Pery Domesticacion de La Yuca y La Batata

7/29/2019 Pery Domesticacion de La Yuca y La Batata

1/16

Starch Granule Size and the Domestication of Manioc (Manihot esculenta) and Sweet Potato(Ipomoea batatas)Author(s): Linda PerrySource: Economic Botany, Vol. 56, No. 4 (Winter, 2002), pp. 335-349Published by: Springer on behalf of New York Botanical Garden PressStable URL: http://www.jstor.org/stable/4256606 .

Accessed: 12/09/2013 09:05

Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at .http://www.jstor.org/page/info/about/policies/terms.jsp

.JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of

content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms

of scholarship. For more information about JSTOR, please contact [email protected].

.

New York Botanical Garden Press and Springerare collaborating with JSTOR to digitize, preserve and extend

access toEconomic Botany.

http://www.jstor.org

This content downloaded from 150.185.172.170 on Thu, 12 Sep 2013 09:05:07 AMAll use subject to JSTOR Terms and Conditions
http://www.jstor.org/action/showPublisher?publisherCode=springerhttp://www.jstor.org/action/showPublisher?publisherCode=nybghttp://www.jstor.org/stable/4256606?origin=JSTOR-pdfhttp://www.jstor.org/page/info/about/policies/terms.jsphttp://www.jstor.org/page/info/about/policies/terms.jsphttp://www.jstor.org/page/info/about/policies/terms.jsphttp://www.jstor.org/page/info/about/policies/terms.jsphttp://www.jstor.org/page/info/about/policies/terms.jsphttp://www.jstor.org/stable/4256606?origin=JSTOR-pdfhttp://www.jstor.org/action/showPublisher?publisherCode=nybghttp://www.jstor.org/action/showPublisher?publisherCode=springer


2/16

STARCH GRANULE SIZE AND THE DOMESTICATION OF MANIOC(MANIHOT ESCULENTA)AND SWEET POTATO (IPOMOEABATATAS)'LINDA PERRY

Perry, Linda (Department of Anthropology, Southern Illinois University, Carbondale, IL62901). STARCH GRANULE SIZE AND THE DOMESTICATION F MANIOC (MANIHOT ESCULENTA) ANDSWEETPOTATO IPoMoEA BATATAS). Economic Botany 56(4):335-349, 2002. Archaeological stud-ies of plant remains have indicated that an increase in seed size is frequently correlated withboth intensive cultivation and domestication of seed crop plants. To test if starch granules ofdomesticated root crops are significantly larger than those of wild or less intensively cultivatedplants, archaeological and modern specimens of manioc and sweet potato were sampled forstarch granules, and granule size was compared across a temporal sequence. The results in-dicate that a gross generalization can be made that modern specimens of both manioc andsweet potato yield larger starch granules than some archaeological specimens. It does appear,however, that modern domesticated manioc roots produce significantly larger-sized starch gran-ules than those of its purported wild ancestor. Additionally, there exist two lines of evidencethat the coastal Peruvian and lowland Neotropical regional types of manioc differ from oneanother and have been separate for several millennia. These findings indicate that manliocmayhave been domesticated more than once.Key Words: Manihot esculenta; Ipomoea batatas; starch granules; plant domestication; rootcrops.

The cultivation of plant species by humansresults in significant changes in the nature ofthe plant. Both biological and geographicalmodifications can be interpreted by archaeolo-gists as indicators of intensive use or even do-mestication of a given taxon. A domesticatedcrop species is frequently cultivated outside itswild range, and, often, mechanisms of seed dis-persal necessary to the survival of wild popu-lations are lost. Further, a domesticated plant isoften discernible from its wild counterpart dueto the morphological differences of one or moreorgans that are favored for use by the cultiva-tors. These differences are often observable inthe plant remains derived from archaeobotani-cal assemblages.One of the most commonly recognized mark-ers of the domestication of seed-bearing plantsis an increase in cultivated seed size over wildtype seeds, and researchers frequently cite thistransformation as evidence of intensive cultiva-tion (Smith 1995:23). Indeed, increase in seedsize concomitant with the processes of domes-

' Received 17 January2000; accepted 15 December2001.

tication has been noted in many species of plantsin several parts of the world.In the Midwest of the United States, sunflow-er (Helianthus annuus) and sumpweed (Iva an-nua var. macrocarpa) seeds from archaeologicalcontexts were designated "domesticates" due totheir size. The size of the charred archaeologicalseeds from prehistoric cultivars is consistentlygreater than that of seeds from modern wild typepopulations of the same species of plant (Yarnell1972, 1978). Archaeological representatives ofcultivated grain crops from the Middle East havebeen described as having both larger sized and"plumper" grains than their non-cultivar con-specifics (Miller 1992:44). The phenomenon oflarger seed size in domesticated seed crops hasalso been noted in common beans (Phaseolusvulgaris) from South America (McClung deTapia 1992:153), as well as from barnyard millet(Echinochloa utilis) from China (Crawford1992). Uniformity of seed size may also be amarker of a domesticated seed crop. This phe-nomenon was noted in an assemblage of sun-flower seeds from the U.S. Midwestem site,Newt Kash (Jones 1936).It is fair to say that a large seed size in com-

Economic Botany 56(4) pp. 335-349. 2002(C2002 by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.

This content downloaded from 150.185.172.170 on Thu, 12 Sep 2013 09:05:07 AM

All use subject to JSTOR Terms and Conditions
http://www.jstor.org/page/info/about/policies/terms.jsphttp://www.jstor.org/page/info/about/policies/terms.jsphttp://www.jstor.org/page/info/about/policies/terms.jsp


3/16

336 ECONOMICBOTANY [VOL. 56

parison to wild-type seeds or a gradual increasein seed size over time can be used as a markerfor the cultivation of and domestication of ar-chaeological seed crop species. With this trendin mind, the question arises as to whether or nota similar morphological change would occurduring the domestication of plants that producestarch-bearing underground storage organs.It is reasonable to assume that, for the sakeof a more bountiful harvest, humans would se-lectively propagate those plants that producedlarger underground storage organs, and, there-fore, more carbohydrate in the form of starch.The meter-long roots of a modern domesticatedVenezuelan bitter manioc plant dwarf the ten-centimeter root of modern Brazilian wild-typemanioc in this author's collection. However,though all manioc roots produce starch granulesof a typical morphology, whether or not a largerroot would store starch in larger-sized grains, orsimply create many more granules of the same,small size is not clear. The data collected in oneinvestigation, however, do suggest that a trendsimilar to that found in domesticated seed cropsmay occur in the starch grains of tuberous crops.In a study designed to delineate diagnosticmorphological characteristics of potato starchgrains, the size of the granules of the domesti-cated species (Solanum tuberosum) was, on av-erage, larger than that of most of the wild spe-cies of Solanum (Ugent and Verdun 1983). Onlyone species of the eighteen wild type tubersyielded larger granules than those of the culti-vated species. Ugent has noted that wild potatofrequently contains "much smaller [starch]grains" than those of a cultivated potato (1982:191). Unfortunately, at this time there exists noarchaeological sequence of data that demon-strates a gradual increase in starch grain size asuse intensifies and domestication occurs. Theabovementioned study, however, indicates thatlooking for such a pattern may be a worthwhileendeavor.

