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Home , Lima bean, Phaseolus vulgaris

Corpoica. Ciencia y Tecnología Agropecuaria ISSN: 0122-8706 [email protected] Corporación Colombiana de Investigación Agropecuaria

Debouck, Daniel G. Colombian Common and Lima : Views on their Origin and Evolutionary Significance Corpoica. Ciencia y Tecnología Agropecuaria, vol. 1, núm. 1, octubre, 1996, pp. 7-15 Corporación Colombiana de Investigación Agropecuaria Cundinamarca, Colombia

Available in: http://www.redalyc.org/articulo.oa?id=449953017002

How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative GUESTAUTHOR ColombianCommon and Lima DanielG.Debouck' Beans:Views on their Origin and Evolutionary Significance

ABSTRACT This article reviewsthe geographical Título:Los Fríjoles Colombianos Limay Común: Puntos deVista de su Origen distribution of wild common and lima y el Significadode su Evolución beansin the Neotropics,their morphologicaland ecologicalattributes, Sepresenta una revisión de la distribución geográficade las formas silvestresdel fríjol and their biochemical and molecular común y del fríjol lima en el Neotrópico,de suscaracterísticas morfológicas y ecológicas, variationalong their ranges.These facts y de la variación bioquímica y molecular a lo largo de estadistribución. Estoshechos revealthe organization of the genetic muestranque la diversidadgenética viene organizadaen tresacervos o grupos de genes, diversity into three major genepools, uno de ellossiendo ancestral, y con subdivisionesadicionales dentro de los acervosde- with one being consideredancestral, and rivados.Se discuten las relacionesfilogenéticas entre esteramal ancestraly las especies additional subdivisionswithin the afines.Más que lugar de contacto entre acervosde materialescultivados, Colombia derived ones.The relationshipsbetween aparececomo corredor biológico donde transitaron las ramas ancestrales,y como lu- the ancestralbranch and related species gar de posibledomesticación. arediscussed. Colombia appears to be more than a placeof contact between PalabrasClaves: ,especies silvestres, distribución geográfica,marcadores mole- genepools of cultivated materials,but culares,evolución de cultivos,filogénia the transit place of the ancestral branches,and a possibleplace of domesticationas well. 1.INTRODUCTION Key words: Phaseolus,landraces, wild T) Dro*, HAVELoNc beenknown as netic diversit¡ and thus in the decision species,geographic distribution, part of the major food usedby the making processof conservationof such a molecular markers,crop Amerindians (DeCandolle,1883; Vavilov, geneticdiversity. evolution, phylogeny r93t,t939).Perhaps under the influenceof When a'second generation'group of their fellow historians who were much scholarshad accessto more archaeologi- impressedby the prestigiouspre-Colom- cal information and data from ex- bian civilizations existingjust at the mo- plorations carried out mostly prior to ment of the Conquest,scholars have long World War II, the view that two centersof focusedtheir attention on diversity for beans,most likely indepen- (whosegeographic boundaries are here dent,was further developed,but wassoon definedas per León, g9z),in particular in need of additional. novel evidence ,and the CentralAndes,in particu- (Heiser,t965,1979). During the r96osand lar .In doing so,other regionsas well r97os,additional material was collected, as the right time perspectivemight have fortunately enough in a broader perspec- been overlooked,in order to bring full tive than before, and with the advent of clarity on eventssuch as crop origin, cul- molecular geneticsand electrophoresis tivar diversit¡ flows of materials,and pat- techniques,many more comparisonshave terns of .Because of their become possible,across gene pools or natural distributionsin both Mesoamerica across closely related species,namely and the Central and beyond where morphological or physiological (Debouckand Smartt,1995), the two Neo- variation was either inexistent or poorly tropical (Phaseolas,Phaseolinae, expressedin phenotypes. )species of worldwide economic Although we are still perhapsfar from importance,P. vulgarisL., the common a completeunderstanding of bean evolu- bean,and P.lunatusL.,the ,of- tion over the past thousandsof years,it fer the possibility to challengefurther might be temptingfor a'third generation' theseearly conceptsof crop origin and student of bean evolution to sum up evolution, and would allow us to more wherewe are,what we know less,and im- fully answer the questions: from which plications for conservationand use of material did thesecrops arise?,where?, beangermplasm. I shallconsider succes- attributes, r. Genetic ResourcesUnit, cIAT, Apartado Aéreo how?,when?, by whom?, all of them of sivelygeographic and ecological 67y,Cali, Colombia critical importance for the shapingof ge- geneticvariation, and evolutionary histo-

