Posted on Authorea 17 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159769281.18634046 — This a preprint and has not been peer reviewed. Data may be preliminary. n oecaua eiw Rne ta. 04 hmcie l,22)a ela hntccmaio wi- comparison a phenetic be a to as well seems as analyses there 2020) 2015), phylogenetic date al., al., To et et including Chomicki (Achigan-Dako 2011a). research, 2014; genus Renner, recent al., the & et to thin (Renner Schaefer According reviews taxonomy. 2011b; nomenclatural mor- complex Renner, and high confounded its & exhibit years Schaefer regarding which many 2007; of consensus for all al., was varieties, et watermelon and Kocyan subspecies, of the species, routes of diversity. among colonization distribution phological confusion the historical taxonomic on the significant focused tracing by have 2020). of et studies neither al., challenge few Mujaju et found The comparatively 2013; Chomicki were al., and al., relatives 2019; et Africa; et wild al., Dane east within Levi et no 2007; variation northern 2011; Renner as Liu, genetic in Nesom, 2015; remains & 2000; nor 2010; Dane Renner, questions Newman, west al., vis-`a-vis 2005; & these & in et uncertainty Thomas, Chomicki Solmaz & Jarret efforts, 2014; 2010; Levi these 1997; Gladis, 2004; al., Despite & al., al., et Hammer et et Dje 2013; Levi 2009; (Jarret al., 2004; Sari, times al., & divergence et Solmaz and Dane 2007; 2001; relatives, origin wild al., of of identify et taxonomy nomenclature, centers Levi and and origin both relationships the million phylogenetic determine regarding 3.2 clarify and from questions to 2018 decades, studies in two numerous production last fuelled fruit the have global Over of tons 2017). metric (Faostat, million ha 103,9 nearly for accounting importance, ( Watermelon investigations phylogeographical and historical Introduction Africa. further for North-East calls mucosospermus, and This C. West of lanatus. in endemism C. samples of to of center of origin mucosospermus a the C. collection as of findings wider search Africa, previous Some West the and support in cordophanus). interest results haplotypes). of The subsp. area (including (16 colocynthis. an C. lanatus colocynthis is haplotypes C. C. lanatus, shared and to C. mucosospermus amarus relative C. was C. The closest C. and the in both diverse continent. lanatus be of the haplotypes most C. those to and the 38 from West-Africa, endemic distinct to and identified appear were exclusive 17 haplotypes) that we nearly and were (5 total, mucosospermus Africa mucosospermus C. in In (trnT-trnL of found C. sequences haplotypes were was accessions. cpDNA 21 species these, 135 non-coding of diverse of using colocynthis; least colocynthis collection C. and haplotype C. global amarus, and and a C. biogeographical north-eastern amarus, mucosospermus, of from infer suggests we C. center regions) study, hypothesis mucosospermus, the this ndhF-rpl32 second C. In suggests a and lanatus, thrives, cultivated. hypothesis mucosospermus is C. first C. melon for a Kordophan species genealogy Sudanese endemic While white-fleshed sister the a debated. where where remains Africa Africa, lanatus) west (Citrullus be to watermelon origin of origin geographical The Abstract 2020 17, August 3 2 1 Achigan-Dako Gbenato Enoch wild species of Citrullus collection cultivated global and a of analysis phylogenetic Georeferenced ebi nttt fPatGntc n rpRsac (IPK) Research Crop and Genetics USA Plant Hampshire, of New Institute Durham, Leibniz Hampshire, New of University available not Affiliation irlu lanatus Citrullus Citrullus Citrullus crd xEk eh soeo 5gnr fCcriaee(ery 2005; (Jeffrey, Cucurbitaceae of genera 95 of one is Zeyh. & Eckl ex Schrad. Tub)Mtu.&Nki sahriutrlseiso iheconomic high of species horticultural a is Nakai) & Matsum. (Thunb.) rtelkl ooiainrue fvrosseiswti h genus. the within species various of routes colonization likely the or 1 ev Degbey Herve , Citrullus nopse h olwn ee pce:1 h widely the 1) species: seven following the encompasses 1 1 aoHale Iago , 2 n rn Blattner Frank and , Citrullus 3 spp. Posted on Authorea 17 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159769281.18634046 — This a preprint and has not been peer reviewed. Data may be preliminary. hssuy ed falacsin eegriae nagenos tIKGtrlbn n approximately of and part IPK-Gatersleben, Germplasm as at Plant greenhouse Africa West National a USDA in throughout germinated the collected were from were accessions received 16 all were and of accessions IPK-Gatersleben, Seeds 53 study. from from of this received total 16 were a and 66 1, Europe, System, Table from in Institute 25 the (IPK-Gatersleben). indicated of 2015) Asia, As herbarium al., the from in et 41 deposited (Achigan-Dako to were Africa, Genetics accessions relative from Plant all closest of of 53 specimens the Voucher including 1). and (Table accessions) assessed, America Europe North were (13 to accessions watermelon spread 135 cultivated has total, of that species Africa and sister Southern accessions); closest from four 2) the the species Africa); West and study wild from were Africa the 14 West only in which to included of out we continents four haplotypes, from accessions watermelon important of economically distribution most geographical the investigate isolation To DNA genomic total agricultural and significant sampling Taxon globally this of methods history and the Materials of understanding evolution our sequence clarify chloroplast of species. help distribution the geographical will of the information light characterize such to specific is shed objective and The haplotypes continents. do four from how collected and of size phylogeography world originated ( chloroplast the cultivated has throughout a one presents distribute it cultigens paper if as extant this such world the types questions, the wild did throughout to How spread relate Africa? they watermelon north-east did or how west from remain: West questions fundamental from not The records was of archaeological 2015). variety farming ( al., contrast, species or when cultivated et Valley subspecies In Achigan-Dako endemic Nile a 2014; one 2000). of the be Hopf, presence 8 in to & the the cultivated deemed are for (Zohary between watermelon been except scanty, Africa period have of are recent southwest to records Africa relatively in archaeological known 2015; a practiced few is Paris, Sudan, to very yet cultigen 2004; in date Veen, studies, a area all Der latter Furthermore, river Van and these A.D. & Nile Africa; (Wasylikowa to (Bates the southern data desert According from especially archaeological Namibian 2019). known on Africa, the based al., north-east in origin et or relatives of Renner northern center wild to likely of point the distribution as others the 1995), on origin Robinson, of based center & Africa, possible southern the from as Africa originated west indicated (2015) Renner of and Chomicki study, phylogenetic understanding recent this of 2004); clarification Lang, recent & Given (Dane sampling. variation extensive intraspecific more little with very change with may Namibia of region desert the 5) eccirhosus genus ; the Citrullus Near-East 7) in and and placed animals; Africa previously desert region, northern feeding apple for throughout bitter used areas the fruits sandy 4) small 1983); in Fursa, growing 1981; species Fursa, ecirrhosus melon perennial 1972; (Fursa, egusi a ago the Schrad., decades 3) (L.) many 1976); rank Bemis, specific & to (Whitaker Africa southern in spermus grows which Mansf., (Bailey) melon tsamma the 2) Fursa; (Ter-Avan.) cultivated .lanatus C. us,peiul eerdt sasbao of subtaxon a as to referred previously Fursa, on,aohrprnilwl pce D-itr 90;6) 1990); (De-Winter, species wild perennial another Cogn., .lanatus C. li tod iherirasrin.Ide,weessm xet eiv watermelon believe experts some whereas Indeed, assertions. earlier with odds at claim a , .mucosospermus C. .colocynthis C. , uc ri on ntoia n utoia lmtsicuigvar. including climates subtropical and tropical in found fruit juicy a , C . rehmii, Citrullus Citrullus idseisfudi otenArc n atAi 2 cesos.In accessions). (22 East-Asia and Africa northern in found species wild a , .colocynthis C. and n w id( wild two and ) Acanthosicyos aooy ti prpit orvstteqeto fgnaoy na In genealogy. of question the revisit to appropriate is it taxonomy, pce:1) species: .naudinianus, C. .amarus C. .lanatus C. or .naudinianus C. .amarus C. .lanatus C. Nsm 01 amr&Gai,21;Rne tal., et Renner 2014; Gladis, & Hammer 2011; (Nesom, crdsyn. Schrad .amarus C. ew xHo.f n itrgopt l te species. other all to group sister and f. Hook. ex Welw. 2 urnl,aecniee nei n etitdto restricted and endemic considered are currently, .lanatus C. ocnrbt oorudrtnigo these of understanding our to contribute To ? ieycliae hogottewrd(78 world the throughout cultivated widely , irlu lanatus Citrullus .caffer C. and Sn. okf rmteNamib-Kalahari the from Hook.f. (Sond.) ymn uhr u hc a raised was which but authors many by .colocynthis C. .rehmii C. crd or Schrad. fC lanatus C. of .mucosospermus C. .mucosospermus C. idana pce,with species, annual wild a , n he eae species, related three and .lanatus C. ,uigalresample large a using ), th yohszn that hypothesizing , ; n 13 and 3) .ecirrhosus C. .amarus C. .colocynthis C. var. cordophanus previously ) th .mucoso- C. restricted , centuries Citrullus citroides (22 a , C. Posted on Authorea 17 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159769281.18634046 — This a preprint and has not been peer reviewed. Data may be preliminary. igegpfr6(T,Afr7(T,adTfr8(T;ad6 GT irstliea oiin1198 position at microsatellite TGATT A (absence). 