhibit L´opez-Pujol, Fu, 1979; Zhang, 5 endemics, 2017; world subtropical predominantly last (Qiu, al., the is species, et the biota plant in (Hou mountains within their north-temperate radiation forests these of rapid the Many such of and of 2012). continuous local type Shi, elements largest vegetation & relictual Tang, the primary Fang, ancient Wang, Zhong, of The harbouring & one for Zhang, 2012). (STEBF), known Chen, complex 2016; Shi, forest to Zeng, al., attributed & broadleaved (Qiu, et been Tang, gradients evergreen has (Gong elevational uniqueness Fang, habitat species steep Wang, local plant or Their 2013; various longitudinal 2019). to for with Zhang, suggested (Hou, correlated & corridors Wu, been directions topography colonization Xiong, have northeast-southwest 2018; or ranges al., or diverse barriers et topographically north-south Tian geographical these either either 2001), in to as Ying, oriented part serve 1992b; large Often Wang, al in 1992a; et Wang, due 2016). (Fan is 1983; ranges Huang, China mountain & southern its Yao, Ruth, of of Dick, & biodiversity heterogeneity (Colin high physiographical isolation The geographic extreme 1964). of the episodes Simpson, because frequent habitats 2019; by montane Muellner-Riehl, afforded in 2006; opportunities diversification ecological rapid increased prompt the to of suggested is heterogeneity physiographical High INTRODUCTION ranges mountainous China China Southern southern contact, in secondary species plant our of advances KEYWORDS: history study This evolutionary the species. affected distributed approaches ecosystems. have narrowly genomic montane of cycles population dynamics that glacial shows population how work the Nan- understanding Our the detecting in began. in expansion effective migration demographic contemporary are the which facilitated from cycles Mts., glacial ling across conditions ecological stable long-term Cercis . 03 h,L,&L,19;Wn tal et Wang 1998; Li, & Li, Shi, 2013; , .P ecl Fbca:Crioda;Aaie l,21)i ml reo hu endemic shrub or tree small a is 2017) al., et Azani Cercidoideae; (Fabaceae: Metcalf P. F. ecschuniana Cercis hlgntcte,wt netmtdaeo .0M Fishe al et (Fritsch Ma 2.40 of age estimated an with tree, phylogenetic Cercis . 02 og hn oe,F,&Kc,20;L,Sa,L,Zag i,2012; Qiu, & Zhang, Lu, Shao, Li, 2008; Koch, & Fu, Dobes, Chen, Gong, 2012; , pce,i a naymtia efbae(hn hn,Lre,&Vincent, & Larsen, Zhang, (Chen, blade leaf asymmetrical an has it species, eorpi oeig egahcioain litcn lca cycles, glacial Pleistocene isolation, geographic modeling, demographic , . 08 sl,17;WnelnLre,&Lre,1981). Larsen, & Larsen, Wunderlin 1975; Isely, 2018; , 2 . . 00.Teecaatrsismk otenChina southern make characteristics These 2010). , 09 eqia ilan end oeod & Rosemond, Bernad, Tillmann, Mesquita, 2019; , º )drn h litcn,ie the i.e. Pleistocene, the during E) . 08 ue l,21;Yang, 2017; al., et Xu 2018; , . 08 i tal., et Liu 2018; , . , Posted on Authorea 22 Jun 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159285541.11256769 — This a preprint and has not been peer reviewed. Data may be preliminary. ihtetsu ooeie iseye-6(hnhiJnxnIdsra eeomn o,Ld.Total beads (Doyle glass method Ltd). with (CTAB) Co., bromide tubes ammonium Development in trimethyl ground cetyl Industrial was modified Jingxin the tissue with (Shanghai Leaf extracted Tissuelyser-96 was -80. 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Glacial resolution. the and Last for Ma), Ma), inferred ( data 0.006 0.12 paleoclimate distribu- ˜ [?] high-resolution spatial 0.14 on (MH, the analysis (LIG, characterize the Interglacial to based Last 2005) We Peterson inferences. for & biological conditions historical Soberon suitable (ENM; of modeling tion niche ecological used We and individuals through modeling NLW1 20 Mts. collected niche Wuyi whereas also We Mts., the Ecological 1). Nanling between Table 2.2 1, the located (Figure of is Mts. of part Nanling LXS2 population Mountains. the respectively. eastern and of one Mts., the Nanling part Mts. from Wuyi western in Luoxiao the and the located in the of are Luoxiao, located of parts are NLE2 Wuyi, NLW5 from western end and the and collected southern NLE1 eastern were Mts. the including the Luoxiao populations in China, in five located located southern to is are one LXS1 in WYS2 from ranges and Anywhere WYS1 mountain 1). Populations Table the of 1, of (Figure individuals each species 112 the and of populations distribution 11 collected We narrowly of and sampling history Population China evolutionary 2.1 the southern on its in METHODS environment ranges elucidate AND this MATERIALS mountain 2) of 2 impact of species, the roles hoped the of thereby the We of understanding species. distribution. infer distributed structure better geographic to and a population diversification gain data its and to driving of the diversity in analyses use fluctuations genetic genetic climatic 3) population Pleistocene investigate nonmodel and for 1) SNPs history, for to genome-wide analysis demographic collected aimed genome-wide and We a GBS . for al used et genotyping, we Davey tool and 2016; Here, cost-effective Hohenlohe, discovery 2017). & Luikart, genome and marker Miller, reference with high-throughput, Good, a genetic (Andrews, combination al require rapid, species investigating In not et a does (SNPs). Niu for GBS provides 2010; polymorphisms approach enzymes, Metzker, GBS nucleotide restriction genomic 2017; with single Li, reduction a detect & genomic as to on Pang, based Zhang, used is Hou, widely it (Chen, Because been structure population has and (GBS) diversity sequencing by Genotyping http://worldclim.org/download .chuniana C. .chuniana C. rmnn ebra(,IE,IK BC U,LG MH EadSF)a well as SCFI) and PE NMNH, LBG, KUN, IBSC, IBK, IBEC, (A, herbaria nine from .chingii C. .chuniana C. and ik&Hjas 07 o h ordffrn tgswt .