Dictionary of cultivated plants and their regions of diversity

Transcription

1 Dictionary of cultivated plants and their regions of diversity

2 Second edition revised of: A.C. Zeven and P.M. Zhukovsky, 975, Dictionary of cultivated plants and their centres of diversity

3 'N -'\:K ~ Li Dictionary of cultivated plants and their regions of diversity Excluding most ornamentals, forest trees and lower plants A.C. Zeven and J.M.J, de Wet K pudoc Centre for Agricultural Publishing and Documentation Wageningen ~ T ^/- / - /+<>?- /

4 CIP-GEGEVENS Zeven, A.C. Dictionary ofcultivated plants andtheirregionsof diversity:excludingmostornamentals,forest treesand lower plants / A.C.ZevenandJ.M.J, de Wet. - Wageningen :Pudoc. - Herz,uitg. van:dictionary of cultivatedplants andtheir centresofdiversity / A.C.Zeven andp.m. Zhukovsky, Metindex, lit. opg. ISBN SISO632UDC633 Trefw.: plantenteelt. ISBN Centre foragricultural Publishing anddocumentation, Wageningen,982. Nopart ofthisbookmaybe reproduced andpublished inany form,by print, photoprint,microfilm oranyother meanswithout written permission from thepublisher.

5 Contents Preface 7 History ofthework 8 Origins of agriculture anddomestication ofplants Cradlesof agriculture and regions ofdiversity 2 Chinese-Japanese Region 32 2 Indochinese-IndonesianRegion 48 3 Australian Region 65 4 Hindustani Region 70 5 Central AsianRegion 8 6 NearEasternRegion 87 7 Mediterranean Region 03 8 AfricanRegion 2 9 European-Siberian Region 48 0 SouthAmerican Region 64 CentralAmerican andmexicanregion 85 2 NorthAmerican Region 99 Specieswithout anidentified region 207 References 209 Indexofbotanicalnames 228

6 Preface Theaimofthiswork istogive thereaderquick reference totheregionsof diversity ofcultivated plants. For important crops, regionsofdiversity of related wild species arealsopresented.wild species areoftenuseful sources ofgenesto improve thevalueof crops. Species cultivated primarily asornamentals and timber crops,anduseful lowerplant species arenot included. Taxaarearranged alphabetically firstby family,secondlyby genusand thirdlyby specieswithin genera.themore commontaxonomiesynonyms, as well asthebetterknown (English)names are listed.taxonomy isbased primarily onwillis'sdictionary (966)and thebailyhortorium, HortusThird (976). Somatic chromosomenumbers andgenome formulae arepresented where known. Most ofthechromosomenumbers arederived frombolkhovskikh et al. (969). Where thechromosomenumber could notbe traced, aspacehasbeen leftopen. Chromosomenumberandgenomeconstitutionsmay indicate therelationships of a species. Thework includedmanymore species than wecould know.corrections, criticisms and additions including dataonchromosomenumberwould behighly appreciated.they should besent tothesenior author,instituteofplant Breeding (I.v.P.), Agricultural University,P.O.B. 386, 6700AJ Wageningen, the Netherlands. Wehope that this workmayhelptheplantbreeder toeaseshortagesof food andother agricultural products. Wehopethat it will alsoencourage theestablishment ofnaturalwild plant reserves inanticipation ofneeds forwildgenes. AntonC.Zeven JanM.J. de Wet

7 History of the work FIRSTEDITION In968 Prof.P.M. Zhukovskijpublished a paper 'Newcentres oforiginand new genecentres ofcultivated plants including specifically endemic microcentresofspecies closely allied tocultivated species'.thispaperwas issued inbotanical Journal, Moskov 53: andwas abstracted inplant BreedingAbstracts (-968). I wrote to Prof.Zhukovskijasking whetherhe would prepare anenglish version. Hewroteback thathe waspreparing a booklet inrussianonthe 'Worldgenofund ofplants forbreeding:world gene centresofcultivated plants andtheirwild progenitors', whichwaspublished in970. Thetextwas translated bydr E.E.Leppik, ResearchBotanist of thenewcropsresearchbranch oftheusdepartment ofagriculture, Beltsville, Maryland, who invited me toeditthemanuscript and toseek a publisher. Thepublishers suggested that the workbeextended to includemore cultivated plants. Prof.Zhukovskij agreed tothisproposal and the workhasnow beenenlarged from 700species toabout 2300 species. A.C. Zeven SECONDEDITION InOctober 975my co-author, Professor DrP.M. Zhukovskijdiedat Leningrad after along and fruitful lifewhichhededicated tocultivated plants:theirbotany,their taxonomy and their agriculture and use. He worked almosttothelast dayofhis lifetospread knowledge ofcultivated plants. Afewmonthsbeforehisdeath, hereceived copiesof the first editionofthisdictionary andheexpressed hishappinesswith the work. In 979, Pudoc informedmethat thestock ofthebookwas almost exhausted and that they considered reprinting the work.however asmany scientists are working oncultivated crops, many newdata were available for a newedition. Furthermore,colleagues and myselfhaddiscovered mistakes andomissions and soit was agoodopportunity toprepare a revision. Iamvery gratefulto those whohave suggested additions andimprovements. Tohelp inpreparing arevised edition, Iaskedhelp fromprofessorj.m.j. de Wet, CropEvolution Laboratory,Department ofagronomy,universityof Illinois,Urbana-Champaign, Illinois, United States. Heisanexcellenttaxonomist ofcultivated plants and I wasextremely happy thathetookupthe invitationdespite a busy lifeasscientist.thepresent editionhas greatly benefited fromhisencyclopaedic knowledgeofcultivatedplants. A.C. Zeven

8 Origins of agriculture and domestication of plants INTRODUCTION Manwasnot always afarmer.onlyduring the last fifteen thousand yearsor sohashe learned tosomedegree tocontrolhis food supply.before theadventof agriculture, manwas a hunter and food gatherer.gradually, however, someofthe animal speciesheused tohuntwereprotected, and selected species of foodplantswerebrought intocultivation.plant and animal husbandry were initiated,and theseplants and animals eventually becameso dependent onman that they couldno longer compete successfully with their wild relatives for natural habitats.theybecame domesticated. The antiquity of thisshift in man's activities from food gatheringto food producing isnot knownwith certainty.plant and animalhusbandry were probably well established longbeforenoticeable phenotypic changes occurred inspecies underdomestication, and became preserved inarchaeologicalrecordsofman'shistory. Bethat asit may, manhad abandoned hisnomadic food-gathering way oflifesome 0 000years ago foroneofsedentary foodproduction inseveral partsofboth theold andnew Worlds. In many regions, evenwith a high levelof agriculture, man still gathers wild and semiwild plants or fruits, such as brambles, blueberries, raspberries, mushrooms, herbs forfood, heath for brooms, wood for buildings, fuelorpaper-making,andgrass fordomestic animals. Howevermandoesnot dependon theseplants; heonly collects them foreconomicor recreative reasons. If hedepended onthem,hewould growthemor find a substitute. Somepeoplemay growplantswhile others collect thesame species inthe wild. Wemay askwhyman started tocultivate plants, whyhestarted todoso only 'recently'andwhy only certainplant speciesorvarieties were domesticated. ORIGINSOF AGRICULTURE Muchhas beenwritten aboutman's shift from plant collecting toplant growing.some authorshaveput forward 'deterministic' hypotheses,such as a highermental orsocial level leading tothecultivation of plants, orclimaticchanges causing aprogressive desiccation ofthecountry and enforcing the applicationof artificial methods of food production (Spinden, 97; MacNeish, 964a). Sauer (952),however,thought that agriculture couldnot have originated solely from chronic food shortage,as fourconditionshadto be fulfilled beforeplant oranimalhusbandry couldbe initiated: - Previously acquired skills inother fields tostartexperiments. - Sedentarywayofliving. - Presenceofwooded lands easiertoclearthansavannasor forests.large river-valleys subject toperiodical flooding areunsuitable, becauseman was not abletocontrolfloods. - Amarked diversity ofplant populationsmustbepresent,sothat alarge reservoir ofgenes isavailable forselection.

9 0 ORIGINSOFAGRICULTURE Sauer concluded that the ancestorsof theearliest agriculturistswere relatively prosperous progressive fishermen living in a mild climate along fresh waters. Little isknown about theskills ofthe first farmers,and information on thecorrelation between earlier fixed dwelling and incipient food production islimited.theearliest siteswith year-round occupationwerediscovered in thenilevalleyofupperegypt (5000to0500 B.C.)but they shownoevidenceofplantof animal domestication (Churcher &Smith, 972). Early occupation sites found insouthern Africadating from B.C. (BorderCave in Zululand), B.C. (Howieson'sPoort nearmontagu,southwest Cape Province),42000 B.C. (RoseCottage Cavenear Ladybrand ineasternorange FreeState) belong tothiscategory. But,itisnot clearwhether thesesites were occupied alltheyearround.thebotanicalmaterial associatedwiththem hasnot yetbeen analysed (Dart & Beaumont, 97; Beaumont & Boshier, 972). Sites where agriculture developed firstmusthavebeen inareaswhereplant collectors/hunters/fishermenroamed. Itismost likely thatthey livedin wooded lands for hunting game, ornearwater forfishing.fishing communities led asedentary life. Nomadsroam,but returntositesknown fortheir richness inanimal andplant food.thismayhave led toannual occupation ofsites for afewweeksuntil the food supplywas depleted.on suchsites,thesoil may havebecomebarebecauseofdisturbanceby man; paths,loam pits, graves, dilapidated mudhouses and abandoned compounds.near water,therewould be natural bare landssuch asriverbanks, gravel, rocks, landslides and esturian plains.plantspre-adapted tosuch environmentswould colonize them.around dwellings, manyplantswould derive fromplant parts collected bymanand brought home. Plants adapted todisturbed habitats areweedy inhabitand prefer 'open'rich soils.they growquickly andhave large food reserves that enables them tosurvive adverse conditions.these characteristics make them suitable forcultivation.theymay growwild in mountainsorhills with a wide topographical diversity. Insuch areaswithmanymicroclimates, variantshavemost chance tosurvive.after theyhadmigrated into the artificial habitat, manmayhave found someuseful types among them.someofthe plantsmayhavebeen genotrophes adapting quicker to man-made conditions thanexpected (Zeven, 975). Other sites where agriculture mayhave arisenwouldbemiddens onthe compounds.manypartsofplants (fruits, seeds, tubers, roots)musthave been accidently orpurposefully thrown away.theymust havedeveloped into plantswith aluxurious growthonthese fertileplaces (Anderson, 952; Burkill,952; Chang, 970; Engelbrecht,96; Flannery, 965; Harlan & de Wet, 965; Hawkes, 969). The sequencebywhich crops arosemaybe summarized as follows: -Wild plants collected by man. -Wild plantsmigrated intotemporary orpermanent dwelling sitesofman eitherby accident asgathered plantparts orspontaneously. Thismusthave continued foranextremely longtime. - Plantpreadapted todisturbed habitats colonized areas arounddwellings. Man gathered wanted plant parts from someof theseweedy plants. -Natural selectionwas reduced and selectionpressures introduced foradaptationto man-madehabitats. Decrease invariationwas counteractedby hybridization and mutation,followed by isolation,protection and conscious selectionby man.selected deviants from thewildphenotypeswould survive. Thisstate canbecalledproto-agriculture. - Thedependence ofmanonselected plants increased insuch a way that when demand exceeded availability,man eradicated undesirable plants andstarted to improveuseful plants. Whenmanmoved outside thenatural rangeof a

10 ORIGINSOF AGRICULTURE speciesonwhichhedepended,hewas forced to plant.thusman learnedto retain seeds andother propagules when theplantwastogrowoutside its natural range,topurposefully prepare a habitat inorder toreap a better harvest of thecolonizer,nowturned into crop.this stagerepresentsincipient agriculture.near-eradication hasled toincipient cultivationof Tabernanthe ibogaand Camassialeightlinii. - Cropswere further improved intentionally by cropping methods.thisstage represents effective agriculture. Thechange-over from food collector/hunter/fisher to full-time agriculturistmust havebeen very gradual.once theprocess started, it became practically automatic (Hawkes, 969)or self-generating.this gradual change - including thechange toanimal husbandry -resulted in - less energy toobtainmore food, - peoplebecoming tied tothe(ir)land - sparetime forotherpursuits (MacNeish, 964). By inventing agriculture,mankind gained more from solarenergy.raising crops (andhusbanding animals)areman'smajormeans ofexploiting that form ofenergy (Rappaport, 97). PLANTDOMESTICATION Twokindsoforganisms, weeds anddomesticates, prosper in man-made habitats. Organisms adapted tohabitatsnotnotably disturbed byman are wild.among wild organisms arekinds adapted tovarious degrees ofnaturaldisturbance. Theyoccupy different positions inserai succession.plants atthepioneer end ofsuccession can also invademan'sdisturbed habitats. When thehabitat iscontinously being disturbed by man, however, adifferent setof species becomes established. Plants that arespontaneous and persistant inhabitats that are continually beingdisturbed byman are weeds.there aredegrees of weediness. Wildcolonizerswill invademan-disturbed habitats, but cannot survive continual disturbancesbymanorbynatural means.ifnot continually disturbed, wavesofspecieswill becomeestablished untildynamicbut essentially stable populations are achieved.whenhabitats arecontinually beingdisturbed,only weeds cansurvive spontaneously.oneneed onlydrive alonghighways toobserve thestrict adaptation of weeds to man-disturbed habitats.roadside species rarely formpart ofthe adjacent natural vegetation,evenofcultivatedfields. Plants tendedbyman arecultivated butnotnecessarily domesticated.they maybe wild,weed ordomestic inadaptation.manmaintains cultivated plants intheman-madehabitatbecause they areof sufficient value. Cultivation includes allkindsof agricultural practices,frommerely protectingindividual plants toactual planting or sowing,and tendingofplanted populations. Villages in WestAfrica areoftenbuilt aroundoneor moregiantbaobab trees (Adansonia digitata). These treesprovide shade as well asfruitsfrom which arefreshing drink isproduced.baobab iscultivated but wild,and survives continuous disturbance bymanofitshabitatbecause it isperennial andnotbecause itis a weed.ontheotherhand, weeds can becultivated. Animal fonio (Brachiariadeflexa)is acultivated cereal inwest Africa (Portères, 976). Itis widely distributed across theafrican savannaand oftenharvested as a wild cereal (de Wet, 979). In Angola, however, a weed raceofbrachiaria deflexais often encouraged by local growers ofsorghum (Sorghumbicolor)to invade theircultivated fields,and forms aconsiderable partof theannual cereal harvest (de Wet, 975). Domesticates arestrictly adapted tohabitats specially created forthem byman.they haveevolved underdomestication tothepointwhere theydepend

11 2 ORIGINSOFAGRICULTURE onmanboth forhabitat andpropagation.domesticated seed-plants havelost thenatural ability todisperseseed efficiently natural seeddispersaland domesticated root and tuber crops areoftenpoorseedproducersorarecompletely sterile sexually. Cropsinanearly stage ofdomestication oftenbecome weedy,andmanyweeds represent species invariousstagesofdomestication. Indeed, weeds differ fromdomesticated primarily in degreeofdependenceon man forsuccess inthecontinually disturbed man-madehabitat. Ecological boundaries,as well as boundariesofusefulness to manbetween wild,weed anddomestic taxa arenot always sharp.higgs &Jarman (969, 972)correctly pointout that thewild classoforganisms merge intothe domestic classby acontinuous seriesofdegreesofintimacywith man. The question thus arises whether domestication occurs fromwild acrossbridging weedy races.thiscertainly hasbeenone route.themajority of weeds, however,donot represent astage intheevolution of crops. Origins and evolution of weeds Weeds evolved and arestill evolving in man-made habitats inoneormoreof fourprincipalways (dewet & Harlan, 975). First,they evolve from colonizer speciesby selection foradaptation tohabitats that are continually beingdisturbed.second,weedsoriginate asderivatives ofhybridsbetween wild andcultivated racesofdomestic species. Third,they evolve from domesticates that areabandoned bymanorescapecultivation,by selection for a less intimate associationwith cultivation.fourth, weedy racesofcrops originate as aresultof introgression with related wild speciesofthe domesticate. Amajority of weedshaveevolved fromwild colonizers that invaded the man-madehabitat.theirwild ancestors are aggressive colonizers ofall disturbed habitats,andmany ofthemhavebeendistributed bymanwellbeyond theirnatural ranges.thecommonweedsofnorthamerica areeurasian in origin (Reed & Hughes, 970). Intheirnewhabitats thesespecies are obligate weeds, while intheirnativehabitats races areknown that formpartof thenaturalhabitat.dandelion (Taraxacum officinale),henbit (Lamium apoplexicaule),crabgrass (Digitariasanguinalis)andmanyother urbanweeds arenativesof Europe,andhave been introduced tothenewworld sincethe Fifteenth Century. Weed racesof domestic speciesoriginatebynatural selection as a byproduct ofdomestication,by selection frompartially domesticated or abandoned domesticates, or fromhybridbetweenwild progenitors and their domestic relatives. Weedy hybrid derivatives areknown for most,ifnot all, crops (Harlan,965, 969). There are,for instance, weed sunflowers (Helianthus annuus),weed carrots (Daucus carota),weedmaize (Zea mays)and weedwatermelons (Citrillus vulgaris). Indeed,evendomestic speciesthat areregularly propagated byvegetative means,such ascassava (Manihot esculenta),sweet potatoe (Ipomoea batatas)orpotato (Solanum tuberosum) ihave weed races. Weed racesthat originate as aresult of introgression '(Anderson,949) betweenwild anddomestic taxaareoften strictlyagricultural weeds. They rarely invadenatural habitatsoftheir wild relatives with any success. Adaptative traits acquired under domestication have little selectivevalue inprimary habitats,and geneexchange inthedirectionof wild taxaisnot extensive.gene flow inthe direction ofthe crop, however, maynotonly produce successful weeds, butmay actuallybenefit thecrop (Harlan, 969). IntheNobogameValley ofchihuahua in Mexico, earsof maize showing tracesof.introgression from teosinte (Zeamays ssp. mexicana) are consciously selected by farmers asseed stock (Lumholz,902; Wilkes, 970).

12 ORIGINSOFAGRICULTURE 3 Weed racesofdomestic speciesoftenmimic thecultivated race they accompany in vegetative and inflorescencetraits, except forretainingnatural seed dispersal.suchweeds areparticularly obvious inpearlmillet (Pennisetum americanum)and arewidely distributed with thiscrop across thewest African savanna.scholz (979)suggests that thesemimeticweeds areactually degeneratedomesticates.brunken et al. (977)indicate, however,that although shibras closely resemble pearlmillet ingrossmorphology, in more detailed inflorescencetraits they arewidely variable,and generally intermediatebetween pearl millet and itswild progenitor.weed-cropmimicrymay also become established between unrelated species.it was already knownin Biblical timesthat darnel (Lolium temulentum) grains were difficult to separate from thoseof thewheat orbarley it accompanies as a weed.similarly, false flax (Camelinasativa ssp.linicola)isanobligateweed of flax fields insouthern Russia, where it mimics racesofcultivated flaxingrowth habit, timeof flowering andseed size (Sinskaja & Beztuzhera, 93).These weeds originated from ssp.sativa by selection in fields ofcultivated flax. Cultivated racesofdomestic species canrevertback toweed habit. Many fenceandhedgerowweeds ofeurope arederived from species oncegrownin herborvegetable gardens.thecommonqueenanne's lace,the weedy spring annualof temperateeurasia andnorth America is a weed raceof thevegetable carrot. Incereals, wherenatural seeddispersalmechanismshavebecome lost underdomestication,new dispersal mechanismsneed to become established after thecrop isabandoned as a cultigen.seed dispersal ingrasses commonly occurs as aresult of anabscission layer that formsbelow theglumesor between theglumes and the florets.inthegenus Sorghum disarticulation is commonlybelow theglumes.this istrueofbothwild andweed racesof Sorghumbicolor in Africa.Mississippi chicken corn (ssp. drummondii)in theunited States, however,disperses itsgrainbybreaking oftherachis below the spikelet.this dispersalmechanism evolved inanescapeddomesticateand isalso encountered inpartsofethiopiawherewild relativesof S. bicolor areabsent (de Wet, 978). There areweed sorghums intheunited Stateswith theusualmeansofseeddispersal.theserepresent derivatives of introgressionbetweendiploid cultivated sorghum andtetraploid Johnson grass (Sorghum halepense),anintroduced Mediterranean weed.thisweed was introduced intotheeastern United States some 00years agoandhasbeen extending itsrangeby absorbing genes from cultivated sorghum.the initial hybrid istriploid andpartially sterile. Whenbackcrossed with thecultivated parent,some fully fertile tetraploid offspring areproduced. Diploid offspring arealsoproduced from such introgression.they resemblecultivated grain sorghum inhabit andhabitat preference,andhaveinrecentyears becomeobnoxiousweeds inthecornbeltoftheunitedstates. Weeds arenot restricted totheplantkingdom.numerous animal species are weedy.thehousefly is acosmopolitan 'weed', European rabbitsare 'weedy'inaustralia,and theholybrahmin cow isanobnoxious though reveredweed in Hindu India.Indeed, bydefinition civilized man istheultimate weed, beingobligately confined tototally disturbed man-createdhabitats. Evolutionary dynamicsofplant domestication Domesticates areadapted topermanently disturbed man-made habitats. Plant husbandry musthavebeen initiated toprotect and increasepopulationand density ofselected food-plants. Phenotypic changes that accompanied adaptationunderdomesticationprobably resulted from acombination ofnatural andconsciousandunconscious selection.sauer (952)andAnderson (960), amongothers, suggested thatwild food-plants invaded theman-disturbed

13 4 ORIGINS OFAGRICULTURE habitat, becameweedy aroundwasteplaces,and eventually adapted to cultivation.thisisprobably trueofaggressive colonizers suchastheseveral cultivated species ofchenopodium (Sauer,950)and probably some vegetatively propagated tuber-crops. Partsof edible tubersmusthavemade their way torubbishheapswhere they sprouted toproducevigorous plants,and suchdumpheaps arenotorious habitats for colonizers.man certainly could nothave failed tonotice theseplantsnor failed torealize the convenience inhaving them readily available forharvesting.planthusbandry isan obviousnext step,and domestication ofselected food-plantswould be initiated. Sowingorplanting, however,isonly partofplant domestication,and thesepracticeswereprobably known to man longbefore theadventof agriculture.domestication isinitiated withplanting or sowing, but the domestication process continuesonly aslong asthecrop isgrownone generation after another in habitats specially created by man.food collecting,sowing, plantingor anyotherhusbandry by food gatherers doesnotnecessarily lead to domestication.digging foredibleunderground parts,and stripping plants ofedible leaves, stemsor fruits areasextensively practiced byother animals than man.plants that areuprooted during food-gathering are frequently allowed tosetseedbeforeharvesting (Burkill,952)and,evenif not allowed tofullymature,seedsproduced inprevious generationswill insure continuation of thegenepoolofthe species. Similarly, whenpopulations areharvested fortheirmature fruitsorseeds, sufficient seeds usually escape theharvester toinsure establishment of a newgeneration. Individuals in a wild population donotallmature atthesame timeandinflorescences onthesame individual usually mature atdifferent times. Among themany thousandsofplants regularly harvested inthewild asfood,few havebecome domesticated.evenharvesting followed by sowingorplantingwill notnecessarily lead todomestication.wild cereals,as an example,areoften sownbynomadic food gatherers to increasenatural populations (Steward, 94). Thispractice, however,will onlymaintainnatural evolutionarydevelopment.these food gatherers rarely trytoimprove the habitat,except perhaps forburning theareato be sown,and very rarely repeat the experimentwith thesamepopulation more thantwice. Adaptation to man-made habitats iscomplex.the ability tocolonize disturbed habitats ischaracteristic of wild progenitors ofall seed-cropsas well asother food plantspropagated by sowing.sowing inprepared habitats selects foranincrease incolonizing ability. Itincreases competition among individuals.seedlings that germinate firstwhen conditionsbecome favourable,and those that growmost vigorously when crowded will provide most seeds fromwhich thenextsown generationwillbecome established.with eachsuccessive man-sown generation,selection pressure forsurvival inthe man-madehabitat increases. Domesticated seed-crops therefore lack dormancy, and tolerate crowdingby theirownkind,but arepoorly adapted tocompete withnaturalcolonizers. Harvesting alone selectstoenforcewild-type survivalmechanisms. The deepest rooted individuals oftuber-bearing speciesorindividualswiththe most efficient meansofseed dispersal escape thefood gatherer,and transfertheirgenes forsurvival tothefollowing generations.wild seed-crops areharvested bybeating,swinging of a basket,hand stripping,by uprooting whole plants, orby cutting inflorescenceswith a knifeorsickle forlater threshing. Wilkeet al. (972) pointout thatharvesting inanyway other thanforlater threshingwillnot encourage lossofseed-dispersal ability, even iffollowed insuccessive generationsby sowing.cereals such assauwi (Panicum sonorum)innorthwestern Mexico,orraishan (Digitaria cruciata)in northeastern India (Singh & Arora,972)arecommonly harvested byuprooting

14 ORIGINS OFAGRICULTURE 5 plantsbefore theindividuals are fullymature.they retain efficient seeddispersal mechanisms although thespecies are fullydomesticated inthe sense that cultivated races cannolonger competesuccessfully with their wild progenitors fornatural habitats. Harvesting and later threshing, however,select forindividualswith the mostpersistent floretsor fruitsat maturity.major seed-crops therefore lack theabilityofnaturalseed-dispersal,anddomesticated racesdepend on manboth for asuitablehabitatand seeddispersal (sowing).they arenever successfully spontaneous (weedy)in theman-made habitat formore than afew generations. Food gatherersharvest anarray ofannual and perennial edible plants, seeds and fruits. Relatively fewoftheseweredomesticated.thesitesof early agriculturemust havelimited thenumberofspecies availablefor cultivation.numerousdesirable food plants, however,donot lend themselves todomestication.somehaveexacthabitat requirements and arenot already pre-adapted to man-made habitats.theprogenitors ofall seed-crops are aggressive colonizers,and theprogenitors ofvegetatively propagated crops readily adapt totransplantingby man. Manydesirable food plants give a poor return fortheeffort and for time invested insowingorplanting,and wereprobably soonabandoned bymanascultigens.still otherswereprobably neverbrought intocultivation. Plant domestication issympatricevolution. Thewild progenitors ofcropsare commonly sympatricwith theirdomestic conspecific races. They usually differ strikingly inphenotype and adaptation,but remain sufficiently relatedgenetically tocross andproduce fertilehybrids. Hybridization iscommonand genesareexchanged,particularly inthedirectionofcultivated races. Divergence and introgression areopposing evolutionary processes. Evolutionary divergence dependson isolation.burkill (952)suggests that the initial isolation thatled todomesticationwasprovided bymanwhenhe transported his favourite food plantsbeyond theirnatural ranges.thiswould alsohave forced man to plant,replant seedharvested from populations planted inspecially prepared habitats. However,selection pressuresassociatedwith domestication themselves act asisolatingmechanisms.in domestication,isolationbetweenwild and cultivated racesbecomesdisruptive. Thoday (972) demonstrates thatunderconditions ofdisruptive selection associated with differences inadaptation,population divergencecontinues, even withhybridization. Whendivergencehasprogressed tothestage where parent and daughterpopulations havedistinctly different adaptive norms, interpopulation geneflow iseffectivelyeliminated, sincehybridsbetween themarepoorly adapted toeitherparentalhabitat.onlywith achangein selection pressures doesdivergent evolution cometoanend.racial isolation in domestic species isachieved by gametophytic and sporophytic barriers,anddifferences inflowering time.theprincipal isolating force between domestic races and theirprogenitors, however,isecological adaptation.wild racesofdomestic species donot successfully invade cultivated fields,andcultivated races aretotally adapted to habitats specially created for thembyman. Hybridsbetweenwild andcultivated races,and derivatives ofsuch introgression survive in 'intermediate' habitats as weeds andprovide a bridgebetween them foroccasional gene exchange. Evolutionary dynamicsofplant domestication Crane (950)andMasefield et al. (969) present classesof selection schemes from wildplant tothepresent cultivated crop. Thepossible changes of aspecies caused bydomesticationwerelisted bypolunin (960)and Purseglove (968).

15 6 ORIGINSOFAGRICULTURE Domesticated plants - spread to a morediverse environment andhave a wider geographic range -may have a different ecological preference - may flower and fruit simultaneously -may lack shattering or scattering ofseeds andmay have lostdispersal mechanism completely - may have larger fruits andseeds,and solowerefficiency of dispersal -mayhavebeenconverted from aperennial toanannual - may have lostseed dormancy -may have lostphotoperiodic controls - may lacknormal pollinating organs -mayhave a differentbreeding system (Usually thechange isfrom complete orpartial cross-fertilization topartial orcomplete self-fertilization. Thischangemay result from achange inflowermorphology,or achangefrom self-incompatibility to self-compatibility.) -may lackdefensive adaptation such as hairs, spines and thorns -may lackprotective coverings and sturdiness -may havebetterpalatability and chemical composition,rendering themmore likely tobe eatenby animals -maybemore susceptible todiseases and pests -may develop seedless parthenocarpic fruits - mayhaveundergone selection fordouble flowers, whichmay involveconversionofstamens intopetals -may havebecome sexually sterile and vegetatively reproduced. Speed ofdomestication depends onthe durationof ageneration andintensityofselection pressures. For cereals, agenerationusually takesone year, whereas forvegetatively propagated plants,fast changes arenot usual.braidwood & Howe (962)estimated that all major changes in wheat andbarley underdomesticationhad takenplacewithin 2000 years. Helbaek (966)suggested even 500 years. Somecropswere domesticated forseveral uses.examplesare: - Sorghumbicolor: annual forage grass,syrup sorghum,grain sorghum,broom corn,popping sorghumused forconfectionary, inflorescenses infloral arrangements, - Cannabis sativus:fibre, drugs, oil seeds, - Brassicanapus: rape, swedes, hungary gap kales, oil-seedcolzas, -Brassica campestris:rapeseed, turnip,leafyvegetables, - Brassicaoleracea: vegetable,forage, ornamental, walking stick, construction material, - Helianthus annuus: oil, cattle feed, ornamental, bird food, ceremonies, - Elaeis guineensis: mesocarp oil, kernel oil, wine - Vicia faba:dry seed,freshseed,forage,green manure. This list caneasilybe extended.someplantsmayhavebeen domesticated foroneusethateventually becameobsolete.ifno alternative use were found,itscultivationwould beabandoned anditwould be lost as a cultigen,butmay survive as a weed.cropsmay havebeen abandoned untiluse was found forthem.for instance,severalmedicinal crops andherbs arealso grown asornamentals,asare Viola tricolor anddigitalis purpurea.some formermedicinal species arenowadays grownonly as ornamentals.similarly reverseplantsused inritual becameornamentals.many fencing orhedging plants,grown tostopdomestic animals from running away andwild animals from enteringprotected areas,areused nowadays asornamentals orfor hedges.anderson (960)andChang (970)supposed that the first cropswere not food plants. Anderson suggested thatplantswere firstdomesticated for body paints, hedges, poisons, chewing,fatiguedrugs and ritualpurposes.

16 ORIGINSOFAGRICULTURE 7 Changbelieves theearly domesticated plantswereused formaking containers (bamboo trunks,fruitsof bottle gourd), cordageor as herbs. Theseplants wereuseful and, whenmanbecamedependent onthem,hestarted tocultivate them.most experts oncrophistory,however,believe that food cropsare among the first domesticants.burkill (952)listed thesequence in which hebelieved cropswere domesticated: - cereals - pulses - greens - oil seeds - 'roots' - herbaceous fruits - fibre -woody plants,chiefly fruit trees - various industrialplants. Numerous species ofwild grasses areadaptable todomestication; theyyield well,growgregariously, sothattheircaryopses could becollectivelyharvested,theyhave edible caryopses,their foliage isexcellent forfodder, and thecaryopses aregood tostore. Mandidnotoverlook these advantages ofgrasses (Burkill, 952).Pulsesmusthave followed grasses in domestication.subsequently, several plants collected fortheleaves came into domestication asoil crops.manywoody plantsreceived particular attention. Purseglove (968)stated that cerealswere first domesticated inaridand semi-arid regions, whereas inthewet tropics cultivation started withroot and tuber crops. Archaeological researchwill eventually elucidate the correct sequenceofdomestication. Itmaydiffer from region toregionand different kindsofcropsmayhavebeendomesticated simultaneously. Theplant familieshavenotcontributed equally tothe present supplyof domesticated species. Among the73families (seetable) 48families are represented by only one item,24 by 2 items, 0by 3items,and 4by more than 00items. The familywithmost itemsisthegramineae (379, 5.2% ofthe totalnumberof items); mostof them coming fromregion 8: Africa.This continentis well known forits foragegrasses. The'Leguminosae followwith 337 items (3.5%);Regions 2,7, 8and 0arethe main sources. Gramineaeand Leguminosae contribute aboutone third ofthenumber of items. Rosaceae rank thirdwith 58items (6.3%), most ofthemcome fromregions and9.solanaceaerank fourth with 5items (4.6%), most ofthem comefromregions 0and. Region 2 hascontributed thehighestnumber: 33items (2.5%), closely followedbyregion (295items,.8%), andregions 8and 0 (each 292,.7%). These fourregions contribute almosthalf ofall items.

17 8 ORIGINS OF AGRICULTURE Table. Number of Family Acanthaceae Aceraceae Actinidaceae Agaraceae Aizoaceae Alismataceae Alliaceae Alstromeriaceae Amaranthaceae Amaryllidaceae Anacardiaceae Annonaceae Apocynaceae Aquifoliaceae Araceae Araliaceae Aristolochiaceae Asclepiadaceae Averrhoeaceae Azollaceae Balanitaceae Balsaminaceae Basellaceae Berberidaceae Bignoniaceae Bixaceae Bombacaceae Boraginaceae Bromeliaceae Burseraceae Cabombaceae Cactaceae Campanulaceae Cannabidaceae Cannaceae Capparidaceae Caricaceae Caryocaraceae Caryophyllaceae Casuarinaceae Celastraceae Chenopodiaceae Chloranthaceae Chrysobalanaceae Cleomaceae Combretaceae Compositae Convulvulaceae Cornaceae Corylaceae Corynocarpaceae Crassulaceae Cruciferae Cucurbitaceae Cupressaceae Cyperaceae Datiscaceae Dioscoreaceae items Reg: per ion family 3 per reg ion, per family and per centre S unidentified 2 2 Total

18 ORIGINS OF AGRICULTURE 9 Family Region Dioscoreaceae Dipsacaceae Dipterocarpaceae Ebenaceae Ehretiaceae Elaeagnaceae Elaeocarpaceae Ericaceae Erythroxylaceae Eucommiaceae Euphorbiaceae Euryalaceae Fagaceae Flacourtiaceae Geraniaceae Ginkgoceae Gnetaceae Gramineae Grossulariaceae Guttiferae Hippocastanaceae Hydrastidaceae Hydrophyllaceae Illiciaceae Iridaceae Juglandaceae Labiatae Laminariaceae Lauraceae Lecythidaceae Leguminosae Lemnaceae Liliaceae Limnanthaceae Limnocharitaceae Linaceae Lythraceae Magnoliaceae Malpighiaceae Malvaceae Marantaceae Martyniaceae Melastomataceae Meliaceae Menispermaceae Moraceae Moringaceae Musaceae Myricaceae Myristicaceae Myrtaceae Nelumbonaceae Nyctaginaceae Oleaceae Onagraceae Orchidaceae Oxalidaceae Paeoniaceae Palmae Pandaceae unidenti- - Total fied

19 20 ORIGINS OF AGRICULTURE Family Region uni- Tot identified Papaveraceae 3 Passifloraceae 7 9 Pedaliaceae Pentaphragmaceae Peperomiaceae Perioplocaceae 2 Phytolaccaceae 3 5 Phytolaccaceae 3 5 Pinaceae 2 Piperaceae Pistaciaceae 2 2 Plantaginaceae 2 5 Polygalaceae Polygonaceae Portulacaceae Protaceae 3 3 Punicaceae Ranunculaceae Resedaceae 3 6 Rhamnaceae Rosaceae Rubiaceae Rutaceae Salicaceae 7 9 Sambucaceae 2 2 Santalaceae Sapindaceae Sapotaceae Saurucaceae Saxiphragaceae Scrophylariaceae 3 5 Simaroubaceae 3 S immonds i aceae Solanaceae Sterculiaceae Stilagninaceae Strychnaceae Styraceae Taccaceae Tamaricaceae 2 Taxaceae Tetragoniaceae 5 Theaceae Thymelaeaceae 2 3 Tiliaceae 2 4 Trapaceae 3 3 Tropaeolaceae 3 3 Typhaceae Ulmaceae Umbelliferae Urticaceae Valerianaceae Verbenaceae 2 5 Violaceae 3 Vitadaceae Zingiberaceae Total % of total

20 Cradles of agriculture and regions of diversity Geographic centresofplantdomestication cannotbe foundwithout studying theorigins, hearthsor 'cradles',andspread of agriculture andofdomestic plants. Wild plants arestill entering cultivation,whereas animportant croplike theoilpalm inwest Africa isstill largely semi-domesticated (Zeven, 967; 973). Other examplesofsemi-domesticates aresecondary crops,i.e.crops thatwere firstweeds inprimary cropsbutwere later themselvesdomesticated. Sitesofprehistoric farmshave beendiscovered inthailand, theneareast and Mexico.They showed that incipient agriculture existed inthailand at about 000 B.C. (Gorman, 969), intheneareast atabout B.C. (Cambel & Braidwood, 970)andMexico atabout 6000 B.C. (MacNeish,964a; 964b). Inotherpartsoftheworld nosuchearly siteshaveyetbeen found, and itisgenerally accepted that from thesecradles agriculture spread to otherpartsofthe world.there aregood arguments for independent origin of agriculture in China (Ho, 969).But agriculturemayhavereached China and Japan,andS.E.Asiafrom Thailand, while agriculture probably reached Europe, Africa, W.andS.Asia from thenear East. Alexander VonHumboldtwasprobably the first author torefer totheorigin of crops.in his workessai surlagéographie des Plantes (807) hesaid: "The origin,the firsthomeoftheplantsmost useful to man andwhichhave accompanied him fromremotest epochs,is asecret asimpenetrable asthe dwellings ofallourdomestic animals. Wedonotknowwhat region produced spontaneously wheat, barley,oats and rye.theplantswhich constitute the natural richesofall inhabitants ofthe tropics,the banana,thepawpaw, thecassava,andmaizehaveneverbeen found inwild state" (citedby Hawkes, 970). If he were alivenow, VonHumboldtwould bedelighted tolearnhow muchweknowof theoriginofcultivated plants. Thenext studywasby AlphonseDeCandolle in "Géographie Botanique Raisonêe" (855). Then camecharlesdarwin (868) withhisbook "Variation of animals andplants underdomestication".however Darwinwasnotinterested intheoriginofcultivated plants, but inevolution of animalsand plants, whether in natureorunderdomestication. DeCandolle'sthoughtsontracing theoriginsofcultivated plants were published in882inhisbook OriginedesPlantes Cultivées. Hisworkis still largely upto date (Harlan, 96). Hebased his investigations on: - classical botany (plant geography,knowledgeof adventive and ruderal species, understanding ofhistory ofdevelopment of whole floras), - bio-archaeology (plant remains, pictorial records, especially from Egypt), - palaeontology, - philology. Heconcluded that theregionwhere aspecieswas abundantwasnotnecessarily itscentreoforigin. PerhapsDeCandolle (882) was thefirst to indicate regionswhereplant domestication might have takenplace (Smith, 968): -China -S.W. AsiaandEgypt

21 22 CRADLESOFAGRICULTURE -tropical Asia. InDeCandolle's time,it wasquitenatural to includeegypt as muchofthe knowledgeofplanthistory came from thatcountry. AfterDeCandolle,Nicolai IvanovióVavilov suggested cradlesofagriculture.at theheight ofhiscareerhehad more facilities than anyonebefore (Harlan, 95).Withhis abundant energy,heexploited them tothe full. During thefifth International GeneticsCongress atberlin in926, Vavilov (928) developed histheory ofcentresoforiginorgene centres indicating thatseveral regionsoftheworldpossess concentrations ofvariationof certain cultivated plants and thattheseregionsoverlap forseveralcultivated plants. Theseregions canbe identified bythedifferential Method, described byburkill (952): - Take a map - selectmajor cultivated plants -mark thesites where recognizablebotanical varieties andracesofthese cultivated plants are found.theidentificationofthebotanical varieties wasdoneby investigating the morphology, cytology,genetics and resistance to diseases, pests andunfavourable climatic conditionsof theplants -Where thosemarksoften coincide is a centreoforigin.insuch centres the greatest diversity ofthecultivated crop isobserved. Vavilov concluded that a centreoforigin wascharacterized bydominant alleles and thatthe frequency ofrecessive alleles increased anddiversity decreased towards theperiphery.thecause was inbreeding,geographical isolation and drift. Attheperiphery,secondary genecentresmaydevelop;new areaswith a greatdiversity conditioned by recessive alleles.in926, Vavilov reported thatasiaminorliesintheasiatic, Mediterranean, Balkan andtranscaucasiangenecentresofwheat andother crops.in 93, heextended thisidea bydistinguishing seven genecentres.in 935, heraised thisnumberto eightby splitting SWAsiaintoCentralAsia andthenear East. Later Zohary (970) proposed toreunite them.the centres recognizedbyvavilov were: I. China II. India IIa. Indo-Malaya III. C. Asia,including Pakistan, Punjab, Kashmir, Afghanistanand Turkestan (USSR) IV. NearEast V. Mediterranean coastal and adjacent regions VI. Ethiopia VII. S.Mexico andc.america VIII. S.America (Peru, Ecuador, Bolivia) Villa. IsleofChiloe (Chile). ousregions andoften inareaswith atemperate climate.they areseparated by greatdesertsorlie ondifferent continents. According to Vavilov,agriculture intheseeight regionsdeveloped independently,becauseofthe differences inagricultural methods,implements anddomesticanimals. Vavilovmayhavebeen influenced bywillis' AgeandAreahypothesis (Willis, 922): incomparingwild species with similar modesof dispersal, those with the wider distribution arethe older,and thatthelonger a specieshas beenpresent inanarea,themorediverse thederived speciesand subspecies found there. Vavilov may alsohave been influenced by theagro-

22 CRADLES OFAGRICULTURE 2 3 geographical work ofengelbrecht (Zeven, 973).However time isnot the only factorthat influences thedispersal of aspecies and its increaseof variation. Inthe 930s, Vavilov established an 'ecological passport' forthe accessionsofhislargecollections by sowing them atvarious sites ('geographical sowing')after whichhe estimated: - differences ingrowth during thevegetativeperiod - differences inlength ofthevariousdevelopment stages, including growth rhythm - economic characters,such assizeof fruitsandseeds - vegetative characters - resistance todifferentkindsofdrought - resistance tocold - differences inflowering - resistance to bacteria andviruses - resistance toinsects - ecological growth form:xerophyte, hydrophyte, mesophyte. Thediversity wasenormousbutwithin limits andwith certainregularities. Vavilov discovered parallelisms that areespecially clear for plantsofthe samegeneral group (annuals, herbaceous),characterized by thesame areaof distribution and following thesamegeographical route intheirevolution. Since itseemed asthougheach species differentiated into differentagroecological andgeographical groups, hewas abletoestablish the 'ecological passport'forannual cereals,grain legumes, oil and fibre flax.in 940, Vavilov divided theoldworld (excluding Africa southof thesaharaand tropical Asia)into9areas,eachcharacterized by theplants withessentially thesame 'ecologicalpassport':.syriangroupagricultural Territory:chiefly foothills of Syria,Palestine and Jordania.Characteristics ofcultivated andwild plants:relatively small;with small leaves,flowers and seeds; thin,stiff stems; nonshattering spikes orindéhiscent pods;highmaturing temperature;short vernalization stage. Examples:typesofwild anddomesticated Triticum species; barley; oats; peas; lentils; grass-peas; chick-peas;domesticated flaxandvetch. 2.Anatolian GroupAgricultural Territory: mountainous partsof Turkey. Characteristics: medium-size;thin,stiff stems; medium-sized spikes,fruits and seeds;resistant todrought;shortdevelopment stages; requiringconsiderablewarmth during laststageofdevelopment. Examples:as Group. 3. ArmenianXerophyticMountainGroupAgriculturalTerritory: arid, mountainous steppesof Soviet andturkisharmenia. Characteristics: markedlyxerophytic (smallnarrow leaves); small seeds. Examples:Triticum vavilovii (alsoresistant toshattering,and winterhardy);early dwarf,small seeded, xerophytic chick-peas; alargenumberofrelatives ofdomesticatedwheat; Secalevavilovii. 4.CaucasianMesophyticHigh-Mountain GroupAgricultural Territory:high mountain plateauxofdaghestan andgeorgia,northern America. Characteristics:thin stems; comparatively smooth awns; smallormedium-sizedseeds; short or mediumvegetationperiod. Examples: original ecotypesofsoft wheat;prototypes ofeuropean steppewinter and springbread-wheats; Triticum carthlicum; aspecific groupof barleywithnarrow leaves; many xerophytic and mesophytic typesofsecalemontanum ands.cereale ssp.segetale (many with agreat diversity of redandbrown forms). 5. Daghestan-Azerbaijan FoothillGroupAgriculturalTerritory: coastal regionsofdaghestan and Azerbaijan.Characteristics: mesophytic;long vege-

23 24 CRADLESOFAGRICULTURE tationperiod; tall; leafy; large seeds; rather resistant toleafrust. Examples: giant formsofsoft and durumwheats; barley,rye; peas,vetch, wintertypes ofdurum. 6.TranscaucasianHumidSubtropicalGroupAgriculturalTerritory:West Georgia andblack Seacoast, humid regionsofturkey and S. Azerbaijan (Lenkoran),N.Iran. Characteristics:hydrophytic, tall, leavy;late; ratherresistant tovariouseuropean fungus diseases, Examples:endemic Triticum ssp.such ast. machaandt.timopheevi,and someotherdiploid and tetraploid Triticum types;late typesofprostrate fibre-flax sown inautumn and winter;transitory andvery latespring varietiesof cereals. 7. Iran-Turkestan Group Agricultural Territory: irrigated andunirrigated regionsof Iran, Afghanistan,Soviet CentralAsia (Uzbekistan,Tadjikistan, Turkmenia).Characteristics:lowto medium-high;rathernon-shattering rough spikes;weak stems subject tolodging;slow growth during early stagesof development;drought-resistant during latestages; high temperaturerequirement atmaturity;extremely susceptible toalleuropean fungus diseaseswhen sowninsteppeorwooded steppe regionsofeurope.twosubgroups: a.khivasubgroup:nearmouthofamu-darya river,latevarieties of wheat, barley, flaxandpeas b.kashgar Subgroup:highplateauxnearthe Pamir,extremely cold-resistant varietiesofsoftwheat andrelatively latevarietiesof flax (frequently withwhite flowersand seeds). 8.Pamir-Badakhstan GroupAgriculturalTerritory: Soviet andafghan Badakhstan (PamirAgricultural District),C.and N. Kafirstan,atveryhighaltitudes (even3000 mor more). Includes types fromupperhimalayas andtibet. Characteristics: mesophytic types;ofmedium height; broad leaves;short vegetative period; extremely susceptible toalleuropean fungal diseases. Furthermore agiant typeofryewith large anthers andpollen grains, big kernels and large spikes;liguleless,soft and compactum wheat;largebroadleaved,naked,six-rowed barley; small seeded,early peas, beans and grasspeas. 9. IndianGroupAgricultural Territory:N. India. Characteristics:asthose forthepamir-badakhstan Group; despite thediversity inecological circumstancesquiteuniform;notbushy;thin,stiffstems;smallnarrowleaves; early;short; development stages and rapiddevelopment rhythm; resistantto drought;need high temperatures,especially during last stagesofdevelopment;rapid filling-out ofseeds; small seeds (in cereals,flaxandgrain legumes); spikesof wheat andbarley smooth withnon-shattering grain. In Kashmir, asubgrouphasbeenestablished based on aspecialwheattype characterized bymedium height,thinstems,long narrow leaves,small kernels,rather smooth awns, winterhabit,and lesssusceptibility tobrown rustthantheplantsofgroup 7. (Thereasonwhy Group 8and 9havebeen separated, despite identity ofcharacteristics, wasnot stated.) 0.Arabian MountainGroupAgriculturalTerritory:Yemen, where high-mountainagriculture issubject tothe influence ofthesurroundingdeserts. Characteristics:short spring annualswith extremely rapid growth;thinstiff stems;narrow leaves;relatively largeseeds. Noexamples aregiven..ethiopian (Abyssian)GroupAgriculturalTerritory:Ethiopia and Eritrea. Divided intotwo subgroups; a.varieties sown at beginningofmain rainy season;cosmopolitan, hydrophytic typesoftall large-seeded varietiesofbarley and peas. (Ethiopian wheats,thoughnotnotably cosmopolitan,maybe included here.) b.varieties sownattheendoftherainy season:flax,chick-peas, lentils, beans,grass-peas,andanarabian typeofpea (xerophytic, early,low, smallleaved,small-seeded).origin:veryprobably linkedwith Indiaand mountain-

24 CRADLESOFAGRICULTURE 25 ous Arabia. 2.Chinese-JapaneseGroupAgricultural Territory: China andjapan.very likely,theoriginal materialwas imported fromasiaminorbyway of India severalmilleniaago,butvery important new characters havedeveloped in this group.characteristics:shortdevelopment stages;lowormediumheight; extremely small seeds;rapid fillingof grains. Examples:rapidly filling wheatswith small kernels,awnless or awnletted. 3. MediterraneanGroup AgriculturalTerritory:Mediterranean Area. Characteristics:rather tall,bush;large spikes;long awns; large light-coloured seeds;high yields; usually solid straw,short first development stage; resistant tolow airhumidity,requiringmuchwarmth at maturity; resistant tofungal diseases. (No exampleswere given.) 4.EgyptianGroupAgriculturalTerritory:Egypt.Characteristics: barley and durumwith short stiff stems, medium-sized spikes and short firstdevelopment stages. Similar typeshavebeen foundon Cyprus. 5.SouthEuropeanGroupAgriculturalTerritory:S.France, N. Italy,part of Yugoslavia,Bulgarian coast. Characteristics:tallplants;large leaves; big fruits; highyields. Examples:Triticum turgidum sensustricto,soft wheats;inlombardy,giant formsof oats,chick-peas, horse beans,and a polonicoidwheat havebeen found. 6.EuropeanSteppeGroupAgriculturalTerritory:European Steppe fromtirol totheurals;transferred to N. America,especially tothe prairies. Examples: xerophytic spring and wintertypesofcereals andgrain legumes,thewinter types winter-hardy,the spring typesdrought-resistant; rather small seeds, weak straw, narrow leaves. (Vavilovdivided thisgroup intotwosubgroups, but gave noground for them.) 7.WestEuropeanGroupAgriculturalTerritory: W.Europe including S. Finland ands.sweden. Characteristics:tallhydrophytic plants;thick;stiff stems;largebroad leaves;large dense, highly productive spikes; mediumsizedorlarge grain;ripening late.local varieties have laxspikes,and aretalland early. 8.CentralEuropeanGroupAgriculturalTerritory: forest and wooded steppe ofc. Europe. Characteristics:high-yielding mesophytes. Examples: longfibre flax, high-yielding peas, awnlesssoftwheats. 9.Northern (Boreal) GroupAgriculturalTerritory: N.EuropeanSovietUnion, Siberia, N.Scandinavia.Characteristics: mesohydrophytic;precocity;mediumsized;low warmth requirement;cold-resistant. Examples:self-compatible rye and veryearly typesof foragebarley. Vavilovworked on his concept ofgene centres, modifying it, untilhis death.these agro-ecological groupsneednot coincide exactlywith gene centres.the purpose of allhiseffort is obvious.there aregroupsof plantspossessing certain characteristics notpresent inother groups. So when looking for acertainproperty in a species,itisnotnecessary to study itsentire areaofdistributuion,but itissufficient tolook forit inthegroup(s) where thisproperty has alreadybeen found. Thegenemicrocentres ofharlan (95)are afurtherbreakdown in geobotanicalpatternsofvariation.they aresmall areas in whichevolution is still proceeding at arapid rate.for wheat, Harlan identified three microcentres inturkey.undoubtedly manymoreexist elsewhere. With theintroductionofhigh-yielding foreignwheat varieties,thesemicrocentres are disappearing.harlan also identified genemicrocentres inturkey for a numberofother crops. Hefound that such centres frequently coincide.theymay be intheplainsor in mountainous regions, nearcivilizationorremote from it,inareas with very primitiveormoreadvanced husbandry.

25 26 CRADLES OFAGRICULTURE With increasingknowledge of cultivated,weedy and wildplants,it is becomingevident that someparts ofvavilov's theoryhad to be changed (reviewedbykuckuck, 962). Nevertheless itstill forms agoodbasis tosearch for wild orsemi-wildrelatives ofparticular crops. The largecollectionsmade byvavilov and his introduction of agenetic element ininvestigation still render thesediscussions pertinent. Onepoint in Vavilov's theorywas that a primary centrewasmarkedby a high frequency ofdominant alleles. Gökgöl (94)showed that it was impossible toindicate such acentre for wheat. Brieger (963)did not findone for maize, Zeven (967; 972) not for oil palm and Hanelt (972) not for Vicia faba. Besides ithasbeenpointedout that agreat diversitymay also arise from thevariationof theenvironment. Hence the relationbetweenmountain regions andcentres of origin.such a greatdiversitymay alsodevelopwhen two populationsof a (partial) crossfertilizing species meet,as has been shown forcarthamustinctorus. Vavilov'stheorythatwhere the greatest diversity isfound isalsothecentre oforigin,isnolonger tenable,as wasshownforcrops such as Triticum dicoccum andhordeum vulgare in Ethiopia.Thesecrops show agreat diversity there, butnowild relatives are present. Kuckuck (963)concluded that Vavilov would certainlyhave alteredhis theorieswith presentknowledge. Indeed, he introduced changes as his researchprogressed. Thenumberofcradlesof agriculture hasbeenextensively discussed. Vavilovbelieved inseveral, others suggested two (Sauer, 952); one forthe OldWorld (Burmaand adjacent area)andone forthenewworld (C. America). Darlington (952,969)alsosuggested two:thefertilecrescent of thenear East and Mexico.From thesenuclear areas agriculturewas supposed to have spread across theoldworld and thenew World, respectively.after the introductionof agriculture,newcentres ofplant domestication developed. Thus, Darlington &JanakiAmmal (945)distinguished twelve 'centresof origin':. Ethiopia 2.Mediterranean coast 3. Iran,incl.theCaucasus and E. Turkey 4.Afghanistan 5.Indo-Burma 6.Siam-Malaya-Java 7. China 8. Mexico 9. Peru 0.Chile. Brazil-Paraguay 2.United States As comparedwith thelistofvavilov (926), theyproposed continental Chile insteadof theisleof Chiloe,andadded the Brazil-Paraguay and theunited States centres. They considered themediterranean centres asdiffuse,forcultural rather than botanical reasons. "The Mediterranean, a barrierto wildplants, has been a meansof dispersal and a bond ofunion for plantsof established cultivation". In956, Darlington addedeurope (fornoindicated reason),centralafrica (perhapsbasedonportères'views -see below)andc.america (already mentionedby Vavilov). He alsoswitched inexplicable to 'region',thoughthe captionsof his tableand figure stillmention 'centres'. Thisresultedin the following centres:

26 CRADLESOFAGRICULTURE 27 Gene centresof cultivated plantsofdarlington &JanakiAmmal (945) derived fromvavilov. 2. 2a. 3. 3a S.W. Asia Mediterranean Europe Ethiopia C.Africa C. Asia Indo-Burma S.E.Asia a. 8b. 9. 9a. 9b. China Mexico United States C.America Peru Chile Brazil-Paraguay In950, Portèressuggested independent cradlesof agriculture inafrica southof thesahara, one in E. Africa,theother in TropicalW.Africa.He divided thelatterinto: - thesenegambian 'Subcradle', - thecentral Niger 'Subcradle', - thebenin 'Subcradle'and - theadamawa 'Subcradle'. OtherAfrican cradlesof agriculture are in N.Africa andethiopia.in 962, hechanged hisconceptbydividing and subdividing Africainto: West African Cradle I.Tropical Sector a.senegambian Subsector b.c.niger Subsector c.chad-nilotic Subsector II. Subequatorial Sector B.Nilo-AbyssinianCradle I. Nilotic Sector II.Abyssinian Sector C. E. AfricanCradle D.C.AfricanCradle TheNilo-Abyssinian Cradlecoincides withvavilov'sethiopian andpartof themediterranean Centreoforigin.Thelasttwocradleshavenotbeen further elaborated.portères (950;962)decided on a W.AfricanCradlebecause ofthepresenceof several cropstypical tothat area.inthishewas supported bymurdock (959), whoestablished fourregional agricultural complexes :

27 2 8 CRADLESOFAGRICULTURE.S.W. AsianAgricultural Complex,developed by Caucasoids 2. SE.AsianComplex,developed by Mongoloids 3.C.AmericanComplex,developed byamerican Indians 4.W.AfricanComplex,developed by the W.African Negroes. His decision onanindependent West African Agricultural Complex isbasedon grounds similar tothoseof Portères. Anderson (960)started fromquite another characteristic,individing agriculture into floral andnon-floral seed-crop agriculture inc.africa and a 'pole'of floral agriculture in Indonesia. Hesupposed that thefloral typeof agriculture spread intooceania, China andjapan,indiaandafghanistan,andthenon-floral typeremained in Africa. Thealmost completelack ofinterest inflowers andornamental plants among theafrican peoplesis really astonishing, whereas intheregionofthe floral typeeven thepoorest mangrows someornamentals (Anderson, 960). This isnot duetoanabsence ofornamental species inafrica.many species arecommonly grown elsewhere now. TheclaimofanAfrican cradleof agriculture hasbeen refutedby Wrighley (960), Clark (962), Baker (962), Harris (967)andHarlan (967). Baker (962)summarized hisobjections as follows: - fewof thedomesticates aredefinitly known tooriginate fromw.africa - several ofthedomesticates havesolittledifferentiated from their wild progenitors thatthey cannotbeofgreat antiquity ascultivated plants - ifcultivationhadbeenpractised locally forsevenmillenia,anassociated weed flora rich inindigenous specieswould haveevolved. Harris (967)concluded that typically W.African crops arelocal additions toanintrusive agricultural complex,rather thancompoundsof anancient indigenous one.after the introduction of agriculture inton.africait spread intothesahara.with thedesiccation ofthis areainthethird millenium BC,agriculturebecame established inthesavanna zone stretching from theatlantic tolakechad and further tocapehorn in E. Africa.In thiscentre,themany typically African plants,listedbyharlan (97) were domesticated. Kupzov (955, citedby Darlington 956)showed regionsoftheworld that belong tocertainhearthsof agriculture. He identified ten,grouped into 5 'mainagricultural regions': Hearth of agriculture Main agricultural region Indian Indonesian Chinese C.Asiatic NearEast Ethiopian Mediterranean Nigerian 9. Mexican 0.Peruvian I. II. III. IV. V. Australoid Mongoloid Europoid Negroid Americanoid Except fornigeria they derive from a neolithicstage. Zhukovskjj (965) was the first torefer tosiberia as agene centrefor crops.many Malus, Prunus, Pyrus andother species weredomesticated there. Further,itis arich sourceofwild relativesof these species.

28 CRADLES OFAGRICULTURE 29 Primary regionsofagriculture ( )and regionsofexpansion ( ) of Darlington (956)derived fromkupzov (955) i V? J i s^~^ ^S»^ S Jk-J g 5 JS ** I? il i s. i \ *v ^ ly _à \i 2 «VysP? \\ 8 w ' / JioS/ Megacentres of cultivatedplantsof Zhukovskij (968) In 968, heheralded his idea about 'megagenecentres'.assomany crops originate outsideoneofvavilov's centresoforigin,itwasnecessary to enlarge the areas in which species weredomesticated. These megacentres engulf much ofthe world's landsurface andextend overvast areas. Theyare:

29 30 CRADLESOFAGRICULTURE.China Indochina-Indonesia 8. 3.Australia-New Zealand IndianSubcontinent 0, 5.C.Asia, 6.W.Asia 2. Mediterranean coastal and adjacent regions Africa (8aEthiopia) Europe-Siberia C.America Bolivia-Peru-Chile N.America Megacentres, 2, 4,5, 6, 7, 8a,0and asrecognized byhim,arebased onvavilov's concepts thoughmuch enlarged.zhukovskijproposed asnewones 3 (Australia-New Zealand), 8 (wholeafrica)and 9 (Siberia).Megacentre 2 (N.America)had already beenpresented by Darlington &Janaki Ammal (945) and 9 (Europe)byDarlington (956). Zhukovskij (968) didnot drawboundaries between 2and 4, 5and6. In 970, Zhukovskijmade some amendments;some megacentreswere enlarged andboundariesweredrawnbetween 2and4,and 5and6. Obviously thegreater thenumber of investigated crops,thelarger theareas.therefore Harlan (97) developed the ideaof centres andnon-centres. He suggested that agriculturebegan independently inthree areas and that therewas asystem composed of acentre and a non-centre, many indigenous plant specieswere domesticated, after agriculturewas introduced.harlan (97) preferred the term 'non-centre' becauseofthelarge area involved.hisclassification was: Centre Al. Near-Eastern Bl. Chinese Cl. C.American Non-centre A2.African B2. SE. Asianand C2. S.American Pacific Major cropsdomesticated inthe 'non-centres'may sometimes havespread totheircentres inearly times. Zhukovskij's (970)classificationhas beenused as basis for thefollowing list,thoughpossibly somemegacentres stillhave tobeenlarged.this Centres andnoncentresof agriculturalbeginnings of Harlan (97)

30 CRADLESOFAGRICULTURE 3 holds especially fors.america, where ashiftof theeasternboundary may includebrazil andparaguay and thelandwestof thesecountries,asproposed bydarlington &JanakiAmmal (945). We preferred the term Region to Megacentre. Futureresearchwill showwhether therehave actuallybeen threecradles ofagriculture:. E.Asia (ChinaandBurma) 2. theneareast (Fertile Crescent) 3.C. America, andhow agriculture spread from thesecradlesover the world.

31 Chinese-Japanese Region Vavilov called the Chinese-Japanese Region the 'East Asian Centre of Origin'. For several crops, Japan is a secondary centre of diversity. The Chinese- Japanese Region is the primary region of diversity for several fruit-crops from the Amur-Ussuri Region. Li (966, quoted by Chang 970) divides China into two regions: () N. China with a seed and vegetable agriculture; (2) S. China, which forms a buffer zone between N. China and Region 2, with its vegetatively produced crops. Chang (970) and Harlan (97) suggest an independent origin of agriculture in the N. China Region, which resulted in a wholly original assemblage of cultivated plants. Harlan calls the Bl North Chinese Centre of Origins for Agriculture. The earliest known site of agriculture in China is at Yang-Shao. It is strictly Chinese, with no appreciable foreign influence before 300 BC. It is at present assessed to date back as far as the 4th millenium BC. Older agricultural sites will probably be found in China (Ho, 977). China contributed several major crops. These include several species of fruit trees, Camélia sinensis, Corchorus sinensis, Glycine max, Panicum miliaceum and Setaria italica. It is a secondary centre of diversity for Oryza sativa and other crops. Actinidiaceae ACTINIDIA ARGUTA Sieb.fc Zucc. Tara vine. 2n= c. 6. China, Japan, Korea and the Primorye Territory, USSR. Very frost resistant. Used in crosses with A. chinensis* (Schroeder & Fletcher, 967). ACTINIDIA CHINENSIS Planch. Chinese gooseberry, Strawberry peach, Yang tao. 2n=c. 6, c. 60. W. and C. China. Extensively cultivated in the Yangtse valley and elsewhere for its large, fragrant, juicy fruits. Luther Burbank used it as a pollen donor with the frost resistant A. arguta*. ACTINIDIA KOLOMICTA Maxim. Kolomikta. 2n=c. 2. NE. China and the Primorye Territory, USSR. Very winterhardy. With delicious berries containing much Vitamin C. It is cultivated. ACTINIDIA POLYGAMA Miq. Silver vine. 2n=c. 58, c. 6. N. and W. China, Korea and Japan. A polygamous, trioecious ornamental. In Japan the leaves are boiled and eaten. Alismataceae SAGITTARIA SAGITTIFOLIA L. Arrowhead. 2n=22. Europe and Asia. A herb cultivated in China and Japan for its edible conns. Alliacéae ALLIUM CHINENSE G. Don (syn. A. bakeri Regel). Rakkyo, Ch'iao T'ou. 2n=(x=S) 6, 24, 32. China (Li, 970). Cultivated in China, Japan, California and elsewhere by the Japanese and Chinese. ALLIUM FISTULOSUM L. Welsh onion, Cibol, Stone leek, Spring onion. 2n=6. Siberia and China (Li, 970). Cultivation started probably in N. China. Cultivated in China and Japan. Related toa. altaicum Pall. (2n=6) from N. Mongolia. A. wakegii Araki. (2n=6) and A. microbulbum Prokh. The latter is considered a

32 ALLIACEAE - BALSAMINACEAE 33 hybrid of A. fistulosum* and A. altaicum. Cultivars with blue-green leaves and white bulb are sometimes separated as A. bouddbae 0. Deb. (2n=6) (Purseglove, 972). ALLIUM LEDEBOURIANUM Roem. & Schult. Asatsuki. 2n=6.From USSR to Japan. Cultivated in Japan (Kihara, 969). ALLIUM MACROSTEMON Bunge. Chinese garlic. Chromosome number varying with parts of the plant from diploid (2n=2x=8) to hexaploid (2n=2x=72). Including aneuploids. Ancient Chinese garden plant with very big bulbs.introduced in W. Georgia (USSR) during the Middle Ages. ALLIUM NIPPONICUM Franch. & Savat. 2n=6, 32. Formerly cultivated in Chinabut now it only grows wild there (Li, 969). ALLIUM RAMOSUM L. Chinese leek. 2n=32. N. China and Siberia. Cultivated in N. China.An autotetraploid. Itdiffers from A. porrum*. ALLIUM SATIVUM L. Garlic. 2n=6, genome formula SS.C. Asia (p. 8). Var. pekinense Makino sometimes considered a native of N. China. Cultivated in N. China and Japan (Li, 969). ALLIUM SCHOENOPRASUM L. Chive. 2n=6, 6 + IB, (24, 32). Europe, Asia and N. America. Very polymorphous. Domesticated inussr (region not given) (Kazakova, 97). Cultivated over the whole world. ALLIUM TUBEROSUM Rottl. ex Spreng, (syn. A. odoratum L.). Kui ts'ai, Nira, Chinese chive. 2n=6, 32. Primary centre of origin unknown, as it easily runswild (Jones & Mann, 963). At present from E. Mongolia to Japan,the Philippines and through Thailand to N. India. Its tetraploid type may derive from an autotetraploidization of a diploid species or from an amphiploidization of a hybrid of two diploid species.cultivated in China for its edible leaves and young inflorescenses, and as an ornamental. Amaranthaceae AMARANTHUS GANGETICUS* Anacardiaceae RHUS SUCCEDANEA L. Waxtree. 2n=30. China and Japan. RHUS VERNICIFERA DC. (syn.r. verniciflua Stokes). Varnish tree. 2n=30. China and Japan. It is the source of a varnish, Japanese lacquer. Aquifoliaceae ILEX INTEGRA Thunb. 2n=. Japan. A tree cultivated for its bark which ispounded and used as bird lime. Araceae C0L0CASIA ESCULENTA (L.)Schott var. antiquorum (Schott) Hubbard & Rehder (syn.c. antiquorum Schott,C. esculenta var. globulifera Engl. & Krause). Eddoe, Taro, Dasheen. 2n=2x= 28, 3x=42. SE.Asia (p. 49). Many socalled wild specimens are probably derivatives of run wild plants.from SE. Asia it spread to China and Japan where var. antiquorum developed. In 500 AD. some cultivars are mentioned in China (Li, 969). At present many cultivars are described. Some of them are triploid (Bai et al., 97). Their vernacular names are alsoused for Xanthosoma spp. Dasheen is a corruption of 'eddo de la China'. In Japan a secondary centre of diversity developed. Araliaceae ARALIA CORDATA Thunb. Udo. 2n=28. Japan. Cultivated in Japan as a vegetable (Kihara, 969). PANAX GINSENG C.A. Meyer. Ginseng, Chinese ginseng, Korean ginseng. Ussuri region, China, Manchuria and Korea. Exterminated in the Chinese provinces Shansi and Shensi. There it was cultivated for along time in SE. Manchuria, N. Korea, Japan and also USA and USSR (Baranov, 966). Radix Ginseng comes from the cultivated ginseng, and Radix Ginseng Sylvestris from thewild (Hu, 976). PANAX PSEUDO-GINSENG Wall. San-ch'i.2n= China. Wild and cultivated for its roots used in medicine (Hu, 976). PANAX REPENS Max. (syn. Aralia repens Max.). China and Indochina. A herb cultivated in Yunaran, China and elsewhere for its medicinal roots. TETRAPANAX PAPYRIFERUM (Hook.) Koch. Ricepaper plant. 2n=24. N. Formosa and S. China (Hunan, Szechwan, Yunnan, Kweichow, Kwangsi and Kwangtung provinces). Cultivated in the (sub)tropics as an ornamental (Perdue & Kraebel, 96). Azollaceae AZOLLA PINNATA R. Brown. Water velvet, Water fern, Mosquito fern. 2n=. Domesticated in China and Vietnam (p. 50) and grown for its symbiosis with then-fixing alga Anabaena axollae Strassburger. In China, several cultivars have been developed: Red azolla, Green azolla, Wild azolla-whole river red and Vietnam azolla. See further p. 50. Balsaminaceae IMPATIENS BALSAMINA L. Balsam, Garden balsamine. 2n=4. Indo-Malaya and China. Cultivated in China as a cosmetic plant and elsewhere as an ornamental.

33 CHINESE-JAPANESE REGION Boraginaceae LITHOSPERMUM OFFICINALE L. Var. erythrorhizon (Sieb. & Zucc.) Hand.-Mazz. (syn. L. murasaki Sieb.,L. erythrorhizon Sieb. & Zucc). 2n= 28. Cultivated in N. China and Japan for ared dye. Ssp. officinale is cultivated in Bohemia (P. ). Burseraceae CANARIUM ALBUM (Lour.) Raeusch. White Chinese olive. 2n=. China. Cultivated in S. China and Cochinchina. Cabombaceae BRASENIA SCHREBERI J.F. Gmel. Watershield, Junsai. 2n=28. Asia, Africa, Australia and N. America. Cultivated in Japan as a vegetable (Kihara, 969). Campanulaceae CODONOPSIS TANGHEN Olivier. Szechuan tangsêng. 2n=. China. Cultivated for seng. PLATYCODON GRANDIFLORUM DC. Chinese bellflower, 2n=(6), 8, (28). Cultivated in China and Japan as a medicinal crop. Cannabidaceae CANNABIS SATIVA L. Hemp. 2n=20. Its origin is described on p. 49. In NE. Asia including Japan, Korea and the Peking area exceptionally tall plants are grown (Small et al., 975). HUMULUS JAPONICUS Sieb. & Zucc.2n=6 in male plants and 7 in female plants with an X-Y-Y2 sex chromosome system. E. Asia (Japan, Taiwan, China, Korea and Manchuria). Naturalized in E. N. America, Europe and sporadically elsewhere. An annual aggressive weed (Small, 978). Chenopodiaceae KOCHIA SCOPARIA (L.) Schrader. Summer cypres. 2n=8. S. Europe and Asia. Cultivated in Japan and China as a potherb, and as an ornamental. SALSOLA KOMAROVII (Iljin.) Oka-hijiki. 2n=36. Japan. Cultivated there (Kihara, 969). SALSOLA SODA L. 2n=8, 36. Mediterranean area and Asia. A herb cultivated in Japan. SUAEDA GLAUCA Bunge. Matsuna. 2n= Cultivated there (Kihara, 969). Chloranthaceae. Japan. CHLORANTHUS SPICATUS (Thunb.) Mak. (syn. Ch. inconspicuus Swartz.). 2n=30. China. Cultivated in China, Indochina and Japan as atea aroma. Compositae ARCTIUM LAPPA L. 2n=32, 36. Europe and Asia. Cultivated in China and Japan as a root vegetable and in China and Europe (p. ) as a medicinal plant. ARTEMISIA CAPILLARIS Thunb. 2n=8, (36). E. Asia.Cultivated there and elsewhere as a medicinal plant. CHRYSANTHEMUM CORONARIUM L. Garland chrysanthemum, Crown daisy. 2n=. China. Cultivated in S. China (Li, 970), later in whole China and Japan and elsewhere. Used as a vegetable. CHRYSANTHEMUM SEGETUM L. 2n=8. Europe and Asia. Cultivated especially in China. Leaves are used as a vegetable in the Near-East, Malaya and Indochina. HUMULUS LUPULUS L. Hop. 2n=2x=20 with X-Y sex chromosome system. Var. cordifolius (Miquel) Maximowicz (syn. H. cordifolius Miquel) is wild and cultivated in Japan and China. A perennial mainly propagated by rhizomes (Small, 978a). HUMULUS YUNNANENSIS Hu. 2n=. Dioecious. Yunnan, China. It looks similar to the female plant of H. lupulus and is often confused with it. CHRYSANTHEMUM SINENSE Sab. (syn. Pyrethrum sinense DC). 2n=. China and Japan. Cultivated there as a vegetable. GYNURA PINNATIFIDA DC. (syn.g. japonica Mak.). San ch'i, Tien ch'i. 2n=20. China. A perennial herb cultivated for its medicinal properties. LACTUCA DENTICULATA Maxim. 2n=0, (20). was cultivated in China. It Celastraceae EUONYMUS JAPONICUS Thunb. (syn. E. pulchellus Carr.). 2n=32, S. Japan. A shrub. Cultivated in Spain and elsewhere for rubber. TRIPTERYGIUM WILFORDII Hook.f. 2n=. China, Japan and Taiwan.Cultivated in Chekiang, China as a source of insecticide. LACTUCA INDICA L. Indian lettuce. 2n=8. India, Japan, Philippines and Indonesia. Cultivated in China, Japan and other countries. Many varieties exist in China. PETASITES JAPONICUS F. Schmidt. Fuhi. 2n=87. Sachalin and Japan. Cultivated for its flower buds and leaf stalks (Uphof, 968; Kihara, 969).

34 BORAGINACEAE - CRUCIFERAE XANTHIUM STRUMARIUM L. Cocklebur. 2n=36.In China it was used as vegetable. Now it is a weed in fields and along roadsides (Li, 969). Convoivulaceae CALYSTEGIA SEPIUM (L.). R.Br. 2n=22, (24). Subtropics and tropics. A perennial herb cultivated in China for its roots which are used as a vegetable. IPOMOEA AQUATICA Forsk. (syn. I. reptans Poir.). 2n=30. Throughout the tropics. Var. aquatica is an aquatic plant and a paddy vegetable in S. India and SE. Asia. Propagated by cuttings. Cultivated in fishponds to provide spinach, pig and fishfood (Purseglove, 968). Var. reptans is an upland vegetable in SE. Asia propagated by cuttings or seed. Corylaceae CORYLUS CHINENSIS Franchot. 2n=. China. Cultivated especially in the Szechuan and Yunnan provinces. CORYLUS HETEROPHYLLA Fischer. Siberian hazel nut. 2n=28, N. China, Japan, Korea and the Primorye Territory, USSR. Cultivated in China. The seed has a medium-good taste. The shell is very hard. CORYLUS MANSHURICA Maxim. Manchurian hazel. 2n=, China,Japan and the Primorye Territory, USSR. Cultivated in China. CORYLUS SIEBOLDIANA Blum. Siebold's hazel. 2n=28. Japan. Cultivated there. Cruciferae BRASSICA CAMPESTRIS L. 2n=20, genome formula AA. See p. 50 for the origin of this species. In Region four (sub)species developed. Ssp. chinense (L.) Makino (syn. B. chinensis*) is an annual, fast-growing, precocious,leafy vegetable. The juicy leaves only contain 3.5-4% dry matter. Ssp. nipposinica (Bailey) Olsson (syn. B. japonica Sieb., B. rapa var. laciniifolia (Bailey) Kitam). It has finely dissected deep-green leaves (7% of fresh leaves is dry matter). It grows slowly and has little winterhardiness. Ssp. pekinensis (Lour.) Olsson (syn. B. pekinensis Rupr.) is one of the oldest vegetables in China. It forms large, compact heads. Ssp. narinosa (Bailey) Olsson, Broad-beaked mustard forms atight rosette of small, curley leaves (see B. narinosa*). BRASSICA CHINENSIS L. (syn. B. campestris L. ssp. chinensis (L.) Makino). Chinese cabbage, Celery cabbage, Pak-choi. 2n=20, genome formula AA, Primary centre China where it was domesticated. Cultivated in SE. Asia and elsewhere. It is a vegetable, a salad and an oil crop (var. oleifera). Var. pekinensis (Rupr.) (syn B. pekinense Rupr.), pe-tsai. (2n=20) has blanched heart (see B. campestris*), Var. parachinensis Bailey (Sinsk.) is B. parachinensis Bailey, mok pak-choi. B. japonica* has also the same genome formula. This genome is related to the Ad genome (n=7)of B. adpressa Boiss. (2n=4), the F genome (n=8) ofb. fruticulosa Cyr. (2n=6) and the T genome (n= 0)of B. tournefortii Gouan (2n=20) (Mizushima, 969). Prakash & Narain (97) concluded that the genomes of B. tournefortii are younger than the A genome, and that this species has evolved from the oleiferous plants of the species carrying the A genome. BRASSICA NARINOSA L. (syn. B. campestris L. ssp. narinosa L.). Kou T'sai, Broad-beaked mustard, Chinese Savoy. 2n=20. Only known as a cultigen. Cultivated in E. China esp.around Shanghai» Introduced to Japan and later to the USA (Helm, 963b). Related to B. chinensis* and other A genome carrying diploid Brassica-species. It has entire, deep darkgreen leaves. BRASSICA OLERACEA L. Chinese kale. 2n=8, genome formula CC. Formerly described as B. alboglabra Bailey,but Phelan & Vaughan (976) showed that Chinese kale belongs to B. oleracea, however Snogerup (979) suggested that this vegetable may derive from introductions of B. cretica* ssp. nivea. NASTURTIUM INDICUM DC.2n=.In China it was cultivated as a vegetable. Near Saigon var. apetala Gagnep. is grown as a medicinal crop. PUGIONUM CORNUTUM Gaertn. 2n=. A herb cultivated as a vegetable in Mongolia. RAPHANUS SATIVUS L. Radish, Small radish.2n= 8, genome formula RR. This is avery polymorphic species including biennials with large, fleshy roots and annual forms. Japan and the opposite coastal areas of the mainland are suggested asprimary centre. If so the radish would have derived from the wild R. raphanistrum L. (2n=8)and spread over the Old World probably introgressing with other ecotypes and otherwild species as R. maritimus Smith (2n=8) and R. rostratus DC. Wein (964) suggested that R. maritimus is the parent of radish, while R. landra Moretti (2n=8) is the parent of the small radish. He indicated the E. Mediterranean region as its gene centre (p. 07). In Japan and China large rooted forms: daikon (R. acanthiformis de la Blanch., R. sativus var. acanthiformis Mak., var. macropus Mak., var. longipinnatus Bailey) have been developed, A giant form, the Sakurajuma Daikon (f. gigantissimus), is cultivated in Japan. The roots weigh up to 20 kg. Vavilov (949/50) called these giant cultivars,the champions of plant breeding. Var. oleiformis* Pers., the oil-seed radish is cultivated in China and Japan and also

35 CHINESE-JAPANESE REGION elsewhere. WASABIA JAPONICA (Miquel) Matsumura (syn. Eutremawasabi (Sieb.) Maxim.). Wasabi, Japanese horseradish. 2n=28,Cultivated for its pungent rhizomes. Cucurbitaceae tivated in Japan, China and neighbouring islands. Some taxonomists include this species ind. opposita*. DIOSCOREA OPPOSITA Thunb. (syn.d. batatas Decne.). Chinese yam, Cinnamon vine, 2n=c. 40, c. 44. China.Cultivated in China,S. Japan, Taiwan and the Ryukyu islands. CUCUMIS MELO L. Melon, Muskmelon, Canteloupe. 2n=24. Centre of origin in Africa (p. 24). Secondary centre in China. Chinese and Japanese melons have small fruit and an unpleasant strong taste. The genotypes are convar. chinensis (Pang.) Greb., convar. monoclinus (Pang) Greb., ssp. conomon (Thunb.) Greb. (syn. C. conomon Roxb.), oriental pickling melon. CUCUMIS SATIVUS L. Cucumber, Gherkin, 2n=4. Centre of origin in India (p. 72). Secondary gene centre a mesophytic type with elongated fruits arose in the Far East. Sources of resistance to powdery mildew are found there. HODGSONIA MACROCARPA Cogn. (syn.h. heteroclita Hook.f. & Thomson, Trichosanthes kadam Miq.). Lard fruit.2n=. Cultivated in Yunnan and elsewhere in China for its oily seeds. TRICHOSANTHES CUCUMEROIDES Max. Japanese snake gourd. 2n=44, The chromosomal number suggests and autoploidization or alloploidization, which may have happened in Japan or China where their roots are used toprepare starch, TRICHOSANTHES JAPONICA Regel. 2n=22. Japan. There starch isprepared from the roots. Cyperaceae CAREX DISPALATA Boott. 2n=78, 84. Japan. Cultivated there in rice fields for its leaves which are made into hats. CYPERUS CEPHALOTUS Vahl, (syn. C. natans Buch-Ham.). 2n=. Trop. Asia and Australia. A perennial herb cultivated in the rice fields injapan for mat making (Uphof, 968). CYPERUS GLOMERATATUS L. Wangul. 2n=, Korea. Old fibre crop. Rarer than C. iwasakii*. CYPERUS IWASAKII M. Wangul.2n=. Korea. Old fibre crop. Much more common than C. glomeratatus*. ELEOCHARIS DULCIS (Burm.f.) Trinius (syn. E. plantaginea R.Br.). Water chestnut.2n= W. Africa,upto India, China, Japan, Philippines, Fiji and New-Caledonia. A herb. Cultivated ins. China for its tubers. Probably derived from E. tuberosa Schultes which grows wild in trop. Asia. Dioscoreaceae Dioscorea opposita (Harris, 972) Ebenaceae DIOSPYROS KAKI L.f. (syn.d. chinensis Blume). Kaki, Japanese persimmon, 2n=c , 90. Mountains of C. China. Centre of origin and primary centre of diversity in China. Secondary centre is Japan, Cultivated in Mediterranean countries andusa for its edible fruits. DIOSPYROS LOTUS L. Caucasian persimmon. 2n=30. Subtropical China, in Talysk and W, Georgia, USSR and adjacent Iran (p. 82). All three are countries of primary diversity. Naturalized in the Balkan peninsula and elsewhere. The fruit is excellent when dried. DIOSPYROS MAJOR (Forst.f.). Bakh. (syn. D. andersonii P.S. Green). 2n=, Pacific islands. Cultivated for its fruits which produce oil that can be used to scent other oils. The seeds are edible (Smith, 97). Elaeagnaceae ELAEAGNUS MULTIFLORA Thunb. Cherry eleagnus 2n=. China, Japan and Korea. Cultivated for its edible nuts (Mansfeld, 959). ELAEAGNUS PUNGENS Thunb. 2n=28. N. China and Japan, Cultivated for its edible fruits. ELAEAGNUS UMBELLATA Thunb. 2n=28. China, Korea and Japan. Cultivated for its edible nuts (Mansfeld, 959). DIOSCOREA JAPONICA Thunb. 2n=40. Japan. Cul-

36 CRUCIFERAE - GRAMINEAE 37 Eucomraiaceae EUCOMMIA ULMOIDES Oliver. Gutta percha tree, Tuchung. 2n=34. The upland regions of W. and C. China, Cultivated as a medicinal plant. A polygamous plant, dioecious forms have been found. Euphorbiaceae ALEURITES CORDATA (Thunb.). R.Br. Tung oil tree. 2n=22. Primary gene centre: Japan. Cultivated there and in Taiwan. Its crossability with A. fordii* and A. montana* points to an affinity. ALEURITES FORDII Herasl. Tung oil tree. 2n=22. C. China, between 26 and 33 N. Hybrids may occur between wild and cultivated forms. Secondary gene centre of cultivated tung trees probably in USA (p. 202). Cultivated in other American countries, inussr and Madagascar. With A, montana* natural hybrids may occur. It also crosses with A, cordata. ALEURITES MONTANA (Lour.) Wils. Tung oil tree. 2n=22. China south of 250N. Cultivated in Malawi, Brazil and elsewhere. With A. fordii* natural hybrids may occur. It also crosses with A. cordata*. SAPIUM SEBIFERUM Roxb. Chinese tallow tree. 2n=36. Cultivated in the tropics. Euryalaceae EURYALE FEROX Salisb. 2n=58, Tropical Asia. Cultivated in S, China. Fagaceae CASTANEA CRENATA Sieb. & Zucc. Japanese chestnut, 2n=22, 24. Japan, Cultivated in Japan and in USA for its nuts. CASTANEA MOLLISSIMA Blume, Chinese chestnut, 2n=24. N. and W. China. Cultivated in China and elsewhere for its nuts. It is source of resistance to Endothia parasitica, a fungus causing damage to chestnut (C. sativa) in USA. QUERCUS ALIENA Blume.2n=. Japan, Korea and C. China.Cultivated as food for the Japanese oak spinner (Mansfeld, 959). QUERCUS DENTATA Thunb. Daimyo oak. 2n=24, 48. Japan, Taiwan, Korea and Manchuria and W. and N. China. Cultivated as food for the Japanese oak spinner and also used for timber (Mansfeld, 959). QUERCUS M0NGOLICA Fisch. Mongolian oak. 2n=24. N. China, Korea and N, Japan. Cultivated as food for the Japanese oak spinner and also for timber. Ginkgoaceae GINKGO BILOBA L. Gingko, Maidenhair tree.2n=. E 0 China. Cultivated in China and Japan as an ornamental.the seeds are eaten. Gramineae ARUNDINARIA AMABILIS McClure. Tonkin bamboo, Tonkin cane, 2n=. Only known as cultigen and may have originated in Vietnam. Secondary gene centre: China - Province Guandun and adjacent regions of Guancy. Cultivated for its stems which have many technical properties. Used for hand work, including fishing rods. AVENA SATIVA L. convar. nuda Nord. (syn. A. nuda L.), Naked oats. 2n=42 t genome formula AACCDD, The origin of oats has been described on p. 09. Cultigen of NE. China and Mongolia, the Tibetan-Himalaya highlands, in Turkestan and W. China. It is characterized by 5to 7 florets per flower and by big seeds. BAMBUSA GLAUCESCENS (Willd.) Sieb, ex Munro. (syn. B. nana Roxb.). Hedge bamboo. 2n=72, China and Japan where it is cultivated. In Indochina it is grown as a border plant. BAMBUSA MULTIPLEX Raeusch. 2n=72. Cochin China and Japan. A shrubby,woody grass. Cultivated in trop. Asia for various purposes, BAMBUSA STRICTUS Nees. 2n=70, 72. India (p. 73) and provinces Guancy, Guandun, China and in Hongkong, in tropical evergreen forests. Stems are about equal to those of the best Indian species B. arundinaceae*. Secondary centres: Indochina (p. 53) and S. China. BAMBUSA TEXTILIS McClure. 2n= Guancy, China.. Province BAMBUSA TULDOIDES Munro. 2n=. S. China. Cultivated for various purposes, CHIMONOBAMBUSA QUANDRANGULARIS (Fenzi.) Mak. 2n=48, Continental China and Taiwan. Cultivated in Japan, China and Taiwan and occasionally on the shores of the Black Sea in Caucasus, USSR, ECHINOCHLOA CRUS-GALLI L. Barnyard grass. 2n= 36, (42, 48), 54, (72). Japan and China. Close affinities with the cultivated E. frumentacea*. The hexaploid type, 2n=6x=54 is an allopolyploid with E. oryzicola Vasing., 2n=36 as one parent. According to Yabuno (968) this species has the same genomic constitution as E. utilis (see E, frumentacea*). ECHINOCHLOA CRUS-PAVONIS Schult. 2n=36, 54. Subtropics and tropics. A grass cultivated in Yunnan, China. ECHINOCHLOA FRUMENTACEA (Roxb.) Link. (syn. Panicura frumentacea Roxb.). Japanese millet.

