list of figures - Terry Sunderland

THE TAXONOMY, ECOLOGY AND UTILISATION OF 

AFRICAN RATTANS 

(PALMAE: CALAMOIDEAE) 

A Thesis submitted for the degree, Doctor of Philosophy (PhD) 

by 

Terence Christopher Heesom Sunderland 

2001 

University College, 

London

ABSTRACT 

THE TAXONOMY, ECOLOGY AND UTLISATION OF AFRICAN RATTANS 

(PALMAE: CALAMOIDEAE) 

This thesis presents the findings of a multi-disciplinary study of the taxonomy, 

ecology and utilisation of the rattans of the lowland tropical forests of Africa. 

Based on extensive field-work and herbarium study, and through the application of a 

morphological species concept, a taxonomic revision is presented. Twenty species, 

including two new species, representing four rattan genera, are described and 

illustrated. 

Ecological studies undertaken as part of this study allows assessments to be made of 

edaphic and climatic factors affecting rattan diversity and abundance. Further field 

study of the life history of these taxa has clarified the occurrence of hapaxanthy and 

pleonanthy within the rattan genera. Additional ecological work has focussed on the 

interaction of rattan species with their wider environment. In particular, the complex 

relationship between forest fauna and rattan is presented in detail. 

The findings of extensive ethnobotanical surveys of rattan use by selected local 

communities are presented. These surveys conclude that very few of the known 

species are actually of any significant use value. This study of local usage enables a 

conceptual framework of the indigenous classification systems for rattan, employed 

by forest-based communities, to be discussed and presented. 

A brief overview of the wider rattan trade in Africa highlights the economic 

importance of rattan as a forest resource. A more detailed analysis of the socioeconomic 

nature of the formal rattan trade Cameroon concludes that, whilst 

contributing significantly to the forest economy, the uncontrolled nature of the trade 

is leading to an increasingly scarce resource base.

The final Chapter presents summarises the findings of the thesis and discusses the 

framework for the sustainable exploitation of rattan in Africa. The ecological, social 

and institutional criteria that need to be met before strategies for sustainability can be 

implemented are discussed in detail.

LIST OF CONTENTS 

ACKNOWLEDGEMENTS 17 

INTRODUCTION 20 

TAXONOMY 

CHAPTER ONE 

MORPHOLOGY AND BIOGEOGRAPHY OF AFRICAN RATTANS 

1.1 Introduction 21 

1.2 Morphology 21 

1.2.1. Growth form 21 

1.2.2. The stem 24 

1.2.3. Cane anatomy 25 

1.2.4. The root system 26 

1.2.5. Rattan leaves 27 

1.2.6. The sheath 28 

1.2.7. Sheath armature 28 

1.2.8. Indumentum 29 

1.2.9. The ocrea 29 

1.2.10. Sheath striations 29 

1.2.11. The knee 30 

1.2.12. The petiole and leaf rachis 30 

1.2.13. Leaflets 31 

1.2.14. Climbing organs 32 

1.2.15. Flowering behaviour 33 

1.2.16. The inflorescence and floral morphology 34 

1.2.17. Calyx and corolla 36 

1.2.18. Stamens 36 

1.2.19. Pollen 36 

1.2.20. The fruit 36 

1.2.21. The seed 37 

1.2.22. Germination 38 

1.3. Fossil record and biogeography 38 

4

CHAPTER TWO 

TAXONOMIC ACCOUNT 

2.1. Introduction 41 

2.2. History of classification 41 

2.3. The wider taxonomic picture 44 

2.4. Species concepts 46 

2.5. Key to the genera 47 

Eremospatha (G. Mann & H. Wendl.) H. Wendl. 48 

Key to the species of Eremospatha 50 

E. hookeri (G. Mann & H. Wendl.) H. Wendl. 52 

E. cabrae de Wild. 56 

E. laurentii de Wild. 61 

E. wendlandiana Dammer ex Becc. 66 

E. barendii sp. nov. 70 

E. macrocarpa (G. Mann & H. Wendl.) H. Wendl. 74 

E. haullevilleana de Wild. 80 

E. tessmanniana Becc. 86 

E. cuspidata (G. Mann & H. Wendl.) H. Wendl. 88 

E. quinquecostulata Becc. 91 

Excluded names and nomina nuda 95 

Laccosperma (G. Mann & H. Wendl.) Drude 97 

Key to the species of Laccosperma 99 

L. opacum (G. Mann & H. Wendl.) Drude 100 

L. laeve (G. Mann & H. Wendl.) H. Wendl. 106 

L. acutiflorum (Becc.) J. Dransf. 109 

L. robustum (Burr.) J. Dransf. 113 

L. secundiflorum (P. Beauv.) Kuntze 119 

Imperfectly-known taxon 127 

Excluded names and nomina nuda 128 

Oncocalamus (G. Mann & H. Wendl.) H. Wendl. 129 

Key to the species of Oncocalamus 131 

O. mannii (H. Wendl.) H. Wendl. 132 

O. macrospathus Burr. 136 

O. tuleyi sp. nov. 140 

5

O. wrightianus Hutch. 144 

Imperfectly-known taxon 147 

Calamus L. 149 

C. deërratus G. Mann & H. Wendl. 150 

Mixed collections 159 

ECOLOGY 

CHAPTER THREE 

RATTAN DIVERSITY AND ABUNDANCE: A COMPARISON BETWEEN 

THREE FOREST SITES IN CAMEROON 


3.2. Rattan inventory 170 

3.2.1. Introduction 170 

3.2.2. Inventory parameters 170 

3.2.2.1. Introduction 170 

3.2.2.2. Taxonomy 171 

3.2.2.3. Stem length 171 

3.2.2.4. Plot shape and size 172 

3.3. Research sites 173 

3.3.1. Campo Ma’an Faunal Reserve 174 


3.3.1.2. Climate 175 

3.3.1.3. Topography, geology and soil type 175 

3.3.1.4. Vegetation 175 

3.3.1.5. Forest exploitation 176 

3.3.2. Mokoko River Forest Reserve 176 


3.3.2.2. Climate 177 




3.3.3. Takamanda Forest Reserve 179 


6

3.3.3.2. Climate 180 




3.4. Methods 182 

3.4.1. Sampling 182 

3.4.2. Plot establishment 183 

3.4.3. Mapping 184 

3.4.4. Enumeration 184 

3.5. Presentation of results 185 

3.5.1. Cumulative summary of vegetative plots 185 

3.5.2. Summary of rattan stocking and abundance by site 186 

3.6. Discussion 187 

3.6.1. Floristic diversity and similarity 187 

3.6.2. Rattan diversity and similarity 188 

3.6.3. Total stem length vs harvestable stem length 191 

3.6.4. Inventory techniques and species capture 192 

3.7. Conclusion 193 

CHAPTER FOUR 

RATTAN / FAUNAL RELATIONSHIPS 


4.2. Herbivory 196 

4.2.1. Stem apex or “palm heart” 196 

4.2.2. Leaflet predation 197 

4.3. Seed dispersal 198 

4.4. Seed predation and caching 201 

4.5. Ant / rattan associations 202 

4.6. Summary 207 

CHAPTER FIVE 

HAPAXANTHY AND PLEONANTY IN AFRICAN RATTANS 


5.2. Hapaxanthy and pleonanthy: a discussion 210 

7

5.3. Geographical distribution of hapaxanthy 212 

5.4. The African genera 213 

5.4.1. Laccosperma 213 

5.4.2. Eremospatha 213 

5.4.3. Oncocalamus 214 

5.4.4. Calamus 215 

5.5. Ecology and evolution 215 

5.6. Summary 217 

UTILISATION 

CHAPTER SIX 

INDIGENOUS NOMENCLATURE, CLASSIFICATION AND UTILISATION OF 

AFRICAN RATTANS 


6.2. Indigenous utilisation of African rattans 220 

6.3. Brief introduction to language and history in Africa 224 

6.4. Berlin’s model of ethnobiological classification 225 

6.4.1. Introduction 225 

6.4.2. The unique beginner 226 

6.4.3. Life form 227 

6.4.4. Intermediates 227 

6.4.5. Generics 227 

6.4.6. Specific and varietal categories 228 

6.5. Methods 228 

6.6. African rattan nomenclature with reference to Berlin’s model 229 

6.6.1. The unique beginner or kingdom 229 

6.6.2. Life form categories 229 

6.6.3. Intermediate and generic categories 230 

6.6.4. Specific categories 233 

6.6.5. Varietal categories 235 



8

CHAPTER SEVEN 

A SOCIO-ECONOMIC PROFILE OF THE COMMERCIAL RATTAN TRADE IN 

CAMEROON 


7.2. The markets for NTFPs in Central Africa 243 

7.3. The rattan trade in Cameroon 244 

7.4. Methodology 245 

7.4.1. Selection of study sites 245 

7.4.2. Sampling methodology 247 

7.4. The rattan sector in Cameroon 248 

7.5.1. Rattan harvest and supply to urban markets 248 

7.5.1.1. The resource base 248 

7.5.1.2. Customary laws and State legislation 248 

7.5.1.3. Production to consumption system 249 

7.5.1.4. Range 250 

7.5.1.5. Frequency of supply and purchase 251 

7.5.2. Rattan artisan enterprises 252 


7.5.2.2. Number of workers and sources of labour 252 

7.5.2.3. Types of enterprise 254 

7.5.3. Socio-economic profile of urban artisans 255 

7.5.3.1. Age range 255 

7.5.3.2. Gender 255 

7.5.3.3. Ethnicity 255 

7.5.3.4. Educational level of rattan artisans 256 

7.5.3.5. Previous occupations 257 

7.5.4. The scale of the trade 258 

7.5.4.1. Amounts and values 258 

7.5.4.2. Profitability 258 

7.5.5. Nature of the trade 259 

7.5.5.1. Decline or growth? 259 


7.6.1. “La crise” and increased reliance on forest products 262 

7.6.2. Sustainability 264 

9

7.6.3. Increased range = increased price? 264 

7.6.4. Profitability and size of urban market 265 

7.6.5. An industry in decline? 266 

7.6.6. Rattan unions 267 


CHAPTER EIGHT 

A BRIEF OVERVIEW OF THE RATTAN TRADE IN AFRICA 

8.1. The international economic context 273 

8.2. The African rattan trade 274 

8.3. The resource base 275 

8.4. The nature of the trade 275 

8.5. Sustainability issues 277 

8.6. Amount and value of the trade 278 


SUMMARY AND CONCLUSIONS 

CHAPTER NINE 

IMPLICATIONS FOR CONSERVATION AND DEVELOPMENT 


9.2. Conservation status of African rattans 281 

9.3. Harvest and management 282 

9.3.1. Growth rates 282 

9.3.2. Management 283 

9.3.2.1. Natural regeneration in high forest 283 

9.3.2.2. Enhanced natural regeneration 283 

9.3.2.3. Agroforestry systems 283 

9.3.2.4. Silvicultural systems 284 

9.3.3. Harvest procedures 284 

9.3.5. Inventory 285 

9.4. Land tenure and socio-economic issues 285 

9.5. Rattan cultivation 286 


10

REFERENCES 290 

APPENDIX ONE 

INDIGENOUS NOMENCLATURE AND UTILISATION OF AFRICAN 

RATTANS BY SPECIES 320 

APPENDIX TWO 

LIST OF EXSICCATAE 337 

APPENDIX THREE 

RATTAN RESEARCH PLOT No. 1: CAMPO FAUNAL RESERVE 349 

APPENDIX FOUR 

SOCIO-ECONOMIC QUESTIONNAIRE 352 

APPENDIX FIVE 

PUBLICATIONS AND DISSEMINATION 356 

LIST OF FIGURES 

Figure 1. Mature clustering stem of Laccosperma robustum (Burr.) 

J. Dransf. 22 

Figure 2. Aerial branching of Eremospatha hookeri (G. Mann & 

H. Wendl.) H. Wendl. 23 

Figure 3. Acanthophylls of Laccosperma acutiflorum (Becc.) J. Dransf. 33 

Figure 4. Elaminate rachis of Oncocalamus tuleyi sp. nov. 34 

Figure 5. Diagram of selected dyads within the Calamoideae 35 

Figure 6. Adjacent-ligular germination of Oncocalamus mannii (H. Wendl.) 

H. Wendl. 38 

Figure 7. Pleistocene refugia and the distribution of African rattans 40 

Figure 8. Schematic tree showing relationships with the Calamoideae 45 

Figure 9. Eremospatha hookeri (G. Mann & H. Wendl.) H. Wendl. 54 

Figure 10. Distribution of E. hookeri (G. Mann & H. Wendl.) H. Wendl. 55 

11

Figure 11. E. cabrae de Wild. 59 

Figure 12. Distribution of E. cabrae de. Wild. 60 

Figure 13. E. laurentii de Wild. 63 

Figure 14. Distribution of E. laurentii de Wild. 64 

Figure 15. E. wendlandiana Dammer ex Becc. 68 

Figure 16. Distribution of E. wendlandiana Dammer ex Becc. 69 

Figure 17. E. barendii sp. nov. 72 

Figure 18. Distribution of E. barendii sp. nov. 73 

Figure 19. E. macrocarpa (G. Mann & H. Wendl.) H. Wendl. 

Figure 20. Distribution of E. macrocarpa (G. Mann & H. Wendl.) 

76 

H. Wendl. 77 

Figure 21. E. haullevilleana de Wild. 82 

Figure 22. Distribution of E. haullevilleana de Wild. 83 

Figure 23. Distribution of E. tessmanniana Becc. 87 

Figure 24. Distribution of E. cuspidata (G. Mann & H. Wendl.) H. Wendl. 89 

Figure 25. E. cuspidata (G. Mann & H. Wendl.) H. Wendl. 90 

Figure 26. Distribution of E. quinquecostulata Becc. 92 

Figure 27. E. tessmanniana Becc. & E. quinquecostulata Becc. 94 

Figure 28. Distribution of Laccosperma opacum (G. Mann & H. Wendl.) 

Drude 102 

Figure 29. L. opacum (G. Mann & H. Wendl.) Drude & 

L. laeve (G. Mann & H. Wendl.) H. Wendl. 

105 

Figure 30. Distribution of L. laeve (G. Mann & H. Wendl.) H. Wendl. 108 

Figure 31. L. acutiflorum (Becc.) J. Dransf. 111 

Figure 32. Distribution of L. acutiflorum (Becc.) J. Dransf. 112 

Figure 33. L. robustum (Burr.) J. Dransf. 116 

Figure 34. Distribution of L. robustum (Burr.) J. Dransf. 117 

Figure 35. L. secundiflorum (P. Beauv.) Kuntze 122 

Figure 36. Distribution of L. secundiflorum (P. Beauv.) Kuntze 123 

Figure 37. Oncocalamus mannii (H. Wendl.) H. Wendl. 134 

Figure 38. Distribution of O. mannii (H. Wendl.) H. Wendl. 135 

Figure 39. O. macrospathus Burr. 138 

Figure 40. Distribution of O. macrospathus Burr. 139 

Figure 41. O. tuleyi sp. nov. 142 

12

Figure 42. Distribution of O. tuleyi sp. nov. 143 

Figure 43. Distribution of O. wrightianus Hutch. 145 

Figure 44. O. wrightianus Hutch. 146 

Figure 45. Calamus deërratus G. Mann & H. Wendl. 154 

Figure 46. Distribution of C. deërratus G. Mann & H. Wendl. 155 

Figure 47. Eremospatha hookeri, Korup N.P., Cameroon 160 

Figure 48. E. laurentii, Campo, Cameroon 160 

Figure 49. E. laurentii, Campo, Cameroon 160 

Figure 50. E. laurentii, Cogo, Equatorial Guinea 160 

Figure 51. E. macrocarpa, Rumpi Hills, Cameroon 161 

Figure 52. E. macrocarpa, Limbe Botanic Garden, Cameroon 161 

Figure 53. E. macrocarpa, ocrea, Limbe Botanic Garden, Cameroon 161 

Figure 54. E. macrocarpa, Mamfe, Cameroon 161 

Figure 55. E. macrocarpa flowers, Mamfe, Cameroon 162 

Figure 56. E. macrocarpa fruits, Kribi, Cameroon 162 

Figure 57. E. wendlandiana, Southern Bakundu, Cameroon 162 

Figure 58. E. wendlandiana knee, Campo, Cameroon 162 

Figure 59. E. cuspidata, Etembue, Equatorial Guinea 163 

Figure 60. E. cuspidata fruits, Etembue, Equatorial Guinea 163 

Figure 61. L. opacum fruits, Nguti, Cameroon 163 

Figure 62. L. laeve fruits, Draw River, Ghana 163 

Figure 63. L. acutiflorum, Nguti, Cameroon 164 

Figure 64. L. acutiflorum, Campo, Cameroon 164 

Figure 65. L. robustum, Campo, Cameroon 164 

Figure 66. L. robustum fruits, Bata, Equatorial Guinea 164 

Figure 67. L. secundiflorum, Tarkwa, Ghana 165 

Figure 68. L. secundiflorum, Nigeria 165 

Figure 69. O. macrospathus, Evinayong, Equatorial Guinea 165 

Figure 70. O. macrospathus fruits, Evinayong, Equatorial Guinea 165 

Figure 71. O. tuleyi, Rumpi Hills, Cameroon 166 

Figure 72. O. tuleyi, Southern Bakundu, Cameroon 166 

Figure 73. O. tuleyi sheath, Southern Bakundu, Cameroon 166 

Figure 74. O. tuleyi, Limbe to Kumba road, Cameroon 166 

Figure 75. O. mannii, Ayameken, Equatorial Guinea 167 

13

Figure 76. O. mannii inflorescences, Etembue, Equatorial Guinea 167 

Figure 77. O. mannii, juvenile, Basile, Equatorial Guinea 168 

Figure 78. O. mannii, inflorescence, Etembue, Equatorial Guinea 168 

Figure 79. C. deërratus, Mungo Bridge, Cameroon 168 

Figure 80. C. deërratus, inflorescences, Mungo Bridge, Cameroon 168 

Figure 81. Map of Cameroon showing protected areas 174 

Figure 82. One hectare plot layout 184 

Figure 83. Cumulative size-class distribution from survey sites 185 

Figure 84. Correlation: total stem length and harvestable stem length 191 

Figure 85. Ant colonisation of Laccosperma laeve 209 

Figure 86. Ant farming of scale insects on Eremospatha hookeri 209 

Figure 87. Hapaxanthy and pleonanthy in palms 212 

Figure 88. Inflorescences of E. cuspidata 214 

Figure 89. Inflorescences of O. mannii 215 

Figure 90. Hapaxanthic inflorescence of L. acutiflorum 217 

Figure 91. Berlin’s theoretical model of ethnobiological classification 226 

Figure 92. Temporary forest camp for rattan harvesting 241 

Figure 93. Cut stems of Laccosperma robustum 241 

Figure 94. Fish trap 242 

Figure 95. Cane bridge 242 

Figure 96. Weaving of farm basket 242 

Figure 97. Market basket 242 

Figure 98. Map of southern Cameroon showing survey sites 246 

Figure 99. Scatterplot of size of market and distance of supply 247 

Figure 100. Production to consumption rattan flow in Cameroon 250 

Figure 101. Mean range (or distance) of rattan supplies 251 

Figure 102. Urban prices paid per unit 251 

Figure 103. Frequency of wholesale rattan purchase 252 

Figure 104. Summary of rattan products 253 

Figure 105. Number of workers per enterprise 253 

Figure 106. Types of rattan workshops in Cameroon 254 

Figure 107. Age range of the primary owner / worker 255 

Figure 108. Educational level of urban artisans 256 

Figure 109. Previous occupations of rattan artisans 257 

14

Figure 110. How many years spent in the trade? 257 

Figure 111. Mean monthly profits of rattan artisans 259 

Figure 112. How much cane is used now compared with 5 years ago 259 

Figure 113. General consumption trend 260 

Figure 114. Reasons for decline in rattan sector 261 

Figure 115. Reasons for growth in rattan sector 261 

Figure 116. Correlation between range and costs of raw cane 265 

Figure 117. Open-air rattan enterprise in Bata, Equatorial Guinea 270 

Figure 118. Enclosed permanent rattan enterprise in Bata 270 

Figure 119. Woven baskets for sale on roadside in Cameroon 271 

Figure 120. Finished rattan products for sale in Bata 271 

Figure 121. Woven rattan products for sale on roadside in Cameroon 272 

Figure 122. Finished rattan products for sale in Bata 272 

Figure 123. Scale and value of rattan trade in selected markets 279 

Figure 124. Cultivated system of Calamus merrillii in Sabah 287 

LIST OF TABLES 

Table 1. African rattan species producing aerial branches 24 

Table 2. Vegetation summary for ecological survey sites 185 

Table 3. Rattan abundance and stocking for Campo 186 

Table 4. Rattan abundance and stocking for Mokoko 186 

Table 5. Rattan abundance and stocking for Takamanda 187 

Table 6. Floristic similarity between survey sites 188 

Table 7. Floristic similarity between study sites for rattans 189 

Table 8. Comparison of rattan stocking by site 189 

Table 9. Sampling and representation 192 

Table 10. Comparison of rattan diversity, abundance and stocking 193 

Table 11. Effects on seed survival after primate predation 200 

Table 12. Herbivory and dispersal of African rattans 201 

Table 13. Rattan hosts and ant colonies 205 

Table 14. Count of domatia and ant species 206 

Table 15. Count of domatia and rattan species 207 

Table 16. Hapaxanthy in the Palmae 213 

15

Table 17. Taxonomy and utilisation of the rattans of Africa 223 

Table 18. Summary of non-cane uses of African rattans 224 

Table 19. Life form, intermediate and generic categories 232 

Table 20. Selected cane product names 233 

Table 21. Folk specific taxa 235 

Table 22. Summary of rattan classification 236 

Table 23. Parity between western and local classification systems 237 

Table 24. Study sites and sampling 248 

Table 25. Percentage of artisans who harvest their own cane 250 

Table 26. Amount and value of cane trade in Cameroon 258 

Table 27. Major constraints to development of rattan sector 262 

Table 28. Recommendations for stimulation of sector 262 

Table 29. Raw cane exports from Cameroon, 1926-29 274 

Table 30. Commercially important rattan species by region 275 

Table 31. Constraints to the development of the African rattan sector 278 

Table 32. Scale and value of rattan trade in selected markets 279 

Table 33. Conservation status of African rattans 282 

Table 34. Growth rates of commercial rattans 282 

LIST OF BOXES 

Box 1. Nomenclatural relationships within the Denya language group 234 

Box 2. The structure of vernacular names 236 

Box 3. Permis d’exploitation 249 

16

ACKNOWLEDGEMENTS 

This work would not have been possible without the assistance and encouragement of 

a great many individuals and institutions. The greatest share of gratitude must go to 

John Dransfield of the Royal Botanic Gardens, Kew and Phil Burnham of the 

Department of Anthropology at University College, London. Dr Dransfield initially 

identified the need for this study and helped develop the work into a coherent (and 

fundable) research programme. His encouragement and inspiration during the course 

of this research has been fantastic. Prof. Burnham provided a great deal of assistance 

and guidance throughout the course of this study and encouraged me to extend my 

research beyond my formal botanical training to make this study as multi-disciplinary 

as I hope it is. Barrie Goldsmith, also of UCL, provided useful guidance and 

comments. 

A great many collaborators provided a huge amount of support and encouragement for 

this study, often beyond the call of professional duty. In Cameroon I would 

particularly like to thank: James Culverwell and Zachary Akum formerly of the Korup 

Project, Nouhou Ndam, Joseph Nkefor and Paul Blackmore of the Limbe Botanic 

Garden, Bahiru Daguma, Tony Simons and Zach Tchoundjeu of ICRAF, Nicodeme 

Tchamou of CARPE, David Mbah of MINREST, Johnson Jato and Augustine 

Njiamnshi of BDCP, Guillaume Akogo Mvogo of the Campo Ma’an Project, Raphael 

Ebot of MINEF Manyu Division, Henry Njalla Quan and Richard Grey of CDC, Steve 

Gartlan of WWF, Anacletus Koufani, Benoit Sabatier and his successor Georges 

Achoundong of the National Herbarium, Louis Defo of APFT and all the staff of 

MINEF and MINREST who have helped with the provision of research permits, 

phytosanitary certificates and other formal documentation over the years. In 

Equatorial Guinea, particular thanks go to Frank Stenmanns, formerly of the CUREF 

Project, for enabling and funding me to undertake my study there and to Crisantos 

Obama who has proved to be an enthusiastic long-term collaborator. In Nigeria, 

thanks to Maurice Iwu of BDCP and Tunde Morakinyo, formerly of Living Earth for 

invaluable advice, support and encouragement during field work there. In Ghana, 

Andrew Oteng-Amoako of FORIG provided logistical support and research 

17

collaboration, as did Emmanuel Ebanyele. Dr Henry Borobou Borobou and Jean- 

Pierre Profizi in Gabon provided access to historical collections and literature. 

A special thank you also to all the Chiefs, Council Members, harvesters, artisans, 

guides, and other members of the communities I have worked with in the past few 

years on this project who have provided hospitality, assistance, support and good 

humour in copious quantities. 

At Kew, I would like to thank Bill Baker for his sound advice during the writing of the 

taxonomic account, and to Lucy Smith who tirelessly produced the wonderful 

botanical illustrations included within this thesis in such a short space of time. Thanks 

are also due Colin Clubbe for his enthusiastic support during my year or so there and 

for administering the INBAR funds so efficiently. Tom Evans, Leng Saw Guan, and 

Hiroshi Ehara provided much stimulating discussion in the Palm Room. 

In the US, Stefan Cover of Harvard University identified the ant samples for me, for 

which I am grateful, whilst Laurent Some and Rose-Marie Gay administered the 

CARPE funds which kept the field work running (almost) to schedule. Thanks also to 

Michael Brown for funding the Mokoko NTFP study. Lisa Jordan and Katy Moran in 

Washington provided advice, tea and considerable sympathy. 

In Malaysia, Supardi Mohammed Nur and Lee Ying Fah proved to be extremely 

informative hosts during a rewarding rattan study tour to both Peninsular Malaysia 

and Sabah, Borneo and helped shape my thinking with regard to the sustainable 

management of the rattan resource in Africa. 

Special mention goes to Steve Ruddy of the Garden Development Unit of the Royal 

Botanic Gardens, Kew who provided extensive advice on “all things computer” and 

Jim Comiskey of the Smithsonian Institution who provided extremely helpful 

guidance on statistics and data analysis. To both of them, I am particularly grateful. 

To undertake long-term field work in West and Central Africa entails a great deal of 

financial support, as does the process of sitting and writing up the results of those 

three years in the field. To that end I am particularly grateful to Mark Buccowich, who 

18

ecommended this project be funded by the Office for International Programmes of 

the United States Forest Service (USFS) and the Central African Regional Programme 

for the Environment (CARPE). Without Mark’s vision and commitment, this work 

would never have been able to be completed. In this same vein, a great many thanks 

are also due to Cherla Sastry, formerly Director of the International Network for 

Bamboo and Rattan (INBAR) who saw the urgent need for a coherent taxonomy of 

the African rattans and agreed to fund the writing of the taxonomic account from 

March 1999 to February 2000. 

Thanks are also due to the curators and directors of the following herbaria who 

allowed me to visit or undertake loans: BM, BR, EG, FHO, FI, G, GC, HBG, K, KRI, 

KUM, LBR, MO, NY, SCA, WAG, YA. 

Things change; people move on and people do not live forever. This work is dedicated 

to Dinga Njingum Franklin, wherever he may now be, and to the memory of Mukete 

Wilfred (1967-1999) who both worked with me in the field over the course of many 

years in Cameroon. Now I am back in the field, their presence is greatly missed. 

Finally, I would like to extend my profound and deep gratitude to my long-suffering 

wife, Jacqui, who has lived with the rattans of Africa for almost as long as I have and 

who willingly sacrificed a year of her own (much more exciting) field work to 

accompany me to the UK to enable me to write up the results of this study. 

Thanks J; my turn! 

19

INTRODUCTION 

Throughout the lowland tropical forest zone of Africa, the climbing palms, or rattans 

(Palmae: Calamoideae) form an integral part of subsistence strategies for many rural 

populations and provide the basis of a thriving cottage industry. 

In this regard, African rattans have long been recognised by donor agencies and 

national governments as having a potential role to play within the domestic and 

regional non-timber forest product (NTFP) sectors, as well as within the significant 

global rattan market. As increased interest is being shown in the potential role of such 

high-value NTFPs to contribute to the paradigm of conservation and development, 

rattan has been one of the oft-mentioned products that could be developed and 

promoted in a meaningful way. However, the development of the rattan resource in 

Africa, unlike that of Asia, has long been hindered by a lack of basic knowledge about 

the exact species utilised, their ecological characteristics and requirements and the 

social context of their utilisation. 

This thesis presents the findings of a four-year study of the rattans of Africa. Funded 

by the United States Forest Service (USFS), the Central African Regional Programme 

for the Environment (CARPE) and the International Network for the Development of 

Bamboo and Rattan (INBAR), this research has concentrated on the study of the basic 

taxonomy, ecology and utilisation of these taxa. The provision of such baseline 

research is critical for the development and promotion of any high value NTFP within 

the framework of forest-based conservation and development initiatives. The 

information presented in this thesis may, it is hoped, feasibly provide the ecological, 

social and institutional contexts in which appropriate management strategies might be 

developed and implemented to ensure the sustainable, and equitable, exploitation of 

the rattans of Africa. 

20

CHAPTER ONE 

MORPHOLOGY AND BIOGEOGRAPHY OF AFRICAN RATTANS 

1.1 INTRODUCTION 

The rattans of Africa are represented by the endemic genera Laccosperma, 

Eremospatha and Oncocalamus, as well as by a single representative of the Asian 

genus Calamus. These climbing palms occur in a wide range ecological conditions 

within the lowland tropical forests of the continent and, throughout their range, play a 

significant role in the forest economy of the region through the utilisation of the inner 

stems, or cane. Despite this economic importance there has persisted considerable 

uncertainty with regard to the natural history of these species. This Chapter discusses 

the morphology and wider natural history of these species and provides the basis of 

the taxonomic revision presented in Chapter 2. 

1.2 MORPHOLOGY 

1.2.1 Growth form 

All the rattan species of Africa are clustering i.e. they produce numerous stems from 

one individual. The suckers that develop to form a cluster are produced from the 

lowermost nodes of the original stem. They, in turn, produce further suckers and a 

large clump then develops (Dransfield, 1992a). Within the taxa of African rattans, the 

suckers are produced in the axils of the basal-most leaves and this is commonly the 

case with most clustering species of rattan. However, in a few instances (e.g. some 

species of the Asian genus Daemonorops), the sucker shoots are produced opposite to 

the leaf, whilst the vegetative buds of Korthalsia, another Asian genus, appear to be 

intermediate between axillary and leaf-opposed (Fisher and Dransfield, 1979). 

Although in general, rattan species are consistently single-stemmed or clustered it is 

possible to encounter individuals of predominantly clustering species with only a 

single stem e.g. the Asian species, Calamus subinermis H. Wendl ex Becc. This is 

sometimes the case with some normally clustering Eremospatha species in Africa, 

particularly in closed-canopy forest where the lack of light penetration could possibly 

be a limiting factor to the development of the cluster. 

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Figure 1. Mature clustering stem of Laccosperma robustum in the Korup National Park, 

Cameroon (Sunderland 2305) 

Aerial branching, as opposed to suckering, is commonly encountered in a number of 

palm taxa (Ridley, 1907; Hodge, 1965; Corner, 1966; Dransfield, 1978; Fisher, 1973; 

Fisher, 1974; Fisher and Dransfield, 1979; Dransfield, 1992a; Fisher and Tomlinson, 

1973; Tomlinson, 1961a; Tomlinson, 1990). Although, in general, rattans lack the 

ability to produce vegetative branches above the few basal nodes, there are exceptions 

to this and aerial branches in the Calamoid palms are well reported (Dransfield, 1978; 

Fisher and Dransfield, 1979; Uhl and Dransfield, 1987; Baker et al., 1999d). These 

branches are axillary in origin, and this type of branching is quite different from the 

regular, two-forked, dichotomous branching of Hyphaene and Nypa (Corner, 1966; 

Fisher and Tomlinson, 1973; Dransfield, 1978; Tomlinson, 1990), the non-axillary 

(abaxial) branching of Dypsis (Fisher, 1973, as Chrysalidocarpus), or the production 

of a sequence of inflorescences and vegetative branches (suckers) along a horizontal 

axis as in Serenoa (Fisher and Tomlinson, 1973). 

Within the Calamoideae, aerial branching has been recorded in a number of species of 

Korthalsia (Dransfield, 1978; Fisher and Dransfield, 1979; Dransfield, 1992a). 

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Similar branching of the stem is also relatively common in some species of both 

Laccosperma and Eremospatha. This type of branching is sympodial and axillary, 

with the branch developing from anywhere on the stem. In the case of the African 

rattans, branching does not necessarily occur in association with flowering or from 

damage to the stem apex, as has been suggested it might by a number of palm workers 

(Corner, 1966; Dransfield, 1978) and appears to be relatively widespread, particularly 

within Laccosperma and Eremospatha. In these genera, branching occurs most 

commonly amongst forest-dwelling species rather than for species preferring more 

light-demanding forest habitats. 

Figure 2. Aerial branching of Eremospatha hookeri in the Korup National Park, Cameroon 

(Sunderland 2302) 

The branching habit has a considerable impact on utility, and those species in which 

aerial branching occurs are not commonly valued as a source of cane. The cane quality 

(and length) is deleteriously affected by the development of axillary branches and the 

cane itself tends to break readily. 

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Table 1. African rattan species producing aerial branches 

Species Habitat Branching 

Laccosperma laeve Closed-canopy forest Common 

L. opacum Closed-canopy forest Common 

Eremospatha hookeri Closed-canopy forest Common 

E. quinquecostulata Closed-canopy forest Common 

E. tessmanniana Closed-canopy forest Profuse 

E. wendlandiana. Closed-canopy forest and forest gaps Relatively uncommon 

Suckering from shoots that fall and make contact with the forest floor occurs in 

several species of Calamus (Dransfield, 1978) and this is where adventitious 

vegetative buds develop from axillary meristems (Fisher and Dransfield, 1979). In 

Africa, the widespread Calamus deërratus develops into dense, often monospecific, 

stands through profuse suckering from the base, as well as from these axillary 

meristems. 

1.2.2 The stem 

The rattan stem, or cane, is concealed at first by the tightly sheathing and often 

densely spiny leaf bases. As the stems develop and grow, the leaves mature and die. 

The leaves are not shed neatly but gradually tatter and fall away. It is only when the 

older leaf sheaths have finally sloughed off that the rattan stem is exposed. It is at this 

stage that the cane is mature enough to be harvested. 

The diameter of rattan stems is relatively constant within species and hence it is 

possible to distinguish between “small-diameter” (20 mm) species. For each species, as the rattan seedling develops, there is a gradual 

increase in the diameter of the stem to the maximum prior to the stem beginning to 

develop apically. This is known as the establishment phase (Dransfield, 1992a). The 

diameter of the lowermost portion of the stems, where the roots emerge, may be 

greater than that just above the base, and when the stem reaches the open canopy, the 

diameter may also increase slightly (Fisher, 1978). However, the stem diameter is 

relatively uniform within each species. 

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The growing point of the stem of rattans is found at a point considerably below the 

apparent stem apex, enclosed by the leaf sheaths, the youngest of which form a 

column at the stem apex (Tomlinson, 1990). The developing leaves represent the 

“palm heart” of the rattan which is often consumed by forest dwellers and herbivores 

(see Chapter 6 and Appendix 1). Damage to this delicate growing point often results 

in the death of the individual stem. 

Although the stem length of some rattans have been recorded as reaching up to 175m 

in length (Burkill, 1935), for most African rattan species the average stem length is 

between 30-50m. In this respect, Eremospatha macrocarpa & E. tessmanniana are 

relatively uncommon in that individual stems of these species can attain lengths of up 

to 150m. 

In general, most stems of African rattans are circular in cross-section. However, the 

stems of Eremospatha laurentii, in particular, are broadly triangular in cross section at 

maturity. Stems of this species are difficult to bend and, as such, are not often utilised 

for cane work. 

1.2.3 Cane anatomy 

The intrinsic property of rattan stems most affecting the cane quality is that of the 

anatomy of the cane itself (Tomlinson, 1961b). Detailed studies of the anatomical 

properties of rattan canes have been undertaken by Weiner and Liese (1988; 1989) 

although some regional studies focussing on the Indian canes (Bhat, 1991) and some 

members of the African rattan species (Weiner and Liese, 1994; Oteng-Amoako and 

Ebanyele, in press) have also taken place. These studies suggest that there are 

significant anatomical differences between the majority of the rattan genera and allow 

the determination of desirable properties for the optimum quality of cane. In general, 

Bhat et al. (1988), Bhat and Verghese (1989) suggest that the three most important 

structural features that appear to determine rattan physical properties are fibre wall 

thickness, the relative proportion of fibrous tissue and metaxylem vessel diameter. 

Anatomical studies of three of the four (Laccosperma, Eremospatha and Calamus) 

African genera have recently been undertaken in Ghana (Oteng-Amoako and 

Ebanyele, in press). The initial results of the study suggest that the thickness of the 

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fibre walls, the proportion of fibre tissues and metaxylem vessel diameter differ 

significantly between the genera, hence influencing the relative utility of the members 

of each. 

The relatively higher proportion of thick-walled fibres and narrower diameter of 

metaxylem vessels suggests that the genus Laccosperma has a greater density and 

hence strength properties than the canes of Eremospatha and Calamus deërratus. 

These latter taxa have a relatively higher proportion of thinner wall fibres and larger 

metaxylem vessels, which contribute to greater void volume of the stems resulting in 

lower density and strength (ibid.). The study also revealed that fibre wall thickness, 

fibre proportion and metaxylem diameter, which are the likely determinants of rattan 

quality, do not differ significantly between Calamus and Eremospatha and therefore 

these genera are included within the same density and strength groupings. These 

findings concur with those of Wiener and Liese (1994) who additionally examined 

material of Oncocalamus. This latter genus was found to have very thin fibre walls 

and very large metaxylem vessels and possessed the least desirable properties of 

density and strength of any of the African rattans. 

These anatomical conclusions generally correspond with those of researchers of rattan 

utilisation in Africa. It is generally accepted that the large-diameter species of 

Laccosperma are particularly durable, whilst Oncocalamus is particularly weak and 

brittle and, as such, is not commonly valued as a source of cane (Profizi, 1986; Defo, 

1997; Defo, 1999; Sunderland, 1999a; 1999b). However, it is surprising that 

Eremospatha and Calamus are found to be anatomically similar, and hence share 

similar cane properties, as most workers note that Calamus deërratus is considered of 

inferior quality to that of the desired species of Eremospatha and is only utilised in the 

absence of other species (see Appendix 1). Further anatomical studies, which are 

currently under way, might shed more light on this anomaly. 

1.2.4 The root system 

The primary root in palms functions for a very short time only in the seedling. 

Subsequent roots are lateral and may also be described as secondary or adventitious. 

These secondary roots are usually borne near the base of the stem, forming large root 

masses at or near ground level (Uhl and Dransfield, 1987). 

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Aerial roots are commonly encountered in the Calamoid palms in particular e.g. 

Pigafetta, Daemonorops and Korthalsia (Dransfield pers. comm). The slender 

inflorescences of Calamus pygmaeus Becc., an acaulescent species of rattan, arch out 

between the leaves, and from their apices sprout new vegetative shoots that, on 

contact with the soil, develop into new shoots. A relatively uncommon species of 

Eremospatha in Africa, E. tessmanniana produces conspicuous aerial roots at the 

point of aerial branching. These roots persist until such time as the stem falls to the 

floor and they are able to anchor themselves and allow the shoot to then develop 

(author pers. obs.). 

A recent study of the root anatomy of some Calamoid genera, whilst limited in its 

scope, has identified distinct relationships based on root anatomy within the subfamily 

(Seubert, 1996). Of particular significance is the fact that the genus 

Eremospatha is, in this respect, closely related to the Asian genera Calamus, 

Calospatha, Daemonorops and Ceratolobus. These genera possess velamen cells that 

are unequally thickened, isodiametric endodermis cells, with thicker cell walls 

throughout, with frequent mucilage cells that are invariably surrounded by fibres. 

Further study of the root anatomy of the other African taxa might shed light on the 

wider relationships between Eremospatha and the other African genera, Laccosperma 

and Oncocalamus, and might also provide further insights into their relationships with 

the other members of the Calamoideae. 

1.2.5 Rattan leaves 

Rattan leaves are produced spirally sequentially at the apex of the stem. They consist 

of a tubular leaf sheath which arises directly from the node on the stem (Corner, 

1966). At its apex, the sheath narrows into a petiole that continues into the rachis or 

portion of the leaf bearing leaflets. In some cases the petiole is absent and the leaf is 

sessile. In many species within the Calamoideae, the rachis is extended beyond the 

terminal leaflets to form a barbed whip, or cirrus; the principal means of climbing for 

the species in which it is present. 

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1.2.6 The sheath 

The leaf sheath develops from a soft meristematic region at the base and hence the 

upper portion of the sheath matures before that of the base. In extreme cases the whole 

leaf may be expanded and fully functional whilst the base of the sheath is still soft and 

meristematic but this soft region is supported by the tightly enclosing sheaths of older 

leaves (Corner, 1966; Dransfield, 1992a). In this regard, perhaps only 25-30% of the 

entire length of the sheath is visible beyond the preceding leaf. Although this is not an 

internode in itself, this length is correspondent with the length of the internode of the 

stem. The sheaths of the rattans of Africa, in common with many of those of SE Asia, 

are tubular and tightly sheathing. 

1.2.7 Sheath armature 

The degree of armature on the leaf sheath is highly diagnostic, and most rattan species 

bear spines on the sheath. Spines are emergences which vary in their stoutness and 

persistence (Tomlinson, 1962a). The presence of spines on the sheath is a common 

feature of the Calamoideae and the sub-family shows an extraordinary diversity in 

spine arrangement. This arrangement is often of diagnostic importance and Dransfield 

(1992a), suggests that armature may be of adaptive significance. The presence of 

spines and armature is commonly assumed to be linked to the climbing process, as 

well as protection against herbivory (Tomlinson, 1962a; Corner, 1966). 

In the African Calamoid palms, spines are not of a characteristic arrangement (e.g. in 

whorls or in vertical series) as they are in other Calamoid palms, particularly some 

members of Daemonorops and Calamus. The spines of Oncocalamus and Calamus 

are broadly triangular, often dark brown in appearance and are particularly 

concentrated on the upper portion of the sheath with considerably lesser armature 

evident on the remainder of the sheath. The spines on the leaf sheath of the genus 

Laccosperma are fine and needle-like and more commonly, dark green or black; these 

spines are more uniformly distributed over the length of the sheath as well as on the 

ocrea itself. 

Spines also vary in persistence. Those of the genera Laccosperma and Calamus are 

rather persistent as they are on a number of species of Oncocalamus and, as such, do 

not slough off readily. However, the genus Oncocalamus is characterised by the 

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sloughing of the sheath spines particularly on drying, leaving bare sheaths. However, 

these sloughed spines may leave faint, yet distinct, scars. 

The total absence of spines on the sheath of Eremospatha is a characteristic feature of 

this genus. 

1.2.8 Indumentum 

Between the spines on the sheath (where they occur) a thin covering of scales or 

indumentum is a characteristic feature of nearly all of the endemic rattan genera in 

Africa, particularly on the young or developing stems. In both Laccosperma and 

Eremospatha, this indumentum is often dark brown to black and non-waxy, and in 

Oncocalamus, the indumentum is white and somewhat waxy. Calamus deërratus does 

not possess a distinctive indumentum. 

1.2.9 The ocrea 

At the mouth of the sheath where it narrows into the petiole or leaf rachis, there is 

often a prolongation of the main part of the sheath. Often this prolongation is tubular 

and encloses the sheath of the leaf above. This is called an ocrea. The ocrea may be 

long, papery and prone to tattering as in Laccosperma, or may be short, saddle-shaped 

and more truncate, as in Oncocalamus and Eremospatha. Calamus deërratus 

possesses an ocrea that is intermediate between these two. The ocrea of two species of 

Eremospatha, E. wendlandiana and E. barendii, are conspicuously longitudinally split 

on the opposite side to the leaf. The resulting cleft is also rather prone to drying. 

A number of species of Eremospatha, (E. macrocarpa, E. wendlandiana and E. 

hookeri) also possess a distinctive linear “wrinkle” or narrow ridge, on the ocrea 

facing the leaf. This wrinkle is particularly evident in the juvenile growth stages. This 

morphological feature is not encountered elsewhere in the Calamoideae and is unique 

to the African rattan taxa. 

1.2.10 Sheath striations 

Striations on the leaf sheath are a common feature of Eremospatha, Laccosperma and 

are also possessed by some members of Oncocalamus. The leaf sheaths of Calamus 

deërratus are not striate. 

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1.2.11 The knee 

A knee-like swelling occurs on the leaf sheath below the insertion of the petiole in 

some rattans. In technical terms, this is called a geniculus (Baker et al., 1999d). The 

function of the knee is not known (Uhl and Dransfield, 1987). The knee develops as 

the leaf expands and matures, and may be associated with the change in the angle 

from vertical as it emerges from the apex to the horizontal (Dransfield, 1992a). The 

knee is clearly not associated with the climbing habit, as many climbing palms do not 

possess a knee of any kind. 

The knee is present in some species of the genus Eremospatha and, where it occurs, is 

unique amongst the Calamoideae in that it is conspicuously vertically-linear. Calamus 

deërratus exhibits a horizontally-wrinkled ridge in common with most Asian members 

of the genus. Species within Laccosperma and Oncocalamus do not posses a knee 

although within the latter genus, O. tuleyi possesses a slight horizontal swelling 

beneath the leaf junction. 

1.2.12 The petiole and leaf rachis 

The petiole in most rattan palms is somewhat variable in length (Dransfield, 1992a). It 

is usually much longer in juvenile individuals than in mature leaves and it may be 

absent altogether. The petiole, where present, and the rachis are often heavily armed 

with spines and it has been speculated are not necessarily adapted for the climbing 

process but appear to function as a means of trapping litter from the forest canopy 

(Corner, 1966; Dransfield, 1992a). In taxa that lack petioles, the lowermost leaflets are 

also suggested to function as litter traps. The primary purpose of such trapping is 

assumed to be associated with the accumulation of nutrients (ibid.) 

In the genus Laccosperma, some species (particularly L. secundiflorum) are 

characterised by a long petiole, whilst other species have a very short petiole (L. 

robustum and L. acutiflorum). In the species of Oncocalamus, the petiole is absent or 

much reduced, particularly in mature stems, although more conspicuous in the 

juvenile stage. 

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For many of the species of Eremospatha, the petiole is absent and the lowermost 

leaflets are modified such that they are much reduced and strongly reflexed so that 

they are swept back and clasp the stem. This clasping can be rather lax, or extremely 

tight, enclosing the stem completely. This enclosure is often colonised by ants (see 

Chapter 4). 

1.2.13 Leaflets 

The rattan leaf is pinnate (Uhl and Dransfield, 1987; Dransfield, 1992a). The leaflets 

result from the splitting of the folded leaf blade which occurs as the leaf expands. In 

almost all species of rattan, the leaflets consist of a single-fold; however, within some 

species of African rattan (L. secundiflorum and Oncocalamus wrightianus), leaflets 

can be composed of two or more folds. 

Although, in general, the leaflets within the Calamoideae are entire, some taxa possess 

jagged, distal margins; these are termed praemorse. This character is particularly 

common within the genus Eremospatha. 

The leaflets of the majority of species of African rattan are linear. However, some 

species of Laccosperma, in particular, are sigmoid. Other leaflet shapes exhibited by 

the African rattans include the distinctive rhomboid leaflets of Eremospatha 

wendlandiana and the more rounded rhomboid leaflets of E. cabrae and E. hookeri. 

The manner in which the leaflets are held on the rachis is often highly diagnostic. In 

particular, the leaflets of Laccosperma robustum are conspicuously pendulous 

differentiating this species from other, closely related species of the same genus. The 

leaflets of Eremospatha macrocarpa are also somewhat pendulous distinguishing this 

species from other species within the genus. In addition, although the majority of the 

African rattan taxa possess leaflets that are arranged uniformly and singly on the 

rachis, the leaflets of Calamus deërratus may be grouped, particularly on the distal 

portion of the leaf. The slender forest species, Eremospatha quinquecostulata has 

characteristically clustered (or plumose) leaflets, arranged in groups of 4-6. 

The presence of spines on the leaflet margins is a common feature of the African 

rattans and, where present, their size and arrangement is a good field characteristic for 

31

distinguishing species. Laccosperma laeve, for example is devoid of leaflet spines, 

and character is a useful means of distinguishing this species from the closely-related 

L. opacum. Accordingly, the long, fine, hair-like leaflet margin spines of L. robustum 

are a useful means of distinguishing this species from L. secundiflorum and L. 

acutiflorum, which both possess rather short and robust spines on the leaflet margin. 

In the majority of the African rattan taxa, the emerging leaf is a mid-deep green. 

However, Oncocalamus is often characterised by the presence of an orange or crimson 

expanding leaf, this turning deep green as it develops. 

1.2.14 Climbing organs 

Calamoid palms climb with the aid of two main organs; they may either have a 

flagellum or posses a cirrus. Flagella only occur in certain species of Calamus, 

including C. deërratus, the sole representative of Calamus in Africa. The flagellum 

arises directly from the sheath and is regarded as a modified inflorescence (Fisher and 

Dransfield, 1977; Dransfield, 1978; Baker et al., 1999d). Indeed, inflorescences of C. 

deërratus are flagellate. 

The remaining taxa within the Calamoideae, particularly those of Asian origin, climb 

with the aid of a cirrus, a whip-like extension at the distal end of the leaf rachis armed 

with short, recurved thorns that often resemble a cat’s claw (Tomlinson, 1990). 

However, the three rattan genera endemic to Africa, Laccosperma, Eremospatha and 

Oncocalamus, possess a vegetative morphology unique within the Calamoideae and 

the cirrus is actually a marked extension between the distal leaflets. The leaflets are 

present as reduced, reflexed thorn-like organs termed acanthophylls. This structure is 

also present in some members of the unrelated genera present only in the New World; 

Chamaedorea (sub-family Ceroxyloxideae; tribe Hyophorbeae) and Desmoncus (subfamily 

Arecoideae; tribe Cocoeae). These taxa also climb through the means of 

reflexed terminal leaflets. Coincidentally, many species of Desmoncus are also 

exploited for their cane-like qualities and are used in the same way as the true rattans 

(Henderson and Chávez, 1993). 

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Figure 3. Acanthophylls of Laccosperma acutiflorum, Campo, Cameroon (Sunderland 1926) 

On the lower portion of the stems of some taxa in the genera Eremospatha and 

Oncocalamus, an undifferentiated rachis is often produced, devoid of any true leaflets 

and possessing acanthophylls only. This unusual, and previously unrecorded, organ 

seems to be more common on younger stems, particularly under a forest canopy. 

However, this organ also occurs in Oncocalamus colonising recently cleared areas as 

well. The adaptive significance of this organ is not known, although it certainly 

facilitates the establishment of the developing stems and assists in the climbing 

process. In the absence of an existing term for this appendage, I have applied the term 

“elaminate” (devoid of leaf blades) to signify the lack of true leaflets on the rachis. 

1.2.15 Flowering behaviour 

In all, 16 genera of palms, the majority of them in the Calamoideae, produce what 

appears to be a massive “terminal” inflorescence that results in the death of the 

primary axis of the palm. In fact, this structure is not terminal (Corner, 1966) but is an 

aggregate inflorescence that consists of a large number of lateral inflorescence units 

borne in the axils of the leaves, which are often reduced. These inflorescence units are 

borne in the distal portion of the stem and mature simultaneously (Tuley, 1965; Uhl 

33

and Dransfield, 1987, Tomlinson, 1990). This condition is termed hapaxanthy. In 

essence, there is no morphological difference between the inflorescences in 

hapaxanthy and pleonanthy, its opposite condition. Of the three endemic African 

rattan genera, Laccosperma is hapaxanthic, whilst Eremospatha and Oncocalamus are 

pleonanthic. Calamus deërratus is also pleonanthic. See Chapter 5 for further 

elaboration of hapaxanthy and pleonanthy in the African rattans. 

Figure 4. Elaminate rachis of Oncocalamus tuleyi, Southern Bakundu FR, Cameroon (Sunderland 

1761) 

1.2.16 The inflorescence and floral morphology 

Fisher and Dransfield (1977) showed that adnation of the inflorescence to the 

internode or the leaf sheath occurs in several rattan genera. In the Calaminae, the 

inflorescence is adnate to the internode and the sheath above its axil of origin such 

that the inflorescence appears to arise from the sheath itself (Baker et al., 1999d). The 

flagellum of Calamus deërratus, being a modified inflorescence, is attached in this 

way. The other rattan genera in Africa do not display any inflorescence adnation 

(although fide Baker et al., 1999d). 

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In common with other members of the genus, Calamus deërratus possesses dyads of 

unisexual flowers, however the inflorescence units of the endemic rattan genera of 

Africa also somewhat unique within the Calamoideae. The genera Eremospatha and 

Laccosperma are composed of dyads of hermaphroditic flowers 1 . Although the dyad 

composed of unisexual flowers is itself a common feature within the Calamoideae, the 

dyad composed of hermaphroditic flowers is unique within the Palmae and is 

considered to be an unspecialised form of flower arrangement (Uhl and Dransfield, 

1987; Baker et al., 1999b; 1999d). 

In contrast, the flower cluster of Oncocalamus is distinctive and complex, not only 

within the Calamoideae, but in the Palmae as a whole. Oncocalamus is monoecious, 

and consists of a central 1-3 pistillate flowers with two lateral cincinni subtended by a 

single bract, with each cincinnus bearing basal 1-3 pistillate flowers and 3-5 distal 

staminate flowers. 

Figure 5. Diagram of selected dyads within the Calamoideae showing arranegments of flowers in 

hermaphroditic, monoecious and dioecious taxa. 

Laccosperma Oncocalamus Calamus 

Eremspatha (Monoecious) (Dioecious) 

(Hermaphroditic) 

With the exception of the genus Eremospatha which, uniquely in the Palmae possess 

an inflorescence that is free of conspicuous bracts, the Calamoideae possess strictly 

tubular bracts that are rather uniform in shape (Baker et al., 1999d). As such, within 

the genera Laccosperma and Oncocalamus, as well as Calamus deërratus, the rachis 

bracts are tightly sheathing. In the former two genera, these bracts are often prone to 

splitting and tattering post-anthesis. 

1 Less commonly, Laccosperma may also possess triads of flowers. 

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1.2.17 Calyx and corolla 

The calyx of male and female flowers of C. deërratus is distinctly tubular, enclosed 

the flower up to ¾ of its length. The calyx of the endemic African genera is less 

tubular and more cup-shaped, or campanulate, often enclosing the corolla for 1/3, or 

more, of its length. In both Laccosperma as well as both the male and female flowers 

of Calamus deërratus, the corolla is fused for only the proxal 1mm with the remainder 

of the corolla divided into 3 corolla lobes that are moderately (Laccosperma) to highly 

(Calamus) divergent at anthesis. In Eremospatha the corolla is fused for ½ to ¾ of its 

length with the corolla lobes forming a minute (trilete) opening at anthesis. The 

corolla of Oncocalamus is intermediate between the other two African genera and is 

commonly united only for ½ of its length. 

1.2.18 Stamens 

The stamens of the endemic African genera are all united to varying degrees. In 

Laccosperma, the filaments are united at the very base only to form a short staminal 

ring. In Eremospatha, the filaments are united into a massive, fleshy epipetalous ring 

and in the staminate flowers of Oncocalamus, the filaments are also united to form a 

thick, fleshy androecial tube that is free from the corolla. The filaments of the 

staminate flowers of Calamus deërratus, however, are free. 

1.2.19 Pollen 

The pollen morphology of the Palmae has been summarised by Ferguson (1986) and 

Uhl and Dransfield (1987). For the African rattan genera the pollen morphology has 

been described in detail by Harley and Hall (1991) and more recently by Harley 

(1996). 

1.2.20 The fruit 

The members of the Calamoideae are characterised by the possession of distinctive 

imbricate scales on the ovary and the fruit, arranged in vertical rows. The number of 

these vertical rows is often used as a diagnostic character to distinguish between rattan 

species in particular (see Dransfield, 1979; 1984; 1992c). However, within the African 

rattans, the number of vertical rows of scales ranges from between 15-22 for all 

species and hence it is not a sound distinguishing character between species. The only 

36

exception to this is Laccosperma opacum, which possesses between 12-14 rows of 

vertical scales. 

Within the outer covering of scales lies the remainder of the fruit wall. In the majority 

of the Calamoideae the fruit wall is rather thin (Uhl and Dransfield, 1987) but in the 

endemic African genera, there is a conspicuous mesocarp (this character is also shared 

by the genus Korthalsia (ibid.)), which is usually somewhat sweet-tasting. The 

innermost layer of the fruit wall, the pericarp, usually consists of a thin membrane 

within which lies the seed. 

A sarcotesta is a fleshy layer developed from the outer seed coat. This fleshy layer is 

common within the Asian members of the Calamoideae, notably among species where 

the fruit wall is thin and not particularly fleshy. Although rare within the African 

rattan taxa, the sarcotesta is particularly conspicuous in Calamus deërratus. Despite 

the presence of a fleshy mesocarp the seeds of the species of Oncocalamus also posses 

a very thin sarcotesta, as does Laccosperma opacum. 

1.2.21 The seed 

Usually within the Calamoideae, each fruit produces only one seed. However, some 

species of Laccosperma and Eremospatha sometimes have two, or perhaps three, 

seeds in each fruit, although this is rather uncommon. The species of both 

Laccosperma (with the exception of L. opacum, which is globose and has a deep cleft 

on one side) and Eremospatha have rounded seeds that are flattened on one side. The 

seeds of Oncocalamus are always globose. Those of Calamus deërratus are globose 

but possess a small conspicuous beak at the apex. 

The seed coat in a number of genera of African rattans shows some remarkable and 

wide variation. In the majority of the genus Laccosperma, the seed is smooth; 

however, Laccosperma opacum has a distinctly pitted and warty surface. Likewise, in 

Oncocalamus, the seed of O. mannii is also pitted, whilst the seed coat of the 

remaining species is smooth. The seeds of Eremospatha and Calamus are always 

smooth. 

37

The endosperm of most Calamoid palms is solid and uninterrupted. However, the 

endosperm of Oncocalamus is penetrated by a smooth-margined mass of inner seed 

coat, giving the endosperm a peculiar moon-like appearance. The unrelated genus 

Pritchardiopsis (sub-family Coryphoideae; tribe Corypheae) also shares this character 

(Uhl and Dransfield, 1987). All members of the Calamoideae, including those from 

the African continent, possess a homogenous endosperm (ibid.) 

1.2.22 Germination 

All African rattan taxa exhibit adjacent-ligular germination and possess a bifid 

eophyll. 

Figure 6. Adjacent ligular germination of Oncocalamus mannii, Limbe Botanic Garden, 

Cameroon 

1.3 FOSSIL RECORD AND BIOGEOGRAPHY 

The paucity of the palm flora in Africa has been long recorded (Richards, 1973; 

Moore, 1973; Brenan, 1978; Moore, 1982; Dransfield, 1988b; Lawson, 1996; Maley, 

1996). This lack of palm diversity along with the lack of diversity in other groups such 

the Orchidaceae, as well within the families Lauraceae, Myrtaceae and the 

Myristicaceae has led the continent being referred to as the “odd man odd”, in terms 

38

of overall diversity, when compared to Madagascar, SE Asia and the Neotropics 

(Richards, 1973). On the African continent, only fifteen palm genera are known, seven 

of which are endemic, representing ca. 72 species (Dransfield, 1988b). When 

compared with the astonishingly rich palm flora of Madagascar (170 species in 16 

genera, with 165 endemic species (Dransfield and Beentje, 1995)) this lack of 

diversity is somewhat surprising. In addition to the paucity of genera and species, the 

morphological disjunction between these taxa is great and they are not particularly 

closely related (Moore, 1973; Moore & Uhl 1982; Dransfield, 1988b). Even within 

closely related taxa such as the endemic rattan genera, the wide differences in floral 

arrangement and morphology are somewhat surprising 2 . In this respect, the palm flora 

of Africa is considered to be the living relic of a previously much richer palm flora. 

Indeed, as Dransfield (1988b) suggests, although the fossil record of palms in Africa 

is somewhat sparse, sufficient pollen records 3 indicate a much richer palm flora in the 

past (Morley and Richards, 1993). 

In particular, there is evidence of a sudden increase in the presence and diversity of 

palm pollen in the Lower Senonian and Maastrichtian periods (Dransfield, 1988b; 

Maley, 1996), suggesting that palms have had a considerable presence in Africa. The 

pollen record also reveals the presence of a number of additional pollen types that are 

known to be of extinct Palmae (Herngreen and Chlonova, 1981; Morley and Richards, 

1993; Maley, 1996) among which were forms close to the present day genus Nypa. 

These extinctions are now known to have taken place within the Oligocene and the 

Miocene, and whilst palms declined considerably in Africa during this time, they 

persisted in large numbers to the present day in the Neotropics and SE Asia. These 

main extinction stages correlate with the main phases of climatic deterioration on a 

global level which also affected the other tropical regions, yet is has been postulated 

that the arid phases that occurred in Africa were much more severe than they were on 

the other continents (Maley, 1996). 

2 As these genera share similar vegetative morphology but are distinct in reproductive structure, this 

suggests that a complex evolution of the Calamoideae alone has occurred in Africa with much 

extinction leaving only these isolated lineages. 

3 The pollen of the Palmae are useful indicators of changes in species composition over time, not only 

of the family itself, but also for the wider forest habitat (Maley, 1996), especially as they produce large 

quantities of fairly resistant pollen (Moore, 1973). 

39

Maley (1996) argues that the extant forest flora of Africa are related to the forest taxa 

of the Eocene, Oligocene and the Miocene such that much of the present day flora of 

tropical Africa must have persisted through the climatic vicissitudes of the 

Quaternary. In order for this to take place, refugia must have existed where the 

climatic conditions remained stable for such taxa, dependent on high rainfall and a 

very short dry season, to occur. Based on floristic evidence, in terms of both diversity 

and endemism, such forest refugia have been long postulated for Africa (Brenan, 

1978; Hamilton, 1981; Pannel and White, 1988; White, 1993; Maley, 1996; 

Robbrecht, 1996; Sosef, 1996; Morat and Lowry, 1997). The speciation patterns, 

endemism and diversity of the African rattans corresponds directly with these refugia. 

The major centre of diversity and endemism for the African rattans is the Guineo- 

Congolian refuge ranging from SE Nigeria to Gabon (17 species), with secondary 

refugia in the Upper Guinea forests of Côte d’Ivoire and Liberia (6 species) and the 

Kivu region of central DR Congo (9 species). Aside from the Guineo-Congolian 

refuge, the levels of endemicity within each refugium are relatively low and there is 

considerable overlap in species composition between each. Interestingly, 

Oncocalamus wrightianus is alone in the fact that it is the sole representative of the 

African rattans present in the eastern Delta region of eastern Nigeria, another 

postulated refuge. 

Figure 7. Pleistocene refugia and the distribution of African rattans (the spotted areas represent 

the major refugia) 

6 species 

(0 endemic spp) 

O. wrightianus 

18 species 

(5 endemic spp) 

40 

9 species 

(2 endemic spp)

CHAPTER TWO 

TAXONOMIC ACCOUNT 

“Taxonomies are not relevant abstractions but are the essential foundations of 

conservation practice” 

2.1. INTRODUCTION 

- Daugherty et al., 1990. 

Despite the economic importance of the rattans of Africa, until recently, the taxonomy 

of this group has been uncertain. Based on recent fieldwork as well as thorough 

examination of herbarium records, this taxonomic account describes 20 species within 

the four genera, including two new species. A neotype is assigned for Laccosperma 

robustum. 

2.2. HISTORY OF CLASSIFICATION 

The first rattan collections from the African continent were from specimens collected 

by the French botanist Baron Palisot de Beauvois. De Beauvois, was appointed in the 

capacity of “gardener” to the mission of Landolphe (1786-88), the aim of which was 

to establish a French trading station at what is, today, Owerri in Nigeria. Although in 

this capacity he was supposedly responsible for the establishment of plantations in the 

area, he instead spent most of his time collecting herbarium specimens. Between 

1786-87, de Beauvois diligently explored Owerri and the region around Benin, 

travelling as far east as Calabar, collecting as he went. Despite considerable hardship 1 , 

de Beauvois’ collections provided many taxa new to science and culminated in the 

publication of his nineteen-volume Flore d’Oware et de Benin en Afrique (1805- 

1820). The publication of the first volume in 1805 included the first description of an 

African rattan species 2 . Although morphologically distinct from its Asian relatives 

particularly in its floral arrangement, de Beauvois included the taxon within the genus 

1 During the duration of the mission, of the 300-strong French contingent, over 250 died within the first 

twelve months (Cuvier, 1928). In 1791, despite being under the safe keeping of Landolphe, the majority 

of de Beauvois’ specimens were destroyed when the British plundered Owerri in 1791. The specimens 

that survive from the voyage are those that were periodically despatched by de Beauvois to Jussieu in 

Paris (Merrill, 1936). 

41

Calamus. Hence, de Beauvois has the distinction of providing the first description of a 

species of African rattan: Calamus secundiflorus. 

Along with other botanical specimens, further collections of rattan, particularly from 

the Niger Delta by Barter (1857-1859) and later from the major estuarine areas and 

coastal forests of Sierra Leone, Nigeria, Cameroon and Gabon by Mann (1859-1863) 

were made (Keay, 1954). These collections provided the basis for the first 

comprehensive treatment of the palms of Africa by Mann and Wendland in 1864. 

Their paper provided the first published treatise of these new discoveries and 

described eight species of rattan, seven of which were described for the first time. In 

recognition of the morphological uniqueness of the African rattan taxa, Mann and 

Wendland described four main rattan sub-genera of the genus Calamus, under which, 

aside from C. deërratus G. Mann & H. Wendl. (or “true” Calamus as they termed it) 

to which all of the remaining African rattan taxa were then consigned and the 

subgenera, Ancistrophyllum, Laccosperma, Oncocalamus and Eremospatha were 

described for the first time. 

The first published recognition of the rattans endemic to Africa to generic rank was by 

Drude (1877) who published the first valid generic use of Laccosperma in 1877. This 

was immediately followed by Wendland who, in Kerchove’s Les Palmiers (1878), 

also elevated the other sub-generic taxa he and Mann described in 1864, Eremospatha 

and Oncocalamus, to generic rank. It is important to note that in this publication 

Wendland continued to recognise Laccosperma and Ancistrophyllum as distinct taxa. 

Bentham and Hooker, in their Genera Plantarum (1883) provided further descriptions 

of the three African genera, but included Laccosperma as a subgenus of 

Ancistrophyllum. The use of this latter name persisted until Dransfield (1982), in 

recognition of Drude’s earlier (1877) publication and with reference to Kuntze’s 

(1891) subsequent observations regarding Laccosperma & Ancistrophyllum 3 gave 

priority to the name Laccosperma, with Ancistrophyllum being reduced to synonymy 4 . 

2 Although there is some confusion regarding the publication of the first volume, the evidence given by 

Merrill (1936) is enough to suggest that 1805 is the correct year of publication and not 1804 as 

published on the frontispiece of Volume 1. 

3 See Dransfield, 1982. 

42

It is somewhat surpising that since Mann and Wendland’s (1864) account, the 

majority of studies of the African rattans have been floristic rather than monographic 

(Drude, 1895; Wright, 1902; Guinea-Lopez, 1947; Renier, 1948; Robyns and 

Tournay, 1955; Irvine, 1961; Russell, 1968; Letouzey, 1978; Dransfield, 1986; 

Berhaut, 1988; Morakinyo, 1995b; Tuley, 1995) or have focussed on the economic 

contribution of the species of rattan to the forest economy (Pynaert, 1911; Mildbraed, 

1913; de Wildeman, 1919; Hédin, 1929; Chevalier, 1936; Dalziel, 1937; Raponda- 

Walker and Sillans, 1961; Profizi, 1986). However, a number of botanists described 

new species of rattan collected from the lowland tropical forests of Africa as the 

botanical exploration of the continent continued (Drude, 1895; de Wildeman, 1904; 

Beccari, 1908; Beccari, 1910; de Wildeman, 1916; Burret, 1942). It was, Beccari, 

however, the Florentine palm specialist who was the first to attempt any monographic 

treatise of the African rattans. His 1908 monographic account of the genus Calamus, 

included the descriptions of five new species from Africa. More specifically, the later 

(1910) monograph of the African Lepidocaryeae provided the first keys to 

identification of the endemic African rattan genera and included full descriptions of all 

the species known at that time. Despite the considerable paucity of material at his 

disposal, Beccari’s 1910 account provides a useful framework for the classification of 

the African rattans. In this regard, it is surprising that his taxonomic treatise of the 

African rattans was not adopted by subsequent workers 5 . 

Since Beccari’s 1908 & 1910 publications, very little taxonomic work has been 

undertaken on the rattans of Africa aside from the floristic studies cited above and 

until recently, there was considerable taxonomic confusion associated with these taxa. 

In particular, the lack of representative fertile material and adequate field observations 

has thwarted any attempt at providing a definitive monograph of this group. The most 

recent attempt by Tuley (1995) at providing a preliminary and somewhat popular 

account of the taxonomy of the palms of Africa, including the rattans, was certainly 

affected by these constraints. 

4 As Dransfield (1982) points out, the use of Ancistrophyllum for a rattan palm is invalid anyhow as the 

name was earlier published by Göppert in 1841 as a fossil Lepidodendron stem. 

5 Indeed, Beccari’s account of Laccosperma ( syn. Ancistrophyllum), in particular, was dismissed 

entirely by Russell (1968) despite considerable morphological evidence and field observations 

supporting Beccari’s taxonomy. 

