ISSN1880-3881

民族植物学ノオト
Ethnobotanical Notes
in collaboration with plants
第9号

No.9

2016

植物と人々の博物館
自然文化誌研究会

Plants and People Museum
The Institute of Natural and Cultural History民族植物学ノオト 第９号

民族植物学ノオト 第 9 号：目次
Ethnobotanical Notes No.9：contents

巻頭言

多様な生活文化の中の雑穀（特集）………………………………………………………………… 1

Preface

Agricultural complex of millets in the Indian subcontinent
M. KIMATA, Y. ISHIKAWA, H. KAGAMI, A. OTSUBO and K. OTSUKA ……… 2

Domestication process and linguistic differentiation of millets in the Indian subcontinent
Mikio KIMATA……… 12

Tertiary domestication process of korati, Setaria pumila (Poaceae)
through the mimicry to other grain crops in the Indian subcontinent
Mikio KIMATA ……… 25

Domestication process of korati, Setaria pumila (Poaceae), in the Indian subcontinent
on the basis of cluster analysis of morphological characteristics and AFLP markers
Mikio KIMATA ……… 39

Domestication and dispersal of Panicum miliaceum L. (Poaceae) in Eurasia
Mikio KIMATA ……… 52

Appendix: International Forum for Promotion and Development of Minor Cereals (1997) ………………………… 67
資料：国際フォーラムの提案（1997）

あとがき …………………………………………………………………………………………………………………… 69
Editorial postscriptⅰ

民族植物学ノオト 第９号

ⅱ

民族植物学ノオト 第９号

巻頭言

多様な生活文化の中の雑穀（特集）

このくにはユーラシア大陸の極東に位置し、て、独立心を喪失した「米悲願単一民族」に仕

海洋に囲まれながら、多数の巨大都市や工業地立て上げた。柳田の思想の一部、微瑕が拡大解帯を構築した今日でさえ、ありがたいことに領釈されて、この政策に与したのだろう。

域の約 64％が山林に被われている。島ぐにであ

人間も生き物であるから、食料の安全・安心

ると同時に山ぐにであるという特異な自然環境確保は第一の欲求である。ところが、この国のに立地している。台風や地震などの自然災害は

政策は食糧安全保障を放棄したのである。水田

多いが、このことを忘れずに、禍福転じて暮ら

稲作のみに集中した結果の米の過剰生産を減反

せば、平常時にはとても穏やかで豊かな自然環の強制によって対応した。生産を放棄すること境にある。

に対して補助金を出したのである。稲の多収よ

大陸から直接東漸して、あるいは花綵列島をりも安全・安定生産、畑作の麦・雑穀の生産も伝って間接的に南下あるいは北上して、次々と維持すべきであった。世界的な人口激増や環境

多様な民族がそれぞれの生活文化をもってやっ

変動のもとで、不安定な食糧生産と需要の拡大てきて、定住した。島々の先は太平洋だから、によって、海外からの食糧輸入に全面的な依存

この先には行けなかった。豊かな自然は行く必はできない。この国が農業生産を軽視し、食料要も感じさせなかったのだろう。約 1 万年前に安全保障を怠っているので、多様な食料の安全・農耕文化が獲得された後には、栽培植物を含む

安心は家族で、地域社会で確保せねばならない。
私の第 2 の人生は、ユーラシア大陸各地およ農耕文化複合も伴って移住してきた。
この見方からすれば、このくには、ヤマト族

び日本各地への旅に多くの年月を費やした。イ

だけではなく、アイヌ、オロッコ、ギリヤーク、ンド（バンガロール、農科大学）とイギリス（カ朝鮮族などの混合民族によって構成され、山住

ンタベリー、ケント大学）の地方都市にはサバの縄文文化の系譜をうけた畑作農耕から、平住ティカルで、タイ（バンコック、ラジャバト・（低地）の弥生文化の系譜による水田稲作農耕

プラナコン大学）には大学院の集中講義などで、へと歴史的に進展してきたと考えられる。農耕しばらく暮らす機会を得た。このため、とりわが始まる以前の無土器文化であれ、その後の農

けインド農村の暮らし、西欧の農村景観から大耕文化であれ、このくにの生活文化は途切れるきな心象を受けた。第 3 の人生では、旅で観たことのない時を経ながら重層的に蓄積されてきこと、考えたこと、旅の後の調査研究の成果をたはずだ。

アーカイヴすることにした。

ところが、第二次世界大戦後、何故か、柳田

この第 9 号はユーラシアの旅行でたくさんの国男による日本の単一民族論、平住（低地）の農家から分譲を受けた栽培植物の種子・標本を弥生文化の系譜のみを重視する水田稲作農耕が用いた実験研究の成果をまとめた。研究の一部優勢になった。これが今日の日本の農業や農耕は東京学芸大学民族植物学研究室のインド雑穀のあり方を著しく歪め、父祖以来の食糧自給の研究ティームの成果である。彼らの卒業論文や努力を捨て去り、食糧輸入に依存する体制に導

修士論文の成果は各自のものであり、いずれ彼いた思想の一つであると考える。江戸時代の米らが公表することを願いながら、ここでは概要

石高制に加えて、明治維新の殖産興業、戦時中を未発表として記録しておく。
木俣美樹男（2015-9-24）の米配給制度、アメリカに敗戦した後の小麦・脱脂粉乳などの食糧援助が、この国の人々をし1

民族植物学ノオト 第９号

Agricultural complex of millets in the Indian subcontinent
M. KIMATA, Y. ISHIKAWA, H. KAGAMI, A. OTSUBO and K. OTSUKA
Research Team of Indian Millets, Laboratory of Ethnobotany,
Field Studies Institute for Environmental Education, Tokyo Gakugei University

The Indian subcontinent is a wonderland for

Stapf. and Oryza sativa L., (c) Echinochloa frumentacea

studying the domestication process of grain crops.

(Roxb.) Link, Panicum sumatrense Poth., Paspalum

Several species of millet are domesticated in thisscrobiculatum L., Digitaria cruciata (Nees) A. Camus.,region. Cooking of cereals forms an important part of

Setaria pumila (Poir.) Roem. & Schult, and Brachiaria

the agricultural complex. The agricultural complex isramosa (L .) Stapf.; and (d) Southwestern China,

composed of their vernacular names, religious function,

Fagopyrum esculentum Moench, Fagopyrum tataricumarchaeological evidence, etc. The first author had(L .) Gaertn.; and (IV) New World, Zea mays L .,participated six times in expeditions for millet research

including Amaranthus hypocondriacus L., Amaranthus

and collected numerous accessions of millets and theircaudatus L., and Chenopodium quinoa Willd. These

relative species, with information on their agricultural

four cereal and pseudocereal groups accompanied bycomplex, from hundreds of farmers in their villages andagricultural complexes have been introduced several

fields. Ancient farmers had originally domesticated six

times during prehistoric and historic ages from many

species of millet from the relative weed species in India.

regions into the subcontinent.

Indian millet species were domesticated in the process

Indian food culture has been a reflection of theof diff usion from humid paddy fields in Eastern India to

people’s heritage. It represents India’s historical

dry upland rice fields in the Deccan Plateau, Southern

development, religious beliefs, cultural practices, andIndia.

above all, geographical attributes (Sahni 1986). In the
Indian subcontinent, staple foods made using grain

Key words: agricultural complex, domestication, Indiancrops are served with various types of spicy curries andsubcontinent, millet, tertiary crop

legume dal stews. Many unique cooking styles can befound for each cereal in any part of the subcontinent(Aziz 1983, Sahni 1986). Cooking of cereals forms an

Introduction
The Indian subcontinent is a wonderland forimportant part of the agricultural complex (Maeshwari

studying the domestication process of grain crops.

1987, Sakamoto 1988). Indian cooking consists of a

Several species of millet are domesticated in this region.

unique combination of special cooking styles developed

The grain crops cultivated in this subcontinent arefor each grain crop (Kimata et al. 2000). Moreover, theclassified into the following four groups on the basis of

agricultural complex is composed of their vernaculargeographical origin: (I) African, Eleusine coracana (L.)

names, religious function, archaeological evidence, etc.

Gaertn., Pennisetum glaucum (L.) R. Br., and Sorghum

This special issue is composed of studies on the

bicolor Moench; (II) Mediterranean, Hordeum vulgare L.

agricultural complex, domestication process, andand Triticum spp.; (III) Asian, including four subgroups,

dispersal of millets, especially Setaria pumila (syn.

(a) Panicum miliaceum L. and Setaria italica (L.) P.

Setaria glauca) and Panicum milliaceum and not major

Beauv., (b) Coix lacryma-jobi L. var. ma-yuen (Roman.)crops such as rice, wheat, barely, and maize, in the2

民族植物学ノオト 第９号

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Indian subcontinent. Setria pumila has been dispersedand Pennisetum glaucum. These species were introducedin only a very limited area of the Deccan Plateau

via the Arabian peninsula in about 2000 BC (Sakamoto(Kimata 2015a, 2015b), while Panicum milliaceum has

1987, Ohnishi 1998). The New World pseudocereals are

been dispersed throughout Eurasia (Kimata 2015d),

Amaranthus caudatus, Amaranthus hypocondriacus, andincluding the Indian subcontinent, and recently North

Chenopodum quinoa, and they were introduced in 19th

America and Australia. It is very fascinating from ancentury (Sauer 1976).

environmental perspective of history and geography thatIndigenous millet species have been domesticatedthe distribution patterns of Setaria pumila and Panicum

in the Indian subcontinent for about 3500 years

milliaceum are remarkably different.

(Fuller 2002). These millet species are Echinochloa
fur ume nta cea , Panicum sumat re nse , Pa s palum

Many species of millet cultivated in the Indianscrobiculatum, Brachiaria ramosa, Setaria pumila,subcontinent

Digitaria cruciata, and Digitaria sanguinalis. The

Many species of millet are still grown by numerous

former three species seemed to be secondary in origin

farmers in the Indian subcontinent. These species arethrough mimic and/or companion weeds with rain-

divided into three groups on the basis of place of origin:

fed paddy and upland rice in Eastern India. The next

(1) Asia, including the indigenous Indian subcontinent,

two species, Brachiaria ramosa and Setaria pumila,Central Asia, Southeast Asia, and Tibet; (2) Africa; andwere domesticated as a secondary crop associated with

(3) the New World (Table 1). The following seven species

the other millet species via their mimic/companionof Asian millet were introduced from Central Asia,

weed types in Southern India. Digitaria cruciata wasSoutheast Asia, and Tibet: Panicum miliaceum, Setaria

domesticated in the late 19th century by Kashi natives

italica, Coix lacryma-jobi var. ma-yuen, Fagophyrumin Meghalaya, and it is cultivated in the Kashi Hills

esculentum, and Fagophyrum tartaricum. The African(Singh and Arora 1972). Unfortunately, Digitaria

millet species are Eleusin. coracana, Sorghum bicolor,sanguinalis has disappeared, and its origin is unclear.

3

民族植物学ノオト 第９号

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Moreover, it is obvious that several words of the old
Indo-Aryan and Dravidian languages are related to thevernacular names of millets. Consequently, Brachiaria
ramosa and Setaria pumila are called ‘tertiary crops’,
which means they are double secondary crops forother millet species and upland rice. The order of firstoccurrence for millet species in historical sites generally
supports this evolutionary process.
Field trips in the Indian subcontinent
The first author had participated six times in
expeditions for millet research and collected numerous
accessions of millets and their relative species, withinformation on their agricultural complex, fromhundreds of farmers in their villages and fields (Table 2).
Fig. 1. Expedition routes in the Himalayas, Western Ghats
and Eastern Ghats between 1983 and 2001.

He mainly visited the southern foot of the Himalayas
and Western and Eastern Ghats in and around the

A domesticated plant is always accompanied by the

Indian subcontinent between 1983 and 2001. The

basic agricultural complex, which includes cultivation

research team used many means of transportation, suchpractices, processing, cookery, religious use, vernacular

as car, train, airplane, and their feet, for frequent fieldnames, and other aspects.

trips (Fig. 1). Particularly, the trips extended widely overA domestication centre for millet covers the Eastern

Karnataka, Andhra Pradesh, Tamil Nadu, Maharashtra,

Ghats and Southern Deccan Plateau on the basis of field

Telangana, Maharashtra, Madhya Pradesh, Orissa,

observation, experimental results, linguistic sources,

Chhattisgarh, Jharkhand, West Bengal, Bihar, Uttar

and archaeological data. Although this process is quite

Pradesh, Uttarakhand, Himachal Pradesh, and Jammu

complicated among millet and its relatives, it is very

and Kashmir in India and the North-West Frontier in

effective for understanding domestication by a secondary

Pakistan and Eastern Nepal.

origin via an insurance crop, a mimic companion weed,
Millets domesticated in the Indian subcontinent

and weed types. The domestication process indicatesthe importance of the weed–crop complex and basic

Ancient farmers had originally domesticated six

agricultural complexes as a plant–man symbiosis.

species of millet from the relative weed species in India

4

民族植物学ノオト 第９号

Fig. 2. Five species of millet domesticated in the Indian
subcontinent and wild rice: a, Panicum sumatrense; b,
Paspalum scrobiculatum; c, Echinochloa frumentacea; d,Brachiaria ramosa; e, Setaria pumila; and f, Oryza rufipogon in
the irrigation canal.

Fig. 3. Five species of millet introduced into the Indian
subcontinent and a field of inter-cropping with Setaria
italica and Grycine max: a, Sorghum bicolor; b, Pennisetum
glaucum; c, Eleusine coracana; d, Setaria italica; e, Panicum
miliaceum; and f, an inter-cropping field.

(Table 1). Then, these plants were distributed over the

Other mil let species grown in the Indian

Indian subcontinent and neighbouring areas.

subcontinent

Panicum sumatrense (samai) is an annual plant (2nIndian farmers introduced six species of millet (Table= 36, tetraploid) derived from Panicum sumatrense

1) from Africa via the Arabian Peninsula, Central Asia

ssp. psilopodium (Fig. 2a). Paspalum scurobiculatum

through the Himalayas, and South-Eastern Asia via(kodo) is a perennial plant (2n = 40, tetraploid, Fig. 2b).

Assam in the Indian subcontinent during the Indus

Echinochloa furumentacea ( jangora) is an annual plant

Civilization Age or before the dawn of history.

(2n = 54, hexaploid, Fig. 2c) derived from the relative

Sorghum bicolor ( jowar) is an annual plant (2n =weed Echinochloa colona. Brachiaria ramosa (korne)

20, diploid, Fig. 3a) derived from Sorghum bicolor var.

and Setaria pumila (kolati) are annual plants (Fig. 2dverticilliflorum in Eastern Africa. Pennisetum glaucum

and 2e). These plants are secondary crops domesticated(bajra) is an annual plant (2n = 14, diploid, Fig. 3b)from their relative weeds in paddy fields. This will bedomesticated from Pennisetum violaceum in Africa.

discussed in detail below. Digitaria cruciata (raishan) is

Eleusine coracana (ragi) is an annual plant (2n = 36,

an annual plant derived from the relative weed growntetraploid, Fig. 3c) domesticated from Eleusine coracana

in maize or vegetable fields (Singh and Arora 1972). In

var. africana in Eastern Africa.

addition, Oryza rufipogon Griff. (wild rice) is used as an

Setaria italica (thenai) is an annual plant (2n =offering for gods and goddesses during festivals. It grows

18, diploid, Fig. 3d) derived from Setaria viridis, ain ponds and irrigation canals near paddy fields (Fig.

cosmopolitan weed in Central Asia. Panicum miliaceum

2f).

(cheena) is an annual plant (2n = 36, tetraploid, Fig. 3e)5民族植物学ノオト 第９号

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domesticated from Panicum miliaceum ssp. ruderale(Fig. 5a, 1st, 2nd, and 3rd from lower left). Mudde isin Central Asia. Coix lacryma-jobi var. ma-yuen is aa popular food made from cereal flour and originatedperennial plant (2n = 20, diploid) domesticated fromfrom the cooking of ugari brought from Eastern Africa

Coix lacryma-jobi var. lacryma-jobi in South-Eastern

via the Arabian Peninsula (Fig. 4d and Fig. 5b). Figure

Asia. These millet species are mostly grown by mixed

4 shows cooking methods for cereals in the Indian

cropping or intercropping, for example, Setaria italica is

subcontinent: (a) a traditional boiled rice with papad

grown and mixed with Glycine max (Leguminosae), as(crispy salted wafer made from dal, vegetables, andshown in Fig. 3f.

cereals); (b) upma and kesari bhat; (c) dosa; (d) muddeand boiled grain made using Brachiaria ramosa; (e) puli;

Moreover, five species of pseudocereals are grown inand (f) idli.

the Indian subcontinent. Fagophylum esculentum andFagophylum tartaricum (both annual, 2n = 16, diploid)

Sorghum bicolor and Pennisetum glaucum are

were introduced from Tibet. Recently, Amaranthus

mainly used for making roti, while Eleusine coracana iscaudatus, Amaranthus hypocondriacus (both annual, 2nmostly used for making mudde (Fig. 5b) and fermented= 32 or 34, diploid), and Chenopodeum quinoa (annual,alcoholic drink chan (Fig. 6a, b, and c). Other millet

2n = 36, tetraploid) were dispersed from the New

species are mainly used for bhat. A special food, mavu,

World.

is made from the raw flour of Setaria italica and Oryza
sativa as offering for gods and goddesses during festivals.
Nine foods are made using Brachiaria ramosa (Fig. 5a),Foods made using millets
People have cooked many types of food using

mudde is made using Eleusine coracana (Fig. 5b), andmillets and cereals. Mainly bhat (meshi in Japanese),chapati is made using wheat, Triticum aestivum (Fig. 5croti (pan), and mudde (oneri) are cooked because theyand Fig. 6d, upper). With respect to fermented foods, a

are frequently made using most of the cereals listed instarter is made from Hordeum vulgare (Fig. 6a, starter;

Table 3 (Kimata 1987). Bhat is the most popular food, a

6b, a jar for fermentation; 6c, alcohol drink, chan, madeboiled grain food made using all the ingredients shown

from Eleusine coracana; and 6d, yogurt, dahi.

in Fig. 4a, 4d right, and 5a (2nd from upper right).

Other cookeries are shown in Table 3, Fig. 4, and Fig.

Bhat originated in ancient China and was brought to

5. Upma is a coarse-ground grain food (Fig. 4b, right).

the Indian subcontinent via Eastern India. Roti is also

Dosa is a thin leavened pancake stuffed with potato

a popular food made from cereal flour and originatedcurry (Fig. 4c). Idli is a leavened pound cake made usingfrom the cooking of wheat bread in the Fertile Crescent

the same ingredients as dosa (Fig. 4f). Vada is a cakeand was brought to the subcontinent via Western India

made from freshly ground dal or millet flour. Ganji is a6

民族植物学ノオト 第９号

Fig. 5. Cookeries of nine foods made from Brachiaria ramosa (a),chapati from wheat (b) and mudde from Eleusine coracana (c).

nucleotide sequence variation of the chloroplast trnK/
matK region divided cultivated accessions into two

Fig. 4. Cookeries of cereal in the Indian subcontinent:a, a traditional boiled rice (bhat) with papad; b, upuma
and khesari bhat; c, dosa; d, mudde and boiled grain
made from Brachiaria ramosa; e, puli; f, idoli.

groups, northern and southern groups. The northern
cultivated accessions were genetically related to weed
accessions collected from upland rice fields in Orissa.

very thin starch-paste made from the same ingredients

However, southern cultivated accessions showed

as mudde.

close relationships to both accessions of upland rice
fields in Orissa and the weed type in southern states.

Dispersal routes of millets in the Indian subcontinent

Furthermore, two alternate hypotheses for the origin

Our team studied the domestication process andof Paspalum scrobiculatum were summarized: (1)dispersal routes of Indian millets. An outline of the

kodo millet was domesticated once in Orissa and thenresearch results is given below.

diffused to inland and southern states and (2) kodo

Kodora, Paspalum scrobiculatum, was domesticatedmillet was domesticated in Orissa and somewhere insince about 2000 BC in India. This species is cultivatedthe southern states of India, independently (Ishikawa

throughout the Indian subcontinent, but mainly2007).

in Madhya Pradesh. Ecological and morphological

Jangora, Echinochloa furumentacea, is cultivated forcharacteristics were compared using 32 accessions

food, fodder, and as an emergency crop in India, Nepal,(including weed forms) of Paspalum scrobiculatum. In

and Pakistan. Its ancestor is a weed, Echinochloa colona,

addition, the relationship between plant pigmentation

found in paddy fields. Morphological characteristics andand mimicry of rice was observed in 16 accessions,

AFLP analysis results of seven accessions of Echinochloa

including six accessions collected from upland rice fields.

colona and 42 accessions of Echinochloa furumentacea

Domestication process of the secondary crop to uplandwere compared. On the basis of the results, the placerice was discussed (Ishikawa unpublished). Th is speciesof origin was assumed to be around Bihar, and then it

shifted from perennial to annual and obtained crop-likewas distributed to Tamil Nadu via Karnataka (Kagami

traits by accessions with rice cultivation. Both amplified

unpublished).
Samai, Panicum sumatrense, is cultivated for foodfragment length polymorphism (AFLP) analysis and7

民族植物学ノオト 第９号

Fig. 7. Main dispersal route of Indian millets.

its domesticated type is mostly grown by mixed cropping
Fig. 6. Fermented foods: a, starter made from Hordeum
vulgare; b, a jar for fermentation, c, alcohol drink (chan) madefrom Eleusine coracana and d, yogurt, dahi.

with Paspalum scrobiculatum or Panicum sumatrense
in only India (Kimata et al. 2000). Th is domestication
process is discussed in detail below.

and fodder in India, Nepal, Sri Lanka, and Myanmar.
Its ancestor is a weed, Panicum sumatrense subsp.