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ries,with main emphasison wild ancestral Bosquehúmedo subtropical templado in ingly enough, a few forms of Southwest- forms of thesebean species, and I havethis Guatemala(de la Cruz S.and Sagastume ern Mexico in Mexico, but also of Costa time focusedthis essaydeliberately on L., 1983),in Bosquemuy húmedomontano Rica and Colombia tend to behave as Colombia. The reasons for doing so bajoin CostaRica (Tosi,1969), in Bosque short-living perennialsregrowing after the should become hopefully obvious to the húmedomontano b aj o inYenezaela(Ewel first seedset, with subwoodylower stems readershipin this essay. et al.,tg7 6), in Bo sque húmedo pr emontano and fibrous roots. Someforms of Costa in Colombia (Anonymous, r988), in Rica, Colombia and Northern Peru are z. Featuresof wild common bean Bosque seco premontano in Ecuador alsoparticularly late for their first - before Humans (CañadasC. and EstradaA., 1978),in ing. Not surprisingl¡ differencesare also 2.1.Distribution of wild commonbean: Bosqueseco montano bajo tropicalin Peru reflectedin physiologicalparameters, such an anomalyin theAndes (ONERN, t976),in Bosqueseco montano as the ones controlling photosynthesis The wild common bean is presentlY bajoín (Becket al.,ryy).It grows (Lynch et al.,rygz). Accordingto thesere- distributedin the westernmountainous rn Distrito de las SelvasMontañas of sults,wild bean populations from Mexico rangesof the Neotropics,from Chihuahua Provincia de las Yungasin that showadaptation to more open,sunny, in Mexico to San Luis in Argentina, gen- (Cabrera,ry76).In thesehabitats, where savannah-habitatsare different from the -2,ooo erally at \4oo-z,2oo masl (Gepts and the amount of rainfall ranges4oo onesof Guatemala,Peru and Argentina. Debouck,r99r; Toro Ch. et al.,r99o). It is mm/year but concentratedat the begin- relatively abundant in western Mexico, ning of the growingseason, soil natureand 23. An impressiveuariation at bio' particularly along the Eje Volcánico fertility vary a lot but with topsoil slightly chemicaland molecularlevels (Nayarit,Jalisco, Michoacán, Guerrero), acidic (pH 6.2-6.2)and rich in organic The variation of wild P. vulgaris re- much lessin Central America (Delgado matter. Given these ecological affinities, vealedby ecological,morphological and Salinaset al.,1988).In theAndes, its range the gap in SouthwesternColombia is less physiologicalcharacteristics is also re- starts in western Venezuela,and from understandable- and certainlymerits fur- flected at the biochemical (seedproteins, there extends into Colombia uP to ther attention, since according to data allozymes)and molecular (RFLPs,AFLPs) Cundinamarca(Debouck et al., 1993). publishedelsewhere (Anonymous, 1988) levels.An important and easy-to-detect There is a gap in southwesternColombia the Bosqueseco premontano exists between polymorphism has been observedin the (Cauca,Nariño) that needsadditional sur- El Cerrito and Praderain the Valle del main storage , phaseolin, with veys.The rangeresumes in Chimborazo, Cauca and in the upper valley of Río more than twenty electromorphsfound in Ecuador,and alongthe Pacificslope ofthe Guaitarain the Nariño department. Mesoamerica,slightly lessin the Central Andes, extends to Cajamarca, Peru The almost continuous habitat of wild and Southern Andes (Toro Ch. et al., (Deboucket al.,1989a). P vulgarisand its abundancein the Neo- r99o).In Mesoamerica,wild beansshow 'S' The distribution turns again to the tropical