Plant-parasitic nematodes parasitizing saffron in Morocco: Structuring drivers and biological risk identification Hinde Benjlil, Khalid Elkassemi, Mohamed Aït Hamza, Thierry Mateille, James Nicholas Furze, Khalil Cherifi, El Hassan Mayad, Zahra Ferji

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Hinde Benjlil, Khalid Elkassemi, Mohamed Aït Hamza, Thierry Mateille, James Nicholas Furze, et al.. Plant-parasitic nematodes parasitizing saffron in Morocco: Structuring drivers and biological risk identification. Applied Soil Ecology, Elsevier, 2020, 147, ￿10.1016/j.apsoil.2019.103362￿. ￿hal-02623449￿

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Distributed under a Creative Commons Attribution - NonCommercial| 4.0 International License Version postprint ae ihlsFurze Nicholas James pc costegoe( globe the across spice eevd1 ac 09 eevdi eie om1 etme 09 cetd1 etme 2019 September 19 Accepted 2019; September 18 form revised in Received 2019; March 15 Received https://doi.org/10.1016/j.apsoil.2019.103362 Introduction 1. factors Abiotic ecology Soil sativus Crocus nematodes Phytoparasitic Keywords: INFO ARTICLE c b a Benjlil Hinde identi risk biological and drivers sa parasitizing nematodes Plant-parasitic ihteipeetto fgorpia niainpoessi the in processes indication geographical of ‘ ( implementation quality the high and With its nationally reputation of great because a gained internationally recently has and centuries, for ubro rwr ewe 00ad21 n h rdcinof production the and 2014 and 2010 between the in growers increase of an encouraged number of initiative designation government protected This a (PDO). became origin region the and registered was liouine 0929-1393/ ©2019 ElsevierB.V.Allrightsreserved. re ooc Plan’ Morocco Green ntttArnmqee ééiar asnI,Cmu d Campus II, Hassan Vétérinaire et Agronomique Institut nvriéInZh,FclédsSine d Sciences des Faculté Zohr, Ibn Université BP R,CRD NA otele uAr,Ui.Mnplir otele,France Montpellier, Montpellier, Univ. SupAgro, Montpellier INRA, CIRAD, IRD, CBGP, Please cite thisarticleas: HindeBenjlil,et al.,AppliedSoi l Ecology, https://doi.org/10.1016/j.apsoil.2019.103362 ⁎ orsodn uhra:Uiest b or aut e cecsd Sciences des Faculté Zohr, Ibn Université at: author Corresponding -aladdress: E-mail Sa ff Plant-parasitic nematodes parasitizing saffroninMorocco: Structuring driversand biological o ( ron Benjlil, H., Elkassemi,K., AïtHamza, M.,Mateille,T., Furze,Cherifi, Mayad,Ferji (2020). risk identification. Applied Soil Ecology, 147. ,DOI :10.1016/j.apsoil.2019.103362 rcssativus Crocus [email protected] a ,c hldElkassemi Khalid , Ka Pa ao et,i 00 sa 2010, in Vert), Maroc (Plan fi . sse stems otyadluxurious and costly most the as seen is L.) ta. 2006 al., et a hllCherifi Khalil , ’ gdr aoaor BR,B 16 00 gdr Morocco Agadir, 80000 8106, BP LBVRN, Laboratoire Agadir, .I a engoni Morocco in grown been has It ). Comment citer cedocument: ora homepage: journal ’ gdr éatmn ePoeto e lne,B 8S 00 gdr Morocco Agadir, 80000 18/S, BP Plantes, des Protection de Département Agadir, c aeadCnrl,2009 Cantrell, and Lage oae ï Hamza Aït Mohamed , ..TloieadTzah ein r h otpoutv ra fti aubenurcuia.Det its to Due neutraceutical. valuable this of areas productive most the economically are sa an pro colorant/medicinal/spice regions are the metabolic Taznakht for They country and productive pathogens. Taliouine highly plant a L.). be all to signi considered of is causing destructive Morocco pathogens, soil most of the group are important (PPN) nematodes Plant-parasitic ABSTRACT 9 8ad3%i h rao aiun,wiei anktte ersn 5 9 3 n 8 respectively. 28% and 33, 69, 95, represent they Taznakht in while di Taliouine, at of diversity area of assessment the the in Regarding 36% and 48 49, Helicotylenchus huhmxmmtmeaueddnthv nimpact. an have not did taxa, PPN temperature and Paratylenchus temperature minimum maximum rainfall, humidity, Soil between though types. links soil indicated infl and (MBPLS) factor analyses communities Multivariate nematode major between the relationship is substantial texture a revealed analyses Co-inertia ( The diversity Shannon the whereas genera, 12 communes to rural belonging genera 11 parameters. PPN from abiotic Fifteen collected regions. identi and samples Taznakht biotic and were soil with Taliouine 163 in assemblages families in gradients Taznakht their altitudinal characterized and by between Taliouine were of characterized links soils communities in of nematode communities PPN understanding Herein, of facilitate diversity the to assess to regions aims work This (PPN). nematodes eaoecnrligapoce a eapidadpeettv esrssol ecniee tnursery at considered be should measures preventative and applied be and may approaches controlling nematode Tylenchorynchus a M ï Hamza). Aït (M. lHsa Mayad Hassan El , fi l level. eld otnslssaalbeat available lists Contents fication ple olEcology Soil Applied ff Applied SoilEcologyxxx(xxxx)xxxx o rmTa- from ron

eerltdt h uiiyadsl olta eeoe nTaliouine. in developed that soil silt and humidity the to related were fi oetaesroslmtn atr nsa in factors limiting serious potentiate , e n rwhfrs sa forms, growth and le, and ’ gdr aoaor BR,B 16 00 gdr Morocco. Agadir, 80000 8106, BP LBVRN, Laboratoire Agadir, Dorylaimus fi www.elsevier.com/locate/apsoil d mn hc h orgenera four the which among ed, ff ). o nMrco Structuring Morocco: in ron ecn h rsneadteaudneo osdrbeprino genera. of portion considerable a of abundance the and presence the uencing a a,b sa rpig notntl,itnepatcsices h rbblt of probability the increase sa practices intense Unfortunately, cropping. growers and parasites deleterious most the ( considered are which (PPNs), nematodes intensi through production crop boost to tempt sa Currently 2010 ( producers non-PDO for 34% reached ar Ferji Zahra , eemr rvln nriylctosadca ol fTzah.Suitable Taznakht. of soils clay and locations rainy in prevalent more were baiadBkoe 2017 Bakooie, and Abtahi ff ff , * o D a mrvdb 6% uigti eid sa period, this During 360%. by improved has PDO ron o en ucpil osi iarsos nldn ln parasitic plant including bioagressors, soil to susceptible being ron ScienceDirect fl her Mateille Thierry , .Teeprstcognssijc ertoswt tltadfeed and stylet a with secretions inject organisms parasitic These ). wrpouto,addces h omyed( yield corm the decrease and production, ower H ff ’ ne aisfo . o15adteEens ( Evenness the and 1.5 to 