Dictionary Cultivated Plants

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 a r e 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 a r e arranged alphabetically by families and secondarily alphabetically by g e -
nera within a family and tertiarily by species. For several species, taxonomie synonyms and common
names a r e 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 a r e presented where known. Most of the
chromosome numbers a r e 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 a g r i -
cultural products. We hope that it will also encourage the establishment of natural wild plant r e s e r v e s in
anticipation of future needs for wild genes.
P . M . Zhukovsky
A . C . Zeven
History of thework
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 andwas 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 onthe 'World genofund of plants for breeding: world gene centres
of cultivated plants andtheir 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 USDepartment of A g r i -
culture, Beltsville, Maryland, who invited me to edit the manuscript andto seek a publisher.
The publishers suggested that the work be extended to include more cultivated plants.
Prof. Zhukovsky agreed to this proposal andthe 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 a r e 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
p a r t s 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, r a s p b e r r i e s , 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 a r e 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 r e s e r v o i r of genes is available for selection.
Sauer concluded that the ancestors of the earliest agriculturists were relatively prosperous, p r o -
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 y e a r -
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
F r e e 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 p r e -
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 r e s e r v e s 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 a r e the refuse heaps (rubbish heaps, dump heaps) on
the compounds. Many parts of plants (fruits, seeds, tubers, roots) must have accidently o r 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 &de 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 p r e s s u r e s were reduced and favourable selection p r e s s u r e s were
introduced so that variation decreased counteracted by an increase of variation by hybridization and mu-
tation, followed by isolation, protection and selection. This caused more deviants from the wild phenotype
to survive. Such deviants belong to the ruderal flora or habitation weed flora. This stage could be called
proto-agriculture.
5. The dependence of man on certain plants increased in such a way that when demand exceeded availabi-
lity, man eradicated the wild weedy plant or started taking measures to improve its development. When
man moved outside the natural range of a species on which he depended he was also forced to take measures.
Thus man learned to retain seeds etc. when the plant was to grow outside its natural range, to disturb
the soil on purpose i. e. to cultivate in order to reap a better harvest from the weed now turned into a
crop. This stage might be called incipient agriculture. Modern examples a r e the eradication of some
herbs, ornamentals and mushrooms. Near eradication has lead to incipient cultivation of Tabernanthe
iboga and Camassia leightlinii.
6. Crops were further improved by a semi-intentional and intentional bettering of agricultural methods
and plant types. This stage could be called effective agriculture.
The change-over from food collector/hunter/fisher to full-time agriculturist must have been very
gradual. Once the process started it became practically automatic (Hawkes, 1969) or self-generating.
This gradual change - including the change to animal husbandry - resulted in 1. less energy required to
obtain more food, 2. people becoming tied to the(ir) land and 3. spare time available for other pursuits
(MacNeish, 1964).
Through the invention of agriculture mankind gained more from solar energy. Raising crops (and
husbanding animals) a r e man's most important means of exploiting this energy (Rappaport, 1971).
DOMESTICATION OF PLANTS
The first fully domesticated plants derive from partially domesticated ones like ruderal.plants, habitation
weeds, refuse heap plants. Those plants with a use for man would be protected or at least damage would
be avoided.
Domestication of plants is the change of the idiotype to adapt them better to man-made environments.
Such changes often render the plants less likely to survive in a wild state. The above definition includes
the origin of habitation weeds. In general the term domestication means the unintentional and intentional
selection by man for idiotypes which a r e useful to him. Then the domesticated idiotypes are less suited
to survive in a wild state.
The ancestors of the first crops must have had a weedy character and large food r e s e r v e s to enable
them to survive in very dry summer conditions in poor thin soil free from competition with perennial
plants.
Due to cultivation some plants have changed very quickly, These changes may have been induced by
cycles of differentiation and hybridization between species, forms, ecotypes and races. During differenti-
ation the plants must have grown isolated by genetic, special, cultural and hybridization b a r r i e r s . For
instance hybridization is hampered by a shift from allogamy to autogamy, a shift in time of flowering or
a shift in ecological adaptation. For generatively propagated diploid crop the period of differentiation will
be much shorter than that for a vegetatively propagated polyploid crop. Special b a r r i e r s may be a few
hundred metres, for instance the slopes where wild types grow and the valley floor where the domesticant
is cultivated. Hybridization occurs between the domesticants and weedy or wild relatives often resulting
in a two-way gene flow. When 'cultivated' genes a r e dominant, they have little chance to survive in weeds
or wild plants as is shown for maize - teosinte.
The impact of weedy and wild relatives can be very dominant (large) as has been shown by Heiser
(1965) for sunflower. Because of hybridization, variability will increase and adaptation becomes wider; the
greater the variability and the wider the adaptation, the larger the area where the crop can be cultivated.
Crane (1950) and Masefield et al. (1969) have presented classes of selection schemes from the wild
plant to the present cultivated crop. Both classifications have been used to develop the following one
which is based on taxonomy. It shows the change from a wild species A to new taxons.
1. Domestication, no apparent influx of foreign genie materialI

1.1. cultivated plant morphologically resembles the wild parent, Species A.
1.2. cultivated plant differs morphologically largely from wild parents, Species A, var. B, or Species B
1. 3. autopolyploidization, the newly originated plant may differ from its parent, Species A or Species C
2. Domesticated with influx of foreign genie material
2 . 1 . cultivated plant still resembles morphologically the wild plant (introgression), Species A
2. 2. cultivated plant differs morphologically from the wild parent, Species A var. B or Species D
3. Amphipolyploidization
3 . 1 . between a cultivated plant and a wild one of a different species. Species E
3. 2. between a cultivated plant and another of a different species, Species F
The possible changes of the plant due to domestication have been listed by Polunin (1960) and Purseglove
(1968). The domesticated plants
1. spread to a greater diversity of environments and a wider geographical range,

2. may come to have a different ecological preference,
3. may flower and fruit simultaneously,
4. may lack shattering or scattering seeds and sometimes may have lost the dispersal mechanism com-
pletely,
5. may have an increased size of fruits and seeds, which often reduces the dispersal efficiency,
6. may have been converted from a perennial to an annual,
7. may have lost seed dormancy,
8. may have lost photoperiodic controls,
9. may lack normal pollinating organs,
10.may have undergone a change in its breeding system. Usually the change is from complete or partial
cross-fertilization to partial or complete self-fertilization. This change may be a result from a change
in flower morphology, or a change from self-incompatibility to self-compatibility.
11. may have lost defensive adaptation such as hairs, spines, thorns e t c . ,
12.may have lost protective coverings and sturdiness,
13.may have undergone an improvement of its palatability and chemical composition thus rendering them
more likely to be eaten by animals,
14.may have an increased susceptibility for diseases and pests,
15.may develop seedless parthenocarpic fruits,
16.may have undergone selection for double flowers which may involve conversion of stamens into petals,
17.may be multiplied vegetatively.
The speed of domestication depends on the duration of a generation. For cereals a generation usually
takes one year, while in vegetatively propagated plants no fast changes may be expected. Braidwood &
Howe (1962) estimated that all major changes in wheat and barley had taken place within 2 000 y e a r s .
Helbaek (1966) suggested that this period is 1 500 y e a r s .
Several crops have been domesticated for several purposes. Examples a r e :
Sorghum bicolor: 1. annual forage g r a s s , 2. perennial forage g r a s s , 3. syrup sorghum, 4. grain sorghum,
5. broom corn, 6. popping sorghum used for confectionary, 7. inflorescenses a r e also used for decora-
tion,
Cannabis sativus: 1. fibre, 2. drugs, 3. oil seeds; there a r e also 4. weedy forms,
Brassica napus: 1. rape, 2. swedes, 3. hungary gap kales, 4. oil seed colzas,
Brassica campestris: 1. rapeseed, 2. turnip, 3. leafy vegetables,
Brassica oleracea: 1. vegetable, 2. forage, 3. ornamental, 4. walking stick, 5. construction material,
Helianthus annuus: 1. oil, 2. silage, 3. ornamental, 4. bird's food, 5. ceremonies,

Elaeis guineensis: 1. mesocarp oil, 2. kernel oil, 3. wine
Vicia faba: 1. dry seed, 2. fresh seed, 3. forage (fresh silage), 4. green manure.
This list can easily be extended. Some plants may have been domesticated for a certain use that b e -
came obsolete. If no alternative use is present its cultivation will be abandoned; it will be lost as a culti-
gen, but may survive as a weed or in a living collection. Several crops had two uses or man found a new
use which made them important again. For instance several medicinal crops and herbs a r e also grown
as ornamentals like Viola tricolor and Digitalis purpurea. Some medicinal species are nowadays orna-
mentals only. Similarly fetish plants also became ornamentals. Many fence or stockade plants, which
were planted to stop domestic and wild animals from running away or entering protected a r e a s , a r e used
nowadays as ornamentals or for hedges. Anderson (1960) and Chang (1970) supposed that the first crops
were not food crops. Anderson suggested that plants were domesticated for body paints, living stockades,
poisons, for chewing, for fatigue drugs and for ritual purposes. Chang believed the plants were used for
making containers (bamboo trunks, fruits of bottle gourd), cordage or as herbs. These plants were
needed and when man became dependent on them, he started to cultivate them. Most scientists, however,
believe that food crops are the first domesticants. Burkill (1952) listed the sequence in which he believed
the crops were domesticated:
1. cereals
2. pulses
3. greens
4. oil seeds
5. 'roots'
6. herbaceous fruits
7. fibre
8. woody plants, chiefly fruit t r e e s
9. various industrial plants.
Several wild g r a s s e s a r e very adaptable to domestication: they form many fruits or seeds; they grow
gregariously so that their fruits or seeds could be collectively harvested; they have fruits or seeds
edible for man; the foliage is very excellent for fodder and the seeds a r e good to store. Man did not over-
look these advantages of grasses (Burkill, 1952). Pulses must have followed quickly. Subsequently, seve-
ral greens were also domesticated as oil crops. Many of the woody plants received individual attention.
Purseglove (1968) stated that cereals were first domesticated in the arid and semi-arid regions whereas
in the wet tropics cultivation started with root and tuber crops. Archeological research must elucidate
the right sequence. It may differ from region to region.
The plant families have not contributed equally to the present supply of domesticated species. Harlan
&de Wet (1965) have prepared a list classifying the families as contributing 1. many major crops, 2. a
few major crops, 3. many minor crops and 4. no important crops.
Among the 167 families (see the table) included in the list of this book 51 families a r e represented by
only one species, 23 by 2 items, 12 by 3 items and 82 by more than 3 items. The family with most items
is the Gramineae (359, 15.6%); most of them coming from Africa. This continent is well-known for its
forage g r a s s e s . The Leguminosae follow with 323 items (14.1%); Centres 2, 7, 8, 10 and 9 a r e the rnain
sources. Gramineae and Leguminosae contribute about one third of the list. Rosaceae rank third with 154
items (6.7%), most of them come from Centres 1, 9 and 6.
Families with 100 to 50 items a r e the Solanaceae (100; 4.4%); the Compositae (75; 3.4%); the Myrtaceae
(73; 3.2%); the Malvaceae (67; 2.9%) and the Labiatae (55; 2.8%).
Most Solanaceae come from Centres 10 and 11. 40 of the 73 items on Myrtaceae come from Centre 3.
38 of them belong to the genus Eucalyptus. One third of the Labiatae comes from Centre 7. The Compositae
and Malvaceae have a more even distribution over the various regions.
Centre 2 has contributed the highest number: 303 items, closely followed by Centre 1 (284 items ) and
Centre 8 (276 items). Together they contributed 37.6%. If Centre 9 is added, almost the half (47.5%) of
the items have been included.
Number of items per family p e r region, per family and p e r centre.

There a r e 167 families.
Centre
Family 10 11 12 Un- Total

iden-
tified
Acanthaceae 3
Aceraceae 1
Actinidaceae 4 4
Agavaceae 15 25
Aizoaceae 4
Alismataceae 1 1
Alliaceae 9 21
Alstroemeriaceae 1
Amaranthaceae 1 3 20
Amaryllidaceae 2
Anacardiaceae 2 7 21
Annonaceae 2 14
Apocynaceae 1 11
Aquifoliaceae 1 2
Araceae 1 7 22
Araliaceae 4 4 9
Aristolochiaceae 1
Asclepiadaceae 1 5
Averrhoeaceae 2 2
Balanitaceae 1
Balsaminaceae 1 1
Basellaceae 1 4
Berberidaceae 1
Bignoniaceae 4
Bixaceae 1
Bombacaceae 4 9
Boraginaceae 1 1 6
Bromeliaceae 1
Burseraceae 1 4 7
Cabombaceae 1 1
Cactaceae 8
Campanulaoeae 1 1 3
Cannabidaceae 2 3
Cannaceae 1 1 2
Capparidaceae 1 1
Caricaceae 3 1 4
Caryocaraceae 1 1
Caryophyllaceae 2 2 4
Casuarinaceae 1
Celastraceae 2 1 3
Chenopodiaceae 4 6 6 5 2 1 30
Chloranthaceae 1 1
Chrysobalanaeeae 1 1
Cleomaceae 1 1
Combretaceae 4 4
Compositae 10 7 3 14 6 17 5 4 75
Convoivulaceae 2 1 1 3 2 10
Cornaceae 1 1
Corvlaceae 4 3 1 8
Crassulaceae 1 2 3
Crueiferae 8 4 12 4 11 1 43
Cucurbitaceae 5 2 2 3 11 2 2 7 46
Cupressaceae 1
Cyperaceae 6 3 16
Datiscaceae 1
Dioscoreaceae 2 8 15 29
Dipsacaceae 2
Dipterocarpaceae 1 1
ORIGIN O F A G R I C U L T U R E 16
Centre
Family 6 7 10 11 12 Un- Total

Iden-
tified
Ebenaeeae 3 2
Ehretiaceae 1
Elaeagnaceae 3 4
Elaeocarpaceae 1 1 3
Ericaceae 4
Eryth r o x y l a c e a e 2
Eucommiaceae 1 1
Euphorbiaceae 4 13 10 41
Euryalaceae 1 1
Fagaceae 5 9
Flacourtiaceae 6 9
Geraniaceae 9
Ginkgoceae 1 1
Gnetaceae 1 1
Gramineae 34 44 27 13 35 33 70 38 22 15 25 2 359
Grossulariaceae 3 1 6 2 12
Guttiferae 9 2 15
Hippocastanaceae 1 1 2
Hydrastidaceae 1 1
Hydrophyllaceae 1 1 2
niiciaceae 2 2
Iridaceae 2 1 2 1 6
Juglandaceae 1 1 1 1 2 4 13
Labiatae 5 7 1 1 17 10 10 3 1 55
Lauraceae 2 3 1 2 2 10
Lecythidaceae 2 2
Leguminosae 13 46 23 7 21 48 42 35 41 23 10 6 323
Lemnaceae 1 1
Liliaceae 8 1 1 1 3 2 2 18
Linaceae 1 1 1 1 1 6
Lythraceae 1 1
Magnoliaceae 1 1 2
Malpighiaceae 3 2 5
Malvaceae 6 4 2 6 2 13 1 9 9 1 3 67
Marantaceae 2 2
Martynaceae 1 1
Melastomataceae 1
Meliaceae 3 4
Menispermaceae 1 3
Moraceae 3 6 21
Moringaceae 2
Musaceae 1 4 13
Myricaceae 1 1
Myristicaceae 2 2
Myrtaceae 10 40 13 73
Nelumbonaceae 1
Nyctaginaceae 1 2
Oleaceae 5 11
Onagraceae 2
Orchidaceae 2
Oxalidaceae 1
Paeoniaceae 1
Palmae 1 11 30
Pandaceae 4 5
Papaveraceae 4
Passifloraceae 12 13
Pedaliaceae 1 6
Pentaphragmaceae 1 1
Peperomiaceae 1
Perioplocaceae 2
Phytolaccaceae 1 6
Centre
Family 1 2 3 4 5 6 7 8 9 10 11 12 Un- Total

iden-
fled
Plnaceae 1 1
Piperaceae 5 2 2 1 10
Pistaciaceae 2 2
Plantaginaceae 1 1 2 4
Polygalaoeae 1 1
Polygonaceae 8 1 1 1 1 5 2 20
Potyulacaceae 3 1 1 1 6
Protaceae 3 3
Punicaceae 1
Ranunculaceae 2 2 3 7
Resedaceae 3 1 5
Rhamnaceae 3 3 2 1 9
Rosaceae 40 2 1 25 22 2 1 37 4 1 19 154
Rubiaceae 1 4 4 6 1 17
Rutaceae 12 12 1 4 5 3 2 2 1 43
Salioaceae 1 7 1 9
Sambucaceae 1 1
Santalaceae 1 1
Sapindaceae 3 5 1 1 3 13
Sapotaceae 3 3 2 5 6 19
Saurucaceae 1 1
Saxiphragaceae 1 1
Scrophularlaoeae 1 1 3 5
Simaroubaceae 1 1 1 3
Simmondsiaceae 1 1
Solanaceae 1 2 6 4 2 11 4 34 31 5 100
Sterculiaceae 4 4 2 10
Stilaginaceae 1 1
Strychnaceae 1 1
Styraceae 1 1
Taccaceae 1 1
Tamaricaceae 1 1 2
Taxaceae 2 2
Tetragoniaceae 1 1
Theaceae 4 1 5
Thymelaeaceae 2 1 3
Tiliaceae 1 2 1 4
Trapaceae 3 3
Tropaeolaceae 3 3
Typhaceae 1 1
Ulmaceae 2 1 3
Umbelliferae 5 2 1 2 3 13 10 1 1 38
Urtioaceae 1 1 2 1 2 7
Valerianaceae 2 6 1 9
Verbenaceae 1 1 2 1 5
Violaceae 1 1 1 3
Vltadaceae 2 1 1 7 11
Zingiberaceae 4 16 9 2 1 32
Total 284 303 66 152 79 129 221 276 229 250 181 104 23 2297
% of total 12.4 13.2 2.9 6.6 3.4 5.6 9.6 12.0 10.0 10.9 7.9 4.5 1.0 100
Cradles of agriculture and centres of diversity
A search for the geographical distribution of centres of plant domestication can not be carried out without
studying the origins of agriculture, hearths or cradles of agriculture and the spread of agriculture. The
latter may include a study of the spread of domesticated plants.
At present wild plants are still taken into cultivation, whereas an important crop like the oil palm in
Africa is still largely semi-domesticated (Zeven, 1967, 1973). Other examples a r e the secondary crops
i. e. crops which were first weeds in primary crop but developed later into a crop.
Sites of early farms have been discovered in Thailand, Near East and Mexico. They showed that in-
cipient agriculture existed in Thailand at about 11 000 BC. (Gorman, 1969), in the Near East at about
9 000 BC. (Cambel &Braidwood, 1970) and Mexico at about 6 000 BC. (MacNeish, 1964a, 1964b). In
other areas no such sites have (yet) been found, and at present it is accepted that from these cradles of
agriculture, agriculture spread to other parts of the world. So agriculture may have reached China and
Japan, and SE. Asia from Thailand, while agriculture may have reached Europe, Africa, W. Asia, SW.
Asia and S. Asia from the Near East.
Probably Alexander Von Humboldt was the first to refer to the origin of crops. In his work 'Essai
sur la géographie des plantes' (1807) he said: 'The origin, the first home of the plants most useful to man
and which have accompanied him from remotest epochs, is a secret as impenetrable as the dwellings of
all our domestic animals. We do not know what region produced spontaneously wheat, barley, oats and
rye. The plants which constitute the natural riches of all inhabitants of the tropics, the banana, the
pawpaw, the manioc, and maize have never been found in wild state' (cited by Hawkes, 1970). If alive Von
Humboldt would be delighted to learn about the present available knowledge on the origin of cultivated
plants.
The next study was by Alphonse De Candolle in 'Géographie Botanique Raisonée' (1855). He was
followed by Charles Darwin in 1868 with his book 'Variation of animals and plants under domestication'.
However, Darwin was not interested in the study of the origin of the cultivated plants, but in the study of
evolution of animals and plants.
De Candolle's first real effect in tracing the origin of the cultivated plants was published in 1882 in
his book 'Origine des plantes cultivées'. This work is still very up-to-date (Harlan, 1961). De Candolle
based his investigations on 1. Classical botany (plant geography, knowledge of adventive and ruderal
species, understanding of history of development of whole floras), 2. Bio-archaeology (plant remains,
pictorial records, especially from Egypt), 3. Palaeontology and 4. Philology. He concluded that the region
where a species is abundant is not necessarily its centre of origin. Perhaps De Candolle (1882) was the
CRADLESOFAGRICULTUREANDGENECENTRES 19
first to indicate regions where the first plant domestication might have taken place: 1. China, 2. SW.
Asia and Egypt, 3. Tropical Asia (Smith, 1968). In De Candolle's time it was quite natural to include
Egypt as much of the knowledge of plant history came from that country.
After De Candolle, Nicolai Ivanoviß Vavilov indicated the cradles of agriculture. Vavilov at the
height of his c a r e e r had more facilities at his disposal than anyone before (Harlan, 1951). His abundant
energy made full use of them. During the Fifth International Genetics Congress at Berlin in 1926, Vavilov
(1927) developed his theory of centres of origin or gene centres: some regions of the world possess a
concentration of variations of certain cultivated plants; for several cultivated plants these regions overlap.
These regions can be identified by the Differential Method. This method is simply described by Burkill
(1952):
1. Take a map,
2. select important cultivated plants,
3. mark on the map the sites where recognizable botanical varieties, races of these cultivated plants
a r e found. The identification of the botanical varieties was done by investigating the morphology, cytology,
genetics and resistance to diseases, pests and unfavourable climatic conditions of the plants.
4. Where those marks accumulate is a centre of origin. In such centre the greatest diversity of the cul-
tivated crop is observed.
Vavilov concluded that a centre of origin was characterized by dominant alleles while towards the
periphery of the centre, the frequency of recessive alleles increased and the diversity decreased. The
cause was inbreeding and geographical isolation (drift).
At the periphery secondary gene centres may develop: new areas with a great diversity conditioned
by recessive alleles. In 1926, Vavilov reported that Asia Minor lies in the Asiatic, Mediterranean, Bal-
kan and Transcaucasian gene centres of wheat and other crops. In 1931, he extended this idea by d i s -
tinguishing seven gene centres. In 1935 he brought this number up to eight by splitting Southwest Asia
into Central Asia and the Near East. Later Zohary (1970) proposed to unite them again.
I. China
II. India
IIa. Indo-Malava
III. Central Asia including Pakistan, Punjab, Kashmir, Afghanistan and Turkestan (USSR)
TV. wear East
V. Mediterranean Sea coastal and adjacent regions
VI. Ethiopia
VII. South Mexico and Central America
VIII. South America (Peru, Ecuador, Bolivia)
Villa. Isle of Chiloe (Chile)
These centres a r e all between 20 and 45 in mountainous regions and often in areas with a t e m p e -
rate climate. They a r e separated by great d e s e r t s . According to Vavilov agriculture in these eight regions
developed independently, because of the differences in agricultural methods, implements and domestic
animals.
Vavilov may have been influenced by Willis' Age and Area hypothesis (Willis, 1922). It states that,
in comparing wild species with similar modes of dispersal, those with the wider distribution a r e older
and that the longer a species has been present in an area, the more diverse will be the derived species
and subspecies found there. Vavilov may also have been influenced by the agro-geographical work of
Engelbrecht (Zeven, 1973). At present, it is known that time is not the only factor that influences the d i s -
persal of a species and its increase of variation.
In the thirties Vavilov established an 'ecological passport' for the accessions of his large collections
by sowing them at various sites ('geographical sowing') after which he estimated:
1. differences in growth during the vegetative period.

2. differences in length of the various development stages, including growth rhythm,
3. economic characters, such as size of fruits and seeds,
4. vegetative characters,
5. resistance to different kinds of drought,
6. resistance to cold,
7. specific differences in flowering,
8. resistance to various fungi,
9. resistance to different bacteria and viruses,
10.resistance to various insects,
11.ecological type of plant: xerophyte, hydrophyte, mesophyte.
The diversity was enormous but within limits and with certain regularity. Vavilov discovered
'parallelisms which a r e especially clear for plants which belong to the same general group (annuals,
herbaceous) and which a r e characterized by the same area of distribution and have followed geographi-
cally the same route in their evolution. As it appeared that each species has differentiated into different
agro-ecological and geographical groups, he was able to establish the 'ecological passport' for annual
cereals, grain legumes, oil and fibre flax. In 1940 Vavilov divided the Old World (excluding Africa south
of N. Africa, tropical Asia) into 19 a r e a s , each characterized by the plants with in general the same
'ecological passport':
1. Syrian Group Agricultural territory: chiefly foothills of Syria, Palestine and Jordania. C h a r a c t e r i s -
tics of cultivated and wild plants: relatively small; with small leaves, flowers and seeds; thin, stiff stems;
non-shattering spikes or indéhiscent pods; high maturing temperature; short vernalization stage.
Examples: types of wild and domesticated Triticum species; barley; oats; peas; lentils; g r a s s - p e a s ;
chick-peas; domesticated flax and vetch.
2. Anatolian Group Agricultural territory: mountainous parts of Turkey. Characteristics: medium-size;
thin, stiff stems; medium-sized spikes, fruits and seeds; resistant to drought; short development stages;
requiring considerable warmth during last stage of development. Examples: same as preceding group.
3. Armenian Xerophytic Mountain Group Agricultural territory: arid, mountainous steppes of Soviet and
Turkish Armenia, Characteristics: markedly xerophytic (small narrow Leaves); small seeds. Examples:
Triticum vavilovii (also resistant to shattering, and winterhard); early dwarf, small seeded, xerophytic
chickpeas; a large number of relatives of domesticated wheat; Secale vavilovii.
4. Caucasian Mesophytic High-Mountain Group Agricultural t e r r i t o r y : high mountain plateaux of Daghestan
and Georgia, Northern Armenia. Characteristics: thin stems; comparatively smooth awns; small or
medium-sized seeds; short or medium vegetation period. Examples: original ecotypes of soft wheat;
prototypes of European steppe winter and spring bread-wheats; Triticum carthlicum; a specific group of
barley with narrow leaves; many xerophytic and mesophytic types of Secale montanum and S. cereale ssp.
segetale (many with a great diversity of r e d and brown forms).
5. Daghestan-Azerbaijan Foothill Group Agricultural t e r r i t o r y : coastal regions of Daghestan and A z e r -
baijan. Characteristics: mesophytic; long vegetation period; tall; leafy; large seeds; rather resistant to
leaf rust. Examples: giant forms of soft and durum wheats; barley, rye; peas, vetch, winter types of
durum.
6. Transcaucasian Humid Subtropical Group Agricultural t e r r i t o r y : West Georgia and Black Sea coast,
humid regions of Turkey-and Southern Azerbaijan (Lenkoran), Northern Iran. Characteristics: hydro-
phytic, tall, leafy; late; rather resistant to different European fungus diseases. Examples: endemic
Triticum ssp. such as T. macha and T. timopheevi, and some other diploid and tetraploid Triticum types;
late types of prostrate fibre-flax sown in autumn and winter; transitory and very late spring varieties of
cereals.
7. Iran-Turkestan Group Agricultural t e r r i t o r y : irrigated and non-irrigated regions of Iran, Afghanistan,
Soviet Central Asia (Uzbekistan, Tadjikistan, Turkmenistan). Characteristics: low to medium high;
rather non-shattering rough spikes; weak stems subject to lodging; slow growth during early stages of
development; drought-resistant during late stages; high temperature requirement at maturing; extremely
susceptible to all European fungus diseases when sown in steppe or wooded steppe regions of Europe.
Two subgroups:
a) Khiva subgroup: near mouth of Amu-Darya river, characterized by late varieties of wheat, barley,
flax and peas;
b) Kashgar subgroup: high plateaux near the Pamir, includes extremely cold-resistant varieties of soft
wheat and relatively late varieties of flax (frequently with white flowers and seeds).
8. Pamir-Badakhstan Group Agricultural territory: Soviet and Afghan Badakhstan (Pamir agricultural
district), Central and North Kafirstan, at very high altitudes (up to 3000 m and more). Includes types
from Upper Himalayas and Tibet. Characteristics: mesophytic types; of medium height; broad leaves;
short vegetative period; extremely susceptible to all European fungus diseases. Furthermore a gigantic
type of rye with large anthers and pollen grains, big kernels and large spikes: liguleless, soft and com-
pactum wheat; large broad-leaved, naked, six-rowed barley; small seeded, early peas, beans and grass -
peas.
9. Indian Group Agricultural territory: Northern India. Characteristics: as those for the P a m i r -
Badakhstan Group; notwithstanding the diversity in ecological circumstances quite uniform; not bushy;
thin, stiff stems; small narrow leaves; early; short; development stages and rapid development rhythm;
resistant to drought; needs high temperatures, especially during last stages of development; rapid
filling-out of seeds; small seeds (in cereals, flax and grain legumes); spikes (of wheat and barley) not
rough; grain (ditto) non-shattering.
In Kashmir a subgroup has been established based on a special wheat type characterized by medium height,
thin stems, long narrow leaves, small kernels, rather smooth awns, winter habit, and less susceptibility
to brown rust than the plants of Group 7. (The reason why groups 8 and 9 have been separated, despite
the identity of the characteristics, is not stated. )
10. Arabian Mountain Group Agricultural territory: Yemen, where high-mountain agriculture is subject
to the influence of the surrounding deserts. Characteristics: short spring annuals with extremely rapid
growth; thin, stiff stems, narrow leaves; relatively large seeds. No examples a r e given.
11. Ethiopian (Abyssian) Group Agricultural territory: Ethiopia and Eritrea. Divided into two subgroups:
a) varieties sown at beginning of main rainy season: cosmopolitic, hydrophytic types of tall, l a r g e -
seeded varieties of barley and peas (Ethiopian wheats, though not outspokenly cosmopolitic, may be in-
cluded here);
b) varieties sown at the end of the rainy season: flax, chickpeas, lentils, beans, grass-peas, and an
Arabian type of pea (xerophytic, early, low, small-leaved, small-seeded). Origin: very probably linked
with India and mountainous Arabia.
12. Chinese-Japanese Group Agricultural territory: China and Japan. Very likely, the original material
was imported, several millenia ago, from Asia Minor by way of India, but very important new characters
have developed in this group. Characteristics: short development stages; low or medium height; extremely
small seeds; rapid filling of grains. Examples: rapidly filling wheats with small kernels, awnless or
awnletted.
13. Mediterranean Group Agricultural territory: Mediterranean area. Characteristics: rather tall, bushy;
large spikes; long awns; large, light-coloured seeds; high yields; usually solid straw, short first develop-
ment stage; resistant to low air humidity, requiring much warmth at maturity; resistant to fungal di-
seases. No examples a r e given.
14. Egyptian Group Agricultural t e r r i t o r y : Egypt. Characteristics: barley and durum with short, stiff
stems, medium-sized spikes, and short first development stages. Similar types have been found on
Cyprus.
15. South-European Group Agricultural territory: Southern France, Northern Italy, part of Yugoslavia,
Bulgarian coast. Charateristics: tall plants; large leaves; big fruits; high yields. Examples: Triticum
turgidum s . s . , soft wheats; in Lombardy giant forms of oats, chickpeas, horse beans, and a polonicoid
wheat, have been found.
16. European Steppe Group Agricultural territory: European steppes from Tirol to the Urals; transferred
to North America, especially to the p r a i r i e s . Examples: xerophytic spring and winter types of cereals
and grain legumes, the winter types winterhard, the spring types drought resistant; rather small seeds,
weak straw, narrow leaves. (Vavilov divided this Group into two subgroups, but he gives no grounds for
this division. )
17. West-European Group Agricultural territory: Western Europe including South Finland and South
Sweden. Characteristics: tall, hydrophytic plants; thick; stiff stems; large, broad leaves; large, dense,
highly productive spikes; medium-sized or large grain; ripening late. Local varieties have lax spikes,
a r e tall and early.
18. Central-European Group Agricultural territory: forest and wooded steppe area of Central Europe.
Characteristics: high yielding mesophytes. Examples: long-fibre flax, highly productive peas, awnless
soft wheats.
19. Northern (Boreal) Group Agricultural territory: Northern European USSR, Siberia, North Scandinavia.
Characteristics: mesohydrophytic; precocity; medium sized; low warmth requirement; cold-resistant.
Examples; self-compatible rye and very early types of forage barley.
Vavilov worked on his concept of gene centres, modifying it, until his death. These agro-ecological
groups need not coincide exactly with the gene centres. The purpose of all his effort is obvious, however:
there a r e groups of plants possessing certain characteristics not present in other groups. So, when
looking for a certain property in a species, it is not necessary to study its entire a r e a of distribution,
but it is sufficient to look for it in the group(s) where this property has already been found.
The gene microcentres as Harlan (1951) called them form another breakdown in the geobotanical
patterns of variation. They a r e areas of relatively small size in which evolution is still proceeding at a
rapid rate.
F o r wheat, Harlan identified three such microcentres in Turkey. Outside this country undoubtedly
many more exist. With the introduction of high-yielding foreign wheat varieties these microcentres have
disappeared.
Harlan also identified gene microcentres in Turkey for a number of other crops. He found that such
centres frequently coincide. They may be located in the plains or in mountainous regions, near civili-
zation or remote from it, in areas with very primitive or more advanced husbandry.
With increasing knowledge of cultivated, weedy and wild plants it became evident that some parts of
Vavilov's theory had to be changed (for a literature review see Kuckuck, 1962). Nevertheless it still
forms a good base with which to search for wild or semi-wild relatives. The large collections made by
Vavilov and his introduction of the genetic element in the investigations, still render these discussions
very valuable. One point in Vavilov's theory is that a primary centre is marked by a high frequency of
dominant alleles. Gökgöl (1941) showed that it was impossible to indicate such a centre for wheat.
Brieger (1961) did not find one for maize, Zeven (1967, 1972) not for oil palm and Hanelt (1972) not for
Vicia faba. Furthermore, it has been pointed out that a great diversity may also a r i s e from the variation
of the environment. Hence the relation between mountain regions and centres of origin. Such a great
diversity may also develop when two populations of a (partial) cross-fertilizing species meet, as has been
shown for Carthamus tinctorus. Vavilov's theory that where the greatest diversity is found is also the
centre of origin, is no tenable, as was shown for crops like Triticum dicoccum and Hordeum vulgare in
Ethiopia: they show a great diversity there, but no wild relatives a r e present.
Kuckuck (1962) concluded that Vavilov certainly would have altered his theories with the present
available knowledge. Indeed he introduced changes during his research in the course of the years.
The number of cradles of agriculture has been much discussed. Vavilov believed in many, others
suggested two (Sauer, 1952): one for the Old World: Birma and adjacent area and one for the New World:
C. America. Darlington (1952, 1969) also suggested two: 1. the Fertile Crescent of the Near East and 2.
Mexico. From these nuclear areas agriculture would have spread over the Old World and the New World,
respectively. After the introduction of agriculture new centres of plant domestication developed. Thus,
Darlington & Janaki Ammal (1945) distinguished twelve 'centres of origin':
1. Ethiopia
2. Mediterranean coast
3. Iran, incl. the Caucasus and Eastern Turkey
4. Afghanistan
5. Indo-Burma
6. Siam-Malaya-Java
7. China
8. Mexico
9. Peru
10. Chile
11. Brazil-Paraguay
12. USA
Gene c e n t r e s of cultivated plants of Darlington &Janaki Amma l (1945) based on Vavilov
As compared with list on p. 22 continental Chile instead of the Isle of Chiloe, the Brazil-Paraguay
and the USA a r e added.
Darlington & Janaki Ammal considered the Mediterranean centre as a diffuse one. It is based on
'cultural rather than botanical considerations. The Mediterranean, a b a r r i e r to wild plants, has been a
means of dispersal and a bond of union for plants of established cultivation'.
In 1956, Darlington added Europe (for no indicated reason), Central Africa (perhaps based on P o r -
t ê r e s ' views - see below) and Central America) already mentioned by Vavilov). He furthermore used,
without explanation, the term 'region', though the captions of his table and figure still mention ' c e n t r e s ' .
This resulted in
1. Southwest Asia 7. China

2. Mediterranean 8. Mexico
2a. Europe 8a. USA
3. Ethiopia 8b. Central America
3a. Central Africa 9. Peru
4. Central Asia 9a. Chile
5. Indo-Burma 9b. Brazil-Paraguay
6. Southeast Asia
In 1950, P o r t ê r e s suggested independent cradles of agriculture in Africa south of the Sahara. One
was located is East Africa, the other in Tropical West Africa. He divided the latter into 1. the Senegam-
bian 'Subcradle', 2. the Central Niger 'Subcradle', 3. the Benin 'Subcradle' and 4. the Adamawa 'Sub-
cradle'. Other African cradles of agriculture stood in North Africa and Ethiopia. In 1962 he changed his
concept by dividing and subdividing Africa into:
West African cradle B. Nilo-Abyssinian cradle

I. Tropical sector I. Nilotic sector
a. Senegambian subsector II. Abyssinian sector
b. Central Niger subsector C. East African cradle
c. Chad-Nilotic subsector D. Central African cradle
II. Subequatorial sector
The Nilo-Abyssinian cradle coincides with Vavilov's Ethiopia and a part of the Mediterranean centre
of origin. The last two cradles have not further been elaborated. P o r t ê r e s (1950, 1962) decided on a West
African cradle because of the presence of several crops typical to that area. In this he was supported by
Murdock (1959), who established four regional agricultural complexes:
1. Southwest Asian agricultural complex-developed by the Caucasoids

2. Southeast Asian complex developed by the Mongoloids
3. Central American complex developed by the American Indians
4. West African complex developed by the West African Negroes.
His decision on an independent West African agricultural complex is based on grounds similar to
those of P o r t ê r e s .
Anderson (1960) started from quite another characteristic in dividing agriculture into floral and non-
floral seed crop agriculture in Central Africa and a pole of floral agriculture in Indonesia. He supposed
that the floral type of agriculture spread into Oceania, China and Japan, India and Afghanistan, while the
non-floral type remained in Africa. The almost complete lack of interest in flowers and ornamental plants
among the African peoples is really astonishing, whereas in the region of the floral type even the poorest
man grows some ornamentals (Anderson, 1960). This is not due to an absence of ornamental species in
Africa: many a r e commonly grown elsewhere now. The claim of an African cradle of agriculture has been
refuted by Wrighley (1960), Clark (1962), Baker (1962), Harris (1967) and Harlan (1967). Baker (1962)
summarized his objections as follows:
1. few of the domesticates a r e definitely known to derive from W. Africa;
2. several of the domesticates have so little differentiated from the wild that they cannot be of great
antiquity as cultivated plants;
3. if cultivation had been practised locally for seven millenia, an associated weed flora rich in indigenous,
species should to have evolved.
Harris (1967) concluded that the typically W. African crops a r e local additions to an intrusive a g r i -
cultural complex, rather than compounds of an ancient indigenous one. After the introduction of agricul-
ture into N. Africa it spread into the Sahara. Owing to the desiccation of this area in the third millenium
BC. agriculture became established in the savanna zone stretching from the Atlantic to the Lake Chad and
further on to the Cape Horn in East Africa. It is in this centre the many typically African plants, listed
by Harlan (1971), were domesticated.
Kupzov (1955, cited by Darlington, 1956) showed which regions of the world belong to certain hearths
of agriculture. He identified ten, grouped into 5 'main agricultural regions':
Hearth of agriculture Main agricultural region
1. Indian
I. Australoid
2. Indonesian
3. Chinese n. Mongoloid
4. Central Asiatic
5. Near East m. Europoid
6. Ethiopian
7. Mediterranean
8. Nigerian rv. Negroid
9. Mexican
10. Peruvian
v. Americanoid
Except that of Nigeria they derive from a neolithic stage. Darlington (1956) who cited Kupzov does
not explain why Kupzov came to this classification.
Zhukovsky (1965) was the first to refer to Siberia as a gene centre. Many Malus, Prunus, Pyrus and
other species have been domesticated there. Further it is a rich source of wild relatives of these species.
Primary regions of agriculture ( )and regions of expansion ( ) of Darlington {1956) based on Kupzov (1955)
In 1968 Zhukovsky heralded his idea about 'megagene c e n t r e s ' . As so many crops originate outside one
of Vavilov's centres of origin it was necessary to enlarge the areas in which species were domesticated.
These megacentres engulf much of world's land surface and adjoin over great distances. They a r e :
1. China 7. Mediterranean coastal and adjacent regions

2. Indochina-Indonesia 8. Africa (8a Ethiopia)
3. Australia-New Zealand 9. Europe-Siberia
4. Indian subcontinent 10. Central America
5. Central Asia 11. Bolivia-Peru-Chile
6. West Asia 12. North America
Megacentres of cultivated plants of Zhukovsky (1968)

Centres and noncentres of agricultural beginnings of Harlan (1971)
The centres 1, 2, 4, 5, 6, 7, 8a, 10 and 11, although much enlarged, a r e based on Vavilov's concepts.
Zhukovsky proposed as new ones 3 (Australia-New Zealand), 8 (whole Africa) and 9 (Siberia), 12 (North
America) has already been presented by Darlington & Janaki Animal (1945) and 9 (Europe) by Darlington
(1956). Zhukovsky (1968) did not draw boundaries between 2 and 4, 5 and 6.
In 1970 Zhukovsky made some amendments: some megacentres were enlarged, and boundaries were
drawn between 2 and 4, and 5 and 6. Obviously the greater the number of investigated crops the larger
the a r e a s . Therefore Harlan (1971) developed the idea of centres and noncentres. He suggested that a g r i -
culture began independently in three areas and that there was a system composed of a centre and a non-
centre, hi a noncentre, agriculture has been introduced after which many indigenous plant species were
domesticated. Harlan (1971) preferred the t e r m noncentre because of the large area involved. His c l a s s i -
fication is:
Centre Noncentre
A l . Near Eastern A2. African

B l . Chinese B2. Southeastern Asian and S. Pacific
CI. Central American C2. South American
Important crops domesticated in the noncentres may in some cases have spread to its centre in early
times.
Zhukovsky's (1970) classification has been used as the base of the following list, though possibly
some megacentres still have to be enlarged till at least they cover most of the world's surface. This holds
especially for South America where a shift of the eastern boundary may include Brazil and Paraguay and
the land west of these countries, as proposed by Darlington & Janaki Ammal (1945).
Zeven (unpublished) preferred the term Region instead of Megacentre.
Future research has to show whether there have been three cradles of the origins of agriculture:
1. Eastern Asia (China and Birma), 2. the Near East (Fertile Crescent) and 3. Central America and
how from these cradles agriculture spread over the world.
A. C. Zeven.
1 Chinese-Japanese N
__ r
/""
Centre
j '-^S^
n
The Chinese-Japanese centre was called by Vavilov the East Asian Centre of Origin. It is mostly situated
in China. For several crops is Japan a secondary centre of diversity. Sometimes this region is associated
with the primary region of diversity of fruit crops in Amur-Ussuri area. Li (1966 quoted by Chang, 1970)
divided China in two a r e a s : 1. N. China with a seed and vegetable agriculture and 2. S. China which forms
a buffer zone between N. China and centre 2 (vegetatively reproduced crops). Chang (1970) and Harlan
(1971) suggested an independent origin of agriculture in N. China, which resulted in a wholly original
assemblage of cultivated plants. Harlan called this centre Bl North Chinese centre.
The earliest site of agriculture lies at Yang-Shao. It is a strictly Chinese centre with no appreciable fo-
reign influence upto 1300 BC. It is assessed at present about to be 4th millenium BC. It is very probably
that still older farming sites will be found (Harlan, 1971).
China is one of the richest regions contributing many important crops particularly fruit t r e e s . Other i m -
portant crops a r e Brassica campestris and related species, Camellia sinensis, Colocasia esculenta,
Corchorus sinensis, Glycine max, Panicum milaceum, Raphanus sativus, Setaria italica etc. It is a s e -
condary centre for Oryza sativa ssp. japonica, Zea mays and other crops.
Actinidiaceae ACTINIDIA POLYGAMA Miq. Silver vine. 2n=c. 58,

c. 116. N. and W. China, Korea and Japan. A
ACTINIDIA ARGUTA Sieb. & Zucc. Tara vine. polygamous, trioecious ornamental. In Japan the
2n=c. 116. China, Japan, Korea and the Primorye leaves a r e boiled and eaten.
Territory, USSR. Very frost resistant; Used in
c r o s s e s with A. chinensis* (Schroeder & Fletcher, Alismataceae
1967).
SAGITTARIA SAGITTIFOLIA L. Arrowhead.
ACTINIDIA CHINENSIS Planch. Chinese goose- 2n=22. Europe and Asia. A herb cultivated in China
berry, Strawberry peach, Yang tao. 2n=c. 116, and Japan for its edible corms.
c. 160. W. and C. China. Extensively cultivated
in the Yangtse valley and elsewhere for its large, Alliaceae
fragrant, juicy fruits. Luther Burbank used it as ALLIUM CHINENSE G. Don (syn. A. bakeri Regel).
a pollen donor with the frost resistant A. arguta*. Rakkyo, Ch'iao T'ou. 2n=16, 32. S. China (Li,
1970). Cultivated in China, Japan, California and
ACTINIDIA KOLOMICTA Maxim. Kolomikta. elsewhere by the Japanese and Chinese.
2n=c. 112. NE. China and the Primorye Territory,
USSR. Very winterhardy. With delicious berries ALLIUM FISTULOSUM L. Welsh onion, Cibol,
containing much Vitamin C. It is cultivated. Stone leek, Spring onion. 2n=16. Siberia and China
(Li, 1970). Cultivation started probably in N.
China. Cultivated in China and Japan. Related to
A. altaicum Pall. (2n=16) from N. Mongolia. A.
wakegii Araki. (2n=16) and A. microbulbum Prokh.
THE CHINESE-JAPANESE CENTRE 28
The latter is considered a hybrid of A, fistulosum* Araceae

and A. altaicum.
COLOCASIA ESCULENTA (L. ) Schott var. anti-
Cultivars with blue-green leaves and white bulb
quorum (Schott) Hubbard &Rehder (syn. C. anti-
are sometimes separated as A. bouddbae O. Deb.
quorum Schott, C. esculenta var. globulifera Engl.
(2n=16)(Purseglove, 1972).
&Krause). Eddoe, Taro, Dasheen. 2n=28, 42.
SE. Asia (p.000). Many socalled wild specimens
ALLIUM LEDEBOURIANUM Roem. & Schult.
a r e probably derivatives of run wild plants. From
Asatsuki. 2n=16. From USSR to Japan. Cultivated
SE. Asia it spread to China and Japan where var.
in Japan (Kihara, 1969). antiquorum developed. In 500 AD. some cultivars
a r e mentioned in China (Li, 1969). At present
ALLIUM MACROSTEMON Bunge. Chinese garlic. many cultivars are described. Some of them are
Chromosome number varying with parts of the triploid (Bai et a l . , 1971). Their vernacular
plant from diploid (2n=2x=18) to hexaploid (2n=2x= names are also used for Xanthosoma spp. Dasheen
72). Including aneuploids. Ancient Chinese garden is a corruption of 'eddo de la China'.
plant with very big bulbs. Introduced in W. Geor-
gia (USSR) during the Middle Ages. In Japan a. secondary centre of diversity deve-
loped.
ALLIUM NIPPONICUM Franch. &Savat. 2n=16,
32. Formerly cultivated in China but now it only Araliaceae
grows wild there (Li, 1969). ARALIA CORDATA Thunb. Udo. 2n=28. Japan.
Cultivated in Japan as a vegetable (Kihara, 1969).
ALLIUM RAMOSUM L. Chinese leek. 2n=32. N.
China and Siberia. Cultivated in N. China. An PANAX GINSENG C.A. Meyer. Ginseng. 2n=44.
autotetraploid. It differs from A. porrum*. Ussuri region, China, Manchuria and Korea. E x -
terminated in the Chinese provinces, Shansi and
ALLIUM SATIVUM L. Garlic. 2n=16, genome Shensi. There it was cultivated for a long time in
formula SS. C. Asia (p. 71). Var. pekinense Ma- SE. Manchuria, N. Korea, Japan and also USA
kino sometimes considered a native of N. China. and USSR (Baranov, 1966).
Cultivated in N. China and Japan (Li, 1969).
PANAX REPENS Max. (syn. Aralia repens Max. ).
ALLIUM SCHOENOPRASUM L. Chive. 2n=16, (24, China and Indochina. A herb cultivated in Yunaran,
32). Europe, Asia and N. America. Very poly- China and elsewhere for its medicinal roots.
morphous. Domesticated in USSR (region not given)
(Kazakova, 1971). Cultivated over the whole world. TETRAPANAX PAPYRIFERUM (Hook. ) Koch.
Rice-paper plant. 2n=24. N. Formosa and S. China
ALLIUM TUBEROSUM Rottl. ex Spreng, (syn. A. (Hunan, Szechwan, Yunnan, Kweichow, Kwangsi
o d o r a t u m L . ) . Kui t s ' a i , Nira, Chinese chive. and Kwangtung provinces). Cultivated in the (sub)
2n=16, 32. P r i m a r y centre of origin unknown, as tropics as an ornamental (Perdue &Kraebel, 1961).
it easily runs wild (Jones &Mann, 1963). At p r e -
sent from E. Mongolia to Japan, the Philippines Balsaminaceae
and through Thailand to N. India. Its tetraploid e
type may derive from an autotetraploidization of a IMPATIENS BALSAMINA L. Balsam, Garden b a l -
diploid species or from an amphiploidization of a samine. 2n=14. Indo-Malaya and China. Cultivated
in China as a cosmetic plant and elsewhere as an
hybrid of two diploid species. Cultivated in China
ornamental.
for its edible leaves and young inflorescenses,
and as an ornamental.
Boraginaceae
Amaranthaceae LITHOSPERMUM OFFICINALE L. Var. erythrorhi-
zon (Sieb. & Zucc. ) Hand.-Mazz. (syn. L. m u r a -
AMARANTHUS GANGETICUS* saki Sieb., L. erythrorhizon Sieb. & Z u c c ) .
2n=28. Cultivated in N. China and Japan for a red
dye. Ssp. officinale is cultivated in Bohemia
Anacardiaceae (p. 129).
RHUS SUCCEDANEA L. Waxtree. 2n=30. China
and Japan. Burseraceae
CANARIUM ALBUM (Lour. ) Raeusch. White Chi-
RHUS VERNICIFERA DC. (syn. R. verniciflua
nese olive. 2n= . China. Cultivated in S. China
Stokes). Varnish t r e e . 2n=30. China and Japan.
and Cochinchina.
It is the source of a varnish, Japanese lacquer.
Cabombaceae
Aquifoliaceae BRASENIA SCHREBERI J. F. Gmel. Watershield,
Junsai. 2n=28. Asia, Africa, Australia and N.
ILEX INTEGRA Thunb. 2n= . Japan. A tree
America. Cultivated in Japan as a vegetable (Ki-
cultivated for its bark which is pounded and used hara, 1969).
as bird lime.
ALLIACEAE - CRUCIFERAE 29
Campanulaceae LACTUCA INDICA L. Indian lettuce. 2n=18. India,

PLATYCODON GRANDIFLORUM DC. Chinese Japan, Philippines and Indonesia. Cultivated in
bellflower. 2n=(16), 18, (28). Cultivated in China China, Japan and other countries. Many varieties
and Japan as a medicinal crop. exists in China.
Celastraceae PETASITES JAPONICUS F. Schmidt. Fuhi. 2n=87.

Sachalin and Japan. Cultivated for its flower buds
EUONYMUS JAPONICUS Thunb. (syn. E. pul- and leaf stalks (Uphof, 1968; Kihara, 1969).
chellus Carr. ). 2n=32. S. Japan. A shrub. Culti-
vated in Spain and elsewhere for rubber. XANTHIUM STRUMARIUM L. Cocklebur. 2n=36.
In China it was used as vegetable. Nov: it is a weed
TRIPTERYGIUM WILFORDII Hook,f. 2n= in fields and along roadsides (Li, 1969).
China, Japan and Taiwan. Cultivated in Chekiang,
China as a source of insecticide. Convolvulaceae
Chenopodiaceae CALYSTEGIA SEPIUM (L.). R . B r . 2n=22, (24).
Subtropics and tropics. A perennial herb cultivated
KOCHIA SCOPARIA (L.) Schrader. Summer cypres. in China for its roots which a r e used as a vegetable.
2n=18. S. Europe and Asia. Cultivated in Japan and
China as apotherb, and as an ornamental. IPOMOEA AQUATICA Forsk. (syn. L. reptans
Poir. ). 2n=30. Throughout the tropics. Cultivated
SALSOLA KOMAROVII (Tljin. ) Oka-hijiki. 2n=36. in China and Hongkong in fish ponds to provide
Japan. Cultivated there (Kihara, 1969). spinach, pig and fish food (Purseglove, 1968). It is
also grown on flooded rice fields and on raised
SALSOLA SODA L. 2n=18, 36. Mediterranean beds.
region and Asia. A herb cultivated in Japan.
Corylaceae
SUAEDA GLAUCA Bunge. Matsuna. 2n= . Japan.
Cultivated there (Kihara, 1969). CORYLUS CHINENSIS Franchot. 2n= . China.
Cultivated especially in the Szechuan and Yunnan
Chloranthaceae provinces.
CHLORANTHUS SPICATUS (Thunb. ) Mak. (syn. CORYLUS HETEROPHYLLA Fischer. Siberian

Ch. inconspicuus Swartz. ). 2n=30. China. Culti- hazelnut. 2n=28. N. China, Japan, Korea and the
vated in China, Indochina and Japan as a tea aroma. Primorye Territory, USSR. Cultivated in China.
The seed has a medium-good taste. The shell is
Compositae very hard.
ARCTRJM LAPPA L. 2n=32, 36. Europe and Asia.
Cultivated in China and Japan as a root vegetable CORYLUS MANSHURICA Maxim. Manchurian hazel.
and in China and Europe (p. 130) as a medicinal 2n= . China, Japan and the Primorye Territory,
plant. USSR. Cultivated in China.
ARTEMISIA CAPILLARIS Thunb. 2n=18. E. Asia. CORYLUS SIEBOLDIANA Blum. Siebold's walnut.
Cultivated there and elsewhere as a medicinal 2n=28. Japan. Cultivated there.
plant.
Cruciferae
CHRYSANTHEMUM CORONARIUM L. Garland BRASSICA ALBOGLABRA Bailey. Chinese kale.
chrysanthemum. 2n= . China. Cultivated in S. 2n=18. China. Cultivation originated in S. China
China (Li, 1970), later in whole China and Japan (Li, 1970).
and elsewhere. Used as a vegetable.
BRASSICA CAMPESTRIS L. 2n=20, genome formu-
CHRYSANTHEMUM SEGETUM L. 2n=18. Europe la AA. See p. 131 for the origin of this species. In
and Asia. Cultivated especially in China. Leaves Centre 1 four (sub)species developed. Ssp. cnmen-
a r e used as a vegetable in the Near-East, Malaya se (L. ) Makino (syn. B. chinensis*) is an annual,
and Indochina. fast-growing, precocious, leafy vegetable. The
juicy leaves only contain 3.5-4% dry matter. Ssp.
CHRYSANTHEMUM SINENSE Sab. (syn. Pyrethrum nipposinica (Bailey) Olsson (syn. B. japonica Sieb.,
s i n e n s e D C ) . 2n= . China and Japan. Cultivated B. rapa var. laciniifolia (Bailey) Kitam). It has
there as a vegetable. finely dissected deep-green leaves (7% of fresh
leaves in dry matter). It grows slowly and has
GYNURA PINNATIFIDA DC. (syn. G. japonica little winterhardiness. Ssp. pekinensis (Lour.)
Mak. ). San ch'i, Tien ch'i. 2n= . China. A p e - Olsson (syn. B. pekinensis Rupr. ) is one of the
rennial herb cultivated for its medicinal p r o p e r - oldest vegetables in China. It forms large, compact
ties. heads. Ssp. narinosa (Bailey) Olsson, Broad-
beaked mustard forms a tight rosette of small,
LACTUCA DENTICULATA Maxim. 2n=10, (20). curley leaves (see B. narinosa).
It was cultivated in China.
BRASSICA CHINENSIS L. (syn. B. campestris L. cultivated in China and Japan and also elsewhere.
ssp. chinensis (L.) Makino). Chinese cabbage,
Celery cabbage, Pak-choi. 2n=20, genome formula Cucurbitaceae
AA. P r i m a r y centre China where it was domesti-
CUCUMIS MELO L. Melon, Muskmelon, Cante-
cated. Cultivated in SE. Asia and elsewhere. It is
loupe. 2n=24. Centre of origin in Africa (p. 110).
a vegetable, a salad and an oil crop (var. oleifera).
Secondary centre in China. Chinese and Japanese
Var. pekinensis (Rupr. ) (syn B. pekinense Rupr. ),
melons have small fruit and an unpleasant strong
p e - t s a i . (2n=20) has a blanched heart (see B. c a m -
taste. The genotypes are convar. chinensis (Pang.)
pestris*). Var. parachinensis Bailey (Sinsk. ) is
Greb., convar. monoclinus (Pang. ) Greb., ssp.
B. parachinensis Bailey, mok pak-choi. B. japoni-
conomon (Thunb. ) Greb. (syn. C. conomon Roxb. ),
ca* has also the same genome formula. This g e -
oriental pickling melon.
nome is related to the Ad genome (n=7) of B. ad-
p r e s s a Boiss. (2n=14), the F genome (n=8) of B.
fruticulosa Cyr. (2n=16) and the T genome (n=10) CUCUMIS SATIVUS L. Cucumber, Gherkin, 2n=14.
of B. tournefortii Gouan (2n=20) (Mizushima, 1969). Centre of origin in India (p. 64). Secondary gene
Prakash and Narain (1971) concluded that the g e - centre (a mesophytic type with very elongated
nomes of B. tournefortii a r e younger than the A fruits) arose there. Sources of resistance to m i l -
genome, and that this species has evolved from dew are found there.
the oleiferous plants of the species carrying the
A genome. HODGSONIA MACROCARPA Cogn. (syn. H. h e t e -
roclita Hook,f. &Thomson, Trichosanthes kadam
Miq.). Lard fruit. 2n= . Cultivated in Yunnan
BRASSICA NARINOSA L. (syn. B. campestris L. and elsewhere in China for its oily seeds.
ssp. narinosa L . ) . Kou T'sai, Broad-beaked m u s -
tard, Chinese Savoy. 2n=20. Only known as a cul-
TRICHOSANTHES CUCUMEROIDES Max. Japanese
tigen. Cultivated in E. China esp. around Shanghai.
snake gourd. 2n=44. The chromosomal number
Introduced to Japan and later to the USA (Helm,
suggests an auto- or alloploidization, which may
1963b). Related to B. chinensis* and other A genome
have happened in Japan or China where their roots
carrying diploid Brassica-species. It has entire,
a r e used te prepare starch.
deep dark-green leaves.
TRICHOSANTHES JAPONICA Regel. 2n=22. Japan.
EUTREMA WASABI Max. (syn. Wasabi japonica There starch is prepared from the roots.
Matsum., W. pungens Matsum., Lunaria japonica
Miq. ). Wasabi. 2n=28. Cultivated in Japan and E. Cyperaceae
Siberia (Kihara, 1969).
CAREX DISPALATA Boott. 2n=78, 84. Japan. Cul-
NASTURTIUM INDICUM DC. 2n= . In China it tivated there in rice fields for its leaves which a r e
was cultivated as a vegetable. Near Saigon var. made into hats.
apetala Gagnep. is grown as a medicinal crop.
CYPERUS CEPHALOTUS Vahl. (syn. C. natans
PUGIONUM CORNUTUM Gaertn. 2n= A herb Buch-Ham. ). 2n= . Trop. Asia and Australia.
cultivated as a vegetable in Mongolia. A perennial herb cultivated in the rice fields in
Japan for mat making (Uphof, 1968).
RAPHANUS SATIVUS-L. Radish, Small radish.
2n=18, genome formula RR. This is a very poly- CYPERUS GLOMERATATUS L. Wangul. 2n=
morphic species including biennials with large, Korea. Old fibre crop. Rarer than C. iwasakii*.
fleshy roots and annual forms. Japan and de oppo-
site coastal areas of the mainland a r e suggested CYPERUS IWASAKn M. Wangul. 2n= . Korea.
as primary centre. If so the radish would have Old fibre crop. Much more common than C. glome-
derived from the wild R. raphanistrum L. (2ftpl8) ratatus*.
and spread over the Old World probably intro-
gressing with other ecptypes and other wild species ELEOCHARIS DULCIS (Burm.f. ). Trinius (syn.
as R. maritimus Smith, (2n=18)andR. rostratus E. plantaginea R. Br. ). Water chestnut. 2n=
DC. W. Africa, upto India, China, Japan, Phillippines,
Wein (1964) suggested that R. maritimus is the Fiji and New-Caledonia. A herb. Cultivated in S.
parent of radish, while R. landra Moretti (2n=18) China for its tubers.
is the parent of the small radish. He indicated
the E. Mediterranean region as its gene centre ELEOCHARIS TUBEROSA Schultes. Water chest-
(p. 95). nut. 2n= . E. India, China and Japan. Cultivated
In Japan and China large rooted forms: daikon in the F a r - E a s t .
(R. acanthiformis de la Blanch., R. sativus var.
acanthiformis Mak., var. macropus Mak., var. Dioscoreaceae
longipinnatus Bailey) have been developed. A giant
DIOSCOREA JAPONICA Thunb. 2n=40. Japan. Cul-
form, the Sakurajuma Daikon (f. gigantissimus),
tivated in Japan, China and neighbouring islands.
is cultivated in Japan. The roots weigh up to 20 kg.
Some taxonomists include this species in D. oppo-
Vavilov (1949/50) called these giant cultivars, the
sita*.
champions of plant breeding.
Var. oleiformis* P e r s . , the oil-seed radish is
CRUCIFERAE - GRAMINEAE 31
DIOSCOREA OPPOSITA Thunb. (syn. D. batatas Euphorbiaceae

Decne. ). Chinese yam, Cinnamon vine. 2n=c. 140,
ALEURITES CORDATA (Thunb.). R . B r . Tung oil
c. 144. China. Cultivated in China, S. Japan, T a i -
t r e e . 2n=22. P r i m a r y gene centre: Japan. Culti-
wan and the Ryukyu islands.
vated there and in Formosa. Its crossability with
A. fordii* and A. montana* points to an affinity.
ALEURITES FORDII Hemsl. Tung oil t r e e . 2n=22.

C. China, between 26° and 33°N. Hybrids may
occur between wild and cultivated forms. Secon-
dary gene centre of cultivated tung t r e e s probably
in USA (p. 175). Cultivated in other American coun-
t r i e s , in USSR and Madagascar. With A. montana*
natural hybrids may occur. It also c r o s s e s with
A. cordata.
ALEURITES MONTANA (Lour. )Wils. Tung oil

t r e e . 2n=22. China south of 25°N. Cultivated in
Malawi, Brazil and elsewhere. With A. fordii*
natural hybrids may occur. It also c r o s s e s with
A. cordata*.
SAPIUM SEBIFERUM Roxb. Chinese tallow t r e e .

2n=36. Cultivated in the tropics.
Dioscorea opposita (Harris, 1972)
Euryalaceae
EURYALE FEROX Salisb. 2n=58. Tropical Asia.
Ebenaceae Cultivated in S. China.
DIOSPYROS KAKI L.f. (syn. D. chinensis Blume).
Kaki, Japanese persimmon. 2n=c. 54-56, 90. Fagaceae
Mountains of Central China. Centre of origin and
CASTANEA CRENATA Sieb. & Zucc. Japanese
primary centre of diversity in China. Secondary
chestnut. 2n=22, 24. Japan. Cultivated in Japan
centre is Japan. Cultivated in Mediterranean coun-
and in USA for its nuts.
tries and USA for its edible fruits.
CASTANEA MOLLISSIMA Blume. Chinese chestnut.
DIOSPYROS LOTUS L. Caucasian persimmon.
2n=24. N. and W. China. Cultivated in China and
2n=30. Subtropical China, in Talysk and W. Geor-
elsewhere for its nuts. It is source of resistance
gia, USSR and adjacent Iran (p. 72). Both a r e
to Endothia parasitica, a fungus causing damaga to
countries of primary diversity. Naturalized in the
chestnut (C. sativa) in USA.
Balkan peninsula and elsewhere. The fruit is ex-
cellent when dried.
QUERCUS ALPENA Blume. 2n= . Japan, Korea
and C. China. Cultivated as food for the Japanese
DIOSPYROS MAJOR (Forst, f.). Bakh. (syn. D.
oak spinner (Mansfeld, 1959).
andersonii P . S . Green). 2n= . Pacific islands.
Cultivated for its fruits which produce oil that can
QUERCUS DENTATA Thunb. Daimyo oak. 2n=24,
be used to scent other oils. The seeds are edible
48. Japan, Formosa, Korea and Manchuria and
(Smith, 1971).
W. and N. China. Cultivated as food for the Japa-
nese oak spinner and also used for timber (Mans-
Elaeagnaceae
feld, 1959).
ELAEAGNUS MULTIFLORA Thunb. Cherry
elaeagnus. 2n= . China, Japan and Korea. Cul- QUERCUS MONGOLICA Fisch. Mongolian oak.
tivated for its edible nuts (Mansfeld, 1959). 2n=24. N. China, Korea and N. Japan. Cultivated
as food for the Japanese oak spinner and also for
ELAEAGNUS PUNGENS Thunb. 2n=28. N. China timber.
and Japan. Cultivated for its edible fruits.
Ginkgoaceae
ELAEAGNUS UMBELLATA Thunb. 2n=28. China,
GINKGO BILOBA L. Gingko, Maidenhair t r e e .
Korea and Japan. Cultivated for its edible nuts
2n= . E. China. Cultivated in China and Japan
(Mansfeld, 1959).
as an ornamental. The seeds are eaten.
Eucommiaceae
Gramineae
EUCOMMIA ULMOIDES Oliver. Gutta percha
tree, Tuehung. 2n=34. The upland regions of W. ARUNDINARIA AMABILIS McClure. Tonkin bam-
and C. China. Cultivated as a medicinal plant. A boo, Tonkin cane. 2n= . Only known as cultigen
polygamous plant, dioecious forms have been found. and may have originated from Vietnam. Secondary
gene centre: China - Province Guandun and adjacent
regions of Guancy. Cultivated for its stems which that the genome formulas of E. utilis and E. c r u s -
have many technical properties. Used for hand galli* a r e the same and that the genome formulas
work, including fishing rods. of E. frumentacea and E. colona a r e also the
same.
AVENA SATIVA L. convar. nuda Nord. (syn. A.
nuda L . ) . Naked oats. 2n=42, genome formula ELYMUS ARENARIUS L. Sea lyme g r a s s , Sand
AACCDD. The origin of oats has been described elymus. 2n=56. Europe and Asia. A perennial
on p. 97). Cultigen of NE. China and Mongolia, the g r a s s . Cultivated in Japan for its culms and e l s e -
Tibetan-Himalaya highlands, in Turkestan and W. where as a dune stabilizer.
China. It is characterized by 5 to 7 florets per
flower and by big seeds. HORDEUM VULGARE L. ssp. humile Vav. &
Bacht. 2n=14. Barley. The origin of barley is d e s -
BAMBUSA GLAUCESCENS (Willd. ) Sieb, ex Munro. cribed on p. 80. Japan and C. China. Ssp. humile
(syn. B. nana Roxb. ). Hedge bamboo. 2n=72. is short, has small leaves, hexastichious e a r s
China and Japan where it is cultivated. In Indo- which a r e apically awned or awnless.
China it is grown as a bbrder plant.
LINGNANIA CHUNGII McClure. 2n= . S. China
BAMBUSA MULTIPLEX Raeusch. 2n=72. Cochin (Provinces Junjan and Guancy) in tropical e v e r -
China and Japan. A shrubby, woody g r a s s . Culti- green forests.
vated in trop. Asia for various purposes.
MISCANTHUS SINENSIS Anderss. 2n=38. Cultiva-
BAMBUSA STRICTUS Nees. 2n=70, 72. India ted as an 'arrow plant'. It may have played a role
(p. 64) and provinces Guancy, Guandun, China and in the origin of the Pansahi group of Saccharum
in Hongkong, in tropical evergreen forests. Stems sinense* (Grassl, 1968).
a r e about equal to those of the best Indian species
B. arundinaceae*. Secondary centres: Indo-China ORYZA SATIVA L. ecospecies japonica (syn. ssp.
(p. 47) and S. China. japonica Kato). Japonica r i c e . 2n=24, genome
formula AA. Indo-China. The origin of rice is d i s -
BAMBUSA TEXTILES McClure. 2n= . Province cussed on p. 65. Ecospecies japonica consists of
Guancy, China. ecotypes japonica and nuda. Spread to Japan,
Korea, N. China, Himalaya region, Egypt, Italy
BAMBUSA TULDOIDES Munro. 2n= . S. China. and Spain.
Cultivated for various purposes.
PANICUM MILIACEUM L. Proso millet, Shu.
CHIMONOBAMBUSA QUANDRANGULARIS (Fenzi.) 2n=36, (40, 54, 72). P r i m a r y centre: N. China.
Mak. 2n=48. Continental China and Taiwan. Culti- From here it has spread upto Italy. In China it
vated in Japan, China and Taiwan and occasionnally was an important cereal till the introduction of
on the shores of the Black Sea in Caucasus, USSR. wheat and barley (Li, 1970). P . spontaneum Lys-
sev. (2n= ) might be a weedy type of this s p e -
ECHINOCHLOA CRUS-GALLI L. Barnyard g r a s s . cies. It grows in Afghanistan, Kazakstan and may
2n=36, (42, 48), 54, (72). Japan and China. Close be wild in Mongolia (Mansfeld, 1959).
affinities with the cultivated E. frumentacea*. The
hexaploid type, 2n=6x=54 is an allopolyploid with PHYLLOSTACHYS BAMBUSOIDES Sieb. & Zucc.
E. oryzicola Vasing., 2n=36 as one parent. A c - Madake, Giant timber bamboo, Japanese timber
cording to Yabuno (1968) this species has the same bamboo. 2n=48. China where its p r i m a r y gene
genomic constitution as E. utilis (see E. frumen- centre is located. Secondary gene centre is Japan.
tacea*). Many forms occur there under the name 'Madake'.
ECHINOCHLOA CRUS-PAVONIS Schult. 2n=36, 54. PHYLLOSTACHYS DULCIS McClure, Sweetshoot
Subtropics and tropics. A grass cultivated in bamboo. 2n= . C. China. It is cultivated there.
Yunnan, China. The young shoots a r e edible.
ECHINOCHLOA FRUMENTACEA (Roxb.) Link, PHYLLOSTACHYS HENONIS Mitf. 2n=48, 54. C.

(syn. Panicum frumentacea Roxb. ). Japanese China (Szechwan). It is cultivated there. Secondary
millet, Billion dollar g r a s s , Sanwa millet. 2n=36, centre is in Japan. One of the forms developed
54, (56). China. P r i m a r y centre in China. Culti- under cultivation, is black bamboo, kenon bamboo
vated in Korea, China, USSR and N. America for ('Nigra', syn. Ph. nigra (Lodd. ) Munro, which is
human consumption, and as a fodder crop. Closely cultivated for its young shoots.
related to E. crus-galli*.
Ohwi and Yabuno separated E. utilis Ohwi &Yabuno PHYLLOSTACHYS MAKINOI Hayata. 2n=48. T a i -
(2n=54) from E. frumentacea because they found wan. It is cultivated in Japan.
differences in the genomic constitution, geographi-
cal distribution and panicle morphology of these PHYLLOSTACHYS MEYERI McClure. 2n=48.
two species (Yabuno, 1968). Yabuno considers China (Chzesian). Apparently cultivated in Japan
where a strain with deformed internodes a r o s e .
GRAMINEAE -JUGLANDACEAE 33
PHYLLOSTACHYS PUBESCENS Mazel ex de L e - Collins originated in E. Asia. Cultivated in China,

haie. 2n=48. Mountains of SE. China. Secondary Japan, Manchuria, Burma and the Philippines.
gene centre in Japan. This species has the largest
plants in the genus. Used in the timber industry ZIZANIA LATIFOLIA Turc. Manchuria waterrice.
and for its shoots. 2n=30, 34. China and adjacent regions. Cultivated
as a cereal in N. China in ancient times. Later on,
PHYLLOSTACHYS VIRIDIS (Young) McClure. its cultivation moved to the south. Its use as a
2n= . China. It is cultivated there for pulping. cereal gradually decreased. Now it is chiefly cul-
tivated in the south for its fleshy shoots which
SETARIA ITALICA L. (syn. Panicum italicum L.). yield the vegetable Chiaopai (Li, 1970).
Foxtail millet, Liang. 2n=18, genome formula AA.
N. China. From here it spread throughout Asia and Grossulariaceae
Europe in prehistoric times. In China it was an RIBES ALPESTRE Decne. 2n= . Himalaya up to
important cereal till the introduction of wheat and 3000 m. In China used as a hedge plant.
barley (Li, 1970).
It derives from S. virides (L.) Beauv. (2n=18), RIBES LONGERACEMOSUM French. 2n= . W.
genome formula AA. It is possible that S. pallidi- China. Cultivated there for its fruits. It could be
fusca Stapf. &C E . Hubbard (2n=36) is an allo- used to improve the strig length of cultivar R.
tetraploid with S. italica as one of the diploid p a - nigrum*.
rents.
RIBES USSURIENSE Jancz. 2n= . E. Asia. It is
SINOBAMBUSA TOOTSIK (Sieb. ) Makino. 2n=48. a source of resistance to black-current gall mite,
Japan (Riu-Kiu Islands). Phytoptus ribes Nal., a pest of R. nigrum*. It
easily hybridizes with R. nigrum.
SINOCALAMUS BEECHEYANUS (Munro) McClure.
2n= . S. China. It is cultivated there. Illiciaceae
SINOCALAMUS EDULIS (Odashima) Kenf. 2n= ILLICIUM ANISATUM L. Japanese s t a r anise.
China. Its shoots a r e edible. 2n=28. China, Korea and Japan. Cultivated for its
medicinal seeds.
SINOCALAMUS OLDHAMII (Munro) McClure.
2n= . Taiwan. ILLICIUM VERUM Hook.f. (syn. L. religiosum
Sieb. &Zucc. ). Star anise. 2n=28. SE. Asia.
SORGHUM BICOLOR (L.) Moench. Chinese Amber Cultivated for its medicinal fruits. It is not known
Canes and Kaoliang. 2n=20. Sorghum was domesti- wild.
cated in Africa (p. 116). Chinese amber canes a r e
found on the coast of China and Korea, and also in Iridaceae
India and Burma. They very likely arrived in China BELAMCANDA CHINENSIS (L.) DC. Blackberry
and the F a r East by sea traffic. They a r e related lily, Leopard flower. 2n=32. China and Japan.
with the E. African sorgo. After introduction of Cultivated there as a medicinal crop.
sorghum from India into S. China it came into con-
tact with S. propinquum (Kunth. ) Hitchc., 2n= , IRIS ENSATA Thunb. 2n=40. Temp. Asia upto
and by hybridization kaoliang arose (Doggett, 1970). Himalaya. Cultivated in China for its leaves (bin-
ding material).
TRITICUM AESTIVUM L. Thell. s s p . vulgare
(Vill.)MK. (syn. T. vulgare Vill. ). Bread wheat, Juglandaceae
Common wheat. 2n=42, genome formula AABBDD.
Centre of origin: Transcaucasia and adjacent r e - CARYA CATHAYENSIS Sarg. Chinese hickory.
gions (p. 82). It arrived through Korea and Japan 2n= . E. China. Cultivated in Yunnan, China
in about 300 B.C. (Kihara, 1969). Secondary cen- (Mansfeld, 1959). It closely resembles C. tonki-
t r e of diversity in both countries. The Chinese nensis Lecomte.
wheats a r e characterized by very broad leaves,
with 5 to 7 florets p e r spikelet, with squarehead JUGLANS AILANTIFOLIA Carr. 2n= . Japan.
e a r s , with daylength neutrality, with fast ripening Var. ailantifolia (syn. J. sieboldiana Maxim.),
grain, and by precocious forms. In the mountains Siebolds walnut. Var. cordiformis (Maxim. ) Rehd.
of the Sinkiang Province of China very frost r e - (syn. J. cordiformis Maxim. ). Cultivated in N.
sistant wheats developed. Chinese and Japanese America.
wheats cross easily with rye, probably because
there was no selection p r e s s u r e against this cha- JUGLANS DUCLOUXIANA Dode. 2n= Mountain
racteristic due to absence of Secale cereale-types. regions of Asia. Cultivated in China.
Some Japanese and Korean wheats are short and
this character was introduced in wheat varieties JUGLANS MANDSHURICA Maxim. Manchurian
of Italy, Japan, USA, Mexico and elsewhere. walnut. 2n=32. NE. China andthe Primorye T e r r i -
tory, USSR. It is winterhardy and is used as root-
ZEA MAYS L. Maize. 2n=20. Maize was domesti- stock.
cated in C. America (p. 166). Secondary centre:
China (Brandolini, 1970). The mutant ceratina
Labiatae ria and adjacent USSR. Domestication may have

taken place in E. half of N. China about the 11th
ELSHOLTZIA CRISTATA Wildd. 2n . China and
century BC. (Hermann, 1962; Hymowitz, 1970).
Japan. A perennial plant introduced in other parts Semi-cultivated and weedy types described as
of Asia, Europe and America as an oil-seed crop. G. gracilis Skvortzow (2n=40), might be hybrids
of G. max and G. ussuriensis.
MENTHA ARVENSIS L. var. piperascens Mai.
Japanese mint. 2n=96, genome formula R a R a SSJJAA.
LESPEDEZA CUNEATA (Dum. Cours. ) G. Don.
Japan. Cultivated in Japan, China and Brazil. It is
(syn. L. sericea Benth. ). Perennial lespedeza.
the main source of menthol. Var. agrestis (2n=72,
2n=(18), 20. E. Asia. Cultivated in the USA for
genome formula R a R a SSJJ) is very likely a hybrid
erosion control.
of var. piperascens and M. japonica Makino
(2n=48). It is a source of early maturity and rust
resistance (Ikeda et a l . , 1970). M. arvensis is one LESPEDEZA STIPULACEA Maxim. Korean l e s -
of the parents of M. x gentilis (see M. cardiaca*). pedeza. 2n=20. E. Asia. Cultivated in the USA
for hay making.
PERILLA ARGUTA Benth. 2n= . China and Japan.
LESPEDEZA STRIATA Hook. Common lespedeza,
Cultivated for various purposes. It is sometimes
King g r a s s . 2n=22. E. Asia. Cultivated in the
included in P. frutescens*.
USA in pastures and for hay making.
PERILLA FRUTESCENS Britt. (syn. P. ocymoides
MUCUNA HASSJOO (Piper &Tracy) Mansf. (syn.
L. ). Suttsu, Perilla. 2n=38, 40. Himalayas, China
Stizolobium hassjoo Piper &Tracy). Yokohama
and Japan. P r i m a r y centre: China. The red-leaved
bean. 2n= . Japan and China. Cultivated there
strains (syn. P. crispa (Thunb. ) Nakai) are some-
and in the USA for the seeds.
times used as ornamentals and the green-leaved
plants (P. crispa var. ocymoides) for the seed oil.
Cultivated in China, Japan and Korea as a drug PHASEOLUS ANGULARIS Wight. Adzuki bean.
2n=22. Primary gene centre Central China. It is
plant (Li, 1969) and formerly as a leafy vegetable.
unknown wild. Cultivated in China, Manchuria,
Korea and Japan. Secondary gene centre Japan.
STACHYS SIEBOLDII Miq. Chinese artichoke. J a -
panese artichoke. 2n= . It is one of the few tuber
PHASEOLUS VULGARIS L. var. chinensis (syn.
crops domesticated in China. Cultivated there,
Ph. chinensis Hort, ex Schur. ). Asparagus bean.
Japan, Belgium and France.
2n=22. Its origin is discussed on p. 168. It reached
China from the Americas after Columbus' voyage.
Lauraceae
In China a secondary gene centre a r o s e . The main
CINNAMOMUM CAMPHORA (L. ) Nees & Eberm. character, a parchment-like layer in the pod wall
Camphor t r e e . 2n=24. China, Japan and Taiwan. was lost and the pod became edible.
Cultivated in these countries and other tropical
countries. PUERARIA THUNBERGIANA (Sieb. & Zucc.)
Benth. Kudzu. 2n=24. China and Japan. Cultivated
CINNAMOMUM ZEYLANICUM Breyn. 2n=24. as a cover crop, green manure and hay crop, in
Ceylon and SW. India. Cultivation started in Ceylon New Guinea and New-Caledonia as a tuber crop
in 1770. Cultivated now in several countries. (p. 51).
Leguminosae VICIA UNLJUGA A. Br. (syn. Orobus lathyroides

ASTRAGALUS SINICUS L. (syn. A. lotoides Lam.). L. ). Two-leaved vetch. 2n=12, (24, 36). E. Siberia,
Genge, 2n=16. China. Cultivated there on rice Manchuria and Japan. Occasionally cultivated
soils as a soil improver.
WISTERIA BRACHYBOTRYS Sieb. &Zucc. 2n=16.
A vine. Cultivated for its fibrous bark.
CANAVALIA GLADIATA (Jacq. ) DC. var. alba
(Makino) Hisauchi (syn. C. ensiformis (L.)
DC. var. alba Makino). Siro-nata-name. 2n=22. Liliaceae
Cultivated in Japan (Sauer, 1964). Characterized ANEMARRHENA ASPHODELOIDES Bunge. 2n=22.
by white seeds. N. China. A medicinal plant occasionally cultiva-
ted.
GLEDITSIA JAPONICA Miq. 2n= . Japan. Culti-
vated for fruit juice, which is used for washing. FRITILLARIA VERTICILLATA Willd. 2n=2<±. Var.
thunbergii Baker. Cultivated in China as a medici-
GLYCINE MAX (L. ) Merr. (syn. Soja hispida nal plant.
Moench, S. max (L.) Piper, Glycine hispida
(Moench) Maxim., G. gracilis Skvortzow). Soya LILIUM AURATUM Lindl. Gold band Lily, Yama-
bean. 2n=(38), 40. It is unknown wild. It is b e - yuri. 2n=24. Japan. Cultivated there for its large
lieved to be a cultigen from G. ussuriensis Regel bulbs (Kihara, 1969).
&Maack. (syn. G, javanica Thunb., G. soja Sieb.
& Zucc.) (2n=40). This latter species is found in LILIUM CORDIFOLIUM Thunb. 2n=24. Japan.
N. and C. China, Taiwan, Japan, Korea, Manchu- Cultivated there for its starchy bulbs.
LABIATAE - PHYTOLACCACEAE 35
LILIUM LANCIFOLIUM Thunb. Oni-yuri. 2n=24. Moraceae

Japan. Cultivated there for its bulbs (Kihara, 1969).
BROUSSONETIA KAZINOKI Sieb. 2n=26, 39. A
t r e e cultivated in Japan and Korea for its bark
LILIUM MAXIMOWICZII Regel. Ko-oni-yuri. which is a source for paper production.
2n=24. Japan. Cultivated there as a food crop
(Kihara, 1969).
BROUSSONETIA PAPYRIFERA (L) Vent. Paper
mulberry. 2n=26. China and Japan. In the Far
LILIUM TIGRINUM Ker-Gawl. Tiger lily. 2n=(24),
East this t r e e is used for making paper and bark-
36. China. Cultivated there and in Japan for its
cloth (Purseglove, 1968).
edible bulbs.
OPHIOPOGON SPICATUS Kunth. 2n= . China. MORUS ALBA L. White mulberry. 2n=28. China.
Cultivated there and elsewhere for its leaves
A herb cultivated in Chekiang as a medicinal plant.
eaten by silk worms, for its fruits and for paper
making. It is often planted as a road-side t r e e ,
Magnoliaceae cv Makado has 2n=3x=42.
MICHELIA FIGO (Lour. )Spr. (syn. M. fuscata
Andr. ). Banana shrub. 2n=38. China. Cultivated Musaceae
there for its banana-scented flowers used for
scenting hair oil. MUSA BASJOO Sieb. & Zucc. 2n=22. Japan.
Species of the Eumusa section. Used for making
fibre.
Malvaceae
ABELMOSCHUS MANIHOT* Myricaceae
MYRICA RUBRA Sieb. & Zucc. (syn. M. nagi
ABUTILON AVICENNAE Gaertn. (syn. A. theo-
Thunb.). Chinese strawberry tree, Ioobai, Yama
phrasti Medic. ). Button weed, Chinese jute, Velvet
momo. 2n=16. Cultivated in China for its fruits.
weed, Butter print chingma. 2n=42. Cultivated in
China (many local varieties), USSR and elsewhere
for its fibre called jute or Indian mallow. Oleaceae
FRAXINUS CHINENSIS Roxb. 2n=92, 138. W. and
GOSSYPIUM ARBOREUM L. Tree cotton. 2n=26, C. China. Especially var. acuminata Lingelsh.
genome formula A..A... Arose in India (p. 68). (syn. F. koehneana Lingelsh. ) is cultivated as a
Race sinense, Chinese cotton, Nanking cotton, host plant of the insect Coccus pela for wax p r o -
developed in E. China. It is the earliest fruiting duction.
form of this species. It has short lint and requires
a long daylength. First cultivated as an ornamental. LIGUSTRUM JAPONICUM Thunb. Japanese privet.
At present is has a low breeding value. 2n=44. A shrub. Cultivated in Japan for its seeds.
HIBISCUS SYRUCUS L. Rose of Sharon. 2n=80, LIGUSTRUM LUCIDUM Ait. 2n=46. A t r e e culti-
80-84, 90, 92. China and Formosa. Cultivated vated in China as a host plant of the insect Coccus
first in China as a hedge plant and later elsewhere pela for wax production.
as an ornamental.
LIGUSTRUM OVAFOLIUM Hassk. 2n=46. Japan.
MALVA SYLVESTRIS L. High mallow. 2n=42. P r o - A shrub widely planted for hedges in Europe and
bably the early vegetable K'uei mentioned in Chi- elsewhere. It may have run wild there.
nese literature. At present a weed in China (Li,
1970). Cultivated in Europe as a medicinal crop OSMANTHUS FRAGRANS Lour. 2n=46. Himalaya,
and ornamental. China and Japan. A t r e e cultivated in E. Asia for
its very scented flowers used to aromatize tea.
MALVA VERTICILLATA L. (syn. M. crispa L.,
M. mohileviensis Graebn., M. pamiroalaica Ilj. ). Palmae
Mallow. 2n=c. 84, c. 112. E. Asia. It was an
early Chinese domesticate there. About 500 A.D. TRACHYCARPUS FORTUNEI (Hook. ) H. Wendl.
Windmill palm, Chusan palm. 2n=36. China.
it was there an important vegetable with several
Often planted in E. Asia for its fibres.
varieties like purple and white stemmed, large
and small leaves. During the 7-10th century the
cultivation in China declined. In 1848 it was only Pedaliaceae
observed in remote a r e a s . Introduced to Japan, SESAMUM INDICUM L. Sesame, Beni seed. 2n=26.
where it is a weed now (Li, 1969). Also in W. Asia P r i m a r y centre is discussed on p. 126. Secondary
and Europe. Cultivated in Europe as a medicinal centre in China/Japan. There ssp. quadricarpel-
crop. This plant has often been described as M. latum developed.
crispa being a cultigen of M. verticillata.
Phytolaccaceae
Menispermaceae
PHYTOLACCA ACINOSA Roxb. 2n=36. Trop.
COCCULUS THUNBERGII DC. 2n= . A woody Asia, China and Japan. A perennial herb cultiva-
vine cultivated in Japan for basket making. ted in India as a vegetable. In the Chinese p h a r m a -
cy the b e r r i e s a r e used.
Plantaginaceae Indian jujube. 2n=48. Tropical Africa and Asia.

PLANTAGO MAJOR L. 2n= . Europe and temp. Cultivated in India, Cochinchina and China since
Asia. Var. asiaticum Dene, is cultivated in China ancient times.
as a vegetable and as a medicinal herb.
Rosaceae
Polygonaceae AMYGDALUS BESSERIANA Schott, (syn. A. nana
FAGOPYRUM TATARICUM (L.) Gaertn. Tatary L. ). Dwarf almond, Russian almond, Steppe a l -
buckwheat. 2n=16. P r i m a r y centre is probably in mond. 2n=16. P r i m a r y gene centre is in E.Europe
East Asia. There, it is often found as a weed of and Siberia (p. 139). Ssp. rosiflora types from
barley and wheat fields. It is a source of rutine. China a r e sources of winterhardiness, precocious
fruiting and prostrate-growing. Disadvantages a r e
POLYGONUM HYDROPIPER L. Tade, Knotweed. its small and white-fleshed fruits.
2n=20, (22). Cultivated in Japan (Kihara, 1969)
and formerly, also in China. Var. maximowiczii AMYGDALUS KANSUENSIS Skeels. (syn. Persica
is the edible type. In China and elsewhere it is a kansuensis (Rehd. ) Kov. &Kost., Prunus kansuen-
weed now (Li, 1969). s i s R e h d . ). Chinese bush peach. 2n= . NW.
China. It tolerates -35°C. It could be useful as a
POLYGONUM MAXIMOWICZII Regel. 2n= rootstock (Zylka, 1970).
Japan. Cultivated there as a vegetable.
AMYGDALUS MIRA (K.) Koch (syn. Prunus mira
POLYGONUM TINCTORIUM Ait. 2n=40. China. Koehne). Smoothpit peach. 2n= . W. China. It
Formerly it was cultivated as a blue dye-plant. might be used as a source of late flowering (Zylka,
It is naturalized for instance in the Ukraine. 1970) and winterhardiness.
RHEUM HYBRUJUM Murray. Rhubarb. 2n= AMYGDALUS PERSICA L. (syn. A. pumila Lour.,
Originating probably in Mongolia. Probably a P e r s i c a vulgaris Mill., Prunus persica (L.)
hybrid of R. rhaponticum and R. palmatum*. Batsch. ). Peach. 2n=16. P r i m a r y centre: montane
a r e a s of Tibet and SW. China (Holub, 1969). Secon-
RHEUM OFFICINALE Baill. Medicinal rhubarb. dary centres: Iran, C. Asia (p. 74), Caucasus,
2n=22, (44). Cultivated in China as a medicinal Crimea (p. 89), Moldavia (USSR) (p. 139), Italy,
crop. Spain (p. 105) and California, USA (p. 177). Culti-
vated for its fruits and as an ornamental. Holub
RHEUM PALMATUM L. East Indian rhubarb, (1969) divided the cultivated varieties according
China rhubarb, Turkey rhubarb. 2n=22, (44). to their morphology and geography into four groups
Mongolia, or W. China. Cultivated formerly as a viz. 1. Chinese, 2. Central Chinese, 3. Western
purgative (root) and at present as an ornamental. Chinese and 4. Yellow-fleshed from Europe.
It is probably one of the parents of R. hybridum*
(syn. R. rhabarbarum L. ). It is related to R. ARMENIACA MANDSHURICA (Koehne) Kost. (syn.
rhaponticum* (syn. R. rhabarbarum L. ). Prunus mandshurica (Maxim.), P. armeniaca var.
mandshurica Maxim. ). Manchurian apricot. 2n=16.
RHEUM UNDULATUM L. 2n=22, 44. China. Cul- S. Ussuria, E. Manchuria upto N. Korea. It resists
tivated as a vegetable. intense cold.
Ranunculaceae ARMENIACA MUME (Sieb. &Zucc. ) Sieb, ex

Carr. (syn. Prunus mume Sieb. & Zucc. ). J a p a -
ACONITUM CARMICHAELI Debeaux (syn. A. wil- nese apricot. 2n=16, 24. Mountains in Central and
sonii Stapf ex Mottet). 2n=64. China. Cultivated N. China. Cultivated as an ornamental in China,
there as a medicinal crop. Korea and Japan. Its kernel is eaten. It is not
frost resistant.
COPTIS CHINENSIS Franch. 2n= . W. China.
A herb cultivated as a medicinal plant. ARMENIACA VULGARIS Lam. (syn. Prunus a r m e -
niaca L., P. tiliaefolia Salisb, ). Apricot. 2n=16.
Rhamnaceae P r i m a r y gene centre: NE. China. Secondary gene
HOVENIA DULCIS Thunb. Japanese raisin t r e e . centre in E. Tian-Shan and in Trans-Tlii and
2n=24. China, Korea and Japan. Cultivated in E. Dzhungar-Alatau. A small area of wild apricots
Asia upto India for its edible inflorescence and as is found in Daghestan (p. 89) and in the Kotur
an ornamental. Bulak canyon in the Alma-Ata region. Evreinoff
(1955) believed that it occurred in a much larger
ZIZIPHUS JUJUBA Mill, non Lam. (syn. Z. vul- region up to Armenia and Iran. The related A.
garis Lam. ). Jujub, Chinese jujube. 2n=24, (40), ansu (Max. ) Kost. (syn. Prunus ansu Komar)
48, (60, 72), 96. P r i m a r y gene centre of the wild (2n= ) could be used as a source of resistance
and cultivated types is probably in Central and S. to fungal diseases.
China. Wild type grows in many Asian countries.
CHAENOMELES SINENSIS (Thouin) Koehne. Chi-
ZIZIPHUS MAURITIANA Lam. (syn. Z. jujuba nese quince, Japanese quince. 2n=32. Its fruits
(L.) Lam. non Mill. ), Z. sosoria Roem. &Schult. ).
PLANTAGINACEAE - ROSACEAE 37
MALUS HALLIANA Koehne. 2n=34. Gene centre in

China and Manchuria. A wild ornamental.
MALUS HUPEHENSIS (Pamp.)Rehd. (syn. Pyrus

hupehensls Rehd. ). Chinese crab apple. 2n=51, 68.
C..and W. China and south into Assam. It is used
as rootstock.
MALUS MICROMALUS Makino. 2n=34. China and

Japan. Cultivated there. Its fruits a r e edible
(Rehder, 1947). It is thought to be a hybrid of M.
spectabilis x probably M. baccata.
MALUS PUMILA var. niedzwetskyana Dieck. 2n=34.

Tian-Shan mountains, China.
MALUS SIEBOLDII Rehd. (syn. Pyrus toringo

Sieb. ). Toringa crab. 2n=34. P r i m a r y centre
Japan. It is a profusely branching salt-tolerant
shrub used as rootstock for dwarf plantings.
MALUS SPECTABILIS (Ait.) Borkh. Zamechatel-

naya. 2n=34, 51. P r i m a r y centre: Central China.
It is not known wild. It is probably a parent of
M. micromalus*.
PRUNUS subgen. cerasus P e r s . There a r e proba-

bly 150 species belonging to ssp. c e r a s u s . The
greatest number is located in the mountain regions
of W. China. Some big fruited species a r e valu-
able for breeding.
PRUNUS CANTABRIGIENSIS Stapf, (syn. P .

pseudocerasus Koids.). Chinese sour cherry,
Yingtao cherry. 2n= . Yangtze Valley in China.
Cultivated in China (Uphof, 1968).
Prunus bessyi
PRUNUS DAVIDIANA (Carr. ) Franch. (syn. P.
have the aroma of quince. Therefore it is m i s - persica var. davidiana Maxim., P e r s i c a davidia-
takenly described as a species of C. donia. na Carr. ). Chinese wild peach. 2n=16. Vladivos-
tok, SW. through Charbln upto Dacin-San and
CRATAEGUS HUPEHENSIS Wils. 2n= Hupeh, Ala-San (China). Cultivated as an ornamental. It
China. Cultivated there for its fruits. is frost, drought and heat resistant. Probably it
is valuable as a rootstock for Amygdalus persica*
CRATAEGUS PENTAGYNA Waldst. &Kit. (syn. and Prunus domestica* (Zylka, 1970).
C. pinnatifida Bunge). Chinese hawthorn. 2n=34.
N. China, Korea and Siberia. Cultivated in China PRUNUS PSEUDOCERASUS Lindl. (syn. P. pani-
for its fruits (Li, 1970). culata Edwards). 2n=32. W. Hupei, China. A t r e e
cultivated there for its fruits.
DUCHESNEA FILIPENDULA (Hemsl. ) Focke.
(Syn. Fragaria filipendula Hemsl, ). 2n= . A PRUNUS SALICINA Lindl. Chinese plum, Japanese
herb of China cultivated for its fruits. plum. 2n=16. P r i m a r y gene centre: the forests of
N. China. Second gene centre: Japan, Cultivated
ERIOBOTRYA JAPONICA (Thunb. ) Lindl. Loquat, in Japan, China and also in California. It crosses
Japanese plum, Japanese medlar. 2n=32, 34. Cul- easily with the North American plum species.
tivated in China, Japan, California and S. Europe. A new stone fruit 'cherry plum' was derived from
crossing the wild P . cerasifera* with P. salicina
MALUS ASIATICA Nakai (syn. M. prunifolia var. var. Burbank. Because of its winterhardiness
rinki). 2n= . NW. China. Used for cultivation. this fruit t r e e can be grown where apricot will not
fruit.
MALUS BACCATA (L.) Borkh. var. mandchurica
(Max.). Schneid, (syn. Pyrus baccata L. ). Man- PRUNUS SARGENTII Rehd. Sargent cherry, Moun-
churian crab apple. 2n=34. P r i m a r y gene centre: tain cherry. 2n=16. Japan, Manchuria, Korea and
the Shansi, Shensi, Kiangsi and other provinces in rarely in the Far East of USSR. Used as an orna-
China. Centre of origin probably Siberia, SE. of mental. It is frost resistant and fast growing. The
Lake Baikal (p. 140). Used for rootstock material. fruits a r e not very palatible.
PYRUS BRETSCHNEIDERI Rehd. 2n=34. Hupei

and Shansi, China. P r i m a r y gene centre: N. China.
There it was domesticated. It is the commonest
cultivated pear in this region. The fruits a r e cha-
racterized by hard, crisp, white sweet flesh.
PYRUS CALLERYANA Dene. 2n=34. China, Japan

and Korea. P r i m a r y gene centre: the Tsinling
mountain range, China. It is used as rootstock.
PYRUS PHAEOCARPA Rehd. 2n=34. N. China.
PYRUS PYRIFOLIA (Burm.) Nakai (syn. P . s e r o -

tina Rehd. ). Sand pear. 2n=34. P r i m a r y gene cen-
t r e : the highlands of N. and C. China. Var. culta
(Mak. ) Nakai is drought resistant, but not very
winterhardy. The leaves reach 15 cm in length.
The fruits a r e outstanding for their preserving
quality. It c r o s s e s easily with the wild European
pear, P. communis*.
PYRUS USSURIENSIS Maxim. Ussuri pear. 2n=34.

SE. Siberia upto the Manchurian-Chinese area,
P r i m a r y gene centre: NE. China and in the P r i m o -
rye Territory, USSR, along the Ussuri r i v e r . The
ancient var. culta is widely distributed over N. and
C. China. It is adapted to cold, dry regions. It is
the most winterhardy wild pear. As this species
originated outside the Chinese centre proper it has
no resistance to scab and other diseases. It p r o -
bably played a part in the origin of P . communis*.
ROSA MULTIFLORA Thunb. 2n=14. E. Asia. Used

as rootstock.
Prunus davidiana ROSA RUGOSA Thunb. 2n= . China and Japan.

The rose hips a r e used by the Ainu. Used as an
ornamental and in hedges as one parent to breed
PRUNUS SIMONII Carr. Apricot plum. Simon plum. for rootstocks.
2n=16. P r i m a r y centre: probably N. China and
Japan. No wild plants a r e found. Also cultivated RUBUS ILLECEBROSUS Focke. Strawberry r a s p -
there. Crossing with P . triflora Roxb.* (2n=16) berry, Balloon berry. 2n=14. Japan. Cultivated
from the same a r e a has led to the development of in N. America.
cultivars which are especially grown in N. A m e r i -
ca. RUBUS PHOENICOLASIUS Maxim. Wine raspberry.
2n=14. Japan and N. China. Cultivated for its
PRUNUS TOMENTOSA Thunb. (syn. P. trichocarpa fruits and as an ornamental.
Bunge). Nanking cherry, Manchur cherry, Chinese
bush cherry. 2n=16. N. and W. China, Japan, RUBUS PUNGENS Oldhami. 2n= . Korea. Used
Himalaya, Turkestan and in the Far East of USSR. in breeding with R. idaeus*.
Cultivated as a fruit t r e e and ornamental in the
F a r East of USSR, N. China and Japan. Rubiaceae
GARDENIA JASMINOIDES Ellis (syn. G. florida
PRUNUS USSURIENSIS Kov. &Kost. (syn. P. t r i - L. ). Cape jasmin. 2n=22. Probably S. China. Cul-
flora Roxb. var. mandshurica Skvoro. ). Ussurian tivated in E. Asia for perfumery oil.
plum. 2n=16. Cultivated or run wild in Manchuria,
E. of USSR and for some years also in Siberia and
Rutaceae
N. Kazakhstan (Zylka, 1970). It is a source of
good fruit flavour and cold resistance. This s p e - CITRUS ICHANGENSIS Swing. 2n=18. S. of Tsin
cies is sometimes considered as a subspecies of 'lin range in W., C. and SW. China. It is very
P. cerasifera*. frost resistant. Therefore it has been crossed
with cultivated Citrus species. Used as a root-
PYRUS BETULAEFOLIA Bgb. 2n=34. N. and C. stock.
China. Used as rootstock. Resistance against
scab (Venturia) is found in this species. CITRUS JUNOS Tan. Juzu. 2n=18. Han'su province,
China at 1372 m altitude. It is frost resistant and
therefore used as a rootstock. Its fruits a r e big
ROSACEAE - THEACEAE 39
but sour. It was already known in the time of Con- Sapindaceae

fucius (about 2500 years ago). Because of intro-
NEPHELIUM LITCHI Camb. (syn. Litchi chinen-
gression, characters of 12 Japanese and 2 Chinese
sis Sonn. ). Litchi. 2n=28, 30. S. China. Cultiva-
wild Citrus species a r e recognized in juzu.
ted there and elsewhere for its 'litchi nuts'.
CITRUS RETICULATA Blanco. (C. nobilis Andr.
NEPHELIUM LONGANA (Lam.) Camb. (syn.
non Lour. ). Mandarin, Tangerine. 2n=18. See for
Euphoria longana Lam. ). Longan. 2n=30. C. and
its possible origin p. 55. Secondary centre: Japan.
S. China. Cultivated there for its fruits.
New types such as the Satsuma (var. unshiu, syn.
'C. unshiu') and the Natsudaudau ('C. natsudaidai')
arose through bud mutation and spontaneous hybrid SAPINDUS MUKOROSSO Gaertn. Chinese soap-
dization. Satsuma tangerine (unshiu mikan) is p r o - berry. Soapnut t r e e . 2n= . China, Burma and
bably a derivative of So-kitsu or Man-kitsu. Himalaya. Cultivated in China, Japan, India and
elsewhere for its fruits.
CLAUSENA LANSIUM (Lour.) Skeels. Wampi.
Scrophulariaceae
2n-18. It is a small fruit-tree of S. China. Culti-
vated in several tropical countries. VERONICA ANAGALLIS L. 2n=36. Temperate
zone. Cultivated in Japan as a lettuce.
FORTUNELLA CRASSIFOLIA Swing. Meiwa kum-
quat. 2n=18. China and Japan. It is occasionnally Simaroubaceae
cultivated.
AILANTHUS VILMORINIANA Dode. 2n= . W.
China. Cultivated there as a source of food for
FORTUNELLA HINDSII (Champ. ) Swing. Wild
kumquat, Kongkong kumquat. 2n=18, (36). The silk worms.
Tziulun mountains of Honkong. It has no great
value. The 4x form originated spontaneously Solanaceae
(Cameron & Soost, 1969). SOLANUM MELONGENA L. var. esculentum Nees.
Chinese eggplant. 2n=24. Domesticated in India
FORTUNELLA JAPONICA (Thunb. ). Swing, (syn. (p. 55). Secondary centre: China. There the small
Citrus japonica (Thunb. ). Round kumquat, Marumi fruited-form already existed before 500 AD (LI,
kumquat. 2n=18. P r i m a r y centre Japan. This 1969).
fruit tree is unknown in a wild state. It is occasio-
nally cultivated. Taxaceae
TORREYA GRANDIS Fort. Chinese torreya.
FORTUNELLA MARGARITA (Lour. ) Swing, (syn.
2n= . China. Cultivated in Szechuan and Anhwei,
Citrus margarita Lour. ). Oval kumquat, Nagami
China.
kumquat. 2n=18. Japan. Cultivated in Japan,
China and Florida, USA. Fortunella species cross
TORREYA NUCIFERA Sieb. &Zucc. Japanese
easily with each other and with Citrus species.
torreya, Kaya. 2n= . Japan. Cultivated there.
PONCIRUS TRIFOLIATA (L. ) Raf. Trifoliata
Tetragoniaceae
orange. 2n=18, (36). N. China, also primary
centre. This area is its centre of diversity. Cul- TETRAGONIA EXPANSA Murr. New Zealand s p i -
tivated in China as an ornamental, and in USSR nach, Tsuruna. 2n=32. Japan. Cultivated there
and Japan it is used as a rootstock. Hybrids with (Kihara, 1969). However, Uphof (1968) restricted
sweet orange (Citrus sinensis*) a r e citranges its native area to Australia and New Zealand.
used as rootstocks, with sour orange (C. auran-
tium*) are citradias, crosses of citrange with Theaceae
kumquat (Fortunella margarita*) resulted in c i -
trangequat. CAMELLIA JAPONICA L. Camellia. 2n=30. From
Taiwan northwards through the Riu Kiu islands and
Yakushima to S. Japan, also in S. Korea and Dage-
TRIPHASIA TRIFOLIA (Burm.f. ) P. Wilson (syn.
let islands (Sealy, 1958). Cultivated for its Tsubaki
T. aurantiola Lour., T. trifoliata DC.). Lime
Oil and as an ornamental. Attempts to cross it with
berry, Trifoliata lime berry. 2n= . Centre of
tea, C. sinensis* have failed so far, although C.
origin possibly China (Mansfeld, 1959). Cultivated
wabiske* might be a natural hybrid of C. sinensis
for its edible fruits.
and C. japonica. If so this species might be used
ZANTHOXYLUM PIPERITUM DC. Japanese p r i c k - to introduce japonica characters such as cold r e -
ly ash, Japan pepper, Sanshô. 2n=70. China and sistance into tea.
Japan. Cultivated there (Uphof, 1968; Kihara,
1969). CAMELLIA OLEIFERA Abel. (syn. C. sasanqua
Thunb. ). Sasanqua camellia. 2n=90. China. Cul-
ZANTHOXYLUM SIMULANS Hance (syn. Z. niti- tivated there and Indo-China to yield 'tea oil'
d u m D C , Z. bungei Planch. ). 2n=32. China. A (Sealy, 1958).
shrub cultivated in C. and S. China for its seeds.
This seed is a source of Chinese pepper. CAMELLIA SINENSIS (L. ). O. Kuntze (syn. C.
thea Link., Thea sinensis L. ).. Tea. 2n=30, (45,
60). P r i m a r y centre: the mountains of China, Thymelaeaceae

north of NE. India (Simura et a l . , 1967). It spread
to SE. China, Indo-China, Assam and later to DAPHNE ODORA Thunb. 2n=(18, 27), 28, 30.
other countries. Secondary centres: Assam, other Japan. A shrub cultivated there for its fragrant
parts of India (p. 70) and Ceylon. The taxonomy flowers.
of Camellia is confusing. Some group the cultiva-
ted escapes and putative wild plants in one species EDGEWORTHIA PAPYRIFERA Sieb. & Zucc. (syn.
C. sinensis, and recognize subspecies, varieties Daphne papyrifera Sieb. ). 2n=36, (72). Japan.
and types; others have given some subspecies/ Occasionally cultivated there.
varieties/types specific rank. Wu et al. (1970)
suggested that the wild tea populations originated Tiliaceae
from heterogeneous populations of species hybrids. CORCHORUS OLITORIUS L. Jute, Tossa jute,
The main types a r e China tea (C. sinensis var. Jew's mallow. 2n=14. S. China and probably taken
sinensis L., syn. C. sinensis L. ) and Assam tea to India/Pakistan. Cultivated as a spinach in the
(C. sinensis var. assamica (Masters) Wight, syn. Ganges-Brahmaputra delta, where it may have run
C. assamica Masters). The latter type includes wild and in the Middle East, Egypt, Sudan and trop.
the Cambodia race (C. assamica ssp. lasiocalyx Africa.
Planch.-MS, syn. Thea lasiocalyx Planch. ). At
Tocklai, Assam Wilson's Camellia (C. i r r a w a - Trapaceae
diensis P . K. Barua) is found. The F o r e s t ' s Ca-
mellia (C. taliensis Sealy) produces tea of low TRAPA BICORNIS L. 2n= . China. A waterplant
quality. It is probably a hybrid of C. sinensis and cultivated there, Korea and Japan.
C. irrawadiensis and should be included in the
latter species. Darjeeling tea also is a possible TRAPA BISPINOSA Roxb. Singhara nut. 2n=
product of C. sinensis introgression in C. a s s a m i - India, China and Japan. A waterplant cultivated
ca (C. irrawadiensis) (Visser, 1969). for the seeds in India.
Leaves a r e widely used to make tea; in Burma
leaves a r e used to prepare pickled tea as food TRAPA NATANS L. Water chestnut, Water c a l -
(Simura et a l . , 1967). trop. 2n=c. 36, 40, 48, c. 48. Europe, Medi-
terranean region and Asia. It was cultivated in
C. wabiske* might be a natural hybrid of C. sinen- S. China (Li, 1970). Elsewhere the seeds have
sis and C. japonica* and so a r e the 'China hybrids' been consumed since neolithic t i m e s .
and Cambodian varieties of tea (Bezbaruah &
Gogoi, 1972). Typhaceae
TYPHA LATIFOLIA L. Cat-tail. Reeds mace,
Bull rush, Marsh beetle. 2n=30. A cosmopolitic
plant cultivated in China.
Umbelliferae
ANGELICA KIUSIANA Maxim. 2n=22. China. An
old Chinese vegetable. It is a weed now (Li, 1969).
ANGELICA POLYMORPHA Maxim. 2n=22. China.