Dispersal route of Indian millets in the Indian

psilopodium, found in paddy fields. Morphologicalsubcontinent

characteristics and A FLP ana lysis results of 38

The integrating hypothesis for the dispersal route ofaccessions and 281 herbarium specimens of Panicum

Indian millets is illustrated in Fig. 7 on the basis of thesumatrense were compared. On the basis of the results,

results. Echinochloa furumentacea, Panicum sumatrense,

the place of origin was assumed to be Eastern India,

and Paspalum scurobiculatum were secondary crops toand it was then distributed in Southern India (Otsuka

upland rice. First, their ancestral plants were companion

unpublished).

weeds derived from the relative weeds that invaded

Korne, Brachiaria ramosa, is grown by very extensive

paddy fields in humid regions of Eastern India. Second,

farming for food in only India. Its ancestor is a weed

the companion weeds became insurance crops in uplandfound in paddy fields. Morphological characteristics

rice fields, and they spread to a dry region in the Deccan

and AFLP analysis results of 70 accessions of Brachiaria

Plateau (Kobayashi 1987, 1989). Brachiaria ramosa andramose, including both weed and domesticated types,

Setaria pumila were so called ‘tertiary crops’ because

collected from Pakistan and India were compared.

they were secondary crops to other millet species

On the basis of the results, the place of origin was

domesticated from their relative weeds in upland fields.

assumed to be the southern part of Orissa, and it was

On the other hand, Digitaria cruciata has been recently

then distributed in the Deccan Plateau via Tamil Nadu

derived from the relative weed grown in maize or(Otsubo unpublished).

vegetable fields, Kashi Hill, Meghalaya, and is limited to

Korati, Setaria pumila, is a cosmopolitan weed, but

the same area (Singh and Arora 1972).

8民族植物学ノオト 第９号

Indian Subcontinent. Ethnobotanical Notes 9: 32-48.

Tentatively, Indian millet species were domesticatedKimata, M. 2015b. Domestication process of korati, Setaria pumila

in the process of diff usion from humid paddy fields in(Poaceae), in the Indian subcontinent on the basis of cluster

Eastern India to dry upland rice fields in the Deccan

analysis of morphological characteristics and AFLP markers.
Ethnobotanical Notes 9:49-64.
Kimata, M. 2015d. Domestication and dispersal of Panicum

Plateau, Southern India.

milliaceum L. (Poaceae) in Eurasia. Ethnobotanical Notes 9: 6582.
Kimata, M., E.G. Ashok and A. Seetharam. 2000. Domestication,cultivation and utilization of two small millets, Brachiaria ramose
and Setaria glauca (Poaceae), in South India. Economic Botany
54(2):217-227 .
Kobayashi, H. 1987. Mimic and associated weeds with millets andcultivation methods of millets in the Indian Subcontinent. Pages
15-40 in S, Sakamoto, ed., A preliminary report of studies on
millet cultivation and its agro-pastoral culture complex in Indian
sub-continent, I. (1985). Kyoto University, Kyoto, Japan.
Kobayashi, H. 1989. Mimic and associated weeds with millets andrice cultivation in Orissa and Maharashtra in India. Pages 1132 in S, Sakamoto, ed., A preliminary report of studies on millet
cultivation and its agro-pastoral culture complex in Indian subcontinent, II. (1987). Kyoto University, Kyoto, Japan.
Maeshwari , J. K. 1987. Interdisciplinary approaches in ethnobotany.
ed. by Jain, S. K. In A Manual of Ethnobotany. Scientific
Publishers, Jodhpur, India.
Ohnishi, O. 1998. Search for the wild ancestor of buckwheat III.
The wild ancestor of cultivated common buckwheat, and of
Tartary buckwheat. Economic Botany 52:123-133.
Sahni, J. 1986. Classic Indian Cooking , Dorling Kindersley,
London.
Sakamoto, S. 1987. Origin and dispersal of common millet andfoxtail millet. Japan Agricultural Research Quarterly 21(2):84-89.
Sakamoto, S. 1988. Origins and Dispersals of Millets – An EurasianEthnobotany. Nihonhoso-shuppan-kyokai, Tokyo, Japan (inJapanese).
Sauer, J. D. 1976. Gra in Amaranthus (Amaranthus sp.,
Amaranthaceae). In Evolution of Crop Plants, ed. Simonds, N. W.,
pp.4-6. Longman, London.
Singh, H. B., and R. K. Arora. 1972. Raishan (Digitaria sp.) – a
minor millet of the Kashi Hills, India. Economic Botany 26:376380.

Acknowledgements
We had many expeditions conducted by Kyoto
University, Tokyo Women’s University and Tokyo
Gakugei University supported by Japanese and Indian
Governments in the Indian subcontinent from 1983 to
2001.The author would like to thank cordially Indian
farmers for kind giving local varieties of millets; Prof. S.
Sakamoto and Dr. Y. Ishikawa, Kyoto University; Dr.
Y. Suyama, Tohoku University; Prof. I. Fukuda, Tokyo
Women’s University for their kind advice; and to thelate Professor H. Kobayashi, for his excellent advice
during the field survey in 1985, 1987 and 1989.
Moreover, the authors wish to express their hearty
thanks to the Indian farmers in the areas of study for
their valuable information and kindness; to Dr. A.
Seetharam, Dr. Ashok, Dr. Madhukeshwara and Mr.
Mantur, Project Coordination Cell (Small Millets),
ICAR, University of Agricultural Sciences, Bangalore,
India, for their collaboration and useful suggestion:
to Dr. M. Nesbitt and Dr. T. Cope, Royal Botanic
Gardens, Kew, UK, for their valuable suggestion on
excellent collections of literatures and herbarium
specimens.

Literature cited
Aziz, K. 1983. Indian Cooking, Publishing Group, New York.
Fuller, D. Q. 2002. Fifty Years of Archaeobotanical Studies in India:
Laying a solid foundation. In S. Setter and Ravi Korisetter (eds.).
Indian Archaeology in Retrospect. Vol. III. Archaeology andInteractive Discipline. pp.247-361. Indian Council of Historical
Research, Manohar.
Ishikawa, Y. 2007. Domestication of kodo millet (Paspalum
scrobiculatum L.) in India. Doctoral thesis for Kyoto University.
Kimata, M. 1987. Grain crop cookery in South India, Research
Team for the Studies on Millet Cultivation and its Agro-pastoral
Culture Complex in the Indian Subcontinent, Kyoto University,
pp.41-55.
Kimata, M. 2015a. Tertiary domestication process of korati, Setaria
pumila (Poaceae) through the mimicry to other grain crops in the

Appendix 1: Reference to graduation and master’s theses by the
Research Team of Indian millet
ISHIKAWA, Yuko 石 川 裕 子 : Undergraduate and graduate
student of Tokyo Gakugei University (1999~2002) ＝修士論文
2001：インド起源の雑穀コドラ Paspalum scrobiculatum L. の栽培化過程
KAGAMI, Hiraku 加 賀 美 啓 : Undergraduate student of Tokyo
Gakugei University (2009~2013) ＝ 卒 業 論 文 2012： イ ン
ド亜大陸におけるインドビエ（Echinochloa furumentacea）の起源と伝播―形態的形質の比較と DNA 断片長多形(AFLP 法 ) から

9

民族植物学ノオト 第９号

MILLET NEWSLETTER 12:18-20.

OHTUBO, Yukino 大 坪 礼 乃 : Undergraduate and graduate
student of Tokyo Gakugei University (2005~2009) ＝ 卒 業 論

Kimata, M. 2000b. 南 イ ン ド に お け る 2 種 の 雑 穀 コ ル ネ

文 2006：南インドの栽培植物コルネにおける種分化
PCR=RFLP 法による分析、修士論文 2008：南インドの雑穀コルネ Brachiaria ramose (L.) Stapf. の栽培化過程
OHTSUKA, Keita 大 塚 啓 太 : Undergraduate and graduate
student of Tokyo Gakugei University (2009~2014) ＝卒業論文
2011：サマイの起源と大陸伝播―形態的形質と DNA 多型による種内比較調査、修士論文 2013：サマイのインド亜大陸における伝播

Brachiaria ramosa とキンエノコロ Setaria glauca の栽培化
過程、Domestication of Brachiaria ramosa and Setaria glauca inSouth India. MILLET NEWSLETTER 12:23-25.
木俣美樹男 2000c. 南インドにおけるイネ科雑穀のコルネ
およびコラリの栽培化過程とその利用（Domestication
and utilization of two small millets, korne (Brachiaria ramosa) andkorali (Setaria glauca), Poaceae in South India）、育種学研究 2（別 1）；245
木俣美樹男 2002、世界の雑穀類と栽培状況、農林水産技
術ジャーナル 25（11）：11-16
木俣美樹男 2003、第 11 章雑穀の亜大陸インド、山口裕文・
河瀬真琴編『雑穀の自然史―その起源と文化を求めて』
pp.145-162、北海道大学図書刊行会
木俣美樹男 2008、シコクビエの酒・チャン（分担執筆）、
山本紀夫編『酒づくりの民族誌』pp.219-226、八坂書房
Kimata, M. and S. Sakamoto. 1992. Utilization of several speciesof millet in Eurasia. Bull. Field Studies Inst., Tokyo Gakugei
University, 3:1-12.

Appendix 2: Reference to Indian millet research by Mikio Kimata
Dr. of Agricultural Sciences, Professor Emeritus of Tokyo Gakugei
University and Fellow of Research Institute for Languages andCultures of Asia and Africa, Tokyo University of Foreign Studies
Fukuda, I. and M. Kimata. 1983. III-3. Legumes in Nepal. In ed. by
I. Fukuda, Scientific Research on the Cultivation and Utilizationof Major Crops in Nepal. pp.40-58. The Japanese Expedition of
Nepalese Agricultural Research, Tokyo.
石 川 裕 子 ･ 木 俣 美 樹 男 2001、 コ ド ラ（Paspalum
scrobiculatum）の栽培化過程における形態的特徴、育種学研究 3（別 1）：154
Kimata, M. 1983. Characteristics of some grain crops, garden crops
and weeds, and methods of cooking grain crops in Nepal. In
ed. by I. Fukuda, Scientific Research on the Cultivation andUtilization of Major Crops in Nepal. pp.40-58. The JapaneseExpedition of Nepalese Agricultural Research, Tokyo.
Kimata, M. 1987. Grain crop cookery in South India, Research
Team for the Studies on Millet Cultivation and its Agro-pastoral
Culture Complex in the Indian Subcontinent, Kyoto University,
pp.41-55.

Kimata, M. and A. Seetharam. 1997. Processing and utilization ofsmall millets in Eurasia. pp.112-114. National Seminar on Small
Millets, Indian Council of Agricultural Research and Tamil Nadu
Agricultural University.
Kimata, M., S.G. Mantur and A. Seetharam. 1997. Cultivation
and utilization of small millets in hill regions, Uttar Pradesh andHimachal Pradesh, India. Environmental Education Research,
Tokyo Gakugei University. 7:33-43.
Kimata, M., M. Kanoda and A. Seetharam. 1998. Traditional andmodern utilizations of millets in Japan. Environmental Education
Studies, Tokyo Gakugei University 8:21-29.
Kimata, M., S. Fuke and A. Seetharam. 1999. The physical andnutritional effects of the parboiling process on the grains insmall millets. Environmental Education Studies, Tokyo Gakugei
University 9:25-40.
Kimata, M., E.G. Ashok and A. Seetharam. 2000. Domestication,cultivation and utilization of two small millets, Brachiaria ramosa
and Setaria glauca (Poaceae), in South India. Economic Botany
54(2):217-227.
Kobayashi, H. and M. Kimata. 1989. Millets in Maharashtra
and Orissa in India. Research Team for the Studies on Millet
Cultivation and its Agro-pastoral Culture Complex in the Indian
Subcontinent, Kyoto University, pp.1-9.

木俣美樹男 1988、雑穀の栽培と調理（分担執筆）、佐々木
高明・松山利夫編『畑作文化の誕生 縄文農耕論へのアプローチ』pp.189-211、日本放送出版協会
木俣美樹男 1988、南インドにおける雑穀の栽培と調理について、生活学 第 13 冊：127-149
木俣美樹男 1989、雑穀のルーツをもとめて、健康医学
437：43-48
Kimata, M. 1989. Grain crop cookery on the Deccan Plateau,
Research Team for the Studies on Millet Cultivation and its Agropastoral Culture Complex in the Indian Subcontinent, Kyoto
University, pp.33-50.
木俣美樹男 1990、インドの穀物料理、教室の窓 中学社
会 新しい社会、No.343:6-8、東京書籍
木俣美樹男 1991、インドにおける雑穀の食文化（分担執筆）、『インド亜大陸の雑穀農牧文化』pp.173-222、学会
出版センター
木俣美樹男 1995、シコクビエの酒・チャン（分担執筆）、
山本紀夫・吉田集而編著『酒づくりの民族誌』pp.219-＜口頭発表＞
木俣美樹男 1987、インド亜大陸における雑穀の栽培と調理、生活学会第 13 回秋季研究発表会
木俣美樹男 1991、南アジアのキビの変異、第 5 回雑穀研究会シンポジウム
木俣美樹男 1993、雑穀の利用、アジア民族文化フォーラ
ム「南アジアの食文化」シンポジウム、アジア民族造形文化研究所木俣美樹男 1997、Brachiaria ramosa と Setaria glauca の栽培

226、八坂書房
木俣美樹男 2000a. インド亜大陸北部における雑穀類の伝播（Cultivation and utilization of small millets in north India）、

10

民族植物学ノオト 第９号

化と利用、第 11 回雑穀研究会シンポジウム
Kimata, M., E.G.Ashok and A. Seetharam. 1998. Domestication,cultivation and utilization of two small millets, Bracharia ramosa(L.)Stapf. and Setaria glauca (L.)P.Beauv. in South India. p.25.
VIIth International Symposium, International Organaization of
Plant Biosystematists.
Kimata, M. 1999. Domestication of Brachiaria ramosa and Setaria
glauca. XVI International Botanical Congress - Abstracts.
Kimata, M. 2000. Domestication process of korali, Setaria glauca(Poaceae). 41st Annual Conference, Society for Economic
Botany. Abstracts pp. 14-15.
木俣美樹男・石川裕子 2002、南インドにおけるコラリ
Setaria glauca の サ マ イ Panicum sumatrense お よ び コ ド ラ
Paspalum scrobiculatum への擬態と混作、第 16 回雑穀研究会シンポジウム
木俣美樹男 ･ 石川裕子 2004、コラリ Setaria glauca のサマイまたはコドに対する擬態と混作、第 19 回雑穀研究会
シンポジウムKimata, M. and Y. Ishikawa. 2004. Mimicry and mixed croppingof Setaria glauca with Panicum sumatrense or Paspalum
scrobiculatum, Ninth International Congress of Ethnobiology.
A58.
Kimata, M. 2013, Domestication process of Setaria pumila (Poaceae)
related to the weed-crop complex in Indian subcontinent,
Conference Handbook p.86, 54th Annual Meeting of the Society
for Economic Botany, Plymouth, UK.

11

民族植物学ノオト 第９号

Domestication process and linguistic differentiation ofmillets in the Indian subcontinent
Mikio KIMATA
Plants and People Museum

The vernacular names of millets were gathered

have been domesticated across their ranges of present-

through field surveys in the Indian subcontinent

day cultivation for some 3500 years (de Wet et al. 1983a;

since 1983. Farmers have an appropriate awareness

Fuller 2002; Pokharia 2008). These millets includeof the status of millets and their relative weeds in

Paspalum scrobiculatum L. (kodo millet), Echinochloa

the domestication process. This symbiotic process

frumentacea Link (Indian barnyard millet), Panicum

between millets and farmers was reconstructed bysumatrense Roth. (little millet), Brachiaria ramosa (L.)

integrating field observations, botanical experiments,

Stapf. (korne), Setaria pumila (Poir.) Roem. & Schult.

archaeological data, and linguistic sources. There were

(korati; syn. Setaria glauca (L.) P. Beauv.), Digitaria

various vernacular names in the Eastern Ghats andcruciata (Nees) A. Camus (raishan), and Digitaria

Southern Deccan Plateau, where Indian millets weresanguinalis (L.) Scop. (Chandra and Koppar 1990; dewidely cultivated with their relative species today. It is

Wet et al. 1983a, b, c). The former three species seem

obvious that the several names in the old Indo-Aryan

to be secondary in origin, through the mimic and/or

and Dravidian languages are related to the vernacularcompanion weeds of the rain-fed paddy and then uplandnames of millets. Brachiaria ramosa and Setaria pumila

rice in Eastern India. The next two species, Brachiaria

have been domesticated from the weeds that grewramosa and Setaria pumila, were domesticated asaround upland rice fields via a mimic companion weed

secondary crops that were associated with the othertype that was mainly related to Panicum sumatrense andmillets via their mimic companion weed types inother grain crops. Brachiaria ramosa has become an

South India (Kimata et al. 2000; Kimata 2015a,

independent crop in pure stands, while Setaria pumila

2015b, Kobayashi 1987, 1989). Digitaria cruciata was

grows as a mixed crop with Panicum sumatrense anddomesticated in the late nineteenth century by Kashi

other millets. Consequently, Brachiaria ramosa andnatives in Meghalaya and is cultivated only in the Kashi

Setaria pumila are so-called “tertiary crops,” meaning,

Hills (Singh and Arara 1972). Unfortunately, Digitaria

they are a double secondary crop for the other millets

sanguinalis has disappeared, and its origin is not clear.

and upland rice. The order of first occurrence of millets

In contrast to other millets, which were probably

from historical sites generally supports this evolutionary

domesticated in humid Eastern India, Brachiaria

process. This domestication center of millets covered

ramosa and Setaria pumila have adapted to the dry

the Eastern Ghats and Southern Deccan Plateau.

climate of the semi-arid tropics. Brachiaria ramosa wascultivated in the hot, arid red soil region of Southern

Key words: dispersa l, domestication, ling uistic

India, whereas Setaria pumila was cultivated in the hot

differentiation, millets, mimic companion weeds

sub-humid ecoregion in red and lateritic soils of Orissa,
as well as in the hot semi-arid ecoregion on red loamy
soils of Southern India (Sehgal et al. 1992). Brachiaria

Introduction

ramosa tolerates drought better than Setaria pumila, it

The indigenous millets of the Indian subcontinent12

民族植物学ノオト 第９号

agricultural complex,” the so-called “from seeds tostomach” idea, was proposed by Nakao (1967) while
studying the origin of agriculture. A domesticated plant
always is accompanied by a cultural complex, which
includes cultivation practices, processing, cookery,religious use, vernacular names, and other aspects(Kimata and Sakamoto 1992). Bellwood and Renfrew(2002) recently proposed and examined their “farming/
language dispersal hypothesis” cooperative across thedisciplines of archaeology, linguistics, and genetics froma broad comparative perspective. These millets andtheir relative weeds also have many vernacular names
in each locality and language. This report is concernedwith the reconstruction of their domestication process,particularly Brachiaria ramosa and Setaria pumila,from the point of view of their vernacular nameswith reference to linguistic archaeology, because good
linguistic data have not yet been sufficient for theindigenous millets (Fuller 2002; Southworth 2005).
Fig. 1. Field surveys in the Indian subcontinent.

Field surveys and methods
has undergone a specializing adaptation to arid regions,

Extensive field surveys were conducted in Karnataka,and it has nearly attained the tertiary domesticatedAndhra Pradesh, and Tamil Nadu in 1985, 1996,

phase (Kimata et al. 2000). On the other hand, the1997, and 2001; Maharashtra in 1987; Orissa inlocal varieties of Setaria pumila have adapted to drier

1987 and 2001; Madhya Pradesh and Bihar in 1989;

fields in Southern India than in Orissa. Setaria pumila

and Himachal Pradesh and Uttar Pradesh in 1996.

was normally grown with Panicum sumatrense, but it

Furthermore, the surveys were added in Nepal in 1983

seemed to grow singly when the latter failed to growand Pakistan in 1985 and 1989 (edited by Sakamotoin severe droughts, which was observed in our 1987

1987, 1989, 1991). The observations that concentrated

survey. This possibly suggests that Setaria pumila couldon Brachiaria ramosa and Setaria pumila were made in

become an independent crop. Brachiaria ramosa is an

the local fields, particularly in 1996 to 1997 and 2001

underutilized millet that is restricted in cultivation(Fig. 1). The vernacular names of cereals and their wild/

today to dry areas in the two border districts of Tumkur

weed relatives were gathered from local farmers in eachand Anantapur in the states of Karnataka and Andhra

locality and language, used to construct a database, andPradesh, respectively. Brachiaria ramosa is cultivatedwere also extracted from the literature about Indianin pure stands as a sole tertiary crop, while Setaria

agriculture. The vernacular names from farmers were

pumila is still cultivated by mixed cropping withgiven an expression that was written in English by

Panicum sumatrense and other grain crops as a minor

local farmers and regional researchers from agriculture

domesticated plant. A tertiary crop is a type of double

extension stations. Moreover, the vernacular names

secondary crop of Panicum sumatrense and others and aof food items were collected from the English menu of

secondary crop of upland rice.

local restaurants and cookbooks from each state.