Starch granule size data collected from botharchaeological manioc and sweet potato speci-mens correspond to an archaeological sequencefrom the Peruvian coast (see below). It is hy-pothesized that starch grain, and, therefore,granule size will increase as the intensity of cul-tivation practices increase or domestication oc-curs. Therefore, if the samples in this study werecollected from a temporal range that encom-passes the intensification of cultivation or do-

mestication of the root crops, it is hypothesizedthat specimens from younger contexts will yieldlarger starch granules than those from oldersites. Modern starch granules from manioc rootscollected in the Upper Rio Negro region of Ve-nezuela, the north coast of Peru, as well asstarch granules from Manihot esculenta ssp. fla-bellifolia, the purported wild ancestor of domes-ticated manioc, will be included in the analysesof the archaeological manioc data for compara-tive purposes. A sample of modem sweet potatostarch will be included in the analyses of thosearchaeological data.

Further, the uniformity of starch granule sizewill be examined across the same independentvariables. If larger roots make more small starchgrains rather than larger grains, perhaps the as-semblage will become more consistent in size,as did the assemblage of sunflower seeds fromNewt Kash.

If the phenomenon of size increase is com-mon to both seeds and starch grains, one wouldgain the ability to document the domesticationof root and tuber crops over time and/or spaceusing the data from starch granule analyses.Considering the significance of root crops in thediets of the prehistoric inhabitants of tropical re-gions, this information could certainly clarifyour understanding of prehistoric subsistence.

DESCRIPTION OF THE COLLECTIONSThe vast majority of the manioc and sweet

potato specimens examined in this study wereon loan from the archaeological collections ofDr. Donald Ugent, Department of Plant Biology,Southern Illinois University Carbondale (desig-nations M.e.A., and I.b.A.). The archaeologicalspecimens were collected and identified by Dr.Ugent during his work in coastal Peru (see Ug-ent et al. 1981, 1985), and were curated in hislaboratory in the Department of Plant Biologyat SIUC. Identifications of the root specimenswere completed by Ugent using a combinationof features including surface morphology, starchgranule morphology, and some vascular anato-my (see Ugent et al. 1981, 1985).

Collections from Pampa de las Llamas-Mox-eke included both manioc and sweet potato rootsthat were sampled in this study. Pampa de lasLlamas-Moxeke is a major early ceramic site lo-cated in the south bank of the Casma Valley onthe North coast of Peru about 18 km from thePacific Ocean (Ugent et al. 1985). A 2 km2 site




4/16

2002] PERRY:STARCHSIZE IN MANIOCAND SWEETPOTATO 337area included mounds and domestic structuresbordered by midden deposits that were sampledfor plant remains. Ceramic material and otherartifacts indicate that Pampa de las Llamas-Moxeke was occupied during the Initial period,and radiocarbon samples yielded dates rangingfrom 1785 to 1120 B.C. (Ugent et al. 1986).Several other species of plants were also iden-tified from the midden deposits (Ugent et al.1986). These collections include peanut, achira,squash, cotton, gourd, beans, potatoes, and othertuberous crop plants. All materials were verywell preserved due to the arid coastal environ-ment, and some material was partially charred.A single specimen of manioc was examinedfrom the collections from Las Haldas. Las Hal-das is another coastal site located about 20 kmto the south of the Casma Valley in Peru (Ugentet al. 1986). Most of the 40 ha site is middendeposit, and the central area contains a moundstructure, a plaza, and smaller, flanking mounds.The manioc root was collected in a sample of a"squatter settlement" midden that dates to afterthe construction of the temple mound structures,and contains the remains of plants that weretransported into the site from further inland(Matsuzawa 1978; Ugent et al. 1986). Three ra-diocarbon samples from the midden yieldeddates ranging from 1040 to 895 B.C.In addition to the manioc root, the middendeposits at Las Haldas yielded other species ofplants. Peanut, pepper, squash, gourd, cotton, av-ocado, bean, and maize as well as lucuma seedswere recovered. Again, plant remains were pre-served under very arid conditions.Manioc roots were also sampled from the col-lections of the Pampa Rosario site. Pampa Ro-sario is located in the Casma Valley about 16km from the Pacific Ocean. The 40 ha site in-cludes about 20 structures, low, narrow platformmounds with small courtyards and connectedrooms, and midden areas ranging from one-halfto two meters in depth (Ugent et al. 1986). Ra-diocarbon dates from the midden excavationranged from 810 to 450 B.C.Other recovered plant remains included pea-nut, pepper, achira, squash, cotton, inga, avo-cado, gourd, cansaboca, bean, and maize. Plantswere preserved under arid conditions.Both sweet potato and manioc roots weresampled from collections from Manchan, orChimu Sur, another site in the Casma Valley.Chimu Sur covers an area of about 70 ha, and

was likely a Chimuadministrative enterratherthan a residential site (Ugent et al. 1986). Thesite includes nine compoundswith courtyards,storerooms,offices, andresidential ooms forthe"residentnobility" (Ugent et al. 1986). Whileplant remains from the other sites were frommidden deposits, the archaeobotanicalamplesfromChimu Surwere collected fromfloorsandplatformmounds.TheChimuoccupationat Chi-mu Sur dates to the 14th to mid 15th centuryA.D.Apart romthe rootremains,soursop,peanut,cansaboca,pepper,squash,coca, cotton,gourd,lucuma, ajipa, avocado, lima bean, commonbean,potato,and maize were collected fromthearchaeobotanicalamples.Again,ariditywas theprimarypreservative.A single archaeologicalmaniocrootwas sam-pled from a collectionmadeby Dr.DonaldUg-ent at a cemetery site in Chancay locatedjustnorthof the PeruviancapitalLima on thecentralcoast (designationPerul). The informationontheacessionlabel indicates hatthe rootwas col-lected from a midden at the "Chancayceme-tery" datingto aboutA.D. 1400.All archaeologicalmanioc and sweet potatospecimens, by modem standards, were rathersmall in size. The size of the dried rootsrangedbetween one and two centimeters n maximumdiameter,and length occasionally reachedfourcentimeters. The specimen from Chancaywassomewhat larger, about three by six or sevencentimeters.All roots sampled for starchgrainswere non-carbonized,desiccatedspecimens.Additionalmodem samples of manioc werederivedfrom threesources.The first is a pack-age of commercial"tapiocastarch"purchasedat a local market n Carbondale, llinois (desig-nationComm.ts.). This samplewas, in effect, alarge plastic bag of white powder,so little canbe said as to the size of the roots that yieldedthe starchtherein. It can be fairly securely as-sumed, however, that a number of differentplantswere processedto produce hecontentsofthe bag.The second set of sampleswerederivedfromcollectionsdepositedby Dr. ShirleyHoffman nthe Herbario Ovalles in the Universidad Centralde Venezuela,ocatedin Caracas,Venezuela,Dr.StephenTillet,curator designationMYF).Hoff-man collected these samples duringher disser-tationworkwith Arawakspeakinggroups n theUpperRio Negro, a blackwaterriver system in




5/16


southernVenezuela(Hoffman 1993). Cross-sec-tional slices of the manioc roots were providedwith each herbarium pecimen, and the diame-ters of the roots rangedfrom six to nine centi-meters.Thefinaltwo samplesof modernmanioc wereprovidedto the authorby Dr. Izumi Shimadawho collected the material n BatanGrandeonthe northcoast of Peru(designationMPeru).Theroots measured about 25 centimeters n length,and from threeto five centimeters n diameter.The single modem specimenof sweet potatowas purchasedat a local market n Carbondale(designationMonahl). The white boniato wasabout nineteen centimeters n length and eightcentimeters n diameterat its widest point.

