GenetResourCropEvol(2012)59:981–997 DOI10.1007/s10722-011-9738-x RESEARCH ARTICLE Genetic resources collections of leafy vegetables (lettuce, spinach, chicory, artichoke, asparagus, lamb’s lettuce, rhubarb and rocket salad): composition and gaps R. van Treuren • P. Coquin • U. Lohwasser Received:11January2011/Accepted:21July2011/Publishedonline:7August2011 (cid:2)TheAuthor(s)2011.ThisarticleispublishedwithopenaccessatSpringerlink.com Abstract Lettuce, spinach and chicory are gener- nl/cgn/pgr/LVintro/. Based on a literature study, an ally considered the main leafy vegetables, while a analysis of the gene pool structure of the crops was fourth group denoted by ‘minor leafy vegetables’ performed and an inventory was made of the distri- includes,amongstothers,rocketsalad,lamb’slettuce, bution areas of the species involved. The results of asparagus, artichoke and rhubarb. Except in the case these surveys were related to the contents of the of lettuce, central crop databases of leafy vegetables newly established databases in order to identify the were lacking until recently. Here we report on the main collection gaps. Priorities are presented for update of the international Lactuca database and the futuregermplasmacquisitionaimedatimprovingthe development of three new central crop databases for coverageofthecropgenepoolsinexsitucollections. each of the other leafy vegetable crop groups. Requests for passport data of accessions available Keywords Chicory (cid:2) Crop database (cid:2) Germplasm to the user community were addressed to all known availability (cid:2) Lettuce (cid:2) Minor leafy vegetables (cid:2) European collection holders and to the main collec- Spinach tion holders located outside Europe. Altogether, passport data of 17,530 accessions from a total of 129 collections were collected. The four separate Introduction databasesweremadeavailableonlineviaacommon entry page accessible at http://documents.plant.wur. Leafyvegetablesconstituteahighlyvariablegroupof crop plants, which are grown for their edible leaves, which are rich sources of minerals and vitamins. R.vanTreuren(&) Lettuce,spinachandchicoryaregenerallyconsidered CentreforGeneticResources,TheNetherlands(CGN), the main leafy vegetables because of the worldwide WageningenUniversityandResearchCentre, P.O.Box16,6700AAWageningen,TheNetherlands humanconsumptionandeconomicalimportance.For e-mail:[email protected] these three crops combined, the world production quantity in 2008 reached 38.11 million tons on a P.Coquin harvested area of 1.95 million hectares (FAO 2010). VarietyStudyDivision,DomainedelaBoisselie`re, GEVES,49250Brion,France Otherleafyvegetablesinclude,amongstothers,rocket salad,lamb’slettuce,asparagus,artichokeandrhubarb U.Lohwasser (Lebeda and Boukema 2001). This fourth group is DepartmentofGenebank,LeibnizInstituteofPlant denoted by ‘minor leafy vegetables’, which refers GeneticsandCropPlantResearch,Corrensstrasse3, 06466Gatersleben,Germany moretothelevelofattentionthesecropshavereceived 123 982 GenetResourCropEvol(2012)59:981–997 in the past, rather than to their market impact. For interest(HodgkinandHajjar2008).Efficientaccessto example, the worldwide harvested area of asparagus thesegeneticresourcesisthereforeessentialinmodern was1.39 millionhectaresin2008,whichexceedsthe plantbreeding. valueof1.06forlettuceandchicorycombined(FAO Inadditiontothepresenceofdesiredtraits,thelevel 2010). Furthermore, crops such as rocket salad and ofinter-fertilitywithcultivatedplantsisoneofthekey lamb’slettucehaveincreasedinproductionbecauseof factors determining the usefulness of wild species in their use in processed products, such as mixed salad plantbreeding.Speciesmaybeclassifiedaccordingto packages that have received growing interest from theirpositionwithinthetotalgenepoolfollowingthe consumersduringthelasttwodecades. principles of Harlan and de Wet (1971) who distin- European breeding companies, next to companies guishedthreecategoriesoftaxabasedonthelevelof located inthe UnitedStates, have a strong position in inter-fertility.Theprimarygenepoolisformedbythe