6) gap (absence). and single 8*(AT); gap a for single or and T (presence) a (absence); T and or gap 7*(AT), a single (presence) as for a T coded A or was 6*(AT), as (presence) for coded T gap a was single as 423 coded position was the at 405 In position microsatellite TTTATA the constructing at In Another of indel TATT polymorphisms. 3) purpose bp A the single 2) for as (absence); alignment, re-coded ”-” were combined indels the and In repeats trn nucleotides. haplotypes, aligned parsimonious 1,611 and coherent to of matrix submitted a were 1) yielding reads generated: merged were alignments these F- into three and combined merging, 2012); and Following colocynthis al., available. edited publicly et of manually alignments them (Kearse were make Species-pairwise 5.5.6 sequences to Geneious reverse GenBank using and NCBI sequence forward single the region, a chloroplast capillary each Biosciences) For (Applied coding XL performed haplotype 3730 was and ABI analysis Sequencing an Sequence buffer. or TE Biosciences) 1x (Amersham warmed 1000 ml sequencer. MegaBACE 28 a in either on re-suspended and for instructions, conditions manufacturer’s Cycling DNA. genomic 72 of at ng 20 30 approximately and of 98 primer, volume each reaction 1.5 of a additional pmol in an Scientific) (10x), buffer (Thermo supply Kit 68 Start at Hot min Phusion 10 of extension and final primer, for 95 a each conditions at of Cycling s pmol DNA. 30 50 Germany), genomic of Hilden, of (QIAGEN, ng polymerase 20 DNA approximately Taq cycler. U thermal 1.5 2.5 Biosystems) (Qiagen), additional (PE Q-solution System an PCR (10x), 9700 buffer Amp Gene a the using For performed were amplifications PCR sequencing and Amplification 3’). hs w ein fteclrpatgnm eeapie sn h olwn rmrpis 1) pairs: investigation. primer in-depth following more the for using selected amplified were were regions genome two chloroplast 2) these the thus of (5’-GAAAGGTATKATCAAYGMATATT-3’) species; regions and two among These and within for However, both inter-specific. exclusively and 16-5’ ae ntewr fSa ta.(07,tefloignn o-oigclrpatrgoswr cho- were regions chloroplast non-coding nine following of the each (2007), accession al. one et of Shaw screening locynthis initial of for work sen the on Based 1x PCR ml regions 50 sub-optimal chloroplast in exhibiting resuspended of Samples and Choice (QIAGEN) bromide. Kit ethidium Concentrations Purification PCR DNA. with buffer. QIAquick the genomic stained the TE Total of step via gels quality gel. washing purified the agarose silica one were increase and with 1% amplification Kit, to dried Plant on instructions and DNAeasy manufacturer’s estimated QIAGEN accession the the were per using to seedling tissues according leaf one added dried from was the collected from was extracted was tissue DNA leaf of mg 100 trn rpl ° o is 5cce f3 t98 at s 30 of cycles 35 mins; 3 for C - ein )amcoaelt CT tpsto 6 a oe sA(eetpeec)o igegap single a or presence) (repeat A as coded was 366 position at ACATA microsatellite a 1) region: T-L trn ° L .AlPRpout eeprfiduigteQAuc C uicto i QAE) following (QIAGEN), Kit Purification PCR QIAquick the using purified were products PCR All C. ndh 2 n )acmie lgmn falseis otiigboth containing species, all of alignment combined a 3) and 32; (UAG) trn K, : rpl F- o h hools region chloroplast the for ° T- atp ,3 t56 at s 35 C, rpl 5-TCTCAGGACT3)and (5’-CTGCTTCCTAAGAGCAGCCT-3’) 32- trn 2rgo:1 oyA,uulysxt ih nt psto 19,wsjs elcdb a by replaced just was 1149), (position units eight to six usually AT, poly a 1) region: 32 I- trn atp ein eue ecinvlm f50 of volume reaction a used we region, L L, ,and H, trn ° ,ad9 t68 at s 90 and C, Q-5’ trn .lanatus C. ° μ .Frthe For C. f1 MMgCl mM 15 of l rps T- trn μ trn 6 3’ 16, f1 MMgCl mM 15 of l ° ,3 t58 at s 35 C, .Frms fteergos oa eeso aito eelow were variation of levels total regions, these of most For L. -;2 h aeseispiws lgmnsfrteregion the for alignments species-pairwise same the 2) T-L; cesoswt hs of those with accessions ndh ° ndh ;3 ylso 0sa 95 at s 30 of cycles 35 C; trn F F- -rpl V- rpl .lanatus C. rpl ndh 2 . Mo ahdoyuloietihsht,10 triphosphate, deoxynucleoside each of mM 0.2 , 2rgo,PRapicto a are u sn the using out carried was amplification PCR region, 32 3 ° 2R(’CATTCTYTTCA-’;and (5’-CCAATATCCCTTYYTTTTCCAA-3’); 32-R ,ad8 t72 at s 80 and C, 2and 32 C, 2 . Mo ahdoyuloietihsht,50 triphosphate, deoxynucleoside each of mM 0.2 , trn ndh trn T C , trn μ (GGU) ossigo 26.6 of consisting l F- uoopru,C amarus, C. mucosospermus, . T- rpl T- .mcssems .amarus C. mucosospermus, C. trn ° ,3 t53 at s 35 C, trn (5’-CCCTTTTAACTCAGTGGTAG- 32, μ ein 95 region: L ° trn ossigo 17.7 of consisting l ;adafia xeso f1 min 15 of extension final a and C; psb ,plmrhsswr observed were polymorphisms L, - and T-L D- trn ° ,ad9 t68 at s 90 and C, ° μ o is 0cycles 10 mins; 3 for C T, H l ndh ndh psb 2 ,5 O, F- F μ rpl J- -rpl H l pet μ and fsupply of l 2regions, 32 2 ,6 O, 2region: 32 ,3’ A, and , ° ;and C; ndh .co- C. μ ndh of l rps C. μ F l Posted on Authorea 17 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159769281.18634046 — This a preprint and has not been peer reviewed. Data may be preliminary. nomtv iewsfudwithin amplified found was amplified site informative the of length The diversity Illustrator. interspecific Adobe and using intra- output variations, TCS Nucleotide the from re-drawn performed was and (2006) network substitutions, The Results al. base et (2011). underlying identical Banfer by rates al. collapses described mutation et and strategy Guicking different two-step sequences the by the followed of we for relationships microsatellites, account genealogical and To indels, estimates (HT). TCS TCS haplotypes using into 2000). haplotypes sequences among al., relationships et phylogeographical (Clement infer v1.21 to constructed comparison were networks for Parsimony implemented was also network test parsimony D Statistical Tajima’s population. equilibrium. overall Fu’s nucleotide population the of the 2016; with and homogeneity within Nunez-Farfan, neutrality of differentiation & selective hypothesis haplotypic Chavez-Pesqueira et null of investigate Burban 2013; Fu’s the to 1996; performed tested al., test Petit, also we et We exact & (1996), Chiu (Pons Fisher’s Templeton chance 2011; Following using by al., mutations 2019). expected sequence et al., be considers Guicking et would Nst 2002; Sun than Gst, Grivet, area to geographical contrast 1999; given (Gst). In al., a frequencies permutations. in 1000 Nst haplotype often with Thus, using more 2013) haplotypes. by al., G the et between and between Chiu difference differences (Nst) 1996; the haplotypes Petit, on based & between based matrix (Pons distance tested steps a was another. mutational computing structure one of by Phylogeographical from estimated number different was esti- genetically the populations differentiation are Carlos among on genetic Monte differentiation populations of on genetic geographical We analysis based addition, the Chi-square method In whether traditional non-parametric sub-populations. understand the simple geographic helped to a among mates, test, compared differentiation statistical method, (1992)’s genetic That al. of permutations. frequency- et allele estimates an Hudson on (1983), used Chesser relies further and popula- Nei that