Cultivated there as a medicinal crop.
CRYPTOTAENIA JAPONICA Hassk. Mitsube.

2n=(18), 20, 22. Cultivated in Japan.
GLEHNIA LITORALIS F. Schmidt. 2n=22. N. and

E. Asia. Cultivated in Shantung, China.
PHELLOPTERUS LITTORALIS Benth. Hama-bôhû.

2n=22. Sachalin, Japan and China. Cultivated in
Japan (Uphof, 1968; Kihara, 1969).
Camellia sinensis
Urticaceae
BOEHMERIA NIVEA (L. ). Gaud. Ramie, Rhea,
China g r a s s . 2n=28, (42). S. and C. China, on
CAMELLIA WABISKE Kitamura. Wabisuke. Taiwan and S. Japan. Cultivated in China since
2n=30. Cultivated in Japan. This species is con- ancient times, where there a r e many varieties,
sidered as a natural hybrid of C. sinensis* and Japan, Philippines and other trop, countries.
C. japonica* L. If so it could probably be used Var. chinensis, Chinese ramie, White ramie,
as a bridge species between these two species, (2n=28) may have originated in SW. China, while
because hybridization of them has failed so far. var. indicum (syn. B. utilis Blume, B. tenacissi-
ma Gaud. (2n= ), Indian ramie. Green ramie
THEACEAE - ZINGIBERACEAE 41
might be a derivative of a cross between var.

chinensis and an unknown species.
Violaceae
VIOLA VERUCUNDA A. Gray. Chin. 2n=24. China.
An old chinese vegetable. Now it is a weed (Li,
1969).
Vitadaceae
VITIS AMURENSIS Rupr. (syn. V. thunbergii
Regel, V. shiragai Makino). Amut grape. 2n=38.
Primorye and Khabarovak and NE. China. This
species belongs to the Chinese centre of origin of
Vitis-species. It withstands -40°C. It is a possible
source of winterhardiness for V. vinifera*.
Occasionally cultivated.
VITIS DAVIDII (Rom.) Foex. Spring vitis. 2n=

The Tziansu and Yunna Provinces, China. Occa-
sionally cultivated there.
Zingiberaceae
ALPINIA CHINENSIS Rose. 2n=48. China and
Indo-China. The rhizomes a r e used in Chinese
medicine and the leaves for fibre. Sometimes
cultivated.
ALPINIA OFFICINARUM Hance. L e s s e r galangal,

Small galangal. 2n= . E. and SE. Asia. Culti-
vated in China.
AMOMUM GLOBOSUM Lour. Round Chinese c a r -

damon. 2n= . China. Cultivated there.
ZINGIBER MIOGA (Thunb. ) Rose. Mioga ginger,

MyÔga. 2n=55. Japan. Cultivated there (Kihara,
1969).
2 Indochinese-
Indonesian Centre
Vavilov called the Indochinese-Indonesian centre the Tropical Asian Centre of Origin. Darlington (1956)
and Li (1966 cited by Chang, 1970) divided this region into S. Asia: Burma, Thailand and Indochina, and
SE. Asia: Malay peninsula and the Malay achipelago. Li described for Centre 2 an agriculture mainly
based on vegetatively propagated crops. Harlan (1971) considered this region as a noncentre B2 Southeast
Asian and South Pacific noncentre, as agriculture may have been introduced into this Centre.
The oldest agricultural remains come from centre 2: the Spirit Cave 60 km N. of Mae Hongson, Nw.
Thailand (Gorman, 1969). This coincides with Sauer's (1959) conclusion that the Old World centre of
development of agriculture was situated in the NW. part of Centre 2 (about present Burma). The Spirit
Cave was inhabitated from ca. 10 000 to 5 600 BC. Solheim (1972) proposed that horticulture may have
developed in 20 000 - 15 000 BC., while during 15 000 - 8 000 BC. there was further domestication of
plants and domestication of animals, resulting in large-scale agriculture and animal husbandry.
Another archeological site is at Non Nok Tha, NE. Thailand. It dates from perhaps 5 000 BC.
In the Spirit Cave remains of Prunus, Terminalia, Areca, Vicia or Phaseolus, Pisum or Raphia,
Lagenaria and Trapa, in another layer Piper, Madhuce, Canarium, Aleurites and Areca had been found,
while in a third layer Canarium, Lagenaria and Cucumis had been discovered (Gorman, 1969). Further
work should support the taxonomie identifications. For instance Schultz-Motel (1972) could not accept that
Vicia faba* could have,been found. Chang (1970) used the presence of Vicia faba/Phaseolus, P i s u m /
(Raphia), Lagenaria, Trapa and Cucumis as proofs that these plants were actually cultivated.
If this conclusion is correct this centre should be according to Harlan (1971) a centre and not a non-
centre.
This region is important for crops such as bamboos, tropical fruit t r e e s , ginger plants, Cocos nuci-
fera, Colocasia esculenta, Dioscorea spp., Musa spp., wild and weedy Oryza spp., Piper spp., and
Saccharum officinalis.
AMARANTHACEAE - ARACEAE
Amaranthaeeae Araceae
AMARANTHUS GANGETICUS L. 2n=34. Asia. ALOCASIA INDICA (Roxb.) Schott. 2n=
Cultivated in India, Malaya, China and Japan as Centre of diversity SE. Asia. Cultivated there
a spinach. See also A. mangostanus*. for its stem which is eaten and as an ornamental.
Introduced to other countries such as India. This
AMARANTHUS MANGOSTANUS Juslen (syn. species is sometimes included in A. macrorrhiza*.
A. tricolor L. v a r . mangostanus Thell. ) 2n=32.
Trop. Asia. Cultivated as a potherb. In A. t r i c o - AMORPHOPHALLUS CAMPANULATUS (Roxb.)
lor are sometimes included v a r . gangeticus and Blume. Elephant yam. 2n=28. SE. Asia. Culti-
var. tricolor (syn. A. melancholicus L.). See vated in C. and E. Java and India (p. 63).
A. gangeticus*.
AMORPHOPHALLUS HARMANDII Engl. &Gehr.
AMARANTHUS PANICULATUS L. 2n=32. Used as 2n= . Occasionally cultivated in Tonkin.
a potherb and grain crop in SE. Asia. It might be
conspecies with A. cruentus* (Sauer, 1950) or a AMORPHOPHALLUS RIVIERI Dur. 2n=24, 26, 32,
synonym. 39. Indo-China. Var. konjac (Schott) Engl. Philip-
pines. Cultivated in China and Japan (Mansfeld,
Anacardiaceae 1959).
BOUEA MACROPHYLLA Griff. 2n= A fruit COLOCASIA ESCULENTA (L.) Schott. Dasheen,
tree of the wet tropics of SE. Asia. Taro, Cocoyam. 2n=28, 42. SE. Asia. It was an
early domestieant introduced into China and
MANGIFERA CAESIA Jack. 2n=40. P r i m a r y centre Japan, where var. antiquorum* developed, the
Indonesia. This fruit t r e e is cultivated there and Mediterranean region, W. Africa, the Pacific
elsewhere. islands, New Guinea, Samoa and New Zealand. In
SE. Asia var. esculenta (syn. C. esculenta s. s . ,
MANGIFERA FOETIDA Lour. Bachang mango. C. esculenta var. t y p i c a A . E . Hill) developed.
2n=40. Indo-Chlna and Malaysia. Cultivated in
Many socalled wild plants are probably d é r i -
Java.
vâtes of plants that have run wild (Purseglove,
1972).
MANGIFERA ODORATA Griff. Kurwlni mango.
2n= . Malaysia. Cultivated in Java. It is
closely related to M. indica* (Rhodes et a l . , 1970).
SEMECARPUS ANACARDIUM L.f. (syn. Ana-

cardium orientale L.). Cashew marking nut tree,
Oriental cashew nut. 2n=60. Trop. Asia and
Australia. Cultivated in the tropics.
SPONDIAS LAOSENSIS P i e r r e . 2n= . Trop.

Asia. At r e e cultivated for its fruits.
SPONDIAS PINNATA (Koen. &L.f. ) Kurz (syn. S.

mangifera Willd. ). Hogplum. 2n= . Trop.
Asia. A fruit t r e e whose flower clusters are also
eaten.
Annonaceae
CANANGA ODORATUM Lamb. Ylang-ylang.
2n=16. Malaysia. Cultivated there and in other
countries for the flowers which a r e a source of
essential oils.
STELECHOCARPUS BURAHOL (Bl. )Hook,f. &

Thorns, (syn. Uvaria burahol Bl.). Burahol.
2n= . Malaya and Java. Cultivated in Java for
its fruits.
Apocynaceae
ERVATAMIA CORONARIA Stapf, (syn. Tabernae-
montana coronaria Willd., T. divaricata R. Br.).
Grape jasmine. 2n=22. Malaya. Cultivated there Colocasia esculenta
for various purposes.
INDOCHINESE-INDONESIAN CENTRE 44
CYRTOSPERMA CHAMISSONIS (Schott. ) Merr. DURIO KUTEJENSIS (Hassk. ) Beccari. Lai.

(syn. C. edule Schott, C. merkusii (Hassk. ) 2n= . Along the foothills of the central ranges
Schott. 2n= . Indo-Malaysian region. Intro- of Borneo. It has spread now to E. and N. of
duced into many Pacific islands. Cultivated for Borneo.
its tubers (Purseglove, 1972).
DURIO OXLEYANUS Griffith, Kerantongan.
PISTIA STRATIOTES L. Water lettuce, Tropical 2n= . Malaya, Sumatra and Borneo. The fruits
duckweed. 2n=28. Subtropics and tropics of the are eaten and seeds a r e dispersed around the
Old en New Worlds. A floating plant. Cultivated (temporary) settlements. Occasionally cultivated.
in Java in fishponds for edible shrimps that live Other wild species of which the fruits a r e eaten
below the plants (Uphof, 1968). a r e D. graveolens Beccari, tabelak (2n= ),
D. dulcis, lahong (2n= ), and D. grandiflorus
Araliaceae (Mast. )Kostermans &Soegeng, durian munjit
(2n= ). D. graveolens grows wild in Borneo,
NOTHOPANAX FRUTICOSUM Miq. (syn. Panax
Malaya and Sumatra, D. dulcis in Borneo.
fruticosum L., Polyscias fruticosa (L. ) Harms).
2n=22, 24. SE. Asia and Polynesia. Cultivated in
Java for its roots and leaves. DURIO ZIBETHINUS Murray. Durian. 2n=56.
W. Malaysia. Unknown wild. Cultivated throughout
SE. Asia, W. Irian, the Moluccas, Celebes, S.
NOTHOPANAX GUILFOYLEI Merr. (syn. Panax
Philippines, Indonesia and Malaya. Introduced to
guilfoylei Cogn., Aralia guilfoylei Bull. ). 2n=
Indochina, Thailand, Burma, Ceylon and elsewhere.
Probably the Pacific islands. Perhaps it is a cul-
It is often semi-cultivated i. e. semi-wild t r e e s
tigen of N. pinnatum* or a closely related species'
result from dispersal of seeds. In this way durian
(Mansfeld, 1959).
groves have arisen.
NOTHOPANAX OBTUSUM Miq. (syn. Panax obtu-
sum Blume, Polyscias obtusa (Bl. ) Harms. Boraginaceae
2n= . Probably SE. Asia. Cultivated in Java. TOURNEFORTIA ARGENTEA L.f. (syn. M e s s e r -
schmidia argentea (L.f. ) Johnston). Velvet leaf.
NOTHOPANAX PINNATUM Miq. (syn. Panax 2n= . Trop. Asia. A shrub cultivated for its
pinnatum Lam., Polyscias rumphiana Harms. leaves which a r e used as smoking tobacco.
2n= . SE. Asia and New Guinea. Cultivated
for its leaves. Burseraceae
CANARIUM COMMUNE L. Java almond. 2n=
Asclepiadaceae
The Moluccas (Indonesia). Cultivated in many
GYMNEMA SYRINGIFOLIUM Boerl. Sajor pepe. other tropical countries. The kernels a r e eaten
2n= . Cultivated in Malaya as a vegetable. and oil is extracted from them. Sometimes they
are planted as shade trees and as ornamentals.
Averrhoaceae
AVERRHOA BILIMBI L. Bilimbi. 2n=22, 24. CANARIUM MOLUCCANUM Blume. 2n= . The
Moluccas, New Guinea and W. Polynesia. Culti-
Malaya. Its fruits are very acid.
vated in Malaysia.
AVERRHOA CARAMBOLA L. Carambola. 2n=22,
24. Indonesia. Cultivated for its fruits. Some CANARIUM OVATUM Engl. Pili, Pili nut.
varieties have been developed. 2n= . S. Luzon (Philippines). The kernels con-
tain 70-80% pili-nut oil.
Basellaceae
CANARIUM PIMELA Koenig. Black Chinese olive.
BASELLA RUBRA L. Indian spinach, Ceylon s p i - 2n= . E. Asia. Cultivated in S. China and
nach, Malabar nightshade. 2n=44, 48. Probably Cochinchina.
S. Asia (Winter, 1963). Cultivated as a vegetable
now throughout the tropics. Formerly it may have Combretaceae
been used for dyeing. The commonest synonyms
a r e B. alba L. (2n=48), with white flowers, and QUISQUALIS INDICA L. Rangoon creeper.
B. cordifolia Lam., (2n= )with heart-shaped 2n= . SE. Asia. A woody vine. Cultivated as
an ornamental, vegetable and as an anthelmintic.
leaves.
TERMINALIA BELLIRICA (Gaertn. ) Roxb. (syn.
Bombacaceae
Myribalan bellirica Gaertn. ). Bellirica, T e r m i -
CEIBA PENTANDRA Gaertn., var. indica (DC.) nalia. 2n=26, 48. India and Malaysia. Cultivated
Bakh. Kapok tree, Silk cotton t r e e . 2n=72, 80, 88. as a source of myrobalan which is used for tanning
Secondary gene centre: SE. Asia. Introduced from leather, for black dye and for making ink.
Africa (p. 108) probably via India. Cultivated in
SE. Asia (Zeven, 1969). The Indonesian cultivar TERMINALLY CATAPPA L. Indian almond, Myro-
Reuzenrandoe (giant kapok) bears some c h a r a c t e - balan, Almendro. 2n=24. Trop. Asia. N. Australia
ristics of the var. caribaea*. and E. Polynesia. Widely cultivated in the tropics.
The kernels contain about 55% oil. Used for manu-
ARACEAE - DIOSCOREACEAE 45
facturing edible fats, cosmetics and pharmaceutic SCIRPODENDRON GHAERI (Gaertn. ) Merr. (syn.
preparations. Used for timber; leaves and bark S. costatum Kurz. ). 2n= . Trop. Asia, Samoa
a r e used for preparation of medicines. The fruits and Australia. Cultivated in Sumatra for mat
a r e edible. making.
TERMINALIA CHEBULA Retz. 2n=14, 24, 26, 48. Dioscoreaceae

India to Malaysia.
DIOSCOREA ALATA L. Greater yam, Water yam,
Winged yam, Ten months yam. 2n=(20), 30, 40,
Compositae 50, 60, 70, 80. SE. Asia, in the Assam-Burma
BLUMEA BALSAMIFERA (L.). DC. 2n=20. Hima- region it is the cultigen of D. hamiltonii Hook.,
laya, India, Malaysia, S. China and Taiwan. Cul- 2n= , or D. persimilis Prain &Burk, 2n= ,
tivated in Java as a medicinal crop. or a similar species (Burkill, 1935). Some types
have been described as D. atropurpurea Roxb.,
BLUMEA MYRIOCEPHALA DC. 2n= . India, 2n= , and D. purpurea Roxb., 2n=
Vietnam, Malaya and Indonesia. Occasionally cul-
tivated in Vietnam.
CARUM ROXBURGHIANUM Benth. (syn. Trachy-

spermum roxburghianum (DC.)Wolff). Ajmud.
2n=18. Trop. Asia, Unknown wild. Cultivated in
Indochina, Ceylon and India.
ENHYDRA FLUCTUANS Lour. (syn. E. helonchu

D C , Hingtsha repens Roxb. ). 2n=22. India, Indo-
china, Thailand, China and Indonesia. A water
plant occasionally cultivated in Cambodia and Ma-
laya for its leaves.
EUPATORrUM STOECHADOSUM Hance. 2n=40.

Vietnam. Cultivated there.
SPILANTHES PANICULATA Wall, ex DC. 2n= Dioscorea alata ( ), D. esculenta (-- -) and D. hispida (...)
SE. Asia and New Guinea. Cultivated as a vege- (Harris, 1973)
table or salad.
VERNONIA ANTHELMINTICA Willd. Kinka oil DIOSCOREA BULBIFERA L. Potato yam, Aerial
iron weed. 2n=20, 54. Trop. Asia. It might be a yam, Bulbil-bearing yam. 2n=36, 40, 54, 60, 80,
source of epoxy fatty acids. 100. Trop. Asia and Africa. Possibly it was do-
mesticated in Asia as well as Africa (p. 111). Cul-
Convolvulaceae tivated in Trop. Asia, Africa, Oceania and the
West Indies. The tubers and bulbils a r e edible.
IPOMOEA MAMMOSA Chois. 2n= . Abouana,
The African form (p. I l l ) has been described as
Philippines. Cultivated in Indochina. Formerly it D. latifolia Benth., 2n= . There a r e many
was erroneously believed that this species was the types which often have been described as species
ancestor of I. batatas*. e.g. D. heterophylla Roxb., 2n=
Cucurbitaeeae DIOSCOREA ESCULENTA (Lour. ) Burk. Lesser
BENINCASA HISPIDA (Thunb.) Cogn. (syn. B. yam, Asiatic yam, Potato yam, Fancy yam, Chi-
cerifera Savi). Wax gourd, White gourd. 2n=24. nese yam. 2n=40, 60, 80, 90, 100. Indo-China.
Java. Cultivated throughout trop. Asia (Purseglove, Cultivated in S. China, and later throughout the
1968). It was already mentioned as a vegetable in tropics.
China in 500 AD (Li, 1969).
DIOSCOREA FLABELLIFOLIA Prain. & Burk.
TRICHOSANTHES ANGUINA L. Edible snake gourd. 2n= . Malaya. Occasionally cultivated there.
2n=22. Trop. Asia from India (p. 64) to Australia.
DIOSCOREA HISPIDA Roxb. (syn. D. hirsuta
Cyperaceae Dennst, D. triphylla L. ). 2n=40, (80). India
(p. 64) and SE. Asia. Closely related to the Afri-
FIMBRISTYLIS GLOBULOSA (Retz.) Kunth. can D. dumentorum (Coursey, 1967).
2n= . Trop. Asia, Ceylon, India, Malaysia and
Mariannes. Cultivated on Malaysia for matmaking
DIOSCOREA NUMMULARIA Lam. 2n= SE.
etc.
Asia. Cultivated there and in Indonesia and
Oceania. It closely resembles D. cayenensis*.
LEPIRONIA ARTICULATA (Retz. ) Domin. (syn.
L. mucronata Rich. ). 2n= . SE. Asia, Malaysia,
DIOSCOREA QUARTINIANA A. Rich. 2n=
Australia and Fiji. Cultivated in Indonesia.
Throughout trop. Asia. Cultivated in E. Nigeria
(Coursey, 1967).
DIOSCOREA PENTAPHYLLA L. 2n=40, 80, 144, PHYLLANTHUS DISTICHUS (L. ) Muell. -Arg.
c. 144. SE. Asia. Cultivated throughout Indonesia (syn. Ph. acides (L.) Skeels. Otaheite gooseberry.
and the Pacific islands. 2n=26, 28. India and Madagascar. Cultivated in
the tropics for its fruits (Purseglove, 1968).
Dipterocarpaceae
PHYLLANTHUS EMBLICA L. Emblic, Myrobolan.
SHOREA STENOCARPA Burck. 2n= . Malaya.
2n=28, 98. Trop. Asia. Cultivated in the Old and
Cultivated for its seeds, a source of a Borneo
New Worlds for its fruits (Uphof, 1968).
tallow.
SAUROPUS ALBICANS Blume (syn. S. androgynus
Ebenaceae Merr. ). 2n= . Cultivated as a vegetable in SE.
Asia.
DIOSPYROS DISCOLOR Willd. (syn. D. blancoi
How. ). Malobo, Velvet apple. 2n= . Malaysia
and Philippines. Occasionally cultivated for its TRIGONOPLEURA MALAYANA Hook.f. Gamber
edible fruits. ooran. 2n= . The Malayan Archipelago. Culti-
vated there for its leaves which substitute for
Uncaria gambir*.
MABA MAJOR Forst, f. 2n= . Cultivated for
its fruits on the Friendship Islands.
Flacourtiaceae
Elaeocarpaceae FLACOURTIA RAMONTCHI L'Hér. Botoko plum,
Madagascar plum, Governor's plum, Ramontchi.
ELAEOCARPUS FLORIBUNDUS Blume. 2n=
2n=22. Malaya and Madagascar. Cultivated in the
From Bangla Desh to Java. Cultivated in Bengal
tropics for its fruits.
and Assam for its fruits.
Euphorbiaceae FLACOURTIA RUKAM Zoll. &Mor. Rukam.

2n= . Malaysia and Philippines. A t r e e culti-
ALEURITES MOLUCCANA Willd. Tung oil t r e e . vated for its fruits.
2n=44. Autotetraploid. Indonesia. Its wild parent
is not known. Crosses between this species and HYDNOCARPUS ALCALAE C. DC. 2n=
A. montana did not succeed (Wit, 1969b). Philippines. Cultivated for its seeds which a r e a
source of oil used to cure leprosy.
ALEURITES TRISPERMA Blanco. Tung oil tree,
Banucalang. 2n=22. The Philippines. Cultivated HYDNOCARPUS ANTHELMINTHICUS P i e r r e ex
there, in Malaya and Indonesia. Crosses with Lanessan. 2n=24. Indochina and Siam. Cultivated
A. montana did not succeed (Wit, 1969). in many trop, countries for the seeds which a r e
a source of oil used to cure leprosy.
BACCAUREA DULCIS Muell. -Arg. Tjoopa.
2n= . Malaya and Indonesia. Cultivated there HYDNOCARPUS KURZII (King) Warb. 2n=
for its fruits (Uphof, 1968). Ssp. kurzii in Burma highlands and Assam. Ssp.
australis in Burma lowlands and N. Siam. Ssp.
BACCAUREA MOTLEYANA Muell. -Arg. Rambai. kurzii is cultivated in many tropical countries for
2n= . SE. Asia. Cultivated there for its fruits the seeds which a r e a source of oil used to cure
(Uphof, 1968). leprosy.
BACCAUREA RACEMOSA Muell. -Arg. 2n= PANGIUM EDULE Reinw. Pangi. 2n=
The Malayan Archipelago. Cultivated there for Malaysia. Cultivated in Java.
its fruits.
Gnetaceae
GLOCHIDION BLANCOI Lowe. 2n= . A tree
GNETUM GNEMON L. Bulso. 2n= . From
cultivated in the Far East, and Philippines for the
Assam to Malaysia and Fidji. Var. ovalifolium
young leaves and shoots (Terra, 1967).
(Poir. ) Bl. is considered the wild type while var.
HEVEA BRASILIENSIS (Willd. ) Muell. -Arg. Bra- gnemon is the cultivated type planted in Java. In-
zilian hevea. P a r a rubber t r e e . 2n=36. Amazon troduced to Java, Sumatra and elsewhere. Culti-
basin (p. 148). A secondary gene centre: Malaya. vated in SE. Asia for its seeds.
Domesticated in SE. Asia at the end of the 19th
Century (Purseglove, 1968). Gramineae
ANDROPOGON ACICULATUS Retz. (syn. Chryso-
MANIHOT ESCULENTA Crantz. Cassava. 2n=36. pogon aciculatus Trin. ). 2n= . The tropics.
America (p. 148 and 165). Secondary centre of Cultivated in Vietnam for its roots which contain
diversity in Indonesia. Chiendent grenille â b r o s s e (Uphof, 1968).
PLUKENETIA CORNICULATA Smith, (syn. BAMBUSA ARUNDINACEA (Retz. )Willd. Spiny

Pterococcus corniculatus Pax & Hoffm. ). Paina- bamboo. 2n=70. P r i m a r y centre: India and Burma
paina. 2n= . SE. Asia. Cultivated as a vege- (p. 64). Secondary centre: Malaysia and E. Java.
table.
DIOSCOREACEAE - 3RAMINEAE 47
BAMBUSA CORNUTA Munro. 2n= . Java. A GIGANTOCHLOA APUS (Schult.) Kurz. 2n=
woody grass cultivated for its tender shoots which Burma and Indochina. Several species a r e culti-
a r e used as a vegetable. vated in Java, Borneo and Philippines and on the
Malaya Peninsula (Tenasserim). Secondary
BAMBUSA SPINOSA Roxb. 2n= . Philippines centre: Java.
and Indonesia. A woody, tall grass cultivated as
a timber bamboo and also for its young shoots GIGANTOCHLOA LIGULATA Gamble. 2n=
used as a vegetable. N. part of Malaya Peninsula and Thailand. The
timber is used and the shoots a r e eaten.
BAMBUSA STRICTUS Nees. 2n=70, 72. India and
Burma {p. 64). Secondary centres: Indochina and GIGANTOCHLOA MAXIMA Kurz. 2n= . Un-
S. China (p. 32). known wild. Secondary centre: Java. Its stems
a r e an excellent material for building.
BAMBUSA TULDA Roxb. 2n= . India, Burma
(p. 64) and Tahiti. Secondary centre: Java. GIGANTOCHLOA SCORTECHINII Gamble. 2n=
The Malaysian Peninsula. Its stems a r e used as
BAMBUSA VULGARIS Schrad. ex Wendl. 2n=72. building material.
Probably Malaysia or India. It is unknown wild.
Cultivated in the tropics for its young shoots and GIGANTOCHLOA SCRIBNERIANA M e r r . 2n=
for its stems. Laos and Cambodia. Its stems a r e used as buil-
ding material.
COIX LACRYMA-JOBI L. Job t e a r s , Adlay. 2n=20.
Trop. Asia. Probably first domesticated as a GIGANTOCHLOA VERTICILLATA (Willd. ) Munro.
cereal in Indochina. Cultivated in the tropics now. 2n= . Unknown wild. Cultivated on the Malay
Archipelago. Its stems a r e used as building m a t e -
CYMBOPOGON CITRATUS (DC.) Stapf (syn. rial and for the paper industry. The sprouts a r e
Andropogon citratus DC. ). Lemon g r a s s . 2n=40, eaten. There a r e several varieties.
60. Probably Malaysia or Ceylon. Unknown wild.
Cultivated in S. Asia, Indochina and elsewhere ISCHAEMUM INDICUM (Houtt. ) M e r r . Batiki blue
for its lemongrass oil. g r a s s . 2n= . SE. Asia. Cultivated in W. Africa,
W. Indies, Fiji and elsewhere (Purseglove, 1972).
CYMBOPOGON NARDUS (L.) Rendle (syn. Andro-
pogon nardus L. ). Citronella g r a s s . 2n=20, (40, ORYZA GRANULATA Nees &A m . (incl. O. meye-
60). Cultivated in Indonesia, Ceylon and elsewhere riana Baill. ). 2n=24, 48. Malaya. It belongs to the
for its citronella oil. There a r e two types of oil: 'officinalis' group of Oryza (p. 47).
1. Ceylon type obtained from var. lenabatu which
is cultivated in S. Ceylon and 2. Java type obtained ORYZA LONGIGLUMIS Jansen. 2n=48. New Guinea.
from var. mahapengiri (syn. C. winterianus
Jowett, 2n=20). The latter was introduced into ORYZA MINUTA P r e s l . 2n=48, genome formula
Java from Ceylon early in 20th Century. It is now BBCC. This wild species has the same genomes
widely distributed throughout the tropics. as the African O. eichingeri*. It belongs to the
'officinalis' group.
DENDROCALAMUS ASPER (Schult. ) Becker ex
Heyne (syn. Bambusa asper Schult. ). 2n= ORYZA NIVARA Sharma &Shastry. 2n=24. S. and
Probably from the Malay Peninsula and adjacent SE. Asia and N. Australia. It may include the
a r e a s . Unknown wild. Secondary centre: Malay 'old' species O. fatua*, O. sativa f. spontanea*
Archipelago. It has strong stems and edible shoots. and O. rufipogon*.
DENDROCALAMUS BRANDISH Kurz. 2n=72. ORYZA OFFICINALIS Wall. 2n=24, genome formula
Burma, Thailand, Cambodia and Vietnam. Its CC. This wild species is the parent of the species
stems a r e used as building material. belonging to the 'officinalis' group
DENDROCALAMUS MERRILLIANUS Elm. 2n= ORYZA PERENNIS Moench. 2n=24, genome for-
Philippines. P r i m a r y centre: Philippines. Its mula AA. The distribution of this wild species is
stems a r e used as building material. discussed on p. 65. In Oceania the Oceanian race
(2n=24) of this species developed. See also O. r u -
DINOCHLOA GIGANTEA Munro. 2n= . Lower fipogon* and index.
Burma. P r i m a r y centre: Lower Burma. Its stems
a r e used as building material. The plant of this ORYZA RIDLEYI Hook,f. 2n=48. SE. Asia.
species is the largest among the bamboos.
ORYZA RUFIPOGON Griff, (syn. O. montana
DINOCHLOA MACLELLANDII Kurz. 2n= L o u r . ) . 2n=24. Several SE.Asian countries. It is
Cambodia, Laos and Vietnam. Its stems a r e used a pernicious weed of rice land. It easily crosses
in the basket industry. with rice. It might be a hybrid product of natural
crosses of rice and O. perennis* and would then
DINOCHLOA PENDULUS Ridb. 2n= . Malaya be of the same nature as O. sativa var. fatua*.
Peninsula. Its stems a r e used for baskets. Recent views of taxonomists include in O. r u -
fipogon: O. perennis*, O. fatua*, O. sativa f. W. Java. From here the chromosome number d e -
spontanea*, O. perennis ssp. balunga*, O. p e - c r e a s e s with distance. Clones with the smallest
rennis ssp. cubensis*. See also O. nivara*. chromosome numbers a r e observed in India. Here
the widest variation of this number is found.
ORYZA SATIVA L. Rice. 2n=24, genome formula
AA. The primary centre: SE. Himalaya region SACCHARUM SPONTANEUM L. 2n=112. Plants
(p. 65). Ecotype Tjereh and Bulu developed in with 2n=112 occur in Indonesia especially in Java
Indonesia. Thereh belongs to the ecospecies and Sumatra. A hybrid origin has been suggested
'aman' (ssp. indica Kato) (Morinaga, 1968). e . g . S. offieinarum* (2n=80) x S. spontaneum
(2n=64).
ORYZA SCHLECHTERI Pilger. 2n= .New
Guinea. SCHIZOSTACHYUS BRACHYCLADUS Kurz.
2n= . Java and E. Malaysia. Secondary centre:
PASPALUM SCROBICULATUM* the Malaysian Peninsula.
SACCHARUM EDULE Hassk. 2n=70-120. P r o b a - SCHIZOSTACHYUS GRANDE Rtdl. 2n= Ma-

bly near Maprik, New Guinea. Cultivated for its laysian Peninsula.
edible inflorescens. It may have developed as a
hybrid of S. robustum and Miscanthus floridulus*. SCHIZOSTACHYUS LULAMPAO Merr. 2n=
A group of closely related cultivars is found in Centre of origin Philippines. Used in the paper
Fiji. It may derive from S. offieinarum* x M. flo- industry.
ridulus (See S. offieinarum*) (Grassl, 1964, 1967,
1968; P r i c e 1963). SCHIZOSTACHYUS ZOLLINGERI Steud. 2n=
Malaysian Peninsula, Java and Sumatra.
SACCHARUM OFFICINARUM L. Sugarcane, Noble
sugarcane. 2n=40II=80. New Guinea. Modern SINOCALAMUS LATIFLORUS (Munro) McClure
clones resulting from hybridization (2n=100-125). (syn. Dendrocalamus latiflorus Munro. ) 2n=
Sugarcane derives from S. robustum Brandes & Burma, Thailand, Taiwan and Philippines. Its
Jeswlet ex Grassl, which grows wild in New stems a r e used as building material. The young
Guinea, Celebes, Borneo upto New Hebrides. shoots a r e eaten. They a r e also canned and e x -
Basic types of northern coast of New Guinea, Ce- ported.
lebes and Borneo have 2n=60, while those from
southern coast of New Guinea have 2n=80. The VETIVERIA ZIZANIOIDES Stapf (syn. V. odorata
first basic type may have originated in Borneo. Virey, Andropogon muricatus Retz. ). Vétiver.
The second basic type may have been the wild 2n=20. A grass of trop. Asia. Cultivated for its
parent from which primitive sugarcane was deve- roots and planted as a hedge.
loped.
S. robustum is cultivated for its large stalks ZEA MAYS L. Maize. 2n=20. Domesticated in C.
which a r e used for fences and for construction. America (p. 166). Secondary centre arose in S. and
During the domestication of sugarcane, geographi- SE. Asia (Brandolini, 1970).
cal types may have hybridized. Due to selection,
sugarcane has a much lower fibre content, in- Guttiferae
creased juiciness and sugar content throughout the
stalk. CALOPHYLLUM INOPHYLLUM L. Alexandrian
S. robustum has been spread from its original laulal, Undi. 2n=32. Coastal regions from E. Afri-
habitat to other places. At higher altitude it hybri- ca upto Australia and Polynesia. Often planted.
dizes with Miscanthus floridulus (Labill. ) Warb. In India it has a rather restricted economic i m -
(2n=38) which has lead to hybrid swarms. From portance. The kernel yields Domba oil.
such a hybrid S. edule* and probably the Hawaiian
Original sugarcanes derive. Some hybrids between GARCINIA ATROVIRIDES Griffith. Gelugur.
S. offieinarum and M. floridulus have been d e s - 2n= . Assam and Malaya. Occasionally culti-
cribed as Erianthus maximus Brongn. and S. pedi- vated.
cellare Trin. The present sugarcane clones have
been derived from artificial hybridization with GARCINIA COCHINCHINENSIS (Lour. ) Choisy.
S. spontaneum*. 2n= . Cochinchina. Cultivated for its fruits.
Moriya (1950) suggested that the sugarcane
GARCINIA DULC1S (Roxb.) Kurz. Baniti. 2n=
plants in SE. Asia and its vicinity can be divided
Philippines to Java. The bark yields a green dye,
into three sections when the chromosome number
while the fruits a r e edible. Occasionally cultivated
is considered:
in Java.
1. Malay Archipelago and South Seas Section
Philippines, Micronesia, New Guinea included
GARCINIA INDICA Choisy. Kokum, Kokan, Ktam-
2. India-Burma Section Turkmenistan in-
bi. 2n=48, c. 54. Trop. Asia. Cultivated for its
cluded, and
fruits. In India it is a minor oil-seed plant (p. 66).
3. Japan islands Section Formosa and
Okinawa included.
GARCINIA MANGOSTANA L. Mangosteen.
He further proposed that clones with the highest
chromosome number a r e found in S. Sumatra and 2n=c. 76, 96. Malaysia. It is considered to be the
GRAMINEAE - LEGUMINOSAE 49
most delecious of all tropical fruits. It is derived LITSEA CALOPHYLLA (Miq. ) Mansf. (syn L. t e -
from wild G. silvestris, which is also found in tranthera Mirb., L. sebifera Blume). 2n=
India (p. 66). Malaya and Indonesia. Cultivated esp. in
Indonesia for its fruits.
GARCINIA MULTIFLORA Champ, (syn. G. tonki-
nensis Vesque). Cây giôc, Bira tai. 2n= Leguminosae
N. Vietnam, Laos, Hainan and Hongkong. Culti-
vated in N. Vietnam for its fruits. ALBIZIA LEBBECK Benth. Lebbek, Indian walnut.
2n=26. Trop. Asia upto N. Australia. Cultivated
GARCINIA PEDUNCULATA Roxb. Tikul. 2n= in the tropics and subtropics as a fodder crop and
Bengal and Silhat (Bangla Desh). Cultivated for as a shade t r e e .
its fruits.
ALBIZIA MOLUCCANA Miq. (syn. A. falcata
GARCINIA TINCTORIA (DC.) W. F. Wight. Matau, (Stickm. ) Backer. 2n= . Malaya. Cultivated
Gamboge t r e e . 2n=c. 80. India (p. 66) and Malaya. there and elsewhere as a shade tree and as a
Cultivated in the tropics for its fruits. green manure.
Hydrophyllaceae ALBIZIA MONTANA (Jungh. ) Benth. 2n=

Malaysia. Cultivated as a green manure and shade
HYDROLEA ZEYLANICA Vahl. 2n= . Trop. tree.
Asia. Cultivated in Java for its young leaves.
ALBIZIA SUMATRANA . 2n= . Indonesia. Cul-
Labiatae tivated in Zaire as a soil cover, green manure and
COLEUS AMBOINICUS Lour. (syn. C. aromaticus shade t r e e .
Benth. ). Indian borage, Daoon ajenton 2n=32.
Indonesia. Cultivated in SE. Asia and West Indies CANAVALIA GLADIATA (Jacq. ) DC. Sword bean.
for its aromatic leaves. These leaves a r e used in 2n=22, 44. The Old World. Probably derived from
stuffings and for flavouring meats. They may s u b - C. gladiolata Sauer (2n=22), which occurs in the
stitute for sage (Salvia officinalis*) and borage Burma-Yunnan region (Sauer, 1964). Wild in t r o -
(Borago officinalis*) (Purseglove, 1968). pical Asia and Africa. Cultivated in Asia especially
in India as a food, forage and cover crop or as a
COLEUS PARVIFLORUS Benth. (syn. C. tubero- green manure. In some areas it has naturalized
sus Benth. ). 2n=56, 64. This tuber crop is culti- (Purseglove, 1968). In Japan the white seeded cul-
vated in SE. Asia. tigen (var. alba) is cultivated (p. 34).
C. polystacha (Forsk. ) Schweinf. (2n= ). Cul-
tivated from SW. China upto Ethiopia/Somaliland
OCIMUM BASILICUM L. (syn. O. americanum L. ).
for its pulses and seed. It is also considered as
Basil, Sweet basil. 2n=48. Trop. Asia. Cultivated
the parental type of C. gladiata.
in China since 500 AD (Li, 1969). It has spread to
many regions now.
CASSIA DIDYMOBOTRYA Fresen. Candelabra t r e e .
OCIMUM GRATISSIMUM L. 2n=40, 48, 64. Trop. 2n=28. A shrub used for green manure in Malaya
Asia, esp. India. Cultivated in India as a medici- and Ceylon.
nal crop.
CASSIA HIRSUTA L. 2n=28, 56. A vigorous bush
OCIMUM SANCTUM L. Holy basil. 2n=64. A shrub used in Malaya, Indochina and Uganda for soil
of the trop. Old World. Cultivated as a sacred cover.
plant in India and elsewhere.
CASSIA LESCHENAULTIANA DC. 2n=48. A shrub
ORTHOSIPHON STAMINEUS Benth. (syn. Ocimum used in India and Indonesia as a green manure.
grandiflorum Blume). 2n= . SE. Africa to
Australia. A shrub cultivated in Java as medicinal CASSIA MIMOSOIDES L. 2n=16, (32). Trop. Asia
plant. and Africa. This t r e e is used in Indochina and In-
donesia as a green manure.
POGOSTEMON CABLIN (Blanco) Benth. Patchouli.
2n= . Philippines. Cultivated for its essential CASSIA OCCIDENTALE L. Coffee senna, Negro
oil. coffee, Stink weed. 2n=26, 28. Tropics. Used in
Indochina for green manure.
Lauraceae
CASSIA PUMILA Lam. 2n= Cover crop in
CINNAMOMUM BURMANI Blume. Batavia cinna- Indochina.
mon. 2n= . Malaysia. Cultivated there.
CASSIA SIAMEA Lam. (syn. C. florida Vahl. ).
CINNAMOMUM CASSIA Blume (syn. C. a r o m a t i - 2n=28. India, Malaya and Indonesia. Cultivated in
cum Nees). Cassia cinnamon, Chinese cinnamon. Malaya and India as a fodder crop. Introduced on
2n= . Cultivated in S. China for its bark and Cuba as a green manure.
flower buds. Cassia oil is obtained from the leaves
(Purseglove, 1968). CASSIA TORA L. Sickle senna, Wild senna. 2n=26,
(28, 56). Tropics. Occasionally cultivated as a MUCUNA COCHINCHINENSIS (Lour.) A. Cheval,

green manure in China and Indonesia. (syn. M. n i v e a D C , Stizolobium niveum O.
Kuntze). 2n=22. Cochinchina. Cultivated in tropics
CLITORIA LAURIFOLIA Poir. (syn. C. cajanifo- as a vegetable, for its seeds, as a green manure
lia Barth). 2n=24. Tropics. Occasionally cultiva- and soil cover.
ted in Ceylon and Indonesia and formerly in Tanza-
nia as a green manure. MUCUNA DEERINGIANUM (Bort.) Small, (syn.
Stizolobium deeringianum Bort. ). Florida velvet
CLITORIA TERNATEA L. Butterfly pea, Kordofan bean. 2n=22. Probably trop, Asia or Malaysia.
pea. 2n=16. Probably trop. Asia. Widespread in Cultivated as a cover crop, green manure and
the tropics and cultivated as a fodder and soil forage crop.
cover crop.
MUCUNA PRURIENS DC. var. utilis Wahl. (syn.
CROTALARIA ALATA Ham. 2n=16. Malaya/Indo- Stizolobium aterrimum Piper &Tracy). Bengal
nesia. It is an excellent green manure. bean. 2n=22. Probably trop. Asia. Widely culti-
vated as a cover crop and green manure in the
DERRIS DALBERGIOIDES Baker. 2n= . It is tropics.
used as a shade t r e e in SE. Asia.
NEPTUNIA OLERACEA Lour. 2n=c. 52, 54. The
DERRIS ELLIPTICA Benth. D e r r i s . 2n=22, 24, Tropics. A water plant cultivated as a vegetable
36. From E. India to New Guinea except in S. Ma- in Indochina.
laya. Clones a r e distributed locally except one
which is found in many places in SE. Asia. This PARKIA SPECIOSA Hassk. 2n= Malaysia. It
clone has a high content of rotenone (Toxopeus, is cultivated.
1952).
PELTOPHORUM PTEROCARPUM Backer (syn.
DERRIS MALACCENSIS Prain. D e r r i s . 2n=22, 24. Cäesalpinia arborea Zoll., Inga pterocarpa DC. ).
Malaysian Archipelago where also cultivated types Soga. 2n= . SE. Asia upto Australia. Cultiva-
a r e found. It is like D. elliptica* a source of r o t e - ted on Java for its bark which is a source of brown
none (Toxopeus, 1952). dye.
DERRIS MICROPHYLLA (Miq. ) Jackson. 2n= PHASEOLUS AUREUS Roxb. (syn. Vigna radiata
Used as shade tree in SE. Asia. Introduced into (L. ) Wilczek, P. radiatus L. ). Green gram, Gol-
Indochina. den gram, Mung bean, Oregon pea. 2n=22. India
and Burma. This crop is probably derived from
DERRIS ROBUSTA Benth. 2n= Used as a var. sublobata (Roxb.) Verde. (2n=22), which
shade tree in SE. Asia. grows wild in India and Burma. Sometimes culti-
vated. Spread to S. China, Indochina and Java and
DESMODIUM GYROIDES DC. 2n=20, 22. Trop. later to other countries. It is closely related to
Asia. A shrub used as a green manure. P. mungo*, which probably also derived from var.
sublobata.
INDIGOFERA TEYSMANNn Miq. 2n=32. SE. Asia.
It is a green manure. PHASEOLUS CALCARATUS Roxb. (syn. Vigna u m -
bellate (Thunb. ) Ohwi &Ohashi, Dolichos umbella-
INOCARPUS EDULIS Forst. Tahiti chestnut. 2n=20. tus Thunb. ). Rice bean. 2n=22. Himalaya and India.
From Malaysia to Polynesia where it is cultivated P r i m a r y centre: India. Cultivated in many Asian
for the seeds and as a shade t r e e . countries and elsewhere. Characterized by late
maturing and dehiscent pods.
MELILOTUS SUAVEOLENS Ledeb. (syn. M.
graveolens Bunge). Daghestan sweet clover. 2n=16. PITHECELLOBIUM BIGEMINUM Mart. 2n=
E. Asia and Indochina. Cultivated in the USA. E. and SE. Asia. Cultivated on Java for its edible
Some annuals a r e found in this biennial plant. seeds.
MIMOSA SEPIARIA Benth. 2n= . Trop. Asia. PITHECELLOBIUM JIRINGA Prain. 2n=
Used for hedges (Mansfeld, 1959). Malaysia and Philippines. On Java it is cultivated
for its edible seeds.
MUCUNA ATERRIMA (Piper &Tracy) Holland
(syn. Stizolobium aterrima Piper &Tracy). Mau- PITHECELLOBIUM LOBATUM Benth. 2n=
ritius bean, Bengal bean. 2n=22. Trop Asia. Cul- SE. Asia. Cultivated for its leaves, fruits and
tivated there and elsewhere as a green manure and flowers which a r e eaten as a vegetable.
soil cover.
PSOPHOCARPUS TETRAGONOLOBUS (L. ) D C .
MUCUNA CAPITATA (Roxb.)Wight & A m . 2n= Goa bean, Asparagus bean. Winged bean, Manila
India and Java. Cultivated as a vegetable and for bean. 2n=26. Probably Tropical Asia. Burkill
its s r e d s . (1935) believed that it came from the African coast
bordering the Indian ocean. However as it is cul-
LEGUMINOSAE - MORACEAE 51
tivated in SE. Asia and not in Africa its African Magnoliaceae

origin is doubtful. MICHELIA CHAMPACA L. 2n=38. Cultivated for
perfumery.
PUERARIA PHASEOLOIDES (Roxb. ) Benth. Tropi-
cal kudzu. 2n=22. Malaysia. Used as a cover crop
Malvaceae
and green manure throughout the tropics.
ABUTILON INDICUM (Torner) Sweet. Country
PUERARIA THUNBERGIANA (Sieb. & Zucc.) mallow. Indian abutilon. 2n=(36), 42. Malaysia,
Benth. (syn. P . lobata (Willd.) Ohwi). 2n=24. India and Philippines. Cultivated in India and
Asia and the W. Pacific islands. Cultivated in elsewhere for its fibres and its oily seed. It is a
Central Highlands of New Guinea and in New Cale- weed in the tropics now.
donia for its edible tubers. Formerly it may have
been a staple crop replaced by Ipomoea batatas* GOSSYPIUM ARBOREUM L. Tree cotton. 2n=28,
(Watson, 1968). See also p. 34. genome formula A2A2. Probably originated in
India (p. 68) of Africa (p. 121). Race burmar.icum
SESBANIA AEGYPTIACA Poir. (syn. S. sesban was selected in Indochina. It is an annual form
( L . ) M e r r . ). 2n=12. E. Africa. S. Asia and with a very short day length requirement and which
Australia. In India and Java it is used as a hedge is characterized by the presence of short hairs
and shade plant. resembling wool.
SESBANIA GRANDIFLORA (L. ) Poir. Agati s e s - GOSSYPIUM HIRSUTUM L. Cambodia. 2n=52,

bania. 2n=14, 24. E. India to Australia. Cultivated genome formula (AADD). The origin of this species
in the tropics for its flowers and green pods used is given on p. 154 and 169. Cambodia type developed
in S. Asia as a vegetable. in Cambodia. Introduced into India where it is
cultivated on a large scale. There it was named
TEPHROSIA CANDIDA DC. White tephrosia. Cambodia.
2n=22. Asia. Used as a green manure and cover
crop. THESPESIA POPULNEA (L.). Sol. ex Correa.
Portia tree, Tulip t r e e . 2n=26. The New Guinean
VIGNA HOSEI (Craib) Backer. Sarawak bean. species T. patellifera B o r s s . , T. robusta B o r s s . ,
2n=20. The origin of this cover crop is not clear. T. fissicalyx B o r s s . , T. multibracteata Borss.
It is only known that material cultivated in Malaya belong together with T. populnea to the section
was obtained from Sarawak. It rarely fruits in Thespedia of this genus. This may point to an E.
these regions. Morphologically it is very similar New Guinean origin of T. populnea. It is widely
to V. parkeri Bak. ssp. acutifolia Verde., which distributed in trop, countries as shade t r e e . This
is found in E. Africa. The number of chromosomes wide distribution is due to the capacity of the
of this subspecies is not given, but another sub- seeds to float in sea water for months and remain
species maraguënsis (Taub. ) Verde., (syn. V. alive.
maraguënsis (Taub. ) Harms.) has 2n=22. Ssp.
maraguënsis also grows in E. Africa (Verdcourt, Meliaceae
1970).
LANSIUM DOMESTICUM Jack. Langsat. 2n=72.
With no evidence for the origins of V. hosei Malaysian Archipelago and Indochina. Cultivated
this species is included in Centre 2. Further r e - there and elsewhere for its fruits.
search into its origin is needed. For instance its
karyotype could be compared with those of other
MELIA AZADIRACHTA L. (syn. Melia indica
species. The cause of its almost lack of fruits
Brand., M. japonica Hassk., M. parviflora Moon,
(propagation by cuttings) could be studied. And
Azadirachata indica A. J u s s . ) . Margosa, Nim,
interspecific crosses should be made to study
Neem. 2n=28. Dry region of Irrawadi valley. Cul-
species affinity.
tivated and naturalized throughout India and in
Pakistan, Ceylon, Burma and Malaya. It is a
Lemnaceae minor oil crop in India; the seeds a r e the source
WOLFFIA ARRHIZA Wimm. Khai-nam. 2n= of margosa oil. '
Burma, Laos, Thailand, Bangladesh and India. An
aquatic plant cultivated as a vegetable by Burmese SANDORICUM KOETJAPE (Burm.f. ) Merrill.
and Laotians. Cultivated in N. Thailand in r a i n - Santol. 2n=22, 44. Malaysia and Indochina. Culti-
fed open ponds. It produces a very high protein vated in this area and elsewhere for its fruits.
yield, much higher than the traditional crops in-
cluding soya bean. Moraceae
ALLAEANTHUS LUZONICUS F. Vill. 2n=
Liliaceae
This t r e e is cultivated in Philippines.
TAETSIA FRUTICOSA (L. ) Merr. 2n= . Paci-
fic islands and Malaya. The leaves were used for ARTOCARPUS ALTILIS (Park. ) Fosberg (syn.
cloths. Cultivated in Samoa for this purpose. A. communis Forst.) Breadfruit. 2n=54, 56, c. 81.
Wild in the forests of Malayan Peninsula and those
of the Moluccas. Secondary gene centre on the
islands of Oceania.
Diverse forms are found in Philippines. They G* ß •

a r e products of hybridization between the bread-
fruit and A. blaneoi (Elm. ) M e r r . , which grows
in that country. Similarly variants are observed
in Micronesia being products of introgression / *— * ^ S H £ '
between breadfruit and the wild A. mariannensis

Trécul.
The introgressed characters of this wild 3
species are seeded fruits, entire leaves and (*-
reddish hairs on veins (Fosberg, 1960a; Coenen &
Barrau, 1961). 'H
ARTOCARPUS CAMANSI Blanco. Kamansi.
2n= . Philippines. Cultivated near Manila for
v J>
its fruits.
The wild Musa balbisiana (- -) and M. acuminata ( ) types
ARTOCARPUS CHAMPEDEN (Lour. ) Spreng. (Simmonds, 1962)
Champedak. 2n= . Malaya. Selected forms are
cultivated in SE. Asia.
M. acuminata is a polymorphous species. The
ARTOCARPUS LAKOOCHA Roxb. 2n= . India primary centre of origin is the Malayan region,
and Malaysia. Cultivated in the tropics for its but at present the greatest diversity is known in
fruits. New Guinea (Simmonds, 1964).
The cultivars of the AA-group a r e found
ARTOCARPUS RIGIDUS Blume (syn. A. dimor- throughout the tropics. In Malaya region the p r i -
phophylla Miq. ). Monkey jack. 2n= . Malaysian mary centre of diversity is observed. A secondary
Archipelago. Cultivated for its fruits. is found in E. Africa (p. 125). The main cultivars
as Gros Michel, Cavendish subgroup and R e d /
Musaceae Green Red belong to the AAA-group. The Dwarf
Cavendish cultivars of the Cavendish group grown
MUSA (Eumusa) edible cultivars. Banana. 2n=22,
e.g. on Samoa and Canary Islands originated from
33, 44. The genus Musa is divided into four main
Indo-China. These cultivars have often been r e -
sections of which Eumusa includes the important
ferred to as M. cavendishii Lambert or M. sinen-
cultivars. The edible cultivars are improved
sis Sweet ex Saget. (Simmonds, 1964).
types of M. acuminata Colla (2n=22, genome for-
mula AA), triploids of this species and diploid, The first M. x acuminata x balbisiana hybrid
triploid and tetraploid hybrids of this species and group is the diploid AB group. This small group
M. balbisiana Colla (2n=22, genome formula BB) is of S. Indian origin (Simmonds, 1964) (p. 68).
(p. 69). The latter types have often been named The second hybrid group is the triploid AAB
M. x sapientum L. and M. x paradisiaca L. group. Its major centre of origin lies in India
^
0 F
A-
/ABB ABBB 7\;'-
\A /AAB—\
\ t \ ÂABV
^v~^—-~
!^l|ëSF^.
\ *% y
AAor~nf' \" \AWA N

AB J
later 3x1
° J7
Distribution of wild bananas ( )and cultivation in Africa (-• -). Origin and movements of Eumusa groups (AA-ABBB) and of Aus-
tralimusa series (open arrow) (Simmonds, 1962)
MORACEAE-PALMAE 53
(p. 68), while a clone (Malo maoll) may have EUGENIA JAMBOS L. (syn. Syzygium jambos L.)
arisen in Philippines (Simmonds, 1964). Alston). Rose apple. 2n=28, 33, c. 42, 44, 46,
The third hybrid group is the triploid AAB c. 54. This tree has been cultivated for a long
group. S. India is a major centre of origin. It is time in Indo-Malaysia. Its exact centre of origin
quite likely that a second centre is found in Philip- is not known.
pines.
The fourth hybrid group, consisting of one EUGENIA JAVANICA Lam. (syn. Syzygium s a m a -
clone, is the tetraploid ABBB group. Its centre rangense (Bl.) Merr. .&P e r r y . 2n=33, 42, 44, 45,
of origin lies very probably in Indochina (Simmonds, 66, 88, 110. Malaysia to India. Much cultivated
1964). in Java.
MUSA* cultivars'of the ABB-group. 2n=33. Most EUGENIA MALACCENSIS L. (syn. Syzygium
ABB-cultivars originated in S. India (p. 68). malaccensis (L.) Merr. &Perry). Pomerac,
However, it is possible that after the cultivated Malay apple. 2n=22. Malay. Some varieties are
M. acuminata (AA) reached Philippines, hybrids known.
arose with M. balbisiana*.
MELALEUCA QUINQUENERVIA L.f. (syn. M.
MUSA* BALBISIANA* leucadendra L. ) Cajéput t r e e . 2n= . Australia
to Burma. Planted in forestry projects in Philip-
MUSA* TEXTILIS Nee. Abaca, Manilla hemp. pines, Hawaii and elsewhere. Also planted for the
2n=20. Philippines. A tall, perennial. Cultivated purpose of drying up swamps, and as an ornamental.
there and elsewhere in the tropics for its fibre.
Canton fibre is obtained from a natural completely PIMENTA ACRIS Kostel. 2n=22. Indonesia. Culti-
sterile hybrid (2n=21) of M. textilis x M. balbi- vated there for its oil which is distilled from the
siana*. leaves (Purseglove, 1968).
Myristicaceae RHODOMYRTUS TORMENTOSA Wight. Downy

rose myrtle. 2n= . India and Malaysia. A
MYRISTICA ARGENTEA Warb. Papuan nutmeg. shrub cultivated in the (sub)tropics for its fruits.
2n= . New Guinea where it is also occasionally
cultivated (Flach and Cruickshank, 1969). Nyctaginaceae
MYRISTICA FRAGRANS Houtt. Banda-nutmeg. PISONIA ALBA Span. Maluko, Lettuce t r e e .
2n=42, 44. Centre of origin: the Moluccas. It is 2n= . Malaya. Wild t r e e is called P . sylvestris
not found there wild. From here it spread through- Teijsm. &Binn. (2n= ) (syn. P. grandis R. Br. ).
out the tropics (Flach & Cruickshank, 1969). Cultivated for its leaves which a r e used as a vege-
table
Myrtaceae
Palmae
EUGENIA AQUEA Burm.f. (syn. Syzygium aqueum
(Burm.f. ). Alston. Water rose apple. 2n= ARECA CATECHU M e r r . Betelnut palm. 2n=32.
Bangla Desh, Burma, Ceylon, Sumatra and Mo- Trop. Asia. Cultivated for its nuts.
luccas. It is also cultivated.
ARENGA PINNATA (Wurmb. ) Merr. Sugar palm.
EUGENIA CARYOPHYLLUS (Sprengel) Bullock & 2n=26, 32. P r i m a r y centre: the Indonesian-Hindu-
Harrison (Syzygium aromaticum (L. ). Merr. & stani gene centre (p. 69). Secondary gene centre:
P e r r y ) . Clove t r e e . 2n= . The wild clove t r e e , possibly India.
sometimes named E. obtusifolia Reinwardt (Caryo-
phyllus sylvestris T. &B. ), grows in many islands BORASSUS FLABELLIFER*
of the Moluccas and in New Guinea. Is has bigger
leaves and flower buds and is less aromatic than CALAMUS CAESIUS Blume. 2n= . .Malaya,
the cultivated t r e e . Cultivated for a long time. Borneo and Sumatra. Cultivated for its stems.
Some variability exists in and outside its
centre of origin. COCOS NUCIFERA L. Coconut palm. 2n=32. Cen-
t r e of diversity SE. Asia, Indonesia and W. P a c i -
EUGENIA FORMOSA Wall. (syn. Syzygium mappa- fic islands. It is not known whether it comes
ceum (Korth. ) Mansf. ). 2n= . Trop. Asia. from S. America or SE. Asia, Indonesia and W.
This tree is cultivated in Cochin-China for its Pacific islands. From its native area it spread
fruits. to all tropical countries. Dennis and Gunn (1971)
suggested that sea currents may have helped in
EUGENIA JAMBOLANA Lam. (syn. Syzygium the dispersal over small distances (from island
cumini (L. ). Skeels, Eugenia obtusifolia Roxb., group to island group), but that man distributed
E. cumini (L.) Druce). Java plum, Jambolan the coconut over the world.
plum. 2n=33, 44, 46, 55. India to Malaysia, China It is possible that in Centre 4 a secondary
and N. Australia. It is cultivated there and e l s e - centre arose (p. 69).
where. In India a large fruited type is cultivated
(Mansfeld, 1959). COELOCOCCUS ARMICARUM Warb. Polynesian
ivory-nut palm. 2n= . Carolina Islands. Cul-
tivated in Philippines for its ivory-like nuts. gether with the betelnut (Areca catechu*).
METROXYLON SAGU Rottb. Sago palm. 2n= PIPER CUBEBA L.f. Cubeb, Cubebe, Tailed
Malaya, Moluccas and New Guinea. Sago is ob- pepper. 2n= . Cultivated there and neighbouring
tained from the marrow of the stem. countries.
This species is occasionally split in M. sagu
- the wild type and M. rumphii (Willd. ) Mart. PIPER METHYSTICUM Forst. Kava pepper.
(2n= ) - the cultigen. 2n= . Polynesia. Cultivated there. The roots
and rhizomes a r e used to prepare a non-alcoholic
NYPA FRUTICANS Wurmb. (syn. Nipa fruticana beverage. In small quantities it is a stimulant, in
Thunb. ). Nipa palm. 2n=16, SE. Asia upto Austra- large quantities a narcotic.
lia. Cultivated on Sumatra for its leaves and for
wine production. Introduced to the mangrove area PIPER RETROFRACTUM Vahl (syn. P. officina-
of S. Nigeria where it has run wild (Zeven, 1973). r u m D C ) . Javanese long pepper. 2n= . Malay-
sia. It resembles P. longum*. Cultivated for its
PRITCHARDIA GAUDICHAUDII H. Wendl. 2n= spike which is used as a spice.
Sandwich Islands. Cultivated there for its leaves
which are used for thatching. PIPER SAIGONENSE C. DC. Lolo. 2n=
Indochina. Cultivated there occasionally. Closely
PRITCHARDIA PACIFICA Seem &Wendl. 2n=36. related to P . lohot C. DC. which comes from the
Fiji and Samoa. Cultivated for its leaves which region of Tonkin.
a r e used for thatching.
Polygonaceae
SALACCA EDULIS Reinw. 2n= . Malaysian
Archipelago. Cultivated on Java for its edible POLYGONUM ODORATUM Lour. 2n= . Indo-
fruits. China. Cultivated as a potherb in Vietnam.
Pandaceae Rosaceae
PANDANUS BROSIMAS M e r r . &P e r r y . 2n= RUBUS ALBESCENS*

Cultivated in the highlands of New Guinea for its
seeds, which have a pleasant flavour and a r e rich RUBUS ROSAEFOLIUS Smith. Cape bramble,
in oil (Purseglove, 1972). Mauritius raspberry. 2n= . Tropical Asia.
Introduced in other continents. Cultivated. It is
considered as a parent of R. probus Bailey,
PANDANUS ODORUS Ridl. 2n= . Cultivated by
Queensland raspberry, a shrub from Australia.
Malays for its fragrant leaves (Purseglove, 1972). The other parent is R. ellipticus Smith, the Yellow
Himalayan raspberry from E. India.
PANDANUS SPURIUS Miq. (syn. P. moschatus
seu laevis Rumph., R. moschatus Rumph. ex Miq.
Rubiaceae
P. tectorius Soland. var. moschatus (Rumph. ex
Miq. ) M e r r . , P. laevis Lour., R. odoratissimus MITRAGYNA SPECIOSA Korth. 2n= . Malaya
L.f., P . inermis Roxb. ). Thatch screw pine, and Lower Siam. Cultivated as a substitute of opium.
Putat, Pudak. 2n=c. 51, 54, 60. Cultivated in SE.
Asia to the extremes of Polynesia for its leaves MORINDA TRIFOLIA L. Indian mulberry. 2n=
for thatch and for its fruits. The cultivar is one Indonesia and Malaya. Cultivated on Java as a dye
clone probably originated as a bud sport on a s t a - crop.
minate plant of some wild species of the section
Pandanus. Maybe this mutation occurred on a OLDENLANDIA UMBELLATA L. Indian madder.
specimen of P. spurius some millenia ago (St. 2n=36. Trop. Asia. Cultivated as a dye plant.
John, 1965). UNCARIA GAMBIR (Hunt. ) Roxb. Gambier.
2n= . Malaya. Formerly cultivated in SE. Asia.
PANDANUS WHITMEEANUS Martelli. Paogo. Its leaves and young branches contain a tannin.
2n= . Cultivated in New Caledonia, New Hebri-
des, and elsewhere. On Futuna it is used only for Rutaceae
personal adornment. The fruit oil is used to p e r -
fume coconut oil (St. John and Smith, 1971). AEGLE MARMELOS (L. ) Corr. Indian bael, Ben-
gal fruit. 2n=18, (36). Cultivated in SE. Asia and
Pentaphragmaceae some other tropical countries for its fruits which
a r e used medicinally.
PENTAPHRAGMA BEGONIAEFOLIUM Wall.
2n= . A fleshy herb cultivated as a vegetable CITRUS AURANTIFOLIA (Christm. ) Swing. Lime.
in Malaya (Terra, 1967). 2n=18, (27). Probably the Indonesian Archipelago
or N. India. Wild t r e e s a r e reported to grow in
Piperaceae N. India. Spread throughout the tropics. The cul-
PIPER BETLE L. Betel peper, Betle vine, Betal, tivar Tahiti is triploid. Interspecific hybrids
Sirih. 2n=32, 64, (78). C. a n d E . Malaysia. Cul- have been obtained. Mandarin lime is probably a
tivated in the tropics. The leaves a r e chewed t o - hybrid of this species with C. reticulata* and
sweet lime with C. medica*, while limequat is a
PALMAE - SAPOTACEAE 55
hybrid with Fortunella margarita*. a natural hybrid of the same parents C. reticulata*.
The nakoor lime (named C. nakoor) is a complex Its origin is in Siam. Tangor has been described
natural hybrid of this species and some Papeda as C. nobilis Lour.
group parentage.
CITRUS SINENSIS (L. ) Osbeck (syn. C. aurantium
CITRUS AURANTIUM L. Sour orange, Seville L. var. sinensis L. ). Sweet orange. 2n=18, (27,
orange, Bigarade. 2n=18. Probably SE. Asia or 36). Probably S. China or Cochin-China. Unknown
Cochin China. Unknown wild. Spread throughout wild. Secondary centres: Israel and Spain (p. 105).
the (sub)tropics. In some a r e a s it has run wild. It was already mentioned in Chinese sources dated
The ssp. bergamia (Risso &Poit. )Wight & Arn., 2200 BC.
Bergamot, (2n=18) is especially cultivated in Ca- Scora and Malik (1970) showed that this species is
labria, S. Italy for the production of bergamot oil not a mutant of C. aurantium* or a hybrid of this
(p. 105). Crosses with C. sinensis* (Sweet latter species and C. reticulata* which had been
Orange) gave Bitter Sweet Orange. The var. suggested. It shows close affinities to C. reticulata*.
myrtifolia Kergawl., Myrtle leaved Orange is a It is widely distributed in the (sub)tropics.
bud mutant. Its fruits Chinottos a r e candied in There a r e many cultivars. Citrange is a hybrid
Italy and S. France. product with Poncirus trifoliata*, while chironja
is a spontaneous hybrid with C. paradiso*. It origi-
CITRUS GRANDIS (L.) Osbeck (syn. C. decuma- nated in Puerto Rico. Owing to apomixis it breeds
nus L. 2n=18, 21; C. maxima (Burm. ) Merr. 2n=18, true.
36), Pummelo, Shaddock. 2n=18, 36. Probably
SE. Asia. P r i m a r y centre of diversity: SE. Asia. MURRAYA EXOTICA L. Limonia. 2n=18. Trop.
Spread to China, India and Iran and later to other Asia. Used for hedges.
tropical countries (by captain Shaddock to Barba-
dos in the 17th Century). Unknown wild. The best MURRAYA PANICULATA (L. ) Jacq. Cosmetic
fruits come from Thailand where the plants a r e barktree, Orange jasmine. 2n=18. SE. Asia. Cul-
cultivated on ridges surrounded by brackish water. tivated in the tropics as an ornamental and for
hedges. The wood (Satinwood) is used in Java to
CITRUS HYSTRIX DC. Mauritius papeda. 2n= make cutlery.
Philippines and Burma to Malaya. A small type
cultivated for its fruits. Santalaceae
SANTALUM ALBUM L. (syn. Sirium myrtifolium
CITRUS LIMETTA Risso. Sweet lemon. 2n=18.
L. ). Sandal wood. 2n=10. E. India to Malaysia.
Trop. Asia. Small t r e e cultivated in some coun-
Cultivated there and elsewhere for its scented
tries.
wood.
CITRUS LIMON (L. ) Burm.f. Lemon. 2n=18, 36.
Centre of origin somewhere in SE. Asia. The Sapindaceae
area east of Himalayas in N. Burma and S. China ERIOGLOSSUM RUBIGINOSUM (Roxb. ) Blume
has been suggested. Unknown wild. A secondary (syn. E. edule Bl. ). 2n= . Trop. Asia to New
centre: the Mediterranean Region (p. 105). Scora Guinea and Australia. Asmall tree cultivated in
and Malik (1970) suggested'that this species might Indonesia and elsewhere.
be a stabilized hybrid of C. medica* - C. auranti-
folia* assemblage. Cultivated in several ( s u b t r o - NEPHELIUM LAPACCEUM L. Rambutan. 2n=22.
pical regions. Rough lemon is probably a hybrid Malaysian Archipelago. Cultivated for its delicious
with C. medica*. It became naturalized in Rho- fruits. Many varieties have been developed.
desia.
NEPHELIUM MUTABILE Blume. Pulasan.
CITRUS MITIS Blanco. Calamondin. 2n=18. Philip- 2n= . Malaysia. Cultivated in SE. Asia and in
pines. A tree occasionally cultivated in (sub)tropics. other countries.
Hybrids of this species have been produced, so
a r e Calarin and Calashu hybrids with C. reticulata* POMETIA PINNATA Forst. Matoa, Taun.
(Satsuma). 2n= . Malaysia, Indonesia, Papua and Pacific
islands. A forest tree used for timber and for its
CITRUS RETICULATA Blanco (C. nobilis Andr. fruits. Cultivated for its edible fruits. On W. Irian
non Lour. ). Manderin, Tangerine. 2n=18. P r o b a - alongside the banks of the Sentani lake.
bly Philippines, or Cochin China. Unknown wild. This culture will probably be replaced by that of
Secondary centre arose in Japan (p. 39). Minessy higher yielding exotic fruit trees (Rappard, 1961).
et al. (1970) found close relationship with C. sinen-
sis*. "Balady Blood". Its relationship with C. SAPINDUS RARAK DC. 2n= . Cochin-China
paradisi* "Duncan" and "March" is moderate and and Malaysia. Planted in Java, India and e l s e -
with C. grandis* distant. Var. austera Swing is where for its fruits.
the sour manderin. It probably includes the Rang-
pur lime (Purseglove, 1968). Sapotaceae
Hybrids with other species have been made.
MANILKARA ELENGI (L. ) Chev. 2n= . Origin
For instance Oranguma is an artificial hybrid of
Satsuma x C. sinensis* (Orange), while Tangor is uncertain (Uphof, 1968). Cultivated in the Malay-
sian Archipelago. ALPINA GALANGA (L. ) Willd. Langwas, Greater

galangal. 2n=48. Trop. Asia. Cultivated for its
PALAQUIUM GUTTA (Hook. ) Burck. Gutta percha. rhizomes. It is a common village plant. Several
2n=24. Malaysia. It is tapped for its latex. In varieties have been observed.
general the tree is first felled.
ALPINIA MALACCENSIS (Burm.f. )Rose. 2n=48.
PAYENA LEERII (Teysm. &Binn. ) Kurz. 2n= Malaysian Archipelago and E. India. This perennial
Burma and W. Malaysia. Cultivated on Java as a herb is cultivated.
source of gutta percha.
AMOMUM CARDAMOMUM Willd. Cardamon.
Saururaceae 2n= . Malaysia. Cultivated there.
HOUTTUYNIA CORDATA Thunb. 2n=56, 96, AMOMUM KEPULAGA Sprague &Burk. Round
c. 96, 100-104. Indochina and China. Cultivated cardamon. 2n= . Cultivated in Malaysia and
in Vietnam for salad and as a medicinal crop. Java.
Solanaceae
AMOMUM KRERVANH P i e r r e . Krervanh. 2n=
LYCINUM CHINENSE Mill. Chinese wolfberry. Cambodia. Cultivated in Indochina.
2n= . E. Asia. Cultivated in Java as a vegeta-
ble. AMOMUM MAXIMUM Roxb. Java cardamon.
2n= . Malaysia. Cultivated in Java.
SOLANUM UPORO Dunal. 2n= . Polynesia.
Cultivated in Fiji for its fruits. BOESENBERGIA PANDURATA (syn. Gastrochilus
pandurata Ridl. ). 2n= . Malaya and Java. Cul-
Stilagninaceae tivated over wide area for its rhizome.
ANTIDESMA BUNIUS (L.) Spreng. Bignay, China CURCUMA HEYNEANA Valeton. 2n= . Java.
laurel. 2n= . India to Australia. Cultivated in
The rhizomes are a source of an arrowroot.
Malaysia and elsewhere for its fruits (Purseglove,
1968).
CURCUMA PIERREANA Gagn. 2n= Malaya.
Cultivated in Annam.
Styraceae
STYRAX BENZOIN Dryander. 2n= . Malaysian CURCUMA XANTHORRHIZA Roxb. 2n=
Archipelago. Planted in Sumatra. Amboina. Occasionally cultivated in Java and
Malaya.
Taccaceae
KAEMPFERIA GALANGA L. 2n=22, 54. Trop.
TACCA PINNATIFIDA Forst, (syn. T. involucrata
Asia. Widely cultivated for its rhizomes.
Schum. &Thonn., T. leontopataloides (L. ) Kuntze).
2n=30. SE. Asia (Massai and Barrau, 1956).
KAEMPFERIA ROTUNDA L. 2n=33, 54. Trop.
Spread to the South Sea islands, Asia and Africa.
Asia. Cultivated for its rhizomes.
Cultivated for its starchy tubers.
PHAEOMERIA MAGNIFICA Schum. (syn. Alpinia
Umbelliferae magnifica R o s e . , A. speciosa D. Dietr., Amomum
LIGUSTICUM MONNIERI Calest. (syn. Selinum magnificum Benth. ). 2n= . Malaya. Cultivated
monnieri L. ). 2n= . E. Europe, Siberia, China there.
and Vietnam. Occasionally cultivated in N. Vietnam.
ZINGIBER CASSUMUNAR Roxb. Cassumunar
OENANTHE JAVANICA DC. (syn. O. stolonifera ginger. 2n=22. Cultivated in Cochinchina and
Wall. ). Oriental celery, Water dropwort, Batja- Malaya. In Malaya as a village medicinal crop.
rongi. 2n=20. From Indochina to Malaya, Philip-
pines, China, Korea, Japan and Australia. Culti- ZINGIBER ZERUMBET (L.) Smith. Zerumbet
vated in Indochina, Japan, China (Kihara, 1969) ginger. 2n=22. Trop. Asia. Cultivated in Cochin-
and in Java. A leafy vegetable that often occurs as china, Cambodia and elsewhere.
a weed.
Urticaceae
LAPORTEA DECUMANA Wedd. 2n= . The
Moluccas. Cultivated as a medicinal plant.
Zingiberaceae
ALPINIA CONCHIGERA Griff, (syn. Languas con-
chigera Burk. ). 2n= . Malaya. It is a common
village plant there.
3 Australian Centre
The Australian Centre was not described by Vavilov, but it was marked out by Zhukovsky (1970) because
of the domestication of several plant species to important crops, or the use of wild species as breeding
parents. The main crops derived from this centre a r e Eucalyptus species. Wild species useful for t o -
bacco breeding a r e Nicotiana debneyi and N. goodspeedii. It is a secondary centre of diversity for Trifo-
lium subterraneum.
Agavaceae ACACIA DEALBATA Link. Silver wattle. 2n=26.

PHORMIUM TENAX J . R . et G. Forst. New Zea- SE. Australia and Tasmania. Cultivated as an o r -
land flax, New Zealand hemp, Harakaka lily, F o r - namental, for its timber and as soil stabilizer. It
mio. 2n=32. New Zealand. Cultivated there. Intro- is the familiar florist's mimosa.
duced into S. America and other countries. The
only other species of this genus Ph. colensoi Hook., ACACIA LONGIFOLIA (Andrews) Willd. (Syn. A.
mountain flax (2n=32) produces a weak fibre. It cibaria F . V . M u e l l . ) . 2n=26. New South Wales,
might be used as a breeding parent. Australia. Cultivated as an ornamental and as a
stabilizer of coastal dunes in Europe.
Casuarinaceae
ACACIA MEARNSn De Wild. Black wattle. 2n=26.
CASUARINA EQUISETIFOLIA Forst. Swamp oak, Cultivated in several countries mainly for its tannin
Bull oak, Polynesian iron wood, Horsetail t r e e . and as an ornamental. Sometimes the names A.
2n=18. It is often cultivated as a soil stabilizer. decurrens (Wendl.) Willd. or A. mollissima Willd.
are wrongly used for black wattle.
Chenopodiaceae
ATRIPLEX SEMIBACCATA R. Br. Australian s a l t - ACACIA PYCNANTHA Benth. Golden wattle. 2n=
bush, Berry saltbush. 2n=18. Australia. Cultivated S. Australia and Victoria, Australia. Cultivated
as a fodder crop on the saline soils of California for tannin and as an ornamental.
and Arizona, USA.
LUPINUS COSENTINI Guss. (syn. L. varius L. ssp.
varius Franco &P.Silva). Western Australia blue
Gramineae
lupin, Sandplain lupin. 2n=32. Along the coast of
ORYZA AUSTRALIENSIS Domin. 2n=24, genome Marocco and scattered in W. Mediterranean region.
formula EE. Australia. All r e s e a r c h into the affini- Introduced into W. Australia about 1850 to be used
ty of the species to other Oryza species uses plants as a source of flour. Cultivated for summer sheep
derived from one collection (Chang, 1970). feed and soil improvement. It is naturalized now
widely (Gladstones, 1970).
Leguminosae
ACACIA CYANOPHYLLA Lindl. 2n=26. Australia. PHASEOLUS LATHYROIDES L. Phasemy bean.
Cultivated as an ornamental and in Europe to s t a - 2n=22. Queensland, Australia. Used in E. Africa
bilize coastal dunes. in pastures ( W h y t e e t a l . , 1953).
AUSTRALIAN CENTRE 58
TRIFOLIUM SUBTERRANEUM L. Sub clover.

2n=16. P r i m a r y centre in the Mediterranean r e -
gion (p. 103). Secondary centre: Australia.
Malvaceae
GOSSYPIUM AUSTRALE V. Muell. 2n=26, genome
formula C3C3. N. Australia.
GOSSYPIUM ROBINSONII V. Muell. 2n=26, genome

formula C2C2. W. Australia
GOSSYPIUM STURTH V. Muell. 2n=26, genome

formula C i C j . C. and S. Australia.
Musaceae
MUSA (Australimusa). Fe'i banana. 2n=20. The
fe'i banana originated from one or more wild Aus-
tralimusa species in New Guinea-Solomon Islands
area. Probably carried by man in an easternly d i -
rection. Cultivated especially in Tahiti, where
many bunches are harvested from semi-wild plants.
Some clones have been described as M. fehi Bert,
ex Vieill., M. aiori Sagot, M. seemanii F.V.
Muell. and M. troglodytarum L. (Simmonds, 1964). Gossypium s t u r t i i
.
• ^ ^ ^
s 0/ » 0 V
\5^-
fa
V\. \ * • •... ••••/

Cj
«.
Wild A u s t r a l i m u s a ( ) and F e ' i bananas ( ) (Simmonds, 1962)
Myrtaceae EUCALYPTUS BOTRYOIDES Smith. Blue gum,

EUCALYPTUS ALBA Reinw. ex Blume. 2n=22. Bangalay eucalyptus, Bastard mahogany. 2n=22.
Timor and Flores and to the south of New Guinea. Coastal areas of SW. Australia. Cultivated in
Cultivated in Brazil. The wood is reddish-brown. Algeria and Zaire. E. trabutii Vilm. (2n=22), is
The cork contains much tanning material. a hybrid of E. botryoides $ and E. camaldulensis*
& a r i s e n in Italy.
EUCALYPTUS AMYGDALINA Labill. (syn. E.
salicifolia Cav. ). Willowleaf eucalyptus, Pepper- EUCALYPTUS BROCKWAYI Gardn. 2n=22. S.
mint t r e e . 2n=22. Tasmania. Cultivated in Chile, Australia. Its area of distribution is limited. Cul-
Zaire and W. Georgia, USSR. Closely related to tivated in N. Africa. Extremely drought resistant.
E. regnans*.
EUCALYPTUS CAMALDULENSIS Dehn. Longbeak
EUCALYPTUS ASTRINGENS Maiden. Brown mallet. eucalyptus, Australian kino, Red gum. 2n=22.
2n=22. SW. Australia. Cultivated in Marocco, S. Australia, excluding Tasmania. It is cultivated
Africa and Cyprus. The bark used for the tanning almost in all countries that grow Eucalyptus. Secon-
industry. It is very drought resistant. dary centres: the Mediterranean region (p. 104),
Brazil (p. 115) and Argentine (p. 115). In cultivation
AGAVACEAE - MYRTACEAE 59
many spontaneous hybrids have arisen. E. trabutii EUCALYPTUS GLAUCESCENS Maiden & Blakely.
Vilm. (2n=22) is a hybrid of E. botryoides Ç and 2n= . Mountains of SE. Australia. Its distribution
E. eamaldalensis (f . A new form developed in is very limited. Used for crossing with species of
Israel (p. 104). poor hardiness.
EUCALYPTUS CINERA F. Muell. 2n=22. S. areas EUCALYPTUS GLOBULUS Labill. Fever t r e e ,

of New South Wales, Australia. Used as an orna- Blue gum. 2n=20, 22, 28. SE. Tasmania. Culti-
mental. It is a valuable source for breeding cold vated. Secondary centre: the Mediterranean region.
resistant forms of Eucalyptus. Used for wood and oil. Spontaneous hybrids are
known in Tasmania under cultivation.
EUCALYPTUS CITRIODORA Hook. Spotted gum,
Lemon scented gum. 2n=(20), 22, (28). The N. EUCALYPTUS GOMPHOCEPHALA A.DC. 2n=22.
coast of Queensland, Australia. Cultivated in many SW. coasts of W. Australia. Cultivated in countries
(sub)tropical countries for an essential oil rich in of the Mediterranean region. Africa esp. Cameroon,
citronellal. Hawaii and New Zealand. It has the heaviest and
strongest wood among all Eucalyptus species. In
EUCALYPTUS CLADOCALYX F. Muell. (syn. Algeria some spontaneous hybrids are known.
E. corynocalyx F . Muell). Sugar gum. 2n=22.
S. Australia. Cultivated in Australia, the Medi- EUCALYPTUS GRANDIS Hill, ex Maiden. 2n=
terranean region and in some African countries. Coast areas of the Northern part of New South
The wood is of excellent technical value. Wales and SE. Queensland up to 650 m. Cultivated
in Cameroon, Nigeria and Madagascar. It is thought
EUCALYPTUS COCCIFERA Hook. f. 2n= . T a s - t h a t E . ' s a l i g n a ' o r E. 'saligna/grandis' a r e African
mania. Because of its hardiness it is used for strains developed after introduction of'Queensland
breeding types for W. Georgia, USSR. material (Larsen &Cromer, 1970).
EUCALYPTUS CREBRA F. Muell. 2n= . Queens- EUCALYPTUS GUNNII Hook.f. 2n=22. Cultivated
land, reaching New South Wales, Australia. Culti- in USSR, Great Britain, Japan and Hawaii. Used
vated in several countries of Africa, India and for industry and breeding on the Caucasus coasts
Argentine. Some spontaneous hybrids a r e known. of the Black Sea.
EUCALYPTUS CYPELLOCARPA L. Johnst. EUCALYPTUS LEUCOXYLON F. Muell. (syn.

(E. goniocalyxpl. anct. ). 2n=22. SW. Australia E. conoidea Benth. ). White ironbark, White gum.
attaining 900-1200 m altitude. Cultivated in the 2n=22. Central areas of Victoria and South Austra-
Mediterranean region, S. America and on Hawaii. lia. In the latter a r e a it is r a r e . Cultivated in the
Mediterranean region, S. America esp. Argentine
EUCALYPTUS DALRYMPLEANA Maiden. Moun- and Cyprus. Used for its wood and oil. Some geo-
tain gum. 2n=22. SE. Australia, attaining 1350 m graphical races and spontaneous hybrids have been
altitude and in C. Tasmania attaining 900 m alti- described.
tude. Cultivated in USSR on the coasts of the Black
Sea in the Caucasus. It is a promising economie EUCALYPTUS MACARTHURI Dean & Maiden.
species on Hawaii and in China. It is considered 2n=22. Central New South Wales, Australia. Cul-
to be of hybrid origin. Natural and artificial hy- tivated in Africa esp. Zaire, Hawaii, New Zealand,
brids are known. It can be used in breeding for S. France and W. Georgia, USSR. It produces an
better types. essential oil. Some spontaneous hybrids are known.
In the USSR many (poly)hybrids have been produced.
EUCALYPTUS DELEGATENSIS R . T . Baker, (syn.
E. gigantea Hook.f. ). Alpine ash, Woolybutt, Red EUCALYPTUS MACULATA Hook.f. (syn. E. v a r i e -
mountain ash, White top stringbark. 2n= . The gata F . Muell. ). Spotted gum. 2n=22. Coastal
mountains of SE. Australia up to 1350 m and in areas of SE. Queensland, New South Wales and
Tasmania up to 900 m. Cultivated in New Zealand, E. Victoria. Cultivated in Africa esp. Cameroon,
Hawaii and W. Georgia, USSR. Used for cultivation Zaire, Kenya and Madagascar; the Mediterranean
and as a breeding parent in the USSR. region esp. Spain, France; Chile and Uruguay.
The wood is very valuable.
EUCALYPTUS DIVERSICOLOR F. Muell. K a r r i .
2n=22. Coasts of SE. Australia. Cultivated in EUCALYPTUS MAIDENII F. Muell. Maiden's gum.
countries of the Mediterranean region, in Africa Spotted blue gum. 2n=22. Mountains of SE. Austra-
and New Zealand. It is one of the most valuable lia. Cultivated in Africa esp. Cameroon, Congo
economic species. and Kenya; the Mediterranean region esp. Italy and
Spain; Brazil and New Zealand. Its wood is valuable,
EUCALYPTUS EUGENIOIDES Sieb. (syn. E . s c a b r a containing essential oil. Some spontaneous and
Dum-Cours.). White stringbark, Pink blackbutt. artificial hybrids have been reported.
2n= . Coasts areas of SE. Australia. Cultivated
in S. Africa, Kenya, India and Hawaii. The wood EUCALYPTUS MELLIODORA A. Cunn. Yellow box.
is used in industry. Some natural hybrids a r e known. 2n=22. Australia. Cultivated in the Mediterranean
region, in Africa esp. Zaire and E r i t r e a . Used for
its wood and as an ornamental t r e e . It is extremely
AUSTRALIAN CENTRE 60
melliferous. There are geographical races and and Brazil. After E. globulus* the most widely d i s -
spontaneous hybrids known. tributed species in cultivation. The wood is ex-
tremely valuable. This is the most rapidly growing
EUCALYPTUS MICROCORYS F. Muell. Fallow species in the genus.
wood. 2n= . Coastal areas of the N. part of New
South Wales and SE. Queensland. Cultivated in the EUCALYPTUS SIDEROXYLON A. Cunn. ex Benth.
Mediterranean region and Africa esp. Zaire and Red ironbark. 2n=22. The W. slopes of New South
E r i t r e a . Used for its wood. Some spontaneous Wales upland and in the N. part of Central Victoria,
hybrids are known. Australia. Cultivated in Africa esp. Cameroon,
Kenya, Zaire and Rhodesia; the Mediterranean
EUCALYPTUS NIPHOPHILA Maiden & Blakely. region, esp. Spain, Portugal, Algeria, Morocco,
2n= . Alpine zone of SE. Australia, up to 2000 m. Cyprus and Israel; Japan, USA and New Zealand.
It tolerates -24°C and hence is of great importance Its wood is economically very valuable. It contains
for hybridization with valuable economic species. essential oil. Some spontaneous hybrids a r e known.
EUCALYPTUS PANICULATA Sm. (syn. E. fergu- EUCALYPTUS TERETICORNIS Smith, (syn.

s o n i R . T . Baker). Grey ironbark. 2n=22. The E. subalatum Cunningh. ) Red gum, Flooded gum,
coasts of New South Wales, Australia. Cultivated Grey gum, Blue gum. 2n=22. Almost the whole
in Africa esp. Kenya and Tripoli; the Mediterranean coast of E. Australia. Cultivated almost in all the
region, esp. Spain and Tripolitania; S. America, countries of the world where Eucalyptus is grown.
esp. Paraguay and Uruguay, and India. The wood The wood is very valuable. In the USSR interspeci-
is especially strong, heavy and durable. fic hybrids a r e produced. The strains 'C' of Zanzi-
bar and 'Mysore Hybrid' of India belong to this
EUCALYPTUS PAUCIFLORA Sieb, ex Spreng. species (Larsen & Cromer, 1970).
2n= . Sub-alpine zone of E. Victoria and the
mountains of New South Wales and Tasmania, up EUCALYPTUS VIMINALIS Labill. (syn. E. manni-
to 1650 m. Cultivated in England, France, Japan fera Cunning, E. persicifolia Lodd. ). Ribbon
and W. Georgia, USSR. On the fringe of its area eucalyptus. White gum, Swamp gum. 2n=22. SE.
it is very hardy. It is valuable in breeding hardy Australia and E. Tasmania. Cultivated in the
strains. Some geographical races and spontaneous Mediterranean region, in countries of C. and S.
hybrids a r e known. Africa, India, New Zealand and USA. In subtropical
areas of the USSR it is the commonest Eucalyptus
EUCALYPTUS PERRENIANA F. Muell. ex Rodway. species. The wood is of moderate value. An e s s e n -
2n= . Tasmania. Cultivated in the USSR. It is tial oil is obtained. Many spontaneous and artificial
hardy (it tolerates -13°C). hybrids a r e known.
EUCALYPTUS REGNANS F. Muell. Mountain ash, LEPTOSPERMUM LAEVIGATUM F. Muell. Aus-

Swamp gum, Australian oak. 2n= . Mountains tralian t e a - t r e e . 2n=22. Australia. Cultivated there
of the S. Victoria up to 900 m and in Tasmania up for the reclamation of moving sand. Dried leaves
to 600 m. Cultivated in Kenya, New Zealand and are used for tea-making.
other countries. This is the biggest and most va-
luable species in this genus. Some t r e e s a r e r e - MELALEUCA PREISSIANA Schan. 2n= . Austra-
corded up to 96 m high. It is closely related to lia. Var. leiostachya Schan. (syn. M. parviflora
E. amygdalina*. Lindl. ) is a soil stabilizer.
EUCALYPTUS RESINIFERA Smith (syn. E. s p e c - Protaceae

tabilis F. Muell., E. hemilampra F. Muell.).
Kino eucalyptus, Red mahogany, Forests mahogany. HAKEA SALICIFOLIA (Vent. ) B. L. Burtt. 2n= .
2n=22. Coastal zone of S. Queensland and central SE. Australia and Tasmania. Cultivated for r e c l a -
part of New South Wales. Cultivated in Argentine, mation of arid land in Spain and Portugal. It has
Ceylon, Ethiopia, Cameroon and other countries. run wild in these countries.
The wood is very valuable.
HAKEA SERICEA Schrader. 2n=20. E. Australia.
Cultivated for reclamation of arid land in Portugal
EUCALYPTUS ROBUSTA Smith. Beakpod eucalyp-
and Spain. It has run wild in these countries.
tus, White mahogany, Swamp mahogany. 2n=
Coasts of S. Queensland as far as S. of New South
Wales, Australia. Cultivated in the Mediterranean MACADAMIA INTEGRIFOLIA L.S. Smith, (syn.
region, Africa esp. Cameroon, Zaire and Kenya; M. ternifolia F.V. Muell., M. ternifolia var.
Argentine, India and other countries. Often culti- integrifolia) and M. tetraphylla L . A . S . Johnson.
vated on swampy grounds. The wood is economi- Queensland nut, Macadamia nut, Australian bush
cally valuable. nut, Australian hazelnut. 2n=28, 56. E. Queens-
land, Australia. Cultivated in Hawaii. M. integri-
folia is known as the smooth-shell type and M.
EUCALYPTUS SALIGNA Sm. Sydney blue gum.
Saligna gum. 2n=22. Coasts and slopes of moun- tetraphylla as the rough-shell type. M. ternifolia
tains in New South Wales and SE. Queensland, is now considered to apply correctly only to a s p e -
Australia. Cultivated in Africa, esp. Cameroon, cies with bitter cyanogenic seeds less than 25 mm
Kenya and Rhodesia; S. America esp. Argentine in diameter, inedible and never cultivated (Kraus &
Hamilton, 1970).
MYRTACEAE-SOLANACEAE 61
Rutaceae
EREMOCITRUS GLAUCA (Lindl. )Swing. 2n=18.
This tree is capable of withstanding 6 months
drought. It easily crosses with Citrus species
giving fertile hybrids.
Solanaceae
DUBOISIA HOPWOODII F. V. Muell. Pituri, Pitche-
ry. 2n= . Australia. Cultivated for some decades
to yield atropine.
DUBOISIA LEICHHARDTIIR.Br. 2n=60. Australia.

Cultivated for some decades to yield atropine.
DUBOISIA MYOPOROIDES R. Br. Corkwood,

Mgmeo. 2n=60. Australia. Cultivated to yield
atropine.
NICOTIANA DEBNEYI Domin. 2n= . Australia.

Used as a source of resistance to blue mold,
caused by Perenospora tabaclna Adam.
NICOTIANA GOODSPEEDII Wheeler. 2n=40. New

South Wales to SE. of W. Australia. Is has a
short growing period. It is very resistant to P e r o -
nospora tabacina Adam. Some natural introgression
with the closely related N. exigua Wheeler, 2n=32,
N. suaveolens Lehm., 2n=(24), 32, (48, 64), and
N. rotundifolia Lindl., 2n=44.
SOLANUM LACIANIATUM Aiton. 2n= . Australia

and New Zealand. Cultivated in Europe and e l s e -
where for the foliage which is a source of steroid
p r e c u r s o r s (Tutin et a l . , 1972).
4 Hindustani Centre
The Hindustani Centre of diversity was included by Vavilov in the Tropical South Asian Centre of Origin.
Zhukovsky (1968) separated this centre only by number (TV), but in 1970 he drew on the map a line b e -
tween both centres. He based this separation on the existence of specific species of this Centre 4.
Although this centre is not far from the old farming sites in Thailand, agriculture must have been in-
troduced from the NW adjacent area. Early farming sites have so far revealed few details of crops culti-
vated. At Müan-jo Daro (Mohenjodaro) and Harappa on the Indus, Pakistan more or less on the boundary
between Centres 4 and 5, a site of the Harappan culture dating 2500 - 2000 BC. was discovered. Some
remains of Gossypium arboreum have been discovered. At a site, Navdatoli-Mahesvar on the Narbada
River, in Central India, dated from 2000 BC. remains of wheat, peas, broad beans, lentils, Lathyrus
sativus and rice have been found. Except rice all these crops have been first domesticated outside India.
Important crops of this region a r e bamboos, fruit trees, Cucurbita sativa, Mangifera indica, Musa
s p . , Oryza sativa, Phaseolus mungo, Piper s p . , Saceharum sinense, Vigna sinensis.
Species of this centre have influenced the development of crops in other areas mainly due to an active
distribution between this region and areas such as Ancient Egypt, Assyria, Sumeria, the Hittite Empire.
Much exchange has existed with Africa while many crops were distributed especially to the Mediterranean
region by the Arabs in the 8th-10th centuries AD. Such crops are citrus trees, cotton species, jute, rice,
sugarcane etc.
Acanthaceae Amaranthaceae
BARLERIA PRIONITIS L. 2n=30, 40. Trop. Africa AMARANTHUS ANGUSTIFOLIUS Lam. 2n=32, (34).
and Asia. Cultivated in India as a medicinal crop. S. and C. Europe, Ante-Asia up to India and T u r -
kestan and to Africa. In India var. polygonoides
Agavaceae Thell. is cultivated.
SANSEVERINIA HYACINTHOIDES (L. ) Druce (syn. CELOSIA ARGENTEA L. Quail g r a s s . 2n=(36),
S. zeylanica Willd. ). Ceylon bowstring hemp. 72. India. Var. cristata Kuntze (syn. C. cristata
2n= . Ceylon. A fibre plant cultivated there. L. ), Cockscomb grass, 2n=36. It is a potherb,
fodder and fibre crop and an ornamental.
Alliaceae
ALLIUM AMPELOPRASUM L. Levant garlic, P e - Annacardiaceae
rennial sweet leek. 2n=16, (24), 32, genome formu- MANGIFERA INDICA L. Mango. 2n=40. Assam and
la AAA'A", (40, 48). S. Europe, Asia Minor, the Chittagong Hills. Spread to many tropical
Caucasus to Iran and N. Africa. Some cultivation countries. Rhodes et al. (1970) classified culti-
in Germany and France (p. 129) and in Kashmir vars into:
(Koul &Gojil, 1970). The wild and cultivated types 1. polyembryonic group with oblong fruits, com-
a r e both extremely variable. This species is r e - mon in SE. Asia,
lated to A. sativum*, A. p o r r u m * a n d A . scorodo- 2. monoembryonic group with roundish fruits
prasum*. common in India and
ACANTHACEAE - CUCURBITACEAE 63
3. a group intermediate in fruit shape, also Convolvulaceae

common in India, IPOMOEA ERIOCARPA R. Br. 2n= . India.
4. the Sandersha-Haden complex consists of hy-
Used as a spinach and as a green fodder.
brids developed in Florida and Hawaii. M. odora-
ta* and M. zeylanica Hook,f, a r e not closely r e -
Cruciferae
lated.
BRASSICA CAMPESTRIS L. 2n=20, genome for-
Apocynaceae mula AA. See p. 135 for the origin of this species.
In Pakistan/India the var. toria Duthrie & Fuller,
NERIUM INDICUM Mill. (syn. M. odorum Soland. ). Indian rape, Toria, and var. sarson Prain, Indian
2n=22. Trop. Asia especially India. Cultivated as colza, Brown sarson a r e cultivated. Brown sarson
a medicinal plant. can be divided into 1. self-compatible and 2. self-
incompatible types. These two types can be diffe-
RAUVOLFIA SERPENTINA Benth. 2n=(20), 22, rentiated by disruptive selection for flowering
(24, 44). India, Ceylon, Burma and from Thailand time, genetic drift in isolated populations, and
to Java. Because of the high demand for this medi- chromosomal inversions suppressing recombina-
cinal crop it became (nearly) extinct in some tion in connection with a recessive mutation of a
a r e a s . To provide sufficient roots some hospitals major gene, independent of the S locus but inacti-
in India set up small gardens of it. Its cultivations vating this locus (Swamy Rao, 1971).
could be extended to India and elsewhere (Dutta et
a l . , 1963).
ERUCA VESICARIA (L. ). Cav. (syn. E. sativa L. ).
2n=22. The Mediterranean region (p. 95) and Asia.
Araceae Cultivated in India for jamba oil.
ALOCASIA CUCULLATA (Lour. ) Schott. Giant
taro. 2n=28. India and Ceylon. Cultivated for its RAPHANUS SATIVUS L. Serpent radish, Snakelike
corms. radish, Rat-tailed radish, Tree radish from Java.
2n=18. Cultivated from Java to NW. India. The
ALOCASIA MACRORRHIZA (L.) Schott. Giant plant requires a short daylength to develop roots.
Alocasia. 2n=26, (28). Probably Ceylon. Spread This group of cultivars var. mougri Helm. R. cau-
in the Malay Archipelago and to India and further datus L., R. sativus var. indicus Sinsk. *, is
to tropical America. Several varieties a r e cyano- characterized by small long fleshy fruits and gla-
genic. A. indica* is often included in this species. brous leaves. Var. oleiformis P e r s . * is also grown
in India.
AMORPHOPHALLUS CAMPANULATUS Blume.
Whitespot giant arum, Oroy. 2n=(14), 28. Trop.
Asia (p. 43). Cultivated in India and elsewhere as
a tuber crop.
Asclepiadaceae
MARSDENIA TINCTORIA R. Br. 2n= . Hima-
laya to China, Malaysia. Cultivated in India as a
dye plant.
Cannabidaceae
CANNABIS SATIVA L. Hemp. 2n=20. Centre of
origin C. Asia (p. 130). Spread to India in early
times. The Indian type is cultivated for its n a r c o -
tic properties. From this country it must have
spread to the Middle East and other countries.
C. ruderalis Janisch, (2n= ), SE. Russia and
C. Asia. This weed perhaps derived from the cul- Raphanus sativus var. mougri (Sinskaya, 1931)
tivated form.
Chenopodiaceae Cucurbitaceae
KOCHIA INDICA Wight. 2n=18. Introduced into CITRULLUS COLOCYNTHIS*
Egypt where it is cultivated as a forage crop.
CITRULLUS LANATUS (Thunb. ) Mansf. Water-
Compositae melon. 2n=22. Var. fistulosus (Stocks) Duthie &
Fuller (syn. C. fistulosus Stocks) is cultivated in
CARUM COPTICUM (L. ) Benth. &Hook. (syn. India for its small round fruits (Purseglove, 1968).
Trachyspermum ammi (L.) Sprague. Ammi. 2n=18. It is only known cultivated.
India. It yields an essential oil.
COCCINIA CORDIFOLIA Cogn. (syn. C. indica
VERNONIA AMYGDALINA Delile. Bitterleaf. W. & A. ). Ivy gourd, Small gourd. 2n=24, (36).
2n= . Trop. Africa. Occasionally cultivated. Trop. Asia, and in the Red Sea area to Sudan. In
S. India forms occur with long less bitter fruits.
HINDUSTANI CENTRE 64
CUCUMIS SATIVUS L. Cucumber. Gherkin. TRICHOSANTHES CUCUMERINA L. (syn. T. an-

2n-14. P r i m a r y gene centre probably the Himala- quina L. ). Snake gourd. 2n=22. India to Australia.
yas. Introduced into Europe, Near East. China Cultivated for a long time in India.
and other countries. Secondary gene centres in
China and Near East (p. 30 and 79). In India Dioscoreaceae
cucumbers have been cultivated at least 3 000
years (Leppik, 1965). The closely related wild DIOSCOREA HISPIDA*.
species C. hardwickii Royle also grows in this
centre. It might, however, be a weedy form of Euphorbiaceae
the cultigen. BACCAUREA SAPIDA Muell. -Arg. 2n=
Sources of resistance to mildew (Erisyphe Malaysia, India and China. Cultivated by Hindus
cichoracearum DC (ex Merat emend. Salm. ) and/ for its fruits.
or Sphaerotheca fuliginea (Schlecht, ex Fr. ) Poll
are found in cucumber material from China, Japan, CROTON TIGLIUM L. Purging croton. 2n=
Indonesia and India. SE. Asia. Cultivated now in India and Ceylon for
its seeds (Purseglove, 1968).
CUCURBITA MAXIMA Duch. ex Lam. Pumpkin,
Winter squash. 2n=40. Its origin is described on Flacourtiaceae
p. 147. Secondary centre in India and adjacent
areas. DORYALIS POVYALIS) HEBECARPA (Gardn.)
Walb. (syn. Aberia gardneri Clos. ). Ceylon goose-
LUFFA ACUTANGULA (L. )Roxb. Sponge gourd, berry. 2n= . Trop. Asia especially India and
Angled loofah, Sinkwa towel gourd. 2n=26. P r o b a - Ceylon. Cultivated for its b e r r i e s .
bly India. P r i m a r y gene centre probably h e r e .
It was found in Karakoram/Hindukush in 1955 HYDNOCARPUS LAURIFOLIUS p e n n s t . ) Sleumer
(Tozu, 1965). Cultivated in China and Japan. (syn. Hydnocarpus wightianus Blume). 2n=24.
Dutt &Roy (1971) suggested the following India. Cultivated in several trop, countries for
evolution of the various related Luffa species. its oil used to cure leprosy.
They considered the wild monoecious L. graveo-
lens Roxb., (2n=26) as the prime species. From Gramineae
this species two species derived: the wild dioecious BAMBUSA ARUNDINACEA (Retz. ) Willd. Spiny
L. echinata Roxb. (2n=26) and the cultivated m o - bamboo. 2n=70. India. Cultivated there. In forests
noecious L. aegyptica*. These authors considered bordering r i v e r s and on mountains as high as 900 m.
dioecism as a derived factor that arose after di- Much cultivated in Java, r a r e l y in Indochina.
vergence from the monoecious L. graveolens. Secondary gene centre in Java (p. 46). Used as
From L. aegyptica the monoecious type of building material and in the paper industry. This
L. acutangula and later the hermaphrodite type of is one of the largest bamboo species. Its stems
this species arose. The latter type is also named can be 30 m long.
L. hermaphrodita*.
BAMBUSA POLYMORPHA Munro. 2n=72. Bangla
LUFFA AEGYPTIACA Tull. (syn. L. cylindrica Desh. Cultivated there and in Burma. Used as
(L. ) Roem. ). Smooth loofah, Suakwa, Sponge building material and in the paper industry.
gourd, Vegetable gourd. 2n-26. Domesticated
probably in tropical Asia, possibly in India. Cul- BAMBUSA STRICTUS Nees. 2n=70, 72. In India
tivated in almost all tropical regions where it may (except Assam) and Burma. Secondary centres in
have run wild. Used for producing vegetables or Indochina (p. 32) and S. China (p. 32). In forests
sponges. It is also used as a medicine, isolation in dry a r e a s . It is drought resistant.
material etc.
This species includes L. racemosa Roxb. BAMBUSA TULDA Roxb. 2n= . India, Burma
(2n= )with L. hermaphrodita* the only two (p. 47) and Tahiti. Secondary centre Java.
'species' with bisexual flowers.
CEPHALOSTACHYUM CAPITATUM Munro.
LUFFA HERMAPHRODITA Singh & Bhandari. 2n= . Burma and Himalaya. Its shoots a r e
Satputia. 2n= . Cultivated in Bihar and Bengal, edible.
India. It c r o s s e s easily with L. acutangula*, the
F i being monoecious. It is similar to this latter CYMBOGON FLEXUOSUS (Nees ex Steud. ) Wats,
species and to L. cylindrica (see L. aegyptiaca*), (syn. Andropogon flexuosus Nees). Malabar g r a s s ,
but has besides bisexual flowers, oblong-ellipsoi- Cochin g r a s s . 2n=20, 40. India. Cultivated for its
dal fruits and smooth shining black seeds. Maybe essential 'East Indian lemongrass' oil.
this species is a hybrid of one of these two species.
Types described as L. racemosa Roxb. and in- CYMBOGON MARTINI (Roxb.)Wats. Rosha g r a s s .
cluded in L. cylindrica also a r e bisexual (Singh & 2n=20, 40. E. India. Cultivated in India and Indo-
Bhandari, 1963). nesia for palmarosa oil. Var. motia (syn. C.
Dutt &Roy (1971) included L. hermaphrodita motia Gupta) (2n=40) produces palmarosa oil, while
in L. acutangula*. They consider it as the h e r m a - var. sofia (2n=20) is the source of gingergrass oil.
phrodite type of this latter species.
CUCURBITACEAE - GRAMINEAE 65
CYNODON DACTYLON (L. ) P e r s . Bermuda g r a s s . MELOCANNA BACCIFORA (Roxb. ). Kurz.