the seed market of leafy vegetables. As in many cultivated and wild species that can be inter-crossed vegetablecrops,breedingactivitiesinleafyvegetables relativelyeasy,generallyresultinginfertileoffspring. aremainlydirectedtoqualitycharactersandimprove- Thesecondarygenepoolconsistsofspeciesthatshow ment of the resistance against pests and diseases. only low levels of fertility with cultivated plants, Currently,resistancetolettucedownymildew(Bremia although gene transfer is possible through consider- lactucaeRegel)andlettuceaphid[Nasonoviaribisnigri able effort. The tertiary gene pool is constituted by (Mosley)] are the main characters in lettuce breeding species that usually do notdisplay inter-fertility with (Lebeda et al. 2007; Mou 2008), while resistance to cultivatedplants,requiringtechniquessuch asbridge spinach downy mildew is of main interest in the crosses or embryo rescue to obtain some fertile improvementofspinach cultivars (Correlletal.2003; offspring.Studiesonspeciesrelationshipswithinleafy Morelock and Correll 2008). The prime characters in vegetable crops have been performed predominantly chicorybreedingvaryamongthedifferentspeciesand forlettuce(forareviewseeLebedaetal.2007). subspecies. The two main species within the genus A survey of genetic resources collections of leafy CichoriumareC.endiviaL.,includingssp.endivia(cut vegetablesin1999showedthatapproximately23,000 and plain endive), and C. intybus L., including var. accessions were conserved on a global scale (Van foliosum Hegi(leaf chicory, witloof) and var.sativum Hintum and Boukema 1999). In 2000, a central crop (Bisch.) Janch. (root chicory). For witloof and root database for lettuce, called the international Lactuca chicory, the latter being used either for roasting or database (ILDB), was developed (Stavelikova et al. inulinproduction,breedingeffortsfocusonproductiv- 2002).However,availabilityoftheincludedmaterial ity and improvement of nutritional and taste compo- to the user community was not considered. Until nents,inadditiontotheimprovementoftheresistance recently, databases for other leafy vegetables were topestsanddiseases(Dore´ andVaroquaux2006).For lacking. Here we report on the recent update of the cut and plain endive and for the Italian leaf chicory ILDBandtheestablishmentofcentralcropdatabases cultivargroups(e.g.ChioggiaandVerona),adaptation for spinach, chicory and minor leafy vegetables, tonovelgrowinghabitsandresistanceimprovementto takingavailabilityoftheaccessionsintoaccount.The pests and diseases are the main characters of interest composition of the databases is presented and (Bellamy 2000). Only little is known about breeding analyzed with respect to species representation and activitiesforminorleafyvegetables.Forlamb’slettuce coverage of the main distribution areas in order to onemainaspectistodevelopastrategyforresistance identify important collection gaps that may serve as against downy mildew (Peronospora valerianellae guidelines for future germplasm acquisition. Fuckel) (Pietrek and Zinkernagel 2002). Other breed- ing programmes are dealing with marker assisted selection for the identification of special agronomic Materials and methods traits in artichoke (Martin et al. 2008, 2010) and with hybrid breeding in asparagus (Scholten and Boonen Database development 1996; Moreno et al. 2010). Particularly in the case of novel resistances, crop related wild relatives are The leafy vegetable crops that were addressed in the increasingly being explored to find the characters of presentpaperfollowedtheworkinggroupstructureof 123 GenetResourCropEvol(2012)59:981–997 983 the European Cooperative Programme for Plant made to verify the actual availability of the acces- Genetic Resources (ECPGR). However, distinction sions, neither was the validity of the supplied data between the different vegetable crop groups is not about taxonomy and geographic origin verified. Fur- precisely defined. For example, the Brassicas are thermore,thedatawerenotcheckedfortheexistence treated as a separate group although many cabbages of potential duplicate accessions