Monkey Orange monograph.pdf - Crops for the Future

Monkey orange 

Strychnos cocculoides 

Author: 

Charles K. Mwamba 

Editors: 

J. T. Williams (chief editor) 

R. W. Smith 

N. Haq 

Z. Dunsiger

First published in 2006 by: 

Southampton Centre for Underutilised Crops, University of 

Southampton, Southampton, SO17 1BJ, UK 

© 2006 Southampton Centre for Underutilised Crops 

Printed at RPM Print and Design, West Sussex, UK 

The text in this document may be reproduced free of charge in any 

format or media without requiring specific permission. This is 

subject to the materials not being used in a derogatory manner or in 

a misleading context. The source of the material must be 

acknowledged as [SCUC] copyright and the title of the document 

must be included when being reproduced as part of another 

publication or service. 

Copies of this handbook, as well as an accompanying manual and 

factsheet, can be obtained by writing to the address below: 

International Centre for Underutilized Crops 

@ International Water Management Institute 

127 Sunil Mawatha, Pelawatte, Battaramulla, Sri Lanka 

British Library Catalogue in Publication Data 

Monkey orange 

1. tropical fruit trees 

i Williams ii Smith iii Haq iv Dunsiger 

ISBN 0854328416 

Citation: C. Mwamba (2005) Monkey orange. Strychnos cocculoides. 

Southampton Centre for Underutilised Crops, Southampton, UK.

THE FRUITS FOR THE FUTURE PROJECT 

This publication is an output from a research project funded by the 

United Kingdom Department for International Development (DFID) for 

the benefit of developing countries. The views expressed are not 

necessarily those of DFID [R7187 Forestry Research Programme]. 

The opinions expressed in this book are those of the authors alone and do 

not imply an acceptance or obligation whatsoever on the part of ICUC, 

ICRAF or IPGRI.

ICUC 

The International Centre for Underutilised Crops (ICUC) is an autonomous, 

non-profit, scientific research and training centre. It was established in 1992, 

based at the University of Southampton in the UK. In 2005 ICUC was 

transferred to the International Water Management Institute (IWMI), Sri Lanka. 

The centre was established to address ways to increase the use of underutilised 

crops for food, nutrition, medicinal and industrial products. The enhancement of 

currently underutilised crops is a key to food security, to the conservation of 

biological diversity and to the preservation and restoration of fragile and 

degraded environments throughout the world. 

World Agroforestry Centre 

The World Agroforestry Centre (ICRAF), established in Nairobi in 1977, is an 

autonomous, non-profit research body supported by the Consultative Group on 

International Agricultural Research (CGIAR). ICRAF aims to improve human 

welfare by alleviating poverty, improving food and nutrition security and 

enhancing environmental resistance in the tropics. 

IPGRI 

The International Plant Genetic Resources Institute (IPGRI) is an international 

research institute with a mandate to advance the conservation and use of genetic 

diversity for the well-being of present and future generations. It is also a centre 

of the Consultative Group on International Agricultural Research. 

Also available in this series of monographs: 

Tamarind - Tamarindus indica by H. P. M. Gunasena and A. Hughes (ISBN 

0854327274) 

Ber - Ziziphus mauritiana by O. P. Pareek (ISBN 0854327525) 

Safou - Dacryodes edulis by J. Kengue (ISBN 0854327649) 

Baobab - Adansonia digitata by M. Sidibe and J. T. Williams (ISBN 

0854327762) 

Annona spp. by A. C. de Q. Pinto, M. C. R. Cordeiro, S. R. M. de Andrade, F. 

R. Ferreira, H. A. de C. Filgueiras, R. E. Alves and D. I. Kinpara (ISBN 

0854327851) 

Forthcoming in this series: 

Jackfruit - Artocarpus heterophyllus by N. Haq 

Mangosteen - Garcinia mangostana by M. bin Osman and M. Abd. Rahman 

Ndjanssang – Ricinodendron heudelotii by Z. Tchoundjeu 

Sapote species - Pouteria sapota, P. campechiana, P. viridis by C. Azurdia

TABLE OF CONTENTS 

PREFACE..................................................................................................i 

CHAPTER 1. TAXONOMY.................................................................... 1 

1.1 The family Loganiaceae................................................................. 2 

1.1.1 Loganiaceae s. l....................................................................... 2 

1.1.2 Strychnaceae s. s..................................................................... 3 

1.2 The genus Strychnos L................................................................... 3 

1.3 Species of the genus....................................................................... 5 

1.3.1 S. cocculoides.......................................................................... 6 

1.3.2 S. spinosa ................................................................................ 9 

1.3.2.1 Subspecies S. spinosa ...................................................... 9 

1.3.3 Other edible fruited species .................................................. 10 

1.3.3.1 S. gossweileri Exell ....................................................... 10 

1.3.3.2 S. innocua Del. .............................................................. 11 

1.3.3.3 S. lucens Bak. ................................................................ 11 

1.3.3.4 S. madagascariensis Poir............................................... 12 

1.3.3.5 S. pungens Solored. ....................................................... 12 

1.3.3.6 S. stuhlmannii Gilg ........................................................ 13 

1.4 Vernacular names......................................................................... 13 

1.5 Key to the species ................................................................. 15 

CHAPTER 2. DISTRIBUTION............................................................. 16 

2.1 Regional distribution of S. cocculoides........................................ 16 

2.2 Sampling the distribution patterns................................................ 19 

2.3 Distribution in relation to habitation ............................................ 19 

2.4 Distribution outside Africa........................................................... 19 

CHAPTER 3. PRODUCTION AREAS ................................................. 20 

CHAPTER 4. PROPERTIES AND USES ............................................. 22 

4.1 Fruits for food use ........................................................................ 22 

4.1.1 Fruit composition.................................................................. 22 

4.1.1.1 Pulp................................................................................ 22 

4.1.1.2 Dried fruits..................................................................... 25 

4.1.2 Toxicity of Fruits .................................................................. 25 

4.1.3 Processing............................................................................. 25 

4.2 Seeds ............................................................................................ 26 

4.3 Leaves .......................................................................................... 27 

4.4 Medicinal uses of Strychnos cocculoides..................................... 27

4.4.1 Fruits ..................................................................................... 27 

4.4.2 Leaves................................................................................... 27 

4.4.3 Bark....................................................................................... 27 

4.4.4 Roots ..................................................................................... 27 

4.5 Medicinal uses of other Strychnos species................................... 28 

4.5.1 S. spinosa .............................................................................. 28 

4.5.2 S. innocua.............................................................................. 28 

4.5.3 S. madagascariensis.............................................................. 28 

4.5.4 S. pungens ............................................................................. 28 

4.5.5 Other wild species................................................................. 29 

4.6 Other uses of S. cocculoides......................................................... 29 

4.6.1 Other uses of other Strychnos fruits...................................... 29 

CHAPTER 5. ECOLOGY...................................................................... 42 

5.1 Introduction.................................................................................. 42 

5.2 Rainfall......................................................................................... 44 

5.3 Light............................................................................................. 44 

5.4 Wind............................................................................................. 46 

5.5 Soil requirements ......................................................................... 46 

5.5.1 Details of soil types............................................................... 51 

5.5.1.1 Siliceous Rock Parent Material ..................................... 51 

5.5.1.2 Orthoclase – Feldspathic Rock (Acid Igneous Rock) 

Parent Material .......................................................................... 51 

5.5.1.3 Ferromagnesian Rock Parent Material .......................... 51 

5.5.1.4 Calcareous Rock Parent Material .................................. 51 

5.6 Ecotypic differentiations .............................................................. 52 

5.7 Mycorrhizae ................................................................................. 52 

5.8 Fire ............................................................................................... 54 

CHAPTER 6. AGRONOMY ................................................................. 55 

6.1 Seed propagation.......................................................................... 55 

6.1.1 Seed collection and handling ................................................ 55 

6.1.2 Seed treatment and germination............................................ 56 

6.2 Vegetative propagation ................................................................ 56 

6.2.1 Grafting................................................................................. 57 

6.3 Orchard establishment.................................................................. 57 

6.3.1 Site preparation..................................................................... 57 

6.3.2 Planting................................................................................. 57 

6.3.3 Seedling survival................................................................... 58 

6.3.4 Mycorrhizae.......................................................................... 58 

6.4 Orchard management ................................................................... 58 

6.4.1 Stand density......................................................................... 58

6.4.2 Fertilizers .............................................................................. 58 

6.4.3 Pruning.................................................................................. 59 

6.4.4 Weeding and intercropping................................................... 59 

6.4.5 Protection from pests and diseases ....................................... 60 

6.4.6 Physical stresses.................................................................... 60 

6.4.7 Growth rates.......................................................................... 61 

6.5 Agroforestry ................................................................................. 61 

6.6 Ongoing research ......................................................................... 61 

CHAPTER 7. REPRODUCTION AND HARVEST ............................. 62 

7.1 Reproduction................................................................................ 62 

7.2 Harvesting .................................................................................... 62 

7.2.1 Harvesting practices.............................................................. 62 

7.2.2 Yields.................................................................................... 62 

7.2.3 Post-harvest handling............................................................ 63 

7.3 Processing .................................................................................... 63 

7.4 Economics.................................................................................... 64 

CHAPTER 8. SELECTION AND GENETIC RESOURCES ............... 67 

8.1 Background .................................................................................. 67 

8.2 Surveys......................................................................................... 67 

8.2.1 Botswana............................................................................... 67 

8.2.2 Malawi .................................................................................. 67 

8.2.3 Tanzania................................................................................ 68 

8.2.4 Zambia .................................................................................. 68 

8.2.5 Zimbabwe ............................................................................. 68 

8.3 Summary selection criteria........................................................... 68 

8.4 Germplasm................................................................................... 69 

8.5 Conservation ................................................................................ 71 

8.5.1 Methods of storage................................................................ 71 

8.5.2 In situ conservation............................................................... 71 

CHAPTER 9. POTENTIAL IMPACT AND MARKETING ................ 73 

9.1 Potential for widening the cultivation of S. cocculoides.............. 73 

9.1.1 Educating farmers................................................................. 73 

9.1.2 Marketing potential............................................................... 74 

9.1.3 Pricing................................................................................... 74 

9.1.4 Markets in the region ............................................................ 75 

9.2 Potential for off-farm income based on products other than fruits .. 

……………………………………………………………………….76 

9.2.1 Prospects for extraction of chemicals from African 

Strychnos....................................................................................... 76

9.2.2 Prospects for enhanced uses in local medicine..................... 76 

CHAPTER 10. RESEARCH NEEDS .................................................... 78 

10.1.1 Understanding the genepool ............................................... 78 

10.1.2 Developing technology ....................................................... 78 

10.2 Backing the R&D....................................................................... 79 

10.3 Adding value to certain products ............................................... 79 

10.4 Development of new products.................................................... 79 

APPENDIX I. INSTITUTIONS WITH GERMPLASM OF 

STRYCHNOS COCCULOIDES.............................................................. 81 

APPENDIX II. INSTITUTIONS AND INDIVIDUALS ENGAGED IN 

STRYCHNOS RESEARCH AND DEVELOPMENT ............................ 83 

REFERENCES....................................................................................... 86 

INDEX.................................................................................................... 97

LIST OF TABLES 

Table 1.1 Vernacular names of Strychnos species………………..…… 

Table 2.1 S. cocculoides in countries of Southern Africa ………..….... 

Table 4.1 Physiochemical composition of fruit pulp……………..…… 

Table 4.2 Mineral composition of fruit pulp ……………………......... 

Table 5.1 Principal biomes in which the genus Strychnos 

is endemic, prevailing climate and dominant growth forms………...... 

Table 5.2 The performance of S. cocculoides and 

S. birrea seedlings under different shading regimes……….................. 

Table 5.3 Soil physical requirements of S. cocculoides......................... 

Table 5.4 Stocking densities of S. cocculoides and S. pungens 

on soils with various properties …………………………………….... 

Table 5.5 Soil types that support S. cocculoides and S. spinosa in 

Botswana………………………….………………….……………....... 

Table 5.6 Chemical soil properties that support S. cocculoides 

and S. spinosa in Botswana……………………………...................... 

Table 5.7 Effect of mycorrhizal inoculation on initial growth 

of outplanted indigenous fruit tree seedlings …..…….………………. 

Table 7.1 Percentage of households consuming indigenous fruits as a 

snack or main meal during normal, bumper or disaster harvest seasons 

for maize in two areas in Zimbabwe…………………………………... 

Table 7.2 Domesticated IFT planting dependent on age to maturity, 

yield increase and collection cost……………………………………... 

Table 8.1 Institutional germplasm collections in Africa………….…... 

LIST OF FIGURES 

Figure 1.1 Flowers and leaves of Strychnos cocculoides………..…… 

Figure 1.2 Fruit and leaves of Strychnos cocculoides………………... 

14 

17 

24 

24 

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47 

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49 

50 

53 

65 

66 

70 

6 

8

LIST OF PLATES 

Plate 1. Fruit and spines of monkey orange……...…………………. 

Plate 2. Flowers and rounded leaves of monkey orange……………. 

Plate 3. Mottled unripe fruit of monkey orange…………………….. 

Plate 4. Fissured bark……………………………………………….. 

Plate 5. Variation in fruit of Strychnos species……………………... 

Plate 6. Variation in leaves of Strychnos species……….. ………… 

Plate 7. Brown jelly-like pulp surrounding the seeds ........................ 

Plate 8. Pale seeds of monkey orange…. …………………………... 

Plate 9. Monkey orange height and form at maturity ………………. 

Plate 10. Monkey orange: A pioneer on abandoned cultivation 

sites………………………………………………………………….. 

Plate 11. Deciduousness in monkey orange………………………… 

Plate 12. Monkey orange growing and fruiting on limestone parent 

material …………………………………………………................... 

Plate 13. Grass providing partial shade to young monkey orange 

trees ……………………………………………………………….... 

Plate 14. Sapling on abandoned cultivation site……………….......... 

Plate 15. Effect of fire on saplings …………..................................... 

Plate 16. Coppice sprouts in monkey orange after fire ………..…… 

Plate 17. Grafted plant flowering after one year……………………. 

Plate 18. Coppice shoots …………….……………………………... 

Plate 19. Coarse and fine roots of mycorrhizal root system………... 

Plate 20. Termite damage of a seedling ……………………………. 

Plate 21. Powdery mildew on fruits………………………………… 

Plate 22. Monkey orange chopped for fire wood and fruits scattered 

on the ground………………………………………………………... 

Plate 23. Extraction of roots for medicinal uses……………………. 

Plate 24. Monkey orange and Masuku (Uapaca kirkiana) 

wine…………………………………………………………………... 

30 

30 

31 

31 

32 

32 

33 

33 

34 

34 

35 

35 

36 

36 

37 

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39 

40 

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PREFACE 

Trees which produce edible fruits or seeds are important in tropical 

regions because they supplement and improve the quality of diets. Only a 

limited number have been fully domesticated and improved through 

selection and breeding; although a large number have been domesticated 

and are cultivated locally in traditional cultures. 

Many of the latter species are considered incipient domesticates and they 

remain genetically wild even through they are protected by rural people, 

often around homesteads and their cultivated fields. 