T he methodolog ica l concept of the “ ba sic

13

民族植物学ノオト 第９号

Table 1. Vernacular names of Brachiaria ramosa, summer annual in India
State

Language

OrissaOriya

StatusVernacular names

Weed with Pas. scrobiculatum

gusara pata, chusara mataWeed/Domesticated?

ghusara pata, lota, ghada langi
3)

Maharashtra

MarathiDomesticatedchama pothaval

Andhra Pradesh

TeluguWeed
Domesticatedandakora, anda korra, pedda sama , disakalu, edurigaddi

Domesticatedkornne, korale, korne, korneki, kornike, bennakki hullu

Mimic companion weed with P.

koothi same, sakkalati same, same melatti

Karnataka

KannadaTamil Nadu

Tamil

Kelarasumatrense

Malayalamakki hullu, votlu kosavu
1)

Domesticatedkam pampul, palapul

Domesticatedchama pothaval

3)5), pil sama, pani varagu

3)

3)

Italics cited from 1) Fuller 2002, 2) Kobayashi 1991, 3) Ambasta 1986.

each locality and language as shown in Table 2. The

Results
Brachiaria ramosa was cultivated mainly in a few

domesticated type was known by a great variety ofstates of South India. This semi-arid area is subject

vernacular names in Orissa and in the border area

to a savanna climate in Deccan Plateau. Brachiaria

between Andhra Pradesh and Karnataka (cf. Chandra

ramosa and its relatives are summer annuals and haveand Koppar 1990; Kawase 1987; Kimata et al. 2000;

many vernacular names in each locality and language

Kobayashi 1987, 1989). These names were usuallyas shown in Table 1. The following tables contain someshortened to a single word, such as nehari in Orissa,

vernacular names that are cited for the conveniencelingudi in Maharashtra, korati in Andhra Pradesh, korlu

of discussion, but the results of surveys are from thein Tamil Nadu, and korin in Karnataka, and the names

author’s own data. This domesticated type has been

were sometimes composed of two words, including

known by various vernacular names in Maharashtra andkuku lange and kukur lange in Orissa, kora samuru

South India (cf. Chandra and Koppar 1990; Kawase

in Andhra Pradesh, and samuru korra in Karnataka.

1987; Kimata et al. 2000; Kobayashi 1987, 1989). The

The mimic companion weed type was known by many

domesticated type was called similar names: hamavernacular names, too. Further, these names were

pothaval in Maharashtra, chama pothaval in Kelara, andusually a single word, such as nauri in Bihar, lingri inkama pampul and palapul in Tamil Nadu. On the other

Orissa, nauri in Madhya Pradesh, korale in Andhra

hand, it was called different names in the border area

Pradesh, and erikorra in Karnataka. They sometimes

between Andhra Pradesh and Karnataka, mainly korne,

have adjectives that indicate the associated plants, forkorneki, and andakora, and sometimes pedda sama andexample, in Andhra Pradesh, varagu korali and varagu

disakalu. The mimic companion weed type was known

sakkalathi indicate a companion weed of kodo millet,

as koothi same, sakalati same, and pil same in Tamilwhile samalu korali and arasama indicate a companion

Nadu. The weed type was known as gusara pata andweed of little millet. The weed type was often calledchusara mata in Orissa, and akki hullu and votlu kosavu

navari in Madhya Pradesh, ghas in Orissa, and uniquein Andhra Pradesh.

names such as ghoda langi, meaning horse tail, in Orissa
and sana korulu, meaning little foxtail millet.

Setaria pumila was cultivated at a few hill sitesthat were mainly in Orissa and South India. ThisThe vernacular names of other indigenous millets

semi-arid area is also subject to a savanna climate inand rice in the Indian subcontinent are shown in Table

Deccan Plateau. Setaria pumila and its relatives are

3. The domesticated type of Panicum sumatrense, asummer annuals and have many vernacular names insummer annual, was usually called samai, same, sama,14

民族植物学ノオト 第９号

Table 2. Vernacular names of Setaria pumila, summer annual in India
State

Language%LKDU+ LQGL0 LP LFFRP SDQLRQZ HHGZ LWKųƄƖőƖƆƕƒƅƌƆƘƏƄƗƘƐQDXULQDYULQHEULQHXULQHYULQLEULKDUULWXWXDP2ULVVD

2 UL\D: HHG

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Status

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Vernacular names



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

Italics cited from 1) Fuller 2002, 2) Kobayashi 1991.
Austin 2006: korai [kora, korali ] (Bengali, Deccan, Hindi, India and Bangladesh), bandra (Hindi, India), varagu korali (varagu , firewood, korali , ear or corn, Tamil)and similar names in South India, while it was called

varagu in Karnataka; and varagu in Tamil Nadu. Thevari and wari in Maharashtra, gurji and koeri in Orissa,

mimic companion weed grew in upland rice fields. It

and gondula in West Bengal. Further, indigenous

was called kodo and kodaira in Madhya Pradesh, kodo

people called it various names, including kutki (Vaiga)

war in Bihar, and kodoghas (Paraja) in Orissa. The wild/

and mejheri (Gobdi) in Madhya Pradesh; gundli

weed type was called kotocha in Maharashtra, khar sami

(Munda) in Bihar; ghantia (Kunda Tading), gurgi

and kodo wani in Bihar, and kodo ghas in Orissa.

(Kunda Dora), and suau (Paraja) in Orissa; and batta

The domesticated type of Echinochloa frumentacea, a(Kotha) in Tamil Nadu. The mimic companion weed

summer annual, was known as jangora in Uttar Pradesh;type was identified and called akki marri hullu, meaning

sawan and similar names in Madhya Pradesh andweed-like rice, kadu same, meaning weed little millet,

Bihar; sankari wari in Maharashtra; jhari, dhatela, andand kosu samalu only in Karnataka, while the weed typegruji suau (Paraja) in Orissa; ooda in Andhra Pradesh;

was sometimes called kadu and fodo in Karnataka, gabat

kudurai vali in Tamil Nadu; and wadalu in Karnataka.

in Maharashtra, and erigola and arasama in Andhra

The ancestral weed species, Echinochloa colona was called

Pradesh.

chichivi in Maharashtra, dhela in Orissa, and probably

The domesticated type of Paspalum scrobiculatum, asain in Bihar. Digitaria cruciata was a summer annual

perennial, was mainly called kodo, kodora, and similar

called raishan only in Kashi Hills. The domesticatednames, but it had different names such as harik in

type of Coix lacryma-jobi was a perennial called re-si in

Maharashtra; arik in Andhra Pradesh; arka, alka, andNagaland (Church 1886), while the other weed species

15民族植物学ノオト 第９号

Table 3. Vernacular names of other indigenous millets and rice in Indian Subcontinent
Language&RXQWU\State
Growth habit

StatusVernacular names (Indigenous people)ųƄƑƌƆƘƐŃƖƘƐƄƗƕƈƑƖƈ ųƄƖƓƄƏƘƐŃƖƆƕƒƅƌƆƘƏƄƗƘƐ ŨƆƋƌƑƒƆƋƏƒƄŃƉƕƘƐƈƑƗƄƆƈƄ
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3DNLVWDQ

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3XQMDE+ DU\DQD
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that often invaded rice paddy fields was called gulru

Oryza sativa L., a perennial, was usually calledin Madhya Pradesh, gurya, meaning small, in Bihar,

chawal or dhan, but the upland rice was called lehi inkorankhar in Orissa, and garemara in West Bengal.

Madhya Pradesh, gora dhan in Bihar, gadeba dhan in16民族植物学ノオト 第９号

Table 4. Vernacular names of Asian and African millets in Indian Subcontinent&RXQWU\State
Growth habit

Language

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2 U\D

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MXQHURP DNDLEDMUDOrissa, and probably gouri in Karnataka. The wildwith those of Indian millets in Table 4. These species

relative O. rufipogon Griff. was used specially for a

are all summer annuals. Panicum miliaceum L. wasfestival food and called pasahi in Madhya Pradesh,

widely called cheena and similar names, while it wasdeobath in Maharashtra and probably balunga in Orissa.

known as wari and tane in Maharashtra and varagu

The vernacular names of Asian and African millets

and similar names in Orissa, Andhra Pradesh, Tamilin the Indian subcontinent are shown for comparison

Nadu, and Karnataka. Setaria italica (L.) P. Beauv. was17

民族植物学ノオト 第９号

Table 5. Vernacular names of other cereals in the Indian subcontinent
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also widely called kangani, kauni, and similar names inin Tamil Nadu, junero in West Bengal, and junero

Sanskrit, while it was called rala and rai in Maharashtra,

makai in Nepal. Pennisetum glaucum (L.) R. Br. waskorra and navane in Andhra Pradesh, korra and thenai

also generally called bajra and similar names, but it

in Tamil Nadu, and navane in Karnataka. Eleusine

was sometimes called kayna in Orissa, sajja in Andhra

coracana Gaertn. was usually called ragi in Madhya

Pradesh, and cumba and similar names in Tamil Nadu.

Pradesh, Orissa, and South India, while it was called

The vernacular names of the other cereals are shown

mandua, marwa, and similar names in Uttar Pradesh

in Table 5. Triticum aestivum L. was called gehun,and Bihar, natuni and similar names in Maharashtra

godi, and similar names. Triticum dicoccum Schübler,and Karnataka, tamada in Andhra Pradesh, kapai in

Char. et Descr. was gangil in Tamil Nadu and aja in

Tamil Nadu, and kodo and similar names in Uttar

Karnataka. Hordeum vulgare L. was called jao andPradesh, West Bengal, and Nepal. Further, indigenous

similar names. Those two species are winter annuals.

people called it various names, such as manje suau

Avena sativa L. was not cultivated in South India. Zea(Paraja), mandia (Kondho), and pahado mandia (Kondmays L., a summer annual, was widely called makai andDora) in Orissa. Sorghum bicolor Moench was generally

similar names, while the relative teosinte was introduced

called jowar and similar names, but it was called cholam

for fodder and was called jenera in Bihar.

18

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民族植物学ノオト 第９号

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to the same species as a result of hybridization between

via a mimic companion weed in upland rice fields. This

the domesticated type and the closely related weed type.

thoroughly conforms to the observations that were

The domestication process is supported by themade in the fields. The vernacular name of Echinochloa

linguistic recognition of various types by farmers,

frumentacea is clearly distinguished from that ofsuch as the weed, companion weed, mimic companion

Echinochloa colona, which is one of the ancestral species

weed, semi-domesticated, and domesticated types(Yabuno 1962). For instance, the former is called jhariof Brachiaria ramosa and Setaria pumila, in theirand the latter is dhela in Orissa (Table 3). Sometimes,

vernacular names (Tables 1 and 2). The linguistic

the same names were used by farmers to name Panicum

differentiation shows a close relationship to thesumatrense and Echinochloa frumentacea, same anddomestication process, for instance, in Jalaripalli

sawan, but the names were not used in the same place

Village, Andhra Pradesh, where Setaria pumila that isand time. In the same way, the vernacular name of

mixed with little millet is called kora samuru, meaning

Eleusine coracana is distinguished from a relative weed,

foxtail millet-like little millet, and tela samuru, meaning

Eleusine indica, and the hybrids. However, the weeds

the grains mixed with little millet, which is sold at

associated with other millets and cereals have no names

a local market. This linguistic recognition suggests(Tables 4 and 5). Interestingly, Panicum miliaceum

clearly the agro-ecological status of Setaria pumila as aand Setaria italica have various names in North-Westsecondary origin (Kimata et al. 2000).

Frontier Province and Gilgit, Pakistan (Kawase 1991).

The vernacular names of Panicum sumatrense and

The vernacular names of Indian cookery-used millets

Paspalum scrobiculatum distinguish three types inare unique, particularly in South India, because rice

their domestication process. The names of the mimic(eastward) and wheat (westward) are staple foods today

companion weed type are called, for example, akki hullu

in the other states (Table 6) (Kimata 1991).

(little millet), meaning a rice-like weed, and kodoghas,

The linguistic archaeological names of millets andmeaning a kodo millet-like weed in upland rice fields

other cereals are summarized in Table 7. The old Indo-(Kobayashi 1991). The linguistic differentiation

Aryan names for Brachiaria ramosa, Setaria verticillata,

indicates that both species were also a secondary crop

Setaria pumila, and Panicum sumatrense are not found20

民族植物学ノオト 第９号

in the ancient literature (cf. Southworth 2005). This

(2001) but modified with additional information (Fuller

might indicate that these millets were domesticated inand Madella 2001; Fuller, personal communication). H.

India relatively recently. In contrast, because Paspalum

vulgare, Triticum species (great many), and Avena sativa

scrobiculatum is named kodorava, this word is considered(a few) were identified in the Early Phase (around 4500

to be the origin of kodo and kodora. The word syamaka

B.C.) of Harappan sites. O. sativa (many) and Panicum

for Echinochloa frumentacea is considered a derivation

miliaceum (a few) were identified in the Mature Phaseof shama and sama. The word cina(ka) of Panicum(around 2600 B.C.). Then, Setaria species (great

miliaceum is also considered to be the origin of cheena,many), Sorghum bicolor (many), and Pennisetum

and the words kanku(ni) and rahala for Setaria italica

glaucum (syn. americanum, trace) were found in theare the origin of kangani, which was widely used, andLate Phase (around 2000 B.C.). The following species

rala, which was used in Maharashtra. The word madaka

were found in early South Indian sites (2300 to 1800

for Eleusine coracana is considered to be the origin of

B.C.): Panicum sumatrense (trace), Brachiaria ramosa

mandua in Uttar Pradesh and the word *bajjara is the(many), Setaria verticillata (many), and Setaria pumila

origin of bajra (*, reconstructed forms by Southworth(trace). Then, traces of Paspalum scrobiculatum and2005). The Dravidian name *var-ak- for Paspalum

many Echinochloa cf colona (possibly Echinochloa

scrobiculatum and Panicum miliaceum is considered tofrumentacea) were identified in the late sites (1800 to

be the origin of varagu, and the names *tinai and *nuv-

1200 B.C.). Asian millets occurred historically in the

an-ay for Setaria italica are the origin of thenai in Tamilfollowing order: Panicum miliaceum; Setaria species;

Nadu and navane in Andhra Pradesh and Karnataka.

then Brachiaria ramosa, Setaria verticillata, Panicum

Because these species have old Indo-Aryan or Dravidiansumatrense, and Setaria pumila; and Echinochloa

names, they might have been introduced from thecf colona and Paspalum scrobiculatum. However,Western areas or domesticated within India a relatively

Brachiaria ramosa, Setaria verticillata, Setaria pumila,long time ago, according to the archaeological evidence

and Echinochloa cf colona might have been gathered as a

(Weber 1992).

wild grain.

The first occurrence of grain crops in South Asia is

The naming scheme of millets and their relative

summarized in Table 8, which is based on Fuller et al.

weeds is summarized in Table 9. Farmers have four21

民族植物学ノオト 第９号

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YDUDJX>VRZ LQJLQ-DQXDU\@stages of awareness of the symbiotic process between

indica, was known as maruani. As a consequence of thisthem and plants. They are unknown (stage I), non-survey, farmers appear to have an appropriate awarenessdistinctive (II), identified (III), and classified into someof the status of millets and their relatives, even though

local varieties (IV). In stage I, the farmers have no namethey sometimes use the same names for millets infor wild/weed plants and call them ghas and hullu. In

different places.

stage II, the farmers use the same name for the crop (ragi)

In conclusion, the domestication process of millets

and weed (ragi). In stage III, the farmers identified

based on field observations (Kimata et al. 2000),

and called millets a specific name, for instance, madua

experimental results (Kimata 2015a, 2015b), andfor Eleusine coracana (domesticated) and khadua forthese linguistic sources is illustrated in Fig. 2. This

Eleusine indica (weed). Furthermore, farmers added

domestication center of millets covered the Easterna few adjective words to the root of the millet name,

Ghats and Southern Deccan Plateau. Although thisfor example, to mean “weed” (ghas lingudi, meaning

process is quite complicated among millets and theirweed of Setaria pumila) and “like another crop” (samerelatives, it is very effective for understanding themelatti, meaning mimic weed like little millet), anddomestication by a secondary origin via weed andto indicate a morphological (bilai lange, meaning cat’s

mimic companion weed types. Oats and rye were thetail) or ecological trait ( yerri arasamulu, meaning

secondary crops of wheat that developed cold tolerance

weed with grain shattering) and a utility (pil sama,(Vavilov 1926), while Indian millets were secondary

meaning Brachiaria ramosa for fodder). In stage IV,

crops of upland rice that developed drought tolerance.

farmers classified the millets into some local varieties,

Bachiaria ramosa tolerates drought better than Setaria

for example, Eleusine coracana was known as marua andpumila, and it became an independent crop. Setaria

was classified into the varieties agat- (early), madhyam-

pumila is almost always grown with little millet, but(medium), and pichhat- (late); and a weed, Eleusine

it seems to grow singly when little millet fails to grow22

民族植物学ノオト 第９号

To Southern
dry area

Tertiary crop
S.pumilamixed

field D

Dispersal

From Easternhumid area

field D

cropping D Sakkalathi-same 㸿㹕

Secondary cropEc.
frumentacea

B.ramosa

Pas.scrobiculatum
Gusara-pata 㹕field D
S.pumila 㸿㹕 Pan.sumatrense field D
Dhe㹪aW

Kodoghas 㹕
Jangle rice W

Panicum sp. 㹕
S.pumilaS.verticillata 㹕El.coracana field ‫ڹ‬Upland rice field D

Rice paddy field ‫ڹ‬Fig. 2. Domestication process of millets in the Indian subcontinent

Research, New Delhi.
Austin, D.F. 2006. Fox-tail millets (Setaria: Poaceae) – Abandoned
food in two hemisphers. Economic Botany 60(2): 143-158.
Bellwood, P. and C. Renfrew. 2002. Foreword. pp.xiii-xiv. in P.
Bellwood and C. Renfrew eds., Examining the Farming/Language
Dispersal Hypothesis, McDonald Institute for Archaeolog yResearch, Cambridge.
Chandra, U., and M. N. Koppar. 1990. Diversity and domesticationof minor millet species in Indian sub-continent. Indian Journal of
Plant Genetic Resources 3(2):47-58.
Church, A.H. 1886. Food-grains of India. The Committee ofCouncil on Education, London.
de Wet, J. M. J., K. E. Prasada Rao, M. H. Mengesha, and D. E.
Brink. 1983a. Diversity in kodo millet, Paspalum scrobiculatum.
Economic Botany 37:159-163.
de Wet, J. M. J., K. E. Prasada Rao, M. H. Mengesha, and D. E.
Brink. 1983b. Domestication of sawa millet (Echinochloa
colona). Economic Botany 37:283-291.
de Wet, J. M. J., K. E. Prasada Rao, M. H. Mengesha, and D. E.
Brink. 1983c. Systematics and domestication of Panicum
sumatrense (Gramineae), Journal d’Agriculture Traditional et
Botanique Applique 30:159-168.
Fuller, D. 2002. An agricultural perspective on Dravidian historicallinguistics: Archaeological crop packages, livestock and Dravidian
crop vocabulary. pp.191-213. in P. Bellwood and C. Renfrew
eds., Examining the Farming/Language Dispersal Hypothesis,
McDonald Institute for Archaeology Research, Cambridge.
Fuller, D. Q. and M. Madella . 2001. Issues in Harappan
achaeobotany: Retrospect and prospect. in Indian Achaeology
in Retrospect, Vol. II. Protohistory. S. Settar and Ravi Korisettar

in severe droughts. Both species are so-called tertiary
crops, meaning, they are a double secondary crop for theother millets and upland rice. The millet domestication
process indicates the importance of weed–cropcomplexes and basic agricultural complexes as a plant–
man symbiosis.
Acknowledgements
The author wishes to express his hearty thanks to theIndian farmers in the areas of study for their valuable
information and kindness; to Dr. M. Nesbite and
Dr. T. Cope, Royal Botanic Gardens, Kew, for theiruseful suggestion and kind arrangement for examining
literature and herbarium specimens; to Dr. D. Fuller,
University College of London and Prof. F. Southworth,
Pennsylvania University, for their valuable advice andcitation permission; and to the late Prof. H. Kobayashi,
for his excellent advice and warm collaboration during
the field survey in the Indian subcontinent.

Literature cited
Ambasta, S.P. 1986. The Useful Plants of India. Publications &Information Directorate, Council of Scientific & Industrial

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民族植物学ノオト 第９号

cultivation and its agro-pastoral culture complex in Indian sub-(eds). Publications of the Indian Council for Historical Research,continent, I (1985). Kyoto University, Kyoto, Japan.

Manohar, New Delhi, 317-390.

Sakamoto, S. 1989. A preliminary report of studies on millet
cultivation and its agro-pastoral culture complex in Indian subcontinent, II (1987). Kyoto University, Kyoto, Japan.
Sakamoto, S. 1991. Agro-pastoral Culture Complex of Millets
in Indian Subcontinent, Gakkai-Shuppan Center, Tokyo (inJapanese).
Sehgal, J. L., D. K. Mandal, C. Mandal, and S. Vadivelu. 1992. AgroEcological regions of India. Oxford and IBH Publishing Co.,New Delhi, India.
Singh, H. B., and R. K. Arora. 1972. Raishan (Digitaria sp.) ---- aminor millet of the Kashi Hills, India. Economic Botany 26:376380.
Southworth, F.C. 2005. Linguistic Archaeology of South Asia.
Routledge Curzon, London.
Vavilov, N. I. 1926. Studies on the origin of cultivated plants. Bull.
Appl. Bot. Plant Breed. (Leningrad), 16(2):1-248.
Weber, S. A. 1992. South Asian Archaeobotanical Variability. In
South Asian Archaeology 1989, C. Jarrige (ed). pp. 283-290.
Prehistory Press, Madison Wisconsin.
Yabuno, T. 1962. Cytotaxonomic studies on the two cultivatedspecies and the wild relatives in the genus Echinochloa. Cytologia
27:296-305.