43

The present treatment recognises twenty species within the four genera (two of which 

are described for the first time), two imperfectly known species, 32 synonyms and 6 

nomina nuda, invalid or unpublished names as such. 

2.3. THE WIDER TAXONOMIC PICTURE 

The Palmae is split into six sub-families (Dransfield and Uhl, 1986; Uhl and 

Dransfield, 1987). The second-largest sub-family, the Calamoideae, is easily 

recognised by the presence of a scaly pericarp. Worldwide, there are estimated to be 

650 species within the Calamoideae belonging to twenty-two genera (Uhl and 

Dransfield, 1987). The Calamoideae is almost exclusively an Old World tribe (Moore, 

1973) and the majority of the Calamoid taxa occur in the humid tropical forests of 

south and SE Asia, Malesia, and the west Pacific. The sub-family is further split into 

two tribes, the Calameae (which includes the Old World members of the tribe) and the 

Lepidocaryeae (which are represented by the New World palms). The Calameae is 

further separated into eight sub-tribes (Uhl and Dransfield, 1987). 

In Africa, the Calamoideae is represented by five genera: Calamus, Eremospatha, 

Laccosperma, Oncocalamus and Raphia (Moore & Uhl, 1982; Uhl and Dransfield, 

1987). These genera currently represent four sub-tribes within the Calamoideae 

(ibid.); the genus Calamus falls within the Calamineae, Raphia within the Raphiinae, 

the genera Laccosperma and Eremospatha within the Ancistrophyllinae and 

Oncocalamus within the Oncocalaminae (ibid.). 

Moore & Uhl (1982), followed by Uhl and Dransfield (1987) placed the 

Ancistrophyllinae and the Oncocalaminae at almost opposite ends of the tribe 

Calameae on the basis of their divergent floral characters, despite their close 

vegetative similarities, as discussed above. However, recent cladistic analysis has 

determined that, despite the variation in the reproductive structure, the two sub-tribes 

are much more closely related than previously thought (Baker et al., 1999a; 1999c; 

1999d). In addition, the relationship between the African rattans and the other 

climbing palms is not at all close (Baker et al., 1999b). This is unsurprising, given the 

biogeographical and morphological differences between the two groups but this 

44

evidence suggests that the climbing habit has evolved within the Calamoideae on at 

least two occasions (Baker et al., 1999b; Lewis et al., 2000). 

The major relationships within the Calamoideae highlighted by Baker et al. (1999a; 

1999c; 1999d) are distinctly influenced by biogeographical considerations. As such, 

strong relationships are known occur between the Ancistrophyllinae and the 

Oncocalaminae 6 , the predominantly African Raphiinae and the Neotropical 

Lepidocaryeae. Indeed these form distinct clade of African-American palms within 

the Calamoideae. 

Figure 8. Schematic tree showing the relationships within the Calamoideae (Baker et al., in press) 

Key 

E Eugeissona 

AR African rattans 

R Raphia 

L Lepidocaryeae 

ES Eleiodoxa-Salacca 

P Pigafetta 

M Metroxylinae 

Pl Plectocomiinae 

CR Rattans of the Calaminae 

45 

OUTGROUP 

6 A revision of Genera Palmarum is currently in preparation. In light of the research cited, this revision 

may place the Ancistrophyllinae and the Oncocalaminae much closer together within the Calameae, or 

may include them in a single sub-tribe (Dransfield, pers. comm.). 

E 

AR 

L 

ES 

M 

R 

P 

Pl 

CR 

The African- 

American Clade The Asian Clade

Interestingly, the position of the sole representative of Calamus in Africa, C. 

deërratus, is as yet unresolved. Baker et al., (1999b), in their study of the molecular 

phylogenetics of Calamus found that this taxon, being flagellate, did not, as would be 

expected, resolve in the same clade as the other flagellate members of the genus and 

seems exclusive to this group. This would indicate that the presence of this taxon in 

Africa is due to vicariance, rather than dispersal as proposed by Dransfield (1988), 

and adds credence to the hypothesis that the Calamoideae are specifically Gondwanan 

in origin (Moore & Uhl, 1982; Dransfield, 1988b; Baker and Dransfield, 2000). 

2.4. SPECIES CONCEPTS 

A review of the plethora of literature on the philosophy and nature of species concepts 

is beyond the scope of this study. However, the importance of explicitly stating the 

concepts and methods used for the recognition of taxa in the production of botanical 

monographs has been highlighted by Luckow (1995) and McDade (1995), and further 

elaborated by Sidwell (1999). The implicit taxonomic convention that the personal 

opinion of an expert botanist is sufficient to delimit taxa is, rightly, unacceptable for 

an unbiased scientific study, and it is essential that revisionists explicitly determine 

the concepts used in monographic study. In palms, the most commonly applied 

species concept that is applied to palm taxonomy is the morphological species concept 

where discontinuities in morphological variation provide the means to separate species 

(Davis and Haywood, 1963). The morphological species concept in palms has recently 

been reviewed for the Old World by Dransfield (1999) and for the Neotropics by 

Henderson (1999). Both studies suggest that the delimitation of species using the 

morphological species concept is valid. 

The species concept applied in this revision is based on morphological discontinuities 

both within and between populations and I have recognised as species only those 

smallest units that can be diagnosed by constant character states. 

46

2.5. KEY TO THE GENERA 

Rattans climbing by means of a cirrus armed with acanthophylls; 

Leaf sheath without spines; stem sometimes with conspicuous knee below leaf 

junction; lowermost leaflets often swept back across stem; leaflets variouslyshaped, 

often with praemorse apices, or leaflets entire. Inflorescence without 

conspicuous bracts: Eremospatha 

Leaf sheath armed with conspicuous spines: 

Ocrea triangular, not tubular drying; spines on sheath long, slender and 

sparsely to densely arranged, not easily detached. Inflorescence units 

produced simultaneously in the axils of the distal leaves (hapaxanthic). 

Hermarphroditic flowers in dyads, rarely triads: Laccosperma 

Ocrea neat, horizontal, not dry; spines on sheath short, irregularlyspaced, 

black or brown, triangular, easily sloughing off. Inflorescence 

units, pendulous, produced in axils (pleonanthic). Monoecious flowers 

in clusters of 7-11: Oncocalamus 

Rattans climbing by means of a flagellum emerging from the leaf sheath: Calamus 

47

EREMOSPATHA (G. Mann & H. Wendl.) H. Wendl. 

Eremo = (Latin) “destitute of”; spatha = (Latin) “spathes” 

H. Wendl. in Kerchove, Les Palmiers 244: (1878). Lectotype: E. hookeri (G. Mann 

& H. Wendl.) H. Wendl. (Calamus hookeri) (G. Mann & H. Wendl.) see H.E Moore 

1963). 

Calamus subgenus Eremospatha G. Mann & H. Wendl. in Trans. Linn. Soc. 

(London), 24: 433 (1864). 

Solitary or clustered, spiny, high-climbing, pleonanthic, hermaphroditic rattan palms. 

Stems circular to triangular in cross section, with short to long internodes, juvenile 

stems much more slender than the adult, sucker shoots axillary. Leaves pinnate, bifid 

in juveniles, becoming pinnate, with a terminal cirrus; sheath strictly tubular, 

unarmed, longitudinally striate, often sparsely to moderately covered with black or 

brown caducous indumentum or indumentum absent; ocrea conspicuous, entire, 

tightly sheathing, neatly or obliquely truncate or somewhat saddle-shaped, or drying 

and becoming longitudinally split; knee absent or conspicuous, vertically linear, 

abrupt or tapering at base, sometimes more linear, ridge-like; elaminate rachis present 

on lower stems, or absent; petiole present in juvenile stems, occasionally in mature 

climbing stems, armed with reflexed spines, sometimes with a caducous indumentum 

below, or indumentum absent; rachis armed as the petiole, with caducous indumentum 

below; cirrus armed as the petiole or unarmed; leaflets few to numerous, single-fold 

except, rarely, in juvenile leaves where lamina undivided, clustered or regularly 

arranged, linear-lanceolate, sub-orbicular to rhomboid, broadly attenuate at the base, 

narrowly to broadly praemorse or entire and apiculate at apex, concolorous or 

somewhat discolorous, usually armed along the margins with conspicuous robust 

reflexed spines; tranverse veinlets moderately to highly conspicuous; proximal few 

leaflets on each side of the rachis often smaller than the rest, shaped as the mature 

leaflets or somewhat linear, strap-like, sparsely to heavily armed along margins, laxly 

or tightly reflexed across the sheathed stem, or not reflexed and more regularly 

arranged; acanthophylls in neat pairs, opposite, rarely sub-opposite, parallel, or at 

varying angles to cirrus. Inflorescence arching outward, branched to 1-order, branches 

48

horizontal, peduncle enclosed within the leaf sheath and emerging from the mouth, 

flattened, not adnate to the internode, the surface glabrous or minutely to profusely 

papillose; bracts throughout the inflorescence inconspicuous or somewhat more 

conspicuous; prophyll absent although vestigial scar visible; peduncular bracts absent; 

rachis somewhat longer or than the peduncle; rachis bracts low, triangular, striate, 

opposite, alternate distally, sometimes united proximally to form an incomplete 

sheathing collar; rachillae distichous, opposite proximally and subtended by a double 

bract, becoming alternate distally, subtended by a single triangular bract, adnate to the 

inflorescence axis a short distance above the bract, bearing ± distichous, minute, 

triangular, incomplete bracts, each subtending a pair of equal flowers without 

bracteoles. Flowers very pale in colour, becoming darker post-anthesis, moderately to 

strongly fragrant; calyx thick, coriaceous, very shallowly 3-lobed distally, obscurely 

veined, minutely papillose, corolla very thick, coriaceous, divided at the apex to ¼ to 

1/3 its length into 3 short, triangular valvate lobes, remaining approximate, even at 

anthesis, the lobes then separating slightly to reveal a discrete trilete opening; stamens 

6, united into a fleshy epipetalous ring, clasping the gynoecium and occluding the 

mouth of the flower, free filaments angled, very short, anthers enclosed within the 

flower, ± medifixed, very short, somewhat sagittate, latrorse; pollen elliptic, 

monosulcate, the sulcus extended, exine foveolate, tectate; gynoecium tricarpellate, 

triovulate, rounded, covered in reflexed scales, tipped by a columnar or tapered, 3angled 

style, apically with 3 stigmatic angles, ovule basally attached, anatropous. Fruit 

1-3 seeded, stigmatic remains minute, apical, perianth whorls persistent; epicarp 

covered in vertical rows of reddish-brown reflexed scales with fringed margins, 

mesocarp fleshy at maturity, endocarp not differentiated, sarcotesta, when present, 

very thin. Seed sub-basally attached, from the shape of 1/3 of a sphere to 

hemispherical or ellipsoid (depending on the number of seeds developing), sometimes 

slightly lobed or grooved, with a conspicuous abaxial ridge opposite the embryo, seed 

coat thin, rarely fleshy, endosperm homogenous; embryo lateral. Germination 

adjacent-ligular; eophyll bifid. 

Distribution 

Eremospatha is represented by ten species confined to the lowland (

Zambia. 

Notes 

The leaves of this genus display a remarkable plasticity of form, from bifid juvenile 

leaves to regularly or irregularly pinnate adult leaves. Historically, this has caused 

much taxonomic confusion and a number of species have been described from 

juvenile material. Although recent collections and field observations have clarified the 

species somewhat, the taxonomic picture may not yet be complete and a number of 

new species may be yet discovered, particularly in the Cameroon-Congo region. 

Key to the species of Eremospatha 

Stem with sheaths ±1 cm in diameter; knee highly conspicuous or absent; leaflets 

regularly arranged, or moderately inequidistant; 

Knee present, conspicuous beneath leaf: 

Ocrea entire, horizontally or obliquely truncate, or somewhat saddleshaped: 

Stem ± circular in cross-section; mature leaflets few (

Knee absent: 

Stem ± triangular in cross-section; mature leaflets many in 

number (up to 30 on each side), linear-lanceolate to ovate, 

lowermost leaflets reduced, linear, strap-like; cirrus unarmed: 

E. laurentii 

Ocrea longitudinally splitting into v-shape, or sometimes tattering; 

Leaflets rhomboid or trapezoid with straight margins; cirrus 

armed with reflexed spines; bracts on inflorescence minute, 

inconspicuous

Leaflet apex entire, terminating in a conspicuous apiculum: E. 

cuspidata 

Stem with sheaths ±1 cm in diameter; knee inconspicuous, linear, ridge-like; leaflets 

conspicuously inequidistant, in groups, clustered or somewhat plumose: E. 

quinquecostulata 

______________________________________ 

E. hookeri (G. Mann & H. Wendl.) H. Wendl. 

Joseph Dalton Hooker (1817-1911), botanist, and former Director of the Royal 

Botanic Gardens, Kew 

H. Wendl. in Kerchove, Les Palmiers 244 (1878); Drude in Engl. Bot. Jarbh. 5: 131 

(1895); C.H. Wright in F.T.A. 8: 112 (1901); Durand & Durand in Fl. Cong. 1: 585 

(1909); Becc. in Webbia 3: 281 (1910); De Wild. in Pl. Thon. Cong. 2: 142 (1911); 

Hutch. in F.W.T.A. 2: 391 (1936); Guinea-Lopez in Ensayo Geobot. de la Guinea 

Cont. Espanola 245: (1946); Fosberg in Principes 4: 129 (1960); Irvine in Woody 

Plants of Ghana 780 (1961); Russell in F.W.T.A. 2(3): 168 (1968); Letouzey in Adan. 

18(3): 314 (1978); Morakinyo in Principes 39(4): 200 (1995); Tuley in Palms of 

Africa 45: (1995); H.A. Burkill in Useful Pl. of W. Trop. Africa 4: 370 (1997); Cable 

& M. Cheek in Pl. of Mt. Cam. 179: (1998). 

Calamus (Eremospatha) hookeri G. Mann & H. Wendl. in Trans. Linn. Soc. 24: 434 

(1864); Type: Nigeria, mouth of the River Niger, Mann 451 (holotype K!). 

Clustering moderate to robust palm climbing to 30 m. Stems often branching, circular 

in cross-section, without sheaths, 15-20 mm in diameter, with 20-30 mm; internodes 

16-20 cm long, commonly less (10-12 cm). Leaf sheath longitudinally striate, sparsely 

to profusely covered with caducous black indumentum, or indumentum absent; ocrea 

entire, obliquely truncate, or with high rounded lobe adaxial to the rachis, often drying 

grey-brown; knee linear, 1.5-3 cm long, somewhat abrupt at base. Juvenile stems up to 

15m long; stem with sheath

15 cm broad, deeply notched with rounded, somewhat rectangular lobes; cirrus up to 

60 cm long, emerging from the centre; elaminate rachis up to 80 cm long. Leaves on 

mature stems sessile, up to 2.2 m long; rachis up to 1.5 m long, abaxially rounded 

adaxially concave, becoming rounded, rarely triangular, in cross section distally, 

armed as the petiole, although spines becoming more sparse distally, underside of 

rachis with sparse black caducous indumentum, absent distally; cirrus 50-70 cm long, 

armed as the rachis; leaflets up to 20 on each side, very variable in shape, obovateelliptic, 

oblanceolate to almost rhomboid, bluntly contracted at base, finely to very 

broadly praemorse at apex, 12-22 cm long x 3.5-5.5 cm broad at the widest point, 

discolorous, adaxially dark green, abaxially mid-green, armed along the margins with 

inequidistant forward and (rarely) backward-facing black-tipped spines, c.5-7 main 

veins radiating from the base; lowermost leaflets smaller than the rest, linear to ovoid, 

reflexed and laxly to tightly clasping sheath; acanthophylls 2-2.5 cm long. 

Inflorescence, glabrous, up to 40 cm long; peduncle 12-18 cm long; rachis 18-22 cm 

long, somewhat straight; rachis bracts 0.5-1.5 mm, bluntly triangular; rachillae 

distichous, 8-10 on each side, 8-12 cm long, decreasing distally, adnate to the 

inflorescence axis for 3-7 mm, with

Figure 1. Eremospatha hookeri (G. Mann & H. Wendl.) H. Wendl. 

Sunderland 2258; a. Stem x 1 (2/3): Sunderland 2261; b. Leaflets x 1 (2/3): 

Sunderland 2256; c. Leaflets x 1 (2/3): Sunderland 2257; d. Juvenile leaf x ½ (1/3): 

Mann 451; e. Fruit on infructescence x 1 ½ (1), f. Seed x 1 ½ (1). Drawn by Lucy 

T. Smith. 

54

Distribution 

This species occurs from Côte d’Ivoire in the Upper Guinea region, through to the 

northern Congo Basin. 

Figure 2. Distribution of E. hookeri (G. Mann & H. Wendl.) H. Wendl. 

Habitat and ecology 

E. hookeri is particularly shade tolerant and is often found under a forest canopy. 

However, this species is also common in gaps and in forest margins and is found in a 

wide range of edaphic conditions, from swamp vegetation to well-drained volcanic 

soil. 

Notes 

E. hookeri is a very variable species exhibiting a great plasticity in leaflet and knee 

shape and ocrea form in particular. However, examinations of the limited collections 

from the Upper Guinea forests suggest this might be a distinct taxon. Further 

collections, particularly of fertile material, will probably provide sufficient 

morphological justification for splitting this taxon into two distinct species. 

55

Specimens examined 

SIERRA LEONE: Gledhill 309, Lake Soufon, sterile, February 14, 1966 (K!, GC!); Scott-Elliot 4442, 

Mofani, sterile, January 12, 1892 (K!); Small 832, Kambui Hills (07.05N:11.20W) sterile, November 4, 

1952 (K!); GHANA: Hall & Abbiw 45124, Subiri F.R. (05.17N:01.43W) sterile, January 2, 1975 

(GC!); Moore & Enti 9888, Ankasa River F.R. (05.15N:02.36W) sterile, March 3, 1971 (GC!); 

Sunderland 2261, Draw River Forest Reserve (05.12N:02.20W) sterile, May 26, 1999 (K!, KUM!); 

Tomlinson s.n., Bobiri F.R.(06.38N:01.17W) juvenile, December 20, 1957 (K!); NIGERIA: Ayewoh 

3852, Ondo Province, Owo (08.25N:03.20E) juvenile, February 24, 1944 (K!); Maggs 160, Kwa Falls 

near Calabar (04.59N:08.20E) sterile, August 26, 1948 (K!); Mann 451, River Niger (05.00N:06.00E) 

Fr., August 1860 (K!); Morakinyo 1005, Cross River National Park (05.15N:08.42E) sterile, August 17, 

1993 (K!); Onochie 7706, Obutung forest, Calabar (05.02N:08.21E) sterile, March 2, 1945 (K!); Tuley 

651, Oban rubber estate, Calabar (05.02N:08.21E) sterile, July 12, 1964 (K!); CAMEROON: Dinklage 

1155, Grand Batanga (02.23N:09.50E) sterile, February 18, 1891 (MO!); Kalbrayer 65, SW Province, 

sterile, July, 1904 (K!); Sunderland 2302, Korup National Park, Chimpanzee Camp (05.02N:08.48E) 

sterile, February 15, 2000 (K!, SCA!); Sunderland 1760, Limbe - Kumbe road: Mile 40 

(04.23N:09.26E) sterile, November 11, 1996 (K!, SCA!, WAG!); Sunderland 1801, Campo Ma'an 

Faunal Reserve (02.10N:09.54E) sterile, March 24, 1997 (K!, YA!, BH!, NY!, MO!, WAG!); 

Sunderland 1890, 30km south of Kribi (02.48N:09.43E) sterile, December 2, 1997 (K!, YA!, BH!, 

NY!, MO!, WAG!); Sunderland 2256, Mokoko River Forest Reserve (04.29N:09.00E) sterile, February 

16, 1999 (K!, SCA!, MO!); Sunderland 2257, Mokoko River Forest Reserve (04.29N:09.00E) sterile, 

February 16, 1999 (K!, SCA!, BR!); Sunderland 2258, Mbanga - Nkongsamba road (04.25N:09.33E) 

sterile, February 23, 1999 (K!, YA!, BH!, NY!); Thomas 5163, Korup National Park (05.01N:08.51E) 

sterile, February 20, 1986 (YA!); Thomas 10059, Mokoko River Forest Reserve (04.25N:09.02E) 

sterile, May 22, 1994 (SCA!); EQUATORIAL GUINEA: Sunderland 1906, near village of Njakem 

(01.42N:09.40E) sterile, March 24, 1998 (K!, EG!, WAG!); Sunderland 1917, 2km WSW of village of 

Basilé (01.10N:09.50E) sterile, April 7, 1998 (K!, EG!); GABON: le Testu s.n., Haute-Ngounye 

(00.22S:10.27E) sterile, s.d. (BR!); CONGO: Bermejo 88, Parc National d’Odzala (00.36N:14.54E) 

sterile, 1993 (BR!); Thomas et al. 8944, Bessié village (01.54N:13.56E) sterile, November 23, 1991 

(MO!) 

______________________________________ 

E. cabrae (De Wild. & Th. Dur.) De Wild. 

Captain E. Cabra, Belgian administrator and explorer 

De Wild. in Ann. Mus. Congo 5(1): 95 (1904); Durand & Durand in Fl. Cong. 1: 585 

(1909); De Wild. in Ann. de Mus. Col. de Marseille 3(7): 20 (1919); Chev. in Rev. de 

56

Bot. Appl. 17: 896 (1936); Renier in Fl. du Kwango 1: 82 (1948); Walker & Sillans in 

Plantes Utilès du Gabon 331: (1961); Letouzey in Adan. 18(3): 314 (1978); Tuley in 

Palms of Africa 45: (1995). 

Calamus cabrae De Wild. & Th. Dur. in Matér. pour la Fl. de Congo 5: 32 (1897); 

Baudon in Rev. de Bot. Appl. 4: 595 (1924); Type: DR Congo, Mayombe, Cabra s.n. 

(holotype BR!). 

E. rhomboidea Burr. in Notizbl. Bot. Gart. Mus. Berlin-Dahlem 15: 747 (1942); 

synon. nov. Type: Angola, Nkanda Mbaku, Gossweiler 10086 (holotype B†; isotype 

K!). 

E. suborbicularis Burr. in Notizbl. Bot. Gart. Mus. Berlin-Dahlem 15: 747 (1942); 

synon. nov. Type: Angola, Nkanda Mbaku, Gossweiler 10088 (holotype B†; isotype 

K!). 

Clustering moderate palm climbing to 50 m, more commonly 20-30 m. Stems ± 

circular in cross-section, without sheaths 10-15 mm in diameter, with to 25 mm; 

internodes 10-15 cm long. Leaf sheath longitudinally striate, sparsely to moderately 

armed with black caducous indumentum, particularly concentrated on the apex of the 

sheath; ocrea entire, somewhat saddle-shaped, with 1-1.5 cm rounded lobe adaxial to 

the leaf; knee narrow, linear, abrupt, up to 2.5 cm long. Leaves sessile, up to 1.5-2 m 

long; rachis up to 1 m long, abaxially rounded, adaxially flattened or slightly convex, 

becoming trapezoid then rounded in cross-section distally, with sparse to moderate 

black caducous indumentum below, armed along the margins with inequidistant 

reflexed, bulbous-based, black-tipped spines, becoming more sparsely armed distally; 

cirrus up to 1 m long, armed as the rachis proximally, becoming unarmed distally; 

leaflets up to 8-10 on each side of the rachis, obovate to trapeziform, narrowly 

contracted at the base, irregularly and broadly praemorse at apex, 7-16 cm long x 4-9 

cm broad at the widest point, concolorous, somewhat coriaceous, with 8 or more main 

veins radiating from base, armed along the margins with stout forward and (rarely) 

reverse-facing angular spines, praemorse apex somewhat ciliate-spiny; lowermost 

leaflets smaller than the rest, erect or reflexed and laxly swept back across the sheath; 

57

acanthophylls up to 3.5 cm long, very fine, slender. Inflorescence with profuse, soft, 

velvety, papillose covering, up to 40 cm long, arching, rarely straight; peduncle up to 

20 cm long; rachis 18-28 cm long, rachis bracts 1.5-2 cm, finely acuminate; rachillae 

distichous, 10-12 on each side, up to 10 cm long, decreasing distally, rarely straight, 

often arching, adnate to the inflorescence for 10 mm, decreasing distally. Flowers 

borne in close pairs, with

Figure 3. Eremospatha cabrae (De Wild. & Th. Dur.) De Wild. 

Leonard 929; a. Stem & inflorescence x 1 (2/3): Louis 3804; b. Leaflets x 3 (2), c. Flower x 3 (2), d. 

Flower section x 3 (2): Louis 5656; e. Fruit x 1 ½ (1). Drawn by Lucy T. Smith. 

59

Distribution 

E. cabrae is restricted to Gabon, southwards to Angola and across to the lowland 

forests of the wider Congo basin. This species is reported to be relatively common 

where it occurs (de Wildeman, 1904). 

Figure 4. Distribution of E. cabrae (De Wild. & Th. Dur.) De Wild. 


E. cabrae is more commonly encountered in swamp forest, and rarely in terra firme 

forest. 


CENTRAL AFRICAN REPUBLIC: Harris 3419, 6km from Bayanga (02.55N:16.16E) sterile, July 

7, 1993 (K!); Harris 4765, 8km ESE of Lidjombo (02.39N:16.11E) sterile, March 16, 1994 (K!); 

Harris 4966, 25km SE of Bayanga (02.47N:16.25E) Fl., May 27, 1994 (K!); GABON: Williamson 128, 

Lopé Reserve (00.30N:11.32E) sterile, January 1985 (K!); de Wilde et al. 11177, road from airport to 

Vera (02.47S:10.06E) Fl., November 23, 1994 (WAG!); CONGO: Hallé 1814, 20km N of Brazzaville 

(04.05S:15.17E) Fl., February 2, 1970 (BR!); DR CONGO: Allard 294, Kanya, sterile, 1910 (BR!); 

Bequaert 953 Bumba (02.12N:22.27E) Fr., October 28, 1913 (BR!); Cabra s.n., Haut-Chiluango, 

60

sterile, 1904 (BR!); Cabra s.n., Mayombe, sterile, January 21, 1897 (BR!); Compère 2182, Kinkosi 

(04.33S:14.35E) sterile, October 17, 1910 (BR!); Demeuse s.n., Mangobo (00.20S:28.06E) sterile, May 

1892 (BR!); Deuse 121, Lac Tumba (00.46S:20.06E) juvenile, May 6, 1955 (BR!); Germain 326, 

Yangambi (00.45N:24.26E) Fl., May 15, 1940, BR!; Gillet 2069, Kimengue, Fl. & Fr., March 1900 

(FI!, BR!); Hulstaert 1417, Bokela (01.07S:21.55E) Fr., s.d. (BR!); Jans 655, Bokoro, Fl., January 17, 

1948 (BR!); Kuasa 48, Lukula (05.23S:12.56E) Fr., November 26, 1960 (BR!); Laurent 912, Bamama, 

Fl., February 28, 1896 (BR!); Laurent 1118, Eala (00.03N:18.18E) Fr., November 10, 1902 (BR!); 

Laurent s.n., Eala (00.03N:18.18E) sterile, July 1, 1902 (BR!); Lejoly 1438, Kisangani (00.30N:25.15E) 

sterile, April 30, 1977 (BR!); Lejoly 82/820, Batekes Plateau (04.35S:16.20E) Fl., December 9, 1982 

(BR!); Leonard 929, Eala (00.03N:18.18E) Fl., October 1946 (K!, WAG!, BR!); Liegeois 88, Tshopo 

(10.12S:24.51E) sterile, July 1943 (BR!); Lisowski 86336, Batekes Plateau (04.35S:16.20E) Fl., July 9, 

1982 (BR!); Louis 3804, Yangambi (00.45N:24.26E) Fl., April 30, 1937 (K!, WAG!, BR!); Louis 5656, 

Yangambi (00.45N:24.26E) Fr., July 27, 1937 (K!, FHO!, BR!, MO!); Louis 15169, between Yangambi 

and Isangi (00.43N:24.23E) Fl., June 12, 1939 (BR!); Louis 16797, Yangambi (00.45N:24.26E) Fr., 

November 17, 1943 (BR!); Nannan 46, Eala (00.03N:18.18E) sterile, August 25, 1914 (K!); 

Nsimundeue 1055, Luki (05.26S:12.44E) Fl., July 17, 1972 (BR!); Pauwels 2322, Kisantu 

(04.25S:14.42E) sterile, April 13, 1959 (BR!); Pynaert 1673, Eala (00.03N:18.18E) Fl., August 20, 

1907 (BR!); Pyneart 1073, Eala (00.03N:18.18E) Fr., 1907 (K!); Sapin s.n., Haut-Chiluango, Fr., 

January 1910 (BR!); Toussaint 2331, Vallée de la Nkula, sterile, May 20, 1947 (BR!, MO!); Wellens 

473, Lubumba (03.57S:29.05E) sterile, December 1923 (BR!); ANGOLA: Gossweiler 10086, 

Mayombe, Luali (05.00S:12.25E) sterile, 1923 (K!); Gossweiler 10088, Mayombe, Luali 

(05.00S:12.25E) sterile, 1923 (K!) 

______________________________________ 

E. laurentii De Wild. 

Marcel Laurent (1879-1924), Belgian botanist 

De Wild. in Bull. Jar. Bot. Brux. 5: 147 (1916); De Wild. in Ann. de Mus. Col. de 

Marseille 3(7): 21 (1919); Renier in Fl. du Kwango 1: 82 (1948); Morakinyo in 

Principes 39(4): 202 (1995); Tuley in Palms of Africa 45: (1995); Cable & M. Cheek 

in Pl. of Mt. Cam. 179: (1998); Type: DR Congo, between Bolobo and Yumbi, 

Laurent 645 (holotype BR!). 

Clustering robust palm climbing to 30 m long. Stems ± triangular in cross-section, 

without sheaths, 18-24 mm in diameter, with 25-30 mm; internodes 10-16 cm. Leaf 

sheath lightly striate, moderately to profusely covered in caducous grey-black 

61

indumentum, or indumentum absent; ocrea entire, obliquely truncate, extending for 1- 

2 cm; knee conspicuous, narrow, linear, 5-8 cm long, rather abrupt at base. Leaves 

sessile, up to 3m long; rachis 1.2-1.5m long, abaxially rounded, adaxially convex to 

concave, becoming trapezoid then triangular in cross-section distally, sparse grey 

indumentum present below, or absent, armed along the margins with robust reflexed, 

bulbous-based, black-tipped, spines, becoming sparsely armed distally; cirrus 1.2-1.5 

m long, unarmed; leaflets up to 30 on each side of the rachis, inequidistant, opposite to 

sub-opposite, linear-lanceolate to ovate, bluntly contracted at the base, very finely 

acuminate at apex, with apex often breaking off giving slightly blunt appearance, 22- 

38 cm long x 2.8-3.8 cm broad at the widest point, concolorous, armed along the 

margins with slender to robust black-tipped yellow spines, with c.6 moderately 

conspicuous transverse veinlets 1-2 mm apart; lowermost leaflets, smaller than the 

rest, linear, strap-like, armed along the margins with robust bulbous-based blacktipped 

yellow to orange spines, laxly swept back across, or tightly clasping stem; 

acanthophylls 3-4 cm long. Inflorescence glabrous, 24-32 cm long; peduncle 10-15 

cm long, somewhat flattened in cross-section; rachis 12-17 cm long, erect, arching; 

rachillae distichous 8-10 on each side, 5-16 cm long, decreasing distally, adnate to the 

inflorescence axis for 3-5 mm; rachis bracts, acuminate,

Figure 5. Eremospatha laurentii De Wild. 

Sunderland 1920; a. Stem x 2/3 (4/9), b. Stem section x 2/3 (4/9), c. Leaf x 2/3 (4/9): Louis 15994; d. 

Flower x 2 (2 2/3), e. Flower section x 2 (2 2/3): Otedoh and Tuley 7258; f. Infructescence section x 1 

(2/3), g. Seed x 1 ½ (1). Drawn by Lucy T. Smith. 

63

Distribution 

This species occurs predominantly in the lowland forests of the northern Congo Basin. 

However, intriguingly, there are outliers of this species found in the forests of Upper 

Guinea, with a pronounced disjunction from Côte d’Ivoire to Benin. 

Figure 6. Distribution of E. laurentii De Wild. 


E. laurentii is found in both open areas as well as in closed-canopy forest. However, 

this species responds particularly well to selective logging and is a common 

component of regrowth vegetation where it occurs. 


SIERRA LEONE: Deighton 4117, Gola forest (07.45N:10.45W) Fl., March 10, 1945 (K!); LIBERIA: 

Harley 2174, Wanana (07.25N:09.31W) Fr., January 24, 1958 (K!); NIGERIA: Onochie 5243, Olujiji 

(08.11N:04.08E) sterile, December 13, 1957 (K!); Otedoh & Tuley 7258, Ologbo, near Sapele 

(07.59N:04.25E) Fr., s.d. (K!); Tuley & Ochie 1682, South of Maraba, sterile, October 4, 1969 (K!); 

CAMEROON: Cheek 5554, Mount Cameroon: Njonji (04.04N:08.59E) sterile, November 24, 1993 

64

(SCA!); de Wilde 2183, 60km S of Eseka (03.39N:10.46E) Fr., March 20, 1964 (WAG!); Dransfield 

7003, Mungo River Crossing (04.08N:09.31E) Fr., June 27, 1991 (K!, SCA!); Letouzey 4278, 40km S 

of Mesamena (03.19N:12.49E) sterile, February 16, 1962 (YA!); Letouzey 4416, 20km E of Somalonyo 

in Dja (03.00N:12.40E) Fl., February 24, 1962 (YA!); Letouzey 11796, 25km NNE of Mintom II 

(02.03N:13.30E) sterile, Janaury 5, 1973 (YA!); Letouzey 12477, Lake Ossa, 8km WNW Edea 

(03.50N:10.02E) Fl., December 22, 1966 (K!, YA!); Letouzey 14522, Rumpi Hills nr Lokando 

(04.54N:09.20E) sheath only, March 23, 1976 (YA!); Letouzey 14748, 25km N of Douala 

(04.18N:09.43E) Fr., August 29, 1976 (YA!); Njingum 4, Mbalmayo (03.31N:11.30E) sterile, June 15, 

1999 (K!); Njingum 8, Akom II (02.47N:10.34E) sterile, July 3, 1999 (K!); Sunderland 1752, Mungo 

River Crossing (04.08N:09.31E), sterile, November 16, 1996 (K!, SCA!, BH!); Sunderland 1766, 

Southern Bakundu Forest Reserve (04.46N:09.29E) sterile, November 24, 1996 (K!, SCA!, NY!); 

Sunderland 1805, Campo Ma'an Faunal Reserve (02.10N:09.54E) sterile, March 27, 1997 (K!, YA!); 

Sunderland s.n., Sud Province (02.24N:09.54E) sterile., s.d. (K!, YA!); Watts 514, Mount Cameroon: 

Njonji (04.04N:08.59E) sterile, October 15, 1992 (SCA!); CENTRAL AFRICAN REPUBLIC: 

Harris & Fay 459, Ndakan (02.22N:16.09E) Fr., April 4, 1988 (MO!, BR!); EQUATORIAL 

GUINEA: Sunderland 1920, on road to Monte Mitra (01.12N:09.59E) sterile, April 7, 1998 (K!, EG!); 

GABON: Klaine s.n., Libreville (00.35N:09.22E) Fr., January 15, 1907 (FI!); van Nek 517, Gamba 

(02.47S:10.03E) sterile, December 31, 1990 (WAG!); DR CONGO: Bequaert 878, between Bolobo & 

Sandy Beach (02.08S:16.15E) sterile, October 14, 1913 (BR!); Couteaux 473, Eala (00.03N:18.18E) 

Fr., October 20, 1908 (BR!); Dubois 912, Maringa (00.07N:21.17E) Fr., August 1938 (K!, BR!); 

Evrard 2984, between Mangania and Lifoku (00.75S:21.03E) Fr., November 19, 1957 (BR!); Evrard 

4511, Parc National de Monkoto (00.08N:19.16E) sterile, August 6, 1958 (K!, WAG!, BR!); Evrard 

7070, Nselé (04.14S15.34E) Fr., May 8, 1975 (BR!, WAG!, MO!); Gerard 2152, Bambesa 

(03.28N:25.11E) Fl., February 20, 1956 (BR!); Germain 1681, Eala (00.03N:18.18E) Fr., October 28, 

1943 (BR!); Germain 4808, Ikelemba river, Fl., February 1949 (BR!); Gilbert 7909, Yangambi 

(00.45N:24.26E) Fl., 1947 (BR!); Hulstaert 747, Bokuma, Fl., March 8, 1942 (BR!); Laurent 645, 

between Bolobo & Yumbi (02.08S:16.15E) Fl. & Fr., April 14, 1903 (BR!); Leonard 55, between 

Bamania & Ilelenge, Fr., September 26, 1945 (K!, MO!, BR!); Leonard 816, Eala (00.03N:18.18E) Fr., 

October 12, 1946 (BR!); Leonard 980, Eala (00.03N:18.18E) sterile, November 11, 1946 (K!, BR!, 

WAG!); Louis 10155, Yangambi (00.45N:24.26E) Fr., July 1, 1938 (K!, BR!, WAG!); Louis 11439, 

between Yangambi and Basoko (01.12N. 23.51E) Fl. & Fr., September 1938 (K!, BR!, WAG!); Louis 

15925, Yangambi (0.45N:24.26E) Fr., August 25, 1939 (K!, BR!); Louis 15944, Yangambi 

(00.45N:24.26E) Fl., August 28, 1939 (K!, BR!, WAG!); Louis 16791, Yangambi (00.45N:24.26E) Fr., 

November 17, 1943 (BR!); Louis 7994, Yangambi (00.45N:24.26E) Fr., February 22, 1938 (K!, BR!); 

Mandango 2970, Basoko (01.15N:23.36E) Fr., May 13, 1981 (BR!); Sapin s.n., sterile, 1912 (BR!); 

Thonet 129, Lac Tumba (00.46S:20.06E) Fr., February 5, 1957 (BR!); CULTIVATED: Java, Furtado 

A113, Bogor Botanic Garden, sterile (K!); Palm House, RBG Kew, 1984-1058 (K!) 

______________________________________ 

65

E. wendlandiana Dammer ex Becc. 

Hermann Wendland (1825-1903) German palm botanist and horticulturist 

Becc. in Webbia 3: 290 (1910); Hutch. in F.W.T.A. 2: 391 (1936); Guinea-Lopez in 

Ensayo Geobot. de la Guinea Cont. Espanola 245: (1946); Russell in F.W.T.A. 2(3): 

168 (1968); Letouzey in Adan. 18(3): 314 (1978); Letouzey in Man. For. Bot. Trop. 

Afr. 2B: 401 (1986); Morakinyo in Principes 39(4): 204 (1995); Tuley in Palms of 

Africa 45: (1995); Cable & M. Cheek in Pl. of Mt. Cam. 179: (1998; Type: 

Cameroon, Lake Barombi, Preuss 460 (holotype B†; isotype FI!). 

Eremospatha korthalsiaefolia Becc. Webbia 3: 292 (1910); Walker & Sillans in 

Plantes Utilès du Gabon 333: (1961); synon. nov. Type: Cameroon, Akoafim near 

Ebolowa, Dusen 292 (holotype B†; isotype FI!). 

Clustering moderate to robust palm climbing to 60 m. Stems ± circular in crosssection, 

without sheaths 12-20 mm, with 15-30 mm; internodes up to 30 cm long. Leaf 

sheath only very lightly striate, with sparse to moderate caducous black indumentum; 

ocrea drying brown and splitting longitudinally, sometimes with horizontal linear 

wrinkle opposite the leaf, extending to up to 8 cm; knee conspicuous, narrowly-linear, 

up to 5-12 cm long, tapering at base. Leaves sessile; rachis up to 2m long, abaxially 

rounded, adaxially flattened or slightly concave on upper surface, becoming rounded 

then triangular in cross-section distally, armed along the margins with inequidistant, 

reflexed, bulbous-based, black-tipped spines, with sparse black indumentum below; 

cirrus up to 2 m long, armed as the rachis, although spines becoming sparse distally, 

indumentum absent; leaflets up to 20 on each side of the rachis, strictly rhomboid or 

trapezoid with conspicuously straight margins, broadly attenuate at the base, broadly 

and irregularly praemorse at apex, very variable in size 12-22 cm long x 8-17 cm 

broad at the widest point; concolorous, armed along the margins with 2 mm long, 

robust, slightly reflexed, black-tipped spines, praemorse apex somewhat ciliate spiny; 

with 5-9 main veins radiating from the base; acanthophylls 2-2.5 cm long, somewhat 

slender, at 30° angle to cirrus. Inflorescence glabrous, up to 80 cm long, peduncle up 

to 30 cm long; rachis up to 50 cm long, arching, rachis bracts incomplete, 1.5-2 cm 

66

long, decreasing distally, rachillae distichous, 10-12 on each side, 25-30 cm long, 

decreasing distally, adnate to the rachis for up to 2 cm, less so distally. Flowers borne 

in close pairs; calyx 2 mm long x 4 mm wide at the mouth, shallowly 3-lobed; corolla 

c. 8 mm long x 3 mm wide divided to ¼ of its length; stamens united into 3 mm-long 

epipetalous ring, free filaments

Figure 7. Eremospatha wendlandiana Dammer ex Becc. 

Sunderland 1798; a. Stem x ¾ (1/2), b. Leaflets x 2/3 (4/9), c. Acanthophylls x 1 (2/3): Nkefor 920; 

d. Fruit on infructescence x 1 (2/3). Drawn by Lucy T. Smith. 

68

Distribution 

E. wendlandiana is distributed from SE Nigeria to Gabon, commonly in coastal forest, 

although with outliers present in the swamp forests of the Central African Republic. 

Figure 8. Distribution of E. wendlandiana Dammer ex Becc. 


This species is a common component of gap vegetation and forest margins, although it 

is commonly present in the juvenile form in closed-canopy forest where it occurs. 


NIGERIA: Aninze 15402, Oban Hills (06.00N:09.15E) Fr., February 9, 1946 (K!); Morakinyo 1001, 

Cross River National Park (05.15N:08.42E) Fr., August 11, 1993 (K!); Tuley 652, Mile 26 on Calabar 

to Okpora road (05.60N:08.02E) sterile, July 13, 1964 (WAG!); CAMEROON: Dransfield 7004, Mile 

40, Buea-Kumba road (04.23N:09.26E) sterile, June 28, 1991 (K!, SCA!); Dundas 8381, Southern 

Bakundu F.R. (04.26N:09.21E) seedling, August 20, 1945 (K!); Harris 3738, Onge (04.21N:08.57E) 

sterile, September 11, 1993 (K!, SCA!); Iquito 61, Fr. only (FI!); Letouzey 4151, 45km NE of Kribi 

(03.15N:10.12E) sterile, January 25, 1962 (YA!); Letouzey 11518, Nkongkengui, 12km NNE of Makak 

(03.34N:11.02E) sterile, July 17, 1972 (YA!); Letouzey 11800, Mintom I, 70km E of Djoum 

69

(02.03N:13.30E) sterile, January 8. 1973 (YA!); Lowe 3443, Edea - Kribi rd nr Elogbatindi 

(03.27N:10.11E) Fr., January 22, 1978 (K!, YA!); Mildbraed 6036, 58km east of Kribi 

(02.56N:10.26E) sterile, July 21, 1911 (HBG!); Nkefor 920, Southern Bakundu Forest Reserve 

(04.46N:09.29E) Fr., s.d. (K!); Preuss 460, Barombi (04.40N.09.23E) sterile, 1890 (FI!); Richards 

5209, Kembong Forest Reserve, SW Province (05.38N:09.14E) sterile, March 16, 1955 (K!); Rosevear 

30138, Kumba (04.38N:09.26E) sterile, October 29, 1937 (FHO!); Sunderland 2304, Korup National 

Park, Chimpanzee Camp (05.02N:08.48E) sterile, February 15, 2000 (K!); Sunderland 1640, Mokoko 

River Forest Reserve (04.29N:09.00E) sterile, May 1, 1994 (K!, SCA!); Sunderland 1701, Southern 

Bakundu Forest Reserve (04.46N:09.29E) sterile, November 8, 1995 (K!, SCA!, NY!); Sunderland 

1712, Onge River valley (04.21N:08.57E) sterile, November 23, 1995 (K!, SCA!, MO!); Sunderland 

1719, 30km north of Mamfe (05.58N:09.20E) sterile, December 2, 1995 (K!, SCA!, BH!) Sunderland 

1927, Campo Ma'an Faunal Reserve (02.10N:09.54E) sterile, October 11, 1998 (K!, YA!); Sylvanus 

s.n., Obonyi I (06.08N:09.16E) sterile, November 11, 1998 (K!); Thomas 9733, Idenau 

(04.16N:09.01E) sterile, September 10, 1993 (K!, SCA!); Thomas s.n., Korup National Park, 

(04.55N:08.50E) sterile, s.d. (SCA!); Webb & Bullock 310, Campo Faunal Reserve (02.24N:09.54E) 

sterile, July 9, 1976 (K!, YA!); CENTRAL AFRICAN REPUBLIC: Harris 2360, 45km S of 

Lidjombo (02.21N:16.09E) Fl., May 21, 1990 (MO!); EQUATORIAL GUINEA: Sunderland 1798, 

2km north of Ayemeken village (02.10N:10.03E) sterile, March 13, 1997 (K!, EG!, WAG!); 

Sunderland 1876, 2km SW of village of Angoma (02.03N:10.10E) sterile, September 15, 1997 (K!, 

EG!, NY!, MO!, WAG!, BR!); GABON: Breteler et al. 11194, 30km E of Latoursville 

(00.40S:13.00E) Fr., April 30, 1992 (WAG!); de Wilde et al. 9301, 22km from Mayumba 

(03.16S:10.46E) Fr., December 11, 1986 (MO!, WAG!, BR!); Gentry & Emmons 33732, M'Passa field 

stn. Makokou (00.33N:12.50E) sterile, July 31, 1981 (MO!); Klaine s.n., Ogoué River (00.59S:09.03E) 

sterile, s.d. (K!, BR!); ANGOLA: Gossweiler 7567, Mayombe, Luali (05.00S:12.25E) sterile, June 

1920 (K!); Gossweiler 8145, Mayombe, Luali (05.00S:12.25E) sterile, 1920 (K!) 

______________________________________ 

E. barendii Sunderland sp. nov. 

affinis E. macrocarpa (G. Mann & H. Wendl.) H. Wendl. sed vagina cum geniculus, 

ochrea fissa, siccus non integra, spadix leviter pileus non glabellus, conspicue 

bracteatus (3.0-5.0 mm longus non 1.0-3.0 mm). Type: Cameroon, Ebom, near 

Lolodorf (03°44’N:10°43’E) 11 th March, 1997, van Gemerden 77 [infructescence] 

(holotype K!; isotypes YA!, KRI!) 

Clustering palm climbing to 25-30 m. Stems circular in cross section, without leaf 

sheaths, c.15 mm in diameter, with, to 25 mm; internodes 10-15 cm long. Leaf sheath 

longitudinally striate, sparsely to moderately covered with dark brown caducous 

70

indumentum; ocrea obliquely truncate, dry, grey-brown, often splitting in conspicuous 

v-shape on abaxial side, extending to 2 cm; knee linear, 3-3.5 cm long, somewhat 

abrupt at base. Leaf up to 2.4m long; rachis 1-1.2m long, abaxially rounded, adaxially 

flattened or convex, becoming trapezoid then triangular in cross-section distally, 

armed along the margins with inequidistant black-tipped, bulbous-based spines, sparse 

brown indumentum present on underside of rachis, absent distally; cirrus up to 1.2 m 

long, unarmed; leaflets, opposite or sub-opposite, up to 26 on each side of the rachis, 

linear-lanceolate, broadly contracted at the base, apex narrowly praemorse, 25-32 cm 

long x 1.5-2 cm broad at the widest point, concolorous, armed along the margins with 

inequidistant, black-tipped spines, with 5-7 moderately conspicuous transverse 

veinlets 1-2 mm apart; lowermost leaflets smaller than the rest, linear-ovate, erect or 

reflexed and laxly swept across stem; acanthophylls 2.4-2.8 cm long. Inflorescence, 

very lightly papillose up to 30 cm long; peduncle 8-10 cm long; rachis 17-20 cm long, 

arching, sometimes straight, rachis bracts, finely to broadly acuminate, united at base 

to form a conspicuous sheathing bract, 3-5 mm long, decreasing distally; rachillae 

distichous, c.10 on each side, 10-14 cm long, decreasing distally, adnate for 5-8 mm 

of the inflorescence axis, arching, rarely straight, with 1-3 mm circular bracts 

subtending each dyad. Flowers not known. Fruit at maturity, 1-seeded, 2.3-2.8 cm 

long x 1.5-1.7 cm broad, broadly cylindrical, with 16 vertical rows of scales. Seed 

compressed, 2 cm long x 1.2 cm wide x 0.7 cm thick, flattened on one side, embryo 

lateral, raised opposite flattened side. 

71

Figure 9. Eremospatha barendii Sunderland 

van Gemerden 77; a. Stem x 1 (2/3), b. Leaflets x ½ (1/3), c. Infructescence x 2/3 (4/9), d. Fruit x 1 ½ 

(1), e. Seed x 1 ½ (1). Drawn by Lucy T. Smith. 

72

Distribution 

E. barendii is known from a single collection in a logging concession near Lolodorf, 

Cameroon. 

Figure 10. Distribution of E. barendii Sunderland 


This is a very poorly-known species but is thought to occur in gaps in high forest. 

Notes 

This is an unusual species characterised by the large, conspicuous bracts on the 

inflorescence. The dry, splitting ocrea is also an unusual character in this genus, 

shared only by the very distinct E. wendlandiana 

Specimen examined 

CAMEROON: van Gemerden 77, Ebom (03.04N:10.43E) Fr., March 11, 1997 (K!) 

______________________________________ 

73

E. macrocarpa (G. Mann & H. Wendl.) H. Wendl. 

(Latin) “large-fruits” 

H. Wendl. in Kerchove, Les Palmiers 244: (1878); C.H. Wright in F.T.A. 8: 113 

(1901); Becc. in Webbia 3: 272 (1910); Unwin in W. Afr. For. 240: (1920); Hédin in 

Rev. de Bot. Appl. 9: 504 (1929); Hutch. in F.W.T.A. 2: 391 (1936); Dalziel in App. 

F.W.T.A. 507: (1937); Burr. in der Tropenfl. 42(5): 204 (1939); Guinea-Lopez in 

Ensayo Geobot. de la Guinea Cont. Espanola 245: (1946); Fosberg in Principes 4: 

129 (1960); Irvine in Woody Plants of Ghana 780: (1961); Russell in F.W.T.A. 2(3): 

168 (1968); Letouzey in Adan. 18(3): 314 (1978); Hall & Swaine in For. Veg. Ghana 

195: (1981); Profizi in RIC Bull. 5(1): 2 (1986); Hawthorn in Trees of Ghana 225: 

(1990); Morakinyo in Principes 39(4): 202 (1995); Tuley in Palms of Africa 45: 

(1995); H.A. Burkill in Useful Pl. of W. Trop. Africa 4: 370 (1997); Cable & M. 

Cheek in Pl. of Mt. Cam. 179: (1998); Aedo et al. in Bases Docs. Fl. de Guinea 

Ecuatorial 375: (1999). 

Calamus (Eremospatha) macrocarpus G. Mann & H. Wendl. in Trans. Linn. Soc. 24: 

435 (1864); Type: Sierra Leone, Bagroo River, Mann 2330 (holotype K!). 

E. sapini De Wild. in Bull. Jard. Bot. Brux. 5: 147 (1916); De Wild. in Ann. de Mus. 

Col. de Marseille 3(7): 23 (1919); Renier in Fl. du Kwango 1: 82 (1948); synon. nov. 

Type: DR Congo, Thibangu, Sapin s.n. (holotype BR!). 

E. haullevilleana sensu Walker & Sillans in Plantes Utilés du Gabon 331: (1961). 

Clustering slender to moderate palm climbing to 50-75 m, rarely to 150 m. Stems 

circular in cross-section, without sheaths, 10-18 mm in diameter, with 22-30 mm; 

internodes 13-16 cm long. Leaf sheath longitudinally striate, sparsely to moderately 

covered with light brown scale-like indumentum; ocrea entire, ± truncate saddleshaped 

with a 2.5-4.0 cm rounded lobe adaxial to the leaf; knee absent. Juvenile stems 

up to 20m long; stem with sheath,

stems sessile, up to 3.5 m long; rachis 1-1.5 m long, abaxially rounded, adaxially 

flattened, becoming trapezoid then rounded in cross-section distally, armed along the 

margins with inequidistant, reflexed thorns, becoming sparsely armed distally, 

underside of rachis with sparse light brown indumentum; cirrus 1.2-2 m long, 

unarmed; leaflets, up to 25 pairs on each side of the rachis, linear-lanceolate, abruptly 

contracted at the base, irregularly and narrowly praemorse at apex, 22-35 cm long x 2- 

2.5 cm broad at the widest point, concolorous, with 5-7 inconspicuous transverse 

veinlets 1-2 mm apart, armed along the margins with inequidistant, curved, forwardfacing 

brown-tan spines; lowermost leaflets, smaller than the rest, linear-ovate, 

reflexed and laxly clasping the stem; acanthophylls c.3 cm long, parallel to cirrus. 

Inflorescence glabrous, up to 55 cm long; peduncle 10-15 cm long; rachis 25-40 cm 

long, arching outwards, rarely straight; rachis bracts, acuminate, opposite proximally, 

alternate distally, 1-3 mm long, decreasing distally; rachillae distichous, arching 

vertically, sometimes horizontal, straight, 10-14 on each side, 12-18 cm long, 

decreasing distally, adnate to the inflorescence axis for 0.5-1.5 mm, with

Figure 11. Eremospatha macrocarpa (G. Mann & H. Wendl.) H. Wendl. 

Sunderland 1886; a. Stem x 1 (2/3), b. Leaflets x ½ (1/3): Sunderland 1901; c. Juvenile leaf x 3/8 (1/4): 

Sunderland 1886; d. Inflorescence x 1 (2/3): Sunderland 1956; e. Fruit x 1 ½ (1). Drawn by Lucy T. 

Smith. 

76

Distribution 

E. macrocarpa is a very widespread and common species and is distributed from 

Sènègal in West Africa through to the lowland forests of the Congo Basin. 

Figure 12. Distribution of E. macrocarpa (G. Mann & H. Wendl.) H. Wendl. 


This species is extremely light demanding, occurring naturally in gap vegetation and 

forest margins. As a result of this, in common with other members of the genus, E. 

macrocarpa responds extremely well to selective logging activities and is a common 

component of regrowth vegetation. This species occurs more commonly in terra firma 

forest, and is rarely encountered in swamp forest. 


SIERRA LEONE: Deighton 4118, Giewahun (07.36N:11.11W) Fr., March 10, 1945 (K!); Mann s.n., 

Bagroo River (07.45N:12.50W) Fr., 1861 (K!); LIBERIA: Linder 1341, Moala, sterile, November 1, 

1926 (K!, MO!); CÔTE D'IVOIRE: Boughey s.n., Taï Forest (05.38N:07.08W) juvenile, May 5, 1954 

(GC!); Jangoux 215, Tien-Oula (06.45N:07.04W) juvenile, August 3, 1962 (BR!); Oldeman 674, 40km 

N of Bereby (04.54N:07.02W) Fr. only, November 14, 1963 (WAG!); GHANA: Adams 2195, Enchi 

77

(05.29N:02.29W) sterile, December 30, 1953 (GC!); Adams 2214, Enchi (05.29N:02.29W) sterile, 

December 30, 1953 (GC!); Enti 1914, Ankasa F.R. (05.15N:02.36W) Fl., January 30, 1979 (MO!, 

WAG!); Hall 42605, Ankasa F.R. (05.15N:02.36W) Fl., February 14, 1971 (MO!); Hall s.n., Ankasa 

River F.R. (05.15N:02.36W) Fl., February 12, 1971 (GC!); Irvine 4861, Kade Agricultural Research 

Station (06.05N:00.50W) sterile, June 1961 (K!); Irvine 4873, Kade Agricultural Research Station 

(06.05N:00.50W) sterile, June 1961 (K!); Irvine 4886, Kade Agricultural Research Station 

(06.05N:00.50W) sterile, June 1961 (K!); Johnson s.n., Aburi (05.51N:00.10W) Fr., February 25, 1901 

(K!); Kisseadoo 441, Bobiri F.R. (06.38N:01.17W) sterile, November 10, 1988 (MO!); Moore & Enti 

9887, Ankasa River F.R. (05.15N:02.36W) sterile, March 4, 1971 (GC!); Moore & Enti 9891, Ankasa 

River F.R. (05.15N:02.36W) Fl., March 4, 1971 (GC!); Moore & Enti 9893, Ankasa River F.R. 

(05.15N:02.36W) Fl., March 4, 1971 (GC!); Sunderland 2260, Draw River Forest Reserve 

(05.12N:02.20W) sterile, May 26, 1999 (K!, KUM!); Sunderland 2264, Draw River Forest Reserve 

(05.12N:02.20W) sterile, May 26, 1999 (K!, KUM!); Tomlinson s.n., Bobiri F.R. (06.38N:01.17W) 

sterile, December 20, 1957 (GC!); Vigne 1829, Juasso (06.32N:01.06W) Fl., February 28, 1938 (K!, 

KUM!); Vigne 4858, Kumasi District (06.40N:01.39W) Fl., May 30, 1945 (K!, KUM!, BR!); West- 

Skinn 11, Juasso (06.32N:01.06W) juvenile, 1957 (K!); BENIN: Aufsess 425, Adjarra (06.32N:05.52E) 

sterile, December 6, 1988 (K!); Aufsess 426, Adjarra (06.32N:05.52E) sterile, December 6, 1988 (K!); 

Aufsess 429, Adjarra (06.32N:05.52E) sterile, December 6, 1988 (K!); NIGERIA: Daramola & 

Adebusuyi 38415, Kabba Province, Kotokerifi (08.08N:06.44N) Fr., October 24, 1958 (K!); Jones & 

Onochie 17237, Oma & Shasha F.R.'s (07.07N:04.23E) Fr., April 4, 1946 (K!, BR!); Letter 8224, Ikan, 

sterile, 1904 (K!); Lowe 2793, 40 miles SE of Benin City (05.59N:06.07E) juvenile, March 28, 1974 

(K!); Maggs 159, Kwa Falls near Calabar (04.59N:08.20E) juvenile, August 26, 1948 (K!); Mann 2330, 

Cross River (05.15N:08.42E) Fl., February 1863 (K!); Morakinyo 1003, Cross River National Park 

(05.15N:08.42E) seedling, August 15, 1993 (K!); Morakinyo 1004, Cross River National Park 

(05.15N:08.42E) Fr., August 18, 1993 (K!); Niger Company s.n., Oban rubber estate Calabar 

(05.02N:08.21E) sterile, December 15, 1991 (K!); Nwambin & Tuley 603, Ojo road (08.19N:04.14E) 

juvenile, May 4, 1964 (K!); Onochie & Jones FHI 17332, Afi River F.R., Fr., May 28, 1946 (FHO!); 

Thomas 338, Agola District, sterile, 1911 (K!); Tuley 1076, Ikom to Obudu road (06.04N:08.56E) 

sterile, December 10, 1964 (K!); Tuley 1077, Ikom to Obudu road (06.04N:08.56E) seedling, August 

24, 1976 (K!); Tuley 650, Aban Rubber Estate (07.36N:08.56E) Fr., July 12, 1964 (K!, WAG!); Unwin 

109, southern Nigeria, Fr., July 28, 1907 (K!); CAMEROON: Asonganyi 729, Tissongo, 16km EES of 

Mouanko (03.24N:09.50E) Fl., January 20, 1984 (YA!); Brunt 137, Ndop Plain (05.47N:10.15E) 

seedling, March 5, 1962 (YA!); Brunt 207, Ndop Plain (05.47N:10.15E) sterile, March 21, 1962 (K!); 

Dransfield 7005, Mile 45, Buea-Kumba road (05.02N:09.24E) sterile, June 28, 1991 (K!, SCA!); 

Dransfield 7002, Mungo River Crossing (04.08N:09.31E) juvenile, June 27, 1991 (K!); Etuge 1393, 

Mount Kupe (04.46N:09.41E) sterile, November 5, 1995 (SCA!); Faden & Mbama 86/60, Kribi- 

Ebolowa road (02.51N:10.00E) sterile, January 31, 1986 (YA!); Harris 2456, 34km W of Nguti 

(05.00N:09.00E) sterile, August 27, 1990 (K!); Harris 2471, 35km W`of Nguti (05.00N:09.00E) sterile, 

August 27, 1990 (K!); Harris & Payne 2470, 35km W`of Nguti (05.00N:09.00E) sterile, August 29, 

1990 (K!); Letouzey 8465, 10km SE of Sangmelima (02.55N:11.58E) Fl. & Fr., November 24, 1966 

78

(YA!); Letouzey 12563, Lac Tissongo (03.34N:09.53E) Fl. only, January 4, 1974 (YA!); Letouzey 

13843, 10km N of Nguti (05.23N:09.23E) Fr., June 15, 1975 (YA!); Meijer 15221, Sangmelima 

(02.55N:11.58E) sterile, March 24, 1981 (K!, WAG!, YA!); Meijer 15251, Dja Forest Reserve 

(03.00N:12.40E) sterile, March 25, 1981 (K!); Njingum 7, Nguti (05.02N:09.24E) sterile, August 5, 

1999 (K!); Raynal 9785, 17km SW Ambam (02.21N:11.12E) sterile, February 18, 1963 (YA!); 

Sunderland 1702, Southern Bakundu Forest Reserve (04.46N:09.29E) sterile, November 8, 1995 (K!, 

SCA!, MO!); Sunderland 1704, Southern Bakundu Forest Reserve (04.46N:09.29E) sterile, November 

8, 1995 (K!, SCA!, NY!, WAG!); Sunderland 1713, Onge River valley (04.21N:08.57E) sterile, 

November 23, 1995 (K!, SCA!, BR!); Sunderland 1717, Kumba to Mamfe road (05.02N:09.24E) 

sterile, December 1, 1995 (K!, SCA!, BR!); Sunderland 1720, 30km north of Mamfe (05.58N:09.20E) 

juvenile, December 2, 1995 (K!, SCA!, MO!); Sunderland 1721, 30km north of Mamfe 

(05.58N:09.20E) Fl., December 2, 1995 (K!, SCA!, BR!); Sunderland 1730, Rumpi Hills Forest 

Reserve (04.54N:09.20E) seedling, May 19, 1996 (K!, SCA!, BH!); Sunderland 1742, Rumpi Hills 

Forest Reserve (04.54N:09.20E) sterile, May 19, 1996 (K!, SCA!, MO!); Sunderland 1758, Limbe - 

Kumbe road: Mile 40 (04.23N:09.26E) juvenile, November 11, 1996 (K!, SCA!, BR!); Sunderland 

1767, Southern Bakundu Forest Reserve (04.46N:09.29E) juvenile, November 24, 1996 (K!, SCA!, 

BH!); Sunderland 1802, Campo Ma'an Faunal Reserve (02.10N:09.54E) Fl., March 24, 1997 (K!, YA!, 

NY!, WAG!); Sunderland 1807, Campo Ma'an Faunal Reserve (02.10N:09.54E) sterile, April 8, 1997 

(K!, YA!, BH!, MO!); Sunderland 1856, 20km south of Kribi (02.34N:09.50E) Fr., August 30, 1997 

(K!, YA!, NY!, WAG!); Sunderland 1881, Southern Bakundu Forest Reserve (04.46N:09.29E) Fl., 

November 26, 1997 (K!, SCA!, NY!, MO!); Sunderland 1883, Kumba to Mamfe road (05.02N:09.24E) 

juvenile, November 26, 1997 (K!, SCA!, WAG!); Sunderland 1886, 30km south of Kribi 

(02.48N:09.43E) Fl., November 28, 1997 (K!, YA!, NY!, BR!); Sunderland 1937, Takamanda Forest 

Reserve (06.06N:09.47E) sterile, November 17, 1998 (K!, SCA!); Sunderland 1999, 15km north of 

Nguti on Mamfe road (05.23N:09.23E) Fl., January 6, 1999 (K!, SCA!); Sunderland 2042, Takamanda 

Forest Reserve (06.08N:09.16E) sterile, January 13, 1999 (K!, SCA!); Sunderland 2043, Takamanda 

Forest Reserve (06.08N:09.16E) sterile, January 13, 1999 (K!, SCA!); Sunderland 2057, Takamanda 

Forest Reserve (06.08N:09.16E) sterile, January 18, 1999 (K!, SCA!); Sunderland 2252, Mokoko River 

Forest Reserve (04.29N:09.00E) sterile, February 16, 1999 (K!, SCA!); Thomas 8182, 15km W of 

Manyemen (05.10N:09.15E) sterile, August 29, 1988 (MO!); Thomas 10058, Mokoko River Forest 

Reserve (04.25N:09.02E) sterile, May 22, 1994 (SCA!); van Gemerden BL, Ebom II (03.04N:10.43E) 

sterile, March 11, 1997 (K!); CENTRAL AFRICAN REPUBLIC: Fay 7018, N'Dele-Pata road 

(08.08N:21.08E) Fr., May 30, 1985 (MO!); EQUATORIAL GUINEA: Lisowski 1263, Mbini 

(02.00N:09.45E) juvenile, September 1997 (EG!); Sunderland 1797, 2km north of Ayemeken 

(02.10N:10.03E) juvenile, March 13, 1997 (K!, EG!, NY!); Sunderland 1800, 1km north of Ayemeken 

(02.10N:10.03E) sterile, March 13, 1997 (K!, EG!, WAG!); Sunderland 1874, 2km SW of village of 

Angoma (02.03N:10.10E) seedling, September 15, 1997 (K!, EG!, BH!, MO!); Sunderland 1901, 10km 

south of Bata (01.45N:09.43E) sterile, March 20, 1998 (K!, EG!, WAG!); Sunderland 1918, 2km WSW 

of village of Basilé (01.10N:09.50E) juvenile, April 7, 1998 (K!, EG!, WAG!); DR CONGO: Bequaert 

7895, Katala (08.35S:25.16E) Fr., June 28, 1915 (BR!); Dewevre 639, Fl., s.d. (BR!); Dewulf 526, 

79

River L'Uele (03.52N:26.28E) Fl., December 20, 1934 (BR!); Holman-Bentley s.n., Upper River 

Congo, sterile, March 23, 1889 (K!); Hulstaert 864, Bamanga (00.16S:25.32E) Fr., November 1, 1942 

(BR!); Hulstaert 1420, Bokela (01.07S:21.55E) juvenile, s.d. (BR!); Laurent s.n., Basounda, sterile, 

January 10, 1904 (BR!); Laurent s.n., River Kivi, Fr., November 6, 1903 (BR!); Liegeois 86, 28km 

from Buta (02.47N:24.43E) sterile, July 1943 (BR!); Louis 15489, Bengamisa (00.57N:25.10E) 

juvenile, July 9, 1939 (BR!); Pauwels 4543, Kimpoko (04.11S:15.34E) sterile, July 4, 1964 (BR!); 

Sapin s.n., Kasai (10.07S:22.20E) sterile, s.d. (BR!); Sapin s.n., Tshibangu (05.56S:20.54E) Fr., 

January 1910 (BR!) 

______________________________________ 

E. haullevilleana De Wild. 

De Wild. in Ann. Mus. Congo, Bot. 5(1): 96 (1904); De Wild. in Miss. Laurent 24: 

(1905); Durand & Durand in Fl. Cong. 1: 585 (1909); Becc. in Webbia 3: 285 (1910); 

Pyneart in Bull. Agric. du Congo Belge 2: 547 (1911); De Wild. in Ann. de Mus. Col. 

de Marseille 3(7): 19 (1919); Staner & Boutique in Mem. l’Inst. Roy. Col. Belge 14: 

(1937); Guinea-Lopez in Ensayo Geobot. de la Guinea Cont. Espanola 245: (1946); 

Renier in Fl. du Kwango 1: 82 (1948); J. Dransf. in F.T.E.A. (Palmae) 35: (1986); 

Kabuye in Monogr. Syst. Bot. Miss Bot. Gard. 25: 360 (1988); Tuley in Palms of 

Africa 45: (1995); Types: DR Congo, Lubamba, Gillet 2026 & Kisantu, Gillet 1385 

(syntypes BR!; Gillet 2026 iso-syntype FI!) 

Clustering slender to moderate palm climbing to 25 m. Stems circular in cross-section, 

without leaf-sheaths 6-15 mm in diameter, with 10-25 mm; internodes ±15 cm long. 

Leaf-sheath longitudinally striate, bearing sparse black caducous indumentum; ocrea 

entire, obliquely truncate, extending to 3-4 cm; knee absent. Juvenile stems up to 15 m 

long, stem with sheath

and sparse distally; leaflets 8-14 on each side of the rachis, sub-opposite to alternate, 

cuneate, spathulate or ovate with an uneven, moderately to strongly praemorse apex, 

broadly contracted at the base, 9-24 cm long x 2-6 cm broad at the widest point, 

concolorous, with ciliate-spiny margins, up to 10 sub-equal main veins, transverse 

veinlets 1-2 mm distant, moderately prominent; lowermost leaflets smaller than the 

rest, sometimes reflexed and laxly clasping the stem, or absent entirely; acanthophylls 

up to 2 cm long, very fine. Inflorescence glabrous, up to 35 cm long; peduncle to 10 

cm long; rachis up to 20 cm long, sometimes arching, more commonly straight, erect; 

rachis bracts up to 2 mm long, broadly acuminate; rachillae distichous, opposite, 

becoming sub-opposite distally, 7-12 on each side, the lowermost c.7 cm long, 

decreasing distally, adnate to the inflorescence for 8 mm, less so distally, arching 

vertically or straight. Flowers borne in close sub-distichous pairs with

Figure 13. Eremospatha haullevilleana De Wild. 