METHODSFor desiccated archaeological roots and tu-bers,a small amountof starchwas scraped romthe cortex using a metal probe that had beencleaned with 95% ethanol. The starchwas thenstirred nto a dropof histomountmountingme-dium on a clean glass slide, and a cover glasswas placed on top. Modern roots and tuberswere sliced into sections, driedin a food dehy-dratorat a temperatureoo low to affect mor-phological changes on the starchgrains 1300 F),and then preparedas were the archaeologicalspecimens. Commercially prepared starcheswere stirred into the histomountwith no otherpreparation.Tocompile the comparativedata, the methodsof Loy et al. (1994) were modified.All obser-vations were made on a compound ight micro-scope equipped with a small numerical grid

printedon the slide stage.To obtain a sample ofstarchgrainsto measure,randomnumbersweregeneratedusing dice, and the slide was placedat these coordinates on the grid. All measure-mentswere collected at 400X magnificationus-ing an ocularmicrometer.The starchgranule nthe centermostposition on the micrometerwaschosen for measurement,and if there was nostarchgranule in contact with the micrometer,the ocular lens was rotatedclockwise until onewas intersected.Because measurementsnvolvethe rotation of the micrometerand the move-ment of the stage as one notes both the lengthand the width of an object, the measurement fone starchgranule usually places the scale incontactwith the next one to be measured.Thus,the sampling strategy employed here can be

very loosely referred to as stratifiedrandomsampling.Maximumwidth (Ml) and maximumlength(M2) measurements n microns were collectedalong with basic morphologicaldata such asshape, position of hilum, extinction cross ar-rangement,faceting, and fissuring patterns,ifany. Granules were observed in three-dimen-sions by pressinga probegentlyontothesurfaceof the cover glass. This pressurecreatesmove-ment in the mountingmedium,andthe granulesrotate. Length and width measurementswerenoted using the ocular micrometerhash markswith the tenth decimal unit estimated by eye(e.g., 10.5 with .5 estimated).Only single, freegranules were measuredto eliminate the possi-bility of a mismeasurementdue to the interfer-ence of anotherover- or underlyingobject.Tenstarchgranules were measuredin the field ofview of each set of coordinates,and five sets ofcoordinateswere chosen for each slide. Thismethod resulted in fifty measured starchgran-ules for each sample slide. If there were notenough granules to measure at a given set ofcoordinates,a new set was generated.Starchgranulemeasurements ndmorpholog-ical characteristicswere collected for the maxi-mum width andlengthof each starchgranule inmicrons),and a width/lengthratiowas calculat-ed fromthese data.Finally,sample standardde-viation for both width andlength measurementswas calculatedand noted for each specimen ofmaniocand sweet potato.The data were entered into a spreadsheetalong with approximatedates of depositionforarchaeological samples and known dates formodernsamples.Radiocarbondates for the Pe-ruviansites of Pampade las Llamas,LasHaldas,Pampa Rosariowere provided by Dr. Tom Po-zorski, and those of ChimuSur were providedby Dr. CarolMackey.The database iles were imported nto the PCSAS program, where various statistical testswere performed.Regressionanalyseshad beenperformed n Lotus 123 priorto SAS analysesin an attempt o correlatedates with starchgran-ule size. Because regression analyses provednon-significant,Tukey'sHighly SignificantDif-ferencetest, a conservativemultiplecomparisontest, was performed n an attemptto discern ifany significantdifferencesexisted between anyof the sample means.Tukey'stest was chosen over other multiple




6/16

2002] PERRY:STARCHSIZEIN MANIOCAND SWEETPOTATO 339

TABLE 1. MEAN WIDTH/LENGTHRATIO OF MANIOC STARCHGRANULESWITHTUKEYGROUPINGS.Sample Locale Date Mean Group

Mperu2 Northcoast Peru 0.93905 AMperul North coast Peru Modern 0.93596 AM.e.A.21 ChimuSur A.D. 1470-1523 0.93216 AM.e.A.7 ChimuSur A.D. 1840 0.92439 A BMYF8731 UpperRio Negro Modern 0.92033 A BM.e.A.ll PampaRosario 810-450 B.C. 0.91733 A BMYF8498 UpperRio Negro Modern 0.91626 A BComm.TS Thailand Modern 0.91127 A BM.e.A.13 ChimuSur A.D. 1470-1523 0.91033 A BM.e.A.22 Chimu Sur A.D. 1470-1523 0.90822 A BM.e.A.15 Chimu Sur A.D. 1470-1523 0.90791 A BM.e.A.19 ChimuSur A.D. 1470-1523 0.90276 A BMYF8360 UpperRio Negro Modern 0.90131 A BM.e.A.23 Chimu Sur A.D. 1470-1523 0.90023 A BM.e.A.18 Chimu Sur A.D. 1470-1523 0.89832 A BM.e.A.12 Pampa Rosario 810-450 B.C. 0.89578 A BM.e.A.10 Pampa Rosario 810-450 B.C. 0.89554 A BM.e.A.14 Chimu Sur A.D. 1470-1523 0.89244 A BM.e.A.20 ChimuSur A.D. 1470-1523 0.89023 A BM.e.A.4 Las Haldas 1040-895 B.C. 0.88585 A BM.e.A.16 Chimu Sur A.D. 1470-1523 0.88578 A BM.e.A.8 Chimu Sur A.D. 1840 0.88577 A BM.e.A.5 ChimuSur A.D. 1840 0.88100 A BPerul Chancaycemetery A.D. 1400 0.88024 A BM.e.flabell Brazil Modern 0.87960 A BM.e.A.6 Chimu Sur A.D. 1840 0.87917 A BM.e.A.17 ChimuSur A.D. 1470-1523 0.87480 A BM.e.A.l Pampade las Llamas 1785-1120 B.C. 0.85646 B

comparisontests (e.g., Least SignificantDiffer-ence, Student Newman Keuls), due to its ten-dency to find fewer significantdifferencesthanthe otherprocedures.The conservativenatureofTukey's test makes it much less likely that theresearcherwill falsely rejectthe null hypothesisthat all sample means are equal (Type I Error)(Toothaker1993).

RESULTSThe mostmodemsamples of bothmanioc andsweet potato, in reference to both length andwidthmeasurements, ieldedsome of the largeststandarddeviationsin the entire data set (Tables2, 3, 5, 6). The sampleof wild type manioc, incomparison o most of the moremodemmanioc

samples, had much more uniform length andwidth measurements Tables2, 3). The overalltrend, that is more clearly defined in the sweetpotato data set (Tables 5, 6) is an increase insamplestandarddeviation as starchgranulesizeincreases.If the data from the wild manioc spec-

imen do not representa samplinganomaly, itcan be saidthatvariabilityn bothstarchgranulelength and width appears to increase after do-mesticationin this particularspecies, with thesingle exceptionof the moderncoastal Peruvianspecimens.Date and starchgranule size was not signifi-cantly correlated.Some of the earliest samplesof manioc yielded several of the largest starchgranules n the entiresample (Tables 1, 2). Be-cause regressionanalysesshowed no significantcorrelationbetween date and increasein starchgranule size in either species, a multiple com-parisontest was employedto assess whether ornot the starchgranules derived from a modernsweet potatodiffered from archaeologicalspec-imens, and if starch granulemeasurementsde-rived from "domesticated"samples of maniocdifferedsignificantly rom those of thepurportedwild ancestor, M. esculenta, subsp. flabellifolia.Beginningwith the manioc data(Tables 1-3),thecomparisonsbetween the means of thestarch




7/16

340 ECONOMIC BOTANY [VOL. 56

TABLE 2. MEAN WIDTH OF MANIOCSTARCHGRANULESWITHTUKEY GROUPINGS.Sample Site Date Mean S.D. Tukey grouping

