1492 and the Loss of Amazonian Crop Genetic Resources. II. Crop Biogeography at ContactAuthor(s): Charles R. ClementSource...
1492 ANDTHELoss OF AMAZONIANCROP GENETICRESOURCES. II. CROP BIOGEOGRAPHYAT CONTACT"2CHARLES R. CLEMENTClement, C.R. (Insti...
204 ECONOMICBOTANY [VOL.531989). This area appearsto have relied moreheavily on fruit crops than was the case else-where i...
1999] CLEMENT:AMAZONIANCROPRESOURCESII 205FJLowlondliIAndeIlaCoostalDesert305 30SOXTemperatePraireUScrub0 00** 160W_______...
206 ECONOMICBOTANY [VOL.53species wereincludedin this centerof diversity.Note also that position of the Brazilianfocuswas ...
1999] CLEMENT:AMAZONIANCROPRESOURCESII 20711.BrazilPeru30S j*30510.ChileOS 160W 60WW6iWFig. 2. Post-Vaviloviandevelopmento...
208 ECONOMICBOTANY [VOL.53tween wild and domesticatedfruit species, be-cause his majorobjective was to identify areasfor t...
1999] CLEMENT:AMAZONIANCROPRESOURCESI 209trade,stealthor duringwarfare(Leon 1987),as well as by domestication;e) the longe...
210 ECONOMICBOTANY [VOL.5360W ,60W~~~~~CentersMinorCentersRegions0 0~~~~60 WFig. 3. Biogeographyof Amazoniaandnorthernlowl...
1999] CLEMENT:AMAZONIANCROPRESOURCESII 211high populationdensities at contact, especiallyalongthevarzeas(Denevan1992)andth...
212 ECONOMIC BOTANY [VOL. 53Xanthosoma sagittifolium, Crescentia cujete,possibly Carica papaya, Cucurbita moschata,Dioscor...
1999] CLEMENT:AMAZONIANCROPRESOURCESII 213CentersMinorCenters4 **. (7)~~~~~~Regions0~~~~~~1030S 30SFig. 4. The biogeograph...
214 ECONOMICBOTANY [VOL.53riculture. Giacomettis (1992) suggestion forthreecentersis transformedinto a single regionof div...
1999] CLEMENT:AMAZONIANCROPRESOURCESII 215Lathrap, D. W. 1970. The upperAmazon. Praeger,New York.. 1977. Ourfatherthe caym...
216 ECONOMICBOTANY [VOL.53of cultivatedplants. CambridgeUniversityPress,Cambridge,UK.. 1992e.The problemconcerningthe orig...
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1492 and the loss of amazonian crop genetic resources. ii. crop biogeography at contact

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1492 and the loss of amazonian crop genetic resources. ii. crop biogeography at contact

  1. 1. 1492 and the Loss of Amazonian Crop Genetic Resources. II. Crop Biogeography at ContactAuthor(s): Charles R. ClementSource: Economic Botany, Vol. 53, No. 2 (Apr. - Jun., 1999), pp. 203-216Published by: Springer on behalf of New York Botanical Garden PressStable URL: http://www.jstor.org/stable/4256180 .Accessed: 11/05/2011 20:56Your use of the JSTOR archive indicates your acceptance of JSTORs Terms and Conditions of Use, available at .http://www.jstor.org/page/info/about/policies/terms.jsp. JSTORs Terms and Conditions of Use provides, in part, that unlessyou have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and youmay use content in the JSTOR archive only for your personal, non-commercial use.Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at .http://www.jstor.org/action/showPublisher?publisherCode=nybg. .Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printedpage of such transmission.JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range ofcontent in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new formsof scholarship. For more information about JSTOR, please contact support@jstor.org.New York Botanical Garden Press and Springer are collaborating with JSTOR to digitize, preserve and extendaccess to Economic Botany.http://www.jstor.org
  2. 2. 1492 ANDTHELoss OF AMAZONIANCROP GENETICRESOURCES. II. CROP BIOGEOGRAPHYAT CONTACT"2CHARLES R. CLEMENTClement, C.R. (InstitutoNacional de Pesquisas da Amaz6nia,Cx. Postal 478, 69011-970Manaus,AM,Brasil). 1492 AND THELoss OF AMAZONIANCROP GENETICRESOURCES.II. CROPBIOGEOGRAPHYAT CONTACT.EconomicBotany53(2):203-216, 1999. Fiftysevenpercentof the138 cultivatedplantspeciespresentinAmazoniaat contactprobablyoriginatedin theAmazonBasin and another27% originatedin lowlandnorthernSouthAmerica.The relationshipbe-tweenprobableindigenoushumanpopulationdensityand resultantagriculturalintensificationand crop diversityis used to propose the existence of a mosaic of crop genetic resourceconcentrationsinAmazoniaat contact,includingtwo centersof diversity,four outlyingminorcenters,andfive regionsof diversity.Thismethodologyis extrapolatedtopresenta synthesisof SouthAmericancropgeneticbiogeographyat contact.1492E A PERDA DOSRECURSOSGENETICOSDA AMAZ6NIA. II.A BIOGEOGRAFIADOSCULTIVOSNOMOMENTODE CONTATO.Cinquentae sete por cento das 138 especies cultivadasna Amaz6niano momentodo contatoprovavelmenteoriginaramna baciaamaz6nicae mais27%originaramnas terras baixas do norte de Americado Sul. A relaV-doentre a densidadepopulacionalindigenahipoteticae a intensifica,cioagricolae a diversidadegeneticados cultivosresultanteseusadapara propora existenciade ummosaicode concentra,ces de recursosgeneticosnaAmazoniano momentode contato,incluindodois centrosde diversidade,quatrocentrosper-ifericos menores,e cinco regioes de diversidade.A mesmametodologiaeextrapoladaparaapresentarumasintese da biogeografiada diversidadegenetica dos cultivosna AmericadoSul no momentode contato.Key Words: Amazonia;centersof diversity;regionsof diversity;SouthAmerica.Seventy-nine (57%) of the 138 crops culti-vated or managed in lowland northern SouthAmerica at contact probably originated in theAmazon Basin and another 37 crops (27%) inadjacent parts of lowland South America (Clem-ent 1999). This represents 45% of the total forthe Americas and has been largely ignored bymost students of Neotropical crop diversity. Inthis paper, an hypothesis of Amazonian cropbiogeography at contact is presented to placeAmazonian diversity squarely in an integratedSouth American context.The centers of crop genetic diversity proposedby Vavilov (1992c) and his successors are oftenrelated to the presence of advanced societies. In1Received 12 February1998;accepted8 December1998.2Dedicatedto VictorManuelPatinlo,one of LatinAmericasforemosteconomic andethnobotanistsandfellow studentof croporigins,domesticationandbio-geography.SouthAmerica,the prehistoryof manyof thesesocietiesandpossiblecentersof cropgeneticdi-versity associatedwith them are being discov-ered,althoughthey arestill farfromcompletelyknown (Pearsall1992). In Amazonia,however,little archeologicalwork with modem methodshas been done (Meggers 1988; Pearsall 1992).Becauseof the lack of architecturalartifacts,therapidpost-contactpopulationdecline and smallrecuperation,the vigor with whichthe forestre-colonizes areas that humansabandon,and thecatastrophicloss of cropgeneticresourcesin thelast 500 years(see Clement1999), it is difficultto identifyconcentrationsof cropgeneticdiver-sity in Amazoniatoday. These difficultiesalsohelp explain why Amazonia has been over-lookedas a cradleof cropdiversityby everyoneexceptSauer(1952), whorecognizedthatit con-taineddiversitybut did not proceedto identifyit as a majorregion of diversity.A center of crop genetic diversity in NWAmazonia was proposed recently (ClementEconomic Botany 53(2) pp. 203-216. 1999C) 1999 by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.