2n=(18, genome formula AA, 27, 30), 36, genome 2n= . Burma from Garo to Arakan. P r i m a r y
formula AABB, 40, (54, genome formula AAAABB), centre: Bangla Desh. Its stems are used as buil-
aneupl., B-chromosomes. P r i m a r y centre: India, ding material.
although Indo-Malaysia and trop. Africa have also
been mentioned. Spread all over the world. Malik NEOHOUZEAUA DULLOSA (Munro) A. Camus.
&Tripathi (1968) report that the A and B genome 2n= . S. and SE. Burma. It has a very long
have segments in common. The hexaploid fibre and produces much paper m a s s .
AAAABB-type is found in Udaipur, India. It has a
recent origin and hence a restricted distribution. OCHLANDRA TRAVANCORICA Bedd. 2n=c. 72.
Harlan &de Wet (1969) suggested two putative P r i m a r y centre: the Travancore mountains and
parents. One of them is the diploid var. aridus Tinneville (900 - 1 650 m altitude). It is used in
Harlan &de Wet, (2n=18), which originally had a the paper industry.
small range but is now found in India, Ceylon,
Israel, Zambia, Tanzania and S. Africa. The ORYZA MALAMPUZHAENSIS Krish. & Chand.
other putative parent is the diploid variety afgha- 2n=48. India.
nistan Harlan &de Wet. From var. aridus arose
var. dactylon, (2n=36). From this race the t e m p e - ORYZA SATrVA L. Rice. 2n=24, genome formula
rate race of var. dactylon arose. Through crossing AA. P r i m a r y centre SE. Himalayan region (Mori-
with tetraploids of var. afghanistan and inter- naga, 1968). The cultivated rice developed from
crossing the seleucidus race of var. dactylon d e - O. perennis Moench (syn. O. rufipogon - perennial,
veloped. Nayar, 1973), 2n=24, genome formula AA. O. p e -
Other botanical varieties a r e var. elegans rennis has a wide distribution in the tropics of
Rendle, (2n=36), of Southern Africa, var. pole- Asia, the Americas, the Mediterranean Region and
vansii (Stent. ) Harlan &de Wet (2n=36) of S.Africa N. Australia. There a r e four main geographical
and var. coursii (A. Camus) Harlan &de Wet races mainly based on breeding behaviour and in-
(2n=36) of Madagascar. terracial sterility: 1. Asian, (floating and p r o -
cumbent habit, partial self-incompatibility),
DENDROCALAMUS HAMILTONII Nees &A m . 2. American (p. 165) (caespitose), 3. African
2n= . C. and E. Himalaya up to 900 m and (p. 115) (erect, well developed rhizomes), self-
Upper and Lower Burma up to 1 200 m altitude. incompatibility), and 4. Oceanian (p. 47). How-
Used as building material and in the paper industry. ever, floating types have been found in the Amazone,
It grows in humid areas along r i v e r s and low places erect types with rhizomes in Asia and non-rhizo-
and forms large thickets. matous, floating types in C. Africa.
The Asian race is the parent of O. sativa (Nayar,
DENDROCALAMUS LONGISPATHUS Kurz. 2n=72. 1973), while annual types (rufipogon-annual, var.
Bangla Desh and Burma (Arakan). Used for paper fatua, f. spontanea, O. nivara*) a r e intermediates
making. It is found in humid mixed forests along of perennial plants and cultivated rice. Much gene
r i v e r s on fertile clay soils. exchange exists between wild-perennial, weedy-
annual and cultivated-annual rice, resulting in
DIGITARIA CRUCIATA (Nees) A. Camus. 2n= forms described as var. fatua or f. spontanea or
Var. cruciata grows wild in a large a r e a of E. as O. rufipogon* and O. nivara*.
India and China. Probably domesticated in the 19th
Century in the Khasi hills, E. India by selecting
var. esculenta Bor with longer stems, longer
spikes, l a r g e r inflorescences and much bolder
grains. It grows slowly and yields little. Its ad-
vantage is the production of straw in an area where
little grass grows (Bor, 1955; Singh &Arora,
1972).
DIGITARIA SANQUINALIS Scop. 2n=(18, 28), 36

(-48, 54, 76). This cereal was cultivated in India
a n d E . Europe (p. 133) (Portêres, 1955).
ECHINOCHLOA COLONA (L. ) Link. Shama millet,

Jungle rice. 2n=(36, 48), 54, (72). Cultivated in
India as a fodder grass and as a cereal. Formerly
it was also used in Egypt.
Yabuno (1968) considered the genome formula Centre of origin of rice (Chatterjee, 1951)
of this species to be the same as that of E. frumen-
tacea*.
Morinaga (1968) classified rice in 4 genecolo-
ELEUSINE COROCANA (L. ) Gaertn. Finger millet, gical groups:
Ragi. 2n=36. Introduced from Africa (p. 113). 1. ecospecies 'aman' (ssp. indica Kato) divided in
Kempanna (1969) recognized various types in India. three ecotypes 1. aman (winter rice), 2. boro
(summer rice) and 3. tjereh
2. ecospecies 'aus' (autumn rice) 2n=40-128, with euploids 40, 48, 64, 80, 96, 104,
3. ecospecies 'bulu' 112, 128 and possibly 54. Probably India at the
4. ecospecies 'japonica' divided in 1. ecotype foothills of the Himalaya Mountains. Now it is d i s -
'japonica' (ssp. japonica Kato) and 2. ecotype tributed in innumerable groups of different ranks
'nuda'. and significance from Africa over Asia to Japan
He based his classification on the extent of their and the Solomon Islands. One group (2n=112) is
affinity as shown by the hybrid fertility. found in Indonesia (p. 48), while another one
The ecotypes aman, boro and aus a r e found (2n=104-128) occurs in E. Africa (p. 116). Recent-
in the Bengal-Assam region of India. The e c o - ly introduced into New Guinea and hence its influen-
types tjereh and bulu developed in Indonesia ce there is still limited.
(p. 48) and ecotypes japonica and nuda together S. spontaneum is used as a source of disease
comprising ecospecies 'japonica' developed in resistance of S. officinarum*. In N. India it has
Indo-China (p. 48). In parts of Asia, notably in hybridized with S. sinense* group Saretha.
Bangla Desh and in Burma floating paddy is c u l -
tivated on a large scale. The floating habit is the SINOCALAMUS GIGANTEUS (Walb.) Keng. 2n=
capability to lengthen the stem with increasing India, Indochina and S. China. Cultivated there.
water depth. It is conditioned by two recessive Used as building material. It is one of the largest
genes. Nayar (1973) showed that O. sativa is the bamboos.
ancestral species of the African rice, O. glaber-
rima*. SORGHUM BICOLOR (L. ) Moench. Juar, Jowar.
2n=20. P r i m a r y gene centre: Africa (p. 116).
PASPALUM SCROBICULATUM L. Koda millet, Secondary centre: India. No hybridization has
Ditch millet. 2n=40. Wild types var. c o m m e r - occurred with wild sorghums as these are t e t r a -
sonii (Lam.) Stapf (syn. P. commersonii Lamk. ) ploids (Doggett, 1970).
and var. polystachyum (R.Br.) Cheval., and the
cultivated type var. acrobiculatum. Var. c o m m e r - TRITICUM AESTIVUM (L. ) Thell. ssp. sphaero-
sonii is found in the tropics of the Old World, and coccum (Perc. ) MK. (syn. T. sphaerococcum
var. polystachyum in Africa (Mansfeld, 1959). P e r c . ). Indian dwarf wheat. 2n=42, genome formu-
Koda millet is cultivated in China, Japan, Ante- la AABBDD. Transcaucasia and adjacent regions
India and Australia. (p. 82). Ssp. sphaerococcum is indigenous to
NW. India and adjacent Afghanistan. It is c h a r a c t e -
SACCHARUM SINENSE Roxb. Chinese sugarcanes, rized by short, non-lodging culms, erect leaves,
North Indian sugarcanes. Mungo group 2n=81-83, globular grains and susceptibility to diseases.
Dhaulu group 2n=82, 83, Saretha group 2n=91,
91 + fragrants, 92, Nargori group 2n=105-119, VETIVERIA ZIZANIOIDES (L. ). Nash. Vétiver.
124, Pansahi group 2n=106-120, and unclassified 2n=20. Ceylon, India up to Burma. A grass culti-
2n=c. 104-121. The first four groups have also vated in the tropics for its essential oil.
been classified as S. barberi J e s w . , the North
Indian sugarcanes, so then the Pansahi group ZEA MAYS L. 2n=20. Probably domesticated in
(Chinese sugarcanes) is the only group of S. sinen- C. America (p. 166). Secondary centre: the S.
se. It is possible that the original North India Himalayas (Brandolini, 1970). Flint maize (indu-
sugarcanes have derived from wild S. spontaneum* rata Sturt. ) is common here.
(Grassl, 1968) while after introduction of S. offi-
cinarum* hybridization may have taken place. The Guttiferae
present North Indian sugarcanes a r e considered as GARCINIA INDICA*.
complex polyploid hybrids of S. officinarum*
(x=17) and types of S. spontaneum* (n=40, 48, 56).
GARCINIA SILVESTRIS Boerl. Wild mangosteen.
This hybridization would have occurred in Bengal, 2n= . Malaysia (p. 49) and India. Parental
Orissa and Bihar, India (Parthasarathy, 1946, species of G. mangostana*.
1948; Bremer, 1966). P r i c e (1968) suggested that
the Mungo group derives from one plant possibly GARCINIA TINCTORIA*.
a mutant of the closely related Dhaulu group. How-
ever, Grassl (1968) suggested an Erianthus sp.
MESUA FERREA L. Nahor, Nagas tree, Indian
introgression in the origin of the Mungo group. The
rose chestnut, Ironwood. 2n=32. Trop. Asia, In-
Saretha group may also originate from one plant
dia and Malaysia. Cultivated in India as a timber
and so may the Nargori group (2n=124). The P a n -
t r e e and for its flowers and fruits. The flowers
sahi group may be a hybrid of S. officinarum* and
a r e used in the perfume industry, the fruits a r e
5. spontaneum* or a Miscanthus species. The latter edible and the seeds contain oil for lighting.
may be M. sinensis*, but no tetraploid types have
yet been found. Leguminosae
S. sinense is found in SE. Asia and N. India.
In India it is a cottage industry crop from which ALBIZIA STIPULATA Boiv. (syn. A. chinensis
a crude brown sugar is obtained (Price, 1963). (Osb. ) Merr. ). 2n= . Cultivated in India and
Ceylon for its high quality fodder. Elsewhere it
SACCHARUM SPONTANEUM L. Wild sugarcane, is cultivated as a shade tree, green manure and
Kassoer, Thatch grass, Bagberi, Dharb, Khus. cover crop.
GRAMINEAE - LINACEAE 67
BAUHINIA PURPUREA L. Camel's foot. 2n=28. MUCUNA CAPITATA*.

China to India. A t r e e cultivated in India for
various purposes and in trop. Africa as a fodder MUCUNA PACHYLOBIA (Piper &Tracy) Rock
plant. (syn. Stizolobium pachylobium Piper & Tracy).
Fleshy pod bean. 2n= . India. Cultivated as a
CAJANUS CAJAN (L. ) Millsp. Pigeon pea. vegetable.
2n=22, 44, 66". Africa (p. 118). Secondary centre:
India (Purseglove, 1968). MUCUNA UTILIS Wall, ex Wight. Velvet bean.
2n= . India. Cultivated as a vegetable, as
CANAVALIA ENSIFORMIS (L. ) DC. Jack bean, cattle food and as a green manure.
Horse bean. 2n=22. S. America (p. 151). Secon-
dary centre: India. PHASEOLUS ACONITIFOLIUS Jacq. (Ph. trilobus
Wall. ). Mat bean, Moth bean. 2n=12. India, Bangla
CASSIA AURICULATA L. Tanner's cassia. 2n=14, Desh and Burma. Cultivated in these countries and
16, 28. Cultivated there for its tanning material also in Ceylon and China. In the USA it is culti-
obtained from the bark (Purseglove, 1968). vated as a fodder crop.
CASSIA FISTULA L. Indian laburnum, Purging PHASEOLUS MUNGO L. (syn. Vigna mungo (L.)
cassia. 2n=24, 26, 28. India. Cultivated in the Hepper. ). Black gram, Urd. 2n=22, (24). Un-
tropics for its pods of which the pulp round the known wild. Closely related to P. aureus* and may
seeds is used as a purgative (Purseglove, 1968). together even form one species (Verdcourt, 1970).
Ph. aureus var. sublobata (syn. P. sublobatus
CASSIA SIAMEA*. Roxb., P . trinervis Wight & Arn. ) is likely the
wild form. (It is certainly the ancestor of P.
CICER ARIETINUM L. Gram, Chickpea. 2n=14, aureus.)
16, (24, 32, 33). Probably W. Asia (p.000). P r o -
bably a secondary centre in India. Introduced in SESBANIA ACULEATA (Pers.) Poir. Dhanchia.
India in early t i m e s . Cultivated much in India now. 2n=(12), 24, 32. Cultivated in Bengal for its fibre
Strains with fine dark-brown and black seed have and especially as a green manure crop.
been cultivated for a long time. 'Kabuli' types have
been introduced from Afghanistan in about 1700 SESBANIA AEGYPTIACA*.
(van der Maesen, 1972).
SESBANIA SPECIOSA Taub, ex Engl. 2n=12.
CROTALARIA BURHIA Buch. -Ham. 2n=16. This India (?). Cultivated there as a green manure of
fibre crop comes from E. India. rice fields.
CROTALARIA JUNCEA L. Sunn hemp, Sann hemp. VIGNA UNGUICULATA (L. )Walp. (syn. V. sinen-
2n=16. Probably India. Unknown wild. A bast fibre sis (L. ) Savi, Dolichos sinensis L. ). Cowpea,
crop. Cultivated in many tropical countries as a Black eye, Southern pea. 2n=22, 24. P r i m a r y
crop (green manure) (Purseglove, 1968). centre of diversity: W. Africa. Secondary centre:
India. Probably originally domesticated in W. and
CYAMOPSIS TETRAGONOLOBA (L. ) Taub. (syn. C. Africa (p. 120).
C. psoralioides DC.). Guar, Cluster bean. 2n=14. From Africa cowpea was taken to the Indian
Probably domesticated in Africa (Anderson, 1960). subcontinent. In Africa the large-seeded type was
However, Hymowitz (1973) described that seeds of selected for, while in India ssp. cylindrica (L.)
C. senegalensis* arrived in Africa as flotsam in Van Eseltine and ssp. sesquipedalis (L. ) Verde,
Arab-Indian (horse) trade. Subsequently it became were obtained. In N. Nigeria a fibre form was
domesticated in India. Cultivated in India, P a k i s - developed. The fibre is obtained from the peduncles.
tan and elsewhere for fodder, food and as a source From W. Africa and India cowpea cultivars
of gum. No wild forms in the Indo-Pakistan conti- spread over the world (Faris, 1965).
nent. The explosive opening of the pod of the wild
type (ssp. dekindtiana (Harms) Verde., syn. V.
DOLICHOS UNIFLORUS Lam. (syn. D. biflorus dekindtiana Harms) distinguishes this type from
auct. non Linn). Horsegram. 2n=20, 22, (24). the cultivated types.
The tropics of the Old World. Especially found in
India and the Himalayas where it is also cultivated. Linaceae
INDIGOFERA PILOSA Poir. 2n=16, 32. Used in LINUM USITATISSIMUM L. Flax. 2n=30, (32).
Ceylon as a green manure. Its possible origin is given on p. 87. In India and
adjacent regions flax of the ssp. indo-abyssinicum
MELILOTUS INDICUS All. Indian clover, Yellow Vav. &Ell. a r e found. It is identical to flax of
annual sweet clover, Sour clover. 2n=16. Punjab, Ethiopia and Eritrea and it may have originated in
India to the Mediterranean region and Turkestan. these countries. This subspecies hybridizes with
Cultivated in N. India as a fodder crop and in the ssp. mediterraneum* resulting in a hybrid ssp.
USA as a cover crop. hindustanicum Vav. &Ell.
Malvaceae GOSSYPIUM STOCKSII Masters. 2n=26, genome

ABELMOSCHUS ESCULENTUS (L. ) Moench (syn. formula EÊ^. Sind, India and SE. Arabia. It is
Hibiscus esculentus L. ). Okra, Lady's finger. drought resistant
2n=72-132. Probably a cultigen developed from a
wild species in trop. Asia. A. tuberculatus Pal & HIBISCUS RADIATUS Cav. 2n=72, genome formula
Singh (2n= ). It could be the ancestral or wild AABB. India to Burma.
form of A. esculentus (van Borssum Waalkes, Introduced into Africa and elsewhere where it
1966; Bates, 1968). It is a N. Indian species is cultivated. Mainly used as an ornamental; cul-
differing from A. esculentus only in having s t r i - tivated in Java as a vegetable and a drug plant.
gose pubescence on the stems and shorter c a p - It is a source of root-knot nematode r e s i s t a n -
sules beset with bristly tuberculate h a i r s . Van ce for H. sabdariffa*.
Borssum Waalkes suggested that A. tuberculatus It is an allotetraploid of H. cannabinus* and
be included in A. esculentus. another diploid species. If this species is an indi-
A. esculentus is commonly known by its syno- genous species of India the amphiploidization must
nym. have taken place in that country after the intro-
duction of H. cannabinus as a fibre crop. How-
ABELMOSCHUS MANIHOT (L. ) Medicus, (syn. ever, the only known diploid B - c a r r i e r is the Afri-
Hibiscus manihot L. ). 2n=60, 66. India, Pakistan, can H. surattensis* (Menzel and Martin, 1971).
through S. China to New Guinea and N. Australia. If the origin of H. furcatus Roxb. non Willd.
Cultivated for its immature fruits. (2n=144), genome formula BBGGWWZZ from India
Ssp. manihot is the cultigen which must have and Ceylon is studied, the origin of H. radiatus
been selected by man from wild, hairy and prickly may also be solved.
types (ssp. tetraphyllus (Roxb. ex Hörnern.)
B o r s s . , syn. Hibiscus tetraphyllus Roxb. ex SIDA RHOMBIFOLIA L. Queensland hemp, Broom-
Hörnern. ). jue sida, Cuba jute. 2n=14, 28. The tropics. Cul-
tivated in India and later in Queensland, Australia
ABELMOSCHUS MOSCHATUS Medicus, (syn. Hi- as a fibre crop (var. retusa L. ). It is an e x t r e m e -
biscus moschatus L. ). Musk mallow. 2n=72. India, ly variable species.
S. China, Indochina to Indonesia and SW. Pacific
islands to New Guinea and N. Australia. Centre of Moraceae
origin possibly E. India (Mansfeld, 1959). Culti- ARTOCARPUS HETEROPHYLLUS Lam. (A. inté-
vated for its seeds which a r e used as perfume, for gra (Thunb. ) M e r r . , A. integrifolia L.f. ). J a c k -
its immature edible fruits and often as an ornamen- fruit. 2n=56. Unknown wild. It has a very ancient
tal. It is commonly known by its synonym. cultivation in India. There and in Ceylon it is popu-
lar, elsewhere in the tropics it is also cultivated
ABUTILON INDICUM*. (Purseglove, 1968).
GOSSYPIUM ARBOREUM L. Tree cotton. 2n=26, FICUS ELASTICA Roxb. Indian rubber t r e e . Karet
genome formula AQAO. P r i m a r y centre: peninsular t r e e . 2n=26, (39). India and Malaya. Cultivated in
of India. Unknown wild. Spread in E. and SE. di- India, Java and elsewhere. It is also cultivated as
rection to Burma, Indochina and the Malaysian an ornamental house-plant (Purseglove, 1968).
Archipelago. Centre of domestication probably
lies in Gujarat which is the westernmost state of FICUS RELIGIOSA L. Pipal tree, Peepal tree,
India (Hutchinson, 1971). Close to this region a Bot t r e e . 2n=26. This strangling fig is sacred to
fragment of textile and a string dated 2500-1700 Hindus and Buddhists. It is propagated by cuttings
BC. have been found in Mohenjo-Daro, Pakistan. and layering. A scion planted at Anuradhapura in
Race indicum of peninsular India is more closely Ceylon in 288 BC. (Purseglove, 1968) died in 1971.
related to cottons belonging to G. herbaceum* than
are other races of G. arboreum. It includes both FICUS ROXBURGHIIWall. 2n= . Cultivated for
perennial and annual forms. It is very likely that its figs.
the perennial forms are primitive while the annuals
were selected later. Moringaceae
In N. India and Pakistan race bengalense was
cultivated. Perennial forms are occasionally found MORINGA OLEIFERA Lam. Horse-radish t r e e .
in remote places in Rajputana and in the Ganges 2n=28. India. Cultivated throughout the tropics.
valley. It spread towards the S., SE. and W.
The African race soudanense* was probably the Musaceae
cotton cultivated by the people of Meroë (an ancient MUSA cultivars of the AB group. Ney poovan and
Nubian civilization), who were the first in Africa synonyms and homonyms. 2n=22. Cultivated on a
to spin and weave cotton. Chowdhury and Buth small scale now (p. 52). See p. 52 for discussion.
(1970) suggested, that this race might be indigenous
to Africa rather than introduced from India as a MUSA cultivars of the AAB group. 2n=33. Mostly
textile crop. India (see p. 52). Only the clone Maia maoli has
probably arisen in Philippines.
MALVACEAE - SAPOTACEAE 69
MUSA BALBISIANA Colla. Pisang bau, Klue Tani. Plantaginaceae

2n=22, genome formula BB. India, Burma, Ceylon
PLANTAGO OVATA Forsk. (syn. P. decumbens
and maybe E. New Guinea. Cultivated for its
leaves as a packing material or as a fibre plant Forsk. ). 2n=8. Mediterranean region, C. Asia
(de Langhe, 1969). It is not very variable. It is and India. Cultivated in India.
one of the parents of the AAB, ABB and ABBB
groups (p. 52) of the cultivated bananas, Polygonaceae
RUMEX VESICARIUS L. 2n=lE , (20). Greece,
Myrtaceae N. Africa, India and Malaysia, Cultivated in India
EUGENIA JAMBOLANA *. as a medicinal plant.
RHODOMYRTUS TORMENTOSA*. Rosaceae

RUBUS ALBESCENS Roxb. Mysore raspberry.
Oleaceae 2n= . The mountains of India, Ceylon, Malaya
and Indonesia. Occasionally cultivated especially
JASMINUM GRANDIFLORUM L. Catalonian j a s -
in Puerto Rico.
mine, Italian jasmine. 2n=26. The Himalayas.
Cultivated for its fragrant flowers and used in
perfumery. Rubiaceae
COFFEA BENGALENSE Heyne ex Willd. 2n=22.
JASMINUM SAMBAC (L. ). Ait. Arabian jasmine. Bengal, Burma and Sumatra. Occasionally culti-
2n=26, 39. India and Ceylon. Cultivated in the vated in India (Purseglove, 1968).
tropics. The flowers a r e used for scenting tea and
a r e a source of an essential oil. MORINDA ANGUSTIFOLIA Roxb. 2n= . Trop.
Himalayas, Assam and adjacent a r e a s . Cultivated
Palmae for its bark and wood which is a source of yellow
ARENGA PINNATA (Wurmb. ) Merr. Sugar palm. dye.
2n= . P r i m a r y centre: possibly Malaysian
Archipelago (p. 53). Secondary centre: India. MORINDA CITRIFOLIA L. ('M. bracteata Roxb. ).
Cultivated in many trop. Asian countries. Indian mulberry. 2n= . S. India and Malaysia
upto Pacific isles. Cultivated in India as a dye crop.
BORASSUS FLABELLIFER*.
RUBIA CORDIFOLIA L. Indian madder. 2n=22.
COCOS NUCIFERA L. Coconut palm. 2n=32. SE. Trop, and temp. Asia. Cultivated in India as a dye
Asia, Indonesia and W. Pacific islands (p. 53). plant.
Secondary centre: possibly India.
Rutaceae
Pedaliaceae CITRUS LATIPES (Swing) Tan. 2n= . The hills
SESAMUM INDICUM L. Sesame, Beni seed. 2n=26. of NE. India. The fruits a r e not edible. It can be
P r i m a r y centre: Africa (p. 126). Secondary centre: crossed with other Citrus species including the
India. From here it is believed to have been spread cultigens.
towards the east, through China, Indo-China into
Japan, and towards the west, through Afghanistan, CLAUSENA DENTATA (Willd. ) Roem. (syn. C.
Asia Minor and Iran to the Mediterranean region willdenowii Wight & A m . ). 2n=18, 36. India. This
to reach N. Africa. small t r e e is known for its edible b e r r i e s .
Perioplocaceae FERONIA LIMONIA (L. ) Swing, (syn. Limonia

acidissima L. ). Indian wood apple, Elephant apple.
CRYPTOSTEGIA GRANDIFLORA*. 2n=18. India and Ceylon. Cultivated now in several
trop, countries for its fruits.
Piperaceae
PIPER LONGUM L. Indian long pepper, Jaborandi MURRAYA KOENIGII (L.) Spreng. Curry-leaf t r e e .
pepper. 2n=24, 48, 52, 96. The foot of the Hima- 2n=18. India. Cultivated for its leaves.
layas. Cultivated in India and Ceylon for its spike
which is used as a spice. It resembles P . r e t r o - Sapindaceae
fractum Vahl*. SAPINDUS TRIFOLIATUS L. Soapnut t r e e , Arceta.
2n= . India, Pakistan and Ceylon. Cultivated
PIPER NIGRUM L. Black pepper. 2n=48, 52, 104, for its fruits.
128. The slopes of mountains in the Ghats, Mala-
bar, SW. India at an altitude between 150 and Sapotaceae
2 400 m. Spread to SE. Asia and Philippines. Now
MADHUCA INDICA Gmelin (syn. M. latifolia
cultivated in other trop, countries (Gentry, 1955;
de Waard &Zeven, 1969). (Roxb. ) Macbr. Moa tree, 2n= . N. and C.
India. Cultivated for its flowers which are rich in
nectar.
MADHUCA LONGIFOLIA (Koenig) Macb. (syn. MAOUTIA PUYA Weddell (syn. Boehmerla puya
M. indica J. P. Gmel.). Mahua, Mowra butter Hassk., Urtica puya Wall. ). 2n= . A perennial
t r e e . 2n= . India. Cultivated there. herb of trop. Himalaya, Khasia and Burma. Occa-
sionally cultivated for its fibres.
MANILKARA HEXANDRA (Roxb. ) Dubard.
2n= . S. Asia. Cultivated in India. Verbenaceae
Solanaceae NYCTANTHES ARBOR-TRISTIS L. Tree of s a d -

ness. 2n=44. C. India. Planted near temples. It
ATROPA BELLADONNA L. Belladonna. 2n=(50), is a source of a saffron-yellow dye. The oil is
72. From Spain, the Balkans, Asia Minor to India. used in perfumery. Cultivated as an ornamental.
A medicinal plant. In India var. acuminata (syn.
A. acuminata Royle, 2n=72) is found. It is culti- Zingiberaceae
vated there.
AMOMUM AROMATICUM Roxb. Bengal cardamon,
CAPSICUM ANNUUM L. Bell pepper, cayenne Nepal cardamon. 2n= . India and Pakistan.
pepper. 2n=24. Mexico (p. 171). Secondary centre: Cultivated there.
Asia.
AMOMUM XANTHIOIDES Wall. 2n= Burma.
DATURA METEL L. Hindu datura. 2n=24. India. Cultivated in India.
This medicinal plant is introduced into many parts
of the (sub)tropics. It is often cultivated as an o r - CURCUMA AMADA Roxb. 2n=42. India and P a k i s -
namental. tan. Cultivated in India for Mango ginger.
SOLANUM MELONGENA L. Eggplant, Aubergine, CURCUMA ANGUSTIFOLIA Roxb. East Indian

Melongene. 2n=24, (36, 48). India. Cultivated arrowroot. 2n=42, (64). Himalayan region. Culti-
t h e r e . Secondary centre: China (p. 70). Cultivated vated for its starchy rhizomes.
now throughout the tropics.
CURCUMA CAESIA Roxb. Kalihaldi. 2n=
Strychnaceae Bengal. Occasionally cultivated for its rhizomes.
STRYCHNOS NUX-VOMICA L. Strychnine t r e e . CURCUMA DOMESTICA Val. (syn. C. longa

2n=24, 44. India, Ceylon and Indonesia. Cultivated Koenig non L. ). Turmeric, Curcuma. 2n=(32),
for its nux vomica used in medicine. 62, (63, 64). A completely sterile triploid which
probably arose with the wild C. aromatica Salisb.,
Theaceae Wild turmeric, Yellow zedoary (2n=42) as one
CAMELLIA SINENSIS (L. ) O. Kuntze (syn. C. parent. C. aromatica grows in India. At first
thea Link., Thea sinensis L. ). The origin of tea turmeric may have been used as a sacred plant;
is discussed on p. 39. Secondary centres: India, l a t e r is was cultivated for its rhizomes, which
Assam and Ceylon. are used for flavouring, and for colouring food
and cloth. It spread in early times to China, SE.
Asia and later to other parts of the tropics. There
Tiliaceae
it may have run wild (Purseglove, 1972).
CORCHORUS CAPSULARS L. Jute, White jute,
2n=14. Unknown wild. P r i m a r y centre: India and CURCUMA ZEDOARIA Rose. Zedoary. 2n=63, 64.
Pakistan. It has been cultivated there for a long E. India. Cultivated in SE. Asia, Ceylon and
time. Spread now throughout the tropics. Madagascar. Rhizomes a r e a source of a condi-
ment. Young flowers are used for flavouring food.
GREWIA ASIATICA L. Phalsa. 2n=36. Salt Range,
Puna, India (Mansfeld, 1959). Cultivated in India, ELETTARIA CARDAMONUM (L. ) Maton. Carda-
Ceylon and elsewhere for its fruits. mon. 2n=48. S. India and Ceylon. Cultivated in
India and Malayan Archipelago. Seeds of cardamon
Umbelliferae are used to flavour coffee and in cooking, the oil
ANETHUM GRAVEOLENS L. (syn. Peucedanum is used for perfumery. The cultivated types can be
graveolens (L.) Hiern. ) Dill. 2n=22. Eurasia. divided into three varieties and two groups viz.
Cultivated in Greece, Rome and Palestine (p. 105). group major Thwaites includes the Ceylon variety
Indian types have longer fruits. This species in- and group minuscula Burkill (syn. var. minor Watt)
cludes A. sowa*. includes the Malaban variety and Mysore variety.
Wild types have also been described as belonging
Urticaceae to major. Plants of this group a r e marked by
anthocyanin pigmentation.
GIRARDINIA HETEROPHYLLA Dene. (syn. Urtica
heterophylla Roxb. ). Nilgeri nettle. 2n=20. From ZINGIBER OFFICINALE Rose. Ginger. 2n=22.
NW. Himalaya to Malaysia. In the Nilgeri area, Unknown wild. Probably domesticated in India. It
India var. palmata is cultivated. was known in China and trop. Asia at an early date.
Cultivated now throughout the tropics.
5 Central Asian Centre r-Ç^
X
^^*~*+J~ "
-,1
- ^^"^s^ ^
/ \
s' ""
'•7 J/T
\ T-Jp"
( ;
V.
> JA
The Central Asian Centre was called by Vavilov Southwestern Asian Centre. Zohary (1970) prefered to
join it with Centre 6: the Near Eastern Centre, like Zhukovsky (1968) did, who separated both areas by
number only. But in 1970 both centres were on the map, separated by a line, while Centre 5 was extended
northerly (Zhukovsky, 1970).
This centre served as a transfer zone between centres 1 and 4. Furthermore, the Himalayas have
provided many species as parental stocks for crops.
Agriculture must have reached this centre from Centre 6 at about 5000 BC.
Important crops of this centre a r e fruit t r e e s , Allium cepa, A. sativum, Daucus carota, Lathyrus
sativus, Spinacea oleracea, Vicia faba etc. It is a secondary centre of diversity for Cucumis melo.
Alliaceae SPINACIA OLERACEA L. Spinach. 2n=12. Iran up

to Manchuria. P r i m a r y centre in Afghanistan and
ALLIUM CEPA L. Onion. 2n=16. Probably C. Tadzikistan. In this area the wild, related S. t e -
Asia and esp. NW. India, Afghanistan, Uzbekistan trandra Stev. (2n=12) also grows.
and W. Tienshan where related species A. p s k e -
mense B. Fedtsch. (2n=16)andA. vavilovi Popov Compositae
&Vved. (2n=16) grew wild (Vavilov, 1949/50).
Secondary centre is Centre 7 (p. 91). Another ARTEMISIA CINA Berg. Levant wormseed plant.
related wild species is A. oschaninii O. Fedtsch. 2n=18. The Orient and Russian Turkestan. Culti-
from Pamir-Alai and Tien Shan regions. This vated in Russia and W. of USA.
species is used as foodi Wendelbo (1971) and
McCollum (1974) suggested this species to be the ARTEMISIA DRACUNCULUS L. Tarragon, E s t r a -
wild A. cepa or the wild ancestor of A. cepa. gon. 2n=18, 36, 54, 72, 90. USSR and from W.
The top onion, t r e e onion or Egyptian onion Asia to Himalaya. Perennial widely cultivated as
(var. viviparum (Metzger) Alefeld) is probably a a condiment. The 'Russian' tarragon (2n=90), a
hybrid of A. cepa and A. fistulosum*. Such hybrids decaploid, is fertile, while the 'French' tarragon
have been described as A. aobanum Araki (2n=16). (2n=36), a tetraploid, is sterile and propagated
vegetatively. The types with (2n=45, 54, 72) may
ALLIUM SATIVUM L. Garlic. 2n=16, genome for- be hybrids (Rousi, 1969).
mula SS. Some consider A. longicuspis Regel A. dracunculoides Pursh (2n=18) from N.
(2n=16) as the wild parent of garlic. This species America has often been included in this species.
occurs in C. Asia. Var. pekinense (Prokh. ) Maki-
no became cultivated in China (p. 28). Secondary CARTHAMUS TINCTORIUS L. Safflower. 2n=24,
centres developed in Centres 6 and 7 (p. 78, 91) genome formula BB. P r i m a r y centre in Centre 6
(Kazakova, 1971). Garlic was already known in (p. 78). Secondary centre near Kabul, Afghanis-
Egypt before 3000 BC. tan. The great variability of the safflower popu-
lation here, that led Vavilov to believe this area
Chenopodiaceae to be a centre of origin, is very likely caused by
the meeting of the Middle Eastern and West P a k i s -
BETA VULGARIS L. Beet. 2n=18, (27, 36). P r i - tan safflower types in this area (Knowles, 1969).
mary centre is discussed on p. 92. Secondary
centre developed in Centre 5.
CENTRAL ASIAN CENTRE 72
INULA HELENIUM L. (syn. Helenium grandiflorum

Gilib. ). Elecampane, Elfdock, Horseheal, Yellow
starwort. 2n=20. Probably C. Asia. Also wild
westwards to C. and S. Europe. It was cultivated
and had various uses.
TARAXACUM KOKSAGHYZ Rodin (syn. T. b i c o r -

S3? ö/^Tr'
ne Dahlst. ). Kok-saghyz. 2n=16. Turkestan,
USSR. Cultivated in USSR as a rubber crop.
Cucurbitaceae
CUCUMIS MELO L. Melon, Muskmelon, Cante- E
loupe. 2n=24. Probably Africa (p. 110). Secondary
centre in Centre 5 in which a r e found ssp. melo
Pang. : convar. chandalak (Pang. ) Greb., convar.
ameri (Pang.) Greb., convar. zard (Pang.) Greb.,
K
Aegilops squarrosa
ssp. flexuosus (L. ) Greb. (snake melon) (2n=24),
var. t a r r a Pang, and ssp. agrestis (Naud. ) Greb.
(2n=24), var. agrestis Pang. The latter is a weedy
field melon. AEGILOPS TRIARISTATA*
AEGILOPS TRIUNCIALIS*
Datiscaceae
DATISCA CANNABINA L. 2n=22. C. Asia. A herb. SECALE CEREALE L. Weedy and cultivated rye.
Cultivated as a source of yellow dye. 2n=14. The origin of this species is discussed on
p. 82. Secondary centre in E. Iran and Afghanis-
Ebenaceae tan where S. afghanicum (Van. )Roshev. (syn.
S. cereale ssp. afghanicum (Vav. ) Khush) origi-
DIOSPYROS LOTUS L. Caucasian persimmon.
nated. The main stream of cultivated rye spreading
2n=30. Subtropical China (p. 31) in Talysk and
over Europe and Asia comes from the secondary
W. Georgia, USSR and adjacent Iran. Both these
centre (Khush, 1963). Khush based his conclusion
areas form primary centres.
on the pigmented e a r s of all the cultivated rye
varieties and those of the weedy rye types in Af-
Elaeagnaceae ghanistan. They occur in grain fields with a habit
ELAEAGNUS ANGUSTIFOLIA L. (var. E. argen- and growth rhythm similar to wheat (Stutz, 1972).
ta Moench). Silverberry, Russian olive. 2n=12, Var. eligulatum Vav., the liguleless rye was
28. From S. Europe to C. Asia, China and Hima- found by Vavilov in this secondary centre.
laya. P r i m a r y centre in C. Asia. Cultivated there
and in Iran for its edible nuts. SECALE TÜRKESTANICUM*
E. orientalis L. has been included as var.
orientalis (L. ) O. Kuntze. in this species. TRITICUM AESTIVUM (L. )Thell, ssp. compactum
(Host.)MK. (syn. T. compactum Host. ). Club
Gramineae wheat. 2n=42, genome formula AABBDD. Ssp.
compactum developed in the mountains of Hindu-
AEGILOPS CAUDATA* Kush.
AEGILOPS CYLINDRICA* TRITICUM TURGIDUM ssp. turgidum conv. polo-
nicum (L. ) MK. 2n=28, genome formula AABB.
AEGILOPS JUVENALIS* The type 'T. ispahanicum Heslot' was found in
Ispahan, Iran where it has adapted to irrigated
AEGILOPS KOTSCHYI* cultivation. It is marked by its narrow and elon-
gated glumes.
AEGILOPS LORENTII*
Hippocastanaceae
AEGILOPS OVATA*
AESCULUS HIPPOCASTANUM*
AEGILOPS SQUARROSA auct. non. L. (syn. Ae.
tauschii Cosson, Triticum tauschii (Coss.) Juglandaceae
Schmalh.). 2n=14, genome formula DD. E. Turkey, JUGLANS REGIA L. Walnut, Persian walnut,
Iraq and Crimea and Caucasus, USSR (Zohary et English walnut. 2n=32, 36. P r i m a r y centre:
a l . , 1969) in the west, and Pakistan and Kashmir Region 5. Secondary centre: Moldavia, SE. E u r o -
in the east. P r i m a r y centre in the South Caspian pe and SW. Europe (p. 135). Many varieties have
area. This wild species is the D genome parent been described.
of T. aestivum*. Often as a weed in wheatfields.
Useful as a source of genes for wheat improvement.
ALLIACEAE - MORACEAE 73
V. faba may have derived from the weed V.

angustifolia L., (2n=14) or V. narbonensis*. The
field bean is divided into three varieties according
to the size of the seed: var. minuta (Alef. ) Mansf.
(syn. var. minor (Pieterm. ) Harz., )the pigeon
bean, var. equina P e r s . , horse bean and var.
faba (syn. var. major Harz. ), broad bean.
From its centre of domestication the field
bean spread to Europe, China and the Mediterra-
nean region. In the Mediterranean region a secon-
dary centre of diversity arose (p. 103). The pigeon
and horse beans are used as animal food, while
the broad bean is a vegetable.
Schultze-Motel (1972) has listed all data on
archeological remains of the field bean. Most of
these remnants belong almost exclusively to var.
minuta. He did not find a sharp distinction b e -
tween long-seeded and roundish types, so the d e -
velopment of the field bean from two original
forms is not very probable. Only once was var.
Juglans regia faba, the broad bean found i. e. in Iraq. It dated
ca. 1000 AD, which may point to a late develop-
ment of the large size of the seed.
Labiatae
Liliaceae
LALLEMANTIA ROYLEANA (Wall. &Benth.)
Benth. 2n=14. From Iran to Himalaya. Cultivated FRITILLARIA IMPERIALIS L. Crown imperial.
in E. India as an oil crop. 2n=24. Iran and Turkestan. The bulbs were a
source of starch and used in medicine. It is an
Leguminosae ornamental.
CICER ARIETINUM L. Chickpeas, Gram. 2n=14, Linaceae
16, (24, 32, 33). Secondary centre in Afghanistan.
From this area the 'Kabuli' types of India derive. LINUM USITATISSIMUM L. Flax, Linseed. 2n=30,
They were introduced there about 1700 (van der (32). Origin of flax is discussed on p. 87. P r i m a -
Maesen, 1972). ry centre of origin probably in Centre 5 (Vavilov,
1957). This conclusion is based on the great d i -
CICER MICROPHYLLUM Royle (syn. C. songari- versity of flax in India and adjacent northerly area.
cum Steph., C. jaquemontii Jaub. &Spach. ). Spread from Centre 5 into Centre 4 (p. 67).
2n=14. Tibet, Afghanistan and W. Himalaya. Cul- In the mountains of C. Asia the 'curly oil
tivated in W. Himalaya for its seeds. flax' was developed. It is characterized by the
large number (140-150) of seed capsules per plant.
LATHYRUS SATIVUS L. Grass pea, Chickling pea.
2n=14. Probably W. Asia. A centre of diversity in Malvaceae
the Mediterranean centre. Cultivated in Europe GOSSYPIUM BARBADENSE L. Sea Islands cotton.
since ancient times and in Centre 7 (p. 101). Un- 2n=52, genome formula (AADD)2. Peru. Spread to
known wild. reach Africa and Asia in historical times. Secon-
dary centre in Turkmenia - Tadzhikistan - S.
MEDICAGO SATIVA L. Lucerne. Blue alfalfa. Uzbekistan, USSR.
2n=16, genome formula SS, 32, genome formula
SSSS. Transcaucasia (p. 86). The centre of diver- GOSSYPIUM HERBACEUM L. Short staple Ame-
sity of the blue lucerne is in NW. Iran and adjacent rican cotton. 2n=26, genome formula AÂj.
regions up to Tibet. S. Africa (p. 121). Introduced into Ethiopia, S.
Arabia and Belutchistan where race acerifolium
MEDICAGO TIANSCHANICA Vass. Tien-shan (p. 88) developed (Hutchinson, 1962).
lucerne. 2n=32, 36. Tien-shan, USSR. In USSR
attempts have been made to domesticate it. Meliaceae
MOGHANIA VESTITA (Benth. ex Baker) O. Kuntze MELIA AZEDARACH L. China berry, Bakayan,
(syn. Flemingia vestita Benth. ex Baker). Soh- Pride of India, Persian lilac, Bead t r e e . 2n=28.
phlong. 2n=22. W. Himalayas, N. India. Cultivated SW. Asia up to W. China. Cultivated in the t r o -
for its edible tubers especially in Assam, E. India. pics as an ornamental and shade t r e e . Seeds p r o -
duce oil. They a r e also used as beads.
VICIA FABA L. (syn. Faba vulgaris Moench).
Field bean. Broad bean, Horse bean, Pigeon bean, Moraceae
Tick bean, Windsor bean. 2n=12, 14. Probably MORUS NIGRA L. Black mulberry. 2n=(89-106),
SW. Asia. Broad bean is unknown wild. 308. C. Asia. Cultivated at higher altitude in the
tropics (Purseglove, 1968), and also throughout centres: W. Kopet-Dagh and W. Tien-Shan. Ex-
S. Eurasia for its fruits. The Black Persian mul- tensively cultivated in S. Europe and California.
berry is probably a variety (Purseglove, 1968). In this centre and in Centre 6 (p. 89) other Amyg-
dalus species a r e found. In Georgia, A. georgica
URTICA CANNABINA L. 2n= . C. and N. Asia. Desf. (2n=16) is found. In Kopet-Dagh and Badkhyz
Cultivated for fibre production. A. turcomanica Lincz. occurs. In Armenia A.
nairica Fed. &Takhi. and A. urartu* with A. fenz-
Oleaceae liana a r e native. In this area and in Nakhichevan
JASMINUM OFFICINALE L. Common white j a s - A. fenzliana* (syn. Prunus fenzliana Fritsch)
mine. 2n=26. From Iran to Kashmir and China. (2n=16) grows. This species is very cold r e s i s -
Cultivated especially in S. France for its flowers tant and c r o s s e s freely with cultivated species.
which contain essential oil used in perfumery. Further A. scoparia Spach. (syn. Prunus scoparia
(Spach) Schneid. ) (2n=16) is found in Kopet-Dagh
Papaveraceae and Iran.
PAPAVER SOMNIFERUM L. Opium poppy. 2n=22. AMYGDALUS PERSICA L. Peach. 2n=16. P r i m a r y

P r i m a r y centre either in Centre 5 or in the Me- centre: China (p. 36). Secondary centre in Iran and
diterranean centre (p. 104). The cultigen ssp. som- C. Asia.
niferum (incl. ssp. hortense (Hussenot) Corb. ) is
suggested to derive from ssp. setigerum (DC) AMYGDALUS PETUNNKOWII Litvin. (syn. Prunus
Corb. (syn. P . setigerum DC. ). In Centre 5 the petunnikowii (Litvin. )Rehd. ). Turkestan almond.
greatest variability is observed. In Turkestan the 2n=16. W. Tien-Shan, in Kazakhstan, Tadshikistan
actual domestication could have taken place. and Uzbekistan and partly in Kirgizistan. USSR.
Another classification of subspecies i s : It is an ornamental. It has a high oil content. It is
1. ssp. subspontaneum N. Bas. Asia Minor, Iran, drought resistant. It easily crosses with A. com-
Afghanistan, India and Europe. It has dehiscent munis*.
fruits,
2. ssp. tianshanlcum N. Bas. N. Kirghiz regions. AMYGDALUS SPINOSISSIMA Bge. (syn. Prunus
It also has dehiscent fruits, spinosissima (Bge.) Franch. ). Thorny peach brush.
3. ssp. persicum N. Bas. Iran, Afghanistan, India, 2n=16. C. Asia from Kopet-Dagh through P a m i r -
Egypt and C. Asia. It has dehiscent fruits, Alai to W. Tien-Shan, in Iran, Afghanistan and
4. ssp. turcicum N. Bas. Anatolia, Turkey. It Kurdestan. It is late flowering and has a high oil
has indéhiscent fruits, content,
5. ssp. chinense N. Bas. China. It has indéhiscent
fruits, AMYGDALUS TANGUTICA Korsh. (syn. Prunus
6. ssp. tabargataicum N. Bas. the mountains of dehiscens Koehne). Tangut Plum. 2n= . W.
Tabargataï. W. China. It has indéhiscent fruits, China. Cultivated in some parts of China for the
7. ssp. songaricum N. Bas. W. Europe, USSR kernels.
and W. China. It has indéhiscent fruits.
AMYGDALUS ULMIFOLIA (Franch. ) M. Pop.
Phytolaccaceae (syn. Prunus triloba Lindl. ). 2n=16. C. Asia
PISTACIA VERA L. Pistachio. 2n=30. It grows on
mountain slopes and sometimes forms thin forests. AMYGDALUS VAVILOVII M. Pop. (syn. Prunus
Cultivated in Centre 5, in the Mediterranean r e - vavilovii M. Pop. ). Vavilov almond. 2n=16. Kopet-
gion and elsewhere for its seeds. In Turkmenia Dagh, W. Tien-Shan and Pamir-Alai. At one time
there a r e relic populations which formerly b e - it was believed that this species was a hybrid b e -
longed to one large area of this species (Kabulov, tween A. spinosissima* and A. communis*.
1969).
ARMENIACA DASYCARPA (Erhr. ) Borkh. (syn.
Polygonaceae A. atropurpurea Lois., Prunus dasycarpa Ehrh. ).
Purple apricot, Black apricot. 2n= . Unknown
RHEUM RHAPONTICUM L. Rhubarb. 2n=44. SE. wild. Cultivated in C. Asia, Transcaucasia and
USSR. Cultivated as a vegetable. Probably one of Iran. The fruits a r e very sour and may be used
the parents of R. hybridum*. Related to R. palma- for marmelades etc. It might be used as a source
tum*. of late flowering, and of cold resistance of the
flower buds.
Rosaceae
AMYGDALUS BUCHARICA Korsh. (syn. Prunus ARMENIACA VULGARIS L. Apricot. 2n=16. P r i m a -
bucharica (Korsh. ) Fedtsch. ). Bokhari almond. ry centre in NE. China (p. 36). Secondary centre
2n=16. W. Tian-Shan and Pamlr-Alai, in Afgha- for the cultivated apricot E. Tien-Shan. For the
nistan. It may form a source of sweet pits, of wild apricot this is a primary gene centre which
high oil content and of a good kernel to shell ratio. was formerly connected with the main part in
China (p. 36).
AMYGDALUS COMMUNIS L. (syn. Prunus amyg-
dalus Batsch. ). Almond. 2n=16. W. Kopet-Dagh, CRATAEGUS AZAROLUS L. (syn. C. aronia
Afghanistan and W. Tien-Shan (p. 89). P r i m a r y Bosc. ). Azarolier. 2n= . S. Europe, Africa
MORACEAE -TAMARICACEAE 75
(p. 105) and the Orient. In Uzbekistan, a l a r g e - MALUS SYLVESTRIS (L. ) Miller. 2n=34. Much of
fruited type, var. turcomanica Popoff, is found. Europe, in Transcaucasia and probably into W.
It is poor in vigour. Turkestan. In the Caucasus this species is not
always distinct from M. pumila*. It is very win-
FRAGARIA BUCHARICA Losinsk. Bokhara s t r a w - terhard. Used as a rootstock. Where it grows t o -
b e r r y . 2n= . Centre 5. gether with M. pumila, hybrids occur and therefore
M. sylvestris must have been involved in the r e -
MALUS KIRGHIZORUM Al. &Fed. 2n= . W. mote origin of the cultivated apple. One subspecies,
Tien-Shan found in the underbush of wild walnut ssp. praecox (Malus praecox (Pall. ) Borkh. ) is
(Juglans regia L. ). P r i m a r y centres a r e in the early maturing. P r i m a r y centre in C. Asia.
basins of the Pskem, Ugam, Kok-Su and other Occasionally cultivated in N. Africa (Uphof, 1968).
r i v e r s . It is a polymorphous species. It is likely
that it introgressed into the cultivated apple (Ma- PRUNUS subgen CERASUS P e r s . In Centre 5 wild
lus pumila*). cherries with small fruits (sect. Microcerasus
Webb. ) occur.
PRUNUS CERASIFERA Ehrh. Cherry plum, Myro-

balan. 2n=16, genome formula CC, (24, 32, 48).
P r i m a r y centre is in the Caucasus (p. 89). Secon-
dary centre: the W. Tian-Shan.
It is characterized by high yield, early flowe-
ring, early ripening, wide adaptability and high
acid content.
PRUNUS FERGANICA Lincz. Ferghana plum.

2n= . Ferghana ridge in Tian-Shan and in
Pamlr-Alai (USSR). It is a true-breeding hybrid
of a spontaneous cross Amygdalus communis* and
Prunus cerasifera*.
PYRUS BUCHARICA Litv. 2n= . W. Tian-Shan

and the mountains of Tadjikistan. It is thought to
be a hybrid of P. regelii* and P. korshinskyi*
(Vavilov, 1930).
PYRUS KORSHINSKYI Nak. &Kik. 2n=

Malus kirghizorum Pamir-Alai and in W. Tian-Shan. It is considered
as one parent of P. bucharica*.
MALUS PUMILA* PYRUS REGELII Rehd. 2n= . W. Tian-Shan,

Pamir-Alai and the Bokhara Uplands. It is very
MALUS SIEVERSn (Ledeb. ) M. Roem. 2n= resistant to drought.
W. Tian-Shan and in Alatan of Jungar in the under-
bush of the wild walnut (Juglans regia L. ). PYRUS SOGDIANA S. Kudr. 2n= Shakhrisibai
region.
PYRUS VAVILOVII M. Pop. 2n= . E. F e r g h a -

na. It is considered as a hybrid of P . communis*
x P . korshinskyi* (Vavilov, 1930).
ROSA MOSCHATA J. Hermann. 2n=14, (28).

Himalaya and Iran. Cultivated as ornamentals. It
is a parent of R. x bite r a*.
Salicaceae
SALIX ALBA L. (syn. S. aurea Salisb. ). White
willow. 2n=76. Europe (p. 141). Asia and N. Afri-
ca. Cultivated in the Kashmir for lopping for
fodder (Heybroek, 1963).
Tamaricaceae
TAMARIX GALLICA L. Tamarisk, 2n=24. W.
Malus sieversii Himalayas and NW. India. Cultivated for shelter
and as an ornamental.
Ulmaceae
ULMUS VILLOSA Brandis ex Gramble. 2n=

Himalayas. Cultivated in sacred places or for
ornamental purposes. It is lopped for fodder
(Heybroek. 1963).
ULMUS WALLICHIANA Planch. 2n= . Large-
leaved elm. Himalayas. Cultivated in Kashmir
and lopped for fodder (Heybroek, 1963).
Umbelliferae
CUMINUM CYMDMUM*
DAUCUS CAROTA L. Carrot. 2n=18. The genus

Daucus occurs mainly in South-West Asia and the
Mediterranean countries but some species are
found in the New World, tropical Africa, Australia
and New Zealand.
D. carota includes many wild (ssp. carota)
and cultivated forms p . sativus (Hoffm. ) Roehl.)
which occur in Europe, N. Africa, SW. and C.
Asia. P r i m a r y centre of the oldest cultivated
forms is in Afghanistan. These types are c h a r a c -
terized by carrots purple coloured with anthocya-
nins. From Afghanistan it spread to Asia Minor.
During its distribution the yellow and white carrots
originated probably through mutation (see however,
p. 142). The white mutants (albus) were used for
fodder and did not participate in the development
of the European carrot (Banga, 1957).
In Asia Minor and during its further distribu-
tion in Europe and Mediterranean countries, intro-
gression occurred with local wild carrots (p. 105).
After having reached Iran it probably spread
from this country to China (Banga, 1962).
Vitadaceae
VITIS VINIFERA L. Common grape. 2n=38. Can-
yons of Tian-Shan and adjacent a r e a s . P r i m a r y
centre for the cultivated grape lies in this centre
(see further p. 90).
6 Near Eastern Centre
">-—•" "ƒ ? Ö
r
V f- "'
> )\
• \ 1
C. """'"-7
5 ^
The Near Eastern Centre was called by Vavilov the Near Eastern Centre of Origin. This last centre in-
cluded a part of Centre V.
Darlington (1956) called this area the SW. Asian region. Zhukovsky (1968) gave it two numbers 5 and
6, but in 1970 he separated both by drawing a line on the map. Zohary (1969) preferred to fuse both r e -
gions into one. In this region lies the Fertile Crescent. Here agriculture may have independently been
developed about 9 000 BC. (Cambal &Braidwood, 1970). Hence, Harlan (1971) called the area of the
Fertile Crescent, a centre Al Near East centre. From this centre agriculture spread to Europe, the
Mediterranean region, Afghanistan, India and possibly Africa.
Important crops a r e fruit t r e e s , Brassica oleracea, Hordeum vulgare, Lens esculenta, Medicago
s p . , Secale s p . , Triticum sp. , Vicia sativa, Vitis vinifera etc.
In Georgia, USSR a secondary centre of diversity developed for Glycine max, Lupinus albus, Phaseo-
lus vulgaris, Setaria italica and Zea mays. Maize entered USSR by way of Georgia.
In Turkey, Harlan (1951) described microcentres for Amygdalus s p . , Cucumis melo, C. sativus,
Cucurbita moschata, C. pepo, Lens esculentum, Lupinus s p . , Malus s p . , Medicago sativa, other annual
Medicago species, Onobrychis viceaefolia, Phaseolus vulgaris, Pistacea s p . , Prunus s p . , Pyrus s p . ,
Trifolium s p . , Vicia faba, Vitis vinifera and Zea mays.
Alliaceae ALLIUM KURRAT Schweinf. Salad leek, kurrat.

2n=32. Probably Arabia and Sinai (Uphof, 1968).
ALLIUM AMPELOPRASUM L. Levant garlic,
The geographical distribution is not known. Culti-
Perennial sweet leek. 2n=16, (24), 32, genome
vated in the Nile region, Arabia and Palestine for
formula AAA'A", (40), 48, genome formula
its leaves (Uphof, 1968). It might be derived from
AAA'A'A"A", AABBBB. Europe, Asia Minor,
A. ampeloprasum* (Kuckuck, 1962). Because of
Caucasus to Iran and N. Africa. A type resembling
its close relationship to A. ampeloprasum* it is
ssp. iranicum P.W. is cultivated near Teheran
often included in this latter species as var. kurrat
(Tahbaz, 1971). At Israel the diploid is r a r e ,
Schweinf. ex Krause.
whereas the triploid is a vegetatively propagated
triploid weed (Kollmann, 1972).
ALLIUM PORRUM L. Leek. 2n=32. Asia Minor.
Probably gene centre for cultivated forms (Kuckuck,
ALLIUM ASCOLINICUM L. Shallot. 2n=16. P r o b a - 1968). Leek is a cultivated form derived from
bly W. Asia. This species is often included in A. ampeloprasum*. In this case it is included as
A. cepa*. var. porrum (L. ) Cav. in the latter species.
NEAR EASTERN CENTRE 78
ALLIUM SATIVUM L. Garlic. 2n=16, genome Compositae

formula SS. Wild type in C. Asia (p. 71). Secon-
dary centre in Region 6 (Kazakova, 1971). CARTHAMUS TINCTORIUS L. Safflower. 2n=24,
genome formula BB.
Boraginaceae Vavilov (1951) and Kupzow (1932) proposed
three a r e a s of origin for the cultivated safflower:
SYMPHYTUM ASPERUM Lepechin (syn. S. a s - in India, based on variability and ancient culture,
perrimum Donn. ). Prickly comfrey. 2n=40 Cauca- in Afghanistan, based on variability and proximity
sia to Armenia and N. Iran. Cultivated as a forage of wild species, and in Ethiopia because of the
crop. It is probably one of the parents of S. X u p - presence of wild safflower species. However
landicum Nyman, (2n=36), a forage crop. The Hanelt (1961), Ashri and Knowles (1960) placed
other is S. officinale L., common comfrey, the centre of origin in the Near East because of
(2n=26, c. 36, 40, c. 40, 48) which is native to the similarity of cultivated safflower to two c l o s e -
Europe, W. Siberia and Asia Minor. ly related wild species: C. flavescens Spreng.
(syn. C. persicus Willd. (2n=24, genome formula
Chenopodiaceae BB), found in Turkey, Syria and Libanon, and
C. palaestinus Eig. (2n=24, genome formula BB),
BETA COROLLIFLORA Zos. 2n=36, genome for-
found in deserts of W. Iraq and Israel (Khidir and
mula CCCC. Iran, Transcaucasia and the Pontian
Knowles, 1970). In this area introgression b e -
range of Asia Minor. It is frost resistant.
tween wild and cultivated safflower may still take
place.
BETA INTERMEDIA Bunge. 2n=36, genome for-
mula possibly LLCC. Plants which a r e probably The great variability of the Afghanistan saf-
hybrids of B. lomatogona* and B. trigyna* have flower population must be caused by the meeting
been described as B. intermedia. They occur of the Middle Eastern and the Pakistani types
where both species grow together. It is a source (Knowles, 1969) (p. 71). The wild safflower of
of resistance to yellow mosaic disease. Ethiopia cannot be the progenitor of safflower in
this centre because it has 32 pairs of chromosomes
BETA LOMATOGONA Fisch. &Mey. 2n=18, while the cultivated species has 12 pairs (Knowles,
genome formula LL (22), 36. Asia Minor and E. 1969). Safflower is or has been cultivated in many
Transcaucasia. It is a weed characterized by areas of the Old World and in North America.
'monogerm fruits'. Where the distribution of this Many improved USA varieties have mainly Sudanese
species and that of B. trigyna overlaps tetraploid material as parent. These varieties are now culti-
B. lomatogona a r e found. The hybrids a r e proba- vated too in Egypt, Spain and some other countries
bly identical to plants described as B. intermedia*. where they may cross with local varieties and so
produce new genotypes or they may replace the
local varieties so that gene material is lost. Cul-
tivated mainly for its flowers which were a source
of pigment. At present it is cultivated for its seeds
which yield an edible oil. Its high level of poly-
unsaturation due to its high content of linoleic acid
makes it very suitable for consumption.
Imrie and Knowles (1970) suggested that C.
palaestinus is a wild species. From this species
the weedy species C. flavescens and C. oxyantha
M. B. (2n=24, genome formula BB) and the culti-
vated species C. tinctorius derive.
CHRYSANTHEMUM COCCINEUM Willd. (syn.

C. roseum Adam). 2n= . Wild in N. Iran,
Caucasia and Armenia. Cultivated as a garden
Beta lomatogona and B. intermedia ( ), B. macrorrhiza (—) plant. The flowers contain the insecticide Pyrethri-
and B. trigyna (...) (Ulbrich, 1934) ne, but its toxity is less than C. cinerariaefolium*.
CHRYSANTHEMUM PARTHENIUM*
BETA MACRORRHIZA Stev. 2n=18. (Sub) alpine
zones of the mountains in Iran, Turkish Armenia Cornaceae
(Lake Van) and the Caucasus. A winterhardy
CORNUS MAS L. (syn. C. mascula Hort. ). C o r -
species with a sugar content (8-12%) and white
nelian cherry. 2n=18, 54. Caucasus and Asia
pulp. Minor as an underbush of deciduous forests. Cul-
tivated for its edible fruits and as an ornamental
BETA TRIGYNA Wald. &Kit. 2n=54, genome
shrub. The fruits a r e also used to produce Vin de
formula LLCCCC. Around the Black Sea with out-
Cornoulle, an alcoholic beverage.
liers to the Caspian Sea, Iran, Ukraine and Hun-
gary.
CHENOPODIUM CAPITATUM (L.)Asch. 2n=16,

18. The Orient. Cultivated (Mansfeld, 1959).
ALLIACEAE - GRAMINEAE 79
Corylaceae Dipsacaceae
CORYLUS AVELLANA L. European hazel, Cob- CEPHALARIA SYRIACA Schrad. Pelemir. 2n=10.
nut, Hazel nut. 2n=22, 28. Europe and Caucasus. This pestweed of wheat fields is occasionally cul-
P r i m a r y centre in the Caucasus. Cultivated wide- tivated on the central-anatolian peneplane as an
ly. oil crop.
In this same centre there is a wealth of other
Corylus species: C. maxima*, C. pontica C. Koch DIPSACUS SATIVUS (L. ) Scholier (syn. D. fullo-
(2n=28), C. colchica Alb., C. iberica Wittm. & num L. ). Teasel. 2n=16, 18. Cultivated in Europe
Kemular, C. imoretica Kemular, C. cervorum and elsewhere. It has developed from D. sylves-
P e t r . and C. colurna*. t r i s Huds. (2n=16, 1 8 ) o r D . ferox Loisel (2n=16,
18).
CORYLUS COLURNA L. Turkish hazel. 2n=28.
Occasionally cultivated in Turkey for its nuts. Fagaceae
CASTANEA SATIVA Mill. (syn. C. vesca Gaertn. ).
CORYLUS MAXIMA Miller. Filbert, White filbert,
Sweet chestnut, Spanish chestnut. 2n=22, 24.
Red filbert. 2n=22, 28. Caucasia, W. Asia and
From Italy northwards to Hungary and eastwards
SE. Europe. Cultivated for its nuts.
up to Asia Minor and W. Georgia, Caucasia. Cul-
tivated for its nuts and timber. Outside the above
Cruciferae
range it is naturalized.
BRASSICA OLERACEA L. Wild and cultivated
cabbages. 2n=18, genome formula CC. In Asia Gramineae
Minor varieties belonging to convar. oleracea*,
convar. capitata* and convar. acephala* a r e AEGILOPS CAUDATA L. (syn. Triticum dicha-
common. sians (Zhuk. ) Bowden, T. caudatum (L. ) Godr. &
Gren. ). 2n=14, genome formula CC. Greece,
Turkey, Iraq and Afghanistan. Its cytoplasm has
CAMELINA SATIVA (L. ) Crantz. False flax.
a male sterilizing action on the T. aestivum*
2n=40. In SE. Europe and SW. Asia the wild parent
nucleus.
form occurs probably C. microcarpa Andrz.
(2n=40). It became a weed in cereal crops and
flax. Later it was cultivated for its oily seeds, the AEGILOPS COLUMNARIS Zhuk. 2n=28, genome
cultigen being called C. sativa. An intermediate formula C U C U M C M C . Turkey, Iraq, Iran and Cau-
form of this latter species and its wild parent is casus. It is a weed of cultivation and looks quite
C. pilosa (DC) Zinger. Another weed of flax fields similar to Ae. triaristata* (Bor, 1970).
is C. alyssum (Miller) Thell. (2n=40). All these
species have also been grouped in one species AEGILOPS COMOSA*
C. sativa this species being divided into subspecies
microcarpa (Andrz. ) Hegi, sativa, pilosa (DC.) AEGILOPS CRASSA Boiss. (syn. Triticum e r a s -
Hegl and alyssum (Miller) Hegi. Its cultivation sum (Boiss. )Aitch. &Hemsl. 2n=28, genome
has almost disappeared. formula D D M c r M c r , 42, D D D 2 D 2 M c r M c r . Turkey,
Iraq, Iran, Afghanistan and Palestina.
Hybrids between 6x forms and T. aestivum* p r o -
CRAMBE CORDIFOLIA Steven, (syn. C. tatarica
duce a few fertile lOx amphiploid plant. This might
Jacq. ). Tatarian sea-kale. 2n= . Highlands of
be useful for introducing useful genes of Ae. c r a s s a
Asia Minor, India and Ethiopia. The perennial
into wheat.
herb is cultivated for the young leaves.
IRATIS TINCTORIA L. (syn. I canescens DC., AEGILOPS CYLINDRICA Host. (syn. Triticum
I. littoralis Steven, I. taurica Bieb. ). 2n=28. cylindricum C e s . , P a s s . &Gib. ). 2n=28, genome
Most of Europe. Cultivated as a source of dye, formula CCDD. Balkan peninsula, Crete, Turkey,
and therefore probably introduced. Caucasus, S. USSR, Iraq, Iran and Afghanistan.
It is a weed in fallow fields and on hillsides.
Cucurbitaceae
AEGILOPS JUVENALIS (Thell. ) Eig. (syn. Ae.
CUCUMIS MELO L. Melon, Musk melon, Cante- turcomanica Roshev., Triticum turcomanicum
loupe. 2n=24. Africa (p. 110). Secondary centre (Rosh. ).Bowden). 2n=42, genome formula DDCU
arose in Centre 4 in which convar. cassaba C U M-'M-'. Iraq, Iran, Turcomania and Turkestan,
(Pang. ) Greb., cassaba melon, winter melon from USSR.
Asia Minor, convar. cantalupa (Pang. ) Greb.,
cantaloupe melons, convar. adana (Pang. ) Greb., AEGILOPS KOTSCHYI Boiss. (syn. Ae. triuncia-
kilik melons and convar, flexuosus (L. ) Greb., lis var. kotschyi (Boiss. ) Boiss., Triticum kot-
t a r r a melons, adjur melons, snake melons, s e r - schyi (Boiss.) Bowden). 2n=28, genome formula
pent melons are found. C U C U S V S V . N. Africa, Palestina, Iraq, Iran,
Afghanistan and Caucasus.
CUCUMIS SATIVUS L. Cucumber, Gherkin. 2n=14.
India (p. 64). Secondary centre (a xerophytic type) AEGILOPS LORENTII Höchst, (syn. Ae. biuncia-
arose in this centre. H s V i s . , Ae. macrochaeta Shuttl. &Huet., syn.
Triticum macrochaetum (Shuttl. & Huet. ex Duval-
Jouve) Richter, Triticum lorentii (Höchst. ) Zeven). both parent species. At present Ae. sharonensis
2n=28, genome formula C u C u M b M b . S. Europe, is effectively reproductively isolated.
USSR, Turkey, Israel, Iraq and Iran. Maan (1973) concluded from his nucleo-cyto-
The name Ae. lorentii antedates Ae. biuncia- plasmic and cytogenetic studies that the cytoplasm
lis Vis. (Bor, 1970). of Ae. speltoides is closely related to that of T.
timopheevi* and differs from that of T. monococcum*
AEGILOPS MUTICA Boiss. (syn. Triticum t r i p - and T. turgidum*.
sacoides (Jaub. &Spach) Bowden. 2n=28 (+B),
genome formula C U C U CC. Anatolia and Armenia. AEGILOPS SQUARROSA*
Jones and Majisu (1968) consider it related to the
A, S or D genomes (p. 83 and p. 72), but not to AEGILOPS TRIARBTATA Willd. (syn. Triticum
the B genome (p. 84). It is a cross-fertilizer. triaristatum (Willd. ) Godr. &Gren. ). 2n=28, g e -
nome formula C"CUM M4, 42, genome formula
AEGILOPS OVATA L. (syn. Triticum ovatum C u C u M t M t M t 2 M . The Mediterranean region, W.
(L. ) Raspail. 2n=28, genome formula C U C U M°M°. Asia, Iraq, Iran and S. of USSR. The hexaploid is
The Mediterranean region, Palestina, Syria, also described as Ae. recta (Zhuk. ) Chenn.
Turkey, Iraq, Iran, Afghanistan. A weed in c u l -
tivation. Its cytoplasm has a male sterilizing AEGILOPS TRIUNCIALIS L. (syn. Triticum t r i u n -
action on the nucleus of T. aestivum* and T. t u r - cialis (L. ) Raspail. 2n=28, genome formula
gidum*. C U C U CC. The Mediterranean region, Turkey,
Palestina, Syria, Lebanon, Iraq, Iran, Turcomania,
AEGILOPS SPELTOIDES Tausch, (syn. Triticum Afghanistan and Caucasus.
speltoides (Tausch. ) Gren. ex Richter. 2n=14, Ae. bushirica Rosh. might be a synonym. How-
genome formula SS (formerly it was believed that ever Bor (1970) suggested that it could be a hybrid
the S genome was identical to the B genome of product of Ae. triuncialis and Ae. cylindrica*.
T. turgidum* a n d T . aestivum*. However, new
investigations showed that this is not so (Johnson, AEGILOPS UMBELLULATA Zhuk. (syn. Triticum
1972; Kimber & Athwal, 1972). P r i m a r y centre: umbellulata (Zhuk. ) Bowden). 2n=14, genome for-
S. Turkey, N. Syria and N. Iraq. It is less com- mula C U C U . Moist steppes, dry hills and as a weed
mon in the remaining part of Turkey and Thrace, in cultivation in the Greek islands, Turkey, N.
Greece and in Syria, N. and C. Israel. It is often Syria, Iraq, N. and W. Iran and Transcaucasia.
found together with the wild T. boeoticum*. Used in wheat breeding as a source of resistance
This species includes Ae. speltoides s . S . , to leaf rust, Puccinia triticina E r i k s .
Ae. ligustica (Savign.) Cosson and probably Ae.
longissima Schweinf. ex Muschl., genome formu- AEGILOPS VARIABILIS*
la S°S° and is synonymous to Ae. sharonensis
Eig., genome formula S b S" and Ae. aucheri Boiss. AGROPYRON INTERMEDIUM*
Ssp. speltoides is a B-chromosome c a r r i e r and
a cross-fertilizer. AGROSTIS TENUIS*
AVENA SATIVA L. Byzantina type, Red oat, Al-

gerian oat. 2n=42, genome formula AACCDD.
The Mediterranean centre (p. 97). Cultivated in
the south and extreme west of Asia Minor, where
it may come in contact with other Avena species.
CYNOSURUS CRISTATUS*
HORDEUM VULGARE L. Barley. 2n=14, genome

formula VV. The wild parent of barley is H. spon-
taneum C. Koch (2n=14). E. Mediterranean region -
- W. Asiatic countries to Turkmenia and Afghanis-
tan. In the peripheral areas it only grows as a
weed. In the Upper Jordan valley catchments robust
types with very large seeds and very long awns a r e
Aegilops speltoides found. A slender type occurs in dry steppes of
Negev, Transjordan-Turkish border a r e a - Iran -
Afghanistan. In E. Galilee small grassy types a r e
Waines and Johnson (1969) considered Ae. s h a - found while everywhere intermediates a r e observed
ronensis (Triticum x sharonense (Eig. )Waines & (Zohary, 1969).
Johnson as a hybrid of Ae. longissima and Ae. H. spontaneum is two rowed and has a brittle
bicornis*. They based this conclusion on the s i m i - rachis. Its domestication probably took place in
larity of grain-protein electrophoretic patterns of the Fertile Crescent resulting in the two-rowed
this hybrid and that of the summation of its puta- type (ssp. distichon, syn. H. distichon L. ). The
tive parents, the intermediate spike characters, start of its domestication might have been about
the intermediate ecological preference and on its 7 000 BC. Helbaek (1966) suggested that the barley
distribution in Israel: the a r e a of overlap between
GRAMINEAE - GRAMINEAE 81
genotype BtBtbt 2 bt 2 is very common (90-100%) in

Y' ^ .--s1' '
^ (? ö the 'Oriental' region: C. and S. Japan, S. Korea,

China, Tibet and N. Central Nepal (naked forms),
while the genotype btbtBt 2 Bt 2 is common (60-95%)
in the 'Occidental' region: N. Japan, N. Korea,
Manchuria, India, Nepal (hulled forms), Kashmir,
Turkey, Europe, USSR, Egypt and Ethiopia. In
Pakistan, Afghanistan, Iran, Iraq and Syria both
«=. types occur (65-75%) BtBtbt 2 bt 2 ).
C. 7 000 BC. 'naked barley' was first identi-
^ "vc, V
Hordeum spontaneum (Harlan & Zohary, 1969)
fied at Beidha, Aceramic Hacilar and Ali Kosh
(Bus Mordesh period). In this material the first
six-rowed types (ssp. hexastichon, syn. H. hexa-
stichon L. ) appears (Braidwood et a l . , 1969).
Formerly H. agriocrithon Aberg was thought
to be the wild parent of the six-rowed barley. It
has a brittle rachis. It is however a hybrid p r o -
found at Beidha dating c. 7 000 BC was cultivated duct of H. spontaneum and six-rowed barley. They
wild barley. were first observed in Tibet as weeds in barley
Braidwood et al. (1967) found 'evidently do- fields. Later it has also been found elsewhere
mesticated barley' at Cayönli, Turkey, in layers where H. spontaneum comes in contact with culti-
dated c. 7 000 BC. vated six-rowed barley. Lagunculiforme and inter-
Domesticated barley has a tough rachis and medium types (H. lagunculiforme Bakhteyev)
the change of brittle to tough rachis may have have very probably the same origin.
taken place in 7 000 - 6 000 BC. The tough-rachis In Centre 6 ssp. humile* arose and in Centre
is conditioned by two recessive alleles bt and bt„. 7 ssp mediterraneum*.
Both genes a r e closely linked and the geno- Lagunciliforme (H. lagunculiforme) and inter-
type for tough rachis is either BtBtbtobt, or medium (H. intermedium Carlet) have a similar
btbtBt 2 Bt 2 . No plants with btbtbt 2 bt 2 have yet been origin. Zohary (1971) suggested that these brittle
found or bred probably due to the close linkage of types a r e ill-adapted to survive under stable, wild
the genes. Takahashi (1964) observed that the conditions (at least in Israel).
Primary centre ( ), secondary centre ( )and distribution of wild and weedy Secale cereale types and ways of their introduction
and that of rye into Europe and N. Asia (...) (Khush, 1962)
POA BULBOSA L. 2n=14. 28, (39), 42, 45, (40- SECALE MONTANUM Guss. Mountain rye. 2n=14,
58, 56>. A grass of Europe, W. Asia and W. of genome formula R m R m . From the C. Atlas Moun-
N. Africa. Cultivated in USA. tains of Morocco and the Sierra Nevada Mountains
Var. vivipara of C. and Ante-Asia is a very of Spain eastward in isolated pockets in the moun-
valuable forage plant of dry steppes. tains of Sicily, Italy, Yugoslavia. Greece, Lebanon,
Turkey, Iran and Iraq (Stutz, 1972). It is a highly
SECALE ANATOLICUM Boiss. (syn. S. monta- polymorphic perennial, cross-fertilizing form.
neum v a r . anatolieum (Boiss. ) Boiss. ). 2n=14. Some of the isolates have been described as d i s -
Turkey and W. Iran and Iraq. It is a highly poly- tinct species or varieties.
morphic weedy form. S. ciliatoglume (Boiss. ) Grossh. (syn. S.
montanum var. ciliatoglume Boiss. ). A weedy
SECALE CEREALE L. Weedy and cultivated rye. population with pubescent culms in orchards and
2n=14, genome formula R C R C . P r i m a r y centre of vineyards near Mardin, SE. Turkey.
annual and perennial rye species and forms: NE. S. dalmaticum Vis., population growing within
Turkey - NW. Iran. Khush (1963) suggested that a the walls of the old St. Johannis fortress above
secondary centre (p. 72) is in Afghanistan and Kotor, S. Jugoslavia.
that the cultivated rye of Europe and N. and C. S. daralgesii Thum., a weedy form with non-
Asia derives from this centre. Only a few come fragile rachis along roadsides and ditchbanks of
from the p r i m a r y centre. In the primary centre Armenia.
S. vavilovii* and S. montaneum* hybridize with S. kuprijanovii Grossh., a broad-leaved form
each other and introgress resulting in a mixture of mountain meadows of the N. Caucasus Moun-
of genetic variants which could be described as tains.
'S. segetale', 'S. afghanicum', S. daralgesii', S. montanum has the same chromosomal
'S. cereale', 'S. turkestanicum' and 'S. dighori- arrangement as S. anatolieum*, S. silvestre*
cum' (Stutz, 1972). Stutz (1972) suggested that and S. africanum*. Its chromosomal arrangement
from this highly variable population the annual differs from that of S. vavilovii* and S. cereale*
S. cereale types invaded cultivated fields to b e - and from its closely related forms in reciprocal
come weeds. translocations involving 6 of the 14 chromosomes.
S. cereale includes cultivated rye and a num- It is the parental species of S. anatolieum*
ber of weedy rye types which occur in grain fields and S. silvestre* (Stutz, 1972). Owing to hybridi-
and along ditchbanks and roadsides throughout the zation with S. vavilovii* weedy types belonging to
Middle East countries (Stutz, 1972). These weedy S. cereale* arose (Stutz, 1972).
types a r e :
S. afghanicum* SECALE SILVESTRE Host. (syn. S. fragile
S. dighoricum (Vav. )Roshev. (syn. S. c e r e a - Marsch.). 2n=14. C. Hungaria eastward through-
le L. ssp. dighoricum (Vav. ) Khush). It is a weed out the sandy steppes of S. Russia up to W. Siberia
in grain fields of N. Ossetia, USSR. and Pamir-Alaj (Bor, 1970; Stutz, 1972). A low
S. segetale (Zhuk. ) Roshev. (syn. S. cereale growing annual psammophyte with fragile rachis.
L. ssp. segetale (Zhuk. ) Khush. ). This is a poly- It has the same chromosome arrangement as S.
morphic weed in grain fields throughout E. Europe montanum*. It is suggested that this species d e -
and the Middle East (Stutz, 1972). rives from S. montanum and that it is the ancestor
S. ancestrale Zhuk. (syn. S. cereale L. ssp. of S. vavilovii* (Stutz, 1972).
ancestrale (Zhuk. ) Khush). It is a robust, tall (up
to 2.4 m) weed with small, invested seeds and SECALE VAVILOVn Grossh. 2n=14. Common to
fragile rachis restricted to sandy ditchbanks and the lower slopes of Mount Ararat and along the
fence rows near Aydin, SW. Turkey. banks of the Araxis River. It is a wild low growing,
S. turkestanicum Bensin. A self-compatible annual, self-compatible psammophyte with fragile
cultigen of C. Asia (p. 72) and Transcaucasia. rachis. It has the same chromosome arrangement
The weedy types have been derived mainly as S. cereale (see S. montanum*). Bor (1970).
from S. vavilovii* by introgression with S. monta- strongly suspected this species to be the same as
num* and its derivative species S. anatolieum*. S. afghanicum*. Khush (1960) suggested that it
In some places suboptimum for wheat and barley, derived from S. montanum, but Stutz (1972) made
rye may have been developed to become fully do- it clear that this species derives from S. silvestre*
mesticated. It is generally proposed that S. an- and that it is the ancestor of S. cereale*.
cestrale and S. segetale a r e the parental types of
S. cereale; however, Stutz (1972) suggested that TRITICUM AESTIVUM (L ) Thell. Wheat. 2n=42,
S. ancestrale derives from S. cereale. There is genome formula AABBDD. P r i m a r y centre: T r a n s -
no introgression into S. ancestrale of other Seca- caucasia and adjacent a r e a s . There natural cross
le genetic material in spite of its contact with pollination within the species and between sub-
other species, because a high incidence of geno- species, other Triticum-species and related
typically controlled chromosomal breaks in out- species Aegilops and Secale is still taking place.
crossed hybrids leads to sterility (Stutz, 1971). This species is a natural amphiploid of emmer
Hybridization between wheat (p. 82) and rye and Aegilops squarrosa*. This amphiploidization
may have increased the variability of wheat. Rye must have taken place after the development of
is a source of resistance to diseases in wheat e m m e r from its wild ancestor T. turgidum ssp.
breeding and a parent of octaploid and hexaploid dicoccoides*. There is a continuous exchange of
triticales. genes from wild species to cultivated and vice v e r -
sa. This has resulted in numerous botanical two awns and a winter habit.
'varieties'. In the Fertile Crescent the wild einkorn was
This hybridization has led to hexaploid types domesticated to become ssp. monococcum, which
with a very brittle ear. This is a 'wild' character has a tough rachis.
and it could be concluded that there a r e wild hexa- Its earliest appearance is from Ali Kosh
ploid wheats. However these segregants can not dating c. 6 500 BC., thus later than the first
really be called wild. appearance of T. turgidum ssp. dicoccum.
The main division of T. aestivum is into ssp. Einkorn was spread over Europe, N. Africa,
spelta (L. ) Thell. (syn. T. spelta L. ), spelt, ssp. Asia Minor, Caucasus, Iraq and Iran. Cultivated
vavilovi (Tum.) Sears (syn. T. vavilovi (Turn.) in some areas as a fodder crop. It forms a com-
Jakubz. ), ssp. macha (Dek. &Men. ) MK. (syn. ponent of the Zanduri wheat which is a mixture of
T. macha Dek. &Men.), makha wheat, ssp. vul- einkorn, T. timopheevi ssp. timopheevi* and T.
gare (Vill. ) MK. (syn. T. vulgare Vill. ), common zhukovskyi*. This population is cultivated in
wheat, bread wheat, ssp. compactum (Host) MK. Georgia, USSR. This species forms an important
(syn. T. compactum Host., club wheat, and ssp. source of disease resistance.
sphaerococcum (Perc. ) MK. (syn. T. sphaero- There is still a natural gene flow between
coccum P e r c . ), Indian dwarf wheat. The origin diploid and tetraploid species (Vardi & Zohary,
of some subspecies has not yet been classified. 1967).
They have originated in another centre.
Spelt wheats have been found in Iran, in Cen- TRITICUM TIMOPHEEVI Zhuk. 2n=28. genome
tral Europe (p. 134) and Africa (p. 117). formula AABB (or AAB'B', AAGG). This species
Ssp. vavilovi wheat is indigenous to Armenia. consists of two subspecies ssp. araraticum
It is characterized by its branching spikelet. (Jakubz. ) Mac Key (syn. T. araraticum Jakubz. )
Makha wheat is indigenous to W. Georgia. It and ssp. timopheevi. Ssp. araraticum grows wild
is often mixed with T. turgidum ssp. paleocolchi- in Transcaucasia, N. Iraq, W. Iran and E. Turkey.
cum*. It was first described as T. dicoccoides ssp.
Bread wheat is now widespread. P r i m a r y armeniacum Jakubz. and as T. armeniacum Mak.
centre in Transcaucasia and adjacent regions. Some types closely resemble T. dicoccoides*.
Secondary centres in Hindukush and adjacent r e - However, this subspecies only crosses easily with
gions (p. 66), in China and Japan (p. 33) and p r o - ssp. timopheevi.
bably in African Sahara (p. 117). Ssp timopheevi is a part of the Zanduri wheat,
Club wheat developed in Afghanistan and adja- which is cultivated in Georgia, USSR. It is an an-
cent regions (p. 72) and probably in Switzerland/ cient cultivated wheat. Its rather brittle rachis
Austria (p. 134). A secondary centre of diversity causes difficulties in threshing. It is difficult to
is in Armenia. cross with other Triticum species; it is a source
Indian dwarf wheat originated in NW. India of disease resistance and (timopheevi) durum and
and adjacent regions (p. 66). Its presence has aestivum plants often are male sterile.
been reported in N. Africa. Maan (1973) concluded from his nucleo-cyto-
Dorofejev (1971) suggested that ssp. macha plasmic and cytogenic studies that T. timopheevi
is the oldest hexaploid. From this subspecies ssp. ssp. timopheevi and ssp. araraticum and T. d i -
vulgare developed. Ssp. spelta and ssp. vavilovii coccoides var. nudiglumis ex Turkey-Iran-Iraq
a r e secondary spelts. They may have been derived area have the same type of cytoplasm and the g e -
from ssp. vulgare. nome formula AAGG. The cytoplasm of Ae. spel-
toides* is closer to the above cytoplasm than to
TRITICUM AESTIVUM (L. ) Thell. ssp. compac- T. turgidum*-cultivated wheats.
tum (Host) MK. (syn. T. compactum Host. ).
Club wheat. 2n=42, genome formula AABBDD. TRITICUM TURGIDUM (L. ) Thell. Wild emmer
This subspecies developed in the Hindu-Kush wheats. 2n=28, genome formula AABB. The d i s t r i -
(p. 72). Secondary centre in Armenia. bution area of the wild ssp. dicoccoides can be
divided into two regions. One northern region is
TRITICUM MONOCOCCUM L. Wild and cultivated S. Turkey - Iran - Iraq where var. nudiglumis is
einkorn. 2n=14; genome formula AA. The wild found, the Southern region is Israel, S. Syria and
ssp. boeoticum (Boiss. ) Mac Key (syn. T. boeoti- Jordan. The cytoplasm of var. nudiglumis is s i m i -
cum Boiss. ) includes the types aegilopoides lar to that of T. timopheevi*, while that of ssp.
(T. aegilopoides (Link) Bal. and thaoudar (T. dicoccoides of the southern region is the same as
thaoudar Reut. ); furthermore 'T. spontaneum the cultivated T. turgidum* (Maan, 1973). Ssp
Flaksb. ' and 'T. urartu Tuman. '. It is spread over dicoccoides (Körn. ) Thell. (syn. T. dicoccoides
Greece, Turkey, Syria, N. Iraq and Transcauca- Körn. ). is derived from a natural amphidiploidi-
sus. Aegilopoides is characterized by one grain zation of an unknown diploid species and T. mono-
and one awn per spikelet, while thaoudar has two coccum ssp. boeoticum*. This amphidiploidization
grains and two awns. There a r e two distribution must have occurred probably in the Syrio-Pales-
centres: in the Fertile Crescent and in Turkey. tine probably at several places. Much research
In peripheral areas it is segetal. In the first has been carried out to identify the unknown diploid
centre thaoudar is found. Aegilopoides type occurs parent. Until 1971 Ae. speltoides* was widely
in the cooler Balkans and W. Anatolia. In Anatolia accepted as the source of the B genome while ssp.
intermediate types and mixtures a r e found. In A r - boeoticum was the A genome donor. However, r e -
menia the type urartu has been described. It has sults of new research showed that Ae. speltoides*
Dek. ). Georgian emmer, Kolchic emmer, s s p .

turgidum including conv. turgidum, Poulard wheat,
English wheat, conv. durum (Desf. ) Mac Key (syn.
T. durum Desf. ), durum wheat, hard wheat, conv.
turanicum (Jakubz. ) Mac Key (T. turanicum
Jakubz., T. orientale P e r c . ), khorassan wheat,
conv. polonicum (L. ) Mac Key (syn. T. polonicum
L. ) Polish wheat, and ssp. carthlicum (Nevski)
Mac Key (syn. T. carthlicum Nevski, T. persicum
Vav. ex Zhuk. ), Persian wheat. They have origi-
nated through cultivation from ssp. dicoccoides*
and maybe by intergeneric and interspecific hybri-
dization.
P r i m a r y centre in 6. Secondary centres in
Wild t e t r a p l o i d T r i t i c u m s p e c i e s : T r i t i c u m tugidum s s p . d i c o c - Ethiopia (p. 117) and in the Mediterranean centre
coides (1, 2, 3), spp. dicoccoides v a r . nudiglumis (3) and T.
timopheevi s s p . a r a r a t i c u m (4) (Johnson, 1972)
(p. 99).
E m m e r is the oldest cultivated wheat. Its cul-
tivation is declining. Until recently it was culti-
cannot be the parent (Johnson, 1972; Kimber and vated in Ethiopia (p. 117), Iran, E. Turkey, T r a n s -
Athwal, 1972). Kimber and Athwal (1972) have caucasia, Volga-area, Yugoslavia, Czechoslovakia
speculated about the most likely origin of the B and India. Its domestication may have taken place
genome. They concluded that the tetraploid wheats c. 8 000 BC. in the 'Fertile Crescent Belt'. The
a r e polyphyletlc in origin i . e . they are hybrids of first appearance of domesticated emmer dates
amphiploids which originated as a result of hybridi- from c. 7 000 BC. at Aceramic Neolithic Beidha,
zation between s s p . boeoticum and the other species. Ali Kosh, Jericho and Ramad. Already in 4 000 -
3 000 BC. it had reached the Atlantic coast from
Scandinavia to Spain and also the Nile Delta. Hel-
baek (1960) was surprised by the uniformity of
emmer. However, plants a r e called dicoccum
when they correspond to a certain morphological
description. It should be investigated whether the
idiotypes of the various dicoccum populations also
correspond to each other for characters other than
morphological ones.
The disease resistant khapli (khapli is the
vernacular name of emmer) of India and Yaroslav
emmer from USSR belong to ssp dicoccum.
Durum wheat is cultivated over a large area.
It is cultivated in the Mediterranean coastal region,
in Ethiopia where a secondary centre (p. 117).
exists, and in areas north of the Black Sea. It is
T r i t i c u m boeoticum (Harlan &Zohary, 1969)
rarely observed in wheats of Iran and Afghanistan.
It is also cultivated in the Americas and elsewhere.
The spread in the Old World is shown by Ciferri
Furthermore, Maan (1973) concluded from his (1939). It is the second most important wheat in
nucleo-cytoplasmic and cytogenetic studies that the world.
possibly the cytoplasm of T. turgidum* is not d e -
rived from ssp. boeoticum, Ae. speltoides* and
Ae. bicornis*. The latter species has also been
mentioned as the possible source of the B genome.
From ssp. dicoccoides several cultivated
subspecies have been derived. These a r e dicoccum*
paleoeolchicum*, turgidum* and carthlicum*. The
ssp. turgidum includes conv. turgidum, conv.
durum, conv. turanicum and conv. polonicum.
Ssp. dicoccoides is an important source of disease
resistance; it also c a r r i e s a r e s t o r e r gene of
(timopheevi) cytoplasmic male sterility.
TRITICUM TURGIDUM (L.) Thell. Cultivated

emmer wheats. 2n=28, genome formula AABB.
The cultivated tetraploid wheat can be subdivided
as follows (Mac Key, 1966): ssp. dicoccum
(Schrank) Thell. (syn. T. dicoccum Schubl. ) . ,
emmer, ssp. paleoeolchicum (Men.) Mac Key Spread of durum wheat in the Old World (Ciferri, 1939)
(syn. T. paleoeolchicum Men., T. georgicum
Turanicum wheat originally involved as an o a s i s - in the centres 4 (p. 67), 5 (p. 73) and 7 (p. 101).
ecotype. Its cultivation is restricted to irrigated Cultivated in S. Europe and N. Africa from the
fields (Mac Key, 1966). Mac Key suggested a wide Atlantic eastwards, the Nile Delta and Ethiopia
occurrence in Asia, but Kuckuck (1970) and Bor and northwards and eastwards up to NW. Burma,
(1970) limited the spread to Iran and Iraq. Kuckuck W. China, Kazachskaya USSR. Some of the Cicer
(1970) suggested that it is a hybrid of durum x species indigenous to Anatolia may have played a
polonicum. role in its ancestry, particularly C. pinnatifidum
Polonicum wheat, marked by its long glumes Jaub. &Spach., (2n=16), (from Anatolia, Armenian
and kernels was cultivated in S. Europe, Turkey, SSR., Syria, N. Iraq and Cyprus), C. echinosper-
Iraq, Iran. Afghanistan and NW. India. According m u m P . H . Davis, 2n= , (from E. Anatolia)
to Kihara et al. (1956) it includes T. ispahanicum and C. bijugum K.H. Rech., 2n=16 (from SE.
Heslot. Anatolia, N. Syria, and N. Iraq) (van der Maesen,
Carthlicum wheat is characterized by the p r e - 1972).
sence of the Q (vulgare) gene. It is not known Race orientale Pop. is characterized by its
whether this gene arose independently in this wheat very small seeds (1000 seed weight 100-120 g). It
or came from vulgare wheat. Carthlicum wheat is common in Ethiopia, Sudan, Egypt, India, P a -
was cultivated in Iraq, Iran and Caucasia. It is a mir, Tadzhikistan and Iran. Those from Ethiopia
source of disease resistance. a r e black-seeded.
Georgian emmer, Kolchic emmer (ssp. paleo- Race asiaticum Pop. has somewhat bigger,
colchicum) was formerly cultivated in a mixture but still small seeds (1000 seed weight 140-200 g).
with T. aestivum ssp. macha* in W. Georgia, It occurs in C. Asia, Afghanistan, W. China,
USSR. Iran and E. Turkey.
English wheat (conv. turgidum) was at one Race eurasiatieum Pop. has medium large
time cultivated in Europe and elsewhere. From seeds, (1000 seed weight 200-300 g). It is cultivated
time to time it was reintroduced into cultivation in the Near East, Armenia, Azerbaydzan, Ukraine
because its branching habit (Osiris wheat, Wonder up to C. USSR, and near large cities in the eastern
wheat) convinced farmers that its yield must be part of the area of cultivation. The seeds a r e white.
high. Race mediterraneum Pop. has the largest
seeds (1000 seed weight over 350 g). It is found in
TRÏTICUM ZHUKOVSKYI Men. &Er. Zanduri. Spain, Italy, Morocco, Algeria, Tunesia andW.
2n=42, genome formula AAAABB. It was cultivated Turkey. The seeds a r e white.
in W. Georgia, USSR. It was composed of einkorn,
T. timopheevi ssp. timopheevi* and T. zhukovskyi. GALEGA ORIENTALIS Lam. 2n=16. Caucasia.
The latter is a hexaploid but its genome formula Cultivated as a fodder and as an ornamental.
differs from those of the common hexaploid sub-
species. It is a natural amphiploid of s s p . t i m o - LENS ESCULENTA Moench (syn. L. culinaris
pheevi and einkorn. Its cytoplasm has an identical Medik., Ervum lens L. ). Lentil. 2n=14. Zohary
male sterilizing action onthe durum and aestivum (1972) suggested L. orientalis (Boiss. ) Hand. -
nuclei as ssp. timopheevi. It c a r r i e s genes for Mazz. as the wild parent of lentil. It looks like a
stem rust and mildew resistance. miniaturized lentil. It grows wild in Centre 6.
In ancient times it was cultivated in Egypt,
ZEA MAYS L. Maize. 2n=20. Secondary centre in S. Europe and W. Asia.
the Near East (Brandolini, 1970). Domesticated The p r i m a r y gene centre of lentil is W. Asia.
in C. America (p. 166). Flint maize - indurata Spread to other parts of Europe, to India, and
Sturt. is common h e r e . China and Ethiopia. Vavilov (1949-50) suggested
that L. lenticula (Schreb. )Alef., L. nigricans
Iridiaceae (M.B.) Godr., L. kotschyana (Boiss. )Alef. or
L. orientalis Hand, is the progenitor of lentil.
CROCUS SATIVUS*
They grow wild in this centre. He classified three
varieties: v a r . macrosperma Baumg. from the
Labiatae
Mediterranean region, var. syrica Barul. a m e -
LALLEMANTIA IBERICA Fisch. &Mey. Lalle- dium-sized seed form from the inner mountainous
mantia. 2n=14. Asia Minor and some regions of region of Asia Minor and var. afghanica Barul.,
USSR. Cultivated in Iran and S. USSR for its oil a small seeded form of the highlands of Afghanis-
seeds. tan. Another division is ssp. maçrosperma
(Baumg. ) Barul. and s s p . microsperma (Baumg.)
Leguminosae Barul. The latter is found in all prehistoric exca-
vations (van Zeist &Bottema, 1971).
ALOPHOTROPIS FORMOSUM (Boiss.) Lamprecht
(syn. Pisum formosum (Stev. ) Boiss., Vavilovia Agro-ecological groups meet each other in
formosa (Stev.) Fed.). Wild perennial pea. 2n=14. Turkey and here microcentres have developed
Alpine and subalpine zones of Asia Minor, T r a n s - (Harlan, 1951).
caucasia, Armenia and Iran.
MEDICAGO CANCELLATA Prod. 2n=48. Cauca-
CICER ARIETINUM L. Chickpea, Gram, Garban- sus, in the Stavropol elevation. A wild perennial
zos. 2n=14, 16, (24, 32, 33). Unknown wild. hexaploid species. It has a rhizome.
Secondary centres of diversity probably developed
MEDICAGO DAGHESTANICA Rupr. 2n=16. Cauca- Because of the extreme variability of this
sus. A wild perennial alpine species. It is a good species, its taxonomy is not yet fully defined. The
fodder plant. following subspecies have also been put on a s p e -
cies level. Ssp. ambigua (Trautv. ) Tutin = M. fal-
MEDICAGO DZHAWAKHETICA Bordz. 2n=16, g e - cata var. ambigua Trautv. = M. trautvetteri
nome formula DD, 32. The (sub)alpine zones of Sumnev (2n=16). It is a source of cold and drought
the Alkhalak uplands, Georgia, USSR and a part resistance. Ssp. coerulea (Less, ex Ledeb. )
of Asia Minor. A wild perennial species. Var. Schmalh. = M. coerulea Less, ex Ledeb. (2n=16).
timofeevii Troitz., (2n=32) is endemic to the It is a source of resistances to salinity of the soil
Transcaucasian Mountains. It crosses fairly easy and drought.
with M. sativa* (Lesins and Lesins, 1966). Some Ssp. glomerata (Balbis) Tutin = M. glomerata
include this species as var. dzhawakhetica Bordz. Balbis (2n=16) = M. glutinosa Bieb., (2n=16, 32)=
in M. papulosa Boiss. (2n=16). M. polychroa Grossh. (2n=32). It is a source of
disease resistance and persistency.
MEDICAGO HEMICYCLA Grossh. 2n=16, 32. The
alpine meadows of Transcaucasia and Daghestan. MEDICAGO SAXATILIS Bieb. 2n=28, genome for-
It is a source of cold resistance, and resistance mula S'S'X X X X . The S'genome of this species
to Verticillium wilt and Bacterial wilt. A wild is related to the S genome of M. sativa*. The X-
perennial species. It has been suggested that it is genome is possibly related to that of M. rhodopaea
a parent of M. sativa*. Velen. (2n=16), genome formula RpRp
MEDICAGO ROMANICA Prod. 2n=16. Caucasus. ONOBRYCHIS ALTISSIMA Grossh. 2n=14. Trans-
A wild variable perennial species caucasia.
MEDICAGO SATrVA L. Lucerne, Blue alfalfa. ONOBRYCHIS VICIIFOLIA Scop. Esparcette. 2n=28.
2n=16, genome formula SS, 32, genome formula Cultivation of this crop started in S. France
SSSS, (64). Transcaucasia. Wild types in Anatolia, (p. 137). In Transcaucasia var. transcaucasia
S. Caucasia and SW. Asia. Secondary centre NW. (syn. O. transcaucasia Grossh.) is endemic.
Iran. From Iran it reached Italy through Greece.
Italy through Greece. PISUM SATIVUM L. (syn. P . arvense L . s . 1 . ) .
Pea. 2n~14. This species may be divided into six
subspecies: ssp. abyssinium (p. 119), ssp. j o m a r -
di (p. 102), ssp. syriacum Berger, ssp. elatius
(Stev. )Alef., ssp. arvense Poir. and ssp. horten-
se Asch, et Graeb. (Gentry, 1971).
Ssp. syriacum (syn. P. humile Boiss. et Noë,
P. sativum var. humile, P . syriacum Berger) is
found in N. Iraq, Jordan, Syria, N., NW. and W.
Iran, Israel, Turkey and Cyprus. Some forms a r e
robust (30-70 cm tall), others slender and small
(20-40 cm) (Ben-Ze'ev and Zohary, 1973).
Ssp. elatius (syn. P. elatius Stev. ) is found in
Syria, N. Israel, Lebanon, S. coast of Turkey,
Aegean belt of Turkey and Greece, Cyprus, A d r i a -
tic coast of Yugoslavia, S. Italy, Sicily, Sardinia
and scattered localities in Morocco, Algeria, Tu-
nesia, S. Spain, S. France, N. Italy and the Black
Sea coast of Turkey, Crimea and Caucasia (Ben
Ze'ev and Zohary, 1973).
In the E. Mediterranean countries (esp. S.
Medicago sativa (Fischer, 1938) Turkey) intermediate types a r e found.
Ben Ze'ev and Zohary (1973) suggested that
ecotypes of Turkey and Syria may have formed
Another route of distribution is from Arabia through the parental material of the domesticated types
N. Africa to Spain and then through S. France ssp. arvense (P. arvense L. ), field pea and ssp.
(Provence) to W. and C. Europe. A third route was hortense (ssp. sativum, P . sativum L.), garden
from Media over Asia Minor through the Balkans to pea.
C. Europe. The French population 'met' the Bal- It may also derive from ssp. elatius, or from
kan population in Thuringia, Germany (p. 137) hybrids of ssp. elatius x ssp. arvense. Secondary
Lucerne also spread eastwards and reached China centre in the Mediterranean area (p. 102). Its do-
in the 2nd Century BC. It has also been used in mestication may have taken place in SW. Asia.
China as a vegetable. Two main introductions were The crop reached the Greeks via the Black Sea,
made to America. One from the Spanish population who passed it on to Latin and Germanic t r i b e s .
and another from C. Europe (p. 137). It spread to India and Chinathrough the Himalayas
'M. medica P e r s . ' originated and still does so and Tibet, and to Ethiopia and E. Africa (Purse-
where M. sativa and M. falcata grow together and glove, 1968).
hybridize.
LEGUMINOSAE - LINACEAE 87
Pisum humile ( ), P. elatius (...) and Alophotropis formosum ( ) (Govorov, 1937)
TRIFOLIUM AMBIGUUM M. B. 2n=16, (32, 48). tive cultivars, and convar. sativa, the cultivated
Caucasus and Crimea, USSR and Turkey. This forms.
valuable fodder plant forms an essential part of The following ancestry of common vetch has been
the pastures and meadows. suggested (Mettin &Hanelt, 1964):
Ancestor (2n=14?) > V. angustifolia ssp.
TRIGONELLA FOENUM-GRAECUM L. Fenugreek, angustifolia (2n=12) > V. ang. ssp. segetalis
Fenugrec. 2n=16. Probably SW. Asia. Cultivated (2n=12) ? > V sativa convar. cosentini
in S. Europe, N. Africa and India as a fodder. (2n=12) > V. sat. convar. sativa (2n=12).
The seeds are also eaten in India and used in medi- It is reasonable to assume that during this develop-
cine. ment introgression with other varieties took place.
V. cordata Wulf. (2n=10) is probably a derivative
VICIA ERVILEA (L. )Willd. Bitter vetch, Ervil. ofV. angustifolia ssp. angustifolia x V. sativa
2n=14. P r i m a r y centre in the Mediterranean r e - convar. consentini.
gion (p. 103). In Asia Minor a characteristic group Plants belonging to ssp. amphicarpa collected
developed. in C. Anatolia a r e very persistant as they can r e -
At present it is cultivated as a fodder, but in p r e - sist severe cold, drought and grazing.
historic times it was cultivated for food by man.
It was already cultivated in Turkey in 5 750 BC. VICIA VILLOSA Roth. Sand vetch, Hairy vetch,
and probably in Greece in about 5 500 BC. (van Winter vetch. 2n=14. W. and C. Europe, the
Zeist &Bottema, 1971). Mediterranean area, N. Iraq, N. Iran and SW. of
USSR. P r i m a r y centre probably in W. and Ante-
VICIA NARBONENSIS L. Narbonne vetch. 2n=14. Asia. Spread to the Mediterranean area and Europe
P r i m a r y gene centre probably in E. Georgia. as a cereal weed.
Secondary gene centre in the Mediterranean. This
species is a weed in wheat and barley fields in Liliaceae
Transcaucasia and other areas in SW. Asia. It HYACINTHUS ORIENTALIS L. Common hyacinth.
is not cultivated there. 2n=16, (24, 32). Syria, Asia Minor, Greece and
Dalmatia. Cultivated in the Netherlands as an o r -
VICIA PANNONICA Crantz. Hungarian vetch. namental and in S. France as a source of an
2n=12. P r i m a r y gene centre in Georgia, USSR, on essential oil used in perfumery.
the plateau of Akhalkalak, where it grows wild and
is cultivated. Secondary gene centre in Hungary.
Linaceae
VICIA SATIVA L. Common vetch. 2n=(10), 12, LINUM USITATISSIMUM L. Flax, Linseed. 2n=30,
(14). P r i m a r y centre of diversity in Centre 6. (32). P r i m a r y centre probably in centre 5 (Vavi-
Widely cultivated in the world as a green manure, lov, 1957).
fodder crop and for hay production. A polymorphic This conclusion is based on the great variation of
species. It may have derived from the weed V. flax in India and adjacent northerly a r e a s . How-
angustifolia L. (2n=12). ever, as the progenitor of flax L. bienne Mill.,
This latter species can be divided into the Pale flax, (2n=30) is not found in this a r e a it can-
wild ssp. angustifolia and the segetal type s s p . not have been domesticated there (Helbaek, 1956).
segetalis (Mettin &Hanelt, 1964). The latter is a Helbaek suggested that flax was more or less do-
weed of cereal fields. It easily c r o s s e s with V. mesticated at the same time as emmer and barley
sativa. This species might be divided into convar. in the mountainous region of the Near East. From
consentini, a segetal type which may include p r i m i - here it spread to other parts in the Old World.
NEAR EASTERN CENTRE
sistant and early maturing.
GOSSYPIUM HERBACEUM L. Short staple cotton.

2n=26, genome formula A..A... S. Africa (p. 121).
Introduced to Ethiopia, S. Arabia and Belutchistan
where race acerifolium* developed. In Iran a cha-
racteristic group of annual forms has arisen,
named race persicum. It spread to W. India where
it was the first annual cotton cultivated. At present
varieties of G. herbaceum a r e often cultivated. In
C. Asia race kuljianum developed. It will mature
in three months from sowing, resulting in a small
crop.
GOSSYPIUM INCANUM (Schwartz) Hillcoat. 2n=26,

genome formula E4E4. Aden. It is drought r e s i s -
tant.
GOSSYPIUM STOCKSII*
Moraceae
FICUS CARICA L. Common fig. 2n=26. Probably
S. Asia. P r i m a r y gene centre in SE. Asia. Spread
to Asia Minor, Mediterranean countries and W.
Europe (Storey &Condit, 1969). Cultivated for a
long time. In 4 000 BC. figs were already culti-
vated in Egypt. In Transcaucasia, Crimea, C.
Asia, Baluchistan and the Mediterranean countries
it ran wild a long time ago.
Nelumbonaceae
'I 2'\ 3\ t, NELUMBO NUCIFERA Gaertn. Indian lotus. 2n=16.
Centre of diversity probably lies in N. Iran, the
Linum usitatissimum, various length and branching types Kura estuary in Transcaucasia and Volga delta.
Cultivated in China, Japan and elsewhere for its
rhizomes and fruits. Formerly it was also grown
L. bienne can be divided into two main geo- in the E. Mediterranean region (Helmqvist, 1972).
graphical r a c e s . The first is the continental win-
t e r annual of the semi-arid foothills of Iraqi Kur- Papaveraceae
distan and Iran. This might be the parent of the
prostrate, multi-stemmed type cultivated since PAPAVER SOMNIFERUM*
ancient times along the N. coast of Turkey, the
Caspian coast of Azerbaijan and some parts of Polygonaceae
Kolchid bordering the Black Sea. According to FAGOPYRUM ESCULENTUM Moench. (syn. F .
Helbaek (1956) this type is the ancestor of the vulgare T. Nees, F . sagittatum Gilib., Polygo-
small-seeded flax cultivated by the prehistoric num fagopyrum L. ). Buckwheat, Silverhull,
C. European pile dwellers. The latter is the p a - 2n=16, 32. C. Asia. Introduced into several coun-
rent of 'Winterlein', a winter-annual cultivated in t r i e s as a grain crop. It is often found as a r u d e -
mountainous S. Germany. The second is the At- ral.
lantic-Mediterranean coastland race, a perennial,
also described as L. angustifolium Huds. (2n=30). Punicaceae
This race has the highest seed oil content and the
highest seed weight of all wild species (Seetharam, PUNICA GRANATUM L. Pomegranate. 2n=16, 18,
1972). 19. Wild in the Near East and C. Asia. An old
fruit t r e e which was even cultivated in the Hanging
During its domestication and further develop- Gardens of Babylon. Cultivated now in many coun-
ment, types for fibre (flax) and oil (linseed) were t r i e s . The only related species is P . protopunica
developed. Ralf, found wild on Socotra in the Indian Ocean.
Malvaceae
Resedaceae
ALTHAEA OFFICINALIS*
RESEDA PHYTEUMA*
ALTHAEA ROSAE*
Rosaceae
GOSSYPIUM AREYSIANUM Deflers. 2n=26, g e - AMYGDALUS BESSERIANA*
nome formula E3E3. S. Arabia. It is drought r e -
LINACEAE - ROSACEAE
AMYGDALUS COMMUNIS L. (syn. Prunus amyg-

dalus Batsch. ). Almond. 2n-16. P r i m a r y gene
centres In C. Asia (p. 74) and in Centre 6.
AMYGDALUS FENZLIANA (Fritsch) Lipsky (syn.