geographic by developed approach among statistic differentiation based differentiation genetic gene to the the considered computing assessed were by We sub-region tions or populations. continent the geographic by distinct study), grouped haplotypes comprise the parameters, differentiation in 2011). genetic (Baier, infer once To estimated only haplotypes structure also of occurs were genetic number diversity and that the haplotype differentiation as (haplotype overall inser- Population such the haplotypes used metrics of and We singleton intra-population haplotypes, number Other effective of 2007). the of 2019). number number al., al., is the et et (h ID population, Zhao Dane diversity diversity 2019; gene per transitions, gene 1992; al., within-population total of et (O’donnell, mean the the Sun number sequence and calculate 2013; the to the al.) 1996) of is Petit, et length & (Ching-Yi Ts (Pons total package transversions, software the PERMUT is of the L number and the tions/deletions, is formula: Tv the region ID)/L following where chloroplast + site, each Ts per for + differences divergences (Tv nucleotide x sequence of Rozas, 100 inter-specific number & and (Librado mean Rozas, 5.10.01 intra- the & version Pairwise as software Librado DnaSP computed 2013). 1997; 1987), using were al., Rozas, computed Nei, et were & 2013) 1983; Chiu (Rozas al., Tajima, 2009; rates et transversion Chiu & 2011; and (Nei Baier, inversion, 2009; diversity substitution, nucleotide and diversity, content, sequence A+T including parameters Statistical diversity genetic of Analysis analysis Data ndh Fs F- rpl . 2rgo agdfo 5-5 pi h pce,as ihn parsimony-informative no with also species, the in bp 650-652 from ranged region 32 Fs F,19)t nlz h xaso ee ftepplto ne h hypothesis the under population the of level expansion the analyze to 1997) (Fu, trn T- .lanatus C. trn einwithin region L u neswr on tpstos22 9,ad26 The 296. and 295, 242, positions at found were indels 3 but , > T s niae htcoeyrltdhpoye r observed are haplotypes related closely that indicates Gst Mri ta. 03 ucige l,21;Ci tal., et Chiu 2011; al., et Guicking 2003; al., et (Martin ) 4 .lanatus C. agdfo 5-5 p oparsimony- No bp. 951-954 from ranged ST n N and ST sn EMT2.0 PERMUT using Posted on Authorea 17 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159769281.18634046 — This a preprint and has not been peer reviewed. Data may be preliminary. ih altps mn hc i eepiae n vrl altp iest s0.81. is 1149. diversity position haplotype at overall and microsatellite colocynthis private; a C. were is six there which and among 1526; haplotypes, eight and 1397, 1149, 918, Goga uolva tl n kan) sa(usa aa,Cia ni) n ot mrc (USA America shared North (India) and Asia India), Europe southern in China, and also to (Egypt) Japan, specific but H4, countries Africa (Russia, haplotype African North Asia West among Ukraine), Canada). only and and not Italy found was Yugoslavia, Asia H1 in (Georgia, haplotype haplotypes ancient of The number singletons. highest 1). the (Fig. contributed Africa Iran Within in species, did this Egypt For as of Africa. 1), haplotypes to some (Fig. specific continent, were that six Across 5). and Asia. - throughout 2 widespread of Figs. found colocynthis Haplotypes (see also region were Africa. that Africa southern Europe, to northern to in were specific specific found were 16 nine appeared two Africa, the types America, of in majority from specific; found recovered of were the haplotypes four eight Haplotypes with 21 Asia, the in the Africa), of found and of (in haplotypes specific; instance, 14 0.85 For were the to six of regions. continent; Europe) certain the (in to to specific 0.57 specific from being p varied haplotypes 0.1958, of (Hs) = diversity (LD gene substitutions Within-continent nucleotide evolution. rejection of the molecular homogeneity suggested of Fu’s population of theory both overall neutral hypothesis the by null within revealed differentiation the haplotypic as of investigate to selection used neutral test exact non suggested polymorphism Tajima’s of expansion pattern population and The haplotypes of analysis, differentiation TCS genetic distribution, to Geographical according detected were (H1), haplotypes haplotype 1). within 38 ancient found (Fig. were regions, most species (50%) cpDNA The cultivated 13 two the detected, the 3). to (Table within exclusive detected accessions (6 is sites informative sampled parsimony polymorphic the were 29 among which the 0.96. of on of 7 diversity Based 1530), haplotype (positions and overall indels 1262, an 11 1200, has were there and 1180, private, addition, 1179, In 972, 1149). 297, 423, within 296, 366, 295, (positions 242, microsatellites 3 199, and 1526) and 1397, for bp 650-653 and L in 1 and 405) 297, 296, 295, (positions in is bp in diversity 651-653 haplotype length overall sequence and combined singletons; The the were three in species, identified this was of in accessions 0.5333. informative) indels sampled two (non-parsimony the as among SNP well found One as 1397), bp. (position overall 1601-1604 an by with singletons, ning sampled were within 10 the and lengths combined which total, 1399) Sequence among The 2). (position In haplotypes, (Table SNP S1). 0.5656 distinct parsimony-informative. 1 (Tables of 12 included were 1198 diversity comprise and polymorphisms haplotype and bp species no 1601-1605 1178, this but to of 1143, equal 366); accessions 1028, found (position 970, was microsatellite regions positions cpDNA 1 at two found the were of length indels 5 though site, trn .lanatus C. D eeseict ieetcutis(i.1.Six 1). (Fig. countries different to specific were T- F 364 .1;D 0588 o ttsia infiat p significant, statistical not -0.59858; D: 0.016; = p -3.624, = (Fs ndh trn a hrceie yacmie eunelnt f19-65b 9894b for bp (948-954 bp 1599-1605 of length sequence combined a by characterized was .mucosospermus C. F- n within 1 and L lhuhalrgossae othpoye,Arc xiie h ihs ubrof number highest the exhibited Africa haplotypes, most shared regions all although , rpl ndh 2 n otie e oyopi ie.O hs,4idl eeosre in observed were indels 4 those, Of sites. polymorphic ten contained and 32) .mucosospermus C. .amarus C. F- rpl 2 htfaue 0SP pstos46 5,47 8,98 4,11,1286, 1111, 949, 918, 882, 487, 455, 406, (positions SNPs 10 features that 32) ndh .amarus C. eeams nqeyrsrce oWs fia and Africa, West to restricted uniquely almost were a hrdb fia onre eg ot-fiaadteDemocratic the and South-Africa (e.g. countries African by shared was , .colocynthis C. .lanatus C. F- ndh rpl trn eesmlr ihtecmie egho h w ein span- regions two the of length combined the with similar, were F- 2.Tecleto fti pce otis1 altps all haplotypes, 16 contains species this of collection The 32). T- agdbten10-64b 9093b in bp (950-953 bp 1602-1604 between ranged rpl trn and 2(oiin 18.Fu Nswr on tpositions at found were SNPs Four 1198). (positions 32 ein(oiin 4 n 9) fte5haplotypes 5 the Of 296). and 242 (positions region L niaigrcn altp iegnei htspecies that in divergence haplotype recent indicating , 5 .mucosospermus C. .colocynthis C. .colocynthis C. .colocynthis C. hrdb aii,Ehoi,and Ethiopia, Namibia, by shared fte2 igeo haplotypes singleton 26 the Of . altpswr pcfi oAsia, to specific were haplotypes .amarus C. > .0.Mroe,Ficher’s Moreover, 0.10). < a hrceie by characterized was ndh .0)floigthe following 0.001) altp 1;and H12; haplotype .amarus C. Fs trn F- rpl ttsi and statistic T- trn 2region 32 trn trn haplo- and L T- T-L trn C. Posted on Authorea 17 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159769281.18634046 — This a preprint and has not been peer reviewed. Data may be preliminary. w pN ein n 3 cesoscrflyslce orpeetawd egahclrgo,w detected we region, geographical wide a represent to selected carefully accessions 135 of and accessions regions 55 cpDNA using two genus, the within haplotypes colocynthis 2018). seven C. al., found wild (2004), et and cultivated (Endl al. the et Africa Dane among in detected and were Asia haplotypes in Thirty-eight twice: least Such at colocynths. domesticated was African Citrullus and and within lineages on Asian two diversity studies of to recent haplotype back the comparison The by go morphological exemplified comprehensive (2017). is a investigation an al. for et calls Levi and evolution by confirmed also within cultivated was detected those haplotypes in that structure revealed genetic 2019). current egusi al., colocynthis et to and C. Renner watermelon contribute type 2015; seed