Bidgood et al., 2924; a. Stem x 1 ½ (1), b. Leaflets x 1 (2/3), c. Juvenile leaf ½ (1/3): Evrard 5511; d. 

Flower x 3 (2): Louis 9560; e. Fruit x 1 ½ (1), f. Seed x 1 ½ (1). Drawn by Lucy T. Smith. 

82

Distribution 

E. haullevilleana is restricted to the lowland forests of the Congo Basin. Unlike the 

majority of the rattan species, it is curiously absent from the coastal forest regions. 

Figure 14. Distribution of E. haullevilleana De Wild. 


E. haullevilleana is found both in closed-canopy forest and in open areas. In common 

with E. macrocarpa, it is a species of terra firme forest and is not associated with 

swamp vegetation. 


CAMEROON: Letouzey 11798, Mintom I (02.03N:13.30E) sterile, January 5, 1973 (YA!); 

CENTRAL AFRICAN REPUBLIC: Carroll 115, 35km NE of Bayanga (03.07N:16.27E) sterile, 

March 13, 1985 (MO!); Harris 3508, 20km SE of Bayanga (02.50N:16.22E) sterile, October 10. 1993 

(K!); Harris 2652, Lidjombo, E of Sangha River (02.39N:16.11E) Fr., November 2, 1990 (K!); 

GABON: Wieringa 1550, 120km on road from Okunju to Makokou (00.08N:13.41E) Fr., September 1, 

1992 (WAG!); CONGO: Harris et al. 3172, 55km SW of Souanké (02.02N:13.49E) sterile, November 

11, 1991 (K!, MO!); Lejoly 96/750, Parc National d'Odzala (00.36N:14.54E) Fl., November 21, 1996 

(BR!); Lerot s.n., Ogoué, sterile, 1894 (K!); DR CONGO: Allard 213, sterile, 1909 (BR!); Apema 217, 

83

Masako, sterile, February 1, 1987 (BR!); Bavicchi 277, Lukabo, seedling, 1913 (BR!); Bequaert 1277, 

Yambuya (01.15N:24.33E) seedling, November 24, 1913 (BR!); Bequaert 1411, Bamalia, Fr., 

December 9, 1913 (BR!); Billiet & Jadin 4054, Bangu Massif (06.41S:19.22E) Fr., February 4, 1987 

(BR!); Callens s.n., Kisantu (04.25S:14.42E) sterile, March 1947 (BR!); Claessens 381, Kalako Kombe 

(00.45S:21.33E) Fr., January 1910 (BR!); Compère 2181, Mputu (04.46S:15.31E) seedling, July 9, 

1960 (BR!); Couteaux 471, Eala (00.03N:18.18E) Fr., October 20. 1908 (BR!); Couteaux 1051, 

Leopoldville (04.22S:15.23E) sterile, July 21, 1944 (BR!); de Graer 297, Mongoli River, sterile, 

September 1, 1934 (BR!); de Graer 327, Doruma (04.43N:27.42E) sterile, October 26, 1924 (BR!); 

Dewevre 581, Fr., s.d. (BR!); Dewevre 986, Fimbo, sterile, s.d. (BR!); Evrard 4041, Ikelemba river, 

Befale (00.26N: 20.48E) sterile, May 7, 1958 (K!, BR!); Evrard 5511, Yalikungu (00.42N:22.35E) Fl., 

January 10, 1959 (K!, BR!); Evrard 5890, Equateur Province, Bomongo (01.22N:18.23E) Fr., March 

11, 1959 (K!); Flamigny 6361, Fr., February 1943 (BR!); Gathy 1639, Kaniama (06.05S:22.20E) Fl., 

August 21, 1958 (BR!); Gerard 1432, Diagbe (04.19N:27.45E) Fr., July 24, 1954 (K!, BR!); Gilbert 

2258, 100km from Abo (03.14N:30.10E) sterile, May 1939 (BR!); Gillet 1385, Kisantu 

(04.25S:14.42E) juvenile, 1900 (BR!); Gillet 2026, Lukamba ( 05.20S:19.14E) Fl. & Fr., 1903 (FI!, 

BR!); Gillet 3505, sterile, 1903 (BR!); Gutzwiller 539, Yangambi (00.45N:24.26) seedling, February 

10, 1955 (BR!); Hendrickx 4157, Hombo (01.52S:28.27E) sterile, August 1946 (BR!); Herman 2138, 

Kaniama, Haut Lomami (06.05S:22.20E) Fl., March 1937 (BR!, WAG!); Hulstaert 1416, Bokela 

(01.07S:21.55E) seedling, s.d. (BR!); Hulstaert 1418, Bokela (01.07S:21.55E) Fr., s.d. (K!, BR!); 

Hulstaert 1421, Bokela (01.07S:21.55E) seedling, s.d. (BR!); Hulstaert 1614, Bokuma, seedling, July 

16, 1953 (BR!); Hulstaert 1616, Bokuma, sterile, July 16, 1953 (BR!); Kitembo 60, Kalima, Kivu 

(02.31S:26.26E) sterile, February 1981 (BR!); Laurent 911, Eala (00.03N:18.18E) sterile, July 1, 1903 

(BR!); Laurent s.n., Eala (00.03N:18.18E) Fl., July 15, 1903 (BR!); Laurent s.n., Eala (00.03N:18.18E) 

Fl., 1905 (K!, FHO!); Laurent s.n., Isaka (01.28S:23.50E) sterile, November 21, 1903 (BR!); Laurent 

s.n., Kasai (10.07S:22.20E) sterile, November 1895 (BR!); Lebrun 1508, Eala (00.03N:18.18E) Fl., 

December 1931 (BR!); Leclercq 736, Doruma (04.43N:27.42E) sterile, March 7, 1958 (BR!); Lejoly 

566, Bawombi (00.42N:26.11E) seedling, November 17, 1976 (BR!); Leonard 932, Eala 

(00.03N:18.18E) Fl. & Fr., October 26, 1946 (K!, BR!, MO!); Leonard 933, Eala (00.03N:18.18E) Fl., 

October 26, 1946 (K!, BR!, WAG!); Leonard 936, Eala (00.03N:18.18E) sterile, October 30, 1946 (K!, 

BR!, WAG!); Leonard 1138, Oriental Province, Weko (01.13N:24.07E) Fr., March 1947 (K!, BR!, 

MO!); Liben 2603, Luisa Territory, Tomba (00.10N:19.18E) Fr., February 1957 (K!, BR!, WAG!); 

Liegeois 87, Oriental Province, Tshopo (10.12S:24.51E) sterile, July 1943 (BR!); Liegeois 87, Oriental 

Province, Tshopo (10.13S:24.50E) sterile, July, 1943 (K!); Lisowski 7160, Kouilou, Bena 

(02.34S:11.27E) sterile, October 8, 1990 (BR!); Louis 772, Yangambi (00.45N:24.26E) Fl., December 

5, 1935 (BR!); Louis 847, Yangambi (00.45N:24.26E) Fl., December 17, 1935 (BR!); Louis 1970, Eala 

(00.03N:18.18E) Fl., May 30, 1936 (K!, BR!); Louis 3395, 6km W of Yangambi (00.45N:24.26E) Fr., 

March 3, 1937 (K!, BR!); Louis 3638, 25km NW of Yangambi (00.49N:24.12E) Fl., March 19, 1937 

(BR!, WAG!); Louis 4218, Yangambi (00.45N:24.26E) seedling, June 22, 1937 (BR!); Louis 7671, 

Yambuya (01.15N:24.33E) Fr., January 24, 1938 (K!, BR!); Louis 8106, Yangambi (00.45N:24.26E) 

seedling, February 25, 1938 (BR!); Louis 9420, Yangambi (00.45N:24.26E) Fr., May 18, 1938 (K!, 

84

FHO!, BR!); Louis 9560, 20km W of Yangambi (00.49N:24.12E) Fr., May 30, 1938 (K!, BR!); Louis 

9731, Yalibwa, river Lubuye (00.56N:24.30E) Fl., June 9, 1938 (BR!); Louis 11850, Yangambi 

(00.45N:24.26E) Fl. & Fr., October 18, 1938 (BR!, MO!, WAG); Louis 12106, 20km W of Yangambi 

(00.49N:24.12E) Fr., October 27, 1938 (K!, BR!); Louis 16775, Yangambi (00.45N:24.26E) Fr., 

November 17, 1943 (BR!); Louis 10045, Oriental Province, Yalibora, Fr., June 1938 (K!, BR!); Louis 

218, Oriental Province, Yangambi (00.45N:24.26E) seedling, June 1937 (K!, FHO!, MO!); Luja 107, 

Gungu (05.45S:19.19E) seedling, November 18, 1898 (BR!); Luja 221, Kasai (10.07S:22.20E) sterile, 

March 11, 1899 (BR!); Luja 288, Lubué (04.08S:19.49E) juvenile, June 16, 1899 (BR!); Luja 297, 

Lubué (04.08S:19.49E) sterile, June 26, 1899 (BR!); Masens 451, Kikwit (05.02S:18.48E) juvenile, 

November, 21, 1990 (BR!); Mullenders 1166, Kaniama, Haut Lomami (06.05S:22.20E) sterile, 

September 1947 (BR!); Nannan 117, Bamania, Fr., August 29, 1914 (BR!); Nsola 621, Bikoro 

(00.44N:18.09E) Fr., May 31, 1984 (BR!); Pynaert 1676, Eala (00.03N:18.18E) Fr., 1907 (BR!); 

Robyns 4300, Kimengua, sterile, February 7, 1957 (BR!); Sapin s.n., Demba (04.27S:23.45E) Fl., 

December 1910 (BR!); Sapin s.n., Tshibangu (05.56S:20.54E) sterile, 1910 (FHO!); Schmitz 3710, 

Kanzenze (10.27S:27.11E) sterile, August 26, 1950 (BR!); Schouten 103, sterile, August 23, 1910 

(BR!); Terashima 94, Nyamakombola (01.41S:28.09E) sterile, October 20, 1989 (BR!); Thiebaud 321, 

Katobo (00.57S:28.58E) Fl., January 1959 (BR!); Thonet 110, Lac Tumba (00.46S:20.06E) Fl., October 

30, 1957 (K!, WAG!, BR!); Troupin 2658, Parc National de Garamba (04.10N:29.28E) sterile, May 21, 

1952 (BR!); Troupin 9162, Kavumu-Walikele (01.28S:28.48E) Fl., September 11, 1958 (BR!); 

Vandenbrand 238, sterile, s.d. (BR!); Vanderyst 2781, Kikwit (05.02S:18.48E) sterile, January 1914 

(BR!); Vanderyst 4906, Dumu (03.20N:18.18E) sterile, August 1914 (BR!); Vanderyst 12664, Fl., 

November 1922 (BR!); Vanderyst 30729, Wula to Kipako, juvenile (May 1932) BR!; ANGOLA: 

Gossweiler 10087, Mayombe, Luali (05.00S:12.25E) Fl., 1923 (K!); Gossweiler 6645, Mayombe, Luali 

(05.00S:12.25E) Fl., 1923 (K!); Gossweiler 7844, Mayombe, Luali (05.00S:12.25E) sterile, 1919 (K!); 

Gossweiler 8129, Mayombe, Luali (05.00S:12.25E) sterile, January 5, 1919 (K!); UGANDA: Dawe 

668, Semliki (00.45N:30.00E) sterile, October 31, 1905 (K!); Makombo et al. s.n., Semliki 

(00.45N:30.00E) sterile, October 24, 1998 (K!); BURUNDI: Lewalle 4016, Kigwena (04.10S:29.32E) 

sterile, November 9, 1969 (BR!); Lewalle 4414, Kigwena (04.10S:29.32E) Fl., February 1, 1970 (BR!); 

Reekmans 11180, Kigwena (04.10N:29.30E) Fr., May 13, 1982 (K!, MO!, BR!); TANZANIA: 

Bidgood & Vollesen 3040, Kigoma to Kasulu road (04.33S:29.52E) sterile, April 1, 1994 (K!); Bidgood 

et al. 2924, Kigoma: Kasye forest (04.47S:29.40E) sterile, March 23, 1994 (K!); Eggeling 6207, 30 

miles S of Kibondo (03.34S:30.46E) sterile, July 1951 (K!); Proctor 470, Western Province, Mpanda 

(06.22S:31.03E) Fl., May 1956 (K!); Proctor 369, Western Province, Mbuti River (04.53S:38.29E) 

sterile, February 1955 (K!) 

______________________________________ 

85

E. tessmanniana Becc. 

Günther Tessmann (born ? - 1926) German botanist and anthropologist 

Becc. in Webbia 3: 278 (1910); Guinea-Lopez in Ensayo Geobot. de la Guinea Cont. 

Espanola 245: (1946); Letouzey in Adan. 18(3): 314 (1978); Aedo et al. in Bases 

Docs. Fl. de Guinea Ecuatorial 375: (1999). Type: Equatorial Guinea, Tessmann 4 

(holotype B†; isotype FI!). 

Clustering slender palm climbing up to 150 m, more commonly to 60-80 m. Stems, 

often branching, circular in cross-section, without sheaths 10-12 mm in diameter, with 

12-15 mm in diameter; internodes 15-20 cm long. Leaf sheath longitudinally striate, 

with black caducous indumentum; ocrea entire, horizontally truncate, extending to 1.5 

cm; knee absent. Juvenile stems with sheath, 0.6 cm in diameter; petiole angular, 15- 

17 cm long armed along the margins with reflexed, bulbous-based, black tipped 

spines; leaves bifid, 20 cm x 24 cm, deeply notched, with somewhat rounded lobes; 

elaminate rachis present on lower section of stems, up to 80 cm long. Leaves on 

mature stems sessile, or very nearly so (petiole

order region of Cameroon and the Rio Muni territory of Equatorial Guinea. Further 

collections might link this disjunction. 

Figure 15. Distribution of E. tessmanniana Becc. 


E. tessmanniana is a forest species found on well-drained soils in closed-canopy 

forest. 

Notes 

In common with E. quinquecostulata, E. tessmanniana has not been recognised as a 

distinct species since Beccari’s original description (Beccari, 1910). Tuley (1995) 

suggested this species represented a juvenile form of an unspecified species, however, 

recent studies of herbarium material and subsequent field collections have proved this 

not to be the case. Fertile material of this species is needed for a more complete 

description to be made. 

87


CAMEROON: Mildbraed 5285, sterile, May 20, 1911 (HBG!); Mildbraed 5879, Ebolowa 

(02.55N:11.08E) sterile, s.d. (HBG!); Sunderland 2021, Takamanda Forest Reserve (06.08N:09.16E) 

sterile, January 11, 1999 (K!, SCA!); Sunderland 2017, Takamanda Forest Reserve (06.08N:09.16E) 

sterile, January 17, 1999 K!, SCA!; EQUATORIAL GUINEA: Tessmann 4, sterile, s.d. (FI!) 

______________________________________ 

E. cuspidata (G. Mann & H. Wendl.) H. Wendl. 

(Latin) refers to finely apiculate leaflet apex 

H. Wendl. in Kerchove Les Palmiers 244: (1878); C.H. Wright in F.T.A. 8: 112 

(1901); Durand & Durand in Fl. Congolanae 1: 585 (1909); Becc. in Webbia 3: 275 

(1910); Pyneart in Bull. Agric. du Congo Belge 2: 547 (1911); Hédin in Rev. de Bot. 

Appl. 9: 504 (1929); Guinea-Lopez in Ensayo Geobot. de la Guinea Cont. Espanola 

245: (1946); Letouzey in Adan. 18(3): 314 (1978); Tuley in Palms of Africa 51: 

(1995). 

Calamus (Eremospatha) cuspidatus G. Mann & H. Wendl. in Trans. Linn. Soc. 24: 

434 (1864); Type: Gabon, Ogooué River, Mann 1043 (holotype K!). 

Clustering slender palm climbing to 12-15m. Stems circular in cross-section, without 

sheaths, 10-15 mm in diameter, with 16-25 mm; internodes 11-15 cm long. Leaf 

sheath longitudinally striate sparsely covered with brown-black indumentum; ocrea 

obliquely truncate, extending c.1 cm above the rachis; knee absent. Leaves sessile, up 

to 2 m long; rachis 1-1.3 m long, flattened on upper surface, rounded below, becoming 

trapezoid then rounded in cross section distally, armed along the margins with 

inequidistant, reflexed, bulbous-based spines, becoming sparsely armed distally, 

indumentum absent; cirrus 50-75 cm long, unarmed; leaflets up to 15-20 on each side 

of the rachis, linear-lanceolate, abruptly contracted at the base, with a fine 0.8-1.2 cm 

long (rarely 3 cm long) apiculum at the apex, 22-30 cm long x 1.6-2 (rarely 3 cm) 

broad at the widest point, discolorous, adaxially mid-green, abaxially light green, 

armed along the margins with inequidistant abrupt black-tipped spines, 5-7 

moderately conspicuous transverse veinlets 2-3 mm apart; lowermost leaflets smaller 

than the rest, linear-ovate, reflexed and laxly swept back across stem; acanthophylls in 

88

pairs c.3 cm long, at 45° angle to cirrus. Inflorescence glabrous, 30-38 cm long, rarely 


E. cuspidata is highly unusual amongst the rattans of African in that it is commonly 

found in the deep white sand savannah areas or “praderas”, characteristic of the 

coastal forests of the Congo Basin where it forms dense, scrambling thickets. 

However, in some areas, E. cuspidata has also been encountered in gap vegetation in 

forest. 

90

Figure 17. Eremospatha cuspidata (G. Mann & H. Wendl.) H. Wendl. 

Sunderland 1909; a. Stem x 1, b. Leaflets x 2/3 (4/9), c. Leaflet apex x 1 ½ (1): Sunderland 1792; d. 

Acanthophylls x 1 (2/3): Sunderland 1922; e. Flower x 4 (2 2/3), f. Flower section x 4 (2 2/3): 

Sunderland 1909; g. Fruit and infructescence. Drawn by Lucy T. Smith. 

91


CAMEROON: Bruneau 1074, Mt Kupe, SW Province (04.48N:09.42E) sterile, October 26, 1995 (K!); 

Mildbraed 9546, Buea – Douala, sterile, June 5, 1914 (K!); CENTRAL AFRICAN REPUBLIC: Fay 

4021, Manovo-St Floris National Park (09.29N:21.17E) sterile, December 30, 1982 (K!); 

EQUATORIAL GUINEA: Eneme & Lejoly 113, Ndote Reserve (01.20N: 09.28E) Fl., August 19, 

1987 (EG!); Sunderland 1792, near village of Etembue (01.16N:09.26E) Fr., March 13, 1997 (K!, EG!, 

BH!); Sunderland 1909, near village of Etembue (01.16N:09.26E) Fr., March 28, 1998 (K!, EG!, 

WAG!); Sunderland 1922, near village of Etembue (01.16N:09.26E) Fl., April 8, 1998 (K!, EG!, 

WAG!); GABON: Breteler & van Raalte 5557, SE of Port Gentil (00.40S:08.50E) Fl., November 16, 

1968 (MO!); Dybowski 140, Fl. (FI!); Mann 1043, Gaboon River (00.19N:09.29E) Fl. & Fr., July 1861 

(K!); Reitsma 2889, 20km N of Libreville (00.35N:09.22E) Fr., January 29, 1987 (WAG!, LBR!); DR 

CONGO: Desenfous 2023, Kaniama, Haut Lomami (06.05S:22.20E) Fl., September 1951 (BR!); 

Malaisse 14159, Mushindi River (09.30S:23.13E) sterile, February 19, 1987 (BR!); Schmitz 5617, 

66km from Kinda (09.17S:25.03E) Fr., September 20, 1957 (BR!); ANGOLA: Milne-Redhead 4219, 

River Monu, sterile, 1937 (K!); ZAMBIA: Loverage 931, Mwinilunga District (11.44S:24.26E) sterile, 

June 12, 1963 (K!); Mutimushi 3372, Mwinilunga District (11.44S:24.26E) sterile, May 19, 1969 (K!) 

______________________________________ 

E. quinquecostulata Becc. 

(Latin) “five main veins” 

Becc. in Webbia 3: 279 (1910); Tuley in Palms of Africa 123: (1995); Type: 

Cameroon, Dja, unknown collector (holotype FI!) 

Clustering slender palm climbing to 10-15 m. Stems ± circular in cross-section, 

without sheaths 4-9 mm, with 5-10 mm; internodes 14-16 cm long. Leaf sheath 

longitudinally striate, with sparse brown caducous indumentum; ocrea entire, 

obliquely truncate, extending to 1.2-2.7 cm; knee somewhat inconspicuous, ridge-like, 

vertically linear, tapering at base, extending to 4 cm. Leaves sessile, or with petiole up 

to 10 cm long, somewhat flattened, armed along the margins with inequidistant, 

reflexed, bulbous-based, black-tipped spines; rachis 60-80 cm long, flattened in crosssection 

proximally, becoming triangular in cross-section distally, armed as the petiole; 

cirrus up to 40-50 cm long, very fine, armed as the rachis although spines becoming 

sparse distally; leaflets, 5-12 on each side, inequidistant, grouped in pairs or in 4’s, 

somewhat irregularly clustered, lanceolate to loosely rhomboid, unequally attenuate at 

92

ase, entire and acuminate to irregularly praemorse at apex, 13-20 cm long x 2-3 cm 

broad at the widest point, discolorous, adaxially dark green, abaxially mid-green, 

margin unarmed or sparsely armed with somewhat straight, forward-facing, blacktipped 

spines, praemorse apex ciliate-spiny; with 5 conspicuous, equal, main veins; 

lowermost leaflets smaller than the rest, although not obviously so; acanthophylls, 

somewhat slender, up to 2 cm long, at 45° angle to cirrus. Flower and fruit unknown. 

Distribution 

This species is known only from Cameroon and SE Nigeria. 

Figure 18. Distribution of E. quinquecostulata Becc. 


E. quinquecostulata is a slender rattan found only in high forest. 

Notes 

This species is relatively uncommon and, along with E. tessmanniana, has not been 

recognised as a distinct species since Beccari’s original account (Beccari, 1910). 

93

Tuley (1995) again suggested that this species represented “a juvenile form of 

unknown provenance”, however recent field work and study of herbarium collections 

have confirmed that this is indeed a distinct species and that Beccari’s original 

recognition of E. quinquecostulata as a distinct species is valid. The collection of 

fertile material would enable a more complete description to be made. 


NIGERIA: Tuley 653, Calabar to mamfe road: Mile 25 (05.18N:08.34E) sterile, July 13, 1964 

(WAG!); CAMEROON: unknown collector, Bezirk Djah (03.00N:12.40E) sterile (FI!); Letouzey 

4285, 40km S of Mesamena (03.28N:12.50E) sterile, February 17, 1962 (YA!); Sunderland 1741, 

Rumpi Hills Forest Reserve (04.54N:09.20E) sterile, May 19, 1996 (K!, SCA!, NY!); Sunderland 1806, 

Campo Ma'an Faunal Reserve (02.10N:09.54E) sterile, April 8, 1997 (K!, YA!); Sunderland 1938, 

Takamanda Forest Reserve (06.06N:09.47E) sterile, November 18, 1998 (K!, SCA!); Sunderland 2054, 

Takamanda Forest Reserve (06.08N:09.16E) sterile, January 10, 1999 (K!, SCA!) 

94

Figure 19. Eremospatha tessmanniana Becc. 

Sunderland 2021; a. Stem x 1 (2/3), b. Leaflets and cirrus x 2/3 (4/9) 

Eremospatha quinquecostulata Becc. 

Sunderland 1938; c. Stem x 1 (2/3), d. Leaflets x 2/3 (4/9), e. Leaflets & cirrus x 1 (2/3) 

Drawn by Lucy T. Smith. 

95

Indet. Eremospatha 

SIERRA LEONE: Jaeger 9222, between Krutown & Siki Koro, Fr. February 4, 1966 (MO!); CÔTE 

D'IVOIRE: Leeuwenberg 2515, 61km N of Sassandra (06.10N: 05.19W) sterile, January 20, 1959 

(WAG!); LIBERIA: Adam 29807, Yekepa (07.34N:08.32W) sterile, October 6, 1975 (MO!); 

NIGERIA: Ayewoh 3851, Ondo Province, Okeluse Reserve (06.43N:05.30E) juvenile, February 23, 

1944 (K!); Imp. Inst. Nigeria 346, Ahoada area (05.03N:06.34E) Fl. only, February 1936 (K!); 

CAMEROON: Bruneau 1071, Mount Kupe (04.48N:09.42E) juvenile, October 25, 1995 (K!); 

Letouzey 3673, NW of Eschienbot nr Abong-Mbang (03.59N:13.11E) juvenile, April 1, 1964 (YA!); 

Letouzey 4206, Nkomo nr Ngoase, S of river Loba (02.32N:11.49E) juvenile (YA!); Letouzey 11133, 

Bafang - Yabassi, 12km NNE of Nkondjock (03.51N:10.33E) sterile, February 6, 1972 (YA!); Letouzey 

11778, Mintom II (02.03N:13.30E) seedling, January 3, 1973 (YA!); Letouzey 11794, Alat Mahay, 

NNE of Mintom II (02.03N:13.30E) sterile (YA!); Sunderland 2301, Korup National Park, Chimpanzee 

Camp (05.02N:08.48E) sterile, February 15, 2000 (K!, SCA!); Sunderland 1738, Rumpi Hills Forest 

Reserve (04.54N:09.20E) seedling, May 19, 1996 (K!, SCA!); Sunderland 1933, Takamanda Forest 

Reserve (06.06N:09.47E) seedling, November 14, 1998 (K!, SCA!); Thomas s.n., Korup National Park 

(04.55N:08.50E) sterile, s.d. (SCA!); Webb 311, Campo Faunal Reserve (02.24N:09.54E) seedling, 

July 10, 1976 (K!); CENTRAL AFRICAN REPUBLIC: Fay 4036, Manovo-St Floris National Park 

(09.29N:21.17E) seedling, December 31, 1982 (K!); Fay & Harris 8795, Dzanga-Sanga 

(02.21N:16.11E) sterile, October 20, 1988 (MO!); GABON: le Testu s.n., Haute-Ngounye 

(00.22S:10.27E) sterile, s.d. (BR!); le Testu s.n., Haute-Ngounye (00.22S:10.27E) seedling, s.d. (BR!); 

CONGO: Harris et al, 3222, base of Mont Nabemba (01.50N:13.58E) sterile, November 20, 1991 

(MO!); DR CONGO: Camp s.n., sterile, s.d. (BR!); Dechamps s.n., Les Sara (04.22N:12.22E) 

juvenile, June 30, 1989 (WAG!); Gillet s.n., juvenile, 1903 (BR!); Gossweiler 8705, Sumba 

(05.37S:14.03E) Fr. only, December 1921 (K!); Hendrickx s.n., seedling, 1903 (BR!); Hulstaert 1428, 

Bokela (01.07S:21.55E) Fr. only, s.d. (BR!); Luja 231, Kasai (10.07S:22.20E) sterile, March 18, 1899 

(BR!); Michel 2957, Mosso, seedling, June 19, 1952 (BR!); Ringoet s.n., seedling, 1923 (BR!); Sapin 

s.n., sterile, s.d. (BR!); Vanderyst 5256, seedling, August 1915 (BR!); Vanderyst 11246, Fl. only, 

November 1921 (BR!); Vanderyst 12670, Fl. only, November 1922 (BR!); BURUNDI: Michel & Reed 

1362, Kiofi-Muzye (04.00S:30.06E) seedling, September 15, 1952 (BR!) 

______________________________________ 

Eremospatha: excluded names and nomina nuda 

E. longehamata Dammer nomen in herb. Berilo. = E. cabrae 

E. trapezoidea Dammer nomen in herb. Berilo. = E. cabrae 

E. schweinfurthii Becc. in Ann. Roy. Bot. Gard. Calc. 11(1): 164(1908), nom. = E. 

haullevilleana De Wild. 

E. lujae Dammer nomen in Herb. Berilo. = E. haullevilleana De Wild. 

E. wildemannii Dammer nomen in Herb. Berilo. = E. haullevilleana De Wild. 

96

LACCOSPERMA (G. Mann & H. Wendl.) Drude 

(Latin) “hole in the seed” 

Drude in Bot. Zeit. 35: 635 (1877); Type: Equatorial Guinea, Bioko, L. opacum (G. 

Mann & H. Wendl.) Drude (Calamus opacus G. Mann & H. Wendl.). 

Calamus subgenus Laccosperma G. Mann & H. Wendl. in Trans. Linn. Soc. (London) 

24: 430 (1864). 

Ancistrophyllum (G. Mann & H. Wendl.) H. Wendl. (non Göppert 1841) in Kerchove, 

Les Palmiers 230 (1878); Type: A. secundiflorum (P. Beauv.) H. Wendl. 

(Calamus secundiflorus P. Beauv.) (= Laccosperma secundiflorum (P. Beauv.) 

Küntze). 

Ancistrophyllum subgenus Laccosperma (G. Mann & H. Wendl.) Hook.f. in Benth. & 

Hook., Gen. Pl. 3: 937 (1883). 

Ancistrophyllum subgenus Ancistrophyllum Hook.f. in Benth. & Hook., Genera 

Plantarum 3:937 (1883). 

Neoancistrophyllum Rauschert, Taxon 31:557 (1982); Type: Neoancistrophyllum 

secundiflorum (P. Beauv.) Rauschert (superfluous substitute name - fide Dransfield, 

1982). 

Clustered, spiny, understorey to high climbing, hapaxanthic, hermaphroditic rattan 

palms. Stems, circular in cross section, rarely oval, with medium to long internodes; 

sucker shoots axillary. Leaf pinnate, with cirrus; sheath strictly tubular, sparsely to 

profusely armed with fine scattered spines, sometimes becoming bare; ocrea 

conspicuous, split opposite the petiole, scarcely sheathing, rarely inflated with inrolled 

edges or reflexed and tattering, armed as the sheath; knee absent; petiole short to long, 

much shorter in the reduced leaves subtending the inflorescences, usually armed with 

inequidistant angular spines along margins, never unarmed; rachis armed as the 

petiole; cirrus armed with reflexed prickle-like spines and bearing neat pairs of 

reflexed acanthophylls; leaflets few to very numerous, 1-4 fold, entire, linear to 

97

sigmoid, regularly or irregularly arranged, often fiercely armed with short to long 

spines along the margins and the main ribs, midribs prominent adaxially, transverse 

veinlets conspicuous or inconspicuous; acanthophylls alternate proximally, subopposite 

to opposite distally. Inflorescences produced simultaneously in the axils of 

the most distal few, frequently reduced leaves, branched to 1-order; peduncle enclosed 

within the leaf sheath and emerging from its mouth, ± hemispherical in cross section; 

prophyll strictly tubular, 2-keeled, enclosed within the subtending leaf sheath; 

peduncular bracts 1-3; rachis longer than the peduncle; rachis bract distichous, strictly 

tubular with a triangular limb, without spines, sparsely to profusely indumentose, 

becoming tattered at apex, each subtending a pendulous or spreading rachilla; rachilla 

prophyll tubular, 2-keeled, included within the subtending bract, rachilla bracts 

distichous, tubular with apiculate triangular limb, striate, sparsely indumentose, the 

margin sometime ciliate, each, except for the basal 1-2, subtending a flower cluster. 

Flowers usually in dyads, rarely in triads, sometimes solitary towards the tip of the 

rachillae, the flower cluster bearing a 2-keeled prophyll and 0, 1 or 2, 2-keeled 

bracteoles (depending on the number of flowers); calyx slightly to strongly stalk-like 

at the base, often bent at right angles, incompletely divided distally into 3- triangular 

striate lobes; corolla tubular at the very base, divided above into 3 oblong , narrow, 

triangular, valvate lobes; stamens 6, epipetalous, filaments distinct, much swollen, 

angular, scarcely narrowed at the connective; anthers medifixed, oblong, latrorse, 

pollen elliptic, monosulcate with finely reticulate, tectate exine; gynoecium 

tricarpellate, triovulate, ovary covered with scales, those at the base of the style 

minute, spine-like, style elongate, 3-angled, stigma minute, pyramidal, ovules basally 

attached, anatropous. Fruit baccate, 1- sometimes 2-seeded, tipped with the base of the 

style, the remainder of the style usually breaking off in early fruit development, calyx 

and corolla persistent at base; epicarp covered in vertical rows of reflexed scales with 

fringed margins, mesocarp white, fleshy and sweet at maturity, endocarp not 

differentiated. Seed attached sub-basally at one side, ovoid and laterally flattened, or 

rounded and deeply scalloped, with a very shallow to deep, lateral pit, seed coat 

fleshy, endosperm homogenous; embryo lateral, opposite the depression or pit. 

Germination adjacent-ligular; eophyll bifid. 

98

Habitat and distribution 

The genus Laccosperma is represented by five (plus one imperfectly known) species 

which occur throughout the lowland forest region of West and Central Africa. The 

species are either shade tolerant and are present as slender climbers under the forest 

canopy, whilst other species are strongly light demanding and are a common 

component of gaps and forest margins. 

Notes 

The provision of a satisfactory taxonomy of this genus has been hindered by a paucity 

of adequate fertile material and inadequate field observations by those working on the 

group. Despite a number of species being described, only very few species names 

have been applied. Most notable is the fact that the name L. secundiflorum has been 

far too widely applied to include what are now known to be closely-related species 

and there are in fact three morphologically distinct species of large-diameter 

Laccosperma. This is an understandable mistake to make, especially as the flowers 

and fruits of all these species show a remarkable similarity, although there are some 

morphological differences (fide Beccari, 1910). In the field, however, as these species 

are often sympatric, they are easily distinguished from each other. 

Key to the species of Laccosperma 

Slender canes, stems with sheaths ±2 cm in diameter, with 10-12 leaflets on each side 

of the rachis; common in forest understorey: 

Leaflet margin armed with forward-facing truncate spines, fruit globose; seed 

sub-globose, covered in polygonal rounded depressions, deeply scalloped on 

one side: L. opacum 

Leaflet margin unarmed, fruit ovoid; seed ovoid, flattened, with a linear 

depression on one side: L. laeve 

Robust canes, stems with sheaths ±2 cm in diameter, with >12 leaflets on each side of 

the rachis; common in forest gaps and open areas: 

99

Stem with sheaths ±3.5 cm in diameter, petiole on mature stems 20 cm long 

petiole, leaflets, sigmoid, elongate, horizontally held on rachis: L. 

secundiflorum 

_______________________________________ 

Laccosperma opacum (G. Mann & H. Wendl.) Drude 

(Latin) “darkened” or “dull”; refers to dark green leaflets 

Drude in Bot Zeit. 35: 635 (1877); Küntze in Rev. Gen. Plant. 2: 729 (1891); Mildbr. 

in Deutsch. Zentr. Afr. Exped. 54: (1914); J. Dransf. in Kew Bull. 37(3): 456 (1981); 

J. Dransf. in F.T.E.A. (Palmae) 36: (1982); Hawthorn in Trees of Ghana 225: (1990); 

Morakinyo in Principes 39(4): 206 (1994); Tuley in Palms of Africa 37: (1995); H.A. 

Burkill in Useful Pl. of W. Trop. Afri. 4:373 (1997); Cable & M. Cheek in Pl. of Mt. 

Cam. 179: (1998). 

Ancistrophyllum opacum (G. Mann & H. Wendl.) Drude in Engl. Bot. Jarbh. 21: 111 

(1895); Cummins in Kew Bull. 137: 80 (1898); C.H. Wright in F.T.A. 7: 116 (1902); 

Becc. in Webbia 3: 257 (1910); Mildbr. in Notizbl. Bot. Gar. Dah. App. 27: 16 (1913); 

Baudon in Rev. de Bot. Appl. 4: 595 (1924); Hédin in Rev. de Bot. Appl. 9: 504 

100

(1929); Hutch. in F.W.TA. 2: 391 (1936); Dalziel in App. to F.T.W.A. 495: (1937); 

Guinea-Lopez in Ensayo Geobot. de la Guinea Cont. Espanola 244: (1942); Fosberg 

in Principes 4: 129 (1960); Irvine in Woody Plants of Ghana 773: (1961); Walker & 

Sillans in Plantes Utiles du Gabon 324: (1961); T.A. Russell in F.W.T.A 3(2): 167 

(1968); H.E. Moore in Principes 15: 112 (1971); Letouzey in Adan. 2(18): 314 (1978); 

Hall & Swaine in For. Veg. Ghana 123: (1981); Aedo et al. Bases docs. Fl. de Guinea 


Calamus (subgen. Laccosperma) opacus G. Mann & H. Wendl. in Trans. Linn. Soc. 

24: 431 (1864); Type: Equatorial Guinea, Bioko, Mann 97 (holotype K!). 

Clustering slender palm climbing to 10-15 m. Stem, often branching, without sheaths 

up to 15 mm in diameter, with c.20 mm; internodes 10-20 cm long. Leaf sheath 

moderately to sparsely armed with black tipped, pale coloured bulbous-based, upward 

pointing or reflexed spines; sheaths near inflorescence occasionally very sparsely 

armed; black caducous indumentum present on mature sheaths; ocrea 12-30 cm long, 

gradually tapering at the apex, papyraceous and tattering, pale straw-coloured without, 

dark shiny brown within, armed with black-tipped spines with pale bulbous bases, 

particularly at apex. Leaves up to 1.2 m long; petiole to 12 cm long x 0.8 cm wide, 

abaxially rounded, adaxially flattened, armed along the edges with up to 1 cm-long, 

inequidistant, black spines with pale bulbous bases, spreading or reflexed; rachis to 60 

cm long, rounded or somewhat angular in cross section, armed as the petiole, the 

spines decreasing in size distally; cirrus to 50 cm long armed as the rachis, although 

spines becoming sparse distally; leaflets composed of 2-4s fold, 10-12 on each side of 

the rachis, inequidistant and unequal in size, usually sub-equidistant proximally and 

borne in pairs distally, sigmoid, 1-5-costulate, the largest 20-30 cm long x 2.5-10 cm 

broad at widest point, ± concolorous, with prominent transverse veinlets and numerous, 

rather distant, small spines on the margins; acanthophylls to 2.5 cm long. 

Inflorescences, numbering 4-8, produced simultaneously in distal 30-50 cm of stem; 

peduncle c.10 cm long; prophyll c.4 cm; lower bracts up to 2 cm long, gradually 

decreasing distally, with a short triangular acuminate lobe, closely adpressed to the 

next bract, finely striate; rachillae to 10 cm long, spreading, densely clothed with 

striate imbricate bracts + 4 mm distant, each with a rather wide mouth and short 

apiculum to1 mm; bracteoles minute. Flowers 8 mm long x 2 mm wide; calyx, tan- 

101

cream, 4 mm long, tubular in the basal 2 mm, with 3 triangular striate lobes 2 mm 

long x 2 mm wide; petals cream or white, 9 mm long x 2.5 mm wide, lanceolate with 

a blunt triangular tip; stamen-filaments to 3 mm long x 1 mm wide, minutely 

epipetalous and united into a very short (


Tolerant of deep shade, L. opacum is commonly found in high forest in the lower to 

mid-canopy. This species prefers well-drained soil and is the only species of rattan 

found on basalt and other volcanic soil types. 


GUINEA-CONAKRY: Wolfert 1910, Ntume, sterile, 1911 (FI!); LIBERIA: van Meer 264, 7km from 

Bababli (06.08N:09.55W) Fl., December 12, 1965 (MO!); GHANA: Adams 2407, Awaso, Western 

Region (06.15N:02.17W) sterile, December 24, 1953 (GC!); Cummins 229, Kumasi (06.40N:01.39W) 

Fr., 1995 (K!); Enti 614, Bimpong F.R., Foso (05.43N:01.28W) Fr., February 16, 1972 (GC!, MO!, 

BR!); Hall 2748, Kakum F.R. (05.26N:01.19W) Fl., November 20, 1964 (K!, GC!); Irvine 502, 

Ashantia, Ashanti, sterile, April 30, 1927 (GC!); Irvine 2075, Awisa (07.49N:02.07W); Fl., December 

1933 (K!, GC!); Irvine 2300, Atwabo, Western Region (05.18N:02.03W) Fl., February 28, 1934 (GC!); 

Moore 2115, SS Forest Reserve, sterile, December 1920 (K!); Morton 3618, Awaso, Western Region 

(06.15N:02.17W) Fl., December 3, 1958 (K!, GC!); Vigne 1365, Banka, S. Ashanti (06.17N:01.14W) 

Fl., September 1928 (K!, FHO!, MO!); Vigne 1875, Amentia F.R., E. Region (06.10N:01.58W) Fr., 

March 30, 1930 (K!, KUM!); West-Skinn 90, Axim Cocoa Station (04.52N:02.14W) sterile, 1957 (K!); 

NIGERIA: Chapman 3681, Mambilla Plateau, sterile, April 15, 1975 (FHO!); Chapman 5202, 

Mambilla Plateau, Fl., February 3, 1978 (FHO!); Chapman 5331, Baissa River (07.14N:10.38E) Fr., 

April 1, 1978 (FHO!); Morakinyo 1006, Cross River National Park (05.15N:08.42E) sterile, August 18, 

1993 (K!); Otedoh & Tuley 7252, Okwabe, near Warri (05.22N:05.48E) sterile, August 15, 1972 (K!); 

Tuley 530, Nsukka (06.50N:07.37E) sterile, January 31, 1964 (K!); Tuley 649, Calabar to Mamfe road, 

Mile 31 (05.18N:08.34E) sterile, July 13, 1964 (K!); CAMEROON: Bos 5162, 19km from Kribi on 

Lolodorf road (03.00N:10.03E) Fr., August 7, 1969 (K!, YA!); Breteler 1241, Bertoua (04.34N:13.40E) 

Fl., April 17, 1961 (WAG!); Breteler 1560, 23km W of Yaounde on Douala road (03.05N:11.17E) Fl., 

July 7, 1961 (WAG!); Breteler et al 1200, 6km along Batouri to Betare Oyi road (04.25N:14.21E) 

sterile March 14, 1961 (WAG!); Breteler et al 2563, Mt Febe nr Yaounde (03.53N:11.31E) Fr., January 

23, 1962 (WAG!, BR!); Bruneau 1093, Kribi to Ebolowa road, Sud Province (02.51N:10.00E) sterile, 

October 30, 1995 (K!); Cheek 5591, Mt Cameroon, Upper Boando (04.04N:09.08E) Fr., November 30, 

1993 (K!, SCA!); Dransfield 6998, Mount Cameroon, Upper Boando (04.04N:09.08E) Fl., June 26, 

1991 (K!, SCA!); Fotius 3074, Djoumbi-Mbidan, 45km St Tiguere, sterile, March 12, 1978 (YA!); 

Lederman 1487, Sanchu, Fr., December 3, 1908 (FI!); Letouzey 3553, Betare Dja (03.00N:12.40E) Fr., 

February 27, 1964 (YA!); Maitland 761, Limbe (04.01N:09.11E) Fr., October 1929 (K!); Mbani 497, 

Boa Plain (04.26N:08.54E) sterile, June 5, 1994 (SCA!); Mildbraed 5264, Lomie (03.09N:13.37E) 

sterile, May 18, 1911 (HBG!); Mildbraed 5310, Lomie (03.09N:13.37E) sterile, May 21, 1911 (HBG!); 

Mpou 338, Mbalmayo (03.31N:11.30E) Fl., June 6, 1959 (YA!); Njingum 11, Nguti (05.02N:09.24E) 

sterile, August 5, 1999 (K!); Nkefor 445, Mabeta-Moliwe (03.58N:09.14E) sterile, s.d. (K!); 

Nkongmeneck 596, Nkobasso, 40km SSE of Ndiki, Fr., November 15, 1983 (YA!); Raynal 9825, 

103

Meyo-Bibilou, 36km SW Ambam (02.16N:11.11E) sterile, February 19, 1963 (YA!); Sunderland 1700, 

Mabeta-Moliwe Forest (03.58N:09.14E) Fr., October 10, 1995 (K!, SCA!, NY!); Sunderland 1711, 

Onge River valley (04.21N:08.57E) sterile, November 23, 1995 (K!, SCA!, MO!); Sunderland 1744, 

Rumpi Hills Forest Reserve (04.54N:09.20E) sterile, May 19, 1996 (K!, SCA!, BH!); Sunderland 1750, 

Mabeta-Moliwe Forest (03.58N:09.14E) sterile, November 13, 1996 (K!, BH!); Sunderland 1762, 

Mabeta-Moliwe Forest (03.58N:09.14E) sterile, November 25, 1996 (K!, SCA!); Sunderland 1885, 

10km south of Nguti (05.02N:09.24E) Fr., November 26, 1997 (K!, SCA!, BH!); Sunderland 1931, 




Takamanda Forest Reserve (06.08N:09.16E) sterile, January 18, 1999 (K!, SCA!); Sunderland 2250, 

Mokoko River Forest Reserve (04.29N:09.00E) sterile, February 16, 1999 (K!, SCA!); Tessmann s.n., 

sterile, October 2, 1909 (FI!); Thomas 6139, SW Province, nr Mundemba (04.58N:08.55E) Fl., May 12, 

1986 (K!, BR!, WAG!, MO!); Watts 511, Mt Cameroon, Njonji (04.04N:08.59E) sterile, October 15, 

1992 (SCA!); Watts 821, Liwenyi (04.23N:08.59E) Fr., October 28, 1993 (K!, SCA!); Wheatley 154, 

Mabeta-Moliwe forest (03.58N:09.14E) Fl., April 10, 1992 (SCA!); CENTRAL AFRICAN 

REPUBLIC: Carroll 33, Dzanga Region, 12km NE of Bayanga (03.07N:16.27E) sterile, April 15, 

1985 (K!, MO!); Harris 5707, Sangha-Mbaere (02.59N:16.13E) Fr., December 16, 1997 (K!); Harris & 

Fay 1494, Dzanga-Sanga (02.21N:16.09E) sterile, October 31, 1988 (MO!); EQUATORIAL 

GUINEA: Barter s.n., Bioko (03.25N:08.40E) Fl., 1857 (K!); Carvalho 2212, Bioko: Malabo to 

Cupapa (03.34N:08.46E) Fr., August 5, 1986 (K!, BR!); Guinea 913, Bioko: Boloko to Luba 

(03.24N:08.34E) sterile, January 9, 1947 (MO!); Mann 97, Bioko (03.31N:08.33E) Fl. & Fr., April 

1860 (K!); Sunderland 1875, 2km SW of village of Angoma (02.03N:10.10E) sterile, September 15, 

1997 (K!, EG!, NY!, BR!); Sunderland 1904, 10km south of Bata (01.45N:09.43E) sterile, March 20, 

1998 (K!, EG!, WAG!); Sunderland 1910, 1km from village of Mfeck-Ayong (02.00N:10.35E) sterile, 

March 18, 1998 (K!, EG!, WAG!); Sunderland 1911, 1km from village of Mfeck-Ayong 

(02.00N:10.35E) sterile, March 30, 1998 (K!, EG!, WAG!); Tessmann 44, Nkolentangau, Fr., 

December 5, 1907 (FI!); GABON: le Testu 9258, Fl., August 26, 1933 (BR!); DEMOCRATIC 

REPUBLIC OF CONGO: Bequaert 1824, Awakubi (01.19N:27.34E) Fr., January 7, 1914 (BR!); 

Evrard 3145, Befale: River Nkoyo (00.24N:20.46E) Fr., December 28, 1957 (BR!); Gerard 3218, 

Bambesa (03.28N:25.11E) Fr., December 11, 1957 (BR!); Gerard 3933, Madabu (Zobia) 

(02.58N:25.22E) Fl., May 22, 1958 (BR!); Gerard 4951, Bambesa (03.28N:25.11E) Fr., Novemver 15, 

1961 (BR!); Germain 8728, Yabahondo (Isangi) (00.42N:23.58E) sterile, March 28, 1956 (BR!); 

Lebrun 2985, Ougodia, Fl., May 1931 (BR!); Louis 9556, 30km E of Yangambi (00.45N:24.43E) Fl., 

May 30, 1938 (K!, BR!); Louis 13176, Oriental Province, Yangambi (00.45N:24.26E) Fr., December 

1938 (K!, BR!); Louis 14729, Yangambi (00.45N:24.26E) Fl., May 6, 1939 (K!, BR!, WAG!); Louis 

16340, Yangambi (00.45N:24.26E) Fr., December 7, 1939 (BR!); Mandango 3067, Ile Mbie: 5km from 

Kisangani (00.34S:29.15E) Fr., March 2, 1982 (BR!); Mildbraed 2818, Beni-Rewenzori 

(00.30N:29.25E) Fr., 1908 (FI!) 

_______________________________________ 

104

Figure 21. Laccosperma opacum (G. Mann & H. Wendl.) Drude & L. laeve (G. Mann & H. 

Wendl.) H. Wendl. 

L. laeve: Sunderland 2266; a. Stem x 1 (2/3); b. Leaf x ½ (1/3); c. Inflorescence x ½ (1/3); d. Fruit x 3 

(2); e. Seed x 3 (2): L. opacum: Sunderland 1700; f. Leaflet section x ½ (1/3); g. Inflorescence x ½ 

(1/3): Mann 97; h. Flower in bud x 4 (2 2/3); Sunderland 1885; i. Fruit x 2 (1 1/3); j. Seed x 2 (1 1/3). 


105

Laccosperma laeve (G. Mann & H. Wendl.) H. Wendl. 

(Latin) “smooth”; refers to the smooth seeds 

H. Wendl. in Kerchove, Les Palmiers 249: (1878); Küntze in Rev. Gen. Plant. 2: 

729(1891); J. Dransf. in Kew Bull. 37(30): 456 (1982); Morakinyo in Principes 39(4): 

205 (1994); Tuley in Palms of Africa 39: (1995); H.A. Burkill in Useful Pl. of W. 

Trop. Africa 4: 373 (1997). 

Ancistrophyllum laeve (G. Mann & H. Wendl.) Drude in Engl. Bot. Jar. 5:111 (1895); 

C.H. Wright in Th. Dyer, Fl. Trop. Afr. 7: 114 (1902); Becc. in Webbia 3: 261 (1910); 

Hutch. in F.W.T.A. 2: 391 (1936); Guinea-Lopez in Ensayo Geobot. de la Guinea 

Cont. Espanola 244: (1942); Walker & Sillans in Plantes Utiles du Gabon 324 (1961); 

T.A. Russell in F.W.T.A. (2): 167 (1968); Aedo et al., Bases Docs. Flora de Guinea 


Calamus (subgen. Laccosperma) laevis G. Mann & H. Wendl. in Trans. Linn. Soc. 24: 

430 (1864); Type: Gabon, Ogoué River, Mann 1045 (holotype K!). 

Clustering slender palm climbing to 10-13 m. Stems often branching, without sheaths 

to 16 mm in diameter, with c.20 mm; internodes 8-17 cm long. Leaf sheath 

moderately to sparsely armed with black-tipped, pale brown spines, spreading or 

lightly reflexed, upper sheaths near inflorescence more sparsely armed; black 

caducous indumentum present on mature leaf sheaths; ocrea 8-20 cm long, gradually 

tapering at the apex, pale straw-coloured without, shiny mid-brown within, armed as 

the sheath. Leaves up to 1.5 m; petiole on lower leaves up to 18 cm long x 0.8 cm 

wide, abaxially rounded, adaxially flattened, armed along the margins with angular, 

inequidistant black tipped, bulbous-based spines up to 1.3 cm long, spreading or 

reflexed; rachis up to 90 cm long, more commonly, 60 cm, distinctly trapezoid in 

cross section at the base, rounded in cross-section distally, armed as the petiole, spines 

decreasing in size distally and becoming regularly reflexed and increasingly bulbousbased; 

cirrus up to 70 cm long, more commonly 50-60 cm, armed as the rachis; 

leaflets composed of 2-4 folds, 10-12 pairs on each side of the rachis, inequidistant 

and unequal in size, usually subequidistant proximally and borne in loose pairs 

distally, sigmoid, 15-20 to 25-30 cm long x 3.5-6 to 8-10 cm broad at the widest point, 

106

± concolorous with prominent transverse veinlets, margin unarmed; acanthophylls 

2.5-2.8 cm long. Inflorescences, numbering 4-8 produced simultaneously in the distal 

to 30 cm of stem; peduncle c.8 cm long; prophyll c.3.5 cm; rachis bracts 1.5-2 cm 

long, gradually decreasing distally, tapering to form a short, triangular lobe, often 

tattering, finely longitudinally striate, closely adpressed to the bract above; rachillae 8- 

12 cm long, spreading, densely covered in imbricate longitudinally striate, rachillae 

bracts ± 3 mm long, each with a wide opening and 1 mm-long apiculum. Flowers at 

anthesis, 6-8 mm x 2-3 mm; calyx tan-cream, 4-4.5 mm long, tubular in the basal 2.5- 

3 mm, with 3 longitudinally striate, 2 mm x 2-2.5 mm triangular lobes; corolla, tubular 

in the basal 0.8-1 mm only, with three valvate lobes, white or pale cream, 

longitudinally striate, 6-7.5 mm x 2-2.5 mm, with a broadly acuminate tip; stamen 

filaments dark brown, 3 mm x 1 mm, united into 1 mm-long basal tube; anthers 3 mm 

x 1 mm; ovary ± 1 mm in diameter, stigma up to 4 mm long. Fruit at maturity, ovoid, 

0.8-1 cm x 0.6-0.8 cm, with 14-18 rows of vertical scales. Seed smooth, ovoid, 0.6-0.8 

cm long, 0.4-0.6 cm wide, ± 0.3 cm thick, flattened on one side, with light linear 

depression running from base to apex. 

Distribution 

L. laeve is distributed in the coastal forests from Liberia and the forest of Upper 

Guinea to Cameroon and south to Cabinda (Angola). However, there is a considerable 

disjunction from Ghana to Nigeria. 


This species, in common with, L. opacum is shade tolerant and is commonly found 

under a forest canopy. In fact, aside from on basaltic soils the two species often 

sympatric. 

107

Figure 22. Distribution of L. laeve (G. Mann & H. Wendl.) H. Wendl. 


LIBERIA: Jansen 1822, Mmal Mining Company Concession, Fr. February 13, 1970 (K!); Linder 676, 

Gbanga (06.59N:09.28W) sterile, September 17, 1926 (K!, MO!); Linder 1228, Gbanga 

(06.59N:09.28W) sterile, October 24, 1926 (K!, WAG!); CÔTE D'IVOIRE; Ake-Assi 9450, Pata 

(04.35N:07.23W) sterile, February 19, 1967 (K!); Boughey 14732, Issia (06.28N:06.33W) sterile, 

August 2, 1954 (K!); Chevalier 22658, Between Abougoua and Yacassi (05.43N:03.57W) Fr., 

December 26, 1909 (K!); de Wilde & Leeuwenberg 3432, 26km W of Abidjan (05.20N:04.16W) Fl., 

February 20, 1962 (BR!, MO!); Hall & Ake-Assi 45442, 2 miles E of Sakre (05.41N:07.21W) sterile, 

August 15, 1975 (GC!); Hepper & Maley 8214, Taï Forest (05.38N:07.08W) sterile, September 18, 

1984 (K!); Oldeman 137, Foret de Banco, sterile, July 6, 1963 (K!, WAG!); GHANA: Adams 2190, 7 

miles from Enchi (05.29N:02.29W) sterile, December 30, 1953 (GC!); Enti 2344, Neung F.R., Tarkwa 

(05.27N:00.50W) Fr., November 30, 1988 (GC!); Kinlock 3237, Tarkwa, Ndumnfri F.R. 

(05.10N:02.09W) sterile, February 3, 1934 (KUM!); Moore & Enti 9882, Ankasa River F.R. 

(05.15N:02.36W) sterile, March 4, 1971 (GC!); Sunderland 2265, Draw River Forest Reserve 

(05.12N:02.20W) sterile, May 26, 1999 (K!, KUM!); Sunderland 2266, Draw River Forest Reserve 

(05.12N:02.20W) sterile, May 26, 1999 (K!, KUM!); CAMEROON: Sunderland 1804, Campo Ma'an 

Faunal Reserve (02.10N:09.54E) sterile, March 20, 1997 (K!, YA!, MO!, WAG!); Sunderland 2251, 

Mokoko River Forest Reserve (04.29N:09.00E) sterile, February 16, 1999 (K!, SCA!); CONGO: Sita 

4642, Chaillu, Fl., April 1982 (BR!); GABON: Arends et al. 671, Mt Doubou (02.15S:10.20E), Fr., 

108

December 6, 1984 (WAG!); le Testu 1712, Region de Nyanga (02.14N:11.27E); Fl., July 3, 1919 (K!, 

BR!, MO!); Mann 1045, Gaboon River (00.19N:09.29E) Fl., 1861 (K!); Rietsma 2047, 15km NW of 

Libreville (00.35N:09.22E) Fr., March 21, 1986 (LBR!); ANGOLA: Gossweiler 7995, Mayombe, 

Luali (05.00S:12.25E) sterile, 1919 (K!) 

_______________________________________ 

Laccosperma acutiflorum (Becc.) J. Dransf. 

(Latin) refers to acuminate calyx lobes 

J. Dransf. in Kew Bull. 37(30): 456 (1982); Tuley in Palms of Africa 37: (1995); 

Cable & M. Cheek in Pl. of Mt. Cam. 179: (1998). 

Ancistrophyllum acutiflorum Becc. in Webbia 3: 255 (1910); Hutch. in F.W.T.A. 2: 

391 (1936); Guinea-Lopez in Ensayo Geobot. de la Guinea Cont. Espanola 243: 

(1942); Aedo et al. Bases Docs. Fl. de Guinea Ecuatorial 375: (1999); Type: 

Cameroon, Limbe to Bimbia, Preuss 1232 (holotype B†; isotype FI!) 

A. secundiflorum sensu T.A. Russell in F.W.T.A. 3(2):167 (1968) partim non Becc.; 

H.E. Moore in Principes 15:112 (1971) partim non Becc. 

Clustering robust to massive palm climbing to 70 m, more commonly to 30-50 m. 

Stems without sheaths 3.5-4.5 cm, with 4.5-5 cm; internodes up to 40 cm long, more 

commonly 18-25 cm. Leaf sheath light green-yellowish, conspicuously striate, 

moderately to sparsely armed with angular, black-tipped, spreading or upward 

pointing, spines, mature sheaths becoming somewhat bare, with only vestigial remains 

of spines, covered with sparse black-brown indumentum; ocrea 12-20 cm long 

(although up to 40 cm long on juvenile sheaths only), broadly sheathing and tapering 

to form a rounded lobe, dry, sometimes splitting longitudinally, light grey-brown 

without, deep crimson brown within, armed as the sheath although spines often 

concentrated in central region of ocrea. Leaves up to 3.5 m long; petiole 6-10 cm long 

x 2-3 cm. wide, much longer on juvenile sheaths (up to 45 cm) light green to dull 

yellow, with sparse light brown indumentum beneath, abaxially rounded, adaxially 

flattened or slightly concave, armed along the margins with inequidistant black-tipped, 

109

ulbous-based spines 0.8-1 cm long, angular, spreading in many directions; rachis 

yellow-green, up to 1.8-2.5m long, shaped as the petiole proximally, becoming 

trapezoid to triangular in cross section distally, armed as the petiole, although spines 

becoming more sparse distally, underside of rachis with sparse light brown 

indumentum; cirrus often bright yellow, 1.2-1.8 m long, scarsely armed, ± triangular 

in cross section; leaflets up to 50 on each side, held horizontally or arching from the 

rachis, rarely strictly pendulous with a single-fold, equidistant and sub-opposite 

proximally, alternate distally, linear lanceolate, bluntly acuminate to apiculate at the 

apex (often breaking off), 30-40 cm long, more commonly 18-30 cm, 3.0-4.5 cm 

broad at the widest point, rarely up to 6 cm wide, discolorous with dark green upper 

lamina, light green lower surface, leaflet margin armed with sub-equidistant robust, 

short, forward-facing black-tipped spines, up to 2mm long, 1-2- or rarely 3-5 

costulate, armed as the margin although spines on primary veins rather longer; 

acanthophylls bright yellow, 4-4.5 cm. long, 0.5 cm. broad, bulbous at base. 

Inflorescences, numbering 6-12 produced simultaneously in the distal 1.5-2.5 m 

portion of stem; peduncle 15-20 cm long; prophyll up to 20 cm; rachis branches up to 

50 cm long, perpendicular to the main axis, rachis bracts 3-3.5 cm long, decreasing 

distally, tapering to form an elongate triangular lobe adaxially, closely adpressed to 

the bract above, upper half dry, grey, longitudinally splitting, at first, lower half 

fleshy, bright yellow-green, then, as fruits develop, becoming dry throughout; 

rachillae 20-30 cm long, pendulous, densely covered with yellow-green imbricate 

bracts 4-5 mm long, with a wide opening and 1 mm long apiculum. Flowers at 

anthesis 1-1.2 cm x 3 mm; calyx 5-6 mm long, excluding 1.5-2.0 mm stalk, 

indistinctly striate, cream, tubular in the basal 2-3 mm with 3 broadly triangular to 

acuminate lobes 3.0-3.5 mm x 2.0-2.5 mm; corolla tubular in the basal 1.5-2 mm, with 

3 valvate lobes, white or pale cream, rarely mottled brown/tan, c.9 mm x 2 mm with 

broadly acuminate tip; stamen filaments dark brown, 4-5 mm x 1 mm, united into a 1- 

1.5 mm-long basal tube; anthers 3 mm x 1 mm, ovary c.1 mm in diameter, stigma up 

to 6 mm. Fruit at maturity ovoid, 1- (sometimes 2) seeded, 1.8-2 cm x 1.3-1.5 mm 

wide, with 17-20 vertical rows of scales. Seed smooth, ovoid, with lightly scalloped 

depression on one side 1.0-1.2 cm long x 0.8-1.2 mm wide x 0.5-0.8 mm deep. 

110

Figure 31. Laccosperma acutiflorum (Becc.) J. Dransf. 

Sunderland 1926; a. Habit; b. Stem x 2/3 (4/9); c. Leaflets x 3/8 (1/4); Leaflet section (underside) x 1 

1/8 (3/4); d. Flowers on rachilla x 1 (2/3); e. Flower x 3 (2): Sunderland 1707; f. Fruit x 4 (2 2/3). 


111

Distribution 

L. acutiflorum is distributed from Sierra Leone to Cameroon, southwards to Gabon 

and DR Congo. 

Figure 23. Distribution of L. acutiflorum (Becc.) J. Dransf. 


L. acutiflorum is a light demanding species commonly found in gap vegetation and in 

open areas. This species often occurs in seasonally-inundated and swamp forest, 

although it is also found in drier, exposed sites. L. acutiflorum responds well to 

selective logging and will colonise recently disturbed soil particularly on skid trails 

and roadsides. 

Notes 

See notes under L. secundiflorum. 


SIERRA LEONE: Gledhill 339, Koruboula to Sokwela (09.12N:10.56W) sterile, February 17, 1966 

(K!); Jordan 2064, Gola forest (07.45N:10.45W) Fl., May 13, 1955 (K!); GHANA: Chipp 643, 

Konongo, Ashanti region (06.37N:01.12W) Fr., February 4, 1914 (K!); Enti 758, Aiyaola F.R. 

112

(06.09N:01.53W) Fl., June 1972 (MO!); Sunderland 2263, Draw River Forest Reserve 

(05.12N:02.20W) sterile, May 26, 1999 (K!, KUM!); NIGERIA: Morakinyo 1000, Cross River 

National Park (05.15N:08.42E) Fl. & Fr., August 10, 1993 (K!); CAMEROON: Dransfield 7001, 

Mungo River Crossing (04.08N:09.31E) Fl., June 27, 1991 (K!); Dusen 292, Ndian (05.00N:09.00E) 

Fl. 1892 (FI!); Preuss 1232, Victoria to Bimbia (03.59N:09.13E) Fl., April 10, 1894 (FI!); Sunderland 

1707, Southern Bakundu Forest Reserve (04.46N:09.29E) Fr., November 8, 1995 (K!, SCA!, WAG!); 

Sunderland 1714, Kumba to Mamfe road (05.02N:09.24E) Fr., November 30, 1995 (K!, SCA!, BH!); 

Sunderland 1723, 30km north of Mamfe (05.58N:09.20E) Fr., December 2, 1995 (K!, SCA!, BR!); 

Sunderland 1764, 15km from Kribi on Campo road (02.34N:09.50E) Fr., December 1, 1996 (K!, YA!, 

MO!); Sunderland 1855, 10km from Kribi at Grande Batanga (02.09N:09.48E) Fl., August 30, 1997 

(K!, YA!, NY!, BR!); Sunderland 1882, Mfakwa-Supe flyover, 20km south of Nguti (05.02N:09.24E) 

Fr., November 26, 1997 (K!, YA!, BH!, MO!, WAG!); Sunderland 1926, Campo Ma'an Faunal 

Reserve (02.10N:09.54E) Fl., October 10, 1998 (K!, YA!, WAG!); Thomas 9738, Idenau 

(04.16N:09.01E) Fr., September 10, 1993 (K!, SCA!); van Gemerden 110, River Lobe, near Kribi 

(02.57N:09.54E) Fr., December 15, 1996 (K!); EQUATORIAL GUINEA: Sunderland 1907, Near 

village of Njakem (01.42N:09.40E) juvenile, March 24, 1998 ( K!, YA!, EG!, WAG!), 

DEMOCRATIC REPUBLIC OF CONGO: de Graer 216, Adjala (02.02S:15.08E) sterile, November 

12, 1933 (BR!); Louis 16995, Oriental Province, Lac Yangambi (00.47N:24.26E) Fr., September 1944 

(K!); Vanderyst 6411, sterile, 1917 (BR!) 

_______________________________________ 

Laccosperma robustum (Burr.) J. Dransf. 

(Latin) “robust” 

J. Dransf. in Kew Bull. 37(30): 456 (1982); Type: Central African Republic, Sangha- 

Mbaera, Harris 5706 (neotype K!). 

Ancistrophyllum robustum Burr. in Notizbl. Bot. Gart. Mus. Berlin-Dahlem 15: 747 

(1942); Letouzey in Adansonia 2 (18): 314 (1978); Types: Cameroon, Moloundou, 

Mildbraed s.n. (holotype B†; isotype HBG†). 

L. secundiflorum sensu Morakinyo in Principes 39(4): 207 (1994); Tuley in Palms of 

Africa, 36: (1995); White & Abernethy in Guide to Veg. of Lopé Res. Gabon 64: 

(1997). 

113

A. secundiflorum sensu Mildbr. in Notizbl. Bot. Gar. Dah. App. XXVII:16 (1913); 

Guinea-Lopez in Ensayo Geobot. de la Guinea Cont. Espanola 244: (1942); Walker & 

Sillans in Plantes Utiles du Gabon 324 (1961); T.A. Russell in F.W.T.A. 3(2):167 

(1968) partim non Burr.; Letouzey in Adansonia 2(18): 314 (1978); Letouzey in Man. 

For. Bot. Trop. Afr. 2B:401 (1986); Aedo et al. Bases docs. Fl. de Guinea Ecuatorial 

375 (1999). 

Clustering robust palm climbing to 30-45 m. Stems without sheaths 30-50 mm in 

diameter, with 45-60 mm; internodes 35-50 cm long, although more commonly 18-25 

cm. Leaf sheath moderately to profusely armed with black-tipped finely triangular, 

upward pointing or spreading spines; sheaths on upper portion of stem more sparsely 

armed; juvenile sheaths particularly profusely armed; dense brown-black indumentum 

present on mature sheaths, more sparse on juvenile sheaths; ocrea 20-30 cm long, dry, 

gradually tapering at the apex, reflexed, tattering longitudinally and disintegrating, 

dark tan mid-brown without, crimson-brown within, armed as the sheath although 

spines often concentrated at the apex, particularly on juvenile sheaths. Leaves up to 

3.5m long; petiole 5-12 cm long, 1.5-2.5 cm. wide, mid to dark green with scattered 

brown-black indumentum, particularly on upper surface, abaxially rounded, adaxially 

lightly to moderately concave proximally becoming flattened in cross section distally, 

armed along the margins with inequidistant black-tipped angular spines up to 1.4 cm 

long, spreading in many directions; rachis 1.5-2 m long, shaped as the petiole, 

becoming more trapezoid in cross section then triangular in cross-section distally, 

armed as the petiole proximally, spines becoming more sparse distally, with sparse 

brown indumentum present below; cirrus up to 1.5-2 m. long, triangular in cross 

section, only very sparsely armed with reflexed bulbous-based black-tipped spines, 

sparse brown indumentum below; leaflets always composed of a single-fold, 45-65 on 

each side of the rachis, equidistant, opposite to sub-opposite proximally, inequidistant, 

alternate distally, papyraceous, conspicuously pendulous, finely linear-lanceolate, 

base abruptly contracted, apex broadly to finely acuminate, often breaking off, 30-45 

cm long (commonly up to 60 cm) 1.2-2.8 cm broad at the widest point, ± concolorous, 

glaucous blue-green, margins armed with inequidistant exceptionally fine, 5-8mmlong, 

forward facing curved or angular black-tipped spines; 1 costulate midrib armed 

as the margins although spines much longer.; acanthophylls 4.5-5 cm long. 

Inflorescences, numbering 6-12 produced simultaneously in the distal 1.5-2.2m 

114

portion of stem; peduncle 12-20 cm long; prophyll ± 15 cm long; rachis branches up 

to 50 cm long, perpendicular to the main axis; rachis bracts glabrous, 1.5-1.8 cm long, 

decreasing distally, dry throughout, covered in dense brown-black indumentum, 

tapering to form a truncate triangular lobe abaxially, closely adpressed to the bract 

above; rachillae 18-25 cm long, pendulous, rachillae bracts campanulate-cylindrical, ± 

5 mm long, dry throughout, triangular to broadly acuminate adaxially, each with a 

wide opening and a 1.5 m.-long apiculum. Flowers in dyads, rarely triads, at anthesis 

± 1 cm x 2.5-3.0 mm wide; calyx ± 6 mm long, excluding 1-1.5 mm long angular 

stalk, 2.5-3 mm wide, tubular in the basal 4 mm only, with 3 dark-tan, lightly striate, 

rounded to triangular, rarely acuminate, apex, 3.5 x 3 mm; corolla, tubular in the basal 

2 mm only, with 3 valvate lobes c.7 x 2 mm, white or cream, with a bluntly acuminate 

apex; stamen filaments, brown, fleshy, angular, 3 mm x 1 mm broad, united into 1.5 

mm-long basal tube; anthers 3 mm x 1 mm broad; ovary ± 1 mm in diameter, stigma 

up to 5 mm. Fruit at maturity ovoid; 1- (rarely 2-) seeded, 1.2-1.5 cm x 0.8-1.2 cm, 

with 17-20 vertical rows of scales. Seed smooth, ovoid, with lightly scalloped 

depression on one side, 0.8-1.3 cm long x 0.6-0.8 cm wide x 0.5 cm thick. 

