Biodiversity of West African Forests:

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Biodiversity of West African Forests An Ecological Atlas of Woody Plant Species Edited by: L. Poorter, F. Bongers, F.Y.N’. Kouamé, W.D. Hawthorne

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CABI Publishing is a division of CAB International CABI Publishing CAB International Wallingford Oxon, OX10 8DE UK Tel: +44 (0)1491 832111 Fax: +44 (0)1491 833508 E-mail: [email protected] Website: www.cabi-publishing.org

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©CAB International 2004. All rights reserved. No part of this publication may be reproduced in any form or by any means, electronically, mechanically, by photocopying, recording or otherwise, without the prior permission of the copyright owners.

Library of Congress Cataloging-in-publication Data A cataloque record for this book is available from the British Library, London, UK Biodiversity of West African forests: an ecological atlas of woody plant species/ edited by L. Poorter, F. Bongers, F.N’. Kouamé, W.D. Hawthorne p. cm Includes bibliographical references and index. ISBN 0-85199-734-1 (alk.paper) 1. Forest plants–Africa, West. 2. Plant diversity–Africa, West. 3. Plant diversity conservation–Africa, West. 4. Forest conservation–Africa, West. I. Poorter, L. (Lourens) QK393.B56 2004 333.95’3’0966–dc22 2003055534

ISBN 0 85199 734 1 Printed and bound in Singapore by MRM Graphics from copy supplied by the editors.

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Biodiversity of

West African Forests An Ecological Atlas of Woody Plant Species Edited by L. Poorter F. Bongers F.N’. Kouamé W.D. Hawthorne

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Table of

Contents Preface and acknowledgements

A

1

Forest

1 2 3 4 5 6

West African forests: introduction

7

What explains the distribution of rare and endemic West African plants?

8

Implications for conservation and management

L. Poorter, F. Bongers and R.H.M.J. Lemmens

Forest cover changes in Côte d’Ivoire and Upper Guinea C. Chatelain, H. Dao, L. Gautier and R. Spichiger

The forest-savanna transition in West Africa L. Gautier and R. Spichiger

The forests of Upper Guinea: gradients in large species composition

5

15

33

41

F. Bongers, L. Poorter and W.D. Hawthorne

Floristic diversity of closed forests in Côte d’Ivoire F. N’. Kouamé, K.E. Kouadio, K. Kouassi and L. Poorter

Biodiversity hotspots in West Africa; patterns and causes J.J. Wieringa and L. Poorter

M. Holmgren, L. Poorter and A. Siepel

F. Bongers, L. Poorter, V. Beligné, W.D. Hawthorne, F. N’. Kouamé, M.P.E. Parren and D. Traoré

53

61

73

87

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B Species 9 10 11

Ecological profiles of rare and endemic species M. Holmgren, L. Poorter, A. Siepel, F. Bongers, M. Buitelaar, C. Chatelain, L. Gautier, W.D. Hawthorne, A.T.F. Helmink, C.C.H. Jongkind, H.J. Os-Breijer, J.J. Wieringa and A.R. van Zoest

Ecological profiles of large timber species A. Siepel, L. Poorter and W.D. Hawthorne

Checklist of Upper Guinea forest species

101

391

447

C.C.H. Jongkind

C Appendices Appendix 1

Colour figures Chapter 2: Forest cover changes in Côte d’Ivoire and Upper Guinea

481

Appendix 2

Forest sites in Upper Guinea

493

Appendix 3

Forest reserves in Upper Guinea

499

Appendix 4

Hotspots of plant biodiversity in Upper Guinean forests

501

Appendix 5

Geographical map of West Africa

503

D Sources & index Literature

507

Authors, photographers and illustrators

517

Index

519

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Preface and acknowledgements

Forests

in West Africa are seriously under threat. Deforestation is continuously going on, and many forest areas have been transformed into agricultural fields. Remaining forests are degrading by poaching, illegal cutting and agricultural planting. This threat has led the European Community to support the ECOSYN project, in which an international group of researchers aimed at compiling available information on the forests of West Africa and on the plants in those forests (project B7-5041/95-02/VIII: ECOSYN: Guides, BIOGIS et Eco-Atlas des arbres et des lianes pour mieux gérer et conserver les forêts d’Afrique Occidentale (de la Guinée au Ghana). The project started in 1996 and was jointly prepared by Renaat van Rompaey, Frans Breteler (Wageningen University), Dossahua Traoré (University of Cocody, Abidjan) and Vincent Beligné (Yamoussoukro). The European Community, Directorate General 8, financed the ECOSYN project. Special thanks to Enrico Pironio for continuous support during the first years, and Matthieu Bousquet for the last period. The project was headed in The Netherlands by Frans Breteler, Roel Lemmens and Frans Bongers, each in different periods. In Côte d’Ivoire Vincent Beligné and Dossahua Traoré guided the project. The book you are reading now is one of the products of this ECOSYN project. It focuses on the distribution and state of the forests and analyses vegetation gradients, determines hotspots of plant diversity, and treats distribution patterns of rare and endemic species. Numerous people have contributed to this book, by providing logistic support, collecting data, data entry, providing photographic material, drawings, commenting on species descriptions and chapters, reviewing, and so on. We wish to thank the following colleagues. Logistic support was provided in Wageningen and Abidjan by Folkert Aleva, Jean Assi, Rob Allaart, Laurent Aké-Assi, Aman Kadio, Vincent Beligné, Frans Breteler, Erik Frederiks, Theo van Hoksbergen, Joke Jansen, Roel Lemmens, Kees-Jan Manschot, Judith van Medenbach de Rooy, Artine van Pouderoijen, Renaat van Rompaey, Barbara van Roosmalen, Tine Ruijsch, Dossahua Traoré, Marina Wassink.

The atlas strongly draws on the use of herbarium collections. Carel Jongkind, Jan Wieringa and Folkert Aleva were responsible for the collection of new specimens in the field, species (re)identification and herbarium mounting & maintenance. Several herbaria in Africa and Europe allowed us to use their herbarium collections; Nationaal Herbarium Nederland, Wageningen branch (Herbarium Vadense, The Netherlands), National Botanical Garden of Belgium (Meise, Belgium), Daubeny Herbarium (Oxford, Great Britain), Royal Botanic Gardens Kew (Kew, Great Britain), Conservatoire et Jardin Botaniques de la Ville de Genève (Genève, Switzerland), Muséum National d’Histoire Naturelle (Paris, France), Herbier National de Côte d’Ivoire, University of Cocody (Abidjan, Côte d’Ivoire), and the Dept. of Botany of the University of Ghana (Legon, Ghana). Data entry was done by Marjo Buitelaar, Stuart Cable, Patrick Ekpe, Hendrikjan van Os Breijer, Jan van Veldhuizen and many students, especially Paul van Esch, Tom van Lokven, Marieke Sandker, Arjan Schoonhoven, Petra Wilbrink and Saskia Woudenberg. Xander van de Burgt and Marieke van Bergen contributed to the selection of rare and endemic species. Denis Filer designed the Brahms herbarium database and tailored it to our specific needs. Several colleagues kindly allowed us to use their databases; Petra De Block (Ixora), Cyrille Chatelain (part of the Genève database), Carel Jongkind (Combretaceae), Marc Pignal (part of the Paris database), Roger Polhill (Loranthaceae), Bonaventure Sonké (Oxyanthus), Piet Stoffelen (Pausinystalia and Corynanthe), Jan Wieringa (Aphanocalyx and Tetraberlinia). GIS support and species distribution maps were provided by Toon Helmink, Gerbert Roerink, Cees de Zeeuw, and Roland van Zoest. Advice on statistics was provided by Cajo ter Braak and on soils by Vincent van Engelen (ISRIC). The species descriptions were refined and checked by Jan-Just Bos (Dracaenaceae), Frans Breteler (Dichapetalaceae), Joost van der Burg (Orchidaceae), François Kouamé, Arnold Pieterse (Podostemaceae), Marc Sosef (Begoniaceae and a number of other species), Laurens Vogelezang helped with editing for some species groups. Species drawings were made by Rosemary Wise, Marjolein Spitteler and Emmelien Jaggar, and photos were kindly provided by William Hawthorne, Herbarium Vadense, and others.

1

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The following authors contributed to the atlas chapters (in alphabetical order): Frans Bongers, Marjo Buitelaar, Cyrille Chatelain, Hy Dao, Laurent Gautier, William Hawthorne, Toon Helmink, Milena Holmgren, Carel Jongkind, K.E. Kouadio, K. Kouassi, François Kouamé, Roel Lemmens, Hendrikjan van Os Breijer, Lourens Poorter, Rodolphe Spichiger, Almira Siepel, Jan Wieringa, Roland van Zoest. David Dunn and Andrea Falke translated some manuscripts into English. The atlas chapters benefited from detailed reviews by (alphabetically) Frans Bongers, Cyrille Chatelain, James Fairhead, William Hawthorne, Milena Holmgren, Sally

Horn, Michael Huston, Carel Jongkind, François Kouamé, Melissa Leach, Adrian Newton, Marc Parren, Lourens Poorter, Stefan Porembski, Hans ter Steege, Mike Swaine, and Claudius van den Vijver. Finally, the beautiful design and layout of the atlas was in the hands of Marjolein de Vette.

Additional acknowledgements for specific chapters may be found in those chapters. We thank all for their contribution to this book.

Lourens Poorter, Frans Bongers, François Kouamé, William Hawthorne

September 2003

The contents of this publication is the sole responsibility of the authors and can in no way be taken to reflect the views of the European Union.

2

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A

Forest Chapters 1-8

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1

L. Poorter, F. Bongers and R.H.M.J. Lemmens

West African forests: introduction

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H A P T E R

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C

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Upper Guinean forests

The rainforests of West Africa have been earmarked as one of the world’s hotspots of biodiversity (Myers et al. 2000). These forests extend from Senegal to Togo, and are referred to as the Upper Guinean forests. These are separated from the rest of the African rainforests by the Dahomey gap: an extension of the woodland savanna of the Sahel to the Gulf of Guinea (Figure 1.2). Because of its isolated position the Upper Guinean forest zone harbours a large number of endemic animal and plant species (Hall & Swaine 1981, Brooks et al. 2001). The Upper Guinean forests are disappearing rapidly. For an effective conservation policy, information is needed on the distribution of rare and endemic species in the region, and on the regions in which they are concentrated (Conservation International 2001). Little of the original forest area is left, and most of the large remaining forest blocks straddle national borders. Only transnational conservation efforts may warrant therefore a successful management of the remaining forest resources, and a regional approach is urgently needed (Conservation International 2001). This book focuses on the biodiversity and ecology of West African forests. It analyses the major ecological factors that give rise to biodiversity, and structure tropical plant communities. Additionally it contains an atlas with ecological profiles of the most important species in view of their conservation value or commercial value. This introductory chapter provides background information of the region. First the location, uniqueness and threats of the area will be indicated and conservation need highlighted. The biogeographical environment of the area will be treated in detail, giving thorough background for the introduction of the forest vegetation and its dynamics. The chapter will end with an outline of the different book chapters.

A distinct biogeographic region The rainforests of West Africa are known as Upper Guinean forests. The origin of the word dates back to the

Figure 1.1 The faith of the Upper Guinean dwellers and their forests is closely intertwined. The photo shows a small boy in a Krou village near Para, southwest Côte d’Ivoire.

Moroccan Berbers, who referred to the lands south of the Sahara as “Akal n-Iguinawen”, or “Land of the Blacks”. This was subsequently modified by early Portuguese travellers to “Guinea” (Martin 1991). Lebrun (1947) was the first one to coin the African tropical forests as the “Région guinéenne”, later renamed by Monod (1957) as “Région guinea-congolaise”. The Guinea Congolian region is a distinct phytogeographic region with a relatively homogeneous flora (White 1979). White defined a phytogeographic region as a regional centre of endemism, when it has more than 50% of its species confined to it, and when it has more than 1000 endemic species. Within the Guinea Congolian region there are tree subcentres of endemism: the Upper Guinea subcentre (corresponding to the Domaine libéro-ivoréen of Aubréville, 1962), the Lower Guinea subcentre, and the Congolian subcentre (Figure 1.2). Upper Guinea is separated from Lower Guinea by the Dahomey gap, a savanna vegetation with few endemic forest species. The Upper Guinean forests straddle nine different countries aligned along the coast: Senegal, Gambia, Guinea Bissau, Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana and Togo (Figure 2.2). Appendix 5 contains a detailed map of the region. Mangrove forest is found as far north as Gambia, near the mouth of the

5

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Gambia river. Another large mangrove area is found south of the Casamence river, along the coastline of Senegal and Guinea Bissau, up to western Guinea. The rainforest belt starts in eastern Sierra Leone, and stretches along the coast to Liberia, Côte d’Ivoire, and Ghana. This rainforest belt is up to 350 km wide, but becomes as narrow as 100 km in Côte d’Ivoire, where a V-shaped savanna intrusion approaches the coast. In Ghana the rainforest finally gives way to savanna. The savanna area is 300 km wide, and covers the coastline of eastern Ghana, Togo and Benin. Here, forests are only found at higher altitudes, on the Togo plateau, and as small islands on the plains. Incidentally, forests penetrate into the savanna, as gallery forests along river courses. The tropical forests of Africa cover c. 2.8 million km2 (Beentje et al. 1994), of which 109,000 km2 belongs to Upper Guinea (1992 data from Sayer et al. 1992). In Upper Guinea, 99,200 km2 is lowland forest, 8700 km2 mangrove forest, 450 km2 swamp forest and 480 km2 montane forest (Sayer et al. 1992, Table 1.1). It is estimated that the African tropical forests contain 12,000 forest plant species, of which 6400 species are endemic (Beentje et al. 1994). The Upper Guinean forests contain 2800 vascular forest plant species, of which 650 species (c. 23%) are endemic, and c. 400 species are considered to be rare (Jongkind, chapter 11). Families with relatively many endemic species are Acanthaceae, Anacardiaceae, Euphorbiaceae, LeguminosaeCaesalpinoideae, Rhizophoraceae, Rubiaceae, Sapindaceae and Zingiberaceae. A few monotypic genera are endemic to the Upper Guinean forests, for example Dinklageodoxa (Bignoniaceae), Habropetalum and Triphyophyllum (Dioncophyllaceae), Apodiscus (Euphorbiaceae) and Polystemonanthus (Leguminosae-Caesalpinoideae) (Jongkind, chapter 11).

Figures for animal species are lacking. Myers et al. (2000) use a wider definition of West Africa, including the forests from Guinea to western Cameroon. This area harbours 514 bird, 551 mammal, 139 reptile and 116 amphibian species, of which respectively 90, 45, 46 and 89 species are endemic.

Deforestation, threats, and conservation Upper Guinea harbours many endemic plant and animal species that are threatened by deforestation, habitat fragmentation, and over-exploitation. In the area from Sierra Leone to Togo between 100,000 and 470,000 km2 of forest has been lost since the turn of the 19th century (Sayer et al. 1992, Fairhead & Leach 1998). Animals might have sought refuge in neighbouring forest areas, but sessile plant species with narrow distribution ranges might have disappeared along with the forest. Many commercial tree species have been exploited to such extent that they have become threatened (Hawthorne 1996). Commercial bushmeat exploitation prospers in most countries, leading to the extinction of some animal species (Oates et al. 2000), or reducing the populations of other animals to such low levels that they may go extinct in the coming decades (Holbech 1998, Caspary et al. 2001). Conservation of the remaining forest habitats therefore deserves ample attention, both at local, regional and international level. The high number of rare and endemic plant species are extremely important in this respect. This high number is partly due to the variety in environments in the area, including topographical variation, a range of soil types, and sometimes very steep climatic gradient.

Table 1.1 Estimated cover of different forest types for nine countries in Upper Guinea (period 1985-1990, data from Sayer et al. 1992). Although there are more recent estimates of forest cover (FAO 2001, Matthews 2001), we prefer not to present those data here. The FAO defines forest as land with more than 10% tree cover of trees more than 5 m tall. Not only closed forest, but also savanna woodland is therefore included in their estimates of forest cover. lowland (km2)

montane (km2)

mangrove (km2)

swamp (km2)

Total (km2)

(%)

Senegal Gambia Guinea Bissau Guinea Sierra Leone Liberia Côte d’Ivoire Ghana Togo

192 0 5368 4482 3925 41177 26890 15839 1360

0 0 0 210 81 55 138 0 0

1853 497 2360 2963 1015 6 29 0 0

0 0 0 0 43 0 407 3 0

2045 497 7728 7655 5064 41238 27464 15842 1360

1.9 0.5 7.1 7.0 4.6 37.7 25.1 14.5 1.2

Total (km2)

99233

484

8723

453

108893

Country

Total (%)

6

90.8

0.4

8.0

0.4

100

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Rivers and lakes The Upper Guinean countries are bounded to the west and south by the Atlantic Ocean. The transition between land and sea is often formed by estuaries and lagoon systems (Kogon in Guinea, Ebrié and Aby in Côte d’Ivoire, and Keta in Ghana). The main water bodies on the continent are formed by lakes Buyo and Kossou in Côte d’Ivoire and lake Volta in Ghana. At the western side of Upper Guinea the rivers run east-west (Gambia, Casamence), whereas in the centre and east the rivers run predominantly north-south. Apart from the Gambia and Casamence, the main rivers are the Konkouré in Guinea, Scarcies, Moa and Mano in Sierra Leone, Lofa, St. Paul, St. John and Cestos in Liberia, Cavally, Sassandra, Comoé in Côte d’Ivoire, and Tano and Volta in Ghana (Appendix 5).

Topography In Senegal, Gambia, and Guinea Bissau the topography is relatively flat. In the other countries a narrow plane coastal strip is followed by a belt of rolling hills, extending from the coast inland, followed by a belt of dissected table lands, varying from 300-600 m in altitude (Figure 1.3A). Further inland the Guinean backbone is formed by an interrupted mountain belt, formed by the Fouta Djalon (1540 m) in Guinea, the Loma (1950 m) and Tingi (1720 m) mountains in Sierra Leone, Mt Ziama (1390 m) in Guinea, Mt Wutivi (1340 m) in Liberia, Mt Nimba (1750 m) at the border between Guinea, Liberia and Côte d’Ivoire, and Tonkui (1190 m) in Côte d’Ivoire. Smaller outliers are formed by the Akwapim and Atewa (750 m) ranges in Ghana and the Togo highlands (300-900 m). Locally, granitic outcrops (inselbergs) occur in an otherwise flat landscape. Many of these inselbergs are only up to 100 m tall, but are nevertheless characterised by a typical vegetation (Swaine et al. 1990, Porembski et al. 1994).

Climate The climate in Upper Guinea is determined by the seasonal movement of the sun, the interplay between maritime and continental winds, and the position of the coastline. When the sun is overhead the irradiance leads to

Figure 1.2. Map of continental Africa, with potential distribution of tropical rain forest in West and Central Africa. Three phytochorical regions: Upper Guinea Lower Guinea and Congo, centres of endemism.

a local uplift of the air, resulting in low pressure near the earth surface, giving rise to maritime and coastal winds. The area where these two winds merge is called the intertropical convergence zone (ITCZ). In December, the ITCZ is situated just behind the coast, between 5 and 7 degrees North. During the first part of the year the sun moves north, attaining its northernmost position at 23.5 degrees. The movement of the sun is accompanied by cool and moist maritime winds from the south, and heavy rainfall. Due to the rotation of the earth, these southern winds are deflected to the east, giving rise to southwestern monsoon winds. During the second half of the year the sun moves to the south, sparking off the dry desert winds from the north. Owing to the rotation of the earth, these winds deflect to the west, thus becoming northeastern winds. These desert winds contain reddish dust particles from the Sahara, and are locally known as the Harmattan. The magnitude and seasonal pattern of rain are to a large extent affected by the shape of the coastline. The moisture-laden monsoon winds hit under a perpendicular angle the Guinean, Sierra Leonean and Liberian coast, giving rise to an annual high rainfall of > 4000 mm/yr (Figure 1.3B). The rainfall decreases rapidly in a northeastern direction, declining 800 mm over a distance as short as 30 km (Voorhoeve 1965). Towards the interior, this rapid decrease is reduced, and the isohyets are spaced further apart towards the interior, attaining 1200 mm at the forest savanna boundary. In mountainous areas such as the Guinean backbone, the rainfall may increase again, due to orographic uplift.

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Altitude (m) 0-100 100-200 200-500 500-1000

A

1000-1500 1500-2000

Rainfall mm/yr 0-1000 1000-1250 1250-1500

2500-3000

1500-1750

3000-3500

1750-2000

> 3500

2000-2500

Rainfall stations

B

Soil class cmk < 4 and ph < 5.5 cmk 4-8 and ph ≥ 5.5 cmk > 8 and ph ≥ 5.5

C

At Cape Palmas, at the Liberian-Ivorian border, the coast deflects towards the northeast. The western part of Côte d’Ivoire is therefore situated in the rain shadow of Liberia, and receives substantially less rain. Only beyond Abidjan, the coast again assumes a southeast direction, at perpendicular angles with the monsoon winds. Consequently rainfall levels increase again up to 2000 mm at the coast. At Cape Three Points (Ghana), the coastline turns northeast again, leading to a decrease in rainfall. This pattern is reinforced by a cold seawater upwelling in front of the Ghanaian coast, giving rise to reduced surface air temperatures, less convective uplift, and less rainfall (Hayward & Oguntoyinbo 1987). Accordingly the rainfall drops from 2000 mm at Cape Three Points to 1200 mm in Takoradi, and to 900 mm at Accra, at the border of the Dahomey gap. Most of the forests in the eastern part of Upper Guinea receive an annual rainfall between 1600 and 2000 mm. Rainfall is strongly seasonal, with a long dry season from December to March when the sun is in the southern hemisphere. In some places there is a small dry season from July to August when the winds change direction from southwest to the west. In that case the winds come from the continent, rather than the sea, thus producing a rain shadow effect (Ojo 1977). Another reason is that during this period the ITCZ and its accompanying rains is positioned furthest north. Sites situated closest to the equator experience therefore two small dry seasons, whereas sites further to the north have one long dry season. During the rainy season cloud formation prevents irradiance to reach the earth surface. The average temperature and daily variation therein increase during the dry season because of the clear skies. The monthly average temperature may therefore vary from 24-28ºC. The relative humidity shows a seasonal pattern, being 70% during the dry season, and over 90% during the wet season.

Cation availability (cmol cations/kg soil) 0-1

4-6

1-2

6-8

2-4

8-50

D Geology and soils

Water Holding Capacity (mm water/m soil) 10-50 51-85

E

86-120

Figure 1.3 Environmental maps, focusing on the high forest zone of Upper Guinea. (A) altitude, (B) rainfall, (C) soil types, (D) cation availability, Ca2+, Mg2+, K+, (E) water holding capacity. The dots on the rainfall map indicate the location of the weather stations.

8

Granites, gneisses, quartz and schists form the main parental bedrock from which rainforest soils develop. Approximately two-thirds of Sierra Leone is granite, with scattered strips of metamorphic rocks (Kambui schists) which occur as steep hilly outcrops (Savill & Fox 1967). The bedrock of Liberia is of Precambrian age and made up of igneous and metamorphic rocks such as granitic gneisses, sandstones and schists (Voorhoeve 1965). This bedrock comes to the surface in river beds and on steep hills. The soils that cover the major part of Liberia are Ferralsols, also called Latosols or lateritic soils. In some areas, often in hilly and rugged land, Lithosols are also

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Figure 1.4 NDVI (Normalized Difference Vegetation Index) image of West Africa. The composite image shows the mean leaf area index (m2 leaf area per m2 soil surface) and the seasonal variation therein (expressed as the amplitude and the periodicity). The composite uses the average NDVI, the yearly amplitude and the half-yearly amplitude. Areas around the equator have a high average leaf area index and two growing seasons, more to the north the savannas have a high average leaf area index and strong yearly amplitude. The dark area in the north indicates no vegetation, desert. The picture is the result of Fourier timeseries analysis using the HANTS program. (Roerink et al. 2000, Roerink & Menenti 2000). 36 ten-day NDVI composed pictures are used from 1995 (Figure provided by G.J. Roerink).

found, while in valley bottoms most soils are of alluvial type. The western part of the forest zone in Côte d’Ivoire consists mainly of granite and migmatite, with intrusions of metamorphic schists near Taï National Parc (Avenard 1971). The eastern part of Côte d’Ivoire consists of schists and these underlie also most of the western and southern parts of Ghana (Hall & Swaine 1976). A band near the coast consists of Tertiary sandy sediments, whereas an even smaller strip of Quaternary sands separates the lagoons from the ocean. The different soil types of the region are displayed in the FAO’s Soil Map of the World - Africa sheet (Figure 1.3C). The coastal strip features a variety of soil types, viz. Fluvisols (Sierra Leone), Regosols (Sierra Leone and Liberia), and Arenosols (Côte d’Ivoire and Ghana). The wet forest areas (Sierra Leone, Liberia, and near the Ivorian and Ghanaian coast) are characterised by Ferralsols. The moderately wet areas further inland are characterised by Acrisols, while in Côte d’Ivoire some inclusions of Cambisols are found. The highland areas are distinguished by their Lithosols. The coastal strip, running from Sierra Leone to

southwest Ghana, contains Gleysols. These soils contain a mixture of clay and sands, but are not free draining. The wet forest areas near the coast contain Acrisols or Ferralsols. These are yellowish or reddish of colour, and contain kaolinitic clay minerals with iron and aluminium oxides (Martin 1991). The high rainfall has leached the soil to such extent that available nutrients are much reduced. The moderately wet areas further inland contain Acrisols of a reddish yellow colour. They are less acidic and richer in nutrients. In southeast Ghana, Vertisols are found. The cation availability of most of these strongly leached soils is low (0-2 cmol/kg), with the exception of a few higher elevation areas with younger soils, such as the Putu Range and Mt Nimba range (Figure 1.3D). The water holding capacity ranges between 10 and 120 mm water per m soil, with southeast Ghana having relatively high values, Liberia having low values and Côte d’Ivoire having a mixture of these (Figure 1.3E).

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Vegetation

A

B

C Figure 1.5 Three different forest types in Ghana: (A) Moist Evergreen forest in Subri Forest Reserve. The big tree is Lophira lanceolata (Ochnaceae). (B) Moist Semi-deciduous forest on the University of Ghana Agricultural Research Station at Kade. The big tree is Antiaris toxicaria (Moraceae). (C) Southeast Outlier dry forest on the Shai Hills Game Production Reserve. This dry forest type receives less than 1000 mm rainfall per year, is short statured (c.10 m) and contains only few species.

10

In West Africa there is a strong rainfall gradient from the coast to the Sahel. This has led to a distinct zonation of the vegetation, which can easily be seen from satellite images. Figure 1.4 shows a composite of 10-days NDVI images (Normalized Difference Vegetation Index), that shows the reflectance of the vegetation. Forest, savanna, Sahel and Sahara can easily be distinguished on the image. Along the precipitation gradient, the vegetation changes from wet evergreen, to moist evergreen, moist semi-deciduous, and dry semi-deciduous forest (Guillaumet & Adjanohoun 1971, Hall & Swaine 1981). Wet evergreen forest occurs in areas > 1750 mm annual rainfall, has an average canopy height of 30 m, and contains few deciduous species. Moist evergreen forests occur between 1500-1750 mm rainfall, the canopy is taller (40 m) but the forest contains fewer species than wet evergreen forest (Figure 1.5A). Moist semi-deciduous forest is found between 1250-1750 mm rainfall. It is the tallest forest type (50 m), and the upper canopy is composed of both evergreen and deciduous species (Figure 1.5B). Very few species are confined to this forest type, but it has the highest density of commercial tree species. The dry semideciduous forest occurs between 1250 and 1500 mm annual rainfall, and is adjacent to the savanna zone. The canopy reaches 30-45 m height, is no longer fully closed and contains many deciduous species. In Ghana two distinct forest types are found (southern marginal and southeastern outlier) in regions with less than 1250 mm rain. The southern marginal forest is shorter than 30 m, has a thick undergrowth, and may include high densities of gregarious species. The southeastern outlier forest is short (< 15 m), has only two strata, a low diversity, and is composed of few species (Figure 1.5C). In Liberia wetter forest types exist. Within the mixed forest there are monodominant stands, dominated by single species such as Cynometra ananta, Gilbertiodendron preusii, Tetraberlinia tubmaniana, Parinari excelsa (Voorhoeve 1965) and Talbotiella gentii (only found in the drier forests of Ghana Swaine & Hall 1981). As elsewhere in Africa, most of these single-dominant species belong to the Caesalpiniaceae. Apart from the zonal vegetation types, that respond to the main climatic gradient, there are azonal vegetation types that occur where local site conditions are overruling. In cloudy areas, upland evergreen forest, or montane forest may occur. In Ghana these forests are found above 500 m altitude, whereas in the other countries these are found at considerably higher altitudes (> 1000 m). These forests may become dominated by Parinari excelsa, but most of the characteristic species are herbaceous rather than woody. Less than 5% of the species is deciduous, and many

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Figure 1.7 Summary palynological diagram showing the major palynomorph groups in offshore Niger Delta boreholes spanning the last 10.5 million years. (from Morley 2000, reproduced with courtesy of John Wiley and Sons).

Figure 1.6 Summary of the main palaeoclimatic and palynological trends from 10.5 Ma to 0.5 Ma for a 7000 m core taken in the offshore Niger Delta (from Morley 2000, reproduced with courtesy of John Wiley and Sons). The palynological diagram, excluding mangrove pollen, shows the major freshwater palynomorph groups in the boreholes. The composite profile covers Miocene (M), Pliocene (P) and Quaternary (Q).

epiphytes are found (Hall & Swaine 1981). Mangrove forests are found in tidal and silty areas in lagoons, and along the rivers, up to 100 km inland (Jones 1992). The mangrove forests are up to 20 m tall, and dominated by Rhizophora species with stilt roots, or Avicennia africana with pneumatophores. Littoral woodland is found in the coastal belt, where the soil consists of nearly pure sand. These areas are flooded in the rainy period, but very dry in the dry period, leading to a kind of open savanna vegetation. Swamp forests are especially extensive in Sierra Leone and Côte d’Ivoire. They are subjected to periodic inundation. They have one tree layer, a well developed herb layer, and many large-leaved species. The most common species are Mitragyna spp. and Symphonia globulifera. Raphia palms replace the trees when the inundation period is longer, or when the swamp becomes permanently waterlogged. Gallery forests are found along the riverbanks. The high moisture availability allows them to penetrate deep into the savanna zone.

Vegetation in the past One peculiarity of the human mind is that we tend to believe that what we see has always been there. For the African forests nothing is less true, as they have waxed and waned with climatic fluctuations. 25-10 million years ago, rainforests were very widespread in Africa, with tropical forests occurring as far south as the Cape (Morley 2000). A 7 km core taken from deep sea sediments off the coast of Nigeria shows how the vegetation and climate have changed during the past 10 million years (Figure 1.6 and 1.7). 10 million years ago, Graminae pollen increased in abundance, reflecting an expansion of the savanna in the Niger catchment, and a parallel reduction in rainforest. Temporal changes in the relative abundance of savanna and rainforest pollen closely paralleled changes in climate

11

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and sea levels. At the same time large areas of the continent underwent uplift, restricting the presence of suitable habitats for lowland rainforest. The combination of drought and reduced habitat probably resulted in widespread extinctions in the rainforest flora from the Late Miocene into the Pliocene (Morley 2000). During the Quaternary rainforests expanded and contracted with the glaciations of polar regions. About 18,000 years ago the last severe ice age climaxed, leading to lower sea water levels, cooler periods, and less precipitation. The rainforest contracted to a few patches, the so-called forest refuges. These were probably situated near Cape Palmas and Mount Nimba in Liberia, and near Cape Three Points in Ghana (Hamilton 1976, Maley 1996). The glaciations have left their imprint, and even nowadays many plants, and even some animal species are confined to these former rainforest refuges (Martin 1991). The last ice age lasted till 12,000 years ago, giving way to warmer and wetter climate, and an expansion of the rainforest zone. 6000 years ago rainforest occupied a much larger area than presently, and extended across the Dahomey gap, thus connecting Upper and Lower Guinea. Africa has often been described as “the odd man out”, because its rainforests have a much lower species richness, and less palm, epiphyte and understorey species compared to Neotropical and Asian rainforests (Richards 1973). One reason is that the African rainforests are less extensive than their Neotropical and Paleotropical counterparts, and thus can harbour a smaller species pool. Another reason is that the African continent is relatively arid, and there is a limited availability of lowland habitat below 500 m altitude. As a consequence, the effect of the glacial periods must have been felt much stronger here than in other continents (Harrison et al. 1981, Morley 2000). West Africa has a long history of human occupation. The first European travellers in the 16th century give account of already large populations and extensive farming activities in large parts of the Upper Guinean forest zone. It is a wideheld believe that, since the early 1900s, rising population densities and unsustainable farming practices have led to a unilateral and accelerated rate of forest loss. This view has recently been challenged by Fairhead and Leach (1998) who suggest that extent of forest loss during the 20th century has been overestimated. Much of the deforestation either took place much earlier, or did not take place at all since the areas in question did not carry forest in recent historical times. Fairhead and Leach argue that in many cases traditional forest management practices actually have contributed to afforestation, and that many of the Upper Guinean forests are of recent origin, because population densities in the 19th and early 20th century have been much lower than before.

12

Outline This book is divided into two parts. The first part introduces the reader to the forests of Upper Guinea; their distribution, composition, biodiversity, conservation and management. The second part presents ecological profiles for a set of rare, endemic, or commercial species. Forests in West Africa, and specifically Côte d’Ivoire, have disappeared rapidly during the last decades. It is important to know the current deforestation rates, and to know where are the main forest blocks left. In Chapter 2, Chatelain et al. address these questions by analysing (changes in) forest cover at different spatial scales, using satellite images and historical analysis. Comparisons are made at subcontinental, national, regional and local scale. The spatial and temporal patterns in deforestation are explained in terms of socio-economic and geographic conditions. The Upper Guinean forests are bounded by extensive savannas. The transition between forest and savanna is surprisingly sharp, and can easily be observed from satellite images. In Chapter 3 Gautier and Spichiger analyse how climatic and edaphic conditions affect this forest-savanna transition at different spatial scales. In turn, the effects of climate and soils are modified by natural fires and anthropogenic disturbances. The forest savanna boundary is by no means stable, and has shifted considerably over time. During the last decades the forests have expanded in many places at the expense of the savanna. It is discussed how the present situation is the result of the simultaneous action of several factors in the past and present and the need for new policies addressing the questions of fire regime and land use is stressed. Within the forest zone, variation in species composition and structure is gradual and continuous. Nowadays, some excellent local- and national-level forest vegetation maps are available, but maps at a subcontinental scale are lacking. Exchange of experience and knowledge between neighbouring countries is hampered because the vegetation classifications differ. The combination of the existing vegetation maps is difficult because of the differences in criteria and legend systems used. In Chapter 4 Bongers et al. provide a forest gradient map for Upper Guinea, using forest inventory data of 40 large tree species. Variation in species composition is subsequently explained in terms of climate and soils. In Chapter 5 Kouamé et al. provide a detailed analysis of the floristic diversity in Côte d’Ivoire. In Côte d’Ivoire there is a strong south-north rainfall gradient and the vegetation composition changes accordingly, from moist forest in the south to semi-deciduous forest in the north. The forests in the eastern and western side of the country are partly isolated by a savanna intrusion, the so-called VBaoulé. It has been hypothesised that this savanna

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Box 1.1 Botanical exploration in Upper Guinea At the end of the 17th and 18th centuries the first botanical collections were made in West Africa, especially along the coast. The first floras for the region were published from the beginning of the 19th century, and from 1868 volumes of the Flora of Tropical Africa (Oliver et al.) became available; this flora was completed in 1937. The completion of the Flora of West Tropical Africa, covering the region east to western Cameroon, in 1936 was a major breakthrough, although it is concise (Hutchinson & Dalziel, second edition completed in 1972). For most West African countries national floras do not exist, and when they do they are incomplete or dormant (Berhaut for Senegal, Flore de Guiné-Bissau). The flora of Togo (Brunel et al., 1984) is the only complete national flora in West Africa, and there is an ongoing project for a flora of Benin. Checklists of varying scope and quality exist for

intrusion has been a main barrier for species exchange, in a similar way as the Dahomey gap forms a barrier for species exchange between Upper and Lower Guinea. In this chapter, Kouamé et al. discuss the role of the V-Baoulé and other environmental factors in determining species composition of Ivorian forests. The rainforests of West Africa have been earmarked as one of the world’s hotspots of biodiversity. This leads to the question: where exactly are the hotspots of biodiversity within this hotspot? The botanical background information necessary to delineate such areas is to date scarce or highly fragmented. In Chapter 6, Wieringa and Poorter explore how, and to what extent, herbarium collections (Box 1.1) can be used to define hotspots of biodiversity in West Africa. They selected 600 species that are rare or endemic to the closed forests of Upper Guinea. For the first time they present a higher resolution map that places the high biodiversity areas in a Guinea-wide perspective, and indicate in a quantitative way which areas are virtually unknown and merit further botanical exploration. Some of the centres of diversity they found have been postulated to be Pleistocene refuges. In an overall analysis, Wieringa and Poorter weigh the relative importance of history and environment in determining the current patterns of diversity. Understanding species distribution patterns is a crucial step for the conservation and management of species, plant communities and ecosystems. This is particularly important for species with small distribution ranges, or occurring at low densities. In Chapter 7 Holmgren et al. analyse the distribution of nearly 300 rare and endemic species. Which species show disjunct distributions, and which ones have an extremely narrow distribution range? How common are these different distribution types amongst the selected species? Finally,

Senegal, Gambia, Guinea-Bissau, Sierra Leone, Liberia and Côte d’Ivoire. It is worth attention that Liberia, where the flora is probably the richest for the region, is among the lesser-known areas concerning plant diversity. Adam (1971-1983) published a flora for the Nimba massif and, together with Jaeger (1980-1981) for the Loma Mountains in Sierra Leone. It was not until after World War I that local herbaria began to be established in West Africa. However, substantial West African collections are held only outside the region, above all in Europe (mainly at Kew, United Kingdom and Paris, France, but also at Geneva, Switzerland, Brussels, Belgium, and Wageningen, The Netherlands). Herbaria harbouring important collections from the Upper Guinean forest zone still exist at Abidjan (Côte d’Ivoire) and Accra (Ghana).

Holmgren et al. explore to what extent these distribution patterns are shaped by climate, soil conditions, and historic events, and indicate priorities for conservation. How can the botanical and ecological information presented here be used for the conservation and management of the remaining forest resources? In chapter 8 Bongers et al. explore the implications of the results of the work outlined above for conservation and management of forests in the area. They also address the current status of the Guinean forests and indicate where the bestpreserved forests are left, and in which forest types preservation is scarce. The hotspots of biodiversity are located and linked to the protection status in the present system of forest reserves and parks. For the selected commercial species they show where the areas with large commercial stock are concentrated. Finally, the question of the potential trade-off between timber exploitation and biodiversity conservation is addressed. The second part of the ecological atlas contains ecological profiles for 286 rare and endemic species (Chapter 9), and 56 large, commercial tree species (Chapter 10). These species represent c. 11% of the total forest flora of Upper Guinea, and include the most important species from a conservation or commercial point of view. For the large, commercial species an up-to-date summary of existing information is given. For the rare and endemic species new relevant ecological information is presented, because hitherto little was known about them. One species is presented per page. Each species account includes a distribution map, photograph, line drawing, and environmental table. First a short botanical description of the species is given, followed by information on its chorology, distribution type, distribution range and its rarity. The environmental requirements of the species are described in terms of light, water and nutrient availability.

13

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Finally, regeneration, growth characteristics and main uses of the species are discussed. Each description is accompanied by the most relevant literature references. This part is ended with Chapter 11 in which Jongkind gives the complete checklist of Upper Guinean forest species. Synonyms are included and also look-up tables of families and genera. These 11 chapters together provide a state-of-the-art review of biodiversity and forest ecology of West African forests. They also provide an in-depth analysis of ecological factors that give rise to biodiversity, and structure tropical plant communities. As such this ecological atlas provides a valuable companion to The woody plants of western African forests. A guide to the forest trees, shrubs and lianas from Senegal to Ghana (Hawthorne & Jongkind 2003). In this plant guide identification keys are provided for over 2000

14

species, and drawings of nearly all are presented. Also, colour photos of 1000 plants are included. The present atlas provides background information on the forests in the region, and gives excellent additional information on the ecology and distribution of a large number of species.

Acknowledgements We thank M.D. Swaine for his thoughtful comments on the manuscript, and V. van Engelen for critically reading the section on geology and soils.

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2

C. Chatelain, H. Dao, L. Gautier and R. Spichiger

H A P T E R

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Of the forest that covered West Africa at the beginning of the 20th century, the large majority has disappeared and the Taï National Park forest in Côte d’Ivoire is now the last important one. Notwithstanding this rampant deforestation, it is necessary to remember that the forest was not always present throughout the centuries: forests took over savanna zones, and vice versa (Guillaumet & Adjanohoun 1971, Maley 1996, van Rompaey 1993). More recently, extensive savanna zones got forested during the 1850 to 1920 period (Richards 1996, Fairhead & Leach 1998). Because of the forest’s natural cycle of regression and progression, it is difficult to speak of an original or primary forest. This is especially the case for the moist semi-deciduous forest domain (Hawthorne 1996) and for the savanna woodland transitional zone (Spichiger & Blanc-Pamard 1973, Spichiger 1975). However, we need to use these terms to differentiate this state from recently developed secondary forests. The FAO/UNEP (1981) study on the state of tropical forest in 76 countries showed that on continental or national scales, deforestation was very high between 1975 and 1980. In order to understand the need to conserve certain forests and the impact of deforestation on environmental and economic changes, we must analyse the actual situation of forest cover. Such an analysis is needed for the development of a management policy for the forests at the sub-continental or national level. In this chapter we analyse in detail the changes in forest cover over the last ten years for Côte d’Ivoire, a country with a long history of forest exploitation (Meniaud 1922, Aubréville 1958, Adjanohoun et al. 1966, Lanly 1969). Since the 1970s, the Ivorian forest has the highest rate of deforestation among tropical countries, 6.5% forest loss per year, compared to a global average of 0.5% per year (FAO/UNEP 1981). For the 1981-1990 period, the average rate of deforestation in Côte d’Ivoire has been assessed at 7.6% per year (850 km2/yr). In 1990 only 11,230 km2 of dense forests were remaining in Côte d’Ivoire, from an original forest cover of 150,000 km2 (FAO 1993, but see Fairhead & Leach 1998), approximately 7.5%. Areas with a formal protection status have become essential to the conservation of natural settings. Except for

Figure 2.1 Azagny National Park in 2000 consists of forests and swampy savannas. Although this is a national park, the borders are strongly degraded by agricultural activities. A full-colour version of this figure can be found in Appendix 1.

national parks, the purpose of most of these areas is a combination of conservation and production and exploitation of wood, as is the case for the “forêts classées” of Côte d’Ivoire or the forest reserves of Ghana. Although these areas have played a significant role in the preservation of forest, in many cases forest management plans have forgotten to take the rural setting of these forests into consideration. For the sake of improved environmental management, forestry policies must integrate rural and state domains. A global approach to deforestation, using low resolution images, allows us to acquire information on the whole region and to identify large remaining forests which are the more important for conservation. But in west tropical Africa, the high degree of fragmentation and the complexity of the landscape mosaic is difficult to resolve (Päivinen et al.1992, Husson 1995). It is necessary to use high resolution images such as LANDSAT in order to describe all the parameters needed to evaluate the extent of deforestation. These parameters are the distribution of small forest patches, the dynamics of secondary forests, the density of cash crop cultivation, and the density of cleared areas bordering on or in protected areas (Chatelain et al. 1996b).

15

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Chapter 2. Forest cover changes in Côte d’Ivoire and Upper Guinea

A

B

Figure 2.2 Forest cover in West Africa. (A) Cover according to NOAA-AVHRR satellite images (Paivinen et al. 1989). The eastern part of Liberia and western part of Côte d'Ivoire, including Taï National Park, are the last large forested areas that remain. (B) Cover according to the "eco-regions" map of Olson & Dinerstein (1998) representing the forest cover zonation with the limits in 1912 according to Chevalier (1920) and the limits in 1923 following Shantz & Marbut (1923). The variability seems to be due to the personal view of the authors rather than to changes in forest cover between 1912-23. A full-colour version of this figure can be found in Appendix 1.

Figure 2.3 Forest cover in Côte d’Ivoire in 1955-58 according to the vegetation map (scale 1:500,000) of Guillaumet & Adjanohoun (1969) and limits of the vegetation domains following Monnier (1983). A full-colour version of this figure can be found in Appendix 1.

16

Figure 2.4 Forest cover in Côte d’Ivoire in 1993 (Dao 1999) based on NOAA-AVHRR images. The Taï forest area represents at least 40% of the total forest area of Côte d'Ivoire. Also the forest area south of Abengourou (the classified forests of Yaya-Bossématié-Mabi) is very important. A full-colour version of this figure can be found in Appendix 1.

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Chapter 2. Forest cover changes in Côte d’Ivoire and Upper Guinea

In this chapter we will report on forest areas, and their changes, at several scales of resolution. Firstly, we will describe the total forest areas in West Africa. Secondly, we will analyse the total forest cover of Côte d’Ivoire, in particular the total protected area. Thirdly, we will analyse this for the region of Abidjan, as an example for the country. Lastly, we will analyse forest fragmentation in detail for eight blocks of 20 x 20 km in that area.

Method Study area We studied the changes in forest cover of the dense forests on four scales. The first scale is the level of West Africa. This takes into account all the countries of West Africa that had part of their territory covered with rainforest and comprises the phytogeographical zone of Upper Guinea from West Senegal to Nigeria (Figure 2.2). The forested zone probably covered between 56,600,000 to 68,000,000 ha at the beginning of the century, representing 20% of the African forest cover (Sommer 1976). We used the vegetation maps of Chevalier (1912) and of Shantz and Marbut (1923) to assess the forest cover at the turn of the century, remaining cautious in the interpretation of their results. The second scale is the national level (Figure 2.3). This allows a more precise vision of deforestation in Côte d’Ivoire. The cartographic data used for this approach are the vegetation map of Guillaumet and Adjanohoun (1969) at 1:500,000, which is based on the interpretation of aerial photographs taken from 1955-58, and the NOAAAVHRR images of 1993 (Dao 1999). The third scale is the regional level. We chose the region including Abidjan in southeast Côte d’Ivoire, as it is the only region for which a time series of images exists. LANDSAT images taken in 1990 and 2000 cover the region between the coast of the Gulf of Guinea, including Abidjan, and the V-Baoulé (Figure 2.4). This represents an area of 180 x 180 km, about a third of the Ivorian forest

zone. This area shows many types of forests and also many types of farming activity, essentially due to the variability in climate and soils. We have also used the topographical maps of the Côte d’Ivoire Geographical Institute (Dao 1999) which are based on aerial photographs taken in 1958 and which are more precise than Guillaumet and Adjanohoun’s maps (1969). The fourth scale is the local level. At this level we analyse changes in forest cover of eight blocks of 20 x 20km extracted from the LANDSAT images. Both the evergreen forest zone and the moist semi-deciduous forest zone are present here. The characteristics of the maps and images are summarised in Table 2.1. Treatment of the 1990 and 2000 LANDSAT images was done with IDRISI software. Digitisation of the old maps was done with ARCINFO software, and the graphic production of the maps was done with ARCVIEW. These data were all connected to SIGIVOIRE (Chatelain & Gautier 2002). Image classification The vegetation map of 1990 was obtained by classifying three infrared channels of LANDSAT, using 45 classes (the visible channels are unusable). The classes that represented identical or similar subjects were then merged to obtain ten classes. The 2000 map, on the other hand, was obtained by automatic classification with the IDRISI “isocluster” method, which uses 32 classes. These classes were then regrouped to arrive at 15 classes that are identical to those of the 1990 map. Verification of the classification was done by utilising the “kappa” index (Congalton & Mead 1983). We did not keep the degraded forest classification because almost all the forests in this region are degraded, making it impossible to distinguish the states of degradation on the 2000 image, as opposed to that of 1990. The retained classes are: Forest on firm land: this groups most of the forests (degraded forests, wetland forests, old secondary forests, etc.). Secondary forest: woody vegetation that reaches 10 m in height, with occasional presence of palms. This includes plantations of cola-nut trees in this region as well as degraded forests with a cover of less than 10%.

Table 2.1 Cartographic data used in the present study. Year

Scale

Type

1958 1958 1990 1993 2000

1:200,000 1:500,000 resolution 30 m resolution 1 km resolution 30 m

Topographical map (IGCI), used for the region around Abidjan. For explanation of the legends, see Dao (1999) Vegetation map (Guillaumet & Adjanohoun 1969), used for the national approach, based on aerial photos from 1955-58 LANDSAT TM (196-56, December 24th 1990) 3 images NOAA AVHRR in 1993 LANDSAT ETM+ (196-56, February 9th 2000)

17

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18

7737 17097 3576.8 2773.9 501.4 2643.4 Total surface

1390.5

139.8

3266.1

2329.4

35.6

1356.

3703.0

3283.9

3261.3

865.2

874.9

32.1 583.8 3575.3 2684.9 474.4 2233.7 Outside zones

1166.6

127.0

2766.5

1984.8

34.1

1248.7

3320.2

2842.0

2856.0

742.3

771.4

194.0 224.6 139.9 277.2 152.7 2100.8 5.6 2.8 4.9 66.1 138.5 52.7 Loviguié Yapo Dogodou

21.0 8.2 35.6

0.5 1.5 2.5

102.4 72.5 33.6

78.4 46.4 25.4

0.1 0.2 0.3

9.2 14.2 15.3

43.2 47.0 40.2

16.5 17.4 64.1

11.6 16.0 85.7

19.5 9.8 6.0 6.8 11.5 10.8 15.2 9.5 22.5

0.0 0.7 0.2

40.9 18.1 227.3 193.3 149.4 377.8 368.7 196.9 128.9 182.3 52.3 0.1 0.5 4.2 11.0 12.3 26.6 6.2 6.2 8.6 1.1 0.9 0.6 4.6 6.5 11.5 0.5 60.3 52.2 27.9 Divo Hiré Go Abié Sikensi

11.8 8.5 90.0 12.0 36.7

1.6 0.7 3.1 0.3 2.7

12.5 6.9 71.6 124.9 75.3

20.0 10.3 18.0 79.3 66.9

0.0 0.0 0.3 0.1 0.4

12.3 22.0 7.8 7.2 20.0

25.8 54.7 34.6 75.6 61.6

102.4 131.4 60.4 15.4 34.4

106.6 126.7 33.1 3.5 22.1

25.8 6.9 10.2 11.0 21.8

0.1 0.1 0.0 0.1 0.3

forest 1990 forest 1958 water palm defor. soil- savanna bush bush refor. refor. SF SF forest Interpretation

defor.

defor.

degrad.

SF

water water palm-soil palm-soil soil-savanna soil-savanna SF bush soil bush bush bush bush SF soil SF soil forest SF SF SF SF forest SF forest soil forest bush forest forest Legend 1990 Legend 2000

Table 2.2 Local and regional approach of forest cover changes (km2) in the Abidjan region between 1990 and 2000 for each of eight 20 x 20km blocks and overall values for the whole Abidjan region. The whole area was 25,931 km2. SF = secondary forest. The classes soil, savanna and palms are lumped, see text.

Chapter 2. Forest cover changes in Côte d’Ivoire and Upper Guinea

Thicket: woody vegetation 1 to 3 m in height. It can be confused with coffee and cacao plantations which are of a similar density and height to thickets. Thickets precede the first stages of secondary forest. Coffee-cacao: plantations of coffee and cacao, with or without forest cover. Cultivated grasslands: fields under annual cultivation and grassy fallow lands. This covers large areas of lowlands where rice is cultivated. Bare ground, village, roads: this is made up of areas with no vegetation. Orchard plantations: this is made up largely of plantations of oil palm trees, pineapple and banana. Rubber and other tree plantations: we retained this class because of the existence of large plantations in the North zone and the Dabou region. It is difficult to distinguish this classification from the previous one. Savanna: we have merged all types of savanna into this classification. Water: this includes the ocean, lagoons, lakes and rivers. Before being superimposed for comparison, the 1990 image needed to be geographically corrected based on the topographic maps. The 2000 image was corrected based on the 1990 image. The map of the forest cover that was made based on the 1955 and 1958 aerial photographs could not be superimposed exactly on the satellite images due to the differences in scale. The identification of palm plantations posed a problem because they were sometimes confused with forest: however, their homogeneous structure and the presence of rectangular trails makes them visible to the naked eye. To deal with them, we manually mapped the plantations and assigned their zones to the corresponding class. The same errors occurred for coffee-cacao plantations, and for the division between secondary forests and very degraded forests. These were of minor importance, however. Verification of the 2000 map was done on the basis of that of 1990 for which we had already done verifications between 1990-93 within the framework of preceding studies (Chatelain 1996, Dao 1999). Comparison of images The 1990 map was qualitatively compared (cross tabulation method) with the 2000 map to obtain 10 x 10 classes (100), then filtered to eliminate isolated pixels. Only 15 classes had a real relevance concerning the development of the forest (Table 2.2). The rest of the classes were merged either because they covered small areas without relevance (e.g. bare ground, water, savannagrasslands, coffee-cacao thicket etc.) or because they remained ambiguous and could not be interpreted (palmground, palm forest). The values of the forest area of 1990 and 2000 and the evolution between 1990 and 2000 are the outcomes of this analysis, so only the forest classification was retained to express the area of forest

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cover. The secondary forest classification was not kept except for certain statistics, and this will be noted when applicable. Verification of such a comparison was done in a quantitative way with the infrared channel (TM7) of the 1990 and 2000 images. This, after the calibration of the two images, enabled us to find the pixels of which the values have been either increased or decreased following the increase in biomass or reforestation. The superposition of the results of this quantitative comparison with the qualitative approach gives a tool for verification (in the case of the deforested zones we found a concurrence of 70%). Fragmentation On the national scale, the pixels on the maps were reclassified into forest and non-forest with the aim to analyse the distance of a forest pixel to the nearest forest limit. For each pixel, the Euclidean distances to the forest limits were calculated. The negative values give the distance from a pixel inside a forested area to the forest limit. The more negative the value the further the distance to the forest limit and the less the external influence on the forest ecosystem. We calculated a “core area” defined as that part of the forest that was more than 1 km away from its border. The positive values give the distance between the blocks of forest. For each block of forest, we calculated a perimeter-area relationship (PA). The PA has a value between 1 (maximum fragmentation, in the case of an isolated pixel) and 0 (theoretical case of an infinite area). The PA index takes into account both the form and the size of the blocks: a square of 4 pixels has a higher index (0.5) than a square of 16 pixels (0.25), but a lower one than a line of 4 pixels (0.62). These methods have been tested and used by Dao (1999).

Results

1.

Scale of West Africa

As difficult as it is to get good estimates of recent forest cover with the tools of satellite imagery, as is shown with the disparity of values in Table 2.3, it is even more difficult to get a good view of the forest cover at the turn of the last century or before. The difficulty is even greater since the terms initial, original or primary forest are all relative considering the dynamism of the vegetation over time (Richards 1996). Fairhead and Leach (1998) showed convincingly that the forest cover in West Africa at the turn of the century was over-estimated. We took this into account in adopting the most precise estimates possible by using the original vegetation maps of Chevalier (1920) and Shantz & Marbut (1923) (Figure 2.2B). Although in Sierra Leone the forest mosaic in the forest savanna transition zone indeed was mapped as forest (and thus gave rise to over-estimation of forest - Fairhead & Leach 1998), this was not the case in the other countries in West Africa. Also the coastal zone, which also is a mosaic, was not mapped as forest along its entire length. The forests along rivers, so characteristic of Guinean savanna, were mapped separately, but the size of these areas was indeed excessively large, for typographical reasons. Excluding mosaic zones or forest gallery, Chevalier’s 1912 map (Chevalier 1920) shows an area of 29,205,000 ha in Côte d’Ivoire, Liberia, Ghana and Sierra Leone (Table 2.3). The same analysis of Shantz’s map (Shantz & Marbut 1923), a less precise map, shows an area of around 20,849,000 ha.

Table 2.3 Forest surface in thousands of ha for different countries and different sources. According to these data, deforestation occurs only in a few countries. (1) Chevalier 1920; (2) FAO/UNEP 1981; (3) Paivinen et al. 1989; (4) SOFO 1997; (5) Odoom 2000; (6) Iremonger et al. 1997; (7): FAO 2000. Countries are ordered from west to east. Country

Senegal Gambia Guinea Bissau Guinea Sierra Leone Liberia Côte d'Ivoire Ghana Togo Benin Nigeria Total

Total surface area of the country

% of protected forest

19200 1000 3600 24500 7160 11000 31800 24000 5400 11300 91000 229960

12.0 3.7 0.0 1.8 5.1 1.5 10.0 3.3 3.4 14.0 4.7 6.0

1912 (1)

1980 (2)

1989 (3)

1990 (4)

1238 8465 13672 5830

2200 60 660 2000 700 2000 4500 1710 340 4700 5900 24770

210 50 690 760 510 4120 310 9600 1360 4200 3860 25670

7600 100 23.6 6700 1520 4600 5600 9600 1330 4900 1430 45740

1990 (5)

7663 1895 4639 10961 9608 1370 2104 2790

1995 (4)

1996 (6)

7300 91 2300 6367 1309 4507 5469 9022 1240 4625 1370 43460

11100 480 2010 7660 1360 6320 7780 5970 420 2110 27900 73110

2000 (7)

6929 1055 3481 7117 6335 510 2650 13517

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If the conservation of the “Congolian” forests is less of a problem due to their extent, the conservation of the forests of Upper Guinea is more and more at risk if an effective management strategy is not put into place. Of the rainforests that at the turn of the last century still covered a small part of Sierra Leone (1,238,000 ha), half of Liberia (8,465,000 ha), half of Côte d’Ivoire, a little under half of Ghana (5,830,000 ha), and a little bit of Togo, today there remains only the Taï National Park in Côte d’Ivoire, a few large forests in Liberia, and some forests in Ghana. The total area of these forests from Guinea to Nigeria totaled 48,783,000 ha in the eighties (FAO 1993) and 41,594,000 ha in the nineties (FAO 2000), that is, around 60% of the forest at the beginning of the century. Aside from the National Parks, these last large forest tracts are degraded by forestry exploitation. The amount of deforestation and the policies of protection (definition of protected areas and respect for these areas) differ largely between countries (Table 2.3). According to the FAO (2000), between 1990 and 1995 the annual percentage of deforestation varied between 0.56% for Côte d’Ivoire and 3% for Sierra Leone. While in Ghana the protected areas are part of a forest network, in Liberia the situation is hazier. The most recent forest cover data are those of Päivinen et al. (1992) (Figure 2.2), De Monza (1992) and Parren & de Graaf (1995). The data of the FAO (1993, 1995a, 2000) are simply extrapolations taken from these data, and there is no recent map of the total forest cover in this region. In the case of Côte d’Ivoire, the data of Odoom (2000) and the FAO (1993) showed an increase in excess of 1,500,000 ha of forested area between 1990 and 2000. This, as we will see, is a flawed result, whether or not we take forest plantations into consideration.

2.

The protected areas The total area of protected forests covers 5,667,405 ha in 1992, which represents 17.6% of the area of Côte d’Ivoire, which is considerable (Table 2.4). The “forêts classées” cover 2/3 of the protected areas, while national parks represent 1/3. The “forêts classées” are located mostly in the forest zone, as whilst only 40% of the area of the national parks are located in forest zone. Two national parks alone, Comoé (in the northeast of the country, 1,167,167 ha) and Taï (southwest, 423,942 ha) total 1,591,109 ha, that is, 81% of the total area of national parks, and 28% of the total protected area. 134 of the 189 “forêts classées” (71%) have an area equal to or less than

National scale: Côte d’Ivoire

The oldest map showing forest cover of Côte d’Ivoire was the one of Chevalier (1920) (Figure 2.2B), which enabled us to estimate the forest cover at approximately 13,670,00 ha in 1912. This value is, in our opinion, indicative, since Chevalier’s map did not consider mosaic savanna forests or the entire coastal zone, which he considered non-forest. In contrast, the mosaic zones in community forest gardens and the areas of towns and villages are indistinguishable from the forest. The map of Guillaumet and Adjanohoun (1969) (Figure 2.3), based on data of 1955-58, shows a forested area of 8,640,000 ha. From 1900 to 1955, the forest cover did not change much compared to later years, despite the creation of numerous pioneering fronts of forest exploitation during that period (Kolibi 1990). In the whole of Côte d’Ivoire, between 1958 and 1993, more than 80% of the rainforest disappeared - about 60,000 km2. According to the 1993 NOAA classification (Dao 1999) (Figure 2.4), the total forest area dropped to

20

1,994,100 ha (of which 1,920,700 ha is within the forest zone, the rest being isolated forest patches in the savanna zone). It means that only 6% of the forest part of Côte d’Ivoire is considered as forest zone. These values differ greatly from those published by the FAO (2000). Thirty-six per cent of the forests are in two administrative regions (Table 2.4) alone - the Southwest and the South - and consist of six main forest areas: the Taï region with the Taï National Park, the Goin-Débé and Cavally “forêts classées”; the Agboville-Abengourou region with the forests of Mabi-Yaya and Yapo, and the Guitry region with the Gô-Bodiénou “forêt classée”. At the same time some regions have no forest at all. This is the case most notably in the Daloa-Gagnoa-Soubré triangle in the Central-West region, which was the first zone of agricultural colonisation that moved towards the country’s southwest during the 1960s. Compared with Monnier’s (1983) ecological zones, forest is found mostly in the evergreen forest zone (25% of the forest, a total of 1,657,100 ha) and is least present in the moist semi-deciduous forest zone (3% of the forest, a total of 263,600 ha) (Table 2.4).

Table 2.4 Forest cover as percentage of total area in the different administrative zones of Côte d’Ivoire in 1993. The forest cover is based on NOAA satellite images. The protected forest percentage is the percentage of the forest in the region. Region

Forest NOAA (ha)

Center Center East Center North Center West North Northeast Northwest West South Southwest Total

3,400 49,400 600 79,700 0 44,400 100 746,200 427,000 643,400 1,994,100

% of total of the zone

Protected forest (%)

0 7 0 3 0 1 0 25 11 25 6

7.9 14.6 15.7 24.4 6.5 28.5 16.3 20.9 36.5 14.7 17.6

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variable area of protected forest cover. The West region of Côte d’Ivoire (28.5% of the area is forest) and especially the Southwest region (36.5%) are most densely covered with protected forests. This high value is principally attributed to the presence of the Taï forest. In the same way, the Northeast region, which includes the Comoé national park has 24.4%. At the scale of Côte d’Ivoire, the regions most poorly covered with protected forests are found in the borders of the North (around Tingréla, in the

20,000 ha, and have a cumulative area of only 16% (878,804 ha) of the total 5,667,405 ha. It is to be noted that the savanna zone contains proportionally more small sized “forêts classées” (less than 10,000 ha), than the forest zone, that is, 66% compared to 43%. The actual forest cover The different regions in Côte d’Ivoire have a highly

Table 2.5 Forest cover of some protected forests. Calculations are based on the NOAA images of 1992 and 1993. The fragmentation index Perimeter / Area (PA) and the core index (distance to the interior of the forest) are shown as well. Name

Classified surface

Forest surface 1992/93 (ha)

Forest surface 1992/93 (%)

Core index area 1993 (%)

PA index (%)

Taï (P.N.) Hte Dodo Cavally-Goin-Débé Mabi-N'To-Songan-Tamin-Yaya Ht Sassandra

422,076 245,438 198,473 184,689 102,347

393,287 160,901 137,846 111,618 69,411

93 66 69 60 68

88 44 55 36 55

0.03 0.07 0.06 0.10 0.06

N'Zo (R.F.) Niégré Scio Duékoué Mt-Péko (P.N.) Okromodou-Diogoro-Bogbo Go-Bodiénou Yapo Tiapleu Niouniourou

76,120 97,468 90,271 51,845 30,985 90,149 61,238 25,373 19,274 18,786

59,164 55,870 54,528 30,253 23,056 19,274 18,176 18,054 17,932 17,688

78 57 60 58 74 21 30 71 93 94

64 37 39 38 42 11 16 41 56 69

0.09 0.07 0.09 0.11 0.12 0.06 0.07 0.10 0.20 0.11

Mt Momi-Sangouiné Béki-Bossématié Asagny (P.N.) Mt Glo Krozialé Dogodou N'Guéchié Banco (P.N.) Mt Bableu Mt Glas

32,693 37,572 19,396 10,613 9,149 6,343 3,538 2,196 4,148 1,464

15,614 15,370 12,199 8,295 7,807 3,294 3,050 2,196 2,196 732

48 41 63 78 85 52 86 100 53 50

10 24 28 47 56 25 38 44 26 8

0.23 0.06 0.19 0.19 0.12 0.13 0.15 0.21 0.16 0.15

Table 2.6 Number of forest islands per size class (ha) for each of eight 20 x 20 km blocks in the Abidjan region in the year 2000. The percentage of the forest surface of some classes is given between brackets. Zones

0.3 -2 ha

2-4 ha

4-8 ha

8-16 ha

16-32 ha

32-64 ha

> 64 ha

Total (ha)

Divo Hiré Gô Yapo Abié Sikensi Loviguié Dogodou Mean

135 34 952 581 1358 1034 (43%) 1099 (18%) 577 (8.5%) 721.2

8 2 141 50 229 127 (21%) 189 (12%) 68 (4%) 101.7

8 1 72 36 135 50 (17%) 81 (10%) 46 (5%) 53.6

2 0 31 15 53 14 (8%) 33 (8.5%) 29 (7%) 22.1

1 0 16 5 18 3 (4%) 18 (9%) 9 (4%) 8.7

1 0 6 1 5 0 11 (11%) 12 (12%) 4.5

1 0 5 2 3 0 5 (28%) 3 (57%) 2.3

560 41 6795 12986 3895 1664 4286 4636 4357

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Fragmentation An analysis of the 1993 NOAA image shows that the forest is very fragmented. A total of 486 tracts were recognised. Measuring the level of fragmentation using the “Perimeter-Area” index (PA), based on the form of the fragments, shows that these forests have an average value of 0.17. The four forests with a PA below 0.1 are Taï, GoinDébé-Cavally, Haute-Dodo and Haut-Sassandra (Table 2.5). The core index represents the distance between the black border and the centre of the fragment. Only 22% of the forest areas are found at more than 1000 m from the edge of the forest, 10% are further than 5000 m. Only 14 fragments represent 75% of the forest cover. Figure 2.5 Area of closed forests in Côte d'Ivoire (1918-90). (Sources: Monnier 1983, Arnaud & Sournia 1978, FAO/UNEP 1981, FAO 1993).

3.

savanna zone) and in the Central-East (around Bondoukou, at the limit of the savanna forest). In the forest zone, the extreme Southeast and most of the Daloa-Gagnoa-Soubré triangle in the Southwest are farthest from all protected forests. The latter zone was the “frontline” of the colonisation of the southwest since the end of the 1960s: a zone of occupation and movement towards the open spaces of the southwest. It is still subject to intense agricultural exploitation today. To a lesser degree, the Bongouanou-Daoukro-Bocanda triangle (known as the “cacao ring”) also has a deficit in forests. The same applies to the area around Adzopé (in the midst of the oldest area of coffee and cacao cultivation) and to the coastal regions of Jacqueville and Bonoua, zones of large industrial plantations, notably of palms and bananas.

The forest covered approximately 2643 km2, or 8% of the Abidjan region’s area (Figure 2.6) in February 2000. The area deforested between 1958 and 1990 was 10,580 km2 while between 1990 and 2000 it was 1530 km2 (Figure 2.6). The area deforested since 1990 seems comparatively small, however more than the half of the remaining forest area has disappeared during this interval and there is hardly any forest left. If we consider the percentage of forest cleared in relation to the total forest area over ten-year periods, the two decades 1970 - 80 and 1980 - 90 show a rate of 30%, while 1990 -2000 shows 20%. This is still considerable, and does not indicate a big change despite the development of a forest management policy. The values given for the deforested areas only consider the transfer from a forest legend to a thicket or cultivated land legend. More of the 1990 forest area has become secondary forest (Table 2.2), which is actually a degradation. However, image interpretation constraints do not allow us to map evolution from thicket to secondary forest.

Development of protected areas In general, while the number of protected forests has diminished since 1949, their area has increased. This is the case both for the “forêts classées” and for the national parks. The savanna zone shows an increase in both number and area of “forêts classées”. In the forest zone, including the national parks, the highest losses of protected forests were in the Central-West and Central-East regions, that is, on the border of the Baoulé people’s original homeland. The largest increase in protected forest area was in the Southwest region, where human influence did not increase much since 1949. In the “forêts classées” of the forest zone, remnant forests represent on average 42% of their area. The area of these “forêts classées” cover 1,218,000 ha, which represent 68% of the total 1,994,000 ha of forests in the zone. This average is misleading because only 17 forests out of 169 have a real forest cover more than 94% (Table 2.5). Most of them are located in the west of the country, with the exception of the Yapo “forêt classée”, the Mabi-SonganTamin-Yaya grouping, and Beki-Bossématié.

22

Regional scale: Abidjan region

Development before 1958 In 1958, the forest cover of the Abidjan region was 27,806 km2. Large forest areas were still numerous and the majority of “forêts classées” were still little touched by agriculture. The map of 1958 showed evidence of deforestation mostly in the east of the region (Adzopé, Agboville, Dabou). The region between Grand-Lahou and Divo was still intact, save for a hole expanding from Divo towards Guitry, clearing centred around the main localities and followed the main axes of communication, as the creation of these routes was linked to the extraction of wood and its transport to the ports (Arnaud & Sournia 1978). Development 1958-1990 A comparison of the 1958 map of land occupation with the 1990 classification (Figure 2.6B) shows that this period of 32 years had a high deforestation, especially between 1975-1980. Between 1958 and 1990, out of a

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A

B

Figure 2.6 Changes in forest cover in the region around Abidjan. (A) Changes between 1990 and 2000, based on LANDSAT images with 30 m resolution. The limits of the classified forests are indicated. Only the Yapo classified forest and Banco National Parc (near Abidjan) are completely covered with forest. (B) Changes between 1958 (topographic map) and 1990 (LANDSAT image) with a resolution of 250 m. The eight blocks of 20 x20 km each that are studied in detail (see Figure 2.8) are indicated. A full-colour version of this figure can be found in Appendix 1.

total zone area of 2,507,982 ha, over 51% (1,274,606 ha) was cleared and only 8% (193,163 ha) was left forested. In 1958, 41% (1,040,213 ha) of the zone was already cut. The 3% (84,900 ha) of land reforested between 1958 and 1990 was very fragmented. On average, 87% of the forested areas of 1958 had disappeared in 1990. Several small forested areas were not mapped as forests in 1958. We are certain that there has been no real reforestation, but that in 1958 an agrarian landscape with a mosaic of wooded fragments grown on the top of slopes was dominant. The zone around Abidjan currently is among the regions with the highest forest cover. The zones that were already fragmented by agriculture in 1958 are the least cut today, and those that were still forested at that time are the most deforested. Contrary to Fairhead and Leach (1998), we think that the old maps strongly underassessed the forest cover of the 1955-58 period because the maps did not take the small and very numerous groves into consideration, for reasons of scale (Chatelain 1996). The study region is covered with evergreen forest and moist semi-deciduous forest. The moist semi-deciduous forests have the highest deforestation rate and this can be explained above all by the development of coffee plantations, especially in the Tiassalé-Divo zone. The evergreen forest zone shows less clearing, and the forests saved are for the largest part situated on sites with difficult access (along the Agneby river, the wetland forest near

Dogodou and the Gô-Bodiénou forest) and where management policies have been established such as for the forests of Yapo. Development 1990-2000 In 1990, forests covered 7740 km2. During 19902000 deforestation was 20% (1530 km2) of that area and was mainly found in the evergreen forest zone, specially near Bandama (Figure 2.6A). Very large areas of forest have disappeared in the Gô-Bodiénou classified forest where deforestation began a few months after the image was taken in 1990. There is actually only 50% of this forest remaining. The Agboville zone, with a long history of human occupation, has paradoxically changed little and still has a significant forest area. The development map (Figure 2.6), shows hardly any large forested areas aside from those of the Yapo classified forest, and to a lesser extent, Banco national park, Azagny national park, the Agneby forest and the Dogodou classified forest. The forests of Mopri and Téné are nevertheless visible because of a small intact forest that is still present. Condition in 2000 Between 1958 and 1990, half of the forest cover disappeared, and in the following ten years half of the remaining forest was destroyed. From 1990 to 2000, about 140 km2 of forest were lost to cultivation and 3200

23

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A

B

Figure 2.7 Comparison between two interpretation maps of LANDSAT images with 30 m resolution. (A) Map from the present study, showing all forested areas independent of their size. Also secondary forests are given showing the density of human occupation. (B) Map from the 1993 "Bilan Forestier" SODEFOR (1993) based on a 1990 image and taking only forest fragments of more than 10,000 ha. The total area is 55 x55 km. A full-colour version of this figure can be found in Appendix 1.

km2 were exploited to such a degree that in 2000 they were identified as highly degraded forest or secondary forest. Secondary forests, mostly in the east of the region, or neighbouring highly degraded forests cover 11,192 km2 (Table 2.2). Only 23% of the residual forest cover is within the state-controlled “forêts classées”. There is almost no information available for 76% of the forest cover, mainly consisting of areas less than 1000 ha in size and not mapped on the Ivorian forestry map (Figure 2.7). In some cases, like at Divo, it is even possible to delimit classified forest “in negative”: more forest exists outside than inside the classified forest. In the case of the Mopri classified forest, the managers have conserved a tiny grove of primary forest in the middle of the forest, which represents at most 10% of the original forest. The rest is occupied by young tree plantations. Irobo classified forest shows the same scenario. Thus, in this region, these two tiny areas represent the last moist semi-deciduous forests.

forest tracts (Yapo forest, Gô-Bodiénou forest), almost the majority of forests are made up of tiny fragments distributed throughout the rural landscape. Their presence certainly plays a significant role in the process of environmental conservation. The many recent clearings indicate a highly dynamic pattern of deforestation in the studied blocks. The fragmentation of the forest can be assessed by surveying areas on the basis of surface classes (Table 2.6). The fragments of less than 4 ha in total represent between 10 and 30% of the total forested area, which is far from negligible and justifies our opinion that these areas should be taken into account in studies of forest cover.

Discussion

1. 4.

Local scale

We chose to analyse forest cover and changes therein using eight blocks of 20 x 20 km (Figure 2.8 and Table 2.2). The distribution of the forested areas is heterogeneous: some blocks have maintained a forested character despite their proximity to urban centres (Agboville region), while some isolated blocks have long ago lost their forests (Hiré region). Some “forêts classées” in the studied blocks are devoid of all forest. Although a large proportion of forest is still found in the form of large

24

Deforestation at various scales

West African scale West Africa is widely recognised as highly significant for the biodiversity conservation of the African dense forests (Conservation International 2001). The rate of deforestation is considerable, but information on forest cover differs widely depending on the sources (Matthews 2001), some of which show an increase of forest in certain countries (Table 2.2). Fairhead and Leach (1998) showed that the extent of the forest area at the turn of the 19th century was over-estimated. This was especially the case for

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A

The Hiré block, in the moist semi-deciduous forest zone. A high reduction of forest cover from 368 km2 in 1958 to 18 km2 (less than 5%) in 1990, and c. 0.4 km2 in 2000. Two thirds of the forests are less than 2 ha in area, and the “forêts classées” were completely cleared in 1990. In 2000 the whole area is characterised by old coffee-cacao plantations.

The Divo block. Deforestation here is quite similar to Hiré (from 337 to 40 to 6 km2). Forests here are a little more abundant due to numerous steep slopes covered with granite outcrops that are less conducive to agriculture. There is a unique tract of forest which is privately owned (ex IRCC). Partial exploitation of it, unfortunately, started in 1991. The Boubo “forêt classée” was completely transformed in oil palm plantations.

The Gô-Bodiénou block. Forest cover reduced from 196 km2 in 1958 to 227 km2 in 1990 to 67 km2 in 2000. This block has much closed broadleaf forest and was highly isolated until 1990. A large part of the block is occupied by the Gô-Bodiénou “forêt classée”. The landscape to the north of this forest is made up of numerous groves. Para-rubber tree and palm plantations have increased heavily. This “forêt classée”, along with those of Yapo and Niegré, are the three last large areas of forest in the South zone of the country. The Lovigué block. Forest cover reduced from 227 km2 in 1958 to 194 km2 in 1990 to 42 km2 in 2000. Most of the forest area is found outside the “forêts classées”. In 1969, deforestation already reached the borders of the “forêts classées”. In 1990, over 40% of the “forêts classées” of Mafé and Lovigué were already cleared, and in the decade to 2000, most clearing was done in the Mafé “forêt classée”. This block, along with that of Abié, shows the highest forest cover in 2000, mostly situated in the rural domain.

B

The Yapo block. Forest cover changed from 152 km2 in 1958 to 224 km2 in 1990 to 129 km2 in 2000. 14,900 ha of forest is situated within the “forêt classée” whose conservation status seems superficially good. However, the forests are extremely degraded due to high logging levels. This block show the lowest rate of deforestation. In spite of the strong human pressure due to the presence of large banana and pineapple plantations south of the forest, numerous small groves of secondary forests are present where cola-nut trees are often abundant. The block shows a coexistence of industrial along with traditional land use. The Abié block north of the Yapo “forêt classée”. Forest cover changed from 129 km2 in 1958 to 139 km2 in 1990 to 39 km2 in 2000. This block was considered completely cleared on the 1958 maps. It is, however, the block with the highest forest cover in the rural areas. Land occupation according to the relief is clearly visible on the images: forests are found on the tops of slopes and rice and taro fields on low-lying lands. The deforested parts are small in area and scattered. This block illustrates that it is meaningless to consider only large areas of forest in the national inventory. This block is the most forested one both in area and in number of forest fragments. The Sikensi block north of Dabou. Forest cover changed from 182 km2 in 1958 to 149 km2 in 1990 to 17 km2 in 2000. Forests having an area less than 4 ha represent 64% of the forest area. This block was crossed by a road a long time ago and this resulted in the establishment of many farms. In addition, the most important oil palm plantations are located in the south of this region. Clearing in this block is high and large areas of cleared land belong to a single landholder. Many areas far from the main road were only recently deforested. The Dogodou block. Forest cover changed from 210 km2 in 1958 to 139 km2 in 1990 to 46 km2 in 2000. Numerous migrants have established themselves along the coastal Abidjan-San Pedro road since it was improved in 1992. Coffee-cacao plantations, however, have been established to the northwest of this main road for a long time. The Yocoboue-Tiegba main road also crosses the block and led to agricultural occupation. Deforestation is essentially in the interior of “forêts classées” (10% of the classified forest area). The zone along the road to Tiegba is a mosaic of forest and agricultural land.

Figure 2.8 Changes in forest cover at a local scale for eight blocks of 20 x 20 km each in the region of Abidjan (see Figure 2.6). (A) Blocks located in the eastern part of the area. Left the changes between 1958 and 1990, right the changes between 1990 and 2000. (B) Blocks located in the western part of the area. Left the changes between 1959 and 1990, right the changes between 1990 and 2000 (for legends see Figure 2.6). A full-colour version of these figures can be found in Appendix 1.

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the savanna-woodland zones, most probably due to the incorrect interpretation of pioneer bush vegetation and of secondary forest. These were often interpreted as a forest degradation phase instead of a forest colonisation phase of the savanna (Spichiger 1975, Blanc-Pamard & Spichiger 1973, Gautier 1992b). Another explanation of this variability is the definition of rainforest, which varies according to each report (e.g. some take plantations into account, others do not). Another aspect is the absence of recent data, since the last maps made on the scale of West Africa date from 1990. This lack of sound information makes it very difficult to evaluate the level of deforestation, which should be a basis for proper management of forest areas. For the conservation of the last few forest areas, it is certainly necessary to redefine protected areas in Liberia and to reassign the roles of the “forêts classées” in Côte d’Ivoire and Ghana. The actual assessment of the distribution of the forests should also support the numerous confirmations and explanations of climate changes in West Africa (Anhuf 1995, L’Hote & Mahé 1995). In Côte d’Ivoire, deforestation has been higher in the moist semi-deciduous forest zone than in the evergreen forest zone. The impact of this is aggravated by the higher sensitivity of degraded moist semi-deciduous forests to fire hazard and infestation by pest plants such as Chromolaena odorata. Although the distinction between the two forest zones is fundamental for proper forest management, it is generally absent on the maps of West Africa such as those of White (1983) or of WWF (EcoRegions from Olson & Dinerstein 1998). This seems equally valid for Ghana where the moist semi-deciduous forest zone represents around 70% of the area but accommodates almost the same quantity of forest as the evergreen forests’ zone. Côte d’Ivoire: national scale Instead of presenting the statistics on the country as a whole, we have tried to show the values of deforestation according to the domains of vegetation and administrative regions, in order to present evidence on the distribution of the last forests. Forests are mostly situated in the southwest of the country (91% of the forest area) and in the evergreen forest zone (86% of the forest area). The forests in the moist semi-deciduous zone cover less and less area, even inside “forêts classées”. This is due to large tree plantations within their borders and to the importance of coffee-cacao plantations in the rural domain. Since 1926, when the French colonial administration started to determine forest areas reserved for forestry exploitation under the form of “forêts classées”, it has not been simple to follow the development of these areas. In fact “forêts classées” appear or disappear from various available lists with no official public notice of declassification (Dao 1999). Of the two categories defined by the 1978 forest decree, the rural forest domain (the area outside the

26

“forêts classées”) has without doubt suffered the highest deforestation. This is mainly the result of the absence of a legal status of these forests. Not all forests in that domain have been cut however. A certain number of forests which are not “forêts classées” still exist today, often as groves that are difficult to access or are unexploitable. In and around “forêts classées” the regulations against clearing for agricultural use, although in principle absolute, have not always been respected. As a result of this, more than half of the “forêts classées” area has been cleared. Numerous factors determine conservation: the existence of real management and protection projects (as for example in the Taï National Park and the Yapo forest), population pressure around the forests, accessibility and exploitability of the forests. For every forest, this mix of factors is different. The quality of the forests in Côte d’Ivoire, in terms of fragmentation measured by the sizes of the core areas, is weak as only 22% of the forest area has a distance of over 1000 m to the forest border. Regional scale One of the most interesting observations concerns the history of the distribution of deforestation: it runs contrary to the general belief that the longer the history of human occupation is, the more the lands are cleared. In almost all cases, the areas most wooded in 1958 have the lowest level of forest cover in 1990 and 2000, and the areas considered under cultivation in 1958 still have a significant forest cover in 2000. This phenomenon of differential development of deforestation is illustrated by the comparison of the east and west parts of the Abidjan region: in the western part, between Divo and Grand-Lahou, the forests have been transformed into plantations of coffee-cacao, palm or rubber trees, while the eastern part (Agboville) shows a landscape composed of a mosaic of numerous small fragments of forest (mainly secondary) and crops. We conclude that the rate of deforestation is related directly to the type of land use rather than to the density of the population. The lands situated along the road axes prior to 1958 were and are still occupied by populations that profit from and manage the forests and fallow fields in a sustainable way. In the peripheral zones, which were composed of dense forest in 1969, the lands have been occupied quickly and extensively with an aim of planting cash crops, often by non-indigenous populations who came from the northern regions of the country. The forests of these areas that are still wooded have certainly become depleted, as is the case in Yapo for example. Thus, the preservation of these forests is not due to the absence of exploitation or a weak need for agricultural land, but actually to the sustainable management of the lands. The percentage of deforestation between 1990 and 2000 varies largely over the administative zones (from 6 to 90%) and the overall statistic for the whole country does not give pertinent information. Overall deforestation is

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Figure 2.9 Causes of deforestation in Côte d'Ivoire. The scheme is based on the models proposed by Turner et al. (1995).

diffuse and touches even the large forest areas that remain in places with difficult access. Some parts in the South zone, on the other hand, show some very large areas recently deforested in order to plant rubber trees. This is particularly the case in the Gô-Bodiénou “forêt classée”. This supports our believe that the role of “forêts classées” as a biological reserve for species is rather weak and that currently their main role is that they are a source of exploitable wood. At the same level, we have shown that deforestation is much less significant in the Abidjan region than in the Taï region where people prefer to clear large portions of forest to appropriate them even if they will not cultivate them right away. Local scale We characterised the changes in the forest cover by analysing the size distribution of the forest areas. The eight blocks of 20 x 20 km under study all showed a significant area occupied by small forest tracts of less than 8 ha in size. These attain 30% of the overall forested area. The

assessment of these forest mosaics, which are typical in tropical landscapes, is essential because the presence or absence of such a fragmented forest cover leads us to the mode of land use and to the consequences of the maintenance of an environmental and ecological equilibrium. In fact, the almost total absence of forest in the Divo region shows that the lands are all occupied by agriculture, which explains that the Gô-Bodiénou forest (situated 20 km to the south) is now under extreme pressure of deforestation. In the case of the Yapo classified forest in the east, the presence of numerous fragments of secondary and “primary” forest leads us to suppose that human pressure is still weak since the manner of land utilisation is still relatively traditional. Unfortunately, the small forest areas in the rural domain, and to some extent in the domain of the “forêts classées”, are ignored in forestry assessment mappings, as is shown in Figure 2.7. This makes it difficult for forestry authorities to properly manage these areas. A proper mapping is needed and technically possible, especially at a local level but also at a regional level.

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the capacity for renewal of the environment and the role of climate change, whether from global causes or inflicted by modification of the vegetation cover in Côte d’Ivoire.

Figure 2.10 Changes in area under cultivation in Côte d’Ivoire. The principal commercial cultures of the forest zone are given.

2.

Causes of deforestation

The causes of deforestation in Côte d’Ivoire are many and play over various spatial and temporal scales. Figure 2.9, inspired by Turner et al. (1995), summarises the respective roles of different direct and indirect factors of deforestation in Côte d’Ivoire. Conditions of the natural environment The natural environment is a combination of conditions that make certain types of vegetation and certain crops possible (Figure 2.9 relation (1)). The actions of agents of deforestation (farmers, lumberjacks, etc.) are conditioned by the natural environment (the presence of species of trees sought, the abilities of the soils to grow cacao and coffee, etc.). Consequently, the forests that present few prospects for cultivation (infertile soils, sloping land, etc.) have been preserved for a longer time. The indirect influences (2) of the natural environment on deforestation are hard to identify. Even on a larger scale (3) this influence is limited. An example is formed by the presence of certain savannas in the southern forest which were residuals of paleoclimates. On the contrary, forests tend to take over the savanna, at least in certain regions like the V-Baoulé in the middle of Côte d’Ivoire, (Spichiger & Blanc-Pamard 1973, Gautier 1992b, Gautier & Spichiger chapter 3). We stress again the importance, for future changes in forested areas, of the role played by

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Direct causes of deforestation Technical advances: Modifications of the Ivorian landscape over the last 40 years have essentially been effected by woodsmen and farmers (6). In concrete terms, the colonisation of the forested south of Côte d’Ivoire, inaccessible for a long time, was made possible by the opening of new roads which permitted the shipping of merchandise to the export ports of Abidjan and San Pedro. In the case of Côte d’Ivoire, during the phase of accelerated forest colonisation, improvements in agrotechnology (5) permitted the establishment of large tracts of industrial plantations (palm, rubber trees). It did little, however, towards an improvement of coffee and cacao cultivation which remain relatively weak systems by international standards. The rejuvenation system of plantations is little developed and the plants are old today (Lee 1980). Expansion of existing plantations and development of new ones was preferred. Farmers as agents of deforestation: Farmers are part of a complex pyramidal system composed of three parts (Leplaideur & Ruf 1981). Firstly, native populations hold customary rights to the land and others must address themselves to them on arrival if they wish to settle. Secondly, non-indigenous people, Ivorians coming from other regions of Côte d’Ivoire, most notably the savanna regions (such as the Baoulé), often constitute the first wave of forest colonisation. Thirdly, foreigners, mainly from Mali and Burkina Faso, come to search for work, and start as employees with the hopes of starting their own plantations once they have accumulated a certain capital. The lightning-fast development of the plantation economy is due to the synergy of these three factors (CIRES 1979) (Figure 2.10). Plantations need large numbers of people to work on the land. The pyramidal system only works when large land reserves exist which permit the farmer to dream of the next instalment. The more farmers who wish to become planters, the more the need for manpower grows, and, by same count, the need for land. This system explains the phenomenon of the race for land and the exponential growth in conversion of forested areas to agriculture during the years 1960-70. Underlying causes of deforestation More fundamentally, however, these agricultural activities result from political choices, from general economic conditions, from a land system that allows the exploitation of the forest, and from the action of the State. Agriculture: Until the second half of the 20th century, Côte d’Ivoire had a low population density, especially in the southern forests. The traditional agrarian system of Côte d’Ivoire was one of subsistence agriculture, in the context of abundant space which permitted the regeneration of the forest. The main subsistence crops are

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rice (west and southwest), Chinese yam (centre and north), cassava (south coastal fringe), plantain banana and taro (southeast of the country). Since its independence in 1960, the economic policy of Côte d’Ivoire has been characterised by an “agrarian” option open to international commerce. In this policy, priority was given to cash crops destined for export (in first place wood, cacao and coffee). Cash crops were introduced in a progressive manner starting at the end of the 19th century. As S. Amin wrote (1971), “Côte d’Ivoire finds itself in 1950 still in the state of a colonial “reserve” where commercialisation has not yet begun.” Paradoxically, initially very few autochthonous farmers adopted this system. At present, this is hard to believe in the light of their success with the Ivorian farmers after 1950. The main commercial crops in the forest zone are cacao, coffee, pineapples, bananas, oil palms, coconut palms, rubber trees and cola nuts. Because of its expropriation of space and its growth (harvested areas multiplied by 5 between 1960 and 1990, Figure 2.9), commercial agriculture is among the principal causes of the disappearance of the Ivorian forest cover, in conjunction with forest exploitation. Exploitation of wood: The most precious wood for woodworking, the famous Bassam mahogany, has been exploited since 1880 (Arnaud & Sournia 1978). The boom in exploitation began at the end of the 1950s and achieved an absolute record in 1977 (Figure 2.11). At first it was the fine woods (such as mahogany) that were exploited, later more and more wood of lesser quality was taken as the forests were gradually depleted. Until the 1980s, most of the wood was destined for export in the form of rough timber (logs with bark still on). Over 85% of the demand came originally from Europe. The export share dropped to 80% in the 1960s, 64% in 1978, 56% in 1985, and 40% in 1987 (Ministry of the Environment 1991) due in part to a drop in demand but also to an Ivorian decision to promote the conversion of wood within Côte d’Ivoire itself. The role of the State: Besides the agrarian option chosen by Côte d’Ivoire many other factors have favoured the development of a “plantation economy”. The State created the myth of the citizen-farmer, symbolised by the figure of the president presented as the first farmer of the country. The value given to the role of the farmer is clearly expressed by the order given by the president as early as the beginning of the 1960s: “the land belongs to those who cultivate it.” At the economic level, the State controls the commercialisation and transformation of export crops through its intermediary the Ministry for Stabilisation and Support of Agricultural Products Pricing (CSSPPA) which fixes the price of products and the payment to those involved to regulate the national market. The CSSPPA constitutes the second revenue source of the State (after tax takings) and it is there that we find all its interest in the development of commercial crops. The State played an even more direct role as initiator and director of development projects at the regional level. Thus, the

Southwest Region Management Authority (ARSO) and the Bandama Region Management Authority (AVB) had considerable impact during the 1970s on the commercialisation of the central and southwest regions, by displacing populations, constructing large infrastructures and making agricultural technologies available. Likewise, the large national agriculture plans (for sugar, palm, rubber trees) aimed at the development of new products and the opening of entire regions. Land rights: The primordial element of the modern mode of production, Roman land rights, needed to be brought in at the same time as new commercial products appeared. After independence, and contrary to the other countries which engaged in socialisation of the land (like Senegal), Côte d’Ivoire conceived a judicial status of land according to the concept of private and liberal ownership. The Ivorian State sought to define a State Domain and a forestry legislation to regulate the exploitation of wood. Besides the conflict brought about by superimposing these rights onto traditional rights, modern land rights have not put a brake on deforestation. On the contrary, because the commercial use of the land was not only the principal criteria to attribute property rights, it is seen as the actual objective of land rights. The commercial use of the land is a condition of ownership: the Binger order of 1893 allowed concessions to be revoked in cases of noncommercialisation; the law of May 3 1946 made commercial use of lands previously acquired obligatory and permanent. As early as 1926, the system of forest classification was established with the creation of the Haut Sassandra Wildlife Reserve, in fact the first forest reserve of the French colonies (Parren & de Graaf 1995, Arnaud & Sournia 1978). It took until 1965, however, for a forestry code to be adopted. Forested areas are by default considered part of State Domain, although they are in principal “vacant and ownerless”. Forest exploitation in the permanent State Domain is subject to a temporary exploitation permit allocated by decree for a unitary area of 2500 ha (5 x 5 km) and for periods of 5, 10, or 15 years. Since 1978, the State Domain forest is divided into a permanent domain of the State (either “forêts classées” or forests awaiting classification) and a rural forest domain (space under economic utilisation). The establishment of legislation aimed at saving and keeping a forest cover has not prevented the clearing of forests (at times complete) which are nominally classified, even if a certain number of important tracts have been effectively protected until today (Taï, Yapo). In many cases, the law simply confirmed the state of affairs by declassifying forests that were overexploited or converted to agricultural land. Retroaction: the end of the race for land Today, the landscape is saturated in many regions of Côte d’Ivoire, an occupation marked by perennialisation and aging of the plantations and by a dwindling of wood reserves. At a social level, the lack of land inhibits the establishment of young farmers and possibly also

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Figure 2.11 Production of wood logs in Côte d’Ivoire: comparison of produced and exported log volumes.

