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Indigenous t ees of Ethiopia:
Biolog , uses and propagation techniques
Legesse Negash
Addis Ababa Universit , Facult of Science
Indigenous trees of Ethiopia: --- 1
INDIGENOUS TREES OF ETHIOPIA:

BIOLOGY, USES AND
PROPAGATION TECHNIQUES
SECOND EDITION (2021)
Cover picture: Mature, yellow fruits of Podocarpus falcatus (Thunb.) Mirb.

on the female tree’s live branches. Data provided in Figure 1.8 of the
present book were obtained from these and other fruits of the same
tree, which had mast-fruited during the time of fruit collection.
2 Biology, uses and propagation techniques
Indigenous trees of Ethiopia:

Biology, uses and propagation
techniques
Legesse Negash
Department of Biology
Addis Ababa, Ethiopia
Printed by the SLU (Sveriges

Lantbruksuniversitet) Reprocentralen
Umeå, Sweden, 1995.
Legesse Negash
Department of Biology
P.O. Box 1176
Addis Ababa, Ethiopia
ISBN 91-7191-105-7
© 2021, 1995 Legesse Negash

Department of Biology, Faculty of
Science
All rights reserved

Dedicated to our children who must not inherit a

deserted landscape only to starve in it and succumb to it
Foreword
One need not visit Ethiopia for long before one realizes that
one of the main problems of the country is the destruction of its forests.
Only some 8-10 decades ago, approximately 40% of Ethiopia was
covered by forests that were composed of various indigenous tree
species including podo, various acacias, the African pencil cedar, wild
olive, as well as many other tree species. But now the cover of real
forest is less than 3%, and a substantial proportion of this consists of
trees of foreign origin, including the various Eucalyptus species.
The importance of forests can hardly be overemphasized.

The wood from the trees is used for fuel (either directly or after
processing to charcoal), for the construction of houses, for furniture, as
well as for other innumerable utilities. On top of this, many of the
indigenous trees have medicinal uses.
But forests do not consist of trees only. They are homes for
many other interdependent organisms, which cannot survive without
the forests, and without which the forests cannot survive in the absence
of significant human intervention. In a natural podo forest, for
instance, animals like the Colobus monkey eat the fruits and process
the seeds in such a way that they may germinate. Once the forest has
been reduced to a few scattered trees, the animals are gone, and
germination becomes more difficult or impossible.
Hidden from our view, below the ground in the forests, are
other important organisms. Mycorrhizal fungi, specific for each tree
species, help the trees acquire the nutrients they need. When the last
tree of a species is gone from an area, its fungus will soon disappear
from the soil, and will not return by itself.
Forests are important not only for the products that can be
harvested from them and for the complex interactions they make with
other organisms to build up and/or maintain the complex fabric of
biodiversity, but also for preventing erosion and for affecting the
climate in a positive way. Moreover, deep roots of many tree species
‘fetch’ nutrients from great depths, nutrients that can later be used by
other members of the biosphere. Trees provide shade and variation to
the landscape, and could assume economic importance also from the
perspective of ecotourism.
The value of trees was appreciated long ago and, as their

number started to dwindle, farsighted persons started afforestation
programs. Not surprisingly, trees of foreign origin were chosen on
grounds that they are easy to establish and are capable of growing fast.
However, not all of them gave the anticipated results. Already there are
signs of tree die-backs, as well as occurrence of adverse effects on the
natural ecosystem. Some of these problems are mentioned by the
author in Chapter I of his book.
In retrospect, we must regret that the indigenous trees were

neglected --- to the point that the survival of some of them are now at
stake and, as a consequence of the interdependence, the survival of
those other organisms that have co-evolved with these natural forests.
The main reason that indigenous trees were not used in the
early afforestation projects is that their seeds do not germinate readily,
and that the seedlings are often not easy to establish. However, this is
not always true since it is shown in this book that there are useful
indigenous tree species whose seedlings can easily be raised from
seeds.
Ethiopia is one of the main centers of species diversity of the

world. A large number of plant and animal species are dependent on
the continued existence of indigenous forests. Although the main
responsibility resides with the Ethiopian communities themselves,
these indigenous forests constitute a heritage of all humankind since
we are all indigenous to planet earth.
Fortunately the author of this book, Dr. Legesse Negash,

through a pioneering research conducted at the Addis Ababa
University, has now solved some of the problems and worked out
methods for the rapid propagation of some important indigenous trees,
including Podocarpus falcatus (Thunb.) Mirb.
To transmit this new knowledge to the wider communities is

the responsibility of those concerned with the rapid disappearance of
not only native trees and shrubs, but also water, soils and biodiversity.
The completion of this book is an important part of this process, and it
is my hope that it will reach a wide readership.
Lars Olof Björn (October, 1995)

Professor of Botany at the University of Lund, Sweden
Member of the Royal Swedish Academy of Sciences
Acknowledgments
The Swedish Agency for Research Cooperation with
Developing Countries (SAREC, through the Ethiopian Science and
Technology Commission), the International Foundation for Science,
the African Academy of Sciences, as well as the Addis Ababa University,
are gratefully acknowledged for supporting research on indigenous
trees of Ethiopia. The printing cost for the book was covered by SAREC.
The author is grateful to Prof. L.O. Björn of the University of

Lund (Department of Plant Physiology), for supporting the idea of
research on indigenous forests of Ethiopia and for participating in one
of the capacity building processes in the Department of Biology
(Faculty of Science, Addis Ababa University). The author is thankful to
Prof. Per Christer Odén of the Swedish University of Agricultural
Sciences (Department of Silviculture, Umeå, Sweden) for facilitating
my two and a half months' stay in his laboratory. The time (August to
mid-October, 1995) was used for both the finalization of the book, as
well as for performing some experiments on seed viability of
Podocarpus falcatus (Thunb.) Mirb.
I thank the following persons for reading one or more of the

draft chapters of the book: Prof. L.O. Björn, Dr. Mona Gussarsson, Dr.
C.J. Geldenhuys, Dr. P.J. Wood, Ato Kassa Semagn and Ato Demel
Teketay. I am also grateful to W/ro Selome Bekele for preparing the
final pages of the book.
Many thanks to my students (both graduates and

undergraduates), as well as to my laboratory and greenhouse
technicians, with whom I have shared some of my excitements about
indigenous forests of Ethiopia. I am indeed grateful to them. Also, the
assistance and information obtained from some of the field-based
young Ethiopian forest technicians, as well as the local communities of
the various regions, are gratefully acknowledged.
My wife, W/ro Tsehaynesh Messele, made the final touch on

the proof-reading of the book.
Responsibility for any errors rests entirely with the author

who shall greatly appreciate if such errors are pointed out, and if
suggestions are forwarded to him on how the book might be improved
and/or simplified.
Legesse Negash
Department of Biology (Faculty of Science)
Addis Ababa University (Addis Ababa, Ethiopia)
About the book