With limited resources for plant breeding in developing countries many 

of the species considered to have potential for focused and relatively 

rapid improvement do not receive the attention they deserve. Plant 

breeders have little choice other than to place their limited resources on 

improving field crops. 

Taking this into account the International Centre for Underutilised Crops 

has developed a series of priority species on which it is felt more effort 

would be justified because it would repay results in generating incomes, 

alleviating poverty and providing more balanced diets. 

In the case of woody fruits of the tropical and subtropical regions there 

are a number of these priority species considered as components of land 

use systems where they can help to stabilise environments in 

agroforestry systems. Strychnos cocculoides is a prime example and this 

monograph attempts to summarise what is currently known about the 

species. 

The preparation and publication of this monograph has been funded by 

the Department for International Development (DFID), UK as part of a 

project entitled “Fruits for the Future”. 

The World Agroforestry Centre (ICRAF) is a partner organisation in this 

endeavour as are numerous national programmes. This book is the 7th in 

a series of monographs; a parallel series of extension manuals is being 

issued. 

i

It will be noted that there are many gaps in our knowledge and further 

basic as well as applied research is needed. It is hoped that making this 

monograph available to teachers, students, extensionists, policy makers, 

growers and others will promote further production and marketing and 

will stimulate scientists to address some of the knowledge gaps. 

We are grateful to the late Dr. C. Mwamba who produced the 

manuscript. We regret that he was not able to see his work published and 

dedicate the book to his memory. We also thank Miss Rosemary Wise 

for the illustration of the plant, and Miss Angela Hughes and Mr. 

Berekhet Berakhy, former staff members of ICUC, for their advice and 

efforts in seeing the manuscript through to finalisation. 

Editors 2005 

ii

CHAPTER 1. TAXONOMY 

Edible fruit-bearing species of Strychnos belong to the family 

Loganiaceae, which includes tree, shrubs and liana species distributed 

throughout the warm tropical and subtropical region of Asia, Africa and 

the Americas and occasionally in the warmer temperate regions. Many 

species of Strychnos produce alkaloids such as strychnine and resinous 

substances such as the South American arrow poison curare. Similarly, 

in Malaysia, species have been used to produce dart poisons. Such 

poisonous properties were alluded to when the name Strychnos was 

coined, recalling the properties of poisonous nightshade, called 

strukhnos in Greek. One species has been domesticated and cultivated in 

several continents: S. nux-vomica L. whose seeds were extracted for 

strychnine, used in medicine since 1640. 

S. cocculoides Baker, the monkey orange, produces a locally traded fruit 

which is especially popular in Eastern and Southern Africa. Another 

African species of monkey orange, S. spinosa Lam, is widely gathered 

throughout Sub-Saharan Africa from drier savannah habitats. 

Few other genera of the family are economically important although 

many are used locally for dyes, medicine and wood. The genus Buddleja 

has provided species traded internationally for ornamental purposes, as 

has Spigelia. Gelsemium has similarly provided ornamentals, such as G. 

sempervirens L. of the Eastern USA but it was probably more important 

as a source of medicinal alkaloids. 

This monograph describes the properties and uses of S. cocculoides 

because it has been widely recognised to be a species worthy of further 

exploitation for its fruit and one recognised as currently neglected 

(Maghembe et al., 1998; Leakey and Newton, 1994). Reference is also 

made to S. spinosa since this has similar recognition (e.g. Haq and 

Atkinson, 1999; a European Union project coordinated by the University 

of Turin, Italy: see http://www.divapra.unito.it/; Maghembe et al., 1998). 

1

1.1 The family Loganiaceae 

The family was originally included in the order Gentianales along with 

Oleaceae, Salvadoraceae, Apocynaceae, Asclepiadaceae and 

Gentianaceae, in the system of Bentham and Hooker. Later many other 

orders were established, that of Loganiales retaining Loganiaceae. 

Hutchinson (1969) then split the Loganiaceae into several families and 

Strychnos with some related genera became Strychnaceae. This split was 

based on long-existing evidence that the original classification was not 

natural. Work by Scott and Brebner (1889) had shown Buddleja was 

different to all the other genera of Loganicaceae in anatomical 

characteristics and some morphological characters; and Spigelia had had 

claim to being part of a separate group since Martius described 

Spigeliaceae in 1827. 

For the purpose of this monograph the short description of the family 

below is for Loganiaceae sensu lato (Hutchinson, 1973). Nonetheless it 

is also helpful to consider the Strychnaceae because, as a family in its 

own right or as a subfamily of Loganiaceae, it contains a limited number 

of genera, all with only 1-5 species, plus Strychnos with about 200 

species distributed through tropical regions. 

1.1.1 Loganiaceae s. l. 

This group includes trees, shrubs, which are often climbing, or herbs. 

Leaves are opposite sometimes whorled, simple, generally connected by 

interpetiolar stipules often much reduced or by a raised line. 

Inflorescence is usually cymose. Flowers are regular, usually 

hermaphrodite, 4-5-merous. Calyx is gamosepulous and usually 

imbricate. Corolla is hypogynous, valvate or imbricate. Stamens are 

epipetalous inserted on corolla tube, alternating with the corolla lobes. 

Ovary is free, 2-celled. Ovules are one or more in each cell. Style is 

simple, stigma is capitate or bi-lobed. 

Fruit is a berry, capsule or drupe, seeds with straight embryo in copious 

albumen. 

2

There are up to 35 genera and over 550 species, mostly tropical but some 

warm temperate. 

1.1.2 Strychnaceae s. s. 

This group of genera is typified by possessing indehiscent drupaceous or 

baccate fruits. Leaves are 3-5 nerved from or above the base. Branchlets 

are often armed with spines, or there are tendrils often in the herbaceous 

species. 

There are 3 groups of genera: 

(i) Baccate fruit and entire leaves 3-5 nerved from or above the 

base, Strychnos L. with species spread across most tropical regions. 

(ii) Drupaceous fruit, with pinnately nerved leaves: Couthovia A. 

Gray, 5 species in Malaysia to Polynesia; Crateriphytum Scheff., ex 

Koord., 1 species in Moluccas. 

(iii) Baccate fruit, climbers with pinnately nerved leaves. Gardneria 

Wall, ex Roxb., 3 species in India and Japan; ScyphoStrychnos Moore, 1 

species in Nigeria; Pseudogardneria Raciborski, 2 species in East Asia. 

When Strychnaceae and 5 other families (Potaliaceae, Antoniaceae, 

Spigeliaceae amd Buddleiaceae) were split from Loganiaceae the 

remaining family (Loganiaceae s.s.) was left with 7 genera and over 90 

species (Hutchinson, 1969). 

1.2 The genus Strychnos L. 

(Species Plantarum 1:159:1753) 

The following description is based on Leeuwenberg (1983), Hutchinson 

and Dalziel (1963) and White (1962). 

Trees or shrubs are erect or climbing by hooked tendrils or lianas with 

curled tendrils. Trees are usually less than 10 m tall in savannah or up to 

35 m in forests. Bark is usually thin and smooth, but thick and corky in 

some species; in lianas often with large lenticels. Branches with axillary 

or a terminal straight spine. Branchlets are terete, sometimes sulcate. 

3

Leaves are opposite, sometimes decussate, stipules are reduced to ciliate 

rim joining petiole bases; petioles are inserted on a leaf cushion. 

Laminae are often variable in shape but broadly orbicular to narrowly 

elliptic and mostly coriaceous. Leaf shape varies between plants in shade 

and open sun and for woody species between original growth and regrowth 

after cutting and/or fire. Leaves are entire, rounded and slightly 

mucronate or emarginate to acuminate at apex; glabrous or pubescent. 

Venation with 1, 2 or 3 pairs of secondary veins from the base curving 

along the margins, usually not as far as the apex and anastomosing with 

the other veins; in rare cases pinnately veined. 

Inflorescence is cymose either terminal or axillary or both tending to be 

thyrsoid and 1 to many flowered, lax or congested, simple or condensed, 

shorter or larger than the leaves. Bracts are small or very small. 

Flowers are 4-5-merous, regular or with the sepals unequal. Calyx lobes 

are triangular, ovate, rarely narrow to linear, imbricate; free or connate 

up to half their length and green or coloured with the outside hairy or 

glabrous. 

Corolla is a tube, rotate or sub-campanulate with lobes valvate in bud 

becoming spreading or reflexes or rarely sub-erect; lobes are triangular, 

oblong, entire, acute; whitish to yellow or pale green, sometimes orange 

in colour; sometimes with a corona at the mouth. Corolla on both sides is 

glabrous or hairy but at inner base always glabrous. 

Stamens are exserted or included, inserted on the corolla tube usually 

adnate to corolla throat, rarely low down in tube. Filaments are filiform, 

usually short, anthers ovate to narrowly oblong, cordate, sometimes 

sagittate at the base. 

Ovary is 2-celled (rarely 1-celled due to breakdown and absorption of 

the dividing wall); style is straight; stigma is terminal, usually capitate. 

Fruit is a globose berry usually 2-celled, mostly yellow or reddish when 

mature but occasionally greenish or blue-black; subtended by the 

persistent calyx. Small fruited species usually have thin, soft walls but 

large fruited species have thicker, harder, indehiscent walls. Pulp is 

juicy; seeds variously compressed, globose or oval in shape, embedded 

in the pulp; seeds are 0.5-3.0 cm long with testa thick or membranous; 

embryo is straight. Cotyledon is flat and leafy. 

4

Estimates of number of species vary greatly: 179 (Leeuwenberg, 1983), 

200 (Willis, 1957) or 400 (Hutchinson, 1969). 

1.3 Species of the genus 

It is not known with certitude how many species comprise the genus, 

although large numbers are quoted from the warm tropical and 

subtropical parts of Australia, Asia, Africa and the Americas. In Africa 

there are probably about 75 species. There has been no continent-wide 

assessment of the species and at present information is drawn from 

national and regional floras, hence it is to be expected that a degree of 

synonymy is built into the estimates. The two most important African 

species for fruits are described below. 

5

1.3.1 S. cocculoides 

Synonym: S. tuberosa T. R. Sim 

Figure 1.1 Flowers and leaves of S. cocculoides 

6

A deciduous tree about 5 (1-8) m tall, occasionally semi-lianoid with a 

spreading open crown and one or several trunks, frequently suckering. 

The bark is thick, corky and ridged, brown and not lenticillate. 

Branchlets are dark brown and densely pubescent when young with pale 

longitudinal corky, non-powdery ridges which later coalesce. Spines 

terminate branches, 1.0-1.5 cm long, occasionally curved but usually 

straight. 

Leaves are up to 6 cm long and about 4 cm wide; orbicular to ovateelliptic, 

pubescent but sometimes hairy on both sides. 1-3 pairs of 

secondary veins from the base curved along the margin. 

Flowers are in terminal inflorescences which are thyrsoid i.e. main 

branch racemose and laterals cymose, seemingly umbellate and 

congested. Male and female flowers are borne on the same tree. Sepals 

are greenish, hairy or glabrous about 2 mm long. Corolla is a tube, green 

to orange in colour, always glabrous at the base but may be hairy above, 

lobed with a white penicillate corona at the mouth. Ovary is 2-celled. 

Fruits are 6-12 cm diameter, round; blue-green mottled white when 

young turning mottled green-yellow to yellow orange when ripe with a 

granular skin, hard-shelled (shell about 2-5 mm wide); containing many 

(10-100) subglobose irregularly curved, flattened seeds. The pulp is the 

edible part; seeds are non-toxic. Pulp has a sweet taste. 

It is regarded as semi-cultivated in numerous parts of northern-southern 

Africa but it remains essentially a wild species. 

7

Figure 1.2 Fruit and leaves of S. cocculoides 

8

1.3.2 S. spinosa 

Synonyms: S. buettneri Gilg, S. cardiophylla Gilg, S. arvalhoi Gilg, S. 

courteti Chev., S. cuneifolia Gilg, S. emarginata Bak., S. gracillima 

Gilg, S. harmsii Gilg, S. laxa Solered., S. leiosepala Gilg, S. lohua A. 

Rich., S. radiosperma Gilg, S. rhombifolia Gilg, S. tonga Gilg, S. 

volkensii Gilg. 

A deciduous shrub or tree about 5 (0.5-10) m tall with rounded crown 

from one or several trunks, frequently suckering. The bark is somewhat 

scaly and grey-buff in colour, shallowly fissured, very rarely slightly 

corky, not lenticellate. Branches can be robust or not, sometimes deeply 

ringed at nodes, often with straight or recurved axillary spines. 

Branchlets are hairy or not, often ending in a straight spine. 

Leaves are extremely variable, to 6-10 cm long and about 4 cm wide; 

petioles 2-10 mm; lamina is coriaceous, obovate to suborbicular to 

ovate-elliptic, apex subacute and very shortly cuspidate to retuse 

occasionally acuminate, glabrous or hairy beneath, 1-3 pairs of distinct 

secondary veins from the base curved along the margin. 

Flowers in terminal cymes often looking rare like an umbel, small, 

greenish-white. Sepals connate at the base, 5 mm long, sub-equal, 

pubescent on the outside. Corolla a tube with lobes hardly split and 

hardly spreading with white penicillate corona at the mouth, lobes of 

tube triangular. Ovary 1-celled, stigma oblong. One basal placenta with 

60-120 ovules. 

Fruits are large, up to 10 (7-15) cm diameter, with 10-100 flattened 

seeds, round often shining; apple-green and somewhat knobbly often 

with yellow spots when young, turning yellow or brown when mature; 

hard shelled (shell about 5 (0.8-8) mm wide). Seeds are toxic when 

eaten. Pulp has an acid-sweet taste. 

It is regarded as semi-cultivated, particularly in West Africa, and 

numerous parts of Central, Eastern and Southern Africa. 

1.3.2.1 Subspecies S. spinosa 

This subspecies has been described with fruit shell only about 4 mm 

thick and with the leaf apex acuminate when the typical forms of S. 

spinosa have leaves less pointed to rounded. It is unlikely to be a valid 

9

taxon and using the name cocculoides is confusing. The species S. 

spinosa shows a great deal of phenotypic variation especially in hairiness 

of various plant parts, leaf shape and size, re-growth after cutting or fire 

and in flowering patterns. Infraspecific taxa based on morphology are 

not helpful: patterns of genetic variation across distribution and 

ecologies need to be assessed. 

1.3.3 Other edible fruited species 

There are not many other large-fruited Strychnos species prized for their 

fruits in Africa. They include S. pungens, S. innocua, S. lucens and S. 

madagascariensis. There are even fewer small-fruited species widely 

prized but two important ones are S. caespitosa Good (now regarded as 

S. gossweileri) and S. stuhlmanii Gilg (now regarded as S. potatorum), 

the latter being prized for cooking but not eaten fresh. 

An additional species, S. dysophylla Benth., thought to be related to S. 

innocua, is now regarded as part of S. madagascariensis. 

These taxa are described below. 