within or between aregrownfortheiredibleleaves,whilerocketsaladis collections. Requesting and processing of the data considered a leafy vegetable although this crop is wascarriedoutbetween2007and2009.Requestsfor often included in Brassica collections. Dill is datawererepeatedincasenoresponsewasreceived. included in the Medicinal and Aromatic Plants In a few cases where no data were received for working group although the leaves form one of the European collections, the data from EURISCO were main use aspects of this herb. The crops considered downloadedandincludedinthedatabase.Forlettuce, by the leafy vegetables working group of ECPGR the data from Iran in the previous version of the was established in 2000 during a meeting of the ILDB were maintained because of missing updates. network coordinating group on vegetables. Lettuce, Theresultspresentedinthepresentpaperwerebased spinach and chicory were considered the main crops, on the data extracted in March 2010 from the inter- while the minor leafy vegetables included, amongst national leafy vegetables databases that can be others, rocket salad, lamb’s lettuce, asparagus, arti- accessed on line via http://documents.plant.wur.nl/ choke and rhubarb (Lebeda and Boukema 2001). cgn/pgr/LVintro/. Each of the four databases also To update the ILDB and to develop databases for include an overview of the contributors to the dat- spinach, chicory and minor leafy vegetables, it was abases, together with links to the institutional web- aimed to include all European collections and the sites and to the contact persons of the collections. maincollectionslocatedoutsideEurope.Arequestto provide accession information was directed to the Wild species and distribution areas members of the ECPGR working group on leafy vegetables (http://www.ecpgr.cgiar.org/Workgroups/ In order to identify main collection gaps within the Leafy_Vegetables/Leafy_Vegetables.htm), the hold- newly established databases an inventory was made ers of collections included in EURISCO (http:// of the crop related wild speciesand their distribution eurisco.ecpgr.org/home_page/home.php), the partici- areas. The International Plant Names Index (IPNI; pants in the EU GENRES project ‘‘Leafy vegetables http://www.ipni.org/index.html) of September 2009 germplasm,stimulatinguse’’(http://documents.plant. was used as the basis for the inventory. Species wur.nl/cgn/pgr/leafyveg/)andthemainnon-European nameswerecheckedforsynonymyandreassignment collection holders listed in the Bioversity Directory to other genera and reported distribution areas were of Germplasm Collections (http://www2.bioversity collected using information provided by IPNI, GRIN international.org/Information_Sources/Germplasm_ Taxonomy for Plants (http://www.ars-grin.gov/cgi- Databases/Germplasm_Collection_Directory/). Col- bin/npgs/html/index.pl), Flora Europaea (http://rbg- lection holders that were not part of the aforemen- web2.rbge.org.uk/FE/fe.html), Mansfeld’s World tioned groups but who were included in the first Database of Agricultural and Horticultural Crops versionoftheILDBwerealsoapproachedinthecase (http://mansfeld.ipk-gatersleben.de), Tropicos (http:// oflettuce.Requestsfordatawereaccompaniedbyan www.tropicos.org/) and the Cichorieae Portal (http:// Excel spreadsheet containing instructions for data wp6-cichorieae.e-taxonomy.eu/portal/). For lettuce, submission and a format for information on 35 theresultsofanearlierinventory(Lebedaetal.2004) passport descriptors largely following the EURISCO were included in the analyses. descriptor list (http://eurisco.ecpgr.org/documents/eu Tostructurethelettucegenepoolwefollowedthe risco_descriptors-update-feb2008.pdf). In the request classification described by Koopman et al. (1998; for data it was emphasized to provide information 2001) and by GRIN Taxonomy for Plants. This only of accessions that in principle are available for classification system is largely in line with the one distributiontousers.Receiveddatawerecorrectedin described earlier by Zohary (1991). However, com- case of obvious spelling and