Figure 2.12 Development of human population and of deforestation in Côte d’Ivoire.

constitutes a brake to the flow of migrants. The “modernisation” of land rights is not without conflicts with the traditional land system. In addition, in times of economic crisis, many people are drawn to agriculture as a means of survival, putting more pressure on the lands as yet unoccupied. At the scale of the entire system (9), the lack of equilibrium of the relationship with the environment that was prevalent during the 1950s is still in place. The relative weight of factors varies according to the time scale and the spatial scale considered. About a hundred years ago, one could distinguish several stages. Before the French colonisation, a system of traditional sustainable cultivation was present with an expansion of natural forest cover due to population decrease caused by tribal and precolonial wars. During the pioneer period of colonisation up until 1920, private landownership developed and new crops were imported. During the 1920-50 period, forest exploitation and the distribution of cash crops started, and the first forests were protected. The period 1950-80 showed a boom in coffee-cacao plantations and the establishment of a plantation economy. Farming was important in almost the entire forest zone, caused amongst other factors by a massive immigration. This lead to disappearance of the forest. During the post-1980 period, the exploitation limits of the ecosystem were reached, and the need to manage the new ecosystem developed. The future development remains unclear: forest regeneration, stabilisation, or disappearance? Until the 1950s, the main attack on the forest was forest exploitation itself. The boom in cash crops was still to come and the population density remained very low. Then the plantation economy started, and in its initial phase the growth of the population had a very negative impact on the forest. Later, population growth lost its impact, partly due to intensification of land exploitation, partly to measures of land management and development.

3.

Population and deforestation

Population growth appears at the centre of all the mechanisms leading to the Ivorian deforestation. Demographic development is both an exogenous factor in deforestation (an external independent variable), and an endogenous factor, in the sense that deforestation in Côte d’Ivoire provokes a demand for the abundant manpower necessary for the cultivation practised, and therefore an influx of population which in turn accelerates deforestation. The population of Côte d’Ivoire has been multiplied by 5 during the 1960-2000 period. This growth is especially high in the forest zone, where density has gone up from 12 to 75 inhabitants per km2 during that period. Until the 1950s, both population density and rate of clearing of the forested zone remained at a very low level (Figure 2.12). Beginning with the boom in coffee-cacao, these two phenomena intensified strongly. Their development is not the same, however. Until the 1980s the increase in clearing is much stronger than that in population density. The population density continues to increase and the deforested area (as a percentage of the total land) does not increase further after the 1980s. The maximum deforestation rate was in 1973. After a population density of around 60 inhabitants per km2 deforestation does not increase anymore. Population density continues to increase, however, as a result of intensification of land exploitation. In this new saturation phase, we need to examine more closely the pressures on the remnant forests. The regions with the highest levels of clearing (the Abidjan agglomeration with 3 million inhabitants in 1994, west of Dabou, and the Divo region) are also those which have the highest population density. This is not the case in the sub-regions situated to the northwest of Agboville (the northern part of Tiassalé, Rubino, and M’Batto), areas with a high deforestation rate but a relatively low population density. These last areas had a

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strong population growth (10% per year) during the years 1955-75. After the 1970s, however, this growth moved towards the southwest of the South region, most notably towards Fresco, which then had a record population growth (10% per year, compare to Aggboville: 3%). This shows that not only the population density plays a role, but also the intensity of population movement.

4.

What can we do?

At the scale of West Africa: All countries in West Africa show a lack of recent cartographic information, but this is stronger for the countries which have been at war for years such as Liberia or Sierra Leone. We need an update of information on the forested areas, and on the state of conservation and delimitation of the protected areas. Such updated maps should have a sufficiently high resolution (100-300 m?) to allow us to distinguish the land-use mosaics and plantations, which currently pose a big problem for a realistic estimate of the area. The zones that get priority should then be mapped at a higher resolution. Currently, the FAO is making a huge effort, but it is necessary that the countries concerned improve the available information (Matthews 2001). With the availability of more extensive data processing and more affordable image prices, detailed cartographic observations should be possible. At the national scale: “forêts classées”. It is urgent that the government of Côte d’Ivoire develops a real policy of conservation regarding the “forêts classées” in the country, even though only 23% of the remaining forest in the country is within these “forêts classées”. This study of the development of deforestation shows that conserving forests by the creation of “forêts classées” or “botanical reserves” is not enough, aside from a few exceptions such as the Yapo forest or partially in the Irobo forest. In fact, crops have replaced the large majority of the forests. For a real policy of conservation it is necessary to encourage farmers to invest in their plantations (e.g. rejuvenation) and to forbid agricultural practice in “forêts classées”. The PAM program for relocation of farmers has not shown results because of mismanagement of the budget available for that. Also the primary forests managed by the European Union’s “coastal forests management project” show continued deforestation. For the survival of the remnant forest fragments, classified or not, a better exploitation of agricultural lands in the surroundings is needed. In fact, subsistence farming and large extensively used plantations, both huge consumers of space and manpower, should improve their yields. Many areas left fallow are practically unused. The question of property rights should also be solved. The conflict between modern and traditional rights as well as the principle of land ownership acquired by clearing are both obstacles in a lasting land management.

At the local scale: community forests. At the local level, improvements are needed in the village socioeconomic infrastructure, support for the development of more intensive systems of planting, and a re-assessment of subsistence cultivation - for a long time devalued in favour of the system of export cultivation. The rural areas have almost 76% of the forest cover. At present, these forests are ignored on maps and in the national statistics because they are very fragmented. Villagers who invest in forest conservation have no property rights according to the constitution and to the manual of forestry practice. They can do nothing against the continued extraction of large amounts of wood by logging operators who penetrate into community plantations and forests with the authorisation of a logging permit. We can cite as an example the case of some community forests in the Taï region (Bonnéhin, 2000), where the attempts of villagers to conserve them has been put in peril by the logging operators. We strongly suggest that the State establishes a policy of conservation based on community management. Villagers who conserve the forests need to get the necessary rights to accomplish this conservation action.

Conclusions At all levels, the rate of deforestation has diminished since the years 1970-80, which was the period of massive exploitation. However, the problem of deforestation is getting more severe because the total forested area continues to diminish. Using deforestation rates instead of the actual forested areas in analyses of deforestation problems (as did Fairhead & Leach 1998) is for this reason a mistake. At the scale of West Africa, according to the sources used, we estimate that between 20% and 50% of the forest cover which existed at the turn of the 19th century remains. This is between 46,324,000 ha in 1995 (SOFO 1997) and 41,594,000 ha (FAO 2000). That these overall estimates are gross and at times totally wrong is shown when the information for each country is taken into account separately. These errors come from the different methods used in the calculations for the years 1990, 1995 and 2000. For tropical Africa, the estimates of forest cover changes between 1980 and 2000 vary more than the actual changes observed during that period (Matthews 2001). Fairhead and Leach (1998) have shown that the total forested area at the beginning of the century most probably was over-estimated, which has contributed to a false view of deforestation levels. Liberia and Côte d’Ivoire will be the principal actors in putting a conservation program into place, as these countries have most of the remaining forests.

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In almost all countries the moist semi-deciduous forest is the most deforested one. These forests thus need extra preservation. In Côte d’Ivoire, deforestation between 1969 and 1990 was the highest of the West African countries since more than 80% of the forested cover disappeared. With the deforestation the remaining large forest tract of Taï National Park thus has a increasing importance, both at the national level (20% of the Ivorian forest) and at the scale of West Africa. Although we cannot retrace the steps precisely with available maps, the statistics on forest exploitation in Côte d’Ivoire all confirm that disappearance was highest between 1977 and 1984 (Arnaud & Sournia 1978, Parren & de Graaf 1995). The zones with the highest deforestation levels are not those surrounding large agglomerations such as Abidjan, but those in the centre of the country, in the moist semideciduous forests, where the flora is made up of a large number of exploitable species. Among the countries of West Africa, Côte d’Ivoire possesses an important reservoir of protected areas. There are numerous “forêts classées” and they represent a considerable area, in particular in the region of Abidjan. But their importance for wood production and as a biological resource is doubtful, because in many cases they are forest only in name (77% of their area is not forested). Despite the almost complete deforestation of many of these “forêts classées” they have played a role in the conservation of the natural setting, because the most cleared regions, such as the central regions of GagnoaSoubré or Daoukro, are characterised by the absence of these protected areas.

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The analysis of the processes of deforestation in relation to the protected areas only represents a part of the problem. The causes and the solutions have to be found at the heart of the plantation economy, that is to say, the agricultural policy, the system of extensive production, and the land right system. While at the national level we analysed the distribution of the large forests, at the local level we identified the small forests (2 to 100 ha), which represent 10 to 30% of the forested area. A local approach is the only one that would permit an evaluation of the scope of deforestation and could link that to forest management. Fortunately, increasing management policies are focused on rural forest. This brings the small forested fragments into consideration, and the conservation of the flora and fauna associated with them.

Acknowledgements

We would like to thank the “Centre Suisse de la Recherche Scientifique en Côte d’Ivoire” with all the people working in this institute, and the “Fond National Suisse de la Recherche Scientifique” who financed our first study from 1989 to 1994. We are grateful to the anonymous reviewers for their help to improve the draft, to the hard work of the translators and specially to Alisdaire Menzies for his help in solving some translation problems.

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The forest-savanna transition in West Africa

3

L. Gautier and R. Spichiger

H A P T E R

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Introduction

Forest-savanna transitions occur in a variety of localities around the tropical world. Furley et al. (1992) provide an extensive review of the processes at the forestsavanna boundary, its dynamics, the role of physical factors, environmental change and vegetation history. However, apart from Nigeria, regional accounts concerned mainly Eastern and Southern Africa, Australia, and the Neotropics. We present here a review on the topic for West Africa. Distribution of vegetation types in West Africa is tightly bound to the climatic gradient from the shores of the Gulf of Guinea north to the Sahara (Bongers et al. chapter 4). In the absence of major orographic features, this gradient is mainly oriented on a south-north direction with the associated vegetation types forming latitudinal belts: from the lowland evergreen rainforest to drier vegetation types like dry forests, the mosaic of woodlands and savanna, and further north to a Sahelian pseudosteppe and finally desert. Phytogeographically, western Africa is the home of two main regional centers of endemism. The lowland rainforests in the south correspond to the western block of the Guineo-Congolian centre of endemism, whereas the savannas, woodlands and dry forests north of c. 9° latitude belong to the western part of the Sudanian centre of endemism (White 1983). Physiognomically and floristically, most steps in this transition are gradual, like the one from evergreen to semievergreen lowland rainforest, or from Guinean to Sudanian savanna and woodlands. On the contrary, the transition from semi-evergreen lowland rainforest to Guinean savanna is very sharp and can be unambiguously mapped at the continental scale by a solid line separating two very distinct landscapes (Figure 2.2). The multistrate closed forest which dominates the whole toposequences on the southern side abuts on a very different and more open landscape to the north, dominated by annually burning vegetation types. These vegetation types share a dense herbaceous layer made mainly of hemicryptophytic gramineous plant species, but whose tree cover may vary widely, ranging from almost pure grasslands, through savannas, to woodlands. In this open landscape, forest elements are limited to a network of gallery forests on the bottom of the slopes, but sometimes also to plateau forests

Figure 3.1 An example of a forest-savanna mosaic: Reserve de Lamto, south of the V-Baoulé, central Côte d’Ivoire.

on hilltops (Figure 3.1). Their structure and floristic composition is very similar to those of the adjacent drier types of the lowland rainforest. In this chapter we describe the forest-savanna boundary at different spatial scales, and how its location has shifted in the past. We discuss the main environmental factors that explain the present distribution of the two vegetation types in West Africa, such as climate, substrate, fire, and men. Finally, we describe the present trend in vegetation dynamics under natural conditions and under the growing influence of man.

The forest-savanna boundary at different spatial scales At the sub-continental scale, the sharp transition between the two landscapes occurs at a latitude of c. 8°N, with notable exceptions in central Côte d’Ivoire, where savanna penetrates as far south as 6°N in a region called “V-Baoulé”, and in Togo and Benin, where a savanna dominated-landscape reaches the coast. Even now that deforestation has cleared most of the lowland rainforest,

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A

B Figure 3.2 Forest-savanna mosaic in Lamto, Côte d’Ivoire. A) the forest-riparian type, B) the savanna-riparian type.

the transition is still obvious when looking at classifications of low-resolution satellite images (Figure 2.2) and these images still reflect the shape of the limit of the previously continuous rainforest zone as mapped for example by White (1983). In a latitudinal belt from Senegal and Sierra Leone eastwards to Uganda, there is a zone where savanna dominates with dense humid forest elements scattered in the landscape. It crosses all West Africa as a strip of 100 to 500 km wide and has been referred to as the “forestsavanna mosaic”, “secteur préforestier”(Monnier 1981) or the “Guineo-Congolian/Sudanian transition zone” (White 1983). This terminology underlies a very important

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characteristic of this landscape: two distinct vegetation types intertwine, as well as two distinct floras, while few species occur on both side of the forest edge (Aubréville 1966). This interpenetration of forest and savanna is evident when looking at aerial photographs or maps at the scale of 1:50,000 (Blanc-Pamard 1979). Swaine et al. (1976) make the distinction between the “forest riparian type” and “savanna riparian type”. In the former strips of middleslope savanna penetrate forest which persists on hilltops and along streams (Figure 3.2A) whereas in the latter the savanna dominates the whole of the topography except for fringing forest by streams (Figure 3.2B). It should be added that included savannas can also be found deep inside the continuous forest zone. At higher resolution, in the field, the transition between forest and savanna is very clear-cut (Blanc-Pamard 1979) (Figure 3.3A). It is only at this last level of analysis that the term “forest edge” makes sense. Most authors stress the abruptness of the change at the field level (Morgan & Moss 1965, Swaine et al. 1976) and define the forest edge as a narrow line where the domination of savanna grasses stops. Spichiger (1975) points out, however, that the forest edge should be interpreted as a transitional belt of varying width where species of the two floras coexist and compete (Figure 3.3B). Detailed field studies of the forest-savanna transition in Côte d’Ivoire by using transects perpendicular to the forest edge have allowed Spichiger (1975) to go beyond a simple forest/savanna species dichotomy. The GuineoCongolian forest species have been classified in four groups of increasing tolerance to the savanna environment based on the occurrence of the species in the transects relative to the position of the boundary and its sharpness. Accordingly, the Sudano-Zambesian species have been classified in four groups of increasing tolerance to the forest environment. The fourth Sudanian group, which consist of species that are commonly found inside the forest, surprisingly include species of dry forest that are otherwise found only much further north in the Sudanian zone. They could thus be considered as relicts of ancient dry forest nowadays absent from the transition zone due to the impact of more frequent fires.

The impact of palaeoclimatic changes

The presence of coastal savannas c. 100 km south from the savanna belt in Côte d’Ivoire (e.g. in Dabou, Grand-Bassam or Grand-Lahou) and the persistence of rainforest outliers in the north of V-Baoulé (e.g. in Bamoro), indicate that the northern limit of the continuous Guinean rainforest has been fluctuating during the late Quaternary. Aubréville (1949) and Mangenot

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(1955) hypothesised that these fluctuations could be congruent with the glaciations of the northern hemisphere, with cooler episodes corresponding to a drier climate (Aubréville 1962). With the development of various techniques such as palynology, sea level interpretation, diatoms and phytolithes, many data have been accumulated on past vegetation and climatic conditions and have confirmed these hypotheses. With the exception of Lake Bosumtwi in Ghana, there is a lack of paleobotanical data in West Africa due to the scarcity of lakes in which pollen accumulation can be studied. Reconstruction of paleoclimate is thus difficult and we have to refer to the general scheme for whole tropical Africa, as summarised below (Maley 1990, Thomas & Thorp 1992): • From 30,000 to 12,000 years BP: a dry and cold phase, with its maximum between 20,000 and 15,000 years BP. The climatic swing has been estimated to be c. 8° latitude to the south. At this time, forest in Africa was restricted to a limited number of refuges, the main one in West Africa being located between present Liberia and Guinea (Maley 1989, Wieringa & Poorter chapter 6). Pollen types have been found of Afromontane species such as Olea hochstetteri or Podocarpus sp. (Frédoux 1994, Salzmann 2000). These species are nowadays absent from West Africa and only found on Mt Cameroon, indicating that this drier episode was also cooler, the temperature fall being estimated to be (3) 4-6 (7)°C. • From 12,000 to 7000 BP: humid phase, extension of the forest to approximately its present limits. • From 7000 to 3500 BP: warmer and wetter climate than at present. The forest extends to its maximum range. • From 3500 BP to date: progressive decrease of temperature and humidity to meet its present values, the forest retreats to its present position. It is worth mentioning that another dry maximum event has been documented for Central Africa c. 3000 BP. During the last millennium the forest has been in another phase of expansion (Vincens et al. 1999). From work on lake levels in Lake Bosumtwi (Maley 1987) and recent unpublished work in the Dahomey Gap, it appears that this dry episode also took place in West Africa (Salzmann & Neumann, unpublished results). Palynological data have a rather coarse temporal resolution. This has to be taken into account when one tries to bridge the gap between these data and more recent historical records. For the last centuries, it is not clear whether an observed afforestation trend reflects a minor oscillation (decadal or secular) or can be fitted into the general scheme presented above (millennial). These paleoclimatic oscillations and the subsequent displacement of forest and savanna have implications at several levels. First, it can explain the relative low diversity of West African forest flora in comparison to Central Africa; many species could have become extinct during the dry periods. Second, it gives very interesting insight into

A

B Figure 3.3 The forest-savanna mosaic as seen from the ground in Lamto, Côte d’Ivoire. A) A clear-cut forest edge between a gallery forest and a herbaceous savanna, B) a smooth forest edge between a plateau forest (back, right) and a dense woody savanna (front, left).

the connections between Upper Guinea and the Congolian basin during humid periods, which might explain the disjunct distribution of certain species (see also Holmgren et al. chapter 7). Third, the present separation by the Dahomey Gap and the past separation of the Upper Guinea forest block itself by the extension of VBaoulé down to the sea could have led to allopatric speciation resulting in sister species on both sides of the separation (see Kouamé et al. chapter 5 for implications for species composition). Fourth, the present extension of the Guinean savannas inside areas with climatic conditions favourable to forest can certainly be interpreted as the resilience of past distribution. A recent modelling study has shown that both savanna and forest could persist under the same conditions. By varying the dry season precipitation the model shows hysteresis between forest and savanna (Da Silveira & Sternberg 2001).

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Main determinants of forest and savanna distribution Present distribution of forest and savanna can be interpreted by looking for parallels with environmental conditions at various spatial scales. Among them, climate and edaphic conditions determine water availability and are of paramount importance in the competition between the two vegetation types. The outcome of competition is modified by disturbances such as fire and shifting cultivation. In the following paragraphs we discuss these main environmental factors, and how they operate at different spatial and temporal scales. The ecology of savanna per se is far beyond the scope of this chapter and can be found elsewhere in the literature (e.g. Huntley & Walker 1982, Bourlière 1983, Walker 1987). Climate There is a general agreement that climate is the primary factor explaining the distribution of forest and savanna, at least at a continental scale (Adejuwon 1971, Swaine 1992, see also Furley 1992 for the Neotropics). The main limiting factor for the presence of forest is water availability, often expressed in terms of total annual precipitation and the length of the dry season. For instance, Aubréville (1962) states that the lowland tropical forest is mainly found in areas receiving more than 1400 1500 mm of annual rainfall, but that this limit can be lowered to 1250 mm if the precipitation is evenly distributed throughout the year. A place with a dry season of 1 - 3 months (with monthly precipitation below 30 mm) can support forest depending on annual rainfall and soil type. According to Adejuwon (1971), the forest-savanna ecotone in western Nigeria is comprised within the 80 100 rainfall days isopleth, and the 1250 mm mean annual rainfall isohyet. The 25 mm mean annual water deficit and the 1300 mm mean annual evapotranspiration isopleths all have the same shape and pass through this zone. Substrate Edaphic conditions are of paramount importance in determining the forest-savanna equilibrium at a regional scale especially in areas where climate conditions are intermediate. In west-central Ghana, the limit between continuous forest and forest savanna mosaic is congruent with substrate: savanna dominates on Voltaian rocks and the forest dominates on the Birrimian basement complex, on which soils are richer in clay and nutrients (Swaine et al. 1976, Swaine 1992). A similar situation is found in Côte d’Ivoire, at the eastern side of the V-Baoulé where the boundary between continuous rainforest and forestsavanna mosaic corresponds to the boundary between schistous and granitic substrates, the former giving rise to soils being also richer in clay and having thus a better water-holding capacity (Spichiger 1975).

36

At the local scale, most authors agree that soil types (which are generally distributed repeatedly along successive toposequences) are of primary importance to explain the patchy distribution of forest and savanna (Morgan & Moss 1965, Aubréville 1966, Latham & Dugerdil 1970, Adejuwon 1971, Furley 1992). Savanna inclusions are always found on sandy, phosphorus-deficient soils, or very shallow, waterlogged soils (Adjanohoun 1964). In westcentral Ghana, vegetation types were found to occupy distinct catenary positions and to be associated with characteristic soil types. Grassy, treeless savannas are often found on alluvial soils that are waterlogged in the wet season and lack water in the dry season. Levels of total phosphorus, exchangeable calcium and potassium, water content, and pH in the topsoil were all found to be higher under forest than under savanna. These differences in soil fertility, which correlate with tree cover and stature, were however suspected to be of secondary importance. The primary factors separating vegetation types are probably effective soil depth, soil texture and water supply (Markham & Babbedge 1979). Soil properties can also be transformed in feedback with the vegetation above it, by changes in soil fauna (e.g. termite species) and by cultivation and settlements (Fairhead & Leach 1996, 1998). Fire Fire is a key element in understanding the distribution and dynamics of savannas and the transition zone with forests. Although natural fires occur through ignition of the herbaceous layer of savannas via lightning, sparks from falling rocks and fermentation (Phillips 1974), almost all savanna fires are currently anthropogenic of origin. Indeed, man’s use of fire has led to an increase of fire frequency in these systems which can hardly be considered natural and each year savanna fires run over almost the whole surface of the transition zone. However, the existence of fireadapted flora confirms the importance of fire as a system determinant, even before man mastered fire (see also Bond & Van Wilgen 1996). Reviews on the effect of fire on the savanna ecosystem can be found in Phillips (1974), Gillon (1983) and Trollope (1984). Forest pioneer species whose propagules originate from the forest elements scattered in the landscape are very aggressive in invading savannas and are frequently found as seedlings or saplings. These are generally found within a few metres from the forest edge but also sometimes several hundred metres apart depending on their dispersal mode (anemochory or zoochory) and site characteristics (longdistance establishment of forest species occurs in savannas with a high tree cover) (Spichiger 1975). Several long-term experiments on the effect of fire have been initiated in the transition zone of West Africa. In Kokondekro, near Bouaké in Côte d’Ivoire, three 1-ha plots have been subjected to different fire regimes since 1937 (Figure 3.4). Annual early burning (beginning of the dry season) has promoted a shrub savanna (Figure 3.4A). Annual retarded burning (mid to late dry season) has

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promoted a savanna with a very loose shrub cover because of a greater fire intensity (drier fuel) and the fact that it destroys tree leaves that establish long before the rainy season (Figure 3.4B). Integral protection from fire has led to a genuine forest (Figure 3.4C) composed mainly of dense semi-evergreen (Guineo-Congolian) forest species (Monnier 1981). A similar experiment is being carried out in Lamto (south of the V-Baoulé) where a c. 60 ha savanna has been protected from fire since 1962. The results clearly show the same trend, although vegetation succession is in a less advanced stage (Vuattoux 1970, 1976, Devineau et al. 1984). In northeast Ghana, another fire protection experiment has led to a clear decrease of grass cover and a clear increase in tree number and stature, but on contrary to the Kokondekro experiment, there has been no establishment of Guineo-Congolian forest species. This is explained by the much more northern position of the plot, north of the transition zone, in a region where dense forest elements are very rare and thus forest seeds unavailable (Brookman-Amissah et al. 1980). A reverse experiment has also been conducted. There has been an attempt to create a savanna inside the forest zone at Adiopodoumé, south Côte d’Ivoire, by felling and burning the original vegetation, sowing savanna species and burning the vegetation regularly. The result was a relative failure: there was a massive colonisation by ruderal forb species, which made it difficult to ignite fire and burn the vegetation (Adjanohoun 1964). There is a general agreement that the savannas of the transition zone are not in equilibrium with climate and are a direct consequence of annual fires. The climatic climax would be semi-deciduous rainforest and further north (from c. 9°N) dry forests and woodlands. Guinean savannas are considered to be of secondary origin. This is also reflected in names such as: “savane de substitution” or “derived savanna” (Keay 1959). They are to be considered as a “fire-climax”. It has been suggested by Aubréville (1966), Monnier (1981), and Swaine (1992) that before the massive use of fire by humans, the transi-tion was much more gradual; “in a primitive state there were quite extensive woodlands […] intermediate between moist forest and myombo woodlands” (Keay 1959). From the evidence accumulated above it is clear that fire plays a very important role in the resilience of savannas to invasion by forest species. It is also clear that fire reduces the density of savanna trees. The fuel biomass is lower under the shade of the tree crowns, and forest pioneer species have therefore better chances to become established. Fire also favours the herbaceous layer of savanna by promoting the resprouting of perennial grasses and the germination of annual savanna species (Hopkins 1963). In turn, a dense herbaceous layer increases fuel availability and thus dry season fire intensity. Fires can also occasionally sweep under the canopy of the drier types of rainforest which are to be found in direct contact with burning savannas (for Ghana, see Swaine 1992). These fires are generally low in intensity

A

B

C Figure 3.4 The Kokondekro experiment (central Côte d’Ivoire) after 55 years. A) The early burning plot, B) the late burning plot, C) the protected plot.

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and are essentially fuelled by litter (ground fires, Phillips 1974). They extend to the canopy only during exceptionally dry conditions (Swaine 1992). Like logged forests, burnt forests are more prone to renewed burning (Hawthorne 1991). In Ghana, fires have recurred in several years since the main outbreak in 1983. Consequently, regrowth becomes predominantly herbaceous, but it is uncertain if such continued pressure will convert forest to true savanna (Swaine 1992). Animals Animals can have an impact on the forest-savanna transition through pollination and the dispersal of plant species, but also through seed predation, browsing, and the regulatory role of top predators on herbivores (Medellin & Redford 1992). Other vertebrates and especially birds play also key roles in these processes. In Africa, large populations of herbivores can open up the landscape (Kortland 1984). Soil fauna is also of great importance. Termites have a large influence on nutrient cycling and soil displacement. Their mounds are often preferred sites for establishment of woody species (Abbadie et al. 1992, Howse 1992). Predation on colonies by specialised mammals or ant species ultimately result in mound decay and release of nutrients to the soil. Anthropogenic influence Apart from the intensification of the fire regime, there has been considerable debate on the impact of human activities and especially traditional shifting cultivation on the distribution of forest and savanna. It was generally assumed that shifting cultivation would lead to an extension of savannas by destroying woody vegetation (Aubréville 1966). In South America slash and burn agriculture, fire and deforestation have led to savannisation (see for example Kauffmann & Uhl 1990, Uhl & Kauffmann 1990, Miller & Kauffmann 1998). However, in the Rio Branco-Rupununi region of the northern Amazon, forest regrowth generally occurs along the forestsavanna boundary, in spite of persistent shifting cultivation (Eden & McGregor 1992). Field studies in Côte d’Ivoire, have even revealed that shifting cultivation at the forest edge actually promotes afforestation (Spichiger & Blanc Pamard 1973); field preparation reduces herbaceous domination in the fallow, and thus limits fuel availability. As the impact of fire is reduced, forest pioneer species can establish. Similarly, in southwest Togo, forests regenerate as well in savanna fallows (Guelly et al.1993). Studies in heavily populated countries like Nigeria have widely questioned the origin of the derived savannas (Keay 1959). It has been suggested that at Ibadan, some 50 km inside the rainforest zone, savanna is derived from secondary forest. “In the absence of continual bush clearing for cultivation it is doubtful if fire alone could maintain the savanna indefinitely” (Clayton 1958). Aubréville (1966) was also convinced that cultivation (and

38

fire) could explain the present distribution of savannas, like for instance for the southern extension of savannas in central Côte d’Ivoire (V-Baoulé). Monnier (1981) argued that this was not likely, given the low population density and the kind of tools that were available. Nevertheless, in some places like the western extension of the V-Baoulé, there is general agreement that the Pennisetum purpureum savannas are derived from cultivation of forests (Adjanohoun 1964). Fairhead and Leach (1995, 1996) have widely questioned the interpretation of the environmental history of the forest-savanna mosaic. According to the colonial and post-colonial governments in Guinea, the once continuous forest over the region of Kissidougou was cleared by local population and Malinke migrants. Based on oral and historical sources they demonstrate that this interpretation is not true. Instead, most of the human activities are promoting afforestation (farming system, deliberate planting of forest species in fallows, pasture, fire-breaks). Based on aerial photographs, they show that this has led to a 40% increase in forest cover during the last 50 years. In the northern part of the mosaic in Ghana, studies have demonstrated that even under an increasing population, forest losses and gains were in a kind of steady-state equilibrium (Amanor 1993, Afikorah-Danquah 1997, both cited in Leach & Fairhead 2000). Increase in population pressure as well as transformation of farming systems toward cash crop production has completely altered the intensity of human impact on vegetation during the last decades. Comparing aerial photographs in the north of the V-Baoulé in the years 1952, 1969, and 1975, Spichiger and Lassailly (1981) indicate that during the first time interval, vegetation changes expressed its “internal dynamics” with a tendency towards afforestation. During the second interval, the area of cultivated land increased at the expense of both savanna and forest. Especially, the duration of the fallow in the farming system is of paramount importance. If the fallow period is reduced from c. 10 years to c. 5 years, as is experienced in many places in the forest-savanna transition zone, then afforestation processes can no longer take place. Large-scale deforestation in the northern part of the forest zone through logging, charcoal production and slash and burn agriculture can dramatically enhance fire hazard, especially when one takes into account the subsequent alteration of microclimatic conditions. Colonisation by grasses is likely to occur, further increasing fire hazard. Such processes have been studied in the Amazon basin (Uhl & Kauffmann 1990). It is uncertain if such processes take place in West Africa. Attempts to correlate occurrence of fires with broad-scale land cover changes using remote sensing data did not give conclusive results (Ehrlich et al. 1997). As pointed out by Leach and Fairhead (2000), “population-forest relationships [are] variable, non-linear and unpredictable”, especially taking into account changes in population pressure, the evolving strategies and

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environmental global changes. It makes no sense to try to explain present distribution of forest and savanna by one of the above factors only. All published works point at the importance of the combination of factors. There are even secondary effects of vegetation on soil at the local scale or on climate at the global scale that are very difficult to take into account. Distribution of vegetation is also of primary importance on human choice of where to set a field or where to build a village (Adjanohoun 1964). One can only get an idea of the effect of a single factor on the forestsavanna boundary, as all factors are intimately intricated in a way that their respective weight depends on the others. Furthermore, it has been widely recognised that historical factors must be taken into account to explain the present distribution of forest and savanna. Climate and human pressure have not been constant in the late quaternary and vegetation tend to show considerable resilience towards environmental changes.

Present dynamics of the forest-savanna boundary Present environmental conditions are favourable for afforestation, even when the effect of fire is taken into account (Aubréville 1966). This has been demonstrated by comparing two sets of aerial photographs separated by 25 years in Lamto reserve, Côte d’Ivoire. Despite the occurrence of annual fires, forest surface is increasing both by direct progression of existing forests and by the appearance of forest islets within the densely wooded types of savannas found on hilltops (Gautier 1989). At the same time, the woody cover of all savanna types has increased, except the herbaceous savannas downslope where soils are too waterlogged for the woody savanna species (Gautier 1990). Given the absence of cattle and the low density of indigenous herbivores, these results can not be attributed to overgrazing and a subsequent reduction of fire intensity, and are hence believed to reflect an internal trend. It can not be excluded, however, that the regular fire regime associated with the management of the reserve has reduced the destructive effect of occasional late season fires. Similar results have been documented in South Benin (Profizi 1982). Near the limit of the continuous forest, to the extreme tip of V-Baoulé, some small savanna patches can be found with the palm Borassus aethiopum and savanna trees, but completely overgrown by forest pioneer species (Adjanohoun 1964). Based on floristic transects perpendicular to the forest edge, Spichiger (1975) has developed several indexes to quantify the intensity of forest progression. He showed that forest was encroaching on savannas especially on the edge of those hilltop forests which are in direct contact with savannas having a dense woody cover.

These results seem to point out that the expansion of forests recorded in Central Africa since c. 1000 years (see above) might also be relevant for West Africa. However, it should be remembered that among major climatic changes there have always been minor oscillations. Because of a lack of more detailed pollen records and more ancient meteorological data, we can only hypothesise that the last centuries have been characterised by climatic conditions promoting afforestation. However, savanna displays resilience towards afforestation. Present distribution of vegetation compared with edaphic conditions as well as fire-protection experiments point at the role of these factors in slowing down the process. As indicated above, changes in demography and socio-economic conditions have drastically altered the environment during the last decades. Population increase and cash crop development have considerably shortened the fallow period, especially in the transition zone where soils suitable for cultivation are scarce. Deforestation has a tremendous impact on the microclimate, and the generalised deforestation of West Africa has probably already altered the global climate of the region. Chromolaena odorata, a Neotropical species, has considerably spread over West Africa since c. 50 years (Gautier 1992a). The introduction of this exotic weedy forb species has led to a radical transformation of the forest-savanna equilibrium. In Côte d’Ivoire, Chromolaena odorata invades forest margins, included savannas, hilltop wooded savannas, logged forests and savannas protected from fires and is now considered to be a key species in the transition zone between forest and savanna (Gautier 1992b, 1994, 1996). The weed suppresses the fire-tolerant savanna species during wet and normal years, but hampers the colonisation by forest species which establish very slowly under the dense thicket. In dry years, Chromolaena odorata dries and becomes very flammable, resulting in intense fires which destroy the few colonising forest elements as well as the fringe of the established forest. As the weed resprouts easily from its rootstock, the whole process leads to a global increase of Chromolaena odorata thickets with low diversity in the landscape. There is an urgent need to re-examine the vegetation dynamics in the forest-savanna transition zone, and to develop new policies addressing the questions of fire regime and land use. It is for instance of primary importance to allow the use of regular dry season fire in all types of savannas to avoid the encroachment by Chromolaena odorata. Infested areas in proximity of savannas should be surrounded by firebreaks thus allowing for a slow succession towards forest. This succession will be much more rapid if there is a pre-existing tree cover, because Chromolaena odorata is a strictly heliophilous species. Removing the weed and its stumps could even accelerated the process of afforestation.

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Conclusions

The present distribution of forest and savanna in West Africa appears to be the consequence of complex interactions between climate, edaphic conditions, and anthropogenic activities such as cultivation and fire. At the sub-continental scale, climate is of primary importance but its effect on water availability is mitigated by local edaphic conditions. The present position of the boundary is also a legacy of past climatic conditions, because vegetation types have an internal stability resulting in delayed response to changes in the environment. At the landscape scale, the forest savanna mosaic is primarily determined by soil conditions. In addition, the natural colonisation potential of forest species is hampered by fire. The human induced increase in fire frequency can probably explain the sharpness of the transition between

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the two vegetation types. Cultivation has probably favoured afforestation when human population densities were low and fallow periods were long. Cultivation has led to savannisation when population densities were high. Under present, natural conditions the balance between the two vegetation types appears to be shifting towards forest. However, due to the growing impact of man, the future of the forest-savanna transition zone is uncertain and should deserve much attention.

Acknowledgements

The authors would like to express their thanks to the Swiss Fund for Scientific Research who has supported their field work, and to two anonymous reviewers who have contributed much to the improvement of this chapter.

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H A P T E R

The forests of Upper Guinea: gradients in large species composition

4

F. Bongers, L. Poorter and W.D. Hawthorne

C

4

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Introduction

The forest belt in Upper Guinea stretches over 2000 km from Senegal in the west to Togo in the east. Variation in floristic composition of these forests is mostly gradual although most vegetation maps are based for practical reasons on vegetation types and focus on classes rather than gradients (e.g. Taylor 1960, Mooney 1959, Guillaumet 1967, Guillaumet & Adjanohoun 1971, White 1983). Generally the vegetation of the region has been described in terms of the most conspicuous (mostly in terms of abundance) species that compose a vegetation type (e.g. Chevalier 1909, Schnell 1952, Taylor 1960, Mangenot 1955, Aubréville 1959, Voorhoeve 1965, Guillaumet & Adjanohoun 1971). Later studies have used mathematical techniques to classify the vegetation types (Hall & Swaine 1976, 1981 for Ghana, de Rouw 1991 for the Taï region, van Rompaey 1993 for southeast Côte d’Ivoire - Fig 4.1 - and east Liberia). In general, information on the forests of the Upper Guinea is scattered and available at a local level only, and international level vegetation descriptions and vegetation maps are scarce (except for Africa as a whole, e.g. White 1983). In this chapter we present a vegetation map for the whole Upper Guinea region. A common classification allows for better comparisons between forests in different countries and a better exchange of information. For such a map we cannot simply combine the existing maps because of differences in criteria used, resulting in different vegetation types. For the present study we used data from forest inventories. Several types of data are available, from large-scale forest inventory data, collected by national forest services, to smaller scale data collected by individual researchers, or research groups. The large-scale data are mainly collected for large trees or smaller trees of large-tree species, especially commercially interesting ones. Other inventories include large trees as well as other life forms, like lianas, shrubs, understorey tree species, herbs and grasses (de Rouw 1991, Hall & Swaine 1976, 1981, Kouamé 1998b). For the purpose of this book we have chosen to use the large-scale data on large trees collected mainly by the national forest services. The main reason is the large coverage: many forest areas in all countries concerned have been inventoried in this way. A drawback is that in most inventories only a restricted number of large tree species are included. However, for a general classification of forests

Figure 4.1 Overview of Taï National Park, from Mt Nienokoué, Côte d’Ivoire.

this seems to be fine: large trees are better known than small-stature species. Another drawback is that most if not all of the selected species are timber species (that was the main reason for the inventories, after all), and thus the abundance of larger individuals of these species may be reduced due to past logging practices. We examine the major environmental factors that could determine the large-scale gradient. The best known factor is the rainfall gradient from c. 4 m per year along the coast to less than 1 m more inland. This rainfall gradient is generally linked to a gradual change from wet evergreen forest along the coast to semi-deciduous forest further inland, to savanna vegetation when the rainfall becomes too low. Small pockets of dry evergreen forest also occur at the lower end of the forest rainfall gradient. Other important environmental factors are soil fertility (Hall & Swaine 1981, Swaine 1996, White 1983) and altitude (Voorhoeve 1965, Hall & Swaine 1981). Most of the area is in the lowlands but in some places (e.g. Mount Nimba) altitude rises above 1000 m with an associated change in climate leading to a different vegetation. In this chapter we describe the main vegetation gradient in the forests of our area, and determine which are the major environmental factors that give rise to these gradients. We focus on large tree species and highlight the larger scale vegetation patterns.

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Methods Study area, plot- and data selection The area covered in this analysis is confined to the forest zone of Sierra Leone, Liberia, Côte d’Ivoire and Ghana. These countries contain most of the forest in Upper Guinea and are relatively well studied. The data we have used are collected by various organisations. For Sierra Leone we used the data from Small (1953), Savill & Fox (1967) and Davies (1987). For Liberia we used data from the German Forestry Mission to

Liberia (GFML 1967a, 1967b, Sachtler & Hamer 1967a, 1967b, Sachtler 1968) and the Liberian Forest Service. For Côte d’Ivoire we drew largely on data collected by SODEFOR (Clément & Guinaudeau 1973, SODEFOR 1978, 1979), and for some sites on species lists (Kouamé et al. Chapter 5). For Ghana we used the national forest inventory data (Hawthorne & Abu Juam 1995, Hawthorne 1995a, 1996, Wong 1989). In Liberia and Côte d’Ivoire only a number of large timber tree species have been surveyed (in each country c. 60 species), whereas in Ghana all species with a diameter at breast height (dbh) over 5 cm were included. To be able to compare the different areas we made a list of species that were inventoried in all three countries.

Table 4.1 Species used in the ordination/classification analysis. In some cases several species in a genus are pooled. Ordination scores of the first two axes are given for an ordination based on abundance data, and one on presence/absence data. Maximum size

Abundance

Height (m)

Axis 1

Axis 2

Axis 1

Axis 2

Presence/absence Species

Afzelia bella & africana Alstonia boonei Amphimas pterocarpoides Anopyxis klaineana Anthonotha fragrans Antiaris toxicaria Berlinia confusa & occidentalis Canarium schweinfurthii Ceiba pentandra Celtis adolfo-friderici & mildbraedii & sp. Daniellia ogea & thurifera Distemonanthus benthamianus Entandrophragma angolense & candollei & cylindricum & utile Erythrophleum ivorense & suaveolens/guianense Funtumia africana Gilbertiodendron preussii Guarea cedrata Guibourtia ehie Heritiera utilis Khaya anthotheca & ivorensis & grandifoliola Klainedoxa gabonensis Lophira alata Lovoa trichilioides Mammea africana Milicia excelsa & regia Nauclea diderrichii Nesogordonia papaverifera Parinari excelsa & sp. Petersianthus macrocarpus Piptadeniastrum africanum Pycnanthus angolensis Rhodognaphalon brevicuspe Ricinodendron heudelotii Terminalia ivorensis Terminalia superba Tetraberlinia tubmaniana Tieghemella heckelii Triplochiton scleroxylon Turraeanthus africanus Zanthoxylum gilletii

42

Species code

Family

Afz.spp Als.boo Amp.pte Ano.kla Ant.fra Ant.tox Ber.spp Can.sch Cei.pen

Leguminosae Apocynaceae Leguminosae Rhizophoraceae Ceasalpiniaceae Moraceae Leguminosae Burseraceae Bombacaceae

35 45 50 45 38 51 40 50 60

>90 140 120 120 120 130 >100 150 200

38.9 21.6 41.9 56.1 69.2 20.2 47.3 51.8 23.4

36.3 57.2 63.5 46.1 33.4 56.2 67.1 54.0 45.4

42.7 34.2 41.0 56.1 68.5 20.9 47.4 52.1 31.1

37.9 50.1 55.9 58.1 16.2 49.9 59.1 56.9 47.8

Cel.spp

Ulmaceae

54

100

6.9

54.3

26.8

42.1

Dan.spp Dis.ben

Leguminosae Leguminosae

45 36

120 97

49.5 32.0

60.8 50.6

50.0 33.3

51.2 40.8

Ent.spp

Meliaceae

60

250

24.5

57.4

20.8

48.5

Ery.spp

Leguminosae

40

120

55.1

28.1

53.4

31.1

Fun.afr Gil.pre Gua.ced Gui.ehi Her.uti

Apocynaceae Leguminosae Meliaceae Leguminosae Sterculiaceae

30 35 40 45 45

52 120 100 >100 300

42.3 77.1 23.6 15.1 66.7

55.7 48.3 66.9 0.0 51.6

34.7 74.0 26.4 21.9 63.5

52.6 0.0 59.4 1.6 46.8

Dbh (cm)

Kha.spp

Meliaceae

>50

>180

20.4

63.6

19.8

55.0

Kla.gab Lop.ala Lov.tri Mam.afr Mil.exr Nau.did Nes.pap Par.spp Pet.mac Pip.afr Pyc.ang Rho.bre Ric.heu Ter.ivo Ter.sup Tet.tub Tie.hec Tri.scl Tur.afr Zan.gil

Irvingiaceae Ochnaceae Meliaceae Guttiferae Moraceae Rubiaceae Sterculiaceae Chrysobalanaceae Lecythidaceae Leguminosae Myristicaceae Bombacaceae Euphorbiaceae Combretaceae Combretaceae Leguminosae Sapotaceae Sterculiaceae Meliaceae Rutaceae

40 50 45 40 >50 50 45 45 45 50 35 45 30 45 45 42 55 50 35 35

120 150 100 100 >150 150 120 150 180 180 120 120 112 124 150 100 250 >136 100 80

48.0 68.0 51.2 49.1 33.4 50.2 1.6 55.3 33.2 40.9 39.4 30.5 12.6 40.7 26.5 100.0 41.4 0.0 22.2 38.1

59.7 41.7 54.4 81.4 42.9 47.5 46.1 59.5 67.3 60.1 55.3 62.0 55.6 37.8 42.8 61.8 68.8 44.0 100.0 59.5

45.6 63.5 53.1 49.6 37.7 47.6 0.6 49.9 33.3 41.3 40.2 11.3 10.2 43.9 27.2 100.0 46.7 0.0 33.7 14.3

55.6 46.3 47.9 100.0 43.6 51.2 40.5 61.1 58.9 52.2 48.8 67.4 49.1 44.0 47.3 45.3 64.9 42.3 80.9 92.8

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For some genera the species are pooled because in some countries the species were identified to genus only (this is the case for Afzelia, Berlinia, Celtis, Entandrophragma, Erythrophleum, Khaya, Milicia, Parinari). In addition some species were added that occur in one country and are absent in the others (for instance Tetraberlinia tubmaniana is found only in Liberia). In total we have selected 40 species or species groups. In Table 4.1 the selected species are listed along with several characteristics. Most of the species have maximum sizes of over 40 m height and over 100 cm in diameter. Almost all selected species have commercial value, mostly as timber species. In total we collected species abundance data for 176 forest sites, 8 for Sierra Leone, 26 for Liberia, 37 for Côte d’Ivoire and 105 for Ghana. The inventoried area of each site is variable and ranges from 10 to 4500 ha (Appendix 2). Inventories vary from complete inventories to strip inventories. The 176 sites cover a wide range in environmental conditions (Table 4.2) (see below for sources of data). Rainfall varies between 1200 and 3500 mm per year, water holding capacity between 10 and 115 mm water/m soil, cation availability (Ca2+, Mg2+, K+) between 0.3 and 40 cmol per kg soil. The altitudes of the sites vary between 50 and 760 m above sea level. Latitude varies between 4.8 and 8.9 degrees and longitude between –11.4 and –0.5 degrees. Additionally, 38 sites were available with small areas (<10 ha), incomplete species abundance data, or only a species presence-absence list (Appendix 2). These sites were excluded from the analyses but included in the final forest classification and forest map. Classification and ordination of forests The main vegetation data consist of a species-plot matrix with 40 species and 176 plots with abundance values for species. For the abundance data the estimated number of trees >30 cm dbh per km2 was used. In Ghana all trees >30 cm dbh were inventoried and data are available on individual trees. All Liberian and Sierra Leonean sites had data for trees over 40 cm dbh and for Côte d’Ivoire various lower limits were available (>10 cm, >15 cm, >20 cm, >40 cm and >60 cm). To be able to compare all sites we estimated for each of the sites the number of trees >30 cm dbh per km2 as follows. Transformation values were based on the Ghanaian

Table 4.2 Values for selected environmental parameters for 176 forest sites in Upper Guinea, rainfall (in mm per year), CMK (cation availability in cmol per kg soil), WHC (water holding capacity in mm water per m soil) and altitude (m above sea level). N Rain CMK WHC Altitude

176 176 165 176

Mean 1780 3.8 68.7 218.5

Std. Dev.

Minimum

450.6 9.3 36.3 111.5

1194 0.3 10 51

Maximum 3422 39.3 112 758

data. For the Ghanaian data, for each species (thus the pooled data for all sites together) we constructed frequency distribution diagrams for size classes. Such a frequency distribution shows an idealised population structure. Most species then show a negative exponential pattern with size. For each species a negative exponential curve was fitted on the frequencies by size class. We used the regression to estimate numbers of trees >30 cm for each of the sites in Sierra Leone, Liberia and Côte d’Ivoire. We thus assumed that in the sites studied the frequency distributions would be similar to the idealised population structure. For the large samples sizes this is a reasonable supposition. We classified these 176 sites using a hierarchical classification algorithm, using Ward’s method for cluster optimisation and squared Euclidean distances. The classification resulted in seven clusters of sites. The log-transformed abundance matrix was also used for a detrended correspondence analysis, in which we reduced the multidimensional vegetation space into a two dimensional one, using Canoco (Ter Braak & Smilauer 1998). The first axis represents the main variation in species composition, the second axis the main variation once the first axis variation is removed. The analysis leads to scores for each site and for each species on the first two axes. The same procedure was followed for the same sites using presence/absence data. This allowed us to establish a relationship between the axis scores based on abundance and based on presence/absence. The results of the hierarchical site classification were overlaid on the ordination diagram to help determine a final classification of all sites into seven groups (forest types). For the 38 additional sites a presence/absence axis score was calculated based on the species presence/absence axis scores. Using the relationship between presence/ absence axis scores and abundance axis scores we calculated abundance axis scores for these 38 sites. Two forest maps of the Upper Guinea forest zone were constructed, one based on the first ordination axis, and one based on the hierarchical classification of the 176 sites into forest types. Interpolation of scores or classes between the sites was done using ArcView GIS (ESRI Inc.). Vegetation-environment relationships The resulting axes scores from the correspondence analysis (abundance values used) were related to environmental factors that are likely to influence large scale vegetation patterns such as total yearly rainfall (mm/yr), soil fertility (CMK, in cmol cations per kg soil), soil water holding capacity (WHC, in mm water/m soil), altitude (m) and geographical position (latitude and longitude). A rainfall map was created based on data from 578 weather stations in the region. Data on soil fertility and water holding capacity were calculated based on the FAO soil map of Africa and a quantitative review of chemical analyses of soil profiles (Batjes 1997). The water

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Figure 4.2 Scatterplots of the first two axes of a species-site ordination. The ordination is based on logtransformed abundance data (number of individuals >30 cm dbh per km2).

A

(A) Species. The abbreviations refer to the species in Table 4.1. (B) Sites. The symbols refer to the country.

B

C

holding capacity was calculated based on soil depth and soil texture. It was presumed that sandy soils have a water holding capacity of 75 mm/m, loamy soils of 100 mm/m, and clayey soils of 125 mm/m. For a detailed account on methods and analyses of the environmental metadata see Chapter 9. All spatial analyses were carried out using ArcView GIS (ESRI Inc.). To estimate environmental variables for a forest site, we used its centre. The ordination axis scores were related to each of the environment parameters using a Pearson’s correlation. To evaluate which of the environmental factors was most important in determining variation in species composition, the ordination axes 1 and 2 are regressed on the environmental parameters (and their squared valued to account for possible non-linear effects) using a stepwise multiple regression. Similarly, the axes were regressed on the geographical parameters latitude and longitude.

44

(C) Sites. The symbols refer to the hierarchical classification of the sites into seven clusters (see also Figure 4.4). The lines in the figure indicate the final classification of the sites (combined results of classification and ordination). HW = Hyper Wet ; WE = Wet Evergreen (type 1, 2, 3); ME = Moist Evergreen; MS = Moist Semi-deciduous; DS = Dry Semi-deciduous; SL = Sierra Leone type (in classification, part of HW).

Species responses to environmental factors The major vegetation gradient should reflect the distribution gradients of the major species involved. We show some examples of species distributions over the area and also their distribution with respect to the most important environmental gradient underlying the vegetation gradient, i.e. rainfall. The species distributions are made in ArcView, based on log transformed abundance data (interpolations and the values for each site). The non-linear species responses were modelled by regressing log-transformed abundance against rainfall and its quadratic term. Also, for each of the species studied we have analysed the major environmental factors determining its distribution and abundance. We used a stepwise multiple regression with the four environmental factors (rainfall, cation availability, water holding capacity and altitude) and their quadrates as the independents and species abundance as the dependent variable.

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A

Figure 4.3 (A) Spatial interpolation of the site ordination axis 1 scores, indicating the most important gradient in the species composition. For the analysis a selection of 40 species (in some cases species groups) is used. The size of the symbols indicates the axis scores. A large symbol (and dark interpolation colour) indicates a high axis 1 score (wet forests), a small symbol (and light interpolation colour) a dry forest.

(B) Spatial interpolation of rainfall in Upper Guinea, based on 580 weather stations in the area.

B

(C) Spatial interpolation of the forest types (based on final classification from Figure 4.2C, the classes between the lines). The symbols indicate the hierarchical classification results (the seven clusters). Sites with crosses are based on transformation from presence/absence ordination scores to abundance axis ordination scores.

C

Results The vegetation gradient The correspondence analysis for sites shows that the forests belonging to the same country are clustered together. Ghana and Liberia have the most distinct forests, and Côte d’Ivoire and Sierra Leone are in-between (Figure 4.2B). The most extreme plots on the first axis are Krahn Bassa South in Liberia and Bandai Hills in Ghana and on

the second axis Bia Shelterbelt and Onuem Nyamib in Ghana. These first two axes explain respectively 37.2% and 7.6% of the total variation in species composition. Axis 1 shows the major gradient in the composition of the forests studied. This means that the sites with the highest and the lowest axis 1 values are most different. This compositional information is used to construct a vegetation gradient map of Upper Guinea, in which for each site the size of the symbol is scaled to the axis score and the area inbetween is interpolated (Figure 4.3A). The general trend is clear and runs from high axis scores near the southwest coast to low axis scores in the northeast and east and is

45

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parallel to the general rainfall gradient in the area (Figure 4.3B). The forests in Liberia and Sierra Leone have the highest axis 1 scores and the forests along the savanna border in Côte d’Ivoire and Ghana have the lowest. The correspondence analysis also shows a clear separation between some of the wetter-forest species (Tetraberlinia tubmaniana, Gilbertiodendron preussii) and some of the drier-forest species (Triplochiton scleroxylon, Nesogordonia papaverifera) on the first axis. On the second axis Guibourtia ehie and Turreanthus africanus stand out, each at one end of the gradient (Figure 4.2A). The ordination based on abundance shows a good correlation to an ordination based on presence/absence data only (first axis Pearson’s r= 0.95, p<0.001; second axis Pearson’s r = 0.81, p < 0.001) Vegetation types The hierarchical clustering of the 176 sites resulted in seven groups, which roughly can be denominated hyperwet evergreen, wet evergreen (with three subtypes), moist evergreen, moist semi-deciduous and dry semi-deciduous. Unfortunately, some very distinct forest types have been effectively excluded from our analysis, in particular Upland types (such as the Upland evergreen forest of Ghana described by Hall and Swaine) and dry evergreen or other extreme dry forest types (Southern Marginal and Southeast outliers). There are a number of reasons, but most

importantly, inventory data (e.g. for Atewa in Ghana) includes the lowland portion with the upland portion of the reserves, and we have not attempted to subdivide the plots according to altitude there. The very dry forests are of little commercial interest and have not been inventoried. Figure 4.4 shows how the groups are divided. In the first separation the wetter forests are split from the drier ones. The wet group is then split into the hyperwet and the wet evergreen groups. From the dry group the moist evergreen is separated and later the dry semi-deciduous and the moist semi-deciduous forests are split. These results of the hierarchical site clustering have been put into the ordination diagram and the final classification is slightly adjusted with respect to their position in the ordination diagram (Figure 4.2C). The final site classification thus is a combination of the results of the hierarchical clustering and of the ordination. Only 22% of the sites have been adjusted, indicating that the hierarchical clustering gave roughly the same results as the (non-hierarchical) ordination. In Figure 4.3C all the sites are shown with their respective final forest classification. The background is an interpolation for the whole area based on these forest types. It is shown that the hyperwet forests are only found in Liberia and the southeastern part of Sierra Leone. The wet evergreen forest type is found in part of Sierra Leone, the eastern part of Liberia, and the western part of Côte

Table 4.3 Pearson correlations between the first two axes of the detrended correspondence analysis (log transformed abundance values) and selected environmental and geographical variables, and between these variables. CMK= cation availability, WHC= water holding capacity (n= 176, water holding capacity n=165). Parameters

DCA axis 1

DCA axis 2

Log CMK

WHC

Altitude

Latitude

Longitude

Rainfall Log CMK WHC Altitude Latitude Longitude

0.84 *** - 0.35 *** - 0.64 *** - 0.17 * - 0.28 *** - 0.75 ***

- 0.02ns - 0.10ns 0.31 *** - 0.31 *** - 0.54 *** 0.47 ***

- 0.26 ***

- 0.64 *** 0.39 ***

- 0.02ns 0.24 ** - 0.15ns

- 0.02ns 0.34 *** - 0.15ns 0.58 ***

- 0.73 *** 0.21 ** 0.74 *** - 0.11ns - 0.18*

Correlations are significant at 0.05 level (*), 0.01 level (**) or at 0.001 level (***) or not significant at 0.05 level (ns) (2-tailed).

Table 4.4 Forward (stepwise) multiple regression of ordination axes as function of selected environmental and geographical parameters. For the environmental parameters both single and squared values are used, to account for non-linear effects. From left to right the significance of the parameters decreases, but all parameters have significant contribution to the whole model. R2 values are increasing values with inclusion of every new parameter. Beta is the standardised value. DCA axis 1 Only environmental vars. DCA axis 1 Only geographical vars.

DCA axis 2 Only environmental vars. DCA axis 2 Only geographical vars.

46

Rain 0.74 0.60 Longitude 0.57 - 0.83

Altitude 0.76 - 0.17 Latitude 0.74 - 0.43

WHC 0.79 - 2.3

Altitude 0.12 - 0.72 Latitude 0.39 - 0.47

WHC 0.19 0.41 Longitude 0.43 0.39

Altitude2 0.27 0.48

WHC2 0.82 2.1

Log CMK 0.84 - 0.14

R2 Beta R2 Beta

Rain 0.29 2.77

Rain2 0.39 - 2.59

R2 Beta R2 Beta

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d’Ivoire (WE3), in a band east of WE3, mainly in the south west of Côte d’Ivoire (WE2) and in the south western part of Ghana (WE1). The moist evergreen forests (ME) are found in a wide band roughly north of the wet evergreen forest, with moist semi-deciduous forest (MS) north of that, in Côte d’Ivoire as well as in Ghana. The dry semi-deciduous forests (DS) are only found along the forest savanna boundary, especially in Ghana. Vegetation gradients: underlying factors Both the first and the second axes of the correspondence analysis are significantly correlated to all environmental and geographical parameters considered, except for the second axis which is not correlated with rainfall and cation availability (Table 4.3). However, these parameters are also strongly related to each other (Table 4.3). To look for the most important parameters we performed a forward stepwise linear regression on the environmental parameters only and the geographical parameters only (Table 4.4). The environmental parameters together explain 84% of the variation in species composition. The geographical variables also explain 74% of the variation, indicating a strong correlation between the environmental and the geographical parameters (Table 4.3). Rainfall is the single most important parameter for the first axis (Figure 4.5). The second axis is significantly correlated with all variables except two (rainfall and cation availability), although the correlations are weaker (Table 4.3). Also in the multiple regressions (Table 4.4) the relations are weaker: environmental parameters account only for 39% and geographical parameters account for 43% of the variation. Latitude is the single most important parameter for the second axis. The forest types that result from our analyses have been characterised with respect to the environmental conditions they experience (Figure 4.6). There is a clear decline in rainfall from the hyperwet evergreen forest to the dry semi-deciduous forest, as expected. In contrast to this, soil fertility and WHC increase along this forest gradient. Species responses to the environment The major vegetation gradient (Figure 4.3) should reflect the distribution gradients of the major species involved. In Figure 4.7 we show some examples of species distributions over the area and also their distribution with respect to the major environmental gradient underlying the vegetation gradient, i.e. rainfall. Some of the species follow the general vegetation gradient, for example Tetraberlinia occurs mainly in high rainfall areas and Nesogordonia occurs mainly in low rainfall areas. Other species show patterns that contrast with the general gradient and cannot be modelled as a response to rainfall (e.g. Amphimas, Piptadeniastrum). The majority of the species studied, for example Petersianthus, show a bellshaped response curve to the rainfall gradient, however.

Figure 4.4 Dendrogram of the hierarchical site classification. HW = Hyper Wet; WE = Wet Evergreen (type 1, 2, 3); ME = Moist Evergreen; MS = Moist Semi-deciduous; DS = Dry Semi-deciduous.

For each of the species studied we have analysed the major environmental factors correlated with its distribution and abundance. For most species several factors are needed for a significant model. Of the four environmental parameters analysed (rainfall, soil water holding capacity, altitude, cation availability) 12 out of the 40 species have only one significant factor, 16 have two, 10 have three and 2 have all four factors. In 24 out of the 40 species studied, rainfall was the most important factor, in an additional six species soil water holding capacity was the most important one, giving a total of 30 out of 40 where water availability was the major independent. In only six species neither rainfall nor water holding capacity is significantly correlated: Afzelia increases with altitude, Amphimas decreases with cation availability, Berlinia decreases with altitude and with cation availability, Piptadeniastrum decreases with cation availability, Terminalia ivorensis increases with altitude and decreases with cation availability, and Tieghemella decreases with altitude.

Discussion What determines the main vegetation gradient in Upper Guinea? Our results show that rainfall is the single most important parameter determining the vegetation gradient for the Upper Guinean forests. However, several other factors are closely correlated with rainfall, notably soil characteristics like cation availability and soil water holding capacity. Also latitude and longitude are strongly correlated to rainfall and these geographical factors, more than soil parameters, account for the rest of the variation not accounted for by rainfall. Longitude and latitude are also the most important underlying parameters for the

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Figure 4.5 Plot of the ordination axis 1 score (indicating the largest axis of variability in the species composition) as function of total amount of yearly rainfall (the parameter explaining most of the variability). The final site classifications (from Figure 4.1C) are given.

second ordination axis. This indicates that simply the location of a forest can give an indication of its position on the gradient. This is possible when location is well correlated to the most important environmental factors. This may be the case, with a strong overriding factor such as rainfall, which is rather unidirectional. Another reason may be that the gradient is a purely bio-geographical one, with a general slow turnover of species over a large distance. Rainfall and soil characteristics strongly co-vary at regional scales (Hall & Swaine 1981, Van Rompaey 1993). Van Rompaey (1993) also showed that local situations (like the catena position) could compensate for larger scale conditions, leading to a wetter type forest in a drier area. Also for the whole Upper Guinea this strong relation between rainfall and soil characteristics holds. Swaine (1996) tried to separate the effects of rainfall and soil, using data for 155 sites in Ghana. He abstracted a composite soil parameter, based on 15 basic soil parameters. Of the 15 soil parameters 11 were significantly related to the major vegetation gradient, and so was the composite soil factor. He also showed that that composite soil factor was strongly related to rainfall. At a larger scale (over more countries) more variable soils could be expected and thus probably a less strong relation to rainfall. The rainfall gradient could also be longer however. In our case the relation between rainfall and soil holds as well, even along the much longer rainfall gradient. Possibly the major soil factors remain largely determined by rainfall. Another possibility is that the large tree species taken into account in this study are not susceptible to the extra part of the gradient (both soil and rainfall). Swaine (1996) argues that both nutrient limitation and moisture determine the distribution limits of large tree species. Of the soil parameters mineral cations seem to be the most important,

48

and phosphorus seems not to be important for determining floristic composition (cf. Burslem et al. 1995). This is rather surprising as low phosphorus availability has been shown to strongly determine vegetation structure and species composition in Guyana (Ter Steege 1993, Raaimakers 1994) and other areas (Vitousek & Sandford 1986). Swaine (op. cit.) found a low impact of phosphorus on species composition in Ghana. In this study we ignored the importance of the seasonality of rainfall. In an earlier study, for southeast Liberia and southwest Côte d’Ivoire, Bongers et al. (1999) showed that water availability, rather than rainfall alone was the most important parameter determining the abundance of a number of species. Length of the dry period and the intensity of that period (calculated as the cumulative water deficit) also had an effect, but less strong than the amount of rainfall. Here we have ignored these factors, but included others that are related to water availability, like soil water holding capacity. Surprisingly, the amount of rainfall alone accounts for 74% of the variation in species composition at the vegetation level. Large trees generally are not the most sensitive plants with respect to gradual changes in environmental conditions. Including other woody species like lianas and shrubs (Hall & Swaine 1976) or herbs (De Rouw 1991) would refine the analysis and enhance the potential for indicating changes in vegetation composition at relatively small scale. De Rouw, for instance, showed that at local scale forest composition changed clearly in southwest Côte d’Ivoire, where Guillaumet (1967), working with larger plants only, did not find changes. Herbs do react to small changes in water availability while large trees probably do not. This indicates that herbs could be used to indicate gradients at small spatial scales. In some areas the floristic gradient is steeper than in others, indicated by the intensity of change. This reflects larger changes in environment over smaller distances (cf. Van Rompaey 1993). For instance, along the line from Haut Sassandra in Côte d’Ivoire to the Krahn Bassa forests in Liberia the rainfall gradient is rather steep and thus the associated change in species composition. The same accounts for the southeast of Ghana. On the other hand, east-west gradients may be very shallow indicating that, for instance, the Haut Sassandra forests are similar to many of the forests in Ghana. Altitude has a significant (negative) correlation to both ordination axes. The altitudinal range included is low however, as the highest forest occurs at 750 m above sea level. In general high altitude sites have high cation availability and are found at higher latitudes. High altitude sites are Lamco, North Lorma, Worobong South and Southern Scarp. In Lamco and North Lorma Heritiera is the most important species, in Worobong South and in Southern Scarp this is Celtis. In Atewa (very close to Worobong South), the data for the high plateau are mixed with data from lowland sites. The high altitude sites do not seem to take a special position in the overall gradient,

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Chapter 4. The forests of Upper Guinea: gradients in large species composition

probably because our data for these sites have been “polluted” by data from the adjacent lowlands. Because we had only very few high altitude sites we have indicated the high altitude areas directly on the vegetation map.

A

The merits of a regional map For Upper Guinea several vegetation maps are available, but the scale is either too large or too small. Our map is one of the first for the whole Upper Guinean forests in one analysis (see also Van Rompaey 2001). We used forests from a large area (the two most distant forests are c. 1130 km apart, compare Hall & Swaine c. 530 km for Ghana). This gave rise to a longer environmental gradient. Our rainfall gradient is from 1200 to 3500 mm while the forest gradient in Ghana is from little under 1000 to 2000 mm. The differences in soil related environmental gradients between Upper Guinea and Ghana are of the same magnitude. However, the relations between the composition and environmental factors remained much the same. Because of the larger area covered it is easier to interpolate for forests where no data are available. For example, in central Liberia hardly any detailed inventories have been done, and neither have botanical collections been made (see Wieringa & Poorter, Chapter 6). As our map is based on a selection of only 40 species, it is possible for the non-specialists (e.g. a forest manager) to locate a forest on the gradient, based on general inventory data. The fact that presence/absence data give similar results, a result found before by Hall and Swaine (1976), shows that not even data on species abundance are needed. This has been applied successfully before (e.g. Hawthorne 1995a, Hall & Swaine 1981). Another advantage is that a common map makes it possible and easy to compare forests in different countries, which was not possible before. On the other hand, of course, we have to realise that with the large scale some of the local scale information is lost. Local environmental conditions can have a strong impact on the vegetation leading to deviations from the general pattern. A specific forest then can be of a drier or wetter type than expected based on interpolations from a larger area (see Hawthorne 1996 for examples in Ghana).

Figure 4.6 Comparison of forest types with respect to (A) rainfall, (B) water holding capacity and (C) cation availability.

Comparison with existing national and local vegetation maps We compared our forest types with other ones that were made for the separate countries Ghana, Côte d’Ivoire, Liberia and Sierra Leone (Table 4.5). Our vegetation map is comparable to the map for Ghana (Hall & Swaine 1981). Their gradient from wet evergreen via moist evergreen to moist semi-deciduous and further to dry semi-deciduous nicely fits into the longer gradient as developed here. The gradient here is longer however, and resulted in an extra wet forest type, that we called the hyperwet evergreen forest. Hall & Swaine (1976, 1981) distinguished four forest types that do not show in our analysis and that are important for biodiversity in Ghana:

upland evergreen, the northwest moist semi-deciduous type (MSNW), the southern marginal type and the southeast outlier type. The upland evergreen did not show up in our analysis as we have only a few sites that fall into one of the other classes (based on our 40 species). As this forest type is important for the whole of Upper Guinea we have put an extra class on the map, based on altitude. Their distinction within the moist semi-deciduous forests does not show in our classification because the most distinguishing species for their MSNW have been excluded (Pericopsis) or merged with relatives (Khaya

B

C

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A

B

T. tubmaniana

N. papaverifera

P. macrocarpus

Figure 4.7 Abundance and response curves to rainfall for the three characteristic species Tetraberlinia tubmaniana, Nesogordonia papaverifera and Petersianthus macrocarpus. (A) Spatial interpolation of the abundance values for three selected species for each of the sites. The size of the symbols indicates the abundance. A dark colour indicates a high abundance, a light colour a low abundance. (B) Response curves of species abundance on rainfall.

anthotheca). The remaining two types do not occur elsewhere in sufficiently large areas (in fact neither in Ghana, cf Hawthorne & Abu Juam 1995). Waterman et al. (1978) made a general vegetation map for Ghana and Côte d’Ivoire. The map for Ghana is based on Hall & Swaine (1976) and they extended that system into Côte d’Ivoire. Wet evergreen forest was found in the southeast and in the southwest of Côte d’Ivoire, which is comparable to our results. Guillaumet & Adjanohoun (1971) recognised four main forest areas, the moist closed evergreen forests, the moist closed semi-deciduous forests, the littoral forest and the moist closed montane forest. The evergreen forests (their “secteur ombrophile”) are found in areas with more than 1700 mm of rainfall and a dry period of between 2

50

and 5 months, while the semi-deciduous forests (their “secteur mesophile”) are found in areas with beween 1200 and 1600 mm rainfall and a 4-6 months dry period. Within the evergreen forests they distinguish five types, based on specific species. Ecologically they are differentiated on a combination of soil characteristics (mainly related to water retention capacity in the soil, rainfall and the length of the dry season). Within the semi-deciduous forest (generally between 1200 and 1600 mm of rain) they recognised three forest types, based on characteristic species, and not on soil or climate: a group with Aubrevillea kerstingii and Khaya grandifoliola, a group with Celtis spp. and Triplochiton scleroxylon (1200-1600 mm/yr), and a group with Nesogordonia papaverifera and Khaya ivorensis. This last

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Chapter 4. The forests of Upper Guinea: gradients in large species composition

forests, where the evergreen forests are determined by high annual rainfall and an absence of a dry period. This coincides with the wetter part of our gradient (the higher axis 1 scores). The forests of Sierra Leone were classified into two broad groups (Savill & Fox 1967), the moist semideciduous forests (consisting of two types) and the tropical rainforest. The difference is mainly based on the length of the dry season, without clear differences in total yearly rainfall. In a forest classification for Nigeria, Hall (1977) found that the major forest division was based on soil characteristics. Only the second level classification was based on rainfall, but that may have been due to the slightly biased sample selection: one of the soil/rainfall categories was effectively missed out (M.D. Swaine personal communication). In our analysis, in contrast, soil is not important compared to rainfall. Probably rainfall is really the overruling factor as long as a large rainfall gradient is covered. At more local scales, with shorter rainfall gradients, soil may become more important. We expected to find a large variation in nomenclature, even in a relatively small area as Upper Guinea. However, we did not expect the cut-off limits between the forest types to be that variable (Table 4.5). Of course this

group is intermediate between semi-deciduous and evergreen forest and mainly located in the east of Côte d’Ivoire, with between 1400 and 1500 mm rainfall per year. In Côte d’Ivoire, the ecotone between the evergreen and the semi-deciduous forest is south of Man in the west, via south of Gagnoa in the centre to south of Abengourou in the east, roughly following the distribution overlap of the species Uapaca guineensis, U. esculenta, Lophira alata, Sacoglottis gabonensis (wet end), Mansonia altissima (dry end) and Celtis species. The northern limit of forest is marked by the distribution limits of Aubrevillea kerstingii, which is north of Man, north of Daloa, north of Gagnoa to just south of Bondokou. Their montane forests are delimited at above 1000 m above sea level, mainly located in the west, near Mount Nimba, and in the massive of Dans. Parinari excelsa is a characteristic species. The littoral forest extends only to a few kilometres from the coast and is slightly more dry than further inland, mainly due to a longer dry season and a less water-holding soil (see also Kouamé et al., chapter 5). Our gradient coincides with the forest types in Liberia (Voorhoeve 1965). Voorhoeve distinguished two major types, the evergreen forests (with a mixed type and a mono-dominant type) and the moist semi-deciduous

Table 4.5 The present classification compared to the most recent ones for the separate countries. Rainfall (mm)

Sierra Leone 1

Liberia 2

Côte d’Ivoire 3

1000

1200

1400

1600

1800

2000

2200

2600

Moist semi-deciduous forest (Tonkoli type and Kasewe type) (>2500, 4 months of dry period) Tropical rain forest (>2500, <3 month dry period) Evergreen (>2000)................. - mixed forest type - mono-dominant type Moist semi-deciduous (ecotone) Semi-deciduous (1600-2000)................. Semi-deciduous (1200-1600) 3 types (dry period 6-8 months, deficit >600 mm) Evergreen (1700 till over 2000) 5 types ................. (dry period 2-5 months)

Ghana 4

Upper Guinea 5

Southeast Outliers (<1000) Southern Marginal (1000-1250) Dry semi-deciduous (1250-1500) Moist semi-deciduous NW type ( 1250-1500) Moist semi-deciduous SE type (1500-1750) Moist evergreen (1500-1750) Wet evergreen (>1750)................. Dry semi-deciduous (1200-1600) Moist semi-deciduous (1250-1750) Moist evergreen (1500-1800) Wet evergreen1 (1400-2300) ................. Wet evergreen 2 (1600-1700) Wet evergreen 3 (1700-2100)................. Hyper wet evergreen (2000-3500)................. .................Upper evergreen forest (above 500m altitude).................

1) Savill & Fox 1967, 2) Voorhoeve 1965, 3) Guillaumet & Adjanohoun 1971, 4) Hall & Swaine 1981, 5) This chapter.

51

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Chapter 4. The forests of Upper Guinea: gradients in large species composition

depends on the available environmental variation in the area under consideration. For instance, the wet forests of Ghana are the wettest forests found in the country but compared to other countries like Liberia and Sierra Leone the Ghanaian forests are relatively dry. Although the local situation sometimes is different, the present classification of forests thus provides a framework for forest classification in the whole forest zone of Upper Guinea. Conclusions We classified the forests of Upper Guinea into eight different types: seven types resulted from a combined classification and ordination of 176 forest sites, one extra class is for the montane evergreen forests. This Guineawide classification was similar in the resulting forest types to earlier smaller-scale forest classifications (mostly for one country or more local situations). The classification was based on a selection of 40 large tree species only. There is a good similarity between a classification

52

based on abundance and one based on presence/absence only. Based on this classification and a spatial interpolation we produced a vegetation map for whole Upper Guinea. Although this regional map is less detailed compared to earlier produced maps for more local situations, the new map provides a nice integration for Upper Guinea as a whole.

Acknowledgements We thank R.S.A.R. van Rompaey for collecting part of the data and for discussions, A. Siepel and T. Helmink for help with data handling and figures, and M.D. Swaine for comments on the manuscripts.

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Floristic diversity of closed forests in Côte d’Ivoire

5

F. N’. Kouamé, K.E. Kouadio, K. Kouassi and L. Poorter

H A P T E R

5

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Introduction

The Ivorian flora is estimated to number 3660 species (Davis et al. 1994) that are found in four main vegetation types (Figure 5.2). Ivorian closed forests, which correspond to the rainforests of White (1986) are divided in evergreen and semi-deciduous forests. In Côte d’Ivoire, the savanna penetrates the forest zone in a region called V-Baoulé (Guillaumet & Adjanohoun 1971, Gautier & Spichiger chapter 3). On each side of V-Baoulé, rainfall increases towards the southwest and southeast (Eldin 1971). Rainfall also increases from lower to higher altitudes. Along these rainfall gradients soil humidity increases, inducing a change in the vegetation from semi-deciduous to wet evergreen forests (Hall & Swaine 1981) and an increase in floristic diversity. The lowland evergreen forests in the southeast (Mabi, Songan, Tamin, Yaya) and southwest (Haute Dodo, Taï), and the mountain forests in the west (Mont Tonkoui) have the highest rainfall in the country (Eldin 1971). They have also been identified as the centres of greatest floristic diversity in Côte d’Ivoire (Aubréville 1949, Guillaumet 1967, Anonymous 1991, Beentje et al. 1994). Guillaumet and Adjanohoun (1971) have distinguished a floristic species group in southwest Côte d’Ivoire which they coined “Sassandrian”. This group includes species that occur between the Cavally river and the Sassandra river, from Tabou to Taï. Another species group, called “Ghanaian”, occurs in southwest Ghana (Hall & Swaine 1981), and appears to penetrate into the east of Côte d’Ivoire. In this chapter, we investigate the environmental factors responsible for the floristic composition in the lowland closed forests of Côte d’Ivoire. The forest blocks of southeast and southwest Côte d’Ivoire are currently separated by the relatively dry region of the V-Baoulé. They have probably been separated by even drier savanna vegetation during the dry glacial periods (Gautier & Spichiger chapter 3). Nevertheless, we expect to find high floristic similarities between these forest blocks, because they experience similar high rainfall levels, and because they have been connected by wetter forest types during the interglacial periods.

Figure 5.1 Ivorian botanist Henry Téré showing the beautiful cauliforous tree Omphalocarpum elatum (Sapotaceae) in Banco National Park, Côte d’Ivoire.

Study sites and methods

The study sites comprise thirteen forests in the centreeast, the southeast, the southern coast, the centre-west, and the southwest of Côte d’Ivoire. The choice of these different forests was based on the intactness of the forest cover and the availability of species lists. The forests vary from semi-deciduous to evergreen (Figure 5.2), they are of various sizes (2590 to 300,000 ha), and have different soil and climatic conditions (Table 5.1). Three soil types can be distinguished: tertiary sands, clay soils derived from schist, or sandy soils resulting from granite weathering (De Rham 1971). The total annual rainfall ranges between 1400 and 2300 mm, while the water deficit is between 150 and 400 mm. The length of the dry season varies between 2 and 5 months (Perraud 1971). Most of the data used in this research come from various studies, in which checklists are made for the forests (Table 5.1). Such inventories aim to identify all the plant taxa encountered while following pre-existing paths in the forest or paths created for this purpose. In addition, we carried out inventories in those forests that had no or

53

Area (ha)

Coordinates

Centre-East

Bossématié

Bm

SD

22,200

6° 20’ - 6° 35’N 3° 20’ - 3° 35’W

1400 - 1500

5° 51’ - 6° 05’N 3° 22’ - 3° 41’W

1650 - 1700

5° 46’ - 6° 12’N 3° 12’ - 3° 26’W

1600 - 1650

5° 40’ - 5° 58’N 3° 11’ - 3° 25’W

1650 - 1700

5° 35’ - 5° 54’N 3° 23’ - 3° 46’W

1700 - 1800

5° 00’ - 5° 07’N 5° 50’ - 5° 57’W

1550 - 1600

4° 44’ - 4° 58’N 6° 14’ - 6° 35’W

1650 - 1750

5° 06’ - 5° 11’N 5° 29’ - 5° 34’W

1550 - 1600

5° 21’ - 5° 25’N 4° 01’ - 4° 05’W

2000

6° 22’ - 7° 24’N 6° 59’ - 7° 10’W

1460 - 1680

6° 53’ - 7° 14’N 5° 46’ - 6° 10’W

1400

4º 41’ - 5º 19’N 7º 01’ - 7º 25’W

1900 - 2300

5° 09’ - 6° 09’N 6° 48’ - 7° 26’W

1800 - 2200

Southeast

Mabi

Songan

Tamin

Yaya

South coast

Dassioko

Monogaga

Port Gauthier

Banco

Centre-West

Southwest

S

T

Y

D

Mo

PG

B

Haut Sassandra

HS

Marahoué

Ma

Haute Dodo

Taï

1. 2. 3. 4.

Mb

Guillaumet & Adjanohoun (1969) Eldin (1971) Perraud & Souchère (1970), De Rham (1971) Eldin & Daudet (1968)

HD

Taï

E

E

E

E

E

E

E

E

SD

SD

E

E

59,616

38,189

24,934

23,877

11,203

39,660

2,590

3,300

102,400

101,000

236,733

300,000

Annual rainfall2 (mm/yr)

Subsoil and soil3

Water deficit4 (mm/yr)

Taxa (#)

Source

Schist

350 - 400

611

Aké Assi (1992) Bakayoko (1999)

Schist

300

640

This study

Schist

300

591

This study

Schist

200

512

This study

Schist

200

617

This study

Sand

450

719

Aké Assi (1997)

Sand

300

859

Kouamé (1998b)

Sand

450

705

Kouamé (1998b)

Sand

200

773

De Koning (1983)

Granite

400

843

Kouamé (1998a)

Schist

350 - 400

475

Jongkind et al. (1999)

Granite

150

906

Kouadio (2000) Kouassi (2000)

Granite

200 - 250

849

Aké Assi & Pfeffer (1975)

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Forest type1

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Abbr.

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Chapter 5. Floristic diversity of closed forests in Côte d’Ivoire

Geographical Zone

22466 layout Chapter 5 p53-60

54

Table 5.1 The forests included in this study (E = evergreen, SD = semi-deciduous) and their environmental characteristics. "Abbr." refers to the abbreviations used in the figures. “Taxa” indicates the number of taxa found in the forests.

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Chapter 5. Floristic diversity of closed forests in Côte d’Ivoire

Figure 5.2 Map of Côte d’Ivoire with four main vegetation types (evergreen forest, semi-deciduous forest, Guinean savanna, sub-Sudanian savanna), the forest reserves (white polygons), and the 13 forests studied (dark grey polygons). The bold V-shaped line indicates the savanna intrusion in the forest zone, or V-Baoulé.

incomplete species lists. Thus, in the southeast forest area (Mabi, Tamin, Songan and Yaya), sixteen 2-km tracks, previously cleared by the forest management services (SODEFOR-GTZ), were used for species inventories. In Haute Dodo, due to the lack of specially cleared tracks, we randomly distributed several sites for inventories, following the pre-existing paths (Kouadio 2000). These inventories were accompanied by an inventory of all species in sample plots (Kouassi 2000). We are aware that the species lists analysed in this research are not exhaustive. In consequence, our discussions are based on the current state of knowledge of the flora of Ivorian forests. The species lists of the forests were pairwise compared using Sørenson’s similarity index (1948). This allowed us to distinguish forests that were more similar in species composition and those that were more dissimilar. The maximum value of the index (100%) indicates that two forests have the same species composition. The minimum value (0%) indicates that two forests have no species in common. Subsequently an ordination of the 13 forests and 2126 taxa was carried out using correspondence analysis (Jongman et al. 1987), based on the presence or absence of

species in the species lists. The ordination score was related to the following environmental parameters: longitude and latitude, soils and subsoil, the annual water deficit and the total annual rainfall. The groups of forests produced by the correspondence analysis were pairwise compared using Sørenson’s similarity index. The taxa which were found in all the forests were considered as being the most common species of Ivorian closed forests. The taxa which were inventoried in all the forests of the same group, but which were not found in any other forest group, were considered as being typical for the group in which they were found.

Results

The floristic diversity of the thirteen forests studied comprises 150 families, 857 genera and 2126 species. 624 species are found in only one of the forests, while 41 species are found in all the forests (Table 5.2). The number of species shared by the forests is inversely

55

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Chapter 5. Floristic diversity of closed forests in Côte d’Ivoire

Table 5.2 Taxa that are common to the 13 forests, or taxa that are characteristic for each of the four forest groups (Fig. 5.4). Species common to the 13 forests

Coastal forests (Group II)

Aganope leucobotrya Agelaea paradoxa Agelaea pentagyna Aidia genipiflora Alstonia boonei Amphimas pterocarpoides Baphia nitida Baphia pubescens Buchholzia coriacea Bussea occidentalis Calycobolus africanus Cnestis ferruginea Cola nitida Costus afer Craterispermum caudatum Culcasia barombensis Diospyros soubreana Funtumia africana Glyphaea brevis Griffonia simplicifolia Klainedoxa gabonensis Landolphia hirsuta Landolphia owariensis Microdesmis keayana Myrianthus arboreus Myrianthus libericus Napoleonaea vogelii Nephrolepis biserrata Neuropeltis acuminata Ochthocosmus africanus Palisota hirsuta Parinari excelsa Piptadeniastrum africanum Psychotria peduncularis Pycnanthus angolensis Strophanthus gratus Strychnos aculeata Treculia africana Ventilago africana Xylopia quintasii Xylopia villosa

Ancistrocladus barteri Bulbophyllum imbricatum Eugenia whytei Eupatorium microstemon Heterotis rotundifolia Salacia pallescens Salacia whytei Tapinanthus belvisii

Southwest forests (Group III) Angraecum podochiloides Anthoclitandra nitida Bertiera fimbriata Bolbitis heudelotii Brieya fasciculata Cercestis ivorensis Clappertonia minor Dalbergia albiflora Delpydora gracilis Didelotia brevipaniculata Drypetes klainei Garcinia granulata Gilbertiodendron robynsianum Gynura sarmentosa Lomariopsis rossii Mapania minor Millettia lucens Mussaenda landolphioides Pauridiantha hirtella Polystemonanthus dinklagei Premna grandifolia Psychotria subglabra Renealmia maculata Scleria vogelii Selaginella versicolor Strychnos icaja Tarenna gracilis Trichilia heudelotii Vitex ferruginea

Semi-deciduous forests (Group I) Acroceras gabunense Aneilema umbrosum Bridelia atroviridis Clerodendrum polycephalum Cyrtococcum chaetophoron Desmodium adscendens var.robustum Dichapetalum madagascariense var. madagascariense Diospyros abyssinica Eugenia tabouensis Grewia carpinifolia Khaya grandifoliola Lagenaria breviflora Landolphia landolphioides Melochia melissifolia Mischogyne elliotianum var. glabra Psychotria kitsonii Psydrax manensis Simirestis dewildemaniana Strychnos congolana Strychnos splendens Telfairia occidentalis Vitex ferruginea subsp. ferruginea

56

Southeast forests (Group IV) Aframomum alboviolaceum Buforrestia mannii Cecropia peltata Costus englerianus Crotonogyne craterviflora Friesodielsia enghiana Guibourtia copallifera Guibourtia tessmannii Licania elaeosperma Marantochloa filipes Memecylon polyanthemos Rutidea dupuisii subsp. occidentalis Sabicea discolor

correlated to the number of forests compared (Figure 5.3). Nine families are each represented by more than 50 species (Table 5.3). Rubiaceae and Euphorbiaceae are the most speciose families. The richest genera are Psychotria and Ficus with 43 and 34 species respectively; they top the list of the six genera represented by at least 20 species each (Table 5.3). The similarity index is lowest (29%) between Banco and Marahoué (Table 5.4), which reveals the floristic dissimilarity of these two national parks. The similarity index is highest (77%) between Songan Forest Reserve and Tamin Forest Reserve, which resemble each other most. The highest values are found either between forests belonging to the same geographical zone (Taï/Haute Dodo, Songan/Tamin, Mabi/Songan, Tamin/Yaya) or between a forest of the southeast and a forest of the southwest (Haute Dodo/Mabi, Haute Dodo/Yaya). The similarity index varies from 38% (Dassioko/Marahoué) to 53% (Bossématié/Songan) for the forests with the same longitudes but different latitudes. The first and second axis of the correspondence analysis explain together 31% of the variation in species composition. 44% of the variation in species composition is explained by the six environmental variables. Four groups of forests can be distinguished (Figure 5.4). Group I, made up of semi-deciduous forests (Bossématié, Haut Sassandra, Marahoué), is characterised by a high water deficit (Figure 5.5A) and a high latitude (Figure 5.5B). The coastal forests (Banco, Dassioko, Port Gauthier and Monogaga) make up group II, which is characterised by a low latitude, intermediate rainfall, and occurrence on tertiary sandy soils (Figure 5.4). Group III corresponds to the forests of the southwest (Haute Dodo, Taï), while group IV corresponds to the forests of the southeast (Mabi, Songan, Tamin, Yaya). Groups III and IV are

Table 5.3 The most common families and genera from all 13 forests. Families with more than 50 species each are shown, and the genera represented by at least 20 taxa. Psychotria is principally herbaceous and Ficus includes mainly shrubs. The genera Salacia, Combretum, Strychnos and Dichapetalum are essentially lianescent. Family

Species (#)

Genus

Species (#)

Rubiaceae

260

Psychotria

43

Euphorbiaceae

107

Ficus

34

Fabaceae

86

Salacia

28

Apocynaceae

77

Combretum

27

Orchidaceae

76

Strychnos

21

Annonaceae

70

Dichapetalum

20

Caesalpiniaceae

69

Moraceae

52

Hippocrateaceae

51

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Chapter 5. Floristic diversity of closed forests in Côte d’Ivoire

centres of greatest floristic diversity in the lowlands in Côte d’Ivoire. They are characterised by an intermediate latitudinal position, quite high rainfall and a low water deficit (Figure 5.4). They differ fundamentally in the type of their subsoils. The subsoil of the southwest is essentially of granite, whereas that of the southeast is composed of schists. The annual water deficit and the latitudinal position are the environmental factors that are best correlated with the first axis (Figure 5.5, Table 5.5). The second axis has a strong positive correlation with tertiary sandy soils, a strong negative correlation with schists (Table 5.5), and a weak negative correlation with latitude (Figure 5.4B). The similarity coefficients between the groups of forests produced by the correspondence analysis are always higher than 50%. This indicates that these groups of forests have many species in common. The southwest and southeast groups have the highest index of similarity (65%). Seventy-two species are characteristic for one of the forest groups: 22 species are typical for the semi-deciduous forests, 8 for the coastal forests, 29 for the southwest forests and 13 for the southeast (Table 5.2).

