Evidence for Hybridization Between Two Sympatric Termite

Posted on Authorea 26 May 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159050496.61216640 — This a preprint and has not been peer reviewed. Data may be preliminary. net n se Abt ta.21;Cplnqe l 05 ens 06.Icmlt aresbetween species, via barriers termite species Incomplete between related 2016). alleles closely species. 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Similar in investigate initial-stage long-differentiated However, to of nature. sympatric, development sequencing two impact. the DNA or between ecological mitochondrial invasion trogression new with species genotyping a by microsatellite caused with combined zones associated we hybrid be on focus to mostly likely bridization is species between Hybridization Abstract 2020 26, May 2 1 Wu nature Jia in species termite sympatric hybridization two for between evidence differentiation: genetic Asymmetric uzogArclua nvriyCleeo ln cec n Technology and Science Plant of College University Agricultural Huazhong available not Affiliation tal. et 03 eioisGnm 02.Hwvr eetsuishv ugse hthbiiaini animals in hybridization that suggested have studies recent However, 2012). Genome Heliconius 2013; 1 tal. et tal. et iyn Huang Qiuying , 03 Meier 2013; 03 umn&Pml 04 Song 2014; Pamilo Kulmuni& 2013; 03 asu&Reeeg21;Zhang 2016; Rieseberg Payseur& 2013; tal. et tal. et 1996). 2 07.I diint t eei osqecs yrdzto a nunethe influence can hybridization consequences, genetic its to addition In 2017). unXu Huan , eiuiemsflaviceps Reticulitermes 1 n asnAli Hassan and , tal. et 1 tal. et 01.Apoiaey1-0 faia n plant and animal of 10-30% Approximately 2011). 06.Ti iegneicue eta diver- neutral includes divergence This 2016). siaand Oshima 1 eiuiem chinensis Reticuliterme Snyder, Posted on Authorea 26 May 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159050496.61216640 — This a preprint and has not been peer reviewed. Data may be preliminary. eol sdsqecsfrwihtetodrcincrmtgaswr elmthd eune were Sequences matched. well was were splicing chromatograms After two-direction (http://www.dnastar.com/t-allproducts.aspx). (Applied v7.1 the v1.0 Lasergene which Scanner DNASTAR those for Peak using on sequences and spliced based Biosystems) used results. were the (Applied only assay of We v4.0 PCR analysis GeneMapper the each for (2020). for and used Ra141 al. conditions were Ra128, et Biosystems) The Ra95, Wu Ra79, S1). Ra50, by Table Rs76, materials, described Rs78, Rs03, supplementary primers: (see microsatellite Ra144 of pairs ten using performed the using 4 performed was 5’-GTTTAAGAGACCATTACTTA-3’ at PCR (mi- (Simon gene, markers B-tLys, stored COII nuclear 5’-CAGATAAGTGCATTGGATTT-3’, and was mitochondrial and A-tLeu, the DNA (COII) For primers sample. gene Co., extracted individual mitochondrial Biotech each The a Gen from for manufacturer. 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R. ° n 15 and o eei analysis. genetic for C .flaviceps R. oee,wehrhbiiainadaatv nrgeso cu in occur introgression adaptive and hybridization whether However, tal. et .chinensis R. .flaviceps R. 06.I em fmrhlgclcaatrsis h colour the characteristics, morphological of terms In 2016). n 15 and 2 .chinensis R. ° ooisfo hnd,10 Chengdu, from colonies (Miura C .chinensis R. and tal. et .chinensis R. 04.Te r ossetyrecognized consistently are They 2014). tal. et sbak h coinadsamn of swarming and eclosion The black. is .flaviceps R. ooisfo ajn clne were (colonies Nanjing from colonies .flaviceps R. 00.FrncerDA C was PCR DNA, nuclear For 2000). hr tbehbtt,including habitats, stable share and .flaviceps R. and .flaviceps R. .chinensis R. .chinensis R. .flaviceps R. n 15 and eclosion , ° until C n to and pop- R. Posted on Authorea 26 May 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159050496.61216640 — This a preprint and has not been peer reviewed. Data may be preliminary. lt agdfo .5 o088 ihama f074 in 0.734, of mean a with 0.848, to 0.657 from for ranged for diversity plots those nucleotide to and similar diversity were haplotype 1) of values in to index found were The similar diversity genetic haplotypes overall of the showing levels within Similar samples diversity clades comprised