Chapters: The book is conveniently divided into eleven
chapters. Each chapter starts with a title and (except for
Chapter I which is rather general) proceeds to
describe the species, discusses its general biological
information (especially its reproductive biology and
physiology), and provides various propagation
techniques for the specific species treated in the book.
Legends: Are often sufficiently explanatory and are

provided just under the pictures or illustrations.
Digressions:Are made when the need for a brief

discussion on certain relevant aspects of a specific
section is felt; e.g., comments on wind pollination,
some chemical/medicinal properties of the species (see
Chapters V and VI), as well as discussions on
biological nitrogen fixation and seed dormancy
(Chapter VII). This is achieved with as much
simplified way as possible.
Glossary: A glossary of some of the technical

terms/phrases used in the book is provided towards the
end of the book.
References: A list of relevant references is provided at the

end of the book.
Contents
Foreword ...................................................................................... 5
Acknowledgements ...................................................................... 7
About the book ............................................................................ 8
Table of contents ......................................................................... 9
CHAPTER I: General ............................................................... 11

Indigenous forests of Ethiopia ......................... 11
The problems .................................................... 15
The aftermaths.................................................. 16
Role of indigenous trees of Ethiopia................ 17
Research on indigenous tree of Ethiopia ........ 26
Concluding remarks ......................................... 36
CHAPTER II: Podocarpus falcatus ..............................................

Description, biology and uses ..............................
Propagation through seeds ..................................
Vegetative propagation .........................................
CHAPTER III: Olea europaea subsp. cuspidata ..........................

Description, biology and uses ..............................
Propagation .....................................................
CHAPTER IV: Cordia africana ....................................................

Description, biology and uses ............................
Propagation ..........................................................
CHAPTER V: Hagenia abyssinica ................................................

Wind pollination ...................................................
Propagation ..........................................................
CHAPTER VI: Millettia ferruginea ..............................................

Propagation ..........................................................
CHAPTER VII: Acacia abyssinica ...............................................

Nitrogen fixation ..................................................
Seed dormancy......................................................
Propagation ..........................................................
CHAPTER VIII: Faidherbia albida .............................................

Propagation ..........................................................
CHAPTER IX: Juniperus procera ................................................
Propagation ..........................................................
CHAPTER X: Dovyalis abyssinica ...............................................

Propagation ..........................................................
CHAPTER XI: Rhamnus prinoides ...............................................

Propagation ..........................................................
Glossary of scientific terms and phrases ................................. 39

References .................................................................................. 47
Index ...............................................................................................
CHAPTER I: General
Citation: Legesse Negash (1995). Indigenous trees of Ethiopia: Biology, uses and
propagation techniques. Printed by the SLU Reprocentralen, Umeå, Sweden.
ISBN 91-7191-105-7. Pp. 11-38
Indigenous forests of Ethiopia
Indigenous forests of Ethiopia (e.g., Figure 1.1) have relentlessly

been deforested (e.g., Figure 1.2) for decades, leaving behind large tracts of
highlands, highland slopes and watersheds unprotected. Consequently, massive
soil erosion, and hence soil nutrient depletion, have occurred (Figure 1.3)
resulting in widespread nutrient deficiency diseases in crop plants, animals, and
humans.
Figure 1.1 An example of indigenous forests from south-eastern

Ethiopia. The picture was taken in May 1990 from an area nearby Kibre-
Menghist (close to the main road, as the town is approached from Addis
Ababa). The greyish crowns are those of Pouteria adolfi-friederici.
Figure 1.2 An example of natural forest deforestation in Ethiopia,

which has been going on for decades. This picture was taken in 1990 from the
vicinity of Kibre-Menghist, close to the area shown in Figure 1.1.
Figure 1.3 The consequence of deforesting the mountain chains

surrounding Addis Ababa. Every year and during the heavy rainy season,
Qebena ‘river’ (one of the highly polluted ‘rivers’ flowing through the city of
Addis Ababa, and barely noticeable during the dry season) swells up with thick
soil solution. Through such seasonal flash floods that occur throughout the
country, Ethiopia loses a huge productive capacity on a regular basis. In most
countries of the world, erosion is localized, but here in Ethiopia it appears that
the whole country is being eroded. The photograph was taken during Ethiopia’s
big rainy season of July, 1989.
Two of the major categories of reasons for the rapid depletion of

native forests have been:
 Land clearing for farming, tree felling for fuel, commercial logging
for timber (see Figure 1.2), tree cutting for house construction, as well as
forest fires; and,
 Difficulty in propagating indigenous forest species through the

conventional tree propagation techniques.
Seeds of many native tree species [e.g., Pouteria adolfi-friederici

(Engl.) Robyns & Gilbert, Podocarpus falcatus (Thunb.) Mirb. (synonym:
Podocarpus gracilior Pilg.), Ekebergia capensis Sparrm., Olea europaea L.
subsp. cuspidata (Wall. ex DC.) Ciffieri (synonym: Olea africana Mill.),
Juniperus procera Endl., Hagenia abyssinica (Bruce) J. F. Gmel.] have either
low percentage germination (which is the result of seed dormancy) or short
viability period (which is caused by seed recalcitrance). Consequently, any
approach in the battle for the regaining of the lost grounds should, among other
things, include the development and/or optimization of techniques for the rapid
propagation of native tree species of Ethiopia.
The two major objectives of this book are:
 To introduce and discuss the biology, physiology, uses, as well as the

propagation techniques, of some economically useful indigenous tree
species of Ethiopia; and,
 To make research results available in as much simplified way as

possible.
In this introductory chapter of the book:
 Problems relevant to the discussion on indigenous forest species of

Ethiopia will be highlighted; and,
 Some examples of ongoing research on indigenous tree species of

Ethiopia in the Department of Biology, Addis Ababa University, will be
presented and discussed.
The problems
It is now about twenty years since scientists and world leaders started
to regard tropical deforestation as one of the Earth's most serious
environmental hazards (emphasis by the present author). This view is fully
justified when reported estimates on the rate of tropical deforestation are
increasing from time to time. Needless to mention, several years have now
elapsed since a reported desertification rate of as large as 6 million hectares per
year has been haunting the minds of many environmental scientists, as well as
the concerned national and international organizations.
Bates (1987) indicated that, in contrast to the past when the world
may have lost one species every millennium, ‘today’ (i.e., in 1987), the world,
on average, is losing at least one species every few hours to ‘development’,
disease or to other causes. Consequently, more than 60,000 species of
organisms will be extinct in the life-time of ‘today's' child (i.e., the child of
1987).
A. W. Wilson (quoted in SPORE, 1994) observed that 17,000
tropical plant and animal species disappear each year. Also, reports by various
environmental institutions warn that, by the year 2000 (in just about five years
after the publication of this book!), between 25,000 and 75,000 species of plants
and animals of the world will be extinct.
According to Janzen (1988), efforts to protect some of the relatively

intact portions of the tropical nature comes too late and too slow for well over
half of the tropics.
The aftermaths
Mathews (1990) wrote: "...An important paradox to bear in mind