A. divaricata Fenzl., A. urartu S. T a m . , Prunus
fenzliana Fritsch. ). Fenzel almond. 2n=16. S.
Transcaucasia and Anatolia, Turkey. An ornamen-
tal. It easily c r o s s e s with A. communis* and it
might be a source of cold and drought resistance
for this species.
AMYGDALUS PERSICA L. Peach. 2n=16. P r i m a r y

centre in China (p. 36). Secondary centre in Cau-
casus and Crimea.
ARMENIACA VULGARIS L. Apricot. 2n=16. P r i -

mary centres in NE. China (p. 36) and in Daghes-
tan on the slopes of the Khunzakh plateau at an a l -
titude of 1 200-1 800 m. The latter centre proba-
bly linked formerly with the main one (p. 36).
The t r e e has a shrubby growth habit. Cultivated
over the entire area of the centre.
CYDONIA OBLONGA Mill. Quince. 2n=34. Talysh,

S. Daghestan, the lor valleys and in Azalan,
Georgia, in the T e r t e r valley, Azerbaijan and in Malus prunifolia
the canyons of Aidero and Yuz-Begi, Kojet-Dagh,
USSR. P r i m a r y centre lies h e r e . Cultivated for a
long time. PRUNUS AVIUM L. (syn. Cerasus avium Moench. ).
Sweet Cherry, Mazzard. 2n=16, (24, 32). P r i m a -
MALUS ORIENTALIS Uglits. 2n= . This is the ry centre in Asia Minor and Transcaucasia. Wild
only wild Malus species in the especially sparse t r e e s also in other parts of Europe, W. Asia and
oak forest of the Caucasus. It is polymorphous. N. Africa. The wild t r e e s of Ukrain could be
Through introgression, characteristics such as grouped into four classes viz. 1. dark-coloured
tallness, late ripening, good transportability of fruit: a. bitter and b. sweet and 2. light-coloured
fruits, high sugar content and unfortunately low fruit: c. bitter and d. sweet. The sweet-fruited
winterhardiness entered the cultivated apple types had elongated stones and longer fruit stalks
(Malus pumila Mill. ), which can still be recognized and petioles than the bitter-fruited types (M'ya-
in Caucasian, Crimean and even Italian cultivars. kushko and M'yakushko, 1970). It is likely that man
selected the sweet-fruited types.
MALUS PRUNIFOLU (Willd. ) Borkh. (syn.-Pyrus Rjadnova (1967) suggested that the domesti-
prunifolia Willd. ) Chinese apple. 2n=34, 51, 68. cation occurred in various places. This resulted
P r i m a r y centre in N. China. Cultivated in E. Asia in several ecotypes differing in resistance to un-
for its fruits. In the USSR this species is r e p r e - favourable conditions, in quality of fruit etc. Con-
sented in wild forms in E. Siberia. It is very r e - stant selection resulted in large-fruited, winter-
sistant.to frost and drought, much used by I. V. hardy types.
Michurin to breed hybrid varieties such as Kandil- P . avium is one of the parents of P . cerasus*.
kitaika, Bellefleur-kitaika, Saffran Peppin, Hybrids with P. cerasus (P. x gondounii (Poiteau
Saffran-kitaika. &Turpin) Rehder a r e known in W. Europe as
'Duke' c h e r r i e s . These hybrids and P . cerasus
MALUS PUMILA* are sources of resistance to bacterial canker
caused by Pseudomonas syringae Van Hall.
MALUS TURKMENORUM Juz. &M. Pop. 2n=
Turkemenia, in the gorges of the Kopet-Dagh PRUNUS CERASIFERA Ehrh. (syn. P . divaricata
Mountains. P r i m a r y gene centre also there. The Led.). Cherry plum, Myrobalan. 2n=16, genome
cultivated form is locally known as 'Baba-arabka', formula CC, (24, 32, 48). Wild in the Caucasus,
meaning: old arab woman. This name refers to Transcaucasia, Iran, Altai, Asia Minor and C.
the dying-down of the main stem at an age of about Asia. P r i m a r y centre C. and southern part of the
20 years and its replacement by soboles p e r m a - Caucasian coast of the Black Sea. From there it
nently rejuvenating the t r e e . spread eastwards and westwards. Secondary cen-
t r e in W. Tian-Shan (p. 75). It is a very poly-
MESPILUS GERMANICA L. Medlar. 2n=34. Cau- morphic species.
casus, N. Iran and Asia Minor. Cultivated e l s e - This species is one of the parents of P . do-
where and run wild there. It crosses with Cratae- mestica*. It is also planted as a rootstock and in
gus oxyacantha* and Sorbus aucuparia*. hedges. Var. pissardii (Carrière) L.H. Bailey
has dark red leaves and flowers tinged with reddish
pink. It is an ornamental. Rubiaceae

PRUNUS CERASUS L. (syn. Cerasus vulgaris COFFEA ARABICA L. Arabica coffee. 2n=22, 44,
Mill. ). Sour cherry. Pie cherry. 2n=32. Unknown (66). The primary centre in SW. Ethiopia. Secon-
wild although t r e e s that have run wild grow main- dary centre in Yemen. This area is the source of
ly in the Caucasus and Asia Minor, but also in the Arabica coffee now cultivated in Latin America,
European USSR. W. Balkan countries and Germa- Kenya, India, Java and elsewhere (Meyer, 1965).
ny. Probably an allotetraploid of P . fruticosa* x
P. avium*. Sour cherry can be divided into the Rutaceae
true Sour cherries and Duke-cherries. The first CITRUS MEDICA L. Citron. 2n=18. Probably
can be subdivided into Morellos (austera L. ) and SW. Asia, although India has often been mentioned
Amarelles (caproniana L. ) (Zylka, 1971a). as the centre of origin. Unknown wild. It has now
A special population 'Vladimirskaya vishnia' spread through the (sub)tropics.
originated in centre 9 (p. 140). The Etrog citron (var. ethrog Engl. ) is used
by Jews at the Feast of Tabernacles, and the finge-
PRUNUS DOMESTICA L. Garden plum, Domestic red citron (var. sarcodactylis Noot. ) Swing) by
plum. 2n=48, genome formula CCSSSS or the Chinese as a medicine and an ornamental.
CdCdSSSiSi or CdCdD 1 D 1 D 2 D 2 . Caucasus. This
species is thought to be a natural hexaploid of Umbelliferae
P. cerasifera* and P. spinosa*. This alloploidi-
zation apparently took place in the Caucasus, where CUMINUM CYMINUM*
both species occur and natural hybrids with 2n=24
and 48 a r e still found. However it may have MALABAILA SECACUL (Mill.) Boiss. Sekakul.
happened elsewhere. Too e.g. Werneck (1958) 2n= . Asia Minor and Syria. Cultivated for its
considered the garden plum to have arisen in roots used as a aphrodisiac.
Upper Austria (p. 140).
It is interesting to note that Rybin (1936) PIMPINELLA ANISUM L. (syn. Anisum vulgare
resynthesized the garden plum. Artificially ob- Gaertn., Anisum officinarum Moench). Anise
tained hexaploids resembled it. plant. 2n=18, 20. Probably the Orient. Cultivated
for aromatic fruits.
PRUNUS SPINOSA L. Blackthorn, Sloe. 2n=32,
genome formula SSSS or SSS.S or SSC C . Wild Vitadaceae
throughout the entire territory of this centre and VITE LABRUSCA L. Fox grape. 2n=38. N. A m e -
in Europe and N. Africa. Volga basin types carry rica (p. 179). Introduced into W. Georgia, USSR as
genes for high winterhardiness. It is one of the a cultivated grape.
parents of P . domestica*. Some natural hybrids
with this latter species a r e described as P. fruti- VITIS VINIFERA L. Common grape, European
cans Weihe (2n=40). grape. 2n=38, (40, 57, 76). P r i m a r y centres:
the Central Asian (p. 76), the Near East and the
PYRUS. This centre is the main geographic centre Mediterranean centres (p. 106). The wild vine ssp.
of formation of Pyrus species. Of the about 60 sylvestris Gmel. is found in regions bordering
Pyrus species in the world about 25 have been the Mediterranean Sea with Libya and Egypt excep-
described for the Caucasus. Some of them also ting, up to Turkestan and Kashmir. P r i m a r y
occur in Iran or in Asia Minor. centre: probably Armenia, USSR and N. Iran. The
western wild types have been called ssp. silvestris,
PYRUS CAUCASICA Fed. 2n= . The entire while the eastern types a r e ssp. caucasia Vav.
forest zone of the Caucasus except the Talysb The wild and cultivated (ssp. sativa DC., ssp.
Mts. A polymorphic species. In open areas it vinifera) types a r e very variable resulting in many
spreads quickly and vigorously. forms. The domestication took place by selecting
natural bud mutants and hybrids.
PYRUS SYRIACA Boiss. 2n= . Armenia. It is The actual domestication may have taken place
cold-resistant. It probably played a part in the in SE. Europe where types with large bunches and
origin of the cultivated pear (Evreinoff, 1944). seedless grapes have developed. Natural hybridi-
Cultivated locally. zation is still taking place in several areas e. g.
the mountains of S. Tajikistan, USSR where many
PYRUS TAKHTADZHIANA Fed. 2n= . It has the new forms a r e observed.
habit of a cultivated t r e e . Cultivated in ancient The common grape has been crossed with the
times, but later it ran wild. North American V. labrusca*. The fruits a r e
used to prepare wine, currants and raisins.
ROSA CENTIFOLIA L. (syn. R. gallica L. var. V. amurensis* is a possible source of winter-
centifolia Reg. ). Provence rose. 2n=28. E. Cau- hardiness.
casia. Cultivated for its flowers. The petals are
used in the perfume industry.
SORBUS DOMESTICA L. Service t r e e , Mountain

ash. 2n=34. Its distribution is given on p. 141.
Large-fruited forms a r e found in forests of Crimea.
7 Mediterranean W
S
^~r-
^\ ) ^< r '
)-.^'>-~^
"-'< ij
Centre /f~~^~~~TÂ "\_ X ^ f J ^»*'
""N. [ r . <—<^^>
v >" ° ' )
\ /=\7T.,^/ °c= cM\
/ • - ^ '
"^xJ • ""^
\J--J
i \
1
The Mediterranean Centre was called by Vavllov the Mediterranean Centre of Origin, while Darlington
(1956) suggested Mediterranean Region of Origin.
Its situation near a cradle of agriculture Centre 6 led to an early introduction of plant cultivation.
Early farming sites were found at Nea Nikomedeia, Greece dating c. 5 470 BC. (van Zeist & Bottema,
1971) and at Fayum, Egypt dating from the 5th millenium, reaching the Atlantic Ocean maybe in c. 3rd
millenium.
A very old site at Kom Ombo, in the Nile Valley, Upper Egypt dated from 15 000-10 500 BC. It is a
non-farming site occupied the whole year round (Churcher &Smith, 1972).
Many crops have been domesticated in this region: Avena s p . , Beta vulgaris, Brassica napus, B.
oleracea, Lathyrus s p . , Linum usitatissimum, L o l i u m s p . , Lupinus s p . , Olea europaea, Raphanus
sativus, Trifolium s p . , Vitis vinifera, etc.
Alliaceae NARCISSUS POETICUS L. Poet's narcissus.

ALLIUM CEFA L. Spanish Onion. 2n=16. Secon- 2n=14, 21. Portugal, Spain, France and Italy.
dary centre in the Mediterranean Centre. Cultivated as an ornamental and in S. France for
its essential oil.
ALLIUM SATIVUM L. Garlic, 2n=16, genome
formula SS. C. Asia (p. 71). Secondary centre in Anacardiaceae
Centre 7 (Kazakova, 1971). RHUS CORIARIA L. Sicilian sumach, 2n=
Mediterranean region. A shrub cultivated in Sici-
Amaranthaceae lia and S. Italy for the leaves which a r e a source
AMARANTHUS LIVIDUS L. 2n=34. Spread through of tanning material
Europe, Asia and to the tropics of the Old and New
World. Var. ascendens Thell. (syn. A. viridis L., Apocynaceae
2n=34) is native to S. Europe and E. Mediterranean NERIUM OLEANDER L. Oleander. 2n=16, 22.
region. Cultivated there in the Middle Ages. Var. A shrub of Mediterranean region. Cultivated as an
lividus is unknown wild. It might be a cultigen of ornamental and for other purposes.
this species. It was cultivated in the 16th and 17th
centuries as a vegetable and medicinal crop and in Asclepiadiaceae
the 18th Century as pig food.
Var. oleraceus Thell. (syn. A. oleraceus L., CYNANCHUM VINCETOXICUM*
2n= ) is probably a cultigen type of var. ascen-
dens. Cultivated in Europe and elsewhere as a Balanitaceae
vegetable (Mansfeld, 1959). BALANITES AEGYPTIACA Del. Betu, Desert
date. 2n=16, 18. This shrub grows wild in N. trop.
Amaryllidaceae Africa, Arabia, Palestine and also in Angola.
Cultivated in Egypt for its edible leaves and
NARCISSUS JONQUILLA L. Jonquille. 2n=14.
flowers (Cufodontis, 1957; T e r r a , 1967).
Europe, Ante-Asia up to Iran and Algeria.
Commonly cultivated as an ornamental and in S.
France for its essential oil.
MEDITERRANEAN CENTRE 92
Boraginaceae ted formerly for its roots which contain saponin.

Now it is an ornamental.
ALKANNA TINCTORIA (L. ) Tausch. Alkanna.
2n=14. S. and E. Europe and Turkey. A herb cul- Chenopodlaceae
tivated as a source of a red pigment.
BETA PATELLARIS Moq. 2n=18, (36). Mediterra-
BORAGO OFFICINALIS L. Borage. 2n=16. Medi- nean and Atlantic coasts of NW. Africa, Canary
terranean region. A herb cultivated for ornament, Islands, the Cape Verde Islands and Madeira. A
as a potherb and for bees. source of nematode and Cercospora resistance
and yellow mosaic tolerance for B. vulgaris*.
Capparidaceae
BETA PROCUMBENS Chr. 2n=18. Canary Islands
CAPPARIS SPINOSA L. Caper bush. 2n=24, 38. and the Cape Verde Islands. A source of nematode
The cultivated forms: (large) flower head, var. resistance for B. vulgaris*.
spinosa and small flower head, var. parviflora
J. Gray probably derived from the wild var.
BETA VULGARIS L. Beet. 2n=18. The parental
aegyptla (Lam. ) Boiss. This variety grows wild
form is the wild sea beet (ssp. maritima (L.)
in S. and SE. Mediterranean region up to the
Thell., syn. B. maritima L. ). P r i m a r y centre
Sudanian and Eritro-Arabian regions. Var. spino-
probably in the E. part of the Mediterranean
sa was developed in the N. Mediterranean region.
region. Spread in a western direction along the
From here this form spread to other areas where
coast of the Mediterranean, W. coast of Europe
it is cultivated for its use as a condiment. Var.
and to Cape Verde Islands and Canary Islands. In
parviflora is also cultivated and might be a mutant
the Mediterranean region, leaves and roots of
type of var. spinosa. Hybrids with C. ovata Desf.
this wild plant may have been collected. This may
a r e found.
have led to the development of the Swiss chard
(var. vulgaris, var. cicla) of which the leaves
Caryophyllaceae and stalks a r e eaten and to the garden beet, table
DIANTHUS CARYOPHYLLUS L. Carnation, Clove, beet and red beet (var. cruenta, var. esculenta).
Pink, Picotes. 2n=30. Mediterranean region. A This development may have been influenced by
perennial herb cultivated as an ornamental and also hybridization with wild types like spp. macrocarpa
as a source of an essential oil. (syn. B. macrocarpa Guss. ) in N. Africa.
The fodder beet (var. rapa) probably developed
GYPSOPHILA PANICULATA L. Baby's breath. in the Netherlands (p. 130) after introduction of
2n=34. S. and C. Europe and Caucasus. Cultiva-
#>"*
/art ^^S^J~Y f t?S
/ i< WlJ [
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/ ^—__^l) ^ / ^ V Vs VS. V>

'1 ? 0 ^
/ 'Œ^'JQ^C*w ""^"•k. ) r '
/ / ks/- °3*w{ 2..!^sfiP^=»--<v ^ ^r >-^r-^~?r'--'-î

a
fc"''" -(
$
v «V^
*V-^'~'^%!N. r
'"
"^-J\^-~'^
/jp4 /
// ~ [ ƒ/
y \
'
,Y
s
\T—N f^^V^ ^ i i «-'X
fi'-'' J~. A
\ ^ Sv
o' J
'^r '•'
^ J ^ x ^\ *• -^\. ' "T

J
K' Jiy- ' "'' '• ' ^"' ' A
^»»»J^—~
' ~\ ^ ^ \
'&^K y i ^ c\ ; i ^ r ^ \--3 /
—"•' i V
! i i .' i A \ -'"*''r \
* * / i i ' i
;M^ _-. — U il
Beta vulgaris (1), B. patellaris (2), B. procumbens and B. webbiana (3), B. patula (4) and B. atriplicifolia (5) (Ulbrich, 1934)
types from Spain, and the sugar-beet in Silecia, Transcaucasia. Syria and Iran. Cultivated f o r m e r -
Poland (p. 140). The wild B. macroearpa Guss. ly in Germany.
from the coasts of the Mediterranean region and
Canary Islands, B. patula Ait. from Madeira and CYNARA CARDUNCULUS L. Cardoon. 2n-34 The
B. atriplicifolia Rouy from S. Spain easily hy- Mediterranean region. Cultivated for its leaf
bridize with B. vulgaris. They may be included stalks. Several varieties a r e known.
as subspecies in this species.
In NW. Europe, hybrid plants of cultivated s u g a r - CYNARA SCOLYMUS L. Artichoke, Globe a r t i -
beet and spp. maritima a r e occasionally observed. choke. 2n^34. The Mediterranean region. Culti-
They derive from material propagated in France vated for its soft fleshy edible receptacles of the
and Italy. Such hybrids bolt in the first year p r o - flowerhead buds and thick bases of the scales
ducing seeds. These seeds drop and may result in around the flowerhead and also as a source of a
a weed (F2 plants) for several years to come. It bitter compound. Several varieties a r e known.
is possible that on a very small scale, wild genes
derived from these hybrids introgress in the cul- LACTUA VIROSA L. Bitter lettuce, Lettuce
tivated population. opium. 2n=18. P r i m a r y centre round the Medi-
The wild plants may form sources of resistance terranean (Linqvist. 1960). Cultivated on a small
to disease such as Cercospora, yellow mosaic and scale for its latex in some parts of Europe. The
to increase the variability to select for new high latex has narcotic properties.
yielding types.
SCOLYMUS HISPANICUS L. Golden thistle, Spa-
BETA WEBBIANA Moq. 2n=18. Canary Islands. nish oyster plant. 2n=20. The Mediterranean r e -
A source of nematode resistance for B. vulgaris*. gion. A root vegetable. Its cultivation is on the
decline.
CHENOPODIUM AMBROSIOIDES L. American
wormweed, Indian wormweed. 2n=16, 32, 36, 64. SCORZONERA HISPANICA L. Scorzonera, Black
Probably S. Europe. Widespread in the tropics salsify. 2n=14. C. Europe, the Mediterranean
and introduced in N. America. The cultivated region, Caucasia and S. Siberia. A vegetable of
type, var. anthelminticus L. is a source of medi- especially S. Europe. Perhaps it was first culti-
cinal and essential oils. vated in Spain (Mansfeld, 1959).
HALOGETON SATIVUS (L. ) Moq. 2n= . NW. SILYBUM MARIANUM (L. ) Gaertn. Holy thistle,
Africa. Cultivated in the Mediterranean region as Milk thistle, Lady's milk. 2n=34. S. Europe.
it yields a base-rich ash when burned. Cultivated as a medicinal plant and as an ornamen-
tal.
Compositae
ANACYCLUS OFFICINARUM Hayne. Bertram. TRAGOPOGON PORRIFOLIUS L. Salsify, Oyster
2n=18. Probably the Mediterranean region. Culti- plant. Purple goats beard. 2n=12. The Mediterra-
vated formerly in C. Europe. nean region. This vegetable was first cultivated
for its roots long ago. It may have been domesti-
cated by the Greeks and Romans (Mansfeld, 1959).
ANACYCLUS PYRETHRUM (L. ) Link. Pellitoria
The wild type is var. australis (Jord. ) Braun-
of Spain. 2n=18. N. Africa, Arabia and Syria.
Blanquet (syn. T. australis Gater. ) and the culti-
Cultivated formerly in Europe as a medicinal
vated one is var. sativus (Gater. ) Braun-Blanquet
plant and now in Algeria for an essential oil.
(syn. T. sativus Gater. ).
ARTEMISIA JUDAICA L. 2n= Cultivated in
the Mediterranean region. Convolvulaceae
CONVOLVULUS SCAMMONIA L. 2n=24. The
CALENDULA OFFICINALIS L. Marigold. 2n=(28), Mediterranean region and Asia Minor. Cultivated
32. Centre of origin probably in the Mediterranean for medicinal purposes.
region. Cultivated as an ornamental, but f o r m e r -
ly as a medicinal plant. Corylaceae
CHRYSANTHEMUM CINERARIAEFOLIUM (Trev.) CORYLUS TUBULOSA Willd. Lambert's filbert,

Brocc. Pyrethrum. 2n=18. Dalmatian coast of Kentish cob. 2n= . Cultivated. Hybrids with
Yugoslavia. Introduced to other countries. Kenya C. avellana* have been cultivated.
is the main producer of the insecticidal Pyrethrine.
Cruciferae
CHRYSANTHEMUM PARTHENIUM (L.) Bernh. BRASSICA CAMPESTRK*
Feverfew, Wild chamomile. 2n=18. The Mediterra-
nean region, Balkans, Asia Minor and Caucasia. BRASSICA CRETICA Lam. 2n=18. Wild in Greece,
Cultivated as medicinal plant in Europe. Crete, Cyprus, Syria and Lebanon. May have
played a role in the origin of B oleracea* var.
CNICUS BENEDICTUS L. (syn. Centaurea bene- botrytis. This species has also been described as
d i c t a L . , Garberia benedicta Adans. ). Blessed var. sylvestris, the wild cabbage.
thistle. 2n=22. The Mediterranean region up to
BRASSICA NAPOBRASSICA (L. ) Mill. Rutabaga, gression with wild species and other cultivated
Swedish turnip. 2n-38. Secondary gene centre in types and by mutation under human selection
Europe. The cultigen developed in the Mediterra- pressure. <-,/>,: (.:.-( -
nean region and further in Europe (p. 131). A simplified pedigree is presented by Helm (1963a).
He suggested that from (1) the wild cabbage (var.
BRASSICA NAPUS L. Rape, Colza. 2n=38, g e - sylvestris, syn. var. oleracea) developed the
nome formula AACC. This species is an amphi- types (2) var. ramosa DC., cottager's kale, (5)
ploid of B. oleracea spp oleracea (2n=18, ge- convar. acephala p C . ) A l e f . , (6) var. medullosa
nome formula CC, see p. 94) and B. campestris Thell. and convar. botrytis (L. )Alef. The cotta-
(2n=20, genome formula AA, see p. 131). The g e r ' s kale was used as forage. It has almost d i s -
first amphiploid was the "primitive leaf rape" appeared now. From this crop (3) var. gemmi-
(spp. pabularia (DC.) Jancken). This primitive fera DC., Brussels sprouts was derived. This
type may have developed in the W. Mediterranean type may have developed in Belgium (p. 131). From
region. From this type cultivated types as spp. the Brussels sprouts, monstrosities such as (4)
oleifera DC., rape and spp. rapifera Metzger var. dalechampii Helm derived. However, Nieuw-
derive, while B. napobrassica* formerly described hof (1969) suggested that the monstrosity pictured
as B. napus var. napobrassica (L. )Reichenb. is by Daleehamp (Helm, 1963a) is a cabbage with the
probably a derivative of B. oleracea* x B. napus, axil buds developed after the head had been r e -
The wild var. napus may be the weedy derivative. moved.
B. napus is one of the parents of the artificially Convar. acephala can be subdivided in many types
made B. napocampestris (2n=58, genome formula as var. acephala, kale, borecole, collard, cow
AAA 1 A 1 C 1 C 1 ). cabbage. This cabbage type belongs to the oldest
cultivated types. The main use is forage crop. On
BRASSICA OLERACEA L. Cabbage. 2n=18, g e - J e r s e y and Guernsey f. exaltata, Cesarean cabba-
nome formula CC. The C genome is related to ge, J e r s e y cabbage was cultivated. When closely
the T or D. genome of B. tournefortii Gouan planted the branches and stalks can be used as
(2n=20). It forms one pair of the genomes of walking sticks (Jersey canes) and in house buil-
B. carinata* and B. napus*. The wild form, var. ding. Other varieties belong to this group. These
sylvestris L. (syn. B. sylvestris (L. ) Miller), a r e var. selenisia L., var. s a b e l l i c a L . , var.
grows on the maritime cliffs of the west coasts of palmifolia DC., var. medullosa Thell. and var.
Britain, France and the Mediterranean coastal gongylodes L.
areas. Var. selenisia, parsley colewort, ornamental
The wild type is very variable. When cultivated kale has been used as an ornamental.
it shows an enormous increase in size. Up to now Var. sabellica is curled kitchen kale, ornamental
this variability has not yet been fully studied. It Scotch kale, curlies.
is possible that through introgression some geo- F. sabellica is mostly cultivated as a vegetable,
graphical r a c e s have a different genetic potency while f. rubra is used as an ornamental.
than other r a c e s . Some related species are: B. Var. palmifolia, palm-leaved kale probably origi-
incana Tenore (2n= )from W. and S. coasts nated in Portugal. It is likely used only as an o r -
of Baly, Sicily and Yugoslavia, B. montana namental.
Pourret (including B. oleracea spp robertiana Var. medullosa, marrow stem kale is a forage
(Gay) Rouy &Fouc. ) (2n= ) from S. Europe, crop. Maybe Pliny's Pompeian cabbage is an a n -
NE. Spain to S. Italy, B. rupestris Raf. (2n=18) cestor of this type. From this same type or from
from Sicily and S. Italy and B. villosa Biv. -Bern. marrow stem kale the (7) var. gongylodes , kohl-
(2n= ) from Sicily. rabi, turnip kale derives.
Other related species a r e B. cretica*, B. balea- From the old cultivars of convar. acephala convar.
rica Persoon (2n=18) from Mallorca, B. insularis capitata (L. )Alef. developed. The oldest type is
Moris. (2n=18) from Corsica and Sardinia, B. probably (8) Tronchuda kale, Portugese kale, var.
macrocarpa Guss. (2n= )from Isole Egadi, costata DC. which developed in Portugal. From
Sicily and B. scopularum Coss. &Dur. (2n= ). this variety (9) var. sabauda L. and (10) var.
Helm (1963a) preferred to include all related wild capitata L. developed.
species into one species group B. oleracea. Var. sabauda, Savoy cabbage, Milan cabbage is
The wild and cultivated types of B. oleracea s. s. probably developed in Italy.
cross with wild related species. By absorbing Var. capitata, cabbage and red cabbage has deve-
genetic material from these species its variability loped from the same stock as var. sabauda. Their
and adaptability has greatly increased. B. oleracea history is not known.
has become a compilospecies (A.C. Zeven, u n - Convar. botrytis exists of (11) var. italica Plenck,
publ. ). This may also have resulted in the v a r i a - sprouting broccoli, asparagus broccoli and (12)
tion of karyotypes. Furthermore, the present var. botrytis L., cauliflower. The latter may have
variation may also have been influenced by neigh- derived from var. italica. Both types developed in
bouring cultivars. the eastern part of the Mediterranean countries.
There a r e several classifications of the cultivated It is possible that B. cretica* has played a role
B. oleracea-types. See e.g. Helm (1963a). The (Jensma, 1957). This author suggested that from
origin of these types is not yet fully understood Cyprus and other E. Mediterranean countries the
and it is supposed that they developed gradually cauliflower was brought to Venice and to Vienna
from several wild cabbage populations by intro- from where it spread to N. countries.
CRUCIFERAE - CYPERACEAE 95
CAPSELLA BURSA-PASTORIS (L. ). Medik. R. caudatus L. ). Var. sativus, the radish, small
Shepherd's purse, Capsell. 2n=32. The Mediterra- radish. Var. niger Kerner, radish, Spanish r a -
nean region. Spread over almost the whole world. dish. Recently fodder radish has been bred. It is
Cultivated in China as a vegetable. a reputed selection of oil-seed radish in France.
More research is needed to ascertain the origin
CHEIRANTHUS CHEIRI L. Wallflower. 2n=14. of radish and the various botanical and agricultural
S. and C. Europe. Herb commonly cultivated as varieties. Through natural (and artificial) hybridi-
an ornamental, formerly as a medicinal crop. zation with Brassica-species genes of this genus
may introgress into R. sativus.
CRAMBE HISPANICA L. 2n=60. The Mediterranean
region. Cultivated in the Ukraine, USSR for its oil. SINAPIS ALBA L (syn. Brassica alba (L. ) Boiss. ).
White mustard. 2n=24. The Mediterranean region.
ERUCA VESICARIA (L. ) Cav. (syn. E. sativa L. ). Weedy or naturalized plants may be found from
Rocket salad, Roquette. 2n=22. The Mediterranean Spain over Asia Minor to E. India. Young seed-
region and Asia. Cultivated since ancient times as lings a r e used as salad. Seeds are the source of
a salad plant. Cultivated in India as a source of white mustard.
jamba oil (p. 63).
Cucurbitaceae
LEPIDIUM LATIFOLIUM L. Dittander. 2n=24. BRYONIA CRETICA*
Europe, temperate Asia and N. Africa. A perenni-
al herb. Once it was cultivated by the Ancient
CITRULLUS COLOCYNTHIS (L. ) Schrad. Colo-
Greeks as a salad plant.
cynth. 2n^22, (34). Arid regions of N. Africa and
trop. Asia. Cultivated in India and the Mediterra-
RAPHANUS SATIVUS L. Radish, Small radish.
nean region for its purgative fruits.
2n=18, genome formula RR. P r i m a r y centre p r o -
bably an E. Mediterranean region (Wein. 1964).
ECBALLIUM ELATERIUM (L.) A.Rich. Squirting
He suggested that the radish is derived from R.
cucumber. 2n=(18), 24. The Mediterranean region,
maritimus Smith (2n=18) and the small radish
Azores, Asia Minor and Crim. Cultivated in
from R. landra Moretti (2n=18). He stated that the
England as a medicinal plant.
radish cannot come from R. raphanistrum L.*
(2n=18) because of the difference in structure of
the fruits. Various varieties have been described Cyperaceae
(Mansfeld, 1959). Var. oleiformis P e r s . (R. chi- CYPERUS ALOPECUROIDES Rottb. Mat sedge.
nensis Mill. ) is the oil-seed radish cultivated in 2n= . Tropics of the Old World. Cultivated in
India, Japan, China (p. 30) and on a small scale Egypt for mat making (Mansfeld, 1959).
in Rumania and Spain. Var. mougri Helm* (syn.
/ % HXu fi /
y w
.**•'Â Y i \v ^ V'/'
Wild and weedy Raphanus species: R. raphanistrum ( ), R. maritimus (black) and R. landra (grey) (Sinskaia, 1931)
CYPERUS ESCULENTUS L. Chufa, Earth almond, 30-40, 36), 40, (48, 54). S. , W. and C. Europe,
Tiger nut, Rush nut, Zulu nut, Yellow nut g r a s s . Mediterranean region, temp. Asia. Cultivated as
2n=(18), 108. White Nile region and in the tropics. sheep food in N. and S. America.
Introduced in S. Europe by the Arabs. Cultivated
in Spain and Italy and elsewhere for its flavour- ERODIUM MOSCHATUM (L. ) L'Herit. ex Ait.
some tubers. Var. aureus (Ten, )Richt, is d e s - White stem filaree. 2n=20. The Mediterranean
cribed as the wild form. Var. esculentus is the region. Cultivated formerly as a medicinal crop.
cultivated form.
Gramineae
CYPERUS PAPYRUS L. Papyrus plant. 2n=c. 102. AEGILOPS BICORNE (Forsk. )Jaub. &Sp. (syn.
Africa. Cultivated formerly in Egypt, Palestine Tritieum bicorne Forsk. ). 2n=14, genome formu-
and in the Mediterranean region. Now occasionally la S°S . Xeric sandy soils of S. Israel. Lower
cultivated. Egypt and Cyrenaica. It is sometimes believed to
be the B donor of tetraploid and hexaploid Tritieum
Ericaceae species (p. 83, 84).
ARBUTUS UNEDO L. Strawberry tree, Arbutus.
2n=26. The Mediterranean region. Occasionally AEGILOPS COMOSA Sibth. &Sm. (syn. Tritieum
cultivated for its edible fruits. comosum (Sibth. &Sm. )Richter). 2n=14, genome
formula MM. Mediterranean Greece, the Aegean
Islands and W. Turkey. Used as a source of r e -
sistance to yellow rust (Puccinia striiformis West. )
AEGILOPS CYLINDRICA*
AEGILOPS KOTSCHYI*
AEGILOPS LORENTÜ*
AEGILOPS OVATA*
AEGILOPS TRIARISTATA*
AEGILOPS TRIUNCIALIS*
AEGILOPS UNIARISTATA Vis. (syn. Tritieum

unlaristatum (Vis. )Richter). 2n=14, genome for-
mula M U M U . The Mediterranean Greece, the Mar-
mara sea area and the Adriatic zone of Yugoslavia.
AEGILOPS VARIABILIS Eig. (syn. Ae. peregrina

Arbutus unedo (Hutchinson, 1969) (Hack. ) Maire & Weill., Tritieum peregrinum
Hack. &F r a s e r ) . 2n=28, genome formula
C U C U S V S V . N. Africa, Egypt, Palestine, Greek
Euphorbiaceae
Islands, Turkey and Iraq.
CHROZOPHORA TINCTORIA (L. )J u s s . Giradol.
The Mediterranean region, France, Yugoslavia, AEGILOPS VENTRICOSA Tausch, (syn. Tritieum
Crim to W. Asia, NW. India, Arabia. Cultivated ventricosum C e s . , P a s s . &Gib. ). 2n=28, genome
formerly in S. France as a source of red and blue formula MVMVDD. The W. Mediterranean region.
dye. The red dye was used for colouring Dutch It is a source of resistance to the wheat disease
cheeses. eyespot caused by Cercosporella herpotricoides
Fron. Natural hybrids with Tritieum turgidum
EUPHORBIA LATHYRUS L. 2n=20. S., W. and C. group durum have been found and described as
Europe. A ruderal and weedy plant occasionally Tritieum rodeti Trabut. Amphiploids with t e t r a -
cultivated as a medicinal. Probably only native to ploid Tritieum species have been named Aegilo-
E. and C. Mediterranean region. tricum.
Fagaceae AGROPYRON JUNCEUM (Jusl. ) Beauv. Sea wheat-

QUERCUS SUBER L. Cork oak. 2n=24. W. part of g r a s s , Bent g r a s s . 2n=28, 42, (84). The coasts
centre 7. A very variable species. Cultivated in of Europe, N. Africa and Asia Minor. Occasio-
S. France, Portugal, Spain, Sardinia, Corsica, nally cultivated to stabilize dunes.
Istria, Dalmatia and Algeria.
AGROSTIS TENUIS*
Geraniaceae
ARUNDO DONAX L. Giant reed. 2n=(c. 60), 110.
ERODIUM CICUTARIUM (L. ) L'Herit. ex Ait. The Mediterranean region to Caucasia and Syria.
Storko bill, Red stem filaree, Alfilaree. 2n=(20, A grass cultivated since ancient times in S.
CYPERACEAE - GRAMINEAE 97
Europe. Also cultivated elsewhere now. la ApAp. The coastal belts of Mediterranean coun-
t r i e s and Moroceo, Portugal and Spain. Medi-
AVENA CANARIENSIS Baum. Rajhathy &Samp- terranean and Negev desert ecogeographic races
son. 2n=14. The uplands of Fuerteventura, Canary a r e recognized (Ladizinsky & Zohary, 1971.
Islands. B a u m e t a l . (1973) discovered that its
genome formula is closely related to the A genome. AVENA PROSTRATA Ladizinsky. 2n=14, genome
They suggested that A. canariensis is the diploid formula ApAp. A species of oat from SE. Spain.
parent of A. magna* and hence is one of the p a - Morphologically similar to the diploid wiestii and
rental species of hexaploid A. sterilis (see A. hirtula forms of A. strigosa*, but can be distin-
sativa*). guished from them by its prostrate habit and by
shorter bristles at the tip of the lemma. Hybrids
AVENA CLAUDA Dur. 2n=14. The whole Medi- between these two species a r e completely sterile
terranean basin from Morocco to Iran. It usually (Ladizinsky, 1971).
grows together with A. sativa type sterilis* and
A. strigosa type barbata* (Ladizinsky & Zohary, AVENA SATIVA L. Oat. 2n=42, genome formula
1971). AACCDD. The origin of A. sativa is not yet fully
This wild species includes type eriantha (syn. understood. It is believed that it derives from the
A. eriantha Dur. = A. pilosa MB, genome formula sterilis type. This type is related to the wild
ApAp) and type elauda (A. clauda Dur. ). tetraploid species A. magna Murphy & Terrell,
found in the Rabat-Tiflet a r e a of Morocco and
AVENA DAMASCENA Rajhathy &Baum. 2n=14, A. murphyi Ladiz. found in the region between
genome formula AdAd. An a r e a 60 km north of Tarifa and Vejer de la Frontera at the southern
Damascus, Syria. It has a high degree of genome tip of Spain. These two species may form the
homology with A. prostrata*. Both species a r e genetic background of A. sativa.
considered relicts of a once common population, According to Ladizinsky and Zohary (1971), this
but a r e now separated by some 2500 km (Rajhathy species forms a complex of wild, weedy and culti-
& Baum, 1972). It resembles A. strigosa. vated types, including A. sterilis L., A. fatua L.,
A. byzantina C. Koch* and A. sativa L. s. str.
AVENA LONGIGLUMIS Dur. 2n=14, genome formu- The last two a r e the cultivated types. The wild
Avena clauda (Ladizinsky &Zohary, 1971)
Avena longiglumis (Ladizinsky &Zohary, 1971)

type sterilis is found in the Mediterranean coun- wiestii Schreb. (2n=14), A. barbata Pott. (2n=28),
t r i e s , where it builds massive stands from the A. vaviloviana Malz. * (2n=28) and the cultivated
Atlantic coast of Morocco and Portugal to the A. strigosa Schreb. (2n=14)andA. abyssinica
western flanks of the Zagros mountains of Iran. Höchst.* (2n=28).
They also a r e agressive pioneers and noxious All over the Mediterranean region, wild and weedy
weeds in wheat and barley fields, etc. Sterilis diploid and tetraploid forms a r e found, hybridizing
type with small spikelets has been described as freely. "A. hirtula*" is common in Spain, Morocco,
A. ludoviciana Dur., and that with bigger spike- Algeria, Italy, Greece, Turkey and Israel. "A.
lets and 3-4 fertile florets as A. macrocarpa wiestii" grows in the d r i e r steppes of the northern
Monch. The fatua type is distinctly weedy. It also fringes of the Sahara and the Arabian d e s e r t s . The
occurs in the Mediterranean countries. cultivated strigosa of W. and N. Europe derives
The main characters of the cultivated types a r e from the weedy forms which a r e common in cereal
non-shattering spikelets, decrease in lemma fields and edges of cultivation in the Iberian penin-
hairiness and reduction in the size and develop- sula. The As and B genome a r e partially homolo-
ment of awns. Nakedness (A. nuda L*) is also gous and many derive from a common parent
associated with domestication. Cultivated oat is (Ladizinsky & Zohary, 1971). The As genome
found in many countries now. might be the prototype of the A genome of the
Rajhathy et al. (1971) concluded from chromato- polyploid species (Rajhathy et a l . , 1971).
graphic studies that A. magna has the genome Introgression between diploid and tetraploid cyto-
formula AACC rather than AADD. If so the C types takes place by means of triploids.
genome would have mediated through A. magna to
A. sativa. AVENA VENTRICOSA Balansa. 2n=14, genome
A diploid sterilis-like type has not yet been found. formula CpCp. This wild species includes spp.
It might be either 2x A. strigosa* or A. longiglu- bruhnsiana (Grüner) Malzew (syn. A. bruhnsiana
mis*. Grüner) and spp. ventricosa (Balansa) Malzew
The parent species of these tetraploids a r e also (syn. A. ventricosa Balansa s. s t r . , genome for-
not determined, although it is thought that one or mula AvAv).
more diploid species (A. clauda*, A. ventricosa*, Spp. bruhnsiana is found in Apsheron Peninsula,
A. longiglumis* and 2x A. strigosa*) have been Azerbeidjan, USSR, while spp. ventricosa is ob-
involved. served is Algeria and Cyprus. The karyotype of
The D genome is probably derived from or related spp. ventricosa is c and of spp. bruhnsiana
to the A genome of A. ventricosa (Steer et a l . , c v s and c v 2 (Rajhathy, 1971).
1970). A. ventricosa is also found in Lybia (Cyrenaica)
Gene exchange exists between the cultivated sativa and Iraq (Ladizinsky & Zohary, 1971).
and byzantina, and sterilis.
Sterilis is a source of resistance to Puccinia c o r o - CHRYSOPOGON GRYLLUS (Tomer) Trin. (syn.
nataCda.f. sp. avenae. Andropogon gryllus T o m e r ) . 2n=20, 40. From
the Mediterranean region to India. Cultivated in
AVENA STRIGOSA Schreb. Black oat, Bristle oat. the Po plain, Italy for its essential oil.
2n=14, genome formula AsAs, 2n=28, genome for-
mula AsAsBB, AABB or AsAsAsAs. The As and HORDEUM VULGARE L. 2n=14. For origin of
B genomes a r e partially homologous and may d e - barley see p. 96. Centre 7 is the centre of origin
rive from a common parent (Ladizinsky & Zohary, of spp. mediterraneum Vav. & Bacht.
1971; Ladizinsky, 1973) The As genome might be
the prototype of the A genome of the polyploid LOLIUM MULTIFLORUM Lam. spp. italicum
species (Rajhathy et a l . , 1971). (A.Br. ) Volkart ex Schinz &Kell. Italian r y e g r a s s .
Ladizinsky and Zohary (1971) included in this 2n=14. The irrigated lands of Lombardy in N.
species the wild A. hirtula Lag. (2n=14), A. Italy. Probably cultivated there in the 13th or 14th
Avena ventricosa (Ladizinsky &Zohary, 1971)

Century (Beddows, 1953). Spread to N. Europe.
LOLIUM PERENNE*
PHALARIS CANARŒNSIS L. Canary g r a s s .

2n=12. The W. Mediterranean region. Canary
Islands, Spain, Portugal. Cultivated for bird seed.
PHALARIS TUBEROSA L. Towoonba grass, Har-

ding g r a s s . 2n=28. The Mediterranean region.
Cultivated in warm countries.
SORGHUM BICOLOR (L. ) Moench. Broomcorn.

2n=20. Sorghum originated in Africa (see p. 116).
The broomcorns were developed in the Mediterra-
nean region from material which came fr om India/
Iran or Africa. Introduced there via the Middle
East from India, Iran or Africa.
SORGHUM HALEPENSE (L. ). P e r s . Johnson

g r a s s . 2n=(20), 40. The Mediterranean region.
A perennial grass. P r i m a r y centre in the east of
Centre 7. Secondary centre runs eastwards to
Indonesia. It has prominent rhizomes and differs
as such from S. bicolor*.
Some new perennial diploid types were selected
from the cross S. halepense x S. bicolor. These Sorghum halepense
A-
I
w
•J - ~ ^ - \ * ^ . y ^ N
~"\ f
Sorghum halepense (deWet & Huckabay, 1967)
types combine high yield and palatibillty with a

degree of frost tolerance and disease resistance.
Their origin is similar to S. almum*.
STIPA TENACISSIMA L. Haifa, Esparto. 2n= ~~^s \ -ft^C\Xl > / »* **l*\JF

The Mediterranean region. In Spain some cultiva- 1 '
tion is done with the cv. Albardin which has a
larger fibre than wild ones. This variety seems In' *'*£5>&>'^V « tv¥ v,
to have developed there. In N. Africa and Spain * i/ i M L / 5 / rvrx^-fr-
wild halfa yields a paper-making fibre. \
v
TRITICUM TURGIDUM spp. turgidum conv. durum ^ V -ô '—~-~^~ x -<P"J
/ ^--' ! \ <\ V -
(Desf. ) Mac Key. 2n=28, genome formula AABB.
It originated under cultivation of emmer (p. 84). Spread of maize in the West European and the Mediterranean
Secondary centre in the Mediterranean region. region, indurata ( ), indentata ( ), century of introduction
(roman number) (Brandolini, 1970)
ZEA MAYS L. 2n=20. Maize was domesticated in
C. America (p. 166). Secondary centres in the
north of the Mediterranean region and in the Nile
basin (Brandolini, 1970).
Grossulariaceae MENTHA AQUATICA L. (syn. M. citrata Ehrh. )

Bergamot mint. 2n=(36, 60), 96, genome formula
RIBES MULTIFLORUM Kitt. 2n=16. The Medi- R a R SSJJA q A q , c. 96. S. Europe, Asia and
terranean region. It is one of the parental species N. Africa. It is a source of an essential oil. Its
of the present-day red currant cultivars (p. 135). A ac l genome is partial homologous to the A genome
of M. arvensis var. piperascens* (Ikedo &Ono,
Hippocastanaceae 1969). It is one of the parents of M. x piperita*.
AESCULUS HIPPOCASTANUM L. Horse chestnut. This species is cultivated as M. citrata in the
2n=40. The central part of the Balkan peninsula, USA for its lavender-like oil for perfumery (Todd
E. Bulgaria, W. Iran and the Himalayas. Culti- &Murray, 1968).
vated as an ornamental, shade t r e e and for its Patented hybrids of this species with M. crispa L.
timber. A. carnea Hayne (2n=40, 80) is a hybrid (syn. M. spicata* var. crispata Schrad. ) a r e also
with the N. American A. pavia L . , Red buckeye cultivated in the USA (M.J. Murray, p e r s . c o m m . ,
(2n=40). 1971).
Iridaceae MENTHA LONGIFOLIA (L. ) Huds. (syn. M.

spicata L. var. longifolia L., M. sylvestris L. ).
CROCUS SATIVUS L. Saffron crocus. 2n=(14, 16), Horse mint. 2n=18, 24, (27, 36, 48). S. and C.
24, (40). The Mediterranean region and Ante- Europe, N. Africa, Ethiopia, Arabia, Ante and
Asia. Cultivated since ancient times for its styles C. Asia. Formerly it was much cultivated. At
which a r e a source of saffron. Cultivated f o r m e r - present only var. crispa Benth. is cultivated. It is
ly for this purpose in Europe and N. America and related to M. rotundifolia* and M. spicata*.
now in S. Europe, Asia Minor, Iran, N. India and
China. MENTHA PULEGIUM L. Penny royal, Pudding
The actual origin of the present cultivars is not g r a s s . 2n=(10), 20, (30), 40, (40-42). The Medi-
known. terranean region, Europe to Iran. Cultivated for-
merly in Europe and elsewhere.
IRIS GERMANICA L. German iris, Flag i r i s .
2n=24, (34, 36), 44, 48, (60). The Mediterranean ROSMARINUS OFFICINALIS L. Rosemary. 2n=24.
region. A perennial herb widely cultivated as an The Mediterranean region. Cultivated for ornament
ornamental and also for its rootstocks used for and for its aromatic oils.
perfumery.
SALVIA OFFICINALIS L. Sage. 2n=14, (16). The
Labiatae Mediterranean region. A culinary herb cultivated
HYSSOPUS OFFICINALIS L. 2n=12. The Medi- now in many gardens in temperate and tropic coun-
terranean region, Asia Minor and Iran. Cultivated tries.
for its essential oil, as a medicinal plant and as an
ornamental. SALVIA SCLAREA L. Clary sage. Clary wort.
2n=22. The Mediterranean region to Iran and
LAVANDULA LATIFOLIA Medik. Broadleaved Transcaucasia. Cultivated formerly in the Medi-
lavender. 2n=54. Cultivated in S. France and terranean region and S. Europe for various p u r -
occasionally in C. Europe for its Oil of Spike. poses, e.g. for flavouring wine and beer.
The cultivated plants are often hybrids with L.
officinalis*. SALVIA VIRIDIS L. Bluebeard. 2n=16. The Medi-
terranean region to Iran. Cultivated here and
LAVANDULA OFFICINALIS Chaix. (syn. L. an- there for its oil to flavour wine and beer.
gustifolia Mill., L. spica L. ). Lavender, 2n=(36),
54. P r i m a r y centre in Centre 7. An old cultivated SATUREJA HORTENSIS L. (incl. S. laxiflora C.
plant for perfumery. First it was used as an in- Koch and S. pachyphylla C. Koch). Summer savory.
sect repellant. Many cultivated varieties a r e 2n=45-48. The Mediterranean region, C. Europe
hybrids with wild plants and L. latifolia*. and Siberia. Cultivated for its oil of savoury and
as a potherb.
MAJORANA HORTENSIS Moench. (syn. Origanum
majorana L. ). Majoram. 2n= . S. Europe, SATUREJA MONTANA L. (syn. S. obovata Lag.,
the Mediterranean region to India. Cultivated in S. illyrica Host). Winter savory. 2n=12, 30. The
Europe and elsewhere as a medicinal plant. Mediterranean region to Ukraine. Cultivated in
S. Europe and in Germany.
MELISSA OFFICINALIS L. Common balm. 2n=32,
64. E. Mediterranean region to Caucasia, SW. TEUCRIUM CHAMAEDRYS L. (syn. T. officinale
Siberia, S. Iran, Turkestan and Syria. Cultivated Lam. ). Common germander. 2n=32, 60, 64. The
formerly in Europe and elsewhere for diverse Mediterranean region, France, C. Germany to
purposes. It is suggested that the commonest cul- S. Ural, Iran, N. Syria and Morocco. Cultivated
tivated type var. officinalis is derived from var. formerly as a medicinal crop.
hirsuta P e r s . (syn. M. hirsuta (Pers. ) Hörnern. ),
a variety from the Balkans. TEUCRIUM MARUM L. 2n= . The W. Medi-
terranean region and S. France. Cultivated in S.
Europe. Cultivated formerly in Germany.
GROSSULARIACEAE - LEGUMINOSAE 101
THYMUS VULGARIS L. Thyme. 2n=30. The Me- LATHYRUS HIRSUTUS L. Rough pea, Caley pea,
diterranean countries. Cultivated now in temperate Singletary pea. 2n=14. The Mediterranean region.
and tropical countries. Cultivated especially in USA as a pasture hay,
winter cover and soil improvement crop.
Lauraceae
LATHYRUS OCHRUS DC. 2n=14. The Mediterra-
LAURUS NOBILIS L. Laurel, True bay, Sweet nean region. Cultivated in Greece occasionally as
bay. 2n=42, 48. The Mediterranean region. P r i - a fodder.
mary centre also there. Cultivated there and
elsewhere for its leaves which a r e used as a con-
LATHYRUS ODORATUS L. Sweet pea. 2n=14. The
diment.
Mediterranean region. Seeds of wild plants were
sent from Sicily to NW. Europe in 1667 by a monk,
Leguminosae Francesco Cupani. It is commonly cultivated as
ASTRAGALUS BOETICUS L. 2n=16, 30. S. Europe an ornamental. Its flowers a r e also used as a
and Mediterranean region. Cultivated as a sub- source of an essential oil.
stitute for coffee.
LATHYRUS SATIVUS L. Grass pea, Chickling pea.
CERATONIA SILIQUA L. Carob, St. John's bread. 2n=14. Probably domesticated in W. Asia (p. 73).
2n=24. The Mediterranean region, Syria and ad- P r i m a r y centre in the Mediterranean region.
jacent countries. P r i m a r y centre in Centre 7.
Cultivated especially on Cyprus as a fodder crop. LATHYRUS TINGITANUS L. Tangier pea. 2n=14.
The fruits a r e eaten, while the seeds a r e used to The Mediterranean region. A micro-centre is in
p r e p a r e carob coffee. More uses a r e given by Morocco. Cultivated as a winter annual, also in
Uphof (1968). USA.
CERCIS SILIQUASTRUM L. Judas t r e e . 2n=14. LOTUS EDULIS L. Asparagus pea, Winged pea.
A t r e e of the Mediterranean region to Crim and 2n=14. The Mediterranean region to Asia Minor
Iran. Cultivated for its leaves (vegetable). and Syria. Occasionally cultivated for its young
pods.
CICER ARIETINUM L. Garbanzos, Chickpea.
2n=14, 16, (24, 32, 33). Probably W. Asia (p. 85). LUPINUS ALBUS L. (syn. L. sativus Gaertn. ).
Secondary gene centre in the Mediterranean area. White lupine. 2n=50. Wild in Corsica, Sardinia,
Especially large-seeded types, race m e d i t e r r a - Sicily and Israel. P r i m a r y centre is in Centre 7.
neum Pop. a r e cultivated. Domesticated in Spain and in N. Africa. All cul-
tivars have white seeds, the production of pigment
CYTISUS CANARIENSIS (L. ). O. Kuntze. Genista. being suppressed by two independent pairs of in-
2n=46. Canary Islands. Cultivated elsewhere. hibitor genes. These genes must already have been
Used in Mexico as hallucinogen. selected for by farmers some 4 000 years ago
(Kazimierski, 1960).
CYTISUS PALLIDUS Poir. 2n= . Canary Islands. L. albus is closely related to L. termis*. Accor-
Cultivated as a forage crop. ding to Kazimierski (1960) both derive from L.
graecum (see L. termis*). This species would
CYTISUS PROLIFER Kit. Tree lucerne, Tree a l - derive from L. jugoslavicus Kazim. &Now.
falfa, Tagasaste, Escabon. 2n=48. Canary Islands (2n=50). It is found in Yugoslavia. Gladstone
(Uphof (1968) says Hungary). Cultivated there as a (1970) considered L. t e r m i s , L. graecus and L.
forage plant. Introduced into New Zealand. jugoslavicus as synonyms of L. albus.
GLYCYRRHIZA GLABRA* LUPINUS ANGUSTIFOLIUS L. (syn. L. varius L.,

L. linifolius Roth, L. reticulatus Desv. ). Narrow-
HEDYSARUM CORONARIUM L. Spanish esparcet. leaved lupine, Blue lupine. 2n=40. The primary
2n=16. The Mediterranean region. Cultivated as a centre in the Mediterranean region. The present
fodder crop. European cultivars probably derive from wild
types from Palestine. Cultivated also in S. Africa
LATHYRUS ANNUUS L. 2n=14. The Mediterranean and Australia. Widely cultivated as an ornamental
region and Portugal. Sometimes cultivated as a too.
fodder.
LUPINUS COSENTINI Guss. (syn. L. varius L . ,
LATHYRUS CICERA L. Vetchling, Flat pod pea spp. varius Franco &P . Silva). Western Austra-
vine, J u r o s s e , Garousse. 2n=14. The Mediterra- lian blue lupine, Sandplain lupine, Geraldton l u -
nean region, Canary Islands, Iraq, Iran and pine. 2n=32. Coastal Morocco and other sites in
Transcaucasia. Cultivated in S. Europe as a W. Mediterranean region. Introduced into W.
fodder crop and as a green manure. Australia about the middle of the 19th Century and
has become naturalized. Since 1910 it has been
LATHYRUS CLYMENUM L. (syn. L. purpureus cultivated for summer sheep food and soil improve-
Desf., L. alatus Sibth. &Sm. ). Cicerchia porpo- ment. Described as L. pilosus L . , L. varius L.
rina. 2n=14. The Mediterranean region and Madei- and L. digitatus Forsk.
ra. Cultivated in S. Europe.
LUPINUS LUTEUS L. (European) yellow lupine. SW. France, N. half of the Iberian Peninsula and
2n-(46, 48. 50). 52. Mediterranean countries. the Azores. Cultivated elsewhere.
P r i m a r y centre in Centre 7. The present European A related species is O. isthmocarpus Coss. (syn.
cultivars derive probably from wild Palestinean O. sativus spp. isthmocarpus (Cosson) Dostal)
plants. Cultivated as a fodder crop, green manure (2n=14). A Mediterranean-Atlantic species, where
and as an ornamental. Closely related to L, h i s - it grows together with O. sativus, hybrids, d e s -
panicus Boiss. & Reut. and L. rothmaleri Klink cribed as O. macrorrhynchus (Willk. ) Klinkowski
(2n=50, 52). &Schwz. (syn. O. sativus v a r . macrorrhynchus
Willk. )a r e found.
LUPINUS PILOSUS L. (syn. L. varius L. spp.
orientalis Franco &P . Silva). Greater blue lupine, PISUM SATIVUM L. spp. hortense Asch. & Graeb.
Mairy lupine. 2n=42, (50). NE. Mediterranean Garden pea. 2n=14. Secondary centre in the Me-
region. It may occasionally be cultivated. It is diterranean region. Spp. hortense is domesticated
characterized by its big seeds, the biggest of all in SW. Asia (p. 86).
Lupinus-species.
PISUM SATIVUM spp. jomardi (Schrank) Alef.
LUPINUS TERMIS Forsk. (syn. L. graecus B o i s s . , (syn. ecotype arvense s s t r . , P . elatius (M.B. )
L. albus spp. albus). Egyptian lupine. 2n= Stev., P . jomardi Schrank, P . transcaucasicum
Palestine and Egypt. Cultivated in Egypt since Stankov). 2n=14. Cultivated in Egypt.
ancient t i m e s . The seeds contain alkaloids, which
have to be removed before consumption. Closely PSORALEA BITUMINOSA L. Asphalt clover.
related to L. albus*. L. termis and L. graecus 2n=20. The Mediterranean region and Canary
may be varieties of L. albus*. Islands. Cultivated for cattle food.
MEDICAGO HISPIDA Gaertn. Bur clover. 2n=14, SPARTIUM JUNCEUM L. Spanish broom, Weaver's
(16). The Mediterranean region. Cultivated as a broom. 2n=48. 52, 52-56. The Mediterranean
green manure, for pasture and as a hay crop. regions and Europe. Some cultivation is done in
France near Aspiran (Hérault).
MEDICAGO SATIVA L. Lucerne, Blue alfalfa.
2n=(16, genome formula SS), 32 genome formula TRIFOLIUM ALEXANDRINUM L. Alexandrian
SSSS, 64, Transcaucasia (p. 86). Secondary cen- clover, Egyptian clover, Berseem. 2n=16. The
t r e in N. Africa, especially Algeria. The great E. Mediterranean regions. Cultivated in the Near
variability mayhave arisen due to introgression East and India. It is the oldest clover cultivated
of M. gaetula which is endemic in N. Africa. and is closely associated with agriculture in Egypt.
P r o s t r a t e types a r e found in the mountains of Secondary centre in Egypt. T. alexandrinum is
Anatolia, Turkey. closely related to T. berytheum Boiss. (syn. T.
alexandrinum var. berytheum) and T. salmoneum
MELILOTUS INFESTUS Guss. 2n=16. A plant of Mout. These species a r e self-incompatible, while
W. Mediterranean region being a source of r e - T. alexandrinum is self-compatible. This may be
sistance to the sweet clover weevil of M. albus* a domestic characteristic.
and M. officinalis*.
TRIFOLIUM FRAGIFERUM L. (syn. T. neglectum
MELILOTUS MACROCARPA Coss. &Dur. 2n=16. Fisch &Mey). Strawberry clover. 2n=16. Europe,
N. Africa. Cultivated in Algeria for its large Canary Islands, Madeira, N. Africa and W. Asia.
fruits used as spice. Cultivated as a fodder crop.
MELILOTUS SULCATUS Desf. 2n=16, (32). S. TRIFOLIUM INCARNATUM L. Crimson clover.

Portugal and the Mediterranean region. Plants b e - 2n=14. C. and S. Europe, in the Balkans andN.
longing to spp. brachystachus Maire a r e coumarin Africa. The cultivated type (var. sativum D u c ,
deficient and resistant to drought, and most pests spp. incarnatum) probably derives from the wild
including the sweet clover weevil. Spp. segetalis var. molinerii (Balbis ex Hörnern. ) Syme. The
(Brot. ) Maire has also been described as M. s e g e - latter is found in Spain. Cultivated since long in
talis Ser. (2n=16). Catalonia, Spain and S. France. Spread to E. and
N Europe and later to N. America.
ORNITHOPUS COMPRESSUS L. 2n=14. Spain,
Portugal andthe Mediterranean regions. Its TRIFOLIUM ISRAELITICUM D. Zoh. &Katzn.
northermost point of occurrence is Bretagne, (syn. T. subterraneum L. var. telavivensis Eig).
France. It has a high leaf and seed production. It 2n=14. N. Israel. It is not a parent of T. s u b -
might be useful as a breeding source for O. s a t i - terraneum as has been suggested. It only has 14
vus*. chromosomes while T. subterraneum has 16.
F o r m e r l y it was considered as the "Israeli r a c e "
ORNITHOPUS SATIVUS Brot. Serradella. 2n=14. of this species.
Wild plants in NWPortugal, N. Spain andSW.
France. From here its cultivation spread over TRIFOLIUM REPENS L. var. giganteum. Lodi
W. and N. Europe, since the beginning of the 19th clover, Ladino clover. 2n=32. Probably, Lodi,
Century. A green manure plant. N. Italy. Cultivated first in N. Italy andthe
Spp. sativus (syn. O. roseus Dufour) is native to Netherlands (p. 137). An excellent fodder crop.
LEGUMINOSAE - MORACEAE 103
TRIFOLIUM SUBTERRANEUM L. Subterranean Schultze-Motel (1972) found no evidence for a

clover, Sub clover. 2n=16. The Mediterranean supposed division of the small-seeded type into
region, in SE. andW. Europe, the Caucasian two geographical r a c e s : a long-seeded type in the
region and N. Iran. It is possible that the west- W. part of the Mediterranean region and a round-
ward migration followed the course of clearing seeded type in the eastern part. Sothere is no
and cropping by man (Katznelson &Morley, 1965a, reason to suppose that the broad bean originated
1965b). Secondary centre in Australia (p. 58). in two separate a r e a s . Whether V. pliniana
Naturalized in Australia, S. Africa and N. and (Trabut) Moratova found in Algeria and Morocco,
S. America. is a type of V. faba or a related species in not
T. subterraneum can be divided into three sub- known.
species: 1. spp. subterraneum (syn. T. blesense
Dodart) which is the commonest taxon sympatric VICIA NARBONENSIS L. Narbonne vetch. 2n=14.
with the species, 2. spp. yanninicum Katzn. & SW. Asia (p. 87). Secondary gene centre in the
Morley, which occurs in Istria, Dalmatia, Alba- Mediterranean areas where it is cultivated.
nia, Serbia and N. Greece and 3. spp. brachy-
calycinum Katzn. &Morley (syn. v a r . oxaloides Liliaeeae
Eig) which occurs from W. Thracia to the C a s -
ALOE BARBADENSIS Mill. (syn. A. vera L.).
pian Sea. These subspecies a r e almost completely
Curapao aloë, Barbados aloë. 2n=(10), 14. The
intersterile (Katznelson &Morley, 1965a).
Mediterranean region, S. Arabia, E. Africa, NW.
The existence of two closely related but more
India and S. China. The wild var. barbadensis of
primitive species (T. batmanicum Katzn. (syn.
the Mediterranean region has run wild in C. A m e -
T. anatolicum Katzn. ) (2n=16) in Diyarbakir P r o -
rica, W. Indies to Bolivia. It probably came there
vince, and T. chlorotrichum Boiss. &Balansa
through Spain. Cultivated in W. Indies.
(2n= ) in Phrygia) in Turkey and the absence
of these and other close relatives elsewhere indi-
cates the origin of T. subterraneum in Turkey. LILIUM CANDIDUM L. Madonna lily, Bourbon
However, the greatest variation is found in Greece lily. 2n=24. The Mediterranean region andSW.
(Katznelson &Morley, 1965a). Asia. Cultivated especially in S. France for its
Railey and Francis (1971) found that the isoflavone flowers. These a r e a source of an essential oil.
pattern of T. batmanicum closely resembles that It is the oldest lily of the European gardens.
of spp. brachycalycinum. They concluded that
T. batmanicum might be the ancestor species of URGINEA MARITIMA (L.) Baker, (syn. U. Scilla
T. subterraneum while spp. brachycalycinum is Steinh. ). Sea onion. 2n=20, (30), 40, 60. Medi-
probably the earliest forms of subterranean clo- terranean coast; the highest concentration in E.
ver. They postulated that spp. subterraneum Algeria. Wild and cultivated plants a r e used for
evolved later and colonized a wider range of e n - their pharmaceutic and rat toxic properties.
vironments.
Linaceae
Furthermore, the isoflavone pattern of T. batma-
nicum is very similar to that of T. globosum L. LINUM USITATISSIMUM L. Flax, Linseed.
(syn. T. radiosum Wahlenb., T. nidificum G r i - 2n=30, (32). For origin see p . 87. In the Medi-
seb. ) (2n=16). However, this species belongs to terranean region the oil flax (spp. mediterraneum
another subsection. Vav. &Ell.) is cultivated. In Italy hybrid forms
(spp. transitorium Vav. &EU. ) of spp. e u r a s i a -
TRIFOLIUM VAVILOVn Eig. 2n=16. Israel. ticum and spp. mediterraneum a r e found. L a r g e -
seeded types a r e cultivated in N Africa. Those
VICIA ARTICULATA Hörnern. One-flowered from Algeria are a source of F u s a r i u m - r e s i s t a n -
vetch. 2n= . The Mediterranean region, Asia ce.
Minor, Madeira and Canary Islands. Cultivated.
Malvaceae
VICIA BENGHALENSIS L. (syn. V. atropurpurea
Desf. ). Purple vetch. 2n=12, 14. The Mediterra- ALTHAEA OFFICINALIS L. Marsh mallow.
nean area. Naturalized in the USA. Cultivated 2n=42, (c. 42, 40-44). Europe, E. Mediterranean
there as a cover crop and green manure. region and W. Asia. Cultivated for its roots which
a r e a source of medicine.
VICIA CALCARATA Desf. Demehi. 2n=12, 14.
The Sahara oasis. There it is cultivated for its ALTHAEA ROSEA (L.) Cav. Garden hollyhock,
seeds. Hollyhock. 2n=(26), 42, (56). Asia Minor, Balkan
and Crete. Ran wild in Italy, S. France and S.
VICIA ERVILEA (L.)Willd. Bitter veteh, Ervil. Tirol. Cultivated in Europe since the 16th Cen-
2n=14. P r i m a r y centre in the Mediterranean area. tury especially var. nigra hort. which has blackish
Cultivated in Spain. A characteristic group deve- purple petals. These a r e used to colour wine and
loped in Asia Minor (p. 87). as medicine. Cultivated now in many types as an
ornamental.
VICIA FABA L. (syn. Faba vulgaris Moench).
Field bean, Broad bean, Horse bean, Pigeon bean, Moraceae
Tick bean, Windsor bean. 2n=12, (14). SW. Asia FICUS SYCOMORUS L. Sycomore fig. 2n=26. A
(p. 73). Unknown wild. t r e e from N. Africa.
Myrtaceae Papaveraceae
EUCALYPTUS CAMALDULENSIS Dehn. Longbeak PAPAVER SOMNIFERUM L. Opium poppy. 2n=22.
eucalyptus. 2n=22. P r i m a r y centre in Australia P r i m a r y centre in Centre 7 or in the Central Asian
(p. 58). Secondary centre in Centre 7 and in S. centre (p. 74). It probably derives from ssp.
America (p. 155). It was believed that the t r e e s setigerum (DC. ) Corb. (syn. S. setigerum DC.)
of this species cultivated in Israel came from S. which occurs wild in W. Mediterranean region and
Australia, but the leaves of the Israeli t r e e s con- elsewhere as a weed. The cultigen (spp. somni-
tain three polyphenols which have not been found ferum incl. spp. hortense (Hussenot) Corb. ) is
in this species anywhere in Australia. cultivated for its medicinal and narcotic latex,
and seeds for its oil and as an ornamental. It may
EUCALYPTUS GLOBULUS Labell. Fever t r e e . escape from cultivation.
Blue gum. 2n=20, 22, 28. SE. Tasmania. Culti-
vated. Secondary centre in Centre 7. Pinaceae
PINUS PINEA L. Stone pine, Pinie. 2n=
MYRTUS COMMUNIS L. Myrtle. 2n=22. The Me- S. Europe. A tree often cultivated for its edible
diterranean region and SW. Europe. Cultivated
seeds.
since ancient times for its fruits and for its medi-
cinal properties.
Pistaciaceae
Oleaceae PISTACIA LENTISCUS L. Lentisk pistache. 2n=24.
The Mediterranean region. A small t r e e cultivated
FRAXTNUS ORNUS L. Flowering ash, Manna ash.
for its chewing gum.
2n=46. A tree of C. and E. Mediterranean region
Cultivated on the N. coast of Sicily.
PISTACIA TEREBINTHUS L. Terebinth pistache.
2n= . The Mediterranean region. On the Greek
OLEA CHRYSOPHYLLA Lam. Golden-leaved islands a type with big fruits and large leaves was
olive t r e e . 2n= . Wild over a large part of cultivated.
the Old World, including the Mediterranean region.
It is possibly the wild ancestor of O. europaea*.
Plantaginaceae
If so, it is a synonym of O. europaea var. sylves-
t r i s Brotero. PLANTAGO INDICA L. 2n=12. C , S. and E.
Europe and W. Asia. Cultivated in S. France as a
OLEA EUROPAEA L. Olive t r e e . 2n=46. The medicinal herb (Mansfeld, 1959).
Mediterranean region. P r i m a r y centre in Centre
7. Its domestication started there in ancient PLANTAGO PSYLLIUM L. Psyllium. 2n=12.
t i m e s . Var. sylvestris Brotero includes the wild Mediterranean region.
forms and the possible naturalized cultivated
types. Var. europaea is the cultivated form. The Ranunculaceae
main difference between these varieties a r e spiny
AQUILEGIA VULGARIS L. Columbine. 2n=14.
lower branches and small leaves and drupes which
S. and C. Europe (p. 138), N. Africa and temp.
a r e present in var. sylvestris. Some cultivars a r e
Asia. Cultivated widely as an ornamental, f o r m e r -
developed as table olive varieties, others for oil.
ly also for medicinal purposes.
See also O. chrysophylla*.
NIGELLA SATIVA L. Black cummin. 2n=12. C.
(p. 138) and S. Europe, N. Africa and W. Asia.
Cultivated for its seed in the Mediterranean region
and in the Orient. Cultivated formerly in C.
Europe.
Resedaceae
RESEDA LUTEOLA L. Weld. 2n=24, (26, 28).
C. Europe (p. 138), Mediterranean region, Iran
and Afghanistan. Cultivated formerly as a source
of deep yellow dye.
RESEDA ODORATA L. Mignonette. 2n=12, (14).

N. Africa. Material was sent to P a r i s between
Olea europea (Polunin &Huxley, 1972)
1733 and 1737. From here its cultivation spread
as a perfumery plant (Mansfeld, 1959).
Palmae
RESEDA PHYTEUMA L. 2n=12. The Mediterra-
CHAMAEROPS HUMULIS L. Dwarf palm. 2n=36. nean region and Asia Minor (p. 38). It is occasio-
Wild in the W. Mediterranean regions. Cultivated nally grown as a vegetable.
in some parts of Morocco. Often planted as an
ornamental. A source of fibre (crin vegetable). Rhamnaceae
RHAMNUS CATHARTICUS*
MYRTACEAE - UMBELLIFERAE 105
RHAMNUS FRANGULA* Ulmaceae

CELTIS AUSTRALIS L. (syn. C. excelsa Salisb.).
ZIZIPHUS LOTUS (L. ) Lam. 2n=24. The Medi- Hackberry. 2n=40. The Mediterranean region. A
terranean region. A t r e e cultivated in Italy, S. tree cultivated there as an ornamental and in Asia
Spain and Egypt. It is probably the lotophagus of Minor for its edible fruits (Mansfeld. 1959).
the ancient peoples of Lybia.
Umbelliferae
Rosaceae
AMMADAUCUS LEUCOTRICHUS (Coss. & Bur.
AMYGDALUS PERSICA L. Peach. 2n=16. P r i m a -
2n=16. N. Africa. Cultivated there.
ry centre in China (p. 36). Secondary centre in
Italy and Spain.
AMMI MAJUS L. (syn. Apium arami Crantz).
Bishop's weed. 2n=22. The Mediterranean region
CRATAEGUS AZAROLUS L. (syn. C aronia
to Iran andto Switzerland and Belgium. Cultivated
B o s c ) . Azerolier. 2n= . S. Europe, N.
Africa and the Orient (p. 74). This shrub or small since the Middle Ages for its aromatic seeds.
t r e e is often cultivated for its edible fruits. Var.
ANETHUM GRAVEOLENS L. (syn. A. s o w a K u r z . ) .
aronia L. is found wild on Crete.
Satapashpi, Sowa, Suwa. 2n=22. Eurasia. Culti-
vated in India. It has longer fruits than the Indian
Rutaceae type (p. 70).
CITRUS AURANTIUM L. spp. bergamia (Risso&
Poit. )Wight & Arn. Bergamot. 2n=18. Calabria, APIUM GRAVEOLENS L. Celery. 2n=22. Culti-
Italy. P r i m a r y centre probably in SE. Asia vation started in the Mediterranean region. The
(p. 55). Cultivated for its bergamot oil in Calabria, wild parent A. graveolens var. silvestre P r e s l .
Italy. (syn. var. graveolens) is cosmopolite. Not much
is known about the development of the three bota-
CITRUS LIMON (L.) Burm. Lemon. 2n=18, (36). nical varieties A. graveolens, var. silvestre f.
P r i m a r y centre probably in SE. Asia (p. 55). secalinum Alef. (syn. var. secalinum Alef. ), the
Secondary centre in this region, especially in leafy celery, smallage or soup celery, var. r a p a -
Sicily. ceum (Mill. ) DC., the celeriac, turnip-rooted
celery or German celery and v a r . dulce (Mill.)
CITRUS SINENSIS (L.) Osbeck (syn. C. aurantium P e r s . , the blanching celery (Becker, 1962).
L. v a r . sinensis L.). Sweet orange. 2n=18, (27,
36). P r i m a r y centre probably in S. China or CORIANDRUM SATIVUM L. Coriander. 2n=22.
Cochin-China (p. 55). Secondary centres in Israel The Mediterranean region and W. Asia. Cultivated
(e.g. the varieties Shamuti, Beladi, Khalili) and for its aromatic fruits.
in Spain (e.g. the variety Valencia, the blood
orange). CRITHMUM MARITIMUM L. Samphire, Sea s a m -
phire, Sea fennel, Piercestone. 2n=20, (22).
RUTA CHALEPENSIS L. Fringed rue. 2n=36. The Canary Islands, Madeira, coasts of Portugal to
Mediterranean region. Cultivated there and e l s e - S. England and those of the Mediterranean region
where as a medicinal plant. and Crimea. Cultivated in the USAas a kitchen
plant.
RUTA GRAVEOLENS L. Common rute, Rue.
2n=72, 81. Wild in the Mediterranean regions. CUMINUM CYMINUM L. Cumin. 2n=14. The
Introduced in many tropical countries. The leaves Mediterranean region to Turkestan. Cultivated in
a r e used as condiment and medicinally. SE. Europe, N. Africa, India and China.
Scrophulariaceae DAUCUS CAROTA L. Yellow carrot. 2n=18. Wild

species from Afghanistan (p. 76)to the Mediterra-
DIGITALIS PURPUREA* nean area. Although yellow carrots may have
arisen in other areas where purple carrots were
Solanaceae cultivated, it is thought that the true yellow c a r -
ATROPA BELLADONNA L. Belladonna. 2n=72. rots developed in Region 7 from crossing with
From Spain, the Balkan, Asia Minor to India the wild D. carota spp. maximum.
(p. 70). Cultivated in Europe, India and USAas a
medicinal plant. A. m a r t i a n a F . Q . is considered FOENICULUM VULGARE Mill. (syn. F. officinale
as a hybrid of A. belladonna andA. baetica Willk. Gaertn. ). Fennel. 2n=22. The Mediterranean
(2n=72). The latter species is found in Spain. countries. Cultivated there for a long time and
has been introduced into many other temperate
HYOSCYAMUS NIGER L. Black henbane. 2n=34. countries.
The Mediterranean region. A medicinal plant Var. piperitum (Ucr.) Cout. (syn. F. piperitum
cultivated in some countries to yield alkaloids. ^ c r . , 2n=22) is the Bitter Fennel. Var. dulce
(Mill. ) Thell. (syn. F. dulce Mill. , 2n=22) is the
Florence Fennel, Sweet Fennel or Roman Fennel.
Cultivated for its blanched petioles in S. France
and Mediterranean region. Var. azoricum (Mill.)
Thell. (syn. F. azoricum Mill. ), Carosella or

Italian Fennel, originated in Italy. It has very
broad leaf-stalk bases.
MEUM ANTHAMANTICUM Jacq. Signel. 2n=22.

A herb of C. and S. Europe, once cultivated in
N. England for its roots.
PETROSELINUM CRISPUM (Mill.) Nym. e x A . W .

Hill (syn. Carum petroselinum Benth. ). Parsley.
2n=22. S. Europe. Widely cultivated there and
elsewhere.
Mansfeld (1959) has classified the wild and culti-
vated types.
SIUM SISARUM L. Skirret, Chervin. 2n=20, 22.

E. Asia and Mediterranean region. Occasionally
cultivated for its edible tuberous roots (var. s i s a -
rum).
SMYRNIUM OLUSATRUM L. Alisander, Alexan-

ders, Maceron. 2n=22. Mediterranean region,
S. andW. Europe and Caucasus. Was much culti-
vated but is now replaced by celery.
Urticaceae
SOLEIROLIA SOLEIROLII (Req. ) Dandy. 2n^
The islands of W. Mediterranean region. Cultiva-
ted.
Valerianaceae
FEDIA CORNUCOPIAE Gaertn. African valerian,
Valériane d'Alger. 2n-32. The Mediterranean
region. Cultivated as a potherb and during famine.
VALERIANA ERIOCARPA Desv. Italian corn s a -

lad. 2n= . The Mediterranean region. Culti-
vated for salad.
Verbenaceae
VITEX AGNUS-CASTUS L. Chaste t r e e . 2n-24,
32. The Mediterranean region. Cultivated in the
Old and New Worlds for various purposes.
Violaceae
VIOLA ODORATA L. (syn. V. officinalis Cr. ).
Sweet violet. Sweet scented violet, Common
violet. 2n=20. Europe and SW. Asia. Var. parma
is cultivated in N. Italy and S. France as a source
of an essential oil for perfumery.
Vitadaceae
VITIS VINIFERA L. Common grape, European
grape. 2n=38, (40, 57, 76). Centre 7 is one of
the three primary centres of diversity. On p. 90
the domestication of the grape is discussed and
other data a r e presented. Several secondary cen-
t r e s a r e observed, e.g. the varieties for currants
in Greece, and the varieties for wine in Italy,
Spain and Algeria.
8 African Centre
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The African Centre includes Vavilov's Abyssian Centre of Origin. Darlington (1956) described two areas
in Africa as region of origin: Ethiopia and C. Africa. Porteres (1950) established three cradles of
agriculture in Africa viz. 1. the Mediterranean cradle which lies in Centre 7, 2. the Ethiopian cradle
3. the East African cradle and 4. the West African cradle. The West African was the most important and
could be subdivided into A. the Senegambian 'subcradle', B. the Central Niger 'subcradle', C. the Benin
'subcradle' and D. the Adamawa 'subcradle'. The independent origin of agriculture in W. Africa has
been supported by Anderson (1960) and especially by Murdock (1960). However, this claim has been r e -
futed by for instance Wrigley (1960), Clark (1962), Baker (1962), Harris (1967) and Harlan (1972). They
found grounds to suppose that the knowledge of how to apply agriculture had been introduced from Centre
6, while the tropical African crops had been locally domesticated.
The African centre has a very important influence on the crops of the world. Many crops have an
African origin. Examples a r e Brassica juncea, Ceiba pentandra, Coffea s p . , C o l a s p . , Cucumis s p . ,
Ensete ventricosum, Gossypium s p . , Hibiscus s p . , Lablab purpureus, Oryza s p . , Pennisetum s p . ,
Phoenix s p . , Ricinus communis, Sesamum indicum, Setaria s p . , Sorghum bicolor, Vigna unguiculata etc.
AFRICAN CENTRE 108
Acanthaceae FUNTUMIA ELASTICA (Preuss) Stapf. Lagos silk

rubber. 2n=22. W. Africa. Large plantations of
JUSTICIA INSULARIS T. And. 2n= Africa. this tree were established in W. Africa, after the
Cultivated in W. Africa for its leaves. discovery that it was a source of rubber. However,
these plantations cannot compete with the Hevea
Agavaceae rubber.
AGAVE FOURCROYDES Lern. Henequen agave.
2n=c. 140. Yucatan (p. 162). Secondary centre TABERNANTHE IBOGA Bâillon. Iboga. 2n=22.
possibly in Africa. Gabon, Republic of Congo and the NW. Zaire.
Cultivated in Gabon. The roots contain several
DRACAENA ARBOREA Link, D. mannii Baker. indol alkaloids. The most important is ibogaine
Asparagus t r e e , Soap t r e e , D. fragrans (L.) which is a stimulant and in large doses a halluci-
Garol. and D. smithii ex Hook,f., Cocked hat. nogen. Roots of wild plants a r e collected which
Cockade bush. 2n- has almost resulted in the extinction of this plant
These four species a r e native to Africa. Cultivated in several districts in Gabon.
as living fences and live sticks.
SANSEVERINIA GUINEENSIS (L. )Willd. Bow-

string hemp. 2n= . Africa. A fibre crop culti-
vated on a small scale in Mexico.
SANSEVERINIA LONGIFLORA Sims. Florida

bowstring hemp. 2n= . Africa. Cultivated in
Trinidad, S. Florida and S. Carolina.
SANSEVERINIA THYRSIFLORA Thunb. 2n=

S. Africa. Cultivated for its fibres in the tropics.
Tabemanthe iboga (Pope, 1969)
SANSEVERINIA TRIFASCIATA Prain. African
bowstring hemp. 2n= . Trop. W. Africa. Cul-
tivated (often as S. guineensis*) in the tropics.
Var. laurentii (De Wildem. ) N.E. Brown is cul- VINCA ROSEA L. Madagascar periwinkle, Cape
tivated as an ornamental in Congo. periwinkle. 2n=16, (32). Probably Madagascar.
A shrub cultivated as a medicinal plant.
Aizoaceae
Bignoniaceae
MESEMBRYANTHEMUM ANGULATUM Thunb.
Marygold. 2n=18. S. Africa. An annual cultivated KIGELIA AFRICANA (Lam.). Benth. 2n=40. Trop.
in Congo and Mediterranean region as a spinach. W. Africa. Cultivated for medicine and witchcraft.
MESEMBRYANTHEMUM CRISTALLINUM L. Bombacaceae

(syn. Cryophytum cristallinum L. ) N . E . Brown. CEIBA PENTANDRA Gaertn. Kapok tree, Silk
Ice plant. Cristalline. 2n=18. S. Africa. Cultiva-
cotton t r e e . 2n=72, 80, 88. Some authors believe
ted as salad vegetable or as a stabilizer.
in an American/African origin of the kapok tree
(see p. 163). If America is the sole centre of
MESEMBRYANTHEMUM EDULE L. (syn. Carpo-
origin than the African centre is secondary. The
brotus edulis (L. ) L. Bolus). Hottentot fig. 2n=18.
African kapok t r e e is divided in the caribaea-
S. Africa. A dune stabilizer.
forest type and the caribaea-savannah type. The
latter type, which has a broadly spreading crown,
Amaranthaceae is planted in market places. It is possible that
CELOSIA TRIGYNA L. 2n=18. Trop. Africa. this type arose from cuttings of plagiotropic
Madagascar and Arabia. Cultivated as a vegetable branches (Zeven, 1969).
in Africa.
Burseraceae
Annonaceae CANARLUM EDULE Hook. f. (syn. Pachylobus
XYLOPIA AETHIOPICA (Dun. )A. Rich. African edulis G. Don, Dacryodes edulis (G. Don. ) Lam).
pepper, Guinea pepper. Ethiopian pepper. Spice Bush butter tree, Native pear. 2n= . Trop.
t r e e . 2n= . Trop. Africa. (Semi-)cultivated Africa. Occasionally cultivated for its edible
in W. Africa. fruits.
Apocynaceae COMMIPHORA OPOBALSAMUM Engl. Mecca

myrrh tree, Harobol myrrh. 2n= . Arabia and
CARISSA GRANDIFLORA A.DC. Natal plum. Somaliland. Formerly (llth-17th century) culti-
2n=22. S. Africa. Cultivated for its fruits and as vated in Egypt and Palestina (Mansfeld, 1959).
an ornamental.
ACANTHACEAE - CUCURBITACEAE 109
Cannaceae tard. Leaf mustard, Indian mustard. 2n=36, g e -

nome formula AABB. Africa. Spread to Asia, E.
CANNA SPECIOSA Rose. 2n= . W. Africa. Europe and China. Now distributed from Europe
Cultivated in Sierra Leone. It is the source of to E. Asia. Often as a weed. It is cultivated for
African T u r m e r i c . The tubers resemble those of its oily seeds.
Curcuma longa*.
This species originated from the amphiploidization
of B. campestris* and B. nigra*.
Celastraceae
Through artificial amphiploidization of hybrids of
CATHA EDULIS Forsk. Kat, Flower of paradise, the parental species it has been possible to intro-
Miraa. 2n= . Ethiopia/Somalia to Cape of duce characteristics of both parents into sarepta
Good Hope, and also in Yemen and Saudi Arabia mustard.
where it probably has been introduced. Cultivated
in Ethiopia and Arabia. Its leaves a r e used to ERUCA PINNATIFIDA (Desf. ) Pomel. 2n=
prepare African, Abyssinian, Arabian or wild tea. The Sahara. Occasionally cultivated in oases for
The leaves and shoots a r e also eaten (Margetts, fodder.
1967).
LEPIDRJM SATIVUM L. Garden c r e s s , Common
Cleomaceae c r e s s . 2n=16, 32. Wild type var. silvestre Thell.
GYNANDROPSE3 GYNANDRA (L. ) Briq. (syn. G. From Sudan a r e a to the Himalayas. Cultivated in
pentaphylla DC). Cat's whiskers. 2n=30, 32, 34. ancient times in Europe as a vegetable. It may
(Sub)trop. Africa and India. Cultivated in Africa, have reached Europe from the Levant as a flax
in the West Indies and in Malaya. Used as vege- weed. In Africa there a r e red, white and black
table and as an ornamental. varieties. There the seeds a r e used for medicinal
purposes and for oil production. Also used as a
vegetable.
Compositae
CRASSOCEPHALUM BIAFRAE S. Moore. 2n= Cucurbitaceae
W. Africa. Cultivated as a vegetable. Several
types a r e known (Terra, 1967). CITRULLUS LANATUS (Thunb. ) Mansf. (syn. C.
vulgaris Schrad. ). Watermelon. 2n--22. Trop,
and subtropical Africa. Still found in the s e m i -
GUIZOTIA ABYSSINICA (L.f. ) Cass. Niger seed.
deserts of the Kalahari. Cultivated since ancient
2n=30. Sporadically from Ethiopia to Malawi.
times in the Mediterranean area and in India. Now
Spread to other countries where it was cultivated
cultivated in many countries of the Old and New
and may have run wild. Cultivated as an oil-seed
Worlds so that secondary centres of diversity have
crop in India and Africa.
arisen. Var. citroides (Bailey) Mansf. is found in
Sudan. Cultivated in USA (citron, preserving
GYNURA CERNUA Benth. 2n=20. W. Africa. A
melon) and USSR.
herb cultivated for its leaves.
The wild form, var. colocynthoides (syn. C.
colocynthoides Pang. ) is characterized by its white
LACTUA TARAXACIFOLIA (Willd. ) Schum.
flesh with bitter or non-bitter flavour while the
(syn. Sonchus taraxacifolius Willd. ). Wild lettuce,
cultivated form var. edulis (syn. C. edulis Pang. )
Langue de Vaches. 2n= . Trop. Africa e s -
has red or yellow flesh with sweet flavour (Shi-
pecially in Sierra Leone, Ghana, S. Nigeria and
motsuma, 1965).
Nile region. Cultivated in W. Africa as a vegetable.
The citron - a fodder melon - is adapted to
SENECIO BIAFRAE Oliv. 2n= . Africa. Occa- very dry regions. It is both a weedy and a culti-
sionally cultivated in W. Africa. vated plant.
SENECIO GABONICUS Oliv. 2n= . Trop. W. COCCINIA ABYSSINICA (W. 6 A. ) Cogn. Anchoté.
2n= . Ethiopia. Sporadic cultivation in SW.
Africa. Occasionally cultivated.
provinces for its tubers. The tubers of the wild
Crassulaceae plants a r e inedible, while the fruits a r e edible.
The fruits of the cultivated anchoté a r e however
BRYOPHYLLUM PINNATUM (Lam. ) Oken. Never not eaten.
die, Resurrection plant. 2n= . Africa. Culti-
vated there as a medicinal crop and elsewhere as CUCUMEROPSIS EDULIS (Hook. f.). Cogn. 2n=
an ornamental. W. Africa. Cultivated in gardens and on roofs.
Cruciferae CUCUMEROPSIS MANNn Naud. 2n= W. and

BRASSICA CARINATA A. Br. Abyssinian mustard. C. Africa. Cultivated there.
2n=34, genome formula BBCC. Unknown wild.
Cultivated in Ethiopia as a vegetable and as an oil CUCUMIS ANGURIA L. West Indian gherkin,
crop. A natural hybrid of B. nigra* and B. o l e r a - Gooseberry gourd. 2n=24. This cultigen (also
cea*. C. anguria var. anguria) derives from the non-
bitter wild species C. longipes*. This last species
BRASSICA JUNCEA (L. ) Czern. &Coss. (syn. was probably taken by slaves to the New World
Sinapsis juncea L. ). Sarepta-mustard, Brown m u s - where the cultigen was developed (Meeuse, 1958).
AFRICAN CENTRE 110
CUCUMIS DIPSACEUS Ehrb. Teasel Gourd. At present it occurs subspontaneously and is cul-
2n=24. Africa. An ornamental. tivated in tropical Africa, tropical Asia and t r o p i -
cal America. It has been shown that gourds float
CUCUMIS LONGIPES Hook,f. (syn. C. anguria L. for a long time while seeds remain viable. This
var. longlpes (Hook,f.) Meeuse). 2n=24. The may explain a very early spread to other continents
wild ancestor of the cultigen C. anguria* which by sea currents. Remains of plant material tenta-
has been developed in the West Indies (Meeuse, tively identified as belonging to Lagenaria have
1958). been found in the Spirit Cave, Thailand and have
been dated 10 000-6 000 BC. (Gorman, 1970).
CUCUMIS MELO L. Muskmelon, Melon, Cante- Remains of the bottle gourd were found in Mexico
loupe. 2n=24. As most Cucumis species come dated 7 000-5 500 BC. (Whitaker &Cutler, 1971)
from Africa this species probably originates from in P e r u 4 000-3 000 B C , in the Egyptian tombs
the tropical part of this continent. From there it dated 3 500-3 000 BC. (Purseglove, 1968) and in
spread to other regions producing secondary gene China 500 AD. (Li, 1969). Material found in
centres in Iran (p. 79), China (p. 30), Iran and Mexico and dated 700-1 300 AD. appears to be
S. USSR (p. 72) (Leppik, 1966). more closely related to the modern races puncta-
tum and latifolium than it is to other races
CUCUMIS METULIFERUS E. Mey. African horned (Whitaker &Cutler, 1971).
cucumber. 2n=24. Cultivated as an ornamental It must be the oldest crop cultivated in the
and in some parts of Africa for its fruits. tropics (Purseglove, 1968). Related species a r e :
L. abyssinica (Hook,f. ) C. Jeffrey (syn. Adeno-
LAGENARIA SICERARIA (Molina) Standi, (syn. pus abyssinicus Hook,f., A. reticulatus Gilg)
L. vulgaris Ser. ) Bottle gourd. White-flowered (2n= ), L. guineënsis (G. Don. ) C. Jeffrey
gourd, Calabash gourd. 2n=22. This species has (syn. Bryonia guineënsis G. Don. Adenopus longi-
most likely a tropical African origin as in this florus Benth., A. guineënsis (G. Don.) Exell,
continent wild plants of this species and closely A. pynaerti De Wild. ) (2n= ) and L. rufa
related species have been found. (Gilg) C. Jeffrey (syn. Adenopus rufus Gilg)
(2n= ) (Jeffrey, 1962).
TELFAIRIA OCCIDENTALIS Hook.f. Fluted

pumpkin. 2n= . Trop. Africa. Cultivated
there for its seeds.
Dioscorea cayenensis and D. rotundata ( ), D. dumetorum

(...) and D.bulbifera (—) (Harris, 1972)
Lagenaria siceraria
CUCURBITACEAE-GERANIACEAE 111
Dioscoreaceae DIOSCOREA ZARA Baudon. 2n= . Cultivated

DIOSCOREA ABYSSINICA Höchst. 2n=40. to some extent in C. Africa. This name is applied
Ethiopia. Savannah regions of Africa. Cultivated to what is possibly a form of either D. sagitti-
to a limited extent as a food crop, especially in folia Pax. or D. lecardii De Wild.
Uganda (Burkill, 1939; Coursey, 1967).
Euphorbiaceae
DIOSCOREA BULBIFERA L. (syn. D. latifolia BRIDELIA MICRANTHA (Höchst. ) Bâillon.
Benth. ). Potato yam, Aerial yam, Bulbil-bearing 2n= . Trop. Africa. Cultivated as food plant
yam. 2n=36, 40, 54, 60, 80, 100. Wild and culti- of the African silk caterpillar.
vated in trop. Asia (p. 31) and Africa. It might be
domesticated in both regions. EUPHORBIA DREGEANA E. Mey. 2n=
Namaqualand, Africa. Sometimes cultivated for
DIOSCOREA CAYENENSIS Lam. Yellow yam, rubber (Uphof, 1968).
Yellow Guinea yam, Twelve month yam, Cut-and-
come yam. 2n=c. 34, c. 54, c. 120. W. Africa. EUPHORBIA KAMERUNICA Pax. Solo. 2n=
Also cultivated in the West Indies and in W. W. Africa. A tall xerophytic t r e e . Cultivated for
Africa, the latex used for tattooing and to poison arrows
(Uphof, 1968).
DIOSCOREA COLOCASIIFOLIA Pax. False water
yam. 2n= . W. Africa. Cultivated in E. Ghana, MANIHOT ESCULENTA Crantz. Cassava. 2n=36.
Cameroon and Mayumbe area of Zaire. S. and C. America (p. 148). Secondary centre in
Africa.
DIOSCOREA DUMENTORUM (Kunth) Pax. Bitter
yam, Cluster yam. 2n=36, 40, 45, 54. Cultivated PLUKENETIA CONOPHORA Muell. Arg. (syn.
throughout Africa between 15°N and 15°S. It is Tetracarpidum conophorum Hutch. &Dalz. ).
closely related to the Asian D. hispida*. 2n= . A woody vine of trop. Africa. Cultivated
as a source of oil.
DIOSCOREA ELEPHANTIDES (L'Her. ) Engl.
Elephant's foot. 2n= . S. Africa, in the rocky, RICINODENDRON HEUDELOTII (Baill. ) P i e r r e
s e m i - d e s e r t s . Collected and eaten by Hottentots. ex Pax. 2n=22. W. Africa to Angola and Usam-
In Europe and N. America it is cultivated as a bara. Fruits and seeds are used as a source of
curiosity (Coursey, 1967). The tuber can grow oil. Also cultivated in Cameroon.
upto 350 kg.
RICINUS COMMUNIS L Castor bean, Castor oil
DIOSCOREA HIRTIFLORA Benth. 2n=40. Savannah plant. 2n=20. Trop. E. Africa and India. Culti-
regions of Africa. Cultivated in N. Nigeria. vated now in most tropical countries where it runs
wild in clearings, roadsides and dumpheaps. It
DIOSCOREA LIEBRECHTSIANA De Wild. 2n= probably originated as a dumpheap-camp-follower
Africa. Cultivated. growing in to an oil plant, a drug and an ornamen-
tal (Anderson, 1952). The only species of this
DIOSCOREA OVfNALA Baker. Ovinala. 2n= genus.
Madagascar. Cultivated there. Almost replaced
by D. alata* and Manihot utilissima* (Coursey, Flacourtiaceae
1967).
ONCOBA ECHINATA Oliver (syn. Caloncoba
echinata (Oliver) Gilg. ). 2n= . Trop. W.
DIOSCOREA PRAEHENSILIS Benth. Bush yam,
Africa. Cultivated in C. and S. America.
Forest yam. 2n=40, 80. W. Africa. Cultivated
there. Probably ancestor of D. rotundata*.
Geraniaceae
DIOSCOREA ROTUNDATA Poir. White yam, PELARGONIUM x ASPERUM Ehrh. ex Willd.
White Guinea yam, Guinea yam, Eboe yam. (syn. P. radula L'Her. var. roseum Willd.,
2n=40. W. Africa. The most important yam culti- Pelargonium roseum Willd. ). 2n=77, 81. S. Africa.
vated t h e r e . Probably derived from D. praehen- Cultivated for its geranium oil. The plant is male
silis*. There exists a great variability. Closely sterile. Autotetraploids (2n=4x=154) a r e fertile.
related to D. cayenensis* and is often included in Crossed with autotetraploid P. denticulatum and
this species. backcrossed with 4x P. x asperum resulted in
plants with 40-55% more oil than P roseum
DIOSCOREA SANSIBARIENSIS Pax. (syn. D. m a - (Tamai and Tokumasu, 1968). P. x asperum is
croura Harms, D. welwitschii Renole). Africa. probably a hybrid of P . radens x P. denticulatum*
Cultivated there. (Clifford, 1958) or P. graveolens* x P. radens
(Moore, 1955). P. radens Moore. (2n= ) is a
DIOSCOREA SEMPERFLORENS Illine. 2n= plant of S. Africa.
Cultivated in Congo. If the first parentage is correct than the
above new oil-rich hybrids a r e results of crossing
DIOSCOREA SOSO Jun. &P e r r . 2n= . Culti- a 4x hybrid (P. radens x P. denticulatum) with
vated formerly in Madagascar, but at present, it 4x P. denticulatum and backcrossing with the 4x
is replaced by Manihot utilissima*. original hybrid.
AFRICAN CENTRE 112
PELARGONIUM CRISPUM (L. ). L'Her. ex Ait. BRACHIARIA BRIZANTHA (Höchst.) Stapf. P a l i -

(syn. P . rigidum Willd. ). 2n= . S. Africa. sade g r a s s . 2n=54. Trop, and S. Africa, esp.
Cultivated for its lemon scented oil. It varies cultivated in Ceylon. A hexaploid g r a s s .
considerably in the wild and there a r e several
forms. BRACHIARIA DECUMBENS Stapf. Sinai g r a s s .
2n=36. Trop. Africa. Cultivated throughout the
PELARGONIUM DENTICULATUM Jacq. 2n=90. tropics for cattle fodder.
S. Africa. Was cultivated as a fragrant pelargo-
nium in Japan where it was replaced in 1954 by BRACHIARIA DEFLEXA C E . Hubbard. 2n=
P . roseum*. It has a pine scent. Ivory Coast to the Red Sea and Yemen and to
Angola, Mozambique, Transvaal and Madagascar.
PELARGONIUM GRAVEOLENS L'Her. (syn. P. Cultivated in Fouta Djalan (Guinea). It often is a
terebinthinaceum (Cav. ) Small). Rose-geranium. companion weed of other c e r e a l s . The cultivated
2n=90 S Africa. A pelargonium with r o s e - s c e n - type var. sativa Port, looks similar to B. r a m o -
ted leaves. Cultivated for its oil. There a r e sa Stapf (Portêres, 1951).
many cultivars. P . capitatum Willd., is a d e r i -
vative of P . graveolens (Moore, 1955). Cultivated BRACHIARIA MUTICA (Forsk. ) Stapf (syn. P a n i -
in Algeria and Isle of Réunion. cum barbinode Trin. ). P a r a g r a s s , Buffalo g r a s s ,
Mauritius g r a s s , Watergrass. 2n=36. Trop. Africa
PELARGONIUM KAROOENSE Kunth. 2n= and S. America. A tetraploid g r a s s . Cultivated
S. Africa. Cultivated for its geranol. as a fodder crop.
PELARGONIUM ODORATISSIMUM (L. ). Ait. BRACHIARIA RUZIZIENSIS Germain &C. Evrard.

2n=16. Trop. Africa. Extensively cultivated for Ruzigrass. 2n= . E. Africa. In Australia
its apple-scented geranium oil. Kennedy ruzigrass is cultivated.
PELARGONIUM TOMENTOSUM Jacq. 2n= CHLORIS GAYANA Kunth. Rhodes g r a s s . 2n=20,

S. Africa. Cultivated for its peppermint-scented (30), 40. Trop, and S. Africa. Cultivated in the
oil. It c r o s s e s readily with P . graveolens*. tropics for forage.
Gramineae CYNODON INCOMPLETUS Nées. var. hirsutus

ACROCERAS AMPLECTANS Stapp. (syn. P a n i - (Stent) De Wet &Harlan (syn. C. bradleyi Stent.
cum zizanoides Hbk. var. angustatum Stapf). 2n=18. S. Africa. Widely cultivated lawn-grass.
2n= . W. Africa. Cultivated in Gambia as a
vegetable (Terra, 1967). CYNODON x MAGENISSII Hurcombe. Sunturf
g r a s s . 2n=27. S. Africa. It is a triploid single
ACROCERAS MACRUM Stapf. 2n=36. S. Africa. clone derived from a putative, natural hybrid of
Cultivated as a pasture g r a s s . a tetraploid and a diploid species, which a r e not
known.
ANDROPOGON GAYANUS Kunth. 2n=20, 40. N.
Nigeria. It has been divided into var. gayanus CYNODON PLECTOSTACHYUS Pilger. Star g r a s s .
(syn. var. genuinus Hack. ), var. squamulatus 2n=18, 54. Trop. Africa. Cultivated for grazing
(Höchst. ) Stapf, var. bisquamulatus (Höchst.) and hay in the tropics. A cultivar of S. Nigeria is
Hack., and var. tridentatus (Höchst. ) Hack. The derived from a cross of this species x C. coursii
second and third varieties have been used for A. Camus.
selection. It is possible that the tetraploid plants
found in var. tridentatus a r e hybrids of A. gaya- CYNODON TRANSVAALENSIS Burtt-Davy. Afri-
nus (2n=20) in the far north of Nigeria and A. t e c - can Bermuda g r a s s . 2n=18. S. Africa. Used as
torum (2n=20) in the S. of N. Nigeria. turf g r a s s . It appears to share the same genome
with C. aethiopicus Clayton &Harlan (2n=18, 36)
ARUNDINARIA ALPINA K. Schum. 2n= and C. nlemfuensis Vanderyst var. nlemfuensis
Kenya, Tanzania, Uganda, Sudan, Ethiopia, (2n=18, 36). The first species occurs in E. of
Rwanda and Urundi in mountain forests at an a l t i - Africa and the second in Angola and E. Africa.
tude of about 2 400-3 000 m. Its stems are used Although C. aethiopicus occurs sympatric with
in paper industry and as building material. the other two species no introgression exists due
to strong crossing b a r r i e r s (Chedda &Rawal,
AVENA STRIGOSA Schreb. abyssinica type. 1971).
Abyssinian oat. 2n=28, genome formula AsAsBB.
This type has also been described as A. abyssini- DIGITARIA ABYSSINICA (Höchst. ) Stapf. Abyssi-
ca Höchst. This is the non-brittle form while the nian finger g r a s s . 2n= . Trop. Africa. Used in
semi-brittle form (vaviloviana type, A. vavilo- S. Africa to control erosion.
viana Malz. ) is also found in Ethiopia The abyssi-
nian type is harvested and threshed together with DIGITARIA DECUMBENS Stent. Digit g r a s s .
barley. Both types probably derive from intro- 2n=27, (37). S. Africa. In 1935, material of this
duced barbata type of A. strigosa* (Ladizinsky & species was collected in E. Transvaal and was
Zohary, 1971). introduced to the USA PI 111110 was vegetably mul-
tiplied to give a clone widely grown in the tropics.
GERANIACEAE - GRAMINEAE 113
This clone was named Pangola g r a s s .
DIGITARIA EX3LIS Stapf. Fonto, Fonia, Fundi,

Hungary rice. 2n=18, 36, 40. W. Africa. Un-
known wild. Its first cultivation may have started
somewhere in C. Africa close to the Sahara about
5 000 BC. It may derive from the wild D. b a r b i -
nodis Henr.
X 1
L-I I
i v. y
i
' l
l/^^s.
i
,,,,_-' ,-
r
~"'^"*~f~-\ ^ \ ^
1 vr ' \
Digitaria exilis ( )andD. iburua ( ) (Portêres, 1950)
DIGITARIA IBURUA Stapf. Iburu, Aburo. 2n= Eleusine africana (Phillips, 1972)
Africa. Cultivated in N. Nigeria, adjacent Niger
Republic, and in Togoland. It may derive from the
wild D. barbinodis Henr.
DIGITARIA PENTZII Stent. Taiwan pangola g r a s s .

2n=(18, 27), 36, (45), 54. S. Africa. Related to
D. decumbens*. It can be used as a source of
virus disease resistance.
DIGITARIA TRICOSTULATA (Hack. ) Henr.

2n= . Africa. Related to D. iburua*.
DIGITARIA VALIDA Stent. Giant pangola g r a s s .

2n=24, 30, 36. S. Africa. A source of disease
resistant for D. decumbens*.
EHRHARTA CALYCINA Smith. Veldtgrass.

2n=24, 30, 48. S. Africa. Used as a soil stabilizer
in W. of USA. Cv. California veldtgrass has an
open panicle and sheds its caryopses. The new cv.
Mission veldtgrass has a compact panicle and is
non-shedding.
ELEUSINE AFRICANA Kennedy-O'Bryan. 2n=36.