genetic major clade nuclear two and first showed Mitochondrial haplotypes The method 18 ML and 2). the Wuhan parsimony-informative in via the (Fig. 78 flaviceps, haplotypes Among constructed species 12 and tree sequences. Nanjing, sampled gene sites phylogenetic in COII collected The variable found mitochondrial singleton were the haplotypes Chengdu. haplotypes on (14 unique in based nine 31 sites individuals plots, samples, variable aligned all sample 67 In different 92 the individuals. the from 67 derived on in sites) resolved based was identified COII of were to bp belonging 661 colonies markers, 38 mitochondrial were There 2). to (Fig. belonging analyses gene COII mitochondrial of colonies sympatric Sixty-seven haplotypes and to to identification identification (LnP(D)) Species species K with each combined used values for We DK data 10,000. used Results of the we burn-in of and a K. independent probability clusters, and optimal 10 of posterior repetitions the number with the MCMC obtain likely 10,000 frequencies, of most with correlated logarithm the performed identify with estimated 10) maximum model to 1 mean admixture of (from the the K populations ran each the We for runs of 2.3. clustering STRUCTURE genetic using species-related was there 2.9.3.2. chinensis v. whether data. FSTAT frequency and determine v5 genetic To DnaSP using termite 6.5, measured the v. indirectly of GenAIEx was levels. differentiation using flow (SSR) calculated genetic gene Gene was nuclear the populations, and low among calculated (COII) variance but we gene the genetic genes Thus, nuclear mitochondrial whereas of the differentiation genes. at species, genetic species (Greiner cytoplasmic high parental recombinant be of different not will differentiation There are the genetic and material. of cytoplasmic mother and product their nuclear by between combined contributed a are Hagstrom are offspring species, carry 2015; the between offspring not occurs of does the hybridization genes gamete When of cytoplasmic male zygote. genes a a whereas nuclear of material, formation the the cytoplasmic to and contrubuting nuclear material both cytoplasmic contributes gamete female flow A gene estimated haplotype and including differentiation v5, analysis. Genetic T-test DnaSP via using species data the mitochondrial between ( ) on diversity based nucleotide diversity and diversity genetic ( investigated locus also per strategies We alleles reproductive effective ( of and heterozygosity of diversity structure expected number genetic genetic mean nuclear the parameters: the evaluating following characterized for We measure populations. important model of an substitution is best diversity estimated retained. 7. Genetic the were maxi- v. length and The se- MEGA same replicates, 2009). using The the Rozas bootstrap obtained of comparison. (Librado& 100 were sequences v5 for with (GTRGAMMA) all DnaSP tree, MEGA7 that using phylogenetic so data into likelihood deleted mitochondrial imported mum were the ends from and estimated both format were at Fasta Haplotypes neat not in were saved that were quences sequences all completed, n h eodcaecmrsdsmlssoighpoye iia to similar haplotypes showing samples comprised clade second the and n hi yrdppltos eepoe aeinapoc o nern ouainstructure population inferring for approach Bayesian a employed we populations, hybrid their and .chinensis R. tal. et 04.Tu,itrpce yrdzto a edt smercgntcdifferentiation genetic asymmetric to lead may hybridization interspecies Thus, 2014). H ohseiswr itiue crs hnd,NnigadWhn o the For Wuhan. and Nanjing Chengdu, accross distributed were species Both . .chinensis R. e .Teeaaye eepromduigGnIxv . n EEO .1.2. v. GENEPOP and 6.5 v. GenAIEx using performed were analyses These ). Reticulitermes h and ( h π 0.854, = .W etdtedffrne ngntcdvriy( diversity genetic in differences the tested We ). emtswr dnie sn eiso opooia and morphological of series a using identified were termites π 3 .flaviceps R. .3,Tbe1.Hpoyedvriy( diversity Haplotype 1). Table 0.034, = .flaviceps R. and R N .flaviceps R. e . .chinensis R. n rm04t .7,wt enof mean a with 0.978, to 0.4 from and ,osre eeoyoiy( heterozygosity observed ), flaviceps F ST tnadzdmaueo the of measure