C H A P T E R
4
Origin, domestication, and spread
R.P. Joshia, A.K. Jaina, Nikhil Malhotrab, and Madhulata Kumaric
a
JNKVV College of Agriculture, Rewa, Madhya Pradesh, India, bICAR-National Bureau of Plant Genetic Resources
Regional Station, Shimla, Himachal Pradesh, India, cAPS University, Rewa, Madhya Pradesh, India
4.1 Introduction
On the basis of a combination of archaeological data and genetic studies of wild and cultivated populations, mil-
lets and pseudo-cereals can be placed in likely, though somewhat debated, diverse geographical regions of origin.
Hence, questions remain regarding when and where these were first originated, used and later, domesticated along
with routes of their dispersal. The origin, domestication and distribution of millets and pseudo-cereals is thoroughly
researched and reviewed by the scientists of past and present era. A taxation of a good number of archaeological
datasets available till now has helped in understanding their role in the ancient crop economy along with diversity
and spatial extent in relation to customary and cultural changes globally. Millets were initially domesticated in dif-
ferent parts of the world including East Asia, South Asia, West Africa, and East Africa. However, the domesticated
varieties have spread well beyond there are of origin. Among the pseudo-cereals, amaranth and chenopod were
important crops for the pre-Colombian culture in Latin America, while buckwheat originated from Central Asia.
Overall, cultivation of millets and pseudo-cereals has been introduced in many countries because of their importance
in human diet. With the time, these crops have gained more interest, thus reaching more people and consequently
more countries.
4.2 Millets
4.2.1 Sorghum
Sorghum (Sorghum bicolor) is among one of the most valuable dry land crops cultivated in poor to medium
soils as a source of feed and fodder. It is a short-day C4 plant and a staple food crop for millions of poor people in
the semi-arid tropics of African and Asian countries. Having an easy adaptability to hot and poor moisture agro-
ecologies, it is widely considered as a climate smart resilient crop. Sorghum formed an integral part of the diet of
most Neolithic and Iron Age grain-producing communities in the Sahelian belt (Harlan, 1992; Bourlag, 1996) and
play an important traditional role in world especially in Indian sub-continent and sub-Saharan Africa. In India, it
established as crop about 4000 years ago (Fuller, 2003; Boivin and Fuller, 2009). Sorghum originated from distinct
centre of domestication in sub-Saharan Africa with the first reported domesticated sorghum in eastern Sudan
(Fuller, 2003). Historical evidences suggest that it was cultivated around 10,000 years ago although crop domesti-
cation and improvement began recently. Evidences from Anthropological studies suggests that hunter-gatherers
started to consume sorghum as a part of their diet as early as 8000 BC. The domestication of sorghum has its origins
in Ethiopia and surrounding places, commencing around 4000–3000 BC. Many varieties of sorghum were created
through the continual practice of disruptive selection, whereby selection for quite one level of a specific character
within a population occurs (Smith and Frederiksen, 2000). This results from a balance of grower selection for cul-
tivated traits and survival for wild characteristics, generating both improved sorghum races, wild races and inter-
mediate races (Smith and Frederiksen, 2000). These improved sorghum races were spread gradually along with
Millets and Pseudo Cereals. https://doi.org/10.1016/B978-0-12-820089-6.00004-5 33 © 2021 Elsevier Inc. All rights reserved.
34 4. Origin, domestication, and spread
the movement of people and trade routes into many other regions of Africa (approx. 1500–1000 BC), the Middle
East (approx. 900–700 BC) and eventually into the Far East (approx. 400 AD). Sorghum was transported to America
during the late 1800s to early 1900s which ultimately generated the diversity of new sorghum types, varieties
and races. Initial domestication of sorghum would have focused primarily on converting wild types with small,
shattering and dehiscence of seed to improved types with larger, non-shattering seed. The meaningful disruptive
selection resulted in sorghum types with vastly different characteristics in plant height, inflorescence type, and
of course, end use (food, fodder, fibre, building materials, etc.). Along with time, sorghum is described under the
family Poaceae, tribe Andropogoneae, sub tribe Sorghinae and genus Sorghum (Clayton and Renvoize, 1986). As
archaeological data and historical records of early sorghum cultivation are almost completely lacking and not
documented, conclusions on the origins and domestication of this crop are based on comparative morphological
studies correlated with distribution patterns.