when examining natural resource trends is that the so-called non-renewable
resources --- such as coal, oil and minerals --- are in fact inexhaustible, while
the so-called renewable resources such as forests can be finite. As a non-
renewable resource becomes scarce and more expensive, demand falls, and
substitutes and alternative technologies appear. For that reason, we will never
pump the last barrel of oil or anything close to it. On the other hand, a fishery
fished beyond a certain point will not recover; a species driven to extinction will
not reappear; and, eroded top soils cannot be replaced (except over geological
times)...." Mathews further continues to elaborate: "...Nature's bill is presented
in many different forms: the cost of commercial fertilizer needed to replenish
[the] once naturally fertile soils; the expenses of dredging rivers that flood their
banks because of soil erosion [which took place] hundreds of kilometers
upstream; the crop failure due to indiscriminate use of pesticides that
inadvertently kill insect pollinators; or the price of worsening pollution, once
filtered from the air by vegetation...". Mathews concluded that it is the zooming
population that lies at the core of most environmental problems (emphasis
by the present author).
Continued decimation of natural forests results in losses of huge

number of plant and animal species (thus seriously diminishing the base of
genetic diversity); it also unleashes soil erosion, increases frequencies of
drought and flooding, degrades water-quality, decreases agricultural
productivity, and, as a result, leads to increased misery and poverty of rural
communities. The massive energy deficit that plagues Ethiopia (as well as other
under-developed nations), coupled with the strong need for financial gains made
through sell of timber, constitute some of the strongest motive forces for
ruthless deforestation.
Pimentel et al. (1987) emphasized that the most fundamental cause of
deforestation and land degradation is the clearing and planting of land to feed
the growing population. Consequently, the magnitude of land destruction had
become alarming, affecting about 35% of the earth's land surface (Mabbutt,
1984, in Pimentel et al., 1987).
In today's Ethiopia, there are fewer medicinal plant species, fewer

nitrogen-fixing plant species and fewer wild plants than there were, say, 50
years ago. As we destroyed more and more indigenous nitrogen-fixing trees,
fewer and fewer of them remained for fixing molecular nitrogen from the
atmosphere into the soil, thus resulting in the impoverishment of the latter. We
now realize that the consequences of soil impoverishment are directly linked to
the amount of money we invest for purchasing nitrogenous fertilizers or, if
money is not available, on the widespread crop failures as a result of nutrient
deficiency diseases in crop plants.
Role of indigenous forests in resource capture and

transformation
Indigenous forests capture radiant energy that comes from the sun.
This radiant energy is converted into chemical energy. Collectively considered,
indigenous forests can be regarded as a gigantic photocell which is capable of
trapping electromagnetic radiation that comes from the sun.
However, the functions achieved by a tree (which is part of the forest)

and by a photocell are incomparable. The role of a photocell is to convert one
form of energy (e.g., the sun's radiant energy) into another form (e.g.,
electricity), with an efficiency of only 15% (UNESCO, 1981). [Note: This
percentage efficiency of the photocell may have been increased by now].
Compared to the photocell, however, a tree performs a variety of
unique functions. For example, a healthy tree containing healthy leaves:
 Traps radiant energy of the sun and then converts it into chemical
energy (with a minimum efficiency of 40%, i.e., of the photons absorbed
by the chlorophyll-protein complex, 40% are effectively converted into

chemical energy. In fact, the efficiency of photoactivation and electron
transport during the primary act of photosynthesis is about 64%; Lawlor,
1987);
 Splits water molecules into oxygen molecules and protons (of which
the former is a gas in the redox processes of organisms, including
humans);
 Synthesizes ATP (Adenosine Triphosphate) from the proton gradient

created between the thylakoid lumen and the stroma of the chloroplast;
 Synthesizes a variety of biologically active molecules such as

vitamins, proteins, carbohydrates, and several secondary products
(including medicines!).
Moreover, again collectively viewed, forests can be considered as

giant sponges, absorbing and storing the stormy rains of the rainy seasons and
releasing them as cool springs and rivers during the hot and dry seasons that
often characterize Ethiopia (see Figure 1.4). More often than not, this vital role
of forests is forgotten when the role of the same in ecosystem-conservation is
discussed in the Ethiopian context.
Glen W. Russ (an American forester who lived and worked in

Ethiopia between 1940 and 1947, and whose reports were compiled by Wolde-
Michael Kelecha in 1979) wrote:
"...The officials of Ethiopia do not seem to realize that the major value of
their forests lies in their stream- and water-conservation qualities rather
than in their commercial uses, and that they should gauge their
administrative policies accordingly. There are large areas of the country
now almost uninhabited, at least partly because of the lack of perennial
water supply, which were obviously once forested and were deforested by
a population which could live there [and] then. Even a bush cover at the
headwaters of a stream may be sufficient to prevent the flash run off of the
very seasonal precipitation, and of far greater economic value to the
country as a whole than the like amount of land under cultivation or
pasture, since each acre here may affect the productivity of hundreds of
acres below. The people of Ethiopia appear to live on an economy and
philosophy of the present, but if they do not soon realize how important
their forests are to their economy and well-being, they will inevitably find
themselves in a very unpleasant situation."
On another page of his report, Russ further reiterated:
"...The greatest value of the forests is in their water-conserving capacities

and unless they are retained, Ethiopia will become a pauper nation
(emphasis by the present author), since her entire economy is based upon
agriculture."
These statements were written in 1944, and, in the opinion of the

present author, there is nothing wrong with them even after they have become
more than half a century old.
The question of water in Ethiopia has remained a question of

survival. Even large cities are now finding it difficult to supply their growing
inhabitants with sufficient amount of clean water simply because the water-
shades of these cities are not properly forested with the right type of
indigenous forest species.
Figure 1.4 One of the most precious gifts of natural forests resides in
their bounteous provision of clean springs and rivers. This is a photograph of a
small river taken in 1991 from Wedo-Ghenet (southern Ethiopia). This segment
of the river is close to the piped water which is used by the community for
recreational purposes. The question is: Would the next or the following
generation get the opportunity to see this river?
The mountain-chain overlooking Addis Ababa on the north-eastern

and the north-western flanks of the city (e.g., Yeka and Intoto), which once
served as a source of numerous clean springs, has now become almost a menace
to the city. Flash floods during the rainy seasons have become common, thus
destroying the upstream landscape and silting the downstream city areas (see the
flood in Figure 1.3).
The inhabitants of the city of Gonder and the peasants farming the
surrounding water-shades appear to have grasped the crux of the problem ---
that rehabilitating the water-shades with vegetation is essential for solving the
chronic problem of water shortage that has been plaguing the city since a long
time. Various localities of the Oromo region have now started to officially
protest about the chronic water shortages. These shortages can directly be linked
to the continued destruction of the water-shades during the last several decades.
Clearly, the government of Ethiopia is obliged to assist these communities
(through crafting smart policies) to re-forest their vital watersheds.
For ages, these and other Ethiopian communities depended on natural

springs and rivers that are produced and maintained by natural forest systems
(see Figure 1.4). As these spring- and river-systems diminished in capacity, the
communities which depended on them were forced to look for subterranean
water systems; and looking for subterranean water systems to get clean water on
a massive scale requires not only economic might but also technical capability.
Moreover, as the water tables sink down due to overuse, the finance and the
technical capability needed for overcoming the gravitational force (for drawing
up the underground water) become more and more difficult to attain. It must be
remembered that “The source of all water is rain”, and that rain is harvested
by native forests.
An important question to be asked is, therefore, how far are we

