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SMALL MILLETS IN GLOBAL AGRICULTURE Proceedings of the First International Small Millets Workshop Banoalore. Ind~a. October 29-November 2. 1986 Editors: A. Seethorarn K. W. Riley G. Harinorayana OXFORD & IBH PUBLISHING CO. PVT. LTD. New Delhi hbav Calcutta

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Page 1: SMALL MILLETS IN GLOBAL AGRICULTURE - ICRISAToar.icrisat.org/4468/1/CP_417.pdf · 2011. 11. 29. · MacNeish and Gall~nat. 1967) Two mtnor cereals, mango (Bromus mango Desv ) in Chile

S M A L L MILLETS IN

GLOBAL AGRICULTURE

Proceedings of the First International Small Millets Workshop

Banoalore. Ind~a. October 29-November 2 . 1986

Editors:

A . Seethorarn K. W. Riley G . Harinorayana

OXFORD & IBH PUBLISHING CO. PVT. LTD. New Delhi h b a v Calcutta

Page 2: SMALL MILLETS IN GLOBAL AGRICULTURE - ICRISAToar.icrisat.org/4468/1/CP_417.pdf · 2011. 11. 29. · MacNeish and Gall~nat. 1967) Two mtnor cereals, mango (Bromus mango Desv ) in Chile

ORIGIN, EVOLUTION A N D SYSTEMATICS O F MINOR C E R E A L S

J M J de Wet

The Poaceae Includes an est~mated 8.000 species belony~ny to sonie 6 0 yenvra Grasses a c u r on all continents. and In all hab~tats that support yrowth of llower~ny plants They serve man In many ways, but It IS the~r use as cereals and feed for livestock that make them essent~al for human surv~val The caryopses of most grasses are ed~ble, and at least 3oU species were harvested dur~ng h~storical tlmes as w~ld cereals by nomadic hunters and I~erders, and by farmers dur~ny times of scarcity Thirty f~ve specles belonylny to 20 yenera are known to have been domest~cated The~r cultivated races rely on man lor seed d~spersal (sawny), and for a su~table hab~tat to reproduce successfully (cult~vated f~eld)

Cereals are globally planted on an estimated 730 rn~lliorr hectares, and yield an est~mated 1.800 m~ll~on metrlc tons of yratn annually Wheat, maue and rlce account for approximately 80 per cent of yratn produced In the world These cereals are followed In Importance by barley, soryhum, oats, rye and pearl milkt w h ~ h towther represent another 19 per cent of the world's cereal p roduc~m The r e m n i n g cereak account for about 1 per cent of the foodgrain produced In the world today These minor cereals are not important In terms of wurld food production, but essential as food crops in their respective ayro- ecosystems They are mostly grown m marginal areas, or under agricultural condit~ons where rnapr cereals fail to consistently produce an acceptable harvest

MINOR CEREALS OF THE AMERICAS

WIW cereals played an important role In the d ~ e t s of nalave Americans until recent hlstwcal tlrnes (Palmer, 1871. Ball. 1984) Fifty species were entensively

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harwsted, but only s u specks were domestkated as cereals Maize (Zea mays L I is the only New World cereal of commeroal sgntfcance Setorfa gcn~culato (Lam ) P &auv Mrtk arassl from arld Mextco (Callen. 19651. and Phororns ~ a r d n h n a Wak (may griss) of the southeastern United States (chomko and Crawford, 1978) are known as cultivated cereals only tn an archaeological context &Wk grass was grown for at least a millennium (Callen, 1967). but was replaced by maize as a cereal some 4000 years ago (Mangelsdorph. MacNeish and Gall~nat. 1967) Two mtnor cereals, mango (Bromus mango Desv ) in Chile and sauwi (Panicurn sonorum Beal) in ~ e i ~ c o were important crovs until recent historical times American unld nce (Z~zonio ocauoltca L ) is arecent domesttcate In north central Canada and adjacent regtbns of the USA (de Wet and Oelke. 1979)

Molina (17821 recorded that the Araucano Indians In Central Chile grew a k n d of rye that was called el Mango, and a kind of barley that was called

ruca Tucs probably refers to Bromus un~olo~des HBK . a wild grass that 1 extennvely harvested as a cereal on the highlands of South America (Ball.