of might of al., histories consumption selection et domestication or for (Achigan-Dako content the protein/oil species lycopene in high diversity, of with bottlenecks sequence cultivars source other red-fleshed within influence as sweet or divergence can for haplotype selection selection factors of Certainly, of lack many melon. result The (Dane While diversity the large 2013). reduce substantially likely al., a (2013). can et is that used bottlenecks Levi al. of we 2004; imposition et Lang, study, the Guo & via this contributor by major In a revealed is also selection origin. as geographical species, or that traits morphological of by number either grouped accessions, the ( than species regions cultivated two the the within that the revealed events of analysis transition sequence resolution and Chloroplast sufficient indels was variability. on intraspecific there mainly reveal However, based to divergence regions 2004). with al., another et one from (Dane separated highly were Taxa o ubro asmn-nomtv ie ihnec ftesuidseis othpoye eefound on were based haplotypes variability Most nucleotide species. low studied revealed the who of (2004) each ( al. within non-cultivated sites et within parsimony-informative 2013). Dane of al., and number et (2004) among low (Levi Lang a variability resistance and cpDNA disease Dane like by low observed traits revealed valuable Oligonucleotide of findings Frequency sources High Our as using accessions study PI among a diversity of instance, genetic importance For high potential revealed genus. (HFO-TAGs) Genes the Active within Target diversity genetic obvious on in diversity 20 genus the Within variation sequence and diversity Genetic and Africa between between lower difference was between pairwise Gst the highest Discussion other; and were the 0.196, was on values 0.374. Gst Europe, differentiation = differentiation and among population Nst genetic Asia of differentiation haplotypes and coefficient pairwise genetic hand, The of total one (0.006). the Coefficients of Asia on level Europe, 5). high and and a Africa revealed 4 differentiation (Tables genetic continents interspecific by of shared Analysis is H2 by Burkina- Armenia), Haplotype shared and (Benin, is Canada). Mongolia, Africa H6 Republic, and West North-Korean haplotype (USA throughout Yemen, found America Japan, was (China, North Asia H2 and in France, Haplotype as well 1). as Ghana) (Fig. and Russia Faso, and Congo) of Republic th etr Hsiuee l 96 eeln aiu rns rvoskoldervae oe genetic lower revealed knowledge Previous trends. various revealing 1996) al. et (Hashizume century nhd altp evolution haplotype .lanatus C. Citrullus F, n oa fsvncDArgos(if,Ra,Tq,Au,HeI,Mo,adBlI.With BglII). and MboI, HaeIII, AluI, TaqI, RsaI, (HinfI, regions cpDNA seven of total a and , xiie eaieyhg ubro asmn-nomtv hrces aee l (2004) al. et Dane characters. parsimony-informative of number high relatively a exhibited ycf Citrullus o reigproe(eie l,20;Lv ta. 04.Rcn tde hdlight shed studies Recent 2004). al., et Levi 2001; al., et (Levi purpose breeding for 6- cesosfo iegorpia ag n bevdlwhpoyedvriywithin diversity haplotype low observed and range geographical wide a from accessions .colocynthis C. psb .mucosospermus C. eei iest nlsshv enivsiae ic h eodmdl fthe of middle second the since investigated been have analyses diversity genetic M, ycf 9- trn ahrta utvtd( cultivated than rather ) ,and G, .colocynthis C. trn and - ein aeadLn 20)aaye orchloroplast four analyzed (2004) Lang and Dane region. T-L ndh atp .amarus C. F- 6 A- rpl .lanatus C. uui melo Cucumis trn eeascae ihgorpia rgnadthat and origin geographical with associated were 2rgo xiiscmaaieyhge variability higher comparatively exhibits region 32 )adfudn aiblt ihncultivated within variability no found and R) pce seFg.2-5). - 2 Figs. (see species .lanatus C. and Citrullus trn htrvae oenmlncultivars melon modern revealed that .mucosospermus C. - and T-L Citrullus .lanatus C. and .lanatus C. .colocynthis C. cesosue nti study. this in used accessions .mucosospermus C. .lanatus, C. ndh p.adhglgtdthe highlighted and spp. and F- and .mucosospermus C. rpl hswsalso was This . .amarus C. ugsscryptic suggests 5acsin of accessions 15 2non-coding 32 species. ) and ; Posted on Authorea 17 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159769281.18634046 — This a preprint and has not been peer reviewed. Data may be preliminary. rgntn rmWsenArc Rne ta. 04.Hwvr otebs forkolde owild no knowledge, our of those best but earlier, the found to been have However, may plants of Spontaneous types 2014). ancient Africa. species al., West two and in the et 2007) confirmed among Liu, been (Renner diversity resulted have & haplotype Africa populations 560006) (Dane limited Western PI fixation very x chlorotype from The Gray with 2004). originating Charleston split, al., are old et (e.g. they an (Gusmini However, them sterility suggests between species. of crosses two levels previous these high between as in relationship species, between close different among relationship that two phylogeny close of indeed infer infer evidence both further to to of provide tomb regions accessions continents, Egyptian gene by the eleven shared from used mucosospermus highly sample who was sampled leaf was (2019) 3500-year-old and haplotype a Benin) ancient al. and This et Nigeria Renner other haplotype. with (i.e. ancient connections Africa most most West the lanatus the has in considered and frequently haplotype be sampled most may frequently most H1 geographic the thus wider evolution- is haplotypes; and H1 associated alleles study, sizes and subs. this older sample lineages of In on that lack descending depend studied. predicts the of may also theory observation number and an coalescence higher histories such ary/domestication a While 1993), Templeton, to & root. due (Crandall the population distributions at a in placed prevail were will Africa West throughout pled of 2015). haplotypes Citrullus regions Renner, Also, different & Chomicki diversification. from 2007; of sub-samples Liu, centers with & likely conclusions, their (2017)’s amarus with C. associated origin al. differentiation et and genetic Levi exhibiting diversification support of of statistics accessions centres differentiation that their observed four 2019). with all (2017) al., population agreement across the et diversity in al. Zhao structure gene 2019; et strong total al., a Levi the permutation et indicating of is Sun value without 0.0320) 1996; significant = calculated Petit, the error parameter & Also one standard differentiation (Pons 0.917, 2011). the the = al., (hT than that et regions fact Guicking greater geographical 2007; The is al., alleles et watermelon. Dane cow between for difference Nst 0.81 the was pairwise (i.e. = account difference the Nst the into on but 0.71, takes 0.374; based = that and Gst 0.196 alleles and to between watermelon equal the respectively, difference and (P found (Gst) pairwise significant were haplotypes not the (Nst) among haplotypes on distances among the based distances account differentiation no al., with of differentiation et coefficient population (Jiang of continents coefficient the The between structure occurred geographical and be differentiation would demographic Genetic distance high a by is there isolation that of indicate significantly might pattern 2016). neutrality produce from to a departure tend expansions the and bottlenecks case Population population expansion our while In non-random. D. D, is of of values observed values positive negative substitution. pattern significantly nucleotide polymorphism produce of to the homogeneity tend that for hypothesis test the Fisher’s supporting the by supported species, test observation other Fu’s an with rates, comparison in different In species in at per 16. species to haplotypes per 5 9.8 haplotypes from found 2.83 varying (2011) species, al. per et haplotypes Guicking Sudanese 9.5 observed the we particularly, average, On sequenced, 2019). are al., Africa et northeast of (Renner or sampling scarce Unfortunately, Sudan melon. from white-fleshed accessions sweet watermelon more among as diversity haplotype higher even an and > Fs hools eune nlsswt C .1rsle nantokweehpoye ieysam- widely haplotypes where network a in resulted 1.21 TCS with analysis sequences chloroplast eemsl rue nSuhr fia hc sasmdt eteoii fta pce (Dane species that of origin the be to assumed is which Africa, Southern in grouped mostly were s)idctsta h olcini hrceie ycergorpi tutr Gie,2002; (Grivet, structure geographic clear by characterized is collection the that indicates Gst) .mucosospermus C. lorjce h ulhptei fnurlt feouino uloiesbttto,further substitution, nucleotide of evolution of neutrality of hypothesis null the rejected also u nig,bsduo ag e feuimlnadwtreo cesosfo four from accessions watermelon and melon egusi of set large a upon based findings, Our . > .5.In 0.05). Citrullus Hordeum hs eut upr h nig fCoik n enr(05 and (2015) Renner and Chomicki of findings the support results These . p.Mjj ta.