0.9 from varies index ’) ff o sssetbet aypatdsae,icuigplant-parasitic including diseases, plant many to susceptible is ron agndcesdb ny1%frPOpoueswis it whilst producers PDO for 17% only by decreased margin rn ie,tegnsrcns ( richness genus the sites, erent ff o eevscnieal teto yfreswoat- who farmers by attention considerable receives ron fi atana aae fu o2%o ol rpproduction. crop world of 25% to up of damages annual cant c ff o rdcin hi rqece r epciey92, respectively are frequencies Their production. ron .PN nuepo eeomn fstigma of development poor induce PPNs ). Ditylenchus iyecu,Hlctlnhs Pratylenchus Helicotylenchus, Ditylenchus, b , , R ne agsfo o1 distinct 10 to 2 from ranges index ) peecods Pratylenchus Aphelenchoides, sróadMgei 2016). Megyesi, and Csurgó Aphelenchoides fi ainadmono- and cation ff E ne ed o1. to tends index ) o ( ron hae tal., et Ahrazem rcssativus Crocus , Tylenchus ff ron and and , Version postprint rdcn onre uha rn( ( Iran as such countries producing ( production crop 2014 world of 25% than 2011 more as valued been td ist vlaetedvriyo h P omnte associated communities PPN the of sa e diversity e with the to increase evaluate contribute to and aims will pests study This of regions. po- management Taznakht the sa cient and on assess risk Taliouine biological to as in tools considered crops be predictive could develop that genera and PPN tential generate this to Consequently, crop. aims quality the work high with of associated country nematodes producing phytoparasitic popular a is sa Morocco 2016). al., et bacteria, and viruses fungi, ( by roots assaults, infecting pathogenic other augments aiun n anktrgosi otenMrco( Morocco southern in regions Taznakht and Taliouine Sampling 2.1. methods and Materials 2. physico-chemical distribution. soil their of on impact parameters the climatic analyse and to and regions), Taznakht nclua otn ( content cellular on al. et Benjlil, H. n pi ( Spain and 2017) al., et Sheikh ff o,btteehsntbe n td drsigtedvriyof diversity the addressing study any been not has there but ron, h apigwscridotdrn ac n uy21 nthe in 2017 July and March during out carried was sampling The .Nmtlgclsuiswr netkni h edn sa ( leading year the per in USD undertaken billion were 157 studies to Nematological 100 ). between ranging ) Plant-parasitic nematodes parasitizing saffroninMorocco: Structuring driversand biological ff Benjlil, H., Elkassemi,K., AïtHamza, M.,Mateille,T., Furze,Cherifi, Mayad,Ferji (2020). o ntecr rdcn ein nMrco(aiun and (Taliouine Morocco in regions producing core the in ron risk identification. Applied Soil Ecology, 147. ,DOI :10.1016/j.apsoil.2019.103362 i.1. Fig. .PNe PPN 2014). al., et Nasiri a n rao sa of area and Map .Mroe,ro decay root Moreover, 2006). Hunt, and Decraemer adkaiadAvn,2013 Alvani, and Mahdikhani hae ta. 2010 al., et Ahrazem ff o rdcinprhcaei rsetdstso aiun n anktrgosi h ot fMorocco. of south the in regions Taznakht and Taliouine of sites prospected in hectare per production ron Comment citer cedocument: ff ff ciessanblt.This sustainability. ective cso il osshave losses yield on ects ; .Seven 1). Fig. Ranzenberger

io tal., et Nicol l tal., et Ali ,India ), ff ff ron ron ffi - 2 anali esta 5m.I ankt h otl vrg tem- average monthly the 40 Taznakht, and In annual 1 and mm. between 75 °C is 32 than perature and the less 10 Taliouine, re- is In between region. Taliouine rainfall are Taznakht the the temperatures in in Taznakht average m m 2856 1965 the monthly to to in 712 1100 from four and from and gion, ranges region altitude Taliouine The the region. in surveyed were sites n usest W) huaa(H,Sru S) zaun(Z)in (IZN) Iznaguen super (SI), Taliouine Siroua ( in (KH), (TA) Taznakht Khousama Taouyalte (WI), (ASK), Ouisselsate Askaouen and (SH), Tassous Hssaine (AM), Melloul Sidi Agadir (TS), (AS), Assaisse (ZA), Zagmouzen munes: ( mm 38 ldcnurnl.Tit om eernol eetdfo each from selected randomly were corms sa in Sa Thirty sample. bag concurrently. homogenized polyethylene one sub-samples pled a 30 obtain the in to mixed We order placed sample. in soil and 1kg-reference a depth form as- cm to 30 order 2cm- the a at with from pathway sampled auger Zig-Zag coming a this diameter sample in of collected composite one sub-samples, 30 Each a of ha. semblage by 163 represented to equal was is hectare which 10% of equivalent Sa ..Si analyses Soil 2.2. ff ff o orchard. ron oa f13si ape eecletdfo lvnrrlcom- rural eleven from collected were samples soil 163 of total A olpyioceia nlsswr are u nteSi Analysis Soil the in out carried were analyses physico-chemical Soil o rhrsae setmtdt oe 60h.W ape the sampled We ha. 1630 cover to estimated is area orchards ron fi i fsa of cie BS,2018). ABHSM, .Tesmldae oestnpreto h total the of percent ten covers area sampled The 1). Fig. ff o rdcini aiun n anktregions. Taznakht and Taliouine in production ron ◦ n h nulrifl sls than less is rainfall annual the and C Applied S ff oil Ecologyxxx(xxxx)xxxx o om eesam- were corms ron fi Version postprint opr ic oim agns,mgeim oasu,chloride spectrophotometry potassium, atomic using ( magnesium, measured were manganese, limestone and content, sodium, ( zinc, method copper, nitrogen 1990 Kjeldahl Clark, the and using by measured described method was the nitrogen using by estimated was o h rsetdstswr olce rmteMorocco the enumeration and characterization from extraction, Nematode Souss 2.4. collected of agency were basin hydraulic ( sites the ABHSM Massa of prospected department Meteorological the for collection data Climatic 2.3. ( measured also were ductivity oto fidvdasi ahgns h vnes(E=H evenness a the in genus; each genera in of individuals number = of (G (H portion richness diversity genus Shannon community); indices: following the diversity Taxonomic the 2.5. in counted were nematodes a the onto after, poured fi hours was small Forty-height suspension into The water. cut water. were (50 sample tap sieve each in paper from blended roots and and pieces corms of aliquot g 10 ln ot) oyamd,Tilnhd n yeciaodr were orders Tylenchida on and feed Triplonchida alternatively identi Dorylaimida, that roots), feeders (fungal plant Aphelenchida to ( belonging technique elutriation malized etto siain( estimation mentation mm 1 of a proportion (0 with determined: sieved were clays characteristics and soil dried following were The samples mesh. soil All Institute Morocco). Veterinary Agadir, and Agronomic the of Laboratory al. et Benjlil, H. rse stenme fidvdaspr10cm 100 per individuals of number the as pressed ( to cording ida n ovl,1978; Norvell, and Lindsay taeadternmeswr xrse n1 fcorms. of g 10 in expressed were numbers their and ltrate eaoe eeetatdfo 0 cm 300 from extracted were Nematodes data rainfall and humidity temperature, minimum and Maximum h aooi iest fPNcmuiiswr tde through studied were communities PPN of diversity taxonomic The a methods: Baermann by corms from extracted were Nematodes fi Plant-parasitic nematodes parasitizing saffroninMorocco: Structuring driversand biological dt eu ne troirsoe(0 magni (40X stereomicroscope a under genus to ed – Benjlil, H., Elkassemi,K., AïtHamza, M.,Mateille,T., Furze,Cherifi, Mayad,Ferji (2020). 