Plant genetic resources of Ethiopia
Plant genetic
resources of Ethiopia
Edited by
J.M.M. ENGELS
Plant Genetic Resources Centre/Ethiopia,
Addis Ababa, Ethiopia
J . G . HAWKES
School of Continuing Studies,
University of Birmingham, UK
MELAKU WOREDE
Plant Genetic Resources Centre/Ethiopia,
Addis Ababa, Ethiopia
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CAMBRIDGE UNIVERSITY PRESS
Cambridge
New York Port Chester
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Published by the Press Syndicate of the University of Cambridge
The Pitt Building, Trumpington Street, Cambridge CB2 1RP
40 West 20th Street, New York, NY 10011, USA
10 Stamford Road, Oakleigh, Melbourne 3166, Australia
© Cambridge University Press 1991
First published 1991
British Library cataloguing in publication data
Plant genetic resources of Ethiopia.
1. Ethiopia. Plants. Genetic engineering
I. Engels, J. M. M. II. Hawkes, J. G.,
III. Worede, Melaku
581.1'5'0963
Library of Congress cataloguing in publication data available
ISBN 0 521 38456 7 hardback
Transferred to digital printing 2002
Contents
Contributors
List of acronyms
Preface
Part I General introduction
1 An Ethiopian perspective on conservation and utilization of
plant genetic resources
Melaku Worede
Part II The Ethiopian centre of diversity
2 The Ethiopian gene centre and its genetic diversity
/. M. M. Engels & } . G. Hawkes
3 Crops with wild relatives found in Ethiopia
Sue B. Edwards
4 Diversity of the Ethiopian flora
Tewolde Berhan Gebre Egziabher
5 Forest genetic resources of Ethiopia
/. de Vletter
6 Plants as a primary source of drugs in the traditional health
practices of Ethiopia
Dawit Abebe & Estifanos Hagos
7 Traditional aromatic and perfume plants in central Ethiopia (a
botanical and ethno-historical survey)
E. Goettsch
8 Spice germplasm in Ethiopia
E. Goettsch
9 A diversity study in Ethiopian barley
J.M.M. Engels
10 Sorghum history in relation to Ethiopia
H. Doggett
11 Prehistoric Ethiopia and India: contacts through sorghum and
millet genetic resources
K. L. Mehra
12 Konso agriculture and its plant genetic resources
/. M. M. Engels & E. Goettsch
Part III Germplasm collection and conservation in Ethiopia
13 Theory and practice of collecting germplasm in a centre of
diversity
/. G. Hawkes
vii-xiii
1
3
21
23
42
75
82
101
114
123
131
140
160
169
187
189
vi
Contents
14 A decade of germplasm exploration and collecting activities by
the Plant Genetic Resources Centre/Ethiopia
Abebe Demissie
15 Collection of Ethiopian forage germplasm at the International
Livestock Centre for Africa
Jean Hanson & Solomon Mengistu
16 Germplasm conservation at PGRC/E
Regassa Feyissa
17 Documentation at PGRC/E
Enyat Sendek & J. M. M. Engels
Part IV Evaluation and utilization of Ethiopian genetic resources
18 Germplasm evaluation with special reference to the role of
taxonomy in genebanks
/. G. Hawkes
19 Crop germplasm multiplication, characterization, evaluation
and utilization at PGRC/E
Hailu Mekbib
20 Evaluation methods and utilization of germplasm of annual
crop species
/. B. Smithson
21 Evaluation and utilization of Ethiopian forage species
/. R. Lazier & Alemayehu Mengistu
22 Improvement of indigenous durum wheat landraces in
Ethiopia
Tesfaye Tesemma
23 Use of germplasm resources in breeding wheat for disease
resistance
Hailu Gebre-Mariam
24 Indigenous barley germplasm in the Ethiopian breeding
programme
Hailu Gebre & Fekadu Alemayehu
25 The role of Ethiopian sorghum germplasm resources in the
national breeding programme
Yilma Kebede
26 Germplasm evaluation and breeding work on teff (Eragrostis
tef) in Ethiopia
Seyfu Ketema
27 Pulse crops of Ethiopia: genetic resources and their utilization
Hailu Mekbib, Abebe Demissie & Abebe Tullu
28 Oil crop germplasm: a vital resource for the plant breeder
Hiruy Belayneh
29 Significance of Ethiopian coffee genetic resources to coffee
improvement
Mesfin Ameha
30 Use of Ethiopian germplasm in national and international
programmes
/. G. Hawkes & Melaku Worede
Index
202
218
226
235
245
247
258
268
278
288
296
303
315
323
329
344
354
360
369
Contributors
ABEBE, Dawit. c/o Coordinating Office for Traditional Medicine, PO
Box 5117, Addis Ababa, Ethiopia.
ALEMAYEHU, Fekadu. Institute of Agricultural Research, Holetta
Research Centre, PO Box 2003, Addis Ababa, Ethiopia.
AMEHA, Mesfin. Institute of Agricultural Research, Jima Research
Centre, PO Box 2003, Addis Ababa, Ethiopia.
BELAYNEH, Hiruy. Institute of Agricultural Research, Holetta
Research Centre, PO Box 2003, Addis Ababa, Ethiopia.
DEMISSIE, Abebe. Plant Genetic Resources Centre/Ethiopia, PO
Box 30726, Addis Ababa, Ethiopia,
de VLETTER, J. German Agency for Technical Cooperation (GTZ),
PO Box 60054, Addis Ababa, Ethiopia.
DOGGETT, H. 38a Cottenham Road, Histon, Cambridge CB4 4ES,
UK.
EDWARDS, Sue B. c/o Asmara University, PO Box 1220, Asmara,
Ethiopia.
ENGELS, Jan M. M. IBPGR Regional Coordinator for South and
Southeast Asia, c/o NBPGR, Pusa Campus, New Delhi
110012, India (permanent address: Worbesgarten 6a, 6239
Eppstein-Ehlhalten, Federal Republic of Germany).
FEYISSA, Regassa. Plant Genetic Resources Centre/Ethiopia, PO
Box 30726, Addis Ababa, Ethiopia.
GEBRE, Hailu. Institute of Agricultural Research, Holetta Research
Centre, PO Box 2003, Addis Ababa, Ethiopia.
GEBRE EGZIABHER, Tewolde Berhan. Asmara University, PO
Box 1220, Asmara, Ethiopia.
GEBRE-MARIAM, Hailu. Institute of Agricultural Research,
viii
Contributors
Holetta Research Centre, PO Box 2003, Addis Ababa,
Ethiopia.
GOETTSCH, E. Scharweg 72, Kiel, Federal Republic of Germany.
HAGOS, Estifanos. c/o Coordinating Office for Traditional Medicine, PO Box 5117, Addis Ababa, Ethiopia.
H A N S O N , Jean. International Livestock Centre for Africa, PO Box
5689, Addis Ababa, Ethiopia.
HAWKES, John G. University of Birmingham, c/o School of Continuing Studies, PO Box 363, Birmingham B152TT, UK.
KEBEDE, Yilma. Institute of Agricultural Research, Holetta
Research Centre, PO Box 2003, Addis Ababa, Ethiopia.
KETEM A, Seyfu. Debre Zeit Agricultural Research Centre/AUA, PO
Box 32, Debre Zeit, Ethiopia.
LAZIER, John R. 187 King Street, West Cobourg, Ontario, Canada
KgA 2M8.
MEHR A, K. L. c/o National Bureau of Plant Genetic Resources, New
Delhi 110012, India.
MEKBIB, Hailu. Plant Genetic Resources Centre/Ethiopia. PO Box
30726, Addis Ababa, Ethiopia.
MENGISTU, Alemayehu. c/o Dr John Lazier, PO Box 5689, Addis
Ababa, Ethiopia.
MENGISTU, Solomon. International Livestock Centre for Africa,
PO Box 5689, Addis Ababa, Ethiopia.
SENDEK, Enyat. Plant Genetic Resources Centre/Ethiopia, PO Box
30726, Addis Ababa, Ethiopia.
SMITHSON, John B. Centro Internacional de Agricultura Tropical,
Apartado Aereo 6713, Cali, Colombia.
TESEMMA,
Tesfaye. Debre Zeit Agricultural Research
Centre/AUA, PO Box 32, Debre Zeit, Ethiopia.
TULLU, Abebe. Debre Zeit Agricultural Research Centre/AUA, PO
Box 32, Debre Zeit, Ethiopia.
WOREDE, Melaku. Plant Genetic Resources Centre/Ethiopia, PO
Box 30726, Addis Ababa, Ethiopia.
List of acronyms used in this volume
AAASA
ADD
AVRDC
CADU
CATIE
CEPGL
CGIAR
CIAT
CIBC
CIMMYT
CIP
CSIRO
ESC
FAL
FAO
FNE
Association for the Advancement of Agricultural
Sciences in Africa
Agricultural Development Department of MOA
(Ethiopia)
Asian Vegetable Research and Development Center
Chilalo Agricultural Development Unit
Centro Agronomico Tropical de Investigation y
Ensenanza (Costa Rica)
Communaute Economique des Pays des Grands Lacs
Consultative Group on International Agricultural
Research
Centro Internacional de Agricultura Tropical
(International Center of Tropical Agriculture)
(Colombia)
Commonwealth Institute for Biological Control
Centro Internacional de Mejoramiento de Maiz y Trigo
(International Centre for Maize and Wheat
Improvement) (Mexico)
Centro Internacional de la Papa (International Potato
Centre) (Peru)
Commonwealth Scientific and Industrial Research
Organisation (Australia)
Ethiopian Seed Corporation
Forschungsanstalt fur Landwirtschaft (Federal Republic
of Germany)
Food and Agricultural Organization of the United
Nations
Forage Network in Ethiopia
x
List of acronyms
GTZ
IAR
IARCs
IBPGR
ICAR
ICARDA
ICRISAT
IDRC
IITA
ILCA
ILRAD
IRAZ
IRGC
IRRI
IRTP
ISNAR
ISTA
IUCN
IUFRO
MOA
NAS
NBPGR
NIHORT
NOAA
NPGS
OECD
ORSTOM
OXFAM
PGRC/E
RRC
SADCC
SAREC
SIDA
Deutsche Gesellschaft fiir Technische Zusammenarbeit
(German Agency for Technical Cooperation)
Institute of Agricultural Research (Ethiopia)
International Agricultural Research Centres
International Board for Plant Genetic Resources
Indian Council of Agricultural Research
International Center for Agricultural Research in the
Dry Areas (Syria)
International Crops Research Institute for the Semi-Arid
Tropics (India)
International Development Research Center (Canada)
International Institute of Tropical Agriculture (Nigeria)
International Livestock Centre for Africa
International Laboratory for Research on Animal
Diseases (Kenya)
Institut de Recherche Agricole et Zootechnique
International Rice Germplasm Center
International Rice Research Institute (Philippines)
International Rice Testing Program
International Service for National Agricultural Research
International Seed Testing Association
International Union for the Conservation of Nature
International Union of Forestry Research Organisations
Ministry of Agriculture (Ethiopia)
National Academy of Sciences
National Bureau of Plant Genetic Resources (India)
National Horticultural Research Institute (Nigeria)
National Oceanographic and Atmospheric
Administration (USA)
National Plant Germplasm System (USA)
Organization for Economic Cooperation and
Development
Office de la Recherche Scientifique et Technique
d'Outre-mer (France)
Oxford Committee for Famine Relief
Plant Genetic Resources Centre/Ethiopia
Relief and Rehabilitation Commission (Ethiopia)
Southern African Development Coordination
Conference
Swedish Agency for Research Cooperation
Swedish International Development Authority
List of acronyms
UNDP
UNEP
UNESCO
United Nations Development Programme
United Nations Environment Programme
United Nations Educational, Scientific and Cultural
Organization
USDA
United States Department of Agriculture
USDA/GRIN United States Department of Agriculture/Genetic
Resources Information Network
WARDA
West African Rice Development Association
WWF
World Wide Fund for Nature (formerly the World
Wildlife Fund)
xi
Preface
Plant genetic resources constitute the building blocks of all
modern plant breeding. They form the raw material from which new
varieties have been systematically bred to meet the growing need for
more food. These traditional genepools are an invaluable asset to the
welfare of mankind and should be preserved, both for current use
and for posterity. Loss of genetic diversity is detrimental to crop
improvement programmes. To prevent this loss countries in all parts
of the world are endeavouring to conserve and utilize these precious
materials. The plant genetic resources (PGR), thus, must be
systematically collected, characterized, evaluated, documented and
conserved, for effective utilization. This is all the more important,
now, since agriculture is becoming more and more industrialized,
thus leading to a narrowing of the genetic base, demanding genetic
uniformity and causing vulnerability to pest and disease attacks, all of
which pose high risks to sustainable agricultural production systems.
The variability in landraces, and the primitive cultivars held by traditional farming societies, can provide the world with appropriate raw
material to stop such unwanted processes. It is therefore essential to
conserve it at all costs.
The urgency and the need to collect and conserve plant genetic
wealth was advocated some three decades back by the Food and
Agricultural Organization of the United Nations (FAO), and since the
1960s the network of activities in this context has spread considerably. Since the establishment of the International Board for Plant
Genetic Resources (IBPGR) in 1974 much has been done to create a
better awareness in plant genetic resources activities and to help
developing countries with training and equipment to this end. The
International Agricultural Research Centres have also been equally
xiv
Preface
instrumental in conserving and utilizing the genetic resources of
several major crops and offering the enhanced materials to Third
World national breeding programmes. The more recent concern of
the FAO Commission on Plant Genetic Resources is equally laudable
in this direction.
With the above international developments, national programmes
have also gradually been strengthened, for instance in Brazil, Ethiopia, India, Indonesia and the Philippines. The Plant Genetic
Resources Centre/Ethiopia (PGRC/E) is an excellent example of such a
functional national programme of a country which in this case is a
well-known centre of diversity and domestication for several world
crops.
The PGRC/E was established in 1976 and with the assistance of the
government of the Federal Republic of Germany has developed into a
full-fledged genebank. PGRC/E, in collaboration with the Deutsche
Gesellschaft fur Technische Zusammenarbeit (GTZ), organized the
First International Symposium on the Conservation and Utilization of
Ethiopian Germplasm in Addis Ababa, Ethiopia, from 13 to 16
October 1986. Sixty-one participants from Ethiopia and 29 from overseas took part in this symposium.
This book presents a synthesis of the activities on plant genetic
resources in Ethiopia and the richness of its plant wealth, laying
emphasis on economic plants of traditional use. The work is dealt
with in four sections, each of which highlights Ethiopia as a centre of
diversity of crop plants and their wild relatives, laying stress on their
collection and conservation and giving detailed accounts of activities
in the evaluation and utilization of these national assets. The international linkages of PGRC/E with International Agricultural Research
Centres and other regional and national programmes have been summarized in the last chapter.
It is felt that this book will add to the existing literature on plant
genetic resources and will be useful not only to scientists, but also to
teachers, policy makers and conservationists. It might in particular be
useful to other national programmes which are being developed at
present and which might require an easy reference source.
The editors are grateful to the contributors, to the secretarial staff
of PGRC/E, to Mrs Caryl Sheffield who was responsible for the copy
editing of the majority of the chapters and to the various organizations which have supported the Organizing Committee of the
symposium in various ways. Special mention should be made here of
the GTZ who financed the production of the proceedings of the
Preface
xv
symposium and of IBPGR for their contribution to the printing of the
cover. Undoubtedly, the book is an outcome of the fruitful cooperation and the sincere efforts of all those involved.
J. M. M. Engels
J. G. Hawkes
Melaku Worede
Parti
General introduction
An Ethiopian perspective on
conservation and utilization of plant
genetic resources
MELAKU WOREDE
Introduction
The Ethiopian region is characterized by a wide range of
agro-climatic conditions, which account for the enormous diversity of
biological resources that exist in the country. Probably the most
important of these resources is the immense genetic diversity of the
various crop plants grown in the country.
The indigenous landraces of the crop plant species, their wild
relatives and the wild and weedy species which form the basis of
Ethiopia's plant genetic resources, are highly prized for their potential value as sources of important variations for crop improvement
programmes.
Populations of these forms of plant species also represent sources
having the greatest potential for genetic diversity and can therefore
serve as invaluable means to fill the gaps that still exist in the available
base of genetic diversity in the world collection of many major crop
species. Among the most important traits which are believed to exist
in these materials are earliness, disease and pest resistance, nutritional quality, resistance to drought and other stress conditions, and
characteristics especially useful in low-input agriculture.
Scientists from many parts of the world have identified highly
desirable genetic characteristics in relatively few germplasm collections of various crop species and they are currently being utilized
intensively in a number of breeding programmes. Preservation of the
indigenous stock has a particular significance in the country's breeding programmes as characters of resistance and adaptation needed by
4
Melaku Worede
breeders to solve acute national problems exist in these materials.
Much of the existing diversity is in constant danger of being
irretrievably lost as the normally lower-yielding indigenous landraces
are being rapidly replaced by introduced or improved cultivars. Seeds
imported as food grain by relief agencies pose an even more serious
threat, as shown during the recent drought when farmers were
forced as a result of the food shortages to eat their own seeds in order
to survive or to sell them for consumption purposes. Entire
ecosystems are being demolished with advances in agriculture and
changes in land use.
Scientists have long realized the dangers of genetic erosion and
have stressed the need for timely action to salvage the country's still
abundant genetic resources. The accumulated efforts of these scientists and those of various concerned national and international
organizations led to the establishment, in 1976, of the Plant Genetic
Resources Centre (PGRC/E) in Ethiopia. The Centre has the following
major objectives:
- to promote the collection, evaluation, documentation and
scientific study of crop germplasm in Ethiopia, East Africa
and adjacent regions;
- to preserve germplasm by long-term storage and maintenance in order to make valuable germplasm available to
breeding programmes;
- to provide germplasm for breeding programmes aimed at the
development of such characters as higher yield, better
quality, and disease and pest resistance;
- to introduce new crop germplasm into Ethiopia by means of
exchange with other institutions.
The story pertaining to the establishment of PGRC/E and the role
played by the International Board for Plant Genetic Resources
(IBPGR) and the German Agency for Technical Cooperation (GTZ) in
attaining this goal are documented in a previous report (Worede,
1983a).
In this review an overall perspective of the various crop genetic
resources activities of the Centre is presented. Other aspects covered
in some detail include problems related to the Centre's germplasm
collections with particular reference to past and present situations as
well as utilization. PGRC/E efforts to coordinate ongoing and projected activities at national and international levels are also discussed.
Ethiopian perspective on conservation and utilization
5
Exploration and collection
In previous years, several scientific expeditions were made in
Ethiopia by a number of international (and national) explorers and
many species identified as being worthy of collection and preservation for their genetic diversity. Much of the work, however, was
confined to roadsides and less remote areas and in many instances
the collections were biased to meet specific needs. Organized missions to collect germplasm throughout Ethiopia started with the creation of the genebank in 1976. More than 115 successful collecting
missions have since been undertaken, involving nearly all
administrative regions of the country and covering a relatively broad
range of crop types and agro-ecological zones. In the field collecting
operation, priority has been given to those species of greatest economic and social importance which are most threatened by genetic
erosion. Areas chosen for collecting are those where the danger of
germplasm loss due to expansion of new varieties, natural disasters
and changes in land use is greatest.
Apart from the conventional germplasm explorations conducted
routinely by the Centre, systematic collecting of large samples of
indigenous landraces in drought-prone areas has been launched in
collaboration with the Ethiopian Seed Corporation (ESC). These samples are being stored at strategic seed reserve centres and subsequently redistributed to farmers as required. This is being done
primarily to avoid drastic losses of genetic variability in situations
where the farmer either is forced to eat his own seed or is replacing
traditional varieties with imported seeds distributed through relief
agencies.
Germplasm collecting in Ethiopia is, therefore, developing into a
major national and international effort responding to emergency situations and the need to build up comprehensive germplasm collections for sustained provision of such materials to breeding
programmes. As large numbers of samples accumulate, however,
questions of efficiency in the collecting and maintenance of the
material will inevitably arise.
To avoid such problems, measures for rational planning of future
collecting activities to explore areas of high genetic diversity are
already being undertaken. A case in point is the series of expeditions
(Seegeler, 1986) carried out recently to study genetic diversity of oil
plants in Ethiopia, which resulted in the identification of areas for
comprehensive collecting of such crops in the future. Such an undertaking is a long-term proposition, however, and will require a more
Melaku Worede
Table 1. Germplasm collecting at PGRC/E (June, 1986)
Crop type
Total number
of collected
and donated
accessions
Cereals
Oilseeds
Legumes
Spices
Coffee
Medicinal
Others
Total
28849
4490
4170
749
702
62
452
39474
%
73.1
11.4
10.6
1.9
1.7
0.1
1.2
100.0
Number of
accessions
collected
by PGRC/E
8219
2355
2890
520
140
61
397
14582
%
56.4
16.2
19.8
3.6
0.9
0.4
2.7
100.0
PGRC/E
collections as
percentage of
total number
of accessions
28
52
69
69
20
98
88
-
extensive survey of large areas representing a broad range of agroecological conditions and the collection of pertinent data over many
years.
Seed conservation
Indigenous landraces, mostly populations, form the bulk of
the existing germplasm collections currently maintained by PGRC/E
(Table 1). Such materials are maintained as active or base collections
at +4°C and -10°C, respectively, following established seed-processing procedures (Krauss, 1983). Seed viability is maintained through
regeneration of material in the field at ecologically appropriate
locations.
Field genebanks
The main focus of the living collection is on coffee. It is now
universally agreed that Ethiopia is the primary centre of diversification for Coffea arabica and perhaps the only region, covering the area
bordering southern Sudan and part of Uganda, where the species
occurs spontaneously. The genetic diversity that exists is tremendous
and this has great significance for the economy of the country and the
rest of the coffee growing world.
In realization of the urgent need for effective measures to preserve
and utilize the existing variability, which at present is being disastrously eroded, a special effort is being made to conserve coffee in
its natural growing environment. This includes conservation of the
semi-cultivated coffee in areas where the forest coffee occurs spon-
Ethiopian perspective on conservation and utilization
7
taneously, and where large variation exists, and maintenance of the
forest coffee in its natural ecosystem in certain protected areas, the socalled genetic reserves (Worede, 1982). A field genebank, comprising
some 700 accessions, is being established within the Kefa administrative region. In the future this genebank will be extended into other
appropriate sites as the size of the collection continues to increase.
Other living collections include Phytolacca spp., commonly known
as 'endod', Ensete ventricosum and several spices and root crops, maintained at different sites in the country in collaboration with existing
agricultural research and other relevant scientific institutions.
Situation with existing collections
In previous years, germplasm collections were maintained
mostly in small holdings by plant breeders who specialized in a few
major food and other crops. In such situations, many of the landraces
and wild types were probably excluded unless they exhibited characters of immediate breeding interest.
The genebank's present holdings (see Table 1) are an assemblage of
old collections that were acquired through transfer from several
breeding centres in Ethiopia, donations by various national and international organizations, and material collected in the field. The collections that were acquired through transfer from breeding centres and
other sources are, for the most part, deficient in documented records.
Many of these old collections are probably not representative of the
populations from which they were sampled and will have been subjected to subsequent natural and artificial selection. Poor storage conditions and improper maintenance of such material may also have
resulted in losses of seed viability.
Germplasm collections in Ethiopia are relatively secure at present
under the improved storage conditions at the Centre. Ethiopian
materials are represented in world collections, with numerous samples being exchanged among genebanks and breeders (Mengesha,
1975; Worede, 1983a). Duplication is inevitable but this serves as a
safeguard against losses and allows populations to be maintained and
studied under more diverse agro-ecological conditions. This may,
however, be of little value unless all passport and evaluation data are
made available by the various centres holding such collections.
Evaluation and documentation
Scientific studies of germplasm at PGRC/E focus primarily on
recording information which is essential in breeding activities. Such
studies include characterization and preliminary and further evalua-
8
Melaku Worede
tion of germplasm and these are carried out in collaboration with
breeders and other scientists. Appropriate descriptors, developed
jointly with the plant breeders, are utilized together with those provided by IBPGR whenever possible.
Further evaluation of material deals mainly with in-depth screening in the laboratory or greenhouse and in the field for disease/pest
resistance and adaptation to stress conditions. Screening under lowinput conditions is also included.
With landraces which are represented by highly heterogeneous
populations, maintenance or handling of accessions is often difficult
and is associated with a number of problems (e.g. genetic drift, contamination of material by foreign pollen) which occur during multiplication and evaluation. With the self-pollinating material, which is
composed of a large number of distinct genotypes, some compromise
is often needed to overcome the practical problems which such diversity poses to the evaluator. At PGRC/E, this is done by subdividing
samples into distinct agro-morphotypes to form the various components, which are then multiplied/characterized separately with
their own accession numbers. With the cross-pollinating types, subdividing is often not necessary as evaluation is dealt with at the
population level. Certain isolation techniques such as multiple bagging are, however, applied in developing the various components.
Data generated in the field and in the laboratory during evaluation
are collected and processed through the computer-assisted documentation system at the Centre for dissemination to breeders or other
users (Engels, 1983).
Genetic resources information available from PGRC/E may, in the
future, be incorporated into a regional or global network of information through the integration of pertinent data into an international
data bank (lead institutes, world directory, etc.).
Utilization of germplasm
The genebank in Ethiopia is designed primarily to provide a
sustained supply of germplasm material required by plant scientists
for the development of improved or new, superior crop varieties. Its
major activities are therefore geared to meet this requirement and are
aligned to follow each other in a logical sequence.
In many genebanks there are certain gaps in those activities linking
breeding with other related aspects which are the responsibility of the
germplasm user. This problem has been largely overcome in Ethiopia
as a result of integration of the Centre's utilization oriented activities
Ethiopian perspective on conservation and utilization
iliz
•o
CU
•S
CO
.2
3
ca
UJS
1
36343230282624222018161412108642-
1981
1982
1983
1984
Year
Fig. 1. Use of PGRC/E germplasm in national yield trials.
into the national breeding programmes. At the regional/international
level, a three-point-contact approach involving the national breeder
in Ethiopia, scientists from the collaborating institution(s) abroad and
PGRC/E has often been successful in conducting such activities to
mutual benefit.
In the past, cooperative efforts of this nature have often been
subject to serious constraints in the national programmes, mainly due
to the lack of evaluation data and insufficient interest among qualified
plant breeders to deal with the often low-yielding primitive landraces
(Worede, 1983b). However, the situation is now changing and as
germplasm data accumulate, breeders are becoming increasingly
interested in the utility of indigenous landraces. Many PGRC/E accessions are already incorporated into the national crop improvement
programmes through the national yield trials and are even utilized in
specific areas of breeding, including those related to resistance and
adaptation. Figure 1 shows the approximate progress of PGRC/E
germplasm utilized in national yield trials since 1981 (Anonymous,
1986).
Much progress in the effective use of the PGRC/E germplasm collection is also attributable to the excellent cooperative relationships
formed with the breeders and their active participation in the screening of germplasm material for specific characters of interest to them.
10
Melaku Worede
Thus breeders at Holetta Research Centre are already looking for
drought resistance in Brassica and Linum and are observing useful
features of earliness in some of the flax lines. The sorghum breeding
team in Ethiopia, together with scientists from the International
Crops Research Institute for the Semi-Arid Tropics (ICRISAT), have
identified lines that are promising for either stalk borer or leaf streak
resistance. Other important characteristics these breeders have
observed in PGRC/E sorghum collections include earliness, lodging
and disease resistance, yield and grain quality. In the indigenous
durum wheat, lines with remarkably high stem rust resistance as well
as good yield potential under low-input conditions have been identified by breeders at the Debre Zeit Research Centre of the National
Agricultural University. In the native castor bean collections, early
maturing as well as short-stemmed types have been identified and
resistance to rust is another outstanding feature.
The value to plant breeders of the existing PGRC/E collections of
indigenous germplasm largely resides in the adaptive complex that is
inherent in such stock, apart from such highly prized characters as
quality, nutritional value and disease/pest resistance. This must also
be viewed from the standpoint of the country's enormous wealth of
germplasm resources which have not yet been adequately explored.
Much of the diversity which exists among the primitive landraces,
their wild relatives or progenitors and the wild and weedy species, is
still untapped.
The potential value of these resources, apart from those with
characters of resistance and adaptation, is still difficult to assess. Such
materials often display crude forms of characters, some desirable and
some deleterious. With the development of new and advanced techniques like the new biotechnologies (Hansen et ah, 1986) it should be
possible to study and efficiently transfer only the useful genes which
would, in effect, increase the value of these primitive/wild gene
pools. It is imperative, however, that indigenous capabilities be
developed for safe and effective employment of such technologies in
scientific studies and utilization of germplasm. Pertinent policies
must also be formulated to reflect national and international interests.
Germplasm distribution!exchange
Samples of germplasm accessions are distributed to crop
improvement programmes in Ethiopia on the basis of specific
requests by breeders and other scientists actively utilizing such
material. Also, other projects receive germplasm for observation,
Ethiopian perspective on conservation and utilization
including adaptive trials, under specific types of farming conditions.
In some instances, excess material from multiplication/evaluation
activities, bulked on a crop-by-crop basis, has been sent to agricultural extension agents for study purposes, including some screening work, in the various localities.
The Centre also acquires material from external sources for storage
and/or distribution as required by the national breeding programmes.
This is usually through donations, including repatriation of germplasm of Ethiopian origin or exchange.
Ethiopian germplasm is actively utilized in current breeding work
worldwide, perhaps a good deal more than the country is given credit
for. Most of these materials were acquired through past explorations
but numerous samples representing a wide range of crop types have
been collected and utilized by regional projects like the International
Livestock Center for Africa (ILCA) (forage grasses and legumes), the
International Center for Agricultural Research in the Dry Areas
(ICARDA) (grain legumes, ICRISAT (sorghum/chickpea) and the
International Development Research Center (IDRC) Oil Crop Network for East Africa and the Indian Region (oilseeds), usually
through cooperative arrangements involving the appropriate national
projects in Ethiopia. PGRC/E has cooperative links with most of these
centres and material is exchanged through specific joint research
work involving national breeding programmes and other projects in
Ethiopia. A training component is often incorporated.
Among the major constraints on germplasm transfer to and from
the genebank are those related to seed health. In the absence of a
national quarantine system in Ethiopia, the Centre solicits the
assistance of the national crop protection programmes to check the
possible introduction of diseases and pests along with incoming
material. Seed dispatched abroad is certified by the appropriate
agency within the Ministry of Agriculture (MOA). The seed health
unit currently being established at PGRC/E is a further development
in the Centre's continuing effort to minimize any risk of introducing
diseases and pests, thereby facilitating a timely and more efficient
flow of germplasm material.
Cooperative links and perspectives
The cooperative role played by PGRC/E at national and international levels has already been mentioned in connection with germplasm utilization and exchange. It should be added that there are
cooperative links established bilaterally within Ethiopia with various
11
12
Melaku Worede
relevant institutions: ILCA (on forage germplasm resources and
exchange of indispensable services); the Pathobiology Institute of the
Addis Ababa University (Phytolacca spp. for medicinal and industrial
use); and the USSR Phytopathology Laboratory at Ambo on disease
resistance study. At the international level, collaborative ties exist
with IBPGR, the Food and Agriculture Organization of the United
Nations (FAO) and various national and international centres working in the same field, namely, ICRISAT, ICARDA, the International
Rice Research Institute (IRRI), the International Institute of Tropical
Agriculture (IITA), Gatersleben Genebank in the Democratic Republic
of Germany, Bari Germplasm Institute in Italy, the USSR Genetic
Resource System, and Forschungsanstalt fur Landwirtschaft (FAL) in
Braunschweig, Federal Republic of Germany, among others.
Genetic resources work on rice, currently being undertaken jointly
with IRRI, represents a new collaborative link at the international
level. It is directly related to a major PGRC/E objective, i.e. to
introduce new germplasm material to Ethiopia. Scientific studies of
wild rice found in the country, carried out by an Ethiopian scientist
trained at IRRI, are also included in such cooperation. This kind of
cooperative development is very significant for Ethiopia as it contributes towards the country's current effort to introduce rice as a
staple crop, especially in marginal areas where rice appears to be
adapted.
Genetic resources work, as Hawkes (1985a) puts it, consists of a
series of distinct activities or stages which follow each other in a
logical sequence. Each stage has an effect, good or bad, on the one
following it according to the efficiency with which it is carried out.
This means that for any genebank to be properly functional, all
aspects of genetic resources work must be adequately encompassed
in its programme and existing gaps filled.
The Ethiopian genebank has yet to claim a fully fledged status in
this regard although most of the activities pertaining to its major
objective are taking a definite shape. Many gaps still exist in the
strategies and scientific approaches currently employed by the Centre
to tackle the enormous qualitative and quantitative dimensions of
those conservation problems unique to Ethiopia. An opportunity
exists, however, to fill these gaps. New approaches and methodologies for scientific preservation can be developed on the basis of priority objectives through a well defined nationally or internationally
integrated network of activities. The Centre will continue to make full
use of such an opportunity and whenever possible seek to apply the
Ethiopian perspective on conservation and utilization
progressively advancing technologies that the international community is providing.
Gaps also exist in the available base of genetic diversity of the
various crops in the collections, as wild gene pools have hardly been
included in PGRC/E collecting operations to date. Nor have the distribution and degree of genetic diversity of crops in the different
ecological areas of the country been adequately explored to allow a
broad enough representation of material. Again based on priority
objectives, a relatively wide array of crop types and locations will
therefore be covered in future exploration/collecting missions. More
comprehensive collections will represent wider crop and plant categories, e.g. food/feed crops, medicinal plants, industrial crops and
other less known but potentially useful plant species. Every effort will
also be made to salvage germplasm material that is threatened with
extinction by the drought prevailing in the country and by factors
such as land clearing in forest areas and ploughing under pastures.
Surveys
Much work still remains to be done to define clearly the
situation of existing PGRC/E collections which comprise both old and
new material. Such a task is significant because it is a prerequisite to
any elaborate future exploration, since the loss of genetic diversity of
material already in storage must be taken into account.
Survey work, involving an assessment of materials for each crop
already in the genebank and for those likely to be still available in the
field, is therefore in order and a consultancy report for this complex
task is already at hand (Hawkes, 1985b).
In devising strategies and approaches for conservation work, one
should take into account emergency situations on the one hand and
the need to be selective or rational on the other. Given this situation,
it would be only logical for the Centre to capitalize on areas where
genetic diversity of a given crop species is concentrated, while
emergency operations in response to drought and changes in land
use continue. A more comprehensive approach to conservation must
also take into account the different requirements of the various plant
categories that the programme seeks to encompass.
Need for a diversified approach
Many new advances have been made in standardizing the
methodologies and approaches in the conservation of germplasm, the
most conventional being ex situ conservation, i.e. conservation as
13
14
Melaku Worede
seed, tissue or pollen or as plants in field genebanks. The Centre will
continue to work on reliable and low-cost, ex situ conservation techniques, including storage of seeds in permafrost areas as has been
suggested for developing countries (Swaminathan, 1983).
Technology for in vitro propagation and preservation of
vegetatively propagated materials is developing rapidly and PGRC/E
will look into the benefits to be gained in terms of plant health and
efficiency of maintenance and distribution of material, not to mention
the employment of such a technique in collecting material whose
seeds are difficult to conserve (Mix, 1983).
There are other, less conventional approaches to the future conservation of germplasm of various crops species, e.g. pollen storage.
Cryogenic (ultra-cold) storage, using liquid nitrogen at a temperature
of —196 °C is a further interesting advance and research is in progress
in the USA on the possible use of such technology in the preservation
of recalcitrant seeds and other plant propagules (National Seed
Storage Laboratory, Fort Collins, Colorado, personal communication). At present, little or no information is available regarding the
possible routine use of these technologies.
Neither of these technologies nor any single system would,
however, represent a mechanism by which safety of genetic diversity
is ensured on a long-term basis. It is, therefore, mandatory for a
country like Ethiopia with genetic reserves to resort to a more diversified conservation approach with a view to minimize losses of germplasm which are likely to occur where only preservation ex situ is
employed. New approaches and strategies, as they apply to the conservation problems unique to prevailing situations, should be considered and adopted by the country.
Conservation in situ, i.e. conservation of landraces and wild relatives in their natural habitats in areas where genetic diversity exists
and where wild/weedy forms are present, often hybridizing with
cultivated forms, represents a vital component of preserving diversity. Such an approach is essential because it relates to continuity of
the evolutionary systems that are responsible for generation of
variability.
As is beginning to happen in Ethiopia, wild relatives of crops could
be preserved in natural parks and biosphere reserves in a state of
continuing evolution through multi-institutional arrangements in
areas where large tracts of land of this nature still exist. Such a conservation approach may provide a less expensive protection of wild
gene pools than ex situ measures for developing countries which
Ethiopian perspective on conservation and utilization
15
often face practical constraints in providing optimal storage and
maintenance facilities for germplasm (Swaminathan, 1983). This type
of preservation also represents deliberate protection of remaining
habitats and the species they include, as entire preservation of vast
tracts with in situ conservation of animals and plants is extremely
important in slowing the rate of species extinction.
Role of farmers
In its broader sense, in situ conservation could include growing out of material without conscious selection on the site where the
seed was collected, although strictly speaking this is applied to
natural populations regenerated naturally (FAO, 1987).
This aspect of a diversified strategy of germplasm conservation
relates to a grass-roots approach involving farmers and community
workers. In Ethiopia as well as in many other developing countries,
farmers play a central role in the conservation of germplasm, as they
hold the bulk of genetic resources. Peasant farmers always retain
some seed stock for security unless circumstances dictate otherwise.
With certain horticultural crops grown by small farmers in Ethiopia, not only are crop species maintained in a dynamic state of evolution under conditions that are almost ideal in respect to sustaining
original population structures, but also new variations are created. In
Arsi region, for example, one often observes new, different forms of
fruits in Brassica carinata and B. nigra on farms where such crops are
grown in mixtures. This is attributable to introgression that may have
resulted from natural intercrossing between the two closely related
species, as presumably also reflected in the relatively high interpopulation diversity that is often observed among PGRC/E Brassica
collections.
With coffee, farmers often plant populations of local types on small
areas usually for safety purposes alongside the more uniform coffee
berry disease (CBD) resistant lines which are distributed by the Coffee Improvement Project in Ethiopia. This is a tremendous support in
the conservation of such a crop which in the first place is difficult to
store safely on a long-term basis as seed.
Similarly, as part of the national coffee conservation programme,
steps are being taken to conserve the semi-cultivated coffee on
peasant farms in areas like Kefa, Ilubabor and part of Welega where
the forest coffee occurred spontaneously or, in the case of backyard
coffee (Harerge, Sidamo and Welega regions), in nurseries maintained by farmer cooperatives.
16
Melaku Worede
Farmers in Ethiopia and other developing countries with similar
backgrounds should, therefore, be encouraged to continue to maintain small holdings of seed stock as this would represent some form
of in situ (on-site) conservation of germplasm across a broad range of
agro-ecological conditions. This may even be extended to encourage
the farmers or farmer cooperatives in Ethopia to play the curator/
farmer's role: grow limited samples of endangered landraces native
to the region (Mooney, 1983), exchange material within a network of
such activity, etc. This becomes even more significant in the long run
as the introduction of modern farming practices progresses in these
countries. Such cooperation by the farming community would provide an additional support to ex situ measures of landrace preservation, at least in providing a long-term protection against extinction of
native cultivars, and would complement in situ conservation in
natural parks or biosphere reserves.
Some allocation of funds should, however, be made through international support for a wider network of such activity to assist the
farmer as he undertakes the curator/farmer's role on account of the
cost-benefit implications that are associated with such an exercise.
Genebanks in cooperation with locally available agricultural extension agents could assist in the sampling of material and in providing
the technical support and monitoring necessary for systematic handling and scientific studies that should accompany such conservation.
Evaluation
Future scientific studies of germplasm will be rather more
comprehensive given the diverse possible uses of the country's wide
resources of plant material. Further evaluation work will be based on
such acute national problems as drought and other stress conditions,
as well as disease and pest resistance. Evaluation of germplasm under
low-input conditions, already initiated with Brassicas, will be
expanded to include various other crop types. New and appropriate
methodologies will aim to make the greatest possible use of the adaptive complex that exists within the indigenous stock.
Germplasm utilization/cooperative roles
The precise role of PGRC/E in international cooperation and
transfer of material will depend largely on the country's future
exchange policy. It is certain, however, that, without prejudice to the
principles of international cooperation, mutualism of benefits from
collaborative activities will continue to be the focal point. This can be
Ethiopian perspective on conservation and utilization
viewed in many different ways depending on the nature of the collaborative work and can be dealt with on a case-by-case basis once
pertinent policy guidelines are provided. In general, however, issues
that promote national capabilities in plant genetic resources, plant
breeding and seed multiplication in Ethiopia would, as has already
been happening, provide some basis for such cooperation. For
institutions working on common mandate crops, the sharing of
responsibilities and cost on the basis of common advantages provides
a useful basis for joint exploration activities, evaluation and subsequent utilization of material.
The three-point-contact arrangement mentioned earlier is one such
venture which has been successful. Already in some use at PGRC/E,
it has proved a useful mechanism in the transfer of technology and
sharing of costs that otherwise might have drawn substantially from
resources available for the national programme. The joint sorghum/
chickpea collecting and characterization work with ICRISAT scientists is one good example of cooperation of this nature (Prasada Rao &
Mengesha, 1981).
With regard to coffee, Ethiopia has already been identified as a
suitable location for a base collection for Africa on account of its
importance as a centre of genetic diversity of the crop. The task of
maintaining active collections should be shared among the other African countries concerned, as was decided at the First Regional Workshop on Coffee Berry Disease, held in Addis Ababa in July 1982.
Based on prevailing national policy, the Centre will pursue this line of
regional cooperation with a view to fostering concerted efforts to
salvage and effectively utilize the dwindling germplasm resources of
this important commodity crop.
Training
Training of Ethiopian scientists and suitable technicians is
another major objective which the Centre has pursued since its inception. As far as possible, training will constitute a vital component of
any ongoing and projected activities, especially in areas where collaboration with other institutions exists.
Training of personnel from other developing countries will also be
included and will continue to form another line of regional cooperation; indeed, PGRC/E is already playing a vital role in this field. An
even better prospect exists with the possibility of a genetic resources
training unit being established at the National Agricultural University
in Ethiopia.
17
18
Melaku Worede
External support/future role
Most of the activities described so far are resource demanding
and are too costly to be handled solely by a developing country like
Ethiopia. External support will therefore be needed, especially for
new areas or those that represent expansion of ongoing activities.
Based on pertinent national policy, such support will be sought
through collaborative links with regional/international institutions
working in the same field or others that are involved in the promotion
of genetic resources activities. The existing bilateral technical cooperative programme with the Federal Republic of Germany provides continued support on a follow-up basis.
It is true that PGRC/E is a national programme, mandated to operate primarily as a nucleus for the conservation of genetic resources in
Ethiopia. But by virtue of the significance of the country as a global
centre, it should, as is beginning to happen already, play a role as a
regional base for genetic conservation in Eastern Africa and
neighbouring regions.
References
Anonymous (1986). PGRC/E activity reports. Plant Genetic Resources
Centre, Addis Ababa (mimeographed).
Engels, J. M. M. (1983). Documentation and information management at
PGRC/E. PGRC/E-ILCA Germplasm Newsletter, 9, 20-7.
Food and Agriculture Organization (1987). Plant genetic resources, their con-
servation in situ for human use. Forest Resources Division, Forestry Department, FAO, Rome.
Hansen, M., Busch, L., Burkhardt, J., Lacy, W.B. & Lacy, L. (1986). Plant
breeding and biotechnology. Bioscience, 36, 29-39.
Hawkes, J. G. (1985a). Plant genetic resources - the impact of the international agricultural centres. CGIAR Study Paper no. 3. World Bank, Washington DC.
Hawkes, J. G. (1985b). Report on a consultancy mission to Ethiopia for GTZ
to advise PGRC/E on germplasm exploration, conservation, multiplication
and evaluation. Birmingham (mimeographed).
Krauss, A. (1983). Organization of the seed processing and storage section at
PGRC/E. PGRC/E-ILCA Germplasm Newsletter, 4, 9-12.
Mengesha, M. H. (1975). Crop germplasm diversity and resources in Ethiopia. In: O. H. Frankel and J. G. Hawkes (eds), Crop Genetic Resources for
Today and Tomorrow. Cambridge University Press, Cambridge, pp. 449-53.
Mix, G. (1983). The importance of in vitro techniques in germplasm conservation. In: K. J. Neddenriep and D. Wood (eds), Genetic Resources and the Plant
Breeder: the Next Ten Years. GTZ, Eschborn, pp. 77-88.
Mooney, P. R. (1983). The law of the seed: another development and plant
genetic resources. Development Dialogue, No. 1-2, pp. 6-173.
Prasada Rao, K. E. & Mengesha, M. H. (1981). A pointed collection of 'Zerazera' sorghum in the Gambela area of Ethiopia. Genetic resources progress
report 33, ICRISAT, Patancheru.
Ethiopian perspective on conservation and utilization
19
Seegeler, C. J. P. (1986). Genetic variability of oilcrops in Ethiopia. Consultancy report by GTZ for PGRC/E. Oosterbeek (mimeographed).
Swaminathan, M.S. (1983). Genetic conservation: microbes to man.
Presidential Address, 15th International Congress of Genetics, New Delhi,
12 December 1983. IRRI, Philippines.
Worede, M. (1982). Coffee genetic resources in Ethiopia: conservation and
utilization with particular reference to CBD resistance. Proceedings, First
Regional Workshop on Coffee Berry Disease. Association for the Advancement
of Agricultural Sciences in Africa, Addis Ababa, pp. 203-11.
Worede, M. (1983a). Crop genetic resources in Ethiopia. In: J. C. Holmes and
W. M. Tahir (eds), More Food from Better Technology. FAO, Rome, pp. 143-7.
Worede, M. (1983b). Issues relevant to genebank management problems in
relation to national and international programmes. In: K. J. Neddenriep
and D. Wood (eds), Genetic Resources and the Plant Breeder: the Next Ten
Years. GTZ, Eschborn, pp. 101-6.
Part II
The Ethiopian centre of diversity
The Ethiopian gene centre and its
genetic diversity
J. M. M. ENGELS AND J. G. HAWKES
Introduction
Based on the concept of gene centres, developed by N.I.
Vavilov in the 1920s, Ethiopia represents one of the eight centres in
the world where crop plant diversity is strikingly high and where
some of the crops concerned became domesticated. The concepts of
centres of origin and diversity have evolved considerably since
Vavilov's days as shown for instance by Harlan (1975) and by Hawkes
(1983). Nevertheless, some of the basic characteristics which apply to
the majority of the world's gene centres generally hold also for Ethiopia though varying from one crop to another. The highly dissected
highland of Ethiopia includes natural barriers formed by mountains
(up to approximately 5000 m above sea level) or ravines (sometimes
more than 1300 m deep) where crop plants would have evolved in
isolation under primitive agricultural conditions. To this must be
added the ancient and very diverse cultural history of its people, thus
providing many thousands of years of artificial selection within the
landrace populations since the early days of agriculture. However,
several crops which possess an extremely high diversity in Ethiopia
do not follow all the basic principles required for a centre of origin in a
given crop. In the case of barley, for instance, no wild relatives are
known within the country nor is there any archaeological evidence to
indicate early cultivation or domestication. In such cases the term
'secondary centre of diversity' has been used by Vavilov and would
be correctly applicable to various other Ethiopian crop plants with an
extremely high diversity (e.g. tetraploid wheats, lentil, faba bean and
others). Schiemann (1951) considered Ethiopia to be an 'accumulation
centre' for genetic diversity of certain crops which had not originated
24
/. M. M. Engels & J. G. Hawkes
there (see under barley, below). For many other crops Ethiopia is a
main centre of diversity as well as the probable area of domestication.
In the following paragraphs an attempt is made to provide information on the status of the Ethiopian centre for each crop in terms of
whether it is a primary or secondary centre of diversity, and whether
or not the crop in question is endemic. Relevant facts and figures are
presented on the important traits for the major crops as well as for
some of the minor crops that are unique to Ethiopia, in order to
illustrate the importance of the Ethiopian gene centre for plant
breeders throughout the world. As stated by Leppik (1970) it is a
generally accepted rule today, though with some exceptions, that the
primary and secondary gene centres of cultivated plants are the best
places to find genuine resistance to common diseases and insect
pests. Furthermore, some factors are mentioned which are causing an
alarming threat to genetic diversity and which give full support and
justification for the operation of a costly genebank within the Ethiopian gene centre.
Finally, the significance of Ethiopia as a source of important diversity in plants can be illustrated by the germplasm flow in and out of
the country since historical times. Not only did the Fertile Crescent
have contacts with the Ethiopian highlands in very early times but the
Egyptians, the Arabs and the Indians (Mehra, Chapter 11) exchanged
plant materials at later periods. From the early days of the European
voyages of discovery, European countries began to profit from the
plant genetic diversity of Ethiopia: first the Portuguese, then the
Italians, Germans, Russians and others.
Cereals
Barley (Hordeum vulgare)
Barley ranks only third in terms of acreage in Ethiopia after
teff and sorghum, but would seem to rank first or second in terms of
phenotypic diversity (Engels, Chapter 9). The crop does not possess
any related wild species in Ethiopia and this might support the idea
that the species was introduced from the Near East in ancient times,
possibly 5000-6000 years ago (Harlan, 1975; Purseglove, 1976). Since
then the crop has formed unique morphotype groups, such as the
deficiens and irregulare barleys. This extreme morphological diversity
led Schiemann (1951) to propose the concept of 'accumulation
centres' for barley in Ethiopia and hexaploid wheats in the Hindu
Kush. She argued that both those regions, where the original wild
prototypes did not exist, could be considered as areas where highly
The Ethiopian gene centre and its genetic diversity
25
diverse natural and artificial selection had been responsible for great
diversity; similar diversity might have occurred elsewhere in ancient
times but had been strongly selected against subsequently. Several
mutations unique to Ethiopia have also been found such as barley
yellow dwarf virus resistance, high lysine gene, resistance to diseases
such as powdery mildew, leaf rust, net blotch, Septoria, scald, spot
blotch, loose smut and barley stripe mosaic virus (Qualset, 1975).
Recently, considerable variation in the degree of drought resistance
has been found, related partly to differences in the root phenotypes
(Hettinger & Engels, 1986). Another example supporting the remarkable status of the Ethiopian barleys is given by Froest, Holm & Asker
(1975) who found a pattern C of flavonoids which was almost completely confined to Ethiopia.
At present, one can still find extensively grown landraces of barley,
some of them consisting of 10 or more clearly distinguishable components. Genetic erosion is caused mainly by replacement of barley
by crops such as bread wheat, teff and, recently, also oats (J.M.M.E.,
personal observation). This replacement is clearly shown by the
steady decrease in acreage since the early 1970s (from approximately
1.8 million hectares in 1971-2 to 0.75 million hectares in 1983-4).
Sorghum (Sorghum bicolor)
The Ethiopian sorghums may well be the most variable of all
crops grown in the country. This could be a reflection of the wide
variation in environments where the crop is being cultivated, ranging
from altitudes below 400 m in a few cases up to almost 3000 m above
sea level. The rainfall range over the whole of the country varies from
approximately 600 mm to well above 2000 mm in the south-western
part.
Of the five morphological sorghum races recognized, i.e. bicolor,
guinea, caudatum, durra and kafir, all except kafir are grown in
Ethiopia. In addition, many of the intermediate forms as well as
several of the wild and weedy forms (e.g. arundinaceum and aethiopicum), can be found (Snowdon, 1955; Stemler, Harlan & de Wet, 1975;
Doggett, Chapter 10; J.M.M.E., personal observation).
The diversity of sorghum encountered by Vavilov in the 1920s led
him to believe that Ethiopia was the centre of domestication of the
crop, and this hypothesis has been supported by many scientists
since then. However, in a detailed account Stemler et al. (1975)
recognized the overwhelming diversity in Ethiopia but presented
arguments that only the race durra might have originated in Ethiopia
26
/. M. M. Engels & J. G. Hawkes
Table 1. Phenotypic diversity expressed in the range, mean and cv of
Ethiopian sorghum germplasm accessions for some quantitative characters
Range
Character
Minimum
Maximum
Mean
cv
(%)
N
Plant height (cm)
Ear length (cm)
Ear width (cm)
Peduncle extension (cm)
Number of days to 50%
flowering
Crude protein content (%)
Thousand-grain weight (g)
19
4
2
1
475
50
30
44
233.7
21.5
9.4
14.7
22.7
36.6
34.1
53.8
2599
2511
2599
2254
76
5.0
6.0
169
15.3
61.1
116.7
9.6
28.5
14.3
13.1
35.5
2603
3644
200
Source: PGRC/E, unpublished data.
and that most probably the race durra-bicolor has further evolved
there subsequently.
Some of the phenotypic diversity for certain quantitative characters
is demonstrated in Table 1. Other important traits reported from
Ethiopian sorghum are: cold tolerance (Singh, 1985); high lysine and
protein content (Singh & Axtell, 1973), glossy seedlings - related to
sorghum shootfly resistance (Maiti et ah, 1984); grain quality and
resistance against grain mould (International Crops Research
Institute for the Semi-Arid Tropics, 1985). Other disease and pest
resistance has been reported from Ethiopia (Plant Genetic Resources
Centre/Ethiopia, unpublished data). Recently, Subramanian et al.
(1987) have reported high sugar content in the stalks of four Ethiopian
accessions of the ICRISAT collection. Drought resistance has been
observed in Ethiopia in several areas (J.M.M.E., personal
observation).
At present the diversity in the sorghum crop is doomed to be
reduced because of the adoption of improved local landraces as well
as imported varieties and the replacement of sorghum mainly by
maize (J.M.M.E., personal observation).
Wheat (Triticum spp.)
Vavilov (1931) was particularly impressed with the diversity
of Ethiopian wheats. He collected very widely there in 1926, and
recognized five species, though modern breeders prefer to class four
of them as one tetraploid species, T. turgidum. Vavilov's species were:
1. T. durum subsp. abyssinicum
The Ethiopian gene centre and its genetic diversity
27
2. T. turgidum subsp. abyssinicum
3. T. dicoccum
4. T. vulgare (now T. aestivum hexaploid)
5. T. polonicum
These 'species' can still be seen in farmers' fields today. Vavilov
was also impressed by the many endemic characters, such as violetgrained, beardless and half-bearded hard wheats (durum types), and
similar forms of turgidum wheats. He was also struck by the parallel
evolution of hard (durum) and soft (turgidum) forms, and considered
very strongly that the divergence of morphological and physiological
characters, clearly seen outside Ethiopia, had in this country not yet
emerged. He noted that the wild tetraploid ancestor, T. dicoccoides,
was not present in Ethiopia even though the Ethiopian wheats were
considered by him to be very primitive.
Vavilov pointed out that the most widespread species of wheat in
Ethiopia was T. durum whilst T. dicoccum was localized in the Harerge
and Addis Ababa regions.
Durum wheat is still one of the major cereals in Ethiopia. It ranks
fifth after teff, maize, sorghum and barley in acreage (Central Statistics Office, 1984; Tesemma, Chapter 22). In terms of genetic diversity it might compete well with barley and sorghum. Compared with
other major wheat producing countries, the Ethiopian durum wheat
accessions in the world germplasm collection, maintained by the
United States Department of Agriculture (USDA), showed the
highest diversity index (Jain et ah, 1975). One of the striking features
of the Ethiopian material was the high percentage of polymorphic
accessions, especially for glossy leaf sheath and kernel colour. A
similar high diversity index was reported by Negassa (1986) and by
Porceddu (1976).
Besides high phenotypic diversity, agronomically important genes
have also been found in Ethiopian germplasm. Resistance or
immunity to Erysiphe graminis f.sp. tritici, Puccinia spp. and Septoria
nodorum are found and Negassa (1986) reported dwarf genes in
several accessions. In other studies very early heading genotypes
have been reported (Qualset & Puri, 1974) as well as very late maturing ones (Porceddu, 1976).
Although Vavilov (1951) described the Ethiopian region as a centre
of diversity and origin for durum wheat, the absence of wild related
species, as well as archaeological findings, strongly suggests that
Ethiopia represents a secondary centre of diversity rather than a primary one. The alternative scheme of Hawkes (1983) distinguishing
28
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Nuclear Centres of agricultural origin (i.e. the Near East) and regions
of diversity (among others Ethiopia) fits the facts well in the case of
durum wheat and also of barley. Genetic erosion does occur, mainly
because of replacement of the crop by teff and because of imported
bread wheat varieties.
Besides durum wheat, emmer (T. turgidum subsp. dicoccoides) was
introduced into Ethiopia some 5000 years ago and is still fairly widespread (Haile/Mariam & Worede, 1988). T. turgidum conv. aethiopicum
is another convariety of subspecies turgidum which can be observed
regularly and about 11% of the wheat accessions in the Ethiopian
genebank belong to this form. Intermediate forms of durum and
aethiopicum are reported (Habtemariam & Worede, 1988). T. turgidum
subsp. turgidum conv. polonicum has been observed to exist in the
central highlands of Ethiopia, frequently interplanted with durum but
also occurring in pure stands. The diversity in the latter is relatively
small (J.M.M.E., personal observation).
Finally, it is worth pointing out that diploid einkorn wheats (T.
monococcum) and hexaploid bread wheats (T. aestivum, etc.) do not
seem to be native to the Ethiopian gene centre. The bread wheats
were in fact all introduced in recent historical times, but einkorn
wheats - as far as is known - never penetrated into the Ethiopian
region.
Teff (Eragrostis tef)
Teff is by far the most important crop in Ethiopia in terms of
acreage, thus, in 1983-4 about 1.38 million hectares were grown
(Central Statistics Office, 1984). It is cultivated from sea level up to
2800 m on all kinds of soil. However, the waterlogged soils of the
central Ethiopian highlands seem to be the 'cradle' of teff, a unique
environment with limited agricultural use. Although E. tef has a wide
distribution in Africa, it is cultivated as a food crop only in Ethiopia
and North and South Yemen (D. Wood & L. Guarino, personal communication). No detailed information is available as to when the
species was brought into cultivation, but it might be several
thousands of years. Shaw (1976) argues that teff must have been
domesticated before the introduction of wheat and barley to Ethiopia
'or else the teff, sorghum and finger millet never would have been
cultivated'. E. pilosa is believed to be the progenitor of the cereal
(Harlan, 1976) though, according to Lester & Bekele (1981), this species does not show any greater similarities to teff in its amino acid
composition than several other wild Eragrostis species. In a later study
The Ethiopian gene centre and its genetic diversity
29
(Bekele & Lester, 1981) in which data from leaf phenolic chromatography and seed protein electrophoresis were compared, it was found
that different teff cultivars showed similarities to several different
wild Eragrostis species, though E. pilosa seemed closest.
The phenotypic diversity in teff is clearly visible but less conspicuous compared with some of the other cereals. The height of the
plant, size and compactness of the culm and colour of the culm and
seeds are the most variable traits. Ebba (1975) described 35 landrace
varieties but undoubtedly more exist. Since diseases and storage
pests do not play an important role in teff cultivation, little research
has been done to find resistance (Ketema, 1986, Chapter 26).
The threat of genetic erosion to the diversity in teff is almost nonexistent. The crop is still expanding its acreage and almost no replacement of the landraces by improved varieties is taking place.
Miscellaneous cereal species
Finger millet (Eleusine coracana) has been mentioned
frequently in the literature as being very likely of Ethiopian origin
(Doggett, 1965; Harlan, 1969; de Wet et al, 1984). Purseglove (1976)
considered it to have originated in Uganda or a neighbouring
country. It is grown mainly in the western part of the country and
does not show such extreme diversity as some of the other cereals.
Seed size, straw length and flowering date do, however, vary considerably (J.M.M.E., personal observation). The wild species E. africana occurs as a weed in the finger millet fields and is regarded as the
progenitor of the cultivated species. The crop is facing some genetic
erosion, due mainly to replacement by other crops.
Pearl or bulrush millet (Pennisetum americanum) is another minor
millet which Doggett (1965) also believes to have originated in Ethiopia; others, however, such as Purseglove (1976) see the centre of
domestication in West Africa. Indeed, in Ethiopia it is not an important crop, since it is only grown in the north-western part of the
country in the marginal lower areas of Eritrea. At present the crop is
expanding quickly in these marginal environments, mainly at the
expense of sorghum and, to some extent, of finger millet (Teferie
Michael, personal communication).
Maize (Zea mays) is a relatively recent introduction into Ethiopia. It
is one of the fastest expanding crops and is causing a real threat to
some of the sorghum diversity (J.M.M.E., personal observation). In
the few hundred years since its introduction it has already built up
quite a remarkable diversity.
30
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In Ethiopia two wild weedy tetraploid oat species (Avena abyssinica
and A. vaviloviana) occur in the 'cereal belt' between 2200 and 2800 m
(Vavilov, 1957; Ladizinsky, 1975). Both are endemic to Ethiopia and
the Yemen, and are related to the Mediterranean A. barbata. Besides
being sometimes noxious weeds and adjusting themselves well to
changing conditions, the Ethiopian oats are of no commercial value.
Finally, a mention should be made of the wild rice species which
have been reported from the western lowlands. Both Oryza barthii and
O. longistaminata are collected from the wild by the local inhabitants in
times of food scarcity (Dadi & Engels, 1986).
Oil crops
Ethiopian mustard (Brassica carinata)
Ethiopian mustard is an important oil crop as well as a leaf
vegetable. It is extensively grown throughout the highlands and
shows a considerable diversity for several vegetative characters,
mainly regarding leaves and growth habit (Engels, 1984). This diversity may be due to the introgression of genes from B. oleracea, since
both species are often found growing in close proximity (Tcacenco,
Ford-Lloyd & Astley, 1985). During the evaluation of the Brassica
germplasm collection many intermediate forms between these two
species have been observed (J.M.M.E., personal observation). No
wild relative of Ethiopian mustard is known, a fact which further
supports the hypothesis that it is an (allotetraploid) hybrid between
B. nigra and B. oleracea. No clear indications for genetic erosion exist.
B. nigra is used mainly for medicinal purposes but also for greasing
the 'injera' pan. The species is predominantly grown in backyards but
can regularly be observed in the same field as B. carinata. B. oleracea is
cultivated mainly for its leaves and is perrenial. Tcacenco et al. (1985)
conducted chromosomal examinations and found that B. carinata and
B. oleracea make up a separate group of 'cabbage' types.
Niger seed, noog (Guizotia abyssinica)
This crop is treated as one of the 'classical Ethiopian crops' in
terms of its origin (Seegeler, 1983). The highlands of central Ethiopia
are the home of noog and because of the high demand for its oil the
crop is expanding its acreage yearly. Regarding its phenotypic diversity, the crop does not show such striking features as some of the
other crops, which might be partly due to its strong outbreeding
behaviour. The highest diversity exists for characters such as number
of days to flower and number of days to maturity as well as for head
The Ethiopian gene centre and its genetic diversity
size (J.M.M.E., personal observation). No systematic research on oil
content and oil quality has been conducted so far.
The frequent presence of the wild/weedy related species G. scabra
around the fields of noog indicates the possible presence of a crop/
weed complex which would be worth further study. Because of the
steadily expanding acreage of the crop and the absence of released
high-yielding varieties genetic erosion for noog will be mild, but
because of the further improvement of agricultural practices it could
be expected for the wild related species G. scabra.
Linseed (Linum usitatissimum)
Linseed is at present the second most important oil crop in
Ethiopia; only noog is more popular. Linseed is grown only for oil
production and its use as flax is hardly known. Vavilov (1957),
however, observed that the seeds were used mainly for the preparation of fodder meals, by grinding after drying. The phenotypic diversity is not very striking: some variation in flower colour, plant height,
number of days to flower and maturity as well as in capsule size has
been noted (J.M.M.E., personal observation). Despite this, Ethiopia
was regarded as a centre of flax diversity, though not of origin, by
Vavilov (1951). There are strong indications that the existing diversity
is seriously threatened by genetic erosion.
Sesame (Sesamum indicum)
The origin of sesame is still under dispute. However, since all
the wild species but one are native to Africa, Ethiopia would seem
very probably to be its centre of origin (Vavilov, 1951; Purseglove,
1968; Harlan, 1969; Nayar, 1976). It is certainly a very ancient cultigen
in Africa. At present sesame is the third most important oil crop in
economic terms in Ethiopia. The phenotypic diversity encountered in
the country is considerable for pod shape and size, for seed size and
colour (Seegeler, 1983) and for number of days to maturity and plant
habit (J.M.M.E., personal observation). The genetic erosion can be
considered as critical and more collecting remains to be done.
Castor bean (Ricinus communis)
Castor bean is widespread as a wild plant through East and
North Africa, the Yemen and the Middle East. According to Purseglove it was cultivated in Egypt from 4000 BC onwards. Although no
commercial production exists in Ethiopia, the plant is widely distributed almost from sea level up to the highlands, generally as a wild
31
32
/. M. M. Engels & J. G. Hawkes
plant, e.g. Eritrea (Bruecher, 1977), or as a weed in disturbed habitats.
'Weedy' plants with dehiscent pods and cultivated types with indehiscent pods can be observed (personal observations). The diversity
for many plant, fruit and seed characters is enormous and this would
fully justify the assumption of Vavilov (1951) and Zeven &
Zhukovsky (1975) that the cultivated castor bean might be of Ethiopian origin. Although no threat of genetic erosion exists at the
moment it would be an exciting study to collect and evaluate castor
bean more extensively. At present, the plant is used mainly for home
consumption, as a medicinal plant and as a producer of oil for
lighting.
Safflozver (Carthamus tinctorius)
Purseglove (1968) mentions three primary centres:
Afghanistan, the Nile Valley and Ethiopia. Since it was probably
derived from the wild C. oxycantha which occurs as a weed from India
to Turkey it is very difficult to be certain of its exact point of origin,
and indeed it may have had several. Vavilov (1951) and Knowles
(1976) considered Ethiopia to be its probable centre of domestication.
However, this assumption was not supported by a diversity study
conducted on a world germplasm collection where the Ethiopian
diversity index was fairly low (Wu & Jain, 1977). In addition, the
observed diversity in the field was relatively small (J.M.M.E., personal observation) and the crop is grown only on a small scale,
frequently in the borders around cereal fields. Since safflower can be
treated as a minor crop its diversity might well be endangered by
genetic erosion.
Crambe (Crambe abyssinica)
This 'new' crop has its home in the Ethiopian highlands (Leppik & White, 1975). Wild populations as well as 'cultivated' fields of
crambe can be found, but relatively rarely. Because of ever expanding
agricultural lands many natural habitats are disturbed and the existing diversity undergoes considerable genetic erosion.
Pulses
Faba bean (Vicia faba)
The faba bean is the most important pulse in Ethiopia, occurring mainly as the small-seeded type, a character also typical for
germplasm from Afghanistan, probably one of the primary centres of
diversification (Lafiandra et al., 1981). In the same study a high pro-
The Ethiopian gene centre and its genetic diversity
tein content in Ethiopian material was reported and, recently, considerable resistance against chocolate spot lias been found in
Ethiopian germplasm (PGRC/E, unpublished data). As in the other
pulses, at present no real threat to the diversity exists.
Field pea (Pisum sativum)
The field pea is an old crop in Ethiopia and is still one of the
dominant pulses in the country. During the several thousand years of
its presence, a unique subspecies developed in Ethiopia (subsp. abyssinicum) and this earlier led Vavilov to seek the origin of the pea partly
in Ethiopia. However, in a later publication (Vavilov, 1957) he admitted that this pea had probably come from the Yemen. From our own
observations it should be noted that the phenotypic diversity was
found to be rather limited. No information about the degree of
genetic erosion exists; however, it is expected to be low.
Chickpea (Cicer arietinum)
After faba beans and field peas, chickpeas are the most
important pulse in terms of acreage. They are eaten immature as a
snack or as matured seeds in a roasted, boiled or ground form. Chickpea is an ancient crop in Ethiopia and archaeological evidence in the
caves of Lalibela has shown an age of 500 BC (Dombrowski, 1969).
Although some authors have mentioned Ethiopia as a centre of origin
and a related wild species has been found in northern Ethiopia (e.g.
C. cuneatum), there are strong indications that the origin lies in southwest Asia (Harlan, 1969). Nevertheless, the diversity encountered in
the Ethiopian fields is considerable and Pundir et al. (1985) reported
unique flower colours, high anthocyanin in the leaves and non-beige
seed colours. Furthermore, some disease resistance and drought
tolerance has been found in initial testing. Genetic erosion is of little
importance and is no real threat to the chickpea diversity.
Lentil (Lens culinaris)
The lentil is one of the crops which was introduced into
Ethiopia in the early days from west Asia, probably via the Yemen,
and which belongs to the 'South-Western Asian Complex' (Harlan,
1969). Another study suggests that lentils might have originated in
India as a selection from a wild form (Williams, Sanchez & Jackson,
1974) but this will require further investigation. Because of the early
introduction a considerable diversity in the crop has built up. Erskine
& Witcombe (1984) reported the following interesting characteristics
33
34
/. M. M. Engels & J. G. Hawkes
found in Ethiopian germplasm: earliness, high seed yield, high
harvest index, high number of seeds per pod and good cold
tolerance. On the other hand, the Ethiopian material showed a low
seed protein content. Some genetic erosion can be expected since the
acreage of lentils is steadily declining.
Miscellaneous pulse species
Of the minor pulses, cowpea (Vigna unguiculata) should be
mentioned first. This African species may have been domesticated in
Ethiopia (Vavilov, 1951; Steele, 1976) and several wild (related) species exist in the country. Not much is known about the diversity in
this crop with its limited cultivation, mainly in Konso and the Gambela region (Engels & Dadi, 1986). The wild species might be
endangered because of drastic changes in land use.
Fenugreek (Trigonella foenum-graecum) is a crop with a long history
in Ethiopia. Although it is considered more as a medicinal plant it is
also used as a pulse. The genetic diversity is considerable (J.M.M.E.,
personal observation).
Grass pea (Lathyrus sativus) is a fairly common crop in the Ethiopian highlands, frequently grown on fallow lands. Its diversity is not
yet systematically studied but considerable drought resistance can be
expected (J.M.M.E., personal observation).
The hyacinth bean (Lablab purpureus) and pigeon pea (Cajanus cajan)
were reported by Vavilov (1951) and Zeven & Zhukovsky (1975) to be
of Ethiopian origin but there is considerable doubt about this. A more
likely hypothesis is that they were domesticated in India (Purseglove,
1968; Royes, 1976; Smith, 1976). Both crops are frequently cultivated
in southern Ethiopia (Konso) where they form an important component in the highly developed and sustainable agricultural system.
Miscellaneous crops of Ethiopian origin
Coffee (Coffea arabica)
From an economic point of view this may be the most important Ethiopian crop. Almost the entire diversity originated in Ethiopia,
mainly in the south-western rainforest area, where almost undisturbed patches of forest with coffee as undergrowth can still be
found. The phenotypic diversity is overwhelming for qualitative as
well as for quantitative characters (Tadesse & Engels, 1986), not to
mention the variation in disease and pest resistance (Wondimu,
1987), quality characteristics and others. Because of extensive deforestation and replacement of primitive coffee populations by maize, chat
The Ethiopian gene centre and its genetic diversity
35
and other crops, together with changing patterns in land use, the
diversity is highly threatened by erosion (Hawkes et al., 1986; Bellachew, 1987).
Ensete or false banana {Ensete ventricosum)
In the more humid highlands, especially in southern Ethiopia, this unique Ethiopian crop species is an important staple. The
starch of the pseudostem is used after fermentation treatment for
several days. Furthermore, the plant or parts of it are used as fodder,
for fuel and packing material, etc. (Olmstead, 1974). Although the
plant is propagated only vegetatively, an astonishing variation in
several characters can be found. Demeke et al. (1986) described 76
named varieties which vary mainly in colour of pseudostem and leaf
midribs, their earliness and quality of the final product as well as for
disease resistance, e.g. bacterial wilt. The most serious factors causing genetic erosion in ensete are bacterial wilt and, to a lesser extent,
drought.
Chat (Catha edulis)
This evergreen bush is the source of the fresh young leaves
which are chewed in Eastern Africa and the Arabian Peninsula for
their mild narcotic properties. The plant, which grows wild in East
Africa, was first domesticated in Ethiopia and is now expanding in
the eastern part of the country at the expense of coffee. Some striking
leaf colour variants have been observed, especially in south-east
Ethiopia (personal observation). No genetic erosion takes place at
present.
Okra (Abelmoschus esculentus)
Okra is grown only in the western lowlands. It was also in
this part of the country that, beside the domesticated species, two
more related species were encountered, one being 'half domesticated'
(A. manihot) and the other, A. moschatus) (?) being collected from the
wild (Engels & Dadi, 1986). These findings support the assumption of
Harlan (1969) and Vavilov (1951) that okra could be of Ethiopian
origin even though Joshi & Hardas (1976) cast some doubt on this.
Roots and tubers
Several species are cultivated or semi-cultivated in restricted
parts of Ethiopia but only for their roots or tubers. Especially in
southern Ethiopia (e.g. Konso), these crops play an important role.
36
/. M. M. Engels & J. G. Hawkes
The better known species are Plectranthus edulis (galla potato), Coccinia
abyssinica (anchote), Amorphophallus abyssinicus (bagana) and
Sauromatum nubicum (Dadi & Engels, 1982). Furthermore, several yam
species (Dioscorea spp.) might have their origin in Ethiopia as well
(Harlan, 1969; Zeven & Zhukovsky, 1975). Leon (1978) reports
Sphenostylis stenocarpa to be a 'yam bean' of possible Ethiopian origin.
Spices and medicinal plants
Several spices and medicinal plants are of Ethiopian origin, or
expected to be so. The most important species are Aframomum
korarima (false cardamom), Carum copticum (nech azmud), Coriandrutn
sativum (coriander), Embelia schimperi (enkoko), Hygenia abyssinica
(koso), Lepidium sativum (garden cress, feto), Nigella saliva (black
cumin) and Rhamnus prinoides (buckthorn, gesho). The bark of the
latter is used in place of hops in beer. Wilson & Gebre-Mariam (1979)
have published a list of 37 species which are used commonly as
medicinal plants on the Ethiopian plateau. Details of a wide variety of
medicinal plants are presented by Abebe & Hagos (Chapter 6).
Fibre plants
Nicholson (1960) presents arguments that cotton (Gossypium
herbaceum L. var. acerifolium) might have been domesticated in Ethiopia. Also kenaf (Hibiscus cannabinus) is reported to be of Ethiopian
origin (Purseglove, 1968; Zeven & Zhukovsky, 1975). In south-west
Ethiopia some wild Hibiscus species have been observed which were
reported to be used for their fibres (personal observation).
Moringa stenopetala (cabbage tree) is an important vegetable tree in
south Ethiopia (Konso). Some other trees are used extensively for
fodder (e.g. Balanites aegyptica and Terminalia brownii), a practice
unique to Ethiopia.
Conclusions
From the foregoing paragraphs it is obvious that Ethiopia is
an important centre of genetic diversity for a wide range of crops.
This is partly due to its very dissected terrain, with consequently an
extremely wide range of agro-ecological conditions. It is also due to its
geographical position at the crossroads between the Near-Eastern
centre of diversity on the one hand and the Indian centre on the
other. It has also benefited from its connections with the mountain
chain and Rift Valley, following southwards in East Africa, and its
connections with West Africa via the Sudan and Sahel regions.
The Ethiopian gene centre and its genetic diversity
37
Many of Ethiopia's most important crops such as barley and the
tetraploid wheats have come from elsewhere several thousand years
ago. Under Ethiopian conditions they have developed much diversity, often of a unique character. Hexaploid or bread wheats came
much later, perhaps not more than 100 years ago. Diploid wheat and
rye do not appear at all, and oats only as weeds.
Sorghum, with its tremendous diversity in Ethiopia, is something
of an enigma. Whilst several authorities such as Vavilov and Doggett
believe that Ethiopia may be the centre of origin of sorghum, others,
such as Harlan and de Wet, are rather doubtful.
However, there is no doubt about teff. This certainly was domesticated in Ethiopia and is still its most important cereal.
Concerning the minor millets (Eleusine coracana and Pennisetum
americanum), doubts exist as to their having originated in Ethiopia.
Of the oil crops, Guizotia abyssinica, Brassica carinata and Sesamum
indicum appear to be native, whilst others are of uncertain origin. Flax
was almost certainly introduced.
Pulses such as faba bean, field pea, chickpea and lentil seem all to
have been introduced, though they probably have been in Ethiopia
for several millennia. Castor bean and safflower may be indigenous
crops but there is insufficient evidence to be really certain. The same
problem is seen with okra.
At least we are on firmer ground with coffee (Coffea arabica), ensete
(Ensete ventricosum) and chat (Catha edulis). These are all Ethiopian in
origin, though from their very nature it is difficult to point to characters in them that are 'domesticated'. In other words, they are still at
the stage of wild species that have been taken into cultivation but
could probably still exist quite well in the wild. Nevertheless, they
were taken into cultivation, even though the boundary between
cultivation and domestication in these cases is not very clear.
The main problem in all the arguments and discussions, as to the
place and time of origin of Ethiopian crops, is that we have very few
hard facts to lean on in terms of reliable carbon-dated archaeological
finds. When and if these appear, as they already have in and near the
Middle East, India and the Americas, we shall be able to back up our
often tentative ideas with greater certainty. However, whether or not
Ethiopia is a centre of origin for cereal crops such as wheat, barley,
sorghum and certain millets, as well as the oil crops and pulses about
which there often is some doubt, one thing is certain: Ethiopia is
undoubtedly a world centre of crop plant diversity of great importance for a number of important domesticates. Agriculture must be
38
/. M. M. Engels & ]. G. Hawkes
very ancient in Ethiopia, and its position at the crossroads of ancient
crop migrations has resulted in the creation of a centre of genetic
diversity which all crop scientists from the time of Vavilov onwards
have considered of the greatest theoretical importance, and one of
very great practical value to plant breeders.
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Crops with wild relatives found in
Ethiopia
SUE B. EDWARDS
Introduction
All our modern crops have been developed from wild plants.
The domestication of a plant passes through stages from intensified
usage of the wild plant to the development of a domesticate so
dependent on Man that it cannot survive in the wild. All stages are
seen in the crop complement of Ethiopia. There are many wild plants
which are used for food, particularly in times of food shortage such as
the period between seed sowing and harvest. It is hardly surprising
that the majority of such plants are those used as leafy vegetables,
followed by those with edible fruits, tubers or roots. Another example
is the grass, Snowdenia polystachya (Fresen.) Pilg., whose seeds are
collected and used in a similar way to teff. The following account
includes only those that are related to domesticates. Examples of
semi-domesticated plants are Avena abyssinica and Coccinia abyssinica,
both of which are discussed further below. There are also wild plants
now attracting attention as potential crops, for example Vernonia
galamensis (Cass.) Less. (Perdue, 1988) and Cordeauxia edulis (Hemsl.)
(Polhill & Thulin, 1989). Ethiopia also has fully domesticated endemic
crops, the best known being teff, Eragrostis tef and ensete, Ensete
ventricosum. For fully domesticated plants the wild species from
which the crop developed has in some cases been identified; in others
it seems to have disappeared after the plant was domesticated.
Both environmental degradation and modern agriculture are putting traditional crops and their wild relatives at risk. The now
inadequate traditional agriculture must change if Ethiopia is to feed
itself and this is one of the major tasks being faced by the Government. However, it is hoped that the following account gives some
Crops with wild relatives found in Ethiopia
43
idea of the size of the task facing conservationists who are working to
preserve the traditional varieties and their wild relatives for use in
developing modern and appropriate cropping systems. There is no
part of the country where some crop or other and/or its wild relatives
do not occur: for example, Thymus spp. in the Afro-alpine regions;
Ensete ventricosum in the medium to higher altitudes and Gossypium
spp. in the lowlands.
But crops are not the only plants used by Man. Any consideration
of the Ethiopian wealth of plants must take into account those other
plants used by Man although it is not possible to cover them in detail
in the present account. An important group of useful plants is those
used in traditional medicine. There is no precise, modern account of
all these plants. However, the Ministry of Health now has a department responsible for studying traditional medicine and assisting its
practitioners. Although some of the better known plants, such as
Brucea antidysenterica J. F. Mill. (Simaroubaceae) and Hagenia abyssinica
(Bruce) J. F. Gmel. (Rosaceae), are left when surrounding vegetation
is cut down and naturally occurring seedlings may be moved and/or
protected, the major medicinal plants in Ethiopia are on the whole
not cultivated. However, some cultivated herbs are also used medicinally and these are mentioned below.
The Ethiopian region, including the Sudan and Somalia, is well
known for its resins and gums. These come mainly from three
genera, Acacia in the Fabaceae (Leguminosae), and Boswellia and Commiphora in the Burseraceae. True gum arabic is extracted from Acacia
Senegal (L.) Willd. which grows most abundantly in the lowlands of
Ethiopia, Somalia and the Sudan (Asfaw Hunde & Thulin, 1989).
Boswellia and Commiphora have a centre of diversity in the SomaliaMassai regional centre (White, 1983) which gives the Ogaden region
of Bale, Gamo Gofa, Harerge and Sidamo its distinctive vegetation.
There are 52 species of Commiphora recorded for Ethiopia, of which 35
(67 per cent) belong to the Somalia-Masai region and are found only
in south and south-eastern Ethiopia (Vollesen, 1989b).
Ethiopian agriculture is heavily dependent on animals for power
and the forage and browse for these all has to come from natural
vegetation and crop residues. Although germplasm taken from
eastern Africa has been developed elsewhere into important forage
plants, for example Chloris gayana Kunth. (Bermuda grass), work on
indigenous forage plants is only beginning. This has already shown
Ethiopia to be a centre of diversity for Trifolium, with 10 of its 26
indigenous species being endemic (Thulin, 1989).
44
Sue B. Edwards
Then there are the many plants used as cosmetics, producing perfume and colouring, as fumigants and cleansers, and as dyes and
inks. There are also the many plants used to construct houses and
furniture, make agricultural tools and provide fuelwood. The
majority of these species are wild plants. They will not be replaced by
plastic and other artificial materials as has happened in developed
countries, because the source for these artificial materials is the fossil
fuels which have a limited world supply. Thus conservationists must
consider the whole Ethiopian environment if these many useful
plants are to be available for future generations.
However, the following account gives only the known occurrences
in Ethiopia of the wild relatives of crops. The crops include both those
grown in Ethiopia and those not grown in this country but which
have some importance in international trade, for example pistachio
nut and the drug senna, and which have wild relatives in the country.
The common name and usage for each of the different crops is given
in Table 1.
Dicotyledons
Amaranthaceae
Amaranthus spp. The account for the Flora of Ethiopia gives 11 common weeds in fields and in open disturbed habitats. In many places
the young plants are eaten cooked and there are records of the seeds
of A. caudatus being used in making 'tala' (Townsend, in preparation).
Anacardiaceae
Pistacia vera L. There are two species in Ethiopia. P. aethiopica Kokwaro is recorded from Eritrea, Gamo Gofa, Sidamo and Bale from a
variety of habitats. P. falcata Mart, grows in Shewa where it is the
dominant tree on recent lava flows. It is also recorded from Eritrea
(Gilbert, 1989b).
Apiaceae (Umbelliferae)
Anethum graveolens L. is semi-cultivated, being also found growing
wild (Heywood et ah, in preparation; personal observation).
Apium graveolens L. is said to occur as an escape from cultivation.
There are two wild species, A. nodiflorum (L.) Lag. which grows in
wet places in Eritrea, Tigray, Gondar and Harerge and A. leptophyllum
(Pers.) Muell. ex Benth., which is a weed (Heywood et al., in
preparation).
Carum carvi L., Coriandrum sativum L., Cuminum cyminum L., and
Table 1. Crops with important gene pools in Ethiopia with their common names and uses
Scientific name
(Family)
Common names in English and Amharic
Part used in Ethiopia
Abelmoschus esculentus
Okra, Ladies Fingers
Young fruit used as cooked vegetable.
Amaranthus caudatus
A. hybridus
Grain Amaranths, 'alma'
Seed for brewing; young plants as cooked vegetable.
Anethum graveolens
Dill, 'insilal'
Apium graveolens
Celery
Whole plant used in traditional medicine and to
flavour alcoholic drinks.
Leaves and stems as herb: fruits as a spice.
Brassica campestris
B. carinata
B. integrifolia
Field mustard
Ethiopian Kale, 'gomen' for young
plants and leaves, 'gomenzer' for seeds
used for oil
Indian mustard
Mustard 'senafetch'
Young plants and leaves used as leafy vegetable;
seeds used for crushing and oiling the baking plate
for cooking 'injera'.
Cabbage, etc.
'tikil gomen', etc.
Jack Bean or Sword Bean
Used as cooked vegetable and salad.
Dicotyledons
(Malvaceae)
(Amaranthaceae)
(Apiaceae)
(Apiaceae)
(Brassicaceae)
Brassica juncea
B. nigra
(Brassicaceae)
Brassica oleracea
(Brassicaceae)
Canavalia ensiformis
(Fabaceae)
Cannabis sativa
(Cannabaceae)
Hemp (fibre)
Cannabis (drug) 'itse faris'
Seeds used to prepare a special fasting dish 'siljo'.
Young pods and seeds may be used as vegetable.
Leaves used medicinally; elsewhere grown for fibre.
Table 1 (cont.)
Scientific name
(Family)
Capsicum spp.
(Solanaceae)
Carthamus tinctorius
(Asteraceae)
Carum carvi
(Apiaceae)
Catha edulis
(Celastraceae)
Cicer arietinum
(Fabaceae)
Citrullus lanatus
(Cucurbitaceae)
Citrus aurantifolia
(Rutaceae)
Coccinia abyssinica
(Cucurbitaceae)
Coffea arabica
(Rubiaceae)
Corchorus oligatorius
(Tiliaceae)
Coriandrum sativum
(Apiaceae)
Common names in English and Amharic
Part used in Ethiopia
Chili pepper, 'karya' for fresh fruit;
'berbere' for red mature fruit
Fruits eaten fresh and dried fruits are the main spice
in 'berbere'.
Safflower, 'suf
Fruits for edible oil.
Caraway
Fruits used to flavour drinks, cakes and bread.
Khat, 'chat'
Leaves used as stimulant.
Chickpea, 'shimbira'
Ripe seed as a pulse: green seeds also eaten fresh.
Watermelon, 'birchik', also 'habhab'
Fruit eaten fresh.
Lime, 'lomi'
Fruits used fresh for sucking, cleaning meat and
treating skin problems.
Tuber eaten as a cooked vegetable.
'anchote'
Arabica coffee, 'buna'
Jute
Coriander, 'dimbilal'
Roasted seeds, as well as dried fruit walls and leaves,
used to make hot drinks.
Young plants used as leafy vegetable. (Elsewhere a
source of fibre.)
Fresh leaves sometimes as a herb: fruits used in
spicing 'berbere'.
Crambe abyssinica
(Brassicaceae)
Cucumis melo
(Cucurbitaceae)
Cucurbita spp.
(Cucurbitaceae)
Cuminum cyminum
(Apiaceae)
Cyamopsis tetragonoloba
(Fabaceae)
Daucus carota
(Apiaceae)
Diplolophium abyssinicum
(Apiaceae)
Eruca saliva
(Brassicaceae)
Ficus carica
(Moraceae)
Foeniculum vulgare
(Apiaceae)
Gossypium spp.
(Malvaceae)
Guizotia abyssinica
(Asteraceae)
Hibiscus cannabinus
(Malvaceae)
Crambe
Only use recorded is medicinal.
Melon
Fruit eaten fresh.
Pumpkin, 'duba'
Fruit eaten as cooked vegetable, both fresh and after
drying.
Cumin, 'cumin7
Cluster Bean
Carrot, 'carrot'
'dog'
Garden rocket, 'jirjir' (Tigrinya)
Fig, 'beles'
Fennel
Cotton, 'tit'
Niger seed, 'noug'
Kenaf, Jute
Fruits as a spice, particularly with 'berbere'.
Young pods may be used as a vegetable.
Root as food.
Used to flavour alcoholic drinks; also medicinal,
usually growing wild.
Young plants as salad.
Fruits eaten fresh.
Leaves and stems as salad and herb; fruits as spice
and flavouring, also medicinal.
The lint covering the seeds spun to make cloth; oil
extracted from the seeds.
Fruits for edible oil.
Elsewhere stems a source of fibre, in Ethiopia young
leaves sometimes eaten as cooked vegetable.
Table 1 (cont.)
Scientific name
(Family)
Common names in English and Amharic
Part used in Ethiopia
Indigofera arrecta
I. tinctoria
Indigo
Indigo
Leafy branches for dye.
Hyacinth Bean, Lablab
Bottle gourd, 'kil'
Whole plant as forage: elsewhere white seeded
varieties as cooked pulse.
Dried fruit as container.
Grass Pea, 'guaya'
Seed as a pulse.
Lentil, 'miser'
Seeds as a pulse.
Cress, 'feto'
Seeds used medicinally; before introduction of
Capsicum used as a spice.
Seeds used as source of edible oil; also used
medicinally.
Young fruits may be eaten as cooked vegetable; dried
fruit used as a cleaning tool.
Seed eaten whole after soaking and boiling; also used
to produce high quality 'araki'.