  3. 3. 204 ECONOMICBOTANY [VOL.531989). This area appearsto have relied moreheavily on fruit crops than was the case else-where in the Americasat contact(Patinio1963:21-52). The pejibaye (Bactris gasipaes), for ex-ample, may have been a staple crop in this re-gion [this inference is based on its degree ofdomestication (Clement 1988) and its impor-tanceto manysocieties as seen in theirlegends,myths, festivities and calenders(Patinio1992)],supplemented by cassava (Manihot esculenta)andmaize (Zea mays).Annualcropsarethe fa-miliarpatternin the temperatezone andthe bi-ases of early and currentbiogeographers,an-thropologistsand archeologiststoward the fa-miliaris probablyyet anotherreasonthatAma-zonia has been ignored.Why is the NW Amazoniancentervisible to-day and others are not? Possible explanationsinclude the fact that the northwestdivides theareas of Spanish and Portuguese authorityinAmazoniaand was one of the most difficulttoaccess during the early post-contactcenturies.Consequently,althoughalso severely impactedby disease, missionization,andlaterby the rub-ber boom, many of the indigenouspeoples sur-vived and preservedtheircrop genetic heritagebetterthanelsewherein Amazonia.The specialcharacteristicsof NW Amazoniaarenot widelyreplicatedin otherareas.Therefore,othermeth-ods, based on hypothesesaboutearly historicaldemography,currentideas of societal organiza-tion as a functionof demography,carryingca-pacity of the various Amazonian ecosystems,andcurrentknowledgeof cropdiversityanddis-tribution,must provide clues to the biogeogra-phy of Amazoniancropsat contact.In this paperI review earlyhypothesesaboutSouth American crop biogeography, suggesthow indirect evidence can be marshalledtoidentify possible concentrationsof crop geneticresources at contact, present a hypothesis ofAmazoniancrop biogeographyat contact, andextrapolatethe methodologyto provide a syn-thesis for SouthAmericaas a whole. These hy-potheses are all testableby archeologicalmeth-ods.HYPOTHESESON SOUTHAMERICANCROP BIOGEOGRAPHYIn this shortreview,I concentrateexclusivelyon SouthAmerica.All of the authorsmentioned,except Clement (1989) and Giacometti(1992),presentedworld-wideanalyses and hypotheses,andall were also interestedin the originsof ag-riculture.As far as possible, I avoid their dis-cussion of the lattersubjectandconcentrateex-clusively on the biogeographyof crop geneticdiversity.The first major student of world-wide cropbiogeographywas Alphonsede Candolle(1882).The informationavailableon cropbiogeographyat thattime was limitedandtaxonomicclassifi-cation of crops was still relativelyconfusedbyan overabundanceof synonymous names inmanycultivatedtaxa.The geneticsof biologicalspecieswere also unknownthen,so thelimitsofsome species groupswere unclear.De Candolleidentified South America as the origin of nu-merouscropsandin some cases identifiedprob-able sub-continentalregions of origin, althoughhe emphasizedthathis informationmightbe un-reliablefor suchfine definitions.His majorcon-tributionwas to apply a multi-disciplinaryap-proach,that is still applied today in expandedandmodifiedform.Forty years later,Nicolai I. Vavilov and hiscolleagues startedexploringthe mountainsandadjacentlowlands of the Neotropicsand incor-poratingthese regions into their hypothesesofworld-widecrop biogeographyand agriculturalorigins.The recenttranslationof Vavilovscon-tributionsto crop biogeographyprovidesa fas-cinatingrecordof the developmentof these hy-potheses.Figure lb-f shows theirdevelopmentwith respectto SouthAmerica.Figure la is anapproximationof the physical geography ofSouthAmerica.The firsthypothesiswas essentiallygeneratedfrom the literature,accordingto Harlan(1992:49). A region of diversity extended along theAndean mountainsand CentralAmerica, fromMexico to Chile. In SouthAmericait followedthe Andes and immediatelyadjacentcoast andinteriorlowlands(Figurelb), fromVenezuelatoChiloe Island, Chile (Vavilov 1992a). Vavilovalways assumed a close relationshipbetweencrop genetic diversity and centers of advancedcivilizations (Vavilov 1992b), which explainslargesectionsof thishypothesis.InSouthAmer-ica he emphasizedthe Incas,the Chibcha(in theColombianmid-to high-elevations),andtheAr-aucanos(in SouthernChile).As informationcollected by Soviet scientistsbecame available,Vavilov generatedmore pre-cise hypotheses.In 1929, he (1992e) hypothe-sizedthatthereweresix centersof cropdiversity
  4. 4. 1999] CLEMENT:AMAZONIANCROPRESOURCESII 205FJLowlondliIAndeIlaCoostalDesert305 30SOXTemperatePraireUScrub0 00** 160W____________ _________________Fig. 1. The developmentof Vavilovshypotheseson the biogeographyof crop genetic diversityin SouthAmerica.A. Physicallvegetationalgeography.B. Thefirsthypothesis(Vaviloy1992a),withone centerof origin.C. The secondhypothesis(Vavilov 1992e), with six centersof origin,each with a focus of type formation.D.The thirdhypothesis(Vavilov 1992d),with one centerof originandfourfoci of type formation.E. Thefourthhypothesis(Vavilov 1992c), with one centerof originand threefoci of type formation(the map in the 1992versiondoes not include VIIIB,but the text does). F.The finalhypothesis(Vav7ilov1992f), with a threepartcenterof origin.in SouthAmerica,each containinga "focus oftypeformation."By this,Vavilovmeanttheareain which a domesticatedcrop shows most clear-ly thetraitssubjectto selection.Thesefoci were:the Colombianhighlands;the Peruviancoast;the mid-elevationAndes;the Peruvian-Bolivianhigh-elevationAndes;theareaaroundChiloeIs-land,Chile;andpartof theParana-Paraguayriv-er basinsin southernBrazil (Figure ic).In 1931 the hypothesis was modified again(Vavilov 1992d), drawingall of the Peruvian-Bolivian foci into one centerof diversitywith ahighlandfocus, while maintainingthe Colom-bian, Chiloean and Brazilianfoci (Figure Id).The latterthreefoci were no longerpartsof in-dependentcentersof diversity,however.The en-larged center of diversity covered parts of thePeruviancoast where the originalhypotheticalfocus hadbeen, as well as partsof the drainagesystems of severalupperAmazonianrivers,no-tablythe UcayaliandtheHuallaga.This appearsto have been deliberate,as several Amazonian
