Amer. J. Bot. 71(4): 550-557. 1984. SYSTEMATICS AND EVOLUTION OF ELEUSINE CORACANA (GRAMINEAE)1 J. M. J. de W et,2 K. E. Prasada Rao,3 D. E. Brink,2 and M. H. Mengesha3 departm ent of Agronomy, University of Illinois, 1102 So. Goodwin, Urbana, Illinois 61801, and international Crops Research Institute for the Semi-arid Tropics, Patancheru, India ABSTRACT Finger millet (Eleusine coracana (L.) Gaertn. subsp. coracana) is cultivated in eastern and southern Africa and in southern Asia. The closest wild relative of finger millet is E. coracana subsp. africana (Kennedy-O’Byme) H ilu & de Wet. W ild finger m illet (subsp. africana) is native to Africa but was introduced as a weed to the warmer parts of Asia and America. Derivatives of hybrids between subsp. coracana and subsp. africana are companion weeds of the crop in Africa. Cultivated finger millets are divided into five races on the basis of inflorescence mor­ phology. Race coracana is widely distributed across the range of finger millet cultivation. It is present in the archaeological record o f early African agriculture that m ay date back 5,000 years. Racial evolution took place in Africa. Races vulgaris, elongata., plana, and compacta evolved from race coracana, and were introduced into India some 3,000 years ago. Little independent racial evolution took place in India. E l e u s i n e Gaertn. is predominantly an African tancheru in India, and studied morphologi­ genus. Six of its nine species are confined to cally. These include 698 accessions from the tropical and subtropical Africa (Phillips, 1972). Indian subcontinent and Sri Lanka, 648 acces­ One species occurs in South America, one sions from Africa, and 63 accessions of un­ species extends from Ethiopia into Arabia, and certain origin..Xwelve. quantitative and nine- one species is a pantropical weed but native to qualitative characters were recorded for each Africa and Asia. Finger millet, E . coracana (L.) accession. Data are derived from observations Gaertn., is cultivated in southern and eastern on at least ten plants in each accession. These Africa, and across most of southern Asia (Hilu observations were compared with field studies and de Wet, 1976a). The cereal is African in in Malawi (Africa) and the states of Andhra origin, was domesticated some 5,000 years B.P. Pradesh and Orissa in India, and with studies in eastern Africa, and introduced into India as of herbarium specimens filed at the Rijksher- a crop 3,000 years ago (Hilu, de Wet and Har- barium, Leiden (L) and Royal Botanic Gar­ lan, 1979). African and Indian cultivated com­ dens, Kew (K). Quantitative characters scored plexes Have been isolated from one another are days to flowering (DFL), plant height (PLH), until historical times. This separation provides number of basal tillers (NBT), number of in­ an opportunity to contrast racial evolution in florescences on the major culm (NIN), flag leaf Africa where the crop is sympatric with its wild length (FLL) and width (FLW), length of flag progenitor, with racial evolution in India where leaf sheath (LFS), peduncle length of terminal wild E . coracana subsp. africana (Kennedy- inflorescence (PDL), inflorescence length (INL), O’Byme) Hilu & de Wet is absent. length of longest inflorescence branch (LIB), width of major inflorescence branch (WIB), M a t e r i a l s a n d m e t h o d s — Germplasm col­ and number of inflorescence branches (NIB). lections of Eleusine coracana filed with the Accessions were classified into six major groups Genetic Resources Unit of the International based upon overall appearance, with the fol­ Crops Research Institute for the Semi-arid lowing sample sizes: Subsp. africana (wild and Tropics (ICRISAT) were planted near Pa- weed) N = 15; subsp. coracana, race coracana N = 206, race elongata N = 88, race plana N = 1 Received for publication 25 June 1983; revision ac­ 287, race compacta N = 202, race vulgaris N cepted 11 October 1983. = 635. Discriminant analysis (direct method, Research supported financially by a grant from the In­ Nie et al., 1975) was used to assess separation ternational Board for Plant Genetic Resources as part of of groups using the above characters. Quali­ its global interest in priority species of millets, and Grant AGR 58-319R-0-165 from the USDA Binational Pro­ tative characters studied are degree of plant grams (US-Spain). The use of facilities at ICRISAT is pigmentation,! growth habit, degree of glume appreciated. prominence, degree of inflorescence compact- 5 5 0 April, 1984] de WET ET AL. — SYSTEMATICS OF ELEUSINE 551 ness, fruit color, degree of lodging at time of maturity, degree of senescence at time of ma­ turity, overall disease resistance, and yield po­ Os — O O tential. Data are filed as permanent descriptors C\ I t~- cs m so cm vi | Irt M —I-OO! | / — COI —I 00I —I —I for each of the Eleusine accessions in the gene — so ec CS in bank at ICRISAT. Specimens of a majority of P o ' S ' o ' ^ r~- o 'r?'