Fuller, D. Q., R. Korisettar and P. C. Venkatasubbaiah. 2001.
Southern Neolithic cultivation systems: A reconstruction basedon achaeobotanical evidence. South Asian Studies 17: 171-187.
Kawase, M. 1987. Variation and distribution of millets in South
India. Pages 5-14 in S. Sakamoto, ed., A preliminary report ofstudies on millet cultivation and its agro-postal culture complex
in Indian sub-continent, I (1985). Kyoto University, Kyoto,
Japan.
Kawase, M. 1991. Millet and their phylog eny in Indian
Subcontinent. pp.33-98. in Sakamoto, S. ed., Agro-pastoral
Culture Complex of Millets in Indian Subcontinent, GakkaiShuppan Center, Tokyo (in Japanese).
Kimata, M. 1991. Food culture of millet in Indian Subcontinent.
pp. 173-222. in Sakamoto, S. ed., Agro-pastoral Culture Complexof Millets in Indian Subcontinent, Gakkai-Shuppan Center,
Tokyo (in Japanese).
Kimata, M. 2015a. Tertiary domestication process of korati, Setaria
pumila (Poaceae) through the mimicry to other grain crops in the
Indian subcontinent. Ethnobotanical Notes 9:32-48.
Kimata, M. 2015b. Domestication process of korati, Setaria pumila(Poaceae), in the Indian subcontinent on the basis of cluster
analysis of morphological characteristics and AFLP markers.
Ethnobotanical Notes 9:49-64.
Kimata, M. and S. Sakamoto. 1992. Utilization of several speciesof millet in Eurasia. Bulletin of Field Studies Institute, Tokyo
Gakugei University 3: 1-12.
Kimata, M., E. G. Ashok and A. Seetharam. 2000. Domestication,cultivation and utilization of two small millets, Brachiaria ramosa
and Setaria glauca (Poaceae), in South India. Economic Botany
54(2):217-227.
Kobayashi, H. 1987. Mimic and associated weeds with millets andcultivation methods of millets in the Indian subcontinent. Pages
15-40 in S. Sakamoto, ed., A preliminary report of studies on
millet cultivation and its agro-postal culture complex in Indian
sub-continent, I (1985). Kyoto University, Kyoto, Japan.
Kobayashi, H. 1989. Mimic and associated weeds with millet andrice cultivation in Orissa and Maharashtra in India. Pages 1132 in S. Sakamoto, ed., A preliminary report of studies on millet
cultivation and its agro-postal culture complex in Indian subcontinent, II(1987). Kyoto University, Kyoto, Japan.
Kobayashi, H. 1991. The origin of secondary crops of millet inIndia. pp. 99-140 in Sakamoto, S. ed., Agro-pastoral Culture
Complex of Millets in Indian Subcontinent, Gakkai-Shuppan
Center, Tokyo (in Japanese).
Nakao, S. 1967. The origin of Agriculture. pp.329-496 in
M.Morishita and T. Kira, eds., The Nature --- ecological studies,
Chuoukouronsha, Tokyo (in Japanese).
Pokharia, A.K. 2008. Palaeoethnobotanical record of cultivatedcrops and associated weeds and wild taxa from Neolithic site,
Tokwa, Uttar Pradesh, India. Current Science 94(2): 248-255.
Sakamoto, S. 1987. A preliminary report of studies on millet

24

民族植物学ノオト 第９号

The tertiary domestication process of korati, Setaria pumila (Poaceae),
through the mimicry of other grain crops
in the Indian subcontinent
Mikio KIMATA
Plants and People Museum

In South India, one domesticated type of Setaria

possibly as long as 12,000 years ago. However, several

pumila is always cultivated in mixed stands along withspecies are threatened and in spite of their potential food

Panicum sumatrense, whilst the others, and the weed

value in their native habitats, have disappeared or havetypes, are grown mostly with Paspalum scrobiculatum,hardly been cultivated. This is because the three major

sometimes Eleusine coracana, and upland rice, Oryza

crops of wheat, rice, and maize have rapidly increased

sativa, in diverse agro-ecological niches in East India.

in their yield and production, due to huge technical

Several botanical traits of S. pumila indicate that

innovation in crop improvement programs. The otherthe domesticated type is descended from the weed

grain crops, that is to say, millets, have decreased

types, and that these traits have become evolutionarily

gradually during the previous century, resulting in

synchronized with those of P. sumatrense and/or

genetic erosion of their local varieties. Today, we needother cereals. This domestication process has beenrecognize their value as exploitable and underutilizedpromoted morphologically through mimicry of othergenetic resources, based on their adaptability to stress-

cereal species and acts as a means of adapting to theprone environments. These species of millet are mostly

arid climate. Moreover, the pigmentation of leaves

C 4 plants, which are early to mature, and can be

and leaf sheaths in terms of anthocyanin composition

cultivated under conditions of severe drought and harsh(HPLC analysis) is involved in mimicry among thesunlight.

other grain crops and the closely-related weeds. The

Local farmers continue to cultivate a few usefulweed and companion weed types are used as fodder forvarieties of millet even today. These indigenous

cattle and as an insurance crop for farmers in times ofvarieties are excellent materials for investigating crop

famine. The grains of S. pumila are always mixed withevolution, particularly the origin and dispersal route

other millet grains, used to make six traditional foodsof domesticated plants. In the Indian subcontinent, a

as a supplementary ingredient, and in turn related to

few small millets are still undergoing the domestication

diversification of the basic agricultural complex. This

process (Kimata et al. 2000; Singh and Arora 1972).

tertiary domestication of S. pumila has proceeded from

While crop evolution can be reconstructed mostly frominter- and intraspecific mimicry by natural hybridization

botanical data, the aspects of geographical origin andand artificial selection in sympatric fields.

dispersal will become clear from the information on thebasic agricultural complex offered by local farmers.

Key words: adaptation, hybridization, millet, mimic

Vavilov (1926) showed the domestication processcompanion weed, tertiary cropfrom weeds associated with wheat to the secondary
crops in two genera, Avena and Secale. For example,

Introduction

Secale cereale L. had built up strong resistance to cold in

Human beings had domesticated more than 30 grasshigh altitude and/or latitude areas, and, subsequently,

species, as cereal crops, in several parts of the world,

this species had been able to grow under more severe

25

民族植物学ノオト 第９号

Fig. 1. Field survey: a) in Karnataka and Andhra Pradesh (1985, 1996, 2001); and b) Orissa (1987, 2001), Indian
subcontinent.

conditions than those under which wheat can grow.

foxtail millet, Setaria pumila was present in limited

Kobayashi (1987, 1989) proposed an integrating

quantities, possibly gathered from the wild (Fuller et al.

model of the domestication process of Indian millets

2001). The first occurrence of cereals in the Harrappan

as secondary crops from mimic companion weeds

Civilization had been wheat, barley, and oats in theassociated with Oryza sativa L.

Early phase (before 2600 BC); Eleusine sp. (problematic,

O. sativa had been spread first from wetlands toE. coracana), Setaria sp., and Panicum sp. in the Mature

uplands, secondarily, in the Indian subcontinent. Theirphase (2600–2000 BC); and Paspalum sp., Echinochloa

ancestral weedy plants had invaded paddy and uplandsp., Sorghum sp., and Pennisetum sp. in the Late phaserice fields in turn. These are Panicum sumatrense Roth.

(more recent than 2000 BC) (Fuller and Madella 2000;(little millet), Paspalum scrobiculatum L. (kodo millet),Weber 1992).

Echinochloa frumentacea Link (Indian barnyard millet),S. pumila is a weedy annual growing 30-60 cm tall.

Brachiaria ramosa (L.) Stapf. (korne), Digitaria crusiata

The inflorescence is cylindrical, densely flowered, spike-(Nees) A. Camus (raishan), and Setaria pumila (Poir.)like raceme, 2.5-10cm long, usually yellow, or more

Roem. & Schult. (korati; syn. S. glauca (L.) P. Beauv.)

rarely purplish or pale green. It is a fairly common grass,(Chandra and Koppar 1990; de Wet et al. 1983a, b, c).

especially in cultivated lands, along the roadsides, andPas. scrobiculatum, P. sumatrense, and E. furumentacea

in cleared forests, up to altitudes of 700 m. The spikelet

were subsequently domesticated by local farmers as(c.a. 3-mm long) is pale green or brownish-green. Cattle

secondary crops to upland rice, because these had put

are fond of it (Achariyar 1921; Singh 1988). S. pumila

up stronger resistance to drought than upland rice inis a multiple polyploid species (2n=18, 36, 72) and an

Eastern India. In this region, several species of millet

unknown genome constitution (almost genome D)

were domesticated.

(Zhao et al. 2013). The present paper is concerned with

Recently, archeological studies have shown very

the tertiary domestication process of S. pumila throughuseful data in the Indian subcontinent. The material ofthe mimicry that is related ecologically and geneticallymillet grasses came from archaeological levels, Phasesto the relative weeds and several grain crops in the

II (2300–1800 cal. BC) and Phases III (1800–1200

Indian subcontinent.

cal. BC) in the Southern Neolithic chronology. They
have been identified as being primarily from twoMaterials and Methods

species, browntop millet, Brachiaria ramosa, andMany local varieties and relative weeds of Setaria

bristly foxtail millet-grass, Setaria vercillata. Yellow

pumila (Poir.) Roem. & Schult.(syn. S.glauca (L.) P.

26

民族植物学ノオト 第９号

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27

民族植物学ノオト 第９号

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Beauv.) have been collected through field surveys of the10, 2002. Two weeks after sowing, germinated plants

Indian subcontinent since 1983. Concentrated field

were transplanted with row spacing of 30 cm andsurveys were conducted in Karnataka, Andhra Pradesh,

plant spacing of 15 cm in the glasshouse. Chemical

and Orissa in 1985, 1987, 1996, and 2001 as illustrated

fertilizer [N:P:K = 8:8:5 (g)] was supplied at 100 g per

in Fig. 1. At the same time, the component species of

m2. Five plants of S. pumila, each of the weed type,millets and weeds were examined in five plots (1 m2)

companion weed, and mimic companion weed types

each in four typical cropping fields (sites) through theassociated with Pas. scrobiculatum and other cereals,

quadrat method. Voucher herbarium specimens andand the domesticated type mixed with P. sumatrense

grain samples were collected along the entire survey

were measured for number of tillers, plant height, spikeroute and deposited at Tokyo Gakugei University andlength, duration to flowering, length and width of thethe University of Agricultural Sciences (Bangalore). The

flag leaf, last internode diameter, number of leaves on

information on agricultural practices, grain processing,the main culm, and number of seeds germinated every

food preparation, and vernacular names was obtained

10 days after sowing. Plant color was assessed using thefrom local farmers.

PANTONE Formula Guide. These data were analyzed
statistically using IBM SPSS (version 21).

As part of these accessions, 60 experimental strains
were selected and grown in a glasshouse at Tokyo

Additionally, the anthocyanin composition

Gakugei University, Japan, in order to compare their

included in the leaf and leaf sheath was analyzed bymorphological and ecological characters in 1998

high-performance liquid chromatography (HPLC)(Table 1) and in 2002 (Table 2). These 10 grains ofin 2013. The materials were S. pumila (5 accessions)

60 strains were sown with a row spacing of 8 cm andfrom site 5 to 8, P. sumatrense (8), Pas. scrobiculatumseed spacing of 2 cm in a box on June 8, 1998 and June(2), B. ramosa (1), and S. viridis (2, fresh and dried). In

28

民族植物学ノオト 第９号

order to identify the retention time (Rt) and measure
the content of each anthocyanin, natural pigments ofcyanidin, pelargonidin, and malvidin were used. The
experimental method was as follows. The extraction ofanthocyanins from leaf and leaf sheath was performed
according to the method reported by Mochioka et al.
(1995), Tsuda et al. (2009), and Chiba et al. (2010),
with some modifications. Brief ly, liquid nitrogen
was added to leaves (300 mg) that had been stored at-30°C and these were then ground using a ceramic
mortar. The ground material was placed into a safetyFig. 2 . Pa n icle t y pe s of S eta r i a pu m i l a: a) a nd b)
d o m e s t i c a t e d t y p e s ( D k ) m i x e d w i t h Pa s p a l u mscrobiculatum; c) domesticated type (Ds) mixed with
Panicum sumatrense and d) weed type shattering seedgrains; e) and f ) cropping fields mixed with Panicum
sumatrense and Setaria pumila: at Illur village near
Chittoor in Andhra Pradesh.

rock tube (1.5 ml), followed by the addition of 300 μl
50% methanol containing 5% acetic acid. The tube
was allowed to stand in a sonicater for 30 min, and thesolution was fi ltered through a membrane fi lter (UST-5,
ADVANTC). The solution of standard pigments was
also filtered through the same membrane filter. Afterthe treatment, the solution was placed in a refrigerator

together as a wild grain for food and forage today.

at 6°C overnight. The anthocyanin composition of

Their spikes show a large variability, for example, in thethe leaves was identified by the retention time and thedomesticated type (Dk) mixed with Pas. scrobiculatum

content was calculated using a calibration curve. The(Fig. 2a and 2b), another domesticated type (Ds) mixed

HPLC system employed to analyze the anthocyaninswith P. sumatrense (Fig. 2c), and the weed type (W) of S.

was a Shimazu system CLASS-M10A equipped with apumila (Fig. 2d)

Shimazu chromatography data station, including pump,

The morphological characteristics and the duration

column oven, and absorbance detector SPD-10A for

to flowering of S. pumila (60 accessions) were compared

monitoring at 520 nm. For the column, Shin-Pak (4.6

among weed, companion weed, mimic companion

× 250 mm, PREP-ODS(H)KIT; Shimazu) was used

weed, and domesticated type mixed with the otherat 40°C. Linear gradient elution was performed withcrops (Fig. 3). The plant height of weed types had asolution A (1.5% phosphoric acid) and solution B (1.5%large range (from 81 to 220 cm), and the height of thephosphoric acid, 20% acetic acid, and 25% acetonitrile)

mimic companion weed and domesticated types weredelivered at a flow rate of 1.0 ml/min as follows: 80%

within the same range. The number of tillers showedof solution A initially; 15% A for the next 40 min;very broad variability from 10 to over 60, but mostly 10

80% A for a further 5 min; and finally 80% A for 15

to 25, irrespective of the type. The spike length was 5 to2min (P.MAX; 200 kgf/cm : P.MIN; 10 kgf/cm ). The

14 cm in weed types, and a little longer (9 cm to over 16

injection volume for the extract was 20 μl.

cm) in mimic companion weed and domesticated types.

2

The duration to flowering of many strains was ranged
Results

from 121 to 140 days, but that of domesticated types

Morphological and ecological diversity of S. pumila

showed a bimodal pattern, 81 to 100 days and 121 to

S. pumila is still cultivated through mixed cropping

150 days. The ratio of length/width of the flag leaf had

mostly with P. sumatrense in restricted regions ina modest range from 21 to 31, but that of domesticatedpeninsular India as shown in Fig. 2e and 2f. The weed

types had a broad variation of range from 21 to 70.

types and mimic companion weed types are also used

The last internode diameter was 1.1 to 2.5 mm in weed

29

14

10
9
8
7
6
5
4
3
2
1
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a

b

9
8

6
5
4
2
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to
10

11
to
15

16
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20

21
to
25

26
to
30

31
to
35

36
to
40

41 46
to to
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56
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0

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7 to 89 to 10 11 to 12 13 to 14 15 to 16 17 to 18

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Tillers

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16

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5
4
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12

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8

Weed

6

Mimic medium

4

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6

4

2

2

2

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0

0
71 to 81to 91 to 101 to 111 to 121 to 131 to 141 to 151 to 161 to 171 to
80 90 100 110 120 130 140 150 160 170 180

f

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Strains

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100 120 140 160 180 200 220 240

6WUDLQV

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10Strains

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民族植物学ノオト 第９号

to 10

11 to 20 21 to 30 31 to 40 41 to 50 51 to 60 61 to 70

0Length/Width ratio of flag leaf

1.1 to 1.5 1.6 to 2.0 2.1 to 2.5

2.5 to 3.0

Diameter (mm)

'XUDWLRQWRIORZHULQJ GD\V

Fig. 3. Morphological and ecological characters of 60 accessions in Setaria pumila: a) plant height (cm), b) number of titters, c)spike length (cm), d) duration of flowering (days), e) length/width ratio of flag leaf, and f) last internode diameter (mm).

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cropping fields (sites)

types, 1.1 to 3.0 mm in mimic companion weed types,

The species components (per m2) of four millet

and 1.1 to 2.0 mm in domesticated types.

cultivation fields are shown in Table 3. At site 1, thepercentages of P. sumatrense, S. pumila, and weedSpecies component of millet and weed grown in four30

民族植物学ノオト 第９号

Fig. 4. Early growing patterns of seedling in S. pumila, P. sumatrense and Pas. scrobiculatum: a/b) plant height, c/d) numberof tillers and e/f) number of leaves on main culm and g) seed germination.

Brachiaria sp. were 59%, 33%, and 8%, respectively,

nearly 100% by 10 days, but the germination of Pas.

while at the nearby site 2, the percentages were 25%,

scrobiculatum (D-pas, W-pasw), Setaria sp. (W-stw)

74%, and trace. At site 3, a single stand of E. coracana,and O. rufipogon (ancestral W-orr) was only starting at

and at site 4, a single stand of P. sumatrense, the weed

5 days and the rates were still increasing at 15 days (Fig.

type of S. pumila, had invaded significantly, but

4g).

represented only 3–5% into both cropping fields.
Mimicry in the color of leaf and leaf sheath
The leaf and leaf sheath colors of S. pumila wereComparison of early growing among companion

the same green (370c~371c) and reddish purple

weeds and crops
With respect to early growing during the period(206c~209c), respectively, using the PA NTONEof 30 to 74 days after sowing, five characteristics were

Formula Guide, as a mixed cropping Pas. scrobiculatum

compared among S. pumila (2, Mk-st56 and Dk-st58)

at site 5 in Karnataka (Table 4 and 5). The colors

and Pas. scrobiculatum (2, Dk-ps1 and Dk-ps2) collectedof S. pumila were also the same green (363c~371c)

at site 5 and 7 in Tumkur, Karnataka, and S. pumila

and reddish purple (206c~208c), respectively, at site

(Ds-st61 and Ws-st62) and P. sumatrense (D-pn3 to

7 in Karnataka. At site 6 in Karnataka, the colors

D-pn6) in Kolar, Andhra Pradesh (Table 2). The earlyof S. pumila and P. sumatrense were the same green

growth of the number of tillers of S. pumila (st58 from(369c~377c) and reddish purple (206c~207c), except for

Karnataka) showed a marked increase, but there was

the leaf sheath color of B. ramosa (green, 374c). Among

little difference in the plant height or the number ofthe control strains used, the colors of P. sumatrense

leaves among the four strains (Fig. 4a-c-e). For theranged from green (366c-370c) to pink (203c), but thosestrains from Andhra Pradesh, the number of tillers of S.

of S. viridis ranged from green (leaf 371c) to purple (leafpumila (st61, st62) was also markedly increased 52 days

sheath 209c-5195c).

after sowing, while there were little difference in plant

The anthocyanin composition of leaf and leaf sheaths

height or the number of leaves among the six strains

was analyzed by HPLC as illustrated in Fig. 5. The leaf

(Fig. 4b-d-f). On the other hand, the seed germination

sheath of S. pumila showed many peaks (present on

rates of O. sativa, B. ramosa, E. coracana, P. sumatrense

each one) on Rt (Fig. 5a), while those of P. sumatrense

and S. pumila were approximately 80% by 5 days, and(Fig. 5b) and Pas. scrobiculatum (Fig. 5c) indicated only31

民族植物学ノオト 第９号

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民族植物学ノオト 第９号

Fig. 5. HPLC analysis and comparison between plant pigmentations through PANTON Formula Guide. a) S. pumila, b) P.
sumatrense, c) Pas. scrobiculatum, d) B. ramosa, e) S. viridis, f) standard samples and g) in a field of Pas. scrobiculatum: ps2,
Pas. scrobiculatum; st58, S. pumila; in a field of P. sumatrense: pn1, P. sumatrense; br, B. ramosa; st57, S. pumila.

a small peak (Rt around 30) except for some occasional

the PANTONE Formula Guide). Sub-cluster IIa (11peaks (around 45.6). However, that of B. ramosa

samples) consisted of the green leaf of S. pumila (1,showed no peak at all around Rt (30) (Fig. 5d). Onlyst57), the green leaf (1, ps1) and leaf sheath (2, ps1 andthe Rt (46.1) of anthocyanin contained in S. viridisps2) of Pas. scrobiculatum, which were reddish purple(dried matter, Fig. 5e) concurred with natural cyanidin’scolor (208/209c), the leaf sheath of P. sumatrense (3,Rt (4 6. 2), but most of the Rts of anthocyanins

pn1-reddish purple, green pn2 and the other), the green

contained in samples did not coincide with the Rt ofleaf sheath of B. ramosa (1, br1) and the green leaf (1)three natural pigments (Fig. 5f). Therefore, cyanidin

and leaf sheath (2) of S. viridis. Moreover, Sub-cluster

was used as an indicator of anthocyanin content. By

IIb (20) consisted of the green leaf of S. pumila (4, st56,

means of cluster analysis (Ward method), these samples

st58, st 59 and st60), Pas. scrobiculatum (1, ps2), S.

were categorized into two groups as illustrated in Fig.

viridis (1), P. sumatrense (8, pn1, pn2 and so on), and B.

6. Cluster I consisted of five samples (st56~st60),

ramosa (1, br1).

which were only the leaf sheath of S. pumila. The

Seed color mimicry was found in the mixed grains

color was reddish purple (198/202/205/206/207c inof P. sumatrense and S. pumila at Illur village near33

民族植物学ノオト 第９号

Fig. 7. Seed mimicry of S. pumila to P. sumatrense and threefoods made from their mixed grains at Jalaripali village,
Andhra Pradesh: a) mixed grains harvested and sold; after
identified them, b) brown seeds of P. sumatrense, c) brown
seeds and d) black seeds of S. pumila; e) annamu (boiled
grain), f) sankati (thick flour porridge) and g) uppitu (semi-solid
porridge).

and insurance crop, used in case of famine, underFig. 6. Dendrogram based on the pigmentation of leaf andleaf sheath in five species.

mixed cropping with Pas. scrobiculatum, E. coracana,and P. sumatrense. The weed types after their invasion

Chittoor in Andhra Pradesh as shown in Fig. 7a. The

into upland rice and millet fields obtained mimicryseed grains of P. sumatrense (Fig. 7b) had a very similar

associated with a particular crop, and made a close weed-

pale brown color to the seed grains of S. pumila (Fig. 7c),

crop complex under the severe weed control measures

except for their black seed grains (Fig. 7d). However,practiced by farmers. In the fourth and fi nal step, theit was possible to distinguish the seed grains of P.

mimic companion weed forms were used not only asumatrense from those of S. pumila in detail, because of

fodder source for cattle, but also as a supplementary

their glossy lemmata. Here, villagers have made threegrain to the main cereal species. In the case of S.

foods from S. pumila. They are boiled grain, annamu

pumila, too strict weeding was avoided as a means(Fig. 7e), flour porridge, sankati (Fig. 7f) and semi-solidof crop insurance in years of extreme drought in the

porridge, uppitu (Fig. 7g).