4.2.2 Pearl millet
Pearl millet (Pennisetum glaucum L.) originated in tropical western Africa some 4000 years ago. From there, it di-
vided into globossum race and moved to western side, and a new race typhoid also appeared that reached eastern
Africa and travelled to India and southern Africa about 2000–3000 years ago (Vilas et al., 2015). In between 3000 and
2500 BC, pearl millet spread rapidly to African countries through pastoralists, spurred by the increasing desiccation
of the Sahara desert at that time. Vavilov in his classic work on ‘Origins of Crops’ allotted pearl millet on the Ethiopian
centre of domestication, the ancient highlands of Ethiopia are modern agriculturally diverse region in which crops
from many parts of the world are continuously grown successfully (Vavilov, 1926). There are two evidences which
argue against an Ethiopian origin of pearl millet. First, the wild progenitor (Pennisetum americanum subsp. monodii) is
rare east of Sudan and has never been collected in the Ethiopian highlands. Wild progenitor is adapted to the sandy
semi-arid conditions of the Sahel and very likely would have been absent from the high rainfall and high altitude en-
vironment of the Ethiopian highlands. Secondly, the pearl millet of Ethiopia lacks sufficient morphological diversity.
Now days, pearl millet is a minor crop in Ethiopia and is probably the product of post domestication introduction. A
second theory as to the origin of pearl millet was proposed by Mudrock (1959). As per initial linguistic evidence, he
proposed that pearl millet was one of the several West African crops domesticated by Mande people near the head
waters of the Niger River between 4000 and 5000 BC. The general hypothesis postulated by Mudrock (1959) has been
the centre of considerable controversy since its publication (Brunken et al., 1977). During the period suggested by
Mudrock, the head waters of the Niger probably exhibited a climax, tropical rain forest type of vegetation.
4.2.3 Finger millet
Eleusine coracana (L.) Gaertn. subsp. coracana (Finger millet or Eleusine) is widely cultivated in Africa and India,
commonly known as ‘ragi’ found to be originated in the beginning of Iron Age about 5000 years ago in the highlands
of Eastern Africa (from western Uganda to Ethiopia) by domestication of wild weedy forms and termed as oldest
known domesticated tropical African cereal. Archaeological records of finger millet are less and not authenticated
although distribution, linguistic and historical evidences seem to suggest an African rather than Indian origin of
finger millet. From eastern Africa, it spread to elsewhere; to India about 3000 years ago, to southern Africa about
800 years ago, before spreading to South-East Asia. It is widespread in warm temperate regions from Africa to Japan
and Australia. Finger millet varieties grown in southern India and African lowlands are morphologically indistin-
guishable (De Wet and Huckabay, 1966). There are also findings that finger millet was domesticated about 5000 years
ago in eastern Africa (possibly Ethiopia) which later on introduced into India, about 3000 years ago (Hilu and De
Wet, 1976). The closest wild relative of finger millet is E. coracana subsp. africana which is native of Africa (De Wet
and Huckabay, 1966). As a cultivated crop, it is at present most important in eastern and southern Africa as well as
in the Indian sub-continent, and is occasionally cultivated elsewhere in the tropics. In South-East Asia, it is grown
on a small scale thereby not attaining commercial importance. Morphological study supplemented with cytogenet-
ical observations and distribution suggested that E. coracana subsp. africana is a wild finger millet. This subspecies
of Eleusine is widely spread along the highlands of East Africa. Consequently, it was concluded that finger millet
originated in the East African highlands and was subsequently introduced and domesticated in India. The species E.
coracana subsp. coracana includes all cultivated finger millets, on the basis of inflorescence shape and its correlation
with geographic distribution (Hilu and De Wet, 1976). Evidences of the patterns of variability in African and Asian
finger millets has by large indicated comparatively larger diversity in African germplasm rather than to Indian
collections, supporting to the view that Africa could be the primary centre of origin. The long history of cultivation
.
4.2 Millets 35
of finger millet in the Indian sub-continent accompanied by natural mutations and human selection has resulted
in the generation of large diversity in landraces and local cultivars in India. Detailed study of various characters in
Indian germplasm has resulted that for economically important characters such as finger length, finger width, finger
number, grain yield, ear weight, total biomass and leaf number, the Indian germplasm possesses large variability
indicating India as the secondary centre of diversity (Naik et al., 1993).