  5. 5. 206 ECONOMICBOTANY [VOL.53species wereincludedin this centerof diversity.Note also that position of the Brazilianfocuswas shiftedslightly.In 1935, Vavilov(1992c) publishedhis majorsynthesis, The Phyto-Geographical Basis ofPlantBreeding,in whichhe recognizedonly onecenterof diversityin SouthAmerica,the Ecua-dor-Peru-BoliviaAndeancenter,with two asso-ciated foci, the Chiloe and the Brazil-Paraguay(Figure le). He does not explain why the Co-lombianfocus was eliminated.Forthefirsttime,crop lists are presentedfor each centerand fo-cus; several Amazoniancrops are included inthe crop list for the centerof diversity,as wellas for the Brazil-Paraguayfocus. This map iswidely published as Vavilovs definitive state-ment.Vavilovs final hypotheis was published in1940, however.The Andeancenterof diversitywas divided into three sections:the Bogotafo-cus, the Ecuador-Peru-Boliviafocus, and theChiloe focus (Figure If). In this final version,the Brazil-Paraguayfocus was eliminated,againwithoutexplanation.Unfortunately,Vavilovwasimprisonedanddied shortlythereafter.The curiousthing about the developmentofVavilovshypotheseswithrespectto SouthAmer-ica is the shiftingimportancethathe attributestothe ColombiaandBrazil/Paraguayfoci. It is un-clearwhy these shiftsoccurred.It is clear,how-ever, that he recognizedthe importanceof thetropicallowlands, even though he visited onlytheirperiphery,at Belem, Para,Brazil (Vavilov1997). The continualinclusionof Chiloe Islandand adjacentChile is a fine exampleof bias to-wardsannualcrops,whichwerethenandcontin-ue to be extremelyimportantto worldfood se-curity.VavilovthoughtthatChiloeIslandprovid-ed theSolanumtuberosumgenotypebestadaptedto Europe;otherwiseit only producedMadiasa-tiva and Bromus mango (Vavilov 1992c), bothessentiallyextinctin cultivationtoday.Shortlyafterwards,Darlington& JanakiAm-mal (1945) publisheda map that they claimedwas based on Vavilovs work (Figure2a). TheAndeancenter,however,was shiftednorthto in-cludeColombiaandexcludeBolivia.TheChiloeIsland area was maintained.The Brazil-Para-guay areawas expandedto encompassthe Bra-zilian savannas,most of Amazonia,the Guianahighlandsandmost of the OrinocoRiverbasin.No explanationwas given. In 1956, Darlington(1973) acceptedKuptsovs(1955) proposedlo-cations of the worldsprimaryregions of agri-cultureandthe regionsinto whichthey diffusedaftertheirinitial development.In this map (3rdedition, 1973), VavilovsAndeancenterequaledKuptsovsprimaryregionandthe areaof expan-sion includedall of SouthAmericanorthof thePlataRiver.In the early 1950s, Sauer (1952) suggestedthatseveralSouthAmericancropsoriginatedinAmazonia.He didnt attemptto delineatecon-centrationsof crop genetic diversity,however,and is therefore seldom cited with respect toSouthAmericancropbiogeography.A decadelater,Zhukovsky(1968, 1975) pro-posed that crop genetic diversitywas so wide-spreadin most of the world that megacenterswere a more appropriateapproximationof cropbiogeography.In South America, Zhukovskysmegacenter 10 (Figure 2b) was essentiallyequivalentto Vavilovsfirsthypothesis(1992a),with anextensionto includenorthernArgentina.At the same time, Brucher(1969) criticizedVavilovshypothesison thebasisthatmanycropsdo not have centersof diversitythatcorrespondwithVavilovianbiogeography.Infact,forBriicher,centersof diversitydo notexist.Rather,eachcropdisplaysone or severalcentersof originand anarcof diffusion(Brucher1990).Immediately thereafter,Harlan (1971) pro-posed that some continents have centers andsome dont. SouthAmericawas an exampleofa non-center(Figure2c), whereagriculturewasso widely distributedthat no clear centerscanbe identified.For the firsttime the northeasterncoast of South America,from Recife, Pernam-buco, Brazil, across the mouthof the AmazonRiverandalongtheGuianacoastandhighlands,was includedin SouthAmericanbiogeography.Why was not clear.Chiloe Islandwas excludedfor the firsttime, also withoutexplanation.Hissubsequentbooks (Harlan1975, 1992) failed toclarifythese issues.A decade later,Hawkes (1983) attemptedtosynthesizeVavilovs,Zhukovskysand Harlansideasinto a mosaic.In SouthAmerica,Hawkesscheme(1983:70-73) positeda nuclearcenterofagriculturaloriginin theAndes(essentiallyVav-ilovs (1992c) Ecuador-Peru-BoliviaAndeancenter),surroundedby a regionof diversityintowhich agriculturediffused [essentiallyZhukov-skysmegacenter(ZevenandZhukovsky1975)],and a couple of outlying minorcenters(essen-tially Vavilovs Chiloean and Brazil/Paraguay
  6. 6. 1999] CLEMENT:AMAZONIANCROPRESOURCESII 20711.BrazilPeru30S j*30510.ChileOS 160W 60WW6iWFig. 2. Post-Vaviloviandevelopmentof SouthAmericanbiogeographichypotheses.A. Darlington& JanakiAmmals(1945) biogeography.B. Zhukovskys(1968) SouthAmericanmegacenter.C. Harlans(1971) SouthAmericannon-center.D. Clements(1989) NorthwesternAmazoniancenter.E. Giacomettis(1992) centersoffruitbiodiversityin Brazil.foci). No mapwas presented.This synthesisap-pearedto offer greatpromisefor furtherdevel-opment, but was not elaborated further byHawkesor otherauthors.Duringthe 1980s, workin Amazoniaallowedme (Clement1989)to proposea Vaviloviancen-ter of crop diversityin NorthwesternAmazonia(Figure2d), basedprincipallyon fruitcrops,al-though various other crops were included assupportingevidence. At that time I also men-tioned thattherewas more crop diversitynorthof the proposedcenter,as well as elsewhereinAmazonia.In 1992,Harlan(1992) restatedhis convictionthatSouthAmericais a non-center,butacknowl-edged that a complex mosaic of crop diversitywas presentin this and other non-centers.Un-fortunatelyhe did not elaborateon the form ofthis mosaic or presenta map.Pearsall (1992) reviewed evidence from ar-chaeologyon theoriginof cropsin SouthAmer-ica. Shedividedthewesternpartof thecontinentintolow, mid-andhighelevationzones andpro-vided exhaustivelists of crops found in manylocalities and their time-depthsof first appear-ance.This database providesconsiderablecon-crete evidence for the diffusion of crop biodi-versity but is still incomplete,covering only athirdof the continent.Giacometti(1992) proposedthatthereare 10centersof fruitspecies diversityin Brazil (Fig-ure2e). Unfortunately,he didnotdistinguishbe-
  7. 7. 208 ECONOMICBOTANY [VOL.53tween wild and domesticatedfruit species, be-cause his majorobjective was to identify areasfor the in situ conservationof wild relativesoffruitcrops and species with potentialfor futuredomestication.Giacomettisspecies lists can beused to identifythose centersthatprobablyhadsignificant fruit crop genetic diversity, as op-posed to fruitbiodiversity,at contact:1-UpperRio Negro/NorthwesternAmazonia;2-LowerAmazonRiver and adjacentAtlanticcoast; 4-Solimoes River/westernAmazonia [an expan-sion of Clements(1989) proposal];9-Atlanticforest, with three subsections;10-Brazil/ Par-aguay(essentiallyVavilovs(1992c) focus of thesamename).The other5 centersarerichin fruitbiodiversityand might once have been sites ofendemicincipientdomestication,butcurrentin-formation is too scarce to confirm them. Allwere populatedto some degree and certainlycontainedsome crop genetic diversity.From this shortreview, it is clear thatPick-ersgill and Heisers(1977) observationthatthevarietyof hypothesesaboutthebiogeographyofcrop genetic resourcesin SouthAmericais dueprincipallyto a lackof informationremainstrue,even