This species was formerly included in E. indica*.
It probably originated as an amphiploid of E. indi-
ca and a closely related taxon, or as an autotetra-
ploid of E. indica. It is suggested that E. c o r a -
cana* African highland type derives from E. afri- Eleusine coracana (Portêres, 1950)
cana.
ELEUSINE CORACANA (L.) Gaertn. Finger millet, one of the parents of E. africana. The Afro-Asiatic
Ragi. 2n=36. Mehra (1963) recognized two types: form derives from the African highland form. It
1. African highland type and 2. Afro-Asiatic type. has some characteristics of E. indica. It is the
The first has longer lemmas, glumes and spike- form that was taken to India (Mehra, 1963)
lets and the spikelets a r e enclosed, while the s e -
cond has naked grains. The African highland type ELEUSINE INDICA (L.) Gaertn. 2n=18. Africa.
probably derives from E. africana*. Wild in Africa. Cultivated on small scale in Ethiopia for fibre
Probably an allotetraploid. Formerly included as production. Wild or weedy types were introduced
the tetraploid form of E. indica*, which may be into India. It grows wild there and as companion
weed of E. coracana*. Probably one of the parents
AFRICAN CENTRE 114
of E. africana*, the putative parent of E. c o r a - MELINIS MINUTIFLORA Beauv. Molasses grass

cana. Brazilian stink g r a s s , Honey g r a s s . 2n=36.
Africa. Cultivated as a fodder plant.
OREOBAMBOS BUCHWALDII K. Shum. 2n=

The mountains of trop. Africa.
ORYZA BRACHYANTHA A. Chev. &Roehr.

2n=24, genome formula F F . W. and C. Africa.
ORYZA BREVILIGULATA vide O. perennis*
ORYZA EICHINGERI Peter. 2n=24, genome for-

mula CC, 48, genome formula BBCC. Africa.
It has the same genomes as the Asian O. minuta*.
It belongs to the 'officinalis' group. As the name
O. eichingeri was also used for a tetraploid
species, now included in O. punctata*. Gopala-
krishnan and Sampath (1966) suggested to rename
this species as O. schweinfurthiana.
ORYZA GLABERRIMA Steud. African rice.

2n=24, genome formula AÂ^. Senegal to Ubangui-
Chari. P r i m a r y centre: Central Niger delta.
Secondary centres: Senegambia and Guinea. It is
generally believed that the cultivated African rice
developed from the wild O. longistaminata A.
Chev. &Roehr. This wild rice is still collected
as food. It is thought to derive from the African
race of O. perennis*.
Eleuame indica (Phillips, 1972)
ERAGROSTIB CURVULA (Schrad. ) Nees. Weeping

loving grass, Boer love g r a s s . 2n=20, 40, 50, 60,
30-70. S. Africa. Used to stabilize sand.
ERAGROSTB SUPERBA Peyr. 2n=40. E. Africa.

Cultivated there.
ERAGROSTIS TEF (Zuccagni) Trotter (syn. E.

abessinica Link, Poa abyssinica Jacq. ). Tef.
2n=40. Ethiopia and E r i t r e a . Cultivated there as
a cereal and elsewhere for forage crop.
Oryza glaberrima (PorteTes, 1950)
However Nayar (1973) showed that O. glaber-

rima derives from an old introduction of rice,
O. sativa*. O. sativa has reached Egypt from the
Indian subcontinent in 4th Century BC. It is not
known whether rice spread to West Africa, but
during the Arab conquest of N. Africa in the 7th
Century rice was distributed. At that time it may
have been taken to the Sudan zone. Depending on
the introduced material, isolation and selection
O. glaberrima evoluated.
Eragrostis tef (Porteras, 1950) O. barthii A. Chev., 2n=24 is according to
Nayar (1973) a hybrid product of Asiatic O. sativa*
introduced in the 16th Century and later. O. s t a p -
HYPARRHENIA RUFA (Nees) Stapf, (syn. Andro- fii Roshev. is included in O. barthii.
pogon rufus Kunth). Yaragua g r a s s . 2n=20, 30,
36, 40. Trop. Africa. Cultivated in N. and S. ORYZA LONGISTAMINATA, see O. glaberrima
America. and O. perennis.
v
L - ' *
L^ L 1 f'
Oryza perennis African race (Portéres, 1950)
ORYZA PUNCTATA Kotschychev ex Steud. 2n=24,

genome formula CC, 48, genome formula BBCC.
Africa. The tetraploid has also been described
O. eichingeri but this name is used for a different
species.
OXYTENANTHERA ABYSSINICA Munro (syn.

Bambusa abyssinica Rich. ). Woody bamboograss.
2n=c. 60. Trop. Africa. Cultivated for its stems
which a r e used for boats and rafts, as well as for
Oryza longistaminata (Harlan, 1973) paper making.
PANICUM COLORATUM L. Small buffalo g r a s s .

2n=18, 36 (54).. S. Africa. A good fodder g r a s s .
Closely allied to P. maximum*.
PANICUM MAXIMUM Jacq. Guinea grass, Panic.

2n=18, (32), 36, (48). Trop, and S. Africa, Mada-
gascar, Mascarene Islands and Yemen. Cultivated
as a pasture grass in many warm countries.
PANICUM SUMATRENSE Roth ex Roem. &Schult,

(syn. P. miliare Lamk. ). Little millet. India and
Ceylon. P. psilopodium Trin. (2n=54) might be its
ancestor species (Mansfeld, 1959). It is possible
that 36 chromosome cytotypes occur.
PASPALUM SCROBICULATUM*
PENNISETUM ANCHYLOCHAETE Stapf &Hubbard.

2n= . Africa. Cultivated as a cereal in N.
Nigeria (Mansfeld, 1959).
PENNISETUM CLANDESTINUM Höchst, ex Chiov.

Kikuyu g r a s s . 2n=36. Trop. Africa. A perennial
g r a s s cultivated for pasture and hay.
PENNISETUM ECHINURUS (K. Schum. ) Stapf &

Hubbard. 2n=14. Africa. Cultivated in Angola and
Oryza barthii (Harlan, 1973) E. Africa.
PENNISETUM GIBBOSUM Stapf & Hubbard.

ORYZA PERENNIS Moench. 2n=24. genome for- 2n= . Africa. Cultivated in N. Nigeria.
mula AA. The distribution of O. perennis is d i s -
cussed on p. 65. The African race includes O. PENNISETUM LEONIS Stapf &Hubbard. 2n=14.
longistaminata A. Chev. & Roehr (syn. O. b r e v i - Africa. Cultivated in Sierra Leone.
ligulataA. Chev. &Roehr). It does not hybridize
with O. glaberrima* and O. sativa*. Grains a r e PENNISETUM MAIWA Stapf &Hubbard. 2n=
collected for food. Africa. Cultivated in N. Nigeria. It has a small
head.
AFRICAN CENTRE 116
PENNISETUM MALACOCHAETE Stapf &Hubbard. Arabia, India and elsewhere. Nigerian plants a r e
2n= . Africa. Cultivated in Tanzania. characterized by yellow endosperm. Hybrids b e -
tween cultivated forms, with wild plants and with
PENNISETUM NIGRITARUM (Schlechtend. ) P. purpureum* result in gene exchange and a high
Durand &Schinz. 2n= . Africa. Cultivated as variability especially in Ethiopia and Sudan.
a cereal in W. Africa. Bono (1973) included all cultivated Pennisetum
species in one species. This species is divided on
PENNISETUM NILOTICUM Stapf & Hubbard. morphological grounds into two groups and four
2n= . Africa. Cultivated in Egypt. subgroups: cultivated from l a Mauritania, Mali
and Ivory Coast, l b . Haute-Volta, 2a. Niger and
PENNISETUM PERSPECIOSUM Stapf & Hubbard. 2b. Senegal. The cultivars from other countries
2n= . Africa. Cultivated in Sudan. in W. Africa were not included in the study.
PENNISETUM PURPUREUM Schum. Napier grass. PENNISETUM UNISETUM (Nees) Benth. Natal
Elephant g r a s s . 2n=28, genome formula A A B B . grass, Drakenberg silky g r a s s . 2n= . N. and
From Senegambia to the Red Sea. A perennial E. Africa. Cultivated there as a forage g r a s s .
pasture g r a s s . Hybrids between cultivated and wild
types and with P. typhoides* often occur resulting PENNISETUM VULPINUM Stapf & Hubbard.
in gene exchange and a high variability. 2n= . Africa. Cultivated in Sudan.
PENNISETUM PYCNOSTACHYUM (Steudel) Stapf SACCHARUM SPONTANEUM L. Wild sugar cane.
&Hubbard. 2n=14. Africa. Cultivated as a cereal 2n=104-128. From India (p. 66) plants with 2n=54
inW. Africa (Mansfeld, 1959). It has brittle spread to Africa. In Uganda and adjacent Tanzania
spines which deter birds. it is actually cultivated. In Africa originally used
as a source of salt by burning, later it became
PENNISETUM ROBUSTUM Stapf & Hubbard. used as hedges for erosion control and for house-
2n= . Africa. Cultivated there. hold. Grassl (1964) suggested that the high num-
ber of chromosomes may derive from hybridization
PENNISETUM SPICATUM (L. )Roem. & Schult. of the original S. spontaneum and an African r e -
P e a r l millet, African millet. 2n=14. Africa. Cul- lated species e.g. Sorghum bicolor*.
tivated as a cereal in C. Africa and elsewhere.
SECALE AFRICANUM Stapf. 2n=14. Eastern part
of the Karoo plateau, S. Africa. Whether this
species is a Pleistocene immigrant to S. Africa
or a derivative of a relatively recent introduction
of seeds of S. montanum from Spain or Italy as a
contaminant of wheat and barley is not known
(Khush, 1960). It has the same chromosome
arrangement as S. montanum* and must have d e -
rived from this species (Stutz, 1972). Owing to
its separation from the Secale-area it adopted
self-fertilization as a means of perpetuation. It
has a fragile rachis and a perennial habit.
SETARIA GLAUCA Beauv. (syn. Pennisetum

glaucum (L. ) R. Br. ). P e a r l millet, Cat-tail millet.
2n=(18), 36, (72). Africa. Cultivated in warm
countries as cattle food.
SETARIA SPHACELATA (Schum. ) Stapf & Hubbard.

Golden timothy g r a s s . 2n=18, 36, 54. Africa.
Cultivated in Africa, Australia and elsewhere.
Some cultivars have a high oxalic acid content
which may result in the death of cattle.
SORGHUM BICOLOR (L. ) Moench. Sorghum.

2n=20. The genus Sorghum is widespread in
Africa, the Americas, Asia, Mediterranean
Europe and Australia. All cultivated sorghums
P e n n i s e t u m spicatum ( ) and P . typhoides ( ) (Portêres,
1950)
belong to the section Sorghum (formerly E u -
sorghum) which is restricted to the Old World.
It is thought that cultivation of sorghum s t a r -
PENNISETUM TYPHOIDES (Burm.f. ) L . C . Rich, ted in one place and spread with the weed s o r g -
(syn. P. spicatum L. ). Bulrush millet, Pearl hums to the other areas poggett, 1970). It is still
millet. 2n=14, genome formula AA. Probably possible that cultivated sorghum spread from one
Africa. Cultivated there for its grain. Spread to centre throughout Africa and in each new site in-
trogressed with the local wild Eusorghum(s) or
are derived from S. arundinaceum, the Kaffir

corn from S. verticilliflorum (Steud. ) Stapf which
occurs from Eritrea to SE. Africa and cultivated
sorghums of Eritrea and Ethiopia from S. aethio-
picum (Hack. )Rupr. ex Stapf and so each has its
centre of domestication. Two sorghum types are
typical for China and the Far East (p. 33).
S. bicolor groups many former species, but as
the cultivated sorghums hybridize easily it is not
correct to put all the different types on a species
level. The same holds good for the wild Eu-
sorghums which a r e interfertile with each other
and with the cultivated ones. They a r e grouped in
S. arundinaceum (Desv. ) Stapf. Dogget (1965)
suggested that man selected sorghum for his spe-
cial needs. This resulted in different types: arun-
dinaceum type in W. Africa, aethiopicum type in
Sudan and verticilliflorum type in E. Africa. As
the cultivated sorghums easily cross with wild and
weedy types there is an interchange of genes b e -
tween them. This results in a big variation of this
species.
A simplified classification of sorghum was
presented by Harlan and de Wet (1972).
TRICHOLAENA ROSEA Nees (syn. Rhynchelytrum

repens (Willd. ) Hubbard. Natal grass, Wire grass,
Ruby g r a s s . 2n= . Trop, and SE. Africa. Cul-
tivated as a forage grass and as an ornamental.
TRITICUM AESTIVUM (L. ) Thell. ssp. vulgare

Sorghum bicolor
(Vill. ) MK. (syn. T. vulgare Vill. ). Bread wheat,
Common wheat. 2n=42, genome formula AABBDD.
The bread wheats in Africa are especially cultiva-
ted in the Sahara oases and Sudan zone. In the
oases special types developed, sometimes called
T. vulgare ssp. oasicolum Due. Other speltoid
types have been referred to as T. spelta var.
saharense, while also inflatum and sphaerococcoid
forms (T. sphaerococcum) have been found. P e r -
haps the Sahara oases could be a secondary centre
of ssp. vulgare.
Spelt wheats may have been introduced by the
Romans. It is remarkable that an African spelt
c a r r i e s an Rf tim. gene like some of the European
spelts (Zeven, 1971).
TRITICUM TURGIDUM (L. ) Thell. Cultivated

tetraploid wheats. 2n=28, genome formula AABB.
Tetraploid cultivated wheats were introduced into
Ethiopia. A secondary centre of T. turgidum*
ssp dicoccum, ssp. turgidum conv. turgidum
conv. durum and conv. polonicum arose h e r e .
Typical Ethiopian types have been classified as
abyssinicum.
Leppik (1968) showed that Ethiopia is also
the primary centre of wheat stem rust (Puccinia
graminis P e r s . ).
Sorghum bicolor (de Wet et a l . , 1972) ZEA MAYS L. Maize. 2n=20. Secondary centres
in W. Africa, S. Africa, E. Africa (Somalia,
Kenya and Ethiopia) (Brandolini, 1970). Domesti-
that domestication of sorghum started indepen- cated in C. America (p. 166).
dently in several places as is believed by P o r - An example of the diversity of maize is given
t e r a s (1962) and de Wet and Huckabay (1967). They by P l a r r e (1972) for the Karamoja district in
suggested that cultivated sorghums in West Africa Uganda.
AFRICAN CENTRE 118
Labiatae nummularifolius (A. nummularifolius DC. ) deve-

ACOLANTHUS PUBESCENS Benth. 2n=36. Trop. loped in the West Indies. It has a low spreading
Africa. Cultivated as a salad. habit with buds located in the root crown. This
makes it an ideal pasture plant (Whyte et a l . ,
COLEUS DAZO Chev. (syn. C. floribundus var. 1953).
longiper Robyns. Plectranthus esculentus N . E .
Br. ). Dazo, Kaffir potato. 2n=24. Africa. Culti- ARACHIS HYPOGAEA L. Groundnut, Peanut.
vated there. 2n=40. S. America (p. 151). Introduced into Africa
where it is cultivated widely. Bunting (1955, 1958)
COLEUS EDULIS Vatke (syn. C. tuberosus R i - has described numerous varieties for trop. Africa,
chard. ). 2n= . E. Africa. Cultivated in Ethio- which points to a secondary centre in this region.
pia (Cufodontis, 1957).
ASPALATHUS CONTAMINATUS (Thunb.) Druce.
COLEUS FORSKOHLII (Poir. ) Briquet, (syn. C. Rooibostee, Rooibos tea bush. 2n= . S. Africa.
barbatus Benth., Ocimum asperum Roth. ) Kaffir There a r e two forms: the prostate one is found on
potato. 2n=28. E. Africa and India. Cultivated In the Cape Penisula and the erect cultivated form
some parts of India (Uphof, 1968). (A. cedarbergensis Bolus) is observed in the
Cedarbergs in the Clanwilliam and Citrusdal
COLEUS LANGOUASSENSB Chev. 2n- regions. The latter is the cultivated type (Cheney
&Scholts, 1963).
Africa. Cultivated there.
COLEUS ROTUNDIFOLIUS Chev. &P e r r o t . (syn. ASTRAGALUS VENOSUS Höchst. 2n=16. E. Africa.
Plectranthus rotundifolius Spreng. ) Hausa potato. Cultivated for horse fodder.
2n= . Trop. Africa and Asia. Cultivated for
its tubers. BAPHIA NITIDA Lodd. Cam wood. 2n=44. W.
Africa. Cultivated formerly as a source of red
HYPTIS SPICIGERA Lam. 2n=32. Trop. Africa. dye. Now it is only cultivated as an ornamental.
Cultivated there for its oily seeds.
CAJANUS CAJAN (L. ) Mill Sp. (syn. C. indicus
OCIMUM KILIMANDSCHARICUM Guerke. 2n=76. Spreng. ) Pigeon pea. 2n=22, 44, 66. Probably
Africa. Cultivated during World War II for the Africa. Wild and naturalized plants have been
preparation of camphor. found there. Spread to India where a secondary
centre arose (p. 67).
ORTHOSIPHON RUBICUNDUS Benth. (syn. P l e c -
tranthus tuberosus Roxb. ). 2n=28. Trop. Africa.
Cultivated for its tubers.
SOLENOSTEMON OCYMOIDES Schum. &Thonn.

2n= . Trop. Africa. Cultivated as a vegetable.
Leguminosae
ACACIA KARROO Hayne. (syn. A. horrida Willd.)
Mimosa thorn, Allthorn acacia, Sweet thorn.
2n=52, 104. S. Africa. This t r e e is planted as an
ornamental, in hedges and as sandbinder.
ACACIA NILOTICA (L. )Willd. ex Del. (syn. A.

arabica (Lam. ) Willd. Babul acacia. 2n=52, 104.
Trop. Africa extending to India. Cultivated there
for its bark tannin. It also yields babul gum
(Purseglove, 1968).
ACACIA SENEGAL (L.)Willd. Sudan gum arabic.

2n=26. Trop. Africa extending to the Red Sea and
NW. India. The gum is mainly obtained from wild
and semi-cultivated t r e e s in the Sudan (Purse-
glove, 1968).
ALYSICARPUS RUGOSUS (Willd. ) DC. (syn. A.

violaceus (Forsk. ) Schindler). 2n=16. Trop. Africa.
Used as a cover crop and fodder crop.
ALYSICARPUS VAGINALIS DC. Alyce clover, Cajanus cajan

One-leaved clover. 2n=16, 20. Trop. Africa.
Cultivated now in all tropics for soil improvement,
as a cover crop and as a fodder crop. Var.
LABIATAE - LEGUMINOSAE 119
CANAVALIA REGALIS Dunn. 2n=22. Probably an are derived from the African stock.
old African domesticant known only in cultivation.
It is probably derived from C. virosa (Roxb.) INDIGOFERA ARRECTA Höchst. Natal indigo.
Wight & A m . (2n=22) which occurs in Africa south 2n=16. E. Africa. Cultivated formerly as a dye
of the Sahara and also in India (Sauer, 1964). crop, and at present as a green manure.
CASSIA ANGUSTIFOLIA Vahl. Indian senna, T e n e - INDIGOFERA ENDECAPHYLLA Jacq. 2n=32, 36.
velly senna. 2n=(26), 28. Somaliland and Arabia. Africa and Asia. Cultivated as a fodder crop
Cultivated in India for senna (laxative) (Purse- usually in pastures.
glove, 1968).
INDIGOFERA TINCTORIA L. (syn. I. indica Lam.,
CASSIA SENNA L. (syn. C. acutifolia Del). I. sumatrana Gaertn. ). True indigo plant. 2n=16.
Alexandrian senna, 2n= . Egypt, Sudan region Probably W. Africa (Mansfeld, 1959). Cultivated
and Sahara. Leaves and pods a r e taken from wild as a dye plant. Also used as a green manure.
and cultivated plants (Purseglove, 1968).
KERSTINGIELLA GEOCARPA Harms. Geocarpa
CROTALARIA CANNABINA Schweinf. 2n= bean, Geocarpa groundnut, Kersting's groundnut.
Sudan. A fibre crop. 2n=20, 22. Wild (var. tisserantii (Pellegrin)
Hepper in Cameroons and possibly in adjacent
CROTALARIA GOREENSIS Guill. &Pur. Gambia regions. Cultivated in W. Africa. The chromosome
pea. 2n=16, (32). Trop. Africa. Cultivated in number was established from this wild material.
Queensland, Australia for green manure. The cultivated types, var. geocarpa Hepper had
2n=22.
CROTALARIA INTERMEDIA Kotschy. 2n=16.
Trop. Africa. Cultivated in N. America and e l s e -
where especially for soil improvement, and also 1 v^
for grazing, hay and silage. L.1 s' Y^v^ y
i S s
i
CROTALARIA SPECTABILIS Roth. (syn. C. retzii
i
A. Hitchc. 2n=16. Trop. Africa. A green manure .1
cultivated in (sub)tropical countries. Wild in India

and elsewhere.
3
Yr.- _\ L
_r >-~4 (
CROTALARIA USARAMOENSIS Baker f. (syn.
/A
/ V S \
C. zanzibarica Benth. ). 2n=(14), 16, (20). x
E. Africa. A cover crop and green manure. •v -—'
a
\ y' ' \
CYAMOPSIS SENEGALENSIS Guill. & P e r r . Kerstingiella geocarpa (Portéres, 1950)
2n=14. The semi-arid savannah zone south of the
Sahara from Senegal to Saudi Arabia. An annual
herb. Probably the parental species of C. tetrago- LABLAB PURPUREUS (L.) Sweet (syn. Dolichos
noloba* (Hymowitz, 1973). lablab L., D. purpureus L., Lablab niger Medik. ).
Hyacinth bean, Bonavit bean, Lablab bean, Seins
CYAMOPSIS TETRAGONOLOBA (L. ) Taub. (syn. bean, Indian bean, Lubia bean, Egyptian bean.
C. psoralioides DC. ). Cluster bean, Guar. 2n=14. 2n=22, 24. Wild type (ssp. uncinatus V e r d e ,
Purseglove (1968) suggested that its origin lies in syn. Lablab uncinatus A. Rich. ) in trop. Africa
India, but Anderson (1960) stated an African do- from W. Africa to Sudan Republic and Ethiopia
mestication of this crop (see further p. 67). Cul- and to S. Africa. The commonly cultivated forms
tivated in S. India (p. 67). belong in general to ssp. purpureus unless they
have linear kidney bean-like pods. Var. purpureus
DESMODIUM SALICIFOLIUM (Poir. ex Lam. ) DC. of this subspecies is a distinct due to all parts of
2n=20, 22. Trop. Africa. Used as green manure. the plant being purple coloured. From Kenya the
cultivated ssp. bengalensis (Jacq. )Verde, (syn.
DIPOGON LIGNOSUS (L. ) Verde, (syn. Dolichos Dolichos bengalensis Jacq. ) is reported (Verd-
lignosus L . , D. benthamii Meisn., D. gibbosum court, 1970).
Thunb., Verdcourtia lignosus (L. ) Wilczek).
2n=22. C. Africa. Cultivated in Africa, S. America LOTONONIS BAINESII Baker. Miles lotononis.
and Australia where it has run wild (Verdcourt, 2n=36. N. Transvaal, S. Africa and Rhodesia.
1970). A perennial cultivated in Australia.
ERYTHRINA SENEGALENSIS DC. Coral flower. PHYSOSTIGMA VENENOSUM Balf. Calabar bean,
2n=42. W. Africa. Used as a hedge plant, as an Ordeal bean. 2n= . W. Africa. Cultivated for
ornamental and for medicinal purposes. its beans which a r e used as an ordeal poison.
GLYCINE WIGHTII Verde. Rhodesian kudzu vine. PISUM SATIVUM ssp. abyssinicum (A. Braun. )
2n=22, 44. Africa. A perennial soya bean. Some Alef. (syn. P. abyssinicum Braun). 2n=14.
forms a r e also found in SE. Asia. The cultivars Ethiopia and Yemen. Rarely found wild.
AFRICAN CENTRE 120
/ I -' N. S
/ ,-J .-'^"^_'^
i \-'^ ) " ^ i J
l---^ 1 ,' J
-'
/'V--~^
,.,_r--^ ;e..
\
^"""-^
Pisum sativum spp. abyssinicum (Govorov, 1937) Vigna sinensis (Harlan, 1973)
PSOPHOCARPUS PALUSTRIS Desv. (syn. P. VOANDZEIA SUBTERRANEA Thouars. Bambara

longepedunculatus Hassk. )• 2n=20, 22. Probably groundnut, Congo eoober. 2n=22. Wild in W.
trop. Africa. Cultivated for its fruits. A perenni- Africa. Distributed throughout Africa and later to
al herb. America and Asia. Hepper (1963) described the
wild type as var. spontanea (Harms) Hepper and
SPHENOSTYLIS SCHWEINFURTHII Harms. Yam the cultivated ones as var. subterranea Hepper.
bean. 2n= . Trop. Africa. A woody plant.
Cultivated for its seeds and tubers. Linaceae
SPHENOSTYLIS STENOCARPA Harms. LINUM USITATISSIMUM L. Flax. 2n=30, (32).

2n=18. For possible origin see p. 87. Ssp. indo-abyssi-
Trop. Africa. Cultivated for its seeds.
nicum Vav. &Ell. is cultivated in Ethiopia and
STYLOSANTHUS FRUTICOSA (Retz. )Alston (syn. Eritrea. Identical types a r e cultivated in India and
S. mucronata Willd. ). Wild lucerne. 2n= may formerly have been introduced there from
Trop. Africa and SE. Asia. Cultivated as a fodder Africa.
crop in Brazil and Australia.
Lythraceae
STYLOSANTHUS HUMILIS H.B. et K. Townsville LAWSONIA ALBA Lam. (syn. L. inermis L. ).
lucerne, Townsville style. 2n=20. S. Africa. Cul- Henna. Camphire. 2n= . A shrub from trop.
tivated in N. Australia in pastures. Asia and Africa with several u s e s .
TAMARINDUS INDICA L. Tamarind. 2n=24. The Malvaceae
savannas of trop. Africa. Introduced to India long GOSSYPRJM ANOMALUM Wawr. & Peyr. 2n=26,
ago and recently to other parts of the tropics. genome formula B . B j . Along the southern fringe
The t r e e and its parts have many uses (Purse- of the Sahara, in SW. Africa and Angola. It can
glove, 1968). be crossed with G. herbaceum* and G. arboreum*.
TEPHROSIA DENSIFLORA Hook,f. 2n= . W. GOSSYPIUM ARBOREUM L. T r e e cotton. 2n=26,
Africa. Cultivated and used to stupefy fish. genome formula A 2 A 2 - Race soudanense. NE. and
Closely related to T. vogelii*. W. Africa. Hutchinson (1962) suggested that it
was introduced from India, but Chowhury and
TEPHROSIA VOGELII Hook,f. Vogel tephrosia. Buth (1970, 1971) thought that it is native to Afri-
2n=22. Trop. Africa. Cultivated and used to ca (p. 121, p. 153). They found material dated
stupefy fish. Cultivated as a green manure and about 3 000 BC at an early Neolithic site. It was
cover crop. probably used as stock feed, because cotton hairs
intermediate between lint hairs and the hairs of
VIGNA UNGUICULATA (L. )Walp. (syn. V. sinen- wild species were found in goat dung. Probably
sis (L. ) Savi). Cowpea, Black eye, Southern pea. the form cultivated by the people of Meroë, an
2n=22, 24. P r i m a r y centre W. and C. Africa. ancient Nubian kingdom, dated about 500 BC.
Probably domesticated in W. Africa. Introduced
into the Indian subcontinent where ssp. sesquipe- GOSSYPIUM BARBADENSE L. Egyptian cotton.
dalis (L.) Verde, (syn. V. sesquipedalis (L.) 2n=52, genome formula (AADD) . Peru (p. 153).
Fruw., Dolichos sesquipedalis L. ), Asparagus Spread to Africa in historical time. A perennial
pea, Yardlong pea (2n=22, 24), and ssp. cylindrica type from S. Nigeria made a 'Green Revolution'
(L. ) Van Eseltine (syn. V. cylindrica Skeels., of cotton growing in the Nile delta possible after
V. catjang (Burm.f. )Walp.), catjang (2n=22), 1820. This lead to introduction of other cottons.
were developed. Ssp. sesquipedalis has a long Only Sea Islands was successful. Vigorous and
flabby pod, and is cultivated as a snap bean while fertile hybrids of these two types occurred from
ssp. cylindrica is developed as a forage crop. It which Egyptian was developed. It combines the
has small seeds (Paris, 1965) (p. 87). annual habit and some of the quality of Sea Islands
and some of the vigour and cropping characteristic
of the perennial. It can be grown twice a year.
LEGUMINOSAE - MALVACEAE 121
GOSSYPIUM HERBACEUM L.Short staple cotton. BC have been identified as G. arboreum*.

2n=26, genome formula A A 1 . Wild G. herbaceum Hutchinson (1962) suggested that cotton must have
var. africanum (Watt) Hutch. & Chose is a perenni- been brought from the present Rhodesian a r e a by
al shrub found in the bushveld across a belt from early t r a d e r s to the north. It is most likely that
Mozambique to Angola and SW. Africa. the earliest domesticants were selected between
Hutchinson (1971) suggested a likely centre of this a r e a and Ethiopia or Arabia. From this
domestication in southern Arabia and Baluchistan. variety the primitive cultivated race acerifolium
Furthermore, as wild G. herbaceum plants were which was formerly found in Ethiopia, S. Arabia
found on the coast of Sind near Karachi domesti- and Belutchistan, was selected.
cation may have taken place within the area of the Chowdhury and Buth (1970, 1971) found cotton
Harappan culture at Mohenjo-Daro. Pakistan at seed and hairs in Egyptian Nubia with an age of
about 2 400 BC. However, fragments of textile and about 2 500 BC. They suggested that because no
a string found at this site and dated 2 500-1 700 textile was found cotton was used as sheep fodder.
^ J 4"^.... • >
C^ r_^_y (3 3-2 t y
i xC2 yj*\ •s
0
k? ^^vy
Distribution of the Old World cottons in the 13th Century: Gossypium herbaceum v a r . africanum (1), G. herbaceum v a r . a c e r i f o -
h
lium (2), G. h e r b a c e u m v a r . p e r s i c u m (3), G. h e r b a c e u m v a r . kuljianum (4) and G. a r b o r e u m v a r . indicum (5) (Hutchinson, 1962)
J _ _ ^ 10 r& 0 <p A•
f o vVfâj J-
\y
tt.
1( N*VW il iL'
Ar( 10 )7 > i
•ft
Y-—-' I
\ ;
w) '
Distribution of annual cottons in the Old World in 1960: Gossypium herbaceum v a r . p e r s i c u m (3), G. herbaceum v a r , kuljianum
b
(4), G. a r b o r e u m v a r . indicum (5), G. a r b o r e u m v a r . bengalense (6), G. a r b o r e u m v a r . s i n e n s e (7), G. herbaceum v a r . wightia-
num (8), G. b a r b a r e n s e Egyptians (9), G. h i r s u t u m uplands and Cambodias (10) (Hutchinson, 1962)
AFRICAN CENTRE 122
Hibiscus acetosella (Menzel & Wilson, 1969) Hibiscus asper (Menzel & Wilson, 1969)
Hibiscus noldeae (Menzel & Wilson, 1969) Hibiscus meeusei (Menzel & Wilson, 1969)
MALVACEAE - MALVACEAE 123
Hibiscus cannabinus (Menzel & Wilson,1969) Hibiscus surattensis (Menzel & Wilson, 1969)
* »o *r-^J <'
v^ ' / ^^
^~~^S ^^~-^^^-^y :
^ Y
r J
.";,« V
" ^
WL"~~,/">/' -'-•4 (
MX^-\ '--/"'V'.\-'.V
\ °'•• \
U
Hibiscus mechowii (Menzel & Wilson, 1969)

AFRICAN CKNTRE 124
This ancient Nubian cotton is related to G. herba- HIBISCUS SABDARIFFA L. Rosella, Jamaica
ceum var. africanum and G. arboreum race s o r r e l . Guinea sorrel. Florida cranberry. 2n=(36),
soudanensis*. 72. genome formula AAYY. Africa. Angola is
From the race acerifolium race persicum* apparently its primary centre of dispersal. P r o -
and race kuljianum* and also G. arboreum race bably first domesticated as a dooryard or weedy
indicum* a r e derived. plant for its seeds. Later it became a vegetable
Either G. herbaceum reached S. America and finally a fibre crop (var. altissima Webster).
from America or G. arboreum reached Peru from It is a ruderal species of Angola, SW. Africa,
Asia by way of the Pacific islands to form the Zaire and Tanzania. Its A genome is derived from
amphiploid G. hirsutum* and G. barbadense*. H. asper* (Menzel &Martin, 1970). Its Y genome
At present many varieties belonging to G. donor is not yet known. Wilson and Menzel (1964)
hirsutum and G. barbadense a r e grown in Africa. noted the relationship of roselle and H. mechowii
G. hirsutum varieties (Upland and Cambodia) a r e Garcke. Var. sabdariffa is used on Jamaica to
cultivated in W. and C. Africa (race punctanum) produce a s o r r e l drink.
in Congo and E. Africa, while G. barbadense
(Egyptian) is found in the Nile valley and delta. HIBISCUS SCHIZOPETALUS Hook,f. 2n=45. E.
Var. africanum has a very simple protein Africa, Used there and elsewhere for hedges and
banding pattern (Cherry et al. , 1970). as an ornamental. Its uneven chromosome num-
ber suggests a hybrid origin. H. rosa-sinensis L.,
GOSSYPrUM LONGIOCALYX Hutch. & Lee. 2n=26, 2n=36. 46, 72, 92, c. 144, 168 being one parent.
genome formula ErE Uganda and Tanzania. It Van Borssum Waalkes (1966) concluded that as
has an entire leaf uke the American G. klotzschi- H. schizopetalus was first collected in E . Africa
anum*. H. rosa-sinensis might have an African origin.
However, negroid people in general do not culti-
GOSSYPIUM SOMALENSE (Guerke) Hutch. 2n=26, vate ornamentals and therefore a domestication of
genome formula E2E2. Sudan, Somaliland and H. rosa-sinensis in Africa is unlikely. More p r o -
Kenya. A variable species. bable is that H. schizopetalus a r o s e in E. Africa
after introduction of H. rosa-sinensis probably
GOSSYPIUM TRIPHYLLUM Hochr. 2n=26, g e - from Asia. Judging from the variable number of
nome formula BgBo. The desert regions of SW. this last species it might have a hybrid origin too.
Africa and S. Angola. It can be crossed with G. H. x archeri W. Watson is an artificial h y -
herbaceum* and G. arboreum*. brid of H. rosa-sinensis and H. schizopetalus.
HIBISCUS ACETOSELLA Welw. ex Hiern. Azedas, HIBISCUS SURATTENSIS L. 2n=36, genome for-
Red-leaved hibiscus. Bronze hibiscus. 2n-72. mula BB, (72). Africa. It also occurs in India,
Genome formula AABB. Tanzania, Zaire, Rhode- SE. Asia, Indonesia and Philippines. The B donor
sia and Angola. Cultivated in SW. Africa as a of H. acetosella* and H. radiatus*.
vegetable. Introduced as H. eetveldeanus De Wild.
&Dur. into Indonesia. Cultivated in the (sub)tro- Melastomataceae
pics as an ornamental. SAKERSIA LAURENTIA Cogn. 2n= . Zaire.
The A genome is almost homologue to the A
Cultivated there for its leaves (Terra, 1967).
genome of H. asper*. H. surattensis* is the B -
genome donor (Wilson &Menzel, 1964; Menzel & Menispermaceae
Martin, 1970). This species grows in western
trop. Africa. The closely related H. radiatus* CISSAMPELOS OWARIENSIS Beauv. Velvet leaf.
comes from Asia. 2n= . W. Africa. Cultivated there as a medi-
H. noldeae Baker f. appears to be a spiny, cinal crop in coastal regions (Dalziel, 1937).
inedible (primitive?) wild or weedy form of H.
acetosella (Wilson &Menzel, 1964). JATEORHIZA PALMATA (Lam. ) Miers. (syn.
J. miersii Oliv. ). 2n= . Trop. Africa. Culti-
HIBISCUS ASPER Hook. f. 2n=36, genome formula vated as a medicinal crop.
AA, 72. Wild in W. and C. trop. Africa. Wild
plants a r e collected for bast fibre. Occasionally Moraceae
cultivated for this purpose. Its A genome is close CHLOROPHORA EXCELSA (Welw. ) Benth. Iroko.
to the A genome of H. cannabinus*. It is one of 2n= . Africa. Cultivated there for its timber.
the parents of H. sabdariffa* and H. acetosella*
(Menzel &Martin, 1970). The A genome is also FICUS TILIAEFOLIA Bak. Voara. 2n= . Ma-
found in the African H. meeusei Exell (2n=26), dagascar. Cultivated there for its fibre.
genome formula AAXX (Menzel &Martin, 1971).
Moringaceae
HIBISCUS CANNABINUS L. Kenaf. 2n=36. Genome
formula AA. Probably (sub)trop. Africa. Also MORINGA PEREGRINA (Forsk. ) Fiori. 2n=
wild in Asia but these plants might derive from Egypt and Somaliland. Cultivated in the (sub)tro-
naturalized plants. Its A genome is related to that pics for the seeds which a r e the source of bennu-
of H. asper*. Kenaf is the A-genome donor of H. oil. This species is closely related to M oleifera*.
radiatus*.
MALVACEAE -PALMAE 125
Musaceae ry centre in Africa. Large areas in Africa a r e

ENSETE VENTRICOSUM (Welw. ) Cheesm. (syn. covered by semi-wild palms. In Africa the oil
E. edule (Horan) Cheesm. ). Ensete, Inset, Abessi- palm was only in a few areas domesticated prior
nian banana. 2n-18. Ethiopia, the mountains of to the establishment of 'European' plantations and
Kordofan and the lower part of the montane forest 'development' f a r m e r ' s plots (Zeven, 1967, 1972).
belt of Ruwenzori. Cultivated for the flour ob- Large plantations are found in Africa, SE. Asia
tained from the pseudostem and also for its fibres. and in C. America.
It is propagated by offshoots and by seeds from
cultivated types or occasionally from wild plants. PHOENIX ATLANTICA Chev. False date. 2n=36.
The cultivated plants a r e grown on higher altitudes Africa. It closely resembles Ph. dactyliSera*.
than the wild plant itself grows. Several cultivated Near Marrakesh in Morocco var. marocana Chev.
types a r e recognized. It is suggested that there is cultivated. It has fairly tasty fleshy fruits while
a r e about forty different ensete types. The back those of this species are, in general, of poor
of the leaf of the Koba type is red to purple c o - quality (Meunier, 1962).
loured. This variety has been described as var.
montbeliardi D. Bois (Smeds, 1955). PHOENIX CANARIENS1B Hort. (syn. Ph. jubae
Christ.) 2n=36. The Canary Islands. Spread as
It is suggested that ensete is one of the oldest an ornamental to N. Africa and S. France, Close-
cultivated plants of Ethiopia. ly related to Ph. dactylifera* and easily hybridi-
zes with this and other Phoenix species.
MUSA cultivars of the AAA group. Banana. 2n=33.
P r i m a r y centre in the Malayan region (p. 52). PHOENIX DACTYLIFERA L. Date palm. 2n-28.
Secondary centre in the uplands of E. Africa. P r i m a r y gene centre: probably N. Africa. One of
There it has been cultivated for a long time. the oldest cultivated plants and cultivated for a
long time from the Atlantic to NW. India.
MUSA cultivars of the AAB group - Plantains s u b - Phoenix species easily hybridize. This may
group. French plantain, Horn plantain. 2n=33. have resulted in an increased variability. P e r -
India (p. 68). Secondary centre in E. Africa. haps all mentioned Phoenix* species should be
included in one species.
Palmae
BORASSUS FLABELLIFER L. Lontar, Palmyra

palm. 2n=36. India and Malayan Archipelago.
V
(p
Cultivated there. Unknown wild. Probably a cul-
^
tigen of the African B. aethiopum Mart. (2n= ). if o v \
ELAEIS GUINEENSIS Jacq. Oil palm. 2n=32. The 3**^ws
coastal belt from Sierra Leone to Angola. P r i m a - fSgtLi»,. <•
f /< - -v vS *w ~?y
V_-^ kfj H
\ «V
Phoenix dactylifera (Oudejans, 1969)
PHOENIX HUMILIS Chev. 2n= . Cameroons.

There palms are wild and semi-wild. In the latter
case they a r e protected, but not planted. They are
in a pre-domestication stage.
PHOENIX RECLINATA Jacq. False date palm.

2n=36. W. Africa. Some reports refer to this
palm as being cultivated as a wine palm. However
these may refer to Ph. humilis* (Porteres,
1955b).
PHOENIX SYLVESTRIS Roxb. Wild date palm.

2n=36. Pakistan, India and S. Iran. Cultivated
there as a source of sugar and wine. Closely r e -
lated to Ph. dactylifera* and may have been d e -
rived from it, or they may have a common p r o -
genitor.
E l a e i s guineensis (Zeven, 1967)
AFRICAN CENTRE 126
RAPHIA HOOKERI Mann & Wendl. Wine palm.

2n= . The 100-250 km wide coastal belt of
W. Africa (Russell, 1965). Highly valued for its ! i •> .
wine and fibre properties and therefore cultivated 1
J---~ '' /''
and cared for. In SE. Nigeria it was observed
that pedlars sold germinated fruits from the Imo /T\V J
/'•'""
--4, f
river banks to farmers up to 100 km away.
VV>6-0; A
Pandanaceae
PANDANUS UTILE Bory. 2n= . Introduced
elsewhere as an ornamental, while leaves are
V Sc -* ' i.ii
-4--; \
used for various purposes. Cultivated in Mauri-
tius for making sugar bags. Polygala butyracea (Portéres, 1950)
Pedaliaceae
Polygonaceae
CERATOTHECA SESAMOIDES Endl. Bungu.
2n=32. W. Africa. Cultivated in some northern RUMEX ABYSSINICUS Jacq. Spanish rhubarb dock.
a r e a s . It yields leaves for soups and oily seeds. 2n= . Ethiopia. Cultivated in the Zaire basin.
SESAMUM ALATUM Thonn. Tacoutta. 2n=26. Portulaeaceae

Trop. Africa or India (p. 69). Occasionally cul- TALINUM CUNEIFOLIUM (Vahl) Willd. 2n=
Africa and Arabia. Cultivated in E. Africa as a
vegetable.
SESAMUM INDICUM L. (syn. S. orientalis L. ).
Oriental sesame, Beni seed. 2n=26, (52, 58). Un- TALINUM PORTULACIFOLIUM (Forsk. ) Aschers.
known wild. Secondary centre: in India (p. 69) and 2n= . Trop. Africa and Asia. Cultivated in
in China/Japan (p. 35). An ancient crop, much Africa as a vegetable.
cultivated, at present, in India, China, Japan,
Burma, NW. Africa, Americas and Europe. As TALINUM TRIANGULÄRE Willd. 2n=48, 72.
all wild Sesamum species but one (S. prostratum Probably C. and/or S. America or Trop. Africa.
Retz. (2n=32), wild in E. India) occur in Africa Cultivated in Brazil (p. 156), West Indies and W.
it is thought that its progenitor(s) a r e African. Africa as a vegetable. The cultivation in forest
Spontaneous tetraploids have been observed. regions may indicate an African origin, but as
Nayar and Mehra (1970) considered S. indi- species of this genus are native to Africa and to
cum var. malabaricum (2n=26) as a possible the New World further investigation is necessary.
'companion weed' of sesame. It may have origi-
nated from hybrids between sesame and some Rosaceae
sympatric wild Sesamum species.
HAGENIA ABYSSINICA J . F . Gmel. (syn. Brayera
SESAMUM RADIATUM Schum. &Thonn. 2n=64. anthelmintica Kunth. ). 2n= . Ethiopia. Culti-
Trop. Africa. Cultivated in C. and W. Africa for vated there for its flowers used as medicine
its seeds.
Rubiaceae
Perioplocaceae COFFEA ARABICA L. Arabica coffee. 2n=22, 44,
CRYPTOSTEGIA GRANDIFLORA Br. 2n=24. (66, 88). P r i m a r y centre: SW. Ethiopia (Meyer,
Trop. Africa or India (p.000). Occasionally cul- 1969). Secondary centre: Yemen (p. 90). Traditio-
tivated for its Palay rubber and often as an orna- nally the arabica coffee has only been known in
mental. cultivation. Cultivated now over large a r e a s .
Arabica coffee is the only known Coffea s p e -
Piperaceae cies being allopolyploid and self-compatible. Its
parental species a r e not known, but closest r e l a -
PIPER CLUSE DC. 2n= . W Africa. Culti- tives occur in C. and W. Africa (Meyer, 1969).
vated as a spice. Kammacher &Capot (1972) suggested that one of
the genomes has a similar structure to the g e -
PIPER GUINEENSE Schum. & Thonn. Guinea nome of C. canephora*.
pepper, Ashanti pepper. 2n= . W. Africa. Various botanical and agricultural varieties
Cultivated there as a spice. a r e known and so a r e many mutants. An example
is the mutant discovered on Réunion, formerly
Polygalaceae Bourbon which became the highly productive Bour-
POLYGALA BUTYRACEA Heck. Cheyi, Numbu- bon coffee (Meyer, 1965).
ni. 2n= . W. Africa. This plant probably does
not exist in the wild. It is probably a relic of an COFFEA CANEPHORA P i e r r e ex Froehner (syn.
ancient tropical West African culture. However, C. robusta Linden). Robusta coffee. 2n=22, 44.
more evidence is needed. Cultivated in W. Africa Africa, western to central (subtropical regions,
for its fibre and seed. from Guinea and Liberia to Sudan and Uganda.
PALMAE - STERCULIACEAE 127
The greatest diversity has been described for (Don. ) Hepper, Vitellaria paradoxa Gaertn. f. ).
Zaire. Karité, Shea butter t r e e . 2n=24. Semiwild in the
Before the arrival of the Europeans in Africa savannas of West Africa.
it was already cultivated there. Cultivated now
especially in Indonesia and because it is used to CHRYSOPHYLLUM AFRICANUM A. DC. African
prepare 'instant' coffee its cultivation increased star apple. 2n=26. Trop. Africa. Cultivated for
in other tropical Asian and African countries. its fruits.
It is a cross-fertilizer and hence very poly-
morphic. This has resulted in several synonymes, Solanaceae
'Congusta' coffee is probably a hybrid of C.
canephora and C. congensis* although the latter is SOLANUM ACULEASTRUM Dunal. 2n=24. Trop.
considered to be a form of C. canephora. Some Africa. This non-tuberous plant is used for g r o -
wing hedges.
botanical and agricultural varieties a r e described.
COFFEA CONGENSIS Froehner. 2n=22, (44). The SOLANUM AETHIOPICUM L. 2n=24. Trop. Africa.
Zaire basin. It resembles C. arabica*. Possibly This non-tuberous plant is cultivated for its
a form of C. canephora*. 'Congusta' coffee is a leaves and fruits.
hybrid product of C. congensis and C. canephora.
SOLANUM ANOMALUM Thonn. Children's tomato.
COFFEA EUGENIOIDES S. Moore. 2n=22. Wild 2n= . Trop. Africa. This non-tuberous plant
in the Lake Kivu area of Zaire, W. Uganda and is sometimes cultivated.
W. Tanzania. Cultivated there. It resembles a
slender form of C. arabica*. SOLANUM BURBANKn Bitter. Wonderberry.
Msoba. 2n= 6x=72. Probably derived from the
COFFEA LIBERICA Bull. Liberica coffee. 2n=22, southern African msoba (Heiser, 1969). It is appa-
44. Guinea to Angola. Cultivated to some extent rently not a hybrid of S. sarachoides (syn. S.
in Liberia, Surinam and a few other countries. It villosum) (2n= ) and S. melanocerasum Allioni
is a cross-fertilizer and hence very polymorphic. (syn. S. guineense Lam. non L. ), Garden Shuckle-
It has been crossed with C. arabica to produce berry (2n=72), but a contaminant.
hybrids which a r e cultivated.
This species includes the Excelsa coffee SOLANUM DUPLOSINUATUM Klotsch. 2n=
(C. excelsa Chev., syn. C. liberica var. dewe- Trop, and southern Africa. Cultivated for its
v r e i De Wild. &Dew., syn. C. arnoldiana De fruits and leaves.
Wild. ).
There is a possibility that in the ancestry of SOLANUM INCANUM L. 2n=24. Africa. Occasio-
this species some introgression with C. canephora* nally cultivated.
has occurred (Chinnappa, 1970).
SOLANUM MACROCARPON L. 2n=36. Mascarene
VANGUERIA MADAGASCARIENSIS J. F. Gmel Islands. Cultivated for its leaves. The cultivar is
(syn. V. edulis Vahl. ). 2n= . Trop. Africa described as var. calvum Bitter. Its triploidy may
and Madagascar. Cultivated for its edible fruits. point to a hybrid origin.
Rutaceae SOLANUM NIGRUM L. Black nightshade. 2n=24,

(36, 40, 48), 72, (96, 144). Native region is un-
ADENANDRA FRAGRANS (Sims. ) Roem. & Schult. known. At present a cosmopolite non-tuberous
2n= . S. Africa. Cultivated there for its leaves weed which is cultivated.
which are used to decoct tea.
SOLANUM NODIFLORUM Jacq. 2n=24, 72. The
BAROSMA BETULINA (Berg. ) Bartl. & Wendl. Sahara and Nigeria. Cultivated for its leaves.
Buchu. 2n= . SW. and S. Africa. Cultivated May have run wild elsewhere.
on a small scale in the Chanwilliam district.
Leaves of wild and cultivated crops a r e used for SOLANUM OLIVARE Paill. 2n= . Cultivated
their medicinal properties (Gentry, 1961). in Ivory Coast, Dahomey and Congo.
CITROPSIS GILLETIANA Swing. &Kell, and other SOLANUM ROTUNDIFOLIUM Moric. ex Dun.
Citropsis species. Trop. Africa. Closely related (syn. S. nelsoni Dun. ). Hausa potato. 2n=
to Citrus. They can be used as citrus rootstocks. Believed to come from Ethiopia. Spread to W.
Africa and other parts of Africa.
Sapindaceae
BLIGHIA SAPIDA Koenig. Akee. 2n=32. Forests Sterculiaceae
of W. Africa. Cultivated in Jamaica and W. Afri- COLA ACUMINATA (P. Brenan) Schott. &Endl.
ca. In Jamaica it has been naturalized. Abata kola. 2n=40. Nigeria to W. Gabon. Spread
to Zaire and Angola, to the West Indies and e l s e -
Sapotaceae where. Cultivated esp. in W. Nigeria, but is
BUTYROSPERMUM PARKÜ (Don. ) Kotschy (syn. second in importance to C. nitida*.
B. paradoxum (Gaertn.f.) Hepper ssp. parkii
AFRICAN CENTRE 128
Aframomum melegueta (van Harten, 1970)
COLA ANOMELA K. Schum. Bamenda kola. Zingiberaceae

2n= . Cameroon, esp. in Bamenda. Cultivated
there. AFRAMOMUM MELEGUETA (Rosc.)K. Schum.
Melegueta pepper. 2n= . West African coastal
COLA NITIDA (Vent. ) Schott &Endl. Gbanja kola. belt from Guinea to Angola, including Fernando
2n=40. Sierra Leone to Dahomey, with its highest Po and San Thomé (van Harten, 1970). Probably
frequency in the forest a r e a of Ivory Coast and not cultivated in W. Africa. After its introduction
Ghana. The genus Cola has its primary centre in into S. America, cultivated in Surinam and Guyana.
W. Africa (van Eijnatten, 1969, 1970). Spread in It is the historical known 'Grains of paradise',
giving its name to the West African Pepper Coast,
W. Africa and fruits were taken to the Caribbean
Grain Coast or Malagueta Coast.
where this kola already grew in 1630. Introduced
to other tropical countries. This is the main kola
of commerce. Subspecies refer to fruit colour,
but this may be caused by some genes conditioning
these colours.
COLA VERTICILLATA (Thonn. ) Stapf ex Chev.

Owe kola. 2n= . From Ivory Coast to lower
Zaire. Often found as stray individuals in plantings
of C. nitida*. On the Mambilla Plateau in N. Ni-
geria it is the only kola found (van Eijnatten, 1970).
Tamaricaceae
TAMARIX ARTICULATA Vahl. 2n= . The
Sahara, Arabia and Iran. Great numbers a r e found
in S. Morocco and Mauritiania. Cultivated as a
windbreak for orange cultivation, as a sandbinder,
for fuel and as ornamental.
Tiliaceae
CORCHORUS TRILOCULARE L. Al Moulinouquia.
2n=14. Senegal to India. Cultivated sometimes as
a vegetable e.g. near Timbuktu, Africa (Uphof,
1968).
Verbenaceae
LIPPIA ADOENSIB Höchst. Gambian tea bush.
2n= . Zaire. A potherb cultivated there. In
W. Africa it is used as a tea substitute.
VITEX CIENKOWSKII Kotschy &Peye. 2n=32.

Trop. Africa. A tree planted on compounds or
semi-cultivated for its fruits.
9 European Siberian Centre
f =&/?/u v, $ 'V 1
\ ^~\
J>
?4i
J\ Ja rJ
VU ..?>*
ö r,.'' -
(; J
~^x
.Z&'X Ï — . ',- - ^ --rt <a
The European-Siberian Centre was not indicated by Vavilov. Darlington (1956) was the first to refer to
Europe as a region of origin of crop plants. Zhukovsky (1968, 1970) recognized it as a megacentre of
diversity of a relatively small importance.
Agriculture reached this region from Centre 6 and arrived at about 4 000 BC. in NW. Europe.
Important crops have been developed in this region: fruit t r e e s , grasses, Brassica s p . , Cannabinus
sativus, Cichorium s p . , Digitaria sanguinalis, Fragaria s p . , Lactuca sativa, Humulus lupulus, Medi-
cago s p . , Ribes s p . , Rubus s p . , Trifolium s p . , etc.
Alliaceae cinal herb in most of Europe, From this cultivation

ALLIUM AMPELOPRASUM L. Levant garlic. it has naturalized.
Perennial sweet leek. 2n=16, 24, 32, genome for-
mula AAA'A", 40, (48, genome formula AAA'A' Asclepiadaceae
A " A " , AABBBB). Europe, Asia Minor, Cauca- CYNANCHUM VINCETOXICUM (L. ). P e r s .
sus to Iran and N. Africa. Cultivated in S. France Swallows wort. 2n=22. Europe to Himalaya and
and around Nürnberg, Germany for its bulbs Altai, and in N. Africa. A perennial herb culti-
(Kuckuck, 1962). Some cultivation also in Kashmir vated formerly in gardens as a medicinal plant.
(p. 62) and Iran (p. 77).
Berberidaceae
ALLIUM SCORODOPRASUM L. Giant garlic, Sand BERBERIS VULGARIS L. European barberry.
leek. 2n=16. C. and S. Europe and Asia Minor. 2n=28. Most of Europe and Caucasus. Difficult to
Cultivated in USSR (Kuckuck, 1962).
a s s e s s its territory because it was planted for-
merly for its edible berries and is now an orna-
Araceae mental. Wood and bark were used to produce a
ACORUS CALAMUS L. Sweet flag, Sweet root, yellow dye. It is an intermediate host of stem rust
Calamus. 2n=18, 24, 36, (44, 48). N. Europe, (Puccinia graminis P e r s . ) and has therefore been
temperate Asia and E. North America. Used as a extirpated on a large scale.
medicinal plant, as an ornamental and for the root
that is used for various purposes such as p r e p a - Boraginaceae
ration of oil. Widely cultivated now, but roots of
LITHOSPERMUM OFFICINALE L. Gromwell.
wild plants are still collected and used.
2n=28. Spp. officinale throughout Europe, W.
Asia, Caucasia and Iran. Cultivated formerly in
Aristolochiaceae Bohemia for preparing Bohemian or Croatian tea.
ARISTOLOCHIA CLEMATIS L. 2n=14. Probably Spp. erythrorhizon is cultivated in China and
E. and S. Europe. Cultivated formerly as a medi- Japan (p. 28).
EUROPEAN SIBERIAN CENTRE 130
Campanulaceae CHENOPODIUM BONUS-HENRICUS L. (syn. Ch.

esculentus Salisb. ). Allgood. Good King Henry.
CAMPANULA RAPUNCULUS L. Rampion, Ramps.
2n=36. A Paleotemperate native. Cultivated for-
2n=20, 102. Europe, N. Africa, SW. Asia and
merly as a potherb.
Siberia. Cultivated in the Middle Ages for its
fleshy roots.
CHENOPODIUM FOLIOSUM Aschers. 2n=18.
Cannabidaceae Europe and the Orient. Cultivated formerly as a
vegetable (Uphof, 1968).
CANNABIS SATIVA L. Hemp. 2n=20. The wild
form (C. ruderalis Janisch) is found in C. Asia. Compositeae
It is marked by a horseshoe-shaped scar at the
base of the achene. C. sativa is one of the e a r - ANTHEMIS NOBILE L. Noble chamomile. 2n=18.
liest cultivated crops. It had reached China by S. andW. Europe. Cultivated as a medicinal
2 500 BC. Cultivated for its fibre and for its plant.
seeds, for food or as a source of hemp seed oil.
The Indian type (p. 63) is cultivated as a source ANTHEMIS TINCTORIA L. Dyer's chamomile,
of narcotics. Special cultivar groups have been Golden chamomile. 2n=18. Europe and W. Asia.
developed for different purposes. Cultivated as a dye plant.
HUMULUS LUPULUS L. Hop. 2n=20. Wild plants ARCTIUM LAPPA L. (syn. Lappa arctium
(var. lupulus) in Europe. Hop is especially culti- Gaertn. ). Great but, Great burdock. Cockle bur.
vated in Europe and N. America. There a r e two 2n=32, 36. Europe and Asia. Cultivated in Europe
cultivated types: convar. europaeus Mandy with as a medicinal plant, and also in China and Japan
divided leaves and convar. cordifolius (Miq.) (p. 29).
Maxim, (syn. H. cordifolius Miq. )with entire
heart-shaped leaves cultivated in E. Asia. ARTEMISIA ABROTANUM L. Southern wood.
2n=18. S. Europe and temp. Asia. Cultivated as
a medicinal crop.
Caryophyllaceae
SAPONARIA OFFICINALIS L. Soapwort, Soap- ARTEMISIA ABSINTHIUM L. Absinthe. 2n=18.
root. 2n=28. Europe and Asia. Cultivated occasio- Europe, S. Siberia, Kashmir and Mediterranean
nally in Germany (Mansfield, 1959). region. Cultivated in S. Europe, N. Africa and
the USA for the production of absinthe.
SPERGULA ARVENSIS L. (syn. Spergularia a r -
vensis Cambess. ). Corn spurrey. 2n=18. Europe. ARTEMISIA LAXA Fritsch. 2n=l£ C. and S.
Var. sativa (Boenningh. ) Mert. &Koch (syn. Europe. Cultivated.
S. sativa Boenningh. ). Cultivated as a fodder crop
or as a green manure. Var. arvensis is a wide- ARTEMISIA MARITIMA L. 2n=18, 36, 54. Europe
spread weed, while var. maxima (Weihe) Mert. to Mongolia. Cultivated as a medicinal crop.
&Koch, is a weed in flax fields.
ARTEMISIA VULGARIS L. Mugwort. 2n=16, 18.
Chenopodiaceae Temp. N. Hemisphere. Cultivated in Indonesia
ARTIPLEX HORTENSIS L. Mountain spinach, and elsewhere.
Garden orach. 2n=18. Wild in temperate Europe
and Asia. Cultivated formerly in Europe as a CARUM CARVI L. (syn. Apium carvi Crantz).
Caraway. 2n=20, 22. Europe and W. Asia. Culti-
vegetable.
vated in temperate regions and in N. India and
Sudan (see C. roxburghianum*, C. copticum*).
BETA VULGARIS L. var. rapa. Fodder beet.
2n=18. Distribution of the the wild type is given
on p. 92. Developed probably in the Netherlands CICHORIUM ENDIVIA L. Endive. Escarolle.
and perhaps from types introduced from Spain. 2n=18. S. Europe to India. P r i m a r y centre in the
Secondary centre in Centre 5 (p. 71). Spread to Mediterranean region.
Germany and elsewhere. It may have played a
role in the development of sugarbeet, var. CICHORIUM INTYBUS L. Chicory, Succory,
saccharifera (syn. var. altissima). The s u g a r - Brussel Witloof. 2n=18. European Siberia, N.
beet probably developed in Silecia, Poland from Africa and the Near East to Iran, Belutchistan
hybridization of an old garden form and fodder and Lake Baikal. Wild type (var. intybus) was
beet. The variety "Weisser schlesicher Zücker- used as a salad and for medicinal purposes. Var.
rübe" is the parent of all sugarbeet varieties. sativum Lam. &DC. is cultivated in Europe and
elsewhere to produce a coffee substitute while
var. foliosum Hegi, the Brussel witloof, was
CHENOPODIUM ALBUM L. Goosefoot, Fat hen,
first developed around Brussels, Belgium.
Lamb's q u a r t e r s . 2n=18, 36, 54. Probably culti-
vated in Europe in Neolithic times. Now it is a
weed. HELIANTHUS ANNUUS L. Sunflower. 2n=34. Wild
in N. America (p. 173). Secondary centre in USSR.
Domesticated and cultivated in N. America. Large-
headed forms introduced in Europe.
LACTUCA QUERCINA L. 2n=18. Europe, esp. in Cruciferae

Germany, France to USSR and the Balkan. A
ARMORACIA RUSTICANA (Lam. ) Gaertner, Mey
biennal. Sometimes cultivated near Clermont-
&Schreb. , (syn. Cochlearia armoracia L. ).
Ferrand (France) for its narcotic properties
Horse radish. 2n=28, 32. Finland, to Poland,
(Uphof, 1968).
the Caspian Sea and the deserts of Cuman and in
Turkey. P r i m a r y centre in temperate E. Europe
LACTUCA SATIVA L. Lettuce. 2n=18. P r i m a r y (Counter &Rhodes, 1969). Cultivated for its culi-
centre: the Middle East. Lettuce derives from nary purposes and hence naturalized.
L. serriola L., the prickle lettuce. This species
occurs in S. and C. Europe to Denmark, Cauca-
sia, Transcaucasia, Iran, Iraq, Syria, Saudi BARBAREA PRAECOX R . B r . (syn. B. verna
Asch. ). Scurvy grass, Winter c r e s s . Upland
Arabia, Siberia to Altai and N. Africa to the
c r e s s . 2n=16. Europe. Cultivated as a vegetable
Canary Islands. However, Lindqvist (1960) b e -
lieved that lettuce probably derives by hybridiza-
tion of other Lactuca species including L. saligna BARBAREA VULGARIS R . B r . Yellow rocket
L. and that L. serriola arose from the same or Common winter c r e s s . Upland c r e s s . 2n=16.
subsequent hybridization. L. serriola is now a Moderate Europe, Asia and N. Africa. Spread
weed. L. saligna like L. serriola has its main throughout the world. Cultivated as a potherb.
distribution centre round the Mediterranean sea
(Lindqvist, 1960). BRASSICA CAMPESTRIS L. Turnip group. 2n=20,
genome formula AA. The wild form spp. sylves-
The first record of lettuce dates from 4 500 BC. ;
t r i s (L. )Jancken grows as a weed and ruderal in
a long-leaved form was depicted on the Egyptian
most of Europe, moderate Asia and N. Africa.
tombs.
The various oily, spp. oleifera (Metzg. ) Sinsk.
The present marked variation of lettuce is proba- (oil seed turnip) and fodder, spp. rapifera (Metzg.)
bly a product of hybridization with L. serriola, Sinsk. ("stubble turnip", Dutch turnip) cultivars
but may also have been induced by some natural have been developed independently.
mutation (Whitaker, 1969). There a r e three main groups: the Asian (p. 29),
Var. asparagina Bailey (syn. L. angustana Vilm., the Mediterranean and the West European.
L. sativa var. angustana Irish), asparagus lettuce, The development of the turnip rape var. oleifera
celtuce forms a single, thickened straight stem of (Metzg.) Sinsk. possibly took place in Belgium.
90 cm or more which is eaten as salad when young. Leafy types of turnip are cultivated especially in
Finland.
MATRICARIA CHAMOMILLA L. Chamomille,
German chamomille. 2n=18. Europe, Iran and The A genome is also found in the diploid B. chi-
Afghanistan. Cultivated in Europe as a medicinal nensis* and the diploid B. japonica*. This genome
and as a source of an essential oil used for fla- is related to the Ad genome of B. adpressa Boiss. ,
vouring and perfumery. A substitute of Anthémis the F genome of B. fruticulosa Cyril, and the D
nobilis*. or T genome of B. tournefortii Gouan (Mizushima,
1969).
TARAXACUM HYBERNUM Steven. Krim sagiz. B. campestris is one of the parents of B. juncea*
2n=32, 40. Italy, Balkan, Asia Minor, Syria and and B. napus*, and also of the artificially made
Crimea. Cultivated in USSR as a rubber crop. B. napocampestris (2n=58, genome formula
AiAiAACiCi).
TARAXACUM OFFICINALIS Weber. Dandelion,
BRASSICA NAPOBRASSICA (L. ) Mill. (syn. B.
Lions-tooth, Milk-gowan, Puffball. 2n=8, 24
napus L. var. napobrassica (L. )Rchb. ). Ruta-
(and others). Europe and W. Asia. In France and
baga, Swedish turnip. 2n=38. Wild unknown. P r i -
elsewhere improved varieties a r e cultivated.
mary gene centre in the Mediterranean region
These varieties "Pissenlit â coeur plein amélioré"
(p. 94). Secondary gene centre in Europe. P r o b a -
and "Pissenlit vert de Montmagny" differ from
bly a derivative of B. oleracea* x B. napus*.
wild plants (pissenlit ordinaire) as they have less
The roots a r e more elongated and oval and larger
bitter leaves. Young etiolated leaves of wild plants
than those of turnip. They a r e consumed as a
covered by mole-hills are collected as dandelion
vegetable.
salad.
BRASSICA NIGRA (L. ) Koch. Black mustard.
Crassulaceae
2n=16, genome formula BB. Europe, especially
SEDUM REFLEXUM L. Jenny stone crop. 2n=34, in C. and S. p a r t s . Cultivated since ancient times.
c. 56, 68, c. 112. S. Europe. Cultivated in W. Seeds a r e p r e s s e d for black mustard-seed oil.
and C. Europe and used to flavour soup and salad. The B genome is related to the F genome of B.
fruticulosa (Mizushima, 1969). Black mustard is
SEMPERVIVUM TECTORUM L. Hen-and-Chickens, one of the parents of B. juncea* and B. carinata*.
Roof houseleek. 2n=(36), 72. Europe. Cultivated An artificial amphiploid of B. tournefortii* and
as a medicinal plant. this species is called B. amarifolia (2n=36), g e -
nome formula TTBB or DDBB).
BRASSICA OLERACEA L. var. gemmifera DC.

Brussels sprouts. 2n=18, genome formula CC.
Developed in Belgium probably from var. ramosa.
COCHLEARIA OFFICINALIS L. Spoonwort, Scor- AGROPYRON INTERMEDIUM (Host. ) Beauv.

bute grass, Scurvy g r a s s . 2n=(14), 24, genome (syn. A. glaucum). Wheat g r a s s . 2n=(28). 42, ge-
formula A„AgAgA„, (28, 36). N. and W. Europe. nome formula BgBgE.EjEgEo. S. and C. Europe
Cultivated formerly as a medicinal plant. to Iran and Caucasia.
CRAMBE MARITIMA L. Sea kale. 2n=60. Sea AGROPYRON REPENS (L.) Beauv. Quackgrass.
coast of Europe. Cultivated in England as a vege- 2n=28, 42, (56). Temperate Eurasia. Cultivated.
table. A tedious pestweed.
HESPERIS MATRONALIS L. Damask. 2n=24. AGROSTIS CANINA L. 2n=14, 28, (35, 42, 56).
C. and S. Europe. Cultivated for its seeds which Europe. Cultivated in the Netherlands.
a r e a source of oil and as an ornamental. E s -
capes a r e common. AGROSTIS GIGANTEA Roth. (syn. A. alba auct,
non L. ). Fiorin, Red top. 2n=42, genome formula
NASTURTIUM OFFICINALIS R. Br. (syn. Rorippa A . A ^ p A g A A„. Europe, Asia and N. America.
nasturtium-aquaticum (L.) Hayek). Watercress. Cultivated as a pasture grass and as a hay crop.
2n=32, (48, 64). W. Asia and S. Europe and Great
Britain. Cultivated. The tips of the leafy stems AGROSTIS TENUIS Sibth. (syn. A. vulgaris With.)
a r e eaten as salad. It is also cooked as a vege- Rhode Island bent, Colonial bent. 2n~28, genome
table. In New Zealand it is a serious river-weed. formula AjAÂÂ Most of Europe, N. Asia
The almost sterile hybrid plants (2n=45) of water- Minor. Armenia, ^Caucasia, Siberia, N. Africa
c r e s s and its relative N. microphyllum (Boenn.) and N. America. Hybrids with A. gigantea* have
Rchb. (2n=64) are also cultivated for salad (Purse- been found in Germany and called A. intermedia
glove, 1968). It is vegetatively propagated. C.A. Weber.
Cucurbitaceae ALOPECURUS PRATENSIS L. Meadow foxtail.

BRYONIA ALBA L. White bryony. 2n=20. C. 2n=28, (42). Most of Europe, N. Asia and Cauca-
Europe. USSR, the Balkans and N. Iran. Cultiva- sia. Cultivated as a meadow g r a s s .
ted formerly as a medicinal plant.
AMMOPHILA ARENARIA Link (syn. A. arundina-
BRYONIA CRETICA L. Red berry bryony. 2n=20. cea Host. ) . Beach g r a s s . 2n=28. The coastal
S., SC. and W. Europe to Britain and N. Africa. areas of Europe. A perennial cultivated as a sand
Cultivated formerly as a medicinal crop. Spp. binder.
cretica is found in the Aegian region, spp. dioica
(Jacq. ) Tutin (syn. B. dioica Jacq. )has a wide ANTHOXANTHUM ODORATUM L. Sweet scented
distribution, while spp. acuta (Desf. ) Tutin (B. vernal grass, Spring g r a s s . 2n=10, 20. Europe,
acuta Desf. ) is found in Tunesia and Libia. Asia, W. of N. Africa. Cultivated as a forage
g r a s s . It has a low food value. The diploid is also
Cyperaceae described as A. alpinum Löve &Löve. Autoploidy
has played an important role in the genesis of the
CAREX ARENARIA L. (syn. C. spadicea Gilib.). tetraploid (Hedberg, 1970). Tepper (1970) sugges-
2n=58, 60-64. Europe, especially the littoral ted the following genome formula for A. alpinum
a r e a s . Cultivated as a soil stabilizer. and A. odoratum:
SCIRPUS LACUSTRIS L. (syn. S. validus Vahl. ). species ploidy

Great bulbrush. 2n=(38, 40), 42. A world wide formula
distribution. Cultivated in the Netherlands and A. alpinum 2x AA general
Germany, to promote land reclamation and i m - A. alpinum 4x AAAA Cantal, France
prove impoldered land. In Germany cultivated to A. odoratum 2x CC Italy
clean polluted water and so it is expected that the A. odorarum 2x DD Italy, Yugo-
planting will increase and better varieties of this slavia, Greece
plant will be bred. Its culms contain 80% a i r taken A. odoratum 2x DE Serbia
from the atmosphere. They absorb a i r pollutant A. odoratum 4x BBDD Southern C.
gases, sodium, phosphorus, zinc and copper. and W. Europe
A. odoratum 4x BBFF W. Europe
Gramineae
A comparison of Austrian, Swiss, Swedish and
AGROPYRON CANINUM P . B . (syn. Roegneria
Polish populations showed that diploids from Aus-
canina (L.) Nevski). 2n=28. Cultivated in the
USSR. tria and Switzerland a r e morphological closer to
those from Poland than to those in Scandinavia
(Hedberg, 1969).
AGROPYRON CRISTATUM L. Gaertn. Crested
wheatgrass. 2n=14, 28, (42). Europe and Asia.
Introduced into N. America. Cultivated there as a ARRHENATHERUM AVENACEUM Beauv. (syn.
hay crop. This species includes a number of other A. eliator Beauv. ). Tall meadow oat g r a s s . 2n=40.
species like A. desertorum (Fisch.) Schult., A. Europe. A valuable pasture g r a s s .
pectiniforme Roem. &Schult., A. michnoi Roshev.
and A. sibiricum (Willd. ) P .B.
CRUCIFERAE - GRAMINEAE 133
ARRHENATHERUM TUBEROSUM Druce (syn. widely distributed.

Avena tuberosa Gilib. , Arrhenatherum avenaceum
Beauv. ). Onion couch grass. 2n=28. In neolithic FESTUCA ARUNDINACEA Schreb. 2n=(28), 42,
times possibly cultivated for its tubers. (70). Europe, N. Africa and Asia (Syria, Siberia,
Japan). Not much cultivated, due to its coarseness
AVENA SEPTENTRIONALIS Malz. (syn. A. fatua although seeds have been commercially available
spp. septentrionalis (Malz. ) Malz. ). 2n=42. N. for a long time.
and NE. European USSR to W. Siberia. There it According to Borrill (1972) the tetraploid and hexa-
usually grows in undisturbed habitats. Baum ploid cytotypes have affinities with F. pratensis*,
(1972) stated that it is probably the most closely while the octoploid and decaploid possess a genome
related taxon to A. sativa* and that it resembles pair of F . scariosa Aschs. & Graebn. (2n=14).
the hypothetical ancestor of the predomesticated This species is endemic in the Spanish Sierra Ne-
oats. vada.
F. arundinacea has been rather widely introduced
BROMUS ERECTUS Huds. (syn. B. arvensis as a meadow and pasture g r a s s in northern USA.
P o l l . ) . 2n=(28), 42, 56. (70, 84, 112). C. and S.
Europe, N. Africa, Ante-Asia up to Caucasia. FESTUCA OVINA L. Sheep's fescue. 2n=14, (21),
Cultivated especially in S. France, Switzerland, 28, 42, (49), 56, 70. Europe, the Caucasus, the
S. Germany and USSR. Some people regard this Himalaya and N. America. Cultivated in Europe.
species and its synonym as two species. An important grass of Australia and S. Africa.
BROMUS rNERMLS Leyss. Awnless brome, Smooth FESTUCA PRATENSIS Huds. (syn. F. elatior L. ).
brome, Hungarian brome. 2n=(28, 42, 49), 56, Fescue grass, Meadow fescue, English bluegrass.
(54-58). N., C. and SE. Europe, Caucasia, t e m - 2n=14, (28, 42, 70). Europe, Caucasia, Iran, the
perate Asia to China. Cultivation started at v a r i - Ural and Siberia. Cultivated in Europe and N.
ous places in Europe. Introduced to N. America. America. Natural hybrids with Lolium perenne*
a r e described as Festulolium loliaceum (Huds.)
CYNOSURUS CRISTATUS L. Crested dogtail, P . Fourn. (syn. Festuca loliacea Huds. ). Artifi-
Dog's tail g r a s s . 2n=14. P r i m a r y centre in C. cial amphiploids have been bred.
and W. Europe, Caucasia and Asia Minor.
FESTUCA RUBRA L. Red fescue. 2n=14, (28),
DACTYLIS GLOMERATA L. Orchard g r a s s . 42, 56, (70 and aneuploids). Europe, temperate
2n=28. Stebbins (1956) suggested that D. glome- Asia, Africa and N. America. Much cultivated as
rata is a tetraploid derived from two related d i - a pasture g r a s s . In New Zealand chewings fescue
ploids. One of them could be D. aschersoniana is cultivated. It is a red fescue of the n o n - c r e e -
Aschers. &Graebn. (2n=14). This species is d i s - ping type (spp. fallax).
tributed over C. Europe, Himalaya and W. China.
Another diploid is D. smithii Link which exists GLYCERIA FLUITANS R. Br. Mannagrass.
in the Canary Islands. It is likely that all diploids 2n=(20), 28, 40. Was collected in a large part of
derive from one common diploid. Hybridization E. Europe.
of diploids and doubling of the number of chromo-
somes and again hybridization within the tetraploid HOLCUS LANATUS L. Soft meadow g r a s s , Woolly
group and with the diploids has led to the very soft g r a s s , Yorkshire fog, Velvet g r a s s . 2n=14.
variable D. glomerata. Cultivated as a pasture Europe and temperate Asia. Cultivated for p a s -
and hay g r a s s . ture and hay.
DIGITARIA SANGUINALIS Scop. 2n=18, 28, 36 LOLIUM MULTIFLORUM Lam. var. westerwol-
(-48, 54, 76). Bluthirse, Millet sanguin. S. dicum Wittm. (syn. spp. multiflorum (Husnot)
Europe, Asia Minor, Central Asia, N. and S. Becherer). Westerwolds r y e g r a s s . 2n=14. Annual
America, in temperate zones. There is a great types derived from populations of spp. italicum
variability of the species. The cultivated type is were selected at Westerwold, NE. Netherlands.
var. esculenta (Gaudin) Caldesi. Among this
variety var. frumentacea Henr. and spp. aegyptia- LOLIUM PERENNE L. Perennial r y e g r a s s . 2n=14.
ca (Retz) Henr. a r e found. P r i m a r y centre is not Not known where and when it was domesticated,
known. Probably first cultivated in Illyria p r e c e e - but probably in Europe. However, the parent
ded by a long time of collection of wild plants. plants may have come from the Mediterranean
Cultivated formerly in a large area in Europe. region or SW. Asia. The first true grass sown in
Another area of cultivation is in India (p. 65). a pure, or relatively pure state. Cultivated now in
Whether the origin of cultivation independently the Old and New Worlds. Tetraploids and amphi-
arose here, or whether this cereal spread to India ploids with Festuca pratensis* are cultivated.
from Europe or the reverse is not known (Por- Natural hybrids between these two species are d e s -
t ê r e s , 1955a). Spp. pectiniformis Henr. o f E . cribed as Festulolium loliaceum (Huds. ) P . Fourn.
Europe, the Near East and NE. Africa. Not culti- Hybrids of L. perenne and L. multiflorum* have
vated. Spp. aegyptiaca has an 'eastern' origin but been called L. x hybridum Hausskn. These last
it is probably not in Egypt. From this subspecies two species a r e closely related.
the cultivated var. frumentacea is derived. Spp.
vulgaris (Schrander) Henr. is very variable and
PHALARIS ARUNDINACEA L. Red canary grass. glected. They a r e probably derivatives of the wheat
2n=14. 28. Most of Europe, W., N. a n d E . Asia. (T. antiquorum Heer) cultivated by the Swiss Lake
Cultivated in the Old and New Worlds. Dwellers in the Neolithicum. They are nearly e x -
tinct.
PHLEUM PRATENSE L. Timothy, Herd's grass.
2n=mostly 42, genome formula NNA.A.A A . TRITICUM AESTWUM (L. ). Thell. spp. spelta
Europe, N. Asia and N. Africa. An amphipfoid (L. )Thell. Spelt. 2n=42, genome formula AABBDD.
of P. alpinum L. (Alpine timothy, 2n=28) and P. Cultivated from the Belgian Ardennes to Switzer-
nodosum L. (syn. P . pratense var. nodosum (L.) land and to Schwaben, Germany and in Spain. F o r -
Richter) (2n=14, genome formula NN(?)). A t e t r a - merly the spelt area in Europe must have been
ploid type similar to this species was developed much larger running from Sweden to Spain and may
from the diploid Ph. nodosum after doubling the be up to Africa (p. 117). In Spain (Asturia) spelt is
number of chromosomes. Ph. pratense is culti- harvested in the same way as in Transcaucasia.
vated in Europe and N. America as a forage and It is remarkable that many German/Belgian spelts,
hay crop. the relic Swedish spelt (from Gotland) and one
from Africa c a r r y an Rf.. -gene (Zeven, 1971).
PHRAGMITES COMMUNIS Trinius. Reed g r a s s .
2n^(36), 48. (54, 84, 96). A cosmopolite grass ZEA MAYS L. Maize, 2n=20, Secondary centres
used for land reclamation and bank protection. in S. Europe and the Mediterranean region (p. 117)
Young sprouts a r e eaten, while the culms have in the European corn belt and the Atlantic and
many u s e s . Continental maize growing regions (Brandolini,
1970). Domesticated in C. America (p. 166). Flint
POA BULBOSA* maize -indurata Sturt. - is common in all these
areas.
POA PALUSTRIS L. Fowl blue g r a s s . 2n=28, (42). Siberian ecotypes are recognized by germination
Arctic zone of Europe, Asia and N. America. at 5-6°C, cold resistance of seedlings to 4-5°C,
Various varieties have been developed in Europe. rapid growth, earliness, high assimilation rate
and protogyny (Gerasenkov, 1968).
POA PRATENSIS L. Blue grass, Kentucky blue
g r a s s , Bird g r a s s . 2n=38-147. Europe, Asia,
N. Africa and northern N. America. The great
variation in chromosome number owing to auto-
ploidization has resulted in many species descrip-
tions, but they can be considered as synonyms.
Furthermore as apomixy of this species is not
constant, types with different chromosome num-
ber may be selected. So it was possible to select
plants similar to P . pratensis from P . trivialis*.
If this proves that P . pratensis derives from P.
trivialis then P . pratensis must have originated
in the Old World. Various varieties have been
bred in Europe and Canada (p. 176) and elsewhere.
POA TRIVIALIS L. Roughish meadow g r a s s .

2n=14, (28). Europe and S. Siberia. Not much cul-
tivated. It might be the parent species of P. p r a -
tensis*.
SPARTINA TOWNSENDII H. et J. Grooves. Cord

g r a s s . 2n=(62, 120, 122, 124), 126. Originated
in W. Europe after introduction of the American
S. alterniflora Lois. (2n=70) and amphiploidization
of this species with the European S. maritima
(Curt. ) Fern. (2n=56). Cultivated for soil r e c l a -
mation and as a stabilizer.
TRISETUM FLAVESCENS (L. ) Beauv. (syn. T.

pratense Bers. ). Yellow cat grass, Golden oat
g r a s s . 2n=24, 28. It probably derives from T.
sibiricum Rupr. (2n=14, 24). This species occurs
in Kamtschatka, Siberia. From here it spread
westwards.
TRITICUM AESTWUM (L.). Thell. spp. compac-

t u m H o s t . ) . Club wheat. 2n=42, genome formula
AABBDD. The club wheats of the Austrian alpines, Ribes acicularis
except for the research of E. Mayr, a r e much n e -
GRAMINEAE - LABIATAE 135
Grossulariaceae Labiatae
RIBES ACICULARIS Smith. 2n= . The moun- MENTHA CARDIACA Gerard ex Baker. Scotch
tains of Siberia especially in the Altai. The most mint, Scotch spearmint. 2n= . Temp. Europe.
precocious Ribes-species with a high winterhardi- Cultivated for its volatile oil. Closely related to
ness and mildew resistance. These c h a r a c t e r i s - M. x gentilis L. (2n=54, 60, 84, 96, 108, 120).
tics a r e useful in Ribes-breeding. It is believed that these two species a r e hybrids
of M. arvensis* and M. spicata*.
RIBES GROSSULARIA L. (syn. R. uva-crispa L.).
(European) Gooseberry. 2n=16. Eurasia and in MENTHA x GENTILIS L. (syn. M. sativa var.
the mountains of W. Asia and the Mediterranean gentilis (L.) Reichenb. ). 2n=54, 60, 84, 96, 108,
countries. Cultivated in temperate zones. Related 120. A hybrid of M. arvensis* and M. spicata*.
N. American Ribes-specles R. oxyacanthoides Usually sterile. Cultivated frequently.
Mill. (2n=16), R. hirtellum Mix. (2n=16), R. d i -
varicatum Dougl. (2n=16), R. c y n o s b a t i L . * MENTHA x PIPERITA L. Peppermint. 2n=(36,
(2n=16), R. pinetorum Greene (2n= )and 48, 64-69), 72, (84, 108, 122, 144). Probably a
R. niveum Lindl. (2n=16) carry resistance to m i l - natural hybrid of M. aquatica* and M. spicata*.
dew, while R. niveum and R. divaricatum may be This hybridization probably took place in England,
used as source of mildew resistance and to i m - f. piperita (Blackmint, black mitcham) is cultiva-
prove fruit characteristics. Resistance to Naso- ted in C. Europe and Great Britain, while f. p a l l e s -
nonia ribisnigri Mosley is found in R. roezlii cens Camus (white mint, white mitcham) is cul-
Regel (2n=16) and R. sanguineum Pursh (2n=16), tivated especially in France. In USA existing
while the latter species and R. cereum Dougl. clones were replaced bythe cultivar Mitcham in
(2n=16) a r e sources of resistance to Hyperomyzus 1890. This is still the main clone cultivated.
lactucae L. (Keep &Briggs, 1971).
MENTHA ROTUNDIFOLIA (L.)Huds. (syn. M.
RIBES NIGRUM L. (European) Black currant. spicata v a r . rotundifolia L.). Apple mint, Wooly
2n=16. Eurasia and sporadically in N. America. mint. 2n=24, genome formula RR. Europe and
The cultivated type was derived from the wild one. Canary islands. Cultivated. Probably the p a r e n -
In N. Scandinavia very precocious, winterhardy tal form of M. spicata* and one of the parents of
types a r e found. The American R. americanum M. japonica Mak., M. arvensis* and M. aquatica*
Mill. (2n= )and the Asiatic R. dikuscha Fish. (Ikeda &Ono, 1969). This species is related to
a r e related to the blackcurrant. They have b r e e - M. longifolia* and M. spicata*.
ding value.
Cultivars of v a r . sibiricum E. Wolf, of this s p e - MENTHA xSMITHIANA R.A. Graham (syn. M.
cies and R. ussuriense* are sources of resistance rubra Sm., non Miller). 2n=54, 120. Rarely cul-
to the blackcurrant gall mite, Phytoptus ribis Nal. tivated (Tutin et a l . , 1972). It is a hybrid of M.
aquatica* x M. arvensis* x M. spicata*. Usually
RIBES PETRAEUM Wulfen. Rock red currant. sterile, spreading vegetatively.
2n=16. The Pyrena to the Carpates and N. Africa.
Cultivated in the Alps. One of the parents of the MENTHA SPICATA (L.) Hudson (syn. M. viridis
present-day-redcurrant (R. sativum*). L. ). Spearmint, Green mint, Lamb mint. 2n=36,
48, genome formula RRSS (48+2B, 64). Temp.
RIBES SATIVUM Syme (R. rubrum L . , R. multi- Europe. It might derive from an autotetraploid
florum Kitt, and R. petraeum Wulf. ). 2n=16. The plant of M. rotundifolia* after which one genome
wild R. sativum grows in W. Europe. In N. A m e - pair RR changed into SS. Tutin et al. (1972)
rica it has run wild. R. rubrum is found wild in suggested that this species arose in cultivation as
W. and C. Europe and N. Asia. R. petraeum* a segmental allopolyploid of M. suaveolens (see
grows in the mountains of Europe and Asia. R. M. x rotundifolia*) and M. longifolia*. Var. c r i s -
sativum is probably the originally cultivated s p e - pata Schrader (syn. M. crispa L.)has genome
cies. Later it hybridized with the other two, so formula RRS c S fc . This species is one of the parents
these three species a r e the parents of the p r e s e n t - of M. x piperita*. It might be one of the parents
day redcurrant. of M. x villosa*.
Murray et al. (1972) artificially crossed M. aqua-
RIBES SPICATUM Robson. 2n=16. NE. Europe. tica* (2n=96) and M. spicata* (2n=48). This r e s u l -
Sometimes cultivated. ted in very variable F 1 due to the heterozygosity
of the pollen parent. Some hybrids resembled the
Juglandaceae natural strains of M. x piperita, others didnot.
JUGLANS REGIA L. Walnut. Persian walnut,
MENTHA SUAVEOLENS Ehrh. (syn. M. rotundi-
English walnut. 2n=32, 36. P r i m a r y centre of
folia auct., non (L.) Hudson). 2n=24. Cultivated
diversity in Centre 5 (p. 72). Secondary centre as a potherb.
in SW. Europe and Moldavia.
Almost all varieties in Germany are apomictic.
MENTHA x VILLOSA Hudson (syn. M. cordifolia
auct., M. gratissima Weber). 2n=36. Nm. alope-
curoides (syn. M. alopecuroides Hull, M. velutina
Lej. )was formerly much cultivated. This species
is a hybrid of M. spicata* and M. suaveolens*. LOTUS CORNICULATUS L. Birds-foot trefoil.

2n=12, 24. Europe, moderate Asia and N. Africa
NEPETA CATARIA L. (syn. Cataria vulgaris to Ethiopia. Formerly and at present in USA in
Moench. ). Catnip, Catmint. 2n=(32), 34, (36). use in seed mixtures for a ley crop and for p a s -
Europe. A perennial herb cultivated for medicinal t u r e s . Landolt (1970) and Somaroo &Grant (1971)
purposes. suggested that the diploid is a hybrid and the t e -
traploid an allotetraploid of L. alpinus Schleicher
ORIGANUM VULGARE L. Wild marjoram. 2n=30, (2n=12) of the Alp and the submediterranean L.
32. Europe, Siberia, Himalaya. Asia Minor and pilosus Jord. (2n=12).
Iran. Cultivated as a medicinal plant. The erect, broad-leaved type probably from C.
European origin is spread now as a contaminant
Leguminosae of grass seed for road sides throughout W. Europe
(Jones, 1973).
ANTHYLLIS VULNERARIA L. Kidney vetch,
Spring vetch, Lady's fingers, Wound-wort, Amer, LOTUS ULIGINOSUS Schkuhr. Greater birds-foot
Tare. 2n=12. Temperate Europe, Caucasia, Ante- trefoil. 2n=12, (24). Europe, N. Africa, Ante-
Asia, N. Africa and Ethiopia. Cultivated since Asia to Tibet. Cultivated in C. Europe and Great
1858 (Mansfeld, 1959) and is now usually mixed Britain as a fodder crop (Mansfeld, 1959).
with pasture g r a s s e s ,
MEDICAGO DENTICULATA Willd. 2n=16. China.
ASTRAGALUS CICER L. Milk vetch. 2n=64. Used as a vegetable. Related to M. sativa*.
Europe. A perennial pasture plant well-adapted
for grass mixtures (Whyte et a l . , 1953). MEDICAGO FALCATA (L. ) Doell. Yellow lucerne.
2n=16, 32. From W. Europe to E. China and E.
ASTRAGALUS FALCATUS Lam. Sicklepod milk of USSR, including the Mediterranean region. P r i -
vetch. 2n=16. W. Asia. A forage plant cultivated mary centre in USSR. There a r e two Siberian sub-
in USSR and France. species described: spp. ruthenica (L.) Ledeb.
(syn. M. ruthenica Trautv. ) and spp. platycarpos
ASTRAGALUS GLYCYPHYLLUS L. Milk vetch. (L. ) Ledeb. (syn. M. platycarpa Trautv. ).
2n=16. Europe and Siberia to Altai. A perennial
herb cultivated as a fodder.
CORONILLA VARIA L. Crown vetch. 2n=24. C.

and S. Europe extending to C. Russia. Cultivated
as an ornamental, a fodder crop and a cover crop.
GALEGA OFFICINALIS L. Galega, European 0%fë

goat's rue. 2n=16. E . , C. and S. Europe, Cauca-
sia, Asia Minor and Iran. Cultivated as a forage
crop and as an ornamental.
GLYCYRRHIZA ECHINATA L. 2n=16. SE. Europe

to Hungary and Italy. Cultivated to produce liquo-
rice.
Medicago falcata (Fischer, 1938)
GLYCYRRHIZA GLABRA (syn. G. glandulifera
Waldst. &Kit.). Common licorice, Liquorice.
2n=16. Europe and the Mediterranean region. A The hybrid described as M. medica P e r s . o r i g i -
perennial herb. Var. typica Regel &Herder is nates where the a r e a of M. sativa and M. falcata
cultivated to produce Spanish or Italian licorice overlap. In the same area M. hemicycla Grossh.
and var. glandulifera Waldst. Russian licorice. (2n=16, 32) is found. It may have a hybrid origin
(Lesin &Lesin, 1964). M. falcata and M. sativa*
HEDYSARUM HEDYSAROIDES (L. ) Schinz. & are closely related. Their karyotypes are similar
Thell. (syn. H. alpinum Jacq. ). 2n=14. S. Europe, (Gillies, 1970). Spp. falcata (L. )Arcangeli (syn.
Asia Minor, America and Caucasia. Cultivated as M. borealis Grossh. ) (2n=16, 32) is a wild perenni-
a fodder crop especially in the Alps. al type growing in meadows of the non-chernozem
zone of European USSR and W. Siberia. It is early
LATHYRUS SYLVESTRIS L. Flat pea, Wood pea. maturing requiring a long-day period. It has a
2n=14. Europe. Cultivated for forage and as an good practical value.
ornamental plant. M. romanica Prodon (2n=16) is a Russian Southern
Steppe type. Other species as M. tenderiensis
LATHYRUS TUBEROSUS L. Groundnut peavine, Opperm. and M. glandulosa David. (2n=32) are
Earth chestnut. 2n=14. Europe and W. Asia. Cul- variants of M. falcata.
tivated for its tubers. In the 16th Century its Wild diploid forms do not readily cross with cul-
flowers were distilled for perfumes (Uphof, 1968). tivated tetraploid types, hence these types may
not have introgressed (Lesin & Lesin, 1964).
LABIATAE - LEGUMINOSAE 137
M. glomerata* resembles hybrids of M. falcata x probably started in S. France resulting in spp.

M. prostrata Jacq. (2n=16, 32) and M. falcata x sativa. There a r e three subspecies: arenaria (Kit
M. sativa*. &Koch. ) Thellung, sand esparcette, montana
(Lam. &D. C. ), the mountain esparcette and s a t i -
MEDICAGO GLOMERATA Balbis. 2n=16. P r o b a - va (Lam. ) Thellung, the cultivated esparcette.
bly the Maritime Alps. It resembles hybrids of The last name is confusing, because spp. arenaria
M. falcata* x M. prostrata Jacq. (2n=16, 32) and (also described as var. transcaucasia, syn. O.
M. falcata x M. sativa*. It easily c r o s s e s with transcaucasia Grossh. )* is cultivated too.
the last-mentioned species being a source of d i -
sease resistance. SAROTHAMNUS SCOPARIUS (L. )Wimm. ex Koch.
Most individuals a r e not winterhardy. Broom. 2n=(14), 46, 48. W. and C. Europe. Cul-
tivated as a soil stabilizer.
MEDICAGO LUPULINA L. Hop clover, Black
medic. 2n=16, genome formula SS, 32, (64). TRIFOLIUM HYBRIDUM L. (syn. T. fistulosum
Europe, N. Africa and temp. Asia. Cultivated Gilib.). Alslike clover. 2n=16. Temperate Europe,
since 1659 in England and since 1785 in France. SW. Asia and N. Africa. Possibly first cultivated
Cultivated now in the Old and New World as a in Sweden. Introduced to other European countries
green fodder, hay crop and green manure. and N. America. Often found in fields of red clo-
Var. lupulina has been described as M. willde- ver. Very likely not the ancestral form of T. r e -
nowii Boenningh and M. stipularis Wallr. and var. pens*. The cultivated type spp. hybridum is p r o -
cupaniana Urb. as M. cupaniana Guss. and M. bably derived from the wild type spp. elegans
leiocarpa Guss. (Mansfeld, 1959). (Savi) Asch. &Graebn. In Anatolia spp. anatoli-
cum (Boiss. ) Hossain is found.
MEDICAGO SATIVA L. Lucerne. Blue alfalfa.
2n=(16, genome formula SS), 32, (48, genome TRIFOLIUM PANNONICUM Jacq. Hungarian clo-
formula SSSSSS, 64). Transcaucasia (p. 86). Two ver. 2n=c. 96, 98, c. 126, c. 130, c. 180, E . ,
populations - one from the Balkans and one from C. and S. Europe. Cultivated.
France - "met" in Thuringia, Germany. This r e -
sulted in a hybrid swarm from which winterhardy TRIFOLIUM PRATENSE L. Red clover. 2n=14,
types were introduced in Minnesota, USA in 1857. genome formula AA, (28). Europe, W. and C.
Asia and N. Africa. P r i m a r y centre in centre 9.
MELILOTUS ALBUS Medik. White sweet clover, It was probably first cultivated in the Netherlands,
Bokhara clover, Honey clover, White melilot. in the beginning of the 16th Century. Spread to
2n=16. Europe and W. Asia. Cultivated in the Old Germany and through Flanders to England. In the
World and particularly in the USA as a fodder beginning of the 17th Century seed of red clover
crop and green manure. was exported from the Netherlands to the Scandi-
It can be divided into two groups 1) the annual navian countries and France. From England red
wild type and 2) the bushy type. The latter might clover was spread to USSR and N. America. The
be a mutant of group 1, or derive from a natural wild type has more leaves and new shoots emerge
cross of M. albus. From both groups cultivars from internodes at the butt end, while the culti-
have been selected. vated type has less leaves and new shoots emerge
The very low variation of this species may point from the leaf rosette. The variable wild type is
to only a few introductions. described as var. pratense Bobr. and the cultiva-
ted as var. sativum (Crome) Bobr. (syn. T. s a t i -
MELILOTUS ALTISSIMUS Thuill. 2n=16. Europe vum (Sturm) Crome).
and temp. Asia. Sometimes cultivated for horse Late red clover (var. serotinum) may have deve-
fodder. loped from contaminants or spontaneous mutants
in USSR from introduced early types (var. p r a e -
MELILOTUS DENTATUS (Waldst. &Kit. ). P e r s . cox).
2n=16. E. and C. Europe to N. Sweden. Coumarin Autotetraploid types a r e widely cultivated now.
deficient and salt tolerant. Used to breed coumarin Var. americanum C O . Hartz was cultivated b e -
free cultivars of M. albus*. tween 1883 and c. 1910 in C. Europe. It originated
from a N. American introduction. It is often e r r o -
MELILOTUS MACRORHIZUS P e r s . 2n=16. Asia neously described as T. expansum Waldst. &Kit.
and Europe. Cultivated in China for its roots which Var. maritimum Zabel (var. villosum Wahlberg)
a r e eaten as a vegetable. Closely related to M. is found wild on the S. coast of the Balkan Peninsu-
altissimus*. la and var. frigidum Gaudin occurs wild in the
Alps.
MELILOTUS OFFICINALIS Lam. Biennal yellow
sweet clover, Field melilot, Yellow melilot. TRIFOLIUM REPENS L. White clover. 2n=32,
2n=16. W. Europe to W. China. Cultivated in (48, 64). Wild type (var. sylvestre). In meadows
Europe and also in the USA. It is a biennal with throughout Eurasia and N. Africa. Cultivation
sporadically some annuals. started probably in N. Italy (p. 102) and in the
Netherlands. Very variable.
ONOBRYCHIS VICIIFOLIA Scop. (syn. O. sativa Brewbaker and Keim (1953) suggest that T. n i -
s . 1 . Lam.). Esparcette. 2n=28. Temp. Europe, grescens Viv., Ball clover (2n=16) is one of the
SW. Asia to Altai and Transbajkal. Cultivation was parents. Chen and Gibson (1971) believe that it is
an autotetraploid while T. nigrescens and T. occi- L. crepitans Dumort. (2n=30), now included in
dentale D. Coombe (2n=16) a r e related to it. L. usitatissimum is probably the weedy type of
T. uniflorum L. (2n=32) might also be a parent. flax.
This species is found in E. Mediterranean region
to Sicilia. It includes T. savianum Guss. of Sici- Malvaceae
lia and Calabria. Italy. It is probably an auto-
tetraploid. ALTHAEA OFFICINALIS*
TRIFOLIUM RESUPINATUM L. (syn. T. suaveo- Paeoniaceae

lens Willd. ). Persian clover. 2n=(14), 16. The PAEONIA OFFICINALIS L. Peony, Piney. 2n=20.
Mediterranean region to Iran, Afghanistan and S. Europe, Asia Minor and Armenia. A perennial
India. Cultivated as a fodder crop. Var. majus herb cultivated for its medicinal m e r i t s .
Boiss. is syn. to T. suaveolens Willd.
Polygonaceae
TRIGONELLA COERULEA (L. ) Ser. Sweet trefoil.
RUMEXACETOSA L. Garden s o r r e l . 2n=14 9 ,
2n=16. Cultivated and also found as a weed or
ruderal. It may be derived from T. procumbens 15CT and other numbers. Temp. Europe and Asia.
(Besser) Reichenb. (syn. T. besserana Ser., T. A perennial herb. Var. hortensis Dierb. (syn.
coerulea spp. procumbens (Besser) Thell. ). This R. ambiguus Gren. ) is cultivated in the Old and
species is a native to EC. and SE. Europe. New Worlds.
RUMEX ALPINUS L. Alpine dock. Monk's r h u -

ULEX EUROPAEUS L. Common corse. 2n=96.
barb. 2n=20. C. Europe, the Balkans and Cauca-
W. Europe to Italy. Cultivated formerly for fodder,
sia. Cultivated formerly in C. Europe as a vege-
bedding and as hedges.
table
VICIA CRACCA L. Gerard vetch. 2n=12, 14, (21,
24), 28. W. Europe to Kamtchaska, E. China and RUMEX OBTUSIFOLIUS L. Broad-leaved dock,
Japan. Cultivated. Bitter dock. 2n=40, (60). Europe and temp. Asia.
Cultivated.
VICIA HIRSUTA (L. ) S. F . Gray. Common tare,
RUMEX PATEENTIA L. Patience dock, Spinach
Hairy t a r e . 2n=14. Europe, N. Africa and W.
dock, Herb patience. 2n=(40), 60. Probably C.
Asia. Cultivated in W. of USSR together with b a r -
Europe to W. Asia. Cultivated as a vegetable.
ley.
VICIA PANNONICA Crantz. Hungarian vetch. RUMEX SCUTATUS L. (R. alpestris Jacq. ).
2n=12. P r i m a r y centre in SW. Asia (p. 87). S e - French s o r r e l . 2n=20, (40). C. and S. Europe,
condary centre in Hungary. Alpine regions, Caucasia, India. Cultivated as a
vegetable (var. hortensis Lam. &DC. ).
Liliaceae
Portulacaceae
ASPARAGUS OFFICINALIS L. Garden asparagus.
PORTULACA OLERACEA L. Common purslane,
2n=20. P r i m a r y centre probably in the s a l t -
Pursley. 2n=(45), 54. Europe. Cultivated as a
steppes of E. Europe (Chittenden, 1951). A. offi-
vegetable (spp. sativa (Haw. ) Celak. ), but is often
cinalis var. prostratus Richter is a tetraploid
found as a weed (spp. oleracea). Now spread over
(Braak &Zeilinga, 1957).
a large part of the world.
CONVALLARIA MAJALIS L. Lily-of-the-valley.
2n=32, 36, 38. Europe, temp. Asia and Japan. Ranunculaceae
A perennial herb cultivated as a medicinal crop ACONITUM NAPELLUS L. Monkshood. 2n=24, 32.
and as an ornamental. C. Europe. Cultivated as a medicinal crop and
also as an ornamental.
Linaceae
LINUM USITATISSIMUM L. Flax, Linseed. 2n=30, AQUILEGIA VULGARIS*
(32). For origin see p. 87. Helbaek (1956) suppo-
sed two ways of introduction of flax. One through NIGELLA SATIVA*
Greece and the Donau valley into C. and W.
Europe and the other west of the Black Sea in a Resedaceae
northern direction into Russia. The first was p r o - RESEDA LUTEOLA*
bably a winter-annual which is the parent of
"Winterlein" cultivated in Germany. The other Rhamnaceae
was probably a summer-annual. In the first mille-
nium B. C. the latter was introduced to C. and W. RHAMNUS CATHARTICUS L. Buckthorn. 2n=24.
Europe. It is at present described as spp. eurasia- Europe up to Transcaucasia and W. Siberia, and in
ticum Vav. &Ell. In NW. USSR there is a centre Algeria. Formerly the fruits of this tree were
of fibre containing some of the finest fibre flax used as a source of yellow dye.
varieties.
In W. Europe and Ukraine the weed rattle flax,
LEGUMINOSAE - ROSACEAE 139
RHAMNUS FRANGULA L. Alder buckthorn. 2n=20, ARMENIACA BRIGANTTNA (Vizi. ) P e r s . (syn.