200 i.2. Fig. risk identification. Applied Soil Ecology, 147. ,DOI :10.1016/j.apsoil.2019.103362 a n uln 1996 Mullin, and Mai μ )adcas (200 coarse and m) μ BS,2018). ABHSM, .Cnet fasmltdpopou,clim iron, calcium, phosphorus, assimilated of Contents ). eh ( mesh) M oiac iga ftenmtd eeaidenti genera nematode the of diagram Dominance egsadOds 1997 Oades, and Hedges on ta. 2010 al., et Coyne ise l,1991 al., et Sims ihrs 1954). Richards, otnrn,1960 Oostenbrink, – .PNpplto eeswr ex- were levels population PPN ). ’ 2000 =- ∑ μ pi* Comment citer cedocument: )snsacrigt sedi- to according sands m) 3 ln lcdit lt with plate a into placed ) fsi yuigtenor- the using by soil of i hr ii h pro- the is pi where pi) .p (H pH ). .Teogncmatter organic The ). 3 .Total 1960). Allison, ffehsoil. fresh of ; S,2007 ISO, “ asnII Hassan 2

)adcon- and O) fi ’ ain ac- cation) / ln Barbano .PPNs ). )that G) fi ” di l rsetdstsadcdsfrnmtd eeaaegvnin given are genera nematode for codes and sites prospected all in ed (IAV, 3 n9%o the of 90% in peecu,Dtlnhs Tylenchus Ditylenchus, Aphelenchus, h 5gnr,fu eeacudb osdrda eiu hetfor threat serious a as considered be Among could frequencies. genera sa low four with genera, Taliouine 15 the in recorded exclusively were eetdi sa in soil detected in nematodes parasitic plant of assessment Diversity 3.1.1. communities nematode plant-parasitic of Diversity 3.1. Results 3. 3.2.4. version R variables. and soil rainfall and (humidity, temperature) variables (MBPLS, climatic with Squares taxa between Least interaction Partial MultiBlock ( (Ade4) method, mbpls multivariate 1 + rqece eersetvl 2 9 8ad3%i h rao Ta- of area the in 36% and 48 49, liouine, 92, respectively were Frequencies iee ob aeyse F<5)ad2%wr ismntdand disseminated were 20% and 5%) < (F be seen to rarely observed (F were be location genera to prospected the sidered the of over 40% spread (F), extensively frequency ( and diagrams (A) dance dominance the to According osuyterlto ewe eeaadsi hsc-hmclfactors physico-chemical soil ( and packages genera applied ADE4 using between was relation the genera study detected to of diagram ( dominance A respectively. vrg muto ahgnsi l h ape)o ahgenus each of analysis Statistical samples) 2.6. the all in genus each (Log ( of abundance and the amount genus) both each by with average represented infested samples was of that (% samples frequency in genera of dominance nity; quanti and rbto fnmtd eeaacrigt hi apigsites. packages sampling using their genera check nematodes to to produced between was correspondence matrix according correlation the and genera map heat a nematode Additionally, of tribution otnradMry 1973 Merny, and Fortuner otnradMry 1973 Merny, and Fortuner ff ron: nteaessree nteTloieadTzah ein,PPNs regions, Taznakht and Taliouine the in surveyed areas the In software source open the by executed were treatments statistical All rnia opnn nlss(C)wsue ode to used was (PCA) analysis component Principal A “ corrplot fi sterglrt ftegnsdsrbto ihntecommu- the within distribution genus the of regularity the es hlti anktte ee9,6,3,ad2%respectively. 28% and 33, 69, 95, were they Taznakht in whilst iyecu,Ahlnhie,Pratylenchus Aphelenchoides, Ditylenchus, ” ff a mlmne oudriethe underline to implemented was 2011) al., et Bougeard ( o rp eogt 2fmle n 5gnr ( genera 15 and families 12 to belong crops ron fi aisr,2012; Falissard, l samples. eld hse ta. 2004 al., et Chessel .AC-nri nlss(O)wsperformed was (COI) analysis Co-inertia A ). ). Criconema aynWiadVla ik,2017 Simko, Viliam and Wei Taiyun and Tylenchorynchus , hc obn abun- combine which 2) Fig. ; Applied S Trichodorus .Ak A 2007). Dufour, and Dray oil Ecologyxxx(xxxx)xxxx and ≥ al 2 Table 0) 0 con- 40% 30%), and Helicotylenchus eeobserved were fi . Rotylenchus etedis- the ne i.2). Fig. “ psy ” ) . Version postprint di ( abundant in given and are Taliouine indices of sites diversity prospected for the codes for and plot regions Score Taznakht (B) indices; diversity the of 3. Fig. al. et Benjlil, H. ynhs Dorylaimus tyenchus, Tassous and Taouayalte di sites. tially remaining ( the set from data the isolated of variance Tassous total the Taouyalte, of The 44.16% region. for each counted in communities PPN of ( region the Taznakht in the greater in were loca- than indices all region diversity across all Taliouine that widespread showed are PCA genera Plotting indicating tions. that sites Tassous ( various in evenness the The seen Askaouen. being in highest lowest the ( index 1.47, diversity to Shannon 0.9 the whereas genera; various ff rn ie hwdagnsrcns ( richness genus a showed sites erent h iest seseto Psascae ihsa with associated PPNs of assessment diversity The ltigPAo h eaoegnr xiie di exhibited genera nematode the of PCA Plotting rnia opnn nlsso iest nie:()PAlaigplot loading PCA (A) indices: diversity of analyses component Principal Plant-parasitic nematodes parasitizing saffroninMorocco: Structuring driversand biological ff Benjlil, H., Elkassemi,K., AïtHamza, M.,Mateille,T., Furze,Cherifi, Mayad,Ferji (2020). rnitdby erentiated Tylenchorynchus risk identification. Applied Soil Ecology, 147. ,DOI :10.1016/j.apsoil.2019.103362 fi and n gdrMlolcmue eepartially were communes Melloul Agadir and Helicotylenchus Tylenchorynchus and Tylenchus E agdbten05t .9for 0.79 to 0.5 between ranged ) n gdrMlolby Melloul Agadir and G ). Comment citer cedocument: fi agn rm2t 0with 10 to 2 from ranging ) s w xso h C ac- PCA the of axes two rst ( ) hr a no was There 4C). Fig. i.3). Fig. fi n saseadthe and Assaisse and ff rn structures erent H’ al 2 Table aidfrom varied ) B). & 4A Fig.