4.2.4 Barnyard millet
Two barnyard millet species namely Echinochloa frumentacea and E. esculenta are under cultivation in the different
parts of world. The original archaeological evidence of the crop is from China, where it was cultivated since 4100
BC. E. frumentacea showed parallel line of evolution both in India and Africa. E. esculenta is annual crop cultivated
mostly in the temperate regions of Japan, Korea, China, Russia, and Germany. It is grown in India, Japan, and China
as a substitute to rice when the paddy crop cannot be grown. The crop has gained attention as a fodder in the United
States and Japan (Vilas et al., 2015). The genus Echinochloa includes about 20 species which are distributed in tropical
parts of the world. Some of them are aggressive colonizers of distributed habitats, and three are cultivated as cereals.
The recognized four morphological races of Echinochloa do not have geographical, ecological, or ethnological unity.
4.2.5 Kodo millet
Kodo millet (Paspalum scrobiculatum L.), is an indigenous cultivated cereal of India and is in cultivation since
time immemorial. It is estimated to have been domesticated in southern Rajasthan and Maharashtra dating back
3000 years ago (Kajale, 1977). Kodo millet is widely distributed in damp habitat across the Old World tropics. It was
harvested as wild cereal in West Africa and India about 3000 years ago. Wild P. scrobiculatum grows in the western
African countries as perennial grass (Joshi, 2014). Kodo millet is either cultivated or picked in Africa and India as
one of the psychotropic cereals (Roland, 1976). Nowadays, it is grown in Uttar Pradesh in the north, and Kerala,
Karnataka, and Tamil Nadu in the south. This cereal is popular as ‘kodo’ in Hindi and ‘varagu’ in Tamil. The species
is widely distributed and spread in damp habitats across the tropics and subtropics of the world. Paspalum scrobicu-
latum var. scrobiculatum is grown in India as an important crop, while Paspalum scrobiculatum var. commersonii is the
wild variety is indigenous to Africa. Paspalum scrobiculatum occurs throughout the Old World tropics (Clayton and
Renvoize, 1986).
4.2.6 Proso millet
Proso millet (Panicum miliaceum) is a small crop with many common names in different regions including proso mil-
let, common millet, broomcorn millet, kashfi millet, hog millet, white millet, and red millet. As per the Archaeological
evidences, the crop was first domesticated before 10,000 BC in Northern China (Lu et al., 2009). The proso millet is
comprehensively cultivated in China, India, Nepal, Russia, Ukraine, Belarus, the Middle East, Turkey, Romania, and
the United States, where approximately it is grown on half a million acres each year. The crop has its importance
among growers because of its extremely short lifespan, with some varieties producing grain only 60 days after plant-
ing, and its low water demanding requirements, producing much more efficiently per unit of soil moisture than any
other grain species (Graybosch and Baltensperger, 2009). Its most popular common name ‘proso millet’ comes from
the pan-Slavic general and generic name for millet (Croatian: proso, Serbian: просо). Initially, weedy forms of proso
millet were found throughout central Asia, covering a widespread area from the Caspian Sea to Mongolia, which
ultimately represent the wild progenitor of proso millet from domesticated production (Zohary and Hopf, 2000). It is
widely grown in northern china, Mongolia, Korea, Russia, Afghanistan, Pakistan, India, and southern Europe. It was
also introduced to North America (Vilas et al., 2015).
4.2.7 Foxtail millet
Foxtail millet (Setarica italica) was considered to be domesticated in the highlands of central China. The main
production regions of the world include China, parts of India, Afghanistan, Central Asia, Manchuria, Korea, and
Georgia. It is also one of the specialty crops in Japan. In Asia, it is mainly grown for human consumption. It was
recently introduced in the United States for hay and silage. Because of its short growth cycle, it is a suitable crop for
nomads, and was probably brought to Europe during the Stone Age (Vilas et al., 2015). It is necessary to understand
.