Table 5.5 Pearson’s correlation coefficients between the environmental factors and the first two ordination axes of the correspondence analysis. ns= not significant; * = P < 0.05; ** = P < 0.01. Variable

Axis 1

Latitude

Axis 2

0.70

**

– 0.61

*

Longitude

– 0.38

ns

– 0.38

ns

Rainfall

– 0.39

ns

0.45

ns

Water deficit

0.70

**

0.15

ns

Granite

0.14

ns

– 0.04

ns

Sand

– 0.11

ns

0.90

**

Schist

– 0.02

ns

– 0.80

**

Table 5.4 Matrix of similarities between forests using Sørensen’s index (1948). The figures in brackets correspond to the cumulative species richness of the two forests that are compared. The similarity indices larger than 50% (indicating that two forests have more than 50% of the species in common) are underlined and those larger than 60% are given in bold. Forest

Banco

Bossématié

Dassioko

Haut Haute Sassandra Dodo

Port Gauthier

Mabi

Marahoue

Monogaga

Songan

Taï

Bossématié

42 (1382)

Dassioko

48 (1495)

51 (1328)

Haut Sassandra

42 (1618)

58 (1447)

48 (1561)

Haute Dodo

48 (1685)

46 (1518)

53 (1632)

50 (1751)

Port Gauthier

48 (1482)

53 (1313)

69 (1427)

49.5 (1546)

53 (1617)

Mabi

46 (1417)

48 (1249)

53 (1363)

45 (1482)

61 (1553)

52 (1348)

Marahoue

29 (1250)

50 (1082)

38 (1195)

57 (1318)

37 (1385)

40 (1182)

36 (1117)

Monogaga

49 (1635)

50 (1467)

73 (1581)

49 (1700)

57 (1771)

69 (1566)

54 (1502)

37 (1335)

Songan

45 (1367)

53 (1200)

53 (1314)

48 (1433)

59 (1504)

52 (1299)

72 (1235)

40 (1067)

54 (1453)

Taï

40 (1625)

44 (1457)

53 (1571)

44 (1690)

58 (1761)

49 (1556)

50 (1492)

33 (1325)

55 (1710)

48 (1443)

Tamin

43 (1289)

46 (1121)

49 (1235)

43 (1354)

56 (1425)

49 (1220)

69 (1156)

37 (989)

49 (1374)

77 (1107)

45 (1364)

Yaya

46 (1392)

46 (1224)

54 (1338)

44 (1457)

60 (1528)

52 (1323)

72 (1259)

34 (1092)

53 (1477)

72 (1210)

50 (1467)

Tamin

68 (1131)

57

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Chapter 5. Floristic diversity of closed forests in Côte d’Ivoire

Figure 5.3 Relationship between the number of species in common and the number of forests compared. Spearman’s rank correlation coefficient is given.

Discussion

Floristic diversity The 9700 km2 of forests included in this research, represent less than 50% of the forest cover of the country before 1985 (Davis et al. 1994) and less than 3.6% of the national territory. However, 2126 species of vascular plants were found in these forests, which represents 58% (out of 3660 species) of the total Ivorian flora (Heywood & Davis 1994). This indicates that the 13 forests contribute to a large extent to the flora of Côte d’Ivoire. The high contribution can be explained by the fact that all principal lowland forest types in Côte d’Ivoire were included in the study. The remaining 42% of the Ivorian flora is found in savannas, the upland evergreen forests, forest islands, gallery forests, granitic domes and human environments. The species richness of these forests resembles that of certain countries such as Benin, Liberia, Senegal or Togo. It is richer than Sierra Leone, whose flora is evaluated at 1700 species (Heywood & Davis 1994). It represents almost two-thirds the diversity of Guinea and Ghana. The relative paucity of flora in Benin, Togo and Senegal can be attributed to their low rainfall and to their limited forest cover. Liberia is considered to harbour the former glacial forest refuges (Morley 2000, Wieringa & Poorter chapter 6). The relatively low richness of Liberia and Sierra Leone may be attributed to the lack of knowledge of the flora of these countries. Haute Dodo is the richest forest in Côte d’Ivoire with at least 906 species. The Rubiaceae (260 species) and Euphorbiaceae (100 species) were the most species rich families. At the national level, these forests contribute to respectively 85 and

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Figure 5.4 Ordination diagram of 13 Ivorian forests, showing their position on the first two axes of the correspondence analysis. Forest sites are indicated with filled symbols, and environmental factors with arrows. The forests are arranged in four groups. Abbreviations are given in Table 5.1.

75% of the species richness of these families (Aké Assi 1984). The Fabaceae, represented in the Guinean savannas by 103 species (Kouamé 1993, Bänninger 1995), are not uncommon in closed forests, where they account for 86 species. The families of Dioncophyllaceae, Hoplestigmataceae, Medusandraceae, Octoknemataceae, Pandaceae and Scytopetalaceae are present in the 13 forests, and represent nearly 70% of the endemic families in the GuineoCongolian region (White 1986). Numerous endemic Guineo-Congolian genera such as Afrobrunnichia, Amphimas, Anopyxis, Anthonotha, Antrocaryon, Aubrevillea, Buchholzia, Calpocalyx, Chidlowia, Coelocaryon, Coula, Crotonogyne, Cyclodiscus, Decorsella, Didelotia, Discoglypremna, Distemonanthus, Duboscia, Heckeldora, Hymenostegia, Gilbertiodendron, Grossera, Monocyclanthus, Ophiobotrys, Tieghemella and Turraeanthus have also been found in these forests. Relationships among forest blocks The distinction in forest types is governed by water availability, which is the combination of the total amount of rainfall, the rainfall distribution over the year, and the water holding capacity of the soil. The largest distinction in species composition in our ordination analysis is between semi-deciduous forests on the one hand, and the evergreen and coastal forests on the other hand (Figure 5.4). The semi-deciduous forests clearly have a lower rainfall which is unevenly distributed over the year. Although the coastal forests have a similar high rainfall as the other two evergreen forest groups, the water availability to the vegetation is substantially lower, because of a high water deficit, and sandy soils with a low water holding capacity. The more subtle separation between the two evergreen forest groups is mainly caused by a higher rainfall amount in the southwest.

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A

B

Figure 5.5 Relationship between A) the first axis score of the forest and the water deficit, B) the second axis score of the forest and the latitudinal position. The regression lines, coefficients of determination, and significance level are given.

Although Banco has a high rainfall, and is relatively far from the coast (13 km) it still has been grouped with the coastal forests (Figure 5.4). The high rainfall is mediated by the low water holding capacity of its sandy soils, resulting in a low water availability. The most important environmental factor that determines this floristic group is the soil; the tertiary sands which underlie the forests, lead to a low water and nutrient availability. Each forest group possesses a fairly well defined group of characteristic species (Table 5.2), e.g., Khaya grandifoliola in semi-deciduous forests, Ancistrocladus barteri in the coastal forest, Mapania minor in the southwest forests, and two species of Guibourtia in the southeast forests. It is striking that almost one third of the

species occurs in only one of 13 forests and only 2% of the species occur in all sites. The floristic similarity between two forests varies from 29 to 77%, and is on average around 50%. In general, the floristic similarity is related to the distance between sites, their similarity of environmental conditions, or a combination of the two. All three cases were found in this study. For example, the neighbouring forests of Bossématié and Songan share almost 50% of the species despite obvious differences in the environment. The forests of the southeast and the southwest that are 400 km apart have a high floristic similarity, because of a high similarity in water availability. Subtle differences in rainfall and water deficit in these two zones are attenuated by the nature of their subsoils. The clay soils of the southeast (Perraud & Souchère 1970) have a greater water retention capacity than the sandy soils of the southwest (De Rham 1971), thus compensating for the lower rainfall in this region. As a consequence, plants experience a similar water availability in these two zones. Finally, forests which are geographically close and have similar soil and climatic conditions, as is the case for Mabi, Songan, Tamin, and Yaya, have the highest floristic similarities (Table 5.4). The paleoclimatic history is another reason why two forests might be similar. It has been postulated that forests in the extreme southwest and southeast Côte d’Ivoire have been part of two former glacial forest refuges (Guillaumet 1967, Wieringa & Poorter chapter 6). Wet forest species might have survived in these refugia during the dry glacial periods. Many species with regional or continental disjunct distribution patterns have populations in both of these two forest blocks (Holmgren et al. chapter 7). Summarising, the differences among Ivorian forests are governed by an intricate interplay between climate, soil and history.

Acknowledgements

We would like to thank SODEFOR for having allowed us to carry out research in its forest reserves. We are grateful to Claude Amani, Jean Assi, Saturnin Dougouné, Amadou Fofana and Patrice Mabéa for their assistance in the data collection. We thank Profs Laurent Aké Assi for identifying the taxa, and Dossahoua Traoré and Frans Bongers for their comments, which have allowed us to improve this chapter.

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6

J.J. Wieringa and L. Poorter

H A P T E R

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Introduction

The rainforests of West Africa have been earmarked as one of the world’s hotspots of biodiversity (Myers et al. 2000). These forests extend from Togo to Senegal, and are referred to as the Upper Guinean forests (White 1983). The Upper Guinean forests are separated from the rest of the African rainforests by the Dahomey gap; a woodland savanna which extends in Togo and Benin from the north to the Gulf of Guinea. Upper Guinean forests harbour a large number of endemic plant and animal species. It is estimated that about 2800 vascular plant species can be found in the Upper Guinean forests, of which 22% are endemic to the region (Jongkind, chapter 11). These forests are disappearing rapidly because of logging activities, shifting cultivation, and conversion into plantations (Chatelain et al. chapter 2). For an effective conservation policy, information is needed on the distribution of rare and endemic species in Upper Guinea, and the places in which they are concentrated. A problem of many tropical countries is that such botanical background information is scarce, or highly fragmented. To rapidly generate the necessary information one may carry out botanical surveys, in which selected areas are screened for their species composition. Hawthorne and Abu-Juam (1995) used such an approach in Ghana. Based on sample plots, systematically distributed over the whole forest zone, they were able to demarcate areas with a high share of endemic species. However, such an approach is labour intensive, and is only possible if a restricted floristic and geographic range is covered with a team of well-trained botanists. Another option is to use existing herbarium collections as a data source. Botanical collections have the advantage that they provide an existing source of information, cover a large geographic range, and that they are likely to be identified correctly. The latter is important, as especially the rare species are not easily recognised by tree spotters in the field. A disadvantage is that collection efforts are not evenly distributed over the area. Several authors have used the distribution patterns of small groups of plants or animals to indicate areas with a high biodiversity (Aubréville 1949, 1962, Hamilton 1976, Grubb 1982, Sosef 1994, Lovett et al. 2000). These studies typically point at three areas in Upper Guinea: the interior

Figure 6.1 The wet evergreen forest of Cape Three Points, Ghana. Cape Three Points is one of the three postulated Pleistocene forest refuges in Upper Guinea.

of Liberia with a diversity centre around Mount Nimba, Cape Palmas at the border between Liberia and Côte d’Ivoire, and Cape Three Points and its surrounding area in southwest Ghana. The exact location of the hotspots may vary, depending of the taxonomic group under concern (Conservation International 2001). A few biodiversity analyses on a larger number of species exist (e.g. Pomeroy 1993, Linder 2001), but they have a rather low resolution. Up to now a detailed, quantitative analysis based on a large number of species is therefore lacking for Upper Guinea. Both environmental and historical factors affect spatial variation in species richness. Species richness is known to vary along environmental gradients of rainfall (Hall & Swaine 1976, Currie 1991, O’Brien 1993), altitude (Hall 1973), and soil fertility (Hall & Swaine 1976, Huston 1980). In general, better site conditions lead to an increased primary productivity, more individuals and niches, and hence, a higher species richness. Species richness may decrease at very productive sites, giving rise to an unimodal relationship between richness and site productivity (Grime 1973, Huston 1979). Current floristic patterns are strongly shaped by largescale climatic disturbances in the past. Throughout the Quaternary, the area of rainforest waxed and waned with

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climatic fluctuations (Hamilton & Taylor 1991). During the dry and cool glacial periods, precipitation levels declined, and the rainforest contracted to small patches. As rainfall levels were low in Africa compared to other continents, the effect of the glacial period has been felt more strongly here than elsewhere (Richards 1973, Morley 2000) and it is likely that its imprint has lasted for a longer time. Testimony of the dynamic changes in vegetation cover are savanna relicts within the rainforest zone in Côte d’Ivoire (Gautier & Spichiger chapter 3), and the observation of fossils of Guineo-Congolean forest trees in Ethiopia (Bonnefille & Letouzey 1976). Some species could not keep up with the rapid expansion, and are still found in a narrow range around former forest refuges. The terrestrial forest herb Begonia mildbraedii, for example, has a disjunct distribution, with two isolated populations in Côte d’Ivoire and Ghana, being as far as 1800 km apart from its main, more widespread, distribution in central Africa (Holmgren et al. chapter 9, Sosef 1994). This chapter focuses on patterns and causes of plant biodiversity hotspots in West Africa. Hotspots are defined as centres with a high richness of rare and endemic species. The chapter explores how, and to what extent herbarium collections can be used to define hotspots of biodiversity. Then it relates biodiversity to environment and distance to postulated former forest refuges, and weighs the importance of environment and history in the current distribution of biodiversity.

Methods

Data collection Based on the 2nd edition of the Flora of West Tropical Africa (Keay 1954, 1958b, 1963, Hepper 1968, 1972), inventories in Ghana (Hall & Swaine 1981, Hawthorne 1995a), taxonomic revisions, and new herbarium collections, we made a compilation of just over 1000 species, which are rare or endemic to the closed forests of Upper Guinea (see Jongkind & Wieringa, chapter 11). All life forms were included (trees, shrubs, lianas, herbs, parasites, saprophytes and epiphytes). 640 species were selected for a shortlist to analyse biodiversity patterns. Care was taken to include species from different families, and to include species with different distribution patterns or ecology. Of these 640 species, herbarium specimens were entered for c. 510 species, and this was complemented by distribution data from taxonomic revisions for c. 130 species. We entered into a database all herbarium specimens collected from Senegal to Togo. For some non-endemic rare species, we also included the herbarium specimens collected in Lower Guinea. We entered all specimens from Herbarium Vadense (Wageningen, The Netherlands),

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Figure 6.2 Relationship between the observed number of species in a cell and the number of herbarium collections in that cell. The cells have a size of 55 x 55 km (n = 329). Two cells with a collection intensity larger than 500 per cell are not shown.

National Botanical Garden of Belgium (Meise, Belgium) and Kew Botanical Garden (Kew, Great Britain). In addition, we added the collections from Côte d’Ivoire present in the Geneva herbarium database (Conservatoire et Jardin Botaniques de la Ville de Genève, Switzerland) (only as far as the data was owned by Geneva), the collections from Côte d’Ivoire and Guinea present in the Paris herbarium database (Muséum National d’Histoire Naturelle, Paris, France), and the collections of 100 species present in the Herbarium of the University of Ghana (Legon, Ghana). For many non-endemic species, specimens from lower Guinea were entered as well. In total the database contained 48,000 records from West Africa, of which over 12,500 records correspond to rare or endemic species from our shortlist. The database covers all major herbaria that have collections from West Africa, with the exception of Paris which herbarium was closed for reconstruction during our data-entry period. For our area, we expect the Paris herbarium to contain mainly collections from Côte d’Ivoire and Guinea. Since we have abundant collections from Côte d’Ivoire from other sources, the real problem was the lack of data from Guinea. Cited collections made by Chevalier in Guinea were retraced using Chevalier (1911, 1920). Although our results for Guinea would have been more accurate with all data available, it is likely that they are good enough for those areas where the largest amount of forest species is to be expected (Fouta Djalon, Mt Ziama and Mt Nimba). The identification of specimens in different herbaria was cross-checked by C. Jongkind, who has ample knowledge of the taxonomy of West African forest species. The nomenclature was updated using Hawthorne & Jongkind (2004).

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Box 6.1. Species-collection curve To allow for comparison between regions, one may construct a species-collection curve, in a similar way as a species-area curve is made. For each grid-cell of half a degree latitude by half a degree longitude, we randomly drew collections, and plotted the cumulative number of species found, against the cumulative number of collections (Figure 6.3). In this way 100 species-collection curves were constructed and subsequently averaged. The number of species found in a given area increases with the number of collections made, until an asymptote is reached (Colwell & Coddington 1995, ter Steege 1998). Therefore we fitted per cell an asymptotic curve through the data, according to the equation Sn = (Smax* n) / (c+n), where Sn is the number of species in a sample of n collections, Smax the estimated maximum number of species in a given area, and c is a constant (Figure 6.3). For this analysis we confined ourselves to collections belonging to our shortlist of 640 species. The biodiversity estimates become more accurate when many specimens (N) have been collected relative to the number of species found (Sobs). We consider our biodiversity estimate to be fairly satisfactory if for a cell the ratio of N / Sobs is larger than 1.5. If N / Sobs < 1.1, then the biodiversity estimates vary considerably, and are too unreliable to use. We only fitted a curve through the data, if we had eight or more collections. For eight cells as many species were found as collections made, so curve fitting was not possible. For another seven cells N / Sobs was smaller than 1.1. This reduced the number of cells with regression results to 154.

How to analyse biodiversity patterns? To describe large-scale patterns in biodiversity, Upper Guinea should be divided into areas of equal sizes. What grid size should be chosen to describe those patterns best? A practical consideration is that there should be sufficient collections in each cell to make meaningful comparisons. A more theoretical consideration is that the grid size should not be too small, else local site conditions are likely to overrule the large-scale picture. As a compromise we selected a gridsize of 0.5 x 0.5 degree. Botanical collections have several advantages; they provide an existing source of information, are likely to be correctly identified (if the herbarium is not too small and if the species groups have been revised recently), and provide permanent records that can always be rechecked. A disadvantage is that specimen collection is not carried out in a stratified or random way. As a consequence, sampling efforts are not evenly (or randomly) distributed, and areas of high measured species density often coincide with areas of high collection intensity (Nelson et al. 1990). The same applies for the herbarium data from West Africa. If West Africa is divided in equal cells of 0.5 x 0.5 degree, then there is a large variation in collection intensity; the number of collections per cell for our shortlist species varies from 0 to 1303. The observed

Figure 6.3 Species-collection curve for the cell that includes Banco forest, Côte d’Ivoire. The species-collection curve is created by randomly drawing collections from the total collection pool in a 55 x 55 km area. The cumulative species number is plotted against the cumulative number of collections (circles). Subsequently a regression curve is fitted through the data using an asymptotic curve. By using the curve, the predicted species richness can be calculated (broken line), and the maximum number of species in the area (Smax) can be estimated.

number of species per cell increases in a curvilinear way with the number of collections (Figure 6.2, second degree polynome, r2 = 0.96, P < 0.001, n = 329). Sites with a larger number of collections appear to be species-rich, but this can largely be attributed to higher sampling effort there. The shape of the curve resembles the species-area curve (Gaston 1996), and the underlying sampling mechanism is somewhat comparable; the larger the area or number of collections sampled, the more species are found. To allow for comparison between regions, one may construct a species-collection curve, in a similar way a species-area curve is made (Box 6.1). From the speciescollection curves several biodiversity measures are derived. The S50 indicates the number of species found when 50 collections are made from the shortlist of 640 rare or endemic species (Box 6.2). The Smax is the estimated maximum number of species from our shortlist that can be found in a cell (Box 6.2). Finally, the rarity-weighted species richness (Srw) indicates areas with a large number of rare and endemic species (Box 6.3). For each of these three biodiversity measures maps have been made using the inverse distance weighting interpolation method in ArcView. Cells for which curve fitting was impossible (N = Sobs), or that were considered to be too unreliable to be used (N / Sobs < 1.1, see Box 6.1) were not included in

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Box 6.2. S50 and Smax; two measures of species richness The diversity of different grid cells can be compared, using a species-collection curve. Figure 6.4 shows such a species-collection curve for three different sites. In Tabou, the species number increases more rapidly with collection numbers than in Taï, which in turn is far more diverse than Banco. Tabou has therefore the largest diversity. Yet, this ranking in diversity is reversed, when one simply compares the total number of species found (Banco 192 species, Taï 160, and Tabou 119). The reason is that in Banco far more collections have been made than at the other two sites. According to our calculations in Banco 84% of the actually occurring species have also been found, while for Tabou this is only 38%. As it would be far too complicated to compare many species-collection curves, we use as a biodiversity measure the total number of species found at a standard number of collections. To this end we calculated, with help of the regression equation, the expected number of species at 50 collections (S50). This reference value of 50 is close to the median number of collections per grid cell (median = 35, range 8 - 1303), so that not too many extrapolations have to be made. S50 can vary from 1 (if all 50 collections in a cell belong to the same species) to 50 (if all 50 collections belong to 50 different species). If we would make an infinite number of collections, we would approach the maximum number of species (Smax) in a cell. The Smax indicates how many rare species are present in an area. Smax and S50 show a strong exponential relationship (r2 = 0.96, P < 0.001); therefore the larger S50, the larger Smax. At high levels of S50, a small increase in S50 will lead to a

the interpolation analysis. The interpolated biodiversity values are only shown for the forest zone. Relationship between biodiversity and environment Water availability, altitude, and soil fertility shape to a large extent the structure and composition of plant communities. Environmental variables used in the analyses were rainfall (in mm/yr), soil water holding capacity (WHC, in mm water/m soil), altitude (in m), soil fertility 2+ 2+ + (Ca , Mg , K , in cmol cations per kg soil), soil pH and cation exchange capacity (CEC in %). A rainfall map was created based on a compilation of 578 weather stations in the region. Data on soil fertility and water holding capacity were calculated based on the FAO soil map of Africa, and a quantitative review of chemical analyses of soil profiles (Batjes 1997). For each grid cell an average soil fertility was calculated, based on the relative cover of the different soil types, and their median soil fertility. Similarly, for each grid cell an average soil water holding capacity was calculated, based on the relative cover of the soil types, their depth, and texture. It was presumed that sandy soils have a water holding capacity of 75 mm/m, loamy soils of 100 mm/m, and clayey soils of 125 mm/m. For a detailed analysis of the environmental metadata, see

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Figure 6.4 Fitted species collection curves for 50 x 50 km areas around Tabou (continuous line), Taï (small broken line), and Banco (broken line). For each site the actual number of species observed in the area is indicated between parenthesis.

disproportionally large increase in Smax. The Smax is therefore a biodiversity measure that provides a large resolution at high levels of species richness. A disadvantage of Smax is that extrapolations need to be made. In incidental cases this may lead to an over- or underestimation of the real number of species present in the area.

chapter 9. All spatial analyses were carried out using ArcView (Esri Inc.). The S50, and Srw were related to environmental variables using a Spearman’s rank correlation and stepwise multiple regression. In addition to these abiotic factors, also the distances to the three proposed glacial forest refuges were used as factors in the analyses. To some extent, it might seem circular to relate the richness of endemic species to the distance to proposed forest refuges, as endemism and refuge theory are closely connected. However, the existence and location of these forest refuges has been postulated mainly on the basis of pollen cores and paleoclimatic data (Maley 2001), and on species with disjunct distributions between Upper and Lower Guinea (Aubréville 1949, Guillaumet 1967). To a lesser extent it has been based on the occurrence of species with restricted distribution ranges.

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Box 6.3. Rarity-weighted species richness S50 and Smax are general measures of species richness that do not take the "specialness" of the species into account. For conservation purposes, one might be interested to know where species are concentrated that are rare, or have a limited distribution. To this end we calculated the rarity-weighted species richness (Srw). It is a measure of species diversity, that gives weight to cells with many rare or endemic species. Each species is weighted for the number of half-degree grid-cells in which it occurs. A common species, which occurs in 100 half-degree grid-cells receives thus a weighting score of 1 / 100 for each cell in which it is occurs. A rare species which occurs in 4 half degree grid-cells receives a weighting score of 0.25. The rarity-weighted species richness is then the sum of the weights of all the species occurring in a cell. Yet, of all the species that occur in a cell (Smax), only a part has been sampled (Sobs). To correct for this, we multiplied the weighted measure with Smax/Sobs For example, if a grid cell A has 10 collections belonging to 4 species of which 1 species occurs in 1 cell, 2 in 2 cells and 1 in 4 cells, then the weighted richness of that cell equals (1 x 1 + 2 x 0.5 + 1 x 0.25) = 2.25. The predicted species richness (Smax) of the cell is 16, four times as much as actually observed. When correcting for sampling intensity, the rarity-weighted species richness equals 2.25 x (16 / 4) = 9. The rationale behind a rarity-weighted species richness is that, at a large scale (of Upper Guinea), each species is equally important, and receives a total weight of 1. We included 640 species in our analysis, so the sum of the rarityweighted richness of our species in all cells world-wide equals

640. Because some species also occur outside Upper Guinea, the sum of all cells in Upper Guinea is 498. Since we subsequently multiplied each cell with Smax/Sobs, the total amount of points assigned to Upper Guinean cells became 1406. Because Upper Guinea has a conservation value of 498 species points, we rescaled the rarity-weighted species richness of each cell by multiplying it with 0.354 (= 498 / 1406), to arrive again at our original score of 498. Table 6.1 gives an example of biodiversity calculations for two different cells. In cell A only a few, but very rare species have been found. In cell B more species have been found but they are also more common. Although less species have been found in A, its rarity-weighted richness is nearly four times as large as in B. The rarity-weighted richness of a cell may vary from close to 0 (if only one species is found that occurs in many grid-cells) to 640 (if all 640 species happen to be found in the same cell and nowhere else). An interesting feature of this biodiversity measure is that it puts the floristic value of the cell into a regional, Upper Guinea-wide perspective. If a cell has a rarity-weighted richness of 10, it might indicate that it contains 10 species that are found only here, and nowhere else. Alternatively, it might contain 20 species, that occur only here, and in another cell somewhere in the world. Each cell has of course a different conservation value for different species, but the rarity-weighted richness is a good measure of the combined value of a cell for the conservation and longterm survival of individual species.

Table 6.1 Example of the calculation of rarity-weighted species richness for a fictive cell with a few but very rare species (cell A), and a cell with many common species (cell B). For eight species it is indicated how many individuals are found in cell A and B, in how many cells they occur, and their weighting score (weight). Cell A

Cell B

N Sobs Smax Σ weight Σ weight x Smax/Sobs rescaled Srw

10 4 16 2.25 9.00 3.20

20 6 15 1.01 2.53 0.89

Species

# ind

# cells

weight

1 2 3 4 5 6 7 8 Σ

4 1 1 4

1 2 2 4

1.0 0.5 0.5 0.25

10

2.25

# ind

# cells

weight

1 3 3 2 1 10 20

2 4 10 10 20 100

0.50 0.25 0.10 0.10 0.05 0.01 1.01

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Figure 6.5 Distribution of herbarium collections (dots) of 640 rare and endemic forest species in West Africa. The potential forest zone is shaded.

Results

Collection efforts; where are the white spots on the map? Most collections of our shortlist of forest species are confined to the southern part of West Africa and indeed, closely follow the forest zone (Figure 6.5). Collection efforts have been particularly high near the capitals (Abidjan, Monrovia, Freetown) and near botanical research stations (see Appendix 5 for a map of West Africa). By far the highest number of collections have been made west of Abidjan. Both the Banco National Park and the research station in Adiopodoumé fall within this half-degree square, resulting in 1303 collections belonging to 190 species from our shortlist. Also the cell north of Abidjan, that includes Yapo forest and Téké forest, and the cell containing the Ecological Station at Taï score rather high. In Liberia, next to the surroundings of Monrovia the area around Mt Nimba has been well collected. In Sierra Leone three cells have relatively many collections: the Peninsula (including the capital Freetown), the forestry research station at Njala, and the Kambui Hills Reserve. In Ghana the Ankasa Forest Reserve and the Atewa Range are the cells with the highest number of collections attributed to our shortlist species. If we divide the observed species number by the predicted maximal species number, we get a measure of sampling intensity. Sampling intensity is particularly low (< 20%) in southeast Liberia (Figure 6.6). This area contains one of the largest remaining forest blocks in West Africa, and receives also a high amount of rainfall. It might therefore harbour a rich and unexplored flora, which definitely merits further attention. Southeast Sierra Leone, and southwest Ghana (north of Ankasa and Cape Three Points to Bia National Park) are two other large areas with a high potential species richness but few species sampled in our database. On a smaller scale it is striking that areas like the southeastern part of the Taï National Park and the Scio

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Figure 6.6 Sampling intensity (100 x Sobs /Smax) of 640 rare and endemic forest species in West Africa.

Forest Reserve are so poorly explored that we can not even estimate the sampling intensity. Hotspots of diversity There is a north-south gradient in species richness, which coincides with the rainfall gradient; the S50 (Fig. 6.7A) and Smax (Figure 6.7B) increase from the Sahel towards the coast. A belt of high rare and endemic species richness is found about 50-100 km inland, starting in Sierra Leone, running through Liberia to southwest Côte d’Ivoire and then fading away towards Sassandra. The climax of this range lies in Liberia and southwest Côte d’Ivoire. Additional rich areas, east of this belt are found around Abidjan and Ankasa (Ghana). A second belt of high rare and endemic species richness can be found more inland around the montane area of Mt Nimba with extensions to Mt Ziama in Guinea and the montane area around Man in Côte d’Ivoire. The Atewa Range in Ghana could be seen as a far-out exclave of this species-rich montane belt. Some areas in species-poor regions are relatively rich compared to their immediate surroundings or other areas with the same latitude. Examples are coastal areas in the Casamance region in Senegal and in Guinea-Bissau, the Fouta Djalon in Guinea, the Peninsula of Freetown in Sierra Leone, and Haut Sassandra Forest and Comoé National Park in Côte d’Ivoire (Figure 6.7A). Patterns in rarity-weighted richness are comparable to the ones for species richness (Figure 6.7C), although the richness of montane areas becomes more pronounced compared to the coastal areas. Examples are Mt Nimba, Mt Ziama, Mt Tonkui and the Atewa range. Within the coastal rich band also the wettest forests become more pronounced, which is expressed in very high values at Tabou and Greenville. Biodiversity vs. environment What environmental factors give rise to a high species richness? S50 shows a curvilinear relationship with rainfall;

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A S50

B Smax

C Srw

Figure 6.7 Biodiversity maps of West Africa showing A) the number of species at 50 collections (S50), B) the maximum estimated number of rare and endemic species (Smax) and C) the rarity-weighted species richness (Srw). Biodiversity values of cells are interpolated over the whole potential forest zone of Upper Guinea. The midpoint of the cell is indicated by an open symbol. For cells that partly cover the sea, the midpoint is positioned on the land surface. The size of the symbol is scaled to its biodiversity value. Cells without open symbols have less than eight collections. Cells for which curve fitting was impossible (N = Sobs) or that were considered to be too unreliable to use (N / Sobs< 1.1, see Box 6.1) were not included in the interpolation analysis, and are indicated by a cross. A full-colour version of this figure can be found in Appendix 4.

it increases up to an annual rainfall of 2000 mm whereafter it remains constant, to decline slightly again above 3000 mm (Figure 6.8A). S50 is negatively correlated with CEC (Figure 6.8B), CMK, and WHC (Table 6.2). Altitude and longitude are the only two environmental and topographical variables that are not correlated with species richness. The species richness decreases with distance to the postulated forest refuges Cape Palmas

(Figure 6.8C) and Mount Nimba, but is not correlated with the distance to Cape Three Points. In general, the environmental correlations are similar for S50, Smax and Srw (Table 6.2). Many environmental factors are closely associated, and show a similar spatial pattern. Rainfall and soil fertility are, for example, negatively correlated, as a significant amount of weathering and leaching has

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A

occurred under high rainfall conditions. It is therefore difficult to distinguish whether a correlation between S50 and soil fertility should be attributed to a causal effect of soil fertility, or is a spurious result because soil fertility and rainfall are closely associated. Moreover, it is likely that many factors act in concert and contribute to a high biodiversity. To evaluate which factors have an independent effect on biodiversity, we carried out a stepwise, multiple regression (Table 6.3). We included the environmental factors and their squares, to account for non-linear relationships. The first, and most important factor that explains variation in S50 is the distance to Cape Palmas. Rainfall emerges as the second-most important factor, and has a strong, non-linear effect on S50. Subsequently, the distance to Cape Three Points is included. In combination, these ecological and historical factors explain 56% of the variation in rare and endemic species richness in West Africa. Regression results are slightly different for Srw. The distance to Cape Palmas and rainfall are here as well the most important variables, but subsequently the distance to Mount Nimba and the interaction between rain and WHC are included. In combination, these factors explain 44% of the variation in rarity-weighted species richness.

Discussion B

C

Figure 6.8 Relationship between species richness (S50) and A) annual rainfall, B) cation exchange capacity, and C) distance to Cape Palmas. The sample units are cells of 55 x 55 km (n = 161). For Figure 6A, the regression line is fitted excluding the cell (represented by a triangle) with a very high rainfall but low species richness.

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Patterns in biodiversity The richness of rare and endemic forest species closely follows the forest zone, with the exception of Sierra Leone, and the Fouta Djalon in Guinea. Within the forest zone, species richness increases towards the wetter forest types, near the coast (cf. Hawthorne 1996). The Srw is especially high in the wettest forests between Greenville and Tabou, which harbour many narrow endemic species (Holmgren et al. chapter 7). This supports the biodiversity analysis made by Beentje et al. (1994) for Africa. They concluded that endemism in Upper Guinea is centred on the LiberiaCôte d’Ivoire border. Several other areas mentioned by them as centres of diversity and endemism (e.g. Gola Forest in Sierra Leone and Loffa-Mano Forest in Liberia) fall within our rich sub-coastal forest band, but in our analysis these areas do not appear to be separate areas; they are part of the same rich band. Given the climatic conditions, more than the half of Sierra Leone could be covered with rainforest (Harcourt et al. 1992). Because of high deforestation rates in the past, only isolated forest patches are left, which still harbour many typical forest species. A second band of high biodiversity follows the mountains of the Guinean belt. The higher richness of these mountain zones is partly explained by a high orographic rainfall. A fair amount of species is restricted to these mountains because they are adapted to high altitude

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conditions such as daily fog and low temperatures. Especially epiphytes will prosper under such conditions. Not less than 13 endemic species and many near-endemic species are found on Mount Nimba alone (Beentje et al. 1994). Montane forests are relatively rare in West Africa compared to lowland forest, or to montane forests in South America or Southeast Asia. For this reason, montane species may have a very small potential distribution range (Figure 7.4C, cf. Gentry 1992). In addition some slow-dispersing lowland forest species may have had their glacial refuge on the relatively wet foothills of these mountains and occur nowadays only in their surroundings. Together, these factors may explain the increased importance of these mountain areas in the rarityweighted richness (Srw) compared to S50. Mountains like Mt. Nimba, Mt. Ziama and Mt. Tonkui harbour not only many highly endemic plant species (Holmgren et al. chapter 7), but also endemic mammals, reptiles, birds and insects (Conservation International 2001). These mountains should receive extra attention in conservation

schemes, because they are essential for the existence of these species. The Atewa Range, and in a lesser extent the Southern Scarp and the Togo Plateau in Ghana, are also more pronounced in the Srw analysis. These mountains share some species, such as Begonia cavallyensis and Vernonia titanophylla with other mountainous areas in Upper Guinea. This strengthens the idea that they could be regarded as a far extension of the mountain high diversity belt. A similar biogeographical connection between Mt. Nimba and the Atewa range can be observed in land snails (A.J. de Winter pers. comm.). The Atewa Range also thanks its richness by providing a habitat for evergreen lowland forest species such as Albertisia scandens and Aframomum atewae. Mt Loma in Sierra Leone is not characterised by a higher diversity in forest species compared to its surroundings. This is in contrast to what has been suggested by Jaeger and Adam (1946) who treat this mountain as similar to Mt Ziama, Mt Nimba and the Man area. Beentje et al. (1994), based on Schnell (1983),

Table 6.2 Spearman’s rank correlation between biodiversity, environmental and topographical variables, and the distance to three postulated forest refuges (n = 148 cells). ns = not significant; * = P < 0.05; ** = P < 0.01; *** = P < 0.001.

S50

Smax

Srw

Variable

rs

P

rs

P

rs

P

Rain WHC Altitude CMK CEC pH Latitude

0.49 - 0.39 0.16 - 0.39 - 0.48 - 0.19 - 0.40

*** *** ns *** *** * ***

0.49 - 0.40 0.12 - 0.38 - 0.47 - 0.19 - 0.45

*** *** ns *** *** * ***

0.43 - 0.35 0.12 - 0.29 - 0.40 - 0.12 - 0.45

*** *** ns *** *** ns ***

Longitude Distance to C. Palmas Distance to C. Three Points Distance to Mt. Nimba Distance to nearest refuge

- 0.06 - 0.56 - 0.07 - 0.51 - 0.56

ns *** ns *** ***

- 0.05 - 0.59 - 0.10 - 0.50 - 0.57

ns *** ns *** ***

0.01 - 0.52 - 0.14 - 0.40 - 0.51

ns *** ns *** ***

Table 6.3 Results of a multiple stepwise regression of S50 and Srw on environmental variables and refuges. Order indicates the sequence in which the variables are included in the regression analysis, slope the coefficient of that variable, r2 the cumulative variance explained by the model, and P the significance level. Thirteen variables were included in the analysis (Rain, Rain2, WHC, WHC2, Altitude, Altitude2, CMK, CMK2, CEC, CEC2, the interaction between Rain and CMK, the interaction between rain and WHC, and the distance to Cape Palmas, Cape Three Points, and Mount Nimba) that might effect species richness. CMK and CEC were log-transformed prior to analysis. N = 148 cells. * = P < 0.05; ** = P < 0.01; *** = P < 0.001.

S50

Srw

Order

1 2 3 4 5

Variable

Slope

rs

C. Palmas Rain Rain2 C. Three Points -

- 1.0 0.3 - 0.5 - 0.8

0.38 0.45 0.50 0.56

P

Variable

*** *** *** ***

C. Palmas Rain Rain2 Mt Nimba Rain x WHC

Slope - 0.62 0.06 - 0.11 0.44 - 5 x 10-7

rs

P

0.29 0.35 0.38 0.42 0.44

*** *** *** *** *

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mention it as a centre of endemics for the GuineoCongolian region. However, all nine endemic species of Mt. Loma are species from mountain grassland or rocky areas (Schnell 1983). Mt Loma may therefore be a centre of endemism for mountain grassland flora, but not for forest flora. Some isolated areas have a higher biodiversity than their immediate surroundings. For the coastal regions of Guinea-Bissau and the Casamance region in Senegal this might be attributed to an extremely high but strongly seasonal rainfall. The Fouta Djalon harbours a relatively large amount of species for its latitude, probably caused by the high rainfall in this elevated area. Similarly, the high diversity in the Peninsula area of Freetown, Sierra Leona, might be due to the coastal fogs that give rise to humid conditions. Another reason for its relatively richness might be that it harbours one of the last forests left in Sierra Leone (Davies 1987). The rest of western Sierra Leone may have been rich, but this has not been well recorded at the time. In Côte d’Ivoire two areas strike as being richer than expected. The Haut Sassandra forest harbours a remarkable range of both wet and dry species, which is not yet explained (Kouamé 1998a). The Comoé National Park is not particularly rich, but is surrounded by areas which do not contain rare species at all. The park harbours riverine species and some rare species that are restricted to dry forests such as Afraegle paniculata (Porembski 1991). The extremely dry forests that border the Dahomey gap in Ghana, harbour few, but highly endemic dry forest species such as Talbotiella gentii, Turraea ghanensis and Hunteria ghanensis (Hawthorne 1996). This challenges the traditional view that only wet forests have a high conservation value and stresses that these forests have unique species that need to be conserved. Biodiversity and rainfall We used fairly large grid cells to describe regional patterns in biodiversity. Local, regional, and landscape diversity contribute to the species richness of the grid-cell. To understand which factors govern variation in regional diversity, we should look at environmental factors that operate at this spatial scale (O’Brien 2000). In Upper Guinea there is a strong rainfall gradient that varies from 1000 mm at the forest-savanna boundary to 4000 mm at the coast. Species richness increases along this rainfall gradient, showing an optimum around 2500 mm, whereafter it may level-off or decline. An increase in tree species richness with rainfall has also been found in other studies (Hall & Swaine 1976, O’Brien 1993, Clinebell et al. 1995). Two mechanisms may give rise to this pattern. Over the first part of the rainfall gradient, an increase in forest height and structural complexity provides more niches, thus allowing more species to coexist. Another reason is that in addition to the drought tolerant species, also more drought intolerant life forms and species are able to persist at higher rainfall levels (Huston 1994). The shape of the biodiversity-rainfall curve depends

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on the section of the rainfall gradient regarded. In southern Africa the species richness of dry vegetation formations shows a strong, linear increase with rainfall between 200 to 1000 mm (O’Brien 1993). For the closed forest formations in the Neotropics this linear relationship is maintained between 1000 and 2000 mm, above which it disappears (ter Steege et al. 2000b). One may argue that under perhumid conditions water availability is not a limiting factor for species richness anymore. However, it is very likely that the biodiversity-rainfall relationship is to a large extent scale-dependent. At a macro scale the potential species richness is determined by the climate (Whittaker et al. 2001), at a meso scale on the configuration of different vegetation types within the landscape, whereas at a micro scale, the realised species richness depends on local site conditions. Ter Steege et al. (2000b) evaluated the diversity of 1-ha forest plots in the Neotropics. The maximum species richness at a given amount of rainfall indeed shows a linear increase with rainfall up to 3500 mm. Adverse local site conditions (such as swamps, flooding, extreme soil conditions) may prevent the forest from attaining its climatic potential, leading to a lower diversity and a specific flora. Biodiversity-rainfall relationships are also likely to vary with the life forms considered. Tree species diversity increases with rainfall, and levels-off at higher rainfall levels, while the diversity of epiphytes, mosses and understorey herbs still increases (Gentry 1988). Our species list includes a diverse array of life forms (trees, shrubs, lianas, herbs, epiphytes) but has some emphasis on woody plants. It is therefore likely that it shows a saturating response to rainfall. Our study is the first to show a potential decline of biodiversity at very high rainfall levels (>3000 mm). It is not clear to what extent this is a spurious result. Many of these extremely high rainfall sites are situated along the coast of Liberia, Sierra Leone and Guinea. The coastal grid cells cover only part of the land surface, so fewer habitats and less landscape diversity might be sampled, resulting in lower diversity values for such cells. This is supported by the fact that coastal areas are in general less rich than adjacent continental areas (Figure 6.7). The coastal zone in Guinea and Sierra Leone also receives a high amount of rainfall which is unevenly distributed over the year. The length and severity of the dry season is likely to be a stronger determinant of humidity and species richness than rainfall per se. If the decrease in diversity with rainfall is not spurious, this might pinpoint to other mechanisms. First, some species might be intolerant to waterlogged conditions. This hypothesis is supported by the fact that a high rainfall, in combination with a high water holding capacity leads to a reduced diversity (Table 6.3). Second, very humid areas, with an annual rainfall larger than 4500 mm do not exist in West Africa. Species that require very wet conditions (i.e. >4000 mm) are likely to have populations that are too small to persist, or even to have gone extinct during the dry glacial periods. Strikingly,

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many species with small populations are found under very high rainfall conditions, at the Liberian coast (Holmgren et al., Chapter 7). These may be examples of species adapted to a rainfall of e.g. 3000-8000 mm, which can now only occur at the lower end of that range. Biodiversity versus soil fertility Species richness decreased with soil fertility. In Ghana, Hall & Swaine (1976) found the same pattern for small 25 x 25 m plots, in which they inventoried all vascular plant species. On fertile Costa Rican soils, tree species richness of 0.1 ha plots also declined with soil fertility. Soil conditions vary considerably over small spatial scales along the catena gradient. It is therefore surprising that we find such a highly significant relationship with soil fertility for large areas (55 x 55 km) and relatively coarse soil data. However, soil fertility explains relatively little in the variation in species richness after rainfall has been accounted for. Species richness is expected to decline with soil fertility, because of a stronger competition for light at high soil fertility. Slow-growing species are outcompeted by fast-growing, dominant species, leading to a lower diversity at the stand level. Comparable trends have been found for temperate grasslands (Grime 1973, Huston 1979). Forest refuges Surprisingly, the distance to postulated refuges was the foremost important factor explaining spatial patterns in biodiversity. This suggests that after 12,000 years, the glacial period has still left its imprint on the African landscape. In fact, several of such glaciations have occurred over the past 10 million years (Morley 2000). Many drought-intolerant species might have gone extinct during such glacial periods, leading to an impoverished flora between former rainforest refuges. At the same time, speciation might have occurred in the isolated forest refuges. Relatively young, vicariant species that occur in West Africa (Schizocolea linderi and Berlinia occidentalis) and central Africa (S. ochreata and B. bracteosa), might still be testimony of that period. Many other vicariant species relationships may exist between Upper and Lower Guinea, however, with no molecular phylogenies and only very little morphological ones available it is not possible to tell how important this phenomenon is for our area. The few phylogenies available (e.g. Wieringa 1999) show that Upper Guinean species are often sister taxa to small groups of Lower Guinean species. This might indicate that speciation set off after the first Pleistocene glaciations dividing the ancestral species into two over the Dahomey gap. Subsequently, splitting up in more species could take place only in central Africa, probably due to the higher number of glacial refuges and a larger forested area. Upper Guinean species are genetically more distinct than their Lower Guinean counterparts. They would, therefore, become more important if phylogeny would be included in biodiversity calculations.

After the last glaciation the rainforest expanded rapidly, attaining its maximal extension around 6000 BC (Hamilton 1976). Some species could not keep pace with this fast expansion, and still have a restricted, disjunct distribution pattern. Examples are Begonia hirsutula, Hemandradenia chevalieri and the two disjunct subspecies of Aphanocalyx microphyllus. Interestingly, there is a tight correlation between drought tolerance and the maximal distribution range of about 300 West African tree species (Holmgren et al., Chapter 7). The most drought intolerant species have the smallest distribution. This might suggest that the “wettest” species, that survived in the former rainforest refuges still have not recuperated their full potential distribution range. Alternatively, it might indicate that the distribution of these species is shaped by present environmental conditions, instead of by the past. Three forest refuges exist, of which Cape Palmas is the most important one. Cape Three Points appears to be another forest refuge (Table 6.3). Yet, in our analysis the forests in the southwest corner of Ghana (i.e., around Ankasa Game Production Reserve) seems to be more species rich than Cape Three Points (Figure 6.7). This is also confirmed by floristic surveys in which Ankasa and Neung forest reserve are richer than Cape Three Points (Hawthorne & Abu Juam 1995). A more detailed analysis, however, shows that the hillside samples in Cape Three Points are the second and third hottest patches of forest yet sampled in Ghana. The strong rainfall gradient at the coast, and topographic relief probably account for this steep gradient in biodiversity (Hawthorne & Abu Juam 1995). Reviewing the current evidence, it must be concluded that there has been a minor forest refuge in southwest Ghana, including Cape Three Points, Ankasa, or both. A third refuge is Mount Nimba, which is especially important for rare species associated with mountain habitats. Mountains are relatively isolated and rare in West Africa. Migration between these mountain areas might have occurred in the past during cooler, glacial periods, or at present, by means of jump dispersal (Hall & Swaine 1981). Summarising, the use of species-collection curves has allowed us to get a first glimpse of large-scale biodiversity patterns in rare and endemic West African species. Such species are to some extent confined to special habitats. We believe, however, that common, non-endemic species are either quite ubiquitous, or show similar responses to the environment as endemic species (e.g. Lovett et al. 2000). The observed biodiversity patterns for rare and endemic species might therefore be representative for biodiversity patterns in general. Plant biodiversity in Upper Guinea is to a large extent shaped by rainfall, orography and history. The relative aridity of Africa, in combination with a strong glacial period might explain why forest refuges are more clearly observed here, than in other continents (Richards 1973, Beentje et al. 1994, Morley 2000). In concert the strong historical imprint and steep rainfall gradients have lead to the remarkable biodiversity patterns currently observed in West Africa.

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Acknowledgements

We thank Marjo Buitelaar, Hendrikjan Os Breijer, Stuart Cable, Patrick Ekpe and many students for the arduous work of entering over 12,500 herbarium

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collections into the database. Without them this work would not have been possible. The herbaria of Paris and Geneva kindly provided part of their data. Carel Jongkind checked many of the species identifications, Toon Helmink and Roland van Zoest prepared the environmental maps for the GIS database, and Frans Bongers, William Hawthorne, Milena Holmgren and Hans ter Steege gave helpful comments on the manuscript.

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M. Holmgren, L. Poorter and A. Siepel

What explains the distribution of rare and endemic West African plants?

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Understanding species distribution patterns is a crucial step for the conservation and management of plant communities and ecosystems. This is particularly true for species with small distribution ranges that grow and reproduce under a very particular set of environmental conditions. West African forests, although less species rich than some other tropical forests, seem to be relatively rich in endemic species (e.g. Myers et al. 2000, White 2001). About 22% of the forest species in Upper Guinea are restricted to this region (Jongkind & Wieringa Chapter 11). Because of the rapid rate at which these forests are disappearing (e.g. Chatelain et al. chapter 2), it is urgent to know the characteristics of the species distribution and the mechanisms responsible for them. In Upper Guinea distinctive forest types occur along a west-east coastal gradient from Guinea to Togo. There is also a clear zonation of forest types from the coastal forests towards the northern savanna (Hall & Swaine 1976, Martin 1991, Bongers et al. chapter 4). The change in forest types along both gradients has been explained mainly by a strong rainfall gradient (Hall & Swaine 1976, van Rompaey 1993). At the species level, we also know how some forest species are distributed along environmental gradients. In most cases, species occurrence or density has been found to be significantly correlated with water availability (Bongers et al. 1999), although soil fertility (Swaine 1996) or, at a smaller spatial scale, light availability (Veenendaal et al. 1996c, Agyeman et al. 1999a), are sometimes better explanatory variables. The interpretation of species distribution patterns along environmental gradients is difficult. This is partly because environmental gradients tend to covary. For example, it is hard to separate the effects of changes in rainfall and soil fertility because under high rainfall conditions, nutrient leaching causes a loss of soil fertility as well (e.g. Swaine 1996). Rainfall effects can be also complicated by the interaction with altitudinal and topographical gradients. For example, tree species largely confined to high rainfall forests can be found in forests with lower rainfall, but are restricted to the lower and therefore wetter, topographical places (Guillaumet & Adjanohoun 1971, de Rouw 1991, van Rompaey 1993). Field experiments can play a crucial role in separating the relative importance of different environmental variables. In

Figure 7.1 Cola attiensis, a small understorey tree species with a disjunct distribution between Upper and Lower Guinea.

general, they support the predominant role of seasonal drought over soil fertility to explain the limits of species distribution in these forests (e.g. Veenendaal & Swaine 1998). Clearly, the distribution of any species along these environmental gradients will be partly the result of its ecophysiological tolerances combined with the effects exerted by the interaction with co-occurring species. The actual distribution of species along present environmental gradients may also reflect historical changes on longer time scales. The Upper Guinea region has been exposed to strong climatic changes and severe humaninduced disturbances (Fairhead & Leach 1998). Rainforest cover has periodically retracted during glacial periods, when the climate was drier and cooler, and expanded during warmer and wetter interglacial episodes, to cover surfaces even greater than the present (Livingstone 1982). In this region, glacial-interglacial climatic changes were probably stronger than in any other tropical forest around the world because of the relatively low rainfall here compared to the other continents (Richards 1973, Morley 2000, Maley 2001, White 2001). Furthermore, these West African forests have been exposed to a longer, and recently faster, rate of deforestation and degradation than other rainforests around the world (FAO-UNEP 1981, FAO 2001). These past forces must have left a deep imprint on the composition and structure of these West African

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forests. Climate change and deforestation have produced not only a change in the general climatic and edaphic conditions for which the ecophysiological tolerances of the species might differ. But they have also increased the fragmentation of the landscape and we could expect differences in dispersal capabilities between species to play a fundamental role in explaining the extension of their distribution range. In summary, the present distribution of a particular species would be the combined result of ecophysiological tolerances, the opportunities to disperse, the interactions with other species, and the effects of past changes in climate and disturbance. Most of our knowledge on the distribution of plant species in the Upper Guinea forests comes from the outstanding work performed in Ghana (e.g. Hall & Swaine 1976, Hawthorne 1996, Swaine et al. 1997), and to a lesser extent from other countries of the region (e.g. Guillaumet 1967, Schnell 1977, Aké Assi 1984, van Rompaey 1993). The only work covering the whole Upper Guinea region is the Flora of West Tropical Africa (Keay 1954-1963), which has a strong taxonomical focus. In general, most of the emphasis has been on taxonomical descriptions or studies confined to one country. A renewed regional perspective can bring new insights into the amplitude of the species distributions and the mechanisms behind them. In this chapter, we describe the spatial distribution patterns and discuss the potential responsible mechanisms for a large set of endemic and rare plant species in the Upper Guinea region. We use herbarium collections from the whole region and relate the inferred distribution patterns to the main environmental variables and species life-history traits.

Methods

Species selection Approximately 2800 species of vascular plants have been recorded in the forests of Upper Guinea, 22% of which are endemic to this region (Jongkind, Chapter 11). As a first step we made a selection of 1000 endemic and rare species of these forests based on the second edition of the Flora of West Tropical Africa (Keay 1954-1963), forest inventories, and taxonomic revisions. The emphasis was on woody species (trees, shrubs and lianas), although some herbs were included as well. From this selection, we prepared a short-list of 600 species that have been taxonomically well described and for which there is enough certainty about their correct identification in the different herbaria. From this short-list we selected a subset of 286 species that are individually described in chapter 9, and for which the distribution patterns are discussed here. In selecting this sample we aimed at preserving as much taxonomical diversity as possible. This means that we

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selected species from all families taking a smaller fraction of species from the large families (with many species in this area). The 286 species of this final subset represent c. 10% of the total forest flora of Upper Guinea (with 223 species considered endemic to Upper Guinea). They belong to 75 different families. Amongst these species, there are 126 trees, 35 shrubs, 79 lianas, 36 herbs, nine epiphytes, and one saprophyte. This means that our sample is relatively rich in trees, since in the Upper Guinean flora the number of trees, shrubs and lianas is rather similar (W.D. Hawthorne, personal communication). Species data We entered into a database (Brahms 4.8, Filer 2000) all herbarium specimens collected from Senegal to Togo, for all 600 short-list species (including our 286 species). For some non-endemic rare species, we also included the herbarium specimens collected in Lower Guinea (the Congo Region). We checked and entered all specimens from Herbarium Vadense (Wageningen, The Netherlands), National Botanical Garden of Belgium (Meise, Belgium) and Kew Botanical Garden (Kew, Great Britain). In addition, we added the collections from Côte d’Ivoire present in the Geneva herbarium database (Conservatoire et Jardin Botaniques de la Ville de Genève, Switzerland), the collections from Côte d’Ivoire and Guinea present in the Paris herbarium database (Muséum National d’Histoire Naturelle, Paris, France), and the collections of 100 species present in the Herbarium of the University of Ghana (Legon, Ghana). These data were complemented by distribution data from taxonomic revisions. In total the database contained 48,000 records from Upper Guinea, of which about 6900 records concern our 286 rare or endemic species. This database is not exhaustive, but gives a fair indication of the state of knowledge for the species in the region. Environmental data Species distributions are usually strongly shaped by water availability, soil fertility, and altitude. Maps of these environmental variables were prepared and included in a Geographical Information System (ArcView, ESRI Inc.). Water availability is determined by rainfall, soil water holding capacity, and the presence of rivers, lakes, and proximity to the coast. Annual rainfall data were compiled from different sources (Myers & Staff 1981, ANAM 1987, ICCARE 1994, Global Historical Climatology Network). In total, 578 weather stations were included from Senegal to Nigeria, Burkina Faso, Mali and Niger. From these data a rainfall map was made, using the inverse distance weighting interpolation method in ArcView. The soil water holding capacity was derived from the FAO Digital Soil Map of the World (FAO 1995b). The soil map has a scale of 1 : 5,000,000. Therefore small-scale differences in soil types due to topographical variation are not visible on this map. For each soil type, we calculated the water holding capacity of the soil (in mm water/m soil) based on the

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depth and the texture of the dominant soil type. The position of perennial rivers, lakes, and coastline was obtained from the Digital Chart of the World (ESRI 1984). As a measure of soil fertility we used the sum of exchangeable cations (CMK; Ca2+, Mg2+, and K+, in cmolc/kg). For each unit of the FAO Digital Soil Map we used a median CMK value of the dominant soil type. The CMK values were based on a review of the chemical properties of soil profiles by Batjes (1997). Elevation data were derived from the Digital Chart of the World (US Geological Service, EROS Data Center, 1984). The elevation map has a spatial resolution of approximately 1 km. In addition, we calculated the percentage of records of each species found in open habitats and in moist habitats. Using the herbarium collector’s notes that had been added to our database, we searched for keywords for open places (e.g. gap, forest edge, roadside, etc.) and moist places (riverside, close to lake, stream, etc.). To estimate the proportions in particular habitats we used only the subset of records for which clear habitat descriptions were present. Individual species occurrence along environmental gradients A logistic regression analysis was carried out to model the species occurrence in Upper Guinea as a function of four environmental factors; altitude, annual rainfall, soil water holding capacity, and available cations. We used stepwise forward regressions with the simple and quadratic variables, but without including interaction terms. A significant simple variable indicates an increasing or decreasing probability to find the species with an increase in the environmental variable. A significant quadratic variable indicates that the species shows a bell-shaped response curve towards that environmental variable (Jongman et al. 1987). Since the species had not been collected at random and since we had only presence data (herbaria records), we needed to assume absence under certain conditions. For this we selected for Upper Guinea only those half-degree grid cells with 50 or more records of all species occurring in the area (thus including only cells that have been botanically well explored). This resulted in a total of 145 cells. For some cells there were no data for all four environmental variables. This reduced the total number of cells to 140. We assumed that a species was absent when no records of this species were found in those cells. A logistic regression analysis has only sufficient resolution to detect significant patterns, when there are sufficient numbers of presence and absence. In order to have a reasonable number of cells with presence, we ran the analysis only on those species present in ten or more cells (i.e. > 7% of the total amount of cells). This resulted in a total of 112 species. Species distribution, range, and commonness along environmental gradients The species were classified into different distribution types based on the continuity of their distribution, their

range, and their commonness. We distinguished between species having a continental disjunct distribution (disjunct between Upper and Lower Guinea), a regional disjunct distribution (disjunct populations within Upper Guinea), and a continuous distribution (a more or less continuous occurrence within its range). The classification was based on a visual interpretation of the species distribution map, and a histogram of multiple neighbour distances of the species. To this end, we calculated for every species the distance between each collection point and all the other collection points of that species. For a species with five observations this would result in ten neighbour distances. A histogram of the frequency of these neighbour distances showing a unimodal distribution was interpreted as an indicator for a continuous distribution (Fig. 7.2A). On the other hand, species with a bi-modal histogram were interpreted as having a disjunct distribution (Fig. 7.2B,C). Every mode or peak represents a clump, with the height corresponding to the number of collections within a clump. The distances between the peaks correspond to the distances between the clumps. Obviously, continental disjunct species (e.g. with a population in Liberia and a population in Gabon) usually have a much larger distance between the two peaks than regional disjuncts (e.g. with a population in Liberia and a population in Ghana). The classification of spatial patterns based on the multiple neighbour distance histograms was in close agreement with a visual interpretation of the maps. For species with a continuous distribution, we estimated the species range within Upper Guinea based on the maximum distance found between collection points. We classified the species range as being very local (maximum distance <100 km), local (100- 300 km), regional (300- 600 km), or widespread (>600 km). Our very local and local categories fit the most widely used criterion of tropical plant species endemism (Gentry 1992). The commonness of a species in Upper Guinea was expressed in two different ways. The abundance is the total number of records found in the region from Senegal to Togo. This measure is potentially influenced by collector’s bias. Rare species are likely to be over-represented, as they might be more interesting to collect by botanists. A more robust measure of commonness is the frequency of halfdegree grid cells in which the species is collected. If a species had been collected many times in the same forest reserve close to a capital or research station, this would result only in one observation of a half-degree grid cell in which it is present. In our dataset, both measures are highly correlated (Pearson’s r = 0.92, p < 0.001, n = 286) and therefore provide consistent estimates of a species commonness. For subsequent analysis of commonness, we only present the results for the total number of records. In order to find general patterns among species with different distribution types, ranges, and commonness we performed a series of statistical analyses and selected different species subsets of the database depending on the questions asked. We compared both species distribution

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A

B

C Figure 7.2 Types of species distribution patterns. A) Continuous distribution (Combretum grandiflorum), B) continental disjunct distribution (Gymnosiphon longistylus), C) Upper Guinea disjunct distribution (Acanthus guineensis). Distribution maps indicate the locations of the species collections (dots), the potential forest cover (grey), areas above 500 m altitude (dark grey), and country boundaries. Next to the map is the histogram of distances between each collection and all other collections for that particular species.

types (i.e. continuous versus disjunct) for several environmental variables. No distinction was made between Upper Guinea and continental disjuncts in order to increase the sample size for the disjunct group. The variables for each species belonging to either distribution group were: rainfall (minimum, maximum, median, range,

76

10-percentile), altitude (% of records of a species found higher than 500 m), habitat openness (% of records found in disturbed open places), habitat moisture (% of records found in moist places). For each variable, we performed a t-test to compare the mean between both distribution groups. We used two criteria to select the species to be

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used in the comparisons. For the rainfall and altitude data we used species with more than five records in the database. For the habitat openness and moisture, we used all species for which we had more than ten records in the database with enough habitat information provided by the specimen collector and calculated the percentage of records in either disturbed open places or moist places. For all subsequent analyses, relating a species range and commonness, we confined ourselves to species that have a continuous distribution, that are endemic to Upper Guinea (i.e. only occurring between Senegal and Togo), and that have more than five records in the database (n = 163). We used a multiple regression to relate a species range (maximum distance between collections) with the various environmental variables. We included only the environmental variables that were not strongly correlated with each other, namely minimum rainfall, maximum rainfall, altitude, percentage of records in open places, and percentage of records in moist places. To select the variables in the final model, we used both stepwise forward and backward selection. In both cases we obtained the same results. In addition, we used one-way Anovas to compare the four distribution range categories (widespread, regional, local, and very local) in relation to each environmental variable. We related species commonness (a species total number of records, and the number of half-degree cells in which the species has been sampled) with the environmental variables using multiple regression analysis. We used the same environmental variables and data selection criteria as for the range analysis. Distribution type, range and commonness in relation to a species life history traits Based on the literature, we classified the species according to life form (herbs, trees, shrubs, lianas, and epiphytes). The main dispersal mechanisms were: animal, wind, water, explosive (active expulsion of the seeds), and barochore (simply falling). We also classified the species into guilds as pioneer, non-pioneer, and shade bearers based on the reported shade tolerance in Ghana (Hall & Swaine 1981, Hawthorne 1995a). Pioneers are species with the highest light demand, of which seedlings are only found in gaps and older plants are absent or very rare in the forest understorey. Non-pioneer light demanders are species with an intermediate light demand, of which seedlings are common in the understorey whereas adults are not (i.e. they require a gap to grow). Shade bearers are those species for which both young and older plants are frequently found in the shaded forest understorey. We compared the percentage of records found in open places in our database with the classification made by Hawthorne (1995a). Indeed, we found that pioneer species had a higher abundance in open places, but we did not find a difference between shade bearers and light-demanding species (t-test, t = – 2.37, p = 0.026). Based on this, we classified the species lacking a literature description on

shade tolerance using the proportion of herbarium records found in open places. We classified them as pioneers (> 65% records in open places), non-pioneer light demanding species (30-65% records in open places), shade bearers (< 30% in open places). We related a species’ life history traits (life form, dispersal mechanism, and shade tolerance) with the occurrence in particular habitats (e.g. % of records in moist places) using one-way Anovas. The effect of these life history traits on the distribution type was done through Chi2 frequency analysis, and the relationship with the range and the abundance was done through one-way Anovas.

Results

Species distribution types Three different types of species distributions were recognised (Fig. 7.2); continuous, continental disjunct, and Upper Guinea disjunct. Combretum grandiflorum is an example of a species with a continuous distribution; it has a regular occurrence all over its range, and a neighbour histogram without clear peaks (Fig. 7.2A). Gymnosiphon longistylus is an example of a species with a continental disjunct distribution. It occurs in Upper Guinea and in Cameroon-Gabon, and has a neighbour histogram with two clear peaks, spaced 2000 km apart (Fig. 7.2B). Acanthus guineensis is a species with an Upper Guinea disjunct distribution. It occurs around Liberia, and in Ghana, and does not occur in between. The neighbour distance histogram shows two clear peaks, which are spaced approximately 1200 km apart (Fig. 7.2C). From a total of 270 species, 86% showed a continuous distribution, 10% a continental disjunct distribution, and 3% an Upper Guinea disjunct distribution. For 16 species, there were not enough data points (< 5 collections in Upper Guinea and a neighbour distance greater than 100 km) to classify their distribution. There were four distinctive patterns among the 28 species with a continental disjunct distribution (sensu White 1979, Table 7.1). The most frequent group of “Guinea wide” (eleven species) occurred in both Upper and Lower Guinea (Fig. 7.3C). A second group of “near endemics” (nine species) occurred mostly in Upper Guinea and very scarcely in Lower Guinea, (from the Bight of Biafra to Gabon) (Fig. 7.3D). A third group of “satellites” (three species) showed the opposite pattern, with very few records in Upper Guinea (concentrated at the borders of Côte d’Ivoire with Liberia and Ghana) and a high concentration in Lower Guinea (Fig. 7.3E). Finally, a fourth group of “Guineo-Congolians” (five species) occurred in all three subcentres of endemisms (Upper Guinea, Lower Guinea, and the Congolian Region) (Fig. 7.3B).

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Nine species had an Upper Guinea disjunct distribution (Table 7.1, Fig 7.3A). Most of them have a population at the border between Côte d’Ivoire and Ghana, and a second population at the border between Côte d’Ivoire and Liberia (Cassipourea hiotou) or further to the west (Monocyclanthus vignei, Pierreodendron kerstingii, Schumanniophyton problematicum, Strephonema pseudocola). Both populations, especially the one between Côte d’Ivoire and Ghana, are concentrated close to the coast. Two other species (Acanthus guineensis and Memecylon aylmeri) show a comparable pattern although their populations tend to be more inland. There are two

species with a completely different distribution; Uapaca chevalieri has a disjunct distribution in Liberia and Sierra Leone that seems to be related to montane habitats, and Anisophyllea laurina has two isolated populations in Sierra Leone and Guinea-Bissau. Species distribution range We calculated the distribution range for all those endemic Upper Guinean species with a continuous distribution type and more than five collections in the database (n = 163). Species differed considerably in their distribution range, varying from widespread to very local

Table 7.1 Species or varieties with disjunct distribution patterns. Life form (H=herb, S=shrub, T=tree, WC=woody climber, SH=saprophytic herb), Distribution (UGD= Upper Guinea Disjunct, CD= Continental Disjunct), country of occurrence (S=Senegal, GB=Guinea Bissau, Gu=Guinea, SL= Sierra Leone, L=Liberia, CdI=Côte d’Ivoire, Gh=Ghana, T=Togo) and distribution pattern are indicated. Species

Family

LF

Countries

Distrib. S

Acanthus guineensis Anisophyllea laurina Cassipourea hiotou Memecylon aylmeri Monocyclanthus vignei Pierreodendron kerstingii Schumanniophyton problematicum Strephonema pseudocola Uapaca chevalieri Englerina gabonensis Guaduella macrostachys Illigera vespertilio Lasiodiscus mannii Vernonia titanophylla Anisophyllea meniaudii Calvoa monticola Chytranthus cauliflorus Cola attiensis Dorstenia turbinata Guaduella oblonga Gymnosiphon longistylus Magnistipula zenkeri Manotes macrantha Mapania rhynchocarpa Puelia olyriformis Cassipourea afzelii Combretum bipindense Dracaena ovata Euadenia eminens Hemandradenia chevalieri Marattia odontosora Okoubaka aubrevillei Pyrenacantha glabrescens Tarenna vignei var. subglabra Begonia hirsutula Begonia mildbraedii Hymenocoleus axillaris

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Acanthaceae Anisophylleaceae Rhizophoraceae Melastomataceae Annonaceae Simaroubaceae Rubiaceae Combretaceae Euphorbiaceae Loranthaceae Gramineae Hernandiaceae Rhamnaceae Compositae Rhizophoraceae Melastomataceae Sapindaceae Sterculiaceae Moraceae Gramineae Burmanniaceae Chrysobalanaceae Connaraceae Cyperaceae Gramineae Rhizophoraceae Combretaceae Liliaceae Capparaceae Connaraceae Marattiaceae Santalaceae Icacinaceae Rubiaceae Begoniaceae Begoniaceae Rubiaceae

H T T S T T T T T S H WC S T T H T T S H SH T WC H H S WC S T T H T WC S H H S

UGD UGD UGD UGD UGD UGD UGD UGD UGD CD CD CD CD CD CD CD CD CD CD CD CD CD CD CD CD CD CD CD CD CD CD CD CD CD CD CD CD

Pattern

GB

Gu

SL

L

CdI

Gh

+ +

+ +

+

+

+

+

+

+

+ +

+ +

+ + + + + +

+

+ + + + + +

+ + +

+ +

+

+ +

+ + + +

+ + + + + + + + +

+ + +

+ + + + + + + +

+ +

+ +

+

+

+ + +

+ + +

+ + + +

+ + + + + + + + + + + + +

+ + + + + + + +

+ + + + +

+ + + +

+ + +

+ + + + +

+ +

+ +

T

+

submontane unclear coastal inland coastal unclear coastal inland montane Guinea congolian wide Guinea congolian wide Guinea congolian wide Guinea congolian wide Guinea congolian wide Guinea wide Guinea wide Guinea wide Guinea wide Guinea wide Guinea wide Guinea wide Guinea wide Guinea wide Guinea wide Guinea wide near endemic near endemic near endemic near endemic near endemic near endemic near endemic near endemic near endemic satellite satellite satellite

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A

D

B

E Figure 7.3 Patterns of Upper Guinea disjunct (A) and continental disjunct (B-E) distributions. A) Upper Guinea disjunct (9 species), B) Guineo-Congolian (5 species) with individuals found in all three subcentres of endemism, C) Guinea wide (11 species), D) near endemic (9 species) with the largest population in Upper Guinea, E) satellite (3 species) with the largest population in Lower Guinea. Distribution maps indicate the locations of species collections (dots), the potential forest cover (grey) and country boundaries.

C

(Fig. 7.4): 67% of the species had a widespread distribution (range > 600 km), 14% of the species had a regional distribution (range 300-600 km), 14% a local distribution (range 100-300 km), and 5% had a very local range (< 100 km, Table 7.2). In addition there were 13 other species with a very local range (< 100 km) but with only two to five records in the database. An extreme example of a highly endemic subspecies with a very small range is Impatiens nzoana ssp. nzoana of which all five known botanical collections were found within a range of 7 km on Mount Nimba (Table 7.2). Most species with a continuous distribution had a range between 800 and 1600 km (Fig. 7.5).

Species commonness and rarity Twenty-six species were collected only once or twice and can be considered extremely rare (Table 7.3). Most of these species are found in Liberia (eight species), Côte d’Ivoire (seven species), or both countries (two species). The rare Liberian species occur often in three locations; around Monrovia and Bomi Hills (e.g. Ancistrocladus pachyrrachis), southern Greensville (e.g. Begonia fusicarpa), and in the mountains (e.g. Sericanthe adamii). The rare Ivorian species are frequently found in two locations, near the border with Liberia (e.g. Clerodendrum sassandrense) and around Abidjan (e.g. Argocoffeopsis lemblinii).

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(average r2 = 0.27, range 0.05–0.63). The species occurrence could be explained by a single environmental factor (57 cases), or by a combination of factors (31 cases). Rainfall was the most important environmental factor (significant for 71% of the species) followed by altitude (36%), water holding capacity (23%) and available cations (14%). The combination of significant factors found most frequently were rainfall and altitude (16 cases), and water holding capacity and altitude (11 cases).

Figure 7.4 Types of distribution ranges. A) Widespread (Berlinia tomen-tella), B) regional (Aphanocalyx microphyllus ssp. compactus), C) local (Begonia quadrialata ssp. nimbaensis). Distribution maps indicate the locations of species collections (dots), the potential forest cover (grey), areas above 500 m altitude (dark grey) and country boundaries.

Individual species occurrence in relation to the environment Of the 112 species for which we had enough data to run the logistic regressions, we found significant relationships with the environment for 88 species. The variation explained by the models was moderate to low

80

Distribution type, range and commonness in relation to the environment We compared the environmental variables between species with continuous versus disjunct distributions. We found that species with disjunct populations had a lower number of records in open places than species with continuous distribution and thus tended to be found in relatively undisturbed habitats (Fig. 7.6, t-test, p = 0.001, t = 4.0, nc = 97, nd = 10). We did not find any significant relationship with rainfall (all indicators), altitude, or habitat moisture. For species with a continuous distribution, we compared the environmental conditions among different distribution range categories (widespread, regional, local, and very local), and found significant differences for all rainfall variables (minimum, maximum, mean, range, and 10-percentile) (one-way Anovas, n = 198, p = 0.001). We did not find significant differences between species range categories for altitude, habitat openness and habitat moisture. We performed a multiple regression analysis to compare the relative contribution of all environmental variables on a species’ range. We found that a species’ range was positively correlated with the maximum rainfall and negatively correlated with the minimum rainfall where the species occurs (multiple regression, regression coefficient b maximum for rainfall = 0.4, p < 0.001; b minimum rainfall = – 0.74, p < 0.001; n = 163; r2 = 0.66). Thus species with larger distribution ranges where found in a larger amplitude of rainfall conditions, and were found also in drier places. We found the same pattern among all life forms (i.e. trees, shrubs, herbs, lianas). The relationship between species range and either minimum rainfall or maximum rainfall did not vary among life forms (Anovas, no significant interaction between life form and rainfall covariate) (Fig. 7.7). We found significant correlations between a species’ total number of records and all rainfall variables (Pearson’s r, p < 0.001 in all cases), and also with the percentage of records found in open places (Pearson’s r, p = 0.01). When we tested the relative contribution of the environmental variables, we found comparable results as with species range, namely that a species frequency was positively related to the maximum rainfall and negatively related to minimum rainfall (multiple regression, b maximum rainfall = 0.018, p < 0.001; b minimum rainfall = – 0.027, p < 0.001; n = 163; r2 = 0.37). This relationship was consistent among different life forms.

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Distribution type, range and frequency in relation to species life-history traits Continuous and disjunct distribution types differed in the proportion of species that are shade bearers or light demanding (i.e. non-pioneer and pioneer) (Chi2 analysis, X2 = 6.6, df = 1, p < 0.01, n = 200). There were more shade tolerant species with disjunct populations than one would statistically expect if there was no association between distribution type and species guild. We found an association between life form and distribution type (Chi2 analysis, X2 = 11.1, df = 3, p = 0.01, n = 261). Herbs had relatively more species with disjunct distribution type. We found no association between distribution type and dispersal mechanism. There were no significant relationships between a species’ range and the life-history traits (i.e. life form, guild, and dispersal mechanism). Neither did the abundance of a species vary with its life form or dispersal mechanism. However, light demanding species (i.e. nonpioneer light demanding ones and pioneers) were more abundant than shade-bearers (t-test, p = 0.02, nsb = 70, nld = 66). We did find that a species’ percentage of records at open places was significantly different among dispersal syndromes. Species dispersed by wind had a higher percentage of records in open places compared to species dispersed by animals or with explosive seeds (Fig. 7.8, oneway Anova, p = 0.02).

Discussion

Species distribution: the role of environmental factors Water availability is probably the most important factor explaining the distribution of individual plant species in West Africa. We found that the probability of occurrence of 71% of these endemic forest species in Upper Guinea was related to annual rainfall. These results are in agreement with the findings at smaller scales in the region. For example, in Liberia and Côte d’Ivoire Bongers et al. (1999) found that the occurrence and abundance of 10 of their 12 selected canopy tree species could be explained fairly well by water availability. Also field studies and experiments in Ghana, comparing the effects of rainfall and soil fertility, have found that water availability is the most limiting factor in the distribution of tree species, although soil fertility might be important for some species (Swaine 1996, Veenendaal & Swaine 1998). Not surprisingly, the distribution of forest types and species richness has been explained mainly by changes in rainfall conditions (see Bongers et al. chapter 4, Kouamé et al. chapter 5, Wieringa & Poorter chapter 6). Beyond rainfall, other environmental factors can play a role for particular species. For example, we found that an altitude effect was significant in 36% of the species tested. This probably

Table 7.2 Highly endemic species or varieties with a continuous distribution pattern, and a distribution range smaller than 100 km. The life form and country of occurrence are indicated. The species are ordered based on their maximum distribution range. For abbreviations see Table 7.1. Species

Family

Countries

LF Gu

Uvaria dinklagei Sericanthe adamii Impatiens nzoana ssp. nzoana Clerodendrum sassandrense Pseudocalyx libericus Dichapetalum dictyospermum Albertisia cordifolia Alafia parciflora Sabicea arachnoidea Millettia leonensis Begonia quadrialata ssp. nimbaensis Tapinanthus praetexta Ixora liberiensis Alsodeiopsis chippii Tapura ivorensis Gilbertiodendron robynsianum Cola umbratilis Suregada ivorensis Zanthoxylum psammophilum Strychnos odorata Beilschmiedia caudata

Annonaceae Rubiaceae Balsaminaceae Verbenaceae Acanthaceae Dichapetalaceae Menispermaceae Apocynaceae Rubiaceae Leguminosae-Pap. Begoniaceae Loranthaceae Rubiaceae Icacinaceae Dichapetalaceae Leguminosae-Caes. Sterculiaceae Euphorbiaceae Rutaceae Loganiaceae Lauraceae

WC WC H S WC WC WC WC WC T H S S S T T T T WC WC T

SL

L + + +

Range (km)

CdI

+ +

+ + + +

+

+ + +

+

+ +

+ + +

+ + +

+ + + +

Abundance (#)

Gh 4 6 7 10 11 11 13 22 23 31 46 46 47 60 65 68 68 73 87 89 92

2 2 5 2 2 53 14 3 3 2 17 11 2 6 3 6 10 2 2 4 6

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reflects the fact that altitude represents a complex environmental gradient where different resources (temperature, irradiance, soil fertility, water availability) change simultaneously.

depends on how successful a species is in dispersal and in tolerating open habitats, and a wide range of rainfall conditions. These results suggest the success of a ruderal strategy behind the main patterns of plant distribution in these forests. This is an interesting result because even within the set of endemic or rare forest species of Upper Guinea the most successful species tend to be ruderal, a pattern that is more commonly found in drier ecosystems. Our results are in line with the early observations of Chevalier (1917) and Mildbraed (1922), who pointed out that a very large proportion of the African forest species had wide ranges (Richards 1973). These patterns likely reflect the importance of disturbances in shaping the composition and distribution of species in West Africa. The climatic fluctuations over the past 10 million years, which have periodically led to expansions and contractions of these rainforests (Morley 2000), might have contributed to the selection of ruderal species able to disperse and colonise new areas. Certainly the long and severe deforestation in this region (e.g. Martin 1991) must have reinforced this pattern. As a result of deforestation, the landscape has become drier both in terms of the microclimate of open

General patterns of distribution and driving forces We found that species with continuous spatial distributions had a higher proportion of records in disturbed open habitats than species with disjunct distributions. In fact, species with disjunct distributions were predominantly shade tolerant. Species with larger ranges (i.e. maximum distance between collections) were found in a larger range of rainfall conditions and had a larger drought tolerance indicated by the minimum rainfall where the species has been collected. Also the commonness of a species (i.e. number of records) was positively correlated with the amplitude of rainfall conditions at which it was found and the openness of the habitat. Light demanding species were more abundant than shade tolerants. We also found that species having a higher proportion of records in open places were mainly wind dispersed. These patterns indicate that how continuous, widespread, and abundant a species is,

Table 7.3 Twenty-six extremely rare forest species (or varieties) with only one or two collections in Upper Guinea in the database. The life form, country of occurrence, abundance (# collections), and frequency (# half-degree grid cells in which the species occurs) are given. Byttneria dahomensis occurs only in Benin, so there is no “+” mark in the Countries list. For abbreviations see Table 7.1. Species

Family

Countries

LF Gu

Impatiens nzoana ssp. bennae Begonia fusicarpa Begonia prismatocarpa ssp. petraea Gilbertiodendron obliquum Hibiscus whytei Diaphananthe suborbicularis Malaxis melanotoessa Argocoffeopsis lemblinii Hymenocoleus axillaris Sabicea bracteolata Tarenna vignei var. vignei Byttneria dahomensis Byttneria ivorensis Pseudocalyx libericus Ancistrocladus pachyrrachis Uvaria dinklagei Suregada ivorensis Leucomphalos libericus Millettia leonensis Ixora liberiensis Keetia obovata Sericanthe adamii Zanthoxylum psammophilum Byttneria guineensis Clerodendrum sassandrense Premna grandifolia

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Balsaminaceae Begoniaceae Begoniaceae Leguminosae-Caes. Malvaceae Orchidaceae Orchidaceae Rubiaceae Rubiaceae Rubiaceae Rubiaceae Sterculiaceae Sterculiaceae Acanthaceae Ancistrocladaceae Annonaceae Euphorbiaceae Leguminosae-Pap. Leguminosae-Pap. Rubiaceae Rubiaceae Rubiaceae Rutaceae Sterculiaceae Verbenaceae Verbenaceae

H H H S H H H S S WC S WC WC WC WC WC T WC T S WC WC WC WC S S

SL

L

+ + + + +

+

+ +

+ + + + + + + + + +

+ +

+ + + +

Abundance (#)

Frequency (#)

1 1 1 1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 2 2 2

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 2 2 1 2 2 2 1 2 2

CdI Gh

+ +

+ + +

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gaps within the forest area, as well as in terms of the general climate conditions experienced in the whole region. This is because of the feedbacks of vegetation on both micro and regional climate. In open gaps irradiance and temperatures are higher and relative humidity is lower compared to conditions under the closed forest canopy (e.g. Agyeman et al. 1999b). At a larger scale, tropical forests modify regional climate by recycling a large proportion of the rainfall mainly through the transpiration of forest trees. Deforestation disrupts this feedback, makes climate drier, and beyond a certain threshold could cause a collapse of forest ecosystems and their replacement by savannas (Da Silveira & Sternberg 2001). In fact, hydrological models predict that in the worst scenario of deforestation in West Africa, in which tropical forests are converted into savannas, we could expect a complete collapse of the monsoon system and a significant reduction of regional rainfall to about half of present conditions (Zheng & Eltahir 1997). Certainly we should interpret our results cautiously because they have been derived from the analyses of non randomly selected samples. We can expect herbaria collections to be biased towards places easily approached by roads and therefore closer to relatively disturbed places. One may also argue that the correlation between a species’ distribution range and the range of rainfall conditions is spurious because a species with a large range would cover a larger part of the rainfall gradient. We believe this is not the case because rainfall is the driving factor for occurrence in 71% of the species, and all life forms responded to it the same way. In general, the clarity of the patterns we found is remarkable in view of the great diversity of genus, life forms, and habitat of the analysed species. Rare species and disjunct distributions: relicts from the past? Most very rare species (with one or two collections) and disjunct species occurred near the coast at the border between Côte d’Ivoire and Ghana, at the border between Côte d’Ivoire and Liberia or further to the west. Others were confined to the mountains. These areas coincide with the three postulated Pleistocene forest refuges: Cape Three Points, Cape Palmas, and Mount Nimba (Fig. 7.3, see Wieringa & Poorter chapter 6). The current distributions of these very rare and disjunct species may therefore be relicts from former glacial periods, when the rainforest contracted to a few high rainfall areas, and the forest disappeared in between (Morley 2000). The V-Baoulé, a savanna intrusion in Côte d’Ivoire (Gautier & Spichiger chapter 3) may have separated the forest of Upper Guinea in an eastern and western block. This was even long enough for some speciation to occur. Cola umbratilis in Ghana and Cola buntingii in Côte d’Ivoire, for example, are clear vicariant species (Hall & Swaine 1981). The importance of Cape Palmas (southwest Côte d’Ivoire) and Cape Three Points (southwest Ghana) is underscored by the fact that 65 species are confined to these areas

Figure 7.5 Frequency distribution of species ranges (length in km) of 163 endemic Upper Guinean forest species. Species included have a continuous distribution type and more than five observations in our database.

Figure 7.6 Proportion of records from open places for species with continuous and disjunct distribution types. Mean and standard error are shown.

Figure 7.7 Proportion of records in open places for species with different dispersal mechanisms: explosive (n = 10), animal (n = 11), and wind (n = 8). Mean and standard errors are shown.

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(Aubréville 1949, Guillaumet 1967, updated by Hall & Swaine 1981). Some rare and disjunct species are confined to montane areas. They may have colonised distant areas in the late Pliocene, when the climate was much cooler and montane forest covered a substantial part of the lowlands (Morley 2000). An alternative explanation is that these species are recent immigrants, and have covered large distances by “jump dispersal” (Hall & Swaine 1981, Hawthorne 1996). An example may be Alsophila manniana, a tree-fern from upland forests, that is dispersed by air-borne spores. Importance of these results for conservation in Upper Guinea To set conservation priorities for plant species, one should select conservation areas based on the following ecological criteria: 1) a high concentration of rare and endemic species, 2) a high species richness, 3) uniqueness of the habitat, 4) opportunities to connect different types of habitat and 5) the minimum forest cover needed to preserve the present climatic conditions in the region. Very local species (with ranges smaller than 100 km) and very rare species (one or two records) occurred in five areas: the Liberian coast, around Abidjan, Mount Nimba, southwest Côte d’Ivoire, and southwest Ghana. The latter three sites are also the ones showing a higher species richness (Wieringa & Poorter chapter 6). Given the high sensitivity of these species to water availability and the likelihood that rainfall may decrease in the future, one should allow for the movement of species under changing climatic conditions. We would need corridors allowing species from wet and dry forest types to move towards habitats where wetter conditions are preserved. This could be done by establishing a corridor network of protected areas covering a fair stretch of the climatic gradient. Rivers should play an important role because these are likely to be the moister places left in the landscape if the regional climate becomes drier. Rivers usually act as refuges for species from wet forests (e.g. Meave & Kellman 1994, Puth & Wilson 2001), and as natural corridors allowing species from dry forests to move towards moister conditions. Such an approach could be taken for each of the two largest remaining forest blocks in Upper Guinea, at the borders of Liberia-Côte d’Ivoire and the borders of Côte d’Ivoire-Ghana (Fig. 2.2). For the Liberian-Côte d’Ivoire forest block, we suggest to (see the map in Appendix 2): • create a network of protected areas between the Cestos river in Liberia and the Sassandra river in Côte d’Ivoire. That would connect the very wet evergreen forests at the coast, via the wet evergreen forests of Krahn Bassa, Sapo, Putu Range, and Haute Dodo to the semideciduous forest of Haut-Sassandra (cf. van Rompaey 1993) • connect Grebo forest in Liberia with Taï National

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Figure 7.7 Species ranges in relation to rainfall conditions: a) minimum rainfall and b) the rainfall range. Relationships are shown for four different life forms (herbs, shrubs, lianas and trees). Regression lines, coefficients of determination, and significance levels are given.

Park in Côte d’Ivoire • connect the semi-deciduous forests of Haut Sassandra with the moist evergreen forests of Taï-N’Zo and Niégré using the Sassandra river as a corridor. For the Côte d’Ivoire-Ghana forest block we suggest to: • connect the well-preserved wet evergreen forests in Ghana between Ankasa and Cape Three Points with the semi-deciduous forests in the northeast using the present forest reserves and the Tano and Ankobra rivers • connect the forests of Mabi-Songan-Tamin-Yaya in Côte d’Ivoire along the Bia river with the forests in southwest Ghana. A special conservation effort should be given to montane areas which provide a unique habitat not only for many rare and highly endemic plant species, but also for

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mammals, reptiles, birds and insects (Conservation International 2001). A network of protected montane habitats in Upper Guinea should cover the Fouta Djalon in Guinea, Mount Loma in Sierra Leone, the ZiamaNimba-Mont Péko mountain range between Guinea, Liberia and Côte d’Ivoire, and the Atewa range in Ghana (cf. Conservation International 2001). Quite a few rare and highly endemic species were found near Abidjan. It would be worth preserving Banco Forest, the only large remnant left of the unique forests on tertiary sands (Guillaumet & Adjanohoun 1971). It is important to emphasise that if we only put our conservation efforts in safeguarding a small set of hotspots in a protected area system today, we compromise the conservation of biodiversity on the long term. Small forest patches might not work for these endemic Upper Guinean species even if initially present in an area selected for protection. A highly fragmented landscape, with relatively small forest patches, would be drier both in terms of the local environmental conditions experienced in gaps and forest edges (Bierregaard et al. 2002), as well as by the precipitation falling in the region. This means that species, although initially present in an area, might end up being unable to cope with a progressive decrease in water availability. Seedling establishment is likely to be the bottle-neck for the survival of these plant populations in drier conditions as has been experimentally shown for other rainforests (e.g. Bruna 1999). The importance of preserving West African forests for their feedback on the regional climate was put forward at least several decades ago (Moor 1939). The importance of this is reinforced by our present knowledge of how ecosystems tend to respond to disturbances. In a great variety of ecosystems,

disturbance rates beyond a certain threshold (e.g. removal of biomass greater than a certain surface area) can bring an ecosystem to a state that is very different and often irreversible (Scheffer et al. 2001). In tropical forests a switch to a savanna vegetation could be expected (Da Silveira & Sternberg 2001). Although we can not answer what this biomass removal threshold might be in West Africa, an increase in the protection of these forests seems the most sensible direction based on the combination of insights we presently have on the role of these forests in regional climate, the present regional climate trends, and the way ecosystems might respond to environmental change. The patterns of plant distribution we have discussed in this chapter are a clear indicator of how endemic species are responding already.