(Fabaceae)
Lablab purpureus
(Fabaceae)
Lagenaria siceraria
(Cucurbitaceae)
Lathyrus sativus
(Fabaceae)
Lens culinaris
(Fabaceae)
Lepidium sativum
(Brassicaceae)
Linum usitatissimum
Linseed, 'telba'
(Linaceae)
Luffa cylindrica
Luffa
(Cucurbitaceae)
Lupinus albus
White lupin, 'gibto'
(Fabaceae)
Meriandra bengalensis
Sage, 'nihba' (Tigrinya)
(Lamiaceae)
Momordica charantia
(Cucurbitaceae)
Bitter gourd
Leaves used as a herb.
Young fruits may be eaten
Moringa oleifera
M. stenopetala
(Moringaceae)
Mucuna pruriens
(Fabaceae)
Nasturtium officinale
(Brassicaceae)
Nigella sativa
(Ranunculaceae)
Ocimum basilicum
O. gratissimum
(Lamiaceae)
Olea europea
subsp. africana
(Oleaceae)
Phaseolus vulgaris
(Fabaceae)
Piper nigrum
(Piperaceae)
Pistacia aethiopica
(Anacardiaceae)
Pisum sativum
var. abyssinicum
(Fabaceae)
Plantago afra
(Plantaginaceae)
Plectranthus edulis
(Lamiaceae)
Horse-Radish Tree
Bengal Bean, Velvet Bean
Watercress
'tikur azmud'
Sweet Basil, 'bessobila'
Holy Basil
Olive, 'weira'
Seeds used as a source of oil; can also be used to
purify water.
Leaves used as a cooked vegetable.
Seeds can be eaten after repeated boiling.
Leafy branches as salad.
Seed used as a spice in bread and in spicing 'berbere'.
Flowering shoots used as herb in cooking and
clarifying butter; same parts used medicinally.
Leaves used medicinally and as fumigant.
Fruits eaten by children: leaves and twigs used as a
fumigant and tooth brush.
Haricot Beans, 'fasolya' for green pods;
'bolokie' for dried beans
Young pods used as cooked vegetable; dried seeds
infrequently used.
Black pepper, 'kundo-berbere'
Fruit a high quality spice.
Ethiopian mastic
Resin for mastic.
Field Pea, 'ater'
Seeds used as a pulse; and unripe seeds eaten fresh.
Psyllium
Seed a source of mucilage, but not used in Ethiopia.
Hausa Potato, 'Oromo dinich'
Tubers eaten as cooked vegetable.
Table 1 (cont.)
Scientific name
(Family)
Common names in English and Amharic
Part used in Ethiopia
Prunus persica
Peach, 'kok'
Ripe fruits eaten fresh.
Punka granatum
Pomegranate, 'roman'
Rhamnus prinoides
'gesho'
Ricinus communis
Castor, 'gulo'
Ruta chalapensis
Rue, 'tenaddam'
Salvia nilotica
S. schimperi
'antate-welakha' (Tigrinya)
'mai-sendedo' (Tigrinya)
Fruit eaten fresh; root, bark, fruit, rind of the fruit
and flowers all used medicinally.
Leaves and smaller branches used to flavour homemade beer 'tala' and honey wine 'tej'.
Used medicinally. Mostly seeds crushed and used to
oil the baking plate for cooking 'injera'.
Leafy and flowering shoots used as a spice and
medicinally.
Seeds used medicinally.
Seeds used medicinally.
Satureja spp.
Savory or Bean herb
Occasionally leafy shoots used as herbs.
Senna alexandrina
Senna, 'sono'
Leaves and fruits used medicinally.
Sesamum indicum
Sesame, 'selit'
Seed a source of high quality edible oil.
Tamarindus indica
Tamarind, 'roka' and 'humer'
Pulp from pods used in cooking and preparing nonalcoholic drinks.
Whole plant used as a herb.
(Rosaceae)
(Lythraceae)
(Rhamnaceae)
(Euphorbiaceae)
(Rutaceae)
(Lamiaceae)
(Lamiaceae)
(Fabaceae)
(Pedaliaceae)
(Fabaceae)
Thymus spp.
(Lamiaceae)
Thyme, 'tossin'
Trachyspermum ammi
'netch azmud'
Fruits used as a spice, important in spicing 'berbere'.
Vicia faba
Horse Bean, 'bekela'
Seeds used as a pulse; young seeds eaten fresh.
Vigna radiata
Mung Bean, Green Gram
Seeds used as a pulse but only found in a few areas.
Vinga unguiculata
Cowpea, 'adenguare'
Ziziphus spina-christi
'geba' and 'qwrqwra'
Young leaves and pods eaten as a vegetable; ripe
seeds used as a pulse.
Ripe fruits eaten both fresh and after drying.
Aframomum korarima
False Cardamom, 'korarima'
Seeds used as a spice important in 'berbere'.
Allium cepa
Onion, 'kei shinkurt'
Shallot, 'kei shinkurt'
Fleshy bulb used as both a vegetable and a spice;
leaves also used sometimes as spice.
Garlic, 'netch shinkurt'
Fleshy bulbils used as a spice and also medicinally.
'hamba guita' (Tigrinya)
Tuber is said to be edible.
Asparagus, 'kestenitcha'
Young shoots eaten fresh or as a cooked vegetable.
Ethiopian Oats, 'senar'
Seed used in admixture with barley as food and for
brewing.
Taro, 'godere'
Tubers are eaten as a cooked vegetable.
Lemon Grass, 'tej sar'
Leaves used as a spice and fumigant.
(Apiaceae)
(Fabaceae)
(Fabaceae)
(Fabaceae)
(Rhamnaceae)
Monocotyledons
(Zingiberaceae)
(Alliaceae)
Allium sativum
(Alliaceae)
Amorphophallus abyssinica
(Araceae)
Asparagus spp.
(Asparagaceae)
Avena abyssinica
(Poaceae)
Colocasia esculenta
(Araceae)
Cymbopogon citratus
(Poaceae)
Table 1 (cont.)
Scientific name
(Family)
Common names in English and Amharic
Part used in Ethiopia
Aerial Yam, 'kota hari' in SW Ethiopia
Yam, 'boyye'
Aerial tubers eaten as cooked vegetable.
Underground tubers can be eaten.
Eleusine coracana
African Finger Millet, 'dagusa'
Eragrostis tef
'teff
Ensete ventricosum
'enset' for food types; 'koba' for types
where leaves are used
Hordeum vulgare
Barley, 'gebs'
Grain used mainly for brewing but also used to make
food.
Grain used mainly to make a flat fermented bread
called 'injera', also used for other types of bread.
Pseudocorm is processed to form a starchy food.
Fibre from the leaves used for rope. Leaf lamina for
wrapping bread during cooking, for eating out and
for wrapping many other materials.
Grain used for both food and brewing.
Hyphaene thebaica
Dum Palm, 'dum' and 'arkokobay' for
plant; 'akaf for fruit; 'lakha' for the
leaves (all Tigrinya names)
Dioscorea bulbifera
Dioscorea alata
D. cayenensis-D.
rotundata
complex
(Dioscoreaceae)
(Poaceae)
(Poaceae)
(Musaceae)
(Poaceae)
(Arecaceae)
Musa spp.
(Musaceae)
Oryza saliva
(Poaceae)
Banana, 'mooz'
Rice, 'rooz'
Outer covering of fruit edible; leaves used to weave
mats and baskets; stems for fuel.
Ripe fruits eaten fresh; leaves used in same way as
'enset'.
Grain used as boiled food.
Pennisetum
glaucum
(Poaceae)
Phoenix dactylifera
P. reclinata
Pearl Millet, 'bultug'
Date Palm, 'temer'
'zembaba'
(Arecaceae)
Sorghum bicolor
(Poaceae)
Sorghum, 'mashila' for 'injera' types;
'zengada' for brewing types
Triticum aestivum
(Poaceae)
Triticum durum
(Poaceae)
Bread Wheat, 'sindi' or 'dabo sindi'
Triticum polonicum
T. spelta, T. turgidum
'adja'
Durum Wheat, 'sindi' or 'habesha sindi'
(Poaceae)
Zingiber officinale
(Zingiberaceae)
Ginger, 'zinjib'
Grain used to make 'injera', usually mixed with other
cereals, and other types of food.
Fruits edible fresh and dried.
Fruits are edible; leaves used for making mats.
White grain used to make 'injera', either alone or
mixed with 'teff and other types of food.
Dark grain used in brewing.
Grain used to make raised bread.
Whole grain eaten boiled or roasted.
Used in same way as bread wheat plus as chipped
grains and in making pasta.
Used to make special food for nursing mothers and
invalids.
Rhizome used as spice; also important medicinally.
Sources: personal notes and Cufodontis (1953-72); Purseglove (1968); Grieve (1976); FAO (1984).
54
Sue B. Edwards
Trachyspermum ammi (L.) Sprague ex Turrill (synonyms Atntni copticum
L. and Carum copticum (L.) Benth. & Hook, ex Hiern) are widely
cultivated throughout the highlands and are also found as escapes.
There are no wild relatives of these species in Ethiopia (Heywood et
ah, in preparation).
Daucus carota L. A wild form, often named as var. abyssinica A. Braun,
is found in grassland and bushland on better drained soil in the
highlands of Eritrea, Tigray, Gondar, Gojam, Shewa and Harerge.
There is a second species, D. hochstetteri Engl. which is endemic to
Ethiopia. It occurs in similar habitats to D. carota and is recorded from
Eritrea, Tigray, Gondar, Shewa and Sidamo (Heywood et al., in
preparation).
Foeniculum vulgare Miller is a recent introduction. It is often confused
with Diplolophium africanum Turcz. because the plants have similar
leaf types and smells. Diplolophium occurs growing wild and can be a
conspicuous member of the natural vegetation in open meadows. The
soft young stems are eaten by children (personal observation).
Asteraceae (Compositae)
Carthamus tinctorius L. According to Cufodontis (1953-72, pp. 1177-8),
C. flavescens Willd. (given as C. persicus Desf. ex Willd.) occurs only in
northern Somalia and north-eastern Sudan. The wild species in Ethiopia is C. lanatus L. which is widespread throughout the highlands. It
is a very aggressive weed in vertisols where C. tinctorius is normally
cultivated and this could be the result of introgression with C. tinctorius (student research project, unpublished).
Guizotia abyssinica (L. f.) Cass. is most likely derived from G. scabra
(Vis.) Chiov. This is a widespread weed often growing in the same
fields as G. abyssinica. However, naturally occurring hybrids are not
common. Both the crop and weedy species show a great deal of
phenotypic variation (Seegeler, 1983, pp. 87-110).
Brassicaceae (Cruciferae)
Brassica spp. According to Seegeler (1983) there are six species of
cultivated Brassica in Ethiopia. These are B. campestris L., B. carinata A.
Braun, B. integrifolia (West) Rupr., B. juncea (L.) Czern., B. nigra (L.)
Koch and B. oleracea L. There are no completely wild species given in
Cufodontis (1953-72, pp. 148-50) but throughout the highlands of
Ethiopia there are weedy forms of Brassica which are gathered to be
eaten as a leafy vegetable. Mature plants of these weedy forms may
also be collected for their seeds which are crushed and used to oil the
Crops with wild relatives found in Ethiopia
55
earthenware plate on which 'injera' is baked. These weedy forms
have not been studied intensively. Seegeler (1983, pp. 85-7) has
recorded species of Erucastrum, which mostly occur wild, being used
in a similar way to those of Brassica.
Crambe abyssinica Rich. Cufodontis (1953-72, p. 151) gives two other
species, C. kilimandscharica Schulz, which occurs throughout East
Africa and C. sinuato-dentata Petri. The latter is endemic and could
well be conspecific with C. abyssinica as both C. abyssinica and C.
kilimandscharica have been included in C. hispanica L. by Jonsell.
However, Jonsell considered the Ethiopian material to form a distinct
group within C. hispanica and this has been confirmed in chromosome
studies. Crambe seems to be a rather difficult crop to find either in
markets or growing in fields (Seegeler, 1983, pp. 82-5).
Eruca sativa Hill, occurs in northern Ethiopia and is cultivated and
eaten as a salad before the inflorescences develop. It also occurs as a
weed (Cufodontis, 1953-72, p. 145; personal observation).
Lepidium sativum L. Cufodontis (1953-72, pp. 140-2) gives four more
species, L. alpigenum Rich, found in Eritrea and Arabia, L. armoracia
Fisch. & Mey. found in Eritrea, Kenya and Arabia, L. divaricatum
Soland. subsp. subdentatum (Burch.) Engl. found in north and probably also central Ethiopia and L. intermedium Rich, found only in
Eritrea.
Nasturtium officinale R. Br. occurs naturally throughout the highlands
and is sometimes gathered and sold to expatriates (Cufodontis, 195372, p. 152).
Cannabaceae
Cannabis sativa L. is most likely found in all regions as a weed and also
sometimes cultivated. It is not used as a source of fibre in Ethiopia but
in traditional medicine to treat epilepsy and similar emotional disorders (Verdcourt, 1989).
Celastraceae
Catha edulis (Vahl.) Forssk. ex Engl. is an important crop but it also
occurs naturally in evergreen montane and medium altitude forest,
usually near the margins or along valleys with rocky slopes (Robson,
1989).
Cucurbitaceae (Jeffrey, in preparation)
Citrullus lanatus (Thunb.) Matsum. & Nakai. The second species in
Ethiopia is the wild C. colocynthis (L.) Schrad. which has small fruits
Table 2. Distribution of Coccinia spp. in Ethiopia
Species name and
altitude range
Region according to Flora of Ethiopia (1989)"
EE
AF
EW
TU
GD
GJ
WU
SU
AR
WG
IL
KF
GG
SD
BA
C. schliebenni Harms.
1220-2000m
C. adoensis
(Hochst. ex A. Rich.)
Cogn.
550-1850m
C. sp. A.
350-760m
C. abyssinica
1300-2360m
C. sp. B
1220-1350m
C. megarrhiza C. Jeffrey
1600-1800m
C. sp. C
1250-1300m.
C. grandis (L.) Voigt
300-1900m
" Regions used to describe the distribution of plants in the Flora of Ethiopia:
EW - Eritrea West, west and above 1000 m contour.
EE - Eritrea East, east and below 1000 m contour.
TU - Tigray region, west and above 1000 m contour.
AF - Afar region, east and below 1000 m contour to Eritrean border in the east and Harerge border in the south.
WU - Welo region, west and above 1000 m contour.
SU - Shewa region, west and above 1000 m contour.
GD - Gondar region.
GJ - Gojam region.
WG - Welega region.
IL - Ilubabor region.
KF - Kefa region.
GG - Gamo Gofa region.
AR - Arsi region.
SD - Sidamo region.
BA - Bale region.
HA - Harerge region.
HA
58
Sue B. Edwards
with bitter flesh. It is found below 1300 m in Eritrea, the Afar and
Harerge.
Coccinia abyssinica (Lam.) Cogn. is an endemic plant found both
cultivated and wild. The genus in Ethiopia needs further study as the
most recent account (Jeffrey, in preparation) has a further seven taxa,
three of them unnamed. The distribution of these taxa is given in
Table 2.
Cucumis melo L. and C. saliva L. There are no wild relatives of C. saliva
in Ethiopia. The wild C. melo subsp. agreslis (Naud.) Grebensc. is
found in open woodland, especially on river margins and also in
cultivation in eastern Eritrea, the Afar, Shewa and Kefa below 1100 m.
The fruits are recorded as being used as food. The cultivated subsp.
melo, is also grown. There are 10 other wild species of Cucumis found
in Ethiopia. C. humifruclus Stent has a subterranean fruit and is
recorded from woodland and wooded grassland in southern Shewa
and Sidamo. C. meluliferus E. Mey. ex Nadu, has been collected only
once from Gondar; elsewhere it is sometimes cultivated. The following eight species - C. figarei Del. ex Naud., C. ficifolius A. Rich., C.
aculealus Cogn., C. prophelarum L., C. insignis C. Jeffrey, C. dipsaceus
Ehrenb. ex Spach and the species of two unnamed taxa - are genetically compatible to varying degrees and can form hybrids. Many of
the wild species are highly poisonous but are used in traditional
medicine.
Cucurbila. There are no wild species found in Ethiopia but four species are recorded as cultivated. C. ficifolia Bouche is a perennial
recorded from Asbe Teferi in Harerge. C. moschala (Duchesne ex
Lam.) Duchesne ex Poir. is an annual recorded from the Lower Omo
Valley. C. pepo L. and C. maxima Duchesne ex Lam. are annuals which
are grown in many parts of the country but herbarium collections do
not exist to give an accurate record of the distribution of these two
species. It is highly likely that at least one of these species has been
cultivated for a long time in Ethiopia (Tewolde Berhan Gebre
Egziabher, 1984).
Lagenaria siceraria (Molina) Standl. is found both cultivated and wild
or escaped in bushland and grassland throughout the country up to
1850 m. A completely wild species, L. abyssinica (Hook, f.) C. Jeffrey,
is found in forest and scrub and is recorded from Gondar, Gojam,
Shewa, Arsi, Kefa and Sidamo between 1600 and 2750 m.
Luffa cylindrica (L.) M. J. Roem. grows wild on riverbanks and in
cultivated areas in the western lowlands where it is recorded from
Eritrea and Ilubabor and also the Webi Shebelli Valley. It may be
cultivated in some areas. The wild species L. echinala Roxb. is found
Crops with wild relatives found in Ethiopia
59
on riverbanks and along irrigation ditches in the Afar and Kefa.
Momordica charantia L. is found in cultivation at lower altitudes; it is
widely cultivated in some other countries such as India. The closely
related wild species M. balsamina L. is found in deciduous bushland
on banks and in dry beds of rivers on sandy soil in Eritrea and
Harerge. There are 11 other species of Momordica found in Ethiopia.
Euphorbiaceae
Ricinus communis L. is the only species in the genus and is generally
recognized as having originated in Africa. It is widespread
throughout Ethiopia, being grown as a house garden plant where the
ripe seeds are crushed and used mainly to oil the baking plate for
'injera'. The seeds are also collected from wild stands, which can
range from small shrubs to fairly robust trees (Seeleger, 1983, pp.
204-38). Some plants have dehiscent (wild type) fruits and others
indehiscent (cultivated type) fruits.
Fabaceae (Leguminosae), subfamily Caesalpinioideae
(Polhill & Thulin, 1989)
Senna alexandrina Mill, previously Cassia senna L., is the source of the
drug senna. Both the commercially exploited var. alexandrina and the
wild var. obtusata (Brenan) Lock occur in Ethiopia. Var. alexandrina
occurs in Eritrea and the Afar in semi-desert scrub and grassland; var.
obtusata is found in Acacia-Commiphora bushland and semi-desert in
Eritrea and Harerge.
Tamarindus indica L. is not cultivated in Ethiopia. It is found
throughout the country most frequently in river valleys, but also in
Combretum woodlands where there is adequate ground water.
Fabaceae (Leguminosae), subfamily Papilionoideae (Thulin,
1989)
Canavalia ensiformis (L.) DC. Its wild relative, C. africana Dunn
(synonym C. virosa (Roxb.) Wight and Arn.), occurs in Eritrea,
Harerge, Ilubabor and Gamo Gofa. In Eritrea it is cultivated as a cover
crop or to give shade.
Cicer arietinum L. The wild species, C. cuneatum A. Rich., is found in
grassland and as a weed in cultivations in Eritrea, Tigray and Shewa.
Cyamopsis tetragonoloba (L.) Taub. is recorded as cultivated in western
Eritrea as a house garden crop. Its wild relative C. senegalensis Guil
and Perr. reaches the north-eastern limit of its distribution in western
Eritrea.
60
Sue B. Edwards
Indigofera. There are four species of Indigofera in Ethiopia which were
once important as the source of the internationally traded dye,
indigo. These are I. articulata Gouan found in dry grassland and
bushland in Eritrea, the Afar, Shewa and Harerge, I. arrecta Hochst.
ex A. Rich, found also in dry grassland and bushland and recorded
from all parts of the country except eastern Eritrea and the Afar, I.
coerulea Roxb. with two varieties (var. coerulea from the arid coastal
plains and var. occidentalis Gillett & Ali from all the drier parts of the
country), and I. tinctoria L. found only in the Lower Omo Valley.
Only I. tinctoria has been found in cultivation in Ethiopia. Indigofera is
a large genus in Ethiopia with 78 species in the most recent account of
the family (Thulin, 1989).
Lablab purpureus (L.) Sweet (synonym Dolichos lablab. L.), has both a
wild subspecies, subsp. uncinatus Verde, which can also be cultivated
and a cultivated subspecies, subsp. purpureus; subsp. uncinatus is
widespread. Dolichos has six species, all wild, in the present treatment
of the family.
Lathyrus sativus L. can also be found growing as an escape. The wild
species found in Ethiopia are L. pratensis L. and L. sphaericus Retz.
growing in upland grassland. There are two more introduced
cultivated species, the ornamental L. odoratus L. and the forage L.
aphaca L. which is found as an escape in Eritrea.
Lens culinaris Medik. The wild species, L. ervoides (Brign.) Grande,
grows in montane grassland and is found in Tigray, Gondar and
Shewa.
Lupinus has six species, L. albus L. (synonym L. termis Forssk.) which
is cultivated, particularly in Gojam and Gondar, and four more introduced and being grown by Soil and Water Conservation Projects. The
sixth, L. princei Harms., is found in grassland in southern Sidamo.
Mucuna pruriens (L.) DC. var. utilis (Wall, ex Wight) Bak. ex Burck.
The endemic M. melanocarpa Hochst. ex A. Rich, (sometimes misidentified as M. pruriens var. pruriens) is found in woodland and forest
margins in western Eritrea, Tigray, Welega, Arsi, Harerge, Kefa,
Gamo Gofa and Sidamo.
Pisum sativum L. var. abyssinicum (A. Br.) Alef. is endemic and only
known as a cultivated plant.
Vicia faba L. has no close relative in Ethiopia. V. villosa Roth, is
cultivated for forage and has escaped in some areas of Shewa and
Arsi. V. sativa L. var. sativa is reported to be cultivated for fodder
while var. angustifolia L. is wild and widespread in upland grassland
and scrub. V. paucifolia Bak. and V. hirsuta (L.) S. F. Gray are wild
species from montane grassland.
Crops with wild relatives found in Ethiopia
61
Vigna unguiculata (L.) Walp. is divided into five subspecies - three
cultivated and two wild. Subsp. unguiculata and subsp. cylindrica
occur as unimproved landraces with good drought resistance in
Eritrea and Harerge; there are no recent records of subsp. sesquipedalis
(L.) Verde. Subsp. dekindtiana (Harms.) Verde, occurs wild in Eritrea,
Tigray, Gondar and Ilubabor and subsp. mensensis (Schweinf.) Verde,
in Eritrea, Kefa and Gamo Gofa.
V. radiata (L.) Wilczek (synonym Phaseolus radiatus L.) has a wild
variety, var. sublobata (Roxb.) Verde, recorded from Tigray and
Gondar. Material of this variety was collected in the early 19th century. There are 15 other wild species of Vigna recorded from Ethiopia.
Lamiaceae (Labiatae)
Ocimum basilicum L. is an important cultivated spice and herb occurring in two varieties, var. basilicum and var. thyrsiflorum (L.) Benth. It
is part of a complex of four species. O. canum Sims is smaller than O.
basilicum and grows both wild and cultivated. O. forskolei Benth. is a
wild species close to O. basilicum with white or light blue flowers
which is found in many drier parts of the country. O. stirbeyi
Schweinf. & Volkens is confined to the Ogaden region of Sidamo,
Harerge, Bale, northern Kenya and southern Somalia (Ryding &
Sebald, in preparation).
O. gratissimum L. has probably been introduced and is found only
in cultivation. However, this species belongs to a group of species
which are all used in a similar way and which are found both wild
and cultivated. The most widespread are O. urticifolium Roth,
(synonym O. suave Willd.) and O. trichodon Baker ex Gurke which can
be difficult to distinguish from O. gratissimum. O. lamiifolium Hochst.
ex Benth. is a more distinctive plant which is important in traditional
medicine. It is usually found in forests and abandoned in fields.
However, it has also been seen in gardens. The remaining two species in this group, O. spicatum Deflers and O. jamesii Sebald, are both
wild and found in drier parts of the country (Ryding & Sebald, in
preparation).
Plectranthus edulis (Vatke) Agnew, synonym Coleus edulis Vatke,
occurs both wild and cultivated for its small irregularly shaped edible
tubers. The crop is found in the wetter south and south-west, but
wild forms are found throughout the country. Plectranthus is a large
genus in Ethiopia with about 30 wild species recorded. The nearest to
P. edulis is P. punctatus L'Herit. which is sometimes considered conspecific with it. However, P. punctatus never forms tubers. The other
species which forms tubers is P. esculentus N.E. Br. but this has not
62
Sue B. Edwards
been recorded as growing in Ethiopia. Solenostemon is closely related
to Plectranthus and S. rotundifolius (Poir.) Morton has tuberous roots
which are edible, but it has not been confirmed as grown in Ethiopia.
There are three other species of Solenostemon found in Ethiopia (Ryding & Morton, in preparation).
Salvia spp. Although none of the indigenous species are cultivated, S.
nilotica Juss. ex Jacq. and S. schimperi Benth. have seeds with a good
oil content (Seegeler, 1983). The ornamental species have all been
introduced. The shrub Meriandra bengalensis (Konig & Roxb.) Benth.
has often been mistaken as belonging to the genus Salvia. It is found
both wild and cultivated in Eritrea and possibly also Gondar (Ryding,
in preparation).
Satureja spp. and Thymus spp. The two species of Thymus, T. schimperi
Ronniger and T. serrulatus Hochst. ex Benth. are extensively collected
for the local market from their natural habitat above 2500 m. They are
never cultivated and the wild populations show considerable
phenotypic variation. There are 10 species of Satureja recorded in
Cufodontis (1953-72, pp. 821-5), the most widespread being S. biflora
(Ham. ex Don) Briquet. None are used as extensively as Thymus.
Other species widely grown in house gardens for their aromatic
foliage are Origanum majorana L. and Rosmarinus officinalis L.
Linaceae
Linum usitatissimum L. Cufondontis (1953-72, pp. 354-6) gives four
other species: L. holstii Engl., found in southern Ethiopia and eastern
Africa; L. keniense Fries, a rare plant found only in southern Ethiopia
and northern Kenya; L. strictum L., found in northern Ethiopia; and
L. trigynum L. var. sieberi (Planch.) Cuf., recorded from Eritrea, Shewa
and Harerge. These are plants of grassland and edges of woodland.
The closest relative of L. usitatissimum, L. bienne Miller, does not occur
in Ethiopia (Seegeler, 1983).
Lythraceae
Punica granatum L. has been grown in the northern and central highlands for a long time but it is not found wild. It is now found
cultivated in most of the larger towns above 1500 m. The only other
species in this genus is P. protopunica Balf. f. which is endemic to
Socotra (Gilbert, in preparation, a).
Malvaceae
Abelmoschus esculentus (L.) Moench. A. ficulneus (L.) Wright & Arn.
Crops with wild relatives found in Ethiopia
63
occurs in lowland Eritrea and Gamo Gofa in grassland on seasonally
waterlogged black cotton soil. It closely resembles A. esculentus and is
probably more widespread than existing records suggest (Vollesen, in
preparation, a).
Gossypium has eight species recognized in the treatment for the Flora
of Ethiopia (Vollesen, in preparation, a). These fall into three groups:
- indigenous wild species whose seeds are glabrous or covered
with short brown hairs;
- indigenous cultivated species which are diploid and have
seeds covered in a cottony lint which does not separate
cleanly from the seed; and
- introduced cultivated species which are tetraploid and have
seeds covered in a cottony lint which separates cleanly from
the seed.
Wild species
Gossypium anomalum Wawra & Peyr. subsp. senarense (Fenzl. ex
Wawra & Peyr.) Vollesen is found in the western Eritrean lowlands at
the eastern end of its range in Africa. It grows on alluvial soil in Acacia
bushland and grassland. This is a B genome species (Saunders, 1961).
G. somalense (Giirke) Hutch, is recorded from the Awash Valley,
Sidamo and Harerge on gravelly granitic, volcanic or limestone soils.
This is an E genome species (Saunders, 1961) also found outside
Ethiopia. G. bricchettii (Ulbr.) Vollesen, known only from Bale
administrative region and southern Somalia, is found in open AcaciaCommiphora bushland on gypsum hills. G. benadirense Mattei has been
collected from the Dolo area of Sidamo, north-east Kenya and
southern Somalia in similar habitats to G. bricchettii. These two species were included in G. somalense by Hutchinson (1947) and Fryxell
(1980) but Vollesen (in preparation, a) considers them to be distinct
although closely related to G. somalense. The latter two species probably have the same genome type as G. somalense and are an example
of the important species-rich flora of the Somalia-Masai region which
has a high level of endemism (White, 1983).
Cultivated species
The use of cotton has a long history in Ethiopia as seen in the
clothing of the woman in the 'Statue of Haoulti' from the Pre-Axumite
period (de Cotenson, 1981, p. 360). The indigenous cultivated species
are G. arboreum L. and G. herbaceum L. According to Seegeler (1983) G.
arboreum seems to have disappeared completely from cultivation
64
Sue B. Edwards
although it is sometimes found in a feral state. Ramanathan (1947) is
doubtful if this species is G. arboreum. He suggests that it may well
have been perennial G. herbaceum. It was last recorded in cultivation
from the lower parts of the Webi Shebele Valley around 1960. G.
herbaceum is still cultivated but to a reduced extent in the Konso area
together with G. hirsutum. It used to be widespread in northern Ethiopia but there are no recent records. It does not seem to have become
established as an escape.
The introduced species, of American origin, are G. hirsutum L. and
G. barbadense L. Both were introduced before the 1830s as there are
records from the time of W. G. Schimper (1837-78). Formerly G. hirsutum was the species most commonly cultivated by peasant farmers
but G. barbadense has become increasingly popular since it was introduced on a larger scale by the Italians around 1910.
Hibiscus is a large genus with 48 species in 10 sections. Both H.
cannabinus L. and H. sabdariffa L. belong to section Furcaria which has
six other wild species in Ethiopia (H. diversifolius Jacq., H. berberidifolius A. Rich., H. sparseaculeatus Bak. f., H. surattensis L., H. rostellatus
Guill. & Perr. and H. noldeae Bak. f.), all of which are widespread, and
H. acetosella Welw. ex Hiern. which is an introduced ornamental.
Hibiscus cannabinus L. occurs wild in a variety of habitats (Acacia
woodland and wooded grassland on grey to black alluvial soil,
swamps, seepages, etc.) where there is sufficient ground moisture.
There are no records of its cultivation except at the research level.
Hibiscus sabdariffa L. is the only cultivated species (Vollesen, in
preparation, a).
Moraceae
Ficus carica L. The closely related F. palmata Forssk. is a common shrub
or small tree in hedgerows, secondary scrub, forest edges and
riverine forest and scrub in Eritrea, Tigray, Gondar, Gojam, Welo,
Shewa, Harerge, Arsi and Kefa (Friis, 1989).
Moringaceae
Moringa oleifera Lam. Of the 14 species recognized in this genus, nine
are more or less endemic to north-east Africa. M. oleifera has only
been recorded from Harerge and Eritrea but could occur more widely.
M. peregrina (Forssk.) Fiori is recorded from eastern Eritrea and the
Afar below 700 m. The other four species are all part of the SomaliMasai regional flora (White, 1983) and include M. stenopetala (Bak. f.)
Cuf. which is an important and conspicuous part of the agricultural
Crops with wild relatives found in Ethiopia
65
system practised in the Konso area of Gamo Gofa (Verdcourt, in
preparation).
Oleaceae
Olea europea L. Subspecies africana (previously O. africana Mill.) is
found throughout the drier parts of the highlands. There have been
two expeditions by American entomologists to Eritrea to find natural
enemies of the Olive Black Scale, Saissetia oleae (Bernard), which
occurs naturally on O. europea subsp. africana. The scale was causing
devastation to citrus orchards in California but after the introduction
of the parasite, Metaphycus helvolus, the Black Olive Scale was reduced
to a minor pest (Andemeskel, 1987).
Pedaliaceae
Sesamum indicum L. is found both cultivated and escaped or wild
below 1800 m. Cufodontis (1953-72, p. 918) records two other species,
S. alatum Thonn. from Eritrea, and S. latifolium Gillett from western
Gojam and Gondar.
Piperaceae
Piper nigrum L. Piper has three species in Ethiopia of which P.
guineense is sometimes eaten and has a fruit similar to P. nigrum
(Gilbert, in preparation, b).
Plantaginaceae
Plantago afra L. (synonym P. psyllium L.) grows wild in northern
Ethiopia and can be abundant on poorer, well drained soil. There are
five other wild species found in Ethiopia (Cufodontis, 1953-72, pp.
980-2).
Ranunculaceae
Nigella sativa L. is reported to occur only in cultivation but it has been
found growing abundantly as an ephemeral on a steep bank beside
the road in the Bale mountains. There are no other species recorded
for Ethiopia (Cufodontis, 1953-72, p. 106; personal observation).
Rhamnaceae
Rhamnus prinoides L'Herit. grows as both a cultivated plant and a
natural component of montane and riverine forest, usually on the
edges or in clearings, in all parts of the country from 1400 to 3200 m.
R. staddo A. Rich., which is used in a similar way to R. prinoides,
66
Sue B. Edwards
occurs only wild, usually at the edges of montane forest, in wooded
and scrub grassland from 1400 to 2900 m in Eritrea, Tigray, Gondar,
Shewa, Arsi, Kefa, Gamo Gofa, Sidamo, Bale and Harerge (Vollesen,
1989a).
Ziziphus jujuba Mill. The preferred fruit, Z. spina-christi (L.) Desf., is
found in wooded grassland on limestone slopes, Acacia bushland, in
and along dry riverbeds, as well as edges of cultivation and gardens
up to 2400 m in Eritrea, the Afar, Tigray, Gondar, Welo, Shewa,
Gamo Gofa, Bale and Harerge. Other species with edible fruits are: Z.
abyssinica Hochst. ex Rich, (sometimes considered a subspecies of Z.
jujuba), with a similar distribution to Z. spina-christi but growing in
woodland, wooded grassland and bushland; Z. mucronata Willd.,
growing in a wide range of dry woodland in all areas except Tigray,
the Afar and Welega; Z. mauritiana Lam., recorded from riverine
thickets and riverbanks in southern Ethiopia; and Z. hamur Engl.,
confined to soils derived from limestone and gypsum in Sidamo, Bale
and Harerge. None of these have as good-tasting a fruit as Z. spinachristi (Vollesen, 1989a; personal observation).
Rosaceae
Prunus persica (L.) Batsch. has been grown in house gardens
throughout the highlands of Ethiopia for a long time and was widespread in the early 16th century (Alvares, 1961). Although suffering
severely from peach curl, with adequate moisture, the trees produce
large crops of small green to yellow fruits.
Rubiaceae
Coffea arabica L. is found throughout the country, mostly between
1500 and 1900 m. It can occur as low as 1000 m in the very wet extreme
south-west and as high as 2500 m in gardens and backyards. It grows
as a genuinely wild, moist montane forest shrub or small tree, as a
semi-wild crop in moist montane forests, as a properly cultivated crop
in shade under rainfed conditions in the moist montane forests, as an
irrigated crop without shade in some drier areas, and as a garden
plant often mixed with fruit trees and herbs in the backyard, or
simply watered from water jars (Tewolde Berhan Gebre Egziabher,
1990).
Rutaceae
Ruta chalepensis L. (often misnamed as R. graveolens L.) is very widely
cultivated but is not known as an escape (Gilbert, 1989a).
Citrus aurantifolia (Christm.) Swingle is the most widely cultivated
Crops with wild relatives found in Ethiopia
67
citrus and is sometimes found naturalized (Gilbert, 1989a). Like
Prunus persica it has been grown in Ethiopia for a long time (Alvares,
1961).
Solanaceae
Capsicum spp. Introduced some time in the 16th or early 17th century
(Tewolde Berhan Gebre Egziabher, 1984), Capsicum now shows a
wide range of types cultivated for different purposes throughout the
country. Cufodontis (1953-72, pp. 859-61) records three species, C.
annum L., C. frutescens L. and C. abyssinicum Rich. C. abyssinicum is
sometimes considered conspecific with C. frutescens.
Solanum is a large genus in Ethiopia with over 50 species recorded by
Cufodontis (1953-72, pp. 861-80). Neither S. tuberosum L. nor S.
melongena L. has wild relatives in the country. The wild species are of
interest because a number of them are used in traditional medicine,
including S. marginatum L., and in removing the hair prior to tanning
animal skins.
Tiliaceae
Corchorus olitorius L. grows in grassland on black cotton soil, along
riverbeds and as a weed of irrigated fields. It is not cultivated in
Ethiopia. There are nine other species found in Ethiopia; none of
them are cultivated but some are collected at a young stage and eaten
as a cooked vegetable (Vollesen, in preparation, b).
Monocotyledons
Alliaceae
Allium ceya L. (including both shallot and onion) and A. sativum L.
The native landraces of A. cepa are all shallots, onion being a recent
introduction. Shallot is variable, ranging from landraces cultivated for
their well developed bulbs to those with virtually no bulbs, the whole
plant being chopped up as a vegetable. There are two wild species, A.
alibile Rich., which is sometimes included in A. ampeloprasum L. from
northern Ethiopia, and A. subhirsutum L. subsp. spathaceum (Steud. ex
Rich.) Duyfjes from Eritrea, Gondar and Harerge (Tewolde Berhan
Gebre Egziabher, in preparation).
Araceae
Amorphophallus abyssinicus (Rich.) N. E. Brown. Cufodontis (1953-72,
p. 1501) gives two more species, both endemic, A. gallaensis (Engl.)
N. E. Brown from Sidamo and A. gomboczianus Pichi-Sermolli from
Gondar, Gojam, south-western Shewa and Kefa.
68
Sue B. Edwards
Colocasia esculenta (L.) Schott is widely grown in the wetter south and
south-west. It is also recorded from Eritrea and Gojam. It easily
escapes and can appear to occur spontaneously (Cufondontis, 195372, pp. 1501-2).
Arecaceae (Paltnae)
The importance of palms as multi-purpose plants in traditional economies is being increasingly realized. The following are
found in Ethiopia.
Borassus aethiopum Mart, is found in the lowlands and river valleys of
western Ethiopia and south-western Sidamo (Cufodontis, 1953-72, p.
1499).
Hyphaene thebaica (L.) Mart, is cultivated in the lowlands of Eritrea,
Tigray, Gondar and Harerge including the Afar. According to
Cufodontis (1953-72, pp. 1496-9) there are two other species in Ethiopia: H. dankaliensis Beccari, found only in eastern Eritrea and Djibouti;
and H. nodularia Beccari, recorded only from western Eritrea and
southern Gamo Gofa. A further 10 species are recorded from Somalia,
nine of them endemic.
Phoenix dactylifera L. is recorded from Eritrea, Tigray and Gamo Gofa.
P. reclinata Jacquin is found throughout the country and also outside
Ethiopia, but P. abyssinica Drude is an endemic found in Eritrea,
Tigray, Gondar, Gojam, Sidamo, Gamo Gofa, Kefa and Ilubabor.
Asparagaceae
Asparagus spp. Cufodontis (1953-72, pp. 1562-6) records 10 species
for Ethiopia. They grow from the dry hot lowlands to the cold, frostprone mountains just below 3000 m. The most widespread species in
the highlands are A. africanus Lam. and A. asiaticus L. The young
shoots are eaten by children and are sometimes found being sold in
Addis Ababa (personal observation).
Dioscoreaceae
Dioscorea. Yams are not a staple crop in Ethiopia although both root
and aerial tubers, with aerial tubers being more common, can be
found in local markets of the wetter, western half of Ethiopia (personal observation). Tubers of wild species are said to be eaten in times
of food shortage.
Miege (1986) has identified seven species and two species groups
from herbarium collections. Dioscorea bulbifera L. has both cultivated
and wild forms, the latter are said to be violently poisonous. The
Crops with wild relatives found in Ethiopia
69
other wild species with aerial tubers is D. schimperana Kunth. It is
recorded as thriving on terraces between 800 and 2100 m where
annual rainfall is between 900 and 1400 mm. It grows wild in Acacia
and Combretaceous woodlands and at forest edges. Most cultivated
yams belong to the section Enantiophyllum and material of both D.
alata L. and the D. cayenensis Lamk.-D. rotundata Poir. complex have
been identified. Some cultivars of D. alata produce both root and
aerial tubers. Enantiophyllum also includes the D. abyssinica Hochst.,
D. lecardii de Wild, and D. odoratissima Pax complex, all of which only
occur wild. D. abyssinica grows in hilly areas covered in wooded
grassland and Combretaceous woodland between 1000 and 1800 m.
Of the remaining four species, the most peculiar is D. gilletti MilneRed, with its nearest relatives in the Pyrenees of south-west Europe.
It is the most drought-resistant of all the Ethiopian species being
found in areas with less than 700 mm of rain a year. The distribution
of the various species according to Miege (1986) is given in Table 3.
Musaceae
Ensete ventricosum (Welw.) Cheesman occurs throughout the country
both wild and cultivated wherever there is adequate moisture. Its
optimal altitudinal range is between 1600 and 2400 m but it can be
found up to 3000 m in the Gurage highlands and below 1000 m in the
wet south-west. Wild stands grow in forests and are found as far
north as Tigray in isolated moist pockets of forest. This crop shows a
very wide range of variation which has not been systematically
studied throughout its range (Food and Agriculture Organization,
1984; personal observation).
Musa spp. are grown both on a large scale and by peasant farmers.
The latter show quite a range of types which do not get into the larger
markets because they do not travel well. There are no wild species of
Musa in Ethiopia, although stands may be found apparently
separated from any habitation. These are either left after a household
has moved or have been deliberately planted in a place, such as a
sheltered valley, which is more favourable for the production of the
crop (FAO, 1984; personal observation).
Poaceae (Gramineae)
Avena abyssinica Hochst. This is the oat often grown in a mixture with
barley. It is closely related to the weedy A. vaviloviana (Malz.) Mordv.
and hybrids are formed where these two species meet. Specimens
from these two species, as well as the hybrid between them have also
Table 3. Distribution of Dioscorea spp. in Ethiopia
Region according to Flora of Ethiopia (1989)*
Species name and
altitude range
D. quartiniana
D. dumetorum
D. cochleariapiculata
D. gillettii
D. bulbifera
D. schimperana
D. alata
D. cayenensisrotundata
D. abyssinica
D. odoratissima
a
EE
X
AF
EW
TU
GD
X
X
X
X
X
X
GJ
WU
SU
X
AR
X
WG IL
KF
X
GG
SD
X
X
BA
HA
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Abbreviations as for Table 2.
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Crops with wild relatives found in Ethiopia
71
been referred to the species A. barbata Pott, from which the Ethiopian
species may have been derived. A. barbata Pott., in the narrow sense,
does not occur in Ethiopia (Phillips, in preparation).
Cymbopogon citratus (DC. ex Nees) Stapf is cultivated in house gardens
throughout the country. According to Cufodontis (1953-72, pp. 13925) there are seven other species, C. commutatus (Steud.) Stapf., C.
excavatus (Hochst.) Stapf., C. floccosus (Schwfth.) Stapf., C. giganteus
(Hochst.) Chiov., C. nervatus (Hochst.) Chiov., C. proximus (Hochst.
ex Rich.) Stapf. and C. schoenanthus (L.) Sprengel, all of which contain
aromatic essential oils. All these wild species grow below 1700 m.
Eleusine coracana (L.) Gaertn. can grow as an escape. It also forms
hybrids with both subspecies of E. indica (L.) Gaertn., subsp. indica
and subsp. africana (Kennedy-O'Byrne) Phillips. Both subspecies are
found as weeds and in open disturbed habitats from sea level to
2400 m. Hybrids of subsp. africana with the crop species usually have
longer and narrower spikes than true E. coracana; also the spikelets
are not so closely packed on the rachis, the grain is intermediate in
size and the spikelets usually shatter (Phillips, 1974 and in
preparation).
Eragrostis tef (Zucc.) Trotter. The closest wild relative is generally
considered to be E. pilosa (L.) P. Beauv. This species is recorded from
Eritrea, Tigray, Gondar and Shewa where it grows as an annual in
open places and as a weed in cultivated fields, often near ditches
(Jones, 1988; Phillips, in preparation).
Oryza sativa L. O. barthii A. Chev. is found in the Gambella plains of
western Ilubabor and O. longistaminata A. Chev. & Roehr. occurs in
the swamps and marshes up to 2500 m around Lake Tana, where it
sometimes forms pure stands (Phillips, in preparation).
Pennisetum glaucum (L.) R. Br. The supposed wild source of pearl
millet is P. violaceum (Lam.) A. Rich, with the eastern extension of its
distribution reaching the lowlands of western Eritrea (Phillips, in
preparation).
Sorghum bicolor (L.) Moench. This species forms hybrids with S.
arundinaceum (Willd.) Stapf., which occurs in Ethiopia, and possibly
other species are found in Ethiopia.
There are no wild relatives in Ethiopia of either Hordeum vulgare L.
or the several species of Triticum.
Zingiberaceae
Aframomum korarima (Per.) Engl. Cufodontis (1953-72, pp. 1594-5)
72
Sue B. Edwards
records two more species, A. polyanthum (K. Schum.) K. Schum. and
A. sanguineum (K. Schum.) K. Schum., both from Kefa and extending
from southern Sudan to north-eastern Zaire.
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and Plant Breeding, 7, 55.
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Ryding, O. (in preparation). Lamiaceae (Labiatae). To be published in Flora
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Diversity of the Ethiopian flora
TEWOLDE BERHAN GEBRE EGZIABHER
Introduction
It is generally accepted that Ethiopia is an important domestication and genetic diversification centre of crop species (Purseglove,
1968; Mooney, 1979). Likewise, it is instinctively felt that it must have
a rich flora. But this is not known quantitatively, partly because
efforts at documenting Ethiopian plants have been sporadic (Friis,
1982) and as a result, many plants remain unrecorded. On the other
hand, hasty recording has often meant that a species goes by different
names, causing double counting. The situation is made more confusing because the plant specimens collected from Ethiopia are scattered
in various herbaria, mostly in Europe. The information published on
them is equally scattered and in numerous European languages
(Cufodontis, 1953-72). Compiling information on Ethiopian plants is
thus a daunting task.
The Ethiopian Flora Project, supported financially by both the
Ethiopian Government and the Swedish Agency for Research
Cooperation with Developing Countries (SAREC), was launched to
meet this challenge. The project is therefore building a reference
herbarium and library so that, in some years' time, information on
Ethiopian plants will be found organized at one reference point in
Ethiopia. The project is training young Ethiopian taxonomic botanists
so that this information can be continually augmented, managed and
updated. It is also writing a Flora of Ethiopia, covering the whole
country, so that plant identification, both in the field and in the
herbarium, becomes possible. When the Flora has been completed, it
will be possible to compile reliable data on the number of species and
their distribution as well as on the interesting question of endemism,
thus quantifying the diversity of the Ethiopian flora.
76
Tewolde Berhan Gebre Egziabher
One volume of the Flora of Ethiopia has now been published
(Hedberg & Edwards, 1989). It should, therefore, be possible to extrapolate from it and obtain impressions of the whole flora. Another
very important source of information is Cufodontis' list of plants of
the Horn of Africa (Cufodontis, 1953-72). He collected written information on the plants of Ethiopia, Somalia and Djibouti. It is a rather
uncritical compilation, which is understandable since it is the work of
one man and was based on literature. Nevertheless, we will try to
juxtapose the information in this work with the more critically compiled information in the completed volume of the Flora of Ethiopia,
and extrapolate. This will give only a rough estimate of both the size
of the Ethiopian flora and its endemism; given the present state of
knowledge, however, this will be the best that can be obtained. But,
before extrapolating, we must look at the affinities of the Ethiopian
flora as these affinities have a direct bearing on the number of species
involved and on the likelihood of a species being endemic: in short,
on the diversity of the flora.
Affinities of the Ethiopian flora
Thulin (1983), in the introduction to his Leguminosae of
Ethiopia, has summarized the information on this issue. The highlands of Ethiopia, together with the highlands of East Africa,
Cameroun and the Sudan, constitute the Afro-montane floristic
region (White, 1978). The flora is uniform and endemism in a given
country is therefore low as many of the Afro-montane taxa of one
country will also be found in another country. Though the Ethiopian
highlands are the most extensive of the African mountainous regions,
the number of species in them is lower than in the less extensive East
African mountains. This is probably because the Ethiopian highlands
are, on the whole, drier than their East African counterparts.
To the west of the Ethiopian plateau, encroaching into it along
river valleys, is the adjoining Sudanian floristic region (White, 1979).
This region extends westwards from Ethiopia all the way to the Atlantic Ocean. Endemism in countries in this region is, therefore, understandably low as the region is divided into many countries.
To the east of the Ethiopian plateau, and thence south to northern
Tanzania, is the relatively small (in terms of area) but very distinct
Somalia-Masai floristic region. This is a region of high endemism. It
has similarities with the Madagascan and Arabian floristic regions
since the bodies of water separating it from them are narrow. It also
bears a similarity with the Kalahari floristic region in spite of the
intervening wide Zambesian floristic region.
Diversity of the Ethiopian flora
77
Since Ethiopia contains these regions, its flora has affinities with all
but Equatorial Africa. It also has affinities with Arabia, both because
of its Somalia-Masai component and because of its Afro-montane
component. In floristic and geological terms, the plateau of southern
Arabia is, in fact, African (Mohr, 1971).
The higher parts of the Ethiopian plateau, with their medium to
low temperatures, have enabled primarily Mediterranean and
temperate Eurasian taxa to occur. With increased human movement,
new taxa are being added, establishing the similarity at even specific
and infraspecific levels. There is a similarity between the floras of
Ethiopia and the Canary Islands, which is difficult to explain. Furthermore, human movement is also introducing Australian and American
taxa into Ethiopia. Who can imagine Addis Ababa without Eucalyptus
globulus, an Australian species?
In the following text some examples are given to illustrate the
affinities summarized above. The genera Cadia and Delonix are Madagascan, but C. purpurea and D. elata occur in Ethiopia. Of the many
taxa found in Arabia and Ethiopia, Rosa abyssinica and Polygala aethiopica can be mentioned, and of course teff (Eragrostis tef) from among
the montane species; Barbeya oleoides, Commicarpus pedunculosus,
Erythrococca abyssinica and a number of Commiphora and Boszvellia spe-
cies from among the Somalia-Masai species.
It is surprising that the Kalahari link exists even at the species level:
Indigofera trigonelloides is found only in Namibia and Ethiopia, and
nowhere in between (Thulin, 1983). Commiphora and Boswellia are two
characteristic genera of the Somalia-Masai floristic region, the centre
of diversity for these genera. Forty-eight Commiphora species and six
Boswellia species occur in Ethiopia (Vollesen, 1989), firmly establishing the Somalia-Masai character of its flora.
The dominant montane forest trees, perhaps Ethiopia's most conspicuous species, occur in other montane areas of Africa; Juniperus
procera, Podocarpus gracilior, Olea africana, Aningeria adolfi-friedericii and
Celtis africana can be mentioned as examples (Eggeling, 1952; Dale &
Greenway, 1961).
The Sudanian floristic character is shown in the savannahs of the
lowlands and river valleys of western Ethiopia, e.g. Panicum maximum
(elephant grass), Anogeissus leiocarpus and many savannah woodland
species of Combretum and Terminalia.
The Mediterranean and temperate Eurasian traits of the Ethiopian
flora can be illustrated with a number of genera, e.g. Festuca, Dianthus, Silene and Trifolium illustrating the temperate connection,
Scorpiurus, Medicago and Satureja illustrating the Mediterranean con-
78
Tewolde Berhan Gebre Egziabher
nection and a number of families, e.g. Cruciferae (Brassicaceae),
Umbelliferae (Apiaceae) and Caryophyllaceae illustrating both connections. Recent European introductions have been deliberate, e.g.
varieties of Brassica oleracea (cabbage), Beta vulgaris (beetroot) and Daucus carota (carrot). Note that wild forms of D. carota are native to
Ethiopia but the cultivated form is an introduction. Introductions
have been accidental and sometimes harmful, e.g. Galinsoga parviflora,
which became a serious weed; and sometimes not harmful - would it
be better to say 'not yet harmful'? - e.g. Dactylis glomerata, Silybum
marianum. Some introductions are ancient, e.g. Arundo donax which
still cannot flower in Ethiopia but keeps growing vigorously,
vegetatively.
The puzzling Canarian connection can be seen from the genera
Canarina, with only two species occurring in Ethiopia and the other
eastern African highlands; Hypagophytum, with one species in Ethiopia only (H. abyssinicutn); and Aeonium, with two species in Ethiopia
and southern Arabia, but each with numerous species in the Canary
Islands and none in the vast intervening African hinterland.
Size and endemism of the Ethiopian flora
Cufodontis' (1953-72) list includes about 6370 species. The
qualifying term 'about' is needed because Cufodontis has included
uncertain records, even only plant names without taxonomic descriptions or voucher specimens. Dealing with such records without going
to the source material involves personal judgement. Going to source
material is a long process and cannot be contemplated when making
only an estimate of species numbers; it forms a major part of the
ongoing work on the Flora of Ethiopia.
Of the 6370 species Cufodontis records, 4865 are Ethiopian, the
other 1505 being endemic to the Somalia-Djibouti region (Cufodontis,
1953-72). The number of endemics to Ethiopia is only 1182, while the
number of endemics to the whole of the Horn of Africa is 2291. These
figures can be used as the basis for estimating the size of the Ethiopian flora.
The method of estimating species numbers to be adopted here is
that of comparing the figures derived from Cufodontis' work for the
whole flora with the more precise estimate obtained from the work of
the Ethiopian Flora Project. When doing this,, the families in the
completed volume of the Ethiopian Flora cannot be taken as a random
sample for simple extrapolation because the choice of the families
involved is systematic in the taxonomic sense, and hence is not a
Diversity of the Ethiopian flora
79
random sample in the statistical sense. The choice had to be systematic because related families must come close together in a volume. A
simple extrapolation from the numbers in the completed volume to
the whole, by calibrating Cufodontis' records against the records of
the completed Ethiopian Flora volume, would thus not be justified.
Instead, a method of arriving at an informed guess is being adopted
as more realistic. Two families from the completed volume of the
Flora are being used to arrive at what intuitively feels correct. The two
families chosen are Leguminosae (Fabaceae) and Burseraceae.
The family Leguminosae is represented all over Ethiopia, and its
estimate of endemism using Cufodontis' records is about average (31
per cent for the Horn of Africa and 22 per cent for Ethiopia for
Leguminosae, compared with 36 per cent for the Horn of Africa and
24 per cent for Ethiopia for the whole flora. Endemism in the family
Burseraceae, on the other hand, is average according to Cufodontis'
records for Ethiopia (24 per cent) but very high for the Horn of Africa
(87 per cent). This suggests a much more complete recording of species for Somalia and Djibouti than for Ethiopia. More new records for
Ethiopia would, therefore, be expected in the Burseraceae than in the
Leguminosae. The work of the Ethiopian Flora has shown this to be
the case; Leguminosae has risen from Cufodontis' records of 492 to
607, an increase of only 23 per cent, while Burseraceae has risen from
Cufodontis' records of 29 to 54, an increase of 86 per cent. In either
case, the increase in the species is proportional to the number of
regional (Horn of Africa) species not found in Ethiopia as it is mostly
from among these species that new Ethiopian records can be expected. Using this formula:
(Regional species - Ethiopian species) x
Ethiopian species
:
:
h Ethiopian species
an estimate of the likely number of Ethiopian species can be obtained
from the numbers in Cufodontis' records. This will be an overestimate because it assumes that the less known regional endemics
and the better recognized widespread species equally have failed to
be recorded in Ethiopia in the past and will be equally represented in
future new records. The opposite view is to assume that only regional
endemics will continue to be discovered in Ethiopia. The reality is
probably somewhere between the two. An average of both figures is,
therefore, being used to estimate the size of the Ethiopian flora.
Predicted and actual values for the two big, and in terms of distribu-
80
Tewolde Berhan Gebre Egziabher
Table 1. Estimates of the Ethiopian flora
Number of Ethiopian species
Leguminosae
Burseraceae
All seed plants in Ethiopia
A
B
C
D
E
492
29
4865
604
50
6014
562
52
5712
583
51
5863
607
54
?
A = Counted from Cufodontis (1953-72).
B = (Regional spp. - Ethiopian spp.) x ^ ^ ^ '
+ Ethiopian spp.
C = (Regional endemic — Ethiopian endemic) x -=—.* .—*-*—
+ Ethiopian spp.
D = Average value of B and C.
E = Counted from the completed volume of Flora of Ethiopia.
tion interesting, families in the completed volume of the Flora of
Ethiopia, Leguminosae and Burseraceae, are used to check this out
(Table 1).
Using the adopted formulae, the actual numbers of species for
Leguminosae and Burseraceae are shown to be greater than the
estimated numbers. This is because, while Ethiopia adjoins the Sudan
and Kenya, Cufodontis' records exclude these two countries; yet
plants already recorded in these countries, but not yet in Ethiopia,
will also be expected to appear as new records in Ethiopia. The figure
of 5863, or roughly 6000, is therefore a low estimate for the Ethiopian
species. Since endemism in the Sudanian zone is low, the rise owing
to contribution from it will not be as large as that from the SomaliaMasai region. However, it could be estimated that it will be about half
as much, thus giving us an estimate for the seed plants of Ethiopia of
approximately 6360, rounding off to a more realistic figure, about
6500. The ferns of Ethiopia will consist of a few hundred species, thus
giving us an estimate of the higher plants of Ethiopia of about 67006900, or, more realistically, between 6500 and 7000.
Endemism in Ethiopia has been estimated by a few authors.
Brenan (1978), counting a sample portion of Cufodontis' list, arrived
at the conclusion that about 21 per cent of the species of Ethiopia are
endemic. Using the whole of Cufodontis' list, it was calculated that 24
per cent of the species are endemic. The endemism according to
Cufodontis' list is 22 per cent for Leguminosae and 24 per cent for
Burseraceae. Endemism in these two families, according to the newly
completed Ethiopian Flora volume, is 11 per cent (Thulin, 1983) and
Diversity of the Ethiopian flora
81
15 per cent (Vollesen, 1989), respectively. These figures are not very
different from each other in spite of the very high regional endemism
of Burseraceae. This is because even the small Somalia-Masai region
is divided among several countries (Ethiopia, Djibouti, Somalia,
Kenya and Tanzania), thus reducing endemism within national
boundaries. The regional (i.e. Ethiopia-Somalia-Djibouti) endemism,
considering only those species found in Ethiopia, gives a different
picture: 15 per cent for Leguminosae and 42 per cent for Burseraceae.
Endemism in the Horn of Africa is thus high, in the order of 20 per
cent, but it is reduced in Ethiopia to about 12 per cent, or about half
for the region.
The implicit belief that has existed hitherto, that the Ethiopian flora
is rich both in species numbers and in endemics is, therefore, valid.
References
Brenan, J. P. M. (1978). Some aspects of the phytogeography of tropical
Africa. Annals of the Missouri Botanical Garden, 65, 437-78.
Cufodontis, G. (1953-72). Enumeratio Plantarum Aethiopiae, Spermatophyta.
Bulletin du Jardin Botanique National de Belgique, Bruxelles.
Dale, I.R. & Greenway, P.J. (1961). Kenya Trees and Shrubs. Buchanans,
Kenya Estates Ltd, Nairobi, in association with Hatchards, London.
Eggeling, W. J. (1952). Indigenous Trees of the Uganda Protectorate, 2nd edn
revised by I.R. Dale. Crown Agents, London.
Friis, I. (1982). A list of botanical collectors in Ethiopia. University of Copenhagen (unpublished).
Hedberg, I. & Edwards, S. (eds) (1989). Flora of Ethiopia, vol. 3. The National
Herbarium, Addis Ababa University, Ethiopia, and the Department of
Systematic Botany, Uppsala University, Sweden. (Further volumes in
preparation.)
Mohr, P. (1971). The Geology of Ethiopia. Haile Selassie I University Press,
Addis Ababa, pp. 7-8.
Mooney, P. R. (1979). Seeds of the Earth. Canadian Council for International
Cooperation, Ottawa.
Purseglove, J. W. (1968). Tropical Crops, vols 1, 2, & 3. Longmans, London.
Thulin, M. (1983). Leguminosae of Ethiopia. Opera Botanica, 68, pp. 7-13.
Vollesen, K. (1989). 123. Burseraceae. In: I. Hedberg and S. Edwards (eds),
Flora of Ethiopia, vol. 3. The National Herbarium, Addis Ababa University,
Ethiopia, and the Department of Systematic Botany, Uppsala University,
Sweden, pp. 442-78.
White, F. (1978). The Afromontane region. In: M.J.A. Werger (ed.), Biogeography and Ecology of Southern Africa, vol. 1. Dr W. Junk, The Hague, pp.
463-513.
White, F. (1979). The Guineo-Congolian region and its relationships to other
phytochoria. Bulletin du Jardin Botanique National de Belgique, 49, 11-55.
Forest genetic resources of Ethiopia*
J. DE VLETTER
Introduction
The Ethiopian Government attaches a high priority to plantation forestry. Through institutions like the State Forests Conservation and Development Department, the Soil Conservation and
Community Forestry Development Department and the revolutionary mass organizations, more than 40 000 ha of degraded lands annually are put under some kind of forest-like vegetation cover, partly as
pure stands, partly in combination with other land uses such as
agriculture or grazing.
This is an effort that requires the introduction of suitable trees,
growing from seeds of known origin, if the forests (or agroforests) are
to fulfil their objectives: erosion control and production of fodder,
fuelwood, construction wood and timber. Tree species suitable for
plantation forestry should have a maximum adaptability to a wide
spectrum of prevailing (and sometimes rapidly changing)
environmental conditions. In order to meet this requirement, the
forester must be able to select from natural or planted tree populations, which must have a certain degree of genetic diversity.
Unfortunately, the Ethiopian natural tree populations have been,
and still are, subject to indiscriminate destruction. Shifting cultivation
and traditional grazing have been practised for centuries in Ethiopia.
This, and the relentless cutting for fuel and building needs by a dense
and rapidly growing population, have led to an almost complete
deforestation of the Ethiopian highlands today.
The remaining forests are very unequally distributed. The northern
* The content of this paper does not necessarily represent the ideas of the State
Forests Conservation and Development Department (SFCDD).
Forest genetic resources of Ethiopia
83
and central parts of the country are almost bare. Most of the forests
are found in the south-western and southern parts of the country,
mainly as closed broadleaved forests ('rainforests'). Elsewhere, in
areas of lower altitude, woodlands, open bush and shrublands occur.
These still cover more or less extensive areas, but overgrazing,
charcoal production and man-made fires pose a serious threat.
The closed broadleaved forests form Ethiopia's best developed forests, representing the most eastern extension of the African
equatorial rainforest belt. These forests are confined to the
administrative regions of Kefa, Ilubabor, Sidamo and Bale. They are
extremely important for the supply of raw material for the saw-milling industry. Moreover, the area decreases year after year, even
before 'domestication' of the commercial tree species could start.
Whereas Ethiopia's agriculture can already benefit from more than 10
years of collection, conservation and evaluation of agricultural crop
germplasm resources, carried out by the Plant Genetic Resources
Centre (PGRC/E) in Addis Ababa, similar work in the field of forestry
is still in its initial stages.
It is true that species elimination trials have a fairly long tradition
and that today, from a moderately wide range, suitable species can be
chosen for the numerous bio-climatic zones of Ethiopia. However,
systematic exploration and conservation of indigenous tree species is
a neglected field of activity which should receive highest priority. All
forest-like vegetation types should be included in the programme,
not only the coniferous forests and the broadleaved forests (where
most of the timber trees are found), but also the woodlands and
riparian forests. Woodlands are composed mainly of droughtresistant Acacia species, which are widely used as a source of fodder
and fuel. They are also used as multi-purpose trees in agroforestry
systems and in plantation forestry in Ethiopia as well as in other
African countries.
Agro-climatic belts in Ethiopia
The characteristics of Ethiopia's natural vegetation are to a
large extent determined by two main factors:
- elevation (and temperature);
- rainfall.
On the basis of elevation, Ethiopia has been divided traditionally into
five agro-climatic belts:
1. Berha
the dry and hot belt below 500 m above sea level
84
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2. K'olla
the dry to moist, warm belt between 500 and
1500 m
3. Weyna Dega the dry to wet, moderately warm belt between
1500 and 2300 m
4. Dega
the moist to wet, temperate belt between 2300
and 3200 m
5. Wurch
the moist to wet, cold belt above 3200 m
'Dry' is defined as having less than 900 mm annual rainfall, 'moist' as
having between 900 and 1400 mm and 'wet' as having more than
1400 mm annual rainfall.
Forests are found in the moist and wet Weyna Dega and to a lesser
extent also in the moist K'olla and the moist and wet Dega. The other
agro-climatic zones carry mainly woodlands, bushlands, savannahs,
steppes or alpine formations.
Vegetation classification
The classification of the Ethiopian vegetation is still in a preliminary stage. In the introduction to his manual of indigenous trees
of Ethiopia, von Breitenbach (1963) gives a scheme of plant associations, listing more than 50 different plant communities. Knapp (1968)
has developed a complicated system of vegetation units on the basis
of dominant species.
Steppe
Nearly treeless grasslands; here and there widely scattered
shrubs occur. This vegetation type covers extensive areas in west
Ethiopia, the Danakil and Ogaden plains and the coastal parts of
Eritrea.
Savannah
Lands covered with perennial grasses, with scattered trees
and shrubs. The trees shed their leaves during the extended dry
season. Fire occurs frequently.
Lowland savannahs occupy vast areas in the Rift Valley, on the
plains surrounding Lake Tana, on the Sudan plain, on the lower
plateau escarpments and in west Kefa, south Gamo Gofa and Welega.
In the hot and dry lowland areas (Berha and K'olla), lowland steppes and lowland savannahs are found.
Steppes and savannahs are also found at the other end of the
altitudinal range, in the moist and wet Wurch zones. Their upper
Forest genetic resources of Ethiopia
85
limits are situated around 4000 m. Their lower limits are not clear on
account of the extensive land clearings and grazing in the area of
mountain woodlands and forests. On abandoned cultivations and
pastures they spread as secondary vegetation. Thus this formation,
originally confined to comparatively small surfaces at high altitudes,
now occupies a major part of the Ethiopian plateau. It is easily
recognized that this vegetation is secondary because small remnants
of former forest or woodland are often found.
Woodlands
These are lands dominated by trees, which are heavily branched and which have a height of up to 20 m. The flat crowns do not
form a closed canopy, but cover more than 20 per cent of the ground
and are leafless for some part of the year. The ground is covered with
grasses, herbs and shrubs. Fires are frequent.
Vegetation types with intermediate characteristics between savannahs and woodlands are shrublands and bushlands.
Shrublands
Lands supporting a stand of shrubs, usually not exceeding
6 m in height, with a canopy cover greater than 20 per cent. Trees are
rare. The ground cover is often poor. Fires are usually infrequent.
Bushlands
Lands supporting an assemblage of trees and shrubs, often
dominated by plants with a shrubby habit but with trees always
conspicuous, with a single or layered canopy, usually not exceeding
10 m in height and total canopy cover greater than 20 per cent.
Ground cover is poor and fires infrequent. Thickets are an extreme
form of bushlands where the woody plants form an impenetrable
closed stand.
Lowland woodlands, bushlands and shrublands include a wide
variety of woody vegetation types, often difficult to separate clearly,
mainly occurring in the upper dry K'olla and lower dry Weyna Dega
zones. They are distributed over large areas in Eritrea, the Awash
region, east and south Harerge, the Rift Valley, south Sidamo, west
Ilubabor, Welega and the slopes of the eastern and central highlands.
The main genera are various Acacias, Boswellia, Commiphora, Balanites,
Euphorbia, Combretum, Croton and many others.
Mountain woodlands are found at the other end of the altitudinal
86
/. de Vletter
range (upper moist and wet Dega, lower moist and wet Wurch). Their
physiognomy is similar to the one of lowland woodlands: an upper
(more or less) open canopy formed by 5-12 m high trees. Poor specimens of Juniperus procera occur. Other genera and species are Acacia
abyssinica, Protea, Cussonia, Hagenia abyssinica, Erica arborea, Hypericum
and Arundinaria alpina (bamboo forest).
Forests
These can be defined as a vegetation type which is dominated
by trees, forming a closed, deep and complex, often multi-storeyed,
canopy. The height of the largest trees may exceed 45 m. Most trees
are columnar in shape, having a straight and clear bole. Many species
are evergreen. The forest floor has a micro-climate which is clearly
moderated by the tree cover and carries a wide variety of herbs,
shrubs, seedlings and saplings (regeneration of the trees which form
the upper storeys).
Forests are found in the moist and wet Weyna Dega, the moist and
wet Dega and (upper) moist K'olla zones (Fig. 1).
Closed broadleaved moist forests are found in south and southwest Ethiopia (mainly Ilubabor and Kefa, partly also in Sidamo and
Bale), at elevations between 1200 and 2200 m. Rainfall is more than
1400 mm (locally even up to 2000 mm) annually and the dry period is
restricted to 2-3 months. These forests are sometimes referred to as
upland rainforests. They form the best developed forests in Ethiopia,
but are less impressive than, for instance, the equatorial rainforests of
West and Central Africa. The upper storey is discontinuous and consists of scattered 40-60 m high 'emergents' dominating the dense and
close canopy of the intermediate storey. Aningeria adolfi-friedericii is
generally the only emergent species. It forms the highest, non-continuous stratum of the forest; therefore, the term Aningeria-forest is
sometimes used. A continuous stratum about 30 m above the forest
floor consists of 10-20 species of trees, all with a comparatively similar
appearance. The following species can be found here: Albizia
schimperiana, Celtis africana, Cordia abyssinica, Croton macrostachys,
Ekebergia spp., Ficus spp., Olea hochstetteri, O. welwitschii, Ocotea
kenyensis, Polyscias spp., Sapium ellipticum, Syzygium spp., Schefflera
abyssinica, Trichilia spp. and some others.
An 8-10 m high lower storey of small trees consists of: Allophyllus,
Apodytes, Bersama abyssinica, Brucea, Teclea nobilis, Coffea arabica, Millettia ferruginea, Galiniera and many others.
Climbers are common along forest edges (where more light can
Forest genetic resources of Ethiopia
87
Fig. 1. Transect through a closed broadleaved forest showing
storeys and different size classes. Aa, Aningeria adolfi-friedericii;
As, Albizia schimperana; CRm, Croton macrostachys; CYm,
Cyathea manniana; Da, Dracaena afrotnontana; Ds, Dracaena
steudneri; Eo, Euphorbia obovalifolia; Ev, Ensete ventricosum; Fs,
Ficus sur; Lg, Lobelia giberroa; Mf, Millettia ferruginea; Mk,
Macaranga kilimandscharica; Mr, Mitragyna rubrostipulata; Pa,
Prunus africanus; Pf, Polyscias fulva; Pr, Phoenix reclinata; Sa,
Schefflera abyssinica; Se, Sapium ellipticum (from Friis ef a/.,
1982).
penetrate), but are not dominant in high forest. The herbaceous
stratum on the forest floor is rich in species, but mostly discontinuous
in mature forest. Several species of ferns, seed plants and broadleaved grasses occur. Epiphytes are frequent, particularly ferns.
Other types of closed broadleaved forests include: broadleaved
semi-humid forests (less developed than the broadleaved moist forests, forming the transition towards the coniferous forests of higher
elevations); the dense Acacia forests (which occur in dry and windy
areas between 1800 and 2400 m, and are characterized by the absence
of an upper storey of big trees and a closed crown cover dominated by
Acacia xiphocarpa trees); and riparian forests (which occur at several
different altitudes along creeks and rivers).
A second main group of forest types consists of the coniferous
forests, which fall into two groups, the Juniperus forests and the
88
/. de Vletter
Podocarpus forests. Juniperus forests occur naturally between 2500 and
3200 m, partly in areas with a long dry period of up to five months.
Their upper storey is 30-45 m high and consists of Juniperus procera
trees. The middle storey is 20 m high and consists of genera such as
Pygeum, Olea, Ekebergia and Bersama. The middle storey is more or less
discontinuous and the undergrowth is poorly developed.
Podocarpus forests occur naturally between 2000 and 2500 m, in
areas with a relatively humid climate and a well distributed rainfall.
There is a 40-45 m high closed canopy with Podocarpus gracilior, often
mixed with Juniperus, dominating over Pygeum, Ekebergia and Celtis.
Present condition of natural forests in Ethiopia
Over the past century, a rapid growth of the already dense
Ethiopian population has led to overexploitation of the land. In areas
with settled agriculture new land has been cleared at the expense of
forests. Fallow periods have tended to become shorter and shorter.
Intensive cutting for fuel and construction wood has taken place.
Pasture grounds suffer from overgrazing. All this has led to deforestation, erosion, land degradation and, connected with this, declining
agricultural productivity.
It is often assumed, though not established as a fact, that 40 per
cent of the Ethiopian highlands were still covered with closed high
forests (mainly mixed and coniferous forests) not more than a century
ago. By 1950 the forest cover had decreased to about 8 per cent.
Today, Ethiopia's forest estate is reduced to 3.6 per cent of the total
land area. Table 1 shows the various environments covering Ethiopia
and their percentage of the total land area. The figures given are
estimates based on Landsat image photo interpretation (Anonymous,
1986).
It can be seen that the total forest area now covers only 3.6 per cent
of Ethiopia's land surface, an alarmingly low figure. Also degraded
and overexploited forests are included here, and it can be safely
assumed that only a minor part of the remaining forest lands consists
of undisturbed, virgin forest.
The distribution of the remaining forests is very unequal. They are
concentrated in the less densely populated southern and southwestern parts of the country. The central and northern parts of Ethiopia are almost completely deforested (Fig. 2).
About 80 per cent of the forests are broadleaved forests ('rainforests') and mixed broadleaved/coniferous forests. The rest consists of
more or less homogeneous coniferous forests (Podocarpus and
Forest genetic resources of Ethiopia
89
Table 1. Present types of land cover in Ethiopia
Environment
Area
(km2)
Percentage
Forest lands
Afro-alpine vegetation
Woodlands
Bushlands
Xerophilous bushlands
Shrublands
Grass/range lands
Riparian vegetation
Wetlands
State farms
Agricultural lands*
Bare lands
Total
45120
1080
59195
207380
234905
264155
166990
19565
11305
4730
263300
20615
1298340
3.6
0.1
4.7
16.6
18.8
21.2
9.4
1.6
0.9
0.4
21.1
1.6
100.0
a
Agricultural lands also include lands partly in combination with other
cover types, such as forests, wood-, shrub- or grassland.
Juniperus forests). All these forests are important for the raw material
supply of the country's saw-milling industry. Although for the larger
part the forests enjoy official status as State Forest or National Priority
Area and have been or are being demarcated as such, effective
management aiming at sustained yield still has to be implemented.
The area and quality of these forests decrease year after year, mainly
due to:
- timber exploitation practices, which destroy the (potentially
sustainable) production capacity of the forest;
- agricultural encroachment, cutting for fuelwood and manmade fire;
- extensions of coffee and tea production areas;
- other changes in land use, including settlement.
The other woody vegetation type of interest in a discussion on
forest genetic resources consists of the woodlands, which cover about
4.7 per cent of the total land area. Although their area probably has
not shown such a spectacular decrease as in the case of forests,
charcoal burning, fuelwood exploitation, overgrazing and man-made
fire form a serious threat to their diversity.
Elsewhere in Ethiopia, in areas of lower altitude and less rainfall,
bushlands and shrublands still occur in sizeable areas, together covering more than 50 per cent of the total land area. These vegetation
types are not considered here.
90
/. de Vletter
SAUDI ARABIA
SOMALIA
UGANDA
Approximate scale
0
50
50
1 Forest
200 km
100 miles
^
Lake
—
Regional boundary
—
International boundary
Fig. 2. Main forest areas in south-west Ethiopia (from Chaffey,
1980).
Forest genetic resources of Ethiopia
For comparison purposes, the present area of man-made forests
can be subdivided as follows:
80 000 ha
Urban plantations
State forest plantations
120 000 ha
Community forests
200 000 ha
Total plantations
400 000 ha
The total plantation area is only 9 per cent of the total natural forest
area and represents only 0.3 per cent of Ethiopia's land area. Plans
exist to extend considerably the plantation area but it will be clear that
even if the present rate of afforestation of 40 000 ha annually could be
increased significantly, it will hardly be possible to balance the losses
which result from destruction of natural forests.
Regarding the figures for the areas occupied by the various vegetation types (Table 1) and the changes occurring at present, the following should be noted:
1. It is very difficult to give exact information on the rates of
deforestation which have taken place, since comparable data
are hardly available. Sources are scarce and these use different vegetation classification systems or are based on very
tentative area estimations. In order to close this gap a project
with foreign assistance is planned, the Woody Biomass
Inventory. The negotiations are still not concluded.
2. The estimates of plantation areas are based on numbers of
seedlings produced and are therefore only tentative.
3. Most plantations are young, in the seedling or sapling stage.
Many have only a low stocking rate. On the Landsat images
they probably do not appear as forests, but as woodlands or
even bushlands.
Despite these possible inaccuracies the conservation of the remaining
natural forests as well as the introduction of sound management
practices should receive highest priority.
Conservation and evaluation of forest genetic resources in
Ethiopia
The Forestry Research Centre of Ethiopia has made a start
with selection and improvement of a number of exotic and indigenous tree species suitable for various plantation objectives. Species
elimination trials have been established on some scale since the mid1950s and today, for a variety of environmental conditions, a choice
can be made from a modest range of suitable species.
91
92
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In view of the strong and continuing human pressure on the
remaining natural forests, and directly connected with this the
danger of irrevocable disappearance of whole populations of indigenous trees, the ongoing selection programme must be supported
and supplemented by a programme of systematic exploration and
conservation of natural forest genetic resources. This broadening of
the scale of activities should receive highest priority.
Conservation and evaluation of forest genetic resources should be
considered as part of one consistent programme, of a long-term
nature, and follow the recognized steps of:
- exploration;
- collection of seed for evaluation;
- evaluation;
- conservation;
- utilization.
Exploration and collection of seed for evaluation
Forest inventories and systematic botanical explorations of
remaining natural forests will lead to a better knowledge of the distribution of the species, will identify the useful provenances and will
indicate the populations of trees which have become endangered in
their natural habitat. Since these populations are often already
depleted as a result of man-made destruction, the number of areas
where successful seed collections can be made will be limited. One
strategy could be simply to sample what is left in a systematic way.
Under optimal growing conditions, a given species will generally
show the largest genetic variation. Populations under marginal conditions often contain valuable genes for adaptability to extreme conditions but are generally less diverse. It is very important to include
such populations as well in order to cover the complete range of
genetic diversity of a species.
Evaluation
Evaluation methods of forest genetic material by means of
comparative, replicated field trials on 'representative' sites are well
developed in Ethiopia. General aspects of species and provenance
evaluation, including experimental design, layout, assessment,
statistical analysis and interpretation have been outlined by Burley &
Wood (1976).
The first species trials in Ethiopia were established about a century
ago. Some Eucalyptus were introduced in order to find suitable species
Forest genetic resources of Ethiopia
to counter the severe shortage of fuelwood around Addis Ababa. In
the mid-1950s and the early 1960s more trials were planted in a number of regions. From 1975 onward, and after the establishment of the
Silvicultural Section of the Forestry Research Centre, more systematic
trials followed. About 130 species, indigenous as well as exotic, have
been tested and evaluated. Today, a range of suitable species for
various environmental conditions is known. In the coming years,
emphasis will not be so much on the introduction of new species as
on new provenances. It is anticipated that high gains in productivity
can be achieved by selection of the best adapted provenances for
prevailing environmental conditions.
Conservation
Conservation of forest genetic resources can be achieved in
the following ways:
- in situ conservation of representative parts of forest
ecosystems;
- ex situ conservation of seeds in specially equipped genebanks
or seed centres;
- ex situ conservation of species in plantations or field
genebanks.
In view of the circumstances prevailing in Ethiopia, in situ conservation of provenances and populations or even samples of whole
ecosystems, will probably be the main approach.
In principle, in situ conservation could well be integrated with the
existing system of national parks, managed by the Wildlife Development and Conservation Organization. A disadvantage of this possibility is that the present nine National Parks incorporate only a few
forest types, such as Podocarpus gracilior and Juniperus procera forest.
The other parks include only several types of Acacia woodlands. Not a
single park represents closed broadleaved forest (rainforest), the
natural forest type which still covers the widest area in Ethiopia.
A better possibility would be to integrate in situ conservation of
forest genetic resources with the existing system of officially
designated State Forests and National Priority Areas. For these forests at least the intention exists to apply some kind of systematic
management, and although this management still has to be
implemented, this intention might provide the guarantee for protection in the long run. For the time being, additional management
provision must be made in order to achieve real protection and
prevent unwanted human disturbance or cattle grazing.
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In situ conservation requires the reservation of larger areas than ex
situ conservation, whereby research has to find an answer to the
question of whether many relatively small areas are to be selected or
few relatively large areas. For management reasons, the second
option seems to be the more realistic.
Conservation of forest tree seeds, for instance within the facilities
of the Plant Genetic Resources Centre in Addis Ababa, and in the
long run in a proper equipped Seed Centre of the Forestry Research
Centre, is another necessary approach to conserve forest genetic
resources. For many forest species, long-term storage under lowtemperature conditions can be used. However, much research is still
needed in order to ascertain the best storage conditions for less
orthodox forest tree seeds. Whether the ex situ conservation of tree
pollen or tissue material in a genebank is a practical proposition at the
moment, is questionable. Again, much research will be needed in
order to reveal the appropriate techniques.
Ex situ conservation by species provenance plantations is justified
only for a rather limited number of important commercial or socially
valuable tree species suitable for growing in plantation monocultures
or in agricultural schemes.
Utilization
It is extremely important that the genetic reserves, once
established, are not only preserved, but also utilized. The management must make possible controlled seed collections in bulk and
allow for scientific research or extensive activities. Sometimes the
management should also allow active human interference in the forest ecosystem. From a silvicultural point of view light-demanding,
fast growing species ('colonizers') are often more interesting than the
extremely slow growing, shade-tolerant climax species. Some genetic
reserves should be kept deliberately at a point somewhere in between
the 'pioneer' and climax stages of succession.
Priorities
The implementation of a consistent, long-term programme of
exploration, evaluation, conservation and utilization of forest genetic
resources is a huge task, which requires careful planning, including
the determination of priorities. Priorities must be allocated to:
- the different operations involved;
- different species or groups of species;
- different regions.
Forest genetic resources of Ethiopia
Establishing priorities means that for each species or group of species,
on the basis of a social and commercial importance rating, a decision
is to be taken concerning the urgency of each possible operation. This
urgency is then expressed in the form of a phasing-in time of the
operation. This process can be repeated for each of the different zones
of Ethiopia; thereby it is conceivable to start the implementation of
the programme first in those zones where the pressure on the remaining vegetation is most serious, e.g. the central highlands and north
Ethiopia.
Concerning the priorities of species or the groups of species it is
important to take the following into consideration. Von Breitenbach
(1963) describes 68 families, 158 genera and 326 indigenous species. It
is clear that priorities have to be set since not all these species or all
ecosystems can be conserved at once.
Under the prevailing conditions, the highest priority should be
attached to the group of multi-purpose tree species. Plantation efforts
will be largely directed towards rural communities through the
establishment of small village woodlots or agroforestry systems.
Ethiopia has a large number of indigenous multi-purpose tree species. Tentatively, the following species can be considered to be of
most importance:
Acacia abyssinica
Acacia albida
Acacia mellifera
Acacia nilotica
Acacia senega!
Acacia seyel
Acacia sieberiana
Acacia tortilis
Balanites aegyptiaca
Cordia africana
Croton macrostachys
Erythrina abyssinica
Erythrina brucei
Terminalia brownii
A study is urgently needed in order to make this list more complete
and to decide which species should receive highest priority.
Once the natural distribution ranges of these species have been
determined, systematic seed collections must be made. At the same
time more information has to be collected about their phenology,
biology and reproduction systems. The next step is to identify the
threatened populations and to implement conservation measures. In
situ conservation of the species in their natural ecosystems should
have highest priority, but complementary ex situ conservation activities in areas under less pressure should also be initiated.
In the case of species for industrial use, construction and fuel
purposes as well as for conservation activities, the priorities should be
set according to their threat by genetic erosion.
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Looking at the merchantability for timber processing wood, the
following species have commercial value:
Juniperus procera
Podocarpus gracilior
Aningeria adolfi-friedericii
Apodytes dimidiata
Albizia schimperianalgummifera
Celtis africana (=kraussiana)
Chlorophora excelsa
Cordia abyssinica
Croton macrostachys
Dalbergia melanoxylon
Ekebergia capensis (=rueppelliaria)
Hagenia abyssinica
Linociera giordanii
Olea africana/hochstetteri/welwitschii
Polyscias fulva
Prunus africana (=Pygeum africanum)
Syzygium guineense
All these species can be conserved in situ by establishing reserves
within the coniferous forests, the broadleaved forests and the mixed
forests. Since the (climax) rainforest species probably have a poor
potential for plantation forestry, special ex situ conservation stands or
provenance trials should not receive a very high priority. For the
important species Podocarpus gracilior and Juniperus procera, and possibly also Chlorophora excelsa, additional ex situ conservation plantations and provenance trials are necessary.
In situ conservation of rainforest species, and species of the mixed
coniferous/broadleaved forests, would conserve at the same time a
number of potentially commercial species, i.e. those species which do
not at present have a high market value but might do so in the future.
Some examples of potentially usable hardwoods are:
Allophyllus abyssinicus
Diospyros spp.
Dombeya spp.
Lepidotrichilia volkensii
Mimusops kumtnel
Ocotea kenyensis
Schefflera volkensii
Warburgia ugandensis
There is also a number of exotic species which are already more or
Forest genetic resources of Ethiopia
less extensively used in plantations, and which belong to the groups
of principal use mentioned earlier. A tentative list comprises the
following species:
Acacia decurrens (construction wood)
Acacia mearnsii (construction wood)
Acacia saligna (=cyanophylla) (B)*
Azadirachta indica (multi-purpose tree)
Casuarina equisetifolia (construction wood)
Cupressus lusitanica A
Eucalyptus camaldulensis (construction wood)
Eucalyptus cladocalyx (construction wood)
Eucalyptus globulus (construction wood)
Eucalyptus grandis (construction wood)
Eucalyptus saligna (construction wood)
Grevillea robusta (A/B)
Melia azedarach (multi-purpose tree)
Leucaena leucocephala (multi-purpose tree)
Pinus patula (A)
Pinus radiata (A)
Parkinsonia aculeata (B)
Prosopis juliflora (B)
Prosopis tamarugo (B)
Sesbania spp. (multi-purpose tree)
Schinus molle (B)
The species with an asterisk are especially suited for industrial processing (A), for conservation purposes (B) or for both (A/B).
Since these species have been used for a reasonable period of time
under different conditions, it can be expected that locally adapted
ecotypes have developed. Therefore, systematic collections should be
made and provenance trials need to be set up.
A considerable amount of research with some of these species has
been carried out on a global scale; much can be gained, therefore, if
participation is sought in ongoing selection and improvement programmes which are carried out abroad.
Present organizational structure
The State Forests Conservation and Development Department and the Soil Conservation and Community Forestry Development Department are the two governmental institutions responsible
for the bulk of the country's reforestation programmes. In this work,
both institutions receive support from a specialized Forestry Research
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Centre. This Centre has a Seed Section, established in 1957, which is
responsible for the collection, storage, testing and distribution of high
quality seed of known origin. There is also a Silvicultural Section,
which carries out general silvicultural research related to forest
establishment and management, such as spacing/increment/thinning
trials and species elimination/testing trials. This last category of
research has been and still is the main component of the programme.
Provenance research has received much less attention. Recently,
'wide range' provenance trials with a few indigenous species were
started {Croton, Cordia). For the future, it will be necessary that systematic provenance trials with all socially and commercially valuable
indigenous species are established, by using certain priorities, for
instance, as indicated in the previous paragraph.
The Forestry Research Centre has for practical reasons put
emphasis on applied research, at the evaluation and utilization end of
the range of the operations which are necessary in the selection and
genetic improvement work. Exploration and conservation have been
neglected areas and a considerable increase in efforts in these fields
will be of highest importance.
A possible outline of a set-up in the coordination of exploration
and conservation activities at the national level could be as follows:
- The Forestry Research Centre would be responsible for the
botanical and phenological explorations and collections.
Cooperation with the Inventory Section of the State Forests
Conservation and Development Department and the University of Addis Ababa would be intensified.
- The Forestry Research Centre would indicate which genetic
reserves have to be established and would develop the
management guidelines for these reserves.
- The State Forests Conservation and Development Department would be made responsible for the management of
these in situ reserves; a special unit within this department
might have to be founded for this purpose.
- The Forestry Research Centre would establish and manage ex
situ conservation plantations.
- The Plant Genetic Resources Centre/Ethiopia would take care
of the ex situ conservation of forest tree seeds. In the future an
upgraded Seed Centre might take over this task.