I&) El Mango (Bromus mongo) was culttvated Gay (1865) recorded that it was a btennial mop, grown with peppers and beans Florets were roasted to fxilitate removal of the lemma and palea, and the grains were ground Into flour to make bread or a fermented drink called chicha Cruz (1972) cited an unpubl~shed manuscript by Arturo Fontec~lla Larrain, a professor of agronomy at the Ca tho l~ University of Santiago in Ch~le during the early twentieth century. who recorded that plants of el Mango produced 40-50 culms, each bearing an Inflorescence with 70.100 grains It requtred etghteen months to mature. however, and el Mango was replaced during the last half of the twent~eth century by wheat for making flour and by apples to make c~der (Brucher. 1979)

Sauw (Panicurn sonorurn) is native to arid western North Amer~ca It formed an tmportant part of the Sonoran desert agriculture of northwestern Mexko (Nabhan and de Wet. 1984) It was wdely grown well tnto the twenneth century by Indian tribes who I~ved along the Colorado rtver delta (GiMord. 1931) Today it 6 groum only in southeastern Sonora and adpcent Chihuahua Although lmle known outs~de this qea, sauwi 1s a promlstng cereal for the semi arid tropics of Africa and Asta It is drought tolerant, and has acceptable y~eld

ential under adverse condtt~ons Plants produce several t~llers, each of which duce an Inflorescence with as many as 2.500 fertile florets The only New World minor cereal of present.day economic Importance

is Arneihan wild ihe Thts is not a true rice It belongs i o the genus Zlzonlo rather than Olyza Amerkan nce is commercially harvested as a wild cereal (Dote, 1969). and since the late 1960's planted on a commercial scale Zizan~o acquaka is the only grass species succersfuliy domest~cated as a cereal in hhtorlcal times (do Wet and Oelke. 1979) It is g o w n In paddies as is rice paddies are fboded and seeded in late fall Germination 1s rapid in spring and the water kvel is maintained until August when the mop matures Fields are

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then dra~ned and harvested w~ th a mod~f~ed rce comb~ne Since the cultlgen retalnssome degree of natural seed dnpersal. no subsequent sowlng is needed Sp~kekts and straw are w r k e d Into the wet so11 alter harvest, where the caryopses be dormant unt~l the soil thaws In the next spnng Natural populations rarely ywld more than 100 kg ha ', wh~le y~elds of l(XK) kg ha- 1 are obtained 1 8 1 planted paddws Yields of at least .W00 kg ha- 1 are posslble from fully non-shatter~ny culttvars

MINOR CEREALS OF AFRICA

At least MI grass swcles were exlens~vely harvested 1x1 Africa as wild cereals unt~l recent h~stor~cal tlnies (Uussvn. 1965. Jard~n, 1967) The widely dlstnbuted Brochiana dejlexa (Schumach ) C E Hubbard. Oryza barthri A Chev . and Paspolunt scrob~cula~urn L were extenstvely collected In West Afrra Sl~payrosl~r punyens (Desl 1 de W~nler, and Cenchrus brjlorus Roxb were harvested by nomad~c tr~bes In the Sahara These specles are often st111 encouraged as weeds in cult~vated f~elds where they are harvested as wild cereals

In West Afrra. Oryza bar1h11 Chev gave rlse under domest~cation to the culllvated 0 glaberr~t?~o Strudel (Porteres. 1976) Tuff, truyrosl!~ tej (Succ ) 11otter. 15 an Important cereal In the Ethiop~an h~yhlands and was probably dertved from E pllosu IL ) P Beauvo~s (de Wet. 1977. Costanza. de Wet attd Harlan. 1979) The weedy Brochiar~a dejlcxa (animal Ionlo) IS cultivated on the Fouta b a b n htyhhnds of the wesl Afrran savanna (Chevalier. 1933, I'ruteres. 1951) An~rnal fon~o differs froli? wlld l3 dejlexa only In havlny sptkelets that d~sarliculate tard~ly at rnaturtty

Two other minor cereals are ~mporlarit crops m the west A f r ~ a n savanna Black funlo. D~gllar~o ~burua Stapt IS grown by the Hausa tr~be of N~geria. and occurs sporadtcally across most of semi-and west Afr~ca (Porteres. 1955. Clayton. 1972) It 15 often planted between rows of sorghum or pearl millet. and commonly as a mtxture with Dtgjlar~a extlls (K!pplst) Stapf (true fon~o) Black Ionlo (D tburua) IS drought tolerant and often y~elds a harvebt when the mapr cereal 11 eccompanles fails to survtve True Ionlo (D exilir) is widely yrown across t l ~ e west Afrran savanna It d~f l rrs from D ~burua which has both glumes conspruously shorter than the sp~kqkt, in having the upper glume at least as long as the sptkelet Fonios are sown In west Afr~ca during May or June. and harvested ~n Seplemkr Harvested lnllorescences need l o be protected from molsture slnce the grams become agglutinated to the lemma and pdea when they yet wet lhreshed yrains are parched or dried ~n the sun before (he chaff IS removed by pound~ng In a wooden mortar Fon~o a used In slews. or the b o k d grams are eaten as rlce w~ th butter or palm 011

The most ~mporlant minor cereal in Afrra IS finger millet. Ekusine coracana (L ) Gaertner It was domest~cated In Africa, but IS also wdely grown in south

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22 Sml Millet.