Acknowledgements

Jan Wieringa selected the rare and endemic species, and he and Carel Jongkind checked species identifications and distribution maps. Denis Filer created the BRAHMS database software used in this project. Marjo Buitelaar and Hendrikjan Os Breijer entered the herbarium collections into this database. Toon Helmink and Roland van Zoest made valuable GIS analyses and maps. Cajo ter Braak gave advice on statistical analysis. Frans Bongers, William Hawthorne, Sally Horn, Michael Huston and Carel Jongkind gave helpful comments on earlier versions of this manuscript.

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8

F. Bongers, L. Poorter, V. Beligné, W.D. Hawthorne, F. N’. Kouamé, M.P.E. Parren and D. Traoré

H A P T E R

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Forests are important as a habitat for plants and animals, for their role in the maintenance of cycles of rainfall and biogeochemical substances like carbon and nitrogen, and as a source of all kinds of forest products (e.g. timber, fuelwood, bushmeat, medicines, lianas for construction) satisfying the needs of human beings. Forests are commonly used for recreation and religious ceremonies (e.g. sacred forests). In short, tropical forests are able to fulfil a lot of these functions for society. Tropical forests are also well-known to be generally species-rich, although there is a great variation in this aspect. Recent studies have localised the areas in the world where biological diversity is high, but which are at the same time threatened by destruction (Mittermeier et al. 1999, Myers et al. 2000). The rainforests of West Africa rate among the 25 most important “biodiversity hotspots” identified. Because of their isolated position the Upper Guinean forests harbour a large number of endemic animal (Brooks et al. 2001) and plant species (Myers et al. 2000, Wieringa & Poorter chapter 6). At the same time these forests are disappearing rapidly (Chatelain et al. chapter 2). A recent overview of deforestation rates in selected areas all over the world (Achard et al. 2002) shows that the overall rate of forest disappearance is declining. This is an average value, however, and in several West African countries, notably Côte d’Ivoire, the deforestation rates are still extremely high. For an effective policy for conservation and sustainable use, information is needed on several aspects. We need to know where forests are still present, what their conservation status is, and what their actual threats are. We need to know the distribution of rare and endemic species in the region, and the regions in which they are concentrated. We also need to know the main uses of these forests and how they can be managed sustainably. A conservation strategy may be based on a large variety both of species (plants, animals) as well as on habitat diversity (Margules & Pressey 2000). Conservation strategies often focus on rarer species, because these are generally more likely to go extinct without specific management. Unusual habitats often support unusual species, so conservations tend to focus on these as well. For a good conservation strategy, apart from these aspects also information is needed on the maintenance of

Figure 8.1 Satellite image of forest cover of the Classified Forest of Mopri, Côte d’Ivoire, in 2000. The central square represents the area that has been conserved in a conservation project. The other areas are in various stages of plantation development and agriculture (figure provided by Chatelain, Hao, Gautier & Spichiger).

ecosystem processes (maintenance of soil fertility, hydrological functions), the maintenance of plant-animal interactions (pollination, dispersal), and of minimum viable population size of selected species. We need to know how animals and plants react to climate change to be able to predict changes in the vegetation as a result of predicted large scale climatic changes. Finally, we need to know how species react to forest fragmentation that results from logging and agricultural activities. The realisation of conservation goals requires strategies for managing whole landscapes looking at both production and protection areas. Forest Reserves and National Parks are often the cornerstones of national protection strategies, but safeguarding their protection and conditions alone generally is not enough for effective conservation. This is mainly due to the fact that in most countries reserves are biased samples of biodiversity, mostly confined to remote places and to areas that are unsuitable for commercial activities (Margules & Pressey 2000). In Ghana, however, forest reserves cover all forest types (Hawthorne 1996). How can botanical and ecological information be used for the conservation and management of the

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remaining forest resources? In this book we present mainly information on the botanical part of the forests. We describe the forests of the area and classify them into a regional system. We focus on rare and endemic species, their characteristics and their distribution. We address the question of where large numbers of these species are found. Further analyses of the data collected may lead to identification of areas where species are concentrated that deliver products that society needs, like timber, rattans, medicinal plants, and dyes. For a large number of species we have analysed the environmental conditions of the locations where these species are found and we have been able to predict where these environmental conditions are optimal for their occurrence. This could lead to strategies of management directed at use of selected species (e.g. in plantations, homegarden farms and/or in sacred forests).

Deforestation and land use changes The extent of forests is diminishing drastically. At the scale of West Africa we estimate that between 20% and 50% of the forest cover which existed at the turn of the 19th century remains (Chatelain et al. chapter 2). The current area with a forest cover is estimated to be between 463,240 km2 (SOFO 1997) and 415,940 km2 (FAO 2000). These overall estimates are necessarily crude, mostly due to the different methods used in the calculations. For tropical Africa, the variation among researchers in the changes in forest cover between 1980 and 2000 is even larger than the average change observed during that period (Matthews 2001). Fairhead and Leach (1998) have shown that the total forested area at the beginning of the century most probably was over-estimated, which has contributed to an incorrect view of deforestation levels. However, no one can realistically deny that over the last 50 years the extent of forest in our region has declined dramatically. We need an update of information on the forested areas, and on the state of conservation and delimitation of the protected areas. All countries in West Africa show a lack of recent detailed cartographic information, but this is stronger for the countries that have seen war for years such as Liberia and Sierra Leone. Priority zones should be mapped at a higher resolution to be able to follow their development over time in detail. As in almost all countries around the globe, the moist semi-deciduous forest is the most disturbed one (Chatelain et al. chapter 2), mainly because these forests harbour a large number of exploitable species and also because the soils of these areas are of the most productive ones in terms of agriculture. These forests thus need more careful sustainable management. Large areas of forest are still found in southwest Côte d’Ivoire and in southeast Liberia. Smaller, but numerous

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tracts occur in west Ghana. Taï National Park contains a large part of the remaining forest block at the LiberianIvorian border. This famous park is currently actively managed but the effectiveness in the long term is doubtful, as poaching is extreme (Caspary et al. 2001) and cocoa plantations are found in some areas. The political situation over the last past years is not very stable, which is a serious drawback for forest conservation (Richards 2001). Taï National Park needs stronger control of mainly illegal activities, in close cooperation with people living in the area. Alternative land uses need to be stimulated, such as planting trees for wood and for all kind of products in arable fields and in abandoned areas. Planting of Tieghemella heckelii is a good example in this respect (cf. Bonnehin 1998). Currently Liberia, together with several international NGOs is trying to preserve forests in the country, like Sapo National Park. Progress is slow however, not in the least due to the political situation in the country. We suggest that the international community increases political pressure for effective preservation of these last remaining vast areas of forest in the region. Small forests (2 to 100 ha) represent 10 to 30% of the forested area in Côte d’Ivoire. The detailed local analyses of Chatelain et al. (chapter 2) show that the rural areas of Côte d’Ivoire have almost 76% of the forest cover, while 23% is in classified state forests. At present, these small forests are ignored on maps and in the national statistics because they are very fragmented. Updated forest cover maps should have a resolution sufficiently high (30 x 30 m) to allow the resolution of the land-use mosaics and tree plantations, which currently pose a large problem for a realistic estimate of the forest cover updates. Ivorian villagers who invest in forest conservation have no property rights according to the constitution and to the manual of forestry practice. They can do nothing against the continued extraction of large amounts of wood by logging operators who penetrate into community plantations and forests with the authorisation of a logging permit. Most forest policies are focused on forests managed by the State (like Forest Reserves). New policies are needed that focus on small forests in a fragmented landscape despite difficulties for their identification and (long-term) management. To be able to tackle problems that exert pressure on such forests a small-scale local approach is needed, in close concert and cooperation with the local population. For example, the socio-economic infrastructure of village development of more intensive systems of agriculture, and a re-assessment of subsistence cultivation may all need support or help to keep abreast of the changing environment. For a long time, subsistence cultivation was devalued in favour of the system of exportoriented cultivation, with, sometimes high, negative impacts on the food security of farmers. Also, corridors may be established to link existing patches of forest (including “sacred” forests) into larger scale forest networks. A relatively small amount of land area then is

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Box 8.1 Forest corridors and networks for elephants Forest elephants Loxodonta africana cyclotis have declined greatly in the moist forests of Upper Guinea and dwindled down to a total number estimated at not more than c. 3000 individuals in 1998. Elephants influence forests at two main levels: as opportunistic frugivores, by directly effecting the dispersal and regeneration of certain species; and by trampling, debarking and otherwise disturbing the forest. They tend to make gaps and keep forest open and declining elephant populations are followed by altered successional dynamics in some forests (e.g. Laws et al. 1975, Sheil 1996). They are dispersers of a large number of woody forest species, giving rise to concerns that without elephants the plant diversity of Upper Guinea forest plant communities will not be maintained. Although several studies (e.g Chapman et al. 1992, Lieberman et al. 1987) have pinpointed plant species that are absolutely elephantdependent for dispersal/germination, two recent studies cannot find hard evidence for that (Hawthorne & Parren 2000, Chifu Nchanji & Plumptre 2003). Nowadays, these elephants live mostly in scattered forest patches in a matrix of cultivated land, except for those in extensive forests in eastern Liberia (Parren & de Graaf, 1995; Appleton, 1997). The elephants of the Côte d'IvoireGhana border area, the total population of which numbered 685-855 in the late 1980s, are contained by the savannas of the Dahomey Gap to the east and by the so-called V-Baoulé, in Côte d'Ivoire to the west. Movement of elephants between forest fragments, often not exceeding 10,000 ha, in the Côte d’Ivoire-Ghana border area would require forest corridors between nearby fragments. Corridors are linear vegetations

needed to be able to improve the existing forest “infrastructure” and thus to improve forest quality and forest functions. For the border area between Côte d’Ivoire and Ghana comparable forest networks (with the elephant as a flagship species, see Box 8.1) have been proposed recently (Parren et al. 2002). Such measures then should lead to a lower pressure on the remaining old-growth forests in the area: planted corridors can be used by the local population for some of their needs (e.g firewood, construction poles). This relief of pressure on old-growth forest may have positive effects on animal and plant populations. The ideal of actually regaining forest area in corridors may seem a long way off, but in the shorter term, and less contentiously, we suggest that the State first strives to establish a widespread policy of conservation based also on community management, as is widely promoted in Ghana, with varying degrees of success (see also Fairhead & Leach 1998). Villagers need forest to fulfil many of their needs (e.g. wood, medicines, religion). Only in a long-term conservation scheme in which advantages for the local population are clear for the short and long term, villagers will (co-) protect forests. And, before even this stage can

connecting at least two isolated habitat fragments that were once connected (Saunders & Hobbs 1991). “Shelterbelts”, strips of forests often not more than 1.5 km wide and up to 20 km long (Foggie 1949), established in the mid-1930s in Ghana, provide examples of potential corridor design. Corridors should be designed to attract forest elephants into them, and important characteristics to take into consideration are the presence of food plants, availability of water, and the size and structure of the corridors. Forest elephants feed opportunistically on certain fleshy fruits when these are available, and rely upon less nutritious foods during periods of fruit scarcity. Favourite forest tree species are Parinari excelsa, Balanites wilsoniana, Duboscia viridiflora, Panda oleosa, Sacoglottis gabonensis and Tieghemella heckelii (Sachtler 1968, Martin 1991, Hawthorne & Parren 2000). In the dry season the water provided by the fruits may be insufficient and elephants come out of the forest in search of water. Artificial waterholes could be created in forest elephant reserves to ensure they stay within the reserve boundaries, as has already been successfully done inside FC Bossematié (Waitkuwait 1992). Corridors on riverbanks, such as the Bia river corridor, would have the advantage of providing readily available drinking water. Corridors along streams and rivers would probably be the most useful and successful as such landscape features have a disproportionately large influence on landscape functions (e.g. population, community, and ecosystem processes) beyond what their relative area on the landscape would suggest (cf. Puth & Wilson, 2001).

be expected to make progress, villagers who conserve the forests need to get the necessary rights to accomplish this conservation action. We therefore recommend managers and politicians in the immediate future focusing on these short-term goals and, when the benefits are obvious to all, the medium and longer term goals outlined above might also be realisable. Fire as a management tool The local population can have a strong direct impact on the extent of forest (Gautier & Spichiger chapter 3) especially in the drier areas. There is a general agreement that savannas of the transition zone are not in equilibrium with climate and are a direct consequence of annual fires, which are, however, more likely in periodically drier periods. The climatic climax would be dry semi-deciduous forest and further north (from c. 9°N) dry forests and woodlands. Most Guinean savannas are considered to be of secondary origin. Protection of fire, cattle ranching with exclosures and shifting cultivation with longer fallow periods would lead to forest expansion, and give opportunities for area expansion to the threatened dry forest type.

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The prevailing environmental conditions may be favourable for afforestation, even when the effect of fire is taken into account (Aubréville 1966). This has been demonstrated by comparing two sets of aerial photographs separated by 25 years in Lamto reserve, Côte d’Ivoire. Despite the occurrence of annual fires, the forest surface is increasing both by direct progression of existing forests and by the appearance of forest islets within the densely wooded types of savannas found on hilltops (Gautier 1989). Hilltops may need continued protection as safe sites for establishment of forest species, and sources of recolonisation of the forest. However, the hilltops on the Accra plains were reserved in the 1930s and still exist. Meanwhile fire continues to rage annually across the lowlands and nothing ever redevelops there. Swaine et al. (1992) demonstrated that forest that recovered in fire protected areas included dry semi-deciduous forest species that had no local seed source, presumably because the majority of them are well dispersed by bats and wind. In Côte d’Ivoire the timing and frequency of fire as a management tool has shown promising results. Gautier and Spichiger (chapter 3) describe experiments that show that annual early burning (beginning of the dry season) has promoted a shrub-dominated savanna, while annual retarded burning (mid to late dry season) has promoted a savanna with a very low shrub cover because of a greater fire intensity (drier and maybe also more fuel) and the fact that it destroys tree leaves and shoots that establish long before the rainy season. Meanwhile an integral protection from fire has led to a genuine forest structure restoration, even in the vicinity of the southern dry outlier forests in the Dahomey gap (Swaine et al. 1992). These results imply that fire can be used as a management tool to create specific desired vegetation types, and that fire protection in particular is crucial to forest conservation in the semideciduous forests. In northeast Ghana, another fire protection experiment has led to a clear decrease of grass cover and a clear increase in tree number and stature, but in contrast to the above, there has been no establishment of GuineoCongolian forest species. This is explained by the much more northern position of the plot, north of the forestsavanna transition zone, in a region where dense forest species are very rare and thus forest seeds unavailable. This indicates that successful restoration of Guineao-Congolian forest is only possible if seed sources are available, although many pioneers can apparently arrive from many kilometres or tens of kilometres (Swaine et al. 1992). The environmental matrix is important: gallery forests may be useful in some savanna areas as corridors for Guineo-Congolian species. Fires can also occasionally sweep under the canopy of the drier types of rainforest (for Ghana, see Swaine 1992). These fires are generally low in intensity and are essentially fuelled by litter (ground fires, Phillips 1974). They extend to the canopy only during exceptionally dry conditions (Swaine 1992). Like logged forests, burnt forests are more prone to renewed burning (Hawthorne 1991). This can be

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dangerous and large fires therefore have to be strictly under control: prevention is better than healing. Impact of shifting cultivation on forest extension There has been considerable debate on the impact of shifting cultivation on the distribution of forest and savanna. It was generally assumed that shifting cultivation would lead to an extension of savannas by destroying woody vegetation (Aubréville 1966). In South America slash and burn agriculture, fire and deforestation have lead to savannisation. In some places like the western extension of the V-Baoulé (see Kouamé et al. chapter 5), there is general agreement that the Pennisetum purpureum savannas are derived from cultivation of forest areas (Adjanohoun 1964). Field studies in Côte d’Ivoire and Togo have even revealed that shifting cultivation at the forest edge promotes afforestation (Spichiger & Blanc-Pamard 1973); field preparation reduces herbaceous domination in the fallow, and thus limits fuel availability. Fairhead & Leach (1995; 1998) have demonstrated that (especially in Guinea) many human activities promote afforestation (farming system, deliberate planting of forest species in fallows, pasture, fire-breaks). Based on aerial photographs, they show that in some areas this has lead to a 40% increase in forest cover during the last 50 years. Although this might be the case for some areas, the results of Achard et al. (2002), however, show that overall deforestation is far larger than afforestation. Increase in population pressure as well as transforming farming systems into cash crop production has completely altered the intensity of human impact on vegetation during the last decades. The duration of the fallow in the farming system is of paramount importance. If the fallow period is reduced from c. 10 years to c. 5 years, as is currently happening in many places in the forest-savanna transition zone, then afforestation processes can no longer take place. We therefore suggest to aim at a fallow period of at least 10 years. We have to realise, however, that such young secondary forests will not help much in long-term biodiversity conservation, and as such it will never be a substitute for old-growth forest. It does help, however, in terms of soil fertility maintenance (increasing agricultural production when it is next farmed), or it provides fuelwood for burning (lowering the pressure on old-growth forests), and it may increase common wild animals that can be used as bushmeat, as an alternative for the rarer wildlife (again lowering pressure on old-growth forests).

Conservation: hotspots and other areas As part of systematic conservation planning a general compilation of data on the biodiversity of a planning

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region is needed, and conservation goals have to be identified for that region. Based on that, existing conservation areas have to be reviewed and additional conservation areas might be selected with the various stakeholders. Sometimes, more detailed analyses of biodiversity data in the planning region are needed for that. The data and results presented in this book should support this part of the systematic conservation planning process. Biodiversity hotspots: consequences for natural reserves Upper Guinea is one of the 25 most important hotspots in the world (areas where biodiversity is very high and threats in terms of deforestation are large). The Upper Guinean forests are estimated to contain 2800 forest plant species of which 650 are endemic, and c. 400 are considered to be rare (chapter 11). These forests also harbour 1320 vertebrate species of which 270 are endemic (Myers et al. 2000). Within Upper Guinea and based on a selected list of endemic and rare plant species for Upper Guinean forests, Wieringa and Poorter (chapter 6) distinguish three areas of high diversity. A belt of rare and endemic species is found about 50-100 km inland, from Sierra Leone, through Liberia to southwest Côte d’Ivoire. The climax of this belt is in Liberia. Additional high diversity areas are found in areas around Abidjan in Côte d’Ivoire and Ankasa in Ghana. A second belt is found in the mountain areas of Mt Nimba/Mt Ziama in Guinea, and around Man in Côte d’Ivoire, and the Atewa Range in Ghana. These areas need protection and attention. Clearly, the mountain ranges have many endemic and rare plants. Within these area the mist-bound summits with all their epiphytes are the more diverse parts, and include many species that are uncommon throughout the more extensive lowland forests. The steeper hills are important because of the variety in special habitats and the fact that they are relatively untouched. Hill sanctuaries, as established in Ghana (Hawthorne & Abu Juam 1995), may provide a good framework for planning general protective management throughout the region, as issues of soil erosion, water supply conservation, migratory bird refuges are at a peak here, and yet the sites are generally the least suitable for logging, and not ideal for farming either. Many of the higher hill forests, such as those around Man in Côte d’Ivoire and Tano Ofin and Atewa in Ghana are also rich in rare species that flourish in the hilltop mists; some of these forests have escaped relatively undamaged until the last few years, but are currently succumbing rapidly to destruction, and deserve a special conservation review. The location of the areas with high plant diversity is strongly related to both water availability and, it is assumed, to the presence of ancient forest refugia. Diversity clearly increases with rainfall and decreases with soil fertility and with distance to presumed ancient forest refuge areas (Wieringa & Poorter chapter 6). In such refuge areas plant species, especially the drought-intolerant ones, are thought to have survived the glacial periods in

the past. Because of their isolation during the glacial periods speciation may have occurred. After the glaciation period plant species supposedly dispersed from these refuge areas, leading to higher numbers of less dispersible species nearby. Supposed refuge areas thus need special conservation management inasmuch as any area that is rich in more localised species demands it. We have to realise that many areas in the Upper Guinea forest block are undersampled with regard to herbarium collections (Figure 6.4 and 6.5). Sampling intensity is particularly low in southeast Liberia. As many of these areas are high rainfall areas we expect that their plant diversity is very high. We recommend herbarium collection expeditions to these areas. Distribution patterns of rare and endemic species The rare and/or endemic species of Upper Guinean forests show three characteristic distribution patterns (Holmgren et al. chapter 7). From the 270 species analysed 86% showed a continuous distribution (the species is found over the whole area), 10% showed a continental disjunct distribution (the species is found somewhere in Upper Guinea and also in central Africa) and 3% showed an Upper Guinean disjunct distribution (the species is found in two separated areas within Upper Guinea). Most species had a wide distribution, and 21 species had a distribution smaller than 100 km (Table 7.2). These latter species need extra conservation attention, as species with a small range are generally the most vulnerable to extinction. In total 26 species in this sample (which we emphasise is not a comprehensive set of all very rare species, many of which are still in the process of being described) are considered very rare as they have been collected only once or twice. Eight of these are found in Liberia, especially around Monrovia and Bomi Hills, in southern Greensville and in the mountains. The rare Ivorian species are concentrated near the border with Liberia and around Abidjan. The occurrence of 88 out of 112 species was correlated with various environmental factors, of which rainfall was the most important (in 77% of the species rainfall had a significant effect), followed by altitude (in 36% of the species a significant effect). Species with disjunct populations tended to be found in undisturbed areas, and were often shade tolerant species. Species with continuous spatial distributions were often found in open disturbed habitats. How continuous, widespread and abundant a species is, depends on how successful a species is in dispersal and in tolerating open habitats, and a wide range of rainfall conditions. Thus, even within the set of endemic or rare forest species of Upper Guinea the most successful species tend to be pioneer type species. Disturbances have obviously played an important role in shaping the composition and distribution of forest species in this area. As for the species with disjunct distributions, specialising on very specific environmental conditions (in

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most cases wet and undisturbed), most need special conservation effort. Three out of the five areas where many rare species and very local species occur also show very high species richness, notably Mount Nimba, southwest Côte d’Ivoire and southwest Ghana (Wieringa & Poorter chapter 6). Areas where environmental gradients are steep (i.e. a long gradient over a relatively short distance) have many environments in a relatively small area. As rainfall is the most important environmental factor determining species diversity, and also determines distribution of most species, areas with a steep rainfall gradient are extra interesting from the point of view of conservation. Climate change thus may have a profound impact on the distribution of species (Holmgren et al. chapter 7). A series of reserves along the gradient, connected by corridors would be very helpful. Due to the gradual changes species can migrate easily over the whole area in periods when this is needed (e.g. during periods of climate change). Two such areas have been identified. One is the gradient in southeast Liberia and southwest Côte d’Ivoire and the other is the area in southwest Ghana (see Figures 1.3 and 4.3). In the Liberian-Côte d’Ivoire zone several areas can be connected, notably between the Cestos and the Sassandra rivers, connecting the wet coastal forests with the semi-deciduous forests, between Grebo and Taï forests, and between Haut Sassandra and Taï-N’Zo and Niégré. In the Côte d’IvoireGhana zone the forests of Ankasa and Cape Three Points would ideally be connected with the northeast using the Tano and Ankobra rivers, and the Mabi-Songan-TaminYaya forests might be connected with the forests in southwest Ghana. Implementation of such corridors would need huge efforts, however. Some of the driest forests, especially those in Ghana at the eastern limit of our atlas, are species poor, but of extreme high conservation concern, as these forests include the last stands of several extremely rare species, and the forests are of extremely small, and diminishing extent.

Time and again, however, calls have been made to secure the highly endangered Talbotiella gentii stands in Ghana, but they are still not fully secured (e.g. Swaine and Hall 1981; Hawthorne 2001). Because these may have slipped through our (selected species) herbarium analysis, efforts should be made to seek other such pockets of dry forest, possibly with endemics as yet undiscovered, elsewhere on the periphery of our region. The mountain areas contain many rare and endemic species and need special conservation effort. The Fouta Djalon in Guinea, Mount Loma in Sierra Leone, the Putu Range in Liberia, the Ziama-Nimba-Mont Péko mountain range between Liberia, Guinea and Côte d’Ivoire, and the Atewa range in Ghana are crucial in this respect (Wieringa & Poorter chapter 6, Holmgren et al. chapter 7, see also Conservation International 2001). Also Banco forest, near Abidjan, ranks high in number of rare and/or endemic species. The scattered dry forests of southern Ghana (Southern Marginal and Southeast outlier types) tend to slip through the analytical net with these sorts of surveys, because they are species poor, but the vanishingly small and highly vulnerable communities include a high proportion of globally rare endemic plants, like the very rare Talbotiella gentii and Hunteria ghanensis trees - and so are amongst the highest priorities for conservation effort in our region (Hawthorne and Abu Juam 1995). The Ghanaian Southern Dry forests are again the target of protective measures by the Ghana Forest Service, but it remains to be seen how secure such small pockets can be made without a major ex situ conservation initiative. We would like to stress the fact that, even though the options are limited for small communities like the Ghanaian Southern dry forests, large stretches of forests are always desirable because small patches are much more vulnerable, especially under current levels of disturbance and climate change.

Table 8.1 Distribution of forest condition, and plant diversity, for areas under protection (national forest parks, forest reserves, classified forests) for Côte d’Ivoire and Ghana. The forest condition is classified into six classes: class 1 (excellent condition), class 2 (good), class 3 (slightly degraded), class 4 (mostly degraded), class 5 (very poor), and class 6 (no significant forest left). Data for Ghana from Hawthorne and Abu Juam (1995), data for Côte d’Ivoire from H. Dao and C. Chatelain, amended by V. Beligné. The plant diversity (S50 values, the number of plant species for 50 random collection) increases from class 1 to class 7 (class 1: 0- 1 5 species; class 2: 15-30 species; class 3: 30-35; class 4: 35-37.5; class 5: 37.5-40; class 6: 40-42.5; class 7: 42.5-45.0). The forest types are DS = Dry Semi-deciduous; MS = Moist Semi-deciduous; ME = Moist Evergreen, WE = Wet Evergreen. Values are percentages of total area under protection.

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A. Forest condition Ghana Côte d’Ivoire

1 (good) 1.7 25.6 (inc PN)

2 15.1 11.3

3 37.2 35.6

4 19.4 21.2

5 16.2 3.6

6 (bad) 10.2 2.6

B. Plant Diversity Ghana Côte d’Ivoire

1 (low) 0.2 0

2 13.7 10.4

3 47.5 27.6

4 23.6 13.5

5 11.7 43.2

6 3.4 4.6

7 (high) 0 0.6

C. Forest type Ghana Côte d’Ivoire

DS 12.7 4.4

MS 35.3 12

ME 31.0 9.4

WE1 19.9 13.4

WE2 0.8 21.8

WE3 0.3 39.1

HW

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Complementarity in selection of areas for conservation In selecting areas for biodiversity conservation the principle of complementarity is very important, apart from high numbers of rare and endemic species, high species richness, uniqueness of the habitat, minimum forest cover and possibilities for connections in networks. Complementarity analysis seeks to include as many species as possible within a network of protected areas, thus sites are selected that add as many species as possible to the protected pool. For Côte d’Ivoire Kouamé et al. (chapter 5) show that many species are very forest specific (out of the 2126 species in the thirteen forests studied, 624 species were found only in a single one of these forests, see also Figure 5.2) and thus it would seem that a large number of specific forests need to be protected to be able to conserve a large number of species. However, it is acknowledged by all proponents of complementarity analysis that it is much diminished in relevance where 100% complete species lists are not available – a fact that is sadly the norm for forests in our area. Where almost complete checklists for large forest reserves have been compiled from special surveys, it is seen that a significant proportion of the flora of our entire area occurs in limited regions. Between about 24 and 33% of the Upper Guinea forest flora is found in Bia South forest in Ghana, for instance, an “average” type of forest (Hawthorne et al. 1998). In Ankasa and Draw Forest Block alone about 1200 species are found, about half the forest flora for the entire region. For these reasons, complementarity probably has less to offer than prioritisation by rarity, but we can still adhere to general principles – to try and protect as wide a variety of forest types as possible. This is a principle that was extended by Hawthorne and Abu Juam (1995) in a bid to protect the “hidden” infraspecific biodiversity in terms of genetic diversity within overexploited and widespread species (see also Box 8.2). With respect to our hotspots of plant diversity we suggest that as much as possible of the hotspot areas will be protected, but also that the number of total protected species will be considered. Kouamé et al. (chapter 5) distinguish four forest types with local endemic species. Consequently, forest management and protection in Côte d’Ivoire must be oriented to have at least a forest reserve/national park in each of these forest types. In three of these, forest reserves have been established already: Taï NP for western evergreen forest, Banco NP for coastal and Marahoué NP for semi-deciduous forests. The creation of biological reserves in the classified forests of Yaya-MabiSongan-Tamin in eastern evergreen forest is the first step towards protection of this type of forest. For this chapter, we have calculated the plant diversity of the forests in Ghana and in Côte-d’Ivoire in several Upper Guinea-wide classes (Table 8.1). In Ghana the majority of the forest reserves are found in areas with intermediate species diversity. Only 3.4% of the reserve areas are found in high diversity areas (classes 6 and 7). The main reason for this is that high diversity areas and

very wet areas form a small proportion in Ghana’s forest, and these are the areas with the highest diversity (note here that diversity values are determined on a regional scale, see chapter 6). As almost 100% of all larger extant forest blocks in Ghana are within forest reserves, a bigger issue is how precisely to manage it, not what to gazette. For protected forests in Côte d’Ivoire diversity is slightly higher, especially class 5 is better represented. Overall the variety in diversity is high (see also the maps in chapter 6 and Appendix 4). Because extremely wet sites only occur in Liberia and Sierra Leone, these hyperwet areas need special protection. The Ghanaian Southern Dry forests, forest that has already been emphasised as a conservation priority, are currently the focus of protective measures by the Ghana Forest Service.

Forest reserves and protected areas A regional classification of forests: consequences for conservation For the Upper Guinea area several forest classifications have been made in the past (Table 4.5). Most of these are at the scale of one country. In Chapter 4 Bongers et al. have produced a regional classification based on over 200 sites in the Upper Guinea region. This is a very useful addition to the already existing classifications. In general the classes are comparable but in some cases differences exist. This is for a large part due to the longer environmental gradient existing in the whole area, compared to the one in a single country. For instance, the wet forests in Ghana are not “very wet” when seen in a regional perspective. Also, our classification (and thus also the resulting map (see Appendix 2) is based on a selection of species only, which makes it easy for decision makers to put “their” forest in an international context. In Chapter 4 Bongers et al. have clearly shown where the new classification is different from the older ones. It is clear that some forests, like the hyperwet forests of Liberia and Sierra Leone do not occur in the other countries. As such forests do not occur in Ghana and Côte d’Ivoire, Liberia and Sierra Leone have a special regional responsibility with regard to specific protection of hyperwet forests. Other forest types, such as the southern marginal outlier forests occur only in Ghana. These forests are extremely dry and have a high proportion of endemic species (and also of endemic individual plants) and thus a high conservation value, despite low species diversity (Hawthorne 1996). Conservation of such forest thus is of a wider importance than for the particular countries alone. The same applies to the upland mountain forests that occur sporadically in Ghana, but mainly in the triangle border of Côte d’Ivoire, Liberia and Guinea.

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Figure 8.2 A large Ceiba pentandra tree is cut in a forest concession at N’Zo (Côte d’Ivoire).

In this book eight broad forest types are characterised. That is not an overview of all the different forest types in the region, only a characterisation of the forest types and forest sites used in this book, at the level of Upper Guinea. The special forest types in Ghana have been mentioned already. Also, swamp forests and mangrove forests, and various kinds of riverine forest and sacred groves in the savanna area are not taken into account. With respect to the mountain forests, all are put in one category here. We have to realise, however, that at a more local and more detailed scale many more differences can and will be found. For a proper conservation of all these forest types, in-depth analyses are necessary. These can better be done at a more local level. The studies in Ghana (for instance Hall & Swaine 1981, Hawthorne & Abu Juam 1995) are starting points for those studies. Also for Côte d’Ivoire such studies are being undertaken, and for Liberia new studies are being performed currently (Conservation International, Flora & Fauna International). Status of the areas under protection In Upper Guinea various kinds of protected areas can be distinguished: national forest parks, forest reserves and classified forests. For the countries of Liberia, Côte d’Ivoire and Ghana we have mapped these forests (Appendix 3). In spite of the fact that some natural areas are protected by law, this legal protection is no guarantee for factual effective protection. The conditions of these forests are quite variable, due to legal and illegal logging (Figure 8.2) for timber but also to clear land for small scale as well as large scale agriculture. In Côte d’Ivoire, for instance, only 11 areas are formally protected as “national parks” or “natural reserves”, with in total 1,874,800 ha: Azagny, Banco, Comoé, Haut-Bandama, Iles Ehotilé, Marahoué, Mt Péko, Mt Sangbé, the complex Taï & N’Zo and Mt Nimba (Bakarr et al. 2001). Only Comoé and Azagny National Parks were considered to have a reasonable level of administrative and management support, enough to

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provide real protection for the parks. All others are considered to have shortage of effective protection. For Taï National Park & N’Zo Reserve effective protection is doubtful, despite numerous international agencies maintaining research projects in the area. Taï is very important as it probably contains the only spots of primary forest for the country, and still contains many of the species that are disappearing from other areas. But currently also Taï (and especially the formerly called buffer zone) is threatened by habitat degradation, poaching, logging, farming, illegal gold mining, and other human activities. For Ghana, the forest condition was determined by Hawthorne and Abu Juam (1995) into six categories: class 1 (excellent condition), class 2 (good), class 3 (slightly degraded), 4 (mostly degraded), class 5 (very poor), and class 6 (no significant forest left). From the total areas of forest reserves 26.4% was in bad condition (their classes 5 and 6), 16.8% in good condition (their classes 1 and 2) and the rest was intermediate. For Côte d’Ivoire this was done by Hy Dao, Chatelain and amended by Beligné. For the classified forests of the forest zone 6.2% was in bad condition, 36.9% in good condition and the rest in intermediate (Table 8.1). The high percentage in good condition is mainly due to the large area of Taï National Park. For Liberia comparable information is scarce and scattered, and we are not able to give a comparable and reliable overview. For some of the forests information is collected by Conservation International and Flora & Fauna International. D. Byler (personal communication) and M.P.E. Parren (personal communication) considered in very good shape (class 1) Sapo National Park, in good shape (class 2) Grebo National Forest (but degrading M.P.E. Parren), and parts of Krahn Bassa National Forest, and as slightly degraded (class 3) Gio and Gbi National Forest (but degrading - M.P.E. Parren), the largest part of Krahn Bassa, and Lorma National Forest. The Lamco concession area was in degraded condition (class 4). Chatelain et al. (chapter 2) show that in the large majority of classified forests in Côte d’Ivoire 40-50% of these forests have been replaced by planted trees and crops. The numerous classified forests in the region of Abidjan, for instance, are not well managed and protected. Their importance for wood production and as a biological reservoir is doubtful, because in many cases they are forest only in name (77% of their area is not forested). In Côte d’Ivoire protection effectively maintains reasonably good quality forests only in national parks (see Table 8.1). To avoid this and to restrict agriculture to areas outside the forests, farmers need to be encouraged to invest in their plantations (e.g. rejuvenation) and to strictly forbid agricultural practice in classified forests. The existing agricultural lands in the surroundings could be more effectively used, and many fallow lands are practically unused. Additionally, conflicts between modern and traditional land rights as well as the principle of land ownership acquired by clearing, are both obstacles in a

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Box 8.2 Integrating forest management with academic analysis of endemism and rarity There are several problems with linking global patterns of rarity to management practice, including the following three examples:

except presence-absence in small areas is highly expensive. Complementarity analysis for instance requires more or less complete species lists for forests to be prioritised.

1. There is a continuum of scale of analysis of pattern and of management implementation of rarity (Hawthorne 1996). Endemism and presence/abundance issues are all dependent on scale. Patterns of concentration of more or less (globally) rare species are sharply demarcated at a local scale, but vary subtly across any landscape, from swamps to ridges and from logged to unlogged forest.

For these and other related reasons, and in order to integrate global trends with local management practice and Environmental Impact Assessments (EIA), Hawthorne and Abu Juam (1995), developed and implemented for Ghana a species (“Star”) and associated forest (“Genetic Heat Index system”) ranking system. This is a fairly crude but equally simple, objective and repeatable method, and has enabled GEF (Global Environment Facility) funded projects to target priority areas (GSBAs) for special management. More detailed examination of these Globally Significant Biodiversity Areas (GSBA) has yielded numerous new species, exemplifying the predictive power of the system. Meanwhile Star and GHI rankings have become part of the common parlance of forestry in Ghana at all scales (tree selection in logged forest; forest EIA – e.g. for the Subri industrial plantations company, seeking low priority parts of Subri FR for Gmelina plantation), and conservation is correspondingly a routine issue in Ghanaian foresters’ minds.

2. There are considerable difficulties with obtaining precise information on all species, even if one can see they are globally rare. Although a large subset of species has been reviewed here, they are not a complete set of all high priority species, and even this work has taken many hundreds of person-hours to compile. 3. For many proposed methods of linking the global to the local, obtaining adequate information about the local community is likely to be a problem; forest inventory of all

lasting land management. Some of the classified forests in Côte d’Ivoire, however, do have an important role to play with respect to biodiversity: many forest species are confined to one or a few forests only (Kouamé et al. chapter 5). The same accounts for the forest reserves in Ghana (Hawthorne 1996) and probably also in Liberia. In Ghana forest reserves are thought to reasonably well represent the variety of forests available a century ago (Hawthorne 1996), but special conservation measures (“GSBAs”, Globally Significant Biodiversity Areas) have recently been instigated by the Ghana Ministry of Lands and Forestry within that portion of the reserved forest with the higher proportion of rare plants, recognising their priority status. Work is underway to demarcate and manage sensitively a modest network of these specially protected GSBAs within the much more extensive and long standing network of forest reserves. The GSBA priorities were selected (using methods described by Hawthorne and Abu Juam (1995) for their high biodiversity conservation priority), with a view to better and sensitive conservation of high priority species, but within a broad concept of ongoing forest management and use. In Liberia effective measures for forest protection are difficult due to the unfavourable political situation over the last decade, but also, construction of national parks with effective measures are currently being planned (e.g. for the Sapo National Park area, D. Byler, J. Suter and E. Waitkuwait personal communications). Much additional work needs to be done

in this field before conservation and protection efforts will be effective. Table 8.1 also shows the percentages of area of protected forests in different forest types. In Ghana the majority of protected forests is in moist semi-deciduous and in moist evergreen forests, with a considerable part also in wet evergreen. In Côte d’Ivoire the wet evergreen forests are well represented with c. 40% in the third type. Forest corridors and animal conservation Many plant species depend in their life history on animals, be it for pollination, for dispersal of their seeds or otherwise. For effective conservation in the long term not only conservation of plants is needed but also of the animals that are connected to them. Hunting is widespread in the region and leads to severe declines in animal populations (e.g. Caspary et al. 2001). Many large animals have large home ranges and as a result of fragmentation of forest, the resulting forest patches may be too small to sustain a viable animal population. Thus for effective protection of plant species and their related animals sizeable forest tracts are needed. An alternative is a combination of smaller patches connected to each other with corridors, forming ecological networks. Networks that link forest reserves and classified forests are particularly interesting in this respect. Such corridors are needed as linkages for animals and plants to cross areas to maintain a larger population that might be more viable. Parren et al. (2002), for example, plea for a network

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Figure 8.3 Distribution of stock of commercial timber out of the selected species of Chapter 4 (Table 4.1). The number of individuals > 30 cm dbh are shown, together with their interpolation for the whole area. Notice that this is a potential map based on earlier inventory data, and not the actual situation.

Figure 8.4 Tree bark is sold on the market of Man, Côte d’Ivoire.

directed to forest elephants. They focus on the border areas between Ghana and Côte d’Ivoire (Box 8.1). Also, some plant species have special value for animals as keystone species. Keystone species provide food for animals when overall food availability is low. Fig trees are known for their overall importance as food source for a large variety of animal species. Such individual species thus need special attention. On the other hand, animals can be extremely important for plants as well, in terms of pollination and seed dispersal for example. The forest elephant is the best-known example (see Box 8.1). Areas that have high value in terms of plant diversity are not necessarily important for their animals. Logically those areas that have biodiversity values for a large number of plant and animal groups will have a larger chance of

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being effectively protected. To be able to characterise different areas in Upper Guinea with respect to overall biodiversity, in 1999 Conservation International brought together a large number of specialists. Together they produced a biodiversity hotspot map for terrestrial ecosystems for the whole of Upper Guinea, based on priority areas for mammals, birds, reptiles and amphibians, insects, and plants. For plants the extremely high conservation priority areas were: Liberia Hygrophilous Evergreen Forest, Taï-Haute Dodo area, Mount Nimba, Gola, Ghana Wet Evergreen Forest. Several of these areas are also hotspots for animals (see maps in Conservation International 2001). The Taï-Haute Dodo-southeastern Liberia area is important for plants, mammals, birds, reptiles and amphibians. The same accounts for the Gola area in western Liberia, and for the Mount Nimba region. Banco National Park is a hotspot for reptiles and amphibians and insects, but less so for mammals and birds. The Guinea highlands of Sierra Leone and Guinea are of high importance for mammals, the Loma mountains in Sierra Leone stand out in terms of insects. Taking all five groups into account, the overall (regional) conservation priority areas for Upper Guinean forests are found in (from west to east) the Fouta Djalon, the Gola-Lofa-Mano complex, Mount Nimba Range, Taï-Grebo-Putu Range, the Cestos-Upper Krahn Bassa area, Mount Peko, Abi Lagoon-Cape Three Points area, Ankasa Jema, KrokosuaSui and finally the Ghana-Togo highlands.

Forest resources and use The forests of West Africa have been used extensively for their resources. Assuming that plant composition is indicative for the whole forest ecosystem we can use the

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forest classification as indicative for the whole system. We thereby assume that a broad cover of vegetation types also implies a broad cover of other forest elements (e.g. animal communities). This means that our overall ecological stratification is based on forest types. By linking for instance forest use to forest types we can predict specific forest uses over a large area. For many areas the main resources were timber. For the forests that have been inventoried for a selection of species (see Bongers et al. chapter 4) we have calculated the total number of commercial trees larger than 30 cm diameter per square km (note that the data used are mostly pre-logging inventory data and thus not indicating the actual but rather the potential timber stock). This stock of commercial species is an indication of the importance of these forests as a timber resource. Figure 8.3 shows that the concentration of timber species was high in Liberia, southwest Côte d’Ivoire (specially Haute-Dodo and HautSassandra) and southwest Ghana. We expected the moist and semi-deciduous forests to be the ones that have most of these trees. However, also some of the wet evergreen forests and hyperwet forests had large stocks of timber species as well (compare Figure 8.3 with Figure 4.2C). This is partly because the moist semi-deciduous forests have been heavily depleted over the last century, much more so than the evergreen forests. When we compare this potential timber map (Figure 8.3) with the diversity map (Figure 6.7) it is clear that there is no simple link between the potential timber stock and the biodiversity. A general relation exists, however: the lower the timber density is, the higher the diversity. This is also shown for Ghana alone (Hawthorne & Abu Juam 1995). At regional level the negative relation between timber density and species density implies that timber use and biodiversity conservation are, in a broad sense, compatible options, based on spatial segregation. However, some concessionaires only have forests in the evergreen forest areas, and will seek to maximise the use of their concession even if on a national level this is not ideal for a conservation perspective. Comparison of this map with the forest classification shows that on a local level some hyperwet areas have high densities of timber species, but also some dry semi-deciduous or wet evergreen forests. For local people non-timber forest products are important as well. Bushmeat, for instance, is highly used and its utilisation is currently one of the most important conservation challenges in the tropical moist forest region, both in Upper Guinea as in Central Africa (Bakarr et al. 2001). In West Africa, bushmeat collection has led to drastic decline in many species, and populations of many species are near the brink of extinction. For instance, the primate subspecies Procolobus badius waldroni (Miss Waldron’s red colobus) disappeared completely from previously occupied localities in Ghana and Côte d’Ivoire (Bakarr et al. 2001). That hunting indeed is severe in most West African forests is shown by Caspary and co-workers (Caspary et al. 2001): during 1996 alone, 35.5 million

animals, with an estimated weight of 120,000 tons of carcasses, were consumed in Côte d’Ivoire. At a smaller scale plants or plant parts are used for construction, artisan products, and for the preparation of medicines (Figure 8.4). For Ghana the non-timber products are listed in Abbiw’s (1990) book. For two forests in Côte d’Ivoire, FC Haut Sassandra and FC Scio, Tra Bi (1998) has shown that a total of 304 species were used, of which 65 as sources of food, 73 for construction purposes, 99 for artisanal craftworks, 14 for hunting and 9 for fishing. No less than 182 species were used for medicines. This exemplifies the local importance of many forest plants. Integrated management guidelines are needed to be able to manage the available resources wisely. For bushmeat the blueprint provided by Bakarr et al. (2001) seems a good way ahead.

Long-term conservation and use The results in this book, and the areas prioritised for conservation, are in line with the recommendations of an international priority-setting workshop in Ghana in 1999. During that workshop a total of 41 regional priorities sites were identified from Guinea to Togo, covering approximately 235,000 km2 of land across six countries (Conservation International 2001). This is equivalent to 23% of the land in those countries. This includes 55,000 km2 of the estimated lowland evergreen forest remaining in Upper Guinea (Sayer et al. 1992b). The priority areas include all existing habitat types, from mountain forest to coastal mangrove, and from moist evergreen to semideciduous forest. For a better and more effective management of Upper Guinean forests, goals directed at conservation need to be combined with goals on short-term and long-term use by local people and state organisations. Needs of local people, and also their best indigenous knowledge of biodiversity and biological management, have to be taken into account. The currently widely advocated approach is to enable local communities to participate in the management of forests. In most West African countries, education curricula are weak in biodiversity information. Against this background, effective environmental education that combines the best of indigenous and modern information is perhaps the single most important activity in ensuring long-term biodiversity conservation. Regional cooperation in this respect may improve use of collective knowledge regarding proper management. The challenge of biodiversity conservation is a dynamic one. Although this book focuses on species and forests, this challenge is as much about people. Integrated management plans are necessary, cutting across sectors in

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Box 8.3 Conclusions and recommendations Plant diversity hotspots • Three prime areas of high plant species diversity are distinguished: a belt 50-100 km inland from Sierra Leone, through Liberia to southwest Côte d’Ivoire; the wet evergreen forests of southwest Ghana, and the area around Abidjan. These are also rich in species of relatively restricted distribution. • Very rare and very local species, worthy of the forest managers’ particular attention, occur specially in five areas: the Liberian coast, around Abidjan, Mount Nimba, southwest Côte d’Ivoire and southwest Ghana. • The supposed glacial refuge areas (Mount Nimba region, Cape Three Points, Cape Palmas) deserve special conservation attention. • Hill sanctuaries, as established in Ghana, may provide a good framework for planning general protective management throughout the region. Forests from a regional perspective • The regional forest classification presented in this book is a valuable addition to the country-based ones developed earlier. Local forests now can be put in a regional context. This is of high relevance for regional forest conservation and management. • Hyperwet forest areas in Liberia and Sierra Leone need extra focus because of their regional scarcity. • Complementarity is a good principle in selection of (additional) sites for forest and species conservation. In Côte d’Ivoire an extra forest reserve (National Park) in the eastern evergreen forests is highly recommended. • Corridor networks in two areas would be highly valuable to facilitate migration, especially of some animals. One in the southwest of Côte d’Ivoire and border with Liberia, the other in southwest Ghana and border with Côte d’Ivoire. • Large forest blocks in southeast Liberia are heavily undersampled with regard to herbarium collections. Collection expeditions are highly needed. Forest condition and preservation • The condition of the protected forests (whether national parks or classified forests) is rather variable. Most protected forests suffer from high disturbance, and several do not exist any more. The condition of national parks is reasonably well. • Protection effectiveness of forests needs to be improved, but this is admittedly difficult, even to promote, in times of war and of economic recession.

society, effectively to reach and maintain conservation of forests and species. Such plans and guidelines must lead to practical measures, at all geographical levels, from the local to the international. We hope that the information compiled in this book will help develop measures that will lead to effective long-term conservation and use of the forests in Upper Guinea.

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• Ghana has most of its forest types sampled in protected forests, in Côte d’Ivoire protected forests are less well distributed. • Corridors between forest patches may improve long-term conservation of forest and forest species. • The last vast areas with moist and wet forests, notable southwest Côte d’Ivoire and southeastern Liberia need effective preservation. • Long term monitoring of forest cover is needed, at a sufficiently high resolution (30 x 30 m) to be able to map forest fragments and corridors. Forest use and management • Instead of focusing on large estate forests only, forest policy should also take into account the management of small and fragmented forest areas. Local community-based management is important in this respect. • Around National Parks alternative land use systems need to be stimulated with a focus on integration of trees in farms. Planting of trees and other innovative or proven agroforestry schemes needs stronger support. • Fire may be used as forest management tool to create specific desired vegetation types. Protection of forests from fire needs to be pursued in drier areas with maximum vigour, and this includes reduction of logging especially near the forest boundaries. • For successful restoration seed sources need to be close by. Remnant forest and gallery forests have an important role to play, wherever forest regeneration is to be promoted in previously disturbed vegetation. • Deforestation has had the upper hand in Upper Guinea, but locally farmers have led to an increase of forest area through their activities, especially in the forest-savanna boundary zone. • Fallow fields need more time for restoration of soil and for longer-term growth and development of forest products. • Forests in Upper Guinea are and have been used intensively. Timber is the major product but non-timber products play a role as well, especially at the local level. Extensive hunting of bushmeat has lead to local extinction of animal species, and may also lead to a cascade of other effects, including regeneration problems of important timber species and needs more effective control. • Biodiversity conservation is challenging. Effective longterm conservation is difficult in areas with high population pressure and political instability. Regional cooperation and involvement of local populations are strongly needed to succeed in the long run.

Acknowledgements We thank Erika van Duyl for calculating the data of Table 8.1 from our GIS database.

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B Species Chapters 9-11

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M. Holmgren, L. Poorter, A. Siepel, F. Bongers, M. Buitelaar, C. Chatelain, L. Gautier, W.D. Hawthorne, A.T.F. Helmink, C.C.H. Jongkind, H.J. Os-Breijer, J.J. Wieringa and A.R. van Zoest

Ecological profiles of rare and endemic species

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Upper Guinean forests still harbour a large number of endemic animal and plant species. However, these forests are disappearing rapidly (Chatelain et al. chapter 2). Species that are rare, or have small distribution ranges are especially vulnerable to such habitat alteration or loss, and may eventually go extinct if conservation measures are not taken soon (Holmgren et al. chapter 7). For an effective conservation policy, we need information on the distribution and ecology of these rare and endemic species (Myers et al. 2000, Conservation International 2001). In this chapter, we present ecological profiles for 286 rare and endemic species, representing c. 10% of the total forest flora of Upper Guinea. Rare species are not easily recognised by tree spotters in the field. We therefore opted to use existing herbarium collections as a data source. Botanical collections have the advantage that they provide an existing source of information, cover a large geographic range, and are likely to be identified correctly. Information on the taxonomy, ecology, and environmental requirements of these species is scarce at best, or sometimes even non-existent. We have made an effort to summarise the existing knowledge on these species, and to give a first idea of their environmental requirements by combining the information on herbarium labels with GIS layers on the environment. In the following pages, we describe the metadata behind the ecological profiles. First we present information on species selection and occurrence, then on the environmental data and the analyses used, and finally we describe the format of the ecological profiles.

Species data Species selection We made a selection of just over 1000 endemic and rare species of the closed forests of Upper Guinea based on the 2nd edition of the Flora of West Tropical Africa (Keay 1954, 1958a, 1963, Hepper 1968, 1972), inventories in Ghana (Hall & Swaine 1981, Hawthorne 1995a), taxonomic revisions, and new herbarium collections (see

Figure 9.1 Campylospermum amplectens, an endemic tree species of wet evergreen forests.

Jongkind Chapter 11). All life forms were included (trees, shrubs, lianas, herbs, parasites, saprophytes and epiphytes). From this list, we selected 286 species that have been taxonomically well described, and for which there is enough certainty about their correct identification in the different herbaria. In this sample we aimed to preserve as much taxonomical diversity as possible, including species with different life forms and distributions. This means that species were selected from all families, taking a smaller fraction of species from the large families (with many species in this area). The 286 taxa of this final subset represent c. 10% of the total forest flora of Upper Guinea. 223 of these 286 taxa are considered to be endemic to Upper Guinea. They belong to 75 different families. Amongst these taxa, there are 126 trees, 35 shrubs, 79 lianas, 36 herbs, 9 epiphytes, and 1 saprophyte. This means that the sample is relatively rich in trees, since in the Upper Guinean flora the number of trees, shrubs and lianas is rather similar (W.D. Hawthorne, pers. comm.). Species occurrence Data on species occurrence come from two different sources; herbarium collections, and rapid botanical surveys. We entered into a herbarium database (BRAHMS 4.8, Filer 2000) all herbarium specimens collected from Senegal to Togo for the 286 selected species. For some non-

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endemic rare species, the herbarium specimens collected in Lower Guinea (the Congo Region) were included as well. We checked and entered all specimens from the National Herbarium The Netherlands, Wageningen branch (WAG: Herbarium Vadense, The Netherlands), National Botanical Garden of Belgium (BR: Meise, Belgium) and Royal Botanical Gardens Kew (K: Kew, Great Britain). In addition, we added the collections from Côte d’Ivoire present in the Geneva herbarium database (G: Conservatoire et Jardin Botaniques de la Ville de Genève, Switzerland) the collections from Côte d’Ivoire and Guinea present in the Paris herbarium database (P: Muséum National d’Histoire Naturelle, Paris, France), and the collections of 100 species present in the Herbarium of the University of Ghana (GC: Legon, Ghana). These data were complemented by distribution data from published taxonomic revisions for c. 20% of the species. Our total database contained 48,000 records from Upper Guinea, of which about 6900 records belonged to our subset of 286 rare and endemic species. For the 286 species, the number of records in Upper Guinea varies from 1 to 171 records per species, with a median of 14. At first sight this may seem low but these are, by definition, uncommon species. The database is of course not exhaustive, but it gives a fair indication of the state of knowledge for these species in the region. In Ghana, information on plant distribution does not only come from herbarium records, but also from rapid botanical surveys. These rapid botanical surveys have been carried out by Hall & Swaine (1981) and Hawthorne & Abu-Juam (1995). With a rapid botanical survey one identifies all vascular plants occurring in a small area (mostly 25 x 25 m). 748 survey samples have been made in Ghana, including 155 Hall & Swaine A-plots, 130 Hall & Swaine B-plots, 8 Hall & Swaine C-plots, and 455 Hawthorne & Abu-Juam plots. The plots of Hall & Swaine were situated in closed forest. The A samples consist of 25 x 25m plots, in which all species present were listed. The B samples consist of lists of the first 30 -60 species encountered. Hawthorne & Abu-Juam’s plots are comparable to the A samples, with the exception that plots were mostly established in swamps, plantations, logged areas and other abnormal forests not included in the Hall & Swaine survey. Each sample contains about 100 plant records (range 50 -175). In total the rapid botanical survey database contains 66,400 records.

Environmental data Plant distributions are usually strongly shaped by water availability, soil fertility, and altitude. Maps of these environmental variables were prepared and included in a Geographical Information System (ArcView, ESRI Inc.).

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Rainfall, water holding capacity, and proximity to rivers Water availability for plants is the combined result of several factors such as rainfall, soil water holding capacity, temperature, and the proximity to rivers, lakes, and sea coast. We compiled annual rainfall data from 578 weather stations distributed from Senegal to Nigeria, Burkina Faso, Mali and Niger using different sources (Myers & Staff 1981, ANAM 1987, ICCARE 1994, Global Historical Climatology Network). We used these data to make a rainfall map using the inverse distance weighting interpolation method in ArcView. Soil water holding capacity was derived from the FAO Digital Soil Map of the World (FAO 1995b) which has a scale of 1:5,000,000. Small-scale differences in soil types due to topographical variation are not visible on this map. For each gridcell (55 x 55 km), we calculated an average soil water holding capacity (in mm water/m soil), based on the relative cover of different soil types, their depth, and texture. It was presumed that sandy soils have a water holding capacity of 75 mm/m, loamy soils of 100 mm/m, and clayish soils of 125 mm/m. The position of perennial rivers, lakes, and coastline was obtained from the Digital Chart of the World (ESRI 1984). Soil fertility We used the sum of exchangeable cations (CMK; Ca2+, Mg2+, K+, in cmol cations per kg soil) and median Cation Exchange Capacity (CEC; %) as indicators of soil fertility. For each grid-cell, we calculated an average soil fertility based on the relative cover of the different soil types and their median soil fertility. The relative cover of the different soil types was derived from the FAO digital soil map of Africa. The median CMK or CEC value of the dominant soil type came from a review of the chemical properties of soil profiles by Batjes (1997). Altitude Elevation data were derived from the Digital Chart of the World (US Geological Service, EROS Data Center, 1984), which has a spatial resolution of c. 1 km. Potential forest cover The potential forest cover represents a mosaic of mature forest, secondary and degraded forest and forest converted to plantations, agricultural fields and urban areas. The potential forest cover of Upper Guinea (SenegalNigeria) was based on the map of Paivinen et al. (1992). They used a NOAA-AVHRR image of 1984 with a spatial resolution of 1 km2 to classify the vegetation. For our forest cover layer we pooled the mature forest, the mosaic of mature and disturbed forest, and the mangroves. The potential forest cover of Lower Guinea was based on the 1992 JRC-TREES map of Central Africa, with an update with SPOT vegetation imagery of 1998 (Eerens 1998). This map has a spatial resolution of 1 km2. For our forest cover layer we pooled the moist forest, secondary forest and rural complex.

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Analysis Species distribution Species were classified into different distribution types based on the continuity of their distribution, their range, and their commonness. A distinction was made between species having a continental disjunct distribution (disjunct between Upper and Lower Guinea), a regional disjunct distribution (disjunct populations within Upper Guinea), and a continuous distribution (a more or less continuous occurrence within its range). The classification was based on a visual interpretation of the species distribution map, and a histogram of multiple neighbour distances for the species occurrence records. For every species, we calculated the distance between each collection point and all the other collection points of that species. For example, for a species with five observations this would result in 10 neighbour distances. A histogram of the frequency of these neighbour distances showing a unimodal distribution was interpreted as an indicator of a continuous distribution (Fig. 7.2A). On the other hand, species with a bi-modal histogram were interpreted as having a disjunct distribution (Fig. 7.2B,C). Every mode or peak represents a clump, with the height corresponding to the number of collections within a clump. The distance between the peaks corresponds to the distance between the clumps. Obviously, continental disjunct species (e.g. with a population in Liberia and a population in Gabon) usually have a much larger distance between the two peaks than regional disjuncts (e.g. with a population in Liberia and a population in Ghana). The classification of spatial patterns based on the multiple neighbour distance histograms was in close agreement with a visual interpretation of the species distribution maps. Species range For species with a continuous distribution, we estimated the species range based on the maximum distance found between collection points. The species range was classified as being very local (maximum distance < 100 km), local (100-300 km), regional (300-600 km), or widespread (> 600 km). The very local and local categories fit the most widely used criterion for tropical plant species endemism (Gentry 1992). Species commonness The commonness of a species in Upper Guinea can be expressed as the total number of records found in the region from Senegal to Togo. This measure is potentially influenced by collector’s bias. Rare species are likely to be over-represented, as they might be more interesting to collect by botanists. A more robust measure of commonness is the frequency of half-degree grid cells in which the species is collected. If a species had been

collected many times in the same forest reserve close to a capital or research station, this would result only in one presence record of a half-degree grid cell. In our database, both measures are highly correlated (Pearson’s r = 0.92, P < 0.001, n = 286) which indicates consistent estimates of a species’ commonness. Environmental requirements To relate a species occurrence to the environmental variables, we used only the records of Upper Guinea, because for this region all herbarium collections had been completely entered into our database. We used two different approaches; a frequency analysis and a logistic regression analysis. For the frequency analysis the continuous environmental variables were divided into categories and the number of records for each species counted and assigned to each category. The environmental variables and categories were: altitude (below or above 500 m), distance to rivers (within or beyond 2 km from each riverside), distance to the coast (within or beyond 5 km from the coast), annual rainfall (< 1500, 1500-2000, 2000-2500, > 2500mm /yr), soil water holding capacity (< 50, 50-85, > 85 mm water/m soil), and CEC (0-4, 4-8, > 8%). A Chi2 analysis was carried out for each combination of species and environmental variable to evaluate whether the observed distribution of the species differed from the expected distribution. The “expected” distribution was based on all records of all species (irrespective of life form, or commonness) found in the database (46,333 records). We analysed 181 species for which there were sufficient records (n ≥ 10). A logistic regression analysis was carried out to model the species occurrence in Upper Guinea as a function of four environmental factors: altitude, annual rainfall, soil water holding capacity, and available cations (CEC: Cation Exchange Capacity). We used stepwise forward regressions with the simple and quadratic variables, but without including interaction terms. A significant simple variable indicates an increasing or decreasing probability to find the species with an increase in the environmental variable. A significant quadratic variable indicates that the species shows a bell-shaped response curve towards that environmental variable (Jongman et al. 1987). Since the species had not been collected at random and since we had only presence data (herbaria records), we needed to assume absence under certain conditions. For this we selected for Upper Guinea only those half-degree grid cells (55 x 55 km) with 50 or more records of all species occurring in the area (thus including only cells that have been botanically well explored). This resulted in a total of 145 cells. For some cells there were no data for all four environmental variables. This reduced the total number of cells to 140. We assumed that a species was absent when no records of this species were found in those cells. A logistic regression analysis has sufficient resolution to detect significant patterns when there are sufficient numbers of presence and

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absence. In order to have a reasonable number of cells with presence, we ran the analysis only on those species present in ten or more cells (i.e. > 7% of the total amount of cells). This resulted in a total of 112 species. The Chi2 analysis and logistic analyses each have their strengths and weaknesses (Jongman et al. 1987). The advantage of the Chi2 analysis is that it remains as close to the original data as possible, and that it presents a simple and straightforward test. The disadvantage is that the definition of the class boundaries is arbitrary, and that the test is not very powerful. The advantage of the logistic regression analysis is that it provides a compact description of species responses to the environment, and that the relative importance of different environmental variables for species occurrence can be weighed. The disadvantages are that absences are assumed but not measured, and that the spatial resolution is not very high as the sample units are quite large (55 x 55 km). Habitat openness A species’ shade tolerance plays a fundamental role in understanding its response to the environment. For many rare species such information is not readily available. As a first step in defining the light requirements of a species, we calculated the percentage of records of each species found in open habitats. Using the herbarium collector’s notes that had been added to the database, we searched for keywords for open places (e.g., bord de piste, border, broken canopy, bush, bushes, bushland, clairière, clearcut, clearing, cultivated, degraded, dénudée, disturbed, edge, farm, fourré, fringe, fringing, gap, jachère, lisièr, open, regrowth, roadside, along (the) road, near the road, savanna, scrub, secondaire, secondary, secondary bush, secondary forest, secondary regrowth, secondary vegetation, shrubland, sunny, thicket(s), etc.). To calculate the proportion of records found in open or recently disturbed habitats we used only the subset of records for which clear habitat descriptions were present. Guilds Species were classified into guilds as pioneer, nonpioneer, and shade bearers mostly based on the reported shade tolerance in Ghana (Hall & Swaine 1981, Hawthorne 1995a). Pioneers are species with the highest light demand, of which seedlings are only found in gaps and older plants are absent or very rare in the forest understorey. Non-pioneer light demanders are species with an intermediate light demand, of which seedlings are common in the understorey whereas adults are not (i.e. they require a gap to grow). Shade bearers are those species for which both young and older plants are frequently found in the shaded forest understorey. We compared the percentage of records found in open places in our database with the classification made by Hawthorne (1995a). Indeed, we found that pioneer species had a higher abundance in open places (t-test, t = – 2.37, p = 0.026), but we did not find a difference between shade

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bearers and non pioneer light demanding species. Based on this, we classified the species lacking a literature description on shade tolerance using the proportion of herbarium records found in open places. We classified them as pioneers (> 65% records in open places), non-pioneer light demanding species (30-65% records in open places), or shade bearers (< 30% in open places). Forest types We used the herbarium labels and literature to describe the forest types in which the species occurs. The vegetation typology differs strongly between the countries in Upper Guinea (Bongers et al. chapter 4), and the interests and background of the collector. To facilitate comparisons we tentatively regrouped the wealth of terminology used in a few classes belonging to zonal forest formations (occurring along a rainfall gradient), azonal forest formations (caused by locally overruling site conditions such as waterlogging or special soils), and forest formations that were subjected to different levels of disturbance. We realise that by reducing the number of categories some information is lost and that inevitably some errors have been made, but we believe it provides the reader with a clearer comparative framework. Zonal forest formations Rainforest: primary forest, dense forest, high forest, dense natural forest, undisturbed forest, sacred woods, old stands, mature forest, residual forest, forest remnant, forest island Wet evergreen forest: wet forest, wet evergreen forest, wet primary forest relict, lowland rainforest, rainforest, evergreen primary rainforest, primary hygrophilous forest, humid evergreen forest, humid rainforest, dense humid forest, deep humid forest Moist evergreen forest: moist forest, moist high forest, subhumid dense forest, dense mesophytic forest (Semi)deciduous forest: semi-deciduous forest, dense semi-deciduous forest, deciduous forest Moist semi-deciduous forest: moist semi-deciduous forest Dry semi-deciduous forest: dry semi-deciduous forest Dry forest: dry evergreen forest, dry forest, southern marginal forest, southeast outlier forest Savanna-Woodland: savanna, savanna forest, savanna woodland, tree savanna, woodland Shrubland: shrubland, bushland, dry thickets Azonal forest formations Coastal forest: coastal rainforest, coastal evergreen forest, littoral forest, coastal forest Coastal shrubland: coastal thickets, coastal bushland, coastal scrub, coastal shrubland, coastal savanna Gallery and Riverine forest: floodplain forest, floodplain, seasonally flooded forest, gallery forest, riverine forest, riverine forest in savanna, gallery forest in savanna

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Swamp forest: marshy forest, swamp forest, swampy thickets, swamps in deciduous forest Montane forest: montane forest, submontane forest, (sub)montane rainforest, misty forest, montane gallery forest Upland evergreen forest: upland evergreen forest, upland forest Forest formations varying in disturbance Disturbed forest: degraded primary forest, logged forest, exploited forest Secondary forest: secondary regrowth, secondary vegetation, secondary rainforest, secondary forest, old secondary forest Thickets: thickets

Explanation of the species descriptions In this chapter we present an ecological description (or profile) of the rare and endemic species. The information for each species is presented in one page. In some cases, though, two subspecies, varieties, or sister taxa are grouped on the same page. Each species profile includes a short botanical and ecological description, a distribution map, a photo or drawing, and an environmental table. The aim of the botanical description is to offer an impression of the species to the general reader. For a more detailed taxonomical description we refer the reader to the Flora of West Tropical Africa, the recently published The Woody Plants of Western African Forests (Hawthorne & Jongkind 2004), or any other recent species revision in journals or books. The botanical description is followed by a summary of the species distribution type, range and abundance. The ecological description of the species summarises its requirements for light, water and nutrient availability. It also indicates the species regeneration and growth characteristics. Each description is accompanied by a table with a quantitative summary of the environmental requirements. We finally describe some of the main uses of the species and present the most relevant literature references. Species distribution maps A species distribution is presented in two, and sometimes three different maps. Obviously, these maps can not be completely comprehensive, and will probably significantly change when our knowledge of today’s poorly explored regions increases, or when more collections get included into our database. Species occurrence at the national level. A small map shows either Upper and Lower Guinea, or Africa completely. Countries where the species occurs are indicated in orange. The rectangular frame indicates the

area for which a larger, more detailed distribution map is shown. Detailed species distribution map. The species distribution map contains several geographical features (rivers, lakes, country borders) to facilitate the reader orientation. The potential forest cover and mountainous areas are shown as coloured layers, and give a first hint of the species’ environmental requirements. To distinguish between lowland and upland areas a threshold level of 500 m was used (cf. Whitmore 1998). Lowland forest areas are indicated in light green, upland forest areas in dark green, lowland non-forest areas in light yellow, and upland nonforest areas in dark yellow. Data on species occurrence come from two different sources: herbarium collections, and rapid botanical surveys (see above). The herbarium collections are represented by red circles. In general they have a geographical precision of 0.1-10 km. Herbarium collections for which the identification is doubtful are represented by open circles. The herbarium collections cover a time span of 100 years. It is possible that a species is shown to occur in an area where the forest has already disappeared. The map gives therefore an impression of the potential distribution of a species. Species occurrences, as observed during rapid botanical surveys are represented by red triangles. Rapid botanical survey data are only available for Ghana, and only those species are shown that could be reliably identified in the field. Predicted species occurrence. Species occurrence in Upper Guinea was modelled as a function of several environmental factors (see above). For those 46 species for which these models have a reasonable predictive power (i.e. the explained variance is more than 25%), the predicted probability of species occurrence is shown in 5 classes (< 5%, 5-25%, 25-50%, 50-75%, and 75100%). This does not necessary mean that a species should be found in those areas where it is predicted to occur. It simply indicates that, given the environmental conditions, the species may be found in that area with certain probability. It can be absent from that area if, for example, dispersal limitation or competition prevents the species from filling that niche. Description Guild: species are classified into four different guilds based on their light requirements for germination and survival; pioneer (pi), non-pioneer light-demander (np), shade-bearer (sb) and unknown (u). Life form: species are classified into six different life forms based on their maximal height and position of the perennating buds; herb, shrub, tree, climber, epiphyte and saprophyte. Herbs are either erect, stoloniferous or rhizomatous. Trees are assigned to different size classes, depending on their maximal height (pigmy < 2 m, small 2-10 m, medium 10-30 m, large > 30 m). Climbers are classified into different length classes (small < 10 m, large > 10 m), climbing mechanisms (winding, climbing with tendrils, etc.) and woodiness classes (woody,

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herbaceous). Lianas (woody climbers) can start their life as shrubs. They climb actively only in a later stage of their life cycle, or when triggered by an environment condition. For those cases, the species is described as being a climber, but a sentence is added about its shrubby nature when young. Max. height: maximum height (in m) according to either literature sources or herbarium labels. Max. diameter: maximum diameter (in cm) according to either literature sources or herbarium labels. Leaf: leaf characteristics are described in a fixed order, starting with their arrangement (e.g. opposite, alternate, whorled), whether they are simple or compound (palmate, paripinnate, imparipinnate), shape (e.g. ovate, obovate, elliptic), leaf size class (for compound leaves we used the size of the leaflets) (leptophyll (< 0.25 cm2), nanophyll (0.25 - 2.25 cm2), microphyll (2.25 - 20.25 cm2), notophyll (20.25 - 45 cm2), mesophyll (45 -182 cm2), macrophyll (182 -1640 cm2), megaphyll (> 1640 cm2)), blade size (minimum and maximum width and length in cm, excluding the petiole), margin (e.g. entire, lobed, dentate, serrate), texture (coriaceous, herbaceous), hairs (this is not always given, glabrous, pubescent), colour (not always given), and incidentally some information about the petioles and stipules. The description of different leaf units is separated by semicolons. Inflorescence: position (terminal, axillary, or cauliflorous), whether it is a simple flower or an inflorescence (unbranched or branched). Sometimes a more detailed, technical description of the inflorescence is given in parenthesis, such as panicle, spike, umbel, etc. This is not always done, as classification into inflorescence types is not always straightforward. Sometimes data on the size are given. Flower: size (tentatively defined as being small < 1 cm length, medium 1- 4 cm, or large > 4 cm), calyx (colour, shape and size), corolla (colour, shape, and size), flower form and colour. The description of different flower units is separated by semicolons. Fruit: dry or fleshy. Sometimes a more detailed, technical description of the fruit is given in parenthesis, such as berry, drupe, samara, etc. Shape (round, ovate), size (width and length, in cm), texture, colour, the amount of seeds (one, few, many). Other: any other striking descriptive features such as changes in life form with age (i.e. erect shrub when young, later a liana) buttresses, latex, crown form, etc. Distribution Continent: occurrence in countries outside Upper Guinea (e.g. Nigeria, Cameroon). Otherwise, the species is described as an Upper Guinea endemic. When the information does not come from our database, the literature source is indicated. Upper Guinea: occurrence in countries within Upper Guinea from Senegal to Togo. Countries are listed according to their west-east position along the coast. When the information does not come from our database, the

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literature source is indicated. Distribution: distribution type based on the continuity of its occurrence: continuous, Upper Guinea Disjunct (disjunct within Upper Guinea), Continental Disjunct (disjunct between Upper and Lower Guinea). Species range based on how widespread the species is (very local, local, regional, widespread). Species commonness (number of half-degree grid cells in Upper Guinea in which the species has been observed). Species range (maximum distance between two collections). The maximum distance is only given for species that have five or more records in our database and that have a continuous distribution or an Upper Guinea Disjunct distribution. The conservation status of a species is given in the following categories: extinct, extinct in the wild (known only to survive in cultivation), critically endangered (facing extremely high risk of extinction in the immediate future), endangered (very high risk of extinction in the near future), vulnerable (high risk of extinction in the medium-term future), lower risk (none of the above), based on the IUCN red list of 2000. Forest type: the forest types in which the species is found according to literature or herbarium descriptions. Forest types are organised from wet to dry forests (wet evergreen, moist evergreen, moist semi-deciduous, dry semi-deciduous, dry forest, savanna), particular habitat (e.g. gallery forest, coastal forest, montane forest), and disturbance degree (e.g. secondary forests, mature forests). Habitat This section reports the environmental conditions where the species has been observed to grow in relation to light (e.g. shady places in the understorey), water (e.g. close to water sources such as a creek, river, lagoon, or waterfall), soils (e.g. sandy soils) and altitude. The environmental requirements are either derived from the literature, or inferred from the logistic regression analysis and Chi2 analysis (see above). Regeneration This section contains details on the initial part of the life cycle of the species, such as germination and establishment, sapling abundance, etc. Seedlings are classified into different seedling types (cf. Garwood 1996), dependent on the exposure of the cotyledons (exposed, phanerocotylar or hidden within the seed, cryptocotylar), the position of the cotyledons (above ground, epigeal or below ground, hypogeal), and whether the cotyledons are photosynthetically active (i.e. leaf-like, foliaceous or not, reserve). Although eight combinations are possible, three of them have never been described or reported. Five types remain: phanerocotylar epigeal reserve (exposed, above ground, storage cotyledons), phanerocotylar epigeal foliaceous (exposed, above ground, leaflike cotyledons), phanerocotylar hypogeal reserve (exposed, below ground, storage cotyledons), and cryptocotylar hypogeal reserve (cotyledons that remain hidden in the seed, below ground).

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Phenology Deciduousness: whether a species is deciduous (loses the leaves during some period of the year) or evergreen (maintains leaves during all seasons). Dispersal: main dispersal mode has been classified as explosive (active expulsion of the seeds), wind, animal, water, and barochore (simply falling). Timing: the flowering and fruiting period reported in the literature. Note that phenological histogram may deviate from this, depending on the latitudinal and altitudinal positions, and the interannual variation in climatic conditions. A phenological histogram was made using the information on flowering and fruiting found on the herbarium labels of every collection. The histogram runs from July to June, so that the dry season with the most phenological activity (December-March) is found in the centre of the graph. The number of reproductive individuals upon which the histogram is based is indicated as well. Uses Commercial or domestic use. Data sources References used for the species description, or where more general information about the species can be found. The references are given in chronological order. Environmental table The table provides a quantitative summary of the environmental conditions where the species has been observed to grow. We indicate which percentage of the herbarium records for that species is found in open, disturbed places (open), above 500 m altitude, within 2 km distance from rivers, or within 5 km from the coast. The rainfall gradient is divided in four classes; Dry (< 1500), Medium (1500-2000), Wet (2000-2500) and Very Wet (> 2500 mm/yr). The soil Cation Exchange Capacity (CEC) is classified as Low (0-4), Medium (4-8) and High (> 8%) and the Water Holding Capacity (WHC) of the soil is subdivided in Low (< 50), Medium (50-85) and High (> 85 mm water/m soil). Note that the number of observations of a species (n) is smaller for “openness” than for other environmental conditions. The reason is that the openness was based on all collections of a species with enough and clear habitat information provided by the specimen collector on the collection label. In many cases, such habitat description was unavailable. The information on the environmental conditions where a species grows is particularly interesting for comparisons with other species. To this end we present in the first line the information on the focal species, and in the second line the environmental requirements of an “average” plant in the database. The environmental requirements of an “average” plant was based on all records of all species (irrespective of life form, or commonness) found in the database (46,333 records).

Acknowledgements This chapter is the result of the joined efforts of many co-authors. Marjo Buitelaar and Hendrikjan Os Breijer entered the herbarium collections into the database. William Hawthorne provided inventory data for Ghana and some species drawings and photos. Cyrille Chatelain and Laurent Gautier provided data for Côte d’Ivoire from the Genève herbarium. Jan Wieringa selected the rare and endemic species to be described and he and Carel Jongkind checked species identifications and distribution maps, and commented on the species descriptions. Toon Helmink and Roland van Zoest made the species distribution maps and did the GIS analysis. Frans Bongers made valuable suggestions that helped shaping the contents of this chapter. Almira Siepel made the graphics and several text revisions. Milena Holmgren and Lourens Poorter designed the chapter, did the statistical analysis and data interpretation, and wrote the species descriptions. We all deeply thank the contributions of a very large number of people along these years. We would especially like to thank the extraordinary work of Denis Filer for creating the user-friendly Brahms 4.8 database and making our task much easier. Folkert Aleva permanently supported the database management work. Marieke van Bergen helped with the selection of rare and endemic species. Numerous people contributed to the arduous work of entering over 12,500 herbarium collections into the database. We like to thank especially Stuart Cable, Patrick Epke, Tom van Lokven, Marieke Sandker, Arjan Schoonhoven, Jan van Veldhuizen, Petra Wilbrink, and Saskia Woudenberg. Several researchers generously allowed us to use their species distributions data: Marc Pignal (Paris database), Roger Polhill (Loranthaceae), Petra De Block (Ixora), Bonaventure Sonké (Oxyanthus), Piet Stoffelen (Pausinystalia and Corynanthe). We were able to beautifully illustrate this chapter thanks to the drawings of Rosemary Wise, Marjolein Spitteler and Emmelien Jaggar, and to the photos provided by many authors listed on page 2. The chapter was reviewed by many specialists in different species groups: Jan-Just Bos (Dracaenaceae), Carel Jongkind (Combretaceae), Jan Wieringa (Aphanocalyx and Tetraberlinia), Joost van der Burg (Orchideaceae), Mark Sosef (Begoniaceae and some additional species from Gabon), Frans Breteler (Dichapetalaceae), Lytton John Musselman (Saxicolella), Heinjo During (Selaginella) and Francois Kouamé. We especially thank the great job of Marjolein de Vette for the chapter layout.

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Key to symbols in maps and tables of chapter 9 Species distribution map

based on herbarium collections

Forest zone altitude < 500 m Forest zone altitude 500-1000 m Forest zone altitude > 1000 m Altitude < 500 m Altitude 500-1000 m Altitude > 1000 m Location of the species (rapid botanical survey - Ghana) Location of the species (herbarium data) Location of the species (herbarium data - uncertain identification) Border Rivers and lakes Ocean

Species occurrence at the national level Species present Species absent Ocean

Predicted species occurrence 0-5 % probability 5-25 % “ 25-50 % “ 50-75 % “ 75-100 % “ Insufficient environmental data

Environmental tables spp n Open (n) Altitude River Coast

species number of records % of records found in open habitats % of records found at altitudes > 500 m % of records found within 2 km of rivers % of records found within 5 km of the coast

Mean annual rainfall

Soil CMK

Soil WHC

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% of records found in (D) relatively dry areas: < 1500 mm/yr, ( M ) intermediate areas: 1500-2000 mm/yr, (W) wet areas: 2000-2500 mm/yr, and (VW) very wet areas: > 2500 mm/yr % of records found on soils with (L) low availability of cation: 0-4 cmolc/kg, (M) intermediate availability of cation: 4-8 cmolc/kg, and (H) high availability of cation: > 8 cmolc/kg % of records found on soils with (L) low water holding capacities: < 50 mm water/m soil, (M) intermediate water holding capacities: 50 - 85 mm water/m soil, and (H) high water holding capacities: > 85 mm water/m soil

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Acanthus guineensis

Heine & P.Taylor

Acanthaceae A

Description

Phenology

Guild: pi Life form: perennial erect herb Max. height: 2 m (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, elliptic, mesophyll (4-9.5 x 13-29 cm), deeply lobed, dentate and spinose, herbaceous; shortly petiolate, without stipule Inflorescence: terminal, not branched (spike) Flower: medium-sized (2 cm long); calyx pale green, reddish veined; corolla white, upper lobe with reddish veins, lower lobe with yellow-green veins; bracts 2-2.5 cm long, hairy Fruit: dry dehiscent (4 cm long) Seed: medium-sized (0.6 x 0.5 x 0.2 cm), brown Other: an erect almost unbranched herb with a light-green smooth stem.

Deciduousness: evergreen

Distribution

FWTA

Data sources

Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: Upper Guinea disjunct, widespread, present in 19 30’ cells, distribution range is 1440 km Forest type: wet evergreen forest, secondary forest

Habitat Species occurrence increases significantly at places with rainfall higher than 1500 mm/yr (Chi2 test). In moist places of the forest (e.g. by streams or rivers). Occasionally along roadsides (under bush shade) or along cultivated fields (by stream banks). On clayish soils (herbarium).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

A.g.

28

25 (8)

11

39

0

0

43

32

25

0

50

50

39

0

H 21

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

109

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Page 110

Acridocarpus alternifolius

(Schum. & Thonn.) Niedenzu

Malpighiaceae A

Habitat The species is more abundant in lowlands (logistic regression analysis, Chi 2 test), and in regions where rainfall is between 1500-2000 mm/yr (Chi 2 test). It is mostly found in disturbed areas (e.g. roadsides, thickets, and secondary forests), although it has also been described as an understorey climber in very dry forests (Hall & Swaine 1981). Occasionally reported near waterfalls or in swamps. On clayish, sandy and sandy-rocky soils (herbarium).

Regeneration

Description

It regenerates in the shade (Hall & Swaine 1981). Guild: pi Life form: slender winding woody climber Max. height: 6 m (Kasparek 2000) Max. diameter: data unavailable Leaf: alternate, simple, obovate, mesophyll (3-10 x 6-28 cm), entire, coriaceous, sessile glands beneath; no stipules Inflorescence: terminal or axillary, subcorymbose to racemose Flower: medium-sized; bowl-shaped; calyx reddish with sessile glands; corolla yellow to pale orange, 5-merous Fruit: dry indehiscent, winged (samara) (wings up to 2 x 6 cm), pinkish to red, up to 3 fruit parts per flower, each part dropping separately; up to 3 seeds, but usually 2, with 1 seed and 1 wing per fruit part Seed: medium-sized Other: it starts as a shrub when young.

Phenology Dispersal: by wind

Distribution Continent: Guinea wide: from Sierra Leone to Nigeria (Hall & Swaine 1981), Nigeria (herbarium) Upper Guinea: Guinea, Sierra Leone, Côte d'Ivoire, Ghana

spp

110

n

Open (n)

Data sources

Distribution type: continuous, widespread, present in 13 30' cells, distribution range is 1780 km Forest type: moist evergreen forest, dry semideciduous forest, secondary forest, thickets, swamp forest. In Côte d'Ivoire, typical of moist evergreen forests (Kasparek 2000), whereas in Ghana, it is mostly found in dry forest types (Hall & Swaine 1981).

Hall & Swaine (1981), Kasparek (2000), Hawthorne & Jongkind (2004)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

A.a.

27

100 (14)

0

41

37

15

78

7

0

26

41

33

19

30

H 41

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 108-118

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Page 111

Acridocarpus plagiopterus

Guill. & Perr.

Malpighiaceae A

Description

Habitat

Guild: pi Life history: large winding woody climber Max. height: 6.1 m (FWTA) Max. diameter: data unavailable Leaf: alternate, simple, very variable in shape and size, from linear to obovate, mesophyll (5-8 x 12-38 cm), entire, coriaceous, young leaves pink on lower surface with red glands, hairy, up to three pairs of glands at the base of the lower leaf surface; petiole 1 cm long, hairy; no stipules Inflorescence: terminal or axillary, unbranched (raceme, approx. 11 cm long) Flower: small (approx. 1 cm long); calyx brown; corolla bright orange yellow, 5-merous Fruit: dry indehiscent, winged (samara), (0.5 x 0.5 cm), up to 3 fruit parts per flower, each part dropping separately, very dark red; 1 seed and 1 wing per fruit part Seed: medium-sized (0.4 x 0.2 cm) Other: it starts as a scandent shrub. Appearing as a rhizomatous shrub when yearly burnt by bush fires. The twigs are hairy and the slash has no exudate.

It is found both in wet (e.g. along rivers, humid forests) and dry places (e.g. dry secondary bush, degraded thickets, dry soil), and often also along roadsides. Nevertheless, species occurrence increases significantly with rainfall (logistic regression analysis), especially in places with an annual rainfall higher than 2500 mm/yr (Chi2 test). We found it to occur very frequently in coastal ranges (Chi2 test), but according to Hawthorne & Jongkind (2004) it is found in wet forests at higher altitudes.

Phenology Dispersal: by wind

Distribution Continent: Upper Guinea endemic Upper Guinea: Senegal, Guinea-Bissau, Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 27 30’ cells, distribution range is 1904 km Forest type: wet forest, dry forest, secondary forest, thickets

spp

n

Open (n)

Data sources FWTA, Hawthorne & Jongkind (2004)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

A.p.

52

54 (24)

13

38

23

17

30

25

26

0

48

52

35

21

H 17

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

111

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Page 112

Adhatoda robusta

C.B.Clarke

Acanthaceae A

Description

Phenology

Guild: np Life form: shrub or small tree Max. height: 8 m (herbarium) Max. diameter: 20 cm (herbarium) Leaf: opposite, simple, elliptic, macrophyll (6-14 x 18-36 cm), entire to crenate; petiole up to 4 cm, pubescent Inflorescence: terminal, branched (large erect panicles) Flower: medium-sized; white or pale yellow to reddish or violet; bracts pubescent Fruit: dry dehiscent, brown-black Seed: medium-sized (0.5 x 0.4 cm) Other: the bark outside is green with globular lenticels. The sapwood is yellow-white.

Data sources FWTA, Hawthorne & Jongkind (2004)

Distribution Continent: Nigeria, Cameroon, Equatorial Guinea, Gabon, Democratic Republic of Congo Upper Guinea: Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 4 30’ cells, distribution range is 3644 km Forest type: montane forest, wet evergreen forest (FWTA), semi-deciduous forest, secondary forest (herbarium)

Habitat Often found in secondary forest along roadsides and plantations (herbarium). Sometimes gregarious on dry rocky hills (Hawthorne & Jongkind 2004). Occasionally found in moist places of the forest (e.g. by waterfalls).

spp

112

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

A.r.

9

50 (4)

11

67

0

33

67

0

0

0

33

67

22

0

22

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 108-118

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Page 113

Aframomum atewae

Lock & Hall

Zingiberaceae A

Description

Phenology

Guild: pi Life form: perennial stoloniferous herb Max. height: 0.5 m Leaf: arranged in two vertical rows, simple, oblanceolate to elliptic, mesophyll (4-5 x 18-25 cm), glabrous above, hairy beneath Inflorescence: separate with 1 to several flowers Flower: medium-sized (2 cm long); calyx purplebrown; corolla white, funnel-shaped, with 1 large central petal (3.5 x 2 cm) Fruit: fleshy, spherical (2 cm in diameter), glabrous, subterranean Seed: medium-sized (0.25 cm in diameter)

Dispersal: by animals (chimpanzees)

Distribution

Data sources

Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire, Ghana Distribution type: continuous, regional, present in 4 30’ cells, distribution range is 471 km Forest type: upland forest, wet evergreen forest

FWTA, Lock & Hall (1973), Hall & Swaine (1981)

Habitat In open places such as forest gaps and along roadsides (Lock & Hall 1973).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

A.a.

8

0 (7)

13

75

0

13

88

0

0

0

38

63

0

13

38

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

113

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Page 114

Afzelia parviflora

(Vahl) Hepper

Leguminosae-Caes. A

Description

Habitat

Guild: np Life form: medium-sized tree Max. height: 30 m (herbarium) Max. diameter: 50 cm (herbarium) Leaf: opposite, paripinnately compound, 6-10 leaflets, shiny above, mat or glaucous below, often with many glands along the lamina; leaflets have twisted petiolules Inflorescence: unbranched (spike) stout, partly draped by 0.4 cm long, reflexed bracts after the flower has fallen Flower: calyx green; corolla red Fruit: dry dehiscent (5 x 7 cm), woody Seed: very large, hard, black with red waxy arils Other: it has buttresses.

Species occurrence increases significantly when rainfall is higher than 2000 mm/yr to reach an optimum around 3100 mm/yr (logistic regression analysis, Chi2 test). It can be found in a wide range of habitats. It occurs in mature forest, but also in secondary forest, coastal thickets, and roadsides. Mostly at the coast but also occasionally in mountains. Often near water (along river borders and in swampy areas). It seems to be mostly a riverine or sea-side tree (Hawthorne & Jongkind 2004). Usually on rocky or gravelly soils (herbarium).

Phenology Dispersal: probably by animals

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire Distribution type: continuous, widespread, present in 18 30’ cells, distribution range is 1031 km Forest type: wet evergreen forest, savanna, gallery forest, coastal shrubland, secondary forest

Data sources Hawthorne & Jongkind (2004)

spp

114

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

A.p.