mid-altitude subhumid moutain and several'M'qpes(Gepts et al.,t986; easternslope of theAndes,from Huanuco forestsraises the following question: has Koeniget a1.,r99o; Toro Ch. et al.,t99o), in Peru through Central EasternBolivia the wild common bean seenits habitat al- while in the Central and SouthernAndes and ends up in San Luis in Argentina teredby human activitiessuch as logging, of Peru(]unín,Apurimac, Cuzco), Bolivia (Berglund-Brücl-rerand Brücher, 1976; periodic fires,clearings for shifting agri- (Cochabamba,Chuquisaca, Tarija) and ToroCh. et al.,r99o). It is so far unknown culture?It seemsthat perhaps up to half Argentina (Iujuy, Salta,Tucumán), they from Chile and likely not present in that of the habitats where wild P vulgaris show'T','Cl'H','A and'f' t1pes(Gepts et country (Debouck,personal observations, thrivesin recentdecades have been altered, al.,1986; Koenig et al.,r99o; Tohme et al., r99ó).One shouldnote that if the rangein so that it has benefitteclfrom periodic 1989;Toro Ch. et al.,r99o). So,given the Mexico and Guatemala,and in Argentina, human alterations(Delgado Salinas et al., attributes ofphaseolin asan evolutionary is known sincethe r94os(Burkart,7g4r; 1988).But along its range,some popula- marker(Gepts and Bliss,r985b), two gene McBryde,1947, respectively), the Pacific tions thrive in true climaxvegetations, that poolswere recognized. More subtlediffer- rangeof distribution of wild P.vulgarisin aredry variantsof Lower montane humid ences were revealed afterwards using has been disclosedonly forest or Dry montane forest.So, many RFLPson mtDNA (Khairallah etal,t99z), recently(Debouck et al.,1993). populations growing in placesso different namely that Guatemalan wild forms and distant asthe statesof Mexico,Nayarit might be different from the ones in forms of Bolivia and z.z.A movingfrom theforest into or Jaliscoin Mexico, or in Tárija, Bolivia, Mexico,and that the savannahs or in luju¡ Argentina, truly belong to cli- Argentina would be much alike. The wild common bean is a viny le- max,original forests. Colombian wild beansdisplay two in- sunnypine-oak forests and In addition to ecologicaldifferences, terestingfeatures. First, they displaysome gumein open, 'S','CH' oak grasslandsin Mexico and Central morphological differencesexist along the phaseolintypes such as (Geptsand Americawith many speciesof Compositae rangeof distribution of 8,oooKm, with Bliss,1986), that are presentelsewhere in and Solanaceae(Delgado Salinaset al., many intermediaryforms betlveenthe two Mesoamerica(Toro Ch. et al.,r99o). Sec- 1988;Gentr¡ 1969),while its SouthAmeri- geographicalextremes: forms with large ond, they alsodisplay phaseolin types such can habitat is somewhatmore shadyand and obovateleaflets, racemes with many as'f that areapparently unique to Colom- humid in montaneforests with speciesof floral insertionsand largeheart-shaped bian materials (Chacón S. et al., 1996). Podocarpus,Alruts, Cebis and Schinus bracteolesin Mexico (Gentr¡ 1969),and Recentwork using AFLPs markers has (Brücher,1988; Debouck et al., 1993).So, forms with small rhomboedric leaflets, shownthat for thosewild beansof Colom- according to a Iife zone classification racemeswith few floral insertions and bia sharing'Mesoamerican'phaseolins dif- (Holdridgeet al.,r97r), it thrivesinBosque small triangular bracteolesin the Central- ferences at the DNA level for húmedo montano baio subtropicaland SouthernAndes (Brücher, 1988). Interest- non-phaseolincoding genes are important