MYF8731 Modern 14.9420 4.63 AMYF8360 Upper Rio Negro Modern 14.3320 4.36 B AM.e.A.5 Chimu Sur A.D. 1840 14.0122 5.78 B A CM.e.A.l Pampa de las Llamas 1785-1120 B.C. 13.4906 5.30 B D A CPerul Chancay cemetery A.D. 1400 13.0478 2.32 E B D A CMYF8498 Upper Rio Negro Modern 12.9052 3.30 E B D A C FMperu2 Peruvian north coast Modern 12.772 3.02 E B D A G C FMperul Peruvian north coast Modern 12.64 2.57 E B D H A G C FM.e.A.4 Las Halas 1040-895 B.C. 12.3689 3.90 E B D H A G C FM.e.A.18 Chimu Sur A.D. 1470-1523 12.3049 4.15 E B D H A G C FM.e.A.7 Chimu Sur A.D. 1840 12.2164 5.19 E B D H A G C FM.e.A.19 Chimu Sur A.D. 1470-1523 12.1229 4.34 E B D H G C FM.e.A.6 Chimu Sur A.D. 1840 11.5030 4.19 E D H I G C FM.e.A.16 Chimu Sur A.D. 1470-1523 11.1782 3.70 E D H I G FM.e.A.8 Chimu Sur A.D. 1840 11.0110 4.60 E J D H I G FComm.TS Thailand Modern 10.9667 3.12 E J D H I G FM.e.A.ll Pampa Rosario 810-450 B.C. 10.9224 3.58 E J D H I G FM.e.A.23 Chimu Sur A.D. 1470-1523 10.7994 3.03 E J D H I G FM.e.A.21 Chimu Sur A.D. 1470-1523 10.6026 2.44 E J H I G FM.e.A.20 Chimu Sur A.D. 1470-1523 10.5288 3.52 E J H I G FM.e.A.17 Chimu Sur A.D. 1470-1523 10.4009 3.52 E J H I G FM.e.A.22 Chimu Sur A.D. 1470-1523 10.3271 3.43 E J H I G FM.e.A.14 Chimu Sur A.D. 1470-1523 10.2582 3.73 E J H I G FM.e.A.15 Chimu Sur A.D. 1470-1523 10.2188 3.12 J H I G FM.e.A.10 Pampa Rosario 810-450 B.C. 9.9876 2.30 J H I GM.e.A.13 Chimu Sur A.D. 1470-1523 9.9630 4.22 J H IM.e.A.12 Pampa Rosario 810-450 B.C. 8.7428 2.81 J IM.e.flabell Brazil Modern 8.3345 2.20 J

granulewidth/lengthratiosresultedin, perhaps,the least complexgroupingsof means(Table1).Tukey test outputsgroup values such that meanssharingthe same letter do not differ significant-ly. Differences are consideredto be statisticallysignificantif P values are less than0.05. Withan F value of 2.03 and P < 0.0014, the meansfrom the two modemPeruvian pecimens(MPe-ru1, 2) and archaeological specimen 21(M.e.a.21) group out and differ highly signifi-cantly from archaeologicalsample 1 (M.e.a.1).Width and length calculations for maniocstarchgranulesresulted in rathercomplex andoverlappinggroups of means (Tables2, 3). Forwidthmeasurements,F = 9.24 andP < 0.0001,while for lengthmeasurements,F = 10.89 and,again,P < 0.0001. Therefore,groups of meansthatare scored with differentletters differ witha very high level of significance.For both length and widthmeasurements, hearchaeological amplesareinterspersed etweenoverlappinggroupsthat do not correlate n any

way with the date of the specimen(Tables2, 3).Theredo, however,appearto be some patternsthat standout with the modern specimens. ThePeruvianmanioc root with the smallest samplemean for both width and length measurementsamongmodernsamples(Mperul) differssignif-icantly in widthfrom archaeologicalsample 12and from the wild manioc (M.e.fla.),while inlength, the same specimen differs significantlyfrom archaeologicalsamples 12 and 13 as wellas the wild formflabellifolia. Modern Peruvianspecimen 2 (Mperu2)differs significantlyfromarchaeologicalsamples 12 and 13 as well aswild maniocin both lengthand width.The three modern samples from the UpperRio Negro, like those fromPeru,also group outin "A" with the largest overall means.MYF8498 differssignificantly romarchaeolog-ical specimens 10, 12, and 13 and wild maniocin both width and length, and also differs sig-nificantly from archaeologicalspecimen 15 inlength. MYF8360 differs significantlyin both




8/16

2002] PERRY:STARCHSIZE IN MANIOCAND SWEETPOTATO 341

TABLE3. MEANLENGTH F MANIOC TARCH RANULES ITH uKEYGROUPINGS.Sample Site Date Mean S.D. Tukeygrouping

MYF8731 UpperRio Negro Modern 16.1081 4.52 AMYF8360 UpperRio Negro Modern 15.9260 4.71 B AM.e.A.5 Chimu Sur A.D. 1840 15.7243 6.06 B A CM.e.A.l Pampade las Llamas 1785-1120 B.C. 15.6308 5.36 B A CPerul Chancaycemetery A.D. 1400 15.0109 3.13 B D A CMYF8498 UpperRio Negro Modern 14.0761 3.47 E B D A CM.e.A.4 Las Haldas 1040-895 B.C. 13.9039 4.11 E B D A C FM.e.A.18 Chimu Sur A.D. 1470-1523 13.6087 4.18 E B D A G C FMperu2 North coast Peru Modern 13.6087 3.06 E B D A G C FMperul North coast Peru Modern 13.5300 2.59 E B D A G C FM.e.A.19 Chimu Sur A.D. 1470-1523 13.2446 4.01 E B D G C FM.e.A.7 Chimu Sur A.D. 1840 13.2397 5.52 E B D G C FM.e.A.6 Chimu Sur A.D. 1840 12.9740 4.22 E D G C FM.e.A.16 Chimu Sur A.D. 1470-1523 12.5263 3.60 E D H G FM.e.A.8 Chimu Sur A.D. 1840 12.3344 4.57 E I D H G FM.e.A.23 Chimu Sur A.D. 1470-1523 12.0343 3.18 E I H G FComm.TS Thailand Modern 11.9654 3.11 E I H G FM.e.A.17 Chimu Sur A.D. 1470-1523 11.8867 3.73 E I H G FM.e.A.11 PampaRosario 810-450 B.C. 11.8228 3.51 E I H G FM.e.A.20 Chimu Sur A.D. 1470-1523 11.8031 3.60 E I H G FM.e.A.14 Chimu Sur A.D. 1470-1523 11.4636 3.89 E I H G FM.e.A.22 Chimu Sur A.D. 1470-1523 11.3160 3.36 E I H G FM.e.A.21 Chimu Sur A.D. 1470-1523 11.2914 1.98 E I H G FM.e.A.15 Chimu Sur A.D. 1470-1523 11.1782 2.92 I H G FM.e.A.10 PampaRosario 810-450 B.C. 11.1192 2.10 I H G FM.e.A.13 Chimu Sur A.D. 1470-1523 10.8043 4.13 I H GM.e.A.12 PampaRosario 810-450 B.C. 9.7170 2.77 I HM.e.flabell Brazil Modern 9.6186 2.77 I

TABLE 4. MEAN WIDTH/LENGTHRATIO OF SWEETPOTATOSTARCHGRANULESWITHTUKEYGROUPING.Sample Site Date Mean Tukeygrouping