standardized a , .flaviceps R. and ( .chinensis. R. h ntemDAsequences. mtDNA the in 081, = .chinensis R. π .flvcp,R. flaviceps, R. n 9colonies 29 and N .3,Table 0.036, = h e ndifferent in ) , H sn the using H e o and and ) tal. et H R. o Posted on Authorea 26 May 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159050496.61216640 — This a preprint and has not been peer reviewed. Data may be preliminary. ut ugse htitrpce yrdzto n eeecag curdi h w emt pce in species termite two the in occurred exchange gene and Additionally, genes, hybridization mitochondrial species. interspecies the nature. of between that divergence genes nuclear genetic suggested of high sults divergence and genetic low flow and identified gene results flow the low gene Our high was was There 2000). there while Watanabe genes. (Setoguchi& nuclear hybridization and in- interspecific chondrial maternally of are evidence that genes provide showed cytoplasmic to whereas considered recombination, repro- (Abbott is gene sexual evolution Greiner nuclear of adaptive 2001; mode in (Birky for same herited only basis the result a through provide can alleles may reproduction of which combinations species, new in with duction variation in indicating results plots, Hybridization all for of 2 populations the = genotypic contrast, K Discussion In species-specific was two observed. K into were three appropriate divided introgressions appropriately chinensis in most gene were R. the introgression Wuhan few identified, and nuclear a Nanjing were although of both clusters, species evidence of the colonies showed As the also that 4). STRUCTURE using (Fig. analysis plots test assignment The approximately words, other clustering In Genetic basis. genetic a flaviceps. on R. colonies the in of 6% individuals of possessing of individuals colonies female mean of 29 a 10% in with Similarly, (Chengdu), 3). 14.9% (Table 10% a of lotypes, mean a with (Chengdu), area sympatric chinensis the in constrained either from between introgression hybridization Nuclear interspecific was there that chinensis. indicated material ( species cytoplasmic the and between flow ( gene species low the and divergence genetic ( high < genes species showed two nuclear analysis the and gene between mitochondrial COII differentiation their genetic between that the found differentiation We results, genetic 2). (Table genetic F-statistics nuclear using and calculated was mitochondrial the on Based flow gene and in differentiation 0.72, Genetic in of 0.69, means with of 0.88, mean to a 0.20 with in from 0.50, ranged of heterozygosity mean expected a with 0.80, to of populations 3B; Fig. : in 4.15, of lee agdfo .5t .4 ihama f43,in 4.39, of mean a with 8.54, to 1.25 from ranged alleles o ula eei iest,teewsn infiatdffrnei h ubro ffcieallsbetween alleles effective of number the in respectively. difference 0.013, of significant mean no a of was with populations there the 0.028, diversity, to genetic 0.002 nuclear from For and 0.014, of mean a with in 0.674, .5 al ) u h ula ee SR)soe o eei iegneadhg eeflwbetween flow gene high and divergence genetic low showed (SSRs) genes nuclear the but 2), Table 0.15, .chinensis R. seicncerhpoyewsosre tpootosrnigfo .%(ua)t 23.7% to (Wuhan) 1.5% from ranging proportions at observed was haplotype nuclear -specific .flaviceps R. .chinensis R. t F .flaviceps R. .flvcp/.chinensis flaviceps/R. R. nCeguwr uhmr ie n hwdltl vdneo pce differentiation. species of evidence little showed and mixed more much were Chengdu in =1.17, .flaviceps R. .chinensis R. st < .flaviceps R. 0.12, p uloiedvriy( diversity Nucleotide . .5 repce eeoyoiy( heterozygosity expected or 0.25) = seicncerhpoyewsfuda rprin agn rm22 Whn to (Wuhan) 2.2% from ranging proportions at found was haplotype nuclear -specific .chinensis R. iial,teewr osgicn ieecsi h bevdhtrzgst ( heterozygosity observed the in differences significant no were there Similarly, . N and and m ae ihmlsof males with mated .flaviceps R. tal. et .chinensis R. > and .chinensis. R. .flaviceps R. .3 al ) h smercgntcdffrnito ewe h nuclear the between differentiation genetic asymmetric The 2). Table 2.63, .flaviceps R. . 