prepared to bear the consequences of this suicidal act of forest destruction?
What alternatives do we have at hand?
Maser (1991) remarked:
"...if society learned anything from the decade of the 1960's, it is that one
cannot unilaterally destroy the ‘establishment’ without offering a viable
alternative to replace it. Before an old paradigm can be cast out, there must
be a new one to take its place."
Plochmann (1989, quoted in Maser, 1991) underscored that forestry solely

oriented towards the maximization of profit can no longer meet the expectations
and needs of the society.
These two environmentalists seem to be pointing to the grave

problems faced by today's Ethiopia: massive destruction of native forest species
followed by (when loan money is available from World Bank) their replacement
by temperate conifers and eucalypt trees. The alternative is neither viable (e.g.,
the prevalence of die-backs in pines and cypresses after some 30 years of
reforestation efforts: see Figures 1.5 and 1.6) nor ecologically acceptable (e.g.,
the negative impacts of the various Eucalyptus species on water, soil, and
biodiversity resources: see Figure 1.7).
Figure 1.5 Dead and dying group of young cypress trees (Cupressus
lusitanica Miller), commonly known as cedro blanco or Mexican white cedar.
The picture was taken in July 1991 from the compound of Sholla milk factory in
Addis Ababa. There are a number of reasons for the mal-adaptation of the
species in Ethiopia, including soil factors, insects, and (in some regions) water
stress. Ethiopia has been spending hundreds of millions of Birr on the
reforestation of its deforested areas with exotic trees such as C. lusitanica only
to find out that these trees produce allelopathic chemicals (hence are inimical
to plant biodiversity: see also Figure 1.7) and that there is heavy tree mortality
(see Figures 1.5 and 1.6).
Figure 1.6 A dead specimen of Pinus radiata D. Don (the Monterey

pine). The picture was taken in May 1989 from within the Science Faculty of
the Addis Ababa University, located just behind the Freshman Building.
Figure 1.7 Introduced to Ethiopia in 1895, eucalypt trees have been widely
planted in rural settings, as well as in urban centers and the corresponding peripheral
areas. As can be seen from the picture (taken in 1993 from Arssi region), there is no
herbaceous or shrub flora beneath the trees of this stand. This is due mainly to the
potent allelochemicals produced by the leaves of these trees. As a result, a eucalypt
stand is not only inimical to biodiversity, but it is also not suitable for erosion
control (clearly seen from the exposed root system of individual trees, where these
have been uncovered by soil erosion). Eucalypt trees are also notorious in guzzling up
water from the ground and depleting soils of their nutrients since they grow all year
round. Many forest technicians in Ethiopia ‘passionately’ argue for eucalypt trees,
citing their fast growth and ‘economic benefits’. However, the purported ‘economic
benefits’ are not for the impoverished rural communities, but for the merchants who
ferry eucalypt poles to urban centers using the 1950s/1960s Italian lorries, which by the
way are notoriously polluting. But what is ‘economic benefit’ which makes sense only
to the few when food and water security are at stake for the majority? When
biodiversity is threatened? When the planting of eucalypt trees is raging like a wildfire,
taking over agricultural lands, drying up springs and wetlands?
Research on indigenous tree species of Ethiopia
It was against the above, rather daunting, challenges that research on

the propagation biology and physiology of indigenous tree species of Ethiopia
was pioneered in 1988 in the Department of Biology, Addis Ababa University.
As has been emphasized above, three of the most important challenges

confronted by indigenous tree species of Ethiopia are:
 Unwise exploitation of these tree species for economic purposes

(Legesse Negash, 1990a,b; see also Figure 1.2);
 Deforestation for the expansion of subsistence or commercial

agriculture; and,
 Lack of scientific knowledge on their reproduction biology and tree

physiology, as well as absence of capable and dedicated institutions
dedicated to the research and development of native trees.
The following are examples of investigations that are currently being

undertaken at the Biology Department of the Faculty of Science (Addis Ababa
University).
Investigations on seed physiology
Many tropical trees produce seeds that are covered with stony or hard
seed coats (e.g., the endocarp of wild olive, the sclerotesta of podo or the hard
seed coat of various acacias) (e.g., Legesse Negash, 1992a, 1993). Under
natural conditions, such a ‘stony’ or hard seed coat requires a set of suitable
environmental/physico-chemical conditions and/or biological actions that
modify the structure of its seed coat in such a way that imbibition by the seed is
facilitated.
Under natural conditions, important environmental factors for seed

germination include: oxygen, adequate moisture, optimum temperature,
‘activation’ light (for some plant species), as well as various types of fungi
and/or other microorganisms that are needed for the modification of the seed
coat. After the seeds have germinated, they may require shade, especially during
the initial moments of their life cycle, in order to protect themselves from the
harmful ultraviolet radiation, wind, low relative humidity, various weeds,
herbivores, as well as other detrimental environmental factors. It must be
remembered that protective shade is available only when there is a vegetation
system that is not seriously disturbed.
Interestingly, certain tree seeds [e.g., Pouteria adolfi-friederici (Engl)

Baehni.; Podocarpus falcatus (Thunb.) Mirb. (Synonym: Podocarpus gracilior
Pilg.)] undergo significant decline in viability with time if stored at room
temperature or are mishandled during the process of fruit collection and
extraction. Such seeds are categorized as either recalcitrant (P. adolfi-
friederici) or semi-recalcitrant (P. falcatus).
Recently, an in vitro method (giving a gain in time of 600%, and an

increase in the percentage germination of 400%) has been developed for the
rapid germination of P. falcatus seeds (Legesse Negash, 1992a; see also
Chapter II of the present book). Using this method, it is now possible to raise
large numbers of seedlings in the nursery within a relatively short period of
time.
Figure 1.8 shows trends in viability of P. falcatus seeds, studied over

a period of 12 months.
Figure 1.8 Trends in the viability of Podocarpus falcatus seeds. Mature,

yellow fruits (see Figure 2.2) were collected from relatively young and mast-fruited
trees, which were flanked by a number of male trees. The fruits were stored at room
temperature for 1 month (empty circles), 2 months (solid triangles), 3 months (empty
triangles), 6 months (inverted solid triangles) and 12 months (inverted empty triangles).
Seed pretreatments were performed according to the method described by Legesse
Negash (1992a). Seeds obtained from mature and partially green fruits, stored for 4
months at room temperature and under moist conditions resulted in close to 100%
germination (uppermost solid circles). Bars indicate + SE (standard errors of the
means). Where no bar is indicated, SE is smaller than the symbol.
Vegetative propagation of Podocarpus falcatus
To complement the in vitro seed germination method for the rapid

propagation of P. falcatus, an alternative procedure is now described by Kassa
Semagn and Legesse Negash (1995) by means of which P. falcatus can be
propagated through rooting of branch cuttings. This method is feasible at even
low technical level and is relatively inexpensive.
The results of the investigation showed that when cuttings were taken
from juvenile saplings (about 3-5 years old stock plants), up to 90% of the
cuttings developed roots and grew into normal plants. It has also been found
that when cuttings of appropriate size were taken from juvenile source plants,
adventitious roots were satisfactorily developed even without the application of
any plant hormone.
Comparison of the growth rates of seedlings and stecklings (i. e.,