ff fi o nthe in ron eepar- were . Para- 4 httergosaeapoiaeydsrbtdaogthe along distributed approximately are regions the that ta. 2016 al., et oasu K,mnaee(n,popou (P phosphorus (Mn), manganese (K), potassium yecu,Hlctlnhs Pratylenchus Helicotylenchus, tylenchus, etrs( features (Tassous ( regions the of plot loading The (H). humidity and (T) the with positively sandy in absent e were no genera had the altitude tribution. of the majority Where The soils. soils. while clay soils, in silt tected with associated calcium strongly (Fe), iron (Zn), ( zinc (Mg) organic magnesium (Cu), with and copper correlated (Ca) (N), negatively nitrogen were (OM), genera matter The (Ec). salinity and 3.3. eeawr ent epstvl orltdwt ietn (CaCo limestone with correlated positively be to seen were genera factors pedoclimatic h opsto,feuny bnac n iest sesetof di assessment sa the diversity with and associated diversity. abundance communities PPNs PPN frequency, of composition, risks. drivers major The nematological the and are parameters parameters These agronomical sa environmental, that discovered by we regions, Taznakht and nematodes plant-parasitic of diversity Taxonomical 4.1. Discussion 4. soils. clay and Taliouine. nakht) in occurs that soil Tylenchorynchus silt and wet to ra hsmyb xlie yteoii ftesa the of origin the by explained Taznakht the be the in may seen in was This observed diversity less area. were Comparatively sites. evenness Melloul Tasouss and in especially area diversity Taliouine Shannon richness, ( axis Tylenchorynchus Helicotylenchus Ditylenchus corms in Diversity 3.1.2. di Aphelenchoides Dorylaimus Tylenchorynchus and Pratylenchus, Helicotylenchus, Ditylenchus, Aphelenchus, mi- from free ( completely nematodes were endoparasitic (TA) (AM) gratory Melloul Taouayalte Agadir and from Wi collected (Ouisselssate) corms All Tailouine. in occurrence Tassous in Contrastingly, in respectively (especially occurrence moderate and Taliouine high with frequent most being orltd( 1). = (r correlated genera. PPN taxa between Co-occurrence 3.2. ff rnebtenhg,mdrt n o liue ( altitudes low and moderate high, between erence ln rwhpoutvt sgetya greatly is productivity growth Plant h BL odn lto h P lutae that illustrates PPN the of plot loading MBPLS The Paratylenchus Helicotylenchus, Aphelenchus, h O nlsshglgtdta h aoiyo h detected the of majority the that highlighted analysis COI The uigorsre odce ntesuho ooc nTaliouine in Morocco of south the in conducted survey our During he P eeawr eoee rmcletdsa collected from recovered were genera PPN Three etmp eeue orpeetteitreainhp between interrelationships the represent to used were maps Heat orsodnebtenpatprstcnmtdstx and taxa nematodes parasitic plant between Correspondence ) lmtcfcos( factors Climatic 7A). Fig. ff rn ie uvyd ihdvriyidcsicuiggenus including indices diversity High surveyed. sites erent fi adie l,2001; al., et Kandji ,temcolmt fec raadsi physico-chemical soil and area each of microclimate the ), , Aphelenchoides .Nmtdsblnigt h re yeciaare Tylenchida order the to belonging Nematodes ). Dorylaimus Ditylenchus , and Pratylenchus oiiecreain eeas on between found also were correlations Positive . and and Rotylenchus Dorylaimus Dorylaimus fi Tylenchus s xs whereas axis, rst ff and smr rqeti anktadhsalower a has and Taznakht in frequent more is o hwdvraiiybtenadwithin and between variability showed ron i.6). Fig. and and ae ta. 2005 al., et Cadet Tylenchorynchus ( )aerifl R) temperature (RF), rainfall are 7B) Fig. i.5 Fig. eepeeti an oain (Taz- locations rainy in present were eepstvl ikdt h second the to linked positively were Paratylenchus Helicotylenchus eepstvl orltdwith correlated positively were al 1 Table fi ff fi c nPN soitdwt dis- with associated PPNs on ect amue,Asis n Agadir and Assaisse Zagmouzen, , n saun n Taznakht. and Askaouen) and ). and Applied S ). Ditylenchus ff Paratylenchus Aphelenchoides ff peecods Tylenchus, Aphelenchoides, ce yPN ( PPNs by ected ron eesrnl positively strongly were eeawr associated were genera ; and 2 auie l,2017 al., et Karuri oil Ecologyxxx(xxxx)xxxx O with fi 5 ,sdu N) pH (Na), sodium ), lsaein are elds Pratylenchus )showed 7C) Fig. i.4 Fig. a anyde- mainly was Aphelenchoides fi eerelated were ff , s axis. rst Ditylenchus o corms: ron ff D). Tylenchus o crop ron fl Mateille uenced were ). Di- 3 ), , , Version postprint .Bnll tal. et Benjlil, H. hlssmls iyecu ipc,Brahlnhsxylophilus, Bursaphelenchus dispaci, Ditylenchus similis, pholus h ito cnmclyms motn Pshsbe modi been has spp PPNs gyne (2013) important al. most et economically Jones of list ( pests the agricultural major as considered sa in nematodes parasitic di plant of genera Prospected 1 Table 4. Fig. amue Z) ++:mr rqet +:feun;+:ls rqet;-: - ; frequent less : + Tassous frequent; : (SI), ++ frequent; Absent). Ouislsate more Siroua : (KH), (+++ (SH), Khosama (ZA); Zagmouzen Hssaine (IZN), Iznaguene Sidi (ASK), (Wi), Askaouane (AS), Assaisse listed is variables environmental and taxa the for encoding and variation altitudinal the for plot Score (D) sites; prospected for plot in Score (C) regions; prospected for Taznakht Taliouine Area ff al 2 Table rnslcto rmTloieadTzah ein:Aai elu (AM), Melloul Agadir regions: Taznakht and Taliouine from location erents rnia opnn nlsso htprstcnmtdsgnr soitdto associated genera nematodes phytoparasitic of analyses component Principal Plant-parasitic nematodes parasitizing saffroninMorocco: Structuring driversand biological Benjlil, H., Elkassemi,K., AïtHamza, M.,Mateille,T., Furze,Cherifi, Mayad,Ferji (2020). ., . Heterodera risk identification. Applied Soil Ecology, 147. ,DOI :10.1016/j.apsoil.2019.103362 i++++ +++ Si Z +++ Kh Wi IZN TA S++ ASK AM AS S+++++ +++ + + Za TS SH Site h eotdta h ucsiespecies: successive the that reported who , spp. –– – –– –– –– –– – –– – iyecu peecodsHelicotylenchus Aphelenchoides Ditylenchus n Globodera and enr ta. 2017 al., et Bernard ++ ++ +++ –– + + Comment citer cedocument: spp. fi T) auyle(A and (TA) Taouayalte (TS), Pratylenchus , ff o om olce in collected corms ron – + – – – –