Deccan Plateau. This possibly led to S. pumila growing
taller with larger spikes and large seeds, accompaniedwith less shattering, and gradually progressing toward

Discussion
The domestication process of S. pumila may havedomestication. Actually, S. pumila has obtained mimicpassed through four steps as illustrated in Fig. 8. The

traits such as long leaf, a few tillers, and tall plant height

first step was a weed that had grown along roadsides

in the field of P. sumatrense as shown in Figs. 3 and 4.

and other unstable habitats and then invaded upland

The morphological and ecological characteristics of therice fields. The second step was an evolutionary process

60 strains in S. pumila were very variable, as shown into obtain an agro-ecological niche out of weed status,

Fig. 3. The plant height (cm) indicated a trend toward

using for fodder, in order to get the companion weed

higher tallness in the domesticated type than in thestatus growing in upland rice and some millet fields.

weed type (Fig. 3a). The number of tillers showed a

The third step was a process of advancing from theslight reduction in the domesticated type compared to

mimic companion weed status to a semi-domesticatedthe weed type, excluding two exceptional strains (Fig.

34民族植物学ノオト 第９号

Fig. 8. Domestication process of Setaria pumila3b). The spike length (cm) clearly increased more in thearid environment. The mimic companion weeds

domesticated type than in the weed type (Fig. 3c). The

were harvested together with other millet, and sown

duration to flowering (days) showed a bimodal pattern:involuntarily again next season. In the third step,

early and late (Fig. 3d). The early domesticated type was

farmers changed their hostile weed control into asimilar to the weed type, while the late domesticatedfriendly one. Recently, during the fourth step, thistype was similar to the mimic companion weed typesituation was followed by mixed cropping. S. pumila ismixed with other cereals. The length/width ratio oftermed a tertiary crop in relation to its associated plants,

the flag leaf showed two trends, wide and slender, as

which were secondary crops, such as P. sumatrense, Pas.

shown in Fig. 3e. The last internode diameter (mm) was

scrobiculatum, with respect to rice.

clearly thick in the domesticated type than in the weed

The species ratio composition of each mixed

type (Fig. 3f). The domesticated type mixed with P.

cropping field was variable annually, according to somesumatrense had a higher ratio than the others, which had

field conditions such as summer precipitation (Table

a medium ratio. It was morphologically clear that spikes

3). At the early growing stage, it was very difficult to

had become longer and the last internode diameter had

recognize morphological differences from each other inbecome bolder in the domestication process.

a set of plant species collected from the same cropping

It was very difficult for farmers to detect thefield. The inter- and intra-specific mimic variation had

dif ference among mimic companion weeds andoccurred not only in the morphological characteristics

domesticated types in sympatric fields during the earlye.g., plant height, leaf size and number of tillers, but also

growing stage, because at this stage they resemble eachin the ecological ones, e.g., seed germination pattern,other very closely. The seed germination of domesticatedseed color, and leaf sheath color. It was clear that theand annual plants was usually rapid and uniform, butleaf sheath color of P. sumatrense and Pas. scrobiculatumthat of Pas. scrobiculatum, perennial plants, and weeds(Sub-cluster IIa) was mimicry related to that of S.

was a little late and irregular.

pumila (Cluster I) as shown in Fig. 5. The glume

At the same time, S. pumila diversified their traits

color of S. pumila was also mimicry of the glume of P.

in totality through hybridization among the types

sumatrense. The domestication process and dispersal ofunder natural and artificial selection in the severely

S. pumila has developed by means of elaborate mimicry35

民族植物学ノオト 第９号

in several characteristics of other crops to itself as shownof other grain crops. The domesticated type of S.

in Fig. 5, 6, and 7.

pumila was always sown, harvested, and consumed

Pas. scrobiculatum is perennial and the seed

together with P. sumatrense as shown in Table 7e, 7f,

germination, tillering, and plant height elongation

and 7g. Farmers called the mixed grains tela samuru

are usually slow. Therefore, Pas. scrobiculatum was(meaning white little millet) in Telugu. The proportion

distinguished relative to the other species. Amongof grains purchased at a local market was 70% of P.

others, because these characters were synchronized tosumatrense and 27% of S. pumila, respectively, in 1997

each other, it was very difficult to distinguish them,(Kimata et al. 2000). These authors are of the opinion

especially plant height and the number of leaves onthat in severe drought, S. pumila provides a reasonable

main culm in the early growing stage. The domesticatedharvest while P. sumatrense might fail completely. This

type of S. pumila was adjustable to its associated mimic

situation recommends the domestication process of

species in morphological characters, ecological traits insecondary crops in other cereal fields against an arid

early growth, and plant pigmentation as shown in Table

climate. These crops are used to make six traditional

4 and Fig. 6. It indicated mimicry of leaf and leaf sheathfoods in total, including ganji (thin flour porridge),by way of color among S. pumila, Pas. Scrobiculatum,

kheer (sweet gruel) and roti (unleavened bread), as aand P. sumatrense, but not B. ramosa. However, bothsupplementary ingredient (Kimata and Sakamoto 1992,

leaf color of S. pumila (Ds) and P. sumatrense were the

Kimata et al. 2000).

same green (371c~377c), while the leaf sheath color of

Furthermore, this domestication process is supported

S. pumila was reddish purple (198c~202c) in spite ofby the linguistic recognition of their various vernaculara finding of green color (206c~207c) in P. sumatrense

names by farmers. For example, a mimic companion

at site 8 in Andhra Pradesh. This case did not indicate

weed or semi-domesticated plant occurring with Pas.

mimicry of leaf sheath color.

scrobiculatum was called varagu korali, meaning justMoreover, S. pumila has been domesticated as

the same as kodo millet, while a mimic companiona tertiary crop, by way of the other millet species,

weed or semi-domesticated plant occurring with P.

to upland rice, because it has built up the strongest

sumatrense was called samuru korali, meaning also

resistance to frequent droughts in the Deccan Plateau.

just the same as little millet. The domesticated typeThe domesticated type of S. pumila is commonly

was called various vernacular names (e.g., korati, kora

cultivated in mixed stands mostly along with P.

samuru, korin) in each local language and region. The

sumatrense in South India today, while the semi-

linguistic differentiation shows a close relationshipdomesticated crop and mimic companion weed types

to the domestication process (Chandra and Koppar

are not only grown with P. sumatrense, but also mainly

1990; Kawase 1987; Kimata et al. 2000, Kimata 2015c;

with Pas. scrobiculatum, E. coracana, and upland rice

Kobayashi 1987, 1989).

in diverse agro-ecological niches in Orissa. The weed

B. ramosa is cultivated under pure single croppingtype of S. pumila grows widely through the India

as a sole tertiary crop, while S. pumila is still cultivatedsubcontinent and tropical and temperate Northern

under mixed cropping with P. sumatrense as a minor

Hemisphere. The various types have adapted to arid

domesticated plant. B. ramosa tolerates drought better

conditions and agro-ecological niches in the Deccan

than S. pumila, and has been undergoing a specializing

Plateau during their distribution from eastern humid

adaptation to arid regions, and has nearly attained

areas to southern dry areas in the Indian subcontinent

the tertiary domesticated phase. On the other hand,(Sehgal et al. 1992).

the landraces of S. pumila have adapted to drier fields

The domesticated types of S. pumila have broadly

in South India than in Orissa. S. pumila was almost

promoted the biocultural diversity through the mimicry

always grown with P. sumatrense, but seemed to grow36

民族植物学ノオト 第９号

Brink. 1983a. Diversity in kodo millet, Paspalum scrobiculatum.

singly when the later fail to grow in severe droughts,

Economic Botany 37:159-163.

as was observed in our survey of 1987. This possiblyde Wet, J. M. J., K. E. Prasada Rao, M. H. Mengesha and D. E.

suggests that S. pumila could become an independent

Brink. 1983b. Domestication of sawa millet (Echinochloa colona).
Economic Botany 37:283-291.
de Wet, J. M. J., K. E. Prasada Rao, M. H. Mengesha and D. E. Brink.
1983c. Systematics and domestication of Panicum sumatrense(Gramineae), Journal d’Agriculture Traditional et BotaniqueApplique 30:159-168.
Fuller, D. Q., R. Korisettar and P. C. Venkatasubbaiah. 2001.
Southern Neolithic cultivation systems: A reconstruction basedon achaeobotanical evidence. South Asian Studies 17: 171-187.
Fuller, D. Q. and M Madella . 2000. Issues in Harappan
achaeobotany: Retrospect and prospect. in Indian Archaeology
in Retrospect, Vol. II. Protohistory. S. Settar and Ravi Korisettar(eds). Publications of the Indian Council for Historical Research,
Manohar, New Delhi, pp317-390.
Kawase, M. 1987. Variation and distribution of millets in South
India. In: S. Sakamoto (ed) A preliminary report of studies on
millet cultivation and its agro-postal culture complex in Indian
sub-continent, I (1985). Kyoto University, Kyoto, Japan, pp5-14.
Kimata, M. 2015b. Domestication process of korati, Setaria pumila(Poaceae), in the Indian subcontinent on the basis of cluster
analysis of morphological characteristics and AFLP markers.
Ethnobotanical Notes 9: 49-64.
Kimata , M. 2015c. Domestication process and ling uistic
differentiation of millets in the Indian subcontinent.
Ethnobotanical Notes 9: 17-31.
Kimata, M. and S. Sakamoto. 1992. Utilization of several speciesof millet in Eurasia. Bulletin of Field Studies Institute, Tokyo
Gakugei University 3: 1-12.
Kimata, M., E. G. Ashok and A. Seetharam. 2000. Domestication,cultivation and utilization of two small millets, Brachiaria ramosa
and Setaria glauca (Poaceae), in South India. Economic Botanycrop.
In this case of multiple polyploidy species, S. pumila
indicated that diversity had been increased mainly bymimicry and inter- and intra-specific hybridizationu nd e r n a t u r a l a nd a r t i f ic i a l s e le c t ion . T h i s
domestication process has been promoted as a meansof imparting adaptation to an arid climate, and also
produced a symbiotic relationship among weeds, othercrops, and farmers, while departing from an antagonistic
one. The domestication process of S. pumila indicatesthe importance of weed-crop complexes and biocultural
diversity as a plant-man symbiosis. The diversity of S.
pumila and its relatives should be conserved both in situ
and ex situ, especially in on-farm conservation sites.
Acknowledgments
The author wishes to express his hearty thanks to theIndian farmers in the areas of study for their valuable
information and kindness; to Dr. A. Seetharam, Dr.
Madhukeshwara and Mr. Mantur, Project Coordination
Cell (Small Millets), ICAR, University of Agricultural
Sciences, Bangalore, India, for their collaboration anduseful suggestion: to Dr. M. Nesbitt and Dr. T. Cope,
Royal Botanic Gardens, Kew, UK, for their valuable

54(2):217-227.
Kobayashi, H. 1987. Mimic and associated weeds with millets
and cultivation methods of millets in the Indian subcontinent.
In: S. Sakamoto (ed.) A preliminary report of studies on millet
cultivation and its agro-postal culture complex in Indian subcontinent, I (1985). Kyoto University, Kyoto, Japan, pp 15-40.
Kobayashi, H. 1989. Mimic and associated weeds with millet
and rice cultivation in Orissa and Maharashtra in India. In:
S. Sakamoto S (ed.) A preliminary report of studies on millet
cultivation and its agro-postal culture complex in Indian subcontinent, II (1987). Kyoto University, Kyoto, Japan, pp. 11-32.
Machioka, R., M. Yamaguchi, S. Horiuchi, H. Matsui and H.
Kurooka. 1995. Chemotaxonomic classification of native wild
Japanese grapes by anthocyanins in berry skin. J.Japan.Soc.Hort.
Sci. 64(3):463-470.
Sehgal, J. L., D. K. Mandal, C. Mandal, and S. Vadivelu. 1992. AgroEcological regions of India. Oxford and IBH Publishing Co.,New Delhi, India.
Singh, H.B., and R. K. Arora. 1972. Raishan (Digitaria sp.) - a

suggestion on excellent collections of literatures andherbarium specimens; and to the late Professor H.
Kobayashi, for his excellent advice during the fieldsurvey in 1985, 1987 and 1989.

References
Achariyar, R. B. K. R. 1921. A handbook of some South Indiangrasses. Government Press, Madras, India, pp 109-113.
Chandra, U. and M. N. Koppar. 1990. Diversity and domesticationof minor millet species in Indian sub-continent. Indian Journal of
Plant Genetic Resources 3(2):47-58.
Chiba, Y., Y. Yamaguchi, K. Hiramoto, S. Yanagi, Y. Saito and
T. Hamana. 2010. Studies on qualitative analysis of natural
food colors in foods. Annual Report of Center for Health andEnvironment, Miyagi Prefecture 28:50-54.
de Wet, J.M.J., K.E. Prasada Rao KE, M. H. Mengesha and D. E.

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民族植物学ノオト 第９号

minor millet of the Kashi Hills, India. Economic Botany 26:376380.
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Tsuda, T., I. Suda and T. Tsushida. 2009. New Aspect of
Anthocyanins Science - Development of Health Benefits andpractical Use -. Kenpakusha, Tokyo (in Japanese).
Vavilov, N. I. 1926. Studies on the origin of cultivated plants. Bull.
Appl. Bot. Plant Breed. (Leningrad), 16(2):1-248.
Weber, S. A. 1992. South Asian Archaeobotanical Variability. In:
C. Jarrige (ed.) South Asian Archaeology 1989, Prehistory Press,
Madison Wisconsin, pp. 283-290.
Zhao, M., H. Zhi, A. N. Doust, W. Li, Y. Wang, H. Li, G. Jia, Y.
Wang, N. Zhang and X. Diao. 2013. Novel genomes and genome
constitutions identified by GISH and 5s rDNA and knotted 1
genomic sequences in the genus Setaria. BMC Genomics 14:244.

38

民族植物学ノオト 第９号

Domestication process of korati, Setaria pumila (Poaceae),
in the Indian subcontinent on the basis of cluster analysis
of morphological characteristics and AFLP markers
Mikio KIMATA
Plants and People Museum

Introduction

The endemic landraces and related weeds werecollected in field surveys around the Deccan in India

Humans have domesticated more than 30 grasssince 1983, to explain the domestication process of

species as grain crops in several parts of the world,

Setaria pumila (Poir.) Roem. et Schult. (Poaceae)

possibly as long as 12,000 years ago. However, several

throug h its mimicr y of other g ra in crops. The

species are threatened and, despite their potential fooddomestication process of Setaria pumila in relation tovalue in their native habitats, have disappeared or havethe weed-crop complex was comparatively investigated

not been extensively cultivated. This is because the yieldusing statistical and AFLP analyses. It was clear on theand production of the three major crops: wheat, rice,basis of these results that the domestication process

and maize, have rapidly increased due to technological

had progressed through the four stages according

innovations in crop-improvement programs. Cultivation

to geographical trends in morphological (artificialof other grain crops (e.g., millets) has decreased

selection) and genetic variation (neutral DNA markers).

gradually during the 20th century, resulting in loss

Under the complex process, the 4 stages were as follows:of genetic diversity of local varieties. It is currently

weed, companion weed, mimic companion weed andnecessary to recognize the value of these neglected

domesticated type to the secondary crop. The paddy

species as exploitable and underutilized genetic

rice had dispersed from Assam, the humid east toward

resources that exhibit adaptability to stress-prone

Deccan, the dry south in the Indian subcontinent.

environments. In this paper, we focus on millet species,

Several species of Indian millet were domesticated bywhich are mostly C 4 plants, are early to mature, and canlocal farmers as the secondary crop of rice along the

be cultivated under conditions of severe drought andclimatic trend and dispersal route. In South India,

harsh sunlight.

one domesticated type of S. pumila was cultivated only

Small-scale farmers continue to cultivate a few

in mixed stands mostly along Panicum sumatrense.

useful local varieties of millet. These indigenous

Around Orissa, the other types and the related weeds

varieties are excellent materials for investigating crop

were grown in the sympatric fields with Paspalum

evolution, particularly the origin and dispersal routes

scrobiculatum, Eleusine coracana, and upland rice (Oryzaof domesticated plants. In the Indian subcontinent, asativa) in diverse agro-ecological niches. Therefore,

few millet species are still undergoing domestication

S. pumila became exactly a tertiary crop to the other(Kimata et al. 2000; Singh and Arora 1972). While crop

Indian millet (secondary crop to rice).

evolution can be reconstructed mostly from botanicaldata, details on geographic origin and dispersal will

Key words: A FLP ma rkers, a r ti f icia l selection,

become clear from information on the basic agricultural

domestication process, mimicry, mixed stand, tertiarycomplex offered by local farmers. This basic agricultural

cropcomplex consists mainly of cultivation, processing andcooking such as biocultural diversity.

39

民族植物学ノオト 第９号

Vavilov (1926) illustrated the domestication process

possibly gathered from the wild (Fuller et al. 2001).

from companion weeds associated with wheat to

The first known occurrences of various cereals in thesecondary crops in two genera, Avena and Secale. For

Harrappan Civilization are reported as wheat, barley,

example, Secale cereale L. acquired strong resistance toand oats in the Early phase (before 2600 BC); Eleusine

cold in high altitude or latitude areas, and subsequently

sp. (problematic, E. coracana), Setaria sp., Panicum sp.

was able to g row under more severe conditionsin the Mature phase (2600-2000 BC); and Paspalum

than those under which wheat can grow. Kobayashisp., Echinochloa sp., Sorghum sp., and Pennisetum sp. in(1987, 1989) proposed an integrating model of thethe Late phase (more recent than 2000 BC) (Fuller anddomestication of Indian millet (e.g. P. sumatrense,

Madella 2000; Weber 1992).

Echinochloa frumentacea) as a secondary crop from

Many new techniques using DNA markers,

mimic companion weeds associated with Oryza sativa

including SSR (simple sequence repeat), RAPD (random

L. Farmers have manipulated the domestication process

amplified polymorphic DNA), R FLP(restriction

by selecting for desired growth, visual, and palatability

fragment length polymorphism), and AFLP (amplifiedtraits, e.g. yield, early maturation, color, sugar

fragment length polymorphism analysis), have been

content. However, natural selection and hybridization

conducted for the genus Setaria (Benabdelmouna et al.

have occurred among closely related weeds during

2001; d’Ennequin et al. 2000; Fukunaga et al. 2002;

domestication.

Lin et al. 2012). Intraspecific polymorphic variability

The growing area of O. sativa expanded fromrevealed with RAPD and RFLP marker systems waswetlands to establish secondarily in uplands in the

negligible. AFLP has gained wide acceptance for

Indian subcontinent. In turn, weedy ancestral plants

enabling a high degree of resolution and reproducibility

invaded paddy and upland rice fields. Local farmers

in genetic analysis (Lakshmi et al. 2002). AFLP has asubsequently domesticated Panicum sumatrense Roth.

number of other relevant applications and advantages(little millet), Paspalum scrobiculatum L. (kodo millet),for analysis of plant genomes in general. A large number

and Echinochloa frumentacea Link (Indian barnyardof DNA loci can be assayed in each reaction, and a large

millet), as secondary crops, because these species

number of fragments can be assayed with a relatively

demonstrated stronger resistance to drought thansmall number of primers. Intergeneric polymorphism

upland rice in Eastern India. Several additional species

revealed by AFLP markers was very high (94.4%). At

of millet were domesticated in this region: Brachiaria

inter-specific level, it was not significant enough AFLP

ramosa (L.) Stapf. (korne, browntop millet), Digitaria

analysis recorded a higher level of variation, 66.5%,

cruciata [Nees] A. Camus (raishan), and Setaria pumila

between Panicum miliaceum and P. sumatrense (Bai(Poir.) Roem. & Schult. (korati, yellow foxtail millet;

et al. 1999). Information on intraspecific diversity andsyn. Setaria glauca [L.] P. Beauv.) (Chandra and Koppar

species relationships could form the basic foundation

1990; de Wet et al. 1983a, b, c).

for further research on crop-improvement programs(Lakshmi et al. 2002).

Recently, archeological studies in the Indian
subcontinent have provided usef u l data on the

GISH (Genomic in situ hybridization) patterns

ancient history of various grains. Millet materials

revealed that two diploid species (2n = 18), S. viridis andwere identified from two archaeological levels in the

S. italica, bore genome AA and a tetraploid species (2nSouthern Neolithic chronology: Phase II (2300–1800= 36), S. verticillata, had genome AABB. The genomic

cal BC) and Phase III (1800-1200 cal BC). These

composition of S. pumila (polyploid species, 2n = 18,

materials were identified primarily as two species, B.

