The Origin of Cultivated Plants in Southeast Asia 1 H U I - L I N LI 2 Cultivated plants represent man's most important heritage, and we cannot afford to loose sight of this fact even in an age of great and rapid technological advancement. Among the world's major problems, none is more urgent than an increase in food crops production. Intellectually, our knowledge of the origins of cultivated plants and their role in the early history of mankind is of interest. Without considering cultivated plants, we cannot hope to understand the nature of conditions under which civilizations arose. A study of the origins of cultivated plants is interdisciplinary, intimately concerned with history, geography, archaeology, anthropology, ethnology, and linguistics. It is, however, basically botanical. Modem plant taxonomy began with Linnaeus, whose classic work, Species Plantarum (1753), is an attempt, though a highly premature one, to present a world flora. He included both spontaneous and cultivated plants, assigning a geographical origin to most of the plants that he treated. There were few facts to guide him. Even his knowledge of geography was often inadequate. Several authors in the early 19th Century worked on this subject, but source material then was based generally on classical writings and statements of the herbalists. The beginning of scientific studies on origins of cultivated plants stems from the work of Alphonso de Candolle, specifically from his Gdographie batanique raisonn~e (1855). The latest botanical methods were employed 1 Adapted from a lecture in Chinese given at the Chinese University of Hong Kong in 1965 and published by the University in 1966, The

Origin of Cultivated Plants in Southeast Asia, 28 pp., Hong Kong, 1966 ( in Chinese with English summary). The author is indebted to Dr. E. H. Walker of the Smithsonian Institution for his kindness in reading the manuscript and his many helpful suggestions for improvement. Submitted for publication November 8, 1968. 2 Morris Arboretum, University of Pennsylvania, Philadelphia, Pennsylvania.

and correlated with evidence from archaeology, history, and linguistics. The cultivated plants were treated only as a chapter in a work on the distribution of plants as a whole. De Candolle's interest in the cultivated plants was motivated by his study of evolution in the Plant Kingdom. Cultivated plants were taken as a model to elucidate larger problems. This chapter was later revised and greatly expanded into his L'origines des ptantes cultivdes (1882), a standard classical treatise on this subject even today. Toward the middle of the 19th Century the general idea of evolution developed among naturalists. De Candolle's first work antedated Darwin's Origin of Species (1858) by three years. Darwin used the origins of cultivated plants as a basis for his thesis on the origin of species, and his interest was further manifested in his Variation in Plants and Animals under Domestication (9). In 1858, selection was the key to Darwin's thesis on the evolution of plants and animals. In 1868, he added to this his untenable theory of "pangenesis," to explain how acquired variation could be maintained. At about the same time, Mendel was using cultivated plants in his experimental studies of the nature of variation. His Versuche iiber Pflanzen.Hybriden (1865) lay unnoticed by the scientific world until 1900, when its rediscovery sparked the science of genetics. Since then, genetics have extensively used cultivated plants as materials for research. Academic interest in the cultivated plants amongst geneticists and cytologists and the practical interest of plant breeders in the first quarter of the 20th Century resulted in the accumulation of much information on the nature of cultivated plants. It was, however, not until Vavilov that genetics was consistently and systematically applied to the elucidation of problems on the origins of cultivated plants. Vavilov's

Studies on the Origin of Cultivated Plants (35) has exerted a great influence on later workers. He refined and brought up to date the botanical methods of de Candolle, while the newer approaches of genetics and cyto-

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genetics were introduced and correlated with the older methods. The noteworthy contributions made by Vavilov and his school stimulated interest in the cultivated plants and brought scholastic research to the problem of their origins. In 1882, de Candolle treated 247 species, some of which were only briefly mentionec]. He was able to trace 199 Old World and 45 New World species to wild ancestral forms. Three additional species were regarded as uncertain and seven as extinct. Since then, the species concept has been considerably modified. The complex nature of species alone renders it difficult to attribute the ancestry.of various cultivated species to simple and precise origins with the same confidence as in de Candolle's time. In the study of the origin of cultivated plants, the determination of wild aneestors is a very complex field still demanding intensive interdisciplinary studies. The geographical origins of many of the cultivated plants have resulted from the investigations of numerous students in recent years and, in general, are clear. Even though we are unable to point to exact locations from which plants have orginiated, we do know the origin of cultivation or domestieation of most plants within definitive regions. These "regions" may be very wide. A number of the oldest cultivated plants were widely dispersed in prehistoric times, the earliest location of cultivation cannot today be readily determined to a limited area. Moreover, the literature pertaining to the origin of cultivated plants is exceedingly extensive and scattered. Although some regional studies are available, comprehensive treatments are very few (40). This paper treats the origin of cultivation of plants in Southeast Asia. "Southeast Asia" is here used broadly to include the area from northern China south to the Indonesian islands. It is limited in the west by the Himalayas and the Tibetan Plateau; in the north by the desert regions. There have been several studies on the major geographical regions of the origin of cultivated plants. De Candolle pointed out the three basic agricultural regions: southwestern Asia, China, and tropical America. But he considered it impossible dearly to delimit natural regions for the cultivated

plants. Vavilov studied this problem most extensively and intensively. From 1922 to 1935, he published many papers on the basic centers of cultivated plants. In 1926, he established five "fundamental world centers of the origin of cultivated plants," namely, southwest Asia, Southeast Asia, the Mediterranean region, Abyssinia, and America (35). These are, in truth, all very extensive geographical areas, not "centers." Following continual revision, his final scheme (36, 37) indicates eight centers: (1) China, (2) India, (3) central Asia, (4) the Near East, (5) the Mediterranean, (6) Abyssinia, (7) Mexico and Central America, and (8) South America. In addition, there are three supplemental centers, the Indo-Malayan center for No. 2, the Chiloe center, and the Brazilian-Paraguayan center for No. 8. Vavilov's research was done thirty years ago. Since then our knowledge of various cultivated plants has grown. There is now, consequently, need for reassessment and reinvestigation of the geography of the origins of cultivated plants. Vavilov's scheme is often cited by later authors and occasionally revised or amended. It is not possible to discuss and criticize here the basic methodology of his scheme and its contents. He was most familiar with Europe and western Asia, and he actually explored and collected in South America, but he seems to have been relatively unfamiliar with eastern Asia. His "Chinese Center" embodies the whole of China from the northern cold-temperate regions all the way to the subtropical south. This is not only not a "center" but it is incompatible with the geographical distribution of plants in general to consider an extensive political area as a natural unit. Vavflov originally professed to use plant geography as the fundamental principle of his division of the origins of cultivated plants' but it does not seem to have been followed in this instance. Under each of the geographical centers, Vavilov listed a number of cultivated plants which presumably originated there. These lists are too general and occasionally are not accurate. Many plants listed under his "Chinese Center," for example, are unimportant or cannot even be called cultivated plants. Plants such as Aconitum wilsonii, Actino-