05.Tu,aymti eeflwbtenctpamcadncerloci nuclear and cytoplasmic between flow gene asymmetric Thus, 2015). .chinensis R. . mn 8clne possessing colonies 38 Among ooispossessed colonies ae ihmlsof males with mated xiie smercgntcdffrnito ewe hi mito- their between differentiation genetic asymmetric exhibited n rm02 o08,wt eno .4 in 0.54, of mean a with 0.85, to 0.22 from and π h bevdhtrzgst tdffrn oirne rm0.22 from ranged loci different at heterozygosity observed The in ) into ( N .flaviceps R. e .chinensis R. i.3A; Fig. : 4 .flaviceps R. .ciessR flaviceps chinensis/R. R. .flaviceps R. H e .chinensis, R. i.3C Fig. : t and 0.44, = or .chinensis R. .chinensis R. R.flaviceps and n rm10 o1.1 ihamean a with 10.01, to 1.06 from and t t ; 10.72, = p .flaviceps R. .chinensis R. .6.Tenme feffective of number The 0.66). = n prxmtl %o female of 6% approximately and 0.78, = tal. et .chinensis R. agdfo .0 o0.02, to 0.008 from ranged tN altps an haplotypes, mtDNA into p ula ee.Teere- These genes. nuclear 03.Hwvr sexual However, 2013). < p n rm00 o0.90, to 0.05 from and ipae asymmetric displayed .4 ewe the between 0.44) = .flaviceps R. .0) h mtDNA The 0.001). .flaviceps R. .chinensis. R. F .flaviceps R. st tN hap- mtDNA > 0.62, a un- was and N The and H R. R. m o Posted on Authorea 26 May 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159050496.61216640 — This a preprint and has not been peer reviewed. Data may be preliminary. h bv-ecie eutosi eei ieecsbtenseisnrseito r bevdi the in observed are speciation nor species between differences genetic termite in the termite (Bernal reductions in effect above-described existed homogenizing introgression the gene genetic Additionally, and a (Abbott hybridization cause 2016). speciation (Pennisi even accelerates species or and of bifurcation taxa on evolution binary between based the tree differences a al. describe phylogenetic reduce through truly existing diffusion the can cannot The described Therefore, to hybridization structure be hybridization. evolution. bifurcation due via reticulate cannot binary namely, re-fused speciation network, a process often or a evolutionary by introgression are but adaptive this species structure in of differentiated result relationship speciation, can exchange phylogenetic which and gene changes, also evolution interspecific environmental are of that or exchange suggested process differentiation. gene results species the of of These During patterns time the 2016). 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(Henrich& not two ago shows the years between species million tapeworm flow 20-25 related gene the is in there reported between that time and species differentiation barriers between the (Bourguignon flow prediction, years gene million that preventing 2-3 with offspring, Inconsistent hybrid of 2009). fertility and viability species (Anderson long-diverged the between affects hybridization However, negatively species 2007). original often Feder the (Dambroski& differentiation between of of stage isolation time initial reproductive short in the prezygotic in to races remain led host they shift because and host species a young example, in For termites occur differentiation. in barriers diversity reproductive genetic incomplete high general, nature hybridization. of In in and maintenance introgression the species that adaptive the speculation to between the exchange related confirmed genetic is further regarding we study, results this the in with Combined still conditions. that is laboratory noted diversity previously We genetic inbreeding. high al to maintained fact, respect be can with In diversity termites chinensis genetic of why inbreeding. explain populations may inevitable colony natural species in closed to isolated One incompletely lead between may introgression 2009). and Husseneder (Huang which Vargo& populations by kin, natural 2005; established in of Vargo are maintained (DeHeer& colonies composed the many colony on be termites, natal depending social can assimilation same For genetic the 2004). facilitate from (Seehausen or impacts founders diversity populations various of genetic cause construction to may contribute and species can size isolated and It incompletely diversity. dealates mate between genetic then between material on (Mizumoto and