.Recently ). spp. Meloido- fi ,Rado- dby ed 5 sativus scienti bessseyi aberrans Naccobus lenchoides index, Xiphinema reniformis, Rotylenchulus cuso ta. 2007 ( al., et McCuiston fTloieTzah sntsmlrt h eease nKhorozan in ( seen Psilenchus genera Merlinius, the cenamus, to similar ( not (Iran) is Taliouine-Taznakht of fi Pratylenchus eeao P nsa in PPN of genera ioyecu,Payecu,Hrcmnel,Psilenchus. Hirschmaniella, ( Pratylenchus, licotylenchus, Ahlnhs peecods oyecu,Helicotylenchus Rotylenchus, lenchus Aphelenchoides, (Aphelenchus, eerpre oprst h sa the parasite to reported were Taznakht nakht. ub swl fsa 2011 of well as bulbs ecl,1903; Metcalf, hihe l,2017 al., et Sheikh aznegre l,2016 al., et Ranzenberger dnsadcntttn h eaooia ikt hscrop. this to risk nematological the constituting and ndings rcssativus Crocus h opsto fPN soitdwt sa with associated PPNs of composition The mn h rsetdnmtd eeai h olo Taliouine- Aphelenchoides of soil the in genera nematode prospected the Among ). fi eeidenti were ) ndvrecutis( countries diverse in adkaiadAvn,2013): Alvani, and Mahdikhani Aphelenchoides n cnmcsigni economic and c Aphelenchoides , Ditylenchus . A C odn lto h eaoegnr;()Soeplot Score (B) genera; nematode the of plot loading PCA (A) L.: cek 1970 Schenk, p.and spp. r h o 0ms aaigPN u otheir to due PPNs damaging most 10 top the are ff a a was ) o rp ( crops ron fi ff dwti P omnte nTaliouine-Taz- in communities PPN within ed a enidenti been has o rp fTloieTzah.Teewere These Taliouine-Taznakht. of crops ron ). , Ditylenchus and Pratylenchus ff 1gnr eelse,and listed, were genera 11 ) Ditylenchus ce by ected ,weei asssutn fpat u to due plants of stunting causes it where ), .Sa fi ekr 1989 Decker, ac.I hssuy edtce three detected we study, this In cance. Aphelenchoides ruoadOo,2002 Oros, and Ortuño ff ff o crop. ron o ntevle fKsmr(India) Kashmir of valley the in ron , fi Pratylenchus fi i evsadcrsof corms and leaves in p.gnrlyatc tm and stems attack generally spp. egnr being genera ve a endtce nsa in detected been has ed mlmriis iecu,Geo- Filenchus, Amplimerlinius, Applied S ; ruoadOo,2002; Oros, and Ortuño uprigteabove the supporting , ff o ntesuyregion study the in ron oil Ecologyxxx(xxxx)xxxx and ; Helicotylenchus yecu,He- Tylenchus, olse al., et Vovlas fi eo them of ve nSpain In and and Aphe- ff ron Ty- C. Version postprint ..C-curnebtentaxa between Co-occurrence 4.2. tions. sa of species Pratylenchus etdby fested itiue nsi hr sa where soil in infection distributed corm no addition, In by genera. nematode the these to of development relate may observations di Such and Ouisselsate nematode-free. Melloul, Agadir were from Taouayalte obtained corms whilst sites, the of a in listed is taxa the for encoding and L. 5. Fig. codes. corresponding and analyzed Variables 2 Table al. et Benjlil, H. ff Helicotylenchus Ditylenchus Dorlaimus Criconema Anguina Aphelenchoides Aphelenchus genera Nematode Variables Heterodera Tylenchorhynchus Tylenchus Trichodorus Rotylenchus Pratylenchus Paratylenchus Zygotylenchus ff cigro rwh( growth root ecting rnei niomna odtoso admngmn restricting management land or conditions environmental in erence Pratylenchus oiiecreain eeeietbtennmtd aai the in taxa nematode between evident were correlations Positive otatnl,i hssuy u ape sa sampled our study, this in Contrastingly, etmpbtendi between map heat A Plant-parasitic nematodes parasitizing saffroninMorocco: Structuring driversand biological Benjlil, H., Elkassemi,K., AïtHamza, M.,Mateille,T., Furze,Cherifi, Mayad,Ferji (2020). iyecu Aphelenchoides Ditylenchus risk identification. Applied Soil Ecology, 147. ,DOI :10.1016/j.apsoil.2019.103362 ff curn nTloieTzah a o eparasitic be not may Taliouine-Taznakht in occurring o n curnei u ohbta eelco rota- crop cereal habitual to due is occurrence and ron a eoddi u td,ee huhi swidely is it though even study, our in recorded was oe ta. 2013 al., et Jones Hel Dit Dor Cri Ang Apo Apu Het Tyc Tyu Trd Rot Pra pTyl Code Zyg ff ff rn P eeaascae with associated genera PPN erent o rpwscliae.Ti ugssthe suggests This cultivated. was crop ron and al 2 Table Helicotylenchus ). Comment citer cedocument: Ouislsate Khosama Inzagune Askaoune Melloul Agadir Taliuoine Taznakht iiHsaine Sidi Climate Altitude Zagmouzen Taouayalte Tassous Siroua Variables evenness diversity local richness species nematodes of number total indices Diversity (mm) rainfall annual (°C) month hottest the of aiu temperature maximum egahcregions Geographic . fi ff o om eein- were corms ron ntemajority the in