115

Figure 24. Laccosperma robustum (Burr.) J. Dransf. 

Sunderland 1757; a. Habit; b. Stem x 2/3 (4/9); c. Leaflets x ½ (1/3); d. Leaflet detail x1 (2/3); e. 

Acanthophylls x ½ (1/3): Sunderland 1791; f. Infructescence x ½ (1/3); g. Fruit x 2 ½ (1 2/3). Drawn 

by Lucy T. Smith. 

116

Distribution 

L. robustum is very common species throughout its range and is distributed from SE 

Nigeria to the central Congo Basin. 

Figure 25. Distribution of L. robustum (Burr.) J. Dransf. 


This species is commonly encountered in forest gaps and regrowth vegetation and 

responds well to selective-logging activities. It is encountered on both terra firma and 

seasonally-inundated forest. 

Notes 

L. robustum was first described by Burret (1942) from specimens collected in 

Moloundou early in the 20 th century by Mildbraed. Although all the material that 

Burret cites is now extinct, destroyed in the allied bombing of Germany in World War 

II, his description clearly matches this taxon. In this respect a neotype is assigned for 

this species (Harris 5706) collected in the same area, although across the border in 

what is now the Central African Republic. Further elaboration on the status of the 

117

large-diameter species of Laccosperma can be found in the notes under L. 

secundiflorum. 


NIGERIA: Smith 53, Calabar (04.59N:08.20E) Fl., June, 1931 (K!); CAMEROON: Bililong & 

Bullock 348, Campo Faunal Reserve (02.14N:09.54E) seedling, s.d. (K!); Bos 4799, 20km from Kribi 

(03.00N:10.03E) Fl., June 10, 1969 (K!, MO!, WAG!, YA!, BR!); Bos 5160, 20km from Kribi, 

(03.00N:10.03E) Fr., August 7, 1969 (K!, WAG!, BR!, YA!); Dransfield 7006, Limbe to Kumba road, 

Mile 40 (05.02N:09.24E) sterile, June 28, 1991 (K!, SCA!); Gentry & Thomas 52727, Korup National 

Park (05.00N:08.30E) sterile, November 12, 1985 (K!, MO!); Gentry & Thomas 52766, Korup 

National Park (05.00N:08.30E) sterile, November 12, 1985 (MO!); Letouzey 7368, Nr Benga on Douala 

to Yaounde rd, Fl., July 7, 1966 (YA!); Letouzey 8479, Nr Kamelon, 10km SE Sangmelima 

(02.55N:11.58E) Fr., November 24, 1966 (YA!); Lowe 3442, Edea - Kribi rd nr Elogbatindi 

(03.27N:10.11E) sterile, January 22, 1978 (K!, YA!); Njingum 1, Nkakanzock, near Edea 

(03.49N:10.14E) sterile, May 15, 1999 (K!); Njingum 3, NW Province, Bagoran, sterile, June 1, 1999 

(K!); Njingum 5, Akom II (02.47N:10.34E) sterile, July 1, 1999 (K!); Njingum 9, Nguti 

(05.02N:09.24E) sterile, August 5, 1999 (K!); Sunderland 2305, Korup National Park, Chimpanzee 

Camp (05.02N:08.48E) sterile, February 18, 2000 (K!, SCA!); Sunderland 2306, Korup National Park, 

Chimpanzee Camp (05.02N:08.48E) sterile, February 18, 2000 (K!, SCA!); Sunderland 1645, Mokoko 

River Forest Reserve (04.29N:09.00E) sterile, May 1, 1994 (K!, SCA!); Sunderland 1708, Southern 

Bakundu Forest Reserve (04.46N:09.29E) Fr., November 8, 1995 (K!, SCA!, BR!); Sunderland 1722, 

30km north of Mamfe (05.58N:09.20E) sterile, December 2, 1995 (K!, SCA!, BH!); Sunderland 1740, 

Rumpi Hills Forest Reserve (04.54N:09.20E) sterile, May 19, 1996 (K!, SCA!, WAG!); Sunderland 

1747, Rumpi Hills Forest Reserve (04.54N:09.20E) sterile, May 19, 1996 (K!, SCA!, MO!); 

Sunderland 1757, Limbe - Kumbe road, Mile 40 (04.23N:09.26E) sterile, November 11, 1996 (K!, 

SCA!, MO!); Sunderland 1928, Campo Ma'an Faunal Reserve (02.10N:09.54E) sterile, October 11, 

1998 (K!, YA!); Sunderland 1930, Takamanda Forest Reserve (06.06N:09.47E) sterile, November 9, 

1998 (K!, SCA!); Sunderland 1935, Takamanda Forest Reserve (06.06N:09.47E) sterile, November 15, 

1998 (K!, SCA!); Sunderland 2058, Takamanda Forest Reserve (06.08N:09.16E) sterile, January 18, 

1999 (K!, SCA!); Sunderland 2253, Mokoko River Forest Reserve (04.29N:09.00E) sterile, February 

16, 1999 (K!, SCA!); Sunderland 2254, Mokoko River Forest Reserve (04.29N:09.00E) sterile, 

February 16, 1999 (K!, SCA!); van Gemerden Bi, Kribi to Lolodorf road (03.13N:10.38E) sterile, s.d. 

(K!); CENTRAL AFRICAN REPUBLIC: Fay 8236, Ndakan (02.21N:16.09E) sterile, February 16, 

1988 (MO!); Fay 8254, Ndakan (02.21N:16.09E) sterile, February 20, 1988 (MO!); Harris 5704, 

Sangha-Mbaere (02.59N:16.13E) Fr., December 16, 1997 (K!); Harris 5706, Sangha-Mbaere 

(02.59N:16.13E) Fr., December 16, 1997 (K!); Harris 5718, Sangha-Mbaere (02.59N:16.13E) Fr., 

December 25, 1997 (K!); Harris 5719, Sangha-Mbaere (02.59N:16.13E) Fr., December 25, 1997 (K!); 

Harris 5720, Sangha-Mbaere (02.59N:16.13E) Fr., December 25, 1997 (K!); EQUATORIAL 

GUINEA: Sunderland 1791, Bata to Mbini road, 17km from Bata (01.45N:09.43E) Fr., March 11, 

118

1997 (K!, EG!, BH!); Sunderland 1799, 4km north of Ayemeken village (02.10N:10.03E) Fr., March 

13, 1997 (K!, EG!, NY!, MO!); Sunderland 1915, 5km from village of Nsork (01.53N:11.06E) Fr., 

April 2, 1998 (K!, EG!, WAG!); Tessmann 2, sterile, s.d. (FI!); GABON: Wilks 1486, 20km N of 

Koumameyong (00.22N:11.54E) Fr., April 1, 1987 (MO!); DEMOCRATIC REPUBLIC OF 

CONGO: Bermejo 19, Equator: zone d'Ikela (00.36S:20.06E) Fr., s.d. (BR!); Breyne 2357, Maluku, 

Plateau des Bateke (03.52S:15.00E) Fr., June 15, 1975 (MO!, BR!); Dubois 911, Boendu, S of Maringa 

(00.56N:20.03E) Fr., August 1938 (K!, BR!, MO!); Evrard 4943, Kodoro-Yekokora (01.16N:20.06E) 

Fr., September 26, 1958 (BR!); Leonard 1671, Popolo (03.06N:20.45E) Fr., August 20, 1955 (BR!); 

Leonard 3817, Kigulube: Shabunda (02.36S:28.00E) Fr., April 11, 1959 (BR!); Liegeois 89, sterile, 

July 1943 (BR!); Louis 6445, Yangambi (00.45N:24.26E) Fr., October 27, 1937 (BR!); Louis 16049, 

River Bokuye (tributary of the Luye), sterile, September 11, 1939 (BR!); Louis 16794, Yangambi 

(00.45N:24.26E) sterile, November 17, 1943 (BR!); Ndjele 732, 44km from Lubutu (00.42N:26.32E) 

sterile, June 22, 1981 (BR!); Toussaint 2294, Vallee de la Nkula, Fr., May 7, 1947 (BR!, FHO!); 

ANGOLA: Gossweiler 7541, Mayombe, Luali (05.00S:12.25E) sterile, 1919 (K!) 

_______________________________________ 

Laccosperma secundiflorum (P. Beauv.) Küntze 

(Latin) refers to the inflorescence structure with hermaphrodite flowers in pairs 

Küntze in Rev. Gen. Pl. 2: 729 (1891); J. Dransf. in Kew Bull. 37(3): 456 (1981); 

Johnson in Principes 28(4): 161 (1984); Profizi in RIC Bull. 5(1): 2 (1986); Hawthorn 

in Trees of Ghana 225: (1990); H.A. Burkill in Useful Pl. of W. Trop. Afri. 4: 374 

(1997); Cable & M. Cheek in Pl. of Mt. Cam. 179: (1998). 

Ancistrophyllum secundiflorum (P. Beauv.) H. Wendl. in Kerchove, Les Palmiers: 230 

(1878); Drude in Engl. Bot. Jarbh. 5: 131 (1895); C.H. Wright in F.T.A. 7: 115 

(1902); Becc. in Webbia 3: 251 (1910); De Wild. in Ann. Mus. Col. de Marseille 7(3): 

28 (1919); Holland in Kew Bull. 9: 727 (1922); Hutch. in F.W.T.A. 2: 391 (1936); A. 

Chev. in Rev. de Bot. Appl. 17: 897 (1936); Staner & Boutique in Mem. l’Inst. Roy. 

Col. Belge 13: (1937); Dalziel in App. to F.W.T.A. 495: (1937); Renier in Fl. du 

Kwango 1: 82 (1948); Irvine in Econ. Bot. 6(23): 31 (1952); Fosberg in Principes 4: 

129 (1960); Irvine in Woody Plants of Ghana 773: (1961); T.A. Russell in F.W.T.A. 

3(2): 167 (1968); H.E. Moore in Principes 15: 112 (1971); Letouzey in Adansonia 

2(18): 314 (1978); Bauchet in Fl. du Sénégal IX:74 (1988). 

119

Calamus (subgen. Ancistrophyllum) secundiflorus G. Mann & H. Wendl. Trans. Linn. 

Soc. 24: 432 (1864). 

Calamus secundiflorus P. Beauv. in Fl. D’Oware et de Benin 1:15, t9, 10 (1805); 

Hook. in Fl. Nigr. 526: (1849); Durand & Schinz. in Consp. Fl. Afr. 5: 456 & Etude. 

Fl. de l’Etat Ind. du Congo 17: (1896); Durand & Durand in Fl. Cong. 1: 584 (1909); 

Pyneart in Bull. Agr. Congo Belge. 2: 550 (1911); Type: Nigeria (Benin) (Palisot de 

Beauvois s.n. (holotype G! (Herb. de Candolle)). 

Laccosperma laurentii (De Wild.) J. Dransf. in Kew Bull. 37(30): 456 (1982). 

Ancistrophyllum laurentii De Wild. in Bull. Jard. Bot. Brux. 5: 148 (1916); synon. 

nov. Type: DR Congo, Laurent s.n. (holotype BR!). 

Laccosperma majus (Burr.) J. Dransf. in Kew Bull. 37(30): 456 (1982) 

Ancistrophyllum majus Burr. in Notizbl. Bot. Gart. Mus. Berlin-Dahlem 15: 747 

(1942); synon. nov. Type: Bioko, Equatorial Guinea, Mildbraed 6873 (holotype B†; 

isotype HBG†). 

Clustering moderate to robust palm, climbing to 25-50 m. Stems without sheaths 20- 

25 mm in diameter; with 30-35 mm; internodes 18-35 cm long. Leaf sheath dark 

green, lightly striate, moderately to sparsely armed with black-tipped finely triangular, 

upward-pointing or spreading spines; sheaths on upper portion of stem more sparsely 

armed; sparse black indumentum present on mature sheaths; ocrea 25-35 cm long, dry, 

often tattering, gradually tapering at the apex, dark tan coloured without, shiny dark 

brown to dull maroon within, armed as the sheath, spines concentrated at apex. Leaves 

up to 3.5 m long; petiole 30-60 cm long, 1.5-1.8 cm wide, commonly at 45-60° angle 

to the sheath, light to mid-green to straw coloured often with scattered brown 

indumentum below, abaxially rounded, adaxially concave, becoming flattened, 

somewhat rectangular in cross section distally, armed along the margins with 

inequidistant black-tipped spines up to 1.8 cm long, angular, spreading in many 

directions; rachis up to 1.2-1.5m long, hexagonal in cross section proximally 

becoming trapezoid then rounded in cross section distally, armed as the petiole, spines 

becoming increasingly short and more sparse distally; cirrus up to 1.5-1.8m long, 

armed on the underside with inequidistant, reflexed, black-tipped spines, with sparse 

120

own indumentum below; leaflets composed of 2-4-folds, 25-40 on each side of the 

rachis, equidistant, often variable in width, arching from the rachis, not strictly 

pendulous, sigmoid, elongate, leaflet apex very finely acuminate with tip often 

breaking off, 35-45 cm long x 3-8 cm broad at the widest point, concolorous or 

somewhat discolorous with a darker green adaxial surface, leaflet margin armed with 

fine to robust, 1-2 mm-long, forward-facing, equidistant black-tipped spines, 1,2 or 3 

costulate, each vein armed as the leaflet margin; acanthophylls, up to 4 cm long. 

Inflorescences, numbering 6-10, produced simulataneously in the distal 1-1.8 m 

portion of stem; peduncle 15-20 cm long; prophyll 8-10 cm; rachis branches 25-35 cm 

long, perpendicular to the main axis; rachis bracts 2.5-3 cm long, decreasing distally, 

dry, often tattering, tapering to form an elongate acutely triangular lobe on the abaxial 

side, closely adpressed to the bract above, covered with a dense brown indumetum; 

rachillae 15-25 cm long, pendulous, densely covered with imbricate bracts c.7 mm 

long, dry and triangular at apex, each with a wide opening and a 1.5 mm-long 

apiculum. Flowers at anthesis 1-1.2 cm x 3.0-3.5 mm; calyx 5-5.5 mm long, excluding 

angular 3 mm-long stalk, 3-3.5 mm wide, dark tan coloured, tubular in the basal 1.5-2 

mm, with 3 longitudinally-striate or mottled rounded to bluntly triangular lobes ± 4.5 

mm x 3 mm; corolla tubular in the basal 1 mm only, with 3 valvate lobes, white or 

pale cream, longitudinally striate or mottled, ± 9 mm x 2 mm with a bluntly acuminate 

tip; stamen filaments dark brown, 4 mm x 1 mm, united into a 2 mm-long basal tube; 

anthers 3 mm x 1 mm; ovary c.1 mm in diameter, stigma up to 5 mm long. Fruit 

ovoid, 1.8-2 cm x 1.3-1.5 cm, with 18-22 vertical rows of scales. Seed smooth, ovoid, 

1-1.2 cm long, 0.8-1.2 cm wide, 0.5-0.7 mm deep, lightly flattened on one side. 

121

Figure 26. Laccosperma secundiflorum (P. Beauv.) Küntze 

Sunderland 2255; a. Stem x 2/3 (4/9): Sunderland 2259; b. Leaflets x 1/3 (2/9); c. Acanthophylls x 1 

(2/3); Tuley photograph (1964); d. Inflorescence: Mann 453; e. Flower x 4 (2 2/3); f. Flower section x 4 

(2 2/3): Tuley photograph (1963); g. Fruit x 3 (2). Drawn by Lucy T. Smith. 

122

Distribution 

This species is distributed from Sénégal to Cameroon south to DR Congo. 

Figure 27. Distribution of L. secundiflorum (P. Beauv.) Küntze 


L. secundiflorum is a species of high forest, and is commonly found under a forest 

canopy. 

Notes 

A number of species of the large diameter members of the genus Laccosperma were 

described in the early part of the 20 th century by Beccari (1910) and de Wildemann 

(1916) and later by Burret (1942). Although Burret, in particular, was prone to 

applying very narrow typological species concepts (see Henderson, 1999) he has been 

proven to be somewhat accurate in the majority of his species accounts within this 

genus and Oncocalamus. Despite this, and obvious morphological discontinuities 

123

within this complex, evident both from herbarium material and field observations, 

many of the species described both by Burret and Beccari were never recognised. 

One of the major reasons for this is the fact that in their original paper in which Mann 

and Wendland (1864) re-described Laccosperma secundiflorum (syn. Calamus 

secundiflorus), the account was accompanied by an illustration by Fitch, probably 

based on Mann’s field sketches, that showed the growth habit of a number of African 

palms including that of L. secundiflorum. However, with its conspicuously pendant, 

and distinctively linear leaflets, the rattan palm labelled in the account as “L. 

secundiflorum” bears little resemblance to the species of de Beauvois, nor indeed to 

the original collections of Mann the account cites. The taxon drawn is undoubtedly a 

species described later by Burret (1942) as L. robustum. This confusion seems to have 

given credence to the oft-mentioned claim of many of the subsequent floristic studies 

that L. secundiflorum is a highly polymorphic taxon both within and between 

populations. As such, these floristic studies continued to group all of the largediameter 

species of Laccosperma into a single species complex: L. secundiflorum 

(Drude, 1895; Wright, 1902; Unwin, 1920; Dalziel, 1937; Renier, 1948; Robyns and 

Tournay, 1955; Irvine, 1961; Russell, 1968; Letouzey, 1978; Dransfield, 1986; 

Berhaut, 1988; Morakinyo, 1995b). 

However, field workers began to note that there were in fact at least two species of 

large-diameter Laccosperma in West and Central Africa. The morphological 

differences between taxa are obvious, particularly as these species are often sympatric. 

Paul Tuley, who was working in Nigeria at the time, wrote to Tom Russell at Kew, 

who was then preparing an account of the Palmae for the revision of the Flora of West 

Tropical Africa. Tuley suggested to Russell that there might be at least two “forms” 

within the L. secundiflorum complex with “one [form] having dropping segments and 

the other with rigid segments that are held horizontally”. Tuley was quite clearly 

describing, in the first instance, the pendant leaflet habit of L. robustum. However, 

Russell discounts his observations, and states that “all the flowering material at Kew, 

including collections of Tuley, are referrable to Ancistrophyllum secundiflorum” 

(Russell, 1968). In addition, Chapin, a botanist active in the then Belgian Congo 

makes the distinction between the true L. secundiflorum and “the one with pendant 

leaflets that grows in swamps” (Chapin 613, herb. BR!) (authors’ translation from the 

124

French). A brief discussion of this is also reported in the recent article by Prance et al. 

(2000). Tuley (1995) went some way in distinguishing between the taxa, in describing 

L. acutiflorum and L. secundiflorum as separate taxa, but no doubt a lack of adequate 

voucher material hindered the preparation of a full account of this group. 

However, since then, further field-work and examination of herbarium material has 

confirmed that this “taxon” is composed of three distinct species as I have described 

above. 


SENEGAL: Vanden-Berghen 7285, Casamance (12.51N:15.17W) sterile, November 26, 1984 (BR!); 

GUINEA-BISSAU: d'Orey 262, Cacine (11.06N:15.00W) Fl., January 30, 1954 (K!); GUINEA- 

CONAKRY: Chillou 1419, Condoya forest, sterile, May 16, 1939 (BR!); Roberty 2903, Irie 

(08.16N:09.12W) Fl., December 21, 1954 (K!); SIERRA LEONE: Deighton 2593, Njala 

(08.06N:10.46W) sterile, January 1933 (K!); Deighton 4119, Gola forest (07.45N:10.45W) Fr., March 

10, 1945 (K!); Deighton 3090, Tabe (08.01N:11.56W) Fr., September 21, 1935 (K!); Jaeger 1802, 

Between Krutox & Siki Koro, sterile, September 21, 1945 (MO!); Jordan 2065, Gola forest 

(07.45N:10.45W) Fr., May 13, 1955 (K!); Jordan 2066, Gola forest (07.45N:10.45W) Fl., May 14, 

1955 (K!); Pyne 39, Joru to Daru road (07.41N:11.03W) sterile, November 2, 1955 (K!); Small 697, 

Gola forest (07.45N:10.45W) Fl., May 24, 1952 (K!); LIBERIA: Adam 20746, Fr., January 25, 1965 

(K!); Bos 2165, St Paul river (06.30N:10.40W) Fl., July 27, 1966 (K!, WAG!, BR!); CÔTE 

D'IVOIRE: Bernardi 8382, Tien-Oula (06.45N:07.04W) Fr., March 2, 1962 (K!); de Wilde 3282, 

Nigbi (06.39N:06.39W) Fr., November 21, 1961 (K!, WAG!, BR!); Hepper & Maley 8178, Taï Forest 

(05.38N:07.08W) sterile, February 9, 1984 (K!); Hepper & Maley 8062a, Foret de Gheoule, sterile, 

February 3, 1984 (K!); Leeuwenberg 3954, 15km NE of Bianouan (06.06N:03.09E) Fr., April 17, 1962 

(K!, MO!, WAG!, BR!); GHANA: Foggie 16/40, Fr., 1940 (K!); Hall 3371, Nkroful, Western Region 

(04.56N:01.44W) Fl., August 18, 1965 (K!, GC!); Moore & Enti 9892, Esiama-Nkroful road 

(04.56N:01.44W) sterile, March 7, 1971 (GC!); Morton 377, Ankobra River, Axim (04.55N:02.16W) 

sterile, August 18, 1965 (GC!); Sunderland 2259, Road from Tarkwa to Axim (05.21N:01.00W) sterile, 

May 25, 1999 (K!, KUM!); Tomlinson s.n., Bobiri F.R. (06.38N:01.17W) sterile, December 20, 1957 

(K!, GC!); Vigne 2410, Mampong Escarpment (07.17N:01.27W) Fl., August 30, 1932 (K!, KUM!); 

BENIN: Aufsess 427, Adjarra (06.32N:05.52E) sterile, December 6, 1988 (K!); Latilo s.n., Sapoba 

F.R. (06.06N:05.52E) Fr., January 25, 1951 (GC!); NIGERIA: Arwaodo 42, Aluu, Niger Delta 

(05.00N:06.00E) sterile, March 16, 1998 (K!); Ayewoh 3853, Ondo Province, Owo (08.25N:03.20E) 

Fl., February 24, 1944 (K!); Barter 4, Niger Delta (05.00N:06.00E) Fl., 1859 (FI!); Brenan 8580, 

Okomu F.R., Fl., December 21, 1947 (K!, FHO!, BR!); Keay FHI 6996, sterile, November 24, 1943 

(K!); Magajie & Tuley 2166, Ankpa Division (07.23N:07.36E) sterile, February 28, 1971 (K!); Maggs 

125

150, Ikot Arna (05.06N:07.36E) Fl., August 23, 1948 (K!); Mann 453, Mouth of the River Niger 

(04.35N:07.00E) Fl., August 1, 1960 (K!, FI!); Tuley 851, Ikorundu Causeway, Lagos (06.28N:03.20E) 

Fl., September 4, 1964 (K!); Tuley 454, Between Nsu and Okigwe (05.39N:07.14E) sterile, January 28, 

1964 (K!, WAG!); Tuley 648, Calabar to Mamfe road, Mile 31 (05.18N:08.34E) Fr., July 13, 1964 

(K!); CAMEROON: Aninze 24732, Kembong F.R., SW Province (05.38N:09.14E) sterile, September 

5, 1951 (K!); Gentry et al. 62566, Sango River, East Province (02.22N:16.09E) sterile, May 8, 1988 

(MO!); Harris & Payne 2469, Korup National Park (05.00N:09.00E) Fl., August 29, 1990 (K!); 

Letouzey 10605, Banks of Ngoko river nr Pandama, 8km ESE of Moloundou (02.03N:15.09E) Fl., 

April 4, 1971 (YA!); Sunderland 1710, Onge River valley (04.21N:08.57E) sterile, November 23, 1995 

(K!, SCA!, NY!); Sunderland 1755, Limbe - Douala road at Mungo Bridge (04.08N:09.31E) sterile, 

November 16, 1996 (K!, SCA!, BH!); Sunderland 1763, 15km from Kribi on Campo road 

(02.34N:09.50E) sterile, December 1, 1996 (K!, YA!, BR!); Sunderland 2048, Takamanda Forest 

Reserve (06.08N:09.16E) Fl., January 18, 1999 (K!, SCA!); Sunderland 2059, Takamanda Forest 

Reserve (06.08N:09.16E) sterile, January 18, 1999 (K!, SCA!); Sunderland 2255, Mokoko River Forest 

Reserve (04.29N:09.00E) sterile, February 16, 1999 (K!, SCA!); Thomas 2292, Korup National Park 

(04.55N:08.50E) Fl., July 16, 1983 (K!, MO!, YA!); CENTRAL AFRICAN REPUBLIC: Harris & 

Fay 449, Ndakan (02.21N:16.09E) Fl., April 9, 1988 (MO!, BR!); GABON: de Wilde et al. 9917 

(01.43S:09.50E) sterile, December 1, 1989 (WAG!); DEMOCRATIC REPUBLIC OF CONGO: 

Bequaert 7076, Barumbu (01.13N:23.30E) sterile, March 12, 1915 (BR!); Chapin 613, Mompoto, 

above Lukolela (01.03S:17.12E) Fr., October 22, 1930 (BR!); Corbisier-Baland 1992, Eala 

(00.03N:18.18E) Fl., June 20, 1933 (BR!); Couteaux 472, Eala (00.03N:18.18E) Fl., October 20, 1938 

(BR!); Evrard 1686, Equator Province, Popolo (03.06N:20.45E) Fr., August 22, 1955 (K!); Germain 

1669, Eala (00.03N:18.18E) Fr., October 1943 (K!, WAG!, BR!); Hulstaert 869, Eala (00.03N:18.18E) 

Fr., November 1, 1942 (BR!); Hulstaert 1419, Bokela (01.07S:21.55E) Fr., s.d. (BR!); Hulstaert 1623, 

Bamanga (00.16S:25.32E) sterile, September 17, 1954 (BR!); Hulstaert 1624, Bamanga 

(00.16S:25.32E) sterile, September 17, 1954 (BR!); Laurent 913, Eala (00.03N:18.18E) Fl., June 11, 

1909 (BR!); Laurent s.n., Equator Province, Eala (00.03N:18.18E) Fl., 1905 (K!, BR!); Laurent s.n., 

Fl., 1906 (BR!); Lejoly 1512, Batikalela, 43km from Kisangani (00.18N: 25.33E) sterile, May 22, 1977 

(BR!); Leonard 54, Bank of the river Bonkele, between Bamania & Ilalenga (near Eala) 

(00.03N:18.18E) Fr., September 26, 1945 (BR!); Leonard 815, Between Eala et Bantoie 

(00.03N:18.18E) Fr. October 12, 1946 (K!, BR!, WAG!); Leonard 1686, Popolo (03.06N:20.45E) Fr., 

August 22, 1955 (BR!); Leonard 3960, River Lomela (02.17S:23.15E) Fl., April 21, 1958 (BR!); Louis 

3646, Yangambi, 5km N de fleuve (00.45N:24.26E) Fl., April 14, 1937 (BR!); Louis 3699, Yangambi: 

5km N of Yaosuka (00.45N:24.26E) Fl., April 18, 1947 (K!, BR!); Louis 3958, Yangambi, 4km N du 

fleuve (00.45N:24.26E) Fl., May 24, 1937 (K!, BR!, FHO!, WAG!, MO!); Louis 11739, Ile Esali, en 

face de Yangambi (00.45N:24.26E) sterile, s.d. (BR!); Louis 16995, Yangambi (00.45N:24.26E) Fr., 

September 1, 1944 (BR!); Sapin s.n., sterile, June 26, 1906 (BR!); Vanderyst 1408, Kwango, sterile, 

1913 (BR!); Vanderyst 9974, Kikwit (05.02S:18.48E) sterile, June 1921 (BR!) 

_______________________________________ 

126

Imperfectly-known taxon 

Laccosperma sp. 1 

This taxon is obviously closely-related to L. laeve and L. opacum, inhabiting the forest 

understorey and with the tendency to produce aerial branches. However, this species 

possesses distinctly lanceolate leaves and has a very sparsely-armed sheath. Even 

more unusual, and the reason why this taxon is distinct, is the fact that acanthophylls 

that are entirely absent from the cirrus, which is armed by short, recurved thorns; most 

unusual in this genus and within the group of rattans endemic to Africa. This taxon is 

distributed from SW Cameroon, where it is abundant in the Korup National Park, to 

southern Cameroon, in coastal forests. The collection of fertile material will allow a 

full description of this species to be made. 


CAMEROON: Collector unknown, Kumba (04.38N:09.26E) sterile, s.d. (FI!); Cheek 5062, Koto II 

(04.21N: 9.02E) sterile, October 2, 1993 (K!, SCA!); Harris 3660, Onge (04.21N:08.57E) sterile, July 

11, 1993 (K!); Harris 3742, Onge (04.21N:08.57E) sterile, September 11, 1993 (K!, SCA!); Harris 

3778, Onge (04.21N:08.57E) sterile, October 11, 1993 (K!, SCA!); Njingum 2, Bidou I, near Kribi 

(02.50N:09.58E) sterile, July 15, 1999 (K!); Sunderland 2303, Korup National Park, Chimpanzee 

Camp (05.02N:08.48E) sterile, February 15, 2000 (K!, SCA!); Thomas 9726, Idenau (04.16N:09.01E) 

sterile, September 10, 1993 (K!, SCA!) 

Indet. Laccosperma 

_______________________________________ 

CÔTE D'IVOIRE: de Wilde 3101, 25km W of Abidjan (05.20N:04.16W) sterile, September 30, 1961 

(WAG!); GHANA: Moore & Enti 9886, Ankasa River F.R. (05.15N:02.36W) Fr. only, March 3, 1971 

(GC!); NIGERIA: Lowe 2792, ca. 40 miles SE of Benin (05.59N:06.07E) sterile, March 28, 1974 (K!); 

CAMEROON: Harris 3739, Onge (04.21N:08.57E) sterile, September 11, 1993 (SCA!); Maruhashi 

171, Ejagham Council Forest Reserve (05.30N:09.00E) Fr. only, 1981 (YA!); Njingum 10, Nguti 

(05.02N:09.24E) sterile, August 5, 1999 (K!); Sunderland 1737, Rumpi Hills Forest Reserve 

(04.54N:09.20E) seedling, May 19, 1996 (K!, SCA!, BH!, NY!); CENTRAL AFRICAN REPUBLIC: 

Fay 8255, Ndakan (02.21N:16.09E) sterile, February 20, 1988 (MO!); Harris 3386, 25km SE of 

Bayanga (02.47N:16.25E) sterile, July 5, 1993 (K!); Harris 4232, 6km from Bayanga (02.55N:16.16E) 

sterile, January 4, 1994 (K!); GABON: Soyeaux 155, Munda, sterile, September 1880 (K!, FI!); DR 

CONGO: Baland 1992, Eala (00.03N:18.18E) Fl. only, s.d. (K!, WAG!); Breyne 2105, Kimpete 

127

(04.09S:15.49E) Fr. only, March 17, 1971 (BR!); Compère 2183, Mputu (04.46S:15.31E) seedling, 

June 8, 1960 (BR!); de Giorgis 174, sterile, July 1908 (BR!); Gillet 3812, Fl. only,1904 (BR!); Laurent 

s.n., Eala (00.03N:18.18E) Fr. only (BR!); Luja 223, seedling, March 13, 1899 (BR!); Szafranski 1578, 

River Zaire: km 635, sterile, June 29, 1989 (BR!); Vanderyst 5139, Dima (03.16S:17.28E) Fl. only, 

1914 (BR!); ANGOLA: Gossweiler 6416, Mayombe, Luali (05.00S:12.25E) Fr. only, 1919 (K!); 

Gossweiler s.n., Mayombe, Luali (05.00S:12.25E) Fl. only, 1924 (K!) 

_______________________________________ 

Laccosperma: Excluded names and nomina nuda 

Eremospatha yangambiensis Louis & Mullenders nom. nud. = Laccosperma opacum 

128

ONCOCALAMUS (G. Mann & H. Wendl.) H. Wendl. 

G. Mann & H. Wendl. ex Hook.f. in Benth. and Hook. Genera Plantarum 3:881, 936 

(1883). Type: O. mannii (H. Wendl.) H. Wendl. (Calamus mannii H. Wendl.). 

Calamus subgenus Oncocalamus G. Mann & H.Wendl. in Trans. Linn. Soc. (London) 

24:436 (1864). 

Clustering, spiny, moderate to high-climbing, pleonanthic, monoecious rattan palms. 

Stem circular in cross section, with short to medium internodes; sucker shoots 

axillary. Leaves pinnate, strongly bifid in juveniles, with a terminal cirrus; sheath 

strictly tubular, bearing scattered, brown or black, bulbous-based triangular, brittle 

spines, sometimes becoming bare, and scattered, thin, white, caducous indumentum; 

ocrea conspicuous, tightly sheathing, neatly horizontally truncate, lobed or somewhat 

saddle-shaped, armed as the sheath, spines often concentrated on ocrea margin; knee 

absent, although rounded horizontal swelling visible at the base of the leaf in some 

species; leaves sessile, or with a very short flattened petiole; rachis unarmed or 

sparsely to profusely armed on the underside; cirrus bearing reflexed acanthophylls; 

elaminate rachis co mmon on lower part of stems, bearing equidistant, alternate to 

opposite acanthophylls; spear leaf deep orange to bright crimson to light green; 

leaflets few to numerous, usually single-fold, sometimes with 2-4 folds, entire, acute, 

linear, lanceolate or sigmoid, regularly arranged, usually armed along the thickened 

margins with robust spines, less so distally, midribs evident, other large veins rather 

distant, transverse veinlets conspicuous, proximal few leaflets smaller than the rest, 

often erect, vertical to rachis and stiffly swept back across stem or arching and 

somewhat pendulous. Inflorescences produced in axils; peduncle enclosed within the 

leaf sheath and emerging from its mouth, hemisperical or flattened and rectangular in 

cross-section; prophyll tubular, tightly sheathing, 2-keeled, 2-lobed at its tip, much 

shorter than the sheath; peduncular bracts c.4, ± distichous, tightly sheathing at first, 

later splitting longitudinally, each with a short triangular or straight lobe; rachis longer 

than the peduncle; rachis bracts like the peduncular, rather close; rachillae pendulous 

with a basal 2-keeled tubular prophyll and numerous distichous, short, tubular, 

somewhat inflated, striate bracts, each enclosing a flower cluster, eventually 

129

longitudinally splitting and tattering post anthesis; flower cluster partially covered by 

a tubular 2-keeled prophyll and consisting of 5, 7, 9 or 11 flowers arranged in a group; 

1-3 pistillate flowers in the centre subtended by 2 lateral cincinni of 0-2 pistillate and 

2-4 staminate flowers, each flower, apart from the central pistillate, bearing an open, 

spathulate, 2-keeled, prophyllar bracteole. Staminate flowers symmetrical; calyx 

membranous, striate basally, stalked, tubular, with 3, short, triangular, apiculate lobes; 

corolla enclosed or only slightly exceeding the calyx, divided almost to the base into 

3, elongate, striate, valvate petals; stamens 6, filaments united to form a thick fleshy, 

androecial tube, free from the corolla, tipped with 6 shallow lobes, bearing pendulous, 

rounded, latrorse anthers on the inside; pollen elliptic, monosulcate, with scabrate, 

tectate exine; pistillode very narrow, conical, slightly exceeding the androecial tube. 

Pistillate flowers superficially very similar to the staminate except slightly broader; 

calyx and corolla similar; staminodial tube bearing tiny empty anthers; gynoecium 

tricarpellate, triovulate, ± ellipsoidal, covered in reflexed scales, style long, narrow, 3angled; 

ovule basally attached, anatropous. Fruit ± spherical, stigmatic remains 

minute, conical; epicarp covered in vertical rows of rather thin reflexed scales, 

mesocarp very thin, almost obsolescent at maturity, endocarp not differentiated. Seed 

single, ± rounded, smooth or warty, basally attached with an oval hilum, covered with 

a thin, papery, sweet sarcotesta; endosperm homogenous, laterally penetrated by a 

smooth-margined mass of inner seed coat; embryo lateral opposite the intrusion. 

Germination adjacent-ligular; eophyll bifid, petiole of seedling dull reddish pink. 

Distribution 

Oncocalamus has a distinct Guinea-Congolian distribution and ranges from SE 

Nigeria to northern Angola, predominantly in coastal forest. 

Notes 

Recent collections have provided considerable insight into the life history and 

morphology of this intriguing genus. Although recent literature has stated that 

Oncocalamus is represented by only one, very variable, species, O. mannii, there are 

in fact four species present in Africa. A new species, O. tuleyi, previously assigned to 

O. mannii, is also described. 

130

Key to the species of Oncocalamus 

Moderate to robust canes; stems with sheaths >20 mm in diameter; ocrea truncate with 

± conspicuous 1.5-2.5 cm long rounded lobe; leaves, including cirrus, >1 m long, 

leaflets linear-lanceolate or only very mildly sigmoid, composed of a single fold: 

Stems with sheaths 30 mm in diameter, unarmed or moderately to 

profusely armed, with visible rounded swelling beneath leaf; leaf with 

>35 pairs of leaflets on each side of the rachis; inflorescence >1 m 

long; flower cluster with constant 3 pistillate flowers in each, seeds 

smooth: 

Leaf sheath moderately to profusely armed, ocrea almost horizontal, 

truncate, with small (8 cm long; rachillae deep crimson; north 

of the Sanaga River: O. tuleyi 

Slender canes; stems with sheaths

O. mannii (H. Wendl.) H. Wendl. 

Gustav Mann (1836-1916), German botanist and horticulturist 

H. Wendl. in Kerchove, Les Palmiers 244: (1878); Drude in Engl. Bot. Jarbh. 21: 111 

(1895); C.H. Wright in F.T.A. 7:111 (1902); Becc. in Webbia 3: 265 (1910); Baudon 

in Rev. de Bot. Appl. 4: 595 (1924); Hédin in Rev. de Bot. Appl. 9: 503 (1929); 

Guinea-Lopez in Ensayo Geobot. de la Guinea Cont. Espanola 244: (1942); Toml. in 

Principes 6: 100 (1962); Letouzey in Adansonia 2(18): 314 (1978); Morakinyo in 

Principes 39(4): 208 (1994); Tuley in Palms of Africa 83: (1995); Aedo et al. Bases 

Docs. Fl. de Guinea Ecuatorial 375: (1999). 

Calamus (subgen. Oncocalamus) mannii H. Wendl. in Trans. Linn. Soc. 24: 436 

(1864)Type: Gabon, Gaboon River, Mann 1044 (holotype K!). 

Oncocalamus acanthocnemis Drude in Engl. Bot. Jahrb. 11: 133 (1895); Durand & 

Durand in Fl. Cong. 1: 585 (1909); Holland in Kew Bull. 9: 727 (1922); Hutch. in 

F.W.T.A. 2: 391 (1936); Dalziel in App. to F.W.T.A. 508: (1937); Guinea-Lopez in 

Ensayo Geobot. de la Guinea Cont. Espanola 244: (1942); Letouzey in Adansonia 

2(18): 314 (1978); Aedo et al. Bases Docs. Fl. de Guinea Ecuatorial 375: (1999); 

synon. nov. Type: Gabon, Büttner 527 (holotype B†). 

Oncocalamus phaeobalanus Burr. in Notizbl. Bot. Gart. Mus. Berlin-Dahlem 15: 748 

(1942); Letouzey in Adansonia 2(18): 314 (1978); synon. nov. Type: Cameroon, 

Ebolowa, Mildbraed 5458 (holotype B†; isotype HBG†). 

Calamus niger (non Willd.) Braun & Schum. in Mitth. Deutsch. Schutzgeb. II:147 

(1889) ex Wright in F.T.A. 8: 109 (1902); Hedin in Rev. de Bot. Appl. 19: 502 

(1929); synon. nov. Type: Cameroon, Braun s.n. (holotype B†). 

Clustering slender to moderate palm climbing to 15-30 m. Stems without sheaths 8-16 

mm in diameter, with 12-28 mm; internodes 12-18 cm long. Leaf sheath 

longitudinally striate, dark green, tan, often dull crimson brown on young sheaths, 

132

moderately to sparsely armed with brown-black spines, concentrated and persistent on 

the ocrea, spines often sloughing off elsewhere on sheath to leave conspicuous, raised, 

circular, blister-like scars; thin, white caducous indumentum present on mature 

sheaths, particularly dense on young sheaths and juvenile individuals; ocrea ± truncate 

or saddle-shaped with a 0.5-1.8 cm high rounded lobe opposite the rachis, armed as 

the leaf sheath, spines concentrated at ocrea margin, extending ± 2 cm. Spear leaf 

deep orange to bright crimson. Juvenile stems up to 6 m long, with sheaths,

narrow rounded depression below, covered with regular polygonal depressions, giving 

a distinctly warty appearance; sarcotesta white, thin (

Distribution 

O. mannii is restricted from southern Cameroon to Gabon. 

Figure 29. Distribution of O. mannii (H. Wendl.) H. Wendl. 


O. mannii is common in open areas, roadside and forest gaps. This species responds 

extremely well to selective logging and is a common component of regrowth 

vegetation. 

Notes 

This species is separated from related taxa by its slender stems and the rather variable, 

and somewhat complicated, flower structure. The presence of a warty seed coat is also 

highly distinctive. 


CAMEROON: Asonganyi 279, Bissombo, 59km SE Akono-linga (03.17N:12.28E) sterile, June 12, 

1981 (YA!); Bililong & Bullock 351, Campo Reserve, Sud Province (02.14N:09.54E) juvenile, s.d. 

135

(K!); Dinklage 1154, Grand Batanga, (02.23N:09.50E) juvenile, February 18, 1891 (HBG!); Letouzey 

11776, NE of Mintom II (02.30N:13.30E) juvenile, January 3, 1973 (YA!); Letouzey 15317, 40km S of 

Kribi (02.28N:09.53E) sterile, December 7, 1979 (YA!); Sunderland 1765, 15km from Kribi on Campo 

road (02.34N:09.50E) sterile, December 1, 1996 (K!, YA!, WAG!); Sunderland 1768, 15km from Kribi 

on Campo road (02.34N:09.50E) Fr., December 1, 1996 (K!, YA!, WAG!); Sunderland 1769, Kribi - 

Ebolowa road, 30km west of Kribi (02.39N:10.09E) Fr., December 1, 1996 (K!, YA!, BH!); 

Sunderland 1790, 15km from Kribi on Campo road (02.34N:09.50E) Fr., March 5, 1997 (K!, YA!, 

BH!, MO!, BR!); Sunderland 1887, 30km south of Kribi (02.48N:09.43E) Fl. & Fr., November 28, 

1997 (K!, YA!, BH!, MO!, WAG!); Sunderland 1888, 30km south of Kribi (02.48N:09.43E) sterile, 

November 28, 1997 (K!, YA!); Sunderland 1929, Kribi to Campo road: 40km south of Kribi 

(02.48N:09.53E) Fr., October 12, 1998 (K!, YA!); van Gemerden BJ, Lolodorf (03.05N:10.25E) sterile, 

s.d. (K!); EQUATORIAL GUINEA: Sunderland 1793, Near village of Etembue (01.16N:09.26E) Fl. 

& Fr., March 13, 1997 (K!, EG!, NY!); Sunderland 1796, 2km north of Ayemeken village 

(02.10N:10.03E) Fr., March 13, 1997 (K!, EG!, NY!); Sunderland 1900, 10km south of Bata 

(01.45N:09.43E) sterile, March 20, 1998 (K!, EG!, WAG!); Sunderland 1902, 10km south of Bata 

(01.45N:09.43E) juvenile, March 20, 1998 (K!, EG!, WAG!); Sunderland 1903, 10km south of Bata 

(01.45N:09.43E) sterile, March 20, 1998 (K!, EG!); Sunderland 1908, Near village of Etembue 

(01.16N:09.26E) Fl., March 28, 1998 (K!, EG!, WAG!); Sunderland 1916, 20km from Sandje along 

Cogo road (01.30N:09.40E) Fl., April 7, 1998 (K!, EG!, NY!); Sunderland 1919, 2km WSW of village 

of Basilé (01.10N:09.50E) juvenile, April 7, 1998 (K!, EG!); Sunderland 1921, On road to Monte Mitra 

(01.12N:09.59E) Fl., April 7, 1998 (K!, EG!); Sunderland 1923, Near village of Etembue 

(01.16N:09.26E) Fr., March 28, 1998 (K!, EG!, WAG!); Tessmann 1, sterile, s.d. (FI!); CONGO: Hens 

170, Belabo (02.09N:16.04E) sterile, May 8, 1889 (K!); GABON: Klaine s.n., Libreville 

(00.35N:09.22E) sterile, October 1893 (FI!); Mann 1044, Gaboon River (00.19N:09.29E) Fl., July 1861 

(K!); Mann 1044a, Gaboon River (00.19N:09.29E) sterile, s.d. (K!); Rietsma 2151, NNW Libreville 

(00.36N:09.22E) Fl., April 25, 1986 (LBR!, WAG!) 

_____________________________________ 

O. macrospathus Burr. 

(Latin) large bracts on inflorescence 

Burr. in Notizbl. Bot. Gart. Mus. Berlin-Dahlem 15:749 (1942); Type: Angola, Luali, 

Gossweiler 9092 (holotype B†: isotype K!). 

Clustering robust palm climbing to 20-35 m. Stems without sheaths, 18-30 mm in 

diameter, with 28-40 mm; internodes 18-24 cm long. Leaf sheaths very lightly striate, 

light moderately to profusely armed with upward-pointing brown spines; armature 

136

often concentrated on the sheath apex and often sloughing off at base leaving very 

feint circular scars; profuse white caducous indumentum present on mature and 

juvenile sheaths; ocrea ± horizontally truncate, somewhat striate, sometimes with a 

very slight (0.5-1 cm) rounded lobe abaxial to the leaf, armed as the sheath, although 

spines often concentrated at apex, extending up to 3 cm. Spear leaf dull yellow to 

bright green. Juvenile sheaths, armed as the mature sheaths only spines more profuse; 

leaves bifid, soon becoming pinnate; elaminate rachis up to 2 m long. Leaves on 

mature stems sessile, or with short (

Figure 30. Oncocalamus macrospathus Burr. 

Sunderland 1913; a. Stem x ½ (1/3); b. Leaflets x ½ (1/3); c. Inflorescence x 1/3 (2/9); d. Portion of 

immature rachilla x 1 ½ (1); e. Flower cluster diagram x 6 (4); f. Pistillate flower with prophyllar bract 

x 4 ½ (3); g. Pistillate flower section x 12 (8); h. Staminate flower section x 12 (8); i. Fruit on rachilla x 

1 2/3); j. Fruit x 1 ½ (1); k. Seed x 1 ½ (1). Drawn by Lucy T. Smith. 

138

Distribution 

O. macrospathus is distributed from Cameroon, south of the Sanaga River, to Cabinda 

(Angola). This species is more commonly encountered in coastal forest although it 

also occurs in the lowland riverine forests of the Congo Basin. 

Figure 31. Distribution of O. macrospathus Burr. 


This species is commonly found in forest margins, tree-fall gaps and other open areas 

and is particularly common in seasonally-inundated forest and alongside water 

courses. 

Notes 

Although previously assigned to O. mannii by Morakinyo (1995) and Tuley (1995), 

recent collections have provided further morphological evidence that this is a distinct 

species. In particular, the presence of the constant 3-central pistillate flowers separates 

this species from O. mannii, as does the smooth seed coat. 

139


CAMEROON: Lejoly 86/1005, 3km S of L'Olonou (02.46N:12.02E), Fr., December 16, 1986 (BR!); 

Letouzey 4556, 35 km S of Bengbis (03.13N:12.28E), Fr., March 17, 1962 (YA!); Letouzey 

11889, Mintom I (02.03N:13.30E) Fl., January 23, 1973 (K!, YA!); EQUATORIAL GUINEA: 

Sunderland 1913, Akonibe to Evinayong road (01.21N:10.43E) Fl. & Fr., April 2, 1998 (K!, EG!, BH!, 

WAG!); Sunderland 1914, Akonibe to Evinayong road (01.21N:10.43E) sterile, April 2, 1998 (K!, EG!, 

WAG!); GABON: Breteler et al. 10957, 5-30km NNW of Ndjale (00.05S:10.45E) Fl., April 21, 1992 

(WAG!); Dibata & Mbouissou 958, Mission Otouma (00.13N:10.56E) Fr., February 10, 1992 (BR!); 

Louis et al. 1350, 32km SE of Sindera (01.14S:10.53E) Fr., July 13, 1983 (K!, WAG!, BR!, HBG); 

Reitsma 1340, Between Cap Santa Clara and Cap Esterias (00.34N:09.22E) Fr., August 15, 1985 

(WAG!); Wieringa 466, 0.5km SE of Tchimbelé (00.36N:10.24E) Fr., January 25, 1990 (WAG!); DR 

CONGO: Couteaux 502, Eala (00.03N:18.18E) sterile, October 29, 1938 (BR!); Evrard 2077, Isangi 

(01.02N:23.41E) Fl. & Fr., December 11, 1956 (BR!); Gossweiler H1039/24, Port Congo, Fr., April 28, 

1924 (K!); ANGOLA: Gossweiler 9092, Mayombe, Luali, (05.00S:12.25E) Fl. & Fr., s.d. (K!) 

_____________________________________ 

O. tuleyi Sunderland sp. nov. 

affinis O. mannii (H. Wendl.) H. Wendl. sed vagina robustus (25-45mm diametrum) 

non tenuis (12-23mm), ramis inflorescentia 1.2-1.8 m longis (non 0.8-1.0 m), rhachis 

bracteatus 13-15 cm longus (non 2.4-4.0 cm), floribus 9-11 (non 5-11), 3 (non 2-5) 

medii foemineis et 6-8 (non 4-6) masculis anterioribus, semen testa laevis non 

tuberculate. Type: Cameroon, Ossing, near Mamfe, Sunderland 1939 (holotype K!; 

iso-types SCA! NY!, MO!). 

O. mannii sensu Holland in Kew Bull. 9: 727 (1922); Russell in F.W.T.A. 2(3): 167 

(1968); Morakinyo in Principes 39(4): 206 (1994) partim non H. Wendl.; Tuley in 

Palms of Africa, 83: (1995) partim non H. Wendl.; Cable & M. Cheek in Pl. of Mt. 

Cam. 179: (1998). 

Clustering robust palm climbing to 30m, rarely to 50 m. Stems without sheaths 13-22 

mm in diameter, with, 25-45 mm; internodes 14-25 cm long, commonly 15-18 cm. 

Leaf sheath lightly striate, light brown to mid-green, very sparsely and patchily armed 

with dark brown to glaucous black spines, often concentrated on the ocrea; sheaths 

often becoming bare with spines sloughing to leave raised, linear blister-like scars; 

thin white caducous indumentum present on young sheaths, absent on mature sheaths; 

ocrea saddle-shaped, with 1.5-2 cm. rounded lobe abaxial to the leaf, armed as the leaf 

140

sheath, spines concentrated on margin, extending for ± 2.5 cm; knee absent, although 

conspicuous horizontal rounded swelling visible beneath leaf. Spear leaf dull reddish 

brown, becoming green. Juvenile stems up to 10 m, with sheaths

Figure 32. Oncocalamus tuleyi Sunderland 

Sunderland 1939; a. Stem x ½ (1/3); b. Leaflets x ½ (1/3): Sunderland 1746; c. Juvenile leaf x ¼ (1/6): 

Sunderland 1939; d. Inflorescence x 1/3 (2/9); e. Portion of immature rachilla x 1 ½ (1); f. Flower 

cluster diagram x 6 (4); g. Staminate flower x 7 ½ (5); h. Pistallate flower x 7 ½ (5): Sunderland 1761; 

i. Fruit x 1 ½ (1); j. Seed x 1 ½ (1). Drawn by Lucy T. Smith. 

142

Distribution 

This species is restricted to coastal forest from SE Nigeria to SW Cameroon, north of 

the Sanaga River and is allopatric with O. mannii. 

Figure 33. Distribution of O. tuleyi Sunderland 

Habitat 

O. tuleyi occurs at the forest edge, adjacent to open areas, and in gap regrowth 

vegetation in forest. This species is an early coloniser of disturbed land and as such is 

a characteristic feature of roadside vegetation in logged forest. 

Notes 

Although previously assigned to O. mannii, this species is clearly sufficiently 

morphologically distinct to warrant designation as a separate species and O tuleyi is 

characterised by very robust stems, large rachis bracts on the inflorescence, a uniform 

flower cluster arrangement and a smooth seed coat. 

143


NIGERIA: Morakinyo 1002, Cross River National Park (05.15N:08.42E) sterile, August 16, 1993 

(K!); Tuley 1078, Calabar to Ikot Opora road (05.00N:08.12E) sterile, December 10, 1964 (K!); 

CAMEROON: Dransfield 7007, Mile 48 Buea-Kumba road (05.02N:09.24E) sterile, June 28, 1991 

(K!, SCA!); Dransfield 7476, 8km S of Nguti (05.02N:09.24E) Fl., November 26, 1997 (K!); Gartlan 

39, Southern Bakundu Forest Reserve (04.46N:09.29E) Fr., November 11, 1968 (K!); Sunderland 

1705, Southern Bakundu Forest Reserve (04.46N:09.29E) sterile, November 8, 1995 (K!, SCA!, BR!); 

Sunderland 1706, Southern Bakundu Forest Reserve (04.46N:09.29E) sterile, November 8, 1995 (K!, 

SCA!, BH!); Sunderland 1731, Rumpi Hills Forest Reserve (04.54N:09.20E) seedling, May 19, 1996 

(K!, SCA!, NY!); Sunderland 1733, Rumpi Hills Forest Reserve (04.54N:09.20E) sterile, May 19, 1996 

(K!, SCA!, BH!); Sunderland 1739, Rumpi Hills Forest Reserve (04.54N:09.20E) sterile, May 19, 1996 

(K!, SCA!); Sunderland 1743, Rumpi Hills Forest Reserve (04.54N:09.20E) sterile, May 19, 1996 (K!, 

SCA!, WAG!); Sunderland 1746, Rumpi Hills Forest Reserve (04.54N:09.20E) juvenile, May 19, 1996 

(K!, SCA!, NY!, MO!); Sunderland 1756; Limbe - Kumbe road: Mile 40 (04.23N:09.26E) juvenile, 

November 11, 1996 (K!, SCA!, NY!, WAG!); Sunderland 1759, Limbe - Kumbe road: Mile 40 

(04.23N:09.26E) sterile, November 11, 1996 (K!, SCA!, WAG!); Sunderland 1761, Southern Bakundu 

Forest Reserve (04.46N:09.29E) Fr., November 25, 1996 (K!, SCA!, BR!); Sunderland 1939, 

Mamfe to Ossing road, 15km south of Mamfe (05.38N:09.17E) Fl., November 20, 1998 (K!, SCA!, 

NY!); Sunderland 2056, Takamanda Forest Reserve (06.08N:09.16E) sterile, January 17, 1999 (K!, 

SCA!) ; Thomas 9732, Idenau (04.16N:09.01E) Fr., September 10, 1993 (K!, SCA!); Thomas s.n. 

Korup National Park (04.55N:08.50E) sterile, s.d. (SCA!) 

_____________________________________ 

O. wrightianus Hutch. 

Charles Henry Wright (1864-1941), British botanist 

Hutch. in F.W.T.A. 2: 391 (1936); Jeffreys in Nig. Field 25(1): 42 (1960); Hutch. in 

Kew. Bull. 27:181 (1963); Russell in F.W.T.A. 2(3): 167 (1968); Profizi in RIC Bull. 

5(1): 2 (1986); Type: Nigeria, Lagos, Barter 2220 (holotype K!). 

sensu O. mannii H.A. Burkhill in Useful Pl. of W. Trop. Afri. 4:376 (1997); Dalziel in 

Hutch. & Dalziel, App. to F.W.T.A. 508: (1937). 

Clustering (?) slender palm climbing to 10 m. Stems without sheaths 6-10 mm in 

diameter, with 8-15 mm; internodes ±15 cm long. Leaf sheaths striate, moderately to 

sparsely armed with caducous black spines, particularly concentrated and persistent on 

144

ocrea, often sloughing off to leave raised, triangular, blister-like scars, sheath 

sometimes becoming bare at base; white caducous indumentum present on mature 

sheaths, profuse on young sheaths; ocrea horizontally truncate, without conspicuous 

lobe, extending to ±3 cm. Leaves sessile, or with a very short (

Figure 35. Oncocalamus wrightianus Hutch. 

Aufsess 430; a. Stem and leaf section x ¾ (1/2). Drawn by Lucy T. Smith. 

146

Notes 

O. wrightianus is distinct from all the other species in this genus by its slender nature 

and relatively broad, sigmoid leaflets. This species was recently reduced to synonymy 

by Tuley (1995), who suggested that this species represented a juvenile form of O. 

mannii. However, this seems highly unlikely as the juvenile leaves of the members of 

this genus are strongly bifid. More extensive field collections, particularly those of 

fertile material, are needed to elaborate on this species. 


BENIN: Aufsess 430, Adjarra (06.32N:05.52E) sterile, December 6, 1988 (K!); NIGERIA: Barter 

2220, Lagos (06.28N:03.20E) sterile, s.d. (K!); Jones & Onochie 17416, Sunmoge to Oshu road 

(06.40N:04.18E) sterile, April 10, 1946 (K!); Miller 18, Lagos (06.28N:03.20E) sterile, s.d., (K!) 

Imperfectly-known taxon 

_____________________________________ 

O. djodu De Wild. 

De Wild. in Bull. Jar. Bot. Brux. V: 146 (1916); Type: DR Congo, Nannan, 65 (BR!). 

This species was described by de Wildeman from, by his own admission, very scanty 

material. The majority of the collections held in BR are mostly of juvenile material 

and the type specimen contains little more than a section of inflorescence and one 

complete leaflet. Although de Wildeman’s original description states that this species 

is characterised by the possession of 3 central pistillate flowers, this is hard to qualify 

give the poor nature of the material. Nonetheless, it is certainly distinct from O. 

macrospathus. Further collections in the central and coastal regions of DR Congo 

would elaborate on this taxon further. 

Indet. Oncocalamus 

_____________________________________ 

NIGERIA: Commonwealth Imperial Institute, s.n., Benin district (05.59N:06.07E) seedling, March 21, 

1906 (K!); Tuley 603, Ojo road (08.19N:04.14E) juvenile, May 4, 1964 (K!); CAMEROON: Harris 

147

3769, Onge (04.21N:08.57E) seedling, October 11, 1993 (K!, SCA!); Sunderland 1716, Kumba to 

Mamfe road (05.02N:09.24E) sterile, December 1, 1995 (K!, SCA!); Sunderland 1718, 30km north of 

Mamfe (05.58N:09.20E) sterile, December 2, 1995 (K!, SCA!, MO!); EQUATORIAL GUINEA: 

Sunderland 1871, Ndote Forest Reserve (01.20N: 09.28E) juvenile, September 13, 1997 (K!, EG!, 

WAG!); DR CONGO: Leonard 817, Eala (00.03N:18.18E) fruit only, October 12, 1946 (BR!); 

Leonard 935, Eala (00.03N:18.18E) sterile, October 30, 1946 (BR!); ANGOLA: Gossweiler 7537, 

Mayombe, Luali (05.00S:12.25E) sterile, October 26, 1918 (K!) 

148

CALAMUS L. 

(Greek = a reed) 

L. in Sp. P1. 325: (1753) & Gen. P1., ed. 5:152 (1754) Type: C. rotang L. 

Rotanga Boehmer in Def. Gen Pl. Ed.3:395 (1760) 

Rotang Adanson in Fam. des Pl. 2(24):599 (1763) 

Palmijuncus Kuntze in Rev. Gen. Pl. 2:731 (1891) 

Zalaccella Becc. in Ann. Roy. Bot. Gard. Calc. 11(1): (1908) & app. (1913). Type: Z. 

harmandii (Pierre ex Becc.) Becc. (= Calamus harmandii Pierre ex Becc.) 

Schizospatha Furt. in Gar. Bull. Sing. 14:525 (1955): Type: S. setigera (Burr.) Furt. (= 

Calamus setiger Burr.) 

Cornera Furtado in Gar. Bull. Sing. 14:518 (1955). Type: C. pycnocarpa Furt. (= 

Calamus pycnocarpus (Furt.) J. Dransf.) 

Solitary or clustering, acaulescent to high-climbing pleonanthic dioecious palms. 

Stems very slender, only a few mm in diameter, to robust (>15 mm), in Africa 

moderate, 10-35 mm in diameter, branching sympodially at the base. Leaf-sheaths 

tightly enclosing the stem, variously armed with spines and spiculae or unarmed, often 

covered with indumentum, often continued into an ocrea. Mature leaf of two kinds; 

terminating either in a long barbed whip (cirrus) or without such a whip (African and 

some Asiatic species), species without a cirrus normally but not always bearing a 

similar barbed whip (flagellum), adnate to the leaf-sheath at the base, equivalent to a 

modified sterile inflorescence; petiole prominent or absent, variously armed with 

spines and hooks; rachis usually armed with reflexed hooks; leaflets narrow to broad 

or rhomboid, single-fold, arranged regularly or irregularly on either side of the rachis 

or variously clustered, fanned or paired, variously hairy, scaly or spiny. Inflorescence 

axillary, with the base of the peduncle adnate to the internode and sheath of the 

following leaf, hence appearing in a non-axillary position, branching to 2-3 orders, 

with or without a long terminal flagellum; bracts variously armed, tubular, tightly 

sheathing, very rarely splitting, sometimes with an expanded limb, never caducous, 

though rarely tattering and decaying before fruiting; prophyll usually 2-keeled and 

empty; other bracts on main axis subtending close to very distant partial inflorescences; 

partial inflorescences bearing bracts subtending rachillae; rachillae usually 

149

with approximate tubular bracts, each, except for the one or more basal-most, subtending 

a flower or flower group; in male inflorescence flowers solitary, borne together 

with a bracteole (‘involucre’); in female inflorescence flowers borne in pairs, a sterile 

male (acolyte) together with a fertile female and 2 bracteoles (‘involucrophore’ and 

‘involucre’). Male flower symmetrical; calyx tubular, 3-lobed; corolla 3-lobed, 

divisions almost reaching the base; stamens 6, epipetalous, with free filaments; 

pistillode minute or absent. Sterile male flower as the fertile male but anthers empty. 

Female flower with calyx and corolla + as in the male flower; staminodes 6; ovary 

covered in vertical rows of reflexed scales and tipped with 3 stigmas; locules 3, 

incomplete, each with a single ovule, normally only one ovule developing. Fruit 

variously shaped, tipped with the remains of the stigma, with the calyx and corolla 

persistent below, covered in vertical rows of reflexed scales. Seed with thick or thin, 

sweet or sour or very astringent sarcotesta and variously shaped hard diaspore; 

endosperm homogeneous or ruminate; embryo basal or lateral. Germination adjacentligular; 

eophyll bifid or pinnate. 

Around 350 species distributed from Africa, India to Eastern Asia, Malesia, to 

Australia and Fiji with the greatest abundance and diversity occurring in the 

archipelagoes of Malesia. Represented in Africa by a single, very variable, widely 

distributed species. Important economically as the source of much of the rattan for the 

cane industry, particularly in SE Asia. 

C. deërratus G. Mann & H. Wendl. 

(Latin) “to go astray” refers to the habit of this species to form expansive clumps 

G. Mann & H. Wendl. in Trans. Linn. Soc. 24: 429 (1864); Drude in Engl. Bot. Jarbh. 

5: 130 (1895); Cummins in Kew Bull. 137: 80 (1898); C.H. Wright in F.T.A. 8: 108 

(1901); Becc. Ann. Roy. Bot. Gard. Calc. 11(1): 151 (1908); Milbr. in Notiz. Bot. Gar. 

Dah. App. 27: 15 (1913); Unwin in W. Afr. For. 240 (1920); Holland in Kew Bull. 9: 

727 (1922); Hédin in Rev. de Bot. Appl. 19: 503 (1929); Hutch. in F.W.T.A. 2: 390 

(1936); Dalziel in App. F.W.T.A. 497: (1937); Burr. in der Tropenfl. 42(5): 204 

(1939); Guinea-Lopez in Ensayo Geobot. de la Guinea Cont. Espanola 244: (1946); 

Irvine in Econ. Bot. 6(23): 31 (1952); Berhaut in Fl. du Sénégal 1: 211 (1954); A. 

150

Robyns & Tournay in Fl. du Parc Nat. Albert 3: 297 (1955); Fosberg in Principes 4: 

129 (1960); Irvine in Woody Plants of Ghana 775: (1961); Toml. in Principes 6: 96 

(1962); Russell in F.W.T.A. 2(3): 166 (1968); Moore in Principes 15: 113 (1971); 

Letouzey in Adan. 18(3): 314 (1978); Hall & Swaine in For. Veg. Ghana 139: (1981); 

Johnson in Principes 28(4): 161 (1984); Profizi in RIC Bull. 5(1): 2 (1986); J. Dransf. 

in F.T.E.A. (Palmae) 43: (1986); Letouzey in Man. For. Bot. Trop. Afr. 2B: 401 

(1986); Bauchet in Fl. du Sénégal 9: 82 (1988); Hawthorn in Trees of Ghana 225: 

(1990); Morakinyo in Principes 39(4): 199 (1995); Tuley in Palms of Africa 50: 

(1995); White & Abernethy in Guide to Veg. of Lopé Res. Gabon 64: (1997); H.A. 

Burkill in Useful Pl. of W. Trop. Afri. 4: 347 (1997); Aedo et al. in Bases Docs. Fl. de 

Guinea Ecuatorial 375: (1999); Type: Sierra Leone, Bagroo River, Mann 891 & 

Cameroun, Mann 2147 (syntypes K!; Mann 891, iso-syntype FI!). 

C. akimensis Becc. in Ann. Roy. Bot. Gard. Calc. 11(1): 162 (1908); Type: Ghana, 

Akim, Kibbi, Johnson s.n. (holotype FI!; isotype K!). 

C. barteri Drude in Engl. Bot. Jarbh. 5: 134 (1895), pro parte; Becc. in Rec. Bot. Surv. 

Ind. 2:199 (1902); C.H. Wright in F.T.A. 8:109 (1901), pro parte, & in Ann. Roy. Bot. 

Gard. CaIc. 11(1): 154(1908); Baudon in Rev. de Bot. Appl. 4: 595 (1924) Type: 

Nigeria, Onitsha, Barter 110 (holotype K!). 

C. falabensis Becc. in Ann. Roy. Bot. Gard. Calc. 11(1): 157 (1908); Type: Sierra 

Leone, Falaba, Scott-Elliot 4460 (holotype FI!). 

C. heudelotii Drude in Engl. Bot. Jarbh. 5: 134 (1895); Becc. in Ann. Roy. Bot. Gard. 

Calc. 11(1); 155 (1908); Type: Gambia, Heudelot 372 (holotype FI!; isotype K!). 

C. laurentii De Wild. in Ann. Mus. Congo, Bot. 5(1): 97 (1904) de Wild in Miss. 

Laurent 24: (1905); Durand & Durand in Fl. Cong. 1: 584 (1909); Pyneart in Bull. 

Agric. du Congo Belge 2: 547 (1911); Becc. in Ann. Roy. Bot. Gard. Calc. 11(5): 

(1913); De Wild. in Ann. de Mus. Col. de Marseille 3(7): 17 (1919); Renier in Fl. du 

Kwango 81: (1948); Type: DR Congo, Eala, Laurent 126 (holotype BR!; isotype FI!). 

C. leprieurii Becc. in Rec. Bot. Surv. Ind. 2: 200 (1902) & in Ann. Roy. Bot. Gard. 

151

Calc. 11(1): 158 (1908); Type: Gambia, Leprieur 1830 (holotype FI!; isotype P). 

C. perrottetii Becc. in Rec. Bot. Surv. Ind. 2: 200 (1902) & in Ann. Roy. Bot. Gard. 

Calc. 11(1): 160 (1908); Type: Sénégal, Casamance River, Perrottet 1826 (holotype 

FI!; isotype G). 

C. schweinfurthii Becc. in Rec. Bot. Surv. lnd. 2: 200 (1902); Drude in Engl. Bot. 

Jarbh. 5: 131 (1895); Types: Sudan, Mansilli, Schweinfurth 2860 & Nabambisso, 

Schweinfurth 3703 (syntypes B†; iso-syntypes FI! K!). 

C. secundiflorus sensu Schweinf., Beitr. Fl. Aeth.: 291 (1867), non P. Beauv. 

Clustering, slender to moderate palm climbing to 20m, often branching sympodially at 

the base. Stem without sheaths, 1.0-2.8 cm in diameter, with, 1.2-3.5 cm; internodes 

8-20 cm long, more commonly 15-20 cm. Leaf-sheaths varied in armature from 

almost unarmed to densely spiny, with a distinct horizontal, sometimes folded, knee 

below the petiole; spines dark brown or black, triangular, flattened at base, up to 3 cm 

long, clusters of upward pointing spines often concentrated around the leaf-sheath 

mouth to form a conspicuous cleft; mature sheaths with brown or grey indumentum; 

ocrea to up 12 cm long, usually 8-10 cm, dry, papyraceous, tongue-shaped, often 

longitudinally splitting and reflexed, becoming unrecognisable, armed on the margins 

with spines more pale and bristle-like than those on the leaf-sheath, rarely unarmed. 