though significant advances have beenmade, especially in the Andes. It also seemsclearthatsometype of mosaicwill be necessaryto describecropbiogeographyin theregion.Thequestionis how to characterizeconcentrationsofdiversity.DEFNING A CROP DIVERSITYMOSAICSince Vavilovs proposals,a mosaic of cropgenetic diversityhas been recognizedin SouthAmerica,althoughVavilovneverusedthisterm.De Candolle, Brucher,and Harlanrecognizedthe difficultyof identifyingdiscretecenters;deCandollefor lack of firmdata,BrucherandHar-lan because modernresearchhad alreadyiden-tifiednumerouscropswithprobableoriginsout-side of Vavilovsmosaic. Hawkes(1983:71-72)was the first to specifically propose an hierar-chical structurefor this crop diversity mosaic.His hypothesis resolved part of the criticismraised by Harlanbut it ignored two thirds ofSouthAmerica.Most of the pre-1980s studentsof crop bio-geographyconsideredagriculturaloriginsas in-tegralto theirhypotheses.Generally,they werealso diffusionists,at leastinitially,believingthatonly a few peoples hadthe idea thatlead to do-mesticationandagriculturaldevelopment,whileothers simply copied. The main exception isBrucher.In the mid- to late 1980s, however,theconceptof a Neolithic revolutionlost favor,be-ing replacedby an evolutionary(Rindos 1984)or a situational(MacNeish 1992) hypothesis,orsome synthesis of these (e.g., Harlan 1992).Some form of synthesis seems most likely,which meansthatagricultureprobablyoriginat-ed in variousplaces at various times in SouthAmerica.The evolutionary aspects of this synthesissuggest that the first steps towardsagriculturewere taken during the late Pleistocene, ratherthan the early Holocene. These first steps willbe difficultto detectin the archeologicalrecordbecausetheyinvolve minorchangesin landuse,settlementbehaviorandplantpopulationgenet-ics. The suddenappearanceof "domesticated"crops in the early Holocene archaeologicalrec-ord (e.g., Manihot esculenta and Ipomoea ba-tatas in coastal Peru, Pearsall 1992) suggeststhatdomesticationstartedmuchearlier,e.g., dur-ing the late Pleistocene. Althoughthese initialstepshave not yet been identifiedarcheological-ly, the final stages in the domesticationcontin-uumremainvisible, even after500 yearsof con-questandacculturation.Consequently,I develophere a mosaic withoutreferenceto agriculturalorigins, placing the emphasis instead on otherculturalandbiotic factors.Specifically,I hypothesizethat:a) an environmentrich in biodiversityis likelyto yield morecropsthananenvironmentpoorin biodiversity,if humansarepresentto usethe diversity(Levi-Strauss1950);b) humans continuallyexperimentwith plantsin theirenvironmentand agriculturalpeoplewill bringanythingespeciallyusefulintodo-mestication(cf. Harlan1992; Hawkes 1983;Rindos 1984), with the possible exceptionofperennialsthatareexceptionallyabundantintheirenvironment(e.g., buriti[Mauritiaflex-uosa], aqai-do-Para[Euterpeoleracea];c) the combinationof swidden/fallowagricul-ture and micro-environmentalheterogeneityin Amazonia allowed for rapid progressindomestication,because of small populationsizes (genetic drift), inbreeding, selection,short generationspans, etc (Clement 1988)(This allows for the same type of extensivevariationobservedby Vavilov and othersinmountainousregions);d) humansocieties accumulateuseful plantsby
  8. 8. 1999] CLEMENT:AMAZONIANCROPRESOURCESI 209trade,stealthor duringwarfare(Leon 1987),as well as by domestication;e) the longer humans continuouslyoccupy anarea,the more crop diversitythey are likelyto createor accumulate(Vavilov 1992a);f) humanpopulationdensitycorrelateswith thecarryingcapacity of the domesticatedland-scape,whichis determinedby theinteractionbetween the physical environmentand hu-man food procurementand processingtech-nologies, social organization,agriculturalin-tensificationand the costs of these (W. M.Denevan,pers. com., 1995);g) agriculturalintensificationdependedprinci-pally uponcrop genetic diversityto producereliably in a biotically aggressive environ-ment like Amazonia,especially on the terrafirme, but possibly less so on the varzeabe-cause periodic flooding could reduce pestpopulations;h) in any given locality, any or all of the aboveassumptionsinteractto maximizegeneticdi-versityas social organization,agriculturalin-tensificationandpopulationdensityincrease;i) the rare observationsabout crops made byearlychroniclersof ecological Amazoniacanbe deciphered(e.g., Patinlo1963, 1964) andno majorchanges in biogeographyof mostAmazoniancrops occurredin the first 100years after contact,i.e., until Europeancol-onizationstarted;j) for crops thatwere not adoptedby Europeansettlersand not lost completely,currentbio-geography can provide another reasonablyreliableguide to pastbiogeography.A majorexample is Bactris gasipaes, whose bioge-ographyhasbeenpartiallymappedin thelastdecade (Clement1988;MoraUrpi 1992).Like Hawkes (1983), I believe that a three-tiered hierarchyis useful. In the following hi-erarchyeach tieris basedexclusively on the ex-pectationof crop genetic diversity,ratherthanon agriculturalorigins and diffusions, eventhougha significantportionof the diversitymaynot be presenttoday(Clement1999).I conservethe terminologydeveloped duringthe last cen-tury,but defineeach termdifferently.The hier-archyincludesthe following levels:1) Centersof diversity-relatively restrictedar-eas with high concentrationsof crop geneticresourcesat contact, includingboth speciesandlandracesor cultivars.The high concen-trationmay be due to one or more of thefollowing factors:greattime depthof humanoccupation;high humanpopulationdensitiesand associatedsocial organizationand agri-culturalintensification;significantmicro-en-vironmentaldiversity that enhancedgeneticdiversificationduringdomestication.2) Minor centers of diversity-restricted areaswithmoderateto high concentrationsof cropgenetic resourcesat or before contact. Theconcentrationis relatedto a specificculturalgroupwith a significantagriculturaltechnol-ogy and may be thoughtof as a center ofaccumulationduring the centuries immedi-atelypre-contact.These arehypothesizedforthe areas with earthworks,some of whichwere abandonedbeforecontact.3) Regions of diversity--extensive areas withmoderateconcentrationsof crop genetic re-sourcesatcontact.The concentrationmaybedue to one or moreof the factorsmentionedfor the centersof diversitybut never actingintensivelyenough to attainhigh concentra-tions.Therewerealso areaswithlow concentrationsof cropgeneticresourcesatcontact,becauseanyareaoccupiedby humanpopulationswith someform of agricultureor landscape managementwould have crop genetic resources.Concentra-tions were low in these areasbecause of a lessfavorableenvironment,low populationdensities,simpler social organization,and/or shorteroc-cupation.An exampleis the ChiloeIslandfocusthat Vavilov and subsequent students main-tained. There were also areasthat were essen-tially uninhabitedby agriculturalpopulations,e.g., thedriersectionsof theBraziliannortheast,the Argentineanpampas,etc.BIOGEOGRAPHYOF AMAZONIANCROPGENETICRESOURCESAt contact, there were probablytwo centersof diversity,fourminorcenters,andfive ormoreregions of diversity in lowland northernSouthAmericaeastof theAndes(Fig. 3). All exhibitedmoderateor high populationdensity, advancedsocial organization,and agriculturalintensifica-tion. Many of them are not visible today, al-though some of the early chronicles attest totheir existence. I discuss each concentrationofdiversityandattemptto builda croplist foreachwith emphasis on domesticates (see Clement1999forbotanicalfamilies,probableorigin,anduses). The majorresourcefor this data set was