37 38 CHINESE-JAPANESE REGION Billion dollar grass,sanwa millet. 2n=36, 54, (56). China. Primary centre in China. Cultivated in Korea, China,USSR and N. America for human consumption, and as a fodder crop. Closely related to E. crus-galli*. Ohwi and Yabuno separated E. utilis Ohwi & Yabuno (2n=54) from E. frumentacea because they found differences in the genomic constitution, geographical distribution and panicle morphology of these two species (Yabuno, 968). Yabuno considers that the genome formulasof E. utilis and E, crus-galli* are the same and that the genome formulas of E. frumentacea and E. colona are also the same, ELYMUS ARENARIUS L. Sea lyme grass,sand elymus. 2n=56. Europe and Asia. A perennial grass. Cultivated injapan for its culms and elsewhere as adune stabilizer. HORDEUM VULGARE L. ssp. humile Vav. & Bacht. 2n=4, Barley. The origin of barley is described on p. 9.Japan andc. China. Ssp. humile is short,has small leaves, hexastichious ears which are apically awned or awnless. LINGNANIA CHUNGII McClure. 2n=. S. China (Provinces Junjan and Guancy) in tropical evergreen forests. MISCANTHUS SACCHARIFLORUS (Maxim.) Hack. 2n= 4x=76. E. Siberia, N., C. and NW. China, Mongolia, Manchuria, Korea, Japan.Erect types near Hongkong were cultivated as 'arrow plants'. This species probably played a role in the development of Chinese sugar-canes (Saccharum officinarum* Sinense group). It is also used as an ornamental (Grassl, 977). ORYZA SATIVA L, ecospecies japonica (syn. ssp. japonica Kato). Japonica rice, 2n=24, genome formula AA. Indochina. The origin of rice is discussed on p. 74. Ecospecies japonica consists of ecotypes japonica and nuda. Spread to Japan, Korea, N. China, Himalaya region, Egypt, Italy and Spain. PANICUM MILIACEUM L. Proso millet, Shu. 2n=36, (40, 54, 72), Primary centre: N, China. From here it has spread upto Italy. In China it was an important cereal till the introduction of wheat and barley (Li, 970). P. spontaneum Lyssev, (2n= ) might be a weedy type of this species. It grows in Afghanistan, Kazakstan and may be wild in Mongolia (Mansfeld, 959). PASPALUM DISTICHUM L. 2n=40., genome formula X5X5WW, (48), 60. Lowlands of the world. In Japan, it is valued as a forage crop in ricefields. See also p PHYLLOSTACHYS BAMBUSOIDES Sieb. & Zucc. Madake, Giant timber bamboo,japanese timber bamboo. 2n=48. China where its primary gene centre is located. Secondary gene centre is Japan,Many forms occur there under the name 'Madake'. PHYLLOSTACHYS DULCIS McClure. Sweetshoot bamboo, 2n=. C. China. It is cultivated there. The young shoots are edible. PHYLLOSTACHYS HENONIS Mitf. 2n=48, 54. C. China (Szechwan). It is cultivated there. Secondary centre is in Japan. One of the forms developed under cultivation, is black bamboo, kenon bamboo ('Nigra', syn. Ph. nigra (Lodd.) Munro, which is cultivated for its young shoots PHYLLOSTACHYS MAKINOI Hayata. 2n=48. Taiwan. It is cultivated in Japan. PHYLLOSTACHYS MEYERI McClure. 2n=48. China (Chzesian).Apparently cultivated in Japan where a strain with deformed internodes arose. PHYLLOSTACHYS PUBESCENS Mazel ex de Lehaie. 2n=48, Mountains of SE. China. Secondary gene centre in Japan. This species has the largest plants in the genus.used in the timber industry and for its shoots. PHYLLOSTACHYS VIRIDIS (Young) McClure. 2n= China. It is cultivated there for pulping. SACCHARUM OFFICINARUM L. Sugar-cane. 2n various. On New Guinea, the New Guinea Noble canes developed (p. 54) and via Indonesia and Philippines they reached S. China where they hybridized with a4x Miscanthus species (probably M, sacchariflorus*)toproduce the Tekcha clones of the new Sinense group.this must have taken place BC. during the Yueh culture.the Tekcha clones have 2n=8. From these basic Sinense clones,younger clones originated with 2n= They crossed with Chinese S. spontaneum*, 2n=96 to produce still younger Sinense clones, including the 'S. sinense Roxb. clone'. During its migration to India it must have hybridized again with S, spontaneum and probably other grasses. Its migration to India (p. 75)may have been promoted by the red rot disease which affected the S. officinarum clones, which had reached N. India via S. India from Indonesia.This must have taken place ca 250 BC, In N, India, the Sinense group was described as the Pansahi group (Grassl, 977). SETARIA ITALICA L. (syn.panicum italicum L.). Foxtail millet, Liang. 2n=8, genome formula AA. N. China,From here it spread throughout Asia and Europe in prehistoric times.in China it was an important cereal till the introduction of wheat and barley (Li, 970), It derives from S. virides (L.) Beauv. (2n=8), genome formula AA. It is possible that S. pallidifusca Stapf & CE. Hubbard (2n=36)is an allotetraploid with S. italica as one of the diploid parents. SINOBAMBUSA T00TSIK (Sieb.) Makino. 2n=48. Japan (Ryu-Kyu Islands). SINOCALAMUS BEECHEYANUS (Munro) McClure. 2n=

38 GRAMINEAE - LABIATAE 39. S. China. It is cultivated there. SINOCALAMUS EDULIS (Odashima) Kenf. 2n= China. Its shoots are edible, SINOCALAMUS OLDHAMII (Munro) McClure. 2n= Taiwan. SORGHUM BICOLOR (L.) Moench. Chinese Amber Canes and Kaoliang. 2n=20. Sorghum was domesticated in Africa (p. 33). Chinese amber canes are found on the coast of China and Korea, and also in India and Burma. They very likely arrived in China and the FarEast by sea traffic.they are related with the E. African sorgo. After introduction of sorghum from India into S. China it came into contact with S. propinquum (Kunth.) Hitchc., 2n=, and by hybridization kaoliang arose (Doggett, 970). TRITICUM AESTIVUM L. The. ssp. vulgare (Vill.) MK. (syn. T. vulgare Vill.). Bread wheat, Common wheat. 2n=42, genome formula AABBDD. Centre of origin: Transcaucasia and adjacent regions (p. 93). It arrived through Korea and Japan in about 300 BC. (Kihara, 969). Secondary centre of diversity in both countries. The Chinese wheats are characterizedby very broad leaves, with 5to 7 florets per spikelet, with squarehead ears, with daylength neutrality, with fast ripening grain, and by precocious forms. In the mountains of the Sinkiang Province of China very frost resistant wheats developed. Chinese and Japanese wheats cross easily with rye, probably because there was no selection pressure against this characteristic due to absence of Secale cereale-types. Some Japanese and Korean wheats are short and this character was introduced in wheat varieties of Italy, Japan, USA, Mexico and elsewhere. ZEA MAYS L. Maize. 2n=20. Maize was domesticated inc. America (p. 90). Secondary centre: China (Brandolini, 970). The mutant ceratina Collins originated in E. Asia. Cultivated in China, Japan, Manchuria, Burma and the Philippines. ZIZANIA LATIFOLIA Turc. Manchuria water-rice, Gau Sun, Chiaopai. 2n=30, 34. China and adjacent regions. Cultivated as acereal in N. China in ancient times. Later its cultivation moved to the south and its use as acereal gradually decreased. It is cultivated as the vegetable Chiaopai or Gau Sun.The fungus Ustilago esculenta P. Henn. infects the leaf bases, which swell and are eaten. Grossulariaceae RIBES ALPESTRIS Decne.2n=. Himalaya up to 3000 m. In China used as a hedge plant. RIBES LONGERACEMOSUM French. 2n=. W. China. Cultivated there for its fruits. It could be used to improve the strig length of cultivar R, nigrum*. RIBES USSURIENSE Jancz. 2n=. E. Asia. It is a source of resistance to black-current gall mite, Phytoptus ribes Nal., a pest of R. nigrum*. Iteasily hybridizes with R. nigrum. Illiciaceae ILLICIUM ANISATUM L. Japanese star anise.2n= 28. China, Korea and Japan. Cultivated for its medicinal seeds. ILLICIUM VERUM Hook.f. (syn. L. religiosum Sieb. & Zucc). Star anise. 2n=28. SE. Asia. Cultivated for its medicinal fruits.it isnot known wild. Iridaceae BELAMCANDA CHINENSIS (L.) DC. Blackberry lily, Leopard flower. 2n=32. China and Japan. Cultivated there as a medicinal crop. IRISENSATA Thunb. 2n=40. Temp. Asia upto Himalaya. Cultivated in China for its leaves (binding material). Juglandaceae CARYA CATHAYENSIS Sarg. Chinese hickory. 2n~. E, China. Cultivated in Yunnan, China (Mansfeld, 959). It closely resembles C. tonkinensis Lecomte. JUGLANS AILANTIFOLIA Carr. 2n=. Japan. Var. ailantifolia (syn.j. sieboldiana Maxim,), Siebolds walnut. Var. cordiformis (Maxim.) Rehd. (syn.j. cordiformis Maxim.), Cultivated in N, America. JUGLANS DUCLOUXIANA Dode.2n=. Mountain regions of Asia. Cultivated in China. JUGLANS MANDSHURICA Maxim. Manchurian walnut. 2n=32. NE. China and the Primorye Territory, USSR, It is winterhardy and is used as rootstock, Labiatae ELSHOLTZIA CRISTATA Wildd. 2n=. China and Japan, A perennial plant introduced in other parts of Asia, Europe and America as an oilseed crop. MENTHA ARVENSIS L. var, piperascens Mai. Japanese mint. 2n=96, genome formula R a R a SSJJAA. Japan. Cultivated in Japan, China and Brazil. It is the main source of menthol. Var. agrestis (2n=72, genome formula R a R a SSJJ) isvery likely a hybrid of var. piperascens and M. japonica Makino (2n=48). It is a source of early maturity and rust resistance (Ikedaet al,, 970), M. arvensis is one of the parents of M. x gentilis (see M. cardiaca*).

39 CHINESE-JAPANESE REGION PERILLA ARGUTA Benth. 2n=. China and Japan. Cultivated for various purposes. It is sometimes included inp. frutescens*. PERILLA FRUTESCENS Britt. (syn.p. ocymoides L.). Suttsu, Perilla. 2n=38, 40. Himalayas, China and Japan,Primary centre : China,The red-leaved strains (syn.p. crispa (Thunb.) Nakai) are sometimes used asornamentals and the green-leaved plants (P. crispa var, ocymoides) for the seed oil. Cultivated in China, Japan and Korea as a drug plant (Li, 969)and formerly as a leafy vegetable. STACHYS SIEBOLDII Miq. Chinese artichoke. Japanese artichoke, 2n=,It isone of the few tuber crops domesticated in China. Cultivated there, Japan, Belgium and France. Laminariaceae LAMINARIA JAPONICA Aresch, Haidai.2n= China. Cultivated as a food plant and as a source of iodine. Lauraceae CINNAMOMUM CAMPHORA (L.) Nees & Eberm. Camphor tree. 2n=24. China, Japan and Taiwan. Cultivated in these countries and other tropical countries. CINNAMOMUM ZEYLANICUM Breyn. 2n=24. Sri Lanka and SW. India, Cultivation started insri Lanka in 770. Cultivated now in several countries. Leguminosae ASTRAGULUS SINICUS L, (syn. A. lotoides Lam.). Genge, 2n=6. China. Cultivated there on rice soils as a soil improver. CANAVALIA GLADIATA (Jacq.) DC. var. alba (Makino) Hisauchi (syn. C, ensiformis (L.) DC. var. alba Makino). Siro-nata-name. 2n=22, Cultivated in Japan (Sauer, 964). Characterizedby white seeds, GLEDITSIA JAPONICA Miq. 2n=. Japan. Cultivated for fruit juice, which is used for washing. GLYCINE MAX (L.) Merr. Soya, soybean. 2n=40. China. Available evidence suggests that soya was domesticated around the th Century BC. in E.N. China,Since the stcentury AD.,it became widely introduced and land races evolved in China, Korea, India and other parts of Asia. The species is today widely cultivated in most agricultural regions of the world (Hermann, 962; Hymowitz, 970). GLYCINE SOJA Sieb. & Zucc. Wild soya. 2n=40. Widely distributed inn., NE.and C. China, adjacent USSR, Korea, Japan and Taiwan. The weedy G, gracilis Skvortz. occurs where G. soja and G. max are sympatric and represent derivatives of hybrids between wild and cultivated soybean (Hymowitz & Newell, 980). LESPEDEZA CUNEATA (Dum. Cours.) G. Don. (syn. L. sericea Benth,), Perennial lespedeza. 2n= (8), 20. E. Asia. Cultivated in the USA for erosion control, LESPEDEZA STIPULACEA Maxim, Korean lespedeza. 2n=20. E. Asia. Cultivated in the USA forhay making. LESPEDEZA STRIATA Hook. Common lespedeza, King grass. 2n=22. E, Asia. Cultivated in the USA in pastures and for hay making, MUCUNA HASSJ00 (Piper& Tracy) Mansf. (syn. Stizolobium hassjoo Piper & Tracy). Yokohama bean. 2n=. Japan and China.Cultivated there and in the USA for the seeds. PHASEOLUS ANGULARIS Wight. Adzuki bean. 2n=22. Primary gene centre C. China. It is unknown wild. Cultivated in China, Manchuria, Korea and Japan. Secondary gene centre Japan. PHASEOLUS VULGARIS L. var. chinensis (syn. Ph. chinensis Hort, ex Schur,). Asparagus bean, 2n=22, Its origin is discussed on p. 94. It reached China from the Americas after Columbus' voyage. In China asecondary gene centre arose. The main character, a parchment -like layer in the pod wall was lost and the pod became edible. PUERARIA THUNBERGIANA (Sieb. & Zucc.) Benth. Kudzu. 2n=24. China and Japan, Cultivated as a cover crop, green manure and hay crop,in New Guinea and New-Caledonia as a tuber crop (p. 58). VICIA UNIJUGA A.Br. (syn. Orobus lathyroides L.). Two-leaved vetch. 2n=2, (24, 36). E, Siberia, Manchuria and Japan. Occasionally cultivated. WISTERIA BRACHYBOTRYS Sieb. & Zucc. 2n=6. A vine. Cultivated for its fibrous bark, Liliaceae ANEMARRHENA ASPH0DEL0IDES Bunge. 2n=22. N. China. A medicinal plant occasionally cultivated, FRITILLARIA VERTICILLATA Willd. 2n=24. Var, thunbergii Baker. Cultivated in China as a medicinal plant. LILIUM AURATUM Lindl. Gold band l i l y, Yamayuri. 2n=24. Japan. Cultivated there for its large bulbs (Kihara, 969). LILIUM CORDIFOLIUM Thunb. 2n=24. Japan. Cultivated there for its starchy bulbs.

40 LABIATAE LILIUM LANCIFOLIUM Thunb. Oni-yuri. 2n=24. Japan Cultivated there for its bulbs (Kihara, 969). LILIUM MAXIMOWICZII Regel. Ko-oni-yuri. 2n= 24. Japan. Cultivated there as afood crop (Kihara, 969). LILIUM TIGRINUM Ker-Gawl. Tiger lily. 2n=(24), 36. China. Cultivated there and in Japan for its edible bulbs. OPHIOPOGON SPICATUS Kunth. 2n=. China. A herb cultivated in Chekiang as amedicinal plant. Magnoliaceae MICHELIA FIGO (Lour.) Spr. (syn. M. fuscata Andr.). Banana shrub. 2n=38. China. Cultivated there for its banana-scented flowers used for scenting hair oil. Malvaceae ABELMOSCHUS MANIHOT* ABUTILON AVICENNAE Gaertn. (syn. A. theophrasti Medic.). Button weed, Chinese jute, Velvet weed, Butter print chingma. 2n=42. Cultivated in China (many local varieties), USSR and elsewhere for its fibre called jute or Indian mallow. GOSSYPIUM ARBOREUM L. Tree cotton. 2n=26, genome formula A^A^. Arose in India (p. 77). Race sinense, Chinese cotton, Nanking cotton, developed in E. China. It is the earliest fruiting form of this species. Ithas short lint and requires along daylength. First cultivated as an ornamental.at present it has a low breeding value. HIBISCUS SYRIACUS L. Rose of Sharon. 2n=80, 80-84, 90, 92. China and Taiwan. Cultivated first in China as a hedge plant and later elsewhere as an ornamental. MALVA SYLVESTRIS L. High mallow. 2n=42. Probably the early vegetable K'uei mentioned in Chinese literature.at present a weed in China (Li, 970). Cultivated in Europe as a medicinal crop and ornamental. MALVA VERTICILLATA L. (syn. M. crispa L., M. mohileviensis Graebn.,M. pamiroalaica IIj.). Mallow, 2n=c. 84,c, 2. E. Asia. It was an early Chinese domesticate there. About 500 AD. it was there an important vegetable with several varieties like purple and white stemmed, large and small leaves. During the 7-0th Century the cultivation in China declined. In 848 itwas only observed in remote areas. Introduced to Japan, where it is a weed now (Li, 969). Also in W. Asia and Europe. Cultivated in Europe as a medicinal crop. This plant has often been described as M. crispa being a cultigen of M. verticillata. Menispermaceae COCCULUS THUNBERGII DC. 2n=. A woody vine cultivated in Japan forbasket making. Moraceae BR0USS0NETIA KAZINOKI Sieb. 2n=26, 39. A tree cultivated in Japan and Korea for its bark which is asource for paper production, BROUSSONETIA PAPYRIFERA (L) Vent. Paper mulberry. 2n=26. China and Japan. In the Far East this tree isused for making paper and barkcloth (Purseglove, 968), MORUS ALBA L. White mulberry. 2n=28. China. Cultivated there and elsewhere for its leaves eaten by silk worms, for its fruits and for paper making. It is often planted as aroadside tree, cv Makado has 2n=3x=42. Musaceae MUSA BASJ00 Sieb. & Zucc. 2n=22. Japan. Species of the Eumusa section.used for making fibre. Myricaceae MYRICA RUBRA Sieb. & Zucc. (syn. M. nagi Thunb,). Chinese strawberry tree, Ioobai, Yama momo. 2n=6. Cultivated in China for its fruits. Oleaceae FRAXINUS CHINENSIS Roxb. 2n=92, 38. W. and C. China. Especially var, acuminata Lingelsh. (syn.f. koehneana Lingelsh.) is cultivated as ahost plant of the insect Coccus pela for wax production. LIGUSTRUM JAPONICUM Thunb. Japanese privet, 2n=44. A shrub. Cultivated in Japan for its seeds. LIGUSTRUM LUCIDUM Ait. 2n=46. A tree cultivated in China as ahost plant of the insect Coccus pela for wax production. LIGUSTRUM OVALIFOLIUM Hassk. 2n=46. Japan. A shrub widely planted for hedges in Europe and elsewhere. It may have run wild there. OSMANTHUS FRAGRANS Lour. 2n=46. Himalaya, China and Japan. A tree cultivated in E. Asia for its very scented flowers used to aromatize tea. Palmae TRACHYCARPUS F0RTUNEI (Hook.)H. Wendl. Windmill palm, Chusan palm. 2n=36. China. Often planted in E, Asia for its fibres.

41 CHINESE-JAPANESE REGION tivated in China (Uphof, 968). PRUNUS DAVIDIANA (Carr.) Franch. (syn.p. persica var. davidiana Maxim., Persica davidiana Carr.). Chinese wild peach. 2n=6. Vladivostok, SW. through Charbin upto Dacin- San and Ala-San (China). Cultivated as an ornamental. It is frost,drought and heat resistant. Probably it is valuable as a rootstock for Amygdalus persica* and Prunus domestica* (Zylka, 970). PRUNUS PSEUDOCERASUS Lindl. (syn.p. paniculata Edwards). 2n=32. W. Hupei, China. A tree cultivated there for its fruits. PRUNUS SALICINA Lindl, Chinese plum, Japanese plum. 2n=6. Primary gene centre: the forests of N. China. Second gene centre: Japan. Cultivated in Japan,China and also in California. It crosses easily with the North American plum species, A new stone fruit 'cherry plum' was derived from crossing the wild P. cerasifera* with P. salicina var. Burbank, Because of its winterhardiness this fruit tree can be grown where apricot will not fruit. PRUNUS SARGENTII Rehd. Sargent cherry, Mountain cherry, 2n=6. Japan, Manchuria, Korea and rarely in the Far East of USSR, Used as an ornamental. It is frost resistant and fast growing. The fruits are not very palatible. PRUNUS SIMONII Carr. Apricot plum, Simon plum. 2n=6 B Primary centre:probably N. China and Japan. No wild plants are found. Also cultivated there. Crossing with P. triflora Roxb.* (2n=6) from the same area has led to the development of cultivars which are especially grown in N. America. PRUNUS TOMENTOSA Thunb. (syn.p. trichocarpa Bunge). Nanking cherry, Manchur cherry, Chinese bush cherry. 2n=6, N. and W, China, Japan, Himalaya, Turkestan and in the Far East of USSR. Cultivated as a fruit tree and ornamental in the Far East of USSR, N. China and Japan. PRUNUS USSURIENSIS Kov. & Kost, (syn.p. triflora Roxb. var. mandshurica Skvoro.). Ussurian plum, 2n=6. Cultivated or run wild in Manchuria, E, of USSR and for some years also in Siberia and N. Kazakhstan (Zylka, 970). It is a source of good fruit flavour and cold resistance. This species is sometimes considered as a subspecies of P. cerasifera*. PYRUS BETULAEFOLIA Bgb. 2n=34. N. and C. China. Used as rootstock. Resistance against scab (Venturia) is found in this species. PYRUS BRETSCHNEIDERI Rehd. 2n=34. Hupei and Shansi, China, Primary gene centre: N. China. There it was domesticated. It is the commonest cultivated pear in this region. The fruits are characterized by hard, crisp, white sweet flesh. PYRUS CALLERYANA Dene. 2n=34. China, Japan and Korea. Primary gene centre: the Tsinling mountain range, China. It is used as rootstock. PYRUS PHAEOCARPA Rehd. 2n=34. N. China. PYRUS PYRIFOLIA (Burm.) Nakai (syn. P. serotina Rehd.), Sand pear. 2n=34. Primary gene centre: the highlands of N. and C, China. Var. culta (Mak.)Nakai is drought resistant, but not very winterhardy. The leaves reach 5cm in length.the fruits are outstanding for their preserving quality. It crosses easily with the wild European pear,p. communis*. Prunus davidiana PYRUS USSURIENSIS Maxim. Ussuri pear. 2n=34. SE. Siberia upto the Manchurian-Chinese area. Primary gene centre : NE. China and the Primorye Territory, USSR, along the Ussuri river. The ancient var. culta is widely distributed over N. and C, China. It is adapted to cold, dry regions. It is the most winterhardy wild pear. As this species originated outside the Chinese centre proper it has no resistance to scab and other diseases. It probably played a part in the origin of P. communis*.

42 SAPINDACEAE ROSA MULTIFLORA Thunb. 2n=4. E. Asia. Used as rootstock, ROSA RUGOSA Thunb. 2n=,. China and Japan. The rose hips are used by the Ainu.Used as an ornamental and in hedges as one parent to breed for rootstocks. RUBUS ILLECEBROSUS Focke.Strawberry raspberry, Balloon berry. 2n=4. Japan. Cultivated in N. America, RUBUS PHOENICOLASIUS Maxim. Wine raspberry. 2n=4, Japan and N. China. Cultivated for its fruits and as an ornamental, RUBUS PUNGENS Oldhami. 2n= breeding with R. idaeus*. Rubiaceae. Korea.Used in GARDENIA JASMINOIDES Ellis (syn. G. florida L.). Cape jasmin. 2n=22. Probably S. China. Cultivated in E. Asia for perfumery oil, Rutaceae CITRUS ICHANGENSIS Swing. 2n=8. S. of Tsin 'lin range in W., C. and SW. China, It is very frost resistant. Therefore it has been crossed with cultivated Citrus species.used as a rootstock. CITRUS JUNOS Tan. Juzu. 2n=8. Han'su province, China at 372 m altitude. It is frost resistant and therefore used as a rootstock. Its fruits are big but sour. It was already known in the time of Confucius (about 2500 years ago). Because of introgression, characters of 2 Japanese and 2 Chinese wild Citrus species are recognized in juzu. CITRUS RETICULATA Blanco. (C. nobilis Andr. non Lour.). Mandarin, Tangerine. 2n=8. See for its possible origin p. 63.Secondary centre: Japan. New types such as the Satsuma (var. unshiu, syn. 'C, unshiu') and the Natsudaudau ('C. natsudaidai')arose through bud mutation and spontaneous hybridization. Satsuma tangerine (unshiu mikan) is probably a derivative of So-kitsu or Man-kitsu. CLAUSENA LANSIUM (Lour.) Skeels. Wampi. 2n= 8. It is a small-fruit tree of S. China. Cultivated in several tropical countries. FORTUNELLA CRASSIFOLIA Swing. Meiwa kumquat, 2n=8. China and Japan. It is occasionnally cultivated, FORTUNELLA HINDSII (Champ.) Swing. Wild kumquat. Hongkong kumquat. 2n=8, (36). The Tziulun mountains of Hongkong. It has no great value. The 4x form originated spontaneously (Cameron & Soost, 969). FORTUNELLA JAPONICA (Thunb.) Swing, (syn. Citrus japonica Thunb,). Round kumquat, Marumi kumquat. 2n=8. Primary centre Japan. This.fruittree is unknown in a wild state. It is occasionally cultivated. FORTUNELLA MARGARITA (Lour.) Swing, (syn. Citrus margarita Lour.). Oval kumquat, Nagami kumquat. 2n=8. Japan, Cultivated in Japan, China and Florida, USA. Fortunella species cross easily with each other and with Citrus species. PONCIRUS TRIFOLIATA (L.) Raf. Trifoliate o- range. 2n=8, (36), N. China, also primary centre. This area is its centre of diversity. Cultivated in China as an ornamental, and in USSR and Japan it is used as a rootstock. Hybrids with sweet orange (Citrus sinensis ) are citranges used as rootstocks, with sour orange (C. aurantium*) are citradias, crosses of citrange with kumquat (Fortunella margarita*) resulted in citrangequat. Other hybrids are citrandarin (P.trifoliata x C. reticulata) and citrangedin (citrange x calamondin (see C. reticulata)). TRIPHASIA TRIFOLIA (Burm.f.)P. Wilson (syn. T. aurantiola Lour.,T. trifoliata DC.). Lime berry, Trifoliate lime berry. 2n=. Centre of origin possibly China (Mansfeld, 959). Cultivated for its edible fruits. ZANTHOXYLUM PIPERITUM DC. Japanese prickly ash, Japan pepper, Sanshô. 2n=70, China and Japan. Cultivated there (Uphof, 968; Kihara, 969). ZANTHOXYLUM SIMULANS Hance (syn. Z. nitidum DC., Z. bungei Planch.). 2n=32. China. A shrub cultivated in C. and S. China for its seeds. This seed is a source of Chinese pepper. Sapindaceae LITCHI CHINENSIS Son. Litchi, Lychee, Leechee. 2n=28, 30. S, China. Leenhouts (978) described three subspecies: ssp. chinensis, ssp. philippensis (Radlk.) Leenh. and ssp. javensis Leenh. Ssp. chinensis is the cultivated litchi. It was first mentioned ca 00 BC. when Emperor Wu Ti tried to introduce it from N. Indochina into S. China. It is grown now in N. India, S, Africa, Florida and Hawaii, There are many races. Those from China can be grouped as Water litchi and Mountain litchi. Water litchis are grown in the lowland and have smooth fruits, while the Mountain litchi is used as stock or as a fruit-tree inhilly regions. Its fruits are smaller and more prickly. Leenhouts (978) thinks that the Mountain litchis are similar towild types. Ssp. philippensis is found in the Philippines, where it isnot cultivated; its fruits bear sharp pyramidal warts and are not eaten. It isnot a wild type (Leenhouts, 978). Ssp javensis has fruits like the original ssp. chinensis. It isoccasionally grown in Indochina and W.

43 2 Indochinese-Indonesian Region Vavilov called the Indochinese-Indonesian Region thetropical Asian Centre of Origin.Darlington (956)andLi (966cited by Chang, 970)divided this regionintos. Asia: Burma, Thailand and Indochina,and SE. Asia: Malayan Peninsula and the MalaysianArchipelago.Li described anagriculture based mainly onvegetatively propagated crops. Harlan (97)considered this regionas B2Southeast Asian and SouthPacific noncentre,asagriculture may havebeen introduced there. Theoldest known agricultural remains fromregion 2come from Spirit Cave 60km N.of Mae Hongson, NW.Thailand (Gorman, 969). This agrees with Sauer's (952) conclusion that theoldworld centre ofdevelopment of agriculture wassituated inthenw.part ofregion 2 (aboutpresent day Burma). Spirit Cavewas inhabitated from ca.0000to BC. Solheim (972) proposed thathorticulture mayhave developed in BC.and there was a further domestication ofplants and alsoof animals in BC., resulting in large-scale agriculture and animal husbandry. Another early archaeological site isatnonnok Tha, NE. Thailand.It dates from around 5000 BC. InSpiritCave remainsof Prunus, Terminalia, Areca, Viciaor Raphia, Lagenaria and Trapa, andin another layer Piper, Madhuce,Canarium,Aleurites and Arecawere found, and in athird layercanarium, Lagenaria andcucumis (Gorman, 969). Further research isneeded tosupport thetaxonomieidentifications.for instance Schultze-Motel (972)doesnot accept thatvicia faba couldhavebeenpresent. Chang (970) used thepresence of Vicia faba, Lagenaria, Trapa andcucumis asproof that theseplants were actually cultivated. The region isimportant forcrops such as bamboos,tropical fruit trees, ginger,cocos nucifera,colocasia esculenta,dioscorea spp., Musa spp., wild andweedy Oryza spp., Piper spp.,andsaccharum officinarum.

44 AGAVACEAE - ARACEAE 49 Agavaceae CORDYLINE TERMINALIS Kunth. Palm lily. 2n=ca 52. SE. Asia, Australia and most of Oceania. Cooked roots will keep for several weeks. During the 7th and early 8th Century, roots were fermented and distilled to produce spirits (Barrau, 96). One clone is extensively cultivated for the fleshy roots that contain laevulose (Ezumah, 970). Amaranthaceae AMARANTHUS GANGETICUS L. 2n=34. Asia. Cultivated in India, Malaya, China and Japan as a spinach.see also A. mangostanus*. AMARANTHUS MANGOSTANUS Juslen (syn. A. tricolor L. var. mangostanus Thell.) 2n=32. Trop. Asia. Cultivated as a pot-herb. In A. tricolor are sometimes included var. gangeticus and var. tricolor (syn. A. melancholicus L.). See A. gangeticus*. AMARANTHUS PANICULATUS L. 2n=32. Used as a pot-herb and grain crop in SE. Asia. It might be conspecies with A. cruentus* (Sauer, 950) or a synonym. Anacardiaceae BOUEA MACROPHYLLA Griff. 2n= of the wet tropics of SE. Asia.. A fruit tree MANGIFERA CAESIA Jack. 2n=40. Primary centre Indonesia.This fruit tree is cultivated there and elsewhere. MANGIFERA FOETIDA Lour. Bachang mango. 2n=40 Indochina and Malaysia. Cultivated in Java. STELECHOCARPUS BURAHOL (Bl.) Hook.f. &Thorns, (syn. Uvaria burahol Bl.). Burahol.2n= Malaya and Java. Cultivated in Java for its fruits. Apocynaceae ERVATAMIA CORONARIA Stapf, (syn. Tabernaemontana coronaria Willd.,T. divaricata R.Br.). Grape jasmine. 2n=22. Malaya. Cultivated there for various purposes. Araceae ALOCASIA INDICA (Roxb.) Schott. 2n=28. Centre of diversity SE. Asia. Cultivated there for its stem which is eaten and as an ornamental. Introduced to other countries such as India. This species is sometimes included ina. macrorrhiza*. AMORPHOPHALLUS CAMPANULATUS (Roxb.) Blume. Elephant yam. 2n=28. SE. Asia. Cultivated in C. and E. Java and India (p. 7). AMORPHOPHALLUS HARMANDII Engl. & Gehr.2n=. Occasionally cultivated in Tonkin. AMORPHOPHALLUS RIVIERI Dur. 2n=24, 26, 32, 39. Indochina. Var. konjac (Schott) Engl. Philippines. Cultivated in China and Japan (Mansfeld, 959). COLOCASIA ESCULENTA (L.) Schott. Dasheen, Taro, Cocoyam. 2n=2x=28, 3x=42. SE Asia or NE. India (p. 7). It was introduced into China and Japan, where var. antiquorum developed. It is also grown in the Mediterranean region, W. Africa, the Pacific islands, New Guinea, Samoa and New Zealand. In SE. Asia, var. esculenta (syn. C. esculenta sensu MANGIFERA 0D0RATA Griff. Kurwini mango.2n=. Malaysia. Cultivated in Java. It is closely related to M. indica* (Rhodes et al. 970). SEMECARPUS ANACARDIUM L.f. (syn. Anacardium orientale L.). Cashew marking nut tree, Oriental cashew nut. 2n=60. Trop. Asia and Australia. Cultivated in the tropics. SPONDIAS LAOSENSIS Pierre. 2n=. Trop. Asia. A tree cultivated for its fruits. SPONDIAS PINNATA (Koen. & L.f.) Kurz (syn. S. mangifera Willd.). Hog plum. 2n=. Trop. Asia. A fruit tree whose flower clusters are also eaten. Annonaceae CANANGA ODORATUM Lamb. Ylang-ylang. 2n=6. Malaysia. Cultivated there and in other countries for the flowers which are a source of essential oils. Distribution of somatic chromosome numbers for taro (Colocasia esculenta)(yen & Weeler,968) 0=3x.

45 50 INDOCHINESE-INDONESIAN REGION. Probably the Pacific islands. Perhaps it is a cultigen of N. pinnatum* or a closely related species (Mansfeld, 959). NOTHOPANAX OBTUSUM Miq. (syn. Panax obtusum Blume, Polyscias obtusa (Bl.) Harms). 2n= Probably SE. Asia. Cultivated in Java. NOTHOPANAX PINNATUM Miq. (syn. Panax pinnatum Lam., Polyscias rumphiana Harms). 2n=. SE. Asia and New Guinea. Cultivated for its leaves. Asclepiadaceae GYMNEMA SYRINGIFOLIUM Boerl. Sajor pepe. 2n=. Cultivated in Malaya as a vegetable. Averrhoaceae AVERRHOA BILIMBI L. Bilimbi. 2n=22, 24. Malaya. Its fruits are very acid. AVERRHOA CARAMBOLA L. Carambola. 2n=22, 24. Indonesia. Cultivated for its fruits. Some varieties have been developed. Colocasia esculenta ^^*rs0%&e t stricto, C. esculenta var. typica A.E. Hill) developed. Many wild populations are probably dérivâtesof escapes from cultivation (Purseglove, 972). Yen & Wheeler (968) showed that the first introduction of taro into the Pacific, including the Philippines, composed of 2n=28 cultigens and 2n=42 cultigens were introduced later into Indochina-China-Ryu Kyu-Japan area and into Timor-New Caledonia-New Zealand area. Chromosome counts are needed of Indonesian taro cultigens. CYRTOSPERMA CHAMISSONIS (Schott) Merr. (syn. C. edule Schott,C. merkusii (Hassk.) Schott.). 2n=, Indo-Malaysian area. Introduced into many Pacific islands. Cultivated for its tubers (Purseglove, 972), PISTIA STRATIOTES L. Water lettuce. Tropical duckweed. 2n=28. Subtropics and tropics of theold and New Worlds. A floating plant. Cultivated in Java in fishponds for edible shrimps that live below the plants (Uphof, 968). Araliaceae NOTHOPANAX FRUTICOSUM Miq. (syn. Panax fruticosum L., Polyscias fruticosa (L.) Harms). 2n=22, 24. SE. Asia and Polynesia. Cultivated in Java for its roots and leaves. NOTHOPANAX GUILFOYLEI Merr. (syn. Panax guilfoylei Cogn., Aralia guilfoylei Bull.).2n : = Azollaceae AZOLLA PINNATA R.Brown. Water velvet, Water fern. Mosquito fern. 2n=. Domesticated in Vietnam and China (p. 33) for its symbiosis with the N-fixing alga Anabaena azollae Strassburger and therfore used to enrich soils with N, especially of rice fields. For this purpose, it has already been grown for several centuries. In Vietnam, the cv.beo Giong was developed. It dies at the time of the maximum tillering of the rice plants and its nutrients become available at this crucial stage. Other cultivars have been developed in China (p. 33). Water fern may also be used to suppress pest weeds and as fodder. In some areas like New Zealand, it is itself a pest weed, blocking water channels and pipes (Lumpkin& Plucknett, 980). Basellaceae BASELLA RUBRA L. Indian spinach. Ceylon spinach. Malabar nightshade. 2n=44, 48. Probably S. Asia (Winter, 963), Cultivated as a vegetable, now throughout the tropics. Formerly it may have been used for dyeing. The commonest synonyms are B, alba L, (2n=48), with white flowers, and B. cordifolia Lam. (2n= ) with heart-shaped leaves. Bombacaceae CEIBA PENTADRA Gaertn,, var. indica (DC.) Bakh. Kapok tree,silk cotton tree. 2n=72, 80, 88. Secondary gene centre: SE. Asia. Introduced from Africa (p. 23) probably via India. Cultivated in SE. Asia (Zeven, 969). The Indonesian cultivar Reuzenrandoe (giant kapok) bears some characteristics of the var, caribaea*.

46 ARACEAE - CYPERACEAE 5 DURIO KUTEJENSIS (Hassk.) Beccari. Lai. 2n=. Along the foothills of the central ranges of Borneo. Ithas now spread to E. and N. of Borneo, DURIO OXLEYANUS Griffith. Kerantongan. 2n=. Malaya, Sumatra and Borneo,The fruits are eaten and seeds are dispersed around the (temporary) settlements. Occasionally cultivated. Other wild species of which the fruits are eaten are D. graveolens Beccari, tabelak (2n= ), D. dulcis, lahong (2n= ), and D, grandiflorus (Mast.) Kostermans & Soegeng, durian munjit (2n= ). D. graveolens grows wild in Borneo, Malaya and Sumatra,D, dulcis in Borneo. DURIO ZIBETHINUS Murray. Durian, 2n=56. W. Malaysia, Unknown wild. Cultivated throughout SE, Asia, W. Irian, Moluccas, Celebes,S, Philippines, Introduced to Indochina, Thailand, Burma, Sri Lanka and elsewhere. It is often semi-cultivated, i.e. semi-wild trees result from dispersal of seeds. In this way, durian groves have arisen, Boraginaceae TOURNEFORTIA ARGENTEA L.f. (syn. Messerschmidia argentea (L.f.) Johnston), Velvet leaf, 2n=, Trop. Asia, A shrub cultivated for its leaves which are used as smoking tobacco. Burseraceae CANARIUM COMMUNE L. Java almond. 2n=. Moluccas. Cultivated inmany other tropical countries. The kernels are eaten and oil is extracted from them. Sometimes they are planted as shade trees and as ornamentals. CANARIUM MOLUCCANUM Blume.2n=. Moluccas, New Guinea and W. Polynesia. Cultivated in Malaysia. CANARIUM OVATUM Engl. Pili, Pili nut.2n= S. Luzon (Philippines). The kernels contain 70-80% pili-nut oil. CANARIUM PIMELA Koenig. Black Chinese olive. 2n=. E. Asia. Cultivated ins. China and Cochin China. Combretaceae QUISQUALIS INDICA L. Rangoon creeper.2n= SE. Asia. A woody vine. Cultivated as an ornamental, vegetable and as an anthelmintic. TERMINALIA BELLIRICA (Gaertn.) Roxb. (syn. Myribalan bellirica Gaertn.). Bellirica, Terminalia. 2n=26, 48,India and Malaysia. Cultivated as asource of myrobalan which is used for tanning leather, for black dye and for making ink. TERMINALIA CATAPPA L. Indian almond, Myrobalan, Almendro. 2n-24. Trop. Asia, N. Australia and E, Polynesia.Widely cultivated in the tropics.the kernels contain about 55% oil. Used for manufacturing edible fats, cosmetics and pharmaceutic preparations.used for timber ; leaves and bark are used for preparation of medicines.the fruits are edible. TERMINALIA CHEBULA Retz. 2n=4, 24, 26, 48. India to Malaysia. Compositae BLUMEA BALSAMIFERA (L.). DC. 2n=20. Himalaya, India, Malaysia, S,China and Taiwan, Cultivated in Java as a medicinal crop. BLUMEA MYRIOCEPHALA DC. 2n=. India, Vietnam, Malaya and Indonesia. Occasionally cultivated in Vietnam. ENHYDRA FLUCTUANS Lour. (syn. E. helonchu DC, Hingtsha repens Roxb.), 2n=22. India, Indochina, Thailand, China and Indonesia. A water plant occasionally cultivated in Cambodia and Malaya for its leaves. EUPATORIUM STOECHADOSUM Hance, 2n=40. Vietnam. Cultivated there. PLUCHEA INDICA Less. 2n=20. Cuttings of this shrub are planted ashedges in Indonesia. Young leaves are eaten as a vegetable or used to prepare a medicinal tea. SPILANTHES PANICULATA Wall, ex DC.2n=. SE. Asia.and New Guinea, Cultivated as a vegetable or salad, VERNONIA ANTHELMINTICA Willd. Kinka oil iron weed. 2n=20, 54. Trop, Asia, Itmight be a source of epoxy fatty acids. Convoivulaceae IPOMOEA MAMMOSA Chois.2n=. Abouana. Philippines. Cultivated in Indochina. Formerly it was erroneously believed that this species was the ancestor of I, batatas*. Cucurbitaceae BENINCASA HISPIDA (Thunb.) Cogn a (syn. B. cerifera Savi). Wax gourd,white gourd. 2n=24, Java. Cultivated throughout Trop, Asia (Purseglove, 968). It was already mentioned as a vegetable in China in 500 AD. (Li, 969), TRICHOSANTHES ANGUINA L. Edible snake gourd. 2n=22. Trop. Asia from India (p.7*3) to Australia. Cyperaceae FIMBRISTYLIS GL0BUL0SA (Retz.) Kunth, 2n= Trop. Asia, Sri Lanka, India, Malaysia and

47 52 INDOCHINESE-INDONESIAN REGION Mariannes,Cultivated onmalaysia for matmaking etc. LEPIRONIA ARTICULATA (Retz.) Domin. (syn. L. mucronata Rieh.). 2n~. SE. Asia, Malaysia, Australia and Fiji. Cultivated in Indonesia. SCIRPODENDRON GHAERI (Gaertn.) Merr. (syn.s. costatum Kurz.). 2n=. Trop. Asia, Samoa and Australia. Cultivated in Sumatra for mat making. Dioscoreaceae DIOSCOREA ALATA L. Greater yam, Water yam, Winged yam, Ten months yam. 2n=(20), 30, 40, 50, 60, 70, 80. SE. Asia, in the Assam-Burma region it is the cultigen of D. hamiltonii Hook., 2n=, or D. persimilis Prain & Burk, 2n=, or a similar species (Burkill, 935). Some types have been described as D u atropurpurea Roxb., 2n=, and D. purpurea Roxb., 2n=.It is virtually sterile (Ayensu& Coursey, 972). Dennst, D. triphylla L.). 2n=40, (80). India (p. 73) and SE. Asia. Closely related to the African D. dumentorum (Coursey, 967). DIOSCOREA NUMMULARIA Lam. 2n=. SE. Asia. Cultivated there and in Indonesia and Oceania. It closely resembles D, cayenensis*. DIOSCOREA QUARTINIANA A. Rich. 2n=. Throughout Trop. Asia. Cultivated in E. Nigeria (Coursey, 967). DIOSCOREA PENTAPHYLLA L. 2n=40, 80, 44,c. 44. SE. Asia. Cultivated throughout Indonesia and the Pacific islands. Dipterocarpaceae SHOREA STENOCARPA Burck. 2n=. Malaya. Cultivated for its seeds, asource of a Borneo tallow. Ebenaceae DI0SPYR0S DISCOLOR Willd. (syn.d. blancoi How.). Malobo, Velvet apple. 2n=, Malaysia and Philippines. Occasionally cultivated for its edible fruits. MABA MAJOR Forst.f. 2n=. Cultivated for its fruits on the Friendship Islands. Elaeocarpaceae ELAEOCARPUS FLORIBUNDUS Blume.2n=. From Bangladesh to Java. Cultivated in Bengal and Assam for its fruits. Euphorbiaceae Dioscorea alata ( ), D. esculenta ( D. hispida ( ) (Harris, 973) )and DIOSCOREA BULBIFERA L. Potato yam, Aerial yam, Bulbil-bearing yam. 2n=36, 40, 54, 60, 80, 00. Trop. Asia and Africa.Possibly it was domesticated in Asia as well as Africa (p. 25). Cultivated in Trop. Asia, Africa, Oceania and the W. Indies. The tubers and bulbils are edible. The African form (p. 25) has been described asd. latifolia Benth., 2n=. There are many types which often have been described as species e.g. D. heterophylla Roxb., 2n= DIOSCOREA ESCULENTA (Lour.) Burk. Lesser yam, Asiatic yam, Potato yam, Fancy yam, Chinese yam. 2n=40, 60, 80, 90, 00. Indochina. Cultivated in S. China,and later throughout the tropics. Most flowers are male (Ayensu & Coursey, 972). DIOSCOREA FLABELLIFOLIA Prain. & Burk. 2n=. Malaya. Occasionally cultivated there. DIOSCOREA HISPIDA Roxb. (syn. D 0 hirsuta ALEURITES MOLUCCANA Willd. Tung oil tree.2n= 44. Autotetraploid. Indonesia. Itswild parent is not known.crosses between this species and A. montana did not succeed (Wit, 969b). ALEURITES TRISPERMA Blanco. Tung oil tree, Banucalang. 2n=22. Philippines. Cultivated there, in Malaya and Indonesia. Crosses with A. montana did not succeed (Wit, 969b). BACCAUREA DULCIS Muell.-Arg. Tjoopa.2n= Malaya and Indonesia.Cultivated there for its fruits (Uphof, 968). BACCAUREA MOTLEYANA Muell.-Arg. Rambai.2n=. SE. Asia. Cultivated there for its fruits (Uphof, 968). BACCAUREA RACEM0SA Muell.-Arg. 2n=. Malaysian Archipelago. Cultivated there for its fruits. GL0CHIDI0N BLANCOI Lowe.2n=. A tree cultivated in Far East and Philippines for the young leaves and shoots (Terra, 967). HEVEA BRASILIENSIS (Willd.) Muell.-Arg. Bra-

48 CYPERACEAE - GRAMINEAE 53 zilian hevea, Para rubber tree. 2n=36. Amazon basin (p. 69). A secondary gene centre : Malaya. Domesticated in SE. Asia at the end of the 9th Century (Purseglove, 968). MANIHOT ESCULENTA Crantz. Cassava. 2n=36. America (p. 70 and 90). Secondary centre of diversity in Indonesia. PLUKENETIA CORNICULATA Smith, (syn. Pterococcus corniculatus Pax & Hoffm.). Painapaina. 2n=. SE. Asia. Cultivated as vegetable. PHYLLANTHUS DISTICHUS (L.) Muell.-Arg. (syn. Ph. acides (L.) Skeels. Otaheite gooseberry. 2n=26, 28.India and Madagascar. Cultivated in the tropics for its fruits (Purseglove, 968). PHYLLANTHUS EMBLICA L. Emblic, Myrobolan. 2n=28, 98. Trop. Asia. Cultivated in the Old and New Worlds for its fruits (Uphof, 968). SAUROPUS ALBICANS Blume (syn. S. androgynus Merr.). 2n=. Cultivated as a vegetable in SE. Asia. TRIGONOPLEURA MALAYANA Hook.f. Gamber ooran. 2n=. Malaysian Archipelago 0 Cultivated there for its leaves which substitute for Uncaria gambir*. Flacourtiaceae FLACOURTIA RAMONTCHI L'Hér. Botoko plum, Madagascar plum, Governor 7 s plum, Ramontchi. 2n=22. Malaya and Madagascar. Cultivated in the tropics for its fruits. FLACOURTIA RUKAM Zoll. & Mor. Rukam. 2n= Malaysia and Philippines. A tree cultivated for its fruits. HYDNOCARPUS ALCALAE C. DC. 2n=. Philippines. Cultivated for its seeds which are a source of oil used to cure leprosy. HYDNOCARPUS ANTHELMINTHICUS Pierre ex Lanessan. 2n=24. Indochina and Thailand. Cultivated in many tropical countries for the seeds which are a source of oil used to cure leprosy. HYDNOCARPUS KURZII (King) Warb. 2n=. Ssp. kurzii in Burma Highlands and Assam. Ssp. australis in Burma Lowlands and N. Siam. Ssp. kurzii is cultivated inmany trop, countries for the seeds which are a source of oil used to cure leprosy. PANGIUM EDULE Reinw. Cultivated in Java. Gnetaceae Pangi. 2n= GNETUM GNEMON L. Bulso. 2n=. From Assam to Malaysia and Fiji. Var, ovalifolium (Poir.) Bl. is considered the wild type while var. Malaysia. gnemon is the cultivated type planted in Java. Introduced to Java, Sumatra and elsewhere. Cultivated in SE. Asia for its seeds and leaves. A large dioecious shrub. Gramineae ANDR0P0G0N ACICULATUS Retz. (syn. Chrysopogon aciculatus Trin.). 2n=. The tropics. Cultivated invietnam for its roots which contain Chiendent grenille à brosse (Uphof, 968), BAMBUSA ARUNDINACEA (Retz.) Willd. Spiny bamboo. 2n=70. Primary centre: India and Burma (p. 73). Secondary centre: Malaysia and E.Java. BAMBUSA CORNUTA Munro. 2n=. Java. A woody grass cultivated for its tender shoots, which are used as a vegetable. BAMBUSA SPINOSA Roxb. 2n=. Philippines and Indonesia. A woody tall grass cultivated as a timber bamboo and also for its young shoots used as a vegetable. BAMBUSA STRICTUS Nees. 2n=70, 72. India and Burma (p. 73). Secondary centres: Indochina and S. China (p. 37). BAMBUSA TULDA Roxb. 2n=. India, Burma (p. 73) and Tahiti. Secondary centre: Java, BAMBUSA VULGARIS Schrad. ex Wendl. 2n=72. Probably Malaysia or India. It is unknown in the wild. Cultivated in the tropics for its young shoots and for its stems. COIX AQUATICA Roxb. 2n=0. S. Asia. Grown as fodder in India, also collected as a wild cereal in Orissa, India. COIX GIGANTEA Koenig ex Roxb. 2n=20, 40. S. Asia. Involucres are used asbeads and poultry are fed on the grains in NE. India, COIX LACRYMA-JOBI L. Job's tears, Adley.2n= 20, genome formula BB. S. Asia. Wild races have indurated involucres of various colours used as beads. Domesticated races have soft involucres and are widely grown as cereals in NE. India and SE. Asia. Cultivated kinds are often referred to as C. Ma-yuen Romanet (Arora, 977; Jain & Banerjee, 974; Kaul, 973). CYMBOPOGON CITRATUS (DC.)Stapf (syn. Andropogon citratus DC.). Lemon grass. 2n=40, 60. Probably Malaysia or Sri Lanka. Unknown wild. Cultivated in S. Asia, Indochina and elsewhere for its lemon grass oil. CYMBOPOGON NARDUS (L.) Rendle (syn. Andropogon nardus L.). Citronella grass.2n : =20, (40, 60). Cultivated in Indonesia, Sri Lanka and elsewhere for its citronella oil. There are two types of oil: ()Sri Lanka type obtained from var, lenabatu which is cultivated in S. Sri Lanka; (2) Java type obtained from var.

49 54 INDOCHINESE-INDONESIAN REGION mahapengiri (syn. C. winterianus Jowett, 2n= 20). The latterwas introduced into Java from Sri Lanka early in20th Century. It is now widely distributed throughout the tropics.the wild type in Sri Lanka has a different oil composition (Wijesekera et al., 973). DENDROCALAMUS ASPER (Schult.) Becker ex Heyne (syn. Bambusa asper Schult.). 2n=. Probably from the Malay Peninsula and adjacent areas. Unknown wild. Secondary centre: Malaysian Archipelago. Ithas strong stems and edible shoots. DENDROCALAMUS BRANDISH Kurz. 2n=72. Burma, Thailand, Cambodia and Vietnam. Its stems are used as building material. DENDROCALAMUS MERRILLIANUS Elm. 2n=. Primary centre: Philippines. Its stems are used as building material. DIN0CHL0A GIGANTEA Munro. 2n=. Lower Burma. Primary centre: Lower Burma. Its stems are used asbuilding material.the plant of this species is the largest among the bamboos. DIN0CHL0A MACLELLANDII Kurz. 2n=. Cambodia, Laos and Vietnam. Its stems are used in the basket industry. DINOCHLOA PENDULUS Ridb. 2n=. Malay Peninsula. Its stems areused for baskets. GIGANTOCHLOA APUS (Schult.) Kurz. 2n=. Burma and Indochina. Several species are cultivated in Java,Borneo and Philippines and on the Malay Peninsula (Tenasserim). Secondary centre: Java. GIGANTOCHLOA LIGULATA Gamble.2n=. N. part of Malay Peninsula and Thailand. The timber is used and the shoots are eaten. GIGANTOCHLOA MAXIMA Kurz. 2n=. Unknown wild. Secondary centre: Java. Its stems are an excellent material for building. GIGANTOCHLOA SCORTECHINII Gamble.2n= Malay Peninsula. Its stems are used as building material. GIGANTOCHLOA SCRIBNERIANA Merr. 2n=. Laos and Cambodia. Its stems are used as building material. GIGANTOCHLOA VERTICILLATA (Willd.) Munro. 2n=. SE. Asia. Cultivated in W. Africa, W. Indies,Fiji and elsewhere (Purseglove, 972). ISCHAEMUM INDICUM (Houtt.) Merr. Batiki blue grass. 2n=. SE. Asia. Cultivated in W. Africa, W. Indies, Fiji and elsewhere (Purseglove, 972). MISCANTHUS FLORIDULUS (Lab.) Warb. 2n=38. New Guinea. This species together with M. sinensis* from Fiji played a role in the origin of theedule group of Saccharum officinarum* (Grassl, 977). ORYZA GRANULATA Nees & Am. (incl. 0. meyeriana Baill.). 2n=24, 48. Malaya. It belongs to the 'officinalis' group of Oryza. ORYZA LONGIGLUMIS Jansen. 2n=48. New Guinea. ORYZA MINUTA Presl. 2n=48, genome formula BBCC. This wild species has the same genomes as the African 0. eichingeri*. Itbelongs to the 'officinalis' group. ORYZA NIVARA Sharma & Shastry. 2n=24. S. and SE. Asia and N. Australia. This isthe wild annual close relative of 0. sativa*. ORYZA OFFICINALIS Wall. 2n=24, genome formula CC. This wild species is the parent of the species belonging to the 'officinalis' group. ORYZA PERENNIS Moench. 2n=24, genome formula AA. The distribution of this wild species is discussed on p. 74.In Oceania, the Oceanian race (2n=24) of this species developed. See also 0. rufipogon* and index. ORYZA RIDLEYI Hook.f. 2n=48. SE. Asia. ORYZA RUFIPOGON Griff, (syn. 0. montana Lour.). 2n=24. Several SE. Asian countries. It is a pernicious weed of rice land. It easily crosses with rice. Itmight be a hybrid product of natural crosses of rice and 0. perennis* and would then be of the same nature as 0. sativa var. fatua*. According to recent views of taxonomists, 0. rufipogon includes 0. perennis*, 0. fatua*, 0. sativa f. spontanea*, 0. perennis ssp. balunga*, 0. perennis ssp. cubensis*. See also 0. nivara*. ORYZA SATIVA L. Rice. 2n=24, genome formula AA. Primary centre: SE. Himalaya area (p. 74). Ecotype Tjereh and Bulu developed inindonesia. Tjereh belongs to the ecospecies 'aman' (ssp. indica Kato) (Morinaga, 968). ORYZA SCHLECHTERI Pilger. 2n= PASPALUM SCROBICULATUM*. New Guinea. SACCHARUM OFFICINARUM L. Sugar-cane, Noblecane. 2n=40II=80. New Guinea. Modern clones resulting (2n=00-25) from hybridization. Sugar-cane derives form S. robustum Brandes & Jeswiet ex Grassl, which growswild in New Guinea, Celebes, Borneo up to New Hebrides. Basic types of N. coast of New Guinea, Celebes and Borneo have 2n=60, while those from southern coast of New Guinea have 2n=80. The first basic type may have originated in Borneo The second basic type may have been the wild parent from which primitive sugar-cane developed.

50 GRAMINEAE - GRAMINEAE 55 Dispersal of races Oryza sativa across Asia. Area of origin (grey), "Indica" ( ), "Javanica" ( ), extend of wild relatives ( ) (Chang, 976). Japonica" S. robustum is cultivated for its large stalks used for fences and for construction. During the domestication of sugar-cane, geographic types may have hybridized. Through selection, sugar-cane has a much lower fibre content, increased juiciness and sugar content throughout the stalk. These are the New Guinea Noble canes which must have originated ca 700 BC. On New Guinea, these Noble canes hybridized with Miscanthus sinensis. With hybridization, and backcrossing and perhaps other intergeneric hybridization (Roach, 972), the Edule group of canes developed (Grassl, 964, 967, 968, 977). This group has also been described as S. edule Hassk. The Edule canes have been subdivided into Vitho canes (syn. Erianthus maximus Brongn. and E. pedicelare Trin.), 2n=87-00 and Nduruka canes (syn. S. edule Hassk.), 2n=70. From Fiji, sugar-cane was taken to Hawaii, which lies outside the Miscanthus area. There the Hawaiian (Original) Noble canes developed, which are thicker and are used as chewing canes and as ornamentals. The New Guinea Noble canes were also transported northwards and northwestwards to other parts of Indonesia, Philippines, S. Japan,S. China (p. 38) and India (p. 75). SACCHARUM SPONTANEUM L. Wild sugar-cane. 2n= (x=0) SE. Asia. In Philippines three groups are found: 2n=56, 2n=72 and 2n=80. Plants with 2n=80 occur in many habitats for instance on mountains and river banks,and in grassland (Rithidech & Ramirez, 974). In the Indo-Gangetic Plain,plants with 2n=40-72 are found (Mehra & Sood, 974). In Indonesia and especially in Java and Sumatra, plants with 2n=2 occur. They may be of hybrid origin, deriving from S. officinarum (2n=80) x S. spontaneum (2n=64). Used as fencing for pigs by the Austronesians from SE. Asia. SCHIZOSTACHYUS BRACHYCLADUS Kurz. 2n= Java and E. Malaysia. Secondary centre: Malay Peninsula. SCHIZOSTACHYUS GRANDE Ridl.2n= Malay the

51 INDOCHINESE-INDONESIAN REGION Peninsula, SCHIZOSTACHYUSLULAMPAO Merr. 2n=. Centre of origin Philippines. Used in the paper industry, SCHIZOSTACHYUS ZOLLINGERI Steud. lay Peninsula, Java and Sumatra. 2n= Ma- SINOCALAMUS LATIFLORUS (Munro) McClure (syn. Dendrocalamus latiflorus Munro.) 2n=. Burma, Thailand, Taiwan and Philippines, Its stems are used as building material. The young shoots are eaten. They are also canned and exported. TRITICUM TURGIDUM (L.) Thell. Durum wheat (syn. T. durum Desf.). 2n=28, genome formula AABB. For origin see p. 94.In India a secondary centre of diversity (Jain et al., 976). VETIVERIA ZIZANIOIDES Stapf (syn c V. odorata Virey, Andropogon muricatus Retz,), Vétiver. 2n=20. A grass of Trop. Asia. Cultivated for volatile oils in its rhizomes and as a hedge plant. ZEA MAYS L. Maize. 2n=20. Domesticated in C. America (p. 90). Secondary centre arose in S. and SE. Asia (Brandolini, 970). Guttiferae CALOPHYLLUM INOPHYLLUM L. Alexandrian laulal, Undi. 2n=32. Coastal regions from E. Africa upto Australia and Polynesia, Often planted. In India it has a rather restricted economic importance. The kernel yields Domba oil. GARCINIA ATROVIRIDES Griffith. Gelugur.2n=. Assam and Malaya, Occasionally cultivated. GARCINIA COCHINCHINENSIS (Lour.) Choisy. 2n=. Cochin China. Cultivated for its fruits. GARCINIA DULCIS (Roxb.) Kurz. Baniti.2n= Philippines to Java, The bark yields a green dye and the fruits are edible. Occasionally cultivated in Java. GARCINIA INDICA Choisy.Kokura, Kokan, Ktambi. 2n=48, c. 54. Trop, Asia. Cultivated for its fruits. In India, it is a minor oil-seed plant (p. 75). GARCINIA MANGOSTANA L. Mangosteen. 2n=c. 76, 96. Malaysia, It is considered tobe the most delicious of all tropical fruits. It is derived from wild G. silvestris, which is also found in India (p. 75). GARCINIA MULTIFLORA Champ, (syn.g. tonkinensis Vesque), Cây giôc, Bira tai, 2n= N, Vietnam, Laos, Hainan and Hongkong. Cultivated in N. Vietnam for its fruits. GARCINIA PEDUNCULATA Roxb. Tikul. 2n= Bengal and Silhat (Bangladesh). Cultivated for its fruits. GARCINIA TINCTORIA (DC.) W.F. Wight. Matau, Gamboge tree. 2n=c.80. India (p. 75)and Malaya. Cultivated in the tropics for its fruits. Hydrophyllaceae HYDROLEA ZEYLANICA Vahl 2n=. Trop. Asia. Cultivated in Java for its young leaves. Labiatae COLEUS AMBOINICUS Lour. (syn. C. aromaticus Benth.). Indian borage, Dacon ajenton 2n=68. Indonesia. Cultivated in SE. Asia and W. Indies for its aromatic leaves. These leaves are used in stuffings and for flavouring meats. They may substitute for sage (Salvia officinalis*)and borage (Borago officinalis*) (Purseglove, 968). COLEUS PARVIFLORUS Benth. (syn. C. tuberosus Benth.). 2n=56, 64. This tuber crop is cultivated in SE, Asia. OCIMUM GRATISSIMUM L. 2n=40, 48, 64. Trop. Asia, esp. India. Cultivated in India as medicinal crop. OCIMUM SANCTUM L. Holy basil. 2n=64. Shrub of trop.old World.Cultivated as a sacred plant in India and elsewhere, ORTHOSIPHON STAMINEUS Benth. (syn. Ocimum grandiflorum Blume). 2n=, SE, Africa to Australia, A shrub cultivated in Java as medicinal plant, P0G0STEM0N CABLIN (Blanco) Benth. Patchouli, 2n=, Philippines. Cultivated for its essential oil. Lauraceae CINNAMOMUM BURMANI Blume. Batavia cinnamon. 2n=. Malaysia. Cultivated there. CINNAMOMUM CASSIA Blume (syn, C. aromaticum Nees), Cassia cinnamon, Chinese cinnamon. 2n=, Cultivated ins. China for its bark and flower buds. Cassia oil isobtained from the leaves (Purseglove, 968), LITSEA CALOPHYLLA (Miq.) Mansf. (syn. L. tetranthera Mirb., L, sebifera Blume). 2n= Malaya and Indonesia.Cultivated esp, in Bangka, Indonesia for its fruits. Leguminosae ALBIZIA LEBBECK Benth e Lebbek, Indian walnut. 2n-26. Trop. Asia to N. Australia, Cultivated in tropics and subtropics as fodder crop and as shade tree.

52 GRAMINEAE - LEGUMINOSAE 57 ALBIZIA MOLUCCANA Miq. (syn. A. falcata (Stickm.) Backer). 2n=. Malaya. Cultivated there and elsewhere as shade tree and as green manure. ALBIZIA MONTANA (Jungh.) Benth. 2n= Malaysia. Cultivated as green manure and shade tree. ALBIZIA SUMATRANA. 2n=. Indonesia. Cultivated in Zaïre as soil cover, green manure and shade tree. CANAVALIA GLADIATA (Jacq.) DC.Sward bean. 2n=22, 44. Old World. Probably derived from C. gladiolata Sauer (2n=22), which occurs in the Burma-Yunnan area (Sauer, 964). Wild in Trop. Asia and Africa. Cultivated in Asia, especially in India as food, forage and cover crop or as green manure. In some areas, it has naturalized (Purseglove, 968). In Japan, the white-seeded cultigen (var. alba) is cultivated (p. 40). C. polystacha (Forsk.) Schweinf (2n= ). Cultivated from SW. China upto Ethiopia/Somalia as pulse and seed. It is also considered the parental type of C. gladiata. CASSIA DIDYMOBOTRYA Fresen. Candelabra tree. 2n=28. A shrub used as green manure in Malaya and Sri Lanka. CASSIA HIRSUTA L. 2n=28, 56. Vigorous bush used in Malaya, Indochina and Uganda for soil cover. C. intermedia Sharma, Vivek. & Rathak (2n= )is a natural hybrid of C. occidentalis L., (2n=26, 28) and C. hirsuta (Sharma et al., 974). CASSIA LESCHENAULTIANA DC. 2n=48. Shrub used in India and Indonesia as green manure. CASSIA MIMOSOIDES L. 2n=6, (32). Trop. Asia and Africa. Tree used in Indochina and Indonesia as green manure. CASSIA OCCIDENTALIS L. Coffee senna, Negro coffee, Stink weed. 2n=26, 28. Tropics. Used in Indochina as green manure. CASSIA PUMILA Lam. Indochina. 2n= Cover crop in CASSIA SIAMEA Lam. (syn. C. florida Vahl.). 2n=28. India, Malaya and Indonesia. Cultivated in Malaya and India as fodder crop. Introduced on Cuba as green manure. CASSIA TORA L. Sickle senna, Wild senna.2n= 26, (28, 56). Tropics. Occasionally cultivated as a green manure in China and Indonesia. CLITORIA LAURIFOLIA Poir. (syn. C. cajanifolia Barth). 2n=24. Tropics. Occasionally cultivated in Sri Lanka and Indonesia and formerly in Tanzania as green manure. CROTALARIA ALATA Ham. 2n=6. Malaysian Archipelago. Excellent green manure. DERRIS DALBERGIOIDES Baker.2n= shade tree in SE. Asia.. Used as DERRIS ELLIPTICA Benth. Derris. 2n=22, 24, 36. From E, India to New Guinea except S. Malaya. Clones are distributed locally except one which is found inmany places in SE. Asia. This clone has a high content of rotenone (Toxopeus, 952). DERRIS MALACCENSIS Prain. Derris. 2n=22, 24. Malaysian Archipelago, where also cultivated types are found. Like D. elliptica*, it is a source of rotenone (Toxopeus, 952). DERRIS MICROPHYLLA (Miq.) Jackson. 2n= Used as shade tree in SE. Asia. Introduced into Indochina. DERRIS ROBUSTA Benth. 2n= tree in SE. Asia. Used as ashade DESMODIUM GYROIDES DC. 2n=20, 22. Trop. Asia. A shrub used as a green manure. INDIGOFERA TEYSMANNII Miq. 2n=32. SE. Asia. It is a green manure. INOCARPUS EDULIS Forst. Tahiti chestnut.2n= 20. From Malaysia to Polynesia, where it is cultivated for seeds and as shade tree. MELILOTUS SAUVEOLENS Ledeb. (syn. M. graveolens Bunge). Daghestan sweet clover. 2n=6. E. Asia and Indochina. Cultivated in USA. Some annuals are found in this biennial plant. MIMOSA SEPIARIA Benth. 2n=. Trop. Asia. Used forhedges (Mansfeld, 959). MUCUNA ATERRIMA (Piper h Tracy) Holland (syn. Stizolobium aterrima Piper & Tracy). Mauritius bean,bengal bean. 2n=22. Trop. Asia. Cultivated there and elsewhere as agreen manure and soil cover. MUCUNA CAPITATA (Roxb.) Wight & Arn. 2n= India and Java. Cultivated as a vegetable and for its seeds. MUCUNA COCHINCHINENSIS (Lour.) A. Cheval, (syn. M. nivea DC., Stizolobium niveum 0. Kuntze). 2n=22. Cochin China. Cultivated in tropics as vegetable, for seeds,as green manure and soil cover. MUCUNA DEERINGIANUM (Bort.) Small, (syn. Stizolobium deeringianum Bort.). Florida velvet bean. 2n=22. Probably trop. Asia or Malaysia. Cultivated as cover crop, green manure and forage crop. MUCUNA PRURIENS DC. var. utilis Wahl. Stizolobium aterrimum Piper & Tracy). (syn. Bengal

53 58 INDOCHINESE-INDONESIAN REGION INDOCHINESE-INDONESIAN REGION Distribution of wild bananas ( ) and cultivation in Africa ( ). Origin and movements of Eumusa groups (AA-ABBB) and ofaustralirausaseries (open arrow) (Simmonds, 962). AAB group. Its major centre of origin lies in India (p, 68), while a clone (Maio maoli)may have arisen in Philippines (Simmonds, 964). The third hybrid group is the triploid AAB group. S. India is a major centre of origin. It is quite likely that a second centre is found in Philippines. The fourth hybrid group, consisting of one clone, is the tetraploid ABBB group. Its centre of origin very probably lies in Indochina (Simmonds, 964), MUSA* cultivars of the ABB group. 2n=33. Most ABB cultivars originated in S. India (p. 68). However it is possible that after the cultivated M. acuminata (AA)reached Philippines, hybrids arose with M. balbisiana*. MUSA* BALBISIANA f^ 3 (+- ^ W j> The wild Musa balbisiana ( ) and M. acuminata ( ) types (Simmonds, 962). MUSA* TEXTILIS Nee. Abaca, Manilla hemp. 2n- 20. Philippines. A tall perennial. Cultivated there and elsewhere in the tropics for its fibre. Canton fibre is obtained from a natural completely sterile hybrid (2n=2)of M. textilis x M. balbisiana*. Myristicaceae MYRISTICA ARGENTEA Warb. Papuan nutmeg. 2n=. New Guinea, where it is also occasionally cultivated (Flach & Cruickshank, 969). MYRISTICA FRAGRANS Houtt. Banda nutmeg. 2n= 42, 44. Centre of origin: Moluccas. It is not found there wild. From there,it spread throughout the tropics (Flach & Cruickshank, 969). Myrtaceae EUGENIA AQUEA Burm.f. (syn.syzygiura aqueum (Burm.f.) Alston). Watery roseapple. 2n= Bangladesh, Burma, Sri Lanka, Sumatra and Moluccas. It is also cultivated. EUGENIA CARYOPHYLLUS (Sprengel) Bullock & Harrison (Syzygium aromaticum (L.). Merr. & Perry). Clove tree. 2n=. The wild clove tree, sometimes named E. obtusifolia Reinwardt (Caryophyllus sylvestris T. & B,), grows on many islands of Moluccas and in New Guinea. It has bigger leaves and flower buds, and is less aromatic than the cultivated tree. Cultivated for a long time. Some variation exists in and outside its centre of origin. EUGENIA FORMOSA Wall. (syn. Syzygium mappaceum (Korth.) Mansf.). 2n=. Trop. Asia.

54 MUSACEAE - PINACEAE 6 Tree cultivated in Cochin China for fruits. EUGENIA JAMBOLANA Lam. (syn.syzygium cumini (L.). Skeels, Eugenia obtusifolia Roxb., E. cumini (L.) Druce). Java plum, Jambolan plum. 2n=33, 44, 46, 55. India to Malaysia, China and N. Australia. It is cultivated there and elsewhere. In India, a large-fruited type is cultivated (Mansfeld, 959). EUGENIA JAMBOS L. (syn.syzygium jambos (L.) Alston). Roseapple. 2n=28, 33, c.42, 44, 46, c.54. Tree cultivated for a long time in Indo-Malaysia. Its centre of origin is not known. EUGENIA JAVANICA Lam. (syn.syzygium semarangense (Bl.) Merr. & Perry). 2n=33, 42, 44, 45, 66, 88, 0. Malaysia to India. Much cultivated in Java. EUGENIA MALACCENSIS L. (syn.syzygium malaccensis (L.) Merr. & Perry). Pomerac,Malay apple. 2n=22. Malaya. Some varieties are known. MELALEUCA QUINQUENERVIA L.f. (syn.m. leucadendra L.). Cajêput tree. 2n=. Australia to Burma. Planted in forestry projects in Philippines, Hawaii and elsewhere. Also planted to drying out swamps, and as ornamental. PIMENTA ACRIS Kostel. 2n=22. Indonesia. Cultivated there foroil distilled from the leaves (Purseglove, 968). RH0D0MYRTUS T0RMENT0SA Wight. Downy rose-myrtle. 2n=. India and Malaysia. A shrub cultivated in the (sub)tropics for its fruits. Nyctaginaceae PISONIA ALBA Span. Maluko, Lettuce tree. 2n=. Malaya.Wild tree is called P. sylvestris Teijsm. & Binn. (syn.p. grandis R. Br.) (2n= ). Cultivated for leaves, which are used as vegetable. Palmae ARECA CATECHU Merr. Betelnut palm. 2n=32. Trop. Asia. Cultivated for nuts. ARENGA PINNATA (Wurmb.) Merr. Sugar palm. 2n=26, 32. Primary centre: the Indonesian- Hindustani gene centre (p. 78). Possible secondary gene centre: India. BORASSUS FLABELLIFER* laya, Moluccas and New Guinea. Sago is obtained from the marrow of the stem. This species isoccasionally split in M. sagu - the wild type and M. rumphii (Willd.) Mart. (2n= ) - the cultigen. NYPA FRUTICANS Wurmb. (syn. Nipa fruticans Thunb.). Nipa palm. 2n=6. SE. Asia upto Australia. Cultivated on Sumatra for its leaves and for wine production. Introduced to the mangrove area of S. Nigeria, where it has runwild (Zeven, 973). PRITCHARDIA GAUDICHAUDII H. Wendl. 2n= Sandwich Islands. Cultivated there for its leaves which are used for thatching. PRITCHARDIA PACIFICA Seem & Wendl. 2n=36. Fiji and Samoa. Cultivated for its leaves, which are used for thatching. SALACCA EDULIS Reinw. 2n=. Malaysian Archipelago. Cultivated on Java for its edible fruits. Pandaceae PANDANUS AMARYLLIFOLIUS Roxb. (syn.p. odorus Ridl.). 2n=. Cultivated in Malaya for its fragrant leaves. PANDANUS BROSIMAS Merr. & Perry. 2n=. Cultivated in the highlands ofnew Guinea for its seeds, which have a pleasant flavour and are rich inoil (Purseglove, 972). PANDANUS SPURIUS Miq. (syn.p. moschatus seu laevis Rumph.,R. moschatus Rumph. ex. Miq.,P. tectorius Soland. var. moschatus (Rumph.ex Miq.) Merr.,P. laevis Lour.,P. odoratissimus L.f.,P. inermis Roxb.). Thatch screw pine, Putat, Pudak. 2n=c.5, 54, 60. Cultivated in SE. Asia to the extremes of Polynesia for its leaves for thatch and for its fruits.the cultivar is one clone, probably originated as a bud sport on a staminate plant of some wild species of section Pandanus. Perhaps the mutation occurred on a specimen of P. spurius some millenia ago (St. John, 965). PANDANUS WHITMEEANUS Martelli. Paogo. 2n= Cultivated innew Caledonia,New Hebrides and elsewhere. On Futuna, it isused only for personal adornment. The fruit oil is used to perfume coconut oil (St. John & Smith, 97). Pentaphragmaceae CALAMUS CAESIUS Blume.2n=. Malaya, Borneo and Sumatra. Cultivated for stems. COELOCOCCUS ARMICARUM Warb. Polynesian ivorynut palm. 2n=. Carolina Islands. Cultivated in Philippines for its ivory-like nuts. METROXYLON SAGU Rottb. Sago palm. 2n= Ma- PENTAPHRAGMA BEGONIAEFOLIUM Wall. 2n=. A fleshy herb cultivated as vegetable in Malaya (Terra, 967). Pinaceae PINUS MERKUSII Jungh & de Vriese (incl. P. merkusiana Corling & Gaussen). Merkus pine.

55 INDOCHINESE-INDONESIAN REGION 2n=. Burma tophilippines and south to Sumatra. Planted in tropics as a source of turpentine and paper, and to control erosion. Piperaceae PIPER BETLE L. Betel pepper,betle vine, Betal, Sirih. 2n=32, 64, (78). C. and E. Malaysia, Cultivated in the tropics. The leaves are chewed together with betelnut (Areca catechu*). PIPER CUBEBA L.f. Cubeb, Cubebe, Tailed pepper, 2n=24. Cultivated there and in neighbouring countries. PIPER METHYSTICUM Forst. Kava pepper.2n=. Polynesia. Cultivated there.the roots and rhizomes are used to prepare a non-alcoholic beverage. In small amounts it is a stimulant; in large amounts a narcotic. PIPER RETROFRACTUM Vahl (syn. P. officinarura DC,). Javanese long pepper, 2n=. Malaysia. It resembles P. longum*. Cultivated for its spike, which is used as a spice. PIPER SAIGONENSE C. DC. Lolo. 2n=. Indochina, Cultivated there occasionally. Closely related top, lohot C. DC., which comes from Tonkin district. Polygonaceae POLYGONUM ODORATUM Lour,2n=, Indochina. Cultivated as a pot-herb invietnam,, Rosaceae RUBUS ALBESCENS* RUBUS ROSAEFOLIUS Smith. Cape bramble, Mauritius raspberry, 2n=, Tropical Asia, Introduced in other continents. Cultivated, It is considered a parent of R, probus Bailey, Queensland raspberry, a shrub from Australia. The other parent isr. ellipticus Smith, the Yellow Himalayan raspberry from E. India, Rubiaceae MITRAGYNA SPECIOSA Korth. 2n=. Malaya and S. Thailand, Cultivated as a substitute for opium. MORINDA TRIFOLIA L. Indian mulberry. 2n= Indonesia and Malaya.Cultivated on Java as adye crop. OLDENLANDIA UMBELLATA L. Indian madder. 2n= 36, Trop. Asia. Cultivated as a dye plant. UNCARIA GAMBIR (Hunt.) Roxb. Gambier. 2n=. Malaya, Formerly cultivated in SE. Asia. Its leaves and young branches contain a tannin. Rutaçeae AEGLE MARMELOS (L.) Corr. Indian bael, Bengal fruit. 2n=8, (36). Cultivated in SE. Asia and some other trop, countries for its fruits, which are used medicinally. CITRUS AURANTIFOLIA (Christm.) Swing. Lime. 2n=8, (27). Probably Malaysian Archipelago or N. India. Itmay derive from a cross of C. medica* with a biotype of the primitive subgenus Papeda (Scora, 975). Wild trees are reported to grow in N. India.Spread througout tropics.the cultivar Tahiti is triploid. Interspecific hybrids have been obtained. Mandarin lime isprobably a hybrid with C, reticulata*, sweet lime with C. medica* and limequat with Fortunella margarita*. The nakoor lime (named C. nakoor) is a complex natural hybrid with some Papeda group parentage. The Rangpur lime belongs toc. reticulata*. CITRUS AURANTIUM L. Sour orange, Seville orange, Bigarade. 2n=8. Probably SE, Asia or Cochin China, Unknown wild. It may derive from C. reticulata* x C. grandis* (Scora, 975). Spread throughout (sub)tropics. In some areas,it has run wild, Ssp, bergamia (Risso & Poit.) Wight & Arn., Bergamot (2n= 8)is cultivated especially in Calabria, S. Italy for the production of bergamot oil (p, 05), Crosses with C, sinensis* (Sweet orange)gave Bitter sweet orange. The var. myrtifolia Kergawl., Myrtle-leaved orange is a bud mutant. Its fruits, Chinottos, are candied in Italy and S. France. CITRUS GRANDIS (L.) Osbeck (syn. C. decumanus L, 2n=8, 2; C. maxima (Burm.) Merr. 2n=8, 36). Pummelo, Shaddock. 2n=8, 36. Probably SE. Asia, Primary centre of diversity: SE. Asia,Spread to China, India and Iran, and later toother tropical countries (by Captain Shaddock to Barbados in 7th Century), Unknown wild. The best fruits come from Thailand where the plants are cultivated on ridges surrounded by brackish water. Self-incompatible and monoembryonic perennial. Introgression with C. reticulata* occurs (Scora, 975). CITRUS HYSTRIX DC. Mauritius papeda,2n= Philippines and Burma to Malaya. A small type cultivated for its fruits. CITRUS JAMBHIRI Lush. Rough lemon.2n= It derives from C, medica* x C. reticulata* (Scora, 978). CITRUS LIMETTA Risso. Sweet lemon. 2n=8. Trop. Asia. Small tree cultivated in some countries. CITRUS LIMON (L.) Burm.f. Lemon. 2n=8, 36. Centre of origin somewhere in SE. Asia.The area east of Himalayas in N. Burma and S. China has been suggested. Unknown wild. Has a complex origin (Torres et al,, 978). A

56 PINACEAE -SOLANACEAE 63 secondary centre:the Mediterranean Region (p. 8). Scoi-a &Malik (970) suggested that this species might be a stabilized hybrid of C. medica* - C 0 aurantifolia* assemblage. Cultivated in several (sub)tropical regions. CITRUS MEDICA L. Citron. 2n=8. Subtrop. Asia. A basic Citrus species (Torres et al., 978). Itmay be one of the parents of C. limon* and C. jambhiri*. Monoembryonic. CITRUS MITIS Blanco. Calamondin. 2n=8. Philippines. It derives from C. reticulata var. austera x Fortunella sp. A tree occasionally cultivated in (sub)tropics. Hybrids of this species have been produced, for instance Calarin and Calashu are hybrids with C. reticulata* (Satsuma), Calamondin is a hybrid of C. reticulata x Fortunella sp. CITRUS PARADISI Macf. Grapefruit. 2n=8. SE. Asia. Unknown wild. Probably derived from C. grandis* x C. sinensis* (Torres et al., 978). Chiranja is a hybrid with C. sinensis and citrumelo with Poncirus trifoliata*. CITRUS RETICULATA Blanco (C. nobilis Andr. non Lour,). Mandarin, Tangerine. 2n=8. Probably Philippines, or Cochin China. Unknown wild. Secondary centre arose in Japan (p. 45). Minessy et al. (970) found close relationship with C. sinensis* 'Balady Blood'. Its relationship with C. paradisi* 'Duncan' and 'March' is moderate and with C. grandis* remote. Var. austera Swing is the sour mandarin. Itprobably includes the Rangpur lime (Purseglove, 968). Hybrids with other species have been made. For instance Oranguma is an artificial hybrid of Satsuma x C. sinensis* (Orange), and Tangor is a natural hybrid of the same parents.its origin is in Thailand. Tangor has been described as C. nobilis Lour. Citrandarin derives from Poncirus trifoliata* x C. reticulata, calamondin (see C. mitis) from C. reticulata x Fortunella sp., and chironja from orange x C. reticulata. CITRUS SINENSIS (L.) Osbeck (syn. C. aurantium L. var. sinensis L.). Sweet orange. 2n=8, (27, 36). Probably S. China or Cochin China. Unknown in wild. It may derive from C. reticulata* x C. grandis* (Scora, 975). Secondary centres: Israel and Spain (p. 8). It was already mentioned inchinese sources dated 2200 BC. It has the same origin as C. aurantium* but the wide genetic variation of C. reticulata causes differences in the derived species. It is widely distributed in the (sub)tropics. There are many cultivars, Citrange is a hybrid with Poncirus trifoliata* and chironja is a spontaneous hybrid with C. paradisi*. It originated inpuerto Rico. By apomixis, it breeds true. MURRAYA EXOTICA L. Limonia. 2n=8. Trop. Asia. Used for hedges. MURRAYA PANICULATA (L.) Jacq. Cosmetic barktree, Orange jasmine. 2n=8. SE. Asia. Cultivated in the tropics as an ornamental and for hedges. Thewood (Satinwood) is used in Java to make cutlery. Santalaceae SANTALUM ALBUM L. (syn. Sirium myrtifolium L.). Sand^el wood. 2n=0. E. India to Malaysia. Cultivated there and elsewhere for scented wood. Sapindaceae ERIOGLOSSUM RUBIGINOSUM (Roxb.) Blume (syn. E. edule Bl.). 2n=. Trop. Asia to New Guinea and Australia, A small tree cultivated in Indonesia and elsewhere. NEPHELIUM LAPACCEUM L. Rambutan. 2n=22. Malaysian Archipelago. Cultivated for its delicious fruits. Many varieties have been developed. NEPHELIUM MUTABILE Blume. Pulasan. 2n= Malaysia. Cultivated in SE. Asia and in other countries. POMETIA PINNATA Forst. Matoa, Taun, 2n= Malaysia, Indonesia, Papua New Guinea and Pacific islands. A forest tree used for timber and for its fruits. Cultivated for its edible fruits. On W. Irian alongside the banks of the Sentani Lake. This crop will probably be replaced by higher -yielding exotic fruit trees (Rappard, 96). SAPINDUS RARAK DC.2n=. Cochin China and Malaysia. Planted in Java, India and elsewhere for its fruits. Sapotaceae MANILKARA ELENGI (L.) Chev. 2n=. Origin uncertain (Uphof, 968). Cultivated in the Malaysian Archipelago. PALAQUIUM GUTTA (Hook.) Burck. Gutta percha. 2n=24. Malaysia, It is tapped for its latex. In general, the tree is first felled. PAYENA LEERII (Teijsm. & Binn.) Kurz. 2n= Burma and W. Malaysia. Cultivated on Java as a source of gutta percha. Saururaceae HOUTTUYNIA CORDATA Thunb. 2n=56, 96, c.96, Indochina and China. Cultivated in Vietnam for salad and as medicinal crop, Solanaceae LYCINUM CHINESE Mill. Chinese wolfberry.2n== 24. E. Asia. Cultivated in Java as vegetable.