Leaves ecirrate, up to 1.75m long, usually 1.2-1.5m; petiole to 20 cm long, rounded 

abaxially, concave adaxially, ± 5 mm broad, variously armed with large black spines 

to 3 cm long and small recurved black thorns; rachis triangular in section distally 

armed as the petiole, spines becoming sparse distally; leaflets up to 30 on each side of 

the rachis, sub-equidistant to equidistant proximally, grouped in 3’s to 6’s distally, 

linear-lanceolate, finely acuminate to apiculate at apex, bluntly compact at the base, 

up to 35 cm long by 2 cm broad at the widest point, + concolorous of with slightly 

darker green upper surface, leaflet margins, main vein and secondary nerves bristly 

throughout. Flagellum up to 3.5m long by 4 mm wide at the base, decreasing very 

gradually above, armed with small recurved thorns. Male and female inflorescences 

similar, up to 3.5m long, with 1-4 partial inflorescences and a long terminal sterile 

flagellum; axis and bracts armed throughout with reflexed, solitary or grouped black 

152

prickle-like spines; bracts tightly sheathing, up to 70 cm long with an expanded, 

papyraceous limb ± 5 cm long; partial inflorescences to 40 cm long, with up to 15 or 

more rachillae on each side, subtended by bracts + 2 cm long (1 cm. exposed), with 

mouths + 7 mm wide and with a short triangular limb to 4 mm; rachillae up to 7 cm 

long, arcuate, arranged distichously; bracts distichous, dull brown in colour, ciliatehairy 

around the mouth. Male flowers solitary, distichous, with a minute involucre to 

1 mm long; calyx 4 mm long, tubular for + 3 mm, with 3 short, triangular, striate 

lobes; corolla-lobes to 7 mm long x 2 mm wide, fused at the base for + 1 mm, widely 

diverging at anthesis; stamens to 4 mm long, minutely epipetalous, with filaments up 

to 3 mm long, anthers + 3 mm. long, medifixed; pollen yellow. Sterile male flower 

very similar to fertile male but slightly shorter and narrower. Female flower with 

calyx tubular at first and then splitting as ovary increases in size, lobes + 3 mm long; 

corolla-lobes + 5 x 2 mm, with 6 minutely epipetalous flattened staminodes; ovary ± 5 

mm long by 2.5 mm wide, tipped by 3 stigmas + 1 mm long, markedly recurved at 

anthesis. Fruit at maturity to 1.5 cm, x 1 cm with a short beak up to 2 mm tipped by 

remains of the style, with 17-20 vertical rows of scales. Seed flattened laterally, + 9 x 

8 x 5 mm, with sarcotesta + 1 mm thick when dry; endosperm homogeneous, embryo 

basal. Germination adjacent -ligular; eophyll pinnate. 

153

Figure 36. Calamus deërratus G. Mann & H. Wendl. 

Sunderland 2262; a. Stem x ¾ (1/2), b. Leaflet section x ½ (1/3): Sunderland 1754; c. Inflorescence x 1 

(2/3), d. Male flower x 4 (2 & 2/3), e. Male rachilla x 1 ½ (1): Eggeling 1626; f. Pistillate flower x 4 (2 

2/3): Deighton 1847; g. Fruit x 1 ½ (1), h. Fruit section x 2 ¼ (1 ½). Drawn by Lucy T. Smith. 

154

Distribution 

C. deërratus is the most widely distributed of the rattans of Africa and is distributed 

across the humid forest zone of Africa; from the Gambia and Casamance in Senegal, 

southwards to northern Angola and Zambia and eastwards to southern Sudan and 

Uganda. 

Figure 37. Distribution of C. deërratus 


C. deërratus has a strong preference for swamp and riverine forest, (Ainslie, 1926; 

Foggie, 1941; Ahn, 1961) and is rather less common in areas with high rainfall (Hall 

and Swaine, 1981). As such, this species is relatively rare in the Guineo-Congolian 

forest of Cameroon and Gabon (Richards, 1963; Letouzey, 1978; author pers. obs.). 

This species is more common in drier gallery forest found in the transition zones 

between Sudanian savanna woodland to the north of the Guineo-Congolian forest 

formation, and Zambezian savanna woodland to the south. C. deërratus occurs in 

lowland forest areas in west and central Africa at altitudes

altitude regions of east Africa >1500m. This species is usually found in forest under a 

canopy, but also occurs in open areas where it often forms dense thickets. 

Notes 

Beccari (1908) admitted the close relationship of the African species of Calamus he 

described, and his treatment of the African representatives of this genus, along those 

of Drude (1895) and de Wildeman (1904), was reasonable given the fragmentary and 

often rather poor quality material at their disposal. The fact that Calamus in Africa has 

been the cause of some taxonomic problems has been undoubtedly due to the 

recognition of poorly-defined infraspecific variation. However, from recent 

examination of herbarium specimens and field observations, it is clear that Calamus in 

Africa is represented by a single polymorphic species. 


GAMBIA: Anderson 131, Fr. July 10, 1975 (MO!); Heudelot 372, stam. 1839 (K!); Ingram s.n. sterile 

s.d. (K!); Starin 136, Aboko F.R., Fr., January 20, 1992 (K!); Starin 28, Aboko F.R., Fr., December 

1979 (K!); SÈNÈGAL: Berhaut 877, Sangalkam (14.47N:17.12W) stam. December, 1950 (BR!); 

Vanden-Berghen 1752, Casamance (12.51N:15.17W) sterile, January 4, 1977 (MO!, BR!); Vanden- 

Berghen 4094, Badioure (12.53N:16.08W) stam. November 11, 1980 (BR!); Vanden-Berghen 5264, 

Bouyouye (12.26N:16.44W) pist. July 19, 1982 (BR!); GUINEA-CONAKRY: Chillou 1905, Kouyaya 

(10.25N:12.37W) sterile, March 19, 1940 (K!, BR!); Ory 216, Fula Kunda (13.15N:16.37W) sterile, 

January 25, 1954 (K!); SIERRA LEONE: Deighton 1847, Taisma, Central Province, Fr. July 8, 1930 

(K!); Deighton 2592, Njala (08.06N:10.46W) sterile, January 1, 1933 (K!); Mann 895, Bagroo River 

(07.45N:12.50W) sterile, April 1861 (K!, FI!); Scott-Elliot 4738, Kambia (08.41N:13.03W) sterile, 

January 8, 1892 (K!); Scott-Elliot 5121, Musaia (09.27N:11.25W), sterile, March 10, 1892 (K!); Small 

455, River Seli, Fr., September 5, 1951 (K!); Thomas 2753, Jigaya, stam. September 28, 1914 (K!); 

LIBERIA: Linder 1078, Piatah (07.12N:09.28W) sterile, October 15, 1926 (K!, MO!, WAG!); Linder 

1116, Piatah (07.12N:09.28W) stam., October 17, 1926 (K!, MO!); Linder 1226, Gbanga 

(06.59N:09.28W) sterile, October 24, 1926 (K!, WAG!); Whyte s.n., near Kabatown (06.21N:10.43W) 

stam., April 1904 K!; CÔTE D'IVOIRE: Hepper & Maley 8041, Mont des Dans near Santa 

(08.16N:08.07W) pist., February 3, 1984 (K!); Hepper & Maley 8177, Taï Forest (05.38N:07.08W) 

sterile, February 9, 1984 (K!); Leeuwenberg 2524, 61km N of Sassandra (06.10N: 05.19W) stam., 

January 21, 1959 (WAG!); Leeuwenberg 2882, 18km NW of Sassandra (06.15N:05.00W) stam., 

November 26, 1960 (K!, WAG!); Oldeman 571, 9km ENE of Bereby (04.34N:07.00W) seedling, 

November 9, 1963 (WAG!); Oldeman 589, 3km E of Bereby (04.34N:07.00W) pist., November 5, 

1963 (K!, WAG!); GHANA: Adams 2025, 2m E of Bibiani (06.20N:02.10W) sterile, December 23, 

1953 (GC!); Cummins 128, Ashanti region, sterile, 1895 (K!); Enti 643, Kade Agricultural Station 

156

(06.05N:00.50W) pist., March 17, 1972 (GC!, MO!, BR!); Enti & Hall s.n., Kade Agricultural Station 

(06.05N:00.50W) Fr., May 30, 1970 (GC!); Hall 2846, Anjakal, stam., January 20, 1965 (K!, GC!); 

Johnson 242, Kibbi-Akkim (06.09N:00.35W) Fr., December 13, 1899 (K!); Kinlock 3326, Tarkwa, 

Ndumnfri F.R. (05.10N:02.09W) sterile, February 15, 1934 (KUM!); Kisseadoo 435, Bobiri F.R. 

(06.38N:01.17W) sterile, November 10, 1988 (MO!); Sunderland 2262, Draw River Forest Reserve 

(05.12N:02.20W) sterile, May 26, 1999 (K!, KUM!); Tomlinson s.n., Bobiri F.R. (06.38N:01.17W) 

stam., December 20, 1957 (K!); Tomlinson s.n., Cape Coast F.R. (05.04N:01.30W) stam., December 

15, 1957 (GC!); Tsiforkor s.n., Bunsu (06.15N:00.28W) sterile, November 22, 1995 (K!); Vigne 1868, 

Amentia F.R., E. Region (06.10N:01.58W) pist., March 30, 1930 (K!, KUM!); Vigne 3951, South 

Formango F.R. (06.35N:01.57W) sterile, July 30, 1935 (FHO!, KUM!); BENIN: Aufsess 424, Adjarra 

(06.32N:05.52E) sterile, December 6, 1988 (K!); NIGERIA: Allison 6994, Kabba Province 

(08.08N:06.44N) stam., November 15, 1943 (K!); Ayewoh 3854, Ondo Province, Ifon (06.54N:05.46E) 

sterile, February 24, 1944 (K!); Barter s.n., Onitsha (06.06N:06.48E) Fr., s.d. (K!); Bennett 8, sterile, 

February 24, 1906 (K!); Chapman 5010, Gangumi Forest Reserve, Gongola State (07.16N:11.23E) Fr., 

May 24, 1977 (K!); Chapman 5423, Baissa F.R., Gongola State (07.14N:10.38E) Fr., April 29, 1978 

(K!); Gledhill 923, Akure F.R. (07.12N:05.11E) sterile, April 5, 1968 (K!, WAG!); Imp. Inst. Nigeria 

347, Ahoada (05.03N:06.34E) sterile, February 1, 1936 (K!); Keay 28091, Kouton Kerifi, 

(08.08N:06.44N) Fl., November 6, 1950 (K!); Lowe 4353, Ilaro Forest Reserve, near Abeokuta 

(08.03N:06.06E) sterile, December 14, 1982 (K!); Tuley 846, Lagos (06.28N:03.20E) Fl., May 4, 1964 

(K!); CAMEROON: unknown collector, Bezirk Djah (03.00N:12.40E) stam., March, 1910 (FI!); 

Buschen 3, Ebolowa (02.55N:11.08E) sterile, s.d. (FI!); Dransfield 6999, Mungo River Crossing 

(04.08N:09.31E) stam., June 27, 1991 (K!, SCA!); Dransfield 7000, Mungo River Crossing 

(04.08N:09.31E) sterile juvenile, June 27, 1991 (K!); Lederman 2428, Tibati (06.27N:12.37E) pist., 

January 29, 1909 (FI!); Mann 2147, Cameroon River (04.04N:09.38E) Fl. & Fr., January 1863 (K!, 

FI!); Meijer 15220, Sangmelima (02.55N:11.58E) sterile, March 24, 1981 (K!, MO!, WAG!, YA!); 

Meijer 15288, 9km W of Sangmelima (02.55N:11.58E) sterile, March 26, 1981 (WAG!); Mildbraed 

4190, 21km northeast of Moloundou (02.03N:15.09E) Fl., June 6, 1911 (HBG!); Mildbraed 9548, Buea 

- Douala at Uham, Fl., May 5, 1914 (K!); Raynal 10150, 17km SE Ambam along the river Kye 

(02.23N:11.16E) sterile, March 1, 1963 (YA!); Raynal 10548, Guerima (7km NE de Bafia), gallery 

forest on river Mbam (04.44N:11.13E) stam., March 28, 1963 (YA!); Sunderland 1754, Limbe - 

Douala road at Mungo Bridge (04.08N:09.31E) stam., November 16, 1996 (K!, SCA!, BH!); 

Sunderland 1864, Djoum (02.48N:12.22E) sterile, September 9, 1997 (K!, YA!, WAG!); CENTRAL 

AFRICAN REPUBLIC: Fay 7020, N'Dele-Pata road (08.08N:21.08E) pist., May 30, 1985 (MO!); 

Fay 4381, Manovo-St Floris National Park (09.29N:21.17E) Fr., April 4, 1983 (K!); Harris & Fay 820, 

Ndakan (02.21N:16.09E) sterile, June 1, 1988 (K!); le Testu 3594, Haute-Kotto (04.11N:22.08E) Fr., 

January 17, 1922, (BR!); EQUATORIAL GUINEA: Tessmann 6, Fr., s.d. (FI!); GABON: Klaine 

3246, Environs de Libreville (00.35N:09.22E) Fr., February 18, 1903 (K!); DR CONGO: Claessens 

989, Mangbetu (02.28N:27.22E) sterile, June 1921 (BR!); de Witte 4066, Parc National de l'Upemba 

(09.04S:26.38E) pist., August, 1948 (BR!); Demeuse s.n., sterile, s.d. (BR!); Evrard 1876, Likimi 

(02.49N:20.44E) stam., September 29, 1955 (BR!); Evrard 3933, River Tshuapa en amont de Boende 

157

(00.44S:19.12E) Fr., April 18, 1958 (BR!); Germain 210, Lileko, river Yoko (00.48S:19.34E) stam., 

February 28, 1940 (BR!); Hart 633, Epulu, Ituri forest (01.25M:28.35E) stam., September 8, 1986 

(MO!, BR!); Hoier s.n., Parc National de Albert (00.46S:29.17E) pist., January, 1930 (BR!); Laurent 

981, Limbuku, Fr., March 16, 1906 (BR!); Laurent s.n., Machofa, Fr. December 17, 1898 (BR!); 

Laurent s.n., Sankuru (04.03S:22.32E) pist., November 18, 1903 (BR!); Laurent s.n., stam., 1903 

(BR!); Lejoly 2912, Ubundu (00.26S:25.28E) stam., March 12, 1978 (BR!); Leonard 832, Eala, on 

banks of the Ikelemba (00.03N:18.18E) Fr., October 14, 1946 (K!, WAG!, BR!); Louis 15541, between 

Lisala and Ukaturaka (02.14N:21.33E) Fr. July 13, 1939 (K!, BR!); Louis 16796, Yangambi 

(00.45N:24.26E) Fr., November 17, 1943 (BR!); Luja 234, sterile, s.d. ( BR!); Malaisse 9453, Kando 

(10.49S:25.44E) sterile, February 22, 1978 (WAG!, BR!); Malaisse 11907, between Kyamasumba and 

Buzange, sterile, September 20, 1981 (MO!, WAG!); Malaisse 13889, Kibwe (08.19S:29.04E) stam., 

July 12, 1986 (BR!); Mandango 2977, Ile Mbo near Lubutu (00.44S:26.34E) Fr., May 13, 1981 (BR!); 

Noirfalise 664, Parc National de la Garamba (04.10N:29.28E) Fr., August 4, 1950 (BR!); Robyns 3250, 

Parc National de la Garamba (04.10N:29.28E) sterile, July 25, 1948 (BR!); Sapin s.n., Milangala, 

sterile, Febuary 1910 (BR!); Schweinfurthii 2860, Manselli, sterile, October 1893 (K!); Troupin 296, 

Parc National de la Garamba (04.10N:29.28E) sterile, March 5, 1982 (BR!); Vanderyst 12343, Luenge 

(06.28S:26.14E) seedling, August 1922 (BR!); Vanderyst 21843, seedling s.d. (BR!); ANGOLA: 

Gossweiler 13644, Luachima, NE of Luanda (07.53S:14.05E) pist. & Fr., May 1938 (K!); SUDAN: 

Andrews 1291, Equatoria Province, stam., May 28, 1937 (K!); Jackson 3406, Equatoria Province, 

Yambio (04.34N:28.23E) Fr., 1950 (FHO!); Myers 6757, Mt Ameringi, sterile, May 28, 1937 (K!); 

Myers 11334, Equatoria Province, stam., May 17, 1939 (K!); UGANDA: Dawe 149, Mabira forest 

(00.30N:33.00E) sterile, 1904 (K!); Eggeling 1626, Budongo Forest (07.40N:31.32E) pist., February 

1935 (K!, FHO!); Katende 702, Nyabisabu River, Budongo (07.40N:31.32E) Fl., October 17, 1970 

(K!); Katende 2783, Budongo Forest (07.40N:31.32E) stam., September 17, 1977 (MO!); Poulson 969, 

Budongo Forest (07.40N:31.32E) pist., September 18, 1995 (K!) 

158

MIXED COLLECTIONS 

A number of collections held in herbaria are comprised of material of more than one 

species, although included within a single accession. These collections are not 

included within the exsiccatae for each taxa and have been determined as follows: 

GHANA: Chipp, 127; Yenalim, Ashanti region, stam. inflorescence of C. deërratus & leaflets of 

Laccosperma sp., May 7, 1912 (K!, KUM!); Irvine 5067, Aiyinase (05.18N:01.58W) July 1961, 

juvenile leaflet of E. macrocarpa & leaflets of L. secundiflorum (K!); NIGERIA: Dunstan s.n., 

southern Nigeria, leaflets of L. secundiflorum & C. deërratus, October 27, 1904 (K!); Gentry & Pilz 

32873, Cross River State (05.15N:08.42E) leaflets of O. tuleyi & E. macrocarpa, June 20, 1981 (K!, 

MO!); Otedoh & Tuley 7253, between Warri and Sapele (06.04N:05.29E) sheath & leaflets of L. 

robustum, sheath of E. macrocarpa, August 18, 1972 (K!); CAMEROON: Bruneau 1116, Lolodorf 

(03.05N:10.25E) leaflets of L. robustum & E. macrocarpa, October 31, 1995 (K!); DR CONGO: Gillet 

167, Kipapashi, leaflets of E. haullevilleana & infructescence of L. opacum, 1903 (BR!); Moureau- 

Cheauvard 129, Lac Tumba (00.46S:20.06E) leaflets of L. secundiflorum & flowers of Eremospatha 

sp., November 5, 1957 (BR!); Nannan 66, Ikelemba river, leaflet of Oncocalamus sp. & stems of 

Draceana, August 26, 1914 (K!, BR!) 

159

Figure 47. Eremospatha hookeri, Korup NP, 


Figure 48. E. laurentii, Campo, Cameroon 


160 

Figure 49. E. laurentii, Campo, Cameroon 

(Sunderland, 1805) 

Figure 50. E. laurentii, Cogo, Equatorial 

Guinea (Sunderland 1920)

Figure 51. E. macrocarpa seedling, Rumpi 

Hills, Cameroon (Sunderland 1730) 

Figure 52. E. macrocarpa juvenile, Limbe 

Botanic Garden, Cameroon 

161 

Figure 53. E. macrocarpa ocrea, Limbe 

Botanic Garden, Cameroon 

Figure 54. E. macrocarpa, Mamfe, 

Cameroon (Sunderland 1999)

Figure 55. E. macrocarpa flowers, Mamfe, 


Figure 56. E. macrocarpa fruits, Kribi, 


162 

Figure 57. E. wendlandiana, Southern 

Bakundu, FR, Cameroon (Sunderland 1701) 

Figure 58. E. wendlandiana, showing knee, 

Campo, Cameroon (Sunderland 1927)

Figure 59. E. cuspidata, Etembue, 

Equatorial Guinea (Sunderland 1792) 

Figure 60. E. cuspidata, immature fruits, 

Etembue, Equatorial Guinea (Sunderland 

1909) 

163 

Figure 61. Laccosperma opacum, fruits, 

Nguti, Cameroon (Sunderland 1885) 

Figure 62. L. laeve, fruits, Draw River FR, 

Ghana (Sunderland 2266)

Figure 63. L. acutiflorum, Nguti, Cameroon 

(Sunderland, 1882) 

Figure 64. L. acutiflorum, Campo, 

Cameroon (Sunderland, 1926) 

164 

Figure 65. L. robustum, Campo, Cameroon 


Figure 66. L. robustum fruits, Bata, 

Equatorial Guinea (Sunderland 1791)

Figure 67. L. secundiflorum, Ghana 

(Sunderland (2259) 

Figure 68. L. secundiflorum, Nigeria (Photo: 

Paul Tuley) 

165 

Figure 69. Oncocalamus macrospathus, 

Evinayong, Equatorial Guinea (Sunderland 

1913) 

Figure 70. Oncocalamus macrospathus 

fruits, Evinayong, Equatorial Guinea 

(Sunderland 1913)

Figure 71. O. tuleyi, seedling, Rumpi Hills, 


Figure 72. O. tuleyi, Southern Bakundu FR, 


166 

Figure 73. O. tuleyi sheath, Southern 

Bakundu FR, Cameroon (Sunderland 1706) 

Figure 74. O. tuleyi, Limbe to Kumba road, 

Cameroon (Sunderland 1756)

Figure 75. O. mannii, Ayemaken, Equatorial Guinea (Sunderland 1923) 

Figure 76. O. mannii, inflorescences, Etembue, Equatorial Guinea (Sunderland 1908) 

167

Figure 77. O. mannii, juvenile, Basilé, 

Equatorial Guinea (Sunderland 1919) 

Figure 78. O. mannii, inflorescence, 

Etembue, Equatorial Guinea (Sunderland 

1908) 

168 

Figure 79. Calamus deërratus, Mungo River, 


Figure 80. C. deërratus, male inflorescence, 

Mungo River, Cameroon (Sunderland 1754)

CHAPTER THREE 

RATTAN DIVERSITY AND ABUNDANCE: 

A COMPARISON BETWEEN THREE FOREST SITES IN 


CAMEROON 

Research in Asia has shown that rattan diversity, abundance and distribution, are often 

related to edaphic and climatic factors (Dransfield, 1992b; Siebert, 1993; Nur Supardi 

et al., 1996; Bøgh, 1996; Nur Supardi 1999; Nur Supardi et al., 1999). It is also 

speculated that the rattan diversity of a sample area is closely related to overall 

species diversity (Dransfield, 1979; Dransfield, 1992b). 

In addition to assessments of diversity, the stocking, growth rates and potential 

harvest yield of rattans is crucial in determining levels of sustainable harvesting. For 

this baseline information to be available, rapid methods of assessment and inventory 

that are both accurate as well as economically and logistically feasible, need to be 

developed and implemented. 

In conjunction with these one-off surveys, the establishment of permanent sample 

plots to monitor mortality, growth and recruitment against this baseline, allows 

additional, and more detailed, information to be gathered. Combined, these techniques 

provide information that is crucial for the determination of sustainable levels of 

exploitation for commercially important rattan species. 

This Chapter presents the results of a survey of the rattan resource of three diverse 

protected forest sites in Cameroon. In addition to an assessment of abundance and 

stocking, the relationships between the rattan flora and vegetation type are discussed. 

This diversity and abundance are compared with other sites in Africa and Asia. 

169

3.2 RATTAN INVENTORY 

3.2.1 Introduction 

In the main, forest inventory techniques have been designed for the exploitation of the 

timber resource and, as such, have concentrated on the enumeration of tree species 

(Philip, 1994). However, it is possible to include a wider range of life forms into the 

inventory process and enumeration techniques are being developed to focus on the 

wider “resource base” of an area (Peters, 1996; Peters, 1999). These techniques are 

being used to provide information on a range of commercially important forest 

products, including timber; information that will hopefully lead to more holistic 

models for sustainable exploitation and management of forest resources (van Dijk, 

1995; Wong, 1997; van Dijk, 1999; Sunderland and Tchouto, 1999). 

Rattan inventory has proved to be a somewhat imperfect science. Initial attempts to 

determine stocking and yield have often been thwarted by a poor taxonomic base; it is 

essential to know which species are being enumerated, and which are of commercial 

value. Furthermore, the lack of sampling within the correct parameters has hindered 

the potential application of such surveys. 

The majority of inventories and resource estimates relating to rattan have been 

undertaken in SE Asia (Stockdale and Wright, 1994; Siebert, 1993; Peters, 1996; 

Bøgh, 1996). Until recently, very little was known about the stocking and potential 

yield of the commercially valuable species of rattan in the forests of West and Central 

Africa. Recent work, notably by the Cross River Forestry Project in Nigeria (CRFP, 

1994), the Tropenbos and ECOFAC programme’s in Cameroon (van Dijk, 1995; 

Nzooh, in press) and the Forestry Department in Ghana (Wong, 1997) have attempted 

to address this shortfall. 

3.2.2. Inventory parameters 

3.2.2.1 Introduction 

Quantification and enumeration of non-timber forests is not as straightforward as 

measuring the diameter, and then calculating the basal area of tree species (Philip, 

1994). With non-timber products, in order to develop sustainable harvesting regimes, 

it is essential that the correct parameters are determined prior to the commencement of 

170

an inventory or survey. These parameters depend on the life form of the plant as well 

as the plant part harvested. This is discussed in more detail by Peters (1996; 1999). 

3.2.2.2 Taxonomy 

It is essential to know which species are being enumerated and, in conjunction with 

studies of economic and ethnobotany, which species have potential utility. Without 

reference to a sound taxonomic base, an accurate assessment of the resource being 

enumerated is not possible. Where there is doubt surrounding the identification of a 

species, voucher specimens should be taken. This process need not be undertaken for 

each individual within a sample area, but representative samples can be taken for 

“morpho-species” (de Walt et al., 1999) 1 . In addition to providing crucial information 

about the resource base, knowledge of the taxonomy of the species concerned also 

provides invaluable information with regard to the relationship between the rattan 

flora and the wider vegetation. 

A number of previous inventories in Africa that have included rattan have been 

somewhat constrained by a poor understanding of the taxonomic base (CRSFP, 1994; 

Wong, 1997), or have relied on local nomenclature that is not always congruent with 

western taxonomy (van Dijk, 1995; van Dijk, 1999). Unfortunately, in such cases the 

lack of a rigorous baseline reference means that such surveys have little application 

for the coherent management of the rattan resource. 

3.2.2.3 Stem length vs harvestable length 

Stem length, as well as the number of stems per unit area, are the most crucial 

parameter for determining potential yield in rattans. However, there is some difference 

between total stem length and harvestable stem length (Bøgh, 1996; Sunderland and 

Dransfield, in press). Rattan stems selected for harvesting are those mature stems 

without the lower leaves (i.e. where the leaf sheaths have sloughed off). It is usual that 

only the basal 10-20m is harvested. The upper “green” part of the cane is too soft and 

inflexible for transformation and is often left in the canopy. Hence, to ascertain the 

true yield per hectare of a species, both total and harvestable stem lengths must be 

measured. 

171

As Peters (1996) rightly points out, however, accurately measuring the length of a 

rattan stem that can climb up to 150m into the forest canopy is fraught with logistical 

problems. Actual measurement of stem length is almost impossible and “rough 

estimation” does not provide the accuracy required for inventory purposes (Nur 

supardi et al., 1996). Shim (1989) and Lee (1993), in their studies of Calamus in Asia, 

found that the internode length for most species of rattan is relatively constant. By 

calculating the mean internode length and multiplying by the number of internodes on 

each stem, it is possible to determine the total stem length in a manner that is 

relatively accurate. 

Unfortunately, the mature (and harvestable) cane length is devoid of leaves and hence 

internodes, and cannot be measured in this way. However, as this represents the 

proximal portion of the stem, this portion can be more accurately measured by more 

conventional means such as with a tape or a calibrated stick. In applying a 

combination of these two methods of stem measurement provides an extremely 

accurate means of determining stem length can be achieved. 

3.2.2.4 Plot shape and size 

Considerable discussion has surrounded the determination of plot shape and size with 

regard to forest mensuration and these have been reviewed in the rattan context by 

Stockdale and Wright (1994) and Nur Supardi (1999) and the wider NTFP context, 

including rattan, by Peters (1996). Stockdale and Wright (1994) conclude that 

rectangular plots, oriented parallel to the direction of the slope, are more cost efficient, 

and within the desired level of precision, than square plots. However, Peters (1996) 

concluded that rectangular enumeration plots are prone to errors in boundary 

identification and area estimation, and advocates the use of square, or circular, 

research plots for NTFPs. 

Beyond the fact that larger individual organisms require larger sample plots, there are, 

in general few guidelines that govern the selection of an appropriate plot size for 

vegetation sampling. In the case of rattans, quite large sample sizes are needed if 

1 Individuals clearly of the same (albeit unidentified) species within the sample area. Only a single 

172

clustering species are to be included in the sample. Nur Supardi (1999) found that 

large, square plots were most effective at capturing not only the larger clustering 

individuals, but also up to 80% of the rattan species present within the sample area. 

Bøgh (1996) also found that 1ha square plots were sufficiently large enough to 

capture these large clustering individuals and provide a representative sample of the 

rattan flora of the area in Thailand he was studying. This 1ha square plot methodology 

has been widely implemented for many resource surveys and, if sited correctly, can 

provide a representative and reasonably homogenous sample of variation in forest 

type (Boom, 1987; Prance et al., 1987; Philips and Gentry, 1993; Philips et al., 1994; 

Wong, 1997; Graham et al., 1998; de Walt et al; 1999; Sunderland and Comiskey, 

2000). In this respect, such 1ha plots, when permanently demarcated, are also 

recommended as being suitable for long-term monitoring of vegetation (Alder and 

Synnot, 1992). 

For the purposes of this study, a series of 1ha permanent sample plots (PSPs) were 

established in three protected areas in Cameroon with the intention that they are 

permanently demarcated for long-term research. Aside from the initial baseline 

measurement, which is presented here, these plots will be monitored on a regular 

basis, which will allow, over time, the calculation of growth rates, mortality and 

recruitment. This, in turn, will enable the potential harvest, and sustainable extraction 

rates to be established for each species within the sample sites. 

3.3. RESEARCH SITES 

Three diverse protected areas, the Campo Ma’an Faunal Reserve, the Mokoko River 

Forest Reserve and the Takamanda Forest Reserve, were chosen for this survey. Site 

selection was based on known diversity and climatic variation (based on a review of 

previous studies) and with regard to logistical feasibility. These sites were enumerated 

from February to April 1997 (Campo); October to December 1998 (Takamanda) and 

January to February 1999 (Mokoko). 

representative voucher of such taxa needs to be made. 

173

Figure 81. Map of Cameroon showing protected areas; the study sites are highlighted. Modified 

from Gartlan, 1989). 

3.3.1 Campo Ma’an Faunal Reserve 


The Campo Ma’an Faunal Reserve (2°09’-2°53’N; 9°48’-10°25’E) was created by 

decree on the 19th November 1932 and is situated in the South Province of Cameroon 

(Gartlan, 1989). The Reserve covers an area of 271,160 ha and is bordered to the 

north by the Lobé River, to the south by the Ntem River, which also marks the 

political border with Equatorial Guinea, to the west by the Atlantic Ocean and to the 

east by the Ntem River near to the rapids at Memve’ele (Sunderland et al., 1997). 

174

3.3.1.2 Climate 

The Campo Reserve has a typical equatorial climate with four distinct seasons: a long 

dry season from November to February, a mini rainy season from March to May, a 

shorter dry period from June to August followed by a protracted period of rain from 

mid-August to November. The average annual rainfall is 2,820mm and the mean 

annual temperature is 26.8°C. 

3.3.1.3 Topography, geology and soil type 

The Reserve is situated on inferior pre-Cambrian formations with varying relief. In the 

west of the area, the topography is relatively and consistently plain-like except for the 

Massif des Mamelles, which rises to 323m altitude. The east of the Reserve however, 

is quite mountainous with altitudes varying between 400-950m. Mount Nkolenengue, 

at 969m, is the highest point in the Reserve (Thomas and Thomas, 1992). 

The parent rock of the area is made up of micaschists, superior and inferior gneiss and 

undifferentiated gneiss. There are essentially two types of soil, ferric soils and 

hydromorphic soils. The ferric soils are somewhat yellow in colour and are derived 

from the metamorphic rocks characteristic of the coastal plain. The hydromorphic 

soils develop in a distinct layer near to the soil surface in area of swamp and seasonal 

inundation. 

3.3.1.4 Vegetation 

The vegetation of the Campo region is defined by Letouzey (1985) as Atlantic Biafran 

forest rich in Caesalpiniaceae 2 . This vegetation type is widespread within 100km of 

the coast, from Nigeria to Equatorial Guinea. Campo is rich in Caesalpiniaceae; there 

are large numbers of individuals, high dominance and high levels of diversity of this 

family. Although Letouzey (1985) described a single vegetation type covering the 

whole range, there is considerable variation in dominant species and species 

composition between localities, there are several sub-types in the area. These are 

characterised by indicator species such as Sacoglottis gabonensis (Humeriaceae) and 

Calpocalyx heitzii (Mimosaceae) (Sunderland et al., 1997). 

175

The vegetation of the sample sites and the surrounding area is classified as Atlantic 

Evergreen Biafran Forest with Caesalpiniaceae and Calpocalyx heitzii (Letouzey, 

1985). This forest type covers a large area of the south of Cameroon and the northern 

regions of Equatorial Guinea. Unlike many areas within Cameroon, this forest 

formation is somewhat unique in its homogeneity and is representative of a large 

forested area from southern Cameroon to northern Equatorial Guinea. 

Along with numerous members of the Caesalpiniaceae, the characteristic species of 

this forest formation are Anthonotha macrophylla (Papillionaceae), Coula edulis 

(Olacaceae), Glossocalyx brevipes (Monimiaceae) Lophira alata (Ochnaceae) and 

Scyphocephalium mannii (Myristicaeae) Other indicator species are Hoplestigma 

klaineanum (Hoplestigmataceae), the abundant understorey tree Meiocarpidium 

lepidotum (Annonaceae) and Podococcus barteri, a slender understorey palm that 

forms dense, often monospecific, stands. 

3.3.1.5 Forest exploitation 

This forest type is highly prized for its timber resources and as such, is heavily 

logged, even within the boundaries of the Reserve. Hence, a substantial proportion of 

the study area supports an intricate mosaic of secondary regrowth vegetation. Nontimber 

forest products are also exploited, particularly for primary health care on which 

the majority of the local population relies, in the absence of formal medical services. 

Rattan is not harvested at levels beyond immediate household requirements. 

3.3.2 Mokoko River Forest Reserve 

3.3.2.1. Introduction 

The Mokoko River Forest Reserve (4°21’-4°28’N; 8°59’-9°07’E) covers an area of 

9,100ha and is situated to the north-west of the Bambuko Forest Reserve. The reserve 

is bordered to the south-east by the Onge River, to the north-east by the Meme River 

estuary, to the north by the Mokoko River and to the west by the River Boaba. The 

Mokoko FR was created in 1952 as a production forest and this legal status has 

remained unchanged by the successive Forestry Laws of 1983 and 1994. Currently, 

2 I have maintained the continental nomenclature of the Leguminosae, where the three sub-families are 

accorded family status, throughout this Chapter. 

176

the reserve has no management plan (Thomas, 1994; ERM, 1998; Sunderland and 

Tchouto, 1999). 


Annual precipitation on the Western flank of Mount Cameroon varies between 

3,000mm and 4,000mm and declines further east of Mokoko in a rain shadow caused 

by the Mount Cameroon massif. There is a single rainy season between March and 

October. December, January and February are all relatively dry months, often with no 

rain falling at all. The annual mean temperature is 27°C. 


The Mokoko FR is situated at an altitude between 120 to 360m, on the north west 

inferior slope of Mount Cameroon. The majority of the reserve is seated on ancient 

volcanic rocks, basalts and trachites. Nearer the sea, there are small areas of more 

recent alluvial deposits. The soils that predominate within the reserve have been 

formed by an association between ancient and recent volcanic material with a small 

area of non-volcanic sedimentary sandstone-derived soil nearer to the Boa Plain 

(ERM, 1998). 


The majority of the lowland forests in the hinterland of Mount Cameroon have been 

converted to industrial plantations, and the forests of the Mokoko area currently 

constitute the only pristine and most extensive forest formation in the region. As such, 

they are extremely important, both in terms of biodiversity value (Cable and Cheek, 

1998) and for indigenous use and management (Sharpe, 1998; Sunderland and 


The vegetation of the Mokoko Reserve was originally described by Letouzey (1985) 

as being of the “Atlantic Biafran evergreen forest with numerous Caesalpiniaceae” 

formation. Whilst this general description certainly encompasses the general 

physiognomy of the area, Letouzey’s classification, at 1:500,000 scale with limited 

ground-truthing, does not allow for more subtle variations in vegetation that are 

characteristic of the forests around Mount Cameroon. As such, subsequent field-work 

177

and observations have built upon Letouzey’s original classification (Gartlan, 1989; 

Thomas, 1994). 

Based on preliminary Mount Cameroon Project inventory results, Thomas (1994) 

suggests that the vegetation of Mokoko is comprised of two forest types that intergrade 

completely. To the south, where the rainfall is higher is what Letouzey (1985) 

described as Atlantic Biafran Forest with Caesalpiniaceae with Oubangia alata 

(Scytopetalaceae) and other coastal indicators such as Protomegabaria stapfiana 

(Euphorbiaceae), Dichostemma glaucescens (Euphorbiaceae), Octoknema affinis 

(Octoknemataceae), Tapura africana (Dichapetalaceae) and many members of the 

Olacaceae. 

Moving progressively northwards, the coastal forest grades into Atlantic Biafran 

forest with Caesalpiniaceae well represented by gregarious genera such as Didelotia, 

Hymenostegia afzelii, Microberlinia bisulcata, Monopetalanthus, Plagiosiphon and 

Tetraberlinia bifoliolata. The Myristicaceae are also an important component of this 

forest, represented in particular by Coelocaryon preussii, Scyphocephalium mannii 

and Staudtia stipitata. The narrow endemic Medusandra richardsiana 

(Medusandraceae) along with Oubangia alata (Scytopetalaceae) dominate the 

understorey along with Garcinia mannii (Guttiferae) and Lasianthera africana 

(Icacinaceae), the latter of which are both important NTFP resources. The genera Cola 

(Sterculiaceae) and Diospyros (Ebenaceae) are also important components of the 

lowland forests of northern Mokoko. Cable and Cheek (1998) suggest that the forest 

types found in the northern Mokoko area exhibits clear affinities with the southern 

Korup area and, to a lesser extent, to the forests immediately south of the Sanaga. 

The vegetation was also described by Gartlan (1989) as having certain semideciduous 

elements at its western edge. These latter elements originate from the rain 

shadow present to the north of Mount Cameroon which has species communities 

present more often associated to those of the drier, eastern parts of Cameroon such as 

Triplochiton (Sterculiaceae) Ceiba (Bombacaceae) and many other representatives of 

the Meliaceae and Sterculiaceae. The presence of these species more commonly 

178

associated with drier forest, was confirmed by an inventory undertaken by the Mount 

Cameroon Project (Thomas, 1994). 

Another interesting component to the flora of the Mokoko area is the presence of the 

sub-Sahelian palm, Borassus aethiopum, which is at the southern-most edge of its 

range. These Borassus-dominated savannahs occur on deep ash soils, and are a highly 

unusual formation in what is, essentially, a forest zone (Sunderland and Tchouto, 

1999). 


The exploitation levels of timber and non-timber forest products in Mokoko is high 

(Sunderland and Tchouto, 1999). There is a thriving Nigerian-led cross-border trade 

in a number of high-value forest products such as chewstick (Garcinia mannii), 

“vegetable” (Lasisanthera africana and Gnetum spp.) and the cattle-stick (Massularia 

acuminata). Rattan is also widely exploited and exported. 

3.3.3 Takamanda Forest Reserve 


The Takamanda Forest Reserve (05º59’-06º21’N: 09º11-09º30’E) covers an area of 

67,600 ha and is situated at the most northern point of Cameroon’s South-West 

Province, north of the extensive Cross River valley. Created by decree in 1934, the 

reserve stretches along the eastern border of Nigeria and this border forms the north 

and north-west boundaries of the reserve itself (Gartlan, 1989). 


The Takamanda area has two distinct seasons. Most rainfall occurs from April through 

to November, with a peak in July and August and a second peak in September. 

Although accurate figures are not available, the total annual rainfall is probably 

similar to that of the Nigerian side of the border in the Okwangwo region (up to 

4500mm) (Groves and Maisels, 1999). From November through to April, the climate 

is mainly dry with some months (January to March) often having no rain at all. The 

mean annual temperature is around 27º C. 

179


The majority of the lowland forest area within the southern and central part of the 

Reserve lies between 100-400m. The terrain is rolling in the lowland areas but rises 

sharply to 1500m altitude in the north of Reserve, where slopes are extremely steep. 

Small hills, up to 725m, lie to the north of the Obonyi villages along the border with 

Nigeria. The hills separating the villages of Kekpani and Basho are similar in height, 

rising to about 600-700m. The soils of the reserve are mainly ferrite, derived from 

Pre-cambrian crystalline rock, although large areas of alluvial soil are found towards 

the southern end of the reserve. 


The majority of the southern and central parts of the Takamanda Forest Reserve are 

covered by lowland (100-700m) Guineo-Congolian forest. The dense sub-montane 

(“highland”) humid forest (700-2,000m) is predominantly found mostly in the 

northern part of the Reserve, although some isolated patches occur further south on 

long ridge tops and isolated hills reaching an altitude of up to 1,500m. Areas further 

east and outside of the Reserve, including the northern part of the Mone Forest 

Reserve, including Mount Oko, which rises to over 1500m, are also of this latter 

vegetation type. 

There are also some isolated patches of wooded savannah and gallery forests, along 

with relics of herbaceous savannah, in the extreme north of the Reserve. Whilst these 

vegetation types are an important component of the highland areas towards Akwaya, 

they are but a small component of the Takamanda Forest Reserve itself. This 

savannah/ woodland complex extends eastwards and northwards along the Nigerian 

border, and in the direction of the Bamenda highlands. Finally, there are three 

enclaved villages within the Reserve, surrounded by extensive areas of secondary 

vegetation and farm bush. Patches of this vegetation type are found near all the 

villages of the area. 

The lowland forest of the Takamanda Reserve forms part of a large contiguous forest 

block that covers large areas of the Cross River Basin north of Mamfe (Letouzey, 

1985). The reserve is unique in that it represents a sharp gradation from lowland forest 

to sub-montane (highland) forest with the associated variations within. Floristically, 

180

the lowland forest is characterised by species commonly associated with the Atlantic 

coastal forest yet the relative paucity of representatives of the family Caesalpiniaceae, 

normally abundant in the closed-canopy forests of Central Africa, make this forest 

formation distinct. Whilst relatively contiguous, there is some variation within this 

forest formation, most notably in seasonally inundated areas and on hill-tops and 

ridges. 

A common component of these forests is the family Irvingiaceae, canopy-emergent 

trees represented by eight species (Harris, 1996). This family includes the 

economically important sweet and bitter bush mango (Irvingia gabonensis and I. 

wombolu respectively) as well as large numbers of Klainedoxa gabonensis and 

Desbordesia glaucescens. Other common canopy trees include: Maranthes 

gabunensis (Chrysobalanaceae), Terminalia ivorensis, T. superba (Combretaceae), 

Andira inermis (Papillionaceae), Poga oleosa (Anisophylleaceae), Coula edulis 

(Olacaceae), Pterocarpus soyauxii (Papillionaceae) and Scyphocephalium mannii 

(Myristicaceae). The mid-canopy is dominated by Tapura africana (Dichapelaceae), 

Treculia obovoidea (Moraceae), Diogoa zenkeri (Olacaceae) and Rauvolfia 

macrophylla (Apocynaceae) along with numerous additional members of the 

Euphorbiaceae and Olacaceae. In seasonally inundated areas and along river-banks, 

Protomegabaria stapfiana (Euphorbiaceae) is abundant, along with large populations 

of Pandanus (Pandanaceae) and members of the utilitarian palm genus, Raphia. Along 

the ridges and lower altitude watershed areas, the coastal affinities become more 

obvious. These long ridges are often characterised by stands of even-aged Lophira 

alata (Ochnaceae) along with numerous Canarium schweinfurthii, and Santiria 

trimera (Burseraceae) as well as by some representatives of the Caesalpiniaceae, 

including Berlinia bracteosa, Afzelia bipindensis, Microberlinia bisulcata, 

Erythrophloem ivorense and many others. The narrow endemic, Napoleana egertonii 

(Lecythidaceae) is a common feature of these higher altitude forest formations, as are 

a number of species Garcinia (Guttiferae). 

Common herbs in the closed-canopy lowland forest include stands of Impatiens 

(Balsaminaceae) along the smaller rocky streams in particular, often accompanied by 

semi-epiphytic Begoniaceace, and many members of the Araceae, Commelinaceae, 

181

Marantaceae and Cyperaceae. Cyanastrum cordifolium (Cyanastraceae) another 

common feature of Atlantic-type forest is also common encountered on the forest 

floor. The highly seasonal nature of the area also means that variations in water levels 

in many rivers and streams expose extensive rocky areas. A common and persistent 

coloniser of these oft-flooded rocks is Biophytum petersianum (Oxalidaceae) a small 

rhizomatous herb widespread in the Korup forest but not commonly encountered 

elsewhere. 

As the altitude increases, especially where the dramatic relief and rocky outcrops 

preclude the ability of forest to fully develop and persist, many open areas are 

commonly encountered in this forest type, most notably on hill sides and ridge tops. 

Whilst not strictly “highland”, at altitudes often less than 700m, these vegetation 

formations, often characterised by dense, monospecific stands of Acanthaceae and 

Costaceae as well as members of the Balsaminaceae also being common, are more 

commonly associated with higher altitude vegetation. 


There is little or no exploitation for timber in Takamanda. The harvest and sale of 

some high-value non-timber forest products comprises a large proportion of the 

household income (Groves and Maisels, 1999; Ifeka, pers. comm.). In particular, bush 

mango (Irvingia gabonensis and I. wombolu) is of significant importance and is 

widely traded as is njansang (Ricinodendron heudelottii) and njabe oil (Baillonella 

toxisperma). Rattan is not harvested at levels beyond immediate household 

requirements. 

3.4 METHODS 

3.4.1 Sampling 

A series of 1ha plots were established in each of the survey sites. The number of plots 

that were established at each site was dictated by both biological and logistical 

factors. A stratified random sampling design was followed to ensure that the plots 

were representative of the forest type within each reserve area and provide a 

reasonably homogenous sample within that variation. In total nine, 1ha plots were 

established: two in Takamanda, three in Campo and four in Mokoko. Although the 

182

sampling intensities were much lower than for standard timber inventories, 0.003%, 

0.001% and 0.045% respectively, and it is probable that some minor forest formations 

were not sampled, a random stratified approach to site selection, ensured that the main 

forest types were represented. Increased resources would facilitate the greater 

sampling intensities for full resource inventories; increased sampling which is needed 

for the development of detailed management plans (Stockdale and Wright, 1994). 

In addition to the quantitative assessment for each survey site, intensive floristic 

inventories of the rattan resource were also undertaken in each area (see Sunderland et 

al., 1997). 

3.4.2 Plot establishment 

Within the 1ha plot configuration, the plot is divided into 25 quadrats, each 20 x 20 

meters in size 3 . To eliminate errors, surveying to establish the quadrat corners 

proceeds from the centre of the plot outwards. A row of quadrats is built north to 

south along the centre line from its midpoint; new quadrats are then added westward 

until rows two and three are completed. The next step is to finish the centre row 

before moving to the eastern portion of the plot and establishing the remaining 

quadrats. 

The quadrats are permanently marked at each of their corners with aluminium or 

plastic stakes and tagged with a letter and number to differentiate their location within 

the plot. Marking starts at the baseline, which is the north/south border on the east 

side of the plot. All stakes protrude well above the forest floor to increase visibility. 

Red stakes are used for the plot corners, and white for marking the quadrats. 

The survey area is surveyed in a horizontal plane using a theodolite. Corrections for 

slope are made, ensuring that each of the quadrats contains 400 square meters 

regardless of topography 4 . Before enumerating the plot, string is tied along the borders 

of the quadrats. This helps in locating plants to be enumerated within each quadrat. 

3 

It is generally accepted that 20 meters is the longest distance that can be accurately surveyed in a 

dense forest (Dallmeier, 1992). 

4 

Correcting for the influence of slope is crucial is establishing a horizontal plane of 1 ha in size. The 

equation: Slope Correction = 1/cos arctan (% slope / 100) is used for slope correction but most hand- 

183

Figure 82. (i) One-hectare plot divided into 25 quadrats. 

(ii) One 20 x 20 m quadrat divided into 16 quadrats. 

3.4.3 Mapping 

It was intended that the rattan individuals on all nine plots would be mapped, 

following the methodology outlined in Dallmeier (1992). However, after the mapping 

of the first plot in the Campo FR (see Appendix 3) it became clear that both time, 

resource and logistical constraints would not enable this activity to take place on the 

remaining plots. 

3.4.4 Enumeration 

To provide a quantitative vegetation description for each site, all of the trees on each 

plot >10 cm dbh were measured and identified. Voucher specimens were collected for 

tree species that could not be identified with confidence in the field 5 . For the rattan 

species, each individual was identified and tagged with numbered aluminium labels 

attached to the rachis, or stem, with wire. Each cluster (ramet) was numbered as a 

single individual with each stem (genet) within the cluster measured, numbered and 

tagged separately. For both ramets and genets, the species and size (height class of 

held clinometers have tables attached to them to allow the calculation of slope correction easily in the 

field. 

5 These voucher specimens are held in the Limbe Botanic Garden Herbarium (SCA). 

184

seedling 6 or length of stem) were recorded. For mature individuals, both the total stem 

length and harvestable stem length were measured and recorded. 

3.5 PRESENTATION OF RESULTS 

Based on the enumeration of the vegetation of each site, the relative density and 

relative dominance were calculated by family and species. These are presented by 

family below: 

3.5.1 Cumulative summary of vegetation plot results by site 

Table 2. Presentation of vegetation summary by plot for all trees > 10cm dbh 

Mokoko Campo Takamanda 

1 2 3 4 

Plot No. 

5 6 7 8 9 

No. of stems 462 525 418 453 397 402 394 525 526 

No. of species 88 90 92 81 74 81 74 75 80 

No. of genera 72 75 78 70 63 67 66 53 65 

No. of families 30 32 33 31 28 25 25 27 26 

Basal area (m²) 30.3 30 30.8 31 30.8 30 35 33.2 34 

Figure 83. Cumulative size-class distribution for sample area compared with the Pantropical 

mean (from Philip, 1994). 

Mean no. trees / ha 

350 

300 

250 

200 

150 

100 

50 

0 

Campo Mokoko Takamanda Pantropical mean 

10-19 20-29 30-39 40-49 50-59 60-69 70-79 80-89 90-99 > 100 

Size class 

6 In the seedling stage, size was measured as total height within a given height class, regardless of the 

number of leaves as used by Bøgh (1996). With Laccosperma secundiflorum, in particular, the 

variation in the number of leaves is not determined by height (or age) and the number of leaves of a 3m 

high seedling varies from between 3 and 7. 

185

3.5.2 Summary of rattan stocking and abundance by site 

Table 3. Rattan abundance and stocking for Campo 

Laccosperma Laccosperma Eremospatha Oncocalamus 

opacum robustum hookeri mannii 

Mean # of 

individuals/ha -1 

47 87 3 5 

Mean # of clumps / ha -1 - 4 1 - 

Mean # of mature stems / 

cluster 

- 14 1 - 


ha -1 

- 56 1 - 

Mean total mature stem 

length (m/ha -1) 

- 1,181 20 - 

Mean stem length / 

individual (m/ha -1) 

- 21 20 - 

Mean total harvestable 

cane length (m/ha -1) 

- 305 0 - 

% total stem length 

harvestable 

- 26% 0% - 

Table 4. Rattan abundance and stocking for Mokoko 


individuals/ha - 1 


clumps / ha - 1 

Mean # of mature 

stems / cluster 

Mean # of mature 

stems / ha – 1 

Mean total 

mature stem 

length (m/ha –1) 

Mean stem 

length / 

individual (m/ha 

–1) 

Mean total 

harvestable cane 

length (m/ha –1) 

% total stem 

length 

harvestable 

Laccosperma Laccosperma Eremospatha Eremospatha Eremospatha 

opacum secundiflorum hookeri wendlandiana macrocarpa 

92 42 4 2 4 

- 3 - - 1 

- 11 - - 13 

- 33 - - 13 

- 316 - - 169 

- 95 - - 13 

- 80 - - 63 

- 25% - - 37% 

186

Table 5. Rattan abundance and stocking for Takamanda 

Laccosperma Laccosperma Eremospatha Eremospatha 

opacum secundiflorum tessmanniana macrocarpa 


individuals/ha -1 

47 330 3 134 

Mean # of clumps / ha -1 - 4 3 3 


cluster 

- 15 9 9 


ha -1 

- 59 9 26 

Mean total mature stem 

length (m/ha -1) 

- 1,011 184 435 

Mean stem length / 

individual (m/ha -1) 

- 17 20.4 20.4 

Mean total harvestable 

cane length (m/ha -1) 

- 256 0 184 

% total stem length 

harvestable 

- 25% 0% 42% 

3.6 DISCUSSION 

3.6.1 Floristic diversity and similarity 

Species diversity for each site was assessed using the Shannon-Weaver index 7 . Based 

on the distribution of individuals among the different species, Mokoko is the most 

diverse site (H = 2.81) followed by the Takamanda FR (H = 2.66) and the Campo FR 

(H = 2.58). This study correlates closely with previous estimates of species diversity 

for Campo (Sunderland et al., 1997) and Mokoko (Thomas, 1994). The figures for 

Takamanda represent the results of the first vegetation study in that area. In many 

respects, the estimates of species diversity by this study follow the general perception 

that forests with greater rainfall and more heterogeneous forest, are known to harbour 

a greater number, and range, of species (Maley, 1996; Sosef, 1996). Hence it is 

unsurprising that Campo, with less annual rainfall (and two distinct dry seasons) is 

characterised by a relatively homogenous forest formation and possesses the least 

diversity of species of the three sites. 

In terms of similarity (i.e. shared taxa), the application of both Sorenson’s coefficient 

and Jaccard’s coefficient suggests that closer floristic affinities are shared between 

Takamanda and Campo, with Mokoko possessing a greater number of species unique 

to that site. The is undoubtedly due to the fact that the lowland forests of the Mount 

187

Cameroon hinterland are well-known for their high levels of endemism and speciation 

(Cheek et al., 1992; Thomas, 1994; Cable and Cheek, 1998) and these forests are 

postulated to represent a Pleistocene refuge of some significance (Brenan, 1978; 

Hamilton, 1981; Pannel and White, 1988; White, 1993; Maley, 1996; Robbrecht, 

1996; Sosef, 1996; Morat and Lowry, 1997). 

Table 6. Floristic similarity between the sample sites (all species) 

Sorenson’s coefficient 8 Jaccard’s coefficient 9 


Mokoko - 0.609 0.560 

Campo 0.757 - 0.818 

Takamanda 0.718 0.900 - 

3.6.2 Rattan diversity and similarity 

It is speculated that, to a certain degree, rattan diversity is positively correlated to 

overall species diversity (Dransfield, 1979; Dransfield, 1992b). The results of this 

study provide evidence to support this hypothesis. The Mokoko FR shows the greatest 

diversity of rattan species (Shannon-Weaver index H = 1.09) followed by the 

Takamanda FR (H = 1.01) and the Campo FR (H = 0.31). From these results, it is 

clear that there is a strong positive correlation between overall floristic diversity and 

rattan diversity (Pearsons correlation analysis r = 0.849, P < 0.01). In this respect, and 

in the African context, preliminary study of the rattan diversity of an area may indeed 

prove to be a reliable indicator of overall species diversity. 

In terms of floristic similarity, however, the affinities between the rattan flora are 

closer between Mokoko and Takamanda, and Mokoko and Campo, with very few 

species shared between Takamanda and Campo. This is in contrast to the similarities 

in the wider vegetation presented above. Although it is not clear why this might be the 

case, the evolutionary history of the palm flora in Africa (as discussed in Chapter 1) 

may account for these affinities. However, unlike the vegetation as a whole, the 

7 Shannon-Weaver Diversity Index: H = -xpi 1n pi where p is the proportion of a particular species in 

a sample which is multiplied by the natural logarithm of itself. H is derived by summing the product for 

all the species in a sample (Hayak and Buzas, 1997). 

8 Sorenson’s coefficient: CN = 2jN / (aN+bN), where aN is the number of individuals in site A, bN is the 

number of individuals in site B and jN is the sum of the lower of the two abundances of species which 

occur at the two sites (Hayak and Buzas, 1997). 

9 Jaccard’s coefficient: Cj = j / (a=b-j), where j is the number of species common to both sites, a is the 

number of species in site A, and b is the number of species in site B (Fowler et al., 1998). 

188

sampling of the rattan population, by either quantitative or qualitative methods, may 

not have been sufficiently intensive for any firm conclusions to be made in this 

respect. 

Table 7. Floristic similarity between the sample sites (rattan species) 

Sorenson’s coefficient Jaccard’s coefficient 


Mokoko - 0.429 0.450 

Campo 0.600 - 0.143 

Takamanda 0.598 0.250 - 

Table 8. Comparison of rattan stocking by site 

Species 

Campo Mokoko Takamanda 

Mean no. Mean stem Mean no. Mean stem Mean no. Mean stem 

individuals / (harvestabl individuals / (harvestable) individuals / (harvestable) 

ha –1 e) m / ha –1 ha –1 m / ha –1 ha –1 m / ha -1 

Laccosperma 

opacum 

47 - 92 - 47 - 

L. robustum 87 1,181 (305) - - - - 

L. secundiflorum - - 42 316 (80) 330 1,011 (256) 

Eremospatha 

macrocarpa 

- - 4 169 (63) 134 435 (184) 

E. hookeri 3 - 4 - - - 

E. tessmanniana - - - 3 - 

E. wendlandiana - - 2 - - - 

Oncocalamus 

mannii 

5 - - - - - 

Totals = 142 1,181 (305) 144 485 (143) 514 1,446 (440) 

Results of 

No. of 

Sum of square df Mean Square F value Pr (F) 

Analysis of individuals Site 657.00 2 328.5 6.750 0.266 * 

Variance 

Residuals 50.00 5 12.89 

(ANOVA) Total stem 

Sum of square df Mean Square F value Pr (F) 

length Site 36486.864 2 18243.432 7.764 0.246 * 

Residuals 2349.886 5 327.89 

Notes: df = degrees of freedom; F value = value of the test; Pr (F) = probability of error; * = probability 

of error at P

(Sunderland, 1998; Sunderland, 1999a; Sunderland, 1999b), and the evidence 

provided above seems to support this hypothesis. This is in direct contrast to the 

situation in Malaysia where Nur Supardi et al. (1996; 1999) found that logged forest 

had a negative impact on the rattan population. The reasons for this discrepancy are 

not related to the dynamics of the rattan population itself, but are undoubtedly due to 

the nature of the logging process. Timber exploitation in SE Asia is characterised by 

the removal of large numbers of individual stems/ha as the commercial timber species, 

predominantly members of the Dipterocarpaceae, occur in high concentrations (Hooi, 

1987; Mok, 1992; Brown and Whitmore, 1992; Pinard et al., 2000). The resulting 

secondary regrowth is not suited to rattan regeneration and it is some while before 

rattans can establish themselves in the secondary forest (Nur Supardi, 1999). 

However, the greater spatial distribution of timber species in the forests of West and 

Central Africa results in a much-reduced level of exploitation (2-3trees / ha) (Jonkers 

and Leersum, 2000). This selective felling, when undertaken responsibly and with 

minimum extraction damage, creates a mosaic of semi-natural gaps within the forest 

that are rapidly colonised by light demanding rattan species (Sunderland, 1999a; 

Sunderland 1999b). 

As discussed above, unlike the other two sites, which are in unlogged forest, the plots 

in the Campo FR are sited in forest that has been selectively logged 20-25 years ago. 

Despite a relatively low number of individuals, the proportion of mature rattan stems 

is higher in this sample area and hence the cumulative (and harvestable) stem length is 

correspondingly greater. Hence, the selective logging activities in Campo seem to 

have had a positive impact on the rattan population. This confirms the observations 

that certain species of rattan in Africa respond positively to moderate reductions in 

canopy cover and subsequent increases in light penetration (Sunderland, 1999a; 

1999b). 

Despite the presence of a robust and diverse rattan population, the relative paucity of 

harvestable cane from Mokoko is undoubtedly a result of the high level of cane 

exploitation from the area. The harvesting cycle is considerably short and the levels of 

exploitation are currently beyond the capacity of the rattan to regenerate sufficiently; 

hence, this exploitation is currently unsustainable. This Nigerian-led exploitation is 

190

concentrated, not only on rattan, but many other NTFPs as well (Sunderland and 


3.6.3 Total stem length vs harvestable cane length 

The tables above show that the percentage of harvestable cane per species is relatively 

constant. In this regard, for all species there is a positive correlation between total 

stem length and the length of cane harvested (Pearson correlation analysis r = 0.848, P 

< 0.01). In addition, the findings of this study clearly show that the mature stems need 

to reach a minimum length of 20m before they provide adequate quantities of 

harvested cane. 

Figure 84. Correlation between total stem length and harvestable length 

30 

Ha 

rve 

sta 

ble 20 

len 

gth 

(m 

) 

10 

0 

0 10 20 30 40 50 60 

Total stem length (m) 

3.6.4 Inventory techniques and sampling intensity (Species capture) 

191

As mentioned in the discussion of methods, this survey took place in conjunction with 

floristic inventories of the three sites. Whilst somewhat subjective, these latter studies 

prove that many more rattan species occur at each site than were included in the 

enumeration process, probably due to the site specific requirements of the different 

species (Wong, 1997; Sunderland, 1999a; 1999b). This lack of representation in the 

sampling is undoubtedly a shortfall in both the methodology and the sampling 

intensity. To enable a more representative sampling of the rattan flora, it might be 

more appropriate, and cost effective, to establish a wider network of smaller plots 

(0.025-0.05 ha), as recommended by Stockdale and Wright (1994). This would ensure 

that important forest formations would be better represented, particularly for 

heterogeneous forest, and hence a greater proportion of the known species would be 

included in the inventory process. 

Table 9. Sampling and representation 

No. of spp. recorded 

in plots 

192 

No. of spp. known 

from botanical 

collections 

% representation 

Campo 4 9 45 

Mokoko 5 11 45 

Takamanda 4 10 40 

When compared to studies that have been undertaken in SE Asia, it is interesting to 

note that both the diversity and abundance of rattans in Africa is relatively low (see 

Table 10). In terms of diversity, the relative paucity of species diversity in the Palmae 

as a whole surely accounts for this discrepancy (see Chapter 1). With smaller 

populations and fewer species, it is then unsurprising that the stocking and abundance 

of rattans in Africa is less than that for comparative sites in SE Asia.

Table 10. A comparison of rattan diversity, abundance and stocking between sites 

Sample Sample 

size 

Khao Chong 

NP, Thailand 

Temburong, 

Brunei 

Darussalam 

Kerinci- 

Sablat NP 

Dugoma Bone 

NP 

(Indonesia) 

Kalimantan, 

Indonesia 

Pasoh FR, 

Malaysia 

Western 

Province, 

Ghana 

Cross River 

NP, Nigeria 

Southern 

Cameroon 

Campo, 

Cameroon 

Mokoko, 

Cameroon 

Takamanda, 

Cameroon 

3.7 CONCLUSION 

Total no. 

of 

species 

Mean no. of 

individuals / 

ha –1 

193 

Mean 

cumulative 

stem length 

/ ha -1 

Harvestable 

stem length 

/ ha -1 (m) 

Reference 

1 ha 11 1,319 2,341 2,066 Bøgh, 1996 

1.5 ha 21 228 ? ? Stockdale & 

Wright, 

2 ha 1 

1 

38 

65 

? 

? 

2,666 

1,910 

1994 

Siebert, 

1993 

1 ha 1 754 ? ? Peters, 1996 

1.92 ha 20 1750 ? ? Nur Supardi, 

1999 

814 ha 5 42.5 ? ? Wong, 1997 

236 ha 2? 350 ? ? CRSFP, 

1994 

32 ha ? 90 ? ? van Dijk, 

1995 

3 ha 4 142 1,181 305 This study 

4 ha 5 144 485 143 This study 

2 ha 4 514 1,446 440 This study 

This study suggests that there is a strong positive correlation between overall species 

diversity and rattan diversity and the richness of the rattan flora may reflect that of the 

wider vegetation of a sample area. As such, rapid assessments of key taxa, such as 

rattan palms, might provide a cost-effective preliminary means of contributing ti the 

assessment of overall forest diversity. 

There does not seem to be a particularly strong influence of edaphic factors on rattan 

diversity and abundance as is reported in SE Asia (Nur Supardi, 1999). Despite the 

variation in soil type between the three sites, there is no significant variation in rattan 

diversity based on this, although this quite probably accounts for the variation in 

species composition between the three sites. Hence, the free-draining basaltic soils of

Mokoko, for example, are sufficiently distinct from the other sites that such the rattan 

flora there has less affinity with that of Campo in particular. This study shows, 

however, that the amount and uniformity of annual precipitation influences overall 

species diversity. The lower rainfall site with two strongly defined dry seasons 

(Campo) is somewhat depauperate in overall species diversity and, to a certain degree, 

the rattan flora. 

The anecdotal observations that suggest that rattans respond very positive to logging 

activities seem to be supported by this study (Defo, 1999; Sunderland, 1999a; 1999b). 

However, it should be noted that the gap partitioning is influential purely in terms of 

abundance and not diversity and the number of species in logged forest does not 

increase. However, the increased light penetration allows those seedlings in the 

seedling bank (that would normally be prone to high levels of mortality) to develop 

beyond the establishment phase into mature individuals. This obviously has 

implications for potential harvest yields. 

This study also confirms that there is a strong positive correlation between total stem 

length and the harvestable stem length. For all commercially valuable species, the 

minimum length of mature stem that is potentially harvestable, are those of >20m in 

total length. 

Whilst the permanent sample plots established as part of this study, through continued 

monitoring, will provide useful information on recruitment, growth and mortality of 

the rattan species contained within, they do not provide an adequate sampling of either 

of the three sites. In this respect, the 1ha plot methodology is not suitable for 

management inventories and it is suggested that a different plot methodology (smaller 

plots at a greater sampling intensity) is implemented for full resource surveys. 

Finally, it should be noted that in comparison to studies in SE Asia, it is clear that in 

terms of both diversity and abundance, the rattan resource in Africa is somewhat 

depauperate. This is undoubtedly due to the paucity of the palm flora, as a whole, in 

Africa. 

194


CHAPTER FOUR 

RATTAN / FAUNAL RELATIONSHIPS 

Herbivory, predation and seed dispersal by animals are widely recognised processes in 

tropical forests (Whitmore, 1990; Happold, 1996). However, the most basic 

information on the natural history of most floral / faunal relationships remained, until 

recently somewhat sketchy and incomplete. In the lowland tropical forests of Africa, a 

number of these relationships have been studied yet these are often rooted in the 

relationship between a single organism and its wider environment, rather than speciesspecific 

interactions. Notable exceptions to this include the study of the relationships 

between lowland gorillas (Gorilla gorilla gorilla) and Cola lizae (Tutin, et al., 1991); 

Barteria fistulosa and Pachysima aethiops ants (Janzen, 1972) and the reliance on 

elephants (Loxodonta africana) to regenerate and disperse seeds of Balanites 

wilsoniana (Chapman et al., 1992) and Strychnos aculeata (Martin, 1990) 1 . 

With regard to rattans, considerable anecdotal information from research in Asia 

suggests that rattans interact widely with their environment. For example, the stems of 

many species of rattan are widely predated by forest hogs and elephants for their 

growing points and the fruits are consumed and dispersed by a range of agents, most 

notably hornbills (Dransfield, 1992b). However, despite the scale of research 

undertaken in Asia, there is surprisingly little information about the specific nature 

and relative importance of such relationships. Accordingly, until recently, very little 

was known about the relationship of the African rattans and their wider environment, 

and even less was known about the importance of species-specific relationships, if 

indeed such relationships occur. Although much of the information contained in this 

chapter is predominantly based on personal observations and information provided by 

local informants, a review of the available literature has supported some of these 

observations. 

1 Although in a recent review of the role of elephants in seed dispersal, Hawthorn and Parren (2000), 

suggested that both species continued to regenerate in forests where elephants are absent and have been 

for some time. 

195

4.2 HERBIVORY 

Being covered in spines is no defence from herbivory from the most wily or persistent 

of animals and there are strong indications that herbivory of rattans in Africa is 

significant, particularly from endemic megafauna. Destructive damage and feeding on 

some rattan species by primates and other organisms is now known to be relatively 

common. 

4.2.1 Stem apex or “palm heart” 

As discussed in Chapters 1, 6 and Appendix 1, the soft, developing stem apex, or the 

“palm heart” is a morphological feature shared by all species of rattan. The size of the 

palm heart itself is dependent on the diameter of the cane and hence varies from 

species to species. The palm heart is widely consumed not only by humans, but also 

by a range of faunal predators (White and Abernethy, 1997; Williamson et al., 1990; 

Moutsambouté et al., 1994; Idani et al., 1994; Yumoto et al., 1994; Tutin et al., 1994; 

author pers. obs.). Due to the rather robust nature of most rattan species, dragging 

down the stem apex, often anchored high up in the forest canopy, and then removing 

the strong leaf sheath to access the developing stem, entails significant strength and 

persistence on the part of the predator. Because of this, it is generally only large 

mammals that are able to feast on this part of the rattan palm. In particular, the stem 

apices of a number of species of Laccosperma are consumed by elephants (Loxodonta 

africana) and larger primates, notably the chimpanzee (Pan troglodytes), the bonobo 

(Pan paniscus), and the western lowland gorilla (Gorilla gorilla gorilla 2 ). Elephants 

consume the palm heart by physically dragging down the individual stems from the 

base until the apex itself is within reach. The apex is then bent to break the enclosing 

sheath and the exposed soft inner pith is consumed. 