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CULTIVATED PLANTS IN SOUTHEAST ASIA

sterna paniculata, Elatostema umbellatum var. convolucratum, Panax ginseng, Peueedanum iaponicum, Smilax China, etc. are all wild species that have not been cultivated. Vavilov's scheme indicates how impossible it is accurately to distinguish the "world centers" of origin of cultivated plants (1). It is more feasible to differentiate the main regions of collective origin of the more important crops. Within each of these regions, a number of significant plants arose in cultivation. The first cultivation of individual plants might have occurred at a number of different localities, but the points all lie within the range of a certain geographical region. Earliest attempts at cultivation must always have occurred within the natural area of distribution of the wild progenitor. Burkill (4) has criticized Vavilov's reliance upon plant evidence alone and his neglect of the human planter factor. He shows also that Vavilov's Old World centers are actually only political divisions along a latitudinal belt through the southern portion of the Eurasian continent. Political divisions create discontinuity in civilization, thus providing barriers to the dispersal of cultivated plants. Burkill does not present any scheme for the geographical division of the origin of cultivated plants. In his discussion of the plants of eastern Asia, he indicates such divisions as India, China, and Indo-China (to encompass the area east of India and south of China), Malaysia, Polynesia, etc. He distinguishes, especially, the difference between the plant geography of China and India: in India, the rainfall is heaviest after the season of highest temperature, while in China, the seasons of heaviest rainfall and highest temperature coincide. He also emphasizes the difference in cultural and historical backgrounds between the two areas, and carefully points out the significance of the region between China and India as a main route of southern migration of peoples from the Asiatic mainland in all periods, and its importance to the dispersal of cultivated plants. More recently Darlington (7), following Vavilov's scheme with slight modification, names nine regions as follows: (1) southwest Asia, (2) the Mediterranean, (3) Abyssinia, (4) central Asia, (5) India-Burma, (6) Southeast Asia, (7) China, (8) Mexico,

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and (9) Peru. Beyond listing these regions and a number of plants for each, no discussion is presented. As to China, he still follows Vavilov in recognizing it as a single unit, making no attempt at north-south differentiation in accord with plant geography. To pursue this study, it is necessary to recognize the innate characteristics of our investigation. Cultivated plants involve two essential factors, man and plant. Thus to study the origin of these plants one must utilize both the ethnobotanical and phytogeographical viewpoints and methods. The chief factors of phytogeography or plant distribution are climate, physiography, and historical geology. The effect of latitude or climate on plant distribution is the most pronounced. In eastern Asia the monsoon winds greatly influence the distribution of plants. Physiographically, in the west, the Himalayan Range, the Tibetan Plateau, and the desert region of central Asia clearly set off eastern Asia from the rest of Eurasia. In eastern Asia proper, the most important physiographic factors influencing plant distribution are the major mountain ranges which extend east and west. They are not high enough, however, to prevent the northsouth migration of plants. Since eastern Asia was not covered by the Quarternary glaciation, many kinds of temperate plants still survive in this area, whereas in other parts of the earth where the overriding continental glacier existed these plants became extinct. Therefore eastern Asia has the richest temperate flora of the world. This is further influenced by its continuous contact with the extensive and exceedingly rich belt of tropical vegetation in the south. While the ranges of these plants may be restricted by latitudinal influences, there are no high mountains barring their migrations, although there is some restriction due to the effect of monsoon winds and ocean current on climate. Thus a part of the tropical flora could spread rather widely and mingle with the temperate floras to a considerable extent. The mixing of temperate and tropical elements and the northern extension of certain tropical plants are significant aspects of the flora of this region. Also significant is its richness in number of species and their great diversity (17). For these reasons, the primary physio-

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67

Fic. 1. The Four Belts of Origin of Cultivated Plants in South-East Asia (I. Northern China. II. Southern China. III. Southern Asia. IV. Southern Islands.) graphical divisions are latitudinal zones. Their boundaries, however, are not precise but only approximate. Although the majority of the plant species are confined to disinct regions, the ranges of many others extend into neighboring regions, thus transcending the physiographical boundaries of

the geographical regions (16). In the distribution of the cultivated plants this phenomenon is especially common due to man's overriding of the subtle latitudinal influences determining plant distribution. There are four main latitudinal zones in eastern Asia in respect to cultivated plants,

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CULTIVATED PLANTS IN SOUTHEAST ASIA

namely, (1) the Northern China Belt, (2) the Southern China Belt, (3) the Southern Asia Belt, and (4) the Southern Islands Belt. These are shown in Fig. 1. Ethnobotanically speaking, the way plants are used by peoples of different regions indicates both the level of their development and the nature of their culture--also the origins and history of the cultivated plants they use. Man utilizes plants first for food, next for clothing; secondary and later uses are for dyes, perfumes, and industrial purposes. Ceremonial uses of plants rank very high among less well developed societies. This may actually account for the introduction of some species into cultivation. The basic requirements for human nutrition are starches, fats and oils, proteins, and vitamins. Starch is obtainable only from plants. Fat, oils, and proteins can be obtained from animals, but they can also be obtained directly from plants. It is imperative that man have a sustained, balanced diet to insure his health. If a population can not obtain at least a minimum standard of nutrition for long periods of time, it will not achieve a high degree of culture through uninterrupted creativity and constructive enterprise. Nor will its production level be sufficiently high to achieve population growth, which seems to accompany cultural development in the New World. The chief sources of starch are cultivated cereal grains and tuber crops. Each has a different role in human culture. Cereals can be preserved longer than tubers, and their nutritional value per volume is higher. Thus they contribute more toward man's welfare. The chief civilizations of mankind were all built on cereal grains. Propagation of cereals is mainly by means of seeds, while tuber crops depend mainly on vegetative reproduction. This difference has resulted in different agricultural techniques. Plant proteins come mainly from leguminous plants. These plants, because of symbiotic nitrogen-fixing bacteria in nodules on their roots, can combine free nitrogen in the air with other materials to form nitrates, a manufacturing process most other plants cannot perform. Thus the legumes not only can be substituted for food of animal origin, but also can enrich the fertility of the soil, tlms promoting the growth of man's other

7

plant foods. As man became aware of these facts, he gradually developed the techniques of fertilizer application and crop rotation, and agriculture became highly organized. The increased use of fat and oils of plant origin parallels the development of special techniques for extracting them. Thus the cultivation of oil-yielding crops in a given region indicates a higher achievement in agriculture and a higher cultural attainment as a whole. When a given region has a number of desirable crop cultivated in each of these three categories, it clearly implies that agriculture of this particular region has become a well rounded system. This is the necessary physical basis for further development of the civilization of a higher order. As civilization develops further, cultivation and utilization of other types of plants such as fruits, vegetables, condiments, and beverage plants follow. The number of plants of each of these types, their different nature and different methods of utilization, etc., are closely related to the particular nature of the culture of any one of these regions. Thus it is possible to discover and examine the background factors in the development of the peculiar culture of each of the regions. In recent years, geographical studies on the origin of cultivated plants, aside from the problem of divisions or centers, have been concerned mainly with plant dispersion between the Old and New Worlds in pre-Columbian times, while relatively little attention has been given to the relation of cultivated plants to the development of cultures. (23, 24, 1, 29, 4, 2). We can now examine the four belts of southeastern Asia and their cultivated plants. The first region is the Northern China Belt. This includes the Yellow River Valley and the southern part of northeastern China (Manchuria). In the north it extends to the desert, in the south to the Tsin-ling Range, and eastward to Korea and the Yellow Sea. This is the famous loess region, and the place of birth and early development of Chinese civilization. The Tsin-ling Range is, in respect to geography and plant distribution, a conspicuous boundary between the north and the south. The mountain ranges east of it are not very high, leaving obscure the divisions between the plant regions. In general, the