  9. 9. 210 ECONOMICBOTANY [VOL.5360W ,60W~~~~~CentersMinorCentersRegions0 0~~~~60 WFig. 3. Biogeographyof AmazoniaandnorthernlowlandSouthAmericain 1492. Centersof Diversity:1.NorthwesternAmazonia;2. CentralAmazonia.MinorCentersof Diversity:3. MarajoIsland;4. LlanosdeMojo;5. MiddleOrinoco;6. Guiana.Regionsof Diversity:7. AmazonEstuary;8. Solimoes;9. UpperNegro/Orinoco;10. UpperAmazon;11. GuianaCoast.Patifio(1963, 1964). I also assumethatseveralcrops were commonto all areas:Bixa orellana,Ananas comosus, Ipomoea batatas, Manihot es-culenta, Zea mays, Gossypium barbadense, Gen-ipa americana, and Nicotiana tabacum. Theselists are certainlyincomplete.Confirmationandexpansionof these croplists by paleoethnobota-nists will be a test of the soundness of theseproposals.1. The Northwestern Amazonian CenterThis center (Clement 1989) extends along theupperAmazon River from below the mouthofthe Putumayo/IIaRiverin the east to abovethemouthof the Napo Riverin the west, as well asextendingup the PutumayoRiver to the north-west and the lower JavariRiver to the south.Populationdensities at contact were probablyhigh, especially along the varzeas throughoutthiscenter.Environmentalvariationis also high,with outcroppingsof Guianashieldremanentsinthe northof the centerandextensive reworkingof the Tertiarysediments by river meanders.This centeris relativelywell preserved,princi-pally because of the large numberof Amerin-diansocieties still presentin the area,especiallyin comparisonwith the CentralAmazoniancen-ter.The crop list probablyincludedthe follow-ing domesticates: Rollinia mucosa, Xanthosomasagittifolium, Crescentia cujete, Canna edulis,Carica papaya, Cucurbita moschata, Dioscoreatrifida, Erythroxylum coca var. ipadu, Poraquei-ba sericea, Persea americana, Arachis hypo-gaea, Canavalia ensiformis, Phaseolus vulgaris,Pachyrrhizus tuberosus, Calathea allouia, Bac-tris gasipaes Putumayo landrace, Pouteria caim-ito, Capsicum chinense, Solanum sessiliflorumandothervegetables,condiments,drugsandpoi-sons. A few of the semi-domesticatesmentionedin Clement (1989) include: Quararibea cordata,Inga edulis, Pourouma cecropiaefolia, Borojoasorbilis, and possibly other semi-domesticatesand incipientdomesticates.2. The Central Amazonian Center. Thiscentermayhaveextendedalongthemainstreamof the Amazon River,from the PurusRiver inthe west to the TapajosRiver in the east, andincludedthe lower sections of the Negro, Ma-deira and TapajosRivers. Therewere probably
  10. 10. 1999] CLEMENT:AMAZONIANCROPRESOURCESII 211high populationdensities at contact, especiallyalongthevarzeas(Denevan1992)andthegreat-est time depthdetectedin Amazoniato date-11000 BP (Roosevelt et al. 1996). There arelarge areas of terrapreto do indio [an anthro-pogenic soil whose originandimportanceis thesubjectof debate(Denevan1996;Meggerset al.1988; Smith 1980), but is generally associatedwith forestsenrichedwith useful species (Smith1995)] and Santaremarea contains the oldestpottery yet found in the Americas-8000 BP(Roosevelt et al. 1991). Lathrap (1977) andLeon (1987:29,31) hypothesizedthatthis regionmay have been a nuclearcenterof agriculturalorigins. Giacometti(1992) recognizedthe east-ern portionof this center as part of his lowerAmazon centerand the westernportionas partof his Manaus/Roraimacenter.A few annualandperennialcrops are still clearly associatedwiththis area, but in general it is severely eroded.The crop list probablyincluded the followingdomesticates: Rollinia mucosa, Xanthosomasagittifolium,Crescentiacujete,perhapsCucur-bita moschata, Dioscorea trifida, Poraqueibaparaensis,Arachishypogaea,Phaseolusvulgar-is, Calatheaallouia, Bactrisgasipaes Paraland-race, Paullinia cupana var. sorbilis, Pouteriacaimito, Capsicumchinense, and othervegeta-bles, condiments,drugs and poisons. Some ofthe semi-domesticateswere: Anacardiumocci-dentale, Spondiasmombin,Inga spp., Loncho-carpus utilis, Psidium guajava, Astrocaryumaculeata, Theobromacacao. Some of the im-portantincipient domesticates were: Caryocarvillosum,Bertholletiaexcelsa, Lecythispisonis,Hymenaeacourbaril,Theobromagrandiflorum,andT.speciosum.Leersiahexandraand/orOry-za glumaepatulamay have been importantce-real crops, althoughtheir placementalong thedomesticationcontinuumis in doubt (Clement1999).At least four minorcentersof diversitymayhave existed also. These were all relatedto thesocieties that built earthworks:MarajoIsland,the Llanos de Mojos, the lower to middle Ori-noco River, and the Guiana coastal lowlands.The societies that createdthe earthworkswereeither acculturatedsoon after contact and theiragriculturaltechnologies were lost, includingtheircrops,or the societies responsiblewere nolongerpresentat contact,althoughthe areawasinhabitated,as occurredon MarajoIsland (cfMeggers and Danon 1988). Consequently,croplists for these areas are extremelyconjectural;they will be filled in as archeologicalresearchadvancesat each site.3. The Marajo Island Minor Center. Roo-sevelt (1991:25-26,113) identifiedthe followingdomesticates:Xanthosomasagittifolium,Manih-ot esculenta, Zea mays. Many of the domesti-cates of the Lower Amazonian Center wereprobablypresentas well. Among the semi-do-mesticatesSpondiasmombin,Platonia insignis,Inga spp., Psidium guajava, Astrocaryumvul-gare, Pouteriamacrophyllaand/orP. macrocar-pa, and Theobromacacao were probablypres-ent. Roosevelt (1991:25) emphasizesthe proba-ble importanceof Leersia hexandraon MarajoIslandbeforeandimmediatelyaftercontact.4. TheLlanos de Mojos Minor Center. Zhu-kovskys(1975) mapof theSouthAmericanme-gacenter suggests that this minor center is itsonly extension into the tropical lowlands, al-thoughthereis no discussion.Its croplist prob-ably included Phaseolus vulgaris, P. lunatus,possibly Cucurbitamoschata and C. maxima,Arachishypogaea,Pachyrrhizustuberosus,Bac-tris gasipaes (which may