Primates, on the other hand, usually access the stem apex by climbing adjacent trees 

and then reaching out to break the sheath at the apex and remove the inner pith. 

However, it is also recorded that gorillas drag down the stems of some of the more 

slender species of rattan such as Eremospatha haullevilleana and E. cabrae to access 

2 Although the Cross River gorillas are now known to represent a distinct sub-species within the 

western lowland gorilla complex (Sarmiento and Oates, in press), for the purposes of this chapter, all of 

the lowland gorillas are referred to as a single taxonomic group. 

196

the palm hearts from ground-level (Yumoto et al., 1994; Tutin et al., 1994). In the 

case of Eremospatha, the lack of spines on the sheath makes it possible for primates to 

directly handle the individual stems without incurring the irritation of numerous 

spines penetrating the skin. In this respect, there is a well-known preference amongst 

primates for the consumption of palm hearts of species within this genus, rather than 

for species with spiny sheaths, such as Laccosperma, Calamus and Oncocalamus 

(Williamson et al., 1990; Moutsambouté et al., 1994; Idani et al., 1994; Yumoto et al., 

1994; Tutin et al., 1994). 

Besides being predated by member of the endemic megafauna, the stem apex of some 

palms in Africa is also colonised by a beetle larva, Rhynchophorous phoenicis 

(Coleoptera: Curculionidae) 3 . This larva, also referred to as “spear borer”, is 

particularly associated with the oil palm (Elaeis guineensis) and is a significant 

problem in commercial plantations where heavy infestations can lead to the death of 

the individual 4 (Opeke, 1997). The adult female lays its eggs in wounds or soft areas 

of tissue, such as the stem apex. The eggs hatch after 4-7 days and the larva then bores 

into the interior of the stem where it remains feeding for around 30 days, often causing 

significant damage. 

Although previously unrecorded, this larva is also known to attack rattan palms and an 

individual of Oncocalamus mannii 5 collected in Equatorial Guinea was found to be 

colonised by Rhyncophorous phoenicis. The colonisation of the slender terminal bud 

of this individual was significant and would have severely affected the development of 

the stem, if not directly leading to its death. The presence of this borer in rattans could 

have potential implications for the establishment of intensively cultivated systems of 

rattan in Africa. 

4.2.2 Leaflet predation 

Although the leaflets of most rattans are rather spiny they are, somewhat surprisingly, 

consumed by local people, both for nutritional and medicinal purposes (see Appendix 

3 A beetle of the same genus, Rhynchophorus schach, is a particular pest of rattans in SE Asia and has 

caused significant damage to commercial plantings of Calamus manan and C. merrillii (Zakaria et al., 

1992). 

4 When commercial plantations are felled, the palm heart of the oil palm is often found to be colonised 

by these larvae. These are removed, roasted and sold at the roadside for human consumption. 

197

1 for further elaboration). The leaflets of certain species of rattan are also consumed 

by primates. In particular, Williamson, et al., (1990) record how gorillas in the Lopé 

reserve in Gabon have been seen to “process” the leaflets of Eremospatha cabrae, by 

“pulling the leaflets through the teeth, leaving the ribs behind”. The young leaves of 

Eremospatha haullevilleana are also eaten by gorillas, albeit whole (Yumoto et al., 

1994). Many informants I have worked with state that the young, delicate spear leaf of 

Oncocalamus tuleyi and O. mannii, is also reported to be consumed by primates, 

particularly chimpanzees. 

4.3 SEED DISPERSAL 

Aside from destructive herbivory, animals also play an important role is the dispersal 

of the seeds of many rattan species. The baccate fruits of the taxa of African rattan, 

some of which can be somewhat sweet-tasting, suggests that they are animal and avian 

dispersed and it is clear that many animals, both mammal and avian, play a role in the 

distribution of rattan (Corner, 1966). 

Elephants are known to consume vast quantities of rattan seed when they are mature 

and will consume mature fruits falling to the ground. Elephants will also drag 

individual fruiting stems from the canopy to retrieve the developing fruits. Initial 

observations suggest that elephant consumption of rattan fruit result in the least 

amount of damage to the ingested seed. In the Korup National Park, Cameroon, I have 

found large quantities of Laccosperma seed in fresh elephant spore, none of which 

were physically damaged by this consumption and exhibited normal germination 

when sown. In areas with high concentrations of elephants, such as the Takamanda 

Forest Reserve in Cameroon, large areas of even-aged Laccosperma secundiflorum 

individuals, reaching the end of their establishment phase, have been observed 

occurring in high concentrations (author pers. obs.). This is a relatively uncommon 

occurrence and, given the nature of dispersal, it is more common that for most rattan 

species the number of individuals in a given area is relatively low. The congregation 

of high numbers of elephants in Takamanda occurs during the rainy season (June- 

August) when the fruits of bush mango (Irvingia gabonensis) mature and fall to the 

ground in large quantities. Aside from being harvested by local people, elephants also 

5 Sunderland collection number 1903; the larva was collected in spirit for later identification. 

198

feed on these fruits for long periods often creating expansive open areas in the forest 

(Groves and Maisels, 1999). It is in these elephant-created “gap” areas that the large 

populations of developing rattan seedlings have been observed by myself. These are 

undoubtedly created by a combination of concentrated dispersal (i.e. large quantities 

of elephant dung containing rattan seed 6 , being deposited in one locality) and gap-like 

conditions being created that are favourable to the development of young rattan 

seedlings. Although rattans occur widely in forest totally devoid of elephants, this 

concentration of individuals 7 has not been encountered, nor recorded in other forest 

areas. Although Hawthorn and Parren (2000) provide strong evidence to suggest that 

few, if any, plant species are totally reliant on elephants for dispersal, for some 

species, such as L. secundiflorum, the characteristics of the dispersal is arguably 

affected in their absence. 

Primates, notably gorillas, chimpanzees and bonobos, being both terrestrial and 

arboreal frugivores (Happold, 1996) are also known to feed on the fruits of some 

rattan species (Tutin et al., 1994; White and Abernethy, 1997) as are drills 

(Mandrillus leucophaeus) and mandrills (Mandrillus sphinx). However, by the very 

nature of their feeding habits it is somewhat uncertain that primate dispersal of seed is 

as benign or beneficial as has been suggested, as the foodstuffs of these larger 

primates are often chewed before swallowing and thus damaging or destroying the 

seed itself (Martin, 1990; Kingdon, 1997). This will undoubtedly affect the ability of 

the swallowed (and subsequently excreted) seed to remain in tact and germinate. 

Although very few studies on this have been undertaken with regard to rattans, to shed 

some light on the impacts of primate feeding on potential dispersal, a series of simple 

(and very informal) experiments were undertaken over a period of 2-3 months in 

1997. The experiments consisted of recording the feeding habits of captive primates 8 

when presented with a sample of mature rattan fruits. Each of the endemic African 

rattan genera was represented in the trials. The results of this experiment are presented 

below. 

6 

Most species of Laccosperma also fruit at the same time as the bush mango. 

7 

See Chapter 3 for further discussion of stocking of rattans in Takamanda. 

8 

All of the primates concerned were born in the wild and were separated from their family group when 

still young through hunting activities. 

199

Table 11. Effects on seed survival (and potential dispersal) of feeding selected rattan fruits to 

captive primates. 

Rattan genus (& fruit 

characteristics) 

Laccosperma (seed 

“coffee-bean” like, 1cm 

long x 0.6cm wide with 

thin fleshy mesocarp) 

n = 150 

Eremospatha (seed, 

flattened, 2-3cm long, 1- 

2cm wide, thick fleshy 

meoscarp) n = 80 

Oncocalamus (seed 

globose 1.5cm in dia., 

wide medium fleshy 

mesocarp) n = 120 

Chimpanzee 

(Pan troglodytes) 

n = 15 

Mesocarp 

removed & eaten; 

seed swallowed 

whole (100%) 

Mesocarp 

removed and 

eaten; seed 

discarded (100%) 

Mesocarp 

discarded; seed 

chewed before 

swallowing 

(100%) 

Gorilla (Gorilla 

gorilla gorilla) 

n = 6 

Mesocarp 


seed swallowed 

whole (100%) 

Mesocarp 

removed and 

eaten; seed 


Mesocarp 


chewed before 

swallowing 

(100%) 

200 

Drill (Mandrillus 

leucophaeus) 

n = 9 

Mesocarp 


seed chewed 

(85%) before 

swallowing or 

seed discarded 

(15%) 

Mesocarp 

removed and 

eaten; seed 


Mesocarp 


chewed before 

swallowing 

(100%) 

Notes 

Good chance of 

seed survival, and 

excellent dispersal 

from parent 

Excellent chance 

of seed survival, 

although poor 

dispersal from 

parent 

No chance of seed 

survival; poor 

dispersal 

The striking aspect of this experiment is the fact that amongst the members of the 

primates represented the feeding behaviour and response to each fruit type was 

significantly uniform. Although the seed survival rates were highest amongst the 

species of Eremospatha, the fact that the seeds are immediately discarded suggests 

that dispersal from the parent is relatively short in terms of distance. As many species 

experience high rates of mortality beneath parents (Chapman and Chapman, 1995), 

primates as a dispersal mechanism for Eremospatha would be relatively ineffective. 

The fruits of Oncocalamus seem to be more highly prized for their endosperm than 

their fleshy mesocarp and all the seeds of this genus were destroyed by the primate 

“dispersers”. Only members of the genus Laccosperma were swallowed whole, 

probably due to their relatively small size. These seeds pass through the intestinal tract 

undamaged and, as such, are able to travel long distances in their host ensuring 

widespread dispersal. 

A number of species of hornbill are well known as being important dispersal agents of 

rattans (White and Abernethy, 1997; Whitney et al., 1998). In particular, species of

the genus Ceratogymna have been recorded to swallow whole seeds of Eremospatha 

macrocarpa and Laccosperma secundiflorum, without causing damage to the seed 

itself 9 (Whitney et al., 1998), although subsequent germination tests revealed a 

slightly deleterious effect on germination rates (ibid.). This ingestion and subsequent 

range movements suggests that dispersal of the seeds occurs for relatively long 

distances. The importance of flowering behaviour and seed dispersal is also discussed 

in Chapter 5. 

Table 12. Summary of recorded herbivory and dispersal of African rattans (from personal 

observations and literature cited above). 

Rattan species Plant part eaten Herbvivore / disperser 

Eremospatha cabrae Leaflets 

Fruit / seed 

Palm heart 

E. haullevilleana Leaflets 

Palm heart 

201 

Gorilla 

Gorilla, chimpanzee 

Chimpanzee 

Gorilla 

Gorilla, chimpanzee, bonobo 

E. hookeri Palm heart Gorilla, chimpanzee 

E. macrocarpa Fruit / seed Hornbill, drill, mandrill 

E. wendlandiana Fruit / seed Hornbill, drill 

Laccosperma 

Fruit 

Gorilla, chimpanzee, hornbill 

secundiflorum Palm heart 

Elephant, bonobo 

L. robustum Fruit / seed 

Gorilla, chimpanzee, mandrill, hornbill 

Palm heart 

Elephant, chimpanzee, bonobo 

L. opacum Fruit / seed 

Gorilla, chimpanzee, hornbill 

Palm heart 

Elephant, gorilla, chimpanzee 

Oncocalamus mannii Fruit / seed 

Mandrill 

Palm heart 

Spear borer 

O. tuleyi Fruit / seed Drill 

Calamus deërratus Palm heart Gorilla, chimpanzee 

4.4 SEED PREDATION AND CACHING 

The fallen fruits of Laccosperma are often predated upon by rodents, particularly 

brush-tailed porcupines (Atherurus africanus). The relatively small size of the seeds of 

most members of this genus makes them ideal for rodent consumption. The tendency 

for these terrestrial frugivores to cache seeds for later consumption can occasionally 

provide the opportunity for some of the seeds to germinate if they are left for long 

enough in the conditions most appropriate for germination. In this regard, it is 

common to find large numbers of seedlings of Laccosperma germinating very closely 

together in cached groupings. However, these caches are never far from the parent 

individual.

4.5 ANT / RATTAN ASSOCIATIONS 

It has been long reported that ants and rattans share, what is speculated to be a 

mutually beneficial relationship (Bequaert, 1922; Ridley, 1910; Corner, 1966; 

Whitmore, 1990; Dransfield, 1979; Tomlinson, 1990; Dransfield and Manokaran, 

1994; Tuley, 1995). However, few studies have attempted to determine the nature of 

the relationship, nor take stock of whether there really is a mutual benefit to having 

ants colonise many rattan taxa. 

There are numerous recorded examples where ants and plants have formed close, 

often mutually beneficial, relationships (Janzen, 1966; 1969; 1972; Bentley, 1976; 

1977; Lock, 1986; Huxley, 1986). Such relationships are often quite distinct from 

other forms of plant/animal interaction. Whilst other organisms may consume, 

pollinate or disperse plants, ants are often, although not always, conspicuously absent 

from these activities. Despite this, it has been argued that ants are the organism most 

commonly found to play an integral role in the interaction between the plant and its 

immediate environment (Huxley, 1986). 

It has been postulated that ants and plants have a relationship based mainly on defence 

(Huxley, 1986) and any rattan collector, be they botanical or commercial, can vouch 

for the ferocious defence that ants provide to protect their rattan hosts. In addition to 

this defence role, some evidence suggests that ants might also play a role in the 

provision of nutrients to the host (Rickson, 1979; Rickson and Rickson, 1986). In 

studies of Daemonorops verticillaris and D. macrophylla in Asia, Rickson and 

Rickson (1986) found that the nutrients from accumulated plant debris, were absorbed 

by the ant nest material (which itself is comprised of a combination of spines and 

plant hairs). A further adaptation which has been identified on rattan palms, is the 

presence of scale insects (Coccus spp.) on the sheath that appear to be “farmed”, by 

the ants (Dransfield, 1979; Whitmore, 1990). The excretions of the sweet honeydew 

of the scale insects is used to nourish the developing ant pupae and both the scale 

insects and plant host are then protected vigorously by the ant colony. In Africa, this 

9 

Although some fragments of the seed coat of Eremospatha macrocarpa were encountered in nest 

traps (Whitney et al., 1998). 

202

latter situation has been observed on Laccosperma robustum and Eremospatha 

hookeri in particular (author pers. obs.). 

The simplest and most widespread adaptations of plants that attract ants are the 

presence of extra-floral nectaries; glands that manufacture rich sugary compounds and 

amino acids outside of the floral structure (Bentley, 1976; 1977). However, many 

plants, including rattans, produce chambers that are used by ants either for feeding, or 

for nest sites, or for both. These “little houses” are called domatia (Huxley, 1986), 

although they are also referred to as formicaria or myrmecodomatia (ibid.). Domatia 

may be either deciduous, where shelters are quite small and delicate in structure, or 

are permanent, and these are often characterised by the colonisation of the stem by 

ants. The colonisation of the hollow stems of Barteria fistulosa by Pachysima 

aethiops ants is a good example of the development of permanent domatia (Janzen, 

1972) as is the swollen stem of Mymecodia tuberosa, colonised by the ant species, 

Iridomyrmex cordatus (Huxley, 1978). 

Although palms in general are not known for their close association with ants, the 

rattan palms have been widely reported as providing, what are now more commonly 

considered as deciduous domatia for ants to colonise (Beccari, 1884-86; Bequaert 

1922; Ridley, 1910; Corner, 1966; Whitmore, 1990; Dransfield, 1979; Tomlinson, 

1990; Dransfield, 1992b; Tuley, 1995). The unique morphology of the scandent palms 

within the Calamoideae provides a number of specific domatia that ants are known to 

inhabit. In the case of some species of Asian rattan, for example, Corner (1966) 

records the presence of ant colonies in the inflated ocrea of Korthalsia echinometra 

and in the interlocking verticillate spines on Daemonorops verticillaris. Dransfield 

(1979) provides further examples of ant colonisation on certain rattan species 

including colonisation of the reflexed lowermost leaflets of Calamus laevigatus and C. 

javensis and the curious leaf sheath auricles of Pogonotium ursinum. 

Despite some fundamental differences in both rattan morphology and the diversity of 

the ant fauna (Brown, 1973), many of the morphological features that provide ant 

domatia are shared between the Asian and African taxa. As part of this study, during 

203

the collection of herbarium vouchers, ant samples were also collected and preserved in 

spirit 10 . Extensive notes were taken on the specific domatia they inhabit. 

The following domatia have been identified on some members of the African rattan 

taxa: 

Casual relationships 

Some species of ant construct domes of “carton” on the leaf sheaths of some rattan 

species. These domes are made from plant debris and ant saliva and are supported by 

the spines of the leaf sheath. The enclosed space below is inhabited by the ant colony, 

or may also be used for the farming of scale insects. This type of colonisation has 

been observed on Laccosperma robustum and L. secundiflorum. A further example of 

a casual relationship is the weaving of a developing leaf to form an in-rolled tubular 

organ, which is then colonised by ants. This has been observed on the developing 

leaflets of L. laeve. 

Adaptations involving leaflets 

In rattan species where the leaf is sessile, particularly in the genus Eremospatha, the 

lowermost leaflets are often reflexed back across the stem. The enclosed chamber 

formed by this leaflet canopy is frequently colonised by ants or is used for scale insect 

husbandry. This structure is also sometimes fortified with “carton”. 

Ocreas 

The dry, elongated ocreas of some species of Laccosperma are often colonised by 

ants. In some species of Oncocalamus, which have much shorter truncate (and rather 

fleshy) ocreas, there is also significant ant colonisation. 

Inflorescence bracts 

The inflorescence of the majority of palms is covered with numerous imbricate bracts. 

In some species of Laccosperma, the tubular nature of the bracts sheathing the 

inflorescence provides a small, enclosed area, which is sometimes inhabited by ant 

colonies. 

10 The ant specimens were identified by Dr Stefan Cover of Harvard University. 

204

Hollow sheaths 

Some species of ant chew into the outer layer of the leaf sheath and inhabit the 

enclosed area between it and the inner stem core. This has been observed in young 

sheaths of L. opacum. 

Hollowed out acanthophylls 

The bulbous base of these organs is entered through a hole created by the colonising 

ants. The ants then inhabit the hollowed-out organ. This has been observed in all three 

large-diameter species of Laccosperma. 

Although it has not yet been possible to represent all the observed situations listed 

above, Table 13 summarises the findings of the samples taken to date. 

Table 13. African rattan hosts and ant colonies 

Rattan species Domatia Ant species 

Eremospatha cuspidata 


Lowermost leaflets Crematogaster sp. 1 

Eremospatha hookeri (Sunderland 

1906) 


Eremospatha hookeri (Sunderland 

1917) 

Lowermost leaflets Oecophylla longinoda 

Eremospatha laurentii 



Eremospatha laurentii 


Lowermost leaflets Polyrachis regesa 

Eremospatha macrocarpa 


Leaf junction Crematogaster sp. 1 



Lowermost leaflets Polyrachis fissa 



Lowermost leaflets Tetramorium aculeatum 

Eremospatha wendlandiana Lowermost leaflets Pheidole sp. (minor workers 


only) 

Eremospatha tessmanniana Leaf junction Crematogaster sp. 1 


Atopomyrmex mocquerysi 

Laccosperma secundiflorum 


Ocrea Crematogaster sp. 1 

Laccosperma acutiflorum Inflorescence (under rachis Crematogaster sp. 2 


bracts) 

Oncocalamus macrospathus 


Ocrea / lowermost leaflets Oecophylla longinoda 

Oncocalamus mannii (Sunderland Ocrea / lowermost leaflets Pheidole sp. 

1768) 

Crematogaster sp. 1 

Oncocalamus mannii (Sunderland 

1923) 

Ocrea / lowermost leaflets Atopomyrmex creyptoceroides 

Oncocalamus tuleyi (Sunderland Ocrea / lowermost leaflets Camponotus sp. 1 

1790) 

Cataulacoccus sp. 1 

Crematogaster sp. 1 

Polyrachis laboriosa 

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The following table shows that each rattan species is associated with between 1 and 4 

different ant species. A chi-square test reveals that there is little significant difference 

in the number of ant species associated with a rattan than would have been expected 

(P 2 = 5.71, P=0.13). 

Table 14. Count of domatia and ant species 

Species Count of domatia Count of ant species 

Eremospatha cuspidta 1 1 

Eremospatha hookeri 2 2 

Eremospatha laurentii 2 2 

Eremospatha tessmanniana 2 2 

Eremospatha macrocarpa 3 3 

Eremospatha wendlandiana 1 1 

Laccosperma acutiflorum 1 1 

Laccosperma secundiflorum 1 1 

Oncocalamus macrospathus 1 1 

Oncocalamus mannii 3 3 

Oncocalamus tuleyi 4 4 

Totals = 21 21 

Given the above data, from the number of rattan species that each ant species visits it 

appears that the majority of the species of ant are only found on one species of rattan. 

There is one exception where there seems to be less of a preference and one species of 

ant, Crematogaster sp. 1, visits up to eight species of rattan. The Chi-square test for 

this data reveals highly significant differences between the ant species in the number 

of rattans species that they use (P 2 = 25.29, P=0.00001). This is clearly due to the fact 

that this species of Crematogaster is a generalist. The remaining species of ant visit 

one or two species, which seems to suggest that there is a fair level of specialisation 

between ants and rattan. 

206

Table 15. Count of domatia and rattan species. 

Ant species Count of domatia Count of rattan species 

Atopomyrmex creyptoceroides 1 1 

Atopomyrmex mocquerysi 1 1 

Camponotus sp. 1 1 1 

Cataulafcccus sp. 1 1 1 

Crematogaster sp. 1 8 8 

Crematogaster sp. 2 1 1 

Oecophylla longinoda 2 2 

Pheidole sp. 2 2 

Polyrachis fissa 1 1 

Polyrachis laboriosa 1 1 

Polyrachis regesa 1 1 

Tetramorium aculeatum 1 1 

Totals = 21 21 

Given the above evidence, it might be argued that ants are more specific about which 

rattans they visit than the rattans are concerned by which species of ants they host. 

This is not surprising given that each rattan genus has its own morphological 

characteristics that influence the colonisation of domatia. Not only does there appear 

to be some level of ant / rattan specificity occurring in the African rattans, the 

relationship also seems to be somewhat symbiotic. The very nature of the morphology 

of rattans means that the ants are provided with hospitable domatia on plant hosts that 

are conveniently spiny and high enough in the canopy to deter all but the most 

persistent of predators. The rattans themselves are then provided with a very effective 

means of defence. The nutrient-related benefits, for example the farming of scale 

insects, are also important for the colonisers. However, whether there are nutrientrelated 

benefits for the rattan species colonised remains, as yet, an unsubstantiated 

possibility. 

4.6. SUMMARY 

As discussed above, there is considerable evidence that rattans in African interact 

widely with their immediate forest environment, particularly with faunal agents. 

Understanding the nature of these relationships is crucial if these taxa are to be 

207

managed in their wide state or promoted through cultivated systems as further 

discussed in Chapter 9. It is important to be able to assess the relative impact, both 

beneficial and potentially detrimental, of these inherent relationships and further 

research could focus on the impacts of some of these relationships in high-stocking 

cultivated systems. 

208

Figure 85. Ant colonisation of leaflets of 

Laccosperma laeve, Ghana 

209 

Figure 86. Ant farming of scale insects on 

Eremospatha hookeri, Ghana

CHAPTER FIVE 

HAPAXANTHY AND PLEONANTHY IN AFRICAN RATTANS 


Hapaxanthy, and its alternate state, pleonanthy, are terms that have long been used to 

differentiate the flowering behaviour in palms. Despite the fact that hapaxanthy and 

pleonanthy have recently been reviewed (Uhl & Dransfield, 1987; Tomlinson, 1990; 

Tucker, 1991; Henderson, in prep.) there has persisted some confusion regarding the 

inflorescence structure and life form of the rattans of Africa. Recent field observations 

have provided further information with regard to the flowering behaviour of the 

African rattans and have confirmed the hapaxanthic nature of Laccosperma and the 

pleonanthic nature of Eremospatha and the sole representative of Calamus in Africa, 

C. deërratus. The genus Oncocalamus, long recorded as being hapaxanthic is now 

known to be pleonanthic. Knowledge of the life form of economically-valuable plants 

such as rattan is essential if rational decisions are to be made about their long-term 

management and sustainable utilisation. 

5.2 HAPAXANTHY & PLEONANTHY: A DISCUSSION 

A number of palms produce what appears to be a massive “terminal” inflorescence 

that, in solitary palms, results in the death of the primary axis. In fact, this structure is 

not terminal (Corner, 1966) but is an aggregate of a large number of lateral 

inflorescence units borne in the axils of often markedly reduced leaves, which may be 

described as antecedent (ibid.). In palms, this condition has been widely termed 

hapaxanthy. In essence, there is little morphological difference between hapaxanthy, 

and pleonanthy, its alternate condition (Uhl & Dransfield, 1987). In pleonanthy, the 

lateral production of inflorescences occurs on the lower portions of the stem which 

continues to grow vegetatively and reproduce over a relatively long period throughout 

its adult life (Tuley, 1965; Uhl & Dransfield, 1987; Tomlinson 1990; pleonanthic 

palms are polycarpic (Dransfield, 1978; Henderson (in prep.)). 

The differences between the two flowering states are ultimately physiological (Uhl & 

Dransfield, 1987; Baker et al., 1999b). In hapaxanthic palms, the primary axis 

undergoes a vegetative phase, which may be up to 50 years in the genus Corypha 

210

(Fisher et al., 1987), followed by a relatively short reproductive phase that results in 

the production of several to many axillary inflorescence units. The true apex of the 

stem aborts and the stem then dies after flowering. In taxa with solitary stems, such as 

in all the species of Corypha, the whole plant dies after a reproductive event and is 

thus monocarpic. However, in multiple-stemmed palms such as some rattans, although 

individual axes can be hapaxanthic, the continued production of basal suckers ensures 

the survival of the individual as a whole (Tomlinson, 1990). Hence hapaxanthy, as 

used by most palm botanists, is not synonymous with monocarpy (ibid.) (i.e. palms 

with hapaxanthic axes can be polycarpic). 

In the majority of palms the inflorescence units expand acropetally i.e. in order of 

their age such that they develop and mature from the base to the apex. However, in 

some palms, notably within the tribe Caryoteae (sub-family Arecoideae), the 

inflorescences develop and mature in the reverse fashion (basipetally) i.e. the younger 

inflorescences develop below the older (often fruiting) axes (Uhl & Dransfield, 1987; 

Tomlinson, 1990). Both acropetal and basipetal inflorescences are encountered in 

hapaxanthic palms, whilst in pleonanthic taxa, the inflorescences are always acropetal 

(ibid.). 

Henderson, (in prep.) has recently applied the terms semelparity and interoparity to 

the palm family. The term semelparity has been applied to describe whole organisms 

that reproduce once only and then die (Cole, 1954; Tomlinson, 1990; Young & 

Augsperger, 1991; Henderson, in prep.). The opposite state, those organisms that 

reproduce many times during their life cycle have been termed iteroparous (ibid.). 

Henderson (in press) argues that the terms hapaxanthy and pleonanthy are somewhat 

obsolete as the lack of morphological difference between the two inflorescence states 

does not warrant such a distinction. However, when clearly defined (e.g. Dransfield & 

Mogea, 1984) hapaxanthy and pleonanthy can be consistently applied to usefully 

describe a distinct feature obvious to all. For this reason, the use of hapaxanthy and 

pleonanthy, are maintained throughout this chapter. 

211

Figure 87. Hapaxanthy and pleonanthy in palms 

A. Single-stemmed pleonanthic palm with acropetal production of inflorescences and potentially 

indeterminate growth. B. The same, but multi-stemmed. C. Single-stemmed (and thus monocarpic) 

hapaxanthic palm with simultaneous production of inflorescences. D. The same, but multi-stemmed and 

thus polycarpic. E. Single-stemmed hapaxanthic palm with basipetal production of inflorescences, the 

distal inflorescences opening before the proximal. F. The same, but multi-stemmed. (Arrows indicate 

continuation of terminal growth; x = termination of stem elongation). 

5.3 GEOGRAPHICAL DISTRIBUTION OF HAPAXANTHY 

With the exception of the Central and South American Raphia taedigera, an otherwise 

African genus, hapaxanthy is limited to the Old World palms. In all, 15 genera of 

palms, the majority of them in the Calamoideae, are wholly or partly hapaxanthic. In 

addition, the majority of hapaxanthic palms are climbers. Within the African rattan 

genera, Laccosperma is hapaxanthic, whilst Eremospatha, Oncocalamus and Calamus 

are pleonanthic. 

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Table 16. Hapaxanthy in the Palmae (adapted from Uhl & Dransfield, 1987) 

Sub-Family Genus No. of spp. No. of hapaxanthic spp. 

Coryphoideae Corypha 8 8 

Nannorrhops 1 1 

Calamoideae Daemonorops 116 1 (D. calicarpa) 

Eleiodoxa 1 1 

Eugeissona 6 6 

Korthalsia 26 26 

Laccosperma 5 5 

Metroxylon 6 5 

Myrialepis 1 1 

Plectocomia 16 16 

Raphia 28 28 

Salacca 15 1 (S. secunda) 

Arecoideae Arenga 22 17 

Caryota 12 12 

Wallichia 7 7 

5.4 THE AFRICAN GENERA 

5.4.1 Laccosperma 

Laccosperma is clearly hapaxanthic and has long been recorded in the literature as 

such (Mann & Wendland, 1864; Wright, 1902; Ridley, 1907; Beccari 1910; 

Hutchinson, 1934; Gineis, 1960; Tomlinson, 1962; Tuley, 1965; Corner, 1966; Moore, 

1971; Dransfield, 1976; Dransfield, 1978; Uhl & Dransfield, 1987; Tucker, 1991; 

Dransfield, 1992; Tuley, 1995; Sunderland, 1998; Sunderland, 1999b). After 

flowering, although the flowering stem itself dies, the individual clump continues to 

produce vegetative growth. From long-term field observations in the Campo Faunal 

Reserve in Cameroon, between 1995-99, it appears that at least one stem from each 

clump produces flowers and fruits each year (author pers. obs.). 

5.4.2 Eremospatha 

Despite being noted as hapaxanthic by Tomlinson (1962b), all species of Eremospatha 

are pleonanthic and the inflorescences are produced laterally, some distance from the 

stem apex. 

213

Figure 88. The lateral inflorescences of Eremspatha cuspidata, near Etembue, Equatorial Guinea 


5.4.3 Oncocalamus 

The inflorescence unit of Oncocalamus was first described as “lateral” (Mann & 

Wendland, 1964). Subsequent descriptions of Oncocalamus also described the genus 

as pleonanthic (Wright, 1902; Hutchinson, 1934). However, almost certainly due to 

the poor quality and incomplete nature of herbarium material, first Tomlinson (1962b) 

subsequently followed by a plethora of other palm workers, described Oncocalamus 

as being hapaxanthic (Dransfield, 1976; Dransfield, 1978; Moore & Uhl, 1982; Uhl & 

Dransfield, 1987; Tucker, 1991; Dransfield, 1992; Dransfield & Manokaran, 1994; 

Tuley, 1995; Henderson in prep.). Recent field work and collection of voucher 

specimens has clarified the morphological picture somewhat and has confirmed that 

all the currently known species of Oncocalamus are indeed pleonanthic and possess 

long, pendulous inflorescences arising laterally some distance from the stem apex. 

214

Figure 89. Lateral inflorescences of Oncocalamus mannii, near Etembue, Equatorial Guinea 


5.4.4 Calamus deërratus 

C. deërratus produces long whip-like inflorescences and, in common with all other 

species of Calamus, is pleonanthic. 

5.5 ECOLOGY AND EVOLUTION 

Hapaxanthy was initially described as an ancestral condition in palms (Holttum, 1955; 

Corner, 1966). However, Dransfield (1976), Moore & Uhl (1982), Uhl & Dransfield 

(1987), and Tucker (1991) consider it a derived condition. The fact that hapaxanthy 

occurs in unrelated genera, as well as the occurrence of both hapaxanthy and 

pleonanthy in the same genus (e.g. Metroxylon, Daemonorops and Salacca), also 

suggests that hapaxanthy arises independently and does not commonly imply a 

common or close ancestor. This has also been proven cladistically by Baker et al., (in 

press). 

215

In ecological terms, Dransfield (1978) and Henderson (in prep.) speculate that the 

hapaxanthic habit has been developed for the colonisation of temporary habitats such 

as light gaps in forest and Dransfield (1978) suggests that: 

“.. hapaxanthy is an adaptation allowing greater possibilities of colonising 

open habitats by the presentation of a large quantity of fruit at one moment”. 

In Africa, however, all three rattan genera (one being hapaxanthic and two, of which 

are pleonanthic) have representative species that colonise forest gaps. In addition, the 

two forest-dwelling species of the hapaxanthic genus, Laccosperma, L. laeve and L. 

opacum occur in deep shade in the lower to mid-canopy. Hence colonisation might not 

be the only advantage of hapaxanthy in particular and the adaptive significance of 

hapaxanthy and pleonanthy might be related to other ecological aspects, such as fruit 

size and the influence of dispersal agents. 

From recent field observations, it appears that one obvious advantage of presenting 

ripe fruit in such a conspicuous manner as with the nature of hapaxanthic 

inflorescences, is that they attract dispersers. In the Rio Muni region of Equatorial 

Guinea, canopy-feeding hornbills (in particular the black and white casqued hornbill, 

Ceratogymna atrata) have been seen by me feeding from the striking hapaxanthic 

inflorescence units of Laccosperma acutiflorum emerging abruptly from the canopy. 

Consequently, very little fruit fall is observed beneath the inflorescence due to the 

high levels of avian feeding of this type (Whitney et al., 1998). Conversely, in 

Eremospatha and Oncocalamus, the ripe fruits are more commonly consumed by 

primates (Gartlan, pers. comm.; author pers. obs.). Being pleonanthic, the 

inflorescence units of these taxa are more commonly produced beneath the vegetative 

structure of the individual plant where primates are able to climb and feed with 

relative ease. Hence, the fruits are not immediately available to hornbills or other 

canopy feeders. This observation is also supported by variation in fruit size. The fruits 

of the hapaxanthic Laccosperma (except for the forest dwelling L. opacum) tend to be 

small (8-12mm in diameter) and more amenable to avian feeders. However, the fruits 

of the pleonanthic Oncocalamus and Eremospatha are relatively large and robust 

(usually up to 15mm in diameter; to between 12-15mm wide and up to 25mm long, 

respectively) and are more likely to be better dealt with by primates. Interestingly, in 

216

the absence of primate dispersers in some forest areas due to over-hunting, there is 

almost 100% fruit fall among the large-fruited pleonanthic species; seedlings are often 

encountered near to the mother plant and there is very little evidence of dispersal of 

any kind. 

Figure 90. Striking hapaxanthic infructescence of Laccosperma acutiflorum emerging from the 

forest canopy, near Mamfe, Cameroon (Sunderland 1714) 

5.6 SUMMARY 

Although, in morphological terms, hapaxanthy and pleonanthy are essentially 

indistinct, the physiological variation in their manifestation means that they remain 

useful features in the description of the habit and life form of palms. These terms, 

when suitably defined, are unambiguous and describe an easily recognised state 

distinguishing many palm species. The clarification of the flowering behaviour of the 

African rattans in this chapter provides useful baseline information that could 

217

effectively contribute to their future management. Knowledge of the life form, 

reproductive nature and the ecological significance of any commercially valuable 

species, such as rattan, is essential if coherent management strategies are to be 

developed and implemented to ensure their sustainable utilisation. 

218

CHAPTER SIX 

INDIGENOUS NOMENCLATURE, CLASSIFICATION AND 


UTILISATION OF AFRICAN RATTANS 

The recording of vernacular names as part of ethnobotanical research has become 

routine so that contemporary works on ethnobotany provide detailed methodologies 

on how to collect local nomenclatural information (Given and Harris, 1994; Martin, 

1995; Alexiades, 1996). Whilst the limitations of studying vernacular names in 

isolation have been recognised (Wilkie and Saridan, 1999), it is known that the careful 

study of plant names reveals a wealth of knowledge on how indigenous people 

perceive and utilise their plant resources (Berlin 1973; Berlin et al., 1973; Berlin, 

1977; Brown, 1977; Hays, 1983; Balée, 1989; Berlin, 1992). It is also proven that 

studies of nomenclature can provide deep insights into the ethnobotanical taxonomic 

structure utilised by indigenous societies (Berlin 1973; Berlin et al., 1973; Berlin, 

1977; Balée, 1989; Berlin, 1992). 

This cognitive approach to the study of ethnobotany has been widely applied and, 

despite considerable regional, ethnic and social differences in the communities 

studied, many researchers have identified similar systems of folk classification found 

in place across the world (Ekandem, 1955; Berlin 1973; Berlin et al., 1973; Friedberg, 

1974; Berlin, 1977; Brown, 1977; Hays, 1983; Balée, 1989; Berlin, 1992). The 

recurrent features of these systems has led to the formulation of basic principles of 

ethnobiological classification utilised by many folk societies (Berlin 1973; Berlin et 

al., 1973; Berlin, 1977; Berlin, 1992) principles that provide useful models for the 

study of how indigenous societies view their plant resources. 

The majority of folk classifications, it is claimed, are based on the universal 

significance of the morphological discontinuities among plants, and rarely, is 

classification based on functional considerations such as cultural utility (Berlin 1973; 

Berlin, 1992). However, utilitarian approaches to classification are also considered of 

significant importance in the understanding of how indigenous peoples relate to 

219

species that are of functional importance and those that are not (Hunn, 1976; Hunn, 

1982; Atran, 1983; Randall and Hunn, 1984; Gianno, 1986; Balée and Daly, 1989). In 

addition, comparison of such “special classifications” with the general-purpose 

classification system utilised by indigenous societies, within the plant kingdom, 

provides significant insight into how useful plants are classified within, and relate to, 

such classifications (ibid.). 

The rattans of Africa are taxa of immense functional value and the classification 

systems of which they are part are based as much in recognition of this functional 

utility as they are on morphological discontinuities. This chapter presents the findings 

of a recent study of a number of rattan and general-purpose classification systems 

within selected languages of the Bantu language sub-group of Central Africa in 

particular, as well as a cursory review of those of the non-Bantu languages within the 

greater Niger-Congo language family of West Africa. Based on extensive fieldwork, 

as well as a broad review of herbarium collections and available literature, a 

comprehensive study of rattan nomenclature and systems of classification is presented 

for the first time. This review is based on the vernacular names listed in Appendix 1. 

6.2 INDIGENOUS UTILISATION OF AFRICAN RATTANS 

As is the case in South East Asia, the rattans of Africa play an integral role in the 

maintenance of the economy of forest peoples and are a crucial component of 

subsistence-based systems (Abbiw, 1990; Davies and Richards, 1991; Falconer, 1994; 

Morakinyo, 1994; Townson, 1995; Defo, 1997; 1999; Defo and Sunderland, 1999; 

Sunderland, 1999a; 1999b; Minga, in press; Tenati, in press). The most important 

product of these rattan palms is cane; this is the stem stripped of its leaf sheaths and 

stem epidermis, although the epidermis is also often utilised for basic weaving. The 

inner stem is solid, strong and uniform, yet it is highly flexible. The canes are used 

either in whole or round form or are split, peeled or cored. This latter raw material can 

then be used as a simple rope or is woven for baskets or other products. The range of 

indigenous uses of rattan canes across Africa is vast. Aside from the extensive use of 

this inner stem, other plant parts of some species of rattan are utilised for a wider 

range of purposes and some plant parts have alimentary, household or medicinal 

purposes. 

220

The extensive nature and wide range of uses for African rattans has led to the 

proliferation of a common misconception, particularly amongst the conservation and 

development community active in many areas in Africa, that all rattans are useful, and 

therefore all rattan species have potential commercial applications. However, this is 

not the case. Whilst there is a substantial use for many African rattan species, and a 

wider, somewhat more spontaneous 1 use for many others, our recent taxonomic and 

ethnobotanical studies indicate that a number of species are not utilised in any way, or 

are only used in the absence of other more desirable species. This lack of utilisation is 

due to inflexibility or a tendency to break when being worked or a tendency of the 

species to produce aerial branches and hence provides cane of limited length. 

1 Spontaneous use indicates immediate short-term usage, regardless of species, such as tying a bundle 

of harvested leaves in the forest, repairing a cutlass handle on the farm. In this case, use is based on 

immediate, or “emergency”, requirements and is not predetermined. 

221

Table 17. The rattans of Africa; taxonomy and utilisation 

Genus Species Description Distribution Cane use Non-cane uses 

Calamus deërratus G. Mann & H. Clustering flagellate species; stems to 20m long up to 35mm in Senegal to Angola, west to Uganda Yes, but only in absence of other, Many 

Wendl. 

diameter; leaves ecirrate 

more desirable species 

Eremosptha barendii sp. nov Clustering; stems to 30m long, up to 25mm in diameter; 

conspicuous knee and bracts on inflorescence 

Southern Cameroon None recorded None recorded 

cabrae de Wild. Clustering; stems to 50m long, up to 25mm in diameter; leaflets Gabon & DR Congo to northern Yes, Few 

obovate; papillose inflorescence 

Angola 

cuspidata (G. Mann & H. Clustering; stems to 15m long, 25cm in diameter; leaflets with Congo Basin Few None recorded 


conspicuous apiculum 

haullevilleana de Wild. Clustering; stems to 25m long, up to 25mm in diameter; ocrea 

striate; leaflets spathulate – ovate 

Congo Basin Yes, highly prized and widely traded Many 

hookeri (G. Mann & H. Clustering; stems to 30m, up to 30mm in diameter; knee Sierra Leone to Gabon Yes, although mostly in absence of Few 


conspicuous, leaflets rhomboid to obovate 

other species 

laurentii de Wild. Clustering; stems to 30m, up to 30mm in diameter; knee Congo Basin with outliers in Upper Few recorded None recorded 

conspicuous; lowermost leaflets clasping stem 

Guinea forest 

macrocarpa (G. Mann & H. Clustering; stems to 50m long, 10-18mm in diameter; juvenile Senegal to DR Congo Yes, juvenile form is the best small- Many 


leaves bifid, adult leaflets linear lanceolate 

diameter cane in Africa. Widely 

traded 

quinquecostulata Becc. Clustering; stems to 15m long, 10mm in diameter Se Nigeria to southern Cameroon Few None recorded 

tessmanniana Becc. Clustering; stems to 100m long (although branching is 

common), up to 15cm in diameter; glaucous grey-green leaflets 

Southern Cameroon to E. Guinea None recorded None recorded 

wendlandiana Dammer ex Clustering; stems to 60m, up to 30mm in diameter; conspicuous SE Nigeria to Gabon Yes, but poor quality cane 

Becc. 

knee and rhomboid leaflets 

Laccosperma acutiflorum (Becc.) J. Dransf. Clustering; stems to 70m, up to 60mm in diameter; yellowish 

appearance; non-pendulous leaflets 

Upper Guinea to DR Congo None recorded None recorded 

laeve (G. Mann & H. Wendl.) Clustering; stems often branching, to 10m, up to 15mm in Upper Guinea to DR Congo None Few 

H. Wendl. 

diameter; leaflet margins unarmed; seeds smooth 

opacum (G. Mann & H. Clustering; stems often branching, to 10m, up to 15mm in Upper Guinea to DR Congo Yes, but poor quality cane Few 

Wendl.) Drude 

diameter; leaflet margins armed; seeds warty 

robustum (Becc.) J. Dransf. Clustering; stems to 45m, 50mm in diameter; leaflets 

conspicuously pendulous, glaucous blue-green 

SE Nigeria to DR Congo Yes, highly prized cane; traded widely Many 

secundiflorum (de Beauv.) Clustering; stems to 30m, up to 35mm in diameter; leaflets Senegal to DR Congo Yes, highly prized cane; traded widely Many 

Kuntze 

sigmoid, dark green 

Oncocalamus macrospathus Burr. Clustering; stems to 35m, up to 30mm in diameter, sheaths well Southern Cameroon to northern No; poor quality cane None recorded 

armed; rachillae bright yellow, seeds smooth. 

Angola 

mannii (H. Wendl.) H. Wendl. Clustering; stems to 30m, 28 mm in diameter, sheaths wellarmed; 

rachillae bright crimson, seeds warty 

Southern Cameroon to Gabon No; poor quality cane None recorded 

tuleyi sp. nov. Clustering; stems to 30m, up to 45mm in diameter, sheaths 

sparsely or unarmed; seeds smooth 

SE Nigera and SW Cameroon No; poor quality cane None recorded 

wrightianus Hutch. Clustering ?; stems to 10m, up to 10mm in diameter; leaflets 

sigmoid 

Southern Nigeria Yes, but for cane rope and twine only Few 

223

Table 18. Summary of the non-cane uses of African rattans (see Appendix 1 for further 

elaboration) 

Species Use Region 

Calamus deërratus Palm heart eaten 

Young shoots roasted and eaten 

Grilled leaves macerated and made into tea to 

promote weight loss and to treat oedema caused 

vitamin deficiencies 

Ash from burned roots used as salt substitute 

Sheath twisted and used to clean cooking pans 

Sheath twisted to make rope 

224 

Ghana, Sierra Leone 

Ghana 

Senegal 

Guinea-Bissau 

Ghana 

Nigeria 

Eremospatha cabrae Base of leaf sheath used as a chewstick DR Congo 

E. haullevilleana Palm heart eaten 

Fruits used for decoration 

Acanthophylls used as fish hooks 

Sap used as arbortifacient 

Congo 

DR Congo 

DR Congo 

DR Congo 

E. macrocarpa Powdered root used to treat syphilis Ghana, Nigeria 

E. wendlandiana Palm heart eaten 

Congo 

Base of leaf sheath used as a chewstick Cameroon 

Laccosperma laeve Roasted roots eaten to improve virility Central African Republic 

L. opacum Sap potable & drunk by forest workers 

Gabon 

Palm heart eaten 

Congo 

L. robustum Palm heart eaten 

Cameroon to Gabon 

Young leaves eaten in stews 

Equatorial Guinea 

L. secundiflorum Palm heart eaten 

Throughout its range 

Young shoots eaten 

Throughout its range 

Sap potable & drunk by forest workers 

Senegal, Gabon 

Tea from young shoots used as vermifuge Ghana, Gabon 

Sap, when mixed with other species, used to 

treat dysentary 

DR Congo 

Oncocalamus tuleyi Base of leaf sheath used as a chewstick Cameroon 

O. wrightianus Base of leaf sheath used as a chewstick Nigeria 

6.3. A BRIEF INTRODUCTION TO LANGUAGE AND HISTORY IN SUB- 

SAHARAN AFRICA 

The Niger-Congo family of languages dominates the forested zone of sub-Saharan 

Africa. From Sénégal to Lake Chad a number of distantly related languages within 

this language family occur (Oliver and Fage, 1975; Iliffe, 1995; Grimes, 1996). These 

non-Bantu languages are differentiated from one another to such an extent that 

through the study of glottochronology, most linguists postulate that they have been 

growing apart for around 8,000 years; at least as long as their speakers have been 

sedentary agriculturists (Oliver and Fage, 1975; Iliffe, 1995). 

In contrast, almost all of the peoples south of a line drawn from the Cross River area 

of SE Nigeria to southern Somalia speak more closely related languages (Vansina,

1990; Iliffe, 1995). These Bantu languages form a sub-group within the Niger-Congo 

family of languages that are relatively homogenous yet are distributed across a vast 

area of sub-Saharan Africa. From linguistic studies, it has been determined that the 

centre of origin of the Bantu languages is probably the Benue region of Nigeria 

(Vansina, 1990; Iliffe, 1995) from which, around 5,000 years ago, this Bantu family 

split into two branches: Eastern and Western (ibid.). The former group moved slowly 

eastwards along the northern edge of the Congo Basin to the great lakes area of East 

Africa whilst western Bantu developed east of the Cross River in the highlands of 

western Cameroon. 

Around 3,000 years ago, the Western Bantu speakers began to migrate slowly 

southwards, reaching as far as, what is now, northern Namibia. During this migration, 

pioneer groups broke off, travelling eastwards up the river valleys through the topical 

forest, settling as far inland as southern Sudan and the western Zambezi. The 

introduction of the banana and plantain, postulated to have occurred around 500 AD 

(Vansina, 1990), made this rapid expansion through the tropical forest possible as it 

provided the means to farm an otherwise inhospitable environment. 

The study of the ancestral Bantu language indicates that at the time of separation, 

Bantu speakers made pottery and had begun to farm, yet had not begun to use metals. 

Both Bantu groups had words for oil palm and yam and their cultivation, although 

there existed no vocabulary for cereal cultivation in Western Bantu until this group’s 

expansion reached the savannah areas of east and southern Africa. Most terms for root 

and tree cultivation are of western Bantu origin and it is clear that the Western Bantu 

vocabulary and system of classification of living things have developed to reflect the 

group’s familiarity with their forest surroundings (Guthrie, 1948; Guthrie, 1969-70; 

Vansina, 1990; Oliver, 1999). 

6.4 BERLIN’S MODEL OF ETHNOBIOLOGICAL CLASSIFICATION 

6.4.1 Introduction 

It has been long recognised that the presence of hierarchically arranged folk 

taxonomies is probably universal and, as such, these are shared by a wide range of 

unrelated human societies (Conklin, 1962; Berlin, 1973; Berlin et al., 1973; Berlin, 

225

1977; Berlin, 1992). This hypothesis has been supported by comparison of the 

classifications of a number of indigenous groups and has led to the development of 

the theoretical model by Berlin and his collaborators (Berlin, 1973; Berlin et al., 

1973; Berlin, 1977; Berlin, 1992). Berlin’s model is based on hierarchy and 

recognises five (sometimes six) mutually exclusive ranks that are logically 

comparable to contemporary scientific approaches to taxonomy. Within this 

framework, the folk classification systems of rattan, encountered through our studies, 

are discussed. 

Figure 91. Berlin’s theoretical model of ethnobotanical classification showing “the schematic 

relationship of the universal ethnobiological ranks and their relative hierarchical positions as 

shown in an idealised system.” (Berlin, 1992). 

6.4.2 The unique beginner (or kingdom) 

Beginning with the highest, or most inclusive, category, the unique beginner (or 

kingdom) is the most general category, which is implicitly recognised by most folk 

societies and distinguishes between the plant and animal kingdoms. However, this 

distinction is commonly of a covert nature for most indigenous societies and explicit 

terms for “plant” or “animal” do not exist in most folk taxonomies (Berlin, 1992). 

Indeed, Berlin (1973) argues that the terms for “plant” or “animal” are quite recent 

constructs citing linguistic investigation that the term for plant in any language is first 

found in Albertus Magnus in the 13 th century, appearing only as recently as 300 years 

ago in French (Berlin, 1973: 267). It is hence unsurprising that in many pre-scientific 

societies the nomenclatural designation for kingdom is covert. 

226

6.4.3 Life form 

Life-form classification has been the subject of much debate, particularly with regard 

to the relative influence of universality versus utility; (Berlin et al., 1973; Brown, 

1977; Berlin, 1977; Hunn, 1982; Atran, 1983; Berlin, 1992; Ellen, 1998). In general, 

life forms are broad classes usually recognised by their distinctive morphology 

(Berlin, 1973; Berlin et al., 1973; Berlin, 1992). Berlin views the life- form categories 

of folk systematics as very similar to those of early classical botanical classification 

(see Atran, 1983) involving the major categories, “tree”, “vine” and “herb” (Berlin, 

1977). Indeed, so universal is the application of these life form categories that Berlin 

suggests that: 

“These three major groupings represent such distinct perceptual discontinuities that their recognition 

may constitute a substantive near-universal in pre-scientific man’s view of the world”. (Berlin, 1977: 

385). 

However, it has been noted that other life forms may also be recognised according to 

their utility or ecological distribution, as well as morphological discontinuities; or 

indeed a combination of these (Hunn, 1982; Atran, 1983; Randall and Hunn, 1984). 

Hence, in most folk taxonomies there may be more than the three major life form 

categories suggested by Berlin. 

6.4.4 Intermediates 

Intermediates are small groupings of generics (see below) that are similar to each 

other through shared morphological or functional characters and are intermediates 

because they fall in between life forms and generics. These are also often referred to 

as “covert categories” as they are often unnamed and not easily undetected in folk 

systematics, or they may be only known by a small subsection of a community 

(Berlin, 1973; Martin, 1995). 

6.4.5 Generics 

These are the most salient categories in folk classification systems and as such are the 

most commonly encountered during initial ethnobotanical surveys. Generics are the 

smallest units in nature that are easily recognised on the basis of relatively large 

numbers of gross morphological characters. The majority of generic taxa within folk 

227

taxonomies are included in (or affiliated with) one of the recognised life form taxa. 

However, there are generic taxa that are sufficiently morphologically distinct or 

economically important to be classified as unaffiliated, that is independent of lifeforms. 

6.4.6 Specific and varietal categories 

Some generics are further divided into specific or even varietal categories. These 

differ from both life-form and generic categories in that they are conceptually 

distinguished from one another on the basis of very few morphological characters. 

The two levels of classification are more widely applied to plants of considerable 

cultural or utilitarian value. 

6.5 METHODS 

Fieldwork for the study of the taxonomy, ecology and utilisation of African rattans, 

was undertaken primarily in Ghana, Nigeria, Cameroon and Equatorial Guinea. The 

field surveys concentrated both on the collection of herbarium voucher specimens to 

provide representative material for the completion of a taxonomic revision of the 

rattans of Africa, as well as the recording of vernacular names and uses for each 

species. Where possible (and where appropriate), vernacular names were recorded in 

the field using a tape recorder and later transcribed and interpreted with the assistance 

of a translator, who were native speakers of the languages in question. English (both 

Standard and Pidgin English) and French were used as lingua francas. 

Further study of existing herbarium voucher material held in herbaria 2 , as well as an 

extensive literature search, provided a plethora of vernacular names that could be 

compared with the actual species the local name referred to. This latter study 

confirmed the presence of significant consistencies within a number of classification 

systems first detected during our own field research. 

For the purposes of this chapter, vernacular names are spelled phonetically but the use 

of the phonetic alphabet has been avoided for ease of reading. Language and tribal 

2 Material was studied from the Libreville, Yaounde, Bata, Kumasi, Gold Coast, New York, Missouri, 

Hamburg, Geneva, Wageningen, Brussels, Florence and Kew herbaria. 

228

names have been standardised by referring to the Summer Institute of Linguistics’ 

Ethnologue (Grimes, 1996) 

6.6 AFRICAN RATTAN FOLK NOMENCLATURE WITH REFERENCE TO 

BERLIN’S MODEL 

6.6.1 The unique beginner or kingdom 

In common with many folk classifications throughout the world, in the Niger-Congo 

language family, there exist no words for “plant” (Guthrie, 1948; Westermann and 

Bryan, 1952; Guthrie, 1969-70) and the classification the plant kingdom is covert i.e. 

this taxonomic rank is not lexically recognised. Although in many languages in the 

western Bantu subgroup a root word for animal, -namma, is commonly encountered, 

this term is polysemous with “meat” (Guthrie, 1948; Guthrie, 1969-70; Sharpe pers. 

comm. 2000). This observation conforms with Berlin’s assertions that in traditional 

societies where a kingdom is named the word s often polysemous with some life form 

class or reflects an overt recognition of function (Berlin, 1973; Berlin, 1992). 

6.6.2 Life form categories 

In the Niger-Congo language family, explicit terms for “tree”, “vine” and “herb” are 

widely recognised life-form categories (Guthrie, 1948; Westermann and Bryan, 1952; 

Guthrie, 1969-70; Oliver and Fage, 1975; Vansina, 1990; Profizi and Makita-Madzou, 

1996) 3 . However, based on both morphological discontinuities and as a reflection of 

cultural utility, a number of additional life-form classifications within the plant 

kingdom are also recognised. In many western Bantu languages for example, at the 

life-form level of classification, domesticated plants are often distinguished from wild 

plants despite shared morphological similarities (Vansina, 1990). In addition to 

domesticated plants, other plants of extreme functional utility are also accorded 

separate life-form categories. For example, “palm” is also recognised as a separate 

life- form category within many folk taxonomies. Despite the fact that the Palmae is 

considered a “natural” plant family and has long been classified by Western science as 

such, many indigenous societies do not share this view and include only arborescent 

3 In the majority of Bantu languages the word for “tree” is polysemous for “wood”; a lexical similarity 

that is extremely widespread (Brown, 1977). Interestingly the Bantu proto-language had a word for 

tree, -ti-, which was also the word for medicine, indicating the historical importance of the plant-based 

approach to medicinal practice (Iliffe, 1995) suggesting that utility has played an important role in the 

development of plant-based vocabularies. 

229

taxa (standing palms) within their palm life-form category (see Ellen, 1998 for a 

discussion of this). Indeed it is commonly the case that the life-form category “palm” 

of many folk taxonomies do not include climbing palms or other palms exhibiting 

significant morphological discontinuities, such as acaulescence (Ellen, 1998). 

For example, Berlin (1977) from his research in Peru recorded that the Aguarana 

recognise “palm” as a life form, distinct from “tree”, “vine” or “herb”. Yet within this 

category, they exclude from this category slender climbing palms (members of the 

genus Chamaedorea) as well as a number of acaulescent species (Berlin, 1977). Ellen 

(1998) also excludes rattans from his discussion of palm life-form categories in a 

number of indigenous societies for the same reason; they are not included within the 

same category in which standing palms occur. As such, rattans are commonly 

classified separately from other palms as a separate life form (Ellen, 1998); indeed the 

word rattan itself is derived from the Malay word rotan, the life form category for all 

climbing palms. 

In the Niger-Congo language family, “palm” is recognised as a separate life form, 

albeit covertly 4 . Yet, as with many other indigenous societies (Berlin, 1977; Ellen, 

1998) this life-form category encompasses the standing palms only (Elaeis, Raphia 5 , 

Phoenix & Borassus) and excludes both the climbing palms, and the small acaulescent 

genus Sclerosperma, and the stoloniferous Podococcus barteri 6 . 

In this Niger-Congo context, climbing palms are recognised as a mutually exclusive 

life form category. This life form category, when applied, is explicit and overt and, as 

such, is commonly named. 

6.6.3 Intermediate and generic categories 

Although rattan as a life-form category is distinct in many non-Bantu languages, the 

Western Bantu languages do not commonly recognise rattans as a distinct life form, or 

if they do, the recognition is covert. In these instances, these societies categorise 

4 Where folk taxonomies have names for this category, it is usually polysemous with the oil palm 

(Elaeis guineensis), reflecting its symbolic and utilitarian importance. 

5 Acaulescent members of this genus are also included under the palm life-form category. 

230

attans within intermediate categories by grouping closely related folk generics (see 

below). The separation of the climbing palms in this manner is based on utilisation, or 

less commonly, morphological discontinuities such as diameter, or the presence or 

absence of spines on the leaf sheath. In these instances, the biological genera 

Eremospatha, Calamus and Oncocalamus, as well as sometimes some small-diameter 

members of Laccosperma are often grouped together at the intermediate level. 

When this intermediate category is recognised, this category excludes the largediameter 

members of the biological genus Laccosperma. These taxa, which represent 

a number of biological species, are highly prized as a source of good quality cane and 

are categorised in many folk classifications only at the generic level. 

6 These latter taxa, whilst possessing considerable utilitarian importance, are unrelated to other palms 

and to each other and can be classified as unaffiliated generics within most Niger-Congo folk 

classification systems in which they are recorded. 

231

Table 19. Life form, intermediate and generic folk classification of rattan canes in selected Niger- 

Congo languages 

Folk name (-root) Language 

(country) 

-ailé (all spp. except large diameter Laccosperma 

spp.) 

-ahike (large diameter Laccosperma spp.) 

232 

Anyin (Côte 

d’Ivoire) 

Language 

sub- 

group 7 

Ethnobiological 

category 

non-Bantu Intermediate 

Generic 

-nwatia (all climbing palms) Akan-Asanti 

(Ghana) 

non-Bantu Life form 

-dekun (all climbing palms) Gun-Gbe (Benin) non-Bantu Life form 

-ikan (all climbing palms) Edo (Nigeria) non-Bantu Life form 

-egbéé (all climbing palms) Yoruba (Nigeria) non-Bantu Life form 

-kogiri (all climbing palms) Fulfulde non-Bantu Life form 

-kwagiri (all climbing palms) Hausa non-Bantu Life form 

-uga (all climbing palms) Igbo (Nigeria) non-Bantu Life form 

e-chié (all spp. except large diameter Laccosperma Denya (Cameroon) Bantu Intermediate 

spp.) 

-gekwiya (large diameter Laccosperma spp.) 

Generic 

-edju (Oncocalamus spp.) 

Oroko language Bantu Generic 

-ndongo (Eremospatha spp.) 

group (Cameroon) 

Generic 

-mekah (large diameter Laccosperma spp.) 

Generic 

-nloun (all spp. except large diameter Laccosperma Bassa (Cameroon) Bantu Intermediate 

spp.) 

-? (large diameter Laccosperma spp.) 

Generic 

-mokolo (small diameter canes) 

Bakossi 

Bantu Intermediate 

-mekah (large diameter Laccosperma spp.) (Cameroon) 

Generic 

-nlon (all spp. except large diameter Laccosperma Bulu (Cameroon) Bantu Intermediate 

spp.) 

-nkan (all Laccosperma spp.) 

Generic 

-nlong (all spp. except large diameter Laccosperma Fang (Equatorial Bantu Intermediate 

spp.) 

-nkan (all Laccosperma spp.) 

Guinea & Gabon) 

Generic 

-mikaana (all climbing palms) Téké (Congo) Bantu Life form 

-kekelé (small diameter canes) 

Zande, Lingala, non-Bantu Intermediate 

-likaw (large diameter Laccosperma spp.) 

Swahili-Zaire (DR 

Congo) 

Generic 

Interestingly, some generic terms are polysemous with the products that are derived 

from the plant itself. Some examples are included in Table 20, below. 

7 Bantu languages are characterised by the possession of root terms that are distinguished into 

singular/plural by independent prefixes. These root terms are commonly shared among related 

languages and it is variation within the prefixes that is reflected in the variation in names for plants, for 

example (see Guthrie 1948; 1953).

Table 20. Selected cane product names (*asterisk marks the product names that are also the 

generic category, or derived directly from them). 

Product Name Language (country) Notes 

Palm heart mekah* Balundu-Bima (Cameroon) Apex of L. robustum 

baa ndanga Téké (Congo) Apex of E. haullevilleana 

mukaana a ngomu* Téké (Congo) Apex of L. secundiflorum 

mukaana a buulu* Téké (Congo) Apex of E. wendlandiana 

ngodji Lomdo (DR Congo) Apex of L. robustum 

Cane and aka Fang (Equatorial Guinea) Cleaned stems of L. robustum / L. 

cane rope 

secundiflorum 

ukpa Ijo-Izon (Nigeria) Split stems of L. secundiflorum 

ukwen Edo (Nigeria) Split stems of L. secundiflorum 

ekwe oya* Igbo (Nigeria) Split stems of L. opacum for tie-tie 

ekwele / akwala Igbo (Nigeria) Split stems of O. wrightianus 

(coarse cordage) 

udo Igbo (Nigeria) Split stems of O. wrightianus (fine 

twine) 

elili Igbo (Nigeria) Split stems of O. wrightianus 

(string or thread) 

apié* Igbo (Nigeria) Cane rope of C. deërratus 

Baskets kenten Akan-Asanti (Ghana) Long baskets made from stems of 

L. opacum 

penja Bakossi (Cameroon) All cane baskets 

mbaka Denya (Cameroon) Farm baskets made from E. 

macrocarpa 

bi-dong Fang (Equatorial Guinea) Fish baskets made from split stems 

of L. robustum & E. macrocarpa 

be-koro Fang (Equatorial Guinea) Fish traps made from split stems 

of L. robustum & E. macrocarpa 

nkeuiñ Fang (Equatorial Guinea) Farm baskets made from split 

stems of L. robustum & E. 

macrocarpa 

maa kutu Téké (Congo) Baskets made from E. 

haullevilleana (baana = small; 

mwana kutu = medium; kiana = 

large) 

6.6.4 Specific categories 

Although some rattan folk genera are monotypic, a considerable proportion of the 

remaining genera are polytypic and are further split into folk specific categories. 

Within the polytypic genera of rattan, the recognition and natural grouping by 

indigenous societies of folk specific taxa often reflects the intrinsic relationship of 

these specifics both to each other and to the ranks superordinate to them. Commonly, 

these are lexically recognised by mean of terms that are based on the language of 

kinship and descent. Such kinship metaphors have been identified in a number of folk 

taxonomies (Hays, 1983; Berlin, 1992) and have been widely encountered in rattan 

classification systems, particularly within the Western Bantu subgroup. 

233

For example, the Anyang of Cameroon, who speak Denya, a language belonging to 

the Western Bantu subgroup, use the intermediate category, e-chié, for all canes not 

included in the group of large-diameter, heavily armed species of Laccosperma. 

However, as commonly the case with intermediate lexicons in other Bantu languages, 

this term is polysemous for “cane rope” and is used, in the folk generic sense, to 

recognise one highly valuable species of small-diameter rattan in particular, 

Eremospatha macrocarpa. All other species of cane in the e-chié intermediate 

category are lexically recognised by the Anyang at the folk specific level, even though 

many are not actually valued as a source of cane, with the lexical designation often 

reflecting kinship metaphors. For example, two other species of Eremospatha, E. 

tessmanniana and E. quinquecostulata, are both referred to as calumé-e-chié, “the 

uncle of cane rope”, in recognition of the poor quality of the cane of these biological 

species. In the same context, E. wendlandiana is referred to the “true” cane rope as 

mua-e-chié or “brother to cane rope”. However, two further unrelated taxa, despite 

being morphologically distinct, are also referred to in relation to the true cane rope: 

Laccosperma opacum; ge-nomé-echié, or “slave to cane rope”: Oncocalamus tuleyi; 

moa-e-chié, or an (undefined) “relative to cane rope”. 

Box 1. Selected nomencalatural relationships within the Denya language group, Cameroon 

calumé-e-chié 

(E. tessmanniana & 

E. quinquecostulata) 

ge-nomé-e-chié 

(Laccosperma 

opacum 

e-chié 

(E. macrocarpa) 

Further examples of such metaphors include the recognition of related species (yet 

separate in biological terms). Laccosperma opacum, a small diameter cane, is called 

npue-nkan by the Fang of Equatorial Guinea; it is considered the “child of” true nkan 

234 

mua-e-chié 

(E. wendlandiana) 

moa-e-chié 

(Oncocalamus 

tuleyi)

(generic term for L. robustum and L. secundiflorum; large diameter canes and widely 

utilised species). Similarly, the same species, L. opacum, is also categorised at the 

specific level by the Mokpwe around Mount Cameroon as liko ko’ko, or “close to 

cane”. 

The application of the language of descent is commonly encountered within taxa that 

have manifestly different morphological characteristics in the juvenile and adult 

phases, particularly reflected in the leaf shape; flabellate when juvenile and pinnate 

when adult. Interestingly, these juvenile stages can be so morphologically distinct, 

they have sometimes been described by Western botanists as “new” species (Drude, 

1895). A number of these species are widely utilised in the juvenile stages, but are too 

inflexible for use when mature; for example Eremospatha macrocarpa and E. 

haullevilleana. The recognition within some folk taxonomies of this within-species 

morphological variation and distinct utilisation is effected by using what are different 

specific level terms for the juvenile and adult growth phases. All of the prefixes listed 

in Table 4 are glossed as “child of”. 

Table 21. Folk specific taxa based on morphological and utilisation characteristics. 

Species Species name Species name (adult Language (country) 

(juvenile phase) phase) 

Eremospatha 

koto-mbalu mbalu Mende (Sierra Leone) 

macrocarpa 

asa-nlong nlong (semi adult); 

adult = ongam 

Bulu (Cameroon) 

bana-ndongo ndongo Balundu-Bima 

(Cameroon) 

6.6.5 Varietal categories 

Although varietal categories are well recorded for a few African palms, particularly 

the oil palm reflecting its long history of farmer-led selection and breeding (see 

Burkill, 1997), varietal categories for rattan have not been encountered. 

Table 22, below, presents a summary of rattan classification in Africa with reference 

to Berlin’s conceptual model. The theoretical structure of vernacular names is 

presented in Box 2, below. 

235

Box 2. The structure of vernacular names (modified from Berlin et al., 1973) 

PRIMARY NAME 

(Semantically unitary) 

SIMPLE 

(composed of single constituent) 

LEXICON COMPLEX 

(name) (composed of 

or more constituents 

SECONDARY NAME 

(Semantically binary expression with higher 

category named) 

236 

PRODUCTIVE 

(Higher category named) 

UNPRODUCTIVE 

(Higher category unnamed) 

Table 22. Summary of rattan classification within the Niger-Congo language group with 

reference to Berlin’s model. 

Rank Type of name Non-Bantu languages Bantu languages 

Kingdom Unnamed Covert Covert 

Lifeform Primary name or Rattan life form Not recognised or 

unnamed 

category named covert 

Intermediate Primary name or Commonly none (but Commonly two 

unnamed 

see Table 2) 

members (but see Table 

2) 

Generic Primary name 2-3 members 4-6 members 

Specific Usually secondary No generics contain Most generics contain 

name 

known specifics specifics 

Varietal None None recorded None recorded 


It is known that rattans are considered by the peoples within the Niger-Congo 

language family as life forms lying outside the standard life-form classifications of 

“tree”, “vine” and “herb” ((Berlin, 1973; Berlin et al., 1973; Berlin, 1977; Berlin, 

1992). As we have seen, this is undoubtedly due both to morphological discontinuities 

and to considerations of functional utility. As is the case with other indigenous 

classification systems that regard the climbing palms as separate from standing palms 

(Berlin, 1977; Ellen, 1998), throughout their range, African rattans are accorded life 

form status of their own, whether covertly or overtly recognised.