-iuespatial 2.5-minute with stages different four the for 2017) Hijmans, & Fick ; hnlctdi hceghni hjagProvince. Zhejiang in Chichengshan in located Chun .chingii C. n oaeptnildsrbtoa ra ncnucinwith conjunction in areas distributional potential locate and eesmldadpae nocnrfg ue,wihwere which tubes, centrifuge into placed and sampled were 3 .chuniana C. rmtruhu h urn geographic current the throughout from . 01 cee,Bte,&Edwards, & Batley, Scheben, 2011; , . 06,weesoe higher scores where 2006), , .chuniana C. . 2019). , Posted on Authorea 22 Jun 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159285541.11256769 — This a preprint and has not been peer reviewed. Data may be preliminary. mlmne nFsSrcuev. Rj tpes rthr,21) eue h eal etn with setting default the (MCMC) used We model Carlo 2014). Monte of Pritchard, individuals Chain & 112 Markov the Stephens, Bayesian on of cross-validation (Raj, a analysis 10-fold v1.0 with hierarchical FastStructure 1984). structure a in Cockerham, genetic implemented and & (AMOVA) Weir population variance 1996; estimated Weir, molecular We 1992; of Quattro, 0.1.16 & analysis vcftools Smouse, used (Excoffier, with we subdivision calculated populations, ( were populations within among SNPs and differences of selected values pairwise the calculating for frequencies ( allele heterozygosity and al et alleles (Petr of number admixture The and assignment population diversity, al Genetic et trees (Bouckaert ( 2.5 remaining v.1.4.3 v2.4.7 FigTree TreeAnnotator the with with edited from intervals were 2014). chronogram trees time Rambaut, final (MCC) confidence The (ESS) credibility 95% BEAST. size in and clade 2014) sample heights maximum effective mean the the nodal that generated with (Rambaut, check then v1.6 to for four Tracer and and BEAST and used convergence option, We in assess model simulations to generation. I MCMC 2014) 1000th + was Drummond, the every and G & sampled ran uncorrelated + Xie, parameters We with Suchard, GTR the with setting covered branches. the Clock generations which Strict the million with the Ma, 10 across conducted and 0.2 variation were prior, of rate Population analyses Constant deviations log-normal-distributed time Coalescent standard divergence a and The categories, distributions rate range. prior HPD normal a 95% with point calibration chuniana C. ( al PhyML-SMS substitution et (Fritsch in nucleotide within 1974) of times Akaike, model Divergence (AIC; The criteria information 2014). Akaike (Alexandros, Gateway the montpellier.fr/phymL/ v8 Science with RAxML CIPRES selected the in was in pseudoreplicates GBS al available the bootstrap cluster et on ML computer Miller based high-performance extracted (www.phylo.org; the SNPs 3.3 on the performed using were trees yses phylogenetic generate of to populations dataset. analysis 11 (ML) the likelihood among mum relationships phylogenetic estimation reconstruct 50% time To of divergence heterozygosity and at observed population locus. of analysis maximum For rate per Phylogenetic a genotyping populations a 2.4 Stacks. specified five and also with of 0.05 We calling number least at minimum populations. SNP of a within for (MAF) and individuals frequency individuals allele of 132 minor 80% by a used least with We assembled SNPs were extracted sorting. we of format and analysis, individual fastq one mapping DNA with in sequence (Illumina, and 2013) Reads for analyses Cresko, platform advanced & sequencing Further reads. Amores, HiSeq Bassham, low-quality sequencing. glabra Hohenlohe, Illumina remove 150 (Catchen, The to v2.2 (PE) STACKS paired-end performed using selected for Ltd. were were used fragments assembly was Co. desired library USA) Novogene the of CA, in and combination Diego, constructed construction San a was library with amplification GBS multiplex digested for after was barcodes libraries DNA to representation genomic ligation reduced al The generating et for workflow (Heffelfinger 2016). streamlined sequencing a Illumina is for (GBS) of sequencing Scientific, concentration (Thermo by DNA spectrophotometer Genotyping final Nanodrop a a and with USA), quantified was CA, concentration Carlsbad, DNA 1986). Doyle, & > ap srfrne W 078(i&Dri,20)adSMol 131(ie l,20)wr used were 2009) al., et (Li v1.3.1 SAMtools and 2009) Durbin, & (Li v0.7.8 BWA reference. as Pamp. 0 o ahprmtr edsaddtefis 0 ftesa uni ihtema oeheights node mean the with burn-in as trees of 10% first the discarded We parameter. each for 200 ecschingii Cercis . 