Asla, partlcularly lnd~a (Hllu and de Wet. 1976a) The closest wild relative o f fingor milkt IS E coracono subsp ofrtcona (Kennedy.O'Byrne) Hllu and de Wet Spontaneous f~nger millet IS widely distr~buted along the eastern and southern highlands of Af r ra (Phill~ps. 1972) Derivatives of hybr~ds between cultvvated and wild taxa are companion weeds of finger mtllet across most of ~ t s d~strlbunon in Afnca

Finger mlllet first occurs In the archaeolog~cal record of early African agrncull&e dat~ny back some 3.oOU years lHllu ;nd de Wet. 197bb): and was lnlroduced ~ n l o lnd~a at least .?.(KK) wars sac, fV~shnu.M~ttrt.. I9flH) C~llt~vatvd > - flnger millet is extensively var~able, and this variation IS recagnireJ as five races by de Wet. Prasada Rao. Brink and Mengesha (1984) Race coracana IS widely dlstrlbuted across the range of flnger millet cultivation in Afrlca and Asla I t resembles w l d flnger mlllet In havlng flve to nlneteen slender ~nflorescence branches that are 6-11 cm long, d~y~tately arranged, and w~th the tips often becoming sluhtly Incurved or reflexed at tlme of maturlty Some genotypes differ phenotyprally from subsp ofrrano primar~ly In belny unable to disperse their spikelets without the help of man Race coracana IS partlcularly well adapted to agrrulture In the eastern h~ghbnds of Afrlca and the Ghats of lndla Some cultivars are drought tolerant and compete agyresrively w~th weeds under conditions of tradlhonal ayr~culture I t la often sowlr as a secondary crop wtth sorghum or pearl m~llet

Races vulgam, elongata, plana and cornpscta probably evolved from race coracana under cult~val~on They probably evolved In Africa and were introduced into lnd~a Little racial evolut~on took place in thts secondary centre of cultivat~on Race elonyala IS characterlzrd by slender ~nflorescence branches that are 10.24 cm lony I t IS grown in eastern Africa and the eastern Ghats of lnd~a Indian and African cultlvars cannot consistently be separated on the basts of lnflorescsnce morphology Race plana 1s prlniarily Afr~can In d~strlbutlon It is grown In Eth~op~a and Uganda, and to some extent In the eastern and western Ghats of India Race plana is character~zed by large. H 15 mm long sp~kelets that are arranged in two more or less regular rows along the rachis, giving the inflorescence branches a flat ribbon-l~ke appearance In some genolypes the ferille florets are x, numerous that they almost surround the rachis at matunty These genotypes somewhat resemble race compacta. except that the ~nflorerence branches are not Incurved Members of race compacta are k n o w as cockscomb ftnger millets In both Afrtca and India Spikelets are up to ten flowered, w~th the irlflorescence branches dlvlded at the base and strongly incuwed to form a f~st-like inflorexence lndtan cultivars commonly have an ~nflorescence branch located some dlstance below the terminal cluster on each primary Inflorescence axls A f r ~ a n cultlvars usually lack this lower inflorescence branch Race compacta IS grown In northeastern India. Ethiopia and Uganda

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Race vulgaris 1s the most common flnger mtllel of A f r ~ a and Asla It 1s yrown In A f r r a f rom Uganda to Eth lop~a and to south A f r r a , and In Asia from ltrdla to Indunes~a Inflorescence branches are twisted or Incurved Some yel>utypes are drouyltt tolerant, others are well adapted l o area5 of hlylr ramfall. and st111 others are sown Itr nurseries and transplanted to l~e lds w ~ t h the lirst rains of the season In rlce yrowtng areas race vulyarts ohen follows irr~yated rice as a r a b ~ crop Gratns are cooked as r r e , or ground In lo flour to make porr~dye or unleavened bread

RlNOR CEREALS OF EURASIA

1 ell minor nilllets ate yrown III Asia 1 heir s ~ ~ r v ~ v a l as cereals In r u m p s t ~ t ~ o n wllh rlce a ~ l d wheat attests to the sayn~f~cance of these cereals In the ayro ecosystems o l Asla Two of llrese Pansutr~ rn~lraceun~ L (broomcorn or proso nrtlletl and Setorlo ~tal,co ( L ) Y &auv (foxla11 ntilletl are yrown across temperate burasla w ~ l h l ox ta~ l mtllet extend~ng Into the semi arid troplcs of Asla Other nrllror cereals are Imponant In special~zed ayrtcultural rrtches In Asla