£> m — collections studied are filed with the Crop Evo­ c*“> cs <s lution Herbarium (CEL). Systematics—Finger millet is variable (Ta­ ble 1). This led to considerable controversy -?r about its origin and evolution. De Candolle fn — 0 0 o O' ItT (1886) and Cobley (1956) suggested that finger l — — — I — s© r - I I SO O I 1 I I O I millet was domesticated in India, and that its ,^_, ^r»1 — — <N « VO ,a5_, ,«_, ,>5-. ,-_, V, wild progenitor is the pantropical weed, E. in­ !/■> —< (S —>’ vO, C-i —Tf —' —_ ££ dica (L.) Gaertner. Porteres(1951,1958,1970) and Mehra (1963) proposed that this cereal Oao so'o Tf <N riSC SO— O ' <No ‘ O ' '•O —'— C"-' originated in Africa from the native E. africana Kennedy-O’Byme. Vavilov (1951) suggested that finger millet could have been domesticated to independently in Africa and India, and Ken- 3 S ro — OO CJ O nedy-O’Byme (1957) and Jameson (1970) <j s Os lm Os«1 0 —I r-I ©1 ©r —I ©(c s 1 —1© I — I identified E. africana and E. indica as the re­ \o ~ (N—< so — r~- so — ■■a- spective progenitors of African and Indian cul- >3 ^ Oj ^ n o vo « ’t r^_ « « tivars. f s ^■OOOMMJiffi'OvovDvcO Phillips (1972) suggested that E. indica znd E. africana are conspecific, and recognized these taxa as subspecies of E. indica. Cytologically E. africana (2n = 36) and E. indica (In = 18) vn are distantly related, with little homology ob­ w is SO = 'S >/-> — © © O N 't vious between their basic genomes (Chenna- os l o t~-<Nsoc*-icncNmso I ONO\ I I ’—1 I' I I — — veeraiah and Hiremath, 1974). In contrast, hy­ £ * 0000 | |oc©(©fS©l | «13 SO,—.P I —. 04 — 90 — — — brids between E. africana (2n = 36) and E. c Os^Oso — ooO rtos — ^fvo coracana {In ~ 36) are fully fertile. These two m^csso^oscN^-trip'OO^ taxa hybridize where they are sympatric in Af­ — vnosts — ro — — od'^’o^oo rica, and derivatives of such crosses often occur as weeds in cultivated fields. Comparative morphological and cytological studies reveal that E. africana is conspecific with E. cora­ fi £ ■=3- cana, and that E. africana is the progenitor of © —< © © fM v£) «-> Os I P I U1NVOM---OS finger millet (Hilu and de Wet, 1976a). These —lOCSf-l 00 | |soo|cool©l 1—1 l—l — two taxa were combined, and E. africana was 'O — —— oe — — vo — <0 recognized as a subspecies of E. coracana by t--’ — ^ —'• v£_ r4_ —’ Hilu and de Wet (1976b). Stabilized, weedy m sovocvfn - o « co n s ■q:fnodrs'^i(Nr4rntsoo — *6 derivatives of crosses between wild and cul­ r -' — {<1 — — c s — — tivated E. coracana resemble either the cul­ tivated or wild complex in inflorescence mor­ phology. so r~>v-i 00 — so vi m © so 1. Eleusine coracana (L.) Gaertn. subsp. cor­ M O — — I — I I [——I acana Fruct. et Sem. 1: 8, t. 1. 1778. Type: soosi 1 ^ i m ^ rr I I — Illustration in Plukenet, Phytographia t. 91, f. O^fj-optJip^poeeov 0\ 5. 1691. Cynosurus coracan L., Syst. Nat. Ed. c^' O P~t ©_ O M m © © r^> 10, 2: 875. 1759. Cynosurus coracanus L., Sp. S so 00 oc ©' od 00 r^' —- so <n PI. ed. 2: 106. 1762. Eleusine cerealis Salisb., « i* -e Prodr. Stirp.: 19. 1795, nom. superfl. Eleusine . c sphaerosperma Stokes, Bot. Mat. Med. 1: 149. a 1812, nom. superfl. Eleusine stricta Roxb., 3 S Hort. Beng.: 8. 1814, nom. nud. Eleusine to- as CL.ZZ-P-.1 552 AMERICAN JOURNAL OF BOTANY [Vol. 71 cussa Fressen., Mus. Senck. 2:141.1837. Eleu­ ognition of five cultivated races with eleven sine coracana (L.) Gaertn. var. stricta (Roxb.) distinct cultivated complexes. Nees, FI. Afr. Austr. 1: 251. 1841. Eleusine luco Welw., in Apont. Phyto. Geog.: 591.1858, 2. Eleusine coracana subsp. africana (Ken- nom. nud. Eleusine dagussa Schimper in Re­ nedy-O’Byme) Hilu & de Wet, Econ. Bot. 30: gel, Garten Flora 21: 205. 1872. Eleusine cor­ 202.1976. Type-. South Africa, Cape Provinqe, acana (L.) Gaertn. vars. alba, atrajusca Koem., Kimberly district, Warrenton-on-Vaal, Wil- Handb. Getreid. 1: 329. 1885. Eleusine cor­ man H.K.1, March 1950 (K, Holotype). Eleu­ acana (L.) Gaertn. var. tocussa (Fresen.) sine africana Kennedy-O’Byme, Kew Bull. 12: Franch., Bull. Soc. Hist. Nat. Autun 8: 377. 65. 1957. Eleusine indica subsp. africana 1895. Eleusine indica var. stricta (Roxb.) Chiov. (Kennedy-O’Bvme) S.M. Phillips, Kew Bull. in Nuov. Giora. Bot. Ital. 26: 83.