01.T esr eei iest,w siae xetdhtrzgst ( heterozygosity expected estimated we diversity, genetic measure To 2011). , . eue ES 247(ocar tal et (Bouckaert v2.4.7 BEAST used we , 08 i tal et Liu 2018; , Ho .W sdAlqi 35(xoe ice,21)t siaegntcdffrnito by differentiation genetic estimate to 2010) Lischer, & (Excoffier v3.5 Arlequin used We ). eot ogeil,&Gsul 2017). Gascuel, & Longueville, Lefort, ; a sdt ottetes sbsdo h eut fFish&Cu 21) Anal- (2012). Cruz & Fritsch of results the on based as trees, the root to used was Cercis . nulse aa 00.Teeoe oetmt h iegnetm within time divergence the estimate to Therefore, 2020). data, unpublished , ae ntefsi eodsgetdthat suggested record fossil the on based . 05.TeM nlsswr efre iutnosywt 1000 with simultaneously performed were analyses ML The 2015). , . 04 lt ya,&Hr,21;Ml,Bral,&Hale, & Bartaula, Melo, 2014; Hare, & Nydam, Ilut, 2014; , .chuniana C. . 4 04 n ple h g f24M stesecondary the as Ma 2.4 of age the applied and 2014) , > 0ng/ 30 Fst etn o uppltos( subpopulations for testing , .T opr oeua iest between diversity molecular compare To ). Mse μ http://tree.bio.ed.as.uk/software/figtree/ a used. was L and I .chuniana C. .chuniana C. Nla I nye.Subsequent enzymes. III eepoe maxi- a employed we , He rgntd24Ma 2.4 originated K http://www.atgc- n observed and ) agn from ranging ) Cercis . ; , Posted on Authorea 22 Jun 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159285541.11256769 — This a preprint and has not been peer reviewed. Data may be preliminary. . hrceiainaddsrbto fSNPs of of distribution range No distribution and suitable Characterization region. LGM for May 3.2 subtropical factor April, climate the Chinese in influential Precipitation the most of current. the of and of S1). be map most (Table MH to distribution between occupying revealed geographical vegetation evident was MH, were June The the distribution a and temperate geographical into during 2001). semi-arid the to widely Adams, in contracted in partially changes expanded & data distribution and have (Ray 7) geographical these to 3) (number continu- (number inferred steppe the compared scrub forest a LGM inside or We yielded mainly the woodland located Results during were areas 2b). contrast, distribution respectively). mod- In (Figure the 0.9966, areas (http://intarch.ac.uk/journal/issue11/2/map/download that 2a). and indicated fragmented (Figure 0.9976 data three LIG sample = the of test (AUC and ing distribution power training geographical predictive both ous high on based had performance els model of Evaluation modelling data. niche the Ecological of fit 3.1 best the (AIC). as criterion information chosen RESULTS Akaike models were the 3 demographic AIC on lowest complex based 10,000 were models the arbitrarily with models best-fit with of using model demographic models by likelihood each calculated The 10 composite was under All SFS The given analyses estimation. the S2-S4). FSC2 under likelihood Tables replicate composite S2, 100 (Figure and for ran compared parameters calculated We been optimal 1999). has for Mao, rate simulations & mutation 1.16 genomic Chen of no 1998; rate because (Aldworth, 2020), mutation data, substitution/site/generation unpublished the al., Cercis We et used (Liu scenario. we research each values, to our subjected real was into that estimates source split the as recent ( used migration ( or alternatively size by were applied. (AMIG) population followed also ES effective ancient were or estimated isolation groups (EXP) including NL two expansion (IS), Models model, demographic the only each and (SEC). migration, with In isolation contact one-way models (NIS), or secondary bidirectional demographic isolation or (RMIG), migration without 10 (BOT), of effect are: construction bottleneck models the (MIG), The as on populations based S2). remaining (SFS), (Figure the and and spectrum group (NL), frequency monophyletic group site ( a one easySFS formed as used which considered We were (ES). Mts., effects, another bottleneck Nanling location, Huerta- contact, geographic the secondary distinct in Dupanloup, divergence, a populations of Excoffier, showed The evidence (FSC2; model expansion. provide fastsimocoal2 demographic to and program 2013) Foll, the over & averaged S-DIVA. with Sousa, are with Sanchez, simulations reconstructions detected ancestral coalescent also the the applied were from in events We node derived With vicariance a tree 2016). or at Favre, credibility area Dispersal & ancestral clade trees. an Sun, maximum all of Muellner-Riehl, the frequencies (Matuszak, and the The TreeAnnotator method, trees 2015). and this MCMC He, BEAST & BEAST with Blair, the analysis Harris, Yu, Bayesian on 1997; based of (Ronquist v3.2 was groups RASP analysis seven in implemented used analysis we analysis) vicariance reconstruction, area ancestral For history demographic of ( Inference Lama Tanya 2.6 of 2012) scripts package. al., the R et Illustrator. 