57 64 INDOCHINESE-INDONESIAN REGION SOLANUM UPORO Dunal. 2n=. Polynesia. Cultivated infiji for its fruits. Stilagninaceae ANTIDESMA BUNIUS (L.) Spreng. Bignay, China laurel. 2n=7,India to Australia. Cultivated in Malaysia and elsewhere for its fruits (Purseglove, 968). Styraceae STYRAX BENZOIN Dryander. 2n=. Malaysian Archipelago. Planted in Sumatra. Taccaceae TACCA PINNATIFIDA Forst, (syn. T. involucrata Schum. & Thonn., T, leontopetaloides (L,) Kuntze). Tacca pin, Tahiti arrowroot. 2n=30, SE. Asia (Massai & Barrau, 956). Possibly domesticated by Polynesians for its starchy roots and introduced into Malaysia and Madagascar. Also found from Ethiopia to W. Africa. Umbelliferae CARUM ROXBURGHIANUM Benth. (syn. Trachyspermum roxburghianum (DC) Wolff). Ajmud. 2n(wild)= (20, 40) 42, (44); (cultivated)=20. Trop. Asia, Cultivated in Indochina, Sri Lanka and India. LIGUSTICUM MONNIERI Calest. (syn. Selinum monnieri L.). 2n=. E, Europe, Siberia, China and Vietnam. Occasionally cultivated in N. Vietnam. OENANTHE JAVANICA DC. (syn. 0. stolonifera Wall.). Oriental celery, Water dropwort, Batjarongi. 2n=20. From Indochina to Malaya, Philippines, China, Korea, Japan and Australia. Cultivated in Indochina, Japan,China (Kihara, 969)and Java. A leafy vegetable that often occurs as a weed. laysia. Cultivated there. AMOMUM KEPULAGA Sprague & Burk.Round cardamon. 2n=. Cultivated in Malaysia and Java. AMOMUM KRERVANH Pierre. Krervanh. 2n= Cambodia. Cultivated in Indochina. AMOMUM MAXIMUM Roxb. Java cardamon.2n= Malaysia. Cultivated in Java. BOESENBERGIA PANDURATA (syn. Gastrochilus pandurata Ridl.). 2n=. Malaya and Java. Cultivated over wide area for its rhizome. CURCUMA HEYNEANA Valeton. 2n=. Java.The rhizomes are a source of an arrowroot.. Malaya. Cul CURCUMA PIERREANA Gagn. 2n^ tivated in Annam (Vietnam). CURCUMA XANTHORRHIZA Roxb. 2n=. Amboina. Occasionally cultivated in Java and Malaya. KAEMPFERIA GALANGA L. 2n=22, 54. Trop. Asia. Widely cultivated for its rhizomes. KAEMPFERIA ROTUNDA L. 2n=33, 54. Trop. Asia. Cultivated for its rhizomes. PHAEOMERIA MAGNIFICA Schuin, (syn. Alpinia magnifica Rose., A, speciosa D. Dietr,, Amomum magnificum Nenth.). 2n=. Malaya. Cultivated there. ZINGIBER CASSUMUNAR Roxb. Cassumunar ginger. 2n=22. Cultivated in Cochin China and Malaya. In Malaya as a village medicinal crop. ZINGIBER ZERUMBET (L.) Smith.Zerumbet ginger. 2n=22. Trop. Asia. Cultivated in Cochin China, Cambodia and elsewhere. Urticaceae LAPORTEA DECUMANA Wedd. 2n=. Moluccas. Cultivated as a medicinal plant. Zingiberaceae ALPINIA CONCHIGERA Griff, (syn. Languas conchigera Burk.). 2n=. Malaya, It is acommon village plant there, ALPINIA GALANGA (L.) Willd. Langwas, Greater galangal. 2n=48. Trop. Asia. Cultivated for its rhizomes. It is a common village plant. Several varieties have been observed. ALPINIA MALACCENSIS (Burra.f.) Rose. 2n=48, Malaysian Archipelago and E. India. This perennial herb is cultivated. AMOMUM CARDAMOMUM Willd. Cardamon. 2n= Ma-

58 3 Australian Region TheAustralian Regionwasnot described byvavilov,but it wasmarked outby Zhukovskij (970) because of the domestication of several plant speciesto important cropsor theuseofwild species as breeding parents. Themain crops derived from thisregion areeucalyptus species. Wild species useful fortobacco breeding arenicotiana debneyi and N.goodspeedii. Itis a secondary centre ofdiversity fortrifolium subterraneum. Agavaceae PHORMIUM TENAX J.R. etg. Forst.New Zealand flax, New Zealand hemp, Harakaka lily, Formio. 2n=32. New Zealand. Cultivated there. Introduced into S, America and other countries. The only other species of this genus Ph 0 colensoi Hook., mountain flax (2n=32) produces a weak fibre. It might be used as a breeding parent. Casuarinaceae CASUARINA EQUISETIFOLIA Forst. Swamp oak, Bull oak, Polynesian iron wood, Horsetail tree. 2n=8. It is often cultivated as soil stabilizer. Chenopodiaceae ATRIPLEX SEMIBACCATA R.Br. Australian saltbush, Berry saltbush. 2n=8. Australia. Cultivated as fodder crop on the saline soils of California and Arizona, USA. Gramineae HOLCUS LANATUS L. Yorkshire fog,soft meadow grass,woolly softgrass, Velvet grass, 2n=4. See p. 53. A secondary centre of diversity has developed in New Zealand (Jacques, 974). ORYZA AÜSTRALIENSIS Domin. 2n=24, genome formula EE. Australia. All research into the affinity of the species toother Oryza species uses plants derived from one collection (Chang, 970). Leguminosae ACACIA CYANOPHYLLA Lindl. 2n=26. Australia. Cultivated as an ornamental and in Europe to stabilize coastal dunes. ACACIA DEALBATA Link. Silver wattle. 2n=26. SE, Australia and Tasmania. Cultivated as an ornamental, for its timber and as soil stabilizer. It is the familiar florist's mimosa. ACACIA LONGIFOLIA (Andrews) Willd. (syn. A. cibaria F.V. Muell.). 2n=26. New South Wales, Australia. Cultivated as ornamental and as stabilizer of coastal dunes in Europe, ACACIA MEARNSII De Wild. Black wattle. 2n=26. Cultivated in several countries mainly for its tannin and as ornamental. Sometimes the names A. decurrens (Wendl.) Willd. or A, mollissima Willd, are wrongly used for black wattle. ACACIA PYCNANTHA Benth. Golden wattle.2n= South Australia and Victoria, Australia. Cultivated for tannin and as ornamental. GLYCINE CANESCENS F.J. Herrn. 2n=40. A close

59 66 AUSTRALIAN REGION relative of G. max*, being a possible source of resistance to powdery mildew, Microsphaera diffusa Cka. & Pk. LUPINUS COSENTINI Guss. (syn. L. varius L. ssp. varius Franco &P. Silva). Western Australia blue lupin, Sandplain lupin. 2n=32. Along the coast of Morocco and scattered in W. Mediterranean region. Introduced into W. Australia about 850 as a source of flour. Cultivated for summer sheep feed and soil improvement.it isnow widely naturalized (Gladstones, 970). PHASEOLUS LATHYROIDES L. Phasemy bean. 2n=22. Queensland, Australia. Used in E. Africa in pastures (Whyte et al,, 953). This species and Vigna radiata*possess a high homology of chromosomes, which points to a close relation (Biswas & Dana, 975). TRIFOLIUM SUBTERRANEAUM L. Subterranean clover, Sub clover, 2n=6. Primary centre in the Mediterranean Region (p. 5). Secondary centre : Australia. Malvaceae GOSSYPIUM AUSTRALE F.V. Muell. 2n=26, genome formula C3C3. N. Australia. GOSSYPIUM BICKII Prokh.2n=, genome formula GG, Queensland, Georgina River, GOSSYPIUM ROBINSONII F.V. Muell. 2n=26, genome formula C2C2. W. Australia. GOSSYPIUM STURTIANUM J.H. Willis. 2n=2x=26, genome formula CC. C. Australia, GOSSYPIUM STURTII F.V. Muell. 2n=26, genome formula C]Ci. C. and S. Australia. Musaceae MUSA (Australimusa). Fe r i banana. 2n=20. The fe'i banana originated from one or more wild Australimusa species in New Guinea-Solomon Islands area. Probably carried by man in an easternly direction. Cultivated especially in Tahiti, where many bunches are harvested from semi-wild plants. Some clones have been described as M. fehi Bert, ex Vieill., M. aiori Sagot, M. seemanii F.V. Muell. and M. troglodytarum L. (Simmonds, 964). Myrtaceae EUCALYPTUS ALBA Reinw. ex Blume. 2n-22. Timor and Flores and to the south of New Guinea,Cultivated in Brazil, Thewood is reddish-brown. The cork contains much tanning material. EUCALYPTUS AMYGDALINA Labill. (syn. E. salicifolia Cav,), Willowleaf eucalyptus, Peppermint tree. 2n=22. Tasmania. Cultivated in Gossypium sturtii Chile, Zaïre and W. Georgia (USSR). Closely related to E, regnans*. EUCALYPTUS ASTRINGENS Maiden. Brown mallet. 2n=22. SW. Australia. Cultivated in Morocco, S. Africa and Cyprus. The bark used for the tanning industry. It is very drought-resistant, EUCALYPTUS BOTRYOIDES Smith. Blue gum, Bangalay eucalyptus, Bastard mahogany. 2n=22. Coastal areas of SW. Australia, Cultivated in Algeria and Zaïre. E. trabutii Vilm. (2n=22), is a hybrid of E. botryoides Q and E. camaldulensis*0*arisen in Italy. EUCALYPTUS BROCKWAYI Gardn. 2n=22. S. Australia, Its area of distribution is limited. Cultivated in N. Africa. Extremely droughtresistant. EUCALYPTUS CAMALDULENSIS Dehn, calyptus, Australian kino, Red Australia, excluding Tasmania, ted almost in all countriesth; lyptus. Secondary centres: the region (p. 7), Brazil (p. 2 tina (p. 27). In cultivation neous hybrids have arisen. E. (2n=22)is a hybrid of E. bot camaldalensis 0*. A new form de Israel (p. 7). Longbeak eugum. 2n=22. It is cultivait grow Euca- Medi terranean 7)and Argenmany spontatrabutii Vilm. ryoides Q and E. veloped in EUCALYPTUS CINERA F.V. Muell. 2n=22. S. areas of New South Wales, Australia,Used as ornamental. It is a valuable source for breeding cold-resistant forms of Eucalyptus. EUCALYPTUS CITRIODORA Hook. Spotted gum, Lemon-scented gum. 2n=(20), 22, (28). N. coast

60 LEGUMINOSAE - MYRTACEAE 6 7. ^u s^ ^ «jj \ i. ^ ùk *5 f a j\ ^ *> x vi ^-" ^ \ s \ X *. " " î> b eu 0» V _^ " / y / / " " " - "*"" Ü». Wild Australirausa ( ) and Fe'i bananas ( )(Simmonds, 962). of Queensland, Australia, Cultivated inmany (sub)tropical countries for essential oil rich in citronellal. EUCALYPTUS CLADOCALYX F.V. Muell. (syn. E. corynocalyx F.V. Muell.). Sugar gum. 2n=22. S. Australia. Cultivated in Australia, Mediterranean area and insome African countries. The wood isof excellent technical value. EUCALYPTUS COCCIFERA Hook.f. 2n=. Tasmania. Because of its hardiness, it is used for breeding types for W. Georgia (USSR). EUCALYPTUS CREBRA F.V. Muell.2n=. Queensland, reaching New South Wales, Australia. Cultivated in several countries of Africa, India and Argentina. Some spontaneous hybrids are known. EUCALYPTUS CYPELLOCARPA L. Johnst. (E. goniocalyx pi. anct.). 2n=22. SW, Australia attaining m altitude. Cultivated in Mediterranean area,s. America and on Hawaii. EUCALYPTUS DALRYMPLEANA Maiden. Mountain gum. 2n=22. SE. Australia, attaining 350 m altitude and in C. Tasmania attaining 900 m altitude. Cultivated on the coasts of the Black Sea in the Caucasus. It is a promising economic species on Hawaii and in China. It is considered to be of hybrid origin.natural and artificial hybrids are known. It can be used in breeding better types. EUCALYPTUS DELEGATENSIS R.T. Baker, (syn. E. gigantea Hook.f.). Alpine ash, Woollybutt, Red mountain ash, White top stringbark. 2n=. The mountains of SE. Australia up to 350 m and in Tasmania up to 900 m. Cultivated innew Zealand, Hawaii and W. Georgia (USSR). Used for cultivation and as breeding parent in USSR. EUCALYPTUS DIVERSIC0L0R F.V. Muell, Karri. 2n=22. Coasts of SE, Australia. Cultivated in countries of Mediterranean area, in Africa and New Zealand. It is oneof the most valuable economic species. EUCALYPTUS EUGENI0IDES Sieb. (syn. E. scabra Dum-Cours.). White stringybark, Pink blackbutt. 2n=, Coasts areas of SE. Australia. Cultivated in S. Africa, Kenya, India and Hawaii. The wood is used in industry. Some natural hybrids are known. EUCALYPTUS GLAUCESCENS Maiden & Blakely. 2n=. Mountains of SE. Australia. Its distribution is very limited. Used for crossing with species of poor hardiness. EUCALYPTUS GLOBULUS Labill.Fever tree, Blue gum. 2n=20, 22, 28. SE. Tasmania. Cultivated. Secondary centre: the Mediterranean region. Used for wood and oil. Spontaneous hybrids are known under cultivation in Tasmania. EUCALYPTUS G0MPH0CEPHALA A.DC. 2n=22. SW. coasts of West Australia, Cultivated in countries of the Mediterranean region. Africa esp. Cameroon, Hawaii and New Zealand. It has the heaviest and strongest wood among all Eucalyptus species. In Algeria, some spontaneous hybrids are known. EUCALYPTUS GRANDIS Hill, ex Maiden. 2n= Coast areas of the N. part of New South Wales and SE. Queensland up to 650 m. Cultivated in Cameroon, Nigeria and Madagascar. Itis thought that E. 'saligna* or E, 'saligna/ grandis' are African strains developed after introduction of Queensland material (Larsen & Cromer, 970). EUCALYPTUS GUNNII Hook.f. 2n=22. Cultivated in USSR, Great Britain, Japan and Hawaii. Used for industry and breeding on the Caucasus coasts of the Black Sea. EUCALYPTUS LEUC0XYL0N F.V, Muell. (syn. E. conoidea Benth.). White ironbark, White gum.

61 AUSTRALIAN REGION 2n=22. C. areas of Victoria and South Australia, In the latter area, it is rare. Cultivated in the Mediterranean area esp, Cyprus and S. America esp. Argentina. Used for its wood and oil. Some geographic races and spontaneous hybrids have been described. EUCALYPTUS MACARTHURI Dean & Maiden. 2n=22. C,New South Wales, Australia, Cultivated in Africa esp. Zaïre, Hawaii, New Zealand, S. France and W. Georgia, USSR. Itproduces an essential oil.some spontaneous hybrids are known. In the USSR many (poly)hybrids have been produced. EUCALYPTUS MACULATA Hook,f. (syn. E. variegataf.v. Muell.). Spotted gum. 2n=22. Coastal areas of SE, Queensland, New South Wales and E. Victoria. Cultivated in Africa esp. Cameroon, Zaïre, Kenya and Madagascar; the Mediterranean region esp. Spain, France; Chile and Uruguay. The wood isvery valuable. EUCALYPTUS MAIDENII F.V. Muell. Maiden's gum, Spotted blue gum. 2n=22. Mountains of SE. Australia. Cultivated in Africa esp. Cameroon, Congo and Kenya; the Mediterranean area esp. Italy and Spain; Brazil and New Zealand. Its wood is valuable, containing essential oil. Some spontaneous and artificial hybrids have been reported. EUCALYPTUS MELLIODORA A. Cunn. Yellow box. 2n=22. Australia. Cultivated in the Mediterranean area, in Africa esp. Zaïre and Eritrea. Used for its wood and as an ornamental tree. It is extreme^ melliferous. There are geographic races and spontaneous hybrids known. EUCALYPTUS MICROCORYS F.V. Muell. Fallow wood. 2n=. Coastal areas of the N. part of New South Wales and SE. Queensland. Cultivated in Mediterranean area and Africa esp. Zaïre and Eritrea. Used for wood. Some spontaneous hybrids are known. EUCALYPTUS NIPHOPHILA Maiden & Blakely. 2n=. Alpine zone of SE, Australia, up to 2000 m. It tolerates -24 C and hence isof great importance for hybridization with valuable economic species. EUCALYPTUS PANICULATA Sm. (syn. E. fergusoni R.T, Baker). Grey ironbark. 2n=22. The coasts of New South Wales. Cultivated in Africa esp. Kenya and Tripoli; Mediterranean area, esp. Spain and Tripolitania; S. America, esp. Paraguay and Uruguay, and India, The wood is especially strong, heavy and durable. EUCALYPTUS PAUCIFLORA Sieb. ex.spreng. 2n= Sub-alpine zone of E, Victoria and the mountains of New South Wales and Tasmania, up to 650m. Cultivated in England, France, Japan and W. Georgia (USSR), On the fringe of its area, it is very hardy. It is valuable in breeding hardy strains. Some geographic races and spontaneous hybrids are known. EUCALYPTUS PERRENIANA F.V, Muell.ex Rodway. 2n-, Tasmania. Cultivated in the USSR. It is hardy (it tolerates -3 C). EUCALYPTUS REGNANS F.V, Muell. Mountain ash, Swamp gum, Australian oak, 2n=, Mountains of S. Victoria up to 900 m and in Tasmania up to 600 m. Cultivated in Kenya,New Zealand and other countries. This is the biggest and most valuable species in this genus, Some trees are recorded up to 96 m high. It is closely related to E, amygdalina*. EUCALYPTUS RESINIFERA Smith (syn. E. spectabilis F.V. Muell., E. hemilampra F.V. Muell.). Kino eucalyptus, Red mahogany, Forests mahogany. 2n=22. Coastal zone of S. Queensland and C. part of New South Wales. Cultivated in Argentina, Sri Lanka, Ethiopia, Cameroon and other countries. Thewood isvery valuable. EUCALYPTUS ROBUSTA Smith. Beakpod eucalyptus, White mahogany, Swamp mahogany. 2n=, Coasts of S. Queensland as far as S,of New South Wales, Australia. Cultivated in Mediterranean area, Africa esp. Cameroon, Zaïre and Kenya; Argentina, India and other countries. Often cultivated on swampy grounds. The wood is economically valuable. EUCALYPTUS SALIGNA Sm. Sydney blue gum, Saligna gum. 2n=22. Coasts and slopes of mountains in New South Wales and SE. Queensland. Cultivated in Africa, esp. Cameroon, Kenya and Zimbabwe ; S, America esp. Argentina and Brazil. After E. globulus*,the most widely distributed species in cultivation. The wood is extremely valuable. This is the most rapidly growing species in the genus, EUCALYPTUS SIDEROXYLON A. Cunn. ex Benth.Red ironbark. 2n=22. The W. slopes of New South Wales upland and in the N. part of Central Victoria, Australia. Cultivated in Africa esp. Cameroon, Kenya, Zaïre and Rhodesia; the Mediterranean area, esp. Spain, Portugal, Algeria, Morocco,Cyprus and Israel; Japan, USA and New Zealand. Its wood is economically very valuable. It contains essential oil.some spontaneous hybrids are known, EUCALYPTUS TERETICORNIS Smith, (syn. E. subalatum Cunningh.) Red gum,flooded gum,grey gum,blue gum. 2n=22. Almost the whole coast of E. Australia. Cultivated almost in all the countries of the world where Eucalyptus is grown. The wood is very valuable. In USSR, interspecific hybrids are produced. The strains 'C r of Zanzibar and 'Mysore Hybrid* of India belong to this species (Larsen & Croner, 970). EUCALYPTUS VIMINALIS Labill. (syn. E, mannifera Cunning, E. persicifolia Lodd,), Ribbon

62 MYRTACEAE - SOLANACEAE 69 eucalyptus t White gum, Swamp gum. 2n-22. SE. Australia and E. Tasmania. Cultivated in Mediterranean area, in countries of C. and S. Africa, India, New Zealand and USA. In subtropical areas of USSR, it is the commonest Eucalyptus species. The wood isof moderate value. An essential oil is obtained. Many spontaneous and artificial hybrids are known. LEPTOSPERMUM LAEVIGATUM F.V. Muell. Australian tea-tree. 2n=22. Australia. Cultivated there for the reclamation of moving sand 0 Dried leaves are used for tea-making. MELALEUCA PREISSIANA Schan. 2n=. Australia. Var. leiostachya Schan, (syn. M, parviflora Lindl.) is asoil stabilizer. NICOTIANA GOODSPEEDII Wheeler. 2n=40. New South Wales to SE. of West Australia. Ithas a short growing period. It is very resistant to Peronospora tabacina Adam. Some natural introgression with the closely related N. exigua Wheeler, 2n=32, N. nuaveolens Lehm., 2n=(24), 32, (48, 64), and N. rotundifolia Lindl., 2n=44. SOLANUM LACIANIATUM Aiton.Cut-leaved nightshade. 2n~92. Australia and New Zealand. Cultivated in Europe and elsewhere forits foliage, which is a source of steroid precursors. Proteaceae HAKEA SALICIFOLIA (Vent.) B.L. Burtt. 2n= SE, Australia and Tasmania. Cultivated for reclamation of arid land in Spain and Portu gal. Ithas runwild in those countries. HAKEA SERICEA Schrader. 2n=20. E. Australia. Cultivated for reclamation of arid land in Portugal and Spain, It has runwild in those countries. MACADAMIA INTEGRIFOLIA L.S. Smith, (syn. M. terni folia F.V. Muell., M. ternifolia var. integrifolia and M. tetraphylla L.A.S. Johnson). Queensland nut, Macadamia nut, Australian bush nut, Australian hazelnut. 2n=28, 56, E. Queensland, Australia. Cultivated in Hawaii. M. integrifolia is known as the smoothshell type and M, tetraphylla as the roughshell type, M, ternifolia isnow considered to apply correctly only to a species with bitter cyanogenic seeds less than 25 mm in diameter, inedible and never cultivated (Kraus & Hamilton, 970). Rutaceae EREMOCITRUS GLAUCA (Lindl.) Swing. 2n=8. This tree is capable of withstanding 6 months drought. It easily crosses with Citrus species giving fertile hybrids, Solanaceae DUBOISIA H0PW00DII F.V. Muell. Pituri, Pitchery, 2n=. Australia. Cultivated for some decades to yield atropine. DUBOISIA LEICHHARDTII R.Br. 2n=60. Australia. Cultivated for some decades for atropine, DUBOISIA MY0P0R0IDES R.Br. Corkwood, Mgmeo. 2n=60 Australia. Cultivated for atropine. NICOTIANA DEBNEYI Domin. 2n=48. Australia. Used as a source of resistance to blue mold caused by Perenospora tabacina Adam.

63 4 Hindustani Region TheHindustani Region of diversity was included by Vavilov inhis Tropical SouthAsianCentre oforigin. Zhukovskij (968) distinguished this centre onlyby number (IV), but in970 he indicated a distinction between thisand the rest ofs. Asia. Hebased his distinction on theexistence of species specific for thisregion 4. Although this region is nearknownold farming sites in Thailand,agriculturemust havebeen introduced fromthenw. adjacent area. Early farming sites have so farrevealed fewdetails ofnative cultivation. AtMüan-jo Daro (Mohen jodaro)andharappa on theriver Indus in Pakistan,almost on theboundary betweenregions 4and 5, asiteof theharappan culturewasdiscovered dating from BC.Someremains ofgossypium arboreumwere discovered.at a site, Navdatoli-Mahesvar on thenarbada River,in C. India,dating from 2000 BC., remainsof wheat, peas,broad beans, lentils, Lathyrus sativusand ricewere found.except for rice,thesecropswere domesticated outside India. Important crops of the region include bamboos,fruit trees, Cucurbita sativa,mangifera indica, Musa sp., Oryza sativa,phaseolus mungo,piper sp., Saccharum sinense and Vignasinensis. Speciesof this region have influenced the development of cropselsewhere, mainly by active distribution between this region and areas such asancient Egypt, Assyria,Sumeria and thehittite Empire. Exchange also took placewith Africa. Crops were distributed especially tothemediterranean areaby the Arabs in the8th-0thcenturies AD.Such crops include citrus trees, cotton species, jute,rice and sugar-cane. Acanthaceae BARLERIA PRIONITIS L. 2n=30, 40. Trop. Africa and Asia. Cultivated in India as amedicinal crop. Agavaceae SANSEVERINIA HYACINTHOIDES (L.) Druce (syn. S. zeylanica Willd.). Ceylon bowstring hemp. 2n=. Sri Lanka. A fibre plant cultivated there. Alliaceae ALLIUM AMPELOPRASUM L. Levant garlic, Perennial sweet leek. 2n=6, (24), 32, genome formula AAA'A", (40, 48). S. Europe, Asia Minor, Caucasus to Iran and N. Africa. Some cultivation in Germany and France (p. 48), Iran (Tarée irani, 2n=32; Tahbaz, 976)and in Kashmir (Koul k Gojil, 970). The wild and cultivated types are both extremely variable. This species is related to A. sativum*, A. porrum*, A. kurrat* and A. scorodoprasum*. Amaranthaceae AMARANTHUS ANGUSTIFOLIUS Lam. 2n=32, (34). S. and C. Europe, SW. Asia up to India and Turkestan and to Africa. In India var. polygonoides Thell. is cultivated. CELOSIA ARGEKTEA L.Quail grass. 2n=(36), 72. India. Var. cristata Kuntze (syn. C. cristata L.). Cockscomb grass. 2n=36. It is a pot-herb, fodder and fibre crop and an ornamental.

64 ACANTHACEAE - COMPOSITAE Anacardiaceae MANGIFERA INDICA L. Mango. 2n=40. Assam and the Chittagong Hills.Spread tomany tropical countries. Rhodes et al. (970) classified cultivars into :. polyembryonic group with oblong fruits,common in SE. Asia, 2. monoembryonic group with roundish fruits common in India and 3. agroup intermediate in fruit shape, also common in India. 4. the Sandersha-Haden complex, consisting of hybrids developed in Florida and Hawaii. M. odorata* and M. zeylanica Hook.f. are not closely related. Natural cross-fertilization ranges from 5 to 62%. A secondary centre of diversity has developed in Florida (p. 99). itscorras. ALOCASIA MACRORRHIZA (L.) Schott. Giant alocasia. 2n=26, (28). Probably Sri Lanka. Spread in the Malaysian Archipelago and to India and further to Trop. America. Several varieties are cyanogenic. A. indica is often included in this species. AMORPHOPHALLUS CAMPANULATUS Blume. Whitespot giant arum, Oroy. 2n=(4), 28. Trop. Asia (p. 49). Cultivated in India and elsewhere as a tuber crop. COLOCASIA ESCULENTA (L.) Schott. Taro, Dasheen. 2n=2x=28, 3x=42. The centre of domestication isnot certain. Some authors suggest Malaysia, others like Kuruvilla & Singh (98) NE. India,Assam or Upper Burma. These authors found 2x cultivars in the plains of India and 2x and 3x cultivars in the hills of NE. India. Asclepiadaceae MARSDENIA TINCTORIA R.Br. 2n=. Himalaya to China, Malaysia. Cultivated in India as a dye plant. Cannabidaceae CANNABIS SATIVA L. Hemp. 2n=20. Centre of origin C. Asia (p. 49). Spread to India in early times.the Indian type is cultivated for its narcotic properties. Thence it must have spread to the Middle East and elsewhere. 'C. ruderalis Janesch' is a weedy non-toxicant type. Chenopodiaceae Centre of domestication and routes of migration of Mangifera indica (Singh, 976). Apocynaceae NERIUM INDICUM Mill. (syn. N. odorum Soland.). 2n=22. Trop. Asia,especially India. Cultivated as a medicinal plant. RAUVOLFIA SERPENTINA Benth. 2n=(20), 22, (24, 44). India, Sri Lanka, Burma and from Thailand to Java. Because of the high demand for this medicinal crop, it became (nearly) extinct in some areas. To provide sufficient roots,some hospitals in India set up small gardens of it. Its cultivation could be extended to India and elsewhere (Dutta et al., 963). Araceae ALOCASIA CUCULLATA (Lour.) Schott. Giant taro. 2n=28. India and Sri Lanka.Cultivated for BETA VULGARIS L. Indian spinach. 2n=8, genome formula VV. For origin see p. 04.Indian spinach iscultivated mainly for its leaves and occasionally for its roots. There is a red-leafed and a green-leafed type. As it differs in chromosome pattern and for its phenolic compounds from table beetroot, Basu & Mukherjee (975)described it as B. palonga. However, more research is needed. It very likely derives from early introductions from the Near East via Afghanistan, whereas the table beetroot was introduced later from Europe (Basu & Mukherjee, 975), KOCHIA INDICA Wight. 2n=8. Introduced into Egypt where it is cultivated as a forage crop. Compositae INULA RACEMOSA Hook.f. 2n=20. Cultivated on a small scale in thelabaul Valley in NW. Himalaya for its aromatic roots. Cultivation is reported from N. Africa, Asia Minor, Ethiopia, Iran, E. India and some European countries. SAUSSUREA LAPPA Clarke. 2n=26. Grown in the

65 72 HINDUSTANI REGION Valley of Kashmir and adjacent area for its aromatic roots,, VERNONIA AMYGDALINA Delile. Bitterleaf. 2n=. Trop. Africa. Occasionally cultivated. Convoivulaceae IPOMOEA ERIOCARPA R.Br. 2n=. India.Used as a spinach and as a green fodder. Cruciferae BRASSICA CAMPESTRIS L. 2n=20. genome formula AA. See p. 50 for the origin of this species. In Pakistan/India the var. toria Duthrie & Fuller, Indian rape, Toria, and var. sarson Prain, Indian colza, Brown sarson are cultivated. Brown sarson can be divided into () self-compatible and (2) self-incompatible types. These two types can be differentiated by disruptive selection for flowering time, genetic drift in isolated populations, and chromosomal inversions suppressing recombination in connection with a recessive mutation of a major gene, independent of the S locus but inactivating that locus (Swamy Rao, 97). A considerable diversity isobserved in Bihar and E. Utter Pradesh (Anand et al., 975). ERUCA VESICARIA (L.). Cav. (syn. E. sativa L.). 2n=22. Mediterranean Region (p. 07)and Asia. Cultivated in India for jamba oil. RAPHANUS SATIVUS L. Serpent radish, Snakelike radish, Rat-tailed radish, Tree radish from Java, 2n=8. Cultivated from Java to NW. India. The plant requires a short daylength to develop roots.var mougri Helm (syn. R. caudatus L.,R. sativus var. indicus Sinsk. is characterized by small long fleshy fruits and glabrous leaves. Var. oleiformis Pers.* is also grown in India. Cucurbitaceae CITRULLUS COLOCYNTHIS* CITRULLUS LANATUS (Thunb.) Mansf. Water melon. 2n=22. Var. fistulosis (Stocks) Duthie & Fuller (syn. C. fistulosis Stocks), Round melon (2n=24)is cultivated in India for its round fruits. It differs in chromosomal number and leaf phenolics from C. lanatus and should probably be raised to a species (Kaur et al., 973). COCCINIA CORDIFOLIA Cogn. (syn. C. indica W. & A.).Ivy gourd, Small gourd. 2n=24, (36). Trop. Asia, and Red Sea area to Sudan. In S. India, forms occur with long less bitter fruits CUCUMUS SATIVUS L. Cucumber, Gherkin. 2n=4, Primary gene centre probably the Himalayas. Introduced into Europe. Near East, China and elsewhere. Secondary gene centres in China and Near East (p.36 and 89). In India, cucum- Raphanus sativus var. mougri (Sinskaya, 93). bers have been cultivated for at least years (Leppik, 965). Three varieties have been described: () var, hardwickii (Royle) Alef. (syn. C. hardwickii Royle) (a weedy type); (2) var. sikkimansis Hook, (the 'Indian type'); (3) var. squamosus Gab. Sources of resistance topowdery mildew Sphaerotheca fuliginea (Schlecht, ex Fr.) Poll are found in cucumber material from China, Japan,Indonesia and India. CUCURBITA MAXIMA Duch. ex Lam. Pumpkin. Winter squash. 2n=40. Its origin isdescribed on p. 69. Secondary centre in India and adjacent areas. LUFFA ACUTANGULA (L.) Roxb. Sponge gourd, Angled loofah, Sinkwa towel gourd. 2n=26. Probably India, Primary gene centre probably there. It was found in Karakoram/Hindu Kush in 955 (Tozu, 965). Cultivated in China and Japan. Dutt & Roy (97) suggested the following evolution of the various related Luffa species. They considered the wild monoecious L. graveolens Roxb. (2n=26) as the primary species. From it, two species derived: the wild dioecious L. echinata Roxb. (2n=26) and the cultivated monoecious L. aegyptica*. They considered dioecism as aderived factor that arose after divergence from the monoecious L. graveolens. From L. aegyptica, the monoecious type of L. acutangula and later the hermaphrodite type of that species arose, which is also called L. hermaphrodita*. LUFFA AEGYPTIACA Tull. (syn. L. cylindrica (L.) Roem.). Smooth loofah, Suakwa, Sponge gourd, Vegetable gourd. 2n=26 Domesticated probably in tropical Asia,possibly in India. Cultivated in almost all tropical regions where it may have run wild. Used for producing vegetables or sponges.it is also used for a medicine and for insulation. This species includes L. racemosa Roxb. (2n = ) with L. hermaphrodita* the only two

66 COMPOSITAE - GRAMINEAE 73 'species' with bisexual flowers. LUFFA HERMAPHRODITA Singh & Bhandari. Satputia. 2n=. Cultivated in Bihar and Bengal, India. It crosses easily with L. acutangula*, the Fi being monoecious. It is similar to L, acutangula* and to L, cylindrica (see L. aegyptiaca*), but has bisexual flowers, oblongellipsoidal fruits and smooth shin}^ black seeds. It may be a hybrid of one of those two species. Types described as L. racemosa Roxb. and included in L, cylindrica are also bisexual (Singh& Bhandari, 963). Dutt & Roy (97) included L. hermaphrodita in L, acutangula*. They condidered it as the hermaphrodite type of that latter species. TRICHOSANTHES CUCUMERINA L. (syn. T. anquina L.). Snake gourd. 2n=22. India to Australia. Cultivated for a long time in India. Dioscoreaceae DIOSCOREA HISPIDA*. Euphorbiaceae BACCAUREA SAPIDA Muell.-Arg. 2n=. Malaysia, India and China, Cultivated by Hindus for its fruits. CROTON TIGLIUM L. Purging croton. 2n=. SE. Asia.Cultivated now in India and Sri Lanka for its seeds (Purseglove, 968). Flacourtiaceae DORYALIS (DOVYALIS)HEBECARPA (Gardn.) Walb. (syn. Aberia gardneri Clos.). Ceylon gooseberry. 2n=. Trop. Asia especially India and Sri Lanka. Cultivated for its berries. HYDNOCARPUS LAURIFOLIUS (Dennst.) Sleumer (syn. Hydnocarpus wightianus Blume). 2n=24, India. Cultivated in several trop,countries for its oil, used to cure leprosy. Gramineae BAMBUSA ARUNDINACEA (Retz.) Willd. Spiny bamboo. 2n=70. India. Cultivated there. In forests bordering rivers and on mountains up to 900 m. Much cultivated in Java, rarely in Indochina. Secondary gene centre in Java (p. 53). Used as building material and in the paper industry. This isone of the largest bamboo species. Its stems can be 30 m long, BAMBUSA POLYMORPHS Munro. 2n=72. Bangladesh. Cultivated there and in Burma. Used asbuilding material and in the paper industry, BAMBUSA STRICTUS Nees. 2n=70, 72.In India (except Assam) and Burma.Secondary centres in Indochina (p. 53) ands. China (p. 37). In dry areas of forests. It is drought-resistant. BAMBUSA TULDA Roxb. 2n=. India. Burma (p. 53) and Tahiti. Secondary centre Java. CEPHALOSTACHYUM CAPITATUM Munro. 2n= and Himalaya. Its shoots are edible.. Burma CYMBOPOGON FLEXUOSUS (Nees ex Steud.) Wats, (syn. Andropogon flexuosus Nees). Malabar grass, Cochin grass. 2n=20(+l-2B), 40( + l-2b), x=0. India. Cultivated for its essential 'East India lemon-grass oil'. Var. coimbatorensis Gupta (2n=40) is a cultigen. CYMBOPOGON MARTINI (Roxb.) Wats. Roshagrass, Palmarosa, Ginger grass. 2n=20, 40(+l-2B), x =0. E. India. Cultivated in India and Indonesia for palmarosa oil and ginger grass oil. The cultigen var. motia (syn, C. motia Gupta, syn. var. martinii), 2n ; =20, produces palmarosa oil, while var. sofia Gupta, 2n=40(+l-2B) produces ginger grass oil. CYMBOPOGON PENDULUS Wats. 2n=20, 60, India. Recently taken into cultivation (Jagadishchandra, 975), CYN0D0N DACTYLON (L.) Pers. Bermuda grass.2n =(x=9) 8, genome formula AA; 36, genome formula AABB; 45, 54, genome formula AAAABB. Widely distributed in the Old World tropics. Malik & Tripathi (968)and Harlan & de Wet (969)indicate that the two diploid genomes of the tetraploid are essentially homologous. Var. aridus Harlan & dewet (2n=8) and var. afghanicus Harlan & de Wet are excellent fodders tolerant to drought. Several selections of var. dactylon are grown as lawn grasses. Var. elegans Rendle is a major natural grass of Africa, and var. coursii (Camus) Harlan & de Wet is a major fodder in Madagascar (Harlan et al., 970). See p. 28. DENDROCALAMUS HAMILTONII Nees & Arn. 2n= C. and E, Himalaya up to 900 m and Upper and Lower Burma up to 200 m altitude. Used as building material and in the paper industry. It grows in humid areas along rivers and in low places,and forms large thickets, DENDROCALAMUS LONGISPATHUS Kurz, 2n=72. Bangladesh and Burma (Arakan). Used for paper making. It is found in humid mixed forests a- long rivers on fertile clay soils. DIGITARIA CRUCIATA (Nees) A. Camus.2n= Var. cruciata grows wild in a large areaof E, India and China.Probably domesticated in the 9th Century in the Khasi Hills, E. India, by selecting var. esculenta Bor with longer stems, longer spikes,larger inflorescences and much bigger grains. Itgrows slowly and yields little. Its advantage is the production of straw in an area where little grass grows (Bor, 955;Singh & Arora, 972). DIGITARIA SANGUINALIS (L.) Scap. Manna.2n= 8, 36, 54, 72. Cultivated in Kashmir and

67 Y 74 HINDUSTANI REGION adjacent Afghanistan, and in SE. Europe, The species is a weed of temperate regions in all continents (Portères, 955a). Cultivated kinds differ little from their close wild relatives in floral morphology. ECHINOCHLOA COLONA (L.) Link. Shama millet, Jungle rice. 2n=(36, 48), 54, (72). Cultivated in India as a fodder grass and as a cereal. Formerly it was grown also in Egypt. Yabuno (968) considered the genome formula of this species tobe the same as that of E, frumentacea*. ELEUSINE CORACANA (L.) Gaertn. Finger millet, Ragi. 2n=36. Introduced from Africa (p. 29). Kempanna (969) recognized various types in India. Hilu & dewet (976) described how the African lowland race must have reached India from the south or south-west.the introduction must have taken place by sea routes c. 000 BC. In S. India,the crop developed a secondary centre of diversity. Thence it spread to the north. In NE. India,the Indian race developed and in N, India the North Indian type. It Isnot clear whether the last taxon evolved from the lowland race or from the Indian race or from both.. ^ -^.- ^ \.^" r^~* '^.. J " T 7 ^xxx ^ <-. (''" > * x- x ^-. i J O * ' " i ^* ' I XV / x " " -, > X o \5' S V 9p O no% Q^W vy] cp J~** <? 0 *f \o 00 0/ D o y \ «of w» B5 \oj V7\ u - ' > ', ; j J i "> '.o r 'v. ~V i. V > \ {^ ~ \ S / /Y V vr Various types of Eleusine coracana in India, (o) lowland race, ( ) Indian race,(x) Indian highland type (Hilu & de Wet, 976). V f is \s r \ mary centre: the Travancore Mountains and Tinneville (altitude m). Itis used in the paper industry. 0RYZA MALAMPUZHAENSIS Krish. & Chand. 2n=48. India. 0RYZA RUFIP0G0N Griff. Perennial wild rice. 2n=24. The more commonly accepted epithet 0. perennis Moench probably does not refer to Asiatic wild race (Tateoka, 964). Widely distributed in Asia and Australia. As recognized by Chang (976a, b),it includes the annual 0.ni vara SharmaSi Shastry (965a, b). The perennials areweakly rhizomatous with extra vaginal branching and are adapted to continuously flooded habitats. Cytogenetic studies suggest that under cultivation this perennial (Nayar, 973) gave rise to the annual cultivated 0. sativa (Chang, 964). Spontaneous annuals of Asia (0. sativa var, fatua Prain) are probably weedy derivatives of hybrids between 0, sativa and 0. rufipogon (Oka and Chang, 96), whereas Oceanic annuals may represent truly wild 0. nivara (Oka, 974). ORYZA SATIVA L. Rice. 2n=24, genome formula AA. S. Asia. The antiquity of rice cultivation is uncertain. Rice was probably collected as a wild cereal across the humid tropics ofasia. In parts of India and in Sri Lanka, perennial wild rice is still harvested as a cereal (Vishnu-Mittre, 974). The oldest known rice remains in the archaeological record (Allchin, 969; Solheim, 972) are from Mohenjodaro in Pakistan (2500 BC.), India (2300BC.) and Thailand (3500 BC.). According to esterase isozyme genotypes,the centre of diversity of rice is in Assam-N. Burma-Yunnan. In Assam, types resembling the African 0. glaberrima* are found (Seetharam & Ghorai, 976). Three races of rice are commonly recognized. () The basic tropical race indica probably originated over a broad region spreading from the Ganges Plains of India to Vietnam and S. China (Chang, 976a, c). It includes the floating rices of South Asia. (2) Race indica was introduced into the Yellow River (Huang Ho) Valley and the lower Chang (Yangtze) River MELOCANNA BACCIFORA (Roxb.) Kurz. 2n=. Burma from Garo to Arakan. Primary centre: Bangladesh. Its stems areused as building material. NEOHOUZEAUA DULLOSA (Munro) A. Camus. 2n= S. and SE. Burma. It has avery long fibre and is used widely to produce paper. OCHLANDRA TRAVANCORICA Bedd. 2n=c. 72. Pri- Centre of origin of rice (Chatterjee, 95).

68 GRAHINEAE - LEGUMINOSAE Basin., where the temperate race japonica e- volve ;d. Race japonica was introduced to Korea, and.aterjapan around the 3rd Century BC. (Mori naga, 968), (3) Race Indica also spread SE. nto the Malaysian Archipelago, where the.arge-grained race javanica evolved, which is widely cultivated across the islands of SE Asia. The wet rice cultivation of Phili ppines, Malaysia and Indonesia is quite a rec :ent introduction (Spencer, 963). PANICUM ANTIDOTALE Retz. Blue panic grass, Giant panic grass. 2n=2x=8. India, W. Saudi Arabia. Introduced into Australia as aforage grass. PANICUM SUMATRENSE Roth ex Roem. & Schult, (syn. P. miliare Lamk.). Little millet. India and Sri Lanka.P. psilopodium Trin. (2n= 54) might be its ancestor (Mansfeld, 959). It is possible that cytotypes with 36 chromosomes occur. PASPALUM SCROBICULATUM L. Khodo millet.2n= 20, 40. C. India. Widely cultivated in the Indian Plains as a cereal. Its closest wild relative, P. orbiculare Forst.,is widely distributed and collected as a wild cereal across S. Asia. SACCHARUM OFFICINARUM L. Sugar-cane. 2n various. Sugar-cane originated innew Guinea (p. 54) about BC. New Guinea Noble canes must have reached S. India via Indonesia. It was transported northwards to reach N. India ca 400 BC. C. 250 BC, it was severely attacked by red rot disease, which must have promoted the introduction of Sinense sugar-cane from China (p. 38). During migration, it hybridized with S. spontaneum*, as also happened in Orissa and Bihar (Parthasarathy, 946; 948; Bremer, i966). Originally the Sinense group of sugar-canes were named Pansahi sugarcane. In N. India (Punjab to Assam), the Barberi group (S. barberi Jesw.) of sugar-canes developed. Its origin isnot yet fully understood.the group can be classed into subgroups: () Mungo, 2n=8, 82, 83, without S. spontaneum introgression; (2) Dhaulo, 2n=82, 83; (3) Nargori, 2n=05-24 with Sinense introgression; (4) Saretha, 2n=9. SACCHARUM SPONTANEUM L. Wild sugar-cane, Kassoer,Thatch grass, Bagberi, Djarb, Khus. 2n =40-28, with euploids 40, 48, 64, 80, 96, 04, 2,28 and possibly 54. Probably India in the foothills of the Himalaya Mountains. Now it is distributed in innumerable groups of various ranks and significance from Africa over Asia to Japan and the Solomon Islands. One group (2n=2) is found in Indonesia (p. 55), while another one (2n=04-28) occurs in E. Africa (p. 33). Recently introduced into New Guinea and hence its influence there is still limited. S. spontaneum is used as a source of disease resistance of S. officinarum*. In N. India,it has hybridized with S. sinense* group Saretha. SINOCALAMUS GIGANTEUS (Walb.) Keng. 2n= India, Indochina and S. China. Cultivated there. Used as building material. It isone of the largest bamboos. SORGHUM BICOLOR (L.) Moench. Juar, Jowar. 2n= 20. Primary gene centre: Africa (p. 33). Secondary centre: India.No hybridization has occurred with wild sorghums, because they are tetraploids (Doggett, 970). TRITICUM AESTIVUM (L.) Thell. ssp.sphaerococcum (Perc.) MK. (syn. T. sphaerococcum Perc.). Indian dwarf wheat. 2n=42 f genome formula AABBDD. Transcaucasia and adjacent regions (p. 93). Ssp. sphaerococcum is indigenous to NW. India, Pakistan and adjacent parts of Afghanistan. It is characterized by short, non-lodging culms, erect leaves, globular grains and susceptibility to diseases. VETIVERIA ZIZANIOIDES (L.). Nash. Vétiver. 2n=20. Sri Lanka, India up to Burma. A grass cultivated in the tropics for its essential oil. ZEA MAYS L. 2n=20. Domesticated in C. America (p. 90). Secondary centre: S. Himalayas (Brandolini, 970), where flint maize (indurata Sturt.) is common. Guttiferae GARCINIA INDICA*. GARCINIA SILVESTRIS Boerl.Wild mangosteen. 2n=. Malaysia (p. 56) and India. Parental species of G. mangostana*. GARCINIA TINCTORIA*. MESUA FERREA L. Nahor,Nagas tree, Indian rose chestnut, Ironwood. 2n=32. Trop. Asia, India and Malaysia. Cultivated in India as a timber tree and for its flowers and fruits. The flowers are used in the perfume industry, the fruits are edible and the seeds contain oil for lamps. Lauraceae CINNAMOMUM TAMALA Nees & Eberm. 2n=. India. A tree, whose bark isused as a substitute for the spice from C. zeylanicum*. Leguminosae ALBIZIA STIPULATA Boiv. (syn. A. chinensis (Osb.) Merr.). 2n=. Cultivated in India and Sri Lanka for its high-quality fodder. Elsewhere it is cultivated as a shade tree, green manure and cover crop. BAUHINIA PURPUREA L. Camel's foot. 2n=28. Chi-

69 HINDUSTANI REGION na to India. A tree cultivated in India for various purposes and in Trop. Africa as a fodder plant. CAJANUS CAJAN (L.) Millsp. Pigeon pea. 2n=22, 44, 66. India, where wild plants of this species and the closely related wild Atylosia cajanifolia Haines,2n= occur (De, 974; van der Maesen, 980). De (976) suggests the forests of Upper W. Ghats as the domestication centre of pigeon pea. A secondary centre of diversity developed in Africa. The cultivars can be grouped into two groups : () var. bicolor DC. primarily large longlived late maturing with red or purple stained flowers and hairy purplish pods containing 4-5 usually purple-mottled or dark seeds; (2) var. flavus DC. mostly smaller and early maturing with all-yellow flowers and lightgreen pods containing 2-3 seeds. CANAVALIA ENSIFORMIS (L.) DC. Jack bean, Horse bean. 2n=22. S. America (p, 73). Secondary centre : India. CASSIA AURICULATA L. Tanner's cassia. 2n=4, 6, 28. Cultivated there for tanning material, obtained from thebark (Purseglove, 968). CASSIA FISTULA L, Indian laburnum, Purging cassia. 2n=24, 26, 28. India. Cultivated in the tropics for its pods, the pulp around the seeds being used as a purgative (Purseglove, 968). CASSIA SIAMEA*. CICER ARIETINUM L. Gram, Chick-pea. 2n=4, 6, (24, 32, 33). Probably W. Asia (p. 96). Probably asecondary centre in India. Introduced into India inearly times and now much cultivated there.strains with fine darkbrown and black seed have been cultivated for a long time. 'Kabuli' types were introduced from Afghanistan about 700 (van der Maesen, 972). CROTALARIA BURHIA Buch.-Ham. crop comes from E. India. 2n=6. This fibre CROTALARIA JUNCEA L. Sunn hemp,sann hemp.2n =6. Probably India. Unknown wild. A bastfibre crop. Cultivated in many tropical countries as a green manure (Purseglove, 968). CYAMOPSIS TETRAGONOLOBA psoralioides DC.). Guar, Probably domesticated in 960). However Hymowitz seeds of C. senegalensis with Arab-Indian horse t came into domestication in India, Pakistan and e food and as a source of in the Indo-Pakistan sub (L.) Taub. (syn. C. Cluster bean. 2n=4. Africa (Anderson, (973)described how * arrived in Africa rade. Subsequently it in India. Cultivated lsewhere for fodder, gum.no wild forms continent. D0LICH0S UNIFLORUS L'am. (syn.d. biflorus auct. non Linn.). Horse gram. 2n=20, 22, (24). Tropics of Old World, especially in India and Himalayas where it is also cultivated. INDIGOFERA PILOSA Poir. 2n=6, 32. Used in Sri Lanka as a green manure, MELILOTUS INDICUS All. Indian clover, Yellow annual sweet clover,sour clover. 2n=6. Punjab, India to the Mediterranean area and Turkestan. Cultivated in N. India as a fodder crop and inusa as cover crop. MUCUNA CAPITATA*. MUCUNA PACHYLOBIA (Piper & Tracy)Rock (syn. Stizolobium pachylobium Piper & Tracy). Fleshy pod bean.2n=. India. Cultivated as vegetable. MUCUNA UTILIS Wall, ex Wight. Velvet bean.2n =. India. Cultivated as vegetable, cattle food and green manure. SESBANIA ACULEATA (Pers u) Poir. Dhanchia. 2n =(2), 24, 32. Cultivated in Bengal for its fibre and especially as green manure. SESBANIA AEGYPTIACA*. SESBANIA SPECIOSA Taub, ex Engl. 2n=2. India (?). Cultivated there as green manure in rice fields, VIGNA AC0NITIF0LIA (Jacq.)Maréchal (syn. Phaseolus aconitifolius Jacq., Vigna lobata (L.) Verde., Phaseolus trilobus Wall.). Mat bean, Moth bean. 2n=22. India, Bangladesh and Burma. Cultivated in these countries and also on Sri Lanka and in China. InUSA it is cultivated as a fodder crop. VIGNA MUNGO (L.) Hepper (syn. Phaseolus mungo L.). Black gram, Urd, Urid. 2n=22, (24). India. Unknown wild. Closely related tov. radiata*. V. radiata var. sublobata (Roxb.) Verde, (syn. Phaseolus sublobatus Roxb.) (2n= 22) is most likely the wild parent. In Pantnagar, strains of this variety are intersterile (Singh & Ahuja, 977). VIGNA UNGUICULATA (L.) Walp. (syn. V. sinensis (L.) Savi, Dolichos sinensis L.). Cowpea, Black eye, Southern pea. 2n=22, 24. Primary centre of diversity: W. Africa, Secondary centre : India. Probably originally domesticated in W. and C. Africa (p. 38). From Africa, cowpea was taken to India and later it spread from both those regions over the world.. Ssp. mensensis Verde, grows wild in the African forest zone and ssp. dekindtiana Verde. wild in the African savanna zone. Most cultivars belong to ssp. unguiculata and the yard-long bean and asparagus bean cultivars to ssp. sesquipedalis (L.) Van Eseltine are rare in Africa. These last two subspecies evolved from ssp, unguiculata (Summerfield et

70 LEGUMINOSAE al., 974). In N, Nigeria, a form developed for the fibre obtained from the peduncles. Other uses are food, green manure, forage and cover crop. Liliaceae IPHIGENIA STELLATA Blatter. 2n= A source of colchicine. Linaceae. W. India. LINUM USITATISSIMUM L. Flax. 2n=30, (32). Its possible origin is given on p. 99. In India and adjacent areas, flax of ssp. indo-abyssinicura Vav. & Ell. is found. It is identical to flax of Ethiopia including Eritrea and it may have originated there. It hybridizes with ssp. mediterraneum* resulting in a hybrid ssp. hindustanicum Vav. & Ell. Malvaceae ABELMOSCHUS ESCULENTUS (L.) Moench (syn. Hibiscus esculentus L.). Okra, Lady's finger, Gombo. 2n= Probably a cultigen developed from a wild species A. tuberculatus Pal & Singh (2n= )in trop. Asia, It could be the ancestral or wild form of A. esculentus (van Borssum Waalkes, 966; Bates, 968). It is a N. Indian species differing from A, esculentus only in having strigose pubescence on the stems and shorter capsules beset with bristly tübercul ate hairs. Van Borssum Waalkes suggested that A. tuberculatus be included in A. esculentus. ABELMOSCHUS MANIHOT (L.) Medikus, (syn. Hibiscus manihot L.). 2n=60, 66. India, Pakistan, through S. China tonew Guinea and N. Australia. Cultivated for its immature fruits. Ssp. manihot is the cultigen that must have been selected by man from wild hairy and prickly types (ssp. tetraphyllus (Roxb. ex Hörnern.) Borss., syn. Hibiscus tetraphyllus Roxb. exhörnern.). ABELMOSCHUS MOSCHATUS Medikus, (syn. Hibiscus moschatus L.). Musk mallow. 2n=72. India, S. China, Indochina to Indonesia and SW, Pacific islands tonew Guinea and N. Australia. Centre of origin possibly E. India (Mansfeld, 959). Cultivated for its seeds, which are used as perfume, for its immature edible fruits,its fibre and often as an ornamental. ABUTILON INDICUM*. GOSSYPIUM ARBOREUM L. Tree cotton. 2n=26, genome formula A0A2. Primary centre: Indian subcontinent. Unknow wild. Spread E, and SE.to Burma, Indochina and the Malaysian Archipelago. Centre of domestication probably Gujarat, which is the westernmost state of India (Hutchinson, 97). Close to this area, afragment of textile and a string dated BC. have been found at Mohenjo-Daro, Pakistan. Race indicum of Indian subcontinent is more closely related to cottons belonging to G. herbaceum* than are other races of 0. arboreum. It includes both perennial and annual forms. It is very likely that the perennial forms are primitive and that the annuals were selected later. In N. India and Pakistan, race bengalense was cultivated. Perennial forms are occasionally found in remote places in Rajputana and in the Ganges Valley. Itspread S,, SE. and W. The African race soudanense* was probably the cotton cultivated by the people of Meroë (an ancient Nubian civilization), who were the first in Africa to spin and weave cotton. Chowdhury & Buth (970) suggested, that this race might be indigenous to Africa rather than introduced from India as a textile crop. GOSSYPIUM ST0CKSII Masters. 2n=26, genome formula EE. Sind, India and SE. Arabia. Itis drought-resistant 0 HIBISCUS RADIATUS Cav. 2n=72, genome formula AABB. India to Burma. Introduced into Africa and elsewhere where it is cultivated. Mainly used as ornamental; cultivated in Java as vegetable and drug plant. It is a sourceof resistance to root-knot nematode for H. sabdariffa.it is an allotetraploid of H. cannabinus* and another diploid species.ifit is indigenous to India, the amphiploidization must have taken place there after the introduction of H. cannabinus as a fibre crop. However the only known diploid B-carrier is the African H. surattensis* (Menzel & Martin, 97). If the origin of H. furcatus Roxb. non Willd. (2n=44), genome formula BBGGWWZZ from India and Sri Lanka is studied, the origin of H. radiatus may also be solved. SIDA RHOMBIF0LIA L. Queensland hemp, Broomjue sida, Cuba jute. 2n=4, 28. Tropics. Cultivated in India and later in Queensland as a fibre crop (var.retusa L.). It is an extremely variable species. Moraceae ARTOCARPUS HETEROPHYLLUS Lara. (A. intégra (Thunb.) Merr., A. integrifolia L.f.). Jack fruit, Jak. 2n=56. Unknown wild. Ithas a very ancient cultivation in India. There and in Sri Lanka it is popular; it is also cultivated elsewhere in the tropics (Purseglove, 968). FICUS ELASTICA Roxb. Indian rubber tree, Karet tree. 2n=26, (39). India and Malaya 0 Cultivated in India, Java and elsewhere. It is also cultivated as an ornamental house-plant (Purseglove, 968). FICUS RELIGIOSA L. Pipal tree, Peepal tree, Bot tree. 2n=26. This strangling fig is sacred tohindus and Buddhists. It is propagated by cuttings and layering. A scion planted at

71 78 HINDUSTANI REGION cuttings and layering. A scion planted at Anuradhapura insri Lanka in 288 BC. (Purseglove, 968)died in 97. FICUS ROXBURGHII Wall, its figs. Moringaceae 2n=. Cultivated for MORINGA OLEIFERA Lam. Horse-radish tree, Drumstick tree. 2n=28. India. Cultivated throughout the tropics as a vegetable, in hedges and for its fruits, whose oil is used for lamps and cosmetics. Musaceae MUSA cultivars of the AB group.ney poovan (and other names). 2n=22.Cultivated on a small scale now. See p. 52 for discussion. MUSA cultivars of the AAB group. 2n=33. Mostly India. Only the clone Mai a maoli has probably arisen in Philippines. MUSA BALBISIANA Colla. Pisang bau, Klue Tani. 2n=22 t genome formula BB. India, Burma, Sri Lanka and E.New Guinea. Cultivated for its leaves as a packing material or as a fibre plant (de Langhe, 969). It isnot very variable. It isone of the parents of the AAB, ABB and ABBB groups (p. 59)of cultivated bananas. Myrtaceae EUGENIA JAMBOLANA*. RHODOMYRTUS TORMENTOSA*. Oleaceae JASMINUM GRANDIFLORUS L. Catalonian jasmine, Italian jasmine. 2n~26. Himalayas. Cultivated for its fragrant flowers and used in perfumery. JASMINUM SAMBAC (L.). Ait. Arabian jasmine. 2n=26, 39. India and Sri Lanka. Cultivated in tropics. Flowers are used for scenting tea and as source of an essential oil. Palmae ARENGA PINNATA (Wurmb.) Merr.Sugar palm. 2n=. Primary centre: possibly Malaysian Archipelago (p. 6). Secondary centre: India. Cultivated inmany trop. Asian countries. BORASSUS FLABELLIFER*. COCOS NUCIFERA L. Coconut palm. 2n=32. SE. Asia, Indonesia andw. Pacific islands. Possible secondary centre:india. Pedaliaceae SESAMUM INDICUM L. Sesame, Beni seed. 2n=26. Primary centre: Africa (p. 44). Secondary centre: India. Thence it is believed to have spread E. through China, Indochina into Japan, and W. through Afghanistan, Asia Minor and Iran to the Mediterranean area including N. Africa. Perioplocaceae CRYPTOSTEGIA GRANDIFLORA*. Piperaceae PIPER LONGUM L. Indian long pepper, Jaborandi pepper. 2n=24, 48, 52, 96. The foot of Himalayas. Cultivated in India and Sri Lanka for its spike, which isused as a spice. It resembles P. retrofractum Vahl*. PIPER NIGRUM L. Black pepper. 2n=36, 48, 52, 60, 04, 28. The slopes of mountains in the Ghats, Malabar, SW, India at an altitude between 50 and m. Spread to SE. Asia and Philippines. Now cultivated in other trop, countries (Gentry, 955;de Waard & Zeven, 969). Plantaginaceae PLANTAG0OVATA Forsk. (syn.p. decumbens Forskc). 2n=8. Mediterranean area, C. Asia and India. Cultivated in India (esp Gujarat) for its seeds used in medicine. Polygonaceae RUMEX VESICARIUS L. 2n=8, (20). Greece, N. Africa, India and Malaysia. Cultivated in India as a medicinal plant. Resedaceae RESEDA 0D0RATA L. Mignonette. 2n=2 (4). For origin see p. 8. Var. neilgherrensis (Muell. -Arg.) Abdallah &de Wit is found on Nilagiri Mt and inbombay area and Decca Peninsula (Abdallah & de Wit, 978). Rosaceae RUBUS ALBESCENS Roxb. Mysore raspberry. 2n=. Mountains of India, Sri Lanka, Malaya and Indonesia. Occasionally cultivated, especially inpuerto Rico. Rubiaceae C0FFEA BENGALENSE Heyne ex Willd. 2n=22. Bengal, Burma and Sumatra. Occasionally cultivated in India (Purseglove,968) MORINDA ANGUSTIFOLIA Roxb. 2n=. Trop. Himalayas,Assam and adjacent areas. Cultivated forbark and wood, as source of yellow dye.

72 MORACEAE -VERBENACEAE 79 MORINDA CITRIFOLIA L. Indian mulberry. 2n= up to Pacific isles, dye crop. (syn. M. bracteata Roxb.).. S. India and Malaysia Cultivated in India as a RUBIA CORDIFOLIA L. Indian madder. 2n=22. Trop, and temp. Asia. Cultivated in India as adye plant. Rutaceae CITRUS LATIPES (Swing) Tan.2n=. Hills of NE, India. The fruits are not edible. It can be crossed with other Citrus species, including the cultigens. CLAUSENA DENTATA (Willd.) Roem. (syn. C. willdenowii Wight & Arn.). 2n=8, 36, India. This small tree is known for its edible berries, FERONIA LIMONIA (L.) Swing, (syn. Limonia acidissima L.). Indian wood apple. Elephant apple. 2n=8, India and Sri Lanka.Now cultivated in several trop, countries for its fruits MURRAYA KOENIGII (L.) Spreng. Curry-leaf tree. 2n=8. India. Cultivated for its leaves. Sapindaceae SAPINDUS TRIFOLIATUS L. Soap-berry tree, Soapnut tree, Arceta.2n=. India, Pakistan and Sri Lanka. Cultivated for its fruits, which yield soap, Sapotaceae MADHUCA INDICA Gmelin (syn. M. latifolia (Roxb.) Macbr. Moa tree.2n=. N. and C, India. Cultivated for its flowers, which are rich in nectar. MADHUCA LONGIFOLIA (Koenig) Macb. (syn. M. indica J, F, Gmel.), Mahua, Mowra butter tree. 2n=. India. Cultivated there. MANILKARA HEXANDRA (Roxb.) Dubard. 2n=, S. Asia.Cultivated in India. Solanaceae ATROPA BELLADONNA L. Belladonna. 2n=(50), 72. From Spain, the Balkans, Asia Minor to India. Medicinal plant. In India, var. acuminata (syn, A, acuminata Royle, 2n=72) is found. Itis cultivated there. CAPSICUM ANNUUM L. Bell pepper, cayenne pepper. 2n=24. Mexico (p. 96). Secondary centre: Asia. DATURA METEL L. Hindu datura. 2n=24. India. Medicinal plant introduced into many parts of (sub)tropics. Often cultivated as ornamental, SOLANUM MELONGENA L. Egg-plant, Aubergine, Bringal, Melongene,2n : probably India. Awild' small fruits,sometime is found on the Bengal haps it is a de-domest Martin & Rhodes (979) some characteristics o point to reproductive fertilizer or to human Primary centres inind: (p. 46). S. melongena S.incanurn*. Strychnaceae =24, (36, 48). Very looking form with many s called var. insanum, Plain of India, Pericated run-wild type, found associations of f cultivars. This could isolation of this selfselection pressure. ia, secondary in China is closely related to STRYCHNOS NUX-VOMICA L, Strychnine tree, 2n= 24, 44, India,Sri Lanka and Indonesia. Cultivated for its nux vomica used in medicine. Theaceae CAMELLIA SINENSIS (L.) 0. Kuntze (syn. C. thea Link., Thea sinensis L.). The origin of tea is discussed on p. 39. Secondary centres: India, Assam and Sri Lanka. Tiliaceae CORCHORUS CAPSULARIS L. Jute, White jute, 2n=4. Unknown wild. Primary centre: India and Pakistan. It has been cultivated there for a long time o Spread now throughout the tropics, GREWIA ASIATICA L. Phalsa, 2n=36. Salt Range, Puna, India (Mansfeld, 959). Cultivated in India, Sri Lanka and elsewhere for its fruits. Umbelliferae ANETHUM GRAVEOLENS L. (syn. Peucedanum graveolens (L.) Hiern.). Dill. 2n=22, Eurasia. Cultivated in Greece, the district of Rome and Palestine (p. 8). Indian types have longer fruits. This species includes A. sowa*. CARUM COPTICUM (L.) Benth. & Hook. (syn. Trachyspermum ammi (L.) Sprague). Ammi, Lorage. 2n=(6), 8. India,It yields an essential oil. Urticaceae GIRARDINIA HETEROPHYLLA Dene, (syn. Urtica heterophylla Roxb.). Nilgeri nettle, 2n=20. From NW. Himalaya to Malaysia. In the Nilgeri area, India var, palmata is cultivated. MAOUTIA PUYA Weddell (syn. Boehmeria puya Hassk., Urtica puya Wall.). 2n=. Perennial herb of trop. Himalaya, Khasia and Burma, Occasionally cultivated for its fibres. Verbenaceae NYCTANTHES ARBOR-TRISTIS L.Tree of sadness. 2n=44. C. India, Planted near temples. Itis a source of a saffron-yellow dye. The oil is used in perfumery. Cultivated as ornamental.

73 80 HINDUSTANI REGION Zingiberaceae AMOMUM AROMATICUM Roxb. Bengal cardamon, Nepal cardamon. 2n=.India and Pakistan. Cultivated there.. Burma. Cul AMOMUM XANTHIOIDES Wall. 2n= tivated in India. CURCUMA AMADA Roxb. 2n=42. India and Pakistan. Cultivated in India for Mango ginger. CURCUMA ANGUSTIFOLIA Roxb. East Indian arrowroot. 2n-42, (64). Himalayan area. Cultivated for its edible starchy rhizomes. CURCUMA CAESIA Roxb. Kalihaldi.2n=. Bengal. Occasionally cultivated for its edible rhizomes. CURCUMA DOMESTICA Val. (syn. C. longa Koenig nonl.). Turmeric, Curcuma. 2n=(32), 62, (63, 64). A completely sterile triploid, which probably arose with thewild C, aromatica Salisb. (Wild turmeric, Yellow zedoary; 2n=42)asone parent. C. aromatica grows in India. At first, turmeric may have been used as a sacred plant; later it was cultivated for its rhizomes, which are used for flavouring, and for colouring food and cloth. It spread in early times to China, SE. Asia and later to other parts of the tropics. There it may have run wild (Purseglove, 972). CURCUMA ZEDOARIA Rose. Zedoary. 2n=63, 64. E. India. Cultivated in SE. Asia, Sri Lanka and Madagascar. Rhizomes are a source of a condiment. Young flowers are used for flavouring food. ELETTARIA CARDAMONUM (L.) Maton. Cardemon. 2n=48, (52). S. India and Sri Lanka. Cultivated in India and Malaysian Archipelago. Seeds of cardamon are used to flavour coffee and in cooking; the oil is used for perfumery. The cultivated types can be divided into three varieties and two groups:group major Thwaites includes the Sri Lanka variety; group minuscula Burkill (syn. var. minor Watt) includes the Malaban variety and Mysore variety. Wild types have also been described as belonging to major. Plants of this group are marked by anthocyanin pigmentation. ZINGIBER OFFICINALE Rose. Ginger. 2n=22. Unknown wild. Probably domesticated in India. It was known in China and Trop. Asia at an early date. Cultivated now throughout the tropics.

74 5 Central Asian Region TheCentral AsianRegionwas called Southwestern AsianCentreby Vavilov. Zohary (970) preferred to join it withregion 6, the NearEastern Centre, asdidzhukovskij (968),who separated both areasbynumber only.but in 970,both centres wereon the map,separatedby a line,andregion 5 was extended northwards (Zhukovskij, 970). This centre served as atransfer zonebetweenregions and 4. Furthermore, thehimalayas haveprovided many species asparental stock for crops. Agriculturemusthave reached this centre fromregion 6about 5000 BC. Majorcropsof thiscentre include fruit-trees, Allium cepa, A. sativum, Daucus carota, Lathyrus sativus,spinacea oleracea andvicia faba.cucumis melohas there asecondary centre of diversity. Alliaceae ALLIUM CEPA L. Onion. 2n=6. Cultivated since 000 BC. Probably C. Asia and esp. NW. India, Afghanistan, Uzbekistan and W. Tien Shan, where related species A. pskemense 0. Fedtsch. (2n= 6) and A. vavilovi Popov & Vved. (2n=6) grewwild (Vavilov, 949/50). Secondary centre is Region 7. Another related wild species is A. oschaninii 0. Fedtsch. (syn. A. cepa var. sylvestre Regel) from Pamirs, Alai and Tien Shan. This species is used as food. Wendelbo (97) and McCollum (974) suggested this species to be the wild A. cepa or thewild ancestor of A. cepa. The top onion, tree onion or Egyptian onion (A. proliferum*, syn. A. cepa viviparum (Metzger) Alefeld) is a hybrid ofa. cepa and A. fistulosum* (Flskesjö, 975). Such hybrids have also been described as A. aobanum Araki, 2n=6 and A. wakegii (see A. chinense*). A. cornutum G.C. Clementi ex Vis., 2n= near Budva, SW. Yugoslavia is probably a bulbilliferous form derived from extinct cultivation (Tutin et al., 976). Var. ophioscorodon (Link) Doli has a stem coiled in one or two wide loops before anthesis, ALLIUM PROLIFERUM. 2n=6. Tree onion, Egyptian onion. Formerly described as A. cepa* var. proliferum and as A. fistolosum*, but a true natural hybrid of both these species (Fiskesjö, 975; Vosa, 976). Propagated by bulbs. Viviparous. ALLIUM SATIVUM L. Garlic. 2n=6, genome formula SS. Some consider A. longicuspis Regel (2n=6) as the wild parent of garlic. This species occurs inc. Asia. Var. pekinense (Prokh.) Makino came into cultivation in China (p. 53). Secondary centres developed in Regions 6 and 7 (Kazakova, 97). Garlic was already known in Egypt before BC. Sterile, propagated with bulbils. Anacardiaceae PISTACEA VERA L. Pistacio. 2n=30. Mountain slopes, sometimes forms sparse forests. Cultivated in this centre and elsewhere for its seeds. In Turkmenia, there are relic populations, which indicate the once wide distribution of this species (Kabulov, 969). Dioeceous. In orchards ofp. atlantica (2n=28) grown for rootstocks in USA, hybridswith P. vera form and develop female and male branches. They could perhaps be used tobreed monoeceous trees. Chenopodiaceae BETA VULGARIS L. Beet. 2n=8, (27, 36). Pri-

75 * ^ i J CENTRAL ASIAN REGION mary centre is discussed on p. 04. Secondary centre developed in Region 5. SPINACEA OLERACEA L. Spinach. 2n=2. Iran to Manchuria. Primary centre in Afghanistan and Tajikistan. Related to S. tetrandra* and S. turkestanica*. An anemophilous species of which the very young seedlings are eaten as a vegetable. SPINACEA TETRANDRA Stev. 2n=2. A wild relative of S. oleracea* occurring in the stony steppes of Caucasia including Armenia, and Kurdistan. A weedy anemophilous species. SPINACEA TURKESTANICA Iljin. 2n=2. A wild relative of S. oleracea* occurring in the loess foothills of the Kara Kum, the southern foothills of Uzbekistan and Turkmenia, and along the Syr Darya. A weedy anemophilous species. Compositae ARTEMISIA CINA Berg.Levant wormseed plant, 2n=8. The Orient and Russian Turkestan. Cultivated in Russia and W. of USA. ARTEMISIA DRACUNCULUS L. Tarragon, Estragon. 2n=8, 36, 54, 72, 90. USSR and from W. Asia to Himalaya. Perennial widely cultivated as a condiment. 'Russian' tarragon (2n=90), a decaploid, is fertile, whereas the 'French' tarragon (2n=36), a tetraploid, is sterile and propagated vegetatively. The types with (2n=45, 54, 72) may be hybrids (Rousi, 969). A, dracunculoides Pursh (2n=8) from N. America has often been included in this species. CARTHAMUS TINCTORIUS L. Safflower. 2n=24, genome formula BB. Primary centre in Region 6 (p. 88). Secondary centre near Kabul, Afghanistan. The great variation of the safflower population there,that led Vavilov to believe that area to be a centre of origin, is very likely caused by the meeting of the Middle Eastern and West Pakistan safflower types there (Knowles, 969). (snake melon) (2n=24), var. tarra Pang, and ssp. agrestis (Naud.) Greb. (2n=24), var. agrestis Pang.The last is a weedy field melon. Datiscaceae DATISCA CANNABINA L. 2n=22. C. Asia. A herb. Cultivated as a source of yellow dye. Ebenaceae DI0SPYR0S LOTUS L. Caucasian persimmon. 2n= 30. Subtrop. China (p. 36) in Talysk andw. Georgia (USSR) and adjacent Iran. Both these areas form primary centres. Elaeagnaceae ELAEAGNUS ANGUSTIFOLIA L. (var. E. argenta Moench). Silverberry, Russian olive. 2n=2 t 28, From S. Europe to C. Asia, China and Himalayas. Primary centre in C. Asia. Cultivated there and in Iran for its edible nuts. E. orientalis L. has been included as var. orientalis (L.) 0. Kuntze. in this species. Gramineae AEGILOPS CAUDATA* AEGILOPS CYLINDRICA* AEGILOPS JUVENALIS* AEGILOPS KOTSCHYI* AEGILOPS LORENTII* AEGILOPS OVATA* AEGILOPS SQUARROSA auct. non, L. (syn. Ae. tauschii Cosson, Triticum tauschii (Coss.) Schmalh.), 2n=4, genome formula DD. E, Turkey, Iraq, Crimea and Caucasia (Zohary et al. 969) in the west, and Pakistan and Kashmir in the east. Primary centre to the south of the Caspian Sea. This wild species is the D INULA HELENIUM L. (syn. Helenium grandiflorum Gilib.). Elecampane, Elfdock, Horseheal, Yellow starwort. 2n=20. Probably C. Asia. Also wild westwards to C. and S. Europe. It used tobe cultivated and had various uses. TARAXACUM KOKSAGHYZ Rodin (syn. T. bicorne Dahlst.). Kok-saghyz. 2n=6. Turkestan. Cultivated inussr as rubber crop. Cucurbitaceae ƒ <3 / / i L / CUCUMIS MELO L. Melon,Musk melon, Canteloupe 2n=24. Probably Africa (p. 24). Secondary centre in Region 5, in which are found ssp. melo Pang., convar. chandalak (Pang.) Greb,, convar. ameri (Pang.) Greb., convar. zard (Pang.) Greb., ssp.flexuosus (L.) Greb. K -x Aegilops squarrosa \/^ si. \

76 CHENOPODIACEAE - LEGUMINOSAE 83 genome parent of T. aestivum*. Ssp. eusquarrosa Eig (syn. Ae. tauschii ssp. tauschii) usually has short anthers and is generally strictly autogamous, whereas ecotypes with long anthers are facultatively allogamous and resemble ssp. strangulata (Eig) Tzvel. (Hammer & Knüpffer, 979). AEGILOPS TRIARISTATA* AEGILOPS TRIUNCIALIS* SECALE CEREALE L.Weedy and cultivated rye. 2n=4. The origin of this species is discussed on p. 9. Secondary centre in E. Iran and Afghanistan, where S. afghanicum (Vav.) Roshev. (syn. S. cereale ssp. afghanicum (Vav.) Khush) originated. The main stream of cultivated rye spreading over Europe and Asia comes from the secondary centre (Khush, 963). Khush based his conclusion on the pigmented ears of all cultivated rye varieties and those of the weedy rye types in Afghanistan. They occur in grain fields with ahabit and growth rhythm similar towheat (Stutz, 972). Var. eligulatum Vav., liguleless rye was found by Vavilov in the secondary centre. SECALE TURKESTANICUM* TRITICUM AESTIVUM (L.) Thell. ssp.compactum (Host.) MK, (syn. T. compactum Host.). Club wheat. 2n=42 f genome formula AABBDD. Ssp.compactum developed in the mountains of the Hindu Kush. TRITICUM TURGIDUM ssp.turgidum conv. polonicum (L.) MK. 2n=28 t genome formula AABB.The type 'T. ispahanicum Heslot' was found in Isfahan (Iran) where it is adapted to irrigated cultivation. It is marked by its narrow and elongated glumes. Hippocastanaceae AESCULUS HIPPOCASTANUM* Juglandaceae JUGLANS REGIA L. Walnut. Persian walnut, English walnut. 2n=32, 36. Secondary centre: Moldavia, SE. Europe and SW. Europe (p. 55). Many varieties have been described. Labiatae LALLEMANTIA ROYLEANA (Wall. & Benth.) Benth. 2n=4. From Iran to Himalayas. Cultivated in E. India as an oil crop. Leguminosae CICER ARIETINUM L. Chick-peas, Gram. 2n=4, 6, (24, 32, 33). Secondary centre in Afghanistan, whence the 'Kabuli' types of India derive. They were introduced there about 700 Juglans regia (van der Maesen, 972). CICER MICROPHYLLUM Royle (syn. C. songaricum Steph., C. jaquemontii Jaub. & Spach.). 2n= 4. Tibet, Afghanistan andw. Himalayas. Cultivated in W. Himalayas for its seeds. FLEMINGIA VESTITA Benth. ex Baker (syn. Moghenia vestita (Benth. ex Baker) 0. Kuntze. Sohphlong. 2n=22. W. Himalayas, N. India. Cultivated for its edible tubers,especially in Assam. LATHYRUS SATIVUS L. Grass pea, Chickling pea. 2n=4. Probably W. Asia. A centre of diversity in the Mediterranean Region.Cultivated in Europe since ancient times and in Region 7 (p. 4). Unkown wild. MEDICAGO SATIVA L. Lucerne, Blue alfalfa. 2n= 6, genome formula SS, 32, genome formula SSSS. Transcaucasia (p. 97). The centre of diversity of blue lucerne is in NW. Iran and adjacent regions to Tibet. VICIA FABA L. (syn. Faba vulgaris Moench.). Field bean,broad bean,horse bean, Tick bean, Windsor bean,faba bean. 2n=2, 4, Probably SW.Asia (Ladizinsky, 975c) or Mediterranean region (Zohary, 977). Wild ancestor uncertain. It may derive from the weed V. angustifolia L. (2n=4). Formerly it was believed to derive from V. narbonensis, but Ladizinsky (975c) and Abdalla &Giinzel (979) showed that this species was not related tothe field bean. Ladizinsky (975c) also showed that V. galilaea Plit. & Zoh. (2n=4) and V. hyaeniscyamus Mount. (2n=4) were not related either. Related species are V. bithynica (L.) L. and V. johannis Tamanschian. The field bean is divided into three varieties according to the size of the seed: var. minuta (Alef u) Mansf. (syn. var. minor (Pieterm.) Harz,,), pigeon bean or tick bean, var. equina Pers., horse

77 CENTRAL ASIAN REGION bean, and var. faba (syn. var. major Harz.), broad bean. From its centre of domestication, the field bean spread to Europe, China and the Mediterranean. In the Mediterranean region, a secondary centre of diversity arose (p. 6). The pigeon and horse beans are used as animal feed, and the broad bean as a vegetable. Schultze-Motel (972) has listed all data on archaeological remains of the field bean. Almost all these remnants belong to var. minuta. He did not find asharp distinction between long-seeded and roundish types,so the development of the field bean from two original forms isnot very likely.only once was var.faba, the broad bean found, in Iraq. It dated from about 000 AD., which may point to late development of the large seed t Liliaceae FRITILLARIA IMPERIALIS L. Crown imperial.2n= 24, Iran and Turkestan. The bulbs were a source of starch and used in medicine. It is an ornamental. Linaceae LINUM USITATISSIMUM L. Flax, Linseed. 2n=30, (32). Origin of flax is discussed on p. 99. Primary centre of origin probably in Region 5 (Vavilov, 957). This conclusion is based on the great diversity of flax in India and adjacent northerly area. Spread from Region 5 into Region 4(p. 76). In the mountains of C. Asia, the 'curly oil flax' developed. It is characterized by the large number (40-50) of seed capsules per plant, Malvaceae GOSSYPIUM BARBADENSE L. Sea Islands cotton. 2n=52, genome formula (AADD)2< Peru, Spread to reach Africa and Asia in historical times. Secondary centre in Turkmenia, Tajikistan and S. Uzbekistan, USSR. GOSSYPIUM HERBACEUM L. Short staple American cotton. 2n=26, genome formula A^Ai. S. Africa (p. 39). Introduced into Ethiopia, S. Arabia and Baluchistan, where race acerifolium (p. 99) developed (Hutchinson, 962). Meliaceae MELIA AZEDARACH L. China berry, Bakayan, Pride of India, Persian lilac, Bead tree. 2n=28. SW. Asia up tow. China. Cultivated in tropics as an ornamental and shade tree. Seeds produce oil. They are also used as beads. Moraceae MORUS NIGRA L. Black mulberry. 2n=(89-06), 308. C. Asia. Cultivated athigher altitude inthe tropics (Purseglove, 968), and also throughout S. Eurasia for its fruits.the Black Persian mulberry is probably a variety (Purseglove, 968). URTICA CANNABINA L.2n= Cultivated for fibre. Oleaceae C. and N, Asia. JASMINUM OFFICINALE L. Common white jasmine. 2n=26. From Iran to Kashmir and China. Cultivated especially in S. France for its flowers, which contain essential oil used in perfumery, Polygonaceae RHEUM RHAPONTICUM L. Rhubarb. 2n=44. SE. USSR. Cultivated as vegetable. Probably one of the parents ofr. hybridum*. Related tor, palmatum* 0 Rosaceae AMYGDALUS BROWICZII Freitag. 2n=. Afghanistan. Related to A. communis*and A, korshinskyi Hand.-Mazz, 2n= AMYGDALUS BUCHARICA Korsh. (syn. Prunus bucharica (Korsh.) Fedtsch.). Bukhara almond. 2n=6. W. Tien Shan, Pamirs and Alai, and in Afghanistan. Itmay form a source of sweet pits, of high oil content and of agood kernel to shell ratio, AMYGDALUS COMMUNIS L. (syn. Prunus amygdalus Batsch.). Almond. 2n=6. W. Kopet-Dagh (Turkmenia), Afghanistan andw. Tien Shan (p. 00). Primary centres: W. Kopet-Dagh (Turkmenia) and W. Tien Shan. Extensively cultivated in S. Europe and California. In this centre and in Region 6 (p. 00) other Amygdalus species are found. In Georgia (USSR) A, georgica Desf. (2n=6)is found. In Kopet-Dagh and Badkhyz A, turcomanica Lincz. occurs. In Armenia A. nairica Fed. & Takhi.and A. urartu* with A. fenzliana are native. In this area and in Nakhichevan A. fenzliana* (syn. Prunus fenzliana Fritsch) (2n=6) grows. This species is very cold resistant and crosses freely with cultivated species. Further A. scoparia Spach. (syn. Prunus scoparia (Spach.) Schneid.) (2n= 6)is found in Kopet-Dagh and Iran. AMYGDALUS PERSICA L. Peach. 2n=6. Primary centre: China (p. 42). Secondary centre in Iran and C. Asia. AMYGDALUS PETUNNIKOWII Litvin. (syn. Prunus petunnikowii (Litvin.) Rehd.). Turkestan almond. 2n=6. W. Tien Shan,in Kazakhstan, Tadshikistan and Uzbekistan and partly in Kirgizistan, USSR. It is an ornamental. It has a high oil content. It is drought resistant. It easily crosses with A. communis*. AMYGDALUS SPINOSISSIMA Bge. (syn, Prunus