rcssativus Crocus 6 oyhgu edn eaius uedsrbto n interspeci and distribution ( huge competition behaviours, feeding polyphagous speci wide ( communities mixed-species 1990; Khan, with ss amn itr n te aaees( organ- parameters other other with and interrelations history host, farming appropriate isms, an habit, speci pagation a of existence The ..Crepnec fpatprstcnmtd omnte oedaphic to communities nematode conditions plant-parasitic of Correspondence 4.3. i h mato neato eutn nices rihbto fne- of inhibition or increase an ( resulting reproduction interaction matode of impact the via ynchus nTloieadTzah einswr oiieycreae with ( studies correlated previous with positively agreement were 2014 in soils, regionss clay and Taznakht loamy and Taliouine in physico-chemical urn td,admyb xlie ytersnritcinteraction synergistic ( their persistence by and behaviours explained feeding be similar may through and study, current yegsi eainhpwsdsusdby discussed was relationship synergystic by studies previous in between ported observed correlation lenchus negative the is ecu dipsaci lenchus 1974 of ( texture fPN eeosre nlaysiswis lysisehbtdlow exhibited soils clay whilst of soils distribution loamy The in abundances. PPN observed were PPNs of nigteocrec fsm pce ( species some of occurrence the encing ( texture soil Pratylenchus, cilacus, oe stemvmnso hs eaoe eedo olporosity, soil on depend nematodes these of movements the as todes rpriso oladsigni and the on soil matter. information a organic essential of to provides properties and particles other soil each of arrangement to The related arranged are particles soil 2018 al., et Melakeberhan TS B,C SI WI KH IZN ASK AM Tal Taz SH ZA TA Code E H G RF T.Max N ’ Pratylenchus eainhp ewe h itiuino P eeaadsoil and genera PPN of distribution the between Relationships ; ; Aphelenchus ï-az ta. 2015 ; al., et Aït-Hamza hmsne l,2010 al., et Thompson hs eut upr h negnrcihbto htwsre- was that inhibition intergeneric the support results These . and .Mroe,hg reproduction high Moreover, 2002). Frederick, and McSorley fi Dorylaimus omnte htaerltdt hi perseverance, their to related are that communities c .PN fe aaiiepat in plants parasitize often PPNs 1991). Niblack, and Norton .Txuei ao atrin factor major a is Texture 1997). Geraert, and Ferji otnrn,1966 Oostenbrink, ( alc,1962 Wallace, a eoddi lysis( soils clay in recorded was , Conductivity Potassium aacollection Data pH Phosphorus Magnesium Iron Copper Nitrogen Clay silt sand Manganese Zinc Sodium Variables olcharacteristics Soil einlArclua eeomn O Development Agricultural Regional matter Organic atr eehglgtd h otdmnn genera dominant most The highlighted. were factors Ditylenchus oyecia n yecia oeae coarse a tolerated Tylenchidae and Rotylenchidae iebc,1993 Eisenback, Interspeci . fi pce nast sascae ihispro- its with associated is site a in species c .Si etr n tutr ee oteway the to refer structure and texture Soil ). os(1964) Ross fi .Te perin appear They 1978). Perry, and Jones , ï az ta. 2018 al., et Hamza Aït .Asmlrtedwsrpre for reported was trend similar A ). atyin cantly .I os regions Souss In ). Helicotylenchus, ). fi oocrec a rvn force driving a has co-occurrence c Tylenchus Pratylenchus .Afurther A ). fl ecstepeec fnema- of presence the uences ae n holue 1998; Thioulouse, and Cadet and Applied S idadJnis(1964) Jenkins and Bird ms (1970) Amosu otn 1978 Norton, rnio n Wallace, and Grandison spstvl correlated positively is Pratylenchus ffi Tylenchus oil Ecologyxxx(xxxx)xxxx ce sdtrie ysoil by determined is peecods Gra- Aphelenchoides, fi .Hg abundances High ). dn fti study this of nding iebc,1993). Eisenback, aaie al., et Salahi and , ; hltthe whilst , Tylenchor- hnand Khan ORMVA pH K Fe Sil San Con Zn Cu N Cla N P Code Mg Mn OM Zygoty- Dity- . fl fi u- c Version postprint vrl,si etrspa usata oei h bnac,dis- abundance, the sa with in associated role communities PPN substantial of a structure and play tribution textures soil Overall, aiu olpyiohmclfcoshv enrltdt eeaand genera to ( related PPNs been some of have gradients omni- density factors of physicochemical abundance nematode soil the ( various on nematodes reduce maturity predatory impact and and reducing adverse nematodes vorous richness, free-living an of genus cause indices decreasing (Zn) structure, zinc community and (Cu) Copper ( poisonous ammoniacal PPNs as and considered nitrate to are of that degradation accumulation nitrogen the through by nitrogen explained be can That abundance nematode of decrease accu- direct OM a (OM): matter for ( organic accounts with correlation mulation negative a showed ( water of moisture thickness retain particles, soil of size 6. Fig. al. et Benjlil, H. hmclcaatrsis C cr ltfrpopce ein;()Soepo o h liuia aito n noigfrtx n olpyioceia char- physico-chemical soil and taxa for encoding and variation altitudinal the for plot Score (D) regions; in prospected listed for is plot acteristics Score (C) characteristics; chemical ansu M)adptsim()hsbe eotd( reported been has 1987 (K) potassium and (Mg) between magnesium correlation ( positive (Ca) a calcium Conversely, and (Mg) magnesium and irgn() xetfrthe for except (N), nitrogen ( 2002 OM with increase that antagonisms rn(e,clim(a n oim( sodium and (Ca) 2012; al., et calcium Yavuzaslanoglu (Fe), iron oiik 1999; Hominick, oto h eeaidenti genera the of Most ; .Ti td con study This ). one l,1993). al., et Zoon oietaaaye ewe P n olpyioceia hrceitc:()PAlaigpo ftenmtd eea B C odn lto olphysico- soil of plot loading PCA (B) genera; nematode the of plot loading PCA (A) characteristics: physico-chemical soil and PPN between analyses Co-inertia Plant-parasitic nematodes parasitizing saffroninMorocco: Structuring driversand biological Benjlil, H., Elkassemi,K., AïtHamza, M.,Mateille,T., Furze,Cherifi, Mayad,Ferji (2020). orge-aaae l,1981 al., et Rodriguez-Kabana risk identification. Applied Soil Ecology, 147. ,DOI :10.1016/j.apsoil.2019.103362 .I hssuy oto h genera the of most study, this In 1989). Decker, rPN r ujce omicrobial to subjected are PPNs or 2007) Hu, and Qi al 2 Table fi mdta Psaudnei orltdwith correlated is abundance PPNs that rmed . Aphelenchoides fi n sa is and 2017) al., et Karuri dso eaiecreainwt total with correlation negative a show ed alc ta. 1993 al., et Wallace .I addition, In 2002). al., et Georgieva urne l,1980 al., et Guiran fi ae n holue 1998 Thioulouse, and Cadet Comment citer cedocument: ; , m,adtesi' blt to ability soil's the and lms, icsadNhr 2002; Neher, and Fiscus Helicotylenchus Ditylenchus orge-aaa 1986 Rodriguez-Kabana, ; hne l,2012 al., et Chen and , oisne al., et Robinson ; imre al., et Widmer ff