1.3.3.1 S. gossweileri Exell 

Synonym: S. caespitosa Good 

A small rhizomatous shrub species with a woody base producing a 

climbing shrub or liana which can reach 1-3 m in height or straggle up to 

20 m. Leaves petiolate (2-5 mm) a little over 2-5 x as long as wide, 

usually about 7 (2-10) cm long and 2.5 (1-5) cm wide, glabrous; very 

variable in shape, oblong-elliptic to narrowly elliptic with apex rounded 

apiculate or sub-acuminate. 

Flowers are 4(-5)-merous, corolla white, lobes oblong. 

Fruits are ellipsoidal, small, 1.5 x 1.0 (2-1.5) cm, orange-yellow in 

colour, soft shelled with a smooth skin, one seeded, pulp is eaten fresh. 

Distributed in woodlands and gallery forests in tropical Africa, 

particularly the northern part of Southern Africa at altitudes up to 700 m. 

10

1.3.3.2 S. innocua Del. 

Synonym: S. alnifolia Bak., S. triclisioides Bak., S. unguacha A. Rich. 

A shrub or much branched tree up to 13 (2-16) m tall. The trunk 

branches from low down. The bark is cream to orange-green and 

powdery and flakes near the base of the trunk. Branches powdery or not, 

branchlets glabrous but hairy in var. pubescens. 

Leaves with short petiole (2-7 mm) elliptic to obovate, 8 (4-16) x 5 (2-9) 

cm rounded (but acute on sucker shoots) at the apex, glabrous or 

pubescent on both surfaces. Venation includes 2 pairs of secondary veins 

from the base curved along the margins and prominent tertiary reticulate 

venation. 

Flowers 4-merous, corolla cream green with a ring of hairs in the throat, 

lobes triangular not thick. Fruits are round 5-7 cm diameter, yellow or 

orange in colour, thick walled, pulp eaten fresh. Seeds toxic (Irvine, 

1961). 

Widespread in savannah from Guinea to Ethiopia and Sudan, through 

Eastern Africa, Congo and northern Southern Africa. 

A variety pubescens Solered. appears to equate to S. unguacha A. Rich. 

recorded as distributed in West Africa and S. triclisioides Bak. recorded 

as distributed in East Africa to Angola and Zimbabwe. This variety has 

pubescent branchlets. 

S. dysophylla Benth., was later considered as a subspecies of S. innocua 

(subsp. dysophylla (Benth.) I.Verd.). The fruit is black, sweet and well 

tasting. This taxon is synonymous with S. randiaeformis Baill. See S. 

madagascariensis below. 

1.3.3.3 S. lucens Bak. 

A woody evergreen climber with bifurcate tendrils. Branches are closely 

lenticellate. 

Leaves are 3.5 x 13.0 cm long and 1.8-5.0 cm wide and glaucous. 

Fruits are round, about 4.0 cm diameter, orange in colour, 7-seeded and 

eaten fresh. 

11

This is a species of northern Southern Africa especially Zambia and 

Zimbabwe. 

1.3.3.4 S. madagascariensis Poir. 

Synonym: S. innocua subsp. Burtoni (Bak.) Bruce and Lewis, S. burtoni 

Bak., S. dysophylla Benth. 

A multi-stemmed and branched deciduous shrub or tree 2-10 (-20) m tall. 

The trunks branch from low down. The bark is pale grey to grey-white 

and smooth. The branches are powdery or not and hairy or not. 

Leaves with short petiole, 1-5 mm, lamina elliptic to obovate 2-10 x 1-4 

(-5) cm rounded, shiny and dark green above distinctly paler beneath, 

glabrous or pubescent on both surfaces. Venation includes 2 pairs of 

secondary veins from the base curved along the margins and not very 

prominent tertiary reticulate venation especially above it. 

Flowers are 4-merous, corolla white or green-yellow with a ring of hairs 

in the throat, lobes triangular and thick, spreading. 

Fruits are round, 2-8 cm diameter blue green turning yellow or orange 

when mature, thick walled; pulp eaten fresh orange and slimy containing 

2-50 seeds. 

Distributed in Eastern (northwards to Malawi) and Southern Africa and 

Malagasy. 

1.3.3.5 S. pungens Solored. 

Synonym: S. occidentalis Solored. 

A deciduous shrub or tree 6-8 (2-16) m tall. The trunk’s bark is greybrown 

and granular, not corky and becomes smooth on the branches. 

Leaves with short petiole, 1-4 mm, lamina 8.0 x 3.5 cm, usually but not 

always glabrous beneath, shining dark green above, slightly paler 

beneath, elliptic to obovate, apex ending in a sharp spine. Venation 

includes one pair of secondary veins from the base curved along the 

margin. 

Inflorescence axillary usually congested. Flowers 5-merous, corolla 

cream-green or yellow with a ring of hairs in the throat, with thick 

triangular spreading lobes. 

12

Fruits are round, large 5-10 (-15) cm diameter, blue black turning orange 

or yellow, calyx lobes notably accrescent. Fruit thick walled; pulp eaten 

fresh, sweet and fragrant containing 20-100 seeds. 

Distributed in Central, northern Southern Africa and Eastern Africa in 

Brachystegia woodland up to 2000 m above sea level. 

1.3.3.6 S. stuhlmannii Gilg 

A deciduous much branched shrub or tree 4-18 m tall. The bark is greybrown, 

very thin and smooth, lenticellate. Dead bark separates in small 

circular scales. Branches many and branchlet spines 1-3mm long. 

Leaves with a petiole 1-7 mm, lamina dark-green above, paler densely 

velvety beneath, elliptic or ovate 12 (6-15) x 6 (3-9) cm. Venation 

includes 2 pairs of distinct secondary veins. 

Flowers 5-merous (occasionally 4), corolla white or yellow, pilose inside 

the tube, lobes oblong spreading. Inflorescences in axis of scales at bases 

of branches. 

Fruits round, small up to 1-2 cm in diameter, blue-black, thin walled; 

purple pulp (only eaten cooked) containing 1 seed. 

Distributed in Eastern and Southern Africa from Malawi to South Africa, 

widespread in miombo woodlands, Brachystegia woodlands along rivers 

and in semi-evergreen bush up to 1600 m above sea level. 

This species has recently been treated as a synonym of S. potatorum L. 

f., which also occurs in India, Sri Lanka and Myanmar (Leeuwenberg, 

1983). 

1.4 Vernacular names 

It has proved difficult to be confident of the usage of many local names. 

Those given below in Table 1.1 appear valid. 

13

Table 1.1 Vernacular names of Strychnos species 

Species Language/ 

Area 

Name 

S.cocculoides English Corky-bark monkey orange, Monkey 

apple, Bush orange 

Afrikaans Klapper Suurklapper 

Zambia Akaminu, Akasangole, Latongo, 

Muhuluhulu, Muwi 

S. spinosa English Kaffir orange, Monkey orange 

Ghana Akankoa, Afankuru Pumpologoru, 

Kampoye, Katupwaga, Pumponsia 

Uganda Lombo, Arwalarwala-Lyech, Shiunwa 

Zimbabwe Mutamba, Muzhumi, Umkomatsane 

Zulu Umhlala 

Zambia and Kampobera, Muhuluhulu, Muwi, 

Malawi Muyimblii, Sansa, Umasaye 

Other West Kokhyo (Hausa), Katenpuanga, 

African Katerpwinga (More), Datokulewi, 

Marbatahi, Norbotahi, Uormatabe, 

Noyabata (Peulh), Ngoba (Serer), 

Ramboet, Tobé (Wolof) 

S. innocua Zambia Kutane, Mongolo, Umulungi 

Uganda Langoro, Unde, Akwalakwala, 

Erwalakawala, Eturukutsuti, Mkukulu 

S. pungens English Kaffir orange 

Zimbabwe Matamba, Umhlale 

Zambia and 

Malawi 

Ifufuma, Muwawa, Umukome 

S. stuhlmanni Zambia Mulombelcombe, Musisi, 

Umubangachulu 

S. potatorum India Nirmali (Hindi), Uriya (Kotako), Tel 

(Chilla), Tam (Tetay Kottai) 

(data from various floras and von Maydell, 1986) 

14

1.5 Key to the species 

1a. Fruit large, 4-12 cm diameter, containing 5 to many seeds 

2a. Small trees without tendrils, with paired spines Inflorescences 

terminal. Calyx lobes narrowly deltoid to linear Corolla campanulate 

3a. Branches and branchlets with corky ridges. Unripe fruit blue 

green……………………………………………………….S. cocculoides 

3b. Branches and branchlets without corky ridges. Unripe fruit apple 

green……………………..…………………………………....S. spinosa 

2b. Small trees without tendrils or spines or evergreen climbing 

shrubs with tendrils. Inflorescences axillary. Calyx lobes ovate or suborbicular. 

Corolla cylindrical 

4a. Leaves with a sharp spine at the apex..………………S. pungens 

4b. Leaves without apical spine 

5a. Scandant shrub with tendrils…………………………....S. lucens 

5b. Erect shrub or tree without tendrils..………………….S. innocua 

1b. Fruits small < 4 cm diameter containing 1 to many seeds 

6a. Fruits 1-2 seeded 

7a. Rhizomatous suffrutex habit……..………..…….. S. gossweileri 

7b. Trees …………………………………………...S. stuhlmannii 

6b. Fruits many seeded………….……………..S. madagascariensis 

15

CHAPTER 2. DISTRIBUTION 

The genus plays an important role in agricultural areas subjected to 

periodic drought (Taylor, 1986). Species are adapted to harsh 

environments such as soils with very poor fertility and dry climates, and 

may even produce fruit during years when traditional crops fail. 

Strychnos cocculoides is ideally suited to the use made of it by rural 

communities. It is widely distributed as a scattered tree in the Miombo 

and Savannah woodlands of tropical Southern Africa and Malagasy, 

occurring at a range of elevations, mainly on sandy areas (Pardy, 1953). 

S. cocculoides is found in warm to hot tropical savannah and other 

woodland regions with 600-1500 mm rainfall, but with a prolonged dry 

season. It occurs between 400 and 2000 m above sea level. Fire is an 

important part of the environment. Since both trees and grasses must be 

resistant to drought and fire, the number of species in the vegetation may 

not be large in sharp contrast to adjacent tropical forests. S. cocculoides 

is adapted to the drier tropical regions by growing only when there is 

adequate moisture in the soil. The species is adapted to drought by its 

ability to become dormant when water is not available, rather than 

wilting and dying as would be the case with drought-sensitive plants. 

The plants lose their leaves and show only green buds and stems thus 

showing high transpiration efficiency (Lange et al., 1969). 

S. spinosa also occurs in savannah forest woodlands and sometimes 

gallery forests all over tropical Africa and Southern Africa, Malagasy 

and the Seychelles, from 0-2200 m above sea level, in areas with more 

than 600 mm annual rainfall. The species tends to occur on hills and 

slopes. 

2.1 Regional distribution of S. cocculoides 

More detailed information on the distribution patterns of S. cocculoides 

in Southern Africa is provided by country in Table 2.1. 

16

Table 2.1 S. cocculoides in countries of Southern Africa 

Country Latitude Longitude Habitat 

Angola 12 º 0’S 18 º 0’E Miombo woodland (and other variants) and grassland savannahs, with 

patches of lowland rainforest in the north. Intermediate elevation forest on 

the western escarpment, Montane forests in the highlands, and desert and 

sub-desert formations in the southwest. 

Botswana 22 º 0’S 24 º 0’E Open wooded grassland and deciduous bush land in the southwest on 

Kalahari sands, Zambezian woodland in the north and east, with extensive 

wetlands in the Okavango Delta and halophytic flora in the Magadigadi 

pan. 

Lesotho 29 º 40’S 28 º 0’E Predominantly Montane grassland with occasional patches of woodland in 

ravines and river valleys. 

Malawi 11 º 55’S 34 º 0’E Predominantly Miombo woodland, with drier Zambezian woodland in the 

south, Montane forest and grassland at higher elevations, and patches of 

lowland forest on the shores of the Northern part of Lake Malawi, Nyika 

Plateau, and the lower slopes of Mount Mulanje. 

Mozambique 19 º 0’S 35 º 0’E Miombo woodland, with Mopane woodland in the Zambezi and Limpopo 

valleys. Montane forests and grasslands found at higher elevations. Mosaic 

of coastal woodlands, as well as forest/mangrove patches. 

17

18 

Country Latitude Longitude Habitat 

Namibia 22 º 0’S 18 º 9’E Dry woodland in the Northeast, becoming drier towards the South and the 

coast, through bushland and wooded grassland to the desert. The 

escarpment transition between the coastal desert and the savannahs of the 

interior. 

South Africa 32 º 0’S 23o 0’E Fynbos and its variants in the southwest, arid (succulent Karoo) and semiarid 

Karoo shrub land and grassy shrub land in northern and central Cape, 

highveld grassland over much of the central plateau, open savannah 

woodland on the eastern plateau, Montane forest and grasslands in enclaves, 

savannah and low-lying forest on the east coast. 

Swaziland 26 º 30’S 31º 30’E North-eastern mountain grassland to the west of the country with pockets of 

Afromontane forest merging eastwards into savannah shrub woodlands 

(mainly sour lowveld bushveld, sweet lowveld bushveld and Lebombo arid 

mountain bushveld). 

Zambia 15 º 0’S 28 º 0’E Miombo woodland, with drier Mopane woodland in the Luangwa and 

Zambezi Valleys and parts of the West on Kalahari sands. Patches of 

lowland forest in the northwest, and Montane forest and grassland in the 

northeast. 

Zimbabwe 19º 0’S 30 º 0’E Dry Miombo woodland, with Mopane woodland and other woodland types 

dominating. Serpentine grasslands in the Great Dyke. Montane forest 

interspersed among high altitude grasslands and heath in the Eastern 

Highlands. 

Sources: Golding (2002) and Hilton Taylor (1996)

2.2 Sampling the distribution patterns 

As expected with such a wide distribution, S. cocculoides exhibits a great 

deal of phenotypic variation. Like most trees the species is expected to 

vary broadly between and within provenances and sites. A limited 

amount of experimentation has been carried out in this area, noteworthy 

being work by Veld Products Research and Development of Botswana to 

identify superior phenotypes (Mateke, 1998); and research on the 

differences in fruit characters of provenances from Zambia, Zimbabwe 

and Tanzania (Mwamba, 1983). Both projects identified patterns of 

genetic variations which can be used in improvement. Recently, 

international provenance trials have been established in Malawi, 

Tanzania, Zambia, and Zimbabwe under the auspices of ICRAF and the 

Danish/FAO Tree Seed Centre. 

2.3 Distribution in relation to habitation 

S. cocculoides is a natural pioneer on abandoned cultivated sites (Plate 

10). It is also a tree typically left growing near farms on cultivated areas 

and in communal woodlands. This is because it provides a sustainable 

source of fruits. Trees of Strychnos are a common feature along the main 

roadsides. Almost certainly the major stimulus for their growth in such 

conditions comes from women, who collect wild fruits with children 

from locally preserved trees and also dominate the selling in markets. 

2.4 Distribution outside Africa 

S. cocculoides has been introduced to South America and India and S. 

spinosa has been tested for adaptability in the USA (Florida) and Israel. 

However, neither species has been exploited in these countries. 