seedlings derived from rooted cuttings) indicated that the latter grew much
faster than the former. However, stecklings developed plagiotropically due to
the synthesis of lignin lagging behind the plant’s growth rate.
Since these comparative experiments were performed in the nursery

of the Biology Department, further assessments on the growth rates of seedlings
and stecklings in the field (and at the appropriate ecological conditions) are
required to arrive at valid conclusions on the merits and demerits of propagating
P. falcatus by vegetative means.
Propagation using plant tissue culture techniques
The present level of plant biotechnology is such that many plant

species that have conventionally been regarded as "difficult" to germinate might
be propagated through the multiplication of suitable explants (e.g., through
axillary bud induction), induction of adventitious meristems (either through the
process of organogenesis or somatic embryogenesis), as well as through
multiplication of calli derived from organs, tissues, cells or protoplasts (Dixon,
1985; Bonga and Durzan, 1987).
Using some of the techniques of plant tissue culture, we were able to

optimize the chemical and physical conditions for axillary bud induction in
Cordia africana Lam. and Hagenia abyssinica.
However, the present author is prompted to caution that it is very

important to be careful in resource allocation on a tissue culture laboratory for

the following three major reasons:
 Tissue culture techniques require skilled manpower and relatively

expensive chemicals and pieces of equipment;
 Woody plants may not readily respond to rooting treatments;
 There is a high degree of somaclonal variation among plantlets

derived from calli; and,
 Plant tissue culture might be powerful when it is coupled with plant

cell genetic engineering. This is because, the expertise and finance
invested on culturing an engineered plant cell, with an eye to getting a
novel plant variety, is far more rewarding than that invested on the
propagation of an already existing plant species through tissue culture,
especially when such a species is known to be a good seeder.
Some observations on physiological responses of some selected native

tree species
Photoinhibition and photooxidation in Podocarpus falcatus
Damage to the photosynthetic apparatus can be caused by either

ultraviolet radiation or visible radiation or by the interaction between the two
spectra (Powels, 1984).
Photoinhibition can be broadly defined as the reduction in the

photosynthetic capacity (independently of gross changes in pigment
concentrations) induced by exposure to visible light (Powels, 1984). In this
deleterious effect of light, Photosystem II is believed to be the site for
photoinhibition (Cleland and Critchley, 1985; Arntz and Trebst, 1986).
Photooxidation, on the other hand, results when plants are exposed to

strong light for a relatively prolonged period of time. During photooxidation,
photodestruction of the photosynthetic pigments occurs (Powels, 1984).
Forest technicians in Ethiopia have been frustrated by the low

percentage of established seedlings of P. falcatus when planted under field
conditions. To begin with, seeds of this tree species were ‘difficult’ to germinate
(i.e., from 0% germination to 1-20% germination) (Legesse Negash, 1992a).
Secondly, the relatively tiny seedlings that were planted in an open field failed
to survive, especially in areas where the cloud cover was minimal and,
consequently, the intensity of light was high. The phenomenon of
photooxidation in P. falcatus and its reversal through the provision of shade has
been noted (Legesse Negash, unpublished observations).
The ‘mystery’ of adaptation in Hagenia abyssinica
Many plants are capable of protecting themselves from the harmful

ultraviolet radiation (UV) by synthesizing flavonoids which selectively filter
UV (Warner and Caldwell, 1983; Wellmann, 1983; Legesse Negash, 1988).
Together with cuticular waxes and other cell-wall components, these
flavonoids attenuate the incident UV radiation by one or two orders of
magnitude (Robberecht and Caldwell, 1978).
Before its decline to the present status, Hagenia abyssinica used to

thrive on the Ethiopian highlands that have altitudinal range of between 2,450
and 3,250 m (Hedberg, 1989). The level of ultraviolet radiation at such altitudes
is rather high (Björn and Murphy, 1985; Legesse Negash and Björn, 1986).
When young trees of H. abyssinica were grown in the glasshouse

where there is almost no UV, they were ‘colorless’ (Legesse Negash,
unpublished observations). When these same young trees were transferred to an
open field where they were exposed to full sunlight, they developed pink
coloration within five to eight days. The pink coloration is due to the synthesis
of flavonoids.
Flavonoids absorb (and hence dissipate) the harmful ultraviolet

radiation, a radiation which at present is threatening the globe because of the
depletion of the ozone layer. Such a rapid response to UV may be one of the
explanations for the adaptation and survival of H. abyssinica on the Ethiopian
highlands.
Indeed, many other indigenous plant species have a variety of

mechanisms for coping with a variety of tropical environmental factors. The
question is, therefore, who warranted us to destroy trees like H. abyssinica
which, not only are our heritages, but are also the products of millions of years
of continuous evolution?
Discovery of midday stomatal closure in Cordia africana
Stomata (Figure 1.9) play a key role in the control of the balance
between water loss and carbon gain. An effective control of this balance is often
reflected in a net biomass production of an individual plant species (Beadle et
al., 1985). In situations where it is important to maximize water-use- efficiency,
investigations on stomatal physiology as related to CO2- and water-vapor-
transfer between the atmosphere and the internal tissue of the leaf are important
(Fischer and Turner, 1978).
Figure 1.9 A typical dicot stoma (plural stomata) from the abaxial
surface of a cucumber leaf The stoma is constituted of its two guard cells that
are ‘fused’ at the polar ends of the pore (courtesy of John Troughton;
reproduced from Frank. B. Salisbury and Cleon W. Ross, Plant Physiology,
1985).
There are a number of factors that affect the physiology of stomata.

These include: relative humidity and plant water status, light (both the
photosynthetically active radiation and ultraviolet radiation, UV), carbon
dioxide, temperature, pathogenic agents, as well as gaseous pollutants.
The effect of UV on both the integrated responses of stomata of

Eragrostis tef (Zucc.) Trotter and ionic relations of guard cell membranes on
Vicia faba L. have been reported by Legesse Negash and Björn (1986), Legesse
Negash (1987), Legesse Negash, Jensén and Björn (1987), Legesse Negash and
Björn (1988), Legesse Negash (1988).
In many parts of Ethiopia, water and nutrient stresses (along with

excess light that contains a lot of UV), are some of the most important limiting
factors that affect plant development and productivity. For example, it is very
difficult to grow coffee plants (Coffea arabica L.) in water and nutrient stressed
regions, as well as in an open field where these plants are exposed to full
sunlight. Similarly, seedlings of Podocarpus falcatus find it very difficult to
establish, grow and develop in open (and degraded) landscapes where there is a
lot of sunlight but limited moisture.
Two contrasting sample data on the effect of increasing water stress,

measured as the daily time courses of stomatal resistances in two different tree
species (Cordia africana Lam. and Pouteria adolfi-friederici (Engl) Baehni.),
are presented in Figures 1.10a and 1.10b).
In Figure 1.10a, daily time courses of stomatal resistance in C.