International cooperation
Long-term conservation and evaluation of forest genetic
resources is very expensive and needs specialized skills. Financial
Forest genetic resources of Ethiopia
99
and skilled manpower resources are both scarce in Ethiopia. Because
of this, and since many of the valuable indigenous tree species mentioned also occur in other African countries, international cooperation
is essential.
IUFRO with its specific working parties is one of the main platforms for international cooperation. Ethiopia participated in the
IUFRO Research Planning Workshop for Africa, Sahelian and North
Sudanian Zones held in Nairobi in January 1986. This workshop was
centred around the topic 'selection and genetic improvement of indigenous and exotic multipurpose woody species; including seed collection, handling, storage and exchange'. Other international
cooperation programmes have been established for a range of exotic
species. Coordinating agencies are, for example, the Commonwealth
Forestry Institute, Oxford, for Central American tropical pines (not
yet very important for Ethiopia); the Commonwealth Scientific and
Industrial Research Organization (CSIRO), Canberra, for Australian
Acacias, Casuarinas and Eucalyptus (highly important for Ethiopia);
the Food and Agriculture Organization (FAO), for drought-resistant
Acacias and Prosopis species (important for Ethiopia); the Centre
Technique Forestier Tropical, Nogent-sur-Marne, France, for African
hardwoods and Pacific insular Eucalyptus (less important for Ethiopia); and NAS, Washington, for nitrogen fixing trees and species for
the Sahel zone (relevant for Ethiopia).
The Forestry Research Centre already receives some outside
assistance. In the near future there will be an extension of outside
assistance by inputs through agencies like the World Bank, Finnida,
FAO and SIDA in connection with projects in the field of forestry
which are presently carried out by these agencies. It is strongly
recommended that the project proposals which have to be prepared
contain components in the field of exploration and conservation of
forest genetic resources.
References
Anonymous (1986). Biomass Energy Sources. Final report of a cooperation in
the energy sector between the Ministry of Mines and Energy of the Provisional Military Government of Socialist Ethiopia, ENEC and CESENAnsaldo/Finmeccanica group, Addis Ababa.
Burley, J. & Wood, P. J. (1976). A manual on species and provenance
research with particular reference to the tropics. Tropical Forests Paper No.
10. CFI.
Chaffey, D. R. (1980). South West Ethiopia Forest Inventory Project. A glossary of
vernacular plant names and a field key to the trees. Land Resources Develop-
ment Centre, UK.
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Friis, I., Rasmussen, N. & Vollesen, K. (1982). Studies of the flora and
vegetation of southwest Ethiopia. Opera Botanica, 63, 8-28.
Knapp, R. (1968). Hoehere Vegetationseinheiten von Aethiopien, Somalia,
Natal, Transvaal, Kapland und einige Nachbargebieten. Geobotanische Mitteilungen (Giessen), 56, 1-36.
von Breitenbach, F. (1963). The Indigenous Trees of Ethiopia, 2nd edn. Addis
Ababa: Ethiopian Forestry Association.
Plants as a primary source of drugs
in the traditional health practices of
Ethiopia
DAWIT ABEBE AND ESTIFANOS HAGOS
Introduction
The maintenance of health by means of various techniques
and substances is almost as old as the history of human evolution
itself. Although the resources available were easily drawn from the
natural environment, their efficacy for solving problems which
reduce life expectancy was established only through rigorous trials
over a considerable period of time, often involving Man himself as
subject of the experiment.
Our ancestors, and millions of people in modern Africa, have
relied heavily on plants, animals and minerals to ward off pathogens
and to maintain the functional balance of each organ. Many species of
plants had to be tested and retested in the endless search for drugs
that could prolong life or, it was believed, even confer immortality.
Many plants have been found to possess the desired effects as a result
of well planned experiments, while some discoveries were just a
product of serendipity. Many sacrifices had to be made, however, not
only in terms of money and time, but also in terms of human lives.
Until a generation or two ago plants were the primary source of
health care for entire populations in most African countries and such
plants still remain important sources of drugs for nearly 80 per cent of
the population in contemporary Ethiopia. In spite of this we still
know very little, whilst many Africans who receive their education
abroad believe that imported drugs are always superior, even if these
are unaffordable to the large majority of the population. The investigation of the efficacy and safety of traditional remedies is no doubt a
102
Dawit Abebe & Estifanos Hugos
big challenge; but to turn our backs on this challenge is not only
abandoning our scientific responsibilities but also an indirect
perpetration of those harmful practices which we always bitterly criticize and which we try to eliminate. A substantial body of knowledge
on the practice of traditional medicine and particularly on medicinal
plants has already been lost because most of it was transmitted by
word of mouth from generation to generation. This situation has been
further aggravated by the expansion of modern education, which is
effectively putting an end to family and traditional ties and hence to
the passing on of traditional knowledge. Thus, we now have the last
generation of practitioners within the age brackets of 40 and 80 years.
The ethno-medical information that has been built around numerous
plants is, therefore, on the verge of collapse. The most unfortunate
consequence of this is not only that we shall lose a crucial guide for
potential sources of new drugs against many diseases that either do
not succumb to or show resistance to modern drugs, but also the
actual medical care that still serves a substantial sector of the population in the developing African countries.
In the industrial countries, with the expansion of conventional
medical services, the herbal health care system has become insignificant. However, it would be unrealistic to assume that the same trend
of events is taking place in the developing nations. With their ailing
economies and ever dwindling external aid, it is extremely difficult
for these nations to achieve reasonable modern health care to satisfy
even the most acute demands of their people. The endless civil wars
and international conflicts, and the recurrent drought, often seriously
affect the distribution of national wealth that is intended for the
improvement of health service facilities. Furthermore, the population
in several of these countries doubles every 25 years, consequently
increasing the competition for the already meagre resources of
modern health services. These and other similar circumstances,
therefore, strongly remind us that we have little or no choice except to
develop scientifically our traditional herbal medical practice, to serve
as a partner to the conventional health care delivery system.
The traditional health care system
Unlike its Western counterpart, which essentially concentrates on symptoms of illnesses, traditional health care often considers a healthy existence or its absence as a product of the social and
physical environments and, therefore, the effort is directed not only
towards curing the malfunctioning organ but also to bring the patient
into harmony with these environments.
Plants as a primary source of drugs
103
It is important to note that the choice or even the preference of
traditional herbal remedies is not dictated by socio-cultural reasons
alone. Modern drugs are not only frequently unavailable to most of
the rural people, who often live where there is little or no means of
communication, but they are also beyond their economic reach. Even
the fortunate few who can get hold of the drugs from clinics or rural
drug vending shops have very little or no idea how to use them and
what dosage to use. The obvious outcome of this is serious sideeffects and resistance to many of the drugs (especially to antibiotics)
by the disease organisms or their vectors. As a result of these we see a
growing mistrust and apprehension towards most of the manufactured drugs and an increasing tendency to resort to traditional herbal
preparations as a sole or a complementary source of treatment.
The herbalists are often general practitioners who can handle a
series of health problems, particularly those that commonly occur
within their own areas, thereby effectively minimizing the wastage of
time and money. Of course, the herbalists have their own limits; they
are honest enough to admit this and refer the patient to a more
experienced practitioner or to a modern medical care centre.
In the herbalogical medical care system prescription charges are
often very small and are not necessarily paid instantly in cash but also
in kind or even in a form of manual work over a long period of time.
In addition to house calls and routine care for the outpatient, the
herbalists sometimes manage with up to 10 inpatients who are provided not only with free prescriptions but also with food and shelter
for several days or even years.
Traditional drugs of plant origin are easily available and the patient
seldom needs to wait for days or even indefinitely before the commencement of treatment, as is now commonly the case with some
modern pharmaceutical preparations. The herbalist informs the
patient about his or her sickness, how the drug should be taken and
the precautions that are needed - all in the local language, which is
easily understood by the patient. Since the practitioners are permanent residents within the community, they can perform follow-up
programmes on a daily basis to evaluate the effect of the drugs. If the
medicine administered is slow in its action or non-effective, the practitioner increases the dosage or may prescribe a more potent drug
with faster action. Therefore, the patient is not simply left at the
mercy of modern prescriptions that have to be obtained from clinics
or drug stores located far away, as a result of which further consultations are difficult or even impossible.
In the traditional medical care system, the patient can have a choice
104
Dawit Abebe & Estifanos Hugos
of one or both of the following sources. The first depends on previous
information or experience about the medical virtues of a particular
species of plant, which the patient is able to obtain from the local
markets or from his backyard, to treat less severe cases. Thus anyone
who suffers from influenza, fever, stomach discomfort or minor
injuries often relies on home treatments by using Artemisia afra, A.
abyssinica, Lippia abyssinica, Plantago lanceolata, etc. The second choice
involves either 'general practitioners', who handle more complicated
cases, or 'specialists', who deal with certain diseases such as rabies,
cataract, jaundice, malaria, haemorrhoids, etc. The experts in this
category, comprising the herbalists, bone setters, traditional birth
attendants and faith healers, acquire their theoretical and practical
knowledge during in-service training of up to 30 years with one of
their parents or teachers who established their own reputations by
undergoing similar apprenticeships. The theoretical aspect of traditional medicine comprises a belief about the etiology and syndromes
of diseases, diagnoses and the drugs employed to cure them. Mythological concepts are sometimes also included and naturalistic
approaches alone are regarded as incomplete, especially in disorders
such as epilepsy, insanity, hysteria, etc. The practical part of the
system consists of all the necessary steps that must be taken to cure or
prevent diseases.
Source of remedies and methods of treatment
The traditional pharmaceutical preparations are of mineral,
animal and vegetable origin. Widely used minerals for external applications include sulphur, copper sulphate, sodium chloride and
thermal spring waters. Drugs of animal origin are mostly obtained
from Coleoptera, chameleon, python, rock rabbit, hyena, greater
kudu and elephant. Ninety-five per cent or more of the traditional
medical recipes, however, are of plant origin (Table 1). Depending on
the availability of materials and their therapeutic actions, fresh or
dried plants may be used (Abebe, 1986). High-altitude specimens are
thought to be milder and slower in their action than comparable
plants from the lowlands. The season and the time of day of plant
collecting are sometimes considered as important factors in determining potency. Thus fruiting plants collected during the dry season are
believed to be more effective than those obtained during rainy
periods. Similarly, dawn collections are preferred to those carried out
later in the day (Abebe, 1987). In general, no specific kinds of collecting tools are required, even though some herbalists believe that
Table 1. Plants used in Ethiopian traditional medicine
Scientific name
Vernacular
name
Uses*
Parts of
plant used
Achyranthes aspera
telenji
ei, me, tn, wo
Leaf
Ajuga remota
anamero
insilal
Leaf
Leaf, seed
Argemone mexicana
madafe
ar, di, ja, ma, rw
ax, bh, du, go,
ka
en, co
Asparagus africanus
sereti
ap, di, go, lc, vo
Leaf, root
Bersama abyssinica
azamir
ar, ha, pw
Root, bark, leaf,
seed
Brassica nigra
senafichi
fl, id
Root
Brucea antidysenterica
abalo
cq, ec, ga, ha,
kd, ku, mi, rp
Seed, leaf
Calotropis procera
tobiawu
digita
am, ha
ad, di, ec, go,
ha, hy, is, ja, ve,
wo
Seed
Seed, leaf
Anethum
graveolens
Calpurnia aurea
Seed
Form of
application
Active substances
Saponin, alkaloid, sterol,
fat, KC1
Leaf juice
Leaf juice, seed
powder
Oil
Powder mixed
with honey
Powder
Fresh root eaten
with salt
Oil of seed,
powder of leaf
Powder
Powder
Limonene, D-carvone
Argemone oil, toxic
alkaloids, sanguinarine,
berberine and protopine
Leaf and root contain
insecticide; mortal to cows
in Uganda
Fixed oil, glucoside
sinigrin
Bruceantine, fixed oil,
volatile acids, bitter
principle, resin,
phytosterol
Calpurnin, tannin
Table 1 (cont.)
Scientific name
Vernacular
name
Uses*
Parts of
plant used
Form of
application
Carissa edulis
agam
by, ep, ia
Root
Powder
Capparis tomentosa
he, Id, Ik, px, so, Root
ta
dt, pb
Leaf
Powder
Cassia occidentalis
arangama
guracha
arda bofa
Clematis sinensis
azo-hareg, fiti
Clerodendrum alatum
Croton macrostachys
misirich,
marasisa
misirich,
marasisa
bisana
cc, ec, el, ha, he, Root, leaf
le, Is, nk, sr, ta
am, as, go, kd, Root, leaf, bark
sm
ap, ep, md
Root, leaf
Cucumis aculeatus
yemdirimbway
C. dipsaceus
anchote
aserkush
tebetebkush
C. myricoides
Cyphostemma niveum
ar, cq, el, ep, fu, Leaf, seed
he, he, ja, kd,
ku, le, sf, wo
di, go, le, mg,
Root
sn, sv, wo
go, nk
Root
it, nk, si, sn, sw, Root, bark, leaf
wo
Powder
Active substances
Root contains cardiac
glycoside carissin
Sulphur oil, stachydrine,
alkaloid
Oxymethyl
anthraquinone, tannin,
fatty substance, gum,
glucose, mucilage,
albumin, emodin, fixed oil
Powder
Powder
Smoke inhaled
Powder
Crotin, crotepoxide resin
Powder
Cucumine-like principle
Juice
Powder
Saponin
Datura stramonium
astenagir
am, da, ec, ep,
es, he, hy, ko,
lh, mi, nb, pk,
so, sq, ta
Root, leaf, seed
Powder
Dracaena steudneri
Erythrina brucei
etse patos
korch
as, ra
ab, hf, it, ja
Bark
Leaf, seed
Heteromorpha trifoliata
Hypericum quartinianum
Jatropha curcas
jib merkuz
amija
habatalumuluk
ja, ra
Root, leaf
Leaf
Seed
Powder
Leaf juice,
powder of seed
Powder
Juice
Powder
Kalanchoe lanceolata
endahula
ar, bs, di, sw, vo Leaf
Pulverized fresh
leaves
K. marmorata
Lepidium sativum
titu
feto
ar, ad, ht, ko, sk All parts
Seed, leaf
et
Powder
Decoction
Lagenaria siceraria
kil
ar, er, ha, md
Root, leaf, fruit,
seed
Powder
Lawsonia inermis
henna
Leaf
Decoction
Momordica foetida
buke seytana
a
j
ar, db, go, he,
hy, me, sa, sr
ap, co, el, ga, kr, Root, leaf
rp, sa, sf
Sap and/or
powder
Daturine, hyoscyamine,
atropine, scopolamine,
solanine, saponin,
carotene, Vitamin C, malic
acid, tomatidine
Saponins, resin
Volatile oil
Fixed oil, phytosterolin,
curcin, tannin, steroidal
sapogenin, hydrocyanic
acid
Cress oil, iodine, uric acid,
fixed oil, sanapin, benzylisothiocyanate
Fruit contains niacin,
riboflavin, thiamine,
amygdaline; seeds contain
saponin andfixedoil
Acid, resin, fats, a yellow
pigment, lawson volatile
oil, fixed oil, brown oil
Momordicin
Table 1 (cont.)
Scientific name
Vernacular
name
Uses*
Parts of
plant used
Form of
application
Myrsine africana
kechemo
ah, ar
Fruit
Powder
Ocimum suave
anchabi
Osyris compressa
keret
ar, as, cc, ed, he, Leaf
ra, sf, sx
Leaf
ga, ja
Powder
Phytolacca dodecandra
endod
Plumbago
zeylanicum
amera
Polygonum
barbatum
Portulaca oleracea
gumamila
kentela
Rumex bequaertii
tult
R. nervosus
embuacho
ad,
go,
ay,
ha,
rp,
wo
ah,
ao,
he,
ar, da, ec,
ja, ra
eg, ec, go,
ku, lb, nk,
sr, sw, sy,
sr
du, fe, ga,
kd, ve
ae, an, ar, as,
bd, co, di, fe,
go, ha, hm, Id,
me, ox, px, ra,
sw, vo, wo
ae, ar, it
Active substances
Embelin, cuercitol,
anmyrsine-saponin
Sap or decoction Eugenol
Root, leaf, fruit
Powder
Bark, root, leaf
Powder
Leaf
Whole plant
Sap or powder
Juice
Root, leaf
Powder
Root, stem, leaf
Infusion
Tannin, sandalwood oil,
sesquiterpene, glucoside
osyritin
Tannin, saponin, alkaloid,
volatile oil
A toxic principle,
plumbagin, fixed oil,
volatile oil
Hydrocyanic acid
Cyanic acid, alkaloid,
saponins, oxalic acid,
Vitamin C, potassium
salts, fixed oil, volatile oil
Securidaca
longipedunculata
etse menhae
bu, ep, go, ne,
le, ma, rp
Spilanthes mauritiana
yemeder
berbere
am, co, It, sa,
Whole plai
sh, sz, ta, tn, vo
Stephania abyssinica
kalala
ar, ch, di, fi
Syzygium guineense
dokma
as, bb, di, sr, wo Bark
Tamarindus indica
roka
ad, ap, ar, bp,
en, hy, sw, ve
Thalictrum
rhynchocarpum
sir bizu
ap, as, co, le, sg, Root
sx
Verbascum sinaiticum
ketetina
ad, ae, ap, as, cl, Root, leaf
di, du, ec, ep,
hf, it, ja, kd, kw,
Id, me, nk, sm,
st, sx, sy, vo
ae, as, di, pa, sa, Root, leaf
ve, vo
Root
Root, leaf
Fruit, seed
Methyl salicylate,
saponin, tannin, steroid,
glucoside
Volatile oil which is toxic
Juice
to fish, acid amide,
spilanthol, sterol
Powder or fresh Morphine
pulverized
Leaf yields 27.5% cellulose
Powder
and 8.4% albumin
Tannin; tartaric, acetic,
Infusion
citric, malic and succinic
acids, sugars, pectin,
Vitamins A, B, B2, C,
active carotenoid
Thalicarpine, which
Powder
shows anti-tumour
activity, has been isolated
Powder
from T. dasycarpum
Verbena officinalis
atuch
Vernonia amygdalina
girawa, ebicha
pa, tn
Stem, leaf
Juice or powder
Ganglion-blocking
alkaloids were isolated
from V. nobile
Juice or powder
Juice
Verbenaline, mucilage,
tannin, essence, bitter
substance
Vernodalin, vernomygdin
Table 1 (cont.)
Scientific name
Vernacular
name
V. hymnolepis
murukruk
Warburgia ugandensis
bifti
Withania somenifera
gizawa
Zehneria scabra
hareg resa
a
Parts of
plant used
Form of
application
ae, am, hf, ka,
sn, ve
co, el, ep, ra, rp,
sx
by, ep, he, md,
me, pd, sa, sq,
su, sx
Root
Powder
Bark
Powder
Tannin, mannitol, resin
Bark
Powder
Scopolamine, withaferin,
somniferin, Vitamin C,
tannin, fatty acids,
pungent volatile oil
di, fe, ma, mn,
mt, sy
Root, leaf
Juice or
decoction
Uses*
Active substances
ab, abortifacient; ad, amoebic dysentery; ae, anti-emetic; ah, anthelmintic; am, anti-asthmatic; ao, abdominal complaints; ap,
aphrodisiac; ar, against Ascaris; as, anti-spasmodic; au, against anus prolapse; ax, against anorexia; ay, vitilago; bb, broken bone; be,
breast cancer; bd, boil dressing; bh, bilharzia; bp, bile problem; bu, burns; by, against evil eye; cc, common cold; eg, anti-chapp; cl,
chill; en, against constipation; co, cough; cq, Tinea corporis; ct, cataract; di, diarrhoea; dt, acute febrile illness; du, diuretic; ec,
eczema; ed, eye disease; ei, epistaxis; el, elephantiasis; ep, epilepsy; es, ear pus; et, against emaciation; fe, fever; fi, fungal infection
of face; f 1, flatulence; fu, fire burn; ga, gastritis; go, gonorrhoea; ha, haemorrhoids; he, habitual miscarriage; he, headache; hf, heart
failure; hm, haemostatic; hp, heart pain; hy, hypotensive; ia, insanity; id, indigestion; is, insecticide; it, insect repellent; ja, jaundice;
ka, heartburn; kd, kidney disease; ko, favus; kr, gastroenteritis; ku, against Tinea versicolor; kw, kwashiorkor; lb, lung tuberculosis;
lc, lactogogum; Id, liver disease; le, leprosy; lh, alopecia; Ik, hysteria; Is, leishmaniasis; It, loose teeth; ma, malaria; md, madness;
me, menorrhagia; mg, migraine; mh, pneumonia; mi, mental illness; mt, chloasma; nb, numbness; nk, tuberculosis and/or cancer;
ox, oxytoxic; pa, poison antidote; pb, poisonous reptile bite; pd, prevention of epidemics; pk, pain-killer; pw, pin worm expellent;
px, placenta expeller; ra, rabies; rp, rheumatic pain; rw, roundworm expeller; sa, stomach-ache; sc, scabies; sf, stomach distention;
sg, spinning head; sh, sore throat; sk, swollen breast; si, swollen scrotum and penis; sm, stomach trouble; sn, snake bite; so,
aphasia; sq, stomach burn; sw, swelling; sx, spasm; sy, syphilis; sz, scorpion bite; ta, tooth-ache; tm, tumour; tn, tonsillitis; up,
uterine prolapse; ve, vermifuge; vo, vomiting; wa, warts; wo, wound.
Plants as a primary source of drugs
111
using a horn-handled knife or olive sticks is more likely to increase
the strength of the drug. Prayers and even sacrifices may be considered as important conditions, not only to enhance the therapeutic
effect but also to remove the poisonous effect of the plant.
The name of a medicinal plant in Ethiopia sometimes takes the
name of the disease or of its operative agent. Two or more allied
species with overlapping distributions are sometimes given the same
name and may be used interchangeably. Among herbalists with a
church education plant species often are given vernacular names
composed of two words. The first is applicable to all species regardless of their affinities or differences. The second is a specific epithet
which usually depicts the characteristic of the taxon when employed
as a drug. Thus, 'etse sioul', which literally means 'plant of hell',
indicating the burning effect or sensation that is produced by applying Ranunculus multifidus. Similarly, 'etse yihayu' means 'restorative
plant', describing the effectiveness of Habenaria spp. in overcoming
impotence.
Depending on its size and its therapeutic action, the whole plant,
or different parts of it, is prepared in powder forms, infusions, decoctions, etc., to be employed in the treatment of a variety of diseases.
The dosages of the preparations are often measured in a glass for
liquids, a pinch or teaspoon for powders or a handful for seeds, roots,
leaves, etc. The patient's age, sex, physical condition and stage of the
illness are the major factors which determine the type of remedy and
the dosages to be prescribed. The formulations prepared from different parts of the same plant may be employed for diseases with
different symptoms or could even be used to produce opposite effects
(Abebe, 1987). The fruit of Ficus vasta, for example, is claimed to have
a laxative property, while its root is believed to stop diarrhoea. Different or similar parts of up to 12 species may sometimes be mixed in
a given proportion to treat diseases with either clear-cut symptoms or
those which manifest mixtures of syndromes. Synergistic and/or
antagonistic effects of the various constituents are, therefore, well
recognized by the herbalists.
Treatment with herbs does not always have to follow the normal
procedures or administrative routes; it may also be applied by, for
instance, just tapping the forehead a few times with a fresh stem of
Malva verticillata to stop epistaxis (nose bleeding), or simply cutting
the stem of Rutnex bequaertii and simultaneously calling the name of
the patient who suffers from excessive menstruation. In the traditional health care system preventive and prophylactic treatments are
112
Dawit Abebe & Estifanos Hugos
also very well known. Preventive remedies against epidemic
diseases, snake and mosquito bites may be carried out by an
individual person, or the plant may be grown around the house to
protect the whole family. Although not common, prophylactic treatments are employed against rabies, malaria and even tapeworms.
Rejuvenants and restorative drugs of plant origin are also known to
counter the effect of ageing and to overcome signs of malnourishment, infertility, amenorrhoea, etc. (Abebe, 1986). Certain plants are
also claimed to have the ability to boost the memory or intellectual
power of teenagers. In fact, there seems to be very little that cannot be
influenced by the application of plant materials, be it to attract the
opposite sex, stop the rain, prevent attack by an enemy or beast, etc.,
although many of these claims have yet to be scientifically tested.
The fields for which plants are employed by the herbalists of Ethiopia are as varied as the species themselves. Among these the most
important and with high potential for future application are the traditionally claimed drugs for human and veterinary health, insecticides,
herbicides and water clarifiers. An insight into traditional medicine
will also bring to light more of the abortifacients, teratogens,
allergenics, hallucinogenics and other toxic plants which are of enormous significance to the health workers.
Concluding remarks
In the traditional medical system, the knowledge of the
plants employed in the cure and prevention of disease is based on
repeated observations and is passed on from one generation to the
next. As a result all those plants or plant parts that have adverse and
serious side-effects are well recognized by the herbalists and are
eliminated from the list of therapeutic agents. Even if their use is
justified, they are given to the patient under strict supervision and
with the antidotes ready to counter their potential harmful effects.
Behind the facade of methods based on superstition, the traditional
healing procedures more often embody rational principles and effective drugs against the major diseases afflicting the large sector of
society. Exaggerating its weak points out of all proportion could
never change the objective situation or the attitudes of over 80 per
cent of the population, that considers the traditional medical practice
to be its vital health care system.
Given their rapid rate of population growth and their weak economic position, many Third World countries seem to have little or no
choice except to develop their traditional medical systems scientifi-
Plants as a primary source of drugs
113
cally, in order to achieve maximum health coverage. With the right
research approaches, effective, safe and cheap drugs of plant origin
will no doubt be relatively easily established as substitutes for
imported and often expensive modern medicines. The possibility of
discovering superior and even completely new therapeutic agents
against the diseases that are less or not at all amenable to existing
pharmaceutical preparations is also very high. Therefore, if we are to
fulfil our immediate objective of maximizing health care coverage and
contributing to the ceaseless worldwide scientific effort directed to
the discovery of new drugs, an open mind towards traditional medicine must certainly be maintained.
References
Abebe, D. (1986). Traditional medicine in Ethiopia: The attempts being made
to promote it for effective and better utilization. Unpublished report,
Coordinating Office for Traditional Medicine, Addis Ababa.
Abebe, D. (1987). Plants in the health care delivery system of Africa. Proceedings of the 14th International Botanical Congress, 24 July-5 August 1987, Berlin.
Bannerman, R. H., Burton, J. & Wen-Chen, C. (eds) (1983). Traditional Medicine and Health Care Coverage. World Health Organization, Geneva.
Traditional aromatic and perfume
plants in central Ethiopia (a botanical
and ethno-historical survey)
E. GOETTSCH
Introduction
The amazing variety of incense, perfumes and other aromatic
materials gained our interest and attention when we collected spices
in the marketplaces of Addis Ababa and its surroundings.
Aside from incense and myrrh very little is generally found in
research literature about the use of plants as perfumes and aromatics
in Ethiopia. In this paper those plants and plant products will be
treated, which were found in markets in central Shewa, the
administrative region around Addis Ababa and in the capital itself.
Some plant products have also been reported from the Bale
administrative region.
The subject will be treated in three sections: the first will deal with
incense and myrrh and will consider their importance in international
trade since ancient times. The other two sections will cover aromatic
plant materials of different uses and perfume plants, respectively.
Incense and myrrh
From time immemorial the fragrant smoke of burning resins
and the aromatic odours of ointments and balms have been used by
Man in religious rituals.
In the ancient Mediterranean civilizations incense (or frankincense,
as it is also called) and myrrh were considered, at times to be more
precious than gold (Gauckler, 1970). The importance of incense in
those days is documented by the fact that the first great trade route in
history is called the 'incense road', covering a distance of about
Traditional aromatic and perfume plants
115
5000 km from the kingdoms of southern Arabia ('Arabia Felix') to the
cultural centres to the east of the Mediterranean Sea. This trade was
already flourishing by about 2000 BC. Both products are mentioned in
the Old Testament and were introduced into church ceremonies at
the beginning of Christianity (Abercrombie, 1985).
In Ethiopia the use of incense and myrrh for ritual purposes goes
back at least to the Aksumite Empire, ca. 500 BC (Goldschmidt, 1970)
and has ever since been continued by the Orthodox Church where it
is still very popular.
In recent years, however, the use of incense in church ceremonies
has considerably decreased worldwide; but a growing amount is
needed in the industrial sector, e.g. in pharmaceutics and cosmetics.
The world markets for incense and myrrh are dominated today by
South Yemen, Ethiopia and Somalia. There is a specially strong
demand for southern Arabian frankincense because of its superior
quality. The best material since ancient times is produced in the
Dhofar province of Oman, where soil and climatic conditions are
ideal (Zohary, 1983; Abercrombie, 1985).
In Ethiopia, trade in these goods is handled by the Ethiopian Forest
and Wildlife Products Processing and Marketing Corporation. The
annual production is estimated to be well over 30000 tonnes (Ahmed
Taib, 1982), most of which is consumed locally. In 1983-4 3300 tonnes
were exported mainly to Western Europe, the Middle East and China.
In commerce myrrh is sold under the trade names gum oppoponax
and gum myrrha, and incense as gum olibanum. From the viewpoint
of strict scientific definition these so-called gums are in fact resins.
The term resin is not easy to define in a precise manner, for natural
resins differ greatly among themselves. They have certain properties
in common, however, which make them easily recognizable: resins
are insoluble in water but dissolve readily in alcohol, ether, carbon
bisulphide and certain other solvents. When heated they first soften
and then melt to a more or less clear, sticky fluid. They burn with a
smoky flame, are resistant to most natural reagents and they do not
decay (Howes, 1949).
Myrrh
Myrrh is a natural exudate of trees of the genus Commiphora.
In Ethiopia about 48 species of Commiphora can be found. The species
used for the production of myrrh are C. myrrha (Nees) Engl., C.
africana (A. Rich.) Engl., C. erythraea (Ehrenb.) Engl., C. gileadensis (L)
C. Chr., C. abyssinica (Berg) Engl., C. hodai Sprague, C. kua (R. Br. ex
116
E. Goettsch
Royle) Vollesen, C. quadricincta Schweinf., C. schimperi (Berg.) Engl.
and C. truncata Engl. (Vollesen, 1989). Except for C. gileadensis, which
occurs only in Eritrea and the Harerge region below an altitude of
750 m above sea level, the rest of the Commiphora species mentioned
are relatively widespread in Ethiopia, occurring in the Eritrea, Tigray,
Gojam, Gondar, Welo, Shewa, Arsi, Sidamo, Harerge, Bale and
Gamo Gofa regions up to an altitude of 2000 m above sea level,
although they are also quite common in the lowlands below 100 m.
The resins yielded by these species differ from one another in taste
and odour. True myrrh is produced by C. gileadensis and C. abyssinica.
Other types of myrrh of different composition are known by their
traditional trade names 'bissabol' (C. erythraea), 'harabol' or 'perfumed bdellium' (C. myrrha) and 'bdellium' (C. africana) (Uphof,
1968).
Myrrh ('kerbe' in Amharic), has a powerful scent when it is burnt.
The traditional use of 'kerbe' as an incense is not very popular
because of this strong scent, and also because of the fact that myrrh is
associated with witchcraft. On the other hand, however, prostitutes
in Addis Ababa are said to attract visitors by burning 'kerbe' in front
of their houses.
Some myrrhs are used by traditional healers as universal remedies.
The oil of myrrh has a rich odour and is used as a balm for ritual
ceremonies, as a disinfecting ointment and as a perfume (Gauckler,
1970; Zohary, 1983).
Incense
Gum olibanum or true frankincense is an oleo-gum-resin
obtained from trees of the genus Boswellia by tapping them. So far six
Boswellia species have been reported to occur in Ethiopia. The two
most common species are B. papyrifera (Del.) Hochst. and B. rivae
Engl. (Cufodontis, 1953-72; von Breitenbach, 1963; Atkins, 1964;
Werner, 1974; Ahmed Taib, 1982; Vollesen, 1989). B. papyrifera is the
most common species, known in Amharic as 'itan zaf (incense tree).
It is found in the lowland areas of Gojam, Shewa, Gondar, Tigray and
Eritrea (up to 1800 m), whereas B. rivae is found between 250 and
800 m in Sidamo and Harerge regions (Maslekar, 1975; Vollesen,
1989). In Konso (Gamo Gofa region) B. rivae has been found by the
author up to an altitude of 1050 m.
There is a great demand for frankincense in the local markets since
large quantities are used in church ceremonies and it is also burnt in
private houses during the coffee ceremony. In certain areas magicians
use it in their rituals.
Traditional aromatic and perfume plants
117
The traditional grading of incense is done by referring to (a) the
colour, (b) the origin or (c) the use of the resin. The following grading
system was and still is used by various merchants in the marketplaces
of Addis Ababa, although it was not possible to determine the species
from which the particular incense derives:
1. nech itan
white incense (best quality)
tikur itan
black incense (said to be produced by old trees)
kai itan
red incense (inferior in quality, contains pieces
of bark)
2. Tigray itan
Ogaden itan
Asmara itan
Bahar itan
used as incense
used as incense and for perfumery
black incense
imported from outside Ethiopia (e.g. from
Aden)
3. set itan
mitan itan
'Ladies' incense'; dresses are dried in its smoke
incense mixture, containing different aromatic
materials and serving as a cheap substitute for
frankincense
Incense also provides the raw material for some manufactured
aromatics, and Ethiopian Muslims, for example, who are influenced
by Arabian culture, have a preference for these. There are two
popular substances of this kind:
1. 'Libanja', the most important incense in this group, is
imported from South Arabia, Djibouti, Somalia and perhaps
Kenya. This substance has a mineral-like appearance. It contains mainly a refined incense, but it was not possible to
obtain any further information on the other ingredients.
2. 'Misketi', a strong aromatic mixture, is produced in Dire
Dawa (Harerge region) and preferred especially by Muslims.
It is made of 'Libanja', sugar, powder of sandalwood (from
India) and 'Miski', a cheap perfume. When burnt 'Misketi'
produces a strong sweet scent; it is mainly used during coffee
or chat (Catha edulis Forssk.) ceremonies.
Other aromatic plant materials
(a) 'Birgud'
Another important group of aromatic plant materials to be
found in Ethiopian marketplaces is called 'birgud' (Amharic).
It was not possible to obtain any clear information on the nature of
'birgud'; the name applies to both a dark resin and a woody bark.
Both materials give a pleasant smell when burnt.
118
E. Goettsch
According to Wolde-Michael (1980) 'birgud' is the Amharic name
for Cinnamomum cassia Blume, which grows in south-east China. Its
bark was already used as an incense in ancient rituals, which are, for
example, repeatedly described in the Bible (Zohary, 1983). In spite of
the fact that 'birgud' bark is quite popular in Ethiopia, there is no
evidence that this material is produced in Ethiopia. It is probably
imported, but the question of its origin remains open. Types of
'birgud' found in the marketplaces are:
1. Small pieces of a crumbling woody bark: not yet identified.
2. A dark resin: the cheaper type of this resin is called 'Chigga
birgud', whereas the other type is one of the most expensive
natural aromatics available.
3. 'Arussi birgud', pieces of a woody bark from Arussi. Regarding appearance and scent it might be the bark of Juniperus
procera Hochst. ex Endl.
(b) Other resins
Other kinds of resin used as an incense include:
1. 'Wunsi' (Amh.): a black resin, not yet identified but because
of its scent it might be produced by Juniperus procera Hochst.
ex Endl.
2. 'Hunsi' (Amh.) or 'Ancha' (Orom.): bought at Goro-market
(Bale); not yet identified. The resin resembles very much that
of Boswellia. According to a local healer the smoke has medicinal properties.
(c) Plants producing scent when burnt
In Ethiopia a number of different plants are known which
give a pleasant odour when put into the fire. Some of these plants are
listed in Table 1.
Perfume plants
Perfume plants contain different types of essential oils. Either
the whole plant is used as a perfume or the essential oil is extracted
from it. Many species of perfume plants have been introduced to
Ethiopia where they are now grown successfully (e.g. lavender,
Geranium spp., mimosa, etc.). In this section only the traditional perfume plants will be listed.
In many parts of Ethiopia it is a traditional fashion to butter the
hair. This habit is mainly restricted to women, but men may also be
accustomed to do so (e.g. Karayu and Afar). In order to overcome the
often rancid odour of the butter, perfume plants are mixed with it.
Table 1. List of aromatic plants which are burnt to produce a pleasant smell
Amharic name
Scientific name
Use and description
1. Karbaricho
Echinops spp.
2. Afer kocher
syn. Nech krinfud
Hedychium spicatum
The root is burnt; the smoke is said to drive out evil spirits and
vermin. Today mainly used because of its pleasant smell.
Tree originating from eastern India growing in Ethiopia. Sliced
roots burnt during coffee ceremony; clothes are dried in the
smoke; expensive.
Mixture of sandalwood powder and 'afer kocher'. The
sandalwood could be imported but could also be produced by
the East African sandalwood {Santalum album L.), a tree that
grows in some areas of eastern Ethiopia. The use is the same as
that of pure 'afer kocher', but this mixture is much cheaper.
Wood, containing an essential oil, pleasant smell when burnt.
Gives pleasant smell when burnt.
3. Bukbuka
4. Ye-Jima inchet
5. Ye-Aden chiraro
Unidentified; 'wood from Jima'
Unidentified; 'dry twigs from
Aden'
6. Weyra
Oka europaea subsp. africana
7. Semat
Unidentified
8. Chiz inchet
'smoke-producing wood'
9. Tinjut
Otostegia integrifolia or O.
steudneri
10. Mitin chito
'Perfume mixture'
Scented when burnt. The pleasant-smelling smoke is led into
containers for milk, home-made beer and yoghurt.
Large tree; bark contains a milky sap. Found in Abbai and
Takazze Gorge; thin strips of bark are plaited into a strand
which is burnt like an incense stick.
Collective term for mixtures of aromatic woods and perfume
plants. The smoke gives a pleasant smell and - depending on
the ingredients - evil spirits can also be driven out. Normally
these mixtures are very cheap. Ingredients can be: Karbaricho,
Kuni, Gizawa, or Weyra, Ades, Birgud and Itan.
Small herbaceous plant. Dried leaves are burnt in containers
for local beer and milk; gnats are expelled by the smoke.
Simply a mixture of wood powder, a little oil and cheap
perfumes. Gives strong odour when burnt. Preferred by
Muslims and Gurage.
Table 2. List of traditional Ethiopian perfume plants
Amharic name
Scientific name
Use and description
1. Koseret or Azkuti
Ocimum spp.
At least five species of Ocimum are found in Ethiopia, O.
basilicum L. being the best known ('basobila'). Some Ocimum
spp. contain an important essential oil which allows them to be
used as perfumes.
Fresh plants are spread on the floor of the house. Also used to
scent butter.
Probably indigenous to Ethiopia; used as an incense, also
expels gnats.
The plant is mostly sold fresh; the crushed leaves of Artemisia
afra Jacq. ex Wild, are used as a perfume. Dried leaves are put
between cloth. Ariti is burnt for its aromatic smoke; fresh
plants are spread on the floor of houses.
The plant contains an essential oil which is of some economic
importance (e.g. used in cheap perfumes, insecticides, etc.).
Traditionally the plant is burnt for its pleasant scent. The fresh
plant is spread on the floor. Tej sar7 is also used as a medicine
and as a flavouring agent.
The roots of this plant are ground and mixed with butter to
improve its smell. 'Kuni' is a cheap perfume in the highlands.
The plant contains myrtle oil. Leaves are ground and mixed
with butter which is put into the hair by traditional women.
Tungug' is probably made of a grass; it has the strong and
pleasant smell of fresh hay. The plant is mixed with butter,
which is rubbed into the hair.
Kasse
Ocimum ladiense
Kasse
Ocimum sacrum
2. Ariti (Tikur, nech)
Artemisia rehan
3. Tej sar
Cymbopogon citratus (Lemon
grass)
4. Kuni
Cyperus bulbosus
5. Ades
Myrtus communis
6. Tungug
Unidentified
Traditional aromatic and perfume plants
121
Most of the plants mentioned here are exclusively, or at least to a
certain extent, used to scent butter (e.g. Koseret, Kasse, Ariti,
Tungug, Ades, Kurd). The perfume plants found in the marketplaces
in Addis Ababa are listed in Table 2.
Acknowledgements
I wish to express my thanks to Ato Wolde Michael Kelecha,
formerly associated with the Forestry and Wildlife Development
Authority. Without his extensive materials the sections on incense
and myrrh could not have been written.
References
Abercrombie, Th. J. (1985). Arabia's frankincense trail. National Geographic
Magazine, 168, 474-512.
Ahmed Taib (1982). The Swiss, Italian and Finnish Markets for Ethiopian Gum
Olibanum. Programme for Development Cooperation, Market Research
Report No. 5, The Helsinki School of Economics.
Atkins, W. S. (1964). The Future of the Natural Resin Industry in Ethiopia. A
report for the Technical Agency of the Imperial Ethiopian Government,
Addis Ababa.
Cufodontis, G. (1953-72). Enumeratio Plantarum Aethiopiae, Spermatophyta.
Bulletin du Jardin Botanique National de Belgique, Bruxelles.
Gauckler, K. (1970). Die kostbarsten Drogen der Alten Welt: Weihrauch,
Myrrhe, Balsam. In: M. Lindner (ed.), Petra und das Koenigreich der
Nabataeer. Abhandlungen der Naturhistorischen Gesellschaft, Nuernberg.
Goettsch, E. (1985). Aromatic and perfume plants in Central Ethiopia. PGRC/EILCA Germplasm Newsletter, 8, 11-16.
Goldschmidt, C. (1970). Die Weihrauchstrasse: Zur Geschichte des aeltesten
Welthandelsweges. In: M. Lindner (ed.), Petra und das Koenigreich der
Nabataeer. Abhandlungen der Naturhistorischen Gesellschaft, Nuernberg.
Howes, F. N. (1949). Vegetable Gums and Resins. Chronica Botanica Company,
Waltham, Massachusetts.
Jansen, P. C. (1981). Spices, Condiments and Medicinal Plants in Ethiopia, their
Taxonomy and Agricultural Significance. PUDOC, Wageningen.
Maslekar, A. R. (1975). A report on rapid aerial survey for Boswellia papyrifera
(incense tree), Tigrai province. Addis Ababa (mimeographed).
Uphof, J. C.T. (1968). Dictionary of Economic Plants, 2nd edn. Verlag von J.
Cramer, Lehre.
von Maydell, H.J. (1981). Baum- und Straucharten der Sahelzone unter
besonderer Berucksichtigung ihrer Nutzungsmoglichkeiten. GTZ, Eschborn (mimeographed).
Vollesen, K. (1989). 123. Burseraceae. In: I. Hedberg and S. Edwards (eds),
Flora of Ethiopia, vol. 3. The National Herbarium, Addis Ababa University,
Ethiopia, and the Department of Systematic Botany, Uppsala University,
Sweden, pp. 442-78.
von Breitenbach, F. (1963). The Indigenous Trees of Ethiopia, 2nd edn. Ethiopian
Forestry Association, Addis Ababa.
Werner, F. (1974). Memorandum on the collection of incense within Ethiopia, Addis Ababa (mimeographed).
122
E. Goettsch
Westphal, E. (1975). Agricultural Systems in Ethiopia. PUDOC, Wageningen.
Wolde-Michael 'Kelecha (1980). A Glossary of Ethiopian Plant Names, 3rd edn.
Addis Ababa (mimeographed).
Zohary, M. (1983). Pflanzen der Bibel. Calwer Verlag, Stuttgart.
8
Spice germplasm in Ethiopia
E. GOETTSCH
Introduction
Although spices are considered as minor crops their significance for Ethiopia can hardly be overestimated. Spices are needed
every day in considerable amounts for the preparation of the main
dish of the day.
Most of the spices needed in Ethiopia are grown as field or garden
crops, although some grow in the wild. Classical spices are also used
but have to be imported, mainly from India. The following 12 spices,
which originated in Ethiopia or were introduced very long ago and
are considered to be of importance, are dealt with in this chapter:
1. Capsicum annuum (red pepper); Amh.: berbere
2. Trigonella foenum-graecum (fenugreek); Amh.: abish
3. Nigella sativa (black cumin); Amh.: tikur azmud
4. Trachyspermum ammi (Ethiopian caraway); Amh.: nech azmud
5. Coriandrum sativum (coriander); Amh.: dimbilal
6. Aframomum korarima (false cardamom); Amh.: korarima
7. Cuminum cyminum (cumin); Amh.: kamun
8. Foeniculum vulgare (fennel)
Pimpinella anisum (anise); Amh. for both: insilal
9. Ruta chalepensis (rue); Amh.: tena-addam
10. Ocimum basilicum (basil); Amh.: basobila
11. Piper longum (Indian long pepper); Amh.: timiz
12. Rhamnus prinoides (buckthorn); Amh.: gesho
Although 'gesho' is not a typical spice, it is included in this list, since
it is of extreme importance in the flavouring of beverages during their
preparation (Jansen, 1981).
In a broader sense, shallots (Allium cepa) and garlic (A. sativum) can
be considered as spices. They were introduced very long ago and
124
E. Goettsch
recently genetic erosion has started in areas where improved varieties
are coming into use. Nevertheless, since the two species can be
regarded as both spice and vegetable, the latter use being the more
important, they are not treated here.
Also not included in this list are Lepidium sativum (garden cress,
feto) and Tamarindus indica (tamarind); both are considered by Westphal (1975) as spices but they are primarily medicinal plants.
Zingiber officinale (ginger) was probably introduced into Ethiopia in
the 13th century but its use was and still is very limited (Jansen, 1981).
Apparently there is only very little diversity (ginger is propagated
vegetatively), so that collecting by the Plant Genetic Resources
Centre/Ethiopia (PGRC/E) is not worth while and the plant is not
included here.
Also the spices Myrtus communis (Amh.: ades), Lippia javanica
(Amh.: kasse), Mentha spp. (Amh.: nana), Rosmarinus officinalis
(Amh.: siga metbesha) and Thytnus schimperi (Amh.: tosign) are not
listed because they grow abundantly in the wild, they are not
endangered and their use is very limited.
Another spice, Brassica nigra (black mustard), Amh.: senafich, is
not treated here, because it is mainly considered to be an oil crop and
collection through PGRC/E has already taken place.
In the past almost all the important classical spices had to be
imported into the country. Trials are now under way by the Institute
of Agricultural Research (IAR) to introduce at least some of them into
Ethiopia as crops.
Turmeric (Curcuma longa) and cardamom (Elettaria cardamomum)
have been cultivated successfully quite recently. Trials with black
pepper (Piper nigrum) carried out at the IAR Station in Jima are promising. Cinnamon (Cinnamomum zeylanicum) and nutmeg (Myristica
fragrans) could be introduced in the future. Only clove (Syzygiutn
aromaticum) does not find a suitable habitat in this country.
General remarks concerning the collection of spice
germplasm in Ethiopia
As mentioned earlier spices play a very significant role in the
daily food preparation of Ethiopia. So far, small-scale production or
harvesting of wild plants has been sufficient to satisfy the demand of
the people. It is only very recently that the social, economic and
technical situation of Ethiopian agriculture has changed drastically.
With the introduction of improved farming methods, the destruction
of natural habitats and the introduction of advanced cultivars into the
Spice germplasm in Ethiopia
125
country, genetic erosion is very likely to occur even in this group of
minor crops.
Species of spices which still also exist as wild types often show
remarkable degrees of disease resistance. For example, a severe attack
of an unidentified fungal disease was observed in cultivated fennel
but did not attack wild plants (Jansen, 1981). Thus, collection of wild
germplasm and its careful screening afterwards should go hand in
hand.
The ecology, use and need for conservation of the main
Ethiopian spices
Regarding production and cultivated area, only Capsicum,
Rhamnus and Trigonella are of significance. There has been little export
of spices so far. Thus in 1981 only about 900 tonnes were exported,
predominantly red peppers (National Bank, 1982). All other spices
are mainly grown as garden crops, although in certain areas there
may be field production.
Five of the 12 spices dealt with also grow in the wild. Aframomum
and Rhamnus are indigenous spices. It should be mentioned that
Rhamnus is widespread in Africa but so far only Ethiopians are known
to use it as a spice. Aframomum may also occur in south Sudan, but so
far no use has been reported from there (Jansen, 1981). Use and
cultivation of Nigella and Trachyspermum are also typical of Ethiopia,
but they are used elsewhere too.
Out of the 12 spices only Capsicum is of New World origin. Besides
Aframomum and Rhamnus, Trachyspermum may also be of Ethiopian
origin (Wolff, 1927). Ocimum and Piper longum originated in southern
tropical Asia. The remaining seven species have at least one suggested centre of origin in the Mediterranean region (including Egypt
and the Near East). Considering this fact, it is becoming clear that
they have probably been used in Ethiopia since ancient times (Uphof,
1968; Zeven & de Wet, 1982). For Coriandrum, Nigella and
Trachyspermum an especially wide variation can be observed in the
country (Jansen, 1981).
In the following paragraphs the ecology and use of selected species
will be described. Some remarks about the extent to which their
genetic diversity is threatened at the moment are also included. If not
mentioned separately, reference has been made to Siegenthaler
(1963); Jansen (1981) and Goettsch (1984). The total number of accessions held by PGRC/E and given in the following pages refers to 30
June 1986.
126
E. Goettsch
Capsicum annuutn. Capsicum is the most important spice in the
country. According to Alkaemper (1972) ca. 2.5 per cent of the total
arable land (ca. 230 000ha) is cropped with Capsicum. Fruits of red
pepper can be found in almost every market in the country.
Capsicum grows chiefly between 1500 and 2000 m above sea level
but is also found from 1000 to 3000 m. The main centres of production
are Ghion, Bako (Shewa) and the state farms in the Middle Awash
Valley. Red pepper is the main constituent of most kinds of 'wot', a
sauce essential in the daily meal. In addition, it is used to flavour
meat, and medicinal uses are also known. Since 1964 the Ethiopian
Spice Extraction Company has been buying an increasing amount of
red pepper to extract the pigment, which is used as a natural colouring agent.
Even in a small indigenous random population of Capsicum bought
in Addis Ababa (Mercato-market) genetic diversity was very high
(Engels & Goettsch, 1984; see also Jansen, 1981). So far, 126 accessions are held by PGRC/E. Considering the importance of this spice
for local consumption and its increasing significance for export, this
might be regarded as insufficient. Further collecting and screening of
red pepper should therefore be regarded as very important.
Trigonella foenum-graecum. Regarding production, Trigonella is the
second most important spice in Ethiopia. It is grown in all provinces
at altitudes between 1800 and 2200 m and is found for sale in almost
every market.
Fenugreek is an important spice for the preparation of 'wot'. It is
prepared as an appetizer, serves as a milk substitute for babies and is
used as a treatment against rheumatism (Westphal, 1974). PGRC/E at
present holds 427 accessions (213 being their own collections and 214
donated). In spite of the large number of collected accessions, important crop areas for fenugreek are under-represented (e.g. Sidamo,
Bale, Eritrea, Gamo Gofa, Harerge, Welega). Further collection
should concentrate on filling these gaps, but since breeding activities
have been stopped for the moment and the plant is not endangered at
all, fenugreek does not require a high priority.
Nigella saliva. Small-scale production of black cumin is widespread all
over the country between 1500 and 2500 m. Nigella is cultivated as a
crop in the provinces of Gondar (Dembia, Gondar), Shewa (AlemGena), Bale (Dinsho), Harerge (Chercher highlands) and Kefa (Jima
region). The seeds are used in Ethiopia in the preparation of bread,
berbere-sauce ('wot') and local beverages. Nigella seed powder is
added to berbere-sauces to reduce the pungency of the pepper. In
addition there are medicinal uses.
Spice germplasm in Ethiopia
127
Twenty-eight accessions have been collected so far, which is not
sufficient. According to Jansen (1981) the genetic diversity is high.
Economically there is an increasing demand for Nigella (including
export possibilities to neighbouring countries). Thus intensified collecting activities are very advisable.
Trachyspermutn ammi. Ethiopian caraway is found in almost every
market. It is grown at altitudes between 1500 and about 2200 m as a
small-scale crop. Cultivation as a field crop is known from Bale,
Gondar, Eritrea, Gojam and Shewa. Trachyspermum seeds are mainly
used in the preparation of berbere-sauce and bread. Some medicinal
uses are also reported.
Vavilov (1951) considered Ethiopia to be a centre of diversity for
Trachyspermum, where the plant was introduced very long ago.
Trachyspermum is definitely one of the more important and typical
Ethiopian spices and is even grown as a field crop. The plant has
some economic future, justifying an increased collecting activity.
Coriandrum sativum. Cultivation of coriander as a garden crop is widespread all over the country (altitude range 1500-2500 m). The plant is
grown as a crop in Eritrea, Harerge, Shewa, Kefa, Welega and
Gondar. Coriander plays an important role in the Ethiopian domestic
spice trade and its seeds are used for the flavouring of berbere-sauce
injera, cakes and bread (Kostlan, 1913). In Kefa, seeds are added to
cheese and to a porridge made of Colocasia esculenta (taro).
Coriander again shows a high diversity. PGRC/E holds 38 accessions, mainly from Gondar and Welega. Coriander has good export
potential if the quality can be improved (fungal resistance, yield,
etc.), a task which can only be fulfilled with a large variety of local
germplasm at hand.
Aframomum korarima. The use of korarima is known only from Ethiopia where it grows in the forests of Kefa, Sidamo, Ilubabor and
Welega. The plant grows naturally at (1350-)1700-2000m altitude,
with high humidity and annual rainfall ranging from 1300 mm to
more than 2000 mm with no real dry season. Korarima grows in
almost the same habitats as natural coffee.
Cultivation of the plant has been reported not only from places
where it grows wild, but also from the Lake Tana area, Eritrea and
Gelemso (Harerge).
Korarima is very important for flavouring foods. It is used in the
preparation of all kinds of 'wot', coffee and sometimes bread. Compared with other Aframomum species the seeds of korarima have a less
pungent, milder and sweeter flavour. This spice could be developed
into an important article of commerce but further experiments with
128
E. Goettsch
cultivation need to be initiated. There is a demand for korarima in the
neighbouring countries and in Arabia where it has long been highly
prized as a spice (Russ, 1945). There is little doubt that markets could
be found in Europe and America as well.
Hardly anything is known about Aframomum. PGRC/E holds only
16 accessions and nothing can be said about the diversity of the
species. Korarima is one of the spices in which genetic erosion could
be a real danger since its natural habitat, the humid mountain forests
of south-western Ethiopia, will be decimated at an increasing rate in
the future. In order to meet these problems the range of diversity
must be known as a precondition for concentrated cultivation. Thus,
comprehensive collection of germplasm from the wild is urgently
needed.
Cuminum cyminum. In Ethiopia cumin seeds are found in almost every
market. Small-scale cultivation is widespread at altitudes ranging
from 1500 to 2200 m. The ground seeds are mainly used to flavour
different kinds of 'wot', and only small quantities are required.
Cuminum was introduced into Ethiopia a long time ago but so far
no reliable information is available on its range of diversity. PGRC/E
holds six accessions. Since Cuminum is produced almost everywhere
in the country and there is no demand for improvement, its collection
is of minor importance.
Foeniculum vulgare and Pimpinella anisum. Both plants are common in
the highland flora of all regions where they are widespread perennial
weeds. They are occasionally cultivated (altitude range 1500-2500 m).
The ground seeds are a constituent of 'wots'. More important is their
use in the preparation of alcoholic beverages such as 'katikala',
'arake', and 'tedj' (a honey wine).
Both plants are common perennial weeds in the highlands, growing abundantly in the wild so that concentrated collection is not
important at the moment. At present PGRC/E does not have any
accessions.
Ruta chalepensis. Rue is a widespread herb cultivated in gardens in
almost every province of the country (altitude range 1500-2000 m).
The plant is used as a culinary herb. The seeds are needed to flavour
'wots', the leaves are also used as a condiment in coffee and tea. Ruta
is important for the local market only. The plant is extremely widespread and at the moment there is no need for improvement, so that
collection is of minor importance.
Ocimum basilicum. Basil is found in Ethiopia in cultivation as well as in
the wild. The plant is cultivated on a small scale near houses in all
Spice germplasm in Ethiopia
129
provinces. It has a wide altitudinal range from sea level to 2500 m and
it even withstands mild frosts (Jansen, 1981).
Basil is an important and frequently used spice for the preparation
of all kinds of 'wot' and for the flavouring of butter. At present no
genetic erosion has to be feared in basil. A future prospect could be
the extraction of perfume oil from the plant but at the moment this is
of lesser importance. PGRC/E at present holds 12 accessions.
Piper longum. This plant is said to be indigenous to Ethiopia, but the
assumption is more than doubtful. Probably in this case P. longum
was confused with P. guineense, which is at least of (West) African
origin. P. longum certainly originates in India (Uphof, 1968; Zeven &
de Wet, 1982), but there is no doubt that it was introduced in ancient
times into Ethiopia, where it is now found to be growing in the wild.
Like korarima the plant grows in almost the same habitats as coffee.
So far there is only small-scale production. No reliable information is
available on the growing season and other aspects of its husbandry.
P. longum is found in Kefa, Ilubabor and Welega, probably also in
parts of Gamo Gofa (altitudinal range ca. 1500-2000 m). The
inflorescence of the plant (a spike) is used for the preparation of 'wot'.
The taste of P. longum is quite equal to that of black pepper, for which
it serves as a substitute. It is preferred by the local consumer because
of its lower price and greater availability. Thus in future P. longum
could play a more important role in the local spice trade. Little is
known about P. longum in Ethiopia. Its range of diversity - if there is
any - has still to be described. Like korarima its natural habitat will be
endangered at an increasing rate in the future. Collecting and screening of this interesting plant seems to be fully justifiable. PGRC/E
holds three accessions.
Rhamnus prinoides. Buckthorn or 'gesho' is found growing in the wild
all over Ethiopia between 1500 and 2500 m, but it is cultivated as well,
sometimes even on a larger scale as a field crop. Rhamnus covers
about 5000 ha of the land under permanent production (Jansen, 1981).
It is a woody bush, whose leaves are used like hops for the preparation of alcoholic beverages such as 'talla' and 'tedj', which are common household drinks in the country. 'Gesho' is widespread all over
the country. It serves the needs of the people so well that at least at
the moment no improvement is needed.
Conclusions
(a) Collecting of Capsicum is required because the introduction of
improved varieties will reduce the local diversity.
130
E. Goettsch
(b) Aframomum korarima and Piper longum should be collected for
three reasons:
1. Conservation aspect: their habitat will become endangered at
an increasing rate.
2. Scientific aspect: very little is known about the diversity and
other properties of these interesting spices.
3. Economic aspect: korarima especially deserves to be concentrated on since it has a promising potential for export. P.
longum is an important substitute for black pepper.
(c) Nigella, Trachyspermum, Coriander and to a lesser extent Ocimum
are important spices which deserve attention even if they are not
endangered. These spices are highly diversified and promising
economically.
(d) Trigonella is well represented in the collection, which represents
the genetic diversity found in the country, so that further activities should be limited to filling existing gaps.
(e) For the remaining spices Cuminum cyminum, Pimpinella anisum,
Foeniculum vulgare, Ruta chalepensis and Rhamnus prinoides there is
no need for immediate or intensified action.
References
Alkaemper, J. (1972). Capsicum - Anbau in Aethiopien fur Gewiirz- und
Farbezwecke. Bodenkultur, 23, 97-107.
Engels, J. & Goettsch, E. (1984). Capsicum in Ethiopia: some notes on its
diversity. PGRC/E-ILCA Gertnplasm Newsletter, 6, 12-15.
Goettsch, E. (1984). Proposal for further spice collecting at PGRC/E. Plant
Genetic Resources Centre, Addis Ababa, 26 pp. (mimeographed).
Jansen, P. C. M. (1981). Spices, Condiments and Medicinal Plants in Ethiopia, their
Taxonomy and Agricultural Significance. PUDOC, Wageningen.
Kostlan, A. (1913). Die Landwirtschaft in Abessinien. I. Teil: Acker- und
Pflanzenbau. Beiheft Tropenpflanzer, 14, 182-250.
National Bank (1982). Annual Report, 1981. Addis Ababa.
Russ, G.W. (1945). Reports on Ethiopian forests. Reprinted by WoldeMichael Kelecha (1979), Forestry and Wild-life Development Authority,
Addis Ababa.
Siegenthaler, I. E. (1963). Useful Plants of Ethiopia. J.E.C.A.M.A. Experimental
Station Bulletin, no. 14. Alemaya Agricultural College, Ethiopia.
Uphof, U. C. T. (1968). Dictionary of Economic Plants, 2nd edn. Verlag von J.
Cramer, Lehre.
Vavilov, N. I. (1951). The origin, variation, immunity and breeding of
cultivated plants. Chronica Botanica, 13, 1-366.
Westphal, E. (1974). Pulses in Ethiopia, their Taxonomy and Agricultural Significance. PUDOC, Wageningen.
Westphal, E. (1975). Agricultural Systems in Ethiopia. PUDOC, Wageningen.
Wolff, H. (1927), Cuminum and Trachyspermum. In: Engler (ed)., Das
Pflanzenreich, vol. 4, Paper 90:228.
Zeven, A. C. & de Wet, J. M. J. (1982). Dictionary of Cultivated Plants and their
Regions of Diversity. PUDOC, Wageningen.
9
A diversity study in Ethiopian barley
J.M.M. ENGELS
Introduction
Barley (Hordeum vulgare L.), one of the oldest of cultivated
plants, has been grown in Ethiopia for at least 5000 years (Harlan,
1969; Doggett, 1970). Generally, Ethiopia is considered as a secondary
gene centre, or a centre of diversity, for barley and not as a centre of
origin (Tolbert etal., 1979). However, in recent studies some evidence
has been presented to suggest that Ethiopia might be a centre of
origin (Bekele, 1983b; Negassa, 1985) as was originally suggested by
Vavilov. The diversity in Ethiopian barley germplasm accessions has
been presented in a number of studies (Ward, 1962; Tolbert et ah,
1979; Bekele, 1983a,b; Negassa, 1985) which were based mainly on
discrete (non-continuous) characters. In studies on disease resistance
in Ethiopian barley it was found that Ethiopian barley germplasm
possesses resistance genes for almost all major diseases (Moseman,
1971; Lehmann, Nover & Scholz, 1976). In addition, high protein and
lysine contents have been found in some Ethiopian genotypes
(Munck, Karlsson & Hagberg, 1971).
In this chapter a detailed analysis is presented of the phenotypic
diversity in the barley germplasm collection of the Plant Genetic
Resources Centre/Ethiopia (PGRC/E), which possesses considerably
more accessions than have been used in earlier diversity analyses.
The results of previous studies on Ethiopian barley germplasm
will also be summarized, particularly the ones on diversity
indices.
Materials and methods
The records of 3765 accessions in the PGRC/E barley collection were surveyed. These records originated from germplasm collecting missions (e.g. passport data) and from the routine
132
/. M. M. Engels
Table 1. Characters used and their respective classes
Character
Character states
Codes used
1. Kernel row number
6 rows
2 rows"
irregular
lax
intermediate
dense
<15
15-20
20-25
25-30
^30
covered
naked
white-brown
purple-black
<25g
25-35g
35-45 g
45-55g
^55 g
<100 days
100-115 days
115-130 days
130-145 days
=2145 days
<60cm
60-90 cm
90-120 cm
120-150 cm
2*150 cm
6
2
irr.
1
2
3
1
2
3
4
5
covered
naked
1
2
1
2
3
4
5
1
2
3
4
5
1
2
3
4
5
2. Spike density
3. Number of spikelets per spike
4. Caryopsis
5. Kernel colour
6. Thousand grain weight
7. Number of days to maturity
8. Plant height
Both categories, sterile and rudimentary lateral florets, are combined.
characterization activities of PGRC/E. The latter have been carried out
since 1982 at Holetta (2400 m above sea level) and in this analysis data
from the years 1982-5 were used. The data originated from unreplicated small plots (up to 2sqm) and were generally based on the
means of five plants or ears per accession, or on plot means. The
majority of the accessions are morphologically rather uniform since
they were selected from landraces based on their agro-morphological
characteristics.
Data were analysed for the characters, presented in Table 1. Some
characters with continuous variation were included in order to
A diversity study in Ethiopian barley
133
examine their value in diversity studies. The choice of the characters
used was based on the following criteria: their use in previous diversity analyses; their consistency over the years in the characterization
work; and their reliability in scoring. The phenotypic frequencies of
the characters were analysed by the Shannon-Weaver information
index (Hf) in order to estimate the diversity of each character within
each administrative region and within each geographic region. The
country was therefore arbitrarily divided into four ecogeographic
zones, the northern administrative regions (Eritrea, Tigray, Gondar
and Welo); the western administrative regions (Gojam, Welega,
Ilubabor and Kefa); the southern administrative regions (Gamo Gofa,
Sidamo and Bale) and the central and eastern administrative regions
(Shewa, Arsi and Harerge).
The index was calculated as presented by Negassa (1985):
where p{ is the proportion of accessions in the i th class of an n-class
character. In order to keep the values of H' in the range of 0-1 each
value of H' was divided by its maximum value, log en. The standard
error was calculated as follows: SE = S2/r — 1 where S2 is the variance
of the means and r the number of means. In order to determine
whether the variance of the diversity was due to differences between
or within administrative regions a hierarchical ANOVA was conducted
with the normalized data for each character.
Results and discussion
The percentages of the phenotypic classes of accessions for
each administrative region and the weighted mean percentages for
the ecogeographic regions are presented in Table 2. In general, the
mean percentages per ecogeographic area do not show marked variation. The same is true for the frequencies per administrative region
for the majority of the characters. However, some exceptions are
Eritrea, Tigray, Welega, Ilubabor and Kefa for kernel row number
which have high frequencies (>72 per cent) for two-rowed barley. The
spike density frequencies increase from north to south and from west
to east, and show a clinal variation. The covered barleys are more
concentrated in northern and western Ethiopia. The purple to black
coloured kernels are more frequent in the south-west (Welega,
Ilubabor, Kefa, Sidamo and Arsi). Gojam and Welega show the
highest grain weights, although the differences are not significant.
The average number of days to maturity is lower than the Ethiopian
134
/. M. M. Engels
Table 2. Percentage of phenotypic classes of entries for each administrative
region and a weighted'mean percentage of each ecogeographic region for eight
characters
Kernel row
number
Number"
of entries
Administrative region
Eritrea
Tigray
Gonder
Welo
Region
Gojam
Welega
Ilubabor
Kefa
Region
6
55
287
635
143
1120
135
72
14
102
323
Gamo Gofa
Sidamo
Bale
Region
88
58
104
250
Shewa
Arsi
Harerge
Region
Unknown
952
389
159
1500
572
ETHIOPIA
3765
Spike
density
2 irr.
Spikelets
per spike
1
2
3
1
2
3
4 5
18
17
47
46
38
82
81
49
37
57
0
2
4
17
5
38
48
52
43
49
58
44
41
48
44
4
8
6
9
7
0
0
2
4
2
18
17
19
31
20
58
52
54
43
52
22
28
23
21
24
2
3
2
1
2
38
13
21
26
28
47
41
44
44
39
46
58
43
43
52 10
87 0
79 0
72 2
67 5
50 3
55 4
44 12
49 7
57 4
48 6
34 8
52 5
52 5
54
53
64
41
50
40
44
36
56
46
6
3
0
3
4
7
3
0
6
5
46
36
49
45
13
5
12
11
2
0
2
2
51
51
57
42
48
45
50
64
54
19
26
43
35
28
35
36
9
24
2
3
0
2
2
41
59
39
44
21
17
0
15
17
14
12
24
18
42
29
39
38
46
63
36
49
12
8
25
13
7
4
8
6
53
13
45
44
49
45
45
20
29
13
22
25
2
5
2
3
34
26
18
28
24
2 23
42
48
10
4
40 55
5
hi
4
2
1
2
5
48 23 3
a
These numbers vary insignificantly from character to character. Only TGW had a total number of
482 entries.
Table 3. Mean squares and percentage of total variance from the
hierarchical analysis of variance for H' of the individual characters
Kernel row
number
Variance source
Ecogeographic regions
Administrative regions
within ecogeographic
regions
Spike
density
Number of
spikelets
per spike
MS
%
MS
%
MS
3
0.0424
29.2
0.0199
48.5
0.0097
10
0.0309
70.8
0.0063
51.5
0.0111
DF
%
Caryopsis
MS
%
6.9
0.0131
24.7
93.1
0.0119
75.3
A diversity study in Ethiopian barley
Caryopsis
covered
naked
100
97
95
98
96
Thousand
grain weight
(TGW)
Kernel
colour
135
Days to
maturity
Plant height
1
2
1
2
3
4
5
1
2
3
4
5
1
2
3
4
5
0
3
5
2
4
87
82
70
76
75
13
18
30
24
25
0
2
6
_
3
50
15
29
20
50
56
59
_
56
0
27
6
_
21
0
0
0
_
0
0
3
2
1
2
71
51
22
71
38
18
34
60
19
46
11
11
15
8
13
0
1
1
1
1
0
2
0
1
1
24
45
37
43
39
76
51
59
52
56
0
2
3
3
3
0
0
1
1
1
95
99
100
95
96
5
1
0
5
4
77
65
64
65
70
23
35
36
35
30
0
0
_
0
12
14
_
13
23
50
_
32
62
36
_
53
4
0
_
2
4
15
20
9
9
31
51
67
51
43
47
25
0
21
32
18
9
13
18
16
0
0
0
0
0
4
0
0
0
2
18
26
29
28
23
65
65
71
68
66
13
9
0
4
9
0
0
0
0
0
99
100
100
100
1
0
0
0
71
67
82
75
29
33
18
25
0
0
6
2
21
8
41
24
41
38
41
40
26
38
6
23
12
16
6
11
5
16
12
10
56
21
36
40
33
40
31
34
6
24
20
16
0
0
1
0
1
0
1
1
18
22
31
34
77
76
63
71
3
2
5
4
0
0
0
0
96
99
98
97
4
1
2
3
72
67
80
72
28
33
20
28
1
0
0
1
30
16
20
26
47
58
80
51
21
24
0
21
1
2
0
1
4
3
1
3
32
17
48
30
43
36
33
40
20
43
18
26
1
1
0
1
1
2
1
1
39
24
35
35
54
65
43
56
6
8
21
8
0
0
0
0
96
4
69
31
4
22
54
18
2
1
23
43
31
2
2
31
60
6
1
97
3
72
28
2
23
50
23
2
3
33
41
22
1
1
34
58
6
1
Kernel
colour
Number of
days to
maturity
1000° gram
weight
Plant
height
MS
%
MS
%
MS
%
MS
%
0.0187
29.9
0.0329
46.3
0.0197
28.1
0.0150
36.2
0.0132
70.1
0.0164
53.7
0.0096
61.9
0.0080
63.8
a
DF of TGW are 3 and 7, respectively.
136
/. M. M. Engels
average in Eritrea, Tigray, Welo, Welega, Ilubabor, Kefa, Gamo Gofa
and Harerge. Apart from a possible drought escape mechanism
through early maturity, which might have evolved in the northern
administrative regions, there seems to be another natural selection
pressure in the southern and eastern administrative regions. Finally,
the barleys from southern Ethiopia, as well as from Eritrea, have a
higher average straw length than the barleys of the rest of Ethiopia.
These differences between administrative regions, and to a certain
extent between ecogeographic regions, are supported by the percentages of the total variance for each of the two areas (Table 3).
The diversity indices showed relatively wide variations between
characters (Table 4). The caryopsis (covered or naked barley) and
plant height were the least diverse characters studied. Spike density
caused the highest diversity, followed by kernel colour. This agrees
with the earlier findings of Negassa (1985) except for spike density,
which showed little variation in his study. This may have been due to
his smaller sample size or to some selection of the germplasm during
collecting in the field.
The pooled diversity indices over characters within administrative
regions and within ecogeographic regions are relatively uniform. The
least diverse administrative regions are Eritrea and Ilubabor (Table 4).
One of the reasons for this could have been the small number of
samples in the study from these regions. On the other hand, there
might be greater natural selection in barley in both regions due to
generally low rainfall in Eritrea and high rainfall in Ilubabor. The
highest diversity index was found for the Shewa administrative
region, followed by Gojam and Arsi. Only the indices for Shewa and
Eritrea are significantly different (t = 2.35, P < 0.05).
Of the four ecogeographic regions, the central and eastern regions
showed the highest diversity index. All the other ecogeographic
regions were almost equally diverse and were not significantly different from each other (t = 1.84, P>0.05). This finding is also confirmed by the low percentage of the total variance caused by the
'among ecogeographic regions' source (Table 5). Although the percentage of the total variance for 'among administrative regions within
ecogeographic areas' is slightly higher (5.7 per cent) it can be concluded from this table that by far the highest variance is due to
'among characters within administrative regions'.
The overall diversity index for Ethiopian barley is relatively high
for almost all the characters as well as for the pooled index over
characters. These results are similar to other studies (Qualset & Mose-
Table 4. Estimates of the diversity indices (H') for the various administrative regions, for the four ecogeographic
regions and of the mean diversity (Hf) and its standard error over all characters as well as results of some other
authors
Administrative region
Kernel
row
number
Spike
density
Spikelets
per spike
Kernel
Caryopsis colour
TGW
Days to
maturity
Plant
height
H'±SE
Eritrea
Tigray
Gonder
Welo
Region
0.43
0.49
0.77
0.93
0.76
0.76
0.83
0.80
0.85
0.82
0.65
0.69
0.70
0.77
0.72
0.00
0.19
0.28
0.14
0.25
0.55
0.69
0.89
0.80
0.81
0.44
0.65
0.61
—
0.67
0.49
0.70
0.66
0.53
0.69
0.34
0.53
0.51
0.56
0.56
0.46±0.08
0.60±0.07
0.65±0.07
0.65±0.10
0.66±0.07
Gojam
Welega
Ilubabor
Kefa
Region
0.85
0.36
0.46
0.60
0.70
0.78
0.74
0.59
0.74
0.77
0.77
0.76
0.43
0.80
0.77
0.28
0.08
0.00
0.28
0.25
0.78
0.94
0.94
0.94
0.88
0.63
0.62
—
_
0.65
0.72
0.74
0.53
0.74
0.77
0.60
0.52
0.38
0.47
0.56
0.68±0.06
0.60±0.10
0.48±0.11
0.65±0.08
0.67±0.07
0.60±0.07
0.66±0.05
Gamo Gofa
Sidamo
Bale
Region
0.75
0.75
0.89
0.82
0.91
0.76
0.89
0.88
0.75
0.66
0.60
0.71
0.08
0.00
0.00
0.00
0.87
0.91
0.69
0.81
0.81
0.77
0.76
0.85
0.63
0.83
0.85
0.78
0.40
0.38
0.53
0.47
0.65±0.10
0.63±0.11
0.65±0.10
0.67±0.11
0.71±0.08
0.54±0.06
0.60±0.04
Shewa
Arsi
Harerge
Region
0.75
0.80
0.81
0.78
0.89
0.77
0.99
0.89
0.81
0.81
0.77
0.81
0.24
0.08
0.14
0.19
0.84
0.91
0.71
0.85
0.70
0.64
0.31
0.79
0.76
0.75
0.67
0.77
0.57
0.55
0.69
0.59
0.70±0.07
0.66±0.09
0.64±0.10
0.71±0.08
0.64±0.03
0.63±0.08
ETHIOPIA
0.77
0.86
0.79
0.19
0.85
0.74
0.77
0.59
0.70±0.08
0.68±0.02
Ethiopia
(Tolbert et ah, 1979)
0.91
0.53
0.82
Ethiopia
(Qualset, 1975)*
0.71
0.53
0.93
Ethiopia
(Qualset & Moseman, 1966)*
0.75
0.59
0.92
a
b
Calculated by author.
This character is in fact heading time.
0.37
0.51±0.01
0.81*
0.67±0.10
0.75±0.10
H'±SE
(Negassa, 1985)
0.65±0.04
0.54±0.11
138
/. M. M. Engels
Table 5. Hierarchical analysis of variance of the diversity index (Hr) and
the percentage of the total variance
Source
Among ecogeographic
regions
Among administrative
regions within
ecogeographic regions
Among characters within
administrative regions
DF
SS
MS
%
F
3
0.1166
0.0389
1.9
1.12
NS
10
0.3470
0.0347
5.7
0.52
NS
84
5.6229
0.0669
92.4
man, 1966; Qualset, 1975; Negassa, 1985) despite their use of different
types and numbers of characters and different sample sizes (varying
from 485 to 3765 samples) and the use of classified quantitative
characters in this study. Tolbert et al. (1979) reported lower values,
probably due to the use of growth habit (winter or spring) and awn
type (rough or smooth), as these characters do not vary much or at all
in Ethiopian barley. Thus the conclusion by Tolbert et al. that Ethiopia
is a secondary centre of diversity may be revised if other more
relevant characters are used in diversity studies. In the present study
awn type was omitted as all accessions were rough.
The use of quantitative characters which were scaled in an arbitrary
way seems to be justifiable and useful as the barley accessions show
considerable variation within and between administrative regions for
these characters (Table 3) and they are, in general, of more interest to
the plant breeder than the discrete or qualitative characters.
However, because of the arbitrary decision on the number of classes
per character and their influence on the magnitude of the diversity
index, a comparison of the indices from different studies is
meaningless.
The results of this analysis have shown that Ethiopia is a centre of
diversity for barley and that this diversity is rather evenly distributed
over the barley growing areas of the country, although there are some
concentrations for individual characters. Furthermore, the initial
results of a study of the diversity index by altitudinal strata of Ethiopia as a whole have shown an obvious relationship between the
diversity index and the altitude. The index is highest around 25002600 m above sea level and decreases with increasing or decreasing
elevation (Engels, 1990).
A diversity study in Ethiopian barley
139
Acknowledgements
The author would like to thank all colleagues involved in the
data collection during the collection and characterization work at
PGRC/E as well as Mrs Karin Ralsgard for the data on thousand grain
weight and Dr John Lazier for his critical comments.
References
Bekele, E. (1983a). Some measures of gene diversity analysis on landrace
populations of Ethiopian barley. Hereditas, 98, 127-43.
Bekele, E. (1983b). A differential rate of regional distribution of barley
flavonoid patterns in Ethiopia, and review on the centre of origin of barley.
Hereditas, 98, 269-80.
Doggett, H. (1970). Sorghum. Longman, London.
Engels, J. M. M. (1990). The genetic diversity in Ethiopian barley in relation to
altitude. In: S. Iyama and G. Takeda (eds), Proceedings, 6th International
Congress of the Society for the Advancement of Breeding Research in Asia and
Oceania, 21-25 August 1989. SABRAO, Tsukuba, Japan, 107-10.
Harlan, J. R. (1969). Ethiopia: a centre of diversity. Economic Botany, 23,
309-14.
Lehmann, C. O., Nover, I. & Scholz, F. (1976). The Gatersleben barley collection and its evaluation. In: H. Gaul (ed.), Barley Genetics, vol. III. Proceedings
3rd International Barley Genetics Symposium, Garching, 1975. Karl Thiemig,
Munich, pp. 64r-79.
Moseman, J. G. (1971). Co-evolution of host resistance and pathogen
virulence. In: R. A. Nilan (ed.), Barley Genetics, vol. II. Proceedings 2nd
International Barley Genetics Symposium, Pullman, 1969. Washington State
University Press, Pullman, Washington, pp. 450-6.
Munck, L., Karlsson, K. E. & Hagberg, A. (1971). Selection and characterization of a high protein, high-lysine variety from the world barley collection.
In: R. A. Nilan (ed.), Barley Genetics, vol. II. Proceedings 2nd International
Barley Genetics Symposium, Pullman, 1969. Washington State University
Press, Pullman, Washington, pp. 544-58.
Negassa, M. (1985). Patterns of phenotypic diversity in an Ethiopian barley
collection, and the Arussi-Bale Highland as a centre of origin of barley.
Hereditas, 102, 139-50.
Qualset, C O . (1975). Sampling germplasm in a centre of diversity: an example of disease resistance in Ethiopian barley. In: O. H. Frankel and J. G.
Hawkes (eds), Crop Genetic Resources for Today and Tomorrow. Cambridge
University Press, Cambridge, pp. 81-96.
Qualset, C O . & Moseman, J.G. (1966). Disease reaction of 654 barley
introductions from Ethiopia. USDA/ARS Progress Report (unpublished).
Tolbert, D. M., Qualset, C O., Jain, S. K. & Craddock, J. C (1979). A diversity analysis of a world collection of barley. Crop Science, 19, 789-94.
Ward, D. J. (1962). Some evolutionary aspects of certain morphological
characters in a world collection of barley. USDA Technical Bulletin 1276.
10
Sorghum history in relation to
Ethiopia
H. DOGGETT
Introduction
Hypotheses on crop development in Africa are long on
theory and short on fact. Let me present my own ideas. Harlan &
Stemler (1976) have presented a theory that sorghum developed in
the southern Sudan-Chad region. My problem with that is the
answer to the question 'how'? It is true that as soon as Man began to
sow the seeds of wild grasses, selection and sowing over the years
would result in cultivated types being developed. That assumes that
Man somehow learnt the idea of agriculture. Perhaps he did, but it is
not clear how this happened in the rainfed savannahs. There were
lots of grasses there anyway, and lots of grass seed. Why sow more?
How would man have learnt to sow seed? How would he have
distinguished between the masses of grass seedlings coming up with
the rains and those which he had put in? By clearing a separate plot of
land for sowing? That presumes the idea of agriculture. Having
worked in the tropics for many years, I find this altogether too difficult to imagine.
It is more likely, to my mind, that agriculture was discovered along
rivers and that the discovery was a rather rare event. One should
always look at the possibility of the spread of the idea of agriculture
from elsewhere before concluding that it had been discovered all over
again. My scenario for the discovery of agriculture in a situation of
this kind is presented below.
Discovery of agriculture
The three oldest civilizations of the Old World all arose along
rivers: in due course, each spread out along its respective river valley
for hundreds of miles (Fairservis, 1971). Rivers and seasonal streams
Sorghum history in relation to Ethiopia
141
provide sites from where the idea of cultivation may have emerged.
Many patches of silt, exposed as the rains ended and the rivers fell,
would have been weed-free at first. People gathering seeds of wild
grasses for food, who also fished, could well have noticed that seeds
dropped on these patches sometimes grew into mature plants on
residual moisture. From this, the use of sickles for harvesting would
have favoured the variants with persistent spikelets. Gradually, the
idea of deliberately sowing these riverine flats with a seed so
harvested, and replanting the following season, would have led to
the accumulation of non-shedding types. People would gradually
have become accustomed to the regular discipline of seed-time and
harvest on silt flats needing no land preparation and no weeding.
This would have provided an additional resource; fishing, foodgathering and hunting would have continued as before. Once seeding became an established practice, it is not difficult to imagine a
gradual awakening of interest in crop improvement as more desirable
types were noticed.
This reconstruction of the possible origins of agriculture also provides an explanation for the way in which people became locked into
the hard labour and drudgery involved. So long as people were using
the natural resources of hunting, fishing and food-gathering, the
population could not increase beyond the number those natural
resources would carry. Improved harvesting and grass-seed processing technology made better use of the resource base but did not
enlarge it. Learning to seed the silt flats deliberately was a different
matter. This enlarged the resource base and provided a way to feed
an expanding population. As the population grew, more silt flats
could be seeded. In due course the population expanded beyond the
point of no return. No longer were hunting, fishing and food-gathering sufficient. From then on, the pressures demanded the extension
of irrigation, preparation and weeding of land to imitate the conditions on the silt flats, leading on eventually to the development of
rainfed agriculture. For that, there was basic crop husbandry to be
learnt: clearing the land, tillage, the time and method of sowing, and
weed control. All this had to be done initially with stone axes and
sticks as the only tools.
Movement of crops
Crop movement presents a problem. Near the beginnings of
agriculture, the idea of agriculture needed to move together with the
new crops. Movement of the agriculturalists themselves is one
142
H. Doggett
obvious method; they may then have acted as focal points for the
teaching of the new technology. Their new neighbours would come
to learn this remarkable new technology and both crops and
methodology would spread from such focal points. Conquest must
also have been a vehicle for the transfer of crops and technology and
the success of the agriculturalists may well have made them liable to
attack from jealous neighbours. The long-distance transfer of
individual crops other than through actual carriage by agriculturalists
is difficult to imagine, unless the recipients were themselves agriculturalists (Gramly, 1979).
It seems most unlikely that settled arable agriculturalists ever
moved until forced by circumstances to do so. They then took their
technology and high-yielding varieties as a package with them, having first located a site which they considered suited to their crops and
methods. Harlan & Stemler (1976), referring to the spread of agriculture to the west and to the east, noted that 'what moved out of the
nuclear area (West Asia) was a complete system including barley,
emmer wheat, einkorn wheat, lentil, vetch, pea, chickpea, faba bean,
rape, flax, vegetables, spices, tree and vine fruits, sheep, goats, cattle
and an array of agricultural techniques'. Doubtless they moved very
much as the people under pressure in the Sahel zone are moving
today. The man of the family goes south, living as best he may. He
prospects, and if he finds a suitable area he returns to help his family
pack up and they move, taking with them their tools, seed and
livestock, together with their accumulated agricultural knowledge
and wisdom. Groups of several households may emigrate together
for mutual protection and support.
Agricultural development in north-east Africa
The people
Language and archaeological studies (Hiernaux, 1974; Ehret,
1979) show that a long-headed, long-faced people had been present
in north-east Africa since the latter Pleistocene, in much of the area
labelled today as Sudan, Ethiopia, Somalia, Tanzania, Kenya,
Rwanda and Burundi. They were Africans, not 'Mediterranean
types'. (In this chapter, 'Ethiopia' will be used for the area south of
the confluence of the Blue Nile and the Atbara, and east of the White
Nile along the rivers of the plain). The Afroasiatic language group
arose in the Ethiopian area, extending roughly from the Amba Farit
mountains on the west, past Lake Tana and reaching almost as far as
the Lake Nasser of today. On the eastern side it followed the Red Sea
Sorghum history in relation to Ethiopia
143
hills and the shores of the Red Sea. This language group later developed into Semitic, Berber, Ancient Egyptian, Cushitic, Omotic and
Chadic. The proto-afroasiatic people lived at least 15000 years ago.
Proto-cushitic was being spoken at least 9000 years ago. Descendants
of these people spread widely across Africa and into the Mediterranean region. The Semitic people moved out from Africa, some of
them returning later. These people were strongly associated with
grass-seed collecting and pottery. It is tempting to suggest that the
value of fermentation in utilizing grass seed for food was known to
them. Injera may have a long history.
The Nile
The Nile has always been a major route into Africa and the
agriculture of south-west Asia developed within travelling distance of
the Nile. It was certainly of greater antiquity than agricultural
development in Africa. There have been big climatic changes along
the Nile. During the last Ice Age, the Mediterranean climate was
forced south into Africa and the Mediterranean plants survived there,
including some of the grasses from which crop plants were later
derived. Wendorf & Schild (1984) have excavated key sites along the
Nile and demonstrated human activities along the river over
thousands of years, including a strong indication of arable agriculture. They thought that a very early cultivated barley had been
discovered but subsequent tests showed that the find was an intrustion. We do know that the climate was growing warmer. We also
know that there was an ancient trade route up the Nile and the
Atbara through to the frankincense and myrrh products of Saba. The
presence of barley has been demonstrated in Egypt in the Fayum,
dating probably to the fifth millennium BC (Arkell & Ucko, 1965). The
Nile has silted some of the areas that archaeologists would like to
explore, and the use of barley in Egypt could be yet earlier.
Agricultural development in Ethiopia
The author suggests that barley was taken into Ethiopia by
people from Egypt, or from cultivators along the banks of the Nile if
barley was indeed grown there. The cool conditions favouring the
cultivation of barley were moving northwards towards the Mediterranean and temperatures along the Nile were rising. Some people
had already become dependent upon barley, using the technology
that was common along the river prior to the climatic change. Small
groups of these people would have moved into the hills, following
144
H. Doggett
the barley climate as it receded. This probably took place along the
Blue Nile, the Atbara and other tributaries of the Nile system. Alternatively, barley may have been introduced up the Nile from West
Asia later. According to Ethiopian tradition, barley is a very ancient
crop in that country. There is a great diversity of barleys in Ethiopia
and Helbaek (1960, 1966) drew attention to the whole series of forms
grown at the beginning of agriculture in Egypt. He also recorded
Hordeum irregulare from the Fayum and this group all seem to have
originated in Ethiopia. Helbaek also noted resemblances between the
ancient emmer wheats of Egypt and some modern Ethiopian types.
Harlan (1969) recorded the great variability in the barleys and
tetraploid wheats in Ethiopia. The barley-with-emmer combination of
the Ethiopian highlands was important in ancient Egypt, dating back
to ca. 4500 BC.
Situation in the hills
Settled cultivators are sitting targets. After harvest, they have
a stock of food which others would gladly seize. The arable agriculturalists, therefore, have, in the past, occupied defensive positions
on the hills. They may well have cultivated in the valleys as well,
returning to their defended communities daily before nightfall.
The early cultivators in the hills were caught between increasing
population size, on the one hand, and the climatic and ecological
limitations of barley culture on the other.
The climate became warmer and drier and population numbers
increased. The early agriculturalists responded to these challenges in
two ways: (a) by domesticating new crops adapted to warmer or more
difficult conditions than those suited to barley; (b) by developing a
more intensive agricultural system.