The odd distribution of S. potatorum L. f. in central provinces, Bihar, 

Orissa and the western peninsula of India, and in Sri Lanka and 

Myanmar, as a species now thought to be synonymous with S. 

stuhlmannii Gilg of Africa (see Chapter 1) and recorded under this name 

in the Flora Zambesiaca requires further elucidation. However, this wild 

species is not of major relevance to this monograph. 

19

CHAPTER 3. PRODUCTION 

AREAS 

Although S. cocculoides is widespread in Africa, most produce is 

gathered from natural populations, supplemented with produce from 

trees which have been locally protected near habitations and on 

abandoned cultivated land. The same is true for S. spinosa, whereas other 

species important to rural people, such as S. gossweileri, S. innocua, S. 

lucens and S. madagascariensis are virtually only gathered from natural 

populations. 

The domestication of S. cocculoides is in its early stages and stems from 

the wider recognition of the needs to ensure household food security and 

to reduce environmental degradation around villages and settlements. 

Local fruit tree species have been accorded priority for satisfying these 

needs in part. Most of the species recognised as priority for enhanced 

domestication, including S. cocculoides, are multipurpose (Kadzere et 

al., 1998; Maghembe, 1995; Maghembe et al., 1998). 

It is to be expected that within a very short time S. cocculoides will be 

integrated into agroforestry systems in at least Botswana, Malawi, 

Mozambique, Tanzania, Zambia and Zimbabwe. The reasons for this 

relate to wide recognition in the early 1990s that the Miombo woodlands 

of Southern Africa contain fruit species that are widely harvested and 

eaten. Moreover some national programmes, e.g. that of Zambia, have 

initiated research on domestication of indigenous fruit tree species and 

have accumulated a number of years of experience. 

ICRAF developed a project, specifically for the Miombo woodland 

species, and donor governments (Germany, Canada, Norway, Denmark 

and Sweden) as well as the Rockefeller Foundation helped the 

partnership with national programmes (Akinnifesi et al., 2002). Under an 

EC project, work started on production potential of S. cocculoides (and 

S. spinosa) on the drylands of Namibia and Botswana. 

Research is now active on propagation (Mateke, 1998), agronomy, 

community processing of fruit (Saka et al., 2001) and selection (see 

subsequent chapters). 

20

The identification of future production areas will depend on combining 

high fruit production genotypes with suitable adaptation of planting 

material. In view of the wide natural and semi-natural distribution of S. 

cocculoides, it is expected that suitable patterns of genetic variation will 

be readily identified. 

21

CHAPTER 4. PROPERTIES AND 

USES 

4.1 Fruits for food use 

Fruits of S. cocculoides are fleshy inside a woody shell (Plate 7). It is 

considered one of the most pleasing of the Strychnos species when eaten, 

as well as S. spinosa. The pulp is eaten when fruits are ripe and the pips 

spat out. S. cocculoides tastes sweet, S. spinosa is more acid-sweet. 

The pulp is brown and gelatinous. Normally fruits are ripe in the period 

of October to December, when they are offered for sale throughout 

Central and Southern Africa. 

4.1.1 Fruit composition 

There are differences in the amount of pulp per fruit depending on 

geographic origin. Pulp equates to 34-48% of total fruit in Zimbabwean 

samples, 47-57% in those from Zambia and 50-57% in Tanzanian 

samples (Mkonda et al., 2002). The rest of the fruit is made up of shell 

and seeds. Mean average in the northern parts of Southern Africa is 44% 

pulp, 41% pericarp (shell) and 15% seeds. 

Unripe fruits are considered somewhat toxic because of their saponin 

content and presence of an alkaloid, the latter apparently being converted 

to non-toxic substances as fruits ripen (Williamson, 1995). There are 

conflicting reports about the presence or absence of alkaloids in the fruit 

pulp of a range of species. This needs to be investigated. 

4.1.1.1 Pulp 

The pulp contains over 30% fat, 45% crude fibre, high carbohydrate 

levels as well as saponin, citric acid and vitamin C (providing the taste). 

The physiochemical composition of pulp of S. cocculoides and two other 

species of Strychnos is shown in Table 4.1. Their mineral content is 

shown in Table 4.2. In both cases, figures are only indicative and more 

systematic analyses are needed. Detailed analyses of fruits have not been 

performed, although they provide good sources of Vitamin C, citric acid, 

zinc and copper. Pulp of S. cocculoides also contains about 0.85% of 

22

fixed reddish oil, with higher content (3.8%) in S. innocua (Laidlaw 

1951). 

23

24 

Species P Ca Mg Fe K 

S. cocculoides 20.2 9.41 26.9 0.18 188 

S. spinosa 22.6 45.8 43.6 0.75 328 

S. pungens 27.1 29.3 38.1 0.62 478 

Source Arnold et al. (1985) 

Table 4.2 Mineral composition of fruit pulp (mg/g) 

(mg/100g) 

DM Ash Crude Fat Fibre Total 

Protein 

Carbohydrates 

S. cocculoides 19.6 0.5 1.3 0.1 0.9 16.8 6.7 308 

S. spinosa 22.2 1.8 2.7 0.1 1.4 15.2 10.6 305 

S. pungens 27.9 1.0 1.1 0.8 6.2 18.9 10.7 367 

Source Arnold et al. (1985) 

Species % Component Ascorbic 

acid 

Energy 

Value 

(kJ/100g) 

Table 4.1 Physiochemical composition of fruit pulp

4.1.1.2 Dried fruits 

There is some evidence that fruits of S. cocculoides can be dried for later 

use (PhytoTrade, 2003). This is also recorded for other species including 

S. pungens and S. innocua (Codd, 1951; Greshoff, 1900). Some wild 

species such as S. gerrardii N.E. Br. and S. shumanniana Gilg - two wild 

species of South Africa - are gathered and buried in sand to ripen, if 

found unripe (Galpin, 1925). 

There appears to be a wide range of local practices which have not been 

thoroughly documented. For instance, in Botswana, dried Strychnos 

fruits are usually used for medicinal rather than food purposes. In 

Swaziland, the Swati people store dried fruits as an emergency food for 

use in times of famine. The Shangana people of Tanzania and the Swati 

of Swaziland only eat fruits of S. spinosa when fresh and not when dried 

(Codd, 1951). In Mozambique, dried ripe pulp of fruits of S. gerrardii is 

valued for making porridge (Almeida, 1930). 

4.1.2 Toxicity of Fruits 

Pulp of ripe fruits appears non toxic but a number of wild species may be 

toxic even when ripe. S. stuhlmannii is an example and local people cook 

the pulp before eating. The cooking must denature any toxins (Miller, 

1948). The fruit of this species when crushed is used as a fish poison in 

the Kruger National Park of South Africa, the poison thought to be due 

to saponin (Codd, 1951). 

4.1.3 Processing 

Fruits of S. cocculoides are processed into juice, jam and fritters of 

mixed fruits (FAO, 1996; Swai, 2001; Saka et al., 2002; Sufi and 

Kaputo, 1977; Mbiyangandu, 1985) especially in Malawi, Zambia and 

Zaire. Preservation of processed juice is achieved through the use of 

benzoic acid and sodium sulphate additives. 

Fruits of S. innocua are also processed into jam. Species which are more 

acid are often processed into marmalade. 

Urgent work is needed on assessing the organoleptic properties, such as 

flavour, texture, consistency and palatability of Strychnos products 

(Watts et al., 1989). 

25

Locally, fruits are used to produce alcoholic beverages, notably pulp of 

S. spinosa in Malagasy (Wildemann, 1946). In Malawi and Zambia 

government research has looked at the potential of S. cocculoides for 

wine production but has emphasised other tree fruits such as Uapaca 

kirkiana and Zizyphus mauritiana. In Namibia a traditional strong 

alcoholic drink is distilled from Strychnos fruits called kashipembe. A 

small enterprise was set up in 1988 in the Kavango region to develop this 

and use juice from Strychnos fruit, which are abundant in the region, to 

flavour a cane and wine-based liqueur. (Schreckenberg 2003). 

4.2 Seeds 

Although seeds are not used as food, when large numbers are present, 

some may get consumed when fruits are eaten fresh. Hence it is 

important to know something about their safety. 

Characteristically, the seeds contain alkaloids, saponins and various 

acids and polysaccharides. Each seed of S. cocculoides weighs ca. 0.6g 

fresh weight (Uronu and Msangu, 2003) but usually less; the weight of 

1000 fresh seeds varies from 382-600 g (Fletcher and Pritchard, 2000). 

Seeds of S. cocculoides contain small quantities of strychnine 

(C21H22N2O2), making the seeds slightly toxic. There is no brucine (an 

alkaloid resembling strychnine) present. 

In contrast the alkaloid is apparently not present in seeds of S. spinosa 

nor in S. madagascariensis. Seeds of S. pungens and S. innocua, though 

bitter tasting, apparently have no alkaloids. 

As stated earlier, the data are not comprehensive enough to gain a clear 

picture of seed composition. Much of the pharmacological interest has 

resulted from the use and cultivation of S. nux-vomica L. from India and 

Sri Lanka as a source of strychnine and brucine. The original species in 

which strychnine was identified was S. ignatii Berg. from the 

Philippines. 

Seeds in some cases contain reddish oil which might be of value 

commercially (Gunstone et al., 1972) and is currently being considered 

for the cosmetic industry by PhytoTrade Africa. Additionally 

galactomannan, a polysaccharide with industrial applications is known in 

seeds of S. potatorum and S. innocua (Corsaro et al., 1995). 

26

4.3 Leaves 

Leaves are not normally eaten, except for cases of S. spinosa leaves 

being used in couscous in the Sahel. Leaves of S. innocua are used as 

stockfeed. 

Interestingly, in Tanzania, the leaves of S. spinosa are reported to be 

toxic and in Mauritius they are said to produce narcotic effects. The 

presence of strychnine is reported by Githens (1949) but no alkaloids 

have been isolated from plants grown in Florida, USA. 

4.4 Medicinal uses of Strychnos cocculoides 

4.4.1 Fruits 

Green, unripe fruits of S. cocculoides are used to induce vomiting. The 

whole fruit is mashed in a mortar, steeped in water and then drunk 

(Leger, 2003; Palgrave, 1992). The emetic effect is probably due to 

toxins in unripe pulp and in the seeds. In Zambia, powder from unripe 

fruits is added to milk and drunk as a purgative. Pulp of ripe fruits, 

mixed with honey or sugar, is used to treat coughing. 

4.4.2 Leaves 

Fresh leaves are pounded and the mash mixed with water, heated and 

simmered to make a leaf porridge. This is applied to wounds to prevent 

infection and promote healing (Leger, 2003). 

4.4.3 Bark 

The middle part of the bark is cooked in water and the decoction is drunk 

as a cure for stomach pains (Fanshawe and Hough, 1960). 

4.4.4 Roots 

Roots are chewed as an alleged cure for gonorrhoea and also to alleviate 

eczema (FAO, 1983). In Zambia, local healers called Nganga use roots 

widely to treat sexually transmitted infections and often combine this 

treatment with regular purging using immature fruits. 

27

4.5 Medicinal uses of other Strychnos species 

In terms of medicinal potential, African Strychnos species have been 

documented as folk medicinal data with no commercial value to date. 

Emphasis on potential medicinal use has focussed on S. spinosa from the 

drier zones of Africa and S. icaja Baill., a liana species of rainforest, 

secondary forest and swamp and gallery forests with rainfall of 1350- 

2500+ mm throughout the year (FAO, 1986, 1988). 

4.5.1 S. spinosa 

The fruits and roots are widely used for gastrointestinal ailments as a 

laxative or emetic throughout Sub-Saharan Africa. Roots and leaves are 

used in treating venereal diseases (Zambia and West Africa) or as a 

febrifuge (Nigeria). A bark decoction is used for fever in the Ivory Coast 

and as ear drops. In Tanzania grated root mixed with coconut oil is 

applied locally to remove jiggers (larvae of mites of the family 

Trombiculidae) (Brennan and Greenway, 1949) and leaves are used in 

other regions for this purpose. In Zambia and neighbouring countries, 

leaf or root decoctions are used as an analgesic. 

Roots and green fruit are used as snake-bite remedies especially in 

Eastern Africa (Bally, 1937), the Sahelian zone of West Africa (von 

Maydell, 1986) and South Africa (Bryant, 1909). 

Leaves are used to treat conjunctivitis in Central Africa and Ivory Coast 

(Wildermann, 1946) and venereal diseases in Zambia. 

4.5.2 S. innocua 

Seeds are used as an emetic (Gilhens, 1949). 

4.5.3 S. madagascariensis 

In Angola, fruit pulp is used as a treatment for dysentery and in Eastern 

Africa the bark is used in obstetrics (Watt and Breyer-Brandwijk, 1962). 

4.5.4 S. pungens 

Leaves are used to make a decoction to ease coughing (Gilges, 1953). 

Unripe fruit pulp is used as an emetic and for snake bites. Roots are used 

for a variety of ailments; stomach ache, fever, inflamed eyes, bronchitis, 

and to make ear drops (Bally, 1937). 

28

4.5.5 Other wild species 

S. icaja is a species of West and Central Africa and the 

pharmacologically active alkaloids are described in Sandberg et al., 

(1969) and Lamotte et al., (1979); and saponins, iridoids and phenolics 

are described in Denoel et al., (1953). 

There are numerous references to medicinal uses of a range of other 

Strychnos species. However, in many cases, the correct or validated 

identity of the species is not confirmed. 

4.6 Other uses of S. cocculoides 

Wood is used in local construction and to make posts and tool handles. 

The wood is somewhat soft, fine pored and a whitish colour. S. spinosa 

is similarly used but is harder and yellowish. 

The fruit shell is used as a resonance board for musical instruments as 

are those of S. spinosa (von Maydell, 1986) and S. pungens. 

The fruit is used to make a dye for colouring fruit trays and containers 

are made from the shells. Fruit pulp can be used as a soap for washing 

clothes due to the saponin content. 

4.6.1 Other uses of other Strychnos fruits 

Pulp of S. potatorum is also used as a soap. 

29

Plate 1. Fruit and spines of monkey orange 

Plate 2. Fruit and rounded leaves of monkey orange 

30

Plate 3. Mottled unripe fruit of monkey orange 

Plate 4. Fissured bark 

31

Plate 5. Variation in fruit of Strychnos species 

Plate 6. Variation in leaves of Strychnos species 

32

Plate 7. Brown jelly-like pulp surrounding the seeds which can be 

processed 

Plate 8. Pale seeds of monkey orange 

33

Plate 9. Monkey orange height and form at maturity 

Plate 10. Monkey orange: A pioneer on abandoned cultivation sites 

34

Plate 11. Deciduousness in monkey orange 

Plate 12. Monkey orange growing and fruiting on limestone parent 

material 

35

Plate 13. Grass providing partial shade to young monkey orange 

trees 

Plate 14. Sapling on abandoned cultivation site 

36

Plate 15. Effect of fire on saplings 

Plate 16. Coppice sprouts in monkey orange after fire 

37

Plate 18. Coppice shoots 

Plate 17. Grafted plant flowering after one year 

38

Plate 19. Coarse and fine roots of mycorrhizal root system 

Plate 20. Termite damage of a seedling 

39

Plate 21. Powdery mildew on bottom fruits while in storage 

Plate 22. Monkey orange chopped for firewood and fruits scattered 

on the ground. 