africana (measured over a period of eight days), are shown as individual
parallelograms for each of the days during the water stress period. There were
two distinct groups of patterns. The first group of patterns (days 1 to 3 on the X-
axis of the graph) was observed between one and three days after watering the
plants with adequate and equal amounts of water. The second group of patterns
(days 5 and 8 on the graph) was observed at days five and eight after
withholding the water.
Figure 1.10 Daily time courses of stomatal resistance in Cordia africana

(a), and Pouteria adolfi-friederici (b), measured over a period of eight days.
In the first group of patterns of Figure 1.10a (i.e., days 1 to 3),

stomata of C. africana closed during the early mornings of the days (7:00 h) and
during the late afternoons (18:00 h). Stomata were wide open between 8:00 and
14:00 h of the days. In contrast, in the second group of patterns stomata closed
vigorously between 10:00 and 16:00 h of the day.
Midday stomatal closure, which is one of the plant’s physiological

mechanisms for responding to increasing water stress, is especially clear in the
parallelogram which corresponds to day eight after withholding water from the
plants.
Figure 1.10b shows stomatal resistances patterns for Pouteria adolfi-

friederici. There was no clear evidence for the presence of midday stomatal
resistances on days 5 and 8 in this species, although resistances between 10:00
and 16:00 h increased moderately during these two days. The early mornings’
and late afternoons’ stomatal resistant were consistently high throughout the
study period.
It is concluded that the extent of stomatal resistances developed

around noon in C. africana (with increasing water stress) are far greater than
those developed in A. adolfi-friederici. Hence, this study discovered for the first
time the presence of midday stomatal closure in Cordia africana.
Details of the methodology, as well as the discussion on the

underlying mechanisms that bring about midday stomatal closure, and the
relevance of the latter to drought-tolerance, were published by Legesse Negash
(1992b).
Concluding remarks
From the foregoing discussion, it is apparent that there are a lot of

challenges and opportunities presented by native trees of Ethiopia. The
challenges require systematic research designs, implemented with vigorous

research and development efforts, with an eye to reversing the alarming trends
in the deforestation of indigenous tree species. The opportunities present the
limitless possibilities of discovering new biological traits (including those
related to medicine) in the diverse native trees and shrubs that are out there.
It is, therefore, suggested that these challenges be tackled in earnest,

and opportunities be harnessed to extent possible so as to generate new
knowledge before native plants are driven to extinction, and the country is taken
over by invasive alien plant species.
Ethiopia should therefore focus on:
 The development and/or optimization of techniques by which its

indigenous forest species can rapidly be propagated and restored;
 The proper investigations on the factors which determine the

distribution and productivity of these tree species, thus
characterizing the specific ecological requirements of the various
species;
 The protection of indigenous tree species both from ruthless

exploitation and from the ‘aggressive’ and/or invasive exotic tree
and shrub species. These efforts must be coupled with the efforts
for seeking solutions for the pervasive ‘energy famine’ through (for
example) developing hydropower, solar devices and geothermal
sources; and, finally,
 The proper perception and evaluation of the linkages (and hence the
compatibility) of the growing human population with the available
(i.e., finite) natural resources. It must be remembered that no
amount of effort and investment are adequate to ease the increasing
pressure on tropical vegetation unless population growth (of both

humans and domestic animals) is controlled. After all: “Life
cheapens in its own abundance”.
Glossary of scientific terms and phrases

[Note: Gk. Greek; L. Latin]
Abscissic acid (L. abscissus, to cut off) - A plant hormone that, e. g., causes
dormancy in buds, maintains dormancy in seeds, brings about stomatal
closing, has effects on fruit growth and development, promotes
resistance to injury.
Anther (Gk. anthos, flower) - A structure that bears the pollen and is located on
the tip of a stamen.
Arelous - A space marked out on a surface; used to describe the raised area on
the surface of seeds of the family Fabaceae/Leguminosae, subfamily
Mimosoideae.
Axillary - A term used to specify buds, fruits, flowers, cones, etc. that occur in
the axils of leaves.
Biological nitrogen fixation - A process by which the conversion of dinitrogen

molecules (N2) into the biologically useful forms such as ammonia
(NH4+), nitrate (NO3-), etc. is achieved. Biological nitrogen fixation is
largely accomplished by symbiotic and/or free-living prokaryotic
microorganisms.
Calyx (Gk. kalyx, a cup or a husk) - A group of the outermost components of

the flower, the sepals.
Chlorophyll (Gk. chloros, green + phyllon, leaf) - A plant pigment molecule

that confers the characteristic green colour to green plants.
Chlorosis - A deficiency disease symptom in green plants that is manifested

through the development of yellow coloration, usually on leaves.
Chlorosis is a characteristic deficiency symptom of, e. g., nitrogen or
magnesium. Also, many plants can become chlorotic when they are
flooded or are waterlogged. Under waterlogged soil conditions, plant
roots suffer from lack of adequate oxygen, which results in poor nutrient
uptake by the cells of the roots.
Coenzymes - Non-protein, complex organic molecules upon which certain

enzymes depend for their effective catalytic activities.
Completion of a life cycle in plants - Involves the complex processes of seed

germination, growth and development, flowering and, finally, the
successful production of viable seeds.
Cone - A dense and often woody series of bract-like reproductive structures

(sporophylls) arranged along an axis. Cones are characteristic
reproductive structures of many gymnosperms. Note that, although this
definition may be fitting for describing the male cone of Podocarpus
falcatus, it is inadequate for describing the female cone of P. falcatus, in
the opinion of the present author.
Corolla (L. corona, crown) - A group of petals that are usually the coloured
components of the flower and are often located next to the sepals.
Cotyledons (Gk. kotyledon, cup-shaped hollow) - Seed leaves that generally

store food in the dicotyledonous plants and absorb food in the
monocotyledonous plants. They may also act as photosynthetic organs.
Two cotyledons occur in dicotyledonous plants, one in
monocotyledonous plants.
Cuticle - A waxy or fatty layer that usually covers the outer wall of epidermal
cells.
Deciduous (L. decidere, to fall off) - Plant (tree) species that shed their leaves
during a specific season of the year; e. g., Erythrina brucei Schweinf.
emend. Gillett, which sheds its leaves during the dry season.
Dehice (L. dehiscere, to split open) - To open and release spores (by sporangia)
or seeds (by fruits) or pollen grains (by anther) or by any other dispersal
units or structures.
Dicotyledonous (Gk. di, two + kotyledon, cup-shaped hollow) - Plant species

that produce two seed leaves in their seedlings or germinants.
Dioecious (Gk. di, two + oikos, house) - Plant/tree species that have the male
and the female elements on two different individuals of the same
species.
Dormancy (L. dormire, to sleep) - A condition in which seeds or buds fail to

germinate or grow as a result of intrinsic or extrinsic dormancy-causing
factors or chemicals. In the case of seeds, it is the ability to retain
viability while having restricted metabolic activity, without observable

growth and development taking place.
Double dormancy - Dormancy caused by both the seed coat (external