Domestication of new crops
Crops that can survive well on difficult soils in the barley
zone include niger seed (noog, Guizotia abyssinica), teff (Eragrostis tef)
and linseed (Linum usitatissimum).
Niger seed was almost certainly ennobled in Ethiopia. Teff could
well have originated from one of the preferred grasses of the grassseed collection days, taken into cultivation as a result of learning the
principles of agriculture and subjected to selection for persistent
spikelets. To the casual observer, teff is a wild grass. Linseed was
probably introduced as an edible oilseed crop. The fibre (flax) was
used by the Egyptians, especially for fabric with which to bind the
dead.
Sorghum history in relation to Ethiopia
145
Two cereal crops extending from below the 'barley line' in the
highlands to the lowlands are finger millet (Eleusine coracana) and
sorghum. There is little doubt that finger millet was developed from
E. africana. One archaelogical find probably dates to the third millennium BC (Mehra, 1962; Phillipson, 1977a; Hilu, de Wet & Harlan,
1979). Thus a range of crops was developed, or introduced from West
Asia, adapted to a wide range of soils and climatic conditions. The
development of sorghum will be considered below.
Development of cultural methods, with soil and water
management
Important developments in soil and water management,
essential to reduce the effects of population pressure on land in the
Ethiopian hills, may be illustrated by looking at the current agriculture of the Konso. These people have lived in south-west Ethiopia
for a long time, although they claim to have inherited at least some of
their practices from the Mado people who preceded them. We may
speculate about the order in which the various practices were developed, but the whole 'package of practices' is impressive. The Konso
lived in relative isolation (apart from contacts through the market
systems) prior to 1896 and may be regarded as inheritors of an ancient
agricultural tradition developed over many centuries.
In order to minimize erosion, the soil is retained by the construction of many hundreds of miles of stone terraces, which follow the
contours. They are built as dry stone walls, the soil being cut away
vertically and the wall built against the vertical face. Only undressed
stones from the ground near the place of construction are used, but
with great skill and neatness. The terrain is steep; terraces are often
about 2.5 m wide and 1.5 m high. The wall projects above the level of
the field it is retaining. After heavy rain, a length of wall may collapse, but is immediately rebuilt by the owner who will rush out
naked in a rainstorm to see what is happening to the water on his
land. The land itself has a ridge on the outside and other ridges are
made at right angles to it, forming a series of boxes to hold the water,
as with tie-ridging or basin listing.
Any streams are used to irrigate the fields and are walled to protect
the fields from flood water. Elaborate stone leats are constructed to
allow the water to pass through a series of walled gardens. Such
irrigable streams are rare: most of the stone-lined drains carry storm
water and are used as paths, especially for cattle. The runoff is carefully channelled through leats on the land and the owner will be there
during heavy rain to see that water is being well distributed over his
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H. Doggett
land. Water for domestic use is obtained from wells or from the few
permanent streams. Huge reservoirs have been constructed to conserve rainwater for cattle; dams may be as much as 12 m high and
more than 60 m in length, containing many hundreds of thousands of
litres. Towns are usually situated on high ground and the stream
beds are in the valleys. Water may be collected from points half-anhour's walk from the town and 60 m below it.
Soil fertility is maintained by the liberal use of manure, which is
applied once before sowing and frequently during the growing
season. Human manure is used. In each town, there are a number of
places, generally along the outer walls, for defecation. The faeces dry
quickly in the sun; they are collected and mixed with animal manure
and then periodically taken to the fields. This may well be a further
indication of the age of agriculture in the area. It is hard to believe that
the organized use of human manure would have been adopted and
retained as an ancient custom if animal manure had been readily
available. The manure is collected outside the homesteads and left to
rot; in some areas, pits are dug in which the dung can mature.
The people live in walled towns with gates built in defensive positions. Only in recent years have the gates been neglected and security
relaxed. The cattle (including sheep and goats) are penned within the
homesteads and are partly stall-fed with fodder cut and carried from
the valleys and lowlands. They are taken out under careful supervision along certain walled paths to the grazing area. Only a few
pastures are found near the town; the greater part of the available
land is situated some distance away and the cattle are grazed there.
Many of the distant fields are terraced, but not manured, and rotation
with fallow is practised; the grazing of the cattle doubtless contributes
to fertility maintenance (Hallpike, 1970, 1972).
Ploughing was introduced by the Amhara. Traditional cultivation
used a three-pronged hoe of a type found formerly in ancient Egypt.
Konso cropping pattern
The plateau of the Takadi area to the west is only a few
hundred feet above the Garati area to the east, yet the cropping is
different. Wheat and barley are the main crops on the Takadi plateau;
linseed, sorghum and finger millet are also grown there. Sorghum is
grown mainly on the lower ground - the Garati area - and ripens
several weeks before the same crop on the plateau. Most sorghum is
interplanted with finger millet. Maize has now also become an
important crop. Some ensete is grown, but it is not very popular. The
long-established ensete cultivation of south-west Ethiopia should be
Sorghum history in relation to Ethiopia
147
noted. This vegeculture could be very ancient and perhaps people
from this area first made contact with the barley crop and seed-crop
agriculture further up the Nile. Barley is a common crop along with
ensete at 3000 m (Hallpike, 1970; Westphal, 1975).
Origin of the sorghum crop
Area of origin
Mann, Kimber & Miller (1983) outlined the current hypotheses on the time and place of the origin of sorghum cultivation.
There can be no doubt that the cultivated sorghums of today arose
from the wild Sorghum bicolor subsp. arundinaceum. There is no
evidence of cultivated types ever having arisen from the rhizomatous
diploid or tetraploid Halepensia. The wild forms of S. bicolor were
confined to Africa until recent historical times and it is certain that the
crop was domesticated on the African continent.
The Saharan and north-eastern regions of Africa (at least) enjoyed
a pluvial period (with interruptions) prior to 3000 BC. The sorghums of
those days were doubtless adapted to wetter conditions; many of the
wild types still are. De Wet, Harlan & Price (1970) listed the distribution and habitat of 16 of Snowden's wild 'species'. Seven of them
belonged to wet or humid areas; five were characteristic of hot, dry
regions; all belonged to damp places, swamp and stream margins, or
irrigation ditches.
Wild sorghum occurs in Ethiopia up to about 2300 m above sea
level. It is fairly common at 1500-1700 m and shatter canes (derivatives of wild x sorghum crosses) are the most serious weed around
1700 m in the central plateau, where they are known as 'keelo' (the
fool). In October 1982, the author sampled wheat fields along the
road from Debre Zeit (1800 m) to Nazret (1600 m) and for some 27 km
down the road towards Awasa. In a distance of 77 km, stops every
few kilometers showed that 97 per cent of these wheat fields contained at least a few plants of wild sorghum. My suggestion that wild
sorghum, occurring as a weed of cultivation, attracted the attention of
early agriculturalists remains a possibility (Doggett, 1965).
South-western Ethiopia provides the type of sites where sorghum
could have been ennobled. The agriculturalists of south-western
Ethiopia would have lived near their main crops and needed the high
altitude, also for defence. Lowland crops would have been cultivated
along the rivers, as is the present-day practice of the people of Gamo
Gofa, Kefa and Ilubabor. The Konso also continue this ancient agricultural way of life. They grow cold-tolerant crops in the highlands.
The Konso grow at least 24 varieties of sorghum, including race
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H. Doggett
guinea. Race caudatum and type bicolor are the main forms grown.
Sorghums are the staple crop and are grown on the higher ground,
but much is also grown on the lower ground. Sorghum is often
ratooned (Hallpike, 1970, 1972).
Gebrekidan (1970), on a trip to the Konso area, noted wild species
along the roads and the abundance of shatter canes. He recorded the
importance of bicolor types and the fact that the cultivated forms of
sorghum all had very thin, grass-like stems with both compact and
loose panicled forms represented. These were all growing in the
highlands and were cold-tolerant when tested at the University in
Harare.
Time of origin
The sorghum races guinea and durra reached India; the more
recent races caudatum and kafir did not. Links through Arabia to
India may have been severed in the third century BC when the 'Abyssinians' invaded south-west Arabia.
Finds of sorghum in India have been reported from Jorwe, ca.
1000 BC, together with finger millet (Kajale, 1977); from Pirak, on the
edge of the Indus plain, ca. 1350 BC; at Ahar near Udaipur, ca. 1500 BC;
and at Imagon, near Ahmadnagar, between 1800 and 1500 BC
(calibrated) (Allchin & Allchin, 1982). These dates give estimates of
the latest date by which sorghum had reached India. The crop may
well have arrived there earlier.
There is one early date for sorghum in the Sudan. Excavations at
Kadero, 18 km north-east of Khartoum and some 6 km to the east of
the Nile, were dated to the second half of the fourth millennium BC.
There, numerous grindstones and abundant sherds carrying
impressions of grass seeds were found. Klichowska (1984) regarded
one group of 15 impressions as S. vulgare, and measured the mean
dimensions as 3.4 x 3.6 mm. A second group, her Sorghum cf. vulgare,
had 11 impressions averaging 3.7 x 3.4 mm. These dimensions lie
well outside the grain sizes of wild sorghum and within the ranges of
several cultivated types. (As a comparison, the bicolor sorghum
c.245, recorded by Clark & Stemler (1975), measured 3.0-3.4 X 2.32.9 mm.) Similarly, the dimensions for 20 impressions of Eleusine
grains averaged 2.1 X 2.0 mm, which lies within the range of dimensions of cultivated finger millet grains. Unless the impressions much
exaggerated the size of the original grains that made them, there were
cultivated sorghums and finger millets in the Kadero material
(Harlan, 1969).
Another discovery of impressions of cultivated sorghum dates
Sorghum history in relation to Ethiopia
149
back some 4500 years and was found in Abu Dhabi, near the mouth of
the Persian Gulf (Cleuziou & Constantini, 1982). Cultivated finger
millet has been found in Ethiopia probably dating to the third millennium BC (de Wet et al, 1984).
Phillipson (1977b) accepted the third millennium BC as a general
estimate of the period during which many African crops were
brought under cultivation. At present, there are no grounds for modifiying the opinion that sorghum was first developed in the north-east
quadrant of Africa some 5000 years ago, probably in the EthiopiaSudan region (Doggett, 1965).
Development of the sorghum crop
The use of the names of the different sorghum races in the
following paragraphs is based on the classification of Harlan & de
Wet (1972).
Bicolor sorghums
The earliest stages of ennoblement would have produced
bicolor types. Bicolor sorghums with small grains and often loose
panicles are frequently found in wet conditions. With these loose
panicles and very dark, small grains often covered by the glumes,
they are adapted to high-altitude, humid conditions. The grains dry
quickly and are little troubled by birds or grain moulds. These forms
were developed in ecological situations similar to those which they
now occupy. They are used for beer, for their sweet stems and for
special kinds of food preparation. We possess these samples from the
past history of the crop because they are still worth growing in
appropriate habitats for particular uses. It is not known when the
bicolors were carried to India, but they were presumably among the
earliest to arrive there.
Guinea sorghums
The wild sorghum race arundinaceum occurs all along the
northern forest margin of western Africa, in northern Uganda and on
through southern Sudan into Ethiopia. Sorghums of Snowden's
Guineensia are found along the same belt. The association between the
wild arundinaceum and many of the cultivated guineense, and the
derivation of one from the other, need not be questioned. Race
guinea contains cultivars adapted to high rainfall areas and others
adapted to the decrue agriculture of Mali. In so far as the Shallu
derivatives are representative of the guinea sorghums, they are adap-
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H. Doggett
ted to humid rather than to dry conditions, with a low level of
drought tolerance.
Where were the guineas developed? There is no evidence of
annual seed crop cultivation in West Africa before 1500 BC, SO the
development of guinea sorghum in western or central Africa, followed by its movement to India, seems unlikely. The most probable
area of origin lies in western and south-western 'Ethiopia', stretching
as far as the White Nile. An island of guinea sorghum survives in the
highlands of south-west Ethiopia, grown by the Konso people practising an ancient agriculture, and isolated today from the nearest
guinea sorghum on the plain to the south by a distance of some
300 km.
The guinea sorghums spread to India and south to South Africa.
They also spread across Africa along a rather narrow belt of land. S.
roxburghii is the principal member of the Guineensia occurring in
eastern Africa and also in India. Coastal sea traffic sailing on the
monsoons has connected the two continents for over 2000 years.
Guinea sorghums occur very widely in coastal areas of the Old
World, having been spread by ship along the coasts of South-East
Asia.
Durra sorghums
The country possessing the best relics of the development of
durra sorghums is Ethiopia. Bicolor types occur in western Ethiopia,
typically in high rainfall, highland areas. Forms with larger, more
'cultivated type', grains are found in the less humid, less rainy parts
of the country. They combine some of the features of bicolor and
durra, for example, the local cultivars 'Fundishu' and 'Zangada'.
Other forms would be classified as Harlan and de Wet's race durracaudatum, especially in Gondar, Simen and Tigray. In the dry areas,
durra types with their large grains and compact panicles predominate, notably in the Harerge administrative region on the Chercher
highlands.
Race durra probably developed through introgression with the
wild type aethiopicum, which occurs in the drier areas of Ethiopia.
Above average levels of drought resistance and seed size are characteristics of this race. As the Ethiopian climate dried and early bicolors
were moved eastwards into drier areas, still under selection by able
agriculturalists, introgression with wild forms adapted to drier conditions occurred. Intermediate types between the races are also found.
K. E. Prasada Rao (personal communication) reviewed the collec-
Sorghum history in relation to Ethiopia
151
tion in Ethiopia (4000 entries) and noted 25 local races of durra, eight
of durra-bicolor, three of durra-guinea and one of caudatum-bicolor.
There were also three races each of bicolor, caudatum and guineacaudatum. All the durras and durra relatives had been collected from
high and medium altitudes. Panicle type is governed by the humidity
of the environment at flowering and ripening time and very dense
panicles are found in types which flower and ripen grain under really
dry conditions. Compact durras are outstanding in such situations,
but open-panicled durras occur in the higher rainfall areas of Ethiopia. The whole sequence, wild type-bicolor-durra-bicolor-durra, can
be seen clearly in Ethiopia. From Ethiopia, durra spread westwards
through the Sudan and across Africa, occupying the dry belt below
the southern margin of the Sahara.
The semi-nomadic peoples of Somalia still grow durra sorghums in
a well developed agricultural system, which is certainly old, and
could be very old. The durras were probably carried from the Horn of
Africa through Yemen and Saudi Arabia. From there, they may have
moved into Iran and Afghanistan or West Asia, or through Oman and
across the Baluchistan. They were then carried either through the
Punjab to northern India, or through Sind to peninsular and southern
India.
K. E. Prasada Rao (personal communication), collecting in the hills
of Madhaya Pradesh occupied by 'Tribals' practising a simple agriculture, noted that the 'pig-mouth' durras grown by the Tribals in the
800 mm rainfall zone are similar to the 'Zurru' durras of Tigray region
in Ethiopia.
Caudatum sorghums
Caudatums are grown by the Konso, as noted above. Both
caudatums and half-caudatums are grown in Ethiopia, especially in
the lowlands. The fact that they do not occur in India shows that this
race is younger than guinea or durra and also suggests that
caudatums could not arise from all the combinations of durra and
guinea crosses that must have occurred there. It is probable that
continued interaction with the wild sorghum gene pool was necessary for caudatums to appear. Caudatums are often associated with
pastoralists.
The caudatums encountered wild and bicolor sorghums in the
Sudan plains, as well as durras and probably guineas, and introgression occurred. Many must have been subjected to bulk mass selection
every time the pastoralists harvested them. Often they grew in the
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H. Doggett
presence of Striga, shoot-fly, other pests, diseases and increasingly
tough environmental conditons as the climate continued to dry. The
casual agricultural standard of the pastoralists encouraged intercrossing. Generations of crude mass selection in these populations,
harvesting the larger grained survivors, resulted in the development
of really tough caudatums, able to yield in spite of the ills listed above
and also possessing a degree of bird resistance.
Stemler, Harlan & de Wet (1975) gave an excellent account of the
association between these sorghums and speakers of the Chari-Nile
languages, linked to their migrations. They noted the importance of
caudatum sorghums to the people of northern Cameroun and around
Lake Chad and suggested that cultivars in the savannah belt from the
eastern side of Lake Chad through the southern half of the Sudan
were probably growing caudatum sorghum by about AD 1000 and in a
belt between 9°50 and 12° N. That must surely be correct. The arrival
of caudatums much further west in Africa could be relatively recent
and they are still spreading southwards into the guinea zone. Essentially, this race occupies the belt across West Africa between the durra
and the guinea races.
On the eastern side, the Chari-Nile speakers carried these
sorghums down to Lake Victoria. The Karamajong still use them in
the traditional way, but some of the Luo in Kenya are now settled
agriculturalists, growing these sorghums in addition to other types.
Further south, some of these caudatums were moved through into
Tanzania, where the Wasukuma use them on heavy land in the
quelea bird areas of the eastern Shinyanga district. The Wasukuma
agriculture contains an important component of Ethiopian agricultural methodology. This is particularly true of the peoples on
Ukara Island in Lake Victoria. Caudatums were well represented in
Snowden's material from Tanzania. More recently, on two collecting
trips in the Dar-es-Salaam-Dodoma-Mwanza-Musoma areas,
Prasada Rao & Mengesha (1979) collected 154 cultivated sorghums
which contained 27 caudatums, 30 durra-caudatums and 5 guineas.
High-altitude durra-caudatums are found in western Uganda,
Rwanda and Burundi and have probably been there for a considerable time.
Kafir sorghum
Harlan & de Wet (1972) took Snowden's S. coriaceum and S.
caffrorum as their race kafir, which gives a rather distinct group
occupying the region from Tanzania to southern Africa. Snowden
Sorghum history in relation to Ethiopia
153
reported kafir collections from Tanzania, Zambia, Zimbabwe, Angola
and South Africa. In the Transvaal, there are some excellent kafir
cultivars growing under high-altitude conditons which set seed
equally successfully when planted in the highlands of Ethiopia. The
sorghum crop is deeply involved in many of the traditional
ceremonies of the Wasukuma and other Bantu tribes.
Race kafir spread neither to West Africa nor to India and it does not
seem to have reached either Ethiopia or the Sudan. This race is associated with the Bantu and their spread into southern Africa.
Spread of the Ethiopian crops and technology
Mid-African drainage basin
Sutton (1974) presented evidence that, between the ninth and
third millennia BC, the wetter conditions prevailing resulted in the
water levels in the lakes and rivers being high with some of the
internal basins temporarily linked, especially in the 'Middle African'
belt. From western Ethiopia, the all-important idea of agriculture,
together with the early cultivated sorghums, moved along the margin
of the aquatic culture towards the west, no doubt often used in the
decrue agriculture which is still practised along the rivers of southwest Ethiopia as they enter the plains.
Eastern and southern Africa
Crop movement through eastern and southern Africa was the
result of human migrations. The Konso are the inheritors of the crops
and technologies of early agriculturalists. These latter were under
relentless pressure and met the challenges through skill and sheer
hard work. Gebrekidan (1970) wrote: 'The Konso are probably one of
the hardest, if not the hardest, working people in Ethiopia.' This
must also have been true of the early agriculturalists. They developed
new crops and intensified their production methods, yet population
pressures on the land grew. Eventually, the point was reached when
some of the people had to move. These people were highly professional agriculturalists whose survival depended on hard work and
'getting it right'. They had their selected seeds and their agricultural
methodology - high-yielding varieties and a package of practices.
When forced to move, they took their seeds, their animals and their
knowledge with them. They looked for sites where familiar types of
agriculture could be practised.
Such movements were probably made by small groups of people
who colonized suitable hills and established scattered nuclei of devel-
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H. Doggett
oped agriculture. As these settlements in turn became overcrowded,
some were forced to move on, so that those further afield were the
most recently settled. Agriculture was new and populations were
sparse.
East Africa
The Cushitic and Omotic speakers of south-western and
southern Ethiopia expanded into East Africa, moving initially down
the Rift Valley from Lake Turkana, and could well have come from
the Kefa, Gamo Gofa and Sidamo areas. They were followed by other
waves of immigrants from southern Ethiopia, moving through the
mountains of western Kenya and adjacent Tanzania, bounded by
Mounts Kenya and Kilimanjaro on the east, Mount Elgon and Lake
Victoria on the west and Lakes Eyasi and Manyara to the south. This
was not an immigration of arable agriculturalists on a broad front.
Much of East Africa was livestock country and many of the
immigrants were pastoralists. The arable agriculturalists, with their
mixed farming, would have consisted of small groups who colonized
scattered defensive positions on hills, where they established their
terraces and settled agriculture. Doubtless they influenced the indigenous inhabitants in due course. Dates on pottery at several sites
from West Kilimanjaro go back to 3000 BC. Phillipson (1977b) wrote: 'It
is tempting to suggest that a gradual spread of Cushitic speakers . . .
began at some time.'
Movement southwards
It is likely that the movement south was continued by small
groups of skilled agriculturalists, carrying the Cushitic/Omotic traditions. Some of the terraced sites appear to have been established
initially in the Stone Age. There are undressed stone terraces at
Inyanga, Zimbabwe, but no dates. Gramly (1979) argued that crop
production must have preceded iron-working, to provide the profit
motive for the ironsmiths. Craftsmen need a fair return for their
products. An initial immigration of small groups of professional agriculturalists, with their crop cultivars and their technology, followed
by the spread of agriculture to the surrounding people and the incoming Bantu, represents the probable prelude to the Iron Age in
southern Africa.
Gramly (1979) suggested that the iron technology carried by
groups of blacksmiths moved independently of any movement of
pottery or crops. The smiths probably did contract work at various
Sorghum history in relation to Ethiopia
155
sites with the local agriculturalist, buying hoes and similar tools from
them. Better tools would have increased the area under cultivation.
Much more remains to be discovered; but clearly race kafir spread
into suitable agricultural areas as the cultivating population of
southern Africa increased. Introgression with the local wild germplasm must have been occurring all the time.
One of the marker crops of this movement was the yellowflowered niger seed or noog (Guizotia), which occurs sporadically
from Ethiopia to Malawi. Niger seed is rapidly disappearing south of
Ethiopia. Finger millet was another crop of the complex carried south.
The Sudan
The Nuba Mountains were probably occupied early on. Settled agriculture there is old, with terracing on the steeper slopes and
animal manure with crop residues in use on the fields. In the hills, the
Nuba have adapted the natural defences of their rocky outposts by
building houses in remote areas and surrounding them with stone
walls. From a distance, southern Nuba hill communities resemble
medieval European castle fortresses built of stone. Cultivated
sorghums show much diversity and the crop is ancient there, as is
sesame (Bedigian & Harlan, 1983). The same is likely to be true for
other suitable hill sites in the Sudan.
The passage to India
The movement of agriculture to Arabia must have occurred at
an early date. The old terraced agriculture, using the same crops, is
still practised there. The overland routes from Saba into Asia are very
ancient and are based on camel caravans. Imports from East Africa
and Somalia usually came to India through Aden. Sea traffic was also
important. There is evidence of sea trade between the Kulli culture of
South Baluchistan and early dynastic Sumer soon after 2800 BC. The
port of Dilmum (probably Bahrain Island) traded extensively along
the coast, possibly as far as Lothal in the Gulf of Cambay. Before the
turn of the era, there was trade between the port of Dhufur in Saba,
and India. Frankincense and myrrh from Saba were highly valued
and spices from India were prized in Arabia and west Asian
countries.
Agriculture in the Indian sub-continent
The Great Indian Desert forms a barrier to movement into
India from the west. The route to the north of this desert goes from
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H. Doggett
the North-West Frontier across northern Punjab to the Ganges and
Jamuna rivers. That to the south skirts between the desert and the
Rann of Kutch and links Sind to Gujarat, Malwa, southern Rajasthan,
Maharashtra and peninsular India. There were important ports for
the coastal traffic at Lothal and Rangpur in the Gulf of Cambay.
Agriculture moved into the north from Iran and was based on the
west Asian crops, with the addition of cotton. The first settled agricultural communities date to the period 8000-5000 BC.
Peninsular India
In the southern Deccan there was an independent indigenous
culture prior to 3000 BC and so contemporary with the urban phase of
the Harappan civilization. A neolithic culture developed in
Karnataka, the first phase of which dated to the period ca. 30002000 BC. From then on, more permanent settlements were discovered,
often located on the crowns and slopes of granitic hills, dated
between ca. 2100 and ca. 1700 BC. There was a terraced agriculture,
with dry stone retaining walls for the terraces. Kodekal and Utnur are
representative of the first period. The people there had domesticated
cattle with sheep and goats. Many rubbing stones and querns were
found, indicating either seed collecting or grain cultivation. The
second period occurs at Utnur, but more information has come from
Piklihal and Hallur, dating from ca. 2500 BC and ca. 2200 BC, respectively, and continuing until the early Iron Age. In the second period,
circular hutments of daub and wattle on a wooden frame were found,
with mud floors. Tool types in the third period are reminiscent of
those found in the Banas culture at the Malwa and Maharashtra sites.
Grinding stones were found, with mullers, as well as large pots,
buried up to their necks, which probably served as storage jars. Cattle
raising continued and crops grown probably included finger millet.
This was identified in Karnataka at Tekkalaksta I. The dates lie
between 2100 BC and 1500 BC (Allchin & Allchin, 1982).
The ancestry of the earliest settlements of the southern Deccan was
independent of those further north. They date to the end of the third
millennium BC and a range of food grains was cultivated. Allchin &
Allchin (1982) noted that local variations in grain utilization at the
present day were already reflected during the Neolithic-Chalcolithic
period. They wrote: 'It is difficult to believe that the Dravidian
languages do not owe their origin to the same people who produced
the neolithic culture there/ Certainly today finger millet is strongly
associated with the Dravidian speakers. Sankalia, Deo & Ansari
Sorghum history in relation to Ethiopia
157
(1971) and Dhavalikar (1979) postulated four claimants as the originators of these cultures: (a) immigrants from west Asia or Iran; (b)
aboriginal tribes who were chased up into the hills by the Aryans ca.
1000 BC; (c) unknown indigenous people who merged completely
with the Sanskrit peoples; (d) a primitive indigenous people from
western Asia who developed these early farming communities. To
this list might be added the possibility of immigrant agriculturalists
from the Horn of Africa, carrying the seeds of their crops with them.
Southern and central Indian agriculture has a strong component of
the Ethiopian crops - finger millet, sorghum, niger and cowpea.
Harlan (1969) noted the resemblance between the Ethiopian emmer
wheats and the southern Indian 'Khapil' wheat. Both have more than
two vascular bundles in the coleoptile. Niger is widely grown among
the Tribals; there were 427 different tribes in India in 1961, with 255
languages. Not all are primitive. Some use a terraced agriculture with
finger millet and niger seed as major crops. Sesame might well have
arrived in the south from Ethiopia, rather than from west Asia.
The following comment of Seegeler (1983) is worth mentioning: Tt
is curious to note an old tradition of the region of Wolcait reported by
Baldrati (1950). According to this legend, an Ethiopian queen had
occupied a vast territory in India in the very remote past. She made
groups of Ethiopians emigrate to India. How far this story has historical background is unknown, but it is striking that there are so many
similarities between the crops of the traditional agriculture in Ethiopia
and India, and that there are groups of Jaferbad in Kathiawar who
consider themselves to be of Ethiopian origin/
One of the oldest of the Afroasian language group's crops must
surely be sesame. Bedigian & Harlan (1983) recorded the ritual use of
this crop for births, marriages and deaths among the peoples of the
Nuba Mountains in the Sudan. The same is true for the speakers of
the Chadic languages in West Africa, for the peoples of Nepal, and
for the Tamils of southern India, where, to this day, sesame seeds are
placed in the mouths of the dead.
References
Allchin, B. & Allchin, R. (1982). The Rise of Civilisation in India and Pakistan.
Cambridge University Press, Cambridge.
Arkell, A. J. & Ucko, P. J. (1965). Review of predynastic development in the
Nile Valley. Current Anthropology, 6, 145-56.
Bedigian, D. & Harlan, J. R. (1983). Nuba agriculture and ethnobotany, with
particular reference to sesame and sorghum. Economic Botany, 37, 384-95.
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H. Doggett
Clark, J. D. & Stemler, A. B. L. (1975). Early domesticated sorghum from
central Sudan. Nature (London), 254, 588-91.
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1181-9.
de Wet, J. M. J., Harlan, J. R. & Price, E. G. (1970). Origin of variability in the
spontanea complex of Sorghum bicolor. American Journal of Botany, 57, 704-7.
de Wet, J. M. J., Prasada Rao, K. E., Brink, D. E. & Mengesha, M. H. (1984).
Systematics and evolution of Eleusine coracana. American Journal of Botany,
73, 550-62.
Dhavalikar, M. K. (1979). Early farming cultures of Central India and early
farming cultures in the Sudan. In: D. P. Agrawal and D. K. Chakrabarti
(eds), Essays in Indian Protohistory. B.R. Publishing Corporation, New
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Hutchinson (ed.), Essays on Crop Plant Evolution. Cambridge University
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History, 20, 161-72.
Fairservis, W. A.,Jr (1971). The Root of Ancient India. Allen and Unwin,
London.
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Bantu language and African culture in situ: an archaeologist's perspective.
South African Archaeological Bulletin, 33, 107-16.
Hallpike, C. R. (1970). Konso agriculture. Journal of Ethiopian Studies, 8, 31^3.
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309-14.
Harlan, J. R. & de Wet, J. M. J. (1972). A simplified classification of cultivated
sorghum. Crop Science, 12, 172-6.
Harlan, J. R. & Stemler, A. B. L. (1976). The races of sorghum in Africa. In:
J.R. Harlan, J.M.J. de Wet and A. B. L. Stemler (eds), Origins of African
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of Eleusine coracana spp. coracana. American Journal of Botany, 66, 330-3.
Kajale, M. D. (1977). On the botanical findings from excavations at
Daimabad, a chalcolithic site in western Maharashtra, India. Current
Science, 46, 818-27.
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Krzyzaniak and M. Kobusiewicz (eds), Origin and Early Development of the
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Sorghum history in relation to Ethiopia
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Krzyzaniak, L. (1984). The neolithic habitation at Kadero (central Sudan). In:
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africana in Uganda. Journal of the Indian Botanical Society, 41, 531-9.
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Hyderabad.
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11
Prehistoric Ethiopia and India:
contacts through sorghum and millet
genetic resources
K.L. MEHRA
Introduction
Several publications have dealt with India's cultural contacts
with western, central and south-east Asian countries, but little information is available on India's contacts with African countries
(Asthana, 1976). This is understandable because little archaeological
work has been done on Neolithic to Iron Age sites in Africa, compared with Asia. Even in India, where several Neolithic-Chalcolithic
sites have been excavated, archaeologists have continued to look for
some kind of west Asian similarity/influence in interpreting their
findings. Thus, even (a) the finds of human skeletons showing
Hamitic-negroid features associated with the Langhanag (Gujarat)
microlithic culture (Sankalia, 1962); (b) terracotta head-rests discovered in Neolithic burials at Narsipur (ca. 1800 BC), Hammige, Hallur (ca. 1800 BC) and Paklihal in the Kaveri and Krishna basins,
showing affinity with similar objects found in Africa and Egypt
(Nagarajarao, 1975); and (c) archaeological finds of African crop
plants (Vishnu-Mittre & Savithri, 1982) have been ignored. Evidences
for indigenous origin(s) of few, or even several, NeolithicChalcolithic cultures of India have been recently discussed but with
bitter controversy (Possehl, 1982). African millets were incorporated
into the cropping system of Chalcolithic farming communities of
India, and these may provide evidence of contacts between India and
Ethiopia where agriculture was practised (ca. third millennium BC).
This chapter deals with recently published archaeobotanical findings
pertaining to Ethiopian cultural contacts with prehistoric India
Prehistoric contacts between Ethiopia and India
161
through the contribution of millet genetic resources. Some aspects of
origin, domestication and evolution of millets in Africa and the
impact of millet introduction on the agricultural history of India are
also highlighted.
Origin, domestication and evolution
Finger millet
Two main groups of cultivars are recognized: (a) the African
highland type, which originated in Africa from its wild progenitor,
Eleusine africana, through selection for plump grains and non-shattering habit, is adapted for cultivation in the highlands, and has long
spikelets, long glumes and grains enclosed within the florets; (b) the
Afro-Asian type, which is adapted for growing in the lowlands, has
short spikelets and short glumes in which the mature grains are
exposed distally, and which evolved from the highland type through
selection of plants with short glumes and exposed grains. This view is
supported by the genetics of glume length. Glume length factors G-l,
G-2 and G-3 are responsible for glume length, with any one or no
factor producing long, any two factors expressing medium-long and
all three factors together producing short glumes (Ayyanger & Warrier, 1936). The African highland type is split into the races coracana
(most primitive), elongata, plana and compacta (de Wet etal., 1984), the
last three races being products of natural and human selection under
domestication, to suit the crop to diverse farming systems and ethnic
preferences. The Afro-Asiatic type is also designated as race vulgaris
(de Wet et al., 1984). All these races occur at present in Africa and in
India. A distinct East Indian group is also recognized (Hilu & de Wet,
1976) and clinal variation occurs among cultivars in India, with
extremes showing a decrease in south Indian influence and an
increase in eastern Indian influence (Hussaini, Goodman & Timopthy, 1977). Indian cultivars also show variation in number of days to
maturity and plant height in relation to increase in latitude, longitude
and altitude, with north-east Indian cultivars being short statured
and of late maturity (Kempanna, 1975).
Introgressive hybridization occurs between E. africana and all races
of cultivated finger millet (African highland and Afro-Asiatic types)
and also between races, wherever sympatric, producing rich variation, in the highlands of Ethiopia southwards to Zimbabwe (Mehra,
1962; de Wet et al, 1984; Apparao & Mushonga, 1987). Stabilized
weedy derivatives of introgressive hybridization between cultivated
and wild types resemble cultivated kinds, except being free-shatter-
162
K. L. Mehra
ing, and occur in disturbed habitats and cultivated fields. Such
hybrids involving E. africana have not so far been reported from India.
In crosses between long glume x long glume types, some hybrids are
likely to have one or two glume length factors but such medium-long
types (with two factors) are likely to segregate or, when crossed with
long-glumed E. africana, would again produce the long glume type.
Thus, evolution of the short glume Afro-Asiatic type would take a
longer time in the African highlands in the presence of E. africana as
compared with the time it would have taken in India, from where this
progenitor has not been reported. It seems more likely that the major
part of the evolution of the Afro-Asian type occurred in India from
where it was taken to East Africa along with durra sorghums during
the Islamic expansions. Thus, India first received the African highland type from Ethiopia, and from this the Afro-Asiatic type was
developed in India.
The Ethiopian highlands (Mehra, 1963a,b), the eastern Sudan
zone, the highlands stretching from Ethiopia to Uganda (Harlan,
1971) and the highlands of East Africa (Hilu & de Wet, 1976) have
been suggested as likely areas of finger millet domestication.
Cultivated finger millet (race plana of the African highland type) has
been identified from archaeological remains found in Axum, in the
northern highlands of Ethiopia (ca. third millennium BC: Hilu, de Wet
& Harlan, 1979) and in Kadero, Sudan (ca. second half of the fourth
millennium BC: Klichowska, 1984). Since the sample belonged to race
plana, an advanced race, the domestication of finger millet must have
occurred much earlier (de Wet et ah, 1984). Finds of finger millet in
India have been reported from Hallur, Karnataka (ca. 1800 BC);
Paiyampalli (ca. 1400 BC), Eastern Ghats; Daimabad (ca. 1400-1100 BC,
topmost Jorwe Ware) and Songaon (wild, 1290 BC) in Maharashtra;
and Surkotda, a Harappan site (ca. 1660 BC), associated with sherds of
the white-painted black and red Ware of the Ahar I culture (Kajale,
1974, 1977; Allchin & Allchin, 1982; Vishnu-Mittre & Savithri, 1982).
Thus finger millet, of the African highland type, seems to have been
domesticated in the fourth millennium BC in the Sudan-Ethiopian
region and it began to be grown in India from about 4000 BP.
Pearl millet
Pearl millet (Pennisetum americanum subsp. americanum) was
domesticated, in a diffuse belt extending from Sudan to Senegal
(Harlan, 1971), from its wild progenitor subsp. monodii, with which it
produces a weedy derivative subsp. stenostachyum, in western Sudan,
Prehistoric contacts between Ethiopia and India
163
northern Nigeria and western Senegal (Brunken, 1977; Brunken, de
Wet & Harlan, 1977; Marchais & Tostain, 1985). Four distinct races,
typhoides, nigritarum, globosum and leonis are known, but only race
typhoides was introduced into India. Racial variation in West Africa is
due mainly to incorporation of genes from closely related wild taxa
with which pearl millet hybridizes (Billiard et al., 1980), but in the
absence of wild relatives in India such a system does not operate.
Variation among populations in India is mainly due to natural and
human selection under domestication. Thus Indian collections are
much less variable than African.
Finds (charred lumps) of pearl millet from archaeological sites in
India are from Rangpur III (1700-1400 BC), Saurashtra, associated with
Lustrous Red Ware (Ghosh & Lai, 1963; Rao, 1963). It seems to have
been introduced into India around 4000 BP.
Sorghum
Harlan (1971) proposed that domestication of sorghum took
place first along the broad band of savannah between the Sudan and
Nigeria. The first domesticated sorghums were bicolor-like and those
originated from its progenitor ssp. arundinaceum race verticilliflorum.
From this region sorghum culture spread to tropical West Africa
where race guinea developed; to Southern Africa where race kafir
developed; to the Sudan-Uganda area where race caudatum developed; and to Asia and Ethiopia where race durra developed, respectively, through hybridization with different close relatives. Durra
sorghums probably developed outside Africa from the race bicolor
that was introduced into Sind, Punjab and North-West India (Stemler
etal., 1977) and the developed durras were reintroduced into Ethiopia
and Arab colonies in East Africa during the Islamic expansion.
Of these sorghums, bicolors were the first to be incorporated in
Indian agriculture, followed probably by guineas, especially S. roxburghii adapted to coastal areas. Next to develop were durras, but
caudatums and kafir were not brought into India.
Archaeological finds of sorghum in India are from:
1. Ahar, Rajasthan, along with rice (ca. 1500 BC: Vishnu-Mittre,
1969), associated with Northern Black Polished Ware;
2. Daimabad, near Pravara river, Maharashtra, late Harappan
phase (ca. 1800-1500 BC: Vishnu-Mittre & Savithri, 1982), and
another sample, topmost Jorwe phase (ca. 1400-1100 BC:
Kajale, 1977);
3. Inamgaon, near Ghod river, Maharashtra, in the early phase
164
K. L. Mehra
of habitation (ca. 2000-1600 BC, associated with Malwa Ware:
Allchin & Allchin, 1982), and another sample associated with
Jorwe Ware (ca. 1100-800 BC, calibrated by the radio-carbon
method: Vishnu-Mittre, 1974), and recent finds of sorghum in
Sangrur district, Punjab (2nd millennium BC: Saraswat, 1986).
These dates are the latest so far known, but sorghum may have been
incorporated into Indian agriculture around 4000 BP.
Mehra (1963b) proposed that finger millet was taken along the
Sabaean Lane route from Ethiopia to India during pre-Aryan times.
Archaeological finds of sorghum (sherd impressions) at Hili, Abu
Dhabi (ca. 2700 BC: Cleuziou & Constantini, 1982) and at Pirak,
Baluchistan (ca. 1900 BC: Constantini, 1979) now confirm that African
millets came to India from Africa through the Sabaean LaneBaluchistan route. Movement through the sea route might also have
been possible because finger millet was identified from Hallur and
Paiyampalli, pearl millet was cultivated at Rangpur III (1700-1400 BC)
and maritime commercial contacts between Gujarat and
Mesopotamia may have existed from at least the beginning of the
second millennium BC (Oppenheim, 1954; During Caspers, 1971).
Discussion
Before the incorporation of African sorghum and millets into
the farming systems of India, the crop-based agricultural economies
of different civilizations/cultures of different regions of prehistoric
India were as follows:
1. The Harappan civilization (ca. 2350-1750 BC) in the plains of the
Indus River system was based on wheat, barley, rape, lentil, pea,
chickpea, sesame and cotton; while in Gujarat, Saurashtra and the
plains of the Yamuna-Ganges rivers (Neolithic-Chalcolithic cultures),
rice was cultivated in addition to these crops (Allchin & Allchin, 1982;
Vishnu-Mittre & Savithri, 1982).
2. Contemporary with the Harappan civilization but independent
of its influence were the Neolithic cultures of Karnataka (30001000 BC), whose subsistence economies were based on Paspalum scorbiculatum and Dolichos biflorus (Vishnu-Mittre, 1969; Kajale, 1977). The
discovery of terracotta head-rests from several sites and finds of
finger millet are suggestive of African contacts and incorporation of
finger millet in the farming system of the region from ca. 1800 BC
(Nagarajarao, 1975).
3. Neolithic-Chalcolithic cultures (2100-1000 BC), late contemporaries of the Harappan civilization in Rajasthan, Madhya
Prehistoric contacts between Ethiopia and India
165
Pradesh and Maharashtra, cultivated wheat, barley, Linum, grass
pea, field pea and lentil in winter and rice, green gram, black gram
and species of Dolichos and Lathyrus during the summer rainy season
(one or more crops at one site: Vishnu-Mittre & Savithri, 1982). The
legumes of Indian origin and African sorghum and millets were
added to the cultivated plants grown in India from the earlier period.
The subsistence economies of Rajasthan, Madhya Pradesh and
Maharashtra were based mainly on minor millets and agro-pastoral
systems. Finds of species of Panicum, Setaria and Echinochloa are from
Surkotda, Maharashtra (ca. 1660 BC: Vishnu-Mittre & Savithri, 1982),
associated with the Ahar culture. When more productive African
millets and sorghum became available for cultivation under different
rainfall regimes and soil types, they rapidly started replacing the
minor millet cultivation, and the subsistence economy gave way to a
food surplus producing economy. African millets can be stored for a
long period without damage in traditional storage systems. They also
produce fodder for cattle besides seed for human consumption. In
areas with higher annual rainfall, sorghum and finger millet were
sown in the summer rainfall season along with Asiatic Vignas, while
in low rainfall areas pearl millet was cultivated. Sorghum and millet
culture moved from Rajasthan to Madhya Pradesh, Maharashtra and
southwards to Gujarat, associated with Ahar, Malwa and Jorwe
cultures and to Gujarat during the late Harappan period. Pearl millet
cultivation was taken up in the drylands of Gujarat and Saurashtra
during phase III of Rangpur, associated with Lustrous Red Ware
(Ghosh & Lai, 1963; Rao, 1963).
African sorghum and millet culture thus played an important role
in the agricultural history of India following the opening up of
opportunities for rainfed agriculture and mixed (agriculture and
animal husbandry) farming systems. This led to a change in the
settlement pattern. Instead of urban centres with neighbouring foodproducing villages, several small villages began to emerge over a
large stretch of the land. The new system progressed rapidly because
the centrally controlled production and distribution system, so
characteristic of the Indus valley civilization, did not operate.
Similarly, the incorporation of pearl millet in the dryland agriculture
of Gujarat seems responsible for the sudden increase in the number
of settlements during Rangpur phases B and C.
African sorghum and millets further evolved under cultivation and
natural selection to suit different ethnic preferences and farming
systems, including the cultivation of sorghum and finger millet dur-
166
K. L. Mehra
ing the winter months in south India and of finger millet in the hilly
areas. But since the wild relatives (progenitors) of sorghum, pearl
millet and finger millet, with which these crops hybridize in nature
even today occur in Africa and not in India, rich variation in several
characters continues to be generated in Africa. Therefore, African
genetic resources of all these crops are more variable than those from
India. Since the wild relatives of these crops are not found in India, it
seems that when these crops were brought into India the seeds of
closely related wild/'companion weedy' species did not accompany
the initial samples of introduction. These crops provide examples of
crop plant evolution in which the evolutionary process has occurred
in Africa in the presence of their wild relatives and in India in their
absence.
We do not, at present, have any non-biological evidence except the
terracotta head-rests for establishing contacts between Africa and
India during the period under reference. The archaeobotanical
evidence is strong, though it needs to be substantiated by future
archaeological findings in East Africa and India. African sorghum and
millets increased the agricultural prosperity of prehistoric India and
continue to do so even today, for which India is grateful to Africa.
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12
Konso agriculture and its plant
genetic resources
J. M. M. ENGELS AND E. GOETTSCH
Introduction
Konso is the name of a relatively small area (approximately
500 sqkm) situated in south-west Ethiopia at a latitude of 5°15'N and
a longitude of 37°30'E, which is populated mainly by the Konso
people. The topography is characterized by rugged and stony highlands, cut by deep valleys that enter into the heart of the country. The
main agricultural area ranges in altitude from 1400 to 2000 m above
sea level and the climate is of the dry montane type with
temperatures ranging from below 15 °Cat night to 32 °Cduring the
day at the hottest time of the year. The Konso Highlands run across
the Rift Valley in an east-west direction and are situated in the dry
belt of Ethiopia with an unreliable rainfall not exceeding 800 mm per
year. There are two rainy seasons: the big rains are concentrated in
March and April and the small rains fall around October and November. In general, the rains come in the form of violent thunderstorms
which seldom last more than two hours (Hallpike, 1972). The Sagan
River forms the eastern and southern borders of Konso, while to the
north the great plains of Gomida and Lake Shamo and, more to the
west, the Gidole mountains and the Woito Valley form natural
boundaries.
The Konso are a small tribe of about 60000 people (Minker, 1986).
Their language belongs to the East Cushitic group (Hallpike, 1970).
The Konso have evolved their remarkable (agri)culture in a high
degree of isolation during the many centuries they have occupied the
area. Their neighbours are mainly pastoralists (e.g. the Borana in the
south) or agriculturalists (among others the Gauwada tribe in the
west) and most of them belong to the Oromos. 'The Konso are
170
/. M. M. Engels & E. Goettsch
markedly shorter and more negroid than the neighbouring Borana.
They are clearly an amalgam, both physically and culturally, in which
other stocks than Galla are represented' (Hallpike, 1972).
The Konso live in densely populated towns, each inhabited by an
average of some 1500 people (Minker, 1986). These towns are surrounded by striking stone walls, with narrow corridors connecting
the walled or fenced homesteads. Within a homestead one can normally find the main round hut, the kitchen, the grinding house,
sleeping huts, stores and one or more stables, all located close
together in order to leave some space for the gardens. Almost all
towns and villages are situated on mountain ridges or on steep slopes
and therefore do not have water. The 6-8 m deep wells are found in
the deep gorges next to the dry riverside and are carefully protected
against floods or dirty river water. During the dry season the women
and girls are frequently occupied for several hours per day in order to
obtain sufficient water (Kuls, 1958).
The agricultural system
The soils in Konso are, in general, of volcanic origin and in
certain parts of the country basalt and tuff layers 100 m or more thick
can be found on top of crystalline formations. The terrain is extremely
mountainous and stony and the paucity of rainfall makes water and
soil conservation of prime importance. The visitor quickly realizes
that the concern for sufficient water for people, cattle and crops has a
dominant place in the daily life of the Konso, and this can also be
frequently observed in the landscaping. The soil is preserved by the
construction of stone terraces, hundreds of kilometres long and often
several metres high. Because of the steep slopes, the terraces are
generally narrow, only a few metres wide. The dry-stone walls are
built along the contour lines of the hillsides and their principal function is to prevent the rainstorms from washing away the soil and
crops. At the same time they assure an adequate supply of water for
the crops by retaining the water within the terraces. In the flatter
areas and in the bigger terraces the land is subdivided into big plots,
up to 9sqm, each surrounded by 10-20 cm high earth walls,
frequently strengthened with sorghum straw in areas where stone is
short (Kuls, 1958). In some parts another practice can be seen where,
in the middle of a terrace, a reinforced hollow some 150 cm wide is
used to concentrate the meagre water supply. The water is distributed through carefully constructed channels and irrigates fields
which are sometimes 100 sqm or more in size. Such terraces are
Konso agriculture and its plant genetic resources
171
separated from each other by carefully constructed stone walls up to
6 m high. The water inlet from the river-bed can be regulated according to the needs of the crop plants. Because of this long lasting and
careful management the soil of the irrigated terraces is generally very
fertile.
One of the striking features of Konso agriculture is the use of
manure, of both animal and, less frequently, human origin (Hallpike,
1972). The connection between animal husbandry (cattle, goats and
sheep) and intensive agriculture is typical of the Konso. Dung is
never used as fuel - as can be observed in many other parts of
Ethiopia - and the practice of stall-feeding (unknown elsewhere in
Ethiopia) all year around is a valuable source of manure. Normally,
cattle are kept on pastures near the periphery of the villages from
where the dung is collected then left to rot in heaps or pits, together
with other organic wastes.
Nearly all the land surrounding the villages is permanently
cultivated, the terraces are richly manured and only a few pastures
are found. Manure is applied not only before sowing, but also during
the growing period of the crops. Since many crop species are interplanted in the same field, crop rotation is not necessary. Only in the
more remote fields where manure is not regularly applied can fallow
land be observed. Because of the terraces and, sometimes, the very
steep slopes, the double-bladed hoe and the less important digging
stick are the most commonly used implements. Ploughing has only
recently been introduced and is not a common practice.
The agricultural activities are determined by the rainfall pattern.
Field preparation starts at the onset of the first rains in January/February and, in general, the fields are worked only once before sowing
(Westphal, 1975). Sowing starts as soon as the big rains begin. The
seeds of the cereals and pulses are mixed, broadcast and lightly
covered with soil. Root and tuber crops as well as cotton seeds are
planted earlier and are carefully protected by soil to prevent them
being eaten by birds. After planting or sowing there is a laborious
period of weeding, and bird and animal scaring as well as protection
against insects. From May onwards the various crops are harvested:
first the roots and tubers, followed by the cereals and pulses and
finally, in mid-September, the sorghum. If sufficient small rains fall,
mainly during October and November, a second sorghum crop can be
harvested in December/January from the ratooned sprouts of the first
season's plants (Kuls, 1958; Hallpike, 1970). From time to time, crop
failures are caused by drought; famine is the unavoidable conse-
172
/. M. M. Engels & E. Goettsch
quence, as was observed in 1984 and 1985. However, because of the
extremely high number of food crop species - domesticated, semidomesticated or wild - even in times of severe drought at least some
food will be available (e.g. Araceae-tubers).
The harvested crops are carefully stored in granaries or kept hanging in the huts. Hallpike (1970) reported that the ashes of burnt cow
dung were used to protect whole sorghum heads against insects in
the granary. The tubers are frequently left in the ground until they are
used.
One important factor for each farmer is that he possesses a piece of
land situated in each of the areas having a different soil type, e.g. in
an irrigable area near to the village, and in a more remote area for the
production of cotton. Kuls (1958) reported an average of at least 10
different fields per farmer, each of them seldom bigger than a quarter
of a hectare.
Plant genetic resources and their uses
The most striking feature of Konso agriculture is the high
number of plant species used and the way they are intercropped (see
Table 1). By integrating multi-purpose trees into this system the
Konso have created an indigenous type of agroforestry, well adapted
to the prevailing dry conditions. Hardly any unused piece of land will
be found around the villages and, throughout almost the whole year,
the soil is covered with crops, crop residues, stones, trees such as
Moringa, Terminalia and Balanites spp., or shrubs (e.g. pigeon pea,
cotton, coffee and yams). Both trees and shrubs are important in
preventing soil erosion.
Some of these plant species demand a careful management (e.g.
Sorghum and the cabbage tree, Moringa stenopetala), whereas others
are wild plants which are tolerated and harvested only in abnormal
years (e.g. some Araceae species). Others are used for medical and/or
ritual purposes. Some of the crop plants in Konso have been grown
since ancient times, e.g. sorghum, Araceae, cabbage tree, cotton and
coffee. The Konso believe that God has given these traditional plants
to their ancestors in the mythological past. In the meantime the
Konso have adapted many crop species from outside. During the
Amharic occupation by the end of the last century important species
such as maize, potato, sweet potato, teff, linseed, pigeon pea, onion,
garlic, chilli pepper, lemon and papaya were introduced. During the
Second World War the Italians introduced, for example, sunflower
and orange. Missionaries and emigrated members of the Konso tribe
Table 1. Plant genetic resources found or reported to be used in Konsoa
Scientific name
Cereals
Amaranthus
caudatus
Eleusine coracana
Eragrostis tef
Hordeum vulgare
Sorghum bicolor
Triticum
durum
Zea mays
Pulses
Cajanus cajan
Local name
Common name or
family name
Cultivation^
Use and remarks
Pasa
Amaranth
(1), 2
Pareja
Kajeta
Boita, poorta
Finger millet
Teff
Barley
1
1
General name: unta
(= cereal). There are
many different names
according to variety
Kaba, kapa
Sorghum, millet
1,(2)
Wheat
1
Paza, pogoloda
Maize
1,(2)
Seeds used for the preparation
of 'dama,c beer. Use of leaves
as a vegetable
Beer, soup, unleavened bread
Cash crop, rare
'Dama, beer, soup, unleavened
bread, seeds sometimes
roasted
Staple food ('dama), beer,
soup, unleavened bread. Stalks
used as fodder; an important
fuel
'Dama, beer, soup, unleavened
bread, seeds sometimes
roasted
Staple food, ('dama), beer,
soup, unleavened bread. Stalks
used as fodder; an important
fuel
Ashakilta, ohota
faranjeta
Pigeon pea
1,(2)
Seeds boiled or eaten raw
Table 1 (cont.)
Local name
Common name or
family name
Cultivation*7
Use and remarks
deer arietinum
Sumpura
Chickpea
1,2
Lablab purpureus
Lens culinaris
Phaseolus lunatus
Okala
Sirota
Bapello
Hyacinth bean
Lentil
Lima bean
1,2
1
1
P. vulgaris
Alkoka
Kidney bean
1, 2
Pisum sativum
Vicia faba
Atara
Bakala
Pea
Horse bean
1, 2
1
Vigna unguiculata
NN
Ohota, okala
Neeqayta
Cow pea
-
1
1,2
Boiled or eaten raw. Ground
for flour in times of famine
Boiled or eaten raw
Boiled
Seeds boiled or eaten raw
when young, ground for flour
in times of famine
Seeds boiled or eaten green.
Ground for flour in times of
famine
Boiled or eaten raw
Seeds boiled or eaten green.
Ground for flour in times of
famine
Boiled or eaten raw
Small green bean
Scientific name
Tubers
Amorphophallus abyssinicus
Arisaema sp.
Saganeida
Burie, lameeta pakana
1,(2)
Indian turnip
1,(2)
Used to make 'dama in times of
famine since these tubers can
survive considerable drought.
Also used to make beer
Contain a poisonous substance
which has to be eliminated
during preparation
Sauromatum
nubicum
Colocasia esculenta
Pansala
Longa
Taro
1/(2)
1
Dioscorea abyssinica
Hidana
Yam
2,3,5
Ipomoea batatas
Manihot esculenta
Dinitscha faranjeta
—
Solanum
Tinassa
Sweet potato
Cassava
Irish potato
2
1,2
1,2
Kaguta
Tuma tima
Tuma ata
Passifloraceae
Onion
Garlic
4(?)
2
2
Brassica carinata
Gomano
Ethiopian mustard
(1), 2
Cucurbita pepo
Potota
Pumpkin
(1), 2, 5
Digera alternifolia
Kogata
Hangoleita
Njannja
Amaranthaceae
Compositae
Tomato
1,2
4
2
Shelagda, tellakata
Halako
Shiferaw
Cabbage tree
(1), 2
tuberosum
Tubers used to prepare 'dama.
Leaves used to flavour beer
Tubers boiled and eaten or
pounded and made into 'dama
Tubers boiled and eaten
Tubers boiled and eaten
Tubers boiled and eaten
Vegetables
Adenia ellenbeckii
Allium cepa
A.
sativum
Launaea taraxacifolia
Lycopersicon
esculentum
Moringa stenopetala
Leaves used as a vegetable
Bulbs used as a vegetable
Bulbs used as a condiment/
vegetable
Leaves are an important
vegetable, seeds are also used
Fruits and leaves are consumed
young
Leaf vegetable
Wild growing vegetable
Occurs naturally, fruits are
eaten
Leaves are a very important
vegetable; eaten boiled with
'dama. Leaves especiallv
py
important during the dry
season, medicine
Table 1 (cont.)
Local name
Common name or
family name
Pergularia daemia
Korroda
Asclepiadaceae
Portulaca quadrifida
NN d
Mereita
Kulbabita
Portulacaceae
NN
Xagalaa
NN
NN
Kutata
Rasota
Scientific name
Spices
Capsicum annuum
Parpara (red)
Qaara (green), mitmita
Tibichota
Cultivation^
Use and remarks
Konso gomen
2,5
2
Leaf vegetable, elsewhere as
edible fruit
Leaf vegetable
Herbaceous 30 cm high plant.
Succulent, leaves are eaten as a
vegetable
Climbing, herbaceous plant.
Leaves are eaten
Leaves are eaten
Leaves are eaten
Red pepper
1,2
1
2
1
—
Coriandmm sativum
Foeniculum vulgare
Linum usitatissimum
Talpa
Coriander
Fennel
Linseed
Ocimum spp.
Iffaya
Basil
2
Buckthorn
2
Rhamnus vrinoides
—
Fruits eaten fresh and used to
flavour food
Seeds used to flavour food
Seeds used as a condiment
Seeds used to flavour food,
also to prepare a kind of dough
Leaves are used to flavour
food, very common
Leaves and wood used as a
condiment to flavour alcoholic
beverages
—
Rue
2
Seeds and leaves used to
flavour food
Carthamus tinctorius
_
Safflower
3
Helianthus annuus
Sufeta
Sunflower
1/(2)
Ricinus communis
—
Castor bean
2
Seeds are consumed and used
for oil
Seeds roasted, also infused and
liquid drunk
Oil used for lighting and for
softening leather
Aureta
Papayata
Loomet
—
Malvaceae
Papaya
Lime
Orange
Tiliaceae
Mulberry
Banana
Anacardiaceae
Prickly pear
4
2
2
2
4
2
2
4
2,5
Saccharum officinarum
Vangueria madagascariensis
Sonkara
Murganta (?)
Sugar cane
Rubiaceae
1,2
2,4
Ximonia coffra
Ziziphus spina-christi
Inginkada
Olacaceae
Christ thorn
4
2
Ruta chalepensis
Oil crops
Fruits
Azanza garckeana
Carica papaya
Citrus aurantifolia
C. sinensis
Grewia tenax
Morus mesozygia
Musa paradisiaca
Rhus natalensis
Opuntia ficus-indica
—
Kotjata
Inch'orre
Museta
Kabudeida
—
Edible fruit
Edible fruit
Edible fruit
Edible fruit
Edible fruit
Edible fruit
Edible fruit
Edible fruit
Fruits are greatly enjoyed by
children; frequently used in
fences and as fodder
Shoots are eaten raw
Wild tree, edible fruits; planted
in villages
Edible fruit
Edible fruit
Table 1 (cont.)
Scientific name
Local name
Common name or
family name
Cultivation*7
Use and remarks
2,4
Wild tree, edible fruits; planted
in villages
Edible fruit
NN
Kenenta
NN
Maderta
-
2
Punitta
Coffee
1,2
Beans are roasted with butter
and cereals; leaves used to
prepare a tea. Important cash
crop. Also used for ritual
purposes
Garatita
Futota
Cotton
1,(2)
G. herbaceum is the older
introduction, G. hirsutum gives
higher yields. Basis of the
important weaving industry.
Cash crop
Catha edulis
Teemahada
Chat
2
Nicotiana tabacum
Tampota
Tobacco
2
Leaves chewed for ritual
purposes
Leaves fermented and smoked
Beverages
Coffea arabica
Fibres
Gossypium herbaceum
G. hirsutum
Narcotics
Miscellaneous
Balanites aegyptica
Hangalta
4,5
Leaves used as browse for
cattle and sometimes as
vegetable. Fruits are eaten
Boswellia rivae
Dangarda itana
Incense tree
4
Commiphora sp.
—
—
2,4,5
Ensete ventricosum
Dupana
False banana
2
Hyparrhenia spp.
-
Lagenaria siceraria
Dahanta
Thatching grass
Gourd
4
1,2
Solarium incanum
Kimbilota
Weybata
Sodom apple
4
1
Terminalia brownii
Lia
a
Resin used in ceremonies;
exported
Tree (often living) is important
to build strong fences,
especially in villages
Starch of pseudostem used as
staple
Thatching of roofs
Fruits produce important
containers
Used for ritual purposes
Widespread cultivated tree,
important timber; leaves are
harvested as browse for cattle
The information is mainly compiled from Goettsch et al. (1984) and Westphal (1975). In addition, Hallpike (1970) has reported the use of
80 wild plant species and trees for food, animal fodder, medicine, building material, magico-rituals and miscellaneous.
b
Key for 'Cultivation': 1, field; 2, backyard in village; 3, border of terraces in fields; 4, 'wild' plant; 5, fences.
c
'dama: sorghum or other starchy products are ground, kneaded into balls and boiled in water.
d
NN = Not known.
180
/. M. M. Engels & E. Goettsch
now living in Kenya introduced new bean varieties. Of the more
recently introduced species, maize is gaining more and more importance at the expense of sorghum (Hallpike, 1970; Minker, 1986).
In the following section, relevant agronomic and botanical details
are given of the important crop plants as well as of some of the
striking species which are typical and traditional in Konso.
Cereals
Sorghum (Sorghum bicolor)
The major staple crop of the Konso is sorghum (Sorghum
bicolor). It is believed to be an ancient crop of the region (Doggett,
1988) and a wide spectrum of varieties exists. Harlan & Stemler (1976)
report an 'unusual assemblage of sorghums' in a few small areas of
Africa. One such region is Konso. The guinea sorghum, one of the
major races, is dominant in West Africa but is also found in Konso
(Stemler, Harlan & de Wet, 1977). Hallpike (1970) mentions that at
least 24 varieties are grown and he listed the names of 17 which are
distinguished according to their uses: ground, then boiled in water in
the form of kneaded balls - 'dama - to be served with boiled cabbage
tree leaves, or made into beer, sour or unleavened bread for travelling
(Hallpike, 1970). Two types of beer are produced, normal beer and an
unmalted variety, named 'erorda'. The Konso are able to describe
each sorghum variety very clearly and some examples are given here,
taken from Hallpike (1970).
1. yedoda
not much good for beer or 'dama. Bitter,
husks difficult to remove. Quick ripening.
Birds do not eat it much.
2. sulida
'dama and beer are good. Beer lasts one
month. Soup and erorda also good. Its
husks are rather difficult to remove. Birds
like it. A slow grower - five months.
3. ha dida
'dama good, but beer is not, goes bad in
two days. A bitter grain. Birds eat it, and it
is rather a slow grower.
4. kulsida
'dama good, erorda good, and the grains
are good roasted. Beer not good, goes off
in two days. If grubs eat it, becomes bitter.
Birds eat it. Quite a quick grower.
5. tisgara
'dama good, erorda good, and good
roasted. Beer is good, lasts two weeks.
Birds cannot get at it since the heads hang
down. Slow grower.
Konso agriculture and its plant genetic resources
6. harboreda
181
not good for erorda or beer since it has
nasty taste. Not much better for 'dama.
Birds eat it. Grows very quickly,
7. hargiti
good for all purposes. Beer only lasts a
week. Birds eat it. Very quick grower,
8. 'gonada
very nice for all purposes. Beer lasts two
weeks. Birds eat it. A slow grower,
9. ken dera
very nice for all purposes. Beer lasts two
weeks. Birds like it. A quick grower,
10. obiyada
quite nice for all purposes. Beer lasts two
weeks. Birds eat it. A slow grower,
very nice for all purposes. Beer lasts two
11. rereda
weeks. Birds eat it. A slow grower,
nice for 'dama, erorda not very nice. Beer
12. hoiriada
is nasty. It is only eaten if unaffected by
grubs. Birds cannot reach grains. A slow
grower.
only 'dama is made from it (not used even
13. magaloda
for this if attacked by grubs). Never used
for beer as far too bitter. Immune to korba.
Birds eat it. A slow grower,
14. 'gamadeda sira nice for all purposes. Beer lasts one
month. Birds eat it. Rather a slow grower,
good for 'dama if grubs do not spoil it. Not
15. ongo/uwada
nice for beer, and erorda is not much good
either. Birds eat it. A quick grower,
nice for all purposes. Beer lasts a week.
16. o jara
Birds eat it. Slightly slow grower,
very nice for 'dama and beer. Beer lasts
17. pijita
two weeks. Birds cannot get at heads
which hang down. Rather slow grower.
The following characteristics of the sorghum 'varieties' are considered to be relevant in deciding which of these varieties to grow:
- taste;
- suitability for 'dama/erorda beer;
- growing time (e.g. earliness);
- effect of grubs on taste;
- effect of korba (a fungal disease);
- difficulty of removing husks;
- bird resistance.
From several recent collecting missions organized by the Plant
Genetic Resources Centre/Ethiopia to the Konso region it was con-
182
/. M. M. Engels & E. Goettsch
eluded that some of the varieties described by earlier workers do not
exist any more and systematic collecting of indigenous germplasm is
therefore an urgent matter.
Finger millet (Eleusine coracana)
Finger millet is another traditional crop in Konso and it can
almost always be found interplanted with sorghum, especially at
lower altitude. The grain of finger millet is mainly used for the
preparation of beer, but also for soup and bread. No details are
known about the diversity of this ancient crop in Konso.
Barley (Hordeum vulgare)
Barley is, next to sorghum, the most important cereal in
Konso and is a traditional component of the cropping pattern. Above
1900 m it is the dominant crop and it is used for the preparation of
beer and food (e.g. dough balls, soup and bread). Although no striking differences from other Ethiopian primitive varieties have been
observed, more systematic collecting and study of the landraces in
Konso is required.
Wheat (Triticum spp.)
The use of wheat is very similar to that of barley and it is
grown under similar conditions. Durum wheat has been collected by
the Ethiopian genebank and during the missions it was frequently
observed that wheat, barley, finger millet and sorghum were grown
in the same field. Minker (1986) has observed that wheat is not as
much favoured by the Konso as barley.
Pulses
The production of pulses plays an important role in the
Konso agriculture. Of the many species reported or collected, horse
bean and the common bean are definitely of pre-Amharic age and
several varieties of both crops are known. Other species commonly
found in the fields or even sometimes in the backyards are chickpea,
garden pea, cowpea, pigeon pea, lentil, mung bean, hyacinth bean
and, more rarely, lima bean. Pulses are eaten raw or boiled and in
times of famine the seeds are ground and used as flour.
Tuber crops
The cultivation and/or use of root and tuber crops in Konso
has a long tradition. They are crushed, ground and formed into flat
Konso agriculture and its plant genetic resources
183
bread or dough balls. Taro and yams are known from other areas in
Ethiopia as well, but the use of wild Araceae is almost entirely restricted to Konso and these species play an important role in the
mythology of the region (Hallpike, 1972). In normal years the tubers
are left in the ground and only in times of food shortage are they
harvested. As a result, famines in Konso are less severe compared
with other areas in Ethiopia. The number of tubers left in the ground
plays an important role when a field is sold (Minker, 1986).
'Pakana' is the common name for the three different Araceae species. The specific local and botanical names for each of the species are:
Pansala:
Sauromatum nubicum
Saganeida: Amorphophallus abyssinicus
Lameeta:
Arisaema (? schimperianum) (Indian turnip)
Amorphophallus is widespread in the villages as well as in the
cultivated fields and their borders. Since its tubers contain a toxic
substance they have to be crushed and exposed to the air for oxidation. After this procedure the product can be consumed without any
problem. Because of the limited knowledge existing on these species
a thorough collection and study would be worth while, also in respect
to its potentialities for other dry areas in the country.
Cabbage tree (Moringa stenopetala)
The most striking characteristic of the Konso agricultural
system is the cultivation of the cabbage tree. The tree is densely
planted within the villages and generally more widely spaced in the
fields and terraces between 1600 and 1800 m. Its light green leaves
and the conspicuous grey bark are characteristic features of the cabbage tree.
Konso can be considered as the area where the tree was first
cultivated. From here the cultivation has spread into neighbouring
areas where it is being used intensively as well. In the whole region
the cabbage tree does not occur in the wild (Minker, 1986). The tree is
raised from seed; it requires relatively good soil conditions and prefers wind-protected places. After 5-6 years the first leaves can be
harvested. They are boiled and eaten as a vegetable with any warm
meal. The leaves are rich in vitamins and are mainly harvested in the
dry season when other vegetables are scarce. During the rainy season
there are only a few leaves left on the tree and they do not taste good.
The leaves are an important trading product in the local markets.
Outside the villages, especially on the terraces, the cabbage tree
plays an important role in reducing soil erosion. There are trials
184
/. M. M. Engels & E. Goettsch
under way to use the tree for this purpose in other areas of the
country. Furthermore, some very promising medicinal uses have
been found. A tea of dried leaves is reported to be very efficient in
treating light cases of diabetes and it is said that the extract of fresh
leaves can cure indigestion and even the cure of an amoebic dysentery has been reported. Eye inflammations are treated and a root
extract helps against unconsciousness (Aschalew Hiude, personal
communication). Recently it has been reported that ground seeds can
be used to clarify muddy water (Jahn, 1981; Goettsch, 1984). Experiments have shown that this powder has the same effectiveness as the
best technical water clarifying agents.
It can be concluded that Moringa stenopetala is a greatly underutilized and relatively unknown tree which deserves further investigation. It could play a much more important role in the nourishment
of people and in the stabilization of the environment in areas with
limited rainfall in the tropical belt between 1400 and 1900 m.
Other traditional crops
Vegetables are highly appreciated by the Konso (Minker,
1986). The traditional group of vegetables is composed of a number of
species, cultivated or wild, of which the leaves are used. Some of the
Brassica species, such as B. carinata, are among the most important.
The gourd (Lagenaria spp.) is a very old and important plant of
which the ripe fruits are used as containers. Capsicum was introduced
by the Amharas. A number of other spices and herbs are in use. Fruitbearing trees are mainly introduced (e.g. papaya, orange, lime)
whereas traditional fruits are collected from the wild. Coffee growing
has a long tradition and trees can be observed quite frequently
despite the prevailing unfavourable ecological conditions.
Two types of tobacco are cultivated: one is very strong and is said
to be pre-Amharic (Minker, 1986). The Konso are famous for their
woven goods. Several varieties of cotton are grown, and at least one
of them is pre-Amharic (Gossypium herbaceum var. acerifolium: Hallpike, 1970).
The remarkable number of different tree species in the man-made
vegetation of Konso should be mentioned. All the trees are used
somewhere and somehow and a systematic study of their uses would
be extremely interesting and promising. The importance of mixed
cropping under the ecological conditions of Konso cannot be overemphasized. It plays a significant role in the food and fodder production security of the region because of the different water and
Konso agriculture and its plant genetic resources
185
temperature requirements. It also is an important factor in soil conservation. Goettsch, Engels & Demissie (1984) reported 40 different
species in one village and collected 24 species in a terraced field of
about 0.2 ha at an elevation of 1750 m. This is quite exceptional when
compared with other montane areas in Ethiopia.
Conclusions
Considering the difficult agro-ecological conditions which
prevail in Konso, it is remarkable how many people can be fed from a
rather limited area when appropriate farming methods are applied.
The ancient terraces and other constructions, as well as the simple
but efficient irrigation methods, are the salient features of Konso
agriculture which allow an optimal use of water throughout the year.
The intercropping of various crop and tree species together with
the cultivation practices seem to be important factors in food and
fodder production security as well as in the soil conservation of the
Konso area.
The diversity of crop species and the genetic diversity within many
of the crop species make Konso an important area from the germplasm conservation and exploration point of view.
The cultivation of the cabbage tree as well as of certain tuber crops
is almost entirely confined to the Konso highlands. These species
may have good potential in other similar areas where rainfall is
limited and where, so far, only relatively small numbers of crops are
grown.
References
Doggett, H. (1988). Sorghum history in relation to Ethiopia. In: J.M. M.
Engels (ed.), The conservation and utilization of Ethiopian germplasm.
Proceedings of an international symposium, Addis Ababa, 13-16 October
1986, pp. 97-115 (mimeographed).
Goettsch, E. (1984). Water-clarifying plants in Ethiopia. Ethiopian Medical
Journal, 22, 219-20.
Goettsch, E., Engels, J. M. M. & Demissie, A. (1984). Crop diversity in Konso
agriculture. PGRC/E-ILCA Germplasm Newsletter, 7, 18-26.
Hallpike, C. R. (1970). Konso agriculture. Journal of Ethiopian Studies, 8, 31-43.
Hallpike, C. R. (1972). The Konso of Ethiopia. A Study of the Values of Cushitic
People. Oxford University Press, Oxford.
Harlan, J. R. & Stemler, A. B. L. (1976). The races of sorghum in Africa. In:
J.R. Harlan, J.M.J. de Wet and A.B.L. Stemler (eds), Origins of African
Plant Domestication. Mouton Publishers, The Hague, pp. 465-78.
Jahn, S. al A. (1981). Traditional Water Purification in Tropical Developing
Countries. Existing Methods and Potential Application. GTZ Publication 117,
Eschborn, Federal Republic of Germany.
186
/. M. M. Engels & E. Goettsch
Kuls, W. (1958). Beitrage zur Kulturgeographie der Suedathiopischen Seenregion.
Frankfurter geographische Hefte 32, University of Frankfurt, Federal
Republic of Germany.
Minker, G. (1986). Birji - Konso/Gidole - Dullay. Reihe F, Bremer Afrika
Archiv, Band 22, Uberseemuseum, Bremen.
Stemler, A. B. L., Harlan, J. R. & de Wet, J. M. J. (1977). The sorghums of
Ethiopia. Economic Botany, 31, 446-60.
Westphal, E. (1975). Agricultural Systems in Ethiopia. PUDOC, Wageningen.
Part III
Germplasm collection and
conservation in Ethiopia
13
Theory and practice of collecting
germplasm in a centre of diversity
J.G. HAWKES
Introduction
In the past, collecting activities have concentrated on particular species or on certain genetic characters which plant breeders were
seeking. At present, because of the rapid rate of genetic erosion of
crops in most parts of the world, exploration trips are now becoming
'rescue operations' in which as much diversity as possible is being
collected. The concept of 'now or never' is in the forefront of the
collectors' minds.
The methodology of collecting and the scientific basis of sampling
have also received considerable attention. Whereas some 30 years ago
it was thought sufficient to collect a few seeds from a single plant,
write one or two words on a label and put them all into a bag, this
method is now thought to be most unsatisfactory.
The genetic resources collector is looking for diversity. Whereas
the botanical or horticultural collector was content to collect a few
herbarium specimens and a small packet of seeds, to serve as a
representative for a species in a particular area, the genetic resources
collector needs not uniformity, but diversity. How is this to be
accomplished?
Studies of the population genetics of wild species by Allard (1970)
and his colleagues showed that more sophisticated methods were
needed if a reasonable amount of the genetic diversity of a species
was to be captured. Marshall & Brown (1975), Bradshaw (1975) and
Jain (1975) developed sampling methods for wild species and stated
that such methods should be applicable to cultigens since all these
were held to possess some kind of population structure.
Before we proceed to set out the generally agreed sampling strate-
190
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gies for genetic resources aims, it will be as well to discuss the general
patterns of diversity within a species. These can be partitioned into
three sections.
Firstly, there is the diversity within a species that results from
broad environmental differences acting on it over different parts of its
distribution range. Where environmental changes over great distances are slight, the changes in genetic diversity will also be slight.
Where environmental changes are abrupt, as with changes in altitude
or soil type, correspondingly abrupt changes in genotype may be
expected.
The second type of diversity lies within a small area where a
mosaic of populations results in what is spoken of as interpopulation
diversity.
The third type is to be seen within populations, necessitating proper population sampling techniques.
Naturally, genetic studies of natural populations show differences
between species in the amount of variation between and within
populations. On the whole, inbreeding species should have greater
interpopulation variability and less intrapopulation variation than
comparable outbreeders. Thus if the breeding system of a species is
known, appropriate modifications to sampling methods can be made.
Instruction manuals for collecting genetic resources materials have
been on the market for some time. Several chapters in Frankel &
Hawkes (1975) and in Hawkes (1983) deal with the practical and
theoretical concepts of field sampling, and a comprehensive work by
Mehra, Arora & Wadhi (1981) is also available. A field collection
manual (Hawkes, 1980), dealing with all practical aspects, was
published in 1980.
Before discussing detailed sampling methods, we must remember
that not all crops or wild species are propagated by seed. Many others
are largely vegetatively propagated and special sampling methods are
needed for them. Another special group is formed by the fruit and
nut trees, which will be discussed separately. First, however, we shall
discuss the exploration and collection of annual seed crops and
annual or herbaceous perennial wild species, which also reproduce
by seed.
Sampling of seed crops and wild species
It is generally agreed that sampling on a population basis
should be random or non-selective, since selective sampling will pick
out only those genes with a clear morphological expression in the
Theory and practice ofgermplasm collecting
191
phenotype and will be likely to omit those controlling disease
resistance and other physiological characters.
The standard procedure for a field crop is to take a sample of 50
seeds from each of 50 plants taken at random and to treat the 2500
seeds as a single sample. The plants are sampled by walking up and
down through the rows, taking a seed sample every so many paces,
the distance between plants being determined by the size of the field
and by the reproductive system of the plants themselves. Thus for a
wind pollinated allogamous plant such as maize, the distance can be
quite great. This method, using 50 seeds from each of 50 plants, was
devised by Marshall & Brown (1975); on population genetics theory it
should give us all the alleles in the population, at 95 per cent
certainty, that are present at 5 per cent frequency or more. This
method can also be easily applied to wild species where the collector
can walk through a population sampling as non-selectively as possible. If time is pressing or the population is small, a minimum of 25
plants will still give reasonably good results.
We have just dealt with intrapopulation sampling. The selecting of
interpopulation sampling sites is also important. In annual crops,
where much mixing is probably very common, the sites should be
evenly dispersed. On the other hand, where wild or weedy species
occur in a series of microhabitats, the sites should be clustered. Thus
the clustering or the evenly dispersed sites will cover the geographical range of a species. One must, however, remember that the sampling sites or clusters of sites must be much closer together when
soils, climate, altitude and other environmental features vary rapidly.
This is especially necessary in highly dissected mountain terrain
where a slight difference of altitude or aspect can bring with it a great
difference in soil or rainfall, or changes from tribe to tribe (Harlan,
1975).
Having strongly advocated the necessity for random sampling, I
may be laying myself open to criticism if I do not suggest that some
selective sampling may be useful under certain circumstances. Harlan
(1975) argues positively for selective as well as random sampling.
Bennett (1970) counsels that after taking a random sample the collector should add extra seeds to the sample from extreme or interesting
phenotypes. Marshall & Brown (1975), however, are not completely
satisfied about this. They recommend that if extreme or interesting
phenotypes are found in a population they should form the basis of a
separate sample. Bogyo, Porceddu & Perrino (1980), although recommending random sampling in general, so as to preserve as much
192
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genetic variation as possible, add: 'Certainly when collecting for specific characters, such as disease resistance, dwarf growth types, stem
strength, etc., collectors will not resort to random sampling. Many of
these traits are rare and only large samples will ensure that genes
responsible for these traits will be represented in the collection.'
Whilst I am inclined to agree with Bogyo and his co-workers in
general terms, it must be pointed out that disease resistance is very
difficult to evaluate with certainty in the field. Thus, the absence of
the disease on the crop may mean that conditions were not right for
the spread of the disease when the sample was made. Again, the
presence of a disease infection on a crop might not mean that the crop
was susceptible to all pathotypes of the disease but only to the one
that happened to be present when the collection was made. An excellent discussion of this subject is given by Dinoor (1975).
Several authors recommend taking a large number of rather small
samples (about 25 seeds each from 50 plants per sample) and so
capturing the diversity from different localities (see Brown &
Munday, 1982).
My conclusion must be that no universal sampling system is perfect but that a generalized random sampling system as proposed by
Marshall and Brown is reasonably good for all situations.
The problem of how to capture rare alleles in a population has not
yet been answered satisfactorily. Thus Chapman (1984) concludes
that alleles which would occur at frequencies of 1 in 10 000 or less are
better sought through mutation breeding than in a genebank.
However, if we do not find them by our present sampling methods,
how are we to know that they exist at all? Nevertheless, he concludes
that we would be likely to find such alleles if the genebanks were big
enough (say 30000 samples). This, however, depends very much on
the spread of samples in the bank and although Chapman argues that
as much diversity might be found by sampling a localized area as by
sampling an extended area, it would appear that he bases this argument on rather few data.
After all, the genebank collector is looking not only for a wide
range of variation, but also for diverse kinds of variation. Thus barley
yellow dwarf virus resistance is found only in Ethiopia and, to a
limited extent, in some neighbouring countries. Virus M resistance in
potatoes is found only in one small valley system in north-west
Argentina. Tungro virus disease resistance in rice is found only in a
few accessions of one wild rice species; bacterial wilt resistance in
cultivated potatoes is found only in one area of Colombia in the
Theory and practice of germplasm collecting
193
northern Andes. Such examples could be multiplied many times.
Thus, I am convinced that careful, evenly spaced sampling of a
crop species throughout its distribution area is an essential part of the
work of a germplasm explorer.
Since today we are talking specifically of germplasm collecting in a
centre of diversity (Ethiopia), we should ask ourselves whether any
special rules apply in this case. Quoting Chapman (1984) again, he
says that 'there are no well-defined centres of diversity in any of the
crops'. This seems to me to be patently untrue and based on insufficient evidence. One has only to look at the clear centre of diversity for
cauliflower in Italy, for Brassica carinata in Ethiopia, for potatoes and
maize in the central Andes of Peru and Bolivia, for teff in Ethiopia, for
chilli peppers in Mexico and many other examples, to realize that
there are so many exceptions as to render Chapman's statement
almost meaningless.
Sampling vegetatively propagated crops and wild species
Having discussed seed crops at length, let us now turn to a
completely different problem, that of vegeculture crops. These are
often quite common in tropical areas, particularly in the New World
tropics; however, they are by no means absent from Africa.
Most vegetatively propagated (vegeculture) crops still possess the
capacity to reproduce by seed, or at least they did so before the
advent of plant breeding and selection, and the very primitive forms
and old landraces are probably still fertile. Related wild species retain
both methods of reproduction as useful alternative strategies, according to whether uniformity (asexual reproduction) or diversity (sexual
reproduction) is needed for survival.
In all vegetatively propagated crops that have come to my notice,
the farmers seem to have carried out such strong artificial selection
that the original population structures have virtually disappeared.
For instance, in the ancient centre of diversity of potatoes in the
Peruvian and Bolivian Andes, each market area or district contains
generally not more than 50-100 distinct morphotypes and sometimes
fewer. The number of morphotypes could have been as high as 200 or
more before genetic erosion took place, though we cannot be certain
of this. Each morphotype probably represents a distinct genotype,
but it is possible that some morphotypes may conceal several different genotypes. Adjacent market areas seem to possess much the
same spectra of morphotypes but not identical ones. In other words,
as we move from market area to market area, we see gradual changes,
194
/. G. Hawkes
some morphotypes disappearing and other new ones appearing.
Market areas some distance from each other will differ very clearly in
their morphotype spectrum (Hawkes, 1975).
This knowledge helps us to build up a sampling strategy which is
in fact the reverse of that advocated for sexually reproducing plants.
Thus, for yams, potato, cassava, sweet potato, ensete, Plectranthus
edulis and all other crops of this sort, it is recommended that a selective sampling of every distinguishable morphotype in each area be
carried out. This undoubtedly means that there will be a large number of duplicates, though not quite as many as might at first be
imagined. In the introduction station the collections are all grown
together and all or most duplicates eliminated. It is best always to
save two or three duplicates of each morphotype to avoid accidental
loss or death through disease or pest infection. At this stage it is
generally possible to identify slight differences between what were, at
first sight, thought to have been identical clones.
Where sexual reproduction takes place in a vegetatively propagated crop, and if the crop is largely outcrossing, the seed should be
sampled on a population basis, even though the plants do not constitute a true population. Most vegetatively propagated crops
(perhaps all?) are very heterozygous and release much diversity
when they reproduce from seed; in addition, the seed collection will
represent a sample of the local gene pool. Such seed samples should
be made as a useful parallel adjunct to the clonal collections but
should be given distinct collection numbers.
When collecting the related wild species the objectives are slightly
different. With these, the population structure is still present and
when seed is available the methods used for seed crops can be used.
When seeds are not available it is useful to take a randomly collected
tuber sample from as many plants as possible (say, up to 25) and treat
them all as a single collection. The plants from this should be sibmated in the experimental field or glasshouse and the seeds mixed in
equal proportions from each plant so as to constitute a single sample.
This advice is perhaps often to be regarded as an ideal procedure
which can seldom be realized in practice. Often one finds only one or
two wild plants in any locality, or what seemed at first sight to be a
population proves to be no more than a single clone.
The general advice given to collectors in these circumstances is
'collect what you can, always striving so far as possible to reach the
ideal procedures'. After all, one collection, however inadequate, is
better than no collection at all.
Theory and practice of germplasm collecting
195
Fruit and nut tree sampling
The biggest problem with fruit and nut tree genetic resources
is that many species possess seeds that cannot be stored in the usual
way (recalcitrant seeds). Thus with these materials, whose seeds are
short-lived, the collecting strategy must always be closely related to
storage. Seeds need to be sown directly or sent back to introduction
stations as soon as possible after collecting. If cuttings are taken they
must be stored carefully during the collecting trip to avoid drying out.
These cuttings must be grafted onto prepared rootstocks or rooted by
mist propagation or similar techniques. It must be borne in mind that
every genotype will have to be grown into a permanent bush or tree
unless multiple grafting techniques are employed.
This means that population sampling is not likely to be possible
since it is quite out of the question to grow 2500 plants for every
collection made in the field (i.e. 50 seeds from 50 plants). In any case,
if collecting is being carried out in tropical forests where the number
of species is large but the number of individuals of each species is
very limited, it may be difficult to identify populations. In fact, it has
been found in South-East Asia that no more than one or two mature
individuals are to be found in every 10 hectares.
In Ethiopia, a special collecting and storage procedure has been
devised for coffee, in which 15 berries from 20 bushes selected at
random within a population are collected, and 10 seeds from each of
the bush samples are taken for sowing (Hawkes, Engels & Tadesse,
1986). Of these, three seedlings are planted out in the field. Thus,
each collection will in the end contain 30 plants, which represents a
fairly reasonable sample of the population (Table 1 and Fig. 1).
In general, for wild fruit and nut trees where the numbers of
individuals are scattered and population size is difficult to estimate,
the following procedure is recommended:
Collect seeds from 10-15 individuals over an area of about 2-5 ha
and combine them into a single sample. If random collections can be
made this will be an advantage, but if the individual trees are very
scarce, seeds from each tree must be collected.
If no seeds are available, suckers or budwood cuttings, one per tree
from 10-15 individuals, should be taken. Repeat this procedure at
intervals depending on climatic, altitudinal or soil differences.
For cultivated fruit and nut trees, the first thing to do is to ask the
farmers if they propagate their trees by means of grafted or rooted
cuttings, or by means of seed. If propagation is by cuttings then treat
these as for vegetatively propagated crops. If propagation is by seed,
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/. G. Hawkes
Table 1. Sowing and planting scheme for each population
Code numbers of bushes
sampled
Berries
Seeds extracted
Seeds sown
Seedlings transplanted
Numbering (assuming
collecting number to
be 153)
1
15
30
10
3
2
15
30
10
3
153-1
153-2
3
15
30
10
3
...
...
...
...
18
15
30
10
3
153-3 ... 153-18
19
15
30
10
3
20
15
30
10
3
153-19 153-20
Finish
Bush to be sampled
Route of collector
Fig. 1. Area of coffee population to be sampled. 20 plants
sampled; 15 berries taken from each plant sampled and kept in
separate bag from others; one collector's number given to the
whole population sample of 20 plants.
Theory and practice of germplasm collecting
197
it is likely that each villager will not possess more than one to five
trees in his garden. If this is so, it is best to treat the whole village as
the collecting site and take a random sample from 10-15 individuals,
bulking all these together as one sample. In this case, the trees
throughout the village are to be regarded as a single population. If
seeds are not available, cuttings should be taken in the same way. It
will be advisable to sample as many sites or villages as possible at
scattered intervals throughout the region.
Market sampling
When it is impossible to do field sampling it may be necessary
to collect market samples. This may occur when lack of time or bad
transport facilities prevent the collector getting to the farms. Where
the crop has already been harvested and sent to market, taking of
market samples may suffice. In any case, a visit to the local markets
may give the collector a good idea of what is available in the general
area.
Market sampling has its drawbacks, however, and must be
approached with caution. In the first place, not all cultivars or populations are sent to market since some will be retained purely for home
consumption. Secondly, many of the market samples may be artificial
mixtures. Thirdly, they may be standard bred cultivars of no interest
to the genetic resources collector. Finally, the information for passport data may be insufficient, lacking, or totally wrong and
misleading.
Having said all this, some market sampling may still be valuable,
but will be nowhere near the value of field sampling, where the
plants in their natural or semi-natural states as population and landraces can be sampled by the collector.