hybridization genetic colony nest of for virgin introgression stable mechanism Adaptive heterospecific a a possible workers. co-establish form a from and to developing provides fuse partner reproductives which workers secondary homosexual into foraging dealates, a develop that heterosexual find possible can may with is workers they it termites, Hence, time; subterranean 2016). long some a Su in for 1981; (Matsuura Additionally, live Howard factors (Haverty& other other. and/or reproductives predators usually each secondary ratio, is encounter sex colony they the initial to when the due Second, partners conspecific repro- species. with al. (Aldrich& prezygotic Thorne termite mate indicated 2008; to species sibling fail Hartfelder the some sibling may (Korb& Thus, in in monogamy absent 2011). similar under largely established Baer is be (Hartke& may pheromone genitalia barrier sex-pairing sclerotized ductive swarm- the external and and possess distribution the females common, of overlap is Capblancq the 2009; species termites, Kambhampati In related hybridization? of causing factors ing possible the are What 09.A h aetm,w on htatfiilhbi ooiscnpouelvn ffpigunder offspring living produce can colonies hybrid artificial that found we time, same the At 2019). . 02.I hscs,termtn ihhtrseicprnr n etbidn aoripoe fitness improved favour building nest and partners heterospecific with mating their case, this In 2002). 07 cirnek21) lentvl,hbiiainrslsi h nrgeso fgntcmaterial genetic of introgression the in results hybridization Alternatively, 2011). Schierenbeck 2017; Reticulitermes oss iia eaiusadlc oseicpeeecsi h rcs frpouto (Wu reproduction of process the in preferences conspecific lack and behaviours similar possess tal. et 00 Brideau 2010; hgltspomonella Rhagoletis fti eetecs,teewudceryb togslcinaanthybridization against selection strong be clearly would there case, the were this If . tal. et tal. et tal. et tal. et 06.Orrslsidctdta hyehbticmlt reproductive incomplete exhibit they that indicated results Our 2016). 06 rce&Bresen21;Ftptik20;Mye Nakazato Moyle& 2008; Fitzpatrick 2013; Bordenstein Brucker& 2006; 05 ate oegu 01.Mr motnl,nihrmlsnor males neither importantly, More 2011). Rosengaus Hartke& 2015; u h otyoi erdciebrir eeivlddet the to due invalid were barriers reproductive postzygotic the but , citcpau solidus Schistocephalus 03 Bendesky 2013; tal. et 03 ag&Hseee 09.Fruaey hybridization Fortunately, 2009). Husseneder Vargo& 2013; 5 tal. et tal. et Reticulitermes 04,addaae htfi omt a also can mate to fail that dealates and 2014), 07 asu&Reeeg21) However, 2016). Rieseberg Payseur& 2017; tal. et and .pungitii S. 99.Mn ooaosindividuals monogamous Many 1999). (Dedeine .flaviceps R. ept hi differentiation their despite tal. et and .flaviceps R. 06.Neither 2016). .chinensis R. and tal. et R. et et et is Posted on Authorea 26 May 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159050496.61216640 — This a preprint and has not been peer reviewed. Data may be preliminary. hntriia)ifre rmaaye fmtcodiladncerloci. nuclear and mitochondrial of analyses S from Guyot inferred development. Rhinotermitidae) S, diapause Dupont of F, stages different Dedeine on adaptation in history variation life diapause 2101-2112. multifaceted latitude-related , for and test plant in a Host speciation : (2007) promotes JL Feder Hybridization HR, (2015) Dambroski J Mavarez D, Rioux butterflies L, lethality Despres hybrid T, cause bacteria Capblancq gut speciation: Nasonia. of genus basis hologenomic the The in (2013) SR Bordenstein RM, Brucker J Wang in HA, lethality Flores NJ, Brideau termites and 20160179. the mechanisms, , of laws, distribution chloroplasts: modern J the and Sobotnik mitochondria N, Lo in T, Bourguignon genes of inheritance The (2001) models. (genus: Jr. evolutionary fishes the CW, reef shaped Birky selection coral and of Introgression sister-species (2017) LA sympatric Rocha of WB, history Simison MR, Gaither MA, Bernal subspecies JM mice. two Lassance amous YM, between Kwon A, zone Bendesky populations. hybrid yeast experimental a in interaction DA of Muller Thompson J, analysis Funt JB, Preliminary Anderson (2009) S Kambhampati of BT, Aldrich S Ansell D, Albach R, Abbott