22, 26. Europe, Asia and N Africa. A t r e e for- Prunus brigantina Vill. ). Brianpon apricot.
merly cultivated. 2n= . Originated in SE. France. The seeds a r e
the source of the perfumed oil "Huile de Marmotte"
Rosaceae (Uphof, 1968). It might be a gene source of late
flowering.
AGRIMONIA ODORATA (Gouan) Mill. 2n=56. It is
included in A. eupatoria L., Agrimony. Cultivated
ARMENIACA SIBIRICA P e r s . (syn. Prunus sibi-
as a medicinal crop.
rica L., P . armeniaca var. sibirica K. Koch.).
Siberian apricot. 2n=16. Intern Mongolia to the
AMYGDALUS BESSERIANA Schott, (syn. A. nana
Sowjet F a r East and Lake Baikal. This species
L . , Prunus nana (L.) Stokes, P . tenella Batsch. ).
has the largest distribution of all apricot species
Dwarf almond, Dwarf Russian almond, Steppe
(Zylka, 1970). It is very cold resistant.
almond. 2n=16. P r i m a r y centre in E. Europe and
Siberia. Also wild in the Balkan, Asia Minor,
FRAGARIA XANANASSA Duch. (syn. F. grandi-
Caucasus and China (p. 36). It is the commonest
flora Ehrh. ). Pineapple strawberry. 2n=56. Arose
wild almond species and it is very frost resistant
spontaneously in W. Europe (possibly in Bretagne,
which makes it extremely valuable as a rootstock
France) after hybridization of F. virginiana*
of A. communis.
from N. America and F . chiloensis* from N. and
S. America and Hawaii. F. ovalis (Rydb. ) Lemm.
AMYGDALUS LEDEBOURIANA Schlecht, (syn. from NW. USA is used as a source of winterhardi-
Prunus ledebouriana Schlecht. ) 2n= . A shrub
from Tarbagatai and Altai.
FRAGARIA MOSCHATA Duch. Hautbois s t r a w -
AMYGDALUS PERSICA L. Peach. 2n=16. P r i m a -
berry. 2n=42. Europe and European USSR. Culti-
ry centre in China (p. 36). Secondary centre in
vated formerly, and run wild in other countries.
Moldavia, USSR.
* ^ j!f#{W
Fragaria moschata
Armerica sibirica
FRAGARIA VESCA L. Wild strawberry, Alpine world today. This shows the very polymorphic
strawberry. 2n-14. genome formula AA. Europe nature of this species which has also arisen due
and Asia. According to Darrow (1955) the var. to introgression with other species.
semperflorens Duch. is the parent form of the
cultivated strawberry. Its domestication occurred PRUNUS CERASUS L. Sour cherry. 2n=32, g e -
in the north of the Italian Alps. nome formula CC. C. Asian centre (p. 90). The
population Vladimirskaya vishnia with large dark-
FRAGARIA VIRIDIS Duch. Polunitsa. 2n=14. claret fruits that a r e very palatable and aromatic,
European part of centre 9. Cultivated formerly. originated in Centre 9, extending westward and
southward to the Rhine and Balkans.
MALUS BACCATA (L. ) Borkh. var. baccata.
Siberian crab apple. 2n=34. Wild in Transbaikal PRUNUS DOMESTICA L. Garden plum, Domestic
and Ante-Baikal t e r r i t o r i e s . P r i m a r y gene centre plum. 2n=48, genome formula CCSSSS or CdCd
in Siberia. Resistant to frost. SSSjSj or CdCdD 1 D 1 D 2 D 2 . For origin see p. 90.
Werneck (1958) considered Upper Austria as a
place where the garden plum has arisen. Bush
seedling would have been transplanted to com-
pounds where further domestication may have
occurred. The Lake Bank Dwellers of neolithic
Switzerland knew the garden plum.
PRUNUS FRUTICOSA Pall. (syn. Cerasus fruti-

c o s a P a l l , ) . Dwarf Cherry, Bush Cherry, Ground
Cherry, Mongolian Cherry, Steppe Cherry.
2n=16, 32. Extended over Europe. It occurs in
great diversity beyond the Volga, in S. Ural, SW.
Siberia and the Bashkir Republic. One of the
parents of P . cerasus*. It withstands -52°C.
PRUNUS INSITITIA L. (syn. P . domestica var.

insititia (L. ) C . K . Schneider, P . domestica spp.
italica (Borkh. ) Hegi). Bullace plum. Damson
plum. 2n=48. S. and SE. Europe and adjacent
parts of Asia. Occurs now throughout temp.
Europe and W. Asia. Probably only known as a
cultigen and naturalized. If so, it is obviously an
allohexaploid. It is frost resistant.
PRUNUS MAHALEB L. Mahaleb cherry, St. Lucie

cherry. 2n=16. C. and S. Europe and W. Asia.
Fruits a r e not edible. Used as rootstock for c u l -
tivated c h e r r i e s .
Malus baccata PYRACANTHA COCCINEA M . J . Roemer. 2n=34.
S. Europe and westwards to NE. Spain. Cultivated
as an ornamental and for its fruits.
MALUS PRUNIFOLIA (Willd.) Borkh. (syn. Pyrus
prunifolia (Willd. ). Chinese crab apple. 2n=34, PYRUS COMMUNIS L. (syn. P . domestica Med.).
(51, 68). Wild and cultivated in the extreme e a s - Common pear. 2n=34, (51). Europe and W. Asia.
tern sector of Centre 9. P r i m a r y centre China. It has been divided into spp. pyraster L. (syn.
P . pyraster Burgsd., P . communis var. achras
MALUS PUMILA Mill. (syn. Pyrus malus L.) Wallr. ), spp. nivalis Jacq. (syn. P. nivalis Jacq. )
Apple. 2n=34, 51, 68. The Balkans and SW. USSR and spp. salvifolia (syn. P . salvifolia DC.). Spp.
(p. 75), eastwards through Transcaucasia, Iran, pyraster is the most important one, it grows in
Turkestan, and northward to the Altai mountains. C. Europe and W. Asia. Spp. nivalis, the snow
It occurs along the ancient and mediaeval routes pear grows in W. Switzerland and France. It is
of commerce and migration between Europe and used as a rootstock. Spp. salvifolia is found in the
E. Asia. Man has greatly promoted its distribution same a r e a s . The cultivars a r e derived from these
(Wilcox, 1962). It is considered as the principal subspecies by selection and by crossing with P.
ancestor of the cultivated apple. M. sylvestris* serotina*, P . ussuriensis*, P . longipes, P. eau-
hybridizes with this species and hence may also casica*, P. amygdaliformis* and P. salicifolia*.
have played a small part as an ancestor. Pavlov (1969a, 1969b) reported that types of W.
European USSR is the primary centre for many and S. Europe derive from crosses with P . niva-
old cultlvars as Antonovka, Aport, Borovinka. In lis* and P. amygdaliformis*. because they have
early 19th Century, Bolotov, described 600 Russi- hairiness and a high number of stomata per area
an cultivars; about 10 000 cultivars exist in the like these two species.
Some cultivars in Caucasus show characteristics
ROSACEAE - SALICACEAE 141
obviously derived from P. caucasia*. The E. Rubus species have also been used in breeding
European cultivars show a direct derivation from work.
the wild P. communis.
RUBUS LACINIATUS Willd. Evergreen blackberry.
PYRUS CORDATA Desv. 2n= . W. Europe. 2n=28. C. Europe. A cultivar was brought to N.
Cultivated in hedges and for its wood. America where hybridization took place with
another European immigrant, R. procerus P . J .
ROSA x ALBA L. French rose. 2n=28, 42. Culti- Muell. (2n=14, 28, 49), Himalaya berry.
vated in Bulgaria and S. France for the perfumery
industry. Probably a hybrid of R. arvensis* x RUBUS SAXATILIS L. Stoneberry. 2n=28. Europe
R. gallica*, and a white flowered member of the and N. Asia. In Sweden a species has been found
Sect. Canina. to be resistant to rust and other diseases. The
fruit has only a few drupelets and lacks flavour
ROSA ARVENSIS Hudson. 2n=14. S., W. and C. (Larson, 1969).
Europe. It is one of the parents of R. x alba*.
SANGUISORBA MINOR Scop. 2n=28. (54, 56).
ROSA x BIFERA (Poiret) P e r s . (syn. R. d a m a s - Europe and temp. Asia. Sometimes cultivated to
cena auct., non Miller). Damask rose. 2n=28. flavour soup or for salads (Mansfeld, 1959).
Probably a hybrid of R. moschata* and R. gallica*.
Cultivated in Bulgaria, S. France and Turkey. SANGUISORBA OFFICINALIS L. Great burnet,
The petals are used to produce oil of roses which Garden burnet. 2n=28, (42. 56, c. 70). Europe,
is used in perfumery. Asia and N. America. Sometimes cultivated as a
vegetable (Mansfeld, 1959).
ROSA CANINA L. Brier, Dog rose, Doghip.
2n=35. Europe, temperate Asia and N. Africa. It SORBUS AUCUPARIA L. (syn. Mespilus aucuparia
is a common rootstock of garden r o s e s . The All. ). Rowan tree, European mountain ash. 2n=34.
named selections a r e often less prickly than the The "Mährische Eberesche" (var. moravica) was
wild ones. found in 1810 in Czechoslovakia. It has been i m -
proved and distributed. Before its domestication
ROSA GALLICA L. French rose. 2n=28. S. and var. rossica and var. rossica-major were already
C. Europe up to Belgium and C. France and W. cultivated in USSR. It is an important source of
Asia. Probably a parent of R. x bifera* and Vitamin C (Mueller-Stoll &Michael, 1949).
R. xalba*. The petals a r e used in perfumery.
SORBUS DOMESTICA L. Service tree, Mountain
ROSA RUBIGINOSA L. (syn. R. eglanteria auct. ). ash. 2n=34. S. Europe, N. Africa and W. Asia.
Sweet b r i a r . 2n=35. W. and C. Europe. Cultivated Cultivated in Europe for its fruits which are eaten
for its flowers and as rootstock. or made into wine and as an ornamental. Large-
fruited forms a r e found in forests in Crimea
ROSA VILLOSA L. Apple rose. 2n=28. Var. p o - (p. 89).
mifera (Herrm. ) Crép. Europe and SW. Asia.
Rubiaceae
RUBUS ARCTICUS L. Arctic bramble, Nectar-
RUBIA TINCTORUM L. Madder. 2n=44. S. Europe
berry. 2n=14. Europe and N. Asia. Used in b r e e -
and Asia Minor. Cultivated in Europe as a dye
ding work with R. idaeus*. Fruits have a distinct
aroma and rich in Vit. C. A hardy, high yielding, plant.
disease resistant plant.
Salicaceae
RUBUS CHAMAEMORUS L. Cloudberry, Yellow- SALIX ACUTIFOLIA L. Caspic willow. 2n=38. A
berry, Salmonberry. 2n=56. Europe and N. Asia t r e e of USSR and Manchuria. Cultivated for twig
used in breeding with R. idaeus*. Easily domes- production.
ticated (Larson, 1969).
SALIX ALBA L. (syn. S. aurea Salisb. ). White
RUBUS IDAEUS L. European red raspberry. willow. 2n=76. In large a r e a of Europe and Asia
2n=14. Spp. vulgatus wild in Europe. It was do- (p. 75) and N. Africa. Introduced into N. America.
mesticated. The present cultivars often a r e hy- Cultivated in Europe for twig production for dike
brids of this subspecies and its NE. American building.
counterpart spp. strigosus.
The tetraploid subspecies, melanolasis Focke SALIX CAPREA L. Goat willow, Common willow.
from NW. America and Siberia, sachalinensis 2n=38, (57, 76). Europe and N. Asia. Cultivated
Léveillé from Sakhalin and sibiricus from Kam- for its twigs.
chatka, have been grouped as R. sachalinensis
Levi. Some cultivars derive from R. idaeus x SALIX FRAGILIS L. Brittle willow, Crack willow.
R. chamaemorus*, cloudberry (2n=56). Crosses 2n=(38), 76, (114). Europe, Asia Minor, Syria,
have also been made between this species and Iran and W. and C. Siberia. Often planted for
R. xanthocarpus Bur. &French from W. China. twig production. It is one of the parents of S. x
R. arcticus* L., Arctic bramble (2n=14)andP. rubens Schrank.
parviflorus L . , Japanese raspberry (2n=14). Other
SALIX PURPUREA L. Purple willow, Purple SOLANUM TUBEROSUM L. Potato. 2n=48. Domes-
osier willow. 2n=38. A large part of Europe, and ticated in S. America. In Europe spp. tuberosum
in Asia to Japan, and in N. Africa. Cultivated for developed. Its genetic basis is very small. This
twig production for dike works and basketry. is probably caused by only a few introductions,
and afterwards by the selection for short-day
SALIX TRIANDRA L. (syn. S. amygdalina L. ). forms and by mass killing during blight epidemics
French willow, Almond-leaved willow. 2n=38, in the 1840's.
44, (57, 88). Spread from W. Europe to E. Asia.
Planted for twig production. One of the parental Umbelliferae
species of the cultivated S. x mollissima Ehrh.
(2n=38). ANGELICA ARCHANGELICA L. Angelica. 2n=22.
Temperate Europe. Himalaya, Siberia and Kam-
tschatka. Cultivated for its aromatic petioles.
SALIX VIMINALIS L. (syn. S. longifolia Lam.).
Spp. archangelica includes the cultivated type.
Twiggy willow, Common osier. Basket willow,
Osier willow. 2n=38. C. Europe and a large part
ANTHRISCUS CEREFOLIUM (Waldst. &Kit.)
of Asia. Much cultivated in N. and S. Europe and
Sprengel. Garden chervil. 2n=18. Probably EC.
elsewhere for twig production. Many of the willows
and SE. Europe. Var. cerefolium has glabrous
planted in the Netherlands for dike work belong to
fruits. It includes the cultivated type.
this species and to S. triandra*. They a r e often
cultivated in mixed stands which leads to cross
BUNIUM BULBOCASTANUM L. (syn. Ligusticum
fertilization and development of hybrids.
S. dasyclados Wimmer (2n=38, 57, 76, 114) is bulbocastanum Crantz). 2n= . W. Europe.
probably a complex hybrid of S. caprea x S. cine- Formerly cultivated for its edible tubers.
r a x S. viminalis. S. helix L. is a hybrid of S.
purpurea* x S. viminalis. CHAEROPHYLLUM BULBOSUM L. Turnip-rooted
chervil. 2n=22. Europe and W. Asia. Its cultiva-
tion as a vegetable is on the decline.
Sambucaceae
SAMBUCUS NIGRA L. European elder. 2n=36. DAUCUS CAROTA L. White carrot, Orange carrot.
Europe. Cultivated. Recently there has been a 2n=18. Afghanistan (p. 76). The origin of the
new interest in this t r e e because of the p r o c e s - white type is not clear. It probably arose as a
sing of alcohol-free beverage (Strauss &Novak, mutant from a yellow type most likely in France.
1971). The orange carrot probably originated in the
Netherlands. This type of carrot is now cultivated
Saxiphragaceae widely by peoples of European stock. It has sup-
p r e s s e d the growth of the purple carrot which
BERGENIA CRASSIFOLIA (L. ) Fritsch. 2n=34.
colours soups and food preparations purple (Banga,
Siberia, Altai and N. Mongolia. A perennial herb
1957, 1962).
cultivated since 1927 in USSR as a tea plant
(Mansfeld, 1959).
LEVISTICUM OFFICINALE Koch. (syn. Angelica
levisticum Ball. ). Garden lovage, Bladderseed.
Scrophulariaceae
2n=22. Cultivated mainly for flavouring.
DIGITALIS LANATAEhrh. 2n=56. SE. Europe.
Elsewhere in Europe it may have run wild. Cul- MYRRHIS ODORATA (L. ) Scop. Garden myrrh.
tivated as a medicinal crop. Sweet scented myrrh. 2n=22. Europe and Caucasia.
Cultivated for flavouring and for fodder.
DIGITALIS PURPUREA L. Purple fox-glove.
2n=56. S. (p. 105) and C. Europe. Cultivated as a PASTINACA SATWA L. Parsnip. 2n=22. Europe.
medicinal plant and as an ornamental. Var. sativa is cultivated there and elsewhere for
its sweet, fresh tap-root. The wild type has a
VERBASCUM THAPSIFORME Schrad. 2n=32. sour root.
Spread throughout Europe. Cultivated for its medi-
cinal properties and as an ornamental. PEUCEDANUM CERVARIA (L.) Lapeyr. Hart's
wort, Much-good, Broad-leaved spignel. 2n=22.
Solanaceae S. and C. Europe. Cultivated formerly as a medi-
cinal.
CAPSICUM ANNUUM L. Bell pepper, Paprika,
Cayenne pepper. 2n=24. Mexico (p. 171). Seconda-
ry centre in Europe. PEUCEDANUM OSTRUTHIUM (L. ) Koch. Master
wort, Pellitory of Spain, Hogfennel. 2n=22.
Europe. Cultivated for its scenting root since the
PHYSALIS ALKEKENGI L. Strawberry tomato,
16th century, as a medicinal and as herb. Its cul-
Winter cherry. 2n=24. C. and S. Europe. A p e -
tivation has almost disappeared now.
rennial herb cultivated for its fruits.
SCOPOLIA CARNIOLICA Jacq. Scopalia. 2n=46- Valerianaceae

48, 48. Europe. Cultivated as a medicinal crop. VALERIANA COLLINA Wallr. 2n=28. Europe.
Formerly cultivated in Germany.
SALICACEAE - VIOLACEAE 143
VALERIANA EXALTATA Mikan.f. ex Pohl.

2n=14, (28). Europe. Cultivated in Germany. This
species might be included in V. officinalis*.
VALERIANA LOCUSTA (L.). Betcke. (syn. V.

olitoria Pollich). Corn salad, Lamb's lettuce.
2n= . Europe, N. Africa, Caucasia. Cultiva-
ted to be used for salads.
VALERIANA OFFICINALIS L. Valerian. 2n=14,

28, (56). Europe and temperate Asia. In Germany
and Poland the three ploidy levels a r e found. In
Poland these ploidy types do not hybridize, but in
Germany hybrids a r e often found. In the Nether-
lands, Belgium, France and Great Britain diploids
a r e not observed, while in USSR octaploids a r e
not seen. These ploidy types and hybrids have led
to many synonyms (Schrantz, 1961).
VALERIANA PROCUMBENS Wallr. 2n=56. Spain,

Great Britain, France and Germany. Cultivated
in Germany. This species might be included in V.
officinalis*.
VALERIANA SAMBUCIFOLIA Mikan.f. ex Pohl.

2n=56. Wild in N., C. and E. Europe. Cultivated
in Thuringia, Germany for its seeds. This s p e -
cies might be included in V. officinalis*.
Violaceae
VIOLA TRICOLOR L. 2n=26. Europe, Siberia up
to Altai and India. Spp. arvensis Gaud. (V. a r -
vensis Murr., 2n=34) is a cosmopolitan weed.
This subspecies and spp. tricolor (2n=26) are cul-
tivated for their medicinal and ornamental purpo-
ses.
10 South American
Centre
The South American Centre was first restricted to the Andes and described by Vavilov (1949/50) as
the Andean Centre of Origin. Vavilov divided it in two a r e a s , 1. Peru, Ecuador and Bolivia, and 2. a
small one on the island of Chiloe, Chile. Darlington & Janaki Ammal (1945) added the zone between the
coast of E. Venezuela and Guyana-Surinam-Cayenne and that of S. Brazil-Paraguay calling it the B r a z i l -
Paraguay Centre of Origin. Zhukovsky (1968) created the megacentre of S. America, while Harlan (1971)
described a large part of S. America as a noncentre C2 South American noncentre.
Agriculture was introduced from C. America before 6 000 BC because remains of domesticated
Phaseolus vulgaris found in the Guitarrero Cave in Peru have been dated about 6 000 BC. (Kaplan et a l . ,
1973).
A number of tuberous crops (Oxalis tuberosa, Solanum, UUucus tuberosus) was domesticated in this
centre. For tuberous Solanum triploid upto hexaploid species were developed. Other examples a r e fruit
t r e e s , Amaranthus s p . , Ananas comosus, Arachis hypogaea, Capsicum s p . , Cinchona ledgeriana, Cucur-
bita maxima, Gossypium s p . , Hevea s p . , Lupinus s p . , Lycopersicon s p . , Manihot esculenta, Nicotiana
s p . , Phaseolus s p . , Solanum s p . , Theobroma cacao etc. Only one cereal Bromus mango was developed
in this region.
A secondary centre of diversity arose for Zea mays.
ACANTHACEAE - AQUIFOLIACEAE 145
Acanthaceae hybrids easily a r i s e : A. braunii Thell. and A.

caracasamus H.B.K. In general A. spinosus is
JUSTICIA PECTORALIS Jacq. 2n= . West
the female parent. Grant (1959) supposed that A.
Indies and trop. America. Var. stenophylla Leo- spinosus is one of the parents of A. dubius, but
nard semi-cultivated in E. Colombia to adjacent Pal (1972) does not support this.
Amazonian Brazil. It is a smaller plant, has
smaller and longer leaves, and has shorter in-
florescences than the common type. Anacardiaceae
ANACARDIUM OCCIDENTALE L. Cashew. 2n=42.
Agavaceae Trop. America from Mexico to Peru and Brazil
and also the West Indies. Cultivated now in many
FURCRAEA GIGANTEA Venth. Piteira. Piteira
tropical countries which may form secondary cen-
gigante. 2n=(18), 60. S. and C. America. The
t r e s of diversity. Thus Northwood (1966) showed
Mauritius hemp comes from var. willemettiana
the great variation in yield and nut size in the
Roem. which is cultivated on Mauritius and e l s e -
cashew populations in Tanzania.
where.
SCHINUS MOLLE L. Californian peper tree, B r a -
FURCRAEA MACROPHYLLA (Hook. ) Baker.
zil pepper t r e e . 2n=28, 30. Mexico to Chile and
Fique. 2n= . Colombia. Cultivated there on a
up to Uruguay. Cultivated in the tropics as a medi-
small scale. Some varieties have developed.
cinal plant, as a shade tree and as an ornamental.
Fique fibre also comes from other Furcraea s p e -
cies such as F. andina Trel. (2n=60), a wild
SPONDIAS MOMBIM L. (syn. S. lutea L. ). Yellow
growing species from Equador and Peru, and T.
mombim, Jobo, Hog plum. 2n=32. Trop. America.
humboldtiana T r e l . , a wild growing species from
A fruit t r e e now cultivated in the tropics.
Venezuela.
Aizoaceae SPONDIAS PURPUREA L. (syn. S. mombim L. ).

Red mombim, Spanish plum. 2n= . Trop.
MESEMBRYANTHEMUM CHILENSE Mol. (syn. America, C. America and Mexico. A small fruit
Carpobrotus chilensis ( M o l . ) N . E . Brown). Sea tree.
fig. 2n= . Chile. A shrub used in N. America
to stabilize dunes. Annonaceae
ANNONA CHERIMOIA Mill. Cherimoya. 2n=14, 16.
Amaranthaceae
Wild in the Andean valleys of Ecuador and Peru.
AMARANTHUS CAUDATUS L. Inca wheat, Qui- There is its primary centre. A small t r e e . Culti-
huicha. 2n=32, 34. S. America, Asia and possibly vated now in the tropics. Its karyotype is similar
in Africa. Cultivated as a grain crop. In Andean to that of A. reticulata* and A. squamosa*. Several
region of Peru, Bolivia and NE. Argentina and in cultivars a r e known. Antemoya is a hybrid with
China, India, Nepal and Afghanistan. Its leaves A. squamosa*.
a r e also eaten. A form with red flower-spikes
used as a garden ornamental ('tove-lies-bleeding') ROLLINIA DELICIOSA Safford. 2n= Brazil.
should not be confused with quinoa (Chenopodium A t r e e cultivated for its fruits.
quinoa*. Quinoa is of S. American origin (Sauer,
1950). It resembles A. edulis* and the wild S. ROLLINIA LONGIFOLIA St.-Hil. (syn. R. dola-
American A. quitensis H. B.K., 2n=32. bripetala (Reddi) St. Hil.). 2n= . Brazil. A
tree cultivated for its fruits.
AMARANTHUS DUBTUS Mart, ex Thell. 2n=64.
Trop. America. Cultivated there as a potherb and ROLLINIOPSIS DISCRETA Safford. 2n=
for its grains. It also is a common weed. It r e s e m - Brazil. A shrub cultivated for its fruits.
bles A. cruentus*. Perhaps it is a tetraploid from
this latter species. See for hybridization with A. Apocynaceae
spinosus (p. 145).
PLUMERIA ACUTIFOLIA Poir. (syn. P. acumi-
nata Roxb., P. obtusa Lour. ). 2n=36. Mexico and
AMARANTHUS HYBRIDUS*
S. America. Cultivated in the tropics as a medi-
cinal t r e e .
AMARANTHUS MANTEGAZZIANUS P a s s e r (syn.
A. edulis Spegazzini). 2n=32. Cultivated in Argen-
THEVETIA NEREIFOLIA J u s s . (syn. T. peruviana
tina. The wild S. American A. quitensis H.B.K.
Schum.). Exile tree, Yellow oleander. 2n=20, 22.
(2n=32) closely resembles it. It is also included in Trop. America and W. Indies. A shrub cultivated
A. caudata* as ssp. mantegazzianus (Passer) in the tropics as a medicinal plant.
Hanelt.
Aquifoliaceae
AMARANTHUS SPINOSUS L. Thorny pigweed.
2n=34. S. and C. America. Widespread tropical ILEX PARAGUENSIS D. Don. (syn. I. paragua-
noxious weed. Cultivated as a vegetable (Mans- riensis St. -Hil. ). Paraguay tea, Yerba maté.
feld, 1959) in Singapore. Because of the spines it 2n=40. S. America. Cultivated for its leaves which
is unlike any of the grain amaranths (Sauer, 1950). are used to prepare tea.
Where it grows together with A. dubius* sterile
SOUTH AMERICAN CENTRE 146
Bombacaceae
QUARARIBEA CORDATA (H, &B. ) Garcia-
Barriga &Hernandez (syn. Matisia cordata H. &
B. ). South American sapote. 2n= . NW. of S.
America. Within this region this fruit is cultiva-
ted. No superior strains have been developed yet.
Bromeliaceae
ANANAS COMOSUS (L. ) Merr. (syn. A. sativus
Schult, f., Bromelia comosa L. ). Pineapple.
2n=50, (75, 100). It is suggested that the Tupi-
Guarani domesticated pineapple in the P a r a n a -
D i s p e r s i o n ( ) and cultivation of Hex p a r a g u e n s i s ( ) Paraguay river drainage area and that from this
(Patifio, 1968)
region pineapple was spread to all (sub)tropics.
However, Brücher (1971) suggested that the do-
mestication of pineapple might have taken place
Araceae in the highlands of Guyana and alongside the r i v e r s
XANTHOSOMA BELOPHYLLUM (Willd.) Kunth. there. In the first area wild related species A.
2n= . Cultivated for its roots in Venezuela. bracteatus (Lindl. ) Schultes, (2n= ), A. ananas-
soides (Bak.) L . B . Smith, (2n= ), A. e r e c t i -
XANTHOSOMA BRASILIENSE (Desf. ). Engl. folius L. B. Smith, (2n= ) and Pseudananas
Yautia, Belembe, Calalou. 2n= . Cultivated sagenarius (Arudda) Camarcq. (2n= ) occur.
from S. Brazil to the West Indies and Panama. A. bracteatus var. typicus is occasionally culti-
The leaves a r e cooked and eaten. vated for its fruits, while A. ananassoides var.
nanus is an ornamental.
XANTHOSOMA CARACU C. Koch &Bouché. C a r a -
cu. 2n= . Cultivated throughout the American ANANAS PARGUAZENSIS Card. -Cam. &Smith.
tropics for its corms and young leaves. 2n= . This species occurs where the Rio P a r -
guazo discharges into the Rio Orinoco, Venezuela.
XANTHOSOMA JACQUINE Schott. Yautia Palma. Brtieher (1971) suggested that primitive fibre and
2n= . Cultivated in trop. America. fruit cultivars have been selected. This selection
work could have been carried out - independently
XANTHOSOMA MAFAFFA Schott. Mafaffa, R a s c a - of each other - in the region Guyana-Orinoco, and
dera, Tärtago, Yautia. 2n= . Probably from between Maranhao and Pernambuco.
S. Brazil. Cultivated now in several areas of
Brazil. PSEUDANANAS MACRODONTES (Harms) Morr.
2n=c. 100. Argentine and Brazil. There its p r i -
XANTHOSOMA SAGITTIFOLIUM (L. ) Schott. mary centre is found. Cultivated on a large scale
Yellow yautia. 2n=24, 26. S. America. There it on Polynesian and Melanesian islands.
was cultivated for its roots and leaves. Now much
cultivated throughout the tropics. Some cultivars Cactaceae
have been developed for their starchy corms, PERESKIA ACULEATA Mill. Barbados cherry,
others for their leaves. Sweet Mary, West Indian goose-berry, Lemon
vine. 2n=22. Trop. America. Cultivated for its
XANTHOSOMA VIOLACEUM Schott. P r i m r o s e fruits.
Malanga, India Kale. 2n=24, 26. S. America.
Roots and leaves a r e eaten. TRICHOCEREUS PACHANOI Britton &Rose.
2n= . Andean parts of Ecuador and Peru.
Basellaceae Apparently widely cultivated throughout the C.
BOUSSINGAULTIA CORDIFOLIA Ten. (syn. B. Andes (Schultes and Hofmann, 1973).
baselloides H. B.K. ). Madeira vine, Mignonette
vine. 2n=c. 20, 36. S. and C. America. Cultivated Cannaceae
as a leafy vegetable or for its tubers. CANNA EDULIS Ker. Achira, Queensland a r r o w -
root. 2n=18, (27). Probably NW. of S. America.
ULLUCUS TUBEROSUS Caldas. Ulluca. 2n=24, Spread to Mexico, C. America, West Indies and
36. Unknown wild. A very ancient crop cultivated the northern of S. America. At present achira is
in C. Andes. The starchy tubers are curved in cultivated in W. Indies, Australia, S. America,
shape. It is very frost-resistant. parts of Asia and Pacific Islands. Remains of
achira have been found at Huaca Prieta, N. Peru
Bixaceae (Bird, 1948). They have been dated c. 2 400 BC.
BIXA ORELLANA L. Annato. 2n=14, 16. Trop. It could not be established whether they had been
America and the West Indies. Introduced into many collected or cultivated.
other tropical countries where it may have run Mukherjee and Khoshoo (1971) suggested that
wild. It is a dye crop. the triploid (2n=3x=27) is probably an intervarietal
hybrid involving rather genetically related varieties.
ARACEAE - DIOSCOREACEAE 147
It is highly vigorous and robust and has large r h i - SPILANTHES OLERACEA L. (syn. S. acmella
zomes. M u r r . ) . P a r a c r e s s , Brazilian c r e s s . 2n=14, 24,
In S. America rhizomes of other Canna s p e - 52. Brazil, W. Indies and also India. Cultivated
cies (C. coccinea Mill., C. paniculata R. &C. as a vegetable or salad.
and C. indica L. )have been collected and eaten
(Gade, 1966). TAGETES MINUTA L. Marigold. 2n= . Trop.
America. Spread to many other countries. Culti-
Caricaceae vated for its medicinal properties (Neher, 1968).
It may reach a height of 3 m or more when cared
CARICA CANDAMARCENSIS Hook.f. (syn. C.
for.
pubescens Lenne &Koch). Mountain papaya.
2n= . The Andes of Colombia and Ecuador. A
t r e e cultivated there and also in E. Africa for its Convolvulaceae
fruits (Mansfeld, 1959). IPOMOEA TILIACEA (Willd. ) Choisy (syn. I.
fastigiata (Roxb. ) Sweet). 2n=30. 60. S. America
CARICA CHRYSOPETALA Heilb. 2n= . Equa- and the West Indies. It has been claimed that it
dor. A t r e e cultivated for its fruits (Mansfeld, was already cultivated in West Indies in pre-Inca
1959). Badilla (1967) suggested that this species times (Uphof, 1968), It has been named as one of
is a natural hybrid product of C. candamarcensis* the parents of I. batatas*.
and C. stipulata Badilla from Ecuador. He further
suggested that this species, C. pentagona* and MERREMIA MACROCARPA (L. ) Roberty. 2n-
C. frutifragans Garcia &Hernandez another hy- Brazil and W. Indies. Cultivated for its medicinal
brid of the same parents (from Colombia) should tubers.
be grouped in C. x heilbornii Badilla.
MERREMIA TUBEROSA (L. ) Rendle (syn. Ipomoea
CARICA PENTAGONA Heilb. 2n= . Equador. tuberosa L., Convolvulus sinuata Ort. ). 2n=30.
A t r e e cultivated for its fruits (Mansfeld, 1959). Brazil, W. Indies, trop. Africa and India. Origin is
Badilla (1967) suggested that this species is a unknown. Cultivated as a medicinal and also as an
natural hybrid product of C. candamarcencis* and ornamental. It may have spread from West Indies
C. stipulata Badilla from Equador. and Brazil because in these areas M. macrocarpa*
grows wild and is cultivated.
Caryocaraceae
CARYOCAR NUCIFERUM L. 2n= Brazil and Cruciferae
Guiana. A tall t r e e cultivated in the W. Indies for LEPIDIUM MEYENII Walp. Maca. 2n= Peru
its edible Suari nuts. and Bolivia. Cultivated in Peru for its root.
CHENOPODIUM PALLIDICAULE Aellen. Canihua. CUCURBITA MAXIMA Duch. ex Lam. Pumpkin,
2n=36. Andes. Cultivated on the Altiplano of Peru Winter squash. 2n=40. Cultivated all over the
and Bolivia as a marginal grain crop (Dale, 1970). world. Secondary gene centre in India and adjacent
areas (p. 64). Whitaker (1962) suggested a com-
CHENOPODIUM QUINOA Willd. Quinoa. 2n=36. mon origin for C. maxima, C. ficifolia*, C.
Cultivated in the Andes as a grain crop. Culti- moschata* and possibly C. pepo* and C. mixta*
vation is on the decline. Closely related to Ch. from C. lundelliana Bailey (2n=40).
nuttalliae*. C. lundelliana grows in S. Mexico, Guatemala and
Honduras. From this parent, C. maxima developed
Chrysobalanaceae in N. Argentina, Bolivia and S. Peru. In this area
the related species C. andreana Naud. grows wild.
CHRYSOBALANUS ICACO L. Icaco plum, Coco
It is probably a weedy derivative of the cultigen.
plum. 2n= . (Sub)trop. America. Cultivated
The wild C. ecuadorensis Cutler &Whitaker
for its fruits.
(2n=40) is closely related to this species and C.
andreana (Cutler & Whitaker, 1969).
Compositae
EUPATORIUM TRIPLINERVE Vahl. (syn. E. aya- SICANA ODORIFERA (Veil. ) Naud. Casa banana,
p a n a V e n t . ) . 2n=51. Trop. America. A perennial Curaba. 2n= . Peru, Brazil to Mexico and W.
herb introduced in Java where it is cultivated as a Indies. This vine is cultivated in trop. America.
medicinal plant.
Dioscoreaceae
MADIA SATIVA Molina. Madia, Tarweed. 2n=32.
Cultivated formerly in Chile as an oil-seed crop. DIOSCOREA PIPERIFOLIA Humb. &Bonpl.
Attempts have been made to grow it elsewhere, 2n= . Brazil. Cultivated there.
but without success. The culture is almost extinct
now. DIOSCOREA TRIFIDA L. Cush-cush yam, Yampi.
2n=54, 72, 81. S. America. Cultivated throughout
the Caribbean area.
POLYMNIA SONCHIFOLIA Poepp. &Endl. (syn.
P . edulis Weddell. ). Yacon strawberry. 2n=60.
Andes. Cultivated there and elsewhere for its tubers.
originated by amphiploidization of two yet unknown

diploid species.
JATROPHA CURCAS L. French purging nut, Phy-

sic nut. 2n=22. Mexico and Bermudas up to Chile
and Paraguay. Cultivated in these and other t r o p i -
cal countries for its curcas oil. In Brazil it is
planted as a living fence.
JATROPHA MULTIFIDA L. ChicaquU, Tortora,

Yuca cimarrona. 2n~22. Trop. America up to
Mexico and West Indies. A shrub cultivated there
as a medicinal crop and elsewhere as an ornamen-
tal.
JATROPHA URENS L. Pendo t r e e . 2n= . Trop.

Dioscorea trifida (Coursey, 1967) America. This t r e e is cultivated in the Philippines
for its leaves which a r e used as a vegetable
(Terra, 1967).
Erythroxylaceae
ERYTHROXYLUM COCA Lam. Coca, Guarigos. MANIHOT ESCULENTA Crantz. Cassava, Manioc,
2n=24. Unknown wild. Probably from high Andes Manihot, Yuca. 2n=36. Cassava is an imported
of Peru and Bolivia. Cultivated at high altitudes food crop troughout the tropics. Secondary gene
in Peru, Bolivia, Argentina, Colombia and Brazil. centres almost certainly exist in Africa (p. I l l )
Some taxonomists include E. novogratense*, and Indonesia (p. 46). Cassava can be divided in-
E. truxillense Rusby and E. bolivianum Burck in to sweet cassava and bitter cassava (M. esculenta
this species. Crantz, 2n=36, M. utilissima Pohl., 2n=36).
The sweet cassava was probably first domestica-
ERYTHROXYLUM NOVOGRANATENSE (Morris) ted in Meso-America (p. 165), from where it
Hieron. Truxillo coca. 2n^ . Andes. Cultiva- spread to S. America. Bitter cassava was domes-
ted at a lower altitude than E. coca*. It was d i s - ticated in northern S. America (Renvoize, 1972).
tributed to the tropics. In Brazil the diversity increased through i n t r a -
species crosses and by hybridization with wild
Euphorbiaceae Manihot species. The weedy M. saxicola Lanj.
HEVEA BENTHAMIANA Muell. - Arg. 2n=36. The

Amazone basin, Brazil, Peru and Bolivia. A t r e e
cultivated for its rubber.
HEVEA BRASILIENSIS (Willd.) Muell.-Arg. B r a -

zilian hevea. P a r a rubber t r e e . 2n=36. The Ama-
zon basin. This is the p r i m a r y gene centre. Se-
condary gene centre in Malaya (p. 46). Cultivated
now in Malaya, Indonesia, Ceylon and in some
other countries. In Africa rubber has been culti-
vated as a f a r m e r ' s crop and as a plantation crop.
A f a r m e r ' s plot often consists of a few t r e e s .
Bouharmont (1960) suggested that H. brasiliensis
Hevea brasiliensis (Dijkman, 1951)

Manihot esculenta {Harris, 1972)
ERYTHROXYLACEAE - GRAMINEAE 149
(2n= ), M. melanobasis Muell. -Arg. and ERIOCHLOA POLYSTACHYA H. B.K. Carib g r a s s .

other weedy species may derive from the cultigen 2n^ . West Indies, Brazil to Ecuador. A grass
(Roger, 1963). The first species is found in NE. cultivated as a forage grass in SE. of USA.
of S. America. M. glaziovii* is a source of r e -
sistance to cassava mosaic disease and drought. GUADUA ANGUSTIFOLIA H.B.K. 2n= . Co-
lumbia and Ecuador up to 1 500 m and in humid
MANIHOT GLAZIOVII Muell. -Arg. Ceara rubber. areas it forms thickets. Cultivated in Puerto Rico,
2n=36. Brazil. Attempts were made to establish Guatemala, Ecuador, Haiti, Honduras, Peru,
plantings for rubber production in Asia and Africa, USA and elsewhere. The stems may grow 18 - 30
but the cultivation was abandoned in favour of para m long. It is the most valuable species in the
rubber (Purseglove, 1968). Western Hemisphere. The wood is strong and
easy to work with. Its stems are used for buil-
OMPHALEA MEGACARPA Hemsl. 2n= . S. dings. N.B. the genus Guadua contains 35 species.
America, especially Brazil and also West Indies. They a r e native to Mexico and S. America. Seve-
A shrub cultivated for its seeds. ral species are cultivated in and beyond the natu-
ral area. This genus is near to the Asiatic genus
PLUKENETIA VOLUBILIS L. (syn. P . peruviana Bambusa.
Muell. - Arg. ). 2n= . N. of S. America and W.
Indies. A vegetable and a fodder crop. GYNERIUM SAGITTATUM (Aubl. ). P . Beauv.
Indian arrowleaf. 2n= . Cultivated in Venezue-
SAPIUM JENMANI Hemsl. 2n= Guiana. Cul- la for its shafts which a r e used for making long
tivated there. arrows.
Gramineae ORYZA ALTA Swallen. 2n=48, genome formula

CCDD. C. and S. America.
BRASILOCALAMUS PUBESCENS (Doell) Nakai.
2n= . Brazil. Related to the Asiatic Bambusa.
ORYZA GRANDIGLUMIS Desv. 2n=48. genome
formula CCDD. C. and S. America. Closely r e -
BROMUS MANGO Desv. 2n= . Chile. Before
lated to O. alta*. Gopalakrishman &Sampath
this country was discovered and European cereals
(1966) suggested that O. grandiglumis derives
were introduced this species was cultivated. Its
from O. alta.
cultivation has disappeared now.
ORYZA LATIFOLIA Desv. 2n=48, genome formu-
BROMUS UNIOLOIDES H.B.K. (syn. B. catharti-
la CCDD. C. and S. America.
c u s V a h l . , B. schraderi Kunth). Rescue g r a s s ,
Schrader's brome. 2n=28, 42, 56. S. of USA, S.
PASPALUM DILATATUM P o i r . Dallis g r a s s .
America, Australia and Europe. It probably deve-
2n=(30), 40, 50, (60). Probably Chaco savanna.
loped in S. America particularly Argentina. In
The common type has 2n=50 and the genome for-
the Andean region it (syn. B. haenkeanus (Presl)
mula AABBC. It probably originated from a cross
Kunth) (2n=42) grows wild.
of an unreduced gamete (AABB) and C genome
donor species. The dilatatum type pauciciliatum
BROMUS WILLDENOWII Kunth (syn. B. unioloi- has 2n=40, genome formula AA.BC,. Type u r u -
des (Willd. )Respail, Festuca unioloides Willd. ). guaiana (2n=60) genome formula AAA.A BB and
Common rescue g r a s s . 2n=28, 42. A plant from type t o r r e s (2n=60) (univalents). These types a r e
S. America cultivated as a fodder g r a s s . all apomlctic. The only sexual type is 'yellow
anther' (2n=40), genome formula AABB. Intro-
CHUSQUEA ANDINA Phil. 2n= . Chile, where duced now into other countries. In Japan sexual
it reaches the snow bounderies. NB. The genus and apomictic types occur. They a r e triploids,
Chusquea Kunth Includes over 70 species native to 2n=3x=30. P . juergensii Hack. (2n=20) or a close-
S. America, Mexico and E. India. It is typical ly related type might be the donor parent of the A
for the Andes, where the species form dense or B genome (Burson &Bennett, 1972).
thickets. Many of them a r e very ornamental and
many be very valuable for introduction to the hu-
PASPALUM NOTATUM Fluegge. Pensacola Bahia
mid subtropical areas of USSR and elsewhere b e -
g r a s s . 2n=20, (30), 40. It is not clear whether
cause they are very hardy.
Pensacola Bahia grass originated in Florida or
that it came from interior of S. America (Burton,
CHUSQUEA CULEON E. Desv. ex C. Grav. 1967), where a wide variation of types on the
2n= .Chile.
Berduc Island of Rio Parana, Uruguay was found.
Widespread in S. USA. There it was first found in
CHUSQUEA DEPAUPERATA Pilg. 2n= . Peru, Pensacola, Florida.
up to 3 400 m.
CHUSQUEA ULIGINOSA Phil. 2n= . Chile. PASPALUM PLICATULUM Michaux. 2n=20, 40,
(60). S. America. Cultivated in Australia.
CORTADERIA ARGENTEA Stapf (syn. Gynerium
argenteum Nees). Pampas g r a s s . 2n=70. S. A m e - SORGHUM ALMUM Parodi. Black sorgo, Colum-
rica. A grass cultivated as a source of pulp. bus g r a s s . 2n=40. Argentine where this forage
crop probably originated from a cross of S. hale-
pense* and a variety of S. bicolor*. It has a Such plants at the silking stage can easily be m i s -
lower sugar content than S. halepense. taken for teosinte (Roberts et a l . , 1957). It is
thought that the cause might be a virus inducing
TRIPSACUM AUSTRALE Cutler &Anderson. profuse branching. Pira might have also occurred
2n- . S. America, extending from Venezuela in Venezuela and Bolivia. It is related to the P e -
to Paraguay. Also found in NC. Peru. ruvian Confite Morocho and Confite Puntiagudo.
At a later stage less primitive and developed
races originated in S. America. In the coastal
eastern S. America the race 'Coastal Tropical
Flint' was cultivated. This race was also found in
the West Indies, it was probably introduced from
the continent. However, it is possible that in both
areas a similar race developed from identical
parents (Hatheway, 1957).
Owing to the high variation of maize in S.
America this region is considered as a secondary
centre of diversity. For instance, Grobman et al.
(1956) concluded that Peru appears the home of
pericarps colour genes. In the dept. Ancash all
three alleles of the A locus and all 7 of the P locus
a r e found. From S. America maize was returned
to C. America and Mexico and was taken to N.
America and the Old World.
Guttiferae
MAMMEA AMERICANA L. Mammey apple. Mamey.
2n= . Trop. America and West Indies. Culti-
vated there for its edible fruits and for its scented
flowers which a r e used to prepare the liquor Eau
de Créole.
Tripsacum australe (Hernandez, 1973)
RHEEDIA ACUMINATA (Ruiz &Pav. ) Planch. &
Triana. 2n= . Colombia up to Peru. Cultivated
TRIPSACUM DACTYLOIDES (L.) L. 2n=36, 72. as a compound t r e e in NW. of S. America.
Described on p. 165. It has a rather thick rachis
and may have been the source of this c h a r a c t e r i s - Juglandaceae
tic of some primitive S. American maize culti-
vars (Zea mays, p. 150) like Cabuya and Sabanero JUGLANS HONOREI Dode. Ecuador walnut.
(Galinat, 1969). 2n= . Rootstock of J. regia*.
ZEA MAYS L. Maize. Corn. 2n=20. Maize d e r i - Lauraceae

ves from teosinte, Z. mexicana*, which grows NECTANDRA CINNAMOMOIDES Nees. 2n=
wild in C. America (p. 167). From here it spread Equatorial Andes. A t r e e cultivated in Ecuador as
south and north. After its introduction into S. a spice plant.
America, its development there became agricul-
turally more advanced than in its area of domesti- PERSEA LEIOGYNA Blake. 2n= . Probably
cation. This might be a result of the absence of trop. America. A fruit t r e e .
its wild and weedy relatives. A secondary centre
in S. America. These S. American primitive Lecythidaceae
maize varieties were taken to C. America. In
Mexico they were described as Pre-Columbian BERTHOLLETIA EXSELSA Humb. &Bonpl. B r a -
Exotic races (p. 166). In the 19th Century a secon- zil nut, P a r a nut. 2n=34. The Amazon forests.
dary centre of diversity arose here when cultivars The kernels and oil a r e eaten. The oil/kernel
from the Amazonian and Paraguayan lowlands ratio is quite high: 60-70%.
mixed with 'coastal tropical' flint from the West
Indies and with flint cultivars from the southern LECYTHIS ZABUCAJO Aubl. Sapucaia nut, P a r a -
slopes of Bolivia (Brandolini, 1970). dise nut, Monkey pot. 2n= . The forests of
Primitive Races - all belonging to the pop- Guianas and Brazil. Not much cultivated.
corn type - have been found in Columbia, Equador,
Bolivia, Peru and Chile. In Colombia the P r i m i - Leguminosae
tive Races a r e called Polio and P i r a . Polio might ACACIA CAVENIA Bert. Cavenia acacia, Espino
be related to the Peruvian Confite Morocho. Col- cavan. 2n=26, 52. S. America. A small t r e e c u l -
lections of Polio made at Medellin, at an altitude tivated as a source of perfume. Related to A.
considerably lower than their normal habitat, farnesiana*.
segregate plants which in their general aspects
a r e almost identical to maizoid teosinte in Mexico.
ACACIA FARNESIANA (L.) Willd. Sweet acacia. Krapovickas (1969) pointed to five S. American
2n=52, (104). Probably trop. America (Purse- centres of diversity: 1. the Guarami region, the
glove, 1968). This shrub is cultivated esp. as an basins of the Paraguay and Parana rivers, which
ornamental and for its perfumery, where it has is the centre of variation for the ssp. fastigiata
run wild in the tropics. In S. France the very Waldron var. vulgaris H a r z . , the Spanish type,
fragrant Cassie Flowers a r e the source of Cassie 2. The region of Goias and Minas Geraes where
Ancienne. ssp. fastigiata var. fastigiata, 3. The region of
Rondonia and NW. Mato Grosso. This region is
AESCHYNOMENE AMERICANA L. (syn. A. glan- not yet fully studied. It includes ssp. hypogaea
dulosa Poir. ). 2n= . Trop. America. Used in var. hypogaea (syn. A. africana Lour., A. nam-
Indonesia and elsewhere as a green manure, soil byquarae Hoehne), the Brazilian or Virginia type,
cover and as a forage crop. and the distantly related cultivated A. villosuli-
carpa*, 4. The region of the eastern foothills of
ALBIZIA CARBONARIA. 2n= . Colombia and the Andes in Bolivia, which contains a great v a r i a -
C. America. Cultivated in Puerto Rico (Whyte et bility of var. hypogaea. Some introgression may
a l . , 1953). occur with var. fastigiata resulting in the forms
Overo, Pintado and Cruceno, and 5. Peru, which
ARACHIS GLABRATA Benth. Arb peanut. 2n=40. is the centre of variability of ssp. hypogaea var.
Brazil, Argentina and Bolivia. Perennial used for hirsuta Kohier (syn. A. asiatica Lour. ).
pastures and hay (Prine, 1964).
ARACHIS HYPOGAEA L. jGrjOugdijut, Peanut.

2n=40, genome formula A A B B . S. America.
P r i m a r y gene centre in Argentine and Bolivia.
Secondary centres in Nigeria, Senegal and Congo
(p. 118). Probably domesticated in the Gran Chaco
a r e a . As it is a tetraploid and no wild groundnut
Arachis monticola
\ "~n r ')
\ ' "~'
V î "\
)Vkr r—l
\- -^
/ ( h f
i /
/1
1
i
Gene centres of Arachis hypogaea (Krapovickasy, 1972)
has been found, an amphiploidization of one or two Arachis monticola (Zhukovsky, 1971)
wild diploid species is suggested, or the ground-
nut may be a cultigen of a wild tetraploid species
that has arisen in this way. This could be A. ARACHIS VILLOSULICARPA Hoehne. 2n=20. Cul-
monticola Krapov. & Rig. This species (2n=40) tivated by the Indians of Juruena and Diamantino
grows wild in the mountains of the Jujuy Province, of the Mato Grosso in Brazil. Perennial and not
NW. Argentina. It crosses easily with the ground- closely related to A. hypogaea* (Krapovickas,
nut. It can be used to improve disease resistance 1969).
etc. of the groundnut. A. villosa Benth. (2n=20)
has the genome formula A A (Raman, 1973). CANAVALIA ENSIFORMIS (L. ) DC. Jack bean,
Horse bean, 2n=22. S. America. Secondary cen-
tre in India. Sauer and Kaplan (1969) mentioned sweet fruit pulp (Uphof, 1968).
C. boliviana Piper (2n= ), C. brasiliensis
Mart, ex Benth. (2n= ), C. dictyota Piper INGA PREUSSII Harms. 2n= . El Salvador.
(2n= ), C. maritima (Aubl. ) Thou. (2n= ) Used as a shade t r e e .
and C. piperi Killip &MacBride (2n=22). as p o s s i -
ble ancestors. An ancient legume, now cultivated INGA PUNCTATA Willd. 2n= . S. and C.
in the tropics as a green manure or fodder crop. America. Used as a shade t r e e .
CANAVALIA PLAGIOSPERMA Piper. 2n=22. P r o - LEUCAENA GLAUCA (L. ) Benth. Jumpy bean.
bably trop S. America. Possible ancestor of the 2n-(36). 104. Trop. America. Used as a shade
Andean C. piperi Killip &MacBride (2n=22) tree and as green manure. Selections have been
(Sauer, 1964). One of the earliest cultivated crops made with a low mimosine content.
in S. America, but not cultivated at present. It
resembles C. ensiformis*. LONCHOCARPUS UTILIS Smith. 2n=44. Peru.
Cultivated as a source of rotenone.
CENTROSEMA PLUMIERI (Turp. ) Benth. 2n=20.
Trop. America. This cover crop and green m a - LUPINUS BOGOTENSIS Benth. 2n= . Bolivia.
nure has been distributed throughout the tropics. Cultivated there.
CENTROSEMA PUBESCENS Benth. 2n-20. Trop. LUPINUS MONTANUS H.B.K. 2n= . Peru,
America. This cover crop and green manure is Bolivia. Guatemala and Mexico. Cultivated in
distributed throughout the tropics. Bolivia.
CROTALARIA ANAGYROIDES H. B.K. 2n=16. LUPINUS MUTABILIS-TAURIS-CUNNINGAHAMII-

Trop. America and West Indies. Cultivated as a CRUCKSHANKSn species group. 2n=48. Andes
cover crop, green manure and fodder. region between Bolivia and Venezuela. This group
of species, also named L. mutabilis Sweet is not
DESMODIUM CINEREUM DC. Trebold. 2n= yet well described. Formerly widely cultivated in
S. America. Cultivated for green manure and as its native centre. Still cultivated in Bolivia.
a vegetable (Terra, 1967). Apparently farmers have not tried to select for
sweet types. At present this species is used as
DESMODIUM DISCOLOR Vog. 2n=22. Brazil. A 'bitter protection rows' around fields of Vicia
forage plant. faba and Pisum sativum to stop animals entering
fields. Types have been developed that grow well
DESMODIUM INTORTUM (Mill. ) Urb. Greenleaf. under tropical short-day conditions, that have
2n=22. C. America and Brazil. Cultivated in pods which do not open and soft-coated seeds rich
Australia (Hutton, 1970). in protein and of low alkaloid content (Brücher,
1970; Hackbarth &Pakendorf, 1970). The big
DESMODIUM UNCINATUM (Jacq. ) DC. Silverleaf. seeds are used to prepare tarwi or ullu. Other
2n=22. S. America. Cultivated in Australia (Hut- wild, but possible valuable Lupinus-species of the
ton, 1970). American continents should be domesticated. They
a r e often shrubby and have small seeds (Brücher,
DIPTERYX ODORATA Willd. Tonka bean, Dutch 1970).
tonka, 2n=32. Forests of trop. America, Venezue-
la, the Guianas and the lower Amazon basin. MIMOSA INVISA*
The tree is cultivated now in Venezuela. Malaya,
West Indies and some other tropical countries MYROXYLON BALSAMUM (L. ) Harms. Balsam of
(Cobley, 1963). Peru. 2n^28. Var. pereira (Royle) Harms is
spread in Guatemala and San Salvador. It is a
ERYTHRINA GLAUCA Willd. 2n=42. S. America. source of balsam. It was cultivated in the impe-
A shade t r e e in cacao plantations. rial gardens of the Aztecs in Mexico (Mansfeld,
1959).
ERYTHRINA MICROPHERYX Poepp. Anauca.
2n= . Peru. A shade tree in cacao plantations. PACHYRHIZUS APIHA (Wedd. ) Parodi. 2n=22.
Probably a cultigen developed by the Indians in
GLIRICIDIA SEPIUM (Jacq. ) Steud. (syn. G. macu- Bolivia and N. Argentina.
lata Benth. ). 2n=20, 22. Mexico, C. America and
N. of S. America. A shade t r e e , green manure PACHYRHIZUS TUBEROSUS (Lam. ) Spreng. Yam
and fodder crop. bean, Potato bean, Jicama. 2n=22. The head-
waters of the Amazon. From there it was d i s t r i -
INDIGOFERA ANIL L. (syn. I. suffruticosa Mill. ). buted to other parts of S. America and parts of
Indigo plant. 2n=12. S. America. Once much cul- the West Indies. The young pods and tubers a r e
tivated in the tropics for its dye (indigo) (Heiser, eaten.
1965).
PHASEOLUS ABORIGINEUS Burk. 2n=22. Forests
INGA FEUILLEI DC. (syn. I. reticulata Spr. ). of northwestern Argentine Andes. Probably exten-
2n= . Peru. This t r e e is cultivated there for ded through Bolivia, Peru, Ecuador up to Hondu-
LEGUMINOSAE - MALVACEAE 153
r a s (Burkart &Bücher, 1953). It might be the GOSSYPIUM BARBADENSE L. (syn. G. vitifolium

progenitor of P . vulgaris* in Peru (Heiser, 1965). Lam., G. peruvianum Cav. ). Sea island cotton.
2n=52, genome formula (AADD)„. It has been p r o -
PHASEOLUS LUNATUS L. Lima bean, Sieva bean, posed that G. barbadense a r o s e r r o m a cross and
Butter bean, Madagascar bean, Burma bean. amphidiploidization of G. arboreum* and G. r a i -
2n=22. C. America, and in the Andes from Peru mondii*. G. arboreum could have been introduced
to Argentine. Kaplan (1965) showed that the big into Peru by way of Asia and the Pacific islands.
lima bean of Peru was first domesticated in the Another hypothesis is that an African diploid
Andean highlands and that the small lima bean of reached S. America by way of Atlantic. This d i -
Mexico may have arisen in the Pacific coastal ploid would probably have been G. herbaceum*.
foothills of Mexico (p. 168). A small-seeded sub- As Bird (1948) found G. barbadense material
species (ssp. microsperma, Sieva or Small Lima) at Huaca Prieta, Peru which was dated 2 400 BC.
originated by natural selection. It spread to the the introduction of the African Gossypium species
Antilles Islands. and its amphidiploidization with G. raimondii must
have taken place long before that time. The main
PHASEOLUS VULGARIS L. Common bean. 2n=22. point is how this African species reached Peru.
For origin see p. 168. The earliest remains of However, the centre of origin N. Peru is in
cultivated common beans have been found in the the arid mountainous interior of the prov. Tumbes.
Guitarrero Cava in Peru. It dates from about The ssp. darwinii is closely related and is ende-
6 000 BC. The 'domesticated' characters a r e mic in the Galapagos Islands. At present it is
especially dark red brown and dark red beans 'contaminated' by hybridization with exotic intro-
(Kaplan et a l . , 1973). ductions. Secondary centre in Peru.
In S. America G. barbadense spread south
PITHECELLOBIUM SAMAN Benth. Rain tree, and eastwards to NW. Argentine. Some other
Saman, Cow tamarind. 2n^26. Trop. America. forms a r e found in S. America. The Tanguis
It is used as a shade t r e e in cacao and coffee plan- variety is a selection from Tumbes. In Chile and
tation. Peru the Pacific assemblage is found, characte-
rized by broad leaves and intense hairiness of the
PROSOPIS JULIFLORA DC. Mesquite. 2n=28, 52, underside of the leaf. This character induces r e -
56. Trop. America. A small t r e e cultivated for
various purposes (Mansfeld, 1959).
RHYNCHOSIA MINIMA (L. ) DC. 2n=22. S. A m e r i -

ca. Cultivated in Australia and elsewhere as a
fodder and pasture crop.
STYLOSANTHUS GUIANENSIS SW. (syn. S. s u r i -

namensis Miq. ). 2n=20. Guiana. Used as a food
plant for livestock and as a soil conservator.
VICIA GRAMINEA Smith. 2n=14. Argentine and

Chile. Occasionally cultivated for its seeds as a
source of anti-N-lecitin (Nijenhuis et a l . , 1961).
This is used as a test serum for the human N-
blood group.
Malpighiaceae
BANISTERIOPSIS CAAPI (Spruce ex Griseb. )
Morton. 2n=20. S. America. A woody vine culti-
vated in the Amazon region as a drug and narcotic.
BUNCHOSIA ARMENIACA (Cav.) DC. 2n=

The Andean region. A shrub cultivated in Ecuador
for its fruits. Distribution of the NewWorld cottons inthe 13th century:
Gossypium barbadense (11), G. hirsutum var. marie-galante
MALPIGHIA GLABRA L. (syn. M. punicifolia L. ). (12) and G. hirsutum punctatum (13) (Hutchinson, 1962)
Barbado cherry, West Indian cherry. 2n=
West Indies and N. S. America. Cultivated there
and elsewhere for its fruits. It also makes a good sistance to jassids, Empoasca ssp. The lint of
hedge, like M. coccigera L. (Purseglove, 1968). G. barbadense is usually coarse with a lenth up to
34. 5 mm. The lint of an Ecuador type, of Sea I s -
Malvaceae lands and Egyptian is fine and silky with a length
up to 37.5 mm. It is possible that the Ecuador
ABUTILON OXYCARPUM F. Von Muell. 2n=14. type is the parent of the Sea Islands/Egyptian com-
S. America and Australia. Cultivated for its plex (p. 73, 120) (Harlan, 1970). The Atlantic
fibres. assemblages include the kidney cottons (seeds
fused in a kidney-shaped cross). They have a wide Grande do Norte, N. Brazil. He suggested that
distribution in northern S. America and the i s - this area is almost certainly the centre of origin
lands of C. America. They have been taken to of the whole Upland group. From here this cotton
Africa, India, Ceylon, Indonesia and elsewhere. dispersed first northward to the Amazon, then
Secondary centres in Egypt (p. 120) and in along the Amazon and across the Andes into Ecua-
Turkmenia - Tadzhikistan - S. Uzbekistan, USSR dor. W. Colombia and possibly still further north.
(p. 73). In another direction this cotton dispersed north-
On the Sea Islands of S. Caroline, USA the ward through the Guyanas passing the West Indies
Sea Island cottons developed after cottons from to E. Colombia and further northward into C.
Bahamas or Jamaica (p. 177) were introduced America via Yucatan. C. America must be con-
(Hutchinson. 1962). sidered as a secondary centre of diversity (p. 169).
Wild and semi-wild marie galante cotton were ob-
served in Florida until some years ago. Upland
, ' ' V. a. cotton also dispersed southward to E. Brazil. At
present this race is also grown in Ghana.
A second important perennial is race puncta-
%& tum. which is found around the coast of the Gulf
of Mexico from Yucatan to Florida and the Baha-
mas and some other islands (p. 169). At present
V\ 14 ~ G. hirsutum Cambodia (a latifolium type) is cul-
tivated in S. India. Their way of spread was p r o -
^¥JDr/^ -16 bably S. America-Philippines-Cambodia-S. India.
GOSSYPIUM KLOTZSCHIANUM Anderson. 2n=26,

^ " ^ genome formula D„ „D . Galapagos Islands.
14 ^ ^ Var. davidsonii is lound on the shores of Gulf of
California and the Revilla Gidego islands (p. 124).
15 C GOSSYPTUM RAIMONDII Ulbr. 2n=26, genome

formula D_D . Formerly N. Peru. Now extinct in
V /HT—\\ its original habitat and only found in collections
)A [)) (Harland, 1970)'. This species is a source of
l°y
hairiness gene Hfi conditioning resistance to j a s -
sid, Empoasca ssp. Probably one of the parental
species of G. barbadense* and G. hirsutum.
k f
\ \ &>
Distribution of annual cottons in the NewWorld at 1960: Gossy-

pium hirsutum var. uplands (14), G. barbadense var. Egyptians
(15) and G. barbadense var. Sea Islands (16) (Hutchinson, 1962)
GOSSYPIUM CAICOENSE. 2n=52, genome formula

AADD. This tetraploid species was discovered in
NE. Brazil in 1967.
GOSSYPIUM HIRSUTUM L. Upland cotton. 2n=52,

genome formula (AADD) . The common theory is
that G. hirsutum arose from an amphiploidization
of the Old World G. arboreum* or G. herbaceum*
and G. raimondii. G. raimondii is the parent of
the D genome and one of the first two the donor of
the A genome. It is not known when this amphi-
ploidization took place, but material collected
from Tehuacan Valley in Mexico and dated 3 500-
2 300 BC. appears to be fully domesticated
(Smith &Stephens, 1971). It is also not known
whether G. barbadense reached Peru by way of Gossypium raimondii
Asia or whether G. herbaceum reached eastern
S. America from W. Africa. Harland (1970) ob-
served wild plants of an exceedingly primitive GOSSYPIUM TOMENTOSUM Nutt. 2n=52, genome
perennial race marie galante in the state Rio formula (AADD),. Hawaii. A fuzzy-seeded but
MALVACEAE - PALMAE 155
not linted species. At one time it was believed EUGENIA UVALHA Camb. Uvalha. 2n- . S.
that if the origin and relationship of this species Brazil. Cultivated for its fruits.
were elucidated the problem of the origin of G.
barbadense* and G. hirsutum* could be solved. FEIJOA SELLOWIANA Berg. Feijoa. 2n=22. S.
However it appears that its origin is independent Brazil, Uruguay, Paraguay and N. Argentine.
of those of the New World species (Hutchinson, Also its primary centre of diversity. Sometimes
1962). cultivated for its fruit in hot countries e. g. the
Caucasian coast of the Black Sea where it grows
WISSADULA CONTRACTA (Link. ) R . E . F r i e s . well.
2n--14. Trop. America. Cultivated in W. Java for
fibre (van Borssum Waalkes, 1966). MYRCIARIA CAULIFLORA Berg. (syn. Eugenia
cauliflora (Berg. ) DC. Jabotica. 2n= . Brazil.
WISSADULA PERIPLOCIFOLIA (L. ) P r e s l ex Cultivated for its fruits (Purseglove, 1968).
Thw. 2n=14. Probably introduced in Ceulon as a
source of fibre for which purpose it is still used MYRCIARIA JABOTICABA Berg. 2n= Brazil.
(van Borssum Waalkes, 1966). The degree of Cultivated in the tropics for its fruits.
variability of this species is very small in Malay-
sia. Some varieties and forms have been described PSIDIUM GUINEENSE SW. 2n= . The West
for American representatives. This may point to Indies and trop. America. Occasionally cultivated.
an American origin. A pantropical weed.
PSIDIUM LITTORALE Raddi (syn. P. cattleianum
Marantaceae Sabine). Strawberry guava. 2n=88. Brazil. A
small t r e e introduced in the tropics and subtropics.
CALATHEA ALLOUIA (Aubl. ) Lindl. Sweet corn Var. lucidum Degener, Chinese strawberry guave
root. 2n= . The W. Indies. A tuber crop cul- yields fruits of improved quality (Uphof, 1968).
tivated there and in S. America.
Nyctaginaceae
MARANTA ARUNDINACEA L. Arrowroot, B e r -
muda arrowroot. 2n=18, 48. N. S. America and MIRABILIS JALAPA L. Marvel of Peru, Four
the L e s s e r Antilles. Cultivated in the tropics for o'clock, False jalap. 2n=(54), 58. S. America.
its rhizomes containing starch. Spread over the whole world and in W. Africa as
a fetish plant. Cultivated as an ornamental. Tube-
Musaceae rous roots were used as jalap. Elsewhere a sub-
tropical weed.
MUSA cultivars of the AAB group - Plantain sub-
group. French Plantain, Horn Plantain. 2n=33. Onagraceae
India (p. 69). Secondary centre: trop. America.
FUCHSIA MAGELLANICA Lam. Fuchsia. 2n=22,
Myrtaceae 44. S. America. Planted as hedges in Azores,
Ireland and W. Britain.
ABBEVILLEA FENZLIANA Berg. 2n= . Bra-
zil. A small t r e e cultivated for its edible fruits. Oxalidaceae
BRITOA ACIDA Berg. P a r a guava. 2n= . Bra- OXALIS TUBEROSA Mol. Oca. 2n=(14), 60, 63-64,
zil. A shrub cultivated for its fruits. 68-70. Cultivated in the Andes from Colombia to
Bolivia for an extremely long time. Introduced
CAMPOMANESIA GUAVIROBA Benth. &Hook. also in Europe where it was cultivated like the
2n= . S. Brazil. Cultivated for its edible Mexican O. deppei Lodd. (2n=14, 56) as a vegeta-
fruits. ble by amateurs (Uphof, 1968). Several colours of
the tubers have been observed. It should not be
CAMPOMANESIA LINEATIFOL1A Ruiz. &Pav. confused with Tropaeolum tuberosum*.
(syn. C. cornifolia H. B.K.). 2n= . E. Andes.
Cultivated as a fruit t r e e in Peru (Mansfeld, 1959). Palmae
COROZO OLEIFERA (H.B.K. ) Bailey, (syn.
EUCALYPTUS CAMALDULENSIS Dehn. Longbeak Elaeis melanococca Gaertn. ). 2n=32. C. America
eucalyptus. 2n=22. P r i m a r y centre: Australia to Colombia and Amazon area. Cultivated for its
(p. 58). Secondary centres: Brazil, Argentine and oily fruits. It can be crossed with the African oil
the Mediterranean region (p. 104). palm, Elaeis guineensis* producing fertile hybrids.
EUGENIA DOMBEYANA DC. Grumichama. GUILIELMA GASIPAES (H.B.K.) L.H. Bailey.
2n= . Peru and S. Brazil. A tree cultivated Peach palm, Peribaye. 2n= . S. and C. A m e -
for its fruits. rica. Cultivated in S. America.
EUGENIA UNIFLORA L. Pitange, Surinam cherry. OENOCARPUS BACABA Martlus. 2n= . Ama-
2n=22. Brazil. Cultivated in the tropics and s u b - zon area to Surinam and Gayana. A palm cultiva-
tropics. ted on compounds for its oily fruits.
Passifloraceae PHYTOLACCA DIOICA L. 2n=36. Temperate and

PASSIFLORA ALATA Dryand. Maracuja. 2n= subtrop. S. America. Cultivated as an ornamental
Peru and Brazil. A woody vine cultivated in B r a - and shade plant.
zil for its fruits.
RIVINA HUMILIS L. Rouge plant. 2n=108. The
PASSIFLORA ANTIQUIENSIS Karst, (syn. P. van- tropics of the Old and New Worlds. Cultivated in
volxemii (Lem. ) Triana &Planch. ). 2n^ Colombia for its b e r r i e s which a r e a source of
Banana passion fruit. Colombia. A woody vine red dye.
cultivated e.g. in New Zealand for its fruits.
Piperaceae
PASSIFLORA CEARENSIS Barb. 2n= . Brazil. PIPER ADUNCUM L. 2n= . Trop. America.
There it is also cultivated for its fruits. Used as a soil conservant.
PASSIFLORA EDULIS Sims. Passion fruit. 2n=18. Portulacaceae

S. Brazil. Widely distributed throughout the t r o -
pics and subtropics. The fruits a r e especially TALINUM TRIANGULÄRE*
used for juice preparation.
Rhamnaceae
PASSIFLORA FOETIDA L. 2n=18, 20, 22. West COLUBRINA RUFA Reiss. 2n= . Brazil. Cul-
Indies and S. America. Weedy. Distributed to tivated for its medicinal bark and other purposes.
many tropical countries in Africa and Asia, where
it has naturalized. Its fruits a r e sometimes eaten. Rosaceae
In Malaya and E. Africa it has been used as a
cover crop. FRAGARIA CHILOENSIS L. Chiloe strawberry,
Ambato strawberry. 2n=56, genome formula AAA'
PASSIFLORA LAURIFOLIA L. Water-lemon. A'BBBB. The Pacific coastal region of N. and S.
Jamaica honeysuckle. Belle apple. Pomme de America and Hawaii. Formerly cultivated t h e r e .
liane. 2n=18. Thickets and forest fringes in the It is one of the parents of F. x ananassa*.
West Indies and NE. S. America. Cultivated for
its fruits in the 17th Century. Spread throughout RUBUS BRASILIENSIS Mart. 2n= Brazil. A
the tropics (Purseglove, 1968). shrub cultivated for its fruits.
PASSIFLORA LIGULARIS J u s s . Sweet granadilla. RUBUS GLAUCUS Berth. 2n= . Costa Rica to
2n^l8. Trop. America. Its sweet fruits a r e much Ecuador. Cultivated in the Andes.
used in the mountainous regions of Mexico and C.
America (Purseglove, 1968). RUBUS MACROCARPUS Benth. Colombian berry.
2n= . Colombia and Ecuador. Cultivated for
PASSIFLORA MALIFORMIS L. Curuba. 2n= its very large fruits (5 cm long).
Trop. America. A vine cultivated for its fruits.
Rubiaceae
PASSIFLORA MOLLISSIMA (H.B.K. ) Bailey. CEPHAËLIS IPECACUANHA (Stokes) Baill. Ipecac,
Banana passion fruit, Tasco, Caruba de Castilla. Ipecacuanha. 2n=22. Brazil. Introduced into India
2n=18. The Andes. Especially cultivated in Ecua- and Malaya. There small plantings were e s t a -
dor and Bolivia. Introduced in other countries. blished. Roots of wild and cultivated plants are
the source of ipecac or ipecacuanha used to treat
PASSIFLORA PSILANTHA (Sodiro) Killip. Gullan. amoebic dysentery.
2n= . Ecuador. A vine cultivated for its fruits.
CINCHONA LEDGERIANA Moens ex Tremen (syn.
PASSIFLORA QUADRANGULARIS L. Giant grana- C. calisaya var. ledgeriana How., C. officinalis
dilla, Barbadine. 2n-18. Trop. S. America. Cul- L., C. calisaya Wedd. and C. succirubra Pav. ex
tivated since 18th Century for its fruits. Now wide- Klotzseh. Quinine. 2n=34 (all species). These
ly distributed in the tropics. species a r e taken together. They all come from the
same centre of diversity: Andes mountains of S.
PASSIFLORA TRIPARTITA (Juss. ) Poir. Tasco. Peru, Bolivia and S. Ecuador. Here many Cin-
2n=18. Ecuador. Cultivated there. chona species a r e found and the great diversity of
botanical varieties is caused by natural hybridiza-
Peperomiaceae tion between the species and varieties. Plantations
PEPEROMIA PELLUCIDA H.B.K. 2n= . S. in Indonesia and Ceylon and recently in E. Africa.
America. In Africa this pantropical weed is cul- The original introductions in the Asian countries
tivated as a vegetable and medicinal crop. were very probably a mixture of true species and
their hybrids. From this material C. ledgeriana
Phytolaccaceae was derived but it is thought to be a variety of
C. calisaya, and is also considered a hybrid of
PHYTOLACCA CHILENSIS Miers. 2n- . Chile. C. calisaya, C. succirubra and C. lancifolia
A perennial herb cultivated for its berries which Mutis. C. succirubra which is used as rootstock
are a source of red dye.
PASSIFLORACEAE -SOLANACEAE 157
is probably a variety of C. pubescens Vahl. (van (Pickersgill, 1969).

Harten, 1969).
CAPSICUM PUBESCENS Ruiz &Pav. 2n=24. Un-
Sapindaceae known wild. Cultivated in the highlands of S. Ame-
rica.
MELICOCCUS BIJUGATUS Jacq. Kanappy tree,
Kinnup tree, Bullace plum, Honey berry, Spanish
lime, Geneps. 2n=32. Trop. America. Cultivated
there for its edible fruits (Mansfeld, 1959).
PAULLINIA CUPANA (H.B.K. ). Guarana. 2n=

S. America. Cultivated in Brazil for its seeds,
used as a coffee.
SAPINDUS SAPONARIA L. Soap wood t r e e , Soap

t r e e , Soap berry t r e e . 2n= . Trop. America.
Cultivated there and elsewhere for its fruits.
Sapotaceae
LUCUMA NERVOSA A. DC. (syn. L. rivicoa
Gaertn.f., Pouteria campechiana (H.B.K.) Baenhi).
Egg fruit, Canistel. 2n= . NE. of S. America.
Cultivated in trop. America for its fruits.
LUCUMA OBOVATA H. B.K. (syn. Pouteria lucu-

ma (Ruiz &Pav. ) O. Kuntze. Lucumo. 2n=
Chile and Peru. This t r e e is cultivated for its Capsicum baccatum var. baccatum (Eshbaugh, 1970)
fruits.
LUCUMA PROCERA Mart. (syn. Urbanella p r o -

cera P i e r r e ) . Macarandiba. 2n= . This fruit
t r e e is cultivated in Brazil.
MANILKARA BIDENTATA (A. DC.)Chev. (syn.

Mimusops balata P i e r r e ) . Balata, Bully, Bullet,
Purgio, Quinilla. 2n= . S. America and T r i n i -
dad. The wild t r e e s are tapped for latex (balata).
POUTERIA CAIMITA (Ruiz &Pav. ) Radlk.

2n= . Peru to E. Ecuador and Guyanas. A tree
cultivated for its fruits.
Simaroubaceae
QUASSIA AMARA L. Surinam quassis, Bitter
wood. 2n= . N. of S. America. Cultivated for
its wood which is used medicinally, and also as
an ornamental t r e e .
Solanaceae
CAPSICUM BACCATUM H.B.K. (syn. C. angulo-
sum Miller). Pepper. 2n=24. The wild type is var.
baccatum (syn. C. microcarpum Cav. ). It occurs
in Peru, Bolivia, Paraguay, N. Argentina and S.
Brazil. It is the parental type of the cultivated
type var. pendulum (Willd. ) Eshbaugh (syn. C.
pendulum Willd.). This cultigen was originally Capsicum baccatum var. pendulum (Eshbaugh, 1970)
found in the same area as var. baccatum and in S.
Columbia, Ecuador and in Chile. Now it is also
CYPHOMANDRA BETACEA (Cav.) Sendt. (syn.
cultivated elsewhere (Eshbaugh, 1970).
C. crassifolia). Tree tomato. 2n=24. Peru. Un-
known wild. Cultivated in the Andean region e s p e -
CAPSICUM CHINENSE Jacq. (syn. C. sinense cially in Ecuador. Other species of this genus a r e
Jacq. ). 2n=24. This pepper was originally cultiva- found in S. America and partly in C. America.
ted in the West Indies and lowland S. America, One of them, is C. hartwegi Sendt. ; its fruits are
from S. Bolivia to S. Brazil. Closely related to harvested in Colombia, Chile and Argentina.
C. frutescens*. It may have originated from it
LYCOPERSICON CHILENSE Dun. 2n= . The A green-fruited species. The glabratum is self-
coastal strip of Peru and northern Chile. A wild compatible. It is characterized by disease r e s i s -
tomato often found growing together with L. p e r u - tance, e.g. tomato mosaic virus (Marmon tabaci
vianum. However they do not c r o s s . This species Holmes).
is characterized as a source for resistance to all
tomato diseases except Phytophthora. LYCOPERSICON PERUVIANUM (L.) Mill. 2n=24.
Chile and Peru. A green, small-fruited wild s p e -
LYCOPERSICON ESCULENTUM Mill. Tomato. cies . Most plants a r e gametophytic self-incompa-
2n=24. The centre of the genus Lycopersicon is a tible, although some plants have been found to be
narrow belt of the S. American west coast limited self-compatible (Hogenboom, 1968). It is a source
by the equator and 30 Sand the Andes andthe of tomato mosaic virus tolerance.
Galapagos Islands. The greatest variability of the
tomato is however outside this area, in the V e r a - LYCOPERSICON PIMPINELLIFOLIUM Mill. (syn.
cruz-Puebla area in Mexico (Jenkins, 1948). This L. esculentum ssp. pimpinellifolium (Mill.)
area was very likely the source of the cultivated Brezhn. ). Currant tomato. 2n=24. P r i m a r y cen-
tomatoes of the OldWorld and probably of other t r e : Chile, Peru and Ecuador. This red fruited
areas in the NewWorld. The putative ancestor of species is cultivated and occurs as a weed. It
the tomato is probably v a r . cerasiforme (Dun.) c r o s s e s easily with L. esculentum*, of which it
Alef. This variety was originally confined to the may be a subspecies. It is a source of tolerance
Peru and Ecuador area from where it spread in to tomato mosaic virus.
pre-Columbian times as a weed of fields and com-
pound yards throughout much of trop. America, METHYSTICODENDRON AMESIANUM R.E. Schut-
either with or without man's active co-operation. t e s . 2n= . S. America. Cultivated as a medi-
In Mexico it became cultivated because of its s i m i - cinal and witchcraft plant,
larity to another food plant, Physalis ixocarpa*.
Outside its primary gene centre the tomato NICOTIANA RUSTICA L. Aztec Tobacco, Makhor-
plant is self-compatible. In Peru and Ecuador it ka, Nicotine Tobacco. 2n=48. Unknown wild, with
spontaneously crosses with L. pimpinellifolium*. a possible exception of var. pavonii (Dunal) Good-
Tomato flowers pollinated by pollen of L. peruvia- speed. This variety occurs as a ruderal in the
num* result in the induction of parthenocarpic Andes. Aztec Tobacco is a tetraploid having p r o -
fruits. Some F . seeds maybe set resulting in hy- bably originated in Peru by amphiploidization of
brid plants with varying degree of fertility. The apparently N. paniculata L. (2n=24) and N. undu-
Galapagos tomato, ssp. minor Rick (syn. L. m i - lata Ruiz &Pavon (2n=24). Both species occur
nutum Rick, L. cheesmanii Riley v a r . minor wild in Peru. Its cultivation is limited to some
(Hook.) Mill., 2n=24) grows wild on the coasts of areas such as the USSR and India. In most other
the Galapagos Islands. They are characterized by areas it is replaced by N. tabacum* which has a
a very dense pubescence, compound, yellow- low nicotine content.
green leaves, yellow or orange fruit (B-carotene
synthesis), a calyx which expands after fertilization NICOTIANA TABACUM L. Tobacco. 2n=48, g e -
and seeds with a deep dormancy. The plants a r e nome formula SSTT. Goodspeed (1954) showed
very resistant to drought. They a r e eaten by the that tobacco originated by amphiploidization oftwo
Galapagos tortoises and seeds become germina- wild diploid species N. sylvestris Speg. & Comes
tive after passing through the digestive t r a c t s of (2n=24, genome formula S'S') and probably N.
these tortoises (Rick &Bowman, 1961). otophora Grisebach (2n=24, genome formula T'T').
Rick (1971) studied.the geographical d i s t r i - This may have happened in NW. Argentina where
bution of the alleles Ge c , Ge^ and Ge . He found the wild parents a r e found. Clausen (1932), how-
that most European and UScultivars have the ever, suggested that tobacco is a natural amphi-
génotype Ge Ge , only a few have Ge Ge or ploid of N. sylvestris and N. tomentosiformis
Ge Ge . The C. American varieties have Ge n Ge n Goodsp. (2n=24). This is supported by isozymic
and occasionally Ge Ge . In Ecuador Ge is also evidence (Sheen, 1972). The occasionally found
common among the cultivars. The C. American wild tobacco plants a r e escapes of cultivation.
sources of var. cerasiforme have Ge Ge , and an Interspecific crosses have been made to intro-
Ecuador source has Ge . So in Ecuador the culti- duce male sterilizing cytoplasms and genes condi-
vars differ from the wild type, but more sources tioning resistance to diseases.
should be investigated. Sources of L. pimpinelli-
folium* ex Ecuador carried Ge . This would PHYSALIS PERUVIANA L. Cape gooseberry.
suggest gene exchange between L. pimpinellifolium 2n=24, 48. Andes. Cultivated in some S. A m e r i -
and the cultivars. The Peruvian cultivars carry can countries for its b e r r i e s . Often observed as a
Ge ;the same allele is found in Peruvian sources weed or semi-wild.
of L. pimpinellifolium. This suggests gene exchange
too. Rick concluded that the European and US tomato SOLANUM ABANCAYENSE Ochoa. 2n= Peru.
cultivars a r e qualitatively closer related to the Tubers a r e very small and white.
cultivars of Peru, and quantitatively to those from
C. America and Mexico. SOLANUM ACAULE Bitt. 2n=48, genome formula
A„A„A A . Wild tetraploid from C. Peru, Bolivia
LYCOPERSICON HIRSUTUM Humb. &Bonpl. and Nw. Argentina. A parent of S. x juzepczukii*.
2n=24. The western slopes of the Andes in Peru. Frost resistant and resistant to X-virus disease,
SOLANACEAE - SOLANACEAE 159
nematodes and the Colorado beetle. Very suscep- SOLANUM HUMECTOPHILUM Ochoa. 2n=
tible to Phytophthora. Peru. With white-hyaline tubers of 8-12 mm
length.
SOLANUM AJANHULRIJuz. &Buk. 2n=24. Culti-
vated in N. Bolivia (dept. La Paz) and S. Peru. SOLANUM xJUZEPCZUKn Buk. 2n=36. The high
Similar to S. stenotomum* that might be its parent Andes of Bolivia and Peru. Cultivated there for a
species after hybridization with S. x juzepczukii* considerable period of time. Probably a sterile
and other species. It is frost resistant, the tubers hybrid of S. acaule* x S. stenotomum* or S. phu-
a r e long and irregularly shaped. Also resistant to reja*.
virus diseases. S. x juzepczukii may have been formed more
than once, with different varieties of its parents
SOLANUM CHACOENSE Bitt. 2n=24, (36). N. and in each case. This may have resulted in its wide
C. Argentina, Paraguay, Uruguay and S. Brazil. morphological variation. However, its sterility
A very polymorphic wild species. Only once an has prevented its use as a source of frost r e s i s -
autotriploid was observed. Rich in tomatine alka- tance (Hawkes, 1962).
loid which is poisonous to the Colorado beetle.
Introgression between this species and S. SOLANUM MURICATUM Ait. Pepino morado.
microdontum exists in Argentine and possibly e l s e - 2n=24. Unknown wild. Probably domesticated in
where. This resulted in an extension of this origi- the Andes. Cultivated in C. America and S. A m e -
nally low altitude species of open places of the rica. Extremely variable and many types of fruits
Argentinean plain to mountainous region (Hawkes, are recognized. There a r e two closely related
1962). species, either of which could be the parental
species. These a r e S. caripense Humb. & Bonpl.
SOLANUM x CHAUCHA Juz. &Buk. (syn. S. tube- (2n=24) and S. tabanoense Correll (2n=24). Both
rosum group chaucha). 2n--36. This species is a occur in Ecuador and Colombia. Pepino is culti-
hybrid of S. tuberosum* ssp. andigena and S. s t e - vated for its fruits (Heiser, 1964).
notomum* or S. phureja*, but Bukasov (1970)
suggested that it was a triploid derivative of S. SOLANUM NUBICOLA Ochoa. 2n=48. The Huän-
phureja*. The hybridization may have occurred reco region, Peru. A tetraploid species of the
several times and because of the variability of the Tuberosum group. (Ochoa, 1970).
parents this species is very polymorphic. Culti-
vated from C. Peru to N. Bolivia. It has a rather SOLANUM PENNELLII Corr. 2n= . Closely
low yield. related to S. lycopersicoides Dunal. (2n= ).
This species has run wild in Simla hills, India. Both species a r e representatives of a transition
Initially it was established by vegetative propaga- between Solanum and Lycopersicon. It can be
tion. The older the population the more plants crossed with L. esculentum* and it is a source of
flower and the more self-incompatibility breaks resistance to Tomato Mosaic Virus.
down (Nayar &Gohal, 1970).
SOLANUM PHUREJA Juz. &Buk. (syn. S. tubero-
SOLANUM COMMERSONII Dun. 2n=24, 36. E . C . sum group Phureja). Criollo potato. 2n=24, genome
Argentina, Paraguay, Uruguay and S. Brazil. A formula A.A . Cultivated in the most lowlands of
source of resistance to potato canker and Colorado Venezuela, Columbia, Ecuador, Peru and N. Bo-
beetle. livia. The tubers a r e the largest among all diploid
species. The rest period is very short (1-13 days).
SOLANUM CONTUMAZAENSE Ochoa. 2n= It matures early. It is a selection for short tuber
N. Peru. With white-yellow tubers of 15-25 mm dormancy of S. stenotomum*.
length.
SOLANUM QUITOENSE Lam. Naranjillo, Lulo.
SOLANUM x CURTILOBUM Juz. &Buk. 2n=60. 2n-24. Unknown wild. Cultivated for its fruits in
The high Andes of Bolivia and Peru, where it has Colombia and Ecuador. Var. septentrionale R. E.
been cultivated. Probably a hybrid of S. x juzepc- Schultes &Cuatrecasan is spineless (Heiser, 1971).
zukii* x S. tuberosum group andigena. It r e p r o -
duces itself vegetatively, although it is moderately SOLANUM RAPHANIFOLIUM Card. &Hawkes.
fertile. It crosses readily with S. tuberosum. Less 2n=24. The dept. of Cuzco, Peru. There it occurs
frost resistant than its parent S. x juzepczukii as a weed. A stabilized hybrid of S. megistacro-
(Hawkes, 1962). lobum Bitt. (2n=24), and S. canasense Hawkes
(2n=24) (Ugent, 1970a).
SOLANUM GONIOCALYX Juz. &Buk. 2n=24. This
potato is cultivated in C. Peru (dept. Junin). It is SOLANUM SPARSIPILUM Bitt. 2n=24. Peru and
a northern derivative of S. stenotomum. It may Bolivia. A weedy species. Probably a clonal mix-
be included in this species as an extreme variant ture of diploid hybrids of S. stenotomum* and S.
(Hawkes, 1958). Bukasov (1970) suggested that it phureja* and diploid related species like S. cana-
was a derivative of S. multi-interruptum. The sense Hawkes, S. raphanifolium Card. &Hawkes,
tubers have a pale-yellow flesh owing to their rich- and others which Ugent (1970a) grouped in one
ness in caretinoids. They have an excellent fla- complex species S. brevicaule* Bitt. This weedy
vour.
species may form a bridge for a gene flow from mainland). They derive from ssp. andigena intro-
S. brevicaule s . 1 . and S. tuberosum* and vice ductions (Brücher, 1971). Sykin (1971) suggested
versa (Ugent, 1970a). an independent origin of ssp. tuberosum in S.
Chile. In Europe and N. America ssp. tuberosum
SOLANUM STENOTOMUM Juz. &Buk. (syn. S. was also selected from ssp. andigena (Hawkes,
tuberosum group Stenotomum). 2n=24. Cultivated 1958). Simmonds (1968) has 'repeated' this evolution.
at very high altitudes from Peru to N. Bolivia. It The distinction between these subspecies is
may derive from S. brevicaule. It is the parent that ssp. tuberosum has less dissected leaves
species of S. tuberosum ssp. andigena*, S. x with wider leaflets, generally arched and set at a
chaucha*, S. phureja* and S. juzepczukii*. Some wider angle to the stem. The tubers a r e formed
forms a r e frost resistant. The yield and quality under long days, or under short days in the t r o -
is good. pics only at lower altitudes.
The parent species of ssp. andigena is p r o -
SOLANUM TOPm o Humbolt &Bonpland ex Dunal. bably S. stenotomum* and S. x chaucha*, a d i -
Jibara, Uvilla, Cocona. 2n= . S. America. ploid and triploid cultivated species, respectively.
Var. topiro is commonly cultivated for its fruits It is interesting to note that in the Canary Islands
in the Upper Amazon valley. There a r e two fruit cultivar Negra has been cultivated. It is a triploid
forms: ovoid named jibara and globose called (2n=36) (Zubeldia L. et a l . , 1955). Sanudo (1970)
uvilla. The latter has been described as S. alibile suggested that it is a hybrid of S. stenotomum and
R. E. Schultes. A common weed in Ecuador is var. ssp. andigena. Zubeldia L. et al. (1955) believed
georgicum (R.E. Schultes) Heiser (syn. S. georgi- that it was introduced from Peru in the early part
c u m R . E . Schultes). Var. topiro has no spines of 17th Century together with 4x material.
and big fruits, while var. georgicum has spines Run wild potatoes grow in the Kilimandjaro moun-
and small fruits. It is believed that these diffe- tains, in Lesotho and Botswana. They probably
rences a r e a result of domestication. derive from cultivars introduced from Europe.
Artificial interparietal hybrids have been Brücher (1966) described 30 types.
made. It has been suggested that in nature such
hybrids also occur (Heiser, 1971). Sterculiaceae
GUAZUMA GRANDIFLORA G. Don. (syn. Theo-
SOLANUM TUBEROSUM L. Potato. 2n=48. There
broma grandiflora Schum. ). 2n= . Brazilian
a r e two geographical regions where the largest
Amazon basin. A t r e e cultivated for its fruits.
number of the wild and cultivated potatoes grow:
THEOBROMA BICOLOR*
THEOBROMA CACAO L. Cacao. 2n=16, 20, 26.

P r i m a r y gene centre: the area of the 'Upper waters
of the Amazon'. Spread by man. Only in Mexico
was the domestication of the cacao completed by
the Maya. Elsewhere cacao was wild or s e m i -
domesticated. Cuatrecasas (1964) described 22
Theobroma species, which all a r e found in trop.
S. and C. America. Relative isolation of cacao
populations and their original parentage resulted
in the development of two more or less uniform
groups distinguished as Criollo (T. cacao ssp.
cacao) and Forastero (ssp. sphaerocarpum).
The Criollo is located in C. America (Central
American Criollo) and in N. Colombia (South A m e -
rican Criollo). The Forastero can be divided into
Solanum t u b e r o s u m group Andigena (Ugent, 19
the Upper Amazonian Forastero, indigenous to
the Upper Amazon basin and the lower Amazonian
Forastero also named and found in the Guianas
1. C. Mexico (p. 171, 172) and 2. Andes of C. (Cheesman, 1944; Toxopeus, 1969).
Peru, Bolivia and NW. Argentine. The greatest
The Trinitario is a recently originated hybrid
number of tuberous Solanum species is found in
Peru where the potato was probably first domesti- swarm of Criollo from C. America and Amelonado.
cated.
THEOBROMA MICROCARPA Mart. 2n=
There are two subspecies of S. tuberosum: Brazil. Cultivated in Bahia.
1. ssp. andigena (syn. S. tuberosum group Andi-
gena, S. andigena Juz. &Buk. ) and 2. ssp. tube- Thymelaeaceae
rosum (syn. S. tuberosum group Tuberosum).
Ssp. andigena is found in the Andes of Venezuela, FUNIFERA BRASILIENSIS (Raddi) Mansf. 2n=
Columbia, Ecuador, Peru, Bolivia and NW. A r - Brazil. Cultivated in the W. Indies for its fibres.
gentine. It was domesticated in the C. Andes.
Ssp. tuberosum occurred formerly in the coastal
regions of SC. Chile (Island of Chiloé and adjacent
SOLANACEAE - VERBENACEAE 161
Tropaeolaceae
TROPAEOLUM LEPTOPHYLLUM G. Don.
2n= . Ecuador and Peru. Cultivated for its
tubers.
TROPAEOLUM MAJUS L. Nasturtium. 2n=28.

S. America. A herbaceous vine cultivated as an
ornamental plant. The flower buds and young
fruits a r e used for flavouring vinegar. They a r e
also used as capers.
TROPAEOLUM TUBEROSUM Ruiz. &Pav. Tuber

nasturtium. 2n=42.A very old cultivated food
plant unknown wild in Peru, Chile and Bolivia. In
Bolivia it is still cultivated in the mountains above
Lake Titicaca.
Umbelliferae
ARRACIA XANTHORHIZA Bancr. (syn. A. escu-
l e n t a D C ) . Arracacha, Apio a r r a c a c i a . 2n=^
The Andean region of S. America. Cultivated in
Bolivia, Peru, Colombia and Venezuela.
Verbenaceae
LIPPIA CITRIODORA H. B.K. (syn. L. triphylla
(L. 'Hér. ) Kuntze). Lemon verbena. 2n=36. S.
America. Formerly much cultivated for its v e r -
bena oil, now as an ornamental.
11 Central American
and Mexican Centre
The Central American and Mexican Centre has been described by Vavilov as the Central American and
South Mexican Centre of Origin. Darlington and Janaki Ammal (1945) only named Mexico as a centre of
origin, while Darlington (1956) added C. America to Mexico. In this centre agriculture developed in the
7th millenium BC. and therefore Harlan (1971) called it a centre CI Mesoamerican centre.
Old sites of farming have been discovered at Tamaulipas and in the Tehuacan Valley, S. Puebla,
Mexico. To the earliest plant remains belong Amaranthus s p . , Avocado persica, Capsicum annuum,
Cucurbita pepo, C. mixta, Gossypium hirsutum and Lagenaria siceraria.
Only a few but important crops have been domesticated in this region e. g. fruit t r e e s , Agave s p . ,
Capsicum s p . , Cucurbita s p . , Gossypium s p . , Ipomoea batatas, Phaseolus s p . , Zea mays etc.
Agavaceae AGAVE LETONAE F.W. Taylor. Letona, Salva-

dor henequen. El Salvador. Cultivated there for
AGAVE AMERICANA L. Century plant. 2n=60, centuries.
120, (180, 240 and aneuploids). Probably from
Mexico. Cultivated in Mexico, Europe, Africa and AGAVE SISALANA P e r r . (syn. A. rigida Mill. ).
N. America as an ornamental, foliage plant, a Sisal agave. 2n=(c. 138, 147, 149), 150. Mexico
hedge plant and for pulp. and C. America. P r i m a r y centre in Yucatan,
Mexico. Introduced to Florida and from here to
AGAVE ATROVIRENS Karw. (syn. A. latissima most sisal-growing countries.
Jacobi). 2n=150, 180. Mexico. Cultivated there.
AGAVE TEQUILANA Weber. Meczal, Chino Azul.
AGAVE CANTALA (Haw. ). Roxb. Cantala. 2n=90. 2n= . Mexico. Cultivated there.
Probably Mexico. There a wild form occurs on
the western coast. Smaller than the cultivated FURCRAEA GIGANTEA*
types. It was taken to the Philippines and later to
Indonesia where it is cultivated for its fibre. In HESPEROYUCCA FUNIFERA (Koch.) T r e l . (syn.
India cultivated as a hedge and anti-erosion plant Yucca funifera Koch. ). 2n= . Mexico. Cultiva-
(Purseglove, 1972). ted for its leaves which a r e a source of fibre.
AGAVE CRASSISPINA Trel. Maguey manso. POLIANTHES TUBEROSA L. 2n=(50), 60. Tube-
2n= Mexico. Cultivated there. rose. Very likely from Mexico p r e s s l e r , 1953).
Unknown wild. It has probably a long history of
AGAVE DEWEANA T r e l . Zapupe verde, Zapupe domesticated ornamental because of its great v a r i a -
de Tantoyuca. 2n= Cultivated for a long time bility. Spread to other countries where it is used
by the Tantoyuca Indians in Mexico. for perfumery and other purposes. It may derive
from P . gracilis Link. (Mansfeld, 1959).
AGAVE FOURCROYDES Lem. Henequen agave.
2n=c. 140. Yucatan, Mexico. P r i m a r y centre also YUCCA ELEPHANTIPES Regel. 2n= . Proba-
there. Secondary centre probably in E. Africa bly Veracruz, Mexico p r e s s l e r , 1953). Cultiva-
(p. 108). Cultivated in many countries. ted for hedges especially in C. America, where it
was apparantly introduced. The flowers a r e used
AGAVE FUNKIANA Koch. &Bouché. Jaumave, as a vegetable.
Loguguilla. 2n= Mexico. Cultivated t h e r e .
AGAVACEAE - BOMBACACEAE 163
Alstroemeriaceae ANNONA RETICULATA L. Bullock's heart,

Common custard apple, Corazon. 2n=14, (16).
BROMAREA EDULIS (Tuss. ) Herb, 2n= Gene centre lies in the Antilles. Spread to trop.
Mexico to S. America. Apparently very variable.
America and later to other tropical countries.
Probably the species of Bromarea cultivated by
the Mexicans for the edible, tuberous roots, and
as an ornamental p r e s s l e r , 1953). ANNONA SCLERODERMA Safford. Posh té.
2n= . C. America especially from S. Mexico
to Guatemala. Cultivated.
Amaranthaceae
AMARANTHUS CRUENTUS L. 2n=32, 34. Culti- ANNONA SQUAMOSA L. Sweetsop, Sugar apple,
vated in Guatemala and other parts of C. America Custard apple. 2n=14, (16). Gene centre in the
as a grain crop. It evidently derives from A. Antilles. Spread to trop. America. Later it was
hybrldus* (Sauer, 1969). brought to other tropical countries. Atemoya is a
hybrid product with A. cherimoia*.
AMARANTHUS HYBRIDUS L. Slim amaranth.
2n=32. C. America. Cultivated in India and neigh- CYMBOPETALUM PENDULIFLORUM Baill.
bouring regions as a grain crop and as an orna- 2n= . Mexico and Guatemala. A shrub cultiva-
mental. Derived from A. cruentus* (Sauer, 1969). ted for its vanilla-scented petals.
Hybridization with other Amaranthus species and
with species of the genus Acnida has been obser- Araceae
ved (Sauer, 1950).
MONSTERA DELICIOSA Liebm. (syn. Philoden-
dron pertusum Kunth). Ceriman. 2n= . Mexico
AMARANTHUS HYPOCHONDRIACUS L. (syn. A.
and Guatemala. A liane cultivated for its fruits.
leucocarpus S. Wats.). Huauhtli. 2n=32. A main
crop in the Colombian times. Still cultivated in
XANTHOSOMA ROBUSTUM Schott. Pixi, Capota,
Mexico and Guatemala, and also in Asia (India,
Quequesque, Marac, Quiscamote. 2n=26. S.
Iran?). An ornamental in Europe and N. America.
Mexico and C. America. There often cultivated as
Most of the Mexican populations a r e pale-seeded,
an ornamental. The huge leaves are used as u m -
as a r e those of Asia although there a r e more
brellas, and the roots as food. Roots and leaves
black-seeded forms than found in Mexico. The
are also used medicinally and as a stimulant.
plants cultivated as ornamentals in Europe and
N. America are invariably black-seeded (Sauer,
1950). It is evidently derived from A. powellii S. Basellaceae
Wats. (2n=34) (Sauer, 1969). It is also morpholo- BOUSSINGAULTIA CORDIFOLIA*
gically close to A. hybridus*.
Bignoniaceae
Anacardiaceae
CRESCENTIA CUJETE L. Calabash tree, Calabazo,
ANACARDIUM OCCIDENTALE* Cujete. 2n=40. Trop. America. Cultivated in the
tropics. The thin shell of the fruits is used for
CYRTOCARPA PROCERA H.B.K. Chupandilla. containers. In Guatemala three varieties are r e -
2n= . Tehuacän Valley of Mexico. A small cognized. It should not be confused with Lagenaria
fruit t r e e . Archeological seed-remains date from siceraria*.
about 6 500 BC. (Smith, 1968).
PARMENTIERA CEREIFERA Seem. Candle t r e e .
SCHINUS MOLLE* 2n= . C. America. Cultivated in the West
Indies and other tropical regions for its fruits.
SPONDIAS PURPUREA*
PARMENTIERA EDULIS DC. Cuachilota, Food
Annonaceae candle t r e e . 2n=40. C. America. Cultivated there
ANNONA DIVERSIFOLIA Safford. Kama. 2n= and in Mexico for its fruits.
S. Mexico, Guatemala, El Salvador and other
countries of C. America. Cultivated there and in Bombacaceae
Florida. CEIBA PENTANDRA Gaertn. Kapok tree, Silk
cotton t r e e . 2n=72, 80, 88. Toxopeus (1950) b e -
ANNONA MONTANA Macf. Mountain soursop. lieved that the kapok t r e e originated in an area
2n=16, The West Indies. Cultivated. which was later divided by the Atlantic Ocean. So
this species is native both to America and Africa
ANNONA MURICATA L. Soursop, Guanabana, (p. 108). He based hin conclusion mainly on the
Corossol. 2n=14, (16). Gene centre the Antilles. great variability of this plant and on the high f r e -
A small tree cultivated from C. America to the quency of dominant inherited characteristics in
coastal valleys in Peru and elsewhere in the t r o - these two continents. However, Bakhuizen van
pics. It develops the biggest fruits of all Annona den Brink (1933) and Chevalier (1949) thought that
species. Some fruits may weigh 2 kg. seeds may have come from America in prehistoric
times and that later introduction increased the
ANNONA PURPUREA Moc. &Sessé. Soncoya. variability.
2n= . S. Mexico and C. America. Cultivated Its chromosome number suggest a polyploid
there.
CENTRAL AMERICAN AND MEXICAN CENTRE 164
origin and if this supposition is correct the kapok TAGETES ERECTA L. Big marigold. 2n=24, g e -
t r e e can only have arisen in that area where its nome formula AeAe. Mexico. Cultivated as an o r -
parents occur. As all other Ceiba species a r e namental and for its medicinal properties. Also
restricted to America this would also indicate an used in religious rituals and celebrations (Neher,
American origin. 1968). Probably a parent of T. patula*. The genus
The variety found in America and Africa is Tagetes extends from SW. USA into Argentina and
C. pentandra var. caribaea P C ) Bakh. the area of the greatest diversity is in SC. Mexico
(Neher, 1968).
Bromeliaceae
BROMELIA PINGUIN L. Pegwe. 2n=96. W. Indies, TAGETES PATULA L. Marigold, Flor del muerto.
C. America and Venezuela. Aperennial herb cul- 2n=48, genome formula ApApBpBp. Mexico. P r o -
tivated as a living hedge. The fruits are edible. bably originated by hybridization of T. erecta*
and T. tenuifolia Cav. (2n=24, genome formula
Cactaceae BtBt) or closely related species. Cultivated as an
ornamental and for its medicinal properties.
HYLOCEREUS UNDATUS (Haw, ) Britt. &Rose. Spread throughout the world. At one time it was
2n=22. Mexico. Cultivated there and in C. A m e r i - thought to have an OldWorld origin because of the
ca for its edible fruits. sacred role in the Hindu religion (Anderson, 1952).
However, its role might have been promoted by
NOPALEA COCHENILLIFERA (L.) Salm-Dyck. the sacredness of the yellow colour in India.
Nopal. 2n=22. Probably S. Mexico. Cultivated in
trop. America. Convoivulaceae
IPOMOEA BATATAS (L.) Lam. Sweet potato.
NOPALEA DEJECTA Salm-Dyck. 2n= Mexl- 2n=90, genome formula BBBBBB. Unknown wild.
co. Cultivated for its fruits.
Probably derived from I. trifida (H.B.K.)Don.
(2n=90, genome formula BBBBBB), which grows
OPUNTIA FICUS-INDICA (L.) Miller. Indian fig,
wild in Mexico. Related to I. littoralis Blume
Nopal. 2n=22, 88. C. America. Cultivated inthe
(2n=60, genome formula BBBB) which is probably
tropics and subtropics for its fruits.
a tetraploid of I. leucantha Jacq. (2n=30, genome
formula BB). Both species also grow wild in Mexi-
OPUNTIA MEGACANTHA Salm-Dyck (syn. O.
co (Nishiyama, 1963, 1971). Purseglove (1968)
castillae Griffith). Tuna, Nopal. 2n= . Mexico.
suggested that I. trifida might be a weed derived
Cultivated there for its fruits.
from the cultigen. It may also derive together with
the cultigen from a common parent. Yen (1963)
SELENICEREUS GRANDIFLORUS (L.) Britt. & proposed I. tiliacea* as the parental species. Ge-
Rose. (syn. Cereus grandiflorus Mill. ). 2n=22.
nome formula is as yet unknown (Nishiyama, 1971).
Mexico. Cultivated as a source of drug.
This species has its greatest variation in S. A m e -
Caricaceae rica.
Sweet potato was already cultivated in Poly-
CARICA PAPAYA L. Papaya, pawpaw. 2n=18. nesia in pre-Columbian times. Whether it was
Lowlands of C. America somewhere in the region brought there as tubers by man or reached it as
between S. Mexico and Nicaragua. Unknown wild. capsules or on drifting material by sea-currents
The history of its domestication is not known. (Purseglove, 1968) is not yet known. Sweet potato
Papaya has now spread to all tropical countries is cultivated in many tropical countries.
and may have run wild as was observed in the
forest fringes in N. trop. Argentina. Closely r e - IPOMOEA PURGA Hayne (syn. Exogonium purga
lated to C. pelta Hook. &A m . , which also occurs (Wender.) Benth. ). Jalap. 2n=24-28. E. Mexico.
in this area. This species may have contributed Cultivated in that country, the West Indies and
by hybridization (Purseglove, 1968). later India for its medicinal tubers.
CHENOPODIUM NUTTALLIAE Saff. 2n=36. Cultiva- CUCURBITA FICIFOLIA Bouché. Malabar gourd,
ted as a vegetable and a grain crop in C. Mexico. Fig-leaf gourd. 2n=40, (42). Highlands of Mexico
Closely related to Ch. quinoa*. and America. This species might be a derivative
ofC. lundelliana* (Whitaker &Davis, 1962).
Compositae
DAHLIA VARIABILIS Desf. (syn. D. rosea Cav. ). CUCURBITA MIXTA Pang. Pumpkin, Winter
Dahlia. 2n=64. Mexico. A tuberous plant intro- squash, Walnut squash. 2n=40. It probably derives
duced a s a food crop into Europe but it is now from C. lundelliana Bailey in C. America and S.
commonly cultivated as an ornamental. Mexico. It appears that it was widely distributed
in N. Mexico and SW. USAin pre-Columbian times
(Purseglove, 1968). It is a primitive horticultural
PARTHENIUM ARGENTATUM A. Gray. Guayule.
crop with little fruit flesh. It c r o s s e s with C.
2n=36, 54, 72, 108and many aneuploids. Mexico
moschata* and has been described as belonging to
and Texas, USA. Cultivated as a rubber producer.
this species. It developed later than C. maxima*
and C. pepo*.
BOMBACACEAE-GRAMINEAE 165
CUCURBITA MOSCHATA Duch. Cushaw, China divided into sweet cassava (M. dulcis, 2n=36,
squash, Pumpkin, Winter squash. 2n=(24), 40. syn. M. palmata Muell. - A r g . , 2n=36) and bitter
From Mexico to Peru. Domesticated in 1800-1400 cassava (M. esculenta Crantz, 2n=36, M. u t i l i s s i -
BC. (Willey, 1962). Cultivated throughout the ma Pohl, 2n=36). The difference is the content and
world. Whitaker (1962) suggested it to be a d e r i - place of HCN in the tuber. Renvoize (1972) stated
vative of C. lundelliana*. that sweet cassava was probably first domestica-
ted in Meso-America. The sweet types may have
CUCURBITA PEPO L. Marrow, Pumpkin. 2n=(24, derived from the wild population. From Meso-
28), 40, (40-42, 44-46). Whitaker (1962) sugges- America it was taken to S. America. In northern
ted that this species is a possible derivative of S. America the bitter cassava would have been
C. lundelliana Bailey, wild gourd, 2n=40, s p r e a - domesticated (p. 148). (Renvoize, 1972). In Brazil
ding into N. Mexico and SW. USA. In Texas the the diversity increased through intra-species
related wild C. texana Gray is found. This species crosses and by hybridization with wild Manihot
is either a weedy off-spring of C. pepo or may species (p. 148).
have been involved in the latter's formation.
Whitaker and Cutler (1969) observed one seed in Gramineae
a layer dated c. 8750-7840 BC. in a cave in Mexi-
co. AXONOPUS COMPRESSUS (Swartz) Beauv. C a r -
pet g r a s s . 2n=40, 50, 60. C. America and the
Var. ovifera (L. )Alef. is cultivated for its
W. Indies. A perennial tropical grass suitable for
ornamental fruits.
lawns and permanent pasture.
CYCLANTHERA PEDATA Schrad. 2n= . Cul-
ORYZA ALTA*
tivated in Mexico for its young fruits and shoots.
ORYZA LATIFOLIA*
POLAKOWSKIA TACACCO Pitt. Tacaco. 2n-
Costa Rica. Semi-cultivated for its fruits.
ORYZA PERENNIS Moench. 2n=24, genome formu-
SECHTUM EDULE Schwartz (syn. Chayota edulis la AA. For distribution see p. 65. In America
Jacq. ). Chayote, Guisquil, Christophine. 2n=24. var. cubensis (O. cubensis Ekman), the American
Mexico and C. America. It was common among race (2n=24, genome formula AA) of this species
the Aztecs prior to the Conquest. Spread now developed. Gopalakrishnan and Sampat (1966)
throughout the tropics. suggested that O. perennis entered America as a
weed of O. sativa in post-Columbian times.
Dioscoreaceae
PANICUM SONORUM Beal. Sauwi. 2n= . This
DIOSCOREA FLORIBUNDA Mart. &Gal. 2n=36, little-known cereal is cultivated in Mexico. One of
54. S. Mexico and adjacent areas of C. America. the few cereals domesticated in the New World
Cultivated in America to yield sapogenin (Coursey, P r e s s l e r , 1953).
1967).
PANICUM VIRGATUM L. Switch g r a s s . 2n=36,
Ebenaceae 72 and aneuploids. N. and C. America. Cultivated
for cattle food.
DIOSPYROS EBENASTER Retz. Black sapote,
Zapote negro. 2n== . Probably Mexico. Culti-
vated for its fruits. TRIPSACUM DACTYLOIDES (L. ) L. 2n=36, 72
and aneuploids. Observed in USA, C. America,
the West Indies and S. America. It has the largest
Ehretiaceae
distribution of all Tripsacum species. It is c r o s s -
CORDIA DODECANDRA DC. Copte, Siricote. compatible with Zea mays* (Randolph, 1970). The
2n= . Mexico. A tall t r e e cultivated for its genus Tripsacum is related to the genus Zea. It
fruits. is believed that it played a role in the evolutionary
history of cultivated maize (Zea mays) (Randolph,
Euphorbiaceae 1970).
JATROPHA ACONITIFOLIA Mill. (syn. Chidosco-
lus chayamansa McVaugh. ). 2n= . A shrub. TRIPSACUM LANCEOLATUM Rupr. ex Fourn.
Cultivated in the Yucatan area, Mexico. Young 2n=72. Arizona (USA), Mexico and C. America.
shoots and leaves a r e eaten as a potherb. During Hybrids possible between T. pilosus* and this
domestication, forms with fewer stinging hairs species have been described as T. lemmoni Vasey
were selected for (Dressier, 1953). (Randolph, 1970).
MANIHOT ESCULENTA Crantz. Cassava, Manioc, TRIPSACUM LATIFOLIUM Hitchc. 2n=36, 72,
Guatemala, Honduras, Belize and the West Indies.
Manihot, Yuca. 2n=36. Unknown wild. There a r e
Cultivated for forage. Typical 4x forms a r e found
two geographical centres: in W. and S. Mexico, in W. Mexico.
and C. America, parts of Guatemala and in NE.
Brazil (Rogers, 1963). He suggested that cassava
could have arisen in both these centres, because TRIPSACUM LAXUM Nash. 2n=(54), 72. Mexico,
there is no reason to exclude root domestication in the West Indies, C. and S. America. Only cultiva-
the Mexican seed type agriculture. Cassava can be ted or has escapes from cultivation. Sterile and
CENTRALAMERICANANDMEXICANCENTRE 166
maize i. e. teosinte was cultivated.