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Table S2, model 5, isolation Figure an 3, indicating Table SECEXP, (EXP; model expansion later. is demographic populations FSC2 and western with (SEC) the analysis and contact demographic and NLE2] secondary the first, by + for followed species vicariance [NLE1 model the six fit of separating the best rest rest Across Mts., The the the NLW5. Nanling from the and through of western diverged NLW2 LXS1 in rest and populations and between NLW1 located eastern the eastern is between the LXS2, V4 is and the events, and V6 between WYS2/LXS2 Mts. WYS2 whereas is Nanling between NLW5], between the through V5 is [NLW1 is in V3 V2 populations. populations the respectively. the populations. and Mts., of the LXS1 Luoxiao of including southern rest populations, and the the and (Figure Mts. analysis WYS1 S-DIVA Wuyi the between western from is inferred were V1 regions geographical 1). the among (V1-V6) events vicariance Six differentiation history genetic demographic significant of indicated Inference NLW5]. 2). 3.5 data through (Table [NLW2 GBS variation and total the NLW1, the on of NLE2], based 96.28% + as ( [NLE1 variance populations LXS2, molecular among LXS1, the the WYS2, the with of of WYS1, together in groups Analysis results populations seven analyses: phylogenetic circumscribed the further and ultimately from PCA, we for separate FastStructure, populations, distinctly the of were Considering locations these geographical S1). and (Figure together Mts. LXS2 Nanling and LXS1 clustered analysis when for clustering groups optimal remaining when the the ( populations from east remaining separated the the and in from populations average, other separated the WYS1 On of those to (0.19). comparable LXS2 highest in The detected 1). Table (0.31; highest of The former spanning differentiation 4). estimated the and (Figure Ma, respectively, diversity 0.51 periods ones, Genetic to Interglacial Population eastern 0.34 3.4 the period. from the and Interglacial calculated 0.29 Glacial Dagu-Lushan in were to the Dagu-Lushan LXS1-2) Ma in 0.20 the and latter 0.37 from both (WYS1-2 the to range east and occurring the 0.35 period Mts. in as Glacial from Nanling diversifications estimated Lushan in and were the diversifications which during populations of Population is 0.58 western which both 4). the Ma, Pleistocene, ca. (Figure 0.51 in Middle period populations Nanling ca. NLW1-NLW5 Ma in Glacial the populations western China Dagu Within western the the in rest 4). Mts. from in the (Figure period Nanling diverged debate from glacial NLW1 under the populations while third still in NLE1/NLE2 Ma, are the populations ranges eastern within the time the arose interglacial and was Mts., and times LXS2 LSX1 glacial and divergence between exact WYS2 Both divergence the between the although occurring Ma. the whereas divergence from 0.68 The Ma, diverging 0.72 period. WYS2/LXS2 ca. Glacial by ca. Poyang followed the as populations, of during estimated time remaining Ma The the 0.85 Mts. from ca. Nanling divergent NLE1 western rest and first the NLE2 was WYS2, in with others WYS1 populations the group, 4). the from monophyletic separate by a clade formed followed a of forming branch, Mts. origin first-diverging Nanling Mts. Nanling the the eastern in at the (Figure populations in values positioned The bootstrap be LXS1. high to and mostly revealed LXS2 with was population WYS1 each in for 3). monophyly deposited yielded been analysis have phylogenetic mean times The data The divergence The and 85.00%. S5). relationships (Table [?] Phylogenetic criteria Q30 3.3 extraction and SNP 92.23% for the XXX). SNPs [?] (DOI: with 61,748 Q20 Figshare detected agreed We with SNPs quality 37.84%. 10,761,958 32,890 high from was which identified content was were C tags data High-quality + sequence reads. G The clean GB 200 reads. and reads GB-PE raw GB 200.8 produced GBS .chuniana C. He a eetdi L2(.8 olwdb L5(.5;telws a eetdi LXS2 in detected was lowest the (0.35); NLW5 by followed (0.38) NLW2 in detected was Fst a siae s23 9%HD=19–.4 adrn h al litcn (Figure Pleistocene early the during Ma 1.97–2.74) = HPD (95% 2.36 as estimated was 0.99, = .chuniana C. 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Data may be preliminary. ouainsz,w ovre h iegnetm ewe LadE notenme fgnrto times, generation of number the into ES and NL between time divergence are the T (ES) converted regions we on eastern size, Based and population (NL) as value. calculated Mts. AIC population was size Nanling lowest effective population the into the of diversity on nucleotide sizes and based of population 57,495 confirmed effective estimates current genome-wide was The the Ma) converted sizes. (2.74 we generations rate, 548,000 mutation the of scale time The uxa t. ihanrhsuhoinain r sue osrea egahcbrirpriual for central particularly the barrier Notably, geographic 2013). a as al., serve et and 3). to Yan isolation 1, assumed geographical 1984; (Figures are cause (Liu, orientation, other to to China north-south each reported attributed southern a been from was in with has populations (V1) Mts., species Pleistocene the WYS2 Luoxiao many early geographic separate and the for to during and WYS1 divergence barrier between Mts. flow genetic Wuyi geographical isolation poorly gene of study, as as rising limited our acting considered The In to are Mts. contributing Wuyi 2018). sometimes thus the al., ranges et Mountain dispersal, 2017). (Oyama long-distance Ree, 2). isolation geographic & facilitating (Table Xing ( when flow for divergence 2018; gene conducive especially population Filatov, high inter-population occur & significantly low Hughes with Hughes, to variance 2017; indicates Contreras-Ortiz, thought