I.b.A.l Chimu Sur A.D. 1470-1523 0.91252 AI.b.A.7 Chimu Sur A.D. 1470-1523 0.97092 A BI.b.A.6 Chimu Sur A.D. 1470-1523 0.89640 A BI.b.A.3 Chimu Sur A.D. 1470-1523 0.89608 A BI.b.A.19 Chimu Sur A.D. 1840 0.89264 A BI.b.A.13 Pampade las Llamas 1785-1120 B.C. 0.88694 A BI.b.A.10 Chimu Sur A.D. 1470-1523 0.88236 A BI.b.A.2 Chimu Sur A.D. 1470-1523 0.87773 A BI.b.A.9 Chimu Sur A.D. 1470-1523 0.87523 A BI.b.A.8 Chimu Sur A.D. 1470-1523 0.87390 A BI.b.A.18 ChimuSur A.D. 1470-1523 0.87201 A BI.b.A.15 Chimu Sur A.D. 1470-1523 0.86973 A BI.b.A.4 Chimu Sur A.D. 1470-1523 0.86321 A BI.b.A.12 Pampade las Llamas 1785-1120 B.C. 0.85563 A BI.b.A.Il Pampade las Llamas 1785-1120 B.C. 0.85200 A BI.b.A.16 ChimuSur A.D. 1470-1523 0.85117 A BMonahl CarbondaleMarket Modern 0.84847 A BI.b.A.17 Chimu Sur A.D. 1840 0.84615 A BI.b.A.14 ChimuSur A.D. 1470-1523 0.83939 B




9/16

342 ECONOMIC BOTANY [VOL. 56

TABLE 5. MEAN WIDTH OF SWEETPOTATOSTARCHGRANULESWITHTuKEY GROUPINGS.Sample Site Date Mean S.D. Tukey grouping

Monah I Carbondale Market Modem 16.9051 5.57 A1.b.A.9 Chimu Sur A.D. 1470-1523 16.5312 4.91 B AI.b.A.l Chimu Sur A.D. 1470-1523 15.9949 5.4 B A Cl.b.A.14 Chimu Sur A.D. 1470-1523 15.9408 5.98 B A CI.b.A.l Pampa de las Llamas 1785-1120 B.C. 15.3258 4.29 B D A CI.b.A.18 Chimu Sur A.D. 1470-1523 14.6173 4.72 E B D A CI.b.A.12 Pampa de las Llamas 1785-1120 B.C. 14.4451 4.09 E B D A CI.b.A.7 Chimu Sur A.D. 1470-1523 14.1450 4.19 E B D A C FI.b.A.10 Chimu Sur A.D. 1470-1523 14.0663 4.36 E B D A C FI.b.A.3 Chimu Sur A.D. 1470-1523 13.7022 3.98 E B D G C FI.b.A.6 Chimu Sur A.D. 1470-1523 13.0823 3.85 E D G C FI.b.A.8 Chimu Sur A.D. 1470-1523 12.9986 3.45 E D G FI.b.A.13 Pampa de las Llamas 1785-1120 B.C. 12.8314 3.60 E D G FI.b.A.19 Chimu Sur A.D. 1840 12.7871 3.09 E D G F1.b.A.4 Chimu Sur A.D. 1470-1523 12.5017 3.50 E D G FI.b.A.2 Chimu Sur A.D. 1470-1523 12.2262 3.62 E G FI.b.A.16 Chimu Sur A.D. 1470-1523 11.3160 3.26 G FI.b.A.15 Chimu Sur A.D. 1470-1523 11.2570 2.93 G FI.b.A.17 Chimu Sur A.D. 1840 10.9568 2.45 G

width and length from fourteen archaeologicalspecimens,the modem commercialgrademan-ioc starch sample (Comm.TS.) and from wildmanioc. MYF8731, the samplewith the largestmeans,has lengthandwidth measurements hatdiffer significantly from those of commercialmanioc starchand wild manioc,while the widthdiffers significantlyfrom fifteen archaeologicalspecimens and the length differs significantlyfrom sixteen archaeologicalspecimens.The data from the sweet potato specimensfallout in a similar pattern to that of the maniocsamples. With an F value of 2.23 and P


10/16

2002] PERRY: STARCH SIZE IN MANIOC AND SWEET POTATO 343

TABLE 6. MEAN LENGTHOF SWEETPOTATOSTARCHGRANULESWITHTuKEY GROUPINGS.Sample Site Date Mean S.D. Tukey grouping

Monahl Carbondale Market Modem 20.2015 7.03 AI.b.A.9 Chimu Sur A.D. 1470-1523 19.0798 6.02 B AI.b.A.14 Chimu Sur A.D. 1470-1523 18.8830 6.13 B AI.b.A. Il Pampa de las Llamas 1785-1120 B.C. 17.8596 3.88 B A CI.b.A.l Chimu Sur A.D. 1470-1530 17.4808 5.43 B D A CI.b.A.12 Pampa de las Llamas 1785-1120 B.C. 16.8559 4.28 B D E CI.b.A.18 Chimu Sur A.D. 1470-1523 16.6985 4.92 B D E CI.b.A.10 Chimu Sur A.D. 1470-1523 15.9654 4.68 F B D E CI.b.A.7 Chimu Sur A.D. 1470-1523 15.6850 4.70 F D E CI.b.A.3 Chimu Sur A.D. 1470-1523 15.3110 4.30 F D E CI.b.A.8 Chimu Sur A.D. 1470-1523 14.9371 3.83 F D E CI.b.A.4 Chimu Sur A.D. 1470-1523 14.6419 4.26 F D EI.b.A.6 Chimu Sur A.D. 1470-1523 14.6173 4.25 F D EI.b.A.13 Pampa de las Llamas 1785-1120 B.C. 14.4648 3.69 F D EI.b.A.19 Chimu Sur A.D. 1840 14.2286 2.75 F EI.b.A.2 Chimu Sur A.D. 1470-1523 13.9679 3.80 F EI.b.A.16 Chimu Sur A.D. 1470-1523 13.3086 3.61 FI.b.A.15 Chimu Sur A.D. 1470-1523 13.0626 3.46 FI.b.A.17 Chimu Sur A.D. 1840 13.0036 2.41 F

two samplesfallingwithingroup"A," thatwiththe largest means.DISCUSSIONSTARCHGRANULEVARIABILITY

For the most part, none of the hypotheses re-garding starch granule size, variability, and dateof deposition were supported by the data. Ingeneral, in both manioc and sweet potato spec-imens, variability in starch granule measure-ments correlatedpositively with an increase insize. It would appear that roots that producelargestarchgranulesalso producesmall ones aswell. Thus, an overall increase in size is alsoaccompaniedby a greaterrangeof sizes of sin-gle granules, or, perhaps, retentionof all sizeclasses of granules. There appears to be no clearrelationship between date of deposition and var-iabilityof starchgranulesize. The single excep-tion is among the data from coastal Peruvianspecimens of manioconly.The two modem Peruvianmanioc specimensyielded less variable measurements of starchgranules han those fromthe vastmajorityof thearchaeological specimens from the Peruviancoast, and, therefore, these data do support thehypothesis that variability in starch granule sizewill decrease over time (Tables 2, 3). However,this trend is confounded by the fact that the var-iability of the starch granules from the modem

Peruvianspecimensis aboutequalto thatof thewild type manioc. The relationshipbetweenthelowland, coastal Peruvian, and wild maniocsampleswill be discussed furtherbelow.