78 LEGUMINOSAE spinosissima (Bge.) Franch.). Thorny peach brush. 2n=6. C. Asia from Kopet-Dagh (Turkmenia) through Pamir-Alai to W 0 Tien Shan, in Iran, Afghanistan and Kurdestan. It is late flowering and has a high oil content. AMYGDALUS TANGUTICA Korsh. (syn. Prunus dehiscens Koehne). Tangut plum. 2n=. W. China. Cultivated in some parts of China for the kernels. AMYGDALUS ULMIFOLIA (Franch.) M. Pop. (syn. Prunus triloba Lindl.). 2n=6. C. Asia. AMYGDALUS VAVILOVII M. Pop. (syn. Prunus vavilovii M. Pop.). Vavilov almond. 2n=6. Kopet Dagh (Iran and Turkmenia), W. Tien Shan, the Pamirs and Alai. At one time it was believed that this species was a hybrid between A. spinosissima* and A. communis*. ARMENIACA DASYCARPA (Erhr,) Borkh. (syn. A. atropurpurea Lois., Prunus dasycarpa Ehrh.). Purple apricot, Black apricot. 2n=. Unknown wild. Cultivated in C. Asia, Transcaucasia and Iran. The fruits are very sour and may be used in marmalades. It might be used as a source of late flowering, and of cold resistance of the flower buds. Malus kirghizorum ARMENIACA VULGARIS L. Apricot. 2n=6. Primary centre in NE. China (p. 42). Secondary centre for the cultivated apricot E. Tien Shan.For the wild apricot, this primary gene centre was formerly continuous with the main part in China (p. 42). CRATAEGUS AZAROLUS L. (syn. C. aronia Bosc.). Azarolier. 2n=. S. Europe, Africa (p. 8) and the Orient. In Uzbekistan, a large-fruited type, var, turcomanica Popoff, is found. It is poor in vigour. FRAGARIA BUCHARICA Losinsk. Bukhara strawberry. 2n=. Region 5. MALUS KIRGHIZORUM Al. & Fed. 2n=. W. Tien Shan in the underbush ofwild walnut (Juglans regia L.). Primary centres are in the basins of the Pskem, Ugam, Kok-Su and other rivers. It is a polymorphous species. It is likely that it introgressed into the cultivated apple (Malus pumila*). MALUS PUMILA* MALUS SIEVERSII (Ledeb.) M. Roem. 2n= W. Tien Shan and in Ala Tau of N e Xingiang Uygur (Sinkiang Province) in the underbush of the wild walnut (Juglans regia L.). Var. hissarica (Kudr.) Ponomarenko is also described as P. hissarica Kudr, MALUS SYLVESTRIS (L.) Miller. 2n=34 u Much of Europe, in Transcaucasia and probably into W. Turkestan. In the Caucasus, this species is not always distinct from M. pumila*. It isvery Malus sieversii winter-hardy. Used as a root-stock. Where it grows together with M. pumila, hybrids occur and so M. sylvestris must have been involved in the distant origin of the cultivated apple. One subspecies, ssp, praecox (Malus praecox (Pall.) Borkh.), is early-maturing. Primary centre in C. Asia. Occasionally cultivated in N. Africa (Uphof, 968). PRUNUS subgen CERASUS Pers. In Region 5, there occurwild cherries with small fruits (sect. Microcerasus Webb.). PRUNUSCERASIFERA Ehrh, Cherry plum. Myrobalan. 2n=6, genome formula CC, (24, 32, 48). Primary centre is in Caucasia (p. 0). Secondary centre: the W. Tien Shan. It is characterized by high yield, early flowering, early ripening, wide adaptability and high acid content, PRUNUS FERGANICA Lincz. Ferghana plum. 2n= Ferghana ridge in Tien Shan and in the Pamirs and Alai (USSR). It is a true-breeding hybrid of a spontaneous cross Amygdalus communis

79 CENTRAL ASIAN REGION and Prunus cerasifera*. PYRUS BUCHARICA Litv. 2n=. W. Tien Shan and the mountains of Tajikistan It is thought to be a hybrid ofp. regelii* and P. korshinskyi* (Vavilov, 930a). PYRUS KORSHINSKYI Nak. & Kik. 2n=. Pamirs, Alai and W. Tien Shan. It is considered as one parent of P. bucharica*. PYRUS REGELII Rehd. 2n=. W. Tien Shan, Pamirs, Alai and Bukhara Uplands. It is very resistant to drought. PYRUS SOGDIANA S. Kudr. 2n= region of Uzbekistan.. Shakhrisyabz PYRUS VAVILOVII M. Pop.2n=. E. Ferghana. It is considered a hybrid ofp. communis* x P. korshinskyi* (Vavilov, 930a). ROSA MOSCHATA J. Hermann. 2n=4, (28). Himalayas and Iran. Cultivated as ornamentals.it is a parent of R. x bifera*. Salicaceae SALIX ALBA L. (syn.s. Aurea Salisb.). White willow. 2n=76. Europe (p. 6), Asia and N. Africa. Cultivated in the Kashmir for lopping as fodder (Heybroek, 963). Tamaricaceae TAMARIX GALLICA L. Tamarisk. 2n=24. W. Himalayas and NW. India.Cultivated for shelter and as an ornamental. 978b). Carrot was domesticated in Afghanistan (primary centre of diversity) and from there it spread over Europe, the Mediterranean area and Asia. During spread, it introgressed with local wild types. The domesticated types are divided into two groups : ()the 'Eastern (or Asian) carrots' (var. atrorubus Alef.), with mainly purple and yellow roots. (2)the 'Western carrots' (var, sativus Hoffm.) with mainly orange roots (Small, 978b). The purple types have a short storage time. InTurkey and Japan, hybrids between the two groups occur, in Turkey because the two groups grow near together and hybridize naturally,turkey is therefore a secondary centre of diversity (p. 0). In Japan, breeders developed varieties from artificial crosses of these two groups. During its spread, the yellow and white carrots probably originated by mutation (but see p. 62). The white mutants (albus) were used for fodder and did not participate in the development of the European carrot (Banga, 957). After reaching Iran,itprobably spread thence to China (Banga, 962) 0 Vitadaceae VITIS VINIFERA L. Common grape. 2n=38. Deep valleys of Tien Shan and adjacent areas. Primary centre for the cultivated grape lies in that Region (see further p. 02). Ulmaceae ULMUS VILLOSA Brandis ex Gramble. 2n= Himalayas. Cultivated in sacred places or for ornamental purposes. It is lopped for fodder (Heybroek, 963). ULMUS WALLICHIANA Planch.2n=. Large-leaved elm. Himalayas, Cultivated in Kashmir and lopped for fodder (Heybroek, 963), Umbelliferae CUMINUM CYMINUM* DAUCUS-CAROTA L. Carrot. 2n=8. Wild types occur in Europe,SW. and C. Asia and N. Africa. These wild types have been grouped into two aggregates : () ssp. agg,gingidium including former sspp. gummifer Hooker f,,commutatus (Paol.) Thell., hispanicus (Goüan) Thell., hispidus (Arcangeli) Heywood, gadecaei (Rouy & Camus) Heywood, drepanensis (Arcangeli) Heywood, and rupestris (Guss.) Heywood, (2) ssp. agg. carota with the former sspp. carota, maritimus (Lam.) Batt., major (Vis.) Arcangeli and maximus (Desf.) Ball. There are some intermediate types (Small,

80 6 Near Eastern Region The NearEasternRegionwas described byvavilov. Itincluded apartof Region5. Darlington (956)called the area thesw. Asian region. Zhukovskij (968) recognised tworegions 5and6, and in 970 he separated theseonhis map. Zohary (969)preferred tocombine these regions into one. Within theregion lies thefertilecrescent. Here agriculture probably evolved around 9000 BC. (Çambal & Braidwood, 970).Harlan (97)called the areaof thefertile Crescent, Al NearEast centre. Agriculture spread from there to Europe,the Mediterranean Region, Afghanistan, Indiaandpossibly Africa. Majorcrops include several fruit-tree species ofbrassica oleracea, Hordeum vulgare, Lens esculenta, Medicago spp., Secale spp., Triticumspp., Vicia faba andvitisvinifera. In Georgia (USSR), asecondary centre of diversity developed forglycine max,lupinus albus, Phaseolus vulgaris, Setaria italica andzea mays.maize enteredussrbyway ofgeorgia. In Turkey, Harlan (95)described microcentres foramygdalus spp., Cucumis melo, C. sativus, Cucurbita moschata, C. pepo, Lensesculentum, Lupinus spp., Malus spp., Medicago sativa, other annualmedicago spp., Onobrychis viceaefolia, Phaseolus vulgaris, Pistacea spp., Prunus spp., Pyrus spp., Trifolium spp., Vicia faba, Vitisvinifera andzea mays. Alliaceae ALLIUM AMPELOPRASUM L. Levant garlic, Perennial sweet leek. 2n=6, (24), 32, genome formula AAA'A", (40), 48, genome formula AAA'A'- A"A", AABBBB. Europe, Asia Minor, Caucasia to Iran and N. Africa. A type resembling ssp. iranicum P.W. is cultivated near Tehran (Tahbaz, 97). In Israel the diploid is rare, but the triploid is a vegetatively propagated triploid weed (Kollmann, 972). Var. babingtonii (Borrer) Syme (syn. A. babingtonii Borrer) of W 0 Ireland and SW. England and var. bulbiferum Syme (syn. var. bulbilliferum Lloyd)of the Channel Islands and W. France are possible relics of former cultivation (Tutin et al., 976). ALLIUM ASCALONICUM L. Shallot. 2n=6. N. Africa, E. Mediterranean area. Closely related to A. deserti-syriaci Feinbrun (2n= ) from Syria and Iraq (de Wilde-Duyfjes, 976). Also described as a variant of A. cepa* (Vosa, 976; and others). ALLIUM KURHAT Schweinfl.Salad leek, kurrat. 2n=32. Probably Arabia and Sinai (Uphof, 968). The geographic destribution is not known. Cultivated in the Nile area, Arabia and Palestine for its leaves (Uphof, 968) Itmight be

81 NEAR EASTERN REGION derived from A. ampeloprasum* (Kuckuck &Kobabe, 962). Because of its close relationship to A. ampeloprasum*, it is often included in that species as var. kurrat Schweinf. ex Krause or in A. porrum* (de Wilde-Duyfjes, 976). ALLIUM PORRUM L. Leek. 2n=32. Asia Minor. Probably gene centre for cultivated forms (Kuckuck, 962). Leek is a cultivated form derived from A. ampeloprasum*. If so,it is included as var. porrum (L.) Cav. in the latter species. ALLIUM SATIVUM L. Garlic. 2n=6, genome formula SS. Wild type in C. Asia (p. 8). Secondary centre in Region 6 (Kazakova, 97). Boraginaceae SYMPHYTUM ASPERUM Lepechin (syn.s. asperrimum Donn.). Prickly comfrey. 2n=40 o Caucasia to Armenia and N. Iran t Cultivated as forage crop. It isprobably one of the parents of S. x uplandicum Nyman, (2n=36), a forage crop. The other is S. officinale L., common comfrey (2n=26, c. 36, 40,c. 40, 48), which is native to Europe, W. Siberia and Asia Minor, Chenopodiaceae BETA COROLLIFLORA Zosimovich ex Buttler,2n= 36, genome formula CCCC and aneuploids of 36 0 Turkey, Georgia (USSR)and Azerbaijan (USSR and Iran). Self-incompatible and frost-resistant. BETA INTERMEDIA Bunge. 2n=36,genome formula possibly LLCC. Plants which are probably hybrids of B. lomatogona* and B, trigyna* have been described as B, intermedia.they occur where the two species grow together. It is a source of resistance to yellow mosaic disease, BETA LOMATOGONA Fisch. & Mey. 2n=8, genome formula LL, (22), 36. Asia Minor and E. Transcaucasia. It is a weed characterized by 'monogerm fruits'. Where the distribution of this species and of B. trigyna overlap, tetraploid Beta lomatogona and B. intermedia ( ), B. macrorrhiza ( )and B. trigyna ( )(Ulbrich, 934). B. lomatogona is foundo The hybrids are probably identical toplants described as B. intermedia*. BETA MACRORRHIZA Stev. 2n=8. (Sub)alpine zones of mountains in Iran, Turkish Armenia (Lake Van) and the Caucasus. A winter-hardy species containing sugar (8-2%) and white pulp. BETA TRIGYNA Wald. & Kit. 2n=54, genome formula LLCCCC. Around the Black Sea with outliers to the Caspian Sea, Iran, Ukraine and Hungary. CHENOPODIUM CAPITATUM (L.) Asch. 2n=6, 8. The Orient. Cultivated (Mansfeld, 959). Compositae CARTHAMUS TINCTORIUS L. Safflower. 2n=24, genome formula BB. Vavilov (95) and Kupzow (932)proposed three areas of origin for the cultivated safflower: in India, based on variation and ancient culture; in Afghanistan,i, based on variation and proximity of wild species; and in Ethiopia, because of the presence of wild safflower species. However Hanelt (96),and Ashri & Knowles (960) placed the centre of origin in the Near East because of the similarity of cultivated safflower to two closely related wild species: C. flavescens Spreng, (syn. C. persicus Willd. (2n= 24, genome formula BB), found in Turkey, Syria and Lebanon; and C, palaestinus Eig. (2n=24, genome formula BB), found in deserts of W. Iraq and Israel (Khidir & Knowles, 970). In that area, introgression may still occur between wild and cultivated safflower. The great variation in the Afghanistan safflower population must be caused by the meeting of the Middle Eastern and the Pakistani types (Knowles, 969) (p. 82). The wild safflower of Ethiopia cannot be the progenitor of safflower in the Near East because it has 32 pairs of chromosomes, whereas the cultivated species has 2 pairs (Knowles, 969). Safflower is or has been cultivated in many areas of the Old World and in North America. Many improved USA varieties derive mainly from Sudanese material. They are now cultivated in Egypt e Spain and some other countries too, where they may cross with local varieties and soproduce new genotypes or they may replace the local varieties so that gene material is lost. Cultivated mainly for its flowers, which were a source of pigment. It is now cultivated for its seeds which yield an edible oil, whose high polyunsaturation due to its high content of linoleic acid makes it very suitable for consumption. Imrie & Knowles (970) suggested that C, palaestinus is a wild species. The weedy species C, flavescens and C. oxyantha M. B. (2n= 24, genome formula BB) and the cultivated species C. tinctorius derive from that species. CHRYSANTHEMUM COCCINEUM Willd. (syn. C, roseum Adam). 2n=8. Wild in N. Iran, Caucasia and

82 ALLIACEAE -GRAMINEAE Armenia, Cultivated as a garden plant. The flowers contain the insecticide pyrethrin,but their toxity is less than C. cinerariaefolium*. TANACETUM PARTHENIUM* Cornaceae CORNUS MAS L. (syn. C. mascula Hort.). Cornelian cherry. 2n=8, 54. Caucasia and Asia Minor as an underbush of deciduous forests. Cultivated for its edible fruits and as an ornamental shrub. The fruits are also used to produce vin de Cornouille, an alcoholic beverage. Corylaceae CORYLUS AVELLANA L. European hazel. 2n=22, 28. Europe and Caucasus,Primary centre in the Caucasus. Cultivated widely for its nuts: hazel nut or cobnut. In this same centre there is a wealth of other Corylus species; C. maxima*, C. pontica C. Koch (2n=28), C. colchica Alb., C 0 iberica Wittm. & Kemular, C. imoretica Kemular, C D cervorum Petr, and C. colurna*, Trazels are hybrids ofco avellana and tree hazels (C, colurna*). They have a high kernel quality, are winter-hardy and vigorous. CORYLUS COLURNA L. Turkish hazel. 2n=28. Occasionally cultivated inturkey for its nuts. It is also used as a rootstock and as a source of resistance of diseases of C. avellana*. Hybrids of that species are called 'trazels'. CORYLUS MAXIMA Miller. Filbert, White filbert, Red filbert. 2n=22, 28. Caucasia, W. Asia and SE. Europe. Cultivated for its nuts. Cruciferae BRASSICA OLERACEA L.Wild and cultivated cabbages. 2n=8, genome formula CC. In Asia Minor, varieties belonging to convar. oleracea*, convar, capitata* and convar. acephala* are common. CAMELINA SATIVA (L.) Crantz. False flax.2n= 40. In SE. Europe and SW. Asia, thewild parental form occurs, probably C. microcarpa Andrz. (2n=40). It became a weed in cereal crops and flax. Later is was cultivated for its oily seeds,the cultigen being called C. sativa. An intermediate form isc. pilosa (DC) Zinger. Another weed of flax fields is C, alyssum (Miller) Thell. (2n=40). All these species have also been grouped in one species, C. sativa, being divided into subspecies microcarpa (Andrz.) Hegi, sativa, pilosa (DC.) Hegi and alyssum (Miller) Hegi. It has almost disappeared from cultivation, CRAMBE ABYSSINICA Höchst, ex. R UE. Vries.2n= 90. Distribution in the wild is obscure. Three introductions from Turkey into USA have been used to develop an oil-producing crop. CRAMBE CORDIFOLIA Steven (syn. C. tatarica Jacq.). Tatar sea-kale.2n=, Highlands of Asia Minor, India and Ethiopia. The perennial herb is cultivated for the young leaves. IRATIS TINCTORIA L. (syn. I. canescens DC, I. littoralis Steven, I. taurica Bieb.). 2n^28, Most of Europe. Cultivated as source of dye, and therefore probably introduced. Cucurbitaceae CUCUMIS MELO L. Melon. Musk melon, Canteloupe. 2n=24. Africa (p. 24). Secondary centre arose in Region 4, in which convar. cassaba (Pang.) Greb. (cassaba melon,winter melon) from Asia Minor, convar. cantalupa (Pang.) Greb, (cantaloupe melons), convar. adana (Pang.) Greb., (kilik melons) convar. flexuosus (L.) Greb. (tarra melons,adjur melons,snake melons, serpent melons) are found. CUCUMIS SATIVUS L. Cucumber, Gherkin. 2n=4, India (p. 72). Secondary centre (bethalpha type) arose in this region. Dipsacaceae CEPHALARIA SYRIACA (L.) Roemer & Schul tes. Pelemir. 2n=0. This pestweed of wheat fields is occasionally cultivated on the C. Anatolian peneplane as anoil crop. DIPSACUS SATIVUS (L.) Scholier. Teasel. 2n= 6, 8, Cultivated in Europe and elsewhere. It has probably derived from D. ferox Loissel (2n=6, 8), which grows in Corsica, Sardinia and some sites in C. Italy or from D, fullosum L, (syn.d. sylvestris Hudson, 2n=6, 8), which grows in S, ( W, and C. Europe to NE. Ukraine. Fagaceae CASTANEA SATIVA Mill. (syn. C. vesca Gaertn u). Sweet chestnut,spanish chestnut. 2n=22, 24. From Italy northwards tohungary and eastwards to Asia Minor and W. Georgia (USSR). Cultivated for its nuts and timber. Outside that range, it is naturalized. Gramineae AEGILOPS CAUDATA L. (syn. Triticum dichasians (Zhuk.) Bowden, T, caudatum (L.) Godr. & Gren.). 2n=4, genome formula CC. Greece, Turkey, Iraq andafghanistan,, Its cytoplasm has a malesterilizing action oi the T. aestivum* nucleus. AEGILOPS COLUMNARIS Zhuk. 2n=28, genome formula C U C U M C M C. Turkey, Iraq, Iran and Caucasia. It is a weed of cultivation and looks quite like Ae. triaristata* (Bor, 970). The C u genome will be renamed U.

83 NEAR EASTERN REGION Primary centre ( ), secondary centre ( )and distribution of wild and weedy Secale cereale types and ways of their introduction and that of rye intoeurope and N. Asia (...) (Khush, 962). 2n=4, genome formula R R. Primary centre of annual and perennial rye species and forms: NE. Turkey - NW. Iran.Khush (963) suggested that a secondary centre (p. 83) is in Afghanistan and that the cultivated rye of Europe and N. and C. Asia derives from that centre. Only afew come from the primary centre.in the primary centre, S. vavilovii* and S, montaneum* hybridize with each other and introgress, resulting in a mixture of genetic variants described as 'S. segetale', 'S. afghanicum', 'S. daralgesii', 'S. cereale', 'S. turkestanicum' and 'S. dighoricum' (Stutz, 972). Stutz (972) suggested that from this highly variable population the annual S. cereale types invaded cultivated fields to become weeds. S. cereale includes cultivated rye and a number of weedy rye types occurring in grain fields and along ditchbanks and roadsides, throughout the Middle East (Stutz, 972). These weedy types are: S. afghanicum* S. dighoricum (Vav.) Roshev 0 (syn.s. cereale L. ssp. dighoricum (Vav.) Khush). It is a weed in grain fields of N. Ossetia (USSR). S. segetale (Zhuk.) Roshev. (syn.s. cereale L. ssp.segetale (Zhuk.) Khush.). This is a polymorphic weed in grain fields throughout E. Europe and the Middle East (Stutz, 972). S. ancestrale Zhuk. (syn<, S 0 cereale L, ssp. ancestrale (Zhuk.) Khush). It is a robust tall (up to2.4 m)weed with small invested seeds and fragile rachis restricted to sandy ditchbanks and fence rows near Aydin, SW. Turkey. It is reproductively isolated from domesticated rye (Stutz, 976). S. turkestanicum Bensin. À self-compatible cultigen of C. Asia (p. 83) and Transcaucasia. Theweedy types have derived mainly from S. vavilovii* by introgression with S. montanum* and its derivative species S. anatolicum*. In some places less favourable for wheat and barley, rye may have been developed to become fully domesticated. It is generally proposed that S. ancestrale and S. segetale are the parental types of S. cereale; however Stutz (972) suggested that S. ancestrale derives from S. cereale. There isno introgression into S. ancestraleof other Secale genetic material, in spite of contact with other species, because a high incidence of geno-typically controlled chromosomal breaks inoutcrossed hybrids leads to sterility (Stutz, 97). Hybridization between wheat (p. 93)and rye may have increased in the variability of wheat. Rye is a source of resistance to diseases used in wheat breeding and is a parent of octaploid and hexaploid triticales. The closest wild relative of cultivated rye is S. ancestrale and this taxon probably gave rise to the weedy S. segetale complex. Wild rye

84 GRAMINEAE -GRAMINEAE 93 is characterized by spikelets that all disarticulate at maturity, while in theweed complex the lower one quarter or more of the spikelets arenot deciduous at maturity. Cultivated rye is known from the Neolithic of Austria, but it became widespread as a cereal in Europe only since the Bronze age. SECALE MONTANUM Guss. Mountain rye. 2n=4, genome formula R m R ra.from the C. Atlas Mountains of Morocco and the Sierra Nevada Mountains of Spain, eastwards in isolated pockets in the mountains of Sicily, Italy, Yugoslavia, Greece, Lebanon, Turkey, Iran and Iraq (Stutz, 972). It is ahighly polymorphic cross-fertilizing perennial. Some of the isolates have been described as distinct species or varieties: S. ciliatoglume (Boiss.) Grossh. (syn.s. montanum var. ciliatoglume Boiss.). A weedy population with pubescent culms in orchards and vineyards near Mardin, SE. Turkey. S. dalmaticum Vis., population growing within the walls of the old St. Johannis Fortress above Kotor,S. Jugoslavia. S. daralgesii Thum., a weedy form with nonfragile rachis along roadsides and ditchbanks of Armenia, S. kuprijanovii Grossh., a broad-leaved form of mountain meadows of the N. Caucasus Mountains. S. montanum has the same chromosomal arrangement as S. anatolicum*, S. silvestre* and S. africanum*. Its chromosomal arrangement differs from that of S,vavilovii and S. cereale and from its closely related forms in reciprocal translocations involving 6 of the 4 chromosomes. It derives from S. silvestre* (Singh & Röbbelen, 975) and is the parental species of S. anatolicum (Stutz, 972), S. africanum, S. vavilovii and S, cereale. SECALE SILVESTRE Host. (syn.s. fragile Marsch.), 2n=4. C. Hungary eastward throughout the sandy steppe of S. Russia up to W. Siberia and the Pamirs and Alai (Bor, 970; Stutz, 972). A low-growing annual psammophyte with fragile rachis. It has the same chromosome arrangement as S, montanum*. This species is believed tobe phylogenetically the oldest species and isprobably the ancestor of S. montanum* (Rimpau & Flavell, 974; Singh & RÖbbelen, 975) from which S. cereale* derives. SECALE VAVILOVII Grossh. 2n=4. Common to the lower slopes of Mount Ararat and along the banks of the Araks River. It is a wild low-growing, annual self-compatible psammophyte with fragile rachis. It has the same chromosome arrangement as S. cereale (see S. montanum*). Bor (970), strongly suspected this species of being the same as S. afghanicum*. Khush (960) suggested that it derived from S. montanum, but Stutz (972) made it clear that is is the ancestor of S. cereale*. TRITICUM AESTIVUM (L.) Thell. Wheat. 2n=42, genome formula AABBDD. Primary centre: Transcaucasia and adjacent areas. There, natural cross-fertilization is still taking place within the species and between subspecies, other Triticum species and related species of Aegilops and Secale. This species is anatural amphiploid of emmer and Aegilops squarrosa*. This amphiploidization must have taken place after the development of emmer from itswild ancestor T. turgidum ssp. dicoccoides* in the area south of the Caspian Sea (Nakai, 979). This hybridization has led to hexaploid types with avery brittle ear. This is a 'wild' character and soone could say that there are wild hexaploid wheats. However these segregants cannot really be called wild. The main division of T. aestivum is into ssp. spelta (L.) Thell. (syn. T. spelta L Q), spelt; ssp. vavilovi (Turn.) Sears (syn.t. vavilovii (Tum.) Jakubz.); ssp.macha (Dek. & Men.) MK. (syn.t. macha Dek. & Men.). Makha wheat, ssp. vulgare (Vill.) MK. (syn. T. vulgare Vill.), common wheat, bread wheat; ssp. compactum Host.)MK. (syn. T. compactum Host.), club wheat; and ssp.sphaerococcum (Perc.) MK. (syn. T. sphaerococcum Perc), Indian dwarf wheat. The origin of some subspecies has not yet been assertained. They have originated in another centre. Spelt wheats have been found in Iran, in C, Europe (p. 54)and Africa (p. 35). The spread of bread wheat during Neolithic times has been described (Zeven, 979; 980). Ssp. vavilovi wheat is indigenous to Armenia. It is characterized by its branching spikelet. Makha wheat is indigenous to W. Georgia (US SR). It is often mixed with T. turgidum ssp. paleocolchicum*. The spread of bread wheat during Neolithic times has been described (Zeven, 979; 980). Bread wheat isnow widespread. Primary centre in Transcaucasia and adjacent regions. Secondary centres in Hindu Kush and adjacent regions (p. 83), in China and Japan (p. 39)and probably in African Sahara (p. 35). Club wheat developed in Afghanistan and adjacent regions (p. 83) and probably in Switzerland/Austria (p. 54). A secondary centre of diversity is Armenia. Indian dwarf wheat originated in NW. India and adjacent regions (p. 75). Its presence has been reported in N. Africa. Dorofejev (97) suggested that ssp. macha is the oldest hexaploid. From it, ssp. vulgare developed. Ssp. spelta and ssp. vavilovii are secondary spelts. They may have derived from ssp. vulgare. There has been much research to identify the donor species of the B genome. However in vain, because itprobably does not exist (seet. turgidum*). TRITICUM AESTIVUM (L.) Thell. ssp. compactum (Host) MK. (syn. T. compactum Host.). Club wheat. 2n=42, genome formula AABBDD. This subspecies developed in the Hindu Kush (p. 83).

85 NEAR EASTERN REGION Secondary centre Armenia, TRITICUM MONOCOCCUM L. Wild and cultivated einkorn. 2n=4, genome formula AA.The wild ssp.boeoticum (Boiss.) Mac Key (syn. T. boeoticum Boiss.) includes the types aegilopoides (T. aegilopoides (Link) Bal.and thaoudar (T. thaoudar Reut.), as well as 'T. spontaneum Flaksb.' and 'T. urartu Tuman.'. It is spread over Greece, Turkey, Syria, N. Iraq and Transcaucasia. Aegilopoides is characterized by one grain and one awn per spikelet, whereas thaoudar has two grains and two awns. There are two distribution centres: in the Fertile Crescent and in Turkey. In peripheral areas, it is segetal. In the Fertile Crescent thaoudar is found, Aegilopoides type occurs in the colder part of Balkans and W. Anatolia. In Anatolia, intermediate types and mixtures are found. In Armenia the type urartu has been described. It has two awns and a winter habit. In the Fertile Crescent, thewild einkorn was domesticated to become ssp. monococcura, which has a tough rachis. Its earliest appearance is from Ali Kosh dating from c BC., thus later than the first appearance to T. turgidum ssp. dicoccum. Einkorn was spread over Europe, N. Africa, Asia Minor, Caucasia, Iraq and Iran. Cultivated in some areas as a fodder crop. It forms a component of the Zanduri wheat, which is a mixture of einkorn. T, timopheevi ssp. timopheevi* and T. zhukovskyi*. This population is cultivated in Georgia (USSR). This species forms a useful source of disease resistance. There is still a natural gene flow between diploid and tetraploid species (Vardi & Zohary, 967) 0 TRITICUM TIMOPHEEVI Zhuk. 2n=28, genome.formula AABB (or AAB'B', AAGG). This species consists of two subspecies ssp. araraticum (Jakubz.) Mac Key (syn. T. araraticum Jakubz.) and ssp. timopheevi. Ssp. araraticum grows wild in Transcaucasia. N. Iraq, W. Iran and E. Turkey. It was first described as T. dicoccoides ssp. armeniacum Jakubz.and as T. armeniacum Mak. Some types closely resemble T 0 dicoccoides. However the subspecies only crosses easily with ssp, timopheevi. Ssp. timopheevi is part of the Zanduri wheat cultivated in Georgia (USSR). It is an ancient cultivated wheat 0 Its rather brittle rachis causes difficulties in threshing. Itis difficult to cross with other Triticum species ; It is a source of disease resistance, (Timopheevi)durum and aestivum plants often are male-sterile. Maan (973)concluded from his nucleo-cytoplasmic and cytogenic studies that T a timopheevi ssp.timopheevi and ssp.araraticum and T. dicoccoides var. nudiglumis ex Turkey-Iran -Iraq area have the same type of cytoplasm and the genome formula AAGG. The cytoplasm of Ae. speltoides* is closer to the above cytoplasm than to T D turgidum*-cultivated wheats. Wild tetraploid Triticum species: Triticum turgidum ssp. dicoccoides (I,II,III,), ssp. dicoccoides var. nudiglunis (III) and T. timopheevi ssp. araraticum (IV) (Johnson, 972). TRITICUM TURGIDUM (L.) Thell.Wild emmer wheats. 2n=28, genome formula AABB. The distribution of the wild ssp.dicoccoides can be divided into two regions. The N 0 region is S. Turkey - Iran - Iraq, where var. nudiglumis is found; the S region is Israel,S. Syria and Jordan. The cytoplasm of var. nudiglumis is similar to that of T. timopheevi*, whereas that of ssp. dicoccoides of the southern region is the same as the cultivated T. turgidum* (Maan, 973). Ssp. dicoccoides (Köm.) Thell. (syn. T. dicoccoides Körn.), derived from a natural amphidiploidization of an unknown diploid species and T. monococcum ssp. boeoticum. This amphidiploidization must probably have occurred in Syria and Palestine,probably at several places. There has been much research to identify the unknown diploid parent. However in vain. Triticum boeoticum (Harlan & Zohary, 969). The research has failed because the B genome of this species and of its donor have evolved of the first tetraploid Triticum species - some years ago. Evolution followed different paths, which branched to give rise to several species with related but different B genomes. From ssp. dicoccoides, several cultivated subspecies have been derived.these are dicoccum*, palaeocolchicum*, turgidum* and carthlicum*. The ssp. turgidum includes conv. turgidum, conv. durum, conv. turanicum and conv. polonicum. Ssp. dicoccoides is a source

86 GRAMINEAE - LABIATAE of disease resistance; it also carries a restorer gene of (timopheevi) cytoplasmic malesterility. TRITICUM TURGIDUM (L.) Thell. Cultivated emmer wheats. 2n=28, genome formula AABB.The cultivated tetraploid wheat can be subdivided as follows (MacKey, 966): ssp. dicoccum (Schrank) Thell. (syn. T. dicoccum Schubl.)., emmer, ssp. palaeocolchicum (Men.) Mac Key (syn.t. palaeocolchicum Men., T, georgicum Dek.), Georgian emmer, Kolchic emmer, ssp. turgidum including conv. turgidum, Poulard wheat, English wheat, conv. durum (Desf.) Mac Key (syn. T. durum Desf.), durum wheat, hard wheat, conv. turanicum (Jakubz.) Mac Key (T. turanicum Jakubz., T. orientale Perc), Khurasan wheat, conv. polonicum (L.) Mac Key (syn. T. polonicum L.) Polish wheat, ssp. carthlicum (Nevski) Mac Key (syn. T. carthlicum Nevski, T. persicum Vav. ex Zhuk.), Persian wheat. They have originated by cultivation from ssp. didoccoides* and perhaps by intergeneric and interspecific hybridization. Primary centre in Near Eastern Region«Secondary centres in Ethiopia (p. 35)and in the Mediterranean Region (p. 2). Emmer is the oldest cultivated wheat. Its cultivation is declining. Until recently, it was cultivated in Ethiopia (p. 35), Iran, E. Turkey, Transcaucasia, Volga Basin, Yugoslavia, Czechoslovakia and India. Itmay have been domesticated c.8000 BC.in the Fertile Crescent. The first appearance of domesticated emmer dates from c.7000 BC.atAceramic Neolithic Beidha, Ali Kosh, Jericho and Ramad. Already in BC., ithad reached the Atlantic coast from Scandinavia to Spain and the Nile Delta. Helbaek (960) was surprised by the uniformity of emmer. However plants are called dicoccum when they correspond to a certain morphological description. It is unknown whether the idiotypes of the various dicoccum populations also correspond to each other for characteristics other than morphological ones. The disease-resistant khapli (khapli is the vernacular name of emmer) of India and Yaroslav emmer from USSR belong to ssp. dicocum. Durum wheat is cultivated over a large area: Mediterranean coastal region, Ethiopia where a secondary centre exists (p. 35) and in areas north of the Black Sea. Another centre of diversity is in India (Jain et al., 976). It is rarely observed in wheats of Iran and Afghanistan. It is also cultivated in the Americas and elsewhere. Distribution in the Old World is shown by Ciferri (939). It is the second most important wheat in the world. Turanicum wheat originally involved as an oasis ecotype. Its cultivation is restricted to irrigated fields (Mac Key, 966). Mac Key suggested a wide occurrence in Asia, but Kuckuck (970) and Bor (970) limited it to Iran and Iraq, Kuckuck (970)suggested that it is a hybrid of durum x polonicum. Spread of durum wheat in the Old World (Ciferri, 939). Polonicum wheat,distinguished by long glumes and kernels was cultivated in S. Europe, Turkey, Iraq, Iran, Afghanistan and NW. India. According to Kihara et al. (956), in includes T. ispahanicum Heslot. Carthlicum wheat is characterized by the presence of the Q (vulgare) gene. It is not known whether this gene arose independently in this wheat or came from vulgare wheat. Carthlicum wheat was cultivated in Iraq, Iran and Caucasia. It is a source of disease resistance. Georgian emmer, Kolchic emmer (ssp. palaeocolchicum) was formerly cultivated in a mixture with T. aestivum ssp. macha* in W. Georgia (USSR). English wheat (conv. turgidum) was atone time cultivated in Europe and elsewhere. From time to time,it was reintroduced into cultivation because its branching habit (Osiris wheat, Wonder wheat) convinced farmers that its yield must be high. TRITICUM ZHUKOVSKYI Men. & Er. Zanduri,2n= 42, genome formula AAAABB or A z A z A t A t ß' E B t. It was cultivated in W. Georgia, USSR. Zanduri was composed of einkorn, T. timopheevi ssp. timopheevi* and T. zhukovskyi. The latter is a hexaploid but its genome formula differs from those of the common hexaploid subspecies.it is a natural amphiploid of ssp. timopheevi and einkorn. Its cytoplasm has the same malesterilizing action on the durum and aestivum nuclei as ssp. timopheevi. It carries genes for resistance to stem rust and mildew. ZEA MAYS L. Maize. 2n=20. Secondary centre in the Near East (Brandolini, 970). Domesticated in C. America (p. 90). Flint maize - indurata Sturt. is common in the Near East. Iridaceae CROCUS SATIVUS* Labiatae LALLEMANTIA IBERICA Fisch. & Mey. Lallemantia. 2n=6. Asia Minor and some regions of USSR.

87 96 NEAR EASTERN REGION Cultivated in Iran ands. USSR forits oil seeds. Leguminosae ALOPHOTROPIS FORMOSUM (Boiss.) Lamprecht (syn. Pisum formosura (Stev.) Boiss., Vavilovia formosa (Stev.) Fed.). Wild perennial pea. 2n= 4. Montane and submontane zones of Asia Minor, Transcaucasia, Armenia and Iran, CICER ARIETINUM L. Chick-pea, Gram, Garbanzos. 2n=4, 6, (24, 32, 33). Unknown wild. Secondary centres of diversity probably developed in Regions 4 (p. 76), 5 (p. 83) and 7 (p. 3). Cultivated in S. Europe and N. Africa from the Atlantic eastwards,the Nile Delta and Ethiopia, and northwards and eastwards to NW, Burma, W. China, Kazakhstan (USSR). Some ofthe Cicer species indigenous to Anatolia may have played a role in itsancestry, particularly C. pinnatifidum Jaub. & Spach., 2n=6, (from Anatolia, Soviet Armenia, Syria, N. Iraq and Cyprus), C. echinospermum P^H. Davis,2n=,(from E. Anatolia)and C. bijugum K.H. Rech,, 2n=6 (from SE. Anatolia, N. Syria,and N. Iraq) (van der Maesen, 972). Ladizinsky (975a) found a wild annual species in Turkey: C. reticulatum Lad., 2n=, which is cross-compatible with chick-pea, producing a fertile Fi» This species resembles chick-pea and could bethe wild parent ofthe cultigen. So Moreno & Cubero (978)proposed to include it as ssp. reticulatum (Lad.) Moreno & Cubero in C. arietinum. Race orientale Pop. is characterized by very small seeds (000-seed weight g). It is common in Ethiopia, Sudan, Egypt, India, the Pamirs, Tajikistan and Iran. Those from Ethiopia are black-seeded 0 Race asiaticum Pop. has somewhat bigger, but still small, seeds (000-seed weight g). It occurs in C. Asia, Afghanistan, W. China, Iran and E. Turkey. Race eurasiaticum Pop.has moderately large seeds (000-seed weight g). Itis cultivated inthe Near East, Armenia, Azerbaijan, Ukraine upto C.USSR, and near large cities in the eastern part ofthearea of cultivation.theseeds are white. Race mediterraneum Pop. hasthelargest seeds (000-seed weight over 350 g) 0 Itis found in Spain, Italy, Morocco, Algeria, Tunisia and W. Turkey, The seeds are white. However Moreno & Cubero (978)grouped the cultivars into two races:the small-seeded inrace microsperma, andthe large-seeded in race macrosperma.the latter isselected from the first, GALEGA ORIENTALIS Lam. 2n=6. Caucasia. Cultivated as afodder and asan ornamental 0 LENS ESCULENTA Moench (syn. L. culinaris Medik., Ervum lens L.). Lentil. 2n=4.Zohary (972) suggested that L. orientalis (Boiss.) Hand.- Mazz., 2n=, isthe wild ancestor of lentil; it grows wild in Region6, L. orientalis is a dwarf lentil,as was confirmed by Williamset al. (974). Ladizinsky (979) toofound aclose affinity between the two species and L. nigricans (M.B.) Godr., 2n=. So Williams et al. (974) concluded that lentil andits wild ancestor belonged to one species L. esculenta, both types being ssp. culinaris (Medik.) Williams, Sanchez & Jackson for lentil,and ssp. orientalis (Boiss.) Williams, Sanchez & Jackson for wild lentil. Lentil had been divided by seed size into varieties : var. macrosperma, a large-seeded form from the Mediterranean Region, var, syrica Barul., a medium-seeded form from the inner mountainous regionof Asia Minor, andvar» afghanica Barul,, a small-seeded form of the highlands of Afghanistan, Another divisionis ssp. macrosperma (Baumg.) Barul.and ssp. microsperma (Baumg.) Barul, However Williamset al. (974)concluded that such divisionis meaningless as macrosperma and microsperma form the extremes of acline forseed size. Microsperma isfound inall prehistoric exca- "a * r- t»> f / / K t i*k u^/ - "K. - ^?T «/ ".~'"" L -r~^ îv*^ y J ) ^ _/ _, > \è, \ l #; ). '"'A '^îv < ^/ \ ' i 0 X \>»' a [- '' \r oo o >r\ \ -7sö ö i \,.J «r?v " - - *\ < ^ Lens nigricans ( ), L. orientalis (o) and early records 6-7th millenia BCof lentil (*)(Zohary, 976; Ladizinsky, 979). \ oo ( oo J ^ < o > o o 0 o o ' - oo3\,. '\ r' " <T- )- '? "i j \ [ 0 V 'S ' J " J _A_. c'

88 LABIATAE - LEGUMINOSAE vations (van Zeist & Bottema, 97). Agro-ecological groups meet in Turkey, where there are microcentres of diversity (Harlan, 95). MEDICAGO CANCELLATA Bieb. 2n=48. SE. European USSR and N. Caucasia. Useful as gene source for adaptation of M. sativa* to poor soils. MEDICAGO DZHAWAKHETICA Bordz. 2n=6, genome formula DD, 32. The (sub)alpine zones of the Alkhalak uplands, Georgia, USSR and a part of Asia Minor. A wild perennial species. Var. timofeevi Troitz., (2n=32) is endemic to Transcaucasia. It crosses fairly easy with M. sativa (Lesins and Lesins, 966) 0 Some include this species as var. dzhawakhetica Bordz. in M. papulosa Boiss. (2n=6). MEDICAGO GLUTINOSA Bieb. 2n=32. Causasus. Useful as gene source for M. sativa*. MEDICAGO ROMANICA Prod. 2n=6. Caucasia. A wild variable perennial species 0 MEDICAGO SATIVA L. Lucerne, Alfalfa. 2n=6, genome formula SS; 32, genome formula SSSS; (48). The species probably evolved around the Caspian Sea, whence it spread as a wild plant. It was the first species to be cultivated as a forage crop,probably in association with the increasing use of horses and the development of light horse-drawn chariots in the first part of the 2nd Millenium BC. It was probably intentionally sown and the more vigorous tetraploid crowded out the diploido During the Medicago sativa (Fischer, 938). Persian-Greek War in the 5th Century BC., lucerne (from Provencal, meaning shining) and its older name alfalfa (from Arabic and ultimately Old Iranian aspo-asti, horse fodder) was introduced into Greece, whence it spread to S. Italy and Europe, and later to other parts of the world (Lesins & Lesins, 979). MEDICAGO SAXATILIS Bieb. 2n=6. Yaila Range of Mountains in Crimea. Useful as gene source for M. sativa*. ONOBRYCHIS ALTISSIMA Grossh. casia. 2n~4. Transcau- ONOBRYCHIS VICIIFOLIA Scop. Esparcette. 2n=28. Cultivation of this fodder crop started ins. France (p. 57). In Transcaucasia, var. transcaucasia (syn. 0. transcaucasia Grossh.) is endemic. PISUM SATIVUM L. (syn. P. arvense L.s..). Pea. 2n=4, This species may be divided into six subspecies: ssp. abyssinicum (p. 37), ssp. jomardi (p. 5), ssp. syriacum Berger, ssp. elatius (Stev.) Alef., ssp. arvense Poir., and ssp. hortense Asch, et Graeb. (Gentry, 97). Ssp. syriacum (syn.p. humile Boiss.et Noë, P. sativum var Q humile,p. syriacum Berger) is found in N, Iraq, Jordan, Syria, N,, NW. and W. Iran, Israel, Turkey and Cyprus. Some forms are robust (30-70 cm tall), others slender and small (20-40 cm) (Ben-Ze'ev & Zohary, 973). This iszohary's (973) 'steppe' type. Itis the primary wild progenitor. Ssp. elatius (syn.p. elatius Stev.)is found in Syria, N. Israel, Lebanon, S. coast of Turkey, Aegean belt of Turkey and Greece, Cyprus, Adriatic coast of Yugoslavia, S, Italy, Sicily, Sardinia and scattered localities in Morocco, Algeria, Tunisia,S. Spain, S. France, N. Italy and the Black Sea coast of Turkey, Crimea and Caucasia (BenZe'ev & Zohary, 973). This is Zohary's (973) tall 'maquis' type. In the E. Mediterranean countries (esp.s. Turkey), intermediate types are found. Ben Ze'ev & Zohary (973) suggested that ecotypes of Turkey and Syria may have formed the parental material of the domesticated types ssp. arvense (P.arvense L,), field pea,and ssp. hortense (ssp. sativum, P. sativum L 0), garden pea. Itmay also derive from ssp,, elatius,or from hybrids of ssp, elatius x ssp. arvense. Secondary centre in the Mediterranean Region (p. 5). Itmay have been domesticated in SW. Asia. The crop reached the Greeks by way of the Black Sea, who passed it on to Latin and Germanic tribes. It spread to India and China through the Himalayas and Tibet, and to Ethiopia and E. Africa (Purseglove, 968). TRIFOLIUM AMBIGUUM M.B. Caucasian clover.2n= 2x=6, (32; allo-6x=48). Caucasia, Crimea and Turkey. This valuable fodder plant forms an essential part of pastures and meadows.in Australia, an allohexaploid cultivar derives from an introduction from USSR. It withstands periods of 6 weeks submerged in water. TRIGONELLA FOENUM-GRAECUM L. Fenugreek. 2n=6, (4x=32). Probably SW. Asia. Cultivated in S u Europe, N. Africa and India as a fodder. The seeds are also eaten in India and used in medicine.

89 NEAR EASTERN REGION Pisum humile ( ), P. elatius ( ) and Alophotropis forraosum ( )(Govorov, 937). VICIA ERVILEA (L.) Willd. Bitter vetch, Ervil. 2n=4. Primary centre in the Mediterranean Region (p. 6). In Asia Minor, a characteristic group developed. It is now cultivated as a fodder but in prehistoric times it was cultivated for food. It was already cultivated in Turkey in 5750 BC 0 and probably in Greece about 5500 BC. (van Zeist & Bottema, 97). VICIA NARBONENSIS L. Narbonne vetch. 2n=4. Primary gene centre probably E. Georgia. Secondary gene centre in the Mediterranean Region. This species is a weed in wheat and barley fields of Transcaucasia and other areas in SE. Asia. It isnot cultivated there. VICIA PANNONICA Crantz. Hungarian vetch. 2n= 2. Primary gene centre in Georgia (USSR), on the plateau of Akhalkalak, where it grows wild and is cultivated. Secondary gene centre in Hungary. VICIA SATIVA L. Common vetch. 2n=0, 2, 4. The V. sativa aggregate is complex because of variation in form, in chromosome number and in karyotype,and of irregular cytogenetic affinities between the types with different karyotypes (Ladizinsky & Temkin, 978). The 3 cytotypes are found among the wild, weedy and domesticated types (Zohary & Plitmann, 979). In the wild types, they are partially isolated by ecological barriers, partial hybrid sterility and predominance of self-fertilization,, Human disturbance of the land has removed these barriers and increased variation, including that of the karyotypes. Zohary & Plitmann (979)divided the aggregate up as follows:. ssp. sativa, 2n=2, 7 karyotypes found. From Atlantic fringe of Europe through Mediterranean to W. India, weedy, escapes and cultivars (seed and hay), run wild in New World 2. ssp. macrocarpa (Moris) Arcang,, 2n=2, 2 karyotypes. W, Mediterranean, cultivars (seed) 3a. ssp. nigra (L.) Ehrh. var. nigra (syn. V. sativa L. var. angustifolia L.), 2n=2, more than 25 karyotypes. Mediterranean Basin and Europe, runwild in the New World, cultivated for hay. 3b. ssp. nigra (L.) Ehrh. var. segetalis (Thuill.) Ser. in DC (syn. V. segetalis Thuill. mostly 2n=2, more than 80 karyotypes. Mediterranean Basin deep into C. Europe, very weedy, recently cultivated for hay. 4. ssp. cordata (Wulfen ex Hoppe) Aschers. & Graebn. (syn. V. cordata Wulfen ex Hoppe), 2n= 0, more than 30 karyotypes. Mediterranean Region, very weedy, some domesticated types (hay) 5. ssp.incisa (M.B.) Arcang. (syn. V. incisa M.B.), 2n=4, karyotypic variation unknown. Near East 6. ssp. amphicarpa (Dorth.) Aschers. & Graebn. (syn. V. amphicarpa Porth.), 2n=4, karyotypic variation unknown. Mediterranean Area and Near East. This type is unique for its subterranean pods and is according toladizinsky (978)an advanced form 7. ssp. pilosa (M.B.) Plitm. & D.Zoh. (syn. V. pilosa M.B.), 2n=4, karyotypic variation unknown. Crimea and Caucasia. By further hybridization of cytotypes and karyotypes, and by natural and human selection, the variation of this species may still increase considerably. VICIA VILLOSA Roth. Sand vetch, Hairy vetch, Winter vetch. 2n=4. W. and C. Europe, Mediterranean Region, N. Iraq, N, Iran and SW.of USSR. Primary centre probably W. Asia and Ante-Asia. Spread to the Mediterranean area and Europe as a cereal weed. Liliaceae HYACINTHUS ORIENTALIS L. Common hyacinth. 2n=6, (24, 32). Syria, Asia Minor, Greece and Dalmatia. Cultivated in the Netherlands as an ornamental and in S. France as a source of an essential oil used in perfumery.

90 LEGUMINOSAE - NELUMBONACEAE 99 coastland race, a perennial,also described asl. angustifolium Huds. (2n=30). Ithas the highest seed oil content and the highest seed weight of all wild species (Seetharam, 972). During domestication and furter development, types for fibre (flax) and oil (linseed) developed. Malvaceae ALTHAEA OFFICINALIS* ALTHAEA ROSAE* GOSSYPIUM AREYSIANUM Deflers. 2n=26, genome formula E3E3. S. Arabia. It is drought-resistant and early maturing. GOSSYPIUM HERBACEUM L. Short-staple cotton. 2n=26, genome formula A-JA-J^ S. Africa (p. 39). Introduced to Ethiopia, S. Arabia and Baluchistan, where race acerifolium* developed. In Iran, a characteristic group of annual forms has arisen,named race persicum. It spread to W. India, where it was the first annual cotton cultivated. Varieties of G. herbaceum are now often cultivated. In C. Asia, race kuljianum developed. It matures in three months from sowing, giving asmall crop. GOSSYPIUM INCANUM (Schwartz) Hillcoat. 2n=26, genome formula E4E4. S. Yemen. It is droughtresistant. Liniun usitatissimum, various length and branching types Linaceae LINUM USITATISSIMUM L. Flax, Linseed. 2n=30, (32). Primary centre probably in Central Asian Region (Vavilov, 957), since flax varies widely in India and adjacent northerly areas. However as the progenitor of flax, L. bienne Mill. (Pale flax, 2n=30) isnot found in this area it cannot have been domesticated there (Helbaek, 956). Helbaek suggested that flax was domesticated about the same time as emmer and barley in the mountains of the Near East. Thence it spread to other parts of the Old World. So itmust have been domesticated before c.6200 BC.(van Zeist & Bakker-Heeres, 975). L. bienne can be divided into two main geographic races.the first is the continental winter annual of the semi-arid foothills of Iraqi Kurdistan and Iran. It might be the parent of the prostrate multi-stemmed type cultivated since ancient times along the N. coast of Turkey, the Caspian coast of Azerbaijan and some parts of Colchis bordering the Black Sea. According to Helbaek (956), this type is the ancestor of the small-seeded flax cultivated by the prehistoric C. European pile-dwellers. The latter is the parent of 'Winterlein', a winter annual cultivated in mountainous S. Germany. The second is the Atlantic-Mediterranean GOSSYPIUM STOCKSII* Moraceae FICUS CARICA L. Common fig. 2n=26.Probably S. Asia. Primary gene centre in SE, Asia. Spread to Asia Minor, Mediterranean countries and W. Europe (Storey & Condit, 969). Long cultivated. In 4000 BC., figs were already cultivated in Egypt. In Transcaucasia, Crimea, C. Asia, Baluchistan and the Mediterranean countries, it ranwild a long time ago. Aweke (979) suggests that Ethiopia or at least Africa might be the area of origin of the fig and that F. palmata* is its ancestor. Var. transcaspica from the Kopet Dagh of Turkmenia is a source of frost resistance. FICUS SYCOMORUS L. Sycomore fig. 2n=26. Its distribution is given on p. 6.Galil et al. (976) suggested that the sycomore was domesticated in the Middle East where man was forced to propagate the tree vegetatively because of the lack of the specific pollinator. Nelumbonaceae NELUMBO NUCIFERA Gaertn. Indian lotus. 2n=6. Centre of diversity probably lies in N. Iran, the Kura Estuary in Transcaucasia and Volga Delta. Cultivated in China,Japan and elsewhere for its rhizomes and fruits. Formerly it was also grown in the E. Mediterranean Region

91 00 NEAR EASTERN REGION (Hjelmquist, 972). Papaveraceae PAPAVER SOMNIFERUM* Polygonaceae FAGOPYRUM ESCULENTUM Moench. (syn.f. vulgare T. Nées, F. sagittatum Gilib., Polygonum fagopyrum L.). Buckwheat, Silverhull. 2n=6, 32. C. Asia. Introduced into several countries as a grain crop. It isoften found as a ruderal. It is insect-pollinated. Punicaceae PUNICA GRANATUM L. Pomegranate. 2n=6, 8, 9. Wild in the Near East and C 0 Asia. An ancient fruit-tree, which was even cultivated in the Hanging Gardens of Babylon. Cultivated now in many countries. The only related species is P. protopunica Ralf, found wild on Socotra in the Indian Ocean. sparse oak forest of Caucasia. It is polymorphous. Through introgression, characteristics such as tallness,late ripening, good transportability of fruits, high sugar content and, unfortunately, lowhardiness entered the cultivated apple (Malus pumila Mill.), as can still be recognized in Caucasian, Crimean and even Italian cultivars. MALUS PRUNIFOLIA (Willd.) Borkh. (syn. Pyrus prunifolia Willd.). Chinese apple. 2n=34, 5, 68. Primary centre in N. China. Cultivated in E. Asia for its fruits.in the USSR, this species is represented in wild forms in E. Siberia. It ishighly resistant to frost and drought, much used by I.V. Michurin to breed hybrid varieties such askandil Kitaika (Kitaika = Chinese), Bellefleur Kitaika, Saffran Peppin, Saffran Kitaika. Resedaceae RESEDA PHYTEUMA* Rosaceae AMYGDALUS BESSERIANA* AMYGDALUS COMMUNIS L. (syn.prunus amygdalus Batsch.). Almond. 2n=6. Primary gene centres in C. Asia (p. 84) and in the Near Eastern Region. AMYGDALUS FENZLIANA (Fritsch)Lipsky (syn. A. divaricata Fenzl., A. urartu S. Tam., Prunus fenzliana Fritsch.). F'enzel almond. 2n=6. S. Transcaucasia and Anatolia. An ornamental.it easily crosses with A. communis*and it might be a source of cold and drought resistance for that species. AMYGDALUS PERSICA L. Peach. 2n=6. Primary centre in China (p. 42). Secondary centre in Caucasia and Crimea. ARMENIACA VULGARIS L. Apricot. 2n=6. Primary centres in NE. China (p. 42) and in Daghestan on the slopes of the Khunzakh Plateau at an altitude of m. The latter centre probably formerly linked with the main one (p. 42). The tree has ashrubby habit. Cultivated over the entire Near East. CYDONIA 0BL0NGA Mill. Quince. 2n=34. Talysh Mountain Range (S. Daghestan), the lor Valleys and Azalan (Georgia), in the Terter Valley (Soviet Azerbaijan) and in the canyons of Aidero and Yuz-Begi, Kopet Dagh (USSR). Primary centre lies there. Long cultivated. MALUS ORIENTALIS Uglits. 2n=. This is the only wild Malus species in the especially Malus prunifolia MALUS PUMILA* MALUS TURKMENORUM Juz. & M. Pop. 2n=. Turkmenia, in the gorges of the Kopet Dagh. Primary gene centre also there. The cultivated form is known in Russian as 'Baba-arabka* (old arab woman). This name refers to the dyingdown of the main stem at an age of about 20 years and its replacement by soboles permanently rejuvenating the tree. MESPILUS GERMANICA L. Medlar. 2n=34. Caucasia, N. Iran and Asia Minor. Cultivated elsewhere and run wild there. It crosses with Crataegus oxyacantha* and Sorbus aucuparia*. PRUNUS AVIUM L. (syn. Cerasus avium Moench.). Sweet cherry, Mazzard. 2n=6, (24, 32). Pri-

92 NELUMBONACEAE -UMBELLIFERAE 0 mary centre in Asia Minor and Transcaucasia. Wild trees also in other parts of Europe, W. Asia and N. Africa.The wild trees of Ukrainia could be grouped into four classes:. darkcoloured fruit: a.bitter and b. sweet and 2. light-coloured fruit: c,bitter and d. sweet. The sweetfruited types had elongate stones and longer fruit-stalks and petioles than the bitter-fruited types (M'yakushko & M'yakushko, 970). It is likely that man selected thesweetfruited types. Rjadnova (967)suggested that domestication occurred in various places. This resulted in several ecotypes differing, for instance, in resistance to unfavourable conditions and quality of fruit. Constant selection resulted in large-fruited hardy types. P. avium is one of the parents of P s cerasus*. Hybrids with P. cerasus (P. x gondounii (Poiteau & Turpin) Rehder are known in W.Europeas 'Duke' cherries,, These hybrids and P. cerasus are sources of resistance tobacterial canker caused by Pseudomonas syringae Van Hall. PRUNUS CERASIFERA Ehrh. (syn a P. divaricata Led.). Cherry plum, Myrobalan. 2n=6,genome formula CC, (24, 32, 48). Wild in Caucasia, Iran, Asia Minor,Altai and C. Asia. Primary centre C. and S. Caucasian coast of the Black Sea, whence it spread eastwards and westwards. Secondary centre in W. Tien-Shan (p. 85). It ishighly polymorphic. This species is one of the parents of P. domestica*. It is also planted as a rootstock and in hedges. Var. pissardii (Carrière) L.H. Bailey has dark red leaves and flowers tinged with reddish pink. It is an ornamental. PRUNUS CERASUS L. (syn. Cerasus vulgaris Mill.). Sour cherry, Pie cherry. 2n=32. Unknown wild, although trees that have run wild grow mainly in Caucasia and Asia Minor, but also in the European USSR, W. Balkan countries and Germany. Probably an allotetraploid of P. fruticosa* x P. avium*.sour cherry can be divided into the true Sour cherries and 'Duke' cherries. The first can be subdivided into Morellos (austera L.) and Amarelles (caproniana L.) (Zylka,97a). A special population 'Vladimir cherry' originated in Region 9 (p. 60). PRUNUS DOMESTICA L. Garden plum, Domestic plum. 2n=48, genome formula CCSSSS orcdcdsss^^sj or CdCdDlDiD2D2. Caucasia. This species is thought to be a natural hexaploid of P. cerasifera* and P. spinosa*. This alloploidization apparently took place in Caucasia, where both species occur and natural hybrids with 2n=24 and 48 are still found. However itmay have happened elsewhere. For instance Werneck (958) considered the garden plum to have arisen in Upper Austria (p. 60). Rybin (936) resynthesized the garden plum. Artificial hexaploids resembled the natural ones. PRUNUS SPINOSA L. Blackthorn, Sloe. 2n=32, ge- nome formula SSSS of SSS]^ or SSC C. Wild throughout the entire territory of this centre and in Europe and N. Africa. Volga Basin types carry genes for high hardiness. It is one of the parents of P. domestica*. Some natural hybrids with P. domestica are described as P. fruticans Weihe (2n-40). PYRUS. The Near East is the main geographic centre of origin of Pyrus species. Of about 60Pyrus species in the world, about 25 have been described for Caucasia. Some of them also occur in Iran or in Asia Minor. PYRUS CAUCASICA Fed. 2n=. The entire forest zone of Caucasia except the Talysh Mountain Range (Soviet Azerbaijan). A polymorphic species. In open areas,it spreads quickly and vigourously. PYRUS SYRIACA Boiss.2n=. Armenia. Itis cold-resistant and probably played a part in the origin of the cultivated pear (Evreinov, 944). Cultivated locally. PYRUS TAKHTADZHIANA Fed. 2n=. Habit of a cultivated tree. Cultivated in ancient times but later ran wild. ROSA CENTIFOLIA L. (syn.r. gallica L. var. centifolia Reg.). Provence rose.2n= : 28. E, Caucasia. Cultivated for its flowers. The petals are used in the perfume industry. SORBUS DOMESTICA L. Service tree. 2n=34. Its distribution is given on p. 6. Large-fruited forms are found in forests of Crimea. Rubiaceae COFFEA ARABICA L. Arabica coffee. 2n=22, 44, (66). The primary centre in SW. Ethiopia. Secondary centre in Yemen.This area is the source of Arabica coffee now cultivated in Latin America, Kenya, India, Java and elsewhere (Meyer, 965). Rutaceae CITRUS MEDICA L. Citron. 2n=8. Probably SW. Asia, although India has often been mentioned as centre of origin.unknown wild. It has now spread through the (sub)tropics. The Etrog citron (var.ethrog Engl.) is used by Jews at the Feast of Tabernacles, and the fingered citron (var. sarcodactylis Noot.) Swing)by the Chinese as a medicine and an ornamental. Umbelliferae CUMINUM CYMINUM* DAUCUS CAROTA L. Carrot. 2n=8. For origin see p. 86.By hybridization between the 'Eastern' and 'Western' carrots in Turkey, a secondary centre of diversity has developed there.

93 02 NEAR EASTERN REGION MALABAILA SECACUL (Mill.) Boiss. Sekakul. 2n=. Asia Minor and Syria. Cultivated for its roots, used as a aphrodisiac, PIMPINELLA ANISUM L. (syn. Anisum vulgare Gaertn.,Anisum officinarum Moench). Anise plant. 2n=8, 20 0 Probably the Orient. Cultivated for aromatic fruits. Valerianaceae VALERIANA PHU L 0 2n=. N. Anatolia. Cultivated for its rhizome, which yields the drug valerian. Vitidaceae VITIS LABRUSCA L. Fox grape. 2n=38. N. America (p. 206). Introduced into W. Georgia (USSR)as a cultivated grape. VITIS VINIFERA L. Common grape, European grape. 2n=38, (40, 57, 76). Primary centres: the Central Asian (p. 86), the Near Eastern and the Mediterranean Regions (p. 9). The wild vine, ssp. sylvestris Gmel., is found in regions bordering the Mediterranean Sea, except Libya and Egypt,up to Turkestan and Kashmir. Primary centre:probably Armenia (USSR) and N. Iran. The western wild types have been called ssp. silvestris, and the eastern types ssp. caucasia Vav. Thewild type is dioecious and the domesticated type (ssp. sativa DC, ssp. vinifera)is hermaphrodite derived from the male wild plants. The vine may have been domesticated in SE. Europe where types with large bunches, and seedless grapes have developed. Natural hybrids are still forming in several areas, e.g. the mountains of S. Tajikistan (USSR), where many new forms are observed. From crosses between the wild grape and cultivated types in Europe, old and new cultivars developed. The common grape has been crossed with the North American V. labrusca*.the fruits are used to prepare wine, currants and raisins. V. amurensis* is a possible source of hardiness.

94 7 Mediterranean Region The Mediterranean Region wasdescribed byvavilov.darlington (956)suggested thenamemediterranean Region of Origin. Itssituation near thecradle of Agriculture inthenear East led toan early introduction ofplant cultivation.early farming siteshavebeen found atnea Nikomedeiain Greece dating c.5470 BC. (vanzeist & Bottema, 97)and atfayum inegypt dating from the 5thMillenium, reaching the coast ofthe Atlantic perhaps c.3rdmillenium. Avery old site atkomombo inthenile ValleyofUpperEgypt dated from BC.Itis a non-farming site occupied thewhole year round (Churcher &Smith, 972). Many cropshavebeen domesticated intheregion including Avena sp., Beta vulgaris,brassica napus, B. oleracea, Lathyrus sp., Linum usitatissimum, Lolium sp., Lupinus sp., Oleaeuropaea, Raphanus sativus, Trifolium sp.and Vitis vinifera. Alliaceae ALLIUM CEPA L; Spanish onion. 2n=6. See p. 8. Secondary centre in the Mediterranean Region. ALLIUM SATIVUM L. Garlic. 2n=6, genome formula SS. C. Asia (p. 8). Secondary centre in the Mediterranean Region (Kazakova, 97). Amaranthaceae AMARANTHUS LIVIDUS L. 2n=34. Spread through Europe, Asia and to the tropics of theold and New World. Var. ascendens Thell. (syn. A. viridis L., 2n=34) is native to S. Europe and E. Mediterranean Region. Cultivated there in the Middle Ages. Var. lividus is unknown wild. It might be a cultigen of this species. It was cultivated in the 6th and 7th Centuries as a vegetable and medicinal crop and in the 8th Century aspig food. Var. oleraceus Thell. (syn. A. oleraceus L., 2n= )is probably a cultigen of var. ascendens. Cultivated in Europe and elsewhere as a vegetable (Mansfeld, 959). Amaryllidaceae NARCISSUS JONQUILLA L. Jonquille. 2n=4. Europe, Ante-Asia to Iran and Algeria. Commonly cultivated as an ornamental and in S. France for its essential oil. NARCISSUS P0ETICUS L. Poet's narcissus. 2n=4, 2. Portugal, Spain,France and Italy. Cultivated as an ornamental and in S. France for its essential oil. Anacardiaceae RHUS CORIARIA L. Sicilian sumach. 2n=. Mediterranean area. A shrub cultivated in Sicily and S. Italy for the leaves, which are asource of tanning material. Apocynaceae NERIUM OLEANDER L. Oleander. 2n=6, 22. A shrub of Mediterranean area. Cultivated as an ornamental. Asclepiadiaceae CYNANCHUM VINCET0XICUM* Balanitaceae BALANITES AEGYPTIACA Del. Betu,Desert date. 2n=6, 8. This shrub grows wild in Arabia, Palestine, N. Trop. Africa and Angola. Cultivated in Egypt for its edible leaves and flowers (Cufodontis, 957; Terra, 967). Boraginaceae

95 04 MEDITERRANEAN REGION ALKANNA TINCTORIA (L.) Tausch. Alkanna. 2n= 4. S. and E. Europe and Turkey. A herb cultivated as asource of ared pigment. BORAGO OFFICINALIS L. Borage. 2n=6. Mediterranean Region. A herb cultivated as an ornamental, as a pot-herb and for bees. Capparidaceae CAPPARIS SPINOSA L. Caper bush. 2n=24, 38. The cultivated forms with large flower-head, var. spinosa, and small flower-head, var. parviflora J. Grey probably derived from thewild var. aegyptia (Lam.) Boiss. This variety grows wild in S. and SE. Mediterranean area to the Sudan and Eritro-Arabia. Var. spinosa developed in the N. Mediterranean Region, whence it spread to other areas, where it is cultivated as a condiment. Var. parviflora is also cultivated and might be amutant of var, spinosa. Hybrids with C. ovata Desf. are found. Caryophyllaceae DIANTHUS CARYOPHYLLUS L. Carnation, Clove, Pink, Picotée. 2n=30. Mediterranean Region. A perennial herb cultivated as an ornamental and also as asource of an essential oil. GYPSOPHILA PANICULATA L. Baby's breath. 2n= 34 0 S. and C. Europe and Caucasia. Cultivated formerly for its roots, which contain saponin. Now it is an ornamental. Chenopodiaceae BETA PATELLARIS Moq. 2n=8, (36). Mediterranean and Atlantic coasts of NW. Africa, Canary Islands, Cape Verde Islands and Madeira. A source of resistance tonematodes and Cercospora and tolerance to yellow mosaic for B. vulgaris*. BETA PROCUMBENS Chr. 2n=8. Canary Islands and Cape Verde Islands. A source of nematode resistance for B. vulgaris*. BETA VULGARIS L. Beet. 2n=8, genome formula VV. The parental form is the wild sea-beet (ssp. maritima (L.) Thell., syn. B. maritima L.). Primary centre probably in the E. part of the Mediterranean Region. Spread in a westerly direction along the Mediterranean, Atlantic coast of Europe and to Cape Verde Islands and Canary Islands. In the Mediterranean Region, leaves and roots of the wild plant may have been collected, perhaps leading to development of Swiss chard and Spinach beet (var, cicla, var. vulgaris), whose leaves and stalks are eaten, and to garden beet, table beet and red beet (var. cruenta, var. esculenta). Development may have been influenced by hybridization with wild types like ssp. macrocarpa (syn s B. macrocarpa Guss.) in N. Africa. In California, such hybridization still continues (McFarlane, Beta vulgaris (), B. patellaris (2) atriplicifolia (5)(Ulbrich, 934). procumbens and B. webbiana (3), B. patula (4)and B.

96 BORAGINACEAE -COMPOSITAE 975). The fodder beet (var. rapa) probably developed in the Netherlands (p. 49), after introduction of types from Spain, and the sugar -beet in Silesia (Poland) (p. 49). The wild B. macrocarpa Guss. from the coasts of the Mediterranean Region and Canary Islands, B. patula Ait. from Madeira and B. atriplicifolia Rouy from S. Spain easily hybridize with B. vulgaris,with which they may be included as subspecies. In NW. Europe,hybrid plants of cultivated sugar-beet and ssp. maritima are occasionally observed. They derive from material propagated in France and Italy. Such hybrids bolt in the first year, producing seed. The seed drops and may result in a weed (F2 plants) for several years. It is possible that, on avery small scale, wild genes derived from these hybrids introgress into the cultivated population. The wild plants may form sources of resistance to disease such as Cercospora, yellow mosaic and increase the variation for selection of new high-yielding types. BETA WEBBIANA Moq. 2n=8. Canary Islands. A source of nematode resistance for B, vulgaris*. CHENOPODIUM AMBROSIOIDES L. American wormweed, Indian wormweed. 2n=6, 32, 36, 64. Probably S. Europe. Widespread in the tropics and introduced into N. America.The cultivated type, var. anthelminticus L. is a source of medicinal and essential oils. HALOGETON SATIVUS (L.) Moq. 2n=. NW. Africa. Cultivated in the Mediterranean Region for the base-rich ash it yields when burned. Compositae ANACYCLUS OFFIÇINARUM Hayne. Bertram. 2n=8. Probably the Mediterranean Region. Formerly cultivated in C. Europe. ANACYCLUS PYRETHRUM (L.) Link. Pellitoria of Spain. 2n=8. N. Africa, Arabia and Syria, Formerly cultivated in Europe as a medicinal plant and now in Algeria for an essential oil. ARTEMISIA JUDAICA L. 2n= the Mediterranean Region. Cultivated in CALENDULA OFFICINALIS L. Marigold. 2n=(28), 32. Centre of origin probably in the Mediterranean Region. Cultivated as an ornamental, but formerly as a medicinal plant. CICHORIUM ENDIVIA L. Endive, Escarolle. 2n=8. So Europe to India. Cultivated as a vegetable. CNICUS BENEDICTUS L. (syn. Centaurea benedicta L., Garberia benedicta Adans.). Blessed thistle. 2n=22. Mediterranean Region to Transcaucasia, Syria and Iran. Formerly cultivated in Germany. CYNARA CARDUNCULUS L. (Wild) cardoon. 2n=34 0 W, and C. Mediterranean area.cultivated for its leaf stalks. Probably together with C. syriaca*, one of the parents of C. scolymus* (Zohary & Basnizky, 97.5). CYNARA SCOLYMUS L. Artichoke, Globe artichoke. 2n=34. Mediterranean area. Cultivated for soft fleshy edible receptacles of young flower heads and thick bases of the scales around the flower heads as well as for a source of a bitter compound. Several varieties are known. If it derives from C. syriaca* with introgression of C. cardunculus*, it originated in W. Mediterranean area; if it derives from C. cardunculus with introgression of C. syriaca,it originated in E. Mediterranean area (Zohary & Basnizky, 975). CYNARA SIBTHROPIANA Boiss. & Heldr. 2n= Mainly on Aegean Islands, Crete and Cyprus. Related to C. cardunculus*and C. scolymus* (Zohary & Basnizky, 975). CYNARA SYRIACA Boiss. Wild Syrian artichoke. 2n=. Levant and S. Turkey. Probably one of the ancestors of C. scolymus*. LACTUCA VIROSA L. Bitter lettuce, Lettuce opium. 2n-8. Primary centre round the Mediterranean (Lindqvist, 960). Cultivated on a small scale in some parts of Europe for its latex, which has narcotic properties. SCOLYMUS HISPANICUS L. Golden thistle, Spanish oyster plant, 2n=20. Mediterranean area. A root vegetable. Its cultivation is declining. SCORZONERA HISPANICA L. Scorzonera, Black salsify. 2n=4. C. Europe, Mediterranean area, Caucasia and S. Siberia. A vegetable especiallyof S. Europe. Perhaps it was first cultivated in Spain (Mansfeld, 959). SILYBUM MARIANUM (L.) Gaertn.Holy thistle, Milk thistle, Lady's milk. 2n=34. S. Europe. Cultivated as a medicinal plant and as an ornamental. TANACETUM CINERARIIFOLIUM (Trev.) Schultz Bip. (syn. Chrysanthemum cinerariifolium (Trev.) Brocc. Pyrethrum. 2n=8. Dalmatian coast including Yugoslavia and Albania. Introduced into many countries. Kenya is the main producer of the insecticide Pyrethrin. TANACETUM PARTHENIUM (L.) Schultz Bip. (syn. Chrysanthemum parthenium (L.) Bernh., Leucantheraum parthenium (L.) Gren. & Godron, Pyrethrum parthenium (L.) Sm). Feverfew, Wild camomile. 2n=8. Mediterranean area, Balkan, Asia Minor and Caucasia, Cultivated as medicinal plant and as an ornamental. TRAG0P0G0N PORRIFOLIUS L. Salsify, Oyster plant, Purple goats beard. 2n=2. Mediterranean Region. This vegetable was first cultivated for its roots long ago. Itmay have been

97 MEDITERRANEAN REGION eties have been described (Mansfeld, 959): var. oleiformis Pers. (R. chinensis Mill,)is the oil-seed radish cultivated in India, Japan, China (p. 35)and on asmall scale in Rumania and Spain; var. mougri Helm* (syn. R. caudatus L.); var. sativus, the radish, small radish ; var. niger Kerner, radish, Spanish radish. Recently fodder radish has been bred.it is a reputed selection from oil-seed radish in France. More research is needed to ascertain the origin of radish and the various botanic varieties and cultivars.through natural (and artificial) hybridization with Brassica spp.,genes may introgress intor. sativus. SINAPIS ALBA L. (syn.brassica alba (L.)Boiss.). White mustard. 2n=24, Mediterranean area. The wild plant is low-growing and much-branched, with siliquae containing browny black seeds (melanosperma) (Hemingway, 976), Weedy or naturalized plants may be found from Spain through Asia Minor to E. India.Young seedlings are used as salad. Seeds are the source of white mustard. Cucurbitaceae BRYONIA CRETICA* CITRULLUS COLOCYNTHIS (L.) Schrad. Colocynth. 2n=22, (34). Arid regions of N. Africa and Trop. Asia. Cultivated in India and the Mediterranean area for its purgative fruits. ECBALLIUM ELATERIUM (L.) A. Rich. Squirting cucumber. 2n=(8), 24. Mediterranean area, Azores, Asia Minor and Crimea. Cultivated in England as a medicinal plant, Cyperaceae CYPERUS ALOPECUROIDES Rottb.Mat sedge.2n=. Trop.Old World, Cultivated inegypt for mat-making (Mansfeld, 959) 0 CYPERUS ESCULENTUS L. Chufa,Earth almond, Tiger nut,rush nut, Zulu nut, Yellow nutgrass. 2n=(8), 08. White Nile region and in the tropics. Introduced to S,Europe by the Arabs. Cultivated in Spain, Italy and elsewhere for its flavoured tubers. The wild form is var. aureus (Ten,) Richt, and the cultivated form is var. esculentus. CYPERUS PAPYRUS L. Papyrus plant. 2n=c.l02. Africa. Formerly cultivated in Egypt, Palestine and the Mediterranean Area. Now rarely cultivated. It could probably remedy and prevent eutrophication of tropical lakes by nutrient extraction. 'More active' extractor genotypes could perhaps be obtained by breeding. Ericaceae ARBUTUS UNEDO L. Strawberry tree, Arbutus.2n= 26. Mediterranean Region. Occasionally cultivated for its edible fruits. Arbutus unedo (Hutchinson, 969). Euphorbiaceae CHROZOPHORA TINCTORIA (L.) Juss. Giradol. Mediterranean Region, France, Yugoslavia, Crimea to W. Asia, NW. India, Arabia. Formerly cultivated in S. France as asource of red and blue dye. The red dye was used for colouring Dutch cheeses. EUPHORBIA LATHYRUS L. 2n=20. S., W. and C. Europe. A ruderal and weedy plant occasionally cultivated as a medicinal. Probably only native to E. and C. Mediterranean Region. Fagaceae QUERCUS SUBER L Cork oak. 2n=24. W. Mediterranean Area. A very variable species 0 Cultivated in S. France, Portugal, Spain, Sardinia, Corsica, Istria, Dalmatia and Algeria. Geraniaceae ERODIUM CICUTARIUM (L.) L'Herit. ex Ait. Storksbill, Red-stem filaree, Alfilaree. 2n=(20, 30-40, 36), 40, (48, 54). S., W. and C. Europe, Mediterranean area, Temp, Asia. Cultivated as fodder for sheep in N. and S. America. ERODIUM MOSCHATUM (L.) L'Herit. ex Ait, Musk storksbill, White-stem filaree. 2n=20 e Mediterranean Region. Formerly cultivated as a medicinal crop u Gramineae AEGILOPS BICORNIS (Forsk.) Jaub. & Sp. (syn. Triticum bicorne Forsk.). 2n=4, genome formula S b S. Xeric sandy soils of S. Israel, Lower Egypt and Cyrenaica (Libya). It is sometimes believed to be the B donor of tetraploid and hexaploid Triticum spp. (p. 93). AEGILOPS C0M0SA Sibth. & Sm. (syn.triticum comosum (Sibth 0 & Sm.) Richter). 2n=4, ge-

98 CRUCIFERAE - GRAMINEAE 09 nome formula MM. Mediterranean Greece, the Aegean Islands and W. Turkey. Used as asource of resistance to yellow rust (Puccinia striiformis West.). AEGILOPS CYLINDRICA* AEGILOPS KOTSCHYI* AEGILOPS LORENTII* AEGILOPS OVATA* AEGILOPS TRIARISTATA* AEGILOPS TRIUNCIALIS* AEGILOPS UNIARISTATA Vis. (syn.triticum uniaristatum (Vis.) Richter). 2n=4, genome formula M^lV The Mediterranean Greece, around the Sea of Marmara and the Adriatic coast of Yugoslavia. AEGILOPS VARIABILIS Eig. (syn. Ae. peregrina (Hack c) Maire & Weill., Triticum peregrinum Hack & Fraser). 2n=28, genome formula C U C U S V S v. N. Africa, Egypt, Palestine, Greek Islands, Turkey and Iraq.Probably identical with Ae a kotschyi*. AEGILOPS VENTRICOSA Tausch, (syn.triticum ventricosum Ces., Pass. & Gib a). 2n=28, genome formula M V M V DD. W. Mediterranean Area. It is a source of resistance to the wheat disease eyespot caused by Cercosporella herpotricoides Fron.Natural hybrids with Triticum turgidum group durum have been found and described astriticum rodeti Trabut. Amphiploids with tetraploid Triticum species have been named Aegilotricum. AGROPYRON JUNCEUM (Jusl.) Beauv. Sea wheatgrass, Bent grass. 2n=28, 42, (84). Coasts of Europe, N. Africa and Asia Minor. Occasionally cultivated to stabilize dunes. AGROSTIS TENUIS* ARUNIX) DONAX L. Giant reed D 2n=(c.60), 0, Mediterranean area to Caucasia and Syria. A grass cultivated since ancient times in S. Europe. Also cultivated elsewhere now. AVENA CANARIENSIS Baum, Rajhathy & Sampson. 20=4, genome formula AcAc u Uplands of Fuerteventura, Canary Islands. It is the donor of the A genome of the evolutionary complex of maroccana*, A. murphyi* and A. sterilis* (Craig et al., 974; Leggett, 980). AVENA CLAUDA Dur. 2n=4. The whole Mediterranean Basin from Morocco, eastwards. It usually grows together with A. sativa type sterilis* and A. strigosa type barbata* (Ladizinsky & Zohary, 97). This wild species includes type eriantha (syn. A. eriantha Dur., A. pilosa MB; genome formula CpCp) and type clauda (A. clauda Dur.) u AVENA DAMASCENA Rajhathy & Baum. 2n=4, genome formula AdAd. An area 60km north of Damascus, Syria. Ithas a high degree of genome homology with A. prostrata*. Both species are considered relics of a once common population, but are now separated by some 2500km (Rajhathy & Baum, 972). Cahana & Ladizinsky (978) consider A. damascena toderive from A. prostrata*. It resembles A. strigosa*. AVENA L0NGIGLUMIS Dur. 2n=4, genome formula AA. The coastal fringe of Mediterranean countries and Morocco,Portugal and Spain. Mediterranean and Negev desert ecogeographic races have been recognized (Ladizinsky & Zohary, 97). It derives from A. prostrata* (Cahana & Ladizinsky, 978). AVENA MAROCCANA Gandog (syn. A. magna Murphy & Terrell). 2n=28, genome formula AACC. Morocco. Probably not an ancestor of A. sativa* (Leggett, 980). An annual belonging to the A. maroccana (magna)-a. murphyi*-a. sterilis complex. It isoften confused with A, sterilis. AVENA MURPHYI Ladizinsky. 2n=28, genome formula AACC. Between Tarifa and Vejer de la Frontera, S. Spain. Probably not an ancestor of A. sativa* (Leggett, 980). It belongs to the A. maroccana* (magna)-a, murphyi-a. sterilis* complex. AVENA PROSTRATA Ladizinsky. 2n=4, genome formula ApAp, SE, Spain.Parental genome donor of A. longiglumis* and A. damascena* (Cahana & Ladizinsky, 978). It isnot the ancestor of the A. maroccana*-a, murphyi* complex (Leggett, 980). AVENA SATIVA L. Oat. 2n=42, genome formula AACCDD. Two species of hexaploid oats are commonly recognized: A. sativa, characterized by florets that separate by fracturing of the rachilla, leaving a section of rachilla attached to each floret after threshing; and A. byzanthina C. Koch in which the basal floret leaves an abscission scar on threshing. Itis widely held that these two complexes were independently domesticated (Bell, 965)but a monophyletic origin of oats is equally likely (Coffman, 946), as becomes obvious if A. sterilis is accepted as the wild progenitor of domesticated oats. Florets of A. sterilis occur among remains of cultivated wheat and barley in agricultural settlements from Europe to China. In the northern extremes of wheat cultivation the better adapted weed was eventually adopted as a cultivated cereal. The cytoplasm comes from adiploid species with A genome (Steer & Thomas, 976). The winter crop Dormoats is an oat with a deep dormancy deriving from A. septentrionalis* (A. fatua) x A. sativa. AVENA STERILIS L. (syn. A 0 athenathera Presl,

99 MEDITERRANEAN REGION Avena clauda (Ladizinsky & Zohary, 97). Avena longiglumis (Ladizinsky & Zohary, 97). A. trichophyllac. Koch), Wild oats. 2n=42. Near East, Mediterranean Region of Europe and North Africa. Wild oats is characterized by dispersal units that disarticulate through abscission callus below the basal floret of each spikelet. Florets separate laterby fracturing of rachilla segments. It is often assumed that the mimetic weed oats (A. fatua L., A. hybrida Petterrct,, A. occidentalis Dur. as recognized by Baum, 977) are progenitors of domesticated oats. This is unlikely. These weeds are characterized by florets that disarticulate individually by formation of abscission callus. Lack of a mechanism for seed dispersal in domesticated oats is genetically dominant over fatua-type seed dispersal. A more likely explanation is that the fatua-type dispersal evolved after oats became domesticated. AVENA STRIGOSA Schreb. Black oat, Bristle oat. 2n=4, genome formula AsAs, 2n=28, genome formula AsAsBB, AABB or AsAsAsAs. The As and B genomes are partially homologous and may derive from a common parent (Ladizinsky & Zohary, 97; Ladizinsky, 973). The As genome might be the prototype of the A genome of the polyploid species (Rajhathy et al., 97). Ladizinsky & Zohary (97) included in this species the wild A. hirtula Lag. (2n=4), A. wiestii Schreb. (2n=4), A. barbata Pott. (2n= 28) and A. vaviloviana Malz 0* (2n=28), and the cultivated A. strigosa Schreb, (2n=4)and A. abyssinica Höchst.* (2n=28). Leggett (980) gives the following genome formulas: A. hirtula-wiestii oat group AsAs and A. barbata AABB. All over the Mediterranean area, wild and weedy diploid and tetraploid forms are found, hybridizing freely. "A. hirtula" is common in Spain, Morocco, Algeria, Italy, Greece, Turkey and Israel "A. wiestii" grows in the drier steppe of the northern fringes of the Sahara and the Arabian Desert. The cultivated strigosa of W. and N, Europe derives from the weedy forms common in cereal fields and edges of cultivation in the Iberian peninsula. The As and B genome are partially homologous and may derive from a common parent (Ladizinsky & Zohary, 97). The As genome might be the prototype of the A genome of the polyploid species (Rajhathy et al., 97) Diploid and tetraploid cytotypes introgress by means of triploids. AVENA VENTRICOSA Balansa. 2n=4, genome formula CvCv. This wild species includes ssp. bruhnsiana (Grüner) Malzew (syn. A. bruhnsiana

100 GRAMINEAE -GRAMINEAE Avena ventricosa (Ladizinsky & Zohary, 97). Grüner) and ssp. ventricosa (Balansa) Malzew (syn. A. ventricosa Balansa s. Str., genome formula AvAv). Ssp. bruhnsiana is found in the Apsheron Peninsula of Soviet Azerbaijan and ssp. ventricosa in Algeria and Cyprus. The karyotype of ssp. ventricosa is c v* and of ssp. bruhnsiana c v3 and c y2 (Rajhathy, 97). A. ventricosa is also found in Cyrenaica (Libya) and Iraq (Ladizinsky & Zohary, 97). CHRYSOPOGON GRYLLUS (Torner) Trin. (syn 0 Andropogon gryllus Torner). 2n=20, 40. Mediterranean area to India, Cultivated in the Po plain, Italy for its essential oil. HORDEUM VULGARE L. 2n=4. For origin of barley see p. 9. The Mediterranean Region is the centre of origin of ssp. mediterranean Vav, & Bacht. LOLIUM MULTIFLORUM Lam. ssp.italicum (A.Br.) Volkart ex Schinz & Kell. Italian ryegrass. 2n=4, The irrigated lands of Lombardy in N. Italy. Probably cultivated there in the 3th or 4th Century (Beddows, 953). Spread to N. Europe, LOLIUM PERENNE* PHALARIS CANARIENSIS L. Canary grass. 2n=2. W. Mediterranean area: Canary Islands, Spain, Portugal, Cultivated for birdseed. PHALARIS TUBEROSA L. (syn. Ph. aquatica L.). Toowoomba grass, Harding grass. 2n=28. Mediterranean area. Cultivated in warm countries. SORGHUM BICOLOR (L.) Moench. Broomcorn. 2n= 20. Sorghum originated in Africa (see p. 33). The broomcorns developed in the Mediterranean area from material that came from India/Iran or Africa through the Middle East. SORGHUM HALEPENSE (L.) Pers. (syn.s. miliaceum (Roxb.) Snowden, S.controversura (Steud.) Snowden). Johnson grass. 2n=40. Mediterranean area topakistan and S. India. This rhizoma- Sorghum halepense. tous perennial was introduced as a fodder to all warmer parts of the world. The leaves and stems contain HCN but make excellent hay. In the Americas, S, halepense has widely introgressed with grain sorghums (Celarier, 958). Derivatives of such introgression are known ass. almum Parodi (Columbus grass) in Argentina (Saez, 949) (p. 7). Some new perennial diploid types were selected from the cross S 0 halepense x S, bicolor. These types combine high yield and palatibility with some frost tolerance and disease resistance. Their origin is similar to S. almum*. STIPA TENACISSIMA L. Haifa, Alfa, Esparto. 2n=32, 40. Mediterranean area. In Spain, some cultivation isdone with cv. Albardin, which has a larger fibre than wild ones. The variety

101 2 MEDITERRANEAN REGION Sorghum halepense (de Wet & Huckabay, 967). seems to have developed there. In N, Africa and Spain, wild halfa yields a fibre for papermaking. TRITICUM TURGIDUM spp. turgidum conv. durum (Desf.) Mac Key. 2n=28, genome formula AABB. It originated during cultivation of emmer (p. 95). Secondary centre in the Mediterranean area. ZEA MAYS L. 2n=20. Maize was domesticated in C. America (p. 98). Secondary centres in Mediterranean area and in the Nile Basin (Brandolini, 970). CROCUS SATIVUS L. Saffron crocus 2n=(4, 6), 24, (40). Mediterranean area and Ante-Asia. Cultivated since ancient times for its styles, which are a source of saffron. Formerly cultivated for this purpose in Europe and N. America, and now in S. Europe, Asia Minor, Iran, N. India and China. The origin of present-day cultivars is not known. IRIS GERMANICA L. German iris, Flag iris. 2n= 24, (34, 36), 44, 48, (60). Mediterranean area. A perennial herb widely cultivated as an ornamental and for its rootstocks, which are used in perfumery. Labiatae / rn / ^ - " &y - ^ "~\ (^^it*^^ h ' \ ^O ^^>w0 *; ^ V - - Spread of maize in the West European and the Mediterranean region, indurata ( ), indentata ( ), century of introduction (roman number) (Brandolini, 970). Grossulariaceae RIBES MULTIFLORUM Kitt. 2n=6. Mediterranean area. It isone of the parental species of present-day red currant cultivars (p. 55). Hippocastanaceae AESCULUS HIPPOCASTANUM L. Horse-chestnut. 2n= 40. C. Balkan Peninsula, E. Bulgaria, W. Iran and the Himalayas. Cultivated as an ornamental or shade tree,and for its timber, A. carnea Hayne (2n=40, 80) is a hybrid with the N. American A. pavia L.,Red buckeye (2n=40). Iridaceae HYSSOPUS OFFICINALIS L. 2n=2. Mediterranean area, Asia Minor and Iran. Cultivated for its essential oil,as a medicinal plant and as an ornamental. LAVANDULA LATIFOLIA Medik. Broad-leaved lavender. 2n=54.Cultivated in S. France and occasionally in C. Europe for itsoil of Spike. The cultivated plants are often hybrids with L. officinalis*. LAVANDULA OFFICINALIS Chaix. (syn. L. angustifolia Mill., L. spica L.). Lavender. 2n=(36), 54. Primary centre. An old cultivated plant for perfumery. First used as an insect repellant.many cultivated varieties are hybrids with wild plants and L. latifolia*. MELISSA OFFICINALIS L. Common balm. 2n=32, 64. E. Mediterranean area to Caucasia, SW. Siberia, S. Iran, Turkestan and Syria. Cultivated formerly in Europe and elsewhere for diverse purposes.the commonest cultivated type var. officinalis is perhaps derived from var. hirsuta Pers. (syn. M. hirsuta (Pers.)Hörnern.), a variety from the Balkans. MENTHA AQUATICA L. (syn. M. citrata Ehrh.). Bergamot mint. 2n=(36, 60), 96,genome formula R a R a SSJJA a qa a <i, c.96. S. Europe, Asia and N. Africa. It is a source of an essential oil. ItsA ac lgenome ispartial homologous with the

102 GRAMINEAE - LEGUMINOSAE A genome of M, arvensis var. piperascens* (Ikeda & Ono, 969). It is one of the parents of M. x piperita*. M. aquatica is cultivated as M. citrata in the USA for its lavender-like oil used in perfumery (Todd & Murray, 968). Patented hybrids with M. crispa L. (syn. M. spicata* var.crispata Sehrad.) are also cultivated inthe USA (M.J. Murray, pers. comm., 97). MENTHA LONGIFOLIA (L.) Huds. (syn. M. spicata L. var. longifolia L., M. sylvestris L.), Horse-mint. 2n=8, 24, (27, 36, 48). S. and C. Europe, N. Africa, Ethiopia, Arabia, Ante- Asia and C. Asia, Formerly it was much cultivated. Now only var. crispa Benth. is cultivated. It is related to M. rotundifolia* and M. spicata*. MENTHA PULEGIUM L. Penny royal, Pudding grass. 2n=(0), 20, (30), 40, (40-42). Mediterranean area and Europe to Iran. Formerly cultivated in Europe and elsewhere, ORIGANUM MAJORANA L. (syn. Majorana hortensis Moench.). Marjoram. 2n=30. Wild on Cyprus,in SW. Turkey, Palestine and E 0 Egypt 0 Subspontaneous in the Mediterranean area 0 Cultivated all over the world. Hybrids with 0. vulgare* have been described as 0. x applii (Domin) Bores, 2n= and 0. x majoricum Cambessedes, 2n=. 0. x applii occurs in gardens in W. and C. Europe. 0. x majoricum grows wild on the Balearic Islands,Spain and Portugal (Ietswaart, 980). Cultivated as medicinal plant. ROSMARINUS OFFICINALIS L Q Rosemary. 2n=24. Mediterranean area. Cultivated as ornamental and for its aromatic oils. SALVIA OFFICINALIS L Sage, Dalmatian sage. 2n=4, (6). Mediterranean area. A culinary herb now cultivated in many gardens in temperate and tropic countries. SALVIA SCLAREA L. Clary sage, Clary wort. 2n= 22. Mediterranean area to Iran and Transcaucasia. Formerly cultivated in the Mediterranean Region and S.Europe for various purposes, e.g. flavouring wine and beer. SALVIA VIRIDIS L. Bluebeard. 2n=6. Mediterranean area to Iran, Cultivated locally for its oil to flavour wine and beer. SATUREJA HORTENSIS L. (incl 0 S. laxiflora C 0 Koch and S. pachyphylla C. Koch). Summer savory. 2n= Mediterranean area, C. Europe and Siberia.Cultivated for oil of savory and as a pot-herb. SATUREJA MONTANA L. (syn. S. obovata Lag., S. illyrica Host). Winter savory. 2n=2, 30. Mediterranean area to Ukraine. Cultivated ins. Europe and Germany. TEUCRIUM CHAMAEDRYS L. (syn. T. officinale LamJo Common germander, 2n-32, 60, 64, Mediterranean area, France, C. Germany to S. Ural, Iran, N. Syria and Morocco, Formerly cultivated as a medicinal crop, TEUCRIUM MARUM L. 2n=. W. Mediterranean area and S. France,Cultivated in S. Europe and formerly in Germany. THYMUS VULGARIS L. Thyme. 2n=30. Mediterranean Region. Now cultivated in temperate and tropical countries. Lauraceae LAURUS N0BILIS L. Laurel,True bay, Sweet bay. 2n=42, 48. Mediterranean Region. Primary centre also there. Cultivated there and elsewhere for its leaves, which are used as a condiment. Leguminosae ASTRAGALUS B0ETICUS L.Milk vetch, Loco.2n= 6, 30. S. Europe and Mediterranean area. Cultivated as a substitute for coffee. CERATONIA SILIQUA L. Carob, Locust tree, St. John's bread. 2n=24. Mediterranean area, Syria and adjacent countries. Primary centre in this Region. Cultivated especially on Cyprus as a fodder crop. The fruits are eaten and the seeds are used toprepare carob coffee. More uses are given by Uphof (968), CERCIS SILIQUASTRUM L. Judas tree. 2n=4. A tree of the Mediterranean Region to Crimea and Iran, Cultivated for its leaves (vegetable). CICER ARIETINUM L Q Garbanzos, Chick-pea. 2n= 4, 6, (24, 32, 33). Probably W. Asia (p. 96). Secondary gene centre in the Mediterranean area. Especially large-seeded types, race mediterraneum Pop., are cultivated. CYTISUS CANARIENSIS (L.). 0. Kuntze. Genista. 2n=46.Canary Islands. Cultivated elsewhere. Used inmexico as hallucinogen. CYTISUS PALLIDUS Poir.2n= Cultivated as a forage crop. Canary Islands. CYTISUS PR0LIFER Kit. (syn. C. pullilans Kit.). Tree lucerne, Tree alfalfa, Tagasaste, Escabon, 2n=48, Canary Islands or, according to Uphof (968), Hungary, Cultivated there as a forage plant. Introduced into New Zealand. GLYCYRRHIZA GLABRA* HEDYSARUM C0R0NARIUM L. Spanish esparcet. 2n= 6. Mediterranean area. Cultivated as a fodder crop. LATHYRUS ANNUUS L. 2n=4. Mediterranean area and Portugal.Sometimes cultivated as a fodder.