36 4. Origin, domestication, and spread
that the wild ancestor of foxtail millet has been securely identified as Setaria viridis, which is inter-fertile with foxtail
millet; wild or weedy forms of foxtail millet also exist. The primary difference between the wild and cultivated forms
is their seed dispersal biology (Zohary and Hopf, 2000). Wild and weedy forms shatter the seeds while cultivars re-
tain them. The most primitive confirmation of the cultivation of this grain comes from the Peiligang culture of China,
which also cultivated proso millet. Out of the all suggestions lately, the Cishan culture of China has been recognized
as the most basic to domesticate foxtail millet around 6500–5500 BC (Stevens et al., 2016). The most primitive ev-
idence for foxtail millet farming outside of its native distribution is at Chengtoushan in the Middle Yangtze River
region, dating back to around 4000 BC (Castillo, 2010). S. viridis is the antecedent of S. italica as confirmed by genetic
comparison (Diao and Guanqing, 2017). During its journey in southern China, foxtail millet reached the Chengdu
Plain around 2700 BC and Gantuoyan, near the Vietnamese border in around 3000 BC (Castillo, 2010). Foxtail millet
also reached Taiwan (Nankuanli, Dapenkeng culture) in around 2800 BC and the Tibetan Plateau (Karuo) in around
3000 BC. Setarica italica likely reached Southeast Asia via multiple routes. In the beginning the evidences for foxtail
millet rises in south east Asia from different sites of central Thailand including Khao Wong Prachan Valley, along
with the site at Non Pa Wai suggesting the primitive date around 2300 BC (Weber, 2010). The most primitive evidence
of origin of foxtail millet in East Siberia found from the archaeological site at Krounovka in Primorsky Krai, dating
to around 3620–3370 BC (Diao and Guanqing, 2017). The earliest direct claim for foxtail millet in Korea come from
Dongsam-dong Shell Midden, a Jeulmun site in southern Korea around 3360 BC (Crawford and Lee, 2015).
4.2.8 Little millet
Little millet (Panicum sumatrense), was originated in Indian peninsula (Weber and Fuller, 2007). There are two
races of little millet (Panicum miliare), namely, nana and robusta (House et al., 2000). Plants in race nana vary from 60
to 170 cm in height. The inflorescence is 14–15 cm long, erect, open, and highly branched. These branches sometimes
droop at maturity. Plants in the race robusta are 120–190 cm tall. The inflorescence is 20–45 cm long, opening com-
pact, and highly branched. It is primarily a self-pollinated crop with ~ 3.5% cross pollination. Little millet is grown
throughout India up to altitudes of 2100 m, but is of little importance elsewhere (Hulse et al., 1980). Although the
origin of little millet is not clearly evidenced, much favours for the Indian origin because of its endemic nature to
India. Out of the two races, P. miliare or commonly known as little millet is grown or naturalized throughout India
and Sri Lanka, and also cultivated in adjoining countries of India. Till date the antiquity of P. sumatrense cultivation
is unknown. This species is nowhere mentioned in the known archaeological records of cereal cultivation in India.
Its wide distribution as a small millet across most agricultural zones of India may suggest an ancient domestication.
4.3 Pseudo-cereals
4.3.1 Buckwheat
Buckwheat (Fagopyrum sp.) is among the three available non-grass starchy grain crops, or pseudo-cereals, the
others being grain amaranth (Amaranthus sp.) and Andean quinoa (Chenopodium quinoa). It was cultivated in traces
nearly in every country that cultivates grain crops and usually consumed locally but now gaining importance due
to its nutraceutical properties. Primitively, it is thought to have originally been found in Southeast Asia, its first real
appearance occurred in the Balkan area of Europe around 4000 BC. Buckwheat has a rich history in the United States
as well unlike other ancient grains. It was much common in colonial times. Though, its popularity peaked in 1866
but due to unawareness it severely declined lately. Central Asia, somewhere between Lake Baikal and Manchuria is
considered to be the origin of buckwheat (Fagopyrum sagittatum syn. F. esculentum) but in research now leans towards
southwest China and the Himalayan region as the centres of origin (Ohnishi, 1998). The two cultivated species of
buckwheat, Fagopyrum esculentum (common buckwheat) and F. tataricum (tartary buckwheat) are Chinese domesti-
cates whose origins are usually thought to lie in upland south west China, outside the major centres of agricultural
origins associated with rice and millet. Synthesis of the macro- and microfossil evidence for buckwheat cultivation
in China found just 26 records across all time periods, of which the majority was pollen finds. There are few or no
identifying criteria distinguishing F. esculentum and F. tataricum for any sample type. The archaeo-botanical record
requires reconciliation with biogeographic and genetic inferences of a southwestern Chinese origin for buckwheat.