ce ypH by ected n acidity, and Criconema. ff ron. ). ). ). 7 sdtrie codn ootmltmeaueo t ieprocesses life its of temperature optimal to species ( nematode according each determined of range is and distribution geographic reproduction optimum. the nematode Moreover, temperature higher precise in a ( result development at may place temperatures take High PPN of activities gical 2007 ( moisture rbto fPNcmuiisaein are communities PPN of tribution a a may ( distribution nematodes uiiscudb ikdt ilgclfaue fnmtdsthat nematodes of features biological to linked be could munities eauehda insigni an had perature ( humidity and communities PPN rainfall structure to through force particularly driving the are conditions climatic while and Taliouine in curred Pratylenchus variables climatic to nematodes parasitic plant of Correspondences 4.4. oreo iessfrarsses lns relvn nml n hu- and animals living free plants, ( agrosystems, mans a for are diseases agents These of agents. source infectious various of spread and development eriae l,2018 al., et Ferreira encon been u ta. 2005 al., et Luc h utbokaayi hwdthat showed analysis multiblock The Dorylaimus ; ff ile ta. 2014 al., et Nielsen avl ta. 2002 al., et Harvell c h itiuino eaoe ic lmt slne othe to linked is climate since nematodes of distribution the ect fi elo n og 1996 ; Boag, and Neilson mdby rmed and zrzkksadTugl,2005 Trudgill, and Tzortzakakis eefudi an oain fTzah.Herein, Taznakht. of locations rainy in found were .Tesrn eainhpbtentmeaueand temperature between relationship strong The ). Paratylenchus ; aue l (2018) al. et Namu aue l,2018 al., et Namu fi oge l,1991 al., et Boag atipc nPNcmuiiswa has what communities PPN on impact cant .I hudb oe htms ftebiolo- the of most that noted be should It ). ; peecods yecu,Tylenchorynchus Tylenchus, Aphelenchoides, art ta. 2013 al., et Garrett eerltdt h uiiyta oc- that humidity the to related were ode l,1999 al., et Todd fl .Cnesl aiu tem- maximum Conversely ). uenced h iest n h dis- the and diversity The . mle htciaechange climate that implies ) Applied S iyecu,Helicotylenchus, Ditylenchus, ï az ta. 2018; al., et Hamza Aït ; vn n er,2009). Perry, and Evans .Tee The ). oil Ecologyxxx(xxxx)xxxx ytmeaueand temperature by ; aoy tal., et Bakonyi ff cso com- on ects Version postprint o aaadsi hsc-hmclcaatrsisi itdin listed cli- is characteristics and physico-chemical chemical soil physico the and soil of taxa sa the for for plot plot of loading Score plot (C) PCA parameters; Loading matic (A) PCA parameters: (B) climatic genera; nematode and chemical physico- soil 7. Fig. al. et Benjlil, H. Plant-parasitic nematodes parasitizing saffroninMorocco: Structuring driversand biological ut-lc nlssbtenpatprstcnmtd communities, nematode plant-parasitic between analysis Multi-block Benjlil, H., Elkassemi,K., AïtHamza, M.,Mateille,T., Furze,Cherifi, Mayad,Ferji (2020). risk identification. Applied Soil Ecology, 147. ,DOI :10.1016/j.apsoil.2019.103362 ff o region ron Comment citer cedocument: fi lssmldadencoding and sampled elds al 2 Table

. 8 lenchus pce,cnieigeeet ftesi omnt nertdit GxE into integrated community expressions soil the of elements considering species, temperature increased to related is ( development and reproduction climatic and in variations temperature environmental The to changes. reactions variable govern civ out/cnrlal iwo ln olitrcin.This interactions. identi soil further and plant communities of PPN of view drivers controllable some detailed / study para- robust further a the and achieve of ( equations predicted elements GxE the in parameterized in be community soil integrating patterns may of recommend series we which a meterized, nematodes indicate results of our behavior as statement this make ae.Smlry h oldyesin dryness a soil directly the which Similarly, moisture, rates. to related were rae teto n intensi and attention creased sa a drive also loss. additionally yield is would distribution infection and root abundance structure, for looking ( soil moisture in PPNs of lifetime ( permeation e through tribution otseis oso h iceesrcueo iarsosand bioagressors of structure discrete the show to species, host sy- or Clar- hierarchical tool. based into Markov-chain a placed within re- i being expression these for before though terms community nergistic ranking soil further the of quire distributions species discrete hne yarclue n yarclueintensi agriculture by use land and by agriculture, improved is by PPNs of changes development the Consequently, plan. liaino h olfrhmnnes eefe,w ekt lrf the clarify to e seek and we causal Hereafter, physiological needs. ex- of human biological nature for continued soil the the of consider ploitation and assessment risk cultivation formulative and sa as production such to species risks valuable the of clarify inter- ultimately of will latter order The ( precise tools ascertain predictive within to actions settings species of feeding environmental emergence the variable