dormancy) and the embryo (internal dormancy). Consequently,
treatments aimed at breaking double dormancy must be applied in
sequence.
Drupe (L. drupa, overripe olive) - A simple fleshy fruit (e. g., as in wild olive
or peach) that is derived from a single carpel and is usually one-seeded
(except in rare cases where ‘twin’ seeds may occur; e. g., in Olea
europaea subspecies cuspidata). The ‘seed coat’ (the endocarp) of a
drupe is usually hard and stony and may be adhered to the seed. In Olea
europaea subsp. cuspidata, there is usually sufficient clearance between
the seed proper and the endocarp. The true seed coat is accessed after
breaking away the endocarp.
Endocarp (Gk. endon, within + karpos, fruit) - The innermost layer of the
ovary wall in a fruit. In wild olive, the endocarp is stony and is one of
the most important factors that seriously hamper successful germination.
Endosperm (Gk. endon, within + sperma, seed) - A tissue that is (usually)

triploid, formed following the fusion of two polar nuclei and a sperm
nucleus in the embryo sac. The endosperm is digested by the growing
sporophyte either before or after the maturation of the seed. Endosperm
is found in the angiosperms only.
Epicotyl (Gk. epi, upon + kotyledon, cup-shaped hollow) - The upper portion of
the axis of an embryo or young seedling, above the point where the
cotyledons (the seed leaves) are attached and below the next leaf or
leaves.
Epigeal (Gk. epi, upon + geo, earth) - When, in a germinating seed, the
cotyledons are raised above the surface of the soil. For example,
germination in Millettia ferruginea is epigeal.
Epimatium - In the context of the present book, it is the fleshy part of the fruit
of P. falcatus, i. e., the pulp of the fruit.
Essential - A term commonly used in the science of plant nutrition for

emphasizing the importance of nutrient elements. The three essentiality
criteria are:
1. The plant cannot complete its life cycle if the nutrient element of
interest is lacking in the soil (or in the experimental system);
2. The nutrient element should be irreplaceable (i. e., there is no other
nutrient element that can exactly replace the role of the nutrient element
in question);
3. The effect of the nutrient elements should be direct. This means that
the element must not be acting by accelerating or retarding the uptake
of some other essential elements or by relieving toxicity caused by still
other elements.
Exfoliation (L. ex, out, from + folium, leaf) - The act of casting or coming off in
scales, laminae, or splinters; scaling or flaking off, e. g., skin, bone,
rock, bark, etc.
Exocarp (Gk. exos, without, outside + karpos, fruit) - The outermost layer of
the mature ovary wall, or pericarp.
Feeder roots - Roots that originate from the main root axis and so are
tributaries to the same.
Female gametophyte/Megagametophyte - The haploid (n) entity formed in

higher plants by the germination of the megaspore; sometimes, as in
many lower plants, a free-living organism, but in seed plants retained on
the sporophyte within the megasporangium or within the ovule. Since
Podocarpus falcatus is recognized as a conifer, the reserve food of its
seed is designated as the female gametophyte.
Flavonoids - Are phenylpropane derivative glycosides. They represent a very

widespread group of water-soluble compounds, and many of them are
brightly coloured. They are commonly red, crimson, purple or yellow.
Flowers owe their colourful appearances to flavonoids.
Habit - The characteristic appearance or form of a tree or any other organism.
Hilum (L. hilum, a trifle) - A scar left on a seed after the separation of the latter
from the funiculus.
Hypocotyl (Gk. hypo, under + kotyledon, cup-shaped hollow) - The portion of

an embryo or a young seedling found between the cotyledons and the
embryonic root, the radicle.
Imbibition - The adsorption of water by hydrophilic surfaces and the swelling-

up of these colloidal surfaces/materials (e. g., starch, protein, clay)

because of the adsorption of water molecules onto the internal surfaces
of the same.
Legume (L. legumen, leguminous plant) - A simple and a dehiscent fruit

composed of two carpels, splitting along two sides (e. g., as in the bean
or Millettia ferruginea or Acacia abyssinica).
Lenticels (L. lenticella, a small lens or window) - A small porous and spongy
region in cork surfaces of stems, roots, and other vascular plant parts
that allow exchange of gases between the internal tissues of the plant
and the atmosphere through the periderm.
Megagametophyte (Gk. mega, large + gamos, marriage + phyton, plant) - The

female gametophyte that produces the egg in heterosporous plants. In
seed plants, the megagametophyte is located within the ovule which is
borne by the sporophyte.
Mesocarp (Gk. mesos, middle, + karpos, fruit) - Layer of the ovary wall in a
fruit that is located between the exocarp and the endocarp. The mesocarp
is usually juicy as in, e. g., the fruits of wild olive.
Monotypic (Gk. monos, single) - A family or genus with a single species.
Nitrogenase - An enzyme complex that catalyses the fixation of N2.

Nitrogenase accepts electrons from reduced ferredoxin or other effective
reducing agents and then catalyses the reduction of N2 into other useful
forms of nitrogen.
Obconic (ob-, a prefix meaning opposite, inverse or against) - A structure that

approximately looks like an inverted cone. The cone is a geometric
structure or solid figure whose bottom is a circle and whose sides taper
evenly up to a point (the apex).
Ovary (L. ovum, egg) - In flowering plants, a closed structure in which one or
more ovules are located.
Ovule (L. ovum, egg) - In seed plants, a female structure consisting of a female
gametophyte enclosed within the megasporangium (the nucellus) which
in turn is covered by one or two protective layers, called integuments.
Panicle - A type of inflorescence with many side branches in which each of the
branches bears two or more flowers. The panicle usually has an

indeterminate axis.
Parthenocarpy (Gk. parthenos, virgin + karpos, fruit) - The development of

fruits without fertilization; parthenocarpic fruits are usually seedless.
Pathogenic (Gk. pathos, suffering, + genesis, beginning) - Disease-causing

organism.
Pedicel (L. pediculus, little foot) - The stalk of an individual flower in an

inflorescence.
Peduncle (L. pedunculus, from pediculus, little foot) - The stalk of an

inflorescence or of a solitary flower.
Petal (Gk. petalon, flower leaf) - One of (the usually) several appendages that
cover the reproductive portions of the flower. When the flower opens,
the petals expand to form delicate, and usually brightly coloured,
structures. Often, petals attract pollinating organisms such as insects.
Phenology - Follow-up of the characteristic duration of e. g., flowering and

fruiting of trees or other plants.
Photoinhibition (Gk. photos, light) - Injurious effects of strong light to the

photosynthetic apparatus of green plants induced by exposure to the
same. Damage to the photosynthetic apparatus may be caused by
wavelengths in the ultraviolet (usually ultraviolet-B) or by light in the
visible part of the spectrum or by the interactions between visible and
ultraviolet radiations.
Photooxidation (Gk. photos, light) - The destruction or the light- and oxygen-
dependent bleaching of pigments as a result of long-term exposure of
plants or photosynthetic organelles to strong light.
Phyllode (Gk. phyllon, leaf) - A flat, expanded, photosynthetic petiole or stem.