supplementary its (and article published References this in included are study this files). during information analyzed or generated data All version. material final and the the data Wrote approved of Q.H. and Availability manuscript and the H.X revised J.W. authors research: interests All the Q.H. Competing Performed and Q.H. H. and A. J.W. J.W. model: paper: of the Foundation designed Science and Natural Conceived National of helps the draft their by for earlier Contributions supported Gao an Author Yongyong was on and work suggestions Wang valuable research Hao for Zhao, This [31772516]. Dong Xingying China Yanan Cheng, collections. Zhen Dr. termite thank and in also Chen We Chun manuscript. Liu, this Min Profs. thank still We will species of mechanisms maintenance the Acknowledgments: on future. studies the Further in nature. required in be integrity species maintain to otrossnevadensis Zootermopsis nuRvGenet Rev Annu . Drosophila o Ecol Mol Nature 544 24 Science . h uhr elr ocmeiginterests competing no declare authors The Science 35 6209-6222. , 434-439. , . 125-148. , netSoc Insect tal. et , 341 tal. et , tal. et , 314 tal. et , 667-669. , tal. et , 21)Hbiiainadspeciation. and Hybridization (2013) 1292-1295. , tal. et , 21)Hsoia igorpyof biogeography Historical (2016) 56 21)Oencdsesl iaineadhmnitouto shaped introduction human and vicariance dispersal, Oceanic (2016) Reticulitermes 20)ToDbhnk-ulrgnsitrc ocuehybrid cause to interact genes Dobzhansky-Muller Two (2006) 439-450. , 21)Dtriat fdvretaatto n Dobzhansky- and adaptation divergent of Determinants (2010) 21)Tegntcbsso aetlcr vlto nmonog- in evolution care parental of basis genetic The (2017) 6 urBiol Curr , Heterotermes Haemulon 20 1383-1388. , and ). Reticulitermes o Ecol Mol o hlgntEvol Phylogenet Mol Coptotermes vlBiol Evol J 26 hgltspomonella Rhagoletis 26 639-652. , . emts(Isoptera: termites rcBo Sci Biol Proc 229-246. , vlBiol Evol J Coenonympha 94 778-790. , 283 20 Posted on Authorea 26 May 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159050496.61216640 — This a preprint and has not been peer reviewed. Data may be preliminary. ol C aaaoT(09 ope psai o ozasyMle yrdicmaiiiyin incompatibility hybrid Dobzhansky-Muller for future epistasis for Complex strategy adaptive (2009) an T as Nakazato pairing solanum. same-sex termite LC, Male nasute Moyle (2016) the K of termites. Matsuura biogeography in T, and reproduction Yashiro phylogeny N, Molecular Mizumoto (2000) T Matsumoto Y, genus Roisin T, Miura S polymorphism Mwaiko DA, DNA radiations. Marques of JI, analysis in Meier termites. herd comprehensive selfish in as behavior for running antipredatory tandem software Homosexual novel (2002) : T a Nishida E, Kuno v5: K, Matsuura DnaSP at (2009) contents J nutrient data. and Plant Rozas weight P, fresh Rare termite ratio, Librado of subterranean sex the of Extinction Dynamics in (2014) alates the QY 51-57. of Huang and stages ZH, developmental Wang Hybridization five CL, (1996) Lei GH, RK Li Jansen males. J, in against Francisco-Ortega Species. selected but DA, females in A Levin favored S is U ants hybrid Sci developmental Acad in understanding Introgression Nat for Proc (2014) framework P Pamilo a J, - Kulmuni development and history termites. lower Life mode in (2008) reproductive plasticity K and structure Hartfelder population J, of parasite Korb analysis two Genetic of (2013) C divergence termite Lei the the C, in of Husseneder mechanisms G, Li isolation Q, prezygotic Huang of role The (2016) species. M Kalbe among T, adaptations Henrich mimicry of exchange labora- promiscuous by reveals genome proportions species. Butterfly (2012) soldier C of Genome Heliconius maintenance and soldiers of Rhinotermitidae). Production of groups (1981) experimental R tory Howard M, between Haverty hybridization and pairing Heterospecific review. comparative (2011) a and RB after termites: Rosengaus mtDNA of TR, biology of Hartke mating recombination The No (2011) B germline. (2014) Baer maternal NG TR, mouse Larsson Hartke the JB, in Stewart generations BJ, 50 for Battersby heteroplasmy C, Freyer E, maternally? transmitted Hagstrom genomes organelle burst-speed most are poor Why 80-94. (2015) by R , Bock