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(ChacónS. et al.,1996, Tohme et al.,1996), the wild, that is the accumulationof small entitiesis quite remote.Data accumulated indicating that these Colombian wild mutations, translocationsand inversions on cpDNA (Llacaet al.,1994; Schmit et al., forms arealready distinct from their Cen- in the DNA molecules (that can now be 1993)among various materialsof P. vul- tral Americancounterparts. revealeddirectly) and so in the genescod- garis show a very low level of polymor- In contrast,Ecuadorian and northern ing for certainproteins or enzymes.There phism, compatible with that observedin Peruvian (Piura, Cajamarca)wild forms are howevercases where undoubtedly in- other speciesof Spermatophyta(Soltis et "mismatches" displaylittle variation in phaseolin,since trogressionhas produced in al.,t99z).TWo genesis scenarios were thus a single'I't1pehas been described for the the expectedpatterns, and it is likely that possible:either i) one genepool derives different populations analyzed so far with circulation of cultivated varieties from the other, or ii) the two genepools (Deboucket aI.,t993; Koenig et al.,r99o; aroundthis phenomenon may increase in arisefrom a third one, that could be seen Neema et al.,:-994).Diversity analysisus- the future.So,'S'phaseolin was observed asthe ancestralpool. Sincem ostPhaseolus ing allozymesconfirms their unique po- in wild and weedy forms present in speciesare today distributed in Mexico sition in comparisonto the two major Apurimac,Peru (Gepts et al.,1986),likely and Central America (Debouck, r99r; genepools (Koenigand Gepts,1989), but coming from the human-introduced DelgadoSalinas, 1985), it lookedplausible 'Panamito'well it was then not possible to conclude distributed this century in that P. vulgaris originated first in whether that uniquenessresulted from Coastaland CentralPeru (Voysest,1983).Mesoamericaand that the Andean gene hybridizations between the two major In spite of the presenceof reproductive pool aroseas a branch deriving from the gene pools becauseof contact through barriersdue to a long evolutionin isola- Mesoamericanpool. The alternatesce- millenia or was due to any other factor. tion (Geptsand B1iss,r985a; Gutiérrez and nario - that the wild common bean arose Later, numerous and unique polymor- Singh,rg8¡), someintrogressive hybridiza- first in South America and then migrated phismshave been revealed using RFLPs on tion betweengene pools is possible(for towards Mesoamerica- was a more re- mtDNA in thesematerials in comparison instancein the Chilean material: Paredes mote possibilit¡ although once consid- to thoseof other regions(Khairallah et al., and Gepts,1995), because the common eredby some(Burkart and Brücher,1953). r99z), giving support to the hypothesis beanis not yet a closed,compartmental- Both scenarioswere however challenged that factorsdifferent from simple contact ized biological system. by the molecular complexity of the and resultinghybridization causeoriginal- phaseolinmarker (Gepts,1988), and the ity of that range of wild materials. 2.5.Two or threemajor gene pools? fact that the dominant Mesoamerican The geneticvariation in wild common Prior to the discoveryof the wild beans phaseolin'S'was slightly more complex bean is thus strongly structured along of the PacificAndean range (Debouck et than the dominantAndean type'T' (Bliss spacegradients, at the regionallevel first al.,r989a), it seemedlogical to organizethe and Brown, 1983).In other words, these (with an Mesoamericanand an Andean diversityof wild P. vulgarisinto two gene phaseolinscould not derive straight from gene pool, and even perhaps a North pools,one centeredin Mesoamerica,and each other. In addition, RFLP polymor- Andean gene pool), at the subregional another one in the Central and Southern phismson mtDNA betweenMesoamerica level next (with differenceswithin each Andes (Gepts et al., 1986;Koenig and and the Central-Southern Andes genepool). The hypothesisthat wild bean Gepts,1989). If subdivisionsand regional (Khairalfah et al., ry92) were important populations growing in mountainous differencescould be recognizedwithin enough as to raisedoubts about a simple rangesevery time separatedby geological eachmajor genepool (for instance,be- migration scenario. There were however fractures are different (Bannerot and tween Mexico west of the Isthmus of two casesreported of a direct similarity Debouck,r99z) has gained some support. Tehuantepecand the entity Chiapas-Gua- betweenArgentinean and Mexican wild As analysisprogresses and more perfor- temala: Bannerot and Debouck, t99z; beangermplasm: i) the collectionDGD- mant markersare becoming available, it is Chacón S. et al., 1996)- and likely will be 6zr of Jujuy displaysthe Mexican alleleof likely that differencesshall be revealed revealedfurther (Tohmeet al.,1996), the diaphorase(Koenig and Gepts,r98q), and eventuallyat the level ofeach population. presenceof two major genepools is today ii) DGD-6zg of Saltadisplays a RFLPpoly- little questioned(Becerra Velásquez and morphism presentin awild Mexicanform a.4.Can we trust the biochemicaland Gepts,1994; Gepts, 1993; Hamann et al., of Guerrero (Khairallah et aI.,tggz). molecularvariation and how did it t995; Zink et aI., ry94). Not surprisingl¡ arise? the organizationof diversityin the host 2.6.In searchof the ancestralbranch Given the level of outcrossingin wild plant wasalso paralleled with that of some In contrast with the situation prevail- common bean (Tiiana et al.,r993;Yander- fully specificpathogens - evidencedon ing in Mesoamericaand in the Central- borght,1983), one wonderswhether such cultivatedforms - suchas angular spot SouthernAndes, we have seenthat the a biochemicaland molecularvariation is (Guzman.et al., ry94) and anthracnose wild beansofEcuador and northern Peru not the result of introgressivehybridiza- (Pastor-Corraleset al., 1995),where two show no variation in phaseolintlpe but a tion with cultivatedbean varieties or even genepools can alsobe recognized. single'I'type. This phaseolintype is out- other Phaseolusspecies (the work by Wall The existenceof two major genepools standingbecause of two characteristics. and Wall, 1975,shows that this possibility - at the level of wild forms - unavoidably First, it lacks the 5zkD high molecular is plausible).The molecular complexity of raisesthe question of their genesis.It weight subunit of phaseolin(Koenig et al., many markers,the uniquenessof many seemsfirst that P. vulgarlsas a speciesdid r99o). In a comparativestudy of nucle- electromorphsand their absencein any not ariseby morphologicalconvergence. otide sequencesresponsible for the spr- 'I','S'and'T'phaseolins, cultivated common bean variety so far In other words, the possibility that two thesisof (Kami et 'I' givessome insurancethat the polymor- wild legumescompletely different from an aI.,99) showedthat phaseolingenes phisms observedare the true reflect of evolutionary perspectivegave rise to two Iack a z7-basepair tandem direct repeat, geneticvariation accumulatedover time in morphologically and geneticallyso close whereasthe'S' and'T'phaseolingenes do