103 4 MEDITERRANEAN REGION LATHYRUS CICERA L. Vetchling, Flat-pod peavine, Jurosse, Garousse. 2=4. Mediterranean area, Canary Islands, Iraq, Iran and Transcaucasia.Cultivated ins. Europe as a fodder crop and as agreen manure. LATHYRUS CLYMENUM L. (syn. L. purpureus Desf., L. alatus Sibth. & Sm.). Cicerchia porporina. 2n=4. Mediterranean area and Madeira. Cultivated in S. Europe. LATHYRUS HIRSUTUS L. Rough pea, Caley pea, Singletary pea t 2n=4, Mediterranean area. Cultivated especially inusa as a pasture hay, winter cover and for soil improvement. LATHYRUS OCHRUS DC. 2n=4. Mediterranean area. Occasionally cultivated in Greece as a fodder. LATHYRUS 0D0RATUS L. Sweet pea. 2n=4. Mediterranean area. Seeds ofwild plants were sent from Sicily to NW. Europe in 667by a monk, Francesco Cupani. It is commonly cultivated as an ornamental. Its flowers are also used as a source of an essential oil. LATHYRUS SATIVUS L. Grass pea, Chickling pea. 2n=4. Probably domesticated in W. Asia (p. 83). Primary centre in the Mediterranean area. LATHYRUS TINGITANUS L. Tangier pea. 2n=4. Mediterranean Region. Ithas a micro-centre in Morocco. Cultivated as a winter annual, also in USA. LOTUS EDULIS L. Asparagus pea,winged pea.2n= 4. Mediterranean area to Asia Minor and Syria. Occasionally cultivated for its young pods. LUPINUS ALBUS L. (syn. L. sativum Gaertn e). White lupin, Mediterranean white lupin.2n= 50. Wild in Balkan, Crete and W. Turkey. Pro- bably domesticated in the Balkans (Gladstone, 977). All cultivars have white seeds, the production of pigment being suppressed by two independent pairs of inhibitor genes. These genes must already have been selected forby farmers some 4000 years ago (Kazimierski, 960). L. albus is closely related to L u termis*. According to Kazimierski (960) both derive from L, graecura (seel. termis*). This latter species would derive from L. jugoslavicus Kazim. & Now. (2n=50) which is found in Yugoslavia. Gladstone (970)considered L. termis, L. graecus and L. jugoslavicus as synonyms of L. albus. LUPINUS ANGUSTIFOLIUS L. (syn. L. varius L., L. linifolius Roth, L. reticulatus Desv,). Narrow-leaved lupin, Blue lupin. 2n=40. Primary centre in the Mediterranean area D The present European cultivars probably derive from wild types of Palestine. Cultivated also in S. Africa and Australia as a forage.widely cultivated as an ornamental too. LUPINUS COSENTINI Guss. (syn. L. varius L., spp. varius Franco &P. Silva). Western Australian blue lupin, Sand-plain lupin, Geraldton lupin. 2n=32. Coastal Morocco and other sites in W. Mediterranean area. Introduced into W. Australia about the middle of the 9th Century and naturalized. Since 90,it has been cultivated for summer sheep food and soil improvement. Described as L, pilosus L., L, varius L. and L. digitatus Forsk. LUPINUS LUTEUS L. (European) yellow lupin.2n =(46, 48, 50), 52. Mediterranean area, where its primary centre lies. The present European cultivars derive probably from wild Palestinean plants.cultivated as a fodder crop, green manure and ornamental. Closely related to L. hispanicus Boiss. & Reut, and L. rothmaleri Klink (2n=50, 52). LUPINUS PILOSUS L. (syn. L. varius L. ssp. orientalis Franco & P. Silva). Greater blue lupin,hairy lupin. 2n=42, (50). NE. Mediterranean Region. Itmay occasionally be cultivated. It is characterized by its big seeds,the biggest of all Lupinus-species.The cultivated type has 2n=50. Distribution of Lupinus albus in the Mediterranean as a wild and cultivated plant. Hatched: cultivated (var. albus only); black: cultivated (var. albus) andnative (var. graecus)(gladstone, 976). LUPINUS TERMIS Forsk. (syn. L. graecus Boiss., L, albus ssp. albus). Egyptian lupin. 2n= Palestine and Egypt. Cultivated inegypt since ancient times and in Nigeria. The seeds contain alkaloids, which have to be removed before consumption. Closely related tol. albus*. L. termis and L. graecus may be varieties of L. albus*. MEDICAGO ARBOREA L. Cytisus u 2n=32, 48. Canary Islands, Balaeric Islands, S. Europe to Asia Minor. The 4x type is common while the 6x type is found on Esparta, Balearic Islands only. A shrub formerly used for fodder and for making

104 LEGUMINOSAE - LEGUMINOSAE 5 baskets (Lesins & Lesins, 979). MEDICAGO POLVMORPHA L. (syn. M. hispida Gaertn. and M. denticulata Willd.). Bur clover. 2n= 4 u Mediterranean area.spread world wide. Cultivated as a green manure, for pasture and as ahay crop in Australia, S. America and S. USA. MEDICAGO SATIVA L. Lucerne, (Blue) alfalfa. 2n=6, genome formula SS, 32 genome formula SSSS, 64. Transcaucasia (p. 97). Secondary centre in N. Africa, especially Algeria. MELILOTUS INFESTUS Guss. 2n=6. A plant of W. Mediterranean area, being a source of resistance to the sweet clover weevil of M. albus* and M. officinalis*. MELILOTUS MACROCARPA Coss. & Dur. 2n=6. N. Africa. Cultivated in Algeria for its large fruits used as spice. MELILOTUS SULCATUS Desf. 2n=6, (32). S. Portugal and the Mediterranean area.plants belonging to ssp. brachystachus Maire are coumarin deficient and resistant to drought,and most pests including the sweet clover weevil, Ssp. segetalis (Brot.) Maire has also been described as M, segetalis Ser. (2n=6). ORNITHOPUS COMPRESSUS L. 2n=4. Spain, Portugal and Mediterranean area. Its northernmost point of occurrence isbrittany in France.It has a high leaf and seed production. It might be useful as a breeding source for0. sativus*. ORNITHOPUS SATIVUS Brot. Serradella. 2n=4. Wild plants in NW. Portugal, N. Spain and SW. France. From here its cultivation spread over W. and N. Europe, since the beginning of the 9th Century. A green manure and fodder. Ssp. sativus (syn. 0. roseus Dufour) is native to SW. France, N. Iberian Peninsula and the Azores.Cultivated elsewhere. A related species is 0, isthmocarpus Coss. (syn. 0. sativus ssp. isthmocarpus (Cosson) Dostal) (2n=4). A Mediterranean-Atlantic species, where it grows together with 0. sativus, hybrids, described as 0. macrorrhynchus (Willk.) Klinkowski & Schwz. (syn. 0. sativus var. macrorrhynchus Willk.) are found, PISUM SATIVUM L. ssp. hortense Asch. & Graeb. Garden pea. 2n=4. Ssp. hortense is domesticated in SW. Asia (p. 97). Secondary centre in the Mediterranean area. PISUM SATIVUM ssp.joraardi (Schrank) Alef. (syn.ecotype arvense s.str., P. elatius (M. B.) Stev., P 0 jomardi Schrank, P. transcaucasicum Stankov). 2n=4. Cultivated in Egypt. Closely related to ssp. abyssinicum (p. 37) (Fouzdar & Tandon, 976). PSORALEA BITUMINOSA L. Asphalt clover. 2n=20. Mediterranean area and Canary Islands. Cul- tivated for fodder. SPARTIUM JUNCEUM L. Spanish broom, Weaver's broom. 2n=48, 52, Mediterranean area and Europe. Cultivated in France near Aspiran (Hérault). TRIFOLIUM ALEXANDRINUM L. Alexandrian clover, Egyptian clover, Berseem. 2n=6. E. Mediterranean area. Cultivated in the Near East and India, It is the oldest clover cultivated and is closely associated with agriculture in Egypt. Secondary centre in Egypt. T. alexandrinum derives from T.berytheum Boiss. (syn. T 0 alexandrinum var. berytheum) and T. salmoneum Mout.,2n=. T. alexandrinum is selfcompatible while its progenitors are self-incompatible. Self-compatibility may be a characteristic of domestication. TRIFOLIUM FRAGIFERUM L. (syn. T, neglectum Fisch &Mey)o Strawberry clover. 2n=6. Europe, Canary Islands, Madeira, N. Africa and W. Asia, Cultivated as fodder. TRIFOLIUM INCARNATUM L. Crimson clover. 2n=4. C 0 and S. Europe, Balkans and N. Africa. The cultivated type (var,sativum Due,, ssp, incarnatum) probably derives from the wild var, molinerii (Balbis exhörnern.) Syme.The latter is found in Spain. Long cultivated in Catalonia (Spain)and S. France. Spread to E, and N. Europe and later to N. America, TRIFOLIUM ISRAELITICUMD. Zoh. & Katzn. (syn. T. subterraneum L. var. telavivensis Eig). 2n =4. N, Israel.It isnot a parent of T. subterraneum ashas been suggested. Itonly has 4chromosomes while T. subterraneum has 6. Formerly it was considered as the "Israeli race" of this species. TRIFOLIUM REPENS L, var. giganteum.lodi clover, Ladino clover. 2n=32, Probably, Lodi, N, Italy. Cultivated first in N. Italy and the Netherlands (p. 57). An excellent fodder crop. TRIFOLIUM SUBTERRANEUM L. Subterranean clover, Sub clover. 2n=6, Mediterranean area, SE,and W, Europe, Caucasian Region and N. Iran. It is possible that the westward migration followed the course of clearing and cropping by man (Katznelson & Morley, 965a, 965b). Secondary centre in Australia (p. 66). Naturalized in Australia, S. Africa, and N. and S. America. T. subterraneum can be divided into three subspecies:. ssp. subterraneum (syn. T. blesense Dodart) which is the commonest taxon sympatric with the species; 2. ssp. yanninicum Katzn. & Morley, which occurs in Istria, Dalmatia, Albania, Serbia and N. Greece; 3, ssp. brachycalycinum Katzn. & Morley (syn. var. oxaloides Eig) which occurs from W. Thrace to Caspian Sea. These subspecies are almost completely intersterile (Katznelson & Morley, 965a), The existence of two closely related but more primitive species (T. batmanicum

105 .6 MEDITERRANEAN REGION Katzn. (syn. T. anatolicum Katzn.) (2n=6)in Diyarbakir Province, and T. chlorotrichum Boiss. & Balansa (2n- )in Phrygia) in Turkey and the absence of these and other close relatives elsewhere indicates the origin of T, subterraneum in Turkey. However, the greatest variation is found in Greece (Katznelson& Morley, 965a). Bailey & Francis (97) found that the isoflavone pattern oft. batmanicum closely resembles that of spp. brachycalycinum. They concluded that T. batmanicum might be the ancestor species of T. subterraneum and ssp, brachycalycinum isprobably the earliest form of subterranean clover.they postulated that ssp. subterraneum evolved later and colonized a wider range of environments. The isoflavone pattern of T,batmanicum is very similar to that of T. globosura L. (syn. T. radiosum Wahlenb., T. nidificum Griseb.) (2n=6) f However, that species belongs to another subsection. TRIFOLIUM VAVILOVII Eig. 2n=6. Israel. VICIA ARTICULATA Hörnern. One-flowered vetch 0 2n=. Mediterranean area, Asia Minor, Madeira and Canary Islands. Cultivated. VICIA BENGHALENSIS L. (syn. V. atropurpurea Desf,)o Purple vetch. 2n=2, 4. Mediterranean, area. Naturalized in the USA. Cultivated as a clover crop and green manure, and as a winter and spring forage. VICIA CALCARATA Desf. Demehi. 2n=2, 4. Sahara Oasis, where it is cultivated for its seeds.also Iran & cultivated in Tripolitania (Libya). VICIA ERVILEA (L.) Willd. Bitter vetch, Ervil. 2n=4, Primary centre in the Mediterranean area. Cultivated in Spain. A characteristic group developed in Asia Minor (p. 98). Used as forage and for grain. VICIA FABA L. (syn. Faba vulgaris Moench). Field bean, Broad bean, Horse bean, Pigeon bean, Tick bean, Windsor bean. 2n=2 (4). SW. Asia (p. 83) or Near East or Mediterranean area (Zohary, 977). Wild ancestor uncertain. See p. 83 for discussion of origin. Schultz-Motel (972) found no evidence for a supposed division of the small-seeded type into two geographical races: a long-seeded type in the W. Mediterranean area and a round-seeded type in the eastern part. So there is no reason to suppose that the broad bean originated in two separate areas. Whether V. pliniana (Trabut) Moratova found in Algeria and Morocco, is a type of V. faba or a related species is not known. VICIA NARBONENSIS L. Narbonne vetch. 2n=4. SW. Asia (p. 98). Secondary gene centre in Mediterranean area where it is cultivated. Liliaceae ALOE BARBADENSIS Mill. (syn. A. vera L.). Curacao aloe, Barbados aloe. 2n-(0), 4. Mediterranean area, S, Arabia, E. Africa, NW. India and S. China. The wild var. barbadensis of the Mediterranean area has runwild in C. America, W. Indies to Bolivia. It probably arrived there through Spain. Cultivated in W. Indies. LILIUM CANDIDUM L. Madonna lily, Bourbon lily 2n=4, Mediterranean area and SW. Asia. Cultivated especially in S. France for its flowers. These are a source of an essential oil. Itis the oldest lily of European gardens. URGINEA MARITIMA (L.) Baker, (syn. U. scilla Steinh.). Sea onion. 2n=20, (30), 40, 60. Mediterranean coast; most common in E. Algeria. Wild and cultivated plants are used for their pharmaceutical properties and in rat poison. Linaceae LINUM USITATISSIMUM L. Flax, Linseed, 2n=30, (32). For origin see p. 99. In the Mediterranean area, the oil-flax (spp. mediterraneum Vav. & Ell.) is cultivated. In Italy, hybrid forms (spp. transitorium Vav. & Ell.) of ssp. eurasiaticum and ssp. mediterraneum are found. Large-seeded types are cultivated in N. Africa, Those from Algeria are a source of Fusarium resistance. Malvaceae ALTHAEA OFFICINALIS L. Marsh mallow. 2n=42, (c.42, 40-44). Europe, E. Mediterranean area and W. Asia. Cultivated for its roots, which are a source of medicine. ALTHAEA ROSEA (L.) Cav. Garden hollyhock, Hollyhock. 2n=(26), 42, (56). Asia Minor, Balkans and Crete. Ran wild in Italy, S 0 France and S. Tyrol. Cultivated in Europe since the 6th Century, especially var, nigra hort., which has blackish purple petals used to colour wine and as medicine. Cultivated now in many types as an ornamental. Moraceae FICUS SYCOMORUS L. Sycomore fig a 2n=26. Its area of distribution can be divided into two parts:. the main part is the E, Coast of Africa from S. Africa to Sudan, where trees produce viable seed and grow wild; 2,the northern area is the Middle East and N. Africa where trees do not produce viable seed and have tovegetatively propagate. The tree was perhaps domesticated in the Middle East (p. 99). The southern boundary of the domesticated sycomores runs through Sudan (Galil et al., 976). Myrtaceae

106 LEGUMINOSAE - RESEDACEAE EUCALYPTUS CAMALDULENSIS Dehn. Longbeak eucalyptus,, 2n=22. Primary centre in Australia (p. 66). Secondary centres in the Mediterranean Region and S. America (p. 77).It was believed that the trees of this species cultivated in Israel came from S. Australia,but the leaves of the Israeli trees contain three polyphenols which have not been found in the species anywhere in Australia. EUCALYPTUS GLOBULUS Labell.Fever tree, Blue gum. 2n=20, 22, 28. SE. Tasmania. Cultivated. Secondary centre in the Mediterranean Region. MYRTUS COMMUNIS L. Myrtle. 2n=22. Mediterranean area and SW. Europe. Cultivated since ancient times for its fruits and for its medicinal properties. Oleaceae FRAXINUS ORNUS L. Flowering ash, Manna ash. 2n=46. A tree of C. and E, Mediterranean area. Cultivated on the N. coast of Sicily. OLEA CHRYSOPHYLLA Lam.Golden-leaved olive tree. 2n=. Wild over a large part of the Old World, including the Mediterranean area. It ispossibly the wild ancestor of 0. europaea*. If so, it is a synonym of 0. europaea var, sylvestris Brotero. OLEA EUROPAEA L. Olive tree. 2n=46. Mediterranean area. Primary centre in the Mediterranean Region. Its domestication started there in ancient times. Var. sylvestris Brotero includes the wild forms and the possible naturalized cultivated types. Var. europaea is the cultivated form. The main differences of var. sylvestris are spiny lower branches and small leaves and drupes. Some cultivars are developed for table olives, others for oil. See also 0. chrysophylla*. Papaveraceae PAPAVER SOMNIFERUM L. Opium poppy. 2n=2x=22, 4x=44. The cultigen ssp, somniferum derives from the wild ssp. setigerum (DC) Corb. (syn. P. setigerum DC), 2n=2x-22, 4x=44, which occurs in the Mediterranean Region from the Canary Isles eastwards. In Greece and Cyprus, tetraploid types are found,which are more ruderal than the diploid and hence spread easier (Hammer & Fritsch, 977). Schiiltze-Motel (979) stated that the poppy was domesticated in the W. Mediterranean area.the cultigen is grown for the dried latex obtained from unripe capsules, which is used in medicine and as a narcotic, for its ripe seeds, which are eaten or pressed forpoppy oil, and as an ornamental. Itoften escapes from cultivation. There are several taxonomie classifications to divide the cultigen into various subspecies on the basis of phenotypic variation. Pinaceae PINUS PINEA L. Stone pine, Pinie.2n=. S. Europe. A tree often cultivated for its edible seeds. Pistaciaceae PISTACIA LENTISCUS L. Lentisk pistache. 2n=24. Mediterranean area. A small tree cultivated for its chewing gum. PISTACIA TEREBINTHUS L. Terebinth pistache. 2n=. Mediterranean area. On the Aegean islands, a type with big fruits and large leaves was cultivated, Plantaginaceae PLANTAGO INDICA L. 2n=2. C., S, and E 0 Europe and W. Asia. Cultivated in S, France as a medicinal herb (Mansfeld, 959). PLANTAGO PSYLLIUM L. Psyllium, 2n=2. Mediterranean area, Ranunculaceae AQUILEGIA VULGARIS L. Columbine. 2n=4. S. and C. Europe (p. 58), N. Africa and temp. Asia. Cultivated widely as an ornamental, formerly also formedicinal purposes. Olea europaea (Polunin & Huxley, 972). Palmae CHAMAEROPS HUMULIS L. Dwarf palm. 2n=36. Wild in the W. Mediterranean area. Cultivated in some parts of Morocco. Often planted as an ornamental. A source of fibre (crin vegetable). NIGELLA SATIVA L. Black cumin. 2n=2. C. (p. 58) and S. Europe, N. Africa andw. Asia. Cultivated for its seed in the Mediterranean area and in the Orient. Cultivated formerly in C. Europe. Resedaceae RESEDA LUTEOLA L. Weld. 2n=24, (26, 28). C. Europe (p. 58), Mediterranean area, Iran and Afghanistan, Formerly cultivated as a source of deep yellow dye.

107 8 MEDITERRANEAN REGION RESEDA ODORATA L. Mignonette. 2n=2, (4). N. Africa. It may derive from R. phyteuma* with introgression ofr. arabica Boiss., 2n-24 and R. orientalis (Muell,-Arg.) Boiss.,2n= R. arabica is found in Africa north of the Sahara, Egypt, Palestine, Syria and upto the Persian Gulf, while R. orientalis occurs in S. Turkey, Cyprus, Lebanon, Syria and Palestine. Between 733 and 737 material was sent toparis and from there its cultivation as a perfumery plant started and spread (Abdallah & de Wit, 978). Var. neilgherrensis is grown in India (p. 78). RESEDA PHYTEUMA L. 2n=2. N. and S.of the W. and C. Mediterranean area. Said to be eaten as a vegetable in Greece. It may be the main ancestor of R. orientalis* (Abdallah & de Wit, 978). Rhamnaceae RHAMNUS CATHARTICUS* RHAMNUS FRANGULA* RHAMNUS PRINOIDES L'Hér. 2n=4. Ethiopia. Cultivated for leaves and branches which are used to flavour beverages, and for medicine (Jansen, 98). ZIZIPHUS LOTUS (L.) Lam. 2n=24. Mediterranean area. A tree cultivated in Italy, S. Spain and Egypt. It isprobably the lotophagus of the ancient peoples of Libya. Rosaceae AMYGDALUS PERSICA L. Peach, 2n=6. Primary centre in China (p. 42). Secondary centre in Italy and Spain. CRATAEGUS AZAROLUS L. (syn. C. aronica Bosc.). Azerolier^ 2n=. S 0 Europe, N, Africa and the Orient (p. 85). This shrub or small tree is often cultivated for its edible fruits. Var. aronical. is found wild on Crete. Rutaceae CITRUS AURANTIUM L. spp. bergamia (Risso & Poit.) Wight & Arn. Bergamot. 2n=8. Calabria (Italy. Primary centre probably in SE 0 Asia (p. 63). Cultivated for bergamot oil in Calabria. CITRUS LIMON (L.) Burm. Lemon. 2n=8, (36). Primary centre probably in SE. Asia (p. 63). Secondary centre in the Mediterranean Region, especially in Sicily. CITRUS SINENSIS (L.) Osbeck (syn. C, aurantium L. var. sinensis L.). Sweet orange. 2n=8, (27, 36) Primary centre probably in S u China or Cochinchina (p. 63). Secondary centres in Israel (e.g. the varieties Shamuti, Beladi, Khalili) and in Spain (e.g. the variety Valen- cia,blood orange). RUTA CHALEPENSIS L. Fringed rue. 2n=36. Mediterranean area. Cultivated there and elsewhere as a medicinal plant. RUTA GRAVEOLENS L. Common rue, Rue. 2n=72, 8, Wild in the Mediterranean area. Introduced into many tropical countries. The leaves are used as a condiment and medicinally. Scrophulariaceae DIGITALIS PURPUREA* Solanaceae ATROPA BELLADONNA L. Belladonna. 2n=72. From Spain, the Balkans, Asia Minor to India (p. 79). Cultivated in Europe, India and USA as a medicinal plant. A. martiana F.Q. is considered a hybrid of A. belladonna and A. baetica Willk. (2n=72), which is found in Spain. HYOSCYAMUS NIGER L. Black henbane. 2n=34. Mediterranean area. A medicinal plant cultivated in some countries for its alkaloids. Ulmaceae CELTIS AUSTRALIS L. (syn. C. excelsa Salisb.). Hackberry. 2n=40. Mediterranean area. Tree cultivated there as an ornamental and in Asia Minor for its edible fruits (Mansfeld, 959). ULMUS spp. Semi-cultivated by the Romans as a support for grapevines and so distributed. The leaves are used as fodder in dry summers. Umbelliferae AMMADAUCUS LEUCOTRICHUS (Coss. & Bur.). N. Africa, Cultivated there. AMMI MAJUS L. (syn. Apium ammi Crantz), Bishop's weed. 2n=22. Mediterranean area to Iran and to Switzerland and Belgium, Cultivated since the Middle Ages for its aromatic seeds and formedicinal purposes. ANETHUM GRAVEOLENS L. (syn. A. sowa Kurz.). Satapashpi, Sowa, Suwa. 2n=22. Eurasia. Cultivated in India. It has longer fruits than the Indian type (p. 79). APIUM GRAVEOLENS L. Celery. 2n=22. Cultivation started in the Mediterranean area. The wild parent A. graveolens var. silvestre Presl. (syn. var. graveolens) is cosmopolitan. Not much is known about the development of the three botanical varieties of A. graveolens: var. silvestre f. secalinum Alef. (syn. var, secalinum Alef,), Leafy celery, Smallage or Soup celery; var. rapaceum (Mill.) DC., Celeriac, Turniprooted celery or German celery; var. dulce (Mill.) Pers., Blanching celery,pascal celery or Stalk celery. 2n=6.

108 RESEDACEAE - VITADACEAE 9 CORIANDRUM SATIVUM L. Coriander. 2n=22. Mediterranean area and W, Asia. Cultivated.for its aromatic fruits. CRITHMUM MARITIMUM L. Samphire,Sea samphire, Sea fennel, Piercestone. 2n=20 t (22). Canary Islands, Madeire, coasts of Portugal to S. England and those of the Mediterranean area and Crimea. Cultivated in the USA as a kitchen herb. CUMINUM CYMINUM L. Cumin. 2n=4. Mediterranean area to Turkistan. Cultivated in SE. Europe, N. Africa, India andchinaa DAUCUS CAROTA L. Yellow carrot. 2n=8. Wild species from Afghanistan (p. 86) to the Mediterranean area. Although yellow carrots may have arisen in other areas where purple carrots were cultivated, it is thought that the true yellow carrots developed in the Mediterranean Region from crosses with the wild D. carota ssp. agg, carota (syn, ssp. maximus (Desf.) Ball). FOENICULUM VULGARE Mill. (syn.f. officinale Gaertn.). Fennel. 2n=22. Mediterranean area u Cultivated there for a long time and introduced into many other temperate countries. Var. piperitum (Ucr.) Cout. (syn,f. piperitum Acr., 2n=22)is Bitter fennel. Var. dulce (Mill.) Thell. (syn.f. dulce Mill,, 2n=22) is the Florence fennel,sweet fennel or Roman fennel. Cultivated for its blanched petioles in S 0 France and the Mediterranean area. Var. azoricum (Mill.)Thell. (syn.f.azoricum Mill.), Carosella or Italian fennel, originated in Italy. It has very broad leaf-stalk bases. MEUM ANTHAMANTICUM Jacq. Signel. 2n=22. A herb of C. and S. Europe,once cultivated in N. England for its roots. PETROSELINUM CRISPUM (Mill.) Nym. ex A.W. Hill (syn. Carum petroselinum Benth.). Parsley. 2n= 22. S, Europe. Widely cultivated there and el- sewhere. Mansfeld (959) has classified the wild and cultivated types. SIUM SISARUM L. Skirret, Chervin. 2n=20, 22. E. Asia and Mediterranean area.occasionally cultivated for its edible tuberous roots (var. sisarum). SMYRNIUM OLUSATRUM L. Alisander, Alexanders, Maceron. 2n=22, Mediterranean area, S. and W. Europe and Caucasia. Formerly much cultivated but now replaced by celery. Urticaceae SOLEIROLIA SOLEIROLII (Req,) Dandy 2n= Islands of W. Mediterranean area. Cultivated. Valerianaceae FEDIA CORNUCOPIAE Gaertn. African valerian, Valériane d'alger, 2n=32. Mediterranean area. Cultivated as a pot-herb and during famine. VALERIANA ERIOCARPA Desv. Italian corn salad. 2n=, Mediterranean area. Cultivated for salad. Verbenaceae VITEX AGNUS-CASTUS L. Chaste tree. 2n=24, 32. Mediterranean area. Cultivated inthe Old and New Worlds for various purposes. Violaceae VIOLA ODORATA L. (syn. V. officinalis Cr,). Sweet violet,sweet scented violet, Common violet. 2n=20, Europe and SW, Asia. Var. parma is cultivated in N. Italy and S. France as a source of an essential oil for perfumery. Vitadaceae VITIS VINIFERA L. Common grape, European grape. ; r ~~> i -*>*.. V / r^-h v.. >^*N ' "'..\ \ ow 0 Y. / V,j!.: ^^^ - Sj?&i0ä*ty '""tp ^r? / ) *N* jpog? / / ^ ^N \ sr ^<^^~S^Mc^k \\ \ /' i c \ I' \ i Wild grape (Vitis vinifera. var. silvestris) (Zohary & Spiegel-Roy, 975). Li A v -

109 20 MEDITERRANEAN REGION 2n=38, (40, 57, 76). The Mediterranean Region is one of the three primary centres of diversity. On p. 02 the domestication of the grape is discussed and other data are presented. The grape reached Greece and Italy c, 000 BC. and spread northwards to enter France c. 55 AD, There by introgression, it absorbed genes for adaptation to cooler and more humid climates (Rives, 975)<, There are several secondary centres, e.g. the varieties for currants in Greece, and the varieties for wine in Italy, Spain and Algeria.

110 8 African Region The African Region includes allofafrica south of thesahara. Portères (950)recognized fourmajor centresofplant domestication in Africa: a Mediterranean cradle, which formspartof themediterannean Region (Chap. 7); an Ethiopian cradle, which correspondswith theabyssinian Centre ofvavilov (928)and theethiopian Centre ofdarlington (956); an East Africancradle; and a West African cradle. Portères divided hiswestafrican cradle into Senegambian,Central Niger, Benin and Adamawa subcradles.an independent origin of agriculture in thewestafrican savannawas alsoproposed by Anderson (960)andMurdock (960), but several research workershavepresented evidence against thishypothesis (Wrigley, 960; Clark, 962; Harris, 967). Agriculture inafricanorth of thesahara istypically NearEastern inorigin. South of thesahara,agriculture is basedprimarily on nativeafrican crops. Anotable exception isthehighlands ofethiopia, wherewheat andbarley have been grown since at least thebeginning of the Christian Era. Theantiquity of native agriculture in Africa is notknown. Archaeological remainsof fingermillet (Eleusine coracana) from Ethiopia suggests that this cereal has been cultivated in Africa foratleast fivemillennia.this

111 AFRICAN REGION archaeological raceof fingermillet has lost the ability todisperse seed naturally, so cereal cultivation in Africa mustbe substantially older than 5000years (Hiluet al., 979). Remains ofpearl millet (Pennisetumamericanum)dating back tothe fifthmillennium BC.were uncovered from lakeedge settlements in Mauritania (Munson, 976). A sequence from gatheringwild grasses to growing cereals such aspearl millet and possibly sorghum (Sorghumbicolor)isobvious in these settlements. However, these crops probably reachedmauritania fully domesticated. Wild racesofpearl millet occurs inc.sahel and Highlands of C. Sahara,andwild sorghum isnativeto thesavanna regions, where these two cerealswere probably domesticated (Brunkenet al., 977;de Wet, 978). The pollen recordshows that thesahara was substantially wetter some 8000 years ago than now.peoplewith cattle,goats and sheep camped along theedgesofnumerous shallow lakes inc. Sahara,andharvested wild cereals and otherplant foods inareas that are now desert (Clark, 976). The Sahara becameprogressively drierover several subsequent millennia, and it wasprobably thesenomadicherdsmen who domesticated thenative cerealsofwest Africa as they migrated south intowhat isnow savanna (Harlanet al., 976). Acanthaceae ADHATODA SCHIMPERIANA (Höchst.) Nées. 2n= E. Africa. In Ethiopia cultivated as a hedgeplant (Jansen, 98). JUSTICIA INSULARIS T. And. 2n=. Africa. Cultivated in W. Africa for its edible leaves. Agavaceae AGAVE F0URCR0YDESLern. Henequen agave.2n= c. 40. Yucatan (p. 85). Secondary centre possibly in Africa. Cultivated for fibre. DRACAENA ARBOREA Link. (syn.d. mannii Baker). Asparagus tree,soap tree; D. fragrans (L 0) Carol; and D. smithii ex Hook.f.,Cocked hat, Cockade bush. 2n=. These four species are native to Africa. Cultivated as living fences and live sticks. SANSEVERINIA GUINEENSIS (L.) Willd. Bowstring hemp. 2n=. Africa. A fibre crop cultivated on a small scale in Mexico. SANSEVERINIA LONGIFLORA Sims. Florida bowstring hemp. 2n=. Africa. Cultivated in Trinidad, S. Florida and S. Carolina. SANSEVERINIA THYRSIFLORA Thunb. 2n=. S. Africa.Cultivated for its fibres in the tropics. SANSEVERINIA TRIFASCIATA Prain. African bowstring hemp. 2n=. Trop. W. Africa. Cultivated (often as S. guineensis*) in the tropics. Var. laurentii (De Wildem.) N.E. Brown is cultivated as an ornamental in Zaïre. Aizoaceae MESEMBRYANTHEMUM ANGULATUM Thunb. Marigold. 2n=8. S. Africa. An annual cultivated in Zaïre and the Mediterranean area as a spinach. MESEMBRYANTHEMUM CRYSTALINUM L. (syn. Cryophytum crystallinum (L.) N.E. Brown). Ice plant, Crystalline. 2n=8. S. Africa. Cultivated as salad vegetable or as a soil stabilizer. MESEMBRYANTHEMUM EDULE L. (syn. Carpobrotus edulis (L.) L. Bolus). Hottentot fig. 2n=8. S. Africa. A dune stabilizer, leaves used as forage and as a source of water. Amaranthaceae CELOSIA TRIGYNA L. 2n=8. Trop. Africa, Madagascar and Arabia. Cultivated as a vegetable in Africa. Annonaceae XYLOPIA AETHIOPICA (Dun.) A. Rich. African pepper,guinea pepper, Ethiopian pepper, Spice tree. 2n=. Trop. Africa. (Semi-)cultivated in W. Africa. Apocynaceae CARISSA GRANDIFL0RA A.DC. Natal plum. 2n=22. S. Africa. Cultivated for its fruits and as an ornamental. FUNTUMIA ELASTICA (Preuss) Stapf. Lagos silk rubber. 2n=22. W. Africa. Large plantations of this tree were established in W. Africa, after the discovery that it was a source of rubber. However, these plantations cannot compete with Hevea rubber. TABERNANTHE IBOGA Bâillon. Iboga. 2n=22. Gabon, Congo and the NW. Zaïre. Cultivated in Gabon. The roots contain several indole alkaloids. The most important is ibogaine which is a

112 ACANTHACEAE -CRUCIFERAE 23 Tabernanthe iboga (Pope, 969). stimulant and in large doses a hallucinogen. Roots of wild plants are collected which has resulted in the almost extinction of this plant in several districts in Gabon. VINCA ROSEA L. Madagascar periwinkle, Cape periwinkle. 2n=6, (32). Probably Madagascar, A herb cultivated as a medicinal plant. Bignoniaceae KIGELIA AFRICANA (Lam.) Benth. 2n=40. Sausage tree. Trop. W. Africa. Cultivated for medicine and witchcraft. Bombacaceae CEIBA PENTANDRA Gaertn. Kapok tree, Silk cotton tree. 2n=72, 80, 88.Some authors believe in an American/African origin of the kapok tree (p. 87). If America is the sole centre of origin, then the African centre is secondary. The African kapok tree is divided in the Caribbean forest type and the Caribbean savanna type. The latter type, which has a broadly spreading crown, is planted in market places 0 It is possible that this type arose from cuttings of plagiotropic branches (Zeven, 969). Burseraceae CANARIUM EDULE Hook.f. (syn. Pachylobus edulis G. Don, Dacryodes edulis (G, Don.) Lam). Bush butter tree, Native pear. 2n=. Trop, Africa. Occasionally cultivated for its edible fruits. COMMIPHORA OPOBALSAMUM Engl. Mecca myrrh tree, Harobol myrrh. 2n=. Arabia and Somalia. Formerly (llth-7th Century) cultivated in Egypt and Palestine (Mansfeld, 959). Cannaceae CANNA SPECIOSA Rose. 2n=. W. Africa. Cultivated in Sierra Leone. It is the source of African Turmeric.The tubers resemble those of Curcuma longa*. Celastraceae CATHA EDULIS Forsk. Khat, Miraa. 2n=. Ethiopia and Somalia, south tonatal and Transvaal in South Africa, and in Yemen and Saudi Arabia where it probably was introduced. Cultivated in Ethiopia and Arabia for its leaves that contain a mild narcotic.the fresh leaves and twigs are used as a stimulant particularly in Arabia, Somalia, Ethiopia and Tanzania 0 Leaves are either chewed, or a refreshing tea is brewed from them a In Ethiopia, the leaf has been used as a protection against pestilence, and especially in Arabia against bubonic plague. In Yemen,khat is an important item at birth, circumcision, and at marriages and funerals ^In Harrar, twigs of khat are placed on graves for seven days. Cleomaceae GYNANDROPSIS GYNANDRA (L.) Briq. (syn.g. pentaphylla DC.). Cat's whiskers. 2n=30, 32, 34. (Sub)trop.Africa and India u Cultivated in Africa, in the West Indies and in Malaya. Used as vegetable and as ornamental. Compositae CRASSOCEPHALUM BIAFRAE S, Moore.2n=. W. Africa. Cultivated as a vegetable. Several types are known (Terra, 967). CRASSOCEPHALUM CREPIDIOIDES (Benth.) S. 2n=40. Vegetable of Nigeria. GUIZOTIA ABYSSINICA (L.f.) Cass. Niger seed. 2n=30. Centre of diversity Ethiopia (Baagoe, 974) and spread southwards to Malawi and to India, where it has run wild. Used for oilseed, GYNURA CERNUA Benth. 2n=20. W. Africa. A herb cultivated for its leaves. LACTUCA TARAXACIFOLIA (Willd.) Schum. (syn. Sonchus taraxacifolius Willd.) ü Wild lettuce, Langue de vaches, 2n=. Trop, Africa especially in Sierra Leone, Ghana, S, Nigeria and Nile region. Cultivated in W, Africa as a vegetable and as fodder, LAUNAEA TARAXACIFOLIA (Willd.) Amin ex C a Jeffrey, Wild lettuce.2n=, Vegetable of Nigeria, SENECIO BIAFRAE Oliv. 2n=. Africa. Occasionally cultivated in W, Africa. SENECIO GABONIÇUS Oliv. 2n=, Trop. W. Africa, Occasionally cultivatedo STRUCHIUM SPARGANOPHORA (L.) 0. Ktze. Water bitterleaf. 2n=, Vegetable of Nigeria, Crassulaceae BRYOPHYLLUM PINNATUM (Lam.) Oken, Never-die, Resurrection plant,2n=, Africa. Cultivated there as a medicinal crop and elsewhere as an ornamental. Cruciferae Moore,

113 24 AFRICAN REGION BRASSICA CARINATA A.Br. Abyssinian mustard. 2n=34, genome formula BBCC 0 Unknown wild 0 Cultivated in Ethiopia as a vegetable and as an oil crop. A natural amphidiploid of Q B. nigra* andd* B. oleracea* (Uchimiya & Wildman, 978). BRASSICA JUNCEA (L.) Czern. & Coss. (syn. Sinapsis juncea L.). Sarepta mustard, Brown mustard, Leaf mustard, Indian mustard. 2n=36, genome formula AABB. Africa. However, Hemingway (976) suggested acentre of domestication in C. Asia-Himalayas with secondary centres of diversity in India, China and Caucasia. Spread to E. Europe and China. Now distributed from Europe to E. Asia. Often as a weed. It is cultivated for its oily seeds and as a condiment. This species originated from the natural amphidiploid of Ç B. campestris* and Çj B nigra* (Uchimiya& Wildman, 978). Through artificial amphiploidization of hybrids of the parental species it has been possible to introduce characteristics of both parents into Sarepta mustard. ERUCA PINNATIFIDA (Desf.) Pomel. 2n=. Sahara. Occasionally cultivated in oases as fodder. LEPIDIUM SATIVUM L. Garden cress, Common cress. 2n=6, 32. Wild type var. silvestre Thell. From Sudan area to the Himalayas. Cultivated in ancient times in Europe as a vegetable. It may have reached Europe from the Levant as a flax weed. In Africa, there are red, white and black varieties and seeds are used for medicinal purposes, oil production and as a vegetable. Cucurbitaceae CITRULLUS LANATUS (Thunb.) Mansf 0 (syn. C. vulgaris Schrad.). Water-melon. 2n=22. Tropical and subtropical Africa,The wild race is commonly eatenby animals. An edible wild race, the tsama occurs in the Kalahari Desert, where it is often the principal source of water for animals and the bushmen.food is often cooked directly in the tsama. Cultivated since ancient times in Mediterranean area and India. Now cultivated in many countries of the Old and New Worlds so that secondary centres of diversity have arisen. Var. citroides (Bailey) Mansf. is found in Sudan.Cultivated inusa (Citron, Preserving melon) and USSR. The citron - a fodder melon - is adapted to very dry areas.ithas both weedy and cultivated races. The wild form, var, colocynthoides (syn. C. colocynthoides Pang.) is characterized by its white or yellow flesh with bitter flavour or not and the cultivated form var. edulis (syn. C. edulis Pang.) has red or yellow flesh with sweet flavour (Shimotsuma, 965). COCCINIA ABYSSINICA (W. & A.) Cogn. Anchoté. 2n=. Ethiopia. Sporadic cultivation in SW. of Ethiopia for its tubers but its fruits are not eaten. The tubers and fruits of thewild plants are inedible. CUCUMEROPSIS EDULIS (Hook.f.). Cogn. 2n= W. Africa. Cultivated in gardens and on roofs. CUCUMEROPSIS MANNII Naud. Africa. Cultivated there. CUCUMIS ACULEALUS Cogn. from Ethiopia. W. and C. 2n=40. Wild perennial CUCUMIS AFRICANUS L.f. 2n=24. Wild in S. Africa, Resistant to cucumis green mottle mosaic virus and powdery mildew (Visser &de Nijs, 980) CUCUMIS ANGURIA L. 2n=24. Gherkin.The wild type is var. longipes A. Meeuse (syn. C. longipes Hook.) occurring in Ethiopia and S. Africa. Introduced to the Caribbean, where the cultigen West Indian gherkin (var, auguria) developed (Meeuse, 958). Esquinas-Alcazar (978) found similar electrophoretic patterns for enzymes of both varieties, but Puchalski et al. (979)did not.some accessions of both varieties resistant to cucumis squash mottle mosaic virus and powdery mildew (Visser & de Nijs, 980). CUCUMIS DIPSACEUS Spach. Teasel gourd. 2n=24. Africa. An ornamental. CUCUMIS FICIFOLIUS A.Rich. 2n=24, (48). A perennial from Ethiopia southwards. CUCUMIS FIGAREI Naudin. 2n=72. Wild in Nigeria. CUCUMIS HEPTADACTYLUS Naudin. 2n=48 e Perennial (Dane et al., 980). Africa. CUCUMIS MELO L u Musk-melon, Melon, Canteloupe. 2n=24.As most Cucumis species come from Africa, the species probably originates from trop. Africa, whence it spread to other regions, producing secondary gene centres in Iran (p. 89), China (p. 36), Iran and S. USSR (p. 82) (Leppik, 966). CUCUMIS METULIFERUS Naudin. African horned cucumber. 2n=24.Cultivated as an ornamental and in some parts of Africa for its fruits e This self-compatible species is asource of resistance to southern root-knot nematode, aphids and squash mosaic virus for C. sativa*. CUCUMIS MYRIOCARPUS Naudin. 2n=24. S. Africa. Wild. Closely related to C. africanus* and C. leptodermis Schweik., 2n=24 (A.P.M. de Nijs, pers, coram,, 980). Resistance to powdery mildew (Visser & de Nijs, 980). CUCUMIS ZEYHERI Sond. 2n=(24), 48 u A perennial from S. Africa (Dane et al., 980). LAGENARIA SICERARIA (Molina) Standi«(syn. L a vulgaris Ser.). Bottle gourd, White-flowered

114 CRUCIFERAE - DIOSCOREACEAE Gilg) (2n= ), L. guineënsis (G. Don.)C. Jeffrey (syn. Bryonia guineënsis G. Don., Adenopus longiflorus Benth., A. guineënsis (G. Don.) Exell, A. pynaerti De Wild.) (2n= ) and L. rufa (Gilg) C. Jeffrey (syn. Adenopus rufus (Gilg) (2n= )(Jeffrey, 962). TELFAIRIA OCCIDENTALIS Hook.f. Fluted pumpkin. 2n=. Trop. Africa. Cultivated there for its seeds. Cupressaceae JUNIPERUS PR0CERA Höchst. 2n=. E. Africa. Mainly a timber tree; also used for soil conservation and for medicinal purposes. Dioscoreaceae DIOSCOREA ABYSSINICA Höchst. 2n=40. Ethiopia, savanna of Africa. Cultivated to a limited extent as a food crop, especially in Uganda (Burkill, 939; Coursey, 967). DIOSCOREA BULBIFERA L. (syn. D. latifolia Benth.). Potato yam, Aerial yam, Bulbil-bearing yam. 2n=36, 40, 54, 60, 80, 00. Wild and cultivated in trop. Asia (p. 52) and Africa. It could have been domesticated in both regions. Lagenaria siceraria gourd, Calabash gourd. 2n=22. Widespread in Africa, now pantropic. Although an extensive variation occurs in Africa both subspecies asiatica Kob. (found in Asia) and afrikana Kob. (found in Africa) also occur in Papua New Guinea, where the penis gourd developed (Heiser, 973a, 973b). It now occurs subspontaneously and is cultivated in trop. Africa, Asia andamerica» It has been shown that gourds float for a long time and seeds remain viable u This may explain very early spread to other continents by sea currents. Remains of plant material tentatively identified as belonging to Lagenaria have been found in the Spirit Cave, Thailand and have been dated B C (Gorman, 970). Remains of the bottle gourd were found in Mexico dated BC. (Whitaker & Cutler, 97) in Peru BC., in the Egyptian tombs dated BC. (Purseglove, 968) and in China 500 AD. (Li, 969). Material found in Mexico and dated AD. appears tobe more closely related to the modern races punctatum and latifolium than it is to other races (Whitaker & Cutler, 97). Itmust be the oldest crop cultivated in the tropics (Purseglove, 968). Related species are: L. abyssinica (Hook.f.) C. Jeffrey (syn. Adenopus abyssinicus Hook.f., A. reticulatus DIOSCOREA CAYENENSIS Lam. Yellow yam, Yellow Guinea yam, Twelve-month yam, Cut-and come yam. 2n=36, 54, 60, 80,40 and aneuploids. W. Africa. Also cultivated in W. Indies. DIOSCOREA COLOCASIIFOLIA Pax. False water yam. 2n=. W. Africa. Cultivated in E. Ghana, Cameroons and Mayumbe area of Zaïre. DIOSCOREA DUMETORUM (Kunth) Pax. Bitter yam, Cluster yam. 2n=36, 40, 45, 54. Cultivated throughout Africa between 5 N and 5 S. Closely related tothe Asian D. hispida*. DIOSCOREA ELEPHANTIDES (L'Hér.) Engl. Elephant's foot.2n=. S. Africa, in the rocky, semi-deserts. Collected and eaten by Hottentots. In Europe and N. America, it is cultivated as a curiosity (Coursey, 967). The tuber can grow to 350 kg. DIOSCOREA HIRTIFLORA Benth. 2n=40. Savanna of Africa. Cultivated in N. Nigeria. DIOSCOREA LIEBRECHTSIANA De Wild. 2n= Africa. Cultivated. DIOSCOREA 0VINALA Baker. Ovinala. 2n=. Madagascar. Cultivated there. Almost replaced by D. alata* and Manihot utllisslma* (Coursey, 967). DIOSCOREA PRAEHENSILIS Benth. Bush yam, Forest yam. 2n=40, 80. W. Africa. Cultivated there. Probably ancestor ofd. rotundata*.

115 26 AFRICAN REGION EUPHORBIA DREGEANA E. Mey. 2n=. Namaqualand, S. Africa. Somethimes cultivated for rubber (Uphof, 968). EUPHORBIA KAMERUNICA Pax. Solo. 2n=. W. Africa. A tall xerophytic tree. Cultivated for the latex used for tattooing and topoison arrows (Uphof, 968). MANIHOT ESCULENTA Crantz. Cassava. 2n=36.S. and C. America (p. 70). Secondary centre in Africa. PLUKENETIA C0NOPH0RA Muell. Arg. (syn. Tetracarpidum conophorum Hutch. & Dalz.). 2n= A woody vine of trop. Africa. Cultivated as a source of oil. RICINODENDRON HEUDELOTII (Baill.) Pierre ex Pax. 2n=22. W. Africa up to Angola and Usambara Highlands (Tanzania). Fruits and seeds are used as a source of oil. Also cultivated in Cameroons. Dioscorea cayenensis and D. rotundata ( ), D. dumetorum (* )and D. bulbifera ( )(Harris, 972). DIOSCOREA ROTUNDATA Poir. White yam, White Guinea yam, Guinea yam, Ibo yam. 2n=40, 60. W. Africa. The most important yam cultivated there. Probably derived from D. praehensilis*. Widely variable. Closey related to D. cayenensis* and often included it. DIOSCOREA SANSIBARIENSIS Pax. (syn. D. macroura Harms,D. welwitschii Renole). Africa. Cultivated there. DIOSCOREA SEMPERFLORENS Illine. 2n= tivated in Congo. DI0SC0REA SOSO Jun. & Perr. 2n=. Formerly cultivated in Madagascar, but now replaced by Manihot utilissima*. DIOSCOREA ZARA Baudon. 2n=. Cultivated to some extent in C. Africa. This name is applied towhat is possibly a form of either D. sagittifolia Pax. or D. lecardii De Wild. Ehretiaceae CORDIA AFRICANA Lam. Sudan teak. 2n=. Trop. Africa, trop. Arabia. Occasionally cultivated in Ethiopia, the leaves being used for medicinal purposes and the wood for building and furniture (Jansen, 98). Euphorbiaceae BRIDELIA MICRANTHA (Höchst.) Bâillon. 2n= Trop. Africa. Cultivated as food plant of the African silk caterpillar. Cul- RICINUS COMMUNIS L. Castor bean, Castor-oil plant. 2n=20, (2). Trop. E. Africa and India. Now cultivated in most tropical countries where it runs wild in clearings, roadsides and dumpheaps. It probably originated as camp-follower, evolving into an oil plant, a drug and an ornamental (Anderson, 952). The only species of this genus. Flacourtiaceae ONCOBA ECHINATA Oliver (syn. Caloncoba echinata (Oliver) Gilg.). 2n=. Trop. W. Africa. Cultivated in C. and S. America for medicinal seed oil. Geraniaceae PELARGONIUM X ASPERUM Ehrh. ex Willd. (syn.p. radula L'Hér. var. roseum Willd., Pelargonium roseum Willd.). Rose geranium. 2n=77, 8. S. Africa. Cultivated for its geranium oil. The plant is male-sterile. Autetraploids (2n=4x= 54)are fertile. Crossed with autotetraploid P. denticulatum and backcrossed with 4xP. x asperum resulted in plants with 40-55%more oil thanp. roseum (Tamai& Tokumasu, 968). P. x asperum is probably a hybrid of P. radens x P. denticulatum* (Clifford, 958) orp. graveolens* x P. radens (Moore, 955). P. radens Moore. (2n= ) is aplant of S. Africa. If the first parentage is correct, the new oil-rich hybrids are a cross of a 4x hybrid (p. radens xp. denticulatum) with 4xP. denticulatum and of backcrossing with the 4x original hybrid. This species,p. quercifolium Ait. (2n=auto 4x=44; Oak-leaved geranium) andp. crispum* have identical zymograms for esterase, peroxidase and acid phosphatase (Tokumasu et al., 977), which points to a close relationship. PELARGONIUM CRISPUM (L.) L'Hér. ex Ait. (syn.

116 DIOSCOREACEAE - GRAMINEAE P. rigidum Willd.). 2n=2x=22. S. Africa. Cultivated for its lemon-scented oil. It varies considerably inthe wild and there are several forms. PELARGONIUM DENTICULATUM Jacq. 2n=90. S. Africa. Was cultivated as a fragrant pelargonium in Japan where it was replaced in 954 byp. roseum*. It has apine scent. PELARGONIUM GRAVEOLENS L'Hér. (syn. P. terebinthinaceum (Cav.) Small). Rose geranium.2n : = 90. S. Africa. A pelargonium with rose-scented leaves.cultivated for it oil. There are many cultivars.p. capitatum Willd., is a derivative of P. graveolens (Moore, 955). Cultivated in Algeria and Isle of Réunion. PELARGONIUM KAR00ENSE Kunth. Cultivated for its geranol. 2n=. S. Africa. PELARGONIUM ODORATISSIMUM (L.) Ait. 2n=6. Trop. Africa. Extensively cultivated for its apple-scented geranium oil. PELARGONIUM TOMENTOSUM Jacq. 2n=. S. Africa. Cultivated for its peppermint-scented oil.it crosses readily with P. graveolens*. Gramineae ACROCERAS AMPLECTANS Stapf, (syn. Panicum zizanoides Hbk. var. angustatum Stapf). 2n=. W. Africa. Cultivated in Gambia as a vegetable (Terra, 967). ACROCERAS MACRUM Stapf. Nilegrass. 2n=36. S. and E. Africa.Cultivated as a pasture grass. ANDR0P0G0N GAYANUS Kunth. Gamba, 2n=20, 40. N. Nigeria. It has been divided into var. gayanus (syn. var. genuinus Hack.), var. squamulatus (Höchst.) Stapf., var. bisquamulatus (Höchst.) Hack., and var. tridentatus (Höchst.) Hack. The second and third varieties have been used for selection. The tetraploid plants found in var. tridentatus are perhaps hybrids of A. gayanus (2n=20) in the far north of Nigeria and A.tectorura (2n=20)in the S. part of N. Nigeria. ARUNDINARIA ALPINA K. Schum. Alpine bamboo. 2n=. Kenya, Tanzania, Uganda, Sudan, Ethiopia, Rwanda and Burundi in mountain forests at an altitude of about m. Its stems are used in paper industry and as building material. AVENA ABYSSINICA Höchst. Ethiopian oats, A- byssinian oats. 2n=28, genome formula AABB. Obligate weed in wheat and barley fields above 2000 m on the Ethiopian Plateau. It resembles domesticated cereals in that it has lost the ability of natural seed dispersal. Ethiopian oats are unintentionally harvested and sown with the crop it accompanies, as a mimictic weed. A spontaneous race (A. vaviloviana (Malz.) Mordv.) also occurs in Ethiopia. Natural seed dispersal mechanisms in A. abyssinica are controlled by two independent loci that are homozygous recessive in the non-brittle, obligately weedy race. Ethiopian oats is derived from the Mediterranean tetraploid (AABB) A. barbata Pott, ex Link. (Ladizinsky, 975b; Ladizinsky & Zohary, 97). AVENA STRIGOSA Schreb. abyssinica type. A- byssinian oat. 2n=28, genome formula AsAsBB. This type has also been described asa. abyssinica Höchst. This is the non-brittle form while the semi-brittle form (vaviloviana type, A. vaviloviana Malz.) is also found in Ethiopia. The abyssinian type is harvested and threshed together with barley. Both types probably derive from introduced barbata type of A. strigosa* (Ladizinsky & Zohary, 97). BRACHIARIA BRIZANTHA (Höchst.) Stapf. Palisade grass. 2n=(x=9), 36, 54. Trop, and S. Africa, cultivated esp. insri Lanka and Brazil. BRACHIARIA DECUMBENS Stapf. Signal grass. 2n= 4x=36. Trop. Africa.Cultivated throughout the tropics for fodder. An obligate aposporous apomict. BRACHIARIA DEFLEXA (Schumach.) C.E. Hubbard. Animal fonio. 2n=20. Guinea coast to Yemen, and south to S. Africa and Botswana. Var. sativa Portères is cultivated as a cereal on the Fouta Djalon Highlands of Guinea in W. Africa. Often invades cultivated fields as a weed, and is frequently harvested as a wild cereal. In Angola, an aggressive colonizer race is a tolerated wild cereal in maize fields (de Wet, 977). This species grades morphologically into B. ramosa (L.) Stapf, a species that is cultivated as a cereal in S. India. BRACHIARIA MUTICA (Forsk.) Stapf (syn. B. purpurascens (Raddi) Henri). Para grass, Mauritius grass, Watergrass. 2n=36. Throughout tropics and subtropics. Grown as a fodder. BRACHIARIA RUZIZIENSIS Germain & C. Evrard. Congo signal grass, Ruzi grass. 2n=8. E. Africa. In Australia Kennedy ruzi grass is cultivated. Itis self-incompatible. CENCHRUS BIFLORUS Roxb. (syn. C. barbatus Schum., C. catharticus Del.) Cramcram. 2n= 30, 34, 36. Aggressive colonizer of disturbed habitats in African savanna, extending toindia. Widely harvested in W. Africa as a cereal, and important fodder in the arid savanna. CENCHRUS CILIARIS L. Buffel grass, African foxtail, Rhodesian foxtail. 2n=mainly 36. Cultivated in Australia. Obligate and facultative apomixis and sexual propagation (Bray, 972). Related to C. setigerus Vehl, Birdwood grass, 2n=34, 36, 37. CHLORIS GAYANA Kunth. Rhodes grass. 2n=20, 30,

117 28 AFRICAN REGION 40. E. Africa from Ethiopia to South Africa. Excellent natural forage. Cultivated widely as a fodder in the tropics and subtropics. CHLORIS ROXBURGHIANA Schult. 2n=20. Kenya. CYN0D0N DACTYLON (L.) Persoon. Kweek grass, Bermuda grass. 2n=8, 36. Africa and Eurasia; introduced to the New World. Several varieties are recognized (Harlan et al., 970). Selections from var. dactylon are widely planted as lawn-grasses. Coastal Bermuda grass (Burton, 947), a widely planted fodder in SE. USA reproduced vegetatively from a hybrid between var. dactylon (2n=36)and var. elegans (2n=36). Var. coursii (2n=36) isan important natural fodder in Madagascar, as is var. elegans in S. Africa. Var. aridus (2n=8) is drought-tolerant, and extends from the S. Karroo (S. Africa), across the NearEast to India. See p. 73. buted inthe tropics. Portères (976) recognizes five races of fonio. However, these races have no geographic unity and grade morphologically into one another. The most primitive is var. gracilis with two racemes and spikelets that are mostly grouped into threes along the rachis. In these traits, var. gracilis resemblesd. longiflora except that the spikelets are glabrous. The lower part of each raceme is devoid of spikelets in var. stricta. The other three varieties have usually more than two racemes per inflorescence. Var. densa is characterized by crowded spikelets. Var. rustica and var. mixta include robust plants that are late to mature. CYNOD0N INCOMPLETUS Nées. var. hirsutus (Stent) De Wet & Harlan (syn. C. bradleyi Stent). 2n= 8. S. Africa. Widely cultivated lawn-grass. CYNODON X MAGENISII Hurcombe. Magenis, Sunturf. This triploid represents a natural hybrid between C. dactylon var. dactylon (2n= 36) and C. transvaalensis (2n=8). Widely planted lawn-grass from a single hybrid clone near Johannesburg in South Africa. Digitaria exilis (- teres, 950). -)and D. iburua ( )(Por- CYN0D0N NLEMFUENSIS Vanderyst. Giant star grass. 2n=8, 36. Ethiopia, Uganda and south to Angola and Zimbabwe.An excellent natural fodder. Cultivated in trop. Africa. A cultivar grown in SW. Nigeria was derived from a cross between this species (2n=36) and C. dactylon* var. coursii (2n=36). CYNODON PLECTOSTACHYUS (Schuroach.) Pilger. Star grass. 2n=8. Ethiopia to Zaïre and Zambia. An excellent natural fodder, but cultivated in Kenya. Often confused with C. nlemfuensis*, but readily distinguished by its minute glumes. CYNODON TRANSVAALENSIS Burtt-Davy. Transvaal Bermuda, African Bermuda grass (US). 2n=8. South Africa. An excellent fine hardy lawngrass. DIGITARIA ABYSSINICA (Höchst.) Stapf. Abyssinian finger grass.2n=. Trop. Africa. Used in S. Africa to control erosion; useful fodder. DIGITARIA DECUMBENS Stent. Digit grass, Pangola (finger) grass. 2n=27. S. Africa. An excellent natural fodder. An introduction from S. Africa is grown as a fodder in the Americas as Pangola grass. DIGITARIA EXILIS (Kippist) Stapf. Fonio, Acha, Fundi. 2n=8, 36. W. Africa. Cultivated as a cereal in the W. African savanna (Portères, 955). Its closest wild relative isd. longiflora (Retz.) Persoon, which is widely distri- DIGITARIA IBURUA Stapf. Iburu, Black fonio, Hungry rice. 2n=36. W. Africa.Cultivated as a cereal by the Hausa of Nigeria between Jos and Zaria, and sporadically around Zinder in Niger, Azagive in the Ivory Coast, Kande and Atalote in Togo, and between Birni and Natitingou in Benin. It is often grown in between rows of sorghum or pearl millet, and frequently as a mixture with D. exilis*. The closest wild relative of black fonio is D. barbinodis Henr., an aggressive natural colonizer in W. African savanna. DIGITARIA PENTZII Stent. Taiwan pangola grass. 2n=(8, 27), 36, (45), 54. S. Africa. Related and perhaps identical with D. decumbens*. Source of resistance to virus diseases. DIGITARIA TRICOSTULATA (Hack.) Henr. Africa. Related tod. iburua*. DIGITARIA VALIDA Stent. Giant pangola grass. 2n=24, 30, 36.S. Africa. A source of disease resistant ford. decumbens*. Introduced in Florida and Surinam. 2n= ECHINOCHLOA COLONA (L.) Link. 2n=54. Widely distributed in the tropics and subtropics. Formerly cultivated in Egypt and Tanzania. Now cultivated as inferior cereal in India. An important wild cereal and good fodder across the dry African savanna and as fodder in N. America.

118 GRAMINEAE - GRAMINEAE 29 EHRHARTA CALYCINA Smith. Perennial veldtgrass. 2n=24, 30, 48. S. Africa. Used as asoil stabilizer inw. of USA. Cv. California veldtgrass has an open panicle and sheds itscaryopses. The new cv. Mission veldtgrass has acompact panicle and is non-shedding. ELEUSINE CORACANA (L.) Gaertn. ssp. africana (Kenn.-O'Bryan) Hilu & de Wet (syn. E. africana Kennedy-0'Bryan).Wild finger millet.2n= 36. Guinea coast of W. Africa to Ethiopia and south to the Cape Province (S. Africa). Differs from E. indica* in being tetraploid, not diploid, and in having more obviously sculptured grains (Phillips, 972). ELEUSINE CORACANA (L.) ssp. coracana. (syn. E. coracana (L.)Gaertn),. Finger millet.2n= 36. Widely cultivated as a cereal along the highlands of E. Africa from Uganda and Ethiopia to South Africa. Mehra (963) recognizes an African highland race with open inflorescence. Hilu & de Wet (976) show that the African highland race is widely cultivated on the E.African highlands and was derived under cultivation from ssp. africana. This race gave rise to an African lowland race that was introduced to India, where it evolved into a morphologically distinct cereal complex.the oldest known domesticated fingermillet occurs in the archaeological record of Ethiopia Eleusine coracana in Africa (grey) African highland race, ( ) African lowland race. south as the Transvaal of South Africa by the beginning of the Bantu Iron Age. ELEUSINE INDICA (L.) Gaertn. Goosegrass. 2n= 8. Eurasia and Africa. Introduced to the New World, where it is an aggressive weed. Jameson (970) proposed that the Indian races of finger millet were derived from E. indica while E. coracana* ssp. africana is the progenitor of domesticated African finger millets. Cytogenetic evidence refutes this hypothesis. Ssp. y - > i **= * & y \/ ^ O ^~^^~ i ~^'f '^"- ^ v ""-"^ V.V"""' -'V *^Ö> '-.- 4> / J, ; ' ^ < '*-' r ;-- 7 ''' % * ' : - y-,--"") '--, / Eleusine africana (Phillips, 972). dating back some 5000 years (Hiluet al., 979). It never spread into the W. African savanna, probably because of competition with other millets such as the fonios (Digitaria sp.). However it became widely cultivated as far y -"-H ( - > - -, ' ' ' ; * / \ * ' X- Eleusine indica (Phillips, 972). '''A (J 3

119 30 AFRICAN REGION africana (2n-36) crosses readily with both African and Indian finger millets to produce fertile hybrids (Chennaveeraiah & Hiremath, 974). ENTER0P0G0N MACROSTACHYUS Schum. Bush rye.2n= 20. E. Africa. ERAGROSTIS CURVULA (Schrad.) Nees. Weeping lovegrass, Boer lovegrass. 2n=20, 30, 40, 50, 60, 70, 80, 30-70, x=0.s. Africa. There is no relation between region of provenance and 2n. Most plants belong to type robusta. The reproduction is sexual,obligate and facultative apomixis. Used assand stabilizer. ORYZA BARTHII A. Chev. (syn. 0. breviligulata A. Chev., 0. stapfii Roshev., 0. perennis Moench. ssp. barthii (A. Chev.) A. Chev.). 2n= 24. From the Guinea coast of W. Africa across the savanna to Zambia. Growing in water, often as a weed in rice fields (Clayton, 972). It probably represents derivatives of hybrids between 0. longistaminata* and0. glaberrima*. ERAGROSTIS SUPERBA Peyr. Tickgrass. 2n=40. Africa. Cultivated there. E. ERAGROSTIS TEF (Zucc.) Trotter, (syn. E. abyssinica (Jacq.) Link.). Teff, Teffgrass. 2n= 40. The large and widely distributed genus Eragrostis includes this one domesticated cereal species. Teff is an endemic cereal crop of the Ethiopian highlands (Tadessa, 975). Widely grown ins. Africa as a fodder for livestock; introduced in USA. The wild progenitor of teff isnot certain. Its closest wild relative E. pilosa (L.) Beauv. is widely distributed across warm temperate parts of the world, and is harvested as a wild cereal in E. Africa (Rozhevicz, 928). Oryza barthii (Harlan, 973). ORYZA BRACHYANTHA A. Chev. & Roehr. (syn. 0. guineensis A. Chev.). 2n=24. From the Guinea coast of W. Africa across N. Zaire to the S. Sudan. This species usually occurs in shallow pools that dry out after the rainy season. In permanent pools it behaves as a perennial. Eragrostis tef (Portères, 950). HEMARTHRIA ALTISSIMA (Poir.)Stapf & CE. Hubb. Limpograss. 2n=8, 36, (54). SE. Africa. Domesticated in Florida, USA as a perennial, vegetatively propagated forage grass. HYPARRHENIA RUFA (Nees) Stapf, (syn. Andropogon rufus Kunth). Jaragua grass. 2n=20, 30, 36, 40. Trop. Africa. Cultivated in N. and S. America. MELINIS MINUTIFLORA Beauv. Molasses grass, (Brazilian) stink grass, Honey grass. 2n=4x =36. Africa. Cultivated as a fodder plant. Naturalized in Brazil. 0RE0BAMB0S BUCHWALDII K. Schum. mountains of trop. Africa. 2n= The ORYZA EICHINGERI A. Peter (syn. 0. latifolia Hook.f. var. collina (Trimen) Hook.f.) 2n=48. This slender species grows in damp places as a forest undergrowth and is distributed from the Ivory Coast touganda and Tanzania, and also occurs in Sri Lanka. The name 0. eichingeri was also used for types now included in 0. punctata*, and 0. schweinfurthiana non Prod. (Gopalakrishnan & Sampath, 966). ORYZA GLABERRIMA Steud. African rice. 2n=24, genome formula ABAS. An indigenous cultivated rice grown in flood plains of savanna. From Senegal to Lake Chad. It is generally accepted that this species evolved under cultivation from 0. longistaminata*, which is widely collected as a wild cereal in the African savanna. Nayar (973) postulated that African rice originated as a derivative of introduced Asiatic rice, 0. sativawhich according to him reached

120 GRAMINEAE - GRAMINEAE which are used for boats and rafts,as well as for paper-making. PANICUM BULBOSUM HBK (syn.p. maximum var. gongylodes Doe ). Texas grass. 2n=6x=54, 70, 8x=72. Distributed in Texas, Mexico,C. and S. America. Forage grass. Oryza glaberrima (Portères, 950). Egypt during the fourth century BC. There is, however, no botanical or genetic evidence to support this hypothesis. Variation is discussed by Portères (956). ORYZA LONGISTAMINATA A. Chev. & Roehr. African wild rice. 2n=24.Throughout trop. Africa including N. Transvaal of South Africa. This perennial occurs in shallow pools and along the banks of rivers. It resembles 0. sativa* (Asiatic rice)in having long ligules on the lower leaves. Oryza longistaminata (Harlan, 973). ORYZA PUNCTATA Kotschy & Steud. (syn. 0. schweinfurthiana Prod.). 2n=24. Swampy streamsides from the Guinea coast to the Sudan, south to Angola. Also in Madagascar and S. Asia. OXYTENANTHERA ABYSSINICA Munro (syn. Bambusa abyssinica Rich.). Woody bamboograss. 2n=c.60. Senegal to Ethiopia. Cultivated for its stems PANICUM COL0RATUML. Small buffalo grass.2n= 8, 36, 44, (54). S. Africa. A good fodder. Closely allied top. maximum*. Var. makarikariense Goossens, makarakeri grass, 2n=44 is a native of Zimbabwe. It is more drought tolerant than var. coloratum and cultivated in S. Africa as a forage grass. PANICUM LAETUM Kunth (syn. albidulum Steud.). Haze. 2n=. An important wild cereal and good fodder forming large stands in thesahel from Senegal to Eritrea. A potential cereal for arid regions. PANICUM MAXIMUM Jacq. Guinea grass, Panic. 2n =8, (32)- 36, (48). Trop, and S. Africa, Madagascar, Mascarene Islands and Yemen. Cultivated as a pasture grass in many warm countries. PENNISETUM AMERICANUM (L.) Leeke ssp. americanum. (syn.p. typhoides (Brum.) Stapf & Hubb., P. spicatum (L.) Koern.). Pearl millet. 2n=4. African savanna, also India and the Near East. This subspecies is recognized to include all cultivated taxa of pearlmillet grown primarily as cereals (Brunken, 977). The cultivated types commonly recognized (Stapf & Hubbard, 934) are divided among four races. Race typhoides is characterized by obovate grains and includes the primitive pearl millets from which other races were derived. Grown across the A- frican savanna and in India. Race nigritarum has grains with angular cross-section and is an important cereal from Sudan to Senegal. Race globosum has spherical grains and is grown from Upper Volta to Sudan. Race leonis is characterized by acute oblanceolate grains. It is primarily a pearl millet of Sierra Leone, but is sporadically grown also ins. Mauritania. Itswild ancestor is ssp. monodii*; aweedy type is ssp. stenostachyum*. PENNISETUM AMERICANUM (L.) Leeke ssp. monodii (Maire) Brunken (syn.p. violaceum Lam.). 2n= 4. Wild pearl millet. W. Africa, Sahel from Dakar toc. Sudan, and the mountains of thec. Sahara. It is an aggressive colonizer of man disturbed habitats, and is thewild progenitor of pearlmillet (P.americanum ssp. americanum*). Distribution suggests that pearl millet was domesticated along the C. Saharan highlands at the onset of the present dry phase in N. Africa, probably between 4000 and 5000 years ago. PENNISETUM AMERICANUM (L.) Leeke ssp. stenostachyum (Klotzsch ex A.Br. & Bouche) Brunken (syn.p. stenostachyum Klotzsch), Shibra, Weed pearl millet. 2n=4. The pearl millet gene pool includes wild (ssp. monodii*) and cultivated

121 32 AFRICAN REGION Pennisetum sect. Pennisetum americanum in Africa. subspecies americanum, 2 subspecies monodii, 3 subspecies stenostachyum, 4P. purpureum (Brunken, 977).

122 GRAMINEAE - GRAMINEAE (ssp. americanum*) kinds, as well as numerous spontaneous, weedy plants that mimic the crop in vegetative and floral morphology. Until maturation, these weeds are often difficult to distinguish from the race of pearl millet they accompany as a weed. However, their inflorescences disarticulates at maturity, and ssp. stenostachyum is spontaneous in man-disturbed habitats. Shibras originate from hybrids between cultivated races of pearl millet and subspecies monodii. PENNISETUM CLANDESTINUM Höchst, ex Chiov. Kikuyu grass. 2n=36. Trop. E. Africa.An excellent natural fodder, and widely cultivated in the tropics and subtropics, as a soil stabilizer, lawn grass and fodder. Hermaphroditic and male-sterile. PENNISETUM PURPUREUM Schumach. Napier grass. 2n=4. Trop. Africa. This species does particularly well in areas with an annual rainfall exceeding 000 mm per year. It is anaggressive natural colonizer ofwet disturbed sites. Napier grass was introduced during the 20th Century tomany parts of the world as a fodder, and has become naturalized in most of the world's wet tropics. Bana grass in Kenya is a hybrid with P. americanum*. PENNISETUM SETACEUM (Forsk.) Chiov. Fountain grass. 2n=27. African savanna. Cultivated as a fodder and ins. Africa also as an ornamental. PENNISETUM UNISETUM (Nees) Benth. Natal grass, Drakenberg silky grass.2n=. African Highlands. Cultivated in S. Africa as a forage grass PENNISETUM VULPINUM Stapf & Hubbard. Africa. Cultivated in Sudan. 2n= SACCHARUM SPONTANEUM L. Wild cane. 2n= From India (p. 75) plants with 2n=54 spread to Africa. In Uganda and adjacent Tanzania it is actually cultivated. In Africa originally used as a source of salt by burning, later it became used as hedges, for erosion control and for household. Grassl (964)suggested that the high number of chromosomes may derive from hybridization of the original S. spontaneum and an African related species e.g. Sorghum bicolor* SECALE AFRICANUM Stapf. 2n=4. E. Karoo Plateau, S. Africa. Whether this species is a Pleistocene immigrant tos. Africa or a derivative of a relatively recent introduction of seeds of S. montanum from Spain or Italy as a contaminant of wheat and barley is not known (Khush, 960). It hasthe same chromosome arrangement as S.montanum*andmust have derived from this species (Stutz, 972). Owing to its separation from the Secale-area, it adopted self-fertilization as a means of perpetuation. It has a fragile rachis and a perennial habit. SETARIA PORPHYRANTHA Stapf. 2n=. Purple pigeon grass. Zimbabwe. Perennial grass cultivated in Australia. SETARIA SPHACELATA (Schum.) Stapf & Hubbard. Setaria, Golden timothy grass. 2n=8, 36, 54. Africa. Cultivated there, Australia and elsewhere. Some cultivars have a high oxalic acid content which may result in the death of cattle. In Australia, it is often called S. anceps Stapf ex Massey, which species isclosely related. Self-incompatible. SORGHUM BICOLOR (L.) Moench ssp. bicolor (syn. vulgare Pers.). Sorghum. 2n=20. This subspecies is recognized to include the 28 cultivated taxa of Snowden (936). It is widely cultivated in Africa, was introduced to India at least 3000years ago, and has become an important crop in the New World during the last century. It is not known when sorghum was first brought into cultivation. Murdock (959) proposes that it is a W. African crop. Doggett (965) proposes that domestication took place somewhere in the NE. quadrant of Africa. Distribution indicated that sorghum must have been domesticated in the savanna zone south of the Sahara, and that the crop must have been cultivated for at least 3000years (Harlan & Stemler, 976). However, known identifiable archaeological remains of cultivated sorghum in Africa date back only to the early centuries of the Christian Era (de Wet, 978). Cultivated kinds of grain sorghums are divided among races bicolor, kafir, durra, guinea, caudatum and several intermediated races (Harlan & de Wet, 972). Race bicolor (subser. Bicoloria Snowden) is characterized by open inflorescences and long clasping glumes that usually enclose the elliptic grain at maturity. Bicolor sorghums resemble spontaneous weedy sorghums, but they lack the ability of natural seed dispersal. Members of race bicolor, particularly those far removed from their African centre of origin, probably represent relics of ancient cultivated kinds. Others may have originated as selections from derivatives of hybrids between modern cultivated races andwild members of S. bicolor. Bicolor sorghums are low yielding as cereals. They are rarely an important crop, but are grown across the range of sorghum cultivation in Africa and Asia. Some selections are grown strictly for their sweet stems, while in Africa other kinds are grown for their bitter grains that are used to flavor sorghum beer. The race is morphologically heterogeneous. It includes S. dochna and S. bicolor of the Bicoloria,S. exsertum of Guineensia, and S. splendidum and S. nervosum of the subseries Nervosum Snowden. Race kafir (subseries Caffra Snowden, excl. S. caudatum). It is characterized by more or less compact inflorescences that are often cylindrical in outline. Sessile spikelets are typically elliptic with the ripe glumes tightly clasping the usually much longer grain. The name is derived from 'kafir' the Arabic for unbeliever, referring to the Bantu who grow this race. Kafir sorghums are important staples

123 AFRICAN REGION Cultivated for its tubers. PLECTRANTHUS ESCULENTUS N.E.Br, (syn. Coleus dazo A. Chev., C. esculentus (N.E.Br.) Tayl., C. langouasiensis A. Chev.). Dazo, Kaffir potato. 2n=24. Africa. Cultivated there for edible tuber. SOLENOSTEMON OCYMOIDES Schum. & Thonn. 2n=. Trop. Africa. Cultivated as a vegetable. Leguminosae ACACIA KARROO Hayne. (syn. A. horrida Willd.). Mimosa thorn, Allthorn acacia, Sweet thorn. 2n=52, 04.S, Africa. This tree is planted as an ornamental, in hedges and as sandbinder. ACACIA NILOTICA (L.) Willd. ex Del. (syn.a. arabica (Lam.) Willd.). Babul acacia. 2n=52, 04. Trop. Africa extending to India. Cultivated there for its bark tannin. It also yields babul gum (Purseglove, 968). BAPHIA NITIDA Lodd. Camwood. 2n=44. W. Africa. Formerly cultivated as a source of red dye, now only as an ornamental. BAUHINIA ESCULENTA Burch. Gemsbuck bean, Tamany berry. 2n=. Arid savanna of S. Africa, often abundant. Seeds are excellent toeat after roasting or boiling as porridge. Young tubers make a good vegetable, and may be boiled or baked. Although not cultivated, seeds and tubers are often for sale in native markets of Botswana. A potential cultivated plant. CAJANUS CAJAN (L.) Millsp. Pigeon pea. 2n=22, 44, 66. Native to India (De, 974; van der Maesen, 980), where wild plants and related species occur. Its early introduction into Africa, its wide variation and run wild plants there confused earlier authors who indicated this area as its native region. A secondary centre of diversity developed in this region. ACACIA SENEGAL (L.) Willd. (Sudan)gum arabic. 2n=26. Dry trop. Africa extending to the Red Sea and NW. India.The gum ismainly obtained from wild and semi-cultivated trees inthe Sudan (Purseglove, 968). ALYSICARPUS GLUMACEUS (Vahl.) DC. 2n=20. U- ganda, Kenya, Tanzania and Mozambique. Cultivated there as fodder. ALYSICARPUS RUGOSUS (Willd.) DC. (syn. A. violaceus (Forsk.) Schindler). 2n=6.Dry trop. Africa. Used as a cover crop and fodder. Sometimes incl. ina. vaginalis*. ALYSICARPUS VAGINALIS DC. Alyce clover, Oneleaved clover. 2n=6, 20. Trop. Africa. Cultivated now in all tropics for soil improvement, as a cover crop and as a fodder crop. Var. nummularifolius (A. nummularifolius DC.) developed in the West Indies. It has a low spreading habit with buds located in the root crown. This makes it an ideal pasture plant (Whyte et al., 953). ARACHIS HYPOGAEA L. Groundnut, Peanut. 2n= 40. S. America (p. 72). Introduced into A- frica where it is cultivated widely. Bunting (955, 958) has described numerous varieties for trop. Africa, which points to a secondary centre in this region. ASPALATHUS CONTAMINATUS (Thunb.) Druce. Rooibos tea-bush. 2n=. S. Africa. There are two forms: the prostrate one is found on the Cape Peninsula and the erect cultivated form (A. cedarbergensis Bolus) is observed in the Cedarbergs in the Clanwilliam and Citrusdal regions. The latter is the cultivated type (Cheney& Scholts, 963). ASTRAGALUS VENOSUS (A. Rich.) Höchst. 2n=6. E. Africa. Cultivated as horse fodder. Cajanus cajan CALPURNIA AUREA (Ait.) Bentham. 2n=. Angola, S. Africa, highlands of trop. E. Africa, Ethiopia ands. India. Hedgeplant in Ethiopia. Shadetree in coffee plantations. Ornamental in S. Europe and elsewhere (Jansen, 98). CANAVALIA REGALIS Dunn. 2n=22. Probably an old African domesticate known only in cultivation. It is probably derived from C. virosa (Roxb.) Wight & Am. (2n=22), which occurs in Africa south of the Sahara and also in India (Sauer, 964).