supported J, Sobanski dysfunction S, Greiner hybrid salamanders. of tiger model hybrid in Dobzhansky-Muller performance (2008) termites the BM in Fitzpatrick depression inbreeding of test indirect An and (2005) EL Vargo CJ, DeHeer .ephratae N. virginicus Reticulitermes Bioinformatics Nasutitermes osr Biol Conserv M vlBiol Evol BMC Nature Genetics a Commun Nat eiuiemschinensis Reticulitermes 487 . netSoc Insect 181 Naturwissenschaften Iotr:Triia)i h aictropics. pacific the in Termitidae) (Isoptera: 25 94-98. , 10 347-351. , 1451-1452. , 16 10-16. , nmBehav Anim 8 111 14363. , 245. , ilRev Biol eiuiemsflavipes Reticulitermes 28 . ea clSociobiol Ecol Behav 12805-12810. , 32-39. , tal. et , 83 98 119 snyder. 295-313. , vlBiol Evol J 745-753. , ho Biol Theor J 179-187. , 21)Acethbiiainfesrpdccldfihadaptive fish cichlid rapid fuels hybridization Ancient (2017) LSOne PLoS Kla)and (Kollar) 59 7 21 753-761. , 214 342-351. , 8 e69070. , 63-70. , o hlgntEvol Phylogenet Mol eiuiemschinensis Reticulitermes eiuiemsvirginicus Reticulitermes uli cd Res Acids Nucleic nmBehav Anim 42 eiuiemssperatus Reticulitermes eiuiemsflavipes Reticulitermes auiemscorniger Nasutitermes 17 1111-1116. , 1-10. , . (Banks)(Isoptera: netSoc Insect 82 Bioessays 927-936. , 62 37 , Posted on Authorea 26 May 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159050496.61216640 — This a preprint and has not been peer reviewed. Data may be preliminary. yteraes re n e ersn h sinn rbblt otetomjrclades. major two the to probability assigning the STRUCTURE. represent by red assigned and samples Green of termite index areas. of diversity their clusters by genetic Genetic the 4. and termites. Fig. plot clusters violin two of The into morphology divided 3. were and Fig. haplotypes of tree haplotypes unique the Thirty-one (ML) with percentages. combined likelihood and bootstrap Maximum the indicate branches 2. 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Endepols reaction Y, chain Song polymerase conserved of compilation 87 a and sequences gene A on Beckenbach (Aquifoliaceae) F, Frati Ilex C, Islands. of Simon Ryukyu endemics the insular between and flow Islands gene Bonin Intersectional the (2000) I Watanabe H, radiation. Setoguchi adaptive and Hybridization (2004) O Seehausen California. introgression of and life. Hybridization of (2011) K tree Schierenbeck the speciation. up and Shaking hybridization (2016) on E Pennisi perspective genomic A (2016) LH 2337-2360. Rieseberg BA, Payseur Reticulitermes 651-701. , Reticulitermes and pce ne aoaoyconditions. laboratory under species eiuiemsaculabialis Reticulitermes Coptotermes tal. et , tal. et , .chinensis R. Reticulitermes . nuRvEntomol Rev Annu .chinensis R. 19)Eouin egtn,adpyoeei tlt fmitochondrial of utility phylogenetic and weighting, Evolution, (1994) 21)Aatv nrgeso fatcauatrdn osnresis- poison rodent anticoagulant of introgression Adaptive (2011) to clEvol Ecol Ethol Bot J Science . and u Biol Cur hso Entomol Physiol Iotr hntriia) eiwo h vdnefrom evidence the of review a Rhinotermitidae): (Isoptera 87 and .flaviceps R. 8 354 eoeBiol Genome 793-810. , n nylpdao ilgclInvasions Biological of Encyclopedia In: . .flaviceps R. Insects 54 21 817-821. , 379-403. , r nsEo Evol Ecol Trends 11 1296-1301. , 149-169. , 11:14. 11(1): .flaviceps R. 39 17 ouain ntreposo China. of plots three in populations . 44-52. , 25. , 19 and 198-207. , h ooisaearranged are colonies The .chinensis R. n noo o Am Soc Entomol Ann h ubr nthe on numbers The o Ecol Mol University , . 25 , Posted on Authorea 26 May 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159050496.61216640 — This a preprint and has not been peer reviewed. Data may be preliminary. 9 Posted on Authorea 26 May 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159050496.61216640 — This a preprint and has not been peer reviewed. Data may be preliminary. 10 Posted on Authorea 26 May 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159050496.61216640 — This a preprint and has not been peer reviewed. Data may be preliminary. 11 Posted on Authorea 26 May 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159050496.61216640 — This a preprint and has not been peer reviewed. Data may be preliminary.