molecular suggesting Hughes, Nevado, the 2018; is 3), 2017; of Reeder, This Analysis 1, & Kadereit, Higham, (Figure 2015; Hollingsworth, diversification. Atchison, regions (Harrington, & role local geographical dominant with via a plays aligned mostly isolation closely evolved and they monophyletic that are os- climatic populations Pleistocene All with ranges associated western mountain divergence and and eastern population cillations the ranges and between mountain isolation variation Geographical the and genetic (NLE1 4.2 of of physiography Mts. pattern short- Nanling and geographic the and orientation the of of to the south populations populations contributes Therefore, the to have in to north NLW5). seen diversification, appear through from is population NLW1 as more allowing retreat divergence and promoting Mts., younger gradual thus NLE2, and Nanling facilitate periods, size the interglacial can population of and larger species, glacial orientation populations during north-south plant eastern migrations the the of distance in Conversely, seen shifts is cooler LXS2). elevational as population of divergence and various increasing older times LXS1 an for in to disadvantageous WYS2, contributed migrations is have (WYS1, southward may which and blocking expansion, diversification, barriers and population geographical size orientations postglacial present different northward the to or be thought climate to are likely al is Mts. west et and China (Chen east ranges the between mountain diversification of of level genetic and of time al divergence (Chen, pattern et distributions general Gong wide with the 2018; species al with Sun, plant The agrees other & in pattern Zhou, were China 4). This western and Deng, group and (Figure monophyly eastern populations. sizes a between recent eastern formed observed population more divergence the Mts. where larger Nanling from Mts., the with distinctly Nanling of Ma) the populations separated that of within (0.37-020 revealed than range also occurred sizes analysis geographic diversification phylogenetic population the population smaller of with rapid and portion and Ma) eastern (0.51-0.34 the older in be divergence populations western population and The eastern between divergence Genetic 4.1 S3). Table 5, Figure DISCUSSION 1980; 3, 4 (Table Wu, 2). constant & (Figure less ( present or Pu, smaller divergence. at more much ES (Duan, than remained be and China sizes higher to rate population 5 in estimated migration ES was period ca. The the NL was Interglacial contrast, of and In Lushan-Tali size increased present. the population temperature effective within when ancestral is 2004), The Jackson, time & This Liu, Zhu, Ma. 0.10 = DIV . 08 i,Ga,F,&Cms 09.Oemi atrcnrbtn otedffrne fpopulation of differences the to contributing factor main One 2009). Comes, & Fu, Guan, Qiu, 2018; , 1,7 eeain g,ie,aot16M.Scnaycnat(E)wsetmtda ca. at estimated was (SEC) contact Secondary Ma. 1.6 about i.e., ago, generations 319,472 = Ne ES .chuniana C. 495 epciey rmtecreteetv ouainsz,teacsrleffective ancestral the size, population effective current the From respectively. 14,955, = M NL-ES 21)wsmc ihrthan higher much was (2.14) . . Ne 08.Tesuhetnrhatoinaino h uiMs n East and Mts. Wuyi the of orientation southwest-northeast The 2018). , ANC . 08 a i,Co,&Km 08 omn ta. 08 uet Lu 2018; al., et Hohmann 2018; Kim, & Choi, Kim, Ha, 2008; , 5,5 Tbe3 iue5 al 3.Uigteacsrleffective ancestral the Using S3). Table 5, Figure 3, (Table 755,955 = 7 M ES-NL 03) ihmgainocrigatrNL after occurring migration with (0.33), .chuniana C. Fst N e-pre-exp 0.99, = 6)ta at than 866) = setmtdto estimated is P .0 also 0.00) = Ne T NL SEC = Posted on Authorea 22 Jun 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159285541.11256769 — This a preprint and has not been peer reviewed. Data may be preliminary. hntepeet(un u u 90 h ta. 04.Ti ugssta h eodr otc may contact secondary the that suggests This 2004). higher al., even still et be it Zhu to Ma), estimated 1980; (0.12˜0.14 was Wu, and LIG & increased the Pu, temperature continuous than (Duan, when present a younger China with the is in when than coincides period period 1980), interglacial Ma) Wu, interglacial the Lushan-Tali (0.10 & within the analysis lies Pu, Although our (Duan, from 2a). Ma) contact (Figure (0.10-0.20 secondary China of of in distribution time geographic period estimated expansion interglacial demographic The Lushan-Tali and contact the 5). secondary by Figure followed other isolation 3, of of migrations (Table model that best-fit west-to-east a to contemporary indicate analyses analogous and FSC2 expansion be demographic not contact, may Secondary refugia conditions glacial 4.3 climatic isolated several the where although ENM, LGM, in cycles. cycles the apparent glacial the glacial also affected Pleistocene during is for Lushan primarily identified populations role and of period were dominant period distribution The Glacial Interglacial geographic diversification. Dagu-Lushan Dagu the population the on The affected whereas northern primarily west, in 4). of and period lowlands (Figure distribution east Glacial to the cycles geographical forests between glacial fir the divergence spruce study, Pleistocene population drive and the