40

Plate 23. Extraction of roots for medicinal uses. 

Plate 24. Monkey orange and Masuku (Uapaca kirkiana) wine 

Plates courtesy of C. Mwamba 

41

CHAPTER 5. ECOLOGY 

5.1 Introduction 

Strychnos cocculoides typically occurs in areas that are characterised by 

having two main seasons, a wet one and a dry one, in tropical Central 

Africa. The wet season lasts from November to April. During this 

period, rainfall occurs for an average period of 114 days and is modified 

by elevation. In January, the rainfall is around 250-500 mm and the mean 

monthly maximum and minimum temperatures are 26°C and 16°C 

respectively. The dry season occurs from May to October. In the first 

cool part of the dry period in May-September, there is no rainfall and the 

mean monthly extreme temperatures are 26°C and 6°C respectively. 

During the second part of the dry season (September to October), warm 

conditions prevail with extreme temperatures of 32°C and 14°C during 

October. 

In other regions, such as in West Africa, the genus Strychnos occurs in 

climatic conditions which do not vary in extremes, but there are three 

geographical zones: a dry desert zone prevalent in the north, a middle 

belt savannah region and a rain forest region of the south. 

Strychnos cocculoides is a remarkably unexacting species. It tolerates 

both the hot and dry summers of the tropics by being deciduous (Plate 

11). It sheds its leaves during the dry season and, unlike many other wild 

fruit trees, it does not produce new leaves until the rains. It occurs from 

high rainfall tropical forests to more desert climates of Southern Africa, 

at a range of altitudes, and can colonise dry sandy soils including 

calcareous ones. It is, therefore, most frequent where more demanding 

trees fail to grow. 

The estimated areas of the natural tropical forests in which the genus 

Strychnos is endemic are presented in Table 5.1. 

42

Table 5.1 Principal biomes in which the genus Strychnos is endemic, prevailing climate and dominant growth 

forms 

(adapted from Burley and Styles, 1976; Jensen and Salisbury, 1972). 

Climate Dominant Growth Form 

Biome Area 

(million ha) 

Temp. (ºC) Rainfall (mm) 

Trees, both evergreen and deciduous 

Trees, broad leaved deciduous 

Trees, grass and shrubs 

560 

308 

588 

Tropical Zone 

Wet evergreen 

Wet evergreen 

Dry deciduous 

Trees, evergreen 

Trees, broadleaved deciduous 

Trees, grass and shrubs 

Shrubs, succulents 

Shrubs, succulents 

Total 1456 18-35 1000-12500 

Subtropical Zone 

Wet evergreen 8 

Subtropical moist 20 

Subtropical dry 196 

Total 224 13-40 250-900 

Desert 

Tropical warm * 

Temperate warm * 

Total 448 2-57 0-250 

* data unavailable 

43

5.2 Rainfall 

Under arid conditions, Strychnos cocculoides and S. spinosa tend to 

develop deep root systems. This confers an ecophysiological advantage 

in accessing the available water and they are able to use periodic rainfall 

quickly and effectively, especially small quantities of water before it 

evaporates. S. cocculoides, S. spinosa and S. pungens, were tested for 

growth under different desert ecozones in the Negev desert, Israel 

(Mizrahi et al., 2002). S. pungens did not survive the arid conditions. S. 

cocculoides, which is considered the best of the three in terms of eating 

quality, survived only in the Besor region because of good quality water 

(not saline) and moderate temperatures. The best of the three species to 

grow under desert conditions was S. spinosa. Mizrahi and his co-workers 

successfully established about 15 fruiting trees, but with a high 

variability in growth, yields, fruit size, ripening season and taste. Some 

of the trees bore astringent, bitter fruits, while others bore very sour 

fruits and only two trees bore fruits which were of excellent taste. 

S. cocculoides occurs in areas where rainfall is restricted for part of the 

year, and the plant survives long periods of drought. Minimum annual 

rainfall per year is 600 mm and maximum annual rainfall per year is 

1200 mm. The seed is desiccation tolerant (Pritchard et al. 2004). 

5.3 Light 

S. cocculoides is a light- demanding species. The effect of light intensity 

on growth and root/shoot ratio in the species were evaluated by Mateke 

(1998) in Botswana, who grew seedlings under 50% and 25% shade and 

full direct sunlight for six consecutive summer months. Seedlings grown 

under 50% shade netting grew fastest, followed by seedlings grown 

under 25% shade netting. Table 5.2 shows the performance of S. 

cocculoides seedlings compared to those of Sclerocarya birrea under 

different light regimes. For S. cocculoides: 

(i) Height increased with increased level of shading; 

(ii) There was a weak negative linear response in stem diameter, 

root and shoot weight with the level of shading; 

(iii) Root/shoot ratio decreased with increased level of shading. 

44

Table 5.2 The performance of S. cocculoides and Sclerocarya birrea seedlings under different shading regimes 

Source: Mateke, 1998 

Parameter Strychnos cocculoides Sclerocarya birrea 

Shade (%) Shade (%) 

0 25 50 0 25 50 LSD 

Survival (%) 100 100 100 100 100 100 NS 

Height (cm) 11.2 12.9 14.9 20.6 29.3 37.8 2.1 

Stem (mm) 3.3 3.7 3.1 9.6 8.3 7.5 1.0 

Shoot weight (g) 1.4 1.0 1.4 11.6 11.4 11.0 1.6 

Root weight (g) 5.5 3.1 2.5 89.9 72.0 67.0 4.3 

Root/shoot ratio 3.9 3.1 1.8 7.8 6.3 6.1 2.7 

45

Shade therefore has a negative effect on the growth of seedlings of S. 

cocculoides. Grasses normally provide limited shade in natural 

communities (See Plate 13). 

The effects of light intensity on height, dry weight and root/shoot ratios 

given in Table 5.2 are typical (Kramer and Kozlowski, 1960). They 

suggest that S. cocculoides has a low capacity to endure shade, in part 

explaining its occurrence as a pioneer tree in old cultivated fields and 

open woodlands. Seedlings appear to develop root biomass before 

producing a lead stem. 

5.4 Wind 

The effect of wind on S. cocculoides is twofold: 

(i) It tends to decrease leaf temperature, which reduces 

transpiration, but also removes water vapour from the vicinity of leaves. 

This helps to maintain a steep vapour-pressure gradient from the leaf to 

air and therefore tends to increase the rate of transpiration. 

(ii) Winds of high velocity sometimes cause reduction in 

transpiration, apparently because they cause closure of stomata. 

5.5 Soil requirements 

S. cocculoides grows well on sites with deep sandy soils and on welldrained 

slopes. It is also found on black to dark grey clays and yellowred 

loamy sands derived from limestone parent material (Plate 12) 

The general soil physical characteristics are summarised in Table 5.3. 

46

Table 5.3: Soil physical requirements of S. cocculoides 

Characteristic Requirements 

Soil type (texture) • Deep sandy soils 

• Black to dark-grey clays 

• Yellow-red loamy sands 

• Red yellow-red loams 

Topography • 0-13% slope 

Rooting depth • Restricted by aeration 

• Restricted by rock outcrops 

Drainage • Well drained 

Terrain • Flat woodlands and savannahs 

• Hilly woodland slopes 

• Rocky slopes 

Soil pH • Acidic, pH 4-6 

In an effort to recognise the potential management advantages and any 

dangers of extending the genus Strychnos beyond its natural distribution 

limits, Mwamba (1983) related the stocking density (or number of trees) 

per hectare with soil properties in areas of natural occurrence of S. 

cocculoides and S. pungens in Zambia. The results are summarised in 

Table 5.4 and reveal that maximum stocking density for Strychnos 

cocculoides is obtained on loamy soil with cation exchange capacity 

(CEC) of 5-10 meq/100 g clay, base saturation of 60-80 % and soil pH 

5.0-6.0. 

Table 5.4: Stocking densities of S. cocculoides and S. pungens on soil 

with varying properties (Source: Mwamba, 1983) 

Soil Property Stocking Density/Hectare 

S. cocculoides S. pungens 

Texture 

Sandy 0 3 

Loamy 3 35 

Clay 0 17 

CEC (meq/100 g clay) 

5-10 4 10 

10-15 0 2 

>15 0 0 

Base Saturation (%) 

80 0 0 

Soil pH 

6.0 0 0 

S. pungens also shows maximum stocking density on loamy soils, but 

with CEC of less than 5 meq/100 g clay, base saturation of 20-60% and 

pH of 5.0-6.0. The high base saturation and CEC requirements of S. 

cocculoides suggest that this species requires richer soils than S. 

pungens, and this may explain why S. pungens is more widely 

distributed on poor sandy soils. However, both species are characteristic 

of nutrient poor soils derived from granites, coarse schists, sandstones, 

quartzites, limestones, Pleistocene sands and gravel parent materials. 

Bonifacio et al., (2000) reported S. cocculoides and S. spinosa on soil 

types as shown in Table 5.5. The chemical characteristics of these soils 

are summarised in Table 5.6. Although optimal conditions for the growth 

of these trees are practically unknown, the species is among the 

dominant woody plants of the vegetation communities in various tropical 

areas (Malaisse et al., 1999). 

48

Table 5.5: Soil types that support S. cocculoides and S. spinosa in Botswana (adapted from Bonifacio et 

al., 2000) 

Soil Parent Material 

Sandstone 

Sands 

Basalts 

Kalahari Sands 

Red Sandstones 

Kalahari Sands 

Sandstone 

Sandstones and conglomerates 

Species Soil Classification Land System 

Division 

S. cocculoides Typic Torripsamments 

Sandveld 

Lithic or Typic Torripsamments Sandveld 

Lithic or Typic Torripsamments Hardveld 

Typic Quartzipsamments 

Sandveld 

Typic Torripsamments 

Hardveld 

Typic Quartzipsamments 

Sandveld 

S. spinosa Typic Torripsamments 

Sandveld 

Typic Torripsamments 

Sandveld 

49

50 

Mean 

Standard deviation 

S. spinosa Serewe 1 

Pilikwe 1 

Mean 

Standard deviation 

* ND: Not determined 

5.8 

±0.1 

4.0 

±1.5 

0.40 

±0.07 

0.45 

±0.07 

10.6 

±12.4 

3.0 

±1.1 

5.9 

5.7 

3.0 

5.1 

0.35 

0.45 

0.4 

0.5 

0.5 

5.8 

±9.1 

1.8 

19.4 

2.3 

3.8 

5.7 

5.7 

±0.8 

7.4 

4.4 

±2.0 

0.21 

0.42 

±0.28 

±0.26 

0.2 

0.42 

1.0 

5.2 

±8.3 

S. cocculoides Serowe 1 

Serowe 2 

Serowe 3 

Shakawe 

Manyana 

Hamoye 

5.9 

5.2 

7.2 

5.4 

4.7 

3.0 

4.0 

4.1 

1.7 

5.9 

0.35 

0.27 

0.92 

0.19 

0.55 

0.4 

0.3 

0.9 

0.2 

0.5 

1.8 

1.4 

22.0 

ND* 

3.7 

2.3 

1.8 

22.0 

0.8 

3.3 

Species Site pH Clay (%) Organic C 

(%) 

N 

(g/kg) 

P (g/kg) CEC 

(C mol./kg) 

Table 5.6: Chemical soil properties that support S. cocculoides and S. spinosa in Botswana (adapted from 

Bonifacio et al., 2000)

5.5.1 Details of soil types 

Strychnos cocculoides trees occur on a variety of soil-forming rocks 

which may be grouped into four major ecological divisions on the basis 

of their contribution to soil fertility (Wilde, 1958) as below: 

5.5.1.1 Siliceous Rock Parent Material 

It occurs on siliceous parent materials, including sandstones, siliceous 

shales, conglomerates and quartzites. These rocks generally produce 

soils which are poor in nutrients and of low exchange capacity. Such 

soils are suitable for less nutrient-exacting tree species. A locally high 

base saturation as a result of the presence of lime in the profile and/or 

higher clay content may considerably raise the productive capacity of 

these soils. 

5.5.1.2 Orthoclase – Feldspathic Rock (Acid Igneous Rock) 

Parent Material 

Soil parent materials, granite, syenite, granitic porphyry, orthoclase 

felsites and gneisses weather into sandy loams or loam soils believed to 

be well supplied with potassium, but low in calcium and magnesium. 

Where rainfall is not intensive, sufficient quantities of phosphates may 

be expected. These soils are generally well adapted to all forest trees, 

except for a few lime-demanding species. 

5.5.1.3 Ferromagnesian Rock Parent Material 

These parent rocks gabbro, diorite, diabase, basalt, andesite and schists 

enriched in augite, amphibole or olivine, upon sufficient weathering, 

produce fine-textured soils with an abundant supply of calcium, 

magnesium, phosphorus and other essential nutrient elements. 

5.5.1.4 Calcareous Rock Parent Material 

These parent rocks include limestone, dolomitic limestone, chalk and 

calcareous shales. These rocks have given rise to soils of greatly variable 

productive potential, determined by climatic conditions, degree of 

weathering and clay content. The deficiency of potassium and 

phosphorous, the high content of carbonates, and the alkalinity are 

among the adverse factors that affect tree production (Wilde, 1958). 

51

5.6 Ecotypic differentiations 

It is thought that there are distinct ecotypes adapted to different 

environments. More research is needed, however, on the patterns of 

genetic variations within and between populations of S. cocculoides and 

any correlations with geographic, climatic or ecological parameters. 

5.7 Mycorrhizae 

The fine root systems of Strychnos fruit trees are reported to form 

symbiotic associations with some beneficial fungi (Munyanziza,1994; 

Mwamba, 1995). The failure to cultivate some of the wild fruit tree 

species in areas where they have never existed before is attributed to the 

lack of natural mycorrhizal symbionts (Bowen, 1980; Munyanziza, 

1994). 

Some research has been carried out in this respect on Kalahari sandy 

soils (McGonigle, 1997; Sinclair, 1998; Mateke, 1998a). Seedlings of S. 

cocculoides when artificially inoculated with isolated cultures of 

arbuscular mycorrhizal fungi (AMF) from soils taken beneath parent 

trees in their natural habitat resulted in better growth (Table 5.7). 

Mateke (1998a) concluded that artificial inoculation of the seedlings 

with the right AMF at planting stage may play a significant role in the 

promotion of initial growth. 

52

Table 5.7: Effect of mycorrhizal inoculation on initial growth of outplanted indigenous fruit tree 

seedlings (Source: Mateke, 1998) 

Species Average Plant Height (cm) at 100 Days 

Control Inoculated Increase over control (%) 

Azanza garckeana 

8.1 17.5 

116 

86 

13.8 

7.4 

Vangueria infausta 

81 

35.3 

19.5 

Sclerocarya birrea 

63 

15.0 

9.2 

Strychnos cocculoides 

53

5.8 Fire 

S. cocculoides grows in environments vulnerable to fire and is capable of 

bearing fruit afterwards. The vulnerability of a species to damage by fire 

is linked to its environment and to the rates at which flammable materials 

accumulate. 