(01 on s .6adNt=04 o sweet for 0.49 = Nst and 0.56 = Gst found (2011) al. et Mujaju spp. In . Citrullus Citrullus .colocynthis C. p.Ti iuto a eepce ocniu oevolve to continue to expected be can situation This spp. 7 cordophanus p. uloiesbtttosapa ohv evolved have to appear substitutions nucleotide spp., .lanatus C. Macaranga . u nig niae htrgoa genetic regional that indicated findings Our eesbdvddit v rusi general in groups five into sub-divided were rmteDru eino ua a been has Sudan of region Darfur the from fo otes fia ntegermplasm the in Africa) northeast (from n ao n ltnr(06 found (2006) Blattner and Jakob and Citrullus C .lanatus C. pce,adalso and species, . mucosospermus and C. C. Posted on Authorea 17 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159769281.18634046 — This a preprint and has not been peer reviewed. Data may be preliminary. re,Mcrnawnlr,M ikeilaadM aais(Euphorbiaceae). tanarius M. and R. winkleriella F. M. Blattner, Halle-Wittenberg, (C. & winkleri, egusi-type Macaranga S., of trees, R. (2011). differentiation Vodouhe, C. and A., Baier, spp.(Cucurbitaceae) Ahanchede, Citrullus C., of Linsoussi, characterization mucosospermus). S., Phenetic E. Avohou, (2015). G., E. Achigan-Dako, Schae- Hanno and R.S. making ac- map Wohlbier, We manuscript. in References Birgit the (IPK). assistant grant Oswald, on for Research under comments Ganse Petra Plant useful Augustin author Koch, Crop thank for We Christina first fer and Ahanchede. from the Genetics Adam support to and Plant technical Vodouhe, Fellowship of Vavilov-Frankel and Institute the contribution Leibniz knowledge by the supported and financially CONT/08/136/RF was study current This origin. the Kordofan of and and places crops melon in major relatives egusi of wild of Acknowledgement routes of relationship colonization diversity close the genetic The regarding extant among questions of haplotypes world. new status chloroplast the raised distinct across watermelon 38 to haplotypes reveal melon those results of Our distribution the data. current and previous the understand genus The fully to cultivars. modern necessary of are history Conclusion continents domestication the all resolve from to regions collections region north-eastern those germplasm among in From More relationships domesticated ago? phylogeographical it years when? 3500 was than and when more If and Asia, watermelon how red-fleshed was haplotype. sweet Africa, unique species cultivating north-eastern were shows which or Egyptians 525083) west Kingdom PI New from as world (e.g. the region with to the clustered spread as from rather accessions 525083 such other PI species while that (H32). endemic showed haplotype (2017) an be unique a how al. to exhibits et indicated watermelon, Levi 525083), dessert of from var with (PI findings away 2015). cluster Previous mutations accessions (Paris, to several Egyptian haplotype. found (2013) is the from been al. accession have of materials et That watermelon one Levi more by of include that observed populations should showed and investigations wild data molecular where published our New newly countries Moreover, on neighbouring knowledge here. and sufficient reported Sudan generate those to investigations including between archaeological relationship results, and close molecular the and further with hand for affinity one genetic the on the cordophanus Africa or West 2015) in haplotype (Paris, ancient region that al., genus et in (Chomicki the watermelon domestication dessert and origin wild of the reported center Authors a (2011)], as al. Africa 2020). a et north-east as Minsart indicates highlighted evidence in be Archaeological 1 should Fig. coat, and soft (2015) yellow from with of al. distinct seeds et genepool Achigan-Dako smaller the in exhibits to 9f that contributor yielded melon [Fig. author type egusi first cultivated ‘neri’ the the another by of mission Africa West collecting in of region-wide population A wild cultivation. no from escaped certainly individuals lanatus . .lanatus C. irlu mucosospermus Citrullus Citrullus fDru nnrhesenArc srvae yRne ta.(09 ntescn ad calls hand, second the on (2019) al. et Renner by revealed as Africa north-eastern in Darfur of nadto,tehptei htwtreo sfo ot-atr fiade o explain not does Africa north-eastern from is watermelon that hypothesis the addition, In . .mucosospermus C. var. eei eore n rpEouin 62 Evolution, Crop and Resources Genetic .mucosospermus C. oprtv hlgorpi n ouaingntcaaye ftretoia pioneer tropical three of analyses genetic population and phylogeographic Comparative nldssvnseista a rgnt rmdffrn at ftewrd codn to according world, the of parts different from originate may that species seven includes cordophanus .mucosospermus C. Citrullus thsbe aeystudied. rarely been has it , .mucosospermus C. asetwieflse utvr Rne ta. 09.Hwvr within However, 2019). al., et (Renner cultivar) white-fleshed sweet (a ean h lss eaieseisto species relative closest the remains oetctd hog htmcaim was mechanisms what Through domesticated? stergo.Wieti eitp ( type neri this While region. the is Citrullus nWs fia(cia-aoe l,21)tog,tepresence the though, 2015) al., et (Achigan-Dako Africa West in .colocynthis C. pce.I fiatefcssol eo ohws and west both on be should focus the Africa In species. 8 hrstesm altp ihdsetwatermelon, dessert with haplotype same the shares 8,1159-1179. (8), n lsrt aemlnadeuimelon egusi and watermelon to closer and .lanatus C. .lanatus C. Martin-Luther-Universitat .lanatus C. .lanatus C. h rsneo an of presence The . .lanatus C. smorphologically is ) Citrullus .amarus C. n subsp. and i indeed did .lanatus C. pedto spread p.and spp. Posted on Authorea 17 Aug 2020 — The copyright holder is the author/funder. 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(4), Citrullus eei eore n rpEouin 58 Evolution, Crop and Resources Genetic p.acsinueu o nacn iesso etrssac in resistance pest or diseases enhancing for useful accession Spp. 207-217. , n-le uiueitorsingntqeapri d’individus partir a genetique introgression une subi ont-elles ) 353–358. , oubaUiest Press University Columbia irlu lanatus Citrullus 11 mrcnJunlo ln cecs 4 Sciences, Plant of Journal American eei eore n rpEouin 48 Evolution, Crop and Resources Genetic 1,47-56. (1), 6) 427-440. (60), e ce uBG 4 BRG, du Actes Les 5,609-617. (5), adae nSuhr Africa. Southern in landraces ) 6,805-814. (6), ora fteAeia Society American the of Journal oeua hlgntc and Phylogenetics Molecular n.Hm Genet., Hum. Ann. 115-126. , irlu lanatus Citrullus 1448-1456. , Citrullus Journal (6), Posted on Authorea 17 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159769281.18634046 — This a preprint and has not been peer reviewed. Data may be preliminary. ho . a,B,&Zag .(09.Pyoegah n osraingntc fteendangered World old the the contributions of in Author plants genetics of conservation Domestication and (2000). Phylogeography M. UK. Hopf, & (2019). China. D., Northwest M. Zohary, Mongolia, Zhang, Inner from & (Asteraceae) B., mongolica Tugarinovia Pan, Y., Cucurbits Zhao, (1976). W. water- B. of Bemis, history 64-69. & the W., to T. contribution Whitaker, archaeobotanical . An primates. (2004). hominoid 13 archaeobotany, M. in Veen, gene der II melon, Van oxidase & rejection cytochrome K., the mitochondrial Wasylikowa, trees: the gene of intra/interspecific evolution using 144 neutrality the Genetics, of for tests neutrality Contingency of (1996). R. Liquidambar of A. structure Templeton, Phylogeographical models. distribution (2019). species Y. Zheng, and phylogeography & chloroplast J., 858. by Lai, X., revealed Ye, Hance P., formosana Huang, Turkish F., Lin, of R., characterization Sun, genetic The (2010). Y. 763-771. N. (5), Yalcin-Mendi, & Y., ( Aka-Kacar, watermelon N., Sari, I., ( Solmaz, watermelon of traits. Characterization morphological (2009). for N. Turkey hare Sari, and & tortoise I., the new Solmaz, chloroplast Angiosperm: a whole in of studies and Comparison phylogenetic Cucurbitales for (2007). regions Small. order III. noncoding L. choose the Randall & to in Schilling, sequences E. genome relationships Edward Lickey, B. Phylogenetic Edgar J., Shaw, (Cucurbitaceae). (2011b). family S. (Ed.), gourd S. the Kubitzki of Renner, K. classification & In H., Myrtaceae Cucurbitaceae Cucurbitales, Schaefer, Sapindales, genetics (2011a). 