124 LABIATAE - LEGUMINOSAE 37 CASSIA ANGUSTIFOLIA Vahl. Indian senna, Tinnevelly senna, 2n=(26), 28. Somali and Arabia. Cultivated in India for senna (laxative) (Purseglove, 968). CASSIA SENNA L. (syn. C. acutifolia Del). Alexandrian senna. 2n=. Egypt, Sudan region and Sahara. Leaves and pods are taken from wild and cultivated plants (Purseglove, 968). CLITORIA TERNATA L.Butterfly pea,kordofan pea. 2n=6. Probably from Madagascar (Lowry & Chew, 974). It is widespread inthe tropics and cultivated as a fodder and soil cover crop. In Malaysia, actinomorphic flowers with bluer anthocyanins than normal are used to colour rice cakes. That type isnot cultivated, but is conserved if noticed (Lowry & Chew, 974). CROTALARIA CANNABINA Schweinf. 2n= A fibre crop. CROTALARIA GOREENSIS Gui. & Pur. Gambia pea. 2n=6, (32). Trop. Africa. Cultivated in Queensland as green manure. CROTALARIA INTERMEDIA Kotschy. Slenderleaf crotalaria. 2n=6. Trop. Africa. Cultivated in N. America and elsewhere especially for soil improvement, and also for grazing, hay and silage. CROTALARIA SPECTABILIS Roth. (syn. C. sericea Retz., C. retzii A. Hitchc). 2n=6. Trop. Africa. A green manure cultivated in (subtropical countries. Wild in India and elsewhere. CROTALARIA USARAMOENSIS E.G. Baker f. (syn. C. zanzibarica Benth.). 2n=(4), 6, (20). E. trop. Africa. A cover crop and green manure. CYAMOPSIS SENEGALENSIS Guill. & Perr. 2n=4. Semi-arid savanna zone south of the Sahara from Senegal to Saudi Arabia.An annual herb. Probably the parental species of C. tetragonoloba* (Hymowitz, 973). Valuable fodder in Senegal. CYAMOPSIS TETRAGONOLOBA (.) Taub. (syn. C. psoralioides DC.). Cluster bean, Guar. 2n=4. Purseglove (968)suggested that its origin lies in India,but Anderson (960) stated an African domestication of this crop (see further p. 76). Cultivated in S. India. DESMODIUM SALICIFOLIUM (Poir. ex Lam.) DC.2n =20, 22. Trop. Africa. Used as green manure. DIP0G0N LIGNOSUS (L.) Verde, (syn.dolichos lignosus L., D. benthamii Meisn., D. gibbosum Thunb., Verdcourtia lignosus (L.) Wilczek). 2n=22. C. Africa. Cultivated in Africa,S. America and Australia where it has run wild (Verdcourt, 970). DOLICHOS AXILLARIS E. Mey. (syn. Macrostyloma. Sudan. axillare (E. Mey.) Verde). Cultivated in E. Africa. ERYTHRINA SENEGALENSIS DC. Coral flower.2n= 42. W. Africa. Used as a hedge plant, asan ornamental and formedicinal purposes. INDIGOFERA ARRECTA Höchst. Natal indigo.2n= 6. E. Africa. Formerly cultivated for dye, and now as a green manure. INDIGOFERA ENDECAPHYLLA Jacq. Wineleaf indigo, Creeping indigo. 2n=32, 36. Africa and Asia. Cultivated as fodder crop, usually in pastures but poisons calves and horses. INDIGOFERA TINCTORIA L. (syn. I. indica Lam., I. sumatrana Gaertn.). True indigo plant.2n= 6. Probably W. Africa (Mansfeld, 959). Cultivated as a dye plant. Also used as a green manure. LABLAB PURPUREUS (L.)Sweet (syn. Dolichos lablab L., D. purpureus L., Lablab niger Medik.). Hyacinth bean,bonavit bean, Lablab bean, Seins bean, Indian bean, Lubia bean, Egyptian bean. 2n=22, 24. Wild type (ssp. uncinatus Verde, syn. Lablab uncinatus A. Rich.) in trop. Africa from W. Africa to Sudan and Ethiopia and tos. Africa. The commonly cultivated forms belong in general to ssp. purpureus unless they have linear kidney bean-like pods. Var. purpureus of this subspecies is a distinct due toall parts of the plant being purple. From Kenya, the cultivated ssp. bengalensis (Jacq.) Verde, (syn. Dolichos bengalensis Jacq.)is reported (Verdcourt, 970). L0T0N0NIS BAINESII Baker. Miles lotononis. 2n=36. N. Transvaal and Zimbabwe. A perennial cultivated in Queensland. MACROSTYLOMA GEOCARPUM (Harms) Maréchal & Baudet (syn. Kerstingiella geocarpa Harms, Voandzeia poissonii Chev.). Geocarpa bean, Geocarpa groundnut, Ground bean, Harms seeds, Kersting's groundnut. 2n=20, 22. W. Africa. Wild plants are classified as var. tisserantii (Pellegrin)Hepper (syn. Kerstingiella tisserantii Pellegrin), 2n=20, and domesticated types as var. geocarpa, 2n=22. NE0N0T0NIA WIGHTII (Arnott) Lackey (syn. Glycine wightii (Arnott) Verde. ). 2n= ; 22, 44. Africa. A perennial relative of soya (Glycine max*). Cultivars in Australia originate from Malawi, 2n=44. PHYSOSTIGMA VENENOSUM Balf. Calabar bean, Ordeal bean. 2n=. W. Africa. Cultivated for its beans which are used as poison for ordeals. PISUM SATIVUM ssp. abyssinicum (A. Braun) Alef (syn.p. abyssinicum Braun). 2n=4. Ethiopia and Yemen. Rarely found wild. Closely related to ssp. jomardi (p. 5).

125 38 AFRICAN REGION mucronata Willd.). Wild lucerne. 2n=. Trop. Africa and SE. Asia. Cultivated as a fodder crop in Brazil and Australia. Also naturalized there. STYLOSANTHES HUMILIS H.B. & K. Townsville lucerne, Townsville stylo. 2n=20. S. Africa. Cultivated in N. Australia in pastures. Pisum savitum spp. abyssinicum (Govorov, 937). PSOPHOCARPUS GRANDIFLORUS Wilczek. 2n= Ethiopia to Uganda and Zaïre in uplands at alt. c. 750 m. Cultivated in Ethiopia (Westphal, 974; Verdcourt & Halliday, 978). Closely related top. tetragonolobus*. PSOPHOCARPUS PALUSTRIS Desv. 2n=6, 8, 20. From Senegal to Sudan. The Ethiopian cultivar Wondo Surprise (Westphal, 974) belongs to P. grandiflorus* (Verdcourt & Halliday, 978). Closely related top. tetragonolobus*. PSOPHOCARPUS SCANDENS (Endl.) Verd. 2n=8. E. Africa. Cultivated in Indonesia, Sri Lanka, India, Burma,Brazil and Jamaica for its fruits Closely related top. tetragonolobus*; it is not its direct parent (Pickersgill, 980). PSOPHOCARPUS TETRAGONOLOBUS (L.) DC. Goa bean, Asparagus bean, Winged bean, Manilla bean. 2n= 8, (20). The origin of this species is much discussed and Hymowitz & Boyd (977) concluded on 'meagre evidence' (as they call it) Papua New Guinea. However like Burkill (935), Smartt (980) advocated E. Africa. Smartt based his conclusion on the absence of related species in Asia and their presence in Africa. Among the last, there is the closest related wild P. grandiflora* found in E. Africa from Ethiopia through Uganda to Zaïre in upland areas at c. 750 m above sea level. Wild material could have been taken to Asia and domesticated there (Smartt, 980). Allparts of this protein-rich vegetable are edible. It can alsobe used as a restorative intercrop and as a cover crop and the stalks can be used as fodder. It is drought-sensitive (Hymowitz & Boyd, 977). TAMARINDUS INDICA L. Tamarind. 2n=24. The savannas of trop. Africa. Introduced to India long ago and recently to other parts of the tropics. The tree and its parts have many uses (Purseglove, 968). TEPHROSIA DENSIFLORA Hook.f. 2n=. W. Africa. Cultivated and used to stupefy fish. Closely related to T. vogelii*. TEPHROSIA VOGELII Hook.f. Vogel tephrosia.2n= 22. Trop. Africa. Cultivated for its rotenoids which are used as insecticides and piscicides. Also cultivated as a green manure and a cover crop. TERAMNUS LABIALIS (Linn.f.) Spreng. 2n=20. Asia and Africa. Cultivated in E. Africa and Australia. TERAMNUS REPENS (Taub.) Bak.f. 2n=. E. and S. Africa, and India. Cultivated in E. Africa and Australia. TRIFOLIUM SEMIPILOSUM Fres. Kenya white clover, E. African white clover. 2n=6. Kenya. Domesticated in Australia as a forage crop. VIGNA UNGUICULATA (L.) Walp. (syn. V. sinensis (L.) Savi). Cowpea,Black eye, Southern pea. 2n=22, 24. Primary centre W. and C. Africa. Probably domesticated in W. Africa. Introduced into the Indian subcontinent where ssp. sesquipedalis (L.) Verde, (syn. V. sesquipedalis (L.) Fruw., Dolichos sesquipedalis L.), Asparagus pea, Yardlong pea (2n=22, 24), and ssp. cylindrica (L.) Van Eseltine (syn.v. cylindrica Skeels., V. catjang (Burm.f.) Walp.), catjang (2n=22), developed. Ssp. sesquipedalis has a long flabby pod, and is cultivated as a snap bean while ssp. cylindrica is developed as a forage crop. It has small seeds (Faris, 965) (p. 76). SPHENOSTYLIS SCHWEINFURTHII Harms. Yam bean. 2n=. Trop. Africa. A woody plant. Cultivated for seeds and tubers. SPHENOSTYLIS STENOCARPA (Höchst, ex A. Rich.) Harms. African yam bean, Yam bean. 2n=8. W. and E. Africa. Cultivated in Central African Republic,Zaïre, Ethiopia and along the Rift Valley of E. Africa for its edible tubers and seeds. Probably domesticated independently at several places across its native range (Okigbo, 973).? f, J L / v -~!-.'- - ^ ' ~~- ^ i i /, y ' T " J \\ J0-* \, " " * * - ' - ^ *$ j r r ' - STYLOSANTHES FRUTICOSA (Retz.) Alston (syn.s. Vigna unguiculata (Harlan, 973) s < ' ^, J i ' s-'' \ I f '

126 LEGUMINOSAE - MALVACEAE VIGNA VEXILLATA (L.) A. Rich. (syn. V. capensis auctt. non (L.) Walp., V. senegalensis A. Chev.). Wild mung. 2n=22. Grows wild in trop. Asia, Africa and Australia. Its tubers are collected in the wild. Cultivated in E. Africa. VIGNA SUBTERRANEA (L.) Verde, (syn. Voandzeia subterranea (L.) DC.). Bambara groundnut, Congo coober. 2n=22. Wild in W. Africa. Distributed throughout Africa and later to the Americas and Asia. Hepper (963) described the wild type as var. spontanea (Harms) Hepper and the cultivated ones as var. subterranea Hepper. Liliaceae TUBAGHIA VIOLACEA Harv. Wild garlic, Wild knoflook. 2n=2. S. Africa. The Zulu of Natal oftenplant this species around their huts to keep snakes away, supposing that the odour repels all vermin. The leaves are cooked as a spinach and thebulb is used as a emetic love medicine (Watt & Breyer-Brandwijk, 962). Linaceae LINUM USITATISSIMUM L. Flax 2n=30, (32). For possible origin see p. 99. Ssp. indo-abyssinicum Vav. & Ell. is cultivated in Ethiopia and Eritrea. Identical types are cultivated in India and may formerly have been introduced there from Africa. Lythraceae LAWSONIA ALBA Lam. (syn.l. inermis L.). Henna, Camphire. 2n=24. A shrub from trop. Asia and Africa with several uses. Malvaceae ABELMOSCHUS ESCULENTUS (L.) Moench. (syn. Hibiscus esculentus L.). Okra, Lady's finger, Gombo. 2n= See for origin p. 77. Siemonsma (980) classified the gombo cultivars in W. Africa into the true type which is cultivated throughout the world, and the Guinean gombo. The first has 2n=c. 30 and the second 2n=c. 94. He suggested that the Guinean type, also called West African gombo, is an amphiploid of A. esculentus (true type) and A. manihot*, 2n=c. 66. GOSSYPIUM ANOMALUM Wawr. & Peyr. 2n=26,genome formula BiBi. Along S. fringe of the Sahara, in SW. Africa and Angola. Itcan be crossed with G. herbaceum* and G. arboreum*. GOSSYPIUM ARBOREUM L. Tree cotton. 2n=26, genome formula A2A2. Race soudanense. NE.and W. Africa. Hutchinson (962) suggested that it was introduced from India, but Chowdhury and Buth (970, 97) thought that it is native to Africa. They found material dated about 3000 BC. at an early Neolithic site. It was probably used as stock feed, because cotton hairs intermediate between lint hairs and the hairs of wild species were found in goat dung. The form was probably cultivated by the people of Meroë, an ancient Nubian kingdom, dated about 500 BC. GOSSYPIUM BARBADENSE L. Egyptian cotton. 2n= 52 genome formula (AADD) 2. Peru (p.76). Spread to Africa. A perennial type from S. Nigeria made a 'Green Revolution' of cotton growing in the Nile Delta possible after 820. This lead to introduction of other cottons. Only Sea Islands was successful. Vigourous and fertile hybrids of these two types occurred from which Egyptian was developed. It combines the annual habit and some of the quality of Sea Islands and some of the vigour and cropping characteristic of the perennial. It canbe grown twice a year. GOSSYPIUM HERBACEUM L. Short-staple cotton.2n= 26, genome formula A^A^. Wild G. herbaceum var. africanum (Watt) Hutch. & Chose is a perennial shrub found in the bushveldt across a belt from Mozambique to Angola and SW. Africa. Hutchinson (97) suggested a likely centre of domestication in S. Arabia and Baluchistan. Aswild G. herbaceum plants were found on the coast of Sind near Karachi, domestication may have taken place within the area of the Harappan culture at Mohenjo-Daro (Pakistan) about 2400 BC. However fragments of textile and a string found at that site and dated BC. have been identified asg. arboreum*. Hutchinson (962) suggested that cotton must have been brought from the area of present Zimbabwe by early traders tothe north.the earliest domesticants were probably selected between there and Ethiopia or Arabia. From that variety, the primitive cultivated race acerifolium was selected, which was formerly found in Ethiopia, S. Arabia and Baluchistan. Chowdhury & Buth (970, 97) found cotton seed and hairs in Egyptian Nubia with an age of about 2500 BC. They suggested that the cotton was used as sheep fodder, since notextile was found. The ancient Nubian cotton resembles G. herbaceum var. africanum and G. arboreum race soudanensis*. Race persicum*, race kuljianum* and G. arboreum race indicum* derived from race acerifolium. Either G. herbaceum reached S. America from America org. arboreum reached Peru from Asia by way of the Pacific islands to form the amphiploid G. hirsutum* and G. barbadense*. Many varieties belonging tog. hirsutum and G. barbadense are grown in Africa.G. hirsutum varieties (Upland and Cambodia) are cultivated in W. and C. Africa (race punctanum) in Congo and E. Africa,while G. barbadense (Egyptian) is found inthe Nile Valley and Delta. Var. africanum has a very simple proteinbanding pattern (Cherry et al., 970). GOSSYPIUM LONGIOCALYX Hutch. & Lee. 2n=26, ge-

127 40 AFRICAN REGION Distribution of the Old World cottons in the 3th Century: Gossypium herbaceum var. africanum (), G. herbaceum var. acerifolium (2), G. herbaceum var.persicum (3), G. herbaceum var. kuljianum (4)and G. arboreum var. indicum (5)(Hutchinson, 962) I ( J (N 0 r& r o v\nr> ^_J gl" t?\_jl. 53" <ß f } ó 4' 3! Distribution of annual cottons in the Old World in 960: Gossypium herbaceum var.persicum (3), G. herbaceum var. kuljianum (4), G. arboreum var. indicum (5), G. arboreum var. bengalense (6), G. arboreum var. sinense (7), G. herbareum var. wightianum (8), G. barbarense Egyptians (9), G. hirsutum uplands and Cambodias (0)(Hutchinson, 962) nome formula E5E5. Uganda and Tanzania. It has an entire leaf like the American G. klotzschianum*. GOSSYPIUM SOMALENSE (Guerke) Hutch. 2n=26, genome formula E 2E 2. Sudan,Somali and Kenya. A variable species. GOSSYPIUM TRIPHYLLUM (Harv. ex Harv. & Sond.) Hochr. 2n=26,genome forumuia B2B2. Desert of SW. Africa and S. Angola. It can be crossed with G. herbaceum* and G. arboreum*. HIBISCUS ACETOSELLA Welw. ex Hiern. Azedas, Red-leaved hibiscus, Bronze hibiscus. 2n=72. Genome formula AABB. Tanzania, Zaïre, Zimbabwe and Angola. Cultivated in SW. Africa as a vegetable. Introduced as H. eetveldeanus De Wild. & Dur. into Indonesia.Cultivated in the (sub)tropics as an ornamental. The A genome is almost homologous to the A genome of H. asper*. The B genome is from H. surattensis* (Wilson & Menzel, 964; Menzel & Martin, 970), which grows in W. trop. Africa. The closely related H. radiatus* comes from Asia. H. noldeae Baker f. is a spiny, inedible (primitive?) wild or weedy form of H. aceto-

128 MALVACEAE - MUSACEAE sella (Wilson & Menzel, 964). HIBISCUS ASPER Hook.f. 2n=36, genome formula AA, 72. Wild in W. and C. trop. Africa. Wild plants are collected forbast fibre. Occasionally cultivated for this purpose. Its A genome is close to the A genome of H. cannabinus*. It is one of the parents of H. sabdariffa* and H. acetosella* (Menzel & Martin, 970). The A genome is also found in the African H. meeusei Exell (2n=72), genome formula AAXX (Menzel & Martin, 97). HIBISCUS CANNABINUS L. Kenaf. 2n=36,genome formula AA. Probably (sub)trop. Africa. Also wild in Asiabut these plants might derive from naturalized plants. Its A genome is related to that of H. asper*.kenaf is the A-genome donor of H. radiatus*. HIBISCUS SABDARIFFA L. Rosella, Jamaica sorrel, Guinea sorrel, Florida cranberry, rozelle, sorrel, red sorrel, Indian sorrel, sour-sour, Queensland jelly plant, jelly okra, lemon bush. 2n=(36), 72, genome formula AAYY. Africa. Angola is apparently its primary centre of dispersal. Probably first domesticated as a dooryard orweedy plant for its seeds. Later it became a vegetable and finally a fibre crop (var. altissima Webster). It is a ruderal species of Angola, SW. Africa, Zaïre and Tanzania. Its A genome is derived from H. asper* (Menzel & Martin, 970). Its Y genome donor is not yet known. Wilson and Menzel (964) noted the relationship of roselle and H. mechowii Garcke. Var. sabdariffa is used on Jamaica to produce a sorrel drink. HIBISCUS SCHIZOPETALUS Hook.f. 2n=45. E. Africa. Used there and elsewhere for hedges and as an ornamental. Its uneven chromosome number suggests a hybrid origin, H. rosa-sinensis L., 2n=36, 46, 72, 92, c.44, 68 being one parent. Van Borssum Waalkes (966) concluded that, as H. schizopetalus was first collected in E. Africa, H. rosa-sinensis might have an African origin. However, Negroid people in general donot cultivate ornamentals and therefore a domestication of H. rosa-sinensis in Africa is unlikely. More probable is that H. schizopetalus arose in E. Africa after introduction of H. rosa-sinensis, probably from Asia. Judging from the variable number of this last species itmight have a hybrid origin too. H. x archeri W. Watson is an artificial hybrid of H. rosa-sinensis and H. schizopetalus. HIBISCUS SURATTENSIS L. 2n=36,genome formula BB, (72). Africa. It also occurs in India, SE. Asia, Indonesia and Philippines. The B donor of H. acetosella* and H. radiatus*. Marantaceae THAUMATOCOCCUS DANIELLII Benth. 2n=. W. African rainforest from Sierra Leone to Zaïre. It produces proteins which are a non-sugar sweetener. Melastomataceae SAKERSIA LAURENTIA Cogn. 2n=. Zaïre. Cultivated there for its leaves (Terra, 967). Menispermaceae CISSAMPELOS OWARIENSIS Beauv. Velvet leaf. 2n=. W. Africa. Cultivated near coast as a medicinal (Dalziel, 937). JATEORHIZA PALMATA (Lam.) Miers (syn.j. columba (Roxb.) Miers,J. miersii Oliv.). 2n=. Trop. Africa. Cultivated as a medicinal. Moraceae CHLOROPHORA EXCELSA (Welw.) Benth. Iroko.2n=. Africa. Cultivated there for its timber. FICUS OVATA Vahl. 2n=26. Africa. Ethiopia as a sacred tree. Planted ins. FICUS PALMATA Forsk. 2n=26. Ethiopia, Sudan, Egypt, Arabia, Yemen. According to Aweke (979), this species could be the ancestor of F. carica (p. 99). FICUS TILIAEFOLIA Bak. Voara. 2n=. Madagascar. Cultivated there for its fibre. Moringaceae MORINGA PEREGRINA (Forsk.) Fiori. 2n= Egypt and Somalia. Cultivated in the (sub)tropics for the seeds which are the source of bennu oil. This species is closely related to M. oleifera*. Musaceae ENSETE VENTRICOSUM (Welw.) Cheesm. (syn. E. edule (Horan) Cheesm.). Ensete, Inset, Abyssinian banana. 2n=8. Ethiopia, the mountains of Kordofan (Sudan)and the lower part of the montane forest belt of Mount Ruwenzori (border of Uganda and Zaïre). Cultivated for the flour obtained from the pseudostem and also for its fibres. It is propagated by offshoots andby seeds from cultivated types or occasionally from wild plants. The cultivated plants are grown at higher altitudes than thewild plant. Several cultivated types are recognized. Itis suggested that there are about forty types of ensete. The back of the leaf of the Koba type is red to purple. This variety has been described as var. montbeliardi D. Bois (Smeds, 955). It is suggested that ensete is one of the oldest cultivated plants of Ethiopia. MUSA cultivars of the AAA group. Banana. 2n= 33. Primary centre in the Malayan region (p. 59). Secondary centre in the uplands of E. Africa. There it has longbeen cultivated. The

129 42 AFRICAN REGION Hibiscus acetosella (Menzel & Wilson, 969). Hibiscus asper (Menzel & Wilson, 969). V^ " " f -~- K - \>-'"\,'''_ \>-'~ -4 /" / y^f -^ '..-.-" '""-N- r \--7'"y r nt tf V '''' * 7 Hibiscus noldeae (Menzel & Wilson, 969). Hibiscus meeusei (MenzelSi Wilson, 969).

130 MALVACEAE 43 Hibiscus cannabinus (Menzel & Wilson, 969). Hibiscus surattensis (Menzel & Wilson, 969). Hibiscus mechowii (Menzel & Wilson, 969).

131 AFRICAN REGION AAB plantain also occurs in Ghana and probably elsewhere in W. Africa. BORASSUS FLABELLIFER L. Lontar, Palmyra palm. 2n=36. India andmalay Archipelago. Cultivated there. Unknown wild. Probably a cultigen of the African B. aethiopum Mart. (2n- ). ELAEIS GUINEENSIS Jacq. Oil palm. 2n=32. The coastal belt from Sierra Leone to Angola. Primary centre in Africa. Large areas are covered by semi-wild palms. The oil palm was domesticated only in a few areas before the establishment of 'European' plantations and 'development' farmer's plots (Zeven, 967, 972). Large plantations are found in Africa, SE. Asia and in C. America. Phoenix dactylifera (Oudejans, 969). Phoenix species easily hybridize. This may have resulted in an increased variation. Perhaps all mentioned Phoenix* species should be included in one species. PHOENIX HUMILIS Chev. 2n=. Cameroons. There palms are wild and semi-wild. Inthe latter case, they are protected, but not planted. They are in a pre-domestication stage. PHOENIX RECLINATA Jacq. False date palm. 2n= 36. W. Africa. Some reports refer to this palm as being cultivated as a wine palm. However these may refer to Ph. humilis* (Portères, 955b). PHOENIX SYLVESTRIS Roxb. Wild date palm. 2n= 36. Pakistan, India and S. Iran. Cultivated there as a source of sugar and wine.closely related to Ph. dactylifera* andmay have been derived from it, or they may have a common progenitor. Elaeis guineensis (Zeven, 967). PHOENIXATLANTICA Chev. False date palm. 2n= 36. Africa. It closely resembles Ph. dactylifera*. Near Marrakesh in Morocco var. marocana Chev. is cultivated. It has fairly tasty fleshy fruits while those of this species are, in general,of poor quality (Meunier, 962). PHOENIX CANARIENSIS hort. (syn. Ph. jubae Christ.) 2n=36. The Canary Islands. Spread as an ornamental to N.Africa and S.France. Closely related to Ph. dactylifera* and easily hybridizes with this and other Phoenix species. PHOENIX DACTYLIFERA L. Date palm. 2n=28. Primary gene centre:probably N. Africa.One of the oldest cultivated plants and cultivated for a long time from the Atlantic to NW. India. RAPHIA HOOKERI Mann & Wendl. Wine palm. 2n=. The km wide coastal belt of W. Africa (Russell, 965). Highly valued for its wine and fibre, and therefore cultivated and cared for. In SE. Nigeria, pedlars sold germinated fruits from the Imo River banks to farmers up to 00km away. Pandanaceae PANDANUS UTLLIS Bory. 2n=. Africa. Introduced elsewhere as an ornamental, while leaves are used for various purposes. Cultivated in Mauritius for making sugar bags. Pedaliaceae CERATOTHECA SESAMOIDES Endl. Bungu. 2n=32. W. Africa. Cultivated in some northern areas.it yields leaves for soups and seeds for oil. SESAMUM ALATUM Thonn. Tacoutta. 2n=26. Trop. Africa or India. Occasionally cultivated for its seeds in W. Sudan zone. SESAMUM INDICUM L. (syn.s. Oriental sesame, Beni seed. orientalis L.). 2n=26, (52, 58).

132 MUSACEAE - RUBIACEAE 45 Unknown wild. Secondary centre in India (p. 78) and in China/Japan (p. 42). An ancient crop, much cultivated, at present, in India, China, Japan, Burma, NW. Africa, Americas and Europe. As all wild Sesamum species but one (S. prostratum Retz. (2n=32), wild in E. India) occur in Africa it is thought that its progenitor(s) are African. Spontaneous tetraploids have been observed. Nayar & Mehra (970) considered S. indicum var. malabaricum (2n=26) as a possible 'companion weed' of sesame. It may have originated from hybrids between sesame and some sympatric wild Sesamum species. SESAMUM RADIATUM Schum. & Thonn. 2n=64. Trop. Africa. Cultivated in C. and W. Africa for its oil seeds. Pe r i op oc ace ae CRYPTOSTEGIA GRANDIFLORA Br. 2n=24. Trop. Africa or India (p. 78). Occasionally cultivated for its Palay rubber and often as an ornamental. Piperaceae PIPER CLUSII DC.2n= as a spice. W. Africa. Cultivated PIPER GUINEENSE Schum. & Thonn. Guinea pepper, Ashanti pepper. 2n-. W. Africa. Cultivated there as a spice. Polygalaceae POLYGALA BüTYRACEA Heck. Cheyi, Numbuni.2n=. W. Africa. This plant probably does not exist in the wild. It is probably a relic of an ancient tropical W. African culture. However, more evidence is needed. Cultivated in W. Africa for its fibre and edible seed. / v --. \ -.-''.--0 Polygala butyracea (Portères, 950). Polygonaceae '; '~--v""" RUMEX ABYSSINICUS Jacq. Spanish rhubarb dock. 2n=. Ethiopia. Cultivated in the Congo basin for its brick-red pigment. /' Portulacaceae TALINUM CUNEIFOLIUM (Vahl) Willd. 2n^ Africa and Arabia. Cultivated in E. Africa as a vegetable. TALINUM PANICULATUM (Jacq.) Gaertn. 2n=24. Vegetable of Nigeria. TALINUM PORTULACIFOLIUM (Forsk.) Aschers.2n=. Trop. Africa and Asia. Cultivated in Africa as a vegetable. TALINUM TRIANGULÄRE Willd. 2n=48, 72. Probably C. or S. America or trop. Africa. Cultivated in Brazil (p. 78), West Indies and W. Africa as a vegetable. The cultivation in forest regions may indicate an African origin,but as species of this genus are native to Africa and to the New World further investigation is necessary. Rosaceae HAGENIA ABYSSINICA J.F. Gmel. (syn. Brayera anthelmintica Kunth). 2n-. Ethiopia. Cultivated there for its flowers used as medicine against tapeworm. Rubiaceae COFFEA ARABICA L. Arabica coffee. 2n=22, 44, (66, 88). Primary centre: SW. Ethiopia (Meyer, 969). Secondary centre: Yemen (p. 0). Traditionally the arabica coffee has only been known in cultivation. Cultivated now over large areas. Arabica coffee is the only known Coffea species being allopolyploid and self-compatible. Its parental species are not known, but closest relatives occur in C. andw. Africa (Meyer, 969). Kammacher & Capot (972) suggested that one of the genomes has a similar structure to the genome ofc. canephora*. Various botanical and agricultural varieties are known and so are many mutants. An example isthe mutant discovered on Réunion, formerly Bourbon, which became the highly productive Bourbon coffee (Meyer, 965). Icatu is a hybrid of C. arabica and C. canephora*. COFFEA CANEPHORA Pierre ex Froehner (syn. C. robusta Linden). Robusta coffee. 2n=22, 44. W. to C. (sub)trop. Africa, from Guinea and Liberia to Sudan and Uganda. It has the highest cafein content of the Coffea species (Charnier & Berthaud, 975). Self-incompatible. Icatu is a hybrid of C. arabica* and C. canephora. The greatest diversity has been described for Zaire. Before the arrival of the Europeans in Africa, it was already cultivated there. Cultivated now especially in Indonesia and, because it is used to prepare 'instant' coffee, its cultivation increased in other tropical Asian and African countries. It is a cross-fertilizer and hence very poly-

133 AFRICAN REGION morphic. This has resulted in several synonyms. 'Congusta' coffee is probably a hybrid of C. canephoraand C. congensis* althoughthe latter is considered to be aform of C. canephora. Some botanical and agricultural varieties are described, COFFEA CONGENSIS Froehner. 2n=22, (44). Congo Basin.It resemblesc. arabica*. Possibly a formof C. canephora*. 'Congusta' coffee is a hybrid product of C. congensis and C. canephora. Form 'de la Nana' is a heterogeneous population most closely resembling this species (Berthaud & Guillaumet, 978). COFFEA EUGENIOIDES S. Moore. 2n=22. Wild in the Lake Kivu area of Zaïre, W. Uganda and W. Tanzania. Cultivated there. It resembles a slender form of C. arabica*. COFFEA LIBERICA Hiern. Liberica coffee.2n= 22, 44. Guineato Angola. Cultivated to some extent in Liberia, Surinam and afew other countries. It is a cross-fertilizerand hence very polymorphic.it has been crossed with C. arabica to produce hybrids which are cultivated. This species includes the Excelsa coffee (C. excelsa Chev., syn. C. liberica var. dewevreide Wild. & Dew., C. arnoldianade Wild.). There is a possibility that in the ancestry of this species some introgression with C. canephora* has occurred (Chinnappa, 970). VANGUERIA MADAGASCARIENSIS J.F. Gmel. (syn. V. edulis Vahl.). 2n=. Trop. Africa and Madagascar. Cultivated for its edible fruits. Rutaceae ADENANDRA FRAGRANS (Sims.) Roem. & Schult. 2n=. S. Africa. Cultivated there for its leaves which are usedto decoct tea. BAROSMA BETULINA (Berg) Bartl. & Wendl.f. Buchu.2n=. SW. and S. Africa. Cultivated on a small scale in the Clanwilliam district. Leavesof wild and cultivated crops are used for their medicinal properties (Gentry, 96). CITROPSIS GILLETIANA Swing. & Kell,and other Citropsis species. Trop. Africa.Closely relatedto Citrus. They can be used as citrus rootstocks. Sapindaceae BLIGHIA SAPIDA Koenig. Akee. 2n=32. Forests of W. Africa. Cultivated in Jamaica and W. Africa. In Jamaica,it is naturalized. Sapotaceae ARGANIA SPINOSA (L.) Skeels (syn. A. sideroxylon Rom. & Schult.). Argan, Arganier. 2n=20. SW. Morocco. A wild tree,communally owned and harvested for their fruits which are a sourceof argan oil,used like olive oil (G.. Barbeau, pers. comm., 979). This tree is suitable for domestication. BUTYROSPERMUM PARKII (Don.) Kotschy (syn. B. paradoxum (Gaertn.f.) Hepper ssp.parkii (Don.) Hepper, Vitellaria paradoxa Gaertn.f.). Karité, Shea butter tree. 2n=24. Semiwild in the savannasof W. Africa, CHRYSOPHYLLUM AFRICANUM A. DC. African star apple. 2n=26. Trop. Africa. Cultivated for its fruits. Solanaceae SOLANUM ACULEASTRUM Dunal. 2n=24. Trop. Africa. This non-tuberous plant is used for hedges. SOLANUM AETHIOPICUM L. 2n=24. Trop. Africa. This non-tuberous plant is cultivated for its edible leaves and fruits. SOLANUM ANOMALUM Thonn. Children's tomato. 2n =. Trop. Africa. This non-tuberous plant is sometimes cultivated for its red berries used as condiment. SOLANUM BURBANKII Bitter. Wonderberry, Msoba. 2n=6x=72. Probably derived from S. African msoba (Heiser, 969). It is apparently not a hybrid of S. sarrachoides (syn. S. villosum) (2n= ) and S. melanocerasura Allioni (syn. S. guineense Lam. non L.), Garden Shuckleberry (2n=72), but a contaminant. SOLANUM DUPLOSINUATUM Klotsch.2n=. Trop, ands. Africa. Cultivated for its edible fruits and leaves. SOLANUM GILO Raddi. 2n=24. Vegetable of Nigeria. SOLANUM INCANUM L. 2n=24. Africa. Occasionally cultivated. Hybridisation with S. melongena* succeeded only whens. incanum was takenas mother. The two speciesare closely related (Baksh, 979). SOLANUM MACROCARPON L. African eggplant. 2n=36. Mascarene Islands. Cultivated for its leaves and fruits (Ghana, Togo, Benin and Nigeria). The cultivartype is described as var.calvum Bitter. Its triploid number of chromosomes may point to a hybrid origin. SOLANUM NIGRUM L. Black nightshade. 2n=(x=2), 24, (36, 40, 48), 72 genome formula AAAASS, (96, 44). Native regionnot known. The 6x is amphidiploid of S. americanum Mill., 2n=2x=24 ands. villosum Mill., 2n=4x=48, genome formula SSXX. The Sgenome derives from S. sarrachoides Sandtn., 2n=24 (Edmonds & Glidewel, 977). The hybrids. nigrum (2n=6x) x S. sarrachoides is named S. x procurrens Leslie (2n=4x=48). It is sterile.

134 ? J fï'^% v \\ fïï IS" yb' tv i/h 0 RUBIACEAE - ZINGIBERACEAE 47 T? /& V f vc? ^. V Aframomum melegueta (van Harten, 970). SOLANUM NODIFLORUM Jacq. 2n=24, 72. The Sahara and Nigeria.Cultivated for its leaves. May have runwild elsewhere. SOLANUM OLIVARE Paill. 2n= Ivory Coast, Benin and Congo. Cultivated in SOLANUM ROTUNDIFOLIUM Moric. ex Dun. (syn. S. nelsoni Dun.). Hausa potato. 2n=. Believed to come from Ethiopia. Spread to W. Africa and other parts of Africa. Sterculiaceae COLA ACUMINATA (P. Brenan) Schott. & Endl. Abata kola. 2n=40. Nigeria to W. Gabon. Spread to Zaïre and Angola, to the West Indies and elsewhere. Cultivated esp.inw. Nigeria,but is second in importance toc. nitida*. COLA ANOMELA K. Schum. Bamenda kola.2n= Cameroon, esp. in Bamenda. Cultivated there. COLA NITIDA (Vent.) Schott & Endl. Gbanja kola. 2n=40. Sierra Leone to Benin, with its highest frequency in the forest area of Ivory Coast and Ghana. The genus Cola has its primary centre inw. Africa (van Eijnatten, 969, 970). Fruits were taken to the Caribbean, where this kola already grew in 630. Introduced to other tropical countries. This is the main kola of commerce. Subspecies refer to fruit colour, but this may be caused by some genes conditioning these colours. COLA VERTICILLATA (Thonn.) Stapf ex Chev. Owe kola. 2n=. From Ivory Coast to lower Congo. Often found as stray individuals in plantings^ ofc. nitida*.on the Mambilla Plateau in N. Nigeria, it is the only kola found (van Eijnatten, 970). for fuel and as ornamental. Tiliaceae C0RCH0RUS TRILOCULARE L. Al Moulinouquia. 2n= 4. Senegal to India. Sometimes cultivated as a vegetable, e.g. near Timbuktu (Mali) (Uphof, 968). Verbenaceae LIPPIA ADOENSIS Höchst. Gambian teabush. 2n=. Zaïre. A pot-herb cultivated there. In W. Africa it is used as a tea substitute. VITEX CIENKOWSKII Kotschy & Peye. 2n=32. Trop. Africa. A tree planted on compounds or semicultivated for its edible fruits. Zingiberaceae AFRAMOMUM C0RR0RIMA (Braun) Jansen. Korarima. 2n=. Ethiopia. Also cultivated there and elsewhere as a condiment and for medicine (Jansen, 98). AFRAMOMUMMELEGUETA (Rose.) K. Schum. Melegueta pepper. 2n=. W. African coastal belt from Guinea to Angola, including Fernando Po and San Thome (van Harten, 970). Probably not cultivated in W. Africa, After its introduction into S. America, cultivated in Surinam and Guyana. It is the historically known 'grainsof paradise', giving its name to thewest African Pepper Coast, Grain Coast or Malagueta Coast. Tamaricaceae TAMARIX ARTICULATA Vahl. 2n=. The Sahara, Arabia and Iran. Great numbers are found ins. Morocco and Mauritania. Cultivated as a windbreak for orange cultivation, as a sandbinder,

135 9 European-Siberian Region TheEuropean-Siberian Region was not indicated by Vavilov. Darlington (956) was the first torefer to Europe as aregion of origin ofcrop plants. Zhukovskij (968)recognized it as a megacentreofdiversity of arelatively small importance. Agriculture reached theregion from the NearEastern Regionand arrived in NW.Europe about 4000 BC. Important cropshavebeen developed inthis region including fruit-trees, grasses,brassica sp.,cannabis sativa,cichorium sp., Digitariasanguinalis, Fragaria sp., Lactuca sativa, Humulus lupulus, Medicago sp., Ribes sp., Rubus sp.andtrifolium sp. Alliacéae ALLIUM AMPELOPRASUM L. Levant garlic. Perennial sweet leek. 2n=6, 24, 32, genome formula AAA'A", 40, (48, genome formula AAA'A^'A", AABBBB), (56). Europe, Asia Minor, Caucasia to Iran and N. Africa. Cultivated in S. France and around Nuremberg, Germany for its bulbs (Kuckuck, 962). Some cultivation also in Kashmir (p. 70)and Iran (p. 87). ALLIUM SCORODOPRASUM L. Giant garlic. 2n= B, 24, 32,38 + IB, B, B. C. and S. Europe and Asia Minor. Tutin et al. (976) describe 4 subspecies: ssp. rotundum (L.)Stearn (syn. A. rotundum L.), 2n=6 +0-2B, 32, 38 + IB, B, 48 + IB, ssp. waldsteinii (G. Don) Stearn (syn. A. waldsteinii G. Don), 2n=6, 32, 40, 48, ssp. jajlae (Vved.) Stearn (syn. A. jajlae Vved.), 2n= and ssp. scorodoprasum, 2n=6, 24. The last is probably derived from ssp.rotundum and its wide and scattered distribution are probably partly due to former cultivation as a culinary plant. According to Kuckuck (962), it is still cultivated in USSR. Araceae ACORUS CALAMUS L. Sweet flag,sweet root, Calamus. 2n=8, 24, 36, (44, 45, 48, 54). N. Europe, temperate Asia and E. North America. Used as a medicinal plant, as an ornamental and for the root that is used for various purposes such as preparation of oil. Widely cultivated now, but roots of wild plants are still collected and used. Aristolochiaceae ARIST0L0CHIA CLEMATIS L. 2n=4. Probably E. and S. Europe. Formerly cultivated as a medicinal herb in most of Europe and now naturalized. Asclepiadaceae

136 ALLIACEAE - COMPOSITAE 49 CYNANCHUM VINCETOXICUM (L.) Pers. Swallowswort. 2n-22. Europe to Himalayas and Altai, and in N. Africa. A perennial herb cultivated formerly in gardens as.a medicinal plant. Berberidaceae BERBERIS VULGARIS L. European berberry. 2n=28. Most of Europe and Caucasia. Difficult to assess its territory because it was formerly planted for its edible berries and now as an ornamental. Wood and bark were used to produce a yellow dye. It is an intermediate host of stem rust (Puccinia graminis Pers.) and has therefore been eradicated in many parts. Boraginaceae LITHOSPERMUM OFFICINALE L. Gromwell. 2n=28. Spp. officinale throughout Europe, W. Asia, Caucasia and Iran. Formerly cultivated in Bohemia for preparing Bohemian or Croatian tea. Spp. erythrorhizon is cultivated in China and Japan (p. 34). Campanulaceae CAMPANULA RAPUNCULUS L. Rampion, Ramps. 2n=20, 02. Europe, N. Africa, SW. Asia and Siberia. Cultivated in the Middle Ages for its fleshy roots. Cannabidaceae CANNABIS SATIVA L. Hemp. 2n=20. The wild form is found inc. Asia. It is marked by a horseshoe-shaped scar at the base of the achene. C. sativa isone of the earliest cultivated crops. It had reached Chinaby 2500 BC. Cultivated for its fibre and for its seeds, for food or as a source of hemp seed oil. 'C. ruderalis Janisch' is a weedy non-toxic type. The Indian type, 'C. indica' (p. 7) is cultivated as a source of narcotics. Special cultivar groups have been developed for different purposes. Exceptionally tall types are found in NE. China (p. 33). HUMULUS LUPULUS L. Hop. 2n=2x=20 with X-Y sex chromosome system. Var. lupulus is cultivated in Europe, Asia and N. America. It has been domesticated in Europe and its present distribution probably reflects dispersion by man. Hop is grown for its cones (female inflorescences) which are used to flavour beer. A perennial mainly propagated by rhizomes. Var. neomexicanus, var. pubescens and var. lupuloides occur in N. America (p. 99), while var. cordifolius is found in Japan and China (p. 34) (Small, 978a). Caryophyllaceae SAPONARIA OFFICINALIS L. Soapwort, Soaproot. 2n=28. Europe and Asia. Occasionally cultivated in Germany (Mansfeld, 959). SPERGULA ARVENSIS L. (syn. Spergularia arvensis Cambess.). Corn spurrey. 2n~8. Europe. Var. sativa (Boenningh.) Mert. & Koch (syn.s. sativa Boenningh.). Cultivated as a fodder crop or as a green manure. Var. arvensis is a widespread weed, while var.maxima (Weihe) Mert. & Koch, is aweed in flax fields. Chenopodiaceae ATRIPLEX HORTENSIS L. Mountain spinach, Garden orach. 2n=8. Wild in temperate Europe and Asia. Formerly cultivated in Europe as a vegetable. BETA VULGARIS L. var. rapa. Fodder beet.2n= 8. Distribution of the wild type is given on p. 04. Probably developed in the Netherlands, perhaps from types introduced from Spain. Secondary centre in Region 5 (p. 8). Spread to Germany and elsewhere. Itmay have played a role in the development of sugar-beet, var. saccharifera (syn. var. altissima). Sugar-beet probably developed in Silesia by hybridization of an old garden form and fodder beet. The land variety "Weisser schlesischer Zückerrübe" is the parent of all sugar-beet varieties. Fuel beets are sugar-beets suitable for alcohol productionbut not for sugar extraction. CHENOPODIUM ALBUM L. Goosefoot,Fat hen, Lambsquarters. 2n=8, 36, 54. Probably cultivated in Europe inneolithic times.now it is a weed. CHENOPODIUM BONUS-HENRICUS L. (syn. Ch. esculentus Salisb.). Allgood, Good King Henry. 2n= 36. Native to the temperate Old World. Formerly cultivated as a pot-herb. CHENOPODIUM F0LI0SUM Aschers. 2n=8. Europe and the Orient. Formerly cultivated as a vegetable (Uphof, 968). Compositae ANTHEMIS TINCTORIA L. (syn. Cota tinctoria (L.) Gay). Dyer's chamomile, Golden chamomile.2n= 8. Europe and W. Asia. Cultivated as a dye plant. ARCTIUM LAPPA L. (syn. Lappa arctium Gaertn.). Great burdoc, Cocklebur. 2n=32, 36. Europe and Asia. Cultivated in Europe as a medicinal plant, and also in China and Japan (p. 34). ARTEMISIA ABROTANUM L. Southern wood. 2n=8. S. Europe and temp. Asia. Cultivated as a medicinal crop for flowers and as an ornamental. ARTEMISIA ABSINTHIUM L. Absinthe. 2n=8. Europe, S. Siberia, Kashmir and Mediterranean area. Cultivated in S. Europe, N. Africa and USA for the production of absinthe. ARTEMISIA LAXA Fritsch. 2n=8. C. and S. Europe. Cultivated.

137 EUROPEAN SIBERIAN REGION ARTEMISIA MARITIMA L. 2n=8, 36, 54. Europe to Mongolia. Cultivated as a medicinal crop. ARTEMISIA VULGARIS L. Mugwort. 2n=6, 8. Temp. N. Hemisphere. Cultivated in Indonesia and elsewhere for several uses. CHAMAEMELUM NOBILE (L.) All. (syn. Anthémis nobilis L.). Noble chamomile. 2n=8. S. and W. Europe.Cultivated as a medicinal and as an ornamental. CHAMOMILLA RECRUTICA (L.) Rauschert (syn. Matricaria chamomilla L.). Chamomile, German chamomile. 2n=8. Europe, Iran and Afghanistan. Cultivated in Europe as a medicinal and as a source of an essential oil used for flavouring and perfumery. A substitute of Chamaemelum nobile*. CICHORIUM INTYBUSL. Chicory, Succory, Brussels witloof, Sugar-loaf chicory. 2n=8. Europe, Siberia, N. Africa and the NearEast to Iran, Baluchistan and Lake Baikal. Wild type (var. intybus) was used as a salad and for medicinal purposes. Var. sativum Lam. & DC. is cultivated in Europe and elsewhere to produce a coffee substitute while var. foliosum Hegi, the Brussels witloof, was first developed around Brussels since c. 830 from var. sativum, whose young leaves had already been traded as lettuce since c. 800 and were known as barba de capucin (Moens, 974). HELIANTHUS ANNUUS L. Sunflower. 2n=34.Wild in N. America (p. 200). Secondary centre in USSR. Domesticated and cultivated in N. America. Largeheaded forms introduced in Europe. LACTUCA QUERCINA L. 2n=8. Europe, esp. in Germany, France to USSR and the Balkans. A biennal. Sometimes cultivated near Clermont- Ferrand (France) for its narcotic properties (Uphof, 968). LACTUCA SATIVA L. Lettuce. 2n=8. Primary centre: the Middle East. Lettuce derives from L. serriola L., prickly lettuce. This species occurs in S. andc. Europe to Denmark, Caucasia, Transcaucasia, Iran, Iraq, Syria, Saudi Arabia, Siberia to Altai, and N. Africa to the Canary Islands. However, Lindqvist (960) believed that lettuce probably derives by hybridization of other Lactuca species including L. saligna L. and that L. serriola arose from the same or subsequent hybridization. L. serriola is now a weed. L. saligna like L. serriola has its main distribution centre round the Mediterranean Sea (Lindqvist, 960). The first record of lettuce dates from 2500 BC.; a long-leaved form was depicted in the Egyptian tombs. The present marked variation of lettuce is probably a product of hybridization with L. serriola, but may also have been induced by some natural mutation (Whitaker, 969). Var. asparagina Bailey (syn.l. angustana Vilm., L. sativa var. angustana Irish), Asparagus lettuce of Celtuce, forms asingle thickened straight stem 90 cm ormore long, which is eaten as salad when young. L. saligna from Israel is a source of resistance to downy mildew, Bremia lactucae Reg. TANACETUM VULGARE L. (syn. Chrysanthemum vulgare (L.) Bernh. non (Lam.) Gaterau, C. tanacetum Karsch. non Vis. incl. T. audibertii (Req.) DC). Common tansy. 2n=8. Almost throughout Europe. Cultivated as a pot-herb, medicinal and ornamental. TARAXACUM HYBERNUM Steven.Krim sagiz. 2n=32, 40. Italy, Balkans, Asia Minor, Syria and Crimea. Cultivated inussr as a rubber crop. TARAXACUM OFFICINALE Weber. Dandelion, Lionstooth, Milk-gowan, Puffball. 2n=8, 24 (and others). Europe and W. Asia. In France and elsewhere, improved varieties are cultivated. These varieties "Pissenlit àcoeur plein amélioré" and "Pissenlit vert de Montmagny" differ from wild plants (pissenlit ordinaire) as they have lessbitter leaves. Young etiolated leaves ofwild plants covered by molehills are collected as dandelion salad. Crassulaceae SEDUM REFLEXUM L. Jenny stonecrop. 2n=34, c. 56, 68,c. 2.S. Europe. Cultivated in W. and C. Europe and used to flavour soup and salad. SEMPERVIVUM TECT0RUM L. houseleek. 2n=(36), 72. a medicinal plant. Cruciferae Hen-and-Chickens, Roof Europe. Cultivated as ARMORACIA RUSTICANA (Lam.) Gaertner. Mey & Schreb., (syn. Cochlearia armoracia L.). Horseradish. 2n=28, 32. Finland, to Poland, the Caspian Sea and the deserts of Cuman and in Turkey. Primary centre in temperate E. Europe (Counter 8s Rhodes, 969). Cultivated as condiment and hence naturalized. BARBAREA PRAECOX R.Br. (syn. B. verna Asch.). Scurvy grass, Winter-cress, Upland cress.2n= 6. Europe. Cultivated as a vegetable. BARBAREA VULGARIS R.Br. Yellow rocket, Common winter-cress, Upland cress. 2n=6. Temp. Europe, Asia and N. Africa. Spread throughout the world. Cultivated as a pot-herb. BRASSICA CAMPESTRIS L.Turnip group. 2n=20, genome formula AA. The wild form ssp. sylvestris (L.) Jancken grows as a weed and ruderal in most of Europe, Asia and N. Africa. The various oily (ssp. oleifera (Metzg.) Sinsk., oil-seed turnip) and fodder (ssp. rapifera (Metzg.) Sinsk., "stubble turnip", Dutch turnip) cultivars have developed independently. There are three main groups: the Asian (p.

138 COMPOSITAE - GRAMINEAE 5 35), the Mediterranean and thewest European. The turnip-rape, var. oleifera (Metzg.) Sinsk., possibly developed in Belgium. Leafy types of turnip are cultivated especially in Finland. The A genome is also found in the diploid B. chinensis* and the diploid B. japonica*. This genome is related to the Ad genome of B. adpressa Boiss., the F genome of B. fruticulosa Cyril, and the D or T genome of B. tournefortii Gouan (Mizushima, 969). B. campestris is one of the parents of B. juncea* and B. napus*, and also of the artificially made B. napocampestris (2n=58, genome formula A A AAC C ). BRASSICA NAPOBRASSICA (L.) Mill. (syn. B. napus L. var. napobrassica (L.) Rchb.). Rutabaga, Swedish turnip. 2n=38, Unknown wild. Primary gene centre in the Mediterranean area (p. 06). Secondary gene centre in Europe. Probably a derivative of B. oleracea* x B. napus*. The roots are more elongated and oval and larger than those of turnip. They are eaten as a vegetable. BRASSICA NIGRA (L.) Koch.Black mustard. 2n= 6, genome formula BB. Europe, especially in C. and S. parts. However, Hemingway (976) suggested a centre of domestication in Asia Minor or Iran. Cultivated since ancient times. Seeds are pressed for black mustard seedoil. The B genome is related to the F genome of B. fruticulosa (Mizushima, 969). Black mustard is one of the parents of B. juncea* and B. carinata*. An artificial amphiploid of B. tournefortii* and this species is called B. amarifolia (2n- 36), genome formula TTBB or DDBB). BRASSICA OLERACEA L. Wild kale. 2n=8, genome formula CC. Atlantic coast of Europe. Occurrence on Heligoland is probably spontaneous. It may have already been present there in medieval times. The wild kales of Atlantic Europe are closely related to the wild kales of the Mediterranean (seeb. oleracea, p. 06). BRASSICA OLERACEA L. var. gemmifera DC. Brussels sprouts. 2n=8, genome formula CC. Developed in Belgium probably from var. ramosa. COCHLEARIA OFFICINALIS L. Spoonwort, Scorbute grass, Scurvy grass. 2n=(4), 24,genome formula AgAgAgAg, (28, 36). N. and W. Europe.Cultivated formerly as a medicinal plant. CRAMBE MARITIMA L. Sea kale. 2n=60. Sea coast of Europe. Cultivated in England as a vegetable. HESPERIS MATRONALIS L. Damask. 2n=24. C. and S. Europe. Cultivated for its seedswhich are a source of oil and as an ornamental. Escapes are common. NASTURTIUM OFFICINALIS R. Br. (syn. Rorippa nasturtium-aquaticum (L.) Hayek). Watercress. 2n=2x=32, (28, 64). W. Asia and S. Europe and Great Britain, where it is cultivated. The leafy stems are eaten as salad. It is also cooked as a vegetable. InNew Zealand,it is a seriousweed of rivers. The almost sterile hybrid (N. x sterile (Shaw) Oefel, 2n=3x=48) of watercress and N. microphyllum (Boenn.) Rchb., 2n=4x=64is also cultivated for salad (Purseglove, 968). It is vegetatively propagated. Watercress andn. x sterile have both runwild in Florida, USA. Cucurbitaceae BRYONIA ALBA L.White bryony. 2n=20. C. Europe, USSR, the Balkans and N. Iran.Cultivated formerly as a medicinal plant. BRYONIA CRETICA L. Red berry bryony. 2n=20. S., SC. and W. Europe togreat Britain and N. Africa. Cultivated formerly as a medicinal crop. Spp. cretica is found in the Aegean region, spp. dioica (Jacq.)Tutin (syn. B. dioica Jacq.) has a wide distribution, while spp. acuta Desf.) Tutin (B. acuta Desf.) is found in Tunesia and Libya. Cyperaceae CAREX ARENARIA L. (syn. C. spadicea Gilib.). 2n=58, Europe, especially the littoral areas. Cultivated as a soil stabilizer. SCIRPUS LACUSTRIS L. (syn.s. validus Vahl.). Great bulbrush. 2n=(38, 40), 42. A world wide distribution. Cultivated in the Netherlands and Germany, to promote land reclamation and improve irapoldered land. In Germany cultivated to clean polluted water and so it is expected that the planting will increase and better varieties of this plant will be bred. Its culms contain 80%air taken from the atmosphere. They absorb air pollutant gases, sodium, phosphorus, zinc and copper. Gramineae AGROPYRON CANINUM P.B. (syn. Roegneria canina (L.) Nevski). 2n=28, genome formula S'S'H'H'. Cultivated in theussr. AGROPYRON CRISTATUM L. Gaertn. Crested wheatgrass. 2n=4 mainly, 28, (42). Europe and Asia. Introduced into N. America. Cultivated there as a hay crop. This species includes a number of other species likea. desertorum (Fisch.) Schult., A. pectiniforme Roem. & Schult., A. michnoi Roshev. and A. sibiricum (Willd.)P.B. AGROPYRON INTERMEDIUM (Host) Beauv. (syn. A. glaucum, Elytrigea intermedia (Host) Nevski). 2n=(28), 42, genome formula B2B2EEE2E2. Intermediate wheatgrass. S. and C. Europe to Iran, Pakistan and Caucasia. Self-compatible. AGROPYRON REPENS (L.) Beauv. (syn. Elytrigia repens (L.) Desv.). Couchgrass, Twitch, Quackgrass. 2n=8, 6x=42, genome formula SSS2S2XX, (56). Temperate Eurasia. Aggressive weed with

139 EUROPEAN SIBERIAN REGION wide adaptation. Spread to all continents. Sometimes used as palatable high-quality forage grass. It easily crosses with A. spicatum*, which introgresses as the F-^ is fertile (Dewey, 976). AGROSTIS CANINA L, 2n=4, 28, (35, 42, 56). Europe. Cultivated in the Netherlands. AGROSTIS GIGANTEA Roth. (syn. A. alba auct. non L.). Fiorin,Red top. 2n=42, genome formula AAA2A2A3A3. Europe, Asia and N. America. Cultivated as apasture grass and as ahay crop. AGROSTIS TENUIS Sibth. (syn. A. vulgaris With.). Rhode Island bent, Colonial bent.2n= 28, genome formula A^A-^A2A 2. Most of Europe, N. Asia Minor, Armenia, Caucasia, Siberia, N. Africa and N. America. Hybrids with A. gigantea* have been found in Germany and called A. intermedia C.A. Weber. ALOPECURUS PRATENSIS L. Meadow foxtail. 2n= 28, (42). Most of Europe, N. Asia and Caucasia. Cultivated as a meadow grass. AMMOPHILA ARENARIA Link (syn. A. arundinacea Host.). Beachgrass. 2n=28. The coastal areas of Europe. A perennial cultivated as a sand binder. ANTHOXANTHUM ODORATUM L. Sweet scented vernal grass, Spring grass. 2n=0, 20. Europe, Asia, W. part of N. Africa. Cultivated as a forage grass. It has a low food value. The diploid is also described as A. alpinum Love & Löve. Autoploidy has played an important role in the genesis of the tetraploid (Hedberg, 970). Teppner (970) suggested the following genome formula fora. alpinum and A. odoratum: species A. alpinum 2x A. alpinum 4x A. odoratum 2x A. odorarum 2x A. A. A. odoratum odoratum odoratum ploidy 2x 4x 4x genome formula AA AAAA CC DD DE BBDD BBFF region general Cantal, France Italy Italy, Yugoslavia, Greece Serbia Southern C. and W. Europe W. Europe A comparison of Austrian, Swiss, Swedish and Polish populations showed that diploids from Austria and Switzerland are morphological closer to those from Poland than to those in Scandinavia (Hedberg, 969). ARRHENATHERUM AVENACEUM Beauv. (syn. A. eliator Beauv.). Tall meadow oatgrass. 2n=40. Europe. A valuable pasture grass. ARRHENATHERUM TUBEROSUM Druce (syn. Avena tuberosa Gilib., Arrhenatherum avenaceum Beauv.). Onion couchgrass. 2n=8. In neolithic times possibly cultivated for its tubers. AVENA SEPTENTRIOLANIS Malz. (syn. A. fatua spp. septentrionalis (Malz.) Malz.). 2n=42. N. and NE. European USSR to W. Siberia. There it usually grows in undisturbed habitats. Baum (972) stated that is is probably the most closely related taxon to A. sativa* and that it resembles the hypothetical ancestor of the predomesticated oats. BROMUS ERECTUS Huds. (syn. B. arvensis Poll.). 2n=(28, genome formula AeAeAeAe), 42, 56, (70, 84, 2). C. and S. Europe, N. Africa, Ante- Asia up to Caucasia. Cultivated especially in S. France, Switzerland, S. Germany and USSR. Some people regard this species and its synonyms as two species. BROMUS INERMIS Leyss. Awnless brome, Smooth brome, Hungarian brome. 2n=(28,genome formula AiAiBiBi, 42, 49), 56,genome formula AiAiAi AiBiBiBiBi, (54-58). N., C, and SE. Europe, Caucasia, temperate Asia and China. Cultivation started at various places in Europe. Introduced to N. America. BROMUS SECALINUM L. Chess,Rye brome. 2n=28. A cultivated hulled cereal of prehistory grown together with emmer (Triticum dicoccum*)and einkorn T. monococcum*). Ithas a very high multiplication factor (2500 caryopses per plant). It isnow a weed mainly of winter cereals, but types with a spring habit are also found (Knörzer, 965). CYNOSURUS CRISTATUS L. Crested dogtail, Dogstail grass. 2n=4. Primary centre in C. and W. Europe, Caucasia and Asia Minor. DACTYLIS GLOMERATA L. Cocksfoot, Orchard grass. 2n=28. Stebbins (956)suggested that D. glomerata is a tetraploid derived from two related diploids. One of them could be D. aschersoniana Aschers. & Graebn. (2n=4). This species is distributed over C. Europe, Himalaya and W. China. Another diploid is D. smithii Link which exists inthe Canary Islands.It is likely that all diploids derive from one common diploid. Hybridization of diploids and doubling of the number of chromosomes and a- gain hybridization within the tetraploid group and with the diploids has led to the very variable D. glomerata. Cultivated as a pasture andhay grass. DIGITARIA SANGUINALIS Scop. 2n=8, 28, 36 (-48, 54, 76). Bluthirse, Millet sanguin.s. Europe, Asia Minor, Central Asia, N. and S. America, in temperate zones. There is a great variation of the species. The cultivated type is var. esculenta (Gaudin) Caldesi. Among this variety var. frumentacea Henr. and spp. aegyptiaca (Retz) Henr. are found. Primary centre is not known. Probably first cultivated in Illyria preceeded by a long time of collection of wild plants. Cultivated formerly in a large area

140 GRAMINEAE - GRAMINEAE 53 in Europe. Another area of cultivation is in India (p. 73). Whether the origin of cultivation independently arose here, or whether this cereal spread to India from Europe or the reverse is not known (Portères, 955a). Spp. pectiniforrais Henr. of E. Europe, the Near East and NE. Africa. Not cultivated. Spp. aegyptiaca has an 'eastern' origin but it is probably not in Egypt. From this subspecies the cultivated var. frumentacea is derived. Spp. vulgaris (Schrander) Henr. is very variable and widely distributed. FESTUCA ARUNDINACEA Schreb. 2n=(28), 42, (70). Europe, N. Africa and Asia (Syria, Siberia, Japan) Not much cultivated, due to its coarseness although seeds have been commercially available for a long time. According to Borrill (972) the tetraploid and hexaploid cytotypes have affinities with F. pratensis*, while the octoploid and decaploid possess a genome pair off. scariosa Aschs. & Graebn. (2n=4). This species is endemic in the Spanish Sierra Nevada. F. arundinacea has been rather widely introduced as a meadow and pasture grass in northern USA. FESTUCA OVINA L. Sheep's fescue. 2n=4, (2), 28, 42, (49), 56, 70. Europe, the Caucasus, the Himalaya and N. America. Cultivated in Europe. An important grass of Australia and S. Africa. Many 4x and 6x types have been described as Festuca species. FESTUCA PRATENSIS Huds. (syn.f. elatior L.). Fescue grass, Meadow fescue, English bluegrass. 2n=4, FpFp, (28, 42, 70). Europe, Caucasia, Iran, the Urals and Siberia. Cultivated in Europe and N. America.Natural hybrids with Lolium perenne* are described as Festulolium loliaceum (Huds.)P. Fourn. (syn. Festuca loliacea Huds.), 2n=2x=4, LpFp, loloid 3x=FpLp Lp, festucoid 3x=FpFpLp. According to Jauhar (975), F. pratensis and Lolium perenne are closely related and probably evolved from a common progenitor, as there is no effective intergeneric barrier to gene flow. FESTUCA RUBRA L. Red fescue. 2n=4, (28), 42, 56, (70 and aneuploids). Europe, temperate A- sia, Africa and N. America. Much cultivated as a pasture grass. In New Zealand chewings fescue is cultivated. It is a red fescue of the non-creeping type (spp. fallax). Var. genuina is creeping red fescue. GLYCERIA FLUITANS R. Br. Manna grass. 2n=(20), 28, 40. Was collected in a large part of E. Europe. HOLCUS LANATUS L. Soft meadow grass, Woolly soft grass, Yorkshire fog, Velvet grass.2n= 4. Europe and temperate Asia. Cultivated for pasture and hay. A secondary centre of diversity is developing innew Zealand (p. 65). LOLIUM MULTIFL0RUM Lam. var. westerwoldicum Wittm, (syn. spp. multiflorum (Husnot) Becherer). Westerwolds ryegrass. 2n=4. Annual types derived from populations of spp. italicum were selected at Westerwolde, NE. Netherlands. LOLIUM PERENNE L. Perennial ryegrass. 2n=4. Not know where and when it was domesticated, but probably in Europe. However, the parent plants may have come from the Mediterranean area of SW. Asia. The first true grass sown in a pure, or relatively pure state. Cultivated now in the Old and New Worlds. Tetraploids and amphiploids with Festuca pratensis*are cultivated. Natural hybrids between these two species are described as Festulolium loliaceum (Huds.)P. Fourn. Hybrids of L. perenne and L. multiflorum* have been called,l. x hybridum Hausskn. These last two species are closely related. PHALARIS ARUNDINACEA L. Red canary grass.2n= 4, 28. Most of Europe, W., N. and E. Asia. Cultivated inthe Old and New Worlds. PHLEUM PRATENSE L. Timothy, Herdsgrass.2n= mostly 42, genome formula NNAAXA2A2. Europe, N. Asia and N. Africa. An amphiploid of P. alpinum L. (Alpine timothy, 2n=28)and P. nodosum L. (syn.p. pratense var. nodosum (L.) Richter) (2n=4, genome formula NN(?)). A tetraploid type similar to this species was developed from the diploid Ph. nodosum after doubling the number of chromosomes. Ph. pratense is cultivated in Europe and N. America as a forage and hay crop. PHRAGMITES COMMUNIS Trinius. Reedgrass.2n= (36), 48, (54, 84, 96). A cosmopolite grass used for land reclamation and bank protection. Young sprouts are eaten, while the culms have many uses. POA BULBOSA* POA PALUSTRIS L. Fowl bluegrass. 2n=8, (42). Arctic zone of Europe, Asia and N. America. Various varieties have been developed in Europe. POA PRATENSIS L. Bluegrass, Kentucky bluegrass, Birdgrass. 2n= Europe, Asia, N. Africa and northern N. America.The great variation in chromosome number owing to autoploidization has resulted in many species descriptions, but they can be considered as synonyms. Furthermore as apomixy of this species isnot constant, types with different chromosome number may be selected. So it was possible to select plants similar top. pratensis from P. trivialis*. Ifthis provesthat P. pratensis derives from P. trivialis then P. pratensis must have originated in the Old World. Various varieties have been bred in Europe and Canada (p. 202) and elsewhere. POA TRIVIALIS L. Roughish meadow grass. 2n=4, (28). Europe and S. Siberia. Not much cultiva-

141 EUROPEAN SIBERIAN REGION ted. Itmight be the parent species ofp. tensis*. pra- SPARTINA ANGLICA CE. Hubbard, Cordgrass. 2n= 22. Originated in W. Europe after introduction of the American S. alternifolia Lois. (2n= 62)and hybridization with the European S. maritima (Curt.) Fern. (2n=60). The hybrid is named S. x townsendii H. &J. Groves (2n=62). From this hybrid the amphiploid S. anglica e- volved, which has ousted out its parent S. maritima (Adema & Mennema, 979). Cultivated for soil reclamation and stabilization. TRISETUM FLAVESCENS (L.) Beauv. (syn. T. pratense Bers.). Yellow catgrass, Golden oatgrass. 2n=24, 28.Itprobably derives from T. sibiricum Rupr. (2n=4, 24). This species occurs in Kamtschatka, Siberia. From here it spread westwards. TRITICUM AESTIVUM (L.) Thell. spp. compactus Host.). Clubwheat. 2n=42, genome formula AAB- BDD. The clubwheats of the Austrian alpines, except for the research of E. Mayr. are much neglected. They are probably derivatives of the wheat (T. antiquorum Heer) cultivated by the Swiss Lake Dwellers in the Neolithicum.They are nearly extinct. TRITICUM AESTIVUM (L.) Thell. spp. spelta (L.) Thell. Spelt. 2n=42, genome formula AABBDD. Cultivated from the Belgian Ardennes to Switzerland and to Schwaben, Germany and in Spain. Formerly the spelt area in Europe must have been much larger running from Sweden to Spain andmay be up to Africa (p.35). In Spain (Asturia) spelt is harvested in the same way as in Transcaucasia. It is remarkable that many German/Belgian spelts,the relic Swedish spelt (from Gotland) and one from Africa carry an Rf tim-gene (Zeven, 97). ZEA MAYS L. Maize. 2n=20. Secondary centres in S. Europe and the Mediterranean Region (p. 2) in the European corn belt and the Atlantic and Continental maize growing regions (Brandolini, 970). Domesticated in C. America (p. 90). Flint maize - indurata Sturt. - is common in all these areas. Siberian ecotypes are recognized by germination at 5-6 C, cold resistance of seedlings to 4-5 C, rapid growth, earliness, high assimilation rate and protogyny (Gerasenkov, 968). Grossulariaceae RIBES ACICULARIS Smith. 2n=. The mountains of Siberia especially in the Altai. The most precocious Ribes-species with a high winterhardiness and mildew resistance. These characteristics are useful in Ribes-breeding. RIBES GROSSULARIA L. (syn.r. uva-crispa L.). (European) Gooseberry. 2n=6. Eurasia and in the mountains of W. Asia and the Mediterranean countries. Cultivated in temperate zones. Re- Ribes acicularis lated N. American Ribes-species R. oxyacanthoides Mill. (2n=6), R. hirtellum Mix. (2n=6), R. divaricatum Dougl. (2n=6), R. cynosbati L.* (2n=6), R. pinetorum Greene (2n= )and R. niveum Lindl. (2n=6)carry resistance to mildew, while R. niveum andr. divaricatum may be used as source of mildew resistance and to improve fruit characteristics. Resistance to Nasononia ribisnigri Mosley is found in R. roezlii Regel (2n=6) andr. sanguineum Pursh (2n=6), while the latter species and R. cereum Dougl. (2n=6)are sources of resistance to Hyperomyzus lactucae L. (Keep & Briggs, 97). Hybrids between R. grossularia andr. sanguineum are namedr. fontenayense Jancz. (2n= ). Spineless types are also found ofr. oxyacanthoides. RIBES NIGRUM L. (European) Black currant.2n= 6. Eurasia and sporadically in N. America. The cultivated type was derived from the wild one. In N. Scandinavia very precocious, winterhardy types are found. The American R. americanum Mill. (2n= )and theasiatic R. dikuscha Fish, are related to the black currant. They have breeding value. Cultivars of var. sibiricumf. Wolf, of this species andr. ussuriense* are sources of resistance to the blackcurrant gall mite. Phy-

142 GRAMINEAE - LEGUMINOSAE 55 toptus ribis Nal. Spontaneous hybrids with R. procumbens (2n= ) occur in the USSR. RIBES PETRAEUM Wulfen. Rock red currant. 2n= 6. The Pyrena to the Carpates and N. Africa. Cultivated in the Alps. One of the parents of the present-day red currant (R. sativum*). RIBES SATIVUM Syme (R. rubrum L., R. multiflorum Kitt, and R. petraeum Wulf.). 2n=6. The wild R. sativum grows in W. Europe. In N. America it has run wild. R. rubrum is found wild in W. and C. Europe and N. Asia. R. petraeum* grows in the mountains of Europe and Asia. R. sativum is probably the originally cultivated species. Later it hybridized with the other two, so these three species are the parents of the present-day red currant. RIBES SPICATUM Robson. Sometimes cultivated. Guttiferae 2n=6. NE. Europe. HYPERICUM PERFORATUM L. Saint Johns wort. 2n= 32, (36). Cultivated on a small scale in the Netherlands as a medicinal crop. Juglandaceae JUGLANS REGIA L. Walnut, Persian walnut, English walnut. 2n=32, 36. Primary centreof diversity in Region5. Secondary centre in SW. Europe and Moldavia. Almost all varieties in Germany are apomictic. Labiatae MENTHA CARDIACA Gerard ex Baker. Scotch mint, Scotch spearmint. 2n=. Temp. Europe. Cultivated for its volatile oil. Closely related to M. x gentilisl. (2n=54, 60, 84, 96, 08, 20). Itis believed that these two species are hybridsof M. arvensis*and M. spicata*. MENTHA x GENTILIS L. (syn. M. sativa var. gentilis (L.) Reichenb.). 2n=54, 60, 84, 96, 08, 20. A hybrid of M. arvensis* and M. spicata*. Usually sterile. Cultivated frequently. MENTHA x PIPERITA L. Peppermint. 2n=(36, 48, 54-69), 72, (84, 08, 22, 44). Probably a natural hybridof M. aquatica*and M. spicata*. This hybridization probably took place in England,f. piperita (blackmint, black mitcham) is cultivated in C. Europe and Great Britain, whilef. pallescens Camus (white mint, white mitcham)is cultivated especially in France. In USA existing Clones were replaced by the cultivar Mitcham in 890. This is still the main clone cultivated. MENTHA ROTUNDIFOLIA (L.) Huds. (syn. M. spicata var.rotundifolia L.). Apple mint, Woolly mint. 2n=24, genome formula RR. Europe and Canary Islands. Cultivated. Probably the parental formof M. spicata*and one of the parents of M. japonica Mak., M. arvensis*and M. a- quatica* (Ikeda & Ono, 969). This species is related to M. longifolia*and M. spicata*. MENTHA x SMITHIANAR.A. Graham (syn. M. rubra Sm., non Miller). 2n=54, 20. Rarely cultivated (Tutinet al., 972). It is a hybrid of M. aquatica* x M. arvensis* x M. spicata*. Usually sterile, spreading vegetatively. MENTHA SPICATA (L.) Hudson (syn. M. viridis L.). Spearmint, Green mint,lamb mint. 2n=36, 48, genome formula RRSS (48+2B, 64). Temp. Europe.It might derive froman autotetraploid plant of M. rotundifolia* after which one genome pairrr changed into SS. Tutinet al. (972) suggested that this species arose in cultivation as a segmental allopolyploid of M. suaveolens (see M. x rotundifolia*) and M. longifolia*. Var, crispata Schrader (syn. M. crispal.) has genome formula RRS C S C. This species is one of the parents of M. x piperita*. It might be one of the parentsof M. x villosa*. Murray et al. (972) artificially crossed M. aquatica (2n=96)and M. spicata (2n=48). This resulted in very variable Fidue to the heterozygosity of the pollen parent. Some hybrids resembled the natural strainsof M. x piperita, othersdid not. MENTHA SUAVEOLENS Ehrh. (syn. M.rotundifolia auct., non (L.) Hudson). 2n=24. Cultivated as a potherb. MENTHA x VILLOSA Hudson (syn. M. cordifolia auct., M. gratissima Weber). 2n=36. This species is a hybrid of M. spicata*and M. suaveolens*. NEPETA CATARIA L. (syn. Cataria vulgaris Moench.). Catnip, Catmint. 2n=(32), 34, (36). Europe. A perennial herb cultivated for medicinal purposes. ORIGANUM VULGAREL. Wild majoram. 2n=30. An extremely variable species from the Azores, Madeira,the Canary Islands, Europe throughout the Mediterranean Region, W., C. and E. China to Taiwan.Its great variation has resulted in numerous synonyms (Ietswaart, 980). Cultivated as a medicinal plant. Leguminosae ANTHYLLIS VULNERARIA L. Kidney vetch, Spring vetch, Lady's fingers, Wound-wort, Amer, Tare. 2n=2. Temp. Europe, Caucasia, Ante-Asia, N. Africa and Ethiopia. Cultivated since 858 (Mansfeld, 959) and is now usually mixed with pasture grasses. ASTRAGALUS CICER L. Milk vetch. 2n=64. Europe. A perennial pasture plant well-adapted for grass

143 56 EUROPEAN SIBERIAN REGION mixtures (Whyte et al., 953). ASTRAGALUS FALCATUS Lam. Sicklepod milk vetch. 2n=6. W. Asia. A forage plant cultivated in USSR and France. ASTRAGALUS GLYCYPHYLLUS L. Milk vetch. 2n=6. Europe and Siberia to Altai. A perennial herb cultivated as a fodder. CORONILLA VARIA L. Crown vetch. 2n=24. C. and S. Europe extending toc. Russia. Cultivated as an ornamental, a fodder crop and a cover crop. l\ I { \ \>V Medicago falcata (Fischer, 938). j vx * rfb'' fa GALEGA OFFICINALIS L. Galega, European goat's rue. 2n=6. E., C. and S. Europe, Caucasia, Asia Minor and Iran. Cultivated as a forage crop and as an ornamental. GLYCYRRHIZA ECHINATA L. 2n=6. SE. Europe to Hungary and Italy. Cultivated toproduce liquorice. GLYCYRRHIZA GLABRA (syn.g. glandulifera Waldst. & Kit.). Common licorice, Liquorice. 2n=6. Europe and the Mediterranean region. A perennial herb. Var. typica Regel & Herder is cultivated to produce Spanish or Italian licorice and var. glandulifera Waldst. Russian licorice. HEDYSARUM HEDYSAROIDES (L.) Schinz. & Thell. (syn. H. alpinum Jacq.). 2n=4. S. Europe, Asia Minor, America and Caucasia. Cultivated as a fodder crop especially in the Alps. LATHYRUS SYLVESTRIS L. Flat pea, Wood pea. 2n=4. Europe. Cultivated for forage and as an ornamental plant. LATHYRUS TUBEROSUS L. Groundnut peavine, Earth chestnut. 2n=4. Europe and W. Asia. Cultivated for its tubers. In the 6th Century its flowers were distilled forperfumes (Uphof, 968). LOTUS CORNICULATUS L. Birds-foot trefoil.2n= 2, 24. Europe, moderate Asia and N. Africa to Ethiopia. Formerly and at present inusa in use in seed mixtures for a ley crop and for pastures. Landolt (970)and Somarov & Grant (97) suggested that thediploid is a hybrid and the tetraploid an allotetraploid of L. alpinus Schleicher (2n=2)of the Alp and the submediterranean L. pilosus Jord. (2n=2). The erect, broad-leaved type probably from C. European origin is spread now as a contaminant of grass seed for road sides throughout W. Eurèpe (Jones, 973). LOTUS ULIGINOSUS Schkuhr. Greater birds-foot trefoil. 2n=2, (24). Europe, N. Africa, Ante- Asia to Tibet. Cultivated in C.Europe and Great Britain as a fodder crop (Mansfeld, 959). MEDICAGO FALCATA L. Yellow lucerne. 2n=6, 32. In Europe and Asia from longitude 0 to 85 E. and latitude 42 to 60 N., and mountains near the S. limits. The 4x is common while 2x is rare but occurs in the whole area. The 2x ssp. romanica (Prod.) Hayek possesses some rare characteristics andmay derive from an older 2x stock (Lesins & Lesins, 979). MEDICAGO GLOMERATA Balbis. 2n=6, (32?). Maritime Alps. Useful as gene source of M. sativa*. MEDICAGO LUPULINA L. Hop clover,black medic. 2n=6, genome formula SS, 32, (64). Europe, most of Asia and N. Africa. It is naturalized in N. America.The 4x type has been found in C. Siberia. Occasionally included inseed mixtures for pastures. Cultivated since 659 in England and 785 in France, and now in the Old and New Worlds as a green fodder,hay crop and green manure (Lesins & Lesins, 979). MEDICAGO SATIVA L. Lucerne, Blue alfalfa. 2n= (6, genome formula SS), 32, (48, genome formula SSSSSS, 64). Transcaucasia (p. 97). Two populations - one from thebalkans and one from France - "met"in Thuringia, Germany. This resulted in a hybrid swarm from which winterhardy types were introduced in Minnesota, USA in 857. MELILOTUS ALBUS Medik. White sweet clover, Bokhara clover, Honey clover, White melilot.2n== 6. Europe and W. Asia. Cultivated in theold World and particularly in the USA as a fodder crop and green manure. It can be divided into two groups ) the annual wild type and 2) the bushy type. The latter might be a mutant of group, or derive from a natural cross of M. albus. From both groups cultivars have been selected. The very low variation of this species may point to only a few introductions. MELILOTUS ALTISSIMUS Thuill. 2n=6. Europe and temp. Asia.Sometimes cultivated for horse fodder. MELILOTUS DENTATUS (Waldst. & Kit.) Pers. 2n= 6. E. and C. Europe to N. Sweden. Coumarin deficient and salt tolerant. Used to breed coumarin free cultivars of M. albus*.