formatting errors, but pleteconsensusaboutspeciesrelationshipswithinthe otherwise remained untreated. No attempts were genus Lactuca is still lacking and discussed in more 123 984 GenetResourCropEvol(2012)59:981–997 detail by Lebeda et al. (2007). Hybrids and taxo- whereas for another 51 the taxomic status and their nomicclassificationsbelowthespecieslevel,suchas belonging to the genus Lactuca was questionable. subspeciesandforms,weredisregardedinthecaseof Of the remaining 130 species, 20 are generally lettucebecauseofthesizeofthegenusLactuca.Also considered to be part of the lettuce gene pool for chicory, species relationships within the genus (Table 1). Cichorium are still debated. Here we followed the Accession data were collected from a total of 30 chicory gene pool structure suggested by Kiers et al. collections, including contributions from Armenia, (2000)andB.Desprez(pers.comm.).Concerningthe Iran and the United States. Altogether, the data of gene pool structure of spinach usually three species 11,643 accessions were included in the ILDB. The are distinguished, namely cultivated spinach (Spina- main part of the ILDB is constituted by landraces, cia oleracea L.), which is inter-fertile with the wild cultivars, research lines and breeding materials of speciesS.tetrandraStevenexM.Bieb.(Sneep1957) cultivated lettuce (Lactuca sativa L.) represented andS.turkestanicaIljin(Handkeetal.2000).Forthe with 10,198 accessions (88%). A highly unbalanced group of minor leafy vegetables the analysis of the distributionwasobservedforspeciesbelongingtothe gene pool structure was restricted to rocket salad croprelatedwildgenepool as96%oftheaccessions (DiplotaxisandEruca)becauseoftherelativelyhigh arecoveredbyonlythreespecies,namelyL.serriola number of accessions involved for this crop and L., L. saligna L. and L. virosa L. Eight species, because of the limited information about species includingL.azerbaijanicaRech.f.andL.scarioloides relationshipsavailablefortheothercrops.Inorderto Boiss. that are part of the primary lettuce gene pool, structure the gene pool of Diplotaxis and Eruca the evenlackasingleaccessionintheILDB.Concerning crossing ability and inter-fertility between species in the primary gene pool, L. serriola L. with 987 the entire family Brassicaceae was studied by means accessions is the only wild species that is well of a literature survey. Firstly, the relationship represented (Table 1). The category ‘‘unknown and between species within the genus Eruca and within other taxa’’ included 167 accessions, comprising the genus Diplotaxis was investigated (Eschmann- specimensofL.biennis(Moench)Fernald,L.canad- Grupe et al. 2003; Warwick et al. 2007). Secondly, ensis L., L. dentata Makino, L. dissecta D. Don, the relationship of each of these genera with other L. floridana (L.) Gaertn., L. homblei De Wild., Brassicaceae genera and species was determined L. indica L., L. livida Boiss. et Reut., L. perennis L., (Harberd and McArthur 1980; Warwick and Black L. raddeana Maxim. and L. tenerrima Pourret, a 1993; Prakash et al. 1999; Snowdon et al. 2007; number of hybrids and undetermined Lactuca spe- Warwick et al. 2009). cies, and a few samples from the genera Chondrilla, Tosummarizegeographicalinformation,countries Cicerbita, Mycelis and Steptorhamphus that are were grouped together according to the regional related to Lactuca (results not shown). subdivision of the United Nations Statistics Division The reported geographic distribution area of the (http://unstats.un.org/unsd/methods/m49/m49regin.htm) species of the lettuce gene pool and the degree of with the exception that for lettuce and spinach coveragebyaccessionsintheILDBisalsopresented the Russian Federation was regarded to comprise inTable 1.Accessionsofcultivatedlettucehavebeen Northern Asia instead of belonging to Eastern Eur- collected from all regions of the world, the majority ope, while Cyprus was considered to belong to originating from Europe and Asia and relatively few Southern Europe instead of Western Asia. from Africa. All the main horticultural types, i.e. ‘butterhead’,‘crisphead’,‘cutting’,‘cos’,‘stalk’,and ‘Latinlettuce’,arewellrepresentedintheILDBwith Results a wide variety in geographic origin (results not shown). The more primitive horticultural type ‘oil- Lettuce seedlettuce’,whichhasarestrictedcultivationinthe Middle East, is represented by six L. sativa L. Survey of the International Plant Names Index accessions, all originating from Egypt. No data on revealed a total number of 538 Lactuca species, of geographic origin was provided for 3,063 accessions which 357 referred to synonyms and basionyms, of L. sativa L. (30%). 