Individual trees are readily killed by fire, but Strychnos cocculoides 

regenerates readily from shoots (see Plate 16). It can also regenerate 

from seed as the woody fruits tend to be held on the tree and survive fire. 

Detailed observations of succession after fire in woodlands containing S. 

cocculoides have not been recorded. In the drier forests, where S. 

cocculoides typically occurs, most fires have relatively little effect on the 

tree stratum; it is the herb and shrub layers and the seedlings of tree 

species that are heavily impacted by fires. 

Fire appears to be a factor essential to maintenance of the natural 

community where S. cocculoides occurs because reduced fire 

frequencies may lead to premature die-back. 

54

CHAPTER 6. AGRONOMY 

6.1 Seed propagation 

Fruit production in S. cocculoides commences between 4 and 6 years of 

age in open-grown stands, but several years later in denser re-growth. In 

general, flowering is continuous, but with light to moderate flowering in 

some years followed by heavier flowering in others. 

The fruits of S. cocculoides contain from 10 to 100 (usually 25-30) pale 

seeds (Plate 8). The average seed weight varies between 0.3-0.6 g with 

moisture content 25-30% of fresh weight. Fruits begin development in 

July-September, mature about June of the following year and are 

available thereafter and even up to December. Hence opening can be ca. 

8 months from pollination. 

6.1.1 Seed collection and handling 

When attempting to collect seed samples over a wide range, the total 

time allowed by the length of the fruiting season can be a severe 

constraint. Determination of the most appropriate sequence of collecting 

sites, difference in fruiting times and also the available means of 

transport between sites can be a complex problem. Adverse climatic 

conditions may greatly alter the time needed at a given site, by 

hampering travel, collection and seed extraction. The preparation of 

alternative plans to cover such contingencies is particularly important 

when synchronisation of the work between separate teams is needed. 

The physiologically mature fruits, light green or yellow, may have to be 

picked from the tree or they may drop on their own (at full maturity) 

after separating from the fruit stalk. Fruits are cracked open using a 

gentle tap on a stone or by hitting the fruits with a stick. The seeds which 

are embedded in the brown pulp do not adhere to the sides of the fruit 

shell and thus can be removed readily. After collection, seeds are 

thoroughly cleaned by scraping them over a wire mesh with sand. Seeds 

may be bulked for each tree or for all the trees of a population. 

55

6.1.2 Seed treatment and germination 

Normally freshly harvested seeds give up to 75-80% germination but 

occasionally this can be lower, usually because some seeds are 

immature. 

There are differences of opinion as to whether seeds should be treated 

prior to sowing. Some say no treatment is necessary, others suggest 

soaking seeds in hot water for 24-48 hours prior to sowing (Taylor and 

Kwerepe, 1995). Seeds appear to have hard coats and in nature annual 

fires are thought to soften seed coats. Treatments by soaking seeds with 

potassium nitrate or thiourea have not had beneficial effects (ICRAF, 

2000). 

Seeds are sown at a depth of 2-3 cm in pots or seedbeds (or even direct 

sowing into an orchard). Time to germination varies greatly. In 

Botswana, germination of 80% has been reported in 3 weeks when sown 

in summer propagation boxes, but seeds sown in winter took more than 9 

weeks to germinate (Taylor, 1983). Seeds stored dry at room temperature 

(23-28°C) for 8-12 months gave good germination (Mateke, 1998). 

Tests show that the seeds are desiccation tolerant and storage at 15-16°C 

after drying results in good germination after 3-6 months (Fletcher and 

Pritchard, 2000). After desiccation to 5% moisture content, germination 

falls to 59% (Uronu et al. 2005). 

Sizes are extremely variable, differing year to year from tree to tree and 

even with location in the fruit. The largest seeds are usually produced in 

the largest fruits and those near the middle of the fruit are larger than 

those at the ends. 

6.2 Vegetative propagation 

S. cocculoides coppices readily (Plate 18). Shoots arise from the stumps 

of trees that have been cut down, or killed-back during burning. It would 

seem, therefore, that propagation should be possible by taking cuttings of 

re-growth shoots. However, cuttings appear not to produce roots, even if 

kept in shade and shoots sprout (Taylor, 1983). 

Additionally there are no reports of air layering. Thus low-cost 

vegetative propagation is not possible. Nonetheless, grafting and 

budding techniques are possible (see below). 

56

6.2.1 Grafting 

Two approaches are being taken at present. In the first, Veld Products 

Research and Development of Botswana has developed techniques of 

grafting seedlings with productive parts of mature trees. This has 

resulted in improvement in the growth of the trees (Mateke, 1998). The 

wedge method is preferred. This research has been focused on 

identifying superior cultivars and development of mass propagation 

methods. 

In Zimbabwe, research has focused on collecting a wide range of 

germplasm from the Southern African region and carrying out grafting 

when evaluated ecotypes or superior genotypes are identified 

(Nyamutowa and Mushonga, 1995). 

6.3 Orchard establishment 

6.3.1 Site preparation 

S. cocculoides is a light-demanding tree. The best preparation of planting 

sites is complete clearing, ploughing and/or ridging, and elimination of 

herbaceous weeds, especially grasses for 2-3 years. 

6.3.2 Planting 

Planting is usually by hand, ideally when the soil is at or near field 

capacity. Planting holes are spaced at 5 x 5 m to provide 400 trees per 

ha. 

If direct seeding is practiced, two seeds are planted per hole and 2 

months after emergence one seedling is rogued. There are no reasons to 

support direct seeding due to the variation in germination and 

establishment rates. 

It is better practice to use nursery-raised seedlings produced from 

containers. Initially seeds are sown in trays at 2-3 cm depth, and resultant 

seedlings transplanted to tubes, sleeves or pots containing sterilized 

nursery soil. Polythene bags are cost effective. 

57

6.3.3 Seedling survival 

Addition of soil from local S. cocculoides sites will enhance the 

possibility of mycorrhizal associations (see 6.3.4 below). Seedlings need 

to be transplanted with minimal damage to the root system. 

Survival is in excess of 80% providing adequate watering is carried out, 

weeds are controlled and seedlings are protected from livestock. A great 

deal of training of smallholder farmers is necessary in this area. 

6.3.4 Mycorrhizae 

Research has shown the beneficial results of artificial inoculation at 

transplanting time with cultures of arbuscular mycorrhizal fungi from 

soils taken from beneath parent trees in their natural habitat. This work 

has been carried out with the University of Pretoria (Mateke, 1995, 

1998a). 

It appears that transplanted seedlings of S. cocculoides can pick up 

indigenous mycorrhizae at the planting site, but attention to this need at 

the nursery stage could reduce any transplanting stress. 

6.4 Orchard management 

6.4.1 Stand density 

Depending on the intensity of management, stands originally established 

at 5 x 5 m spacing may be thinned to 10 x 5 m or 10 x 10 m providing 

200 or 100 trees per ha. This would be done after the second year 

assuming the trees start bearing fruit at about 5 years (but as early as 3 

years) following transplanting. Intensity of management relates to the 

use of fertilizers (6.4.2) and pruning (6.4.3). 

6.4.2 Fertilizers 

S. cocculoides responds well to inorganic fertilizers. Seedlings in nethouse 

nurseries treated with super phosphate and nitrosol grew rapidly in 

height in 3 months compared to untreated seedlings (38-76 cm compared 

to 20 cm untreated: Taylor and Kwerepe, 1995). More detailed analysis 

of fertilizer regimes has not been reported. 

Such treated nursery-raised stock should be transplanted to the field 

holes to which a little organic manure has been added to aid the 

58

transplanting. Treatment with inorganic fertilizers at the start of the wet 

season and half way through it is recommended. 

However, it is essential that water stress is avoided and that any water 

deficit in dry summers is not exacerbated by the presence of recent 

inorganic fertilization. 

At present, less than 1% of local smallholders apply inorganic or organic 

fertilizers to farm-forestry trees (Mango and Akinnifesi, 2000). 

Additionally, severe deficiency of nitrogen or phosphorus reduces 

mycorrhizal formation and overall growth of the trees although a 

moderate deficiency can stimulate mycorrhizal formation. It is better to 

consider this at the nursery stage rather than changing orchard 

management. 

6.4.3 Pruning 

In natural stands, ‘pruning’ in S. cocculoides involves several sequential 

phases. These include weakening of branches, infection with wooddecomposing 

fungi, wood decay, severance of the branch from the tree, 

and wound healing. Natural pruning is set in motion in dense stands with 

low rates of photosynthesis in leaves of heavily shaded lateral branches. 

Since S. cocculoides is widely and variably spaced in the forests, natural 

pruning takes place within the canopy, starting with more shaded 

branches and is generally very slow. Water deficits also contribute to 

natural pruning. Death of branches is preceded by an extended period of 

retarded growth. After the branch is severed, a protection layer develops 

between the outer dead stub and the inner living part of the branch. 

Pruning in orchards should remove any dying side shoots and create a 

good crown shape permitting light entry. At present information is not 

available on the effects of decreasing the numbers of fruit spurs to 

increase size of fruits. Current efforts are under way to use genotypes for 

orchard planting that inherently produce larger and better quality fruits. 

6.4.4 Weeding and intercropping 

Weeding is crucial at the transplanting and establishment phase. In 

subsequent years attention might focus on the possibilities of 

intercropping. 

59

ICRAF has adopted a holistic approach to intercrop indigenous fruit 

trees, including S. cocculoides, with agricultural crops on farmers’ fields. 

Since 1998-99, 35,000 selected, but unimproved, seedlings have been 

planted by almost 3,000 farmers in pilot areas in southern Malawi 

(Mango and Akinnifesi, 2000). This was part of a project in which 

seedlings of different indigenous fruit trees were raised at Makoka 

Agricultural Research Station for distribution to farmers. Farmers were 

selected on the basis of their interest in growing these trees. The farmers 

were all drawn from areas where ICRAF already undertakes agroforestry 

work in Machinga and Zomba districts of the Machinga Agricultural 

Development Division and part of the Blantyre Agricultural 

Development Division, Chiradzulu district. It is expected that clear 

recommendations for intercropping packages will be forthcoming from 

this experience. 

6.4.5 Protection from pests and diseases 

There appear to be no major pests and diseases other than at seed 

germination (Taylor and Kwerepe, 1995). Termites can be a problem, 

however, for both seedlings and mature trees - usually when trees are in 

a poor physiological condition. Termites feed on dead bark hence good 

husbandry practices go a long way to eliminate such damage (Plate 20). 

The species does not appear to be commonly attacked by insect 

defoliators. Animal browsing can be a problem with young trees but is 

not serious for mature ones. 

The major fungal problems are root rot and damage from fungi during 

germination and emergence, as for any other tree or crop species. 

Fruits may be attacked by powdery mildews when in storage (Plate 21). 

6.4.6 Physical stresses 

S. cocculoides is killed by fire at the small sapling stage but mature trees 

are more resistant and tend to recover. Wind causing stem breakages can 

be a problem on shallow soils but this can be avoided by good site 

selection. 

Frost may be a limiting factor since light frosts are experienced in some 

parts of the natural distribution of the species. However, fertilized 

seedlings can withstand temperatures a few degrees below freezing. 

60

6.4.7 Growth rates 

Not enough data are available on growth rates particularly for species 

which can grow in areas with long dry periods, e.g. the Kalahari sands of 

Southern Africa (Mateke, 1998). 

However, major differences have been noted across regions and these 

appear to depend on differences in soil types and temperatures during the 

winter months. In reasonable sites trees can reach over 2 m in 4 years 

and have a crown size of 1.5 m per tree, similar to S. spinosa. 

6.5 Agroforestry 

A range of agroforestry systems can be developed using S. cocculoides 

as a component. 

In Botswana, a trial was established in a 6 year old orchard of 

Sclerocarya birrea trees which were in rows 15 m apart with trees 12 m 

apart. Interplanted in the 12 m between trees were S. cocculoides and 

Vangueria infausta within a contour bund system. Arable crops such as 

sorghum, cowpea and watermelon were incorporated. Results so far 

seem promising and this type of dryland agroforestry provides a range of 

fruits. 

6.6 Ongoing research 

Agronomy practices, plant soil relationships, mycorrhizae and other 

aspects relevant to this chapter are under active investigation through an 

EU cooperative programme with researchers in Holland, Germany, 

Israel, Botswana and Namibia. See 

http://www.divapra.unito.it/res/en0035 

61

CHAPTER 7. REPRODUCTION 

AND HARVEST 

7.1 Reproduction 

S. cocculoides bears female (pistillate) and male (staminate) flowers on 

the same plant. Much more research is needed on breeding systems of 

the African species of Strychnos: most appear to be naturally outcrossing 

through the development of mechanisms to minimise self fertilising but 

mating systems can include up to 20% self fertilisation. 

It is thought that flowers are mostly pollinated by a range of insects. 

Fruit (and seed) dispersal is by mammals. Fruits in the wild simply drop 

from the trees and are gathered, particularly by monkeys. 

7.2 Harvesting 

7.2.1 Harvesting practices 

The cycle of harvesting varies from a few weeks to longer because of 

variation in time of ripening. There are two methods of harvesting fruits. 

The first is to wait for physiological maturity and natural separation of 

the fruit from the fruit stalk when the fruit drops to the ground. The 

mature fruit may be yellow or still green at the time it drops. The second 

method is to use an implement to knock the green fruit onto the ground. 

The fruits are collected from the ground, mostly by women and children, 

since they are usually eaten raw immediately after collection. The green 

fruits may require incubation until ripe, before they are to be eaten. Both 

people and monkeys incubate the fruits under the sand. 

7.2.2 Yields 

The total weight of fruits produced per tree is influenced by the 

provenance. When grown in favourable conditions total weight of fruits 

can vary from 40-100 kg. Fruit numbers vary from 300 to 700 per tree. 

S. spinosa can produce up to 180 kg per tree and 350-850 fruits per tree. 

Comparisons between four populations in Zambia showed variation from 

62

158-296 g weight of fruit. Pulp made up 47.6 to 52.8% respectively. The 

heaviest seeds were also found in the heaviest fruits (Mkonda et al. 

2003). 

7.2.3 Post-harvest handling 

After harvest the fruits may be washed with water to remove any debris 

from the soil, and used immediately or air-dried and stored for 

processing. Air-drying fruits can prevent some surface infection by 

powdery mildew and extends shelf-life. 

Storage facilities are often lacking: the fruit industry needs to be 

transformed so that such durable fruits can be kept for longer periods. 

Currently for example in Zambia, there is a lot of wastage due to lack of 

storage technology. Hence, many fruits rot in the field or while in 

storage. At present, fruits which could be consumed throughout the year 

are actually consumed in 3-4 months before many rot and go to waste. 

The other constraints include lack of accessibility by roads and high 

transportation costs because the fruits are heavy and large trucks are 

needed. 

7.3 Processing 

In some areas processing is not recommended because fresh values are 

so high and there is a reasonable shelf-life. In Botswana, each fruit is 

worth US $ 0.45 (Taylor et al. 1996). As a result Zimbabweans drive 

truckloads across the border to sell them fresh rather than providing 

fruits for processing. 