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From ancient to present times, the chief southern limit of this belt is along the north legume crop of northern China has been the side of the Yangtze River. The loess is of loose texture and rich in soybean, Glgcine max. It was called in anmineral content and especially suitable to cient times, Shu, and it is now generally called the development of agriculture. But the Ta Tou (Great Bean). Not only does it climate of this belt is the most severe of the furnish the necessary proteins for nutrition, four belts of Southeast Asia. The precipita- but it is also the chief oil crop in China. It tion is the least and most unevenly dis- has long been exploited in multitudinous tributed, and it has fewer kinds of plants. ways, not only as a staple every day food but However, the cultural level of the people also as a diet supplement. Today, it is the here is developed to a higher degree, and its most wide spread and the most diversely system of agriculture is also the most com- utilized crop in all China. Thus it is very plete. This situation seems to agree with closely related to the nutritional physiology Toynbee's theory that civilization develops as and physical nature of the Chinese. This is a very interesting problem in their cultural a result of a challenge (34). One of the cereal crops that originated in history and is worthy of further investigathis belt is millet, Panicum miliaceum, called tion. by the ancient Chinese Chi or Shu and at the The cultivation and utilization of the soypresent time Shu or Mi Tzu or Huang Mi bean in ancient China also greatly affected (Yellow Grain). This has been cultivated in the development of agriculture. The ancient northern China since earliest Neolithic times peoples are known to have been at first igno(3). The oldest literature extant clearly rant of the use of fertilizers, but eventually shows that this was the most important discovered that the cultivation of soybeans cereal in earliest times. The significant posi- increased the productivity of the soil. Thus tion of millet in history and religion is shown the soybean is responsible for the developby the use of the name She Shu as a term or ment of crop rotation and the application of symbol for the country or the people. This fertilizers. Among all the legume crops, the also proves that millet was cultivated in this soybean has the most complete protein comregion in the earliest and most ancient times. plex, approaching most nearly that of aniThe foxtail millet, Setaria italica, called mal protein. Thus, in human nutrition it can Liang or Su, now colloquially known as completely replace animal food. Hsiao Mi (Small Grain), is also of very The relatively small development of aniearly appearance in northern China. Its dis- mal husbandary and fishery in northern tribution in Neolithic times was very wide, China could be the effect of the use of soyas it has been found with the lake dwellers bean. Soybeans were introduced into Japan of the Alps in central Europe and in northern at an early time but were not carried to exChina. Its actual place of original cultivation tra-Chinese areas in southern Asia because of cannot be exactly ascertained, but its ap- its photoperiodism. The soybean is a longpearance in northern China in a very remote day plant and cannot be easily grown in age is a proven fact. Thus, it is quite pos- areas of different latitudes. In general, a sible that its cultivation began in China. specific variety of soybean will ripen either In northern China, barley, Hordeum vul- too early or too late if grown more than a gate, and wheat, Tritic~tmaestivum, were in- hundred miles north or south of its original troduced from the West at very early times. habitat. The surmounting of this difficulty Rice, Oryza sativa, was also brought from is an achievement of plant breeding in recent the south. When the use of wheat and barley times (39, 13). became more common in northern China, The soybean is also the most important millet and foxtail millet, inferior in both food edible oil crop in China. The ancient people, value and productivity, were gradually re- because of their limited knowledge and techduced in importance. They are still grown nology, did not know how to extract oil from today because of their earlier harvesting and seeds. Thus they preferred more slippery drought endurance, in spite of the unfavor- food. In northern China, the earliest cultiable climatic conditions and the limited Jr- vated vegetables were not the present-day riga(ion in northern China. common Chinese cabbage, Brassica chinch-

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CULTIVATED PLANTS IN SOUTHEAST ASIA

s/s, celery cabbage, Brassica pekinensis, and radish, Raphanus sativus. These are all of later cultivation or introduction and developed gradually. The most important of the earliest cultivated vegetables, according to records in the early classics, is the K'uei, probably the mallow, Malva sylvestris. Now a weed of fields and dooryards, it was anciently the most widely planted and preferred vegetable. The leaves are rich in a mucilaginous substance. However, it gradually lost its importance, and by the time of the Ming dynasty was nearly forgotten. As the mallow is a perennial plant, relatively difficult to cultivate, it was gradually replaced by more easily cultivated crops. At the same time, its slippery nature also lost its value when oil crops become more widely used. In ancient times the tender leaves of the soybean, called Huo, were also used as a leafy vegetable. Besides the mallow and soybean leaves, other cultivated and commonly used vegetables were lost because of their gradual replacement by other crops. Some of these anciently used vegetables are Angel-

ica kiusiana, Lactuca denticulata, Nasturtium indicum, Polygonum hydropiper (knotweed), Viola verucunda (violet), Xanthium strumarium (cocklebur). They are now not only no longer cultivated but found only as weeds or wild plants of the fields or dooryards. Among these the knotweed and the violet, like the mallow, are also mucilaginous. The work Ch'i-min-yao-shu "Essential Arts for the People" by Chia Ssu-yeh, of the Later Wei dynasty in the 6th Century A. D., is the most ancient complete treatise on agriculture still extant. From its account of the vegetables, we know that in northern China one thousand and five hundred years ago the mallow was still the most important leafy vegetable. Those vegetables mentioned above that have now degenerated into weeds were then in cultivation. At the same time there were other kinds introduced from faraway regions. Those that came from the West included coriandaer, Coriandrum sativum, melon, Cucumis melo, alfafa, Medicago sativum, and radish, Raphanus sativus; from the south were white gourd, Benincasa cerifera, cucumber, Cucumis sativus, eggplant, Solanum melongena, and ginger, Zin-