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'simple'phaseolin haveit. This study togetherwith resultsof typeswould existin the 84oo7'W-1o8o3o'W,while of 64"r9'W- phaseolin gene sequencing(Kami and Northern Andes among more comPlex 79"24'W in the Andes (Toro Ch. et al., Gepts,1994) indicates that genesequences ones,with the possibility that the latter r99o).Second, P. vulgarisas it cameinto responsiblefor the synthesisof phaseolin would have derived from the former by being did not migrate alone further into have gained complexity by endorepli- duplication of nucleotide sequences Mesoamerica,but wiü comPanionspecies cation,instead of loosing sequences(the (Kami and Gepts, 1994).On the other such as P. costaricensi.sand P polyanthus, alternatescenario, less plausible because hand, aswe haveseen, all presentlyknown that would eventually share the same random deletions would have prevented speciesbelonging to the samecpDNA lin- cpDNA polymorphisms (Schmit et al., or hinderedphaseolin formation - appar- eageas P. vulgaris,exist today in Central 1993),or evenP. coccineuswith which it ently a selectiveadvantage in this species). America. With presently availableevi- would share some mtDNA sequences Theseresults have been confirmed else- dence,one could say that the common (Hervieu et al., tg94), while for still un- where (Schumannand Nagl, 1995).The bean evolved as P. vulgarisin an arealo- known reasonsit did migrate alone into nucleotide sequencesresponsible for'I' catedbetween Guatemala and Northern the Andes.Worth mentioning here is the phaseolincan thus be consideredas primi- Peru.Because speciation is an evolution- presenceof certain peptidesin seedstor- tive or ancestralin comparison to other ary process,it doesnot automaticallytake ageproteins of common bean that could onesin common bean.Second, the two- place in a single locality or geographic comefrom P coccineus(Lioi and Hamme6 'I' dimension structure of phaseolin is spot. The hypothesisthat the speciesP 1989),a speciesthat in its very early evo- closeto the one of a globulin presentin vulgarisdid not havea static distribution lution was closeto P. vulgaris(Hervieu et wild P.polyanthusGreenman (Schmit and at the moment of its formation,but rather al, t994), indicating that some gene ex- Debouck, r99r). CpDNA analysis has moved back and forthwards in an area changethrough natural outcrossingin- shown a close similarity betweenwild P between what is today Guatemala and deed took place.It is important to note 'I' vulgarisof Ecuador (with phaseolin) Northern Peru,can be supportedby the lhat P. coccineusand P. polyanthus wete and wild P.polyanthus of Central Guate- presenceof common phaseolintypes such probably introduced by Man into the mala astheywould belongto the samelin- as'CH' in CentralAmerica and Colombia Andes, yet as very primitive materials, eage(Schmit etal.,9y). One shouldnote (Toro Ch. et al.,r99o), or the fact that on thousands of years after the Andean that another species, P costaricensis the basisof allozymeevidence the Colom- branch of P. vulgaris separatedfrom the Freytag& Debouck,belonging to the same bian wild forms are closeto the Central bulk of migrating P vulgarisand started lineage(Schmit et al.,1993) and possibly Americanones (Koenig and Gepts,1989). colonizingthe Andes(and alreadyin com- involved in the parentage of the While nucleotidesequences responsible plete isolation from the Chimborazo- syngameonincluding P. vulgaris,P. poly- for phaseolinsynthesis were conserved, Cajamarcarange). anthusand P. coccineusL., exists in moun- other sequencescontinued to evolve,re- tainousranges of easternCosta Rica and sulting in basepair changesand substitu- 3. Common bean domestication in a westernPanamá (Freytag and Debouck, tions as revealed by AFLPs analysis Colombian perspective r996);phaseolin genes ofthis specieshave (ChacónS. et al.,1996). Finall¡ migration We don't know when the common not been sequencedso far. towardshigher latitudesclose to the Tiop- bean has been domesticated,neither the From the above,it seemsthat the wild ics parallelsand slightly beyond would be exact place, nor the identity of its beanswith'I'phaseolin representan an- a more recentphenomenon. domesticators,nor the reasonsbehind it. cestralbranch of the speciesP. vulgaris Someevidence has been however accumu- prior to its separationinto two major gene 2.7.A higher diversity in Mesoamerica? lated in recentyears which showsthat: pools.One could thus assumethe exist- A slightlyhigher diversityhasbeen long i.Bean domestication is likely ancient, (Kaplan, enceof a nuclear material that was grow- observedin wild beans of Mexico and even if a novel dating method ing in one area from which two major Central America in comparison to the 1994)has recently pleaded for a revisionof branchesprogressively separated, special- Central-SouthernAndes, on the basisof earlyassumptions (see Kaplan and Kaplan, ized and migrated, likely hundreds of seedprotein markers(Gepts et al., 1986; 1988;McClung de Thpia,r99z; McNeish, thousandsof yearsago. What can we in- Ishimotoef al.,l-9g1;ToroCh. et al.,r99o), r99z;Pearsall, :'992, for reviews). fer about that area?The present geo- or RFLP markers on different DNAs ii.Beandomestication surely happened 'I' graphic location of thesebeans with (Becerra Velásquez and Gepts, 1994; in different locations of the range of the phaseolinis somewhatptzzlíng, sinceit Khairallah et al., t99z; Sonnanteet al., wild common bean, independently in falls outside the regular range of wild 1994),and hasbeen long claimedas evi- Mesoamericaand in the Andes (Gepts, beanson the easternslope ofthe Andesin dencesupporting a singleCentral Ameri- 1993;Gepts and Debouck,t99r; Geptset South America. It is not sure that they can (or Mexican) origin for the P vulgaris al.,1986). Further, the existenceofraces in have always been in that part, from species.TWo different featurescould have common bean (BecerraVelásquez and Chimborazo to southwesternCajamarca, actedtogether and explain that higher di- Gepts,1994; Singh et al.,r99ra) would sug- which might havefunctioned asrefugia at versity in Mesoamericanwild beans:first, gestthe possibility of at leasttwo domes- the moment of migrationsof flora during under relatively less changing though tication eventsin Mesoamerica,and two the late Pliocene - early Pleistocene favourableecological conditions, the wild in the Central-SouthernAndes. (Haffer, t987; van der Hammen, r99z). It beanscolonize in Central America and iii.Bean domestication might have is worth mentioning here that another Mexico an expandedniche along a longi- taken placeat different moments in time, phaseolintype called'Mu'which displays tude gradient that is not so wide in the perhapsrepeatedly in some parts,while a simplestructure in SDS-PAGEelectro- Andes(Debouck, 1986). So, the longitude unique in time in others.This meansthat phoresisexists in Cundinamarca,Colom- gradient in Central America and Mexico certain groups of beansmight be more bia (Gutiérrez et al., 1994). So, a few would have been of the masnitude of ancientthan others.For Mesoamericaex-