WIDTH/LENGTH RATIOSThe results of the multiplecomparisontestscompletedon the width/length atios of boththemanioc andsweet potatostarchgranulesarenotentirelyunexpected.Only one to threesamplesdiffered significantlyfrom one another n bothsamplepopulations.Thisresultcan be explainedby the basic morphologicalcharacteristicsand

tendencies of plant starches.No matter the overall size of a manioc orsweet potatostarchgrain,the basic morphologyof the granules tends to be the same. Thus, theratio between the maximumlength and maxi-mum width of any given starchgranule shouldnot differ greatly from that of anothersampledfrom the samespeciesof plant.Further,he mostsignificantdifferenceamong the maniocsamplesamountsto eight one-hundredths f a micron,while in the sweet potato samples, the greatestdifference in means is seven one-hundredths fa micron (Tables 3, 6). Although statisticallyspeaking, these values differ significantly. alength this small is not detectable by the re-searcher working with a light microscope.Therefore,this difference may be the result of




11/16


the researcher's estimation of the "tenth" divi-sion between micrometer lines.

STARCHGRANULESIZEThe initial hypothesis, that starch granule sizewould increase from early to later samples of

cultivated manioc and sweet potato, is onlysomewhat supported by these data. Some of thelargest mean values occur in samples from theearliest sites. It cannot be ascertained whether ornot the results of this study are clouded by sam-pling bias, and if the addition of further sampleswould place the earliest samples as anomalousoutliers within the groups with the largest meansize. This author, however, thinks that is notlikely the case. Although a gross generalizationcan be made that later samples of both maniocand sweet potato have larger starch granulesthan earlier ones, the data reveal a rather com-plex relationship between starch granule sizeand time. Because the only significant relation-ships appear between manioc and its wild an-cestor, the sweet potato data will not be dis-cussed further.

A complicating factor in the manioc data setis the placement of the modern commercialgrade manioc starch in Tukey groupings far fromthe other modern samples, and in a group whosemeasurements do not differ significantly fromthose of the wild manioc. This sample was theonly one that was not collected directly by thisauthor from an actual root, and it is likely thatthis collection bias is responsible for the locationof this sample's means in the Tukey groupings.

Most commercially prepared starch is sievedthrough fine screens to remove debris and otherunwanted material and to increase uniformity ofstarch granule size. This process results in amore desirable and palatable starch product. Itis likely that such a process skewed the sampleby removing large granules, and, therefore, it isdifficult to assess whether or not this sampleshould be included in the analyses at all. If thissample is removed from the data set, only oneconclusion can be drawn with certainty: modernsamples of manioc from both lowland and coast-al Peruvian samples yield significantly largerstarch granules than those of its supposed wildancestor. Several factors could account for thesedifferences.

Root size was considered as a possible causalfactor. Larger starch granules could be correlatedwith larger roots. The modern samples in group

"A," thatwith the largestmeans, (MYF 8731,8360, and8498, Mperu1 and2) have maximumroot diameters ranging from about four centi-meters to eight centimeters. Archaeologicalspecimensin the same group (M.e.a.5, 1, 4, 18,7, 19, Perul) have maximum root diametersofabout one centimeter.Further, oots of the samesize as the archaeological amplesin this groupdo not differsignificantly rom the wild manioc(e.g., M.e.a.10-15). The wild sample itself ap-pears to have had a diameterof between one andtwo centimeters.Therefore, t wouldappear hatroot size is not correlated with starchgranulesize.The second possibilityto consideris soil fer-tility. Notably, the manioc roots yielding thelargeststarchgranuleswere collected in swiddenforest plots in the Upper Rio Negro, a black-water river system where soil fertility is quitelow. While the conditions under which the ar-chaeological samples were cultivated are notknown, one would expect that Peruvian soilswould be more fertile than those of the UpperRio Negro due to the sediment-richdrainagefrom the Andes. And, yet, some of the archae-ological coastal Peruviansamples do not havemeans that differ significantly rom those of thewild manioc.Thus,it wouldappear hat soil fer-tility is not a causal factor in determining hesize of starchgranules n these samples.A host of other factorscannot be consideredhere because they are not known. Effectivemoisture,cultivationpractices,cultivarspecifics,andthe amountof time that a root was allowedto remain n the groundaccumulating tarchareall unknowns.Whatis clear is thatstarchgran-ule size appears o be the result of some factoror factors that were not accountedfor in thisstudy. If, in fact,manioccultivationbegan at theonset of the Holocene, perhaps t has takenuntilmoderntimes for the starchgranules o increasesignificantlyenough in size over those of thewild ancestor for a mathematicaldifferencetobe detected.THE PERUVIANCOAST AND THELOWLANDS

To complicatemattersfurther,while collect-ing measurements f the starchgranules nclud-ed in this study,the authornoticeddiscrepanciesbetweenher observationsandthose of other re-searchers.Piperno and Holst (1998) noted thatboth modern and archaeological tarchgranulesfrom Panamanianmanioc roots were distin-




12/16

2002] PERRY:STARCH SIZE IN MANIOCAND SWEETPOTATO 345

20Oum

Fig. 1. Light micrographs f coastalPeruvian A, B) and lowland(C, D) types of maniocstarchgranulesshowing distinctiveanddiagnosticfeaturesof each. All photographs re at the same magnification.A. Starchgranulesof manioc roots from Chimu Sur. Note the typicalhemispherical orms with linear centralfissuring.B. CoastalPeruvianstarchgranulesfrom modernmanioc roots with typical Peruvian orm. Note the smalleroverall size andless "inflated"qualityof hemispherical ranulescompared o lowlandtypes.C. Modernmanioc starchgranules rom roots from the UpperRio Negro regionof Venezuela.Note the large,centralstellatefissure and hollow hilum.D. Compound tarchgrainof lowlandmaniocshowinghow basal facetingoccursduringdevelopmentas smallgranules orm with the largergranule.Note the inflatedqualityof the large,diagnosticgranule.

guishable from those of other species due to alarge, diagnostic hemispherical granule withbasal faceting and a central stellate fissure. Noneof the 39 archaeological samples or two modemsamples from Peru (in the author's comparativecollection) yielded starch granules with this dis-tinctive morphology. Although hemisphericalforms with basal faceting were common occur-rences, and some samples yielded starch gran-ules with small central fissures, no stellate fis-sures could be located (Fig. 1A, B). In contrast,all 31 modem manioc starch samples from the

Upper Rio Negro yielded starch granules iden-tical to those described from Panama (Fig. IC,D).