36, 72) was unknown (Benabdelmouna et al. 2001).

ramosa and Setaria verticillata (bristly foxtail millet-

S. pumila is a cosmopolitan weed distributed

grass). S. pumila was present in limited quantity,

worldwide. This weed grows sympatrically on roadsides,

40

民族植物学ノオト 第９号

uplands, and the levees of lowlands; four intraspecifictypes of S. pumila have been identified based on
ecological habit: weed type (W), companion weed typeaccompanied by crops (Wx), mimic companion weed
type accompanied by crops (Mx), and domesticatedtype mixed with crops (Dx). Kimata et al. (2000
and 2015a) have shown the biocultural diversityof morphological and ecological characteristics inBrown
panicleS. pumila, and the intraspecific differentiation ofWhite
paniclevernacular names (linguistic data). The present paper
concerns the domestication process of S. pumila, whichis related ecologically to weeds and several grain crops
in the Indian subcontinent, based on cluster analysis ofmorphological characteristics and AFLP markers.
Materials and Methods
Many local varieties and relative weeds of Setaria
pumila (Poir.) Roem. & Schult. (syn. S. glauca [L.]P. Beauv.) have been col lected from the Indian
subcontinent since 1983 in field surveys (Fig. 1).
Concentrated surveys were conducted in Karnataka,
Andhra Pradesh, and Orissa (Fig. 2). At the same time,
accompanying millet and weed species were examined

Fig. 1. Fields of Setaria pumila mixed with Pas.
scrobiculatum (a) and with P. sumatrense (b) in
South India, and spikes of S. pumila (c): weed;
mimic companion weeds; Ds-K, domesticated typemixed with P. sumatrense, and Dk-K mixed with
Pas. scrobiculatum in Karnataka.

in five plots (1 m2) in each of four typical cropping fields(sites) using the quadrat method. Voucher herbarium
specimens and grain samples were collected along thesurvey route and deposited at Tokyo Gakugei University(Tokyo, Japan) and University of Agricultural Sciences(Bangalore, India). Information on agricultural

sowing, germinated plants were transplanted into

practices, grain processing, food preparation, andthe greenhouse, with 30-cm row spacing and 15 cm

vernacular plant names was gathered from local farmers.

between plants. Chemical fertilizer (N:P:K = 8:8:5) was

The experimental strains (n = 78, Table 1) were

supplied at 100 g·m-2. The following parameters of five

selected from these accessions and grown in thetypes of S. pumila were measured at the each full-ripe

greenhouse at Tokyo Gakugei University to compare

stage: number of tillers, plant height , length and width

their morphological and ecological characteristics . Inof spike, length and width of flag leaf, last internode

addition, three relative species of S. pumila: S. italica (ndiameter, and duration to flowering. These types were

= 6, from Japan), S. viridis (n = 2, from Kazakhstan andthree weed types; W, Wx, Mx associated with otherUzbekistan), and S. verticillata (n = 3, from India) were

grain crops, and two domesticated types; Dx mixed

grown using the same methods.

with Paspalum scrobiculatum and Panicum sumatrense.

Ten grains each of 60 strains were sown in a

The lowercase character “x” indicates a main crop mimic

seeding box with row spacing of 8 cm and seed spacingof S. pumila as follows: “p” (paddy, O. sativa L.), “k”

of 2 cm on early June 6, 2002 . Two weeks after(kodora, P. scrobiculatum), “s” (samai, P. sumatrense),41

民族植物学ノオト 第９号

Fig. 2. Sites from which S. pumila and the related species were collected in the Indian subcontinent.

and “o” (others, e.g., Eleusine coracana). These data

The supernatant (approximately 700 μl) was transferred

were analyzed statistically using partial correlation

to a new tube containing 700 μl of chloroform/isoamyl

coefficients and hierarchical cluster analysis (Wardalcohol (24/1), and the tubes were shaken gently for 5

method) by SPSS version 21 (IBM Corp).

min and then centrifuged for 10 min at 10,000 g. The

Ten grains of 78 strains were sown by the same

supernatant (approximately 650 μl) was transferred tomethod above-mentioned on Oct. 4, 2007. DNA

a new tube containing 700 μl of chloroform/isoamyl

extraction was performed on young leaf tissue groundalcohol, and the tubes were shaken for 5 min and thenin liquid nitrogen and incubated in 1.5-ml tubes

centrifuged for 10 min at 10,000 g. The supernatantcontaining 0.5 ml of buffer A for 10 min at 60 ºC using(approximately 600 μl) was transferred to a new tube

CTAB (hexadecyl-trimethyl-ammonium bromide)containing 700 μl of isopropanol to precipitate DNA for

methods (Murray and Thompson 1980). Buffer A

20 min. DNA pellets were collected by centrifugation

contained 2% CTAB, 1% PVP (polyvinylpyrrolidone),at 14,000 g for 10 min, and then were washed in 70%

1.4 M NaCl, 0.25 M sucrose, 0.2% 2-mercaptoethanol,

ethanol, dried twice, and resuspended in 30 μl TE buffer

20 mM EDTA, and 100 mM TRIS-HCl (pH 8.0);(1×) containing RNAse at 4 ºC for 3 to 4 d, followed byultra pure water was added to bring the solution to

storage at –20 ºC. TE buffer (800 μl) was added to 16 μl

100 ml. We then added 0.5 ml of buffer B into theof RNAse (10 mg/ml).

tube, suspended the mixture and then centrifuged
AFLP procedure

it for 10 min at 10,000 g. Buffer B contained 2%
CTAB, 1% PVP, 1.4 M NaCl, 1.1 M sucrose, 0.2%

The AFLP procedure was performed according

2-mercaptoethanol, 20 mM EDTA, and 100 mM TRIS-

to Applied Biosystems (2005), Bai et al. (1999), andHCl; water was added to bring the solution to 100 ml.

Suyama (2001) with some modifications. Briefly, 1 μl of42

民族植物学ノオト 第９号

Table 1. Materials used of Setaria pumila
6DPSOHQR
0DLQFURSDQGUHPDUNV6WDWXV' V$
ųƄƑƌƆƘƐ ƖƘƐƄƗƕƈƑƖƈ P L[HGZ LWKŨƏƈƘƖƌƑƈ ƆƒƕƄƆƄƑƄ
: V0ųőŃƖƘƐƄƗƕƈƑƖƈ
: V0ųőŃƖƘƐƄƗƕƈƑƖƈ
' V0ųőŃƖƘƐƄƗƕƈƑƖƈ
: 0
QRQH
: V0ųőŃƖƘƐƄƗƕƈƑƖƈ
: 0ŲƕƜƝƄŃƖƄƗƌƙƄ
: 0ŶƈƗƄƕƌƄŃƌƗƄƏƌƆƄ
: 0ŶőŃƌƗƄƏƌƆƄ
0 V2ųőŃƖƘƐƄƗƕƈƑƖƈ
0 V2ųőŃƖƘƐƄƗƕƈƑƖƈ
0 V2ųőŃƖƘƐƄƗƕƈƑƖƈ
0 V2ųőŃƖƘƐƄƗƕƈƑƖƈ
: VN2ųő ƖƘƐƄƗƕƈƑƖƈ P L[HGZ LWKųƄƖƓƄƏƘƐ ƖƆƕƒƅƌƆƘƏƄƗƘƐ
: V2ųőŃƖƘƐƄƗƕƈƑƖƈ
: 2QRQH
: V2ųőŃƖƘƐƄƗƕƈƑƖƈ
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: V2ųőŃƖƘƐƄƗƕƈƑƖƈ
' V2ųőŃƖƘƐƄƗƕƈƑƖƈ
' V2ųőŃƖƘƐƄƗƕƈƑƖƈ
' V2ųőŃƖƘƐƄƗƕƈƑƖƈ
: V2ųőŃƖƘƐƄƗƕƈƑƖƈ
0 N2ųƄƖőŃƖƆƕƒƅƌƆƘƏƄƗƘƐ
' N2ųƄƖőŃƖƆƕƒƅƌƆƘƏƄƗƘƐ
0 N2ųƄƖőŃƖƆƕƒƅƌƆƘƏƄƗƘƐ
: V2ųőŃƖƘƐƄƗƕƈƑƖƈ
0 V2ųőŃƖƘƐƄƗƕƈƑƖƈ
: 2
QRQH
: 2QRQH
: V2ųőŃƖƘƐƄƗƕƈƑƖƈ
: V2ųőŃƖƘƐƄƗƕƈƑƖƈ
0 V2ųőŃƖƘƐƄƗƕƈƑƖƈ
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43

民族植物学ノオト 第９号

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each genomic DNA (was digested with 0.3 μl of EcoRI

associated with MseI (M+CAG; M+CTG; M+CTA;

adapter (TAK AR A), 0.3 μl of MseI adapter (New

M+CAT; M+CAA). Five microliters of amplification

England), 2.0 μl of 10× reaction buffer H, and 7.4 μl of

products were loaded onto a 5.75% denaturing

H 2O in a fi nal volume of 20 μl for 3 h at 37 ºC. Afterpolyacrylamide gel (LONZA) and electrophoresed inincubation, ligation of adapters corresponding to the

1× TBE for 1 h. Bands were detected using the silver

sticky ends of both enzymes was performed. Adapter

staining protocol described by Cho et al. (1996).

mix (EcoRI, MseI, 2.0 μl), 0.5 μl of T4 DNA ligase
Data analysis(TAKARA), 1.5 μl of 10× reaction buffer, and 3.5 μl
H 2O were added to 7.5 μl of the digested DNA. The

The bands were detected on the gel at the finest level

resulting reaction mix was incubated for 5 min at 95 ºC.

of sensitivity by Lane Analyzer (ATTO), the raw data

A preselective amplification step using non-selective

were adjusted, and then the visible and reproducible

primers was then performed in a total volume of 20

bands were scored for accessions as present (1) or

μl containing 15 μl of the AFLP Amplification Coreabsence (0). The dendrogram of the AFLP markers

Mix, 1 μl of preselective primer pair (Eco A/Mse C),

was constructed using the neighbor-joining method

and 4 μl of ligation mixture (diluted 5 times in 10

and bootstrap analysis (PAUP* version 4.0) on all datamM Tris buffer, pH 8.0). The core mix contains all of

matrices (Nei and Kumar 2000).

the components necessary to amplify modified target
sequences, e.g. buffer, nucleotides, and AmplTaq DNA

Results

polymerase. The polymerase chain reaction (PCR)

Morphological characteristics

program was the same as that described by AFLP Plant

Results of statistical analyses of partial correlation

Mapping Protocol (Applied Biosystems 2005) using a

coefficients of the ten characteristics (number of tillers,

PCR thermal cycler (TAKARA TP3000). Selectiveplant height, length (pl) and width (pw) of spike, theamplifications were performed in a 20-μl final volume

ratio of pl/pw, length (fll) and width (flw) of flag leaf,

containing 3 μl of pre-amplification products (diluted in

the ratio of fl l/flw, last internode diameter, and duration

10 mM Tris buffer) with primers having three additional

to flowering, are shown in Table 2. Those characteristics

3′nucleotides. Amplification reactions were performed

have been strongly affected by artificial selection duringaccording to the same protocol. Five primers associatedthe domestication process. The controlled variables werewith EcoRI (E+A AC; E+A AG; E+AGG; E+ACT;

seed size and seed shattering in this analyses. Statistical

E+ACA) were used in combination with 5 primers

significance at the 1% level was found for the following44民族植物学ノオト 第９号

S. pumila

6Ws-M
22Ds-O
9W-M

Ia

42Ws-O
8W-M
11Ms-O
10Ms-O

I

57

74

60Ds-A
59Ds-A
4Ds-M

Ib

79

} S. verticillata

57Ds-K
1Ds-A

S. pumila W-P

23Ds-O
20Ws-O
53Mk-O
52Ms-O

53

S. viridis
52

47W-O
25Ws-O
51Ws-O
50Mk-O

64

49Mk-O
56Dk-K
48W-O

IIa

II

92

2Ws-M
46W-O
44Mk-O

IIIb

IIIc

28Mk-O
29Ws-O
32W-O
43Wk-O
16W-O

Fig. 4. Dendrogram of neighbor-joining method based on
AFLP markers of genus Setaria

36Ws-O
19Ws-O
18Ws-O
35Ws-O
15Ws-O

last internode diameter (0.882) to width of flag leaf;

45Mk-O
7W-M
38Mp-O

flag leaf (-0.508), and the duration to flowering (-0.561)

plant height (-0.517), length of flag leaf (0.720), width ofto the ratio of length/width of the f lag leaf; spike
length (0.716) and width of flag leaf (0.882) to the last

26Mk-O
12Ms-O
31W-O

internode diameter; and length of flag leaf (-0.544) andthe ratio of length/width of the flag leaf (-0.561) to the

37Mp-O
24Ds-O

20

15

10

5

S. verticillata} S. pumila ssp. pallide-fusca

63

5W-M

27Dk-O
39Mp-O
34Ws-O

25

S. italica

74

S. pumila Do-A
S. pumila W-A
S. pumila Wo-A

21Ms-O
17Ws-O

IIIa
III

59

3Ws-M
58Dk-K
13Ms-O

IIb

Ws-O
S. pumila Ds-T
S. pumila W-A
S. pumila W-P
S. pumila Dsk-K
S. pumila Do-T
}S. pumila Ms-T

duration to flowering. There were no significant (p <00.01) correlations between the number of tillers and thelast internode diameter.

Fi g . 3 . D e nd ro g r a m of c lu s t er a n a l y si s b a s e d onmorphological characteristics.

Cluster analysis of six morphological characteristics(number of tillers, plant height, spike length, length

results: ratio of length/width of flag leaf to plant height

and width of f lag leaf, and f lag leaf length/width(-0.517); ratio of spike length/width (0.739), length ofratio) and the duration to flowering are illustrated inflag leaf (0.664), width of flag leaf (0.584), and diameter

Fig. 3. Using the Ward method, 60 accessions wereof last internode (0.716) to spike length; spike lengthcategorized into three clusters and several sub-clusters.

(0.739) and length of flag leaf (0.704) to the ratio ofCluster I contained sub-clusters Ia and Ib. Subcluster

spike length/width; spike length (0.664), the ratio of

Ia (7 accessions) included weed type (W2); companion

spike length/width (0.704), the ratio of length/width

weed type (Ws1) from Maharashtra; companion weedof the flag leaf (0.720), and the duration to flowering

type (Ws1); mimic companion weed type (medium,(-0.544) to length of flag leaf; spike length (0.584), theMs2); and domestication type mixed with samai

ratio of length/width of the flag leaf (-0.508), and the(P. sumatrense, Ds1) from Orissa. Sub-cluster Ib

45

民族植物学ノオト 第９号

7DEOH$)/3DQDO\VLVRIDFFHVVLRQVRIŶƈƗƄƕƌƄŃƓƘƐƌƏƄXVLQJFRPELQDWLRQVRISURPHUSDLUV(FR5,0 VH, 7RWDOQRRIEDQGV$$& & $*$& 7& $7$* * & 7$$$* & 7*
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2Ws-M
3Ms-M

66

[I]
4Ds-M
5W-M
7W-M
8W-M
6Ws-M
9W-M

[IIa]

10Ws-O[IIb]
11Ws-O
13Ms-O
15Ws-O
14Wsk-O
19Ws-O
18Ws-O[IIc]
20Ws-O
22Ds-O
23Ds-O
24Ds-O[IId]
25Ws-O
26Mk-O
28Mk-O
29Ws-O
32W-O
30Ms-O
27Dk-O
33Ws-O[III]
34Ws-O
35Ms-O[IVa]
36Ws-O
37Wp-O
39Mp-O
40Ws-O
41Ws-O
43Wk-O
42Ws-O[IVb]
49Mk-O
44Mk-O
53Mk-O
50Mk-O
51Ms-O
52Ms-O
45Mk-O
46W-O
47W-O
48W-O
56Dk-K
57Ds-K
58Dk-K
59Ds-A
60Ds-A
66Ds-A
71Do-A
72Do-A
73W-A

80

64

86
65
57
66
62

53
61

3HUFHQWSRO\P RUSKLVP

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

78

66

[IVc]
[Va]
[Vb]
[Vc]

75W-A
81W-K
82W-K
76W-P
77Dsk-K
84Do-T
85Ms-T
86Ms-T
762W-P

96
78
92

83

12Ms-O
16W-O
17Ws-O

61W-U
69Ws-O
63Ws-A
64Ws-A
70Ds-T

Fig. 5. Dendrogram of neighbor-joining method based on AFLP markers of S. pumila.

46

[Vd]

[Ve]

[Vf]

[VI]

民族植物学ノオト 第９号(5 accessions) included Ds5 from Andhra Pradesh

pallide-fusca (2) from Karnataka and S. verticillata (1)(3), Karnataka (1), and Maharashtra (1). Cluster IIfrom Andhra Pradesh formed a cluster. W1 of S. pumila

contained sub-clusters IIa and IIb. Sub-cluster IIa (17from Pakistan was located as the neighbor of S. viridis.

accessions) included: Ds1, Ms2, Mk4, Ws3, and W3

S. verticillata (2) was located in the cluster of S. pumila,from Orissa; Dk2 from Karnataka; and Ws2 frombut S. viridis (3) and S. italica (8) were located in the

Maharashtra. Sub-cluster IIb (2 accessions) included

same cluster. The location of species within clusters was

Ms1 and Ws1 from Orissa. Cluster III contained sub-not significant at p ≤ 0.05 based on the bootstrap test,

clusters IIIa–c. Sub-cluster IIIa (1 accession) comprised

but the species were clearly categorized.

W1 from Maharashtra. Sub-cluster IIIb (10 accessions)S. pumila (72 accessions) were divided into six

included Mk1, Ws6, Wk1, and W2 from Orissa. Sub-clusters including 16 sub-clusters based on AFLP

cluster IIIc (11 accessions) included Ds1, Dk1, Ms1,

marker data as shown in Fig. 5. Cluster I contained Ws1

Mk2, Mp3, Ws1, and W1 from Orissa, and W1 fromand Ms1 from Maharashtra; Cluster III consisted of

Maharashtra. The "W" type of S. pumila was distributed

Ws2; and Cluster VI contained W1, Ws1, and Ms1 fromaround the Indian subcontinent as a cosmopolitan

Orissa. These clusters did not containea domesticated

weed.

type.
Cluster II (4 sub-clusters, 23 accessions) consistedof sub-cluster I Ia (6), W4 , Ws1, and Ds1 from

Variation of AFLP markers
The results of AFLP on 72 accessions from theMaharashtra; sub-cluster IIb (6), Ws (4), Ms1, andIndian subcontinent are shown in Table 3. Most

Wsk1 from Orissa; sub-cluster IIc (4), Ws2 and Ds2

bands showed polymorphic more than 81.7% tofrom Orissa; and sub-cluster IId (8), W1, Ws2, Ms1,

94.1% polymorphisms, excluding the main bands

Ds1, Mk2, and Dk1 from Orissa. Cluster IV (three sub-were detected more than 70% of all accessions. Eachclusters, 18 accessions from Orissa) consisted of sub-combination of EcoRI and MseI detected from 59 to 76

cluster IVa (7), Wp1, Mp1, Wk1, Ws3, and Ms1; sub-fragments based on the four primer pairs selected, andcluster IVb (7), Ws1, Ms2, and Mk4; sub-cluster IVc (4),were used for analysis as follows: E+AAG/M+CTG,

W3, and Mk1. Cluster V (6 sub-clusters, 23 accessions)

E+AGG/M+CTA, E+ACT/M+CAT, and E+AAC/

consisted of sub-cluster Va (1), Dk1 from Karnataka; Vb

M+CAG. Those primer combinations revealed 263(3), Ds1, Dk1 from Karnataka, and Ds1 from Andhra

visible polymorphic bands. The dendrogram accounted

Pradesh; Vc (5), W1, Do2 and Ds2 from Andhra

for accessions of related species on one hand (Fig. 4) andPradesh; Vd (3), W1 from Andhra Pradesh and W2

the accessions of S. pumila on the other (Fig. 5).

from Karnataka; Ve (6), W2 from Pakistan, Ms2, Dsk1,

The diversity of AFLP markers was compared amongand Do1 from Tamil Nadu; Vf (5), W1 from Utter

relative species (28 accessions) of S. pumila (14 including

Pradesh, Ws1 from Orissa, Ws2 from Andhra Pradesh,ssp. pallide-fusca 2), domesticated S. italica (8 fromand Ds1 from Tamil Nadu.