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giber officinale. Another group of plants began their cultivation in northern China, including spring onion, Allium ~tulosum, leek, Allium ramosum, garlic, Allium sativum f. pekinense, Chinese cabbage, Brassica chinensis, and other species of Brassica. They are still cultivated there and some have also been spread to other places. Among these, the radish and the Chinese cabbage, after long and intensive cultivation, have developed many varieties and moved from secondary positions in former times to become the most important vegetables of today. Among the cultivated plants, the vegetables have changed the most historically. Those that have been carried to other places may have developed new centers of variations in cultivation. As the Chinese historical records are exceedingly rich in plant records, they are worthy of intensive investigation. Noteworthy is the paucity of tuber crops among the cultivated plants originated in China. The taro and the yam, commonly cultivated since ancient times, came from the south. The only plant of this type that came into cultivation is Chinese artichoke, Stachys sieboldii, but it is of relatively little importance. Its range and extent of cultivation through the dynasties remained limited. The great number of fruit trees is significant. Those first cultivated in China include peach, Prunus persica, Chinese plum, Prunus salicina, apricot, Prunus armeniaca, Japanese apricot, Prunus mume, Chinese cherry, Prunus pseudocerasus, sand pear, Pyrus pyrifolia, Chinese crabapple, Malus prunifolia, and Chinese hawthorn, Crataegus pinnati[/z/a, all in the Rose Family. Besides these are persimmon, Diospyros kaki, Chinese jujube, Zizyphus iuiuba, and other relatively less important fruit trees. This abundance of temperate cultivated fruits in a region with relatively few species of plants indicates the emphasis placed on horticulture by the Chinese since ancient times, and it also indicates the special fondness of the Chinese for fruit trees. This fondness for cultivating fruit trees has affected the origin of cultivation of plants in general, especially in southern China. The hemp, Cannabis sativa, was cultivated very early in China, and ever since has been a most important fiber crop. Even the seed was used as grain in early times. But the

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precise origin of hemp is not easy to determine. It probably originated in Central Asia, but in Neolithic times it was already widely distributed in the temperate northern hemisphere. A very distinct cultural practice in China is the raising of the silkworm and the cultivation of the mulberry tree, Morus alba. The fact that China has since ancient times been known throughout the world for its silk needs no further discussion here. The cultivation of the varnish tree, Rhus verniciflua, is another peculiarity of this region. The utilization of silk and lacquer varnish has no counterpart in any other areas of the world. The cultivation of these two special crops demonstrates the creativity in the utilization of plants by the Chinese. To summarize: this many-faceted cultivation of plants of North China constitutes a well rounded, complete system of agriculture, the physical basis for cultural development. The horticultural type of cultivation is a characteristic of Chinese agriculture, practiced obviously since ancient times. Most of the plants prefer high dry grounds. This region is relatively poor in the number of plant species, yet a very significant number of cultivated plants developed here. This fact indicates the mutual relationship between a high cultural level and the origin of cultivated plants. Also, even in very early time China had already introduced several kinds of cereal and vegetable crops from the outside. These have been naturalized and improved upon, and some of them have become leading crops, replacing native kinds. This demonstrates a special character of the Chinese people, namely their ability to absorb cultures from the outside. The second region is the Southern China Belt. This starts in the north along the south side of the Tsin-ling Range in the west and extends eastward along its extensions in Honan to northern Kiangsu on the northern side of the Yangtze River to cover the greater part of the Chinese territory. In respect to plant distribution, the Nan-ling Range is a natural boundary in the south. South of this and westward to the southern part of Yunnan the vegetation is tropical. In respect to the cultivated plants this boundary is not a distinct one. The cultivated plants of the extreme southern part of China

are in general similar to those of the rest of the vast Southern China Belt. Apparently this is due to the human factor involved. In this Southern China Belt, the vegetation is of a warm temperate to subtropical nature. In plant geography, this is the richest temperate flora of the world. The large number of woody genera is especially distinctive (19, 21). This is mainly due to the fact that the region was not devastated by the Quartenary continental glaciation. The flora of this region shows great similarity to that of the eastern part of North America. There are many genera of plants that exist only in these two distinct regions and nowhere else in the world, a phenomenon that has been much emphasized in plant geography (17). The chief cereal crop, rice, and the chief tuber crops, taro and yam, came from the south in very ancient times. Many of the cultivated plants that originated in Northern China are also cultivated in this region. The cultivated plants show clearly that Chinese culture moved from the north to the south. The cultivated flora as well as other cultural aspects are 'sinicized' in the Southern China Belt. The cultivation of many special kinds of vegetables originated here. Moisture-laden monsoon winds greatly influence the climate. Thus precipitation is heavy and water habitats are abundant, with aquatic vegetation common and luxuriant. Among the cultivated aquatic plants, some, such as lotus, Nelumbo nueffera, and water chestnut, Eleoeharis tuberosa, came from the south in very early times. Among those that originated in this region are eurayle, Euryale ferox, arrowhead, Sagittaria sinensis, and water caltrop, Trapa natans. The large number of aquatic tuber and other crops is a special feature (25). Besides these, there are other aquatic vegetables used for their leaves, such as water mustard, Brassica iaponiea, water dropwort, Oenanthe stolonifera, watershield, Brasenia sehreberi and water spinach, [pomoea aquatica, and those used for their tender shoots, such as Zizania latffolia (Zi-

zania caduciflora ) . Among tuber crops, Chinese yam, Dioscorea batatas, has been cultivated since ancient times. For oil crops, there is oil cabbage, Brassica chinensis var. oleifera. Others

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CULTIVATED PLANTS I N SOUTHEAST ASIA

include many kinds of legumes and vegetables whose cultivation originated in this region such as adsuki bean, Phaseolus angularis, Chinese kale, Brassica alboglabra, garland chrysanthemum, Chrysanthemum coronarium, and scallion, Allium bakeri. The bulb of the lily, LiIium tigrinum, the flower bud of the daylily, Hemerocallis fulva, and young shoots from many kinds of bamboos are relatively unusual plant foods. The most important contributions of the Southern China Belt to the cultivated flora are the fruit trees. Among these are the world-known citrus fruits such as sour orange, Citrus aurantium, sweet orange, Citrus sinensis, Mandarian orange, Citrus reticulata, kumquat, FortuneUa iaponica, and wampi, Clausena lansium (33). Others include loquat, Eriobotrya iaponica, and Chinese strawberry tree, Myrica rubra. In the more southern region are lychee, Litchi chinensis, longan, Euphoria longana and Chinese olive, Canarium pimela. These are all the results of the utilization of special endemic plants. Among fiber crops, ramie, Boehmeria nivea, Chinese jute, Abutilon avicennae, and kudzu vine, Pueraria lobata were first cultivated in this region. These crops, although not widely cultivated in other parts of the world, each have some characteristics that make them suitable for certain special purposes. Among the industrial crops are tea oil tree, Camellia oleifera, Chinese tallow tree, Sapium sebiferum, and tung oil trees, Aleurites montana and Aleurites fordii. The large number of woody crops is also a special feature of this region. It has been mentioned earlier that this region has more woody genera and species than has any other region in the temperate world. The most famous native South China crop is the tea plant, Camellia sinensis. Its earliest cultivation was in western China around Szechuan, and it probably spread eastward to northern China in the Warring States period, 5th to 3rd Century B. C. Thereafter, its cultivation gradually extended to the central and southeastern region, including Anhwei and Fukien provinces. The use of plants as stimulating beverages is common all over the world. But most of these numerous plants are only used in a limited way by the natives. Only three have won world-

