Dictionary of cultivated plants and their centres of diversity Excluding ornamentals, forest trees and lower plants A.C. Zeven and P.M. Zhukovsky ,$ Centre for Agricultural Publishing and Documentation Wageningen -1975 ISBN 90 220 0549 6 O Centre for Agricultural Publishing and Documentation, Wageningen, 1975 No part of this book may be reproduced and published in any form, by print, photoprint, microfilm or any other means without written permission from the publishers Cover design: Pudoc, Wageningen Printed in the Netherlands by Krips Repro, Meppel Contents Preface 7 History of the work 8 Origins of agriculture and plant domestication 9 Cradles of agriculture and centres of diversity 18 1. Chinese-Japanese Centre 27 2. Indochinese-Indonesian Centre 42 3. Australian Centre 57 4. Hindustani Centre 62 5. Central Asian Centre 71 6. Near Eastern Centre 77 7. Mediterranean Centre 91 8. African Centre 107 9. European-Siberian Centre 129 10. South American Centre 144 11. Central American and Mexican Centre 162 12. North American Centre 173 Species with a not identified centre 180 References 182 Index of botanical names Preface The aim of the work is to give the reader quick reference to the Centre of diversity of a cultivated plant species. For some important crops, the Centre of diversity of related wild species is also presented. These wild species are important sources of genes useful to man when incorporated in cultivated crops. Hence such wild species have to be included in genitor collections for breeding. For some cultivated species, the region of diversity could not be identified. Future research may show where they have come from. Species only cultivated as ornamentals or for timber and lower plants have not been included, as in previous work by Zhukovsky. The cultivated species are arranged alphabetically by families and secondarily alphabetically by ge­ nera within a family and tertiarily by species. For several species, taxonomie synonyms and common names are given, if they seemed to be used. The taxonomie classification of families and genera is based on Willis's dictionary (1966). Somatic chromosome numbers and genome constitutions are presented where known. Most of the chromosome numbers are derived from Bolkhovskikh et al. (1969). Where the chromosome number could not be traced, a space has been left. This work concerns many more species than we could know. Corrections, critisisms and additions inclu­ ding data on chromosome number would be highly appreciated. They should be sent to A. C. Zeven, In­ stitute of Plant Breeding, I.v. P., Agricultural University, Lawickse Allee 166, Wageningen, the Nether­ lands. We hope that this work may directly help the plant breeder and ease shortages of food and other agri­ cultural products. We hope that it will also encourage the establishment of natural wild plant reserves in anticipation of future needs for wild genes. P.M. Zhukovsky A.C. Zeven History of thewor k In 1968Prof . P. M. Zhukovsky published a paper 'New centres of origin and new gene centres of cultivated plants including specifically endemic microcentres of species closely allied to cultivated species'. This paper was issued in Botanical Journal, Moskov 53:430-460 and was abstracted in Plant Breeding Abstracts (1968). I wrote to Prof. Zhukovsky asking whether he would prepare an English version. He wrote back that he was preparing a booklet in Russian on the 'World genofund of plants for breeding: world gene centres of cultivated plants and their wild progenitors', which was published in 1970. The text was translated by Dr E.E . Leppik, Research Botanist of the New Crops Research Branch of the US Department of Agri­ culture, Beltsville, Maryland, who invited me to edit the manuscript and to seek a publisher. The publishers suggested that the work be extended to include more cultivated plants. Prof. Zhukovsky agreed to this proposal and the work has now been enlarged from 700 species to about 2300 species. A. C. Zeven. Origin of agriculture and plant domestication INTRODUCTION The origins of agriculture and plant domestication are, in general, closely related phenomena. How­ ever, plants may be domesticated without actually being cultivated; man may intentionally or unintentio­ nally select for characters which are useful to him. Furthermore, agriculture can be carried out with wild plants. Helbaek (1966) suggested that about 7000 BC. wild barley was cultivated. Ruderal plants (campfollowers, dumpheap plants, habitation weeds) grow in an anthropogen environment which may result in idiotype changes of the plants making them less suited to survive in a wild environment. They have been, however slightly, domesticated. The oldest findings of man and human dwellings point to man being a hunter and collector of plant parts and of small animals like snails, larves, eggs and small birds. Nowadays in many regions, even with a high level of agriculture man still gathers wild and semiwild plants or fruits, such as brambles, blueberries, raspberries, mushrooms, herbs for