123 GenetResourCropEvol(2012)59:981–997 985 ctuca rosa Laal Lvi.L. 67 2 3 23 39 15 7 1 7 0 0 0 1 0 1 0 58 141 n o ati na n g nter sali 7 6 3 7 1 8 3 4 0 1 0 0 0 0 0 0 0 8 3 ei L.L. 3 2 4 4 1 10 h t dedin rriola u e incl Ls.L. 458 119 19 116 204 344 77 0 24 9 234 14 1 13 0 16 0 16 0 0 155 987 ol po es ne oid ge ol lettuce Lscari.Boiss. 0 0 0 0 0 s) e n li a cal gic verti georossh. by L.Gr 0 0 0 0 0 1 1 d y(separate dregeanaC. ar L.D 0 0 4 4 erti t yandarea anica y,secondardistribution Lazerbaij.Rech.f. 0 0 0 0 primaretotal et heth h. tn s ciesofnwithi aicaFiMey. Lactucaessionsofspeortedgeographicregio LaculeataLalt..Boiss.C.A. 21 0 0 0 00 21 06 27 cp mberofacDB)perre Lsativa.L. 5,702 1,107 406 2,096 2,093 1,029 8 152 83 108 14 664 64 1 3 58 1 1 322 12 3 281 26 18 3063 10,198 Table1Nudatabase(IL Region Europe East North South West Asia Central East North South Southeast West Africa Central East North South West America Caribbean Central North South Oceania Unknown Total 123 986 GenetResourCropEvol(2012)59:981–997 a n a ni o s at wel. L.Tr 0 0 0 0 Lviminea.(L.)J.PresletC.Presl 10 2 4 4 7 0 0 0 7 0 0 3 20 ricaMey. a tatA. L.C. 2 0 0 2 4 0 0 1 0 3 0 0 2 8 a oliet cifh. ac xa Ltara.SchumThonn. 0 0 0 0 0 0 Lsibirica.Benth.exMaxim. 1 0 1 0 0 0 0 0 1 inbold d e cina denot r e e qu ar L.L. 2 2 0 0 0 2 0 0 2 0 4 ent n s nti ali co Lorient.Boiss. 0 0 0 0 0 0 0 0 0 otalsper T ntata y‘0’. e b d longioris noted L.M 0 0 0 0 de e ar ´c) B xˇci D ap.ePans IL Laure.(Sch.BiVis.etStebbin 0 0 0 0 0 0 0 dinthe e nt hifoliaBoiss. eprese antd.) otr ued acWill sn ntin L.( 0 0 0 0 0 0 gion Table1co Region Europe East North South West Asia Central East North South Southeast West Africa Central East North South West America Caribbean Central North South Oceania Unknown Total Reportedre 123 GenetResourCropEvol(2012)59:981–997 987 The majority of wild species of the primary gene Chicory pool have their distribution area in Asia, with the exception of L. dregeana DC. that is restricted to Thirty seven Cichorium species were found in IPNI South Africa and L. serriola L. that can be found on including 32 synonyms. The chicory gene pool all continents (Table 1). For L. serriola L., being the appearedquitelimitedintermsofnumberofdifferent only wild species of the primary gene pool that is species as it is represented by the two cultivated well represented inthe ILDB, relatively few samples species C. endivia L. and C. intybus L. and the three were collected from Africa (14) and America (16), wild species C. calvum Sch. Bip., C. spinosum Jacq. and none from Oceania. Like L. serriola L., also andC.botaeDeFlers(Table 3).Duetotheverylow L. saligna L. and L. virosa L., constituting the fertility resulting from interspecific crosses and the secondary gene pool, show a cosmopolitan distribu- very diverse use aspects of the cultivated types, the tion.Thepredominanceofaccessionsofthesespecies species C. endivia L. (cut and plain endive) and from Europe and Asia in the ILDB was even more C.intybusL.