From C. America maize reached S. America
where its development became more advanced
(p. 150). Advanced varieties were returned to C.
America. For Mexico they have been described
as Pre-Columbian Exotic Races comprising of 4
varieties; Cacahuacinthe, Harinoso de Ocho, Olo-
tdn and Maize Dulce (Wellhausen et a l . , 1952).
Where they hybridized with primitive Mexican
Ancient Indigenous Races: Palomero Toluqueno,
Arrocillo Amarillo, Chapalote and Nal-Tel. This
resulted in a new group of varieties called: P r e -
historic Mestizos. Introgression with teosinte
most likely has taken place too. Wellhausen et al.
(1952) described 13 r a c e s .
T r i p s a c u m latifolium ( . . . ) , T. lanceolatum ( ) and T. laxum
Modern incipient Races have developed since
( — ) (Randolph, 1970) the Conquest. Wellhausen et al. (1952) described
four of them. Some being very recently developed.
The same development took place in Guatemala
hence propagated vegetatively for forage. Some (Wellhausen et a l . , 1957).
variation is observed which suggests some p r o - From C. America maize also spread to N.
duction of viable seed (Randolph, 1970). Sponta- America (p. 176).
neous mutations might be another source, perhaps, De Wet et al. (1972) suggested that the ' t r i p -
the only one, of this variation. sacoid' races of maize in S. America were origi-
nally introduced from Mexico or C. America where
TRIPSACUM MAIZAR Hernandez &Randolph. they inherited their 'tripsacoid' characteristics
2n^36, (72). SW. Mexico. A typical 2x and 4x through introgression with teosinte. It is also
forms occur elsewhere in Mexico and Guatemala. possible that these races represent relics which
Probably one of the parents of the allotetraploid retain some original teosinte-like characteristics
Tripsacum species (Randolph, 1970). inherited from the maize progenitor.
The oldest domesticated maize varieties had
TRIPSACUM PILOSUM Scribn. & Merr. 2n=72. a string (slender) cob. This primitive characte-
Mexico and C. America. ristic is still found in some relict cultivars like
Confite Morocho, which is the most primitive
TRIPSACUM ZOPILOTENSE Hernandez &Ran- living cultivar (Galinat, 1969). It comes from
dolph. 2n= . SW. Mexico. Probably one of the Peru. The domesticated recessive character thick
parents of the allotetraploid Tripsacum species. cob is conditioned in the Corn Belt dent by 3 major
This region is near to the centre of diversity of loci. One allele derives from northern flint which
the genus Tripsacum which is in S. Mexico and obtained it from Maize de Ocho. It is possible that
neighbouring Guatemala (Randolph, 1970). this allele introgressed from Tripsacum sp. proba-
bly T. dactyloides*. Perhaps this introgression
ZEA MAYS L. Maize, Corn. 2n=20. The 'origin' could be traced to S. America by way of the culti-
of maize has been subject to much discussion and vars Cabuya and Sabanero (Galinat, 1969). How-
research. The present conclusion is that teosinte ever, de Wet et al. (1972) believed that no intro-
(Z. mexicana*) is the wild parent of maize (de Wet gression exists between maize and Tripsacum
et a l . , 1971; de Wet &Harlan, 1972; Galinat, species. So cv. Chococena is not a hybrid of cv.
1971; J. G. Waines, see Galinat, 1971). Other Confite (ex Peru) and a Columbian Tripsacum as
theories were that maize derived from hybridiza- has been suggested.
tion of primitive maize, teosinte and Tripsacum The source(s) of other two recessive alleles
ssp. when the first reached C. America from S. is not fully understood. One of these alleles p r o -
America. If so wild maize - a podcorn - tunicata duces high condensation of staminate spikelets in
Sturt. - would have been extinct by now. This the tassel branches and the other increases
complete disappearance of wild maize would have tassel branching. If the first allele is absent the
been caused by a number of factors such as intro- expression of the second is complete resulting in
gression of genes of domesticated maize, the use profuse branching like the mutant ramosa. Some
of the habitats of wild maize for maize cultivation cultivars of the southern dent and 'bear paw' pop-
and the grazing of Old World animals. corn appear to have this high condensatioii-ramosa
The earliest finds of maize were tiny cobs type of thick cob. The degree of fasciation with
(2-3 cm) in the Bat Cave of Tehuacan, Mexico. which this type of thick cob may be associated
They have been dated about 3 600 BC. Two types seems to have been modified by teosinte intro-
have been observed; 1. a. podcorn and 2. a pop- gression, teosinte gene(s) suppressing fasciation.
corn - everata Sturt. (syn. praecox) (MacNeish, It is suspected that the Corn Belt dents obtained
1964; Mangelsdorf et a l . , 1964). In another cave, these two pairs of recessive alleles from the
early Bat Cave-like maize and teosints have been southern dents (Galinat, 1969).
identified. The material dates from about 2 200 BC. Other morphological changes due to domesti-
Early tripsacoid maize dates from 2 300-1 500 BC. cation a r e a development of a complete husk cove-
MacNeish (1964) supposed that about 5 000 BC wild rage of the mature ear, the development of female
GRAMINEAE - LABIATAE 167
inflorescense, a reduction of the glumes of the teosinte to maize, while that of maize to teosinte
female inflorescense, an arrangement of spike - is very small (see p. 166). This may happen b e -
lets in a higher row number, a development of the tween the earliest flowering teosinte plants and
cupules and an increase of the length of the styles the latest of maize. Teosinte plants may grow un-
(silk). Some cultivars have an ear length up to 45 noticed in a maize field and may be harvested t o -
cm. Hybrid maize varieties may produce more gether with its leader crop. Seeds may be spread
than 1 000 kernels per cob while the Tehuacan either during the transport or storage of the crop
maize has about 40 kernels per cob. or in manure (Wilkes, 1967). Attempts have been
The terminal inflorescense becomes entirely s t a - made to cultivate teosinte as a fodder, but it
minate being a lax plume with waving branches yields less than sorghum.
(Galinat, 1969).
A flow of teosinte genes to maize still
exists where maize cultivation is primitive and
teosinte is present. Maize x teosinte hybrids a r e
actually cultivated. Maize may show pronounced
signs of 'tripsacoid' i . e . teosinte germ plasm
such as induration of the lower glume and a
straight rigid ear.
Less genes flow from maize to teosinte since
the genetic incorporation of a maize-like rachis
results in the inability to disperse seed and so to
the extinction of teosinte introgressed with maize
(Wilkes, 1970). Extensive gene exchange in both
directions is evident around Chalco, S. of Mexico
City, where the weedy teosinte race mimics the
local race of maize in size, colour and pubes-
cence. These weeds remain teosintoid with r e s -
pect to female inflorescence structure. In many
other areas of Mexico, particular the Rio Balsas Zea mexicana (Wilkes, 1967)
Valley on the W. escarpment, W. of Mexico City,
teosinte behaves essentially as a wild grass, but
modern development leads to an increased infil-
tration of maize genes into teosinte (de Wet, p e r s . Iridaceae
comm. 1971). TRIGIDIA PAVONIA (L.f. ) DC. Cacomite, Tiger
Several types of maize are cultivated. An flower. 2n=26, 28. Mexico. Naturalized in most
improved popcorn is being cultivated in USA, of C. America, Colombia, Bolivia, Peru and B r a -
Mexico and elsewhere. Softcorn - amylacea zil. Easily cultivated. Soon escapes into maize
Sturt. - predominates in the Andean region. Flint fields etc. as a weed. The Aztecs cultivated this
maize, flint corn - indurata Sturt. - predominates species for almost a 1 000 y e a r s . Cultivated now
in N. Colombia and Eastern S. America. Sweet as an ornamental which resulted in spontaneous
corn - saccharata Sturt. (syn. rugosa Bonof. ) - variations in colour and size (Molseed, 1970).
was cultivated for the preparation of South A m e r i -
can and Mexican beer. At present it is mainly cul-
Juglandaceae
tivated in USA. Waxy maize - ceritina Kulesh.
cultivated in the Americas and in E. Asia. Dent CARYA PECAN (Marsch. ) Engl. &Graebn. Pecan.
maize - indentata Sturt. is the main type of the 2n=32. N. Mexico. Cultivated there. It resembles
Corn Belt of USA and N. Mexico. A hybrid of a the walnut, Juglans regia* but the seed has a
late-maturing dent Gourd Slide cultivated in the better taste.
south, and an early maturing flint maize, mainly
cultivated in the north. The latter derives from JUGLANS MOLLIS Engelm. Guatemala walnut.
Maiz Ocho. Mexico and Guatemala. Similar to the walnut (J.
From C. and S. America maize was taken to regia*).
Europe (p. 99), Asia (p. 48) and Africa (p. 85).
where secondary centres of diversity developed. Labiatae
At present flint maize, flint corn - indurata HYPTIS SUAVEOLENS Poit. 2n=28, 32. Cultivated
Sturt. - is quite common in C. America. mainly in Mexico. A variable and weedy plant now
Derivatives of Z. mays x Z. mexicana are occurring in many parts of the tropics.
used for fodder. These are called maisinte (Pra-
sad &Chaudhuri, 1968). SALVIA CHIA Fern. Chia. 2n= . Mexico.
Seeds a r e used to prepare a beverage and for pain-
ZEA MEXICANA (Schrad. ) Kuntze (syn. Euchlaena ting or medicine.
mexicana Schrad. ). Teosinte. 2n=20. SW. Chihua-
hua, Mexico to S. Honduras. MacNeish (1964) SALVIA DIVINORUM Epling & Jativa-M. 2n=
suggested that about 5 000 BC wild maize i. e. t e o - Wild unknown. Cultivated in NE. Oaxaca, Mexico.
sinte was cultivated. It is the ancestor of maize, Vegetatively propagated (Schultes & Hofmann,
Zea mays*. Owing to natural hybridization b e - 1973). Perhaps one clone. Closely related to S.
tween maize and teosinte there is a gene flow from cyanea Lindl. (2n= .) of C. Mexico.
Lauraceae LEUCAENA LEUCOCEPHALA. Leucaena.

PERSEA AMERICANA Miller. Avocado. 2n=24. 2n= . C. America. Spread to the Caribbean
Mexico and C. America. There a r e three geo- islands, the Philippines, SE. Asia and elsewhere.
graphical r a c e s : (1) Mexican (also named P. a m e - Used as a shade tree, for cattle food and to con-
ricana var. drymifolia Mez. syn. P . drymifolia trol soil erosion.
Cham. &Schlecht. ) from the Mexican highlands
where wild progenitors have been found. Its anise - MIMOSA INVISA Mart. 2n=24, 26. C. and S.
scented leaves, hardiness and small fruits are America. In Java and elsewhere it is used as a
characteristics. (2) Guatemalan, from the Guate- cover and green manure. The spiny stem is a
malan highlands. Wild progenitors have been found disadvantage. Var. inermis Adelb. is a useful s e -
in this area, (3) West Indian, from the Guatemalan lection.
lowlands which has only spread to the West Indies
in post-Columbian times (Purseglove. 1968). PACHYRHIZUS EROSUS (L.) Urban, (syn. P .
Where these races grow together - either in their angulatus Rich, ex DC., P . bulbosus (L. ) Kurz. ).
native region or elsewhere - hybrids originate Yam bean, Jicama. 2n=22. Mexico and N. C.
(Bergh, 1969). America. Cultivated there in pre-Columbian t i m e s .
Taken to Asia and cultivated there. In S. China and
PERSEA SCHIEDEANA Nées. Coyo avocado. Thailand it has run wild (Clausen, 1944). Var.
2n= . From Guatemala to Mexico. Cultivated palmatilobus (DC. ) Clausen is probably P . palma-
on a small scale - chiefly near Orizaba in Mexico tilobus Benth. &Hook.
for its fruits (Bergh, 1969).
PHASEOLUS ACUTIFOLIUS Gray. var. latifolius
Leguminosae Freeman. Tepary bean. 2n=22. Mexico. Texas and
Arizona. A very polymorphic species especially
CALOPOGONIUM MUCUNOIDES Desv. 2n= characterized by its drought-resistance. The c u l -
Trop. America. A cover crop and green manure. tivated variety is var. latifolius.
Cultivated in the tropics where it has naturalized.
PHASEOLUS COCCINEUS L. Scarlet Runner,
CANAVALIA CAMYLOCARPA Piper. Babricon Runner bean. 2n=22. Mexico and Guatemala. Do-
bean. 2n= . The West Indies. Cultivated as a mesticated t h e r e . Cultivated in the Americas and
green manure. Eurasia for food, for fodder and as an ornamental.
For Ph. vulgaris* it is a source of halo blight
CROTALARIA LONGIROSTRATA Hook. &A m . resistance, absence of parchment and string.
Much of Mexico and C. America (Dressier, 1953). Smartt (1973) suggested that the cultigen ssp.
A large herb cultivated in Guatemala as a potherb. darwinianus Hdz. X. &Miranda C. may have an
independent domestication of a yet unknown an-
DESMODIUM TORTUOSUM (SW.)DC. (syn. Mei- cestral form. He stated that a special taxonomie
bomia purpurea (Mill. )Vail. ). Florida clover. treatment may be necessary.
Giant beggarweed. 2n=22. C. America, Florida,
West Indies and northern S. America. A perennial PHASEOLUS LUNATUS L. Sieve bean, Lima bean.
herb used as a green manure and forage crop. 2n-22. C. America and in the Andes from Peru to
Argentine. Kaplan (1965) showed that the small
DESMODIUM UNCINATUM* lima bean of Mexico may have arisen in the P a c i -
fic coastal foothills of Mexico and that the big lima
ENTEROLOBIUM CYCLOCARPUM (Jacq. ) Griseb. bean of Peru was first domesticated in the Andean
2n=26. Mexico, C. America, N. of S. America highlands (p. 153).
and West Indies. Cultivated as a shade t r e e .
PHASEOLUS RETUSUS Benth. Metcalf bean.
INGA DULCIS. 2n= . Mexico. Used as a shade 2n= . Texas, Arizona and New Mexico, USA.
tree and hedge plant. Occasionally cultivated.
INGA EDULIS Mart. Food inga. 2n=26. Mexico PHASEOLUS VULGARIS L. Common bean. 2n=22.
and C. America. Used as a shade t r e e . Kaplan (1965) suggested a multiple domestication
within C. America from a widespread and poly-
INGA GOLDMANH Pittier. 2n= . C. America. morphus species. Willey (1962) suggested that
Used as a shade t r e e . this domestication may have taken place between
5 000 and 3 000 BC. From C. America it would
INGA LAURINA (Sw.)Willd. Sackysac. 2n= have spread to other parts of America (Kaplan,
C. America and the West Indies. It has edible 1965). However, Heiser (1965) believed in an in-
fruits. Used as a shade tree in coffee plantations dependent domestication from the closely related
in the New World (Purseglove, 1968). Ph. arborigineus Burkart which occurs wild in
Peru. It is quite possible that when Ph. vulgaris
INGA LEPTOLOBA Schlecht. 2n= . Mexico, reached Peru it introgressed with Ph. arborigi-
C. and S. America. Used as a shade t r e e . neus. Similarly in other a r e a s introgression of
related wild material may have taken place.
INGA PITTIERI Micheli. 2n= C. America. Recently Gentry (1969) pointed out that a wild type
Used as a shade t r e e . of Ph. vulgaris is growing in C. America. From
LAURACEAE - MORACEAE 169
GOSSYPIUM HARKNESSH Brandagee. 2n=26, g e -

nome formula D „D„_„. The islands and coasts
of Gulf of California.
GOSSYPIUM HTRSUTUM L. Upland cotton. 2n=52,

genome formula (AADD).. The current theory is
that this species originated in C. America. H a r -
land (1970) suggested that its centre of origin is in
S. America (p. 154), while C. America is an i m -
portant secondary centre of diversity. Upland
cotton is cultivated in USA, in Africa except for
the Nile Delta (p. 154), in C. Asia, India (Cam-
bodia type, p . 51), S. America, SE. China,
Indochina and elsewhere. In C. America and the
West Indian islands race marie galante is found
Phaseolus vulgaris (Gentry, 19 while race punctatum is found around the coasts of
the Gulf of Mexico from Florida to Yucatan in the
Bahamas and on some West Indian islands. It was
this wild type the cultivated forms derive. taken to W. Africa where it spread in the zone
Ph. arborigineus was reduced by Burkart and south of the Sahara, to Réunion, the Malabar coast
Bücher (1953) to a subspecies level of Ph. vulgaris. of India, Polynesia, the Marquesas, Fiji and N.
They suggested that this subspecies was not taken Australia. A great diversity was found in N.
to S. America. A study of the wild and cultivated Australia.
beans of this area should clarify whether this is Race yucatense is probably a cotton that has
so or whether ssp. arborigineus is an escape of run wild and is now naturalized into natural vege-
early cultivated forms (Gentry, 1969). Its p r i m a - tation of the coastal sand dunes of the Progeso
ry centre of diversity is in Mexico. Here intro- area, Mexico.
gression between cultivated and wild Ph. vulgaris Race morilli is found in Oaxaca, Peubla and
and P . coccineus* occurs (Wall, 1970). Morelos, C. Mexico. A perennial cotton with a
Secondary centres in Eurasia (p. 34). bushy form and broad, intensely hairy leaves.
Race palmeri in the state Guerrero, Mexico.
PITHECELLOBIUM DULCE Benth. Manila t a m a - It has deeply dissected leaves and strong antho-
rind. 2n=26. Mexico to N. of S. America. The cyanin pigmentation.
arillus is edible. A t r e e planted in hedges. Race latifolium is found in Chiapas, Mexico
and neighbouring regions of Guatemala. An annual
SOPHORA SECUNDIFLORA (Ort. ) Lagasca ex De cotton. Throughout its t e r r i t o r y a small-fruited
Candolle. Mescal bean, Red bean, Coral bean. form is found. A large-fruited form grows in the
2n=18. N. Mexico to Texas and New Mexico. Used vicinity of Acala, Chiapas. This race appears to
as hallucinogen. Often planted as an ornamental. be the foundation stock of all the annual G. h i r s u -
tum cottons (Hutchinson, 1962).
TEPHROSIA SINGAPOU (Buc'hoz) A. Cheval,
(syn. T. toxicaria (Sw.) P e r s . ). 2n=22. Mexico GOSSYPIUM KLOTZSCHIANUM Anderson var.
and C. America to Peru and N. Brazil. Cultivated davidsonii (Kelloggs) Saunders. 2n=26, genome
as a fish poison. formula D 3 _ D D„ Shores of the Gulf of Califor-
nia and the Revflla Gidego islands. Related to the
Malpighiaceae plants of this species found on the Galapagos I s -
lands (p. 154).
BUNCHOSIA COSTARICENSIS Rose. 2n=
Costa Rica. Cultivated for its fruits. GOSSYPIUM LOBATUM Gentry. 2n=26, genome
Mexico.
MALPIGHIA URENS L. Barbados cherry. 2n=
The West Indies. Cultivated for its fruits. GOSSYPIUM THURBERI Todaro. 2n=26, genome
formula D-.D.. Arizona, USA and Sonora and SW.
Malvaceae Chihuahua, Mexico. At one time it was thought to
GOSSYPIUM ARIDUM (Rose &Standley) Skovsted. be the American parent of G. herbaceum* and G.
2n=26, genome formula D4D4. Mexico. barbadense*.
GOSSYPIUM ARMOURIANUM Kearney. 2n=26, GOSSYPIUM TRILOBUM (Moc. &Sess. ex DC.)

ge- Skovsted. (syn. Ingenhouzia triloba Moc. &Sess.
nome formula D„ . D , , . San Marcos Islands.
Gulf of California^ ex D C ) . 2n=26. C. Mexico (Fryxell & Parks,
1967).
GOSSYPIUM GOSSYPIOIDES (Ulb. ) Standley.
2n=26, genome formula D g D fi . Mexico. It c r o s s e s Moraceae
poorly with most of the species of the D genome. CASTILLA ELASTICA Cerv. Arbol del Hule. 2n=28.
Its seed-protein pattern is different from the D S. Mexico and C. America. Cultivated in C. America,
genome species. However, it is similar to the Trinidad and Tobago, and Java at the end of the 19th
patterns of G. klotzschianum* (Cherry et al. ,1970). Century. Now replaced by hevea rubber.
CENTRALAMERICANANDMEXICANCENTRE 170
Myrtaceae 2n=100. SW. of USA and adjacent Mexico. A p e -

PIMENTA DIOICA (L. ) Merr. (syn. P. officinalis rennial herb occasionally cultivated.
Lindl.). Allspice. Pimento. 2n^22. The West
Portulacaceae
Indies and C. America. Spread to other countries.
CLAYTONIA PERFOLIATA Donn. ex Willd. (syn.
PSIDIUM FRIEDRICHSTHALIANUM (Berg. ) Nied. Montia perfoliata How. ). Winter purslane, Miner's
Costa Rican guava. 2n= . C. America. A lettuce. 2n=36. N. America and Mexico. Cultiva-
small tree cultivated for its acid fruits. ted as a vegetable.
PSIDIUM GUAJAVA L. Guava. 2n-22. Trop. Rosaceae

America. Cultivated there. In 1526 Oviedo r e p o r -
ted that improved forms were cultivated in the CRATAEGUS PUBESCENS (H.B.K. ) Steud. (syn.
West Indies. Spread through the tropics, where C. stipulosa (H.B.K. ) Steud. ). 2n=34. A t r e e
guava may be found naturalized. widely cultivated in Mexico and Guatemala for its
fruits.
PSIDIUM MOLLE Bertol. Guisare. 2n= . S.
Rutaceae
Mexico and C. America. Cultivated for its fruits.
CASIMIROA EDULIS La Llave &Lex. White sapo-
PSIDIUM SARTORIANUM (Berg. ) Nied. Pichiché, te, Zapote Blanco. 2n=36. Highlands of Mexico
Arrayan, Guayabillo. 2n= . Mexico. Cultivated and C. America. A fruit t r e e introduced to other
there. subtropical countries.
Onagraceae CITRUS PARADISI Macf. Grapefruit. 2n=18. Un-

OENOTHERA BIENNIS L. (syn. Onagra biennis known wild. Closely related to C. grandis and it
Scop.). Evening primrose. 2n=14. N. America to probably is a bud mutation or a hybrid product of
Mexico. Run wild over a large part of the world. C. grandis and sweet orange (C. sinensis*). This
Cultivated as a fodder crop. must have occurred in the West Indies some time
before 1750 (Purseglove, 1968). It is widely cul-
Orchidaceae tivated in the (sub)tropics.
Hybrids with other Citrus-species have been
VANILLA FRAGRANS (Salisb. ) Ames. (syn. V. obtained, Sopomaldin is a hybrid with C. mitis
planifolia Andrews). Vanilla. 2n=(28-31), 32. C. (Calamondin), Siamelo with C. reticulata* (King
America, SE. Mexico and the Antilles. A perenni- Orange), Tangelo with the same species var. deli-
al vine cultivated in the tropics and S. Mexico for ciosa (Tangerine), Satsumelo with the same s p e -
its aromatic fruits. cies (Satsuma) and Chironja with C. sinensis*
(Sweet Orange). Tangelolo is a hybrid of g r a p e -
VANILLA POMPONA Schiede (syn. V. grandiflora fruit with Tangelo (see above).
Lindl.). West Indian vanilla. 2n=32. SE. Mexico,
C. America and N. of S. America. A perennial Sapotaceae
vine on Tahiti, Martinique and Guadeloupe for its
aromatic fruits. CALOCARPUM SAPOTA (Jacq. ) Merr. (syn. C.
mammosum P i e r r e , Achras mammosa L. ).
Palmae Mamey sapote, Sapote, Marmelade plum. Mamey
Colorado. 2n= . C. America. A tree cultivated
CHAMAEDOREA TEPEJILOTE Liebm., 2n=32, in the tropics for its fruits.
Ch. wendlandiana (Oerst. ) Hemsl. 2n= . At
least one, and probably several, species of this CALOCARPUM VIRIDE Pitt. Green sapote.
genus a r e cultivated in S. Mexico and C. America. 2n= . Guatemala to Costa Rica. At r e e culti-
The young staminate flower clusters are used as vated for its fruits.
a vegetable (Dressier, 1953).
CHRYSOPHYLLUM CAINITO L. Star apple. 2n=52.
GUILIELMA UTILIS Oerst. 2n= . C. America. West Indies and C. America. The pulp is edible.
A palm cultivated there. Also an ornamental.
Papaveraceae LUCUMA BIFERA Mol. Egg fruit. 2n= Chile

ARGEMONE MEXICANA L. Mexican prickly poppy. and Peru. Cultivated there for its fruits.
2n=28. SW. of USA and Mexico. Cultivated in Mali.
Seeds are used to prepare oil. Also an ornamental. LUCUMA SALICIFOLIA H. B.K. (syn. Pouteria
campechiana (H.B.K. ) Baenhi). Yellow sapote,
Polygonaceae Zapote amarillo. 2n= . Mexico and C. America.
Cultivated for its fruits.
COCCOLOBA UNIFERA L. Seaside grape. 2n=
Trop. America. A shrub or t r e e cultivated for its MANILKARA ACHRAS (Mill.) Fosbérg (syn.
edible fruits. Achras zapota L., M. zapotilla (Jacq.) Gilly).
Sapodilla, Chiku. 2n=26. Mexico and C. America.
RUMES HYMENOSEPALUS Torr. Canaigre, Wild Cultivated now in the tropics for its fruits and gum
rhubarb, Pie dock. Sour dock, Tanner's dock. (chickle) for chewing-gum production.
MYRTACEAE -SOLANACEAE 171
Simaroubaceae as S. canasense Hawkes, S. brevicaule Bitt. s . S . ,

S. raphanifolium Card. &Hawkes, S. leptophyes
SIMAROUBA GLAUCA DC. Aceituno. 2n= Bitt., S. soukupii Hawkes, S. multiinterruptum
S. Florida to Costa Rica. In El Salvador and Bitt., S. abbottianum J u z . , S. liriunianum Card.
elsewhere attempts are made to cultivate it as an &Hawkes, S. ochoae Vargas, S. multidlssectum
oil crop. Hawkes, S. spegazzinii Bitt. and S. vidaurrei
Cardenas. It is likely that from this complex
Simmondsiaceae species the diploid S. stenotomum* arose.
SIMMONDSIA CALIFORNICA Nutt. Jojoba, P i g -
nut, Goatnut. 2n= . California and adjacent SOLANUM BULBOCASTANUM Dun. 2n=24, g e -
Mexico. A preliminary domestication was done in nome formula BB, (36, BBB). At medium altitudes
California. The product is a liquid wax, jojoba from C. Mexico to Guatemala in grassland, waste
oil. places and forest glades and clearings. There are
three subspecies (Hawkes, 1966):
Ssp. Bulbocastanum (2n=24, 36) is found in
S. Mexico, ssp. dolichophyllum (2n=24) in the
Mexican states Morelos and Cuerrero and ssp.
partitum (2n=24) in S. Mexico and Guatemala.
This B genome also constitutes two of the
genomes of S. polytrichon* AÂ^BB. The B g e -
nome is related to the A 4 genome and to the A^
genome of S. phureja*. S. bulbocastanum crosses
with S. tuberosum* and is used as a source of
resistance to Phytophthora, X-virus, Y-virus,
Colorado beetle and several aphids (vectors of
virus diseases).
SOLANUM CARDIOPHYLLUM Lindl. 2n=24, 36.

S. Mexico. In dry stony grassy and waste places,
often as a weed in maize and bean fields. Ssp.
Simmondsia californica (Gentry, 1958) ehrenbergii is found in NC. to W. Mexico, ssp.
cardiophyllum occurs in C. Mexico and ssp. lan-
ceolatum in SE. to S. Mexico. Triploid forms are
Solanaceae frequent in ssp. cardiophyllum. Ssp. ehrenbergii
is one parent of S. sambucinum*.
CAPSICUM ANNUUM L. Bell pepper, Cayenne
pepper, Mexican chili. 2n=24. Wild variety in SOLANUM CLARUM Corr. 2n=24. Guatemala, in
Southern USA, West Indies, Mexico, C. America the high mountains. Probably a link between Bul-
and Colombia. P r i m a r y centre in Mexico. Secon- bocastana and Morelliformia s e r i e s .
dary centre centres in Europe (p. 142) and Asia
(p. 70). Originally the cultivars were limited to SOLANUM DEMISSUM Lindl. 2n=(36, 48), 72,
C. America. The wild bird pepper (C. annuum genome formula A]A,A«A.BB. NW. Mexico to
v a r . minimum) is the most probable ancestor of Guatemala. An important source of disease r e s i s -
the cultivated varieties (var. annuum). It is a very tance (Phytophthora, X virus, Y virus). The A^-
polymorphic species. genome may have come from S. verrucosum*.
One of the parental species of S. x edinense* and
CAPSICUM FRUTESCENS L. Tobasco pepper. S. x semidemissum.
2n=24. This species consists of wild types and
derived cultivars, like the Tobasco peppers. SOLANUM x EDINENSE Berthault. Mexican weed
Widespread as a weed and as cultivars in Mexico, potato, Papa morado. 2n=60. Clones occur in and
C. America and lowland S. America. It might be along the edges of cultivated field, irrigation
the parental species of C. chinense*. ditches, roadsides thickets and forest fringes in
the Central Volcani Cordillera of Mexico between
PHYSALIS rXOCARPA Brot. Tomatl, Miltomate, 2 000 and 3 500 m. A source of Phytophthora and
Husk tomato, Tomatillo. 2n=24. Mexico. There frost resistance. This 5x hybrid is a cross b e -
and in Guatemala this vegetable is cultivated. tween S. tuberosum (group Andigena, 2n=4x=48)
and S. demissum Lindl. (2n=6x=72) (Hawkes, 1966).
SOLANUM AGRIMONIFOLIUM Rydb. 2n=48. From S. demissum genes may introgress in the cultiva-
C. Mexico to Bolivia. ted potato by backcrossing. Harvest of potato may
contain tubers of S. x edinense and by trading the
SOLANUM BRACHISTOTRICHUM (Bitt. ) Rydb. potato this weedy potato is also spread (Ugent,
2n=24. NW. Mexico, in open pine and juniper fo- 1967).
r e s t s and amongst bushes and rocks. Resistant to
the green peach aphid, Myrus persicae Sulzer. SOLANUM GUERREROENSE Correll. 2n=72, g e -
nome formula A.A..A A.BB. SW. Mexico. The
SOLANUM BREVICAULE Bitt. 2n=24. Ugent
A. genome may nave come from S. verrucosum*.
(1970a) grouped in this species wild diploid species
SOLANUM HJERTINGII Hawk. 2n- . NE. may block introgression of S. demissum into the
Mexico, in pinon scrub and cultivated fields. Very potato.
similar to S. fendleri*. A source of resistance to
the potato aphid, Macrosiphum euphorbiae Thomas. SOLANUM STOLONIFERUM Schlechtd. &Bouché.
2n=48, genome formula A . A . B B . N. Mexico. A
SOLANUM HOUGASII Corr. 2n-72. genome for- source of resistance to Y virus, A virus, Phytoph-
mula AiAjA 4 A 4 BB. WC. Mexico. The A, genome thora, Pseudomonas and the potato aphid, Macro-
may have come from S. verrucosum*. siphum euphorbiae Thomas. One of the parents of
S. x vallis-mexici*. One genome is the same as
SOLANUM IOPETALUM (Bitt. ) Hawk. 2tt=72, g e - that of S. acaule*.
nome formula A T AAT A . A ^ B B W. and S. Mexico.
n v
It includes S. brachycarpum Corr. The Aj g e - SOLANUM x VALLIS-MEXICI Juz. 2n=36, genome
nome may have come from S. verrucosum*. formula AjA^B, 60, A ^ A ^ B , 72, A 1 A 1 A 4 A 4 BB.
The Valley of Mexico. A natural hybrid of S. s t o -
SOLANUM JUGLANDIFOLIUM Dunn. 2n=24. Cos- loniferum* and diploid S. verrucosum*. Amphi-
ta Rica. Little value to potato breeders because plolds have also been found. Seed collected from
it does not bear stolons or tubers. 5x plants produced aneuploid plants resembling
either S. stoloniferum* or S. demissum*. Plants
SOLANUM LEPTOSEPALUM Correll. 2n= of the first group had 2n=47-55, while those of the
NE. Mexico and possibly in USA. second group had 2n=61-69. Intercrossing between
the hybrid, the aneuploids and the species could
SOLANUM LESTERI Hawkes &Hjerting. 2n=24. account in part, for the extensive polymorphism
Oaxaca, Mexico. found in S. stoloniferum and S. demissum popu-
lations (Marks &Montelongo-Escobedo, 1970).
SOLANUM MICHOACANUM (Bitt. ) Rydb. (syn.
S. trifida Corr. ). 2n=24. Michoacan and Jalisco, SOLANUM VERRUCOSUM Schlechtd. 2n=24, g e -
Mexico. In the pine forests and fields. Resistant nome formula AjA,, (36, 48, 72). NE., C. and
to the green peach aphid, Myrus persicae Sulzer. S. Mexico. Also cultivated there. The tubers have
a good flavour (Abdalla &Hermsen, 1973). A
SOLANUM MORELLIFORME Bitt. &Muench, source of resistance to Phytophthora infestans,
2n=24. C. Mexico, southwards to Guatemala. virus X, virus Y, and several insects. The diploid
forms a link between the Demissa and the Tube-
SOLANUM OXYCARPUM Schiede. 2n=48. EC. rosa series of S. America. It is similar to the
Mexico, Honduras, Costa Rica and adjacent Pana- Colombian S. andreanum*.
ma.
SOLANUM WOODSONn Corr. 2n= . Dry s p e -
SOLANUM PAPITA Rydb. 2n= Similar to cimens came from Costa Rica, Panama and Vene-
S. fendleri*. zuela. No living material has been collected yet.
SOLANUM PINNATISECTUM Dun. 2n=24. NC. Sterculiaceae
Mexico. A maize field weed. A source of r e s i s -
tance to Phytophthora, Y virus and Colorado THEOBROMA BICOLOR H. &B. Nicaraguan
beetle. A parent of S. sambucinum*. cacao. 2n=20. From Mexico to Brazil. Cultivated
outside its natural range for the edible pulp round
SOLANUM POLYADENrUM Greenm. 2n=24. C. the seeds. The seeds themselves a r e used like
Mexico. A source of Phytophthora resistance. those of Th. cacao* (Purseglove, 1968).
SOLANUM POLYTRICHON Rydb. 2n=48, genome THEOBROMA PANTAGONA Bern. Cacao lagarto.
formula A4A4BB. NW. to NC. Mexico. In waste 2n= . Costa Rica to Panama. Cultivated t h e r e .
places, shrubland and cultivated fields. Its g e -
nomes a r e related and also to the Ajgenome of
S. phureja*. S. bulbocastanum* has the genome
formula BB. S. polytrichon is used as a source
of resistance to Phytophthora and potato aphid.
SOLANUM x SAMBUCINUM Rydb. 2n=24. In maize

fields in NC. Mexico. A natural hybrid of S. c a r -
diophyllum ssp. ehrenbergii* (2n=24) and S. pinna-
tisectum* (2n=24) (Hawkes, 1966).
SOLANUM x SEMIDEMISSUM Juz. 2n=60. Clones

of this 5x plant in roadside thickets in Mexico. A
source of Phytophthora and frost resistance. It is
thought that it is a hybrid of S. demissum x S.
tuberosum, the reciprocal of the cross yielding
S. x edinense. It is sterile (Ugent, 1967). This
12 North American
Centre
Darlington & Janaki Animal (1945) established the USA Centre of Origin. Zhukovsky (1968) enlarged
this a r e a to the southern half of N. America. Agriculture must have been introduced from Centre 11 in
the third millenium BC., although there a r e indications that chenopods and amaranths were deliberately
cultivated (Haury, 1962). However, they may have belonged to the ruderal flora the seeds being collected
as food. To the earliest introduced crops belong Zea mays.
A few but important crops have been domesticated in this centre: Fragaria virginiana, Helianthus sp.
Prunus s p . , Rubus s p . . Vaccinium s p . , Vitis sp. etc.
Aceraceae cause it has been discovered to be a source of the

ACER SACCHARUM Marsh. Sugar maple, Rock lobeline used in anti-smoking preparations.
maple. 2n=26. E. of N. America. The sap is a
source of maple sugar and maple syrup. Also cul- Chenopodiaceae
tivated. Mansfeld (1959) suggested that A. saecha- ATRIPLEX CANESCENS (Pursh. ) Nutt. American
rinum L. is a synonym, but this species has shad scale, Hoary saltbush. Wing scale, Fourwing
2n=52. saltbush. 2n= . W. of N. America up to Mexi-
co. Cultivated as a fodder plant on saline soils and
Amaranthaceae as a hedge plant (Mansfeld, 1959). A. gardnesi
(Moq.) Standi. (2n^ ) is from the same area and
AMARANTHUS HYBRIDUS*
A. truncata (Torr.) A. Gray. (2n= ) is from
Utah, USA.
AMARANTHUS POWELLII see A. leucarpus*
AMARANTHUS SPINOSUS* Compositae

HELIANTHUS ANNUUS L. Sunflower. 2n=14, g e -
Araceae nome formula Ba^Baj. N. America. It is difficult
ACORUS CALAMUS* to be more specific because of its spread as a
food plant and weed. The wild type may have r e -
Araliaceae sembled ssp. jaegeri, a small-headed type found
in SW. Utah and NE. Arizona to S. California in
PANAX QUINQUEFOLIA L. American ginseng. USA. In new areas the sunflower has introgressed
2n= . E. part of N. America. Cultivated for with related species so that the morphological
its roots, which are used as a stimulant. variability has increased. An important species is
H. bolanderiA. Gray (2n=34) of C. and N. Califor-
Asclepiadaceae nia. Other species H. agrophyllus Torr. &A. Gray
ASCLEPIAS SYRIACA L. (syn. A. cornuti Decne). (2n=34) of Texas, H. debilis Nutt. ssp. cucumeri-
2n=(20), 22, (24). E. of N. America. Run wild in folius (2n=34) of Texas and H. petiolaris Nutt.
cultivated ground and dry grassland (Tuti n e t a l . , (2n=34) of W. of N. America have also hybridized
1972). Cultivated formerly in Europe for fibre and with the sunflower.
as a food-plant for bees. Various subspecies and varieties a r e recognized:
ssp. lenticularis (Dougl.) Ckll, which is near to
the original wild type and is found from W. Canada
Campanulaceae
to N. Mexico. Ssp. texanus grows mainly in Texas.
LOBELIA INFLATA L. Indian tobacco. 2n=14, It may have arisen by hybridization of ssp. lenti-
(16). N. America. Cultivated there as a medicinal cularis and H. debilis.
plant. It has attracted attention again recently b e - Ssp. annuus L. is a ruderal weed, a weed sun-
NORTH AMERICAN CENTRE 174
Km
\)\i —~s~?y
0
17 <ö
1 3 ^*~
yl / x_ftX—-——
^ 2
s
^"sVe
\ T CT
V
Distribution of Helianthus spp. in pre-human (above), pre-columbian (middle) and modern (below) times. (H. annuus ( ), H. pe-
tiolaris (speckled), H. exilis (A), H. argophyllus (B), H. debilis var. cucumerifolius (...), H. debilis var. debilis (C), H. bolan-
deri ( ), cultivated sunflower (1), campflower (2), Great plains annuus (3), weed petiolaris (4) and weed cucumerifolius (5) (An-
derson, 1956; based on Heiser's research)
ACERACEAE - GRAMINEAE 175
flower, found in the settlements in the Midwest. those of other wild species like V. myrtillus L . .
It possibly derives from ssp. lenticularis. Var. (European) blueberry, Whortleberry, Billberry
macrocarpus (DC.) Ckll is probably the p a r e n - (2n-24).
tal from of the cultivated types. It grows in NE.
of USA and Canada. It probably originated from VACCINIUM MACROCARPON Ait. Cranberry.
ssp. annuus or this subspecies and var. m a c r o - 2n=24, genome formula MaMa. NE. N. America.
carpus developed together from ssp. lenticularis. Cultivars selected and cultivated in N. America
A weedy sunflower may have reached the and on Terschelling, the Netherlands. Related to
Middle West where no other annual Helianthus the N. American wild V. oxycoccos L. (2n=48)
species are found. Here the giant, large-headed genome formula Mi 0 Mi°Ma 0 Ma°. The Mi 0 genome
sunflower may have developed (Heiser, 1955, is related to Mi of V. mlcrocarpum (Turcz. ) Hook.
1965). (2n=24) (Ahokas, 1971).
From N. America the sunflower was intro-
duced into Europe where in USSR a secondary Euphorbiaceae
centre of diversity arose (p. 130).
H. x laetiflorus P e r s . (2n=102), a wild p e r e n - ALEURITES FORDII Hemsl. Tung oil tree, 2n=22.
nial species of USA is probably a hybrid of H. SC. China (p. 31). Secondary gene centre proba-
subrhomboideus Rydb. (2n=102) and H. tuberosus. bly in the large planting.
The first parent is also native to USA (Clevenger
and Heiser, 1963). Fagaceae
CASTANEA DENTATA (Marsh. ) Borkh. American
HELIANTHUS TUBEROSUS L. Jerusalem a r t i - chestnut. 2n=24. E. of N. America. A t r e e culti-
choke. 2n=102, genome formula At.At At„At Bt 1 vated for its edible sweet nuts.
Bt,. N. America. Run wild in S. UTcrame ana N.
Caucasus. A perennial species introduced to CASTANEA PUMILA (L. ) Mill. Common chinqua-
Mexico and Eurasia. pin. 2n= . N. America (from Pennsylvania to
The Bt.genome is related to the Ba1 genome Florida and Texas). A t r e e cultivated for its nuts.
of H. annuus*. H. tuberosus is probably an amphi-
ploid of a species with genome formula At.At..At Gramineae
At„ and a B genome donor. This might be H.
annuus or else a closely related species. AGROPYRON PAUCIFLORUM (Schweinitz) Hitchc.
(syn. A. trachycaulum (Link. ) Malte). Slender
Cupressaceae wheatgrass, Bald wheatgrass. Western r y e g r a s s .
2n=28. N. America. A forage crop.
JUNIPERUS VIRGINIANA L. Eastern red cedar.
2n= . E . N . America. Much variation is due AGROPYRON SMITHII Rydb. Western wheatgrass.
to introgressive hybridization with other Juniperus 2n^28. N. America. Used to control erosion and
species (Hemmerly, 1970). as a forage crop.
Ebenaceae AGROPYRON SPICATUM (Pursh) Scribn. &Smith.
DIOSPYROS VIRGINIANA L. Common persimmon. Bluebunch wheatgrass. 2n=14, genome formula
2n= . E. of N. America. Cultivated for its SS, 28. N. America. Cultivated as a forage crop.
fruits. Also used as a rootstock of D. kaki*. Var. inerme is Beardless bluebunch wheatgrass.
A. latiglume (Scribn. &Smith) Rydb., (2n=28),
Elaeocarpaceae A. scribneri Vasey (2n=28), A. trachycaulum (Link.
Malte ex H. F. Lewis (2n=28) and Sitanion hystrlx
MUNTINGIA CALABURA L. Panama berry, Capu- (Nutt. )J. C. Smith (2n=^28) also possess a pair of
lin, 2n= . Widely cultivated for its sweet, S genomes.
edible fruits.
AGROSTIS GIGANTEA*
Ericaceae
VACCINRJM ASHEI Reade. Rabbiteye. 2n=72. AGROSTIS TENUIS*
N. America. Cultivated there. Wild plants are
also harvested. According to Camp (1945) the wild BOUTELOUA CURTIPENDULA (Michx. ) T o r r .
types derive from hybridization of the tetraploid Side-oats grama. 2n=20-103. N. America. An i m -
species V. arkansanum, V. australe Small, South- portant pasture g r a s s .
eastern highbush blueberry, V. darrowi-4x and
V. myrsinites Lam. , Ground blueberry. BOUTELOUA ERIOPODA T o r r . Black grama,
Grama g r a s s . 2n=20, 21, 28. SW. of USA and N.
VACCINIUM CORYMBOSUM L. Highbush berry. Mexico. Used as a pasture g r a s s .
2n=72. N. America. Cultivars have developed
from the wild type which arose from hybridization BOUTELOUA FILIFORMIS (Fourn. ) Griff. Grama
of the tetraploid species V. lamarckii Camp. (syn. g r a s s . 2n=14, 20, 21, 22, 46. W. of USA and N.
V. angustifolium Ait. ), V. alto-montanum Ashe, Mexico. A pasture grass.
V. simulatum and V. australe Small, Southeastern
highbush blueberry. BOUTELOUA GRACILIS (H.B.K. ) Lag ex Steud.
Fruits of wild plants are still picked, as a r e
Blue grama. 2n=20-84. N. America. ranging from adaptation to dispersal by floating (Galinat, 1969).
Canada to Mexico.
BROMUS MARGINATUS Nees. Mountain brome.

2n=28, 42, 56. 70. N. America. Cultivated in
Australia, E. Africa and N. America.
DESCHAMPSIA INSIGNIS P i e r r e (syn. D. caespi-

tosa (L. ) Beauv. ). Tufted hairgrass, Tussock
g r a s s . 2n=(24, 25), 26. (27), 28, (49). W. of N.
America. Cultivated for hay and pasture.
Putman &Klein (1971) suggested that from this
species D. elongata (Hook. ) Munro (2n=26) and
D. holciformis P r e s l (2n=26) have been derived
From D. elongata comes the annual D. danthoni-
oides (Trin. ) Munro. These four species a r e found
in N. America and Europe.
DIGITARIA ADSCENDENS (H.B.K. ) Henrard.

T r i p s a c u m dactyloides (— I and T. floridanum {• • )
2n=54. N. America. A weedy grass sometimes (Randolph, 1970)
included in D. sanguinalis*, but Gould (1963)
classified 2n=6x=54 plants as D. adscendens and
2n=4x=36 plants as D. sanguinalis. These species TRIPSACUM FLORIDANUM P o r t e r ex Vasey.
are closely related. 2n=36. S. Florida and Texas, USA. Cross compa-
tible with Zea mays*.
ELYMUS CANADENSIS L. Canada wild-rye.
2n=28, (42). N. America. A grass used as pasture
UNIOLA PANICULATA L. Sea cats. 2n=40. S. of
and hay crops.
N. America. A dune stabilizer.
PANICUM VIRGATUM* ZEA MAYS L. Maize, Corn. 2n=20. Secondary
centres in N. America. Domesticated in C. Ame-
PASPALUM DISTICHUM L. 2n=40, (48), 60. N. rica (p. 165). There local races and those introdu-
and S. America. Used in the USA as a soil stabili- ced from the Mexican lowlands and the West Indies,
zer. hybridized (Brandolini, 1970). Dent maize, dent
corn - indentata Sturt. - is common here.
POA COMPRESSA L. Canada blue g r a s s . Wire
g r a s s . 2n=(14), 42, 56 and aneuploids. Europe to
ZIZANIA AQUATICA L. Indian rice. Wild rice,
the Urals and Caucasia, and in N. America.
Tuscarora r i c e . 2n=30. S. Canada and USA in the
Selection work was done particularly in N. A m e r i -
a r e a of the Great Lakes. Seeds are collected. Not
ca.
cultivated.
POA NEMORALIS L. Wood meadows g r a s s . 2n=28,
Grossulariaceae
42, 56, c. 70 and aneuploids. Europe, N. Africa,
temperate Asia, N. America. Cultivated in N. RIBES CYNOSBATI L. American wild gooseberry,
America. Frickle gooseberry. 2n=16.E. N. America. Cul-
tivated there. Its characteristic mildew resistance
POA PRATENSIS L. Blue grass, Kentucky blue is useful in breeding European gooseberry culti-
g r a s s , Bird g r a s s . 2n=38-147. For origin see v a r s (p. 135).
p. 134. Europe, Asia, N. Africa and northern N.
America. Various varieties have been developed RIBES ODORATUM Wendland. Buffalo currant.
in Europe, Canada and elsewhere. Natural hybrids 2n=16. N. America. Types with large fruits are
with P . ampla Merr. have been found in W. N. cultivated.
America.
Hydrastidaceae
PUCCINELLIA NUTTALLIANA (Schult. ) Hitchc.
HYDRASTIS CANADENSIS L. Golden seal, Orange
Nuttal alkali grass, Meadowgrass. 2n=42, 56. N.
root. 2n=26. E. of N. America. Cultivated there
America. Cultivated as a forage grass especially
as a medicinal crop and as an ornamental.
on alkaline soils.
Hydrophyllaceae
STIPA VIRIDULA Trin. Green needle g r a s s .
2n=82. N. America. This perennial bunch grass PHACELIA TANACETIFOLIA Benth. Valley v e r -
has been adapted to the N. Great Plains for r a n g e - venia. 2n=(18), 22. California, USA. Cultivated
land. as a crop for honey bees.
TRIPSACUM DACTYLOIDES (L.) L. 2n=36, 72.

See p. 165. Tetraploid types of Florida, USA have
a thick rachis which apparently developed as an
GRAMINEAE - ROSACEAE 177
Juglandaceae SESBANIA MACROCARPA Muhl. 2n=12. USA.

Cultivated there. Closely related to S. exaltata*
CARYA ILLINOINENSIS (Wangenh. ) K. Koch,
or is a synonym of this species.
(syn. Carya pecan Engler &Graebn. ). Pecan.
2n=32. N. America. A t r e e cultivated for its nuts.
One of the parents of C. x lecontei Little, bitter VICIA LEAVENWORTHII Torr. &Gray. Leaven-
pecan (2n= ). The other parent is probably C. worth vetch. 2n=14. Missouri and Arkansas to
Texas in USA. Occasionally cultivated.
aquatica (Michx.f. ) Nutt., water hickory (2n=32).
CARYA OVATA (Mill.) K. Koch (syn. C. alba Liliaceae

Nutt,). Shagbark hickory, Shellbark hickory. CAMASSIA LEICHTLEMII (Baker) S. Wats. Camas.
2n= . N. America. Cultivated also as an orna- 2n^30. E. part of N. America. Uncultivated bulb
mental. beds a r e divided into family plots, which have
been passed down from generation to generation.
JUGLANS HINDSIIJepson. C. Californian black These plots a r e not farmed, but stones, weeds and
walnut. 2n= . C. California, USA. Cultivated shrubs a r e removed every year. In most cases the
as rootstock of J. regia*. plants a r e marked in bloom so that the bulb can be
harvested when it is fully grown (Chapman Turner
JUGLANS NIGRA L. Black walnut. 2n=32. Most & Bell, 1971).
of the eastern half of the USA and southern Ontario The camas is semi-domesticated i . e . the
between Lake Huron and Lake Ontario. Pure, n a - wild plant is protected and the growing c i r c u m -
tural stands are r a r e and usually small, it gene- stances a r e improved. The latter may result in
rally occurs as individual t r e e s scattered through more sites for the plant to grow.
the forest. This walnut is mainly selected for its
fruits, but also some selections have been made CAMASSIA QUAMASH (Pursh) Greene. Blue c a -
for distinctive foliage and for wood (Funk. 1969). mas. 2n= . E. part of N. America. See fur-
ther C. leichtlinii*.
Labiatae
Malvaceae
MENTHA CANADENSIS L. (syn. M. arvensis L. ).
American wild mint. 2n=54, 96. N. America. GOSSYPIUM BARBADENSE L. Sea Islands cotton.
Occasionally cultivated. 2n=52, genome formula (AADD)„. Peru (p. 153).
Sea Islands cotton developed on the Sea Islands of
Leguminosae S. Carolina, USA after introduction from Bahamas
or Jamaica. Cultivated in E. USA and some Ca-
AMPHICARPAEA MONOICA (L. ) Ell. (syn. F a l -
ribbean islands. Maybe similar types in Ecuador
cata comosa (L. ) Kuntze). Hog pea, Ground p e a -
are its parental material (Harlan, 1970).
nut. 2n=20. E. and S. of USA. Formerly cultiva-
ted for its seeds.
Martynaceae
APIOS TUBEROSA Moench. (syn. A. americana PROBOSCIDEA LOUSIANICA (Mill. ) Thell. (syn.
Medik., Glycine apios L. ). Potato bean, Ground Martynia proboscidea Glox. ). Unicorn plant,
nut. 2n= . E . N . America, Florida and Texas. Ram's horn, Double claw, Proboscis flower.
Cultivated for its edible tubers and as an ornamen- 2n= . N. America. A herb cultivated for its
tal or as a curiosity. fruits and as an ornamental.
GLEDITSIA TRIACANTHOS L. Honey locust, Moraceae

Sweet bean. 2n=28. C. a n d E . N. America. This
MACLURA POMIFERA (Rafin.)C.K. Schneider.
t r e e is cultivated in hedges and as an ornamental.
2n= . N. America. Used for hedges and as an
ornamental.
LUPINUS PERENNIS L. 2n=48, 96. E. N. A m e r i -
ca. Cultivated as an ornamental, fodder crop and
green manure. Passifloraceae
PASSIFLORA INCARNATA L. May-Pop, Apricot-
LUPINUS POLYPHYLLUS Lindl. 2n=48. Pacific Vine. 2n=18, 36. E. N. America, Florida and
N. America. Cultivated as an ornamental, fodder Texas. It was cultivated by Indians in Virginia.
crop and green manure.
Phytolaccaceae
ROBINIA HISPIDA L. 2n=30. SE. of N. America.
PHYTOLACCA AMERICANA L. 2n=36. USA. It has
Cultivated as an ornamental and in hedges.
run wild in S. Europe. Cultivated as a dye plant
(berries) and ornamental.
ROBINIA PSEUDACACIA L. Locust, Black locust.
2n=20, c. 20, 22. C. a n d E . N. America. This
t r e e is planted as an ornamental and as a soil s t a - Rosaceae
bilizer. AMYGDALUS PERSICA L. Peach. 2n=16. P r i m a -
ry centre: China (p. 36). Secondary centre: Cali-
SESBANIA EXALTATA (Raf. )Rydb. 2n=12. USA. fornia, USA.
Cultivated there as a green manure.
FRAGARIA CHILOENSIS* PRUNUS PENSYLVANICA L. (syn. P. p e r s i c i -

folia Desf. ). Bird cherry, Pigeon cherry, Pin
FRAGARIA VIRGINIANA Duch. Virginian s t r a w - cherry, Wild red cherry. 2n=16. N. America,
berry. 2n=56. N. America, especially in the from New Foundland to British Columbia and
eastern part. Cultivated. One of the parents of Colorado. A source of late flowering and cold
F. ananassa* Duch. (syn. F. grandiflora Ehrh. ), resistance.
the pineapple strawberry (2n=56).
PRUNUS SEROTINA Ehrh. Black cherry, Rum
PRUNUS. The American Prunus species are cherry. 2n=32. N. America from Ontario and N.
valuable because of their longevity and winter- Dakota to Texas and Florida. The fruits are un-
hardiness. palatable. It could be a source of late flowering
and frost resistance.
PRUNUS AMERICANA Marsh. American Plum.
2n=16. A large t e r r i t o r y of N. America between PRUNUS VIRGINIANA L. Common choke cherry,
Manitoba and Texas. This small t r e e usually Eastern choke cherry, Choke cherry. 2n=30, 32.
grows slowly. Cultivated. Valuable because of its W. of N. America. Micurin selected Vinogradnaja
longevity and winterhardiness. Cultivars have from this species (Zylka, 1971b).
been developed from interspecific c r o s s e s .
RUBUS ACAULIS Michx. 2n=14. The Canadian
PRUNUS ANGUSTIFOLIA Marsh, (syn. P. chica- counter part of R. arcticus (Larsson, 1969). It
sa Mich. ). Chickasaw plum, Florida sand plum, could be one parent of R. stellatus*. Also named
Mountain cherry. 2n=16. S. Delaware to Florida ssp. acaulis of R. arcticus.
and westward to the Texas and S. Oklahoma. A
small t r e e with a dense crown. The cultivated RUBUS FLAGELLARIS Willd. (syn. R. villosus
species lack hardiness. It hybridizes easily with Ait.). Northern dewberry. 2n=63. E. N. America.
P . munsoniana*. Cultivated for its fruits. Var. roribaccus Bailey
is the Lucretia dewberry (Uphof, 1968).
PRUNUS BESSYI Bailey. Western sand cherry.
2n=16. N. America on sandy and saline soils. RUBUS IDAEUS L. (syn. R. strigosus Michx. ).
This perennial shrub is characterized by a good American red raspberry. 2n=14. The NE. A m e r i -
longevity, frost-resistance and sweet, edible can counterpart of this species. Present cultivars
fruits. A source of a bush-type habit for ease of are often hybrids between this subspecies and ssp.
mechanical harvesting. It hybridizes with A r m e - vulgatus*, the European red raspberry. Natural
niaca vulgaris* and Prunus ssp. hybrids between ssp. strigosus and R. occidenta-
lis* have been described as R. neglectus Peck.,
PRUNUS HORTULANA Bailey. Hortulan plum. Purple cane raspberry (2n=14). They have been
2n=16. C. USA. This t r e e is very cold resistant occasionally cultivated. In N. America such hybri-
and has good fruits. It flowers late (Zylka, 1970). dization has lead to introgression between the two
It hybridizes easily with other American Prunus parental species. See for other hybrids p. 141.
species. According to Bailey (1898) it is a hybrid Cultivars are often unarmed or have simple leaves.
of P. angustifolia* and P. americana*. The loganberry (2n=42), genome formula V, V.V„
V„II, is derived from the cross of a tetraploid
PRUNUS MARITIMA Marsh, (syn. P . maritima form R. ursinus Cham. &Echt. (2n=28), genome
Wangh., P . acuminata Michx. ). Beach plum, formula ViV-i V 2 V 9 a n d R - i d a e u s (2n=14). May-
Sand plum. 2n=16. E. of USA and adjacent Canada. berry is aliybrid product of R. palmatus Thunb. x
Some cultivars have been selected (Zylka, 1970) ssp. strigosus and Youngberry (2n=42, 49) of l o -
in the USA. Also used as a source of late flowe- ganberry x Mayes dewberry (R. baileyanus x R.
ring, cold resistance and very high fertility in the argutus). Other Rubus species like R.arcticus* '
breeding of other cultivars. It is an ornamental. have also been used in breeding work.
PRUNUS MUNSONIANA Wight &Hedrick. Wild RUBUS OCCIDENTALES L. Black raspberry.

Goose Plum. 2n=16. N. Texas, E. Oklahoma and 2n=14. NE. America, Colorado and British Colum-
Missouri. It flowers late and the fruits a r e of bia. It is cultivated. Present cultivars often derive
good quality. It has a good longevity and winter- from hybrids with R. idaeus*. Natural hybrids
hardiness. It hybridizes easily with P. angustifo- with the N. American ssp. strigosus of R. idaeus
lia* and other species. Cultivars were bred from have been named R. neglectus Peck., purple cane
such interspecific c r o s s e s . raspberry (2n=14). This hybridization has lead to
introgression between these two parental species.
PRUNUS NIGRA Ait. Canada plum. 2n=16. The It is for R. idaeus* a source of resistance to the
t e r r i t o r y between S. New Foundland to the Strait rubus aphid, Amphorophora rubi Kalt.
of Mackinac and southward to Lansing, Michigan.
Cultivated there to some extent. RUBUS STELLATUS Sm. 2n=14. Wild from Alaska,
This species is valuable because of its longe- Aleutian Islands and.Kamtchatka. Closely related
vity and winterhardiness. Cultivars have been to R. arcticus* and has been described as ssp.
developed from interspecific c r o s s e s . stellatus of this latter species. It could be a hybrid
of R. arcticus and R. acaulis* (Larson, 1969).
ROSACEAE - VITADACEAE 179
Salicaceae VITIS ROTUNDIFOLIA Michx. Muscadine grape,

Southern fox grape. 2n=40. Florida, the southern
SALIX RIGIDA Mühlenberg (syn. S. cordata Mühl. ).
coast of USA and the east coast of Mexico. Var.
American willow. 2n^44. N. America. Introduced
scuppernong is cultivated in S. part of USA. Hy-
in Europe for twig production. brids of this species and V. vinifera* a r e comple-
tely resistant to Phylloxera, Erysiphe and Oidium
Solanaceae
DATURA STRAMONIUM L. Thorn-apple, Jimson VITIS RUPESTRIS Scheele. 2n-- . SW. of USA.
weed. 2n=24. N. America. Pantropical now. A Used as a rootstock. It can be crossed with V.
poisonous plant cultivated as a medicinal plant vinifera* to introduce resistance to phylloxera,
yielding stramonium. Viteus vitifolii Shimer.
NICOTIANA QUADRWALVIS Pursh. 2n=48. SW.

of the USA along r i v e r s and in rocky soils. The
Indians cultivated it for smoking leaves and flo-
w e r s . An annual, frost-resistant, early maturing
species. It hybridizes easily with N. tabacum*.
PHYSALIS PUBESCENS L. Strawberry tomato.

Dwarf Cape gooseberry. 2n=24. N. America.
Cultivated in Ukraine and elsewhere.
SOLANUM FENDLERI A. Gray. Navajo potato.

2n=48, genome formula A.A ,BB. Arizona. New
Mexico and W. Texas, USA and NW. Mexico.
Very similar to S. hjertingii* and similar to S.
papita*. It is resistant to Y virus.
SOLANUM JAMESII Torr. 2n=24, 36. US states

Arizona, New Mexico and Colorado, and in Mexi-
co near the Arizona boundary.
Valerianaceae
VALERIANA EDULIS Nutt. 2n= . N. America.
This perennial herb is cultivated for its roots.
Vitadaceae
VITIS BERLANDIERI Planch. 2n-38. SE. of USA
to Texas. Used as a rootstock. Rootstocks of V.
riparia* x V. berlandierl are also used. It can be
crossed with V. vinifera*.
VITIS CINEREA Engelm. Downy grape, Sweet

winter grape. 2n=38. SE. of USA. Resistant to
fungal diseases and to phylloxera, Viteus vitifolii
Shimer, but it cannot be crossed with V. vinifera*.
VITIS CORDIFOLIA Michx. 2n=30, 38. SE. of

USA. It can be crossed with V. vinifera* to intro-
duce resistance to fungal diseases.
VITIS LABRUSCA L. Fox grape. 2n=38. E. N.

America. It appears to have run wild in Georgia,
USSR (p. 90). Introduced into the Old World. It is
cultivated. It has been crossed with V. vinifera*
to improve it and for this species it is used as a
rootstock.
VITIS RIPARIA Michx. (syn. V. vulpina L. ).

2n=38. E. and C. USA and in Ontario, Canada.
Used as a rootstock. It can be crossed with V.
vinifera* to introduce resistance to phylloxera,
Viteus vitifolii Shimer. Rootstocks of V. riparia x
V. berlandieri* a r e also used.
Species with a not identified centre
Some species could not be listed in one of the gene centres. They have a very wide geographical distribu-
tion. Either their locality of cultivation has not been reported or they a r e cultivated at several places.
Thus it is not known whether their wide distribution occurred before their domestication or not, i. e. it is
not mentioned whether the wild species grew in a large area where it has been domesticated at several
places, or whether wild plants were domesticated on one site after they had been spread by man.
Amaranthaceae MOMORDICA CHARANTIA L. Bitter gourd, Bitter

GOMPHRENA GLOBOSA L. Batchelor's button. cucumber, Balsam pear. 2n=22. Tropics of the
2n=32, 40-44, 44-48. The tropics. Cultivated in Old World. Naturalized in nearly all tropics and
some villages of mainly coastal districts in W. subtropics. An edible, medicinal and toxic plant
Africa as a curiosity and as a fetish plant. In (Morton, 1967).
Europe and elsewhere it is an ornamental.
Cyperaceae
Anacardiaceae CYPERUS ARTICULATUS L. 2n= Cultivated for
its sweet scented roots.
SPONDIAS CYTHEREA Sonner (syn. S. dulcis
Forst, f.). Otaheita apple, Ambarelia, Hog plum.
2n= . Cultivated for its fruits. Mansfeld (1959) MARISCUS UMBELLATUS Vahl. 2n= . The.tro-
reported the presence of var. cythera on the So- pics of the Old World. Cultivated for its rhizomes.
ciety Islands, Tahiti, Fidji, Samoa and Madagas-
car, var. mucroniserrata (Engl. ) Mansf. in Mexi- Gramineae
co, var. macrocarpa (Engl. ) Mansf. in Brazil, CENCHRUS CILIARES L. (syn. Pennisetum cen-
var. acida (BL. ) Mansf. in Malaysia and var. chroides Rich. ). Buffel g r a s s . 2n= mainly 36.
intégra (Engl. ) Mansf. in Amboina. The tropics and subtropics. Cultivated in Australia.
Apocynaceae HETEROPOGON HIRTUS P e r s . (syn. H. contortus

Beauv. ex Roem. &Schult., Andropogon contortus
APOCYNUM VENETUM L. (syn. A. sibiricum Pall,
L. ). Spearhead, Tangle g r a s s . 2n=20, 40, (42-44,
ex Roem. &Schult. ). Kendyr. 2n=16, 22. Italy to
50), 60, (80, c. 70-80). The Old and New World,
E. Asia. A perennial fibre crop brought into culti-
subtropics and tropics. Cultivated as a fodder for
vation in USSR.
live stock. Plants with 2n=20 come from India, with
2n=40 from Java, India, Madagascar, Tanzania,
Chenopodiaceae Kenya, Uganda, Rhodesia and Zaire, and with 2n=60
CHENOPODRJM BOTRYS L. Jerusalem oak. 2n=18. from S. Africa, Mexico and N. America.
S. Europe, Orient and C. Asia. Occasionally cul-
tivated (Mansfeld, 1959). Leguminoseae
CROTALARIA MUCRONATA Desv. (syn. C. s t r i a -
Cucurbitaeeae
ta DC. ). 2n=16. Distributed in tropics and s u b -
MOMORDICA BALSAMINA L. Balsam apple. 2n=22. tropics as a cover crop and green manure.
Tropics of the Old World. Leaves and stem a r e used
for fodder.
AMARANTHACEAE-URTICACEAE 181
CROTALARIA RETUSA L. 2n=16. This fibre crop

is widespread throughout the tropics. In E. Africa
it is used for its pigments and in Florida as an
ornamental.
DESMODIUM ADSCENDENS DC. 2n=22. Tropics.

Cultivated in many regions as a cover crop and
green manure.
DESMODIUM GANGETICUM DC. 2n= . Trop.

Asia and Australia. Cultivated as a fodder plant
and green manure.
INDIGOFERA HIRSUTA L. Hairy indigo. 2n=16.

Many tropical countries. It is cultivated.
TEPHROSIA PURPUREA P e r s . Purple tephrosia.

2n=22, (24, 44). The tropics. Cultivated as a
green manure and cover crop. Grows wild in N.
India on waste places and road sites.
Malvaceae
ABUTILON GRAVEOLENS Sweet. 2n=14, 36. The
tropics of the Old World. Cultivated especially in
USSR for its oily seeds.
MALACHRA CAPITATA L. 2n=56. The tropics.

A herb cultivated for its fibre.
URENA LOBATA L. Aramina fibre, Congo jute.

2n=28, 56. Wild or naturalized in the tropics and
subtropics. Centre of origin very likely in the Old
World (Purseglove, 1968).
Rutaceae
EVODIA HORTENSIS J . R . &G. Forst, (syn. F a -
gara euoda L.f., F . e v o d a L . f . , Zanthoxylum
varians Benth. ). 2n= . A widespread shrub.
Cultivated in many parts of the Pacific. Leaves
a r e used for medicinal purposes.
Umbelliferae
CONIUM MACULATUM L. Hemlock, Poison h e m -
lock. 2n=16, 22. Europe, Asia, N. Africa and
Ethiopia. Occasionally cultivated. Often occurring
as a ruderal. Used as a poison and medicinal plant.
Urticaceae
BOEHMERIA STIPULARIS Wedd. Hawaian false
nettle, Akola. 2n= . Probably from the Mascarene
islands. According to Uphof (1968) this fibre crop
grows wild on Hawaii, where it was formerly cul-
tivated. This has resulted in many varieties.
TOUCHARDIA LATIFOLIA Gaudich. Olona. 2n=

The Hawaian Islands. At one time cultivated for
its fibre (Hutchinson, 1969).
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Index of botanical names
After the page number the centre of diversity is given between brackets.
ABBEVILLEA fenzliana 155 (10) ADENANDRA fragrans 128 (8)

ABELMOSCHUS esculentus 68 (4) ADENOPSUS abyssinicus, see Lagenaria s i c e -
- manihot 35, 68 (1, 4) r a r i a 110 (8)
- moschatus 68 (4) - guineensis, see Lagenaria siceraria 110 (8)
- tuberculatus, see A. esculentus 68 (4) - longiflorus, see Lagenaria siceraria 110 (8)
ABERIA gardneri, see Doryalis hebecarpa 68 (4) - pynaerti, see Lagenaria siceraria 110 (8)
ABUTILON avicennae 35 (1) - reticulatus, see Lagenaria siceraria 110 (8)
- graveolens 181 (?) - rufus, see Lagenaria siceraria 110 (8)
- indicum 51, 68 (2, 4) AEGILOPS aucheri, see A. speltoides 79 (6)
- oxycarpum 153 (10) - bicornis, see A. speltoides, Triticum t u r g i -
- theophrasti, see A. avicennae 35 (1) dum 96 (7)
ACACIA arabica, see A. nilotica 118 (8) - biuncialis, see A. lorentii 79 (6)
- cavenia 150 (10) - bushirica, see A. t r i a r i s t a t a 79 (6)
- cibaria, see A. longifolia 57 (3) - caudata, see also A. c r a s s a , A. triuncialis
- cyanophylla 57 (3) 72, 79 (5, 6)
- dealbata 57 (3) - columnaris 79 (6)
- decurrens, see A. mearnsii 57 (3) - comosa 79, 96 (6, 7)
- farnesiana, see also A. cavenia 150, 151 (10) - c r a s s a 79 (6)
- horrida, see A. karroo 118 (8) - cylindrica, see also A. triuncialis 72, 79, 96
- karroo 118 (8) (5, 6, 7)
- longifolia 57 (3) - juvenalis 72, 79 (5, 6)
- mearnsii 57 (3) - kotschyi 72, 79, 96 (5, 6, 7)
- mollissima, see A. mearnsii 57 (3) - ligustica, see A. speltoides 80 (6)
- nilotica 118 (8) - longissima, see A. speltoides 80 (6)
- pycnantha 57 (3) - lorentii 72, 79, 96 (5, 6, 7)
- Senegal 118 (8) - macrochaeta, see A. lorentii 79 (6)
ACER saccharinum, see A. saccharum 173 (12) - mutica 80 (6)
- saccharum 173 (12) - ovata 72, 80, 96 (5, 6, 7)
ACHRAS mammosa, see Calocarpum sapota 170 - peregrina, see A. variabilis 96 (7)
(11) - recta, see A. t r i a r i s t a t a 79 (6)
- sapota, see Manilkara achras 170 (11) - sharonensis, see A. speltoides 80 (6)
ACNIDA, see Amaranthus hybridus 170 (11) - speltoides, see also Triticum timopheevi,
ACOLANTHUS pubescens 118 (8) T. turgidum 80 (6)
ACONITUM carmichaeli 36 (1) - squarrosa, see also A. crassa, Triticum
- napellus 138 (9) aestivum 72, 80 (5, 6)
- wilsonii, see A. carmichaeli 36 (1) triaristata, see also A. columnaris 72, 80, 96
ACORUS calamus 129, 173 (9, 12) (5, 6, 7)
ACROCERAS amplectans 112 (8) triuncialis, see also A. kotschyi 72, 80, 96
- macrum 112 (8) (5, 6, 7)
ACTINIDIA ARGUTA 27 (1) turcomanica. see also A. juvenalis 79 (6)
- chinensis 27 (1) umbellulata, see also A. ovata, A. triuncialis
- kolomicta 27 (1) 80 (6)
- polygama 27 (1) uniaristata 96 (7)
INDEX OF BOTANICAL NAMES 194
AEGILÜPS variabilis 80, 96 (6, 7) ALLIUM ascolinicum 77 (6)

- ventrieosa 96 (7) - bakeri, see A. chinense 27 (1)
AEGIl.OTRICUM, sec Acgilops ventrieosa 96 (7) - cepa, see also A. ascolinicum 71, 91 (5, 7)
AEGLE marmelos 54 (2) - chinense 27 (1)
AESCHYNOMENE americana 151 (10) - fistulosum, see also A. cepa, A. chinense
- glandulosa, see A. americana 151 (10) 27 (5)
AESCULUS earnea, see A. hippocastanum 100(7) - kurrat 77 (6)
- hippocastanum 72, 100 (5, 7) - ledebourianum 28 (1)
- pavia, see A. hippocastanum 100 (7) - longicuspus, see A. sativum 71 (5)
AEROMOMUM melegueta 128 (8) - macrostemon 28 (1)
AGAVE americana 162 (11) - microbulbum, see A. chinense 27 (1)
- atrovirens 162 (11) - nipponicum 28 (1)
- cantala 162 (11) - odoratum, see A. tuberosum 28 (1)
- crassispina 162 (11) - porrum, see A. ampeloprasum, A. ramosum
- deweana 162 (11) 62, 77 (4, 6)
- fourcroydes 108, 162 (8, 11) - pskemense, see A. cepa 71 (5)
- funkiana 162 (11) - ramosum 28 (1)
- latissima, see A. atrovirens 162 (11) - sativum, see also A. ampeloprasum 28, 71,
- letonae 162 (11) 78, 91 (1, 5, 6, 7)
- rigida, see A. sisalana 162 (11) - schoenoprasum 28 (1)
- sisalana 162 (11) - seorodoprasum, see also A. ampeloprasum
- tequilana 162 (11) 129 (9)
AGRIMONLA eupatoria, see A. odorata 139(9) - tuberosum 28 (1)
- odorata 139 (9) - vavilovii 71 (5)
AGROPYRON caninum 132 (9) - wakegii, see A. chinense 27 (1)
- cristatum 132 (9) ALOCASIA cucullata 63 (4)
- desertorum, see A. cristatum 132 (9) - indica, see also A. macrorrhiza 43, 63 (2, 4)
- glauca, see A. intermedium 132 (9) - macrorrhiza, see also A. indica 43, 63 (2, 4)
- intermedium 80, 132 (6, 9) ALOË barbadensis 103 (7)
- junceum 96 (7) - vera, see A. barbadensis 103 (7)
- latiglume, see A. spicatum 175 (12) ALOPECURUS pratensis 132(9)
- michnoi, see A. cristatum 132 (9) ALPINIA chinensis 41 (1)
- pauciflorum 175 (12) - conchigera 56 (2)
- peetiniforme, see A. cristatum 132 (9) - galanga 56 (2)
- repens 132 (9) - malaccensis 56 (2)
- seribneri, see A. spicatum 175 (12) - magnifica, see Phaeomeria magnifica 56 (2)
- sibiricum, see A. cristatum 132 (9) - officinarum 41 (1)
- smithii 175 (12) - speciosa, see Phaeomeria magnifica 56 (2)
- spicatum 175 (12) ALTHAEA officinalis 88, 103, 138 (6, 7, 9)
- trachycaulum, see A, pauciflorum, A. spicatum - rosea 88, 103 (6, 7)
175 (12) ALYSICARPUS nummularifolius, see A. vaginalis
AGROTIS alba, see A. gigantea 132 (9) 118 (8)
- canina 132 (9) - rugosus 118 (8)
- gigantea, see also A. tenuis 132, 175 (9, 12) - vaginalis 118 (8)
- intermedia, see A, tenuis 132 (9) - violaceus, see A. rugosus 118 (8)
- tenuis 80, 96, 132, 175 (6, 7, 9, 12) AMARANTHUS angustifolius 62 (4)
- vulgaris, see A. tenuis 132 (9) - braunii, see A. spinosus 145 (10)
AILANTHUS vilmoriniana 39 (1) - caracasamus, see A. spinosus 145 (10)
ALBIZIA carbonaria 151 (10) - caudatus, see also A. mantegazzianus 145 (10)
- chinensis, see A. stipulata 66 (4) - cruentus, see A. dubius, A. hybridus, A. pani-
- falcata, see A. moluccana 49 (2) culatus 43, 145, 163 (2, 10, 11)
- lebbeck 49 (2) - dubius, see also S. spinosus 145 (10)
- moluccana 49 (2) - edulis, see A. caudatus, A. mantegazzianus
- montana 49 (2) 145 (10)
- stipulata 66 (4) - gangeticus 28, 43 (1, 2)
- sumatrana 49(2) - hybridus, see also A. cruentus, A. hypochon-
ALEURITES cordata 31 (1) driacus 145, 163, 173 (10, 11, 12)
- fordii 31, 175 (1, 12) - hypochondriacus 163 (11)
- moluccana 46 (2) - leucocarpus, see A. hypochondriacus 163 (11)
- montana 31 (1) - lividus 91 (7)
- trisperma 46 (2) - mangostanus 43 (2)
ALKANNA tinctoria 92 (7) - mantegazzianus 145 (10)
ALLAEANTHUS luzonicus 51 (2) - melancholicus, see A. mangostanus 43 (2)
ALLIUM altaicum, see A. chinensis 27 (1) - oleraceus, see A. lividus 91 (7)
- aobanum, see A. cepa 71 (5) - paniculatus 43 (2)
- ampeloprasum, see also A. kurrat, A. porrum - powellii, see A. hypochondriacus, A. leucocar-
62, 77, 129 (4, 6, 9) pus 163, 173 (11, 12)
AMARANTHUS quitensis, see A. caudatus, ANETHUM sowa, see A. graveolens 70, 105
A. mantegazzianus 145 (10) (4, 7)
- spinosus, see also A. dubius 145, 173 (10, 12) ANGELICA archangelica 142 (9)
- tricolor, see A. mangostanus 43 (2) - kiusiana 40 (1)
- viridus, see A. lividus 91 (7) - levisticum, see Levisticum officinale 142 (9)
AMMADAUCUS leucotrichus 105 (7) - polymorphs 40 (1)
AMMI majus 105 (7) ANISUM officinarum, see Pimpinella anisum 90
AMMOPHILA arenaria 132 (9) (6)
- arundinacea, see A. arenaria 132 (9) - vulgare, see P. anisum 90 (6)
AMOMUM aromaticum 70 (4) ANNONA cherimoia, see A. squamosa 145, 163
- cardamomum 56 (2) (10, 11)
- globosum 41 (1) - diversifolia 163 (11)
- kepulaga 56 (2) - montana 163 (11)
- krervanh 56 (2) - muricata 163 (11)
- magnificum, see Phaeomeria magnifica 56 (2) - purpurea 163 (11)
- maximum 56 (2) - reticulata, see also A. cherimoia 145, 163
- xanthioides 70 (4) (10, 11)
AMORPHOPHALLUS campanulatus 43, 63 (2, 4) - scleroderma 163 (11)
- harmandii 43 (2) - squamosa 145, 163 (10, 11)
- rivieri 43 (2) ANTHEMIS nobilis, see also Matricaria chamo-
AMPHICARPAEA monoica 177 (12) milla 130 (9)
AMYGDALUS besseriana 36, 88, 139 (1, 6, 9) - tinctoria 130 (9)
- bucharica 74 (5) ANTHOXANTHUM alpinum, see A. odoratum
- communis, see also A. besseriana, A. vavilovii 132 (9)
and Prunus ferganica 74, 89, 139 (5, 6, 9) - odoratum 132 (9)
- divaricata, see A. fenzliana 89 (6) ANTHRISCUS cerefolium 142 (9)
- fenzliana, see also A. communis 74, 89 (5, 6) ANTHYLLIS vulneraria 136 (9)
- georgica, see A. communis 74 (5) ANTIDESMA bunius 56 (2)
- kansuensis 36 (1) APIOS americana, see A. tuberosa 177 (12)
- ledebouriana 139 (9) - tuberosa 177 (12)
- mira 36 (1) APIUM ammi, see Ammi majus 105 (7)
- nairica, see A. communis 74 (5) - carvi, see Carum carvi 130 (9)
- nana, see A. besseriana 36, 139 (1, 9) - graveolens 105 (7)
- persica 36, 74, 89, 105, 139, 177 (1, 5, 6, 7, APOCYNUM sibiricum, see A. venetum 180 (?)
9, 12) - venetum 180 (?)
- petunnikowii 74 (5) AQUILEGIA vulgaris 104, 138 (7, 9)
- pumila, see A. persica 36 (1) ARACHIS africana, see A. hypogaea 151 (10)
- scoparia, see A. communis 74 (5) - asiatica, see A. hypogaea 151 (10)
- spinosissima, see also A. vavilovii 74 (5) - glabrata 151 (10)
- tangutica 74 (5) - hypogaea, see also A. villosulicarpa 118, 151
- turcomanica, see A. communis 74 (5) (8, 10)
- ulmifolia 74 (5) - monticola, see A. hypogaea 151 (10)
- urartu, see A. communis, A. fenzliana 74, 89 - nambyquarae, see A. hypogaea 151 (10)
(5, 6) - villosa, see A. hypogaea 151 (10)
- vavilovii 74 (5) - villosulicarpa, see also A. hypogaea 151 (10)
ANACARDIUM occidentale 145, 163 (10, 11) ARALIA cordata 28 (1)
- orientale, see Semecarpus anacardium 43 (2) - guilfoylei, see Nothopanax guilfoylei 44 (2)
ANACYCLUS officinarum 93 (7) - repens, see Panax repens 28 (1)
- pyrethrum 93 (7) ARBUTUS unedo 95 (7)
ANANAS ananassoides, see A. comosus 146 (10) ARCTIUM lappa 29, 130 (1, 9)
- bracteatus, see A. comosus 146 (10) ARECA catechu, see also Piper betle 53, 54 (2)
- comosus 146 (10) ARENGA pinnata 53, 69 (2, 4)
- erectifolius, see A. comosus 146 (10) ARGEMONE mexicana 170 (11)
- parguazensis 146 (10) ARISTOLOCHIA clematis 129 (9)
- sativus, see A. comosus 146 (10) ARMENIACA ansu, see A. vulgaris 36 (1)
ANDROPOGON aciculatus 46 (2) - atropurpurea, see A. dasycarpa 74 (5)
- citratus, see Cymbogon citratus 47 (2) - brigantea 139 (9)
- contortus, see Heteropogon hirtus 180 (?) - dasycarpa 74 (5)
- flexuosus, see Cymbogon flexuosus 64 (4) - mandshurica 36 (1)
- gayanus 112 (8) - mume 36 (1)
- gryllus, see Chrysopogon gryllus 98 (7) - sibirica 139 (9)
- muricatus, see Vetiveria zizanioides 48 (2) - vulgaris, see also Prunus bessyi 36, 74, 89,
- nardus, see Cymbogon nardus 47 (2) 178 (1, 5, 6, 12)
- rufus, see Hyparrhenia rufa 114 (8) ARMORACIA rusticana 131 (9)
- tectorum, see A. gayanus 112 (8) ARRACIA esculenta, see A. xanthorhiza 161 (10)
ANEMARRHENA asphodeloides 34 (1) - xanthorhiza 161 (10)
ANETHUM graveolens 70, 105 (4, 7) ARRHENATHERUM avenaceum 132 (9)
ARRHENATHERUM eliator, see A. avenaceum AVENA pilosa, see A. clauda 97 (7)

132 (9) - prostrata, see also A. damascena 97 (7)
- tuberosum 133 (9) - sativa, see also A. canariensis, A. clauda
ARTEMISIA abrotanum 130 (9) 32, 80, 97 (1, 6, 7)
- absinthium 130 (9) - septentrionalis 133 (9)
- capillaris 29 (1) - sterilis, see A. canariensis, A. sativa 97 (7)
- cina 71 (5) - strigosa, see also A. clauda, A. damascena,
- dracunculoides, see A. dracunculus 71 (5) A. prostrata, A. sativa 97, 98, 112 (7, 8)
- dracunculus 71 (5) - tuberosa, see Arrhenatherum tuberosum 133 (9)
- judaica 93 (7) - vaviloviana, see A. abyssinica, A. strigosa
- laxa 130 (9) 98, 112 (7, 8)
- maritima 130 (9) - ventricosa, see also A. sativa 97, 98 (7)
- vulgaris 130 (9) - wiestii, see A. strigosa 98 (7)
ARTOCARPUS altilis 51 (2) AVERRHOEA bilimbi 44 (2)
- blancoi, see A. altilis 51 (2) - carambola 44 (2)
- camansi 52 (2) AXONOPUS compressus 165 (11)
- champeden 52 (2) AZADIRACHTA indica, see Melia azadirachta
- communis, see A. altilis 51 (2) 51 (2)
- dimorphophylla, see A. rigidus 52 (2)
- heterophyllus 68 (4) BACCAUREA dulcis 46 (2)
- intégra, see A. heterophyllus 68 (4) - sapida 64 (4)
- integrifolia, see A. heterophyllus 68 (4) - motleyana 46 (2)
- lakoocha 52 (2) - racemosa 46 (2)
- mariannensis, see A. altilis 51 (2) BALANITES aegyptiaca 91 (7)
- rigidus 52 (2) BAMBUSA abyssinica, see Oxytenanthera abyssi-
ARUNDINARIA alpina 112 (8) nica 115 (8)
- amabilis 31 (1) - arundinacea, see also B. sinospinosa 46, 64
ARUNDO donax 96 (7) (2, 4)
ASCLEPIAS cornuti, see A. syriaca 173 (12) - asper, see Dendrocalamus asper 47 (2)
- syriaca 173 (12) - cornuta 47 (2)
ASPALATHUS cedarbergensis, see A. contami- - glaucescens 32 (1)
natus 118 (8) - multiplex 32 (1)
- contaminatus 118 (8) - nana, see B. glaucescens 32 (1)
ASPARAGUS officinalis 138 (9) - polymorpha 64 (4)
ASTRAGALUS boëticus 101 (7) - spinosa 47 (2)
- cicer 136 (9) - strictus 32, 47, 64 (1, 2, 4)
- falcatus 136 (9) - textilis 32 (1)
- glycyphyllus 136 (9) - tulda 47, 64 (2, 4)
- lotoides, see A. sinicus 34 (1) - tuldoides 21 (1)
- sinicus 34 (1) - vulgaris 47 (2)
- venosus 118 (8) BANISTERIOPSIS caapi 153 (10)
ATRIPLEX canescens 173 (12) BAPHIA nitida 118 (8)
- gardnesi, see A. canescens 173 (12) BARBAREA praecox 131 (9)
- hortensis 130 (9) - verna, see B. praecox 131 (9)
- semibaccata 57 (3) - vulgaris 131 (9)
- truncata, see A. canescens 173 (12) BARLERIA prionitis 62 (4)
ATROPA acuminata, see A. belladonna 70 (4) BAROSMA betulina 127 (8)
- baetica, see A. belladonna 105 (7) BASELLA alba, see B. rubra 44 (2)
- belladonna 70, 105 (4, 7) - cordifolia, see B. rubra 44 (2)
- martiana, see A. belladonna 105 (7) - rubra 44 (2)
AVENA abyssinica, see A. strigosa 98, 112 (7, BAUHINIA purpurea 67 (4)
- barbata, see A. strigosa 98 (7) BELAMCANDA chinensis 33 (1)
- bruhnsiana, see A. ventricosa 98 (7) BENINCASA cerifera, see B. hispida 45 (2)
- byzantina, see A. sativa 97 (7) - hispida 45 (2)
- canariensis 97 (7) BERBERIS vulgaris 129 (9)
- clauda, see also A. sativa 97 (7) BERGENIA crassifolia 142 (9)
- damascena 97 (7) BERTHOLLETIA excelsa 150 (10)
- eriantha, see A. clauda 97 (7) BETA atriplicifolia, see B. vulgaris 92 (7)
- fatua, see A. sativa, also A. septentrionalis - corolliflora 78 (6)
97, 133 (7, 9) - intermedia, see also B. lomatogona 78 (6)
- hirtula, see A. strigosa 98 (7) - lomatogona, see also B. intermedia 78 (6)
- longiglumis, see also A. sativa 97 (7) - macrocarpa, see B. vulgaris 92 (7)
- ludoviciana, see A. sativa 97 (7) - macrorrhiza 78 (6)
- macrocarpa, see A. sativa 97 (7) - maritima, see B. vulgaris 92 (7)
- magna, see A. canariensis, A. sativa 97 (7) - patellaris 92 (7)
- murphyi, see A. sativa 97 (7) - patula, see B. vulgaris 92 (7)
- nuda, see A. sativa 97 (7) - procumbens 92 (7)
BETA trigyna, see also B. intermedia, B. loma- BRASSICA parachinensis, see B. chinensis 30 (1)
togona 78 (6) - pekinensis, see B. campestris, B. chinensis
- vulgaris, see also B. patellaris, B. procum- 29, 30 (1)
bens, B. webbiana 71, 92, 130 (5, 7, 9) - rapa, see B. campestris 29 (1)
- webbiana 93 (7) - rupestris, see B. oleracea 94 (7)
BIXA orellana 146 (10) - scopularum, see B. oleracea 94 (7)
BLIGHIA sapida 127 (8) - sylvestris, see B. oleracea 94 (7)
BLUMEA balsamifera 45 (2) - tournefortii, see B. campestris, B. chinensis,
- myriocephala 45 (2) B. oleracea 30, 94, 131 (1, 7, 9)
BOEHMERIA nivea 40 (1) - villosa, see B. oleracea 94 (7)
- stipularis 181 (?) BRAYERA anthelmintica, see Hagenia abyssinica
- tenacissima, see B. nivea 40 (1) 126 (8)
- utilis, see B. nivea 40 (1) BRIDELIA micrantha 111 (8)
BOESENBERGIA pandurata 56 (2) BRITIA acida 155 (10)
BORAGO officinalis, see also Coleus amboinicus BROMAREA edulis 163 (11)
49, 92 (2, 7) BROMELIA comosa, see Ananas comosus 146 (10)
BORASSUS aethiopum, see B. flabellifer 125 (8) - pinguin 164 (11)
- flabellifer 53, 69, 125 (2, 4, 8) BROMUS arvensis, see B. erectus 133 (9)
BOUEA macrophylla 43 (2) - catharticum, see B. unioloides 149 (10)
BOUSSINGAULTIA baselloides, see B. cordifolia - erectus 133 (9)
146 (10) - haenkeanus, see B. willdenowii 149 (10)
- cordifolia 146, 163 (10, 11) - inermis 133 (9)
BOUTELOUA curtipentula 175 (12) - mango 149 (10)
- eriopoda 175 (12) - marginatus 176 (12)
- filiformis 175 (12) - schraderi, see B. unioloides 149 (10)
- gracilis 175 (12) - unioloides, see also B. willdenowii 149 (10)
BRACHIARIA brizantha 112 (8) - willdenowii 149 (10)
- decumbens 112 (8) BROUSSONETIA kazinoki 35 (1)
- deflexa 112 (8) - papyrifera 35 (1)
- mutica 112 (8) BRYONIA acuta, see B. cretica 132 (9)
- ramosa, see B. deflexa 112 (8) - alba 132 (9)
- ruziziensis 112 (8) - cretica 95, 132 (7, 9)
BRASENIA schreberi 28 (1) - dioica, see B. cretica 132 (9)
BRASILOCALAMUS pubescens 149 (10) - guinensis, see Lagenaria siceraria 110 (8)
BRASSICA, see also Raphanus sativus 95 (7) BRYOPHYLLUM pinnatum 109 (8)
- adpressa, see B. campestris, B. chinensis BUNCHOSIA armeniaca 153 (10)
30, 131 (1, 9) - costaricensis 169 (11)
- alba, see Sinapis alba 95 (7) BUNIUM bulbocatanum 142 (9)
- alboglabra 29 (1) BUTYROSPERMUM paradoxum, see B. parkii
- amarifolia, see B. nigra 131 (9) 127 (8)
- balearica, see B. oleracea 94 (7) - parkii 127 (8)
- campestris, see also B. chinensis, B. juncea,
B. narinosa, B. oleracea 29, 30, 63, 93, 94, CAESALPINIA arborea, see Peltophorum p h e r o -
109, 131 (1, 4, 7, 8, 9) carpum 50 (2)
- carinata, see also B. nigra, B. oleracea 94, CAJANUS cajan 67, 118 (4, 8)
109, 131 (7, 8, 9) - indicus, see C. cajan 118 (8)
- chinensis, see also B. campestris and B. n a r i - CALAMUS caesius 53 (2)
nosa 29, 30, 131 (1, 9) CALATHEA allounia 155 (10)
- cretica, see also B. oleracea 93 (7) CALENDULA officinalis 93 (7)
- fruticulosa, see B. campestris, B. chinensis, CALOCARPUM mammosum, see C. sapota 170 (11)
B. juncea 30, 131 (1, 9) - sapota 170 (11)
- insularis, see B. oleracea 94 (7) - viride 170 (11)
- japonica, see B. campestris and B. chinensis CALONCOBA echinata, see Oncoba echinata 111 (8)
29, 30, 131 (1, 9) CALOPHYLLUM inophyllum 48 (2)
- juncea, see also B. campestris, B. nigra 109, CALOPOGONIUM mucunoides 168 (11)
131 (8, 9) CALYSTEGIA sepium 29 (1)
- macrocarpa, see B. oleracea 94 (7) CAMASSIA leichtinii, see also C. quamash 177 (12)
- napobrassica, see also B. napus 94, 131 (7, 9) - quamash 177 (12)
- napocampestris, see B. campestris, B. napus CAMELINA alyssum, see C. sativa 79 (6)
94, 131 (7, 9) - pilosa, see C. sativa 79 (6)
- napus, see B. campestris, B. napobrassica, - sativa 79 (6)
B. oleracea 94, 131 (7, 9) CAMELLIA assamica, see C. sinensis 39 (1)
- narinosa 29, 30 (1) - irrawadiensis, see C. sinensis 39 (1)
- nigra, see also B. campestris, B. carinata, - japonica, see also C. wabiske 39 (1)
B. juncea 109, 131 (8, 9) - oleifera 39 (1)
- oleracea, see B. carinata, B. cretica, B. napo- - sasanqua, see C. oleifera 39 (1)
brassica, B. napus 79, 93, 109, 131 (6, 7, - sinensis, see also C. japonica and C. wabiska
8, 9) 39, 70 (1, 4)
CAMELLIA taliensis, see C. sinensis 39 (1) CARPOBROTUS edilis, see Mesenbrysanthemum

- thea, see C. sinensis 39, 70 (1, 4) edule 108 (8)
- wabiske, see also C. japonica and C. sinensis CARTHAMUS flavescens, see C. tinctorius 78 (6)
39, 40 (1) - oxyacantha, see C. tinctorius 78 (6)
CAMPANULA rapunculus 130 (9) - palaestinus, see C. tinctorius 78 (6)
CAMPOMANESIA cornifolia, see C. lineatifolia - persicus, see C. tinctorius 78 (6)
155 (10) - tinctorius 71, 78 (5, 6)
- guaviroba 155 (10) CARUM carvi 130 (9)
- lineatifolia 155 (10) - copticum, see also C. carvi 63, 130 (4, 9)
CANANGA odoratum 43 (2) - petroselinum, see Petroselinum crispum 106 (7)
CANARIUM album 28 (1) - roxburghianum, see also C. carvi 45, 130 (2, 9)
- commune 44 (2) CARYA alba, see C. ovata 177 (12)
- edule 151 (10) - aquatica, see C. illinoinensis 177 (12)
- moluccanum 44 (2) - cathayensis 33 (1)
- ovatum 44 (2) - illinoinensis 177 (12)
- pimela 44 (2) - lecontei, see C. illinoinensis 177 (12)
CANAVALIA boliviana, see C. ensiformis 151 (10) - ovata 177 (12)
- brasiliensis, see C. ensiformis 151 (10) - pecan, see also C. illinoinensis 167, 177 (11, 12)
- campylocarpa 168 (11) - tonkinensis, see C. cathayensis 33 (1)
- dictyota, see C. ensiformis 151 (10) CARYOCAR nuciferum 147 (10)
- ensiformis, see also C. gladiata, C. plagio- CARYOPHYLLUS sylvestris, see Eugenia c a r y o -
sperma 34, 67, 151 (1, 4, 10) phyllus 53 (2)
- gladiata 34, 49 (1, 2) CASIMIROA edulis 170 (11)
- gladiolata, see C. gladiata 49 (2) CASSIA acutifolia, see C. senna 119 (8)
- maritima, see C. ensiformis 151 (10) - angustifolia 119 (8)
- piperi, see C. ensiformis, C. plagiosperma - auriculata 67 (4)
151, 152 (10) - didymobotrya 49 (2)
- plagiosperma 152 (10) - fistula 67 (4)
- polystacha, see C. gladiata 49 (2) - florida, see C. siamea 49 (2)
- regalis 119 (8) - hirsuta 49 (2)
CANNA coccinea, see C. edulis 146 (10) - leschenaultiana 49 (2)
- edulis 146 (10) - mimosoides 49 (2)
- indica, see C. edulis 146 (10) - occidentalis 49 (2)
- paniculata, see C. edulis 146 (10) - pumila 49 (2)
- speciosa 109 (8) - senna 119 (8)
CANNABIS ruderalis, see C. sativa 63, 130 (4, 9) - siamea 49, 67 (2, 4)
- sativa 63, 130 (4, 9) - tora 49 (2)
CAPPARIS ovata, see C. spinosa 92 (7) CASTANEA crenata 31 (1)
- spinosa 92 (7) - dentata 175 (12)
CAPSELLA bursa-pastoris 95(7) - mollissima 31 (1)
CAPSICUM angulosum, see C. baceatum 157 (10) - pumila 175 (12)
- annuum 70, 142, 171 (4, 9, 11) CASTANEA sativa 79 (6)
- baceatum 157 (10) - vesca, see C. sativa 79 (6)
- chinense, see also C. frutescens 157, 171 CASTILLA elastica 169 (11)
(10, 11) CASUARINA equisetifolia 57 (3)
- frutescens, see also C. chinense 157, 171 CATARIA vulgaris, see Nepata cataria 136 (9)
(10, 11) CATHA edulis 109 (8)
- microcarpum, see C. baceatum 157 (10) CEIBA pentandra 44, 108, 163 (2, 8, 11)
- pendulum, see C. baceatum 157 (10) CELOSIA argenta 62 (4)
- pubescens 157 (10) - cristata, see C. argentea 62 (4)
- sinense, see C. chinense 157 (10) - trigyna 108 (8)
CAREX arenaria 132(9) CELTIS australis 105 (7)
- dispalata 30 (1) - excelsa, see C. australis 105 (7)
- spadicea, see C. arenaria 132 (9) CENCHRUS ciliaris 180 (?)
CARICA candamarcensis, see also C. chrysopeta- CENTAUREA benedicta, see Cnicus benedictus
la, C. pentagona 147 (10) 93 (7)
- chrysopetala 147 (10) CENTROSEMA plumieri 152 (10)
- frutifragans, see C. candamarcensis 147 (10) - pubescens 152 (10)
- x heilbornii, see C. chrysopetala 147 (10) CEPHAELIS ipecacuanha 152 (10)
- papaya 164 (11) CEPHALARIA syriaca 79 (6)
- pelta, see C. papaya 164 (11) CEPHALOSTACHYUM capitatum 64 (4)
- pentagona, see also C. chrysopetala 147 (10) CERASUS avium, see Prunus avium 89 (6)
- stipulata, see C. chrysopetala, C. pentagona - fruticosa, see Prunus fruticosa 140 (9)
147 (10) - vulgaris, see Prunus cerasus 89 (6)
CARISSA grandiflora 108 (8) CERATONIA siliqua 101 (7)
CARPOBROTUS chilensis, see Mesembryanthemum CERATOTHECA sesamoides 126 (8)
chilense 145 (10) CERCIS siliquastrum 101 (8)
CEREUS grandiflorus, see Selenicereus grandi- CITRULLUS colocynthoides, see C. lanatus 109 (8)
florus 164 (11) - colocynthis 63, 94 (4, 7)
CHAMAEDOREA tepejilote 170 (11) - edulis, see C. lanatus 109 (8)
- wendlandianum, see C. tepejilote 170 (11) - fistulosus, see C. lanatus 63 (4)
CHAMAEROPS humulis 104 (7) - lanatus 63, 109 (4, 8)
CHAENOMELES donia, see C. sinensis 36 (1) - vulgaris, see C. lanatus 109 (8)
- sinensis 36 (1) CITRUS aurantifolia, see also C. limon 54, 55 (2)
CHAEROPHYLLUM bulbosum 142 (9) - aurantium, see also C. sinensis, Poncirus
CHAYOTE edulis, see Sechium edule 165 (11) trifoliata 39, 55, 105 (1, 2, 7)
CHEIRANTHUS cheiri 95 (7) - decumanus, see C. grandis 55 (2)
CHENOPODIUM album 130 (9) - grandis, see also C. paradisi, C. reticulata
- ambrosioides 93 (7) 55, 170 (2, 11)
- bonus-henricus 130 (9) - hystrix 55 (2)
- capitata 78 (6) - ichangensis 38 (1)
- esculentus, see C. bonus-henricus 130 (9) - japonica, see Fortunella japonica 39 (1)
- foliosum 130 (9) - junos 38 (1)
- nuttalliae, see also C. quinoa 147, 164 (10, 11) - latipes 69 (4)
- pallidicaule 147 (19) - limetta 55 (2)
- quinoa, see also C. nuttalliae, Amaranthus - limon 55, 105 (2, 7)
caudatus 145, 147, 164 (10, 11) - margarita, see Fortunella margarita 39 (1)
CHIDOSCOLUS chayamansa, see Jatropha aconiti- - maxima, see C. grandis 55 (2)
folia 165 (11) - medica, see also C. aurantifolia, C. limon
CHIMONOBAMBUSA quandrangularis 32 (1) 54, 55 (2, 6)
CHLORANTHUS spicatus 29 (1) - mitis, see also C. paradisi 55, 170 (2, 11)
- inconspicuus, see C. spicatus 29 (1) - nakoor, see C. aurantifolia 54 (2)
CHLOROPHORA excelsa 124 (8) - nobilis, see C. reticulata 39, 55 (1, 2)
CHLORIS gayana 112 (8) - paradisi, see also reticulata and C. sinensis
CHROZOPHORA tinctoria 96 (7) 55, 170 (2, 11)
CHRYSANTHEMUM cinerariaefolium, see also - reticulata, see also C. aurantifolia, C. mitis,
C. coccineum 78, 93 (6, 7) C. paradisi, C. sinensis 39, 55, 170 (1, 2, 11)
- coccineum 78 (6) - sinensis, see also C. aurantifolia, C. paradisi,
- conorarium 29 (1) C. reticulata, Poncirus trifoliata 39, 55,
- parthenium 78, 93 (6, 7) 105, 170 (1, 2, 7, 11)
- segetum 29 (1) CLAUSENA dentata 69 (4)
- sinense 29 (1) - lansium 39 (1)
CHRYSOBALANUS icaco 147 (10) - willdenowii, see C. dentata 69 (4)
CHRYSOPHYLLUM africanum 127 (8) CLAYTONIA perfoliata 170 (11)
- cainito 170 (11) CLITORIA cajanifolia, see C. laurifolia 50 (2)
CHRYSOPOGON aciculatus, see Andropogon - laurifolia 50 (2)
aciculatus 46 (2) - ternatea 50 (2)
- gryllus 98 (7) CNICUS benedictus 93 (7)
CHUSQUEA andina 149 (10) CNIDOSCOLUS chayamansa, see Jatropha aconiti-
- culeon 149 (10) folia 165 (11)
- depauperata 149 (10) COCCINIA abyssinica 109 (8)
- uliginosa 149 (10) - cordifolia 63 (4)
CICER arietinum 67, 73, 85, 101 (4, 5, 6, 7) - indica, see C. cordifolia 63 (4)
- bijugum, see C. arietinum 85 (6) COCCOLOBA uvifera 170(11)
- echinospermum, see C. arietinum 85 (6) COCCULUS thunbergii 35 (1)
- jaquemontii, see C. microphyllum 73 (5) COCHLEARIA armoracia, see Armoracia r u s t i -
- microphyllum 73 (5) cana 132 (9)
- pinnatifidum, see C. arietinum 85 (6) - officinalis 132 (9)
- songaricum, see C. microphyllum 73 (5) COCOS nucifera 53, 69 (2, 4)
CICHORIUM endivia 130 (9) COELOCOCCUS armicarum 53 (2)
- intybus 130 (9) COFFEA arabica, see also C. congensis, C. euge-
CINCHONA calisaya, see C. ledgeriana 156 (10) noides, C. liberica 90, 126, 127 (6, 8)
- lancifolia, see C. ledgeriana 156 (10) - arnoldiana, see C. liberica 127 (8)
- ledgeriana 156 (10) - bengalense 69 (4)
- officinalis, see C. ledgeriana 156 (10) - canephora, see also C. arabica, C. congensis,
- pubescens, see C. ledgeriana 156 (10) C. liberica 126, 127 (8)
- succirubra, see C. ledgeriana 156 (10) - congensis, see also C. canephora 126, 127 (8)
CINNAMOMUM aromaticum, see C. cassia 49 (2) - eugenioides 127 (8)
- burmani 49 (2) - excelsa, see C. liberica 127 (8)
- camphora 34 (1) - liberica 127 (8)
- cassia 49 (2) - robusta, see C. canephora 126 (8)
- zeylanicum 34 (1) COIX lacryma-jobi 47 (2)
CISSAMPELOS owariensis 124 (8) COLA acuminata 127 (8)
CITROPSIS gilletiana 127 (8) - anomela 128 (8)
COLA nitida, see also C. verticillata 127, 128 (8) CROTALARIA goreensis 119 (8)
- verticillata 128 (8) - intermedia 119 (8)
COLEUS amboinicus 49 (2) - juncea 67 (4)
- aromaticus, see C. amboinicus 49 (2) - longirostrata 168 (11)
- barbatus, see C. forskohlii 118 (8) - mucronata 180 (?)
- dazo 118 (8) - retusa 181 (?)
- edulis 118 (8) - retzli, see C. spectabilis 119 (8)
- floribundus, see C. dazo 118 (8) - spectabilis 119 (8)
- forskohlii 118 (8) - striata, see C. mucronata 180 (?)
- langouassensis 118 (8) - usaramoensis 119 (8)
- parviflorus 49 (2) - zanzibarica, see C. usaramoensis 119 (8)
- rotundifollus 118 (8) CROTON tiglium 64 (4)
- tuberosus, see C. edulis, C. parviflorus 49, CRYOPHYTUM cristallinum, see Mesembryanthe-
118 (2, 8) mum cristallinum 108 (8)
COLOCASIA antiquorum, see C. esculenta 28 (1) CRYPTOSTEGIA grandiflora 69, 126 (4, 8)
- esculenta 28, 43 (1, 2) CRYPTOTAENIA japonica 40 (1)
COLUBRINA rufa 156 (10) CUCUMEROPSIS edulis 109 (8)
COMMIPHORA opobalsamum 108 (8) - mannii 109 (8)
CONIUM maculatum 181 (?) CUCUMIS anguria, see also C. longipes 109, 110
CONVALLARIA majalis 138 (9) (8)
CONVOLVULUS scammonia 93 (7) - conomon, see C. melo 30 (1)
- sinuata, see Merremia tuberosa 147 (10) - dipsaceus 110 (8)
COPTIS chinensis 36 (1) - hardwickii, see C. sativus 64 (4)
CORCHORUS capsularis 70 (4) - longipes, see also C. anguria 109, 110 (8)
- olitorius 40 (1) - maxima 64 (4)
- trilocularis 128 (8) - melo 30, 72, 79, 110 (1, 5, 6, 8)
CORDIA dodecandra 165 (11) - metuliferus 110 (8)
CORIANDRUM sativum 105 (7) - sativus 30, 64, 79 (1, 4, 6)
CORNUS mas 78 (6) CUCURBITA andreana, see C. maxima 147 (10)
- mascula, see C. mas 78 (6) - ecuadorensis, see C. maxima 147 (10)
CORONILLA varia 136 (9) - ficifolia, see C. maxima 147, 164 (10, 11)
COROZO oleifera 155 (10) - lundelliana, see C. ficifolia, C. maxima, C.
CORTADERIA argentea 149 (10) mixta, C. moschata, C. pepo 147, 164, 165
CORYLUS avellana, see C. tubulosa 79, 93 (6, 7) (10, 11)
- cervorum, see C. avellana 79 (6) - maxima, see also C. mixta 64, 147, 164 (4,
- chinense 29 (1) 10, 11)
- colchica, see C. avellana 79 (6) - mixta, see C. maxima 147, 164 (10, 11)
- colurna, see also C. avellana 79 (6) - moschata, see also C. maxima, C. mixta 147,
- heterophylla 29 (1) 164 (10, 11)
- iberica, see C. avellana 79 (6) - pepo, see also C. maxima, C. mixta 147, 164
- imoretica, see C. avellana 79 (6) (10, 11)
- manshurica 29 (1) - texana, see C. pepo 165 (11)
- maxima, see also C. avellana 79 (6) CUMINUM cyminum 76, 90, 105 (5, 6, 7)
- pontica, see C. avellana 79 (6) CURCUMA amada 70 (4)
- sieboldiana 29 (1) - angustifolia 70 (4)
- tubulosa 93 (7) - aromatica, see C. domestica 70 (4)
CRAMBE cordifolia 79 (6) - caesia 70 (4)
- hlspanica 95 (7) - domestica 70 (4)
- maritima 132 (9) - heyeana 56 (2)
- tatarica, see C. cordifolia 79 (6) - longa, see also C. domestica, Canna speclosa
CRASSOCEPHALUM biafrae 109 (8) 70, 108 (4, 8)
CRATAEGUS aronia, see C. azarolus 74, 105 - pierreana 56 (2)
(5, 7) - xanthorrhiza 56 (2)
- azarolus 74, 105 (5, 7) - zedoaria 70 (4)
- hupehensis 37 (1) CYAMOPSIS psoralioides, see C. tetragonoloba
- oxyacantha, see Mespilus germanica 89 (6) 67, 119 (4, 8)
- pentagyna 37 (1) - senegalensis 119 (8)
- pinnatifida, see C. pentagyna 37 (1) - tetragonoloba 67, 119 (4, 8)
- pubescens 170 (11) CYCLANTHERA pedata 165 (11)
- stipulosa, see C. pubescens 170 (11) CYDONIA oblonga 89 (6)
CRESCENTIA cujeta 163 (11) CYMBOGON citratus 47 (2)
CRITHMUM maritimum 105 (7) - flexuosus 64 (4)
CROCUS sativus 85, 100 (6, 7) - martini 64 (4)
CROTALARIA alata 50 (2) - motia, see C. martini 64 (4)
- anagyroides 152 (10) - nardus 47 (2)
- burhia 67 (4) - winterianus, see C. nardus 47 (2)
- cannabina 119 (8) CYMBOPETALUM penduliflorum 163 (11)
CYNANCHUM vincetoxicum 91, 129 (7, 9) DESMODIUM gyroides 50 (2)