have originatednear-by), Passiflora edulis, Capsicumbaccatum,Si-cana odorifera and other vegetables, condi-ments, drugs and poisons [Denevans (1966)crop list confirmspart of this list]. Semi- andincipientdomesticatesmay have included:Ingaspp.,Bertholletiaexcelsa,Bunchosiaarmeniaca,Psidiumguajava, Lucumaobovata, Theobromabicolor, Caryocarglabrum,Hymenaeacourbar-il, Acrocomiasclerocarpa,and others.5. The Middle Orinoco Minor Center. Thisminor centerhas also not been surveyedfor apaleoethnobotanicalcrop list. Down-riverfromthe areawith earthworks,Roosevelt (1980) dis-cussedthe arrivalof Zea maysatabout2800 BPandhow it supplantedManihotesculentaas themajorstaple. Otherdomesticatesfrom northernSouth America may have included: Annonamuricata,Xanthosomasagittifolium,Ananaser-ectifolius, possibly Carica papaya, Dioscoreatrifida,Canavaliaensiformisand C.plagiosper-ma, Phaseolus vulgaris,Marantaarundinacea,Capsicumchinense,andothers.6. The Guiana Minor Center. This minorcenteris proposedbecause of probablepopula-tion density and presence of earthworks,andwas probablyinfluencedby bothAmazonianandCaribbeancrop complexes. Domesticates thatmay have been presentwere Annonamuricata,
  11. 11. 212 ECONOMIC BOTANY [VOL. 53Xanthosoma sagittifolium, Crescentia cujete,possibly Carica papaya, Cucurbita moschata,Dioscorea trifida, Arachis hypogaea, Canavaliaensiformis and C. plagiosperma, Phaseolus vul-garis, Gossypium hirsutum, Capsicum chinenseand others. Some importantsemi-domesticateswere probably Annona reticulata, Mammeaamericana, Byrsonima crassifolia, Psidium gua-java, andothers,while incipientdomesticatesin-cluded Caryocar nuciferum, Hymenaea cour-baril, Melicoccus bijugatus and others.The five regions of diversitywere the Ama-zon River estuary,the middle Solim6es River,the upperRio Negro and Orinoco Rivers, theupperAmazonRiver,includingthe Ucayali andthe HuallagaRivers, and the Guianacoast. Allmay have been densely populatedat contact,withthepossibleexceptionof theupperRio Ne-gro andOrinocoRiverregion,whichhas an ap-parently less favorable physical environment(Sponsel 1986).7. The Amazon Estuary Region includestheestuaryand adjacentAtlanticcoast, both northandsouth.Itwas proposedby Giacometti(1992)as a centerandis conservedhereas a regionofdiversity.ItencompassestheMaraj6MinorCen-ter and extends up-riverto the Lower Amazo-nian Center.Most of the crops listed for theLowerAmazonianCenterandtheMarajoMinorCenteroccurredhere also.8. The Solim6es Region extended from theLower Amazonian Center in the east to theNorthwesternAmazonianCenterin thewest andincludedthe lowerreachesof the SolimoesRiv-er tributaries.This was the region occupiedbythe Machiparaand Omaguachiefdoms at con-tact. It probablycontained many of the cropsfrombothcentersandwas in theprocessof cre-atingits own advancedlandracesof Bactrisgas-ipaes and Solanum sessiliflorum, among others(Clement 1988; D. F Silva Filho, INPA, 1998,pers.com.).9. The Upper Negro/Orinoco Region prob-ably extendedfrom above modernBarcelos,onthe Rio Negro, throughthe Casiquiareto theconfluenceof the OrinocoandGuaviareRivers,perhapseven as far as the Meta River,in Ven-ezuela, and includedat least the lower reachesof the Inirida,Ilana, and VaupesRiver basins.Partof it was proposedby Giacometti(1992) asa center and is conserved here as a region ofdiversity.This region probablycontainedmanyof the crops of the NorthwesternAmazonianCenter,but was creating its own landraces.Itcontainedat least two landracesof Bactrisgas-ipaes, the Vaupes and the Inirida (Clement1988), one of Solanumsessiliflorum,andwas intheprocessof domesticatingHeveaspp.fortheirseeds. This region is probablythe origin of theSmooth Cayenne variety of Ananas comosus,possibly of A. erectifolius,andof Paullinia cu-pana.10. The Upper Amazon Region, extendedsouthfromthe NorthwesternAmazonianCenteralongtheAmazonRiver,andincludedtheupperNapo River and Ucayali and HuallagaRivers.As well as containingmanyof the crops of theNorthwesternAmazoniancenter,it mayhaveex-periencedconsiderablein-putfromthe mid-ele-vationAndeanregion.Because the upperNapoRiveris adjacentto a low section of the Andes,it may be where crops such as Caricapapayaand Persea americanaenteredAmazoniafromthe northernPacific coast. There are severallandracesof Bactrisgasipaes, only one of which(the Pampa Hermosa) has been adequatelymapped (Clement 1988). It is probablywhereEugeniastipitataoriginated(Clement1989).11.TheGuiana Coastal Region probablyex-tendedfromthe AmazonEstuaryRegion in thesouthto the OrinocoRiverestuaryin the north,and includedthe GuianaMinorCenter.Conse-quentlyit probablycontainedmanyof thecropsof the GuianaMinor Center and some of theAmazonEstuaryRegion, as well as cropsfromthe Caribbean.A SOUTHAMERICANSYNTHESISAssuming that these Amazonian concentra-tions of crop genetic diversity existed asmapped,how do they fit into a SouthAmericanmosaic?In Fig. 4, I extrapolatemy thinkingforAmazoniato the rest of the continent,althoughwithoutsupplyingcrop lists. Only a shortjusti-ficationis given for each SouthAmericancon-centration.The Andean mosaic. Vavilovs (1992f) finalapproximationof two Andeancentersserves asthe basis for anchoringthe Andeanmosaic.TheColombiancenter is supportedby analysis ofmaize races in South America (Brieger et al.1958), as well as by the large numberof cropsthatarerestrictedto the northernAndes (Patinio1963, 1964). The Peru/Boliviacenterwas wide-ly acceptedbeforeHarlan(1971) andis retainedhere because thereare numerouscrops that are
  12. 12. 1999] CLEMENT:AMAZONIANCROPRESOURCESII 213CentersMinorCenters4 **. (7)~~~~~~Regions0~~~~~~1030S 30SFig. 4. The biogeographyof SouthAmericain 1492. Centersof Diversity:1. NorthwestemAmazonia;2.CentralAmazonia;3. Peru-Bolivia;4. Colombia.MinorCentersof Diversity:5. PeruvianCoast. Regions ofDiversity:6. MarajoIsland;7. Llanosde Mojos;8. MiddleOrinoco;9. Guiana;10.Mid-elevationAndeanColom-bia-Ecuador-Peru-Bolivia;11.Guarani;12. AmazonEstuary;13. Solimoes;14. UpperNegro/Orinoco;15.UpperAmazon;16. GuiananCoast;17.NorthernArgentina;18. AtlanticForest;19.NortheasternBrazilianCoast.endemic there or show exceptional variabilitythere(Harlan1992:217-236). Zhukovskysme-gacenteris transformedinto a region of diver-sity, following Hawkes (1983), but is restrictedto the mid-elevationAndes,becauseof the croplists presentedby Pearsall(1992) that are spe-cific to the mid-elevations,ratherthanincludingthe Llanos de Mojos and the NW Argentineanlowlands(see lowlandmosaicbelow). ChiloeIs-land and adjacentmainlandare not conservedbecauseonly two very minorcropsoccurthere,and other crop genetic variationwas probablyinsufficientto justify its status as a region ofdiversity.Partof the Peruviancoast is consid-ered a minorcenter,following Vavilovsearlierproposal(1992e) andgiven the greatnumberofspecies foundin archaeologicalsitesthere(Pear-sall 1992). This is definitelyan areathataccu-mulatedcrops,ratherthandevelopingthemfromendemicspecies,however,becauseall of thedo-mesticatesin the croplists arenon-nativeto thearea.The lowland, non-Amazonianmosaic. Vavi-lovs originalBrazil/Paraguayfocus is associat-ed with high Guaranipopulation densities atcontactand variouscrops (Vavilov 1992c) andis conserved here as the Guaraniregion of di-versity, in agreementwith Giacometti(1992).NorthwesternArgentinais anotherregionof di-versity,supportedby maize (Briegeret al. 1958)and recent archaeologicaldata (Pearsall 1992).Severalsectionsof coastalBrazilwereoccupiedby relatively dense Tupinambapopulationsatcontact (Hemming 1978) and had efficient ag-