food, heath for brooms, wood for buildings, fuel or paper-making and grass for domestic animals. However, man does not depend on these plants; he only collects them for economic or recreational reasons. If he depended on them he would grow them or find a substitute. Some people may grow them, while others still collect them. We may ask why man started to cultivate plants, why he started to do so only 'recently' and why only certain plant species or varieties have been domesticated. THE ORIGIN OF AGRICULTURE Much has been written about man's change-over from plant collecting to plant growing. Some authors have put forward 'deterministic' hypotheses, such as a higher mental or social level leading to the culti­ vation of plants, or climatic changes causing a progressive desiccation of the country enforcing the application of artificial methods (Spinden, 1917, see MacNeish, 1964). Sauer (1952) however, thought that agriculture cannot have originated solel from chronic food shortage, as four conditions have to be fulfilled first: 1. Previously acquired skills in other fields to start experiments. 2. A sedentary way of living. 3. The presence of wooded lands which are easier to clear than savannahs or forests. Large river valleys subject to periodical flooding are unsuitable, because man was not able to manage the water problems. ORIGIN OF AGRICULTURE 10 4. A marked diversity of the plant populations so that a large reservoir of genes is available for selection. Sauer concluded that the ancestors of the earliest agriculturists were relatively prosperous, pro­ gressive fishermen living in a mild climate along fresh waters. Actually, not much is known about the skills of the first farmers, and the information on the relation between pre-existing sedentism and incipient food production is limited. Extremely old sites with a year- round occupation have only been discovered in the Nile Valley of Upper Egypt (15 000 to 10 500 BC); they show no evidence of plant or animal domestication (Churcher &Smith , 1972). Perhaps the sites found in southern Africa dating from 47 000 BC. (Border Cave in Zululand), 43 000 BC. (Howieson's Poort near Montagu, southwest Cape Province), 42 000 BC. (Rose Cottage Cave near Ladybrand in eastern Orange Free State) belong to this category. However, no evidence has been found yet to show whether these sites were occupied all year round. The botanical material has not yet been analysed (Dart &Beaumont , 1971; Beaumont &Boshier , 1972). The sites where agriculture developed first must have been in areas where plant collectors/hunters/ fishermen roamed. It is most likely that they lived in the wooded lands for hunting game or near water for fishing. Fishing communities lead a sedentary live; nomads return to sites known to them for the richness in animal and plant food. This may have led to annual occupation of sites, each site for several weeks. On such sites the soil may have become bare because of disturbance by man; paths, loam pits, graves, dilapidated mudhouses, and abandoned compounds in general. Near water natural bare lands such as riverbanks, gravel, rocks, screes, landslides and esturial plains may have occurred. Plants pre- adapted to such environments would colonize such sites. Around dwellings many plants would derive from plant parts collected by man and brought to his house. Pre-adapted plants have a weedy tendency and prefer 'open' rich soil conditions. They grow quickly and have large food reserves which enables them to survive in adverse conditions. These make them very suitable for cultivation. They may grow wild in mountainous or hilly areas with a wide topographical diversity. In such areas with many microclimates variants have most chance to survive. After having moved to a disturbed area, man may have found some useful types among them. Other sites where agriculture may have arisen are the refuse heaps (rubbish heaps, dump heaps) on the compounds. Many parts of plants (fruits, seeds, tubers, roots) must have accidently or purposefully been thrown away. They must have developed into plants with a luxurious growth on these fertile places (Anderson, 1952; Burkill, 1952; Chang, 1970; Engelbrecht, 1916; Flannery, 1965; Harlan &d e Wet, 1965; Hawkes, 1969). The sequence in which agriculture arose may be summarized as follows: 1. Wild plants collected by man. 2. Wild plants moved into the temporary or (semi-)permanent dwelling site either by accident or as a gathered plant part, after which fruits, seeds, parts of tubers etc. were lost or thrown away on purpose. This must have continued for an extremely long time. 3. Only pre-adapted highly variable wild plants established and colonized disturbed soil around the dwel­ ling. Man gathered wanted plant parts from some of these weedy plants. 4. Disadvantageous natural selection pressures were reduced and favourable selection pressures were introduced so that variation decreased counteracted by an increase of variation by hybridization and mu­ tation, followed by isolation, protection and selection.