The author considered that the differencesmight have been due to the drying of the coastalPeruvian roots in the arid Peruvian environment.However, when the archaeological starch gran-ules were soaked in deionized water for twoweeks, no changes in morphology occurred. Fur-ther, the two modem samples from Peru yieldedstarch granules that exhibit the same morphol-ogy as those from the Peruvian archaeological




13/16


samples that date from 3000 years B.P., and amodernsample from coastal Ecuadorprovidedto the authorby Dr. DeborahPearsallalso yield-ed this distinctcoastalmorphology.When the samplesfrom the UpperRio Negrowere examined, starch granules of the type de-scribed by Pipernoand Holst (1998) were ob-served and documented. Notably, this starchgranule form was found in sweet varieties ofmanioc from Panama(D. Piperno pers. comm.2000), and in the bittervarietiesfrom the UpperRio Negro, so this distinction does not accountfor the differencein starchgranulemorphology.Further, tarchgranuleswith this distinctive ow-landmorphologyhave been recovered fromthe

Aguadulcerocksheltern Panama Pipernoet al.2000), and they are dated to about 8000 yearsB.P.Every lowland sample contained the largegranuleswith stellatefissures,and no coastal Pe-ruvian or Ecuadoransample yielded any suchgranules.These observations ead to the conclu-sion that there exist two distinct and diagnosticmorphologiesof starchgranules that are indic-ative of different lowland and coastal Peruviancultivars of manioc. These forms have signifi-cant time depth,8000 years in the lowlands and3000 years on the coast of Peru, and may berepresentative f processesthat occurredduringthemigrationof theseplants n prehistoricSouthAmerica.It is quite well establishedandwidely agreedupon that manioc was originallya lowland do-mesticate, and the root-producing plant wasbrought romthe lowlandsto the Andes at somelaterdate (see Pearsall1992). It is entirely pos-sible that the environmental onditionsdiffer sosignificantly between the neotropicallowlandsandthe coast thatphenotypic differences in thestarchgranulesof the plants cultivated in bothregions will differ.The phenotypicplasticity ofstarchgranules s not known, however, t shouldbe pointedout thatno differences n starchgran-ule morphologies have been found in maizesamplesfrom both regions.Another possible cause of this difference instarchgranulemorphology s selectivepressuresthatwere presentin the coastalPeruvianregionafter manioc arrived. The climatological andedaphic differences between the lowland andhighland regions of South America have beennotedfor quitesome time. It is entirelypossiblethat certainmaniocplantswere betteradapted o

the coastal Peruvianconditions,and these cul-tivarscontinuedto be plantedwhile othersdidnot. The starch granulemorphologiesvisible inboth the archaeological and modem samplesmay representa small selection of the morpho-logical diversitypresent n theoriginalmigrants.This same phenomenonwould be presenthadtheoriginalmigrantplantsbeena smallselectionof the entirepopulationof the lowland manioccultivars: a founder effect. If the movement ofmanioc from the lowlandsto the Andes occurredin a single event,andthe cultivar hatwas trans-portedin this event happenedto contain starchgranulesof this distinctivemorphology, henthedescendantsof this plantwould also yield thesediagnosticgranules.Another possible explanationfor the differ-ences in starchgranulemorphologiesmaybe in-dependentdomestication.It has been suggestedthat manioc was domesticated more than oncein variousregions of South America(Ugent etal. 1986). The most recent research on the do-mesticationof manioc has generatedbuzz in thepopularpress that modern cultivatedManihotesculenta is derivedfrom a single wild species,Manihot esculenta subsp. flabellifolia. However,the researchers hemselvesadopta more conser-vative view of theirfindings (Olsen and Schaal1999).Olsen and Schaal were seeking data thatwould assist in the understandingof both thegeographiclocation of the original domestica-tion of manioc,and whetheror not manioc wasderived from a single ancestorrather han fromcontinuoushybridizationwith wild species. Thedata were sought by comparingmoderndomes-ticated specimensto two wild populationsthatmay have been ancestral o the cultivated man-ioc. Genetic studies yielded data that suggest"[T]he crop seems to be derivedfrom popula-tions of subspecies labellifolia along the south-ern border of the Amazon Basin" and further,"subspeciesflabellifolia alone can account forthe genetic variationobserved in cassava" (Ol-sen and Schaal 1999:5591).Several issues are still not well understood,despite this recent work. First, the geographicorigin of the modem germplasmused in thestudy is not disclosed as it is by other authors(e.g., Fregeneet al. 1994). Thus, the readercan-not assess the breadthof samplingfor the mod-ern data. Given the data found in this author'sdissertationwork, it is unclearwhetheror not all




14/16

20021 PERRY:STARCHSIZE IN MANIOCAND SWEETPOTATO 347

moderncultivarsof manioc represent he sametaxon.Second, the authorsonly sampledfrom Bra-zilian populationsof the wild manioc(OlsenandSchaal 1999:5587). The authors indicate thatwild manioc occurs primarilyin west centralBrazil andeasternPeru(Olsenand Schaal 1999:5586). However, herbarium pecimens of Ma-nihot esculenta ssp. flabellifolia have been col-lected from as far northas centralFrenchGuiana(NYBG #s 1004000, 1004001-2, 1004008-9,1004011, 1004014), as far south as Paraguay(MOBOT D. Brunner 1244), and the originaltaxon was typed using materialcollected in theDominican Republic (Rogers and Fleming1973). Further, considering that manioc waslikely domesticatedat the onset of theHolocene,it is difficult to assess what the geographicdis-tributionof wild maniocmighthave been at thattime compared o modern times.Finally, it should not be forgottenthat, ex-ceptingcollections from the arid Peruviancoast,archaeological specimens of manioc are justnow beginning to accumulate,and an under-standingof the prehistoricdistributionand useof the plantis only beginningto emerge.Givenboth the demographicand environmental hang-es that occurred n South America aftercontact,it would be a leap to assume that modern vari-eties of cultivatedmanioceffectively andappro-priately represent he makeupof archaeologicalpopulations.Althoughthis study (Olsen and Schaal 1999)is certainly well conceived and carefully exe-cuted, it is quite geographicallyand temporallyconfined,and from this work alone, it would bedifficult to understandor trace widespreaddo-mestication romanarchaeological tandpoint. twouldcertainlybe useful to see geneticanalysescompletedon collectionsof archaeologicalman-ioc from different regions, particularlythoseyieldingstarchgranules hatdiffersufficiently nmorphology that this authorwould score themas differenttaxa. The data from this study sup-portthe multipledomesticationhypothesis (Ug-ent et al. 1986) rather han the more commonlycited single origin model.

CONCLUSIONSInitial hypotheses were formulated o inves-tigate whether domestication in manioc andsweetpotatocrops could be tracedusing the sizeand uniformityof starchgranules. Presumably,

starchgranule size would both increase andbe-come more uniformover time as cultivationanduse intensified.However,with very few excep-tions, dates in neithersweet potato nor maniocsamples were correlatedwith either starchgran-ule size or uniformity.A gross generalizationcan be made that modem specimens of bothmanioc and sweet potato yield larger starchgranules han some archaeological pecimens.Itdoes appear,however,that modernmanioc rootsproducesignificantlyargersized starchgranulesthan those of the wild typeflabellifolia.It should be noted that the authormade useof those manioc and sweetpotato specimensthatwere readilyavailable.That s to say, no attemptwas made to collect samples representativeofthe entire populationsof these cropsin highlandandlowland SouthAmericafromarchaeologicalor modem contexts. Thus, fewer modem sam-ples were examined than archaeological sam-ples, and all samples are geographicallyandtemporallyrestricted.Any or all of these sam-plingbiases may have affectedtheresultsof thisstudy.At this time it would appear hat starchgran-ule size may not be a useful tool for assessingthe date of depositionof a given archaeologicalmaniocor sweet potatoroot. An altemativehy-pothesisthatmaybe worthexploring s whetheror not starchgranulesize is correlatedwith in-tensityof cultivationor, perhaps,the degree ofrelianceof the populationuponthecropin ques-tion.The largest modern manioc starch granuleswere obtainedfromroots collected in the UpperRio Negro where the crop is the major staplecarbohydrate ource in the diet (see Hoffman1993). Pampade las Llamas, though one of theearliest sites from which manioc and sweet po-tato roots were sampled,yielded specimenswithvery large starchgranules(Tables 2, 3, 5, 6).Although the species composition of the ar-chaeobotanicalassemblage from this site doesnot differ from thatof the otherPeruviansites,the numbers of manioc roots recovered fromeach collection sample (29, 5, 29, 20, 2) werelarger han thoserecovered n excavationsat LasHaldas(1), PampaRosario(estimateof 10), andChimuSur (2, 12, 11, 4, 28, 5, 2). Perhaps heresidents of Pampa de las Llamaswere relyingmoreheavily upon manioc thanwere the inhab-itantsof the othersites,andthis trend s reflectedin the large starchgranulesof the manioc roots.