CYNARA cardunculus 93 (7) - intortum 152 (10)
- scolymus 93 (7) - salicifolium 119 (8)
CYNODON aethiopicus, see C. transvaalensis - tortuosum 168 (11)
112 (8) - uncinatum 152, 168 (10, 11)
- bradleyi, see C. incompletus 112 (8) DIANTHUS caryophyllus 92 (7)
- coursii, see C. plectostachyus 112 (8) DIGITALIS lanata 92 (7)
- dactylon 65 (4) - purpurea 105, 142 (7, 9)
- incompletus 112 (8) DIGITARIA abyssinica 112 (8)
- x magenissii 112 (8) - adscendens 176 (12)
- nlemfuensis, see C. transvaalensis 112 (8) - barbinodis, see D. exilis, D. iburua 113 (8)
- plectostachyus 112 (8) - cruciata 65 (4)
- transvaalensis 112 (8) - decumbens, see also D. pentzii, D. valida
CYNOSURUS cristatus 80, 133 (6, 9) 112, 113 (8)
CYPERUS alopecuroides 95 (7) - exilis 113 (8)
- articulatus 180 (?) - iburua, see also D. tricostulata 113 (8)
- cephalotus 30 (1) - pentzii 113 (8)
- esculentus 96 (7) - sanguinalis, see also D. adscendens 65, 133,
- glomeratus 30 (1) 176 (4, 9, 12)
- iwasakii 30 (1) - tricostulata 113 (8)
- natans, see C. cephalotus 30 (1) - valida 113 (8)
- papyrus 96 (7) DINOCHLOA gigantea 47 (2)
CYPHOMANDRA betacea 157 (10) - maclellandia 47 (2)
- crassifolia, see C. betacea 157 (10) - pendulus 47 (2)
- hartwegi, see C. betacea 157 (10) DIOSCOREA abyssinica 111 (8)
CYRTOCARPA procera 163(11) - alata, see also D. ovinala 45, 111 (2, 8)
CYRTOSPERMA chamissonis 44 (2) - atropurpurea, see D. alata 45 (2)
- edule, see C. chamissonis 44 (2) - batatas, see D. opposita 31, 111 (1, 8)
- merkusii, see C. chamissonis 44 (2) - bulbifera 45, 111 (2, 8)
CYTISUS canariensis 101 (7) - cayenensis, see also D. nummularia, D. rotun-
- pallidus 101 (7) data 45, 111 (2, 8)
- proliter 101 (7) - colocasiifolia, see also D. hispida 45, 111
(2, 8)
DACRYODES edulis, see Canarium edule 108 (8) - dumentorum 111 (8)
DACTYLIS aschersoniana, see D. glomerata 133 - elephantides 111 (8)
(9) - esculenta 45 (2)
- glomerata 133 (9) - flabellifolia 45 (2)
- smithii, see D. glomerata 133 (9) - tloribunda 165 (11)
DAHLIA rosea, see D. variabilis 164 (11) - hamiltonii, see D. alata 45 (2)
- variabilis 164 (11) - heterophylla, see D. bulbifera 45 (2)
DAPHNE odorata 40 (1) - hirsuta, see D. hispida 45 (2)
- papyrifera, see Edgeworthia papyrifera 40 (1) - hirtiflora 111 (8)
DATISCA cannabina 72 (5) - hispida 45, 64 (2, 4)
DATURA metel 40 (4) - japonica 30 (1)
- stramonium 179 (12) - latifolia, see D. bulbifera 111 (8)
DAUCUS carota 76, 105, 142 (5, 7, 9) - lecardii, see D. zara 111 (8)
- sativus, see D. carota 75 (5) - liebrechtsiana 111 (8)
DENDROCALAMUS asper 47 (2) - macroura, see D. sansibariensis 111 (8)
- brandisii 47 (2) - nummularia 45 (2)
- hamiltonil 65 (4) - opposita 31 (1)
- latifolius, see Sinocalamus latiflorus 48 (2) - ovinala 111 (8)
- longispathus 65 (4) - pentaphylla 46 (2)
- merrillianus 47 (2) - persimilis, see D. alata 45 (2)
DERRIS dalbergioides 50 (2) - piperifolia 147 (10)
- elliptica, see also D. malaccensis 50 (2) - praehensilis, see also D. rotundata 111 (8)
- malaccensis 50 (2) - quartiniana 45 (2)
- microphylla 50 (2) - rotundata, see also D. praehensilis 111 (8)
- robusta 50 (2) - sagittifolia, see D. zara 111 (8)
DESCHAMPSIA caespitosa, see D. insignis 176 (12) - sansibariensis 111 (8)
- danthonioides, see D. insignis 176 (12) - semperflorens 111 (8)
- elongata, see D. insignis 176 (12) - soso 111 (8)
- holciformis, see D. insignis 176 (12) - trifida 147 (10)
- insignis 176 (12) - triphylla, see D. hispida 45 (2)
DESMODIUM adscendens 181 (?) - welwitschii, see D. sansibariensis 111 (8)
- gangeticum 181 (?) - zara 111 (8)
- cinereum 152 (10) DIOSPYROS andersonii, see D. major 31 (1)
- discolor 152 (10) - blancoi, see D. discolor 46 (2)
DIOSPYROS chinensis, see D. kaki 31 (1) ELEOCHARIS tuberosa 30(1)

- discolor 46 (2) ELETTARIA cardamonum 70 (4)
- ebenaster 165 (11) ELEUSINE africana, see also E. coracana 113 (8)
- kaki, see D. virginiana 31, 175 (1, 12) - coracana, see also E. africana, E. indica
- lotus 31, 72 (1, 5) 65, 113 (4, 8)
- major 31 (1) - indica, see also E. africana, E. coracana 113 (8)
- virginiana 175 (12) ELSHOLTZIA cristata 34 (1)
DIPOGON lignosus 119 (8) ELYMUS arenarius 32 (1)
DIPSACUS ferox, see D. sativus 79 (6) - canadensis 176 (12)
- fullonum, see D. sativus 79 (6) ENHYDRA fluctuans 45 (2)
- sativus 79 (6) - helonchu, see E. fluctuans 45 (2)
- sylvestris, see D. sativus 79 (6) ENSETE edule, see E. ventricosum 125 (8)
DIPTERYX odorata 152 (10) - ventricosum 125 (8)
DOLICHOS bengalensis, see Lablab purpureus ENTEROLOBIUM cyclocarpum 168 (11)
119 (8) ERAGROSTIS abessinica, see E. tef 114 (8)
- benthamii, see Dipogon lignosus 119 (8) - curvula 114 (8)
- biflorus, see D. uniflorus 67 (4) - superba 114 (8)
- lablab, see Lablab purpureus 119 (8) - tef 114 (8)
- gibbosum, see Dipogon lignosus 119 (8) EREMOCITRUS glaucus 61 (3)
- lignosus, see Dipogon lignosus 119 (8) ERIANTHUS, see Saccharum sinense 66 (4)
- purpureus, see Lablab purpureus 119 (8) - maximum, see Saccharum officinarum 48 (2)
- sesquipedalis, see Vigna sinensis 120 (8) ERIOBOTRYA japonica 37(1)
- sinensis, see Vigna sinensis 67 (4) ERIOCHLOA polystachya 149 (10)
- uniflorus 67 (4) ERIOGLOSSUM edule, see E. rubiginosum 55 (2)
- umbellulatus, see Phaseolus calcaratus 50 (2) - rubiginosum 55 (2)
DORYALIS hebecarpa 64 (4) ERODRJM cicutarium 96 (7)
DOVYALIS hebecarpa, see Doryalis hebecarpa - moschatum 96 (7)
64 (4) ERUCA pinnatifida 109 (8)
DRACAENA arborea 108 (8) - sativa, see E. vesicaria 63, 95 (4, 7)
- fragrans, see D. arborea 108 (8) - vesicaria 63, 96 (4, 7)
- mannii, see D. arborea 108 (8) ERVATAMIA coronaria 43 (2)
- smithii, see D. arborea 108 (8) ERVUM lens, see Lens esculenta 85 (6)
DUBOISIA hopwoodii 61 (3) ERYTHRINA glauca 152 (10)
- leichhardtii 61 (3) - micropheryx 152 (10)
- myoporoides 61 (3) - senegalensis 119 (8)
DUCHESNEA filipendula 37 (1) ERYTHROXYLUM bolivianum, see E. coca 148 (10)
DURIO dulcis, see D. oxleyanus 44 (2) - coca 148 (10)
- grandiflorus, see D. oxleyanus 44 (2) - novogratense, see also E. coca 148 (10)
- graveolens, see D. oxleyanus 44 (2) - truxillense, see E. coca 148 (10)
- kutejensis 44 (2) EUCALYPTUS alba 58 (3)
- oxleyanus 44 (2) - amygdalina, see also E. regnans 58 (3)
- zibethinus 44 (2) - astringens 58 (3)
- botryoides, see also E. camaldulensis 58 (3)
ECBALLIUM elaterium 95 (7) - brockwayi 58 (3)
ECHINOCHLOA colona, see also E. frumentacea - camaldulensis, see also E. botryoides and
32, 65 (1, 4) E. trabutii 58, 104, 155 (3, 7, 10)
- crus-galli, see also E. frumentacea 32 (1) - cinerea 59 (3)
- crus-pavonis 32 (1) - citriodora 59 (3)
- frumentacea, see also E. colona, E. crus-galli - cladocalyx 59 (3)
32, 65 (1, 4) - coccifera 59 (3)
- oryzicola, see E. crus-galli 32 (1) - conoidea, see E. leucoxylon 59 (3)
- utilis, see E. crus-galli and E. frumentacea - corynocalyx, see E. cladocalyx 59 (3)
32 (1) - crebra 59 (3)
EDGEWORTHIA papyrifera 40 (1) - cypellocarpa 59 (3)
EHRHARTA calycina 113 (8) - dalrympleana 59 (3)
ELAEAGNUS angustifolia 72 (5) - delegatensis 59 (3)
- argentea, see E. angustifolia 72 (5) - diversicolor 59 (3)
- multiflora 31 (1) - eugenioides 59 (3)
- orlentalis, see E. angustifolia 72 (5) - fergusoni, see E. paniculata 60 (3)
- pungens 31 (1) - gigantea, see E. delegatensis 59 (3)
- umbellata 31 (1) - glaucescens 59 (3)
ELAEIS guineensis, see also Corozo oleifera - globulus 59, 104 (3, 7)
125, 155 (8, 10) - gomphocephala 59 (3)
- melanococca, see Corozo oleifera 155 (10) - grandis 59 (3)
ELAEOCARPUS floribundus 46 (2) - gunnii 59 (3)
ELEOCHARIS dulcis 30 (1) - hemilampra, see E. resinifera 60 (3)
- plantaginea, see E. dulcis 30 (1) - leucoxylon 59 (3)
EUCALYPTUS macarthuri 59 (3) FESTUCA arundinacea 133 (9)

- maculata 59 (3) - elatior, see F. pratensis 133 (9)
- .iiaidenii 59 (3) - loliacea, see F. pratensis 133 (9)
- mannifera, see E. viminalis 60 (3) - ovina 133 (9)
- melliodora 59 (3) - pratensis, see also F. arundinacea, Lolium
- mierocorys 60 (3) perenne 133 (9)
- niphophila 60 (3) - rubra 133 (9)
- paniculata 60 (3) - scariosa, see F. arundinacea 133 (9)
- pauciflora 60 (3) - unioloides, see Bromus willdenowli 149 (10)
- perreniana 60 (3) FESTULOLIUM loliaceum, see Festuca pratensis,
- persicifolia, see E. viminalis 60 (3) Lolium perenne 133 (9)
- regnans, see also E. amygdalina 58, 60 (3) FICUS carica 88 (6)
- resinifera 60 (3) - elastica 68 (4)
- robusta 60 (3) - religiosa 68 (4)
- salicifolia, see E. amygdalina 58 (3) - roxburghii 68 (4)
- saligna, see also E. globulus 60 (3) - sycomorus 103 (7)
- seabra, see E. eugenioides 59 (3) - tiliaefolia 124 (8)
- sideroxylon 60 (3) FIMBRISTYLIS globulosa 45 (2)
- spectabilis, see E. resinifera 60 (3) FLACOURTIA ramontchi 46 (2)
- subalatum, see E. tereticornis 60 (3) - rukam 46 (2)
- tereticornis 60 (3) FLEMINGIA vestita, see Moghania vestita 73 (4)
- trabutii, see E. botryoides 58 (3) FOENICULUM azoricum, see F. vulgare 105 (7)
- variegata, see E. maculata 59 (3) - dulce, see F. vulgare 105 (7)
- viminalis 60 (3) - officinale, see F. vulgare 105 (7)
EUCLAENA mexicana, see Z. mexicana 167 (11) - piperitum, see F . vulgare 105 (7)
EUCOMMIA ulmoides 31 (1) - vulgare 105 (7)
EUGENIA aquea 52 (2) FORTUNELLA crassifolia 39 (1)
- cariophyllus 53 (2) - hindsii 39 (1)
- cumini, see E. jambolana 53, 69 (2, 4) - japonica 39 (1)
- cauliflora, see Myrciaria cauliflora 155 (10) - margarita, see also Citrus aurantifolia, Pon~
- dombeyana 155 (10) cirus trifoliata 39, 54 (1, 2)
- formosa 53 (2) FRAGARIA x ananassa, see also F. chiloensis,
- jambolana 53, 69 (2, 4) F. virginiana 139, 156, 178 (9, 10, 12)
- jambos 53 (2) - bucharica 75 (5)
- javanica 53 (2) - chiloensis, see also F. x ananassa 139, 156
- malaccensis 53 (2) (9, 10)
- obtusifolia, see E. caryophyllus and E. j a m - - filipendula, see Duchesnea filipendula 37 (1)
bolana 53 (2) - grandiflora, see F. x ananassa, F. virginiana
- uniflora 155 (10) 139, 178 (9)
- uvalha 155 (10) - moschata 139 (9)
EUONYMUS japonicus 29 (1) - ovalis, see F. x ananassa 139 (9)
- pulchellus, see E. japonicus 29 (1) - vesca 140 (9)
EUPATORIUM ayapana, see E. triplinerve 147 - virginiana, see also F . x ananassa 139, 178
(10) (9, 12)
- stoechadosum 45 (2) - viridis 140 (9)
- triplinerve 147 (10) FRAXINUS chinensis 35 (1)
EUPHORBIA dregeana 111 (8) - koehneana, see F. chinensis 35 (1)
- kamerunica 111 (8) - ornus 104 (7)
- lathyrus 96 (7) FRITILLARIA imperalis 73 (5)
EUPHORIA longana, see Nephelium longena 39 (1) - verticillata 34 (1)
EURYALE ferox 31 (1) FUCHSIA magellanica 155 (10)
EUTREMA wasabl 30 (1) FUNIFERA brasiliensis 160 (10)
EVODIA hortensis 181 (?) FUNTUMIA elastica 108 (8)
EXOGONIUM purga, see Ipomoea purga 164 (11) FURCRAEA andina, see F. macrophylla 145 (10)
- gigantea 145, 162 (10, 11)
FABA vulgaris, see Vicia faba 73, 103 (5, 7) - humboldtiana, see F. macrophylla 145 (10)
FAGARA euoda, see Evodia hortensis 181 (?) - macrophylla 145 (10)
- evoda, see Evodia hortensis 181 (?)
FAGOPYRUM esculentum 88 (6) GALEGA officinalis 136 (9)
- sagittatum, see F . esculentum 88 (6) - orientalis 85 (6)
- tataricum 36 (1) GARBERIA benedicta, see Cnicus benedicta 93 (7)
- vulgare, see F . esculentum 88 (6) GARCINIA atrovirides 48 (2)
FALCATA comosa, see Amphicarpaea monoica - cochinchinensis 48 (2)
177 (12) - dulcis 48 (2)
FEDIA cornucopiae 106 (7) - indica 48, 66 (2, 4)
FEIJOA sellowiana 155 (10) - mangostana, see also G. silvestris 48 (2)
FERONIA limonia 69 (4) - multiflora 48 (2)
GARCINIA pedunculata 49 (2) GOSSYPIUM somalense 124 (8)

- silvestris, see also G. mangostana 49, 66 - stocksii 68, 88 (4, 6)
(2, 4) - sturtii 58 (3)
- tinctoria 49, 66 (2, 4) - thurberi 169 (11)
- tonkinensis, see G. multiflora 49 (2) - tomentosum 154 (10)
GARDENIA florida, see G. jasminoides 38 (1) - trilobum 169 (11)
- jasminoides 38 (1) - triphyllum 124 (8)
GASTROCHILUS pandurata, see Boesenbergia - vitifolium, see G. barbadense 153 (10)
pandurata 56 (2) GREWIA asiatica 70 (4)
GIGANTOCHLOA apus 47 (2) GUAZUMA grandiflora 160 (10)
- ligulata 47 (2) GUADUA angustifolia 149 (10)
- maxima 47 (2) GUILIELMA gasipaes 155 (10)
- scortechinii 47 (2) - utilis 170 (11)
- scribneriana 47 (2) GUIZOTIA abyssinica 109 (8)
- verticillata 47 (2) GYMNEMA syringifolium 44 (2)
GINKGO biloba 31 (1) GYNANDROPSIS gynandra 109 (8)
GIRARDINIA heterophylla 70 (4) - pentaphylla, see G. gynandra 109 (8)
GLEDITSIA japonica 34 (1) GYNERIUM argenteum, see Cortaderia argentea
- triacanthos 177 (12) 149 (10)
GLEHNIA litoralis 40 (1) - sagittatum 149 (10)
GLIRICIDIA maculata, see G. sepium 152 (10) GYNURA cernua 109 (8)
- sepium 152 (10) - japonica, see G. pinnatifida 29 (1)
GLOCHIDION blancoi 46 (2) - pinnatifida 29 (1)
GLYCERIA fluitans 133 (9) GYPSOPHILA paniculata 92 (7)
GLYCINE apios, see Apios tuberosa 177 (12)
- gracilis, see G. max 34 (1) HAGENIA abyssinica 126 (8)
- hispida, see G. max 34 (1) HAKEA salicifolia 60 (3)
- javanica, see G. max 34 (1) - sericea 60 (3)
- max 34 (1) HALOGETON sativus 93 (7)
- soja 34 (1) HEDYSARUM alpium, see H. hedysaroides 136 (9)
- ussuriensis 34 (1) - coronarium 101 (7)
- wightii 119 (8) - hedysaroides 136 (9)
GLYCYRRHIZA echinata 136 (9) HELENIUM grandiflorum, see Inula helenium
- glabra 101, 136 (7, 9) 72 (5)
- glandulifera, see G. glabra 136 (9) HELIANTHUS agrophyllus, see H. annuus 173 (12)
GNETUM gnemon 46 (2) - annuus, see also H. tuberosus 130, 173, 174
GOMPHRENA globosa 180 (?) (9, 12)
GOSSYPIUM anomalum 120 (8) - bolanderi, see H. annuus 173 (12)
- arboreum, see also G. anomalum, G. barba- - debilis, see H. annuus 173 (12)
dense, G. herbaceum, G. triphyllum 35, 51, - x laetiflorus, see H. annuus 173 (12)
68, 120, 121, 124, 153, 154 (1, 2, 4, 8, 10) - petiolaris, see H. annuus 173 (12)
- areysianum 88 (6) - subrhomboideus, see H. annuus 173 (12)
- aridum 169 (11) - tuberosus, see also H. annuus 173, 175 (12)
- armourianum 169 (11) HESPERIS matronalis 132 (9)
- australe 58 (3) HESPEROYUCCA funifera 162 (11)
- barbadense, see also G. herbaceum, G. h i r s u - HETEROPOGON contortus, see H. hirtus 180 (?)
tum, G. raimondii, G. tomentosum, G. thur- - hirtus 180 (?)
beri 73, 120, 121, 153, 154, 169, 177 (5, 8, HEVEA benthamiana 148 (10)
10, 11, 12) - brasiliensis 46, 148 (2, 10)
- caicoense 154 (10) HIBISCUS acetosella, see also H. asper, H.
- gossypioides 169 (11) surattensis 124 (8)
- harknessii 169 (11) - x archeri, see H. schizopetalus 124 (8)
- herbaceum, see also G. anomalum, G. a r b o - - asper, see also H. acetosella, H. cannabinus,
reum, G. barbadense, G. hirsutum, G. thur- H. sabdariffa 124 (8)
beri, G. triphyllum 68, 73, 88, 120, 121, - cannabinus, see also H. asper, H. radiatus
124, 154, 169 (4, 5, 6, 8, 10, 11) 68, 124 (4, 8)
- hirsutum, see also G. herbaceum, G. raimon- - eetveldeanus, see H. acetosella 124 (8)
dii, G. tomentosum 51, 121, 154, 169 (2, 8, - esculentus, see Abelmoschus esculentus 68 (4)
10, 11) - furcatus, see H. radiatus 68 (4)
- incanum 88 (6) - manihot, see Abelmoschus manihot 68 (4)
- klotzschianum, see also G. gossypioides, G. - mechowii, see H. sabdariffa 124 (8)
longiocalyx 124, 154, 169 (8, 10, 11) - meeusei, see H. asper 124 (8)
- lobatum 169 (11) - moschatus, see Abelmoschus moschatus 68 (4)
- longiocalyx 124 (8) - noldeae, see H. acetosella 124 (8)
- peruvianum, see G. barbadense 153 (10) - radiatus, see also H. acetosella, H. cannabi-
- raimondii, see also G. hirsutum 154 (10) nus, H. surattensis 68, 124 (4, 8)
- robinsonii 58 (3) - rosa-sinensis, see H. schizopetalus 124 (8)
HIBISCUS sabdariffa, see also H. radiatus 124 (8) INOCARPUS edulis 50 (2)
- schizopetalus 124 (8) INULA helenium 72 (5)
- surattensis, see also H. radiatus 68, 124 (4, 8) IPOMOEA aquatica 29 (1)
- syriacus 35 (1) - batatas, see also I. tiliacea, P u e r a r i a thun-
- tetraphyllus, see Abelmoschus manihot 68 (4) bergiana 51, 147, 164 (2, 10, 11)
HINGTSHA repens, see Enhydra fluctuans 45 (2) - eriocarpa 63 (4)
HODGSONIA heteroclita, see H. macrocarpa - fastigiata, see I. tiliacea 147 (10)
30 (1) - leucantha, see I. batatas 164 (11)
- macrocarpa 30 (1) - littoralis, see I. batatas 164 (11)
HOLCUS lanatus 133 (9) - mammosa 45 (2)
HORDEUM agrlocrithon, see H. vulgare 80 (6) - purga 164 (11)
- distichum, see H. vulgare 80 (6) - reptans, see I. aquatica 29 (1)
- hexastichon, see H. vulgare 80 (6) - tiliacea, see I. batatas 147, 164 (10, 11)
- intermedium, see H. vulgare 80 (6) - trifida, see I. batatas 164 (11)
- lagunculiforme, see H. vulgare 80 (6) - tuberosa, see Merremia tuberosa 147 (10)
- spontaneum, see H. vulgare 80 (6) IRATIS canescens, see I. tinctoria 79 (6)
- vulgare 32, 80, 98 (1, 6, 7) - littoralis, see I. tinctoria 79 (6)
HOUTTUYNIA cordata 56 (2) - taurica, see I. tinctoria 79 (6)
HOVENIA dulcis 36 (1) - tinctoria 79 (6)
HUMULUS cordifolius, see H. lupulus 130 (9) IRIS ensata 33 (1)
- lupulus 130 (9) - germanica 100 (7)
HYACINTHUS orientalls 87 (6) ISCHAENUM indicum 47 (2)
HYDNOCARPUS alcalae 46 (2)
- anthelminthicus 46 (2) JASMINUM grandiflorum 69 (4)
- kurzii 46 (2) - officinale 74 (5)
- laurifolius 64 (4) - sambac 69 (4)
- wightianus, s e e H. laurifolius 64 (4) JATEORHIZA miersli, see J . palmata 124 (8)
HYDRASTIS canadensis 176 (12) - palmata 124 (8)
HYDROLEA zeylanica 49 (2) JATROPHA aconitifolia 165 (11)
HYLOCEREUS undatus 164 (11) - curcas 148 (10)
HYOSCYAMUS niger 105 (7) - multifida 148 (10)
HYPARRHENIA rufa 114 (8) - urens 148 (10)
HYPTIS spicigera 118 (8) JUGLANS ailantifolia 33 (1)
- suaveolens 167 (11) - cordiformis, see J. ailantifolia 33 (1)
HYSSOPUS officinalis 100 (7) - duclouxiana 33 (1)
- hindsii 177 (12)
ILEX intégra 28 (1) - honorei 150 (10)
- paraguariensis, see I. paraguensis 145 (10) - mandshurica 33 (1)
- paraguensis 145 (10) - mollis 167 (11)
ILLICIUM anisatum 33 (1) - nigra 177 (12)
- religiosum, see I. verum 33 (1) - regia, see Carya pecan, J. hindsii, J. honorei,
- verum 33 (1) J. mollis, Malus kirghizorum, M. sieversii
IMPATIENS balsamina 28 (1) 72, 135, 167, 177 (5, 9, 11, 12)
INDIGOFERA anil 152 (10) - sieboldiana, see J . ailantifolia 33 (1)
- suffruticosa, see I. anil 152 (10) JUNIPERUS virginiana 175 (12)
- a r r e c t a 119 (8) JUSTICIA insularis 108 (8)
- endecaphylla 119 (8) - pectolaris 148 (10)
- hirsuta 181 (?)
- indica, see I. tinctoria 119 (8) KAEMPFERIA galanga 56 (2)
- sumatrana, see I. tinctoria 119 (8) - rotunda 56 (2)
- pilosa 67 (4) KERSTINGIE LLA geocarpa 119(8)
- teysmannii 50 (2) KIGELIA africana 108 (8)
- tinctoria 119 (8) KOCHIA indica 63 (4)
INGA dulcis 168 (11) - scoparia 29 (1)
- edulis 168 (11)
- feuillei 152 (10) LABLAB niger, see L. purpureus 119 (8)
- goldmanii 168 (11) - purpureus 119 (8)
- laurina 168 (11) - uncinatus, see L. purpureus 119 (8)
- leptoloba 168 (11) LACTUA denticulata 29 (1)
- pittieri 168 (11) - indica 29 (1)
- preussii 152 (10) - quercina 131 (9)
- pterocarpa, see Peltophorum pterocarpum 50 - saligna, see L. sativa 131 (9)
(2) - sativa 131 (9)
- punctata 152 (10) - serriola, see L. sativa 131 (9)
- reticulata, see I. feuillei 152 (10) - taraxacifolia 109 (8)
INGENHOUZIA triloba, see Gossypium trilobum - virosa 93 (7)
169 (11) LAGENARIA abyssinica, see L. s i c e r a r i a 110 (8)
LAGENARIA guineënsis, see L. siceraria 110 (8) LINUM angustifolium, see L. usitatissimum 87 (6)
- rufa, see L. siceraria 110 (8) - bienne, see L. usitatissimum 87 (6)
- siceraria, see also Crescentia cujeta 110, 163 - crepitans, see L. usitatissimum 138 (9)
(8, 11) - usitatissimum 67, 73, 87, 103, 120, 138 (4, 5,
- vulgaris, see L. siceraria 110 (8) 6, 7, 8, 9)
LALLEMANTIA iberica 85 (6) LIPPIA adoensis 128 (8)
- royleana 73 (5) - citriodora 161 (10)
LANGUAS conchigera, see Alpinia conchigera - triphylla, see L. citriodora 161 (10)
56 (2) LITCHI chinensis, see Nephelum litchi 39 (1)
LANSIUM domesticum 51 (2) LITHOSPERMUM erythrorhiza, see L. officinale
LAPORTEA decumana 56 (2) 28 (1)
LAPPA arctium, see Arctium lappa 130 (9) - murasaki, see L. officinale 28 (1)
LATHYRUS alatus, see L. clymenum 101 (7) - officinale 28, 129 (1, 9)
- annuus 101 (7) LITSEA calophylla 49 (2)
- cicera 101 (7) - sebifera, see L. calophylla 49 (2)
- clymenum 101 (7) - tetranthera, see L. calophylla 49 (2)
- hirsutus 101 (7) LOBELIA inflata 173 (12)
- ochrus 101 (7) LOLIUM x hybridum, see L. perenne 133 (9)
- odoratus 101 (7) - multiflorum, see also L. perenne 98, 133 (7, 9)
- purpureus, see L. clymenum 101 (7) - perenne, see also Festuca pratensis 99, 133
- sativus 73, 101 (5, 7) (7, 9)
- sylvestris 136 (9) LONCHOCARPUS utilis 152 (10)
- tlngitanus 101 (7) LOTONONIS bainesii 119 (8)
- tuberosus 136 (9) LOTUS alpinus, see L. corniculatus 136 (9)
LAURUS nobilis 101 (7) - corniculatus 136 (9)
LAVANDULA angustifolia, see L. officinalis 100 - edulis 101 (7)
(7) - pilosus, see L. corniculatus 136 (9)
- latifolia, see also L. officinalis 100 (7) - uliginosus 136 (9)
- officinalis, see also L. latifolia 100 (7) LUCUMA bifera 170 (11)
- spica, see L. officinalis 100 (7) - nervosa 157 (10)
LAWSONIA alba 120 (8) - obovata 157 (10)
- inermis, see L. alba 120 (8) - procera 157 (10)
LECYTHIS zabucajo 150 (10) - rivicoa, see L. nervosa 157 (10)
LENS culinarus, see L. esculenta 85 (6) - salicifolia 170 (11)
- esculenta 85 (6) LUFFA acutangula, see also L. hermaphrodita
- kotschyana, see L. esculenta 85 (6) 64 (4)
- lenticula, see L. esculenta 85 (6) - aegyptiaca, see also L. acutangula 64 (4)
- nigricans, see L. esculenta 85 (6) - cylindrica, see L. aegyptiaca 64 (4)
- orientalis, see L. esculenta 85 (6) - echinata, see L. acutangula 64 (4)
LEPIDIUM latifolium 95 (7) - graveolens, see L. acutangula 64 (4)
- meyenii 147 (10) - hermaphrodita, see also L. aegyptiaca 64 (4)
- sativum 110 (8) - racemosa, see L. aegyptiaca 64 (4)
LEPIRONIA articulata 45 (2) LUNARIA japonica, see Eutrema wasabi 30 (1)
- mucronata, see L. articulata 45 (2) LUPINUS albus 101 (7)
LEPTOSPERMUM laevigatum 60 (3) - angustifolius 101 (7)
LESPEDEZA cuneata 34 (1) - bogotensis 152 (10)
- sericea, see L. cuneata 34 (1) - cosentini 57, 101 (3, 7)
- stipulacea 34 (1) - cruckshanksii, see L. mutabilis 152 (10)
- striata 34 (1) - cunninghamii, see L. mutabilis 152 (10)
LEUCAENA glauca 152 (10) - digitatus, see L. consentini 101 (7)
- Ieucocephala 168 (11) - graecum, see L. albus, L. t e r m i s 101, 102 (7)
LEVISTICUM officinale 142 (9) - hispanicum, see L. luteus 102 (7)
LIGUSTICUM bulbocastanum, see Bunium bulbo- - jugoslavicus, see L. albus 101 (7)
castanum 142 (9) - linifolius, see L. angustifolius 101 (7)
- monnieri 56 (2) - luteus 102 (7)
LIGUSTRUM japonicum 35 (1) - montanus 152 (10)
- lucidum 35 (1) - mutabilis 152 (10)
- ovalifolium 35 (1) - perennis 177 (12)
LILIUM auratum 34 (1) - pilosus, see also L. consentini 101, 102 (7)
- candidum 103 (7) - polyphyllus 177 (12)
- cordifolium 34 (1) - reticulatus, see L. angustifolius 101 (7)
- lancifolium 35 (1) - rothmaleri, see L. luteus 102 (7)
- maximowiczii 35 (1) - sativus, see L. albus 101 (7)
- tigrinum 35 (1) - taurus, see L. mutabilis 152 (10)
LIMONIA acidissima, see Feronia limonia 69 (4) - t e r m i s , see also L. albus 101, 102 (7)
LINGNANIA chungii 32 (1) - varius, see L. angustifolius, L. cosentini
L. pilosus 57, 101, 102 (3, 7) MANILKARA zapotilla, see M. achras 170 (11)
LYCINUM chinense 56 (2) - bidentata 157 (10)
LYCOPERSICON cheesmanii, see L. esculentum - elengi 55 (2)
158 (10) - hexandra 70 (4)
- chilense 158 (10) MAOUTIA puya 70 (4)
- esculentum, see also L. pimpinellifolium, MARANTA arundinacea 155 (10)
Solanum pennelll 158 (10) MARISCUS umbellatus 180 (?)
- hlrsutum 158 (10) MARSDENIA tinctoria 63 (4)
- minutum, see L. esculentum 158 (10) MATISIA cordata, see Quararibea cordata 146 (10)
- peruvianum, see also L. chilense, L. esculen- MATRICARIA chamomilla 131 (9)
tum 158 (10) MARTYNIA proboscidea, see Proboscidea louisia-
- pimpinellifolium, see also L. esculentum nica 177 (12)
158 (10) MEDICAGO borealis, see M. falcata 136 (9)
- cancellata 85 (6)
MABA major 46 (2) - coerulea, see M. sativa 86 (6)
MACADAMIA integrifolia 60 (3) - cupaniana, see M. lupulina 137 (9)
- ternifolia, see M. integrifolia 60 (3) - daghestanica 86 (6)
- tetraphylla, see M. integrifolia 60 (3) - denticulata 136 (9)
MACLURA pomifera 177(12) - dzhawakhetica 86 (6)
MADHUCA indica, see also M. longifolia 69 (4) - falcata, see also M. glomerata, M. sativa
- latifolia, see M. indica 69 (4) 86, 136, 137 (6, 9)
- longifolia 70 (4) - gaetula, see M. sativa 102 (7)
MADIA sativa 147 (10) - glandulosa, see M. falcata 136 (9)
MAJORANA hortensis 100 (7) - glomerata, see also M. falcata, M. sativa
MALABAILA secacul 90 (6) 86, 136, 137 (6, 9)
MALACHRA capitata 181 (?) - glutinosa, see M. sativa 86 (6)
MALPIGHIA coccigera, see M. glabra 153 (10) - hemicycla, see M. falcata 86, 136 (6, 9)
- glabra 153 (10) - hispida 102 (7)
- punicifolia, see M. glabra 153 (10) - leiocarpa, see M. lupulina 137 (9)
- urens 169 (11) - lupulina 137 (9)
MALUS asiatica 37 (1) - medica, see M. falcata, M. sativa 86, 136
- baccata, see also M. micromalus 37, 140 (1, 9) (6, 9)
- halliana 37 (1) - papulosa, see JVL. dzhawakhetica 86 (6)
- hupehensis 37 (1) - platycarpa, see M. falcata 136 (9)
- kirghizorum 75 (5) - polychroa, see M. sativa 86 (6)
- micromalus, see also M. spectabilis 37, 140 - prostrata, see M. falcata, M. glomerata
(1, 9) 136, 137 (9)
- praecox, see M. sylvestris 75 (5) - rhodopaea, see M. saxitilis 86 (6)
- prunifolia 89, 140 (6, 9) - romanica, see M. falcata 86, 136 (6, 9)
- pumila, see also Prunus orientalls, M. kirghi- - ruthenica, see M. falcata 136 (9)
zorum, M. sylvestris 37, 75, 89, 140 (1, 5, - sativa, see also M. denticulata, M. dzhawakheti-
6, 9) ca, M. falcata, M. glomerata, M. hemicycla,
- sieboldii 37 (1) M. saxatilis 73, 86, 102, 136, 137 (5, 6, 7, 9)
- sieversii 75 (5) - saxatilis 86 (6)
- spectabilis 37 (1) - stipularis, see M. lupulina 137 (9)
- sylvestris, see also M. pumila 75, 140 (5, 9) - tenderiensis, see M. falcata 136 (9)
- turkmenorum 89 (6) - tianschanica 73 (5)
MALVA crispa, see M. verticillata 35 (1) - trautvetteri 86 (6)
- mohileviensis, see M. verticillata 35 (1) - willdenowii, see M. lupulina 137 (9)
- pamiroalaica, see M. verticillata 35 (1) MEIBOMIA purpurea, see Desmodium tortuosum
- sylvestris 35 (1) 168 (11)
- verticillata 35 (1) MELALEUCA leucadendra, see M. quinquenervia
MAMMEA americana 150 (10) 53 (2)
MANGIFERA caesia 43 (2) - parviflora, see M. preissiana 60 (3)
- foetida 43 (2) - preissiana 60 (3)
- indica, see also M. odorata 43, 62 (2, 4) - quinquenervia 53 (2)
- odorata, see also M. indica 43, 62 (2, 4) MELIA azadirachta 51 (2)
- zeylanica, see M. indica 62 (4) - azedarach 73 (5)
MANIHOT dulcis, see M. esculenta 148 (10) - indica, see M. azadirachta 51 (2)
- esculenta 46, 111, 148, 165 (2, 8, 10, 11) - japonica, see M. azadirachta 51 (2)
- glaziovii, see also M. esculenta 148, 149 (10) - parviflora, see M. azadirachta 51 (2)
- melanobasis, see M. esculenta 148 (10) MELICOCCUS bijugatus 157(10)
- palmata, see M. esculenta 148, 165 (10, 11) MELILOTUS albus, see also M. dentatus, M.
- saxicola, see M. esculenta 148 (10) infestus 102, 137 (7, 9)
- utilissima, see M. esculenta, Dioscorea ovinala, - altissimus, see also M. macrorhizus 137 (9)
D. soso 111, 148, 165 (8, 10) - dentatus 137 (9)
MANILKARA achras 170 (11) - indicus 67 (4)
MELILOTUS infestus 102 (7) MICHELIA figo 35 (1)

- macrocarpa 102 (7) - fuscata, see M. figo 35 (1)
- macrorhizus 137 (9) MIMOSA invisa 152, 168 (10, 11)
- graveolens, see M. suaveolens 50 (2) - sepiaria 50 (2)
- officinalis, see also M. infestus 102, 137 (7, 9) MIMUSOPS balata, see Manilkara bldentata 157 (10)
- suaveolens 50 (2) MIRABILIS jalapa 155 (10)
- sulcatus 102 (7) MISCANTHUS, see Saccharum sinense 66 (4)
MELINIS minutiflora 114(8) - floridulus, see Saccharum edule, S. officinarum
MELISSA hirsuta, see M. officinalis 100(7) 48 (2)
- officinalis 100 (7) - sinensis, see S. sinense 32, 66 (1, 4)
MELOCANNA baccifora 65 (4) MITRAGYNA speciosa 54 (2)
MENTHA alopecuroides, see M. x villosa 135 (9) MOGHANIA vestita 73, 54
- aquatica, see also M. x piperita, M. x rotun- MOMORDICA balsamina 180 (?)
difolia, M. spicata, M. smithiana 100, 135 - charantia, see also M. balsamina 180 (?)
(V, 9) MONSTERA deliciosa 163 (11)
- arvensis, see also M. aquatica, M. canadensis, MONTIA perfoliata, see Claytonia perfoliata 170
M. cardiaca, M. x gentilis, M. x rotundifolia (11)
34, 100, 135, 177 (1, 7, 9, 12) MORINDA angustifolia 69 (4)
- canadensis 177 (12) - bracteata, see M. citrifolia 69 (4)
- cardiaca, see also M. arvensis 34, 135 (1, 9) - citrifolia 69 (4)
- citrata, see M. aquatica 100 (7) - trifolia 54 (2)
- cordifolia, see M. x villosa 135 (9) MORINGA oleifera, see also M. peregrina 68, 124
- crispa, see M. aquatica, M. spicata 100, 135 (4, 8)
(7, 9) - peregrina 124 (8)
- x gentilis, see also M. arvensis, M. cardiaca MORUS alba 35 (1)
34, 135 (1, 9) - nigra 73 (5)
- gratissima, see M. x villosa 135 (9) MUCXJNA aterrima 50 (2)
- japonica, see M. arvensis, M. x rotundifolia - capitata 50, 67 (2, 4)
34, 135 (1, 9) - cochinchinensis 50 (2)
- longifolia, see also M. x rotundifolia, M. s p i - - deeringianum 50 (2)
cata 100, 135 (7, 9) - hassjoo 34 (1)
- x piperita, see M. aquatica 100, 135 (7, 9) - nivea, see M. cochinchinensis 50 (2)
- pulegium 100 (7) - pachylobia 67 (4)
- rotundifolia, see M. longifolia", M. suaveolens - pruriens 50 (2)
100, 135 (7, 9) - utilis 67 (4)
- x rotundifolia 135 (9) MUNTINGIA calabura 175 (12)
- rubra, see M. x smithiana 135 (9) MURRAYA exotica 55 (2)
- sativa, see M. x gentilis 135 (9) - koenigii 69 (4)
- x smithiana 135 (9) - paniculata 55 (2)
- spicata, see also M. aquatica, M. x gentilis, MUSA 52, 53, 58, 68, 125, 155 (2, 3, 4, 8, 10)
M. longifolia, M. x piperita, M. x rotundi- - acuminata, see Musa 52, 53, 58 (2, 3)
folia, M. x smithiana, M. x villosa 100, 135 - aiori, see Musa 58 (3)
(7, 9) - balbisiana, see also Musa, M. textilis 52, 53,
- suaveolens, see also M. spicata, M. x villosa 69 (2, 4)
135 (9) - basjoo 35 (1)
- sylvestris, see M. longifolia 100 (7) - cavendishii, see Musa 52 (2)
- velutina, see M. x villosa 135 (9) - fehi, see Musa 58 (3)
- x villosa, see also M. spicata 135 (9) - x paradisiaca, see Musa 52 (2)
- viridis, see M. spicata 135 (9) - x sapientum, see Musa 52 (2)
MERREMIA macrocarpa, see also M. tuberosa - seemanii, see Musa 58 (3)
147 (10) - sinensis, see Musa 52 (2)
- tuberosa 147 (10) - textilis 53 (2)
MESEMBRYANTHEMUM angulatum 108 (8) - troglodytarum, see Musa 58 (3)
- chilense 145 (10) MYRCIARIA cauliflora 155 (10)
- cristallinum 108 (8) - jaboticaba 155 (10)
- edule 108 (8) MYRIBALAN bellirica, -see Terminalia bellirica
MESPILUS aucuparia, see Sorbus aucuparia 141 44 (2)
(9) MYRICA nagi, see M. rubra 35 (1)
- germanica 89 (6) - rubra 35 (1)
MESSERSCHMIDIA argentea, see Tournefortia MYRISTICA argentea 53 (2)
argentea 44 (2) - fragrans 53 (2)
MESUA ferrea 66 (4) MYROXYLON balsamum 152 (10)
METHYSTICODENDRON amesianum 158 (10) MYRRHIS odorata 142 (9)
METROXYLON rumphii, see M. sagu 54 (2) MYRTUS communis 104 (7)
- sagu 54 (2)
MEUM anthamanticum 106 (7) NARCISSUS jonquilla 91 (7)
MICHELIA champaca 51 (2) - poeticus 91 (7)
NASTURTIUM indicum 30 (1) 100 (7)

- microphyllum, see N. officinalis 132 (9) ORIGANUM vulgare 136 (9)
- officinalis 132 (9) ORNITHOPUS compressus 102 (7)
NECTANDRA cinnamomoides 150 (10) - isthmocarpus, see O. sativus 102 (7)
NELUMBO nucifera 88(6) - macrorrhynchus, see O. sativus 102 (7)
NEOHOUZEAUA dullosa 65 (4) - roseus, see O. sativus 102 (7)
NEPETA cataria 136 (9) - sativus, see also O. compressus 102 (7)
NEPHELIUM lapacceum 55 (2) OROBUS lathyroides, see Vicia unijuga 34(1)
- litchi 39 (1) ORTHOSIPHON rubicundus 118 (8)
- longana 39 (1) - stamineus 49 (2)
- mutabile 55 (2) ORYZA alta, see also O. grandiglumis 149, 165
NEPTUNIA oleracea 50 (2) (10, 11)
NERIUM indicum 63 (4) - australiensis 57 (3)
- odorum, see N. indicum 63 (4) - barthii, see O. glaberrima, 114 (8)
- oleander 91 (7) - brachyantha 114 (8)
NICOTIANA debneyi 61 (3) - breviligulata, see O. perennis 115 (8)
- exigua, see N. debneyi 61 (3) - cubensis, see O. perennis 165 (11)
- goodspeedii 61 (3) - eichingeri, see also O. minuta 47, 114 (2, 8)
- octophora, see N. tabacum 158 (10) - fatua, see O. nivara and O. rufipogon 47 (2)
- paniculata, see N. rustica 158 (10) - glaberrima, see also O. breviligulata and
- quadrivalvis 179 (12) O. sativa 66, 115 (4, 8)
- rustica 158 (10) - grandiglumis 149 (10)
- rotundifolia, see N. goodspeedii 61 (3) - granulata 47 (2)
- suaveolens, see N. goodspeedii 61 (3) - latifolia 149, 165 (10, 11)
- sylvestris, see N. tabacum 158 (10) - longiglumis 47 (2)
- tabacum, see also N. quadrivalvis 158, 179 - longistaminata, see also O. glaberrima, O.
(10, 12) perennis 114, 115 (8)
- undulata, see N. rustica 158 (10) - malampuzhaensis 65 (4)
NIGELLA sativa 104, 138 (7, 9) - meyeriana, see O. granulata 47 (2)
NIPA fruticans, see Nypa fruticans 54 (2) - minuta, see also O. eichingeri 47, 114 (2, 8)
NOPALEA cochenillifera 164 (11) - montana, see O. rufipogon 47 (2)
- dejecta 164 (11) - nivara, see also O. rufipogon, O. sativa 47,
NOTHOPANAX fruticosum 44 (2) 65 (2, 4)
- guilfoylei 44 (2) - officinalis 47 (2)
- obtusum 44 (2) - perennis, see also O. glaberrima, O. rufipogon,
- pinnatum 44 (2) O. sativa 47, 65, 114, 115, 165 (2, 8, 11)
NYCTANTHES a r b o r - t r i s t i s 70 (4) - punctata, see also O. eichingeri 114, 115 (8)
NYPA fruticans 54 (2) - ridleyi 47 (2)
- rufipogon, see also O. nivara, O. perennis,
OCHLANDRA travancorica 65 (4) O. sativa 47, 65 (2, 4)
OCIMUM americanum, see O. basilicum 49 (2) - sativa, see also O. glaberrima, O. nivara,
- asperum, see Coleus forskohlii 118 (8) O. perennis, O. rufipogon 32, 47, 48, 65,
- basilicum 49 (2) 114, 115, 165 (1, 2, 4, 8, 11)
- grandiflorum, see Orthosiphon stamineus 49 (2) - schlechteri 48 (2)
- gratissimum 49 (2) - schweinfurthiana, see O. eichingeri 114 (8)
- kilimandscharicum 118 (8) - stapfii, see O. glaberrima 114 (8)
- sanctum 49 (2) OSMANTHUS fragrans 35 (1)
OENANTHE javanica 56 (2) OXALIS deppei, see O. tuberosa 155 (10)
- stolonifera, see O. javanica 56 (2) - tuberosa 155 (10)
OENOCARPUS bacaba 155 (10) OXYTENANTHERA abyssinica 115 (8)
OENOTHERA biennis 170 (11)
OLDENLANDIA umbellata 54 (2) PACHYLOBUS edulis, see Canarium edule 108 (8)
OLEA chrysophylla 104 (7) PACHYRHIZUS angulatus, see P. erosus 168 (11)
- europaea 104 (7) - apiha 152 (10)
OMPHALEA megacarpa 149 (10) - bulbosus, see P . erosus 168 (11)
ONAGRA biennis, see Oenothera biennis 170 (11) - erosus 168 (11)
ONCOBA echinata 111 (8) - palmatilobus, see P. erosus 168 (11)
ONOBRYCHIS altissima 86 (6) - tuberosus 152 (10)
- sativa, see O. viciifolia 137 (9) PAEONIA officinalis 138 (9)
- transcaucasia, see O. viciifolia 86 (6) PALAQUIUM gutta 56 (2)
- viciifolia 86, 137 (6, 9) PANAX fruticosum, see Nothopanax fruticosum
OPHIOPOGON spicatus 35 (1) 44 (2)
OPUNTIA castillae, see O. megacantha 164 (11) - ginseng 28 (1)
- ficus-indica 164 (11) - guilfoylei, see Nothopanax guilfoylei 44 (2)
- megacantha 164 (11) - obtusum, see Nothopanax obtusum 44 (2)
OREOBAMBOS buchwaldii 114 (8) - pinnatum, see Nothopanax pinnatum 44 (2)
ORIGANUM majorana, see Majorana hortensis - quinquefolia 173 (12)
PANAX repens 28 (1) PELARGONIUM radula, see P. x asperum 111 (8)

PANDANUS brosimas 54 (2) - rigidum, see P . crispum 112 (8)
- inermis, see P. spurius 54 (2) - roseum, see P. x asperum, P . denticulatum
- laevis, see P . spurius 54 (2) 111, 112 (8)
- mosehatus, see P. spurius 54 (2) - terebinthinaceum, see P . graveolens 112 (8)
- odoratissimus, see P . spurius 54 (2) - tomentosum, see also P . graveolens 112 (8)
- odorus 54 (2) PELTOPHORUM pterocarpum 50(2)
- spurius 54 (2) PENNISETUM anchylochaete 115 (8)
- tectorius, see P. spurius 54 (2) - cenchroides, see Cenchrus ciliarus 181 (?)
- utilis 126 (8) - clandestinum 115 (8)
- whitmeeanus 54 (2) - echinurus 115 (8)
PANGIUM edule 46 (2) - gibbosum 115 (8)
PANICUM barbinode, see Brachiaria mutica 112 - leonis 115 (8)
- coloratum 115 (8) - maiwa 115 (8)
- frumentacea, see Echinochloa frumentacea 32 - malacochaete 116 (8)
(1) - nigritarum 116 (8)
- italicum, see Setaria italica 33 (1) - niloticum 116 (8)
- maximum 115 (8) - perspeciosum 116 (8)
- miliaceum 32 (1) - purpureum, see also P . typhoides 116 (8)
- miliare, see P . sumatrense 115 (8) - pycnostachyum 116 (8)
- psilopodium, see P. sumatrense 115 (8) - robustum 116 (8)
- sonorum 165 (11) - spicatum, see also P . typhoides 116 (8)
- spontaneum, see P. miliaceum 32 (1) - typhoides, see also P. purpureum 116 (8)
- sumatrense 115 (8) - unisetum 116 (8)
- virgatum 165, 176 (11, 12) - vulpinum 116 (8)
- zizanoides, see Acroceras ampleetans 112 (8) PENTAPHRAGMA begoniaefolium 54 (2)
PAPAVER setigerum, see P . somniferum 74, PEPEROMIA pellucida 156 (10)
104 (5, 7) PERILLA arguta 34 (1)
- somniferum 74, 88, 104 (5, 6, 7) - crispa, see P . fruteseens 34 (1)
PARMENTIERA cereifera 163 (11) - fruteseens 34 (1)
- edulis 163 (11) - ocymoides, see P . fruteseens 34 (1)
PARKIA speciosa 50 (2) PERESKIA aculeata 146 (10)
PASPALUM commersonii, see P. scrobiculatum PERSEA americana 168 (11)
66 (4) - drymifolia, see P . americana 168 (11)
- dilatum 149 (10) - leiogyna 150 (10)
- distichum 176 (12) - schiedeana 168 (11)
- juergensii, see P . dilatum 149 (10) PERSICA davidiana, see Prunus davidiana 37 (1)
- notatum 149 (10) - kansuensis, see Amygdalus kansiensis 36 (1)
- plicatulum 149 (10) - mira, see Amygdalus mira 36 (1)
- scrobiculatum 48, 66, 115 (2, 4, 8) - vulgaris, see Amygdalus persica 36 (1)
PASSIFLORA alata 156 (10) PETASITES japonicus 29 (1)
- antiquiensis 156 (10) PETROSELINUM crispum 106 (7)
- cearensis 156 (10) PEUCEDANUM cervaria 142 (9)
- edulis 156 (10) - graveolens, see Anethum graveolens 70 (4)
- foetida 156 (10) - ostruthium 142 (9)
- incarnata 177 (12) PHACELIA tanacetifolia 176 (12)
- laurifolia 156 (10) PHAEOMERIA magnifica 56 (2)
- ligularis 156 (10) PHALARIS arundinacea 134 (9)
- maliformis 156 (10) - canariensis 99 (7)
- mollissima 156 (10) - tuberosa 99 (7)
- psilantha 156 (10) PHASEOLUS aborigineus, see also P. vulgaris
- quadrangularis 156 (10) 152, 168 (10, 11)
- tripartita 156 (10) - aconitifolius 67 (4)
- van-volxemii, see P. antiquiensis 156 (10) - acutifolius 168 (11)
PASTINACA sativa 142 (9) - angularis 34 (1)
PAULLINIA cupana 157 (10) - aureus, see P. mungo 50, 67 (2, 4)
PAYENA leerii 56 (2) - calcaratus 50 (2)
PELARGONIUM x asperum 111 (8) - chinensis, see P . vulgaris 34 (1)
- capitatum, see P. graveolens 112 (8) - coccineus, see also P . vulgaris 168 (11)
- crispum 112 (8) - lathyroides 57 (3)
- denticulatum, see also P. x asperum 111, 112 - lunatus 153, 168 (10, 11)
(8) - mungo, see also P . aureus 50, 67 (2, 4)
- graveolens, see also P . x asperum, P. tomen- - radiatus, see P. aureus 50 (2)
tosum 111, 112 (8) - retusus 168 (11)
- karooense 112 (8) - sublobatus, see P. mungo 67 (4)
- odoratissimum 112 (8) - trilobus, see P . aconitifolius 67 (4)
- radens, see P . x asperum 111 (8)
PHASEOLUS trinervis, see P . mungo 67 (4) PISUM humile, see P . sativum 86 (6)
- vulgaris, see also P . aborigineus, P . cocci- - jomardi, see P . sativum 102 (7)
neus 34, 152, 153, 168 (1, 10, 11) - sativum, see also P. formosum, Lupinus muta-
PHELLOPTERUS littoralis 40 (1) bilis 86, 102, 119, 152 (6, 7, 8, 10)
PHILODENDRON pertusum, see Monstera deli- - syriacum, see P . sativum 86 (6)
ciosa 163 (11) - transcaucasium 102 (7)
PHLEUM alpinum, see P . pratense 134 (9) PITHECELLOBIUM bigeminum 50 (2)
- nodosum, see P . pratense 134 (9) - dulce 169 (11)
- pratense 134 (9) - jiringa 50 (2)
PHOENIX atlantica 125 (8) - lolatum 50 (2)
- canariensis 125 (8) - saman 153 (10)
- dactylifera, see also P . atlantiea, P . cana- PLANTAGO decumbens, see P . ovata 69 (4)
riensis, P. sylvestris 125 (8) - indica 104 (7)
- humilis, see also P . reclinata 125 (8) - major 36 (1)
- jubae, see P . canariensis 125 (8) - ovata 69 (4)
- reclinata 125 (8) - psyllium 104 (7)
- sylvestris 125 (8) PLATYCODON grandiflorum 29 (1)
PHORMIUM tenax 57 (3) PLECTRANTHUS esculentus, see Coleus dazo
PHRAGMITES communis 134 (9) 118 (8)
PHYLLANTHUS acides, see P. distichus 46 (2) - rotundifolius, see Coleus rotundifolius 118 (8)
- distichus 46 (2) - tuberosus, see Orthosiphon rubicundus 118 (9)
- emblica 46 (2) PLUKENETIA conophora 111 (8)
PHYLLOSTACHYS bambusoides 32 (1) - corniculata 46 (2)
- dulcis 32 (1) - peruviana, see P . volubilis 149 (10)
- henonis 32 (1) - volubilis 149 (10)
- makino 32 (1) PLUMERIA acuminata, see P. acutifolia 145 (10)
- meyeri 32 (1) - acutifolia 145 (10)
- nigra, see P . henonis 32 (1) - obtusa, see P. acutifolia 145 (10)
- pubescens 33 (1) POA abyssinica, see Eragrostis tef 114 (8)
- viridis 33 (1) - ampla, see P. pratensis 176 (12)
PHYSALIS alkekengi 142 (9) - bulbosa 82, 134 (6, 9)
- ixocarpa, see also Lycopersicon esculentum - compressa 176 (12)
158, 171 (10, 11) - nemoralis 134 (12)
- peruviana 158 (10) - palustris 134 (9)
- pubescens 179 (12) - pratensis, see also P. trivialis 134, 165 (9, 12)
PHYSOSTIGMA venenosum 119 (8) - trivialis, see also P. pratensis 134 (9)
PHYTOLACCA acinosa 35 (1) POGOSTEMON cablin 49 (2)
- americana 177 (12) POLAKOWSKIA tacacco 165 (11)
- chilensis 156 (10) POLIANTHES gracilis, see P . tuberosa 162 (11)
- dioica 156 (10) - tuberosa 162 (11)
PIMENTA acris 53 (2) POLYGALA butyracea 126 (8)
- dioica 170 (11) POLYGONUM fagopyrum, see Fagopyrum esculen-
- officinalis, see P . dioica 170 (11) tum 88 (6)
PIMPINELLA anisum 90 (6) - hydropiper 36 (1)
PINUS pinea 104 (7) - maximowiczii 36 (1)
PIPER aduncum 156 (10) - odoratum 54 (2)
- betle 54 (2) - tinctorium 36 (1)
- clusii 126 (8) POLYMNIA edulis, see P . sonchifolia 147 (10)
- cuceba 54 (2) - sonchifolia 147 (10)
- guineense 126 (8) POLYSCIAS fruticosa, see Nothopanax fruticosum
- lohot, see P . saigonense 54 (2) 44 (2)
- longum, see P . retrofractum 54, 69 (2, 4) - obtusa, see Nothopanax obtusum 44 (2)
- methysticum 54 (2) - rumphiana, see Nothopanax pinnatum 44 (2)
- nigrum 69 (4) POMETIA pinnata 55 (2)
- officinarum, see P . retrofractum 54 (2) PONCIRUS trifoliata, see Citrus sinensis 39, 55
- retrofractum, see also P. longum 54, 69 (2, 4) (1, 2)
- saigonense 54 (2) PORTULACA oleracea 138 (9)
PISONIA alba 53 (2) POUTERIA caimita 157 (10)
- grandis, see P . alba 53 (2) - campechiana, see Lucuma nervosa, L. s a l i c i -
- sylvestris, see P. alba 53 (2) folia 157, 180 (10, 11)
PISTACIA lentiscus 104 (7) - lucuma, see Lucuma obovata 157 (10)
- terebinthus 104 (7) PRITCHARDIA gaudichaudii 54 (2)
- vera 74 (5) - pacifica 54 (2)
PISTIA stratiotes 44 (2) PROBOSCIDEA louisianica 177 (12)
PISUM abyssinicum, see P . sativum 119 (8) PROSOPIS juliflora 153 (10)
- arvense, see P. sativum 86 (6) PRUNUS 37, 75, 178 (1, 5, 12)
- elatius, see P . sativum 86, 102 (6, 7) - acuminata, see P . maritima 178 (12)
PRUNUS americana, see also P. hortulana 178 PRUNUS vavilovii, see Amygdalus vavilovii 74 (5)
(12) - virginiana 178 (12)
- amygdalis, see Amygdalis communis 74, 89 (5, 6) PSEUDANANAS macrodontes 146 (10)
- angustifolia, see also P. hortulana, P. munso- - sagenarius, see Ananas comosus 146 (10)
niana 178 (12) PSIDRJM cattleianum, see P. littorale 155 (10)
- ansu, see Armeniaca vulgaris 36 (1) - friedrichsthalianum 170 (11)
- armeniaca, see Armeniaca mandshurica, A. - guajava 170 (11)
sibirica, A. vulgaris 36, 139 (1, 9) - guineense 155 (10)
- avium, see also P . cerasus 89 (6) - littorale 155 (10)
- bessyi 178 (12) - molle 170 (11)
- brigantina, see Armenica brigantina 139 (9) - sartorianum 170 (11)
- bucharica, see Amygdalus bucharica 74 (5) PSOPHOCARPUS tetragonolobus 50 (2)
- cantabrigiensis 37 (1) - longepedunculatus, see P . palustris 120 (8)
- cerasifera, see also P . domestica, P. fergani- - palustris 120 (8)
ca, P. salicina, P . ussuriensis 37, 38, 75, PSORALEA bituminosa 102 (7)
89 (1, 5, 6) PTEROCOCCUS corniculatus, see Pluketia corni-
- cerasus, see also P . avium, P. fruticosa 90, culata 46 (2)
140 (6, 9) PUCCINELLIA nuttalliana 176 (12)
- chicasa, see P . angustifolia 178 (12) PUERARIA lobata, see P. thunbergiana 51 (2)
- dasycarpa, see Armeniaca dasycarpa 74 (5) - phaseoloides 51 (2)
- davidiana 37 (1) - thunbergiana 34, 51 (1, 2)
- dehiscens, see Amygdalus tangutica 74 (5) PUGIONUM cornutum 30 (1)
- divaricata, see P . cerasifera 89 (6) PUNICA granatum 88 (6)
- domestica, see also P. cerasifera, P . davidiana, - protopunica, see P . granatum 88 (6)
P. insititia, P. spinosa 37, 90, 140 (1, 6, 9) PYRACANTHA coccinea 140 (9)
- fenzliana, see Amygdalus communis, A. fenz- PYRETHRUM sinense, see Chrysanthemum sinen-
liana 74, 89 (5, 6) se 29 (1)
- ferganica 75 (5) PYRUS 90 (6)
- fruticosa, see also P. cerasus, P . spinosa - amygdaliformis, see P . communis 140 (9)
90, 140 (6, 9) - baccata, see Malus baccata 37 (1)
- x gondounii, see P . avium 89 (6) - betulaefolia 38 (1)
- hortulana 178 (12) - bretschneideri 38 (1)
- insititia 140 (9) - bucharica, see also P. korshinskyi 75 (5)
- kansuensis, see Amygdalus kansuensis 36 (1) - calleryana 38 (1)
- ledebouriana, see Amygdalus ledebouriana - caucasia, see P . communis 90, 140 (6, 9)
139 (9) - communis, see also P . pyrifolia, P . u s s u r i e n -
- mahaleb 140 (9) sis, P . vavilovii 38, 75, 140 (1, 5, 9)
- mandshurica, see Armeniaca mandshurica - cordata 141 (9)
36 (1) - domestica, see P . communis 140 (9)
- maritima 178 (12) - hupehensis, see Malus hupehensis 37 (1)
- mume, see Armeniaca mume 36 (1) - korshinskyi, see also P. bucharica 75 (5)
- munsoniana, see also P . angustifolia 178 (12) - longipes, see P . communis 140 (9)
- nana, see Amygdalus besseriana 139 (9) - malus, see Malus pumila 140 (9)
- nigra 178 (12) - nivalis, see P. communis 140 (9)
- paniculata, see P . pseudocerasus 37 (1) - phaeocarpa 38 (1)
- pensylvanica 178 (12) - prunifolia, see Malus prunifolia 89, 140 (6, 9)
- persica, see Amygdalus persica, P. davidiana - pyraster, see P. communis 140 (9)
36, 37 (1) - pyrifolia 38 (1)
- persicifolia, see P. pensylvanica 178 (12) - regelii 75 (5)
- petunnikowii, see Amygdalus petunnikowii 74 (5) - salicifolia, see P . communis 140 (9)
- pseudocerasus, see also P . cantabrigiensis - salvifolia, see P. communis 140 (9)
37 (1) - serotina, see P . communis, P. phaeocarpa
- salicina 37 (1) 38, 140 (1, 9)
- sargentii 37 (1) - sogdania 75 (5)
- scoparia, see Amygdalus communis 74 (5) - syriaca, see P. communis 90 (6, 9)
- serotina 178 (12) - takhtadzhiana 90 (6)
- sibirica, see Armeniaca sibirica 139 (9) - toringo, see Malus sieboldii 37 (1)
- simonii 38 (1) - ussuriensis, see P . communis 38, 140 (1, 9)
- spinosa, see also P . domestica 90 (6) - vavilovii 75 (5)
- spinosissima, see Amygdalus spinosissima 74 (5)
- tenella, see Amygdalus besseriana 139 (9) QUARARIBEA cordata 146 (10)
- tiliaefolia, see Armeniaca vulgaris 36, 74 (1, 5) QUASSIA amara 157 (10)
- tomentosa 38 (1) QUERCUS aliéna 31 (1)
- trichocarpa, see P . tomentosa 38 (1) - dentata 31 (1)
- triflora, see P . simonii, P . ussuriensis 38 (1) - mongolica 31 (1)
- triloba, see Amygdalus ulmifolia 74 (5) - suber 96 (7)
- ussuriensis 38 (1) QUISQUALIS indica 44 (2)
RAPHANUS acanthiformis, see R. sativus 30 (1) ROLLINIA longifolia 145 (10)

- caudatus, see R. sativus 63, 95 (4, 7) ROLLINIOPSIS discreta 145 (10)
- chinensis, see R. sativus 95 (7) RORIPPA nasturtium-aquaticum, see Nasturtium
- landra, see R. sativus 30, 95 (1, 7) officinalis 132 (9)
- maritimus, see R. sativus 30, 95 (1, 7) ROSA x alba, see also R. arvensis, R. bifera,
- raphanistrum, see R. sativus 30, 95 (1, 7) R. gallica 141 (9)
- rostratus, see R. sativus 30 (1) - arvensis 141 (9)
- sativus 30, 63, 95 (1, 4, 7) - x bifera, see also R. gallica, R. moschata
RAPHIA hookeri 126 (8) 75, 141 (5, 9)
RAUVOLFIA serpentina 63 (4) - canina 141 (9)
RESEDA luteola 104, 138 (7, 9) - centifolia 90 (6)
- odorata 104 (7) - damascena, see R. x bifera 141 (9)
- phyteuma 88, 104 (6, 7) - eglanteria, see R. rubiginosa 141 (9)
RHAMNUS catharticus 104, 138 (7, 9) - gallica, see also R. x alba, R. x bifera, R.
- frangula 104, 139 (7, 9) centifolia 90, 141 (6, 9)
RHEEDIA acuminata 150 (10) - moschata, see also R. x bifera 75, 141 (5, 9)
RHEUM hybridum, see also P . rhaponticum 36, - multiflora 38 (1)
74 (1, 5) - rubiginosa 141 (9)
- officinale 36 (1) - rugosa 38 (1)
- palmatum, see R. hybridum, R. rhaponticum - villosa 141 (9)
36, 74 (1, 5) ROSMARINUS officinalis 100 (7)
- rhabarbarum, see R. palmatum, R. rhaponti- RUBIA cordifolia 69 (4)
cum 36 (1) - tinctorum 141 (9)
- rhaponticum 36, 74 (1, 5) RUBUS albescens 54, 69 (2, 4)
- undulatum 36 (1) - acaulis, see also R. stellatus 178 (12)
RHODOMYRTUS tormentosa 53, 69 (2, 4) - argutus, see R. idaeus 178 (12)
RHUS coriaria 91 (7) - arcticus, see also R. acaulis, R. idaeus, R.
- succedanea 28 (1) parviflorus, R. stellatus 141, 178 (9, 12)
- vernicifera 28 (1) - baileyanus, see R. idaeus 178 (12)
- verniciflua, see R. vernicifera 28 (1) - brasiliensis 156 (10)
RHYNCHOSIA minima 153 (10) - chamaemorus, see also R. idaeus 141 (9)
RIBES alpestris 33 (1) - ellipticus, see R. rosaefolius 54 (2)
- americanum, see R. nigrum 135 (9) - flagellaris 178 (12)
- acicularis 135 (9) - glaucus 156 (10)
- cereum, see R. grossularia 135 (9) - idaeus, see also R. arcticus, R. chamaemorus,
- cynosbati, see also R. grossularia 135, 176 R. occidentalis, R. pungens 38, 141, 178
(9) a , 9, 12)
- dikuscha, see R. nigrum 135 (9) - illecebrosus 38 (1)
- divarieatum, see R. grossularia 135 (9) - laciniatus 141 (9)
- grossularia 135 (9) - macrocarpus 156 (10)
- hirtellum, see R. grossularia 135 (9) - neglectus, see R. idaeus, R. occidentalis 178
- longeracemosum 33 (1) (12)
- multiflorum, see also R. sativum 100, 135 (7, - occidentalis, see also R. idaeus 178 (12)
9) - palmatus, see R. idaeus 178 (12)
- nigrum, see also R. longeracemosum, R. u s s u - - parviflorus, see R. idaeus 141 (9)
riense 33 (1) - phoenieolasius 38 (1)
- niveum, see also R. grossularia 139 (9) - probus, see R. rosaefolius 54 (2)
- odoratum 176 (12) - procerus, see R. laciniatus 141 (9)
- oxyacanthoides, see R. grossularia 135 (9) - pungens 38 (1)
- petraeum, see also<*l. sativum 135 (9) - rosaefolius 54 (2)
- pinetorum, see R. grossularia 135 (9) - sachalinensis, see R. idaeus 141 (9)
- roezlii, see R. grossularia 135 (9) - saxatilis 141 (9)
- rubrum, see R. sativum 135 (9) - stellatus, see also R. acaulis 178 (12)
- sanguineum, see R. grossularia 135 (9) - strigosus, see R. idaeus 178 (12)
- sativum 135 (9) - ursinus, see R. idaeus 178 (12)
- spicatum 135 (9) - villosus, see R. flagellaris 178 (12)
- ussuriense, see also R. nigrum 33, 135 (1, 9) - xanthocarpus, see R. idaeus 141 (9)
- uva-crispa, see R. grossularia 135 (9) RUMEX abyssinicus 126 (8)
RICINODENDRON heudelotii 111 (8) - acetosa 138 (9)
RICINUS communis 111 (8) - alpestris, see R. scutatus 138 (9)
RIVINA humilis 156 (10) - alpinus 138 (9)
ROBINIA hispida 177 (12) - ambiguus, see R. acetosa 138 (9)
- pseudoacacia 177 (12) - hymenosepalus 170 (11)
ROEGNERIA canina, see Agropyron caninum - obtusifolius 138 (9)
132 (9) - patientia 138 (9)
ROLLINIA deliclosa 145 (10) - scutatus 138 (9)
- dolabripetala, see R. longifolia 145 (10) - vesicarius 69 (4)
RUTA chalepensis 105 (7) SATUREJA pachyphylla, see S. hortensis 100 (7)
- graveolens 105 (7) SAUROPUS albicans 46 (2)
RYNCHELYTRUM repens, see Trichlaena rosea - androgynus, see S. albicans 46 (2)
117 (8) SCHINUS molle 145, 163 (10, 11)
SCHIZOSTACHYM brachycladus 48 (2)
SACCHARUM barberi, see S. sinense 66 (4) - grande 48 (2)
- edule 48 (2) - Iulampao 48 (2)
- officinarum, see also S. edule, S. sinense - zollingeri 48 (2)
S. pontaneum 48, 66 (2, 4) SCIRPODENDRON costatum, see S. ghaeri 45 (2)
- pedicellare, see S. officinarum 48 (2) - ghaeri 45 (2)
- robustum, see S. edule, S. officinarum 48 (2) SCIRPUS lacustris 132 (9)
- sinense, see also S. spontaneum, Miscanthus - validus, see S. lacustris 132 (9)
sinensis 32, 66 (1, 4) SCOLYMUS hispanicus 93 (7)
- spontaneum, see also S. officinarum, S. sinen- SCOPOLIA carniolica 142 (9)
se 48, 66, 116 (2, 4, 8) SCORZONERA hispanica 93 (7)
SAGITTARIA sagittifolia 27 (1) SECALE afghanicum, see also S. céréale, S.
SAKERSIA laurentia 124 (8) vavilovii 72, 82 (5, 6)
SALACCA edulis 54 (2) - africanum, see also S. montanum 82, 116
SALIX acutifolia 141 (9) (6, 8)
- alba 75, 141 (5, 9) - anatolicum, see also S. montanum 82 (6)
- amygdalina, see S. triandra 142 (9) - ancestrale, see S. cereale 82 (6)
- aurea, see S. alba 75, 141 (5, 9) - cereale, see also S. montanum, S. vavilovii,
- caprea, see also S. viminalis 141, 142 (9) Triticum aestivum 33, 72, 81, 82 (1, 5, 6)
- cinerea, see S. viminalis 142 (9) - ciliatoglume, see S. montanum 82 (6)
- cordata, see S. rigida 179 (12) - dalmaticum, see S. montanum 82 (6)
- dasyclados, see S. viminalis 142 (9) - daralgesii, see S. cereale, S. montanum 82 (6)
- fragilis, see also S. x rubens 141 (9) - dighoricum, see S. cereale 82 (6)
- longifolia, see S. viminalis 142 (9) - fragile, see S. silvestre 82 (6)
- x mollissima, see S. triandra 142 (9) - kuprijanovil, see S. montaneum 82 (6)
- purpurea, see also S. viminalis 142 (9) - montanum, see also S. africanum, S. anatoli-
- rigida 179 (12) cum, S. cereale, S. silvestre, S. vavilovii
- x rubens, see S. fragilis 141 (9) 82, 116 (6, 8)
- triandra, see also S. viminalis 142 (9) - segetale, see S. cereale 82 (6)
- viminalis 142 (9) - silvestre, see also S. montanum, and S. vavi-
SALSOLA komarovi 29 (1) lovii 82 (6)
- soda 29 (1) - turkestanicum, see also S. cereale 72, 82 (5, 6)
SALVIA chia 167 (11) - vavilovii, see also S. cereale, S. silvestre,
- divinorum 167 (11) S. montanum 82 (6)
- officinalis, see also Coleus amboinicus 49, 100 SECHIUM edule 165 (11)
(2, 7) SEDUM reflexum 131 (9)
- sclarea 100 (7) SELENICEREUS grandiflorus 164 (11)
- viridis 100 (7) SELINUM monnieri, see Ligusticum monnieri
SAMBUCUS nigra 142 (9) 56 (2)
SANDORICUM koetjape 51 (2) SEMECARPUS anacardium 43 (2)
SANGUISORBA minor 141 (9) SEMPERVIVUM tectorum 131 (9)
- officinalis 141 (9) SENECIA biafrae 109 (8)
SANSEVERINIA guineensis, see also S. trifasciata - gabonicus 109 (8)
108 (8) SESAMUM alatum 126 (8)
- hyacinthoides 62 (4) - indicum 35, 69, 126 (1, 4, 8)
- longiflora 108 (8) - prostratum, see S. indidWm 126 (8)
- thyrsiflora 108 (8) - radiatum 126 (8)
- trifasciata 108 (8) SESBANIA aculeata 67 (4)
- zeylanica, see S. hyacinthoides 62 (4) - aegyptiaca 51, 67 (2, 4)
SANTALUM album 55 (2) - exaltata, see also S. macrocarpa 177 (12)
SAPINDUS mukorosso 39 (1) - grandiflora 51 (2)
- rarak 55 (2) - macrocarpa 177 (12)
- saponaria 156 (10) - sesban, see S. aegyptiaca 51 (2)
- trifoliatus 69 (4) - speciosa 67 (4)
SAPIUM jenmani 149 (10) SETARIA glauca 116 (8)
- sebiferum 31 (1) - italica 33 (1)
SAPONARIA officinalis 130 (9) - pallidifusca, see S. italica 33 (1)
SAROTHAMNUS scoparius 137 (9) - sphacelata 116 (8)
SATUREJA hortensis 100 (7) - viridis, see S. italica 33 (1)
- illyrica, see S. montana 100 (7) SHOREA stenocarpa 46 (2)
- laxiflora, see S. hortensis 100 (7) SICANA odorifera 147 (10)
- montana 100 (7) SIDA rhombifolia 68 (4)
- obovata, see S. montana 100 (7) SILYBUM marianum 93 (7)
SIMAROUBA glauca 171 (11) 158, 159, 160 (10)

SIMMONDSIA California 171 (11) SOLANUM lacianiatum 61 (3)
SINAPIS alba 95 (7) - leptophyes, see S. brevicaule 171 (11)
- juncea, see Brassica juncea - leptosepalum 172 (11)
SINOBAMBUSA tootsik 33 (1) - lesteri 172 (11)
SINOCALAMUS beecheyanus 33 (1) - liriunianum, see S. brevicaule 171 (11)
- edulis 33 (1) - lycopersicoides, see S. pennellii 159 (10)
- giganteus 66 (4) - macrocarpon 127 (8)
- latiflorus 48 (2) - megistacrolobum, see S. raphanifolium 159 (10)
- oldhamii 33 (1) - melanocerasum, see S. burbankii 127 (8)
SIRIUM myrtifolium, see Santalum album 55 (2) - melongena 39, 70 (1, 4)
SITANION hystrix, see Agropyron spicatum 175 - michoacanum 172 (11)
(12) - microdontum, see S. chacoense 159 (10)
SIUM sisarum 106 (7) - morelliforme 172 (11)
SMYRNIUM olusatrum 106 (7) - multidissectum, see S. brevicaule 171 (11)
SOJA hispida, see Glycine max 34 (1) - multi-interruptum, see S. brevicaule, S. gonio-
- max, see Glycine max 34 (1) calyx 159, 171 (10, 11)
SOLANUM abancayense 158 (10) - muricatum 159 (10)
- abbottianum, see S. brevicaule 171 (11) - nelsoni, see S. rotundifolium 127 (8)
- acaule, see also S. x juzepczukii 158, 159 (10) - nigrum 127 (8)
- aculeastrum 127 (8) - nodiflorum 127 (8)
- aethiopicum 127 (8) - ochoae, see S. brevicaule 171 (11)
- agrimonifolium 171 (11) - olivare 127 (8)
- ajanhuiri 159 (10) - oxycarpum 172 (11)
- alibile, see S. topiro 160 (10) - papita, see also S. fendleri 172, 179 (11, 12)
- andigena, see S. tuberosum 160 (10) - pennellii 159 (10)
- andreanum, see S. verrucosum 172 (11) - phureja, see also S. bulbocastanum, S. chaucha,
- anomalum 127 (8) S. x juzepczukii, S. sparsipilum, S. steno-
- brachistotrichum 171 (11) tomum 159, 160, 171, 172 (10, 11)
- brachycarpum, see S. iopetalum 172 (11) - pinnatisectum, see also S. x sambucinum 172
- brevicaule, see also S. sparsipilum, S. steno- (11)
tomum 159, 160, 171 (10, 11) - polyadenium 172 (11)
- bulbocastanum, see also S. polytrichon 171, 172 - polytrichon, see also S. bulbocastanum 171,
(11) 172 (11)
- burbankii 127 (8) - quitoense 159 (10)
- canasense, see S. brevicaule, S. raphanifolium, - raphanifolium, see also S. brevicaule, S. s p a r -
S. sparsipilum 159, 171 (10, 11) sipilum 159, 171 (10, 11)
- cardiophyllum, see also S. x sambucinum 171, - rotundifolium 127 (8)
172 (11) - sarachoides, see S. burbankii 127 (8)
- caripense, see S. muricatum 159 (10) - x sambucinum, see also S. cardiophyllum, S.
- chacoense 159 (10) pinnatisectum 171, 172 (11)
- x chauca, see also S. stenotomum, S. tubero- - x semidemissum, see also S. demissum 171,
sum 159, 160 (10) 172 (11)
- d a r u m 171 (11) - soukupii, see also S. brevicaule 171 (11)
- commersonii 159 (10) - sparsipilum 159 (10)
- contumazaense 159 (10) - spegazzinii, see S. brevicaule 171 (11)
- curtilobum 159 (10) - stenotomum, see also S. ajanhuiri, S. b r e v i -
- demissum, see also S. x edinense, S. x s e m i - caule, S. chaucha, S. goniocalyx, S. x
demissum 171, 172 (11) juzepczukii, S. phureja, S. sparsipilum,
- duplosinuatum 127 (8) S. tuberosum 159, 160, 171 (10, 11)
- x edinense, see also S. demissum, S. x s e m i - - stoloniferum 172 (11)
demissum 171, 172 (11) - tabanoense, see S. muricatum 159 (10)
- fendleri, see also S. hjertingii, S. jamesii, - topiro 160 (10)
S. papita 172, 179 (11, 12) - trifida, see S. michoacanum 172 (11)
- georgicum, see S. topiro 160 (10) - tuberosum, see also S. bulbocastanum, S. chau-
- goniocalyx 159 (10) cha, S. x edinense, S. x juzepczukii, S. phu-
- guerreroense 171 (11) reja, S. sparsipilum, S. stenotomum 142,
- guineense, see S. burbankii 127 (8) 159, 160, 171 (9, 10, 11)
- hjertingii, see S. jamesii 172, 179 (11, 12) - uporo 56 (2)
- hougasii 172 (11) - x vallis-mexici, see also S. stoloniferum 172
- humectophilum 159 (10) (11)
- incanum 127 (8) - verrucosum, see also S. demissum, S. houga-
- iopetalum 172 (11) sii, S. iopetalum, S. stoloniferum, S. x
- jamesii 179 (12) vallis-mexici 171, 172 (11)
- juglandifolium 172 (11) - vidaurrei, see S. brevicaule 171 (11)
- x juzepczukii, see also S. acaule, S. ajanhuiri, - villosum, see S. burbankii 127 (8)
S. curtilobum, S. stenotomum, S. tuberosum - woodsonii 172 (11)
SOLEIROLIA soleirolii 106 (7) SYZYGIUM samarangense, see Eugenia javanica

SOLÈNOSTEMON ocymoides 118 (8) 53 (2)
SONCHUS taraxacifolius, see Lactua taraxacifolia
109 (8) TABERNAEMONTANA coronaria, see Ervatamia
SOPHORA secundiflora 169 (11) coronaria 43 (2)
SORBUS aucuparia, see Mespilus germanica 89, - divaricata, see Ervatamia coronaria 43 (2)
141 (6, 9) TABERNANTHE iboga 108 (8)
- domestica 90, 141 (6, 9) TACCA involucrata, see T. pinnatifida 56 (2)
SORGHUM aethiopicum, see S. bicolor 112 (8) - leontopetaloides, see T. pinnatifida 56 (2)
- almum, see also S. halepense 99, 149 (7, 10) - pinnatifida 56 (2)
- arundinaceum, see S. bicolor 116 (8) TAETSIA fruticosa 51 (2)
- bicolor, see also S. almum, S. halepense, TAGETES erecta, see also T. patula 164 (11)
Saccharum spontaneum 33, 66, 99, 116, 149 - minuta 147 (10)
(1, 4, 7, 8, 10) - patula, see also T. erecta 164 (11)
- halepense, see also S. almum 99, 149 (7, 10) - tenuifolia, see T. patula 164 (11)
- propinquum, see S. bicolor 33 (1) TALINUM cuneifolium 126 (8)
- vertieilliflorum, see S. bicolor 116 (8) - portulacifolium 126 (8)
SPARTINA alterniflora, see S. townendsii 134 (9) - trianguläre 126, 156 (8, 10)
- junceum 102 (7) TAMARINDUS indica 120 (8)
- maritima, see S. townendsii 134 (9) TAMARIX gallica 75 (5)
- townendsii 134 (9) - articulata 128 (8)
SPERGULA arvensis 130 (9) TARAXACUM bicorne, see T. koksaghyz 72 (5)
- sativa, see S. arvensis 130 (9) - hybernum 131 (9)
SPERGULARIA arvensis, see Spergula arvensis - koksaghyz 72 (5)
130 (9) - officinalis 131 (9)
SPHENOSTYLIS schweinfurthii 120 (8) TELFAIRIA occidentalis 110 (8)
- stenocarpa 120 (8) TEPHROSIA Candida 51 (2)
SPILANTHES acmella, see S. oleracea 147 (10) - densiflora 120 (8)
- oleracea 147 (10) - purpurea 181 (?)
- paniculata 45 (2) - singapou 169 (11)
SPINACIA oleracea 71 (5) - toxicaria, see T. singapou 169 (11)
- tetrandra, see S. oleracea 71 (5) - vogelii 120 (8)
SPONDIAS cytherea 180 (?) TERMINALIA bellirica 44 (2)
- dulcis, see S. cytherea 180 (?) - catappa 44 (2)
- laosensis 43 (2) - chebula 45 (2)
- lutea, see S. mombim 145 (10) TETRACARPIDUM conophorum, see Plukenetla
- mangifera, see S. pinnata 43 (2) conophora 111 (8)
- mombim, see also S. purpurea 145 (10) TETRAGONIA expansa 39 (1)
- pinnata 43 (2) TETRAPANAX papyriferum 28 (1)
- purpurea 145, 163 (10, 11) TEUCRIUM chamaedrys 100 (7)
STACHYS sieboldii 34 (1) - marum 100 (7)
STELECHOCARPA burahol 43(2) - officinale, see T. chamaedrys 100 (7)
STIPA tenacissima 99 (7) THEA lasiocalyx, see Camellia sinensis 39 (1)
- viridula 176 (12) - sinensis, see Camellia sinensis 39, 70 (1, 4)
STIZOLOBIUM aterrimum, Mucuna pruriens 50 THEOBROMA bicolor 160, 172 (10, 11)
(2) - cacao, see also T. bicolor 172 (11)
- deeringianum, see Mucuna deeringianum 50 (2) - grandiflora, see Guazuma grandiflora 160 (10)
- hassjoo, see Mucuna hassjoo 34 (1) - microcarpa 160 (10)
- niveum, see Mucuna cochinchinensis 50 (2) - pantegona 172 (11)
- pachylobium, see Mucuna pachylobia 67 (4) THESPESIA fissicalyx, see T. populnea 51 (2)
STRYCHNOS nux-vomica 70 (4) - multibracteata, see T. populnea 51 (2)
STYLOSANTHUS fruticosa 120 (8) - patellifera, see T. populnea 51 (2)
- guianensis 153 (10) - populnea 51 (2)
- humilis 120 (8) - robusta, see T. populnea 51 (2)
- mucronata, see S. fruticosa 120 (8) THEVETIA nereifolia 145 (10)
- surinamensis, see S. guianensis 153 (10) - peruviana, see T. nereifolia 145 (10)
STYRAX benzoin 56 (2) THYMUS vulgaris 101 (7)
SUOEDA glauca 29 (1) TORREYA grandis 39 (1)
SYMPHYTUM asperrinum, see S. asperum 78 (6) - nucifera 39 (1)
- asperum 78 (6) TOUCHARDIA latifolia 181 (?)
- officinale, see S. asperum 78 (6) TOURNEFORTIA argentea 44 (2)
SYZYGIUM aqueum, see Eugenia aquea 53 (2) TRACHYCARPUS fortunei 35 (1)
- aromaticum, see Eugenia caryophyllus 53 (2) TRACHYSPERMUM ammi, see Carum copticum
- cumini, see Eugenia jambolana 53 (2) 63 (4)
- jambos, see Eugenia jambos 53 (2) - roxburghianum, see Carum roxburghianum 45 (2)
- malaccensis, see Eugenia malaccensis 53 (2) TRAGOPOGON australis, see T. porrifolius 93 (7)
- mappaceum, see Eugenia formosa 53 (2) - porrifolius 93 (7)
- sativus, see T. porrifolius 93 (7) - sibiricum, see T. flavescens 134 (9)

TRAPA bicornis 40 (1) TRITICUM aegilipoides, see T. boeoticum 83 (6)
- bispinosa 40 (1) - aestivum, see also Aegilops c r a s s a , A. ovata,
- natans 40 (1) A. speltoides, T. monococcum, T. turgidum
TRICHOCEREUS pachanoi 146 (10) 33, 66, 72, 83, 117, 134 (1, 4, 5, 6, 8, 9)
TRICHOLAENA rosea 117 (8) - antiquorum, see T. aestivum 134 (9)
TRICHOSANTHES anguina, see also T. cucumeri- - araraticum, see T. timopheevi 83 (6)
na 45, 64 (2, 4) - armeniacum, see T. timopheevi 83 (6)
- cucumerina 64 (4) - bicorne, see Aegilops bicornis 96 (7)
- cucumeroides 30 (1) - boeoticum, see also Aegilops speltoides, T.
- japonica 30 (1) monococcum 83 (6)
- kadam, see Hodgsonia macrocarpa 30 (1) - carthlicum, see T. turgidum 84 (6)
TRIFOLIUM alexandrinum 102 (7) - cuadatum, see Aegilops caudata 79 (6)
- ambiguum 87 (6) - comosum, see Aegilops comosa 96 (7)
- anatolicum, see T. subterraneum 103 (7) - compactum, see T. aestivum 72, 82, 83, 134
- batmanicum, see T. subterraneum 103 (7) (5, 6, 9)
- berytheum, see T. alexandrinum 102 (7) - crassum, see Aegilops c r a s s a 79 (6)
- blesense, see T. subterraneum 103 (7) - cylindricum, see Aegilops cylindrica 79 (6)
- chlorotriehum, see T. subterraneum 103 (7) - dichasians, see Aegilops caudata 79 (6)
- expansum, see T. pratense 137 (9) - dicoccoides, see T. timopheevi, T. turgidum
- fistulosum, see T. hybridum 137 (9) 83 (6)
- fragiferum 102 (7) - dicoccum, see T. turgidum 84 (6)
- globosum, see T. subterraneum 103 (7) - durum, see T. turgidum 84 (6)
- hybridum 137 (9) - georgicum, see T. turgidum 84 (6)
- incarnatum 102 (7) - ispahanicum, see T. turgidum 84 (6)
- israeliticum 102 (7) - kotschyi, see Aegilops kotschyi 79 (6)
- neglectum, see T. fragiferum 102 (7) - lorentii, see Aegilops lorentii 79 (6)
- nidificum, see T. subterraneum 103 (7) - macha, see T. aestivum 83 (6)
- nigrescens, see T. repens 137 (9) - macrochaetum, see Aegilops lorentii 79 (6)
- occidentale, see T. repens 137 (9) - monococcum, see also T. speltoides, T. t u r g i -
- pannonicum 137 (9) dum, T. zhukovskyi 83 (6)
- pratense 137 (9) - orientale, see T. turgidum 84 (6)
- radiosum, see T. subterraneum 103 (7) - ovatum, see Aegilops ovata 80 (6)
- repens, see also T. hybridum 102, 137 (7, 9) - paleocolchicum, see T. turgidum 84 (6)
- resupinatum 138 (9) - peregrinum, see Aegilops variabilis 96 (7)
- salmoneum, see T. alexandrinum 102 (7) - persicum, see T. turgidum 84 (6)
- sativum, see T. pratense 137 (9) - polonicum, see T. turgidum 84 (6)
- savianum, see T. repens 137 (9) - rodeti, see Aegilops ventricosa 96 (7)
- suaveolens, see T. resupinatum 138 (9) - x sharonense, see Aegilops speltoides 80 (6)
- subterraneum, see also T. israeliticum 58, - spelta, see T. aestivum 82, 117, 134 (6, 8, 9)
102, 103 (3, 7) - speltoides, see Aegilops speltoides 80 (6)
- uniflorum, see T. repens 137 (9) - sphaerococcum, see T. aestivum 66, 83, 117
- vavilovii 103 (7) (4, 6, 8)
TRIGIDIA pavonia 167 (11) - tauschii, see Aegilops squarrosa 72 (5)
TRIGONELLA besserana, see T. coerulea 138 (9) - thaoudar, see T. monococcum 83 (6)
- coerulea 138 (9) - timopheevi, see also Aegilops speltoides, T.
- foenum-graecum 87 (6) monococcum, T. zhukovskyi 83 (6)
- procumbens, see T. coerulea 138 (9) - triaristatum, see Aegilops t r i a r i s t a t a 80 (6)
TRIGONOPLEURA malayana 46 (2) - tripsacum, see Aegilops mutica 80 (6)
TRIPHASIA aurantiola, see T. trifolia 39 (1) - triuncialis, see Aegilops triuncialis 80 (6)
- trifolia 39 (1) - turanicum, see T. turgidum 84 (6)
- trifoliata, see T. trifolia 39 (1) - turcomanica, see Aegilops juvenalis 79 (6)
TRIPSACUM, see Zea mays 166 (11) - turgidum, see also Aegilops ovata, A. ventri-
- australe 150 (10) cosa, T. aestivum, T. monococcum, T.
- dactyloides, see also Zea mays 150, 165, 166, timopheevi 72, 83, 96, 98, 117 (5, 6, 7, 8)
176 (10, 11, 12) - umbellulatum, see Aegilops umbellulata 80 (6)
- floridanum 176 (12) - uniaristatum, see Aegilops uniaristata 96 (7)
- lanceolatum 165 (11) - urarta, see T. monococcum 83 (6)
- latifolium 165 (11) - vavilovii, see T. aestivum 82 (6)
- laxum 165 (11) - ventricosum, see Aegilops ventricosa 96 (7)
- lemmoni, see T. lanceolatum 165 (11) - vulgare, see T. aestivum 33, 82, 117 (1, 5, 8)
- maizar .166 (11) - zhukovskyi 85 (6)
- pilosum, see also T. lanceolatum 165, 166 (11) TROPAEOLUM leptophyllum 161 (10)
- zopilotense 166 (11) - majus 161 (10)
TRIPTERYGIUM wilfordii 29 (1) - tuberosum, see also Oxalis tuberosa 161 (10)
TRISETUM flavescens 134 (9) TYPHA latifolia 40 (1)
- pratense, see T. flavescens 134 (9)
ULEX europaeus 138 (9) VICIA ervilea 87, 103 (6, 7)

ULLUCUS tuberosus 146 (10) - faba, see also Lupinus mutabilis 73, 103, 152
ULMUS villosa 76 (5) (5, 7, 10)
- wallichiana 76 (5) - graminea 153 (10)
UNCARIA gambir, see Trigonopleura malayana - hirsuta 138 (9)
46 (2) - leavenworthii 177 (12)
UNIOLA paniculata 176 (12) - narbonensis, see also V. faba 73, 87, 103
URBANELLA procera, see Lucuma procera 157 (5, 6, 7)
(10) - pannonica 87, 138 (6, 9)
URENA lobata 181 (?) - pliniana, see V. faba 103 (7)
URGINEA maritima 103 (7) - unijuga 34 (1)
- Scilla, see U. maritima 103 (7) - sativa 87 (6)
URTICA cannabina 74 (5) - villosa 87 (6)
- heterophylla, see Girardinia heterophylla 70 (4) VIGNA catjang, see V. unguiculata 120 (8)
- puya, see Maoutia puya 70 (4) - cylindrlca, see V. unguiculata 120 (8)
UVARIA burahol, see Stelechocarpus burahol 43 - dekindtiana 67 (4)
(2) - hosei 51 (2)
- maraguënsis, see V. hosei 51 (2)
VACCINIUM alto-montanum, see V. corymbosum - mungo, see Phaseolus mungo 67 (4)
175 (12) - parkeri, see V. hosei 51 (2)
- angustifolium, see V. corymbosum 175 (12) - radiata, see Phaseolus aureus 50 (2)
- arkansanum, see V. ashei 175 (12) - sesquipedalis, see V. unguiculata 120 (8)
- ashei 175 (12) - sinensis, see V. unguiculata 67, 120 (4, 8)
- australe, see V. ashei, V. corymbosum 175 - umbellata, see Phaseolus calcaratus 50 (2)
(12) - unguiculata 67, 120 (4, 8)
- corymbosum 175 (12) VINCA rosea 108 (8)
- darrowi, see V. ashei 175 (12) VIOLA arvensis, see V. tricolor 143 (9)
- lamarckii, see V. corymbosum 175 (12) - odorata 106 (7)
- macrocarpon 175 (12) - officinalis, see V. odorata 106 (7)
- microcarpum, see V. macrocarpon 175 (12) - tricolor 143 (9)
- myrsinites, see V. ashei 175 (12) - verucunda 41 (1)
- myrtillus, see V. corymbosum 175 (12) VITELLARIA paradoxa, see Butyrospermum
- oxycoccos, see V. macrocarpon 175 (12) parkii 127 (8)
- simulatum, see V. corymbosum 175 (12) VITEX agnus-castus 106 (7)
VALERIANA collina 142 (9) - cienkowskii 128 (8)
- edulis 179 (12) VITIS amurensis, see also V. vinifera 41 (1)
- eriocarpa 106 (7) - berlandieri, see also V. riparia 179 (12)
- exaltata 143 (9) - cinerea 179 (12)
- locusta 143 (9) - cordifolia 179 (12)
- officinalis, see also V. exaltata, V. p r o c u m - - davidii 41 (1)
bens, V. sambucifolia 143 (9) - labrusca, see also V. vinifera 90, 179 (6, 12)
- olitoria, see V. locusta 143 (9) - riparia, see also V. berlandieri 179 (12)
- procumbens 143 (9) - rotundifolia 179 (12)
- sambucifolia 143 (9) - rupestris 179 (12)
VANGUERIA edulis, see V. madagascariensis - shiragai, see V. amurensis 41 (1)
127 (8) - thunbergii, see V. amurensis 41 (1)
- madagascariensis 127 (8) - vinifera, see also V. amurensis, V. berlan-
VANILLA fragrans 170 (11) dieri, V. cinerea, V. cordifolia, V. labrusca,
- grandiflora, see V. pompona 170 (11) V. riparia, V. rotundifolia, V. rupestris
- planifolia, see V. fragrans 170 (11) 41, 76, 90, 106, 179 (1, 5, 6, 7, 12)
- pompona 170 (11) - vulpina, see V. riparia 179 (12)
VERBASCUM thapsiforme 142 (9) VOANDZEIA subterranea 120 (8)
VERDCOURTIA lignosus, see Dipogon lignosus
119 (8) WASABI japonica, see Eutrema wasabi 30(1)
VERNONIA amygdalina 63 (4) - pungens, see Eutrema wasabi 30 (1)
- anthelmintica 45 (2) WISSADULA contracta 155 (10)
VERONICA anagallis 39 (1) - periplocifolia 155 (10)
VETIVERIA odorata, see V. zizanioides 48 (2) WISTERIA brachybotrys 34 (1)
- zizanioides 48, 66 (2, 4) WOLFFIA arrhiza 51 (2)
VICIA angustifolia, see V. faba, V. sativa 73, 87
(5, 6) XANTHIUM strumarium 29 (1)
- articulata 103 (7) XANTHOSOMA belophyllum 146 (10)
- atropurpurea, see V. benghalensis 103 (7) - brasiliense 146 (10)
- benghalensis 103 (7) - caracu 146 (10)
- calcarata 103 (7) - jacquinii 146 (10)
- cordata, see V. sativa 87 (6) - robustum 163 (11)
- cracca 138 (9) - sagittifolium 146 (10)
XANTHOSOMA mafaffa 146 (10)

- violaceum 146 (10)
XYLOPIA aethioplca 108 (8)
YUCCA elephantipes 162 (11)

- funifera, see Hesperoyucca funifera 162 (11)
ZANTHOXYLUM bungei, see Z. simulans 39 (1)

- nitidum, see Z. simulans 39 (1)
- piperitum 39 (1)
- simulans 39 (1)
- varians, see Evodla hortensis 181 (?)
ZEA mays, see also Tripsacum dactyloides 33,
48, 66, 85, 99, 117, 134, 150, 165, 166
(1, 2, 4, 6, 7, 8, 9, 10, 11, 12)
- mexicana, see also Z. mays 150, 166, 167
(10, 11)
ZINGIBER cassumunar 56 (2)
- mioga 41 (1)
- officinale 70 (4)
- striolabum, see Z. mioga 41 (1)
- zerumbet 56 (2)
ZIZANIA aquatica 176 (12)
- latifolia 33 (1)
ZIZIPHUS jujuba, see also Z. mauritiana 36 (1)
- lotus 105 (7)
- mauritiana 36 (1)
- sosoria, see Z. mauritiana 36 (1)
- vulgaris 36 (1)

Dictionary Cultivated Plants

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Dictionary Cultivated Plants

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Dictionary of cultivated plants and

their centres of diversity

A.C. Zeven and P.M. Zhukovsky

Centre for Agricultural Publishing and Documentation

O Centre for Agricultural Publishing and Documentation, Wageningen, 1975

Cover design: Pudoc, Wageningen

Printed in the Netherlands by Krips Repro, Meppel

THE ORIGIN OF AGRICULTURE

1. Previously acquired skills in other fields to start experiments.

1. Wild plants collected by man.

1. Domestication, no apparent influx of foreign genie materialI

1. spread to a greater diversity of environments and a wider geographical range,

Helianthus annuus: 1. oil, 2. silage, 3. ornamental, 4. bird's food, 5. ceremonies,

Number of items per family p e r region, per family and p e r centre.

Family 10 11 12 Un- Total

Family 6 7 10 11 12 Un- Total

Family 1 2 3 4 5 6 7 8 9 10 11 12 Un- Total

1. differences in growth during the vegetative period.

Gene c e n t r e s of cultivated plants of Darlington &Janaki Amma l (1945) based on Vavilov

1. Southwest Asia 7. China

West African cradle B. Nilo-Abyssinian cradle

1. Southwest Asian agricultural complex-developed by the Caucasoids

1. China 7. Mediterranean coastal and adjacent regions

Megacentres of cultivated plants of Zhukovsky (1968)

Centres and noncentres of agricultural beginnings of Harlan (1971)

A l . Near Eastern A2. African

Actinidiaceae ACTINIDIA POLYGAMA Miq. Silver vine. 2n=c. 58,

The latter is considered a hybrid of A, fistulosum* Araceae

Campanulaceae LACTUCA INDICA L. Indian lettuce. 2n=18. India,

Celastraceae PETASITES JAPONICUS F. Schmidt. Fuhi. 2n=87.

CHLORANTHUS SPICATUS (Thunb. ) Mak. (syn. CORYLUS HETEROPHYLLA Fischer. Siberian

DIOSCOREA OPPOSITA Thunb. (syn. D. batatas Euphorbiaceae

ALEURITES FORDII Hemsl. Tung oil t r e e . 2n=22.

ALEURITES MONTANA (Lour. )Wils. Tung oil

SAPIUM SEBIFERUM Roxb. Chinese tallow t r e e .

ECHINOCHLOA FRUMENTACEA (Roxb.) Link, PHYLLOSTACHYS HENONIS Mitf. 2n=48, 54. C.

PHYLLOSTACHYS PUBESCENS Mazel ex de L e - Collins originated in E. Asia. Cultivated in China,

Labiatae ria and adjacent USSR. Domestication may have

Leguminosae VICIA UNLJUGA A. Br. (syn. Orobus lathyroides

LILIUM LANCIFOLIUM Thunb. Oni-yuri. 2n=24. Moraceae

Plantaginaceae Indian jujube. 2n=48. Tropical Africa and Asia.

Ranunculaceae ARMENIACA MUME (Sieb. &Zucc. ) Sieb, ex

MALUS HALLIANA Koehne. 2n=34. Gene centre in

MALUS HUPEHENSIS (Pamp.)Rehd. (syn. Pyrus

MALUS MICROMALUS Makino. 2n=34. China and

MALUS PUMILA var. niedzwetskyana Dieck. 2n=34.

MALUS SIEBOLDII Rehd. (syn. Pyrus toringo

MALUS SPECTABILIS (Ait.) Borkh. Zamechatel-

PRUNUS subgen. cerasus P e r s . There a r e proba-

PRUNUS CANTABRIGIENSIS Stapf, (syn. P .

PYRUS BRETSCHNEIDERI Rehd. 2n=34. Hupei

PYRUS CALLERYANA Dene. 2n=34. China, Japan

PYRUS PHAEOCARPA Rehd. 2n=34. N. China.

PYRUS PYRIFOLIA (Burm.) Nakai (syn. P . s e r o -

PYRUS USSURIENSIS Maxim. Ussuri pear. 2n=34.

ROSA MULTIFLORA Thunb. 2n=14. E. Asia. Used

Prunus davidiana ROSA RUGOSA Thunb. 2n= . China and Japan.

but sour. It was already known in the time of Con- Sapindaceae

60). P r i m a r y centre: the mountains of China, Thymelaeaceae

ANGELICA POLYMORPHA Maxim. 2n=22. China.

CRYPTOTAENIA JAPONICA Hassk. Mitsube.

GLEHNIA LITORALIS F. Schmidt. 2n=22. N. and

PHELLOPTERUS LITTORALIS Benth. Hama-bôhû.