our to dry with In 2017; shown The be associated been al., to ranges. have 1988). distribution tended et Pleistocene contracted (Liu, periods (Foster with Middle glacial elevations China estimates the lower The during in to time climate FSC2. shifting period The smaller in populations glacial setting the 2017). generate favor Barker, log-uniform to would & the the which thought Hong, under when cool, Zhang, is range estimation 1980), Kong, Wu, time time BEAST i.e. Bayesian 2017; & wider in of Kong, Pu, older, the results used (Duan, to is the attributed period calibration between FSC2 Glacial Periglacial partially discrepancy secondary Xixiabangma in Sizishan be The the the may model present. geographical of or FSC2 at fit the glacial 2016), and than Quaternary best lower al., shaped known 10 et the and was (Wan earliest genetic of east, temperature Ma the the time 1.6 with the drove overlaps divergence ca. to glaciation which 1980). estimated period those (last) 5), Wu, The and (Figure third & Glacial Ma Middle the Pu, variation. Mts. Kunlun 1.60 primarily the (Duan, genetic Nanling 1980), Wu, uncertain, Interglacial in the of Poyang-Dagu & & is China in patterns Pu, or Xu, glaciations in populations Duan, Zheng, 2011), the between period Ma; Wang, Ma; for divergence (0.5–0.6 glacial (0.5–0.7 & the time period period third Shi, in the Glacial Glacial Zhao, last populations Although Naynayxungla Dagu Ma; the the the 1998), (0.62–0.78 with between in Li, period fall Ma) & coincides may Zhou (0.68 east time 2002; al divergence the The have Shen, et of of to 4). Qu time those thought (Figure 2010; the are and Pleistocene (Knowles, and estimation, isolation habitat Bayesian Mts. geographic suitable on Nanling with changed Based coupled data periodically expansion and cycles 2011). and markers These contraction molecular range cycles. of traditional promoted glacial divergence with the population analyzed by that affected were as suggests which such study of China, Our all subtropical in 2019), range Li, distribution & wide analysis. Li, a with Cheng, species (Shen, directly plant isolation have other historical pauhoi many the Mts. Machilus the in to Nanling contributing from found and factor been distant Luoxiao, barriers major have geographically Wuyi, a geographical and considered the the are including Therefore, of Mts. and ranges colonization V6. Nanling populations mountain long-distance involving the the associated limited vicariance of with the the contrast ridge by in In one formed resulting on thus V5. alone populations, involving has that disadvantageous the remaining as also vicariance located (V4) infer within infer events to we is populations east We which contributing NLW1 (V6) Vicariance orientation, the northwest NLW2˜5, thus north-south and to general 3). orientation. colonization, middle a those the present east-west west-to-east 1, as and for Mts. a well (Figures Nanling as with The V3) (V5) Mts. Mts. corridors eastern and Nanling Nanling and geographical currently western (V2 the the of are between of populations which lack exist Mts., populations western the Luoxiao the or through the between in eastern arisen LXS2 isolation their and geographical of LXS1 the to each apply from to isolated appears This colonization. east-west .chuniana C. ouain Jag u eg 09 ie al et Li 2019; Deng, & Xu, (Jiang, populations Zue al et (Zhu .chuniana C. . 2017), , ooeau chinense Loropetalum ln h onanrne nsuhr hn a eetdb ENM by detected was China southern in ranges mountain the along .chuniana C. 8 Gn tal et (Gong . 09 age al et Yang 2019; , curdi h litcn n a been has and Pleistocene the in occurred .chuniana C. . 06 and 2016) , . nsuhr hn salso is China southern in 09.Smlrpatterns Similar 2019). , iidnrnchinense Liriodendron . , Posted on Authorea 22 Jun 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159285541.11256769 — This a preprint and has not been peer reviewed. Data may be preliminary. kie .(94.Anwlo ttesaitclmdlidentification. model statistical the at look new A 19 Control, (1974). H. Akaike, XXX). (DOI: Figshare REFERENCES in authors deposited All been manuscript. have analyzed the data F. wrote PW. The and and G., results sample W. the statement and K., discussed Accessibility HH. investigation G. L., Data field manuscript. W. WZ. conducted the and work. with L. F. lab agree PW. the WZ. and out L., carried read research. WZ. Z. the data. H. designed and genomic X. the and JG. supervised L., WZ. H. collection. G. for and Yan information. G. Haifei and fossil W. and sampling providing Yi, for with Wang Huiqin help Qi Liu, for Tongjian contributions Prof. Zhang 31470319; Meier, to Author Zhong Joana goes (31470312; Ravinet, and thank (2016A030303048). Mark Huang, China Special Sun, China Yemei analyses. of Zhixia Li, Province, data thank Foundation Bo We Zhou, Science of trips. Juan Natural field Wang, Project National Xiaolan Planning thank Technology the We and by Science supported 31970231), was work This the facilitate across may areas which stepping-stone adjacent ones, ACKNOWLEDGEMENTS closely eastern via the ones than for lower ranges. ecological to Mts. refugia mountain elevations Nanling stable the glacial higher the long-term more the from in diverged and migration of