10, population S. molecular vol. S. for plants, Renner, program vascular integrated & an H., 3: Schaefer, version DnaSP analysis. G. evolution (1997). cultivating Chomicki, molecular were R. & and Egyptians Rozas, M., Kingdom & Hofreiter, New J., M., that Rozas, reveals Preick, tomb M., Pharaonic Nesbitt, a V., watermelon. from M. domesticated leaf name Silber, 3500-year-old the A., A O. conserve (2019). Perez-Escobar, to S., Proposal S. Renner, (2014). W. Greuter, 63 & G., unordered Chomicki, versus ( S., ordered with S. differentiation genetic Renner, testing and Measuring watermelon, (1996). phy- J. dessert allells. R. the sweet Petit, the & in O., of divergent Pons, emergence highly and are Origin 116 spacers (2015). botany, transcribed S. H. internal Paris, DNA Ribosomal ascomycete topathogenic (1992). K. O’Donnell, irlu lanatus Citrullus 4,941-942. (4), mrcnJunlo oay 94 Botany, of Journal American . irlu lanatus Citrullus eeis 144 Genetics, 133-148. , irlu lanatus Citrullus 1263–1270. , wtreo,Ccriaee,wt osre ye against type, conserved a with Cucurbitaceae), (watermelon, ) 4,213-217. (4), 1237-1245. , Tub)Mtu.&Nki(syn. Nakai & Matsum. (Thunb.) uaimsambucinum Fusarium bioRxiv cesosuigRP markers. RAPD using accessions ) eei eore n rpEouin 56 Evolution, Crop and Resources Genetic iifrais 2 Bioinformatics, 642785. , 3,275-288. (3), ( ibrlapulicaris Gibberella AO,60 TAXON, 12 1) 174-175. (15), imnsN ed,Eouino rppat 6 plants crop of Evolution (e.d), W N. Simmonds .vulgaris C. 1-7 eln Springer. Berlin: 112-174 . irlu lanatus Citrullus 1,122-138. (1), eei eore n rpEouin 57 Evolution, Crop and Resources Genetic ). ur ee. 22 Genet., Curr. Schrad.). xodUiest rs,Oxford, Press, University Oxford lSoe 14 one, PloS 2,173-188. (2), irlu lanatus Citrullus h aiisadgnr of genera and families The cesoscletdfrom collected accessions ) irlu battich Citrullus eeainhsoyand history Vegetation oodc lanata Momordica 213-220. , 2,e0211696. (2), oet,10 Forests, . nasof Annals . Taxon, (10), , Posted on Authorea 17 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159769281.18634046 — This a preprint and has not been peer reviewed. Data may be preliminary. oTxnHpoyenme ceso ubrOii oreo olcinNC ubrfor number 27 NBCI 26 collection of Source 25 24 23 22 21 20 19 18 17 16 Origin 15 number Accession 14 number Haplotype 13 12 11 10 9 8 7 6 5 4 3 2 1 Taxon No 1: Table 5 Fig. 4 Fig. Africa 3 distribution: Fig. geographical indicate (blue). colors 2 America the North Fig. and and (red), S2), Europe Table (yellow); (see Asia Clv: numbers (green), ID letters. haplotype three arbitrary or two with subsp. abbreviated are names Taxon 1 Fig. S1. Table in provided captions are Figures numbers accession Genbank read NCBI authors All sequences: DNA analysis. and E.G.AD. collection E.G.AD., data accessibility materials. the data. Data plant analysed supervised manuscript. H.D. F.R.B assembled final and E.G.AD manuscript. the E.G.AD approved the and wrote work. research. I.H. laboratory the and and designed H.D., field conducted and experiments, planned performed F.R.B. and E.G.AD irlu amarus Citrullus amarus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus mucosospermus Citrullus lanatus Citrullus mucosospermus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus mucosospermus Citrullus mucosospermus Citrullus mucosospermus Citrullus mucosospermus Citrullus lanatus Citrullus mucosospermus Citrullus mucosospermus Citrullus mucosospermus Citrullus mucosospermus Citrullus lanatus Citrullus mucosospermus Citrullus mucosospermus Citrullus mucosospermus Citrullus lanatus Citrullus lanatus itiuinadfeunisof frequencies and Distribution : of frequencies and Distribution . of frequencies and Distribution . C ewr f38 of network TCS . itiuinadfeunisof frequencies and Distribution : itof List Cm: ; Citrullus .mucosospermus C. var. var. var. var. var. var. var. var. var. var. var. cesos hi egahcloii,adacsinnumbers. accession and origin, geographical their accessions, lanatus lanatus lanatus lanatus lanatus lanatus lanatus lanatus lanatus lanatus lanatus Citrullus p.hpoye.Crl iei rprinlt altp frequency. haplotype to proportional is size Circle haplotypes. spp. Citrullus Citrullus Citrullus Cam: ; Citrullus RF186Rsi SAK739 KX773744 KX773742 KX773738 KX773739 KX773736 KX773595 KX773593 KX773589 KX773590 USDA KX773587 KX773730 USDA KX773729 KX773728 KX773725 KX773727 USDA KX773724 USDA KX773723 USDA KX773722 KX773720 KX773581 Africa Russia South KX773719 KX773580 KX773718 KX773579 KX773576 KX773578 Prospection KX773575 Africa 15896 South GRIF KX773574 596662 Nigeria Prospection PI KX773573 Prospection Prospection Ghana KX773571 271778 PI USDA 249010 KX773570 PI USDA KX773569 USDA 186975 PI USDA 4 3 USDA Benin USDA Benin USDA 1 Benin 1 045 Benin SN 1068 008 MAT Nigeria 977 6 007 MAT Nigeria 975 001 BSN Nigeria 849 Nigeria 560018 PI 560013 Nigeria PI 560010 PI Nigeria 1 560008 Nigeria PI 1 1 560000 PI 1 559994 PI 559993 PI 1 1 1 1 9 26 1 RF590IdaUD X752KX773741 KX773740 KX773737 KX773735 KX773734 KX773592 KX773733 KX773732 KX773591 KX773731 KX773588 USDA KX773586 KX773726 USDA KX773585 KX773584 KX773583 USDA KX773582 USDA KX773721 USDA USDA USDA KX773577 KX773717 India USDA India 55990 China GRIF KX773572 55960 USDA GRIF China China 192937 PI KX773568 China India 17330 GRIF USDA 17310 GRIF China 1 17300 GRIF 10 14199 GRIF 12336 USDA GRIF Nigeria 1 1 2 560024 PI 1 Nigeria 1 19 Nigeria 560002 PI 1 494527 PI 17 9 .amarus C. p.hpoye nNrhAmerica. North in haplotypes spp. Asia. in haplotypes spp. Africa. in haplotypes spp. p.hpoye nEurope. in haplotypes spp. 13 n Cco: and ; .lanatus C. .colocynthis C. subsp. vulgaris h ubr are numbers The . Cll: ; .lanatus C. ndh F- rpl 2NC ubrfor number NBCI 32 trn T-L Posted on Authorea 17 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159769281.18634046 — This a preprint and has not been peer reviewed. Data may be preliminary. 77 76 75 74 73 72 71 70 69 68 67 66 65 64 63 62 61 60 59 68 57 56 55 for number 54 NBCI 53 collection of Source 52 51 50 49 48 47 46 45 44 43 Origin 42 number Accession 41 number Haplotype 40 39 38 37 36 35 34 33 32 31 30 29 28 Taxon No irlu lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus amarus Citrullus amarus Citrullus amarus Citrullus amarus Citrullus amarus Citrullus amarus Citrullus amarus Citrullus amarus Citrullus amarus Citrullus amarus Citrullus amarus Citrullus amarus Citrullus amarus Citrullus amarus Citrullus amarus Citrullus amarus Citrullus amarus Citrullus amarus Citrullus amarus Citrullus amarus Citrullus subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. vulgaris vulgaris vulgaris vulgaris vulgaris vulgaris vulgaris vulgaris vulgaris vulgaris vulgaris vulgaris vulgaris vulgaris vulgaris vulgaris Vulgaris Vulgaris vulgaris vulgaris vulgaris vulgaris vulgaris vulgaris vulgaris vulgaris vulgaris vulgaris vulgaris vulgaris I 4 S P X769KX773798 KX773797 KX773796 KX773795 KX773793 KX773794 KX773792 KX773649 KX773791 KX773648 KX773790 KX773788 KX773647 KX773646 KX773787 KX773786 KX773644 IPK KX773784 KX773645 KX773785 KX773643 KX773783 IPK KX773780 KX773642 KX773779 IPK KX773641 IPK KX773778 KX773639 KX773777 KX773638 KX773776 KX773775 IPK KX773637 KX773774 IPK IPK KX773635 IPK KX773636 KX773773 KX773772 KX773634 KX773770 IPK KX773631 KX773769 KX773630 IPK USA KX773629 IPK USA IPK KX773768 Japan KX773628 KX773627 KX773626 USA KX773766 Korea KX773767 North KX773625 IPK IPK 242 KX773765 CIT IPK Russia 239 IPK KX773764 CIT KX773624 Georgia IPK KX773623 237 KX773621 CIT Georgia IPK 235 KX773620 CIT Georgia 167 KX773763 KX773761 IPK CIT KX773762 Bulgaria IPK 164 IPK CIT KX773619 Bulgaria KX773760 160 IPK CIT Bulgaria 158 KX773617 CIT KX773618 1 Yugoslavia 156 IPK KX773616 CIT IPK Mongolia IPK 1 143 KX773615 CIT Armenia KX773759 IPK 2 142 CIT Ukraine 1 KX773758 135 CIT Russia KX773757 2 KX773755 IPK KX773614 KX773612 130 KX773613 Russia KX773756 1 CIT 128 CIT Ukraine 1 IPK IPK 126 KX773611 CIT Russia KX773754 1 IPK 112 CIT USA KX773753 1 IPK 109 China CIT KX773752 1 107 Hungary CIT KX773751 KX773610 1 105 Greece CIT KX773750 IPK 1 IPK KX773609 103 IPK Italy CIT KX773749 KX773608 18 KX773606 102 KX773746 Italy CIT KX773748 KX773607 IPK 1 99 CIT Croatia 2 97 Yugoslavia CIT KX773745 KX773605 1 86 Ukraine USDA CIT KX773604 1 69 CIT France KX773603 1 USDA 67 USDA CIT Yemen USDA KX773602 1 Africa USDA 60 South KX773601 1 CIT 44 CIT KX773747 Zimbabwe KX773600 1 31 KX773597 USDA CIT Russia KX773599 1 207 USDA CIT 1 Ukraine 313 USDA CIT KX773596 1 Africa 310 South USDA CIT 1 152 Africa USDA CIT South 1 Africa 139 South USDA CIT 18 USDA Africa South USDA 101 1 CIT 596676 PI Egypt KX773598 1 Africa South USDA 596671 2 PI Africa 596669 South PI 2 Africa 596659 South PI 3 Africa South 525083 3 PI Africa 271779 South PI 4 Africa 270563 South USDA PI 15 Africa 255137 South Congo PI of 3 Republic 244019 Democratic PI 14 244018 India PI 8 299379 PI 8 Russia 299378 PI 32 189225 PI 6 179881 PI 4 15897 GRIF 4 3 3 4 3 4 6 4 14 ndh F- rpl 2NC ubrfor number NBCI 32 trn T-L Posted on Authorea 17 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159769281.18634046 — This a preprint and has not been peer reviewed. Data may be preliminary. 