Scrutiny of the genetic data indicates limitations related to sampling, molecular markers and analytical approaches.
Common buckwheat may have been domesticated at the range margins of its wild progenitor before its cultiva-
tion expanded in the north, mediated by changing ranges of wild species during the Holocene and/or by cultural
.
References 37
e xchange or movement of early agriculturalists between southwest China, the Chengdu Plain, and the southern
Loess Plateau. As per the modern Italian botanist Valerio Giacomini, who has made a detailed study of buckwheat,
there are three possible ways buckwheat arrived in Europe: southern Russia, the first route goes through Poland,
Germany reaching Belgium and France where buckwheat cultivated; the second way runs across Turkey, to Greece,
Hungary, and southern Russia; and the last one through the maritime contacts of Venice. Overall, the origin of buck-
wheat is unknown even if its name in Latin languages recommends an Arab provenance. There is a myth that Joost
van Gistele brought it back with him from his journey to the Holy Land in 1485. Ultimately the Arab and the central
Asian origin has been contradicted by recent archaeological research findings. It has come to knowledge through pol-
len analysis that buckwheat grew in the Netherlands and north-west Germany long before the beginning of present
era. During promotion and conservation of neglected crops after knowing there neutraceutical values, much atten-
tion has been given on buckwheat to decide its origin and way of spread along with domestication (Campbell, 1997).
4.3.2 Amaranth
Amaranth (Amaranthus caudatus) is evidenced to be a native to the New World and has been initially found in
the Old World as part of an archaeological excavation in Narhan, India, dated to 1000–800 BC (Saraswat et al.,
1994). The genus is native to Mexico and Central America (Tucker, 1986). A. hybridus is a native riverbank pioneer
of eastern North America and parts of Mexico, Central America, and northern South America (Sauer, 1967). It is
much more common in the eastern than the western half of the United States. Its range has expanded to Africa,
south-central Asia and Australia, possibly because of its use as a green vegetable, commonly known as smooth
pigweed. The three grain-producing species (A. caudatus, A. cruentus, and A. hypochondriacus) have differences in
agronomic performance. A. caudatus is grown at high elevations in South America and Asia; and most accessions
of it are prone to diseases and late maturity in the temperate zone. The most primitive and logical dating regarding
amaranth as a domesticated grain crop is from archaeological digs at a cave in Tehaucan, Puebla, Mexico, where
seeds and plants of A. cruentus were found as 6000 years old, whereas Sauer (1993) noted that initial domestication
could have happened much earlier and in different places. Although the timing and location of domestication are
unknown, A. caudatus has been used in South America's Andean highlands (Peru, Bolivia, and Argentina) for cen-
turies. A. caudatus may have been domesticated from an early introduction of A. cruentus which later on crossed
with a wild Amaranthus in the region. As evidence, archaeological record also displays domesticated amaranth
seeds appearing over a wide range from Argentina into the southern United States. Grain amaranth was used by
various Indian groups in the southwestern United States, and light coloured A. hypochondriacus seeds dated to 1100
AD have been found in the Ozarks. Use of amaranth leaves, both from cultivated and wild amaranths, also seems
to have been widespread. India has been regarded as one of the two main centres of distribution of grain amaranth
besides tropical America. Dioecious forms are absent from the Old World in general, whereas they are found in the
New World (Thakur, 1964).
4.3.3 Chenopods
Chenopod (Chenopodium quinoa) is undoubtedly nutritious food plant cultivated both in the Andes and in the
Himalayas. It is a member of the Amaranthaceae family. Quinoa initially originated from the Andean regions and
remained an important food crop, which encompass Bolivia, Peru, Ecuador, Colombia, Argentina, and Chile. It has
been reported to be related with the Incas and Tiwankans as a primary food for thousands of years before the first
Spanish invasion. C. quinoa of the Andes and C. nuttalliae of Mexico morphologically appears to be very closely re-
lated species. Artificial hybridization has been successfully completed between these two species of chenopods. It
is suggested that both species may have been derived from the same original wild type, a weedy quinoa, C. quinoa
var. melanospermum from South America (Heiser and Nelson, 1974). Many other species of genus Chenopodium are
important wild food sources in parts of North America and some of them are domesticated (Yarnellr, 1963). In Asia
in general and in Himalayan region in particular, the chenopods are cultivated since last 160 years.
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