Japan), the ancestral weed S. viridis (3 from Central
DiscussionAsia), and the weed S. verticillata (3 from India). The
dendrogram constructed with the neighboring-joint

The domestication process for each species was a

method is illustrated in Fig. 4. The clusters of S. pumila

complex combination of natural and artificial selection,

were composed, successively, of Ws1 from Orissa; Wo1,

mimicry, hybridization, and polyploidy. Pioneer farmers

Do1, and W1 from Andhra Pradesh; Ds1 from Tamilrequired plants some to have some degree of tolerance

Nadu; and W1 from Andhra Pradesh. The other clusters

to conditions (e.g., cold, hot, drought, harsh sunlight).

included W1 from Pakistan, Dsk1 from Karnataka,

Farmers continue to gather wild cereals in dry areas

and Do1 and Ms2 from Tamil Nadu. S. pumila ssp.

of Africa and the Indian subcontinent. For example,

47

民族植物学ノオト 第９号

Secale cereale L. has acquired strong resistance to cold

During the evolutionary process from companion

in high altitude or latitude areas, and farmers have been

weed to secondary crop, which involved morphologicalable to grow S. cereale mixed with wheat as a secondary

mimicry of other species (Mo), S. pumila (Ds) became

crop as a companion weed under severe conditions

a slender-type mimic with long-narrow leaves as in(Vavilov 1926). Kobayashi (1987, 1989) proposed an

P. sumatrense, while S. pumila (Do) became a thick-integrated model of the domestication process of several

type mimic with wide leaves as in Pas. scrobiculatum

millet species as secondary crops derived from weeds

and other species. Based on the PANTONE Formulaby mimicking companion weeds associated with Oryza

Guide (Pantone Inc.), it was clear that the leaf, leaf

sativa in the Indian subcontinent.

sheath, culm, and glume of S. pumila exhibited mimetic

Increasing the size and shattering resistance in seeds

coloration among species and demonstrated mimicry

are important factors in the domestication process. The

of coloration of P. sumatrense and Pas. scrobiculatum,

partial correlation coefficients that describe control

according to anthocyanin composition revealed byof seed size and shattering (Table 2) explain that the

HPLC analysis (Kimata 2015a).

cylindrical spike has become longer, the last internode

From the cluster analysis, S. pumila cluster I clearly

diameter of the main culm has thickened, and the flag

showed that the domestication process of S. pumila

leaf has widened for effective photosynthesis as a result

has occurred continuously in fields of P. sumatrenseof artificial selection by farmers.

and other grain crops around the Deccan. Cluster II

The low coefficient between the number of tillers

consisted of sub-cluster IIa; Ds1, Ms2, Mk4, Ws3 andand the other characteristics reveals that the number

W3 from Orissa, Dk2 from Karnataka, Ws2 fromof tillers in Dk has decreased during domestication by

Maharashtra, and sub-cluster IIb; Ms1 and Ws1 fromprocesses such as mimicry of Pas. scrobiculatum, while

Orissa. Cluster II gives an example of the domestication

the number of tillers of Ds has increased as the mimicry

process by mimicry, which S. pumila has become Dsk

of P. sumatrense. Separate selection processes functioned

mixed with P. sumatrense and Pas. scrobiculatum in eachto both decrease and increase the number of tillers

fields. Cluster III revealed that the W type of S. pumila(Kimata 2015a). The low coefficients for length of flag

was distributed around the Indian subcontinent as aleaf and ratio of length/width of the flag leaf to duration

cosmopolitan weed.

of f lowering indicate that artificial selection has

The domestication process of S. pumila, Dsk, hasoperated on the flag leaf, causing it to become narrower

taken a route from weed type to companion weed

and to mature early under domestication.

and then to mimic companion weed with O. sativa,

The negative correlation between ratio of length/

P. sumatrense, and Pas. scrobiculatum in Orissa.

width of the f lag leaf to plant height demonstrated

Therefore, the domestication process of S. pumila hasthat the f lag leaf has become longer and narrower,

moved forward as follows. First, the mimic companion

while plant height has increased, as in Ds. The Ds of

weeds (M ks, mainly in Orissa) and second, theS. pumila matures early and has a relatively long anddomesticated type (Do) evolved and moved south tonarrow flag leaf due to artificial selection, reflected in

the Deccan Plateau via Andhra Pradesh. After that,the significant negative correlations between length ofthe domesticated type progressed from Dk to Ds inflag leaf and the ratio of length/width of the flag leaf in

Karnataka and Tamil Nadu.

relation to duration to flowering. In addition, Ds has

The natural intraspecific hybrids of S. pumila

acquired a relatively long and narrow flag leaf as a result

occurred continually in sympatric fields among weeds,of taller plant height, as seen in the significant negative

companion weeds, mimic companion weeds, andcorrelation between plant height and length/width ofdomesticated types, as revealed by a geographic bias inflag leaf.

both morphological characteristics and AFLP markers

48

民族植物学ノオト 第９号

Hybridization

InterspecificIntraspecific

Mimicry

Weed

Mimicry

MimicCompanion
Weed

Crop
Mixed

With kodo, samai
and others

With samai(little millet)

Companion
Weed
With paddy

W

Wx

Farmers

Mx

Do, Dk < Ds

Insurance
crop

FodderSymbiotic

Antagonistic
Natural selection

Food

Artificial selection

Fig. 6. Domestication process of S. pumila in relation to the weed-cropcomplex.

(Figs. 3 and 5). There were two cases of mimicry,

fusca from Karnataka and S. verticillata from Andhra

interspecific and intraspecific, in this domestication

Pradesh made a cluster. The irregular positions in which

process. Interspecific mimicry was found in the mimic

S. pumila was located related to its multiple ploidy levels

companion weed with O. sativa, Pas. scrobiculatum,and obscure genomic constitution.

and P. sumatrense and other species. The intraspecific

The AFLP methodology gave highly reproducible

mimicry occurred by continuous natural hybridization

bands, and polymorphisms among individuals within

bet ween weed a nd domesticated t y pe a nd a lso

accessions were very low (d’Ennequin et al. 2000).

though natural or artificial selection by farmers. The

Small millet species including S. pumila have shown

mimic companion weed type was quite similar to theremarkable genetical variation (Lakshmi et al. 2002)

domesticated type, but farmers were able to differentiate

because of its polyploidy and natural hybridization.

between the two by seed shattering.

The AFLP variation in S. pumila was generally highThe diversity of AFLP markers was compared among

because of the grouping of many sub-clusters, but

relative species of S. pumila, domesticated S. italica,

the bootstrap values were low in each sub-cluster.

the ancestral weed S. viridis, and another weed, S.

Intraspecific morphological differentiation was easily

verticillata (see dendrogram in Fig. 4). Recently, Wang

detected, but the variation in AFLP was reduced by

et al. (2009) and Zhao et al. (2013) indicated that thenatural hybridization. Therefore, based on the AFLP

genome constitution of S. verticillata had diploid (BB)dendrogram, which was not directly inf luenced byand tetraploid (AABB) forms based on GISH, while S.

the artificial selection by farmers, it was obvious thatglauca (syn. S. pumila) was identified genome ‘D,’ butthere was a regional bias; many accessions of mimic

its genomic constitution was not known. Based on thecompanion weed type were located in sub-cluster IVb,

dendrograms by Bayesian analyses for 5s rDNA andand the most accessions of the domesticated type were

kn1 sequences, the A genome included S. italica, S.

in Cluster V with little significance in bootstrap value.

viridis, and S. verticillata; the B genome comprised S.

Moreover, Cluster IV, from Orissa only, did not include

verticillata, and the D genome consisted of S. glauca (syn.

the domesticated type but contained the most accessions

S. pumila). S. pumila (W-P) from Pakistan was locatedof mimic companion weed type. Cluster II fromthe neighbor of S. viridis in Fig. 4. S. pumila ssp. pallide-

Maharashtra and Orissa indicated that the companion/49

民族植物学ノオト 第９号

mimic companion weeds coexist with the domesticatedas P. sumatrense and Pas. scrobiculatum, with respect totype mixed with other crops.

rice. The domestication process of S. pumila, a tertiary

The domestication process of S. pumila may havecrop mixed with other grain crops, proceeds from inter-

passed through four steps as illustrated in Fig. 6. The

and intraspecific mimicry by natural and artificial

first step consisted of a weed that had grown along

selection in sympatric fields. Th is process has occurredroadsides and other unstable habitats moving to invade

by adaptation to aridity as a result of the spread ofupland rice fields. The second step was an evolutionary

S. pumila from the east to the south in the Indianprocess of obtaining an agro-ecological niche as use

subcontinent.

for fodder, to attain companion weed status in upland
rice and millet fields. The third step was a process ofReferences
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Bai G, M. Ayele, H. Tefera, H. T. Nguyen. 1999. Amplified
Fragment Length Polymorphism Analysis of Tef [Eragrostis tef(Zucc.) Trotter]. Crop Sci 39: 819-824.
Benabdelmouna, A., M. Abrirached-Darmency, and H. Darmency.
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archaeobotany: Retrospect and prospect. in Indian Archaeology
in Retrospect, Vol. II. Protohistory. In: S. Settar and Ravi
Korisettar (eds) Publications of the Indian Council for Historicaladvancing from mimic companion weed status to asemi-domesticated insurance crop in case of famine,
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control measures practiced by farmers, weeds evolved
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fourth step, mimic companion weeds were used as botha fodder source for cattle and as a supplementary grain
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overly strict weeding was avoided as a means of crop
insurance in years of extreme drought in the Deccan.
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has obtained mimetic traits such as a long leaves, a few
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environments. Mimic companion weeds were harvested
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again involuntarily the following season. Recently, atthe fourth step, this situation was followed by mixed
cropping. S. pumila is termed a “tertiary crop” inrelation to its associated plants, secondary crops such

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民族植物学ノオト 第９号

http://www.biomedcentral.com/1471-2164/14/244. Accessed 10

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August 2013

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millet of the Kashi Hills, India. Economic Botany 26:376-380.
Suyama, Y. 2001. AFLP analysis, In: The Society for the Study ofSpecies Biology (ed.), Molecular Ecology of Woody Species (inJapanese). Bun-ichi Sogo Shuppan Co., Tokyo, pp 251-262.
Vavilov, N. I. 1926. Studies on the origin of cultivated plants. Bull.
Appl. Bot. Plant Breed. (Leningrad), 16(2):1-248.
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Jarrige C (ed.). South Asian Archaeology 1989. Prehistory Press,
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244.

51

民族植物学ノオト 第９号

Domestication and dispersal of Panicum miliaceum L. (Poaceae)
in Eurasia
Mikio KIMATA
Plants and People Museum

Introduction

Common millet (Panicum miliaceum L.) was themost important grain crop in Eurasian civilization for

The genus Panicum (Poaceae) consists of about 500

several thousand years starting from the Neolithic era.

species. These species have been used for wild grains,

It is still cultivated and has various uses around thefodder, and medicine up to the present day, while three

world. Although it is one of the oldest domesticatedcereals, P. miliaceum L. (common millet), P. sumatrense

plants in Eurasia, the ancestral plant and place of origin

Roth. (samai, little millet), and P. sonorum Beal. (saui,

have yet to be definitively determined. A series ofpanic grass), were domesticated in different places andstudies was started based on the plant’s morphologicaltimes. These are C4 plants endowed with strong drought

and ecological characteristics, followed by studies ofresistance, early maturation, and high nutrient content.

its genetic characteristics and secondary compounds,

P. sumatrense was domesticated from an ancestralto elucidate its domestication process and dispersal inspecies, P. sumatrense subsp. psilopodium, after aroundEurasia. Accessions (650 local varieties obtained from

2200 BC on the Indian subcontinent (Weber 1992),

local famers) and herbarium specimens collected by field

while P. sonorum was domesticated from P. hirticaulesurveys were used for observations and experiments

around 600 BC in northwestern Mexico (Nabhan andon morphological and ecological characteristics,

de Wet 1984). On the other hand, common millet

crossability, amplified fragment length polymorphism

was the most important grain crop that supportedof total DNA, phenol and iodine color reactions ofcivilization around Eurasia over several thousand

seeds, fatty acid component in seeds, traditional foodyears starting in the Neolithic era. It is still cultivatedstyles, and archeolinguistic data. The botanical origin,and has various uses around the world. Although it

domestication process, and geographical dispersal

is one of the oldest domesticated plants in Eurasia,

of common millet are discussed and then integrated

the ancestral plant and place of origin have yet to bethrough the characteristics mentioned above. In

definitively determined. The place of origin of common

conclusion, common millet was domesticated frommillet has been discussed for many years (e.g., Bellwood

a wild population of P. miliaceum subsp. ruderale in2005, Church 1886, de Candolle 1886, Gerard 1597,

Central Asia, specifically from the Aral Sea to theHarlan 1995, Jones 2004, Kimata 2009, Sakamoto

Southwest Tien Shan Mountains. Since the Neolithic

1987, Vavilov 1926). Vavilov (1926) proposed that theera, the millet has been dispersed eastward to China,

original place was North China, and Chun et al. (2004)

west wa rd to Europe , southwa rd to the I nd ia nsuggested that common millet was domesticated in thesubcontinent, and northward to Siberia by nomadic

southern part of the middle reaches of the Yellow River

groups.

8000–7000 years BP.
Harlan (1975) suggested that the two homelands

Key words: common millet, dispersal, domestication,

were North China and eastern Europe. Nesbitt (2005)

Eurasia, local varieties, original placealso suggested that it might have been domesticated52

民族植物学ノオト 第９号

independently in each area. Although both the wild

morphological and ecological characteristics, followed

ancestor and the place of domestication of common

by studies of the secondary compounds in the grain

millet are unknown, it first appeared as a crop in both

(Kimata and Negishi 2002, Kimata et al. 2007).

Transcaucasia and China about 6000 BC. Zohary and

The present paper is concerned with the ancestor,

Hopf (2000) suggested that common millet may havedomestication, and geographical dispersal of common

originated somewhere between the Caspian Sea andmillet in Eurasia. The purpose is to examine these

Xinjiang. The earliest sites bearing remains of common

questions using all botanical characteristics, including

millet are in China and Europe from the seventh

biocultural diversity (traditional food styles andmillennium BC, on opposite sides of the Eurasianarcheolinguistic data) and genetic characteristics

continent (Jonse 2004). Furthermore, Sakamoto (1987)(crossability, F1 hybrids and AFLP markers).

indicated that the area was located within Central Asiaand the northwestern parts of the Indian subcontinent.

Materials and Methods

Common millet was found from the sixth millennium

Many endemic varieties and relatives of Panicum

BC at Tepe Gaz Tavilla in southeastern Iran (Meadow

miliaceum L. have been collected from all of Japan and1986). One explanation for its domestication could be

the Eurasian continent through field surveys since 1973.

an altered climate earlier in the Holocene. Meadow

Grain samples (650 accessions) were collected along the(1986) suggested that it might have been grown using

survey route and the voucher herbarium specimens were

floodwater runoff to supplement rainfall in this dry

deposited at Tokyo Gakugei University (Tokyo, Japan).

area as a spring or autumn crop. Similarly, farmers grow

Information on agricultural practices, grain processing,common millet in the Ganga area after winter floods.

food preparation, and vernacular names was gathered

Detailed local analyses of this kind should underpin

from local farmers.

future considerations of common millet’s origin in

Some of these accessions, 441 local varieties, wereCentral Asia to determine its possible routes of spread

selected and grown at the greenhouse of Tokyo Gakugei

through this critical but underexplored area (Hunt andUniversity, Japan to compare their morphological andJones 2006). However, the ancestor and original place

ecological characteristics starting on July 10, 1986.

of common millet have yet to be clearly determined.

These local varieties included 132 from Japan, 39

Bellwood (2005) summarized recent thinking on

from eastern Asia, 78 from the former USSR, 90 fromthe origin and spread of common millet based on recent

southern Asia, 26 from western Asia, 43 from Europe,archaeological contributions as follows. Common

two from Africa, and one from Canada (Kimata andmillet perhaps originated in Central Asia (Sakamoto

Negishi 2002).

1987, Zohary and Hopf 2000). Neolithic settlers may

Ten grains of each strain were sown in a seeding

have migrated from there to Afghanistan, the Russian

box with a row spacing of 8 cm and seed spacing of 2

steppes, or even western China. Common millet has

cm. Two weeks after sowing, germinated plants werebeen reported widely in the Neolithic cultures oftransplanted into the greenhouse, with 30-cm row

Europe and the Eurasian steppes, but it would appear

spacing and 15 cm between plants. Chemical fertilizer

that the oldest known dates of cultivation are from

(N:P:K = 8:8:5) was supplied at 100 g/m2. Five plants

North China from about 6500 BC onward. The firstof each strain were measured for traits, including the

known occurrence in southeastern Iran was in the

duration to flowering (days), number of leaves on thesixth millennium BC (Meadow 1986), c. 1550 BC inmain culm, number of productive tillers, hairiness ofnorthwestern Iran (Nesbitt and Summers 1988), andthe uppermost internode, panicle type, lemma color,

about 2600 BC in South Asia (Fuller et al. 2001).

pistil stigma color, and others. These morphological andA series of studies was started on common millet’s

ecological data were analyzed statistically using partial53民族植物学ノオト 第９号

Table 1. Materials used of Panicum miliaceum and the relative species$UHDFROOHFWHG-DSDQ. RUHD& KLQD
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correlation coefficients and hierarchical cluster analysis
in SPSS (ver. 21, IBM Corp).
Moreover, 70 local varieties, including six pollen

Fi g. 1. Food s f rom com mon m i l let i n
Uzbekistan and Inner Mongolia. a, milk
tea with roasted grains in Inner Mongolia;b, colored grains for a topping of bread in
Uzbekistan; c, milk porridge for healthfullunch at a nursery school in Uzbekistan.

testers, were selected and grown in the greenhouse from1990 to 1995. These accessions included 21 from eastern
Asia, 8 from Central Asia, 19 from southern Asia,
21 from Europe, and one weed, P. miliaceum subsp.
ruderale, from Romania. The crossability among the 70
Eurasian varieties and the morphological characteristics

associated with EcoRI (E+AAC, E+AAG, E+AGG,

of their F1 hybrids were examined in the six pollen

E+ACT, and E+ACA) were used in combination withtesters from France, Central Asia, India, China, Japan,

5 primers associated with MseI (M+CAG, M+CTG,

and a weed.

M+CTA, M+CAT, and M+CAA). Five microliters

Ten grains of each of 75 accessions were sown byof amplification products were loaded onto a 5.75%

the same method as above on Oct. 4, 2007 (Table 1).

denat u r i ng polya cr yla m ide gel (LON Z A) a nd

DNA extraction was performed on young leaf tissue

electrophoresed in 1× TBE for 1 h. Bands were detected

ground in liquid nitrogen and incubated in 1.5-ml

using the silver staining protocol described by Cho et al.

tubes containing 0.5 ml of buffer A for 10 min at 60(1996). The bands were detected on the gel at the finest°C by using CTAB (hexadecyl-trimethyl-ammonium

level of sensitivity by Lane Analyzer (ATTO), the raw

bromide) methods (Murray and Thompson 1980). The

data were adjusted, and then the visible and reproducible

AFLP procedure was performed according to Appliedbands were scored for accessions as present (1) or

Biosystems (2005), Bai et al. (1999), and Suyama (2001)

absent (0). The dendrogram of the AFLP markers waswith some modifications. Amplification reactions wereconstructed using the neighbor-joining and UPGMA

performed according to the same protocol. Five primers

methods (Nei and Kumar 2000) with the bootstrap

54民族植物学ノオト 第９号

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analysis (PAUP* ver. 4.0) and the hierarchical cluster

The distribution of glutinous varieties of common millet

analysis (group average method, SPSS ver. 21) on all

and Setaria italica were restricted to eastern Asia. On

data matrices of 75 local varieties.

the contrary, the geographical distribution of phenol
color reaction to seed coats in S. italica was very similar

Results

to that of Oryza sativa, but the distribution in common

Food preparation and secondary compounds in grain

millet was different from the trends in S. italica and O.

The Eurasian foods made from common millet aresativa (Sakamoto 1982, Kawase and Sakamoto 1982,
Kimata and Negishi 2002).

classified into four processing methods: grain, coarseground flour, fine flour, and drinks. Asian people cook

The four types of local varieties of common millet

boiled grain and porridge from the polished grains ofwere categorized by the composition of the minor

non-glutinous varieties (Table 2). Especially, East Asians

fatty acids arachidic, behenic, and eicosapentaenoic

cook steamed grain and mochi (a kind of cake) fromacid. If the ancestral prototype was the weedy AE typethe polished grains of glutinous varieties and ferment

containing arachidic and eicosapentaenoic acids, the AB

alcoholic drinks from polished grains of both non-type (arachidic and behenic acid) may have been bredglutinous and glutinous varieties. Inner Mongolians

both in Europe and Asia, while the ABE (all three fatty

drink daily milk tea with roasted grains (Fig. 1a). Uzbeks

acids) and O (no fatty acids) types may have originated

top non (a kind of bread) with colored grains (Fig. 1b)around Central Asia and then spread to both Europe

and cook milk porridge from non-glutinous varieties

and Asia (Kimata et al. 2007).

for lunch at a nursery school (Fig. 1c). Europeans cook
Vernacular names

milk porridge from coarse-ground flour, bread from fineflour, and ferment non-alcoholic drinks from polishedThe linguistic data are as follows (Table 3). Prefi xes

grains of only non-glutinous varieties. Based on thefor the word for “common millet” were mainly “ki-,”endosperm starch in seed grains, the varieties were“che-,” “va-,” or “ba-” in East and South Asia, but several

divided into two glutinous or non-glutinous categories.

variations were noted in China. The prefi xes were widely55

民族植物学ノオト 第９号

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via Nepal.

diverse in Central Asia and the mountainous area ofPakistan. It was mainly “d-” in western Asia and Egypt.

Taxonomy

There were also many European prefi xes, including “mi-”and “proso.” Because the vernacular names of common

Fig. 2 shows typical panicle types such as sparse (a1),millet were remarkably diverse around all Eurasia,

compact (a2), and dense (a3). It also shows a domestic

this indicates that the crop was domesticated and/type (b1) and an escaped weed (b2) in Pakistan; aor broadly dispersed starting in a very ancient period.

domestic type (b3) and a weed, ssp. ruderale (b4), in

However, common millet is called “cheena,” “chiena,”

Romania; a domestic type (b5) and a weed, subsp.

or “chin” in the Indian subcontinent. Based on the

agricolum (b6), in Uzbekistan; a crop-like weedy

Farming/Language Dispersal Hypothesis (Bellwood

biotype, subsp. miliaceum (c1); and an F1 hybrid between

and Renfrew 2002), these vernacular names might bea domestic type and the subsp. ruderale in Pakistan

derived from China and “Qin” (an ancient Chinese(c2), with both sparse and shattering panicles. It also

Empire), indicating that common millet was dispersedshows a weed, subsp. ruderale, in Inner Mongolia (d,from China to the Indian subcontinent through a routetaken in 2004) and European common millet (e1 and56

民族植物学ノオト 第９号

Fig. 2. Morphology of common millet, Panicum miliaceum. Types of panicle:
a1, sparse; a2, compact; a3, dense. Domestic type and mimic weed in Central
Asia: b1, b3 and b5, subsp. miliaceum; b2, an escaped weed; b4 and d, subsp.
ruderale; and b6, subsp. agricolum. c1, a crop-like weedy biotype in Pakistan
and c2, a F1 hybrid between subsp. miliaceum and subsp. ruderale. e1 and e2,
European common millet in 17th century (Gerarde 1597).

e2) illustrated in a book (Gerarde 1597). The panicles

Japan. Race contractum had a droopy, compact panicleof common millet can be divided into five types: sparse,(a2). Race compactum had a cylindrical, erect panicle.

compact, dense, and two intermediate types (relatively

Race ovatum had an oval, dense panicle (a3). Because

sparse or dense). Common millet is generally a densely

these morphological characteristics did not clearly

piliferous plant, but the hairiness of the uppermost

show a geographical cline, this classification was notinternode is very variable. This trait can be divided into

indicative of their taxonomical characteristics. The

four types: glabrous, sparsely, moderately and densely

taxonomy of common millet needs to identify intra-

piliferous.

specific differentiation through a matrix of various

Lyssov (1968, 1975) classified P. miliaceum L. into

characteristics. Scholz and Mikolāš (1991) classified P.

five groups based on the panicle types as follows. The

miliaceum into three subspecies: miliaceum, ruderale,panicle of race miliaceum was similar to that of a wild

and agricolum. Subsp. miliaceum consisted of thespecies. Race patentissimum had long, slender, and sparse

cultivar form (b1, b3, b5) and crop-like weedy biotype

panicles (a1), but it was very difficult to divide these two(c1) in Pakistan, and also in Austria, Slovakia, andraces, which were distributed from Eastern Europe to

Canada, respectively. Subsp. ruderale (b4, d) was an57

民族植物学ノオト 第９号

Table 4. Partial correlation coefficients of 14 characteristics
Days forflowering

No. oftillersPlantheightNo. ofLength of Width ofleaves onflag leaf flag leaf
main culm

FL/FW

Paniclelength

DF
1.000
0.005
0.835**
0.916** 0.501** 0.503**-0.032 -0.400**
TN
0.005
1.000
-0.203-0.173-0.216 -0.347** 0.297*-0.259PH0.835** -0.203
1.000
0.907** 0.746** 0.736**-0.095-0.024LN
0.916** -0.173
0.907**
1.000
0.594** 0.640**-0.145-0.310*FL
0.501** -0.216
0.746**
0.594**
1.000
0.787**
0.164
0.179
FW
0.503** -0.347**
0.736**
0.640** 0.787**
1.000-0.451**
0.170
FL/FW
-0.0320.297*-0.095-0.1450.164-0.451**1.000-0.052PL-0.400** -0.259-0.024-0.3100.179
0.170-0.0521.000
DI0.569** -0.375**
0.804**
0.713** 0.726** 0.814**-0.2540.169
PT0.363**-0.1570.543**
0.372** 0.382** 0.515**-0.2260.235
LC-0.055-0.118-0.030-0.0090.039-0.127
0.217
0.061
SC
0.131
-0.0450.015
0.172-0.012-0.1230.123-0.240HI
0.078-0.008
0.036
0.056
0.049
-0.104
0.202
0.052
SH
0.027
-0.048
0.057
0.066
0.221
0.186
0.002
-0.116
Controlled value is Iodine color reaction; * 5%, ** under 1% level significance.