15/16


In contrast o the complex issues surroundingstarchgranulesize and intensityof cultivationor domestication,there exist two clear lines ofevidence suggest that the coastal and lowlandregionalvarieties of manioc differ from one an-other,and have been dissimilar for quite sometime. First, unlike the case with the lowlandmodern manioc starchgranulesin which vari-abilityincreasedwith starchgranulesize, in thePeruvianspecimens,the starchgranulesize var-iability of the modem manioc specimens waslower than most of the archaeologicalspeci-mens, and was about equal to that of the wildmanioc. Thus, there is a different patternofstarch granule formation in coastal Peruvianmanioc roots: when starchgranulesize increasesoverall, small size granulesdo not occur at ashigh a rate as in lowland roots.Second,there is the distinctive anddiagnosticmorphologyof each type. This distinction hasbeen notedin every sampleexaminedby the au-thor,and in specimens from Panamaniancon-texts (Piperno and Holst 1998). The distincttypes have at least 8000 years of time depthinthe lowlands,and at least 3000 years in Peru.It should be noted thatif the coastal and low-landtypes of manioc are, in fact, different axa,the hypotheses put forth in the introductionofthis chapterhave not been adequately estedforbothregions. First,it is unknown f Manihotes-culenta subsp. flabellifolia is the wild ancestorof both types. Based solely uponmorphologicalcharacteristics bservedby the author, t wouldappear hat labellifolia is more likely relatedtolowlandmanioc than to highlandmanioc. If thisscenarioprovesto be accurate, henonly modernsamplesof lowlandmanioc have beencomparedto a wild type, andonly archaeological amplesof coastal Peruvianmanioc have been comparedto the moderntype.Therefore,the only conclusions that can bedrawnare thatmodern owland maniocproducesmore variable but much larger starchgranulesthan its ancestor,and Peruvian manioc appearsto produce more uniform starch granules inmodernspecimens comparedto archaeologicalspecimens. This result is in contrastto the ar-chaeological Peruvian sweet potato specimensthat, in general, exhibit higher levels of vari-ability as starchgranule size increases. In thecase of the Peruvianmanioc,theredoes not ap-pear to be a clear correlationbetween the pas-sage of time and an increase in manioc starch

granule size. This is not the case in the lowlandmanioc, where the modem starchgranules aremuch largerthan those of the wild sample. Ofcourse,no archaeologicalowlandsampleswereincluded,so it is difficult to assess the accuracyof this claim for prehistoricmanioc.Clearly,ifthese issues are to be clarified,a good deal ofresearchstill lies ahead.Whether or not the differences between thedistinctive and diagnostic starch granule mor-phologies of the lowland and Peruvian ypes ofmanioc can be sufficientlyexplainedat this time,what is clear is that we have a very useful ar-chaeological markerfor manioc cultivars fromthese two regions. While starch granule sizeanalyses may or may not prove to be effectivetools for understandingdomestication,the dif-ference in manioc starch granule morphologymay be very useful in tracing prehispanicplantmigration and, therefore, may eventually assistin the comprehensiveunderstanding f the ori-gins of maniocin the Neotropics.

LITERATURE CITEDCrawford,G. 1992. Prehistoric lantdomesticationnEast Asia. Pages 7-38 in C. W. Cowan and P. J.

Watson,eds., The origins of agriculture: n inter-nationalperspective.Smithsonian nstitutionPress,Washington,D.C.Fregene,M. A., J. Vargas, J. Ikea, F. Angel, J. Toh-me, R. A. Asiedu, M. 0. Akoroda, and W. M.Roca. 1994. Variabilityof chloroplastDNA andnuclear ribosomal DNA in cassava (Manihot es-culenta Crantz)and its wild relatives.TheoreticalandAppliedGenetics89:719-727.Hoffman, S. D. 1993. Subsistence n transition:ndig-enous agriculture n Amazonas,Venezuela.Doc-toraldissertation,Universityof California,Berke-ley.Jones, V. H. 1936. The vegetalremainsof Newt KashHollow Shelter.Pages 147-167 in W. S. WebbandW. D. Funkhouser,ds., Rock shelters n MenifeeCounty,Kentucky.Universityof KentuckyReportsin ArchaeologyandAnthropology3.Matsuzawa, T. 1978. The formativesite of Las Hal-das, Peru:architecture,hronology,and economy.AmericanAntiquity43:652-672.McClung de Tapia, E. 1992. The originsof agricul-ture in Mesoamericaand CentralAmerica.Pages143-171 in C. W. Cowan and P. J. Watson, eds.,The origins of agriculture:an internationalper-spective.Miller, N. F. 1992. The origins of plantcultivation nthe Near East.Pages39-58 in C. W.Cowan andP.J. Watson,eds., The origins of agriculture: n in-ternational erspective.




16/16

2002] PERRY:STARCHSIZE IN MANIOCAND SWEETPOTATO 349Olsen, K. M., and B. A. Schaal. 1999. Evidence onthe originof cassava:Phylogeography f Manihotesculenta.Proceedingsof theNationalAcademyofScience 96:5586-5591.Pearsall, D. M. 1992. The originsof plantcultivationin South America. n C. W. Cowan and P.J.Watson,eds., The origins of agriculture:an intemationalperspective.SmithsonianInstitutionPress, Wash-ington,D.C.Piperno, D. R., and I. Holst. 1998. The Presence ofstarch grains on prehistoricstone tools from thehumid neotropics: indications of early tuber useandagriculturen Panama.Journalof Archaeolog-ical Science 25(8):765-776.Piperno, D. R., A. J. Ranere, I. Hoist, and P. Han-sell. 2000. Starchgrainsrevealearlyrootcrop hor-

ticulture n the Panamanian ropicalforest. Nature407:894-897.Rogers, D. J., and H. S. Fleming. 1973. A mono-graphof Manihotesculenta withan explanationofthe taximetricmethodsused.EconomicBotany 27:1-113.

Smith, B. D. 1995.Theemergenceof agriculture. ci-entificAmericanLibrary,New York.Toothaker, L. E. 1993. Multiplecomparisonproce-dures.Sage Publications,NewburyPark.Ugent, D., S. Pozorski, and T. Pozorski. 1981. Pre-historicremainsof the sweet potatofrom the Cas-ma Valley of Peru.Phytologia49(5):401-415.. 1982. Archaeologicalpotato tuber remainsfrom the CasmaValley of Peru. EconomicBotany36(2):182-192.. 1986. Archaeologicalmanioc (Manihot) romcoastalPeru.Economic Botany40(l):78-102., and M. Verdun. 1983. Starchgrainsof thewild and cultivatedMexican species of Solanum,SubsectionPotatoe.Phytologia53(5):351-363.Yarnell, R. A. 1972. Iva annuavar.macrocarpa:Ex-tinctAmericancultigen?AmericanAnthropologist74:335-341.. 1978. Domesticationof sunflowerandsump-weed in easternNorth America.Pages 285-299 inR. Ford, ed., Thenatureand statusof ethnobotany.AnthropologicalPapers No. 67, Museum of An-thropology,Universityof Michigan,Ann Arbor.

Pery Domesticacion de La Yuca y La Batata

Documents

Transcript of Pery Domesticacion de La Yuca y La Batata