REVISTACORPOICA . VOt | . Nol . OCTUBRE1996 ColombianCommon and Lima Beans | |

tending to the SouthwesternUSA, for The reasonsfor suchan in masseintro- in Colombia is possible (Toro Ch. et al., which more data areavailable, a south-to- duction of Central American and Andean 1993),but requires additional surveys.It north gradient showsclearly that beans bean cultivarsin Colombia are still un- occupiesniches at slightly higher altitudes were used much earlier in South-Central clear,and perhapsare related to the peo- and more xeric habitatsin comparisonto Mexico in comparison to other places pling of Colombia. In this regard, one the tropical'lowland' wild lima bean. (Carter,1945; Kaplan, t956, t967, t985). should note that most of the materialsof The presenceofa hydrocyanicacid pre- Other data (Kaplanand McNeish,196o) as CentralAmerican origin concentratein cursor in mature dry seedsespecially high well asthe unfinite nature of the archaeo- the North and in the Cauca and in wild forms (Baudoin et al., r99r) Ieaves logical argument might howeverinvite us Magdalenavalleys below r,zoo masl,while little possibility for their active transpor- to considerthis point carefully. thoseof Central-SouthernAndean origin tation by humans or animals. Seedsof iv.Bean domestication might have are grown in the South and generally wild forms, either grey mottled or solid taken placefor other reasonsthan con- abover,4oo masl (Debouck et al., 1993; black, as in most Phaseolusspecies, are sumptionas a boiled food item.Purposes Geptsand Bliss,1986). Materials with'B' unconspicuousand likely not noticed by diverseenough as consumption as snap or'CH' or'Ii phaseolindo not seemto rodentsor birds. Thesefeatures would al- bean(Debouck, 1989) or astoasted beans cumulate negativecharacters that would low to consider the present range of dis- (Tohme et al., 1995; Toro Ch. and havebeen culled out during selection.At tribution asresulting rather from natural Debouck,1995) or asplay (Debouck, 1989) this stage,given the lack of data, these mechanismsof seeddispersal with brutal have been proposed. Given the fact that materials might equally representan old pod dehiscence,hard seedcoat, etc, than 'center' beanshave appeared as fully domesticated of domestication,later on domi- from the intervention of human factors. in severalpre-ceramic contexts especially nated by bean introductions from other in the Andes (Engel,1987; Lynch et al., parts,or a recentone. Not surprisingl¡ the 4.2.Two gene pools, probably ancient 1985;McNeish,DTT), it is not surethat race'Nueva Granada'defined elsewhere The wild lima bean distributed in the consumptionas a boiled food item wasthe (Singhet al.,r99rb; Singh et al.,r99rc) still lowland Neotropics displays,on the basis prime reason for domesticatingbeans. lacksclear-cut boundaries, as it would be of resultsavailable so far in polymorphism Such a way of cooking beansmight have made of severalgenetical bean entities of seedlectins and arcelins,a distinct and come later,once was well estab- with different evolutionaryhistories. Since more important diversityin comparison lished, perhaps becauseof other food sampling of truly traditional is to the Andean wild lima bean (Debouck plants. still fragmentar¡ and might thus intro- et al.,r989b; Gutiérrez Salgado etal.,r995; Datafrom differentdisciplines are criti- duce a dangerousbias, these results are Maquet et al.,r99o). These results would cally lacking in the caseof Colombia. Ar- very preliminar¡ and more material of allow to concludeto the existenceof two chaeological data are few (Kaplan and Colombia,wild and cultivated,needs to be genepools in wild lima bean. The exist- Smith,1985), and not conclusive,but from studied,also paying particular attention to ence of such gene pools in the wild was historical records(Patiño, 1964), we know materials from Ecuador and Venezuela further supported by allozyme analysis that common beans are ancient in that (from where little has been thoroughly (Maquetet a1.,994)and RFLPanalysis of country. For reasonsexplained above,it studied, particularly for cultivated mate- rRNA genes(Jacob et aL,ry95). Not sur- seemsthat at least a few wild common rials). prisingl¡ giventhe extentofits rangeover beanspresent in North-EasternColombia 7,ooo Km, the lowland Neotropical wild arenot weedyescapes, but true wild forms. 4. Featuresabout wild lima bean lima beanshows a wider geneticvariation, The comparison of Colombian landraces before Humans particularly in Mesoamerica(Gutiérrez with sympatricwild forms with the useof 4.t. Twofamilies of wild lima beans Salgadoet al.,1995; Maquet et al.,r994),in the phaseolinmarker has shown the mas- Wild lima beanswith small triangular, spite of insufficient sampling. siveintroduction, likely in pre-Columbian parchment-like, almost glabrous leaflets, times, of cultivars from Central America small ,small with greenish 43. Veryearly evolution and from the Central-SouthernAndes, and hairy standardand purplish wings,are Differencesat the morphological level with the exceptionof a few landracesap- widespreadin the Neotropics,generally betweenthe two wild forms of lima bean parently domesticated in Boyacá- below r,6oo masl to sealevel (Debouck are strongerin comparison to thosesepa- Cundinamarca(Gepts and Bliss, 1986). and Smartt,rggt). The rangeof distribu- rating genepools in wild common beanif Thesefindings were confirmed with the tion of this form extendsfrom Sinaloain we assumeequal speeds of differentiation. screeningof more landraceswith the useof Mexico to Panamá,and from Thmaulipas However we presently do not have evi- AFLPs(Chacón S. et al.,1996). Such analy- in Mexico through the Caribbean up to dencethat thesetwo families of wild lima sesshow further that some Colombian coastal Colombia: elsewhere in South beanwould representtwo different evolu- 'S' 'CH' landraceswith or phaseolinof sus- America,it is known from Bahia,Braztl,and tionary pathwaysin American Phaseolinae pectedCentral American origin group to- easternPeru, and extendsup to Saltain Ar- which by morphological convergence gether with Colombian wild forms gentina(Gutiérrez Salgado et al.,1995). would constitute an artificial spe- (ChacónS. et al.,r996), indicating either In contrast,a group of wild lima bean cies.Rathe6 present evidence could be in- their true origin in Colombia or a re- with larger somewhat hairy leaflets, terpreted as if separation of the two ticulate origin with extensive wide- slightly largerseeds, deep purple standard families of wild lima bean took placeear- crossing.The presenceof'L'phaseolin in and wings, has been disclosedin inter lier in comparisonto what occurred in both wild and cultivated forms of Co- Andeanvalleys on the westernslope of the common bean.From a nuclear stock,two lombia is an additional indication of a Andes, from Imbabura in Ecuador to branches separated,one colonizing the separatedomestication event (Chacón S. SouthwesternCajamarca in Peru (Debou- Neotropical savannahs(of both Central et a1.,1996). ck et al.,rpSz). The presenceofthis form America and South America), while the