  13. 13. 214 ECONOMICBOTANY [VOL.53riculture. Giacomettis (1992) suggestion forthreecentersis transformedinto a single regionof diversity,because theredoesnt appearto bea break in crop distributionsalong the south-easterncoast. Consequently,a region of diver-sity may have extendedalong the SE Braziliancoast, the Atlanticforest ecosystem, frommod-em Sao Paulo to Pernambuco,while a secondextends from Piaui west to the Amazon alongthe NE coast, wherecashewmay have originat-ed (Cundall1995).Infact,Brucher(1969)andHarlan(1971)werecorrect:SouthAmericahastoo muchdiversitytoconsiderVavilovsoriginalhypothesisa sufficientcharacterization.Figure 4 also suggests thatBruchersarcsof diffusionwere probablybasedon the distributionof humanpopulationdensitiesalongthemajorriversystems,coastsandtheAn-des, wherediffusionfromone humanpopulationto the next facilitatedcrop distribution.Further-more,most of the areasnot includedin this mo-saic probablycontainedcrop genetic resources,eitherat lowerconcentrationsor at levels notyetidentified.The hypothesispresentedherewill re-quireconsiderableworkto confirm,butevenafter500 yearstheremainsof theAmerindianheritage,both biological and archaeological,are impres-sive enoughto permitthis.ACKNOWLEDGMENTSI thankWilliamBal6e,JamesL. Brewbaker,RolandC.Clement,Dan-iel G. Debouck,WilliamM. Denevan,JackR. Harlan,JohnG. Hawkes,CharlesB. Heiser,Jr.,JorgeLe6n,PauloS. Martins,BettyJ. Meggers,VictorM. Patiino,DeborahPearsall,BarbaraPickersgill,DoloresR. Pi-perno,GhilleanT. Prance,AnnaC. Roosevelt,Marciode M. Santos,Nigel J. H. Smith,Les E. Sponsel,LyndonWester,DavidE. Williamsandan anonymousreviewerfornumeroussuggestionsandcriticismsonearlyversionsof thismanuscript.Errorsof factandinterpretationare,ofcourse,theauthorsresponsibility.LITERATURE CITEDBrieger, F. G., J. T. A. Gurgel, E. Paterniani, A.Blumenschein,and M. R. Alleoni. 1958.Racesofmaize in BrazilandothereasternSouthAmericancountries.Publication593. NationalAcademy ofSciences/NationalResearchCouncil,Washington,DC.Brucher, H. 1969. Gibtes gen-zentren?Naturwissen-schaften56:77-84.1990. Difusion transamericanade vegetalesutiles del neotropicoen la epoca pre-colombina.Pages 265-283 in D. A. Posey, andW.L. Overal,eds., EthnoBiology:implicationsand applications.Proceedingsof the FirstInternationalCongressofEthnoBiology. Museu Paraense Emnlio Goeldi/CNPq/SCT,Belem, PA, Brazil.de Candolle, A. 1882/1908. Origin of cultivatedplants.D. Appleton,New York.Clement, C. R. 1988. Domesticationof the pejibayepalm (Bactris gasipaes): past and present. Pages155-174 in M. J. Balick, ed., The palm-tree oflife. Advancesin EconomicBotany6. New YorkBotanicalGarden,New York.. 1989. A centerof crop genetic diversityinwesternAmazonia.BioScience39:624-631.. 1999. 1492 andthe loss of Amazoniancropgenetic resources.I. The relationbetweendomes-ticationand humanpopulationdecline. EconomicBotany53(2):188-202.Cundall, E. P. 1995. Cashew,Anacardiumocciden-tale (Anacardiaceae).Pages 11-13 inJ.Smartt,andN. W. Simmonds,eds., Evolutionof crop plants,2nd ed. Longman,London.Darlington, C. D. 1973.Chromosomebotanyandtheoriginof cultivatedplants.3rded. Allen & Unwin,London.Darlington, C. D., and E. K. Janaki Ammal. 1945.Chromosomeatlas of cultivatedplants. Allen &Unwin,London.Denevan, W. M. 1966. The aboriginalculturalgeog-raphyof the Llanos de Mojos de Bolivia. Ibero-Americanano. 48. Universityof CaliforniaPress,Berkeley.. 1992. Native Americanpopulationsin 1492:recentresearchanda revisedhemisphericestimate.Pages xvii-xxxvii in W.M. Denevan,ed. The na-tivepopulationof theAmericasin 1492.Universityof WisconsinPress,Madison,Wisconsin.. 1996. A bluff modelof riverinesettlementinprehistoricAmazonia.AnnalsAssociationAmeri-can Geographers86(4):654-681.Giacometti, D. C. 1992. Recursosgeneticos de fru-teirasnativasdo Brasil.Pages 13-27 in Anais dosimposionacionalde recursosgeneticosde fruteir-as nativas.CentroNacionalde Pesquisasde Man-diocae Fruticultura-EMBRAPA,CruzdasAlmas,BA, Brasil.Harlan, J. R. 1971. Agriculturalorigins:centersandnoncenters.Science 174:468-474.. 1975. Cropsand man. AmericanSociety ofAgronomy/CropScienceSocietyof America,Mad-ison, Wisconsin.. 1992. Cropsandman.2nd ed. AmericanSo-ciety of Agronomy/CropScienceSocietyof Amer-ica, Madison,Wisconsin.Hawkes,J. G. 1983.Thediversityof cropplants.Har-vardUniversityPress,Cambridge,Massachusetts.Hemming,J. 1978.Redgold:theconquestof theBra-zilian Indians. HarvardUniversity Press, Cam-bridge,Massachusetts.Kuptsov, A. I. 1955.Geographicaldistributionof cul-tivatedfloraandits historicaldevelopment.Bulle-tin of theAll UnionGeographicalSociety87:220-231.
  14. 14. 1999] CLEMENT:AMAZONIANCROPRESOURCESII 215Lathrap, D. W. 1970. The upperAmazon. Praeger,New York.. 1977. Ourfatherthe cayman,ourmotherthegourd:Spindenrevisited,ora unitarymodelfortheemergenceof agriculturein the New World.Pages713-753 in C. A. Reed,ed., Originsof agriculture.MoutonPubl.,The Hague.Leon, J. 1987. Botanicade los cultivostropicales.In-stituto Interamericanopara la Cooperaci6nAgri-cola-IICA, SanJose, CostaRica.Levi-Strauss, C. 1950.Theuse of wild plantsin trop-ical SouthAmerica.Pages465-486 in J. H. Stew-ard, ed., Handbookof South AmericanIndians.SmithsonianInstitution,Bureauof AmericanEth-nology, Washington,DC.MacNeish, R. S. 1992.The originsof agricultureandsettledlife. Universityof OklahomaPress,Norman,Oklahoma.Meggers, B. J. 1988. The prehistoryof Amazonia.Pages53-62 in J. S. Denslow,andC. Padoch,eds.,People of the tropicalrainforest.Univ. CaliforniaPress,Berkeley., and J. Danon. 