elevations phytogeography former west-to-east higher one the distinct relatively whereas with be at east, Mts., to present the Nanling thought towards The is Mts. condition, latter. Nanling the the than from recently inferred is direction migration in STEBF of history long a has ridges, distinct five of al composed with et are inferred (Fan diversification which was population Mts., China Mts. local Nanling high Nanling southern indicating The the 5), in Figure 3). expansion 3, (Table (Figure demographic size model, population intensified effective same increased monsoon the notably al In summer et Asia S1). (Liu (Table East period precipitation that interglacial the considering in the that especially during thought habitat, precipitation is more al with it et temperature Moreover, Meng increased to time. due warmer then this during occur rce rmrl rmteNnigMs otees.Mn xmlso ln pce nEs saehbta al exhibit et Asia Wang East 1992b; in species 1992a; plant Wang, of 2018; examples as al., Many such et east. route, (Tian migration the and to with pattern Mts. distribution divergence, Nanling similar ES the and from NL primarily relatively after proceed with occurring species migrations the bidirectional M for indicated even analysis dynamics, FSC2 using sampling, Additionally, population generation conducted population next the is the intensive detect research on to more previous based distribution. ways approaches the with narrow genomic effective of population methods, more Most consider are analysis We sequencing study. 2009). data current Huang, and the & with markers compared Lowe, molecular Kang, traditional Gao, Wang, the China, Mts. 2018; subtropical in increase al., Nanling species size et widespread the population in climatic in documented in resulting been Jiang, as from conditions have finally & such cases diversity, buffer ecological (Tian Similar population current a preserve stable expansion. as to demographic as long-term period and thought glacial served are plus last cycles physiography has the glacial during complex and across precipitation Therefore, periods annual same glacial the 2016). during almost had mountains It al these change. et to (Shen conditions shifts ecological longitudinal or elevational NL-ES uyoybscavaleriei Eurycorymbus 21)hge than higher (2.14) . 6,7673 doi:10.1109/TAC.1974.1100705 716–723. (6), 08 age l,19;Wn ta. 07 age l,21) hspoiigmr suitable more providing thus 2012), al., et Wang 2007; al., et Wang 1999; al., et Wang 2018; , . M 08 ue al et Xu 2018; , ES-NL , ooeau chinense Loropetalum .chuniana C. 03)(al ,Fgr ) h irtosin migrations The 5). Figure 3, (Table (0.33) . . 09 h tal et Zhu 2019; , 07.Isvgtto scaatrzdb ihyvaried highly by characterized is vegetation Its 2017). , saatdt ei niomnsadms nune by influenced most and environments mesic to adapted is . 9 05.Teqeto rssa owycontemporary why to as arises question The 2015). , ersim hemsleyanum Tetrastigma and .chuniana C. oeo chionantha Eomecon . 07,wihcnesrltvl stable relatively confers which 2017), , ouain of Populations . EETascin nAutomatic on Transactions IEEE and Gn ta. 06 Tian 2016; al., et (Gong .chuniana C. oeo chionantha Eomecon .chuniana C. . emto seem 2018; , are Posted on Authorea 22 Jun 2020 — The copyright holder is the author/funder. All rights reserved. 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Wuyi Eastern range Mountain at n lntr cec Letters Science Planetary and Earth oeua hlgntc n vlto,87 Evolution, and Phylogenetics Molecular 17 Mgoica)rvae yclrpatDA east-west DNA: chloroplast by revealed (Magnoliaceae) populations. 426.00 26.78 14 9 27.94 27.82 9 8 N caGorpiaSinica Geographica Acta utrayInternational Quaternary er,7 Peerj, iifrais(xod England), (Oxford, Bioinformatics aiueLongitude Latitude ahlspauhoi Machilus 65.doi:10.7717/peerj.6355 e6355. , º º º º 114.15 N 113.90 N 117.77 N 119.85 N hns idvriy,9 Biodiversity., Chinese , 66 , 7,867-884. (7), 221 nSuhChina. South in , º º º º 6-2 0.31/0.19 369-722 0.32/0.30 E 264-509 E 0.34/0.28 460-712 0.32/0.31 E 264-727 E 97 14,55–69. (1-4), Quaternary Ho 93–101. , 46–49. , and ) ag (m) range Elevation He/Ho Posted on Authorea 22 Jun 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159285541.11256769 — This a preprint and has not been peer reviewed. Data may be preliminary. mn rus636.8 706896.28% 1.38% 38.49802 3458.143 2.34% 0.53215** 37.06608 111 3367.488 53.747 0.89979** 101 36.908 6 Indices Fixation Total 4 populations Within groups within populations Among groups Among variation of Source 2 TABLE Mao’ershan NLW5 Nanshan NLW4 11 Dupoxiang, NLW3 10 Tianpingshan, 9 NLW2 Yindianshan 8 NLW1 Dadongshan 7 NLE2 Mangshan 6 NLE1 Location Population 5 No. nlsso oeua aine(MV)rslsfrglobal for results (AMOVA) Variance Molecular of Analysis , Reserve, Nature National , Yard, Log Hunan hua, Huai- Guangxi Guilin, Reserve, Nature ingshan Huap- Guangxi Guilin, Reserve, Nature Guangdong , Reserve, Nature Hunan zhou, Chen- Reserve, Nature Fsc d.f. 0.6284** : Mts. Nanling Western Mts. Nanling Western Mts. Nanling Eastern Mts. Nanling Eastern range Mountain Mts. Nanling Western Mts. Nanling Western Mts. Nanling Western 18 Fst variation of Percentage components Variance squares of Sum 0.9862** : 25.89 26.18 7 26.07 14 13 25.61 5 24.91 11 24.92 10 24.98 12 N Fct Fst aiueLongitude Latitude 0.9628 : ttsisof statistics º º º º º º º 110.38 N 110.19 N 109.47 N 109.90 N 110.96 N 112.72 N 112.89 N .chuniana C. º º º º º º º 0-5 0.35/0.25 700-852 0.32/0.30 E 611-703 0.32/0.30 E 612-657 E 0.38/0.33 852 0.33/0.34 E 700 0.32/0.32 E 774 0.32/0.21 E 716 E . ag (m) range Elevation He/Ho Posted on Authorea 22 Jun 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159285541.11256769 — This a preprint and has not been peer reviewed. Data may be preliminary. ttebto fe h pi ewe h aln t.