Data recording
Minimum data sheets have been advocated by the International Board for Plant Genetic Resources (IBPGR) and these can be
printed and bound up in books of 100 before starting on a collecting
expedition. One side should show essential descriptors and have
tear-off labels at the base; the other side should be used for optional
descriptors (Table 2).
Specialized sheets for particular crops may also be devised but it is
inadvisable to ask for too much information on the data sheet. Either
the collector will not fill in the information, or he will try to do so and
thus spend too much time on this to the detriment of his collecting
198
/. G. Hawkes
Table 2. Notes on field data recording
1. Expedition and organization (name, year, etc.).
(Note: Descriptors 1, 2 and 3 can easily be printed on all collection sheets
before beginning the mission.)
2. Collecting team. The names of all team participants.
3. Collector's name. The name of the leader or taxonomist.
4. Collector's number (note: a single sequence is strongly recommended for
the whole expedition and any others in which the leader may be
participating. The numbers can be stamped in sequence on to the
collecting sheets before beginning the mission).
5. Date of collection (day, month, year).
6. Photograph numbers of specimen, habitat, farm field, etc.
7. Latin names of genus, species, subspecies, etc. (written in full).
8. Vernacular or cultivar name.
9. Precise locality (e.g. political division, province, department, etc., including
compass direction and approximated distance from nearest village or
geographical feature - distance in kilometres along road between two
inhabited places).
10. Latitude (degrees and minutes).
11. Longitude (degrees and minutes).
12. Altitude (in metres above sea level).
13. Types of material (seeds; inflorescences, spikes, panicles, etc.; vegetative
storage organs, such as rhizomes, corms, roots, tubers; whole living
plants; herbarium specimens).
14. *Sample type (whether populations, pure lines or individuals sampled and
whether by random, non-random or both methods).
15. *Status (cultivated, weed, wild).
16. *Sources (e.g. from field, farmer's store, market, shop, garden, wild
vegetation, etc.).
17. ^Original sources of sample (if from market or store, or if obtained from
another genebank, where grown originally. Farmers may know the
source of their material, also).
18. Frequency (a rough estimate of frequency of wild species).
19. Habitat (mostly relevant for wild species).
20. Descriptive notes (scoring of morphological features of interest for the
particular crop or wild species; thus - plant height, branching, etc.,
colour of stems, leaves, flowers, fruit. Amount of diversity in population
and range shown in voucher specimens).
Note: tear-off labels at the bottom of the page with collector's name and
number are to be placed with the collections (these can be printed and
stamped on to the labels before beginning the mission).
In addition, the following specifications may be recorded on the reverse side of
the sheet.
21. Uses (i.e. for human or animal food, medicinal purposes, etc.).
22. Cultural practices (irrigated, dry, with or without fertilizer, fungicides,
pesticides, etc.).
23. Approximate dates of sowing and harvesting (for instance, 'early April to
mid-July').
24. Soil observations (texture, stoniness, depth, drainage, approximate soil
colour if thought desirable to record).
25. Soil pH (if thought desirable to record).
Theory and practice of germplasm collecting
199
Table 2 (cont.)
26. Land form observation (aspect, slope).
27. Topography (swamp, flood plain, level, undulating, hilly, hilly dissected,
steeply dissected, mountainous, other - specify).
28. Plant community (natural vegetation; for wild species).
29. Other crops grown in surrounding fields or in the rotation.
30. Field observations of pest and pathogen infections.
31. Name and address of farmer (if thought desirable to record).
32. Taxonomic identification (fill in later if identity checked by expert).
33. Expert's name and date.
34. Name of Institution and accession number (where these differ from the
collector's name and number).
* Ring appropriate words.
work. When crop genetic diversity is disappearing so quickly, it is
best to push on with collecting genetic resources and add only the
information required by the minimum data sheet.
Duplication of collections
Many of the early collections made in Ethiopia by Vavilov and
his colleagues some 60 years ago have probably been lost. Other
collections made by expatriate explorers may still exist and subsamples of many of these are being returned to the Ethiopian Plant
Genetic Resources Centre (PGRC/E).
Although it would seem logical to 'repatriate' materials in this way
it may not always be necessary to do so. Thus, if Ethiopian materials
are known to exist in other genebanks, are being well looked after,
and arrangements are available for obtaining them when required,
much duplication of effort would be saved simply by leaving them
where they are. The inventories of the other genebanks might indicate that certain samples could be of immediate value to Ethiopian
breeders and such samples could be recalled for immediate use.
However, the rest could be left where they are, providing that the
relevant passport data are available. If no passport data are now
attached to them they could just as easily be put into a 'reserve'
collection in the other genebank rather than use up much needed
space at PGRC/E.
Conclusions
Although we have dealt in this chapter with the theory and
practice of germplasm collecting, it must be remembered that before
200
/. G. Hawkes
setting out into the field, the collector needs to know what has
already been collected by himself and by previous collectors. In view
of the fact that the Centre is now celebrating 10 very successful years
of its existence, a survey of the present situation should be made for
each crop, as follows:
- to assess how many samples are in the genebank;
- to call up information from the data base on where they were
collected;
- to produce hand-plotted maps of distribution of the collection
sites as well as the altitudinal and ecological data associated
with them;
- to identify areas or eco-geographical zones where collections
with useful characters of resistance have been found, such as
insect, virus and fungus resistance, and adaptation or
tolerance to stress conditions. A thorough knowledge of the
literature would be an advantage here;
- to identify blank areas on the maps and check from previous
experience whether the crop may be found in them, and at
what altitudes and ecological zones;
- to sift the international literature for further information on
Ethiopian collections, especially if provenance data are
available;
- to use the information obtained to formulate a coordinated
strategic plan for collecting genetic resources in Ethiopia during the next 5-10 years.
In this way, one can be certain that a major part of the genetic
resources of the Ethiopian centre of diversity will be conserved for
plant breeding use, now and in the future.
References
Allard, R. W. (1970). Population structure and sampling methods. In: O. H.
Frankel and E. Bennett (eds), Genetic Resources in Plants. Blackwell, Oxford,
pp. 97-107.
Bennett, E. (1970). Tactics of plant exploration. In: O. H. Frankel and E.
Bennett (eds), Genetic Resources in Plants. Blackwell, Oxford, pp. 235-7.
Bogyo, T. P., Porceddu, E. & Perrino, P. (1980). Analysis of sampling strategies for collecting genetic material. Economic Botany, 34, 160-74.
Bradshaw, A. D. (1975). Population structure and the effects of isolation and
selection. In: O. H. Frankel and J. G. Hawkes (eds), Crop Genetic Resources
for Today and Tomorrow. Cambridge University Press, Cambridge, pp.
37-51.
Brown, A. H. D. & Munday, J. (1982). Population-genetic structure and
optimal sampling of landraces of barley from Iran. Genetica, 58, 85-96.
Theory and practice ofgermplasm collecting
201
Chapman, C. G. D. (1984). On the size of a genebank and the genetic variation it contains. In: J. H. W. Holden and J. T. Williams (eds), Crop Genetic
Resources: Conservation and Evaluation. Allen and Unwin, London, pp.
102-119.
Dinoor, A. (1975). Evaluation of sources of disease resistance. In: O.H.
Frankel and J. G. Hawkes (eds), Crop Genetic Resources for Today and Tomorrow. Cambridge University Press, Cambridge, pp. 201-10.
Frankel, O. H. & Hawkes, J. G. (eds) (1975). Crop Genetic Resources for Today
and Tomorrow. Cambridge University Press, Cambridge.
Harlan, J. R. (1975). Practical problems in exploration. Seed crops. In: O. H.
Frankel and J. G. Hawkes (eds), Crop Genetic Resources for Today and Tomorrow. Cambridge University Press, Cambridge, pp. 111-15.
Hawkes, J. G. (1975). Practical problems in exploration. Vegetatively propagated crops. In: O. H. Frankel and J. G. Hawkes (eds), Crop Genetic
Resources for Today and Tomorrow. Cambridge University Press, Cambridge,
pp. 117-21.
Hawkes, J. G. (1980). Crop Genetic Resources Field Collection Manual. IBPGR
and Eucarpia, Wageningen.
Hawkes, J. G. (1983). The Diversity of Crop Plants. Harvard University Press,
Cambridge, Massachusetts.
Hawkes, J.G., Engels, J. M. M. & Tadesse, D. (1986). Suggested sampling
and conservation system for coffee in Ethiopia. PGRC/E-ILCA Germplasm
Newsletter, 11, 25-8.
Jain, S. K. (1975). Population structure and the effects of breeding system. In:
O. H. Frankel and J. G. Hawkes (eds), Crop Genetic Resources for Today and
Tomorrow. Cambridge University Press, Cambridge, pp. 15-36.
Marshall, D. R. & Brown, A. H. D. (1975). Optimum sampling strategies in
genetic conservation. In: O.H. Frankel and J. G. Hawkes (eds), Crop
Genetic Resources for Today and Tomorrow. Cambridge University Press,
Cambridge, pp. 53-80.
Mehra, K. L., Arora, R. K. & Wadhi, S. R. (1981). Plant Exploration and Collection. National Bureau of Plant Genetic Resources, New Delhi.
14
A decade of germplasm exploration
and collecting activities by the Plant
Genetic Resources Centre/Ethiopia
ABEBE DEMISSIE
Introduction
The richness of Ethiopia's biological resources is well known.
It has been mentioned by several scientists that the country exhibits
an extraordinary genetic diversity in cereals such as barley (Hordeum
vulgare), wheat (Triticum spp.), sorghum (Sorghum bicolor) and teff
(Eragrostis tef), oil crops such as castor bean (Ricinus communis),
sesame (Sesamum indicum), and other lesser known but potentially
valuable species of plants. Eleven cultivated crop species have been
identified as having their centre of diversity in Ethiopia (Zohary,
1970). Vavilov (1951) indicated that some 38 species are connected
with Ethiopia as a primary or secondary gene centre.
Owing to the potential and uniqueness of the biological resources
of this country, numerous exploration expeditions have been undertaken in the past. The earliest was probably the one made by
Schimper in 1840, a year which appears to mark the beginning of
botanical collecting in Ethiopia (Gentry, 1971). However, it was after
the establishment of the Plant Genetic Resources Centre/Ethiopia
(PGRC/E) that systematic collecting was launched on a large scale.
Agents of genetic erosion
The valuable genetic diversity in Ethiopian crop species, as
well as in their related wild species, has been built up over the centuries by the natural selective forces of the environment and the
farming community. Such diversity exists not only among the different agricultural areas of Ethiopia, but also within each area and
Germplasm exploration and collecting by PGRC/E
203
even within one farmer's field. This wealth of biological and genetic
diversity is seriously threatened by a number of factors, as follows.
Natural calamities
In the last decade there have been several catastrophic
droughts which have led to complete crop failures and subsequently
severe genetic erosion has taken place in the landraces that have been
maintained through many generations by the farmers. During the last
few years several regions in Ethiopia have been affected by severe
famine and farmers have been forced to consume the seeds normally
kept for planting the next season. This erosion is particularly aggravated by the distribution of food grain (mainly cereals) by relief
agencies because such grain can replace the native landraces.
Introduction of high-yielding varieties
It has been reported that the percentage of Ethiopian farmers
using improved seeds has reached 2 per cent, although in Arsi
administrative region (Anonymous, 1984), where research work has
been going on for many years, the figure is 21 per cent. Worede (1983)
stated that the traditional varieties have been almost completely
replaced by modern, uniform, advanced cultivars in areas such as
Chilalo in Arsi and Adaa in the central highlands. According to
unpublished data from the Ethiopian Seed Corporation (ESC), well
over 24 wheat and 9 barley varieties have entered commercial production in Ethiopia since 1968. A recent publication by ESC indicates that
7 wheat, 7 barley, 4 teff, 11 sorghum and 2 faba bean varieties have
been released for commercial production (Anonymous, 1981). This is
a welcome development approach although, paradoxically, it is a
threat to the genetic diversity on which future improvement work is
based. A number of recent varietal introductions of other cereals,
namely, sorghum, teff, and oilseeds such as Ethiopian mustard and
sesame, have been reported (Institute of Agricultural Research, Addis
Ababa, unpublished data).
Crop replacement
The traditional cereal crop of the Ethiopian highlands, barley,
is currently being replaced or complemented by oats (Avena sativa)
(Jutzi & Gryseels, 1984) and, at slightly lower altitudes, by wheat.
Maize is currently becoming a menace to sorghum in most traditional
sorghum growing areas. Moreover, while sorghum is being replaced
by bulrush millet (Pennisetum americanum), which is early maturing
204
Abebe Demissie
and drought-resistant (Michael, 1983), pearl millet is also becoming
popular at the expense of finger millet in northern Ethiopia.
Change in cropping patterns
Owing to the soaring prices of some agricultural crops in
recent years, the acreage of crops such as teff has increased at the
expense of other crops, such as wheat and millet. This subsequently
reduces the chance of maintaining the landraces of other traditional
crops, as local fanners favour the more profitable crops.
Change in land uses
Currently it is not difficult to see the deforestation process
that is taking place in the country. The semi-wild coffee is seriously
threatened by this process and also because it is being replaced by
crops, such as maize. Furthermore, large-scale agriculture is expanding, new roads are opening up remote areas, bushland is being taken
into cultivation and several other developments are taking place at
the expense of the wild and primitive crop germplasm.
Exploration and collecting activities
PGRC/E embarked on a systematic field collecting operation
in the 1977 crop season. Since then, many collecting expeditions have
been undertaken in all administrative regions of the country, covering a wide range of ecological zones.
Based on relevant and available information, collecting activities
were initiated according to well defined priorities for both crops and
areas. The priorities were based on criteria such as the economic and
social importance of the crops and their respective degree of genetic
erosion. Revisions of the priority list are made during regular workshops with the plant breeders on the basis of factual and up-to-date
data. Based on the criteria mentioned, the following list was formulated at the start of the collecting activities (Ebba, 1978):
1. Wheat
7. Vegetable crops
2. Barley
8. Forage crops
3. Teff
9. Coffee
4. Sorghum
10. Medicinal plants
5. Legume and oil crops
11. Forest trees
6. Root crops
12. Others
In the first 10 years of its operation, the Centre has conducted more
than 76 collecting expeditions. As a result, some 15000 accessions of
about 75 species or groups of species of mainly crop plants have been
collected (Table 1).
Table 1. Crop samples collected by PGRC/E from the various administrative regions of Ethiopia
Species
v
2
Brassica spp.
Capsicum spp.
Cicer arietinum
Coffea arabica
Eleusine coracana
Eragrostis tef
Guizotia abyssinica
Hordeum vulgare
Lathyrus sativus
Lens culinaris
Linum usitatissimum
Phaseolus spp.
Phytolacca dodecandra
Pisum sativum
Ricinus communis
Sesamum indicum
Sorghum bicolor
Trigonella foenum-graecum
Triticum spp.
Vicia faba
Other crops
42
6
16
0
0
16
12
462
2
21
51
1
0
51
9
1
17
9
225
30
21
34
5
11
0
0
20
8
142
0
13
29
7
0
44
13
0
10
9
121
21
23
1
0
1
0
5
8
25
78
0
0
10
1
0
5
4
1
52
1
31
5
1
13
10
6
0
12
10
7
152
0
6
7
32
0
23
9
9
179
2
51
16
110
72
15
106
0
161
204
172
278
60
25
48
32
0
56
27
13
47
59
429
70
100
Grand total
992
510
229
654
1974
3
4
5
7
8
90
17
110
0
107
223
123
584
20
1
116
4
1
148
10
1
119
55
204
152
130
36
4
19
0
4
11
1
215
0
16
19
61
0
20
49
5
70
10
107
29
84
25
6
0
0
8
19
7
5
0
1
3
6
0
0
3
1
176
0
1
2
26
22
4
0
39
3
44
5
37
22
0
6
3
0
14
1
4
50
1
5
15
17
2215
760
289
292
6
9
14
11
12
108
8
203
0
4
81
102
689
0
52
67
9
143
93
27
11
44
17
672
121
74
24
3
22
0
4
34
1
120
0
1
5
32
0
13
12
1
48
3
37
12
41
4
0
15
0
16
59
5
81
12
14
27
2
0
17
0
0
31
6
114
16
8
123
18
5
101
88
97
80
123
5
3
19
50
0
55
14
27
45
8
57
44
110
74
12
30
0
3
197
78
252
20
71
109
6
0
129
17
96
67
36
344
179
41
0
4
184
0
8
44
12
70
13
9
22
10
0
34
15
48
189
8
42
13
46
668
112
728
140
423
1067
638
3288
154
233
538
256
144
702
210
218
1144
224
2440
725
832
2525
413
427
1072
1761
771
14884
10
13
15
Total
a
Administrative regions: 1, Arsi; 2, Bale; 3, Eritrea; 4, Gamo Gofa; 5, Gojam; 6, Gondar; 7, Harerge; 8, Ilubabor; 9, Kefa; 10, Shewa; 11, Sidamo; 12, Tigray; 13,
Welega; 14, Welo; 15, Unknown.
206
Abebe Demissie
Collection strategy and sampling technique
Collection strategy
The collection strategy was based largely on broad-based or
non-crop specific missions rather than on crop specific or so-called
pointed collecting missions. During a non-crop specific mission the
crops that mature at more or less the same time in a given region are
collected in the farmer's field or in the markets. This strategy was
favoured particularly in drought-stricken areas where the ultimate
objective was to rescue whatever germplasm was still available.
However, in order to make better use of the foreign crop-specialist
collectors or to concentrate on the species of interest to any collaborating collectors (e.g. national plant breeders or scientists), pointed collection missions have been undertaken. In this way, a closer and
more detailed view of a given crop can be obtained and thus exploration missions, diversity studies and preliminary surveys of Ethiopian
oil crops (Demissie, 1984; Seegeler, 1986), as well as of other important crops such as chickpea, sorghum and finger millet, have been
carried out in collaboration with interested international agencies.
Furthermore, several missions have been launched to collect spices
and lesser known but potentially valuable plant species such as
Moringa stenopetala, Amorphophallus spp. and Sauromatum sp.
Sampling technique
The optimum sample size per collecting site is the number of
plants required to obtain, with 95 per cent certainty, all the alleles in a
population that occur in 5 per cent frequency or more (Marshall &
Brown, 1975). Hawkes (1976) suggested that bulked seed samples
from up to 50 individual plants, and certainly not more than 100,
should be collected non-selectively to obtain optimum sample size.
These plants are taken at random by walking backward and forward
through the field taking a sample every so many paces until the 50
plants have been sampled. Whenever rare types, i.e. plants which
show characters not included in the random sampling, are noticed a
selective sampling technique is adopted. The sample should be given
a different collection number (Hawkes, 1985).
The sampling of vegetatively propagated material such as sweet
potato, yams, taro, ensete, etc. requires distinct sampling techniques
since such crops do not occur as large populations, but are highly
selected individual genotypes. These crops are sampled on the basis
of information obtained from local farmers on the type and number of
varieties they grow. In general, it should be pointed out that
Germplasm exploration and collecting by PGRC/E
PGRC/E COLLECTION RECORD SHEET
COLLECTION NO.
DATE
COUNTRY
ADM. REGION
AWRAJA
WEREDA
VILLAGE/SITE
SPECIES
1
2
3
4
LONG.
(m)
1 Swamp
2 Flood plain
3 Plain level
4 Undulating
5 Hilly
6 Hilly dissected
7 Steeply dissected
8 Mountainous
9 Other (specify)
SITE:
1 Level
2 Slope
3 Summit
4 Depression
SOIL TEXTURE:
1 Sand
2 Sandy loam
3 Loam
4 Clay loam
5 Clay
6 Silt
SOIL COLOUR:
ACC. NO.
CROP
GENUS
LOCAL/VARIETY NAME
ETHNIC GROUP
LANGUAGE
SAMPLE TYPE:
MAP NO.
UT.
ALTITUDE
TOPOGRAPHY:
7 Highly organic
1 Black
2 Brown
3 Red
4 Orange
5 Yellow
6 Other (specify)
207
Single plant
Pure line/clone
Population/mixture
Other (specify)
GENETIC STATUS:
1 Wild
2 Weed
3 Primitive c u l t l v a r /
landrace
4 Breeders l i n e
5 Advanced cultivar
SOURCE OF COLLECTION:
1 Field
2 Backyard
3 Farm store/
threshing place
4 Market
5 Agricultural i n s t i t u t e
6 Natural vegetation
7 Other (specify)
MATERIAL:
1 Seed
2 Spike
3 Pods
4 Other (specify)
No
HERBARIUM SPECIMEN:: Yes
Yes
No
PHOTOGRAPH:
SOWING MONTH: 1 2 3 ^ 5 6 7 8 9 10 11 12
Early/Mid/Late
HARVEST MONTH: 1 2 3 ^ 5 6 7 8 9 10 11 12
Early/Mid/Late
ORIGIN OF SEED:
Local/Elsewhere
USAGE ( s p e c i f y )
STONINESS:
DRAINAGE:
0
1
2
1
2
None
Low
Medium
Poor
Moderate
3 Well drained
4 Excessive
DISEASES AND PESTS
NOTES: ( A s s o c i a t e d wild-weedy crop
species, local f l o r a , disturbance f a c t o r s ,
morphological v a r i a t i o n , husbandry, e t c . )
SOIL PH:
Fig. 1. PGRC/E's collecting fonn.
208
Abebe Demissie
vegetatively propagated material is often encountered in isolated conditions and sampling is often determined by the availability of
material. During collecting, relevant data are collected on the sampling site (ecology, soil, vegetation, etc.) and on the germplasm itself.
The data format in use has been adapted from the International Board
for Plant Genetic Resources (IBPGR) in order to meet the specific local
conditions (Fig. 1).
International cooperation in collecting activities
In 1981 and 1984, PGRC/E and the International Crops
Research Institute for the Semi-Arid Tropics (ICRISAT) organized
joint collecting expeditions which resulted in the acquisition of a wide
range of material. The first expedition focused on collecting Zera-zera
sorghum types which were identified as good in terms of their
agronomic desirability and their tolerance to diseases and drought
(Prasada Rao & Mengesha, 1981). The second mission was mainly
geared to collecting chickpeas (Pundir & Mengesha, 1982).
IBPGR has provided partial funding to a number of collecting
expeditions that have been carried out in the last 10 years. In July
1980, IBPGR assisted PGRC/E to collect 'belg' or small rainy season
crops (Toll, 1980). A similar type of assistance was offered by IBPGR
to collect the 'meher' or main rainy season crops which are associated
with the June-August rains of the Ethiopian highlands (Toll, 1982).
Furthermore, IBPGR co-sponsored a Brassica spp. collecting expedition in Ethiopia (Astley, Mahteme & Toll, 1982) as well as the visit of a
barley expert.
The German Agency for Technical Cooperation (GTZ) financed a
consultant-collector to assess the degree of genetic erosion in Ethiopian oil crops and to advise on priorities of collection (Seegeler, 1986).
Germplasm collected
In view of the importance of cereals as a staple food and the
degree of genetic erosion, PGRC/E gave top priority to the collection
of cereal germplasm. Well over 56 per cent of the material collected
consisted of the major cereals, i.e. wheat, sorghum, teff and barley.
The results of these cereal collection missions are discussed below,
together with facts and figures on the collection of some other traditional Ethiopian crops. A summary, by administrative region, of all
crops collected by PGRC/E is presented in Table 1.
In order to show, in a general way, the intensity of collection for
some of the major crops, Table 2 gives the calculated hectarage per
Table 2. Intensity of collecting of some of the major crops by administrative regions of Ethiopia, expressed in hectares
of production per collected accession (July 1986)
Cereals
Pulses
Oil crops
Administrative region
Teff
Sorghum
Barley
Wheat
Faba bean
Field pea
Chickpea
Noog
Linseed
Arsi
Bale
Eritrea
Gamo Gofa
Gojam
Gondar
Harerge
Ilubabor
Kefa
Shewa
Sidamo
Tigray
Welega
Welo
Overall mean for PGRC/E
collections
Overall mean for total
Ethiopian collections
Total hectarage of major
crops in Ethiopia ('000 ha)
1631
380
3275
1350
1198
950
364
2574
2216
3927
482
1107
1760
279
1270
80
1325
31
392
515
2564
80
578
2996
151
1429
1509
1182
309
461
369
163
300
167
9
740
514
246
343
349
152
24
716
296
526
35
64
160
73
3500
1920
284
268
353
84
67
1073
119
520
87
464
309
7
950
887
891
242
600
175
131
296
250
860
113
234
128
20
—
607
295
69
153
60
27
83
9
5800
250
73
505
11
—
—
266
186
1087
280
87
42
625
336
14
265
385
400
143
300
83
100
920
339
3
122
172
840
14
321
23
21
67
17
148
20
119
68
8
1224
638
222
232
393
166
208
236
100
575
111
99
89
238
104
208
163
62
728.3
565.0
297.8
116.8
157.6
136.3
53.3
1305.6
766.1
210
Abebe Demissie
collected sample for each of the administrative regions. Since the total
PGRC/E holdings per crop are generally much higher, due to donations and selections, the hectarage per accession of some crops in the
genebank has also been calculated. In both cases, the actual number
of hectares under a given crop per administrative region, as far as is
known, is used to calculate the intensity.
The figures in Table 2 do not allow any conclusions to be drawn on
the genetic value of the collected (or non-collected) samples. Aspects
such as the genetic diversity within and between accessions, and the
coverage of any given area, are important as well. If the collection
sites are plotted on a map of Ethiopia, it can easily be seen that by far
the majority of accessions have been collected along accessible roads
and only rarely have collections been made far from the roadsides.
Wheat (Triticum spp.)
In Ethiopia, it is common for a farmer to grow durum and
poulard wheat in the same fields. More recently, bread and durum
wheat have frequently been found together in the same field and
even a combination of these with T. polonicum or T. compactum can
occasionally be found.
Wheat is one of the most important cereals in Ethiopia, both in
terms of production and of acreage. According to the latest statistics,
a total of ca. 565 000 hectares are under wheat production
(Anonymous, 1984). From the genetic resources point of view also,
wheat is an important crop since several species have a secondary
centre of diversity in Ethiopia and the majority of the fields are still
planted with landrace populations. Wheat collections on a regional
basis ranged from one sample in Ilubabor to 675 in Shewa. The number of hectares per sample ranged from one sample for every 35
hectares in Gamo Gofa to one for every 3500 hectares in Ilubabor.
Overall, when considering the total holdings of PGRC/E, including
donations and selections, this figure comes to 89 hectares per accession, which is close to the figure reported by Chapman (1985) who
probably considered other Ethiopian material not yet in PGRC/E's
possession. This figure shows that wheat is one of the best collected
crops in Ethiopia in terms of hectarage per accession. The other
important aspect is the genetic diversity between accessions, but this
has not yet been assessed in a systematic way.
The altitudinal range for wheat in the areas covered varies from
1200 to 3300 m above sea level with the majority of the accessions
collected from altitudes above 2500 m (Table 3). A general estimate of
Germplasm exploration and collecting by PGRC/E
211
diversity of various crops was provided in a study by Mengesha
(1975). A detailed analysis of the regional pattern of phenotypic diversity in a limited number of Ethiopian durum and bread wheat accessions was carried out by Bekele (1984a). He reported that the total
variation differed from character to character and that the total phenotypic variation was highest within populations followed by differences
among populations within a region, and the least between regions.
Sorghum (Sorghum bicolor)
Sorghum is a major cereal in Ethiopia and is the third most
important in terms of production (Anonymous, 1984). Apart from
being a staple food crop, sorghum has several other uses, e.g. the
stalks are used for fuel and for house construction.
The first major effort to collect sorghum germplasm was launched
in 1967 after the formation of the Ethiopian Sorghum Improvement
Project. Since then, numerous collecting expeditions have been
undertaken by PGRC/E, partly in collaboration with ICRISAT
(Prasada Rao & Mengesha, 1981). Emphasis was mainly on areas that
had been less extensively collected.
The number of hectares per sample is ca. 638, when considering
the PGRC/E collection alone. The overall figure, considering all Ethiopian accessions held by PGRC/E, shows a much more favourable
figure, namely 111 hectares per accession (Table 2).
The altitudinal range for sorghum in the areas covered is from 400
to 2940 m (Table 3), which shows the wide ecological amplitude of
sorghum in Ethiopia.
Teff (Eragrostis tef)
Teff is the most important cereal crop and stands first in
terms of acreage, with 1 305 600 hectares of land under cultivation
(Anonymous, 1984). Its cultivation as a cereal crop is confined
almost entirely to Ethiopia and it is mainly used to make 'injera', a
local bread and an important part of the national dish.
Teff is known to have been domesticated in Ethiopia and a wealth
of diversity exists. Efforts have been made to collect landraces
systematically from each of the ecological zones. The germplasm collections made by PGRC/E in the last 10 years exceed 1100 accessions.
In this case also, the majority of the accessions are landraces.
The altitudinal range of the collected teff varies from 1120 to
2950 m. The intensity of collections from the different administrative
regions and the overall mean are shown in Table 2.
Table 3. Altitudinal range and frequency of occurrence of the various crops
Niger
Altitude
(m)
%
^1300
1301-1500
1501-1700
1701-1900
1901-2100
2101-2300
2301-2500
2501-2700
^2701
1.11
3.54
9.50
19.87
21.42
16.12
17.22
10.60
0.67
Altitude range (m)
1100-2950
Brassica
spp.
Altitude
(m)
%
^1300
1301-1500
1501-1700
1701-1900
1901-2100
2101-2300
2301-2500
2501-2700
2701-2900
5*2901
1.07
1.70
7.86
17.20
16.78
15.50
19.96
14.87
3.87
1.28
1050-3170
Chickpea
Altitude
(m)
%
^1300
1301-1500
1501-1700
1701-1900
1901-2100
2101-2300
2301-2500
2501-2700
^2701
0.28
3.54
7.88
22.01
24.19
11.69
16.85
12.50
1.09
1200-2880
Field pea
Altitude
(m)
%
s=1600
1601-1800
1801-2000
2001-2200
2201-2400
2401-2600
2601-2800
2801-3000
3001-3200
^3201
0.30
0.60
3.29
6.87
17.62
25.97
22.09
15.53
5.68
2.09
1560-3380
Faba bean
Altitude
(m)
%
^1800
1801-2000
2001-2200
2201-2400
2401-2600
2601-2800
2801-3000
^3001
0.79
5.24
9.43
15.71
27.23
24.87
13.62
3.15
1300-3150
Table 3 (cont.)
Wheat
Altitude
(m)
%
=£1500
1501-1700
1701-1900
1901-2100
2101-2300
2301-2500
2501-2700
2701-2900
2901-3100
523101
0.19
0.49
3.27
5.70
11.82
25.50
30.41
16.60
5.21
0.85
Altitude range (m)
120O-3300
Teff
Barley
Altitude
(m)
%
=£1700
1701-1900
1901-2100
2101-2300
2301-2500
2501-2700
2701-2900
2901-3100
3101-3300
3301-3500
^3501
0.63
2.60
6.91
8.67
15.58
21.34
20.41
15.27
6.39
1.93
0.32
1500-3750
Altitude
(m)
%
s=1200
1201-1400
1401-1600
1601-1800
1801-2000
2001-2200
2201-2400
2401-2600
2601-2800
^2801
0.33
1.48
7.38
12.63
20.50
13.28
15.74
20.66
6.73
1.32
1120-2950
Sorghum
Altitude
(m)
%
=£950
951-1150
1151-1350
1351-1550
1551-1750
1751-1950
1951-2150
2151-2350
2351-2550
2551-2750
5*2751
400-2940
5.07
1.77
8.81
20.27
21.81
17.85
9.26
8.59
3.75
1.55
1.33
Linseed
Altitude
(m)
%
=£1700
1701-1900
1901-2100
2101-2300
2301-2500
2501-2700
2701-2900
2901-3100
3101-3300
5^3301
1.06
2.46
9.13
12.28
20.00
26.32
14.39
8.78
3.86
1.76
1470-3430
214
Abebe Demissie
Barley (Hordeum vulgare)
Barley was given high priority since it is an important crop in
Ethiopia, the third in terms of acreage. It possesses a high genetic
diversity (Ethiopia is a secondary gene centre for cultivated barley)
and considerable genetic erosion is being observed. The number of
accessions collected, as well as the size of the total barley collection
maintained at PGRC/E, reflect this priority (Tables 1 and 2).
The potentialities of the barley collections have been demonstrated
by Qualset (1975), who listed an impressive number of accessions
which possess resistance genes against one or more important
diseases, as well as some important quality characters, e.g. high protein and lysine content. Bekele (1983a,b, 1984b) investigated the
diversity existing in Ethiopian barley for the allozyme genotypic composition, the genetic distances between populations based on morphological characters and flavonoids. Significant differences between
regions were found and conclusions for future collection and conservation were drawn. Engels (1987) compared the diversity indices
for the administrative regions and for the country as a whole, for a
large collection, with the results of earlier diversity studies and concluded that the diversity indices for the different regions are generally
not significant and that the overall index for Ethiopia is high.
Pulses
A number of important food crops belong to this section.
Some species (e.g. Vicia faba, Pisum sativum and Cicer arietinum) have
built up a significant diversity in Ethiopia and form an important
source for the local and international breeding programmes. Therefore, considerable efforts have been put into the collection of these
crops and a total of almost 3000 accessions have been collected
throughout Ethiopia (Table 1). The collection intensity of some of the
major pulses is comparable with the other major crop species (Table
2). The altitudinal range for some of the pulse crops can be observed
in Table 3.
Root and tuber crops
At present, only a modest collection of root and tuber crops is
maintained by PGRC/E. A total of some 100 accessions of various
species has been collected so far, including a sweet potato collection
of 42 accessions, as well as yams (Dioscorea spp.), Coccinia abyssinica
and Colocasia esculenta). This collection includes species which are of
regional importance in times of drought and food shortage (e.g.
Table 4. Some of the minor native species with potential value collected by PGRC/E
Scientific name
Common name
Uses
Distribution
Abelmoschus spp.
Arisaema sp.
Amorphophallus abyssinicus
Oryza longistaminata
Brassica oleracea
Coccinia abyssinica
Plectranthus edulis
Ensete ventricosum
Okra (incl. wild species)
Burie
Bagana
Wild rice
Gurage gomen
Anchote
Oromo dinich
Ensete
Vegetable
Root crop
Root crop
Food grain
Leaf vegetable
Tuber crop
Tuber crop
Edible pseudostem
Ipomoea batatas
Sweet potato
Root crop
Moringa stenopetala
Sauromatum nubicum
Trigonella foenum-graecum
Cabbage tree, Shiferaw, Haleko
Banshalla
Fenugreek
Leaf vegetable
Root crop
Baby food
Ilubabor, Welega
Gamo Gofa, Sidamo
Gamo Gofa
Gojam, Ilubabor
Shewa, Sidamo
Welega, Shewa
Shewa, Welega
Shewa, Kefa, Gamo Gofa,
Sidamo
Harerge, Gamo Gofa, Sidamo,
Shewa
Gamo Gofa
Gamo Gofa, Sidamo
Spread over the country
216
Abebe Demissie
Amorphophallus abyssinicus and Sauromatum nubicum). The sweet
potato collection is being systematically screened for reaction to some
prevalent diseases and pests and increasing use is being made of
traditional varieties, especially for their adaptability to local stress
conditions.
Oil crops
Oil crops are another important group of food plants which
are grown extensively in the country. Some species are native to
Ethiopia and were first taken into domestication in the Ethiopian
highlands, e.g. niger seed or noog (Guizotia abyssinica) and Ethiopian
mustard or gomen (Brassica carinata). Other oil crops such as linseed
(Linum usitatissimum), sesame (Sesamum indicum) and safflower (Carthamus tinctorius), have been given due priority (Table 1).
The intensity of collecting for noog ranges from one sample for
every 3 hectares in Welo to one sample per 625 hectares in Bale. The
overall acreage per collected accession for the whole of Ethiopia is 163
hectares and this is comparable to the intensity for other crops. The
considerable altitudinal range, from 1100 to 2950 m, is worth mentioning and shows the broad adaptability of the crops in question.
Miscellaneous and under-utilized crop plants
During the last 10 years, PGRC/E has also explored for and
collected some lesser known but potentially valuable indigenous crop
plants. These include some regionally important crops which are
utilized by local people in times of food shortages (Table 4).
References
Anonymous (1981). Some technical information on seeds. Ethiopian Seed
Corporation, Addis Ababa (mimeographed).
Anonymous (1984). General agricultural survey. Preliminary report 1983-4,
vol. I. Planning and Programming Department, Ministry of Agriculture,
Addis Ababa (mimeographed).
Astley, D., Mahteme, H. G. & Toll, J. (1982). Collecting brassicas in Ethiopia.
Plant Genetic Resources Newsletter, 51, 15-20.
Bekele, E. (1983a). Allozyme genotypic composition and genetic distance
between the Ethiopian landrace populations of barley. Hereditas, 98,
259-67.
Bekele. E. (1983b). A differential rate of regional distribution of barley
flavonoid patterns in Ethiopia and a view on the centre of origin of barley.
Hereditas, 98, 269-80.
Bekele, E. (1984a). Analysis of regional pattern of phenotypic diversity in the
Ethiopian tetraploid and hexaploid wheats. Hereditas, 100, 131-54.
Bekele, E. (1984b). Relationships between morphological variance, gene
Germplasm exploration and collecting by PGRC/E
217
diversity and flavonoid patterns in the landrace populations of Ethiopian
barley. Hereditas, 100, 271-94.
Chapman, C. D. G. (1985). The Genetic Resources of Wheat: a survey and strategies
for collecting. IBPGR, Rome.
Demissie, A. (1984). Oilcrops exploration and collection in Ethiopia. PGRC/E,
Addis Ababa (mimeographed).
Ebba, T. (1978). Plant Genetic Resources Centre/Ethiopia activities and programs. Crop genetic resources in Africa. Proceedings of a workshop jointly
organized by AAASA and IITA. International Institute of Tropical Agriculture, Ibadan, pp. 25-30.
Engels, J. M. M. (1987). A diversity study in Ethiopian barley. In: J. M. M.
Engels (ed.), The conservation and utilization of Ethiopian germplasm.
Proceedings of an international symposium, Addis Ababa, Ethiopia, 13-16
October 1986, pp. 124-32 (mimeographed).
Gentry, H. S. (1971). Pea picking in Ethiopia. Plant Genetic Resources Newsletter, 26, 20-4.
Hawkes, J. G. (1976). Sampling gene pools. Proceedings of Nato conference on
conservation of threatened plants. Series 1, Ecology. Plenum, London.
Hawkes, J. G. (1985). Report on a consultancy mission to Ethiopia for GTZ to
advise PGRC/E on germplasm exploration, conservation, multiplication
and evaluation. Birmingham (mimeographed).
Jutzi, S. & Gryseels, G. (1984). Oats, a new crop in the Ethiopian highlands.
PGRC/E-ILCA Germplasm Newsletter, 5, 22-4.
Marshall, D. R. & Brown, A. H. D. (1975). Optimum sampling strategies in
genetic conservation. In: O. H. Frankel and J. G. Hawkes (eds), Crop
Genetic Resources for Today and Tomorrow. Cambridge University Press,
Cambridge, pp. 53-80.
Mengesha, M. H. (1975). Crop germplasm diversity and resources in Ethiopia. In: O. H. Frankel and J. G. Hawkes (eds), Crop Genetic Resources for
Today and Tomorrow. Cambridge University Press, Cambridge, pp. 449-53.
Michael, T. (1983). Germplasm conservation in Eritrea. PGRC/E-ILCA Germplasm Newsletter, 4, 7-9.
Prasada Rao, K. E. & Mengesha, M. H. (1981). A pointed collection of 'ZeraZera' sorghum in the Gambella area of Ethiopia. Genetic resources progress
report 33, ICRISAT, Patancheru.
Pundir, R. P. S. & Mengesha, M. H. (1982). Collection of chickpea germplasm
in Ethiopia. Genetic resources progress report 44, ICRISAT, Patancheru.
Qualset, C. D. (1975). Sampling germplasm in a centre of diversity: an example of disease resistance in Ethiopian barley. In: O. H. Frankel and J. G.
Hawkes (eds), Crop Genetic Resources for Today and Tomorrow. Cambridge
University Press, Cambridge, pp. 81-94.
Seegeler, C. J. P. (1986). Genetic variability of oil crops in Ethiopia. Consultancy report for GTZ by PGRC/E. Oosterbeek (mimeographed).
Toll, J. (1980). Collecting in Ethiopia. Plant Genetic Resources Newsletter, 43,
36-9.
Toll, J. (1982). Collecting in Ethiopia. Plant Genetic Resources Newsletter, 48,
18-22.
Vavilov, N. I. (1951). The origin, variation, immunity and breeding of
cultivated plants. Chronica Botanica, 13, 1-366.
Worede, M. (1983). Crop genetic resources in Ethiopia. In: J. C. Holmes and
W. M. Tahir (eds), More Food from Better Technology. FAO, Rome, pp. 143-7.
Zohary, D. (1970). Centres of diversity and centres of origin. In: O. H.
Frankel and E. Bennett (eds), Genetic Resources in Plants, their Exploration
and Conservation. Blackwell, Oxford, pp. 33-42.
15
Collection of Ethiopian forage
germplasm at the International
Livestock Centre for Africa
JEAN HANSON AND SOLOMON MENGISTU
Introduction
Ethiopia is an area rich in germplasm of many plant species
and was considered as a primary centre of crop diversity by Vavilov
(1951). Among the most important plant genetic resources of the East
African region, and indeed of all of Africa, are forages and especially
forage grasses (Zeven & Zhukovsky, 1975). Large areas of Africa are
covered with tropical savannah with a great diversity of grasses
which are vigorous and polymorphic (Clayton, 1983). Tropical forage
legumes and browse species are also endemic. In particular, Africa
has been described as the centre of diversity of the browse shrubs of
the subfamily Caesalpinioideae (Williams, 1983).
The genetic resources of forages are usually found in wild populations since they have only been cultivated on a commercial scale for
about 50 years and no landraces are available (Williams, 1983). This is
very different from crop species and therefore the collection strategies
for forages differ to accommodate the population structures found in
the wild. Marshall & Brown (1983) have defined the objective of
forage plant exploration as the collection of material with the maximum amount of useful genetic variability within a strictly limited
number of samples. The strategy of the International Livestock
Centre for Africa (ILCA) is to collect representative population samples from the wild, although in some cases only a few plants can be
found growing together as a population and sampling is therefore
limited (Lazier, 1984). Collection is always a compromise between
capturing the greatest amount of variation using theoretical collection
Collection of Ethiopian forage germplasm
219
procedures and practical constraints imposed by field conditions.
Reid & Strickland (1983) support the concept of Harlan (1975) which
considers that, given the restrictions and practical constraints found
in the field, it is more important to sample the maximum number of
sites than to collect the theoretically ideal number of plants per site,
because more variation is likely to be captured by sampling in different ecological zones. Whenever possible, large seed samples are
preferred because these are more representative of the total variation
in the wild and more likely to capture genes which occur with low
frequency in the population (Marshall & Brown, 1983). Also the seed
sample can be placed into the genebank without a regeneration cycle
prior to storage.
Potential of Ethiopian forage species
Large numbers of grass and legume species found in Ethiopia
have the potential to be developed as forages. Many species have
been collected whose true value is still unknown and these must be
evaluated before they can be utilized. When collecting forages for
genetic resources, all plants which could prove useful as forage crops
should be collected in the field. Plants which may not seem directly
useful may have unseen characteristics such as disease, insect or
drought tolerance. However, in addition, there are certain target species which have proven forage potential and are therefore given priority during collecting missions.
A large number of grass genera which are endemic to Africa are of
special interest during collecting missions. Bogdan (1977) has identified 45 grasses which he considers to be the most important for
forage. Of these, 27 are endemic to Africa, indicating the importance
of collecting in this region (Clayton, 1983). Zeven & Zhukovsky (1975)
have considered that Brachiaria brizantha, B. decumbens, B. mutica,
Chloris gayana, Melinis minutiflora, Pennisetum dandestinum and species
of Cynodon, Digitaria and Setaria are endemic grasses whose centre of
origin lies within the Ethiopian centre of diversity.
Several important genera of forage legumes are native to Ethiopia
and have been identified as targets for collecting. Zeven &
Zhukovsky (1975) consider Acacia, Crotalaria, Indigofera, Lablab
purpureus and Stylosanthes fruticosa to have their centres of diversity in
Africa. The variation found within these genera and species in Ethiopia supports this. Other important legumes showing considerable
variation belong to the genera Aeschynomene, Alysicarpus, Medicago,
Neonotonia (i.e. N. wightii) and Trifolium. Browse species, including
220
Jean Hanson & Solomon Mengistu
Acacia, Cordeauxia, Cassia, Sesbania and Erythrina, are also native to
East Africa and show considerable potential for development as forages, especially in drier areas where their deep roots provide some
tolerance to periods of low rainfall.
Forage legumes are especially important in the native agricultural
system because they both provide fodder and increase soil fertility by
nitrogen fixation in association with Rhizobia. In Ethiopia, the native
genus Trifolium is of interest due to its abundance and observed
potential in highland areas. The Trifolium species collected by ILCA
are being evaluated to identify suitable genotypes for use in the
diverse environment and soil types found in the Ethiopian highlands
(Kahurananga, 1982; Kahurananga & Tsehay, 1984).
ILCA collecting missions
The forage agronomy programme of ILCA has made 22 collecting missions in Ethiopia in the last six years. The target genera for
these missions have been for the most part those which have shown
potential in work under way at ILCA or in other forage research
organizations. Initially, forage collecting was concentrated largely in
accessible parts of the Ethiopian highlands because the potential of
native African Trifolium species had already been recognized during
some preliminary screening in Kenya (Strange, 1958; Bogdan, 1977)
and in Ethiopia (Chilalo Agricultural Development Unit, 1972).
Although these earlier ILCA missions were primarily for Trifolium,
other highland legumes of potential were also collected. Later, the
collecting missions were extended to the lowlands for more general
collecting and also for browse species, an area of interest and research
at ILCA. These missions covered a large area of the central and
southern parts of the country. The major genera collected are listed in
Table 1.
Most of these ILCA collecting missions were funded by the International Board for Plant Genetic Resources (IBPGR) who recognized
the need to collect forages in Ethiopia. ILCA collections in 1985 were
also carried out in cooperation with staff of the Centro International
de Agricultura Tropical (CIAT) with a special emphasis being given to
the grass genus Brachiaria which has shown potential in Latin
America.
Highland collections
There are about 64 endemic legume species in Ethiopia and
most of these can be found in the highlands. The most abundant
221
Collection of Ethiopian forage germplasm
Table 1. Number of collections of major forage genera from the
administrative regions of Ethiopia
Administrative region0
Genus
Alysicarpus
Argyrolobium
Cassia
Crotalaria
Desmodium
Eriosema
Heteropogon
Indigofera
Lablab
Lotus
Lupinus
Medicago
Neonotonia
Rhynchosia
Stylosanthes
Tephrosia
Teramnus
Trifolium
Vicia
Vigna
Zornia
Ar
Ha
n
Ke
6
4
8
3
3
1
3
2
1
1
Sh
Si
4
1
3
1
2
2
We
Wl
Total
8
1
31
1
11
6
8
3
1
3
5
1
14
11
29
13
100
56
20
1063
33
26
19
1
2
2
1
1
2
2
1
1
1
1
12
1
2
1
108
4
2
Acacia
Aeschynomene
Albizia
Dichrostachys
Entada
Erythrina
Pseudarthria
Sesbania
1
2
3
1
3
1
a
Gn
Gj
1
Andropogon
Brachiaria
Cenchrus
Chloris
Digitaria
Echinochloa
Festuca
Panicum
Pennisetum
Setaria
Total
Ga
Ba
4
2
1
3
1
105
2
1
14
1
2
1
217
2
2
3
5
1
6
33
3
2
3
6
2
1
2
14
1
1
1
2
40
1
4
1
6
1
1
1
30
6
409
11
7
2
2
1
1
1
1
2
1
1
1
3
1
2
1
1
28
2
4
1
4
122
32
259
1
38
4
3
17
41
5
1
41
10
8
4
1
2
4
1
4
25
1
2
13
1
1
1
1
1
2
1
5
7
44
2
7
1
3
1
3
2
38
6
38
1
1
1
66
1
6
6
2
4
532
307
92
2
92
12
11
5
2
2
5
9
13
46
6
1
3
1
13
5
15
1
1
1
123
2
2
10
6
95
15
82
1697
Ar, Arsi; Ba, Bale; Ga, Gamo Gofa; Gj, Gojam; Gn, Gondar; Ha, Harerge; II, Ilubabor; Ke, Kefa; Sh,
Shewa;, Si, Sidamo; We, Welega; Wl, Welo.
222
Jean Hanson & Solomon Mengistu
Table 2. Number of accessions of Trifolium collected in Ethiopia from
1982 to 1986 showing the range of altitudes
Altitude range (m)
Species
Trifolium acaule
T. baccarinii
T. bilineatum
T. burchellianum
T. calocephalum
T. crytopodium
T. decorum
T. mattirolianum
T. multinerve
T. pichisermollii
T. polystachyum
T. quartinianum
T. rueppellianum
T. schimperi
T. semipilosum
T. simense
T. steudneri
T. tembense
Total
a
<1500
1500-2000 2001-2500 2501-3000 <3000
2
11
17
2
1
2
1
5
33
1
9
14
18
5
5
20
5
1
21
25
11
3
10
43
12
1
8
24
15
53
18
56
27
43
57
3
16
3
24
14
43
17
1
12
12
10
34
4
71
37
10
75
16
6
13
1
10
2
12
Total*
4
50
45
53
23
66
67
48
13
21
43
30
105
27
142
66
73
149
1025
Thirty-eight of the 1063 accessions of Trifolium do not have altitude data.
genus found in the cool highlands is Trifolium, and 28 of the 40
species of African Trifolium are found in Ethiopia of which nine are
endemic (Gillett, Polhill & Verdcourt, 1971; Thulin, 1983).
Collecting began in the highlands in 1980 when the first mission
collected Trifolium species from Debre Zeit for immediate evaluation
(Kahurananga, 1982). Collecting was intensified in 1982-4 when 17
missions covered all accessible areas of the central highlands
(Kahurananga & Mengistu, 1983, 1984). A large number of accessions
were collected during these missions, the majority belonging to 18
species of the genus Trifolium from a wide range of altitudes (Table 2).
Accessions from other highland genera including Vicia, Medicago and
Lotus were also collected.
Lowland collections
Two general collecting missions were made in 1984 and 1985
for lowland grasses and legumes. The mid-highland and lowland
savannah and bushland of the Harerge, Bale, Sidamo, Gamo Gofa,
Collection of Ethiopian forage germplasm
223
Welega and Bale administrative regions were a good source of forage
grasses including Brachiaria, Cenchrus, Chloris, Andropogon, Panicum
and Cynodon. Forage legumes were also abundant, including Vigna,
Desmodium, Alysicarpus, Macrotyloma, Zornia, Lablab, Stylosanthes and
Neonotonia. Neonotonia is very widespread and was found at a wide
range of altitudes, growing in diverse vegetation types. Both
Neonotonia and Stylosanthes fruticosa are vigorous and abundant in the
Rift Valley and are showing considerable promise in the preliminary
forage evaluation plots in that area.
Three collecting missions were undertaken in 1984 to collect
browse tree and shrub germplasm and to identify browse species
which could be used as fodder. These missions covered the lowlands
of Harerge, Sidamo, Gamo Gofa, Kefa and Ilubabor. The collection
routes covered more than 7000 km across varied vegetation types,
differing terrain and a wide range of altitudes and reached as far as
the Kenyan frontier in the south and the Sudan frontier in the southwest. In all, 103 accessions were collected, most of which belonged to
the genera Acacia, Aeschynomene, Albizia, Eryihrina, Sesbania and
Tamarindus (Mengistu, 1985). Data on palatability, growth season and
traditional feeding method of native forage species were collected
from the rural people. This information, together with visual observations in the field, has been extremely useful to determine which
genera and species are of particular interest for collecting and which
are likely to show potential for development as forages.
Priorities for future collecting
Additional collecting of forages in Ethiopia is necessary to
broaden further the genetic base of the forage crops available and
identify promising genera and species for further study. Forage collecting at ILCA must fill gaps in current collections, salvage
endangered germplasm, fulfil the needs of ILCA, national and international forage programmes and, to some extent, concentrate on the
species of known forage potential.
Collection objectives are closely linked to the needs and goals of
the ILCA forage programme and the need to complement the available germplasm with species of potential forage value. Native Ethiopian legumes of value include, among others, Macrotyloma axillare,
Neonotonia wightii and Stylosanthes fruticosa. Ecotypes with high dry
matter yield, high nitrogren fixing capacity, vigorous growth in
minimal rainfall areas and wide adaptation beyond their normal
environmental range will be sought. Greater emphasis will be placed
224
]ean Hanson & Solomon Mengistu
on the collecting of associated Rhizobia with the different legumes, so
that maximal productivity is achieved in the field. Other priority
genera for collecting include Acacia, Trifolium, Erythrina, Eriosema,
Galactia and Lotus.
Collecting will continue in specific environments where potentially
important genetic variation can exist and in areas not yet fully sampled. Certain genera and species have not yet been extensively collected by ILCA. This is especially true of the grasses, where only a
few collections have been made, but a large amount of variation
occurs in the field. Grasses, legumes and browse are all of interest
from environments which are swampy, seasonally inundated, arid or
semi-arid.
ILCA will continue to collect forages in areas where germplasm is
endangered and genetic erosion is occurring due to such factors as
drought, land clearing, intensive cultivation and heavy grazing. Collecting around the periphery of such areas can yield very promising
types because of the extreme selection pressures. Collecting can both
salvage germplasm and provide a gene pool of ecologically adapted
material for further development for use in similar areas.
References
Bogdan, A. V. (1977). Tropical Pasture and Fodder Plants. Longman, London.
Chilalo Agricultural Development Unit (1972). Report on Surveys and Experiments Carried Out in 1972. Publication No. 80, CADU, Ethiopia.
Clayton, W. D. (1983). Tropical grasses. In: J. G. Mclvor and R. A. Bray (eds),
Genetic Resources of Forage Plants. CSIRO, Australia, pp. 39-46.
Gillett, J. B., Polhill, R. M. & Verdcourt, R. (1971). Leguminoseae (part 3). In:
E. Milne-Redhead and R. M. Polhill (eds), Flora of Tropical East Africa.
Crown Agents, London, pp. 1016-36.
Harlan, J. (1975). Seed crops. In: O. H. Frankel and J. G. Hawkes (eds), Crop
Genetic Resources for Today and Tomorrow. Cambridge University Press,
Cambridge, pp. 111-15.
Kahurananga, J. (1982). ILCA's forage legume work in the Ethiopian Highlands. ILCA Newsletter, 1, 5-6.
Kahurananga, J. & Mengistu, S. (1983). ILCA native forage germplasm collection in Ethiopia for 1982-1983. PGRC/E-ILCA Germplasm Newsletter, 3,
6-9.
Kahurananga, J. & Mengistu, S. (1984). ILCA native forage germplasm collection in Ethiopia during 1983. PGRC/E-ILCA Germplasm Newsletter, 5,
8-18.
Kahurananga, J. & Tsehay, A. (1984). Preliminary assessment of some
annual Ethiopian Trifolium species for hay production. Tropical Grasslands,
18, 215-17.
Lazier, J.R. (1984). Theory and practice in forage germplasm collection.
Paper presented at PANESA Workshop on Pasture Improvement Research
Collection of Ethiopian forage germplasm
225
in Eastern and Southern Africa, Harare, Zimbabwe, 17-21 September 1984.
IDRC proceedings series 237-e.
Marshall, D. R. & Brown, A. H. D. (1983). Theory of forage plant collection.
In: J. G. Mclvor and R. A. Bray (eds), Genetic Resources of Forage Plants.
CSIRO, Australia, pp. 135-^8.
Mengistu, S. (1985). ILCA browse collection in Ethiopia 1984. PGRC/E-ILCA
Germplasm Newsletter, 8, 19-23.
Reid, R. & Strickland, R. W. (1983). Forage plant collection in practice. In:
J. G. Mclvor and R. A. Bray (eds), Genetic Resources of Forage Plants. CSIRO,
Australia, pp. 149-56.
Strange, R. (1958). Preliminary trials with grasses and legumes under grazing. East African Agricultural Journal, 24, 92-102.
Thulin, M. (1983). Leguminoseae of Ethiopia. Opera Botanica, 68.
Vavilov, N. I. (1951). The origin, variation, immunity and breeding of
cultivated crops. Chronica Botanica, 13, 1—366.
Williams, R. J. (1983). Tropical legumes. In: J. G. Mclvor and R. A. Bray (eds),
Genetic Resources of Forage Plants. CSIRO, Australia, pp. 17-37.
Zeven, A. C. & Zhukovsky, P. M. (1975). Dictionary of Cultivated Plants and
their Centres of Diversity. PUDOC, Wageningen.
16
Germplasm conservation at PGRC/E
REGASSA FEYISSA
Introduction
Genetic conservation has arisen as a solution to some of the
problems caused by Man in his social and agricultural relationship
with the environment (Simmonds, 1979). Unwise exploitation of
nature has caused an irreversible loss of variability and has become
the major cause of worldwide genetic erosion. The seriousness and
rapid expansion of the problem has created a universal need to collect
and conserve genotypes that would no longer be available if not
conserved today. This can best be achieved by maintaining a wide
range of plant materials covering the maximum variability existing at
present.
Taking into account these needs, and being aware of the enormous
diversity of crops in Ethiopia, the Plant Genetic Resources Centre
(PGRC/E), is currently working on the conservation of both orthodox
and recalcitrant crops. At present, the centre holds 40000 accessions
of 78 different species, including the germplasm material preserved in
field genebanks.
Facilities, personnel and organization of the Conservation
Division
The longevity of any conserved material depends upon the
system of conservation used and this, in turn, is affected by the
facilities existing at any given genebank and the quality of technical
knowledge available. The inadequacy of the infrastructures for the
maintenance and utilization of plant genetic resources remains the
major limiting factor in the establishment of a genebank in a developing country. Storage facilities require large inputs in terms of construction, equipment and maintenance costs, as well as capable
Germplasm conservation at PGRC/E
227
technicians and a reliable electricity supply. The size of accessions to
be stored, their safety, frequency of rejuvenation and flow of samples
in evaluation and exchange are dependent on the quality and size of
the storage facilities. Similarly, the equipment required by a
genebank varies according to the number of samples and species to
be stored.
The diversity of species maintained at PGRC/E requires an assortment of equipment. At present the Conservation Division is equipped with germination room, maintained at about +20°C with 80 per
cent relative humidity (RH), and a germinator with alternating
temperature. The drying room is fitted with a dehumidifier and an air
conditioner which allow operation at 18-20 °Cwith 15-18 per cent RH.
The seed processing and germination test laboratories are equipped
with seed blowers, several types of scales, a vacuum system with
counting heads used to place seeds in petri dishes, a metal can sealer,
and a sealer for aluminium foil bags. The three cold stores possess a
total volume of about 225 m 3 of which 175 m 3 is used for storage at
—10 °C.The remaining 50 m 3 store is maintained at +4°C and 35 per
cent RH for temporary storage of accessions with insufficient seeds for
long-term storage. The cold rooms are fitted with mobile shelves
which give convenient working conditions and economize on space.
With the help of a computerized numbering system each of the accessions can be located without problems. Electricity is supplied by two
independent lines and a standby generator automatically switches on
if the power supply fails or the voltage varies too much. Personnel in
the Conservation Division includes two physiologists, five trained
technical assistants, eight laboratory assistants and eight supporting
staff.
Efficiency and caution must be given high priority during the processing and storage of the seeds and during the compilation of information. This can be achieved by organizing the activities in an orderly
and logical manner. Figure 1 shows the organization and distribution
of activities between units in the Germplasm Conservation Division
at PGRC/E.
The process of seed deterioration begins immediately after the
seeds have matured on the plant itself. Thus, seeds collected from the
farmers' fields, stores and marketplaces, as well as from seed
increase, have to be processed and stored at the genebank as quickly
as possible to avoid further deterioration. A general scheme developed by PGRC/E, which helps to perform these activities efficiently,
is presented in Fig. 2.
228
Regassa Feyissa
UNITS
ACTIVITIES
Processing unit
Seed drying unit
- Fumigation
- Cleaning
-Total and TSW determination
- Data recording
- Moisture content determination
- Drying
- Sub-sampling
- Data recording
Seed testing unit
Storage unit
Evaluation unit
- Germination test
- Dormancy breaking
- Tetrazolium test
- Viability monitoring of stored seeds
- Receiving samples and storage
-Seed delivery
- Inventory work
- Monitoring sample size in the genebank
- Data recording
- Determination of constituents (protein, oil, fibre, fat)
- Screening germplasm for physiological stress conditions
- Health testing (nutrient deficiency)
Fig. 1. Organization of PGRC/E's conservation units and their
respective activities.
Sources of germplasm
Collection, selection, donation and repatriation are the main
sources of our germplasm material (Table 1). Details of the major crop
species are presented in Table 2 and in a paper on Documentation at
PGRC/E (Sendek & Engels, 1988, Chapter 17).
Germplasm conservation at PGRC/E
—I
• Flow of material
High=High seed viability
229
uisiriDUuon
Flow of information
Low=Low seed viability
TSW
Thousand seed weight
Fig. 2. Organizational chart of the seed conservation division at
PGRC/E.
230
Regassa Feyissa
Table 1. Source and number of accessions held at PGRC/E
Source
Number of accessions
Percentage of total
Collection
Selection
Donations
Repatriation
Total
15126
5840
15227
3785
39978
37.84
14.60
38.09
9.47
100.00
Table 2. Major crop types and corresponding number
of accessions kept by PGRC/E
Crop type
Number of accessions
Brassica spp.
Cicer arietinum
Coffea arabica
Eleusine coracana
Eragrostis tef
Guizotia abyssinica
Hordeum vulgare
Lens culinaris
Linum usitatissimum
Pisum sativum
Sesamum indicum
Sorghum bicolor
Trigonella foenum-graecum
Triticum spp.
Vicia faba
954
902
662
796
2270
924
9316
427
1820
1133
376
8145
438
8444
1198
2173
39978
Others
Total
Full details on the collection and exploration activities of PGRC/E
are presented elsewhere in this volume (Demissie, 1988, Chapter 16).
Selection as a source of germplasm refers mainly to the splitting of
populations or mixtures into agro-morphological components and
details are discussed by Mekbib (1988, Chapter 21).
During its early years, PGRC/E actively tried to incorporate as
many working collections of Ethiopian plant breeders as possible into
the genebank collection. Several of these included duplicate samples
of germplasm which had been collected by reputable collectors or
institutes, mainly during the 1960s.
Since the International Board for Plant Genetic Resources (IBPGR)
Germplasm conservation at PGRC/E
231
Table 3. Sample size required for long-term storage
Sample type
TSW 5-200 g
TSW >200 g
Heterogeneous
Total
B.C.
A.C.
8000 seeds
3000 seeds
5 x 1000 seeds
4000 seeds
1500 seeds
5 x 500 seeds
Homogeneous
Total
B.C.
A.C.
3200 seeds
800 seeds
6 x 400 seeds
1600 seeds
400 seeds
6 x 200 seeds
B.C., base collection; A.C, active collection; TSW, thousand seed weight.
assigned global or regional responsibilities to PGRC/E for a number of
crops, duplicate samples are regularly received for conservation
purposes. Major collections in several countries, e.g. USA, USSR,
Japan, Italy and the Netherlands, form an important source of Ethiopian germplasm. This germplasm was collected in the past and the
duplicate samples left behind in Ethiopia have sometimes been lost,
partly because of inadequate storage.
Preparing seeds for long-term storage
Registration and cleaning
The first step in seed preparation is the registration of samples. Identification data related to each sample must be recorded and
care taken in order to avoid duplications and/or errors. As a
phytosanitary measure, seeds coming into the genebank are fumigated for 72 hours with phosphine to control further infestation and
damage. Due to the heterogeneity of some of the samples, mechanical selection of seeds is likely to occur if seed cleaning equipment is
used. To avoid this, any debris or seeds from other species are
cleaned away by hand.
Sample size and thousand seed weight determination
Thousand seed weight (TSW) and the total available number
of seeds are two of the important characters which need recording for
each accession held by a genebank. TSW is determined electronically;
data are recorded in a crop specific 'seed processing file' and later
transferred to the computerized documentation system. The required
sample size for long-term storage is 8000 seeds for genetically hetero-
232
Regassa Feyissa
geneous and 3200 seeds for genetically homogeneous material. For
pragmatic and economic reasons, the final sample size of species with
a TSW greater than 200 g is reduced to a smaller number of seeds.
Samples fulfilling the minimum requirements are subdivided according to their storage status (Table 3).
Seed drying
Seed drying is a complex process and its impact varies according to the nature of the seed and the drying conditions applied. The
purpose of drying the seeds is to minimize the rate of seed deterioration during storage.
The viability of seeds can be significantly affected by overdrying or
exposure to high temperatures. The equilibrium seed moisture content at a given drying temperature depends on the relative humidity
of the ambient air, air flow rate, and the oil content of the seeds. The
higher the oil content the lower the equilibrium seed moisture content due to the hydrophobic nature of lipids (Cromarty, Ellis &
Roberts, 1982). Nevertheless, seeds of various species have different
equilibrium hygroscopic relationships and thus, when exposed to a
given relative humidity, will have different moisture contents after
the equilibrium is reached. By extending the drying period and lowering the relative humidity for those species which have the highest
equilibrium seed moisture content, it is possible to achieve identical
moisture content for seeds of different species (Cromarty et ah, 1982).
When seeds are dried, it is important that the uniformity of the
equilibrium moisture content within a seed lot is kept and this can be
achieved by thin-layer drying or through adequate ventilation.
At PGRC/E seeds are dried under forced ventilation in a drying
room maintained at 18-20 °Cand 18 per cent RH. Samples are dried
in cloth bags or plastic net bags in thin layers on shelves made of wire
mesh in which the air can circulate freely. Initial and final moisture
content is determined in compliance with the International Seed Testing Association (ISTA, 1985) rules, by oven method and, more
recently, with a near-infrared reflectance analyser.
The desired moisture content for cereals and pulses is 4-7 per cent
of the seed weight and for oil crops 3-5 per cent. Dried seeds are
packed and sealed in moisture-proof aluminium foil bags and all
relevant data are entered into the corresponding data file.
Viability test
Viability is the most important attribute of any seed stored in
a genebank. The initial viability test is made after drying just before
Germplasm conservation at PGRC/E
233
long-term storage and this viability is monitored at regular intervals
during storage. ISTA rules (ISTA, 1985) and IBPGR recommendations
(Ellis, Hong & Roberts, 1985) are followed as a guide for selecting
viability assessment procedures for the various species.
The most commonly used method at PGRC/E is the standard
germination test. This is conducted either in a room maintained at a
temperature of about ±20 °C or in an incubator with an alternating
temperature facility. The result of the test is taken as a percentage of
4 x 50 seeds that are able to produce normal seedlings. In cases where
viability is thought to be affected by dormancy, a tetrazolium test is
conducted.
Viability during the course of storage is monitored by using the
sequential germination test method (Ellis, Roberts & Whitehead,
1980) for an 85 per cent regeneration standard using groups of 40
seeds.
Seed storage
The longevity of seeds under storage depends mainly on the
moisture content of the seed and the storage temperature. According
to Harrington's rule of thumb, longevity of seed is doubled for every
1 per cent reduction in moisture content and 5°C in temperature
(Harrington, 1963). Various genebanks maintain their germplasm
under different storage conditions for a number of reasons. At
PGRC/E, seeds meant for both base and active collections are dried to
3-7 per cent moisture content and are kept at — 10°C in laminated
aluminium foil bags. Germplasm accessions which are too small to
fulfil the sample size required for long-term storage are kept in paper
bags at ±4°C and 35 per cent RH. According to the established criteria
these accessions are increased as soon as possible in order to meet the
minimum sample size to allow long-term storage.
Seed distribution
Seeds held in the genebank are distributed mainly for
research work, increase and rejuvenation. Full details on PGRC/E's
germplasm flow policy are given by Kebebew (1988).
At PGRC/E, germplasm is distributed only from active collections
and the number of seeds distributed for research purposes depends
on several factors such as seed size, homogeneity, precise purpose,
etc. For seed increase and rejuvenation, the number of seeds to be
sown depends on the genetic composition of the sample, the amount
of seed available and the required sample size for long-term storage.
234
Regassa Feyissa
Too frequent exposure of germplasm to either rejuvenation or
multiplication is also avoided.
References
Cromarty, A. S., Ellis, R. H. & Roberts, E. H. (1982). The Design of Seed Storage
Facilities for Genetic Conservation. IBPGR, Rome.
Demissie, A. (1988). A decade of germplasm exploration and collection activities by PGRC/E. In: J. M. M. Engels (ed.), The conservation and utilization
of Ethiopian germplasm. Proceedings of an international symposium,
Addis Ababa, 13-16 October 1986, pp. 28-41 (mimeographed).
Ellis, R. H., Hong, T. D. & Roberts, E. H. (1985). Handbook of Seed Technology
for Genebanks, vol. I, Principles and methodology. IBPGR, Rome.
Ellis, R. H., Roberts, E. H. & Whitehead, J. (1980). A new, more economic
and accurate approach to monitoring the viability of accessions during
storage in seed banks. Plant Genetic Resources Newsletter, 41, 3-15.
Harrington, J. F. (1963). Practical instructions and advice on seed storage.
Proceedings of the International Seed Testing Association, 28, 289-94.
International Seed Testing Association (1985). International rules for seed
testing. Seed Science and Technology, 13, 432-63.
Kebebew, F. (1988). Germplasm exchange and distribution by PGRC/E. In:
J. M. M. Engels (ed.), The conservation and utilization of Ethiopian germplasm. Proceedings of an international symposium, Addis Ababa, 13-16
October 1986, pp. 276-84 (mimeographed).
Mekbib, H. (1988). Crop germplasm multiplication, characterization, evaluation and utilization by PGRC/E. In: J. M. M. Engels (ed.), The conservation
and utilization of Ethiopian germplasm. Proceedings of an international
symposium, Addis Ababa, 13-16 October 1986, pp. 170-78
(mimeographed).
Sendek, E. & Engels, J. M. M. (1988). Documentation at PGRC/E. In: J. M. M.
Engels (ed.), The conservation and utilization of Ethiopian germplasm.
Proceedings of an international symposium, Addis Ababa, 13-16 October
1986, pp. 87-96 (mimeographed).
Simmonds, N.W. (1979). Genetic Conservation: an Introductory Discussion of
Needs and Principles. Seed technology for genebanks. IBPGR, Rome, pp. 1-2.
17
Documentation at PGRC/E
ENYAT SENDEK AND J. M. M. ENGELS
Introduction
The quantity and complexity of the information acquired by
the Plant Genetic Resources Centre/Ethiopia (PGRC/E) through active
collecting, donation and repatriation of germplasm require comprehensive and efficient data management systems. Plant genetic
resources can only be successfully utilized if detailed and reliable data
on each accession are available to genebank users, breeders, research
workers and policy makers.
In 1979, a proposal was made to base such a documentation system
on edge-punched cards (Engels, 1979) and this manual system was
used until 1982, by which time the amount of data had reached such
proportions that the system was overcharged and it became necessary to computerize the documentation activities (Engels, 1985).
At present the documentation system at PGRC/E is based on electronic data processing technology and this has greatly facilitated the
handling of the enormous amount of data currently being generated
at the centre.
-
Information sources and descriptor development
The major sources of information in PGRC/E are:
exploration and collection;
germplasm introduction and accessioning;
temporary storage (at +4°C and 30-40 per cent RH);
field genebanks;
multiplication and rejuvenation;
characterization and preliminary evaluation (in field and
laboratory);
seed processing;
Table 1. Sources of information, recording method, number of descriptors involved and relationship with the
documentation division
Source
Exploration/collection
Germplasm introduction and accessioning
Temporary storage (+4°C, 40% RH)
Field genebanks
Multiplication/rejuvenation
Characterization and preliminary evaluation
(field)
Characterization and preliminary evaluation
(laboratory)
Seed processing:
(a) 1000 grain weight and total weight
(b) seed drying and packing
Seed testing:
(a) germination
(b) seed moisture content
Seed storage of active and base collection at
-10°C
Germplasm dispatch
Further evaluation:
(a) cytogenetic studies
(b) Near-Infra-Red analyses
(c) disease resistance screening
(d) environmental stress resistance
screening
(e) taxonomic studies
(f) literature, reports
Utilization
Computer services to users
Library indexing
Miscellaneous
" Computerized.
b
Partly computerized.
Source: Engels, 1986.
Relationship with
documentation
and remarks
Method of recording
original data
Number of
descriptors
Form
Master book
Form
Form
To be developed
Form
24
13
10
8
Crop specific
36
Forms
Crop specific
Passport data"
Reference6
Internal use for redistribution
Short-term storage data"
Field management data"
Climatic, cultivation and soil
data"
Direct entry"
Forms
Crop specific
Direct entry"
Form
Form
7
—
Internal use only
Form
Form
Form
Form
Forms (3)
23
7
7
6
7
Form
Computerized
1
2
Direct entry"
Direct entry"
Long-term storage data"
Partly registration, partly
Report
Lists
Open
2
Direct entry"
Computers not compatible; re
entry"
Extraction''
Direct entry"
Reports
Various
Reports
Computer file
—
Open
Open
Open
—
—
—
Extraction*7
External information1'
External information
Internal use only"
Planned
Mainly internal services
Lists, reports
238
Enyat Sendek & J. M. M. Engels
- seed testing;
- seed storage (base and active collection at —10 °C);
- germplasm flow;
- further evaluation;
- utilization;
- miscellaneous activities.
Further details are presented in Table 1 and can also be found
elsewhere in this volume (Chapters 14, 16, 19).
Data from all these sources are forwarded to the documentation
division, generally on specially designed forms to facilitate further
handling. The information is transformed and handled in the form of
descriptors and their corresponding states. Each descriptor is properly defined and its possible states are chosen in such a way that no
overlapping occurs (International Board for Plant Genetic Resources,
1984). As far as possible, the international norms and recommendations are followed to define and compile descriptors for the various
activities. The descriptors and their definitions for characterization,
preliminary and further evaluation are developed in consultation
with the scientists who utilize germplasm, mainly plant breeders.
The amount of data generated at each stage, i.e. from collection,
through processing and multiplication/characterization, to final
storage, is considerable. To give an example, one accession of wheat
will have 88 items of information or descriptors (32 for passport data,
35 for seed storage and processing and 21 for characterization). This
means that for wheat alone, with a collection of 8500 accessions, a
total of 748 000 data items will be generated. For the complete germplasm holding at PGRC/E it will amount to some 4000000 data items.
To get useful information out of such a vast amount of data, it is
necessary to handle the data systematically in order to meet the needs
of germplasm users as well as the different sections within the
genebank.
Activities of the documentation division
The day-to-day activities of the documentation division are
organized in the following units:
- germplasm accessioning and data acquisition;
- data compilation, preparation, data entry and correction;
- data processing, retrieval and research.
The first unit deals with material newly arrived at the genebank. It
is the responsibility of the germplasm collector or donor to supply
proper and complete data sets to the genebank. The filled data form-
Documentation at PGRC/E
239
sheets are then sorted by genus and species as well as by collection or
donation number before the assignment of a unique number in the
master book. A copy of the information will be forwarded to the
conservation division for the necessary follow-up action. Since the
establishment of PGRC/E, some 40000 accessions have been
registered. They comprise new collections and selections out of the
populations, as well as donations from national and international
institutions (Table 2).
The main task of the second unit is the compilation of the received
and/or actively collected data. These data are converted, combined,
completed, etc., according to the standards established at PGRC/E, by
defining each of the descriptors and their respective states. Furthermore, reference files for the Latin names of the genera and species and another for the administrative units (e.g. regions, awrajas
and woredas) have been developed for easy checking. After the data
set is completed it is entered into the computer and a first printout is
made, to be corrected as necessary by the division concerned.
The third and most important component of the documentation
division is the data management system. Processing and retrieval
activities as well as provision of data requested by plant breeders are
handled and assistance for research operations is given. This unit is
also responsible for the production of seed lists and catalogues in
order to disseminate information to potential users. At the same time,
it greatly facilitates the monitoring function of the conservation division and supports the collection division in planning new collecting
activities. The research activities include, among others, diversity
studies in the various crops based on the evaluation data, as well as
services provided to scientists for the analysis of experimental data.
In general, it can be said that the information management system
supplies the genebank administration and management with the
necessary facts and details for optimal operation.
Data management system
The computers in use at PGRC/E are two HP 125 micro-computers each with 64 Kbytes of memory. Accessories include double
disc drives with 5i-inch floppy diskettes of 248 Kbytes each, a hard
disc Winchester drive, a T-switch and a daisy wheel printer. The
software comprises dBASE II (Ashton-Tate, 1982), WORD/125 (Hewlett
Packard, 1982), MICROSTAT (Anonymous, 1981a), WORDSTAR/125
(Anonymous, 1981b), BASIC/125 (Hewlett Packard, 1981) and STATPAK
(Anonymous, 1982).
240
Enyat Sendek & ]. M. M. Engels
Table 2. Germplasm accessions donated to or collected and/or selected by
PGRC/E as at 30 June 1986
Species
Abelmoschus esculentus
Alframomum korarima
Allium spp.
Amaranthus spp.
Amorphophallus sp.
Arachis hypogaea
Arisaema sp.
Avena spp.
Brassica spp.
Cajanus cajan
Capsicum spp.
Carthatnus tinctorius
Carum copticum
Celosia sp.
Cicer arietinum
Coccinia abyssinica
Coffea arabica
Colocasia sp.
Corchorus olitorius
Coriandrum sativum
Crambe abyssinica
Cucurbita spp.
Cuminum cyminum
Curcuma longa
Cyphomandra betacea
Datura stramonium
Dioscorea sp.
Eleusine africana
Eleusine coracana
Embelia schimperi
Eragrostis tef
Fagopyrum esculentum
Gossypium spp.
Guizotia abyssinica
Guizotia scabra
Helianthus annuus
Hordeum vulgare
Ipomoea batatas
Lablab purpureus
Lagenaria spp.
Lathyrus sativus
Lens culinaris
Lepidium sativum
Linum usitatissimum
Lupinus spp.
PGRC/E
collected
accessions
9
16
28
30
8
15
1
23
668
27
112
75
16
1
728
5
140
21
1
42
1
54
6
1
1
1
13
1
423
8
1067
1
5
638
6
20
3290
46
37
11
156
293
57
538
25
Donated
accessions
0
0
0
0
0
4
0
0
286
0
14
56
0
0
12
0
522
0
0
1
0
12
0
0
0
0
0
6
373
1
1203
0
0
286
0
0
4088
0
0
0
61
134
0
1250
0
Selected
accessions
0
0
0
0
0
0
0
0
0
0
0
0
0
0
162
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1938
0
0
0
0
0
0
32
0
Total
9
16
28
30
8
19
1
23
954
27
126
131
16
1
902
5
662
21
1
43
1
66
6
1
1
1
13
7
796
9
2270
1
5
924
7
20
9316
46
37
11
217
427
57
1820
25
Documentation at PGRC/E
241
Table 2 (cont.)
Species
PGRC/E
collected
accessions
Donated
accessions
Selected
accessions
Total
Lycopersicon spp.
Medicago sativa
Moringa stenopetala
Myrsine africana
Nicotiana tabacum
Nigella sativa
Ocimum spp.
Oryza spp.
Oxytenanthera abyssinica
Pennisetum typhoides
Phaseolus spp.
Phytolacca dodecandra
Pimpinella anisum
Piper longum
Pisum sativum
Plectranthus edulis
Raphanus sativus
Ricinus communis
Rumex abyssinica
Ruta chalepensis
Sesamum indicum
Solarium incanum
Sorghum bicolor
Tamarindus indica
Trigonella foenum-graecum
Triticum spp.
Vernonia spp.
Vicia faba
Vigna unguiculata
Voandzeia subterranea
Zea mays
Zingiber officinale
Unknown
Grand total
4
2
2
1
0
28
13
24
1
5
256
144
3
3
702
6
5
210
1
1
218
8
1144
0
224
2376
22
725
35
1
179
48
6
15 062
15
3
0
0
24
0
0
0
0
10
7
0
0
0
420
0
0
62
0
0
158
0
5419
1
214
3890
0
473
0
0
7
0
0
19 012
0
0
0
0
0
0
0
0
0
0
0
0
0
0
11
0
0
0
0
0
0
0
1582
0
0
2114
0
0
0
0
0
0
0
5839
19
5
2
1
24
28
13
24
1
15
263
144
3
3
1133
6
5
272
1
1
376
8
8145
1
438
8380
22
1198
35
1
186
48
6
39 913
In order to increase efficiency in the data management, a general
data base system has been designed in which each description has its
own fixed place and which allows extension of the number of descriptors and the number of accessions crop wise all the time. The main
characteristics of the general data base are:
- the accessions are grouped by crop;
- the descriptors are grouped according to main activities (e.g.
passport data, conservation and evaluation);
242
Enyat Sendek & J. M. M. Engels
Table 3. Files and types of data presently kept by the documentation
division of PGRC/E
File name
Number of
files
Type of data
PGRCE
SEED:STR
SEED:CHR
1
1
30
PGRCCOLN
1
SEED:REF
LOCIREF
SEEDIDES
SEED:REC
DESC:DEFN
1
1
1
1
1
Passport
Seed storage and testing
Characterization and
evaluation
Crop specific
Summary of PGRC/E
holdings
Crop reference
Locality reference
Seed dispatch
Seed acquisition
Descriptor definitions
Number of Number of
descriptors records
24
25
9-29
39978
27755
14110
6
78
5
3
12
11
8
119
552
41
28
233
- new accessions can be added without restrictions (vertical
increase);
- new descriptors can be added without limitations (horizontal
increase).
The files at present in use within the PGRC/E documentation division are listed in Table 3. Each contains a specific type of data.
The passport data base at present contains data on almost 35 000
accessions of approximately 75 different crops or groups of crop
types. For each accession, 24 descriptors are recorded: accession
number, crop name, genus, species, subspecies, local name/cultivar,
country of origin, administrative region/state, awraja/district,
woreda/area, village/locality, latitude, longitude, altitude, collecting
institute, collection team/collector, collector's number, donor number, collection date (day, month, year), collection source, sample type
and genetic status.
Some applications of the data management system
The importance of the documentation system for the
genebank can be readily illustrated by listing its uses:
- seed inventory, monitoring and handling (seed storage and
testing file);
- assisting the exploration and collection division to plan collecting missions by area, crop and time (passport file);
- assisting the respective divisions with planning and decision
Documentation at PGRC/E
-
-
243
making in germplasm rejuvenation, multiplication and/or
characterization (seed storage and characterization files);
facilitating exchange of germplasm through the publications
of germplasm lists (passport file);
supporting plant breeders' use of germplasm by analysing
the data and publishing the available information in crop
catalogues (characterization/evaluation files);
helping the users of germplasm to make a first (rough) selection from the available germplasm based on specific criteria
(characterization/evaluation files);
analysing the available information on accessions and collection sites to predict possible areas where germplasm with
specific traits can be found (passport and characterization/
evaluation files);
carrying out taxonomic analyses and classification using the
characterization data;
identifying of duplicate accessions, unknown accessions, etc.
Publications and library
The PGRC/E-ILCA Germplasm Newsletter is published
jointly with the Forage Legume Agronomy Group (FLAG) of the
International Livestock Centre for Africa (ILCA) and produced on the
PGRC/E computing system. Its first issue appeared in December
1982. The newsletter is circulated both nationally and internationally
to more than 2000 addresses and has already shown its value as a link
between the germplasm centres and the users of these genetic
resources worldwide.
The PGRC/E library has a considerable number of specialized
books in stock, as well as a collection of reprints in the field of plant
genetic resources.
References
Anonymous (1981a). MICROSTAT User's Manual. Lifeboat Associates, New
York.
Anonymous (1981b). WORDSTAR/125 Reference Manual. Micropro International
Corporation, San Rafael, California.
Anonymous (1982). TheNWA STATPAK, Version 2.1 Preliminary Manual. North-
west Analytical Incorporated, Portland, Oregon.
Ashton-Tate (1982). dBASE II Version 2.3B Assembly-Language Relational Database
Management System. Ashton-Tate, Culver City, California.
Engels, J. M. M. (1979). Proposal for a documentation system for the Plant
Genetic Resources Centre at Addis Ababa, Ethiopia. GTZ, Eschborn
(mimeographed).
244
Enyat Sendek & J. M. M. Engels
Engels, J. M. M. (1985). Documentation and information management at
PGRC/E. PGRC/E-ILCA Germplasm Newsletter, 9, 20-7.
Engels, J. M. M. (1986). The documentation at the Plant Genetic Resources
Centre/Ethiopia. Ada Horticulturae, 182, 387-92.
Hewlett Packard (1981). BASIC/125. Hewlett Packard, Sunnyvale, California.
Hewlett Packard (1982). WORD/125. Hewlett Packard, Cupertino, California.
International Board for Plant Genetic Resources (1984). Annual Report.
IBPGR, Rome.
Part IV
Evaluation and utilization of
Ethiopian genetic resources
18
Germplasm evaluation with special
reference to the role of taxonomy in
genebanks
J.G. HAWKES
Introduction
Of all the varied activities of genetic resources centres, that of
evaluation is probably the most neglected. Genetic resources centres
in general carry out excellently their tasks of exploration, accession of
samples and all the complex processes of seed storage and data
management. They perhaps do not pay enough attention to surveys
and many only undertake the initial stages of evaluation, thus to
some extent hindering the subsequent processes in the management
chain (Fig. 1). This means that the important function of a genetic
resources centre may be deflected, that is to say, the material in it may
not be made completely available to the breeders and thus may run
the risk of not being incorporated into their new selections and
varieties. In this way much of the money and effort expended in
exploration and conservation is in danger of being wasted and the
very existence of the genebank itself may be threatened (Hawkes,
1985a).
Evaluation
To understand the above assertion we should look more
closely at the activities of germplasm evaluation in the broad sense.
The International Board for Plant Genetic Resources (IBPGR) has
usefully divided the process of evaluation into three main categories
(Erskine & Williams, 1980).
- Characterization. This includes the scoring of morphological
and agronomic characters of high heritability, not likely to
248
/. G. Hawkes
Genetic resource surveys
Exploration
Conservation
Research
Evaluation
Germplasm enhancement
Breeding: trials
National programmes
Release of varieties
Well-being and economic advancement of 3rd World farmers and countries
Fig. 1. Genetic resources impact chain. In the central sequence
each activity exerts a direct impact on the ones following it. Thus
the development of the lower levels of activity is influenced
positively or negatively in accordance with the efficient or nonefficient development of those above them (Hawkes, 1985a).
change very much under different environmental conditions.
Preliminary evaluation. This category comprises a limited
number of agronomic traits thought to be desirable by a consensus of users of the crop in question.
Full evaluation (also termed secondary, in-depth or further
evaluation). This concerns the scoring of characters conferring tolerance, resistance or immunity to the pests and
diseases that attack a particular crop. It also includes characters of adaptation and resistance to stress conditions such as
Germplasm evaluation and taxonomy
249
frost, cold, heat and drought, and to adverse soil conditions
such as high acidity, salinity, aluminium, sulphates, etc.
The work in each category needs to be discussed in detail and
certain difficulties noted.
Under the first category - characterization - a series of committees
convened by IBPGR has advised on the publication of a set of crop
descriptor lists. These enable standardized characterization of crop
after crop to be made in different parts of the world in a uniform
manner by the use of the descriptor and descriptor states listed. Such
standardization is clearly to be welcomed, but some words of warning should be given.
Many of the descriptor lists are very long and genebank staff may
perhaps spend too much time on these activities at the expense of
others that ought to be given a higher priority. Genebank staff may
feel uneasy about omitting some of the descriptors. However, IBPGR
itself has reduced the number of recommended descriptors in its
second edition lists. Thus the first wheat and Aegilops list of 1978
included 26 morphological and 29 agronomic descriptors - a total of
55. The second edition (1981) reduced the list to 12 (6 plus 6), less than
a quarter of the original descriptor number. The change has been
brought about largely by the breeders themselves, who became aware
that many of the characters scored were of very little value to them in
selecting initial material for their breeding programmes. This point
will be referred to later.
It has also been said that a long list of morphological characters will
help the genebank staff to recognize a particular sample if it loses its
label or if it is suspected that it has become mixed with, or been
replaced by, another sample. In general, however, a series of voucher
herbarium specimens, cereal spikes and tubes of seeds or grains will
be more useful than any written descriptions to identify possible
errors and mixtures.
In addition, it has been argued that a large morpho-botanical descriptor list will help genebank staff identify duplicates with the aid of
a computer-generated study. The International Potato Centre (CIP)
used this method with great success to identify identical morphotypes in their clonal potato collection. Fifty-four morpho-botanical and 14 agronomic characters were used to reduce a collection of
some 15000 entries to about one-third. It must be stressed that the
method would be difficult to apply to outbreeding seed crops but
would probably be useful in certain cases for inbreeders. It could not
easily be applied to population samples.
250
/. G. Hawkes
In the preceding paragraphs it has been assumed that both the
morphological and agronomic characters were of high heritability.