Nonetheless, processing adds value. Immediate processing includes 

washing, shelling and pulping. Juices and jams are produced using 

heating for sterilization. Alcoholic drinks are produced in Tanzania and 

Malawi. 

Availability of skilled labour to apply appropriate technology, 

management expertise and capital for investment and marketing of 

processed products is still at a low level in Southern Africa. 

Almost certainly the major processing will relate to making juices. Other 

products, such as dry fruit rolls, have potential but have not yet been 

explored. 

63

A pilot enterprise set up in Namibia in 1988 used monkey orange fruit to 

flavour liqueur. The fruit was exported to South Africa for processing, 

which required five permits, including a collection permit from the 

Ministry of Environment and Tourism, a phytosanitary certificate and an 

import permit from the South African Ministry of Agriculture. Costs 

included transport to South Africa, covered by a Small Business Credit 

Guarantee Scheme. Promotion of the business is now needed to increase 

monthly turnover and allow development of a factory in Namibia. 

7.4 Economics 

A case study on marketing is under way in Zimbabwe which will 

generate baseline information on the incentives and constraints of the 

current marketing system. This includes assessment of the market 

structure and assessment of consumers’ willingness to pay. A parallel 

study is assessing the economics of returns on plantings, what levels of 

tree improvement are needed to induce farmers to invest and what 

competition there is with other on-farm activities. 

These studies are being carried out of at the Institute of Economics in 

Horticulture at Hannover University, Germany and greatly adds to 

technology generation in Africa (See: http://www.unihannover.de/en/internat/kooperat/int_forschproj/e_proj_wiwi04.htm) 

The majority of rural households were shown to benefit from the 

consumption and sale of indigenous fruits, collected from wild sources. 

Indigenous fruit is generally consumed as a snack, with children being 

the main consumers (See Table 7.1). 

64

Table 7.1 Percentage of households consuming indigenous fruits as a 

snack or main meal during normal, bumper and disaster harvest 

seasons for maize in two areas in Zimbabwe. 

Success 

of maize 

harvest 

Murhewa Takawirwa 

No 

consumption 

Main meal 

Snack 

No 

consumption 

65 

Main meal 

Snack 

Normal 22.6 0.5 76.9 0 0 100 

Bumper 21.7 0.5 77.8 0 0 100 

Disaster 21.7 0.5 77.8 0 34.1 65.9 

Source: Mithöfer and Waibel (2003) 

The activity of wild fruit collection was compared in this study to 

production of field crops, household gardens, livestock rearing, growing 

exotic fruit trees and indigenous fruit trees and household tasks such as 

brick-making. Collection of tree products was found to be an efficient 

activity relative to other income generating activities. Gross margins for 

the collection of indigenous fruits were lower than for livestock and crop 

production. Collection provided greater returns for labour allocation than 

from crop growing and livestock rearing, demonstrating that it is an 

efficient use of labour. Since labour is generally the limiting factor in 

rural Africa (Upton 1987), it is an important criterion in the choice of 

activities. 

Fruit sale is done by women of a household to provide cash income to 

purchase household goods. Fruit selling also bridges a gap in cash 

supply, which can be used for farming and household activities. 

Indigenous fruits are available during the dry season, when food 

availability is low, and at the beginning of the wet season when labour 

for agricultural activities reach a peak. However fruit collection can be 

combined with other activities in the field. The average farm gate price 

for Strychnos fruit was 1.5 ZWD per fruit. 37% of households in 

Murehwa District Zimbabwe receive 0-500 ZWD for fruit selling. In 

addition the wood is also valued for timber and the leaves as a source of 

nitrogen for growing vegetables. Fruit selling is related to the proximity 

of wild grown fruits to the village and to the market.

Currently collecting fruits is more profitable than planting trees. 

Technical change and reduced tree density would lead to greater 

incentive to plant indigenous fruit for its products and for biodiversity 

conservation. It was found that collection costs would need to rise or the 

performance of domesticated trees would need to improve to make 

planting a viable option. Improved performance consisted of, increased 

yield, reduced time to fruit bearing, and improved fruit quality. 

Increasing population pressure, and intensification of land use have 

caused severe deforestation of woodlands in southern Africa. 

Traditionally indigenous fruit trees were preserved when land was 

cleared for agricultural production (Clarke et al. 1996), but they are 

under increasing pressure. Farmers have rarely planted indigenous fruit 

trees and continued collecting from communal areas (Campbell 1996). 

Deforestation of natural forests increases the collection costs and reduces 

labour productivity. 

Expected returns from planting depend on the number and value of 

products that could be obtained from the tree. Investment in tree planting 

is long-term. In this study costs of land were assumed to be nil because it 

can be borrowed from neighbours or allocated from the village chief free 

of charge. Planting could be an alternative in areas of low abundance of 

fruit trees particularly in southern Africa, and if breeding efforts were to 

reduce the age and maturity and significantly increase the yield. 

Table 7.2 Domesticated IFT planting dependent on age to maturity, 

yield increases and collection cost. 

Maturity (years) 2 4 6 8 

Yield increase 

(times non-domesticated level) 

9-30 10-40 12-56 16-80 

Collection cost 

(times level of survey year) 

Source: Mithöfer et al. (2004) 

2.7-3.0 - - - 

66

CHAPTER 8. SELECTION AND 

GENETIC RESOURCES 

8.1 Background 

Unfortunately research on patterns of genetic variation between and 

within populations has not been carried out. Additionally chromosome 

counts are woefully lacking, not only for S. cocculoides, but many of the 

other edible species. 

To-date, phenotypic selection has been used to identify superior 

phenotypes. In order to proceed with this a series of surveys were carried 

out in Botswana, Malawi, Tanzania, Zambia and Zimbabwe using 

participatory appraisal tools and structured questionnaires (Kadzere et 

al., 1998). 

8.2 Surveys 

8.2.1 Botswana 

Veld Products Research and Development organized a series of 

countrywide competitions among school children to identify the largest 

and sweetest fruits. These competitions were a resounding success. 

The provenances which produced the biggest and sweetest fruit on 

healthy trees were selected as baseline materials for germplasm 

collection. Variations in tree height, branching characteristics, time of 

fruiting, fruit production, fruit size and quality were recorded. 

8.2.2 Malawi 

In Malawi, farmers were requested to give preferences and prioritise 

wild fruits found in Malawi (Maghembe et al., 1995). The most 

important criteria used in selecting priority species were the contribution 

made to household food security, market potential, length of fruiting 

period, fruit quantity and quality, ease of management and possibility of 

preserving the fruits. See also Maghembe et al. (1998). 

67

8.2.3 Tanzania 

Ethnobotanical surveys using farmer’s choice and participatory rural 

appraisal tools helped determine the importance of the species to the 

local people based on the criteria of taste, multiple use marketability, 

food security in famine or hunger periods and yield potential. 

(Maghembe et al., 1998) 

8.2.4 Zambia 

The National Institute for Scientific and Industrial Research started work 

on indigenous fruits in 1978. In association with surveys in other 

countries, priority-setting surveys identified tree size and height, fruit 

size, yield and precocity as the characteristics which require 

improvement in S. cocculoides. Bigger fruit was the trait most desired. 

8.2.5 Zimbabwe 

S. cocculoides was the most preferred wild fruit tree (Kadzere et al. 

1998). This could be due to its versatility in processing or its ability to do 

well in marginal areas where rainfall is erratic. Farmers wanted better 

fruit size and yield, taste, flesh-to-seed ratio and fruiting precocity. Some 

farmers, however, did not see the need to improve wild fruit trees and 

preferred them in their present state. From surveys of local communities 

in five districts throughout Zimbabwe it was recommended that S. 

cocculoides could be prioritised for domestication, due to its ease of 

establishment and also for the improvement of the crop (Rukuni and 

Mukwekwerere, 2002). 

8.3 Summary selection criteria 

The surveys gave clear evidence for the following criteria: 

• Good fruit size 

• Sweet pulp taste 

• High flesh to seed ratio 

• Higher yields per tree 

• More rapid tree growth 

• Easily accessible crown 

• Fruiting precocity 

68

8.4 Germplasm 

The use of seed from identified superior trees selected from wild 

populations can produce notable improvement in S. cocculoides. Seed 

orchards are then established using grafts. However, the conditions of 

selection, particularly phenotypic selection from the wild, limit the use of 

these favourable results for predicting the amount of genetic 

improvement to be obtained (Burley and Styles, 1976). There is 

therefore, no firm assurance of genetic improvement in S. cocculoides 

and this can only be achieved by genetic evaluation and selection: 

however, the constraints to this have been noted above. Once sufficiently 

variable germplasm has been collected and the breeding system studied, 

then genetically-based improvement can be put in place. Table 8.1 shows 

the current holdings of Strychnos species in Africa. 

In addition, germplasm is also held by ICRAF in Kenya and institutes 

linked to the ARC Plant Genetic Resources Unit in South Africa. A list 

of the institutions in Table 8.1 and others is to be found in Appendix 1. 

69

70 

Strychnos spinosa HRC, Zimbabwe 

VPR, Botswana 

80 

50 

Seed 

Seed/grafting 

Species Institute Accessions Propagation 

Strychnos cocculoides HRC, Zimbabwe 

30 Seed 

ICRAF, Malawi 

8.5 Conservation 

There is a regional agreement on the transfer of germplasm (especially 

seed, vegetative clones and mycorrhizae, amongst the SADC countries 

and this determines “who pays” and “who benefits” from the germplasm 

and its information. Following an agreed regional plan, each country is 

responsible for collecting and evaluating its own germplasm. All the 

SADC countries should share the germplasm equally on a progressive 

basis. Following regional initiatives to promote Strychnos planting and 

use, ICRAF is acting as a trustee of the germplasm and has no ownership 

rights. 

However, comprehensive storage of genetic resources samples is hardly 

in place. Some aspects to be considered are noted below. 

8.5.1 Methods of storage 

It appears that seeds can be stored when dried to 6.2% and kept at -20ºC 

(Fletcher and Pritchard, 2000). However seed viability can be reduced in 

storage and Uronu and Msanga (2005) recommended that seed be kept at 

10-30% moisture content, above 0 ºC. This is probably going to be most 

useful for storage of planting materials rather than long-term 

conservation. For a genepool naturally distributed over a vast area, in 

which the patterns of diversity are poorly understood, there will be the 

need for targeted ecogeographic collecting and maintenance of materials 

in field genebanks (orchards) complemented by a strategy for in situ 

conservation (Smith et al., 1992). 

8.5.2 In situ conservation 

This will take a two-pronged approach. Firstly, materials will be 

conserved in areas designated as reserves. These will need 

comprehensive documentation with data relevant to Strychnos and its 

improvements being maintained by the relevant national programme. 

Secondly, there will be populations managed intentionally by villagers 

and societies. At present, intentional customary conservation practices 

include widespread social conservation that often prevents the felling of 

fruit-bearing trees and the maintenance of favoured types. Campbell 

(1986) noted this in Zimbabwe; but in considering this to be widespread 

in Africa, Cunningham (1989) stressed that for a resource to be of value 

71

to society it must be of value and must be perceived to be in short supply 

and/or vulnerable to over exploitation. 

72

CHAPTER 9. POTENTIAL 

IMPACT AND MARKETING 

This monograph has shown that there is interest in expanding the 

production of S. cocculoides, and to some degree also S. spinosa. To 

date, technology development has focused on identifying suitable 

germplasm and the promotion of the most suitable methods of 

propagation. This technology development has benefited from a regional 

approach under the guidance of ICRAF and interested donors (e.g. 

SIDA, CIDA, GTZ and DFID). 

This chapter summarises the approaches needed to reach the goal of 

wider cultivation of S. cocculoides and the sustainable use of its 

products, whether fruits or other plant parts. This is followed by a 

discussion of research gaps and what needs to be done to bridge them. 

9.1 Potential for widening the cultivation of S. 

cocculoides 

A major constraint in production of fruits lies in the locally accepted 

concept of tenure. The right to harvest fruits is determined by a complex 

of factors which vary from region to region. In some areas all the trees 

are held to be common property and the fruit is not owned until 

harvested. Such a system reduces incentives for planting of the trees, and 

this problem has to be tackled by extension agents. 

9.1.1 Educating farmers 

Before a farmer will invest in cultivating S. cocculoides the following 

have to be considered: 

• Will the returns on sales outweigh the costs and give a suitable 

profit? 

• What agricultural inputs are required and what are the costs? 

• What is the price for fruits likely to be in 3-4 years time? 

• Will cultivation of the fruit be the most efficient way to use the 

land and what options (e.g. agro-forestry) are available? 

• Will there be continued market potential? 

73

There is more to be done in educating rural people on the short and long 

term benefits of sustainable production of indigenous fruit trees. 

9.1.2 Marketing potential 

The presence of local markets selling S. cocculoides fruits can be used as 

an indicator of the commercial potential of the species. It is important, 

however, to establish the size of the market, the target consumer and 

whether processed products are in demand. Preliminary marketing 

studies conducted in Central and Southern Africa (Minae et al., 1995) 

have shown some interesting gender and age factors. For instance, in 

Malawi 37% of traders are female collectors, 32% are middlemen and 

31% are male collectors. About 63% of the traders were young, being 

less than 30 years old. In most cases the harvesting is done by poor and 

marginalised groups. 

The fruits are currently harvested from farmers’ fields, homestead 

gardens and neighbouring forests. In Malawi, the farmers sell their fruits 

along the roadsides or at local markets. About 20% of the fruit is sold 

directly to consumers, and the remaining 80% is sold to traders. 

The main marketing cost is transportation. Marketing margins indicate 

that the mark-up by traders is usually higher compared to marketing 

costs. This could be due to the current small quantities of fruits handled 

and wastage. 

Consumer perception is also important. Many urban populations 

consider it backward to eat “wild” fruits. In rural areas such fruits may 

be considered to be children’s food or food to be eaten only in times of 

hardship or drought. However, perceptions change. Twenty years ago 

these fruits were not acceptable in Gaborone, Botswana; now there is a 

demand (Taylor, 1995). In South African townships, where people have 

lived with little contact with the countryside, there remains a substantial 

demand due to the perception that traditional foods are strength-giving 

and the white man’s food results in weakness (Taylor, 1983). 

9.1.3 Pricing 

Current pricing and variations in price levels do not accurately reflect the 

supply and demand situation. Competition among suppliers is minimal 

and is not a determinant of price. Extension activities will need to 

support the sales of fruits in markets as an incentive to reinforcing 

74

sustainable production and income distribution. Farmers need to be 

advised on how to develop production in an incremental way without 

putting other parts of the farm system at risk (Arnold, 1996; Scherr, 

1995). 

At the same time, forest departments need to consider policy constraints. 

In markets, farmers can encounter forms of competition and policy 

constraints that can make it difficult for them to compete. In many 

situations, urban markets for most wild fruit products are still supplied 

by harvesting natural stocks, with minute payments to harvesters of the 

raw materials, so that the cost of the product delivered to the market 

consists mainly of transport costs. 

In addition, in many countries, products from state forests and 

plantations can be sold at “administered” prices. Private producers are 

also frequently subjected to produce controls originally designed to 

protect against illegal felling from state forests. The resulting 

bureaucratic procedures lead to private producers having to depend on 

intermediaries to market their produce. Unless impediments can be 

reduced or removed, urban markets are likely to be less important than 

local markets. 