(L n h atr onan E) h migration The (ES). box mountains grey eastern shown the top and are The (NL) sizes Mts. ( ES. population right. Nanling rates current the the corresponding The between on and split size. shown directions the population after periods bottom effective interglacial the with or at populations glacial ancestral ( our corresponding the divergence represents the in of with investigated Time lines, models (Ma). dashed demographic time shows fit line best braces, bar. vertical the the the by above of indicated latter representation are study. the frame. periods Schematic line while interglacial dotted bar 5 and the the FIGURE glacial within below Various distinguished is bottom. are which [?] the of Mts. (P at former Nanling shown probabilities the the is high in bar with Populations scale (V1–V6) nodes. events Time for Vicariance corresponding shown areas. are in ancestral 95% 0.70), shown and indicate are regions bars FastStructure geographical blue in different light clusters The different estimation. with to time colours assigned Bayesian Individuals the struc- intervals. in population reconstruction. simulations confidence estimates, (MCMC) area time Carlo divergence ancestral Monte for chain and tree Bayesian clustering the tural of Chronogram 4 FIGURE outgroup. the as used factors influential of method. most tree The Phylogenetic study. 3 present FIGURE simulated the The for S1. (c) localities Table collected representing steppe- samples in steppe. dots and black dry 4, listed with 8, data are scrub; thorn range, and specimen or distribution tropical steppe; herbarium potential woodland forest current on 1, 7, temperate The based LGM: desert; (d) semi-arid temperate MH. the 3, at 6, at range desert; woodland; distribution alpine types boreal and vegetation open polar different 5, 2, tundra; contact represent woodland; secondary 1-8 scrub during Numbers and temperature scrub LGM. The at the ranges. map in mountain suitability indicated relevant vegetation are habitat to period the point each represents arrows in ( LIG; red variations (a) where temperature respectively, The boxes. and grey, gray 44.93%. Suitable and than current. red higher and probability) in (MH), (occurrence Holocene indicated Middle are the habitats (LGM), Maximum unsuitable Glacial Last the (LIG), temperature. Interglacial in variation corresponding and (ENM) **: ae ihtinls iaineeet eetdi -IAaesonwt lelns h e tbottom at key for The 1. areas Table indi- lines. suitable in are blue given Potentially east with is the 2 shown information in FIGURE are accession populations S-DIVA Additional circles; in ranges. black detected elevational with events indicates indicated Vicariance right are triangles. Mts. of with Nanling locations cated the sampling in study. the sampled this showing Populations in China, southeastern used of ulations Map 1 FIGURE LEGENDS FIGURE collections; all among differences T SEC P sidctd b oetal utbeaespoetdi oprsnwt ae fGIS-based of layer a with comparison in projected areas suitable Potentially (b) indicated. is ) < oe ae n h aaeescrepn otoei alsS n 3 epciey h left The respectively. S3, and S2 Tables in those to correspond parameters the and names Model 0.001; otta ecnae ( percentages Bootstrap K ,3ad7 ih7a h pia au.I h -IAaayi,clu eed indicate legends colour analysis, S-DIVA the In value. optimal the as 7 with 7, and 3 2, = Fct ieec mn groups; among difference , h onanrne novdi h td r hw noag shadows. orange in shown are study the in involved ranges mountain The d.f. > M 0 nteM reaeidctdaoetebranches. the above indicated are tree ML the in 50) .chuniana C. ere ffreedom. of degrees , ES nidvda irnsprgnrto r hw ewe Land NL between shown are generation per migrants individual in ) .chuniana C. Fsc T ouain ae nmxmmlklho (ML) maximum-likelihood on based populations 19 DIV ieecsaogcletoswti groups; within collections among differences , n eodr otc ( contact secondary and ) ordffrn eid eeapid .. h Last the i.e., applied, were periods different Four rnhlntswr rnfre i Markov via transformed were lengths Branch rdce yeooia ih modeling niche ecological by predicted T SEC .chuniana C. ecschingii Cercis r niae as indicated are ) pop- Fst was , Posted on Authorea 22 Jun 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159285541.11256769 — This a preprint and has not been peer reviewed. Data may be preliminary. 20 Posted on Authorea 22 Jun 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159285541.11256769 — This a preprint and has not been peer reviewed. Data may be preliminary. 21 Posted on Authorea 22 Jun 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159285541.11256769 — This a preprint and has not been peer reviewed. Data may be preliminary. 22