144 LEGUMINOSAE - LILIACEAE 57 MELILOTUS MACRORHIZUS Pers. 2n=6. Asia and Europe. Cultivated in China for its roots which are eaten as a vegetable. Closely related tom. altissimus*. MELILOTUS OFFICINALIS Lam. Biennal yellow sweet clover, Field melilot, Yellow melilot. 2n=6. W. Europe to W. China. Cultivated in Europe and also in the USA. It is a biennal with sporadically some annuals. ONOBRYCHIS VICIIFOLIA Scop. (syn. 0. sativa s.l.lam.). Esparcette, Sainfoin. 2n=8. Temp. Europe, SW. Asia to Altai and Transbaikal. Cultivation was probably started ins.france resulting in spp. sativa. There are three subspecies: arenaria (Kit & Koch.) Thellung, sand esparcette, montana (Lam. & D.C.), the mountain esparcette and sativa (Lam.) Thellung, the cultivated esparcette. The last name is confusing, because spp. arenaria (alsodescribed as var. transcaucasia, syn.0. transcaucasia Grossh.)* is cultivated too. SAROTHAMNUS SCOPARIUS (L.) Wimm. ex Koch. Broom. 2n=(4), 46, 48. W. and C. Europe. Cultivated as asoil stabilizer. TRIFOLIUM HYBRIDUM L. (syn. T. fistulosum Gilib.). Alslike clover. 2n=6. Temp. Europe, SW. Asia and N. Africa.Possibly first cultivated in Sweden. Introduced to other European countries and N. America. Often found in fields of red clover. Very likely not the ancestral form of T. repens*. The cultivated type spp. hybridum is probably derived from the wild type spp. elegans (Savi) Asch. & Graebn. In Anatolia spp. anatolicum (Boiss.) Hossain is found. TRIFOLIUM PANNONICUM Jacq. Hungarian clover. 2n=c. 96, 98,c. 26,c. 30,c. 80. E., C and S. Europe. Cultivated. TRIFOLIUM PRATENSE L. Red clover. 2n=4, genome formula AA, (28). Europe, W. and C. Asia and N. Africa. Primary centre in Region 9. It was probably first cultivated in the Netherlands, in thebeginning of the 6th Century. The Classics already mentioned 2000 years ago that local ecotypes were developed in SE. Europe and Asia Minor.Spread togermany and through Flanders to England. In the beginning of the 7th Century seed of red clover was exported from the Netherlands to the Scandinavian countries and France.From England red clover was spread toussr and N. America.The^ wild type has more leaves and new shoots emerge from internodes at the butt end, while the cultivated type has less leaves and new shoots emerge from the leaf rosett. The variable wild type is described as var. pratense Bobr. and the cultivated as var. sativum (Crome) Bobr. (syn.t. sativum (Sturm) Crome. Late red clover (var. serotinum) may have developed from contaminants or spontaneous mutants in USSR from introduced early types (var. praecox). Autotetraploid types are widely cultivated now. Var. americanum CO. Hartz was cultivated between 883and c. 90 in C. Europe. It originated from a N. American introduction. It is often erroneously described as T. expansum Waldst. tit Kit. Var. maritimum Zabel (var. villosum Wahlberg) is found wild on the S. coast of the Balkan Peninsula and var. frigidum Gaudin occurs wild in the Alps. Red clover is closely related to the annual T. diffusum Ehrh., 2n=6, tothe annual T. pallidum Waldst. & Kit.,2n=6 and the perennial T. noricum Wulf., 2n=6. TRIFOLIUM REPENS L.White clover. 2n=32, (48, 64). Wild type (var. sylvestre). In meadows throughout Eurasia and N. Africa. Cultivation started probably in N. Italy (p. 5)and in the Netherlands. Very variable. Brewbaker & Keim (953)suggest that T. nigrescens Viv., Ball clover (2n=6) isone of the parents. Chen & Gibson (970) believe that it is an autotetraploid while T. nigrescens and T. occidentale D. Coombe (2n=6) are related to it. T. uniflorum L. (2n=32) might also be a parent. This species is found in E. Mediterranean area to Sicily. It includes T. savianum Guss. of Sicily and Calabria, Italy. It isprobably an autotetraploid. Navalikhina (977) suggested that white clover is an allopolyploid of T. nigrescens, T. occidentale and T. uniflorum. TRIFOLIUM RESUPINATUM L. (syn. T. suaveolens Willd.). Persian clover. 2n=(4), 6. The Mediterranean area to Iran, Afghanistan and India. Cultivated as a fodder crop. Var. majus Boiss. is syn. to T. suaveolens Willd. TRIGONELLA C0ERULEA (L.) Ser. Sweet trefoil. 2n=6. Cultivated and also found as a weed or ruderal. Itmay be derived from T. procumbens (Besser) Reichenb. (syn. T. besserana Ser., T. coerulea spp. procumbens (Besser) Thell.). This species is a native to EC. and SE. Europe. ULEX EUROPAEUS L. Common corse. 2n=96. W. Europe to Italy. Cultivated formerly for fodder, bedding and as hedges. VICIA CRACCA L. Gerard vetch. 2n=2, 4, (2, 24), 28. W. Europe to Kamtchaska, E. China and Japan. Cultivated. VICIA HIRSUTA (L.)S.F. Gray. Common tare, Hairy tare. 2n=4. Europe, N. Africa and W. Asia. Cultivated in W. of USSR together with barley. VICIA PANN0NICA Crantz. Hungarian vetch. 2n= 2. Primary centre in SW. Asia (p. 98). Secondary centre in Hungary. Liliaceae ASPARAGUS OFFICINALIS L. Garden asparagus.2n

145 EUROPEAN SIBERIAN REGION =20. Primary centre probably in the saltsteppes of E. Europe. A. officinalis var. prostratus Richter is a tetraploid (Braak & Zeilinga, 957). CONVALLARIA MAJALIS L. Lily-of-the-valley. 2n =32, 36, 38. Europe, temp. Asia and Japan. A perennial herb cultivated as a medicinal crop and as an ornamental. Linaceae LINUM USITATISSIMUM L. Flax, Linseed. 2n=30, (32). For origin ;ee p. 99. Helbaek (97) supposed two ways of introduction of flax. One through Greece and the Donau valley into C. and W. Europe and the other west of the Black Sea in a northern direction into Russia,The first was probably a winter-annual which is the parent of "Winterlein" cultivated in Germany. The other was probably a summer-annual. Inthe firstmillenium B.C. the latter was introduced to C. and W. Europe. It is at present described as spp. eurasiaticum Vav. & Ell. In NW. USSR there is a centre of flax containing some of the finest fibre flax varieties. In W. Europe and Ukraine the weed rattle flax, L. crepitans Dumort. (2n=30), now included in L. usitatissimum is probably the weedy type of flax. Malvaceae ALTHAEA OFFICINALIS* Paeoniaceae PAEONIA OFFICINALIS L. Peony, Piney. 2n=20. S. Europe, Asia Minor and Armenia. A perennial herb cultivated for its medicinal merits. Plantaginaceae PLANTAGO LACEOLATA L. Rib grass. 2n=2, (3, 2 + 2B). Cultivated on a small scale in the Netherlands and elsewhere as a medicinal crop. Polygonaceae RUMEX ACETOSA L. Garden sorrel. 2n=4 Q, 5 Cfand other numbers. Temp. Europe and Asia. A perennial herb. Var. hortensis Dierb. (syn. R. ambiguus Gren.) is cultivated in the Old andnew Worlds. RUMEX ALPINUS L. Alpine dock, Monk's rhubarb. 2n=20. C. Europe, the Balkans and Caucasia. Cultivated formerly in C. Europe as a vegetable. RUMEX OBTUSIFOLIUS L. ter dock. 2n=40, (60). Cultivated. Broad-leaved dock, Bit- Europe and temp. Asia. RUMEX PATIENTIA L.Patience dock, Spinach dock, Herb patience. 2n=(40), 60. Probably C. Europe to W. Asia. Cultivated as a vegetable. RUMEX SCUTATUS L. (R. alpestris Jacq.). French sorrel. 2n=20, (40). C. and S. Europe, Alpine regions, Caucasia, India. Cultivated as a vegetable (var. hortensis Lam. & DC). Portulacaceae PORTULACA OLERACEA L. Purslane. 2n=2x=8,4x= 36, 6x=54. Purslane is a cosmopolitan weed whose origin is doubtful (Daninet al., 978). The cultivated type (ssp. sativa (Haw.) Celak., 2n=54)is a vegetable, which probably developed in Europe from the Eurasian weedy type ssp. oleracea, 2n=54. The distribution of the 2x, 4x and 6x wild and weedy subspecies indicates that one of the centres of diversity is Mexico (p. ) (Daninet al., 978). Ranunculaceae ACONITUM NAPELLUS L. Monkshood. 2n=24, 32. C. Europe. Cultivated as a medicinal crop and alsoas an ornamental. AQUILEGIA VULGARIS* NIGELLA SATIVA* Resedaceae RESEDA LUTEOLA* Rhamnaceae RHAMNUS CATHARTICUS L. Buckthorn. 2n=24. Europe up to Transcaucasia and W. Siberia, and in Algeria. Formerly the fruits of this tree were used as a source of yellow dye. RHAMNUS FRANGULA L. Alder buckthorn. 2n=20, 22, 26. Europe, Asia and N. Africa. A tree formerly cultivated. Rosaceae AGRIMONIA ODORATA (Gouan) Mill. 2n=56. Itis included in A. eupatoria L., Agrimony. Cultivated as a medicinal crop. AMYGDALUS BESSERIANA Schott, (syn. A. nana L., Prunus nana (L.) Stokes, P. tenella Batsch.). Dwarf almond, Dwarf Russian almond, Steppe almond. 2n=6. Primary centre in E. Europe and Siberia. Also wild inthe Balkan, Asia Minor, Causasus and China (p. 42). It is the commonest wild almond species and it is very frost resistant which makes it extremely valuable as a rootstock of A. communis. AMYGDALUS LEDEBOURIANA Schlecht, (syn. Prunus ledebouriana Schlecht.) 2n=. A shrub from Tarbagatai and Altai. AMYGDALUS PERSICA L. Peach. 2n=6. Primary centre in China (p. 42). Secondary centre in Moldavia, USSR.

146 LILIACEAE ARMENIACA BRIGANTINA (Vizl.) Pers. (syn. Prunus brigantina Vill.). Briançon apricot. 2n=. Originated in SE. France. The seeds are the source of the perfumed oil "Huile de Marmotte" (Uphof, 968). It might be agene source of late flowering. ARMENIACA SIBIRICA Pers. (syn. Prunus s i b i r i- ca L., P. armeniaca var. sibirica K. Koch.). Siberian apricot. 2n=6. Intern Mongolia to the Sowjet Far East and Lake Baikal. This species has the largest distribution of all apricot species (Zylka, 970). It is very cold resistant. ~K. vu A y^ JÉWt- " y\ o^ Fragaria moschata strawberry. 2n=4, genome formula AA. Europe, Asia and N. America (p. 204). Darrow (955) stated that var. semperflorens Duch. is the parent of the cultivated strawberry. It was domesticated N. of the Italian Alps. Cultivated from seed and vegetatively. FRAGARIA VIRIDIS Duch. Polunitsa. 2n=4. European part of Region 9. Cultivated formerly. Armeniaca sibirica FRAGARIA X ANANASSA Duch. (syn.f. grandiflora Ehrh.). Pineapple strawberry. 2n=56. Arose spontaneously inw. Europe (in a garden near Haarlem, the Netherlands) after hybridization of F. virginiana* from N. America and F. chiloensis* from S. America. F. ovalis (Rydb.) Lemm. from NW. USA is used as a source of winterhardiness. FRAGARIA MOSCHATA Duch. Hautbois strawberry. 2n=42. Europe and European USSR. Cultivated formerly, and runwild in other countries. FRAGARIA VESCA L. Wild strawberry, Alpine MALUS BACCATA (L.) Borkh. var. baccata. Siberian crab apple. 2n=34. Wild in Transbaikal and Ante-Baikal territories. Primary gene centre in Siberia.Resistant to frost. MALUS PRUNIFOLIA (Willd.) Borkh. (syn. Pyrus prunifolia (Willd.). Chinese crab apple. 2n= 34, (5, 68). Wild and cultivated in the extreme eastern sector of Region 9. Primary centre China. MALUS PUMILA Mill. (syn. Pyrus malus L.) Apple. 2n=34, 5, 68. The Balkans and SW. US SR (p. 85), eastwards through Transcaucasia, Iran, Turkestan, and northwards to the Altai mountains. It occurs along the ancient and mediaeval routes of commerce and migration between Europe and E. Asia. Man has greatly promoted its distribution (Wilcox, 962). It is considered as the principal ancestor of the

147 60 EUROPEAN SIBERIAN REGION spp. italica (Borkh.) Hegi). Bullace plum, Damson plum. 2n=48. S. and SE. Europe and adjacent parts of Asia. Occurs now throughout temp. Europe and W. Asia. Probably only known as a cultigen and naturalized. If so,it is obviously an allohexaploid. It is frost resistant. PRUNUS MAHALEB L.Mahaleb cherry, St. Lucie cherry. 2n=6. C. and S. Europe and W. Asia. Fruits are not edible. Used as rootstock for cultivated cherries. Mainly self-incompatible. PYRACANTHA COCCINEA M.J. Roemer. 2n=34. S. Europe and westwards to NE. Spain. Cultivated as an ornamental and for its fruits. Malus baccata cultivated apple. M. sylvestris* hybridizes with this species and hence may also have played a small part as an ancestor. European USSR is the primary centre for many old cultivars as Antonovka, Aport, Borovinka. Inearly 9th Century, Bolotov, described 600 Russian cultivars; about cultivars exist in the world today. This shows the very polymorphic nature of this species which has also arisen due to introgression with other species. PRUNUS CERASUS L. Sour cherry. 2n=32, genome formula CC. C. Asian centre (p. 0). The population Vladimirskaya vishnia with large darkscarlet fruits that are very palatable and aromatic, originated in Region9, extending westward and southward to the Rhine and Balkans. PRUNUS DOMESTICA L. Garden plum, Domestic plum. 2n=48, genome formula CCSSSS or CdCdSS SJSJ^ or CdCdDiD D 2D 2. For origin see p. 0. Werneck (958) considered Upper Austria as a place where the garden plum has arisen. Bush seedling would have been transplanted tocompounds where further domestication may have occurred. The Lake Bank Dwellers of neolithic Switzerland knew the garden plum. PRUNUS FRUTIC0SA Pall. (syn. Cerasus fruticosa Pall.). Dwarf Cherry, Bush Cherry, Ground Cherry, Mongolian Cherry, Steppe Cherry. 2n= 6, 32. Extended over Europe. It occurs in great diversity beyond the Volga, in S. Ural, SW. Siberia and Bashkirskaya. One of the parents ofp. cerasus*.it withstands -52 C. PRUNUS INSITITIA L. (syn. P. domestica var. insititia (L.) C.K. Schneider, P. domestica PYRUS COMMUNIS L. (syn.p. domestica Med.). Common pear. 2n=34, (5). Europe and W. Asia. It has been divided into spp. pyraster L. (syn. P. pyraster Burgsd., P. communis var. achras Wallr.), spp. nivalis Jacq. (syn. P. nivalis Jacq.) and spp. salvifolia (syn.p. salvifolia DC.). Spp. pyraster is the most important one, it grows inc. Europe and W. Asia. Spp. nivalis, the snow pear grows in W. Switzerland and France. It isused as a rootstock. Spp. salvifolia is found inthe same areas. The cultivars are derived from these subspecies by selection and by crossing with P. serotina*,p. ussuriensis*, P. longipes, P. caucasica*,p. amygdaliformls* andp. sallcifolia*. Pavlov (969a, 969b) reported that types of W. and S. Europe derive from crosses with P. nivalis* and P. amygdaliformis*, because they have hairiness and a high number of stomata per area like these two species. Some cultivars in Caucasus show characteristics obviously derived from P. caucasica*. The E. European cultivars show adirect derivation from the wild P. communis. PYRUS C0RDATA Desv. 2n=. W. Europe. Cultivated in hedges and for its wood. ROSA x ALBA L. French rose. 2n=28, 42. Cultivated in Bulgaria and S.France for the perfumery industry. Probably a hybrid ofr. arvensis* xr. gallica*, and a white flowered member of the Sect. Canina. ROSA ARVENSIS Hudson. 2n=4. S., W. and C. Europe. It is one of the parents ofr. x alba*. ROSA x BIFERA (Poiret) Pers. (syn.r. damascena auct., non Miller). Damask rose. 2n=28. Probably a hybrid ofr. moschata* and R. gallica*. Cultivated in Bulgaria, S. France and Turkey. The petals are used toproduce oil of roses which is used in perfumery. ROSA CANINA L. Brier,Dog rose, Doghip. 2n=35. Europe, temperate Asia and N. Africa. It is a common rootstock of garden roses. The named selections are often less prickly than the wild ones.

148 ROSACEAE - SALICACEAE 6 ROSA GALLICA L. French rose. 2n=28. S. and C. Europe up tobelgium and C. France and W. Asia. Probably a parent ofr. x bifera* and R. x alba*. The petals are used in perfumery. ROSA RUBIGINOSA L. (syn. R. eglanteria auct.). Sweet briar. 2n=35. W. and C. Europe. Cultivated for its flowers and as rootstock. ROSA VILLOSA L. Apple rose. 2n=28. Var. pomifera (Herrm.) Crép. Europe and SW. Asia. RUBUS ARCTICUS L. Arctic bramble, Nectarberry. 2n=4. Europe and N. Asia. Used in breeding work with R. idaeus*. Fruits have a distinct aroma and rich in Vit. C. A hardy, high yielding, disease resistant plant. RUBUS CHAMAEMORUS L. Cloudberry, Yellowberry, Salmonberry, Bake apple. 2n=56. Europe and N. Asia used in breeding with R. idaeus*. Easily domesticated (Larson, 969). Seed and esp. subterranean runners are used to propagate this dioecious species. RUBUS IDAEUS L. European red raspberry. 2n=4. Spp. vulgatus wild in Europe. It was domesticated. The present cultivars often are hybrids of this subspecies and its NE. American counterpart spp. strigosus. The tetraploid subspecies: melanolasis Focke from NW. America and Siberia, sachalinensis Léveillé from Sakhalin and sibiricus from Kamchatka, have been grouped asr. sachalinensis Levi. Some cultivars derive from R. idaeus x R. chamaemorus*, cloudberry (2n=56). Crosses have also been made between this species and R. R. xanthocarpus Bur. & Franch from W. China, R. arcticus*l., Arctic bramble (2n=4)and P. parviflorus L., Japanese raspberry (2n=4). Other Rubus species have also been used in breeding work. Everbearing types have been developed. RUBUS LACINIATUS Willd. Evergreen blackberry. 2n=28. C. Europe. A cultivar was brought to N. America where hybridization took place with another European immigrant,r. procerus P.J. Muell. (2n=4, 28, 49), Himalaya berry. In 925, a mutant was found in Oregon and named 'Thornless Evergreen'. This cultivar and its minor mutants are commonly grown in the USA. RUBUS SAXATILIS L. Stoneberry. 2n=8. Europe and N. Asia. InSweden a species has been found to be resistant to rust and other diseases. The fruit has only a few drupelets and lacks flavour (Larsson, 969). SANGUISORBA MINOR Scop. 2n=8, (54, 56). Europe and temp. Asia. Sometimes cultivated to flavour soup or for salads (Mansfeld, 959). SANGUISORBA OFFICINALIS L. Great burnet, Garden burnet. 2n=8, (42, 56,c. 70). Europe, Asia and N.'America. Sometimes cultivated as a vegetable (Mansfeld, 959). SORBUS AUCUPARIA L. (syn. Mespilus aucuparia All.). Rowan tree,european mountain ash.2n= 34. The "Mährische Eberesche" (var. moravica) was found in 80 in Czechoslovakia. It has been improved and distributed. Before its domestication var. rossica and var.rossica-major were already cultivated in USSR. It is an important source of Vitamin C (Mueller-Stoll & Michael, 949). SORBUS DOMESTICA L. Service tree, Mountain ash. 2n=34. S. Europe, N. Africa and W. Asia. Cultivated in Europe for its fruits which are eaten or made intowine and as an ornamental. Largefruited forms are found in forests in Crimea (p. 0). Rubiaceae RUBIA TINCTORUM L. Madder. 2n=44. S. Europe and Asia Minor. Cultivated in Europe as a dye plant. Salicaceae SALIXACUTIFOLIA L. Caspic willow. 2n=38. A tree of USSR and Manchuria. Cultivated for twig production. SALIX ALBA L. (syn. S. aurea Salisb.). White willow. 2n=76. In large area of Europe and Asia (p. 86) and N. Africa. Introduced into N. America. Cultivated in Europe for twig production for dike building. SALIX CAPREA L. Goat willow, Common willow. 2n=38, (57, 76). Europe and N. Asia. Cultivated for its twigs. SALIX FRAGILIS L. Brittle willow, Crack willow. 2n=(38), 76, (4). Europe, Asia Minor, Syria, Iran and W. and C. Siberia.Often planted for twig production. It isone of the parents of S. x rubens Schrank. SALIX PURPUREA L. Purple willow, Purple osier willow. 2n=38. A large part of Europe, and in Asia to Japan, and in N. Africa. Cultivated for twig production for dike works and basketry. SALIX TRIANDRA L. (syn.s. amygdalina L.). French willow, Almond-leaved willow. 2n=38, 44, (57, 88). Spread from W. Europe to E. Asia. Planted for twig production. One of the parental species of the cultivated S. x mollissima Ehrh. (2n=38). SALIX VIMINALIS L. (syn. S. longifolia Lam.). Twiggy willow, Common osier, Basket willow, Osier willow. 2n=38. C. Europe and a large part of Asia. Much cultivated in N. and S. Europe and elsewhere for twig production. Many of the willows planted in the Netherlands for dike work belong to this species and to S. triandra*. They are often cultivated in mixed stands which leads to cross fertilization and development of hybrids.

149 62 EUROPEAN SIBERIAN REGION S. dasyclados Wimmer (2n=38, 57, 76, 4) is archangelica includes the cultivated type, probably a complex hybrid of S. caprea x S. cinera x S. viminalis. S. helix L. is a hybrid ANTHRISCUS CEREFOLIUM (Waldst.& Kit.) Sprengel, of S. purpurea* x S. viminalis. In Great Bri- Garden chervil. 2n=8. Probably EC. and SE. tain, this willow is planted as a windbreak Europe. Var. cerefolium has glabrous fruits, and to shelter cattle. It includes the cultivated type. Sambucaceae SAMBUCUS EBULUS L. Dwarf elder. 2n=36. From Netherlands and Ukraine southwards. Formerly cultivated as a medicinal plant. Naturalized elsewhere. SAMBUCUS NIGRA L. European elder. 2n=36. Europe. Cultivated. Recently there has been a new interest in this tree because of the processing of alcohol-free beverage (Strauss & Novak, 97). Saxiphragaceae BERGENIA CRASSIFOLIA (L.) Fritsch. 2n=34. Siberia, Altai and N. Mongolia. A perennial herb cultivated since 927 inussr as a tea plant (Mansfeld, 959). Scrophulariaceae DIGITALIS LANATA Ehrh. 2n=56. SE. Europe. Elsewhere in Europe it may have run wild. Cultivated as a medicinal crop. DIGITALIS PURPUREA L. Purple fox-glove. 2n= 56. S. (p. 8) and C. Europe. Cultivated as a medicinal plant and as an ornamental. VERBASCUM THAPSIFORME Schrad. 2n=32. Spread throughout Europe. Cultivated for its medicinal properties and as an ornamental. Solanaceae CAPSICUM ANNUUM L. Bell pepper, Paprika, Cayenne pepper. 2n=24. Mexico (p. 96). Secondary centre in Europe. PHYSALIS ALKEKENGI L. Strawberry tomato, Winter cherry. 2n=24. C. and S. Europe. A perennial herb cultivated for its fruits. SCOPOLIA CARNIOLICA Jacq. Scopalia. 2n=46-48, 48. Europe. Cultivated as a medicinal crop. SOLANUM TUBEROSUM L. Potato. 2n=48. Domesticated in S. America. In Europe spp. tuberosum developed. Its genetic basis isvery small. This is probably caused by only a few introdutions, and afterwards by the selection for short-day forms and by mass killing during blight epidemics in the 840's. Umbelliferae ANGELICA ARCHANGELICA L. Angelica. 2n=22. Temperate Europe, Himalaya, Siberia and Kamtschatka. Cultivated for its aromatic petioles. Spp. BUNIUM BULBOCASTANUM L. (syn. Ligusticum bulbocastanum Crantz). 2n=. W. Europe. Formerly cultivated for its edible tubers. CARUM CARVI L. (syn. Apium carvi Crantz). Caraway. 2n= 20, 22, (23, 25). Europe Äid W. Asia. Cultivated in temperate regions, N. India and Sudan (seec. roxburghianum*, C. copticum*). CHAEROPHYLLUM BULBOSUM L. Turnip-rooted chervil. 2n=22. Europe and W. Asia. Its cultivation as a vegetable is on the decline. DAUCUS CAROTA L.White carrot, Orange carrot. 2n=8. Afghanistan (p. 86). The origin of the white type isnot clear. It probably arose as a mutant from a yellow type, most likely in France. The orange carrot probably originated inthe Netherlands. This type of carrot is now cultivated widely by peoples of European stock. It has suppressed the growth of the purple carrot, which colours soups and food preparations purple (Banga, 957, 963). The poor storage quality of the purple types (Small, 978b) may alsohave encouraged their replacement by other types. Even before the introduction of domesticated carrots, wild plants were grown in gardens as medicinal crops (W. Brandenburg, pers. comm., 980). LEVISTICUM OFFICINALE Koch. (syn. Angelica levisticum Ball.). Garden lovage, Bladder seed. 2n=22. Cultivated mainly for flavouring. MYRRHIS ODORATA (L.) Scop. Garden myrrh, Sweet scented myrrh. 2n=22. Europe and Caucasia. Cultivated for flavouring and for fodder. PASTINACA SATIVA L. Parsnip. 2n=22. Europe. Var. sativa is cultivated there and elsewhere for its sweet, fresh tap-root. The wild type has a sour root. PEUCEDANUM CERVARIA (L.) Lapeyr. Hart's wort, Much-good, Broad-leaved spignel. 2n=22. S. and C. Europe. Cultivated formerly as a medicinal. PEUCEDANUM OSTRUTHIUM (L.) Koch. Master wort. Pellitory of Spain, Hogfennel. 2n=22. Europe. Cultivated for its scenting root since the 6th Century, as a medicinal and as herb. Its cultivation has almost disappeared now. Valerianaceae VALERIANA LOCUSTA (L.) Betcke. (syn. V. olitoria Pollich). Corn salad, Lamb's lettuce. 2n=. Europe,N. Africa, Caucasia. Cultivated to be used for salads.

150 SALICACEAE - VITIDACEAE VALERIANA OFFICINALIS L. Valerian. 2n=2x=4, 4x=28, 8x=56. Most of Europe and temp. Asia. Subsp. officinalis (syn. V. exaltata Mikan f. ex Pohl), 2n=4, ssp. collins (Wallr.) Nyman (syn.v. collina Wallr.), 2n=28, and ssp. sambucifolius (Mikan f.) Cecak (syn. V. sambucifolius Mikan f., V. exelsa Poiret), 2n=56, are recognized, but other chromosomal numbers occur within a subspecies. These ploidy levels and occasional hybrids have led tomany synonyms (Schrantz, 96). Cultivated for its rhizome which yields the drug valerian. VALERIANA PROCUMBENS Wallr. 2n=56. Spain, Great Britain, France and Germany. Cultivated in Germany. This species might be included in V. officinalis*. VALERIANA SAMBUCIFOLIA Mikan f. ex Pohl. 2n= 56. Wild in N., C. and E. Europe. Cultivated in Thuringia, Germany for its seeds. This speciesmight be included in V. officinalis*. Violaceae VIOLA TRICOLOR L. 2n=26. Europe, Siberia up to Altai and India. Spp. arvensis Gaud. (V. arvensis Murr., 2n=34) is a cosmopolitan weed. This subspecies and spp. tricolor (2n=26) are cultivated for their medicinal and ornamental purposes. Vitidaceae VITISVINIFERA L. Common grape, European grape. 2n~38, (57, 76). For primary centres see p. 02). In Europe, the wild grape played a role in the development of old and modern cultivars. The wild type is declining (Schumann, 977).

151 0 South American Region The SouthAmerican Regionwas restricted totheandesby Vavilov (949/50) and recognized as theandeancentrewhich he divided into two areas: () Peru, Ecuador andbolivia; (2)the island ofchiloe in Chile. The area between the coastofvenezuela, Guyana,Surinam and Cayenne,andS.Brazil and Paraguay was addedby Darlington & JanakaAmmal (945)as athirdcentre. Zhukovskij recognized a megacentre for the whole ofs. America,and Harlan (97)demonstrated the lack of well defined centres oforigin forcultivatedplants in this region. Theoldestknown remainsof cultivated plants ins.america arephaseolus vulgaris fromguitarrero Cave in Peru,datingback some 8000years (Kaplan et al., 973). This,however,isnotnecessarily the centre oforiginof agriculture inthenewworld. Plantswere domesticated ontheandean Highlandsas well as the tropicallowlands (Reed, 977), and a knowledge of food production may have evolved independently in these regions.tuberous crops such asxanthosomawere domesticated inthehumid tropics while otherssuch as Oxalis,Solanum and Ullucus are more typically cropsof thehigh Andes. S.Americaprovided the worldwith numerous fruits and vegetables, but a single cereal, Bromus mango endemic to Chiloe ofchile (Cruz, 972). Maize (Zea mays) was introduced fromc. Americaearly initsevolution intos. America,and thereevolved a major secondary centre of diversity.

152 ACANTHACEAE - ANACARDIACEAE 65 Acanthaceae JUSTICIA PECTORALIS Jacq. 2n=. West Indies and trop. America. Var. stenophylla Leonard semi-cultivated in E. Colombia to adjacent Amazonian Brazil. It is a smaller plant, has smaller and longer leaves, and has shorter inflorescences than the common type. Agavaceae FURCRAEA FOETIDA (L.) Haw. (syn.f. gigantea Venth.). Piteira, Piteira gigante. 2n=(8, 9, 34), 60.S. and C. America.The Mauritius hemp comes from var. willemettiana Roem. which is cultivated on Mauritius and elsewhere. FURCRAEA MACROPHYLLA (Hook.) Baker. Fique. 2n =. Colombia.Cultivated there on a small scale. Some varieties have developed. Fique fibre also comes from other Furcraea species such asf. andina Trel. (2n=60), a wild growing species from Equador and Peru, andt. humboldtiana Trel., a wild growing species from Venezuela. Aizoaceae MESEMBRYANTHEMUM CHILENSE Mol. (syn. Carpobrotus chilensis (Mol.) N.E. Brown). Sea fig. 2n=. Chile. A shrub used in N. America to stabilize dunes. Amaranthaceae AMARANTHUS CAUDATUS L. Inca wheat, Quihuicha. 2n=32, 34. S. America, Asia and possibly in Africa. Cultivated as a grain crop. In Andean region of Peru, Bolivia and NE. Argentina and in China, India, Nepal and Afghanistan. Its leaves are also eaten. A form with red flowerspikes used as a garden ornamental ('love-lies -bleeding') should not be confused with quinoa (Chenopodium quinoa*. Quinoa is of S. American origin (Sauer, 950)). Itresembles A. edulis* and thewild S. American A. quitensis H.B.K., 2n=32. AMARANTHUS DUBIUS Mart, ex The. 2n=64. Trop. America. Cultivated there as a potherb and for its grains. It also is a common weed. It resembles A. cruentus*.perhaps it is a tetraploid from this latter species. See for hybridization with A. spinosus (p. 65). AMARANTHUS HYBRIDUS* AMARANTHUS MANTEGAZZIANUS Passer (syn.a. edulis Spegazzini). 2n=32. Cultivated in Argentina. The wild S. American A. quitensis H.B.K. (2n= 32)closely resembles it.it is also included in A.caudata* as ssp. mantegazzianus (Passer)Hanelt. AMARANTHUS QUITENSIS H.B.K. Sangorache. 2n= Cultivated in Ecuador for its brilliant red inflorescences, which are a source of dye. Selection for bigger flowers (Heiser, 964). Amaranthus caudatus (Sauer, 976). AMARANTHUS SPINOSUS L. Thorny pigweed. 2n=34. S. and C. America. Widespread tropical noxious weed. Cultivated as a vegetable (Mansfeld, 959) in Singapore. Because of the spines it is unlike any of the grain amaranths (Sauer, 950). Where it grows together with A. dubius*sterile hybrids easily arise:a. braunii Thell. and A. caracasamus H.B.K. In general A. spinosus is the female parent. Grant (959) supposed that A. spinosus is one of the parents of A. dubius, but Pal (972) does not support this. Anacardi aceae ANACARDIUM OCCIDENTALE L. Cashew. 2n=42. Trop. America from Mexico toperu and Brazil and also the West Indies. Cultivated now in many tropical countries which may form secondary centres of diversity. Thus Northwood (966) showed the great variation in yield and nut size in the cashew populations in Tanzania. SCHINUS MOLLE L. Californian pepper tree, Brazilpepper tree. 2n=28, 30. Mexico to Chile and Uruguay. Cultivated inthe tropics as a medicinal plant, as a shade tree and as an ornamental. SPONDIAS MOMBIM L. (syn. S. lutea L.). Yellow mombim, Jobo, Hog plum. 2n=32. Trop. America.

153 SOUTH AMERICAN REGION A fruit tree now cultivated in the tropics. SPONDIAS PURPUREA L. (syn.s. mombim L.). Red mombim, Spanish plum. 2n=. Trop. America, C. America and Mexico. A small fruit tree. SPONDIAS RADLKOFERI J. Donn. Sm. 2n=. Lower C. America and Panama. Derived from S. mombin* under selection for late fruiting. Annonaceae ANNONA CHERIMOLA Mill. Cherimoya. 2n=4, 6. Wild in the Andean valleys of Ecuador and Peru. There is itsprimary centre. A small tree. Cultivated now inthe tropics. Its karyotype is similar to that of A. reticulata* and A. squamosa*. Several cultivars are known. Atemoya is a hybrid with A. squamosa*. ROLLINIA DELICIOSA Safford. 2n= tree cultivated for its fruits.. Brazil. A ROLLINIA LONGIFOLIA St. Hil. (syn.r. dolabripetala (Reddi) St.-Hil.) 2n=. Brazil. A tree cultivated for its fruits. ROLLINIOPSIS DISCRETA Safford. 2n= A shrub cultivated for its fruits. Apocynaceae. Brazil. PLUMERIA ACUTIFOLIA Poir. (syn. P. acuminata Roxb., P. obtusa Lour.). 2n=36. Mexico and S. America. Cultivated inthe tropics as a medicinal tree. THEVETIA NEREIFOLIA Juss. (syn. T. peruviana Schum.). Exile tree, Yellow oleander. 2n=20, 22. Trop. America and West Indies. A shrub cultivated in the tropics as a medicinal plant. Aquifoliaceae ILEX PARAGUENSIS D. Don. (syn. I. paraguariensis St.-Hil.). Paraguay tea, Yerba maté.2n= 40. S. America. Cultivated for its leaves which areused to prepare tea. Dispersion ( ) and cultivation of Ilex paraguensis ( )(Patiîïo, 968). Araceae XANTHOSOMA ATROVIRENS C. Koch et Bouche (syn. X. peregrinum Griseb.). Yautia Amarilla, Nut Edo., 2n=. Wild in the llanos of Venezuela. Widely cultivated for its tubers in the Antilles and N. South America. It is characterized by intensely dark green upper leaf-blades, and tubers with yellow flesh. A race grown in Cuba has small tubers, and the race temba taiaof Brazil is characterized by soft tubers. A race grown in Puerto Rico with somewhat hairy leaves is known as jengibrilla. XANTHOSOMA BELOPHYLLUM (Willd.) Kunth. (syn. Caladium belophyllum Willd., X. versicolor Hort, ex Schott). Ocumo, Carouany. 2n=. Cultivated in the coastal mountains of Colombia and Venezuela, and east to Guyana. The tubers of this species is short and stocky,with white flesh. A race, d'espinagas (spinach) is grown in the mountains of Venezuela and adjacent Colombia for its edible leaves. XANTHOSOMA BRASILIENSE Engler. Belember, Herbe a Calalou. 2n=. Wild in S. Brazil. Cultivated for its leaves that are cooked as a vegetable in Puerto Rico and other Caribbean islands. XANTHOSOMA CARACU C. Koch et Bouche. Rolliza, Yautia blanca (Puerto Rico), Lampaza, Rejolgar (Mexico), Manola (Jamaica), Malanga (Cuba), Ocumo (Venezuela). 2n=. Widely grown, also in Africa. Tubers are almost cylindrical with white or rarely orange flesh. XANTHOSOMA MAFAFFA Schott, (syn.x. blandum Schott, X. poeppigii Schott). Mafaffa. 2n=. Tropical forests of Colombia, Brazil, Peru and Bolivia. This species is typically a crop of the wet tropics. It is grown for its tubers in the Amazon Basin of Colombia. XANTHOSOMA SAGITTIFOLIUM (Linn.) Schott, (syn. X. edule (Mey.) Schott., X. xanthorrhizon (Jacq.)C. Koch, Arum sagittaefolium Linn.). Yautia palma. 2n=24, 26. Mountains of Venezuela. Widely cultivated in the Antilles and S. America. Stem up to m tall bearing tubers with white flesh. In Jamaica, it is sometimes known as Yautia panama, although probably introduced to the Caribbean from S. rather than C. America. XANTHOSOMA UNDIPES (C. Koch et Bouche) C. Koch (X. jacquinii Schott.). Yautia palma, Malanga, Chau milon, Grande tayove.2n=. This species resembles X. sagittifolium in developing and aerial stem. But, the leaves are purplish rather than green, and the tubers are white or orange inside. Yautia palma is cultivated, and occurs as a weed from C. Mexico throughout the Antilles to Equador. XANTHOSOMA VIOLACEUM Schott, (syn.x. nigrum Veil.). Jamaica tanier (Trinidad), Oto (Pana-

154 ANACARDIACEAE - CACTACEAE 67 ma), Pica uncucha (Peru), Ocumo (Venezuela), Yautia morada, Prieta (Puerto Rico), and Malanga (Cuba). 2n=24, 26. Widely grown in the Antilles, C. and S. America.The species is characterized by its usually violet pink to purple leaves,and tubers with white flesh but usually purplish outside. Basellaceae BOUSSINGAULTIA CORDIFOLIA Ten. (syn. B. baselloides H.B.K.). Madeira vine, Mignonette vine. 2n=c. 20, 36.S. and C. America. Cultivated as a leafy vegetable or for its tubers. ULLUCUS TUBEROSUS (Lindl.) Lozano. Ulloco (Peru, Bolivia), Chiqua (Colombia), Melloco (Equador), Timbo (Venezuela), Papa lisa (Spanish). 2n=24. Cultivated for its tubers from Venezuela to Argentina. Its wild ancestor, U. a- borigineus Brücher 2n= is a high Andean species of Argentina and Chile. Tubers are eaten fresh or dehydrated, and insome Andean villages ulloco ranks second as a staple only to potatoes. It is beautifully depicted on pottery of the Tiawanako culture (Leon, 964, illustrations). Tubers are of various colours and sizes, and numerous local races are recognized. Bixaceae BIXA ORELLANA L. Annato. 2n=4, 6. Trop.America and the West Indies. Introduced into many other tropical countries where it may have run wild. It is a dye crop. Bombacaceae QUARARIBEA CORDATA (H. & B.) Garcia-Barriga & Hernandez (syn. Matisia cordata H. & B.) South American sapote.2n=. NW. South America. Within this region this fruit is cultivated. No superior strainshave been developed yet. Bromeliaceae ANANAS C0M0SUS (L.) Merr. (syn. A. sativus Schult.f., Bromelia comosa L.). Pineapple.2n- =50, (75, 00). It is suggested that the Tupi- Guarani domesticated pineapple in the Parana- Paraguay river drainage area and that from this region pineapple was spread to all (sub)- tropics. However, Brücher (97) suggested that the domestication of pineapple might have taken place in the highlands of Guyana and a- longside the rivers there. Inthe first area wild related species A. bracteatus (Lindl.) Schultes (2n= ), A. ananassoides (Bak.) L.B. Smith (2n= )and Pseudananas sagenarius (Arudda) Camarcq. (2n= ) occur. A. bracteatus var.typicus is occasionally cultivated for its fruits, while A. ananassoides var. nanus is an ornamental. ANANAS PARGUAZENSIS Card.-Cam. & Smith. 2n=. This species occurs where the Rio Par- V T \ \ ^ ~ (^ i \ ^ \ r ]V / v / ~~ ~* ( i f \ \ \\, ) ^ """"'^ ) r~~i ^,/ V-ï r \ _ N \ ' r I t > } f I ---iv ' / ; ' / > y - / / ^~y Putative area of domestication of Ananas comosus (Pickersgill, 976). guazo discharges into the Rio Orinoco, Venezuela. Brücher (97) suggested that primitive fibre and fruit cultivars have been selected. This selection work could have been carried out - independently of each other - in the region Guyana-Orinoco, and between Maranhao and Pernambuco. PSEUDANANAS MACRODONTES (Harms) Morr. 2n=c. 00. Argentina and Brazil. There its primary centre is found. Cultivated on a large scale on Polynesian and Melanesian islands. Cactaceae CEREUS HEXAGONUS (L.) Mill. 2n=. Venezuela, West Indies, N. South America. Cultivated there for its fruits and as a hedge plant. CEREUS PERUVIANUS (L.) Mill. 2n=24. Probably S. America (Hammer, 976). Cultivated in the tropics as a hedge plant and ornamental. ERIOCEREUS MARTINII (Lab.) Riccob. 2n=. Cultivated in Argentina as source of an alkaloid. HYLOCEREUS POLYRHIZUS (Weber) Britt. & Rose. 2n=. Equador, a hedge plant. OPUNTIA BOLDINGHII Britt. & Rose.2n= NW. coast of Venezuela, Trinidad and Curacao. Cultivated in Venezuela and on Curacao as a hedge plant and for its fruits. OPUNTIA ELATIOR Mill. 2n=. Panama, Colombia, Venezuela. Cultivated there and in Mexico, India, Indonesia and Australia for its fruits. The wild types bearmany long spines, whereas inthe domesticated types fruits are naked or bear a few short thorns (Hammer, 976). OPUNTIA EXALTATA Berger.2n=. Peru. Cultivated as a hedge plant, and there and elsewhere as an ornamental. OPUNTIA VULGARIS Mill. 2n= 22, 33, 66. Brazil,

155 SOUTH AMERICAN REGION Uruguay, Argentina and probably Paraguay, where it may have runwild (Hammer, 976). Cultivated as a hedge plant in India, S. Africa and Australia. In Australia it has run wild. Formerly cultivated in India,Sri Lanka, Indonesia and E. Africa for cochenille. PERESKIA ACULEATA Mill. Barbados cherry, Sweet Mary,West Indian gooseberry, Lemon vine.2n= 22. Trop. America. Cultivated for its fruits and as a hedge plant. Naturalized in Florida, USA.. Brazil. Cul PERESKIA BAHIENSIS Gurke. 2n= tivated in hedges. PERESKIA BLEO (H.B.K.) DC. 2n=. Colombia, Panama, Venezuela and Brazil. Cultivated there and elsewhere. PERESKIA GUAMACHO Weber. 2n=. Venezuela, Marguerita Island. Cultivated there and on Curacao for hedges (Hammer, 976). PERESKIA SACHAROSA Griseb. 2n=22. Paraguay and Argentina. Cultivated there for hedges. RITTEROCEREUS DEFICIENS (Otto & Dietr.) Backeb. 2n=. Venezuela and Curacao. Hedges on Curacao. RITTEROCEREUS GRISEUS (Haw.) Backeb.2n= Venezuela. Cultivated in trop. America and Mexico for hedges and fruits. TRICHOCEREUS BRIDGESII (Salm-Dyck) Britt. & Rose. 2n=. Bolivia. Cultivated there as hedge plant. TRICHOCEREUS CUZCOENSIS Britt. & Rose. Peru. Cultivated there as a hedge plant. TRICHOCEREUS PACHANOI Britton & Rose.2n= Andean parts of Ecuador and Peru. Apparently widely cultivated throughout the C. Andes (Schultes & Hofmann, 973). Cannaceae CANNA EDULIS Ker. Achira,Queensland arrowroot. 2n=8, (27). Probably NW. South America. Spread to Mexico, C. America, West Indies and N. South America. Achira iscultivated inthe West Indies, Australia, S. America, parts of Asia and Pacific Islands. Remains of achira have been found at Huaca Prieta,N. Peru (Bird, 948). They have been dated c BC. It could not be established whether they had been collected or cultivated. Mukherjee & Khoshoo (97)suggested that the triploid (2n=3x=27) is probably an intervarietal hybrid involving genetically related varieties. It isvigorous and robust and has large rhizomes. In S. America rhizomes of other Canna species (C. coccinea Mill., C. paniculata R. & C. and C. indica L.) have been collected and eaten (Gade, 966). Caricaceae CARICA CANDAMARCENSIS Hook.f. (syn. C. pubescens Lenne & Koch). Mountain papaya. 2n=. The Andes of Colombia and Ecuador. A tree cultivated there and also in E. Africa for its fruits (Mansfeld, 959). CARICA CHRYSOPETALA Heilb. 2n=. Ecuador. A tree cultivated for its fruits (Mansfeld, 959). Badilla (967) suggested that this species is a natural hybrid product of C. candamarcensis* and C. stipulata Badilla from Ecuador. He further suggested that this species, C. pentagona* and C. frutifragans Garcia & Hernandez another hybrid of the same parents (from Colombia) should be grouped in C. x heilbornii Badilla. CARICA PENTAGONA Heilb. 2n=. Ecuador. A tree cultivated for its fruits (Mansfeld, 959). Badilla (967) suggested that this species is a natural hybrid product of C. candamarcencis* and C. stipulata Badilla from Ecuador. Caryocaraceae CARYOCAR NUCIFERUM L. 2n=. Brazil and Guyana. A tall tree cultivated inthe West Indies for its edible Suari nuts. Chenopodiaceae CHENOPODIUM PALLIDICAULE Aellen. Canihua. 2n= 36. Andes. Cultivated on the Alti Piano of Peru and Bolivia as a marginal grain crop (Dale, 970). CHENOPODIUM QUINOA Willd. Quinoa, Andean grain chenopod. 2n=allo 4x=36. Andes. Cultivated in the Andes as a grain crop. This cultivation is on its decline. Theweedy C. hircinum Schrad. (sensu Aellen), 2n=36 forms with quinoa a weedcultigen complex (Wilson, 976). Quinoa is closely related to C. nuttalliae*, which may derive from imported quinoa. Chrysobalanaceae CHRYSOBALANUS ICACOL. Icaco plum, Coco plum. 2n=. (Sub)trop. America.Cultivated for its fruits. Compositae EUPATORIUM TRIPLINERVE Vahl. (syn. E. ayapana Vent.). 2n=5. Trop. America. A perennial herb introduced in Java where it is cultivated as a medicinal plant. MADIA SATIVA Molina. Madia, Tarweed. 2n=32. Cultivated formerly in Chile as an oil-seed crop. Attempts have been made togrow it elsewhere, but without success. The culture is almost extinct now.

156 CACTACEAE - EUPHORBIACEAE 69 POLYMNIA SONCHIFOLIA Poepp. & Endl. (syn.p. edulis Weddell.)- Yacon strawberry. 2n=60. Andes. Cultivated there and elsewhere for its tubers. SPILANTHES OLERACEA L. (syn. S. acmella Murr.). Para cress, Brazilian cress. 2n=4, 24, 52. Brazil, West Indies and also India. Cultivated as a vegetable or salad. STEVIA REBAUDIANA Bertoni. 2n=22. N. Paraguay. Annual and biennial. The leaves contain stevioside, which is 300 times sweeter than canesugar. However it acts female sterilizing. TAGETES MINUTA L. Marigold. 2n=. Trop. A- merica. Spread to many other countries. Cultivated for its medicinal properties (Neher, 968). It may reach a height of 3 m or more when cared for. Has runwild in France, Yugoslavia and Greece. Convoivulaceae SCIRPUS CALIFORNICUS (C.A. Meyer) Steudel. (syn. S. tatara Kunth). Totora. 2n=64, 68, 70. Widely distributed in the Americas, on Easter Island and Hawaii. Material of wild plants are used for many purposes like making rafts,house and roofs, and as food for man and animals. Sometimes cultivated in Peru (Heiser, 978). Dioseoreaceae DIOSCOREA PIPERIFOLIA Humb. & Bonpl. Brazil. Cultivated there. 2n= DIOSCOREA TRIFIDA L.f. Cush-cush yam, Yampi. 2n=54, 72, 8. S. America and Antilles, cultivated throughout Caribbean. A racewith purple tubers is grown on Puerto Rico. e-i/ MERREMIA MACROCARPA (L.) Roberty. 2n=. Brazil andwest Indies. Cultivated for its medicinal tubers. MERREMIA TUBEROSA (L.) Rendle (syn. Ipomoea tuberosa L,). 2n=30. Brazil, West Indies, trop. Africa and India. Origin is unknown. Cultivated as a medicinal and also as an ornamental. It may have spread from West Indies and Brazil because in these areas M. macrocarpa* grows wild and is cultivated. Cruciferae LEPIDIUM MEYENII Walp. Maca. 2n=. Peru and Bolivia. Cultivated in Peru for its root. A relic crop. Cucurbitaceae CUCURBITA MAXIMA Duch. ex Lam. Pumpkin, Winter squash. 2n=40. Cultivated all over the world. Secondary gene centre in India and adjacent areas (p. 72). Whitaker & Davis (962) suggested a common origin for C. maxima, C. ficifolia*,c. moschata* and possibly C. pepo* and C. mixta* fromc. lundelliana Bailey (2n=40). C. lundelliana grows in S. Mexico, Guatemala and Honduras.From this parent, C. maxima developed in N. Argentina, Bolivia and S. Peru. In this area the related species C. andreana Naud. grows wild. It is probably a weedy derivative of the cultigen. The wild C. ecuadorensis Cutler & Whitaker (2n=40)is closely related to this species and C. andreana (Cutler & Whitaker, 969). SICANA 0D0RIFERA (Veil.) Naud. Casa banana, Curaba. 2n=. Peru, Brazil to Mexico and West Indies. This vine is cultivated in trop. America. Cyperaceae if.. / N - ^^^v,- ^ " y \ K Dioscorea trifida (Coursey, 967). Erythroxylaceae ERYTHROXYLUM COCA Lam. Coca, Guarigos. 2n=24. Unknown wild. Probably from high Andes of Peru and Bolivia. Cultivated at high altitudes in Peru, Bolivia, Argentina, Colombia and Brazil. Some taxonomists include E. novogratense*, E. truxillense Rusby and E. bolivianum Burck in this species. ERYTHROXYLUM NOVOGRANATENSE (Morris) Hieron. Truxillo coca. 2n=. Andes. Cultivated at a lower altitude than E. coca*. It was distributed to the tropics. Euphorbiaceae HEVEA BENTHAMIANA Muell.-Arg. 2n=36. The Amazone basin, Brazil, Peru and Bolivia. A tree cultivated for its rubber. HEVEA BRASILIENSIS (Willd.) Muell.-Arg. Brazilian hevea, Para rubber tree. 2n==36. The Amazon basin. This is the primary gene centre. Secondary gene centre in Malaya (p. 52). Cultivated now in Malaya, Indonesia, Sri Lanka and in some other countries. In Africa rubber has been cultivated as a farmer's crop and as kas, 969).

157 72 SOUTH AMERICAN REGION SOUTH AMERICAN REGION DESMODIUM DISCOLOR Vog. plant. 2n=22. Brazil. A forage DESMODIUM INTORTUM (Mill.) Urb. Greenleaf. 2n= 22. C. America and Brazil. Cultivated in Australia (Hutton, 970). DESMODIUM UNCINATUM (Jacq.) DC. Silverleaf. 2n =22. S. America.Cultivated in Australia (Hutton, 970). DIPTERYX ODORATA Willd. Tonka bean, Dutch tonka. 2n=32. Forests of trop. America, Venezuela, the Guyanas and lower Amazon basin. The tree is cultivated now in Venezuela, Malaya, West Indies and some other tropical countries (Cobley, 963). ERYTHRINA GLAUCA Willd. 2n=42. S. shade tree in cacao plantations. ERYTHRINA MICROPHERYX Poepp. Anauca. 2n= Peru. A shade tree in cacao plantations. America. A GLIRICIDIA SEPIUM (Jacq.) Steud. (syn.g. maculata Benth.). 2n=20, 22. Mexico,C. America and N. South America. A shade tree,green manure and fodder crop. INDIGOFERA ANIL L. (syn.i. suffruticosa Mill.). Indigo plant. 2n=2. S.America. Once much cultivated in the tropics for its dye (indigo) (Heiser, 965). INGA FEUILLEI DC. (syn. I. reticulata Spr.). 2n=. Peru. This tree is cultivated there for sweet fruit pulp (Uphof, 968). INGA PREUSSII Harms, as ashade tree. INGA PUNCTATA Willd. 2n= Used as a shade tree. LEUCAENA LEUCOCEPHALA* 2n= El Salvador. Used S. and C. America. LONCHOCARPUS UTILIS Smith. 2n=44. Peru. Cultivated as asource of rotenone.. Bolivia. Cul LUPINUS B0G0TENSIS Benth. 2n= tivated there. LUPINUS MONTANUS H.B.K. 2n=. Peru, Bolivia, Guatemala and Mexico. Cultivated in Bolivia. LUPINUS MUTABILIS-TAURIS-CUNNINGHAMII-CRUCK- SHANKSII species group. 2n=48. Andes region between Bolivia and Venezuela. This group of species, also named L. mutabilis Sweet/L. tauris Hook, isnot yet well described. Formerly widely cultivated in its native centre. Still cultivated in Bolivia. Apparently farmers have not tried toselect for sweet types. At present this species is used as 'bitter protection rows' around fields of Vicia fabaand Pisum sativum tostop animals entering fields. Types have been developed that grow well un- der tropical short-day conditions, that have pods which do not open and soft-coated seeds rich in protein and of low alkaloid content (Brücher, 970; Hackbarth & Pakendorf, 970). Thebig seeds are used toprepare tarwi or ullu. Other wild, but possible valuable Lupinus -species of the American continents should be domesticated. They are often shrubby and have small seeds (Brücher, 970). MIMOSA INVISA* MYR0XYL0N BALSAMUM (L.) Harms. Balsam of Peru. 2n=28. Var. pereira (Royle) Harms is spread in Guatemala and El Salvador. It is a source of balsam. It was cultivated in the imperial gardens of the Aztecs in Mexico (Mansfeld, 959). PACHYRHIZUS APIHA (Wedd.) Parodi. 2n=22. Probably a cultigen developed by the indians in Bolivia and N. Argentina. PACHYRHIZUS TUBEROSUS (Lam.) Spreng.Yam bean, Potato bean, Jicama. 2n=22. The headwaters of the Amazon.From there itwas distributed to other parts of S. America and parts of the West Indies. The young pods and tubers are eaten. PHASEOLUS ABORIGINEUS Burk. 2n=22. Forests of NW. Argentina Andes. Probably extended through Bolivia, Peru, Ecuador up to Honduras (Burkart & Bücher, 953). It might be the progenitor ofp. vulgaris*in Peru (Heiser, 965). PHASEOLUS LUNATUS L. Lima bean, Sieva bean, Butter bean,madagascar bean, Burma bean. 2n= 22. C. America, and in the Andes from Peru to Argentina. Kaplan (965) showed that the big lima bean of Peru was first domesticated in the Andean highlands and that the small lima bean of Mexico may have arisen in the Pacific coastal foothills of Mexico (p. 93). A smallseeded subspecies (ssp. microsperma, Sieva or Small Lima) originated by natural selection. It spread to the Antilles. PHASEOLUS VULGARIS L. Common bean. 2n=22. For origin see p. 94. The earliest remains of cultivated common beans have been found in the Guitarrero Cave in Peru. It dates from about 6000 BC. The 'domesticated' characters are especially dark red brown and dark red beans (Kaplan et al., 973). PITHECELLOBIUM SAMAN Benth. Rain tree, Saman, Cow tamarind. 2n=26. Trop. America. It is used as ashade tree in cacao and coffee plantation. PROSOPIS CHILENSIS (Molina) Stuntz emend. Burkart. 2n=. From Peru and Bolivia to C. Chile and NW. Argentina. An unarmed tree suitable for domestication as a shade, timber and fuel tree, and for its sweet pulpy fruits eatenby man and cattle (Burkart, 976). PROSOPIS JULIFLORA DC. Mesquite. 2n=28, 52, 56. Cultivated in Brazil, India and elsewhere as a

158 LEGUMINOSAE -MALVACEAE shade and timber tree,and for forage. The shrubby type is an aggressive invader in Hispaniola and Pakistan (Burkart, 976). PROSOPIS STROMBULIFERA'(Lam.) Bentham. 2n=28. W. Argentina and N. Chile. At one time cultivated in Chile for its fruits, which ease toothache.wild fruits are gathered for this purpose. Var. ruiziana Burkart, perhaps a tetraploid (2n=4x=56) (Burkart, 976), has fruits twice the size of var. strombulifera. PROPOSIS TAMARUGO T. Philippi. Tamarugo tree. 2n=. Tree for desert forestation and for raising Merino sheep and Angora goat, which feed on falling leaves and legumes. The leaves absorb atmospheric moisture (Burkart, 976). RHYNCHOSIA MINIMA (L.) DC. 2n=22. S. America. Cultivated in Australia and elsewhere as a fodder and pasture crop. STYLOSANTHUS GUIANENSIS SW. (syn.s. surinamensis Miq.). 2n=20. Guyana. Used as a food plant for livestock and as asoil conservator. VICIA GRAMINEA Smith. 2n=4. Argentina and Chile. Occasionally cultivated for its seeds as asource of anti-n-lecitin (Nijenhuis et al., 969). This is used as a test serum for the humann-blood group. Malpighiaceae BANISTERIOPSIS CAAPI (Spruce ex Griseb.) Morton. 2n=20. S. America. A woody vine cultivated in the Amazon region as a drug and narcotic. BUNCHOSIA ARMENIACA (Cav.) DC. 2n=. The Andean region. A shrub cultivated in Ecuador for its fruits. MALPIGHIA GLABRA L. (syn. M. punicifolia L.). Barbado cherry, West Indian cherry. 2n= West Indies and N. South America. Cultivated there and elsewhere for its fruits.it also makes agood hedge, like M. coccigera L. (Purseglove, 968). Malvaceae ABUTILON OXYCARPUM F. Von Muell. 2n=4. South, America and Australia. Cultivated for its fibres. GOSSYPIUM BARBADENSE L. (syn. G. vitifolium Lam., G. peruvianum Cav.). Sea island cotton. 2n=52, genome formula (AADD)2. It has been proposed that G. barbadense arose from a cross and amphidiploidization of G. arboreum* and G. raimondii*. G. arboreum could have been introduced into Peru by way of Asia and the Pacific islands. Another hypothesis is that an African diploid reached S. Americaby way of Atlantic. This diploid would probably have been G. herbaceum*. As Bird (948) found G. barbadense material Distribution of the New World cottons in the 3th century: Gossypium barbadense (), G. hirsutum var. marie-galante (2)and G. hirsutism punctatum (3)(Hutchinson,962). at Huaca Prieta, Peru which was dated 2400 BC. the introduction of the African Gossypium species and its amphidiploidization with G. raimondii must have taken place long before that time. The main point is how this African species reached Peru. However, the centre of origin N. Peru is in the arid mountainous interior of the prov. Tumbes. The ssp. darwinii is closely related and is endemic in the Galapagos Islands. At present it is 'contaminated' by hybridization with exotic introductions. Secondary centre in Peru. In S. America G. barbadense spread south and eastwards to NW. Argentina. Some other forms are found ins. America. The Tanguis variety is a selection from Tumbes. In Chile and Peru the Pacific assemblage is found, characterized by broad leaves and intense hairiness of the underside of the leaf. This character induces resistance to jassids, Empoasca ssp. The lint of G. barbadense isusually coarse with a length up to 34.5 mm. The lint of an Ecuador type, of Sea Islands and Egyptian is fine and silky with a length up to 37.5 mm. It is possible that the Ecuador type is the parent of the Sea Islands/Egyptian complex (p. 84, 39) (Harlan, 970). The Atlantic assemblages include the kidney cottons (seeds fused in a kidney-shaped cross). They have a wide distribution in N. South America and the islands of C. America. They have been taken to Africa, India, Sri Lanka, Indonesia and elsewhere. Secondary centres in Egypt (p. 39) and in Turkmenistan - Tadjikistan -S. Uzbekistan, USSR (p. 84). On the Sea Islands of S. Carolina,USA the Sea Island cottons developed after cottons from Bahamas or Jamaica (p. 94) were introduced (Hutchinson, 962).

159 SOUTH AMERICAN REGION V\ 4 ~D) 5 (u «y \ \ * Distribution of annual cottons in the New World at 960: Gossypium hirsutum var. uplands (4), G. barbadense var. Egyptians (5) and G. barbadense var. Sea Islands (6)(Hutchinson, 962). G. mustelinum (7)and G. raimondii (8)(Pickersgill et al., 975). ^ It is a source of resistance to Verticillium. A second important perennial is race punctatum, which is found around the coast of the Gulf of Mexico from Yucatan to Florida and the Bahamas and some other islands (p. 99). At present G. hirsutum Cambodia (a latifolium type) is cultivated in S. India. Their way of spread was probably S. America-Philippines- Cambodia-S. India. GOSSYPIUM KLOTZSCHIANUM Andersson. 2n=26, genome formula D 3_ifD 3_ K. Galapagos Islands. Var. davidsonii is found on the shores of Gulf of California and the Revilla Gigedo islands (p. 39). GOSSYPIUM MUSTELINUM Miers ex Watt. (syn.g. caicoense Aranha, Leitao & Gridi-Papp). 2n= 52, genome formula AADD. NE. Brazil. A wild species distinct from G. hirsutum* and G. barbadense*, and not derived from either of them (Pickersgill et al., 975). GOSSYPIUM RAIMONDII Ulbr. 2n=26,genome formula D gdg. Formerly N. Peru. Now extinct in its original habitat and only found in collections (Harland, 970). This species is a source of hairiness gene Hg conditioning resistance to jassid, Empoasca ssp. Probably one of the parental species of G. barbadense* and G. hirsutum. GOSSYPIUM HIRSUTUM L. Upland cotton. 2n=52, genome formula (AADD)^. The common theory is that G. hirsutum arose from an amphiploidization of the Old World G. arboreum* org. herbaceum* and G. raimondii.g. raimondii isthe parent of the D genome and one of the first two the donor of the D genome. It is not known when this amphiploidization took place, but material collected from Tehuacan Valley in Mexico and dated BC. appears to be fully domesticated (Smith & Stephens, 97). It is also not known whether G. barbadense reached Peru by way of Asia or whether G. herbaceum reached eastern S. America from W. Africa. Harland (970)observed wild plants of an exceedingly primitive perennial race marie galante in the state Rio Grande do Norte, N. Brazil. He suggested that this area is almost certainly the centre of origin of the whole Upland group. From here this cotton dispersed first northward to the Amazon, then along the Amazon and across the Andes into Ecuador, W. Colombia and possibly still further north. In another direction this cotton dispersed northward through the Guyanas passing thewest Indies to E. Colombia and further northward into C. America via Yucatan. C. America must be considered as a secondary centre of diversity (p. 94). Wild and semi-wild marie galante cotton were observed in Florida until some years ago. Upland cotton also dispersed southward to E. Brazil. At present this race is also grown in Ghana. Gossypium raimondii GOSSYPIUM T0MENT0SUM Nutt. ex Seem (syn. G. sandvicense Pari.). 2n=52, genome formula (AA DD)i. Hawaii. A fuzzy-seeded but not limited species. At one time it was believed that if the origin and relationship of this species were elucidated the problem of the origin of G. barbadense* and G. hirsutum* could be solved. However it appears that its origin is in-

160 MALVACEAE 77 dependent of those of the New World species (Hutchinson, 962). WISSADULA CONTRACTA (Link.) R.E. Fries.2n= 4. Trop. America. Cultivated in W. Java for fibre (van Borssum Waalkes, 966). WISSADULA PERIPLOCIFOLIA (L.) Presl ex Thw. 2n=4. Probably introduced insri Lanka as a source of fibre for which purpose it is still used (van Borssum Waalkes, 966). The degree of variability of this species is very small in Malaysia. Some varieties and forms have been described for American representatives. This may point to an American origin. A pantropical weed. Marantaceae CALATHEA ALLOUIA (Aubl.) Lindb.Sweet corn root, Leren.2n=. Native to Hispaniola, Puerto Rico, Lesser Antilles, the Guyanas, Brazil, Venezuela, Colombia, Ecuador and Peru. Introduced into Asia. A minor tuber crop. MARANTA ARUNDINACEA L. Arrowroot, Bermuda arrowroot. 2n=8, 48. N. South America and the Lesser Antilles. Cultivated in the tropics for its rhizomes containing starch and as a medicinal crop. Musaceae MUSA cultivars. 2n=22, 33. Not much is known yet about the distribution of the clones of the various genome groups. In Venezuela, the relative frequency of these groups isaa, AAA 8, AAB c. 0 and ABB c. 3 (Borges, 972). Trop. America is a secondary centre for the Plantain subgroup French Plantain or Horn Plantain, 2n=33, AAB. Myrtaceae ABBEVILLEA FENZLIANA Berg. 2n=. Brazil. A small tree cultivated for its edible fruits. BRITOA ACIDA Berg. Para guava. 2n= A shrub cultivated for its fruits.. Brazil. CAMPOMANESIA GUAVIROBA Benth. & Hook. 2n= S. Brazil. Cultivated for its edible fruits. CAMPOMANESIA LINEATIFOLIA Ruiz. & Pav. (syn. C. cornifolia H.B.K.). 2n=. E. Andes. Cultivated as a fruit tree in Peru (Mansfeld, 959). EUCALYPTUS CAMALDULENSIS Dehn.Longbeak eucalyptus. 2n=22. Primary centre: Australia (p. 66). Secondary centres: Brazil, Argentina and the Mediterranean area (p. 7). EUGENIA DOMBEYANA DC. Grumichama. 2n=. Peru and S. Brazil. A tree cultivated for its fruits EUGENIA UNIFLORA L. Pitange, Surinam cherry. 2n=22. Brazil.Cultivated in the tropics and subtropics. EUGENIA UVALHA Camb. Uvalha. 2n= Cultivated for its fruits. S. Brazil. FEIJOA SELLOWIANA Berg. Feijoa. 2n=22. S. Brazil, Uruguay, Paraguay and N. Argentina. Also its primary centre of diversity. Sometimes cultivated for its fruit in hot countries e.g. the Caucasian coast of the Black Sea where it grows well. MYRCIARIA CAULIFLORA Berg. (syn. Eugenia cauliflora (Berg.) DC). Jabotica.2n=. Brazil. Cultivated for its fruits (Purseglove, 968). MYRCIARIA JABOTICABA Berg. 2n=. Brazil. Cultivated in the tropics for its fruits. PSIDIUM GUINEENSE SW. 2n=. The West Indies and trop. America. Occasionally cultivated. PSIDIUM LITTORALE Raddi (syn. P. cattleianum Sabine). Strawberry guava. 2n=88. Brazil. A small tree introduced in the tropics and subtropics. Var. lucidum Degener, Chinese strawberry guave yields fruits of improved quality (Uphof, 968). Nyctaginaceae MIRABILIS JALAPA L. Marvel of Peru, Four o'clock, False jalap. 2n=(54), 58. S. America. Spread over the whole world and inw. Africa as a fetish plant.cultivated as an ornamental. Tuberous roots were used as jalap. Elsewhere a subtropical weed. Onagraceae FUCHSIA MAGELLANICA Lam. Fuchsia. 2n=22, 44. S. America. Planted as hedges in Azores, Ireland and W. Britain. Oxalidaceae OXALIS TUBEROSA Mol. Oca. 2n=(4), 60, 63-64, Cultivated in the Andes from Colombia to Bolivia for an extremely long time.introduced also in Europe where it was cultivated like the Mexican O. deppei Lodd. (2n=4, 56) as a vegetable by amateurs (Uphof, 968). Several colours of the tubers have been observed. Itshould not be confused with Tropaeolum tuberosum*. Palmae COPERNICIA PRUNIFERA Moore. Carnauba wax palm. 2n=36. Brazil. Mostly semi-wild; some plantations for wax production. C0R0Z0OLEIFERA (H.B.K.) Bailey, (syn. Elaeis melanococca Gaertn.). Nolipalm. 2n=32. C. A- merica to Colombia and Amazon area. Cultivated for its oily fruits. It canbe crossed with

161 78 SOUTH AMERICAN REGION the African oil palm, Elaeis guineensis* ducing fertile hybrids. GUILIELMA GASIPAES (H.B.K.)L.H. Bailey. Peach palm, Peribaye. 2n=. S. and C. America. Cultivated in S. America. OENOCARPUS BACABA Martius. 2n=. Amazon a- rea to Surinam and Guyana. A palm cultivated on compounds for its oily fruits. Passifloraceae PASSIFLORA ALATA Dryand. Winged passion flower, Maracuja. 2n=. Peru and Brazil. A woody vine cultivated in Brazil for its fruits. PASSIFLORA ANTIOQUIENSIS Karst, (syn.p. vanvolxemii (Lem.) Triana & Planch.). 2n=. Banana passion fruit. Colombia. A woody vine cultivated e.g. in New Zealand for its fruits. P. x militaris hort. derives from P. antioquiensis xp. manicata (Juss.) Pers., 2n=8, andp. x exoniensis Bailey derives from P. antioquiensis xp. mollissima* (Ohle, 975). PASSIFLORA CAERULEA L. 2n=8. Blue passion flower. Brazil, Argentina, Paraguay. Cultivated for its fruits and as an ornamental. PASSIFLORA COCCINEA Aubl. 2n= livia, Amazon district of Peru. pro- Brazil, Bo- PASSIFLORA EDULIS Sims. Passion fruit, Purple granadilla. 2n=8. S. Brazil. Widely distributed throughout the tropics and subtropics. The fruits are especially used for juice preparation.the mountain form. f. edulis occurs from Brazil to N. Argentina and in the tropics. It has run wild in Assam. The lowland form flavicarpa Degener is a mutant of it (Ohle, 975). PASSIFLORA FOETIDA L. Stinking passion flower, Love-in-a-mist. 2n=8, 20, 22. West Indies and S. America. Weedy. Distributed to many tropical countries in Africa and Asia, where it has naturalized. Its fruits are sometimes eaten. In Malaya and E. Africa it has been used as a cover crop. PASSIFLORA LAURIFOLIA L. Water-lemon, Golden apple, Yellow grandilla, Jamaica honeysuckle, Belle apple, Pomme de liane. 2n=8. Thickets and forest fringes in the West Indies and NE. South America. Cultivated for its fruits in the 7th Century. Spread throughout thetropics (Purseglove, 968). Cultivated on Java and in India as an ornamental and as a medicinal plant. PASSIFLORA LIGULARIS Juss.Sweet granadilla. 2n=8. Trop. America. Spread toc. America. Its sweet fruits aremuch used in the mountainous regions of Mexico and C. America (Purseglove, 968). PASSIFLORA MALIFORMIS L. Curuba. 2n=8. Trop. America. A vine cultivated for its fruits. PASSIFLORA MIXTA L.f. 2n=8. Venezuela, Colombia, Ecuador, S. Peru,P. x rosea (Karst.) Killip, 2n=. Derives from P. mollissima* x P. mixta. PASSIFLORA MOLLISSIMA (H.B.K.) Bailey. Banana passion fruit, Tasco, Caruba de Castilla. 2n= 8. The Andes. Especially cultivated in Ecuador and Bolivia. Introduced in other countries. PASSIFLORA PINNATISTIPULA Cav. 2n=. Andes. Cultivated Colombia, Chile and Peru. It hybridizes with P. mollissima*. PASSIFLORA P0PEN0VII Killip. 2n=. Andes (Ecuador). Resembles P. laurifolla* (Ohle, 975). PASSIFLORA PSILANTHA (Sodiro) Killip. Gullan. 2n=. Ecuador. A hybrid of P. mollissima* x P. partita* (Ohle, 975). A vine cultivated for its fruits. PASSIFLORA QUADRANGULARIS L. Giant granadilla, Barbadine. 2n=8. Trop.S. America. Cultivated since 8th Century for its fruits. Now widely distributed in the tropics. Possibly a hybrid (Ohle, 975). PASSIFLORA SERRATO-DIGITATA L. (syn. P. cearensis Barb.). 2n=. Trop. S. America, Amazon Basin. PASSIFLORA TRIPARTITA (Juss.) Poir. Tasco.2n= 8. Ecuador. Cultivated there. Peperomiaceae PEPEROMIA PELLUCIDA H.B.K. 2n=. S. America. In Africa this pantropical weed is cultivated as a vegetable and medicinal crop. Phytolaccaceae PHYTOLACCA CHILENSIS Miers.2n=. Chile. A perennial herb cultivated for its berries which are asource of red dye. PHYTOLACCA DIOICA L. 2n=36. Temp, and subtrop. S. America.Cultivated as an ornamental and shade plant. RIVINA HUMILIS L. Rouge plant. 2n=08. The tropics of the Old and New Worlds. Cultivated in Colombia for its berries which are a source of red dye. Piperaceae PIPER ADUNCUM L. 2n=. Trop. America. Used as asoil conservant. Portulacaceae TALINUM TRIANGULÄRE*

162 SOLANACEAE 79 Rharanaceae COLUBRINA RUFA Reiss.2n=. Brazil. Cultivated for its medicinal bark and other purposes. Rosaceae FRAGARIA CHILOENSIS L. Chiloe strawberry, Beach strawberry, Ambato strawberry. 2n=56, genome formula AAA'A'BBBB. The Pacific coastal region ofn. and S. America and Hawaii. Dioecious; polygamodioecious or hermaphroditic types are occasionally found. Formerly cultivated there. It is one of the parents off. x ananassa*. RUBUSBRASILIENSIS Mart.2n=. Brazil. A shrub cultivated for its fruits. RUBUS GLAUCUS Benth. 2n=. Costa Rica to Ecuador. Cultivated in the Andes. RUBUS MACROCARPUS Benth. Colombian berry. 2n=. Colombia and Ecuador. Cultivated for its very large fruits (5cm long). Rubiaceae CEPHAËLIS IPECACUANHA (Stokes) Bai. Ipecac, Ipecacuanha. 2n=22. Brazil. Introduced into India and Malaya. There small plantings were established. Roots ofwild and cultivated plants are the source of ipecac or ipecacuanha used to treat amoebic dysentery. CINCHONA LEDGERIANA Moens ex Tremen (syn. C. calisaya var. ledgeriana How., C. officinalis L., C. calisaya Wedd. and C. succirubra Pav. ex Klotzsch.). Quinine. 2n=34 (all species). These species are taken together. They all come from the same centre of diversity: Andes mountains of S. Peru, Bolivia and S. Ecuador. Here many Cinchona species are found and the great diversity of botanical varieties is caused by natural hybridization between the species and varieties.plantations in Indonesia and Sri Lanka and recently in E. Africa. The original introductions in the Asian countries were very probably a mixture of true species and their hybrids.from this material C. ledgeriana was derived but it is thought to be a variety of C, calisaya, and is also considered a hybrid of C. calisaya, C. succirubra and C. lancifolia Mutis. C. succirubra which isused as rootstock isprobably avariety of C. pubescens Vahl. (van Harten, 969). SAPINDUS SAPONARIA L. Soap wood tree,soap tree, Soap berry tree. 2n-. Trop. America. Cultivated there and elsewhere for its fruits. Sapotaceae LUCUMA NERVOSA A. DC. (syn.l. rivicoa Gaertn. f., Pouteria campechiana (H.B.K.) Baenhi). Egg fruit, Canistel. 2n=. NE. South America. Cultivated in trop. America for its fruits. LUCUMA OBOVATA H.B.K. (syn. Pouteria lucuma (Ruiz & Pav.)0. Kuntze). Lucumo. 2n=, Chile and Peru. This tree is cultivated for its fruits. LUCUMA PROCERA Mart. (syn. Pierre). Macarandiba.2n : = is cultivated in Brazil. Urbanella procera. This fruit tree MANILKARA BIDENTATA (A. DC.) Chev. (syn. Mini us op s balata Pierre). Balata, Bully, Bullet, Purgio, Quinilla. 2n=. S. America and Trinidad. The wild trees are tapped for latex (balata). POUTERIA CAIMITA (Ruiz & Pav.) Radlk. 2n= Peru to E.Ecuador and Guyanas. A tree cultivated for its fruits. Simaroubaceae QUASSIA AMARA L. Surinam quassis, Bitter wood. 2n=. N. South America. Cultivated for its wood which is used medicinally, and also as an ornamental tree. Solanaceae CAPSICUM BACCATUM H.B.K. (syn. C. angulosum Miller). Pepper. 2n=24. The wild type is var. Sapindaceae MELIOCOCCUS BIJUGATUS Jacq. Kanappy tree, Kinnup tree, Bullace plum, Honey berry, Spanish lime, Geneps. 2n=32. Trop. America. Cultivated there for its edible fruits (Mansfeld, 959). PAULLINIA CUPANA (H.B.K.) Guarana. 2n=. S. America. Cultivated in Brazil for its seeds, used as a coffee. Capsicum bacatum var. baccatun (o)and var. pendulum ( )(Eshbough, 975).

163 80 SOUTH AMERICAN REGION V-' * \ l ~'~ \ [T "" \ ] V^* N r I / ') \ ' V/ ".*. *. ^ * n ) ( \ i. \ / j /,< f / I J l ( i Capsicum eximium ( ) and C. cardenasii (*) (Eshbough, 980) CAPSICUM PUBESCENS Ruiz & Pavon. 2n=24. Cultivated in the highlands of S. and C. America. It is related to thewild self-incompatible C. cardenasii Heiser & Smith, 2n=24, and the wild pseudo-self-compatible C. eximium Hunziker, 2n =24; these species together with C. tovari,2n =24, form the purple-flowered group of Capsicum (Jensen et al., 979). C. cardenasii is an endemic of the La Paz district of Bolivia; C. eximium is widely distributed from N. Argentina to C. Bolivia (Eshbaugh, 977, 980). Both species hybridize naturally and the hybrids are fully fertile. They also cross to a lesser extent with C. pubescens and may form one species complex deriving from a common wild ancestor (McLeod et al., 979). baccatum (syn, C.microcarpum Cav.). It occurs in Peru, Bolivia, Paraguay, N. Argentina and S. Brazil. It is the parental type of the cultivated type var. pendulum (Willd.) Eshbaugh (syn. C. pendulum Willd.). This cultigen was originally found in the same area as var. baccatum and in S. Colombia, Ecuador and in Chile. Now it is also cultivated elsewhere (Eshbaugh, 970). CAPSICUM CHINENSE Jacq. (syn. C. sinense Jacq.). 2n=24. This pepper was originally cultivated in the West Indies and lowland S. America, from S. Bolivia to S. Brazil. Closely related toc. frutescens*. Itmay have originated from it (Pickersgill, 969). \ ~^^ "OC!j \^ 0 o * Capsicum pubescens (Eshbough, 975). ir-. CYPHOMANDRA BETACEA (Cav.) Sendt. (syn. C. crassifolia). Tree tomato. 2n=24. Peru. Unknown wild. Cultivated in the Andean region especially in Ecuador. Other species of this genus are found in S. America and partly in C. America.One of them, isc. hartwegi Sendt.; its fruits are harvested in Colombia, Chile and Argentina. \ * ' l / - * - - '' / '' ~ '/' Capsicum frutescens (o)andc. chinense ( ) (Eshbough, 975) *r * } LYCOPERSICON CHEESMANII Riley (syn.l. esculentum Mill. ssp. minor Rick, L. minutum Rick). Galapagos tomato. 2n=24. Coasts of Galapagos Islands. Characterized by a very dense pubescence, compound yellow-green leaves, yellow or orange fruit (rich in beta-carotens), a calyx that expands after fertilization and seeds with adeep dormancy. The plants are drought and salt tolerant. They are eatenby the Galapagos tortoises and seeds can germinate after passing through their digestive tracts (Rick & Bowman, 96).

164 SOLANACEAE - SOLANACEAE 8 LYCOPERSICON CHILENSE Dun.2n=. The coastal strip of Peru and N. Chile. A wild tomato often found growing together with L.peruvianura. They do not cross. This species is characterized as asource for resistance to all tomato diseases except Phytophthora. LYCOPERSICON CHMIELEWSKII Rick, Kesicki, Fobes & Holle. 2n=. C. and N. Peruvian Andes. Sympatric with its derivative L. parviflorum*. Formerly belonging with this species to 'L. minutum'. Cross-fertilizer (Rick et al., 976). LYCOPERSICON ESCULENTUM Mill. Tomato. 2n=24. The centre of the genus Lycopersicon is a narrow belt of the S. American west coast limited by the equator and 30 S and the Andes and the Galapagos Islands. The greatest variation of the tomato is however outside this area, in the Veracruz-Puebla area in Mexico (Jenkins, 948). This area was very likely the source of the cultivated tomatoes of the Old World. The putative ancestor of the tomato is probably var. cerasiforme (Dun.) Alef. This variety was originally confined to the Peru and Ecuador area from where it spread in pre-columbian times as a weed of fields and compound yards throughout much of trop. America, either with or without man's active co-operation. In Mexico it became cultivated because of its similarity to another food plant, Physalis ixocarpa*. Outside its primary gene centre the tomato plant is self-compatible. In Peru and Ecuador it spontaneously crosses with L. pimpinellifolium*. Tomato flowers pollinated by pollen of L. peruvianum* result inthe induction of parthenocarpic fruits. Some Fi seeds may be set resulting in hybrid plants with varying degree of fertility. Rick (97) studied the geographical distribution of the alleles Ge c, GeP, and Ge n. He found that most European and US cultivars have the genotype Ge n Ge n, only a few have Ge c Ge c or GePGeP. The C. American varieties have Ge n Ge n and occasionally Ge c Ge c. In Ecuador Ge is also common among the cultivars. The C. American sources of var. cerasiforme have Ge n Ge n, and an Ecuador source has Ge c. So in Ecuador the cultivars differ from the wild type, but more sources should be investigated. Sources of L. pimpinellifolium* ex Ecuador carried Ge. This would suggest gene exchange between L. pimpinellifolium and the cultivars. The Peruvian cultivars carry GeP; the same allele is found inperuvian sources of L. pimpinellifolium. This suggests gene exchange too. Rick concluded that the European and US tomato cultivars are qualitatively closer related to the cultivars of Peru, and quantitatively to those from C. America and Mexico. LYCOPERSICON HIRSUTUM Humb. & Bonpl. 2n=24. The western slopes of the Andes in Peru. A green-fruited species. The glabratum is selfcompatible. It is characterized by disease resistance, e.g. tomato mosaic virus (Marmon tabaci Holmes). LYCOPERSICON PARVIFLORUM Rick, Kesicki, Fobes & Holle. 2n=. C. and N. Peruvian Andes. Compatric with its ancestor L. chmielewskii*. Formerly classed with this species to 'L. minutum'. Strict self-fertilizer (Rick et al., 976). LYCOPERSICON PERUVIANUM (L.) Mill. 2n=24. Chile and Peru. A green, small-fruited wild species. Most plants are gametophytic self-incompatible, although some plants have been found to be selfcompatible (Hogenboom, 968). It is asource of tomato mosaic virus tolerance. LYCOPERSICON PIMPINELLIFOLIUM Mill. (syn.l. esculentum ssp. pimpinellifolium (Mill.) Brezhn.). Currant tomato. 2n=24. Primary centre: coastal Ecuador and Peru. This red-fruited species crosses with L. esculentum* and its genes introgress into tomato (Rick & Fobes, 975). Because of the high frequency of outcrossing, it is highly heterogenous (Rick et al., 977). It is cultivated and occurs as a weed. A source of tolerance for tomato. METHYSTICODENDRON AMESIANUM R.E. Schultes. 2n =. S. America.Cultivated as a medicinal and witchcraft plant. NICOTIANA RUSTICA L. Aztec Tobacco, Makhorka, Nicotine Tobacco. 2n=48. Unknown wild, with a possible exception of var.pavonii (Dunal) Goodspeed. This variety occurs as a ruderal in the Andes. Aztec Tobacco is a tetraploid having probably originated in Peruby amphiploidization of apparently N. paniculata L. (2n=24)and N. undulata Ruiz & Pavon (2n=24). Both species occurwild in Peru. Its cultivation is limited to some areas such asthe USSR and India. In most other areas it is replaced by N. tabacum* which has alow nicotine content. NICOTIANA TABACUM L. Tobacco. 2n=48, genome formula SSTT. Clausen (932)showed that tobacco is a natural amphitetraploid of N. sylvestri Speg. & Comes, 2n=24, genome formula S*S' and N. tomentosiformis Goodsp., 2n=24, genome formula T'T'. Isozymic (Sheen, 972) and cytoplasmic (Gray et al., 974; Kawashima et al., 976) evidence supports his view. The occasional wild plants are escapes of cultivation. Interspecific crosses have been made to introduce male-sterilizing cytoplasma and genes conditioning resistance to diseases. PHYSALIS PERUVIANA L. Cape gooseberry. 2n=24, 48. Andes. Cultivated in some S. American countries for its berries. Often observed as a weed or semi-wild. SOLANUM ABANCAYENSE Ochoa. 2n= Tubers are very small and white. SOLANUM ACAULE Bitt. 2n=48, genome formula Peru.

165 82 SOUTH AMERICAN REGION A 2 A 2 A 3 A 3- w ild tetraploid from C. Peru, Bolivia and NW. Argentina. A parent of S. x juzepczukii*. Frost resistant and resistant to X-virus disease, nematodes and the Colorado beetle. Very susceptible to Phytophthora. SOLANUM AJANHUIRI Juz. & Buk. 2n=2x=24. C u l t i vated in N. Bolivia (dept. La Paz) and S. Peru. It derives from S. tuberosum* group stenotomum x S. megistacrolobum Bitt., 2n=24 (Huaman et al., 976). It is frost resistant, the tubers are long and irregularly shaped. Also resistant to virus diseases. SOLANUM CHACOENSE Bitt. 2n=24, (36). N. and C. Argentina, Paraguay, Uruguay and S. Brazil. A very polymorphic wild species. Only once an autotriploid was observed. This triploid has been described as 'S. calvescens Bitter'. 'S. muelleri Bitter' also belongs tos. chacoense (Brücher, 975). Rich in tomatine alkaloid which is poisonous to the Colorado beetle. Introgression between this species and S. microdontum exists in Argentina and possibly elsewhere. This resulted in an extension of this originally low altitude species of open places of the Argentinean plain to mountainous region (Hawkes, 962a). Used as a source of resistance to common scab, virus diseases and Colorado beetle. SOLANUM xchaucha Juz. & Buk. (syn. S. tuberosum group chaucha). 2n=3x=36. This species is a hybrid of S. tuberosum* group Andigena x group Stenotomum or group Phureja, but Bukasov (970) suggested that it is atriploid derivative of group Phureja. The hybridization may have occurred several times and because of the variation of the parents this species is very polymorphic. Cultivated from C. Peru to N. Bolivia. It has a rather low yield (Jackson et al., 976). This species has run wild in Simla hills, India. Initially it was established by vegetative propagation. The older the population the more plants flower and the more self-incompatibility breaks down (Nayar & Gohal, 970). SOLANUM COMMERSONII Dun. 2n=24, 36. E. Central Argentina, Paraguay, Uruguay and S. Brazil, 'S. acroleucon Bitter' belongs to this species (Brücher, 975). A source of resistance to potato cancer and Colorado beetle. It withstands frost of -6 C. SOLANUM CONTUMAZAENSE Ochoa. 2n=. N. Peru. With white-yellow tubers of 5-25 mm length. SOLANUM x CURTILOBUM Juz. & Buk. 2n=60. The high Andes of Bolivia and Peru, where it has been cultivated. Probably a hybrid of S. x juzepczukii* and x S. tuberosum group Andigena. It reproduces itself vegetatively, although it is moderately fertile. It crosses readily with S. tuberosum group Tuberosum. Less frost resistant than its parent S. x juzepczukii (Hawkes, 962b). SOLANUM x FLAVOVIRIDENS Ochoa. 2n=. Humid trop. Bolivia. Its glandular hairs provide resistance to virus-spreading aphids. Its parents arenot yet fully known. SOLANUM GONIOCALYX Juz. & Buk. 2n=24. This potato is cultivated in C. Peru (dept. Junin). It is a northern derivative of S. tuberosum* group Stenotomum. It may be included in this species as an extreme variant (Hawkes, 958). Bukasov (970) suggested that it was a derivative of S. multi-interruptum. The tubers have a pale-yellow flesh owing to their richness in Carotinoids. They have an excellent flavour. SOLANUM HUMECTOPHILUM Ochoa. 2n=. Peru. With white-hyaline tubers of 8-2 mm length. SOLANUM IMPROVIDUM Brücher. 2n=24. La Rioja and Catamarca, W. Argentina. Wild potato of deserts (Brücher, 979). SOLANUM x JUZEPCZUKII Buk. 2n=36. High Andes of Bolivia to Mendoza Prov. of W. Argentina. Wild potato of deserts. Self-incompatible. Source of resistance to nematode and tolerance to frost (Brücher, 979). SOLANUM KURTZIANUM Bitter & Wittmack. 2n= S. Bolivia to Mendoza Prov. of W. Argentina. Wild potato of deserts. Self-incompatible. Source of resistance to nematode, and tolerance to frost (Brücher, 979). SOLANUM MACULAE Bukasov. 2n=24. Mendoza Prov. of W. Argentina. Wild potato of deserts, with high content of solanin in tubers (Brücher, 979). SOLANUM MAGLIA Molina 2n=24, 36. The 2x 'andinum' occurs wild in the Cordillera Foothills of Mendoza, W. Argentina, and the 3x 'pacificum' grows wild on the Pacific coast between Valpraiso and Loquimbo. Both are partially male-sterile (Brücher, 979). SOLANUM MURICATUM Ait. Pepino morado. 2n=24. Unknown wild. Probably domesticated in the Andes. Cultivated inc. and S. America. Extremely variable and many types of fruits are recognized. There are two closely related species, either of which could be the parental species. These are S. caripense Humb. & Bonpl. (2n=24) and S. tabanoense Correll (2n=24). Both occur in Ecuador and Colombia. Pepino is cultivated for its fruits (Heiser, 964). SOLANUM NUBICOLA Ochoa. 2n=48. The Huânuco region, Peru. A tetraploid species of the Tuberosum group. SOLANUM OCEANICUM Brücher. 2n=36. A weedy potato of the Pacific coast of Chiloe Island, Chile.