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wide acclaim, namely, tea, coffee, and cocoa. The use of a plant as beverage can be regarded as an achievement in human culture. Most primitive men learn how to ferment starch and sugar to produce alcohol, and thus wine drinking is a very early and wide spread phenomenon. Alcohol has stimulating as well as anaesthetic effects, and can bring quick and violent physiological reactions. In the beginning and during the early development of culture, alcohol undoubtedly played a part in promoting its advancement. It had certain effect on religious, military, and political activities. When a culture reaches a certain level of maturity and advancement, however, it seems there is a need for a milder beverage for use in the regulation and enrichment of life. The ancient Chinese used millet in making wine. Furthermore, wine was used in great quantities. After the Ch'in and Han periods, the use of tea gradually replaced wine as the chief drink. In the West, there is a similar situation. The northern peoples of western Asia and Europe in early times used grape wine as their only drink and also later gradually adopted coffee, similarly of southern origin, as their leading drink. In summary, it can be seen that in Southern China there was no occurrence of indigenous staples such as cereals and important starchy tuber crops. Indigenous cultivated crops in this region included many kinds of fruit trees and vegetables. These plants supplemented nutrition by the major food staples and aided variations to the daily diet of the people. These plants and the several kinds of fiber crops, industrial crops, and the beverage, tea, are apparently the result of the utilization of the very rich flora of this region, after the southern expansion of the higher northern culture. This Southern China Belt has a considerable number of cultivated plants introduced from the north. Most of the legumes, vegetables, and fruit trees of northern origin are also cultivated here. Many plants were also introduced from still further south, including rice, tuber plants, vegetables, etc. At the same time the cultivated plants which originated in this region have also been introduced into the neighboring northern and the still more southern regions. But from the standpoint of culture, this region has the

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closest relationship with the Northern China Belt, and this cultural effect on the cultivated plants is also the most significant. Therefore as a region of origin of cultivated plants, this Southern China Belt ought to be considered a secondary region in reference to the more primary Northern China Belt. It is mentioned above that the cultivation and utilization of the many aquatic plants is a distinctive feature of this region. This distinctive feature, as well as the effect of special environmental conditions upon cultivated plants can be illustrated by an interesting case, that of the Manchurian waterrice, Zizania latifolia. The Manchurian water-rice is distributed south to Taiwan and north to Harbin and is especially cultivated in the marshy regions south of the Yangtze valley. Its range of cultivation nearly corresponds to that of the lowland rice plant. The Manchurian water-rice was originally a cereal crop in ancient China. Its earliest cultivation may have been started in northern China. Although valued in ancient times as a cereal grain, it did not become an important crop, and it seems that both the wild and cultivated state existed in the earlier times. Later on, its cultivation gradually migrated to the south, and its use as a cereal also gradually diminished. In the south, it is chiefly grown for its tender fleshy shoots eaten as vegetables, called Chiao-pai or Chiao-sun. The origin of this Chiao-pai is very peculiar. The expansion of the stem of Zizania is not a natural condition of the plant but a pathological one. It is the result of the invasion of a parasitic smut fungus, Ustilago esculenta, into the tender stem, stimulating the growth of its cellular tissues into a swollen condition. In the early stage of the life history of the fungus, the young stem of Zizania enlarges into a tumor. This is the Chiao-sun ( Zizania shoot), loose-textured and sweet-tasting. After the spores of the fungus are ripened, there are clusters of black spores. At that time, the structure of the Zizania shoot becomes tougher, and it is no longer fit for food. The plant itself, after the infection of the parasitic fungus, does not flower and fruit. This prevalent pathological condition may possibly be the result of the transplanting of a northern plant to a far more moist environment than was its original habitat.

Among all the cultivated plants, the use of a pathological condition like this is rather unique. The third region is the Southern Asia Belt, the southernmost region of continental Asia. It extends from Burma, Thailand, to the Indo-Chinese Peninsula, with the exception of the southern part of the Malay Peninsula which penetrates into the southern seas and should be included in the fourth region. The extreme southern border of China, including Hainan Island, is a transitional zone lying between this belt and the Southern China Belt. This belt is already within the boundaries of the tropics. The temperature is forever mild, and the rainfall is abundant. The effect of moist monsoon winds is even more pronounced than in the former belt. The growth of plants is very luxuriant, and the species are exceedingly numerous. The main staple crop that originated in this Southern Asia Belt is also the most important cereal of all Southeast Asia, rice, Oryza sativa. The utilization of rice forms the backbone of the culture of this entire region. Thus, some call this the rice civilization. Although this view is not substantiated by archaeological evidence, the extreme importance of rice cultivation on the economy of this region is obvious. The beginning of cultivated rice, whether monophyletic or polyphyletic, is as yet an unsettled question, but the monophyletie theory seems to have received more support (6, 14, 30). There is little doubt, from a botanical point of view, that its cultivation originated in this region. Besides rice, other cereals originated here such as Job's-tear, Coix lachryma-jobi. Job'stear is not of any importance at present, and its cultivation is very limited in extent, but in anicent times it was once an important cereal crop. The grains of Job's-tear are large in size but small in number, and it is difficult to remove the husk. It is apparently a more primitive type of cultivated cereal crop. Its cultivation by mankind may even antedate that of rice. Another primitive type of cereal is a kind of millet, Echinochloa

[rumentacea. The large number and great importance of tuber crops arc distinctive features of this region. Among those whose cultivation originated here or those which were very early

LI:

C U L T I V A T E D P L A N T S I N SOUTHEAST ASIA

introductions from India are taro, Colocasia antiquorum, ape, Alocasia macrorrhiza, yam, Dioscorea esculenta, greater yam, Dioscorea alata, water chestnut, Eleocharis tuberosa, etc. These are all important tuber crops. Their importance to human nutrition and economics, however, was far greater in the past than in more modern times. With the introduction to the Orient of the sweet potato, Ipomoea batatas, after the discovery of America, these endemic tuber crops became gradually less important and are still continuously diminishing (4, 5). Some suggest that in the beginning of agriculture vegetative propagation was used rather than seed propagation (12). As a result of the greater number and importance of tuber crops in southern Asia, Saner (29) regards it as the "hearth" of the vegetatively propagated crops of Old World agriculture, and at the same time the very location of the origin of agriculture of the whole world. No important legumes or oil crops originated in this region. This may possibly be due to the abundance of fish and other animals that make the cultivation of such plants less urgent. The origin of the use for food of chickens, man's most important domesticated fowl, and of pigs, his most important domesticated food mammal, began in this region. Thus from the standpoint of nutrition, proteins and oils from vegetative sources do not assume here such importance as in North China. Man's most important fiber crop in the Old World, cotton, Gossypium arboreum and Gossypium herbaceum, originated in India, the neighboring region in the west. Thus, its cultivation and utilization were introduced into this region at a very early date. Similarly iute, Corchorus capsularis, originated in India and has also been cultivated in this region since ancient times. There are many vegetables cultivated in southern Asia. These crops have been widely disseminated and cultivated in many, lands, and thus it is not easy to ascertain exact location of their origin of cultivation. Those with ranges generally running from the latitude of India to Indo-China in southern Asia are balsam pear, Mormodica charantia, white gourd, Benincasa cerifera, serpent gourd, Trichosanthes anguina, luffa, Luffa