In the case of many of the non-Bantu language groups that predominate in the forests 

of Upper Guinea, the hierarchical classification of rattans is relatively straightforward 

and closely adheres to Berlin’s model. The fact that the Upper Guinea forests are 

relatively species-poor in terms of rattan means that somewhat parsimonious 

classification systems are encountered, with very few categories recognised below the 

generic. Life-form categories within these classifications are named, are clear and 

unambiguous, and refer to all palms with the climbing habit. At the generic level in 

particular, it is also clear that there is considerable correspondence between 

indigenous classifications and Western taxonomy (see Table 6). In this case it may be 

argued that morphological considerations take precedence in the non-Bantu folk 

taxonomies for rattan 8 . 

Table 23. Selected examples of parity of generic categorisation of African rattans between folk 

taxonomic systems and Western classification. 

Language (Country) Laccosperma Eremospatha Calamus Oncocalamus 9 

Loko (Sierra Leone) kafo mbalu tambe n/a 

Akan-Asanti (Ghana) ayie mfia demmere n/a 

Ijo-Izon (Nigeria) ukpa boru apie n/a 

In contrast, the Bantu language subgroup, the use of intermediate categories is more 

widespread and the life-form category for rattan is covert and, as discussed, the 

climbing palms are essentially subdivided into two main categories and commonly, 

one intermediate and one to several generic categories are recognised. This may be a 

reflection of both increased rattan diversity (biologically) as well as increased ethnic 

complexity (Brown, 1977), particularly the region from SE Nigeria to northern 

Gabon, the most biologically and linguistically diverse area of Africa. 

Berlin (1973) suggests that intermediate categories are rare in folk taxonomies and 

that, because they often lack names, some doubt has been expressed as to whether 

they might be included as an ethnobiological rank at all. However, the explicit use of 

8 However, examples that lean towards functional utility being a major factor are also encountered. For 

example, within the genus Laccosperma, L. opacum and L. laeve are treated as generics within most 

non-Bantu classifications and, despite their close morphological similarities are often distinguished 

based on their functional utility. In Twi, L. laeve, which is not used by that ethnic group, is called tehan 

muhunu or “it lives in the world for nothing” whilst L. opacum is used for making specific baskets and 

is called sayai. 

9 The genus Oncocalamus does not occur in the Upper Guinea forest region. 

237

intermediate categories for African rattans, particularly in the western Bantu language 

subgroup, is evidence to the contrary. It is likely that the widespread use of the 

intermediate category is indicative of the recognition of functional utility over 

morphology. 

More specific examples can further illustrate this point. The use of an intermediate 

category for what are biologically separate groups is widespread within the Western 

Bantu subgroup, and it is important to note that the name given to this intermediate 

category is commonly polysemous with the generic root name for the most widely 

utilised rattan species, Eremospatha macrocarpa. Further splitting at the folk specific 

level relates each folk taxon with this species, often through kinship metaphors or 

other relational terminology, regardless of often significant, morphological 

differences between these folk specifics. The nomenclature used is based on relating 

each species to the main taxa that are useful (in this case Eremospatha macrocarpa) 

and those others that are not as widely used. 

The most obvious example of the importance of functional utility in indigenous 

classification of rattan is with regard to the large-diameter species of Laccosperma. 

Although Laccosperma was previously considered a single species, it is now 

recognised that there are in fact three biologically distinct species of large-diameter 

Laccosperma: L. secundiflorum, L. acutiflorum and L. robustum. However, most folk 

taxonomies recognise only one, and sometimes two, generics for this group of 

biological species, despite significant and easily observable morphological 

discontinuities obvious in the field. This is because L. secundiflorum and L. robustum 

provide the best quality cane in Africa. They are widely utilised at the subsistence 

level as well as providing the basis of a thriving trade. Although traded in the same 

way, these species are obviously different when encountered in the field and, being 

sympatric, these differences are clear to see. L. secundiflorum has broad sigmoid 

leaflets that are held more or less horizontally and possesses a relatively long petiole; 

it is a species found in the forest understorey and at forest margins. L. robustum has 

fine linear leaflets that are almost completely pendulous on the rachis; this species 

occurs in forest gaps and is extremely light demanding. Yet over 100 informants have 

told me that the species “are the same” and as such are included under a single generic 

category in local classifications. 

238

It is clear that within these two main classification systems discussed above both 

morphology and functional utility are considered in relation to African rattans. 

However, the differences between the two approaches can be considered cognitive. In 

the case of the non-Bantu classifications, it is “climbing palm”, and the morphological 

differences within that group that are the important criteria for classification; in the 

Western Bantu classifications, it is the aspects of functional utility i.e. “cane”, and 

how taxa relate to each other in that functional context, that take precedence. 

Although equal correspondence has been found to occur between folk classification 

and scientific taxonomy at the generic level in some classifications, this parity is not 

commonly encountered in the case of African rattans. As discussed, within rattan folk 

systematics, under-differentiation of western species recognised in Western taxonomy 

tends to be more commonly observed. However, limited over-differentiation is also 

encountered, particularly when juvenile stages are lexically distinguished from adult 

stages in specific folk taxa. 


Understanding how local people both view and use their resources is often a greatly 

under-valued means of developing integrated sustainable strategies for forest and 

resource management. When complimented by biosystematic data, the study of local 

classification systems of indigenous groups can provide considerable insights into 

which species are used and why, and how, in these functional respects, species relate 

to each other. In the case of African rattans, such an approach has been critical in 

providing clear information on the most utilised taxa and which species could be 

targeted for promotion and development in the context of sustainable development. 

This is particularly pertinent for the plethora of development and government 

agencies that are currently calling for the promotion of “rattan” to contribute to the 

improved livelihoods and wider conservation of the forests of West and Central 

Africa. It is essential that such initiatives first identify which species are of particular 

importance and then focus on the few species worthy of such investment. Now that a 

sound taxonomic base has been provided through the study of nomenclature and use, 

the utilisation of each biological species is now more comprehensively known, it is 

239

hoped that any future development of the rattan resource might be able to take place 

in a coherent and structured manner. 

240

Figure 92. Temporary camp for forest product exploitation in Mokoko, Cameroon. Note the 

rattan basket being fabricated in the foreground 

Figure 93. Cut stems of Laccosperma robustum, tied and ready for transportation. Mokoko, 

Cameroon 

241

242

Figure 94. Fish trap made from split stems 

of Laccosperma robustum, near Bata, 


Figure 95. Cane bridge made from juvenile 

stems of Eremospatha macrocarpa, Nyang, 

Cameroon 

242 

Figure 96. Weaving farm basket from split 

stems of Laccosperma spp. Mfeck-Ayong, 


Figure 97. Village-fabricated market basket, 

made from stems of Eremospatha 

macrocarpa Cogo, Equatorial Guinea

CHAPTER SEVEN 

A SOCIO-ECONOMIC PROFILE OF THE COMMERCIAL RATTAN 


TRADE IN CAMEROON 

Rattan and rattan products have long been regarded as one of the major non-timber 

forest products of the forest zone of Cameroon (Shiembo, 1986; Pokam-Wadja, 1987; 

Sunderland, 1999a; 1999b; Defo, 1999). Although both unprocessed rattan and 

finished products are widely traded and form the basis of a thriving cottage industry, 

this trade is a component of the “invisible” markets for forest products that are only 

now being understood (Clark and Sunderland, 1999; Ruiz-Perez et al., 2000). It is 

only in recent years that non-timber forest products (NTFPs) such as rattan have 

become the focus of research and development initiatives that are concerned with the 

increased valorisation of a wide range of these “minor” forest products. Indeed, this 

paradigm shift has been so marked that non-timber forest products are now regarded 

as having a significant role to play in contributing to conservation and development 

initiatives through product promotion and sustainable development (Wilkie, 1999). It 

has been argued, however, that in order for this to happen, the promotion and 

development of high value NTFPs must take place in the context of appropriate forest 

legislation. This can then provide the framework that allows the equitable distribution 

of benefits, community participation in resource management and the realisation of 

forest product-generated revenues 1 (Wilkie, 1999; Laird, 1999; Profizi, 1999; 

Cunningham, 1999). 

7.2 THE MARKETS FOR NON-TIMBER FOREST IN CENTRAL AFRICA 

Although the historical importance of the trade in forest products in West and Central 

Africa has been well recorded (Oliver and Fage, 1975; Liniger-Gomez, 1986; 

Vansina, 1990; Iliffe, 1995; Oliver, 1999), it is only relatively recently that the scale 

and importance of contemporary markets for forest products in Central Africa has 

1 Indeed, the 1995 National Forestry Action Programme of Cameroon (Project 59) proposes that the 

“...inventory, silviculture and further development of the rattans of the rain forests of Cameroon” takes 

place to improve income generation for both rural and urban communities that rely on rattan for their 

livelihoods. 

243

een realised (Ndoye, 1994; Falconer, 1994; Clark and Sunderland, 1999; Sunderland 

and Obama, 1999; Liengola, 1999; Yembi; 1999; Kimpouni, 1999; Tabuna, 1999; 

Ruiz-Perez et al., 2000). Indeed, studies of selected markets within Ghana (Falconer, 

1994; Holbech, 2000) Cameroon (Ndoye et al., 1999), Equatorial Guinea (Sunderland 

and Obama, 1999), Congo (Kimpouni, 1999), Gabon (Yembi, 1999) and DR Congo 

(Liengola, 1999) show that the trade in non-timber forest products is significant. 

Furthermore, this trade is often regional, rather than national, in scope, with a great 

deal of cross-border trade in forest products, particularly focussed on Cameroon. For 

example, there is significant NTFP trade from Cameroon to Nigeria, (Bokwe and 

Ngatoum, 1994; Sunderland and Tchouto, 1999), Cameroon to Equatorial Guinea 

(Sunderland and Obama, 1999) and from Cameroon to Gabon (Yembi, 1999; Ndoye 

et al., 1999). There is also substantial export of non-timber forest products from the 

wider Central African region to supply ex-patriot Africans based in Europe and North 

America. The cultural importance of these products is such that consumers are willing 

to absorb the extra costs of packaging, export and transportation despite the local 

availability of cheaper substitute products (Tabuna, 1999). 

In short, the markets for forest products in West and Central Africa are robust, 

demand-led and support the livelihoods of many thousands of people; from the forest 

harvester to the urban trader (Ruiz-Perez, et al., 2000). As such, the need to “develop 

the markets for NTFPs” as has been advocated for forest conservation efforts in Latin 

America (Padoch, 1987; Padoch, 1992; Clay, 1992; Gentry, 1992; Richards, 1993) 

and SE Asia (Peluso, 1992; Gan and Weinland, 1996) is unnecessary in the African 

context as there is considerable evidence that the sector is significantly market driven 

(Clark and Sunderland, 1999; Ruiz-Perez et al., 2000). What is essential, however, is 

the recognition and evaluation of those markets allows the capture of benefits such 

that these forest resources are able to contribute to the formal forest sector of the 

countries in which they originate. 

7.3 THE RATTAN TRADE IN CAMEROON 

Recent studies of the African rattan trade have concentrated on the importance of the 

trade within individual countries (Falconer, 1994; Morakinyo, 1995a; Defo, 1997; 

Sunderland 1998; Defo 1999; Oteng-Amoako and Obiri-Darko, in press). 

244

Interestingly, the patterns of exploitation and trade of the rattan resource are 

remarkably similar in each of the countries studied and distinct similarities within the 

sector are apparent (see Chapter 8). In Cameroon, rattan has long been identified as 

an extremely important forest product, both at the household level (Shiembo, 1986; 

Thomas et al., 1989; Defo, 1997; Trefon and Defo, 1998; Sunderland, 1999a; 1999b; 

Defo, 1999) as well as being widely traded (Shiembo, 1986; Pokam-Wadja, 1987; 

Sunderland, 1999a; 1999b; Defo, 1999). This chapter presents the findings of a study 

of the commercial rattan trade in Cameroon, and the socio-economic conditions under 

which this trade operates. 

7.4 METHODOLOGY 

7.4.1 Selection of study sites 

Concentrated in the southern half of the country, the forest zone of Cameroon 

comprises an area of some 20 million hectares and encompasses a wide range of 

cultural and socio-political conditions. The rattan trade, artisan industry and markets 

in 15 of the most representative urban areas were studied. The study sites were 

selected on a combination of cultural and economic importance, as well as a 

consideration of the proximity and importance of the rattan resource. In addition, two 

towns outside of the forest zone (Bamenda and Bafoussam) where the transformation 

of rattan is of particular importance were also sampled. Additional surveys were 

undertaken in towns where passing trade is important (e.g. Bertoua and Yokadouma, 

situated along the Central African Republic to Cameroon trade route). Likewise, 

Abong-Mbang was also included due to its rapid population increase (nearly 5-fold 

from 1967 to 1987) and continuing economic development. 

The survey sites selected were based on the following categories of rattan 

consumption identified by Shiembo (1986), Pokam-Wadja (1987), Defo (1997) and 

Defo (1999). 

· Small local markets with a high level of self-sufficiency 2 . These markets 

sometimes act as exchange places and supply the regional and national 

markets (Yokadouma, Abong-Mbang, Ebolowa and Bertoua); 

2 As related to range, or distance, of supply. 

245

· A group of medium-sized markets of regional importance with a moderate 

level of self-sufficiency. These markets often act as assembly points for the 

two large urban markets (Kribi, Limbe, Sangmelima, Mamfe, Kumba, Edea 

and Mbalmayo); 

· Large regional markets with a weak degree of self-sufficiency, having to rely 

on more distant supply areas (Bamenda and Bafoussam); 

· The two large urban markets of Douala (Marché des Fleurs) and Yaounde 

(Mvog-Mbi) with weak degrees of self-sufficiency and relying on distant 

supply areas. Combined, these markets comprise the majority of the rattan 

trade in Cameroon, and can be regarded as national in proportion. 

Figure 98. Map of southern Cameroon. Sites surveyed during this study are ringed. 

246

Figure 99. Scatterplot of size of market and distance of supply 

Range (km) 

250 

200 

150 

100 

50 

0 

LOCAL 

0 

1000000 

REGIONAL 

MEDIUM-SIZED 

2000000 

3000000 

4000000 

247 

5000000 

NATIONAL 

6000000 

Mean monthly value (CFA) 

7000000 

7.4.2 Sampling methodology 

Information for this study was gathered using a standard questionnaire modified from 

techniques developed through the implementation of a number of well-known general 

marketing studies (Padoch, 1987; Falconer, 1994; Martin, 1995; Alexiades 1996; 

Ndoye et al., 1999). The questionnaire was developed and modified to enable the 

gathering of both qualitative and quantitative information as a means of assessing the 

socio-economic nature of the rattan industry. The methodology was successfully 

tested in a pilot study of the rattan industry in the city of Bata, Equatorial Guinea 

(Sunderland, 1998). In total, 174 artisans in 15 urban markets 3 were interviewed over 

a two-month period from July to September 1998, representing a mean sampling 

percentage of 81%. 

3 There is a strong correlation between population and the number of artisan units per urban market 

(Pearson correlation coefficient r = 0.96, P = 0.01) and there is, on average, one processing unit for 

each 6,150 head of population.

Table 24. Study sites and sampling 

Population 4 Registered 5 

Number % (percentage) 

number of enumerated for sampling 

artisans this study 

Douala 809,852 96 24 25% 

Yaounde 649,252 124 31 25% 

Bafoussam 112,681 20 20 100% 

Bamenda 110,142 25 22 88% 

Kumba 70,112 16 13 81% 

Edea 50,609 11 6 55% 

Limbe 44,561 8 7 88% 

Bertoua 43,402 8 8 100% 

Mbalmayo 35,390 13 13 100% 

Ebolowa 34,771 2 2 100% 

Sangmelima 23,261 8 8 100% 

Kribi 21,507 7 7 100% 

Mamfe 13,844 8 7 88% 

Abong-Mbang 12,565 3 3 100% 

Yokadouma 11,235 2 2 100% 

353 174 81% 

7.5 THE RATTAN SECTOR IN CAMEROON 

7.5.1 Rattan harvesting and supply to the urban markets 

7.5.1.1 The resource base 

Although eighteen species of rattan occur in Cameroon, only three form the basis of 

the trade; the large diameter canes, Laccosperma secundiflorum and L. robustum and 

the juvenile stems of Eremospatha macrocarpa. As discussed in Chapter 6 and 

Appendix 1, although there is considerable spontaneous use for most species, only a 

very few species possess the qualities suitable for the commercial production of 

processed products. 

7.5.1.2 Customary laws and State legislation 

Without exception, rattan artisan workshops rely on a regular supply of unprocessed 

cane from the forest. Throughout its range, rattan is considered an open-access 

resource and, as such, is generally not subject to customary laws relating to land and 

resource tenure. This is also reflected in the formal State forestry sector; although, in 

theory, a permis d’exploitation is required for the commercial harvest of non-timber 

4 Accurate figures for population are only available from the 1989 census (GoC, 1989); the populations 

of Douala and Yaounde in particular are thought now to be >1,000,000 and 800,000 respectively 

(Horta, 1991). 

248

forest products (see Box 3) these are rarely, if ever, issued for the harvest of rattan. 

This shortfall provides a convenient mandate for forestry officials to apply an informal 

taxation system when unprocessed rattan is transported. Hence, the harvest of rattan, 

in common with many other forest products, is unregulated and uncontrolled, and the 

benefits are felt in the informal forestry sector, rather than the formal sector. The 

social conditions under which rattan is harvested are discussed in detail in Chapter 8. 

Box 3. Permis d’exploitation 

In Cameroon, the large scale exploitation of non-timber forestry products is subject to the obtention of a 

permis d’exploitation. This permit determines the quantities to be exploited or collected within a 

specified geographic area. The volume or amount of material allowed to be exploited depends on the 

desired material (e.g. fruits, bark, leaves etc.). This quota is set by the Department of Forestry, although 

base line and monitoring data for estimating potential sustainable yield is woefully incomplete for most 

taxa. The length of the exploitation permit would not usually exceed one year (National Forestry Law 

no. 94/01; article 56; October 1994), except by special arrangement. For example, Plantecam formally 

possessed permits for Prunus africana exploitation issued for a period of up to three years duration 

(Cunningham and Mbenkum, 1993). Exploitation permits also apply to special products, which could 

include medicinal species or those of particular interest (Sunderland et al., 1999). Even if special 

products are found on lands belonging to private individuals, they remain the property of the State, except 

where the said products have been acquired by the individual concerned. 

7.5.1.3 Production to consumption 

The supply of cane to urban markets is predominantly undertaken by local villagers 

and farmers. Although artisans also harvest rattan themselves, this is more commonly 

the case only where the urban market is in close proximity to a significant supply of 

rattan (such as Yokadouma and Abong-Mbang). The “production to consumption” 

(Belcher, 1999) of rattan harvest and supply in Cameroon is presented below. 

5 Registered refers to those enterprises actually recognised in urban council records. It is very difficult 

for most enterprises to operate without formally registering with the local council. Figures were 

obtained through reference to local council registers. 

249

Figure 100. Production to consumption rattan flow in Cameroon 

Production → → → → → → → Consumption 

Forest 

Rural 

harvester 

Urban 

harvester 

Urban 

artisan / 

harvester 

Trader (point 

of sale) 

Rural milieu → → → → → Urban markets 

Unprocessed rattan → → → → Transformation → Finished 

products 

Table 25. Percentage of artisans who harvest their own cane 

Town / city Artisans / Artisans Source of cane if not harvested by artisans 

harvesters only 

Mbalmayo 92% 8% Bought directly from villagers / traders 

Bertoua 62.5% 37% Bought directly from villagers / traders 

Yokadouma 100% 0 Collected directly from forest 

Abong-Mbang 100% 0 Collected directly from forest 

Limbe 0 100% Bought directly from villagers / traders 

Kumba -0 100% Bought directly from villagers / traders 

Mamfe 50% 50% Bought directly from villagers / traders 

Tiko - 100% Bought directly from villagers / traders 

Sangmelima 37% 63% Bought directly from villagers / traders 

Ebolowa 50% 50% Bought directly from villagers / traders 

Kribi 29% 71% Bought directly from villagers / traders 

Edea 34% 66% Bought directly from villagers / traders 

Yaounde 0 100% Bought from formal urban market 

Douala 0 100% Bought from formal urban market 

Bafoussam 0 100% Bought from suppliers, buying from Douala 

Bamenda 0 100% Bought from suppliers, buying from Douala 

35% 65% 

7.5.1.4 Range 

The range, or distance, that unprocessed rattan is transported to each urban market 

varies significantly across the forested zone. To some extent, this is reflected in the 

unit price of the cane. 

250 

Formal markets 

(re-sale) 

Transformation 

(urban artisans) 

Sale

Figure 101. Mean range (or distance) of rattan supplies 

Mean range 

(km) 

250 

200 

150 

100 

50 

0 

8.4 8.5 10 13.4 15.5 18 20 

Figure 102. Urban price paid per unit 

Mean unit price (CFA) 

5,000 

4,500 

4,000 

3,500 

3,000 

2,500 

2,000 

1,500 

1,000 

500 

0 

251 

25 28.8 

120 

110 115 

100 

72 

180 

Mbalmayo 

Ebolowa 

Yokadouma 

Sangmelima 

Mamfe 

Abong-Mbang 

Bertoua 

Kribi 

Edea 

Kumba 

Bamenda 

Douala 

Limbe 

Mutengene 

Yaounde 

Bafoussam 

Sangmelima 

Kumba 

Mutengene 

Mbalmayo 

Mamfe 

large dia. Cane small dia. Cane 

Ebolowa 

Kribi 

Yokadouma 

Abong-Mbang 

Bertoua 

Limbe 

Edea 

Yaounde 

Bafoussam 

Douala 

7.5.1.5 Frequency of supply and purchase 

There is a regular supply of cane from the harvest sites to the urban markets. In the 

main, artisans are not able to store large quantities of cane for long periods as it 

deteriorates and becomes unworkable. In addition, most enterprises lack the capital to 

buy bulk quantities of cane even if they could store the unprocessed cane prior to 

Bamenda 

220

transformation. Hence, more commonly, small quantities of cane are purchased on a 

regular basis. For the same reasons of lack of storage and capital, some artisans will 

not buy unprocessed rattan on a regular basis at all without actually having 

commissioned work. These artisans will only purchase raw material “on command”, 

i.e. when rattan products are commissioned. 

Figure 103. Frequency of wholesale rattan purchase by artisans (n = 174) 

Once a month 

20% 

On command 

8% 

Twice a month 

23% 

252 

Twice a week 

6% 

Thrice a month 

13% 

Once a week 

30% 

7.5.2 Rattan artisan enterprises 


Almost without exception, the majority of commercial rattan enterprises currently 

operating in Cameroon are privately owned. These enterprises produce a wide range 

of rattan products, mainly concentrating on furniture and other household items such 

as woven lamp shades and flower baskets. The production of temporary market 

baskets for “buyam sellums” is an important component of the rattan artisan trade. 

7.5.2.2 Number of workers and sources of labour 

In general, artisanal enterprises are small operations and invariably consist of a single 

owner / artisan. However, some craftsmen are supported by a number of apprentices. 

These apprentices are trained by the owner/artisan, and they are primarily responsible 

for the processing of the cane prior to transformation 6 . In general, apprentices are not 

6 In the African context, processing of raw rattan essentially entails the removal of the epidermis (skin) 

from the stem and the drying of the raw cane prior to its use. Immature stems, or the very apex of 

mature stems, where the leaf sheath is also present are not used, and are often left or discarded at the

aid a wage. In fact, quite the opposite is the case and, ordinarily, the families of 

apprentices pay the artisan for the training provided. Other sources of labour include 

that provided by family members, particularly from children, who contribute to the 

more mundane tasks. 

Figure 104. Summary of the types of products produced by Cameroonian artisans (n = 174) 

Tie-tie for 

thatching 

1% 

Furniture & 

baskets 

8% 

Furniture only 

62% 

Figure 105. Number of workers per enterprise 

No. of enterprises (n = 174 ) 

100 

90 

80 

70 

60 

50 

40 

30 

20 

10 

0 

Lamp shades only 

1% 

Baskets only 

9% 

253 

Baskets & flower 

jars 

5% 

Farm & market 

baskets 

14% 

1 worker 2 workers 3 workers 4 workers 5+ workers 

time of harvest. The processing of raw cane is undertaken manually, with the stems being scraped with 

kitchen knives to remove the skin followed by drying, usually undertaken in the open air. This 

rudimentary means of processing is extremely labour intensive. For a more detailed discussion of the 

processing of rattans in SE Asia and potential application to the African context, see Sunderland and 

Nkefor (1999).

7.5.2.3 Types of enterprise 

Rattan enterprises in Cameroon, as they are in much of West and Central Africa, are 

housed in conspicuously modest surroundings. Many of these enterprises are 

essentially “open-air” workshops, although the working area may be covered with a 

simple roof structure of palm fronds or zinc to protect the workers from the 

vicissitudes of the climate. However, there is generally little protection for the 

unprocessed rattan, or the finished products. Because of this, many open-air 

workshops do not operate during the rainy season. Enclosed, permanent enterprises 

are also common. These may be cement-block houses, which also serve as residences 

or timber structures (with both walls and roofs) utilised solely for the use of the 

enterprise. 

The location of the workshop is crucial to eventual sale of finished products and 

many, if not all, are situated along roads, where finished goods are displayed. In the 

majority of the urban areas, the rattan enterprises are generally grouped together in 

one area of the town. Where rattan is sold in a central market, such as Mvog-Mbi in 

Yaoundé, or the Marché des Fleurs in Douala, this area is usually exclusive to the 

formal central markets where other forest products are traded. This is also the case for 

the sale of finished products; rattan is traded a distinct, and separate, commodity from 

other forest products 7 . 

Figure 106. Types of rattan workshop in Cameroon 

No. of enterprises (n = 174) 

90 

80 

70 

60 

50 

40 

30 

20 

10 

0 

Enclosed permanent Open permanent Enclosed temporary Open temporary 

7 Although a relationship between rattan harvesting hunting for bushmeat has been highlighed by 

Trefon and Defo (1998) and Defo (1999). 

254

7.5.3 Socio-economic profile of the urban artisans 

7.5.3.1 Age range 

In general, rattan enterprises are operated by relatively young to middle-aged men, the 

majority of whom are married (70% +). Many of the older artisans interviewed 

suggested that the work was very much for younger, fitter men and that as they have 

become older, their work outputs, and subsequent profits, have decreased. Unlike 

many rural artisans for whom rattan transformation is often a secondary activity to 

farming, urban-based artisans are almost always engaged on a full-time basis. 

Figure 107. Age range of the primary owner / worker 

no. of respondents (n = 174 ) 

70 

60 

50 

40 

30 

20 

10 

0 

60 

Age range 

7.5.3.2 Gender 

Although Ndoye (1994) reports that women are sometimes involved in harvesting 

activities, this is relatively rare (Defo, 1997; 1999; Sunderland 1998; 1999a; 1999b). 

Correspondingly, the processing and transformation of rattan is predominantly a male 

activity, however, some women are involved in the weaving of baskets, notably in 

Bamenda. 

7.5.3.3 Ethnicity 

The ethnic background of the urban artisans is extremely variable and in all of the 

urban markets studied, there is no domination by any single ethnic group in rattan 

processing and transformation. This is particularly the case in the larger towns and 

cities, which have a tendency to be more cosmopolitan. However, somewhat 

unsurprisingly, a pattern does emerge that a greater proportion of artisans within an 

255

urban market, originate from there. This is particularly the case in smaller urban 

markets. 

8.5.3.4 Educational background 

As would be expected from a semi-skilled manual labour force such as rattan artisans, 

the educational level of the majority of the artisans in Cameroon is relatively poor. 

However, there are exceptions to this and some artisans are educated to secondary or 

even to first-degree level. Interestingly, artisans who state they have originated from a 

“technical” background often includes those who have undertaken rattan artisan 

training in the context of rehabilitation, such as that provided by the prison service. 

Rattan artisan training is one of the major rehabilitation programmes of the Cameroon 

prison service as it is both cost-effective 8 and enables the prisoner to enter gainful 

employment upon release. The fact that many of the artisans in Cameroon are exprisoners 

is testament to the success of this rehabilitation. For obvious reasons, many 

artisans would not admit having undertaken such a training programme, however, and 

it is not possible to ascertain the proportions of those having gone through the 

rehabilitation process. 

Figure 108. Educational level of rattan artisans 

technical (incomplete) 

1% 

secondary (complete) 

6% 

secondary 

(incomplete) 

27% 

technical (complete) 

6% 

university 

3% 

256 

no formal education 

2% 

primary (incomplete) 

8% 

primary (complete) 

47% 

8 

The prisoners harvest, transport and transform the rattan themselves (under close supervision, of 

course).

7.5.3.5 Previous occupations 

A large number of the artisans sampled have been previously employed, commonly in 

unskilled and skilled labour occupations, although some have also come from a 

professional background. In addition, a substantial proportion of artisans have come 

directly from full-time education. 

Figure 109. Previous occupation of rattan artisans 

Skilled manual 

21% 

Unskilled manual 

18% 

Professional 

11% 

Agriculture 

2% 

Military 

1% 

257 

Commerce 

11% 

Driver 

7% 

Straight from 

education 

29% 

In terms of longevity, the majority of rattan artisans have been active in the industry 

for less than 10 years. 

Figure 110. How many years spent in the trade? 

16-20 years 

16% 

21-25 years 

6% 

11-15 years 

16% 

26-30 years 

6% 

>30 years 

5% 0-5 years 

27% 

6-10 years 

24%

7.5.4 The scale of the trade 

7.5.4.1 Amounts and values 

The amount and value of rattan being transformed in the forest zone of Cameroon is 

significant. In this respect, the rattan sector in Cameroon is similar in scale to that of 

other countries in West and Central Africa (see Chapter 8). 

Table 26. Amount (in metres) and value (in CFA) of unprocessed rattan consumed each month 

per urban market. 

Amount of cane used / month (m) Value of cane used / month (CFA) 

Large dia. cane Small dia. cane Large dia. cane Small dia. cane 

Douala 107,700 115,500 4,138,740 2,230,400 

Yaounde 92,660 119,060 2,342,740 2,833,340 

Bafoussam 10,800 7,380 374,500 187,250 

Bamenda 5,700 20,990 250,710 450,150 

Kumba 5,900 10,785 50,480 85,950 

Edea 8,820 4,805 126,050 93,000 

Limbe 3,315 2,510 56,000 24,500 

Bertoua 4,880 1,040 97,600 10,400 

Mbalmayo 9,290 2,550 124,000 25,600 

Ebolowa 2,000 1,350 33,300 9,000 

Sangmelima 7,275 5,480 84,085 76,975 

Kribi 4,095 9,250 71,500 62,500 

Mamfe 4,390 6,585 58,800 98,200 

Abong-Mbang 1,760 300 35,200 29,800 

Yokadouma 400 140 8,000 14,000 

Totals = 285,805 307,725 7,851,365 6,231,070 

Based on these figures, the annual consumption of rattan in the forest zone of 

Cameroon is estimated to be 3,225,660m of large diameter cane with a market value 

of US $157,020, and 3,692,700m of small diameter cane with a market value of US 

$124,620. Thus, the trade in unprocessed rattan alone is an estimated US $281,640. As 

this figure does not capture the value of finished products, nor the substantial 

household and rural utilisation of rattan (and therein lies scope for future study) this 

trade, as postulated, is significant. 

7.5.4.2 Profitability 

Despite the lamentations of many of the artisans interviewed, incomes and profit 

margins from the transformation and sale of rattan products are relatively high. Given 

that the average monthly income for an employed semi-skilled or skilled labourer in 

most regions of Cameroon is 30,000 CFA, most artisans are comparatively well off. 

However, there are considerable fluctuations in monthly profits, particularly during 

258

the rainy season when transport difficulties increase the costs of rattan entering the 

market, if indeed the rattan can be transported at all. 

Figure 111. Mean monthly profits of rattan artisan per urban market 

Mean monthly profit (CFA) 

90000 

80000 

70000 

60000 

50000 

40000 

30000 

20000 

10000 

0 

13000 16000 

Yokadouma 

Mutengene 

31045 

25250 25340 

33440 

41750 

42150 

39000 40000 

Bafoussam 

Mamfe 

Bamenda 

Mbalmayo 

Limbe 

Abong-Mbang 

259 

Kumba 

Kribi 

Ebolowa 

50000 

Edea 

63100 

54400 

50600 

7.5.5 The nature of the trade 

7.5.5.1 Decline or growth? 

Although some studies have suggested that the trade in rattan in Cameroon is growing 

(Defo, 1997), very little evidence is provided in this regard. During the interview 

process, the artisans sampled were requested to provide information regarding the 

growth or decline in their own consumption of rattan. Regarding this growth or 

decline, if they responded in the positive or the negative they were also asked to 

provide reasons why, in their experience, this is the case. 

Figure 112. Is there more cane being used this year (1998) than in the previous five years? (Based 

on numbers of responses n = 174) 

The same 

10% More 

34% 

Less 

56% 

Bertoua 

Sangmelima 

Yaounde 

81550 

Douala

In order to determine the general trend for each urban market, and for the sector as a 

whole, the responses outlined above were ranked as follows: 

r more cane used now than previously +1 

r less cane used now than previously -1 

r unchanged 0 

These responses were then added together for each urban market to detect qualitative 

trends in the amounts of unprocessed rattan being consumed. 

Figure 113. General trend of rattan sector per urban market (positive = growth of sector; 

negative = decline of sector; 0 = no change) 

6 

4 

2 

0 

-2 

-4 

-6 

-8 

-10 

-12 

Bafoussam 

Bamenda 

Kumba 

Douala 

Limbe 

Sangmelima 

Yokadouma 

Abong-Mbang 

The cumulative figure, when all of the individual urban rankings are added together, (- 

24) provides strong evidence that the rattan sector in Cameroon is currently in some 

decline. 

260 

Kribi 

Edea 

Mamfe 

Yaounde 

Ebolowa 

Bertoua 

Mbalmayo

Figure 114. Cited reasons for decline in rattan sector in certain urban markets. (Based on 

numbers of responses n = 125). 

Lesser share of 

market (more 

competition) 

52% 

261 

Personal 

circumstances 

17% 

Scarcity of cane 

31% 

Figure 115. Cited reasons given for growth of rattan sector in certain urban markets (figures 

given are per response) 

Increased 

demand for 

products 

60% 

Increased 

labour 

availability 

15% 

Technical 

advances 

25%

Table 27. Major constraints to the development of the rattan sector in Cameroon. (Based on 

number of responses). 

Rank Constraint No. of responses 

(n = 363) 

1 Scarcity of raw material (rattan) 125 

2 Competition between artisans (lack of custom) 52 

3 Lack of technology for processing & transformation (labour intensive 44 

4 Lack of capital 35 

5 Open workshops and storage (adversely affected by weather) 33 

6 High costs of inputs (nails, plywood etc) 28 

7 High taxation (formal and informal) 19 

8 Rattan considered “poor man’s furniture” 8 

9 Transport of finished products to market 6 

9 Poor quality of cane 6 

11 Dangers of cutting cane in forest 3 

12 Lack of artisanal union 2 

12 Poor state of National Economy 2 

Table 28. Recommendations of artisans for the stimulation of the rattan sector in Cameroon. 

Rank Recommendation No. of responses 

(n = 279) 

1 Form artisan unions (price fixing) 56 

2 Greater access to raw material 55 

3 Provision of machinery for processing and transformation 38 

4 Access to credit 27 

5 Exhibition / promotion of rattan products 21 

6 Central enclosed workshops 19 

7 Training in improved artisan techniques for better quality products 18 

8 Increased markets 16 

9 State support in sector 12 

10 Develop export markets 10 

11 Lower prices for inputs (subsidies) 9 

12 Cultivation of rattan 4 

12 End of harassment by police and forestry officials 4 


7.6.1 “La crise” and the increased reliance on forest products 

It is clear from the surveys results presented above that a number of artisans originate 

either from a high educational level and / or from previously successful professional 

trades. 

The main reason for this is due to the economic crisis that hit Cameroon in the late 

1980’s and early 1990’s. The general world recession of the 1980’s that resulted in 

falling prices of oil and agricultural produce (which accounts for 90% of GDP) 

affected Cameroon deeply. This was compounded by a large and top heavy civil 

262

service. Unemployment increased from 12% to 25% during the late 1980’s, civil 

service salaries were reduced by 70% in November of 1993 followed by a 50% 

devaluated of the CFA franc in January 1994. The immediate result of this economic 

downturn has been an increased reliance on subsistence farming by all sections of the 

population, as well as the uncontrolled exploitation of medicinal plants, bushmeat, and 

other forest products. Although the economy has picked up significantly since the 

mid- to late 1990’s, the reliance on forest products remains strong. 

For this reason, many of those leaving education and unable to find employment, or 

are forced to leave education to support the family, or those simply finding themselves 

unemployed, have turned to the forest sector for employment. Aside from the 

relatively high educational and professional levels of some rattan artisans, further 

evidence of this is illustrated by our survey results. These also show many of those 

involved in rattan transformation are relatively recent entrants into the sector < 10 

years and are of an increasingly young age. 

Aside from the economic crisis, although perhaps related to it, the fact that many 

rehabilitated prisoners enter the rattan artisan trade directly from serving their prison 

terms also accounts for the wide variation of the social background of rattan artisans. 

Again, this factor certainly accounts for some of the higher education and professional 

levels of some of the artisans operating in the sector, although due to the sensitive 

nature of this issue, this is difficult to assess. 

Hence, due to this economic crisis, the growth of the rattan sector in the late 1980’s 

first reported by Shiembo (1986) and Pokam-Wadja (1987) led to the establishment of 

many artisan operations in Cameroon. This has had two major impacts: 

•r The increased number of artisans in a market that, despite also undergoing 

some growth for the same reasons 9 , has resulted in the saturation of artisans 

enterprises in some urban markets. This has led to increased competition to the 

extent that profit margins have been cut significantly; 

9 Increased rural and urban poverty meant that consumers had to purchase furniture they could afford 

and they could no longer afford expensive wood products, but began to rely on substitute products such 

263

• This increase in processing and transformation has accordingly resulted in 

increased harvesting of rattan from the wild. 

7.6.2 Sustainability 

There is no doubt that the current exploitation of rattan in Cameroon is unsustainable 

and it is clear that the present intensity of harvest is exceeding that of regeneration and 

growth. The reported scarcity of supplies for many urban markets (Shiembo, 1986; 

Pokam-Wadja, 1987; Defo, 1997; Defo, 1999; this study) and the fact that the harvest 

range is increasing steadily is strong evidence to support this. The over-exploitation of 

rattan has been facilitated by the provision of greater access to the forest through 

increased logging activities during the same period. 

7.6.3 Increased range = increased price? 

As the distance that rattan has to travel increases due to this scarcity, there is also a 

corresponding increase in the opportunity and transport costs of harvesting rattan. 

Whilst there is a correlation between the distance rattan travels from the harvest site to 

the area of transformation and the wholesale price of unprocessed rattan the 

correlation is not as strong as might be expected (Pearsons correlation coefficient, r = 

0.524, P < 0.01). The reasons for this are complex, but are, in the main, due to two 

main factors: 

•r The type of road along which the cane is transported (100km on a tar road is a 

much easier journey, and with more transport opportunities, that 10km on a 

very poor un-graded road); 

• The number of police checkpoints on a given stretch of motorable road. 

“Informal taxation” by members of the police, gendarmerie, military, and 

forest services is common when transporting rattan (and indeed, many other 

products). Transport routes, such as the Douala-Yaounde road, are 

characterised by a high concentration of checkpoints and result in increased 

costs of transportation of rattan, along with the inevitable delays this causes to 

the journey. 

as rattan. Hence, for many Cameroonians, the stigma has persisted that rattan products are a “poor 

man’s furniture”. 

264

Figure 116. Correlation between range (distance km) and costs of rattan (CFA) 

UNIT 

PRICE 

5000 

4000 

3000 

2000 

1000 

0 50 100 150 200 250 

DISTANCE 

7.6.4 Profitability and size of urban market 

Although, to some degree, profits are determined by the size of the urban market 

concerned, and there is a correlation to suggest as much (Pearson’s correlation 

coefficient r = 0.729, P < 0.01), cultural and socio-economic factors are also 

somewhat influential in this regard. For example, towns with little economic status 

where the rattan market is aimed primarily at the indigenous, and often poorer, 

proportion of the population, have smaller profit margins (Yokadouma and Mamfe). 

Towns with greater economic status, higher incomes and greater concentrations of expatriots 

10 have greater profit margins (Kribi and Limbe). Cosmopolitan cities such as 

Douala (the economic capital of Cameroon) and Yaoundé (a large proportion of the 

population of which is comprised of civil servants and their dependants) provide rattan 

10 In contrast to the indigenous market, rattan is a favoured product for ex-patriots working in 

Cameroon and hence the increased profit margins in towns with high concentrations of resident, or 

visiting (for tourist areas), ex-patriots. 

265

artisans with the highest incomes. Both these latter cities also have large 

concentrations of ex-patriots. 

7.6.5 An industry in decline? 

This study has concluded that the rattan sector in Cameroon is currently in some 

decline. However, it should be noted that this decline is currently only relative to the 

quantities of rattan being transformed and not currently in terms of a decline in the 

overall value of the sector. The reason for this is simple and unanimously agreed upon 

by the artisans interviewed for this study. At present, demand is such that local 

scarcity has forced gradual increases in the price of unprocessed cane which are 

currently being absorbed by the consumer. Whilst the quantities of finished products 

are decreasing, the corresponding increases in the price of raw material, which are 

then passed to the consumer, is not as yet manifesting in a significant reduction in 

income for most artisans. As such, most enterprises remain profitable. Obviously, this 

situation cannot continue ad infinitum and eventually, if present harvest rates 

continue, the price of raw material will exceed more than the “willingness to pay” of 

most consumers. 

However, some decrease in profitability has been identified by this survey. This is not 

ultimately due to scarcity of supplies, but to increased competition. Price wars 

between enterprises have recently been in operation, particularly in the large urban 

markets, and these have led to a small reduction in real incomes as enterprises struggle 

to maintain competitiveness. One positive action that most artisans identified that 

could have an effect on this is the establishment of a rattan union that would be 

responsible for setting a pricing framework for standard finished products. This would 

then avoid being under-cut by competitors. Whilst, in theory, this is a wonderful idea 

from the perspective of artisan enterprises, given the variation in the quality of 

finished products by most artisans, this might not be workable in practice. 

Changes in personal circumstances also account for the decline in rattan enterprise. 

For example, some artisans stated that as they get older, they begin to rely on other 

sources of income. Other artisans are somewhat peripatetic in their activities and 

revert to part- or full-time farming, particularly as commodities such as cocoa and 

coffee have increased in value in recent years. Without question, despite the fiscal 

266

enefits rattan brings, those artisans who are also involved in harvesting unanimously 

state that the unpleasant and risky nature of the harvesting process make alternative 

income generating activities far more attractive. Correspondingly, some artisans 

suggest that this is one of the main reasons for local scarcity. In this regard, they state 

that village-based harvesters, who were formerly supplying urban markets, are 

reverting to farming and other occupations for their livelihoods and are less inclined to 

continue with these difficult harvesting activities, particularly as the harvest range 

increases. Hence, rattan harvesting may be described as a livelihood “safety-net” 

when other occupations are not economic. This situation has been also described by a 

number of previous studies (Ndoye, 1994; Defo, 1997; Defo, 1999; Sunderland, 

1999a; 1999b). 

In contrast to the general decline in the rattan sector, some artisans suggest that their 

own circumstances have improved in recent years and their share of the market in 

finished products is increasing. Although these are in the minority, it certainly seems 

that some artisanal enterprises are indeed expanding, particularly in the employment 

of additional employees. In general, these enterprises are based in Douala and 

Yaounde where they have permanent workshops along with access to innovative 

designs and a large ex-patriot market. The increased labour availability can also be 

considered a direct result of the economic crisis; with more unemployment, available 

labour has increased dramatically. 

7.6.6 Rattan unions 

Whilst the use of a formal artisan union to establish a pricing framework might be 

somewhat counter-productive, the sector could certainly benefit from some level of 

organisation and formalisation. The benefits that could be brought about by creating a 

formal union could possibly include the provision of credit facilities, increased 

efficiency through the adoption of appropriate technologies, opportunities for training 

and product development and the promotion of rattan products. In this regard, 

government intervention in the sector might bring about such benefits, although an 

NGO or development agency might also be an appropriate catalyst for such a network 

of organisations. 

267


As this survey has shown, the trade in unprocessed rattan alone is extremely valuable 

and contributes significantly to the livelihoods of rural harvesters as well as artisans 

that rely on unprocessed rattan. However, very little of this income benefits the 

production side of the sector and for this reason, very little effort is being made to 

generate sustainable management regimes for the exploitation of rattan. This is 

additionally hindered by a lack of adequate resource tenure for a product that has 

traditionally regarded as an open-access resource. 

Although there is considerable “informal” taxation of the rattan trade, as well as 

limited (formal) revenue collection, in the form of council dues, the formal forestry 

sector receives very little if any revenue from this extremely profitable industry. Part 

of the problem, as discussed, is the fact that revenues from forest products are traded 

in invisible markets. However, this need not be the case, and, it would not be 

prohibitively complicated to establish a system of harvesting licenses and quotas 

based on known and actual sustainable yields (once this baseline information becomes 

available). It is often argued that the monitoring of such a system is a particular 

problem given the limited resources of the forestry services. However, it could be 

argued that if forest officers are actively gathering informal taxes, how much more 

regulation would it take to collect taxes on an official basis? 

Perhaps if some of the income generated from a high value NTFP such as rattan is 

formally realised by the forestry administration, this might provide the necessary 

means of supporting such initiatives. Revenues generated by such high value NTFPs, 

could be realised through collecting fees for permits and direct product taxation, based 

on volume (equivalent to “stumpage” for timber). 

The realisation of the true value of such resources is critical in the pursuit of a holistic 

conservation management approach for a wide range of forest products rather than for 

timber resources alone. In this respect, moves towards formal Community Forestry 

legislation throughout much of West and Central Africa provide the legislative 

framework for such an ideal. However, in terms of ecological, social and institutional 

criteria that need to be satisfied for this to happen, it seems that we still are a long way 

268

from enabling the sustainable exploitation of NTFPs to contribute both to 

conservation objectives and to livelihoods. 

269

Figure 117. Open-air permanent rattan enterprise, Bata, Equatorial Guinea 

Figure 118. Enclosed permanent rattan enterprise, Bata, Equatorial Guinea 

270

Figure 119. Woven basket products made from Eremospatha macrocarpa on sale on the 

Mbalmayo to Yaounde road 

Figure 120. Finished rattan furniture for sale in Bata, Equatorial Guinea 

271

CHAPTER EIGHT 

A BRIEF OVERVIEW OF THE RATTAN TRADE IN AFRICA 

8.1 THE INTERNATIONAL ECONOMIC CONTEXT 

The international trade in rattan dates from the mid-19 th century (Corner, 1966) and 

this trade is currently worth an estimated $US 6.5 billion a year (ITTO, 1997). A 

conservative estimate of the domestic markets of SE Asia alone by Manokaran (1990) 

suggests a net worth of $US 2.5 billion. This latter market includes the value of goods 

in urban markets and rural trade, as well as the value of the rural usage of the material 

and products. Dransfield and Manokaran (1994) estimate that 0.7 billion of the 

world’s population use, or are involved in, the trade of rattan and rattan products. 

By the 1970s, Indonesia had become the supplier of about 90% of the world’s cane, 

with the majority of this going to Singapore for processing and conversion (from 

which Singapore earned more than US$21 million / annum). In 1977, Hong Kong 

imported some $US26 million of rattan and rattans products which, after conversion, 

was worth over US$68 million in export value. By comparison, Indonesia’s share of 

the trade, mainly of unprocessed canes, was a mere US$15 million (Manokaran, 

1990). 

In the last twenty years the international trade in rattan and rattan products has 

undergone rapid expansion. By the late 1980’s, the combined value of exports of 

Indonesia, Thailand, Malaysia and the Philippines alone had risen to an annual figure 

of almost US$400 million. The net revenues derived from the sale of rattan goods by 

Taiwan and Hong Kong, where raw and partially finished products were imported and 

then processed, together totalled around US$200 million / annum by the late 1980s 

(Manokaran, 1990). 

During this same period, these same four countries banned the export of rattan, except 

as finished products. These bans were imposed to stimulate the development of rattanbased 

industries in each country to ensure that the value of the raw product was 

increased, and (theoretically) to protect the wild resource. Recently, however, the 

economic recession that has hit many countries in SE Asia has meant that countries 

273

such as Indonesia have lifted the ban on the export of raw cane and are currently 

flooding the market with relatively cheap supplies of cane. This is negatively 

impacting the cultivation industry of Malaysia in particular (Loke et al., 1996; 

Sunderland and Nkefor, 1999). 

8.2 THE AFRICAN RATTAN TRADE 

The relatively recent restrictions in the trade of raw cane by some of the larger supply 

countries outlined above has encouraged some rattan dealers and gross users to 

investigate non-traditional sources of rattans. Other sources of supply have 

concentrated predominantly on Indo-China, Papua New Guinea and more recently, 

Africa. Some raw cane has recently been exported from Ghana and Nigeria to SE Asia 

and there is a flourishing export trade of finished rattan products from Nigeria to 

Korea (Morakinyo, 1995a). In addition, trade within and between countries is known 

to be growing significantly across West and Central Africa (Falconer, 1994; 

Morakinyo, 1995a; 1995b; Sunderland 1999a; 1999b). 

During the colonial period, there also existed a significant trade in African rattans. In 

particular, Cameroon and Gabon supplied France and its colonies (Hédin, 1929), and 

Ghana (formerly the Gold Coast) supplied a significant proportion of the large UK 

market during the inter-war period (Anon, 1934). The export industry was not 

restricted to raw cane and in 1928 alone over 250,000 FF worth of finished cane 

furniture was exported from Cameroon to Senegal for the expatriate community there 

(Hédin, 1929). More recently, an initiative promoted by UNIDO in Senegal was 

exploiting wild cane for a large-scale production and export (Douglas, 1974), although 

this enterprise folded not long after its establishment due to problems securing a 

regular supply of raw material. 

Table 29. Raw rattan cane exports from Douala and Kribi to France 1926 to 1928 (Hédin, 1929) 

Year Tonnes Exported Value (FF) 

1926 100 250,000 

1927 58 137,000 

1928 (Douala) 32 80,000 

1928 (Kribi) 34 85,000 

274

In recent years, rattan as been identified as one of the most important non-timber 

forest products of West and Central Africa (Wilkie, 1999). To reflect this importance, 

a number of studies have attempted to quantify the trade in rattan and rattan products 

and to assess the role of rattan in contributing to rural and urban livelihoods. In order 

to determine the potential that rattans might have in this regard, it is also essential to 

understand the nature of the trade, as well as understand the socio-economic 

characteristics of the industry itself (see Chapter 7). 

8.3 THE RESOURCE BASE 

Although numerous studies have concentrated on evaluating the importance of rattan 

in these markets, very few have attempted to adequately define the resource base. As 

discussed in Chapter 6 and Appendix 1, the utilisation of rattan to supply the thriving 

cottage industry is limited to a few species. Table 30 presents a breakdown of the 

major commercial species of rattan per region. 

Table 30. Commercially important rattan species by region 

Region Commercially utilised species 

West Africa (Senegal, Côte d’Ivoire, Ghana, 

Benin, W. Nigeria) 

West/Central Africa (E. Nigeria, Cameroon, 

Congo, Gabon, E. Guinea) 

Central Africa (DR Congo, CAR) 

Southern/East Africa (Zambia, Uganda, 

Kenya, Tanzania) 

* Indicates primary commercial species 

8.4 THE NATURE OF THE TRADE 

*Laccosperma secundiflorum 

*Eremospatha macrocarpa 

Eremospatha hookeri 

Calamus deërratus 

Laccosperma secundiflorum 

*Laccosperma robustum 

*Eremospatha macrocarpa 

*Laccosperma robustum 

*Eremospatha haullevilleana 


*Calamus deërratus 

*Eremospatha haullevilleana 

Large quantities of raw cane enter the urban centres of West and Central Africa each 

day (Morakinyo, 1994; Ndoye, 1994; Falconer, 1994; Townson, 1995; Trefon and 

Defo, 1998; Defo, 1997; Sunderland, 1998; Defo, 1999; Sunderland 1999a; 1999b; 

Kialo, 1999; Minga, in press; Holbech, 2000; Oteng-Amoako and Obiri-Darko, in 

275

press; this study) and the conditions and circumstances under which rattan is harvested 

and transported are similar throughout its range. 

The harvesting of rattan is currently undertaken solely from wild populations. It is an 

unpleasant and often dangerous occupation with dead branches being dislodged as 

well as ants (see Chapter 4) and wasps being disturbed. Raw cane is cut, and bundled, 

and then head-portered out of the forest to the roadside. 

The majority of the harvesting that is undertaken for commercial trading is undertaken 

by individuals, usually farmers or hunters or other rural people primarily involved in 

other occupations. Rattan harvesting provides many these individuals with extra 

revenue, particularly in times of need such as for medical expenses or the payment of 

annual school fees (Trefon and Defo, 1998; Sunderland, 1998). Many cash-crop 

farmers also harvest rattan to obtain extra capital to purchase chemicals, planting 

stock and other necessary items for their primary occupation (ibid.). However, despite 

the capital returns, given the unpleasant and difficult nature of rattan harvesting, most 

would concentrate on their primary occupations given the opportunity. 

In general, rattan harvesters tend to work in the same forest area, and return each time 

they need to cut cane. If the harvester is not an indigene of the area, the chief of the 

local village is paid a small retainer for providing access to the forest. The harvesters 

usually prefer to collect as close to a motorable road as possible to avoid headportering 

the bundled canes too far. However, local scarcity near many urban centres 

now forces many harvesters further into the forest (Sunderland, 1998; Defo, 1999; 

Profizi, 1999). The added porterage resulting from this increased range is slowly 

generating an increase in the raw cane prices, which is being felt at the market level 

(see Chapter 7). 

Village-based harvesters transport the harvested rattan to the urban markets 

themselves, or they may sell at the village to a local trader who then transports the 

cane for sale to urban artisans. Some urban-based artisans harvest rattan themselves, 

although this is often only the case where there is close proximity to the wild resource. 

Falconer (1994), and Oteng-Amoako and Obiri-Darko (in press) provide a good 

276

overview of the production to consumption system of rattan in Ghana, as does Defo 

(1999) for selected sites in Cameroon (see also Chapter 7). 

Although many of the commercial species of rattan respond well to selective logging 

activities, logging has also resulted in increased rattan exploitation. The development 

of a wide network of logging roads throughout many forest areas in West and Central 

Africa has enabled greater access to otherwise inaccessible areas of forest. Indeed, the 

logging trucks themselves are often known to be responsible for the transport of 

recently harvested rattan (Defo, 1997; Sunderland, 1998). 

Indigenous management systems for the rattan resource in Africa are unknown, and, 

throughout its range, rattan is considered an “open-access” resource; there are very 

few, if any customary laws regulating the harvest of rattan from the wild. This is also 

mirrored in the National legislation for most countries. Those States that require the 

exploitation of forest products to be governed by the issue of licenses and permits, 

often do not adequately monitor the exploitation of these resource, nor receive the full 

forestry taxes related to that exploitation. In general though, many national forestry 

codes still do not include the exploitation of non-timber forest product in their 

regulations and the over-harvesting of many commercially important products, 

including rattan, continues unabated and uncontrolled. 

8.5 SUSTAINABILITY ISSUES 

It is reported that the demand for rattan is increasing and much greater amount of cane 

is being processed in many areas of Africa today than was being worked five or ten 

years ago (Morakinyo, 1994; Ndoye, 1994; Falconer, 1994; Townson, 1995; Trefon 

and Defo, 1998; Defo, 1997; Sunderland, 1998; Defo, 1999; Sunderland 1999a; 

1999b; Kialo, 1999; Minga, in press; Holbech, 2000; Oteng-Amoako and Obiri- 

Darko, in press). This has led to a significant decline in wild stocks and has resulted in 

considerable local scarcity. This scarcity and the associated irregular supply of 

unprocessed rattan have been identified as one of the major constraints to the 

continued development of the industry. 

277

Table 31. Findings and recommendations of selected socio-economic and market-related surveys 

of the rattan sector in Africa 

Reference Methodology Constraints to rattan 

development as identified by 

Shiembo, 1986 Interviews with 

traders and artisans in 

Cameroon (n = 768) 

Falconer, 1994 Interviews with urban 

artisan operations (n = 

39); and with rural 

harvesters & traders (n 

Morakinyo, 

1995a; 1995b 

= 1008) 

Interviews of 

harvesters, traders and 

artisans in Nigeria (n 

= not specified) 

Ndoye, 1994 Interviews with 

farmers/harvesters (n 

= 52) 

Townson, 1995 Internviews with 

entrepreneurs (all 

NTFPs) in Ghana (n = 

955) 

Defo, 1999; 

Trefon and Defo, 

1998 

Interviews with rattan 

harvesters in 

Cameroon (n = 84) 

Sunderland, 1998 Interviews with 

harvesters and artisans 

in Bata, Equatorial 

Guinea (n = 25) 

This study Interviews with urban 

artisans in Cameroon 

(n = 174) 

stakeholders 

Huge fluctuations in prices 

realised (scarcity? transport?); 

monopolistic nature of trade 

(middle-men); rotting of cane 

before it reaches market; crude 

tools available for 

transformation. 

Scarcity of cane; lack of adequate 

equipment for processing 

Scarcity of cane; open access 

nature of harvesting (“outsiders” 

harvesting); lack of control by 

Forestry Department over quotas; 

rotting of cane before reaching 

market 

Increasing scarcity of wild 

resource; would like to plant 

rattan but do not know how 

Scarcity of raw material 

(including cane) due to forest 

clearance 

Scarcity of cane = alternative 

income generating activities are 

necessary (bushmeat hunting) 

Seasonal scarcity of cane; poor 

quality of transformation and 

processing (need for training); 

Scarcity of cane; need for 

cultivation; poor quality of 

transformation and processing 

(lack of training); lack of artisan 

union (high competition); lack of 

credit facilities; lack of 

machinery/technology for 

processing. 

8.6 AMOUNT AND VALUE OF THE TRADE 

278 

Summary of 

recommendations 

Need to undertake socioeconomic/marketing 

research 

on rattan to capture real value; 

need to “cure” cane after 

harvesting; improve methods of 

processing 

Include cane in rural forestry 

programmes, planting of cane in 

buffer zones; plantation 

development; better processing 

methods 

Plantation development; 

enrichment planting; 

sustainable forest management 

in context of community forest 

management; village-based 

techniques of curing cane 

introduced 

Introduce rattan into 

agroforestry systems 

Interventions should include 

community forest management 

and investigate ways of 

increasing supplies through 

cultivation. 

Regular supply of cane to 

artisans will reduce need for 

other forms of destructive forest 

use. 

Ensure regular supply of cane 

through better forest 

management and cultivation; 

training in improved processing 

and transformation 

Ensure regular supply of cane 

through better forest 

management and cultivation; 

training in improved processing 

and transformation; encourage 

development of artisan union; 

invest in central processing 

plant 

Although it has been speculated by all of the studies cited above that the rattan trade is 

extremely lucrative, very little quantitative or comparative study has been undertaken 

in this regard. Table 27 (below) summarises the findings of some of these studies 

where quantification of the field data has been possible.

Table 32. The scale and value of the African rattan trade in selected urban markets 

City (country) Population 

(sample size) 

Lagos (Nigeria) 10,712,800 

(not known) 

Accra (Ghana) 1,512,800 

(27 enterprises) 

Kumasi (Ghana) 602,000 


Ankasa (Ghana) not known 

(12 markets) 

Bata (Equatorial 

80,000 

Guinea) 


Douala 

1,262,000 

(Cameroon) 


Yaounde 

1,157,400 

(Cameroon) 

Kinshasa (DR 


Congo) 


Estimated amount 

of cane used / 

month (m) 

279 

Estimated mean 

annual value 

(US$) 

Reference 

180,000m 1,041,180 Morakinyo (1994) 

not known 64,080 Falconer (1994) 

not known 95,475 Falconer (1994) 

4,300 m (all species) 62,000 Holbech (2000) 

20,550m (all 

species) 

26,955m (large dia.) 

28,875m (small dia.) 


29,765 (small dia.) 


14,448 (small dia.) 

27,400 Sunderland 

(1998) 

127,405 This study 

103,500 This study 

Figure 123. The scale and value of the African rattan trade in selected urban markets 

Annual value (US$) 

160,000 

140,000 

120,000 

100,000 

80,000 

60,000 

40,000 

20,000 


0 

27,400 

Bata (E Guinea) 

56,600 

Kinshasa (DR 

Congo) 

62,000 

Ankasa region 

(Ghana) 

64,080 

Accra (Ghana) 

95,475 

Kumasi (Ghana) 

103,500 

Yaounde 

(Cameroon) 

127,405 

Douala 

(Cameroon) 

56,600 Minga (in press) 

141,180 

Undoubtedly, the commercial rattan sector in Africa is thriving. However, in common 

with much of the forest exploitation of the region, the rattan trade is characterised by 

the inequitable distribution of benefits, over-exploitation and the lack of contribution 

Lagos (Nigeria)

of this trade, in fiscal terms, to the formal forestry sector. Although significant 

headway has been made with regard to the development of forestry legislation to 

encourage a more holistic approach to forest management, it will take a considerable 

paradigm shift before forest resources are indeed managed on such a basis. There is 

considerable potential for rattans to be managed on a sustainable basis (Sunderland 

and Dransfield, in press) and there is, additionally, considerable potential for the 

rattans of Africa to contribute to the thriving global rattan trade. However, the 

ecological, social and institutional conditions needed for the development of 

sustainable strategies, whilst ensuring the equitable distribution of benefits, must be in 

place before this can be considered. 

280

CHAPTER NINE 

IMPLICATIONS FOR CONSERVATION AND DEVELOPMENT 

9.1. INTRODUCTION 

It is widely reported that the thriving domestic trade in rattan and rattan products in 

Africa has undoubtedly led to substantial over exploitation of the wild rattan resource 

(Morakinyo, 1994; Ndoye, 1994; Falconer, 1994; Townson, 1995; Trefon and Defo, 

1998; Defo, 1997; Sunderland, 1998; Defo, 1999; Sunderland 1999a; 1999b; Kialo, 

1999; Minga, in press; Holbech, 2000; Oteng-Amoako and Obiri-Darko, in press; this 

study). This exploitation, coupled with the loss of forest cover through logging and 

subsequent agricultural activities, could, in future, threaten the very survival of the 

rattan industry in Africa as has been pessimistically forecast in SE Asia (Dransfield, 

1988a). The detrimental impact of the decline of wild rattan resources is most clearly 

realised by local rattan collectors and urban artisans that rely on forest products; 

individuals and communities already at the lower end of the economic scale. 

In essence, the sustainable harvesting and management of the African rattan resource 

is primarily hindered by a paucity of a sound information on stocking, growth, yield 

and harvest intensity. In addition, the lack of resource tenure precludes any attempts at 

long-term and sustainable harvesting and the fact that rattan is considered an “openaccess” 

resource throughout much of its range mitigates the prospects for long-term 

sustainable management. 

9.2. CONSERVATION STATUS OF AFRICAN RATTANS 

Now that the taxonomic base of the African rattan sector has been established it is 

now possible to determine the global conservation status of the species concerned. 

This has been calculated using the IUCN (1998) conservation categories where among 

other criteria, geographical limits in species distribution are used to determine the 

conservation status. These categories are as follows: 

Endangered (species distribution

Not threatened (species distribution > 20,000 km²) 

Table 33. The conservation status of African rattan species 

Species Geographical range 

(km²) 

IUCN Category 

Calamus deërratus 8,049,170 Not threatened 

Eremospatha barendii One collection only Endangered 

E. cabrae 1,918,050 Not threatened 

E. cuspidata 1,891,190 Not threatened 

E. haullevilleana 2,703,930 Not threatened 

E. hookeri 1,102,420 Not threatened 

E. laurentii 2,731,880 Not threatened 

E. macrocarpa 4,259,660 Not threatened 

E. quinquecostulata 9,276 Vulnerable 

E. tessmanniana 5,899 Vulnerable 

E. wendlandiana 604,086 Not threatened 

Laccosperma acutiflorum 1,485,230 Not threatened 

L. laeve 1,226,210 Not threatened 

L. opacum 1,807,940 Not threatened 

L. robustum 1,537,390 Not threatened 

L. secundiflorum 3,195390 Not threatened 

Oncocalamus macrospathus 701,830 Not threatened 

O. mannii 129,432 Not threatened 

O. tuleyi 18,423 Vulnerable 

O. wrightianus 2,872 Endangered 

9.3. HARVEST AND MANAGEMENT 

9.3.1. Growth rates 

Rattans are vigorous climbers with relatively high growth rates, and are thus able to be 

harvested on a short cycle. For the majority of rattans, stem production from the 

rosette stage (and the seedling bank) is initiated by exposure to adequate light. Stem 

elongation is also affected by light and, whilst continuous, varies, usually on a 

seasonal basis. Whilst no data on the growth rates of African rattans is available as 

yet, results of long-term studies of the growth rates of selected Asian taxa are known. 

Table 34. The growth rates of some commercial rattans in cultivation (Modifed from Dransfield 

and Manokaran, 1994). 

Species Growth rates (m / year) 

Calamus caesius 2.9-5.6 

Calamus egregius 0.8 

Calamus hainanensis 3.5 

Calamus manan 1.2 

Calamus scipionum 1.0 

Calamus tetradactylus 2.3 

Calamus trachycoleus (5.0) 

Daemonorops margaritae (2.0-2.5) 

(Figures in brackets are estimates) 

282

9.3.2. Management 

In general, the wild rattan resource is rarely managed. In many areas, rattan canes are 

considered an “open-access” resource and the lack of resource tenure of this product is 

undoubtedly contributing to its reputed scarcity in areas of high exploitation. 

However, in SE Asia in particular, a number of recognised management systems are 

in place (Sunderland and Dransfield, in press). In this respect, four main types of 

management have been identified: 

9.3.2.1. Natural regeneration in high forest 

This level of management requires the development and implementation of 

management plans based on sound inventory data and an understanding of the 

population dynamics of the species concerned. This is particularly appropriate for 

forest reserves, community forests and other low-level protected areas. These 

“extractive reserve” models are highly appropriate for rattan: a high value, high 

yielding product that relies on the forest milieu for its survival. 

9.3.2.2. Enhanced natural regeneration 

This management system in undertaken through enrichment planting and canopy 

manipulation, in natural forest. This is especially appropriate where forest has been 

selectively logged (more so in the Dipterocarp forest of SE Asia, rather than the more 

selective logging regimes practised in Africa). Management inputs are fairly high, 

with the clearance of competing undergrowth vegetation and subsequent selective 

felling to create “artificial” gaps has been practised in India, with some success for the 

rattan resource. Rattan planting in forest in East Kalimantan has also proved 

successful. 

9.3.2.3. Rattan cultivation as part of shifting cultivation or formal agroforestry 

systems 

The incorporation of rattan into traditional swidden fallow systems in some areas of 

SE Asia is well known (Connelly, 1985; Siebert and Belsky, 1985; Peluso, 1992; 

Weinstock, 1983). The general principle is that, on harvesting ephemeral or annual 

crops, rattan seedlings are planted and the land is then left fallow. When the rotation is 

repeated, usually on a 7-15 year cycle, the farmer first harvests the rattan and then 

283

clears the plot again to plant food crops. The income generated from the harvesting of 

rattan in this way is significant. 

9.3.2.4 Silvicultural systems 

Intensive cultivated forestry-based systems have concentrated on the incorporation of 

rattan into tree-based plantation-type systems. The need for a framework for the rattan 

to grow on is imperative and the planting of rattan in association with tree cash crops 

was begun in the 1980s. In particular, planting under rubber (Hevea brasiliensis) and 

other fast-growing tree crops has proven relatively successful and both silvicultural 

trials and commercial operations, under a wide rage of parent crops are commonlyencountered 

throughout SE Asia. 

9.3.3. Harvest procedures 

The harvesting techniques employed in the extraction of rattan, and which are 

generally the same despite geographic differences, have an impact on potential 

sustainability, particularly for clustering species. The mature stems selected for 

harvesting are those without lower leaves (i.e. where the leaf sheaths have sloughed 

off) and usually only the basal 10-20m is harvested; the upper “green” part of the cane 

is too soft and inflexible for transformation and is often left in the canopy. In many 

instances, all the stems in a clustering species may be cut in order to obtain access to 

the mature stems, even those that are not yet mature enough for exploitation and sale. 

This is particularly an issue where resource tenure is weak. 

In general, two simple interventions can be implemented to improve upon rattan 

harvesting practices: 

For clustering species; 

· Younger stems, often indiscriminately cut during harvesting should be left to 

regenerate and provide future sources of cane. Rotational harvesting systems 

could be increased if this was the case. However, better “stool management” 

relies on adequate resource tenure. 

For all species (including solitary species): 

284

· Harvest intensity and rotation should be based on long-terms assessments of 

growth rates and recruitment. 

9.3.4. Inventory 

As discussed in Chapter 3, rattan inventory has proved to be a somewhat imperfect 

science. Initial attempts to determine stocking and yield have often been thwarted by a 

poor taxonomic base from which to begin. Furthermore, lack of sampling the correct 

parameters also led to much inventory information being simply discarded. When 

planning a rattan inventory it is essential to: 

· Know the species concerned (collect herbarium specimens when in doubt). 

· Measure the correct parameters. These include: 

· Number of stems per clump 

· Number of clumps per hectare, or for solitary canes, stems per hectare 

· Total stem length 

· Harvestable stem length (the lower stem portion with the leaves 

sloughed off. 

· Establish a protocol for measuring over time to determine growth rates and 

recruitment; this will determine the potential harvest and hence sustainable 

extraction rates. 

9.4. LAND TENURE AND SOCIO ECONOMIC ISSUES 

Rattan management of whatever kind will only be a success if those involved have 

clear access to the forest, and/or long, and easily renewable resource rights on it. 