27

50 (20)

7

41

15

4

26

15

53

0

93

7

70

4

H 15

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 108-118

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Page 115

Alafia parciflora

Stapf

Apocynaceae Description

A

Phenology

Guild: pi Life form: winding woody climber Max. height: data unavailable Max. diameter: data unavailable Leaf: opposite, simple, elliptic, notophyll (2-6 x 5-11 cm), entire, herbaceous; glabrous on both sides Inflorescence: terminal or axillary, 2-3 cm long, 2-3 flowered Flower: medium-sized (up to 12 mm long); corolla pinkish, tube red, lower part yellow, centre red; glabrous outside, pubescent inside Fruit: dry dehiscent (0.5 x 23 cm), grey-brown, glabrous Seed: unknown Other: it starts as a shrub. The branchlets are glabrous and have pale brown lenticels.

Data sources FWTA, Leeuwenberg (1997)

Distribution Continent: Upper Guinea endemic Upper Guinea: Liberia (herbarium). According to Leeuwenberg (1997) only known from Liberia, but Hawthorne (pers. comm.) considers that it occurs in Ghana as well. Distribution type: continuous, very local, present in 2 30’ cells Forest type: secondary forest

Habitat Recorded only in disturbed places (secondary forests) at low altitudes (herbarium, Leeuwenberg 1997).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

A.p.

3

100 (2)

0

100

33

0

0

0

67

0

100

0

67

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

115

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Page 116

Albertisia cordifolia

(Mangenot & Miège) F

Menispermaceae A

Distribution

Data sources

Continent: Upper Guinea endemic (De Koning 1983) Upper Guinea: Côte d’Ivoire Distribution type: continuous, very local, present in 1 30’ cell, distribution range is 13 km Forest type: rainforest, secondary forest

Troupin (1962), Guillaumet & Debray (1964), De Koning (1983), Hawthorne & Jongkind (2004)

Habitat It is commonly found in lowland secondary forests close to lakes (De Koning 1983).

Regeneration It has a hypogeal cryptocotylar reserve seedling type (De Koning 1983).

Phenology

Description

Dispersal: probably by animals Timing: flowering period from April to June; fruiting period from June to July and January to February (De Koning 1983)

Guild: np Life form: winding woody climber Max. height: 15 m (De Koning 1983) Max. diameter: 1 cm (De Koning 1983) Leaf: alternate, simple, ovate to elliptic, mesophyll (7-17 x 9-25 cm), entire, coriaceous to herbaceous; petiole 3-12 cm long, hairy; no stipules Inflorescence: axillary, male inflorescence unbranched (cyme), pubescent, 1-3 flowers; female inflorescence unbranched (cyme) Flower: unisexual; male flowers very small, 9 sepals (0.1-0.6 cm), 6 petals; female flowers smaller than the male flowers Fruit: fleshy (drupe), ovoid (3 x 3 cm), yellow to orange, pubescent; 1 seed Seed: ellipsoid, very large (3 x 2 x 1 cm), brown Other: a slender woody climber that occurs in the form of a shrub when young.

spp

116

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

A.c.

14

9 (11)

0

100

0

0

7

93

0

21

79

0

0

21

79

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 108-118

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Page 117

Albertisia cuneata

(Keay) Forman

Menispermaceae A

Description

Phenology

Guild: sb Life form: winding woody climber Max. height: data unavailable Max. diameter: data unavailable Leaf: alternate, simple, ovate to elliptic, microphyll (2-5 x 5-9 cm), entire, with 6-8 lateral nerves, glabrous; no stipules Inflorescence: dioecious, axillary, unbranched, 1-3 flowers in cyme Flower: small; corolla yellowish Fruit: unknown Seed: unknown Other: a slender, glabrous, woody climber.

Dispersal: probably by animals

Data sources Troupin (1962), Hall & Swaine (1981)

Distribution Continent: Upper Guinea endemic Upper Guinea: Senegal, Côte d’Ivoire, Ghana. It was originally described for Ghana, but has been subsequently found in Senegal (Hall & Swaine 1981) and Côte d’Ivoire (herbarium). Distribution type: present in 4 30’ cells Forest type: In Ghana, only found in the South Marginal dry forest, where it can be locally abundant (Hall & Swaine 1981). It has also been recorded in dry secondary forests (herbarium).

Habitat It occurs in the understorey of dry forests (herbarium).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

A.c.

4

0 (4)

0

25

0

75

25

0

0

0

0

100

0

0

50

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

117

22466 layout page 108-118

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Page 118

Albertisia ferruginea

(Diels) Forman

Menispermaceae A

Description

Data sources

Guild: u Life form: winding woody climber Max. height: data unavailable Max. diameter: data unavailable Leaf: alternate, simple, ovate to lanceolate, notophyll (3-7 x 9-18 cm), entire, herbaceous; petiole (and branchlets) covered with white to brownish hairs; no stipules Inflorescence: axillary, cymose Flower: data unavailable Fruit: fleshy, sub-globose (2-2.5 cm in diameter), orange, with long white hairs; 1 seed Seed: large (1.8 x 1.3 x 0.8 cm)

Troupin (1962), Hawthorne & Jongkind (2004)

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia Distribution type: continuous, local, present in 4 30’ cells, distribution range is 274 km Forest type: rainforest, secondary forest

Habitat Data unavailable.

Phenology Dispersal: probably by animals

spp

118

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

A.f.

7

100 (2)

0

43

0

0

0

0

100

0

100

0

100

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 119-128

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Page 119

Albertisia mangenotii

(Guillaumet & M.M.Debray) Forman

Menispermaceae A

Description

Phenology

Data sources

Guild: np Life form: shrub Max. height: 1 m (Guillaumet & Debray 1964) Max. diameter: data unavailable Leaf: alternate, simple, ovate, notophyll (4-8 x 5-12 cm), entire, herbaceous, long palmate ascending nerves, pubescent; petiole 4-8 cm long; no stipules Inflorescence: male inflorescence axillary, unbranched, with 3-5 flowers; female inflorescence axillary, unbranched, with 1-3 flowers Flower: small Fruit: fleshy, 6 ovoid fruit parts (2 x 3 cm), yellow to orange, velutinous; 1 seed per fruit part Seed: medium-sized (1 x 0.8 cm) Other: the branchlets and petioles have erect, yellow hairs.

Dispersal: probably by animals

Guillaumet & Debray (1964), Hawthorne & Jongkind (2004)

Distribution Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire Distribution type: present in 2 30’ cells. Guillaumet & Debray (1964) mention at least 8 collections for Côte d’Ivoire. Forest type: wet evergreen forest, secondary forest

Habitat The species is found in the forest understorey, as well as along roadsides, abandoned plantations, and secondary regrowth (Guillaumet & Debray 1964). On sandy soils (herbarium).

spp

A.m All

n

3 46333

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

100 (1)

0

100

0

0

33

66

0

0

33

67

33

0

0

37

4

39

7

37

29

24

10

6

69

25

36

13

39

119

22466 layout page 119-128

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Page 120

Albertisia scandens

(Mangenot & Miege) Forman

Menispermaceae A

Habitat Species occurrence increases in places where rainfall reaches 2000-2500 mm/yr (Chi2 test). Found in a wide variety of forest types. Sometimes in mature forest but particularly common in open and disturbed places (e.g. along roadsides, forest edges, secondary forests) (herbarium, De Koning 1983). Sometimes close to rivers and lakes (herbarium, Chi2 test). It grows on sandy, clayish, and lateritic soils.

Description

Regeneration

Guild: np Life form: medium-sized, winding woody climber Max. height: 30 m long (herbarium) Max. diameter: data unavailable Leaf: alternate, simple, ovate to elliptic, mesophyll (4-10 x 6-18 cm), entire, herbaceous, young leaves covered with white hairs; petiole 1-3 cm long Inflorescence: dioecious, male inflorescence axillary, cyme (peduncle 1.5-4 cm long); female inflorescence reduced to one solitary flower (peduncle 0.8-1 cm long) Flower: small; calyx with 9 sepals; corolla yellowish, 6 petals Fruit: fleshy (drupe) (1.8 x 3 cm), orange, beaked and bristly with almost white hairs; 1 seed Seed: ellipsoid, large (2 x 1 x 0.9 cm) Other: a slender, hairy, woody climber. The bark is covered with lenticels. The branchlets and petioles are covered with white to brownish, stiff hairs.

It has a cryptocotylar hypogeal reserve seedling type (De Koning 1983).

Phenology Dispersal: probably by animals Timing: flowering period from June to July; fruiting period throughout the year (De Koning 1983)

Distribution Continent: Upper Guinea endemic Upper Guinea: Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 10 30’ cells, distribution range is 800 km Forest type: In Ghana, it is found in a wide variety

spp

120

n

Open (n)

of forests: evergreen (wet, moist, upland), semideciduous (moist, dry), dry south marginal (Hall & Swaine 1981). It is commonly recorded in secondary forests (herbarium, De Koning 1983), and occasionally in coastal savannas (herbarium).

Data sources Troupin (1962), Hall & Swaine (1981), De Koning (1983)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

A.s.

38

35 (23)

0

71

13

0

32

68

0

37

55

8

8

29

47

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 119-128

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Page 121

Alsodeiopsis chippii

Hutch.

Icacinaceae Description

A

Phenology

Guild: sb Life form: shrub or small tree Max. height: 1.8 m (herbarium) Max. diameter: data unavailable Leaf: alternate, simple, oblong-elliptic to oblongoblanceolate, notophyll (3-5 cm x 7-13 cm), herbaceous; petiole hairy (0.3-0.4 cm long) Inflorescence: on the leafy shoots, branched, hairy Flower: small; yellow Fruit: fleshy (drupe), ellipsoid (0.9 cm long), red, pubescent; 1 seed Seed: data unavailable

Distribution

Data sources

Continent: Upper Guinea endemic Upper Guinea: Ghana Distribution type: continuous, very local, present in 2 30’ cells, distribution range is 60 km. Restricted to southwest Ghana. Forest type: wet evergreen forest

FWTA

Habitat Found in the shaded understorey of dense forests (herbarium).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

L

M

H

L

M

H

17

83

0

33

67

0

0

33

67

24

10

6

69

25

36

13

39

A.c.

6

0 (6)

0

33

0

0

All

46333

37

4

39

7

37

29

Soil CMK

Soil WHC

121

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Page 122

Amorphophallus baumannii

N.E.Br.

Araceae A

inside; sterile part spadix from purple-brown to lemon-yellow; stem smooth; green with large purple dots Fruit: green Seed: data unavailable Other: it has a large and flat tuber.

Distribution Continent: Burkina Faso, Benin, Nigeria (Hetterscheid) Upper Guinea: Sierra Leone (Hetterscheid), Côte d’Ivoire, Ghana, Togo (herbarium) Distribution type: present in 4 30’ cells Forest type: data unavailable

Habitat Data unavailable.

Phenology

Description Guild: u Life form: large tuberous perennial herb Max. height: 1.8 m (herbarium) Leaf: compound, leaflets obovate, notophyll (2-6 x 5-16 cm), entire, petiole green with purple dots; only one leaf, from the soil surface like an umbrella Inflorescence: spike of flowers on a swollen fleshy axis (spadix) up to 27 cm long Flower: spathe pale green with some red near base

spp

122

n

Open (n)

Data sources Hetterscheid

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

A.b.

4

- (0)

0

0

0

100

0

0

0

0

75

25

50

25

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 119-128

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Page 123

Ancistrocladus pachyrrachis

Airy Shaw

Ancistrocladaceae A

Description

Data sources

Guild: u Life form: woody climber Max. height: 3 m long (FWTA) Max. diameter: data unavailable Leaf: alternate, simple, mesophyll (3-7 x 10-30 cm), entire, coriaceous; sessile Inflorescence: terminal, branched (panicle, up to 30 cm long) Flower: medium-sized Fruit: dry indehiscent, winged; 1 seed Seed: subglobose Other: it starts as a scandent shrub, climbing with hooks which are found especially on the short lateral shoots (spur shoots). These spur shoots have one bundle of crowded leaves.

FWTA, Hawthorne & Jongkind (2004)

Distribution Continent: Upper Guinea endemic Upper Guinea: Liberia Distribution type: continuous, very local, present in 1 30’ cell Forest type: data unavailable

Habitat Data unavailable.

Phenology

spp

n

Open (n)

A.p.

2

- (0)

All

46333

37

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

L

M

H

L

M

H

100

100

0

0

0

100

0

100

0

0

0

0

39

7

37

29

24

10

6

69

25

36

13

39

0 4

Soil CMK

Soil WHC

123

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Page 124

Anisophyllea laurina

R.Br. ex Sabine

Anisophylleaceae A

Seed: large (1.5 x 1 cm) Other: it has a straight bole and no buttresses. The bark is grey to brown, hard and smooth.

Dispersal: probably by animals Timing: flowering period: mainly in November to December but can be prolonged to June (Savill & Fox 1967); fruiting period: April to May (Savill & Fox 1967)

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea-Bissau, Guinea, Sierra Leone Distribution type: Upper Guinea disjunct, regional, present in 7 30’ cells, distribution range is 538 km Forest type: montane forest, rainforest, coastal savanna, secondary forest

Habitat In Guinea Bissau it is often reported in dense forests. In Sierra Leone it extends across the coastal savanna where it can be locally abundant and occasionally dominant. It is a rapid coloniser of Gmelina arborea plantations as an understorey tree, and can also colonise lateritic subsoil after gravel is removed for road maintenance (Savill & Fox 1967). In general, it is usually found in disturbed forests, sometimes close to lakes, and occasionally on sandy soils near the coast (herbarium).

Description

124

n

Open (n)

The fruits are edible. Poles are used to build houses and are said to be termite proof. Leaves are used as a mouth rinse for toothache and are said to have medicinal properties (Savill & Fox 1967).

Data sources

Regeneration

Guild: pi Life form: medium-sized tree Max. height: 24 m (herbarium) Max. diameter: data unavailable Leaf: alternate, simple, elliptic-ovate lanceolate, notophyll (3-7 x 7-15 cm), entire, coriaceous, glabrous, 1-2 pairs of main longitudinal nerves, tertiary nerves markedly parallel Inflorescence: axillary, not branched Flower: small; greenish; calyx 4-lobed, 4 petals; sweet scented Fruit: fleshy, oblong (3 x 5 cm), yellow or reddish; 1 seed

spp

Uses

FWTA, Aubréville (1950), Savill & Fox (1967)

It is locally abundant in secondary regrowth (herbarium).

Phenology Deciduousness: most leaves are shed in November, but some always remain. New foliage is produced by April, but the trees in closed forests are scarcely deciduous and new leaves appear with the flowers (Savill & Fox 1967)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

A.l.

10

17

(6) 10

0

60

0

30

40

30

0

40

60

20

50

H 10

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 119-128

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Page 125

Anisophyllea meniaudii

Aubrév. & Pellegr.

Anisophylleaceae Description

A

Phenology

Guild: np Life form: medium-sized tree Max. height: 21 m (FWTA) Max. diameter: 29 cm (herbarium) Leaf: alternate, simple, elliptic to lanceolate, notophyll (2.2-6 x 6-13 cm), coriaceous, purple red hairs on young leaves and twigs, tertiary nerves not markedly parallel Inflorescence: axillary, not branched (spike) Flower: small; greenish Fruit: fleshy, subglobular (2 cm in diameter), yellow to orange Seed: medium-sized (1 x 0.6 cm) Other: the bole is straight. The slash is brown and stinks when sawn. Saplings and flushing twigs have conspicuous prominent ‘stipules’ (actually a much reduced leaf).

Data sources FWTA, Aubréville (1959), Savill & Fox (1967), Keay (1989), Hawthorne (1995a), Hawthorne & Jongkind (2004)

Distribution Continent: Guineo-Congolian (Keay 1989), Nigeria, Cameroon, Democratic Republic of Congo (herbarium) Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana (herbarium) Distribution type: continental disjunct, in Upper Guinea present in 11 30’ cells Forest type: evergreen forest, secondary forest (Hawthorne 1995, herbarium)

Habitat It occurs in shaded places and is occasionally reported on steep rocky hills (herbarium).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

A.m

19

14 (14)

0

37

0

0

58

21

21

0

68

32

37

0

63

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

125

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Page 126

Anisotes guineensis

Lindau

Acanthaceae A

Description

Data sources

Guild: u Life form: shrub or small tree Max. height: 4 m (FWTA) Max. diameter: data unavailable Leaf: opposite, simple, oblong to oblanceolate, microphyll (2-4 x 5-8 cm), entire, coriaceous Inflorescence: axillary, solitary Flower: large (corolla up to 4 cm long); corolla dark red, 2 lipped Fruit: dry dehiscent Seed: data unavailable

FWTA

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea Distribution type: continuous, local, present in 3 30’ cells, distribution range is 174 km Forest type: wet evergreen forest, gallery forest

Habitat It is found by streams (FWTA).

Phenology

spp

126

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

A.g.

5

- (0)

20

20

0

0

40

40

20

0

20

80

0

0

20

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 119-128

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Page 127

Anthocleista nobilis

G.Don

Gentianaceae A

Description Guild: pi Life form: medium-sized tree Max. height: 30 m (De Koning 1983) Max. diameter: 80 cm (herbarium) Leaf: opposite, simple, oblanceolate, macrophyll (418 x 6-64 cm), leaves of saplings up to 2 m long, coriaceous Inflorescence: terminal, branched Flower: medium-sized (corolla 3.5 cm long); calyx and corolla creamy white Fruit: fleshy (berry), globose (3 x 4 cm), dark green Seed: ovate-orbicular, medium-sized (0.22 x 0.16 cm), brown Other: only ramified close to the terminal point of the tree. Stilt roots develop at the base of some trees. The adventitious stilt roots enter the ground before growing horizontally, 10-20 cm below the surface. In swampy soils, “peg roots” less than a millimetre in diameter grow upwards and protrude a few centimetres above the ground. The branches have short, paired spines. The species is most likely to be confused with A. vogelii, which also has sharpspiny twigs, but has less slender leaves, and leaves which do not dry black. A. nobilis’ corolla tubes are 2-3 times as long as the lobes, whereas for A. vogelii this is 0.9-1.5 times as long. The wood density is 0.56 g/cm3.

Phenology Dispersal: by animals (bats)

Habitat Species occurrence increases significantly in zones with rainfall higher than 2000 mm/yr to reach an optimum around 2500 mm/yr (logistic regression analysis, Chi2 test). It can be locally abundant. Usually, in moist to swampy places (herbarium, Kasparek 2000). Very often in open places such as gaps and roadsides, and in abandoned and degraded forests (herbarium, Kasparek 2000). On sandy, sandy clay, loamy, and rocky soils (herbarium).

Distribution Continent: Upper Guinea endemic Upper Guinea: Senegal, Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 22 30’ cells, distribution range is 1815 km Forest type: rainforest, secondary forest, swamp forest, gallery forest

Data sources

Regeneration

FWTA, Leeuwenberg (1961), De la Mensbruge (1966), Jeník (1971), Hawthorne (1981, 1995a), De Koning (1983), Kasparek (2000)

It has a phanerocotylar epigeal foliaceous seedling type (cf. de la Mensbruge 1966).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

A.n.

52

70 (47)

15

46

12

12

33

36

18

15

60

25

33

17

H 25

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

127

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Page 128

Anthonotha explicans

(Baill.) J.Léonard

Leguminosae-Caes. A

Description

Phenology

Data sources

Guild: u Life form: large tree Max. height: 33 m (herbarium) Max. diameter: 60 cm (herbarium) Leaf: paripinnately compound, 4-6 leaflets, ovate, notophyll (2.5-5 x 4.5-12 cm), entire Inflorescence: branched (pendulous, up to 50 cm long), glabrous Flower: pale yellow to orange; strongly fragrant like coconut; flowering in the early morning, dropping at noon; glabrous; small stipules Fruit: dry dehiscent, brown, pubescent, with conspicuous raised wavy lines (longer than 8 cm); 1 seed Seed: very large Other: the height of the species varies from a shrub in savannas to a large tree in moist forests. It has a brown crown due to the abundance of dense hairs on the leaves. The slash is pinkish brown.

FWTA, Voorhoeve (1965), Lock (1989), Hawthorne & Jongkind (2004)

Distribution Continent: Upper Guinea endemic Upper Guinea: Senegal, Guinea, Sierra Leone, Liberia, Côte d’Ivoire Distribution type: continuous, widespread, present in 8 30’ cells Forest type: moist evergreen forest, savannawoodland

Habitat It grows in dry places (herbarium).

spp

128

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

A.e.

9

60 (5)

11

67

22

11

11

11

66

0

56

44

33

0

H 44

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 129

Anthonotha vignei

Hoyle

Leguminosae-Caes. A

Description

Phenology

Guild: np Life form: medium-sized tree Max. height: 30 m (herbarium) Max. diameter: 40 cm (herbarium) Leaf: paripinnately compound, ovate to elliptic, notophyll (3-5 x 7-18 cm), entire, herbaceous, redbrown pubescent beneath, rachis hairy Inflorescence: terminal, branched (up to 15 cm long) Flower: small; corolla white, 4 petals Fruit: dry dehiscent, oblong (5 x 11 cm), velvety brown with many transverse wrinkles; 1-2 seeds Seed: very large (4 x 3.5 x 1 cm) Other: the slash is red-brown and brittle.

Timing: flowering period from February to March; fruiting period from September to November (De Koning 1983)

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 22 30’ cells, distribution range is 1096 km, Red List species (vulnerable) Forest type: wet evergreen forest, moist evergreen forest, coastal forest, riverine forest, secondary forest

Data sources FWTA, De Koning (1983), Lock (1989), Hawthorne (1990, 1995a), IUCN Red List (2000), Hawthorne & Jongkind (2004)

Habitat Species occurrence increases with rainfall to reach a very wide optimum range between 1500-3100 mm/yr (logistic regression analysis, Chi2 test). Generally found in the lowlands and close to the coast (logistic regression analysis, Chi2 test). Most often along riversides and riverbanks (Chi2 test, Hawthorne 1995). In both mature forest and disturbed places (e.g. roadsides, exploited forests, open forests). On gravel, white sandy, and clayishsandy soils (herbarium).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

H

A.v.

56

50 (32)

5

55

18

0

34

39

26

7

82

11

39

7

50

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

129

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Anubias afzelii

Page 130

Schott.

Araceae A

Data sources

Flower: scape light green, somewhat tinged with pink; spatha light green, glossy; spadix greenishcreamy Fruit: fleshy, depressed-globose; many seeds Seed: irregularly ovoid, small (0.2 x 0.1 cm), dark brown

FWTA, Crusio (1979)

Distribution Continent: Upper Guinea endemic Upper Guinea: Senegal, Guinea, Sierra Leone, Ghana (doubtful record) (herbarium) Distribution type: continuous, regional, present in 5 30’ cells, distribution range is 380 km Forest type: data unavailable

Habitat Usually found in very moist places (e.g. riverbank, edge of streams, and swamps) (herbarium). Growing on wet, shady places, sometimes completely submerged (Crusio 1979).

Phenology Timing: flowering period from April to July; fruiting period from April to September (Crusio 1979)

Description Guild: u Life form: perennial rhizomatous herb Max. height: data unavailable Leaf: alternate, simple, elliptic, macrophyll (4-16 x 12-39 cm), entire, glabrous; petiole up to 20 cm long, pulvinate at the apex, glabrous Inflorescence: spike of flowers on a swollen fleshy axis (spadix, up to 10 cm long)

?

spp

130

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

A.a.

5

0 (1)

20

20

20

0

0

40

60

0

40

60

40

0

20

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 131

Aphanocalyx microphyllus

(Harms) Wieringa ssp.

compactus

(Hutch. ex Lane-Poole) Wieringa

Leguminosae-Caes. A

Description

Habitat

Guild: sb Life form: large tree Max. height: 42 m (Voorhoeve 1965) Max. diameter: 100 cm (Voorhoeve 1965) Leaf: alternate, paripinnately compound, 16-52 leaflets, rhombic, leptophyll (0.1-0.4 x 0.3-1.8 cm), entire, coriaceous, glabrous at both sides, basal leaflet with 0-3 glands; rachis puberulous, grooved above, stipules Inflorescence: axillary or terminal, unbranched Flower: small; male or bisexual; corolla yellow Fruit: dry dehiscent (pod), flat, oblong to obovate (2 x 5 cm), woody, wine-red when immature, later brown; 1-2 seeds Seed: large (1.3 x 1 x 0.2 cm) Other: the base of large trees sometimes has buttresses. The slash is pink.

The subspecies regenerates in the forest. As some secondary forests might consist completely of this subspecies, the single dominant forests might be relics of shifting cultivation (Voorhoeve 1965). The subspecies is found in areas with an annual rainfall > 2000 mm, and its abundance increases with rainfall (Bongers et al. 1999). In drier areas it only occurs in riverine forest (Wieringa 1999).

Regeneration Trees of 8 cm dbh are already flowering (Voorhoeve 1965). It has a phanerocotylar epigeal reserve seedling type. Regeneration can be abundant; the forest floor can be covered by saplings of 1 m height (Wieringa 1999). In single dominant forests of the subspecies, it is also present as mediumsized trees, unlike other species which are known to form single dominant forests (e.g. Monopetalanthus compactus, Gilbertiodendron preussii) (Voorhoeve 1965). This feature facilitates the design of silvicultural selection system (Voorhoeve 1965).

Distribution Continent: Upper Guinea endemic (Wieringa 1999) Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire (Wieringa 1999) Distribution type: continuous, regional, present in 12 30’ cells. It occurs over a narrow band, 100 km inland, only reaching the coast at Tabou (which belongs to a doubtful record). Outside this band it exists in an isolated population on Mount Nimba (Wieringa 1999). The subspecies is gregarious, and constitutes of single-species dominant forest of vast extension in rolling areas (Voorhoeve 1965). The other subspecies, A. microphyllus spp. microphyllus is only found in central Africa (Wieringa 1999). Forest type: wet evergreen forest, gallery forest

Phenology Deciduousness: evergreen Timing: flowering period from February to April, just before the rainy season; mature pods are present in October (Wieringa 1999).

Data sources Voorhoeve (1965), Bongers et al. (1999), Wieringa (1999)

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

A.m

21

0 (6)

0

33

5

0

5

48

48

0

71

29

71

0

10

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

131

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Page 132

Aphanocalyx pteridophyllus

(Harms) Wieringa

Leguminosae-Caes. A

Phenology

Fruit: dry dehiscent (pod), oblong-obovate (5 x 3 cm), green to reddish, brown hairy; 1-2 seeds Seed: large (1.6 x 1.4 x 0.4 cm) Other: the crown is broad but irregular due to the early branching trunk. Ants build their nests in between the persistent stipules.

Timing: flowering period in July and August, during the rainy season (Wieringa 1999)

Distribution Continent: Upper Guinea endemic (Wieringa 1999) Upper Guinea: Sierra Leone, Liberia (Wieringa 1999) Distribution type: continuous, regional, present in 10 30’ cells, distribution range is 447 km Forest type: wet evergreen forest, gallery forest

Habitat

Data sources

It occurs in the lowlands, near the coast (Wieringa 1999). The species is found near rivers, creeks, and along rivers with brackish water (Wieringa 1999), but sometimes in upland areas, where it often grows in small groups (Voorhoeve 1965).

Voorhoeve (1965), Wieringa (1999)

Regeneration Epigeal germination with cotyledons clasping the base of the first leaf pair (Wieringa 1999).

Description Guild: u Life form: large tree Max. height: 45 m (herbarium) Max. diameter: 35 cm (herbarium) Leaf: alternate, paripinnately compound, nanophyll (0.3-1.3 x 0.8-4.3 cm), entire, herbaceous; primary nerve ending in a conspicuous gland Inflorescence: axillary, in the axils of young shoots, unbranched Flower: small; corolla white

spp

132

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

A.p.

20

0 (2)

0

55

20

0

0

0

100

0

85

15

70

5

20

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 133

Apodiscus chevalieri

Hutch.

Euphorbiaceae A

Description

Data sources

Guild: u Life form: small tree Max. height: 10 m (herbarium) Max. diameter: data unavailable Leaf: alternate, simple, oblong to elliptic, mesophyll (3-8 x 8-17 cm), entire, coriaceous, glabrous Inflorescence: axillary, not branched (spike), hairy Flower: small; white Fruit: pendulous, 3-5 lobed, green Seed: data unavailable

FWTA, Hawthorne & Jongkind (2004)

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea, Liberia, Ghana Distribution type: continuous, widespread, present in 4 30’ cells, distribution range is 1196 km Forest type: wet evergreen forest

Habitat It is commonly observed on riverbanks on sandy soils (herbarium).

Phenology

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

A.c.

7

0 (3)

0

86

0

0

0

29

71

0

86

14

71

0

14

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

133

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Page 134

Argocoffeopsis afzelii

(Hiern) Robbr.

Rubiaceae A

Regeneration It has a phanerocotylar epigeal reserve seedling type (cf. De Koning 1983)

Phenology

spp

134

Description

Distribution

Guild: sb Life form: woody climber, climbing with hooked side-branches Max. height: 25 m long (De Koning 1983) Max. diameter: data unavailable Leaf: opposite, simple, elliptic, microphyll (2.5-4 x 4-10 cm), entire, coriaceous, glabrous; interpetiolar stipules Inflorescence: axillary, solitary Flower: large; calyx dark green; corolla white, tubeshaped, with the lobes having about the equal length as the tube, stamens exerted; fragrant Fruit: fleshy (drupe), globose (0.8 cm in diameter), black; 2 seeds Seed: hemispherical, medium-sized (0.6 x 0.4 x 0.2 cm), white Other: sometimes in the form of a shrub when young. It climbs by means of horizontal or recurved lateral branches. The bark is papery, and peels off characteristically.

Continent: Upper Guinea endemic (Robbrecht 1981a) Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana (herbarium, Robbrecht 1981a) Distribution type: continuous, widespread, in Upper Guinea present in 22 30’ cells. Rare in Ghana (Hall & Swaine 1981). Forest type: wet evergreen forest, moist evergreen forest, dry semi-deciduous forest, secondary forest, thickets

n

Open (n)

Deciduousness: evergreen (Robbrecht 1981a) Timing: flowering period from September to February; fruiting period from November to April (De Koning 1983)

Habitat Although it can be found in dry and wet forests, species occurrence increases sharply with rainfall higher than 2000 mm/yr (logistic regression analysis, Chi2 test). It is usually found in the shaded forest understorey (De Koning 1983).

Data sources Hall & Swaine (1981), Robbrecht (1981a), De Koning (1983)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

A.a.

48

54 (28)

2

65

2

4

16

44

33

10

79

10

46

6

H 40

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 135

Argocoffeopsis lemblinii

(A.Chev.) Robbr.

Rubiaceae A

Description

Phenology

Guild: u Life form: shrub Max. height: 1 m (Hawthorne & Jongkind 2004) Max. diameter: data unavailable Leaf: opposite, simple, oblong to elliptic, microphyll (1.5-5.5 x 3-10 cm), hairy on midrib; interpetiolar stipules Inflorescence: axillary, solitary, on short shoots Flower: small (corolla tube 0.6 cm); calyx lobes rounded; corolla white, corolla-tube as long as lobes Fruit: fleshy; 2 seeds Seed: hemispherical Other: the bark is papery, peeling off characteristically.

Deciduousness: evergreen Timing: only one record is known, and this specimen flowered in January (FWTA).

Data sources FWTA, Robbrecht (1981a), Hawthorne & Jongkind (2004)

Distribution Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire (Robbrecht 1981a) Distribution type: continuous, very local, present in 1 30’ cell. Very rare; only known from the type specimen Chevalier 17180 from Valleé de l’Agniéby, Voquié. Forest type: moist evergreen forest

Habitat The single record of this species was found in the forest.

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

A.l.

1

- (0)

0

0

0

0

100

0

0

0

0

100

0

0

100

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

135

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Page 136

Asystasia scandens

(Lindl.) Hook.

Acanthaceae A

Phenology

Uses Sometimes it is cultivated as an ornamental plant (FWTA).

Data sources FWTA

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire Distribution type: continuous, widespread, present in 14 30’ cells, distribution range is 884 km Forest type: data unavailable

Description Guild: u Life history: woody climber Max. height: 2 m long (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, oblong-obovate to elliptic, notophyll (3.5-5.5 x 8-16 cm), coriaceous, glabrous Inflorescence: terminal, unbranched Flower: large (corolla 5 cm long); calyx pale green; corolla bell-shaped, white with purplish tinges Fruit: dry dehiscent Seed: data unavailable

spp

136

n

Open (n)

Habitat Found in forest or along roads. Sometimes near streams or riverbanks on sandy soils (herbarium). Species occurrence increases very significantly with rainfall (logistic regression analysis) and is highest in places where rainfall is higher than 2500 mm/yr (Chi2 test).

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

A.s.

23

75 (4)

0

35

17

4

0

34

60

0

61

39

43

9

H 13

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 137

Baissea zygodioides

(K.Schum.) Stapf

Apocynaceae Description

Phenology

Guild: np Life form: large woody winding climber Max. height: 20 m long (herbarium) Max. diameter: data unavailable Leaf: subopposite, simple, ovate to oblong, microphyll (0.5-2.5 x 0.5-7 cm), entire, herbaceous to coriaceous, sparsely pubescent to glabrous on both sides Inflorescence: axillary or terminal, few to many cymes Flower: medium-sized (1.4 cm long); corolla white to yellowish, funnel-shaped, pubescent outside, glabrous inside; fragrant Fruit: dry dehiscent (generally 1 x 24 cm, but old fruit valves can be up to 1 x 53 cm), brown, densely pubescent Seed: large (1.7 x 0.24 x 0.13 cm), with parachute of hairs Other: the bark is brownish grey and smooth and the slash exudes a white latex. With tendrils, pubescent.

Timing: flowering period from March to September; fruiting period from January to October (Van Dilst 1995)

B

Data sources FWTA, Hall & Swaine (1981), De Koning (1983), Van Dilst (1995)

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 32 30’ cells, distribution range is 1960 km Forest type: secondary forest, bush, thicket, savanna, riverine forest, gallery forest, swamp forest (Van Dilst 1995). In Ghana, mostly found in dry semi-deciduous forests but also in the dry South Marginal forests (Hall & Swaine 1981).

Habitat Often found in open places such as forest edges, roadsides, or in secondary forests. On sandy to clayish soils (herbarium).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

B.z.

59

60 (15)

7

32

5

40

23

26

10

3

53

44

36

8

H 27

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

137

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Baphia spathacea

Page 138

Hook.f. ssp.

spathacea

Leguminosae-Pap. Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia Distribution type: continuous, regional, present in 6 30’ cells, distribution range is 472 km Forest type: secondary forest, coastal forest, coastal savanna Other: the subspecies is Upper Guinea endemic, and has a strong disjunct distribution from the other subspecies B. spathacea polyantha, which is found in Cameroon, Gabon and the Democratic Republic of Congo.

B

Habitat It is often found in secondary vegetation, at the edge of forests, or along streams or swamps. It occurs on laterite and sandy soils (herbarium).

Phenology

Description Guild: pi Life form: woody climber Max. height: 5 m (Hawthorne & Jongkind 2004) Max. diameter: data unavailable Leaf: simple, oblong to elliptic, notophyll (4-6 x 6.5-13 cm), entire, coriaceous, hairy on the midrib Inflorescence: axillary, branched (panicle), brown tomentose Flower: small; calyx rusty brown; corolla yellow to whitish Fruit: dry dehiscent (pod), oblong (2 x 8 cm), woody, dark brown; 2-3 seeds Seed: large (1.3 x 1.2 cm), brown Other: the branchlets are brown pubescent.

spp

138

n

Open (n)

Data sources FWTA, Soladoye (1984), Lock (1989), Hawthorne & Jongkind (2004)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

B.s.

28

75 (16)

0

57

46

0

0

0

85

0

100

0

50

0

4

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 139-148

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Page 139

Begonia cavallyensis

A.Chev.

Begoniaceae Description Guild: u Life form: perennial epiphytic herb Max. height: 0.9 m (De Wilde 1983) Leaf: alternate, simple, ovate, mesophyll (2-10 x 416 cm), entire, coriaceous, scarcely hairy; red petiole; large stipules (1.4 x 3.5 cm), soon falling off Inflorescence: monoecious, axillary; male inflorescence branched, 4-15 flowers; female inflorescence unbranched, 2-3 flowers Flower: corolla white-pink Fruit: fleshy, dehiscent, fusiform (0.4 x 3.5 cm), red; many seeds, surrounded with aril-like structure Seed: ovoid, small (0.13 x 0.07 cm), brown Other: it has an erect succulent stem with scattered conspicuous white trichomes. The margin has a contrasting band of dark red of approx. 0.5 cm wide. The nerves are wine-red.

B

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 15 30’ cells, distribution range is 1346 km Forest type: (sub) montane rainforest

Phenology Deciduousness: most leaves are shed in the dry season Dispersal: probably by animals

Habitat Species occurrence increases significantly with rainfall higher than 2000 mm/yr (logistic regression analysis, Chi2 test), and is most often found at higher altitudes (Chi2 test). It grows on trees and fallen logs, in shade but sometimes also at forest edges (herbarium). Found at elevations between 500 and 1500 m (De Wilde 1983).

Data sources Johansson (1974), De Wilde (1983)

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

H

B.c.

25

40 (5)

32

36

0

0

24

48

28

0

72

28

48

0

32

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

139

22466 layout page 139-148

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Page 140

Begonia fusicarpa

Irmsch.

Begoniaceae Description Guild: u Life form: perennial epiphytic herb Max. height: data unavailable Max. diameter: data unavailable Leaf: alternate, simple, narrowly obovate to narrowly elliptic, mesophyll (4-6.5 x 13-21 cm), mostly entire with teeth at the upper half; petiole up to 0.6 cm long Inflorescence: monoecious, male inflorescence in the axils of terminal leaves, branched (cymose), female inflorescence lower down the stem Flower: small, corolla pink, 4 petals Fruit: fleshy (2.3 x 4.8 cm), erect; many seeds Seed: small (0.13 x 0.06 cm)

B

Distribution Continent: Upper Guinea endemic Upper Guinea: Liberia Distribution type: continuous, very local, present in 1 30’ cell. It is an extremely rare species. Only known from the type specimen J.T. Baldwin 11417 from Sino County, Kulo. Forest type: data unavailable

Habitat Confined to lowlands in Liberia under 500 m altitude (De Wilde 1983).

Phenology Data unavailable.

Data sources De Wilde (1983)

spp

140

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

B.f.

1

- (0)

0

0

0

0

0

0

100

0

100

0

100

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 139-148

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Page 141

Begonia hirsutula

Hook.f.

Begoniaceae Description Guild: sb Life form: perennial rhizomatous herb Max. height: 0.3 m Leaf: rosette, simple, ovate, mesophyll (1-12 x 419 cm), entire to dentate, lower surface green to wine red, nerves hairy; petioles purplish, red and hairy Inflorescence: axillary, branched with unisexual flowers, 1-3 terminal female flowers Flower: small, yellow with red veins Fruit: fleshy, obovate (0.8 x 1.5 cm), reddish brown, winged Seed: small (< 0.04 cm) Other: a hairy plant.

B

Distribution

Phenology

Continent: Nigeria, Cameroon, Equatorial Guinea, Gabon, Congo (Brazzaville), Democratic Republic of Congo (Sosef 1994) Upper Guinea: Ghana Distribution type: continental disjunct with a small population in Upper Guinea, present in 3 30’ cells in Upper Guinea Forest type: montane forest, rainforest, swamp forest

Dispersal: barochore

Habitat It is found in shaded places in undisturbed forest. It occurs usually in damp sites and along streams. Usually found at up to 1000 m altitude. Soils can be clayish, sandy, or rocky (Sosef 1994).

Data sources Sosef (1994)

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

B.h.

3

- (0)

0

33

0

0

67

33

0

0

67

33

0

0

100

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

141

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Page 142

Begonia mildbraedii

Gilg

Begoniaceae

B

Description

Phenology

Guild: sb Life form: rhizomatous perennial herb Max. height: 0.3 m Leaf: rosette, simple, ovate, notophyll (1.5-10 x 215 cm), entire to undulate, succulent to fleshy; petiole hairy red brown, margin red, lower leaf surface pale green to reddish Inflorescence: axillary, branched Flower: small male and female flowers, red outside, and yellow with red lines inside Fruit: dry, obovate (0.8 x 1.1 cm), green, 3-4-winged Seed: small (<0.04 cm)

Dispersal: barochore

Data sources Sosef (1994)

Distribution Continent: Cameroon, Gabon, Congo (Brazzaville), Democratic Republic of Congo (Sosef 1994) Upper Guinea: Côte d’Ivoire, Ghana (herbarium, Sosef 1994) Distribution type: continental disjunct with a small population in Upper Guinea, present in 3 30’ cells in Upper Guinea Forest type: rainforest, riverine forest, secondary forest

Habitat Usually found in shaded places in mature forest, but sometimes also found in open vegetation. It occurs typically in damp habitats near streams, rivers, waterfalls, or swamps. The soils can be clayey, sandy, or rocky (Sosef 1994).

spp

142

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

B.m.

5

- (0)

0

60

0

0

0

100

0

20

40

40

20

20

20

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 139-148

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Page 143

Begonia prismatocarpa

Hook. ssp.

petraea

(A.Chev.) Sosef

Begoniaceae Description

Phenology

Guild: sb Life form: rhizomatous perennial herb Max. height: 0.1 m Max. diameter: data unavailable Leaf: alternate, simple, ovate and sometimes lobed, microphyll (0.6-3.5 x 1.5-4 cm), dentate, herbaceous Inflorescence: axillary, not branched, unisexual flowers Flower: small, orange to yellow Fruit: dry, elliptic (0.7 x 0.4 cm), dark brown, 3-4-winged Seed: small (<0.04 cm) Other: this is one of three subspecies. The plant is covered with minute glandular hairs.

Dispersal: barochore

B

Data sources

Distribution

Sosef (1994) Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire Distribution type: continuous, very local, present in only 1 30’ cell. This is one of three subspecies, and extremely rare; only known from the type specimen Chevalier 19596 found between Nékaougnié and Grabo. The other two subspecies are found in Equatorial Guinea and Cameroon (Sosef 1994). Forest type: rainforest

Habitat It occurs in very deeply shaded places of mature lowland forests. On granite rocks (Sosef 1994).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

B.p.

1

0 (1)

0

0

0

0

0

100

0

0

0

100

0

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

143

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Page 144

Begonia quadrialata

Warb. ssp.

nimbaensis

Sosef

Begoniaceae Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea, Liberia, Côte d’Ivoire Distribution type: continuous, very local, present in 3 30’ cells, distribution range is 46 km. Begonia quadrialata nimbaensis is one of three subspecies. This subspecies is strictly confined to Mount Nimba and its surroundings. The other subspecies occur in Upper Guinea and Central Africa (Sosef 1994). Forest type: montane forest, rainforest

B

Habitat It is found in half-shaded to shaded places at an altitude between 350 and 1600 m. It occurs near streams or on moist to relatively dry sites. On rocks (Sosef 1994).

Phenology Description

Dispersal: barochore

Guild: sb Life form: rhizomatous perennial herb Max. height: 0.3 m Max. diameter: data unavailable Leaf: alternate, simple, elliptic to ovate to circular, notophyll (1.2-11 x 1.7-12.5 cm), entire to lobed, succulent, upper surface pale green with dark green to purple brown zones around the main nerve and secondary nerves Inflorescence: axillary, not branched, unisexual flowers, 2-5 male flowers and 1-2 terminal female flowers Flower: medium-sized; yellow to orange Fruit: dry, obovate (1.5 x 1 cm), green-brown, 3-4-winged Seed: small (<0.04 cm) Other: the plant is covered with minute glandular hairs.

spp

144

n

Open (n)

Data sources Sosef (1994)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

B.q.

17

0 (3)

65

18

0

0

0

100

0

0

76

24

71

0

6

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 139-148

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Page 145

Beilschmiedia caudata

(Stapf ) A.Chev.

Lauraceae Description

Data sources

Guild: u Life form: tree Max. height: data unavailable Max. diameter: data unavailable Leaf: alternate, simple, long acuminate, mesophyll (5-9 x 17-25 cm), coriaceous; no stipules Inflorescence: branched (panicle) (10-18 cm long), hermaphrodite, hairy Flower: small (0.3 cm long); corolla violet Fruit: fleshy, pendulous; 1 seed Seed: data unavailable Other: the leaves, bark and stem have oil glands and are sweetly scented.

FWTA, De Koning (1983), Mabberley (1987), Hawthorne & Jongkind (2004)

B

Distribution Continent: Upper Guinea endemic Upper Guinea: Liberia Distribution type: continuous, very local, present in 3 30’ cells, distribution range is 92 km Forest type: wet evergreen forest

Habitat Data unavailable.

Phenology Deciduousness: evergreen

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

B.c.

6

- (0)

0

33

17

0

0

0

51

0

83

17

83

0

17

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

145

22466 layout page 139-148

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Page 146

Beilschmiedia chevalieri

Robyns & R.Wilczek

Lauraceae

B

Description

Data sources

Guild: u Life form: small to medium-sized tree Max. height: 15 m (herbarium) Leaf: alternate, simple, oblong-elliptic to elliptic, mesophyll (4-9 x 10-22 cm), coriaceous, glabrous; no stipules Inflorescence: branched (panicle), hairy Flower: small (0.3 cm long); hermaphrodite; calyx pink; corolla greenish yellow, hairy Fruit: fleshy, pendulous; 1 seed Seed: data unavailable Other: the leaves, stem and bark have oil glands and are sweetly scented.

FWTA, Mabberley (1987), Hawthorne & Jongkind (2004)

Distribution Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire, Liberia Distribution type: continuous, regional, present in 6 30’ cells, distribution range is 528 km Forest type: wet evergreen forest, forest remnant, secondary forest

Habitat Data unavailable

Phenology Deciduousness: evergreen

spp

146

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

B.c.

9

17 (6)

11

56

0

0

67

22

11

0

89

11

89

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 139-148

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Page 147

Berlinia occidentalis

Keay

Leguminosae-Caes. Description Guild: np Life form: large tree Max. height: 40 m (herbarium) Max. diameter: 100 cm (herbarium) Leaf: alternate, paripinnately compound, 10 leaflets, elliptic, mesophyll (4-7 x 9-22 cm), entire, coriaceous, fine hairs flat against the lower surface Inflorescence: terminal or axillary, unbranched (up to 40 cm long), large coriaceous green bracts Flower: large; calyx greenish; corolla white with green midrib; fragrant Fruit: dry dehiscent (pod) (up to 10 x 42 cm), black with a narrow wing along the upper edge at both sides Seed: very large (5 x 4 x 3 cm) Other: an unbuttressed, evergreen tree with rounded, dense crown. The wood density is 0.72 g/cm3.

1995a). However, the herbarium records for the whole Upper Guinea region indicate a wider habitat, with incidental observations close to water (e.g. rivers, streams) but not restricted to them. On sandy, sandy clay, and rocky soils (herbarium).

B

Regeneration It has a phanerocotylar epigeal reserve seedling type (cf. de la Mensbruge 1966).

Phenology Deciduousness: evergreen Dispersal: explosive Timing: flowering period from March to May; fruiting period from October to November (Taylor 1960)

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 21 30’ cells, distribution range is 1188 km, Red List species (vulnerable) Forest type: wet evergreen forest, coastal savanna, secondary forest

Habitat

Data sources

It is mostly found in areas with rainfall higher than 2500 mm/yr (Chi2 test), at lowlands close to the coast (logistic regression analysis, Chi2 test). In both mature forest and open places (e.g. along roads in coastal savanna, in villages, plantations). In Ghana, seems to be restricted to evergreen forest, where it is most typical of swampy places (Hawthorne

FWTA, Taylor (1960), De la Mensbruge (1966), Voorhoeve (1965), Lock (1989), Hawthorne (1995a), IUCN Red List (2000), Kasparek (2000)

spp

n

Open ( n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

H

B.o.

41

38 (16)

0

44

20

2

30

25

38

10

90

0

56

7

27

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

147

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Page 148

Berlinia tomentella

Keay

Leguminosae-Caes. Description

places (e.g. riversides or swamp edges). On sandy, sandy clay, rocky, and lateritic soils (herbarium).

Guild: np Life form: medium-sized tree Max. height: 25 m (herbarium) Max. diameter: 50 cm (herbarium) Leaf: alternate, paripinnately compound, 6-10 leaflets, elliptic, mesophyll (3-10 x 7-25 cm), entire, coriaceous, glabrous, venation reticulate and prominent on both surfaces Inflorescence: terminal or axillary, branched Flower: large; one prominent petal, white, green in throat; fragrant Fruit: dry dehiscent (pod) (9 x 35 cm), woody with soft, dense, orange-brown hairs Seed: very large Other: it has a spreading crown. Seedlings and saplings of Berlinia tomentella can not easily be distinguished from Berlinia confusa.

B

Phenology Dispersal: explosive pods (Hall & Swaine 1981) Timing: not clearly seasonal (Hall & Swaine 1981)

Distribution Data sources

Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 27 30’ cells, distribution range is 1234 km Forest type: wet evergreen forest, moist evergreen forest, moist semi-deciduous forest, secondary forest, swamp forest

FWTA, Hall & Swaine (1981), Lock (1989), Hawthorne (1995a)

Habitat Species occurrence is higher where rainfall is between 1500-2000 mm/yr (Chi2 test), and in the lowlands (logistic regression analysis). It has been classified as a shade bearer (Hawthorne 1995a) although most herbarium records are from disturbed places (e.g. forest borders, secondary bushes and along roadsides). Frequently at wet

spp

148

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

B.t.

37

73 (15)

5

49

5

14

44

25

19

5

62

32

30

5

H 43

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 149-158

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Page 149

Bertiera spicata

(P.Gaertn.) Wernham

Rubiaceae Description Guild: np Life form: shrub Max. height: 5 m (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, ovate to oblong, mesophyll (4-9.5 x 11-27 cm), entire, coriaceous, brown hairy on both sides; interpetiolar stipules Inflorescence: terminal, unbranched (raceme) (up to 20 cm long), with sessile or subsessile cluster of crowded flowers Flower: small (corolla tube 0.1 cm across); 5-merous; calyx green; corolla tube green, lobes white Fruit: fleshy, globose (0.5 cm in diameter), brown; many seeds Seed: small (0.1 x 0.1 cm) Other: it has hairy twigs and petioles.

borders, along roadsides, and secondary vegetation. Usually on humid, humus-rich soils with high water holding capacity (Chi2 test) but sometimes also on gravel and lateritic soils (herbarium).

B

Phenology

Distribution Continent: Upper Guinea endemic Upper Guinea: Senegal, Guinea-Bissau, Guinea, Sierra Leone, Liberia, Côte d’Ivoire Distribution type: continuous, widespread, present in 28 30’ cells, distribution range is 1380 km Forest type: coastal forest, coastal shrubland, mangrove forest, swamp forest, gallery forest, secondary forest

Data sources FWTA, Hawthorne & Jongkind (2004)

Habitat Species occurrence increases with rainfall to reach an optimum around 3000 mm/yr (logistic regression analysis, Chi2 test). It is often found near the coast (Chi2 test). Generally found in the forest understorey, close to streams, rivers, and swamps (herbarium). Especially in drier regions it is found near rivers (Hawthorne & Jongkind 2004). Sometimes also in open places such as forest

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

H

B.s.

50

52 (21)

0

36

26

4

22

8

56

2

82

16

50

14

14

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

149

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Bonamia vignei

Page 150

Hoyle

Convolvulaceae Description

Phenology

Guild: u Life form: large winding woody climber Max. height: data unavailable Max. diameter: data unavailable Leaf: alternate, simple, oblong to lanceolate-elliptic, notophyll (2-5 x 5-12 cm), ciliate Inflorescence: terminal; cyme Flower: medium-sized; corolla white (up to 1.5 cm long) Fruit: dry dehiscent, with 4 valves, ovoid (1 cm in diameter), bright red; 2 seeds Seed: medium-sized (0.7 cm long), orange to red with blackish aril, seeds exposed after pericarp has been dropped Other: it has no exudate. It has dark-brown pilose stems, inflorescences, and petioles.

B

Data sources FWTA, Hoyle (1934), Hall & Swaine (1981), Breteler (1992)

Distribution Continent: Upper Guinea endemic Upper Guinea: Ghana Distribution type: continuous, local, present in 3 30’ cells, distribution range is 103 km Forest type: moist semi-deciduous forest, dry forest, thickets

Habitat Found in forest.

spp

150

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

B.v.

8

0 (7)

0

0

0

25

76

0

0

0

0

100

0

0

100

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 149-158

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Page 151

Brachystegia leonensis

Burtt Davy & Hutch.

Leguminosae-Caes. Description

Distribution type: continuous, widespread, present in 13 30’ cells. The tree may be very common locally (Voorhoeve 1965). The genus Brachystegia is most widely represented in the woodland savanna, with only two high forest species in West Africa (Voorhoeve 1965). Forest type: rainforest

Guild: sb Life form: large tree Max. height: 50 m (FWTA) Max. diameter: 180 cm (Voorhoeve 1965) Leaf: alternate, paripinnately compound, 14-26 leaflets, elliptic, microphyll (0.5-4.5 x 1.5-12 cm), entire, fairly coriaceous, young leaves densely pilose, later glabrescent Inflorescence: terminal or axillary, branched Flower: small; greenish yellow Fruit: dry dehiscent (6 x 20 cm), purplish brown, thick woody Seed: flattened disc, large (2 x 2 x 0.4 cm) Other: the tree has low, heavy root swellings when young and may have thick buttresses later on. It has drooping twigs and leaves.

B

Habitat It occurs on ridges and in valleys near water (Hawthorne 1995a).

Regeneration It has a phanerocotylar epigeal reserve seedling type, with cotyledons pressed against the epicotyl (Voorhoeve 1965). Regeneration is common in open places where the soil has been disturbed, such as logging roads (Voorhoeve 1965).

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire

Phenology Deciduousness: evergreen (Voorhoeve 1965) Timing: flowering period from February to April; fruiting period from July to November (Voorhoeve 1965)

Uses It is a timber species (Voorhoeve 1965).

Data sources FWTA, Voorhoeve (1965), Lock (1989), Hawthorne (1995a)

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

B.l.

20

11 (9)

0

60

5

0

0

50

50

5

75

20

35

5

35

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

151

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Page 152

Buforrestia obovata

Brenan

Commelinaceae Description

Phenology

Guild: sb Life form: stoloniferous herb Max. height: 0.35 m (FWTA) Leaf: alternate, simple, obovate, notophyll (4.5-7 x 7-11 cm), entire, herbaceous; a pseudopetiole surrounds the stem Inflorescence: axillary, branched, emerging through a hole at the base of the leaf-sheath Flower: small; reddish brown (white, FWTA) Fruit: dry dehiscent (capsule) Seed: data unavailable

B

Distribution

Data sources

Continent: Upper Guinea endemic (herbarium) Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana (herbarium) Distribution type: continuous, widespread, present in 7 30’ cells, distribution range is 1231 km Forest type: locally abundant in lowland rainforest.

FWTA

Habitat It is found on moist to damp forest floors (herbarium).

spp

152

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

B.o.

8

0 (4)

25

38

13

0

38

50

13

0

75

25

63

0

H 25

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 153

Bussea occidentalis

Hutch. ex Chipp.

Leguminosae-Caes. Description

Habitat

Guild: np Life form: large tree Max. height: 45 m (De Koning 1983) Max. diameter: 100 cm (Voorhoeve 1965) Leaf: alternate, bipinnately compound, 8-10 pinnae each with 12-22 leaflets, elliptic, microphyll (1.5-4 x 4-10 cm), entire, herbaceous, glossy green above, dull green beneath Inflorescence: axillary or terminal, unbranched Flower: large; calyx inside green, outside brown velutinous; corolla yellow Fruit: dry dehiscent, flattened, narrowly obovate, thick (3 x 20 cm), woody and velvety, reddish brown, standing erect on the surface of the crown; 1-2 seeds Seed: flat, very large (2 x 3.5 cm), yellowish-brown, edible Other: it has a dense, rounded and small crown. The bole is usually crooked, without buttresses but it can be fluted. The leaf-bearing branchlets, petioles and rachae are brown tomentose.

More frequently occurring in areas where rainfall is between 1500-2000 mm/yr (Chi2 test). Although in Ghana, the species seems to be mainly associated with upland evergreen forests (Hall & Swaine 1981, Hawthorne 1995a), herbarium records for the whole Upper Guinea region indicate no preference for certain altitudinal ranges. Occasionally along streams and rivers (herbarium).

B

Regeneration Germination is normal (Hawthorne 1995a). It has a phanerocotylar epigeal reserve seedling type (cf. Voorhoeve 1965). Seedlings are most often seen in the shade, and are not common in large gaps. The tree is a marked shade-bearer, benefiting from increased light levels in small to medium-sized gaps. Taylor (1960) records a height increment of c. 15-35 cm per year.

Phenology

Distribution

Deciduousness: evergreen Dispersal: seeds are shot from bursting pods, and eaten by colobus monkeys (Hall & Swaine 1981) Timing: flowering period from June to November; fruiting period from September to the end of the dry season (Hawthorne 1995)

Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 35 30’ cells, distribution range is 1188 km Forest type: upland evergreen forest, wet evergreen forest, moist evergreen forest, moist semi-deciduous forest, dry semi-deciduous forest, gallery forest, secondary forest. In Ghana, present in all forest types, but especially so in upland evergreen forest (Hall & Swaine 1981). In Liberia, more common in northern forests (Voorhoeve 1965).

Uses Fresh or roasted seeds are edible. The bark is used for treating sleeping sickness and jaundice. The hard and heavy wood is locally used for making axehandles (Voorhoeve 1965).

Data sources Taylor (1960), Voorhoeve (1965), Hall & Swaine (1981), De Koning (1983), Hawthorne (1995a), Hawthorne & Jongkind (2004)

spp

B.o. All

n

60 46333

Open (n)

22 (41) 37

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

3

42

5

20

45

23

8

0

68

32

43

2

42

4

39

7

37

29

24

10

6

69

25

36

13

39

H

153

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Page 154

Byrsanthus brownii

Guill.

Flacourtiaceae Description

Phenology

Guild: u Life form: small tree or shrub Max. height: 12 m (herbarium) Max. diameter: data unavailable Leaf: alternate, simple, elliptic, notophyll (4-6 x 611 cm), entire, coriaceous, glabrous; no stipules Inflorescence: axillary, branched (paniculate) Flower: small; fragrant Fruit: dry dehiscent (capsule) Seed: data unavailable Other: the twigs are reddish with very conspicuous small pale lenticels.

B

Data sources FWTA, Berhaut (1975), Hawthorne & Jongkind (2004)

Distribution Continent: Upper Guinea endemic. Upper Guinea: Senegal, Gambia, Guinea Bissau, Guinea, Sierra Leone Distribution type: continuous, widespread, present in 19 30’ cells, distribution range is 788 km. In Brussels, there are specimens identified from the Democratic Republic of Congo, but these have long pedicels and no short ones and are therefore considered to be Byrsanthus epigynus Mast. Forest type: riverine forest, gallery forest

Habitat It is exclusively found in wet habitats (river borders, riverbanks) (herbarium, FWTA).

spp

154

n

Open (n)

B.b.

26

20 (5)

All

46333

37

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

L

M

H

L

M

0

58

19

27

8

12

54

0

62

38

19

27

27

39

7

37

29

24

10

6

69

25

36

13

39

4

Soil CMK

Soil WHC H

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Byttneria dahomensis

Page 155

N.Hallé,

B.guineensis, B.ivorensis Keay & Milne-Redh. Sterculiaceae

Description

Habitat

Guild: u Life form: prickly woody climber Max. height: data unavailable Max. diameter: data unavailable Leaf: alternate, simple, lanceolate, microphyll (1-2.5 x 3.5-5.5 cm), blade at least markedly crenate in upper part, glabrous leaves with a gland on the midrib beneath near the leaf-base; petiole 5-10 mm, pubescent Inflorescence: in groups of 2-9 flowers Flower: small (0.7 cm in diameter); yellowish; petals pubescent at the base Fruit: glabrous (capsule), with many spines, 6-13 mm long Seed: data unavailable Other: it starts as a shrub. B. guineensis has petioles that are 0.2-0.5 cm long. The spines of the fruit are very unequal in length, 0.1-0.9 cm long. B. ivorensis has petioles of 0.5-1.6 cm long. The spines on the fruit are approx. equal in length, 0.20.4 cm long.

All three Byttneria species occur near rivers, streams, or on seasonally inundated areas (Hallé 1962).

Distribution

Hallé (1962), IUCN Red List (2000), Hawthorne & Jongkind (2004)

B

Phenology

Data sources

B. dahomensis occurs in Benin and Nigeria. It has a very local distribution, is present in 1 30’ cell, and has a distribution range of 130 km. B. guineensis is an Upper Guinea endemic that occurs only in Sierra Leone. It has a continuous, very local distribution, and is present in 1 30’ cell. It occurs in floodplain areas (herbarium). B. ivorensis is an Upper Guinea endemic that only has been found in Côte d’Ivoire. It has a continuous, very local distribution, and is present in 1 30’ cell. It is a Red List species (extinct).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

Byttneria spp.

4

- (0)

0

75

0

50

0

0

50

0

75

25

50

0

50

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

155

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Page 156

Callichilia subsessilis

(Benth.) Stapf

Apocynaceae Fruit: fleshy, dehiscent, pendulous, narrowly ovoid, 2.5 x 4.5 cm (measurements from one carpel), yellow to orange; 4-36 seeds Seed: medium-sized (0.4 x 0.45 x 0.4 cm) Other: it is repeatedly dichotomously branched. The bark has lenticels and exudes a white latex.

C

Phenology Timing: not distinctly seasonal (Hall & Swaine 1981)

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 59 30’ cells, distribution range is 1360 km Forest type: wet evergreen forest, riverine forest, shady gallery forest, sacred wood, old secondary forest (herbarium). In Ghana, it is found mainly in moist semi-deciduous forests, but it is also present in dry semi-deciduous forests and in wet and moist evergreen forests (Hall & Swaine 1981).

Habitat Species occurrence increases significantly where rainfall is higher than 1500 mm/ yr (Chi2 test) to reach an optimum around 2700 mm/yr (logistic regression analysis). It is often found close to rivers (Chi2 test), in moist and very shady places (e.g. under the tree canopy along streams, riverbanks, waterfalls, and swampy areas). Occasionally also along roads or paths but often under shade (herbarium). It flowers mainly near clearings (Beentje 1978). Soils can be lateritic, clay-sandy, or sandy (herbarium), often with a high water holding capacity (Chi2 test).

Description Guild: sb Life form: shrub or pigmy tree Max. height: 3 m (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, elliptic to obovate, mesophyll (2-10 x 3-25 cm), entire, coriaceous, glabrous; shortly petiolate Inflorescence: terminal, solitary Flower: large (corolla 3.4-8 cm long, tube 2.2-4.9 cm long); calyx pale green; corolla white, pale yellow inside the tube; funnel shaped; no fragrance

spp

156

n

Open (n)

C.s.

166

31 (107)

All

46333

37

Regeneration It regenerates under closed canopy (Hall & Swaine 1981).

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

L

M

52

7

4

49

25

19

4

70

25

43

3

41

39

7

37

29

24

10

6

69

25

36

13

39

4

Soil WHC H

Uses The bark was formerly used to make clothes. It is cultivated in medicinal plant gardens (herbarium).

Data sources FWTA, Mangenot (1965), Beentje (1978), Hall & Swaine (1981), De Koning (1983)

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Page 157

Calpocalyx aubrevillei

Pellegr.

Leguminosae-Mim. Desription

Regeneration

Guild: sb Life form: medium-sized tree Max. height: 32 m (herbarium) Max. diameter: 110 cm (Voorhoeve 1965) Leaf: alternate, bipinnately compound, each pinnae with 6-12 opposite leaflets, ovate to elliptic, mesophyll (2-8 x 3.5-20 cm), entire, coriaceous; flat gland between the upper pairs of leaflets Inflorescence: terminal, branched (panicle), erect (up to 40 cm long with lateral spikes 15-20 cm long), peduncle hairy Flower: small; calyx golden yellow; corolla white; fragrant Fruit: dry dehiscent (pod), flat, wider at tip (9 x 25 cm), with 2 valves, thick woody, black; 1-4 seeds Seed: very large (4 x 2.5 cm), dark purplish brown Other: a buttressed tree with a small crown. The bark is grey and smooth, the slash dull yellow.

It has a phanerocotylar epigeal reserve seedling type (cf. Voorhoeve 1965). A gregarious tree, often locally abundant (herbarium). Regeneration needs light for healthy development (Voorhoeve 1965).

C

Phenology Deciduousness: evergreen Dispersal: by explosive pods. Seeds are eaten by chimpanzees (Kasparek 2000). Timing: fruits are found the whole year round (Voorhoeve 1965)

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Distribution type: continuous, widespread, present in 11 30’ cells, distribution range is 642 km Forest type: wet evergreen forest, semi-deciduous forest. In Liberia, widespread (scattered or in rich stands). Most abundant in evergreen forests and slightly less abundant in semi-deciduous forest (Voorhoeve 1965).

Habitat Mostly found where rainfall is higher than 2500 mm/yr (Chi2 test). Often found in the vicinity of water (herbarium, Voorhoeve 1965).

Uses It has edible seeds (Kasparek 2000). The species was formerly used to obtain country salt. The wood was burned and the ashes leached; the evaporated leach gave a residue of salt (Voorhoeve 1965).

Data sources FWTA, Voorhoeve (1965), Lock (1989), Kasparek (2000)

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

H

C.a.

21

19 (16)

5

33

0

0

29

15

57

0

95

5

90

0

10

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

157

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Page 158

Calpocalyx brevibracteatus

Harms

Leguminosae-Mim.

C

Distribution

Phenology

Continent: Sierra Leone to Nigeria (Hall & Swaine 1981), Nigeria, Cameroon (herbarium) Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana (herbarium) Distribution type: continuous, widespread, present in 35 30’ cells, distribution range is 1945 km Forest type: wet evergreen forest, moist evergreen forest, moist semi-deciduous forest, dry semideciduous forest, gallery forest, secondary forest. In Ghana, mostly found in wet and moist evergreen forests (Hall & Swaine 1981).

Deciduousness: sometimes briefly deciduous (Hall & Swaine 1981) Dispersal: seeds are shot explosively from dehiscing pods during the dry season (Hawthorne 1994). Seeds are eaten by chimpanzees (Kasparek 2000).

Habitat Description

Species occurrence increases with rainfall to reach an optimum around 2500 mm/yr (logistic regression analysis, Chi2 test). It is usually found in mature evergreen forests, but sometimes also in disturbed sites and along roadsides (herbarium, Hawthorne 1995a). Often found at the edge of rivers, streams, river banks and swamps (herbarium, Hawthorne 1995a). It is more abundant in unburned than in burnt forest (Hawthorne 1994). It occurs on sandy-loamy, sandy-clayish, and lateritic soils (herbarium). In Ghana, the species distribution is very significantly correlated with acidic, base-poor soils (Hall & Swaine 1981).

Guild: sb Life form: large tree Max. height: 40 m (herbarium) Max. diameter: 60 cm (Voorhoeve 1965) Leaf: alternate, bipinnately compound, 2 leaflets, lanceolate to elliptic, notophyll (2-5 x 7-15 cm), without strongly recurved margin when dry, coriaceous; young shoots with thread-like stipules Inflorescence: terminal or axillary, branched (panicle, up to 25 cm long, with lateral spikes) Flower: small, corolla pink to brownish orange Fruit: dry dehiscent, S-shaped (4 x 17 cm), woody; 5-10 seeds Seed: large Other: the tree is densely crowned, often with foliage in plumes, spreading out from low boughs. It is shortly buttressed. The bark is grey to dark brown, the slash hard-fibrous and often peelable. The sapwood is hard, orange striate, with sweet then bitter taste and has raised oval lenticels in longitudinal rows. The wood density is 0.83 g/cm3. In Ghana, the crown exposure profile is mostly of an understorey tree, but with a potentially significant tendency to exposure of approx. 20 cm dbh trees.

spp

158

n

Open (n)

Uses It has edible fruits (Vivien & Faure 1985).

Data sources FWTA, De la Mensbruge (1966), Voorhoeve (1965), Hall & Swaine (1981), Vivien & Faure (1985), Lock (1989), Hawthorne (1994, 1995a), Kasparek (2000), Hawthorne & Jongkind (2004)

Regeneration It has a phanerocotylar epigeal reserve seedling type (cf. de la Mensbruge 1966). It regenerates in shade (Hall & Swaine 1981). In evergreen forests, recruitment occurs in relatively exposed and open areas, whereas in moist semi-deciduous forests, trees are often completely shaded, possibly because of intolerance to desiccation (Hawthorne 1995a). It is often gregarious (Hawthorne & Jongkind 2004).

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

C.b.

62

33 (33)

2

34

3

8

47

21

23

5

66

29

44

3

52

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 159

Calvoa monticola

A.Chev. ex Hutch. & Dalziel

Melastomataceae Description

Phenology

Guild: sb Life form: small epiphytic or terrestrial herb Max. height: 0.23 m (herbarium) Leaf: opposite, simple, elliptic to obovate, microphyll (2-4.5 x 3.5-7 cm), serrate, succulent, glabrous, greenish-white beneath with 3 contrasting nerves; stems and petiole purple Inflorescence: branched Flower: small; corolla pinkish to pale purple Fruit: fleshy (0.8 cm long), brown; many seeds Seed: very small

C

Distribution

Data sources

Continent: Cameroon, Gabon Upper Guinea: Guinea, Liberia, Côte d’Ivoire, Ghana Distribution type: continental disjunct, present in 11 30’ cells in Upper Guinea Forest type: mountain rainforest (herbarium). A common epiphyte in West Africa (Johansson 1974).

FWTA, Johansson (1974)

Habitat It is terrestrial or epiphytic (on trunks or rocks). Usually, found in very moist and shady places (herbarium).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

C.m.

19

0 (6)

26

47

0

0

32

58

10

0

89

11

58

0

H 32

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

159

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Page 160

Calycobolus insignis

(Rendle) Heine

Convolvulaceae Distribution Continent: Upper Guinea endemic Upper Guinea: Liberia Distribution type: continuous, local, present in 4 30’ cells, distribution range is 286 km Forest type: mature forest, old secondary forest

C

Habitat Found in hilly and flat areas, sometimes close to water (e.g. island in mouth of river) (herbarium).

Phenology Dispersal: by wind

Description Guild: u Life form: winding woody climber or shrub Max. height: 2 m long (herbarium) Max. diameter: data unavailable Leaf: alternate, simple, obovate to elliptic, notophyll (3.5-7 x 7-12 cm), entire, coriaceous, surface beneath with many very small glandular dots; petiole small with brown hairs Inflorescence: terminal or axillary, not branched Flower: medium-sized; calyx purple-green; corolla white Fruit: dry indehiscent, with two wings, pale green; 1 seed Seed: small (0.2 x 0.1 cm) Other: the stem has brown hairs.

spp

C.i. All

160

n

Open (n)

Data sources FWTA

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

6

33 (3)

0

67

17

0

0

0

84

0

100

0

83

0

0

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 161

Campylospermum amplectens

(Stapf ) Engl.

Ochnaceae Description Guild: sb Life form: small tree or shrub Max. height: 10 m (herbarium) Max. diameter: data unavailable Leaf: alternate, simple, elongate-oblanceolate to elongate-obovate, mesophyll (6-8 x 20-32 cm), serrate, stiff coriaceous, with leaf base clasping stem and collecting debris there Inflorescence: axillary, branched Flower: medium-sized; bowl-shaped; calyx brown turning scarlet-red when older in fruit; corolla bright yellow Fruit: fleshy, black on an enlarged red calyx Seed: medium-sized (0.8 x 0.5 x 0.4 cm) Other: it has aerial roots that sometimes penetrate the leaf litter, and may help with nutrition on the infertile evergreen forest soils (Hawthorne 1995a).

C

Regeneration It regenerates in shade (Hall & Swaine 1981).

Distribution

Phenology

Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 11 30’ cells, distribution range is 988 km, Red List species (vulnerable) Forest type: evergreen forest. In Ghana, it is mostly found in wet evergreen forests and scarcely in moist evergreen forests (Hall & Swaine 1981).

Deciduousness: not clearly seasonal (Hall & Swaine 1981)

Habitat Species occurrence increases sharply when rainfall reaches more than 2000 mm/yr (logistic regression analysis), and it is mostly found in places where rainfall is higher than 2500 mm/yr (Chi2 test). Usually, found in the understorey of mature forests (herbarium), often close to rivers (Chi2 test).

spp

n

Open (n)

Data sources FWTA, Hall & Swaine (1981), Hawthorne (1995a)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

C.a.

16

10 (10)

6

88

6

0

31

6

63

0

100

0

69

0

31

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

161

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Page 162

Campylospermum subcordatum

(Stapf ) Engl.

Ochnaceae Distribution

Phenology

Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire Distribution type: continuous, widespread, present in 21 30’ cells, distribution range is 934 km Forest type: montane forest, wet evergreen forest, swamp forest, secondary forest.

C

Habitat Species occurrence increases with rainfall to reach an optimum around 2500 mm/yr (logistic regression analysis, Chi2 test). Commonly found in the understorey of dense evergreen forests, and only occasionally in open gaps or secondary forests. On sandy laterite and loamy soils (herbarium).

Description Guild: sb Life form: small tree or shrub Max height: 5 m (herbarium) Max diameter: data unavailable Leaf: alternate, simple, spatulate to oblanceolate, macrophyll (6-14 x 22-51 cm), finely serrate, coriaceous; sessile Inflorescence: axillary, branched Flowers: medium-sized; calyx red; corolla bright yellow Fruit: fleshy, black on an enlarged red calyx; 1 seed Seed: medium-sized (0.8 x 0.5 x 0.5 cm)

spp

162

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

C.s.

35

17 (23)

11

43

0

3

34

17

45

0

86

14

74

0

H 23

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

Data sources FWTA

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Page 163

Cassipourea afzelii

(Oliv.) Alston

Rhizophoraceae Description

Phenology

Guild: sb Life form: shrub or small tree Max. height: 15 m (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, broadly oblong to elliptic, notophyll (3-6 x 7-14 cm), entire to crenate, herbaceous, glabrous, interpetiolar stipules Inflorescence: axillary, arranged in sessile clusters Flower: small; calyx green; corolla white, anthers prominent Fruit: indehiscent, green; 1 seed Seed: small (0.2 x 0.2 cm)

C

Distribution

Data sources

Continent: Nigeria Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continental disjunct with the largest population in Upper Guinea; in Upper Guinea present in 23 30’ cells Forest type: montane forest, wet evergreen forest, gallery forest, secondary forest

FWTA, De Rouw (1991), Hawthorne & Jongkind (2004)

Habitat The occurrence increases very sharply when rainfall is higher than 2500 mm/yr (logistic regression analysis, Chi2 test). The species is usually found in the forest understorey close to water (e.g. riverbanks, along rivers and creeks), and it is often observed close to the sea, near the beach (Chi2 test, Hawthorne & Jongkind 2004). Very often on sandy, sandy-loamy soils (herbarium).

Regeneration It regenerates in primary forests (De Rouw 1991).

spp

C.a. All

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

H

23

29 (7)

4

43

26

4

4

21

69

0

87

13

70

0

9

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

163

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Page 164

Cassipourea hiotou

Aubrév. & Pellegr.

Rhizophoraceae

C

Description

Phenology

Guild: sb Life form: small tree Max. height: 13 m (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, elliptic, notophyll (4-4.5 x 9.5-12.5 cm), entire, herbaceous, pubescent, interpetiolar stipules Inflorescence: axillary, several simple flowers on long stalks Flower: medium-sized, bowl-shaped; corolla greenish white, hairy Fruit: indehiscent Seed: data unavailable Other: it has hairy twigs and petioles and a redbrown slash.

Deciduousness: evergreen

Data sources Aubréville & Pellegrin (1958), Hall & Swaine (1981), Hawthorne (1995a), IUCN Red List (2000)

Distribution Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire, Ghana Distribution type: Upper Guinea disjunct, regional, present in 3 30’ cells, distribution range is 554 km, Red List species (vulnerable), although it is fairly common in evergreen forest in Ghana (Hall & Swaine 1981) Forest type: wet evergreen forest

Habitat Found in the lower strata of rainforests.

spp

164

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

C.h.

3

100 (1)

0

67

0

0

33

66

0

0

100

0

67

0

33

all

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 165

Cathormion rhombifolium

(Benth.) Keay

Leguminosae-Mim. Description

Phenology

Guild: u Life form: medium-sized tree Max. height: 12 m (herbarium) Max. diameter: data unavailable Leaf: opposite, bipinnately compound, 4-6 pinnae, each with up to 14 leaflets, oblong to obovate, microphyll (up to 2.5 x 4.5 cm), hairy Inflorescence: flowers in a head (capitate) Flower: corolla white; fragrant Fruit: dry indehiscent, flat but thick, oblong (1.5 x 9 cm); breaking up into 1-seeded segments; 10 seeds Seed: medium-sized (0.6-1 cm long), lenticular

C

Data sources

Distribution

FWTA, Brenan & Brummitt (1965), Voorhoeve (1965), Hawthorne & Jongkind (2004)

Continent: Upper Guinea endemic Upper Guinea: Senegal (FWTA), Guinea Bissau, Guinea, Sierra Leone (herbarium) Distribution type: continuous, regional, present in 5 30’ cells, distribution range is 434 km Forest type: swamp forest, gallery forest

Habitat Found in swamp forest and along riversides (herbarium, Hawthorne & Jongkind 2004).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

C.r.

7

33 (3)

0

71

14

14

0

14

72

0

71

29

29

14

57

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

165

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Page 166

Cavacoa baldwinii

(Keay & Cavaco) J.Léonard

Euphorbiaceae Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia Distribution type: continuous, local, present in 3 30’ cells, distribution range is 185 km Forest type: wet evergreen forest, moist evergreen forest

C

Habitat It is found in the understorey of moist evergreen forests, near creeks or rivers. Apparently it is not strongly restricted to swamps (Hawthorne & Jongkind 2004). It occurs on loamy soils (herbarium).

Phenology

Description Guild: sb Life form: shrub Max. height: 2.4 m (herbarium) Max. diameter: data unavailable Leaf: alternate, simple, narrowly elliptic, mesophyll (up to 5 x 13 cm), entire, coriaceous, several small glands on the lower leaf surface, glabrous beneath; petiole hairy above Inflorescence: terminal, racemose, young, with cone-like overlapping bracts, dioecious Flower: small; male flowers yellow; female flowers green Fruit: data unavailable Seed: data unavailable

spp

166

n

Open (n)

Data sources FWTA, Hawthorne & Jongkind (2004)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

C.b.

11

17 (6)

0

45

0

0

0

0

91

0

100

0

100

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 167

Chassalia corallifera

(A.Chev. ex De Wild.) Hepper

Rubiaceae Description

Phenology

Guild: sb Life form: shrub or pigmy tree Max. height: 1 m (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, elliptic, mesophyll (4-8 x 8-18 cm), entire, herbaceous, glabrous; interpetiolar stipules Inflorescence: terminal, branched (panicle) Flower: small; bowl-shaped with tube, corolla creamy; on bright red branches Fruit: fleshy (berry), globose (0.7 cm in diameter), black, on coral-pink fleshy pedicel Seed: medium-sized (0.4 x 0.4 x 0.2 cm)

Dispersal: probably by animals. Timing: not clearly seasonal (Hall & Swaine 1981)

C

Distribution Data sources

Continent: Upper Guinea endemic (Hall & Swaine 1981) Upper Guinea: Liberia, Côte d’Ivoire, Ghana (herbarium) Distribution type: continuous, widespread, present in 22 30’ cells, distribution range is 1139 km Forest type: wet evergreen forest, moist evergreen forest, moist semi-deciduous forest, secondary forest. In Ghana, it is equally common in all three non-secondary forest types (Hall & Swaine 1981).

FWTA, Hall & Swaine (1981)

Habitat Species occurrence tends to be higher in places where rainfall reaches 1500-2000 mm/yr (Chi2 test). Usually in the forest understorey, where it can grow under dense shade (Hall & Swaine 1981). Sometimes found near streams and rivers. On sandy clay (herbarium).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

C.c.

35

33 (21)

0

43

0

6

80

9

6

6

43

51

29

6

60

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

167

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Page 168

Chazaliella cupulicalyx

Verdc.

Rubiaceae Description

Phenology

Guild: u Life form: scandent shrub Max. height: 5 m (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, elliptic, mesophyll (1.5-8 x 5-26 cm), entire, coriaceous, glabrous; interpetiolar stipules Inflorescence: terminal, branched Flower: small (corolla tube 0.3 cm); calyx pale green with strongly reflexed lobes; corolla white and hairy Fruit: fleshy, oblong (0.5 x 1 cm), red Seed: medium-sized (0.6 x 0.6 x 0.15 cm) Other: it has a corky bark.

C

Data sources

Distribution

Verdcourt (1977), Hawthorne & Jongkind (2004)

Continent: Upper Guinea endemic Upper Guinea: Liberia (herbarium, Hawthorne & Jongkind 2004) Distribution type: continuous, local, present in 3 30’ cells, distribution range is 189 km Forest type: moist evergreen forest, secondary forest

Habitat Sometimes found near rivers (herbarium).

spp

168

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

C.c.

5

0 (2)

0

60

0

0

0

0

100

0

100

0

100

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 169

Chytranthus cauliflorus

(Hutch. & Dalziel) Wickens

Sapindaceae Description Guild: sb Life form: small tree Max. height: 8 m (herbarium) Max. diameter: 4 cm (herbarium) Leaf: alternate, pinnately compound, 6-14 leaflets, broadly elliptic, mesophyll (4-6.5 x 7.5-35 cm), entire, coriaceous, very hairy Inflorescence: cauliflorous, not branched (spike), probably polygamous Flower: small; bowl-shaped; calyx and bracts with brown hairs; corolla creamy Fruit: fleshy (berry) (7 x 11 cm), yellow, glabrous; 2 seeds Seed: large (1.9 x 0.7 x 0.2 cm), brown Other: rather unbranched, with hairy stems, petioles and midribs of leaves.

C

Distribution Phenology

Continent: Gabon Upper Guinea: Liberia, Côte d’Ivoire, Ghana Distribution type: continental disjunct, in Upper Guinea present in 6 30’ cells Forest type: wet evergreen forest, moist evergreen forest, secondary forest

Habitat In Ghana, its distribution is very highly correlated with acidic, base-poor soils (Hall & Swaine 1981).

Regeneration Data sources

It regenerates in shade (Hall & Swaine 1981).

FWTA, Hallé & Aké Assi (1962), Aubréville & Leroy (1973b), Hall & Swaine (1981), de Koning (1983)

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

C.c.

12

11 (9)

0

33

0

0

67

25

8

8

33

58

8

8

83

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

169

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Page 170

Clerodendrum sassandrense

Jongkind

Verbenaceae Habitat Data unavailable

Phenology

C

Description Guild: u Life form: small creeping shrub Max. height: 0.3 m (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, herbaceous Inflorescence: branched (cymose) Flower: medium-sized (approx. 1 x 1 cm); calyx green; corolla tubular with asymmetric lobes, white Fruit: dry dehiscent, shiny black, orange inside, calyx red; 4 seeds Seed: data unavailable Other: most branches creep on the forest floor and root

Data sources Hawthorne & Jongkind (2004)

Distribution Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire Distribution type: continuous, very local, present in 2 30’ cells, distribution range is 10 km. It is only known from southwest Côte d’Ivoire Forest type: secondary forest

spp

170

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

C.s.

2

100 (1)

0

0

0

0

0

100

0

50

50

0

50

50

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 171

Cola attiensis

Aubrév. & Pellegr.

Sterculiaceae Description Guild: sb Life form: small tree Max. height: 7 m (herbarium) Max. diameter: 8 cm (herbarium) Leaf: alternate, simple, elliptic, mesophyll (4-12 x 15-38 cm), entire; stipules up to 1.5 cm long, falling off prematurely Inflorescence: cauliflorous; flowers in clusters (fasciculate) Flower: small (approx. 1 cm), calyx pubescent Fruit: fleshy, globose (1 cm in diameter), dark red, glabrous; in bundles on trunk; 1-2 seeds Seed: large

C

Distribution

Phenology

Continent: Cameroon, Gabon Upper Guinea: Côte d’Ivoire Distribution type: continental disjunct, in Upper Guinea present in 6 30’ cells, Red List species (endangered) Forest type: rainforest, evergreen forest

Dispersal: by animals (primates)

Habitat In closed lowland forest, on sandy clay (Hawthorne & Jongkind 2004, herbarium).

Data sources FWTA, Aubréville (1959), IUCN Red List (2000), Hawthorne & Jongkind (2004)

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

C.a.

14

0 (10)

0

29

7

57

43

0

0

7

79

14

43

7

50

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

171

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Cola boxiana

Page 172

Brenan & Keay

Sterculiaceae

C

Description

Phenology

Guild: sb Life form: medium-sized tree Max. height: 30 m (herbarium) Max. diameter: data unavailable Leaf: alternate, simple, oblanceolate to elliptic, mesophyll (4-9 x 10-24 cm), entire, herbaceous to coriaceous Inflorescence: in dense clusters (fascicles) Flower: corolla yellow, 5-lobed; fragrant Fruit: irregularly subglobose (1.8 x 2.3 cm), red, dehiscent; many follicles, with 2 seeds per follicle Seed: large

Deciduousness: evergreen Dispersal: by animals (primates)

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 6 30’ cells, distribution range is 1171 km, Red List species (endangered). It can be locally common in some upland forests (Hawthorne & Jongkind 2004). Forest type: montane forest, rainforest

Data sources FWTA, IUCN Red List (2000), Hawthorne & Jongkind (2004)

Habitat It is found in the understorey or in open places.

spp

C.b. All

172

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

8

14 (7)

0

50

0

0

88

0

13

0

38

63

38

0

38

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 173

Cola caricifolia

(G.Don) K.Schum.

Sterculiaceae Description

secondary vegetation) but also present in dense forests. It has been classified both as a pioneer (De Koning 1983, Hawthorne 1995a) and as a shade bearer (Hall & Swaine 1981). More abundant in unburned forests (Hawthorne 1994). The species has a preference for infertile soils (Swaine 1996), and occurs on sandy and sandy-clay soils (herbarium).

Guild: pi Life form: small tree Max. height: 10 m (De Koning 1983) Max. diameter: 7 cm (herbarium) Leaf: alternate, palmately compound, 5-lobed, macrophyll (6-30 x 10-35 cm), entire, herbaceous, with white hairs; petiole 2-30 cm long; stipules up to 1 cm long, falling off prematurely Inflorescence: cauliflorous Flower: large; monoecious; bowl-shaped; corolla tube pink, lobes deep red Fruit: fleshy, dehiscent, sessile, oblong (2 x 10 cm), yellow to orange, 4-10 fruit parts, strongly beaked; 4-10 seeds Seed: large

Regeneration It has a phanerocotylar epigeous foliaceous seedling type (De Koning 1983).

Phenology Deciduousness: evergreen Dispersal: by animals (primates)

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continental disjunct, with the largest population in Upper Guinea, present in 29 30’ cells in Upper Guinea. Forest type: wet evergreen forest, moist evergreen forest, moist semi-deciduous forest, dry semideciduous forest, coastal forest, secondary forest. In Ghana, mostly found in moist and wet evergreen forests (Hall & Swaine 1981). According to Hawthorne (1995a), a dry forest tree, occurring in moister forests on better-drained sites.

Uses

Habitat

It has edible seeds (De Koning 1983).

Species occurrence increases significantly with rainfall to reach an optimum around 2500 mm/yr (logistic regression analysis, Chi2 test). Often in moist places of the forest or close to rivers (Chi2 test), on well-drained soils. Often in open places (e.g. clearcuts, forest edges,

spp

n

C

Open (n)

Data sources FWTA, Hall & Swaine (1981), De Koning (1983), Hawthorne (1994, 1995a), Swaine (1996)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

C.c.

71

38 (37)

1

52

8

21

31

32

14

11

68

21

34

13

H 52

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

173

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Cola reticulata

Page 174

A.Chev.

Sterculiaceae Forest type: upland evergreen, moist semideciduous forest, dry semi-deciduous forest. In Ghana, most often in upland evergreen forests (Hall & Swaine 1981). It can be locally abundant (herbarium).

C

Habitat Species occurrence increases significantly with altitude (Chi2 test, Hawthorne & Jongkind 2004) and is highest where rainfall reaches 1500-2000 mm/yr (Chi2 test). It is usually found in the forest understorey of mature undisturbed forests. Occasionally along riversides. On loamy soils (herbarium).

Description

Phenology

Guild: sb Life form: small tree Max. height: 10 m (Hawthorne & Jongkind 2004) Max. diameter: data unavailable Leaf: alternate, simple, obovate to elliptic, mesophyll (3-8 x 7-17 cm), entire, herbaceous, glabrous Inflorescence: monoecious; axillary; solitary Flower: medium-sized; bell-shaped; perianth pale yellow Fruit: fleshy (berry), ovoid (3 x 4 cm), red to orange Seed: large Other: it has a yellow slash.

Deciduousness: evergreen (Hawthorne 1995a) Dispersal: by animals (primates) Timing: not clearly seasonal (Hall & Swaine 1981)

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea (FWTA), Liberia, Côte d’Ivoire, Ghana (herbarium) Distribution type: continuous, widespread, present in 15 30’ cells, distribution range is 1119 km, Red List species (vulnerable)

spp

174

n

Open (n)

Data sources FWTA, Hall & Swaine (1981), De Koning (1983), Hawthorne (1995a), IUCN Red List (2000), Hawthorne & Jongkind (2004)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

C.r.

24

0 (14)

33

42

4

8

67

25

0

0

54

46

25

0

58

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 175

Cola umbratilis

Brenan & Keay

Sterculiaceae Description Guild: sb Life form: small tree Max. height: 7 m (herbarium) Max. diameter: data unavailable Leaf: alternate, palmately compound, 7 leaflets, elliptic, mesophyll (4.5-9 x 14-28 cm), entire, herbaceous Inflorescence: cauliflorous, flowers in clusters (fasciculate) Flower: monoecious; medium-sized; pink to red, bowl-shaped Fruit: fleshy (berry), dehiscent (up to 4 x 10 cm), red with white pulp Seed: very large

C

Distribution Phenology

Continent: Upper Guinea endemic Upper Guinea: Ghana Distribution type: continuous, very local, present in 3 30’ cells, distribution range is 68 km, Red List species (vulnerable) Forest type: wet evergreen forest Other: given its distribution, it is probably also present in Côte d’Ivoire.

Dispersal: probably by animals

Habitat It is found in forest understorey.

Regeneration It regenerates in shade (Hall & Swaine 1981).

Data sources FWTA, Hall & Swaine (1981), IUCN Red List (2000)

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

C.u.