13

acutangula, amaranth, Amaranthus mangostanus, etc. Fruit trees which originated in this region include pummelo, Citrus grandis, and lemon, Citrus limon. The other numerous cultivated trees cultivated here are mostly very early introductions from the Southern Island Belt in the south and India in the west. The richness of the tropical flora in contrast to the paucity of cultivated fruit trees is worthy of note. The use of the widespread masticatory in southern Asia, the betel-nut, Areca catechu, is a peculiar local phenomenon. The utilization of this tropical plant is parallel with the use of tea in temperate China. The cultivation of two distinct plants, betel-nut, Areca catechu and betel-leaf, Piper betle, utilized together for the same purpose, both originated in southern Asia. Among the spice plants is cassia, Cinnamomum cassia. The condiment, pepper, Piper nigrum, is probably an introduction from India. The Southern Asia Belt is contiguous with India in the west. From ancient times communication between the two regions has continued uninterrupted, both by land and by the sea. Therefore many cultivated plants that originated in India were introduced in very early times, and at the same time those of this region have spread westward to India. Because of the close connection between these two regions, it is not easy to determine on which side lies the origin of cultivation of some of the plants. The same situation also exists with the island belt in the south. The endemic floras of these three regions are all very rich, but each has its own distinct genera and species. In the distribution of the natural flora, the geographical divisions are rather clear-cut. Thus the basis of the cultivated floras that originated in these three regions is apparently also distinct. But, because of the human factors and the early widespread dispersion of individual crop plants, the geographical divisions of the cultivated floras have become relatively vague and indistinct. Aside from the major staple food crops, this extensive region of continental southern Asia is a relatively poor one. The important legumes and oil crops and many other secondary food crops like fruit trees and vegetables all came from outside. This is probably

14

ECONOMIC

because of its geographical position between the three major regions, China, India, and the Southern Island Belt, which enables it to utilize the cultivated plants that they originated. At the same time this can also explain why the culture of the Southern Asia Belt, while started quite early, has not continuously developed to the same level as those of the neighboring regions India and China. Burkill points out that this region has always been the thoroughfare for southern migration of men from Central Asia. This continuous movement undoubtedly has had great effect on the origin and dispersion of cultivated plants. The fourth region is the Southern Island Belt. This is the archipelago region beyond the southern border of the Asiatic mainland, but it includes also the Malay Peninsula. The main portion comprises the Indonesian Islands, while the Philippines on the east and Sumatra on the west are along the periphery. The Southern Island Belt lies astride the Equator and is in the very midst of the tropics. Because of the oceanic climates of the islands, especially their abundant precipitation, this Belt contains the richest tropical flora in the eastern hemisphere. So far as cultivated plants are concerned, a fairly large number of species originated here and are quite different from those of the several belts mentioned above. The most distinctive feature in the cultivated flora of the Southern Island Belt is the large number of fruit trees. Rice, the most abundantly planted cereal, is a very early introduction from southern continental Asia. The cultivation of tuber crops is also very abundant. Different kinds of taro and yam have been widely planted since ancient times in tropical Asia all the way from India, southern mainland Asia, to the Southern Island Belt. Some of these may have originated in cultivation here. Besides these starchy tuber crops, there are two special vegetatively propagated crops that originated in this region, the banana and sugar cane. The multiplication of these plants is not achieved by using underground enlarged tubers, but by using plant divisions. This indicates that the technique of vegetative propagation has advanced further in this Belt, and vegetative propagation has become

BOTANY

the basic characteristic of the agriculture here. The banana, Musa X parad~iaca and the variant, plantain, are, among the cultivated plants, very unique in many respects. The sweet fruits of banana are edible raw as are other fruits. But the plantain or cooked bananas are starchy and must be cooked like tubers. They comprise a staple food of this region. The edible part used is the fruit, but this fruit is sterile, and the plant cannot be propagated by seeds like other fruit trees. In the cultivated bananas, there are many clones, comprising many polyploids. The origin and history of the cultivation of the bananas is an exceedingly complex one. Some understanding is now being reached after thirty years of research at Trinidad (32, 31). The banana developed from two wild species, after many evolutionary changes, into the peculiar crop plant of today. At first it was a normal diploid, which developed parthenocarpy and sterility. Hybridization between two kinds of diploids and artificial selection followed. This was again followed by the development of triploids and further hybridization, to be followed again by the development of tetraploids and hybridization. Finally, there was somatic mutation, common to vegetatively propagated crops in general. This hybridization possibly occurred in various localities. Thus, geographically, the range of initial cultivation of banana may be very extensive. There are probably a number of widespread centers of differentiation scattered from India to the Philippines. But the earliest cultivation was probably in Malaya or the nearby islands (26, 31). The other important crop is the sugarcane, Sacoharum officinarum. Sugarcane is the most important sugar crop of mankind. The use of above-ground stems as a source of sugar occurs otherwise only in Sorghum, but the latter is of much less importance. Only in modern times has another plant, the beet, Beta vulgaris, of temperate region, developed as an important sugar crop. Sugarcane, used since ancient times as a source of sugar even today, has very strong relationships with the culture and history of mankind. After long periods of cultivation, sugarcane, like banana, also developed numerous clones. The sugarcane has a very high degree

L'f:

C U L T I V A T E D P L A N T S I N SOUTHEAST ASIA

of polyploidy, from tetraploid to 12-ploidy. The complexity of its genetic nature is equal to that of the banana. The complex nature and history of its development from the wild plants to the cultivated crop we have today has also been partially revealed through investigation in recent times (38, 28). Based on the evidence we have today, its origin ought to lie in the general area of the Indonesian Islands. The most important and distinctive feature of the indigenous cultivated flora in the Southern Island Belt is the large number of tropical fruit trees. Besides the banana mentioned above, there are a number of fruits that can be used as staple foods, such as breadfruit, Artocarpus altilis, jackfruit, Artocarpus integra, carambola, Averrhoa carambola, camias, Averrhoa bilimbi, etc. These fruit trees have been cultivated all over tropical Asia since very early times, so it is not easy to ascertain their original home. Their range extends from India through southern Asia to this Southern Island Belt. Coconut, Cocos nucifera, is widely cultivated in the tropics, and was cultivated in both the eastern and western hemispheres in preColumbian times. Its center of origin cannot be positively determined at the present, but there are suggestions that its cultivation began in the Southern Island Belt. Among the plants that can be more definitely placed for the origin of their cultivation are lime, Citrus aurantifolia, mangosteen, Garcinia mangostana, rambutan, Nephelium lappaceum, lansone, Lansium domesr and the renowned durian, Durio zibethinus. Besides these are many other fruit trees of secondary importance that have their origin of cultivation in this region, such as kotamba, Terminalia catappa, iambu, Eugenia javanica, etc. There is another distinctive feature of the cultivated flora, that is the lack of leafy vegetables. In this area with its tropical vegetation luxuriant throughout the whole year, there are numerous wild plants that have young leaves or fruits that are utilized as vegetables. Osche & Brink (27) recorded 298 species in the Malaysian Islands that can be used as such. Under this condition of easy access to wild plants, the incentive toward cultivation is absent. Tropical southern Asia is the world-