This cannot always be assumed, since the differences between high
and low heritability characters are not very clearly defined. A decision
'can only be made on the basis of a profound knowledge of a particular crop' (Erskine & Williams, 1980). Evidence of the difficulties
encountered may be quoted from Tyler (1985) working with outbreeding forage grasses. He states that because of the quantitative
nature of inheritance, the characters are often not easily seen, and
most of them are influenced, often quite strongly, by environmental
factors. He goes on to say that the majority of characters used for the
registration of new varieties have 'little relevance to agricultural performance or indeed to breeding objectives'. Traits such as
inflorescence emergence date, vegetative habit and habit at flowering
have more relevance, for instance, than the length and width of the
flag leaf, ear length and other characters. Differences in the expression of many agronomic characters due to season and locality may
also be strongly marked. Such high genotype-environment interactions are inevitable for many characters. To overcome this to some
extent, all screening must bear full site data as well as seasonal or
weather data wherever possible.
The second category - preliminary evaluation - may include other
characters that are not used in characterization, but the difficulties
encountered in the selection of characters of high heritability still
remain.
Full, secondary or in-depth evaluation is considered by IBPGR to
be the task of the breeders, the two previous activities being undertaken by genebank staff.
In an ideal world, the breeders might be able to undertake secondary evaluation. In the real world, this is not often attainable because
of pressure of work from existing programmes or through lack of
interest in genebank material. A happy situation exists in Ethiopia
where breeders are encouraged to examine grown-out genebank
material in the experimental field and to make notes of promising
lines for crossing and performance trials. They take advantage of this
opportunity in a most satisfactory manner. Nevertheless, it has to be
said quite frankly that this is an exception, and in any case it refers
only to field inspections and not to laboratory or glasshouse
screening.
Normally, the genebank staff do not possess the equipment or the
time to undertake secondary evaluation. Nevertheless, it should be
one of the clear responsibilities of a genebank manager to arrange for
Germplasm evaluation and taxonomy
251
secondary evaluation to be carried out. Admittedly, this is not always
easy, but where national facilities exist in universities, sister institutes
or even in commercial companies, screening for resistance to pests,
diseases and environmental stress can be arranged free of charge. The
collaborators receive publishable data and the genebank manager
obtains results to add to his inventories.
A more difficult problem arises when materials need to be sent
abroad for screening because of lack of national facilities. Political
pressures may have prevented this in the past but with the new Food
and Agriculture Organization (FAO) convention on the free international exchange of materials, few or no difficulties should arise in
transmitting materials to other countries for screening. Most
laboratories in developed countries would probably charge a fee for
this work, but it should not be too difficult to obtain bilateral or
multilateral aid funding to cover the costs of screening and postage.
When the screening results are sent back to the genebank they
must be entered into the computerized data base and inventories
with such results should be published with the least possible delay.
Some authorities question the need for the publication of inventories,
stating that if a breeder wants material with certain characters he can
always ask for the appropriate information from the data base. In
theory this could be so, but in practice the genebank manager is the
one who needs to take the initiative. After contacts have been made
and the breeder has received useful material he will probably ask for
more, but the genebank manager should make the first move by
printing and distributing inventories. Simple accession lists will not
do; the hard facts of the screening results must also be available.
These points are stressed by Roelofsen (1985), Hawkes (1985b) and
van Soest (1985) who, incidentally, all took part in a recent
symposium entitled 'Evaluation for the Better Use of Genetic
Resources Materials' held in Prague, Czechoslovakia (Rogalewicz,
1985), and all stress the importance of secondary evaluation as seen
by European breeders and genetic resources personnel. It is of interest to note that Peeters & Williams (1984) also stress the need for
secondary evaluation, based on data provided by breeders.
Pre-breeding
When a breeder is convinced that a certain genebank accession contains characters of value to his programme, he will have no
hesitation in using it, providing, however, that its agronomic characters are on or near the same level as the materials he is already using.
If the valuable characters are to be found only in unproductive or
252
/. G. Hawkes
poor quality landraces, or in related wild species, he may hesitate to
use them because the high-level features of his breeding stocks will be
diminished by contamination with the unacceptable low-level characters which will be introduced from the non-elite line together with the
single feature in which he is interested. Such contamination would
set back his programme for several years and he will, understandably, be reluctant to use this material. It would then be the responsibility of the genebank personnel to look for the same useful feature in
other genebank accessions that might at the same time possess better
agronomic characters. If this is not possible, then a programme of
pre-breeding, or 'germplasm enhancement' as it is often called, will
be needed. By this is meant the transference of the useful gene or
genes into good agronomic lines that would be more acceptable to the
breeder. But who is to do this pre-breeding? It depends very much on
the circumstances of the country concerned. Often a university
department or laboratory can be asked to cooperate and the work can
be undertaken partly by research students. Sometimes a breeding
institute is interested or the breeders themselves are enthusiastic
enough to carry out the work. On a world scale, however, but with
notable exceptions, it must be admitted that secondary evaluation
and pre-breeding constitute 'bottlenecks', thus preventing the
adequate utilization of genetic materials.
Taxonomy
A taxonomic approach, in the wide sense, to genetic
resources work is essential. Beginning with the survey work (Fig. 1),
it is essential to know the nature and names of the materials for which
the genebank is responsible. This perhaps hardly needs saying.
Secondly, it is essential to know where they are to be found and
whether cultivated or wild. Thirdly, we must know the taxonomic
system of a crop. Is it a single species, as in Secale; is it a group of
species at different ploidy levels, as in Triticum; does it possess a
series of wild or weedy forms related to it, evolving in a parallel
manner and exchanging genes from time to time through natural
hybridization, as in Sorghum; does it have no closely related wild
relatives, as in Vicia faba; or does it have something of all these situations? To answer some, at least, of these questions the crop plant
taxonomist must possess not only a solid background of classical
taxonomy but also experience and/or an interest in crop plant taxonomy and in the related disciplines of cytogenetics, phytochemistry,
numerical taxonomy, reproductive biology and ecology. In short, tax-
Germplasm evaluation and taxonomy
253
onomy functions as a filing system in which all other data on
resistance, adaptation, crossability, cytogenetics, etc. can be stored.
For such a system to be reasonably useful it should be simple, logical
and stable (Hawkes, 1980).
On an immediately practical level the taxonomist must identify the
materials accessed into the genebank and if necessary also classify
them at the infraspecific level. This may be done by using the categories of formal taxonomy (Parker, 1978) or by confining these to the
species and subspecies only and using an informal group method for
the lower categories at infraspecific level (Hawkes, 1986). Whatever
infraspecific classification is used for cultivated plants it is certain that
a large amount of genetic diversity at this level cannot be encompassed in such categories. For this we must use the descriptor and descriptor state methods as mentioned above.
Formal taxonomy is obviously necessary, but we must go beyond it
to an understanding of reproductive biology which develops from the
methods of what is generally known as 'experimental taxonomy'. For
instance, breeders need to know about the possibility of gene transfer
and the presence, if any, of genetic incompatibility or 'incongruity'
barriers between species. Such barriers exist, for instance, between
maize and Tripsacum, which can cross with difficulty but have many
problems in their hybrids. On the other hand, maize and teosinte
cross readily and produce viable offspring - a fairly predictable result
if we assume (as many people now do) that they belong to the same
genus (Zea mays and Z. mexicana) and that the latter may even be the
wild prototype of the former. We can see a similar situation in diploid
wheats. The cultivated species, Triticum monococcum, crosses very
readily with T. boeoticum, its wild progenitor, and the hybrids suffer
no loss of fertility. In fact, some authorities class them as two subspecies of the same species.
It will be of value at this point to mention Harlan & de Wet's (1971)
gene pool hypothesis (see also Harlan & de Wet, 1986). Their idea is
that for any species of cultivated plant the primary gene pool (GP-1)
represents the concept of the biological species {Triticum in the example mentioned in the previous paragraph). The secondary gene pool
(GP-2) includes individuals that can be crossed with GP-1 although
there are distinct partial genetic barriers present (maize and Tripsacum, quoted above). Tertiary gene pool species (GP-3) could only
yield hybrids with GP-1 that were completely sterile. Related species
with different genome compositions from those of GP-1 might fit into
this category.
254
/. G. Hawkes
It would be most interesting to investigate some of the lesser
known Ethiopian species and genera from this point of view and, in
particular, the assumed wild prototypes. In this way the materials in
GP-2 and GP-3 might be made more available to breeders.
Chemotaxonomic studies are also of great interest and could form
some of the research projects of genebank personnel. Relationships
between species might be investigated by comparative serology and
immuno-electrophoresis and by two-way electrophoresis of leaf
phenolics. Isozyme analyses may prove illuminating by showing
areas where the greatest amount of isozyme diversity occurs and
linking these with results for disease and pest resistance screening.
Results from all aspects of this kind of work can be assessed by means
of the usual computer-aided techniques (see also Yndgaard &
Hoskuldsson, 1985).
Two words of warning are necessary here. First, although the
diversity of isozymes may be intensive in certain areas, does this
mean that useful genes for resistance to pests and diseases may also
be found with greater intensity in these same areas? No-one has ever
looked into this problem and indeed few have even asked the question. Yet the question must be asked and answered. Otherwise,
isozyme studies, useful as they are from a theoretical viewpoint, may
turn out to be of no practical value whatsoever.
Secondly, no matter how interesting scientifically these results
may be, the acid test is to show whether gene transfer from one
species to another can be accomplished. Thus, if by principal components analysis of isozyme results we can show that a certain accession of a possibly related wild species seems to be closely similar to
the cultigen, the question still remains. Will the two taxa hybridize
and if so can the transfer of useful genes be effected from the wild
relative to the crop itself? For this answer we must always go back to
crossability studies, the cytogenetical analysis of the progenies and, if
necessary, pre-breeding work.
Apart from this extremely important function of taxonomy the
morpho-geographical approach is still essential. If we find useful
resistance in one area, as Qualset (1975) describes for barley yellow
dwarf virus disease resistance in Ethiopia, then this is the area to
which we should return to collect further samples. It seems highly
likely that the evolution of this type of resistance took place in such an
area, and samples should be made at points where the proportion of
resistant to susceptible plants is greatest - in the case of barley yellow
dwarf virus, apparently at elevations higher than 3600 m above sea
Germplasm evaluation and taxonomy
255
60
120
40
40'
30-
20-
-20
10-
— —
Chief concentration of
Phytophthora hypersensitivity
genes
Subsidiary areas of
hypersensitivity genes
—
Chief concentration of genes
conferring resistance to
virus Y and various insects
Eelworm resistance genes
500
1000
1500 miles
Fig. 2. Gene mapping in potatoes. This map represents an early
attempt at mapping some of the genes conferring resistance to
certain potato diseases and pests (Hawkes, 1958).
level, according to Qualset. Evolution of character complexes linked
to the resistance gene also points to materials on which maximum
efforts should be concentrated.
A similar example can be quoted for the resistance of wild potatoes
to various biotypes of the potato round cyst nematode. Since potatoes
are outbreeders, we are here concerned with population sampling
techniques. Populations from some sites may possess low frequencies
of resistance genes, others medium and yet others quite high
256
/. G. Hawkes
frequencies. The breeder is much more interested in high frequencies
of resistance alleles, particularly if they occur in the homozygous
state, because these can be transmitted to a large number of plants in
his crossing progenies.
The lesson we learn from this is to identify carefully the species,
subspecies, etc. and the geographical area where genes for resistance,
adaptation, etc. have been found. This, of course, follows along the
lines of Vavilov's Law of Homologous Series (1926). When such species and areas have been identified we should then make really
detailed collections (fine-grid) from those areas and from related species in the same places, if such species exist. Vavilov's Law points out
the geographical regularities to be found in such situations and the
author has found it to be true for potatoes in the Americas (Fig. 2).
Thus, we can begin to build up an edifice of knowledge and understanding with the interest and close cooperation of specialist evaluators and breeders. The value of genetic resources work becomes
clear to all and its function in the breeding of new varieties is greatly
enhanced and firmly established.
In such ways we are able to coordinate theory and practice to the
advantage of genebank work and to show that a genetic resources
centre is not merely a place where seeds are stored but a centre of
excellence in plant evolution and breeding research.
References
Erskine, W. & Williams, J.T. (1980). The principles, problems and responsibilities of the preliminary evaluation of genetic resources samples of
seed-propagated crops. Plant Genetic Resources Newsletter, 41, 19-33.
Harlan, J. R. & de Wet, J.M.J. (1971). Toward a rational classification of
cultivated plants. Taxon, 20, 509-17.
Harlan, J. R. & de Wet, J. M. J. (1986). Problems in merging populations and
counterfeit hybrids. In: B.T. Styles (ed.), Infraspecific Classification of Wild
and Cultivated Plants. Oxford University Press, pp. 71-6.
Hawkes, J. G. (1958). Significance of wild species and primitive forms for
potato breeding. Euphytica, 7, 257-70.
Hawkes, J. G. (1980). The taxonomy of cultivated plants and its importance in
plant breeding research. In: Perspectives in World Agriculture. Commonwealth Agricultural Bureaux, Farnham Royal, UK, pp. 49-66.
Hawkes, J. G. (1985a). Plant genetic resources. The impact of the international agricultural research centres. CGIAR Study Paper No. 3. World Bank,
Washington DC.
Hawkes, J. G. (1985b). Genetic resources evaluation. An overview. In: V.
Rogalewicz (ed.), Evaluation for the Better Use of Genetic Resources Materials.
Research Institute of Plant Production, Prague.
Hawkes, J. G. (1986). Problems of taxonomy and nomenclature in cultivated
Germplasm evaluation and taxonomy
257
plants. In: L. J. G. van der Maesen (ed.), First International Symposium on the
Taxonomy of Cultivated Plants. Ada Horticulturae, 182, 41-52.
Parker, P. F. (1978). The classification of crop plants. In: H. E. Street (ed.),
Essays in Plant Taxonomy. Academic Press, London, pp. 97-124.
Peeters, J. P. & Williams, J. T. (1984). Towards better use of genebanks with
special reference to information. Plant Genetic Resources Newsletter, 60,
22-32.
Qualset, C O . (1975). Sampling germplasm in a centre of diversity: an example of disease resistance in Ethiopian barley. In: O. H. Frankel and J. G.
Hawkes (eds), Crop Genetic Resources for Today and Tomorrow. Cambridge
University Press, Cambridge, pp. 81-96.
Roelofsen, H. (1985). Using evaluation data: an information problem. In: V.
Rogalewicz (ed.), Evaluation for the Better Use of Genetic Resources Materials.
Research Institute of Plant Production, Prague, pp. 167-73.
Rogalewicz, V. (ed.) (1985). Evaluation for the better use of genetic resources
materials. Proceedings of the Eucarpia Genetic Resources Section international
symposium. Research Institute of Plant Production, Prague.
Tyler, B. F. (1985). Evaluation of forage grass genetic resources for characterization and breeding potential. In: V. Rogalewicz (ed.), Evaluation for the
Better Use of Genetic Resources Materials. Research Institute of Plant Production, Prague, pp. 215-24.
van Soest, L. J. M. (1985). Some aspects concerning the better use of germplasm collections. In: V. Rogalewicz (ed.), Evaluation for the Better Use of
Genetic Resources Materials. Research Institute of Plant Production, Prague,
pp. 189-206.
Vavilov, N.I. (1926). Studies on the origin of cultivated plants. Bulletin of
Applied Botany, Genetics and Plant Breeding, 16, 1-248.
Yndgaard, F. & Hoskuldsson, A. (1985). Electrophoresis: a tool for
genebanks. Plant Genetic Resources Newsletter, 63, 34-^10.
19
Crop germplasm multiplication,
characterization, evaluation and
utilization at PGRC/E
HAILU MEKBIB
Introduction
The majority of the germplasm accessions maintained by the
Plant Genetic Resources Centre/Ethiopia (PGRC/E) are landraces
which have evolved under local conditions in the farmers' fields since
time immemorial. Such gene pools are the reservoirs of variation
which provide the raw material for crop improvement. Samples in the
form of seeds or whole plants, representing the spectrum of genetic
variation within cultivated species and their wild relatives, are currently being collected and maintained in seedbanks and field
genebanks throughout the world (Frankel & Hawkes, 1975; Williams,
1984). Of fundamental importance in the management of these
resources is the determination of the variation they represent. To this
end, characterization of the various crop germplasm collections is
undertaken by the multiplication, characterization and evaluation
division of PGRC/E in close collaboration with the plant breeders.
In the past, characterization activities were limited in scope and
greater attention was given to collection and conservation activities.
During the last three or four years, however, the priorities have
changed and now include the extension and intensification of characterization and evaluation work, as well as support for the utilization
of germplasm. Highest priority has been given to the major economic
crops (e.g. cereals, pulses and oil crops) with the aim of providing
useful materials for the breeding programmes.
Crop germplasm multiplication and rejuvenation
Because of the earlier priorities, a systematic increase of the
Crop germplasm multiplication and characterization
259
collected germplasm accessions did not start until 1982. In order to
cope with the considerable backlog, the division has had to handle
more than 8000 accessions annually up to now. Since the size of the
collected samples is generally not large enough to meet the minimum
requirements for long-term storage, seed increase is a necessity. Furthermore, during the course of storage, a considerable number of
seeds will be used for viability monitoring, research and evaluation
work, etc. and this will lead to the next cycle of increase. By knowing
the multiplication factor for the various crop species, as well as the
initial amount of seed and the genetic structure of a given accession,
the actual required sample size for multiplication can be determined
in order to fulfil the need for safe and adequate conservation. The
larger the original sample, the easier and the safer its multiplication
and evaluation will be (Frankel, 1970).
Where germplasm accessions need only to be multiplied or
rejuvenated, only a few characters, such as flower colour, flowering
date and growth habit, are recorded to ensure the identity of the
accession with the original sample and to avoid or trace possible
mistakes.
Rejuvenation of germplasm accessions is needed when there is a
significant decrease in the germination percentage. This activity is
conducted under the same conditions used for seed increase, i.e.
growing conditions, with maximum care being taken so that no, or as
little as possible, natural selection can take place to alter the identity
of the accession. Multiplication methods appropriate to the species
and cultural practices that maximize the yield of qualitative seeds are
being used.
Germplasm multiplication and characterization sites
The increase and rejuvenation of germplasm should take
place principally where the accessions were collected, or under
similar conditions (Hawkes, 1985). The gene frequencies of each
character should remain unchanged from generation to generation,
i.e. selection pressure should be minimized.
PGRC/E is in a favourable situation to deal with germplasm which
is almost entirely of Ethiopian origin. Therefore, and because of the
Ethiopian topography, no real problems exist in the multiplication
and characterization of crop germplasm in its natural habitat or under
very similar conditions (Table 1). The accessions of a given crop are
divided into groups according to the altitude obtained from the passport data and are evaluated accordingly at sub-stations in corresponding altitudinal zones. For example, highland sorghum is evaluated at
260
Hailu Mekbib
Table 1. Multiplication sites of PGRC/E with altitude, average daily
temperature and average annual precipitation
Sites
Altitude
(m)
Temperature
(°Q
Precipitation
(mm)
Addis Ababa
Arsi Negele
Asmara
Awassa
Bekoji
Debre Zeit and Dembi
Fitche
Ginchi
Holetta and Kuyu
Jima
Kulumsa
Melkassa
Melka Werer
Mieso
2450
1960
2325
1750
2850
1900
2800
2240
2390
1577
2200
1558
737
1320
16.0
18.4
16.6
19.2
12.9
18.7
12.9
17.0
13.8
18.8
16.3
21.5
26.3
22.7
1163
1763
525
961
1203
866
1283
1075
1097
1469
938
806
471
694
1960 m above sea level at Arsi Negele, intermediate sorghum types at
1580 m at Melkassa and lowland types at 1320 m in Mieso.
Procedures followed and difficulties encountered
The most important goal of any seed increase activity is to
produce healthy seeds without changing the genetic composition of
an accession. Therefore, different approaches have to be followed
according to the flower biology of a species, the genetic composition
of an accession, the total number of accessions to be multiplied, the
initial status of an accession (sample size, health condition), etc.
The biological characteristics of self-pollinated crops make the task
of handling germplasm relatively easy. In general, the plants of a
given accession do not require any additional input and the seeds can
be harvested from the whole plot. However, some of the selfpollinated crops have shown a low percentage of cross-pollination.
Therefore, plans have been made to adjust the procedures for
self-pollinated crops in such a way that the chance of occasional
outcrossing will be reduced to nearly zero. A second group of species
comprises those crops which are predominantly cross-pollinators.
These species require special isolation techniques and, in addition,
extra attention has to be paid to the (artificial) pollination of sufficient
plants per accession (Demarly, 1981). Some of the problematic crops
are presented below as an illustration.
Crop gertnplasm multiplication and characterization
261
Niger seed or noog (Guizotia abyssinica)
This annual oil crop belongs to the Compositae family. The
species is pollinated predominantly by insects and is predominantly
genetically self-incompatible. This is mainly because the receptive
part of the stigma rarely touches the pollen of the floret (Seegeler,
1983). Since selfing of this species would cause serious inbreeding
depression and might expose the accessions to unwanted natural
selection, a form of sibbing within each accession is applied. A sufficiently large number of plants are covered with cheesecloth bags and
at regular intervals hand pollination between plants within the accession takes place. The use of pollinating insects within a bag is under
study and could increase the degree of seed setting.
Brassica species
Brassica carinata, B. nigra and B. oleracea are species either
indigenous to Ethiopia or widely grown. All are predominantly crosspollinated and therefore require special attention to avoid gene flow
between accessions. In this case also, cheesecloth bags are used to
isolate a group of plants of the same accession and natural pollination
within a bag is used to obtain the required seed amount. Problems
with fungi and low seed setting are still hampering an adequate
increase procedure.
Faba bean (Vicia faba)
This generally self-pollinating species shows, under ideal
conditions, up to 50 per cent outcrossing, caused mainly by bees. The
isolation method presently in use is based on a Brassica fence around
each accession. The assumption is that the pollinating bees stay
mainly within a plot and before approaching another plot (=another
accession) they will be attracted by the Brassica plants where the faba
bean pollen will be 'brushed off. Although this method does not
ensure the avoidance of gene flow between accessions it is relatively
simple and cheap and thus large numbers of accessions can be
increased.
Castor bean (Ricinus communis)
This monoecious species is a wind pollinator. Although selfing might cause inbreeding depression to a certain extent, individual
inflorescences are bagged to avoid cross-pollination. The same procedure is followed with sorghum.
262
Hailu Mekbib
Characterization and evaluation
Since the value of the conserved germplasm will depend
greatly upon the information available on each accession, high priority is given to the systematic characterization work. More than 20
descriptor lists for the different crop types have been discussed and
agreed upon by the respective plant breeders and are presently in
use. The descriptor lists issued by the International Board for Plant
Genetic Resources (IBPGR) are followed, as far as they are relevant to
Ethiopian conditions, to record morphological and preliminary evaluation data on various crop types.
By 1986 a total of about 35000 accessions of more than 25 crop
types had been multiplied and characterized. Details of the number of
descriptors used for each of the crops as well as the respective
multiplication sites are presented in Table 2. Priorities for the crops to
be multiplied were based on various criteria which were defined
ahead of time. Major aspects taken into consideration were the
degree of dependency of the plant breeder on local germplasm, the
economic importance of the crop in Ethiopia, the amount of diversity
found or expected in the crop, the sample size and viability condition
of the individual accessions, etc.
The genetic composition of an accession has far-reaching consequences for the characterization and evaluation of germplasm.
Genetically homogeneous accessions of self-pollinating crops are easy
to handle and are not any different from medium uniform varieties.
However, genetically heterogeneous accessions cause many complications to a genebank and they frequently require an extra treatment.
To overcome such complications and difficulties as far as possible,
PGRC/E has started to separate heterogeneous accessions or landraces of self-pollinated crops, such as wheat and barley, into agromorphological components. Each component has some obvious and
important agronomic and taxonomic characters in common although
the individuals may vary in other characters. Such components are
treated as pure lines following the procedures described below.
During the first year of increase, spikes with highly heritable
characters in common within an accession are combined to form a
component. The next season, five ears from each component are
planted ear to row. Rows and components are compared with each
other and where there is no difference they will be lumped. Distinguishable components will receive new accession numbers and be
treated as independent samples.
The characters of highest interest to the plant breeders are
Table 2. Crop germplasm multiplication and characterization by PGRC/E in the period 1982-6
Total number of accessions
planted and characterized
Crop type
1982
1983
1644
225
Barley
Sorghum
Teff
Finger-millet
Pearl millet
424
2036
—
—
—
Oil crops
Rapeseed
Noog
Linseed
Safflower
Sunflower
Sesame
Castor bean
Cereals
Wheat
Legumes
Faba bean
Field pea
Lentils
Chickpea
Fenugreek
Lathy rus
Phaseolus
Cowpea
Lablab
Total
1984
Total
Number of
descriptors
employed
Multiplication/
characterization
sites
1985
1986
1777
2898
2595
9139
20
716
2991
350
—
—
4201
500
2400
—
—
2943
953
36
210
11
2510
1777
472
316
8
10794
8257
3258
526
19
20
21
20
16
27
Asmara, Holetta, Debre Zeit,
Combolcha
Holetta, Kulumsa
Arsi Negele, Melkassa
Debre Zeit
Melkassa
Melkassa
303
243
193
75
—
90
-
264
184
300
103
11
104
-
438
344
177
35
8
81
200
280
376
298
7
2
120
49
191
300
83
—
—
92
—
1476
1447
1051
220
21
487
249
27
23
20
21
4
28
26
Holetta
Holetta
Holetta
Debre Zeit, Melka Werer
Awasa, Arsi Negele
Melka Werer
Arsi Negele, Awassa
102
153
168
321
—
—
—
—
—
5752
143
34
—
—
24
—
—
—
—
5449
176
125
145
335
—
—
—
—
—
10942
80
415
214
224
99
74
157
23
30
9499
262
296
130
150
—
—
93
20
25
9320
763
1023
657
1030
123
74
250
43
55
40962
13
14
13
14
13
15
19
19
19
Debre Zeit, Bekoji
Bekoji
Debre Zeit
Debre Zeit, Ginchi
Debre Zeit
Debre Zeit
Melkassa
Melkassa
Melkassa
264
Hailu Mekbib
generally less heritable or their determination needs special
experimental designs or complicated equipment. The activity to
record such characters is called further evaluation and is the principal
responsibility of the plant breeder. However, PGRC/E has initiated
several further evaluation activities, in close contact with the breeder
concerned, in order to increase the value of its germplasm. Examples
include the evaluation of some 1700 sorghum accessions for their
agronomic performance and for possible resistance or tolerance to
bacterial leaf streak and stalk borer. These activities were conducted
in collaboration with scientists from the International Crops Research
Institute for the Semi-Arid Tropics (ICRISAT) and Addis Ababa
University.
In 1983, PGRC/E and the Scientific Phytopathological Laboratory in
Ambo started a pilot project to screen durum wheat accessions for
resistance to stem rust, stripe rust and leaf rust. Of 502 genotypes,
only 110 were found to be susceptible to all three rust diseases. The
other 392 accessions showed resistance to one, two or all three of the
rust populations under investigation. For further screening and field
verification, the genotypes showing resistance to stem rust were
selected. Of the 101 accessions, 63 were also stripe rust resistant, 23
were both stripe rust and leaf rust resistant and 15 were stem rust
resistant only. They were planted at the Addis Ababa University's
experimental site at Debre Zeit in cooperation with the durum wheat
breeders. This site is known as a 'hot spot' for stem rust and has a
high disease pressure.
In order to allow mass screening of germplasm under abiotic stress
conditions (e.g. drought and aluminium toxicity tolerance) laboratory
screening methods are now being developed with encouraging
results so far. The installation of a near-infrared-reflectance analyser
allows easy and precise determination of chemical constituents, such
as protein, oil, water and fibre content, of seeds and other organic
parts.
Utilization activities
The ultimate goal of any germplasm evaluation is its utilization. The national plant breeders form the main target group and
PGRC/E tries to involve the breeders as much as possible in its routine
activities. Examples of close cooperation include the workshops held
with plant breeders and other potential users to discuss and agree on
priorities regarding germplasm collection and evaluation, the participation of breeders in the collection missions and the cooperation that
Table 3. Results of an evaluation of the sesame collection. The best 11 accessions are compared with the standard
1
2
3
Accession
number
Days
to 50%
flowering
Days
to podsetting
Days
to
maturity
111505
111518
111809
111815
111823
111824
111829
111833
111834
111859
111861
79
40
36
36
34
34
33
21
15
31
47
91
79
44
41
38
39
40
63
50
39
57
Standard T85
47
Experimental
average
47
8
5x7x8:1000
1000 seed
weight
(8)
Theoretical
yield"
(g)
70.2
75.3
69.1
84.1
57.6
64.4
70.9
68.8
77.6
79.4
62.0
2.28
3.91
3.47
3.71
3.70
3.87
4.27
3.73
3.00
3.31
2.97
23.8
17.1
43.9
20.4
17.1
23.4
49.0
27.9
17.8
31.4
20.7
22.6
67.8
3.52
21.0
22.1
64.2
3.23
15.5
6
4
5
Plant
height
(cm)
Number of
capsules
Capsule
per plant length
(mm)
Seeds
per
capsule
145
104
104
99
98
98
98
107
111
112
107
224
119
98
104
121
118
111
155
132
181
132
149.0
58.2
183.2
65.2
80.4
93.8
161.8
108.8
76.4
119.4
112.2
17.0
23.5
24.9
27.4
20.9
23.8
24.5
26.8
23.8
26.1
24.7
53
101
156
87.8
55
120
148
74.9
7
Calculated as a product of number of capsules per plant x number of seeds per capsule x thousand seed weight :1000.
266
Hailu Mekbib
Table 4. Number of accessions included in the Pre-national
(PNYT) and National Yield Trials (NYT) as well as in
observation trials (OT) in the period 1983-6
Crop type
Number of
accessions used
Type of trial
Faba bean
Field pea
Chickpea
Linum
Niger
Brassica
Sesame
Niger
Brassica
Linseed
Sesame
Groundnut
Sorghum
Barley
Niger
Lentil
Chickpea
Faba bean
Field pea
Castor bean
16
1
4
1
4
3
4
7
2
3
4
3
12
50
7
13
9
2
5
14
PNYT/NYT
NYT
PNYT/NYT
PNYT
PNYT
PNYT/NYT
PNYT
NYT
NYT
NYT
NYT
NYT
OT
OT
OT
OT
OT
OT
OT
OT
occurs during the characterization and multiplication of germplasm.
Furthermore, PGRC/E presents as many results as possible from
evaluation activities in the PGRC/E-ILCA Germplasm Newsletter or
during meetings with potential users. (An example of the characterization work on sesame is presented in Table 3.) This approach has
led to intensive utilization of germplasm by the breeders and some of
the results can be observed from Table 4. A more detailed account of
the procedures followed is given by Engels & Mekbib (1987).
References
Demarly, Y. (1981). Theoretical problems of seed regeneration in cross-pollinated plants. In: Seed regeneration in cross-pollinated species. Proceedings of the
CEC/Eucarpia Seminar, Nyborg, Denmark, 15-17 July 1981. Balkema, Rot-
terdam, pp. 7-31.
Engels, J. M. M. & Mekbib, H. (1987). The utilization of germplasm in Ethiopia and role of PGRC/E. PGRC/E-ILCA Germplasm Newsletter, 13, 21-5.
Frankel, O. H. (1970). Evaluation and utilization - introductory remarks. In:
Crop germplasm multiplication and characterization
267
O. H. Frankel and E. Bennett (eds), Genetic Resources in Plants, their Explora-
tion and Conservation. Blackwell, Oxford, pp. 395-^01.
Frankel, O. H. & Hawkes, J. G. (eds) (1975). Crop Genetic Resources for Today
and Tomorrow. Cambridge University Press, Cambridge.
Hawkes, J. G. (1985). Report on a consultancy mission to Ethiopia for GTZ to
advise PGRC/E on germplasm exploration, conservation, multiplication
and evaluation. Birmingham (mimeographed).
Seegeler, C. J. P. (1983). Oil Plants in Ethiopia, their Taxonomy and Agricultural
Significance. PUDOC, Wageningen, pp. 122-46.
Williams, J. T. (1984). A decade of crop genetic resources research. In:
J. H. W. Holden and J. T. Williams (eds), Crop Genetic Resources: Conserva-
tion and Evaluation. Allen and Unwin, London, pp. 1-16.
20
Evaluation methods and utilization
of germplasm of annual crop species
J.B. SMITHSON
Introduction
Proper evaluation of the very large germplasm collections
now assembled for many crop species presents major problems.
These arise principally from the effects of the environment on the
expression of plant characteristics. For qualitative characteristics there
is little difficulty as their expression is usually affected little by
environment. Examples are seed coat and flower colour and colour
pattern. A single evaluation is all that is required to characterize a set
of materials for such characteristics.
It is, however, the quantitative characteristics, and these are of
most interest to the breeder, that are especially intransigent as their
expressions are always modified by environment to some degree, so
that the separation of the contributions of genotype and environment
to the phenotype requires special techniques.
Environment and genotype X environment interaction
The modification of plant characteristics by environment
takes two forms. First, there is a general reduction or increase in
expression of a character across all genotypes. Environmental
features such as soil fertility, moisture availability, temperature and
pathogens, pests and weeds may all affect plant characters in this
way. The result is what is often termed 'field variability' and this will
always occur in a single evaluation at a single location in a single
season. It also occurs across locations and seasons.
Secondly, there is the situation where all genotypes are not affected equally by differences in environment, normally described as
'genotype x environment (g x e) interaction'. Such interaction is usu-
Evaluation methods and utilization of germplasm
269
ally associated with a set of materials being grown in more than one
location or season and, indeed, is only detectable in this way. It also
occurs in a set of materials at a single location and season but is in that
case not separable from general environmental effects.
The two features of the relationship between plants and their
environments pose serious problems in the evaluation of large numbers of materials, whether germplasm or breeding lines, and in the
interpretation of the collected data. Two examples from chickpeas
illustrate the importance of environmental effects in evaluation and
the misinterpretations that can arise as a result.
The first concerns seed protein percentages (International Crops
Research Institute for the Semi-Arid Tropics, unpublished data). Seed
protein percentages were routinely determined on seeds from successive evaluations of different germplasm accessions over a period of
five years. The seeds of 100 accessions representing the largest and
smallest seed protein percentages in each of the five years were
grown in a replicated trial at ICRISAT in 1982-3 and 1983^1 and the
seed protein percentages of their produce determined.
Examination of the data revealed very poor correlations between
the seed protein percentages from the germplasm evaluation and
those from the replicated trials in either of the two years. Large and
small values from the germplasm evaluation tended towards the
mean and many of the differences that had been demonstrated earlier
disappeared. There were, however, good correlations between the
protein percentages obtained from the trials in the two years, indicating that the discrepancies arose from environmental differences
within or between seasons or both but illustrating that repeatable
results can be obtained with appropriate methods.
The second example concerns the number of days to flowering
(ICRISAT, 1980). Based on germplasm evaluation data, five groups of
lines, flowering respectively in 45, 45-56, 57-68, 69-80 and more than
80 days after sowing, were included in five replicated trials, again at
ICRISAT, to examine their adaptability to sowing one month earlier
than normal. Mean flowering times for the trials were roughly according to expectation. The ranges of flowering times were, however, 3380, 34-66, 41-80, 40-106 and 44-124, overlapping completely, and
with very poor correlations with the earlier records, and this is exactly
what would be expected from the effects of different day lengths and
temperatures on the flowering times of sets of materials differing in
their photoperiod and temperature responses.
As usual, there is no complete solution. None the less, methods of
270
/. B. Smithson
measuring and controlling environmental variation are available and
some of these will be described and discussed in this chapter.
The questions of the need for evaluation of genetic resources and
of the characters to be evaluated are not considered here. It is
assumed that accessions are maintained discretely in order to retain
character combinations and that some kind of evaluation is required
for the purposes of utilization. In the text, materials being evaluated
will be termed test entries or materials. Performance refers to the
value assigned to any quantitative character, be it size, concentration
or number of any plant component, or rate of any plant process. It is
also assumed that performance is being measured in field nurseries
and with the necessary precision and accuracy. Disease and pest
reactions are not considered since they require specialized
techniques.
Field variability
First, we will consider means of handling a set of test entries
in a single location and season. Where numbers are relatively small
(say 500 or less), orthodox statistical designs such as randomized
blocks or lattices may be employed, utilizing techniques such as
randomization, replication and sub-grouping within replicates to
measure and/or control environmental effects.
But germplasm collections and early generation breeding materials
are usually too numerous to allow replication and, were replication
even feasible, and with the most uniform field environment conceivable, the replicates would be too large to cope with field variability by
orthodox analysis of variance. What then can be done to measure
and/or control environmental effects in such situations?
Regular check entries
The simplest means of obtaining a measure of environmental
variability in an unreplicated set of entries is the inclusion of check
entries. This is common practice in germplasm evaluation, comprising the inclusion of two or three different checks in the nursery at the
rate of one check for every 10 test entries. Unfortunately, the mere
inclusion of checks is not sufficient. They must also be used to assess
and reduce field variability by some form of adjustment of the performances of the test entries according to the performance of the
checks, and this is rarely practised.
The simplest form of adjustment is to express the performance of
each test entry as a percentage or proportion of some measure of the
Evaluation methods and utilization of germplasm
271
performance of the checks in the same sector of the field. Where
several different checks are included, the best measure is likely to be
the mean of the performances of the nearest full set or sets of checks.
Alternatively, the performance of each test line may be adjusted by
subtracting the deviation of the mean of the performances of the
nearest full set of checks from the mean of the performance of all the
checks. For example, if the mean number of seeds per pod of the
nearest full set of checks is 9.5 and the mean for all checks is 9.0 seeds
per pod, the number of seeds per pod of each test entry in the same
sector of the field is adjusted downwards by 9.5 — 9.0 = 0.5.
Conceptually, the latter adjustment is more desirable than the first
in that the actual and the adjusted data are of the same units. Both
have the disadvantage of providing no estimate of error for a comparison of the differences among the test lines.
Augmented designs
An extension of the regular check system is the augmented
design (Federer, 1956). The test materials are again unreplicated but
are randomized and grouped in blocks of convenient size (say 20-50)
for the number of materials, size of field and number of checks. An
appropriate number of different check entries (2-5 according to block
size) is then randomized within each block. Check performance can
then be analysed in the form of a complete randomized block and the
estimate of error so derived used to compare the performances of the
unreplicated test lines.
The performances of the test lines may also be adjusted according
to the deviation of the mean of the checks in the block in which they
occur from the mean of all the checks, so that S ; — y . . is the adjustment for the performance of each of the test lines in the jth block,
where Bj is the mean of the checks in the jth block and y.. is the
overall check mean. Note that the variance of the difference of two
test entries in the same block will be twice the error mean square,
while that of two test entries in different blocks will contain an additional quantity for block differences.
The method assumes that the random components associated with
the checks and the test lines are similar. This may not be so, but is
more likely if the checks are chosen to represent the range of variability in the collection. However, it does provide a measure of
environmental variation and a means of adjusting the performances
of the test lines to remove some of the variability, and thus is an
improvement on other methods.
272
/. B. Smithson
One further point should be mentioned. It is common practice to
include test lines in order of origin, or to group them in some other
way, so that similar materials are compared more accurately. But in
evaluating a set of materials we are also interested in the relative
performances of dissimilar materials, so in the absence of very compelling reasons for grouping, less biased comparisons are obtained by
randomizing the test lines.
Nearest neighbour analysis
A third and less often used method of handling field variability is adjustment according to the performances of neighbouring
plots, first proposed by Papadakis (1937) and described with a
worked example in Pearce (1983). Adjustment by neighbouring plots
may be applied to any replicated field layout and is especially useful
for large sets of materials. It adjusts the performance of each plot
according to the mean performance of its neighbours. In most cases,
the four plots adjoining the ends and sides of each plot are used for
the adjustment. Where plots are long and narrow, it is more
appropriate to use only the plots along each side. In the case of end
plots there are only three neighbours and corner plots have only two.
It has been found that the first cycle of adjustment is often erratic, but
it is possible to iterate (i.e. repeat the calculation with the adjusted
values until the adjustments remain similar) as is usually done when
estimating values for more than a single missing plot.
The analysis proceeds as follows:
- compute the deviations of each plot from the mean of all plots
of that treatment;
- compute the mean deviations (X) of the neighbours of each
plot;
- compute treatment totals and means for the X values;
- compute an analysis of covariance of the actual values (Y) on
X;
- compute the regression (b) of Y on X;
- adjust the Y value for each plot subtracting b(X - x);
- iterate the above steps until the adjustments are the same;
- compute the analysis of variance of the adjusted values.
Since the test entries must be replicated the area required will be
large but the method allows for adjustment for patchy field variability, which is not possible in an orthodox analysis of variance, in
addition to providing an estimate of error.
Evaluation methods and utilization of gertnplasm
273
Summary
It should be noted that these methods of analysis are not
mutually exclusive. Common checks ought always to be included to
enable adjustment across seasons and, provided there is replication
and randomization, both types of adjustment are theoretically possible. Augmented designs and nearest neighbour analysis both provide estimates of error for comparison of differences among entries;
nearest neighbour analysis takes up more land because at least two
replicates are required, but this may be accommodated to some extent
by reducing plot size. Computer facilities are desirable for all because
of the large volume of material to be examined. Nearest neighbour
analysis can be expected to produce more accurate results because of
replication and the opportunity to adjust for patchy field variation
and (see next section) to assess the magnitude of g X e interaction.
Evaluation across sites and seasons
Because of g x e interaction, the relative performance of a set
of materials in one environment is unlikely to reflect its relative performance in other locations and seasons. Therefore the value of a set
of data obtained in one environment is doubtful, especially so in a
country as varied as Ethiopia.
Furthermore, the very large numbers now present in germplasm
collections virtually preclude the possibility of evaluating all materials
at a single time. The whole available Centro Internacional de Agricultura Tropical (CIAT) bean collection (more than 17000) was evaluated for resistance to angular leaf spot in hill plots this year (CIAT,
1987) but this is a special case. There is also the case of continuing
collection and the need to evaluate newly assembled groups of
materials. For these reasons, the evaluation of germplasm collections
in successive seasons and/or at more than a single location is
inevitable.
Care should be taken to ensure that environments are as uniform
as possible. For example, in the evaluation of the International
Institute of Tropical Agriculture (IITA) cowpea collection in two successive years, sowing was on the same date and exactly the same
cultural practices were used, with sufficient fertilizer and irrigation to
reasonably eliminate any soil nutrient or moisture stress (IITA, 1974).
Nevertheless, it must be accepted that the removal of all environmental variability likely to affect performance is impossible. General
environmental effects can be accommodated to some degree by the
274
/. B. Smithson
inclusion of common checks and by using augmented designs to
adjust performance across seasons in the same manner as within
seasons.
G x e interactions cannot be handled in this way. Several methods
of assessing their magnitude and of characterizing them have been
developed. They include: combined analysis across locations and
seasons; regression of individual entry performance on environment
mean performance or some other environmental measure; and, more
recently, multivariate analysis. A comprehensive account of these
techniques is given by Hill (1975). Such methods have helped in the
understanding of g x e interaction but are not appropriate to the
evaluation of large numbers of materials. There appears, therefore, to
be no escape from the need to evaluate the same test materials in
more than a single environment. Seasons are unsatisfactory as they
are unpredictable, so this means testing at a number of locations.
Multivariate analysis can be used to choose locations that represent
the range of environments in which the test entries are likely to be
utilized. The greater the number of locations, the more complete will
be our characterization, but practically, three or four (say, two
extremes and two intermediates) may be all that is feasible or, for
some characters, even necessary. For example, based on growth
cabinet studies with chickpeas and lentils, Roberts, Hadley & Summerfield (1985) and Summerfield et al. (1985) concluded that evaluation in field nurseries at three properly chosen locations is sufficient
to characterize accessions of these species for their flowering responses to photoperiod and temperature. Alternatively, the number of
environments may be increased and the number of materials reduced
by selecting representatives of the total variability by some form of
multivariate analysis.
Finally, environmental data are every bit as important as plant
character data in any evaluation if we are to understand variation in
performance across environments. It is vital, therefore, to characterize the physical and biological environments in which evaluations
are conducted as thoroughly as we characterize our plants. Physical
factors should include, at least: latitude; altitude; maximum and
minimum temperatures and rainfall on 10-day mean bases during the
growing season; physical and chemical properties of soils. The biological environment will include diseases, insects and weeds.
Suggested procedures
Based on the above considerations it is possible to suggest
optimal procedures for germplasm evaluation.
Evaluation methods and utilization ofgermplasm
275
Whole collections should be evaluated at three or four locations
selected to represent a range of situations. Operationally, this will
have to proceed in groups of around 2000 entries. These should be
selected at random from those available for evaluation. They should
be deliberately grouped only if absolutely necessary, e.g. bush and
climbing types. They should be sown in an augmented design with a
set of frequent, common checks, chosen to cover the total variation in
the collection as far as possible. If possible, two replicates should be
sown at each location to allow a nearest neighbour analysis. Plant and
environment data should be collected on each evaluation.
Multivariate analysis of the data should be conducted to select sets
of different sizes representing the variability in the collection, as was
done with the IITA cowpea collection in the late 1970s (Rawal, Kaltenhauser & Snyder, 1977). These sets should be of different sizes (say,
100, 500, 1000 and 2000) to accommodate different capacities. The
evaluation of these sets by breeders in other environments and the
return of the data for continuous updating of information should be
vigorously encouraged.
Finally, a note of caution should be added. The techniques described are merely tools to aid evaluation. In most circumstances they
can be expected to be useful and we have a duty to use them. But
there is always the possibility that they may distort differences rather
than reduce environmental variation. There is, therefore, no substitute for knowledge of the crop, careful observation of the test
materials in the field, careful examination of the actual and adjusted
data and the application of common sense in their interpretation.
Dissemination of information
A further area requiring thought is the method of presentation of data. Pulse germplasm catalogues include those for cowpea
(IITA, 1974), beans (CIAT, 1983) and chickpea (Singh, Malhotra &
Witcombe, 1983). All comprise long lists of accessions and characters
recorded. For cowpea, there are 46 characters for 4224 accessions; for
chickpea, there are 29 characters for about 3300 accessions.
Such forms of presentation are very difficult for an aspiring breeder
to assimilate and are therefore not very useful. The information is
important for the institutions conducting the evaluation but can be on
computer file for manipulation as it is unlikely that a hard copy of the
complete information is ever going to be required.
For the breeder, it would be more useful to have a summary for
each character, perhaps in the form of a histogram showing its
frequency distribution together with an estimate of the total variation,
276
/. B. Smithson
such as the coefficient of variation. This should be accompanied by
important characters (e.g. disease resistance) or important combinations of characters. Data on the environments in which the evaluations are carried out and a summary of the check performance should
also be included. The intending breeder can then see easily what kind
of variation is available and can request whatever number of
materials, having the range of characteristics and adaptation in which
he is interested, that he is capable of handling.
These procedures presuppose that the intending breeder knows
what he requires. But in many cases this may not be so. In other
situations, materials adapted to a wide range of environments are
needed. These are additional reasons why performance in different
environments is important and environmental information is
required for every evalution.
In this situation, the different sized sets representing the variation
are important. The breeder evaluates a set of the size he can handle in
his own environment, identifies the most promising materials,
requests additional accessions of similar origin and character from the
distributing institution and returns the data for updating the information base.
The development of these kinds of relationships between genetic
resource specialists and breeders is vital if we are to make the most
effective use of our genetic resources.
References
Centro Internacional de Agricultura Tropical (1983). Catalogo de Frijoles.
CIAT, Call.
Centro Internacional de Agricultura Tropical (1987). Annual Report 1986.
CIAT, Cali.
Federer, W. T. (1956). Augmented (or Hoonuiaku) designs. Hawaiian Planters
Record, 55, 191-207.
Hill, J. (1975). Genotype x environment interactions - a challenge for plant
breeding. Journal of Agricultural Science, Cambridge, 85, 477-93.
International Crops Research Institute for the Semi-Arid Tropics (1980).
Annual Report 1979. ICRISAT, Hyderabad.
International Institute for Tropical Agriculture (1974). Cowpea Germplasm
Catalogue no. 1. IITA, Ibadan, Nigeria.
Papadakis, J. (1937). Methode statistique pour des experiences sur champ.
Bulletin de VInstitute d'Amelioration des Plantes, Salonika 23.
Pearce, S. C. (1983). The Agricultural Field Experiment. Wiley, Chichester.
Rawal, K. M , Kaltenhauser, J. & Snyder, M.J. (1977). Agronomy abstracts.
69th Annual Meeting, American Society of Agronomy 68.
Roberts, E. H., Hadley, P. & Summerfield, R. J. (1985). Effects of temperature
and photoperiod on flowering in chickpeas (Cicer arietinum L.). Annals of
Botany, 55, 881-92.
Evaluation methods and utilization of germplasm
277
Singh, K. B., Malhotra, R. S. & Witcombe, J. (1983). Kabuli Chickpea Germplasm
Catalogue. ICARDA, Aleppo.
Summerfield, R. J., Roberts, E. H., Erskine, W. & Ellis, R. H. (1985). Effects of
temperature and photoperiod on flowering in lentils (Lens culinaris
Medic). Annals of Botany, 56, 659-71.
21
Evaluation and utilization of
Ethiopian forage species
J. R. LAZIER AND ALEMAYEHU MENGISTU
Introduction
Plants which are utilized as fodder for livestock are confined
to those which can provide maximum yields in animal production
with minimum management inputs. Such plants are usually members of the families Gramineae and Leguminosae and are mainly
herbs or subshrubs. In recent years leguminous shrub and tree species have been receiving increasing attention, particularly for small
farmers in developing countries.
The Gramineae and Leguminosae are major sources of human
nutrition as cereals and pulses while the by-products of these crops
are major sources of nutrition for livestock. This report will be confined, however, to plants which are planted primarily as livestock
fodder.
Ethiopia, as part of the African continent, shares many genera and
species of grasses and legumes with the rest of the continent.
However, its great variations in climate and relief and its heavily
dissected landscape have provided the opportunity for further evolution of species and genotypes. About 64 species of legumes, mainly
montane (10-11 per cent of the total), have been reported as probably
being endemic to Ethiopia, while 30 species of grasses are endemic
(Thulin, 1983).
African grasses are the main source of cultivated commercial grass
species and cultivars in the tropics and subtropics worldwide and are
almost all represented in Ethiopia. While Africa is not the major
centre of diversity in legumes it is a major centre of diversity for such
genera of fodder potential as Aeschynomene, Alysicarpus, Indigofera,
Lablab, Lotononis, Macrotyloma, Neonotonia, Trifolium and Vigna. Other
Evaluation and utilization of Ethiopian forage species
279
genera of importance which are represented include Cajanus, Clitoria,
Galactia, Stylosanthes and Zornia. Browse species of potential in Ethiopia include Acacia, Albizia, Bauhinia, Cassia, Dichorostachys, Eriosema,
Erythrina, Flemingia, Lonchocarpus, Millettia, Parkinsonia, Piliostigma,
Sesbania and Tamarindus. Considerable diversity is present in Acacia
and Sesbania. Thirteen of 24 important legume genera are represented
and 14 of 26 browse genera (Skerman, 1977). Undoubtedly potential
exists in other leguminous genera as little work has yet been done on
collection and evaluation.
Early work
Early work in the screening of native forage germplasm was
done by a Food and Agriculture Organization (FAO) project which
included a number of local collections in unreplicated adaptation
plots at Adami Tulu and Abernosa Ranch in the Rift Valley. The local
germplasm, which was not named, was less successful than exotic
germplasm (Ibrahim, 1975).
The Chilalo Agricultural Development Unit (CADU) has screened
a number of native lines of commercialized grass species (Carlsson,
1972; Froman, 1975). Eight Cenchrus ciliaris ecotypes were found to be
more productive than a Kenyan variety in the lowlands of Chilalo but
were not sufficiently productive to be of interest. Sixteen Chloris
gayana lines were screened and found to be very variable but to form a
good source of germplasm for development as both creeping, stoloniferous and upright, tufted, many-headed types were identified. Six
similar accessions of Phalaris arundinacea, which were collected 2200 m
above sea level in wet areas, were found to be more productive than
exotic lines. Germplasm of Hyparrhenia species and Panicum maximum
was also screened. H. hirta was less stemmy than other Hyparrhenia
species though all reportedly had low palatability. Considerable variation was found in 13 ecotypes of P. maximum germplasm which were
collected from 1500 to 1700 m; four lines were regarded as promising.
Indigenous lines of Setaria sphacelata were found to be better adapted
to higher altitudes than cv. Nandi. Local lines were also more
rhizomatous.
An investigation of the leguminous species of the Arsi region by
the same project resulted in the identification of some 90 taxa of
which about 20 were considered promising for fodder development.
Indigenous material of Trifolium semipilosum var. semipilosum was
more vigorous than the cultivar Safari and one line of Neonotonia
wightii was earlier flowering, heavier seeding and perhaps slightly
lower yielding than commercial lines.
280
/. R. Lazier & Alemayehu Mengistu
Recent work
Over the past five years the International Livestock Centre for
Africa (ILCA) has been involved in the evaluation of native Ethiopian
germplasm as part of the normal procedures of the ILCA Genetic
Resources Section. Agronomic description and evaluation are done
for all lines in the collection at the same time as the germplasm is
multiplied. Multilocation evaluation is then done by the Forage Network in Ethiopia (FNE) in a range of environments. In addition,
national and ILCA scientists have been pursuing their own evaluation programmes on ILCA germplasm. This report will consider the
screening procedures used and the results obtained in the regular
screening exercises undertaken by ILCA and FNE.
Highland species
Annual Trifolium
The initial description, agronomic evaluation and seed
multiplication of the annual native Trifolium species is done at ILCA
headquarters in Addis Ababa (9°02'N, 38°42'E) at an altitude of
2400 m on a transitional soil lying between nitosols and vertisols
which is somewhat more free-draining than the vertisols. Unreplicated strips, 5 m long, at a spacing of 1 m, are planted with a single
line of scarified seed sown at 0.6 g per plot. As the strips are unreplicated, control strips are planted in each fifth plot. The soil is slightly
mounded along each strip and abundant small drains are provided in
the trial to prevent water transporting or burying the seed. Fertilizer
is applied at 10 kg P/ha, banded below the planting level beside the
line of planting. The amount applied is calculated on a 0.5 x 5m plot
basis. The plots are weeded while the pathways are allowed to
accumulate weeds, which are mown periodically.
Regular three-weekly observations are taken on general characteristics, with more frequent observations on germination and date of
flowering. All parameters observed are defined. The most promising
lines for planting in yield trials are determined by using grouping
techniques on the observational data. Principal component analysis
has been found to be the most useful technique.
Screening has been undertaken in this manner since 1983 and more
than 500 lines have been screened to date. The strips normally provide sufficient seed for long-term and duplicate storage, as well as for
the second stage evaluation.
The second level of screening (stage 2) is done in small replicated
plots (5 x 2 m) in which selected lines of the annual species are plan-
Evaluation and utilization of Ethiopian forage species
281
Table 1. Agronomic characteristics of selected lines of promising Ethiopian
annual Trifolium species. 1984 replicated yield trial
Species
Number
of lines
Range of
DM yields
(kg/ha)
Range of
seed yields
(kg/ha)
Trifolium decorum
T. quartinianum
T. rueppellianum
T. steudeneri
T. tembense
5
4
5
5
5
4200-7500
5500-7700
3700-5300
4400-7000
5000-6100
860-1590
150-560
1100-1480
71fl-790
lOMO"
Days to 50%
flowering
>122fl
80-102
72-87
67-93
85-123
a
Dried before maturity due to cessation of rain.
Source: Kahurananga & Tsehay (n.d.).
ted in pure stand. Seed and hay yields are recorded. The most recent
form of this trial includes a plot which is left unplanted and a plot
planted to oats. These plots provide controls in the second year of the
trial when oats are planted in all plots. N fertilizer is then applied to
half of each plot at 40 kg/ha. The trials are run for two or three years to
determine the long-term effects of N fixation by the Trifolium lines
planted in the first year.
In these replicated trials hay yields have generally been good, with
yields varying greatly within species. Generally T. quartinianum, T.
decorum and T. steudeneri have provided the best yields, with the most
productive lines in years of good rainfall producing 7 t/ha or more of
dry matter (DM) and up to 1.5 t/ha of seed (Table 1).
Other studies have indicated that a T. tembense content of 20-25 per
cent in teff straw increased intake of the mixture by 20-30 per cent
compared with the pure teff straw control (Butterworth, Mosi &
Preston, 1985). Similarly, the addition of T. tembense to a number of
cereal straws increased the apparent digestibility of the DM, crude
protein and phosphorus of the diets (Mosi & Butterworth, 1985).
Since 1985 FNE has planted multilocation initial evaluation trials of
the elite lines from stage 2 screening. The trials are planted as unreplicated strips of 20 genotypes belonging to five species (T. quartinianum,
T. decorum, T. rueppellianum, T. steudeneri and T. tembense) with T.
quartinianum ILCA 6301 as control in each fifth plot and about the trial
perimeter. The 7.5 m strips have three P treatments imposed: 0, 10
and 20 kg P/ha. The trials have been planted by ILCA, the Institute for
Agricultural Research (IAR) and the Arsi Rural Development Unit at
282
/. R. Lazier & Alemayehu Mengistu
Addis Ababa, Debre Berhan (Enwari), Holetta, Kulumsa and Gobe in
a range of soil types and altitudes. Standard observations are made
(Anonymous, 1985). ILCA has also recently started screening of
native germplasm at Agew Midir, in Gojam, on a nitosol.
Screening is currently under way in other countries using ILCAsupplied Ethiopian germplasm. These include six other African
countries, five in Central/South America, three each in Europe and
Asia, as well as Australia, New Zealand, Canada and the USA.
Stage 2 multilocation screening of native annual highland Trifolium
species has been under way since 1985 (Anonymous, 1985). Three
promising lines of native annual Trifolium species (T. quartinianum
ILCA 6301, T. tembense ILCA 5774 and T.rueppellianum ILCA 9690) are
planted in mixture with the two most productive grasses from earlier
FNE multilocation trials, Festuca arundinacea cv. Demeter and Phalaris
aquatica cv. Sirocco. These trials are being undertaken at sites at or
above 2400 m by ILCA, the Arsi Rural Development Unit and IAR at
Holetta, Addis Ababa, Gobe and Debre Berhan (Enwari).
Perennial Trifolium
The difficulty of maintaining pure lines of the outcrossing
perennial highland Trifolium species has inhibited seed multiplication
and the amount of screening done.
In 1985, at ILCA headquarters in Addis Ababa, 98 accessions of
perennial Trifolium species (T. burchellianum 26, T. cryptopodium 13 and
T. semipilosum 59), mainly from Ethiopia, were planted as single
plants transplanted from pots to the field in unreplicated rows with
10 plants per row at spacings of 1 X 1 m. Fertilizer was applied once at
40 kg/ha.
Standard parameters are observed every three weeks as well as
such morphological parameters as petiole length, middle leaf length
and width, stolon width, internode length and time of flowering.
Harvests are taken to obtain dry matter yields.
Vicia spp.
Initial screening of 100 accessions of exotic Vicia saliva germplasm and eight collections of native Vicia species was done at ILCA
headquarters in 1984 using the same procedures as previously described for the annual highland Trifolium species. The native lines had
comparable performance to the introduced lines, with two lines
(ILCA 8324, 8047) being grouped among the most promising.
Evaluation and utilization of Ethiopian forage species
Tropical and subtropical
Acid soils, rainfed
283
screening
Stage 1 screening of mainly perennial species of tropical and
subtropical exotic and native germplasm on acid soils in the Sodo area
(1850 m, 1100 mm rainfall) has been under way for more than two
years and the initial 1984 plantings have thus completed their third
growing season.
Screening in Sodo is done using unreplicated microplots
(2 x 1.5 m), each with six plants spaced at 0.5 m and established by
seed or transplanting from pots. The plots are slightly mounded for
uniform drainage and natural vegetation is allowed to establish in the
plots and pathways to provide a more natural environment for the
growth of the plants. The weeds are cut back from time to time and
regular observations are taken at six-week intervals on defined morphological and agronomic parameters on standard ILCA observation
sheets. P fertilizer is added at 10 kg/ha at planting and at the same rate
annually as a split application after alternate observations.
The lines planted in 1984 for initial screening were mainly single
representatives of a wide range of leguminous species. As there had
been little collection of native lowland species at that time, few
introductions were of native germplasm. An early assessment of the
results (Lazier, 1986) indicated that of the six species rated as excellent, three are native to Ethiopia: Argyrolobium ramosissimum,
Macrotyloma axillare and Eriosema psoraleoides. Of these lines only E.
psoraleoides was collected in Ethiopia. Included in the 11 lines ranked
as good were one line each of Neonotonia wightii, Indigofera arrecta and
a Crotalaria species, all of Ethiopian origin.
The 1985 initial evaluation microplot plantings included more than
1000 lines. Major plantings were made of lines of Stylosanthes fruticosa
and Neonotonia wightii, both of which are important, widely represented species in Ethiopia. Of these, the lines of S. fruticosa being
screened are all native. The germplasm is relatively similar in
appearance and thus analyses are being done on details of morphological and agronomic characters in an effort to group accessions
which are similar.
Thirty-seven Neonotonia wightii lines were screened (36 native and
one exotic). Five native lines were outstanding in adaptation and
performed better than an earlier planting of a commercial line (cv.
Cooper). In limited plantings of grass germplasm, mainly commercial, a local collection of Melinis minutiflora performed as well as a
284
/. R. Lazier & Alemayehu Mengistu
commercial line. Ten lines of native Zornia species are being screened
along with 161 lines of mainly South American origin in a 1985 planting. Only one line of the native Zornias is recorded as having good
vigour.
Basic soils, irrigated
All tropical and subtropical germplasm of grasses, herbaceous legumes and browse which can grow reasonably well on basic
soils and at medium altitude is multiplied at the 4 hectare ILCA seed
multiplication site on the Ministry of State Farms, Horticulture Division's site at Zwai (1650m, pH8.0). Here, since 1983, germplasm has
been planted and replanted as required to provide seed for the
genebank and early stage agronomic evaluations.
Plantings are done in 5 X 5 m plots either by seed or by transplanting pot-raised plants. The size of the plantings varies, from one plant
to a 5 x 5 m plot, depending on the amount of seed available. P
fertilizer is applied at 50 kg/ha to all plots and N at 20 kg/ha to grass
and non-legume plots. The applications are made after alternate sixweekly observations. Observations are made of defined characters on
standard ILCA seed multiplication observation forms.
Ideally, all germplasm of one species would be planted at the same
time to provide more uniform screening conditions. However, such
organized plantings are unlikely to be possible until the large backlog
of germplasm awaiting urgent multiplication is reduced. Despite the
variations in time of planting, considerable useful data are acquired
from the plots as the moisture availability is uniform year-round, due
to irrigation, and the plants are normally in the plots for more than
one year.
The number of native Ethiopian grass and legume lines which have
been and are being multiplied in this manner is very large, approaching 3000 lines, and a huge amount of observational data has been
acquired. These data are currently being computerized for analysis.
Mainly vegetative collections of native materials of Erythrina have
been made and these have been screened under irrigation at Zwai.
Six accessions, which are leafy, relatively thornless, vigorous and
with a more prostrate growth habit, have been selected as elite lines
and are currently being further screened at Sodo and at ILCA headquarters at Addis Ababa. The species involved have not yet been
definitely identified as the plants have yet to flower. E. abyssinica and
E. brucei appear to be the species utilized as forage locally.
Evaluation and utilization of Ethiopian forage species
285
Feeding trials
Leguminous trees are important in the local farming system
as sources of forage and as important agents in maintaining soil
fertility and stability (e.g. Lazier, 1985). Their feeding value is currently being studied by the ILCA Nutrition Unit. Acacia seyal has been
found to be as good a feed for sheep as Sesbania sesban, once the
animals have become accustomed to it. The animals lost weight on A.
cyanophylla, an exotic species commonly planted in reafforestation
projects. Goats appeared to be better adapted to the digestion of
Acacia albida pods than sheep (Anonymous, 1986).
Current utilization
While no commercialized or large-scale plantings of native
forage germplasm have been carried out in Ethiopia, considerable use
is made of native forage germplasm. The valley bottoms of the highlands are commonly used as reserves or common grazing lands, or as
sources of hay. Native Trifolium species are frequently abundant in
these areas and undoubtedly make a considerable contribution to the
diet of livestock. A farming system has been discovered in the Gojam
administrative region in which dense stands of native Trifolium
decorum appear in the fallow year after a crop of teff. Teff and the
native legume are thus grown in alternate years as the farmers recognize the feed value of the legume and its beneficial effect on the crop
the following year (J. Kahurananga, personal communication).
In the Welayita awraja of Sidamo administrative region above
1600 m, human population densities are high with farm sizes averaging about 0.5 ha. Here, native Stylosanthes fruticosa stands can be a
major component of the vegetation of eroded areas, sometimes being
the dominant component, while Neonotonia wightii, Zornia spp. and
Indigofera spp. also commonly occur. It has been reported from this
region (Anonymous, 1983; Ochang, 1985) that areas of such legumes
are not grazed but are hand-picked to feed to productive animals,
particularly milking cows. The legumes are commonly cooked before
feeding.
Erythrina species are commonly planted as browse hedges in the
middle altitude (1400-2000 m) areas of western Ethiopia on acid
nitosol soils. Such hedges have been reported as high as 2800 m at
Chencha in Gamo Gofa administrative region. Decreasing rainfall and
lower human population densities limit the distribution at lower
elevations. The leaves are normally fed at the beginning of the dry
286
/. R. Lazier & Alemayehu Mengistu
season. Reports on palatability vary markedly from area to area;
however, animals appear not to graze it in the wet season, suggesting
either that hunger drives the animals to graze it, or that its palatability
improves in the dry season. One farmer claimed that the cuttings
used locally were of a selection which was dwarf, leafy and sterile
(Lazier & Mengistu, 1984).
Sesbania sesban, a palatable browse plant of considerable potential
which is native to Ethiopia, is currently being planted widely in Ethiopia by the Soil and Water Conservation Department of the Ministry of
Agriculture. Exotic and native germplasm is being used. It is also
commonly used by farmers to shade young coffee plants. The origin
of this material is not known.
ILCA's collections of native Sesbania germplasm show considerable
potential in initial evaluation plots. S. sesban (an exotic line) has done
well in the ILCA Highland Programme trials in stabilizing terraces
and as a green manure for cereal crops (Anonymous, 1986). S. sesban
germplasm is despatched regularly by ILCA, as part of a basic initial
evaluation package, to researchers in the tropics and subtropics
worldwide.
References
Anonymous (1983). Natural Stylosanthes pastures in Ethiopia. PGRC/E-ILCA
Germplasm Newsletter, 3, 10-11.
Anonymous (1985). FNE [Forage Network in Ethiopia] trial protocols 1985.
FNE Newsletter, 8, 8-15.
Anonymous (1986). ILCA Annual Report 1985/6. ILCA, Addis Ababa.
Butterworth, M.H., Mosi, A. & Preston, T. R. (1985). Molasses/urea and
legume hay as supplements to poor quality roughage in Ethiopia. XIII
International Congress of Nutrition, Brighton (abstract).
Carlsson, J. (1972). Inventory of indigenous ecotypes of some grass species in
Chilalo Awraja, Ethiopia. Asela (mimeographed).
Froman, B. (1975). Pasture management in Ethiopia with special reference to
conditions in the Chilalo Awraja. Agricultural College of Sweden. Report
and dissertation 32.
Ibrahim, K. M. (1975). Pasture and forage crops research programme report.
Project ETH/74/002/a/01/12. FAO, Rome (mimeographed).
Kahurananga, J. & Tsehay, A. (n.d.). Interspecific and intraspecific dry matter and seed yield and flowering variation of five annual Ethiopian Trifolium species (mimeographed).
Lazier, J. R. (1985). Acacia, an important natural forage plant in Ethiopia. FNE
Newsletter, 10, 20-2.
Lazier, J. R. (1986). Forage germplasm introduction in Sodo, Welayita: 1984
plantings. FNE Newsletter, 12, 17-22.
Lazier, J. R. & Mengistu, S. (1984). Erythrina, a genus with browse potential.
PGRC/E-ILCA Germplasm Newsletter, 8, 20-2.
Evaluation and utilization of Ethiopian forage species
287
Mosi, A.K. & Butterworth, M. H. (1985). The voluntary intake and
digestibility of diets containing different proportions of teff (Eragrostis tef)
straw and Trifolium tembense hay when fed to sheep. Tropical Animal Production (in press).
Ochang, J. (1985). Traditional use of native legumes for supplementary feeding of milking cows in the Welayita region of Ethiopia. PGRC/E-ILCA
Germplasm Newsletter, 8, 38-9.
Skerman, P.J. (1977). Tropical forage legumes. FAO Plant Production and
Protection Series No. 2, Rome.
Thulin, M. (1983). Leguminoseae of Ethiopia. Opera Botanica, 68.
22
Improvement of indigenous durum
wheat landraces in Ethiopia
TESFAYE TESEMMA
Importance of wheat in Ethiopia
Wheat has been and continues to be one of the most important cereal crops in Ethiopia in terms of both area under cultivation
and production. In 1983, the area under wheat production was
estimated at 625 590 ha with an average production of 1065 kg/ha
(Central Statistics Office, 1984). The demand for wheat as a staple
food grain is increasing, especially in the urban areas, while its
utilization will be high even in the rural sector in the near future. At
present, consumer demand for wheat as a staple food grain is increasing, especially in the urban areas, while its utilization will be high
even in the rural sector in the near future. At present, consumer
demand far exceeds domestic production and wheat imports are costing the country millions of dollars in foreign exchange. Wheat constitutes a large portion of the daily diet of the population and
contributes significantly to the calorie and protein intake. It is consumed in several different forms such as leavened bread, pancakes,
macaroni and spaghetti, biscuits and pastries. The most common of
the Ethiopian recipes are dabo (Ethiopian home-made bread), hambasha (home-made bread from northern Ethiopia), kitta (unleavened
bread), injera (thin bread, part of the national dish and prepared
mainly from teff), nifro (boiled whole grains, sometimes mixed with
pulses), kolo (roasted whole grains), dabo-kollo (ground and
seasoned dough, shaped and deep fried) and kinche (crushed
kernels, cooked with milk or water and mixed with spiced butter).
All wheat in Ethiopia is produced under rainfed conditions and the
important production areas are Arsi, Bale, Shewa, Gojam, Gondar,
Eritrea, Tigray, Welo and the highlands of Harerge and Sidamo
Improvement of durum wheat landraces in Ethiopia
289
administrative regions. Durum wheat (Triticum durum) is by far the
most predominant species and occupies 60-70 per cent of the total
area under cultivation, while bread wheat (T. aestivum) constitutes the
remaining 30-40 per cent (Tesemma & Mohammed, 1982).
Durum wheat is a traditional crop in Ethiopia. It is traditionally
grown on the heavy black clay soils (vertisols) of the highlands at
altitudes between 1800 and 2800 m above sea level. On the other
hand, bread wheat, although of recent introduction, has a wider
environmental adaptation than durum wheat. At present, however,
it is grown primarily in Arsi and Bale administrative regions.
Genetic diversity in Ethiopian wheats
Nearly all the wheat varieties grown at present are landraces
consisting of a large number of different genetic lines; even different
species are often found grown as mixtures in wheat fields. It is not
uncommon to find three or occasionally four species and often as
many as 8-15 botanical varieties in the same field.
The variation in the Ethiopian wheats is so great that N. I. Vavilov
was misled and classified Ethiopia as one of the centres of wheat
origin, before other genetic resources authorities (Harlan, 1971)
located the origin in the Near East and confirmed Ethiopia as a centre
of diversity because none of the wild relatives had been found in the
country.
According to Porceddu, Perrino & Olita (1973), Ethiopian wheats
belong to the following species: Triticum turgidum, T. durum, T. dicoccum, T. aestivum, T. polonicum, T. pyramidale and T. abyssinicum. Of the
tetraploid wheat species, durum wheat (T. durum) is the most extensively cultivated. It is believed that the bulk of the wheat types grown
in Ethiopia have undergone very little change over the past centuries
(Nastasi, 1964).
Characteristics of the Ethiopian tetraploid wheats
In general, the Ethiopian durums differ in spike form, spike
density, spike, awn and kernel colours (brown, amber and violet) and
awn condition (awnless, short-awned and long-awned varieties).
Other distinguishing characteristics include hairy or waxy leaves,
hairy or glabrous glumes and presence of pigmentation in glumes,
awns and kernels of seedlings and mature plants (Tables 1 and 2).
The author has observed that the kernel structure and pigmentation of the glumes and awns are influenced by altitude. The same
variety when grown at different elevations may produce seeds that
290
Tesfaye Tesemma
Table 1. Phenotypic diversity in Ethiopian drum wheat germplasm
accessions for some quantitative characters, expressed in their range, mean
and coefficient of variation (N=1120)
Character
Number of kernels per spike
Number of spikelets per spike
Days to 50% flowering
Days to 75% maturity
Plant height (cm)
Range
Minimum
Maximum
11
2.6
49
86
15
63
40
109
151
135
Mean
CV
(%)
30.8
18.5
74.3
108.1
93.2
22.8
16.2
13.1
11.4
15.0
Source: PGRC/E (unpublished data).
look different from each other. At higher elevations (e.g. Cheffe
Donsa at 2450 m) the pigmentation of the glumes and awns is more
intense and kernels are vitreous. At lower elevations (e.g. around
Debre Zeit at 1900 m) the pigmentation may be completely absent on
glumes and awns, and the kernels may be affected by yellow-berry.
At Akaki (2200 m) different degrees of pigmentation and a mixture of
kernels with yellow-berry and amber colours can be noticed.
Ethiopian wheats are important because of their rust resistance,
long coleoptile, short culm, low tillering, early ripening and drought
resistance (Porceddu et ah, 1973). However, from the author's personal observations, it can be said that Ethiopian durum wheat
cultivars generally have very location-specific adaptabilities. They
also have weak straw which is prone to lodging. As a result, they do
not respond very well to fertilizer application and hence are adapted
to low soil fertility conditions. They lack satisfactory resistance to leaf
rust but appear to be highly resistant to stem rust. They have the
ability to survive deep seeding, better than exotic varieties, and tend
to be rather low-yielding but more reliable for yield under adverse
conditions.
Diseases and insect pests of wheats in Ethiopia
Wheats in Ethiopia are attacked by a number of fungal
diseases, such as leaf rust (Puccinia recondita), stem rust (P. graminis
tritici) and stripe rust (P. glumarum). Leaf blotch (Septoria tritici) and
glume blotch (S. nodorum) cause considerable damage while bunt or
stinking smut (Tilletia foetida or T. caries), Helminthosporium spp.,
Fusarium spp. and powdery mildew (Erysiphe graminis) are problems
Table 2. Phenotype diversity in Ethiopian durum wheat germplasm accessions for some qualitative characters,
expressed as frequency distribution over the respective classes (N=1123-1173)
Character states and the frequency distribution (in %)
Character
Awnedness
Glume hairiness
Glume colour
Spike density
Kernel colour
Kernel size
Kernel vitreousness
Awned
99.0%
Absent
87.4%
White to yellow
84.5%
Very lax
1.5%
White to yellow
47.4%
<6mm
32.3%
Soft and starchy
4.5%
Source: PGRC/E (unpublished data).
Awnless
0.8%
Low
3.6%
Red to brown
14.5%
Lax
9.5%
Red to brown
26.2%
6-8 mm
64.8%
Partly vitreous
3.8%
Mixture
0.2%
High
9.0%
Purple to black
1.0%
Intermediate
64.3%
Purple to black
26.4%
>8mm
2.9%
Vitreous
91.7%
Dense
24.6%
Very dense
0.1%
292
Tesfaye Tesemma
in some areas. Recently bacterial stripe (Xanthomonas translucens) has
also been reported as causing some damage.
The extent of damage caused by these diseases depends on the
weather conditions during the crop season. However, because the
cultivars grown are landrace varieties consisting of genotypes which
differ in their reaction to diseases and pests, some lines may be
resistant or tolerant to certain races of the pathogen and others to
different races. Consequently, wheat diseases rarely reach epidemic
proportions because of the mixture of resistant and susceptible
genotypes in the population which provides a buffer against rapid
disease development and helps to extend the life of the resistance
genes (Harlan, 1975).
Among the insects that commonly attack wheat is the wheat aphid
(Rhopalosiphum maydis) which sucks on the leaves. Occasionally, ladybird beetle larvae (Chnootriba similis) feed on the leaves, leaving only
the fibre, while stem borer (Sesamia epunotifera) kills the plants after
heading in scattered spots in wheat fields.
Status of indigenous wheat landrace improvement in
Ethiopia
Attempts to improve Ethiopian indigenous wheat landraces
started in 1949 at the Paradiso Experimental Station near Asmara
(Tesemma & Mohammed, 1982). Among several local wheat collections tested for productivity, stem rust and leaf rust resistance, four
selections, namely A10, R18, P20 and H23, were selected and
released to farmers in Eritrea in 1952.
In 1956 and 1957, several crosses were made between local and
exotic varieties mainly for the purpose of transferring stem rust
resistance of A10 and R18 to Mindum, a variety of excellent quality
from the USA. Two outstanding lines, Mindum x A10 and
Mindum x R18 were selected but were later found to be susceptible
to leaf rust. An attempt to incorporate leaf rust resistance into the
above two lines from an exotic variety (Senator Capelli) resulted in a
selection with good resistance to stem and leaf rusts but too late in
maturity for the Eritrean environment (Nastasi, 1964). For some
unknown reason, work at Paradiso has been discontinued.
Although the Debre Zeit Agricultural Experimental Station was
established in 1953, emphasis in wheat improvement has, until very
recently, been on exotic varieties. Work on indigenous landraces consisted mainly of making collections and maintaining the germplasm.
Subsequently, all of the then 1145 wheat accessions at hand were
Improvement of durum wheat landraces in Ethiopia
293
transferred to the Plant Genetic Resources Centre/Ethiopia (PGRC/E)
soon after its creation in 1976. Through mass selection, two local
varieties, DZ04-118 and Marou (DZ-688), were released to farmers in
Yerer-Kereyu Awraja in 1968 (Tesemma & Mohammed, 1982). On
the other hand, as a result of introduction and extensive testing, four
high-yielding and adapted exotic varieties of durum wheat (Cocorit
71, Gerado, Ld57/CI8155 and Boohai), have been released to farmers
by the experimental station. These varieties are mainly grown in
Yerer and Kereyu and Menagesha awrajas where the extension programme of the experimental station is strong. Although these
varieties are higher yielders than the local cultivars, they are equally
susceptible to diseases, except for the variety Boohai which has good
resistance to leaf and stem rust.
Work on durum wheat hybridization at Debre Zeit was initiated in
1974. Since then, numerous crosses have been made within local
parents and local-exotic parents with a view mainly to improve yielding capacity, disease resistance and straw strength. So far no superior
line has been identified. Nevertheless, the programme continues.
Consequently, there are at present quite a large number of genotypes
at an advanced stage of yield testing.
Prospects
Genetic erosion is not yet far advanced, except in Arsi
administrative region where the adapted local landraces have been
virtually replaced by exotic bread wheat varieties. The administrative
region of Bale is the next most vulnerable area for genetic erosion
because of the rapid expansion of bread wheat production in the state
farms. In general, genetic erosion is bound to accelerate with the
release of high-yielding varieties from research stations, improvement of the infrastructure, increase in population and an overall
advance in agriculture. Cognizant of this, PGRC/E has identified
wheat as a priority crop for collection and preservation. According to
Worede (1985) about 7400 wheat samples have been collected and are
preserved at PGRC/E for future use. Nevertheless, to be properly and
effectively utilized by the breeders, these and other samples must be
characterized, screened and evaluated and the information obtained
must be properly documented and made easily available, together
with the seed samples.
Ethiopia has a wide range of climatic and ecological conditions.
The indigenous cultivars are the outcome of the evolutionary processes in response to the heterogeneous environment that prevails in the
294
Tesfaye Tesemma
wheat growing areas of the country. They are staunch survivors in
the struggle for 'survival of the fittest' in the vagaries of nature where
intermittent 'normal' seasons have forced them to undergo natural
selection for centuries (Chavan, 1975). Consequently, the genetic
variability in Ethiopian wheats has tremendous potential which, if
properly exploited, could be a vital and very useful source of germplasm not only for the country but also for the rest of the world.
In a country like Ethiopia, where research in plant breeding is in its
infancy because of a lack of adequately trained personnel and
inadequate facilities, it is only logical that the first step in breeding
should be maximum utilization of indigenous material. The dissemination of newly developed uniform varieties in any region is
highly risky. Due to their narrow genetic base, such varieties are
potentially vulnerable to diseases and pests that can cause extensive
yield losses. Broad genetic variability is believed to have a buffering
effect against the diverse production environments prevailing in the
country. Therefore, desirable results could be obtained through a
distribution and production programme of some of the better
varieties selected from indigenous landraces or segregations of their
most productive and valuable forms in hybridization programmes.
One way of improving the general productivity of wheat landraces
grown by the farmers is through phenotypic selection of the best
plant types from genetically mixed populations (positive mass selection) and subsequent bulking for further multiplication and distribution to the farmers. A slight modification of this would be to make
pure line selections of the different genetic lines through yield testing
and bulking two or more superior genetic lines for further multiplication and distribution to farmers.
Another strategy for the improvement of landrace populations of
tetraploid wheats in Ethiopia is based more or less on the multiline
concept, using the isogenic lines that exist in nature. In this approach,
it is assumed that a genetic line that grows in certain parts of the
country may have a gene(s) for resistance to certain races of a pathogen or pest prevailing in that particular area. When seeds of such
'identical genetic lines', each possessing different genes for
resistance, are composited, the resulting mixture will form a multiline
variety which will provide a partial protection or buffer against a
broad spectrum of races of pathogens or pests. This approach takes
the uniformity of seed colour into consideration and is now under
investigation in cooperation with the Swedish Agency for Research
Cooperation (SAREC).
In spite of their outstanding merits, Ethiopian wheats also have
Improvement of durum wheat landraces in Ethiopia
295
shortcomings such as low yield potential, unsatisfactory resistance to
leaf rust and weak straw. Therefore, the incorporation of desirable
characteristics of the most promising exotic varieties into the adapted
best local varieties, keeping the background of the local varieties,
should be the long-term objective of the durum wheat varietal
development programme. Along with an improvement of the characters mentioned, due consideration should be given to such research
as the development of:
- varieties that are responsive to small fertilizer inputs;
- varieties that are tolerant to drought, desiccating wind and
frost;
- better cropping systems that would increase soil fertility;
- better tillage practices that will help soil and moisture
conservation;
- varieties with improved grain quality.
To accomplish this task, the wheat breeding programme should
utilize the concerted efforts of breeders, agronomists, soil scientists,
protection experts and others relevant to the field. Such a holistic and
integrated team approach, it is hoped, would lead to self-sufficiency
in the ever increasing demand for wheat in general, and durum in
particular, thereby saving the nation substantial foreign exchange.
Acknowledgement
The writer is grateful to Dr Mesfin Abebe for his useful
suggestions on the preparation of this manuscript.
References
Central Statistics Office (1984). Ethiopian Statistical Abstract. CSO, Addis
Ababa.
Chavan, V. M. (1975). The importance of indigenous varieties in plant breeding. The Indian Journal of Genetics and Plant Breeding, 17(1), 1-6.
Harlan, J. R. (1971). Agricultural origins: centres and noncentres. Science, 174,
468-73.
Harlan, J. R. (1975). Our vanishing genetic resources. Science, 188, 618-21.
Nastasi, V. (1964). Wheat production in Ethiopia. Information Bulletin on the
Near East Wheat and Barley Improvement and Production Project, vol. 1 (13), pp.
13-23.
Porceddu, E., Perrino, P. & Olita, G. (1973). Preliminary information on
Ethiopian wheat germplasm collection mission. Proceedings, symposium on
genetics and breeding of durum wheat. University of Bari, Italy, pp. 181-99.
Tesemma, T. & Mohammed, J. (1982). Review of wheat breeding in Ethiopia.
Ethiopian Journal of Agricultural Science, 1, 11-24.
Worede, M. (1985). Crop genetic resources activities in Ethiopia. International Symposium on South East Asian Plant Genetic Resources, Jakarta,
20-24 August, 1985.
23
Use of germplasm resources in
breeding wheat for disease resistance
HAILU GEBRE-MARIAM
Introduction
Wheat (Triticum spp.) covers 12 per cent of the total area of
about 6 million hectares of land, producing approximately 7 million
tonnes of food grain, in Ethiopia. Both tetraploid and hexaploid
wheats are important in the farming systems; the former takes more
than 50 per cent of the share. Comprehensive wheat improvement
programmes for durum and bread wheat have been developed to
contribute to the overall effort of increasing food grain production in
the country.
Diseases are among the major constraints limiting wheat production. The wheat rusts (Puccinia spp.), Septoria diseases and Fusarium
spp. are the economically important diseases that therefore require
resistance breeding programmes. In particular, with the increase of
bread wheat acreage under the management of state farms, the
potential danger of yellow rust epidemics (caused by P. striiformis) in
Arsi and Bale highlands in southern Ethiopia warrants a well
organized breeding programme.
This paper deals with the approaches in breeding for disease
resistance with particular emphasis on the use of indigenous and
exotic germplasm resources in the bread wheat breeding programme
in Ethiopia.
Guiding principles
Considering the complexity of the biotic and abiotic environments of the Ethiopian agro-ecosystems, three guiding scientific
principles have been essential in planning and implementing the
wheat breeding programme for disease resistance. These are (a) care-
Wheat germplasm in breeding for disease resistance
ful consideration of the genetic variation phase of the breeding programme; (b) critical analysis of the crop ecosystem; (c) proper
evaluation of the pathosystem.
As more and more experience has been gained in breeding for
disease resistance, the importance of the acquisition, selection, evaluation and introgression of diverse germplasm to create as broad a
genetic base as possible has been recognized. The use of resistant
cultivars is the most economic and practical approach for the control
of cereal diseases, and this can only be attained by developing and
employing an effective resistance breeding programme with sufficient genetic variation. Current advances in the techniques of evaluating, selecting and incorporating germplasm into breeding stocks
have been essential in successfully developing disease-resistant highyielding wheat varieties. As the complexities of the host-pathogen
system in relation to the agro-ecosystem increase, sources of
resistance genes become limited.
Within a given agro-ecological zone of adaptation, crop breeding
involves the incorporation of a specific gene or gene complexes governing pest resistance and other essential traits. The Ethiopian crop
ecosystems are very diverse, consisting of a multiplicity of interacting
environmental factors such as altitude, climate, soils, etc. as well as of
a wide range of crop genetic diversity, ranging from primitive but
well adapted landraces to modern, highly uniform, high-yielding
varieties. The latter require high inputs whereas the primitive landraces frequently have to perform without any inputs. This has led the
wheat improvement programme to use different strategies in the
disease resistance breeding work to serve this diversity of conditions.
Sharp (1973), after investigating the significant influence on yellow
rust of the diverse agro-ecosystems of north-western USA, stressed
the difficulty of evaluating sources of resistance. In Ethiopia, the
complexity of these biotic and environmental factors is compounded
by the traditional subsistence farming systems which add a socioeconomic dimension to the problem. Because there is no organized
system for the transfer of farming technology, varieties released for a
specific agro-ecological condition frequently end up in the wrong
domain of recommendation. This results in lower yields caused by
the different wheat rusts. This problem is mainly due to the unavailability of sufficient seed of the recommended varieties and a lack of
essential extension information on existing varieties. Thus it is
necessary to consider the totality of the agro-ecosystems and the
socio-economic settings of the wheat producing areas in the country.
297
298
Hailu Gebre-Mariam
This requires not only the testing of breeding materials across diverse
agro-ecological zones, but also different scales of rating among
genetic sources and types of resistance and an assessment of traits
that show correlation between disease intensity and yield loss
(Roebelen & Sharp, 1978). Investigations such as this help to
determine the effectiveness, or buffering capacity, of resistance to
minimize yield losses when varieties are grown under different agroecological farming conditions.
In the traditional Ethiopian farming setting, both the Triticum spp.
and their pathogens are ancient and indigenous. Therefore, the relative levels of resistance or susceptibility of landraces and the virulence
of the pathogens fluctuate only to a limited extent. This avoids pressure due to a high level of damage or resistance, thus allowing the
host-pathogen balance to be maintained (Buddenhagen & de Ponti,
1984). The breeding programme, however, introduces into such a
system high-yielding homogeneous wheat varieties that remove the
suppressing effect on an epidemic that is provided by the mixtures of
genotypes in landraces. In other words, the successful introduction of
high-yielding wheat varieties has resulted in increased acreage of
extensive state farms which in turn has resulted in the buildup of
pathogen populations and an increasing level of inoculum sufficient
to cause outbreaks of epidemics. A good example of this is the 1981
yellow rust outbreak in Arsi and Bale highlands. In addition, an
epidemic level of yellow rust has been observed in 1986 and two
newly recommended varieties became susceptible. Whether this is
due to the appearance of a new race(s) or not is under investigation.