(rootchicoryandleafchicory,thelatter clear-cut than observed for L. serriola L. as no including witloof and the Italian chicory cultivar accessionsfromAfricaandOceaniaareincludedand groupssuchasChioggiaandVerona)areseparatedin America is only represented by a single sample of the primary and secondary gene pool, respectively. L. virosa L. The ten species of the tertiary gene pool C. endivia L. is a self-compatible annual, while are mainly distributed in Europe and Asia, and are C. intybus L. is a predominantly self-incompatible represented by low numbers of accessions, or not at biennial to perennial species (Rick 1953; Eenink all, in the ILDB. 1981; Cichan 1983). Inthe international chicory database C.endivia L. Spinach andC.intybusL.arerepresentedwith585(34%)and 947 accessions (55%), respectively. The taxonomic In the International Plant Names Index only 14 status of 184 accessions (11%) was found to be Spinacia species were reported, of which seven were unknown. The primary gene pool also includes the foundtobesynonymsandfourwerereassignedtothe wild subspecies C. endivia L. ssp. pumilum (Jacq.) genusAtriplex.Therefore,thegenepoolofspinachis Cout. (9 accessions) and the wild species C. calvum quite small, consisting of cultivated spinach (Spina- Sch. Bip. that is not represented in the ICDB. The cia oleracea L.) and the wild species S. tetrandra secondary gene pool also includes the wild species Steven ex M. Bieb. and S. turkestanica Iljin that are C. spinosum Jacq. that has also been regarded as a interfertile with each other. Data of 2,017 accessions subspecies of C. intybus L. and that is lacking from were collected from a total of 25 collections located the ICDB. The tertiary gene pool is constituted by mainly in Europe, with the exception of Armenia, C. botae De Flers that occurs in Saudi Arabia and Azerbaijan and the United States. YemenandthatiscompletelyinfertilewithC.endivia The international spinach database includes the L. and C. intybus L.. No accessions of this species data of 1,769 accessions of cultivated spinach orig- occur in the ICDB. Altogether the data of 1,716 inating from a wide variety of geographic regions accessionswerecollectedfrom21collectionslocated (Table 2).Aswasobservedforcultivatedlettuce,the mainlyinEurope.Accessions fromtheUnitedStates majority of accessions of cultivated spinach origi- representlessthan5%oftheICDB.Themainpartof nated from Europe and Asia. The wild species are the ICDB is formed by C. intybus L. var. foliosum poorly represented in the database including only 12 Hegi (36%), C. endivia L. ssp. endivia (32%) and accessionsofS.tetrandraStevenexM.Bieb.and14 C. intybus L. var. intybus (11%) that are mainly of S. turkestanica Iljin. Although for each of these landraces and cultivars of European origin. species a single accession was included with Ger- manyasdocumentedorigincountry,theirdistribution Minor leafy vegetables areas are restricted to Asia (Uotila 1997). The databasecontains222accessions(11%)withmissing The database of the minor leafy vegetables (IMDB) taxonomic data. consists of 163 accessions of artichoke, 356 of 123 988 GenetResourCropEvol(2012)59:981–997 Table2 Numberof S.oleraceaL. S.tetrandra S.turkestanica Unknown accessionsofSpinacia StevenexM.Bieb. Iljin speciesbelongingtothe primaryspinachgenepool Europe 636 1 1 96 includedintheinternational East 79 23 spinachdatabase(ISDB) perreportedgeographic North 82 25 regionwithinthetotal South 158 10 distributionarea West 317 1 1 38 Asia 741 8 5 95 Central 4 4 1 East 133 37 North 20 1 1 10 South 111 0 0 16 Southeast 1 1 West 472 7 0 30 Africa 10 5 Central East 3 North 7 5 South West Americas 14 26 Caribbean Central 1 North 13 26 South Reporteddistributionareas notrepresentedintheISDB Oceania 3 aredenotedby‘0’.Totals Unknown 365 3 8 percontinentaredenotedin Total 1,769 12 14 222 bold asparagus, 77 of lamb’s lettuce, 326 of rhubarb, and which D. tenuifolia (L.) DC. (Table 5B) and D. mu- 1,232 of rocket salad, with a total number of 2,154 ralis(L.)DC.