15

famous home of many spices and condiments. These were one of the main motives of the maritime explorations and trade of 15th-Century Europeans. The discovery of the New World, the colonization of the Old World, the many subsequent wars of all magnitudes, the political competition and struggles, all had this same beginning. Even today, the instability of the situation in Southeast Asia affects the whole world. All these are the aftereffects of the relentless search for these plant species. The extensive utilization of these plants in this area is a necessity of the tropical climate. That the origin of these plants occurred in this region and not in Africa or Australia demonstrates the relation between the cultivation and utilization of plants and a higher cultural level. Among crop plants of this kind are those that have been cultivated since ancient times besides those which were originally used in the wild state but have come into cultivation in more modern times. A number of plants, such as ginger, Zingiber officinale, turmeric, Curcuma domestiea, and pepper, Piper nigrum, have been so long cultivated all over the southern part of Asia that it is difficult to ascertain their original homes. Their original cultivation most probably lay within the area from southern India to this region. Among the plants whose original homes are more or less definitely known are nutmeg, Myristica fragrans and clove, Eugenia caryophyUus, these having originated in Molucca. For this reason, Molucca is known as the Spice Island. The most distinctive features of the cultivated plants that originated in this Southern Island Belt are the presence of tuber plants and the absence of cereals and legumes among the staple crops. Among the secondary food crops, fruit trees are numerous, but no leafy vegetables originated here. Besides these, a number of spice and condiment crops are cultivated. The cultivation and utilization of the banana and sugarcane is a peculiar phenomenon. All these distinctive features are related to the tropical environment. Although many vegetatively propagated crops originated in this region, the kinds of cultivated plants are limited. In other words, the agricultural foundation here is not well rounded, and thus it is not suffici-

16

E C O N O M I C BOTANY

TABLE I DISTRIBUTION OF I~VfPORTANT CULTIVATED PLANTS WITI-I ORIGIN IN SOUTHEAST ASIA.

Note: Names in parentheses are ancient crops no longer in cultivation. L Northern China Cereals

II. Southern China

Panicum miliaceum Setaria italica

Root & Stachys sieboldii Tuber Crops

HL Southern Asia

IV. Southern Islands

Oryza sativa Coix lachryma-jobi Echinochloa frumentacea Dioscorea batatas Sagittaria sinensis

*Colocasia *Colocasia antiquorum antiquorum *Alocasia *Alocasia macrorrhiza macrorrhiza Dioscorea alata Dioscorea esculenta Eleocharis tuberosa

Legumes

Glycine max

Phaseolus angularis

Edible Oil Crops

Glycine max

Brassica chinensis var. oleifera

Vegetables

Bulb Bulb Leaf Allium sativum f. Lilium tigrinum *Amaranthus pekinense mangostanus Leaf Stem (Malva vertieilZizania latifolia Fruit lata) *Momordica charantia ( Angelica Leaf kiusiana ) Brassica *Benincasa cerifera ( Lactuca alboglabra *Trichosanthes denticulata) Brassiea japonica anguina ( Nasturtium Oenanthe *Luffa acutangula indieum ) stolonifera ( Polygonum Brasenia sehreberi hydropiper) Ipomoea aquatica ( Viola Chrysanthemum verueunda ) coronarium ( Xanthium Allium bakeri strumarium ) Allium fistulosum Flower Allium ramosum Hemerocallis Brassica chinensis fulva Brassica pekinensis

Fruit Trees

Prunus persica Prunus salicina Prunus armeniaea Prunus mume Prunus pseudocerasus Pyrus pyrifolia Malus prunifolia Crataegus pinnatifida Diospyros kaki Zizyphus jujuba

Citrus aurantium Citrus grandis Citrus sinensis Citrus limon Citrus reticulata Fortunella japonica Clausana lansium Eriobotrya iaponica Myriea rubra Litchi chinensis Euphoria ]ongana Canarium pimela

Artocarpus altilis Artocarpus integra Averrhoa carambola Averrhoa bilimbi *Cocos nucifera Citrus aurantifolia Gareinia mangostana Nephelium lappaceum

LI" CULTIVATEDPLANTS IN SOUTHEASTASIA

17

TABLEI (continued) I.

IL

III.

IV.

Northern China

Southern China

Southern Asia

Southern Islands Lansium domesticum Durio zibethinus Terminalia catappa Eugenia javanica

Other Special Food Crops

Musa • paradisiaca Saccharum officinarum

Beverage and Masticatories

Camellia sinensis

Spices and Condiments

Areca catechu Piper betle Cinnamomum cassia *Piper nigrum

Fiber Crops

t Cannabis sativa

Boehmeria nivea tGossypium Abutilon avicennae arboreum Pueraria lobata ~Grossypium herbaceum ~'Corchorus capsularis

Other Industrial Crops

Morus alba Rhus verniciflua

Camellia oleifera Sapium sebiferum Aleurites montana Aleurites fordii

*Zingiber officina]e *Curcuma domestica *Piper nigrum Myristica fragrans Eugenia caryophyllus

* Plants with uncertain geographical origin. t Crops introduced in ancient times from neighboring regions. ent to sustain the development of a culture at a high level. In summary, the geographical divisions of the origin of cultivated plants in Southeast Asia are here outlined in four latitudinal belts. The cultivated floras of the northern and southern belts are entirely distinct from each other (see Table I). In the north the emphasis is on cereals and propagation by seeds. Legumes and oil crops are more important in the north than in the south. A fairly large number of vegetables are cultivated in the north, and these vegetables have changed considerably through the course of history. Farther south, the cultivation of vegetables gradually decreases, and at the

extreme southern region few vegetables were introduced into cultivation. In the south there are a certain number of peculiar food crops such as banana and sugarcane that have no counterparts in the north. Fruit trees are extensively cultivated in both the north and the south, but the species are widely different. In the north most of the fruit trees belong to the Rose Family. In the south are tropical species in many families, no one being dominant. In the Northern China Belt, many kinds of cultivated plants developed. Among the cereals, legumes, oil crops, supplementary food plants such as vegetables and fruits, fiber crops, and other industrial crops, are