10

10 (10)

0

20

10

0

80

20

0

0

70

30

0

0

100

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

175

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Page 176

Combretum bipindense

Engl. & Diels

Combretaceae

C

Description

Phenology

Guild: np Life form: large winding woody climber Max. height: data unavailable Max. diameter: 18 cm (herbarium) Leaf: opposite, simple, oblong-ovate to ovate, mesophyll (3.5-9 x 6.5-25 cm), entire, coriaceous, when young almost white below, later more yellowish. It can be easily recognised by the small red scales on the undersurface of the leaf. Inflorescence: axillary or terminal, unbranched (spike) Flower: small to medium; green and white, turning yellow; trumpet-shaped Fruit: dry indehiscent, round or elliptic (approx. 3 cm), pale green to brown, 4-winged; 1 seed Seed: small (< 0.1 cm) Other: when young it is also found as scrambling shrub. The exudate is colourless sticky and scarce. The wood is yellowish.

Dispersal: by wind

Data sources FWTA, Mabberley (1987), Jongkind (in prep.)

Distribution Continent: Cameroon Upper Guinea: Guinea Bissau, Côte d’Ivoire, Ghana Distribution type: continental disjunct (it occurs mostly in Upper Guinea, with a single observation in south Cameroon), in Upper Guinea present in 9 30’ cells Forest type: riverine forest, swamp forest, degraded forest

Habitat It is often found along riverbanks, streams and swamps.

spp

176

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

C.b.

10

0 (2)

0

40

10

60

30

10

0

10

50

40

30

30

30

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 177

Combretum blepharopetalum

Wickens

Combretaceae Description

Phenology

Guild: u Life form: large winding woody climber Max. height: data unavailable Max. diameter: data unavailable Leaf: opposite, simple (up to 14 x 26 cm), entire Inflorescence: axillary, unbranched (spike) Flower: small to medium-sized; green and white; flowers with a conspicuous long and slender stipelike part, with small petals Fruit: dry indehiscent, elliptic (3.5 x 3 cm), 4-winged; 1 seed Seed: data unavailable Other: not much is known from this species, all collections in the herbarium lack plant and vegetation descriptions.

Dispersal: by wind

C

Data sources

Distribution

Wickens (1971), Mabberley (1987), Hawthorne & Jongkind (2004)

Continent: Upper Guinea endemic Upper Guinea: Sierra Leone Distribution type: present in 3 30’ cells Forest type: data unavailable

Habitat Data unavailable.

spp

n

C.b.

4

All

46333

Open (n)

- (0) 37

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

0

25

0

0

0

0

100

0

75

25

25

0

50

4

39

7

37

29

24

10

6

69

25

36

13

39

177

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Page 178

Combretum calobotrys

Engl. & Diels

Combretaceae Fruit: dry indehiscent, round or elliptic (3 cm in diameter), 5-winged; 1 seed Seed: small (0.2 cm long) Other: when young also found as a scandent shrub.

C

Distribution Continent: Upper Guinea endemic Upper Guinea: Liberia, Côte d’Ivoire Distribution type: continuous, regional, present in 6 30’ cells, distribution range is 318 km Forest type: wet evergreen forest

Habitat Found in disturbed habitats or marshy places (herbarium).

Phenology Dispersal: by wind

Description Guild: u Life form: small winding woody climber Max. height: 1.8 m long (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, obovate to oblong, notophyll ( 4-4.5 x 6-10), entire, coriaceous, nerves dark green beneath; red petioles Inflorescence: axillary or terminal, not branched (spike), 10-15 cm long Flower: medium-sized; calyx red; corolla bright red, 5 petals, prominent stamens

spp

178

n

Open (n)

Data sources FWTA, Mabberley (1987), Jongkind (in prep)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

C.c.

8

100 (3)

0

0

13

0

13

13

76

0

88

13

88

0

13

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 179

Combretum grandiflorum

G.Don

Combretaceae Description

Phenology

Guild: pi Life form: large winding woody climber Max. height: over 20 m long (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, elliptic, mesophyll (5-6 x 7.5-15 cm), entire, coriaceous, glabrous, shiny dark green above, light green below, reddish veins, prominent below Inflorescence: axillary, branched Flower: medium-sized (approx. 3 cm long); corolla bright red, 5 petals; fragrant Fruit: dry indehiscent, round or elliptic (2 x 2.5 cm), golden brown, 5-winged; 1 seed Seed: small (0.1 cm long) Other: when young found as a scandent shrub. The stems are dark brown and hollow. The bark peels off in fibrous strips.

Dispersal: by wind

Distribution

C

Data sources Hawthorne & Jongkind (2004)

Continent: Upper Guinea endemic Upper Guinea: Gambia, Senegal, Guinea Bissau, Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 56 30’ cells, distribution range is 1858 km Forest type: wet evergreen forest, swamp forest, coastal shrubland, secondary forest

Habitat Species occurrence increases with rainfall to reach a very wide optimum range between 1800 to 3300 mm/yr (logistic regression analysis, Chi2 test). It frequently occurs near the coast (Chi2 test). It is almost exclusively found in disturbed habitats such as forest edges, forest gaps, secondary forests, plantations, and along roadsides. Sometimes found along riverbanks and swamps. On sandy or lateritic soils.

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

C.g.

72

70 (30)

0

39

18

18

41

21

18

3

53

44

31

10

H 47

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

179

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Page 180

Combretum tarquense

J.J.Clark

Combretaceae

C

Description Guild: pi Life form: small winding woody climber Max. height: 3.5 m long (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, oblong, notophyll (2.5-6 x 7-10 cm), entire, herbaceous Inflorescence: terminal, not branched (spike) Flower: small; bright red; 5 petals Fruit: dry indehiscent, red, 5-winged; 1 seed Seed: small (0.1 cm long) Other: when young it can be found as a scandent shrub. The branchlets have long pale green to brown pilose hairs.

small range between southeast Côte d’Ivoire and southwest Ghana. Forest type: wet evergreen forest, secondary rainforest, thickets

Habitat It is often found in disturbed habitats; secondary forest, forest edges, roadside thickets (herbarium).

Phenology Dispersal: by wind

Distribution

Data sources

Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire, Ghana Distribution type: continuous, local, present in 6 30’ cells, distribution range is 159 km. It has a

spp

180

n

Open (n)

C.t.

11

60 (5)

All

46333

37

FWTA, Mabberley (1987), Hawthorne & Jongkind (2004)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

L

M

H

L

M

H

55

9

0

73

27

0

0

82

18

0

18

82

39

7

37

29

24

10

6

69

25

36

13

39

0 4

Soil CMK

Soil WHC

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Page 181

Combretum zenkeri

Engl. & Diels

Combretaceae Description

Phenology

Guild: pi Life form: large winding woody climber Max. height: 10 m long (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, oblong to elliptic, mesophyll (2-9 x 5-18 cm), entire, herbaceous, softly pubescent beneath, veins prominent beneath; long hook-like base of petiole Inflorescence: axillary, branched, globose head (1 cm in diameter) Flower: small; whitish; fragrant Fruit: dry indehiscent, round (approx. 2 cm in diameter), pale brown, 5-winged; 1 seed Seed: data unavailable Other: when young found as a scandent shrub. The twigs and petioles are pubescent. In the flowering season, there are abundant whitish leaves surrounding the inflorescences.

Dispersal: by wind

C

Data sources FWTA, Mabberley (1987), Hawthorne & Jongkind (2004)

Distribution Continent: Benin, Nigeria, Cameroon Upper Guinea: Guinea, Côte d’Ivoire, Ghana, Togo, Benin Distribution type: continuous, widespread, present in 29 30’ cells, distribution range is 2345 km Forest type: high forest, dry forest, savanna woodland, secondary forest, thickets

Habitat Species occurrence is highest at places where rainfall is around 1000 mm/yr and decreases strongly with increasing rainfall (logistic regression analysis and Chi2 test). It is found almost exclusively in disturbed areas, such as roadside thickets, cultivated fields, and secondary vegetation. Usually on rocky soils (herbarium).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

C.z.

47

75 (12)

0

30

2

81

19

0

0

0

38

62

26

6

47

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

181

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Page 182

Commelina macrosperma

J.K.Morton

Commelinaceae Description

Phenology

Guild: sb Life form: perennial decumbent herb Max. height: 0.6 m (FWTA) Leaf: alternate, simple, ovate to elliptic, microphyll (1.8-2.2 x 3-8 cm), entire, herbaceous; pseudopetiole surrounds stem base Inflorescence: not branched Flower: medium-sized, bowl-shaped; white; bracts pale green Fruit: dry dehiscent; 2 seeds Seed: medium-sized (0.4 x 0.25 cm)

C

Data sources FWTA, Hall & Swaine (1981), De Rouw (1991)

Distribution Continent: Guinea wide: Sierra Leone to Nigeria (Hall & Swaine 1981), Nigeria (herbarium) Upper Guinea: Sierra Leone, Côte d’Ivoire, Ghana (herbarium) Distribution type: continuous, widespread, present in 9 30’ cells, distribution range is 2311 km Forest type: rainforest, moist semi-deciduous forest, dry semi-deciduous forest, southern marginal forests (Hall & Swaine 1981)

Habitat In forests and sometimes also in plantations. Often in shady (De Rouw 1991) and moist places (e.g. river and cascades). On rocky hillsides or valley bottoms on loamy lateritic soil (herbarium).

Regeneration It regenerates in shade (Hall & Swaine 1981).

spp

182

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

C.m.

9

0 (3)

0

33

0

11

78

0

11

0

33

67

22

11

56

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 183

Commiphora dalzielii

Hutch.

Burseraceae Description Guild: pi Life form: shrub or small tree Max height: 8 m (herbarium) Max diameter: data unavailable Leaf: alternate, compound, 3 leaflets, obovate to rhomboid, microphyll (1.5-3 x 2.5-7 cm), entire, herbaceous, glabrous Inflorescence: axillary, branched Flower: small, yellowish to red, 4-merous Fruit: fleshy, ellipsoid to globose (1 cm long), pericarp falling off Seed: black with red aril Other: a scrambling spiny shrub. The bark is silvery red, peeling off in papery scales.

C

Distribution Continent: Upper Guinea endemic Upper Guinea: Ghana Distribution type: continuous, local, present in 3 30’ cells, distribution range is 163 km Forest type: coastal forest, savanna, thicket

Phenology

Habitat It is found on inselbergs, and in coastal vegetation (herbarium).

Data sources FWTA, Hall & Swaine (1981), Mabberley (1987)

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

C.d.

9

100 (9)

0

22

0

100

0

0

0

0

56

44

0

0

100

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

183

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Page 184

Copaifera salikounda

Heckel

Leguminosae-Caes. Phenology

fibrous. It has a fragrant resin with distinctive sweet scent like almonds or pipe tobacco. The crown is heavily branched and spreading when in the canopy. The young branches are pubescent. Wood density is 0.75 g/cm3.

C

Deciduousness: briefly deciduous at the beginning of the rainy season, before flowering (Voorhoeve 1965) Dispersal: probably by birds, although many drop straight to the ground because ripening fruits split open on the tree (Hawthorne 1995a) Timing: flowering period from March to May (Taylor 1960); fruits ripen during the dry season (Hawthorne 1995a)

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea Bissau, Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 16 30’ cells, distribution range is 1654 km, Red List species (vulnerable). It is a rare species. Forest type: wet evergreen forest, moist evergreen forest, moist semi-deciduous forest, dry forest, riverine forest (herbarium). In Ghana, it is found in evergreen and in moist semi-deciduous forest (Hall & Swaine 1981). In Liberia, probably more common in drier forest types (Voorhoeve 1965).

Habitat

Description

It seems to prefer well-drained sites (Voorhoeve 1965), but is often found at river borders, creeks, and lagoons that can be periodically inundated (herbarium). The basal area (large trees) is greatest in flat, disturbed sites (Hawthorne 1995a).

Guild: sb Life form: large tree Max. height: 40 m (Voorhoeve 1965) Max. diameter: 117 cm (inventory data Ghana) Leaf: alternate, paripinnately compound, with 6-16 leaflets, oblong to elliptic, microphyll (0.8-3 x 1.5-5 cm), entire, coriaceous, glossy above, dull green beneath, new leaves flush red in the dry season Inflorescence: axillary or terminal, unbranched (spike), pale green and flat Flower: small; in two series along the puberulous peduncle; corolla white; fragrant Fruit: dry dehiscent (pod), thick, elliptic (3 x 4 cm), woody, reddish; 1 seed Seed: flat, very large (1.5 x 2.5 cm), black with red aril Other: a buttressed tree. The bark is rough, the slash orange to pinkish brown, very hard and

spp

184

n

Open (n)

Uses The bark is stripped from the tree, ground, and used as a perfume (Voorhoeve 1965).

Regeneration

Data sources

It has a phanerocotylar epigeal reserve seedling type (cf. Voorhoeve 1965). Seedlings of all sizes are, generally, very common in the shaded understorey around mother trees, and even in only small gaps saplings and older trees can be found. In Ghana, there is only a low density of adults, but very high densities of seedlings, indicating that, clearly, the mortality of trees of all size classes is very high (Hawthorne 1995a).

Taylor (1960), Savill & Fox (1967), Voorhoeve (1965), Hall & Swaine (1981), Hawthorne (1995a), IUCN Red List (2000), Hawthorne & Jongkind (2004)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

C.s.

23

0 (13)

4

43

0

9

39

34

17

26

52

22

43

17

H 26

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 185

Cordia vignei

Hutch. & Dalziel

Boraginaceae Description

Phenology

Guild: pi Life form: shrub or small tree Max. height: 6 m (herbarium) Max. diameter: data unavailable Leaf: alternate, simple, oblong to lanceolate, mesophyll (5 x 20 cm), entire, lower surface with long orange hairs, upper surface with white spots Inflorescence: branched Flower: medium-sized; calyx green; corolla white with reflexed lobes; fragrant Fruit: fleshy, ovoid (1.5 x 2 cm), yellow-green; 1-4 seeds Seed: data unavailable Other: the young twigs are densely hairy. The leaf margins are very often rugged due to insect attack.

Data sources

Distribution

FWTA, Mabberley (1987), Hawthorne (1990), Hawthorne & Jongkind (2004)

C

Continent: Nigeria Upper Guinea: Sierra Leone, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, in Upper Guinea present in 8 30’ cells, distribution range is 2292 km Forest type: disturbed forest, savanna

Habitat It is found in very disturbed forest, fallow vegetation, and thickets.

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

C.v.

12

60 (5)

8

50

8

50

8

0

41

0

58

42

17

0

58

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

185

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Costus deistelii

Page 186

K.Schum.

Zingiberaceae Habitat Species occurrence increases significantly with rainfall to reach an optimum around 2500 mm/yr (logistic regression analysis, Chi2 test). It is frequently reported in relatively open places (e.g. forest gaps, paths, and edges, and secondary forests) (De Rouw 1991), but also in deep humid forests. Usually in very moist conditions (e.g. along rivers, damp forest creeks, and swampy valleys). On clayish or sandy soils (herbarium).

C

Phenology Deciduousness: evergreen

Description

Data sources

Guild: np Life form: large herb Max. height: 2.5 m (herbarium) Leaf: spirally, simple, obovate to elliptic, mesophyll (3-7 x 9-21 cm), entire, coriaceous, glabrous Inflorescence: terminal, cylindrical Flower: corolla yellow to orange, tinged red at the base of the petals Fruit: data unavailable Seed: data unavailable

spp

186

n

Open (n)

C.d.

29

32 (19)

All

46333

37

FWTA, De Rouw (1991)

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 19 30’ cells, distribution range is 1421 km Forest type: wet evergreen forest, secondary forest

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

14

38

3

0

41

34

23

7

69

24

52

7

17

4

39

7

37

29

24

10

6

69

25

36

13

39

H

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Page 187

Crossostemma laurifolium

Planch. ex Benth.

Passifloraceae Description

Regeneration

Guild: pi Life form: large woody climber Max. height: 20 m long (herbarium) Max. diameter: data unavailable Leaf: alternate, simple, elliptic to obovate, notophyll (3.5-5.5 x 7.5-16 cm) or palmately lobed, entire, coriaceous; with stipules Inflorescence: axillary, branched, 2-6 flowers Flower: hermaphrodite; medium-sized; corolla white and yellow Fruit: dry dehiscent, ellipsoid (3 x 8 cm), orange, smooth, opening with 3 valves; many seeds Seed: medium-sized (0.9 x 0.6 x 0.4 cm), with an aril Other: it climbs with tendrils. Possibly starts as a shrub when young.

It has a phanerocotylar epigeal foliaceous seedling type (De Koning 1983).

C

Phenology Dispersal: probably by animals

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 33 30’ cell, distribution range is 1305 km Forest type: montane evergreen forest, wet evergreen forest, moist evergreen forest, secondary forest (herbarium). In Côte d’Ivoire, it is often found in coastal forests (Kasparek 2000).

Data sources De Koning (1983), Kasparek (2000)

Habitat Typically, found in open places of disturbed evergreen forests (e.g. forest gaps, edges, roadsides) and even degraded forests. Often in moist places (e.g. near lagoons or rivers). Soils can be clayish, sandy or rocky (herbarium). Species occurrence is associated to base-poor soils (logistic regression analysis).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

C.l.

63

50 (46)

3

48

6

22

31

36

10

16

67

17

25

19

H 52

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

187

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Page 188

Culcasia glandulosa

Hepper

Araceae Description

Phenology

Guild: sb Life form: small herbaceous root climber (hemiepiphyte) Max. height: 10 m length Leaf: alternate, simple, elliptic, notophyll (3-7 x 7-14 cm), entire, herbaceous, lower surface with dark strikes Inflorescence: spike of flowers on a swollen fleshy axis (spadix) 3-4 cm long, flowers unisexual Flower: spadix pinkish green; top male; centre pale yellow and sterile; bottom pink female; spatha glossy waxy pale brown Fruit: fleshy, globose, orange to red Seed: medium-sized (approx. 0.5 cm in diameter) Other: it has tough green stems clasping the bole with pinkish furry roots.

C

Data sources FWTA, De Rouw (1991)

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone (FWTA), Liberia, Côte d’Ivoire (herbarium) Distribution type: Continuous distribution, widespread, present in 12 30’ cells, distribution range is 871 km Forest type: evergreen forests, riverine forest, secondary forests

Habitat Species occurrence increases significantly with rainfall (logistic regression). Most often found where rainfall is between 1500-2000 mm/yr (Chi2 test), near rivers and creeks (Chi2 test), and in the forest understorey (herbarium, De Rouw 1991).

spp

188

n

Open (n)

C.g.

25

0 (14)

All

46333

37

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

L

M

H

L

M

H

0

72

4

4

52

28

16

0

96

4

68

4

20

39

7

37

29

24

10

6

69

25

36

13

39

4

Soil CMK

Soil WHC

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Page 189

Cussonia bancoensis

Aubrév. & Pellegr.

Araliaceae Description

most abundant in disturbed evergreen forests and in rocky or mildly sloping sites (Hawthorne 1995a). Sometimes found as spare trees in plantations (herbarium).

Guild: pi Life form: large tree Max. height: 40 m (FWTA) Max diameter: 90 cm (FWTA) Leaf: alternate, digitally compound, 5-9 leaflets, elliptic, mesophyll (5-8 x 11-21 cm), entire, glabrous; petiole up to 1 m long; stipules 3-7 mm long, persistent, triangular Inflorescence: clustered at the end of the branches, not branched (spike) Flower: small, whitish; sessile; glabrous Fruit: fleshy, ellipsoid (0.5 x 0.7 cm), red, borne on leafless branches Seed: medium-sized (0.6 x 0.35 x 0.25 cm) Other: it has a thick bark with deep fissures.

C

Regeneration It has a phanerocotylar epigeal foliaceous seedling type (cf. de la Mensbruge 1966).

Phenology

Distribution Continent: Nigeria (herbarium), Cameroon (uncertain record, according to Bamps, 1974) Upper Guinea: Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 8 30’ cells, distribution range is 1461 km, Red List species (vulnerable) Forest type: wet evergreen forest, moist evergreen forest, moist semi-deciduous forest, dry semideciduous forest, secondary forest

Uses The wood is used to make drums (Hawthorne 1995a).

Habitat

Data sources

Thought by Aubréville (1959) and De Koning (1983) to have been planted in Banco, Côte d’Ivoire and not to occur there naturally, possibly having survived in the forest from a long-since abandoned farm or village (De Koning 1983). They also noted that this species has been collected mostly in Ghana. Even in Ghana, however, this tree is not common, but the species regenerates freely in many areas (e.g. Atewa Forest Reserve) (Hawthorne 1995a). It is

spp

n

Open (n)

FWTA, Aubréville (1959), De la Mensbruge (1966), Bamps (1974), De Koning (1983), Hawthorne (1995a), IUCN Red List (2000), Kasparek (2000), Hawthorne & Jongkind (2004)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

C.b.

12

29 (7)

8

58

8

33

41

25

0

17

25

58

8

17

H 58

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

189

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Page 190

Cynometra ananta

Hutch. & Dalziel

Leguminosae-Caes. sapwood has ripple marks and the wood density is 0.96 g/cm3.

canopy is interrupted by only small gaps. The regenerating trees of the species have relatively exposed average crown scores, which is in line with the greater density of smaller trees in logged forest but is rather surprising considering the abundance of small seedlings and saplings in the shade (Hawthorne 1995a).

Distribution

C

Continent: Upper Guinea endemic Upper Guinea: Liberia, Côte d’Ivoire, Ghana. Distribution type: continuous, widespread, present in 12 30’ cells, distribution range is 952 km Forest type: wet evergreen forest, moist evergreen forest. In Ghana and Liberia, it is found predominantly in wet evergreen forests (Hall & Swaine 1981, Voorhoeve 1965).

Phenology Deciduousness: evergreen, leaf flushes during March to May and August to September (Taylor in Hawthorne 1995a) Dispersal: explosive Timing: flowering period from September to November (Liberia, Voorhoeve 1965); fruiting period from December to January (Hawthorne 1995a)

Habitat Species occurrence is higher where rainfall is between 1500-2500 mm/yr (Chi2 test). In Liberia, however, the abundance increases sharply above an annual rainfall higher than 2500 mm, and a dry period shorter than 4 months (Bongers et al. 1999). Experiments have shown that it is not especially sensitive to drought and does not show a strong preference for base-poor soils typical of wet evergreen forests (Swaine & Veenenedaal 1994). In Liberia and Côte d’Ivoire, it forms gregarious stands on the slopes towards creeks, in valleys, and on plateaux (Voorhoeve 1965, C. Chatelain, pers. comm.). On sandy soils (herbarium).

Description Guild: sb Life form: large tree Max. height: 45 m (herbarium) Max. diameter: 120 cm (Voorhoeve 1965) Leaf: alternate, paripinnately compound, 2 leaflets, sickle-shaped, microphyll (1-4 x 3.5-10 cm), entire, coriaceous, glabrous, bright red when newly flushed Inflorescence: axillary or terminal, branched, up to 7 panicles Flower: small; corolla white to pale grey, bowlshaped, densely crowded Fruit: dry dehiscent (pod), flat, oblong (4 x 9 cm), pale brown; 1-2 seeds, explosive Seed: disc-shaped, very large (2.2 x 2 cm), light brown Other: it has a straight bole becoming orange with age, and thin buttresses creeping along the ground. The bole is forked at a low height and the crown is large and spreading. The leaf flush is red. The slash is reddish over yellow, hard fibrous, or leathery. The

spp

190

n

Open (n)

Regeneration

Uses

It has a phanerocotylar epigeal reserve seedling type (cf. Voorhoeve 1965) and regenerates in shade (Hall & Swaine 1981). Seedlings are locally very abundant near the mother trees. In Liberia, saplings grow rarely larger than 2 m (Voorhoeve 1965). In Ghana, it is locally common and often gregarious (Hawthorne 1995a). Stands dominated by older saplings are apparently not uncommon where the

A timber tree (Hall & Swaine 1981).

Data sources Voorhoeve (1965), Hall & Swaine (1981), Swaine & Veenendaal (1994), Hawthorne (1995a), Bongers et al. (1999), Hawthorne & Jongkind (2004)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

C.a.

23

8 (13)

4

61

0

0

56

39

4

4

74

22

17

0

74

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 191

Cynometra leonensis

Hutch. & Dalziel

Leguminosae-Caes. Description

Regeneration

Guild: np Life form: medium-sized tree Max. height: 30 m (Voorhoeve 1965) Max. diameter: 100 cm (Voorhoeve 1965) Leaf: alternate, paripinnately compound, 6-12 leaflets, narrowly oblong, microphyll (0.4-1.8 x 0.8-5 cm), entire, fairly coriaceous, rachis slightly winged with few or no hairs; stipules small, persistent Inflorescence: axillary or terminal, branched, pale brown, tomentose Flower: small; corolla white; fragrant Fruit: dry dehiscent (pod), flat (7 x 3 cm), woody; 1-2 seeds Seed: very large Other: the tree is usually forked at a low height, with a spreading, dense crown. The stem is buttressed, and has a pale grey-brown and smooth bark. The twigs are lenticellate, with brown hairs on young parts only.

It has a phanerocotylar epigeal reserve seedling type (cf. Voorhoeve 1965). Seedlings are common in the forest (Voorhoeve 1965), but experience a high mortality in dense shade (Savill & Fox 1967).

C

Phenology Deciduousness: briefly deciduous (Savill & Fox 1967) Dispersal: explosive pods eject the seeds up to 30 m from the mother tree. Young pods are eaten by monkeys and squirrels (Savill & Fox 1967). Timing: flowering period from March to May (Savill & Fox 1967); fruiting period from April to June (Savill & Fox 1967)

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia Distribution type: continuous, local, present in 8 30’ cells, distribution range is 231 km Forest type: rainforest, secondary forest

Habitat

Data sources

It is very common and typical of riverbanks, where it may be dominant with Monopetalanthus pteridophyllus (Voorhoeve 1965). Cynometra is an associate of Heritiera utilis on the well-drained soils of Gola West in Sierra Leone, where the species dominate the upper canopy (Savill & Fox 1967).

spp

n

Open (n)

Voorhoeve (1965), Savill & Fox (1967), Hawthorne & Jongkind (2004)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

C.l.

14

75 (4)

0

43

0

0

0

0

93

0

86

14

86

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

191

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Page 192

Cyrtorchis hamata

Schltr.

Orchidaceae Distribution Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire, Ghana, Togo Distribution type: continuous, widespread, present in 7 30’ cells, distribution range is 816 km Forest type: unknown

C

Habitat Natural habitat unknown. Only few records from gardens or mango trees in town (herbarium).

Phenology Deciduousness: evergreen Dispersal: by wind

Description Guild: u Life form: epiphytic herb Max. height: data unavailable Leaf: alternate, simple, notophyll (1.5-4 cm x 9-24 cm), succulent Inflorescence: axillary, branched (panicle), lax, (5-17 cm long) Flower: medium-sized, white to yellowish, sweet scented, spur typically curved backwards Fruit: dry dehiscent (capsule), elliptic, brown Seed: very small

spp

192

n

Open (n)

Data sources FWTA

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

C.h.

9

- (0)

0

11

0

77

22

0

0

0

33

67

0

11

33

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 189-198

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Page 193

Dactyladenia dinklagei

(Engl.) Prance & F.White

Chrysobalanaceae Description

Phenology

Guild: sb Life form: small tree Max. height: 10 m (herbarium) Max. diameter: 10 cm (herbarium) Leaf: alternate, simple, ovate, notophyll (3.5-6 x 7-15 cm), entire, coriaceous, conspicuous spots along veins, glands by midrib on lower leaf surface, mainly at the leaf base, persistent stipules Inflorescence: axillary, branched (panicle) Flower: small; white, hairy Fruit: fleshy, heart-shaped (2.5 x 2.7 cm), hard, orange-brown, velutinous; 1 seed Seed: very large (2.3 x 1.3 x 1.3 cm) Other: the bole is cylindrical and greyish green.

D

Data sources

Distribution

FWTA, Hawthorne & Jongkind (2004)

Continent: Upper Guinea endemic Upper Guinea: Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 9 30’ cells, distribution range is 1153 km Forest type: rainforest, secondary forest

Habitat It occurs in the understorey, often close to water (e.g. riverbanks) (Hawthorne & Jongkind 2004)

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

D.d.

9

50 (2)

0

33

22

33

22

33

11

22

56

22

22

11

H 44

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

193

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Page 194

Dactyladenia smeathmannii

(Baill.) Prance & F.White

Chrysobalanaceae

D

Habitat Found along secondary roads, riverbanks, and the sea-side (herbarium, Hawthorne & Jongkind 2004).

Description

Phenology

Guild: u Life form: woody climber Max. height: data unavailable Max. diameter: data unavailable Leaf: alternate, simple, elliptic, notophyll (2.5-5 x 6-10 cm), entire, coriaceous Inflorescence: axillary, unbranched Flower: small; with a calyx tube and protruded long filament tube Fruit: fleshy; 1 seed Seed: very large Other: it starts probably as a shrub.

spp

194

n

Open (n)

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia Distribution type: continuous, widespread, present in 4 30’ cells, distribution range is 720 km Forest type: secondary forest

Data sources FWTA, Hawthorne & Jongkind (2004)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC

D.s.

6

50 (2)

0

50

0

0

0

17

83

0

83

17

All

46333

37

4

39

7

37

29

24

10

6

69

25

L 83 36

M

H

17

0

13

39

22466 layout page 189-198

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Page 195

Dactyladenia whytei

(Stapf ) Prance & White

Chrysobalanaceae Description

Phenology

Guild: np Life form: woody climber Max. height: 11 m long (herbarium) Max. diameter: 4 cm (herbarium) Leaf: alternate, simple, elliptic, notophyll (2.5-5.5 x 6-14 cm), entire, coriaceous, hairy midrib, stipules Inflorescence: axillary, unbranched, compact heads on peduncle Flower: medium-sized; calyx light green; corolla whitish inside; long protruded filament tube Fruit: fleshy, heart-shaped (2.5 x 4 cm), brown, hairy; 1 seed Seed: very large (2.5 x 1.5 x 1.5 cm) Other: it starts as a shrub. The twigs are pubescent.

D

Data sources FWTA, Hawthorne & Jongkind (2004)

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia (herbarium) Distribution type: continuous, regional, present in 16 30’ cells, distribution range is 418 km Forest type: rainforest, coastal forest, secondary forest

Habitat Species occurrence increases very sharply with rainfall higher than 2700 mm/yr (logistic regression analysis, Chi2 test). It can be found both under forest shade but also along paths, roads, and forest edges. In open places, it is sometimes found under the shade of taller trees. Often on soils with low water holding capacity (Chi2 test) such as white sands (herbarium).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

D.w.

32

67 (12)

0

44

13

0

0

3

89

0

91

9

69

9

6

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

195

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Page 196

Deinbollia voltensis

Hutch. ex Burtt Davy & Hoyle (nom. nud.)

Sapindaceae Description

Phenology

Guild: u Life form: shrub Max. height: 0.6 m (herbarium) Max. diameter: data unavailable Leaf: alternate, paripinnately compound, 4-6 leaflets, elliptic to oblanceolate, microphyll (2-3.2 x 6.5-9 cm), entire, herbaceous, winged rachis Inflorescence: terminal, branched (panicle) Flower: small, white Fruit: fleshy, globose (0.8 cm in diameter), reddish orange Seed: data unavailable

D

Distribution

Data sources

Continent: Upper Guinea endemic Upper Guinea: Ghana Distribution type: continuous, very local, present in 1 30’ cell. It has only been recorded near the Volta River and is possibly extinct. Forest type: gallery forest

FWTA, Hawthorne & Jongkind (2004)

Habitat Found on seasonally flooded grasslands (Hawthorne & Jongkind 2004).

spp

196

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

D.v.

3

- (0)

0

0

0

100

0

0

0

100

0

0

0

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 189-198

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Page 197

Dennettia tripetala

Baker f.

Annonaceae Description

Phenology

Guild: sb Life form: small tree Max. height: 15 m (FWTA) Max. diameter: data unavailable Leaf: alternate, simple, oblong to elliptic, mesophyll (4-6 x 10-15 cm), entire, coriaceous, peppered with minute gland spots below Inflorescence: cauliflorous, solitary Flower: medium-sized (2 cm long); petals dark wine-red with cream-coloured centre; bowl-shaped Fruit: fleshy (1.5 x 1 cm), reddish; 3-5 seeds Seed: medium-sized to large (1 cm long) Other: the slash is strongly and distinctively aromatic. Young twigs are black with diamondshaped narrow channels.

Deciduousness: evergreen (Hawthorne 1995a) Dispersal: probably by animals

Distribution

Uses

Continent: Nigeria, Cameroon Upper Guinea: Sierra Leone, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 13 30’ cells, distribution range is 2534 km, Red List species (vulnerable) Forest type: dry forest, dry semi-deciduous forest, gallery forest (Hawthorne 1995a, Hall & Swaine 1981). In Ghana, it is found mainly in the south marginal forest type, but also in dry semi-deciduous forests, and in the Southeast-outlier forest type (Hall & Swaine 1981). In Côte d’Ivoire, it has been found in open forests of the rainforest region, in very degraded forests, and in gallery forests (herbarium).

The fruits, known as pepper fruits, are edible and appreciated for their fragrance (Osisiogu 1975). The oil from the seed is toxic to insects and has a high antifungal activity (Ejechi et al. 1999). It can, therefore, be used as an insecticide or as a natural preservative in food processing.

D

Data sources FWTA, Rendle (1913), Osisiogu (1975), Hall & Swaine (1981), Hawthorne (1995a), Cable & Cheek (1998), Ejechi et al. (1999), IUCN Red List (2000)

Habitat The species occurs mainly in places where rainfall is below 1500 mm/yr (Chi2 test). Usually not close to large rivers (Chi2 test), but it is clearly associated with small streams in the savanna zone. It occurs on shallow rocky soils (Hall & Swaine 1981, herbarium).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

D.t.

16

27 (11)

6

6

6

75

19

6

0

0

75

25

25

19

H 56

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

197

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Page 198

Dialium aubrevillei

Pellegr.

Leguminosae-Caes. Regeneration It has a phanerocotylar epigeal reserve seedling type (cf. Voorhoeve 1965) and regenerates in shade (Hall & Swaine 1981). Seedlings and saplings are common in deep shade in the vicinity of parental trees. Regeneration is higher in “undisturbed” forest than in logged forest (Hawthorne 1995a).

D

Phenology Deciduousness: evergreen Dispersal: probably dispersed by animals (Hawthorne 1995a), fruits are eaten by monkeys (herbarium) Timing: fruiting period around the end of the dry season (Hawthorne 1995a)

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 32 30’ cells, distribution range is 1204 km Forest type: upland evergreen forest, wet evergreen forest, moist evergreen forest, moist semi-deciduous forest, secondary forest. In Ghana, the species is most common in evergreen forests (Hall & Swaine 1981).

Description Guild: sb Life form: large tree Max. height: 65 m (herbarium) Max. diameter: 100 cm (herbarium) Leaf: alternate, imparipinnately compound, up to 5 alternate leaflets, obovate, notophyll (2-5 x 4-11 cm), recurved, herbaceous, glabrous Inflorescence: axillary or terminal, branched Flower: small; creamy-white Fruit: fleshy (2 x 1.5 cm), black with an orange, sweet tasting pulp; 1-2 seeds Seed: medium-sized (0.9 x 0.7 x 0.3 cm), glossy brown Other: the bole is often slightly fluted, usually with large high buttresses. The bark has large plate scales and the slash exudes a red latex. The tree has a very small, half-globular crown.

spp

198

n

Open (n)

Habitat

Uses

More often present where rainfall is between 15002000 mm/yr (Chi2 test). It is usually found on uplands (Chi2 test): mostly on hills, less frequent in valleys, and avoiding swamps (Taylor 1960). In Ghana, it is very significantly associated with basepoor acid soils of high clay content (Hall & Swaine 1981). Reported also on slopes with rocky eroded soils (herbarium).

The timber is very hard and not used (Hall & Swaine 1981).

Data sources Taylor (1960), Voorhoeve (1965), Hall & Swaine (1981), Hawthorne (1995a), Hawthorne & Jongkind (2004)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

D.a.

47

10 (31)

4

21

2

11

71

10

8

2

68

30

43

2

H 45

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 199

Diaphananthe suborbicularis

Summerh.

Orchidaceae Description

Data sources

Guild: u Life form: epiphytic herb Max. height: data unavailable Leaf: alternate, simple, linear to oblanceolate, microphyll (0.7-1.3 x 5-7.5 cm), succulent Inflorescence: axillary (up to 4.5 cm long), up to 6 flowered Flower: small, purplish or pinkish Fruit: dry dehiscent (capsule), elliptic, green-brown Seed: very small

FWTA, Summerhayes (1958)

D

Distribution Continent: Upper Guinea endemic Upper Guinea: Ghana Distribution type: continuous, very local, present in 1 30’ cell. Only known from the type specimen Westwood 158 from the Togo Plateau (Kpandu to Wurupong) Forest type: unknown

Habitat Natural habitat unknown. Only one record on a cocoa tree (herbarium).

Phenology Deciduousness: evergreen Dispersal: by wind

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

D.s.

1

- (0)

0

100

0

0

100

0

0

0

0

100

0

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

199

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Page 200

Dichapetalum albidum

A.Chev. ex Pellegr.

Dichapetalaceae

D

Description

Habitat

Guild: np Life form: large winding woody climber Max. height: 20 m long (herbarium) Max. diameter: 5 cm (Breteler 1973) Leaf: alternate, simple, elliptic to obovate, notophyll (2-6 x 5-16 cm), entire, herbaceous, dark green above, whitish beneath; with stipules Inflorescence: axillary, branched, small Flower: small; creamy; calyx hairy, the sepals are outside somewhat purplish; 5 petals, bilobed Fruit: fleshy, globose (1.5 cm in diameter), with mealy mesocarp, yellow to orange; up to 3 seeds Seed: obovoid, medium-sized (0.7 x 0.5 cm), brown Other: in the first stage, it grows as a shrub. The bark is whitish. It is usually confused with D. pallidum.

Species occurrence strongly increases with altitude (logistic regression analysis, Chi2 test), and is higher where rainfall is over 2000 mm/yr (Chi2 test). Often in rather humid places (e.g. near rivers or on marshy soil). On sandy clayish soils (herbarium).

Regeneration It has a cryptocotylar hypogeal reserve seedling type (cf. Breteler 1973).

Phenology Dispersal: most probably by birds

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire (Breteler 1973), Ghana (herbarium) Distribution type: continuous, widespread, present in 20 30’ cells, distribution range is 1475 km Forest type: montane forest, rainforest, semideciduous forest, gallery forest, secondary forest

Data sources Breteler (1973), Hall & Swaine (1981)

spp

200

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

D.a.

40

37 (19)

20

55

0

3

20

41

38

0

75

25

65

0

H 15

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 199-208

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Page 201

Dichapetalum barteri

Engl.

Dichapetalaceae Description

and old streambeds. Usually found on sandy soils (herbarium).

Guild: np Life form: medium-sized tree or shrub Max. height: 17 m (herbarium) Max. diameter: 14 cm (herbarium) Leaf: alternate, simple, elliptic, mesophyll (3-8 x 7-17 cm), entire, coriaceous, with 2 large glands at base lower surface Inflorescence: axillary, bowl-shaped, branched Flower: small; corolla white to creamy or light yellow, 5-merous, petals retuse to bilobed Fruit: fleshy, subglobose (2.5 cm in diameter), strongly tuberculate, yellow; 1 seed Seed: ellipsoid, large (1.5 x 1 cm), brown Other: a hardwood species with a dense bushy crown. The branchlets are hairy when young, later on purple, with many lenticels.

Regeneration It has a cryptocotylar hypogeal reserve seedling type (cf. Breteler 1973) and regenerates in shade (Hall & Swaine 1981) but occurs also in thickets and secondary forest (herbarium).

D

Phenology Dispersal: probably by animals Timing: flowering is not clearly seasonal (Hall & Swaine 1981)

Distribution Continent: Benin, Nigeria, Gabon, Democratic Republic of Congo, Congo (Brazzaville) Upper Guinea: Côte d’Ivoire, Ghana Distribution type: continuous, widespread, in Upper Guinea present in 3 30’ cells, distribution range is 3703 km Forest type: rainforest, dry semi-deciduous forest, dry forest, secondary forest, gallery forest, coastal shrubland and thickets. The species occurs in very dry forests, especially near the coast and savanna (Hawthorne & Jongkind 2004). In Ghana, it is mostly found in the southern marginal outlier forest (Hall & Swaine 1981).

Uses The leaves are toxic to goats (Nwude et al. 1977). The toxic compounds have potential use as insecticides.

Habitat

Data sources

Often growing as a shrub at the coast, roadsides, or thicket transition woodland, but sometimes also as undergrowth of dense forests such as gallery forests or rainforest (at the drier range). On slopes

spp

n

Open (n)

FWTA, Breteler (1973), Nwude et al. (1977), Hall & Swaine (1981), Hawthorne & Jongkind (2004)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

D.b.

9

86 (7)

0

11

56

100

0

0

0

0

78

22

0

0

100

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

201

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Page 202

Dichapetalum dictyospermum

Breteler

Dichapetalaceae

D

Description

Regeneration

Guild: np Life form: small winding woody climber Max. height: 2 m long (herbarium) Max. diameter: 2 cm (herbarium) Leaf: alternate, simple, elliptic, notophyll (2-5 x 4-15 cm), entire, herbaceous, glabrous; glands conspicuous or absent; stipules 1-4 mm long, falling off soon Inflorescence: axillary, branched (cyme, up to 20 flowers) Flower: small; calyx grey to bright green, hairy; corolla white to yellowish, glabrous; slightly perfumed Fruit: fleshy, obovoid (2.3 x 3.3 cm), yellow to orange, glabrous; 1-2 seeds Seed: ovoid, large (1.3 x 1.8 cm), brown Other: when young it occurs in the form of a shrub. The bark is dark brown-reddish with creamy or brown lenticels, peeling off.

It has a cryptocotylar hypogeal reserve seedling type (cf. De Koning 1983) and can regenerate abundantly (herbarium).

Phenology Dispersal: probably by animals Timing: flowering period from July to March; fruiting period from October to April (De Koning 1983)

Distribution Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire (herbarium, Breteler 1978) Distribution type: continuous, very local, present in 1 30’ cell, distribution range is 11 km Forest type: rainforest, secondary forest, swamp forest

Data sources Breteler (1978), De Koning (1983), Hawthorne & Jongkind (2004)

Habitat In the forest it is locally abundant. Usually on sandy soil (herbarium).

spp

202

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

D.d.

53

38 (24)

0

92

0

0

0

100

0

2

98

0

0

2

98

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 203

Dichapetalum filicaule

Breteler

Dichapetalaceae Description Guild: np Life form: large winding woody climber Max. height: 20 m long (herbarium) Max. diameter: 1 cm (herbarium, Breteler 1978) Leaf: alternate, simple, ovate to elliptic, notophyll (1-5 x 3-13 cm), entire, herbaceous, with prominent nerves beneath; with small stipules (1-3.5 mm long) Inflorescence: axillary, branched, with up to 10 flowers Flower: small; corolla bowl-shaped, pale yellow to bright brownish green, 5-merous Fruit: fleshy, obovoid to ellipsoid (2 x 3 cm), smooth, mesocarp sweet and juicy, orange; 1-2 seeds Seed: ovoid, large (1.7 x 0.7 cm), brown Other: it has a shrubby habit when young or when growing in thickets. Later in life it occurs as a slender climber. The branches have lenticels.

D

Phenology Dispersal: by animals (Breteler 1973) Timing: flowering period from September to December and May to June; fruiting period from December to January (De Koning 1983)

Distribution

spp

Continent: Upper Guinea endemic (Breteler 1978) Upper Guinea: Côte d’Ivoire, Ghana (herbarium, Breteler 1978) Distribution type: continuous, widespread, present in 11 30’ cells Forest type: wet evergreen forest, moist evergreen forest, moist semi-deciduous forest, dry semideciduous forest, secondary forest. In Ghana, mainly found in moist evergreen forest (Hall & Swaine 1981). Locally abundant.

different stadia (open clearcuts to very old ones). Also at roadsides, around houses, and sometimes, on dry and very eroded hillsides. Usually on sandy or sandy-clayish but also on lateritic soils.

Habitat

Regeneration

Species occurrence increases at low altitudes (logistic regression analysis). It is mostly found in areas where rainfall is around 1500-2500 mm/yr (Chi2 test), and closer to rivers or lagoons (Chi2 test). Frequently found in secondary forests of

It has a hypogeal germination, cotyledons remain enclosed in the endocarp (Breteler 1973), and it regenerates in shade (Hall & Swaine 1981). However, most herbarium records come from very open areas (herbarium).

n

Open (n)

Data sources Breteler (1973, 1978), Hall & Swaine (1981), De Koning (1983)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

D.f.

63

77 (31)

0

63

10

3

40

57

0

27

65

8

8

25

59

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

203

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Page 204

Dichapetalum toxicarium

(G.Don) Baill.

Dichapetalaceae village forests, disturbed areas, and roadsides. It is usually found in moist places (e.g. river borders and banks). It can grow on a range of soil types (e.g. lateritic, sandy clay, stony sand, and sand). In Ghana, it is highly correlated with base-poor soils (Hall & Swaine 1981).

Regeneration

D

It has a hypogeal germination, cotyledons remain enclosed in the endocarp (Breteler 1982), and it regenerates in shade (Hall & Swaine 1981).

Description

Phenology

Guild: np Life form: large winding woody climber Max height: 100 m long (herbarium) Max. diameter: 12 cm (herbarium, Breteler 1982) Leaf: alternate, simple, elliptic, mesophyll (2-10 x 5-25 cm), entire, coriaceous, with distinct extrafloral nectaries usually on both sides Inflorescence: axillary, sub-umbellate Flower: small; corolla white, bowl-shaped; peduncle usually fused with petiole; perfumed Fruit: fleshy (3 x 2 cm), juicy, yellow to orange; 1-3 seeds Seed: large (1.8 x 1 cm) Other: it has a shrub habit when young or when growing in thickets. It has a characteristic deeply 5-lobed woody cylinder and is sparsely branched.

Dispersal: probably by animals

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea Bissau, Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 52 30’ cells, distribution range is 1594 km Forest type: wet evergreen, moist evergreen, riverine forest, moist semi-deciduous forest, secondary

spp

204

n

Open (n)

forest (herbarium). In Ghana, most common in wet and moist evergreen forests (Hall & Swaine 1981).

Uses

Habitat

The sprouts, fruits and seeds contain the toxic compound monofluoroacetate, and are very poisonous (Breteler 1982, Hall & Swaine 1981).

Species occurrence increases at lower altitudes (logistic regression analysis) and with increasing rainfall to reach a wide optimum range between 2000-3000 mm/yr (logistic regression analysis, Chi2 test). It grows in a wide range of conditions such as dense forests, forests actively exploited for timber,

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

D.t.

142

62 (63)

2

45

11

6

47

20

25

6

69

25

42

6

H 43

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

Data sources Breteler (1973, 1982), Hall & Swaine (1981)

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Page 205

Didelotia engleri

Dinkl. & Harms

Leguminosae-Caes. Description

Phenology

Guild: u Life form: small tree Max. height: 10 m (herbarium) Max. diameter: data unavailable Leaf: alternate, paripinnately compound, 10-16 leaflets, ovate to asymmetric, nanophyll (0.9-1.2 x 2-3 cm), entire, coriaceous, rachis with small scalelike glands close to and above leaflets, and with hairs red brown and curly Inflorescence: axillary, unbranched (raceme), densely pubescent, branching racemes from the axils of very young and small leaves Flower: small; corolla dark-red; fragrant Fruit: dry dehiscent (pod), rhomboid to oblong (0.1 x 2.5 cm) (only young pods seen), with two distinct longitudinal veins Seed: data unavailable

D

Data sources FWTA, Oldeman (1964), Savill & Fox (1967), Lock (1989), Hawthorne & Jongkind (2004)

Distribution Continent: Upper Guinea endemic Upper Guinea: Liberia, Côte d’Ivoire Distribution type: present in 3 30’ cells Forest type: wet forest

Habitat A rare species of which few collections are known. It is found near the coast, or near rivers (herbarium).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK

D.e.

4

0 (1)

0

50

50

0

0

50

50

All

46333

37

4

39

7

37

29

24

10

L 0 6

Soil WHC M

H

L

M

H

100

0

50

0

0

69

25

36

13

39

205

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Didelotia idae

Page 206

Oldeman, de Wit & J.Léonard

Leguminosae-Caes. Regeneration

pendant, densely hairy Flower: small; corolla purplish red; with long stamens Fruit: dry dehiscent (pod), inside of the valves papillate (4 x 11 cm), yellowish brown; 3 seeds Seed: disc-shaped, large (1.4 x 1.8 cm), dark brown, smooth Other: the slash is red fibrous with a reddish and sticky exudate. An unbuttressed tree with a wood density of 0.64 g/cm3.

D

It has a phanerocotylar epigeal reserve seedling type (cf. Voorhoeve 1965). Seedlings are prolific after a seed year, and seem capable of withstanding the dense shade (Savill & Fox 1967).

Phenology Deciduousness: evergreen Timing: flowering does not occur annually (Savill & Fox 1967); fruiting period from October to November (Liberia, Voorhoeve 1965)

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 12 30’ cells, distribution range is 1034 km, Red List species (lower risk) Forest type: wet evergreen forest, coastal thicket. It is gregarious in parts of Sierra Leone, associated with Brachystegia leonensis and Heritiera utilis (Fox 1968). In Liberia, it grows scattered throughout the evergreen forests, often in the single dominant forests of Tetraberlinia tubmaniana (Voorhoeve 1965). In Ghana, it is rare (Hawthorne 1995a).

Description Guild: sb Life form: large tree Max. height: 45 m (Voorhoeve 1965) Max. diameter: 120 cm (Voorhoeve 1965) Leaf: alternate, simple, narrowly ovate, microphyll (1.5-6 x 2-10 cm), entire, coriaceous, twigs and nerves hairy, new leaves red, the leaves tend to grow in two rows along the slender twigs Inflorescence: axillary or terminal, branched,

spp

206

n

Open (n)

Habitat

Uses

Species occurrence increases with rainfall to reach an optimum around 2600 mm/yr (logistic regression analysis, Chi2 test). In Liberia, it is associated with evergreen forests growing on either flat valleys or steep rocky slopes but never on marshy soil (Voorhoeve 1965). In Sierra Leone, the species does not occur near swamps or water courses (Savill & Fox 1967). However, in Ghana, where it is a rare species, it seems to be found mainly on riverbanks and wet places of evergreen forests (Hawthorne 1995a, Hawthorne & Jongkind 2004).

Timber species (Voorhoeve 1965).

Data sources Oldeman (1964), Voorhoeve (1965), Savill & Fox (1967), Fox (1968), Lock (1989), Hawthorne (1995a), IUCN Red List (2000), Hawthorne & Jongkind (2004)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

D.i.

18

8 (13)

6

61

6

0

39

17

44

0

89

11

72

0

22

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 207

Diospyros chevalieri

De Wild.

Ebenaceae Description

Regeneration

Guild: sb Life form: shrub or pigmy tree Max. height: 5 m (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, oblong, mesophyll (6-11 x 14-27 cm), entire, herbaceous Inflorescence: axillary, solitary Flower: medium-sized; calyx brown velvet; corolla white, dish-shaped, dioecious Fruit: fleshy (berry) (2.5 x 4.2 cm), orange with orange hairs; few seeds surrounded by pulp Seed: strongly angular, very large (0.7 x 3 cm), brown Other: branching in whorls, with densely pubescent twigs.

It regenerates in the shade (Hall & Swaine 1981).

Phenology Dispersal: probably by animals

D

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 25 30’ cells, distribution range is 1076 km Forest type: wet evergreen forest, moist evergreen forest, coastal forest, secondary forest. In Ghana, most abundant in wet evergreen forests (Hall & Swaine 1981).

Data sources FWTA, White (1978), Hall & Swaine (1981), Hawthorne & Jongkind (2004)

Habitat Species occurrence increases significantly with rainfall to reach an optimum around 2500 mm/yr (logistic regression analysis). The species occurs mainly at lower altitudes (logistic regression analysis). It is frequently found in the understorey of dense mature forests, but it occurs in secondary forests as well. On clayish or lateritic soils.

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

H

D.c.

79

28 (53)

1

35

3

0

71

11

19

1

68

30

61

1

38

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

207

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Diospyros cooperi

Page 208

(Hutch. & Dalziel) F.White

Ebenaceae

D

Description

Regeneration

Guild: sb Life form: small to medium-sized tree Max. height: 13 m (herbarium) Max. diameter: 15 cm (herbarium) Leaf: alternate, simple, ovate, mesophyll (4.5 x 11 cm), entire, herbaceous Inflorescence: unbranched, 1-4 flowered, dioecious with unisexual flowers Flower: medium-sized; calyx green; corolla pale yellow, dish with tube Fruit: fleshy (berry), globose (approx. 1.5 cm in diameter), orange; 6-8 seeds Seed: large Other: it has a dark bark, branching in whorls. The slash is reddish.

It regenerates in the shade (Hall & Swaine 1981).

Phenology Dispersal: probably by animals

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 24 30’ cells, distribution range is 1126 km Forest type: rainforest, moist evergreen forest, moist semi-deciduous forest, secondary forest. In Ghana, mostly in moist evergreen forest and only very scarcely present in moist semi-deciduous forest (Hall & Swaine 1981).

Data sources White (1978), Hall & Swaine (1981), Hawthorne & Jongkind (2004)

Habitat Species occurrence increases significantly with rainfall (logistic regression analysis, Chi2 test). It occurs mainly in the shaded understorey of mature evergreen forests, and only occasionally in secondary forests. Often near streams and rivers (herbarium).

spp

208

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

D.c.

35

19 (21)

3

34

3

6

43

18

35

3

83

14

83

3

H 11

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 209

Diospyros liberiensis

A.Chev. ex Hutch. & Dalziel

Ebenaceae Description

Phenology

Guild: pi Life form: small tree Max. height: 5 m (herbarium) Max. diameter: 5 cm (Hawthorne & Jongkind 2004) Leaf: alternate, simple, elliptic to ovate, mesophyll (3-11 x 7-22 cm), entire, herbaceous, pubescent beneath, large glandular spots Inflorescence: axillary, unbranched Flower: small unisexual flowers; male flower with 4 lobes; female flower unknown Fruit: fleshy (berry) (3.3 x 4 cm), velvet, yellow to orange or black; containing two parallel seeds in colourless pulp Seed: long, very large (2.6 x 1.8 x 0.9 cm), brownish black

Dispersal: probably by animals

Distribution

FWTA, White (1978), Hawthorne & Jongkind (2004)

D

Data sources

Continent: Upper Guinea endemic Upper Guinea: Liberia, Côte d’Ivoire Distribution type: continuous, local, present in 4 30’ cells, distribution range is 293 km Forest type: rainforest, secondary forest

Habitat It is typically found in disturbed habitats (e.g. along roadsides, in secondary forests, and under plantations) (herbarium). The species is mostly confined to the coastal zone.

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

D.l.

6

50 (4)

0

33

50

0

50

34

17

0

100

0

67

0

33

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

209

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Diospyros vignei

Page 210

F.White

Ebenaceae

Habitat

D

The species is most frequently found in the lowlands (logistic regression analysis), where rainfall is between 1500-2000 mm/yr (Chi2 test). It occurs in the understorey of mature forests and old secondary forests (herbarium).

Description

Regeneration

Guild: sb Life form: small tree Max. height: 10 m (herbarium) Max. diameter: 3 cm (herbarium) Leaf: alternate, simple, elliptic-lanceolate, notophyll (3-4 x 7-13 cm), entire, herbaceous, glabrous, very pale green beneath Inflorescence: male flowers axillary and cauliflorous; female flowers cauliflorous; dioecious; unbranched Flower: medium-sized; bell-shaped; corolla white, 3-lobed Fruit: fleshy (berry), conically shaped (2 x 3 cm), orange; up to 4 seeds in scarce whitish sweet pulp Seed: oblong, very large (2.5 x 0.9 x 0.6 cm), dark reddish brown Other: it has black bark and branches in whorls. All leaves of a single branch are in one plane. The twigs and the underside of the midribs are pubescent. The species is very similar to Diospyros heudelotii.

It regenerates in the shade (Hall & Swaine 1981).

Phenology Dispersal: probably by animals

Distribution

in 20 30’ cells, distribution range is 761 km Forest type: wet evergreen forest, moist evergreen forest, moist semi-deciduous forest, secondary forest. In Ghana mostly found in moist evergreen forest (Hall & Swaine 1981)

Continent: Upper Guinea endemic Upper Guinea: Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present

spp

210

n

Open (n)

Data sources FWTA, White (1963, 1978), Hall & Swaine (1981), Hawthorne (1995a)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

D.v.

37

34 (35)

3

24

3

19

56

24

0

5

65

30

35

8

57

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 211

Dissomeria crenata

Hook.f. ex Benth.

Flacourtiaceae Description

Phenology

Guild: u Life form: medium-sized tree Max. height: 12 m (FWTA) Max. diameter: 15 cm (herbarium) Leaf: alternate, simple, elliptic to obovate-elliptic, notophyll (4.5-5.5 x 8-13 cm), crenate, coriaceous, pale green beneath with pinkish midrib and nerves Inflorescence: axillary, not branched (spike) Flower: small to medium-sized; white Fruit: dry indehiscent; many seeds Seed: medium-sized (0.3 x 0.3 cm), brown Other: the young twigs are rough with conspicuous, dense but small, rounded lenticels.

Dispersal: probably by water (Hawthorne & Jongkind 2004)

D

Distribution Data sources

Continent: Nigeria Upper Guinea: Guinea, Sierra Leone, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 20 30’ cells, distribution range is 2202 km Forest type: gallery forest

FWTA, Keay (1989), Hawthorne (1996), Hawthorne & Jongkind (2004)

Habitat Most frequently found where rainfall is less than 1500 mm/yr (Chi2 test). It is exclusively confined to streams in the savanna region (Hawthorne 1996) and can be found along channels, riverbanks, and on river islands (herbarium).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

D.c.

30

40 (5)

0

43

3

74

10

7

9

7

63

30

47

13

17

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

211

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Page 212

Dorstenia astyanactis

Aké Assi

Moraceae Description

Phenology

Guild: u Life form: perennial, non-woody, succulent epiphyte Max. height: 1.5 m (Aké Assi 1967) Max. diameter: 1 cm (Aké Assi 1967) Leaf: alternate, simple, ovate, notophyll (3-7 x 5-13 cm), dentate, fleshy, glabrous, with glands; petiole 0.5-1 cm long Inflorescence: numerous male and female flowers packed on an open receptacle Flower: small (approx. 0.15 cm long) Fruit: fleshy, ovoid, small (0.1 cm in diameter), light brown Seed: data unavailable Other: it is a pendulous epiphyte.

D

Data sources

Distribution

FWTA, Aké Assi (1967), Hawthorne & Jongkind (2004)

Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire Distribution type: continuous, very local, present in 1 30’ cell. Collected only on Mt Tonkoui (Hawthorne & Jongkind 2004). Forest type: mountainous rainforest

Habitat It is reported to grow on large trees (herbarium).

spp

212

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

D.a.

3

- (0)

100

0

0

0

100

0

0

0

100

0

0

0

100

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 209-218

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Page 213

Dorstenia embergeri

Mangenot

Moraceae Description

Phenology

Guild: u Life form: perennial herb Max. height: 0.2 m (herbarium) Max. diameter: data unavailable Leaf: alternate, simple, oblanceolate, microphyll (2-4 x 3-11 cm), entire, herbaceous, hairy; with stipulate petiole (1.0-2.5 cm), pubescent Inflorescence: axillary or terminal, unisexual flowers Flower: green Fruit: fleshy, membranous, with 2 dehiscent valves, yellow; 2 seeds Seed: ovoid, small (0.2 cm long) Other: an erect herb.

D

Data sources Mangenot (1957), De Koning (1983), Hawthorne & Jongkind (2004)

Distribution Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 5 30’ cells, distribution range is 779 km. It is extremely rare (Mangenot 1957, De Koning 1983). Forest type: wet evergreen forest, riverine forest (herbarium)

Habitat It is practically confined to the understorey of very dense forests (Mangenot 1957), but occasionally found at forest margins. Often beside streams and in riverine forests (herbarium, Hawthorne & Jongkind 2004).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

D.e.

12

0 (6)

0

75

0

8

17

75

0

17

58

25

8

17

67

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

213

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Page 214

Dorstenia turbinata

Engl.

Moraceae Description

Phenology

Guild: sb Life form: small shrub or small tree Max. height: 4 m (herbarium) Max. diameter: 10 cm (Hawthorne 1996) Leaf: alternate, simple, oblanceolate, notophyll (2.5-4 x 8.5-15 cm), herbaceous to coriaceous Inflorescence: axillary, receptacle, female flowers solitary in the centre of the receptacle Flower: light green Fruit: fleshy (1 cm long), yellow, with 2-4 long stringlike horns; 2 seeds Seed: ovoid, medium-sized (0.5 cm in diameter) Other: the slash exudes a white latex.

D

Data sources

Distribution

FWTA, Hawthorne (1996), Hawthorne & Jongkind (2004)

Continent: Nigeria, Cameroon, Equatorial Guinea, Gabon, Democratic Republic of Congo Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continental disjunct, present in 33 30’ cells in Upper Guinea Forest type: wet and moist evergreen forest, old secondary forest, gallery forest (herbarium)

Habitat Species occurrence increases significantly with rainfall to reach an optimum around 2500 mm/yr. Frequently found under deep shade in the understorey of dense evergreen forests (Hawthorne & Jongkind 2004). It occurs in moist to marshy places. Often on sandy soils and sometimes on lateritic soils (herbarium).

spp

214

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

D.t.

61

12 (42)

7

41

3

3

41

24

32

2

72

26

46

3

H 44

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 215

Dracaena calocephala

Bos

Dracaenaceae Description

Phenology

Guild: sb Life form: rhizotomatic shrub Max. height: data unavailable Max. diameter: data unavailable Leaf: alternate, simple, obovate to oblong, mesophyll (3-7 x 18-51 cm), entire, coriaceous, with inrolled cuspidate tip Inflorescence: terminal, unbranched, globose (6 x 7 cm), densely flowered Flower: large (up to 5 cm long); white Fruit: fleshy, globose (1.5 x 2 cm), orange Seed: lack dry testa, embedded in fruit pulp Other: The rhizomatic stems give rise to unbranched short erect stems.

Deciduousness: evergreen Dispersal: probably by animals Timing: it appears not to have a fixed flowering period

D

Distribution

Data sources

Continent: Upper Guinea endemic Upper Guinea: Guinea, Liberia Distribution type: continuous, local, present in 6 30’ cells, distribution range is 253 km Forest type: wet forest, secondary forest

Bos (1984)

Habitat It occurs terrestrial in forest undergrowth, presumably in undisturbed forests (Bos, pers. comm.).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

D.c.

8

0 (3)

38

50

0

0

0

63

38

0

75

25

75

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

215

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Page 216

Dracaena cristula

W.Bull

Dracaenaceae Habitat Species occurrence increases with rainfall to reach an optimum around 2500 mm/yr (logistic regression analysis, Chi2 test). Often found at higher altitudes (Chi2 test). Usually found in the understorey of mature forests, near water (e.g. streams, waterfalls, lakes). Sometimes also in swampy forests in the savanna region (herbarium). It is frequently found on soils with high water holding capacity (Chi2 test).

D

Regeneration Presumably by seed only.

Phenology Deciduousness: evergreen Dispersal: probably by animals Timing: it does not appear to have a fixed flowering period

Description Guild: sb Life form: unbranched pigmy tree Max. height: 3 m (herbarium) Max. diameter: approx. 1 cm Leaf: alternate, simple, ovate to rounded, mesophyll (3-13 x 3-17 cm), entire, herbaceous to coriaceous, unfolded young leaves dark red; with a winged pseudo-petiole Inflorescence: terminal, unbranched, many flowers together Flower: medium-sized; white Fruit: fleshy, orange Seed: lens-shaped, medium-sized (0.8 x 0.7 x 0.4 cm), white Other: it has an unbranched single stem that is terminated by a pseudowhorl of leaves. After each flowering, growth is continued sympodially by a single axillary bud.

spp

216

n

Open (n)

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana (herbarium, Bos 1984) Distribution type: continuous, widespread, present in 33 30’ cells, distribution range is 1417 km Forest type: montane forest, wet evergreen forest, swamp forest, secondary forest

Data sources Bos (1984)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

D.c.

72

23 (35)

15

44

6

8

29

47

12

3

68

29

39

4

H 35

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 217

Dracaena ovata

Ker-Gawl.

Dracaenaceae Description Guild: sb Life form: shrub Max. height: 5 m (herbarium) Max. diameter: less than 2 cm Leaf: alternate, simple, obovate, mesophyll (2-9 x 8-28 cm), entire, coriaceous; pseudopetiole Inflorescence: terminal, not branched, approx. 5 cm long with crowded flowers; stems often clad in sheathing prophylls leaving angular scars Flower: large; creamy-white; fragrant Fruit: fleshy, subglobose (2 cm in diameter), orange, with orange pulp; 1-3 seeds Seed: lens-shaped, medium-sized (1.0 x 0.9 x 0.6 cm), white Other: the stem is terminated by a pseudowhorl of leaves.

D

Distribution Continent: Nigeria, Cameroon (Bos 1984) Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana (herbarium, Bos 1984) Distribution type: continental disjunct, in Upper Guinea present in 31 30’ cells. The species has a remarkable disjunct distribution, with only three collections found at the border between Nigeria and Cameroon (Bos 1984). Forest type: wet evergreen forests, mature forest, secondary forest

Phenology Deciduousness: evergreen Dispersal: probably by animals Timing: it does not appear to have a fixed flowering period

Data sources FWTA, De Koning (1983), Bos (1984)

Habitat Species occurrence decreases with altitude (logistic regression analysis, Chi2 test). It increases with annual rainfall, to reach an optimum at 2500 mm/yr (logistic regression analysis, Chi2 test). It is frequently found near streams, rivers, or lakes (herbarium). On sandy, clayish, rocky, and lateritic soils (herbarium)

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

D.o.

82

42 (45)

1

39

15

4

54

20

18

7

56

37

28

6

H 5

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

217

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Page 218

Dracaena phrynioides

Hook.

Dracaenaceae

Habitat

D

It is only known from undisturbed forest sites, where it grows terrestrial in the undergrowth (Bos, pers. comm.). Mainly in moist forests where rainfall is around 1500-2000 mm/yr (Chi2 test). Generally in wet shady places in the forest understorey (e.g. near streams and rivers). On soils with a large water holding capacity (logistic regression analysis, Chi2 test).

spp

218

Description

Phenology

Guild: sb Life form: perennial herb Max. height: 0.8 m (Bos 1984) Max. diameter: less than 1 cm Leaf: alternate, most leaves arising close to the base, simple, ovate to lanceolate, mesophyll (6-10 x 15-25 cm), entire, coriaceous, dark green with light green transversely oriented oval dots; long petiole Inflorescence: terminal, branched, in clusters of many flowers Flower: medium-sized; white, narrowly funnelshaped Fruit: fleshy (berry), flattened (1.5 cm in diameter), 3-lobed, orange to red, with an orange pulp Seed: medium-sized (0.8 x 0.7 x 0.4 cm), brown Other: the plant is unbranched and almost stemless. It has orange roots.

Deciduousness: evergreen Timing: it appears not to have a fixed flowering period

n

Open (n)

Distribution Continent: Nigeria (Bos 1984), Cameroon, Equatorial Guinea (herbarium), Fernando Po (Bos, 1984), Gabon (herbarium) Upper Guinea: Liberia, Côte d’Ivoire, Ghana (herbarium) Distribution type: continuous, widespread, in Upper Guinea the species is present in 21 30’ cells, distribution range is 2468 km Forest type: wet evergreen forest, moist evergreen forest, moist semi-deciduous forest, dry semideciduous forest, riverine forest

Data sources FWTA, Hall & Swaine (1981), Bos (1984)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

D.p.

30

7 (15)

10

33

3

10

73

17

0

3

37

60

20

0

H 60

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 219-228

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Page 219

Dracaena praetermissa

Bos

Dracaenaceae Description

Phenology

Guild: sb Life form: small shrublet Max. height: 0.5 m (herbarium) Leaf: spirally, simple, ovate, notophyll (2-6 x 5-19 cm), entire, coriaceous, with 4-6 sharply impressed nerves; pseudo-petiole Inflorescence: terminal, unbranched, small (< 5 cm), hidden under the leaves and not surpassing them Flower: small Fruit: fleshy, globose (1.5 cm in diameter), orange Seed: rhomboid, medium-sized (0.9 x 0.8 x 0.6 cm), pale brown Other: the rhizomes may be repeatedly forked. Arial stems erect up to 5 cm. It is usually mistaken for juvenile D. aubryana.

Deciduousness: evergreen Dispersal: probably by animals Timing: it does not appear to have a fixed flowering period

D

Distribution Data sources

Continent: Upper Guinea endemic to the Cavally river basin in SW Côte d’Ivoire Upper Guinea: Côte d’Ivoire, presumably also on the Liberian bank Distribution type: continuous, local, present in 3 30’ cells, distribution range is 194 km. Forest type: wet evergreen forest, secondary forest

Bos (1984), Hawthorne & Jongkind (2004)

Habitat It is found in deep shade.

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

D.p.

29

20 (5)

0

7

3

0

3

97

0

7

86

7

86

7

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

219

22466 layout page 219-228

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Dracaena scabra

Page 220

Bos

Dracaenaceae

D

Description Guild: np Life form: small tree Max. height: 7 m (herbarium, Bos 1984) Max. diameter: not over 3 cm Leaf: alternate, simple, obovate, macrophyll (6-9 x 40-110 cm), entire, coriaceous, smooth, slightly glossy, widened sheathing base Inflorescence: terminal, branched (erect scabrid panicle), with many flowers crowded together, green to purple Flower: medium-sized; white Fruit: fleshy, subglobose (2 cm in diameter), orange Seed: flat, bi-lobed, large (1.2 x 1.0 x 0.5 cm), pale brown Other: the plant is unbranched and single stemmed. It has stout stems with a terminal tuft of leaves. After flowering, growth is continued by an axillary bud.

spp

220

n

Open (n)

Phenology Deciduousness: evergreen Dispersal: probably by animals Timing: it appears not to have a fixed flowering period

Distribution Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire (herbarium, Bos 1984) Distribution type: continuous, local, present in 7 30’ cells, distribution range is 257 km Forest type: degraded forest, secondary forest, riverine forest, swamp forest

Habitat

Data sources

It seems to be confined to the high water marks of swamps and on riverbanks (Bos 1984).

Bos (1984)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

D.s.

13

40 (10)

0

46

0

8

93

0

0

8

38

54

0

8

85

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 219-228

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Page 221

Ehretia trachyphylla

C.H.Wright

Boraginaceae Description

Phenology

Guild: pi Life form: medium-sized tree Max. height: 20 m (Hawthorne & Jongkind 2004) Max. diameter: 30 cm (herbarium) Leaf: alternate, simple, elliptic, mesophyll (4-8 x 820 cm), entire, coriaceous, pubescent at lower surface Inflorescence: axillary or terminal, branched Flower: small; white Fruit: fleshy, narrowly obovoid (0.6 x 1 cm), white to greyish; 4 seeds Seed: medium-sized (0.3 x 0.3 cm) Other: it is low branching, often with epicormic shoots.

Timing: flowering period from April to August, fruiting period from May to September (De Koning 1983)

E

Distribution Data sources

Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire, Ghana Distribution type: continuous, regional, present in 10 30’ cells, distribution range is 555 km Forest type: evergreen forest, semi-deciduous evergreen forest, secondary forest

FWTA, Hall & Swaine (1981), De Koning (1983), Hawthorne (1990), Hawthorne & Jongkind (2004)

Habitat It occurs in mature and secondary forests, and sometimes along river borders (herbarium).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

E.t

17

11

(9) 0

41

6

47

35

18

0

6

41

53

24

6

H 59

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

221

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Page 222

Elytraria ivorensis

Dokosi

Acanthaceae Distribution Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 8 30’ cells, distribution range is 761 km Forest type: wet evergreen forest, old secondary forest (herbarium)

Habitat

E

Found in old forest gaps, paths, and edges. Usually found in moist shady places of the forest. It occurs on sandy clay or rich humus (herbarium).

Phenology Deciduousness: evergreen

Description Guild: sb Life form: perennial rosette-forming herb Max. height: 0.3 m (herbarium) Leaf: rosette, simple, obovate, microphyll (2-3 x 46 cm), entire, pubescent above, glabrous beneath Inflorescence: axillary, not branched Flower: small; white Fruit: dry dehiscent Seed: small (0.1 cm), brown

spp

222

n

Open (n)

Data sources FWTA, Dokosi (1970), Hawthorne & Jongkind (2004)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

E.i.

11

36 (11)

0

27

9

0

72

27

0

9

82

9

64

9

27

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 219-228

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Page 223

Elytraria maritima

J.K.Morton

Acanthaceae Description

Phenology

Guild: pi Life form: small rhizomatous annual herb Max. height: 0.15 m (herbarium) Max. diameter: data unavailable Leaf: rosette, simple, elliptic to obovate, microphyll (1.5-2.5 x 3-6 cm), entire, herbaceous, pubescent Inflorescence: terminal, not branched (slender pedunculate spike) Flower: small (corolla tube 3 mm long, lobes 2 mm long); white to pale lilac; calyx with 5 sepals Fruit: dry dehiscent, ovate (c. 0.6 cm x 0.15 cm), glabrous, brown; approx. 10 seeds Seed: small (0.05 x 0.03 cm); brown Other: a diffusely branched, prostrate herb with a bright reddish green stem.

E Data sources FWTA, De Koning (1983)

Distribution Continent: Benin (doubtful record) Upper Guinea: Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 3 30’ cells, distribution range is 706 km Forest type: secondary forest (herbarium).

Habitat It is found in disturbed areas (e.g. paths, roadsides, forest edges) usually under shade but occasionally in the open. Also found under coconut plantations at the coast. On sandy soils.

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

E.m.

15

67 (9)

0

67

0

7

14

80

0

40

47

13

0

20

53

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

223

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Page 224

Englerina gabonensis

(Engl.) S.Balle

Loranthaceae Distribution Continent: Nigeria, Cameroon, Central African Republic, Equatorial Guinea, Gabon, Congo (Brazzaville), Democratic Republic of Congo, Angola (Polhill & Wiens 1998, herbarium) Upper Guinea: Sierra Leone, Liberia (herbarium) Distribution type: continental disjunct, in Upper Guinea present in 4 30’ cells Forest type: riverine forest, gallery forest in savanna

E

Habitat It is found along rivers (herbarium), and is parasitic on Ficus and Platysepalum species (FWTA).

Phenology Dispersal: by birds

Description Guild: pi Life form: hemi-parasitic shrub Max. height: 2 m (Polhill & Wiens 1998) Max. diameter: data unavailable Leaf: opposite, simple, ovate, notophyll (2-6 x 7-16 cm), entire, coriaceous Inflorescence: pedunculate umbel, with 6-12 flowers standing up like candles Flower: large (corolla up to 4.8 cm); orange-yellow, sometimes banded red or brownish at or above base Fruit: fleshy (berry), globose (0.5 cm in diameter), red with a sticky pulp; 1 seed Seed: small (0.2 x 0.1 cm)

spp

224

n

Open (n)

Data sources FWTA, Polhill & Wiens (1998)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

E.g.

4

- (0)

25

50

0

0

0

25

75

0

100

0

100

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 219-228

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Page 225

Englerina parviflora

(Tiegh.) Balle

Loranthaceae Description

Phenology

Guild: pi Life form: hemi-parasitic shrub Max. height: 1 m (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, lanceolate to elliptic-oblong, microphyll (1.5-4 x 4-12 cm), entire, herbaceous Inflorescence: pedunculate umbel, with 8-16 flowers, standing up like candles Flower: medium-sized (corolla up to 2.2 cm); red Fruit: fleshy (berry), globose (0.5 cm in diameter), red or purple with a sticky pulp; 1 seed Seed: data unavailable

Dispersal: by birds Timing: flowering period from December to April (Polhill & Wiens 1998)

E

Distribution Continent: Upper Guinea endemic (Polhill & Wiens 1998) Upper Guinea: Guinea Bissau, Guinea, Sierra Leone, Liberia, Côte d’Ivoire (Polhill & Wiens 1998), Ghana (herbarium) Distribution type: continuous, widespread, present in 15 30’ cells, distribution range is 1714 km Forest type: data unavailable

Data sources FWTA, Polhill & Wiens (1998)

Habitat Data unavailable.

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

E.p.

22

- (0)

23

41

5

5

59

28

10

0

45

55

9

5

36

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

225

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Page 226

Euadenia eminens

Hook.f.

Capparaceae

E

Description

Ghana Distribution type: continental disjunct with a small population in Lower Guinea, in Upper Guinea present in 13 30’ cells Forest type: moist evergreen, moist semideciduous, dry semi-deciduous, secondary forest (herbarium). In Ghana, most common in moist evergreen and moist semi-deciduous forests (Hall & Swaine 1981).

Guild: sb Life form: pigmy tree Max. height: 3 m (Hawthorne & Jongkind 2004) Max. diameter: data unavailable Leaf: alternate, digitately compound, 3 leaflets, obovate, mesophyll (5-9 x 8-29 cm), herbaceous, upper and lower side glossy and with a purple midrib Inflorescence: not branched Flower: medium-sized; yellow-green, pale purple at the base Fruit: fleshy dehiscent (up to 1 x 30 cm), orange Seed: ellipsoid, medium-sized (0.5 x 1.0 cm) Other: an unbranched stem. The species is easily confused with E. trifoliolata, however, E. eminens can be distinguished based on its subsessile folioles (De Koning 1983).

Species occurrence tends to increase slightly with rainfall (logistic regression analysis) but the species is found mainly in moist to dry forests. It can be found in the understorey of dense shady forests. Usually on sandy soils. It is occasionally found at the coast (herbarium).

226

Open (n)

Data sources

Regeneration

Continent: Gabon (herbarium), but according to Hall & Swaine (1981) an Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire,

n

Deciduousness: deciduous (herbarium)

Habitat

Distribution

spp

Phenology

FWTA, Hall & Swaine (1981), De Koning (1983), Hawthorne (1995a), Hawthorne & Jongkind (2004)

It regenerates in shade (Hall & Swaine 1981).

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

E.e

14

14 (7)

0

57

14

14

43

29

7

14

50

36

21

0

H 57

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 227

Ficus ottoniifolia

(Miq.) Miq. ssp.

multinervia

C.C.Berg

Moraceae Description

Phenology

Guild: pi Life form: shrub or tree, starts epiphytic Max. height: 15 m (FWTA) Max. diameter: 15 cm (herbarium) Leaf: alternate, simple, elliptic, mesophyll (3-8 x 9-20 cm); coriaceous Inflorescence: cauliflorous, mainly on older branches, flowers in a closed receptacle Flower: very small Fruit: fleshy (fig), ellipsoid (1 x 1.5 cm), smooth, yellowish outside and reddish brown on cross section, many seeds Seed: small (0.1 x 0.1 cm) Other: the slash exudes a watery, whitish latex.

Dispersal: probably by animals (monkeys, birds, bats)

F

Distribution

Data sources

Continent: Cameroon, Gabon Upper Guinea: Senegal, Gambia, Guinea-Bissau, Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 14 30’ cells, distribution range is 1821 km Forest type: gallery forest

FWTA, De Rouw (1991)

Habitat Often found in very wet (e.g. creek and riverbanks, alluvial plain), open (e.g. forest edges, close to villages), and disturbed (secondary forests) places (De Rouw 1991).

Regeneration It regenerates in secondary forest (De Rouw 1991).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

F.o.

18

33 (3)

0

44

33

51

22

11

6

11

56

33

39

11

H 28

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

227

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Ficus scott-elliotii

Page 228

Mildbr. & Burrett.

Moraceae

F

Description

Phenology

Guild: u Life form: tree, starts epiphytic Max. height: 15 m (herbarium) Max. diameter: 50 cm (herbarium) Leaf: alternate, simple, oblong, notophyll (3-6.5 x 6-14 cm), entire, coriaceous Inflorescence: axillary, flowers in a closed receptacle Flower: very small Fruit: fleshy (fig), subglobose (2.5 cm in diameter), green; many seeds Seed: small (0.2 x 0.2 cm) Other: the slash exudes a white latex.

Dispersal: probably by animals (monkeys, birds, bats)

Distribution Data sources

Continent: Upper Guinea endemic Upper Guinea: Senegal, Guinea-Bissau, Guinea, Sierra Leone, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 12 30’ cells, distribution range is 1670 km Forest type: moist forest, riverine forest

Hawthorne & Jongkind (2004)

Habitat Usually, found along rivers in moist forests (herbarium).

spp

228

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

F.s.

14

25 (4)

0

50

21

72

14

7

7

0

43

57

36

29

14

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 229

Garcinia elliotii

Engl.

Guttiferae Description

Phenology

Guild: u Life form: shrub Max. height: 2 m (Hawthorne & Jongkind 2004) Max. diameter: data unavailable Leaf: opposite, simple, oblong to oblong-elliptic, notophyll (3-4 x 10-12 cm), entire, coriaceous, glabrous; distinctly petiolate Inflorescence: axillary, fascicle Flower: data unavailable Fruit: fleshy, ellipsoid (1.5 x 2.5 cm), densely verrucose; 2 seeds Seed: large (0.8 x 1.5 cm) Other: the slash exudes a yellow latex.

Dispersal: by animals (elephants)

Data sources FWTA, Engler (1908), Kasparek (2000), Hawthorne & Jongkind (2004)

Distribution

G

Continent: Upper Guinea endemic Upper Guinea: Guinea Bissau, Sierra Leone Distribution type: continuous, widespread, present in 4 30’ cells, distribution range is 617 km Forest type: data unavailable

Habitat It is sometimes found by streams (herbarium).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

G.e.

6

- (0)

0

100

0

50

0

0

51

0

100

0

50

50

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

229

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Gardenia nitida

Page 230

Hook.

Rubiaceae Distribution

Uses

Continent: Nigeria, Cameroon (herbarium) Upper Guinea: Guinea-Bissau, Guinea, Sierra Leone, Côte d’Ivoire, Ghana (herbarium) Distribution type: continuous, widespread, present in 20 30’ cells, distribution range is 2977 km Forest type: moist evergreen forest, dry forest, savanna-woodland, gallery forest, secondary forest

The fruits are edible (herbarium). In Ghana, the roots are used to increase the fertility of women.

Data sources FWTA, Hawthorne & Jongkind (2004)

Habitat Species occurrence increases in places where rainfall is less than 1500 mm/yr (Chi2 test). Often near rivers, especially in drier areas (herbarium, Hawthorne & Jongkind 2004). Usually in dry forests but also common in villages and botanical nurseries due to its medicinal and practical uses. It can also be found in the rainforest, both in clearings or deep shade.

G

Phenology Description

Dispersal: probably by animals

Guild: sb Life form: shrub or small tree Max. height: 6 m (herbarium) Max. diameter: 4 cm (herbarium) Leaf: opposite, simple, elliptic, mesophyll (3-8.5 x 7-25 cm), entire, becoming dentate towards the apex, glabrous; interpetiolar stipules Inflorescence: axillary, solitary Flower: large (corolla up to 11 cm long, and lobes up to 4.5 cm long); corolla creamy, long tubular; very fragrant Fruit: woody, long and narrow (2.5 x 8 cm), brownish; many seeds Seed: small (0.2 x 0.2 cm) Other: it has pale corky twigs, usually forking into trees.

spp

230

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

G.n.

37

20 (15)

3

30

19

65

17

0

19

3

62

35

16

35

41

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 229-238

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Page 231

Gilbertiodendron bilineatum

(Hutch. & Dalziel) J.Léonard

Leguminosae-Caes. Description

Phenology

Guild: u Life form: medium-sized to large tree Max. height: 33 m (herbarium) Max. diameter: 95 cm (herbarium) Leaf: opposite, paripinnately compound, 8-14 leaflets, elliptic, notophyll (2.5-5 x 6-15 cm), entire with glandular notches, short hairs on midrib and twigs, sometimes with remarkable leafy stipules Inflorescence: axillary, branched Flower: medium-sized; bracts pink; petals white to yellow Fruit: dry dehiscent, in pairs, flat with 2-3 ridges (6.5 x 28 cm), velvety, 3-4 seeds Seed: very large Other: the bark is flaky, the slash hard and pink.

Deciduousness: deciduous

G

Data sources

Distribution

FWTA, Savill & Fox (1967), Lock (1989), Hawthorne (1995a), IUCN Red List (2000), Hawthorne & Jongkind (2004)

Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 14 30’ cells, distribution range is 1323 km, Red List species (Vulnerable) Forest type: wet evergreen forest. It is gregarious in some parts of Sierra Leone (herbarium), but very rare in Ghana. There, it is occasionally found along rivers in the evergreen forest zone (Hawthorne 1995a).

Habitat It is mostly found where rainfall is higher than 2500 mm/yr (Chi2 test). Generally, along rivers and in moist sites of wet evergreen forests (Savill & Fox 1967). In Ghana, usually along rivers and swamps (herbarium, Hawthorne & Jongkind 2004).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

G.b.

17

0 (4)

0

29

6

12

30

30

30

0

65

35

41

0

H 53

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

231

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Page 232

Gilbertiodendron limba

(Scott-Elliot) J.Léonard

Leguminosae-Caes. Phenology

Seed: flat, very large (3.5 x 4.5 cm), bright brown Other: it is unbuttressed, branching low and the crown is deep and close to the stem. Probably ectomycorrhizal, like others in the genus.

Deciduousness: evergreen Dispersal: animals (e.g. monkeys) might eat the seeds (De Koning 1983, Hawthorne 1995a), otherwise short-distance explosive dehiscence (Aubréville 1959) Timing: flowering period from February to June (De Koning 1983); fruiting period from July to September (Ghana, Taylor 1960)

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leona, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 22 30’ cells, distribution range is 1412 km Forest type: evergreen forest, swamp forest, riverine forest, secondary forest

Habitat

G

Mostly found where rainfall is between 1500-2500 mm/yr (Chi2 test). Usually close to rivers and swamps (herbarium, Hawthorne 1995a). Not uncommon in uplands (Chi2 test) as suggested by Hawthorne (1995a). He indicated that although this species is seen to thrive in swamps, there are many records on slopes and upland sites in evergreen forest, which most likely were the result of misidentification of G. preussii.

Description Guild: sb Life form: medium-sized tree Max height: 25 m (herbarium) Max diameter: 20 cm (herbarium) Leaf: opposite, paripinnately compound, 2-6 leaflets, obovate, mesophyll (4-10 x 10-25 cm), entire, coriaceous, glabrous, yellow-green below Inflorescence: axillary or cauliflorous, branched, hairy Flower: medium-sized; corolla cream coloured, one large petal, white inside and pink outside Fruit: dry dehiscent (pod) (5 x 28 cm), brown, woody, with only 1 ridge (and many fainter diagonal lines); several seeds

spp

232

n

Open (n)

Data sources Aubréville (1959), Taylor (1960), Voorhoeve (1965), De Koning (1983), Alexander (1989), Lock (1989), Hawthorne (1995a)

Regeneration It has a phanerocotylar epigeal reserve seedling type (cf. Voorhoeve 1965) and a fast and high germination rate (Taylor 1960). Individuals of all size-classes are found regenerating in the shade (Hawthorne 1995a).

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

G.l.

36

0 (19)

17

53

3

8

44

39

9

3

47

50

22

0

56

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 229-238

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Page 233

Gilbertiodendron obliquum

(Stapf ) J.Léonard

Leguminosae-Caes. Description Guild: u Life form: shrub Max. height: data unavailable Max. diameter: data unavailable Leaf: alternate, paripinnately compound, 14 leaflets, mesophyll (up to 4.5 x 15 cm) Inflorescence: branched (panicle) Flower: medium-sized, white Fruit: dry dehiscent (pod) Seed: unknown

Distribution Continent: Upper Guinea endemic Upper Guinea: Liberia Distribution type: continuous, very local, present in 1 30’ cell. An extremely rare species; only one record known from A. Whyte found at Sinoe Basin. Forest type: unknown

G

Data sources FWTA, Lock (1989)

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

G.o.

1

- (0)

0

100

0

0

0

0

0

0

100

0

100

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

233

22466 layout page 229-238

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Page 234

Gilbertiodendron robynsianum

Aubrév. & Pellegr.

Leguminosae-Caes. Description

Data sources

Guild: u Life form: small tree Max. height: 5 m (herbarium) Max diameter: 10 cm (herbarium) Leaf: paripinnately compound, 10-12 leaflets, elliptic, mesophyll (4-7 x 10-25 cm), herbaceous to coriaceous; very large stipules (> 10 cm long) Inflorescence: axillary, branched (up to 15 cm long, with large inflorescence bracts) Flower: data unavailable Fruit: dry dehiscent (pod), green with brown indumentum Seed: data unavailable

Aubréville (1959), IUCN Red List (2000), Hawthorne & Jongkind (2004)

Distribution

G

Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire Distribution type: continuous, very local, present in 2 30’ cells, distribution range is 68 km, Red List species (Vulnerable). The species has a very restricted distribution in southwest Côte d’Ivoire. Forest type: rainforest

Habitat It has been observed along roadsides. Two out of five records are from trees occurring near water (herbarium).

Phenology

spp

234

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

G.r.

6

0 (5)

0

50

0

0

0

100

0

0

83

17

50

0

33

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 229-238

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Page 235

Globimetula assiana

(Balle) Wiens & Polhill

Loranthaceae Description

Phenology

Guild: pi Life form: hemi-parasitic shrub Max. height: data unavailable Max. diameter: data unavailable Leaf: opposite, simple, lanceolate to ovate, mesophyll (2.5-10 x 5-15 cm), entire, coriaceous, second pair of lateral nerves steeply ascending from near the base to near the apex Inflorescence: axillary or at nodes below, umbels with 1-6 flowers Flower: medium-sized (corolla 2.5-4 cm long); bright pink on lower half, grey-green above Fruit: fleshy (berry), globose (0.5 cm in diameter), red to yellow; 1 seed Seed: brightly coloured, medium-sized Other: it has flattened twigs.

Dispersal: by birds Timing: flowering period from November to December and April to June (Polhill & Wiens 1998)

Distribution

Data sources

Continent: Upper Guinea endemic (Polhill & Wiens 1998) Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire (Polhill & Wiens 1998) Distribution type: continuous, widespread, present in 11 30’ cells, distribution range is 730 km Forest type: montane forest

Polhill & Wiens (1998)

G

Habitat It occurs between 0-1200 m altitude (Polhill & Wiens 1998).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

G.a.

25

- (0)

76

60

4

12

12

68

8

0

24

76

12

0

H 8

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

235

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Page 236

Globimetula cupulata

(DC.) Danser

Loranthaceae Description

Phenology

Guild: pi Life form: hemi-parasitic shrub Max. height: data unavailable Max. diameter: data unavailable Leaf: opposite, simple, ovate to broadly elliptic, mesophyll (3-13 x 6-20 cm), entire, coriaceous, reddish when young, 5-7 pairs of lateral nerves Inflorescence: at older nodes, umbels distinctly papelose Flower: medium-sized (corolla 3-4.5 cm); corolla pinkish red with a green head Fruit: fleshy (berry) Seed: data unavailable Other: the twigs are flattened.