Hence, the wheat improvement programme is conscious of the fact
that a newly released variety will influence and be influenced by a
multitude of factors occurring in the local cropping system. Selection
for yellow rust resistance is further complicated because different
biotypes of P. striiformis and both specific and non-specific forms of
resistance may occur in Ethiopia.
Germplasm resources and introgression
To supplement the classical taxonomic classification of Linnaeus, Harlan & de Wet (1971) devised a gene pool or biological
species concept to group cross-compatible taxa. Of the three gene
pool categories (primary, secondary and tertiary) we have so far used
only those materials which fall into the primary gene pool. Nevertheless, germplasm of related wild species will be briefly mentioned in
this discussion because of its future potential.
Wheat germplasm in breeding for disease resistance
299
The three major germplasm sources for the national bread wheat
breeding programme are: modern advanced varieties and breeding
stocks which originate from national programmes, international
research centres, and sometimes genebanks; landraces which include
local primitive/traditional cultivars obtained from direct field collections, the Plant Genetic Resources Centre/Ethiopia (PGRC/E) and
other genebanks; and hybridizations.
Within the cultigen sources of germplasm, advanced varieties and
various breeding stocks are the economic and ready sources of desirable genes. Since the breeding programme is directed towards developing disease-resistant varieties for use in the short-term and
intermediate periods, the focus is on already improved gene sources
from national and international programmes. This includes both
selection under local conditions for direct use and the introgression of
unadapted germplasm into adapted genetic backgrounds to increase
useful genetic variability. This is a very valuable practice since favourable genes or gene complexes in unadapted germplasm may be
masked by major genes for adaptation to different environments
(Stuber, 1978). In this respect, the international research centres have
been invaluable sources of germplasm resources. In 1985-6 the programme handled a total of 19490 entries in this group. These were
150 genotypes in variety trials, 274 lines in advanced observations,
2831 nursery lines and 16235 in segregating populations. Most of the
segregating populations were generated from the local hybridization
programme.
During the past decade, interest in slow rusting or tolerant types of
resistance has increased. This type of resistance is of great potential
value because it may be more stable than that recognized by clear-cut
infection types (Andres & Wilcoxson, 1986). As a result of a study
carried out at Holetta, several high-yielding lines tolerant to Septoria
tritici blotch were identified (Gebre-Mariam & Gebeyehu, 1985) and
are presented in Table 1.
The bulk of the Ethiopian wheat germplasm resources are mostly
Triticum turgidum and T. dicoccum types. The diversity in hexaploid
wheat is limited. Therefore, a large proportion of 2384 local collections evaluated at Kulumsa and Ginchi in 1985 by the wheat programme were durum types. Of these, 24 lines were found to be
resistant to leaf rust (caused by Puccinia recondita) under field conditions. The major constraints in using these materials directly,
however, are their susceptibility to other diseases and undesirable
morphological characteristics.
300
Hailu Gebre-Mariam
Table 1. Comparative evaluation of three groups of bread wheat varieties
for their response to Septoria tritici blotch at Holetta
G7(R)
PY
HI
KQ
GY
lines
1.3
1.2
1.4
1.6
1.2
1.5
1.2
1.2
0.4
1.2
0.37
0.40
0.38
0.33
0.36
21
2+
11
6530
6140
6000
5760
5340
lines
2.4
2.3
2.2
2.4
1.8
1.8
2.3
1.5
2.3
1.8
0.31
0.34
0.36
0.32
0.34
1+
111
1-
6440
6290
6080
4900
5230
4.1
2.6
3.4
3.6
4.5
0.33
0.35
0.31
0.28
0.29
22
223+
5040
5400
5240
4650
4080
Variety
HT
Group 1
HAR499
HAR487
KCJ29
HAR503
HAR544
Resistant
105
100
105
125
90
Group 2
HAR466
YAL564
ET620BI
TAS879
ET12D4
Tolerant
135
142
105
113
88
Group 3
HAR118
PAN 504
EDC1138
Olaf
Derese
Susceptible lines
100
3.1
101
2.3
126
2.8
3.4
98
2.7
106
HT, Plant height (cm); G7(R), Septoria scoring using 0 (= no infection) to 5 (= high
level of flag leaf infection) at growth stage 7, i.e. milk development stage (after
Zadoks et al, 1974); PY, pycnidia count/mm 2 of the flag leaf; HI, harvest index; KQ,
kernel quality on a 1-3 scale with 9 classes, i.e. 1+ (excellent quality), 1 , 1 - , 2+, 2,
2 - , 3+, 3 and 3 - (very poor quality); GY, grain yield (kg/ha).
Currently, there is a programme of crossing and selection under
sub-optimum soil fertility conditions involving landraces or old/local
bread wheat cultivars and improved varieties. It is a study attempting
to answer some questions concerning the possibility of developing
wheat varieties with an economic level of grain yield under relatively
low soil fertility conditions so that they may be safely recommended
for low-input conditions.
Triticum aestivum is a classic example of utilizing wild species for
crop improvement via interspecific hybridization and genome building. The genus Triticum, containing more than 20 species, includes a
polyploid series ranging from diploids (2n = 14) to hexaploids
(2n = 42). Resistance to several diseases has been incorporated into
bread wheat from related species of Triticum, Aegilops and Agropyron
(Riley, 1965; Sears, 1969).
Among several methods developed to transfer genetic material
Wheat germplasm in breeding for disease resistance
from wild relatives to wheat, induction of homoeologous pairing and
crossing-over is by far the easiest (Sears, 1972). Substitution lines
have also been successfully used in transferring chromosomes carrying genes for disease resistance from Agropyron species to those of
wheat genomes (Schulz-Schaeffer and McNeal, 1977).
Although the wheat programme has not so far used wild relatives
of cultivated wheat species, it would be valuable to collect such germplasm for future use.
PGRC/E and the wheat improvement programme
A comprehensive and coordinated wheat improvement programme has been developed to assist in the urgent need for increased
food grain production in the country. This effort needs the strong
support of the genebank in the following areas:
- greater availability of different types of germplasm from
various sources;
- conservation of various types of germplasm for future use;
- registration of useful germplasm;
- preliminary evaluation and classification of available
landraces;
- collection of wild/weedy relatives of cultivated wheat in the
region.
The genebank can act as an important medium for acquiring cultigen germplasm for use by breeders, including advanced varieties and
breeding stocks from genebanks throughout the world and from
national and international breeding programmes. Secondly, the
genebank's role of storing/conserving and registering germplasm
introduced and/or developed by breeders could be of great assistance
to the national crop improvement efforts. During the last 20 years,
research institutions in the country have introduced and developed a
large number of germplasm accessions but none of these, with the
exception of those still under production, are available any more
because no proper long-term storage facilities existed in the country.
In the conservation of landrace germplasm, PGRC/E is doing a
commendable job. Nevertheless, acceleration of the classification and
preliminary evaluation work would encourage breeders to utilize
existing genetic resources more fruitfully. Since the genetic erosion of
wild relatives of cultivated species is equally serious, the landrace
collecting efforts of PGRC/E should also include the wild and weedy
species. As the existing sources of pest resistance in cultigens are
exhausted, the wild-weed-cultigen complex will be very essential.
301
302
Hailu Gebre-Mariam
Therefore, the germplasm resource collection, conservation and
utilization activities must be looked at holistically in terms of both
space and time.
References
Andres, M. V. & Wilcoxson, R. D. (1986). Selection of barley for slow rusting
resistance to leaf rust in epidemics of different severity. Crop Science, 26,
511-14.
Buddenhagen, I. W. & de Ponti, O. M. B. (1984). Crop improvement to
minimize future losses to diseases and pests in the tropics. FAO Plant
Production and Protection Paper No. 55, pp. 23-49.
Gebre-Mariam, H. & Gebeyehu, G. (1985). Evaluation of bread wheat
genotypes for their resistance to Septoria tritici blotch. In: A. L. Scharen
(ed.). Proceedings, Workshop on Septoria of Cereals, Bozeman, 2-4 August 1983.
Bozeman, Montana, pp. 27-30.
Harlan, J.R. & de Wet, J.M.J. (1971). Toward a rational classification of
cultivated plants. Taxon, 20, 509-17.
Riley, R. (1965). Cytogenetics and the evolution of wheat. In: Sir J. B. Hutchinson (ed.), Essays on Crop Plant Evolution. Cambridge University Press,
London, pp. 103-22.
Roebelen, B. & Sharp, E. L. (1978). Mode of inheritance, interaction and
application of genes conditioning resistance to yellow rust. Fortschritt der
Pflanzenziichtung, Beihefte zur Zeitschrift fur Pflanzenziichtung, Supplement 9,
pp. 1-88.
Schulz-Schaeffer, J. & McNeal, F. H. (1977). Alien chromosome addition in
wheat. Crop Science, 17, 891-6.
Sears, E. R. (1969). Wheat cytogenetics. Annual Review of Genetics, 3, 451-68.
Sears, E. R. (1972). Chromosome engineering in wheat. Stadler Genetics
Symposium, University of Missouri, Colombia, 4, 23-38.
Sharp, E. C. (1973). Wheat. In: R. R. Nelson (ed.), Breeding Plants for Disease
Resistance, concepts and applications. Pennsylvania State University Press,
University Park, pp. 111-31.
Stuber, C. W. (1978). Exotic sources for broadening genetic diversity in corn
breeding programs. In: H. D. Loden and D. Wilkinson (eds), Proceedings,
33rd Annual Corn and Sorghum Research Conference, Chicago, Illinois, 12-24
December 1977. American Seed Trade Association. Washington, DC, pp.
Zadoks, J. C , Chang, T. T. & Konzak, C. F. (1974. A decimal code for the
growth stages of cereals. Eucarpia Bull. 7, 42-52.
24
Indigenous barley germplasm in the
Ethiopian breeding programme
HAILU GEBRE AND FEKADU ALEMAYEHU
Introduction
Barley in Ethiopia is used for human food, home-made
beverages and beer. Its straw is used for animal feed and mattresses.
It is produced in the highlands at altitudes ranging from 1800 to
3300 m above sea level, where poor soil fertility, frost, waterlogging
and moderate soil acidity are major problems. It occupies an area of
about 0.85 million hectares out of a total crop land of 6.0 million
hectares with a productivity of about 1.2 t/ha (Central Statistics Office,
1984). At high altitudes it may be the only crop grown, with or
without oats; and among the small grains it is the earliest to become
available for consumption at the end of the rainy season.
Barley is grown in the main rainy season of June-September
('meher') on sloping and better drained clay soils. Some barley is
grown in the short rainy season ('belg') on bottom lands in some
regions. A few areas also grow barley from October to January
('bega'), sometimes with supplementary irrigation. Double cropping
is a common practice in the major barley growing regions: barleybarley in 'belg' and 'meher' seasons in Shewa, Welo, Arsi, and Bale;
barley-barley in 'meher' and 'bega' in Gojam; and barley-pulses in
Gondar.
The level of management is traditional. Farmers use their own
landraces in most cases. Application of inorganic fertilizers and use of
insecticides and herbicides are fairly low. The landraces are tolerant
to marginal soil conditions. At high altitudes fallow, sometimes
accompanied by soil heating, is a common practice to improve soil
fertility for barley production.
Table 1. Characters used and their respective classes (N varying from 3592 to 3759 accessions)
Frequency
distribution
Character
Character states
1. Kernel row number
6 rows
2 rows"
irregular
lax
intermediate
dense
<15
15-20
20-25
25-30
3=30
covered
naked
white-brown
purple-black
<25g
25-35g
35-45 g
45-55g
5=55 g
<100 days
100-115 days
115-130 days
0.42
0.57
0.01
0.42
0.48
0.10
0.03
0.22
0.49
0.24
0.02
0.97
0.03
0.72
0.28
0.08*
0.50
0.37
0.05
0.00
0.03
0.33
0.42
130-145 days
3=145 days
<60cm
60-90 cm
90-120 cm
120-150 cm
3=150 cm
0.21
0.01
0.01
0.34
0.59
0.06
0.00
2. Spike density
3. Spikelets per spike
4. Caryopsis
5. Kernel colour
6. Thousand grain weight
7. Days to maturity
8. Plant height
Mean
Minimum
Maximum
42
22.0
33.9
14
55
119.4
74
193
94.0
45
180
" Both categories, sterile and rudimentary lateral florets are combined.
b
These data originated from a smaller but representative sample of 2480 accessions of barley.
306
Hailu Gebre & Fekadu Alemayehu
Some characteristics of Ethiopian barleys
Ethiopia is considered to be a centre of diversity for barley
with many genotypes available. It has even been considered to be a
centre of origin (Negassa, 1985b). Ethiopian barleys are mostly tworowed (including deficiens), with lax to intermediate spike density,
white to brown or black coloured, covered kernels and with a diversity of time to maturity (Engels, 1988). Six-rowed genotypes are concentrated in Arsi, Bale and Shewa highlands because of their genes
conferring frost resistance (Negassa, 1985b). Ethiopian barleys are
distinctive in their range of grain colour, short-haired rachilla and the
deficiens and irregulare spike types (Ward, 1962; Qualset, 1975). Some
germplasm accessions have useful traits, especially for resistance to
diseases such as powdery mildew (Negassa, 1985a), barley yellow
dwarf virus, net blotch, scald, loose smut (Qualset, 1975) and high
protein quality (Munck, Karlsson & Hagberg, 1971). Evaluation of
some collections at the International Center for Agricultural Research
in the Dry Areas (ICARDA) indicates moderate resistance to lodging
and higher kernel weight when compared with collections from some
countries (Somaroo et ah, 1984).
Other useful characteristics of Ethiopian barleys include high tillering capacity; tolerance to marginal soil conditions, barley shoot fly,
aphids and frost; vigorous seedling establishment; and quick grain
filling period. On the other hand, they tend to show sensitivity to
lodging due to weak straw, low grain to straw ratio due to tall growth
of straw, and, in some, fragile rachis. The variations for some selected
characters are presented in Table 1.
Utilization of indigenous germplasm in the breeding
programme
The major centre for barley breeding and coordination since
1967 has been the Holetta Research Centre. The early breeding programme at Holetta focused mainly on the introduction and evaluation
of exotic lines using selection under optimum management conditions to develop food and malting barleys. A large number of
introductions were evaluated at Holetta but most lines were found to
be highly susceptible to scald, leaf blotch and barley shoot fly. Some
of the better genotypes were further tested in yield trials at various
locations and a few cultivars were recommended for commercial production. However, their adoption was slow because under traditional
management practices the farmers' landraces performed as well as
Barley germplasm in the Ethiopian breeding programme
307
Table 2. Mean grain yield, heading date and disease data of the
elite selections from Ethiopian collections at Bedi, 1977
Variety
Heading date
CI10389
IAR/H/199
IAR/H/165
IAR/H/189
CI12954
CI2376
HB-30
IAR/H/177
IAR/H/138
IAR/H/117
CI4375-1
IAR/H/195
Local check
Mean
100
100
101
104
105
96
97
100
100
102
106
108
101
102
Scald
(0-9)
Yield
(kg/ha)
6
7
8
8
6
8
8
7
8
7
4
6
7
2342
1883
2093
1993
2050
1781
1970
2066
2142
1846
1606
2105
1961
1988
174
353
12
SE(M)
LSD 5%
CV %
Table 3. Mean grain yield, heading date and disease data of the elite
selections from Ethiopian collections at Sheno, 1977
Variety
Heading
date
816/70
751/70
602/70
814/70
WGA44-4
WGA68-1
435/70
CI9724
Local check
Mean
83
86
89
85
91
77
83
87
74
84
Leaf
blotch
(0-9)
Leaf
rust
(%)
Reaction
type
Yield
7
7
7
8
8
8
8
8
7
80
15
80
90
100
80
60
80
40
MS
MS
MS
S
MS
S
MS
MS
S
1729
1527
1831
1250
1165
1345
1591
1186
1682
1479
445
NS
31
SE(M)
LSD 5%
CV %
MS, moderately susceptible; S, susceptible.
(kg/ha)
308
Hailu Gebre & Fekadu Alemayehu
the new cultivars. Based on this experience selection of indigenous
germplasm and a hybridization programme were initiated.
Selection of cultivars from landraces
In the 1970s, 3300 entries of local collections which were
made available to Holetta by the United States Department of Agriculture (USDA) and others collected by national staff were screened
at Holetta for disease resistance, plant vigour, medium maturity, stiff
straw, good grain set and high kernel weight. Additional collections
from the Plant Genetic Resources Centre/Ethiopia (PGRC/E) were
evaluated in the early 1980s. Some high-yielding selections were
identified from this activity. The elite material was further evaluated
from grain yield at Holetta and some other locations. A few cultivars
such as IAR/H/485 and ARDU 12-60B were recommended for commercial production.
About half of the original 3300 collections were also screened at
Bedi and Sheno, where barley is the only crop together with oats that
gives a dependable grain yield. Most selections were highly susceptible to scald. The best selections from the material were not superior
to established farmers' varieties (checks) at both locations as shown in
Tables 2 and 3.
A similar performance was observed at Mekele where soil moisture
stress is a limiting factor for crop production. The collections were
also screened for grain malting quality. Most of the grain samples
were found to be coarse, wrinkled and thin, and some were pigmented, with high nitrogen content, hence unsuitable for malting
purposes.
Within the collections about 200 hull-less types were isolated and
evaluated in comparison with hulled barleys for grain yield and
disease resistance. The naked barleys were found to be highly susceptible to scald, leaf blotch and lodging, with fairly low kernel weight
and grain yield, the latter being about half that of hulled barleys.
Hybridization and selection
This programme comprised breeding for malting and food
barley. The main objective in the malting barley breeding programmes was to incorporate disease resistance from local germplasm into
introduced malting barley cultivars, namely Proctor, Beka, Kenya
Research and Zephr using the backcross method. Initially these
cultivars were crossed to Holetta local which has good resistance to
scald and leaf blotch. In later years other satisfactory donor parents
Barley germplasm in the Ethiopian breeding programme
309
Table 4. Mean grain yield, heading date (HD) and disease data for seven
Holetta hybrid selections in comparison with ARDU 12-9C (local selection)
and the local check as tested at Holetta and Bekoji, 1985
Holetta
Bekoji
Variety
Origin
HD
Scald
(0-9)
Yield
(kg/ha)
HD
Scald
(0-9)
Yield
(kg/ha)
HB32
HB78
HB99
HB100
HB98
HB71
HB91
ARDU 12-9C
Local check
Mean
Hybrid
Hybrid
Hybrid
Hybrid
Hybrid
Hybrid
Hybrid
Arsi
87
96
83
83
74
86
90
85
86
86
7
2
4
7
6
8
3
4
8
5262
4177
5561
5005
4764
4007
4475
4929
3650
4648
290.4
847.7
12.5
92
103
93
92
98
91
96
97
78
93
7
5
6
8
3
8
7
6
8
5386
4356
5062
5616
5077
4237
4847
6066
3226
4875
332.9
971.8
13.7
SE (M)
LSD 5%
CV %
from other barley growing regions, as well as introduced lines, were
included. Some high-yielding selections such as Holkr, Balkr, HB-16,
HB-28 and others now featuring in yield trials were identified from
this effort.
In the food barley breeding project the aim was to improve yielding
capacity, disease and pest resistance, and nutritional quality of promising selections. Three methods of breeding were applied: backcross
for improving nutritional quality; modified pedigree; and composite
crosses for grain yield and disease resistance.
The activity on nutritional quality was a part of the
FAO/SIDA/SAREC Project. Five high lysine lines - Hiproly, Riso
1508, SV 73608, SV 75240 and SV 791582 - were used as donor parents
to which 20 cultivars from Ethiopia were crossed. Since 1978 about
1411 segregating populations have been evaluated at Holetta for plant
vigour, grain set, resistance to scald and leaf blotch, and for protein
quality. A large percentage of the material was highly susceptible to
diseases, together with poor plant vigour and grain set.
In the other approach to food barley breeding, cultivars from major
barley growing areas and promising introduced lines were included.
In addition, segregating materials from the International Centre for
Maize and Wheat Improvement (CIMMYT) were evaluated. The best
Table 5. Mean grain yield, heading date (HD) and disease data for selections (local and crosses) as tested at three
sites, 1986 (fertilized at 57/26 NIP kg/ha)
Debre Tabor
Variety
Origin
HD
HB42
HB43
ARDU12-60B
ARDU12-9B
HB37
A/HOR 880/61
Local check
Mean
Hybrid
Hybrid
Arsi
Arsi
Hybrid
88
82
85
82
85
95
60
82
SE(M)
LSD 5%
cv%
Spot
blotch
(0-9)
5
7
7
8
8
6
9
Gohatsion
Yield
(kg/ha)
HD
4458
4000
4725
4313
4238
3138
2375
3890
2488
1034
10.9
80
68
71
71
76
79
80
75
Spot
blotch
(0-9)
4
3
3
3
3
3
1
Shambu
Yield
(kg/ha)
HD
Scald
(0-9)
Yield
(kg/ha)
4638
4513
4963
4913
4100
4088
3163
4339
1564
541.3
5.09
85
71
54
71
88
86
57
73
5
5
5
6
5
6
6
4875
6875
5538
5500
4250
5000
3520
3937
363.8
1258.8
10.4
Table 6. Mean grain yield, heading date (HD) and disease data for selections (local and crosses) as tested at three
sites, 1986, without fertilizers
Gohatsion
Debre Tabor
Variety
Origin
HD
HB42
HB43
ARDU12-60B
ARDU12-9B
HB37
A/HOR 880/61
Local check
Mean
Hybrid
Hybrid
Arsi
Arsi
Hybrid
90
87
87
87
87
101
76
88
SE (M)
LSD 5%
cv %
Spot
blotch
(0-9)
5
7
6
7
8
7
9
Yield
(ka/ha)
HD
3750
4013
3538
3150
2938
2125
2250
3109
777.2
NS
35.4
81
81
80
80
79
80
80
80
Spot
blotch
(0-9)
3
4
5
3
3
3
2
Shambu
Yield
(ka/ha)
HD
Scald
(0-9)
Yield
(kg/ha)
3725
3438
4413
3313
3763
3563
2738
3593
257.8
89.25
10.2
91
84
90
87
90
93
75
87
5
5
5
6
5
4
6
2250
4500
3713
3000
1625
2750
1750
2798
963.3
3333
48.7
Table 7. Malting barley and food barley cultivars with their year of recommendation, trial average yield and suitable
areas
No.
Origin
Cultivar
Trial yield
Year
(t/ha)
Suitable areas
Beka
Kenya Research
Proctor
Holkr
Balkr
HB-16
HB-28
1973
1973
1973
1979
1979
1982
1982
3.9
2.7
2.4
2.4-3.8
2.5-3.7
3.5
3.3
Chilalo
Wolmera
Togulet & Bulga
Wolmera, Chilalo, Togulet & Bulga, and Degem
Wolmera, Chilalo, Togulet & Bulga, and Degem
Wolmera
Wolmera
IAR/H/485
Composite 29
A/Hor 880/61
HB-7
HB-15
HB-42
ARDU-60B
1975
1975
1978
1980
1980
1984
1985
3.6
4.1
4.5
3.9
3.7
4.2
3.8
Wolmera, Gondar, Chilalo and Alemaya
Wolmera, Gondar, Chilalo and Alemaya
Wolmera, Selale, Robe, Shambu
Debre Tabor, Mota
Wolmera
Wolmera, Chilalo
Chilalo, Wolmera, Robe
Malting barley
1.
2.
3.
4.
5.
6.
7.
France
Kenya
UK
ETH
ETH
ETH
ETH
FooaI barley
1.
2.
3.
4.
5.
6.
7.
ETH
USA
ETH(?)
ETH
ETH
ETH
ETH
Barley germplasm in the Ethiopian breeding programme
selections from this programme were advanced to yield trials and
were tested at several locations. The performance of such a group of
selections in comparison to local selections and cultivars (checks) is
shown in Tables 4, 5 and 6 for some barley growing areas. There was
not much difference among selections from crosses and local collections in grain yield; however, some selections from both groups were
superior to farmers' cultivars (local checks) which were earlier maturing but gave less response to nitrogen and phosphorus fertilizers.
Current emphasis in the breeding programme
The breeding programme from 1967 to date has generated a
number of high-yielding cultivars of malting and food barley (Table
7). So far, the uptake of malting barley releases has been satisfactory,
with state farms being the major producers, using a fairly high level
of management. On the other hand, the adoption of food barley
releases by peasants has been rather poor. As a result of this setback
the food barley breeding programme has been restructured. The programme now focuses on improvement of local landraces by mass
selection and cultivar mixtures without fertilizer application. A crossing programme between indigenous and exotic lines is under way
using the F2-progeny method with yield testing under low and
optimum fertilizer applications. The objective is to generate highyielding uniform bulk material.
References
Central Statistics Office (1984). Time series data on area, production, and yield or
principal crops by regions 1979/80-1983/84, vol. I. Central Statistics Office,
Addis Ababa.
Engels, J. M. M. (1988). A diversity study in Ethiopian barley. In: J. M. M.
Engels (ed.), The conservation and utilization of Ethiopian germplasm.
Proceedings of an international symposium, Addis Ababa, 13-16 October
1986, pp. 124-32 (mimeographed).
Munck, L. K., Karlsson, E. & Hagberg, A. (1971). Selection and characterization of a high protein, high-lysine variety from the world barley collection.
In: R. A. Nilan (ed.), Barley Genetics, vol. II. Proceedings of the 2nd International Barley Genetics Symposium, Pullman, 1969. Washington State Univer-
sity Press, Pullman, Washington, pp. 544-58.
Negassa, M. (1985a). Geographic distribution and genotypic diversity of
resistance to powdery mildew of barley in Ethiopia. Hereditas, 102, 113-21.
Negassa, M. (1985b). Patterns of phenotypic diversity in an Ethiopian barley
collection, and the Arsi-Bale Highland as a centre of origin of barley.
Hereditas, 102, 139-50.
Qualset, C. O. (1975). Sampling germplasm in a centre of diversity: an example of disease resistance in Ethiopian barley. In: O. H. Frankel and J. G.
313
314
Hailu Gebre & Fekadu Alemayehu
Hawkes (eds), Crop Genetic Resources for Today and Tomorrow. Cambridge
University Press, Cambridge, pp. 81-96.
Somaroo, B., Mekni, M., Adham, Y., Humed, B. & Kawas, B. (1984). Evaluation of barley germplasm at ICARDA. Rachis, 3, 12-15.
Ward, D.J. (1962). Some evolutionary aspects of certain morphological
characters in a world collection of barley. USDA Technical Bulletin 1276.
25
The role of Ethiopian sorghum
germplasm resources in the national
breeding programme
YILMA KEBEDE
Introduction
Sorghum is one of the crop types for which Ethiopia has been
credited as being a Vavilovian centre of origin or diversity (Harlan,
1969). In the different ecological zones of the country, germplasm
resources representing the major and intermediate races of sorghum
are found. In addition, the existence of wide variation in plant, grain,
inflorescence and fruit characteristics in the Ethiopian sorghum germplasm is well documented (Gebrekidan, 1973; Gebrekidan & Kebede,
1977). Among the sorghum growing population in the rural areas, the
importance of this crop is exemplified not only by its use as a staple
food and for other purposes, but also in the folklore, songs and some
of the local names by which the sorghum varieties are known.
As one of the leading traditional food cereals in Ethiopia, in terms
of both total production and area, major research efforts have been
directed towards the improvement and stabilization of sorghum
yields. At a national level, sorghum improvement involves the
manipulation of indigenous and introduced germplasm to develop
adapted types for the various ecological zones. In crop improvement
work the indigenous germplasm has been found invaluable
(Gebrekidan, 1981).
Periodic sorghum germplasm collections made throughout the
country have provided the sources of breeding material necessary for
the sorghum improvement programme. In the high altitude areas the
indigenous germplasm has often been the only adapted material suitable for use. From evaluations of germplasm collections, potential
varieties have been identified. Other accessions, which were found to
316
Yilma Kebede
be outstanding in certain features, have been used in crossing programmes. To date about 5-7 per cent of the evaluated collection has
been used in various breeding programmes (Kebede, Menkir &
Deressa, 1985). Development of new and better cultivars with
improved yield potential is a continuous process whether it be to
meet the needs of consumers or to stay ahead of yield-limiting factors. Through the use of conventional breeding methods - selection
and progeny testing and various crossing schemes - progress has
been made in utilizing local sorghum germplasm.
The objective of this chapter is to review work on sorghum breeding in Ethiopia with reference to the variation and utilization of indigenous germplasm.
Collection and characterization
The concern that some populations may become extinct
because of habitat destruction and other factors, as well as the expressed need for making useful germplasm readily available to the crop
improvement programme, led to the development of an organized
collection and conservation system.
Over the years, in addition to the Ethiopian sorghum programme,
contributions have been made to the sorghum collections by
individuals, research organizations, national establishments and the
Plant Genetic Resources Centre/Ethiopia (PGRC/E) staff. The current
number of indigenous sorghum germplasm accessions stands at more
than 8000 (PGRC/E, 1986). These accessions represent a wide array of
diversity as well as the major sorghum growing areas in the country.
These collections are routinely characterized and screened for
characters useful in crop improvement. In cooperation with PGRC/E,
germplasm collections are grown at appropriate adaptation sites for
evaluation of some agronomic and taxonomic characteristics (Table
1). The major characteristics evaluated fall into vegetative,
inflorescence and fruit and grain categories. These characterization
activities have been important in identifying desirable types with
useful traits either for direct use or in crossing programmes
(Gebrekidan & Kebede, 1977).
Moreover, attempts to classify the Ethiopian sorghum types into
recognizable taxonomic working groups or races have resulted in
recognition of 46 morphotypes representing four out of the five basic
races recognized by Harlan & de Wet (1972). Kafir sorghums are not
found in Ethiopia (Stemler, Harlan & de Wet, 1977). Data in Table 2
show that the dominant type of sorghum is the durra race with
Ethiopian sorghum germplasm in breeding programme
317
Table 1. Predominant characteristics in the Ethiopian sorghum
germplasm collection
Plant
Plant height
Inflorescence and fruit
Days to flowering
Inflorescence
Glume colour
Grain covering
Inflorescence exsertion
Grain
Grain colour
Grain size
Endosperm texture
Threshability
>2.0m
>100 days
loose to semi-loose erect branches
purple
a quarter covered
<10cm
red, white, brown
medium
mostly starchy
freely threshable
Table 2. Frequencies and distribution of races and sub-races in the
Ethiopian sorghum germplasm collection
Race
Number of
sub-racesa
Percentage^
Distribution
Durra
25
66
3
20
1
1
16
8
1
5
North-eastern, central and
eastern plateau
South and west of central
plateau and Rift Valley
West of Rift Valley (Metekel)
Konso
Variable
Caudatum
Bicolor
Guinea
Intermediate
a
b
Based on Abebe & Wech (1982).
Based on Gebrekidan & Kebede (1977).
characteristically large panicles and good grain types. Race caudatum
with asymmetrical grain shape is dominant in the Gambela area of
south-western Ethiopia. Variations as a result of intercrossing among
the basic races have resulted in intermediate races among which the
durra-bicolor sub-race is the most dominant.
A selected group of sorghum types, identified by local farmers for
attributes such as end use, quality, morphology and other traits, is
presented in Table 3. Admittedly most information on the value of
accessions must come from studies of experimental plantings in
which economically important traits can be observed. However, the
318
Yilma Kebede
Table 3. Local names and meaning of some Ethiopian sorghum landraces
grouped according to their most striking characteristics
Characteristic
End use
Quality
Morphology
Other
Ethiopian
sorghum
number
Local name
Meaning
1347
2861
1771
2390
3133
2624
2970
3149
—
—
3870
2611
3252
4762
Fendisha
Tinkish
Yeshet Ehil
Sinde Lemine
Gan Seber
Wetet Begunche
Marchuke
Dirb Keteto
Rejim Genbo
Shufun
Alequay
Hafukagne
Wof Aybelash
Kitgn Ayfere
Pop sorghum
Sweet stalk
Consumed green
Equals wheat
Good fermentation
Mouthful of milk
Oozing honey
Twin seed
Large sink
Large glumes
Faba bean-like seeds
Always heads
Bird-tolerant
Unafraid of syphilis (striga)
Adapted from Gebrekidan (1982).
first evaluation of an accession takes place in the natural habitat
where one can observe variation and other features of the population.
In this respect, the farmer's time-tested knowledge, as shown by the
names assigned to the various sorghums, becomes invaluable.
Utilization
In sorghum improvement work, some of the high priorities
are stand establishment, seed set, grain yield, tolerance to drought,
pests, Striga and food-type sorghums. Germplasm resources
represent a unique potential that could have an impact on these
aspects. The vital importance of genetic resources in crop improvement programmes has been amply demonstrated by the successes in
plant breeding using such resources. Our dependence on germplasm
resources is even more credible when we consider the achievements
of the past.
The Ethiopian sorghum germplasm has been found useful locally
and elsewhere as a source of cold tolerance, high protein (lysine),
good grain quality (zera-zeras), disease resistance and diversity, as
indicated by its use in the US Sorghum Conversion Program. Some of
the phenotypic diversity is illustrated in Table 4.
Ethiopian sorghum germplasm in breeding programme
319
Table 4. Phenotypic diversity expressed in range, mean and CV for some
quantitative characters of Ethiopian sorghum germplasm accessions
Range
Character
Minimum
Maximum
Mean
(%)
N
Plant height (cm)
Ear length (cm)
Ear width (cm)
Peduncle extension (cm)
Days to 50% flowering
Crude protein content (%)
Thousand seed weight (g)
19
4
2
1
76
5.0
6.0
475
50
30
44
169
15.3
61.1
233.7
21.5
9.4
14.7
116.7
9.6
28.5
22.7
36.6
34.1
53.8
14.3
13.1
35.5
2599
2511
2599
2254
2603
3644
200
Source: PGRC/E, unpublished data.
In the Ethiopian sorghum improvement programme, the utilization of germplasm focuses on manipulation of genetic stocks, specific
searches for useful genes and direct utilization of landraces.
Manipulation of genetic stocks
This includes hybridization, population improvement and
backcross breeding. Every season indigenous accessions selected for
desirable traits are used as parents in a crossing programme. In any
one year these parents comprise 20-30 per cent of the total number of
parents used in the crossing programme. The resulting 100-200 F2
generations having at least one indigenous parent are routed through
the pedigree system through planting in areas of adaptation (Kebede
et a\., 1985).
For population improvement, Ms3 and other genetic male sterile
carrier stocks are generally poorly adapted to the Ethiopian highlands. Thus an alternative approach in the Ethiopian sorghum programme is the use of the indigenous high-lysine, hi (Gebrekidan &
Kebede, 1979) and dented seed marker in identifying crossed seed.
Open pollination of the hi and dented (recessive) seeds with pollen
from regular (plump) seed parents results in a few (<10 per cent)
plump Fa seeds which can be visually identified and picked from the
mother panicle. The pollen source is a composite of elite high elevation adapted indigenous lines. The resulting plump Fx seeds are
selfed and segregate into a ratio of 3 plump :1 dented seeds on the
panicle. Through visual evaluation the best panicles are selected from
which the dented seeds are picked for the next cycle of random
320
Yilma Kebede
mating. At different stages, single plant selections have been evaluated in progeny rows and some promising lines have been advanced
to yield trials. This system enables the gathering of genes from
several different local sources into the hi germplasm. This gives rise
to several genotypes which, by reselection, produce further improvement. It has also opened an avenue for the intermating of tall
sorghum as well as the possible accumulation of high lysine genes
associated with the dented seed character (Gebrekidan, 1983).
A backcross breeding programme initiated to transfer resistance to
elite indigenous sorghums (ETS2111, ETS2113, ETS3235, WB-77)
that were found susceptible to anthracnose (Colletotrichum graminicola), using sources of known resistance, has resulted in resistant
types and a good agronomic performance. Details of this programme
have been given by Menkir, Kebede & Gebrekidan (1986).
Specific searches for useful genes
The Ethiopian sorghum germplasm resources have proved to
be useful sources for desirable genes. As a result of pointed evaluation of some of the collections or from information based on farmers'
own experiences, some useful genes reported to exist in the Ethiopian
germplasm are listed below.
Disease and pest resistance
- Stalk borer
- Downy mildew
- Smuts
- Bacterial streak
- Anthracnose
- Striga
Adaptation
- Agronomic desirability (height, maturity, panicle size
and shape, etc.)
- Yield potential
Kernel traits
- Endosperm texture
- Grain colour
- Threshability
- Injera quality
- Protein quality (hi)
Ethiopian sorghum germplasm in breeding programme
321
Direct utilization of germplasm
In general, sorghum landraces are a mixture of related pure
lines. Testing and seed multiplication and further maintenance
require some sort of varietal identity. Thus, panicle selections from
promising accessions are put in progeny rows from which elite types
are selected and put into the Advanced Sorghum Selections Nursery
(Kebede et al., 1985). This nursery serves as an intermediate evaluation stage for entries selected from new accessions before advancing
to national and pre-national yield trials. In the past, in any given
season, approximately 2 per cent of the accessions were advanced to
such a nursery.
Over the years, about a dozen entries derived from the collections
have been recommended for release and currently five such entries
are on the recommended list. They are Alemaya 70 and ETS 2752 for
areas of high elevation, Dedessa 1057 and Asfaw White for intermediate elevation and Gambela 1107 for low elevation (Kebede et al., 1985).
Summary and projections
The remarkable diversity in crop germplasm in Ethiopia is
now widely recognized but these resources have only recently started
to be tapped. The inability to reconstitute lost germplasm underlines
the necessity for conservation before this natural wealth is depleted
completely. Evaluation and characterization of available germplasm
based on needs and requirements of the crop improvement programme would aid in better utilization of available resources. Some of
the future activities should take into account that:
- evaluation and documentation of sorghum genetic resources
have to be accelerated in order to achieve the desired goal of
utilization;
- collecting missions have to concentrate on hitherto unexplored areas since expansion of production into new environments may require attributes not presently considered
important;
- evaluation of germplasm has to emphasize screening for
resistance to drought, pests and Striga.
The importance of germplasm availability for the continued
improvement of sorghum has now been recognized and one of the
most significant developments in this field in Ethiopia in the past
decade has been the establishment of PGRC/E. The Centre has fulfilled its functions in developing appropriate systems of conservation
and characterization and we shall look to PGRC/E to help breeders
322
Yilma Kebede
find useful germplasm as the need arises. The joint efforts of PGRC/E
and the crop breeders will enable all to cope with the varied production environments through the development of germplasm that could
increase and stabilize production.
References
Abebe, B. & Wech, H. B. (1982). The 1981 activities of the Plant Genetic
Resources Centre/Ethiopia. In: Proceedings of the Regional Workshop on
Sorghum Improvement in Eastern Africa, 17-22 October, 1982, Addis Ababa, pp.
31-45.
Gebrekidan, B. (1973). The importance of the Ethiopian sorghum germplasm
in the world sorghum collection. Economic Botany, 27, 442-5.
Gebrekidan, B. (1981). Salient features of the sorghum breeding strategies
used in Ethiopia. Ethiopian Journal of Agricultural Science, 3, 97-104.
Gebrekidan, B. (1982). Utilization of germplasm in sorghum improvement.
In: Sorghum in the Eighties. Proceedings of the International Symposium
on Sorghum, 2-7 November 1981. ICRISAT, Patancheru, pp. 335-45.
Gebrekidan, B. (1983). New breeding concepts in partially self pollinated
crops with special emphasis on sorghum. In: J. C. Holmes and W. M. Tahir
(eds), More Food from Better Technology. FAO, Rome, pp. 186-92.
Gebrekidan, B. & Kebede, Y. (1977). Ethiopian Sorghum Improvement Project. Progress Report No. 5. Addis Ababa University (mimeographed).
Gebrekidan, B. & Kebede, Y. (1979). The traditional culture and yield potentials of the Ethiopia high-lysine sorghums. Ethiopian Journal of Agricultural
Science, 1, 29-40.
Harlan, J. R. (1969). Ethiopia: a centre of diversity. Economic Botany, 23,
309-13.
Harlan, J. R. & de Wet, J. M. J. (1972). A simplified classification of cultivated
sorghum. Crop Science, 12, 172-6.
Kebede, Y., Menkir, A. & Deressa, A. (1985). A review of sorghum research
work in Ethiopia. Paper presented at Workshop on Review of Field Crops
Research in Ethiopia, 25 February-1 March 1985, Addis Ababa
(mimeographed).
Menkir, A., Kebede, Y. & Gebrekidan, B. (1986). Incorporating anthracnose
resistance into indigenous sorghum. Ethiopian Journal of Agricultural Science,
8, 73-84.
Plant Genetic Resources Centre/Ethiopia (1986). Ten years of collection, conservation and utilization, 1976-1986. PGRC/E, Addis Ababa.
Stemler, A.B.L., Harlan, J. R. & de Wet, J.M.J. (1977). The sorghums of
Ethiopia. Economic Botany, 31, 446-60.
26
Germplasm evaluation and breeding
work on teff (Eragrostis tef) in
Ethiopia
SEYFU KETEMA
Introduction
Ethiopia is the only country that produces teff as a cereal
crop. Teff occupies the largest area of cultivated land under cereal
production in Ethiopia, and as such it is the most important crop.
According to the statistical information of five years' average from
1979-80 to 1983-4, teff was cultivated each year on 1.385 million
hectares, followed by barley 0.851, wheat 0.609, maize 0.780 and
sorghum 0.994 million hectares. The national average grain yield of
these cereals for the same five-year period was 9.1 quintals per hectare (q/ha) for teff, barley 11.83, wheat 11.26, maize 17.35 and
sorghum 14.57q/ha (Central Statistics Office, 1984).
Teff is mainly cultivated as a single crop. However, in a few areas it
is cultivated under a multiple cropping system. In such cases it is
usually grown as an intercrop with Brassica carinata, Carthamus tinctorius or Helianthus annuus. It is also relay cropped with Zea mays and
Sorghum bicolor.
Teff is mainly used for making a pancake-like bread called 'injera'.
In some cases it is used to make porridge and native alcoholic drinks
called 'tella' and 'katikala'. Its straw is highly valued and is used as
feed for cattle. In addition, the straw is incorporated with mud to
reinforce it and used for plastering walls of houses.
Teff is on average as nutritious as any of the major cereals. According to Rouk and Mengesha (n.d.), the Ethiopian Nutrition Survey
reported that four unspecified teff varieties when analysed were
found to contain an average of 300 calories, 11.6 g protein, 0.65 g fat
324
Seyfu Ketema
Table 1,. Content of seed grain (proximate analysis) as percentage of grain
Item
Teff
Protein
11.0
Fat
2.6
3.5
Fibre
Carbohydrate 73.0
Minerals (ash) 3.0
Wheat : Rice
11.0
1.9
1.9
69.3
1.7
9.7
1.8
8.8
64.7
5.0
Maize
Sorghum Barley
9.4
4.4
2.2
69.2
1.3
8.6
3.8
1.9
71.3
2.4
8.5
1.5
4.5
67.4
2.6
Oats
Rye
9.5
4.8
10.3
58.4
3.1
10.7
1.7
1.9
69.8
2.0
Source: B. Tareke, unpublished data, Alemaya University of Agriculture, Dire
Dawa, Ethiopia.
and 70.56 g carbohydrate per 100 g, and that teff supplied an average
of two-thirds of the protein in the Ethiopian diet. Table 1 shows the
nutritional status of teff compared with some other cereals.
Teff also contains more calcium, copper, zinc, aluminium and
boron than winter wheat, winter barley and sorghum (Mengesha,
1966).
Some of the reasons why present-day farmers grow teff are given
as follows.
1. It can be grown in areas experiencing moisture stress.
2. It can be grown in waterlogged areas and withstands
anaerobic conditions better than many other cereals including
maize, wheat and sorghum.
3. It is suitable for use in multiple cropping systems such as
double, relay and intercropping.
4. Its straw is a valuable feed during the dry season when there
is an acute shortage. It is highly preferred by cattle over the
straw of other cereals and demands higher prices in the
markets.
5. It has acceptance in the national diet, has high demand and
high market value and hence enables farmers to earn more.
6. It is a reliable and low-risk crop.
7. In moisture stress areas farmers use it as a rescue crop. For
example, around Kobo and Zeway, which are areas with low
and erratic rainfall, farmers first plant maize around April. If
this fails after a month or more due to moisture stress or pest
problems they plough it under and plant sorghum. If this also
fails after a month or more then they sow teff as a last resort,
which often survives on the remaining moisture in the soil
and yields some grain for human consumption and straw for
feed.
Teff germplasm evaluation and breeding in Ethiopia
325
8. It is not attacked by weevils and other storage pests and
therefore is easily and safely stored under local storage conditions. This results in reduced post-harvest management
costs.
9. Compared with any other cereal grown in Ethiopia it has
fewer disease and pest problems (Stewart & Degnachew,
1967).
Domestication and diversity
According to N. I. Vavilov, Ethiopia is the centre of origin for
teff (Mengesha, 1966; Tadesse, 1975). Its domestication is believed to
have taken place first in the northern highlands of Ethiopia (Tadesse,
1975). Although there are no exact records on the history of its
domestication, one hypothesis to explain the situation that led to its
domestication is as follows.
The word teff is said to have originated from the Amharic word
'teffa' which means lost, because the grain size of teff is so small that
if one grain is dropped it is difficult to find (Rouk & Mengesha, n.d.).
Other sources say that the word teff was derived from the Arabic
word 'tahf, a name given to a similar wild plant (Eragrostis sp.) used
by Semites in South Arabia in times of food scarcity (Constanza, 1974;
Tadesse, 1975; Endeshaw, 1978). According to Endeshaw (1978)
Ciferri and Baldrati stated in 1939 that E. pilosa, which is believed to
be the ancestor of teff, is collected as food by people in many parts of
Africa other than Ethiopia in times of famine. This suggests that the
domestication of teff in Ethiopia might have started in times of food
scarcity (Tadesse, 1975; Endeshaw, 1978).
Teff was introduced to other parts of the world by the Royal
Botanic Gardens at Kew, which imported seed from Ethiopia in 1866
and distributed it to India, Australia, California and South Africa. In
1916 Burt Davy introduced teff into California, Malawi, Zaire, India,
Sri Lanka, Australia, New Zealand and Argentina; Skyes introduced
it in 1911 into Zimbabwe, Mozambique, Kenya, Uganda and
Tanzania; Horuity in 1940 introduced it into Palestine (Tadesse, 1975).
Teff is a sexually propagated, self-pollinating annual grass species
(Tadesse, 1975; Tareke, 1975). It is tetraploid with a chromosome
number of 2n = 40 (Tareke, 1975; Endeshaw, 1978) and an
allopolyploid (Tareke, 1981). It is cultivated from sea level up to
2800 m on soils with varying physical and chemical properties, in
waterlogged and in well drained soils, in moisture stress areas having
less than 300 mm of rainfall as well as in areas having 1000 mm
326
Seyfu Ketema
seasonal rainfall. This gives an idea of the tremendous ecological
diversity under which the crop can be grown.
The Plant Genetic Resources Centre/Ethiopia has itself made 1050
germplasm collections (PGRCE, 1986). More than one thousand collections made by Debre Zeit Agricultural Research Centre have been
given to PGRC/E also. However, many of these collections have not
been made from representative sites nor were they collected in a
systematic way and with adequate passport data. All this is now
being corrected and a systematic collection, characterization and
utilization programme is in progress, the last named in conjunction
with the breeding programme. In Ethiopia 54 Eragrostis spp. are listed
of which 14 (or 26 per cent) are endemic (Constanza, 1974). However,
so far no collection of the wild species has been made. The total
number of accessions of teff germplasm at present is 2175. Tadesse
(1975) has characterized and recognized 35 cultivars, and Endeshaw
(1978) mentioned that minor variations still exist within many of the
cultivars. Some of the variations noted so far include maturity period
60-120 days, plant height 45-150cm, culm thickness 1.2-3.1 cm,
panicle varying from very loose to very compact, lemma colour
whitish green, purple, olive grey, pink and various types of these
colour combinations, seed colour white, yellowish brown, dark
brown and intermediate types.
Utilization
Improvement work on teff through plant breeding began in
the late 1950s. This was done only through pure line selection from
landraces, since it was then not possible to produce hybrids. In 1972,
in order to create variability, mutation breeding was started. This line
of investigation established that useful mutations could be induced
through the use of physical mutagens such as X-rays at 100-130 krads
and Gamma rays at 150 krads, and chemical mutagens such as
ethylmethyl-sulphonate at 2.5^.7 per cent concentration. Until 1974
artificial pollination was attempted in the morning after 0800 h with
no success. The observation by Tareke that teff flowers open during
the early morning (0645-0745 h) and that they have only a brief pollination time enabled him to make the first successful intraspecific
crosses towards the end of 1974 (Tareke, 1975). Now it is realized that
artificial crosses may be made either early in the morning between
0600 and 0730 h, or any time during the day, provided that the natural
pollination time is delayed through the use of low temperature 4-5 °C
Teff germplasm evaluation and breeding in Ethiopia
327
or dark treatment, which can be achieved by putting potted plants
under cold or dark conditions overnight. Artificial hand pollination is
time consuming and cumbersome. The latest available technique for
such work, which was suggested by Seyfu (1983); is given as follows.
- Grow one or two plants in a pot of about 13 cm diameter.
- Eight to 18 days after anthesis begins on the central tiller (or
any other particular tiller), put the seed parent plant and the
pollen donor plant into separate light-tight dark boxes at
around 1400 h. Keep the boxes away from direct sunlight at a
temperature well below 28 °C (lower temperature improves
degree of control over flowering). Next day, crossing may be
done at any time before the early afternoon.
- Take donor plant out first. As soon as it starts to open its
flowers, detach those spikelets with open florets using
forceps and attach them to the moist inner wall of a vial.
Label vial with code number of plant.
- Take out seed plant, lay it horizontally under a binocular
microscope (xl5) and begin to emasculate as soon as flowers
start to open (and before the anthers dehisce). Only the basal
florets should be emasculated, the other florets on the
spikelet being removed. This serves to identify the treated
flowers.
- Keeping the emasculated floret under observation, remove a
spikelet from the vial and, with forceps, detach an individual
anther while observing it under the binocular microscope,
and gently squeeze it to release pollen directly onto the
stigma of the emasculated floret.
- Label plant for crossing record.
Achievements
The major objective of the breeding programme was and still
is to develop lodging resistant, high-yielding and stable varieties. In
the major teff growing areas no epidemic disease or pest problems
exist. Therefore, the development of disease and pest-resistant
varieties or ones which have high nutritive value (e.g. high protein
content) is not the major focal point at the moment. The national
average grain yield of teff for landraces is 9.1 q/ha. Improved varieties
that outyield the landraces have been developed through pure line
selection from germplasm as well as through hybridization following
the pedigree method of selection. These varieties give a grain yield of
328
Seyfu Ketema
17-22 q/ha on the farmer's field. Some of these released and recommended varieties are DZ-01-354, DZ-01-99, DZ-01-196, DZ-cross-44,
DZ-cross-82 and DZ-01-787.
References
Central Statistics Office (1984). Time series data on area, production and
yield of principal crops by regions 1979/80-1983/84. Central Statistics
Office, Addis Ababa, 219 pp.
Constanza, S. H. (1974). Literature and numerical taxonomy of teff (Eragrostis
tef). MSc thesis, Cornell University, Urbana, Illinois, USA.
Endeshaw, B. (1978). Biochemical and morphological studies of the relationships of Eragrostis tef and some other Eragrostis species. MSc thesis, University of Birmingham, UK.
Mengesha, M. H. (1966). Chemical composition of teff {Eragrostis tef) compared with that of wheat, barley and grain sorghum. Economic Botany, 20,
268-73.
Plant Genetic Resources Centre/Ethiopia (1986). Ten years of collection, conservation and utilization 1976-1986. PGRC/E, Addis Ababa.
Rouk, H. F. & Mengesha, M. H. (n.d.). An introduction to teff (Eragrostis
abyssinica Schad.). A nutritious cereal grain of Ethiopia. Debre Zeit Agricultural Research Centre Bulletin No. 26. Alemaya University of Agriculture,
Dire Dawa, Ethiopia.
Seyfu, K. (1983). Studies of lodging, floral biology and breeding techniques
in teff (Eragrostis tef (Zucc.) Trotter). PhD thesis, University of London.
Stewart, R. B. & Degnachew, Y. (1967). Index of plant diseases in Ethiopia.
Debre Zeit Experimental Station Bulletin No. 30. Alemaya University of Agriculture, Dire Dawa, Ethiopia.
Tadesse, E. (1975). Teff (Eragrostis tef) cultivars; morphology and classification. Part II. Debre Zeit Agricultural Research Centre Bulletin No. 66, pp. 1-73.
Alemaya University of Agriculture, Dire Dawa, Ethiopia.
Tareke, B. (1975). A breakthrough in teff breeding technique. FAO Information bulletin on cereal improvement and production 12, 11-13.
Tareke, B. (1981). Inheritance of lemma color, seed color and panicle form
among four cultivars of Eragrostis tef (Zucc.) Trotter. PhD thesis, University
of Nebraska, Lincoln, Nebraska.
27
Pulse crops of Ethiopia: genetic
resources and their utilization
HAILU MEKBIB, ABEBE DEMISSIE AND ABEBE TULLU
Introduction
Ethiopia is known as a centre of diversity and/or origin of
numerous cultivated crop plant species. This was first recognized by
N. I. Vavilov in the late 1920s and later confirmed by several other
scientists. Vavilov (1951) indicated that some 38 crop plants have their
centre of diversity in the Ethiopian region. Zohary (1970) mentioned
11 crop species which had their centre of diversity in Ethiopia. Primitive cultivars or landraces and wild relatives of some of the world's
major crops are found in the country. Pulse crops form a significant
portion of the available genetic resource base for plant breeding
programmes.
In this chapter an attempt is made to describe the situation for the
most important pulse crops cultivated in Ethiopia regarding their
diversity and the germplasm kept in the national collection, and their
conservation, evaluation and utilization.
Collection
Owing to the richness and potential of the biological
resources of the country, numerous plant expeditions have been
undertaken by scientists in the past. However, it was only after the
establishment of the Plant Genetic Resources Centre/Ethiopia
(PGRC/E) in 1976 that systematic collecting missions were launched.
The total holding of pulse accessions by PGRC/E is about 4300. The
bulk of the germplasm was acquired from field collecting (ca. 2900) on
the basis of a well defined strategy, and some was acquired through
repatriation and acquisition from national and international sources.
The sampling procedure and techniques are based on a field col-
330
Hailu Mekbib, Abebe Demissie & Abebe Tullu
lecting manual (Hawkes, 1980), which advises that seeds from up to
50 individual plants, and certainly not more than 100, should be
collected non-selectively and bulked to obtain an optimum sample.
Whenever rare types, i.e. plants which show characters not included
in the random sampling, are noticed, a selective sampling technique
is adopted. Such a sample is given a different collection number.
Conservation
Generally, the pulse crops under review are conserved in
temperature-regulated storage units. Seeds intended for both base
and active collections are dried to 3-7 per cent moisture content and
maintained at —10 °C in laminated aluminium foil bags. Accessions
which are too small to meet the sample size required for long-term
storage are maintained in paper bags at 4 °Cand 35 per cent relative
humidity (Feyissa, 1988). The plans for in situ conservation of yeheb
nut (Cordeauxia edulis) are based on the absence of factual data on its
storage behaviour and need for its immediate conservation
programme.
In Ethiopia farmers play a pivotal role in the conservation of landraces as they hold the bulk of genetic resources. Peasant farmers
always retain some traditional seed stock for security even at difficult
times unless circumstances dictate otherwise (Worede, 1987). This
strategy of conservation is the second major option considered in
Ethiopia.
Distribution and diversity
The distribution and the degree of genetic diversity of pulse
crops in different agro-ecological zones of the country has not yet
been adequately studied. A modest programme of work on crops
such as Lathyrus and Vigna has been initiated with a view to identifying areas of maximum diversity. This will help in subsequent rational
planning of collecting missions. An estimate of the diversity of some
Ethiopian pulse crops is presented in Table 1 which was compiled by
pooling available data (Mengesha, 1975) with data generated during
collecting and preliminary evaluation activities.
Both intra- and infraspecific diversity in legumes are relatively
large in Ethiopia. Crops such as faba bean, field pea, chickpea and
lentil have their (secondary) centres of diversity in Ethiopia. There are
ca. 600 species of legumes recorded in Ethiopia (Thulin, 1983) of
which only a handful, namely, peas, lentil, chickpea, common bean,
Table 1. Estimate of crop diversity in Ethiopia
Estimate of crop diversity
Administrative
region
Faba
bean
Field
pea
Chickpea
Lentil
Arsi
Bale
Eritrea
Gamo Gofa
Gojam
Gondar
Harerge
Ilubabor
Kefa
Shewa
Sidamo
Tigray
Welega
Welo
H
M
M
M
M
H
M
L
L
H
L
M
M
H
M
M
M
L
H
H
L
—
T
H
L
M
M
H
M
L
M
T
H
H
M
—
—
H
M
H
—
M
L
L
L
L
M
H
L
—
—
H
—
M
—
H
Grass
pea
_
M
H
H
—
_
M
M
—
M
Fenugreek Lablab
L
L
M
M
H
M
—
—
M
—
M
M
H
H, M and L, high, medium and low diversity; T, trace; —, not enough data available.
Based on Mengesha (1975) and field observations of the authors.
Pigeon
pea
Common
bean
_
_
H
—
—
—
—
M
—
—
H
M
M
H
M
—
L
M
M
L
—
—
—
M
-
M
M
—
—
L
332
Hailu Mekbib, Abebe Demissie & Abebe Tullu
faba bean and cowpea are largely grown as grain legumes for human
consumption.
Faba bean (Vicia faba L.)
The origin of faba bean is so far not clearly established. In
spite of claims by Abdella (1979) that faba beans originated in Egypt,
most recent studies have indicated that the crop perhaps originated in
west (Cubero, 1974) or central Asia (Ladizinski, 1975), but both the
progenitor and place of origin remain uncertain. It is believed that
faba beans were cultivated at an early date in the Nile Valley as far
south as the Ethiopian highlands and as far east as Afghanistan. In
both these regions primitive forms with small and sometimes black
seeds are still cultivated (Hawtin & Hebblethwaite, 1983). Furthermore, Westphal (1974) indicated that the origin of V. faba is in the
Mediterranean region or south-western Asia where it has been
cultivated for centuries. V. pliniana, which grows wild in Algeria, is
said to be its close relative.
Faba bean is one of the most common and major pulse crops in
the highlands of Ethiopia, occupying 6 per cent of the total area
under cultivation by the major crops. It was collected from areas
ranging in altitude from 1600 to 3200 m above sea level; a few accessions were made between 3800 and 4000 m, although the majority
come from 2200 to 2800 m. It is generally planted in June when the
main rainy season commences and harvesting is carried out in
December-January.
Faba bean is usually planted alone. It is sometimes cultivated
together with field pea for support and it is usually incorporated in a
rotation scheme, coming immediately after cereals.
Utilization
Varietal improvement efforts are at rather an early stage. The
first national yield trial was initiated only in 1972 and had to be
discontinued in 1974 due to inadequacy of the working collection
(Telaye, 1988).
Recently, 349 faba bean germplasm accessions have been evaluated
for grain yield and other agronomic characters at several locations. Of
these, 28 high-yielding accessions were identified and selected for
further multilocation yield testing in the national programmes
(Ghizaw et ah, 1986). Yield trials have shown promising results and
new potential cultivars are being selected and recommended for
various agro-ecological zones within the country.
Pulse crop genetic resources
333
PGRC/E has collected 744 accessions of faba bean from various
agro-ecological zones. About 450 of these accessions have been evaluated for a number of agro-morphological characteristics on the basis
of International Board for Plant Genetic Resources (IBPGR) descriptors, synthesized by the PGRC/E in collaboration with the national
crop breeders. A summary of the evaluation data for faba bean is
presented in Table 2.
Field pea (Pisum sativum L.)
Field pea was perhaps domesticated in central or western
Asia, spread to China and India, and reached Africa and the mountain regions of Ethiopia before the arrival of the Europeans. In recent
work (van der Maesen et ah, 1988) four possible centres of diversity
are mentioned, namely, the Near East, the Mediterranean, central
Asia and Ethiopia.
Extensive areas of the central and northern Highlands of Ethiopia
are cultivated with field pea. Although it is a typical plant of high
elevation, some varieties thrive well at lower elevations. Field pea has
been collected in areas as low as 1560 m and as high as 3560 m, but the
bulk of the PGRC/E collection comes from areas between 2160 and
2760 m. The total holding of field pea is close to 1060 accessions.
There are two main cultivar groups, abyssinicum and sativum. The
former has leaves with one pair of leaflets, the flowers are small and
pink or purple, the seeds are globose, brownish or grey, often with
blotches and a black hilum. The latter is usually more robust but less
hardy, the flowers are white, pods and seeds are larger and the seeds
are yellowish round and smooth or wrinkled. The two forms of
cultivated pea are sometimes regarded as separate species, P. arvense
and P. sativum respectively, but there is little taxonomic ground for
treating them so. They are genetically similar and interbreed readily,
producing fertile progeny. In Ethiopia, the abyssinicum type is predominantly grown.
Utilization
Since the early days of research efforts in Ethiopia, agronomy
work has been largely based on local landraces procured from the
local markets or farmers' fields. As a result cultivars such as CS436,
FPEXDZ, and G2276-2c were released (Telaye, 1988). Recently the
Ethiopian field pea germplasm collection has been evaluated for
several morphological and agronomic characters. It has shown considerable diversity for the characters considered (Table 2).
Table 2. Summary of evaluation data for chickpea, field pea, faba bean,
lentil, grass pea, fenugreek and common bean
Crop
Chick pea
Days to flowering
Primary branches
Secondary branches
Days to maturity
Plant height
Pods per plant
Seeds per pod
Field pea
Days to flowering
Days to maturity
Seeds per pod
Seed weight
Faba bean
Days to flowering
Flowers per plant
Days to maturity
Plant height
Pods per plant
Seeds per pod
Seed weight
Lentil
Days to flowering
Primary branches
Secondary branches
Days to maturity
Pods per plant
Seeds per pod
Grass pea
Days to flowering
Days to maturity
Plant height
Pods per plant
Seeds per pod
Thousand seed weight
Fenugreek
Days to flowering
Days to maturity
Plant height
Pods per plant
Seeds per pod
Thousand seed weight
Common bean
Days to flowering
Days to maturity
Plant height
Pods per plant
Seeds per pod
N
Max.
Min.
Mean
STD
cv
639
640
639
639
638
639
332
63.0
3.0
6.2
123.0
48.4
141.0
2.4
36.0
1.0
1.8
84.0
16.0
10.0
1.0
46.5
2.0
2.9
97.1
29.5
33.8
1.5
4.7
0.2
0.7
5.8
6.1
14.7
0.2
10.1
11.2
26.7
6.0
20.6
43.5
15.9
1063
1052
876
203
180.0
183.0
11.0
227.4
32.0
92.0
1.0
10.8
74.0
131.8
5.5
123.3
12.9
17.9
0.9
18.5
17.4
13.5
16.3
15.0
452
410
392
452
452
452
97
88.0
87.0
182.0
195.0
50.0
5.0
521.6
33.0
20.0
95.0
50.0
3.0
1.0
208.3
62.9
42.5
141.5
100.7
14.3
2.3
317.1
7.9
14.9
19.1
24.2
5.9
0.5
73.2
12.6
35.1
13.5
24.0
41.5
24.1
23.1
683
516
516
670
516
516
79.0
6.0
28.0
132.0
159.0
2.2
33.0
2.0
2.0
78.0
13.0
1.0
53.5
2.6
7.4
98.8
49.8
1.8
8.0
0.5
5.8
13.2
21.8
0.2
15.0
21.9
78.0
13.4
43.9
14.8
114
114
114
75
75
40
73.0
160.0
94.8
121.0
4.0
681.6
30.0
92.0
37.6
14.2
1.0
61.8
52.4
116.5
59.1
43.8
2.4
253.2
9.1
27.3
11.3
25.7
0.6
113.2
17.3
23.4
19.1
58.8
24.6
44.7
170
169
170
110
168
71
55.0
137.0
40.6
47.4
33.4
22.6
31.0
87.0
9.0
4.7
5.8
11.7
42.5
99.5
19.7
16.7
16.9
16.1
5.0
6.0
8.3
10.4
9.0
2.6
11.9
6.0
42.2
62.4
53.6
16.6
153
153
114
134
140
102.0
151.0
50.0
83.0
15.0
40.0
77.0
17.6
3.8
0.4
54.0
88.1
31.7
17.6
5.5
7.5
13.9
6.0
8.6
1.6
13.9
15.8
19.0
48.1
29.2
Pulse crop genetic resources
335
Chickpea (Cicer arietinum L.)
Chickpea is believed to have originated in south-west Asia,
although the wild form has never been found. An escape in the wild
state in the Mediterranean region has been reported. The crop has
been in cultivation in India, the Middle East and Ethiopia for centuries (Westphal, 1974). Ethiopia and India are centres of diversity for
the cultivated chickpea (van der Maesen et ah, 1988). A wild related
species (C. cuneatum) is reported to occur in northern Ethiopia.
Chickpea is an important pulse crop, ranking second among the
pulses in Ethiopia. The annual average production reaches ca. 150000
tonnes. There are basically two types of chickpea, Desi and Kabuli.
The former is predominantly grown in Ethiopia. Ethiopia is a treasury
of variability for chickpea and is considered to be a secondary centre
of diversity.
Chickpea is sown after the main rainy season, in SeptemberOctober on residual moisture, and harvested in January-February. In
general it follows teff (Eragrostis tef) or wheat, or precedes wheat in
the case of red clay soils (Westphal, 1974). It is to a large extent a
monoculture crop, but it is sometimes found mixed with safflower,
niger seed or noog {Guizotia abyssinica), sorghum or maize, depending
on the region. Chickpea germplasm is collected from areas ranging in
altitude from 1200 to 3000 m. However, it is largely grown between
1400 and 2300 m where the annual precipitation ranges from 700 to
2000 mm. The crop is grown mainly on black clay soils of pH between
6.4 and 7.9 (Murphy, 1963).
Utilization
Evaluation data of chickpea revealed the existence of wide
genetic diversity (Table 2). The national crop improvement programme has taken advantage of the local germplasm and developed
some cultivars such as Dubie, DZ-10-11 and DZ-10-4 by direct
incorporation of the local landraces in the selection programme.
Germplasm enhancement and utilization efforts have been initiated
on a sizeable number of accessions and production of new cultivars is
already under way. Recently Debre Zeit Agricultural Research Centre
and the International Crops Research Institute for the Semi-Arid
Tropics (ICRISAT) initiated a collaborative germplasm enhancement
programme with 1000 accessions (900 accessions of Ethiopian origin)
for subsequent utilization in the breeding programme in areas where
the material proves to be useful.
336
Hailu Mekbib, Abebe Demissie & Abebe Tullu
Lentil (Lens culinaris L.)
Lentil is one of the oldest leguminous crops, believed to be
indigenous to south-western Asia and the Mediterranean regions.
From these areas the crop spread east to India, south to Ethiopia and
north to Europe (Purseglove, 1968). It is now widely cultivated in
temperate and subtropical regions as well as at higher elevations in
the tropics. Recent collecting work revealed the existence of L. ervoides
in central Ethiopia (A. Demissie, unpublished).
In acreage it is one of the major pulse crops of Ethiopia, surpassed
only by faba bean, field pea and chickpea. Lentils are cultivated under
rainfed conditions from 1500 to 3500 m. About 80 per cent of the
PGRC/E collections were assembled from areas ranging in altitude
from 2100 to 2900 m and receiving an annual rainfall of 950-1500 mm.
Generally, lentil is sown in a pure stand though sometimes it is
grown in association with linseed, which can be separated at harvest.
Lentil greatly resembles chickpea in habit and cultural requirements.
It is sown in late June to early July in poor soil and in AugustSeptember in vertisols where double cropping is sometimes practised
in certain regions.
Utilization
Lentil spread from its primary gene centre in south-west Asia
to Europe, China, India and Ethiopia (Zeven & Zhukovsky, 1975).
Important genetic variation has developed in the secondary centres of
diversity. National crop breeders have recently taken full advantage
of the genetic diversity occurring in lentil. As a result of direct exploitation of local landraces, one accession (EL-142) has been released as a
variety. Recently, efforts have been initiated to evaluate 200 accessions of lentil landraces with the ultimate objective of identifying
high-yielding lines and genetic stocks with acceptable disease/pest
resistance or tolerance.
The work on lentil is mainly on cultivar development by combining
desirable traits with useful genetic characters brought in from other
regions through international organizations. A summary of the evaluation data is presented in Table 2.
Common bean, haricot bean or kidney bean (Phaseolus
vulgaris L.)
Phaseolus vulgaris is believed to have originated in the New
World and has been cultivated throughout North, Central and South
America since ancient times. The common bean was introduced into
Pulse crop genetic resources
337
Europe by the Spaniards and Portuguese in the 16th century and was
later brought to Africa. Now it is widely cultivated in the tropics,
subtropics and temperate regions. The crop has achieved major
importance in Ethiopia.
The total holding by PGRC/E is close to 300 accessions. It is collected from a wide range of soils (from light to heavy clay and loam
soils) and climatic conditions. The altitude of collecting sites ranges
from 600 to 2230 m with a high frequency of occurrence in areas of
altitude 1700-1900 m. It can be seen from this how wide the ecological
amplitude of the crop is. Several types, differing in seed size and
colour, habit, flower colour, pod colour and size have been recorded.
The common bean collections have been evaluated for some agromorphological characters in appropriate agro-ecological sites (Table
2).
Utilization
The haricot bean is one of the important Ethiopian pulse
crops, both as a protein source for local consumption and for export
earning. The research work is based on introduced Mexican cultivars
and those developed from exotic sources. A few local cultigens (Red
Wolayita and Black Dessie) are important at subsistence farming
level.
Though an introduced species, P. vulgaris has developed wide
variation in a number of agro-morphological characters in Ethiopia.
The PGRC/E collections assembled from various agro-ecological
zones are currently incorporated in the breeding programme.
Minor pulse crops
Fenugreek (Trigonella foenum-graecum L.)
Fenugreek is indigenous to southern Europe and Asia and its
cultivation now extends from the Mediterranean to western India and
China, and south as far as Ethiopia; it is also found on the west coast
of the USA (Westphal, 1974). It has been cultivated around Saharan
oases since very early times.
To date a total of 248 accessions of fenugreek have been assembled
from various regions, particularly from Gondar and Welo where the
soils are predominantly black. The altitude of the collecting sites
ranges from 1520 to 2750 m. Seeds are sown in August and harvested
3-4 months later. Plants are uprooted and dried for a few days before
threshing and storing.
338
Hailu Mekbib, Abebe Demissie & Abebe Tullu
Utilization
Evaluation work has been initiated on the germplasm collected by PGRC/E. A total of 88 accessions have been characterized
and the summary of the data is presented in Table 2. There is no
major work on breeding and yield improvement aspects.
Grass pea (Lathyrus sativus L.)
The cultivated species belongs to the large genus Lathyrus,
with about 130 species, which are widely distributed in the Northern
Hemisphere and South America (Purseglove, 1968), with a few species in Africa. In Ethiopia five species, including L. sativus and L.
pratensis, have been identified. The latter appears to be indigenous
(Thulin, 1983).
A total of 245 samples has been assembled from fields, farm stores
and village markets. Of these, 177 were collected from defined sites.
The altitude of the collecting sites ranges from 1685 to 2700 m though
the bulk of the material was collected from an area with the altitude
ranging from 2200 to 2600 m.
Preliminary evaluation work has recently been initiated by
PGRC/E. The study indicated the presence of significant diversity for
the characters considered. The results of the evaluation work are
summarized in Table 2.
Utilization
Grass pea is endowed with many properties that make it an
attractive pulse crop in drought-stricken areas where soil quality is
poor and extreme environmental conditions prevail. Several accessions collected by PGRC/E have been put under observational trials
for screening and subsequent varietal development suitable to
moisture stress areas. Prospects for selection of superior components
within the landraces and identification of non-toxic strains are under
consideration.
Cowpea (Vigna unguiculata (L.) Walp.)
To date the origin of Vigna is not equivocally established. In
the opinion of earlier scientists, the cultivated cowpea originated in
the Indian sub-continent (Vavilov, 1951); however, more recent
studies (Faris, 1965) have gathered evidence to indicate an African
origin for this pulse crop. Steele (1976) proposes a solely Ethiopian
centre of origin followed by subsequent evolution predominantly in
the ancient farming systems of the African savannah (Duke, 1981).
A total of 47 accessions of cowpea landrace material have been
Pulse crop genetic resources
339
collected by the Ethiopian genebank for conservation and subsequent
utilization.
Utilization
The national crop improvement effort of the cowpea programme utilizes exotic material that has come through the various
international institutions, despite the existence of remarkable diversity in the indigenous material. The potential for varietal development for local germplasm is high. The passport data which include
the altitude and soil characters are indicative of the existence of a
wide ecological tolerance and this can possibly be equated with the
existence of a wide range of diversity in cowpea. This apparently
provides the basis for selecting suitable types with desirable traits for
breeding programmes.
Groundnut, peanut (Arachis hypogaea L.)
Although the cultivated groundnut is widely found in tropical and warm temperate regions throughout the world, it is native to
South America. The migration route of groundnut to Ethiopia is not
known for certain. There is little, if any, worthwhile evidence for any
pre-Columbian introduction of groundnut to Africa (Smartt, 1976).
Groundnut, though a legume crop, is categorized as a lowland oil
crop in Ethiopia, usually grown at low to mid-altitudes in the warm
regions. It is mostly intercropped with cereals and if planted alone is
assigned to marginal lands. The cultural practices and varieties used
by farmers are basically traditional types.
Although not much germplasm has been collected by PGRC/E,
efforts have been made to assemble materials from certain localities.
As a result, 14 accessions have been gathered during the course of
general collecting operations from the eastern lowland areas with
altitudes ranging from 500 to 1890 m.
Utilization
The research work on groundnut is based on introductions
and local collections. A few local collections, e.g. Bisidimo, Olole and
Sartu have shown good agronomic performances at a number of
research sites, under both irrigated and rainfed conditions.
Lupin (Lupinus albus L.)
Lupinus as a genus has two centres of distribution, in both the
Old World and the New World. The Old World centre is principally
around the Mediterranean (Gustafsson & Gadd, 1964) and North
340
Hailu Mekbib, Abebe Demissie & Abebe Tullu
Africa as far as the mountains of Kenya and the Horn of Africa. L.
albus belongs to the Old World centre and it is found in Sudan and
Ethiopia.
In Ethiopia the cultivation of lupin is limited to the northern
regions such as Gojam, Gondar and Tigray. Usually it is cultivated in
marginal soils where other pulse crops do not perform very well. It is
generally sown during the main rainy season (July-September) and
harvested in December. Minimum tillage and cultural practices are
followed for the cultivation of the crop.
The economic importance and agricultural potential of lupin is not
widely recognized in the country except in the northern regions
where the seeds are used to make 'katikala', a local drink. There are
very few accessions conserved at PGRC/E.
Lesser known but potentially valuable pulse crops
The genetic resource base of pulse crops in Ethiopia is not
limited to the species indicated above. There are several species
which are lesser known but potentially valuable with significant
regional importance; these are listed in Table 3.
General considerations
It is evident from this account that there is wide ecological
amplitude for most of the pulse species in Ethiopia. Moreover, from
this and from previous evaluation data (Mekbib, 1988) it can be seen
that there is a high diversity for the characters recorded. The widely
grown species have types well adapted to specific habitats. The
recently introduced pulses such as Phaseolus vulgaris have become
important in both acreage and production, while P. lunatus and P.
coccineus are limited in distribution and acceptance.
Utilization of the available genetic diversity in the country is
relatively low compared with cereals and oil crops. Work on the
exploitation of the variability in pulse crops has only recently been
initiated. The genetic exploitation of pulses differs from species to
species and the effort initiated on utilization of cool season legumes
appears to be encouraging. The joint utilization effort between
national institutions and international organizations is a step in the
right direction in maximizing the benefit of landraces and primitive
cultivars available in the country.
Several of these pulse species possess specific attributes of utility in
crop breeding programmes. These cultigens have desirable traits that
can be crossed with high-yielding cultivars. Preliminary observations
Table 3. List of pulse crops occasionally cultivated and rarely encountered in Ethiopia
Scientific name
Common name
Genetic resources and utilization
Voandzeia subterranea (L.) DC
Bambara groundnut
Phaseolus coccineus L.
Scarlet runner bean
P. lunatus L.
Lima bean
Jack bean
Sword bean
Locally grown in south-western Ethiopia, few accessions are
collected and conserved
Rarely encountered as garden plant, few accessions are
collected so far
Fairly recent introduction, found in south-western regions
Recorded as found in south-west and part of Northern
Ethiopia
Probably sometimes cultivated in some regions, reported in
Westphal (1974)
Cultivated in Welega (Thulin, 1983)
A rare occurrence, almost unknown (Westphal, 1974)
Fairly recently introduced but becoming quite common, few
accessions assembled
Highly threatened species, endemic to Eastern Ethiopia and
part of Somalia
Recent introduction and sometimes cultivated for human
consumption, mostly as a garden plant
Recent introduction and restricted cultivation
Recent introduction and grown in limited areas
Recent introduction and grown in limited areas
Canavalia ensiformis (L.) DC
C. virosa (Roxb.) Wight & Arnott
Mucuna pruriens (L.) DC
Psophocarpus palustris Desv.
Vigna radiata (L.) Wilczek Syn.
Cordeauxia edulis Hemsl.
Lupinus mutabilis Sweet
Cajanus cajan (L.) Millsp.
Glycine max (L.) Merr.
Lablab purpureus (L.) Sweet
Velvet bean
Goa bean
Green gram
Mung bean
Yeheb nut
Tarwi
Pigeon pea
Soya bean
Hyacinth bean
342
Hailu Mekbib, Abebe Demissie & Abebe Tullu
have shown that some accessions are tolerant to stress conditions and
diseases and are sometimes preferred by local farmers. Being particularly adapted to diverse agro-ecological conditions and extreme climatic stress, they provide useful germplasm for introducing tolerance
to drought, cold and diseases.
In general, with the exception of recently introduced species, the
major improvement work, particularly of the cool season legumes
such as faba bean, field pea, lentil and chickpea, is largely based on
indigenous germplasm. There are still untapped resources of genetic
material available for breeding work. This considerable genetic
wealth has to be fully exploited in national efforts to ameliorate the
farming situation.
It will be essential to establish more comprehensive collections of
the different pulses, in order to represent the existing diversity in
Ethiopia in these collections. So far pulse crops have only enjoyed
lesser priority in terms of collecting. Field exploration/collection,
survey and basic diversity studies are essential to identify high diversity areas for subsequent collecting operations and conservation.
Evaluation is exceedingly important and constitutes a prerequisite
for the utilization of landraces. In-depth studies of the material to
hand, with respect to characters of adaptation and resistance to stress
conditions such as frost, cold, heat, drought and to adverse soil conditions, are required and should be strengthened. Landrace improvement endeavours on pulses, such as the exemplary undertaking on
durum wheat, should be looked at as one of the facets of improvement programmes for pulse crops.
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their Centres of Diversity. PUDOC, Wageningen.
Zohary, D. (1970). Centres of diversity and centres of origin. In: O. H.
Frankel and E. Bennett (eds), Genetic Resources in Plants: Their Exploration
and Conservation. Blackwell, Oxford, pp. 33-42.
28
Oil crop germplasm: a vital resource
for the plant breeder
HIRUY BELAYNEH
Introduction
According to Seegeler (1983), 328 oil plant species are known
to exist in Ethiopia. Of these, 15 are cultivated and the rest may have
uses other than for oil and may be cultivated or wild. Oil-bearing
plants having oil contents in excess of 10 per cent, but which are not
yet cultivated commercially, have been catalogued by Goshe &
Hamito (1983) (Table 1).
Ethiopia is known to be either a centre of origin or a centre of
diversity for many cultivated oil crops. Several of the cultivated oilseed crops play an important role in the nutrition of the Ethiopian
population and in foreign exchange earnings. The oil crops currently
in production in the country are niger or noog, rapeseed, Ethiopian
mustard or gomenzer, linseed, sunflower, sesame, groundnut, safflower and castor bean.
The overall objective of the research programme is to increase the
production of oil seeds for food and to provide raw materials for agroindustrial development. This can be achieved by the development of
high-yielding, stable cultivars with the necessary package of practices
required for sustained high yields. The programme therefore falls
into three sections:
- the improvement of noog, linseed, sesame and safflower
which possess a wide range of variability and a wealth of
unutilized indigenous germplasm;
- the improvement and popularization of oil seed Brassica and
groundnut for which a wide range of indigenous germplasm
is also available;
- the popularization of the introduced sunflower crop, for
which probably no indigenous germplasm exists.
Oil crop germ-plasm
345
Table 1. The oil or fat content on dry seed basis of some selected Ethiopian
tree and shrub species
Scientific name
Common or local
name
Oil content of
seeds/kernels
(%)
Gossypium spp.
Argemone mexicana
Ricinus communis
Jacaranda spp.
Schinus molle
Ximenia americana
Schefflera abyssinica
Maesa lanceolata
Trichilia roka
Bersama abyssinica
Allophylus abyssinicus
Sterculia africana
Pittosporum mani
Sapium ellipticum
Terminalia macroptera
Balanites aegyptica
Salvadora persica
Melia azedarach
Myrica salcifolia
Erithyrina abyssinica
Trema guineensis
Acanthus spp.
Cocos nucifera
Cotton
Prickly poppy
Castor bean
Jacaranda
Pepper tree
Inkoy (Amharic)
Ketema (Amharic)
Kelewa (Amharic)
Ethiopian mahogany
Azamir (Amharic)
Imbus (Amharic)
Fua (Gambella)
—
Gancho (Sidamo)
Kokora (Oromo)
Desert date
Hadia (Tigray)
Neem tree
Shinet (Amharic)
Red-hot-poker tree
Sendo (Amharic)
Kosheshila (Amharic)
Coconut
10-24
38.7
51.7
36.6
10.9
49°
10.3
22.3
44-56
35.8
26.4
30.8
23.4
55*
52-64"
10-50°
19-34
33-45*
20
15-20
28
23
65-68
a
Represents oil content of kernels.
Source: Goshe & Hamito (1983).
Germplasm collection
Ethiopia is known to be a gene centre for many cultivated
crops. Much has been done in the past to exploit this germplasm
wealth. The collection of oil seed crops has been carried out with the
objective of enriching the existing gene pools of those crops utilized
in breeding and selection programmes.
The collection and characterization of indigenous germplasm were
initiated in the early 1940s, but lost momentum before being resumed
in the early 1970s. A few Italian documents exist showing the work
done on noog landraces between 1940 and 1946. The period 1974-85
saw an intensification of effort in the search for accessions with the
required characteristics, with the intention of exploiting promising
local cultivars to the fullest extent. The first step in this approach was
346
Hiruy Belayneh
Table 2. Oil crop germplasm collections maintained at PGRC/E
Oil crops
Total number
of accessions
Oil seed Brassica (Brassica carinata, B.
nigra, B. oleracea and B. napus)
964
Cultivated niger or noog (Guizotia
abyssinica)
939
Wild noog (G. scabra)
7
Sunflower (Helianthus annuus)
20
Linseed (Linum usitatissimum)
1832
132
Safflower (Carthamus tinctorius)
Castor bean (Ricinus communis)
350
Sesame (Sesamum indicum)
376
Groundnut (Arachis hypogaea)
19
Number of
accessions
characterized
910
826
—
—
470
99
49
195
16
to establish a collection of the local germplasm before it was irretrievably lost as a result of changes in land use, drought, etc.
Initially, the collection of oil crop germplasm was undertaken by
extension agents and oil crop subcommittee members, most of the
collections being samples from farmers' stores or local markets. In
recent years, collections have been made primarily from standing
crops in farmers' fields. However, many of the expeditions were
confined to the roadsides. The oil crop team members and invited
experts have participated in several expeditions which substantially
increased the available germplasm, particularly of Ethiopian mustard,
linseed, noog and sesame. The collecting trips were made in collaboration with the Plant Genetic Resources Centre/Ethiopia (PGRC/E).
The germplasm holdings collected and preserved at PGRC/E so far
are presented in Table 2.
Traditional utilization
In Ethiopia, oil seeds are the main source of oil in the diet of
the majority of the population. Noog is the prime supplier of cooking
oil and usually commands a premium over other available oils.
Sesame and safflower oils are appreciated in regions where the crops
can be produced. Linseed oil is used when other oils are in short
supply. Mustard seed oil is also produced commercially. The cake
remaining after oil extraction is rich in protein and can provide a
valuable livestock feed.
In the local cuisine, dried and/or ground seeds from linseed, noog
Oil crop germplasm
347
and mustard are used to prepare a stew or 'wot'. Mustard leaves are
an important leaf vegetable in the highlands where they are boiled
and served as 'gomen wot'. Roasted and crushed seeds of noog,
linseed and safflower may be mixed with water to prepare beverages.
Noog and sesame flours are sometimes mixed with flour for baking
bread and the whole seeds are often sprinkled on top prior to baking.
Seed oil from castor, linseed and mustard plants can also be used
for tanning leather and for varnishes. The use of mustard, safflower
and castor oils for lighting is widespread and they are also commonly
used to grease the 'metad' (pan) before 'injera' (teff bread) is baked.
Preparations from noog, mustard, sesame, linseed and castor
plants are traditionally used for one or more of the following medical
problems:
- to treat diarrhoea (sesame, linseed, niger);
- as a diuretic (linseed);
- to treat eye irritation due to dust (linseed, castor, sesame);
- as a purgative (linseed, safflower);
- for birth control (mustard, niger);
- to treat sores and rheumatism (safflower, sesame);
- to treat syphilis (niger);
- to treat coughs (niger);
- to treat an upset stomach (mustard).
Oil crops also have many other uses in Ethiopia, e.g. as condiments, as snack food (when roasted), as a lubricant, as an additive for
soap and paints and as a fertilizer.
Variety development
The task of the oil crop breeders is concentrated on the
development of high-yielding varieties, taking into consideration
resistance to major diseases and improved quality. Depending on the
adaptability of the species, the indigenous germplasm was initially
characterized for agronomic and morphological characters at the
Holetta, Melka Werer, Awasa and Debre Zeit Research Centres of the
Institute of Agricultural Research (IAR).
Further evaluation tests were carried out at several sites. Large oil
seed introductions from different countries have also been evaluated
at a number of sites. The multilocation trials have resulted in the
release of several improved varieties of oil crops. The names of the
released and/or recommended varieties, their desirable characteristics
and other information are presented below on a crop basis.
348
Hiruy Belayneh
Table 3. Number of indigenous entries in different variety trials, 1986
Crop
Micro-trial
Pre-national
variety trial
National
variety trial
Niger
Ethiopian mustard
Linseed
Sesame
Groundnut
Safflower
19
10
21
14
—
2
7
8
9
5
2
2
7
2
3
4
3
_
Extension
yield trial
Niger or noog (Guizotia abyssinica)
There is evidence that noog orginated in the highlands of
Ethiopia, north of 10° N latitude (Baagoe, 1974). Harlan (1975) considers noog to be among the earliest of the crops domesticated in
Ethiopia. It may have orginated from the wild species Guizotia scabra
through selection by Ethiopian farmers over thousands of years. The
crop has always been one of the most important oil crops in Ethiopia
in terms of both area and production. All cultivars are local and raised
under rainfed conditions.
The objective of the noog breeding programme is to develop
improved cultivars which produce consistently and give a high yield
of good quality. Special attention is being given to lodging resistance,
uniform ripening, minimal shattering, frost tolerance and resistance
to Septoria disease.
As the research effort in noog has not been as extensive as in other
oil seeds, only one variety (Sendafa) has been released to growers.
However, recent accessions have been evaluated for desirable characters and promising, indigenous lines have been advanced to the different levels of replicated variety trials (Table 3).
Two selection programmes on noog landraces are in progress now.
The mass selection programme aims at improved plant type through
usual selection while the half-sib recurrent selection programme uses
an evaluation of progeny performance as a basis of selection. Each
method is geared towards developing early and medium maturing
composites with the desirable characteristics.
Oil seed Brassica (Brassica carinata, B. nigra, B. napus and B.
oleracea)
Ethiopian mustard or gomenzer {Brassica carinata) is believed
to have originated in Ethiopia from the natural crossing of Brassica
Oil crop germplasm
349
Table 4. Oil crop varieties that have been released from introduced
germplasm
Crop
Variety
Year of release
Country of origin
Rape seed
Target
Tower
Pura
Tower Sel3
Victory
Concurrent
CI-1525
CI-1652
Russian Black
Hesa
Pop 158
Shulamith
NC4X
NC343
T-85
S
E
1976
1984
1984
1986
1978
1978
1984
1984
1974
1974
1974
1976
1986
1986
1976
Canada
Canada
W. Germany
Canada/Ethiopia
North Dakota, USA
The Netherlands
France
Ireland
Yugoslavia/Russia
W. Germany
W. Germany
Israel
USA
USA
Uganda/India
Uganda
Uganda
Linseed
Sunflower
Groundnut
Sesame
nigra (senafitch) and Brassica oleracea (Gurage or Wollamo gomen).
Ethiopian mustard is the fourth most important oil crop after niger,
linseed and sesame and is widely grown only in Ethiopia. It is also
used extensively as a vegetable.