18

ECONOMIC BOTANY

plants whose cultivation originated in this region, forming together a well rounded agricultural complex. This constituted the basis of development of a high-level cultural system. Its influence extended to the Southern China Belt, making it a secondary region of the origin of cultivated plants. In the two belts in the south, while rice had its origin here, the chief staple crops are the many kinds of tuber crops and other vegetatively propagated crops. Relatively few legumes, vegetables, and fiber crops originated here, but there were many early plant introductions from outside regions. Because of factors of climate, geography and history, most of these introductions were from India. These two belts of the southern tropical region, although the site of many indigenous cultivated plants, do not produce crops of all kinds. Their systems of agriculture, while highly developed, are not well rounded. This is probably in part due to their close proximity to the two highly civilized regions, India and China. India, like China, has a complete complex of indigenous cultivated plants, hence its agricultural system has been a well rounded one from ancient times, resulting in a very high cultural level at a very early date. The domestication of animals and plants is a most significant step in the evolution of mankind. Thus man passed from barbarism into civilization. In the process of selecting and improving different food plants for cultivation, undoubtedly man's ways of living had to be modified to a certain extent to adjust to the food source available to him. In other words, man needed to adapt himself to his cultivated plants. Thus when man domesticated animals and plants, he himself underwent a process of domestication (1, 22). By artificial selection cultivated plants were adapted to the new environment of domestication. Similarly, the ability of mankind, after domestication, to adapt gradually to its ever changing artificial environment will determine his fate. From the above, it has been shown that a close mutual relationship exists between culture and cultivated plants. To further supplement this evidence, we can point to the situation in lands bordering Southeast Asia. In these border areas almost no cultivated plants originated. The Japanese Archipel-

ago, situated beyond the edge of the continent, was not in the main current of the development of the civilization of eastern Asia. Although its flora is fairly rich, no important cultivated crops originated here. The cultivated plants of Japan are all introductions from outside. Similarly the Philippine Archipelago is situated on the extreme eastern periphery of southern Asia. There an exceedingly luxuriant tropical flora exists but scarcely any cultivated crop originated. The only notable plants are a relatively unimportant fiber crop, the Manila hemp, Musa textilis, and another, a nut crop, the pill-nut, Canarium ovatum, which grows wild and is occasionally cultivated. L i t e r a t u r e Cited 1. Ames, O. 1939. Economic Annuals and Human Culture. Cambridge, Mass. 2. Anderson, E. 1960. The evolution o~ domestication. In Evolution after Darwin, vol. III. The Evolution of Man, pp. 67--84. 3. Bishop, C. W. 1933. The neolithic age in northern China. Antiquity 7: 389-404. 4. Burkill, I. H. 1950. The rise and decline of the greater yam in the service of man. Advan. Sci. 7: 443-448. 5. . 1953. Habits of man and the origins of the cultivated plants of the Old World. Proc. Linn. Soc., Sess. 164: 1242. 6. Chatteriee, D. 1951. Notes on the origin and distribution of wild and cultivated rices. Ind. Journ. Genet. & PI. Breed. 11: 18. 7. Darlington, C. D. 1963. Chromosome Botany and the Origin of Cultivated Plants. London. 8. Darwin, C. 1858. Origin of Species. 9. , 1868. Variation in Plants and Animals under Domestication. 10. De Candolle, A. 1855. G6ographie botanique raisonn6e. 11. ,1882. L'origines des plantes cultiv6es. 12. Haudrieourt, A. G. & L. Hedin. 1943. L'homme et les plantes euItiv6es. Paris. 13. Johnson, H. W. & R. L. Bernard. 1962. Soybean genetics and breeding. Advan. Agron. 14: 149--221. 14. Kihara, H. 1959. The origin of the cultivated rice. Kihara Inst. Biol. Res. 10: 63-83. 15. Kitamura, 8. 1951. The origin of the cultivated plants of China. Acta Phytotax. Geobot. 14: 81-86.

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17. 18.

19. 20. 21. 22. 23. 24. 25.

26. 27. 28.

Li, H. L. 1944. The phytogeographic divisions of China, with special references to the Araliaceae. Proc. Acad. Nat. Sci. Phila. 96: 249-277. . 1948. Floristic significance and problems of eastern Asia. Taiwania 1: 1--6. . 1952. Floristic relationships between eastern Asia and eastern North America. Trans. Amer. Philos. Soe., N. S. 42(2) : 371--429. . 1953. Endemism in the ligneous flora of eastern Asia. Proc. 7th Pacific Sci. Congr. 5: 212-216. (1949). . 1960. The cultivation of trees by mankind. Morris Arb. Bull. 11: 15-22. . 1964. The Origin and Cultivation of Shade and Ornamental Trees. Philadelphia. Mangelsdorf, P. C. 1952. Evolution under domestication. Amer. Nat. 86: 6577. Merrill, E. D. 1936. Plants and civilizations. Sci. Month. 43: 430--439. .1938. Domesticated plants in relation to the diffusion of culture. Bot. Rev. 4: 1-20. . 1954. The botany of Cook's voyages and its unexpected significance in relation to anthropology, biogeography and history. Chron. Bot. 14: i-iv, 161384. Moore, Jr., H. E. 1957. Musa and Ensete. The cultivated bananas. Baileya 5: 167194. Osche, J. J. & R. C. Backhauizen van den Brink. 1931. Vegetables of the Dutch East Indies. Buitenzorg. Parthasarathy, N. 1948. Origin of the noble sugarcane (Saccharum officinale L.) Nature 161: 608.

29. 30.

31. 32.

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39.

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Sauer, C. O. 1952. Agricultural Origins and Dispersals. (Bowman Mem. Lect. Ser. II.) New York. Seetharaman, R. 1962. Studies on hybridization between Asian and African species of cultivated rice and their significance. Sci. & Culture 28: 286-289. Simmonds, N. W. 1962. The Evolution of the Bananas. London. . & K. Shepherd. 1955. The taxonomy and origin of the cultivated bananas. Journ. Linn. Soc. Bot. 55: 302312. Swingle, W. T. 1943. The botany of Citrus and its wild relatives of the orange subfamily (family Rutaceae, subfamily Aurantioideae). In Weber & Batchelor, The Citrus Industry 1: 129--474. Toynbee, A. J. 1955. A Study of History. Vol. I. Oxford. 2nd Ed. Vavilov, N. I. 1926. Studies on the origin of cultivated plants. Bull. Appl. Bot. 16: (2) : 139--248. - - . 1935. Theoretical Basis of Plant Breeding. (In Russian). Moscow. . 1949--1950. The origin, variation, immunity, and breeding of cultivated plants. (Transl. by K. S. Chester). Chron. Bot. 13: 1-366. Venkatraman, T. S. 1948. Hybridization in and with the genus Saccharum. Pres. Address in Proc. 25th Ind. Sci. Congr. Calcutta 1-16. Withrow, R. B. 1959. Photoperiodism and Related Phenomena in Plants and Animals. Publ. 55, Amer. Assoc. Advan. Sci. Zudovskij, P. N. 1962. Cultivated Plants and their Wild Relatives. (Abridged translation by P. S. Hudson). Commonwealth Agricultural Bureau.