Crabyrass or manna Dtg~lorro so r~yu~nu l~s ( L bcop 1s a cornmoll weed In all temperat; parts of the world 1 he specvs IS morpholoy~cally var~able and var~ously classtf~ed Into subyenera and varlelles (Gould 196.3) It IS a n annual yrass w ~ t h prostrate or decumbent stems and f l o w e r ~ t ~ y culms that can reach well over one metre In he~yh t 1 he speoes was harvested as a spontaneous srnil domesttcate 1s southern Europe u r i t~ l the f~rs l quarter of t h ~ s century (Wetlh. 19371 It 8s cultivated as a cereal In the Caucasus and Kaslrrnir (Henrard 1950 Bor 1955) Crabyrass has beerr a crop for at least 2.000 years Pltn~us wl io l ~ v e d In the f~rst century A L) referred to the species as ~ x l i a c m o n and sugyested ttrat 11 was o l b lav~c ortylns (Korntcke a ~ ~ d Werner 18851 Matth~olus (15651 recorded that this cereal was yrown tn Bohen i~a blavon~a and the Ukralne K o r n ~ ~ k e and Werner (1885) reported that in t l u~ tya ry one hectare ylelded 4 2 0 520 k g of yraln and 780 1200 k g o f hay LJurlny the late ntneteenth century the specles was an Important cereal in southeastern tu rope (Ascherson and Graebner 18'Xl) Wi ld manna was harvested w ~ t h a ackle k f u r r the pldnts wete fully matured ( b r ~ k e r D~ l l~ t l yen , 1Y271 I t never lost i l te a b ~ l ~ l y o f t~a lura l w e d d~spe~sd l Little is k ~ l o w n ah,ul its present cultlvatlon i n the Caucasus and Kashm~r

Another crabqrass. V~g t lo r~a cruc~olo (Nees) A Camus. IS cult~vated hy the Khas~ trlbes In Assam. where 11 IS known as ralshan Hooker and Stapf (18'96) recognuad t h ~ s cereal as a spr'les o f Pospolunr and &r (19401 ntlstakenly ~ncluded 11 tn 1)8y1larto coryrriboso (Koxb 1 Merrlll Bor (1955) correctly transferred ratshall to D cruc~afo and r e c o y ~ r ~ r r d the cult~vated kinds as var ewulerlto Bor Veldkamp (197.31 described vat peclrnato Veldkamp to ~rrclude a cult~var w t h ylutlnous gratns tlial was yrown at Cha Pa tn northern Vl r lnam

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Ralshan IS an annual grass w~ th prostrate to decumbent culms that root at the lower nodes, and that produce flowertng culms up to 1 3 rn tall lnflorerences conslst of two to ten racemes arranyed on a 1 4 cm long central axls. w~ th the racemes up to 1H cm long reflexed at maturity 1 he chartaceous lemma and palea lightly enclose the grain at malurtty, but the gmln is read~ly freed from th~s fru~tcase by poundtng tn a mortar

Raishan pensts as a cereal in Assam probably because it prov~des excellent and essential feed for livestock durtng winter when graelng is scarce It 1s commonly sown on land from whtch potatoes or other crops have been harvested It ts planted between early Aprll and late June and harvested In November In October the culms are-tted together into hunches and allowed to mature lndivtdual plants vroduce as manv as 30 flower~nu culms with each . . inflorescence malurtng at a sltghtly different time Spikelets are collected by hand about a month after the matn inflorescence niatures (Singh and Arora. 1972) Harvested sp~kelels are dried in the sun bpfore they are stored, and usually parched over a flre belore they are pounded in a mortar to remove the lemma and palea The clear>ed grains are botled in a mixture with rice Stnsh and Arora (1972) re~orter l that ra~bl~nn v~elds up HOO ku h a 1 The species was probably domt*sticaterl by ht11 tribes-in ~ s s a i n and soubeast Asla

The genus tch~norttloo 15 wtdely d~strihuted, arid includes wme 20 species

several of which are aygresslve weeds The most obnoxtous weed 15 t oryzondes (Ard ) I'ritscti It trtvades paddy rice, mln~~cs llle crop In veyetat~vr morphology, and flowers a few days earl~er thar~ the cult~var i t accnrnpanws as a weed Natural seed d~spersal before the crop is harvested, and seed dormancy ensure a new populatton of Echinocl~loo weeds when rice is planted in the same field during the next growing season

Barnyard millet. E crusgollr (L ) P Beauv LS a commori weed of temperate and warm regtons of the Old and New Worlds The specles is cultivated i n Chtna, Korea and Japan, where I! 1s con~monly known a5 Japanese millet Archaeologtcal records tnd~cate that 11 was grown In Japan dur~ng the Yayoi penod dattng back some 5,MK) years (Watanabe. 1970) Cultivated plants are erect, tufted annuals up to 1 m tall lnflorerences are erect or sltylitly bent at maturity, with the axc.ndrng rdcemes oftera ~nrurvccl at the ttp Sp~kr.lc.ts are perststent and typ~cally cuspidate

The related Echrnochloo colo,m (L ) Ltnk was harvested a5 a wtld cereal tn predynastic Egypt (Ihxon, 1969) lntestlnal contents of mummies excavated at Naga ed-Der tnclude, among other plant rematns, recogn~rable gratns of E colono It IS a minor wild cereal in Central Afr~ca where the grains are fermented to make beer (Ttswrant. 1053) Thts cereal has as yet not been Identifled from among plant rematns of the numerous farming sites excavated In lndia (Vtshnu-Mittre. 1977) The species 15, however, extensively grown In central lndia where 11 1s commonly known as sawa (de Wet et al . 1983a)