Currently, rattan collectors rationally maximise their income by harvesting the best 

and most accessible canes, because they are paid on a per item basis. Larger canes 

bring the best prices and it is also important to minimise the opportunity costs of 

collection (i.e. the rattans closer to the community will be harvested first, and 

probably more intensively). 

Traditionally, many communities in SE Asia and Africa have benefited from the 

harvesting of their rattan resource. Many of these groups are dependent on high-value 

285

forest products, such as rattan for access to the cash economy. However, local scarcity 

caused by uncontrolled harvesting is denying many local people access to this 

traditional means of income, let alone access to the resource for their own subsistence 

needs. 

However, the recent paradigm shift towards community-based forest management 

provides a unique opportunity to impart significant levels of tenure to non-timber 

forest resources in particular. Formal legislation to reflect this is being introduced 

throughout West and Central Africa and it is hoped that the more holistic approach to 

forest management that this legislation enables will provide the conditions under 

which long-term sustainability can operate. 

9.5. RATTAN CULTIVATION 

In SE Asia, many examples exist of rattans being cultivated in agroforestry systems in 

forest lands controlled by local communities (Connelly, 1985; Siebert and Belsky, 

1985; Peluso, 1992; Weinstock, 1983). For example, In Kalimantan, Indonesia, smallscale 

rattan cultivation has long been practised by forest dwellers. Rattan seedlings are 

removed from the forest and then incorporated into abandoned farm fallows soon after 

the harvest of the annual crops. The rattans grow and develop along with the 

surrounding secondary vegetation. When the land is cleared again after a fallow 

period of up to 15 years, the rattan is harvested and either sold or is retained for 

household use (Peluso, 1992). 

However, despite its local importance, the scale of such cultivation is negligible, and 

the majority of the cane entering international trade originates from the wild. Along 

with this wild harvesting, intensive logging activities and the associated increase in 

forest conversion led to shortages of rattan became apparent in the early 1970's and 

forest departments in the SE Asian region began to investigate the possibilities of 

commercial rattan cultivation. The first steps taken towards this aim resulted in a 

complete taxonomic inventory of wild Malaysian rattans and the establishment of 

research plots that have been used to determine optimum conditions and subsequent 

guidelines for the cultivation of rattans (Dransfield and Manokaran, 1992). To date, 

commercial plantations have been established in Peninsular Malaysia, Sarawak, 

286

Sabah, China, the Philippines, Indonesia and Papua New Guinea. Whilst there remain 

many unsolved problems concerning the cultivation of rattan, over 20,000ha of 

plantations have been established in Malaysia alone. 

In many areas of SE Asia, the large-scale cultivation of rattan has concentrated on: 

· The development of plantation systems with the use of a parent tree crop 

(rattan is a climber than needs arboreal support). Rubber (Hevea brasiliensis) 

has been predominantly used, but latterly a wider variety of tree crops have 

been experimented with. The majority of the commercial cultivation schemes 

favour this type of cultivated system. 

· Enrichment planting with rattan after logging in secondary forest. Rattan is 

planted and maintained in logged-over lowland Dipterocarp forest. 

Figure 124. Three-year old planting of Calamus merrillii with parent crop of five-year old Albizia 

falcata in Sabah, Malaysia. 

It is important to note, however, that many of these plantations and cultivated sources 

of cane are owned and managed by sovereign forestry departments or private 

companies. Hence, the revenues accrued do not often find their way to local 

communities at it would if they were harvesting directly from their own agroforestry 

systems. In many ways, commercial cultivation leads to the removal of a resource 

287

from the informal forest economy and into the formal forestry sector; a system 

renowned for its inequity (de Fretas, 1990). 

In Africa, until recently, no cultivation of any of the species of commercial 

importance has been undertaken, nor has there been any proven history of indigenous 

cultivation. However, the regular supply of raw cane to a thriving domestic cottage 

industry has been affected by over-harvesting and poor management of the wild 

resource to such an extent that, in order to mitigate the effects of this scarcity, many 

calls have been made by both government and development agencies for the inclusion 

of African rattans into cultivated systems. It is only recently that the commercially 

important species have been identified and subsequent work into the cultivation of 

rattans of Africa has led to the development of a rattan arboretum at the Limbe 

Botanic Garden, Cameroon. More recently, in collaboration with the Cameroon 

Development Corporation, this propagation work has also resulted in the 

establishment of a one-hectare trial of Laccosperma secundiflorum in an obsolete 

rubber plantation. Now that the cultivation requirements of the commercial species are 

understood, feasibly, it is now possible to incorporate these species into a wider range 

of cultivation systems. 

9.6. CONCLUSION 

The crux of the sustainability issue may be summarised as follows: How is it 

possible to create the conditions to enable the sustainable utilisation of the rattan 

resource, whilst maintaining, or increasing, profits and benefits? 

Given an appropriate management regime, the ecology and nature of rattans make 

them one of the few forest products that can be harvested sustainably. Rattan 

exploitation, be it in natural forest or in agroforestry systems, relies on an intricate and 

multi-layered ecological balance between the rattan resource and the trees that are 

needed to support it. There are very few products of such high value that require the 

maintenance of a forest analogue. The fact that rattan is fast growing, high yielding 

and can be harvested on relatively short rotations also makes it attractive for 

sustainable management regimes; the impact of harvesting can be measured and 

288

monitored within a few years and the harvesting regime and adjusted accordingly to 

ensure a sustained yield. 

Not only should ecological sustainability be a guiding factor in determining the 

inclusion of rattans within a forest management system, but the social acceptability of 

the exploitation system should also be a prime consideration. In terms of ensuring that 

the benefits of rattan exploitation benefit forest dwellers and forest-based 

communities, conservation and development efforts should concentrate on the 

sourcing of rattans from managed high forest and/or agroforestry systems. Not only 

does this ensure a level of ecological integrity is maintained, but also, those 

traditionally associated with rattan harvesting, can benefit from the wise management 

of the resource. Forest management programmes could perhaps be the stimulus behind 

initiatives towards better forest management and the attainment of adequate resource 

tenure. This is a seemingly impossible goal at present, especially with the present 

situation of disenfranchisement through the establishment of commercial rattan 

monocultures under a parent mono-crop. 

Harvesting levels and quotas need to be determined based on growth rates and 

estimates of recruitment. The establishment of these quotas is hindered by the 

considerable paucity of baseline information for most, if not all taxa. Undoubtedly, a 

great deal more basic ecological and applied research is needed before management 

regimes for most species of commercial interest can be suggested. 

Finally, the rattan resource is being regarded as being a prime candidate for forest 

certification initiatives that are now concentrating on a wider range of forest products 

other than timber (Sunderland and Dransfield, in press). The certification process is 

perhaps the most beneficial means of ensuring the above conditions need for the 

sustainable management of the rattan resource in Africa, and SE Asia, are fulfilled. 

289

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319

APPENDIX ONE 

INDIGENOUS NOMENCLATURE AND UTILISATION OF 

INTRODUCTION 

AFRICAN RATTANS BY SPECIES 

Rattan canes are used extensively across West and Central Africa by local 

communities and play an important role in indigenous subsistence strategies for many 

rural populations. As such, the range of indigenous uses of rattan canes is vast. These 

uses are listed below by species. Full descriptions of the species concerned can be 

found in Chapter 2. The vernacular names recorded are from the examination of 

herbarium records and notes, a literature review (cross-referred to herbarium records) 

and from my own field observations. A detailed discussion of the nomenclature and 

classification of the African rattans, based on the information listed below, is provided 

in Chapter 6. 

___________________________ 

Calamus deërratus G. Mann & H. Wendl. 

Vernacular names 

SENEGAL: ki tid (Balanta); kintem (Bainouk); mantampa da sera (Crioulo, Upper 

Guinea); bu kètao bu ketav, fu fiaf, ka kèt, ka tay, ke hiya, kékiya (Jola-Fogny); 

tambem (Fula-Pulaar); tambi (Tukulor); tambo (Mandinka); tãbi (Malinke); e kapat 

(Mandyak); ratlan (Wolof): GAMBIA: tambo (Mandinka): GUINEA-BISSAU: 

quitite (Balanta); batanou (Biafada); mantampa de sera (Crioulo, Upper Guinea); 

tambem (Fulfulde-Pulaar); tambo (Mandinka); ecapate (Mandyak); quito (Papel): 

GUINEA: tambo (Mandinka); tâbi (Malinke): SIERRA LEONE: lumboinyo-lando 

(Kisi); kanga-mese (Kono); tambe (Loko); tambi (Maninka); tamba (def. tembui) 

(Mende); tambi (Susu); ra-gbet (Themne); tambu-na (Yalunka): LIBERIA: kpa kala 

(Mano): CÔTE D’IVOIRE: ailé-mlé (Anyin); gapapa (Godié): GHANA: dem4eré 

(Twi, also trade name); néné, (Akan); ayeka (Anufo); ayeka (Sehwi); keteku (Éwé); 

ayeké (Nzema): BENIN: akete (Defi); dekun wéwé (Gun-Gbe): NIGERIA: water cane 

320

(Pidgin); erogbo, erugbo (Edo); ekwe-oji, iye (Igbo); apié (the plant itself, or the canerope 

made from it) (Ijo-Izon), bwálàm (a cane) (Pero); erogbo, erugbo (Yoruba): 

CAMEROON: nding (Bulu); ndié (Badjué): EQUATORIAL GUINEA: nzing (Fang): 

CENTRAL AFRICAN REPUBLIC: bioh (Banda-Yangere): DR CONGO: kpude 

(Zande); ma-ndakele (Ngbaka-Ma’bo); ikonga (Lombo); babio (Mongo-Nkundu); mukolo 

(Lega-Mwenga); lekwe (BaMbuti): UGANDA: bi-lekwe (Amba). 

Uses 

The stems of C. deërratus are not as widely used as other species. Throughout their 

range, they are considered inferior to other species of rattan as they do not attain as 

great a diameter and hence are not as robust as many of the larger species of 

Laccosperma. Accordingly, they are often too thick and rather inflexible for weaving, 

for which a number of species of Eremospatha are preferred. However, the use of C. 

deërratus increases in the absence of other rattan canes (Profizi, 1986; Morakinyo, 

1995; Tenati, in press) and, aside from a wide range of construction and weaving 

applications, C. deërratus also contributes significantly to other household needs. 

For example, in Ghana, the whole leaves are often used for thatching (Abbiw, 1990). 

In Sierra Leone the apical bud (palm heart) is eaten by the Mende people as a dietary 

supplement and they are in Ghana (Dalziel, 1937; Irvine, 1952; Irvine, 1961). The 

young shoots are roasted whole and eaten in Ghana (Abbiw, 1990). In lower 

Casamance of Senegal the leaves are grilled over a fire and then macerated; the liquid 

is drunk to promote weight loss is stout persons (Berhaut, 1988) and the same 

preparation is also given for oedema caused by vitamin deficiencies (ibid.). Ash from 

burning the roots is used as a kitchen salt in Guinea (Portères, s.d. cited in Burkhill, 

1997) and in Ghana (Abbiw, 1990). In Sierra Leone, the rind is twisted into little 

sponges and used to clean pans (Irvine, 1961). Williamson & Timitimi (1983) state 

that the leaf sheath may be peeled off and twisted to make a rough rope. 

Throughout West Africa in particular, C. deërratus has a wide range of uses for 

furniture construction and basketry to the extent that in the dry savannas of northern 

Nigeria, in the absence of other species, canes of C. deërratus were formerly supplied 

to prisons for workshop activities and the rehabilitation of prisoners (Burkill, 1997). 

Whole canes were widely used for fences and house-building in other areas of Nigeria 

(Unwin, 1920) as well as in Ghana (Abbiw, 1990). Elsewhere in Africa, in Zambia 

321

and Uganda, in the absence of large-diameter canes, rattan furniture frames are made 

of two or three stems of C. deërratus joined together (Tenati, in press). When the 

canes are split and woven, they have a multiplicity of purposes such as the fabrication 

of “tie-ties” for the staking of yams (Morakinyo, 1995), they are woven and used for 

fencing in Ghana (Abbiw, 1990) and as binding materials for hut-making and to tie 

thatch, (Berhaut, 1988; Dalziel, 1937). In addition, both whole and split stems are also 

used to make hammock suspension bridges (Dalziel, 1937; Abbiw, 1990) as well as 

making a foot-loop for climbing palm trees (Dalziel, 1937). More everyday uses 

include the fabrication of fish traps and weirs, kola baskets, bow strings household 

matting and screens and the seat of chairs (Dalziel, 1937; Irvine, 1961; Abbiw, 1990). 

In Nigeria, the split stems were formerly used to tie timber tree together prior to them 

being floated down river for sale (Sylvic Sap, 1935) and in the continental region of 

Equatorial Guinea they were formerly used in the fabrication of temporary market 

baskets (Guinea-Lopez, 1946), although this is not the case today (Sunderland, 1998). 

References: SENEGAL: Dalziel (1937); J. Berhaut (1988); Burkill (1997): GAMBIA: Anderson 131 

(herb. MO!): GUINEA-BISSAU: Espirito Santo, 1963 ex auctt. cited in Burhill, 1997): GUINEA: 

Dalziel (1937): SIERRA LEONE: Dalziel (1937); Deighton, vocabulary (1957) - cited in Burkhill, 

(1997); Scott-Elliot 4738, 5121 (herb. K!); Thomas 2753 (herb. K!): LIBERIA: Dalziel (1937): CÔTE 

D’IVOIRE: ECOSYN (1999): GHANA: Dalziel (1937); Irvine (1961); Burkhill (1997); Vigne 1868 

(herb K!); Chipp 127 (herb. KUM!): BENIN: Profizi (1986); Aufsess 425 (herb. K!): NIGERIA: 

Dalziel (1937); Dunstan s.n. (herb. K!); Russell unpubl. notes; Williamson & Timitimi, (1983) - cited in 

Burkhill (1997); Bennet (herb. K!); Unwin 223 (herb. K!): CAMEROON: Mildbraed (1913): 

EQUATORIAL GUINEA: Guinea-Lopez (1946); Tessmann 6 (herb FI!); Sunderland (1998): 

CENTRAL AFRICAN REPUBLIC: Harris 820 (herb. K!): DR CONGO: Robyns & Tournay (1955); 

Evrard 1876 (herb. BR!); Louis 15541 (herb. BR!); Germain 210 (herb. BR!); Leonard 832 (herb. BR!); 

Troupin 296 (herb. BR!); Hart 633 (herb MO!): UGANDA: Makombo s.n. (herb. K!). 


None recorded. 

________________________________ 

Eremospatha barendii sp. nov. 

322

Uses 


________________________________ 

E. cabrae de Wild. 


GABON: osono (Tsogo); osono (Pinji); ozono (Myene); li-bamba (Vili); nkolé (Kélé); 

nkolu (Seki); du-bamba (Barama); du-bamba (Lumbu); ivéta (Duma); iló-lóngo 

(Kota); u-lóngo (Benga); lé-mbumu (Ndumu); nlong (Fang): DR CONGO: li-findo 

(Lombo); lu-bambi (Kituba); e-safa (Mongo-Nkundu); ki-sakata (Kete): ANGOLA: 

m’bamba (Mbundu-Luanda). 

Uses 

In Gabon, the split stems of this species are widely used for the fabrication of 

temporary market baskets (Raponda-Walker and Sillans, 1961). In the Congo 

Republic, the complete stems of this species are used for furniture frames, or are split 

and used for weaving, particularly by forest dwellers (Profizi and Makita-Madzou, 

1996). The stems are also employed for the construction of cane bridges (ibid.). 

However, in the absence of E. haullevilleana, the split stems and epidermis of E. 

cabrae are used for weaving and basketry in DR Congo. In Angola, Gossweiler 

reported that, particularly in the absence of other species, the whole stems of E. 

cabrae, were formerly used for the fabrication of armchairs that were “much 

appreciated by the colonists” and many were also exported to Portugal (Chevalier, 

1936). Another main use of this species is the use of the leaf sheath as a toothbrush by 

the Lombo of DR Congo. 

References: GABON: Raponda-Walker and Sillans, 1961: DR CONGO: Compère 2182 (herb BR!); 

Germain 326 (herb. BR!); Hulstaert 1417 (herb BR!); Jans 655 (herb BR!); Kuasa (herb BR!); Leonard 

929 (herb BR!); Louis 16797, 3804, 5656, 15169 (herb BR!); Toussaint 2331 (herb BR!): ANGOLA: 

Chevalier (1936). 

________________________________ 

323

E. cuspidata (G. Mann & H. Wendl.) H. Wendl. 


EQUATORIAL GUINEA: ndera (Fang) 

Uses 

The split stems are used for light basketry and weaving, particularly in the absence of 

other species. 

References: EQUATORIAL GUINEA: Guinea-Lopez (1946); Sunderland (1998) 

________________________________ 

E. haullevilleana De Wild. 


CENTRAL AFRICAN REPUBLIC: pongbo (Ngombe): CONGO: mbaama (Téké): 

DR CONGO: li-findo (Lombo); mbowe (Zande); lu-popi ((Nandi)); n’kele (Bangala); 

m’bio (Bangi); lo-koli (Kele); ke-kele (Lingala); lu-kodi (Luba-Shari); lu-busi 

(Tembo); lu-bubi (Lega-Mwenga); yofoko (Mungo-Nkundu); lo-keko (Lusengo); kodi 

(Luba-Kasai); tukpuru (Bhele): UGANDA: bibbobbi (Amba); enga (Luganda): 

TANZANIA: urugage (Ha) 

Uses 

The preferred species for basketry, weaving and furniture manufacture throughout its 

range, even more so than E. macrocarpa. Profizi and Makita-Madzou (1996) state this 

species as being the best for weaving, possessing important qualities of strength, 

durability and resistance to insect attack. Because of these qualities, the Téké Tsaaya 

of Congo-Brazzaville use this species widely and it is considered second only in 

importance to the oil palm (Elaeis guineensis) (ibid.). The stems are used whole for a 

wide range of products including for use as cables for cane bridges, furniture 

framework and building frames. The split stems are used for the fabrication of fish 

traps, noose-type snares catch to small terrestrial mammals and for the handrails of 

river bridges made from the felled stems of Musanga cecropioides, as well as for a 

wide array of baskets. In the Bambama district, the apical bud of this species is widely 

324

consumed. During certain traditional rites, the stem of this species is respected as 

providing protection for the bearer. When the village is expected to be visited by 

malevolent spirits, the stems of this species are placed through the settlement, adorned 

with powerful fetishes (ibid.). 

In the DR Congo, the fruits are used by the Bangi for decoration, particularly in the 

manufacture of traditional collars (Dewevre, herb. BR!). The Lombo use the 

acanthophylls as fish hooks (Louis, herb BR!) and use the sap (the watery exudate 

collected when the stems are cut) as an abortifacient (Staner and Boutique, 1937). 

In Tanzania, the split stems are used by the Ha for sewing bark (Xylopia sp?) beehives 

and food containers (Proctor, herb. K!). 

References: CENTRAL AFRICAN REPUBLIC: Carroll 115 (herb. MO!); Harris 2360 (herb. MO!): 

CONGO: Profizi & Makita-Madzou (1996): DR CONGO: Couteaux 471 (herb. BR!); Dewevre 581 

(herb. BR!); Gathy 1638 (herb. BR!); Gerard 1432 (herb. BR!); Gillet 167 (herb. BR!); Gutzwiller 539 

(herb. BR!); Herman 2138 (herb. BR!); Hulstaert 1418 (herb. BR!); Kitembo 60 (herb. BR!); Leclercq 

736 (herb. BR!); Leonard 932, 933, 936, 1138 (herb. BR!); Liben 2603 (herb. BR!); Louis 3395, 1970, 

772, 847, 3638, 11850, 9731, 12106, 9560, 8106, 7671, 9420, 16775 (herb. BR!); Nannan 117 (herb. 

BR!); Terashima 94 (herb. BR!); Troupin 9162, 2658 (herb. BR!); Gerard 1432 (herb. BR!); Minga (in 

press): UGANDA: Makombo s.n. (herb. K!); Kabuya (1988): TANZANIA: Proctor 369 (herb. K!) 

________________________________ 

E. hookeri (G. Mann & H. Wendl.) H. Wendl. 


SIERRA LEONE: balu (Kono); mbalu (def. -ui) (Mende); ra-thamp (Themne): 

NIGERIA: epa-emele (Yoruba); inima ború (Ijo-Izon); itomi (Ekit): CAMEROON: 

ki-yince (Balundu-Bima); mbunden (Bakundu-Balue): EQUATORIAL GUINEA: 

alua-nlong (Fang): GABON: gigorula (Sira): 

Uses 

The stem epidermis is easily peeled and is used in Sierra Leone by the Mende as a 

rope and for making baskets (Deighton, 1956; Russell, s.d.). The split cane itself is 

325

also a strong binding material (Russell, s.d.; Small, herb. K!). In Nigeria, the Ekit use 

the base of the leaf sheath as a chewstick (Morakinyo, herb K!). 

References: SIERRA LEONE: Deighton (vocabulary, 1956 cited in Burkill, 1997); Small 832 (herb. 

K!); Deighton 2591 (herb. K!): NIGERIA: Ainslie (1937); Okigbo (1980); Williamson & Timitimi 

(1983); Morakinyo 1005 (herb. K!); Ayewoh 3852 (herb. K!): CAMEROON: Thomas et al., 1989); 

Sunderland (unpubl. notes): EQUATORIAL GUINEA: Sunderland, 1998: GABON: le Testu s.n. 

(herb. BR!). 

________________________________ 

E. laurentii de Wild. 


SIERRA LEONE: bongei (Mende): CAMEROON: kpakpa (Ewondo): CENTRAL 

AFRICAN REPUBLIC: bo-kondi (Banda-Yangere): EQUATORIAL GUINEA: ebuat 

(Fang): DR CONGO: bo-ngale (Mongo-Nkundu); ikonga (Lombo); nkelele mo-none 

(Lingala); nkoli (Bali). 

Uses 

This species is rarely used for either furniture manaufacture or basketry as the cane is 

known to be of somewhat poor quality. 

References: SIERRA LEONE: Deighton 4117 (herb. K!): CAMEROON: Letouzey 11796 (herb. YA!): 

CENTRAL AFRICAN REPUBLIC: Harris and Fay 459 (herb. BR!): EQUATORIAL GUINEA: 

Sunderland (1998): DR CONGO: Leonard, 55, 816, 980 (herb. BR!); Louis 15925, 15944 (herb. BR!); 

Dubois 912 (herb. BR!); 

________________________________ 

E. macrocarpa (G. Mann & H. Wendl.) H. Wendl. 


SIERRA LEONE: penden (Kissi); balu (Kono); mbalu (Loko); mbalu, koto mbalu = 

juvenile (Mende); ra-thamp (Themne): LIBERIA: b�l� de b�l� (Mano): CÔTE 

D’IVOIRE: ailè-mlé (Anyin): GHANA: mfia (Akan-Asanti); néné (Nzima): BENIN: 

dekon (Defi); dekun vovo (Gun-Gbe): NIGERIA: white cane (Pidgin); �kan (Edo); 

odu-a�� (Igbo); b�r� (Ijo-Izon); ukan (Yoruba); ekakieri = male (i.e. with no 

326

fruits), irrumka = female (with fruits) (Ekit); iro (Esan); obong (Efik); dugwah 

(Iwuru): CAMEROON: filet (Trade); cane rope (Pidgin); e-chié (Denya); nlong 

(indef.) melong (def.) (Bulu); bana ndongo = young cane (bana = child) (Balundu- 

Bima); yo-chori (Korop); nloun (Baasa); lo’o (Badjué): EQUATORIAL GUINEA: 

nlong (indef.) mi-long (def.) = juvenile stems, ongam = adult (Fang): GABON: kegèma 

(Lumbu); nyèvila (Sira); ongam (Fang); ndètèse (Kota); iganga-tsungu (Punu); 

songu (Vumbu); tongo (Tsogo); mbubi (Ndumu) 

Uses 

This species is reputed to be the best source of cane in Africa (Unwin, 1920) and is 

reputed to be of comparable quality to the small-diameter canes of SE Asia 

(Mildbraed, 1913; Irvine 1961). E. macrocarpa is widely used for furniture 

construction, basketry, weaving and tying wherever it occurs: Sierra Leone (Burkill, 

1997), Côte d’Ivoire (ECOFAC, 1999), Ghana (Irvine, 1961; Abbiw, 1990), Benin 

(Profizi, 1989), Nigeria (Morakinyo, 1994; Tuley, 1995), Cameroon (Defo, 1998; 

Sunderland, 1999), Equatorial Guinea (Sunderland, 1998; Guinea-Lopez, 1946), 

Gabon (Raponda-Walker and Sillans, 1961). However, E. haullevilleana seems to be 

the preferred species for basketry and weaving in the DR Congo. 

In Côte d’Ivoire, the Anyi utilise the split stems for baskets, cages for transporting 

chickens to market and for bows to play stringed musical instruments (ECOSYN, 

1999). In Nigeria, the split stems of this species were formerly used to tie log rafts 

together for floating downstream to market (Sylvic Sap, 1935) as well as for making a 

kind of string that the Yoruba formerly used for tying lath pieces together in 

housebuilding an in tying the cloth in making coverings for canoes (Unwin, 1920). 

The powdered root is taken as a medicine for the treatment of syphilis by the Akan- 

Asanti in Ghana (Irvine, 1961; Abbiw, 1990) and the Yoruba of Nigeria (Ainslie, 

1937). The long flexible stems of this species make it ideal for the construction of 

cane bridges and these are commonly encountered in Ghana (Irvine, 1961) and by the 

many indigenous groups in SW Province, Cameroon (Sunderland, unpubl. notes). 

References: SIERRA LEONE: Deighton (vocabulary, 1956 cited in Burkill, 1997): LIBERIA: Dalziel 

(1937): CÔTE D’IVOIRE: ECOSYN (1999): GHANA: Irvine (1961); Burkill (1997); Sunderland 

327

(unpubl. notes): BENIN: Profizi (1989): NIGERIA: Dalziel (1937); Williamson & Timitimi (1983); 

Russell (unpubl. notes - cited in Burkill 1997); Morakinyo 1003 (herb. K!); Nwambin 6942 (herb, K!); 

CAMEROON: Sunderland (unpubl. notes); Balinga (1999): EQUATORIAL GUINEA: Guinea-Lopez 

(1946): Sunderland (1998): GABON: Raponda-Walker and Sillans (1961) 

________________________________ 

E. quinquecostulata Becc. 


CAMEROON: calumé-e-chié (Denya): GABON: di-bula (Sira) 

Uses 

Not used in Cameroon due to the presence of other, more desirable, species of rattan. 

In Gabon however, the stem is split and employed for basic weaving, particularly in 

the absence of other species. 

References: CAMEROON: Sunderland (unpubl. notes): GABON: le Testu s.n. (herb. BR!) 

________________________________ 

E. tessmanniana Becc. 


CAMEROON: calumé e-chié (Denya): EQUATORIAL GUINEA: ongam-akot 

(Fang). 

Uses 


References: CAMEROON: Sunderland (unpubl. notes): EQUATORIAL GUINEA: Tessmann 4 (herb. 

FI!); Guinea-Lopez (1946). 

________________________________ 

E. wendlandiana Dammer ex Becc. 

328


NIGERIA: eghounka (Ekit): CAMEROON: cane basket (Pidgin); mua-e-chié 

(Denya); nkonlo lo’o (Badjué): EQUATORIAL GUINEA: akot (Fang): GABON: 

égoo (Tsogo); ngundju (Punu); ngundju (Vumbu): CONGO: ma-bulu (Téké) 

Uses 

This species is not of much value commercially and is rarely traded. However, in the 

Cross River area of Nigeria, the split stems are used to tie bamboo and stick-framed 

houses prior to plastering with clay (Morakinyo, herb K!). The stem epidermis is also 

used for tying yams (ibid.). In Cameroon, the split stems are also used for coarse 

basketry, particularly for temporary market baskets used to transport produce to 

market which are then discarded. The Balundu-Bima also used the base of the leaf 

sheath as a chewing stick (Sunderland unpubl. notes). In the Congo, the Téké use the 

inner stem for basketry and furniture construction and also consume the apex of the 

young emerging stems (Profizi and Makita-Madzou, 1996). 

References: NIGERIA: Morakinyo 1001 (herb. K!): CAMEROON: Sunderland (unpubl. notes); 

Sylvanus s.n. (herb. K!): EQUATORIAL GUINEA: Sunderland, 1998: GABON: Raponda-Walker and 

Sillans (1961): CONGO: Profizi and Makita-Madzou (1996). 

________________________________ 

Laccosperma acutiflorum (Becc.) J. Dransf. 


NIGERIA: ukpekpe (Ekit): CAMEROON: giant cane (Pidgin): EQUATORIAL 

GUINEA: ekwass (Fang) 

Uses 

Despite its size, this species is reported to possess very poor quality cane and hence it 

is rarely used. 

References: CAMEROON: Sunderland (unpubl. notes): EQUATORIAL GUINEA: Sunderland (1998). 

_______________________________________ 

329

L. laeve (G. Mann & H. Wendl.) H. Wendl. 


CÔTE D’IVOIRE: ailé-mla (Anyin): GHANA: nguni (Wasa); tenan muhunu = “it 

lives in the world for nothing” (Twi): NIGERIA: itunibia (Ekit): CAMEROON: genomé-echié 

= “slave to cane rope” (Denya): CENTRAL AFRICAN REPUBLIC: gao 

(Banda-Yangeri): EQUATORIAL GUINEA: ndele (Fang): GABON: munyengi 

(Sira); tèkè (Tsogo) 

Uses 

The poor cane quality due to the extensive branching habit makes this species of little 

use. However, in Gabon, L. laeve is sometimes used to make a rope (Raponda-Walker 

and Sillans. 1961). In the Central African Republic, the BaBinga roast the roots of this 

species on axes, which are then eaten to improve virility (Carroll, herb. K!). 

References: CÔTE D’IVOIRE: ECOFAC (1999): GHANA: Kinlock 3237 (herb. KUM!): NIGERIA: 

Morakinyo 1001 (herb. K!): CAMEROON: Sunderland (unpubl. notes): EQUATORIAL GUINEA: 

Sunderland (1998): CENTRAL AFRICAN REPUBLIC: Carroll 33 (herb. K!): GABON: Raponda- 

Walker and Sillans, 1961) 

_______________________________________ 

L. opacum (G. Mann & H. Wendl.) Drude 


GHANA: eholobaka (Nzema); sayai (Akan-Asanti); edem (Kwawu): NIGERIA: abu 

(Edo); �kw� �ya = cane for tie-tie (Igbo): CAMEROON: liko ko’ko = “close to 

cane” (Mokpwe); ge- nomé-echié = “slave to cane rope” (Denya); aka’lo (Badjué): 

EQUATORIAL GUINEA: npue-nkan (Fang): GABON: ibulu (Myene); di-bulu 

(Sira); di-bulu (Lumbu); abulo (Kele); éboa (Tsogo) ulóngó-mwa-iki (Benga): 

CONGO: kimbana ki mukaana (Téké) 

Uses 

The tendency for this species to produce aerial branches affects the cane quality 

significantly, and L. opacum is not an important source of cane. However, in Ghana, 

330

the stem is sometimes split and is used as a binding and for making the Akan-Asanti 

baskets, kenten. In Gabon, forest workers drink the potable sap of this species, 

particularly during the long dry season when many small streams have dried up 

(Raponda-Walker and Sillans, 1961). In the Congo, the Téké eat the apical bud of this 

species; these are prepared by roasting the whole stems over, or next to, a fire (Profizi 

and Makita-Madzou, 1996). 

References: GHANA: Irvine 2300, 502 (herb. GC!); Irvine (1961): NIGERIA: Dalziel, 1937; Burkill, 

1997): CAMEROON: Cheek 5591 (herb. K!); Sunderland (unpubl. notes): EQUATORIAL GUINEA: 

Sunderland (1998): GABON: Raponda-Walker and Sillans, 1961): CONGO: Profizi and Makita- 

Madzou (1996); 

_______________________________________ 

L. robustum (Burr.) J. Dransf. 


NIGERIA: willow, hard cane (Pidgin); CAMEROON: makak, maraca (Trade); eka 

(Ewondo); nkan, aka = cleaned cane (Bulu); dikah (indef.) mekah (def.) (Bakundu- 

Balue); lo-ntong (Korop); gekwiya (Denya): CENTRAL AFRICAN REPUBLIC: gao 

(Banda-Yangere): EQUATORIAL GUINEA: nkan, aka = cleaned cane (Fang): 

GABON: asperge (nom forestier) DR CONGO: ekpale-ekpale (Bwa): li-sele 

(Lombo); nkao (Ngbaka-Ma’bo); ikoonga (Lombo) 

Uses 

This species is widely used throughout its range for its high quality cane which, in the 

main, is used whole. Previously assigned under the name, L. secundiflorum, the clear 

morphological and ecological differences between the species are marked and as a 

result most local people distinguish between these two species in their folk 

classifications. The cane of this species is used mainly for furniture and house frames 

and forms the basis of a thriving cottage industry and is widely traded (Morakinyo, 

1994; Defo, 1997; Defo and Sunderland, 1999; Sunderland 1998; 1999a; 1999b). 

331

The “palm heart” is eaten widely, from Cameroon through to Gabon (Rapona-Walker 

and Sillans, 1961; Sunderland, unpubl. notes) and contributes to the diets of many 

forest dwellers. The young leaves are eaten in stews by the Fang in Rio Muni, 

Equatorial Guinea (Guinea-Lopez, 1946). 

References: NIGERIA: Morakinyo, 1000 (herb. K!): CAMEROON: Mildbraed (1913); Hédin (1929); 

Letouzey 8479 (herb. YA!); Thomas et al., (1989); Sunderland (unpubl. notes): CENTRAL AFRICAN 

REPUBLIC: Fay 8236, 8254 (herb. MO!): EQUATORIAL GUINEA: Tessmann 2 (herb. FI!); 

Sunderland (1998): GABON: Wilks 1486 (herb. MO!): DR CONGO: Gerard, 3218, 3933, 4951 (herb. 

BR!); Germain 8278 (herb. BR!): Leonard 1671 (herb. BR!). 

_______________________________________ 

L. secundiflorum (P. Beauv.) Küntze 


SENEGAL: ka-likut (Jola-Fogny): GUINEA-BISSAU: tambem-hadje (Fulfulde- 

Pulaar); tambendjom (indef.), tambendjom-� (def.) (Mandinka): SIERRA LEONE: 

lumboinyo-piando (Kisi); kangane (Kono); kafo (Loko); kavo (def. kavui) (Mende); 

ka-gbesu = whole stems, e-gbak = leafless part of the stem (Themne): CÔTE 

D’IVOIRE: kumh (Attié); agu� (Ebrié); djoho, djolo (Krumen); ahika (Anyin); gblé 

(Godié): GHANA: willow (Trade); ayi� (Akan-Asanti); ayik� = large rattan 

(Nzema): BENIN: kpanon (Defi); kpacha (Gun-Gbe): NIGERIA: willow, hard cane 

(Pidgin); ohwara (Urhobo); okankan = whole cane, ukwen = when split (Edo); 

��5� (Efik); ukp� = cane rope made of this species (Ijo-Izon); iga (Ekpeye); 

a�� (Igbo); epe-nla, ikan-ik� = a hook (Yoruba): CAMEROON: makak, maraca 

(Trade); ka-kawa (Baka): ekwos (Balundu-Bima); ekot (Ejagham); nde-gekwiya 

(Denya); mekan (Badjué): GABON: nkan (Fang); nkanda (Kélé); ikandji (Kota); 

okana (Ndumu); mokangé (Pinji); mokangé (Tsogo); mukanda (Sira); mukanda 

(Duma); mukanda (Lumbu); nkogu (Myene); nkanyi (Seki); CONGO: mukaana a 

nguomo (Téké): DR CONGO: ma-kauw, bo-kauw (def.) (Lingala); bo-nganga 

(Mongo-Nkundu); nkau (Kongo); lu-bubi (Lega-Mwenga): ANGOLA: mi-cau 

(Mbundu-Luanda). 

332

Uses 

This species is the most desired large diameter cane species in West Africa and, along 

with L. robustum, one of the two main commercial species in Central Africa 

(Hédin,1929; Irvine, 1961; Bauchet, 1988; Profizi, 1989; Abbiw, 1990; Tuley, 1995; 

Burkill, 1997; Sunderland, 1999a; 1999b). 

Although the stems are used, in the main, whole, throughout west Africa, the split 

stems are also used for binding house frames together as well as for the construction 

of fish traps, baskets and other woven products (Bauchet, 1988; Profizi, 1989; Abbiw, 

1990; Tuley, 1995; Burkill, 1997). The leaves were formerly used by the Mende of 

Sierra Leone (Dalziel, 1937) the Nzeme of Ghana (Abbiw, 1990) and the Edo of 

Nigeria (Burkill, 1997) as a thatching material, but have since been replaced by metal 

roofing (Abbiw, 1990; Davies and Richards, 1991). In Cote d’Ivoire, the leaves are 

also used for thatching whilst the split stems are used for basketry and in the 

fabrication of traditional drums (securing the goat-skin onto the wooden frame) 

(ECOSYN, 1999). In the Congo, the Téké use the split stems of this species as a 

mooring line for fasting their canoes to the river bank. In much the same way, these 

split stems can also be suspended across a river to help with crossing on foot (Profizi 

and Makita-Madzou, 1996). The plant itself has significant symbolic value and is 

reputed to conjure up ghosts (ibid.). 

The young emerging shoots are often cut and roasted whole before the inner soft pith 

is eaten (Raponda-Walker and Sillans, 1961; Berhaut, 1988; Abbiw, 1990) or they 

may be boiled and then fried in butter to remove the bitterness (Chevalier, 1934). 

However, the Balundu-Bima of Cameroon state that the consumption of the young 

shoots can lead to impotence and hence it avoided by males (Mallson pers. comm.). 

The stem apex is eaten is widely eaten and is thought to contain an active heartstimulant 

which promotes a sense of well-being akin to that caused by kola nuts 

(Jordan, herb. K!). Holland (1922) records how many local people in Nigeria 

transported a portion of the upper stem with them when they entered the forest for 

long periods to harvest wild rubber; the whole stems were roasted on the fire and the 

soft pith then removed and eaten. The cut stems also provide a potable sap drunk by 

forest travellers (Berhaut, 1988; Raponda-Walker and Sillans, 1961). 

333

A tea made from the young shoots is used as a vermifuge by the Akan-Asanti of 

Ghana (Abbiw, 1990) as it is in Gabon (Raponda-Walker and Sillans, 1961). The 

Lombo from the Yangambi region of DR Congo, use the sap from the cut stems of 

this species, mixed with the bark Hua gabonii, Quassia africana and Pterocarpus 

soyauxii, to make a tea which is then used to treat dysentary (Staner and Boutique, 

1937). 

References: SENEGAL: Berhaut (1988); Jordan 2064 (herb. K!): GUINEA-BISSAU: Burkill (1997): 

SIERRA LEONE: (Burkill, 1997): CÔTE D’IVOIRE: (Adanohoun & Ak� Assi, 1972 – cited in 

Burkill (1997)); (Bouquet & Debray, 1974); ECOFAC (1999): GHANA: Sunderland unpubl. notes); 

Dalziel (1937): BENIN: Profizi (1989); Aufsess (herb. K!): NIGERIA: Dalziel (1937); Otedoh 7251 

(herb. K!); Burkhill (1997): CAMEROON: Letouzey 10605 (herb. YA!); Thomas et al., (1989): 

GABON: Raponda-Walker and Sillans (1961): CONGO: Profizi and Makita-Madzou (1996): DR 

CONGO: Leonard 815 (herb. BR!); Minga (in press): ANGOLA: Chevalier (1936) 

_____________________________________ 

Oncocalamus macrospathus Burr. 

Vernacular name 

CAMEROON: eboti (Ewondo) 

Uses 


References: CAMEROON: Letouzey 11889 (herb. YA!) 

_____________________________________ 

O. mannii (H. Wendl.) H. Wendl. 


CAMEROON: mfop n’lon (Bulu); EQUATORIAL GUINEA: asa-nlong (juvenile), 

ndoro (adult) (Fang): CONGO: mituo (Téké) 

334

Uses 

The cane of this species is poor in quality. It is rather inflexible and prone to breaking. 

However, particularly in the absence of other species, O. mannii can be employed for 

coarse weaving. In the Congo, the stem epidermis is used by the Téké for tha 

manufacture of coarse baskets and local weaving. In Equatorial Guinea, the juvenile 

stems are sometimes used for weaving in the same manner as the young stems of E. 

macrocarpa, which they resemble; hence the same name for the juvenile form 

(Sunderland, 1998). 

References: CAMEROON: Hédin (1929); van Gemerden BJ (herb. K!): EQUATORIAL GUINEA: 

Sunderland (1998): CONGO: Profizi & Makita-Madzou (1996). 

_____________________________________ 

O. tuleyi sp. nov. 


NIGERIA: iboh (Ekit): CAMEROON: madame (Trade/Pidgin); mo’ap (Balundu- 

Bima); edju (Bakundu-Balue); moa-echié (Denya) 

Uses 

This cane is not used commercially at all due to the poor quality of the cane 

(Sunderland pers. obs.; Morakinyo, 1994). It is rather weak, inflexible and rather 

prone to breakage on bending. However, the base of the leaf sheath is often used by 

the indigenous communities of SW Cameroon as a chew-stick (Sunderland unpubl. 

notes; Thomas et al., 1989; Sunderland & Tchouto, 1999). In Nigeria, the stem 

epidermis is often used for tying yams to their climbing poles (“yam-barns”) as it rots 

fairly quickly and does not constrict the developing climbing stems (Morakinyo, 

1994). 

References: NIGERIA: Morakinyo, 1002 (herb. K!): CAMEROON: Thomas et al., (1989); Sunderland 

(unpubl. notes). 

_____________________________________ 

335

O. wrightianus Hutch. 


BENIN: hofle (Defi); gbe-dekun (Gun-Gbe) NIGERIA: akwal� (Igbo); pank�r� 

(Yoruba). 

Uses 

The Igbo in Nigeria split the stem to make tying material of different sorts: a coarse 

cordage ekwel� or akwal�; a finer twine ud�, and string or thread, elili (Jeffreys, 

1960). A specialised twine called akwal� anya is made for tying bulky packages 

(ibid.). This also serves for binding hut frames together. The leaf sheath base is also 

used by the Igbo as a chew stick (Dalziel, 1937). 

References: BENIN: Profizi (1986); Aufsess 430 (herb. K!): NIGERIA: Jeffreys (1960); (Dalziel, 

1937); Holland (1922). 

_____________________________________ 

336

Adam: Liberia; 20746, 29807 

APPENDIX TWO 

LIST OF EXSICCATAE 

Adams: Ghana; 2025, 2190, 2195, 2214, 2407 

Ake-Assi: Côte d'Ivoire; 9450 

Allard: DR Congo; 213, 294 

Allison: Nigeria, 6994 

Anderson: Gambia; 131 

Andrews: Sudan; 1291 

Aninze: Cameroon; 24732: Nigeria; 15402 

Apema: DR Congo; 217 

Arends et al.: Gabon; 671 

Arwaodo: Nigeria; 42 

Asonganyi: Cameroon; 279 

Aufsess: Benin: 424, 425, 426, 427, 429, 430 

Ayewoh: Nigeria; 3851, 3853, 3852, 3854 

Baland: DR Congo; 1992 

Barter: Equatorial Guinea; s.n.: Nigeria; 4, 2220, s.n. 

Bavicchi: DR Congo; 277 

Bennett: Nigeria; 8 

Bequaert: DR Congo; 878, 953, 1277, 1411, 1824, 7076, 7895 

Berhaut: Senegal; 877 

Bermejo: DR Congo; 19, 88 

Bernardi: Côte d'Ivoire; 8382 

337

Bidgood & Vollesen: Tanzania; 3040 

Bidgood et al.: Tanzania; 2924 

Bililong & Bullock: Cameroon, 348, 351 

Billiet & Jadin: DR Congo; 4054 

Bos: Cameroon, 4799, 5160, 5162: Liberia: 2165 

Boughey: Côte d'Ivoire; 14732, s.n. 

Brenan: Nigeria; 8580 

Breteler: Cameroon, 1241, 1560 

Breteler & van Raalte: Gabon; 5557 

Breteler et al.: Cameroon; 1200, 2563 

Breteler et al.: Gabon; 10957, 11194 

Breyne: DR Congo; 2105, 2357 

Bruneau: Cameroon; 1116, 1071, 1074, 1093 

Brunt: Cameroon; 137, 207 

Buschen: Cameroon; 3 

Cabra: DR Congo; s.n., s.n. 

Callens: DR Congo; s.n. 

Camp: DR Congo; s.n. 

Carroll: Central African Republic; 33, 115 

Carvalho: Equatorial Guinea; 2212 

Chapin: DR Congo; 613 

Chapman: Nigeria; 3681, 5010, 5202, 5331, 5423 

Cheek: Cameroon; 5062, 5554, 5591 

Chevalier: Côte d'Ivoire; 22658 

Chillou: Guinea-Conakry; 1905, 1419 

338

Chipp: Ghana; 127, 643 

Claessens: DR Congo; 381; 989 

Commonwealth Imperial Institute: Nigeria; s.n. 

Compère: DR Congo; 2181, 2182, 2183 

Corbisier-Baland: DR Congo; 1992 

Couteaux: DR Congo; 471, 472, 473, 502, 1051 

Cummins: Ghana; 128, 229 

Daramola & Adebusuyi: Nigeria; 38415 

Dawe: Uganda; 149, 668 

de Giorgis: DR Congo; 174 

de Graer: DR Congo; 216, 297, 327 

de Wilde: Cameroon; 2183; Côte d'Ivoire; 3101, 3282 

de Wilde & Leeuwenberg: Côte d'Ivoire; 3432 

de Wilde et al.: Gabon; 9301, 9917, 11177 

de Witte: DR Congo; 4066 

Dechamps: DR Congo; s.n. 

Deighton: Sierra Leone; 1847, 2592, 2593, 3090, 4117, 4118, 4119 

Demeuse: DR Congo; s.n., s.n. 

Desenfous: DR Congo; 2023 

Deuse: DR Congo; 121 

Dewevre: DR Congo; 581, 639, 986 

Dewulf: DR Congo; 526 

Dibata & Mbouissou: Gabon; 958 

Dinklage: Cameroon; 1154, 1155 

d'Orey: Guinea-Bissau; 262 

339

Dransfield: Cameroon; 6998, 6999, 7000, 7001, 7002, 7003, 7004, 7005, 7006, 7007, 

7476 

Dubois: DR Congo; 911, 912 

Dundas: Cameroon; 8381 

Dunstan: Nigeria; s.n. 

Dusen: Cameroon; 292 

Dybowski: Gabon; 140 

Eggeling: Uganda; 1517: Tanzania; 6207 

Eneme & Lejoly: Equatorial Guinea; 113 

Enti: Ghana; 614, 643, 758, 1914, 2344 

Enti & Hall: Ghana; s.n. 

Etuge: Cameroon; 1393 

Evrard: DR Congo; 1686, 1876, 2077, 2984, 3145, 3933, 4041, 4511, 4943, 5511, 

5890, 7070 

Faden & Mbama: Cameroon; 86/60 

Fay: Central African Republic; 4021, 4036, 4381, 7018, 7020, 8236, 8254, 8255 

Fay & Harris: Central African Republic; 8795 

FHI: Nigeria; 6996 

Flamigny: DR Congo; 6361 

Foggie: Ghana; 16/40 

Fotius: Cameroon; 3074 

Furtado: Java (Cultivated); A113 

Gartlan: Cameroon; 39 

Gathy: DR Congo; 1639 

Gentry & Emmons: Gabon; 33732 

Gentry & Pilz: Nigeria; 32873 

340

Gentry & Thomas: Cameroon; 52727, 52766 

Gentry et al.: Cameroon; 62566 

Gerard: DR Congo; 1432, 2152, 3218, 3933, 4951 

Germain: DR Congo; 210, 326, 1669, 1681, 4808, 8728 

Gilbert: DR Congo; 2258, 7909 

Gillet: DR Congo; 167, 1385, 2026, 2069, 3505, 3812, s.n. 

Gledhill: Sierra Leone; 309, 339; Nigeria; 923 

Gossweiler: DR Congo; 8705; H1039/24; Angola; 6416, 6645, 7537, 7541, 7567, 

7844, 7995, 8129, 8145, 9092, 10086, 10087, 10088, 13644, s.n. 

Guinea-Lopez: Equatorial Guinea; 913 

Gutzwiller: DR Congo; 539 

Hall: Ghana; 2748, 2846, 3371, 42605, s.n. 

Hall & Abbiw: Ghana; 45124 

Hall & Aké-Assi: Côte d'Ivoire; 45442 

Hallé: Congo; 1814 

Harley: Liberia; 2174 

Harris: Cameroon; 2456, 2471, 3660, 3738, 3739, 3742, 3769, 3778: Central African 

Republic; 2360, 3386, 3419, 3508, 4232, 4765, 4966, 5704, 5706, 5707, 5718, 5719, 

5720, 2652 

Harris & Fay: Central African Republic; 449, 459, 820, 1494 

Harris & Payne: Cameroon; 2469, 2470 

Harris et al: Congo; 3172, 3222 

Hart: DR Congo; 633 

Hendrickx: DR Congo; 4157, s.n. 

Hens: Congo; 170 

Hepper & Maley: Côte d'Ivoire; 8178, 8041, 8062a, 8177, 8214 

Herman: DR Congo; 2138 

341

Heudelot: Gambia; 372 

Hoier: DR Congo; s.n. 

Holman-Bentley: DR Congo; s.n. 

Hulstaert: DR Congo; 747, 864, 869, 1416, 1417, 1418, 1419, 1420, 1421, 1428, 

1614, 1616, 1623, 1624 

Imp. Inst. Nigeria: 346, 347 

Ingram: Gambia; s.n. 

Iquito: Cameroon; 61 

Irvine: Ghana; 502, 2075, 2300, 5067, 4861, 4873, 4886 

Jackson: Sudan; 3406 

Jaeger: Sierra Leone; 1802, 9222 

Jangoux: Côte d'Ivoire; 215 

Jans: DR Congo; 655 

Jansen: Liberia; 1822 

Johnson: Ghana; 242, s.n. 

Jones & Onochie: Nigeria; 17237, 17416 

Jordan: Sierra Leone; 2064, 2065, 2066 

Kalbrayer: Cameroon; 65 

Katende: Uganda; 702, 2783 

Keay: Nigeria; 28091 

Kinlock: Ghana; 3237, 3326 

Kisseadoo: Ghana, 435, 441 

Kitembo: DR Congo; 60 

Klaine: Gabon; 3246, s.n., s.n., s.n. 

Kuasa: DR Congo; 48 

342

Latilo: Nigeria; s.n. 

Laurent: DR Congo; 645, 911, 912, 913, 981, 1118, s.n., s.n., s.n., s.n., s.n., s.n., s.n., 

s.n., s.n., s.n., s.n., s.n., s.n., s.n. 

le Testu: Central African Republic; 3594 

le Testu: Gabon; 1712, 9258, s.n., s.n., s.n. 

Lebrun: DR Congo; 1508, 2985 

Leclercq: DR Congo; 736 

Lederman: Cameroon; 1487, 2428 

Leeuwenberg: Côte d'Ivoire: 2515, 2524, 2882, 3954 

Lejoly: Cameroon; 86/1005: Congo; 96/750: DR Congo; 566, 1438, 1512, 2912, 

82/820 

Leonard: DR Congo; 54, 55, 815, 816, 817, 832, 929, 932, 933, 935, 936, 980, 1138, 

1671, 1686, 3817, 3960 

Lerot: Congo; s.n. 

Letouzey: Cameroon; 3553, 3673, 4151, 4206, 4278, 4285, 4416, 4556, 7368, 8465, 

8479, 10605, 11133, 11518, 11776, 11778, 11794, 11796, 11798, 11800, 11889, 

12477, 12563, 13843, 14522, 14748, 15317 

Letter: Nigeria; 8224 

Lewalle: Burundi; 4016, 4414 

Liben: DR Congo; 2603 

Liegeois: DR Congo; 86, 87, 88, 89 

Linder: Liberia; 676, 1228, 1078, 1116, 1226, 1341 

Lisowski: DR Congo; 7160, 86336: Equatorial Guinea; 1263 

Louis: DR Congo; 218, 772, 847, 1970, 3395, 3638, 3646, 3699, 3804, 3958, 4218, 

5656, 6445, 7671, 7994, 8106, 9420, 9556, 9560, 9731, 10045, 10155, 11439, 11739, 

11850, 12106, 13176, 14729, 15169, 15489, 15541, 15545, 15925, 15944, 16049, 

16340, 16775, 16791, 16794, 16796, 16797, 16995 

Louis et al.: Gabon; 1350 

Loverage: Zambia; 931 

343

Lowe: Nigeria; 2792, 2793, 4353, Cameroon; 3442, 3443 

Luja: DR Congo; 107, 221, 223, 231, 234, 288, 297, 302 

Magajie & Tuley: Nigeria; 2166 

Maggs: Nigeria; 150, 159, 160 

Maitland: Cameroon; 761 

Makombo et al.: Uganda; s.n. 

Malaisse: DR Congo; 9453, 11907, 13889, 14159 

Mandango: DR Congo; 2970, 2977, 3067 

Mann: Sierra Leone; 895, s.n.: Nigeria; 451, 453, 2330; Cameroon; 2147: Equatorial 

Guinea; 97: Gabon; 1043, 1044, 1044a; 1045 

Maruhashi: Cameroon; 171 

Masens: DR Congo; 451 

Mbani: Cameroon; 497 

Meijer: Cameroon; 15220, 15221, 15288, 15251 

Michel: DR Congo; 2957 

Michel & Reed: Burundi; 1362 

Mildbraed: Cameroon; 4190, 5264, 5285, 5310, 5879, 6036, 9546, 9548: DR Congo; 

2818 

Miller: Nigeria; 18 

Milne-Redhead: Angola; 4219 

Moore: Ghana; 2115 

Moore & Enti: Ghana; 9882, 9886, 9887, 9888, 9891, 9892, 9893 

Morakinyo: Nigeria; 1000, 1001, 1002, 1003, 1004, 1005, 1006 

Morton: Ghana; 377, 3618 

Moureau-Cheauvard: DR Congo; 129 

Mpou: Cameroon; 338 

344

Mullenders: DR Congo; 1166 

Mutimushi: Zambia; 3372 

Myers: Sudan; 6757, 11334 

Nannan: DR Congo; 46, 65, 66, 117 

Ndjele: DR Congo; 732 

Niger Company: Nigeria; s.n. 

Njingum: Cameroon; 1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11 

Nkefor: Cameroon; 445, 920 

Nkongmeneck: Cameroon; 596 

Noirfalise: DR Congo; 664 

Nsimundeue: DR Congo; 1055 

Nsola: DR Congo; 621 

Nwambin & Tuley: Nigeria; 603 

Oldeman: Côte d'Ivoire; 137, 571, 589, 674 

Onochie: Nigeria; 5243, 7706 

Onochie & Jones: Nigeria; FHI 17332 

Otedoh & Tuley: Nigeria, 7252, 7253, 7258 

Pauwels: DR Congo; 2322, 4543 

Poulson: Uganda; 969 

Preuss: Cameroon; 460, 1232 

Proctor: Tanzania; 369, 470 

Pyneart: DR Congo; 1073, 1673, 1676 

Pyne: Sierra Leone; 39 

Raynal: Cameroon; 9785, 9825, 10150, 10548 

Reekmans: Burundi; 11180 

345

Reitsma: Gabon; 1340, 2047, 2151, 2856 

Richards: Cameroon; 5209 

Ringoet: DR Congo; s.n. 

Roberty: Guinea; 2903 

Robyns: DR Congo; 3250, 4300 

Rosevear: Cameroon; 30138 

Sapin: DR Congo; s.n., s.n., s.n., s.n., s.n., s.n., s.n., s.n., s.n. 

Schmitz: DR Congo; 3710, 5617 

Schouten: DR Congo; 103 

Schweinfurth: DR Congo; 2860 

Scott-Elliot: Sierra Leone; 4442, 4738, 5121 

Sita: Congo; 4642 

Small: Sierra Leone; 455; 832; 697 

Smith: Nigeria; 53 

Soyeaux: Gabon; 155 

Starin: Gambia; 28, 136 

Sunderland: Ghana; 2259, 2260, 2261, 2262, 2263, 2264, 2265, 2266: Cameroon; 

1640, 1645, 1700, 1701, 1702, 1704, 1705, 1706, 1707, 1708, 1710, 1711, 1712, 

1713, 1714, 1716, 1717, 1718, 1719, 1720, 1721, 1722, 1723, 1730, 1731, 1733, 

1737, 1738, 1739, 1740, 1741, 1742, 1743, 1744, 1746, 1747, 1750, 1752, 1754, 

1755, 1756, 1757, 1758, 1759, 1760, 1761, 1762, 1763, 1764, 1765, 1766, 1767, 

1768, 1769, 1790, 1801, 1802, 1804, 1805, 1806, 1807, 1855, 1856, 1864, 1881, 

1882, 1883, 1885, 1886, 1887, 1888, 1890, 1926, 1927, 1928, 1929, 1930, 1931, 

1933, 1934, 1935, 1936, 1937, 1938, 1939, 1999, 2017, 2021, 2042, 2043, 2048, 

2054, 2055, 2056, 2057, 2058, 2059, 2250, 2251, 2252, 2253, 2254, 2255, 2256, 

2257, 2258, 2301, 2302, 2303, 2304, 2305, 2306, s.n.: Equatorial Guinea; 1791, 1792, 

1793, 1796, 1797, 1798, 1799, 1800, 1871, 1874, 1875, 1876, 1900, 1901, 1902, 

1903, 1904, 1906, 1907, 1908, 1909, 1910, 1911, 1913, 1914, 1915, 1916, 1917, 

1918, 1919, 1920, 1921, 1922, 1923 

Sylvanus: Cameroon; s.n. 

Szafranski: DR Congo; 1578 

346

Terashima: DR Congo; 94 

Tessmann: Cameroon; s.n.: Equatorial Guinea; 1, 2, 4, 6, 44 

Thiebaud: DR Congo; 321 

Thomas: Cameroon; 2292, 5163, 6139, 8182, 9726, 9732, 9733, 9738, 10058, 10059, 

s.n., s.n., s.n. 

Thomas et al: Congo; 8944 

Thomas: Nigeria; 338 

Thomas: Sierra Leone; 2753 

Thonet: DR Congo; 110, 129 

Tomlinson: Ghana; s.n., s.n., s.n., s.n., s.n. 

Toussaint: DR Congo; 2294, 2331 

Troupin: DR Congo; 296; 2658, 9162 

Tsiforkor: Ghana; s.n. 

Tuley: Nigeria; 454, 530, 603, 648, 649, 650, 665, 652, 653, 846, 851, 1076, 1077, 

1078, 

Tuley & Ochie: Nigeria; 1682 

Unwin: Nigeria; 109 

van Gemerden: Cameroonl; 77, 110, Bi, BJ, BL, 

van Meer: Liberia; 264 

van Nek: Gabon; 517 

Vanden-Berghen: Senegal; 1752, 4094, 5264, 7285 

Vandenbrand: DR Congo; 238 

Vanderyst: DR Congo; 1408, 2781, 4906, 5139, 5256, 6411, 9974, 11246, 12343, 

12664, 12670, 21843, 30729 

Vigne: Ghana; 1365, 1829, 1868, 1875, 2410, 3951, 4858 

Watts: Cameroon; 511, 514, 821 

Webb: Cameroon; 311 

347

Webb & Bullock: Cameroon; 310 

Wellens: DR Congo; 473 

West-Skinn: Ghana; 11, 90 

Wheatley: Cameroon; 154 

Whyte: Liberia; s.n. 

Wieringa: Gabon; 466, 1550 

Wilks: Gabon; 1486 

Williamson: Gabon; 128 

Wolfert: Guinea; 1910 

348

APPENDIX 3: 

RATTAN RESEARCH PLOT No. 1: 

CAMPO FAUNAL RESERVE, CAMEROON 

349

APPENDIX FOUR 

SOCIO-ECONOMIC SURVEY OF CAMEROON RATTAN SECTOR 

STANDARD QUESTIONNAIRE 

352 

Page ___ 

Form ID No. LQ_ /_ _ _ Interviewer’s name___________________ 

Date____________ Settlement name____________________ House No._______ 

1. Name of interviewee?____________________________ 

2. Rattan-related activity______________________ 

(Harvests rattan, processes rattan, makes furniture/baskets, 

transports cane, sells cane/baskets/ furniture etc.) 

3. Are there any other people in this household who are engaged in the same work? 

4.If yes to Q31, give name(s) of other people in this household involved in this 

activity 

a.______________________ b.____________________ 

c ______________________ d.______________________ 

5. Why did you (respondent) get involved?__________________ 

(family/friends involved; good market prospects; no alternative source of income; 

needed money; easy to start; other (specify) 

6. How did you get involved?_______________________________ 

(inherited skills from family; started from scratch; apprenticeship, training, other 

(specify) 

7. What are your most important problems with 

this activity? 

8. Can you rank them? 

9. Is any special equipment required for this work? specify______________________ 

10. What are your main expenses for this business?_______________ 

(tools; labour; transport; raw materials; collection fees (official/unofficial); other 

(specify).

11. Do you employ anybody to work in your business? no/yes_______ 

If no � to Q13. 

12. If yes, record the a)number of workers, b)timing and c)type of activity in the table 

below: 

a) Number of workers b) Timing of work (which 

season? No. of days, weeks, 

months?) 

13. How many apprentices do you work with?_______________ 

(possible answers: none, any number) 

22. Is any rattan cane originally obtained from yes/no: 

353 

c) Type of activity 

14.Gender of workers and apprentices? a) How many females are there?_____ 

b) How many males are there?_______ 

15. How many children (under 14) are there involved in this enterprise?____ 

16.What are the types of rattan cane you use?______________________________ 

17. How do you obtain the rattan cane needed for this enterprise?______________ 

(harvests rattan cane; buys rattan cane; hires someone to collect rattan cane; 

other(specify). 

If he or she does not collect his own rattan � to Q22. 

18. Is any of the rattan cane obtained from: (no/yes) 

a) high forest 

b) farm 

c) fallow 

19. Which, if any, of these areas do you harvest most of your cane?______________ 

20. Is any of the rattan cane obtained from: (no/yes) 

a) inside reserved forest 

b) outside reserved forest 

21. Which, if any, of these areas is the most important source of rattan 

cane?______________

a) high forest 

b)farm 

c)fallow 

d) don’t know 

23. How does the availability of rattan cane compare with five years 

ago?___________ 

(more available; no change; don’t know; less available) 

If DK ---> skip to Q27 

24. If more available why?_________________________ 

(more people collecting; easy access to source) 

25. If less available why?__________________________ 

(local people using more; outside people using more; destructive harvesting practices 

(local people); destructive harvesting practices (strangers); fires; forest guards 

restricting access; logging operations damaging resources; logging workers harvesting 

rattan cane; clearance of sources areas for agriculture; invasive weeds; other (specify). 

26. If less available, can you think of any way(s) to improve the supply of cane? 

________________________________________________________ 

27. Do you, or other people you know, know of a way of harvesting cane to maintain 

supplies?______________________________________________________ 

28. If there was more rattan cane available would you use more of it?_________ 

(yes; no; don’t know) 

If DK � skip to Q?? 

29. If no, why?___________________________________________________ 

(possible answer: no time) 

30. During which periods in the year are you operating the business? (tick) 

J F M A M J J A S O N D 

31. Is there any high season for this activity? If yes, tick months of high season in 

table below: 

J F M A M J J A S O N D 

32. Who is your major customer? ____________________________ 

(Local individuals (final consumer); local traders; outside traders; processing 

enterprise; government institutions, schools etc; other (specify)) 

33. How has the volume of your own business changed in the last five years? 

354

________________________ 

(increase; no change; decrease; don’t know). 

34. Are there other people in this area that are engaged in the same activity? 

yes/no____ 

If no � Q38 

35. If yes, where are they located?_________________ 

(businesses in the settlement; business outside the settlement; other). 

36. If yes to Q34, how has the number of competitors changed in the last five years? 

_________________________________ 

(increase; no change; decrease; don’t know) 

If no increase � Q38 

37. If there has been an increase, what has been the cause?________________ 

(local people; people outside; both; other; don’t know) 

38. Do you consider this activity to be one of your main sources of income? 

Yes/no____ 

39. If you could choose between expanding this business or starting another business, 

which would you choose ?_________________ 

(expand this business; start new business; don’t know). 

40.Do you belong to any rattan-related business support group or collecting 

group? yes/no_____ 

41. If yes, name_______________ 

THANK-YOU FOR YOUR ASSISTANCE 

Remember to fill in another supplementary sheet if anyone is involved in another 

type of rattan-related enterprise 

355

APPENDIX FIVE 

PUBLICATIONS AND DISSEMINATION 

The following publications and other dissemination have resulted directly from the 

work presented in this thesis. 

PUBLICATIONS 

1. Sunderland, T.C.H. 1998. The rattans of Rio Muni, Equatorial Guinea: utilisation, 

biology and distribution. A report for the European Union Project No.6 ACP-EG- 

020: Proyecto Conservacion y Utilizacion Racional de los Ecosistemas Forestales 

de Guinea Ecuatorial (CUREF). 

2. Sunderland, T.C.H. 1999a. The rattans of Africa. In: R. Bacilieri & S. Appanah 

(eds.) 1999. Rattan cultivation: Achievements, Problems and Prospects. CIRAD- 

Forêt & FRIM, Malaysia. pp 237-236 

3. Sunderland, T.C.H. 1999b. New research on African rattans: an important nonwood 

forest product from the forests of Central Africa. In: T.C.H. Sunderland, 

L.E. Clark & P. Vantomme (eds). The non-wood forest products of Central Africa: 

current research issues and prospects for conservation and development. Food 

and Agriculture Organisation. Rome. pp 87-98 

4. Chapter 6, along with Appendix 1, is currently in press: Sunderland, T.C.H. [in 

press]. Indigenous nomenclature, classification and utilisation of African rattans. 

In: L. Maffi & T. Carlson (eds.) [in press]. Ethnobotany, and Conservation of 

Biocultural Diversity. Advances in Economic Botany. New York Botanical 

Garden 

5. A version of Chapter 9 is currently in press: Sunderland, T.C.H. & J. Dransfield. 

[in press]. Certification guidelines for rattans. In: P. Shanley, S. Laird, A. Pierce & 

A. Guillen (eds). The Management and Marketing of Non-Timber Forest 

356

Products: Certification as a Tool to Promote Sustainability. RBG 

Kew/WWF/UNESCO People and Plants Series no. 5. 

6. Chapter 8 has been submitted to the Overseas Development Institute’s (ODI) 

Occasional Paper Series and is currently under review. 

7. Chapters 1 & 2 have been combined and submitted to the Kew Bulletin under the 

following heading: A revision of the rattans of Africa. The paper is currently under 

review. 

8. Sunderland, T.C.H., M. Balinga & M.A. Dione. [in press]. A socio-economic 

profile of the rattan sector of Equatorial Guinea. In: B. Belcher & O. Ndoye (eds). 

NTFP case comparisons: Africa. CIFOR, Bogor, Indonesia. 

9. In June 2000, the Kew Publications Committee approved the publication of this 

thesis as part of the Scientific Publication Series. The manuscript is currently 

undergoing review and editing to comply to the Kew publishing framework, and 

will be published as: The rattans of Africa: taxonomy, ecology and utilisation. 

10. The proceedings of the International Expert workshop on African rattans is 

currently in press: New research on African rattans edited by T.C.H. Sunderland 

& J.P. Profizi. INBAR. Beijing. This proceedings includes the following authored 

papers. 

r Sunderland, T.C.H. The rattans of Africa: an overview 

r Sunderland, T.C.H., & J.P. Nkefor. Technology transfer between Asia and 

Africa: Rattan cultivation and processing 

r Sunderland, T.C.H. & J. Dransfield. The sustainability of rattan exploitation 

357

DISSEMINATION 

1. Sunderland, T.C.H. 1998a. The rattan palms of Central Africa and their economic 

Importance. Paper presented to the Society of Economic Botany meeting “Plants 

for people”, Aarhus, Denmark, 13 th -17 th July. 

2. Sunderland, T.C.H. 1998b. The Rattans of Africa: An Overview. Paper presented 

to the “Expert consultation for the development of bamboo and rattan in Africa”. 

Johannesburg, South Africa, 20 th -23 rd September. 

3. Sunderland, T.C.H. 1998c. Recent research in African rattans. Paper presented to 

the CIRAD-Forêt international meeting “Rattan cultivation: achievements, 

problems and prospects”. Kuala Lumpur, 12 th - 14 th May. 

4. Sunderland, T.C.H. 1998d. Recent research of African rattans: an important nonwood 

forest products from the forests of Central Africa. Paper presented to the 

CARPE/FAO international expert meeting on the NWFP’s of Central Africa. 

Limbe Cameroon, 10-14 th May. 

5. Sunderland, T.C.H. 2000a. Technology transfer between Asia and Africa: rattan 

processing and transformation. Paper presented to the CARPE international NTFP 

workshop, Limbe Cameroon, 16-18 th July. 

6. Sunderland, T.C.H. 2000b. The rattans of Africa: biology, utilisation and 

prospects for sustainability. Paper presented to the INBAR/FAO meeting 

“International expert consultation on rattans”. Rome, Italy, 4 th -7 th December. 

7. Sunderland, T.C.H. 2000c. A new taxonomy of the African rattans: implications 

for development. Poster presented to the XVIIth AETFAT meeting, Brussels, 

Belgium 1-3 rd September. 

358

8. Sunderland, T.C.H. 2000d. The rattans of Africa: an overview. Paper presented to 

the FAO/INBAR International Expert Consultation on Rattan. Rome, Italy, 4-6 

December. 

9. Much of this work presented in this thesis has also been disseminated through the 

following website: africanrattanresearch@fsnet.co.uk 

359

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