166 SOLANACEAE - SOLANACEAE SOLANUM PELOQUINIANUM Ochoa. 2n=24.Wild potato species from the Peruvian Andes. SOLANUM PENNELLII Corr.2n=. Closely related to S. lycopersicoides Dunal. (2n= ). Both species are representatives of a transition between Solanum and Lycopersicon. Itcan be crossed with L. esculentum* and it is a source of resistance to Tomato Mosaic Virus. SOLANUM PHUREJA see S. tuberosum. SOLANUM QUITOENSE Lam. Naranjillo, Lulo. 2n= 24. Unknown wild. Cultivated for its fruits in Colombia and Ecuador. Var. septentrionale R.E. Schultes & Cautrecasan is spineless (Heiser, 97). Closely related tos. pectinatum Dun. (syn. S. hirsutissimum Standi.), 2n=24. SOLANUM RAPHANIFOLIUM Card. & Hawkes. 2n=24. The dept. of Cuzco, Peru. There it occurs as a weed. A stabilized hybrid of S. megistacrolobum Bitt. (2n=24), and S. canasense Hawkes (2n=24) (Ugent, 970a). Brazil) (von der Pahlen, 977). Self and cross compatible. Artificial intervarietal hybrids have been made. It has been suggested that in nature such hybrids also occur (Heiser, 97). SOLANUM TUBEROSUM L. Potato. 2n=24, 48. There are two geographical regions where the largest number of the wild and cultivated potatoes grow;.c. Mexico (p. 97, 98) and 2. Andes ofc. Peru, Bolivia and NW. Argentina. The greatest number of tuberous Solanum species is found in Peru where the potato was probably first domesticated. This species is divided into 4 groups :. group Stenotomum (syn. S. stenotomum Juz. & Buk.), 2n=24, a cultivated type of very high altitudes from Peru to N. Bolivia. It is the parent of groups Andigena and Phureja, S. x chauca* and S. x juzepczukii*. Some provenances SOLANUM RUIZ-LEALII Brücher. 2n=24. Mendoza Prov. of W. Argentina. Wild and self-incompatible. Source of resistance to nematode and tolerance to drought. Brücher (979) concluded that it isnot ahybrid of S. chacoense* and S. kurtzianum*. SOLANUM SPARSIPILUM Bitt. 2n=24. Peru and Bolivia. A weedy species. Probably a clonal mixture of diploid hybrids of S. tuberosum groups Stenotomum and Phureja* and diploid related species like S. canasense Hawkes,S. raphanifolium Card. & Hawkes, and others which Ugent (970a) grouped in one complex species S. brevicaule* Bitt. This weedy species may form a bridge for a gene flow from S. brevicaule s.. to S. tuberosum* and vice versa (Ugent, 970a) SOLANUM STENOTOMUM see S. tuberosum. SOLANUM SUCRENSE Hawkes. 2n=48. Bolivia. A weedy potato in potato crops. It is a hybrid of S. tuberosum ssp. andigena and S. oplocense, 2n=48. Source of resistance to several pathotypes of Globodera nematodes (Astley & Hawkes, 979). SOLANUM TOPIRO Humbolt & Bonpland ex Dunal. Jibara, Uvilla, Cocona. 2n=24. S. America. Var. topiro is commonly cultivated for its fruits in the Upper Amazon valley. There are two fruit forms: ovoid named jibara and globose called uvilla. The latter has been described ass. alibile R.E. Schultes. A common weed in Ecuador is var. georgicum (R.E. Schultes) Heiser (syn. S. georgicum R.E. Schultes). Var. topiro has no spines and big fruits, while var. georgicum has spines and small fruits. It is believed that these differences are a result of domestication. The greatest variation in size, shape and content of anthocyanins of the fruits is found in the W. Amazon Basin (W. Solanum tuberosum group Andigena (Ugent, 968) are frost-resistant, have a high yield of good quality. If introduced into Europe, its selfincompatibility may have hindered its smallscale propagation at that time (Zeven, 980a). 2. group Phureja (syn. S. phureja Juz. & Buk.), criollo potato, 2n=24, genome formula A^A^, cultivated in most lowlands of Venezuela, Colombia, Ecuador, Peru and N. Bolivia. The tubers are the largest of the diploid species. Dormancy is very short (-3 days) andmay have prevented its introduction into Europe in the 6th Century and later (Zeven, 980). It matures early. It is a selection of group Stenotomum with a short dormancy. 3. group Andigena (S.tuberosum ssp. andigena, S. andigena Juz. & Buk.), 2n=48, found in the Andes of Venezuela, Colombia, Ecuador, Peru, Bolivia and NW. Argentina. Itevolved from group Stenotomum inc. Andes. Parental type of group Tuberosum. 4. group Tuberosum (syn.s. tuberosum ssp. tuberosum, S. tuberosum sensu stricto), derived from group Andigena introduced into the coastal region of S. Central Chile (Island of Chiloe and adjacent coastal mainland) (Brücher, 97; Glendinning, 975)and into Europe and N. A- merica (Hawkes, 958). Naturalized and culti-

167 SOUTH AMERICAN REGION vated Tuberosum from Chile and elsewhere have identical cytoplasmic factors (Grun, 979). The evolution of Andigena into Tuberosum was simulated by Simmonds (968). The distinction between Andigena and Tuberosum is that Tuberosum has less dissected leaves with wider leaflets, generally arched and set at a wider angle to the stem. The tubers form with long days, or in the tropics with short days only at lower altitudes. In the Canary Islands, cultivar Negra has been cultivated. It is a triploid (2n=36) (Zubeldia et al., 955). Sanudo (970) suggested that it is ahybrid of group Stenotomum and group Andigena. Zubeldia L. et al. (955) believed that it was introduced from Peru in the early part of the 7th Century together with tetraploid material. Naturalized potatoes grow in the Kilimandjaro Mountains of Tanzania, in Lesotho and Botswana. They probably derive from cultivars introduced from Europe (Brücher, 966). He described 30 types. Andigena is asource of resistance to potato virus Y. Some plants have tubers with avery black flesh,which can be used as a source of pigments. SOLANUM VIARUM Dunal. (syn. S. khasianum C.B. Clarke var. chatterjeeanum Sen Gupta). 2n=24. S. America.Fruits are source of solanidin. Related species are S. myriacanthum Dunal (syn. S. khasianum C.B. Clarke), 2n=24, naturalized in the Khasia Mountains of Assam; S. retroflexion Schrank, 2n=48; and S. platanifolium Hook., 2n=24, which is rare. S. viarum is naturalized in several places in trop. Asia (Babu & Hepper, 977). A shrub. Sterculiaceae GUAZUMA GRANDIFLORA G. Don. (syn. Theobroma grandiflora Schum.). 2n=. Brazilian Amazon basin. A tree cultivated for its fruits. THEOBROMA BIC0L0R* THEOBROMA CACAO L. Cacao. 2n=6, 20, 26. Primary gene centre:the area of the 'Upper watersof the Amazon'. Spread by man. Only in Mexico was the domestication of the cacao completed by the Maya. Elsewhere cacao was wild or semidomesticated. Cuatrecasas (964) described 22 Theobroma species, which all are found in trop.s. and C. America. Relative isolation of cacao populations and their original parentage resulted in the development of two more or less uniform groups distinguished as Criollo (T. cacao ssp. cacao) and Forastero (ssp. sphaerocarpum). The Criollo is located in C. America (Central American Criollo) and in N. Colombia (South American Criollo). The Forastero can be divided into the Upper Amazonian Forastero, indigenous to the Upper Amazon basin and the lower Amazonian Forastero, alsonamed and found in the Guyanas (Cheesman, 944; Toxo- peus, 969) and in Africa where it is known as the West African amelonado. The Trinitario is a recently originated hybrid swarm of Criollo from C. America and A- melonado. THE0BR0MA MICROCARPA Mart.2n=. Brazil. Cultivated in Bahia. Thymelaeaceae FUNIFERA BRASILIENSIS (Raddi) Mansf. 2n= Brazil. Cultivated in the West Indies for its fibres. Tropaeolaceae TROPAEOLUM LEPTOPHYLLUM G. Don.2n=. Ecuador and Peru. Cultivated for its tubers. TROPAEOLUM MAJUS L. Nasturtium. 2n=28. S. A- merica. A herbaceous vine cultivated asan ornamental plant. The flower buds and young fruits are used for flavouring vinegar. They are also used as capers. TROPAEOLUM TUBEROSUM Ruiz & Pav. Tuber nasturtium. 2n=42. A very old cultivated food plant unknown wild in Peru, Chile and Bolivia. In Bolivia it is still cultivated in the mountains above Lake Titicaca. Umbelliferae ARRACACIA XANTHORRHIZA Bancr. (syn. A. esculenta DC). Arracacha, Apio arracacia.2n= Unknown wild. Cultivated mainly in Venezuela and Colombia, also in Bolivia and Peru, and introduced into Brazil, C. America, E. Africa and India. Verbenaceae LIPPIA CITRIODORA H.B.K. (syn.l. triphylla (L. 'Hér.) Kuntze). Lemon verbena. 2n=36. S. America. Formerly much cultivated for itsverbena oil, now as an ornamental.

168 Central American and Mexican Region TheCentralAmerican andmexican Regionhasbeen describedbyvavilov asthe CentralAmerican andsouthmexicancentre oforigin.darlington &Janaki Ammal (945) named Mexico as acentre of origin, while Darlington (956)addedC.America to Mexico.Inthis region agriculture developed since the 7th Millenium BC. Harlan (97)called itcentre CI Mesoamerican centre. Old farming siteshavebeen discovered at Tamaulipas and inthe Tehuacân Valleyof Mexico. To theearliestplant remainsbelongamaranthus sp., Avocado persica,capsicum annuum,cucurbita pepo,c. mixta,gossypiumhirsutum andlagenaria siceraria. Relatively afewbut important cropshave beendomesticated in this region e.g. fruit trees, Agave sp., Capsicum sp.,cucurbita sp., Gossypium sp., Ipomoea batatas, Phaseolus sp., Zeamays etc. Agavaceae AGAVE AMERICANA L. Century plant, Magvey. 2n= 60, 20, (80, 240 and aneuplolds). Mexico. Cultivated in Mexico, Europe, Africa and N. America as an ornamental, foliage plant, a hedge plant and for pulp. AGAVE ATBOVIRENS Salm-Dyck (syn. A. latlssima Jacobi). Pulque. 2n=50, 80. Mexico. Extensively cultivated in Mexico. The inflorescence iscut off and the sweet juices that collect in the cavity produced are allowed to ferment to produce the alcoholic drink pulque or mexcal de pulque. AGAVE CANTALA (Haw.). Roxb. (syn.a. candelabrum Tod.). Cantala. 2n=90. Probably Mexico. There a wild form occurs on the western coast. Smaller than the cultivated types. It was taken to the Philippines and later to Indonesia where it is cultivated for its fibre. In India cultivated as a hedge and anti-erosion plant (Purseglove, 972). AGAVE COMPULVIATA Trel. Pulque. 2n=. Mexico. Cultivated around Comitân in Mexico. Comiteca, a fine liquor, is distilled from a mixture of fermented sugar-cane juice and a mash made from the stems and leaves of this species. AGAVE CRASSISPINA Trel.Maguey manso.2n= Mexico. Cultivated there. AGAVE DEWEANA Trel. Zapupe verde, Zapupe de Tantoyuca. 2n=. Cultivated for a long time by the Tantoyuca Indians in Mexico. AGAVE FOURCROYDES Lem. Henequen agave. 2n=c. 40. Yucatan, Mexico. Primary centre also there. Secondary centre probably in E. Africa (p. 22). Cultivated in many countries for its excellent fibre. AGAVE HETERACANTHA Zucc. (syn. A. funkiana Koch & Bouche). Ixtle de Juamava. 2n= Arid NE. Mexico. Cultivated for its fine fibre, used to make cordage and brushes. AGAVE LECHEGUILLA Torr. Lechuguilla, Tula Istle. 2n=. Texas and N. Mexico. Cultivated for fibre,used to make rugs, mats and brushes. AGAVE LETONAE F.W. Taylor. Letona, Salvador henequen.el Salvador. Cultivated there for centuries. AGAVE LOPHANTA Schiede. Lechuguilla. 2n= Mexico. Cultivated by natives for its fibre. AGAVE SCHOTTII Engllm. Amole. 2n=. Sonora of Mexico and arid Arizona. Cultivated by A- merindians. Dried pulp of leaves used as soap. AGAVE SISALANA Perr. (syn. A. rigida Mill.): Sisal agave. 2n=(c. 38, 47, 49), 50. Mexico and C. America.Primary centre in Yucatan,

169 88 CENTRAL AMERICAN AND MEXICAN REGION OPUNTIA CRYSTALENIA Griff. 2n=. Mexico. Cultivated there for its edible fruits. OPUNTIA DILLENIA (Ker-Gawler) Haw. Pest pear. 2n=22, 66.S. USA, Bermuda, West Indies, N. South America.Cultivated there and on Tenerife (Canaries), Italy, W. Africa, India, E. Asia and Australia for its edible fruits. It has often run wild. OPUNTIA FICUS-INDICA (L.) Miller. Indian fig, Nopal, 2n=22, 88. Probably Mexico (Hammer, 976). Cultivated in (sub)tropics for its edible fruits,and as a forage and hedge plant. OPUNTIA FUSICAULIS Griff, 2n=. Not known wild. Thornless species cultivated in Mexico and S. USA as forage. OPUNTIA HYPTIACANTHA Weber. 2n= Cultivated for its fruits.. Mexico. OPUNTIA LANCEOLATA (Haw.) Haw. 2n=88.. Mexico. Occasionally cultivated in the (sub)trop. Americas for its fruits. OPUNTIA LEUCOTRICHA DC. 2n=44. Mexico. Cultivated there and around the Mediterranean area for its fruits, and as an ornamental and hedge plant. OPUNTIA LINDHEIMERI Eng. 2n=. SW. Louisiana, SE. Texas (USA), and NE. Mexico. Cultivated for its fruits and as aforage. OPUNTIA MAXIMA Mill. 2n=. Probably Mexico. Occasionally cultivated in the (sub)trop.americas and Asia. Not known wild. OPUNTIA MEGACANTHA Salm-Dyck (syn. 0. castillae Griffith). Tuna, Nopal.2n=. Mexico. Cultivated there and in Jamaica, S. California and Hawaii. Thornless strains cultivated as forage. Naturalized on Hawaii (Hammer, 976). OPUNTIA ROBUSTA Wendl. 2n= vated there for its fruits. Mexico. Culti- OPUNTIA STREPTACANTHA Lem. 2n=. Mexico. Cultivated there. Var. streptacantha is thorny and isused as an animal déterrant, whereas var. pachona (Griff.) Hammer (syn. 0. pachona Griff.) is thornless and is cultivated for its fruits. OPUNTIA TOMENTOSA Salm-Dyck. 2n=44. Mexico. Cultivated there.formerly used as feed for cochenilla. OPUNTIA UNDULATA Griff. 2n=. Mexico. Cultivated there for its edible fruits. PACHYCEREUS PECTEN-ABORIGINUM (Eng.) Britt. & Rose. 2n=. Mexico. Hedge plant. PACHYCEREUS PRINGLEI (S. Wats.) Britt. & Rose. 2n=. Mexico. Cultivated as a hedge plant. PERESKIOPSISCHAPISTLE (Weber) Britt. & Rose. 2n=. Mexico. Cultivated as a hedge plant. PERESKIOPSIS SPATHULATA (Otto) Britt. & Rose. 2n-.S. Mexico. Hedge plant. RITTEROCEREUS PRUINOSUS (Otto) Backeb. 2n= C. and S. Mexico. Cultivated as a hedge plant and for its edible fruits. SELENICEREUS GRANDIFLORUS (L.) Britt. & Rose (syn. Cereus grandi florus Mill.). 2n=22. Jamaica, Cuba, Mexico. Cultivated as asource of drug, and widespread as an ornamental. STENOCEREUS STELLATUS (Pfeiff.) Riccob. 2n=. Mexico. Cultivated for its edible fruits and as a hedge plant. Caricaceae CARICA PAPAYA L. Papaya, Pawpaw. 2n=8. Lowlands of C. America somewhere in the region between S. Mexico and Nicaragua. Unknown wild. The history of its domestication is not known. Papaya has now spread to all tropical countries andmay have runwild as was observed in the forest fringes in N. trop. Argentina. Closely related toc. pelta Hook. & Arn., which also occurs in this area. This species may have contributed by hybridization (Purseglove, 968). Chenopodiaceae CHENOPODIUM NUTTALLIAE Safford. 2n=allo 4x=36. This self-compatible species is grown as a vegetable (inflorescence)and a grain crop in C. Mexico. It isclosely related to C. quinoa* andmay derive from it. The weedy C. berlandieri Moq., 2n=36, forms a weed-cultigen complex with C. nuttalliae. C. berlandieri occurs from C. Mexico to W. USA where it isvery polymorphic (Wilson, 976). See for additional information Wilson & Heiser (979) and Wilson (98). Compositae DAHLIA VARIABILIS Desf. (syn.d. rosea Cav.). Dahlia. 2n=64. Mexico. A tuberous amphiploid ofd. pinnata, 2n=32and D. coccinea Cav., 2n =32, 64. Probably domesticated as a food crop but now grown as an ornamental. PARTHENIUM ARGENTATUM A. Gray. Guayule. 2n= 36, 54, 72, 08andmany aneuploids. Mexico and Texas,USA. Cultivated as a rubber producer. TAGETES ERECTA L.Big marigold. 2n=24, genome formula AeAe. Mexico. Cultivated as an ornamental and for its medicinal properties. Also used in religious rituals and celebrations (Neher, 968). Probably a parent of T. patula*. The genus Tagetes extends from SW. USA into Argentina and the area of the greatest diversity is in S. Central Mexico (Neher, 968).

170 CACTACEAE - EUPHORBIACEAE TAGETES PATULA L Marigold, Flor del muerto. 2n=48, genome fo rmula ApApBpBp. Mexico. Proby hybridization of T. erecta* bably originated and T. tenuifoli a Cav.,(2n-24, genome formula BtBt) or closely related species. Cultivated as an ornamental and for its medicinal properoughout the world. At one time ties. Spread thr itwas thought t o have an Old World origin bered role in the Hindu religion cause of the sac (Anderson, 952) However, its role might have been promoted by the sacredness of the yellow colour in India. Convoivulaceae IPOMOEA BATATAS (L.) Poir. in Lam. Sweet potato. 2n=6x=90, genome formula BBBBBB. Unknown wild.with long cultivation, mutation and hybridization, sweet potato is very variable and many types have been classed as separate species, like I. tiliacea (Willd.) Choisy and I. triloba L. So descriptions of Ipomoea species before Austin (977, 978) should be treated with care. Nishiyama's I. trifida is a feral I. batatas; Nishiyama's K233, Jones* both from Mexico and Jones' 7.3 from Colombia and Jones' 73. from Ecuador are 4x derivatives of I. batatas x I. trifida* (Austin, 977). Sweet potato was already cultivated in Polynesia in pre-columbian times. Whether it was brought there as tubers by man or reached it as capsules or on drifting material by seacurrents (Purseglove, 968) is not yet known. Sweet potato is cultivated inmany tropical countries. IPOMOEA PURGA (Wender.) Hayne (syn. Exogonium purga (Wender.) Benth.). Jalap. 2n= E. Mexico, S. Mexico and C. Panama.Cultivated in Mexico, the West Indies and later India for its medicinal tubers. IPOMOEA TRIFIDA (H.B.K.). G. Don. 2n=2x=30. Mexico and Caribbean to N. Colombia and Venezuela. Common weed. Many chromosome counts attributed to this species refer to I. batatas*. According to Austin (977), Nishiyama's I. trifida is a feral I. batatas. Cucurbitaceae CUCURBITA FICIFOLIA Bouché. Malabar gourd, Fig-leaf gourd. 2n=40, (42). Highlands of Mexico and America. This speciesmight be a derivative ofc. lundelliana* (Whitaker & Davis, 962). CUCURBITA MIXTA Pang. Pumpkin, Winter squash, Walnut squash. 2n=40. It probably derives from C. lundelliana Baily in C. America and S. Mexico. It appears that it was widely distributed in N. Mexico and SW. USA in pre-colombian times (Purseglove, 968). It is a primitive horticultural crop with little fruit flesh. It crosses with C. moschata* and has been described as belonging to this species. It developed later than C. maxima* and C. pepo*. CUCURBITA MOSCHATA Duch. Cushaw, China squash, Pumpkin, Winter squash. 2n=(24), 40. From Mexico to Peru. Domesticated in BC. (Willey, 962). Cultivated throughout the world. Whitaker & Davis (962) suggested it to be a derivative of C. lundelliana*. CUCURBITA PEPO L. Marrow, Zucchini, Pumpkin. 2n=(24, 28), 40, (40-42, 44-46). Whitaker & Davis (962) suggested that his species is a possible derivative of C. lundelliana Bailey, wild gourd, 2n=40, spreading into N. Mexico and SW. USA. In Texas the related wild C. texana Gray is found. This species is either a weedy offspring of C. pepo ormay have been involved in the latter's formation. Whitaker & Cutler (969)observed one seed in a layer dated c BC. in a cave in Mexico. Var. ovifera (L.) Alef. iscultivated for its ornamental fruits. CYCLANTHERA PEDATA Schrad. 2n=32. Cultivated in Mexico for its young fruits and shoots. POLAKOWSKIA TACACCO Pitt. Tacaco.2n= Rica. Semi-cultivated for its fruits. SECHIUM EDULE Schwartz (syn. Chayote edulis Jacq.). Chayote, Guisquil, Christophine. 2n= 24. Mountanous America and Mexico. Centre of variation is from Guatemala to Panama. It was a common crop among the Aztecs before Spanish conquest. A perennial vine now grown insubtrop. N. America, Africa and Europe. Insect pollination, resulting in selfing, crossing and vivipary. Dioscoreaceae DI0SC0REA FLORIBUNDA Mart. & Gal. 2n=36, 54. S. Mexico and adjacent areas of C. America. Cultivated in America to yield sapogenin (Coursey, 967). Ebenaceae DIOSPYROS EBENASTER Retz. Black sapote, Zapote negro. 2n=. Probably Mexico. Cultivated for its fruits. Ehretiaceae CORDIA DODECANDRA DC. Copte, Siricote. 2n= Mexico. A tall tree cultivated for its fruits. Elaeocarpaceae MUNTINGIA CALCABURA L. Panama berry, Capulin. 2n=. Widely cultivated for its sweet edible fruits. Euphorbiaceae JATR0PHA AC0NITIF0LIA Mill. (syn. Cnidoscolus chayamansa McVaugh.). 2n=. A shrub. Cultivated in the Yucatan area, Mexico. Young shoots and leaves are eaten as a pot herb. During do- Costa

171 90 CENTRAL AMERICAN AND MEXICAN REGION mestication, forms with fewer stinging hairs were selected for (Dressier, 953). MANIHOT ESCULENTA Crantz. Cassava, Manioc, Manihot, Yucca. 2n=36. Schmidt (95, cited by Nassar, 978)stated that cassava was taken by Amer-indians from the N. Amazon to Mexico some 000 years ago. But Roger (963) mentioned that cassava could have been domesticated in Mexico too, because there is no reason to exclude domestication of roots in the Mexican seed-type agriculture. If so,one would expect wild ancestral species there. Cassava iscultivated in W. and S. Mexico, C. America including parts of Guatemala. There are only sweet cultivars in Mexico, where they have been developed and whence they have spread. However sweet types in Brazil could have developed locally. More research is needed. 970; dewet & Harlan, 974). TRIPSACUM LANCEOLATUM Rupr. ex Fourn. (syn. T. lemmoni Vasey). 2n~72. Mountains of arid S. Arizona and adjacent Sonora. Excellent fodder. TRIPSACUM LATIFOLIUM Hitchc. 2n=36, 72. C. America. Excellent trop, fodder. Gramineae AXONOPUS COMPRESSUS (Swartz) Beauv. Carpet grass. 2n=40, 50, 60. C. America and the West Indies. A perennial tropical grass suitable for lawns and permanent pasture. Tripsacum latifolium ( ), T. lanceolatum ( ) andt. laxum ( )(Randolph, 970). ORYZA ALTA* ORYZA LATIFOLIA* ORYZA PERENNIS Moénch. 2n=24, genome formula AA. For distribution see p. 74. In America var. cubensis (0. cubensis Ekman), the American race (2n=24, genome formula AA) of this species developed. Gopalakrishnan & Sampat (966) suggested that 0. perennis entered America as a weed of 0. sativa in post-columbian times. PANICUM SONORUM Beal. Sauwi. 2n=. Sonora of of Mexico and adjacent Arizona. Cultivated by Guarijio Indians as a cereal (Dressier, 953; Gentry, 942). PANICUM VIRGATUM L. Switch grass. 2n=36, 72 and aneuploids. N. and C. America. Cultivated ascattle food. SETARIA GENICULATA (Lam.) Beauv. 2n=36, 72. An early crop in Tehuacan Valley, Mexico; now replaced by maize and European food crops. SETARIA MACROSTACHYA HBK. 2n=54, 72. An early crop in Tehuacan Valley, Mexico; now replaced by maize and European food crops. TRIPSACUM ANDERSONII Gray. Guatemala grass. 2n=64. C. and S. America. This species combines 54 Tripsacum and 0 Zea chromosomes in its genome. It is widely cultivated in trop. America as a fodder, and to indicate property boundaries (dewet et al., 976). TRIPSACUM DACTYLOIDES (L.) L. Gamma grass.2n= 36, 72. C. USA to Paraguay. Excellent cultivated fodder grass,often irrigated. Gene exchange with Zea mays is possible (Randolph, TRIPSACUM LAXUM Nash. 2n=36, 72. Veracruz Oaxaca Mexico. Excellent fodder, for the wet tropics. TRIPSACUM MAIZAR Hern. & Randolph. 2n=36, 72. C. Mexico. Robust species, widely grazed when young. TRIPSACUM PILOSUM Scribn. & Merr. 2n=36, 72. C. America. Excellent fodder. TRIPSACUM Z0PIL0TENSE Hern, fe Randolph. 2n=72, 76. Arid W. Central Mexico. The only Tripsacum species without rhizomes. Extensively grazed. ZEA DIPLOPERENNIS litis, Doebley & Guzman. Diploid perennial teosinte. 2n=20. Jalisco, Mexico. Crosses with maize toproduce fertile hybrids (litis et al., 979). ZEA LUSURIANS (Durieu & Ascherson) Bird. Guatemala teosinte. 2n=20. Guatemala and Honduras. Crosses readily with maize to produce fertile hybrids. ZEA MAYS L. ssp. mays. Maize, Corn. 2n=20. The 'origin* of maize has been subject to much discussion and research. The present conclusion is that teosinte (Z. mexicana )is the wild parent of maize (de Wet et al., 97;de Wet & Harlan, 972; Galinat, 97;J.G. Waines, see Galinat, 97). Other theories were that maize derived from hybridization of primitive maize, teosinte and Tripsacum ssp. when the first reached C. America from S. America. If so wild maize - a podcorn - tunicata Sturt. - should have been extinct by now.this complete disappearance of wild maize would have been causedby a number of factors such as intro-

172 EUPHORBIACEAE - GRAMINEAE 9 gression of genes of domesticated maize, the use of the habitats of wild maize for maize cultivation and the grazing of Old World animals. The earliest finds of maize were tiny cobs (2-3 cm) in the Bat Cave of Tehuacan, Mexico. They have been dated about 3600 BC. Two types have been observed:. a podcorn and 2. a popcorn - everata Sturt. (syn. praecox) (Mac- Neish, 964; Mangelsdorf et al., 964). In another cave,early Bat Cave-like maize and teosints have been identified. The material dates from about 2200 BC.Early tripsacoid maize dates from BC.MacNeish (964a, b) supposed that about 5000 BC. wild maize i.e. teosinte was cultivated. From C. America maize reached S. America where its development became more advanced (p. 7). Advanced varieties were returned to C. America. For Mexico they have been described as Pre-Columbian Exotic Races comprising of 4 varieties: Cacahuacinthe, Harinoso de Ocho, Olotón and Maize Dulce (Wellhausen et al., 952). Where they hybridized with primitive Mexican Ancient Indigenous Races: Palomero Toluqueno, Arrocillo Amarillo, Chapalote and Nal- Tel. This resulted in a new group of varieties called: Prehistoric Mestizos. Introgression with teosinte most likely has taken place too. Wellhausen et al. (952) described 3 races. Modern incipient Races have developed since the Conquest. Wellhausen et al. (952) described four of them. Some being very recently developed. The same development took place in Guatemala (Wellhausen et al., 957). From C. America maize also spread to N. A- merica (p. 203). De Wet et al. (972)suggested that the 'tripsacoid' races of maize ins. America were originally introduced from Mexico orc. America where they inherited their 'tripsacoid' characteristics through introgression with teosinte. It is also possible that these races represent relics which retain some original teosinte-like characteristics inherited from the maize progenitor. The oldest domesticated maize varieties had a string (slender) cob. This primitive characteristic is still found insome relic cultivars like Confite Morocho, which is the most primitive living cultivar (Galinat, 969).It comes from Peru. The domesticated recessive character thick cob is conditioned inthe Corn Belt dentby 3 major loci. One allele derives from northern flintwhich obtained it from Maize de Ocho. It is possible that this allele introgressed from Tripsacum sp. probably T. dactyloides*. Perhaps this introgression could be traced tos. America by way of the cultivars Cabuya and Sabanero (Galinat, 969). However, dewet et al. (972) believed that no introgression exists between maize and Tripsacum species. So cv. Chococena isnot ahybrid of cv. Confite (ex Peru) and a Columbian Tripsacum as has been suggested. The source(s) of other two recessive alleles is not fully understood. One of these alleles produces high condensation of staminate spikelets in the tassel branches and the other increases tassel branching. If the first allele is absent the expression of the second is complete resulting inprofuse branching like the mutant ramosa. Some cultivars of the southern dent and 'bear paw' popcorn appear to have this high condensation-ramosatype of thick cob. The degree of fasciation with which this type of thick cob may be asociated seems to have been modified by teosinte introgression, teosinte gene(s) suppressing fasciation. It is suspected that the Corn Belt dents obtained these two pairs of recessive alleles from the southern dents (Galinat, 969). Other morphological changes due to domestication are a development of a complete husk coverage of themature ear, the development of female inflorescense, a reduction of the glumes of the female inflorescense, an arrangement of spikelets in a higher row number, a development of the cupules and an increase of the length of the styles (silk). Some cultivars have an ear length up to 45 cm.hybrid maize varieties may produce more than 000 kernels per cob while the Tehuacan maize has about 40 kernels per cob. The terminal inflorescense becomes entirely staminate being a lax plume with waving branches (Galinat, 969). A flow of teosinte genes to maize still exists where maize cultivation is primitive and teosinte is present.maize x teosinte hybrids are actually cultivated. Maize may show pronounced signs of 'tripsacoid' i.e. teosinte germ plasm such as induration of the lower glume and a straight rigid ear. Less genes flow from maize to teosinte since the genetic incorporation of a maize-like rachis results in the inability to disperse seed and so to the extinction of teosinte introgressed with maize (Wilkes, 970). Extensive gene exchange in both directions is evident around Chalco, S. of Mexico City, where the weedy teosinte race mimics the local race of maize in size, colour and pubescence. These weeds remain teosintoid with respect to female inflorescense structure. In many other areas of Mexico, particular the Rio Balsas Valley on the W. escarpment, W. of Mexico City, teosinte behaves essentially as a wild grass, but modern development leads to an increased infiltration of maize genes into teosinte. Several types of maize are cultivated. An improved popcorn is being cultivated in USA, Mexico and elsewhere. Softcorn - amylacea Sturt. - predominates in the Andean region. Flint maize, flint corn - indurata Sturt. - predominates in N. Colombia and E. S. America. Sweet corn - saccharata Sturt. (syn. rugosa Bonof.) - was cultivated for the preparation of South American and Mexicanbeer'. At present it is mainly cultivated in USA. Waxy maize - ceritina Kulesh. - cultivated in the Americas and in E. Asia. Dent maize - indenta Sturt. is the main type of the Corn Belt of USA and N. Mexico. A hybrid of a late-maturing dent Gourd Slide cultivated in the south, and an early ma-

173 CENTRAL AMERICAN AND MEXICAN REGION turing flint maize, mainly cultivated in the north. The latter derives from Maiz Ocho. From C. and S. America maize was taken to Europe (p. 35), Asia (p. 33)and Africa (p. 7), where secondary centres of diversity developed. At present flint maize, flint corn - indurata Sturt. - is quite common in C. America. Derivatives of Z. mays /2* Z. mexicana are used for fodder. These are called maisinte (Prasad & Chaudhuri, 968). ZEA MAYS ssp. mexicana (Schrader) litis (syn. Z. mexicana (Schrader) Kunze, Euchlaena mexicana Schrader). Mexican teosinte. 2n=20. C. Plateau and Valley of Mexico. MacNeish (964a, b) suggested that about 5000 BC. wild maize i. teosinte was cultivated. It is the ancestor of maize, Zea mays*. Owing to natural hybridization between maize and teosinte there is a gene flow from teosinte to maize, while that TRIGIDIA PAVONIA (L.f.) DC. Cacomite, Tiger flower. 2n=26, 28. Mexico. Naturalized in most ofc. America, Colombia, Bolivia, Peru and Brazil.Easily cultivated. Soon escapes into maize fields etc. as a weed. The Aztecs cultivated this species for almost a 000 years. Cultivated now as an ornamental which resulted in spontaneous variations in colour and size (Molseed, 970). Juglandaceae CARYA PECAN (Marsch.) Engl. & Graebn. Pecan. 2n=32. N. Mexico. Cultivated there. It resembles the walnut, Juglans regia* but the seed has a better taste. JUGLANS MOLLIS Engelm. Guatemala walnut. 2n=. Mexico and Guatemala. Similar to the walnut (J. regia*). Labiatae HYPTIS SUAVEOLENS Poit. 2n=28, 32. Cultivated mainly in Mexico. A variable and weedy plant now occurring in many parts of the tropics. SALVIA CHIA Fern. Chia. 2n=. Mexico. Seeds are used to prepare a beverage and for painting or medicine. SALVIA DIVINORUM Epling & Jativa-M. 2n= Wild unknown.cultivated in NE. Oaxaca, Mexico. Vegetatively propagated (Schultes & Hofmann, 973). Perhaps one clone. Closely related to S. cyanca Lindl. (2n= )of C. Mexico. Lauraceae Zea mexicana (Wilkes, 967). gene flow from teosinte to maize, while that of maize to teosinte is very small (see p. 90). This may happen between the earliest flowering teosinte plants and the latest of maize. Teosinte plants may grow unnoticed in a maize field and may be harvested together with its leader crop. Seeds may be spread either during the transport or storage of the crop or in manure (Wilkes, 967). Attempts have been made tocultivate teosinte as afodder,but it yields less than sorghum. This subspecies is often found as a weed in fields of maize, and introgresses naturally with domesticated maize. See also Z. mays ssp. parviglumis*. Iridaceae PERSEA AMERICANA Miller. Avocado. 2n=24, (36, 48). Mexico and C. America. There are three geographical races: () Mexican (also named P. americana var. drymifolia Mez. syn.p. drymifolia Cham. & Schlecht.) from the Mexican highlands where wild progenitors have been found. Its anisescented leaves, hardiness and small fruits are characteristics. (2) Guatemalan, from the Guatemalan highlands. Wild progenitors have been found in this area. (3) West Indian, from the Guatemalan lowlands which has only spread to the West Indies in post- Columbian times (Purseglove, 968). Where these races grow together - either in their native region or elsewhere - hybrids originate (Bergh, 969). Electrophoretic studies showed that the Guatemalan race is the most ancient form (Garcia & Tsunewaki, 977). The Mexican and West Indian races have been described as var. americanum. PERSEA FLOCCOSA. 2n=24. Semi-cultivated in Puebla-Veracruz area of Mexico. PERSEA SCHIEDEANA Ness. Coyo avocado. 2n=24. From Guatemala to Mexico. Semi-cultivated chiefly in Orizaba, Mexico for its edible fruits (Bergh, 969). Leguminosae CAL0P0G0NIUM MUCUNOIDES Desv. 2n=. Trop. A- merica. A cover crop and green manure. Cultivated inthe tropics where it has naturalized. CANAVALIA CAMYL0CARPA Piper. Babricon bean.

174 GRAMINEAE - LEGUMINOSAE 93 2n=. The West Indies. Cultivated as agreen manure. CROTALARIA LONGIROSTRATA Hook. & Arn. Much of Mexico and C. America (Dressier, 953). A large herb cultivated in Guatemala as a potherb. DESMODIUM TORTUOSUM (Sw.) DC. (syn. Meibomia purpurea (Mill.) Vail.). Florida clover, Giant beggarsweed. 2n-22. C. America, Florida,West Indies and N. South America. A perennial herb used as a green manure and forage crop. DESMODIUM UNCINATUM* ENTEROLOBIUM CYCLOCARPUM (Jacq.) Griseb. 2n= 26. Mexico,C. America, N. South America and West Indies. Cultivated as a shade tree. INGA DULCIS. 2n=. Mexico. Used as a shade tree and hedge plant. INGA EDULIS Mart. Food inga. 2n=26. Mexico and C America. Used as a shade tree. INGA GOLDMANII Pittier.2n= Used as a shade tree. C. America. INGA LAURINA (Sw.) Willd. Sackysac. 2n= C. America and the West Indies. It has edible fruits. Used as a shade tree in coffee plantations in the New World (Purseglove, 968). INGA LEPT0L0BA Schlecht, and S. America. Used as INGA PITTIERI Micheli. 2n= as a shade tree. 2n=. Mexico, C. i shade tree.. C. America. Used LEUCAENA LEUCOCEPHALA (Lam.)de Wit (syn. L. glauca (Willd.) Benth.,L. latisiliqua (L.) Gillis. Leucaena. 2n=04. C. America, spread to the Caribbean islands,the Philippines, SE. Asia and elsewhere. Var. glabrata is L. glabrata Rose. Three types are recognized: () Hawaiian type, a short bush with year round flowering causing a high seed production and easily becoming a pest weed. (2) Salvador type, a tall tree of the inland forests of C. America. Mexican material coming through the Philippines to Hawaii was the source of the Hawaiian Giants cultivar group bred for timber. (3) Peru type,an extensively branching tree extremely rich in foliage. Forage cultivars derive from this type; forage cultivar Cunningham is a hybrid of Salvador type and Peru type. Cultivars poor in mimosin are bred for. Leucaena is autogamous. Since 900 hybrids (2n=80)ofL. leucocephala and L. pulverulenta (Schlecht.) Benth., 2n=56, have developed in Indonesia and shade trees and forage cultivars low in mimosin. They are partially sterile and cannot become a pest weed (Vietmeyer & Cottons. (977). MACROPTILUM ATROPURPUREUM (DC) Urb. (syn. Pha- seolus atropurpureus DC). Siratro. 2n=22. S. Texas, New Mexico (USA), Mexico,C. America, Colombia, Ecuador, Peru and Argentina. Material from Mexicohas recently been domesticated in Australia for tropical pasture. MIMOSA INVISA Mart. 2n=24, 26. C. and S. America. In Java and elsewhere it isused as a cover and green manure. The spiny stem is a disadvantage. Var. inermis Adelb. is a useful selection. PACHYRHIZUS EROSUS (L.) Urban, (syn.p. angulatus Rich, ex DC, P. bulbosus (L.) Kurz.). Yam bean, Jicama. 2n=22. Mexico and N.Central A- merica. Cultivated there in pre-columbian times. Taken to Asia and cultivated there. In S. China and Thailand ithas run wild (Clausen, 944). Var. palmatilobus (DC.)Clausen is probably P. palmatilobus Benth. & Hook. PHASEOLUS ACUTIFOLIUS Gray. Tepary bean. 2n= 2x=22. From N. Central Arizona (USA) to Guatemala. Found in Mexico in layers dating from 3000 BC. A very polymorphic species with drought resistance,high protein content and high productivity. Wild specimens are annuals with indeterminate climbing habit and cleistogamous flowers. They belong to var. tenuifolius Gray; cultivated types belong to var. latifolius Freeman (Baudet, 977b). However Nabham & Feiger (978) stated that some wild types belong to the latter variety. PHASE0LUS COCCINEUS L. Scarlet Runner, Runner bean. 2n=22. Mexico and Guatemala. Domesticated there before 300 BC, perhaps for its thick roots. The cultivated form is var. coccineus, thewild form has not yet been described. Cultivated in the Americas and Eurasia for food, for fodder and as an ornamental. For Ph. vulgaris* it is a source of halo blight resistance, absence of parchment and string. Smartt (973) suggested that the cultigen ssp. darwinianus Hdz. X. & Miranda C may have an independent domestication of a yet unknown ancestral form. He stated that a special taxonomie treatment may be necessary. Baudet (977b)suggested that this subspecies is also described as Ph. polyanthus Greenm. but Smartt (973) stated that this latter species is related to Ph. vulgaris*. It sometimes crosses with Ph. vulgaris*. PHASEOLUS LUNATUS L. Sieva bean, Lima bean. 2n=22. C America and in the Andes from Peru to Argentine. Kaplan (965)showed that the small lima bean of Mexicomay have arisen in the Pacific coastal foothills of Mexico and that the big lima bean of Peru was first domesticated in the Andean highlands (p. 74). Baudet (977a)divided the species into a wild form var. Silvester Baudet and the cultivated form var. lunatus, which is subdivided into three cultivar groups: () Lima (Big Lima/ True Lima) with large flat seeds corresponding to Ph. inamoenus L.; (2) Sieva (Small Lima) with medium-sized seeds corresponding to Ph. lunatus

175 CENTRAL AMERICAN ANDMEXICAN REGION L. sensu stricto;and (3) Potato with small glo- PITHECELLOBIUM DULCE Benth. Manila tamarind, bular seeds corresponding to Ph. bipunctatus 2n=26. Mexico ton. South America. The arillus Jacq. is edible. A tree planted in hedges. PHASEOLUS RETUSUS Benth. Metcalf bean. 2n= Texas, Arizona and New Mexico, USA. Occasionally cultivated. PHASEOLUS VULGARIS L. Common bean. 2n=22. Kaplan (98)suggested a multiple domestication within C. America from a widespread and polymorphus species. Ithas been suggested that this domestication may have taken place between 5000 and 3000 BC. From C. America it would have spread to other parts of America (Kaplan, 965). However, Heiser (965) believed in an independent domestication from the closely related Ph. aborigineus Burk. (syn. Ph. vulgaris L. var. aborigineus (Burk.) Baudet), 2n=22, which occurs wild in N. Argentina, Bolivia, Peru, Ecuador, Colombia and Venezuela. In Argentina and Venezuela, the wild plant is harvested (Berglund- Brücher & Brücher, 976). When Ph. vulgaris reached the area of Ph. aborigineus, aborigineus genes may have introgressed into vulgaris. However, Gentry (969)and Baudet (977b) suggested that Ph. aborigineus could be a naturalized type. Gentry (969)pointed out that a wild type of Ph. vulgaris grows inc. America and is the parent of the cultivated forms. From this wild type the cultivated forms derive. Ph. arborigineus was reduced by Burkart & Bücher (953) to a subspecies level of Ph. vulgaris. They suggested that this subspecies was not taken to S. America. A study of the wild and cultivated beans of this area should clarify whether this is so or whether ssp. arborigineus is an escape of early cultivated forms (Gentry, 969). Itsprimary centre of diversity is in Mexico. Here introgression between cultivated andwild Ph. vulgaris and P. coccineus* occurs (Wall, 970). The purple-marbled culti- Phaseolus vulgaris (Gentry, 969). vars like Kievitsboon in the Netherlands may derive from such introgression. Secondary centres in Eurasia (p. 40). SOPHORA SECUNDIFLORA (Ort.) Lagasca ex De Candolle. Mescal bean, Red bean, Coral bean. 2n= 8. N. Mexico to Texas and New Mexico.Used as hallucinogen. Often planted as an ornamental. TEPHROSIA SINGAP0U (Buc'hoz) A. Cheval, (syn. T. toxicaria (Sw.) Pers.) 2n=22. Mexico and C. America to Peru and N. Brazil.Cultivated as a fish poison. Malpighiaceae BUNCH0SIA COSTARICENSIS Rose.2n= Rica. Cultivated for its fruits. MALPIGHIA URENS L. Barbados cherry. 2n= The West Indies. Cultivated for its fruits. Malvaceae GOSSYPIUM ARIDUM (Rose& Standley) Skovsted. 2n=26, genome formula DD Mexico. GOSSYPIUM ARMOURIANUM Kearney. 2n=26, genome formula D2_iD2-l. San Marcos Islands, Gulf of California. Costa GOSSYPIUM G0SSYPI0IDES (Ulb.) Standley. 2n=26, genome formula DgDg. Mexico. It crosses poorly with most of the species of the D genome. Its seed-protein pattern is different from the D genome species. However, it is similar to the pattern of G. klotzschianum* (Cherry et al., 970). GOSSYPIUM HARKNESSII Brandg. 2n=26, genome formula D2_2^2_2 The i- s l anas ana coasts of the Gulf of California. Var. davidsonii is, according to Bahavandoss & Jayaraman (973), the progenitor of the D genome found in 4x species. GOSSYPIUM HIRSUTUM L. Upland cotton. 2n=52, genome formula (AADD). The current theory is that this species originated in C. America. Harland (970) suggested that its centre of origin is in S. America (p. 76), while C. America is an important secondary centre of diversity. Upland cotton is cultivated in USA, in Africa except for the Nile Delta (p. 76), in C. Asia, India (Cambodia type, p. 59), S. America, SE. China, Indochina and elsewhere. In C. America and the West Indian islands race marie galante is found while race punctatum is found around the coasts of the Gulf of Mexico from Florida to Yucatan in the Bahamas and on some West Indian islands. It was taken to W. Africa where it spread in the zone south of the Sahara, to Réunion, the Malabar Coast of India, Polynesia, the Marquesas, Fiji and N. Australia. A great diversity was found in N. Australia. Race yucatense is probably a cotton that has run wild and is now naturalized into natural vegetation of the coastal sand dunes of the

176 LEGUMINOSAE - PORTULACACEAE 95 Progeso area, Mexico. Race morilli is found in Oaxaca,Peubla and Morelos, C. Mexico. A perennial cotton with a bushy form and broad, intensely hairy leaves. Race palmeri in the state Guerrero, Mexico. It has deeply dissected leaves and strong anthocyanin pigmentation. Race latifolium is found in Chiapas, Mexico and neighbouring regions of Guatemala. An annual cotton. Throughout its territory a smallfruited form is found. A large-fruited form grows in the vicinity of Acala, Chiapas. This race appears to be the foundation stock of all the annual G. hirsutum cottons (Hutchinson, 962). GOSSYPIUM KLOTZSCHIANUM Andersson var. davidsonii (Kellogs) Saunders. 2n=26,genome formula D3_DD3_D. Shores of the Gulf of California and the Revilla Gigedo islands. Related to the plants of this species found on the Galapagos Islands (p. 76). GOSSYPIUM LOBATUM Gentry. 2n=26, genome formula DyDy. Mexico. GOSSYPIUM THURBERI Todaro. 2n=26, genome formula DjD^. Arizona, USA and Sonora and SW. Chihuahua, Mexico. At one time it was thought to be the American parent ofg. herbaceum* and G. barbadense*. GOSSYPIUM TRILOBUM (Sess. & Moc. ex DC.) Skovsted. (syn. Ingenhouzia triloba Moc. & Sess. ex DC). 2n=26. C. Mexico (Fryxell & Parks, 967). Moraceae BROSIMUM ALICASTRUM Swartz. Ramon, Ramon breadnut tree. 2n=26. Trop.Araerica. Seeds are edible when roasted. Probably planted by the Maya (Lundell, 937). CASTILLA ELASTICA Cerv. Arbol del Hule. 2n=28. S. Mexico and C. America. Cultivated in C. A- merica, Trinidad and Tobago, and Java at the end of the 9th Century. Now replaced by hevea rubber. Myrtaceae PIMENTA DIOICA (L.) Merr. (syn.p. officinalis Lindl.). Allspice, Pimento. 2n=22. The West Indies and C. America. Spread toother countries. PSIDIUM FRIEDRICHSTHALIANUM (Berg.) Nied. Costa Rican guava. 2n=. C. America. A small tree cultivated for itsacid fruits. PSIDIUM GUAJAVA L. Guava. 2n=22, 4x=44. Trop. America. Cultivated there. In 526 Oviedo reported that improved forms were cultivated in the West Indies. Spread through the tropics, where guava may be found naturalized. PSIDIUM MOLLE Bertol. Guisare. 2n=5x=55. S. Mexico and C. America. Cultivated for its fruits. PSIDIUM SARTORIANUM (Berg.) Nied. Pichlché, Arrayan, Guayabillo. 2n=. Mexico. Cultivated there. Onagraceae OENOTHERA BIENNIS L. (syn. Onagra biennis Scop.). Evening primrose. 2n=4. N. America to Mexico. Run wild over a large part of the world. Cultivated as a fodder crop. Orchidaceae VANILLA FRAGRANS (Salisb.) Ames. (syn. V. planifolia Andrews). Vanilla. 2n=(28-32), 32. C. A- merica, SE. Mexico and the Antilles. A perennial vine cultivated in the tropics and S. Mexico for its aromatic fruits. VANILLA POMPONA Schiede (syn.v. grandiflora Lindl.). West Indian vanilla. 2n=32. SE. Mexico, C. America and N. South America. A perennial vine on Tahiti, Martinique and Guadeloupe for its aromatic fruits. Palmae CHAMAEDOREA TEPEJILOTE Liebm. 2n=32. Ch. wendlandiana (Oerst.) Hemsl. 2n=. At least one, and probably several, species of this genus are cultivated in S. Mexico and C. America. The young staminate flower clusters are used as a vegetable (Dressier, 953). GUILIELMA GASIPAES* Papaveraceae ARGEMONE MEXICANA L. Mexican prickly poppy. 2n=28. SW. USA and Mexico. Cultivated in Mali. Seeds are used toprepare oil. Also an ornamental. Passifloraceae PASSIFLORA SEEMANNII Griseb. 2n=8. Panama and Andes of Colombia. Polygonaceae COCCOLOBA UNIFERA L. Seaside grape.2n= Trop. America. A shrub or tree cultivated for itsedible fruits. RUMES HYMENOSEPALUS Torr. Canaigre, Wild rhubarb, Pie dock, Sour dock, Tanner's dock. 2n= 00. SW. USA and adjacent Mexico. A perennial herb occasionally cultivated. Portulacaceae CLAYTONIA PERFOLIATA Donn. ex Willd. (syn. Montia perfoliata How.). Winter purslane, Miner's lettuce. 2n=36. N. America and Mexico. Cultivated as a vegetable.

177 96 CENTRAL AMERICAN AND MEXICAN REGION PORTULACA OLERACEA L. Purslane. 2n=2x=8, 4x= 36, 6x=54. Widespread. The distribution of the 2x, 4x and 6xwild and weedy subspecies shows that Mexico is one of the centres of diversity (Danin et al., 978). The cultivated type probably developed in Europe (p. 58). Rosaceae CRATAEGUS PUBESCENS (H.B.K.) Steud. (syn. C. stipulosa (H.B.K.) Steud.). 2n=34. A tree widely cultivated in Mexico and Guatemala for its fruits. Rutaceae Simaroubaceae SIMAROUBA GLAUCA DC. Aceituno. 2n=. S. Florida to Costa Rica. In El Salvador and elsewhere attempts are made to cultivate it asan oil crop. Simmondsiaceae SIMMONDSIA CHINENSIS (Link) Nutt. (syn.s. californica Nutt). Jojoba, Pignut, Goatnut. 2n=56, c. 00. California and adjacent Mexico. Dioecious. It is now being domesticated in California. It yields a stable liquid wax, jojoba oil. CASIMIROA EDULIS LaLlave & Lex. White sapote, Zapote Blanco. 2n=36. Highlands of Mexico and C. America. A fruit tree introduced to other subtropical countries. CITRUS PARADISI Macf. Grapefruit. 2n=8. Unknown wild. Closely related toc. grandis and it probably is a bud mutation or a hybrid product of C. grandis and sweet orange (C. sinensis*). This must have occurred in the West Indies some time before 750 (Purseglove, 968). It is widely cultivated in the (sub)tropics. Hybrids with other Citrus-species have been obtained, Sopomaldin is a hybrid with C. mitis (Calamondin), Siamelo with C. reticulata* (King Orange), Tangelo with the same species var. deliciosa (Tangerine), Satsumelo with the same species (Satsuma) and Chironja with C. sinensis* (Sweet Orange). Tangelolo is a hybrid of grapefruit with Tangelo. Sapotaceae CALOCARPUM SAPOTA (Jacq.) Merr. (syn. C. mammosum Pierre, Achras mammosa L.). Mamey sapote, Sapote, Marmelade plum, Mamey Colorado. 2n= C. America. A tree cultivated in the tropics for its fruits. CALOCARPUM VIRIDE Pitt. Green sapote.2n= Guatemala to Costa Rica. A tree cultivated for its fruits. CHRYSOPHYLLIUM CAINITO L. Star apple. 2n=52. West Indies andc. America.The pulp is edible. Also an ornamental. LUCUMA BIFERA Mol. Egg fruit. 2n=. Chile and Peru. Cultivated there for its fruits. LUCUMA SALICIFOLIA H.B.K. (syn. Pouteria campechiana (H.B.K.) Baenhi). Yellow sapote, Zapote amarillo. 2n=. Mexico and C. America. Cultivated for its fruits. MANILKARA ACHRAS (Mill.) Fosberg (syn. Achras zapotal., M. zapotilla (Jacq.) Gilly). Sapodilla, Chiku. 2n=26. Mexico and C. America. Cultivated now in the tropics for its fruits and gum (chickle) for chewing-gum production. Simmondsia chinensis (Gentry, 958). Solanaceae CAPSICUM ANNUUM L. Bell pepper, Cayenne pepper, Mexican chili. 2n=24. Wild variety in S. USA, West Indies, Mexico, C. America and Colombia. Primary centre in Mexico. Secondary centres in Europe (p. 62) and Asia (p. 79). Originally the cultivars were limited to C. America. The wild type is the bird pepper, var, aviculare (Dierbach)D'Arcy & Eshb. (syn. glabriusculum (Dunal) Heiser & Pickersgill). The domesticated type is var. annuum. CAPSICUM FRUTESCENS L. Tobasco pepper. 2n=24. This species consists of wild types and derived cultivars, like the Tobasco peppers. Widespread as a weed and as cultivars in Mexico, C. America and lowland S. America. It might be the parental species of C. chinense*. PHYSALIS IXOCARPA Brot. Tomatl, Miltomate, Husk tomato, Tomatillo. 2n=24. Mexico. There and in Guatemala this vegetable is cultivated. SOLANUM AGRIMONIFOLIUM Rydb. 2n=48. From C. Mexico to Bolivia. SOLANUM BRACHISTOTRICHUM (Bitt.) Rydb. 2n=24. NW. Mexico, in open pine and juniper forests and amongst bushes and rocks. Resistant to the green peach aphid, Myrus persicae Sulzer.

178 PORTULACACEAE - SOLANACEAE 97 SOLANUM BREVICAULE Bitt. 2n=24. Ugent (970a) grouped in this species wild diploid species as S. canasense Hawkes, S. brevicaule Bitt. s.s., S. raphanifolium Card. & Hawkes,S. leptophyes Bitt., S. soukupii Hawkes,S. multiinterruptum Bitt.,S. abbottianum Juz., S. liriunianum Card. & Hawkes,S. spegazzinii Bitt. and S. vidaurrei Cardenas. It is likely that from this complex species the diploid S. stenotomum* arose. SOLANUM BULBOCASTANUM Dun. 2n=24, genome formula BB, (36, BBB). At medium altitudes from C. Mexico toguatemala in grassland, waste places and forest glades and clearings. There are three subspecies (Hawkes, 966): Ssp. Bulbocastanum (2n-24, 36) is found in S, Mexico, ssp. dolichophyllum (2n=24) in the Mexican states Morelos and Guerrero and ssp. partitum (2n=24) in S. Mexico and Guatemala. This B genome also constitutes two of the genomes of S. polytrichon* A4A4BB. The B genome is related to the A4 genome and to the Ai genome of S. phureja*. S. bulbocastanum crosses with S. tuberosum* and is used as a source of resistance to Phytophthora, X-virus, Y-virus, Colorado beetle and several aphids (vectors of virus diseases). SOLANUM CARDIOPHYLLUM Lindl. 2n=24, 36.S. Mexico. Indry stony grassy and waste places, often as a weed in maize and bean fields. Ssp. ehrenbergii is found in N. Central to W. Mexico, ssp. cardiophyllum occurs in C. Mexico and ssp. lanceolatum in SE. tos. Mexico. Triploid forms are frequent in ssp. cardiophyllum. Ssp. ehrenbergii is one parent of S. sambucinum*. SOLANUM CLARUM Corr. 2n=24. Guatemala, in the high mountains.probably a link between Bulbocastana and Morelliformia series. SOLANUM DEMISSUM Lindl. 2n=(36, 48), 72, genome formula AAA4A4BB. Nw. Mexico to Guatemala. An important source of disease resistance (Phytophthora, X-virus, Y-virus). The A^-genome may have come from S. verrucosum*. One of the parental species of S. x edinense* and S. X semidemissum. SOLANUM X EDINENSE Berthault. Mexican weed potato, Papa raorado. 2n=60. Clones occur in and along the edges of cultivated field, irrigation ditches, roadsides thickets and forest fringes in the Central Volcani Cordillera of Mexico between 2000 and 3500 m. A source of Phytophthora and frost resistance. This 5x hybrid is a cross between S. tuberosum (group Andigena, 2n=4x=48)and S. demissum Lindl. (2n=6x=72) (Hawkes, 966). S. demissum genesmay introgress in the cultivated potato by backcrossing. Harvest of potato may contain tubers of S. X edinense andby trading the potato this weedy potato isalso spread (Ugent, 967). SOLANUM GUERREROENSE Correll. 2n=72, genome formula AAA4A4BB. SW. Mexico. The Al genome may have come from S. verrucosum*. SOLANUM HJERTINGII Hawk. 2n=. NE. Mexico, in pinon scrub and cultivated fields.very similar to S. fendleri*. A source of resistance to the potato aphid, Macrosiphum euphorbiae Thomas. SOLANUM HOUGASII Corr. 2n=72, genome formula A.A.A A BB.W. Central Mexico. The Ai genome may have come from S. verrucosum*. Source of resistance to potato Y-virus. SOLANUM IOPETALUM (Bitt.) Hawk. 2n=72, genome formula AAA4A4BB. W. and S. Mexico. It includes S. brachycarpum Corr. The Ai genome may have come from S. verrucosum*. SOLANUM JUGLANDIFOLIUM Dunn. 2n=24. Costa Rica. Little value to potato breeders because it does not bear stolons or tubers. SOLANUM LEPTOSEPALUM Correll.2n=. NE. Mexico and possibly in USA. SOLANUM LESTERI Hawkes & Hjerting. 2n=24. Oaxaca, Mexico. SOLANUM MICHOACANUM (Bitt.) Rydb. (syn.s. trifida Corr.). 2n=24. Michoacan and Jalisco, Mexico. In the pine forests and fields.resistant to the green peach aphid, Myrus persicae Sulzer. SOLANUM MORELLIFORME Bitt. & Muench. 2n=24. C. Mexico, southwards to Guatemala. SOLANUM OXYCARPUM Schiede. 2n=48. E. Central Mexico, Honduras, Costa Rica and adjacent Panama. SOLANUM PAPITA Rydb. 2n= fendleri*. Similar to S. SOLANUM PINNATISECTUM Dun. 2n=24. N. Central Mexico. A maize field weed. A source of resistance to Phytophthora, Y-virus and Colorado beetle. A parent of S. sambucinum*. SOLANUM POLYADENIUM Greenm. 2n=24. C. Mexico. A source of Phytophthora resistance and, owing to its glandular hairs, of resistance to tarsonemid mite (Polyphago tarsonemus latus Banks). SOLANUM POLYTRICHON Rydb. 2n=48, genome formula A4A4BB. NW. to N. Central Mexico. In waste places, shrubland and cultivated fields. Its genomes are related and also to the Ai genome of S. phureja*. S. bulbocastanum* has the genome formula BB. S. polytrichon is used as a source or resistance to Phytophthora and potato aphid. SOLANUM X SAMBUCINUM Rydb. 2n=24. In maize fields in N. Central Mexico. A natural hybrid of S. pinnatisectum* x S. cardiophyllum* ssp. ehrenbergii* (2n=24) and S. pinnatisectum* (2n=24) (Hawkes, 966).

179 MrvTr'AM Dtmmw NORTH AMERICAN REGION ATRIPLEX CANESCENS (Pursh.) Nutt. American shad scale, Hoary saltbush,wing scale, Fourwing saltbush. 2n=. W. North America up to Mexico. Cultivated as a fodder plant on saline soils and as a hedge plant (Mansfeld, 959). A. gardnesi (Moq.) Standi. (2n= )is from the same area and A. truncata (Torr.)A. Gray (2n= ) is from Utah, USA. CHENOPODIUM NUTTALLIAE Safford. 2n=4x=36. This species is named now Ch. berlandieri Moq. ssp. nuttalliae (Safford) Wilson & Heiser. It is cultivated in the highlands of C. Mexico and Arkansas and Missouri (USA). The weedy Ch. bushianum Aellen of E. North America is closely related (Wilson & Heiser, 979; Wilson, 98). Compositae HELIANTHUS ANNUUS L. Sunflower. 2n=4, genome formula Ba^Bai. N. America. It is difficult to be more specific because of its spread as a food plant and weed. The wild type may have resembled ssp. jaegeri, a small-headed type found in SW. Utah and NE. Arizona tos. California in USA. Innew areas the sunflower has introgressed with related species sothat the morphological variationhas increased. An important species is H. bolanderi A. Gray (2n= 34)of C. and N. California. It may derive from crosses of H. annuus with H. exilus Gray, 2n=. Other species H. agrophyllus Torr. & A. Gray (2n=34)of Texas, H. debilis Nutt. ssp. cucumerifolius (2n=34)of Texas and H. petiolaris Nutt. (2n=34)ofW. of N. America have also hybridized with the sunflower. Various subspecies and varieties are recognized: ssp. lenticularis (Dougl.) Ckll, which isnear to the original wild type and is found from W. Canada to N. Mexico. Ssp. texanus grows mainly in Texas. It may have arisenby hybridization of ssp. lenticularis and H. debilis. Ssp. annuus L. is a ruderal weed, a weed sunflower, found in the settlements in the Midwest. It possibly derives from ssp. lenticularis. Var. macrocarpus (DC.) Ckll is probably the parental form of the cultivated types. Itgrows in NE.USA and Canada. It probably originated from ssp. annuus or this subspecies and var. macrocarpus developed together from ssp. lenticularis. A weedy sunflower may have reached the Middle West where no other annual Helianthus species are found. Here the giant, large-headed sunflower may have developed (Heiser, 955, 965). From N. America the sunflower was introduced into Europe where in USSR a secondary centre of diversity arose (p. 30). HELIANTHUS EXILIS A. Gray. Serpentine sunflower. 2n=. Moist serpentine soils of Inner Coastal Range in California, USA. Almost extinct. H. bolanderi A. Gray, 2n=34, resulted from introgression of H. exilis genes into H. annuus* (Jain et al., 977). Cold-tolerant. HELIANTHUS TUBEROSUS L. Jerusalem artichoke. 2n=02, genome formula At-, At-, AtoAtoBt-jBt-,. N. America. Run wild in S. Ukraine and N. Caucasus. A perennial species introduced to Mexico and Eurasia. The Bti genome is related to the Ba^ genome ofh. annuus*. H. tuberosus isprobably an amphiploid of a species with genome formula AtiAt]At2At2and a B genome donor. This might be H. annuus or else a closely related species. H. x laetiflorus Pers. (2n=02), a wild perennial species of USA is probably a hybrid of H. subrhomboidus Rydb. (2n=02) and H. tuberosus. The first parent is also native to USA (Clevenger & Heiser, 963). IVA ANNUA L. (syn.i. ciliata Willd.). Sump weed, Marsh elder. 3n=34. A large-*seeded' type (var. macrocarpa)is an extinct oil-crop cultivated from the early st Millenium BC. to the first half of the 2nd Millenium AD. (Yarnell, 972). STOKESIA LAEVIS (Hill) Green. Stokes aster. 2n=. A perennial native to Georgia, Florida, Alabama, Mississippi and Louisiana (USA), grown as an ornamental. A possible oil-crop? Cucurbitaceae CUCURBITA FOETIDISSIMA HBK. Feral buffala gourd, Fetid gourd, Missouri gourd, Calabazilla. 2n=. Will probably be domesticated (Bemis et al., 978). Cupressaceae JUNIPERUS VIRGINIANA L. Eastern red cedar. 2n =. E. North America.Much variation is due to introgressive hybridization with other Juniperus species (Hemmerly, 970). Ebenaceae DIOSPYROS VIRGINIANA L. Common persimmon. 2n=. E. North America. Cultivated for its fruits. Also used as arootstock ofd. kaki*. Ericaceae VACCINIUM ASHEI Reade. Rabbiteye. 2n=72. N. America. Cultivated there. Wild plants are also harvested. According tocamp (945) the wild types derive from hybridization of the tetraploid species V. arkansanum, V. australe Small, Southeastern highbush blueberry, V. darrowi-4x and V. myrsinites Lam., Ground blueberry. VACCINIUM C0RYMBOSUM L. Highbush berry. 2n= 72. N. America. Cultivars have developed from the wild type which arose from hybridization of the tetraploid species V. lamarckii Camp (syn.v. angustifolium Ait.), V. alto-montanum Asche, V. simulatum and V. australe Small, Southeastern highbush blueberry.

180 Distribution of Helianthus spp. in pre-human (above), pre-colurabian (middle)and modern(below) times. (H. annuus ( ), H. petiolaris (speckled), H. exilis (A), H. argophyllus (B), H. debilis var. cucumerifolius ( ), H. debilis var. debilis (C), H. bolanderi ( ), cultivated sunflower (), campflower (2), Great plains annuus (3), weed petiolaris (4) and weed cucumerifolius(5) (Anderson, 956; based on Heiser's research). COMPOSITAE 20

181 NORTH AMERICAN REGION crop and green manure. ROBINIA HISPIDA L. 2n=30. SE. North America. Cultivated as an ornamental and in hedges. ROBINIA PSEUDACACIA L. Locust, Black locust. 2n=20, c. 20, 22. C. and E. North America. This tree is planted as an ornamental and as asoil stabilizer. SESBANIA EXALTATA (Raf.) Rydb. 2n=2. USA. Cultivated there as a green manure. SESBANIA MACROCARPA Muhl. 2n=2. USA. Cultivated there. Closely related to S. exaltata*or is a synonym of this species. VICIA LEAVENWORTHII Torr. & Gray. Leavenworth vetch. 2n=4. Missouri and Arkansas to Texas in USA. Occasionally cultivated. Liliaceae CAMASSIA LEICHTLINII (Baker)S. Wats. Camas. 2n=30. E.North America. Uncultivated bulb beds are divided into family plots, which have been passed down from generation to generation. These plots are not farmed,but stones, weeds and shrubs are removed every year. In most cases the plants are marked in bloom so that the bulb can be harvested when it is fully grown (Chapman Turner & Bell, 97). The camas is semi-domesticated i.e. the wild plant isprotected and the growing circumstances are improved. The latter may result in more sites for the plant to grow. CAMASSIA QUAMASH (Pursh) Greene. Blue camas. 2n=. E.North America. See further C. leichtlinii*. Limnanthaceae LIMNANTHES ALBA Benth. Meadow foam. 2n=0. NW. Pacific states of USA. Potential oil-crop. Erect types have been developed for cultivation. Self- and cross-compatible. Malvaceae GOSSYPIUM BARBADENSE L. Sea Islands cotton. 2n=52, genome formula (AADD) 2. Peru (p. 76). Sea Islands cotton developed on the Sea Islands of S. Carolina, USA after introduction from Bahamas or Jamaica. Cultivated in E.USA and some Caribbean islands. Maybe similar types in Ecuador are its parental material (Harlan, 970). Martyniaceae PROBOSCIDEA LOUISIANICA (Mill.) Thell. (syn. Martynia proboscidea Glox.). Unicorn plant, Ram's horn, Double claw, Proboscis flower. 2n =. N. America. A herb cultivated for its fruits and as an ornamental. Moraceae MACLURA POMIFERA (Rafin.) CK. Schneider (syn. M. aurantiaca Nuttall). Osage orange. Texas, Oklahoma and Arkansas, USA. Widely planted as living fence inusa from Naturalized. Also used as a source of bows, yellow dye and building material. Dioecious, wind-pollinated. Macludrania hybrida André is a natural hybrid of Cudrania trisuspidata x Maclura pomifera var. inermis (Smith & Perino, 98). Passifloraceae PASSIFLORA INCARNATA L. May-Pop, Yellow-fruited Virginian passion flower, Apricot-Vine. 2n=8, 36. E.North America, Florida and Texas.It was cultivated by indians in Virginia. Resembles P. edulis*. Phytolaccaceae PHYTOLACCA AMERICANA L. 2n=36. USA. It has run wild ins. Europe. Cultivated as adye plant (berries) and ornamental. The berries are also used to colour wine. Rosaceae AMYGDALUS PERSICA L. Peach. 2n=6. Primary centre : China (p. 42). Secondary centre : California, USA. FRAGARIA CHILOENSIS* FRAGARIA OVALIS Lehm. 2n=8x=56. W. North America. Used in breeding F. x ananassa*. FRAGARIA VESCA L. Wild strawberry. 2n=4, genome formula AA. Europe (p. 59), Asia and N. America. In California, it is found in a broad range of environments from the coastal fog belt to the Sierra Nevada and Cascade Mountains and from San Jacinto Mountains near the Mexican border to Oregon. Hermaphroditic, self-compatible, reproducing sexually and asexually by runners. There are many ecotypes, which aid survival under this wide range of environments (Hancock & Bringhurst, 978). Some clones are used to test for viruses inf. x ananassa*. FRAGARIA VIRGINIANA Duch. Virginian strawberry. 2n=56. N. America, especially in the eastern part. Cultivated. One of the parents off. x ananassa* Duch. (syn.f. grandiflora Ehrh.), the pineapple strawberry (2n=56). PRUNUS. The American Prunus species are valuable because of their longevity and winterhardiness. PRUNUS AMERICANA Marsh. American Plum. 2n=6. A large territory of N. America between Manitoba and Texas. This small tree usually grows slowly. Cultivated. Valuable because of its longevity and winterhardiness. Cultivars have been developed from interspecific crosses.

182 LEGUMINOSAE - SOLANACEAE 205 PRUNUS ANGUSTIFOLIA Marsh. (syn. P. chicasa Mich.). Chickasaw plum, Florida sand plum, Mountain cherry. 2n-6. S. Delaware to Florida and westward to thetexas and S. Oklahoma. A small tree with a dense crown. The cultivated species lack hardiness. It hybridizes easily with P. munsoniana*. PRUNUS BESSYI Bailey. Western sand cherry. 2n =6. N. America on sandy and saline soils. This perennial shrub is characterized by a good longevity, frost resistance and sweet, edible fruits. A source of a bush-type habit forease of mechanical harvesting.lt hybridizeswith Armeniaca vulgaris*and Prunus ssp. PRUNUS HORTULANA Bailey. Hortulan plum. 2n=6. C. USA. This tree is very cold resistant and has good fruits. It flowers late (Zylka, 970). It hybridizes easily with other American Prunus species. According to Bailey (898) it is a hybrid ofp. angustifolia* andp. americana*. PRUNUS MARITIMA Marsh, (syn.p. maritima Wangh., P. acuminata Michx.). Beach plum, Sand plum. 2n=6. E.USA and adjacent Canada. Some cultivars have been selected (Zylka, 970) in the USA. Alsoused as asource of late flowering, cold resistance and very high fertility in the breeding of other cultivars. It is an ornamental. PRUNUSMUNSONIANA Wight & Hedrick. Wild Goose Plum. 2n=6. N. Texas, E. Oklahoma and Missouri. It flowers late and the fruits are of good quality. It has agood longevity and winterhardiness. It hybridizes easily with P. angustifolia* and other species. Cultivars were bred from such interspecific crosses. PRUNUS NIGRA Ait. Canada plum. 2n=6. The territory between S. New Foundland to the Strait of Mackinac and southward to Lansing, Michigan. Cultivated there to some extent. This species is valuable because of its longevity and winterhardiness. Cultivars have been developed from interspecific crosses. PRUNUS PENSYLVANICA L. (syn.p. persicifolia Desf.). Bird cherry, Pigeon cherry, Pin cherry, Wild red cherry. 2n=6.N. America, from New Foundland to British Columbia and Colorado. A source of late flowering and cold resistance. PRUNUS SEROTINA Ehrh. Black cherry, Run cherry. 2n=32. N. America from Ontario and North Dakota to Texas and Florida. The fruits are unpalatable. It could be a source of late flowering and frost resistance. PRUNUS VIRGINIANA L. Common choke cherry, Eastern choke cherry, Choke cherry. 2n=30, 32. W. North America. Micurin selected Vinogradnaja from this species (Zylka, 97b). RUBUS ACAULIS Michx. 2n=4. The Canadian counterpart ofr. articus (Larsson, 969). It could be one parent of R. stellatus*. Also named ssp. acaulis ofr. arcticus. RUBUS FLAGELLARIS Willd. (syn.r. villosus Ait.). Northern dewberry. 2n=63. E.North A- merica. Cultivated for its fruits. Var. roribaccus Bailey is the Lucretia dewberry (Uphof, 968). RUBUS IDAEUS L. (syn.r. strigosus Michx.). American red raspberry. 2n=4. The NE. American counterpart of this species. Present cultivars are often hybrids between this subspecies and ssp. vulgatus*, the European red raspberry. Natural hybrids between ssp. strigosus and R. occidentalis* have been described asr. neglectus Peck., Purple cane raspberry (2n=4). They have been occasionally cultivated. In N. America such hybridization has led to introgression between the two parental species.see for other hybrids p. 4. Cultivars are often unarmed or have simple leaves. The loganberry (2n=42), genome formula V-jViV^V^H, is derived from the cross of a tetraploid form R. ursinus Cham. & Echt. (2n=28), genome formula VVV2V2 and R. idaeus (2n=4). Mayberry is a hybrid product ofr. palmatus Thunb. x ssp. strigosus and Youngberry (2n=42, 49) of loganberry x Mayes dewberry (R. baileyanus xr. argutus). Other Rubus species like R. arcticus* have also been used in breeding work. RUBUS OCCIDENTALIS L. Black raspberry. 2n=4. NE. North America, Colorado and British Columbia. It is cultivated. Present cultivars often derive from hybrids with R. idaeus*. Natural hybrids with the N. American ssp. strigosus of R. idaeus have been named R. neglectus Peck., purple cane raspberry (2n=4). This hybridization has led to introgression between these two parental species. It is forr. idaeus* a source of resistance to the rubus aphid, Amphorophora rubi Kalt. RUBUS STELLATUS Sm. 2n=4. Wild from Alaska, Aleutian Islands and Kamchatka. Closely related to R. arcticus*and has been described as ssp. stellatus of this latter species.it could be a hybrid of R. arcticus and R. acaulis* (Larsson, 969). Salicaceae SALIX RIGIDA Mühlenberg (syn.s. cordatamiihl. ) American willow. 2n=44. N. America. Introduced in Europe for twig production. Solanaceae DATURA STRAMONIUM L. Thorn-apple, Jimson weed. 2n=24. N. America. Pantropical now. A poisonous plant cultivated as a medicinal plant yielding stramonium. NICOTIANA QUADRIVALVIS Pursh. 2n=48. SW. USA along rivers and in rocky soils. The Indians cultivated it for smoking leaves and flowers.

183 NORTH AMERICAN REGION An annual, frost-resistant, early maturing species. It hybridizes easily with N. tabacum*. PHYSALIS PUBESCENS L. Strawberry tomato, Dwarf Cape gooseberry. 2n=24. N. America. Cultivated in Ukraine and elsewhere. SOLANUM FENDLERI A. Gray. Navajo potato. 2n= 48, genome formula A4A4BB. Arizona, New Mexico and W. Texas,USA and NW. Mexico.Very similar to S. hjertingii* and similar to S. papita*. Itis resistant to Y virus. SOLANUM JAMESII Torr. 2n=24, 36. Arizona, New Mexico and Colorado (USA) and in Mexico near the Arizona boundary. Valerianaceae VALERINA EDULIS Nutt. 2n=. N. America. This perennial herb is cultivated for its roots. Vitadaceae VITIS BERLANDIERI Planch. 2n=38. SE.USA to Texas. Used as a rootstock. Rootstocks of V. riparia* x V. berlandieri are also used. It can be crossed with V. vinifera*. VITIS CINEREA Engelm. Downy grape, Sweet winter grape. 2n=38. SE. USA. Resistant to fungal diseases and to phylloxera. Viteus vitifolii Shimer, but it cannot be crossed with V. vinifera*. VITIS CORDIFOLIA Michx. 2n=30, 38. SE. USA. It canbe crossed with V. vinifera* to introduce resistance to fungal diseases. VITIS LABRUSCA L. Fox grape. 2n=38. E. North America. It appears to have runwild in Georgia,USSR (p. 02). Introduced into the Old World. It is cultivated. It has been crossed with V. vinifera* to improve it and for this species it is used as a rootstock. VITIS RIPARIA Michx. (syn.v. vulpina L.). 2n=38. E. and C. USA and in Ontario, Canada. Used as a rootstock. It can be crossed with V. vinifera* to introduce resistance to phylloxera, Viteus vitifolii Shimer. Such hybrids occur wild in USA andhave been described asv. bourquina. Rootstocks of V. riparia x V. berlandieri* are also used. VITIS ROTUNDIFOLIA Michx. Muscadine grape, Southern fox grape. 2n=40. Florida, the southern coast of USA and the east coast of Mexico. Dioecious, but hermaphrodite cultivars have been developed. Cv. Scuppernong was developed in 584 from wild material in NE. North Carolina. VITIS RUPESTRIS Scheele.2n=. SW. USA. Used as a rootstock. It can be crossed with V. vinifera* to introduce resistance to phylloxera, Viteus vitifolii Shimer.

184 Species without an identified region Some species could not be listed inoneof theregions.they have avery wide geographical distribution. Either theirlocality of cultivation hasnot been reported or they arecultivated atseveral places.thus itis notknown whethertheir wide distributionoccurred before their domestication or not, i.e.it isnot mentioned whether the wild species grew in alarge area where ithasbeen domesticated atseveral places, or whetherwild plantswere domesticated on one site afterthey had been spread by man. Agavaceae YUCCA GLORIOSA L. (USSR). Amaranthaceae 2n=50. Cultivated in Georgia, GOMPHRENA GLOBOSA L. Batchelor's button.2n= 32, 40-44, The tropics. Cultivated in some villages of mainly coastal districts in W. Africa as a curiosity and as a fetish plant. In Europe and elsewhere it is an ornamental. Anacardiaceae SPONDIAS CYTHEREA Sonner (syn.s. dulcis Forst.f.). Otaheita apple, Ambarella,Hog plum. 2n=. Cultivated for its fruits. Mansfeld (959)reported the presence of var. cytherea on the Society Islands, Tahiti, Fidji, Samoa and Madagascar, var. mucroniserrata (Engl.) Mansf. in Mexico, var. macrocarpa (Engl.) Mansf. in Brazil, var. acida (BL.) Mansf. in Malaysia and var. intégra (Engl.) Mansf. in Amboina. Apocynaceae APOCYNUM VENETUM L. (syn. A. sibiricum Pall, ex Roem. & Schult.). Kendyr. 2n=6, 22.Italy to E. Asia. A perennial fibre crop brought intocultivation in USSR. Chenopodi aceae CHENOPODIUM BOTRYS L. Jerusalem oak. 2n=8. S. Europe, Orient and C. Asia. Occasionally cultivated (Mansfeld, 959). Corynocarpaceae CORYNOCARPUS LAEVIGATUS J.F. Forst. & G. Forst. Kopi, karaka. 2n=44. This species occurs wild on thenew Hebrides and New Caledonia asc. similis and C. dissimilis respectively. The large kernels can be eaten after removing the poisonous karakin by heating. Therefore they were taken to New Zealand (Stevenson, 978) where the species ran wild and became semidomesticated. Cruciferae CRAMBE ABYSSINICA Höchst, ex. R.E. Vries.2n= 90. Its wild distribution is obscure, but it may have developed in Turkey. Cultivars have been released in USA. See p. 89. Cucurbitaceae MOMORDICA BALSAMINA L. Bitter gourd, Bitter cucumber, Balsam pear, Balsam apple. 2n=22. Tropics of the Old World, especially India and SE. India. Leaves and stem are used for fodder. MOMORDICA CHARANTIA L. Bitter gourd, Bitter cucumber, Balsam pear. 2n=22. Tropics of the Old World. Naturalized in nearly all tropics and subtropics. An edible, medicinal and toxic plant (Morton, 967). Cyperaceae CYPERUS ARTICULATUS L. 2n= its sweet scented roots.. Cultivated for MARISCUS UMBELLATUS Vahl. 2n=. The tropics of the Old World. Cultivated for its rhizomes. Euphorbiaceae EUPHORBIA TIRUCALLI L. 2n=20. Trials at Arizona (USA) to grow this plant for its sap as a raw material for petrol. Gramineae EREMOCHLOA OPHIUROIDES (Munro) Hack. 2n=28. Some cultivation on Puerto Rico.

185 208 SPECIES OF A NOT IDENTIFIED REGION HETEROPOGON HIRTUS Pers. (syn. H. contortus Beauv. ex Roem. & Schult., Andropogon contortus L.). Spearhead, Tangle grass. 2n=20, 40, (42-44, 50), 60, (80,c ). The Old and New World, subtropics and tropics. Cultivated as a fodder for live stock. Plants with 2n=20 come from India, with 2n=40 from Java, India, Madagascar, Tanzania, Kenya, Uganda, Zimbabwe and Zaïre,andwith 2n=60 from S. Africa, Mexico and N. America. Leguminoseae CROTALARIA MUCRONATA Desv. (syn.c. striata DC.). 2n=6. Distributed in tropics and subtropics as a cover crop andgreen manure. CROTALARIA RETUSA L. 2n=6. This fibre crop is widespread throughout the tropics. In E. Africa it is used for its pigments and in Florida as an ornamental. DESMODIUM ADSCENDENS DC. 2n=22. T r o p i c s. Cultivated in many regions as a cover crop and green manure. found in the savanna of Venezuela and Colombia, and it is widely used as fish poison and ornamental, and in fetish. In Panama, it is cultivated as a fish poison (Nee, 979). Umbelliferae C0NIUM MACULATUM L. Hemlock, Poison hemlock. 2n=6, 22. Europe, Asia, N. Africa and Ethiopia. Occasionally cultivated. Often occurring as a ruderal. Used as a poison and medicinal plant. Urticaceae BOEHMERIA STIPULARIS Wedd. Hawaian false nettle, Akola. 2n=. Probably from the Mascarene islands. According to Uphof (968) this fibre crop grows wild on Hawaii, where it was formerly cultivated. This has resulted in many varieties. TOUCHARDIA LATIFOLIA Gaudich. Olona. 2n= The Hawaian Islands. At one time cultivated for its fibre (Hutchinson, 962). DESMODIUM GANGATICUM DC.2n=. Trop. Asia and Australia. Cultivated as a fodder plant and green manure. INDIGOFERA HIRSUTA L. (syn. I. schimperi Jaub. & Spach.). Hairy indigo. 2n=6. Many tropical countries. It is cultivated. TEPHROSIA PURPUREA Pers. Purple tephrosia. 2n =22, (24, 44). The tropics. Cultivated as a green manure and cover crop.grows wild in N. India on waste places and road sites. Malvaceae ABUTILON GRAVEOLENS Sweet. 2n=4, 36. The tropics of the Old World. Cultivated especially inussr for its oily seeds. MALACHRA CAPITATA L. 2n=56. The tropics. A herb cultivated for its fibre. URENA LOBATA L. Aramina fibre, Congo jute. 2n=28, 56. Wild or naturalized in the tropics and subtropics. Centre of origin very likely in the Old World (Purseglove, 968). Rutaceae EVODIA HORTENSIS J.R. &G. Forst, (syn. Fagara euoda L.f., F. evoda L.f., Zanthoxylum varians Benth.). 2n=. A widespread shrub. Cultivated in many parts of the Pacific. Leaves are used for medicinal purposes. Solanaceae SOLANUM MAMMOSUM L. 2n=. It spread as a campfollower over C. and S. America. Its native habitat is not clearbut it appears to be wild in the Caribbean. Primitive types are

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