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J M J de Wet 25

Sawa dtffers from Japanese mllkt pnmarlly In be~ng a more tropral grass, and m lackmy the beak to the spikelet that characterues E crusgollr Both species have 211 - 54 clirornosomes but hybrtds between them are sterile (Vabuno 1'966) Sawa 1s an lnd~yenous crreal In Indta The spectes Is weedy. spat~taneously tnvades cult~vated f~elds and IS ofteti uri~rite~it~o~tally harvehted wth other mltwr ri?~llrls bawa ts yrowl, 111 It>dla Ironi Kaalimtr lo btkklm In the north and to 1 an111 Nadu tn the south Cult~vated k~nds are extensively varlabk (de Wet el a1 1YXJa) The strongly branched racemes of some cull~vars suggest afftn~t~es wllh Japanese mtllet, but E crusgollr does not occur tn lnd~a as a cereal

Kodo mllkt Pospolun~ scrob~culotum L . IS another lndlyenous cultlvated crreal of indla The spectes IS w~dely dlstrlbuted In damp hab~tats across the lroplcs and subtrop~cs o l the Old World It IS known to have been grown In

1 southern Rajasthan and Maharastitra for at least 3 . 0 years (Kajale. 1977) It ts grown today Iron> Utlar Pradesh to Bangladesh 11, the north, and Kerala and Tam11 Nadu In the suuth 1 h ~ s crreal I5 known as kodo In H lnd~ and varayu tn lam11 A small seeded and larye seeded klnd are recogr~lred by farmers In Tam11 Nadu

Raceme morpholoyy allows lor tlie rec~yri~t lon of three cultlvated complexes l h e most commori kodo m~llets are character~red by racemes wlth the spakelets arranged 111 two rows on one side of a flattened rach~s as 1s also typral of wild P scrob,culotun~ Two varlatlons on tills sptkelet pattern often occur ~n the same ftdd as the more conlrnon phenotype In the one complex. rp~kelets are arranged Ira two to lour ~rreyular rows along the rachls In the other complex, the lower part of each racrmr, has four ~rreyularly arranged ruws of sp~kekts wlr~le sptkelet arrangemelit becomes more regularly two rowed In the upper part of the raceme Ide Wet et a1 1983b) Hybr~dtaatton butween cultlvaied kinds, and between weedy and cultlvated races IS common Thls explalns the absence of clear rac~al dlflerent~atlon, even after some 3 000 years of cult~vat~on as a cereal tn lnd~a

Farmers belleve that kodo millet IS polsonous after a rain It Is known to produce unconulousness. or d r l ~ r ~ u m w ~ l h v~olent tremors of the voluntary muwles Kodo mlllet 1s cooked a5 r r e Bh~de and A~men (1959) suggested that the glumes, lemmas and paleas contaln polsonous alkaloids It is more lkely that the polsomng results from a fungus (ha! often Invades and eventually replaces the developing gram The spore masses are about the same s~ze as mature grams and are not easy to detect at harvest tlme Removlny the husks and wnnowng scatter the spores and only healthy grains remaln to be used as food Potronlng only occurs when the grams are damp at threshing, and the soore5 are not wnnowed away

Adlay. Cou locryma pba L . a grown under shifting cultivstion asa rainfed oop by the hill trlbes of troplcal Asla from Assam to the Philippines (Arora.

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1977) The gralns of wild adlay are enclosed In Indurated lnvolucres Wlld C lacryma.)obl IS called Job's tears. and the ~nvolucres of wlld taxa are used to make rosaries and m k b c e s Involucres are papery In most cultlvars allowing for the ready removal of the grain In Assam. the gram IS ground Into flour. and used to make bread, or a sweet d~sh IS prepared by frylng the grain and adding sugar The whole grain is also eaten raw as a snack, or fermented to produce beer

Two minor millets. Setorla pumilo (Poa ) Roem and Schult , and Brochrana rarnosa (L ) Stapf. are ~nd~genous as cereals to the hills of central lndia As ~ l l d specles, however, they are w~dely d~stributed In tropical Afr~ca and Asla Cult~vated kinds are dlst~ngu~shed from their close w~ld relatives only In the absence of efflc~ent natural seed d~spersal Complexes w~th various degrees of sp~kelet d~sartrulat~on commonly occur In the same f~eld Both specles often occur as encouraged weeds In f~elds of finger or foxtail millets

Two Euras~an cereal specles are of commerc~al Importance These are ' Setario italrco (L ) P Beauv (foxtall millet), and Pon~curn m#l~aceum L (broomcorn mtlletl Both species are extens~vely grown across temperate Euras~a, with foxtall m~llet extendmy Into the troplcb and sub tropics of Asia