Dispersal: by birds Timing: flowering period from December to May (Polhill & Wiens 1998)

Data sources FWTA, Polhill & Wiens (1998)

G

Distribution Continent: Upper Guinea endemic (Polhill & Wiens 1998) Upper Guinea: Senegal, Guinea Bissau, Guinea, Sierra Leone (Polhill & Wiens 1998) Distribution type: continuous, widespread, present in 12 30’ cells, distribution range is 842 km Forest type: bushes on coastal dunes

Habitat Commonly found on Parinari trees, but also other trees (Polhill & Wiens 1998).

spp

236

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

G.c.

22

100 (3)

14

64

41

50

18

23

9

0

23

77

0

59

H 18

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 229-238

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Page 237

Guaduella macrostachys

(K.Schum.) Pilger

Gramineae Description Guild: sb Life form: perennial herb (dwarf bamboo) Max. height: 1.5 m (herbarium) Max. diameter: 0.9 cm (Botanischer Jahrbucher 1897) Leaf: simple, lanceolate, mesophyll (2-5 x 10-32 cm), entire Inflorescence: terminal, branched (raceme), (5-7 cm long), up to 20 green flowers per spikelet Flower: small; far exerted stamens Fruit: small Seed: data unavailable Other: it is violet at the base.

Distribution

G

Continent: Nigeria, Cameroon, Democratic Republic of Congo (herbarium) Upper Guinea: Ghana (herbarium, Hall & Swaine 1981) Distribution type: continental disjunct, in Upper Guinea present in 3 30’ cells Forest type: wet evergreen forest

Phenology

Habitat

Dispersal: no obvious dispersal mechanism (Hall & Swaine 1981)

It is found under deep forest understorey shade, old gaps, and occasionally along roadsides (herbarium).

Regeneration It regenerates in the shade (Hall & Swaine 1981).

Data sources Hall & Swaine (1981), Van der Zon (1992)

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

G.m

3

0 (3)

0

0

0

0

66

33

0

0

33

67

0

0

100

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

237

22466 layout page 229-238

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Page 238

Guaduella oblonga

Hutch. ex W.D.Clayton

Gramineae Description

Phenology

Guild: sb Life form: perennial rhizomatous herb Max. height: 1.2 m (Van der Zon 1992) Max. diameter: data unavailable Leaf: simple, oblong to elliptic, mesophyll (3-8 x 10-25 cm), entire, herbaceous to coriaceous Inflorescence: terminal, unbranched (spikelet), basal flowers are masculine with esserted stamens, the others hermaphrodite, spikelets pale greenish (4-5 cm long), up to 15 flowers per spikelet. Flower: small Fruit: data unavailable Seed: data unavailable

Data sources FWTA, De Koning (1983), Van der Zon (1992), Poilecot (1995)

Distribution

G

Continent: Cameroon, Equatorial Guinea (herbarium), Gabon (Van der Zon 1992) Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire (herbarium) Distribution type: continental disjunct, in Upper Guinea present in 24 30’ cells Forest type: wet evergreen forest

Habitat Species occurrence increases with rainfall to reach an optimum around 2700 mm/yr. It is found mainly in mature forest, in densely shaded places (herbarium, De Koning 1983, Poilecot 1995). Occasionally also found in logged forest and along roadsides. Incidentally found near rivers or streams (herbarium).

spp

238

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

G.o.

42

31 (13)

10

48

2

0

23

31

45

2

86

12

69

7

H 12

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 239

Guibourtia dinklagei

(Harms) J.Léonard

Leguminosae-Caes. Description Guild: u Life form: small tree Max. height: 5 m (Hawthorne & Jongkind 2004) Leaf: lower leaves on tree simple, ovate, notophyll (4-7 x 7-13 cm), upper leaves bipinnately compound, sickle-shaped, notophyll (3-9 x 11.5 cm), entire, coriaceous with translucent points, glabrous, young leaflets occasionally with a few hairs on nerves below Inflorescence: terminal or axillary, branched (up to 20 cm long) Flower: small; corolla white Fruit: dry indehiscent Seed: data unavailable Other: it has buttresses up to 1.5 m high.

G

Distribution Continent: Upper Guinea endemic Upper Guinea: Liberia Distribution type: continuous, local, present in 4 30’ cells, distribution range is 227 km Forest type: wet evergreen forest, secondary forest

Habitat It is often associated with riverbanks in wet evergreen forests (Hawthorne & Jongkind 2004). On red-yellow clay with gravel of lateritic origin (herbarium).

Phenology Data sources FWTA, Voorhoeve (1965), Lock (1989), Hawthorne & Jongkind (2004)

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

G.d.

4

67 (3)

25

0

0

0

25

0

50

0

100

0

100

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

239

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Page 240

Gymnosiphon longistylus

(Benth.) Hutch.

Burmanniaceae Distribution Continent: Nigeria, Cameroon, Gabon Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana (herbarium) Distribution type: continental disjunct, in Upper Guinea present in 10 30’ cells Forest type: evergreen forest, mangrove

Habitat Found in shady places in mature forest. It occurs at the forest floor, at the base of forest giants or on roots. It occurs often in very moist habitats such as swamps, along creeks, or in mangroves (herbarium).

G

Phenology

spp

240

Description

Data sources

Guild: sb Life form: saprophytic herb Max. height: 0.2 m (herbarium) Leaf: minute and reduced leaf, lacking chlorophyll, completely white, membranous Inflorescence: terminal, branched (lax cyme) Flower: small; calyx white; corolla yellow at heart with white lobes, 3 petals Fruit: subglobose Seed: data unavailable

FWTA

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

G.l.

14

0 (12)

14

43

7

0

28

35

35

0

71

29

43

0

H 50

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 239-248

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Page 241

Gymnostemon zaizou

Aubrév. & Pellegr.

Simaroubaceae Description

Phenology

Guild: sb Life form: medium-sized tree Max. height: 30 m (herbarium) Max. diameter: 100 cm (herbarium) Leaf: alternate, imparipinnately compound; 13-25 subopposite leaflets, oblong, notophyll (2-4.5 x 8-13.5 cm), entire, coriaceous, glabrous Inflorescence: terminal, branched (panicle) Flower: small; yellow Fruit: fleshy (drupe), ovoid (8 x 10 cm); 1 seed Seed: very large Other: the leaves are clustered at the ends of branches. The wood density is 0.55 g/cm3.

Dispersal: by animals Timing: flowering period in October (Kasparek 2000)

Distribution

G

Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire Distribution type: continuous, local, present in 4 30’ cells, distribution range is 201 km, Red List species (Vulnerable). Recorded only in southwest Côte d’Ivoire Forest type: moist evergreen forest

Data sources FWTA, Aubréville (1959), De la Mensbruge (1966), De Rouw (1991), IUCN Red List (2000), Kasparek (2000)

Habitat Data unavailable.

Regeneration It has a cryptocotylar hypogeal reserve seedling type (cf. de la Mensbruge 1966), and regenerates in undisturbed, primary forests (De Rouw 1991).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

G.z.

7

0 (3)

0

43

0

0

100

0

0

0

100

0

100

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

241

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Page 242

Hemandradenia chevalieri

Stapf

Connaraceae

Regeneration It has a phanerocotyal epigeal reserve seedling type (De la Mensbruge 1966). The cotyledons are thick and red. Germination occurs in 2-3 weeks.

Phenology

H

Description

Dispersal: by animals

Guild: u Life form: medium-sized tree Max. height: 15 m (herbarium) Max. diameter: 26 cm (herbarium) Leaf: alternate, simple, oblong to elliptic, notophyll (3-6 x 7-14 cm), entire, herbaceous, shining dark green above, glabrous above, sparsely pubescent beneath; no stipules Inflorescence: terminal, branched (panicle, manyflowered) Flower: small; calyx pale brown hairy outside, glabrous inside; corolla white, hairy Fruit: fleshy indehiscent, obovoid (2.8 x 1.7 cm), yellow-brown to orange; 1 seed Seed: large (2.1 x 1.1 cm), cream-coloured and fleshy on the outside Other: seedlings with glabrous cotyledons turning red-brown upon exposure. The slash is dark reddish.

spp

242

n

Open (n)

Distribution Continent: Gabon Upper Guinea: Côte d’Ivoire Distribution type: continental disjunct, in Upper Guinea present in 3 30’ cells, Red List species (Endangered) Forest type: coastal rainforest

Data sources

Habitat

FWTA, De la Mensbruge (1966), Eimunjeze (1976), Breteler (1989), IUCN Red List (2000), Hawthorne & Jongkind (2004)

It is regularly found in forests near streams, and close to the coast (FWTA).

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

L

M

H

75

8

17

17

67

25

36

13

39

H.c.

12

0 (2)

0

75

8

0

0

100

0

17

All

46333

37

4

39

7

37

29

24

10

6

Soil WHC

69

22466 layout page 239-248

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Page 243

Heritiera utilis

Sprague

Sterculiaceae Description Guild: np Life form: large tree Max. height: 45 m (herbarium) Max. diameter: 300 cm (herbarium) Leaf: alternate, simple, elliptic to ovate, mesophyll (2-10 x 5-25 cm), undulate, herbaceous; in young trees palmately compound (5-7 leaflets) Inflorescence: axillary, branched (panicle, up to 20 cm long) Flower: small; monoecious; bowl-shaped; corolla yellow to white Fruit: dry indehiscent (samara) (1.5 x 2.5 cm), winged, green-brown; 1 seed Seed: very large Other: it has a golden-brownish sheen to the crown due to the scales and stellate hairs on the leaves. Mature trees have characteristic stilt-like buttresses which are especially well-developed in swamp forests. The wood density is 0.66 g/cm3.

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana (herbarium) Distribution type: continuous, widespread, present in 28 30’ cells, distribution range is 1324 km, Red List species (Vulnerable) Forest type: upland evergreen forest, wet evergreen forest, moist evergreen forest, moist semi-deciduous forest, gallery forest, secondary forest. In Ghana, mostly found in wet evergreen forests (Hall & Swaine 1981).

Habitat Species occurrence increases significantly with rainfall to reach an optimum around 2500 mm/yr (logistic regression analysis, Chi2 test). It is usually

found near creeks, riverbanks, and occasionally in swamps (herbarium). It is very drought-sensitive, but shows no strong preference for either base-poor or base-rich soils (Swaine & Veenendaal 1994).

Regeneration It has a phanerocotylar epigeal foliaceous seedling type (De Koning 1983). Germination is normal, or rather rapid. Taylor (1960) records that early growth is very erratic, and that there is no obvious requirement for either shade or sun in nurseries, although growth is rapid under suitable circumstances (2 m after 4 yr in TSS plots), and the tree is best regarded as a ‘light demander’. Savill & Fox (1967) record that seedlings and saplings can tolerate high shade for years and still begin to grow immediately once the canopy is opened. Young trees higher than 1.5 m of this species are less strongly favoured by gaps than many other tree species, but still are disproportionately common there (De Klerk 1991). Growth rings of this species have been analysed in Côte d’Ivoire (Detienne & Mariaux 1975). In Sierra Leone, plantation trees realised an average annual diameter increment of 1.7 cm/yr over the first 15 years and they attained a height of 20 m (Savill & Fox 1967).

Uses

H Phenology Dispersal: by wind Timing: flowering period from October to November; fruiting period from January to March (Voorhoeve 1965). Fruits are produced towards the end of the dry season. Local mast years may occur (Voorhoeve 1965).

It is an excellent timber species (Voorhoeve 1965).

Data sources Taylor (1960), Voorhoeve (1965), Savill & Fox (1967), Britwum (1976), Detienne & Mariaux (1975), Hall & Swaine (1981), De Koning (1983), De Klerk (1991), Swaine & Veenedaal (1994), Hawthorne (1995a), Poorter et al. (1996), IUCN Red List (2000)

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

H

H.u.

62

11 (38)

3

44

6

0

50

24

26

3

74

23

45

2

48

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

243

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Hibiscus whytei

Page 244

Stapf

Malvaceae Description

Data sources

Guild: u Life form: tall herb Max. height: data unavailable Max. diameter: data unavailable Leaf: alternate, simple, ovate to triangular (3-9 cm long), irregularly dentate Inflorescence: solitary Flower: large; corolla yellow with purple centre Fruit: data unavailable Seed: data unavailable

FWTA

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea (FWTA), Liberia (herbarium), Côte d’Ivoire (FWTA) Distribution type: continuous, very local, present in 1 30’ cell Forest type: data unavailable

H

Habitat Data unavailable.

Phenology

spp

244

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

H.w.

1

- (0)

0

100

0

0

0

0

100

0

100

0

100

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 239-248

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Page 245

Hippocratea vignei

Hoyle

Celastraceae Description

Phenology

Guild: np Life form: large irregularly winding woody climber Max. height: 30 m long (herbarium) Max. diameter: 10 cm (herbarium) Leaf: opposite, simple, oblong, mesophyll (8 x 12 cm), dentate, coriaceous Inflorescence: branched Flower: small; white Fruit: dry dehiscent, with 3 flat carpels, 6 cm long, woody, green; up to 24 seeds Seed: very large (4-5 cm long), 1-winged Other: the wood is pale yellow with yellowish to pinkish sapwood. The stem and main branches are quadrangular to strongly fluted. Resinous threads appear when the leaves are torn apart.

Dispersal: by wind

Data sources

Distribution

Hallé (1962), Hawthorne (1995a), Hawthorne & Jongkind (2004)

Continent: Nigeria Upper Guinea: Côte d’Ivoire, Ghana Distribution type: continuous, widespread, in Upper Guinea present in 9 30’ cells, distribution range is 1384 km Forest type: rainforest, semi-deciduous forest, savanna, secondary vegetation

H

Habitat It is found along roads and clearings, but also in mature forest (herbarium).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

Soil CMK

Soil WHC

VW

L

M

H

L

M

H

H.v.

11

22 (9)

9

27

0

27

73

0

0

0

55

45

45

9

45

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

245

22466 layout page 239-248

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12:01 PM

Hugonia rufipilis

Page 246

A.Chev. ex Hutch. & Dalziel

Linaceae

H

Description

Phenology

Guild: np Life form: large woody liana, climbing with hooks Max. height: 50 m long (herbarium) Max. diameter: 3 cm (herbarium) Leaf: alternate, simple, elliptic, mesophyll (5-10 x 11-32 cm), serrate, herbaceous, pubescent; nerves pale brown and hairy Inflorescence: terminal or axillary, branched (cyme) (up to 25 cm long) Flower: medium-sized; calyx brownish; corolla yellow Fruit: fleshy, globose (1.3 x 1.3 cm), yellowish; 1 seed Seed: stony Other: it starts as a scandent shrub. The tendrils and young twigs are brown and hairy.

Timing: flowering period from August to September; fruiting period from December to January (De Koning 1983)

Distribution

Data sources

Continent: Senegal to the Democratic Republic of Congo (De Koning 1983), but according to Hall & Swaine (1981) it is found from Sierra Leone to Ghana and the Democratic Republic of Congo. The species is also reported from Congo (Brazzaville), but this determination is uncertain. Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana (herbarium) Distribution type: continuous, widespread, present in 8 30’ cells, distribution range is 1168 km Forest type: rainforest, secondary forest

FWTA, Badré (1971), Hall & Swaine (1981), De Koning (1983)

Habitat It is often found near rivers.

spp

246

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

H.r.

11

29 (7)

0

55

18

0

9

55

36

0

100

0

36

0

64

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 239-248

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Page 247

Hunteria ghanensis

J.B.Hall & Leeuwenb.

Apocynaceae Description Guild: sb Life form: small to medium-sized tree Max. height: 12 m (Omino 1996) Max. diameter: 25 cm (Omino 1996) Leaf: opposite, simple, narrowly elliptic, microphyll (0.8-3.5 x 3.5-14 cm), entire to slightly crenate, fairly coriaceous Inflorescence: terminal, almost compound umbellike, many flowers (30-150) Flower: medium-sized (2-4 x 2.5-4 cm); corolla 0.60.9 cm long, pale yellow; tube with 5 lobes Fruit: fleshy (1.5 x 1 x 1 cm), smooth, yellow to orange; 1-2 seeds Seed: medium-sized (0.9 x 0.6 x 0.4 cm) Other: the bark is pale grey, shallowly fissured, but dark brown on section. It exudes a scanty white latex.

Distribution

H

Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 6 30’ cells, distribution range is 677 km, Red List species (Endangered); rare in the world (Hall & Leeuwenberg 1979) Forest type: dry forest, secondary forest. In Ghana, only in extremely dry forests (Hall & Swaine 1981, Hawthorne 1995a, 1996).

Regeneration It regenerates in shade.

Uses

Phenology

The timber is tough and fine-grained (Hall & Swaine 1981).

Timing: not clearly seasonal (Hall & Swaine 1981), flowering and fruiting periods between December and July in both countries (Omino 1996)

Habitat

Data sources

It is found in dry and open places, on rocks, and inselbergs. Generally found in dry forests where rainfall is between 100-1200 mm/yr and the altitude is between 50-100 m (Omino 1996).

spp

n

Open (n)

Hall & Leeuwenberg (1979), Hall & Swaine (1981), Hawthorne (1995a, 1996), Omino (1996), IUCN Red List (2000)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H.g.

10

20 (5)

0

60

20

60

30

10

0

0

80

20

10

0

H 80

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

247

22466 layout page 239-248

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Page 248

Hutchinsonia barbata

Robyns

Rubiaceae

H

spp

248

Description

Distribution

Guild: np Life form: straggling shrub Max. height: 3 m (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, ovate to lanceolate, microphyll (1.5-3 x 4-8 cm), entire, herbaceous, hairy; almost sessile; interpetiolar stipules Inflorescence: axillary, solitary Flower: medium-sized (tube approx. 2.5 cm long); corolla tube yellow, lobes orange and reflexed, shorter than tube Fruit: fleshy, globose (0.5 cm in diameter), whitish, 2-lobed; 1-2 seeds Seed: medium-sized (0.4 x 0.4 cm) Other: occasionally with spines on the lower parts of the stems. The twigs are covered with dense curly hairs.

Continent: Upper Guinea endemic Upper Guinea: Guinea (Hawthorne & Jongkind 2004), Sierra Leone, Liberia, Côte d’Ivoire (herbarium) Distribution type: continuous, widespread, present in 20 30’ cells, distribution range is 842 km Forest type: montane forest, moist evergreen forest, coastal forest, swamp forest, secondary forest

n

Open (n)

Phenology

Habitat Data sources

Species occurrence increases significantly at places where rainfall is higher than 1500 mm/yr (Chi2 test). It can be found in the understorey but also along forest edges, logging roads, and secondary forests. It is frequently found along rivers, flooded grounds, and riverbanks (herbarium). On clayish, sandy and lateritic soils.

FWTA, Hawthorne & Jongkind (2004)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H.b.

29

43 (21)

3

34

17

0

41

24

34

0

86

14

66

0

H 34

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 249-258

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Page 249

Hutchinsonia glabrescens

Robyns

Rubiaceae Description Guild: sb Life form: straggling shrub Max. height: 2 m (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, ovate-oblong to oblongelliptic, microphyll (1.3-2 cm x 3.5-5.5 cm), entire, glabrous, midrib hairy; interpetiolar stipules Inflorescence: axillary or terminal, solitary Flower: medium-sized; corolla yellow to orange Fruit: fleshy; 1-2 seeds Seed: data unavailable Other: it has spines.

Distribution Continent: Upper Guinea endemic Upper Guinea: Liberia Distribution type: present in 3 30’ cells Forest type: wet evergreen forest

H

Habitat Data unavailable.

Phenology

Data sources FWTA

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

H.g.

3

0 (1)

33

33

33

0

33

33

33

0

100

0

67

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

249

22466 layout page 249-258

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Page 250

Hymenocoleus axillaris

Robbr.

Rubiaceae Description

Phenology

Guild: sb Life form: shrub Max. height: 0.6 m (Robbrecht 1977) Max. diameter: data unavailable Leaf: opposite, simple, obovate, mesophyll (4.5-6 x 13-17 cm), entire, herbaceous; petiole and nerves pubescent; interpetiolar stipules Inflorescence: axillary, 2-7 pairs of glomerous inflorescences each with approx. 20 flowers Flower: small (corolla tube 0.4 cm); calyx pale green; corolla white Fruit: fleshy (berry), subglobose (1 cm in diameter), orange; 2 seeds Seed: hemispherical, medium-sized (0.3 x 0.3 x 0.2 cm)

Data sources Robbrecht (1977)

Distribution

H

Continent: Cameroon, Gabon, Congo (Brazzaville) (herbarium, Robbrecht 1977) Upper Guinea: Liberia (herbarium, Robbrecht 1977) Distribution type: continental disjunct with only a satellite population in Upper Guinea, in Upper Guinea present in 1 30’ cell Forest type: wet evergreen forest, secondary forest

Habitat It occurs in the shaded understorey (Robbrecht 1977).

spp

250

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

H.a.

1

- (0)

0

0

0

0

0

0

100

0

100

0

100

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 249-258

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Page 251

Hymenostegia gracilipes

Hutch. & Dalziel

Leguminosae-Caes. Description

Phenology

Guild: sb Life form: medium-sized tree Max. height: 16 m (herbarium) Max. diameter: data unavailable Leaf: alternate, paripinnately compound, 6-8 leaflets, oblong to triangular, microphyll (2-3.5 x 4-8 cm), entire, dull green below, rachis deeply grooved, and young twigs sometimes grooved as well; leaflets occasionally emarginate, rachis often with a small pair of tiny, waxy, gland-like structures attached to rachis at base of middle pair(s) of leaflets, leaflets sometimes gland-dotted at base of lamina as well Inflorescence: terminal, unbranched (raceme) Flower: medium-sized; corolla white Fruit: dry dehiscent (pod), flat Seed: unknown Other: a tree with a spreading crown of drooping branches.

Dispersal: probably explosive Timing: flowering period: one record in August (herbarium)

Data sources FWTA, Hall & Swaine (1981), Lock (1989), Hawthorne (1995a), IUCN Red List (2000), Kasparek (2000), Hawthorne & Jongkind (2004)

H

Distribution Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire (one doubtful record), Ghana Distribution type: continuous, local, present in 5 30’ cells, distribution range is 257 km, Red List species (Endangered). In Ghana, it is an uncommon species (Hawthorne 1995a). Forest type: wet evergreen forest, riverine forest

Habitat It is mostly seen close to rivers in evergreen forest (Hawthorne & Jongkind 2004). On sandy soils (herbarium).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

H.g.

6

0 (3)

0

83

0

0

34

67

0

33

33

33

0

17

67

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

251

22466 layout page 249-258

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Page 252

Illigera vespertilio

(Benth.) Baker f.

Hernandiaceae Description

Phenology

Guild: np Life form: small winding woody climber Max. height: 8 m long (herbarium) Max. diameter: data unavailable Leaf: alternate, compound, with 3 leaflets in West Africa and 5 leaflets in Gabon, obovate, notophyll (2-7 x 3.5-9 cm), entire, coriaceous, glabrous Inflorescence: axillary, branched Flower: small; corolla pale pink Fruit: dry indehiscent (3 cm long), 2 or 3 opposite wings Seed: data unavailable Other: it climbs with a winding petiole.

Dispersal: probably by wind

Distribution

Data sources

Continent: Cameroon, Gabon, Democratic Republic of Congo Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continental disjunct, widespread, present in 5 30’ cells Forest type: rainforest, gallery forest, secondary forest

I

Kubitzki (1969), De Koning (1983), Hawthorne & Jongkind (2004)

Habitat Data unavailable.

spp

I.v. All

252

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

11

29 (7)

0

18

0

9

72

18

0

18

27

55

18

18

64

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 249-258

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12:05 PM

Page 253

Impatiens nzoana

A.Chev. ssp.

bennae

Jacq.-Fél.

Balsaminaceae Description

Phenology

Guild: sb Life form: perennial herb Max. height: 0.6 m Max. diameter: data unavailable Leaf: spirally arranged, simple, narrowly elliptic to oblong, notophyll (1.8-4 x 6-17 cm), finely dentate, glabrous Inflorescence: axillary, solitary Flower: lilac, mauve or pink with a whitish spur which is 5.5-7 cm long in subsp. bennae, and 3.54.7 cm long in subsp. nzoana Fruit: dry dehiscent (0.4 x 1.5 cm), glabrous Seed: small (0.08-0.11 cm), ellipsoid, covered in long hairs Other: the stem is relatively erect, glabrous, usually unbranched. It is rooting at the lower nodes.

Dispersal: probably by wind

Data sources Grey-Wilson (1980)

Distribution I. nzoana bennae is an Upper Guinea endemic that occurs in Guinea. It has a very local distribution and is present in only 1 30’ cell. I. nzoana nzoana is an Upper Guinea endemic that occurs in Liberia and Côte d’Ivoire. It has a very local distribution and is present in 2 30’ cells. It has a distribution range of only 7 km and is found in gallery forest.

I

Habitat I. nzoana grows in moist shaded places of riverine forests and along watercourses.

Regeneration Spreading by thin underground rhizomes.

spp

I.n. All

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

6

0 (2)

50

50

0

0

0

100

0

0

83

17

66

0

17

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

253

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Isolona cooperi

Page 254

Hutch. & Dalziel ex Coop. & Record

Annonaceae Distribution

Data sources

Continent: Upper Guinea endemic Upper Guinea: Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 16 30’ cells, distribution range is 1058 km Forest type: evergreen forest, semi-deciduous forest, swamp forest, secondary forest (herbarium). In Ghana, mostly in wet and moist evergreen forests, but it is also present in moist semideciduous forests (Hall & Swaine 1981).

Hall & Swaine (1981), Hawthorne (1995a), Hawthorne & Jongkind (2004)

Habitat It is found most often where rainfall is between 1500-2000 mm/yr (Chi2 test). Frequently found along river borders, and occasionally in swampy places. Often under shade, but also under broken canopy in disturbed habitats. On sandy or clayish soils (herbarium).

Phenology

I

Deciduousness: evergreen Dispersal: probably by animals Timing: not clearly seasonal (Hall & Swaine 1981)

Description Guild: sb Life form: shrub or small tree Max. height: 9 m (herbarium) Max. diameter: data unavailable Leaf: alternate, simple, elliptic, mesophyll (6-12 x 14-28 cm), entire, coriaceous, very shining and dark-green above Inflorescence: axillary, solitary Flower: medium-sized (approx. 2 cm); green to yellowish, bowl-shaped Fruit: fleshy, oblong (9 x 2.5 cm), yellow to orange with white spots; many seeds Seed: medium-sized (1 x 1 cm) Other: it is a very aromatic tree.

spp

I.c. All

254

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

39

38 (29)

3

31

13

3

79

16

3

5

41

54

23

3

72

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 249-258

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Page 255

Isonema smeathmannii

Roem. & Schult.

Apocynaceae Description

Habitat

Guild: pi Life form: large winding woody climber Max. height: 8 m high, 30 m long (De Koning 1983, herbarium) Max. diameter: 2 cm or more (Van der Ploeg 1983) Leaf: opposite, simple, oblong to obovate, notophyll (2.3-5 x 4-12 cm), entire, coriaceous, pubescent on both sides, especially on the main veins; petiole short and pubescent Inflorescence: terminal (occasionally axillary), branched (6-28 cm long) Flower: medium-sized (corolla 1.2-1.8 cm, tube 0.7-1.1 cm); corolla 5-merous, pubescent; lobes inside red-brown to red-pink and with four yellow longitudinal stripes, outside mainly at the base yellowish and pubescent; reddish pedicels Fruit: dry dehiscent, double horn-shaped (0.7 x 16 cm), pubescent, brown; many seeds Seed: very large (0.8-2.2 cm long), plumed, hairs simple Other: it occurs in the form of a shrub when young. It has a smooth dark red-brown bark on older stems. The twigs are dull green-brown, with coarse rusty hairs. The bark has white lenticels and the slash exudes a scarce white latex.

Species occurrence increases significantly with rainfall higher than 2000 mm/yr to reach an optimum around 3000 mm/yr (logistic regression analysis). It is more often found in coastal ranges and close to rivers (Chi2 test). Frequently climbing in open (e.g. along roadsides and fringing forest) and moist to swampy places (e.g. edge of mangrove swamps, creeks, rivers, and lagoons). Often found on sandy soils, but also on clay and lateritic soils (herbarium).

Phenology Dispersal: probably by wind Timing: flowering during the whole year (Van der Ploeg 1983)

I

Distribution Continent: Upper Guinea endemic Upper Guinea: Senegal, Guinea Bissau, Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 39 30’ cells, distribution range is 1764 km Forest type: coastal forest, gallery forest, swamp forest, savanna-woodland, secondary forest

Uses The young leaves are used as a vegetable in Sierra Leone (Van der Ploeg 1983).

Data sources FWTA, De Koning (1983), Van der Ploeg (1983)

spp

I.s. All

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

145

81 (52)

1

49

35

3

25

28

39

33

46

21

14

27

H 21

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

255

22466 layout page 249-258

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12:05 PM

Ixora hiernii

Page 256

Scott-Elliot

Rubiaceae

I

Description

Phenology

Guild: sb Life form: shrub or small tree Max. height: 7 m (De Block 1998) Max. diameter: data unavailable Leaf: opposite, simple, elliptic to obovate, mesophyll (3-8 x 8-21.5 cm), fairly coriaceous, glabrous; petiole 0.3-1.5 cm long, glabrous; with interpetiolar stipules Inflorescence: terminal, branched (corymbose), inflorescence axes reddish and densely hairy, sessile or shortly pedunculate Flower: medium-sized; calyx glabrous; corolla white to pink, a long corolla tube (2-3 cm long) with short lobes; very fragrant Fruit: fleshy (drupe), bilobed, subglobose, somewhat transversally flattened (0.8 x 1 cm); reddish when ripe; persistent calyx; 1-2 seeds Seed: medium-sized (0.7 x 0.8 cm), reddish brown

Dispersal: probably moth-pollinated. Dispersed by birds and small mammals (De Block 1998) Timing: flowering period from November to April; fruiting period from November to February, April, August (De Block 1998)

Distribution

Data sources

Continent: Upper Guinea endemic (De Block 1998) Upper Guinea: Guinea, Sierra Leone, Côte d’Ivoire, Ghana, Togo (herbarium, De Block 1998) Distribution type: continuous, widespread, present in 18 30’ cells, distribution range is 1523 km Forest type: rainforest, gallery forest, savanna

FWTA, De Block (1998), Hawthorne & Jongkind (2004)

Habitat It occurs up to 900 m altitude, in forest understorey, often near streams (herbarium, De Block 1998).

Regeneration Germination and seedling establishment unknown (De Block 1998).

spp

I.h. All

256

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

28

0 (1)

54

43

4

11

25

50

15

0

32

68

21

7

H 7

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 249-258

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Page 257

Ixora laxiflora

Sm.

Rubiaceae Description

Habitat

Guild: pi Life form: shrub or small tree Max. height: 15 m Max. diameter: data unavailable Leaf: opposite, simple, elliptic to obovate, mesophyll (3-8 x 8-21.5 cm), entire, fairly coriaceous, glabrous; petiole 0.3-1.5 cm long, glabrous; with interpetiolar stipules Inflorescence: terminal, branched (corymbose), inflorescence axes reddish and densely hairy, sessile or shortly pedunculate Flower: medium-sized (corolla tube 2-3 cm long); calyx glabrous, green; corolla white to pink, a long corolla tube with short lobes; very fragrant Fruit: fleshy (drupe), bilobed, subglobose, somewhat transversally flattened (0.6 x 1 cm); reddish when ripe; persistent calyx; 1-2 seeds Seed: medium-sized (0.7 x 0.8 cm), reddish brown Other: I. laxiflora var. laxiflora differs from I. laxiflora var. linderi in its long pedicels.

Species occurrence increases with rainfall to reach a wide optimum between 2200-3500 mm/yr (logistic regression analysis, Chi2 test). It is mostly found in the lowlands (logistic regression analysis) and occurs frequently near the coast (Chi2 test). It occurs in relatively open places of the forest (e.g. forest roads or edges) or in relatively open plant communities such as coastal savannas or near the beach. Often also close to rivers and swamp edges. Usually on sandy soils, but sometimes also on loamy and lateritic soils (herbarium).

Phenology Dispersal: by birds and small mammals (De Block 1998) Timing: flowering period from July to October; fruiting period from November (De Koning 1983)

Distribution I. laxiflora laxiflora is an Upper Guinea endemic that occurs in Senegal, Guinea Bissau, Guinea, Sierra Leone, Liberia, Côte d’Ivoire, and Ghana. It has a continuous distribution, is widespread and is present in 41 30’ cells. Its distribution range is 1809 km. It can be found in wet evergreen forest, moist evergreen forest, savanna-woodland, gallery forest, swamp forest, beach forest, coastal savanna, and secondary forest. I. laxiflora linderi is an Upper Guinea endemic that occurs in Liberia and Côte d’Ivoire. It has a continuous distribution, is widespread and is present in 2 30’ cells. Its distribution range is 772 km. The last record of this species was collected in 1962.

I

Data sources FWTA, De Koning (1983), Hawthorne (1995a), De Block (1998)

spp

I.l. All

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

112

39 (23)

5

40

46

5

29

24

36

17

51

33

24

14

H 22

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

257

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Ixora liberiensis

Page 258

De Block

Rubiaceae Distribution Continent: Upper Guinea endemic (De Block 1998) Upper Guinea: Liberia (herbarium, De Block 1998) Distribution type: continuous, very local, present in 1 30’ cell, distribution range is 47 km. Known from only two localities around Mt Nimba. Forest type: probably montane or submontane forest

Habitat Found in the forest understorey (De Block 1998).

Phenology Dispersal: probably by birds and small mammals (De Block 1998) Timing: flowering period in December (De Block 1998)

I Description Guild: u Life form: shrub Max. height: data unavailable Max. diameter: data unavailable Leaf: opposite, simple, elliptic, microphyll (1.5-4 x 5.5-10 cm), glabrous, 10-15 pairs of lateral veins; petiole 0.2-0.5 cm long, glabrous, with interpetiolar stipules Inflorescence: terminal, branched (compact triads, up to 3 cm long, sessile) Flower: medium-sized (corolla tube 1.5 cm long); 4merous; corolla white to creamy, glabrous outside, hairy inside Fruit: unknown Seed: unknown

spp

258

n

Open (n)

Data sources De Block (1998)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

Soil WHC H

L

M

H

I.l.

2

- (0)

50

50

0

0

0

100

0

0

100

0

100

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 259

Ixora tenuis

De Block

Rubiaceae Description Guild: sb Life form: shrub Max. height: 2 m (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, narrowly elliptic, notophyll (1.8-5 x 9-14 cm), glabrous, 10-16 pairs of lateral veins; petiole 0.2-0.6 cm long, glabrous, with interpetiolar stipules Inflorescence: terminal, branched (triads, up to 5 cm long, sessile or shortly pedunculate) Flower: medium-sized (1.1-1.6 cm long), tubeshaped, glabrous outside and inside, white; no fragrance Fruit: fleshy, bilobed (0.8 x 0.9 cm), red when ripe, calyx persistent; 1-2 seeds Seed: medium-sized (0.6 x 0.6 cm), reddish brown

Distribution Continent: Upper Guinea endemic (De Block 1998) Upper Guinea: Ghana (herbarium, De Block 1998) Distribution type: continuous, local, present in 4 30’ cells, distribution range is 254 km. Known only from Ghana, although Hawthorne & Jongkind (2004) expect it to occur in Côte d’Ivoire as well. Forest type: upland evergreen forest, wet evergreen forest

Phenology

I

Dispersal: probably moth pollinated. Dispersed by birds and small mammals (De Block 1998) Timing: flowering period in February, and from September to October, December; fruiting period in September (De Block 1998)

Habitat It is found in the forest understorey (herbarium).

Data sources De Block (1998), Hawthorne & Jongkind (2004)

spp

I.t All

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

10

0 (2)

10

30

0

0

90

10

0

0

10

90

0

0

30

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

259

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Keetia bridsoniae

Page 260

Jongkind

Rubiaceae Habitat Sometimes found along roads or near riverbanks.

Phenology

Description Guild: u Life form: large woody climber Max. height: 30 m long (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, elliptic, microphyll (1-2 x 2-5.5 cm), entire, herbaceous, very discolorous, midrib red-brown, hairy beneath on the bigger nerves and in the axils of the laterals; interpetiolar stipules Inflorescence: axillary; flowers in heads Flower: calyx light green; corolla tube pale green, lobes white; fragrant Fruit: fleshy, subglobose (0.6 cm in diameter), green; infructescence on stalks; 1-2 seeds Seed: medium-sized (0.3 x 0.3 x 0.2 cm) Other: the main stem is quadrangular.

Data sources Hawthorne & Jongkind (2004)

K Distribution Continent: Upper Guinea endemic Upper Guinea: Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 10 30’ cells, distribution range is 1132 km Forest type: wet evergreen forest, moist evergreen forest, secondary forest

spp

260

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

K.b

13

50 (8)

8

46

0

0

61

15

23

0

54

46

54

0

23

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 261

Keetia obovata

Jongkind

Rubiaceae Description Guild: u Life form: woody climber Max. height: data unavailable Max. diameter: data unavailable Leaf: opposite, simple, obovate, very pale beneath with brownish triangular domatia, with approx. 5 pairs of laterals; interpetiolar stipules with a long and narrow appendage Inflorescence: axillary, many flowers in a head Flower: data unavailable Fruit: green; 1-2 seeds Seed: data unavailable

Distribution Continent: Upper Guinea endemic Upper Guinea: Liberia, Côte d’Ivoire Distribution type: present in 2 30’ cells Forest type: wet evergreen forest

Habitat Data unavailable.

Phenology

K

Data sources Hawthorne & Jongkind (2004)

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

K.o.

2

50(2)

0

50

0

0

50

50

0

0

100

0

50

0

50

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

261

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Page 262

Landolphia micrantha

(A.Chev.) Pichon

Apocynaceae Habitat It is most often found where rainfall is between 1500-2000 mm/yr (Chi2 test). Although in Ghana the species is mainly found in upland evergreen forests (Hall & Swaine 1981), we actually found a negative correlation with increasing altitude (logistic regression analysis). It is often found in gaps, cleared forests, or along forest edges. Occasionally found near rivers (herbarium).

Regeneration It regenerates in shade (Hall & Swaine 1981).

Phenology Dispersal: by animals (monkeys)

Description Guild: np Life form: large winding woody climber Max. height: 30 m long (herbarium) Max. diameter: 2 cm (Hawthorne & Jongkind 2004) Leaf: opposite, simple, elliptic to oblong, notophyll (2-6 x 5-17 cm), entire, herbaceous to coriaceous Inflorescence: axillary and terminal Flower: small (approx. 0.3 cm long); dish-shaped with tube; corolla creamy-orange in bud, creamy at anthesis; fragrant Fruit: fleshy, oblong to cylindrical (up to 3 x 10 cm), yellow to orange and red with dark spots; many seeds Seed: large Other: the slash exudes a sticky white latex.

L

spp

262

n

Open (n)

Distribution Data sources

Continent: Upper Guinea endemic Upper Guinea: Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 24 30’ cells, distribution range is 1134 km Forest type: rainforest, secondary forest, thicket. In Ghana, only found in evergreen forest, especially upland forests but also present in moist and wet evergreen forests (Hall & Swaine 1981).

FWTA, Hall & Swaine (1981), Persoon et al. (1992), Hawthorne & Jongkind (2004)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

L.m.

71

29 (42)

6

51

1

0

51

35

10

4

73

23

24

3

62

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 263

Landolphia togolana

(Hallier f.) Pichon

Apocynaceae Description

Habitat

Guild: np Life form: small winding woody climber Max. height: data unavailable Max. diameter: 15 cm (Persoon et al. 1992) Leaf: opposite, simple, ovate or elliptic, occasionally obovate, notophyll (1.9-7 x 3.5-13 cm), entire, coriaceous, glabrous on both sides Inflorescence: axillary (or occasionally terminal), branched, 1-12-flowered Flower: small (0.25-1.2 x 0.3-1.1 cm); whiteyellowish; dish with tube; fragrant; glabrous or occasionally with some hairs inside Fruit: fleshy, indehiscent, globose, greenish (4 x 5 cm), pulp fleshy and juicy, yellow, orange, or brown, with brown lenticellate spots; 3-40 seeds Seed: large (1.2 x 0.8 x 0.5 cm) Other: it starts as a shrub. The slash exudes a white latex. It grows with curled tendrils.

Species occurrence is highest where rainfall is around 1000 mm/yr and decreases with increasing rainfall (logistic regression analysis, Chi2 test). It is more often found in coastal ranges (Chi2 test) and occurrence tends to decrease with increasing altitude (logistic regression analysis). It is often found in secondary vegetation (e.g. forest edges and roadsides thickets) and moist places (e.g. riverbanks, streams, lagoons) (herbarium). Usually found on sandy soil (herbarium).

Regeneration It regenerates in shade (Hall & Swaine 1981).

Phenology

Distribution

Dispersal: by animals (monkeys) Timing: flowering period mainly at the beginning of the rainy season; mature fruits can be found throughout the year (Persoon et al. 1992)

Continent: Guinea wide: Benin, Nigeria (herbarium, Persoon et al. 1992) Upper Guinea: Côte d’Ivoire, Ghana, Togo Distribution type: continuous, widespread, present in 18 30’ cells, distribution range is 1163 km Forest type: dry semi-deciduous forest, secondary forest, riverine forest, gallery forest in savanna, coastal forest (herbarium, Persoon et al. 1992). In Ghana, it is most abundant in South Marginal forests and dry semi-deciduous forests (Hall & Swaine 1981).

L

Uses The rubber is said to be of good quality in Benin (Persoon et al. 1992).

Data sources FWTA, Hall & Swaine (1981), Persoon et al. (1992)

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

L.t.

29

50 (8)

0

34

24

76

20

3

0

14

48

38

21

14

H 38

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

263

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Page 264

Lasiodiscus mannii

Hook.f.

Rhamnaceae Description

Habitat

Guild: sb Life form: shrub or small tree Max. height: 10 m (herbarium) Max. diameter: 25 cm (herbarium) Leaf: opposite, simple, elliptic to oblanceolate, mesophyll (4.5-9 x 12-23 cm), dentate, coriaceous, glabrous or slightly hairy on the midrib above, hairy on the whole surface or on the nerves beneath; petiole hairy (0.4-1.6 cm long); with interpetiolar stipules Inflorescence: axillary, branched (10 cm long) Flower: small (0.6-1 cm in diameter); white with brown hairs Fruit: dry dehiscent (explosive capsule), globose (1.4 cm in diameter), dull yellow to green, densely covered in short hairs Seed: medium-sized (1 x 0.8 x 0.5 cm), reddish brown Other: the bole is black, the bark red-stained. It has hairy twigs and young branchlets with conspicuous white lenticels.

Found in deep shade. It occurs frequently along riversides and in swamps (herbarium).

Regeneration It regenerates in shade, and sometimes occurs gregarious in the understorey of dry forests (Hall & Swaine 1981).

Phenology Dispersal: explosive

Distribution Continent: Guineo-Congolian wide: from Côte d’Ivoire to Democratic Republic of Congo and Tanzania (Hall & Swaine 1981). Central African Republic, Cameroon, Equatorial Guinea, Sao Tomé and Principe, Gabon, Congo (Brazzaville), Democratic Republic of Congo (herbarium) Upper Guinea: Liberia, Côte d’Ivoire (herbarium) Distribution type: continental disjunct, in Upper Guinea present in 4 30’ cells Forest type: moist evergreen forest, moist semideciduous forest, dry semi-deciduous forest, gallery forest, swamp forest, secondary forest. In Ghana, most abundant in Southern Marginal forests (Hall & Swaine 1981).

L

spp

L.m. All

264

n

Open (n)

Data sources FWTA, Hall & Swaine (1981)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

4

0 (2)

0

75

0

25

25

25

25

0

100

0

50

25

25

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 265

Leptoderris miegei

Aké Assi & Mangenot

Leguminosae-Pap. Description

Phenology

Guild: np Life form: winding woody climber Max. height: 15 m long (Aké Assi & Mangenot 1975) Max. diameter: data unavailable Leaf: alternate, imparipinnately compound, 9-13 opposite leaflets, elliptic, microphyll (1.5-3 x 5-10 cm), entire, coriaceous, with long hairs at the lower leaf surface Inflorescence: axillary or terminal, branched, hairy Flower: small; corolla white to purple Fruit: dry indehiscent, flat, wing-like (8 x 3 cm), brown; 1-2 seeds Seed: medium-sized (0.8 cm), brown Other: a climbing shrub when young. The end of young twigs is covered with golden-brown hairs.

Data sources FWTA, Aké Assi & Mangenot (1975), Hawthorne (1995a), Lock (1989)

Distribution Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire, Ghana Distribution type: continuous, regional, present in 5 30’ cells, distribution range is 511 km Forest type: rainforest, secondary forest

Habitat In Côte d’Ivoire, it is found in the lagoon region, though not always near water (Aké Assi & Mangenot 1975). It is fairly common in Ghana in swamps of evergreen forest zones (Hawthorne 1995a).

spp

n

Open (n)

L

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

L.m.

12

50 (6)

0

83

8

0

16

83

0

50

50

0

8

42

42

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

265

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Page 266

Leucomphalos libericus

Breteler

Leguminosae-Pap.

Habitat

Description

Data unavailable. Guild: u Life form: winding woody climber Max. height: 10 m long (herbarium) Max. diameter: data unavailable Leaf: alternate, simple, elliptic with a caudate apex, mesophyll (3.5-8 x 7-17 cm), entire, coriaceous; triangular stipules, falling soon Inflorescence: axillary, unbranched (raceme, up to 10 cm long) Flower: small; calyx pale brown with rusty pubescence; corolla white with yellow spots near the base Fruit: dry dehiscent (pod); 1-2 seeds Seed: data unavailable

L

Phenology

Distribution Data sources

Continent: Upper Guinea endemic Upper Guinea: Liberia, Côte d’Ivoire Distribution type: present in 2 30’ cells Forest type: wet evergreen forest

spp

266

n

Open (n)

FWTA, Breteler (1994b), Hawthorne & Jongkind (2004)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

L.l.

2

100 (3)

0

50

0

0

50

0

50

0

100

0

50

0

50

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 267

Loesenera kalantha

Harms

Leguminosae-Caes. Description

Phenology

Guild: u Life form: medium-sized tree Max. height: 30 m (Voorhoeve 1965) Max. diameter: 70 cm (herbarium) Leaf: alternate, paripinnately compound, 6-8 leaflets, oblong to lanceolate, notophyll (3-4 x 8-10 cm), entire, coriaceous, rusty brown hairs below, twisted petiolule, knotted vein glands (especially on midrib) Inflorescence: terminal, unbranched (raceme) Flower: medium-sized; bracts and calyx reddish brown; corolla pinkish Fruit: dry dehiscent, oblong (7 x 17 cm), brown, woody, with short pale brown hairs; 1-2 seeds Seed: very large (3-5 cm long), lenticular Other: the bole is cylindrical, straight or swollen at the base. The bark is greenish grey, smooth and hard, the slash light brown. New leaves appear in drooping, pink-red flushes. The species resembles Tetraberlinia tubmaniana.

Data sources FWTA, Voorhoeve (1965), Lock (1989), IUCN Red List (2000), Hawthorne & Jongkind (2004)

Distribution Continent: Upper Guinea endemic Upper Guinea: Liberia, Côte d’Ivoire Distribution type: continuous, regional, present in 5 30’ cells, distribution range is 315 km, Red List species (Vulnerable). In eastern Liberia, it may form gregarious stands (Voorhoeve 1965). Forest type: swamp forest, in savanna near rivers

L Habitat It is found mainly along creeks, in swamps or in depressions that become inundated during the rainy season (herbarium).

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

L.k.

10

50 (4)

20

20

40

0

20

20

50

0

100

0

60

0

H 0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

267

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Page 268

Macropodiella garrettii

(C.H.Wright) C.Cusset

Podostemaceae Description

Data sources

Guild: u Life form: aquatic herb Max. height: 0.1 m (FWTA) Leaf: dimorphic (0.2 cm long), irregularly lobed, herbaceous Inflorescence: terminal, branched (fascicle), on a long stalk Flower: small; 2 tepals, bright green Fruit: dry dehiscent (capsule), ellipsoid (0.18 cm long), with 2 valves; many seeds Seed: small (0.008 x 0.013 cm), slightly compressed Other: a branched herb with horny, compressed bright green stems. It is soon deciduous and appears leafless.

FWTA, Cusset (1977)

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Côte d’Ivoire Distribution type: present in 4 30’ cells Forest type: rainforest

Habitat Growing submerged or at water level on rocks, near waterfalls or in fast-flowing rivers.

Phenology

M

spp

268

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

M.g.

4

100 (1)

0

50

25

0

50

0

50

0

25

75

25

50

25

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 269

Maesa nuda

Hutch. & Dalziel

Myrsinaceae Description

Phenology

Guild: u Life form: woody climber Max. height: 5 m long, 5 m high (herbarium) Max. diameter: 2 cm (herbarium) Leaf: alternate, simple, obovate, notophyll (1.5-6 x 2.5-13 cm), entire, herbaceous Inflorescence: axillary, branched (approx. 11 cm long) Flower: small (0.2 cm long); corolla white Fruit: fleshy (0.5 cm long), white; 1 seed Seed: medium-sized (0.2-0.3 cm) Other: a climber that probably starts as a shrub.

Distribution

Data sources

Continent: Cameroon Upper Guinea: Senegal, Guinea, Côte d’Ivoire Distribution type: continental disjunct, in Upper Guinea present in 2 30’ cells Forest type: gallery forest, swamp forest

FWTA

Habitat Usually, found in periodically inundated forests on sandy soils (herbarium). It has also been recorded in open (Côte d’Ivoire) to very degraded (Cameroon) mountain forests (herbarium).

M

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

M.n.

5

25 (4)

20

0

0

60

0

40

0

0

40

60

40

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

269

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Page 270

Magnistipula cupheiflora

Mildbr. ssp.

leonensis

F.White

Chrysobalanaceae Description

Data sources

Guild: u Life form: medium-sized tree Max. height: 30 m (herbarium) Max. diameter: 40 cm (herbarium) Leaf: alternate, simple, elliptic, mesophyll (3-6 x 8.5-22 cm), entire, coriaceous, glabrous, usually without glands at base, stipules Inflorescence: axillary, raceme, hairy Flower: small; calyx greenish; corolla white Fruit: fleshy, large, edible; 1 seed Seed: large to very large Other: it has a dark-red slash and a rough, brown bark.

White (1976), Hawthorne & Jongkind (2004)

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Côte d’Ivoire Distribution type: continuous, very local, present in 3 30’ cells Forest type: rainforest

Habitat Data unavailable.

Phenology Dispersal: by animals

M

spp

270

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

M.c.

3

0 (1)

0

33

0

0

33

0

66

0

100

0

100

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 271

Magnistipula zenkeri

Engl.

Chrysobalanaceae Description

Phenology

Guild: u Life form: medium-sized to large tree Max. height: 20 m (Vivien & Faure 1985) Max diameter: 100 cm (Vivien & Faure 1985) Leaf: alternate, simple, elliptic to oblong, macrophyll (8-22 x 20-44 cm), entire, coriaceous, glabrous, 6-12 lateral nerves, several glands along the lamina, leafy stipules, the new leafs flush purple Inflorescence: terminal, unbranched (raceme) Flower: medium-sized; calyx pale pink; corolla white-blue Fruit: fleshy, ovoid (5 x 8 cm), pale green, glabrous; 1 seed Seed: very large (2.8 x 5.5 cm), hairy

Data sources

Distribution

White (1976), Vivien & Faure (1985), Hawthorne & Jongkind (2004)

Continent: Cameroon, Equatorial Guinea, Gabon Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continental disjunct, in Upper Guinea present in 5 30’ cells, a rare species (Hawthorne & Jongkind 2004) Forest type: wet evergreen forest

Habitat Found at the border of rivers and swamps (Vivien & Faure 1985, Hawthorne & Jongkind 2004).

M

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

M.z.

6

0 (1)

0

50

0

17

50

17

17

0

67

33

17

0

50

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

271

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Page 272

Malaxis melanotoessa

Summerh.

Orchidaceae Description

Data sources

Guild: u Life form: terrestrial herb Max. height: data unavailable Leaf: alternate, simple, lanceolate, microphyll (1-2 x 2-5 cm), finely crenate Inflorescence: terminal, not branched, erect and gracious (3-6 cm long) Flower: small; corolla pale yellow; lip dark purple at insertion and becoming creamish at length Fruit: dry dehiscent (capsule), elliptic, green-brown Seed: very small

FWTA, Cribb (1978), Summerhayes (1958)

Distribution Continent: Upper Guinea endemic Upper Guinea: Liberia Distribution type: continuous, very local, present in 1 30’ cell. It has been found only in the Gola Forest of Liberia approx. 28 km south of Ba by Bunting Forest type: data unavailable

Habitat It has been found growing on humus on very wet rocks (herbarium).

Phenology Dispersal: by wind

M

spp

272

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

M.m

1

- (0)

0

100

0

0

0

0

100

0

100

0

100

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 273

Mangenotia eburnea

Pichon

Asclepiadaceae Description

Regeneration

Guild: np Life form: small winding woody climber Max. height: 5 m long (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, oblong, microphyll (1-2.3 x 2.5-7 cm), entire, herbaceous Inflorescence: axillary or terminal, branched Flower: medium-sized (1-1.5 cm long); calyx pale green; corolla creamy white; tube with lobes Fruit: dry dehiscent, oblong (0.8 x 13 cm); many seeds Seed: medium-sized, plumed (long hairs) Other: an erect shrub when young. The slash exudes a white latex.

It regenerates in the shade (Hall & Swaine 1981).

Phenology Dispersal: by wind (Hall & Swaine 1981)

Distribution Continent: Nigeria Upper Guinea: Senegal, Guinea-Bissau, Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 22 30’ cells, distribution range is 2296 km Forest type: In Côte d’Ivoire and Senegal, it has been frequently found in coastal forests, and secondary forests (herbarium). In Ghana, however, it is only found in the moist and dry semi-deciduous forests and the dry South Marginal forest (Hall & Swaine 1981).

Data sources FWTA, Hall & Swaine (1981)

Habitat It is found in the forest understorey and also in open places (e.g. clearings, forest edges, low bushlands). Most often at lowlands (herbarium) and usually not close to water (Chi2 test). Often on sandy soils (herbarium).

spp

n

Open (n)

M

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

M.e.

31

30 (20)

0

16

16

42

26

10

22

19

45

35

23

35

H 32

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

273

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Page 274

Manotes macrantha

(Gilg) Schellenb.

Connaraceae Description

Distribution

Guild: np Life form: large woody climber Max. height: 30 m long (herbarium) Max. diameter: 10 cm (herbarium) Leaf: alternate, imparipinnately compound, 5-9 leaflets, ovate, mesophyll (4-7 x 7.5-19 cm), entire, coriaceous, glabrous; no stipules Inflorescence: axillary, branched (approx. 8 cm long) Flower: medium-sized; corolla yellow Fruit: dry dehiscent (follicle), 1-5 fruits per flower (2 x 1 cm), yellowish, velvety, after opening the seed remains attached by the top of the aril; 1 seed Seed: medium-sized (1 x 0.7 cm), outer layer fleshy and red, aril bright yellow Other: mainly climbing with leaves that have been modified to form strong woody hooks.

Continent: Gabon Upper Guinea: Liberia, Côte d’Ivoire Distribution type: continental disjunct, in Upper Guinea present in 9 30’ cells Forest type: wet evergreen forest, rainforest, coastal forest, secondary forest, gallery forest, savanna

Habitat It regenerates in disturbed (savanna, forest edge, secondary forest, roadsides) as well as in undisturbed habitats. It is often found by rivers on sandy soils (herbarium).

Phenology Dispersal: probably by animals

M

Data sources FWTA, Jongkind (1989), Hawthorne & Jongkind (2004)

spp

274

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

M.m

16

50 (10)

0

25

63

0

6

19

43

0

100

0

31

0

6

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 269-278

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Page 275

Mapania baldwinii

Nelmes

Cyperaceae Description

Habitat

Guild: sb Life form: perennial rhizomatous herb Max. height: 0.8 m Leaf: basal, simple, linear to oblong, mesophyll (4-8 x 19-53 cm), entire or scabrid, coriaceous, 3-nerved, dark-green with reddish nerves and dark green to purple leaf sheaths, young leaves are redbrown with a red petiole Inflorescence: solitary shoot or culm terminating in an inflorescence; terminal, globose (1.8-3.3 cm wide), light brown, with spikes; 4 floral bracts, individual florets monoecious Flower: green to dull red, glabrous Fruit: dry indehiscent, subglobose to globose, black, exocarp hard and strongly tuberculated Seed: small (0.13 x 0.12 cm)

Species occurrence is highest where rainfall is between 1500-2000 mm/yr (Chi2 test). Usually in moist (to damp) and very shady places in wet evergreen forests (e.g. stream banks) (herbarium, Simpson 1992). Generally found in mature forests but sometimes also along logging roads in exploited forest. In Côte d’Ivoire, this species is especially found on slopes (Lorougnon 1972). In Côte d’Ivoire, wet forest types on clay soils are characterised by Mapania species (Mangenot, in Lorougnon 1972). The species is found up to 60 m altitude.

Distribution

Phenology

Continent: Upper Guinea endemic (Hall & Swaine 1981) Upper Guinea: Liberia, Côte d’Ivoire, Ghana (herbarium, Hall & Swaine 1981, Simpson 1992) Distribution type: continuous, widespread, present in 23 30’ cells, distribution range is 979 km Forest type: wet evergreen forest, moist evergreen forest, secondary forest. In Ghana mostly in wet evergreen forests (Hall & Swaine 1981). It is locally common.

Deciduousness: evergreen Pollination: probably pollinated by wind (Hall & Swaine 1981) and insects (Simpson 1992) Dispersal: not known (Hall & Swaine 1981) Timing: not clearly seasonal (Hall & Swaine 1981)

Regeneration It regenerates in the shade.

Data sources Lorougnon (1972), Hall & Swaine (1981), Simpson (1992)

M

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

M.b.

43

13 (24)

2

42

0

2

77

19

2

2

63

35

49

2

40

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

275

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Page 276

Mapania ivorensis

(Raynal) Raynal

Cyperaceae Description

Phenology

Guild: sb Life form: perennial rhizomatous herb Max. height: 1.8 m (Simpson 1992) Leaf: basal, simple, linear, macrophyll (3.5-6 x 43-90 cm), entire to finely serrate, coriaceous, 3nerved with dark brown veins; leaf converts at the base in a pseudopetiole Inflorescence: solitary shoot or culm, erect and central (4-6 x 0.2-0.5 cm), green and glabrous, arising from the axil of the basal leaves and terminating in bracts and an inflorescence; terminal, globose (2-5.5 cm wide), brown, with numerous spikes, 4 floral bracts Flower: white to bright brown Fruit: dry indehiscent (0.15 x 0.1 cm), obovoid to globose, black, exocarp hard and strongly tuberculate; 1 seed Seed: small (0.11 x 0.09 cm)

Dispersal: pollination probably by insects (Simpson 1992)

Data sources Lorougnon (1972), Simpson (1992)

Distribution Continent: Upper Guinea endemic Upper Guinea: Liberia and Côte d’Ivoire (herbarium, Simpson 1992) Distribution type: continuous, widespread, present in 12 30’ cells, distribution range is 826 km Forest type: wet evergreen forest, old secondary forest

Habitat Species occurrence is highest at places where rainfall is between 1500-2000 mm/yr (Chi2 test). Found in moist and shady places (Simpson 1992, herbarium). Commonly recorded in old secondary forests, usually under deep shade and at very humid places. Occasionally also in open forests and forest edges (herbarium). Occurs at altitudes up to 60 m (Simpson 1992).

M

spp

276

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

M.i.

29

45 (11)

0

28

3

0

83

10

6

0

41

59

31

0

69

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 277

Mapania linderi

Hutch. ex Nelmes

Cyperaceae Description

Phenology

Guild: sb Life form: perennial rhizomatous herb Max. height: 1.3 m (Simpson 1992) Leaf: basal, simple, linear (1.7-3.5 x 31-70 cm), finely serrate, coriaceous, 3-nerved; leaf blade dark green with veins brown to black; leaf base transformed in a pseudopetiole Inflorescence: erect and lateral shoots or culms (0.1-0.15 x 8-21 cm), green to reddish brown, glabrous, arising from the axil of the basal leaves and terminating in bracts and an inflorescence, subglobose, terminal (1.5-2.4 cm wide), with spikes, 4 floral bracts Flower: whitish to bright brown Fruit: dry indehiscent obovoid (0.13 x 0.09 cm), black, exocarp hard and weakly tuberculate; 1 seed Seed: small Other: the rhizome grows always vertical. This species is often confounded with Maschalocephalus dinklagei, which shares the same habitat.

Pollination: probably by insects (Simpson 1992)

Uses In Liberia it is used for thatching (Simpson 1992).

Data sources Lorougnon (1972), Simpson (1992)

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire (herbarium, Simpson 1992) Distribution type: continuous, widespread, present in 18 30’ cells, distribution range is 825 km Forest type: rainforest, riverine forest, secondary forests. It is locally common.

Habitat

M

It is found mainly at high altitudes (logistic regression analysis, Chi2 test). In very humid and shady places of the forest (Simpson 1992, herbarium), often also in swampy valley bottoms (Lorougnon 1972). On white sand or sandy clay soils (Simpson 1992, herbarium). It occurs at altitudes between 80 and 500 m.

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

M.l.

38

30 (20)

21

37

5

0

42

45

13

0

76

24

58

0

H 32

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

277

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Mapania minor

Page 278

(Nelmes) J.Raynal

Cyperaceae Description

Phenology

Guild: sb Life form: perennial rhizomatous herb Max. height: 1.4 m Leaf: basal, simple, linear macrophyll (2.1-3.6 x 66-128 cm), entire to finely serrate, coriaceous, 3nerved; leaves above glossy and dark green, beneath dull and bright green, with black veins; at base transformed in a pseudopetiole Inflorescence: solitary shoot or culm, erect and central (0.2-0.35 x 22-53 cm), arising from the axil of the basal leaves and terminating in bracts and an inflorescence, green to mid-brown, glabrous, terminal, globose (2.5-4 cm wide), dull brown, with numerous spikes, floral bracts 4 Flower: data unavailable Fruit: dry indehiscent, obovate (0.1 x 0.1 cm), black, exocarp hard and rugulose; 1 seed Seed: small Other: the rhizome grows vertical.

Pollination: probably by insects (Simpson 1992)

Data sources Lorougnon (1972), Simpson (1992)

Distribution Continent: Upper Guinea endemic Upper Guinea: Liberia, Côte d’Ivoire (herbarium, Simpson 1992) Distribution type: continuous, regional, present in 10 30’ cells, distribution range is 384 km Forest type: rainforest, secondary forest

Habitat Species occurrence is higher where rainfall is between 1500-2000 mm/yr (Chi2 test). It is found in moist and shady places of the forest (Simpson 1992, herbarium).

M

spp

278

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

M.m

13

13 (8)

0

62

15

0

69

30

0

0

69

31

46

0

38

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 279-288

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Page 279

Mapania rhynchocarpa

Lorougnon & J.Raynal

Cyperaceae Description

Phenology

Guild: sb Life form: perennial rhizomatous herb Max. height: 1.1 m Leaf: basal, simple, linear, macrophyll (2.9-3.2 x 60-93 cm), entire to finely serrate, coriaceous, 3-nerved Inflorescence: solitary shoot or culm, erect and central (0.35 x 43 cm), arising from the axil of the basal leaves and terminating in bracts and an inflorescence, green, glabrous; terminal, globose (4-6 cm wide), whitish, with numerous spikes, floral bracts 4 Flower: data unavailable Fruit: dry indehiscent, obovoid (0.19 x 0.14 cm), exocarp hard and tuberculate, the whole style may remain attached to the fruit, dark greenish-brown; 1 seed Seed: small Other: this species resembles M. ivorensis (Lorougnon 1972).

Pollination: probably by insects (Simpson 1992)

Data sources Lorougnon (1972), Simpson (1992)

Distribution Continent: Nigeria (herbarium, Simpson 1992) Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire (herbarium), Ghana (Simpson 1992) Distribution type: continental disjunct, in Upper Guinea present in 4 30’ cells Forest type: rainforest

Habitat

M

Found in moist and shady places of the forest.

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

M.r.

5

- (0)

20

80

0

0

80

20

0

0

20

80

20

0

60

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

279

22466 layout page 279-288

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Page 280

Maranthes aubrevillei

(Pellegr.) Prance

Chrysobalanaceae Distribution Continent: Upper Guinea endemic. According to Hall & Swaine (1981) it also occurs in Cameroon but this may be a doubtful sterile record Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana (herbarium) Distribution type: continuous, widespread, present in 10 30’ cells, distribution range is 1141 km Forest type: wet evergreen forest, moist evergreen forest. In Ghana, mainly present in wet evergreen forest (Hall & Swaine 1981). In Liberia, the species is locally common (Voorhoeve 1965).

Habitat Found on sandy clay and laterite soils (herbarium).

Regeneration It can regenerate in the shade (Hall & Swaine 1981).

Phenology Dispersal: probably by primates (Hawthorne & Jongkind 2004), and elephants

Description Guild: sb Life form: medium-sized to large tree Max. height: 30 m (Voorhoeve 1965) Max diameter: 80 cm (Voorhoeve 1965) Leaf: alternate, simple, elliptic, notophyll (3.5-6.5 x 6.5-13.5 cm), entire with dentate apex, coriaceous, with loose cottony hair on underside, very young leaves pale pinkish, two extrafloral nectaries at leaf base; conspicuous stipules Inflorescence: terminal or axillary, branched (manybranched cyme) Flower: medium-sized, bowl-shaped, creamcoloured, pubescent

M

Data sources

Fruit: fleshy Seed: data unavailable Other: it has root swellings at the tree base. The slash is deeply red. The crown is dark and dense, and the twigs are hairy.

FWTA, White (1976), Voorhoeve (1965), Hall & Swaine (1981), Hawthorne & Jongkind (2004)

?

spp

280

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

M.a.

21

11 (9)

0

33

0

5

53

10

33

0

81

19

71

0

29

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 279-288

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Page 281

Marantochloa cuspidata

(Rosc.) Milne-Redh.

Marantaceae Description

Habitat

Data sources

Guild: pi Life history: large perennial rhizomatous herb Max. height: 2 m (herbarium) Max. diameter: data unavailable Leaf: only 2 leaves, simple, ovate to oblong, macrophyll (7-20 x 15-50 cm), entire, coriaceous, glabrous Inflorescence: branched (up to 102 cm long) Flower: medium-sized (1.5 cm long); tube-shaped with lobes; calyx translucent and brownish with red tips when young, hairy; corolla yellow Fruit: fleshy, globose (0.8 x 0.8 cm), yellow, with white hairs; 3 seeds Seed: medium-sized (0.5 x 0.4 cm), with an aril Other: an erect herb with a creeping rhizome

Species occurrence increases with rainfall to reach an optimum around 2700 mm/yr (logistic regression analysis, Chi2 test). It is found in a wide range of moisture conditions. It is usually found in very moist places such as river borders, along streams, and swamps, but in Ghana it is common in dry semi-deciduous forests (Hall & Swaine 1981). Very often, it is found in shady places in the understorey but it is also common in forest gaps. Soils can be alluvial, clayish, loamy, sandy lateritic, or rocky (herbarium).

FWTA, Hall & Swaine (1981), Hawthorne (1995a), Dhetchuvi (1996)

Distribution Continent: Upper Guinea endemic Upper Guinea: Senegal (Dhetchuvi 1996), Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana (herbarium) Distribution type: continuous, widespread, present in 46 30’ cells, distribution range is 1554 km Forest type: rainforest, moist semi-deciduous forest, dry semi-deciduous forest, dry forest, secondary forest, riverine forest. In Ghana, it is only found in semi-deciduous forests (Hall & Swaine 1981), especially in burnt or otherwise disturbed dry forests (Hawthorne 1995a).

Regeneration In Ghana, it regenerates mainly in gaps, and on the forest floor after ground fire (Hall & Swaine 1981). It tends to form clumps in the forest (herbarium).

Phenology Timing: not clearly seasonal (Hall & Swaine 1981)

M

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

H

M.c.

87

29 (52)

6

44

2

8

30

29

30

1

71

28

56

3

23

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

281

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Page 282

Marattia odontosora

Christ

Marattiaceae Description Guild: u Life form: large perennial herb (fern) Max. height: data unavailable Leaf: bipinnately compound (4 pairs of pinnae on either side), up to 30 cm long, serrate Spores: minute Other: it has fleshy mycorrhizal roots and multicellular root hairs. The leaf stems are covered with small spines nearly 0.5 mm long.

Distribution Continent: Cameroon Upper Guinea: Guinea, Sierra Leone Distribution type: continental disjunct with the largest population in Upper Guinea, present in 5 30’ cells in Upper Guinea Forest type: data unavailable

Habitat It is common above 600 m (FWTA)

Phenology Data unavailable.

Data sources FWTA

M

spp

282

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

M.o.

6

- (0)

50

17

0

0

17

84

0

0

17

83

17

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 283

Maschalocephalus dinklagei

Gilg & K.Schum.

Rapateaceae Description

Phenology

Guild: sb Life form: rhizomatous herb Max. height: data unavailable Leaf: rosette, simple, linear to lanceolate, mesophyll (1-5 x 20-75 cm), entire, coriaceous, dark green above, pale green beneath; with darker veins Inflorescence: axillary, sessile, flowers arranged in a head Flower: small; corolla yellow with white pedicel; anthers bright yellow Fruit: dry dehiscent, triangular Seed: data unavailable

Data sources FWTA

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire Distribution type: continuous, widespread, present in 11 30’ cells, distribution range is 959 km Forest type: wet evergreen forest, moist evergreen forest, savanna

Habitat Found in shaded places. It prefers moist to wet habitats (e.g. streams, swamps, marshy places). Sometimes reported in open savanna, but always in swamps on loamy sand (e.g. Liberia, Côte d’Ivoire).

M

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

M.d.

15

29 (7)

7

27

7

0

47

26

20

0

73

27

67

0

27

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

283

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Page 284

Memecylon aylmeri

Hutch. & Dalziel

Melastomataceae Description

Regeneration

Guild: sb Life form: pigmy tree or shrub Max. height: 3.5 m (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, oblong to elliptic, notophyll (3.5-6 x 7-13 cm), entire, coriaceous; petiole 0.5 cm long Inflorescence: axillary, umbel formed by small groups of flowers (approx. 0.5 cm long) Flower: small (approx. 0.3 cm); corolla dishshaped, cream to pale blue Fruit: fleshy, ellipsoid (0.6 x 1.5 cm), blue; 1 seed Seed: medium-sized (0.6 x 0.4 cm) Other: the branches are quadrangular.

It has a phanerocotylar epigeal foliaceous seedling type (cf. Jacques-Félix 1978).

Phenology

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: Upper Guinea disjunct, widespread, present in 17 30’ cells, distribution range is 1307 km Forest type: rainforest, swamp forest, secondary forest. In Ghana, it is mostly found in evergreen forest (wet, moist, and upland), and scarcely in (moist and dry) semi-deciduous forests (Hall & Swaine 1981).

Data sources FWTA, Jacques-Félix (1978), Hall & Swaine (1981)

Habitat Species occurrence increases at higher altitudes (logistic regression analysis, Chi2 test) and with rainfall higher than 2000 mm/yr (logistic regression analysis, Chi2 test). Often found in wet places in the rainforest such as along rivers and even swamps, but occasionally in relatively dry forests (herbarium).

M

spp

284

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

M.a.

30

25 (12)

27

43

3

3

23

47

23

0

63

37

53

0

H 20

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 285

Mendoncia combretoides

(A.Chev.) Benoist

Acanthaceae Description

Phenology

Guild: np Life form: large woody winding climber Max. height: 30 m long (herbarium) Max. diameter: 20 cm (De Koning 1983) Leaf: opposite, simple, elliptic to ovate, notophyll (3-7.5 x 6.5-12 cm), entire, coriaceous; hairy on the nerves and the petiole Inflorescence: axillary, unbranched Flower: medium-sized (corolla tube 2 cm long); calyx and bracts green with dark brown hairs; corolla white Fruit: fleshy indehiscent, ovoid (2.3 x 1.7 cm), black; 1 seed Seed: large (1 x 2 cm) Other: the stem is green with brown hairs.

Dispersal: probably by birds and monkeys (Van Roosmalen 1985)

Distribution

Data sources

Continent: Upper Guinea endemic Upper Guinea: Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 14 30’ cells, distribution range is 889 km Forest type: moist and wet evergreen forest, secondary forest (herbarium)

FWTA, De Koning (1983), Van Roosmalen (1985)

Habitat Species occurrence is higher in forests where rainfall is between 1500 - 2500 mm/yr (Chi2 test). Often found near rivers or even swampy areas (herbarium). On soils with high water holding capacity (Chi2 test).

M

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

M.c.

32

24 (21)

3

53

0

0

57

43

0

0

59

41

25

0

66

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

285

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Milicia regia

Page 286

(A.Chev.) C.C.Berg

Moraceae Phenology

(0.6 x 15 cm long), female inflorescence (0.5 x 1-4.5 cm long) Flower: dioecious; male flowers small (1 x 2 cm), yellow, densely pubescent; female flowers green, hairy Fruit: fleshy (8 x 2.5 x 1.2 cm), sweet and with strong fermenting smell; many seeds Seed: small (approx. 0.2 cm long) Other: it has a straight and cylindrical bole. The crown has outward sweeping layers of dark foliage. The bark is rough, grey, with large, reddish-brown conspicuous lenticels. The slash is very gritty and exudes a rapid, watery white latex.

Deciduousness: apparently it is deciduous for a short period during the dry season (Voorhoeve 1965) Dispersal: probably by animals Timing: flowering period from January to March, later (till June) in Guinea and Senegal (Berg 1982)

Distribution Continent: Upper Guinea endemic Upper Guinea: Senegal, Gambia, Guinea-Bissau, Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 26 30’ cells, distribution range is 1982 km, Red List species (Vulnerable) Forest type: rainforest, moist semi-deciduous forest, coastal savanna, gallery forest (especially in the northwestern part of the range), secondary forest

Uses The bark is used for several medical purposes. The tree produces excellent timber for construction and railway sleepers, floors, etc. The wood is resistant against fungus and boring insects, though not immune to termites (Voorhoeve 1965). The sweet infructescences attract game (Berg 1982).

Habitat Description Guild: np Life history: large tree Max. height: 61 m (herbarium) Max. diameter: 200 cm (De Koning 1983) Leaf: alternate, simple, oblong to elliptic, mesophyll (3-13 x 4-28 cm), almost entire to crenate, herbaceous, glabrous above, sparsely hairy on the veins beneath, midrib prominent, reddish, pubescent below; petioles 0.8-3.6 cm; stipules Inflorescence: dioecious, axillary, unbranched (pendulous spike) pendulous, male inflorescence

M

spp

286

It is commonly found in open places (De Koning 1983) and has been described as a light demander (Voorhoeve 1965). Often found in cleared or even very degraded hydrophile forests, but occasionally also recorded in dense forests (herbarium).

n

Open (n)

Data sources Voorhoeve (1965), Hall & Swaine (1981), Berg (1982), De Koning (1983), UCN Red List (2000), Hawthorne & Jongkind (2004)

Regeneration It has a phanerocotylar epigeal foliaceous seedling type (cf. Voorhoeve 1965). Regeneration is fairly common, especially in open places such as gaps, logging roads, and low bushes (Voorhoeve 1965).

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

M.r.

48

39 (31)

8

46

10

31

26

25

17

8

52

40

29

10

H 42

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 279-288

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Page 287

Millettia leonensis

Hepper

Leguminosae-Pap. Description

Phenology

Guild: u Life form: small tree, sometimes climbing Max. height: data unavailable Max. diameter: data unavailable Leaf: alternate, imparipinnately compound, 5-7 opposite leaflets, ovate to elliptic, microphyll (2.53.5 x 5-9 cm), herbaceous, pubescent beneath, rachis with appressed reddish to purplish hairs Inflorescence: axillary, branched Flower: calyx brown; corolla purple, petals on the outside covered with purplish hairs Fruit: dry dehiscent, flat, woody, young pods densely pilose Seed: data unavailable

Data sources

Distribution

FWTA, Lock (1989), Hawthorne & Jongkind (2004)

Continent: Upper Guinea endemic Upper Guinea: Sierra Leone Distribution type: continuous, very local, present in 2 30’ cells Forest type: high forest

Habitat One of the two known specimens was collected along a river’s edge (herbarium).

M

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

M.l.

2

50 (2)

0

100

0

0

0

0

100

0

100

0

50

0

50

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

287

22466 layout page 279-288

11/11/03

12:15 PM

Millettia liberica

Page 288

Jongkind

Leguminosae-Pap. Distribution Continent: Upper Guinea endemic Upper Guinea: Liberia, Côte d’Ivoire Distribution type: continuous, local, present in 4 30’ cells, distribution range is 209 km Forest type: moist evergreen forest, secondary forest

Habitat It occurs in undisturbed forest, as well as in gaps. Found in secondary forest and along roadsides. Sometimes found near rivers or swamps.

Phenology

Description Guild: sb Life form: small tree Max. height: 10 m (herbarium) Max. diameter: 8 cm (herbarium) Leaf: alternate, imparipinnately compound, 5-7 opposite leaflets, elliptic, herbaceous, almost glabrous beneath Inflorescence: axillary, branched Flower: calyx whitish with purple spots; corolla violet Fruit: dry dehiscent, flat, woody Seed: data unavailable Other: it has twigs with many small lenticels.

M

spp

288

n

Open (n)

Data sources Hawthorne & Jongkind (2004)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

M.l.

6

67 (6)

0

0

0

0

100

0

0

0

100

0

83

0

17

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 289-298

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Page 289

Millettia pallens

Stapf

Leguminosae-Pap. Description

Phenology

Guild: pi Life form: shrub or small tree Max. height: 5 m (herbarium) Max. diameter: 8 cm (herbarium) Leaf: alternate, paripinnately compound, 11-13 leaflets, ovate to obovate, notophyll (2.8-5 x 5.511.5 cm), coriaceous, glabrous Inflorescence: axillary, unbranched (up to 14 cm long) Flower: medium-sized; calyx with brownish spots; young flowers reddish purple, later turning pale purple or pink Fruit: dry dehiscent (1.5 x 5 cm), woody, dark green with a wavy margin; 4-5 seeds Seed: disk-shaped with palmate scratches

Data sources

Distribution

Hawthorne & Jongkind (2004)

Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire Distribution type: continuous, widespread, present in 24 30’ cells, distribution range is 684 km Forest type: secondary forest, thicket, beach forest, gallery forest, savanna, savanna-woodland

Habitat Species occurrence increases strongly in places where rainfall is 2500 mm/yr or higher (logistic regression analysis, Chi2 test). It is often found in disturbed places (e.g. along roadsides, farm bushes, and villages), and sometimes found near rivers or near the coast. Usually, on soils with low water holding capacity (Chi2 test) such as sandy or lateritic soils (herbarium).

M

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

H

M.p.

59

82 (33)

3

41

12

0

2

8

90

0

68

32

58

12

12

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

289

22466 layout page 289-298

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Page 290

Monanthotaxis whytei

(Stapf ) Verdc.

Annonaceae Habitat Species occurs most often where rainfall is between 2000-2500 mm/yr (Chi2 test), in lowlands close to the coast (logistic regression analysis and Chi2 test) and rivers (Chi2 test). It is often found in wet places (e.g. wet cliffs, river borders, swamps, lagoons), in older secondary forest as well as in the understorey of mature forest (herbarium). On loamy soils (herbarium).

Regeneration It regenerates in shade (Hall & Swaine 1981).

Phenology Dispersal: probably by animals

Description Guild: sb Life form: large winding woody climber Max. height: 50 m long (herbarium) Max. diameter: data unavailable Leaf: alternate, simple, obovate to elliptic, notophyll (3-6 x 8-13 cm), entire, coriaceous, shining dark green above, beneath grey-green with yellow nerves Inflorescence: cauliflorous, compound Flower: small to medium-sized; corolla yellowish, bowl-shaped Fruit: fleshy (1 x 0.7 cm), orange, several long fleshy sausage-shaped fruit parts per flower, each with a row of seeds inside Seed: medium-sized (0.7 x 0.5 cm) Other: sometimes found as an irregularly winding shrub in thickets. It has a quadrangular stem and tendril forming branchlets.

M

spp

290

n

Open (n)

Distribution Continent: Upper Guinea endemic Upper Guinea: Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 10 30’ cells, distribution range is 1047 km Forest type: moist and wet evergreen forest, semideciduous forest, gallery forest, secondary forest. In Ghana, it is found mostly in wet evergreen forest, but it is also present in moist evergreen and dry semi-deciduous forests (Hall & Swaine 1981, herbarium).

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

M.w

27

47 (17)

0

78

19

7

18

63

4

19

70

11

15

11

H 59

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

Data sources Hall & Swaine (1981)

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Page 291

Monocyclanthus vignei

Keay

Annonaceae Description

Phenology

Guild: sb Life form: small tree Max. height: 9 m (herbarium) Max. diameter: 15 cm (Keay 1953) Leaf: alternate, simple, oblong to elliptic, mesophyll (5-8 x 15-27 cm), entire, coriaceous, glabrous Inflorescence: cauliflorous, solitary Flower: medium-sized (1.4-1.6 cm long), corolla yellow Fruit: with 7 free carpels, hairy Seed: data unavailable Other: it has a straight stem. The bark is smooth with some large lenticels.

Data sources FWTA, Keay (1953), Achenbach & Frey (1992), Hawthorne (1995a), IUCN Red List (2000)

Distribution Continent: Upper Guinea endemic Upper Guinea: Liberia, Ghana (herbarium). Once believed to be endemic to Ghana, now known from wet forest in Liberia. Very rare in Ghana (Hawthorne 1995). Distribution type: Upper Guinea disjunct, widespread, present in 4 30’ cells, distribution range is 1005 km, Red List species (Endangered) Forest type: rainforest (herbarium)

Habitat An understorey tree found in rainforests (herbarium).

M

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

M.v.

7

25 (4)

0

57

0

0

14

14

72

0

86

14

71

0

29

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

291

22466 layout page 289-298

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Page 292

Monosalpinx guillaumetii

N.Hallé

Rubiaceae Description

Phenology

Guild: u Life form: shrub Max. height: 2 m (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, elliptic, notophyll (3.5-5 x 10-14.5 cm), entire, herbaceous; interpetiolar stipules Inflorescence: axillary Flower: medium-sized; yellow-white, 5-lobes Fruit: fleshy, oblong (2 x 4 cm), with longitudinal ridges, orange with persistent calyx; 10 seeds Seed: angular, large (1.2 x 0.8 x 0.5 cm)

Distribution

Data sources

Continent: Upper Guinea endemic Upper Guinea: Liberia, Côte d’Ivoire Distribution type: continuous, local, present in 3 30’ cells, distribution range is 193 km Forest type: wet evergreen forest, secondary forest

Hallé (1968), Hawthorne & Jongkind (2004)

Habitat Data unavailable.

M

spp

292

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

M.g.

6

33 (3)

0

33

0

0

0

83

17

0

100

0

100

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 289-298

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Page 293

Mostuea hymenocardioides

Hutch. & Dalziel

Gelsemiaceae Description

Phenology

Guild: u Life form: shrub Max. height: 5 m (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, microphyll (0.8-2.4 x 1.5-5 cm) Inflorescence: branched (panicle) Flower: corolla white to pink, red inside; unpleasant scent Fruit: dry dehiscent (capsule), medium-brown, shining Seed: data unavailable

Distribution

Data sources

Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone Distribution type: continuous, regional, present in 9 30’ cells, distribution range is 451 km Forest type: rainforest, gallery forest

FWTA

Habitat Found in wet places, near rivers and in flooded areas (herbarium).

M

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

M.h.

15

0 (1)

0

73

0

7

47

20

26

0

40

60

40

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

293

22466 layout page 289-298

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Page 294

Mussaenda afzelii

G.Don

Rubiaceae

Habitat It is mostly found in places where rainfall is around 3000 mm/yr (logistic regression analysis, Chi2 test), but it has been occasionally recorded in dry secondary bush. It is nearly always found in secondary vegetation, along forest edges, or along the road. Usually close to water such as streams, rivers, lakes, swamps, and marshes (herbarium). It is found on valley bottoms, in swamps (De Rouw 1991).

Phenology

Description Guild: pi Life form: winding woody climber Max. height: 7 m (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, elliptic to ovate, notophyll (2.5-5.5 x 6.5-16 cm), entire, coriaceous, with stiff hairs on upper surface, softly beneath; interpetiolar stipules Inflorescence: terminal, branched (flowers in a globose head) Flower: large; calyx green with enlarged calyx lobe white with green veining; corolla tube yellow Fruit: fleshy, globose (1 x 2 cm), ridged, orange to yellow, densely pubescent; many seeds Seed: small (0.05 cm), brown Other: the whole plant is softly pubescent.

M

spp

294

n

Open (n)

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire Distribution type: continuous, widespread, present in 39 30’ cells, distribution range is 967 km. It has a widely scattered distribution, probably because of its association with streams and swamps. Forest type: rainforest, riverine forest, swamp forest, secondary forest

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

M.a.

69

77 (35)

10

42

25

0

24

23

47

1

87

12

57

3

H 23

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

Data sources FWTA, De Rouw (1991)

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Page 295

Mussaenda grandiflora

Benth.

Rubiaceae Description Guild: np Life form: winding woody climber Max. height: 6 m (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, elliptic to ovate, mesophyll (8-11.5 x 9.5-15.5 cm), entire, petiole and veins brown and pubescent; interpetiolar stipules Inflorescence: terminal, flowers in a globose head Flower: medium-sized, calyx green with a large white lobe; corolla bright yellow, hairy outside; scent of primrose Fruit: fleshy, ellipsoid (1.5 x 4.5 cm), green, hairy; many seeds Seed: small Other: the stems are coarsely hairy.

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire Distribution type: continuous, widespread, present in 23 30’ cells, distribution range is 725 km. It can be locally common. Forest type: moist evergreen forest, secondary forest, thickets

Phenology

Habitat Species occurrence increases significantly with rainfall to reach an optimum around 2700 mm/yr (logistic regression analysis, Chi2 test). It is commonly found in open places (e.g. roadsides, and forest edges) and in secondary vegetation. Occasionally found in closed forest. Sometimes found along rivers. Soils can be clayish or loamy (herbarium).

M Data sources FWTA, Hawthorne & Jongkind (2004)

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

L

M

H

36

23

39

2

91

7

82

2

11

24

10

6

69

25

36

13

39

M.g.

44

81 (26)

7

43

5

0

All

46333

37

4

39

7

37

29

Soil WHC

295

22466 layout page 289-298

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Page 296

Mussaenda tristigmatica

H.A.Cummins

Rubiaceae Habitat Species occurrence decreases with altitude (logistic regression analysis) and is higher where rainfall reaches 1500-2000 mm/yr (Chi2 test). It grows commonly in open places of disturbed forests (e.g. forest edges, roadside thickets, low secondary bushes, and occasionally open savanna), but it can sometimes be found in dense wet evergreen forests (herbarium). It has a preference for wet sites such as streams, ponds, or swampy forests (De Koning 1983). On sandy to sandy-clayish soils (herbarium).

Regeneration It has epigeal germination (De Koning 1983).

Phenology Timing: flowering period in July and December; fruiting period from July to August, and December to February (De Koning 1983)

Description Guild: pi Life form: large winding woody climber Max. height: 25 m long (De Koning 1983) Max. diameter: data unavailable Leaf: opposite, simple, usually obovate, mesophyll (4-8.5 x 8.5-14 cm), entire, pilose on the nerves beneath and with scattered hairs above; interpetiolar stipules Inflorescence: terminal, flowers in a globose head Flower: with a conspicuous white flag-like calyxlobe, all other calyx-lobes small leaf-like at least 3 mm wide and with many spreading long hairs; corolla with ascending hairs on the outside, yellow Fruit: fleshy (1.8 x 3.5 cm), orange, densely hairy; many seeds Seed: small, brown Other: it has no exudate. The species is very similar to M. grandiflora.

M

spp

296

n

Open (n)

Distribution Continent: Nigeria Upper Guinea: Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 22 30’ cells, distribution range is 642 km Forest type: wet evergreen forest, moist evergreen forest, savanna, swamp forest, secondary forest, thickets, coastal shrubland

Data sources FWTA, De Koning (1983), Hawthorne & Jongkind (2004)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

M.t.

74

41 (44)

4

50

9

2

62

35

0

12

46

42

4

9

74

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 289-298

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Page 297

Myrianthus libericus

Rendle

Moraceae Description

analysis, Chi2 test). Recorded in both dense forests (De Rouw 1991) and open areas (e.g. roadsides, forest edges). Commonly found in secondary forests (De Koning 1983). Sometimes along rivers. On sandy-stony or sandy-clayish soils (herbarium). In Ghana, its occurrence is strongly associated with base-poor soils (Hall & Swaine 1981, Swaine 1996).

Guild: np Life form: small to medium-sized tree Max. height: 15 m (herbarium) Max. diameter: 25 cm (herbarium) Leaf: alternate, simple, elliptic-lanceolate to oblongobovate, or 3-5 palmately lobed with a distinctly longer middle lobe, macrophyll (8-33 x 17-41 cm), serrate to dentate, coriaceous, sparse hairs above and below, white pubescent; petioles varying in length (up to 24 cm long) Inflorescence: dioecious, axillary, branched; male inflorescence (3 x 11 cm), with flowers in globose groups; female inflorescence (1-2 cm) Flower: male flowers small (approx. 3 mm in diameter); female flowers small (4 x 3 mm), yellowish Fruit: fleshy, a cluster of several fruit parts, with quite different dimensions depending on the number of fruits (up to 11.5 cm in diameter, each fruit part approx. 2.5 cm in diameter), orange-brownish; 1 seed per fruit part Seed: large (1.5 x 1 x 1 cm) Other: occasionally with stilt roots.

Regeneration It regenerates in shade (Hall & Swaine 1981).

Phenology Deciduousness: evergreen Dispersal: by animals (monkeys, elephants) Timing: not clearly seasonal (Hall & Swaine 1981); flowering period mainly from October to March (or May) (De Ruiter 1976)

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 34 30’ cells, distribution range is 1406 km Forest type: evergreen (wet, moist, upland), semideciduous (moist, dry), dry forest, secondary forest (herbarium). In Ghana, it is found mostly in moist evergreen forests (Hall & Swaine 1981).

Data sources

M

FWTA, De Ruiter (1976), Hall & Swaine (1981), De Koning (1983), De Rouw (1991), Swaine (1996), Hawthorne & Jongkind (2004)

Habitat Species occurrence increases significantly with rainfall higher than 1500 mm/yr (logistic regression

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

M.l.