In certain genera of vascular plants, phyllodes replace leaf blades as
effective photosynthetic organs.
Phytochrome (Gk. phyton, plant + chroma, colour) - A biliprotein pigment of
plants which controls diverse physiological activities including induction
of seed germination (e. g., germination of lettuce seeds), flowering,
plumular unhooking, de-etiolation, induction of rapid chlorophyll
synthesis, as well as inhibition of hypocotyl elongation.
Pistil (L. pistillum, pestle) - The ovule-forming structure in a flower, typically

consisting of an enlarged basal portion (the ovary) in which the ovules
are located, as well as a surface (the stigma) that is receptive to pollen
and is elevated on a stalk (the style).
Pyrene - A small nut or kernel of a fruit or a drupe.
Recalcitrance - Refers to seeds that are difficult to germinate once the natural
moisture content of the seeds drops below the minimum percentage
required for maintaining the viability of the seeds. This phenomenon is
common in tropical plants. The moisture content of such recalcitrant
trees/plants is usually about 30%.
Relic (In the context of this book) - Remains (of forests, tree species) that have
survived past age destructions. Relic forests or trees remind people of
their past (widespread) occurrence in the region where they are now
observed.
Riparian - Occurring in or inhabiting the banks of rivers, lakes, etc.
Saprophyte (Gk. sapros, rotten, + phyton, plant) - A heterotrophic organism

that derives its food from the dead tissues, as well as from products, of
plants and animals.
Scarification - The process of etching or cutting or softening a seed coat for the
purpose of enhancing percentage germination.
Sceptre - Staff or rod carried by a ruler (a king) as a sign of royal power at, e.
g., a coronation ceremony.
Sclerotesta - Is the woody seed coat of P. falcatus.
Seed coat - The protective outer layer of the seed that develops from the
integuments (i. e., from the outermost layer or layers of tissue
enveloping the nucellus of an ovule).
Stamen (L. stamen, thread) - A pollen-forming structure in flowers. The stamen
usually consists of terminal sporangia that are borne at the ends of
slender stalks (the styles). Sporangia are hollow unicellular or
multicellular structures in which spores are produced.
Steckling - A term that is equivalent to seedling. According to Professor Chris

Bornman of the University of Lund (1987), who himself appears to have

accepted the suggestions made by Dr W.J. Libby, the following terms
are used to distinguish and establish equivalence among the processes of
deriving plants from seeds, from cuttings in vivo and from explants in
vitro:
Seeds germinants seedlings
Cuttings rooted cuttings stecklings
Explants plantlets plantlings
Stigma (L. stigma, spot, mark) - A glandular surface which is terminally located
on the pistil, is receptive to pollen and is capable of stimulating the
formation of pollen tubes.
Stomata (singular, stoma) (Gk. stoma, mouth) - Pores located on the surfaces
of the aerial organs (e. g., leaves) of most higher plants, and used for
controlling the entry of CO2 (into the photosynthetic sites of green
plants) as well as the exit of water vapour and other gases out of the
plant. Stomata are surrounded by a highly specialized group of cells
(called guard cells) that are involved in the formation and control of the
same. The stomatal complex, which includes the pore, the pair of guard
cells, as well as the subsidiary cells (if any) is one of the key structures
that transitioned aquatic plants to land.
Style (Gk. stylos, column) - A slender mass of tissue that originates from the top
region of the ovary, and through which the pollen tube grows.
Suberin - Fatty material that is quite impermeable to water and is found in the
cell walls of cork tissue, in the Casperian strips of the endodermis, as
well as in the seed coat of certain plant/tree species.
Symbiosis (Gk. syn, with + bios, life) - A form of association between two or
more different kinds of organisms. Often, a symbiotic association is
beneficial to both organisms.
Ultraviolet radiation - Short-wavelength radiation that is conveniently divided

into UV-A (ultraviolet-A, 315-400 nm), UV-B (ultraviolet-B, 280-315
nm and UV-C (ultraviolet-C, 250-280 nm). The short-wavelength
components of UV-B and the whole UV-C are harmful to most
organisms.
Viable (L. vita, life) - Is a seed that is capable of germinating and successfully
developing into a plant.
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Indigenous trees of Ethiopia: Biology , uses and propagation techniques

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INDIGENOUS TREES OF ETHIOPIA:

SECOND EDITION (2021)

Cover picture: Mature, yellow fruits of Podocarpus falcatus (Thunb.) Mirb.

Indigenous trees of Ethiopia:

Printed by the SLU (Sveriges

© 2021, 1995 Legesse Negash

All rights reserved

Dedicated to our children who must not inherit a

The importance of forests can hardly be overemphasized.

The value of trees was appreciated long ago and, as their

In retrospect, we must regret that the indigenous trees were

Ethiopia is one of the main centers of species diversity of the

Fortunately the author of this book, Dr. Legesse Negash,

To transmit this new knowledge to the wider communities is

Lars Olof Björn (October, 1995)

The author is grateful to Prof. L.O. Björn of the University of

I thank the following persons for reading one or more of the

Many thanks to my students (both graduates and

My wife, W/ro Tsehaynesh Messele, made the final touch on

Responsibility for any errors rests entirely with the author

About the book

Legends: Are often sufficiently explanatory and are

Digressions:Are made when the need for a brief

Glossary: A glossary of some of the technical

References: A list of relevant references is provided at the

CHAPTER I: General ............................................................... 11

CHAPTER II: Podocarpus falcatus ..............................................

CHAPTER III: Olea europaea subsp. cuspidata ..........................

CHAPTER IV: Cordia africana ....................................................

CHAPTER V: Hagenia abyssinica ................................................

CHAPTER VI: Millettia ferruginea ..............................................

CHAPTER VII: Acacia abyssinica ...............................................

CHAPTER VIII: Faidherbia albida .............................................

CHAPTER X: Dovyalis abyssinica ...............................................

CHAPTER XI: Rhamnus prinoides ...............................................

Glossary of scientific terms and phrases ................................. 39

Indigenous forests of Ethiopia

Indigenous forests of Ethiopia (e.g., Figure 1.1) have relentlessly

Figure 1.1 An example of indigenous forests from south-eastern

Figure 1.2 An example of natural forest deforestation in Ethiopia,

Figure 1.3 The consequence of deforesting the mountain chains

Two of the major categories of reasons for the rapid depletion of

 Difficulty in propagating indigenous forest species through the

Seeds of many native tree species [e.g., Pouteria adolfi-friederici

The two major objectives of this book are:

 To introduce and discuss the biology, physiology, uses, as well as the

 To make research results available in as much simplified way as

In this introductory chapter of the book:

 Problems relevant to the discussion on indigenous forest species of

Ethiopia will be highlighted; and,

 Some examples of ongoing research on indigenous tree species of

According to Janzen (1988), efforts to protect some of the relatively

Mathews (1990) wrote: "...An important paradox to bear in mind

Continued decimation of natural forests results in losses of huge

In today's Ethiopia, there are fewer medicinal plant species, fewer

Role of indigenous forests in resource capture and

However, the functions achieved by a tree (which is part of the forest)

by the chlorophyll-protein complex, 40% are effectively converted into

 Synthesizes ATP (Adenosine Triphosphate) from the proton gradient