The closest wild relative of foxtall millet 1s the weedy green foxtail. Setaria itahco subsp u~ridrs (L ) Thellung Green foxtall IS natlve to temperate Eurasia. but was introduced and became w~deiy establ~shed as a weed In temperate parts of the Amer~cas W ~ l d and cult~vated S ~tal~ca cross naturally to produce fert~le hybrids (Li, Pao and LI 1942. LI. Li and Pao. 1945. de Wet Oestry- Stidd and Cubero. 10791 Derivatives o l such hybrids are obnoxious weeds In the Amencan corn belt (Pohl. 1966)

The antlqulty of foxtall m~llet cultivat~on 1s uncertain The species could have been domesticated anywhere across ~ ts natural range extending from Europe to Japan It has been grown In Cl~lna for at least 5.(Y)O years (Ho. 1975) Jars f~lled with husks of foxtail m~llet were found at Ban po in Shanxi prwlnce dating from the fang shao per~od (Nai. 1'963. Chang. 1973) Foxtail m~llet also occurs in early agricultural sates from Switzerland and Austria dating back some 3.000 years (Werth. 1937) The species became widespread as a cereal In Europe during the Bronze age (van Zeist. 1970) It is absent from known early farming sites in lndia (Vishnu-M~ttre. 1968) This, however, does not necessarily Indicate a late introduction of fortail millet into the tropics and subtropks of South Asia Its wide distribution and morphological variation suggost a long hlstory of cultivation in tropical Asia

Foxtail mtllet is commonly classlfled into a European complex (race moharla) and a Far Eastern complex (race maxims) Race moharia includes cultivan with relatively small and erect inflorescences (Kornicke and Werner. 1885), while race maxima is character~zed by large and pendulous inflorescences (Dekaprelevkh and Kasparlan, 1928) Two inflorescence types

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J M J de Wet 27

of race maxlma are recognized by Gr~trenko (1900) Plants w ~ t h small. essentially erect. and compact lnflorerences occur In northwestern C h ~ n a and Monyolaa Plants from eastern Ch~na. Japan and Korea lyplcally have larye. compact. and pendulous ~nflorescerrces Cult~vars from India are morphokgually d~st~nct from those of Europo and the Far East, and are recognned as race lnd~ca by Prasada Rao et a1 (1987) Plants are typically robust, with lnflorerences bearlng branches that are loosely arranged along the primary ants Some collect~ons from northern l n d ~ a resemble race maxlma and probably represent ~nhoducllons from Chlna Collect~ons from Meghalaya have long, slender lnflorescences w ~ t h small sp~kelets Two collect~ons of the lnternat~onal Crops Research lnst~tute for the b e m ~ Arad Troplcs ( ICHISAl) lrom Karnataka have slender, erect, lnflorescences with short lateral branches. somewhat resembllny S purntla In ~nf lorerence and sp~kelet structure These collect~ons, however, have one to three bristles below each splkelet, whereas S purn~lo has four or more brlslles supporting each sp~kelet

The progenitor of broomcorn millet (Panlcum m~l~aceuml IS natlve to Manchur~a The specles was probably introduced Into Europe as a cereal at least 3 000 yearsago Splkekts and florets of broomcorn mlllet occur together w~th remalns of foxta~l millet In early farm~ny sites of the European neol~th~c Neuweller (1%) dated these s~tes to around 1600 B C During the Bronze aye the specles rap~dly spread across Europe as a cereal (H)elmqulst. 1955)

Cult~vated klnds o f P m~ltoceum are commonly subdivided into f ~ v r subspec~es (Lyssov. 19751 These are here recognued as races without taxonombc va l~d~ty Race mll~aceunr resembles wild P mil~oceum In mllorexe~xre rnorpholoyy i t a charactenzed by large, open ~nflorescerices w ~ l h suberect branches that are sparingly subd~vided Race patentlsslmum with 11s slender and diffused pan~cle branches IS often d~fflcult to dlstlngu~sh from race mli~aceum These two races occur across the range of broomcorn millet cult~vat~on from eastern Europe to Japan H~gh ly evolved cult~vars o f broomcorn m~llet have more or less compact ~nllorescences These are class~f~ed into races contractum, cornpacturn and ovatum Cultivars Included in race contractum have compact. drooptny ~nflorexences Those belonylng to race cornpactum have cyllndr~cal shaped ~nflorexences that are essentially erect Cu l t~ars with compacl and sightly curved inflorescences that are ovate in shape are Included In race ovatum

Races have n o ecogeographk unity Thls probably IS due to extensive movement of seed of the crop across Euras~a, particularly slnce early In this century Lyssov (1975) illustrated 2 1 ~nflorescence types that are commonly grown In Eurasa

A d~fferent Pan~cum specles (samal IS grown as a cereal in the eastern Ghats d i n d ~ a (Rangaswarn: Ayyangar and Achyutha Warlar. 1941) This specles, P sumatrense Roth ex Room and Schult , represents the

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domeshcaled complex o f the weedy P p s ~ l o p o d i u n ~ Tr tn ide Wet . Prasada