The objective of the breeding programme is to develop improved
cultivars of the oil seed Brassica, especially rape seed and mustard.
Resistance to diseases, especially to Alternaria leaf spot, low erucic
acid in the oil and low glucosinolate and fibre concentrations in the
meal are receiving special attention.
Evaluation of Ethiopian landraces has led to the recommendation
of five Brassica carinata varieties (S-67, S-71, S-115, Awasa population
and Dodolla-1) for the highlands of Ethiopia. Four napus varieties,
namely Target, Tower, Pura and Tower Sel3, were identified from
introductions and released for large-scale production (Table 4). The
latter three varieties have low erucic acid and low glucosinolate and
have replaced Target, which has high erucic acid and high glucosinolate similar to the recommended high-yielding Ethiopian mustard
varieties.
The major effort in the crossing programme has been towards
incorporating earliness, low erucic acid and low glucosinolate characters into local mustard selections. Interspecific crosses have been
350
Hiruy Belayneh
achieved and are being used to broaden the genetic basis of Ethiopian
mustard.
The existing germplasm lines and selections of Ethiopian mustard
are being screened for low erucic acid and glucosinolate and evaluated for oil yield (Getinet, Rakow & Downey, 1986). Low erucic acid
oil and low glucosinolate meal are now essential if an international
trade is to be developed and the meal is to be fed to animals.
Linseed (Linutn usitatissimum)
Linseed is also an important oil crop occupying a wide production area. The crop has built up considerable diversity in Ethiopia
after its early introduction from Asia. The objective of the breeding
programme is to develop improved cultivars of linseed, and disease
resistance to wilt, powdery mildew and Septoria are important.
A large number of local collections have been evaluated and
characterized (Table 2). Single plant selections were made in the nurseries to capture 'within' plot variation and uniform lines were produced. A number of varieties and lines are being advanced through a
four-stage hierarchy of yield trials (Table 3).
Varietal testing of introduced linseed germplasm has resulted in
the release of four varieties, Victory, Concurrent, CI-1525 and CI1652. The latter two bold-seeded varieties were released in 1984 after
fulfilling the pre-release requirements. A hybridization programme
based on the pedigree method and using early, wilt resistant and
high-yielding lines is in progress. The existing local linseed collections, as well as introductions, will be screened for low linolenic acid
content. Reduction in linolenic acid content in linseed should overcome problems like rapid flavour deterioration, low nutritive value,
etc.
Sunflower (Helianthus annuus)
The purpose of the sunflower breeding programme is to
identify well adapted cultivars of this crop, also giving consistent,
enhanced yields of oil per hectare. Resistance to disease, especially to
downy mildew, as well as to Sclerotinia and rust require more
attention.
More than 300 varieties of sunflower have been introduced, mainly
from North America, the USSR and Eastern Europe, and evaluated
while in quarantine. A number of promising lines were advanced
through a four-stage hierarchy of yield trials. Before 1976, three
varieties of sunflower, Russian Black, Hesa and Pop 158, were recom-
Oil crop germplasm
351
mended for general release. Of the varieties tested in the last five
years, the long-maturing Argentario was identified as being widely
adaptable. The two promising early maturing lines in the programme
are Charnianka x Gene pool I and Gene pool II.
Sesame (Sesamum indicum)
Ethiopia is considered as either the centre of origin or a centre
of diversity for sesame (Seegeler, 1983), hence the collection and
characterization of local germplasm is very important in the overall
improvement programme for the crop.
The purpose of the breeding programme is to identify the most
adaptable sesame varieties for the various agro-ecological zones
through selection and hybridization. Development of varieties with
high oil content, resistance to bacterial blight disease, uniform ripening and minimum shattering characteristics, receives special
attention.
Of the landraces assembled and evaluated, some have proved to be
useful and were advanced through a four-stage hierarchy of yield
trials (Table 3). Selection of progenies was also carried out and some
have shown promising performances under multilocational tests.
From the landraces collected earlier, 'Kelafo 74', a variety collected
from the lower Wabe Shebelle Basin of Kelafo district, has proved to
be among the best yielders, especially under irrigated conditions at
Melka Werer, Tendaho and Gode. One progeny from a collection of
the western region, SPS-111519, is under consideration for release in
the rainfed zones.
Several exotic sesame germplasm accessions have been tested and
three varieties, T-85, S and E (introduced from Uganda), were recommended for general release and are under production at several sites.
A hybridization programme to evolve lines with partial shattering
characteristics, disease resistance and high seed and oil yield, is in
progress at Melka Werer Research Centre. Preliminary results show
that some lines with uniform stand, good capsule setting, partially
opening capsules at harvest and bacterial blight resistant types are
attainable.
Groundnut (Arachis hypogaea)
In Ethiopia, groundnuts are consumed roasted or are crushed
for the production of edible oil. The overall target of the breeding
work is the increase of the productivity of the crop for local consumption, agro-industrial projects and export. Furthermore, the identifica-
352
Hiruy Belayneh
tion of the most suitable groundnut varieties for the different agroecological zones is an important objective. Special attention is being
given to disease resistance, in particular to leaf spot and rust.
From earlier local accessions, 'Asmara' and 'Dire Dawa', which
were collected from the respective regions, were recommended for
general release. Of the few recent local accessions, Bisidimo, Olole
and Sartu have shown good performance at Melka Werer, Bisidimo
and Babile.
Evaluation of earlier introductions has led to the release of
Shulamith. This variety is under cultivation over a wide region. Of
the recent introductions from the USA, two Virginia type varieties,
namely NC 4X and NC 343, were released in 1986 for irrigated areas of
the Awash Valley. The two varieties had shown a maximum yield of
7000 kg/ha at the irrigated site of the Middle Awash Valley and
2500 kg/ha at the marginal rainfall sites of Bisidimo and Babile.
Moreover, moderately disease-resistant and early maturing varieties
have been obtained from the recent introductions from the International Crops Research Institute for the Semi-Arid Tropics (ICRISAT).
Safflower (Carthamus tinctorius)
Ethiopia is considered to be the centre of origin and/or of
diversity for safflower. It is grown as a companion crop with cereals,
mainly teff. Research work on the crop was discontinued for some
time and resumed only a few years ago. The objective of the breeding
programme is to develop improved cultivars of safflower for low,
mid- and high-altitude areas. Disease resistance to Ramularia and
Alternaria caratami is important.
About 100 new landraces have been collected and characterized for
morphological and agronomic performance and some outstanding
lines have reached the micro-trial and pre-national variety trial stages
(Table 3). A few introductions of safflower lines have been received
from Indore, India.
Some 600 progeny selections were made from the earlier collections to get pure and uniform lines. Outstanding progenies have been
advanced to the pre-national variety trial.
Castor bean (Ricinus communis)
The castor bean research programme started two years ago,
after a long period of interruption. The purpose is to evaluate indigenous and exotic varieties mainly for adaptability and high seed and
oil yield.
Oil crop germplasm
353
So far, about 200 local collections and 90 introductions have been
evaluated and characterized. Some outstanding lines were observed
in the preliminary observation nursery and have been advanced for
further studies.
References
Baagoe, J. (1974). The genus Guizotia (Compositae). A taxonomic revision.
Botanisk Tidsskriff, 69, 1-39.
Getinet, A., Rakow, G. & Downey, R. K. (1986). Seed colour and quality
characteristics in Ethiopian mustard (Brassica carinata). PGRC/E-ILCA Germplasm Newsletter, 12, 12-15.
Goshe, B. A. & Hamito, D. S. (1983). Preliminary survey of oil bearing plants
in some regions of Ethiopia. Ethiopian Journal of Agricultural Science, 5,
89-96.
Harlan, J. R. (1975). Crops and Man. American Society of Agronomy,
Madison, Wisconsin.
Seegeler, C. J. P. (1983). Oil Plants in Ethiopia, their Taxonomy and Agricultural
Significance. PUDOC, Wageningen.
29
Significance of Ethiopian coffee
genetic resources to coffee
improvement
MESFIN AMEHA
Introduction
Among the economic species of coffee grown in more than 50
countries in different parts of the world, Coffea arabica L. is the only
tetraploid species of the genus. It contributes over 80 per cent of the
world's coffee production. In many scientific reports Ethiopia is considered to be the centre of origin and diversification of coffee (Sylvain,
1958; Fernie, 1966; Food and Agriculture Organization, 1968; Carvalho et al., 1969; Ameha, 1980; Rodrigues, 1981; Worede, 1982; Watkins, 1985). The question of whether the south-west mountain moist
evergreen forest, the farmer's field or the low altitude river banks is
the primary habitat is not discussed here, although the issue is of
primary interest to geneticists and breeders, for conservation
purposes and in the search for primitive genes in the wild
progenitors.
Arabica coffee is an evergreen shrub of variable size. The tree
grows up to 14 m in height and about 2 m in width under forest strata
and up to 6 m in height and about 12 m in width in canopy under
farmers' holdings (a tree this size was observed in Wanago near Dilla
in Sidamo administrative region). Naturally, it has a dominant central
orthotropic stem and horizontally growing plagiotropic branches
with pairs of secondary, tertiary, etc. branches originating from preceding branches in the hierarchy. The leaves are borne in opposite
pairs along the side of the branches. The flowers emerge as
inflorescences on all forms of lateral branches in each leaf axil of the
nodes. Every flower normally develops into a two-seeded berry. The
Significance of Ethiopian coffee genetic resources
central stem gives rise to a number of orthotropic stems when
stumped, wounded or bent.
Climate and soil
Coffee is a hardy plant which thrives well in almost all types
of soils under a wide range of climatic conditions. In Ethiopia, it
grows in almost all administrative regions in conditions ranging from
the semi-savannah climate of the Gambela plain (550 m above sea
level) to the continuously wet forest zone of the south-west (2200 m).
It generally grows on sloping land of different gradients.
The soil varies from sandy loam to heavy clay while the general soil
types, which are acidic (pH 4.2-6.8), are red to reddish-brown lateritic
loams or clay loams of volcanic origin (Sylvain, 1958; Fernie, 1966).
Annual rainfall varies from 750 to 2400 mm. The upper limit for day
maximum temperature recorded is 36 °Cand the lower limit for night
minimum is 7°C, with varying ranges of diurnal temperature.
Botanical cultivars
Ethiopian coffee production generally comes from botanically
unidentified cultivars of Arabica populations. In the early years the
local names of this coffee reflected either the area of production or the
names of influential landlords or aristocrats and, even today, these
local names are widely used.
Chevalier (1947) and Ciferri (1940) were the pioneers in grouping
Ethiopian coffee by bean characteristics. Sylvain (1958) grouped them
into Ennarea, Jimma, Agaro, Chochie, Yirgalem, Dilla, Arba Gugu,
Harer, Loulo, Wolkitte and Welayita types using bean shape and
colour, leaf size and colour, leaf tip colour and other obvious characters. Recently, a number of distinct morphotypes have been
recognized. In the Harer type, for example, Abadiro, Buna Guracha,
Buna Kella and Shimbre, among others, are quite different (Watkins,
1985). From the accessions conserved in the National Coffee Collection and in the collection of coffee berry disease (CBD) resistant types,
different coffee strains can be distinguished both qualitatively and
quantitatively. The Mettu types (74110, 7412, 74148, etc) possess very
narrow shiny green leaves, somewhat compact in nature. Accessions
like 2370, 3170, 7440, and 741 are robust and of a spreading type
(Institute of Agricultural Research, 1971-84). This considerable variation requires a systematic approach by a taxonomist and geneticist in
order to develop a practical classification.
355
356
Mesfin Amelia
Variation in coffee
In Ethiopia, research findings over 18 years have revealed the
presence of enormous genetic variation for different agronomic traits
(Brownbridge & Gebre-Egziabher, 1968; IAR, 1971-84; Ameha, 1980,
1983; van der Graaff, 1981; Worede, 1982; Ameha & Belachew,
1986a,b; Ameha, Belachew & Shimbr, 1986; Belachew & Ameha,
1986). Yield was almost normally distributed with mean yield per
accession varying from 10 to 840, 55 to 895, 165 to 720, 150 to 775 and
365 to 1250 g per tree clean bean at Melko, Mettu, Agaro, Wonago and
Gera, respectively. Similarly, when the yield variation of individual
accessions over a number of years of production was analysed, the
coefficients of variation ranged from 34 per cent to 211 per cent.
Frequency distributions for resistance to CBD, leaf rust and liquoring quality were skewed but all in favourable directions (IAR, 197184; Ameha, 1980, 1983; Worede, 1982; Ameha & Belachew, 1984;
Belachew & Ameha, 1986). The negative bimodal skewedness
observed for bean grade is a typical character of Arabica coffee.
Generally, there are more discard and remainder beans in the processed coffee as a result of Man's preference for well shaped, large beans
of AA, A and B grades. When bean sizes were measured, most
frequent dimensions varied from 4.8 to 14.5, 4.0 to 10.5 and 2.5 to
8.5 mm for length, width and depth, respectively. The bean colour is
classified into green olive-green, green yellow, green bluish, yellow
amber and pale yellow and the bean shape as small oval, almost
round, large rectangular and large oval (Ciferri, 1940; Sylvain, 1958;
FAO/IBP, 1973; Belachew & Ameha, 1986). The degree of variation for
other characters, such as tree shape, branching habit, resistance to
diseases and pests, persistent sepals, etc. is also striking. As expected, it appeared that most of the traits are quantitatively heritable
and, in general, considerably influenced by environment (Tadesse &
Engels, 1986).
Several researchers in the Western Hemisphere have attributed the
limited variation there to mutation and restricted genetic recombination. The coffee varieties in Latin America evolved from a few seeds
which were derived from only a few trees grown in the Botanical
Garden of Amsterdam in the middle of the 18th century. However,
here in Ethiopia, nature has played a vital role in the selection, distribution and adaptation of Arabica coffee and it was only 20 years
ago that Man entered the picture with his ideas of systematic selection. What is being grown now is the result of tens of thousands of
years of natural selection, selection that has occurred in situ. It is from
Significance of Ethiopian coffee genetic resources
this background that the hybridization and heritability study for CBD
revealed new findings in the predominantly self-fertile coffee species
(Ameha & Belachew, 1984; 1986a). Three to five recessive genes controlling resistance to CBD and a hybrid vigour for yield and yield
components were observed in the indigenous coffee. These recent
findings also indicate that Arabica genotypes are generally locationspecific. No one cultivar performed consistently well across locations
for yield and vigour, suggesting how specific Arabica 'ecotypes' are in
their adaptability.
Loss of genetic diversity
In the last 40 years, a significant reduction of genetic diversity
has occurred in the Ethiopian coffee. According to FAO reports
almost 90 per cent of the Ethiopian forest cover had vanished by 1965
(FAO, 1968, 1973). Added to this deforestation are the effects, in
subsequent years, of the development of new roads bringing
expanded agriculture and forest utilization, particularly in the rainy
forests of the south-west where coffee occurs naturally in association
with forest. Consequently, coffee genetic erosion has gone far beyond
the point of no return. Many coffee forests are no longer intact.
Persistent drought in the last 15 years has further aggravated the
situation. Hundreds of hectares of coffee forest have been replaced by
food crops since farmers want more food for their families. Furthermore, food crops pay farmers more than coffee does when production per unit area is compared in times of drought. Even under
normal conditions, when food is abundant, the income from coffee is
discouraging. This, together with the drought, has led to the destruction of the coffee forests.
With the advent of CBD in 1971 and the subsequent identification
of CBD-resistant selections, the distribution of resistant cultivars
resulted in the retention of relatively invariable individuals in some
typical coffee forests where they were replanted after forest clearing.
This has definitely caused significant losses in genetic diversity. It has
been estimated that between 25 000 and 35 000 hectares of semi-forest
coffee have so far been replaced by CBD-resistant cultivars, leading to
at least 10 per cent loss.
Observations suggest that if the present pace of agricultural
development, forest utilization and population growth continue, by
the year 2000 about 120000 hectares of the estimated 350000 hectares
of semi-forest coffee will be replanted with the advanced cultivars
and about 80 per cent of the remaining 230 000 hectares will be lost
357
358
Mesfin Ameha
through other factors. The loss of the heterogeneous coffee populations, which represent the gene pool for hundreds of agronomic
traits, will be catastrophic. The well known quality coffee of Limu,
Nekemte, Gimbi, Harer and Yirga Chefe, which fetches a high
premium, will no longer exist unless immediate ways are found to
preserve it. The conservation work of PGRC/E which has started at
Chochie, Kefa, in cooperation with the Jima Research Centre of the
Institute of Agricultural Research, is highly appreciated. It needs to
be strengthened with better facilities and more manpower thus allowing more accessions to be conserved. In the same manner, and as a
matter of considerable urgency, collection and in situ conservation of
Harer coffee must resume. Watkins (1986) and the IAR team concluded that at present 80 per cent of the coffee from Habro Awraja,
which produces 40 per cent of the Harer coffee, is rapidly declining in
production. Prompt action is required to save the germplasm of this
coffee which has a tremendous, worldwide reputation for quality and
fetches twice the price of coffee from other regions.
A project with comprehensive conservation strategies including (a)
ecosystem conservation for the wild and semi-wild coffee and (b)
collection and in situ conservation, was prepared and submitted for
implementation some years back. PGRC/E, together with other
governmental institutions and organizations affiliated with coffee,
must be encouraged to facilitate and implement those strategies as a
matter of top priority, by law if necessary.
References
Ameha, M. (1980). Yield assessment of indigenous coffee collection grown at
Jima Research Centre. Ethiopian Journal of Agricultural Science, 11, 69-77.
Ameha, M. (1983). Variabilities of indigenous coffee collection to rust
resistance. Simposio Sobre Ferrugens do Caffeeiro, Oeiras, Portugal.
Ameha, M. & Belachew, B. (1984). Resistance of the F! to coffee berry disease
in six parent diallel crosses in coffee. Proceedings, First Regional Workshop on
Coffee Berry Disease. Association for the Advancement of Agricultural
Sciences in Africa, Addis Ababa, pp. 167-77.
Ameha, M. & Belachew, B. (1986a). Field evaluation of resistance to stress
conditions in crosses and their parents of coffee, Coffea arabica L. First
Ethiopian Coffee Symposium, Institute of Agricultural Research, Addis Ababa,
August 1986. IAR, Addis Ababa.
Ameha, M. & Belachew, B. (1986b). Genotype-environmental interactions in
coffee, Coffea arabica L. First Ethiopian Coffee Symposium, Institute of Agricultural Research, Addis Ababa, August 1986. IAR, Addis Ababa.
Ameha, M., Belachew, B. & Shimbr, T. (1986). Yield assessment of CBD
(coffee berry disease) resistant progenies of coffee under different environments. First Ethiopian Coffee Symposium, Institute of Agricultural Research,
Addis Ababa, August 1986. IAR, Addis Ababa.
Significance of Ethiopian coffee genetic resources
359
Belachew, B. & Ameha, M. (1986). Variation among national coffee collections for some agronomic characteristics. First Ethiopian Coffee Symposium,
Institute of Agricultural Research, Addis Ababa, August 1986. IAR, Addis
Ababa.
Brownbridge, J. M. & Gebre-Egziabher, E. (1968). The quality of some of the
main Ethiopian mild coffees. Turrialba, 18, 361-72.
Carvalho, A., Ferwerda, F.P., Frahm-Leliveld, J.A., Medina, D.M.,
Mendes, A.J.I. & Monaco, L.C. (1969). In: F. P. Ferwerda and F. Wit
(eds), Outlines of perennial crop breeding in the tropics. Miscellaneous Papers
4, Landbouwhogeschool, Wageningen, pp. 189-241.
Chevalier, A. (1947). III. Systematique des cafeiers et faux-cafe"iers. Maladies
et insectes nuisibles. Cyclopedia diologique, No. 28, Paris.
Ciferri, R. (1940). Primo reporto sul caffe nell - Africa Orientale Italiana.
Firenze, Regio Instituto Agronomico per 1'Africa Italiana. Relazioni e
monografie agrario coloniali, No. 60.
Food and Agricultural Organization (1968). Coffee Mission to Ethiopia 1964-65.
FAO, Rome.
FAO/IBP (1973). Survey of crop genetic resources in their centres of diversity. First
report. FAO, Rome.
Fernie, L. M. (1966). Impressions of Coffee in Ethiopia. Coffee Research Station,
Lyamungu, Tanzania.
Institute of Agricultural Research (1971-84). Coffee Progress Report (annual).
Jima Research Centre, IAR, Addis Ababa.
Rodrigues, C.J. Jr (1981). Coffee Leaf Rust in Ethiopia (consultant report, Eth.
78/004). FAO, Rome.
Sylvain, P. G. (1958). Ethiopian coffee - its significance to world coffee problems. Economic Botany, 12, 111-39.
Tadesse, D. & Engels, J. M. M. (1986). Phenotypic variation in some fruit
characters in coffee collected from Chora wereda. PGRC/E-ILCA Germplasm
Newsletter, 12, 2-8.
van der Graaff, N. A. (1981). Selection of Arabica coffee types resistant to
coffee berry disease in Ethiopia. Mededelingen Landbouwhogeschool
Wageningen, 11-1. University of Wageningen.
Watkins, R. (1985). Coffee Coffea arabica L. genetic resources and breeding.
Consultant Report, Wye College, University of London.
Watkins, R. (1986). Proposed Actions for Maintenance and Production of Harer
Coffee. Coffee Improvement Project, Addis Ababa.
Worede, M. (1982). Coffee genetic resources in Ethiopia: conservation and
utilization with particular reference to CBD resistance. Proceedings, First
Regional Workshop on Coffee Berry Disease. Association for the Advancement
of Agricultural Sciences in Africa, Addis Ababa, pp. 203-11.
30
Use of Ethiopian germplasm in
national and international
programmes
J. G. HAWKES AND MELAKU WOREDE
Introduction
It will have become evident from the previous chapters in this
book that the crop genetic resources of Ethiopia are very diverse and
constitute an invaluable base for plant breeding both within and
outside the country. Ethiopia is one of the world centres of diversity,
identified by N. I. Vavilov some 60 years ago. Not only does it possess
important diversity in crops domesticated elsewhere, such as wheat,
barley, grain legumes and several oil plants; it also has developed its
own indigenous cultigens, such as teff, sorghum, niger seed (noog),
ensete, Ethiopian mustard and coffee, many of which are now of
great international importance. Ethiopian breeders have taken full
advantage of the crop genetic diversity in their own country, combining it with useful genetic characters brought in from other regions.
Clearly, the importance of Ethiopian crop diversity has not gone
unnoticed amongst world breeders. Vavilov, who visited Ethiopia in
1927, pointed out the value, particularly to wheat and barley
breeders, of the Ethiopian landraces and their extraordinary morphoagronomic variation (Vavilov, 1931).
In this final chapter we shall attempt to summarize the value of
Ethiopian crop genetic diversity both nationally and internationally.
Wheat
Ethiopia is unique in containing a very wide diversity of
tetraploid wheat, but very little hexaploid wheat diversity; this latter
was probably introduced in recent times.
The use of Ethiopian germplasm
361
Improvement of indigenous landraces is described by Tesfaye
Tesemma (Chapter 22) in terms of higher productivity, and stem and
leaf rust resistance; promising lines are often crossed with exotic
varieties in order to improve yield and quality. Prospects for the
selection of superior components within landraces are also under
consideration.
Breeding for resistance to Septoria tritici is being dealt with by Hailu
Gebre-Mariam (Chapter 23), using groups of bread wheat varieties
from Ethiopia and other sources. Useful resistance was found in these
durum varieties checked for leaf rust under field conditions. In
general, wheat breeders in Ethiopia recognize the need to involve
Ethiopian varieties and landraces in their programmes. Most of this
material is, of course, provided by the Plant Genetic Resources Centre
(PGRC/E).
Concerning the international value of Ethiopian wheats, we have
already mentioned Vavilov's 1927 visit and his collections of wheat,
which were used widely by breeders in the Soviet Union. Much of
this material was also made available, we understand, to breeders in
Germany and other European countries.
The genebank base collection at the International Centre for Maize
and Wheat Improvement (CIMMYT) earlier possessed 16 bread
wheat and 69 durum wheat accessions (Sencer, 1988), and latterly
1800 Ethiopian wheat entries were introduced from genebanks in the
USA, Germany and Italy. Six Ethiopian durum wheats are currently
being used in the CIMMYT breeding programme.
Perrino (1988) also reported on Ethiopian wheats collected on three
separate expeditions from Italy in the early 1970s and currently stored
in the Germplasm Institute at Bari. These wheats were mainly
tetraploid, and amounted to over 400 accessions, in which high levels
of diversity were found.
Barley
Nearly 5000 Ethiopian barley accessions are stored at
PGRC/E. Their high diversity was confirmed by Engels (Chapter 9),
with the diversity fairly evenly spread throughout the areas where
the crop is grown. Breeding for this crop in Ethiopia has turned away
from the exclusive use of exotic materials and towards a greater
emphasis on the autochthonous germplasm (Hailu Gebre and Fekadu
Alemayehu, Chapter 24). Current breeding concentrates on population improvement of landraces by mass selection and other appropriate methods. The evaluation of Ethiopian barleys reveals useful
362
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characters such as resistance to barley yellow dwarf virus, powdery
mildew, net blotch and loose smut as well as high protein quality,
high tillering quality, tolerance to marginal soil conditions and
vigorous seedling establishment. Breeding trials have shown promising results, and new potential cultivars are being recommended. Both
malting and food barley have received attention among Ethiopian
breeders and the prospects are highly satisfactory.
On an international scale, the Ethiopian barleys became famous
ever since the expeditions of H. V. Harlan in 1923 (see H. V. Harlan,
1957), N. I. Vavilov in 1917 and E. L. Smith and C. Thomas in 1963-4.
Qualset (1975) has described the high degree of resistance to barley
yellow dwarf disease in Ethiopia, particularly in the area just north of
Addis Ababa where the concentration of resistance alleles is very
high, according to screening carried out in the USA.
According to Witcombe (1983) more than one-third of the total
barley collections available worldwide originated in Ethiopia - a total
of nearly 4500 accessions. Barley materials are noted also for CIMMYT
(Sencer, 1986) and Bari, Italy (Perrino, 1988). Furthermore, Somaroo
& Holly (1988) report that the International Center for Agriculture
Research in the Dry Areas (ICARDA) possesses nearly 2500 barley
collections from Ethiopia, mostly 6-rowed and deficiens types; these
have shown considerable promise in the search for earlier heading
and maturing characters, as well as high protein content.
Recent barley collections were described by Toll (1980, 1981), totalling 675 samples, of which duplicates were to be sent to Braunschweig, West Germany, and the rest stored at PGRC/E.
Thus the quality of Ethiopian barleys both in terms of total diversity and, most importantly, the types of resistance and adaptation
genes found, shows their extremely valuable importance to breeders
in Ethiopia and in the international community.
Sorghum and millets
The primary centre of origin and diversity of sorghum is
assumed to be the Sudan and Ethiopia (see also Doggett, Chapter 10),
and it is thus not surprising that useful genetic traits have been found
among Ethiopian landraces.
Sorghum breeders in Ethiopia have found sources of cold
resistance, high protein (lysine), good grain quality, resistance to
stalk borer, downy mildew, smuts, bacterial streak, anthracnose and
Striga, as well as useful agronomic and kernel characters (Yilma
Kebede, Chapter 25). Landrace selections from the sorghum collec-
The use of Ethiopian germplasm
363
tions have been valuable and five entries are on the recommended
list, two for high elevations, two for intermediate and one for low
elevation areas. It is confidently expected that more screening and
more germplasm exploration in Ethiopia will reveal a wider range of
valuable traits than is known at present.
Ethiopian sorghums are also well known internationally. Thus
Mengesha & Remanandan (1988) report that nearly 4500 accessions
are stored at the International Crops Research Institute for the SemiArid Tropics (ICRISAT). These authors draw particular attention to
the sorghum line E35-1 which has been selected from a Zera-zera
landrace from Ethiopia, introduced into West Africa for direct cultivation and also used in several African breeding programmes. Other
important characteristics, which were identified in a recent expedition to south-west Ethiopia (Ahluwalia et ah, 1987) in Ethiopian
sorghum (cultivated, wild and weedy types of forage sorghum),
include lodging resistance, stem sweetness and high tillering
capacity.
Nearly 300 accessions of Ethiopian minor millets also figure in the
ICRISAT collections. Characters of interest include morphology,
height, number of spikes per inflorescence, size and shape of the
fingers, resistance to lodging and tillering capacity.
Since Ethiopia is the only country that grows teff as a cereal crop
this section will be concerned only with Ethiopia. Its importance in
terms of area cultivated is greater than any other cereal crop in Ethiopia (Seyfu Ketema, Chapter 26) and improvement has been initiated
through selection from landraces and mutation breeding. The main
aims are to develop lodging resistant, high-yielding stable varieties;
so far, higher yielding materials have been obtained through pure
line selection and hybridization (for further details see Seyfu Ketema,
Chapter 26).
Grain legumes (pulses)
Of 13 million hectares currently under cultivation in Ethiopia,
pulses occupy about 13-14 per cent of the cultivated area, cereals ca.
83 per cent and oil crops ca. 4 per cent. Faba bean is the major pulse
crop, occupying about 6 per cent of the total area under major crops,
followed by field pea, chickpea and lentil.
Ethiopia has high inter- and infraspecific diversity of pulse crops.
Several species of pulse crops have been identified (Westphal, 1974),
and their genetic resources and utilization are described by Mekbib,
Demissie & Tullu (Chapter 27). Recent work indicates that Ethiopia is
364
/. G. Hawkes & Melaku Worede
an important centre of diversity for some cool season legumes,
namely, field pea and chickpea (van der Maesen et al., 1988), and
Ethiopia is considered the secondary centre of diversity for Lens
culinaris (Zeven & Zhukovsky, 1975).
Varietal improvement efforts at national level are at rather an early
stage with regard to pulse crops. However, a considerable number of
accessions of faba bean, field pea, chickpea, lentil, etc. are being
incorporated in crop improvement programmes. Breeding for pulse
crops in Ethiopia is largely dependent on the use of indigenous germplasm. Evaluation of various pulse crops reveals useful characters
such as earliness, high number of pods and tolerance to some adverse
soil conditions. Further details are provided in Chapter 29 (Hailu
Mekbib et al).
On an international dimension the Ethiopian grain legume germplasm has received considerable attention. ICRISAT stores over 900
accessions of Ethiopian chickpea and 14 of pigeon pea. These are
currently being screened for resistance characters (Mengesha &
Remanadan, 1988).
The ICARDA genetic resources unit contains a total of 375 Ethiopian lentil accessions which, when screened in Syria, showed early
flowering and maturing characters. These lines were small-seeded,
with black or dark brown testa (Somaroo & Holly, 1988).
Perrino (1988), describing the Italian expeditions, reported 96
entries of Pisum, 38 of faba bean and 68 of other legumes from Ethiopia. Screening data are not mentioned.
Oil crops
Ethiopia is considered to be the centre of origin or diversity of
a number of important oil crops, such as niger seed (Guizotia abyssinica), Ethiopian mustard (Brassica carinata), safflower (Carthamus
tinctorius), linseed (Linum usitatissimum) and castor bean (Ricinus communis). There are national breeding schemes for all of these and
several others of slightly lesser importance (Hiruy Belayneh, Chapter
28).
Oil-crop breeders are concentrating on developing varieties with
higher yields, improved quality and disease resistance. For Guizotia,
in addition, special attention is being given to lodging resistance,
uniform ripening, minimal shattering, frost tolerance and resistance
to Septoria disease. Many indigenous lines are doing well in variety
trials, and landrace selection is also in progress.
Brassica carinata, B. nigra, B. rapa and B. oleracea are all under con-
The use of Ethiopian germplasm
365
sideration for the production of better cultivars with low erucic add,
low glucosinolate and fibre concentrations and resistance to diseases,
especially Alternaria leaf spot. Useful selections of B. carinata have
already been derived from Ethiopian landraces, and a crossing programme is under way.
Linseed breeding is focussing on oil yield as well as the general
characters mentioned above, with resistance to wilt, powdery mildew
and Septoria. Promising selections have already been made from the
national germplasm collection.
Safflower (Carthamus) work includes the selection of promising
lines from the national collection and progeny selections.
The castor bean (Ricinus) programme is still only two years old but
already some outstanding lines have been observed from the national
germplasm collection and have been advanced for further study.
Finally, work on the non-indigenous oil crops, sunflower, sesame
and groundnut, is in progress.
The Ethiopian oil plants have engendered considerable interest
internationally. There are collections stored in a number of seed
banks in Europe (Holland, Sweden, UK, etc.) and also in other continents (Kebebew, 1988). Some of these collections have recently been
'repatriated' to Ethiopia. A collection of 76 Brassica accessions was
made recently by Astley, Haile Giorghis and Toll (1982), of which a
duplicate set of samples was sent for cytotaxonomic screening to the
Vegetable Gene Bank of the Institute of Horticultural Research,
Warwick, UK. These include nearly 60 samples of B. carinata.
Coffee
Coffee (Coffea arabica) originated in Ethiopia and for that
reason its genetic diversity is considered to be higher in that country
than in other regions where it was introduced in more recent times.
The significance of Ethiopian coffee genetic resources in coffee
improvement is dealt with in detail by Mesfin Ameha (Chapter 29)
and thus need not be described at length in this chapter. It is worth
reiterating, however, that coffee in Ethiopia has become adapted to a
very wide eco-climatic range and with excellent traits for yield,
quality and resistance to diseases and pests. However, grave alarm is
being expressed at the rapidly diminishing wild resources caused by
forest destruction and other changes in land use.
International interest in Ethiopian coffee has been high since the
first decades of this century; the works of Chevalier (1929) and Carvalho (1956, 1959) especially, have drawn attention to the need for
366
/. G. Hawkes & Melaku Worede
further international investigations on this crop, particularly as some
resistance to Hemileia was reported in Ethiopian collections (see also
Meyer, 1965, and Vayssiere, 1961, for further details).
Forage grasses and legumes
Work on Ethiopian forage genetic resources at the International Livestock Centre for Africa (ILCA) is described by Hanson and
Solomon Mengistu (Chapter 15) and by Lazier and Alemayehu
Mengistu (Chapter 21). Although African grasses are of considerable
importance, in general the emphasis in Ethiopian germplasm work
lies with the legumes. Evaluation in plot and field trials of Trifolium
and Vicia species for higher altitudes is in progress, with certain
Argyrologium, Macrotyloma, Eriosema, Neonotonia, Indigofera, Crotalaria
and Stylosanthes accessions also showing promise. Among the grasses
only Melina minutiflora and a Zornia species performed well.
In feeding trials several Acacia species and Sesbania sesban have
seemed to perform reasonably well.
Many accessions of highland and lowland forage species have also
been collected, but it seems that no landraces exist, since cultivation
has been in progress for only 50 years. Thus collecting has been
restricted to wild populations, and in the last six years some 1700
accessions were collected and entered into the seed bank as well as
being set out in observation trials and field plots. No breeding and
selection work has yet been attempted, as far as we are aware.
Final remarks
In this chapter we have attempted to set out some of the
highlights of Ethiopian germplasm utilization in national programmes within Ethiopia. Clearly, much more detailed information is
given in the appropriate chapters.
The information on the use of Ethiopian germplasm internationally
is not so precise, since it has been impossible to scan through the
plant breeding literature of the whole world in the hope that one or
two relevant pieces of information might be revealed. We have thus
relied chiefly on materials provided from international and national
centres at the 1986 conference (Engels, 1988) which were not featured
in the present book. There are also a limited number of easily accessible references on collecting in Ethiopia by foreign missions, germplasm exchange data and general papers and books by Vavilov, H. V.
Harlan, Chevalier and others known to us in the literature.
Despite this limited amount of information from international
The use of Ethiopian germplasm
367
sources it is quite clear that Ethiopian genetic resources have been
valuable in the past, are valuable in the present, and, it is hoped, will
become even more valuable in the future. Future development will
only be possible, however, if the free exchange and flow of germplasm internationally between Ethiopia and other countries, carried
out to mutual advantages, is maintained and strengthened (see also
Worede, Chapter 1). Such a movement of germplasm would help to
develop screening and utilization programmes, to the ultimate benefit of both Ethiopia and the world at large.
References
Ahluwalia, M., Dabas, B. S., Seme, E. N., Demissie, A. & Nafie, N. A. (1987).
Exploration and collecting landraces of cultivated wild and weedy types of
forage sorghum in Kenya, Ethiopia, Sudan. IBPGR Mission Report No.
87/43.
Astley, D., Mehateme, Haile Giorghis & Toll, J. (1982). Collecting Brassicas in
Ethiopia. Plant Genetic Resources Newsletter, 51, 15-20.
Carvalho, A. (1956). O cafe selvagem da Abissinia. Boletin Superintend. Sew.
Cafe, Sao Paulo, 31, 13-15.
Carvalho, A. (1959). Preliminary information on the genetics of Ethiopian
coffee. Nature (London), 183, 906.
Chevalier, A. (1929-47). Les Cafeiers du Globe, 3 vols. Paris.
Engels, J. M. M. (ed.) (1988). The conservation and utilization of Ethiopian
germplasm. Proceedings of an international symposium, Addis Ababa, 1316 October 1986 (mimeographed).
Harlan, H. V. (1957). One Man's Life with Barley. Exposition Press, New York.
Kebebew, F. (1988). Germplasm exchange and distribution by PGRC/E. In:
J. M. M. Engels (ed.), The conservation and utilization of Ethiopian germplasm. Proceedings of an international symposium, Addis Ababa, 13-16
October 1986, pp. 276-84 (mimeographed).
Mengesha, M. H. & Remanandan, P. (1988). The gene bank at ICRISAT and
its significance for crop improvement in Africa with special reference to
Ethiopian germplasm. In: J. M. M. Engels (ed.), The conservation and
utilization of Ethiopian germplasm. Proceedings of an international
symposium, Addis Ababa, 13-16 October 1986, pp. 333-49
(mimeographed).
Meyer, F. G. (1965). Notes on wild Coffea arabica from southwestern Ethiopia
with some historical associations. Economic Botany, 19, 136-51.
Perrino, P. (1988). Country report on plant genetic resources in Italy. In:
J. M. M. Engels (ed.), The conservation and utilization of Ethiopian germplasm. Proceedings of an international symposium, Addis Ababa, 13-16
October 1986, pp. 377-89 (mimeographed).
Qualset, C. O. (1975). Sampling germplasm in a centre of diversity: an example of disease resistance in Ethiopian barley. In: O. H. Frankel and J. G.
Hawkes (eds), Crop Genetic Resources for Today and Tomorrow. Cambridge
University Press, Cambridge, pp. 81-96.
Sencer, H. A. (1988). CIMMYT's wheat germplasm bank and its significance
for crop improvement in Africa with special reference to Ethiopia. In:
368
/. G. Hawkes & Melaku Worede
J. M. M. Engels (ed.), The conservation and utilization of Ethiopian germplasm. Proceedings of an international symposium, Addis Ababa, 13-16
October 1986, pp. 353-62 (mimeographed).
Somaroo, H. B. & Holly, L. (1988). The significance of plant genetic resources
for crop improvement at ICARDA with special reference to Ethiopian
barley and lentil germplasm. In: J. M. M. Engels (ed). The conservation and
utilization of Ethiopian germplasm. Proceedings of an international
symposium, Addis Ababa, 13-16 October 1986, pp. 340-52
(mimeographed).
Toll, J. (1980). Collection in Ethiopia. Plant Genetic Resources Newsletter, 43,
36-9.
Toll, J. (1981). Collection in Ethiopia. Plant Genetic Resources Newsletter, 48,
18-22.
Van der Maesen, J.J.G., Kaiser, W.J., Marx, G. A. & Worede, M. (1988).
Genetic basis for pulse crop improvement: collection, preservation and
genetic variation in relation to need traits. In: R.J. Summerfield (ed.),
Proceedings of the International Food Legume Research Conference on pea, lentil,
faba bean, chickpea, Spokane, Washington, USA, 6-11 July 1986, pp. 55-66.
Kluwer, Dordrecht.
Vavilov, N. I. (1931). The wheats of Abyssinia and their place in the general
system of wheats. Bulletin of Applied Botany, Genetics and Plant Breeding,
supplement 51, p. 233.
Vayssiere, P. (1961). L'Ethiopie, pays d'origine du cafeier d'Arabic Cafe,
Cacao, The, 5, 77-81.
Westphal, E. (1974). Pulses in Ethiopia, their Taxonomy and Agricultural Signifi-
cance. PUDOC, Wageningen, p. 261.
Witcombe, J. R. (1983). A provisional world list of barley expeditions. Plant
Genetic Resources Newsletter, 53, 25-40.
Zeven, A. C. & Zhukovsky, O. M. (1975). Dictionary of Cultivated Plants and
their Centres of Diversity. PUDOC, Wageningen, p . 219.
Index
abalo see Brucea antidysenterica
Abelmoschus spp. 215
A. esculentus 35, 45, 62-3, 240
A. ficulneus 62-3
A. manihot 35
A. moschatus 35
abish see Trigonella foenum-graecum
Acacia spp. 43, 85, 95
A. abyssinica 86
A. albida 285
A. cyanophylla 285
A. decurrens 97
A. mearnsii 97
A. saligna 97
A. Senegal 43
A. seya/ 285
A. xiphocarpa 87
use as forage 219, 220, 221, 223, 224
Acanthus spp. 345
accumulation centre concept 23-4
Achyranthes aspera 105
adenguare see Vigna unguiculata
Adenia ellenbeckii 175
ades see Myrtus communis
adja see Triticum polonicum
Aeonium 78
aerial yam see Dioscorea bulbifera
Aeschynomene sp. 219, 221, 223
afer kocher see Hedychium spicatum
Aframomum spp.
A. korarima 36, 51, 71-2
germplasm resource study 123, 125,
127-8, 129-30, 240
A. polyanthum 72
A. sanguineum 72
african millet see Eleusine coracana
Afro-montane floristic region 76
agam see Carissa edulis
agriculture
crop history 141-2, 144-5
crop spread 153-7
history of development 140-1
management techniques 145-6
regional development 142-3
role of the hills 144
role of the Nile 143
agro-climatic belts 83
Ajuga remota 105
akat see Hyphaene thebaica
Albizia spp. 221, 223
A. schimperiana 86, 96
aleqnay see Sorghum spp.
alkoka see Phaseolus vulgaris
Alliaceae 51, 67
Allium spp. 240
A. alibile 67
A. cepa 51, 67, 175
A. sativum 51, 67, 123, 175
A. subhirsutum 67
Allophyllus spp. 86
A. abyssinicus 96, 345
alma see Amaranthus caudatus
Alternaria leaf spot 349, 352
Alysicarpus sp. 219, 221, 223
Amaranthaceae 44, 45, 175
Amaranthus spp. 44, 240
A. caudatus 44, 45, 173
A. hybridus 45
amera see Plumbago zeylanicum
amija see Hypericum quartinianum
Ammi copticum 54
Amorphophallus spp. 240
A. abyssinicus 36, 51, 67, 174, 183, 215
A. gallaensis 67
A. gomboczianus 67
Anacardiaceae 44, 177
anamero see Ajuga remota
anchabi see Ocimum suave
anchote see Coccina abyssinica
Andropogon 221, 223
370
Index
Anethum graveolens 4A, 45, 105
angular leaf spot resistance 273
Aningeria adolfi-friedericii 77, 86, 96
anise see Pimpinella anisum
Anogeissus leiocarpus 77
antate-welakha see Salvia nilotica
anthracnose resistance 320
Apiaceae 44, 45, 46, 47, 54
Apiutn spp.
A. graveolens 44, 45
A. leptophyllum 44, 45
A. nodiflorum 44, 45
Apodytes spp. 86
A. dimidiata 96
Arabian floristic province 76, 77
arabica coffee see Coffea arabica
arable crop spread in prehistory 153-7,
161-4
Araceae 51, 67-8, 183
Arachis hypogaea
crop development 351-2
germplasm stores 240, 266, 346
origins 339
arangama see Capparis tomentosa
arda bofa see Cassia occidentalis
Arecaceae 52, 68
Argemone mexicana 345
Argyrolobium spp. 221
A. ramosissimum 283
Arisaema spp. 174, 183, 215, 240
ariti see Artemisia rehan
arkokobay see Hyphaene thebaica
aromatic plants 114-21
Artemisia spp.
A. abyssinica 104
A. afra 104
A. rehan 120
Arundinaria alpina 86
Arundo donax 78
Asclepiadaceae 176
aserkush see Cyphostema niveum
ashakilta see Cajanus cajan
Asparagus spp. 51, 68
A. africanus 68, 105
A. asiaticus 68
astenagir see Datura stramonium
Asteraceae 46, 47, 54
atara see Pisum sativum
ater see Pisum sativum
atuch see Verbena officinalis
augmented design in germplasm
evaluation 271-2
aureta see Azanza garckeana
Avena spp. 240
A. abyssinica 30, 42, 51, 69
A. barbata 30, 69
A. sativa 203
A. vaviloviana 30, 69
Azadirachta indica 97
azamir see Bersame abyssinica
Azanza garckeana 177
azkuti see Ocimum
azo-hareg see Clematis sinensis
bacterial streak resistance 320
bacterial stripe 292
bagana see Amorphophallus abyssinicus
bakala see Vicia faba
Balanites spp. 85
B. aegyptiaca 95, 178, 345
bambara see Vigna subterranea
banana see Musa
banshalla see Sauromatum nubicum
bapello see Phaseolus lunatus
Barbeya oleoides 77
barley see Hordeum vulgare
barley yellow dwarf virus 254-5, 306
basil see Ocimum
basobila see Ocimum basilicum
bean herb see Satureja sp.
beetroot see Beta vulgaris
bekela see Vicia faba
beles see Ficus carica
bengal bean see Mucuna pruriens
berbere see Capsicum annum
Berha agro-climatic belt 83-4
Bersame spp. 88
B. abyssinica 86, 105, 345
bessobila see Ocimum basicilicum
Beta vulgaris 78
bifti see Warburgia ugandensis
birchik see Citrullus lanatus
birgud see Cinnamomum cassia
bisana see Croton macrostachys
black cumin see Nigella sativa
black mustard see Brassica nigra
black olive scale see Saisetia oleae
black pepper see Piper nigrum
boita see Hordeum vulgare
bolokie see Phaseolus vulgaris
Borassus aethiopum 68
Boswellia spp. 43, 77, 85, 116-17
B. rivae 178
bottle gourd see Lagenaria siceraria
boyye see Dioscorea alata
Brachiaria spp.
B. brizantha 219, 220, 221, 223
B. decumbens 219
B. mutica 219
Brassica spp.
B. campestris 45, 54
B. carinata
diversity 15, 30, 37
germplasm resources 346, 348-50
origins 30, 54
role in Konso agriculture 175, 184
uses 45
B. integrifolia 45, 54
Index
B. juncea 45, 54
B. napus 346, 348-50
B. nigra
diversity 15, 30
germplasm resources 124, 346, 34850
origins 30, 54
uses 45, 105
B. oleracea
diversity 30, 78
germplasm resources 215, 346, 34850
origins 30, 54
uses 45
conservation work 205
drought research 10
germplasm multiplication 261, 266
PGRC/E stock 230
resource value 364r-5
Brassicaceae 45, 47, 48, 49, 54^-5
breadwheat see Triticum aestivum
broadleaved forest types 86-7
Brucea spp. 86
B. antidysenterica 43, 105
buckthorn see Rhamnus prinoides
buke seytana see Momordica charantia
bulrush millet see Pennisetum americanum
bultug see Pennisetum americanum
buna see Coffea arabica
bunt see Tilletia
burie see Arisaema
Burseraceae 79-80
bushland distribution 85-6
cabbage see Brassica oleracea
cabbage tree see Moringa stenopetala
Cadia purpurea 77
Caesalpinoideae 218
Cajanus cajan 34, 173, 182, 240, 331, 341
Calotropis procera 105
Calpurnia aurea 105
Canarian floristic province 77, 78
Canarina 78
Canavalia spp.
C. africana 59
C. ensiformis 45, 59, 341
C. virosa 59, 341
Cannabis sativa 45, 55
Capparis tomentosa 106
Capsicum spp. 46, 47, 205, 240
C. abyssinicum 67
C. annuum 123, 125-6, 129, 176, 184
C. frutescens 67
caraway see Carum carvi
cardamom see Eletteria cardamomum
Carica papaya 177
Carissa edulis 106
carrot see Daucus carota
371
Carthamus spp.
C. flavescens 54
C. lanatus 54
C. oxycantha 32
C. persicus 54
C. tinctorius
germplasm documentation and
development 240, 263, 346, 352
origins 30, 54
uses 46, 177
Carum spp.
C. carvi 44, 46
C. copticum 36, 54, 240
cassava see Manihot esculenta
Cassia spp. 220, 221
C. occidentalis 106
C. senna 50, 59
castor bean see Ricinus communis
Casuarina equisetifolia 97
Catha edulis 35, 37, 46, 55, 178
Celastraceae 46, 55
celery see Apium graveolens
Celosia 240
Celtis spp. 88
C. africana 77, 86, 96
Cenchrus spp. 221
C. ciliaris 279
centre of diversity concept 23, 202
Centre Technique Forestier Tropical 99
Centro International de Agricultura
Tropical (CIAT) 220
cereals
conservation and exploration 6,
208-14
diversity 24^-30
germplasm multiplication 263
modern crop replacements 203-4
resource value 360-3
role in Konso agriculture 173, 180-2
chat see Catha edulis
check entries in germplasm evaluation
270-1
chemotaxonomy 254
chickpea see Cicer arietinum
chili pepper see Capsicum
chiz inchet 119
Chloris spp. 221, 223
C. gayana 43, 219, 279
Chlorophora exelsa 96
Chnootriba similis 292
chocolate spot resistance 33
Christ thorn see Ziziphus spina-christi
Cicer spp.
C. arietinum
conservation work 205, 230, 240,
263, 266, 269
crop production details 209, 213
diversity 33, 331
origins 33, 59, 335
372
Index
deer spp. (cont.)
role in Konso agriculture 182
use 46, 174
C. cuneatum 33, 59, 335
Cinnamomum spp.
C. cassia 117-18
C. zeylanicum 124
cinnamon see Cinnamomum zeylanicum
Citrullus spp.
C. colocynthis 55
C. lanatus 46, 55
G'frws spp.
C. aurantifolia 46, 66-7, 177
C. sinensis 177
Clematis sinensis 106
Clerodendrum spp.
C. fl/a£um 106
C. myricoides 106
clove see Syzygium aromaticum
cluster bean see Cyamopsis tetragonoloba
Coccina abyssinica
conservation 240
origins 36, 42, 58
use 46, 215
coconut see Cocos nucifera
Cocos nucifera 345
Coffea arabica
conditions of growth 355
conservation efforts 6-7, 15, 195-7, 205
distribution 46, 66, 86
diversity 34-5, 37, 355-7
origins 34, 354r-5
PGRC/E stock 230, 240
problems of losses 357-8
resource value 365-6
role in Konso agriculture 178, 184
coffee see Coffea arabica
Coleus edulis see Plectranthus edulis
collecting methods for germplasm 206-8
Colletotrichum graminicola 320
Colocasia spp. 240
C. esculenta 67-8, 175, 183
Combretum 77, 85
Commicarpus pedunculosus 77
Commiphora spp. 77
C. africana 115-16
C. erythraea 115-16
C. gileandensis 115-16
C. abyssinica 115-16
C. hodai 115-16
C. kua 115-16
C. myrrha 115-16
C. quadricincta 115-16
C. schimperi 115-16
C. truncatum 115-16
common bean see Phaseolus vulgaris
Commonwealth Forestry Institute 99
Commonwealth Scientific and Industrial
Research Organization 99
Compositae 54
coniferous forest types 87-8
conservation facilities 226-7
PGRC/E system 229-34
conservation methods for genetic
resources
forests 9 1 ^
fruits and nuts 195-7
seeds 190-3, 229
vegetative parts 193—4
Corchorus oligatorus 46, 67, 240
Cordeauxia spp. 220
C. edulis 42, 330, 341
Cordia spp.
C. abyssinica 86, 96
C. africana 95
coriander see Coriandrum sativum
Coriandrum sativum
conservation 123, 125, 127, 130, 240
origins 36, 44
use 176
cotton see Gossypium herbaceum
cowpea see Vigna unguiculata
Crambe spp.
C. abyssinica 32, 47, 55, 240
C. hispanica 55
C. kilmandscharica 55
C. sinuato-dentata 55
cress see Lepidium sativum
Crotalaria spp. 219, 221, 283
Croton spp. 85
C. macrostachys 86, 95, 96, 106
Cruciferae 54r-5
Cucumis spp.
C. aculeatus 106
C. dipsaceus 58, 106
C. figarei 58
C. humifructus 58
C. insignis 58
C. me/o 47, 58
C. metuliferus 58
C. srfzzu 58
Cucurbita spp. 47, 240
C. /za/b/w 58
C. maxima 58
C. moschata 58
C. pepo 58, 175
Cucurbitaceae 46, 47, 48, 55, 58-9
cumin see Cuminum cyminum
Cuminum cyminum 44, 123, 128, 130, 240
Cupressus lusitanica 97
Curcuma longa 124, 240
Cussonia 86
Cyamopsis spp.
C. senegalensis 59
C. tetragonoloba 46, 59
Cymbopogon spp.
C. cifrafws 51, 71, 120
C. commutatus 71
Index
C. excavatus 71
C. floccosus 71
C. giganteus 71
C. nervatus 71
C. proximus 71
C. schoenanthus 71
Cynodon sp. 219, 223
Cyperus bulbosus 120
Cyphomandra betacea 240
Cyphostema niveum 106
dabo sindi see Triticum aestivum
Dactylis glomerata 78
dagusa see Eleusine coracana
dahanta see Lagenaria siceraria
Dalbergia melanoxylon 96
dangarda itana see Boswellia rivae
date palm see Phoenix dactylifera
Datura stramonium 107, 240
Daucus spp.
D. carota 47, 54, 78
D. hochstetteri 54
Dega agro-climatic belt 84
Delonix elata 77
desert date see Balanites aegyptiaca
Desmodium sp. 221
Dianthus 77-8
Dichrostachys sp. 221
dicotyledons, diversity of 44—67
Digera alternifolia 175
digita see Calpurnia aurea
Digitaria sp. 219, 221, 223
dill see Anethum graveolens
dimbilal see Coriandrum sativum
dinitscha faranjeta see Ipomoea batatas
Dioscorea spp. 36, 51, 68-9, 70, 183,
240
D. abyssinica 68-9, 70
D. alata 68-9, 70
D. bulbifera 51, 68-9, 70
D. cayensis 68-9, 70
D. cochleari-apiculata 70
D. dumetorum 70
D. g*7/ettz 68-9, 70
D. /ecflrdn' 68-9, 70
D. odoratissima 68-9, 70
D. auartiniana 70
D. schimperana 68-9, 70
Dioscoreaceae 51, 68-9, 70
Diospyros spp. 96
Diplolophium spp.
D. abyssinicum 47
D. africanum 54
dirb keteto see Sorghum spp.
disease resistance studies
barley 25, 306
linseed 350
safflower 352
sorghum 10, 26, 320
373
wheat 10, 27, 264, 290-2, 296-8, 298301
diversity, documentation of 190
diversity centre concept 23, 202
diversity index 133-9
dog see Diplolophium abyssinicum
dokma see Syzygium guineense
Dolichos lablab see Lablab purpureus
Dombeya spp. 96
downy mildew 320
Dracaena steudneri 107
drought
plant resistance studies 25, 26, 34
role in genetic erosion 203
drugs from plants 104, 178
duba see Cucurbita
dum palm see Hyphaene thebaica
dupana see Ensete ventricosum
durum see Triticum durum
ebicha see Vernonia amygdalina
Echinochloa sp. 221
Echinops spp. 119
einkorn see Triticum monococcum
Ekebergia spp. 86, 88
E. capensis 96
elephant grass see Panicum maximum
Eletteria cardamomum 124
Eleusine spp.
£. africana 29, 71, 161-2, 240
E. compacta 161
£. coracana
origins 29, 37, 71, 145, 161
role in Konso agriculture 182
use 51, 173
E. coracana conservation 205, 230, 240,
263
£. elongata 161
£. indica 71
£. plana 161
£. vulgaris 161
Embelia schimperi 36, 240
embuacho see Rubia cordifolia
emmer see Triticum turgidum
endahula see Kalanchoe lanceolata
endemism estimates 78-9, 80-1
endod see Phytolacca dodecandra
enkoko see Embelia schimperi
enset (ensete) see Ensete ventricosum
Ensete ventricosum
diversity 35, 37, 42, 69
use 52, 179, 215
Entada sp. 221
environmental classification 88-91
environmental impact on germplasm
268-70, 273-1
Eragrostis spp.
E. pilosa 28-9, 71, 325
374
Index
Eragrostis spp. (cont.)
E.tef
conservation 205, 230, 240, 263
crop breeding 323-5
crop production 204, 209, 211, 212
diversity 28-9, 42, 71, 77
history of cultivation 144, 325-6
origins 28, 325
use 51, 173, 326-7
Erica arborea 86
Eriosema spp. 221, 224
E. psoraleoides 283
Erythrococca abyssinica 77
Eruca sativa 47, 55
Erucastrum 55
Erysiphe graminis 25, 27, 290, 350
Erythrina spp. 220, 221, 223, 224
E. abyssinica 95, 284, 345
E. brucei 95, 107, 284, 285-6
Ethiopian caraway see Trachyspermum
ammi
Ethiopian Flora Project 75-6
Ethiopian kale see Brassica carinata
Ethiopian mahogany see Trichilia roka
Ethiopian mastic see Pistacia aethiopica
Ethiopian mustard see Brassica carinata
Ethiopian oats see Avena abyssinica
etse menhae see Securidaca
longipedunculata
etse patos see Dracaena steudneri
Eucalyptus spp. 92-3
E. globulus 77
Euphorbia 85
Euphorbiaceae 50, 59
Eurasian floristic province 77-8
faba bean see Vicia faba
Fabaceae 45, 46, 47, 48, 49, 50, 59-61
Fagopyrum esculentum 240
false banana see Ensete ventricosum
falsolya see Phaseolus vulgaris
faranjeta see Cajanus cajan
farmers, role in conservation of 15-16
fendish see Sorghum spp.
fennel see Foeniculum vulgare
funugreek see Trigonella foenum-graecum
Festuca sp. 77-8, 221
feto see Lepidium sativum
fibre plants 36, 178
Ficus spp. 86
F. carica 47, 64
F. palmata 64
F. vasta 111
field genebanks 6-7, 13-14
field mustard see Brassica campestris
field pea see Pisum sativum
fig see Ficus carica
finger millet see Eleusine coracana
fiti see Clematis sinensis
flavonoids pattern 25
flax see Linum usitatissimum
Foeniculum vulgare 47, 54, 123, 128, 176
Food and Agriculture Organization 99
forage
conservation 218-19
diversity 278-9
evaluation 279-86
resource value 366
Forestry Research Centre 98, 99
forests
classification 84-6
broadleaved 86-7
coniferous 87-8
conservation 91-4
development 94^7
distribution 82-3, 88-91
re-establishment 97-9
frankincense see incense
fruit
conservation methods 195-7
role in Konso agriculture 177-8, 184
fua see Sterculia africana
Fusarium spp. 290, 296
futota see Gossypium hirsutum
Galactia sp. 224
Galiniera 86
Galinsoga parviflora 78
galla potato see Plectranthus edulis
gamadeda sira see Sorghum bicolor
gan seber see Sorghum spp.
gancho see Sapium ellipticum
garatita see Gossypium herbaceum
garden cress see Lepidium sativum
garden rocket see Eruca sativa
garlic see Allium sativum
geba see Ziziphus spina-christi
gebs see Hordeum vulgare
gene centre concept 23-4, 202
gene mapping 255
gene pool concept 253-4
genebanks
creation and maintenance 5-6, 7
distribution 10-13
evaluation 7-8
future uses 13-18
role in forestry 93-4
utilization 8-10
genetic erosion 202-4
genotype x environment effects 268-70,
273-4
German Agency for Technical
Cooperation 208
germplasm conservation
characterization 262-4
collection (collecting)
methodology 189-90
recording methods 197-9
375
Index
role of markets 197
sampling strategies 190-7
disease studies 298-301, 306, 320
documentation 235-44
genebanks 5-13
PGRC/E role 13-18, 235-43
enhancement 252
evaluation
augmented design 271-2
check entries 270-1
data analysis 274-6
nearest neighbour analysis 272
pre-breeding 251-2
role of IBPGR 247-51
site/season effects 273-4
taxonomy 252-6
multiplication 258-60
gesho see Rhamnus prinoides
gibto see Lupinus albus
ginger see Zingiber officinale
girawa see Vernonia amygdalina
gizawa see Withania somenifera
glume blotch see Septoria nodorum
Glycine max 341
goa bean see Psophocarpus palustris
godere see Colocasia esculenta
gomano see Brassica carinata
gomen see Brassica carinata
gomenzer see Brassica integrifolia
gonada see Sorghum bicolor
Gossypium spp. 240, 345
G. anomalum 63
G. arboreum 63-4
G. barbadense 64
G. benadirense 63
G. bricchettii 63
G. herbaceum 36, 63-4, 179, 184
G. hirsutum 64, 178
G. somalense 63
gourd see Lagenaria siceraria
grain amaranth see Amaranthus caudatus
Gramineae 69-71
grass pea see Lathyrus sativus
grasses
conservation 218-19
evaluation 279
green gram see Vigna radiata
Grevillea robusta 97
Grewia tenax 177
groundnut see Arachis hypogaea
guaya see Lathyrus sativus
Guizotia spp.
G. abyssinica
conservation 205, 230, 240, 261, 263,
266,346
crop production 20, 213, 348
diversity 30-1, 37, 54
origins 30-1, 144
use 47, 347
G. scabra 31, 54, 240, 346
gulo see Ricinus communis
gum myrrha see myrrh
gum olibanum see incense
gum oppopanax see myrrh
gumamila see Polygonum barbatum
gums 43
guracha see Capparis tomentosa
gurage gomen see Brassica oleracea
ha dida see Sorghum bicolor
habatalumuluk see ]atropha curcas
Habenaria spp. I l l
habesha sindi see Triticum durum
habhab see Citrullus lanatus
hadia see Salvadora persica
hafukagne see Sorghum spp.
Hagenia abyssinica 43, 86, 96
halako (haleko) see Moringa stenopetala
hamba guita see Amorphophallus
abyssinicus
hangalta see Balanites aegyptiaca
hangoleita see Launaea taraxacifolia
harboreda see Sorghum bicolor
hardwood potential 96-7
hareg resa see Zehneria scabra
hargiti see Sorghum bicolor
haricot bean see Phaseolus vulgaris
hausa potato see Plectranthus edulis
health care and plants 101-12
health regulation in plant conservation
11
Hedychium spicatum 119
Helianthus annuus 177, 240, 263, 346, 350-1
Helminthosporium spp. 290
hemp see Cannabis sativa
henna see Lawsonia inermis
herbs and health care 104-12
Heteromorpha trifoliata 107
Heteropogon sp. 221
Hibiscus spp.
H. acetosella 64
H. berberidifolius 64
H. cannabinus 36, 47, 64
H. diversifolius 64
H. noldae 64
H. rostelatus 64
H. sabdariffa 64
H. sparseaculeatus 64
H. surattensis 64
hidana see Dioscorea abyssinica
hoiriada see Sorghum bicolor
holy basil see Ocimum basilicum
Hordeum vulgare
breeding 206, 208-9, 313
characteristics 304-6
conservation 205, 230, 240, 263, 266
crop production 209, 212, 214, 303
diversity 24-5, 71, 131-8
376
Index
Hordeum vulgare (cont.)
history of cultivation 131, 143-4
resource value 361-2
role in Konso agriculture 173, 182
use 52
yields 307, 309, 310-12
horse bean see Vicia faba
horse-radish tree see Moringa oleifera
humer see Tamarindus indica
hunsi 118
hyacinth bean see Lablab purpureus
Hygenia abyssinica 36
Hypagophytum abyssinicum 78
Hyparrhenia spp. 179, 279
H. hirta 279
Hypericum spp. 86
H. quartinianum 107
Hyphaene spp.
H. dankaliensis 68
H. nodularia 68
H. thebaica 52, 68
iffaya see Ocimum
imbus see Allophyllus abyssinicus
incense
history of use 114-15
production 116-17
incense tree see Boswellia rivae
inch'orre see Morus mesozygia
India and the history of crop movemeni
164-6
Eleusine africana 161-2
Pennisetum americanum 162-3
Sorghum bicolor 155-7, 163-4
Indian long pepper see Piper longum
Indian millet see Sorghum bicolor
Indian mustard see Brassica juncea
Indian turnip see Arisaema
Indigofera spp. 60, 219, 221, 285
I. arrecta 47, 60
I. articulata 60, 283
I. coerulea 60
I. tinctoria 48, 60
I. trigonelloides 77
inginkada see Ximonia coffra
inkoy see Ximenia americana
insilal see Anethum graveolens
insilal see Pimpinella anisum
International Board for Plant Genetic
Resources 208
International Crops Research Institute
for the Semi-Arid Tropics 208
International Livestock Centre for Afria
conservation work 218, 220
forage evaluation work 280-6
introduced species 78
Ipomoea batatas 175, 215, 240
Irish potato see Solarium tuberosum
isozyme studies 254
itan zaf see Boswellia
itse faris see Cannabis sativa
Jacaranda spp. 345
jack bean see Canavalia ensiformis
Jatropha curcas 107
jib see Heteromorpha trifoliata
jirjir see Eruca sativa
Juniperus procera 77, 86, 88, 96, 118
jute see Corchorus oligatorus
jute see Hibiscus cannabinus
kaba see Triticum durum
kabudeida see Rhus natalensis
kaguta see Adenia ellenbeckii
kajeta see Eragrostis tef
Kalahari floristic province 76, 77
kalala see Stephania abyssinica
Kalanchoe spp.
K. lanceolata 107
K. marmorata 107
kamun see Cuminum cyminum
kapa see Triticum durum
karbaricho see Echinops
karya see Capsicum
kasse see Lippia javanica
kasse see Ocimum ladiense
kechemo see Myrsine africana
keelo see Sorghum bicolor
kei shinkurt see Allium cepa
kelawa see Maesa lanceolata
ken dara see Sorghum bicolor
kenaf see Hibiscus cannabinus
kenenta 178
kentela see Portulaca oleracea
kerbs see myrrh
keret see Osyris compressa
kestenitcha see Asparagus
ketema see Schefflera abyssinica
ketetina see Verbascum sinaiticum
khat see Catha edulis
kidney bean see Phaseolus vulgaris
kil see Lagenaria siceraria
kimbilota see Solanum incanum
kitgn ayfere see Sorghum spp.
koba see Ensete ventricosum
kogata see Digera alternifolia
kok see Prunus persica
kokora see Terminalia macroptera
K'olla agro-climatic belt 84
Konso
geography 169
people 169-70
plant genetic resources 172-80
system of agriculture 153, 170-2
korch see Erythrina brucei
korroda see Pergularia daemia
koseret see Ocimum
kosheshila see Acanthus
377
Index
koso see Hygenia abyssinica
kota hari see Dioscorea bulbifera
kulbabita 176
kulsida see Sorghum bicolor
kundo-berbere see Piper nigrum
kuni see Cyperus bulbosus
kutata 176
Lablab purpureus
conservation 240, 263
diversity 331
origins 34, 60
role as forage 219, 221, 223
role in Konso agriculture 182
use 48, 174
ladies fingers see Abelmoschus esculentus
ladybird beetle larva see Chnootriba similis
Lagenaria spp. 240
L. abyssinica 58
L. siceraria 48, 58, 107, 179, 184
lakha see Hyphaene thebaica
lameeta see Arisaema
Lamiaceae 49, 50, 61-2
Lathyrus spp.
L. aphaca 60
L. odoratus 60
L. pratensis 60
L. sativus
conservation 205, 240, 263
diversity 34, 331
origins 60, 338
use 48
L. sphaericus 60
Launaea taraxacifolia 175
Lawsonia inermis 107
leaf blotch see Septoria tritici
leaf rust see Puccinia recondita
legumes
conservation 6, 218-19, 263
diversity 278-9
forage value 280-2, 366
Leguminosae 59-61
lemon grass see Cymbopogon citratus
Lens spp.
L. culinaris
conservation 205, 230, 240, 263, 266
diversity 331
origins 33-4, 60, 336
resource value 364
role in Konso agriculture 182
use 174
L. ervoides 60
lentil see Lens culinaris
Lepidium spp.
L. alpigenum 55
L. armoracia 55
L. divaricatum 55
L. intermedium 55
L. sativum 36, 55, 107, 124, 240
Lepidotrichilia volkensii 96
Leucaena leucocephala 97
lia see Terminalia brownii
libania 117
lima bean see Phaseolus lunatus
lime see Citrus aurantifolia
Linaceae 48, 62
Linociera giordanii 96
linseed see Linum usitatissimum
Linum spp. 10
L. bienne 62
L. holstii 62
L. keniense 62
L. strictum 62
L. trigynum 62
L. usitatissumum
conservation 205, 230, 240, 263, 266,
346
diversity 31, 37, 62
domestication and cultivation 144,
209, 212, 350
resource value 365
use 48, 176, 347
Lippia spp.
L. abyssinica 104
L. javanica 124
lomi see Citrus aurantifolia
longa see Colocasia esculenta
loomet see Citrus aurantifolia
loose smut 25, 306
Lotus sp. 221, 224
Luffa spp.
L. cylindrica 48, 58-9
L. echinata 58-9
Lupinus spp. 48, 60, 221, 240
L. a/fcws 48, 60, 339-40
L. mutabilis 341
L. princei 60
L. termzs 60
Lycopersicon esculentum 175
lysine levels, selection for 309, 319
Lythraceae 49, 62
Macrotyloma spp. 223
M. axillare 223, 283
Madagascan floristic province 76,
77
maderta 178
Maesa lanceolata 345
magaloda see Sorghum bicolor
mai-sendedo see Salvia schimperi
maize see Zea mays
Malva verticillata 111
Malvaceae 45, 47, 62-4, 177
Manihot esculenta 175
marasisa see Clerodendrum alatum
marchuke see Sorghum spp.
markets, role in conservation of 197
mashila see Sorghum bicolor
378
Index
Medicago spp. 77-8, 219, 221
M. sativa 241
medicinal plants 36, 43, 104-12, 347
Mediterranean floristic province 77S
Melia azedarach 97, 345
Melinis minutiflora 219, 283-4, 366
melon see Cucumis melo
Mentha spp. 124
mereita see Portulaca quadrifida
Meriandra bengalensis 48, 62
merkuz see Heteromorpha trifoliata
Metaphycus helvolus 65
metbesha see Rosmarinus officinalis
millet see Sorghum bicolor
Millettia ferruginea 86
Mimusops kummel 96
minerals in health care 104
misirich see Clerodendrum alatum
misketi 117
mitin chito 119
mitmita see Capsicum annuum
Momordica spp.
M. balsamina 59
M. charantia 59
M. foetida 107
monocotyledons, diversity of 67-72
mooz see Musa
Moraceae 47, 64
Moringa spp.
M. oleifera 48, 64
M. peregrina 64
M. stenopetala
conservation 215, 241
diversity 36, 64
role in Konso agriculture 183-4
use 48, 175
Moringaceae 48, 64
Morus mesozygia 177
Mucana spp.
M. melanocarpa 60
M. pruriens 48, 60, 341
mulberry see Morus mesozygia
mung bean see Phaseolus radiata
murganta see Vangueria madagascariensis
murukruk see Vernonia hymnolepis
Musa spp. 52, 69
M. paradisiaca 177
Musaceae 52, 69
museta see Musa
mustard see Brassica nigra
Myrica salcifolia 345
Myristica fragrans 124
myrrh 114-16
Myrsine africana 108, 241
Myrtus communis 120, 124
nana see Mentha
narcotics from plants 104, 178
Nasturtium officinale 49, 55
national yield trials 9
nearest neighbour analysis in crop
evaluation 272
nech azmud (netch azmud) see Carum
copticum, Trachyspermum ammi
nech krinfud see Hedychium spicatum
nech see Artemisia rehan
neem see Melia azedarach
neeqayta 174
Neotonia spp. 219, 221, 223
N. wightii 223, 279, 283, 285
net blotch 25, 306
netch shinkurt see Allium sativum
Nicotiana tabacum 178, 184, 241
Nigella sativa
diversity 36, 65
role in conservation programmes 123,
125, 126-7, 130, 241
use 49
niger seed see Guizotia abyssinica
nihba see Meriandra bengalensis
Nile River, role in agricultural
development of 143
njannja see Lycopersicon esculentum
noog (noug) see Guizotia abyssinica
nutmeg see Myristica fragrans
nuts, conservation of 195-7
o jara see Sorghum bicolor
oats see Avena abyssinica
obiyada see Sorghum bicolor
Ocimum spp. 120, 176, 241
O. basilicum 49, 61, 120, 123, 125, 1289,130
O. canum 61
O. forskolei 61
O. gratissimum 49, 61
O. jamesii 61
O. ladiense 120
O. lamiifolium 61
O. sacrum 120
O. spicatum 61
O. stirbeyi 61
O. suave 61, 108
O. trichodon 61
O. urticifolium 61
Ocotea kenyensis 86, 96
ohota see Cajanus cajan
ohota see Vigna unguiculata
oil crops
conservation 6, 209, 216, 263, 345-6
crop development 348-52
diversity 30-2
origins 344
resource value 364-5
role in Konso agriculture 177
uses 346-7
okala see Lablab purpureus, Vigna
unguiculata
Index
okra see Abelmoschus esculentus
Olacaceae 177
Olea spp. 88
O. africana 49, 77
O. europea 49, 65, 96, 119
O. hochstetteri 86, 96
O. welwitschii 86, 96
Oleaceae 49, 65
olive black scale see Saisetia oleae
olive see Olea europea
ongo see Sorghum bicolor
onion see Allium cepa
Opuntia ficus-indica 177
orange see Citrus sinensis
Origanum mayorana 62
oromo dinich see Plectranthus edulis
Oryza spp. 241
O. barthii 30, 71
O. longistaminata 30, 71, 215
O. Sato 12, 30, 52, 71
Osyris compressa 108
Otostegia spp.
O. integrifolia 119
O. steudneri 119
Oxytenanthera abyssinica 241
pakana see Araceae a/so Arisaema
Palmae see Arecaceae
Panicum spp. 221, 223
P. maximum 77, 279
pansala see Sauromatum nubicum
papaya see Carica papaya
papayata see Carica papaya
pareja see Eueusine coracana
Parkinsonia aculeata 97
parpara see Capsicum annuum
pasa see Amaranthus caudatus
Passifloraceae 175
paza see Zea mays
pea see Pisum sativum
peach see Prunus persica
peanut see Arachis hypogaea
pearl millet see Pennisetum glaucum
Pedaliaceae 50, 65
Pennisetum spp. 221
P. americanum
cultivation methods 203-4
domestication 162-3
origins 29, 37, 52, 71
P. clandestinum 219
P. glaucum 29, 52, 71, 204, 263
P. typhoides 241
pepper tree see Schinus molle
perfume plants 114r-21
Pergularia daemia 176
Phalaris arundinacea 279
Phaseolus spp. 205, 241, 263, 331, 336-7
P. coccineus 341
P. lunatus 174, 182, 341
379
P. radiata 50, 61, 182, 341
P. vulgaris 49, 174
Phoenix spp.
P. abyssinica 68
P. dactylifera 52, 68
P. reclinata 52, 68
Phytolacca dodecandra 108, 205, 241
pi jita see Sorghum bicolor
pigeon pea see Cajanus cajan
Pimpinella anisum 123, 128, 130, 241
Pinus spp.
P. patula 97
P. radiata 97
Piper spp.
P. guineense 65,129
P. longum 123, 125, 129
P. nigrum 49, 65, 124, 241
Piperaceae 49, 65
Pistacia spp.
P. aethiopica 44, 49
P. falcata 44
P. vera 44
Pisum spp.
P. abyssinicum 334
P. sativum
conservation 205, 230, 241, 263, 266
crop production 209, 213
diversity 331
origins 33, 60, 334
resource value 364
use 49, 174
Pittosporum mani 345
Plant Genetic Resources Centre/Ethiopia
conservation facilities 226-7
conservation systems 228, 229-34
data management 239-43
documentation systems 235-9
exploration and collecting work 204-10
germplasm characterization 262-4
germplasm multiplication 259-60
improvement trials
oilseeds 346
pulses 329-30
wheat 301-2
objectives 4
role in forest conservation 98
yield trials work 266
Plantaginaceae 49, 65
Plantago spp.
P. afra 49, 65
P. lanceolata 104
P. psyllium 65
Plectranthus spp.
P. edulis 36, 49, 61-2, 215, 241
P. esculentus 61-2
P. punctatus 61-2
Plumbago zeylanicum 108
Poaceae 51, 52, 69-71
Podocarpus spp. 88
380
Index
Podocarpus spp. {cont.)
P. gracilior 77, 96
pogoloda see Zea mays
Polygala aethiopica 77
Polygonum barbatum 108
Polyscias spp. 86
P. fulva 96
pomegranate see Punica granatum
poorta see Hordeum vulgare
Portulaca spp.
P. oleracea 108
P. quadrifida 176
potato round cyst 255-6
potota see Cucurbita pepo
powdery mildew see Erysiphe graminis
pre-breeding techniques 251-2
prickly pear see Opuntia ficus-indica
Prosopis spp.
P. juliflora 97
P. tamarugo 97
Protea 86
Prunus spp.
P. africana 96
P. persica 49, 66
Psudarthia sp. 221
Psophocarpus palustris 341
psyllium see Plantago afra
Puccinia spp. 25, 27, 290, 296, 299
P. glumarum 290
P. graminis 290
P. recondita 25, 290
P. striiformis 296
pulses
conservation 209, 214, 329-30
diversity 32-4, 331
origins 332^40
resource value 363-4
role in Konso agriculture 173-4, 182
pumpkin see Cucurbita pepo
Punica spp.
P. granatum 62
P. protopunica 62
punitta see Coffea arabica
Pygeum spp. 88
P. africanum 96
qaara see Capsicum annuum
Ramularia 352
Ranunculaceae 49, 65
Ranunculus multifidus 111
rape seed 263, 349
Raphanus sativus 241
rasota 176
red-hot-poker tree see Erythrina abyssinica
rejum genbo see Sorghum spp.
rereda see Sorghum bicolor
resins 43
Rhamnaceae 50, 51, 65-6
Rhamnus spp.
R. prinoides
conservation 123, 125, 129, 130
diversity 36, 65—6
use 50, 176
R. staddo 65-6
Rhopalosiphum maydis 292
Rhus natalensis 177
Rhynchosia sp. 221
rice see Oryza sativa
Ricinus communis
conservation 205, 241, 261, 263, 266,
346
crop development 352-3
disease resistance 10
diversity 31-2
oil content 345
origins 59
use 50, 177, 347
roka see Tamarindus indica
roman see Punica granatum
root crops
conservation 193-4, 214-16
diversity 35-6
rooz see Oryza sativa
Rosa abyssinica 77
Rosaceae 49, 66
Rosmarinus officinalis 62, 124
Rubia spp.
JR. cordifolia 108
R. nervosus 108
Rubiaceae 46, 66, 177
rue see Ruta chalepensis
Rumex spp.
R. abyssinica 241
R. bequaertii 111
rust see Puccinia
Ruta chalepensis 50, 60, 123, 128, 130, 177,
241
Rutaceae 46, 50, 66-7
Saccharum officinarum 177
safflower see Carthamus tinctorius
saganeida see Amorphophallus abyssinicus
sage see Meriandra bengalensis
Saisetia oleae 65
Salvadora persica 345
Salvia spp. 62
S. nilotica 50, 62
S. schimperi 50, 62
Sapium ellipticum 86, 345
Satureja sp. 50, 62, 77-8
S. biflora 62
Sauromatum nubicum 36, 175, 215
savannah distribution 84-5
savory see Satureja sp.
scald resistance 25, 306
scarlet runner bean see Phaseolus coccineus
Schefflera spp.
Index
S. abyssinica 86, 345
S. volkensii 96
Schinus molle 97, 345
Scorpiurus 77-8
Securidaca longipedunculata 109
seed conservation
methods 190-3
PGRC/E system 229-34
semat 119
senafetch (senafich, senafitch, senafichi)
see Brassica nigra
senar see Avena abyssinica
sendo see Trema guineensis
Senna alexandrina 50, 59
Septoria spp. 25, 27, 290, 296, 300, 350
S. nodorum 290
S.tritici 290, 300,306
sereti see Asparagus
sesame see Sesamum indicum
Sesamia epunotifera 292
Sesamum indicum
conservation 205, 230, 241, 263, 346
crop development 265, 266, 351
diversity 31, 37, 65
use 50, 347
S. latifolium 65
Sesbania spp. 97, 220, 221, 223
S. sesban 285, 286
Setaria spp. 219, 221
S. sphacelata 279
shallot see Allium cepa
shelagda see Moringa stenopetala
shiferaw see Moringa stenopetala
shimbira see Cicer arietinum
shinet see Myrica salcifolia
snootily resistance 26, 306
shrubland distribution 85
shufun see Sorghum spp.
Silene 77S
Silybum marianum 78
sinde lemine see Sorghum spp.
sindi see Triticum aestivum
sir bizu see Thalictrum rhynchocarpum
sirota see Lens culinaris
Snowdenia polystachya 42
sodan apple see Solanum incamum
Soil Conservation and Community
Forestry Development
Department 97-8
soil management techniques 145-6
Solanaceae 46, 67
Solanum spp. 67
S. incamum 179, 241
S. melongena 67
S. tuberosum 67, 175
Solenostemon sp., S. rotundifolius 62
Somalia-Masai floristic province 76, 77
sonkara see Saccharum officinarum
sono see Senna alexandrina
381
Sorghum spp.
characteristics 316-18
germplasm utilization 318-21
resource evaluation 362-3
role in plant economy 315-16
S. aethiopicum 25
S. arundinaceum 25, 71, 149-50
S. bicolor
conservation 205, 230, 241, 263, 266
crop production methods 209, 211,
212
development 149-53
diversity 25-6, 37, 71
domestication history 163-4
origins 147-9
pest resistance 10
resource value 362—3
role in Konso agriculture 173, 180-2
spread of cultivation 153-7
use 52
S. caffrorum 152-3
S. caudatum 25, 148, 151-2
S. coriaceum 152-3
S. durra 25, 148, 150-1
S. guinea 25, 148, 149-50
S. roxburghii 150
soya bean see Glycine max
Sphenostylis stenocarpa 36
spices
conservation 6
diversity 36
origins 123-4
role in Konso agriculture 176-7, 184
use 125-9
Spilanthes mauritiana 109
spot blotch 25
stalk borer 320
State Forests Conservation and
Development Department 97-8
stem borer 292
stem rust see Puccinia graminis
Stephania abyssinica 109
steppe distribution 84
Sterculia africana 345
stinking smut see Tilletia
Striga resistance 320
stripe mosaic virus resistance 25
stripe rust see Puccinia glumarum
Stylosanthes spp. 221, 223
S. fruticosa 219, 223, 283. 285
Sudan floristic province 76, 77-8
suf see Carthamus tinctorius
sufeta see Helianthus annuus
sugar cane see Saccharum officinarum
sulida see Sorghum bicolor
sumpura see Cicer arietinum
sunflower see Helianthus annuus
sweet basil see Ocimum basilicum
sweet potato see Ipomoea batatas
382
Index
sword bean see Canavalia ensiformis
Syzygium spp. 86
S. aromaticum 124
S. guineense 96, 109
talpa see Linum usitatissimum
Tamarindus spp. 223, 241
T. indica 50, 59, 109, 124
tampota see Nicotiana tabacum
taro see Colocasia esculenta
tarwi see Lupinus mutabilis
taxonomy, role in conservation of 252-6
tebetebkush see Cyphostema niveum
Teclea nobilis 86
teemahada see Catha edulis
teff (tef) see Eragrostis tef
tej sar see Cymbopogon citratus
telba see Linum usitatissimum
telenji see Achyranthes aspera
tellakata see Moringa stenopetala
temer see Phoenix dactylifera
tenaddam (tena-addam) see Ruta
chalepensis
Tephrosia sp. 221
Teramnus sp. 221
Terminalia spp. 77
T. brownii 95, 179
T. macroptera 345
Thalictrum rhynchocarpum 109
thatching grass see Hyparrhenia
Thymus spp. 50, 62
T. schimperi 62
T. serrulatus 62
tibichota see Coriandrum sativum
tikil gomen see Brassica oleracea
tikur azmud see Nigella sativa
tikur see Artemisia renan
Tilaceae 46, 67
Tilletia spp. 25, 290
timber production potential 96
timiz see Piper longum
tinassa see Solanum tuberosum
tinjut see Otostegia integrifolia
tinkish see Sorghum spp.
tisgara see Sorghum bicolor
tit see Gossypium herbaceum
titu see Kalanchoe marmorata
tobacco see Nicotiana tabacum
tobiawu see Calotropis procera
tomato see Lycopersicon esculentum
tosign see Thymus schimperi
tossin see Thymus
Trachyspermum ammi 50, 54, 123, 125, 127,
130
trees
leguminous 285
role in Konso agriculture 184-5
Trema guineensis 345
Trichilia spp. 86
T. roka 345
Trifolium spp.
conservation 219, 220, 221, 222, 224
diversity 43, 77-8
forage evaluation 279, 280-2, 285
T. burchellianum 282
T. cryptopodium 282
T. decorum 281, 282
T. quartinianum 281, 282
T. rueppellianum 281, 282
T. semipilosum 279, 282
T. steudeneri 281, 282
T. tembense 281, 282
Trigonella foenum-graecum
conservation 205, 215, 230, 241, 263
diversity 34, 331
origins 337-8
role in spice cultivation 123, 125, 126,
130
Triticum spp.
conservation 205, 230, 241, 263
crop production 209, 210-13
diversity 26-8, 37, 71
resource value 360-1
role in Konso agriculture 182
T. abyssinicum 289
T. aestivum 52, 289, 300
T. boeoticum 27, 253
T. diococcum 27, 28, 289, 299
T. durum
breeding 296-8
characteristics 289-90, 291
disease resistance 10, 264, 290-2, 299
diversity 26-7
improvement experiments 292-3
role in diet 288-9
use 53, 173
T. monococcum 28, 253
T. polonicum 27, 53, 289
T. pyramidale 289
T. spelta 53
T. turgidum 27, 28, 53, 289, 299
T. vulgare 27, 28
tuber crops
conservation 193-4, 214-16
diversity 35-6
role in Konso agriculture 174r-5, 182-3
tukur azmud see Nigella sativa
tult see Rubia cordifolia
tuma ata see Allium sativum
tuma tima see Allium cepa
tungung 120
turmeric see Curcuma longa
Umbelliferae 44
Uwada see Sorghum bicolor
Vangueria madagascariensis 177
383
Index
vegetables, role in Konso agriculture of
175-6, 184
velvet bean see Mucuna pruriens
Verbascum sinaiticum 109
Verbena officinalis 109
Vernonia spp. 241
V. amygdalina 109
V. galatneisis 42
V. hymnolepis 110
viability testing 232-3
Vicia spp. 221, 282
V. faba
conservation 205, 230, 241, 261, 263,
266
crop production 209, 213
origin 32-3, 60, 331, 332
use 50, 174
V. hirsuta 60
V. paucifolia 60
V. villosa 60
Vigna spp. 221, 223
V. radiata 50, 61, 341
V. subterranea 341
V. unguiculata
conservation 241, 263
origin 34, 61, 338-9
role in Konso agriculture 182
use 50, 174
Voandzeia subterranea 241, 341
Warburgia ugandensis 96,110
water management techniques 145-6
watercress see Nasturtium officinale
watermelon see Citrullus lanatus
weira see Olea europea
wetet begunche see Sorghum spp.
weybata see Terminalia brozvnii
Weyna-Dega agro-climatic belt 84
weyra see Olea europea
wheat aphid 292
wheat see Triticum spp.
white lupin see Lupinus albus
wild gene pools 13, 42-72
wild rice see Oryza longistaminata
Withania somenifera 110
wof aybelash see Sorghum spp.
wollamo gomen see Brassica oleracea
woodland distribution 85
wunsi 118
Wurch agro-climatic belt 84
xagalaa 176
Xanthomonas translucens 292
Ximenia americana 345
Ximonia coffra 177
yam bean see Sphenostylis stenocarpa
yam see Dioscorea alata
ye-aden chiraro 119
ye-jima inchet 119
yedoda see Sorghum bicolor
yeheb nut see Cordeauxia edulis
yellow dwarf virus 25, 254-5, 306
yellow rust see Puccinia striiformis
yemdirimbway see Cucumis aculeatus
yemeder herbere see Spilanthes mauritiana
yeshet ehil see Sorghum spp.
yield trials 266
Zea mays 29,173, 203, 241, 253
Zehneria scabra 110
zengada see Sorghum bicolor
Zingiber officinale 53,124, 241
Zingiberaceae 53, 71-2
zinjibi see Zingiber officinale
Ziziphus spp.
Z. abyssinica 66
Z. hamur 66
Z. jujuba 66
Z. mauritiana 66
Z. mucronata 66
Z. spina-christi 51, 66, 177
Zornia sp. 221, 223, 284, 366