Diameter ofuppermostinternode

Panicle
type

Lemma
color

Stigmacolor

0.569**-0.375**0.804**
0.713**
0.726**
0.814**-0.2540.169
1.000
0.548**-0.033-0.0790.082
0.081

0.363**-0.1570.543**
0.372**
0.382**
0.515**-0.2260.235
0.548**
1.000-0.043-0.335*-0.128-0.142-0.055-0.118-0.030-0.0090.039-0.1270.217
0.061-0.033-0.0431.000
0.358**
0.102
0.043

0.131
-0.0450.015
0.172-0.012-0.1230.123-0.240-0.079-0.335
0.358
1.000
0.124
-0.011

Hairiness ofuppermost Shatteringinternode
0.078-0.008
0.036
0.056
0.049
-0.104
0.202
0.052
0.082
-0.128
0.102
0.124
1.000
0.053

0.027
-0.048
0.057
0.066
0.221
0.186
0.002
-0.1160.081-0.1420.043-0.0110.053
1.000

escaped weed from subsp. miliaceum (b2) around theAsia), and Europe indicated very broad variation (1–6),

world, with the small grains shattering easily on its

up to an extreme of 9 in 6.9% of samples from India.

sparse panicle. Subsp. agricolum (b6) was a mutant race

Common millet is a densely piliferous plant. The

with characteristics intermediate between the domestic

hairiness of the uppermost internode was divided into

form and subsp. ruderale. This subspecies grew in maize

four types: glabrous, sparse, moderate, and dense. Most

fields because of its strong tolerance to herbicide. The

varieties were glabrous or sparse, while the others were

two types of European common millet in the sixteenthdense in Hokkaido (40.0%), western Asia (26.1%), andcentury might have been the races ovatum (e1) andEurope (20.0%). The panicle was divided into five types:

patentissimum (e2). A F1 hybrid (c2) between subsp.

sparse, dense, compact, and intermediate values (Kimata

miliaceum and subsp. ruderale was grown.

unpublished). Most of the local varieties from Japan(Hokkaido), China, India, western Asia, the former
USSR, and Europe were the sparse type, while theMorphological characteristics
The heading of common millet often occurred

remaining varieties from Japan, Korea, and Nepal were

irregularly. Because the panicle flowered inside the leaf

the dense type. Only a few varieties from western Asia,

sheath 4–5 days before heading, the duration (days)

the former USSR, and Europe were the compact type.

to flowering from sowing was observed instead of the

The lemma color on mature plants was classified into

heading date. Generally, the duration of local varieties

six colors: dark brown, brown, pale brown, ivory, orange,from high latitude areas was brief, but the number

and grayish-green. The varieties from the former USSRof days was remarkably variable. The varieties fromand Europe showed large variations in color. Most grains

China, Mongolia, the former USSR, Europe, andfrom Japan (Hokkaido) and China were dark brown,

Japan (Hokkaido) flowered very early, by 40 days after

but others from southern Japan were brown, pale brown,sowing, while those from India and southern Japanor ivory. In India, the grain color included grayish-green

flowered late, a third of them by 80 days. The varieties(45.6%) in addition to pale brown and ivory. The stigma

from China, Mongolia, the former USSR, Europe, andcolor of the pistil in the mature stage was one of three

Japan (Hokkaido) had fewer leaves (5–10) on the maincolors: white, faint purple, or purplish-red. About 70%

culm than those of southern and western Asia, Korea,of the varieties showed the former two colors. Especially,and southern Japan (11 to 16). All of the varieties fromin southern Japan and Nepal all the varieties had white

Japan, Korea, and Nepal had only a few productive

stigmas except one. However, in Japan (Hokkaido),

tillers (1 to 3), while the varieties from southern andIndia, and western Asia over 73% of varieties had

western Asia, the former USSR (including Central

purplish-red stigmas. In Europe 28% of the varieties had

58

民族植物学ノオト 第９号

effective photosynthesis during the growing season, anda bold culm that holds a heavy panicle.
T he h iera rch ica l cluster a na lysis of eig ht
morphological characteristics and earliness is illustrated
in Fig. 3 (by using the group average method of SPSS).
The 75 local varieties were divided into two majorclusters, I, with five sub-clusters, and II, with two subclusters. Sub-cluster Ia consisted of 11 varieties, fromCentral Asia (3 varieties, former USSR), Uzbekistan(1), China (3), Spain (1), Germany (2), and Canada(1); Ib came mostly from Western Europe, including
a few from Japan (Hokkaido), Mongolia, Uzbekistan,and Pakistan; Ic came mostly from Eastern Europe,
including a few from Uzbekistan and the Indian
subcontinent; Id consisted of three varieties fromAfghanistan, Greece, and Pakistan; and Ie consisted ofonly one variety from India. Sub-cluster IIa consistedof 20 varieties, mostly from East Asia but including a
few from Nepal (3) and Bulgaria (1). IIb consisted of 11varieties, mostly from the Indian subcontinent and alsoa few from China (2), Japan (2), and Romania (1). The
distribution of morphological characteristics generally
showed two geographical trends, from Central andSouth Asia toward Europe via Asia Minor, and fromFig 3. Cluster analysis of morphological characteristics.

China toward India via Nepal (to the south) and Japan
via Korea (to the far east).

purplish-red stigmas (Kimata unpublished).
T he pa r tia l correlation coef f icients of 14

Crossability among Eurasian varieties andcharacteristics are shown in Table 4. The coefficientsmorphological characteristics of F1 hybrids

greater than 0.6 under a 1% significance level were PH

The crossability among six testers was estimated(plant height) and LN (number of leaves on the mainby their fructification rates. The f lorets (range 5 toculm) to days for flowering (DF). The DF, LN, FL and50, average 17.0) on panicles (1 to 3, average 1.2) wereFW (length and width of flag leaf), and diameter ofartificially crossed with the tester pollen of each variety,uppermost internode (DI) to PH; DF, PH, FW andyielding an average fructification rate of 4.8%. Crossing

DI to LN; PH, LN, and FL and FW to each other; DI

tests were conducted between 351 combinations yielding

to FL & FW; and PH, LN, FL, and FW to DI. The

117 F1 hybrids fructified. The artificial cross pollination

others, namely, number of tillers (TN), panicle lengthof common millet was technically very difficult because(PL), panicle type (PT), lemma color (LC), stigma

the quite irregular flowering happened often before

color (SC), hairiness of uppermost internode (PI), andheading, and anther dehiscence was very sensitive toSH (shuttering) were not highly statistical significant.

daily weather conditions and it did not open entirely

Therefore, the domestic varieties with late maturity are

under wet conditions on rainy days. Because of this, thetall with many leaves, a large flag leaf that maintains

observed crossability was relatively low, ranging from 059

民族植物学ノオト 第９号

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to 63.9%. The F1 hybrid was obtained from 18 varieties.

Central Asian varieties were infertile when crossed

The crossability of ovum parents was lower than that ofwith Indian or Chinese ovum parents similar to thatpollen parents among the testers. One to 15 seed grains

when crossed with a weed (p32). These data suggested

were obtained from each variety, and the germinationthat common millet was dispersed from Central Asia

ratio was observed in only 105 strains of the F1 hybrid.

to China and Europe, respectively , and then dispersed

Most seeds germinated well, while the others did notindirectly to South Asia and East Asia. The weed (p32)

germinate or necrotized immediately after germination.

was not a crop-like weedy biotype because it was isolated

All of the F1 hybrid plants had good pollen fertility of

reproductively and made no fertile hybrids as an ovum

over 78%.

parent, notwithstanding the assured fructification

Crossability among varieties was summarized to eachamong varieties from all regions in the reverse as acountry and region as shown in Table 5. The Frenchpollen parent. However, it might still be possible that

tester had the largest number of sound F1 plants (41.2%)

subsp. ruderale was an ancestor, since it made fertile F1with European varieties. The Central Asian tester madehybrids between the other varieties.

F1 plants (58.8%) only with East Asian varieties. The

The panicle type of F1 hybrids with the sparse-

Indian tester made F1 plants with East Asian (33.3%)

panicled Indian tester was also sparse. The F1 hybrids

and South Asian (30.8%) varieties. The Chinese testerwith the dense-panicled Japanese tester (p60) had sparse(p51) made F1 plants with East Asian (45.0%) andpanicles when combined with sparse varieties and dense

European (45.0%) varieties. The Japanese tester (p60)panicles when combined with dense varieties. The F1made F1 plants with East Asian (45.0%), Central Asianhybrids between varieties with middle-type panicles(37.6%), and South Asian (38.9%) varieties. The weed

generally also had middle-type panicles.

tester (p32, subsp. ruderale from Romania) made F1

Common millet is generally a densely piliferous

plants with South Asian (35.7%) and Central Asianplant, but the hairiness of the uppermost internode was(28.6%) varieties. The pollens of subsp. ruderale could

highly variable. This trait was divided into four types:

artificially fertilize the ovum of domestic varieties,

glabrous, sparsely, moderately and densely piliferous.

but the counter practices could not at all. Notably, a

The F1 hybrids between the moderate varieties (e.g.,domestic variety with sparse and shattering panicles

p9, p11, and p56) had a moderate internode except for(PC57-2 from Hokkaido, Japan) made F1 hybrids withp8 (glabrous). The F1 hybrids with the glabrous testers

the testers from Central Asia, India, and Japan, but notfrom Central Asia, China, and Japan mostly had awith the others.

glabrous internode except for a few in p2, p53, and p9

The French tester made the largest number of fertile(moderately). The F1 hybrids between the moderate or

F1 hybrids with European varieties, the Central Asian

dense varieties and the dense Indian tester were varied

tester with East Asian varieties, the Indian tester withwidely between glabrous, sparsely and moderately,

East and South Asian varieties, and the Chinese tester

while an F1 hybrid between a glabrous variety fromwith East Asian and European varieties (Table. 5).

Japan (Hokkaido) and the moderate Indian tester had a60

民族植物学ノオト 第９号

Fig. 4. Interspecific dendrogram of three domestic species in genus Panicum by
AFLP markers.

of the former USSR (USSR-CA, 2) and Poland. Ic2

glabrous internode.

consisted of ten varieties from Canada, USSR-EU (2),USSR-CA, China, Nepal, Bangladesh, and India (3).

AFLP markers
The AFLP markers of 75 local varieties were

Ic3 consisted of eight varieties from India (3), Pakistan

analyzed by PAUP* ver. 4.0 and SPSS ver. 21, including(4), and Japan. Id consisted of seven varieties, including

neighboring joint and UPGMA methods with the

Pakistan (4, with two weed types) and Uzbekistanbootstrap test. The interspecific differentiation of(3, with one weed type). Sub-cluster IIa consisted of

Panicum miliaceum, P. sumatrense, P. sonorum, andonly two varieties from Afghanistan. IIb consisted of

their relatives is illustrated in Fig. 4 (neighboring joint

9 varieties from China (2), Nepal (2), Romania (2),tree, PAUP*). Clear interspecific differentiation among

France, Spain, and Belgium. IIc consisted of 11 varieties

there species of Panicum were noted, including thefrom China, Korea, Mongolia (2), Turkey, Bulgaria,

domestic and weed types of P. miliaceum from Pakistan

Romania, Czechoslovakia, and Germany (3). The

and Uzbekistan, and the other species, P. sumatrense

distribution of AFLP markers generally showed two

and P. sonorum in the bootstrap test. However, thegeographical trends, from Afghanistan and Mongolia

phylogenic differentiation of common millet was nottoward Europe and Nepal via China (to the west andas clear among varieties based on the bootstrap testeast), and from Uzbekistan and Pakistan toward India(200 replicates), as shown in Fig. 5 (UPGMA tree,

and Eastern Europe via USSR-CA/EU (to the south

PAUP*), although there was a geographical trend in theand west).
On one hand, based on the hierarchical cluster

dendrogram.
The 75 varieties were divided into two major

analysis (group average method, SPSS), only two clusters

clusters: I with six sub-clusters and II with three sub-were detected among 51 varieties. Cluster I consisted ofclusters. Sub-cluster Ia consisted of five varieties fromfive varieties, including three weed types from Pakistan

Germany, Romania (subsp. ruderale), China, and Japanand Uzbekistan, while Cluster II consisted of 46(2 varieties). Ib consisted of seven varieties from Turkey,

varieties from the other regions.

Greece, Romania (2), and Japan (3). Ic1 consisted of six
Discussion

varieties from Yugoslavia, the European portion of the

T he bota nica l orig i n, domestication, a ndformer USSR (USSR-EU, 2), the Central Asian portion

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民族植物学ノオト 第９号

Fig. 5. Intraspecific dendrogram of common millet by AFLP markers.

geog raphica l d ispersa l of common mi l let werewith a continuous process of dispersal toward China.

discussed and then integrated through the results

The domestic type then dispersed from Central Asia

mentioned above. The following working hypothesis

to South Asia, directly to Europe, and indirectly tomight well-explain the place of origin and dispersal ofsoutheastern Europe via West Asia. On the other hand,common millet with respect to recent archaeologicalthis grain crop might have dispersed from China tocontributions (e.g., Fuller et al. 2001, Hunt and Jones

Japan and Southeast Asia. The ancient farmers who had

2006, Jones 2004, Nesbitt 2005). This hypothesis

cultivated barley and wheat in the Near East area had

is supported by the crossability among varieties in

not necessarily accepted common millet. However, theEurasia and the geographical variation of several genetic

nomads who had moved around the Eurasian steppe

characteristics (Table 5), although this needs further

had gladly accepted the millet as the food source, thedetailed study, especially the phylogeny of common

same as the present day Mongolian herdsmen, because

millet and its close relatives. The early domesticationof its early maturation within the short summer season

process began in Central Asia and then progressed

and its value as fodder for the livestock. They dispersed

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民族植物学ノオト 第９号

Fig. 6. Dispersal routes of common millet through the Eurasia. I, original place and II,
secondary dispersal center; solid lines, ancient routes from Middle Asia; dotted lines, the 13th
century AD routes from China.

common millet from Central Asia to China anddiversity, particularly foods and vernacular names

Europe. Common millet might have dispersed faster(Tables 2 and 3) to reconstruct the domestication

to Europe in an east/west direction at similar latitudes

process and dispersal routes (Kimata 2015c).

than to southeastern Europe in a south/north direction

An ancestral form of common millet might have had

across different latitudes. It matured early in summer,

early maturation, remarkable grain shattering, sparse

but barley and wheat grew slowly in winter.

panicles, small grains, many tillers, pale brown lemmas,

The traditional varieties cultivated by the Ainu

white stigmas, glabrous uppermost internodes, and non-people in Hokkaido, Japan are similar to the varieties

glutinous starch. Usually, the domestic form of cereals

from North China and Mongolia in their panicle typehas fewer productive tillers than wild forms. Many

and the duration to flowering, while the other Japanesevarieties from Central Asia and the Indian subcontinent

varieties are similar to the varieties from Korea andshow many of the ancestral characteristics. There is only

Nepal in their panicle type and stigma color (Kimata

a little information on the mimic weed type associatedet al. 1986). The same large variability in lemma color

with the domestic type of common millet (Sakamotoand panicle type was shown in varieties from both the

1988, Scholz and Mikolāš 1991), but several weed types

former USSR and Europe. Indian and West Asianwith remarkable grain shattering have been foundvarieties had very large variation in many characteristics

in Pakistan, Uzbekistan, and Kazakhstan (Kimata(Fig. 3 and Table 4).

1994, 1997). These seeds were mixed with those of the

As compared with the other varieties throughdomestic type. Because the varieties around Central

Eurasia, the varieties around Afghanistan, Pakistan,

Asia show large variation and their related weedy

India, and Central Asia had a large diversity ofsubspecies still grow today, this area is appropriate to be

characteristics, including sparse panicles and many

the place where common millet had been domesticated.

tillers. The geographical distribution of characteristics

Moreover, the weed types were classified into two

was useful information including the biocultural

subspecies, ruderale and agricolum, and a crop-like

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weedy biotype escaped from the domestic type. It would

research institutes, also for the distribution of importantseem that ruderale was an ancestor, while agricolum

local varieties; to Dr. M. Nesbitt and Dr. T. Cope, Royal

became a weed by hybridization between these two

Botanic Gardens, Kew, for their valuable suggestion andsubspecies.

arrangement for examining literature and herbarium
specimens; and to Professor Emeritus S. Sakamoto,

Common millet was domesticated from a wild

Kyoto University, for his excellent advice.

variety of P. miliaceum subsp. ruderale in Central Asia
including the northern mountains of Afghanistan
and Pakistan, especially from the Aral Sea to the

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Acknowledgements
The author wishes to express his hearty thanks tonumerous farmers for their kindness given their valuable
seeds and their millet cultivation still today; to many64

民族植物学ノオト 第９号

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Appendix:
International Forum for Promotion and Developmentof Minor Cereals(1997)
資料：国際フォーラムの提案 (1997)67民族植物学ノオト 第９号

68民族植物学ノオト 第９号

あとがき

私の第２の人生は、環境教育学を確立するという義務を優先し、教育実践研究に多くの時間をかけた。雑草や雑穀の調査研究・実験は、趣味という位置づけだった。義務と趣味によって、私生活は経済的にも時間的にも、さらに精神的にも甚だしく犠牲にした。自分の家族の世話をしないで、他家の子どもたちのお世話に時間をかけた。
「アンパンマンの正義は、自己犠牲だ。
自己犠牲を伴わない正義はない」と、やなせたかしは言っている。自己の信条や信仰に従うとはそのことであろう。すべては自己の内にあり、足るを知ることなのだろう。第 3 の人生
では義務はほとんどなく、さらに自由になり、趣味と家族に暮らしている。人生に何の恨みも残さず、足るを知って彼岸に行くことだと思うので、今は調査研究などのアーカイヴズ作り、すなわち売らない文筆作業、３家族分の有機無農薬の食料作り、刺繍など、趣味の仕事に勤しんでいる。
第９号の特集は、主にインド亜大陸の雑穀についてである。本来、雑草の進化研究を志したが、老師阪本寧男の誘導により、雑穀研究にも憑りつかれてしまった。初めは文化人類学の野外調査程度だったが、次第に深間にはまり、キビの起源と伝播の研究を進めることになった。
インド亜大陸調査では、優秀な隊員たちとの研究分担のバランスで、食文化を主な対象とした。ところが、畏友小林央往が、西アフリカの野外調査で、マラリアに斃れた。インドで一緒に調査旅行を繰り返した親友の志を継ぎ、インド起源の雑穀の研究を追加することにした。
彼の卓見に導かれながらも、能力不足で、十分な成果を挙げられたとは思えない。
この特集の拙い論文は、定年退職を間近に控えながら、国際誌に投稿する努力によって生まれたものである。残念ながら受理されるまで、あるいは投稿するまでには及ばなかった。しかし、第２の人生において膨大な労力を費やし、拙いとはいえ、二度と余人がなせる仕事量ではないので、残しておくべき研究成果であると考えた。また、小林の遺志であるとも思った。トルストイが言っているように、人はいずれ死を遁れられず、仕事は忘れ去られて行く。
しかし、内村鑑三が言ったように、小さな個人の最大遺産としての「人生」ではある。
「読む
人知らず」であっても、第３の人生の足るを知るために書いておきたい。

木俣美樹男

69

民族植物学ノオト 第９号

民族植物学ノオト

第 9 号（2016）

ISSN 1880-3881

発行日：2016 年 2 月 29 日
発行所：特定非営利活動法人 自然文化誌研究会
発行責任者：植物と人々の博物館 木俣美樹男
所在地：〒 409-0211 山梨県北都留郡小菅村 4115
小菅村中央公民館内 自然文化誌研究会
編集協力 & レイアウト＝宮本幹江［時遊編集舎］

印刷＝有限会社サンプロセス

Ethnobotanical Notes No.9（2016）

ISSN 1880-3881

edited by Mikio Kimata
Plants and People Museum, The Institute of Natural andCultural History, c/o Community Center , 4115 Kosuge,
Kitatsuru-gun, Yamanashi Prefecture 409-0211