127 126 125 124 123 122 121 120 119 118 117 116 115 114 113 112 111 110 109 108 107 106 105 for number 104 NBCI 103 collection of Source 102 101 100 99 98 97 96 95 94 93 Origin 92 number Accession 91 number Haplotype 90 89 88 87 86 85 84 83 82 81 80 79 78 Taxon No irlu lanatus Citrullus colocynthis Citrullus colocynthis Citrullus colocynthis Citrullus colocynthis Citrullus colocynthis Citrullus colocynthis Citrullus colocynthis Citrullus colocynthis Citrullus colocynthis Citrullus colocynthis Citrullus colocynthis Citrullus colocynthis Citrullus colocynthis Citrullus colocynthis Citrullus colocynthis Citrullus colocynthis Citrullus colocynthis Citrullus colocynthis Citrullus colocynthis Citrullus colocynthis Citrullus colocynthis Citrullus lanatus Citrulus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus lanatus Citrullus subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. subsp. lanatus vulgaris vulgaris vulgaris vulgaris vulgaris vulgaris vulgaris vulgaris vulgaris vulgaris vulgaris 1P 327Pksa SAK730 KX773856 KX773855 KX773853 KX773854 KX773857 KX773852 KX773851 KX773850 KX773849 KX773848 KX773847 KX773707 KX773706 KX773704 KX773705 KX773708 KX773703 KX773846 KX773702 KX773843 KX773845 KX773844 KX773701 KX773842 KX773841 KX773700 KX773840 Prospection USDA USDA KX773699 KX773698 USDA USDA KX773839 USDA USDA KX773838 USDA KX773828 USDA KX773697 KX773837 KX773694 KX773827 USDA KX773696 KX773695 KX773693 KX773836 KX773692 USDA KX773829 Faso Burkina KX773691 KX773826 KX773825 KX773824 Pakistan KX773690 KX773823 Egypt USDA KX773822 Cyprus 60 AE KX773689 824 USDA USDA Iran KX773821 IPK IPK KX773820 537277 KX773679 Iran PI IPK Iran KX773688 525082 KX773678 IPK PI KX773819 KX773687 Iran 432337 KX773680 PI Iran 386026 KX773818 KX773677 PI IPK KX773676 Iran 386024 KX773675 PI 386021 Iran PI KX773817 KX773674 KX773815 IPK 386018 Afghanistan PI KX773673 KX773814 2 386016 IPK Afghanistan PI 31 KX773672 KX773811 IPK 386015 IPK KX773812 PI KX773671 Ethiopia IPK 34 Namibia 386014 IPK PI 37 Cyprus KX773670 346082 IPK PI IPK 29 Egypt 220778 IPK KX773810 PI KX773669 27 India IPK 195927 7 PI Morocco IPK KX773809 307 KX773668 KX773666 Verde 5 CIT KX773808 Cape IPK 281 KX773665 5 CIT Turkmenistan IPK 199 Island 5 CIT KX773807 KX773803 KX773662 Canary KX773663 KX773802 192 KX773806 IPK Africa 5 CIT South KX773805 KX773801 190 USA 7 CIT KX773800 IPK 166 USA 7 CIT KX773799 KX773661 154 Canada 30 CIT IPK IPK 150 13 CIT USA IPK KX773660 309 USA 38 CIT KX773659 278 CIT IPK USA IPK 12 273 CIT USA 12 KX773658 KX773654 271 CIT KX773653 Israel KX773657 35 KX773656 Prospection KX773652 270 USA IPK 33 Prospection CIT KX773651 264 CIT China 28 KX773650 260 CIT Tunisia Prospection 36 234 CIT Prospection Tajikistan 16 Prospection 230 CIT Georgia 1 226 CIT IPK Ukraine 1 IPK 206 CIT Mongolia 1 IPK 203 CIT IPK 21 Italy 200 Korea CIT IPK North 2 195 CIT 1 193 CIT 1 Mali 182 Mali CIT 1 Ghana 175 1 CIT 168 CIT Benin 2 032 SE 1005 Mali Portugal 1 BAX1 846 Turkey 1 995 NIA 07 1 567 Japan NIA 06 1 224 USA NIA 06 2 USA 306 CIT 24 303 CIT 1 253 1 CIT 259 1 CIT 244 2 CIT 2 1 1 1 22 11 11 15 ndh F- rpl 2NC ubrfor number NBCI 32 trn T-L Posted on Authorea 17 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159769281.18634046 — This a preprint and has not been peer reviewed. Data may be preliminary. rpl ndh trn rpl ndh for number NBCI collection of Source trn regions CpDNA Origin number Accession number Haplotype the on based statistics Genetic 2: Table 135 134 133 132 131 130 129 128 Taxon No 32 32 - & T-L T-L F- F- irlu aau vulgaris lanatus Citrullus vulgaris lanatus Citrullus lantus Citrullus lantus Citrullus lantus Citrullus colocynthis Citrullus lanatus Citrullus lanatus Citrullus cynthis colo- C. amarus C. mus cososper- mu- C. lanatus Citrullus groups Taxonomic amarus C. mus cososper- mu- C. lanatus C. cynthis colo- C. amarus C. mus cososper- mu- C. lanatus C. v ei10 I 1 hn rseto X774KX773863 KX773862 KX773861 KX773714 KX773713 KX773712 Prospection Prospection Prospection Ghana Ghana Ghana 111 NIA 06 103 NIA 06 095 NIA 06 1 20 1 neri cv. neri cv. neri cv. subsp. subsp. 2948- 950- 22 950- 22 951- 16 78 accessions of Number 21602- 1601- 22 1601- 16 650- 78 651- 22 651- 22 650- 16 78 vulgaris vulgaris ua ay2GI 59 S SAK731 KX773865 KX773864 KX773716 KX773715 USDA USDA USA Canada 15898 GRIF 15895 GRIF 2 baby 2 sugar baby sugar 954 953 953 954 (bp) Length Total 1604 1604 1605 653 653 652 652 trn T-L, 592K 2 ukn aoPopcinK731 KX773860 KX773859 KX773858 KX773711 KX773710 KX773709 Prospection Prospection Prospection Faso Burkina Faso Burkina Faso Burkina 026 KU 962 031 AE 831 60 AE 825 25 2 23 ndh .400275.8 76.1 2 3 x 28 0 x 1 0 0.92 0 0.52 0 12 0.34 5 0.44 6 3 0 4 0 0 sites mative infor- Parsimony 205 . 04005876.2 x .8 4 8 x 0.8 0.81 0.025 10-4 0.53 x 0.2 8 x 7 0.56 5 x 10.5 2 0.80 12 x 1.9 0.71 0 x 0.4 11 0.25 0 6 0.24 1 3 2 8 0 0 16 F- rpl 2adtercmiainin combination their and 32 haplotypes of Number diversity Haplotypes 10 10 (Pi) sity diver- Nucleotide 10-4 10-4 10 10 10 10 Citrullus -4 -4 -4 -4 -4 -4

spp. .5576.0 75.9 5 4 2.65 0.09 (k) ence differ- cleotide nu- of ber num- Average .5676.3 76.8 6 76.8 1 0.45 76.3 0.68 0 5 0.125 0.027 .8676.2 76.2 6 0.78 2 0.125

events Indel A+T (%) ndh F- rpl 2NC ubrfor number NBCI 32 trn T-L Posted on Authorea 17 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159769281.18634046 — This a preprint and has not been peer reviewed. Data may be preliminary. DHpoyeSeisOii Frequency Origin Species T– T–TT-TGTGTAAACACAAA—ATTAGA 7 6 T– 5 T– 4 T- 3 T–TT-TGTGTAAACACAAA—ATTAGA- 2 1 Haplotype ID 3: Table (%) T + A events Indel population. given a within SNPs) or differences nucleotide of number Average 1990). Crease and (Lynch diversity Nucleotide populations. or regions different two diversity: Haplotype sites: Non-informative population. the in twice least sites Parsimony-informative regions CpDNA 9T- 19 T– T– 16 T–TT- 15 T 14 T 13 T–TT-TGTGTAAAC 12 T–TT-TGTGTAAACACAAA—11 T 10 T–TT-TGTGTAAACACAAA—ATTA 9 8 four the of collections 8T T–TT-TGTGTAAACACAAA—ATTA18 17 - – G G G G - T T - - - - - T –T - -TT-TGTGTAAACACAAA—ATTAGA- -TT-TGTGTAAACACAAA— T T-TGTGTAAACACAAA—ATTA T-TGTGTAAACACAAA—ATTA T-TGTGTAAAC T-TGTGTAAACACAAA—ATTA TT-TGTGTAAACACAAA—ATTAGA- TT-TGTGTAAACACAAA—ATTAGA - A T A altp oe o h combined the for codes Haplotype A TGTGT – A cynthis colo- C. groups Taxonomic TGTG + otn ntegnmcregion. genomic the in content A+T : TGTAAACACAAA—ATTA TGTGTAAACACAAA—ATTA TGTG h ubro netosdltosi h eoi region. genomic the in insertions/deletions of number The : GT T GT AA Citrullus AA h vrg ubro uloiesbtttosprst ewe w sequences two between site per substitutions nucleotide of number average The : h rbblt httogvnsqecsfo w ieethpoye eogto belong haplotypes different two from sequences given two that probability The 21599- 22 accessions of Number AA A oyopi ie htaeuiu ntepplto snltnsites). (singleton population the in unique are that sites Polymorphic - A - CAAA—ATTA ACAAA- CAAA—ATTAGA- A ACAAA- ACAAA- pce nti study. this in species oyopi ie ihamnmmo w lee htaeec rsn at present each are that alleles two of minimum a with sites Polymorphic : T 1605 (bp) Length Total - T -A TTAGA- T - -A TTAGA- - -A TA TCC - T TA TCC TC T - TCC T A- TA T A- TC A- T trn A- - h vrg ubro uloiedffrne ete Indels (either differences nucleotide of number average The : C T A- C 609 95x 19.5 0.96 16 7 sites mative infor- Parsimony - A- - and T-L 17 .mcssems;C amarus C. ; mucosospermus C. colocynthis C. amarus C. amarus amarus C. C. ; lanatus mucosospermus C. C. ; lanatus C. .lanatus C. amarus C. colocynthis C. lanatus C. lanatus C. mucosospermus C. colocynthis C. .lanatus C. amarus C. colocynthis C. amarus C. .lanatus C. .mucosospermus C. ndh haplotypes of Number F- rpl 2clrpatrgosfrteglobal the for regions chloroplast 32 diversity Haplotypes 10-4 (Pi) sity diver- Nucleotide fia;Asia ; Africa Eastern Near Asia ; Africa Africa America Southern Europe, Asia, ; America Africa Europe, Asia, ; Africa Asia Africa Southern Africa Northern America Asia ; Europe West-Africa Eastern Near Europe Europe Africa Southern South-Africa otenAfrica Southern Africa