(Table 5C)areusedascultivatedcrops accessions (Table 4). A gap analysis was carried out in addition to their use as wild plants. D. tenuifolia forrocketsalad,whichwasstudiedseparatelyforthe (L.)DC.canalsobefoundinthesecondarygenepool genera Eruca and Diplotaxis that both are used as ofEruca,andD.harra(Forssk.)Boiss.belongstothe salad (Table 5). In the International Plant Names tertiary gene pool of D. muralis (L.) DC. An Index 50 records were found for Eruca and 112 for important position is occupied by many Brassica Diplotaxis. For Eruca, 46 records could be ascribed species as they occur in the secondary and tertiary to synonyms and basionyms or displayed a question- gene pool of Eruca as well as of Diplotaxis, while able status. Only four names are accepted, of which mostoftheotherspeciesofthegenusDiplotaxisand Eruca sativa Mill. together with E. vesicaria (L.) Erucastrumarecloselyrelatedbutnotinvolvedinthe Cav.thatissynonymoustoE.sativaMill.,andEruca gene pool. pinnatifida (Desf.) Pomel, which is sometimes The genera Eruca and Diplotaxis are represented described as a subspecies of both other species in the IMDB with high numbers of accessions represent the primary gene pool of Eruca, whereas belonging to the primary gene pool and with even theothersarewildspeciesthatarenotcrossablewith higher numbers of secondary and tertiary gene pool the species ofthe primary gene pool(Table 5A). For accessions,thelatterbeingduetotheinvolvementof Diplotaxis,14acceptednamescouldberecognizedof Brassicaspecies.Thenativedistributionofcultivated 123 GenetResourCropEvol(2012)59:981–997 989 nal wn o o ernati Unkn 121 3 26 92 7 5 2 2 1 1 1 1 53 184 nt i edinthe Cbotae.DeFlers 0 0 0 d u cl m olin osu o n genep Cspi.Jacq. 0 0 0 y or s c u n s)chi intybvar.know line C.L.un 42 11 17 14 10 3 6 1 1 1 6 5 1 2 61 byvertical usL.var.(Bisch.) bold parated Cintyb.sativumJanch. 41 10 4 27 2 2 6 4 2 20 69 notedin e e s d ( yandtertiaryea Cintybus.L.foliosumvar.Hegi 427 2 2 158 265 5 2 3 6 6 187 625 continentare mary,secondardistributionar Cintybus.L.var.intybus 126 25 7 49 45 3 1 2 1 1 62 192 0’.Totalsper ngtotheprihinthetotal Ccalvum.Sch.Bip. 0 0 0 denotedby‘ choriumspeciesbelongidgeographicregionwit aCendivia.L.pumilumssp.(Jacq.)Cout. 3 3 3 2 1 0 0 3 9 sentedintheICDBare Table3CiNumberofaccessionsofchicorydatabase(ICDB)perreporte CendiviaCendivi..L.ssp.L.ssp.endiviaunknown 32613Europe East2 North4 South1319 West1894 81Asia Central East4 North South1 Southeast West31 6Africa Central East1 North5 South West 2115Americas Caribbean Central North1415 South7 Oceania Unknown186 Total54729 Reporteddistributionareasnotrepre 123 990 GenetResourCropEvol(2012)59:981–997 Table4 Numberofaccessionsofspeciesrelatedtoartichoke(A),asparagus(B),lamb’slettuce(C),rhubarb(D)androcketsalad (E)includedintheinternationalminorleafyvegetablesdatabase(IMDB) Species IMDB Species IMDB (A)Artichoke(Cynara)—163accessions C.albaDC. 1 C.humilisL. 3 C.cardunculusL. 131 C.scolymusL.a 27 C.cornigeraLindl. 1 (B)Asparagus(Asparagus)—356accessions A.acutifoliusL. 8 A.maritimusMill. 1 A.africanusLam. 2 A.officinalisL. 224 A.albusL. 2 A.oligoclonosMaxim. 1 A.aphyllusL. 1 A.pastorianusWebbetBerthel. 1 A.arborescensWilld.exSchult.etSchult.f. 1 A.plumosusBakerb 1 A.asparagoides(L.)Druce 4 A.pseudoscaberGrecescu 1 A.brachyphyllusTurcz. 1 A.racemosusWilld. 8 A.bucharicusIljin 1 A.scopariusLowe 2 A.caspicusHohen. 1 A.scandensThunb. 2 A.cochinchinensis(Lour.)Merr. 3 A.setaceus(Kunth)Jessop 9 A.dauricuslink 3 A.stipularisForssk. 2 A.declinatusL. 3 A.tenuifoliusLam. 1 A.densiflorus(Kunth)Jessop 19 A.umbellatusLink 2 A.falcatusL. 4 A.verticillatusL. 14 A.gonocladosBaker 1 A.virgatusBaker 4 A.laricinusBurch. 4 A.sp. 24 A.litoralisSteven 1 (C)Lamb’slettuce(Valerianella)—77accessions V.carinataLoisel. 1 V.lusitanicaFontQuer 1 V.coronata(L.)DC. 2 V.pumila(L.)DC. 4 V.dentata(L.)Pollich 2 V.rimosaBastard 2 V.eriocarpaDesv. 2 V.sp. 4 V.locusta(L.)Laterr. 59 (D)Rhubarb(Rheum)—326accessions R.altaicumLosinsk. 3 R.palmatumL. 12 R.australeD.Don 6 R.rhabarbarumL. 162 R.compactumL. 3 R.rhaponticumL. 23 R.crispumHort.exG.Don 1 R.ribesL. 2 R.emodiWall.exMeisn.c 1 R.tanguticum(Maxim.exRegel)Maxim.exBalf. 3 R.hybridumMurray 6 R.tataricumL.f. 7 R.macrocarpumLosinsk. 1 R.tibeticumMaxim. 2 R.maculatumC.Y.ChengetT.C.Kao 1 R.undulatumL.d 1 R.maximowicziiLosinsk. 3 R.webbianumRoyle 1 R.moorcroftianumRoyle 1 R.wittrockiiC.E.Lundstr. 2 R.officinaleBaill. 13 R.sp. 72 E.Rocketsalad—1,232accessions BuniaserucagoL. 2 Diplotaxissp. 41 Diplotaxisacris(Forssk.)Boiss. 1 Erucapinnatifida(Desf.)Pomel 13 Diplotaxisassurgens(Delile)Gren. 15 ErucasativaMill. 495 123
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