1988. Identificationand im-plicationsof a hiatusin thearcheologicalsequenceon Maraj6Island,Brasil.J. WashingtonAcademyof Sciences78:245-253., 0. F. Dias, E. T. Miller, and C. Perota.1988.Implicationsof archeologicaldistributionsinAmazonia.Pages 275-294 in W. R. Hayerand P.E. Vanzolini,eds., Proceedingsof a workshoponNeotropicaldistributionpatterns.AcademiaBrasi-leirade Ciencias,Rio de Janeiro.Mora Urpi, J. 1992.Pejibaye(Bactrisgasipaes).Pag-es 209-220 in J. E. HernmndezBermejo,and J.Le6n, eds., Cultivos marginados:otraperspectivade 1492. Food and AgricultureOrganization-FAO/JardinBotanicode C6rdoba(Espafia),Rome.Patifio, V. M. 1963.Plantascultivadasy animalesdo-mesticosen AmericaEquinoccial.TomoI.Frutales.ImprentaDepartamental,Cali,Colombia.. 1964. Plantascultivadasy animalesdomesti-cos en AmericaEquinoccial.TomoII. Plantasali-menticias.ImprentaDepartamental,Cali, Colom-bia.. 1992. An ethnobotanicalsketchof the palmBactris (Guilielma) gasipaes. Principes 36(3):143-147.Pearsall, D. M. 1992.The originsof plantcultivationin SouthAmerica.Pages 173-206 in C. W.Cowan,andP.J. Watson,eds., Theoriginsof agriculture-an internationalperspective.SmithsonianInstitu-tion Press,Washington,DC.Pickersgill, B., and C. B. Heiser, Jr. 1977. Originsanddistributionof plantsdomesticatedin theNewWorldTropics.Pages 803-836 in C. A. Reed,ed.,Originsof agriculture.MoutonPubl.,The Hague.Rindos, D. 1984. The originsof agriculture:an evo-lutionaryperspective.AcademicPress,SanDiego,CA.Roosevelt, A. C. 1980. Parmana:prehistoricmaizeandmaniocsubsistencealongtheAmazonandOri-noco. AcademicPress,New York.. 1989.Lostcivilizationsof thelowerAmazon.NaturalHistory98:74-83.. 1991. Moundbuildersof the Amazon:geo-physicalarchaeologyon Maraj6Island,Brazil.Ac-ademicPress,SanDiego., R. A. Housley, M. I. da Silveira, S. Mar-anca, and R. Johnson. 1991. Eighthmillenniumpotteryfroma prehistoricshell middenin the Bra-zilianAmazon.Science 254:1621-1624., M. Lima da Costa, C. Lopes Machado, M.Michab, N. Mercier, H. Valladas, J. Feathers,W. Barnett, M. Imazio da Silveira, A. Hender-son, J. Silva, B. Chernoff,D. S. Reese, J. A. Hol-man, N. Toth, and K. Schick. 1996. Paleoindiancave dwellersin the Amazon:the peoplingof theAmericas.Science 272:373-384.Sauer, C. 0. 1952.Agriculturaloriginsanddispersals.AmericanGeographySociety,New York.Smith, N. J. H. 1980.Anthrosolsandhumancarryingcapacityin Amazonia.Annals of the Associationof AmericanGeographers70:553-566.. 1995. Human-inducedlandscapechangesinAmazoniaandimplicationsfordevelopment.Pages221-251 in B. L. TurnerII,A. GomezSal,F Gon-zalez Bermnldez,andF di Castri,eds., Globallanduse change-a perspectivefromtheColumbianen-counter.ConsejoSuperiorde InvestigacionesCien-tificas,Madrid.Sponsel, L. E. 1986.Amazonecology andadaptation.Ann. Rev. Anthropol.15:67-97.Vavilov, N. I. 1992a. Centersof origin of cultivatedplants(Firstpublishedin 1926 in Tr.po prikl.bot.i selek. (Papers on Applied Botany and PlantBreeding)16(2)).Pages22-136 in V. F Dorofeyev,ed., Origin and geography of cultivatedplants.CambridgeUniversityPress,Cambridge,UK.. 1992b.Ontheoriginof cultivatedplants(Firstpublishedin 1926 in Novoye v Agronomii(Newsin Agronomy)).Pages 14-21 in V. F Dorofeyev,ed., Origin and geography of cultivated plants.CambridgeUniversityPress,Cambridge,UK.. 1992c.Thephyto-geographicalbasisforplantbreeding(Firstpublishedin 1935in Teoreticheskieosnovy selektsii (Theoretical Bases for PlantBreeding) 1). Pages 316-366 in V. F Dorofeyev,ed., Origin and geography of cultivated plants.CambridgeUniversityPress,Cambridge,UK.. 1992d.The problemconcerningthe originofagriculturein thelightof recentresearch(Firstpub-lishedin 1931in Sots.rekonstruktsiyai nauka(So-cialist reconstructionand Science) 1). Pages 173-183 in V. F Dorofeyev,ed., Originandgeography
  15. 15. 216 ECONOMICBOTANY [VOL.53of cultivatedplants. CambridgeUniversityPress,Cambridge,UK.. 1992e.The problemconcerningthe originofcultivatedplants as presentlyunderstood(Speechgiven atthe All-SovietCongressof Genetics,PlantBreeding, Agriculture and Livestock Breeding,Leningrad,1929).Pages 158-172 in V. F Dorofey-ev, ed., Originandgeographyof cultivatedplants.CambridgeUniversityPress,Cambridge,UK.. 1992f. The theoryof the originof cultivatedplants after Darwin (First publishedin 1940 inNauka (Science) 2). Pages 421-442 in V. F Do-rofeyev, ed., Origin and geographyof cultivatedplants. CambridgeUniversity Press, Cambridge,UK.. 1997.Five continents.InternationalPlantGe-netic ResourcesInstitute,Rome.Zeven, A. C., and P. M. Zhukovsky. 1975. Dictio-naryof cultivatedplantsandtheircentresof diver-sity, excludingornamentals,foresttreesandlowerplants.CentreforAgriculturalPublishingandDoc-umentation,Wageningen.Zhukovsky, P. M. 1968. New centresof origin andnew gene centres of cultivatedplants, includingspecificallyendemicmicrocentresof speciesclose-ly allied to cultivatedspecies. BotanicalJournal(Moscow)53:430-460.. 1975.Worldgenepool of plantsforbreeding.Mega-genecentersandendemicmicro-genecenters.USSR Academyof Sciences,Leningrad.BOOK REVIEWAncient Ammonites & Modern Arabs 5000 Yearsin the Madaba Plains of Jordan. London,G. A.andClark,D. R., editors.(1997). AmericanCenterfor OrientalResearch, P.O. Box 2470, Amman11181,Jordan.iix + 71 pp.Hardcover.$27 includ-ing postage."This books describesthe past and presentpeopleof the Amman/Madabaregionin centralJordan,butitis also a studyof ourselveswithaneye to thefuture,"state the authorsin the conclusion.I cant find a lotaboutourselvesandourfuture,butthe book does saya greatdeal about the MadabaPlains, a fertile areaabout25 km southof the capitolcity of Amman.Am-man derivesfromAmmonites,a people mentionedinthe Bible who inhabitedtheseplains.The twelve chaptersare well written,well edited,informativeandinteresting.Readersof EconomicBot-any will be particularlyinterestedin chaptersVII andVIII thatinclude agricultureand foods. As a studentof Middle East ethnobotany,especially plantsof theBible, I foundthesetwo chaptersinformative.Not sur-prisingly,barley,flax, lentils, andwheatarefoundinthe oldest sites. Otherseeds includeolive, pomegran-ate, wild pistachio (probably Pistacia palaestina,though no scientific names are given), grape,vetch,pea, andchickpea-all plantsstill usedextensivelyinJordan.Buckwheat"seeds"werefoundin a Hellenis-tic age site. Interesting,as I amunawareof buckwheatgrownanywherein the region.Whatis unusualaboutthevolumeis a skillfulweav-ing of ancientand modernpractices,effectively por-trayinglife in the MadabaPlains for five millennia.Attractivecolored illustrationsgrace the book. Quo-tationsfromthe QuranandBible drawupontherichtraditionsof bothreligionsin the region.A glossary,chronology,and bibliographyare included.Unfortu-nately,thereis no index. Despite this, the book willbe of value to ethnobotanistsworkingin the MiddleEast.The level of presentationis clearandsimple sothatI wouldrecommendit for touristsandothervisi-torsto Jordan.And at $27, the book is a bargain.LYTrONJOHNMUSSELMANDEPARTMENTOF BIOLOGICALSCIENCESUNIVERSITYOF JORDAN11954AMMAN, JORDAN

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