R a o and Brmk . 1984) The common ly cu l t ~va ted k~nd d~ f f e r s f r o m w i l d P psi lopodturn wi th w h ~ h it crosses t o produce frrt t le hybrids, primari ly I n havtny

lost the abllity of natural seed dtspersal Th ls race of sama is htyhly tolerant t o heat and drought stress In more favourabk agricultural hah~tats o f the eastern

Ghats a robust race o f sama is g rown The inflorescences of thts race are strongly

branched a n d compact Sama IS often y rown as a mtxture wi th foxtatl mil let.

pear l rntllet o r sorghum

LITERATURE CITED

Arrna H K 1'177 Adky ICuuil rrup I%> Mr!qtroby., Jt~~~rttulcd1l~t Irtclw,i l I ~ r~u r~ ru l . *~~x r t y 52: 05 9 K

Auherwn P ancl I' Graebnet IW'XI Syt~opsrr drr Matrktrmpnwrl~r,~ I lure I clma ti Rrrlart

445 PP 6.8 I1 lldW Contrbulmn, lo Ihu flora uf Nurtlt f'nlay,rta and nr lp~rnt l.rrttc,ry Lrlnoear>

Swr ry Journal 01 Eiola,ly 21: 203 240 Be rk r Mlllngrn J 1027 llondbucll drr Geln~~drbatrrr Rt.rlh, Bor N L 1940 Gramwear Fbra 01 Auorrt 5: 205 207 Bor N L 1955 The genus I)lgUona Hest n l i)ilb Webhn 1 I: 101 407 Bhde N V and H H Amsn 1'J5LJ l'harmarok~gy 01 n tmnqt~tl~rlny p r ~ n r ~ p k 10 Pospalum

~ rmbrub rum gralnr Nulure 153: 17 1') 17 Y1 Brucher. I1 1979 Ilar sngeblclje C~ertrr.nuun> Chduu 50 .lahrr nach Vov$luv L~ l r ch r~ l r lur

Plbnsenzurhlcr 83: 113 147 Bu-n f 1965 Pbrrlr, Ahrnmlaaes de IO l~r r r Alrkast I r<c?hlr Mnrwtlle* C a l m E 0 1965 bowl habnrol some I'ru Columbmn Muxkan lrdmrlr I ccanonta Lloluny 19:

3.35 34.3 Csl rn L 0 1Yb7 lhr Inst New Work1 'errol /\,,~ri, Ar~ra(88IIy 32. ', 15 $8 Chevslrr A 1933 Hewurcer v r ~ ~ l a b k > du hl iara el dr u.r cor,llrlr Nvrd ul 5ud Hrvur

Inlernolanal Boran~lur Applique d Agrruhure lrupkol 12: h(r'J ')Ic) C h q . K 1973 Rsdocmtxn dater horn Chna %me omlal t~lrrpretanr,n, Lunrnl A t i l l ~ r o ~ b g y

14: 525 528 Cbylon. W V 1972 Grarnlnew In J flucltnwm and J M IJalrul I d s J l b r o 01 rropral

Weu AN. 31 349 574 Chomko. S A and G W Ctadord t1)7Ii Ilant hurbar,rlry m prrhislorr enstern Nonh Arnvnca

New evidence In I s drvrbpmrnl Amorrnn Anrrqulty 43: 405 408 k ls rua . 5 H J M J de W n ssd J H tlarbu 197') 1 lleralurr rrvrw alld nurnnkal tnxonrlrny

of trogroub t d It el) Cronomt Eioeny 33: 41 i 424 Crur A W 1972 El Bmmur mongo pbnta derpareclda Ideala 2: 127 111 Vek.p.IPvkh, L L d A S Kaqxwan 1928 A cconIrbupn 10 the audy 01 loxtal mUkf l5elorrr

rtoka (L ) P B wbrp w x h a Ak l I cukesled In Grayls iwesforn Tranuaucasbl Bullelln o j Apphd 5 n m y and Pbnl &cedng 19: 53.i 572

de W a J M J 1977 DarPrtkaoon 01 A h a n cerrak Ahran Fconon~k blulory '3 (spring) I'J77 15 32

dr Wul J M J and C A CJekt, 1979 Ibmuara~,n of Amerran wlld rkr I/rronla erquorro L . Gaminow) Journol dAghukure lro~lllx~rtul r t llolormkqur Apphqorc 25: 07 X4

& Wet. J M J . K E R a d a H a o and LJ t Ikmk I'M4 5y,trrnstcr and dorne.lkatlon 01 Panrum SurmDenu iGrsmkese1 Journal d Aghukure Trndllonal el Holonque Appllque 30: 159 168

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J M J de Wet 29

d e W . J M J . K E R.lsdaRso.DE h k n d M H Mcmgo*la 1984 Sqmvmahesanded- I c n of EL-• comano (Gramn .~ I Amchon Journd of Baanu 7 1: 550.557

d. Wel. J M J . K E R d a R.o, M H Mcngcrha and D E Brtnk 1Y8h b m l k a l i o n 01 -a Mdkl IEchnahbo cDbnol Emm- Baonu 37: 2U3 291

d. wet J M J . K t ~ d a ~.o. M H t.+eng.Aa and D E &Ink M-uy in kodo MJkl, P-bm vmbrubtum Economic B&ny 37: 159.163