continual further in to and required is strategies functions based based behavior life-history within ordination al- ( 2019 have nematodes al., we of et Although studies Hoogen tools. mapping the functional from commenced results ready accurate proceed re obtain we highly to before as chains required approach are community this soil with the of members other iusadmv om rmone from corms move and niques ( pro- sustainability a crop health and conventional system of soil, methods degrade cultivation duction Intensive 2008). al., et hudb eeoe tnreyand nursery at developed be should previous supports work Our texture. soil fi and practices soil by impacted ( loss study yield This by increased. infected is corms risk a demonstrated agronomic As of materials. emergence and soils the corms, consequence, contaminated in PPN disperse practices oeta ietosfrivsiain nld h clari the include investigations for directions potential levels abundance and composition communities ( PPN to linked heavily n ouaindvlpeto h otdmgn Pssc as such PPNs damaging most the Pratylenchus Aphelenchoides, of spread Ditylenchus, restrict development ra- that develop population practices to agricultural and is suitable investigations and future measures of tional focus the Consequently, tion. eoyeadEvrnetl(x)epesoso sa of expressions (GxE) Environmental and Genotype sa in management nematode to approach agroecological 2018 fi uc ta. 2012 al., et Duyck aoae-ise l,2015 al., et Palomares-Rius dnsta olpyiohmclfcosadciai aibe are variables climatic and factors physicochemical soil that ndings aino taeia n ucinlgop ihnnmtdsand nematodes within groups functional and strategical of cation Sa h ucm fti td rvdsabssfrdvlpn an developing for basis a provides study this of outcome The ). ff o rp nTloieadTzah ein hudrciein- receive should regions Taznakht and Taliouine in crops ron pce speci species .Ipcso eprtr nnematode on temperature of Impacts 2011). Colagiero, ai ta. 2013 al., et Latif oehneterslto cuayo h x eut We result. GxE the of accuracy / resolution the enhance to oet ta. 1981 al., et Roberts .h atri otayt u eut u to due results our to contrary is latter ).The ff ,adtoa eonto fseiseegneand emergence species of recognition additional ), fi cietraso edn ( feeding of threats ective hrceitc.I hssuy oto h genera the of most study, this In characteristics. c .Nmtdsaudneaddvriyare diversity and abundance Nematodes ). ff ff ; .Fresuetaiinltech- traditional use Farmers 2011). Lal, fl c idvriycnevto n agro- and conservation biodiversity ect o.Sc ol ieanvldmninto dimension novel a give tools Such ron. fi ï az ta. 2018; al., et Hamza Aït ecsnmtd bnac n dis- and abundance nematode uences uz ta. 2017 al., et Furze ainacrigt Morocco to according cation fi Ditylenchus .teicesdrt of rate increased the ). ere n on,1996 Pooni, and Kearsey e nu utb e noMarkov into fed be must input ned fi fl ff l oaohr ec agricultural Hence another. to eld ecstesria fPN;the PPNs; of survival the uences co ucin fPN n their and PPNs of functions ector fi l rdcinlvl.Further, levels. production eld Applied S and hc assu o62% to up causes which , ff csnmtd infection nematode ects Helicothylenchus og 1980 Boag, oil Ecologyxxx(xxxx)xxxx ; aade l,2019). al., et Basaid fi ain( cation ff o n other and ron ff aue al., et Namu ,wihwill which ), Pratylenchus o produc- ron fi n root and ) ff ainof cation ce by ected a Den Van ’ Mateille which , green s Praty- ff fi ron ed Version postprint id . ekn,W,16.Ocrec,prsts,adptoeiiyo nematodes E 1980. of B., pathogenicity Boag, and parasitism, Occurrence, 1964. W., Jenkins, G., Bird, .Bnll tal. et Benjlil, H. enr,GC,Enn . os,C,21.TeIpc fPatPrstcNmtdson Nematodes Plant-Parasitic of Impact The 2017. C., Bonsi, M., Egnin, G.C., Bernard, aad . hbi . uz,JN,Myd .. ohrod . 09 h au fsi- of value The 2019. R., Bouharroud, E.H., Mayad, J.N., Furze, B., Chebli, K., Basaid, on,D,Ncl . luisCl,B,Qééev,P,21.Lsnmtdsdsplantes: des nématodes Les 2010. P., Quénéhervé, B., Claudius-Cole, J., Nicol, D., Coyne, abn,D,Cak . 90 jlalmto o eemnto fttlntoe content nitrogen total of determination for method Kjeldahl 1990. J., Clark, D., Barbano, of Interaction 1970. J.O., Amosu, inorganic and organic for procedure and apparatus Wet-combustion 1960. L., Allison, oair,M,21.Ciaecagsadnmtds xetde expected nematodes: and changes Climate 2011. M., Colagiero, aoy,G,Ng,P,Kvc-ag . oas . aaá,S,Rps,V,Srs . 2007. A., Seres, V., Répási, S., Barabás, E., Kovacs, E., Kovacs-Lang, P., Nagy, G., Bakonyi, associated nematodes Plant-parasitic 2014. T., Mateille, J., Tavoillot, E., Chapuis, N., Ali, hn . Shea S., Chen, Identi 1998. J., Thioulouse, P., Cadet, hse,D,Dfu,A,Tiuos,J,20.Teae akg.I n-al ehd.R methods. One-table I. package. ade4 The 2004. J., Thioulouse, A., Dufour, D., Chessel, ï-az,M,Fri . l,N,Tvilt . hpi,E,E ulai . okl,A., Moukhli, A., Oualkadi, El E., Chapuis, J., Tavoillot, N., Ali, Z., Ferji, M., Aït-Hamza, ï az,M,Muhi . ej,Z,FsaiGshgad . aolo,J,Ai N., Ali, J., Tavoillot, O., Fossati-Gaschignard, Z., Ferji, A., Moukhli, M., Hamza, Aït ae,P,Mse . holue . 05 eainhp ewe ln-aaii nema- plant-parasitic between Relationships 2005. J., Thioulouse, D., Masse, P., Cadet, hae,O,RboMrg,A,Csil-óe,R,TaeoMzs . óe-óe,L., Gómez-Gómez, A., Mozos, Trapero R., Castillo-López, A., Rubio-Moraga, O., Ahrazem, ogad . anr,EM,Lp,C,Hana C., Lupo, E.M., Qannari, S., Bougeard, Boag, bene bai . aoi,M,21.MdcnlPatDsae asdb eaoe,Medicinal Nematodes, by Caused unpublished Diseases data Plant Interne Medicinal 2017. Massa. 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