Population Table 1. Genetic diversity indices for CO R. chinensis R. flaviceps Chengdu Chengdu Nanjing Nanjing Wuhan Wuhan s

15 17 38 10 14 2 N 6 5 9

Number of Haplotypes ( 18 13 N 9 7 4 9 2 4 h

)

Ⅱ 12

in R. chinensisR.

Haplotype diversity 0.848±0.088 0.657±0.138 0.679±0.112 0.854±0.041 0.978±0.054 0.400±0.237 0.644±0.152 0.81±0.079 ( h )

and

R. flavicepsR. Nucleotide diversity 0.02013±0.00371 0.01329±0.00626 0.00871±0.00499 0.03415±0.00249 0.00221±0.00131 0.02848±0.00794 0.01083±0.0026 0.0359±0.00222 .

( π )

Table S Chengdu Nanjing Wuhan ampling plots 2

- Statistics andestimates of gene flow between M itochondrial DNA 0.6206 0.7435 0.6912 F st

(mtDNA) 0.15 0.09 0.11 N m

N 0.087±0.013 0.039±0.016 0.117±0.022

R. flavicepsR. uclear F st

gene

and

(nDNA, SSR) 13.804±4.209 3.294±0.591 2.639±0.517 R. chinensis N m

Table 3. Percent of sampled R. flaviceps R. chinensis R. Population Chengdu Chengdu Nanjing Nanjing Wuhan Wuhan

(possessing mtDNA haplotypes mtDNA (possessing s (possessing

Number colonies Number

colonie mtDNA haplotypes mtDNA 17 1 15 10 5 6 4

s assigned to genotypes based on ten

14 Percent of colonies assigned to genotype to assigned colonies of Percent R. chinensis

of of 0.015 0.04 0.23 0.978 0.974 0.85

R. flaviceps R. R. chinensis 8 7 1

) )

R. flaviceps 0.98 0.952 0.76 0.02 0.02 0.149 5 3 2 6 SSR

markers s