In Botswana the street value for Strychnos was 40 US cents each, giving 

a value of 140 US$ per tree (Taylor et al. 1996). In Zimbabwe the 

average price was 1.5 ZWD per fruit (Mithöfer and Waibel, 2003). 

9.1.4 Markets in the region 

Local markets generally share similar characteristics throughout the 

Eastern and Southern African region, with the possible exception of 

those in Botswana, where informal markets did not exist until after 

Independence (Taylor et al., 1996). Marketing channels for informal 

markets vary according to the product and from place to place, but in 

general, they will still have commonalities. 

Regional and sub-regional markets comprise a level that have received 

very little attention. In these cases shared preferences and common tastes 

often make it feasible to develop products for a larger market. Examples 

of two regional markets are: SADC (Southern African Development 

Community), and EADC (East Africa Development Community). 

It is widely recognised that producers, extension agents and research and 

development for any plant product, have to work against a background 

75

often distorted by policy barriers. High priority has to be accorded by 

official organisations to changing policies and practices which constrains 

farmers’ access to markets for tree products. Removing barriers is 

usually more effective than subsidies to promote tree planting and 

husbandry. 

9.2 Potential for off-farm income based on 

products other than fruits 

9.2.1 Prospects for extraction of chemicals from 

African Strychnos 

The occurrence of chemicals such as strychnine and brucine in Strychnos 

species raises potential interest in use of these and many other 

components such as polysaccharides and saponins for their 

pharmaceutical and industrial uses. Ongoing research, usually in 

developed countries, has a spin-off to local use in Africa especially in 

promotion of traditional medicinal treatments. However, the prospects 

for provision of raw material for extraction are not high. 

Most research appears to focus on the synthesis of the Strychnos 

alkaloids (e.g. Diez et al., 1994) or the testing of such alkaloids for antimalarial 

use, especially in the light of growing resistance to chloroquine. 

In such research, plant sources are taken from all areas of distribution of 

the many Strychnos species. The indolomonoterpenoid alkaloids exist in 

many chemical forms and S. potatorum and S. pungens have been used 

as sources, along with several other African wild species (e.g. Frédérich 

et al., 2002). 

There is less interest in the chemical strychnine as extracted from Asian 

S. nux-vomica. Once used medicinally for its central nervous system 

(CNS)-stimulating properties, to treat chronic heart disease, as a 

stomachic and in the treatment of asthma and epilepsy, it is now only 

used as bait to trap animals. 

9.2.2 Prospects for enhanced uses in local medicine 

In KwaZulu-Natal (South Africa) commercialisation of indigenous 

plants is well developed in the informal sector, with a large and active 

trade. An extensive network exists, which harvests large volumes of 

plants from wild populations throughout the sub-region and distributes 

them to the consumers, who may be both rural or urban. The trade 

76

network has a number of key components, it includes collectors, 

transporters, hawkers, wholesalers, retailers, mail-order companies, 

traditional healers and exporters (Mander et al., 1996). 

In the medicinal plant trade, several large urban street markets exist 

within the Durban metropolitan areas of South Africa and numerous 

stores or informal street sellers are found in almost every town and 

village in the region. Some 60-70% of urban communities are believed to 

make use of traditional medicine. 

However, traditional medicinal uses of Strychnos are still poorly 

documented, and require validation and the development of guidelines 

for the production of standard preparations with known efficacy. This 

requires cooperation between traditional healers, agronomists and local 

universities with capacity for such work. The framework for the 

production and marketing of traditional remedies has been well 

promoted by the World Health Organization, which has highlighted the 

great need for quality control and standardisation of herbals. 

Information that currently exists on the indigenous knowledge of S. 

cocculoides as a medicinal includes old colonial records and the writings 

of early settlers often endlessly repeated. Indigenous knowledge needs to 

be gathered from the communities noting that gender balance in 

gathering information is important, as men and women often value and 

use plants differently, and, even when available, the information can be 

site-specific, depending on ethnic groups and ecogeographic zones. 

Moreover, the success of medicinal products may be affected by 

psychological and cultural factors (Leeson and Frankenberg, 1977) and 

cures used by local healers are often made by mixing products of several 

plants species together with a synergy between them. In some cases a 

synthesis of information already exists, e.g. in the case of treating 

sexually transmitted diseases in Zambia (Ngubani and Hojer, 1999). 

77

CHAPTER 10. RESEARCH NEEDS 

A list of the institutions involved in S. cocculoides research and 

development is shown in Appendix 2. There are currently several 

African countries pursuing cooperative research, with the help of 

international organisations and collaborators in developed countries. 

In essence, the collaboration to date has resulted in research on rapidly 

identifying a limited number of superior forms, and making them 

available as planting materials. Much more collaborative research is 

needed to fill gaps in information and appropriate technology. These are 

grouped in the discussion below. 

10.1.1 Understanding the genepool 

Chapter 1 has pointed out that the taxonomy of the species is imperfectly 

known and it is not yet possible to say which species are closely related 

to the major genepools of S. cocculoides or S. spinosa nor to know how 

other useful species such as S. innocua, S. potatorum, or S. 

madagascariensis, fit into the overall classification and relationships. 

Filling this gap will require extensive field work backed by laboratory 

research on counting chromosomes and Restriction Fragment Length 

Polymorphisms (RFLP) analysis of seedling materials. 

There is a need to understand the patterns of variation in a target 

genepool. This necessitates survey and assessment of patterns of ecogeographical 

variation backed by laboratory analysis using molecular 

methods. The results could be used to identify targets for germplasm 

collecting. Collected germplasm should then be used to obtain 

information on variation within and between populations so that 

materials used for selection would be genetically based. In order for this 

to be as useful as possible, research is needed on the plant’s reproductive 

biology since the extent of inbreeding is not known. 

10.1.2 Developing technology 

Attention has to be given to rooting cuttings, a technique which is not 

successful at present. This would greatly aid germplasm collection and 

permit evaluation trials using specific mother trees as entries. This could 

result in rapid identification of superior genotypes. The biggest gains 

were made in this way for rubber and oil palm and it has been proposed 

78

for perennials such as bamboo. Identification of superior genotypes 

could result in the development of cultivars and this would enhance any 

cooperative work on species improvement. 

More research is needed on the range of agroforestry models which can 

be applied by farmers. 

10.2 Backing the R&D 

Education, particularly of farmers, is essential. The accompanying 

practical manual to this book will be helpful in this respect. Nonetheless 

this cannot take the place of extension which can discuss, within the 

limits of farm production budgets, the most efficient management 

systems for production, distribution and marketing of fruits and/or 

products. 

10.3 Adding value to certain products 

Strychnos products will continue to play an important role in local health 

care. There is a need to assess all available indigenous knowledge and 

current uses by traditional healers and to focus on a limited number of 

remedies especially in relation to primary health care. The collaboration 

of a university department of pharmacology is needed to make 

recommendations on the standard preparation of these remedies. 

There will also be the need to educate collectors of plant products for 

medicine, in association with the healers, to ensure quality material. 

When Strychnos is cultivated, farmers need to be aware of the value 

added for particular plant products and to know the actors in the supply 

chain in order to programme off-farm income. 

10.4 Development of new products 

Technology is in place for the normal processed products from fruits 

(jam, juice, wine, etc.). The major imperative remains the raised 

production of fresh fruits to satisfy demands. However, there may be 

opportunities to develop new products particularly if these can be sold in 

more affluent urban areas. 

79

Apart from herbal remedies (section 9.5 above), there may be cosmetic 

uses and this is certainly the case in parts of Asia for Strychnos (Riswan 

and Sangal-Roemantyo, 1991). 

80

APPENDIX I. INSTITUTIONS 

WITH GERMPLASM OF 

STRYCHNOS COCCULOIDES 

Botswana 

• Department of Integrated Agricultural Research 

Gaborone 

Botswana 

• Veld Products Research and Development 

P.O. Box 2020 

Gaborone 

Botswana 

Malawi 

• Malawi Plant Genetic Resources Centre 

Forestry Research Institute of Malawi 

P.O. Box 270 

Zomba 

Malawi 

South Africa 

• ARC Plant Genetic Resources Unit 

P.O. Box X05, Lynn East 

South Africa 

Tanzania 

• Horticultural Research Institute 

P.O. Box 1253, Tengeru 

Arusha 

Tanzania 

• National Plant Genetic Resources Centre 

P.O. Box 3024 

Arusha 

Tanzania 

81

Zimbabwe 

• SADC – Tree Seed Centre Network 

Private Bag: BW 6238 

Borrowdale 

Harare 

Zimbabwe 

82

APPENDIX II. INSTITUTIONS 

AND INDIVIDUALS ENGAGED IN 

STRYCHNOS RESEARCH AND 

DEVELOPMENT 

Botswana 

• University of Botswana (Baone Kwerepe) 

Botswana College of Agriculture 

P. O. Box 0027 

Gaborone 

Botswana 

• Veld Products Research and Development (Frank W. Taylor, 

Karen J. Butterworth, S.M. Mateke) 

P.O. Box 2020 

Gaborone 

Botswana 

Germany 

• Institute of Economics in Horticulture 

Faculty of Business Administration and Economics 

Hannover University 

Germany 

Israel 

• Institute for Agriculture and Applied Biology 

Ben Gurion University of the Negev (G. Bohrer, V. Kugan Zur, 

Y. Mizhrahi) 

Israel 

Kenya 

• World Agroforestry Centre (ICRAF) 

PO Box 30677 

00100 GPO 

Nairobi 

Kenya 

icraf@cgiar.org 

83

Malawi 

• Department of Forestry 

P.O. Box 30048 

Zomba 

Malawi 

• Forest Research Institute of Malawi (L. A. Sitaubi, L. 

Mwabumba) 

P. O. Box 270 

Zomba 

Malawi 

• ICRAF – Agroforestry Research Project (S. Minae) 

P.O. Box 31188 

Lilongwe 3 

Malawi 

• ICRAF, SADC Agro-forestry Project (F.K. Akinnifesi, J. Moyo) 

P.O. Box 134 

Zomba 

University of Malawi 

Malawi 

• Bunda College of Agriculture (M. Kwapata, S.S. Munthali) 

P.O. Box 219 

Lilongwe 

Malawi 

South Africa 

• Agricultural Research Council of South Africa 

Institute for Tropical and Sub-tropical Crops 

Nelspruit – 1200 

South Africa 

• Plant Genetic Resources (PGR) Unit 

P.O. Box X05, Lynn East, 0039 

South Africa 

Swaziland 

• Malkerns Research Station 

P.O. Box 4, 

Malkerns 

84

Swaziland 

Sri Lanka 

• International Centre for Underutilised Crops 

International Water Management Institute 

127 Sunil Mawatha 

Pelawatte 

Battaramulla 

Sri Lanka 

Zambia 

• Division of Forestry Research 

P.O. Box 22099 

Kitwe 

Zambia 

• Tree Improvement Research Centre 

National Institute for Scientific and Industrial Research (NISIR) 

P. O. Box 21210 

Kitwe 

Zambia 

• Zambia/ICRAF Agroforestry Project (P. L. Mafongoya, S. 

Lungu, S. Mwanza) 

Msekera Agricultural Research Station 

P. O. Box 510046 

Chipata 

Zambia 

Zimbabwe 

• Department of Research and Specialist Services (E.Nyamutowa, 

J. N. Mushonga) 

Horticultural Research Centre 

P.O. Box 3748 

Marondera 

Zimbabwe 

85

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96

agroforestry....11, 20, 60, 61, 79 

bark. 10, 7, 9, 11, 12, 13, 14, 27, 

28, 31, 60 

breeding.........11, 62, 66, 69, 86 

chemicals...............................76 

clay ............................47, 48, 51 

climate ...............................9, 43 

conservation.....4, 66, 71, 86, 93 

cowpea...................................61 

cutting................................4, 10 

diseases......................28, 60, 77 

distribution....10, 16, 19, 21, 47, 

60, 75, 76, 79 

economics..............................64 

ecotypes...........................52, 57 

environment.....................16, 54 

establishment .......57, 59, 90, 91 

fertilizers..........................58, 59 

flowers...............................7, 62 

fruits 10, 11, 3, 5, 10, 19, 22, 25, 

26, 27, 28, 29, 40, 44, 54, 55, 

59, 61, 62, 63, 64, 65, 66, 67, 

68, 73, 74, 76, 79, 87, 89, 91, 

92, 93 

genepool ..........................71, 78 

genetic resources ...................71 

germination..........56, 57, 60, 89 

germplasm 9, 57, 67, 69, 70, 71, 

73, 78 

grafting ......................56, 57, 70 

grasses .............................16, 57 

habitat ..............................52, 58 

leaf ...............4, 9, 27, 28, 46, 90 

light....44, 46, 55, 57, 59, 60, 76 

Loganiaceae...............1, 2, 3, 90 

marketing.12, 63, 64, 74, 77, 79 

medicine ................1, 76, 77, 79 

mildew.......................10, 40, 63 

INDEX 

97 

mycorrhizae...............58, 61, 71 

nutrient.............................48, 51 

pests.......................................60 

planting... 21, 52, 57, 58, 59, 66, 

71, 73, 76, 78 

pollination..............................55 

products .... 4, 25, 63, 65, 66, 73, 

74, 75, 76, 77, 79, 90, 92, 94, 

95 

propagation... 20, 55, 56, 57, 70, 

73 

pruning.............................58, 59 

rainfall ..... 16, 28, 42, 44, 51, 68 

RFLP .....................................78 

ripening............................44, 62 

root ... 10, 28, 39, 44, 46, 52, 58, 

60, 92 

S. cocculoides ..................49, 50 

S. innocua10, 11, 12, 14, 15, 20, 

23, 25, 26, 27, 28, 78 

S. madagascariensis..10, 11, 12, 

15, 20, 26, 28, 78 

S. potatorum . 10, 13, 14, 19, 26, 

29, 76, 78 

S. pungens 9, 10, 12, 14, 15, 24, 

25, 26, 28, 29, 44, 47, 48, 76 

S. spinosa 9, 1, 9, 14, 15, 16, 19, 

20, 22, 24, 25, 26, 27, 28, 29, 

44, 48, 49, 50, 61, 62, 73, 78 

Sclerocarya birrea 44, 45, 53, 61 

seed... 13, 26, 54, 55, 56, 60, 62, 

68, 69, 71, 89 

selection.. 11, 20, 60, 67, 68, 69, 

78 

shade.... 10, 4, 36, 44, 45, 46, 56 

soil 9, 16, 46, 47, 48, 49, 50, 51, 

57, 58, 61, 63 

sorghum.................................61

storage ...........40, 56, 60, 63, 71 

Strychnaceae........................2, 3 

taxonomy ...............................78 

timber.............65, 86, 88, 90, 95 

Uapaca kirkiana..10, 26, 41, 92, 

94 

98 

Vangueria infausta ..........53, 61 

water. 16, 27, 44, 46, 56, 59, 63, 

90 

watermelon ............................61 

wind.......................................46 

yield.................................66, 68

Monkey Orange monograph.pdf - Crops for the Future

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