REVISTACORPOICA. VOt r. Nor. OCTUBREr996

L l2 ColombianCommon and Lima Beans

caseof lima bean.There are other was restricted to the montane dry particular distribution according to alti- Andes in the preliminary that forestsof the NorthwesternAndes. Present tude, the small-seededmaterials being indications although took place in Colombia or evidencealthough scarcewould locatethis majorily distributed on the Atlantic coast theseevents Both P. vulgaris and P. lunatus nuclear stock in an area extending from of Colombia and in inter-Andeanvalleys closeto it. wild forms in the other re- CostaRica to Ecuador.Indeed wild mate- below r,zoomasl, and the large-seededcul- expandedas to most Phaseolus sPecies, rials showingboth setsof alleleshave been tivarsbeing distributedin the Souü and in gion, in contrast of man-made alterations disclosedin that area (on allozymeevi- the cordillerasabove r,6oo masl (Gutiérrez in part because habitats, that in turn they were dence:Maquet et al., 1994),but it is diffr- Salgadoet al.,1995). As an examplethat the in their (asa preambleto their do- cult at this stage to assert that they do two genepools still belongto the samebio- ableto exploit Hawkes,1969; Heiser, 1969), representthe nuclear stock (that would logical species,and thereforedo not merit mestication: aseolusspecieswerenot. The haveevolved since anyway) or are instead a specialLatin nomenclature(Debouck, while most Ph Andesstill harbour the two the resultof wide hybridizations,although t99t), a few materials with hybrid pattern ñorthwestern both speciesor better say AndeanBig Limabeansare unknown in ar- existin Colombiaas a resultof pasthybrid- branchesfor so that it would be pos- chaeologicaltimes north of Panamá izations(Gutiérrez Salgado et al.'1995). The their descendants, earlyformation of (GutiérrezSalgado et aI.,r995;Kaplan and questionof the specificorigin for the Co- sibleto understandthe pools in the wild. IQplan,r988).It seems that: i) a truenuclear lombian cultivars is still quite open, since the gene parallelism finally extendsto the stock has not been formally identified in wild materialsof both genepools may ex- The histories, where different wild lima beans,and ii) in contrastto what ist in that countr¡ opening the possibility domestication growing in different ecological happenedwith the Andean branch of P of separatedomestications, although sepa- wild forms domesticatedseparately result- vulgaris,P. Iunatushas migrated or wasin rate introductions might be a more likely zoneswere (Debouck, r99z). One the Andeswith companionspecies such as scenariofor historic reasons(Patiño, r964). ing in vicariance is found in the range extending P augustiHarms, P.p achyrrhizoidesHarms More analysisis required,in relationto the example to Cajamarcawhere the (if theset'rvo are different!), and P. mollis germplasmof Panamá,Venezuela and Ec- from Chimborazo lima bean was probably do- Hooker whose range restricts to the uador,but first of all a better samplingof large-seeded while the wild common bean GalapagosIslands (Wiggins and Porter, the wild forms in Colombia. mesticated, phaseolingrowing in a comple- r97r). The little differencebetwe en P.augusti with'I' habitat in the samerange was and P.pachyrrhizoides might indicate that 6. Colombia:at the crossroads mentary explanation may lie in the non- speciationprocess still goeson. Finally,one tor sure not. The early farmersto domesticate has to note the extendedpossibilities for Colombia is a placeof particular rel- necessityfor (that were not neces- widecrossingwith lima bean,wiü species evancein the studyof the two most im- two wild American origin, since sarily sympatric) at the same time for such as P.ialiscanus Piper' P maculatus portant pulsesof (or similarpurposes. On the other hand,do- Scheele,P. ritensis Jones, P salicifoliusPiper it might havebeen the scene very close processesof common and (Katangaand Baudoin,r99o), all today dis- to it) of: i) the processesleading to the for- mestication and within each cultigen al- tributed in Mexico and the Southwestern mation of both speciesas such from a bulk lima beans independent were conducted USA (Debouck,r99r). Suchpossibilities let of companion species,ii) the separationof though the sameprinciples (e.g. soon us to consider genetic afñnities between their original stocksinto Mesoamerican much along with and squashes' thesespecies and the nuclearstock leading and Andean branches(all wild) asfurther growing beans beanswith ceramics),leading to P.Iunatus. stepsof the speciationprocess' and iii) or cooking (detailed separatedomestication events or selection to the sameevolutionarychanges in cultivarsdomesticated by Smartt,1988) and consequences (e.g. an 5. Gene pools in cultivated of recombinants reduction of diversity upon lima bean elsewhere.Evidence is coming that Co- important Yet,markers have revealed Probably as the result of independent lombia is more than a simple zoneof con- domestication). amounts of diversity among domesticationsin the two different fami- tact,because of its geographicalposition' different within eachcultigen (fot P. vul- lies of wild lima bean, there are tlvo gene betweenthe major genepools for these groups and Gepts,:-994; pools in cultivated lima beans,as evi- two pulses. garisi BecerraVelásquez and Tohme, 1989,and for P. dencedon seedproteins (Debouck et al., There is someparallelism between the Debouck Gutiérrez Salgadoet aI', t995; r989b; Gutiérrez Salgado ef al, t995; situationsprevailing in the wild common Iunatus; although Nienhuis et al., 1995),indicating that on Maquet et al., 99o),allozymes (Maquet et bean and the wild lima bean, these grouPs are not at the aI., t994), RAPDs markers on total ge- much evidenceis still lacking,particularly marker base stage,and so opening nomic DNA (Nienhuis et al., 1995)and for the latter. But there are alsomajor dif- sameevolutionary to better use the original poten- RFLPsofrRNA genes(|acob et a1.,t995). ferencesin such a parallelism,namely in the way American Phaseolinae. We lack evidenceas to ascertainthat there the timing of the events,that maintains tial of these countries where these were more than one domestication event thesetwo taxa well apart in the evolution All Neotropical are presentdo all harbour ge- within eachfamily of wild forms,but such of the genus(Debouck, r99r; Maréchal et wild beans some of which havebeen possibilitymight existat leastin the Andes al., ry78). Both specieswere formed by netic resources, past evolution of thesespe- (GutiérrezSalgado et al.,1995). This needs separationfrom a bulk of companion spe- critical in the wild forms or ascultigens' As per- additional support sincethe presenceof cies,which today representthe secondary cies,as these Neotropical countries are high cyanogenicglucoside in the wild and tertiary genepools (Debouck'r99r) haps all in enhancing their agrobio- would rather be a limiting factor for inde- and possibilities for promiseful wide- interested priority is to collect,document pendentmultiple domestications. crossing.These companion speciesmi- diversit¡ a further these resourcesbefore The two genepools of cultivated ma- grated to Central America and Mexico in and study terials are present in Colombia, with a the caseof common bean, and to the their extinction.