&Wet. J M J . L L (kwry.5ldd and J I Cube10 1979 OIlglnsand evolulton of foxlaU mllkts (Setom *aTcal Journal d Aphukure Tmd#unal el Botonlpw AppYque 26: 53-64

Dam. D M 1969 A nde on cereah m anc~n t Egypt In J P Ucho and F W Mmbkby k d t I The Domelacaben and Expbddlon of h n U and Anmob Akin. Chcago p 131.142

Dorr W G 1%9 WHd Prcc Gnada Dppsrtmnl 01 Agrkukuro. Research Branch PuMkatan 1393 04 UP

and Ethndogy 31: 257,334 u Y . F W 1%3 Cylaaxonomy of &#om wnguhab and D k e n d e n s Bntonh 15:

- 241.244 Grlnnko. R J 1%0 Chumira l lelan mllktl tsionomy (Setorb Ilolco IL I P B s u b moirrno

Ak f I Bulean of Appbcd B&ny Genetkr ond Pbnt B~ecdmg 32: 145-182 Henrard, J Th 1950 Mondrsph of the genus &#om Unhren(ly of Lelden Preu. Lelden

B99 PP Hiu. K W ud J M J &Wet 1976a DmwsUcamn 01 EkuMc mrrona (L I Gurma Emmmk

Bourny 30: 199-208 Hdu. K W and J M J de We1 1976b Racbl evolution In Ekwine comcana t u b corwana

(tinget mYkl A w r l o n J o m l of h n y 63: 1311.1318 H r h u t . H 1955 D* d a s Cuthkhk dnKulurpbnnn h Schw&n Amqubl md Wrl.

Stockholm Ho. P 1975 The Cmdk of the East Chkago UnhreWy Prev C h e a p . IL H o o k . J D and 0 %pi 1896 Carheae h J D Hodur k d I Fbroo1BrI.h hdb Vdume

7 Reeve and Co , London Jwdn. C 1%7 Lbt of Food. b e d m Ahko Food and Agrkukural Organlution of the Un~ted

b r a n s P u b l r d h , R- Kapk. M P 1977 Ancrnt gahs from eicauatbnl at Ncvoua. Maharashha Geophytobgy 7:

98 lW Kunrke. F and H Werwr 1885 Handbwh det CHlrcldebow. Volumes l and ll Verbs Paul

Psrey. Par* LI. H . W K Pao and H W L 1W2 inknpecllk cr-t h Setarb Journd of Herrduy 33:

351.355 . H . C H L and W K Pa0 1W5 Cyldogkd and pene(la1 Uudkl 01 the mlerrpcb'k cross o l the cukkakd dkl. SemYI t a k a IL 1 h a u v . and the meen IoxinY d k t S uktdb L J o d of thz Amerran Sockly 01 Apmnomy 37: 32 54

L y r w . B H 1975 Row lt'onrum L h A S K r w led I Fbra of Cukualed PbnU Vol U1 GMI Gopr Kdar Lcnkprd

ManaekIod. P C . R S M r M and W C C a ~ n a l 1%7 Prehbtonc WJd and cultbaled mase In D S &m Ied ) T)n Prehblorv of Tehuocan Vdey UnhrersUy of Texa Prsu. Avrm pp 178-200

M.ahbbr. P A 1565 Corninenton in Lbros Scr Pcdoch h u o n d b A n d l & k d k a M.tcrrs Venre

Mdna. G J 1702 Saggk~ Sulo SM*, Nmurak del Chll Bdogna. h l a N.bhn. G and J M J de Wet 1984 Ponkurn sonorum h the Sawran M agl*uhure

EconomY Botany 38: 65-82

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Na. H 1%3 hhudogy h New Chha Annputy 37: 176 185 NNuah. E 1% k h w w -hkhllklcher FTb- Vkrlulhhmhrlm dur Nuturalmchung

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mn and s)RlemnIksd culkaled ScflrM Ualco IGrammerl trommlr Dorony 41: 108 I lb Ranglawml Awangar. C N and U B Arhyulha Warw 1% 1 Samas !he llttk nlllkt IPonkunt

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d h t cowmk Urnany 16: S7b :YIO Tburanl. R P Ch 1 W Lbyrtuhure dans k r cavanpr drl'Oubanqut Bulletrn D'lnsrulr Elude

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h w d y 01 Tokyo 5: 357.379 Wenh. E 1937 Lur Glo(paphtp und G.uhrhlr der Hklen Angeuondre Bolank 19: 42 W Yabuno. T I966 L*on/rtrmslx sludy of the gvrlur Echhochloo IGramlnunrl Japanese Jour.

n d 01 Baony 19: 777.323