(k) ence differ- cleotide nu- of ber num- Average .01 76.1 12 3.10

events Indel 3 4 7 7 17 60

1 1 2 2 2 2 3

1 1 1 2

1

1

A+T (%) Posted on Authorea 17 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159769281.18634046 — This a preprint and has not been peer reviewed. Data may be preliminary. DHpoyeSeisOii Frequency h (2011). al. et Guicking from adapted and Origin (1996) Petit and Pons to according haplotypes, cpDNA 4: Table Species T– T– 30 T– 29 T–TT 28 T– 27 T- 26 T 25 T- 24 T- 23 T–TT-TGTGTAAACA 22 T–TT-TGTGTAAACACAA21 20 Haplotype ID al :Piws eei ieetainbtencniet a,btenArcnrgos(b) regions African between (c) (a), continents regions between Asian differentiation between genetic and Pairwise 5: Table N V V G h Europe). and Asia, T- 31 3T T– 33 32 8T T– 38 T– 37 T 36 T– 35 34 eei aaeesVleSadr error Standard Value (h diversity gene within-population mean Expected parameters Genetic bevdma ihnpplto eedvriy(s .6 0.1878 0.668 (Nst) (V differentiation haplotypes genetic among of similarities coefficient for Observed accounting diversity, gene (Vs) total diversity Observed gene within-population (Gst) mean differentiation Observed genetic of coefficient (h Expected diversity gene total Expected S T T S St St : : h vrg emtdvleo eedvriywti h orgorpia ein Arc,America, (Africa, regions geographical four the within diversity gene of value permuted average The : h emtdvleo eedvriyars l orgorpia regions. geographical four all across diversity gene of value permuted The : : h emtdvleo eei ieetainaogtefu egahclregions. geographical four the among differentiation genetic of value permuted The : h vrg bevdvleo eedvriywti h orgorpia regions. geographical four the within diversity gene of value observed average The h bevdvleo eedvriyars l orgorpia regions. geographical four all across diversity gene of value observed The h bevdvleo eei ieetainaogtefu egahclregions. geographical four the among differentiation genetic of value observed The GT G G G T T T T - - - – – –A –A –A - - - T T T TT-TGTGTAAACAC TT TT-TGTGTAAACACAAA—A TT -TGTGTAAACACAAA—ATTAGA- -TGTGTAA –A –A –A iest n ieetainsaitc o h four the for statistics differentiation and Diversity A A A TT-TGTGTAAACACAAA—ATTAGA- A T T T TGTGT TGTG TGTG A A T TGT T A-A A-A A-A TGTG A-A A-A TGTGTAAACACAAA—ATTAGA- TGTGTA TAA TAA TAA A GT GT TAA TAA TAA GT T G G GA GA AACACA AA AA CACAAA AA GA GA- G C GA - A A ACAAA– ACAAA ACACAAA- CAAA A ACAAA ACAAA- ACAAA– - ACAAA CAAA– ACAAA- AAA—ATTAGA- - T AA—ATTAGA- C .1 0.0320 0.917 ) - ATA AT —ATTAGA- A- ATA A ATA A T ATA -A T A A T -A A T A -A - - A -A A - - TA TAGA - - - TA TAGA- – TA - A - –CT TA A - TA TAGA- TA TA - T - TA T T TA T A- T T A- T A- A- A- C T A- A- T A- A- A- S 18 .3 0.0671 0.737 ) .mucosospermus C. colocynthis C. lanatus C. lanatus C. lanatus C. lanatus C. .colocynthis C. colocynthis C. colocynthis C. lanatus C. .colocynthis C. .colocynthis C. .amarus C. .colocynthis C. .colocynthis C. colocynthis C. colocynthis C. colocynthis C. colocynthis C. Citrullus

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0.374 0.196 Africa Africa Europe Africa Asia Africa Asia Asia Asia America Africa Asia Africa Africa Europe Europe Europe Africa Africa 0.1274 0.0812 1 1 1 1 1 1

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1 1 1 1 1 Posted on Authorea 17 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159769281.18634046 — This a preprint and has not been peer reviewed. Data may be preliminary. saAeia07 .734 0.014 0.0079 = 0.038 Chi2 0.023 0.035 3.43 2.12 0.006 4.41 2.92 3.84 0.77 0.57 4.78 0.73 0.81 0.76 1993) (Hudson, 0.77 0.57 0.85 regions 0.73 African between 0.81 0.76 ferentiation America America dif- genetic Europe 0.85 Pairwise America 5-b: Europe Europe Asia Asia Asia Africa Africa Africa 1 Region 1993) (Hudson, continents between ferentiation dif- genetic Pairwise 5-a: etArc ot-fia07 .237 .2Ci 84.02 = Chi2 0.05 0.043 0.12 9.34 9.02 3.79 1993) 3.88 (Hudson, 3.14 regions Asian between 1.92 ferentiation 0.85 dif- genetic 0.72 Pairwise North-Africa South-Africa 5-c: South-Africa 0.73 West-Africa South-Africa West-Africa 1 Region ot-saNrhAi .741 .10.07 65.75 0.08 = Chi2 0.014 0.09 0.06 5.11 6.64 0.04 6.20 2.37 4.73 4.19 4.97 6.30 6.20 1.30 2.65 0.77 0.78 3.50 0.89 North-Asia 0.64 North-Asia 0.76 South-Asia South-Asia North-Asia West-Asia South-Asia 0.77 West-Asia East-Asia East-Asia West-Asia East-Asia 1 Region ein2H sKyGtChi-square Gst 1993) (Hudson, regions African Kxy between ferentiation dif- genetic Pairwise 5-b: Ks Hs 2 Region 1993) (Hudson, continents between ferentiation dif- genetic Pairwise 5-a: ein2H sKyGtChi-square Gst 1993) Kxy (Hudson, regions Asian between ferentiation Ks dif- genetic Pairwise 5-c: Hs 2 Region ein2H sKyGtChi-square Gst Kxy Ks Hs 2 Region 1993) (Hudson, regions African between ferentiation dif- genetic Pairwise 5-b: 1993) (Hudson, continents between ferentiation dif- genetic Pairwise 5-a: 1993) (Hudson, regions Asian between ferentiation dif- genetic Pairwise 5-c: 19 1993) (Hudson, regions African between ferentiation dif- genetic Pairwise 5-b: 1993) (Hudson, continents between ferentiation dif- genetic Pairwise 5-a: 1993) (Hudson, regions Asian between ferentiation dif- genetic Pairwise 5-c: 1993) (Hudson, regions African between ferentiation dif- genetic Pairwise 5-b: 1993) (Hudson, continents between ferentiation dif- genetic Pairwise 5-a: 1993) (Hudson, regions Asian between ferentiation dif- genetic Pairwise 5-c: 1993) (Hudson, regions African between ferentiation dif- genetic Pairwise 5-b: 1993) (Hudson, continents between ferentiation dif- genetic Pairwise 5-a: 1993) (Hudson, regions Asian between ferentiation dif- genetic Pairwise 5-c: -au 0.05 = P-value 135.067 1993) (Hudson, regions African between ferentiation dif- genetic Pairwise 5-b: 1993) (Hudson, continents between ferentiation dif- genetic Pairwise 5-a: 0.0001 = P-value 1993) (Hudson, regions Asian between ferentiation dif- genetic Pairwise 5-c: 0.0047 = P-value Posted on Authorea 17 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159769281.18634046 — This a preprint and has not been peer reviewed. Data may be preliminary. rirr altp Dnmes(e al 2,adteclr niaegorpia itiuin Africa distribution: geographical indicate (blue). colors America the North and and (red), S2), Europe Table (yellow); (see Asia Clv: numbers (green), ID letters. haplotype three arbitrary or two with subsp. abbreviated are names Taxon 1 Fig. GST Kxy: Ks: Hs h enwti-otnn eediversity. gene within-continent mean The : egtdaeaeo h ubro ieecsbtensqecsfo continents from sequences between differences of number the of average weighted A h offiin fgntcdffrnito ewe continents between differentiation genetic of coefficient The : h vrg ubro ieecsbtentosmls eadeso hi provenance. their of regardless samples, two between differences of number average The lanatus C ewr f38 of network TCS . Cm: ; .mucosospermus C. Citrullus p.hpoye.Crl iei rprinlt altp frequency. haplotype to proportional is size Circle haplotypes. spp. Cam: ; .amarus C. 20 n Cco: and ; .lanatus C. .colocynthis C. subsp. vulgaris h ubr are numbers The . i Cll: ; and j .lanatus C. . Posted on Authorea 17 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159769281.18634046 — This a preprint and has not been peer reviewed. Data may be preliminary. i.3 Fig. 2 Fig. itiuinadfeunisof frequencies and Distribution . of frequencies and Distribution . Citrullus Citrullus p.hpoye nAsia. in haplotypes spp. Africa. in haplotypes spp. 21 Posted on Authorea 17 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159769281.18634046 — This a preprint and has not been peer reviewed. Data may be preliminary. i.5 Fig. 4 Fig. itiuinadfeunisof frequencies and Distribution : itiuinadfeunisof frequencies and Distribution : Citrullus Citrullus p.hpoye nNrhAmerica. North in haplotypes spp. p.hpoye nEurope. in haplotypes spp. 22