REVISTACORPOICA . V0[ | ' Nol ' ocTUBRE1996 ColombianCommon and Lima Beans l¡

ACKNOWLED GEMENTS Gepts,P. (ed.), I(uwer AcademicPublishers, Debouck, D. G., Maquet, A. and Posso,C. Dordrecht, Holland, pp. 185-214. E. r989b. Biochemicalevidence for two This essayincludes parts of the conference Burkart,A. r94r. Sobrela existenciade different genepools in lima beans,Phaseolus presentedby the author in La Selva,Medellín, razassilvestres de Phaseolusvulgarisy lunatusL. Annu. Rept.Bean Improvement Colombia on February 6th 1996.The author Phaseoluslunatus. Resoluc.& Resumenes Coop.32:58-59. thanks the Centro Internacional de Botánica,Primera Reunión Argentina de Debouck, D. G. and Smartt, f. 1995. Agricultura Tlopical, the International Board Agronomía, BuenosAires: 52. Beans,Phaseolusspp. for Plant Genetic Resources(now IPGRI), the Burkart, A. and Brücher, H. 1953. (Leguminosae-Papilionoideae).In: "Evolution World ConservationUnion and the USDA Phaseolusab origineus Brtkart, die of crop plants.Second Edition', for generousgrants for field work on several mutma3liche andine Stammform der Smartt, j. and Simmonds,N.W. (eds.), occasions.The author would like to express Kulturbohne. Zichter 23 (3): 65-7 2. Longman Scientific& Technical,London, '1976. specialthanks to programmesof genetic Cabrera,A.L, Regiones United Kingdom, pp. 287-29 4. resourcesand agricultural researchservices of fitogeográficasargentinas. Editorial ACME Debouck, D. G, and Tohme, I, tg8g. Mexico, Guatemala,Costa Rica, Colombia, S.A.C.I.,Buenos Aires, RepublicaArgentina, Implications for bean breedersof studieson Ecuador,Peru, Bolivia, Chile and Argentina 8sp. the origins of common beans,Phaseolus "Current for technicalsupport during field CañadasC., L. and EstradaA.,W. 1978. tulgarisL. Ír: topics in breeding of explorations.Thanks are due to Sandra Ecuador.Mapa ecológico,Escala l: 1.000.000. common bean",Beebe, S. (ed.), Bean Albarracín for the final ryping ofthe Ministerio de Agricultura y Ganaderia, Program, Centro Internacional de manuscript. Pronareg,Acuerdo MAG-ORSTOM, Agricultura Tiopical, Cali, Colombia, pp. Instituto GeográficoMilitar, Quito, Ecuador, 3-42. one sheet. Debouck, D. G., Toro, O., Paredes,O. M.,

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