79

44 (34)

8

48

1

12

37

33

19

5

63

32

25

5

H 63

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

297

22466 layout page 289-298

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Page 298

Napoleonaea heudelotii

A.Juss.

Lecythidaceae Description

Data sources

Guild: u Life form: small tree Max. height: 7 m (herbarium) Max. diameter: data unavailable Leaf: alternate, simple, elliptic, notophyll (3.5-8 x 10-17 cm), coriaceous, glabrous, with glands at the base of the laminae Inflorescence: solitary or in axillary clusters Flower: medium-sized (4 cm in diameter); purple Fruit: fleshy, ellipsoid (3.5 x 5 cm), green to orange to brown Seed: data unavailable Other: it has whorled branches.

FWTA, Liben (1971b), Hawthorne & Jongkind (2004)

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia Distribution type: continuous, regional, present in 11 30’ cells, distribution range is 538 km Forest type: high forest, secondary regrowth, gallery forest

Habitat Data unavailable.

Phenology

N

spp

298

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

N.h.

15

33 (3)

7

87

7

0

7

7

87

0

73

27

33

7

47

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 299-308

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Page 299

Neostenanthera hamata

(Benth.) Exell

Annonaceae Description Guild: sb Life form: medium-sized tree Max. height: 25 m (herbarium) Max. diameter: data unavailable Leaf: alternate, simple, elliptic, notophyll (2.5-4.5 x 6-15 cm), entire, coriaceous, veins underneath brownish, pubescent Inflorescence: axillary, on long stalks Flower: large (petals up to 7 cm long); calyx and corolla green to yellowish Fruit: fleshy, ellipsoid, pubescent, brown; 1 seed Seed: large (1.5 x 1 cm) Other: The twigs are pubescent.

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 25 30’ cells, distribution range is 1267 km, Red List species (Vulnerable) Forest type: rainforest, swamp forest (herbarium), evergreen forest (Hawthorne & Jongkind 2004)

Phenology

Habitat Species occurrence increases significantly with rainfall to reach an optimum around 2700 mm/ yr (logistic regression analysis and Chi2 test). Generally found in the rainforest understorey, but sometimes also at forest borders. Usually close to rivers (Chi2 test) and moist places (e.g. humid forests with many brooks, along rivers and even swampy forests) (herbarium).

Data sources

N

FWTA, Irvine (1961), IUCN Red List (2000), Hawthorne & Jongkind (2004)

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

H

N.h.

39

40

(20) 5

69

5

0

26

52

20

5

85

10

56

5

31

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

299

22466 layout page 299-308

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Page 300

Newtonia aubrevillei

(Pellegr.) Keay

Leguminosae-Mim. Description

regression analysis, Chi2 test). It is found in mature rainforests. Often on sandy soils (herbarium).

Guild: sb Life form: large tree Max. height: 33 m (Voorhoeve 1965) Max. diameter: 90 cm (Voorhoeve 1965) Leaf: alternate, bipinnately compound with 4-8 opposite pinnae, each pinna with 6-10 opposite leaflets, elliptic to ovate, microphyll (0.4-4 x 1.5-7 cm), entire, with or without glands between the pinnae; petiole and rachea hairy Inflorescence: axillary or terminal, unbranched (spike, erect, up to 10 cm long), above the foliage Flower: small; corolla creamy Fruit: dry dehiscent, linear-oblong (22 x 13 x 3 cm), thin-woody, opening along one edge Seed: thin, flat, very large (including wing 1.5 x 8 cm), bright red-brown, winged Other: it has narrow steep buttresses sometimes extending in spreading surface roots. The crown is fairly small and open. The slash exudes a yellow exudate.

Regeneration It has a cryptocotylar hypogeal reserve seedling type (cf. Voorhoeve 1965).

Phenology Dispersal: by wind Timing: flowering period from August to September; fruiting period from December to January

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 17 30’ cells, distribution range is 1162 km. Subspecies aubrevillei, rare in Ghana, and Upper Guinea endemic; but the species as a whole (inc. subsp. lasiantha) scattered through GuineoCongolian region (Lock 1989). Forest type: wet evergreen forest, rainforest, riverine forest, secondary forest

Habitat

N

300

n

Open (n)

The inner bark is used as an aphrodisiac (Voorhoeve 1965).

Data sources FWTA, Brenan & Brummitt (1965), Voorhoeve (1965), Lock (1989), Hawthorne & Jongkind (2004)

Species occurrence increases with rainfall to reach an optimum around 2900 mm/yr (logistic

spp

Uses

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

H

N.a.

25

7 (15)

8

40

0

0

44

16

40

4

72

24

60

4

28

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 301

Newtonia elliotii

(Harms) Keay

Leguminosae-Mim. Description

Phenology

Guild: u Life form: medium-sized tree Max. height: 20 m (herbarium) Max. diameter: data unavailable Leaf: alternate, bipinnately compound, 2 pinnae, each with 6-8 leaflets, obovate, microphyll (2-2.5 x 3-4.5 cm), with numerous looped lateral nerves Inflorescence: axillary or terminal, unbranched Flower: corolla white; fragrant Fruit: flat, somewhat curved (2 x 8 cm), smooth Seed: winged

Dispersal: by wind

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone Distribution type: continuous, local, present in 6 30’ cells, distribution range is 187 km Forest type: gallery forest

Data sources FWTA, Savill & Fox (1967), Lock (1989), Hawthorne & Jongkind (2004)

Habitat It is found along riverbanks, in swamps, and in seasonally flooded areas (herbarium).

N

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

N.e.

9

0 (2)

0

67

0

0

0

11

89

0

100

0

56

0

44

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

301

22466 layout page 299-308

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Page 302

Okoubaka aubrevillei

Pellegr. & Normand var.

aubrevillei

Santalaceae Phenology

Flower: small; greenish Fruit: fleshy (drupe), ellipsoid (5 x 9 cm); 1 seed Seed: very large Other: a tree with a cylindrical bole and without buttresses.

Dispersal: the fruits are like those of Balanites and, hence, might be dispersed by elephants, although there is no evidence for this. The saplings were found to be common in large gaps in the Krokosua hills (where elephants occur) (Hawthorne 1995a).

Distribution Continent: Cameroon (herbarium). From Côte d’Ivoire to Cameroon, with a different variety in the Democratic Republic of Congo (Keay 1989, Hawthorne 1995a) Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana (herbarium) Distribution type: continental disjunct with the largest population in Upper Guinea, present in 9 30’ cells in Upper Guinea Forest type: wet evergreen forest

Data sources FWTA, Aubréville (1959), De la Mensbruge (1966), Keay (1989), Tailfer (1989), Hawthorne (1995a), Veenendaal et al. (1996a), Hawthorne & Jongkind (2004)

Habitat Sometimes found on rocky hills. It is often found in forest openings (Veenendaal et al. 1996a). It seems to prefer N-fixing legume hosts, like Pericopsis elata (Veenendaal et al. 1996a).

Regeneration Germination is slow, and takes 3-6 months (De la Mensbruge 1966). It has a cryptocotylar hypogeal reserve seedling type (cf. de la Mensbruge 1966). Within 6 months after germination the seedling is able to infect neighbouring host seedlings. The infection leads to a decreased growth of some of the host species, but does not lead to an increased growth of Okoubaka. In the initial seedling phase, the large seed mass enables Okoubaka to realise a slow growth, independent of its hosts (Veenendaal et al. 1996a).

Description Guild: np Life form: large hemiparasitic tree Max. height: 40 m (Hawthorne 1995a) Max. diameter: 100 cm (Tailfer 1989) Leaf: opposite, simple, ovate to elliptic, mesophyll (3.5-8 x 8-15 cm), entire, herbaceous, densely pubescent beneath; 3-4 pairs of nerves Inflorescence: on older wood, branched (panicle, 15 cm long)

O

spp

302

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

O.a.

11

0 (3)

9

36

0

18

72

0

9

0

45

55

45

0

36

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 303

Oleandra ejurana

Adams

Davalliaceae Description Guild: u Life form: epiphytic herb Max. height: data unavailable Leaf: simple (1.3-2.2 x 6-25 cm), herbaceous, hairy, covered with stalked glands Spores: data unavailable Other: The main stem is densely pubescent.

Distribution Continent: Upper Guinea endemic (Hall & Swaine 1981) Upper Guinea: Ghana (herbarium, Hall & Swaine 1981) Distribution type: present in 2 30’ cells Forest type: dry forest, woodland

Habitat On trees and in shady and very moist sheltered places on steep rocks.

Phenology

Data sources FWTA, Hall & Swaine (1981)

O

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

O.e.

4

0 (3)

0

25

0

75

25

0

0

0

75

25

0

75

H 25

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

303

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Oncoba brevipes

Page 304

Stapf

Flacourtiaceae Distribution

Data sources

Continent: Upper Guinea endemic Upper Guinea: Guinea (FWTA), Sierra Leone, Liberia, Côte d’Ivoire (herbarium) Distribution type: continuous, widespread, present in 26 30’ cells, distribution range is 609 km Forest type: wet evergreen forest, rainforest, secondary forest, swamp forest

FWTA, Aubréville (1959), Sleumer (1974), Hawthorne & Jongkind (2004)

Habitat Species occurrence increases with rainfall to reach an optimum around 2700 mm/yr (logistic regression analysis, Chi2 test). It is usually found in dense forests and close to water (e.g. creeks, rivers, or near swamps). It occurs in mature forest, as well as in clearings, along roadsides, forest edges and secondary forest.

Phenology Dispersal: probably by animals

Description Guild: np Life form: medium-sized tree Max. height: 35 m (Hawthorne & Jongkind 2004) Max. diameter: 20 cm (herbarium) Leaf: alternate, simple, obovate, mesophyll (7-10 x 15-22 cm), entire, clustered at terminal axis, coriaceous, dark green above and paler beneath Inflorescence: axillary, solitary Flower: bisexual and male flowers; corolla white, conspicuously large (petals 3 x 6 cm); sweet smelling Fruit: ellipsoid (4 x 7 cm), yellow; 1 seed Seed: medium-sized (0.7 x 0.5 cm), with aril

O

spp

304

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

H

O.b.

52

59 (27)

4

44

19

2

25

22

41

2

98

0

65

2

13

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 305

Pararistolochia goldieana

(Hook.f.) Hutch. & Dalziel

Aristolochiaceae Description

Phenology

Guild: np Life form: medium-sized winding woody climber Max. height: 6 m long (Poncy 1978) Max. diameter: 3 cm (Hawthorne & Jongkind 2004) Leaf: alternate, simple, heart-shaped, mesophyll (12-15 cm long), entire, glabrous above, pubescent beneath Inflorescence: cauliflorous, solitary Flower: very large (up to 60 cm long); purplish; cylindrical; with 3 lobes; rotten scent Fruit: fleshy, green to brown Seed: large

Data sources

Distribution

FWTA, Poncy (1978), Hall & Swaine (1981), Hawthorne & Jongkind (2004)

Continent: Guinea wide: from Sierra Leone to Cameroon (Hall & Swaine 1981). Nigeria, Cameroon, Equatorial Guinea (herbarium) Upper Guinea: Sierra Leone, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 9 30’ cells, distribution range is 2932 km Forest type: In Ghana, only found in dry semideciduous forest (Hall & Swaine 1981). In Sierra Leone, Nigeria, and Cameroon, it has been recorded in rainforests and secondary forests (herbarium).

Habitat Often found in shady places in the forest understorey (herbarium).

Regeneration It regenerates in shade (Hall & Swaine 1981).

P

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

P.g.

11

29 (7)

9

36

0

45

18

0

36

0

55

45

27

9

36

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

305

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Page 306

Pararistolochia mannii

(Hook.f.) Keay

Aristolochiaceae Distribution Continent: Benin, Nigeria, Congo (Brazzaville) Upper Guinea: Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 9 30’ cells, distribution range is 2379 km Forest type: rainforest, secondary forest

Habitat Found in dense and open forests, sometimes near lagoons. On sandy soil (herbarium).

Phenology

Description Guild: u Life form: large winding woody climber Max. height: 40 m long (herbarium) Max. diameter: 3 cm (Hawthorne & Jongkind 2004) Leaf: alternate, palmately compound, 5 leaflets (12-17 cm long), entire, coriaceous, glabrous above, pubescent below Inflorescence: cauliflorous, 4-5-flowered Flower: very large (9-13 cm); outside brown to pink, inside white to yellow, with white hairs Fruit: dry indehiscent (28 cm long), brown; many seeds Seed: data unavailable

Data sources FWTA, Poncy (1978), Hawthorne & Jongkind (2004)

P

spp

306

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

P.m.

12

50 (4)

8

42

17

42

33

25

0

42

17

42

0

50

H 33

All

46333 3

7

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 307

Pavetta akeassii

J.B.Hall

Rubiaceae Description

Phenology

Guild: sb Life form: shrub Max. height: 4 m (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, oblanceolate, mesophyll (7.5-8 x 18-24.5 cm), entire, coriaceous, glabrous above, pubescent below; interpetiolar stipules Inflorescence: terminal, branched (umbel) (4-7 cm in diameter), with 15-25 flowers Flower: medium-sized; white Fruit: fleshy (drupe), globose (0.9 cm in diameter), black; 1-2 seeds Seed: medium-sized Other: it is difficult to distinguish from other Pavetta species.

Data sources

Distribution

FWTA, Hawthorne (1995a), Hawthorne & Jongkind (2004)

Continent: Upper Guinea endemic Upper Guinea: Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 8 30’ cells, distribution range is 1151 km Forest type: upland evergreen forest, wet evergreen forest, dry forest, secondary forest

Habitat Typically found in the forest understorey. On sandy or clayish soils (herbarium).

P

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

P.a.

13

0 (12)

23

54

0

8

85

0

8

0

31

69

23

0

23

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

307

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Page 308

Pavetta micheliana

J.G.Adam

Rubiaceae Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea (Adam 1973), Liberia, Côte d’Ivoire (herbarium) Distribution type: present in 3 30’ cells Forest type: evergreen forest, secondary regrowth

Habitat Sometimes close to streams (herbarium).

Phenology

Description Guild: u Life form: shrub or small tree Max. height: 9 m Max. diameter: 12 cm (herbarium) Leaf: opposite, simple, elliptic to ovate, macrophyll (9-11 x 25-27.5 cm), entire, coriaceous, glabrous; interpetiolar stipules Inflorescence: axillary, branched (panicle, 10 cm long) Flower: medium-sized (corolla tube 0.5 cm long); corolla red with pale greenish edges, tube-shaped with 4 lobes of approx. 1 cm long; fragrant Fruit: fleshy (drupe), globose (1 cm in diameter), grey; 2 seeds Seed: data unavailable

Data sources Adam (1973), Hawthorne & Jongkind (2004)

P

spp

308

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

P.m.

4

33 (3)

50

25

0

0

25

75

0

0

100

0

75

0

25

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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12:40 PM

Page 309

Penianthus patulinervis

Hutch. & Dalziel

Menispermaceae Description

Regeneration

Guild: sb Life form: shrub Max. height: 2 m (herbarium) Max. diameter: data unavailable Leaf: alternate, simple, in tufts at the end of the twigs, obovate to elliptic, mesophyll (4-13 x 15-30 cm), entire, coriaceous, glabrous; petiole 1-8.5 cm long, with a shallow groove laterally at both sides Inflorescence: dioecious, cauliflorous, unbranched (umbel, globose head with 3-7 male and female flowers) Flower: small; creamy; male flowers with 8-11 tepals; female flowers with 8 tepals Fruit: fleshy (drupe), ellipsoid (1.3 x 3 cm), orange; 1 seed Seed: very large (1 x 2.7 cm)

It has a cryptocotylar hypogeic reserve seedling type (De Koning 1983), and a high germination rate (De Koning 1983).

Phenology Timing: fruiting period the whole year round (De Koning 1983)

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 25 30’ cells, distribution range is 1185 km Forest type: rainforest, secondary forest (herbarium)

Uses The roots are chewed as an aphrodisiac (Hall & Swaine 1981).

Habitat

Data sources

Species occurrence tends to increase in lowlands (logistic regression analysis), and in places where rainfall reaches 1500-2000 mm/yr (Chi2 test). It is common in the shaded and moist understorey (herbarium, Dekker 1983, De Koning 1983). Often found close to water (e.g. rivers, streams, pools, and even swamps) (Chi2 test). Very frequently on sandy soils (herbarium, Dekker 1983), but sometimes also on clay (herbarium).

Hall & Swaine (1981), Dekker (1983), De Koning (1983)

P

spp

P.p. All

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

88

27 (59)

1

57

5

3

48

40

9

0

75

25

16

2

H 80

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

309

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Page 310

Phragmanthera vignei

Balle

Loranthaceae Description

Phenology

Guild: pi Life form: hemi-parasitic shrub Max. height: data unavailable Max. diameter: data unavailable Leaf: opposite, simple, elliptic to ovate, notophyll (3.5-8 x 4.5-10 cm), entire, coriaceous, whitish hairs beneath Inflorescence: axillary, umbel, mostly in axils of current leaves, 2-6 flowered Flower: medium-sized (corolla 3.5-4.5 cm long); corolla with brownish hairs outside, yellow inside Fruit: fleshy, oblong (0.8 x 0.6 cm), villous, reddish brown; 1 seed Seed: medium-sized Other: the internodes of branchlets are initially tomentose, but soon glabrescent.

Dispersal: by birds Timing: flowering period from December to March (Polhill & Wiens 1998)

Data sources FWTA, Balle (1956), Polhill & Wiens (1998)

Distribution Continent: Upper Guinea endemic (Polhill & Wiens 1998) Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana (herbarium, Polhill & Wiens 1998) Distribution type: continuous, widespread, present in 9 30’ cells, distribution range is 1283 km. Quite widely distributed, but still known from only very few collections, so apparently rather rare (Polhill & Wiens 1998). Forest type: wet evergreen forest

Habitat It occurs from the lowlands up to 1600 m altitude (Polhill & Wiens 1998).

P

spp

310

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

P.v.

17

0 (1)

65

65

12

0

36

59

6

6

24

71

6

0

18

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 311

Pierreodendron kerstingii

(Engl.) Little

Simaroubaceae Description

Habitat

Guild: sb Life form: medium-sized tree Max. height: 21 m (herbarium) Max. diameter: 40 cm (herbarium) Leaf: alternate, imparipinnately compound, 11-31 subopposite leaflets, oblong to elliptic, mesophyll (3-9 x 10-30 cm), entire, coriaceous, hairs on midrib Inflorescence: several clustered spikes Flower: small; red Fruit: oblong to ellipsoid (4.5 x 7 cm), woody, yellow Seed: data unavailable Other: it has a rough bark in square scales, with a strong almond scent.

Sometimes found on riverbanks.

Phenology Data unavailable.

Data sources FWTA, Aubréville (1959), Keay (1989), Hawthorne (1995a), IUCN Red List (2000), Hawthorne & Jongkind (2004)

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Côte d’Ivoire, Ghana (herbarium), Togo (Hawthorne 1995a) Distribution type: Upper Guinea disjunct, widespread, present in 6 30’ cells, Red List species (Vulnerable) Forest type: rainforest, wet evergreen forest, semideciduous forest Other: a genus of two (or one very variable) species (Keay 1989): P. africanum occurs from Nigeria to Angola; P. kerstingii is known in Sierra Leone, Côte d’Ivoire, Ghana (where it is uncommon) and Togo. Although the trees in the two extremes are remarkably similar, the discontinuous distribution in Ghana between specimens in very dry forest (e.g. Awura F.R.) and in very wet forest (e.g. Ankasa GPR), with few cases in between, does suggest that two species may be involved (Hawthorne 1995a).

P

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

P.k.

8

0 (1)

0

50

38

13

38

38

13

50

13

38

0

13

38

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

311

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Page 312

Piptostigma fugax

A.Chev. ex Hutch. & Dalziel

Annonaceae Habitat It is found in the understorey of both dense and disturbed forests. Occasionally reported on sandy soils (herbarium).

Regeneration It regenerates in shade (Hall & Swaine 1981).

Phenology Dispersal: probably dispersed by animals

Description

Data sources

Distribution

Guild: sb Life form: shrub or small tree Max. height: 5 m (herbarium) Max. diameter: data unavailable Leaf: alternate, simple, elliptic, mesophyll (4-7 x 9-21 cm), entire, herbaceous, hairy Inflorescence: cauliflorous Flower: large; calyx with brown hairs; corolla cream, pinkish, or orange, bowl-shaped Fruit: fleshy (1.8 cm long), yellow Seed: large (1.3 x 1 x 0.6 cm) Other: the young twigs are hairy.

FWTA, Hall & Swaine (1981), IUCN Red List (2000) Continent: Upper Guinea endemic Upper Guinea: Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 12 30’ cells, distribution range is 1133 km, Red List species (Vulnerable) Forest type: evergreen forest, semi-deciduous forest, disturbed forest. In Ghana, it is mostly found in upland evergreen forests, but it is also present in wet evergreen forests, and moist semi-deciduous forests (Hall & Swaine 1981).

P

spp

312

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

P.f.

16

23 (13)

13

44

0

19

57

19

6

0

81

19

56

0

H 31

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 313

Platysepalum hirsutum

(Dunn) Hepper

Leguminosae-Pap. Description Guild: np Life form: medium-sized woody climber Max. height: 8 m (herbarium) Max. diameter: 4 cm (herbarium) Leaf: imparipinnately compound, 7-9 opposite leaflets, obovate to oblong, notophyll (2-5 x 4-11 cm), herbaceous, brownish hairs on the midribs, soft, dense yellow-orange spreading hairs falling with age to leave short whitish hairs below the leaf Inflorescence: terminal, branched (panicle, up to 35 cm long) Flower: calyx brown, hairy; corolla white with deep pink slashes Fruit: dry dehiscent, flat (2.5 x 7 cm), deep brown, woody, densely covered by stiff hairs Seed: disk-shaped, large (1.2 x 0.9 x 0.1 cm) Other: the slash exudes a brownish gum. The stem of medium-sized lianas develops two lobes.

Regeneration It regenerates in shade (Hall & Swaine 1981).

Phenology Dispersal: explosively dehiscent

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 38 30’ cells, distribution range is 1418 km Forest type: wet evergreen forest, moist evergreen forest, dry semi-deciduous forest, secondary forest. In Ghana, it occurs predominantly in drier forest types (Hall & Swaine 1981).

Data sources

Habitat

FWTA, Hall & Swaine (1981), Lock (1989), Hawthorne & Jongkind (2004)

We did not find any significant correlation between the species occurrence and the environmental variables tested. It is found in a wide variety of forest types. Often found under the forest canopy but also in disturbed forest understorey, at forest edges, and occasionally along roads. Sometimes near streams (herbarium).

P

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

P.h.

65

17 (29)

5

35

6

38

32

13

19

2

55

43

38

6

51

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

313

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Page 314

Pleiocoryne fernandensis

(Hiern) S.Rauschert

Rubiaceae Description

Habitat

Guild: np Life form: large woody climber Max. height: 15 m long (De Koning 1983) Max. diameter: data unavailable Leaf: opposite, simple, ovate, notophyll (3-6 x 5-10 cm), entire, coriaceous to fleshy, glabrous; long petiole (> 2 cm); interpetiolar stipules Inflorescence: terminal or axillary, unbranched (cyme, 5 cm in diameter) Flower: medium-sized (corolla tube 2-3.5 cm long); corolla green and white, long tube-shaped with 5 lobes; very sweetly scented Fruit: fleshy, globose (4 x 5.5 cm), orange; many seeds Seed: flat, medium-sized (approx. 1 cm across) Other: it probably starts as a shrub when young. The branchlets are angular and glabrous.

Often found in open and very wet places (e.g. along creeks, riverbanks, marshy places, and swamps). It can also be found on sandy beaches at the edge of open coastal rainforest. Apparently, more commonly reported in disturbed than in mature lowland rainforest. Often on white sandy soils (herbarium).

Regeneration It has a phanerocotylar epigeal foliaceous seedling type (cf. De Koning 1983).

Phenology

Distribution Continent: Nigeria, Cameroon, Equatorial Guinea, Fernando Po, Gabon, Congo, Democratic Republic of Congo (Kirkbride & Robbrecht 1984), Angola (herbarium) Upper Guinea: Guinea (Kirkbride & Robbrecht 1984), Sierra Leone, Liberia, Côte d’Ivoire, Ghana (herbarium), Togo (Kirkbride & Robbrecht 1984) Distribution type: continuous, widespread, present in 20 30’ cells, distribution range is 4265 km Forest type: wet evergreen forest, moist evergreen forest, coastal forest, dry forest, swamp forest, secondary forest, thickets

Data sources De Koning (1983), Kirkbride & Robbrecht (1984), Hawthorne & Jongkind (2004)

P

spp

314

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

P.f.

26

45 (11)

4

35

15

19

30

31

20

31

31

38

19

23

27

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 315

Poecilocalyx stipulosa

(Hutch. & Dalziel) N. Hallé

Rubiaceae Description

Phenology

Guild: sb Life form: shrub Max. height: 2 m (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, narrowly oblong to elliptic, mesophyll (5-6 x 15-18 cm), herbaceous; interpetiolar stipules, leafy, >1cm long; midrib with a crest of orange hairs, pilose beneath Inflorescence: axillary, flowers in a globose head Flower: corolla white Fruit: fleshy; many seeds Seed: data unavailable Other: the young twigs are softly hairy.

Data sources

Distribution

FWTA, De Rouw (1991), Hawthorne & Jongkind (2004)

Continent: Upper Guinea endemic Upper Guinea: Liberia, Côte d’Ivoire Distribution type: continuous, regional, present in 4 30’ cells, distribution range is 358 km Forest type: wet evergreen forest

Habitat It is found in the forest understorey (herbarium, De Rouw 1991).

P

spp

n

Open (n)

P.s.

6

0 (1)

All

46333

37

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

L

M

H

L

M

H

0

67

0

0

0

33

66

0

83

17

83

0

0

39

7

37

29

24

10

6

69

25

36

13

39

4

Soil CMK

Soil WHC

315

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Page 316

Polycephalium capitatum

(Baill.) Keay

Icacinaceae Description

Phenology

Guild: pi Life form: small winding woody climber Max. height: 3 m long Max. diameter: data unavailable Leaf: alternate, simple, elliptic, mesophyll (5-12 x 9-21 cm), entire, herbaceous, rusty-brown hairs on the lower side and on the veins on both sides; hairy petiole Inflorescence: compound, many small flowers in a cup surrounded by leaf-like bracts Flower: small (0.4-0.6 cm in diameter); creamy Fruit: fleshy (1.1 x 3.3 cm), orange, pubescent; 1 seed Seed: data unavailable

Distribution

FWTA

Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire Distribution type: continuous, widespread, present in 18 30’ cells, distribution range is 1067 km Forest type: secondary forest, thicket

Habitat Species occurrence is highest in zones where rainfall is around 2700 mm/yr (logistic regression analysis, Chi2 test). It is only found in secondary vegetation such as degraded forests, and roadside thickets (herbarium).

P

spp

316

n

Open (n)

Data sources

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

H

P.c.

25

67 (9)

8

48

8

0

36

20

44

8

80

12

44

0

36

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 317

Polystachya bancoensis

Burg

Orchidaceae Description

Phenology

Guild: pi Life form: epiphytic herb Max. height: 8.5 cm long Leaf: alternate, simple, elliptic, microphyll (0.7-1.5 x 2-6 cm), coriaceous, greyish green to reddish blue; 1-2 leaves Inflorescence: terminal, not branched (up to 7 cm long), pendulous, up to 13 flowers Flower: small (petals 0.5 cm long), pubescent; calyx green to yellowish; corolla greenish yellow; fragrant Fruit: dry dehiscent (capsule), elliptic, green-brown; many seeds Seed: very small Other: the pseudobulb is elliptic, dull bluish green sometimes tinged purplish.

Deciduousness: deciduous, it drops the leaves during the dry season (Arends et al. 1980) Dispersal: by wind Timing: flowering period from October to November (De Koning 1983)

Data sources FWTA, Arends et al. (1980), De Koning (1983)

Distribution Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 6 30’ cells, distribution range is 657 km Forest type: evergreen forest (Arends et al. 1980)

Habitat It is found on the bare bark of fully exposed branches of large forest trees. Usually in forests at lower altitudes (Arends et al. 1980).

P

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

P.b.

8

0 (4)

0

63

0

0

38

63

0

25

63

13

25

25

50

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

317

22466 layout page 309-318

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Page 318

Polystachya dalzielii

Summerh.

Orchidaceae Description

Phenology

Guild: u Life form: small epiphytic herb Max. height: 7 cm Leaf: alternate, simple, lanceolate, microphyll (0.22 x 6-12 cm), entire, thin, purplish green Inflorescence: terminal, unbranched (raceme) (up to 15 cm long), rising from a leafless pseudobulb Flower: small; mauve to pink with yellow on lip margin, glabrous Fruit: dry dehiscent (capsule), elliptic, green-brown; many seeds Seed: very small Other: the pseudobulb is around 1 cm, conical, very hard, green.

Deciduousness: deciduous Dispersal: by wind Timing: flowering period from February to March (Johansson 1974)

Data sources FWTA, Jaeger et al. (1968), Johansson (1974)

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire Distribution type: continuous, widespread, present in 9 30’ cells, distribution range is 756 km Forest type: montane forest

Habitat It grows on the middle or outer parts of large tree branches, in full sun and on minor humus deposits. Usually at altitudes over 1000 m (Johansson 1974).

P

spp

318

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

P.d.

17

25 (4)

88

18

0

6

24

53

18

0

71

29

41

0

H 29

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 319

Polystachya pseudodisa

Kraenzl.

Orchidaceae Description Guild: u Life form: epiphytic herb Max. height: data unavailable Leaf: alternate, simple, narrowly strap-shaped, microphyll (up to 1 x 10 cm), entire Inflorescence: terminal, unbranched (spike, 30 cm or more) Flower: small, pink with yellow centre Fruit: dry dehiscent (capsule), elliptic, green-brown Seed: very small Other: it has a pseudobulb.

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire Distribution type: continuous, regional, present in 4 30’ cells, distribution range is 581 km Forest type: data unavailable

Habitat It grows on trees or rocky slopes (herbarium).

Phenology Dispersal: by wind

Data sources FWTA

P

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

P.p.

8

- (0)

38

63

0

0

25

0

76

0

50

50

13

0

H 38

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

319

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Page 320

Premna grandifolia

A.Meeuse

Verbenaceae Description

Data sources

Guild: u Life form: shrub Max. height: 1 m (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, elongate to oblanceolate, mesophyll (6-9 x 25-30 cm), finely dentate in the upper half, pilose on the midrib beneath Inflorescence: a small subsessile cyme Flower: small; white Fruit: unknown Seed: unknown

FWTA, IUCN Red List (2000), Hawthorne & Jongkind (2004)

Distribution Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire Distribution type: present in 2 30’ cells. It is only known from southwest Côte d’Ivoire. A Red list species (Vulnerable). Forest type: data unavailable

Habitat Data unavailable.

Phenology

P

spp

320

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

P.g.

2

- (0)

0

50

0

0

50

50

0

0

50

50

50

0

H 0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

22466 layout page 319-328

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Page 321

Pseudocalyx libericus

Breteler

Acanthaceae Description

Phenology

Guild: u Life form: winding woody climber Max. height: data unavailable Max. diameter: data unavailable Leaf: opposite, simple, elliptic to obovate, mesophyll (3-8 x 11-20 cm), entire, coriaceous to herbaceous, glabrous to sparsely hairy on venation Inflorescence: terminal or axillary, not branched, erect Flower: medium-sized (3.2 cm long), yellowish, laterally flattened Fruit: dehiscent, subovoid (2.2 x 1.5 cm), hairy, green Seed: data unavailable Other: it has grey-green stems.

Data sources

Distribution

Breteler (1994a, 1998), Hawthorne & Jongkind (2004)

Continent: Upper Guinea endemic Upper Guinea: Liberia (Breteler 1994a, herbarium) Distribution type: continuous, very local, present in 1 30’ cell Forest type: wet evergreen forest (herbarium)

Habitat Found along forest roads or forest edges. It occurs on sandy laterite soils (herbarium).

P

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

P.l.

2

67 (3)

0

50

0

0

0

0

100

0

100

0

100

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

321

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Page 322

Pteleopsis habeensis

Aubrév. ex Keay

Combretaceae

Description Guild: pi Life form: small to medium-sized tree Max. height: 15 m (herbarium) Max. diameter: 40 cm (herbarium) Leaf: alternate, simple, ovate with a blunt apex, microphyll (1-2 x 1.5-5 cm), entire, herbaceous, hairy midrib Inflorescence: terminal, not branched Flower: small; white; fragrant Fruit: dry indehiscent, pendulous (0.7 x 1 cm), yellow-brown, 3-winged Seed: medium-sized (0.3 x 0.2 cm) Other: a twisted tree with flaking bark.

Habitat

Uses

In the Yankari Game Reserve, in Nigeria, it occurs in more or less dense stands on slopes along watercourses. These slopes are very dry, sandy, and well-drained. The species is never found in or near the actual streambed. It is apparently fire sensitive and occurs in either places where it is protected from fire, such as on rocky hills, or where it shades out the fire-providing heliophilous grasses (Geerling 1984). In the Yankari Game Reserve, it covers several square kilometres and forms pure stands of coppice-like woodland. This is truly remarkable in a savanna where only exceptionally a single species dominates (Geerling 1984). The two sites in Ghana where this species was collected (see map) are both close to a river (Volta and Black Volta) but also above the wet season’s high water level.

It is relatively unpalatable to herbivores, and being deciduous, does not provide shade in the dry season (Geerling 1984).

Data sources FWTA, Geerling (1984), Hawthorne (1995a), Hawthorne & Jongkind (2004)

Phenology Deciduousness: deciduous (Geerling 1984) Dispersal: by wind

Distribution Continent: Mali, Nigeria Upper Guinea: Ghana Distribution type: continuous, widespread, in Upper Guinea present in 2 30’ cells. This species is known only from four localities in the world: the Bandiagara scarps in Mali; the Yankari Game Reserve in Nigeria; and the dry forests at Bui and Akosombo in Ghana. Forest type: dry forest, savanna woodland, savanna

P

spp

P.h. All

322

n

6 46333

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

40 (5)

0

17

0

100

0

0

0

0

0

100

0

0

0

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 323

Ptychopetalum anceps

Oliv.

Olacaceae Description Guild: sb Life form: shrub or small tree Max height: 9 m (herbarium) Max diameter: data unavailable Leaf: alternate, simple, oblong to elliptic, notophyll (2.5-4 x 6.5-14 cm), herbaceous; subsessile Inflorescence: axillary, not branched Flower: small to medium-sized; funnel-shaped; creamy white Fruit: fleshy (drupe), irregularly globose (0.75 x 1.8 cm), red, velvety; 1 seed Seed: large (1.5 x 0.6 x 0.6 cm), grooved Other: the slash exudes an orange exudate.

Distribution

Phenology

Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 39 30’ cells, distribution range is 1055 km Forest type: evergreen (wet, moist, upland), moist semi-deciduous forest (herbarium, Hall & Swaine 1981). In Ghana, most often found in wet and upland evergreen forests (Hall & Swaine 1981).

Dispersal: probably by birds (De Koning 1983) Timing: flowering period from March to April; fruiting period from November to August (De Koning 1983)

Habitat Species occurrence increases with rainfall to reach a very wide optimum range between 1500-3500 mm/yr (logistic regression analysis, Chi2 test). Most often in lowlands (logistic regression analysis, Chi2 test), although, in Ghana, it is very frequently recorded in upland forests (Hall & Swaine 1981). It can be found in a wide variety of evergreen and semi-deciduous forests. Usually under shade, and very often along creeks or rivers. Soils can be sandy, loamy, clayish, or lateritic (herbarium). In Ghana, plot records are very significantly associated

with acidic, base-poor soils (Hall & Swaine 1981). In Côte d’Ivoire, it is typical for coastal wet evergreen forests (Kasparek 2000).

Regeneration

Data sources Michaud (1966), Hall & Swaine (1981), De Koning (1983), Kasparek (2000), Hawthorne & Jongkind (2004)

It regenerates in shade (Hall & Swaine 1981).

P

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

H

P.a.

119

33 (91)

0

40

6

0

52

24

23

4

77

18

45

6

45

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

323

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Page 324

Puelia olyriformis

(Franch.) Clayton

Gramineae Inflorescence: terminal and drooping, branched (panicle, up to 15 cm long) Flower: pale green Fruit: data unavailable Seed: data unavailable Other: the stem is slightly purplish. The leaves are only found on the top of the plant.

Distribution Continent: Cameroon (Van der Zon 1992), Gabon (herbarium), Tanzania (Van der Zon 1992) Upper Guinea: Senegal, Guinea, Sierra Leone, Liberia (herbarium) Distribution type: continental disjunct, in Upper Guinea present in 4 30’ cells Forest type: rainforest, secondary forest

Habitat It is often found in deep shade in mature forest. It can be found along riverbanks or in hills and mountains (herbarium).

Phenology

Description Guild: sb Life form: erect herb Max. height: 1.8 m (herbarium) Max. diameter: data unavailable Leaf: simple, linear, microphyll (2-7 x 15-35 cm), entire, herbaceous, smooth, glossy dark green

Data sources FWTA, Van der Zon (1992)

P

spp

324

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

P.o.

5

67 (3)

0

20

60

20

0

20

60

0

0

100

0

0

60

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 325

Pycnanthus dinklagei

Warb.

Myristicaceae Description Guild: np Life form: large woody climber Max height: 25 m long (herbarium) Max diameter: data unavailable Leaf: alternate, simple, elliptic, mesophyll (4-6 x 10-17 cm), margin upturned, entire to dentate towards the apex, coriaceous, glabrous; petiole approx. 1 mm long Inflorescence: axillary, branched (panicle, 6-8 cm long) Flower: small; corolla yellow; fragrant Fruit: dry dehiscent, 2-valved, ellipsoid (1.8 x 3 cm), orange; 1 seed Seed: large (1.2 x 1.6 cm), brown with a red aril Other: a climber that probably starts as a straggling shrub. The slash is pale brown, soon turning reddish, it exudes a red latex. The bark has lenticels.

Phenology

Data sources FWTA, De Koning (1983)

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 18 30’ cells, distribution range is 1017 km Forest type: wet evergreen forest, rainforest, secondary forest, coastal shrubland.

Habitat Species occurrence increases significantly in lowlands (logistic regression analysis) and in places with rainfall higher than 2000 mm/yr (logistic regression analysis, Chi2 test). Usually, found close to rivers and streams (Chi2 test, De Koning 1983). Generally, growing on forest trees in shady and wet places (De Koning 1983). On sandy, sandy-clayish, or alluvial soils (herbarium).

P

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

H

P.d.

39

29 (28)

0

64

13

0

33

36

21

3

77

21

23

3

64

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

325

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Page 326

Pyrenacantha glabrescens

(Engl.) Engl.

Icacinaceae Description

Phenology

Guild: np Life form: large winding woody climber Max. height: 10 m long (herbarium) Max. diameter: 10 cm (herbarium) Leaf: alternate, simple, obovate to elliptic, mesophyll (2.5-9 x 6-20 cm), entire, coriaceous, pubescent; petiole (1-3 cm long), hairy Inflorescence: axillary, unbranched (6-15 cm long), dioecious, rachis hairy Flower: small (0.15 x 0.25 cm); pubescent outside; no calyx; corolla pale yellow, 4 free petals Fruit: fleshy, ellipsoid (1.8 x 3.3 cm), orange, pubescent; 1 seed Seed: large (2.0 x 1.2 x 0.8 cm), irregularly grooved and ridged

Timing: flowering period from March to June (De Koning 1983); fruiting period from December to April (De Koning 1983)

Distribution

Data sources

Continent: Cameroon Upper Guinea: Liberia, Côte d’Ivoire, Ghana Distribution type: continental disjunct with the largest population in Upper Guinea, present in 9 30’ cells in Upper Guinea Forest type: wet evergreen forest, secondary forest

FWTA, De Koning (1983)

Habitat Most commonly found in established secondary forest, and along forest borders (Kuzee pers. comm., herbarium).

P

spp

326

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

L

M

H

0

65

35

29

0

71

69

25

36

13

39

P.g.

17

21 (14)

0

53

0

0

65

35

0

All

46333

37

4

39

7

37

29

24

10

6

Soil WHC

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Page 327

Raphia palma-pinus

(Gaertn.) Hutch.

Palmae Description

Phenology

Guild: u Life form: palm tree Max. height: 9 m (herbarium) Max. diameter: data unavailable Leaf: in a terminal cluster, pinnately compound (up to 10 m long), leaflets linear, spinose Inflorescence: branched (lax panicle, sticking out at 4 m height) Flower: data unavailable Fruit: fleshy, elongate-ovoid (4 cm x 8.8 cm), orange-brown, with scales Seed: very large (4.5 x 2.5 x 2.5 cm)

Dispersal: by birds

Distribution Data sources

Continent: Upper Guinea endemic Upper Guinea: Senegal, Gambia, Guinea, Sierra Leone, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 12 30’ cells, distribution range is 1859 km Forest type: wet evergreen forest, swamps (herbarium, Hawthorne 1995a)

FWTA, Russell (1965), Hawthorne (1995a)

Habitat It seems to be confined to permanent swamps in the wet evergreen forest area or behind the mangrove zone.

R

spp

n

Open (n)

R.p.

16

67 (3)

All

46333

37

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

L

M

H

L

M

H

56

0

37

44

6

12

0

44

56

19

31

44

39

7

37

29

24

10

6

69

25

36

13

39

0 4

Soil CMK

Soil WHC

327

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Page 328

Renealmia battenbergiana

Cummins ex Baker

Zingiberaceae Description

Phenology

Guild: sb Life form: large herb Max. height: 1 m (herbarium) Max. diameter: data unavailable Leaf: simple, elliptic, mesophyll (5-5.5 x 20 cm), entire, coriaceous, reddish beneath Inflorescence: at the top of leafy shoots, branched (paniculate) Flower: data unavailable Fruit: fleshy, oblong (2 cm long), red Seed: shiny brown Other: sterile specimens are easily confounded with R. africana.

Data sources

Distribution

FWTA, Hawthorne (1995a) Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire, Ghana Distribution type: present in 4 30’ cells Forest type: wet evergreen forest

Habitat Found in moist places of dense forests (e.g. by streams). On sandy soils (herbarium).

Regeneration It tends to form clumps.

R

spp

328

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

R.b.

4

0 (3)

25

50

0

0

75

25

0

0

25

75

0

0

H 75

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 329

Renealmia longifolia

K.Schum.

Zingiberaceae Description Guild: u Life form: large herb Max. height: 1.2 m (herbarium) Max. diameter: data unavailable Leaf: simple, elongate to lanceolate, mesophyll (4-5 cm x 35 cm), entire, coriaceous Inflorescence: at the top of leafy shoots, branched (paniculate) Flower: pink Fruit: fleshy, ellipsoid (1.2 cm long), black Seed: medium-sized (0.9 x 0.6 cm), reddish brown

Distribution Continent: Upper Guinea endemic Upper Guinea: Liberia, Côte d’Ivoire (FWTA) Distribution type: local, present in 3 30’ cells, distribution range is 230 km Forest type: data unavailable

Habitat There is little data available on this species, but it seems to prefer wetter areas (near rivers and swamps) (herbarium).

Phenology

Data sources FWTA

R

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

R.l.

3

100 (1)

0

33

0

0

33

33

33

0

100

0

100

0

H 0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

329

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Page 330

Rhaptopetalum beguei

Mangenot

Scytopetalaceae

Habitat Found in wet areas (e.g. around lagoons, near rivers, and especially in swamps) (herbarium, Hawthorne & Jongkind 2004).

Phenology Dispersal: by animals (chimpanzees)

Description Guild: np Life form: small tree Max. height: 10 m (herbarium) Max. diameter: 20 cm (herbarium) Leaf: alternate, simple, lanceolate to ovate, mesophyll (6-8 x 13-20 cm), entire, coriaceous, glabrous; 5 pairs of lateral nerves Inflorescence: cauliflorous, solitary flowers in clusters together Flower: small, corolla pink, 3-4 lobes Fruit: fleshy (drupe), partially dehiscent, subglobose (1.8 cm in diameter), 4-5 valves, orange-red outside, white sweet pulp inside, persistent green calyx; 1 seed Seed: medium-sized (1 x 0.5 x 0.5 cm), yellow

Distribution Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire, Ghana Distribution type: continuous, regional, present in 10 30’ cells, distribution range is 562 km Forest type: rainforest, evergreen forest, deciduous forest, swamp forest, secondary forest

Data sources FWTA, Mangenot (1957), Letouzey (1961), Hawthorne & Jongkind (2004)

R

spp

330

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

R.b.

37

21 (9)

0

70

0

0

46

54

0

16

57

27

8

11

H 62

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 331

Rinorea aylmeri

Chipp

Violaceae Description

Phenology

Guild: u Life form: shrub or small tree Max. height: 5 m Max. diameter: data unavailable Leaf: alternate, simple, elliptic, mesophyll (4-7 x 1220 cm), serrate, coriaceous, glabrous; petiole up to 5 cm long Inflorescence: terminal or axillary; unbranched (3-6 cm long) Flower: small; corolla white Fruit: dry dehiscent, triangular to globose (2 cm in diameter), dark brown, sparsely puberulous to glabrous Seed: data unavailable Other: the young stems are very dark brown. When slightly older (i.e. in the region where mature leaves are borne) the dark coloured epidermis abscises and is replaced by a whitish or very pale green to fawn coloured bark.

Timing: flowering period in April

Data sources Hawthorne & Jongkind (2004)

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea-Bissau, Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 23 30’ cells, distribution range is 1670 km Forest type: wet evergreen forest

Habitat Species occurrence increases significantly with rainfall to reach an optimum around 2800 mm/yr (logistic regression analysis, Chi2 test). It is found usually near water by streams and riverbanks. Very often under forest shade but occasionally also in open places (herbarium).

R

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

R.a.

36

22 (9)

3

42

14

3

37

17

43

0

83

17

53

8

H 31

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

331

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Page 332

Sabicea arachnoidea

Hutch. & Dalziel

Rubiaceae Description

Data sources

Guild: u Life form: small winding woody climber Max. height: data unavailable Max. diameter: data unavailable Leaf: opposite, simple, obovate, notophyll (4-5 x 10-12 cm), whitish beneath, pubescent above; interpetiolar stipules Inflorescence: axillary, flowers in a dense spherical head Flower: corolla tube-shaped, yellow Fruit: fleshy, subglobose or ellipsoid (2 cm or less in diameter), soft and sweet; several seeds Seed: data unavailable Other: a rather slender, pubescent climber.

FWTA, Hawthorne & Jongkind (2004)

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone Distribution type: continuous, very local, present in 1 30’ cell, distribution range is 22.7 km Forest type: data unavailable.

Habitat Data unavailable.

Phenology Dispersal: probably by animals

S

spp

332

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

S.a.

3

- (0)

0

33

0

0

0

0

100

0

100

0

67

0

H 33

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 333

Sabicea bracteolata

Wernham

Rubiaceae Description

Data sources

Guild: u Life form: small winding woody climber Max. height: data unavailable Max. diameter: data unavailable Leaf: opposite, simple, elliptic, notophyll (2.5-5 x 4-10 cm), entire, pubescent on the nerves below; interpetiolar stipules Inflorescence: axillary, solitary Flower: medium-sized; densely pubescent outside Fruit: fleshy, subglobose or ellipsoid (2 cm or less in diameter), soft and sweet, red; several seeds Seed: data unavailable Other: it probably starts as a shrub. The stems are densely pubescent when young.

FWTA, Hawthorne & Jongkind (2004)

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea Distribution type: continuous, very local, present in 1 30’ cell, only known from 3 collections in Guinea Forest type: data unavailable

Habitat Data unavailable.

Phenology Dispersal: probably by animals

S

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

S.b.

1

- (0)

0

100

0

0

100

0

0

0

0

100

0

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

333

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Sabicea discolor

Page 334

Stapf

Rubiaceae Description

Phenology

Guild: pi Life form: small winding climber Max. height: 5 m long (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, elliptic to oblong, notophyll (3-4.5 x 6-11.5 cm), entire, leaves covered with stiff apressed hairs on both sides, reflexed interpetiolar stipules Inflorescence: axillary, branched (cyme, 2-6 cm long) Flower: medium-sized; star-like; corolla tube red, lobes white; hairy on the throat Fruit: fleshy, silvery-greyish outside with reddish pulp, some with pale hairs, slightly bitter tasting; many seeds Seed: data unavailable

Dispersal: probably by animals

Distribution

FWTA, Hawthorne & Jongkind (2004)

Continent: Upper Guinea endemic Upper Guinea: Guinea-Bissau, Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 39 30’ cells, distribution range is 1776 km Forest type: montane rainforest, wet evergreen forest, moist evergreen forest, swamp forest, secondary forest, thickets

Habitat Species occurrence increases significantly with rainfall, reaching an optimum around 2700 mm/yr (logistic regression analysis, Chi2 test). It is frequently reported in disturbed places (e.g. roadsides, logged forests, and farm bushes) and in very wet conditions (e.g. near rivers, and swampy areas). Soils can be sandy, clayish, loamy, or lateritic (herbarium).

S

spp

334

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

H

S.d.

61

82 (34)

10

33

18

5

33

28

31

2

75

23

49

0

36

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

Data sources

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Page 335

Sabicea harleyae

Hepper

Rubiaceae Description

Phenology

Guild: pi Life form: small winding woody climber Max. height: data unavailable Max. diameter: data unavailable Leaf: opposite, simple, ovate, notophyll (3-6 x 5-10.5 cm), entire, lower leaf surface densely hairy; interpetiolar stipules, erect Inflorescence: axillary, globose (3 cm in diameter), with a long peduncle Flower: medium-sized; calyx and bracts pink to violet; corolla tube violet outside, white inside Fruit: fleshy, globose (0.8 cm in diameter), red; sweet-sour Seed: data unavailable Other: a slender climber with hairy stems.

Dispersal: probably by animals

Data sources

Distribution

FWTA, Hepper (1958), Hawthorne & Jongkind (2004)

Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire Distribution type: continuous, regional, present in 8 30’ cells, distribution range is 401 km Forest type: upland evergreen forest, wet evergreen forest, secondary forest

Habitat Practically only reported in disturbed open places (e.g. roadsides, logged and open forests) (herbarium).

S

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

S.h.

14

73 (11)

29

36

0

0

7

58

36

0

100

0

79

0

21

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

335

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Page 336

Salacia columna

N.Hallé

Celastraceae Habitat Species occurrence increases very significantly with rainfall (logistic regression analysis), and is highest at a rainfall over 2500 mm/yr. It can be found in both relatively undisturbed forests and secondary vegetation (herbarium).

Regeneration Description

It regenerates in shade (Hall & Swaine 1981).

Guild: sb Life form: medium-sized irregularly winding woody climber Max. height: 10 m long (Hallé 1962) Max. diameter: data unavailable Leaf: opposite, simple, elliptic, mesophyll (6 x 15 cm), entire, coriaceous, glabrous Inflorescence: axillary Flower: small; whitish, dish-shaped Fruit: fleshy, triangular (7.5 x 4 cm), orange; 4-8 seeds in a slimy, whitish, sweet edible pulp Seed: very large (3 x 4.5 cm), brown

Phenology Dispersal: probably by animals

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 18 30’ cells, distribution range is 1255 km Forest type: upland evergreen forest, wet evergreen forest, moist evergreen forest, secondary forest. In Ghana, mostly found in wet evergreen forest (Hall & Swaine 1981). In Liberia, often found in dry secondary vegetation and bushes (herbarium).

S

spp

336

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

H

S.c

23

42 (12)

4

39

9

0

31

9

51

0

91

9

78

0

13

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

Uses The fruits are edible (herbarium).

Data sources Hallé (1962), Hall & Swaine (1981)

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Page 337

Salacia howesii

Hutch. & M.B.Moss

Celastraceae Description

Habitat

Guild: np Life form: large irregularly winding woody climber Max. height: 20 m long (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, mesophyll (5 x 14 cm), entire, coriaceous, densely pubescent underneath Inflorescence: axillary, branched Flower: small (0.6 cm in diameter); yellowish; dishshaped Fruit: fleshy, globose (approx. 2.5 cm in diameter), orange Seed: data unavailable Other: the twigs are densely pubescent and not lenticellate. It has no exudate. The wood on crosssection is pale reddish with many darker dots.

Sometimes observed near the coast.

Regeneration It regenerates in shade (Hall & Swaine 1981).

Phenology Data unavailable.

Data sources FWTA, Hallé (1962), Hall & Lock (1975), Hall & Swaine (1981)

Distribution Continent: Nigeria, Gabon Upper Guinea: Côte d’Ivoire, Ghana Distribution type: continuous, widespread, in Upper Guinea present in 4 30’ cells, distribution range is 1985 km Forest type: wet evergreen forest, moist evergreen forest, moist semi-deciduous forest

S

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

S.h.

4

25 (4)

0

0

50

25

75

0

0

0

50

50

0

0

100

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

337

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Page 338

Sapium aubrevillei

Leandri

Euphorbiaceae Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 10 30’ cells, distribution range is 1207 km, Red List species (Vulnerable). It is rare (Hawthorne 1995a). Forest type: wet evergreen forest, gallery forest

Habitat Few records found in both dense and open forests. We did not find any significant relationship with environmental variables.

Regeneration It has a phanerocotylar epigeal foliaceous seedling type (De la Mensbruge 1966).

Description

Phenology

Guild: pi Life form: large tree Max. height: 36 m (herbarium) Max. diameter: 120 cm (FWTA) Leaf: alternate, simple, oblong to elliptic, mesophyll (5-7 x 10-16 cm), serrate, two glands on margin at base, fairly coriaceous Inflorescence: axillary, unbranched (spike), male flowers below and female flowers above Flower: small Fruit: fleshy (drupe), globose (1.8 cm in diameter), yellow; 2 seeds Seed: data unavailable Other: a tree with swollen base and root ridges. The bark is flaking in long fibrous strips. Branches with white latex. The species is similar to S. ellipticum, but occurs in evergreen forest.

Deciduousness: deciduous Dispersal: probably by animals

Data sources FWTA, Aubréville (1959), De la Mensbruge (1966), Savill & Fox (1967), Hawthorne (1990, 1995a), IUCN Red List (2000)

S

spp

338

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

S.a.

11

0 (7)

18

64

0

9

45

36

9

0

64

36

18

9

45

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 339

Scadoxus multiflorus (Martyn) Raf. ssp. longitubus (C.H.Wright) Friis & Nordal Amaryllidaceae Description Guild: np Life form: perennial bulbous ground herb Max. height: 0.5 m (herbarium) Leaf: opposite, simple, elliptic to obovate, mesophyll (7-8 x 25-30 cm), entire, herbaceous, glabrous Inflorescence: branched (in a globose head) Flower: large; long, slender, bright red Fruit: fleshy (berry) (up to 1.5 cm in diameter), orange to red; 1-3 seeds Seed: medium-sized, pale yellow Other: it has a maroon-brown bulb. The exposed part is greenish white with maroon-brown dots.

Distribution Phenology

Continent: the species has a Guineo-Congolian range, but the subspecies is restricted to Upper Guinea (Hall & Swaine 1981) Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana (herbarium) Distribution type: continuous, widespread, present in 34 30’ cells, distribution range is 1350 km Forest type: wet evergreen forest, moist evergreen forest, secondary forest, gallery and riverine forest. Occasionally, reported in savanna, and cocoa plantations (herbarium). In Ghana, mostly found in south marginal and dry semi-deciduous forests (Hall & Swaine 1981).

Deciduousness: leaves die off completely. Flowers appear before the new leaves. Timing: flowering period: January to April (De Koning 1983)

Habitat Species’ presence is significantly correlated with rainfall (logistic regression), and found mainly where rainfall is between 2000-2500 mm/yr (Chi2 test). It grows often under forest shade, but also along paths and roadsides. Usually found close to water (e.g. creeks, rivers, and waterfalls). Soils can be clayish, sandy, lateritic or granite (herbarium).

Regeneration Data sources

The seeds germinate after 3-6 weeks, with a high germination success (90-100%) (De Koning 1983). It regenerates in shade (Hall & Swaine 1981).

Hall & Swaine (1981), De Koning (1983)

S

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

S.m.

60

42 (31)

5

48

7

9

35

47

8

3

82

15

35

3

H 58

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

339

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Page 340

Scaphopetalum amoenum

A.Chev.

Sterculiaceae Description

to rivers or in swamps (Hawthorne & Jongkind 2004). On humid and sandy soils (herbarium).

Guild: sb Life form: small arching tree Max. height: 15 m (FWTA) Max. diameter: data unavailable Leaf: alternate, simple, oblong, mesophyll (3.512.5 x 10-34 cm), entire Inflorescence: axillary, several flowers clustered together Flower: small to medium-sized; bowl-shaped; corolla yellowish with purple streaks Fruit: dry dehiscent (capsule), ellipsoid (1.5 cm long), 5-ribbed, greenish white; longitudinally furrowed Seed: medium-sized Other: it has a peculiar life form. Branches bend and take root when they touch the ground, thus forming a dense monospecific thicket in the forest understorey, within which no herbs occur and natural regeneration of the emergent trees is impossible. It has distinctive hairy twigs.

Regeneration It regenerates in shade both from seed, and vegetatively by layering of the branches (Hall & Swaine 1981).

Phenology Timing: not clearly seasonal (Hall & Swaine 1981)

Distribution Continent: Upper Guinea endemic Upper Guinea: Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 22 30’ cells, distribution range is 1040 km Forest type: wet evergreen forest, semi-deciduous forest, secondary forest. It can be locally abundant in wet evergreen forest and in swamps of semideciduous forests (Hawthorne & Jongkind 2004).

Data sources FWTA, Jenìk (1969), Hall & Swaine (1981), Hawthorne (1995a), Hawthorne & Jongkind (2004)

Habitat Species occurrence increases significantly with rainfall higher than 1500 mm/yr (Chi2 test). Usually under the rainforest understorey where it can form thickets by vegetative spread, especially so on slopes (herbarium, Hawthorne 1995a). Often close

S

spp

340

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

S.a.

47

33 (30)

0

51

9

2

60

10

27

4

53

43

49

4

H 43

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 341

Schizocolea linderi

(Hutch. & Dalziel) Bremek.

Rubiaceae Description

Phenology

Guild: sb Life form: small shrub Max. height: 3 m (Hawthorne & Jongkind 2004) Max. diameter: data unavailable Leaf: opposite, simple, elliptic, notophyll (2-5 x 615.5 cm), entire, coriaceous, hairy midrib above; interpetiolar stipules Inflorescence: axillary, solitary flowers in clusters Flower: medium-sized; calyx light green, densely hairy; corolla white with a long tube; fragrant Fruit: fleshy, ovoid (0.7 x 1 cm), red with a white pulp, hairy; 1 seed Seed: medium-sized (0.8 x 0.4 x 0.4 cm) Other: the stems, petioles and midribs are hairy.

Dispersal: probably by animals

Distribution

Data sources

Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire Distribution type: continuous, regional, present in 15 30’ cells, distribution range is 596 km Forest type: wet evergreen forest, moist evergreen forest

Hawthorne & Jongkind (2004)

Habitat Species occurrence increases with rainfall higher than 2500 mm/yr to reach an optimum around 2800 mm/yr (logistic regression analysis, Chi2 test). It is found under deep shade and in wet places such as along streams and riverbanks (Chi2 test) (herbarium).

S

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

H

S.l.

26

12 (17)

8

65

0

0

8

31

62

0

96

4

85

0

15

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

341

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Page 342

Schumanniophyton problematicum

(A.Chev.) Aubrév.

Rubiaceae Description

Phenology

Guild: sb Life form: small tree Max. height: 12 m (FWTA) Max. diameter: 5 cm (herbarium) Leaf: opposite, simple, obovate, macrophyll (25-40 x 35-45 cm), dentate, herbaceous, in whorls of 3 at the extremity of branches, rather glabrous or with few hairs on midrib; interpetiolar stipules Inflorescence: axillary, flowers and fruits clustered on horizontal branches Flower: large (corolla tube approx. 5 cm long); corolla tube cream-coloured, approx. 10 lobes, mottled brown Fruit: fleshy, ellipsoid to ovoid (4 x 5.5 cm), pale brown outside, yellow inside, in clusters of 5 or 6; many seeds Seed: dark brown Other: it has hard yellow wood. The stems are triangular at the nodes.

Deciduousness: deciduous, which is a very rare phenomenon for understorey species of evergreen forests (Hall & Swaine 1981)

Data sources FWTA, Savill & Fox (1967), Hall & Swaine (1981), Houghton (1988)

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Côte d’Ivoire, Ghana Distribution type: Upper Guinea disjunct, widespread, present in 9 30’ cells, distribution range is 1086 km, Red List species (Vulnerable) Forest type: wet evergreen forest, moist evergreen forest, semi-deciduous forest

Habitat Found in the understorey, especially in wet places in the forest (Savill & Fox 1967).

Regeneration It regenerates in shade.

S

spp

342

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

S.p.

13

0 (10)

0

46

0

38

30

15

15

8

62

31

38

0

H 54

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 343

Scytopetalum tieghemii

Hutch. & Dalziel

Scytopetalaceae Description

Regeneration

Guild: sb Life form: large tree Max. height: 40 m (Letouzey 1961) Max. diameter: 100 cm (Letouzey 1961) Leaf: alternate, simple, elliptic to oblong, microphyll (2-4 x 5-12 cm), entire, coriaceous, glabrous, with about 5 pairs of lateral nerves Inflorescence: axillary, not branched (short racemes) Flower: small ( 0.5 cm long); perianth white with corolla tube reddish at the base; 12-16 tepals; fragrant Fruit: fleshy inside, endocarp dehiscent (drupe), ovoid (2 cm long), woody and ribbed outside, red Seed: large (1.2 x 1.0 x 0.6 cm) Other: the wood is yellowish, hard, and fibrous.

It is a gregarious species (herbarium), that can regenerate in the shade (Hall & Swaine 1981).

Phenology Deciduousness: evergreen (Savill & Fox 1967) Dispersal: Hawthorne (1995a) commented that although one would expect some kind of animal dispersal, Taylor (1960) referred to the apparent lack of it. Timing: flowering period from March to April; fruiting period in July (Savill & Fox 1967)

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 33 30’ cells, distribution range is 1184 km Forest type: wet evergreen forest, moist evergreen forest, semi-deciduous forest, swamp forest, secondary forest. In Ghana, it is mostly found in wet evergreen forests (Hall & Swaine 1981). It can be locally abundant (herbarium), but it is rather rare (Savill & Fox 1967).

Habitat Species occurrence increases with rainfall to reach an optimum around 2200 mm/yr (logistic regression analysis, Chi2 test). It is found in the understorey of dense forests but often also along forest borders. Usually in very moist places of the forest or close to rivers (Chi2 test). In Ghana, it is highly correlated with acidic, base poor soils (Hall & Swaine 1981). On sandy, clay, or rocky soils (herbarium).

Uses The gum from the wood is locally used as a glue. The wood is not easy to work and not resistant to decay (Savill & Fox 1967).

Data sources FWTA, Taylor (1960), Letouzey (1961), Savill & Fox (1967), Hall & Swaine (1981), Hawthorne (1995a)

S

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

L

Soil WHC M

H

S.t.

75

40 (40)

0

51

3

3

50

26

19

7

71

23

41

4

49

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

343

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Page 344

Sericanthe adamii

(N.Hallé) Robbr.

Rubiaceae Description

Data sources

Guild: u Life form: large woody climber, climbing with recurved spines Max. height: 35 m long Max. diameter: data unavailable Leaf: opposite, simple, elliptic, notophyll (3-6 x 6-11 cm), entire, herbaceous, slightly hairy on the nerves; with interpetiolar stipules Inflorescence: axillary, solitary Flower: small; 8-merous; pedicel pubescent Fruit: fleshy (drupe), globose (2.5 cm in diameter), hard woody pericarp, rough, orange to red, covered with distinct muricate ridges and persistent calyx; 2-4 seeds Seed: large (approx. 1.2 cm in diameter)

Robbrecht (1981a,b), Hawthorne & Jongkind (2004)

Distribution Continent: Upper Guinea endemic Upper Guinea: Liberia Distribution type: continuous, very local, present in 2 30’ cells, distribution range is 6 km Forest type: wet evergreen forest

Habitat It occurs around 500 m altitude (Robbrecht 1981a).

Phenology

S

spp

344

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

S.a.

2

0 (2)

50

0

0

0

0

100

0

0

100

0

100

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 345

Sherbournia calycina

(G.Don) Hua

Rubiaceae Description Guild: pi Life form: large winding woody climber Max. height: 50 m long, 10 m high (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, elliptic, mesophyll (3-7 x 7-18 cm), entire, coriaceous, sometimes with red spots underneath; petiole first pubescent, thereafter glabrous; interpetiolar stipules Inflorescence: axillary, solitary Flower: large (tube 3 cm long); campanulate; large calyx lobes; corolla white to yellowish outside, pink to mauve inside; not fragrant Fruit: fleshy, ovate to ellipsoid (2.5 x 3 cm), yellow to orange; with persistent calyx; many seeds Seed: small (0.2 x 0.1 cm)

Phenology

Distribution

Deciduousness: probably deciduous Timing: flowering period from October to May; fruiting period from November to August (De Koning 1983)

Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana, Togo Distribution type: continuous, widespread, present in 48 30’ cells, distribution range is 1530 km Forest type: wet evergreen forest, gallery forest, secondary forest, thickets

Habitat Species occurrence increases with rainfall to reach an optimum around 2700 mm/yr (logistic regression analysis, Chi2 test). Commonly found in disturbed places (e.g. forest gaps and edges, severely logged forests, and often in secondary thickets along roadsides). However, it has also been recorded in dense rainforests. Often close to rivers (Chi2 test). Soils can be lateritic or sandy-clayish (herbarium).

Regeneration

Data sources

It has a phanerocotylar epigeal foliaceous seedling type (cf. de Koning 1983).

FWTA, De Koning (1983), Hawthorne & Jongkind (2004)

S

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

H

S.c.

125

44 (79)

4

51

8

3

46

26

24

2

68

30

36

6

46

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

345

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Page 346

Sorindeia collina

Keay

Anacardiaceae Description

Phenology

Guild: sb Life form: small tree Max. height: 10 m (Keay 1956) Max. diameter: 5 cm (herbarium) Leaf: alternate or sub-opposite, imparipinnately compound, approx. 9 leaflets, lanceolate, mesophyll (3-13 x 12-30 cm), herbaceous, glabrous Inflorescence: cauliflorous, branched (panicle, up to 30 cm long) Flower: small, reddish Fruit: fleshy (drupe), ellipsoid (2 cm long), glabrous Seed: data unavailable Other: it has aerial roots.

Data sources

Distribution

FWTA, Keay (1956), Savill & Fox (1967), Hawthorne & Jongkind (2004)

Continent: Upper Guinea endemic Upper Guinea: Sierra Leone Distribution type: continuous, local, present in 5 30’ cells, distribution range is 252 km Forest type: montane high forest (FWTA). It is locally common (herbarium).

Habitat An understorey tree in montane high forests (herbarium).

S

spp

346

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

S.c.

6

0 (3)

50

33

0

0

0

33

66

0

33

67

33

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 347

Soyauxia grandifolia

Gilg & Stapf

Medusandraceae Description Guild: sb Life form: small to medium-sized tree Max. height: 25 m (herbarium) Max. diameter: data unavailable Leaf: alternate, simple, elongate to oblong, mesophyll (4-11 x 12-32 cm), entire, coriaceous, glabrous; with paired stipules (approx. 0.2 x 1cm) Inflorescence: axillary, unbranched (densely flowered, 6-12 cm long) Flower: small; calyx green to brownish; corolla white Fruit: dry dehiscent, subglobose (2.3 x 2.8 cm), reddish; 1 seed Seed: large (up to 2 cm in diameter) Other: the bark has many small lenticels.

Regeneration

Distribution

It regenerates in the shade (Hall & Swaine 1981).

Continent: Upper Guinea endemic Upper Guinea: Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 12 30’ cells, distribution range is 1004 km Forest type: wet evergreen forest, coastal forest, coastal savanna, secondary forest. In Ghana, only in wet evergreen forests (Hall & Swaine 1981).

Phenology

Habitat The species is more common in the lowlands (logistic regression analysis) where it can be found in coastal forests and savannas. Its occurrence increases with rainfall (logistic regression analysis), especially when rainfall is higher than 2500 mm/yr (Chi2 test). It is occasionally found on riverbanks. In Ghana, it is found only on very acid, base-poor soils (Hall & Swaine 1981), but in Liberia and Côte d’Ivoire, it has been recorded on lateritic, loamy sands, sandy clay, and sandy soils (herbarium).

Data sources FWTA, Aubréville (1959), Hall & Swaine (1981), Hawthorne & Jongkind (2004)

S

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

H

S.g.

19

44 (18)

0

47

21

0

26

26

42

5

74

21

47

11

32

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

347

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Page 348

Strephonema pseudocola

A.Chev.

Combretaceae Description

(e.g. rivers, waterfalls, and even swamps). Often on sandy soils (herbarium).

Guild: sb Life form: large tree Max. height: 30 m (herbarium) Max. diameter: 90 cm (herbarium) Leaf: alternate, simple, oblanceolate, mesophyll (5-10 x 16-32 cm), entire, coriaceous, often with glands in the margin, glabrous, young leaves pubescent Inflorescence: axillary, unbranched, with brown hairs Flower: medium-sized (petals 5-7 mm long); bisexual; corolla white; filaments white and glabrous; anthers yellow and septate Fruit: dry indehiscent, resembling a large gall (5.5 x 4 x 3.5 cm), subglobose, sessile, succulent, glabrous, brown; 1 seed Seed: very large (5 cm), red, resembles cola nut Other: a large cylindrical tree with a narrow, dense crown. The bark is grey and exudes a translucent jelly. The young epicotyl of seedlings is wine-red. The wood density is 0.69 g/cm3.

Regeneration The heavy seeds are not easily transported by animals. Therefore, regeneration occurs mainly below the parent tree. In Banco forest the species can form clumps (De Koning 1983). It regenerates in the shade (Hall & Swaine 1981). Germination occurs after 2-3 months. It has a cryptocotylar hypogeal reserve seedling type (cf. De Koning 1983).

Phenology Deciduousness: evergreen Dispersal: barochore Timing: flowering period from August to February; fruiting period from October to March

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continental disjunct, in Upper Guinea present in 25 30’ cells Forest type: wet and moist evergreen forest, semideciduous forest, secondary forest

Habitat

Data sources

Species occurrence increases strongly with rainfall higher than 1500 mm/yr (Chi2 test) to reach an optimum around 2500 mm/yr (logistic regression analysis). It is found in mature forest or old secondary forests. Sometimes found close to water

FWTA, Voorhoeve (1965), Hall & Swaine (1981), De Koning (1983), Vivien & Faure (1985), Tailfer (1989), Jongkind (1995)

S

spp

348

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

S.p.

53

20 (25)

6

51

0

2

40

38

19

13

66

21

32

6

H 47

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 349

Strombosiopsis nana

Breteler

Olacaceae Description

Phenology

Guild: u Life form: shrub, treelet Max. height: 3 m Max. diameter: data unavailable Leaf: alternate, simple, elliptic to obovate-elliptic, notophyll (3-5 x 9-15 cm), revolute, coriaceous, glabrous; petiole winged; no stipules Inflorescence: axillary, branched (fasciculate, up to 6 together) Flower: small; corolla creamy white Fruit: fleshy, obovoid to ellipsoid (1.5 x 2 cm), red with yellow stripes, smooth Seed: data unavailable Other: the branchlets are green and slightly compressed to narrowly winged.

Data sources

Distribution

FWTA, Breteler (2001), Hawthorne & Jongkind (2004)

Continent: Upper Guinea endemic Upper Guinea: Liberia Distribution type: continuous, very local, present in 2 30’ cells. Recorded only in Liberia. Forest type: data unavailable

Habitat Found in rainforest at altitudes between 0-100 m (Breteler 2001).

S

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

S.n.

3

0 (1)

0

33

67

0

0

0

33

0

100

0

33

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

349

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Page 350

Strychnos dinklagei

Gilg

Loganiaceae

Regeneration It regenerates in primary forest (De Rouw 1991)

Phenology Description

Distribution

Guild: np Life form: large woody climber Max. height: 40 m long (herbarium) Max. diameter: 13 cm (herbarium) Leaf: opposite, simple, elliptic, notophyll (2-7 x 3-13 cm), revolute, coriaceous, glabrous on both sides Inflorescence: terminal or axillary, branched (panicle, up to 10 cm long, many flowered) Flower: small (corolla 0.3 cm long); calyx greenish white; corolla greenish white, 5-merous Fruit: fleshy, disk-shaped (2 x 1.5 cm), orange, slimy pulp; 1 seed Seed: elliptic, large (1.7 x 1.1 x 0.5 cm), white with thick very short erect hairs Other: it starts as a scandent shrub. The branches are lenticellate.

Continent: Upper Guinea endemic Upper Guinea: Guinea, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 15 30’ cells, distribution range is 1399 km Forest type: rainforest, coastal forest, mangroves, secondary forest

Dispersal: by animals (monkeys, elephants)

Habitat The species is more abundant in lowlands (logistic regression analysis, Chi2 test), and in places where rainfall reaches 1500-2000 mm/yr (Chi2 test). It is usually found near the coast (Chi2 test) or near rivers (Chi2 test). Sometimes in swampy places. Often on white sand or on (at the edge of) granite rocks (herbarium).

S

spp

350

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

H

S.d.

50

46 (28)

0

10

58

14

60

4

10

36

44

20

18

34

28

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

Data sources FWTA, Leeuwenberg (1969), De Rouw (1991)

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Page 351

Strychnos melastomatoides

Gilg

Loganiaceae Description Guild: u Life form: woody climber Max. height: data unavailable Max. diameter: data unavailable Leaf: opposite, simple, elliptic to narrowly ovate, mesophyll (2-8 x 5-19 cm), coriaceous, glabrous, two pairs of secondary veins curved along the margin Inflorescence: axillary or terminal, branched (up to 6 cm long and 3 cm wide, lax, several-flowered) Flower: small (corolla tube 0.4 cm long); calyx glabrous on both sides; corolla white, glabrous outside, hairy inside; fragrant Fruit: fleshy, ellipsoid (3 cm in diameter), orange; 1-3 seeds Seed: flattened, obliquely elliptic, large (1.8 x 1.4 x 0.5 cm) Other: the branchlets are pale to dark brown and glabrous. It climbs with tendrils.

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone Distribution type: continuous, local, present in 5 30’ cells, distribution range is 247 km Forest type: rainforest, gallery forest

Phenology Dispersal: by animals (monkeys, elephants)

Habitat It occurs along riverbanks or in gallery forest, at low elevations (Leeuwenberg 1969).

Data sources FWTA, Leeuwenberg (1969), Hawthorne & Jongkind (2004)

S

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

S.m.

7

- (0)

0

43

0

0

0

57

43

0

57

43

14

14

H 43

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

351

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Page 352

Strychnos millepunctata

Leeuwenb.

Loganiaceae Description

Distribution

Guild: sb Life form: medium-sized woody climber Max. height: 25 m long (herbarium) Max. diameter: 4 cm (herbarium) Leaf: opposite, simple, obovate, notophyll (2-8 x 4-15 cm), entire, coriaceous Inflorescence: axillary, solitary to few flowered (up to 1.5 cm long) Flower: small (corolla tube 0.3 cm long); corolla yellow, 5-merous Fruit: fleshy, globose (1.3 cm in diameter), orange; 1 seed Seed: ellipsoid, medium-sized (0.9 x 0.8 x 0.7 cm), pale brown, densely pubescent Other: it has paired tendrils and climbs with hooks.

Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire Distribution type: continuous, regional, present in 9 30’ cells, distribution range is 392 km, Red List species (Vulnerable) Forest type: rainforest, coastal forest, riverine forest, secondary forest

Habitat It is usually found on riverbanks, on sandy soils (herbarium).

Regeneration It regenerates in primary forest (De Rouw 1991).

Phenology Dispersal: by animals (monkeys, elephants)

Data sources Leeuwenberg (1969), De Rouw (1991), IUCN Red List (2000)

S

spp

352

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

S.m.

14

18 (11)

0

14

36

36

64

0

0

36

57

7

43

36

21

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 353

Strychnos odorata

A.Chev.

Loganiaceae Description

Phenology

Guild: u Life form: large woody climber Max. height: 40 m high, 100 m long (herbarium) Max. diameter: 6 cm (herbarium) Leaf: opposite, simple, elliptic, notophyll (2-4 x 7-12), entire, herbaceous, aromatic Inflorescence: axillary, compound, glabrous, manyflowered Flower: medium-sized (up to 1.5 cm long); corolla pale yellow Fruit: fleshy, obliquely ellipsoid (2.3 x 1.9 x 1.6 cm); 1-3 seeds Seed: flattened, ellipsoid, large (1.6 x 1 x 0.7 cm), at one side with a deep pit Other: it climbs with hooks.

Dispersal: by animals (monkeys, elephants)

Data sources

Distribution

FWTA, Leeuwenberg (1969) Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire Distribution type: continuous, very local, present in 3 30’ cells, distribution range is 89 km Forest type: rainforest, secondary forest

Habitat Found in moist places in rainforest at low elevations, usually, on riverbanks (Leeuwenberg 1969).

S

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

S.o.

4

0 (2)

0

75

0

0

100

0

0

0

50

50

0

0

100

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

353

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Page 354

Strychnos soubrensis

Hutch. & Dalziel

Loganiaceae Habitat Frequently, found in moist places (e.g. along creeks, and riverbanks), but also reported in dry thickets on hillsides and along roads. In both mature and secondary forests. On sandy, clayish, rocky and lateritic soils (herbarium).

Regeneration It has a phanerocotylar epigeal foliaceous seedling type (cf. De Koning 1983), and regenerates in the shade (Hall & Swaine 1981).

Description

Phenology

Guild: np Life form: large woody climber Max. height: 100 m long (herbarium) Max. diameter: 10 cm (herbarium) Leaf: opposite, simple, elliptic, microphyll (1-5 x 2-14 cm), entire, herbaceous, smooth Inflorescence: axillary or terminal, unbranched (up to 1.5 cm long) Flower: small; calyx pale green with almost whitish margin; corolla white, hairs inside, 5-merous; fragrant Fruit: fleshy, globose (2.5 cm in diameter), orange; 2-10 seeds Seed: irregularly ovate, flattened, large (1.3 x 0.9 x 0.5 cm), brown Other: it starts as a straggling shrub and climbs with paired hook-like tendrils. It has pale and conspicuous lenticels.

Dispersal: by animals (monkeys, elephants) Timing: flowering period from December to April; fruiting period in December (De Koning 1983)

Data sources

Distribution type: continuous, widespread, present in 47 30’ cells, distribution range is 2071 km Forest type: wet evergreen forest, moist evergreen forest, moist semi-deciduous and dry semideciduous forest, secondary forest, thickets. In Ghana, it is common in a wide variety of forest types (Hall & Swaine 1981).

Distribution Continent: Benin, Nigeria (Hall & Swaine 1981) Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana, Togo

Leeuwenberg (1969), Hall & Swaine (1981), De Koning (1983)

S

spp

354

n

Open (n)

S.s.

101

53 (70)

All

46333

37

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

L

M

H

L

M

H

46

4

11

44

35

11

7

69

24

31

7

58

39

7

37

29

24

10

6

69

25

13

39

9 4

Soil CMK

Soil WHC

36

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Page 355

Suregada ivorensis

(Aubrév. & Pellegr.) J.Léonard

Euphorbiaceae Description

Phenology

Guild: u Life form: medium-sized to large tree Max. height: 25 m (herbarium) Max. diameter: 40 cm (herbarium) Leaf: alternate, simple, entire Inflorescence: data unavailable Flower: data unavailable Fruit: data unavailable Seed: data unavailable

Distribution Continent: Upper Guinea endemic Upper Guinea: southeast Côte d’Ivoire Distribution type: continuous, very local, present in 2 30’ cells. The species is endangered (C. Chatelain, pers. comm.). Forest type: wet evergreen forest, secondary forest

Data sources Léonard (1958), Aubréville (1959), Hawthorne & Jongkind (2004)

Habitat Data unavailable.

S

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

S.i.

2

100 (1)

0

100

0

50

50

0

0

0

50

50

50

0

H 50

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

355

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Page 356

Tabernaemontana africana

Hook.

Apocynaceae Distribution type: continuous, widespread, present in 82 30’ cells, distribution range is 1803 km Forest type: wet evergreen forest, moist semideciduous forest, dry semi-deciduous forest

Habitat Species occurrence increases with rainfall to reach a very wide optimal region between 2000-3000 mm/yr (logistic regression analysis, Chi2 test). It can be found in the forest understorey but also in disturbed areas and forest edges (herbarium).

Phenology Timing: flowering and fruiting occurs mainly in the dry season (Leeuwenberg 1991)

Description Guild: sb Life form: small tree Max. height: 8 m (herbarium) Max. diameter: 5 cm (herbarium) Leaf: opposite, simple, mesophyll (> 15 cm long), coriaceous, glabrous Inflorescence: data unavailable Flower: large; calyx light green; corolla tube yellowish outside, inside yellow at apex, corolla lobes white; fragrant Fruit: fleshy (berry), dehiscent, double (approx. 10 cm in diameter), yellow to orange Seed: medium-sized to large Other: the slash exudes a white latex. The flowers are open during daytime and closed at night.

Data sources Hall & Swaine (1981), Leeuwenberg (1987, 1991), Hawthorne (1995a), Hawthorne & Jongkind (2004)

Distribution Continent: Upper Guinea endemic Upper Guinea: Senegal, Guinea Bissau, Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana

T

spp

356

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

H

T.a.

171

36 (33)

7

36

11

17

33

21

29

3

60

37

29

11

39

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 357

Talbotiella gentii

Hutch. & Greenway

Leguminosae-Caes. Description

Habitat

Guild: sb Life form: medium-sized tree Max. height: 20 m (herbarium) Max. diameter: data unavailable Leaf: paripinnately compound, 8-14 opposite leaflets, nanophyll (1 x 2 cm), youngest twigs with leafy stipules at nodes when flushing and with brown hairs, but soon glabrous Inflorescence: axillary, unbranched (raceme, approx. 3 cm long) Flower: small; corolla pink and white Fruit: dry dehiscent (pod), almost triangular, flatended (5 cm long), explosive Seed: large Other: the slash is rather fibrous and peelable, reddish with orange-brown inner bark. It is not ectomycorrhizal.

It is found in dry and rocky forest.

Regeneration It regenerates in shade. It is a gregarious species that often forms monospecific stands in which most of the understorey consists of young plants of Talbotiella (Hall & Swaine 1981).

Phenology Deciduousness: evergreen Dispersal: explosive

Distribution Continent: Upper Guinea endemic Upper Guinea: Ghana Distribution type: continuous, local, present in 6 30’ cells, distribution range is 199 km. It is confined to eastern Ghana. This is a very rare, albeit gregarious, species on the edge of extinction, and Ghana’s greatest conservation priority (Hall & Swaine 1981). The record for Cameroon (e.g. Lock 1989) seems to be an error, as no specimen like this has been seen from Cameroon at Kew. It would not be surprising to find isolated pockets of this species in dry, rocky forest remnants in the Guinea sub-region, however, but this would not be likely to alter its threatened status (Hawthorne 1995a). Forest type: found in dry evergreen forests along the Afram or Volta basins (Hawthorne & Jongkind 2004), often forming a belt between savanna and deciduous forests. Found in Southern Marginal and Southeast outlier forest types (Hall & Swaine 1981).

Data sources FWTA, Hall & Swaine (1981), Lock (1989), Hawthorne (1995a), Hawthorne & Jongkind (2004)

T

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

T.g.

23

10 (20)

13

30

0

74

26

0

0

0

22

78

0

22

48

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

357

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Page 358

Tapinanthus belvisii

(DC.) Danser

Loranthaceae Description

Phenology

Guild: pi Life history: hemi-parasitic shrub Max. height: 1 m Max. diameter: data unavailable Leaf: opposite, simple, ovate, notophyll (2-8 x 4-11 cm), entire, coriaceous, glabrous, the lower 2-3 pairs of lateral nerves stronger and more ascending, dull green, reddish when flushed Inflorescence: axillary, umbel (sometimes several around old nodes, with 8-16 flowers in a ring) Flower: medium-sized (corolla tube 2.7 cm long); corolla tube mostly red, lobes green, turning violetblack outside, greenish inside Fruit: fleshy (berry), subglobose (0.7 cm in diameter), shiny red; 1 seed Seed: medium-sized Other: the branchlets are velutinous.

Dispersal: by birds Timing: flowering period mostly from January to April, also in August (Polhill & Wiens 1998)

Data sources FWTA, Balle & Hallé (1961), Polhill & Wiens (1998)

Distribution Continent: Upper Guinea endemic (Polhill & Wiens 1998) Upper Guinea: Sierra Leone, Côte d’Ivoire, Ghana (herbarium, Polhill & Wiens 1998) Distribution type: continuous, widespread, present in 8 30’ cells, distribution range is 767 km. Only one observation in Sierra Leone, without precise location, which is an ancient record (Polhill & Wiens 1998). Forest type: coastal shrubland

Habitat Found in coastal savanna and bushlands, and at forest edges. Inland found along rivers. It grows on various hosts, including cultivated trees (Polhill & Wiens 1998). On sandy soils (herbarium).

T

spp

358

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

T.b.

20

50 (2)

0

25

65

15

60

25

0

45

35

20

5

5

45

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 359

Tapinanthus praetexta

Polhill & Wiens

Loranthaceae Description

Phenology

Guild: pi Life form: hemi-parasitic shrub Max. height: 0.7 m (Polhill & Wiens 1998) Max. diameter: data unavailable Leaf: opposite, simple, elliptic to lanceolate, microphyll (2-5 x 5-9 cm), entire, herbaceous Inflorescence: axillary, umbel (with 8-12 flowers) Flower: medium-sized (corolla tube 2.2 cm long); corolla tube pale pink, lobes red to purple Fruit: fleshy (berry), globose (0.6 cm in diameter), red; 1 seed Seed: medium-sized

Dispersal: by birds Timing: flowering period from July to February (Polhill & Wiens 1998)

Distribution Continent: Upper Guinea endemic (Polhill & Wiens 1998) Upper Guinea: Côte d’Ivoire (Polhill & Wiens 1998) Distribution type: continuous, very local, present in 3 30’ cells, distribution range is 46 km. It has a very restricted distribution; only known from the Banco and Adiopoudome forest in southeast Côte d’Ivoire (Polhill & Wiens 1998). Forest type: coastal forest

Data sources Polhill & Wiens (1998)

Habitat Data unavailable.

T

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

T.p.

11

100 (2)

0

91

0

0

45

55

0

64

18

18

0

55

27

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

359

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Tapura fischeri

Page 360

Engl.

Dichapetalaceae Description

Regeneration

Guild: u Life form: medium-sized tree Max. height: 20 m (Breteler 1986) Max. diameter: 45 cm (Breteler 1986) Leaf: alternate, simple, elliptic, notophyll (1-7 x 2-16 cm), entire, herbaceous Inflorescence: axillary, umbel (very distinctive as if growing from the apex of the leaf stalk in clusters of about 1.2 cm across) Flower: small (0.2-0.5 cm long); corolla white, 2 petals; somewhat fragrant Fruit: dehiscent, ovoid (approx. 0.4 cm long), 3-lobed, densely hairy; 1-3 seeds Seed: medium-sized (0.4 x 0.15 cm), brown Other: a small, well-branched tree with a grey and rough bole. The branches grow horizontal or drooping.

In a study of the effects of fire in Asukese and other Ghanaian forest reserves, regeneration of this species was found to thrive in burnt forest (Hawthorne 1994).

Phenology

Distribution Continent: Nigeria, Cameroon, Congo (Brazzaville), Democratic Republic of Congo, Sudan, Ethopia, Uganda, Kenya, Tanzania, Malawi, Zimbabwe, Mozambique, South Africa (Breteler 1986) Upper Guinea: Côte d’Ivoire, Ghana (herbarium, Breteler 1986) Distribution type: continuous, widespread, in Upper Guinea present in 7 30’ cells Forest type: semi-deciduous forest, deciduous forest, dry forest, savanna, gallery forest

Data sources FWTA, Breteler (1986), Keay (1989), Hawthorne (1990, 1994, 1995a), Carr (1998), Hawthorne & Jongkind (2004)

Habitat In Ethopia, it is found in riverine forest, usually in clearings and on clayey soils (Carr 1998).

T

spp

360

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

T.f.

11

0 (6)

0

9

0

90

9

0

0

0

55

45

18

45

27

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 361

Tapura ivorensis

Breteler

Dichapetalaceae Description

Phenology

Guild: sb Life form: small tree Max. height: 10 m Max. diameter: data unavailable Leaf: alternate, simple, elliptic, mesophyll (3-8 x 9-17 cm), entire, coriaceous, with glands below Inflorescence: axillary, many flowers together (glomerulous) Flower: small; corolla light cream to yellow Fruit: fleshy, subovoid, laterally compressed (1.5 x 3 cm), slimy pulp, yellow; 2 seeds Seed: large (0.7 x 1.8 cm), brown Other: it has a knobbly bole. The branches grow horizontal, then drooping, to form an untidy tangledlooking crown.

Dispersal: probably by animals

Data sources Breteler (1986), Hawthorne (1990, 1995a), IUCN Red List (2000), Hawthorne & Jongkind (2004)

Distribution Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire, Ghana Distribution type: continuous, very local, present in 2 30’ cells, Red List species (Vulnerable) Forest type: wet evergreen forest

Habitat It occurs especially along streambanks (Hawthorne 1995a).

T

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

T.i.

3

0 (1)

0

33

0

0

67

33

0

0

100

0

0

0

100

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

361

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Tarenna vignei

Page 362

Hutch. & Dalziel

Rubiaceae Data sources

occurs in wet evergreen forest, gallery forest and secondary forest. T. vignei vignei is an Upper Guinea endemic that occurs only in Ghana. It has a very local distribution and is found in only 1 30’ cell. It can be found in evergreen forest.

FWTA, De Koning (1983), Hawthorne & Jongkind (2004)

Habitat T. vignei subglabra is mostly found where rainfall reaches 2000-2500 mm/yr (Chi2 test). Usually under dense shade (De Koning 1983), and close to streams and rivers (Chi2 test). On loamy soils (herbarium).

Description Guild: sb Life form: shrub or small tree Max. height: 7 m (Hawthorne & Jongkind 2004) Max. diameter: data unavailable Leaf: opposite, simple, rectangular to elliptic, notophyll (1.5-7 x 5.5-20 cm), entire, herbaceous, glabrous; lower surface of petiole has a channel; interpetiolar stipules Inflorescence: terminal, laxly branched cyme Flower: small (corolla tube 1 cm long, lobes 0.7 cm); tube-shaped, 5 lobes, white Fruit: fleshy (drupe), globose (1.1 cm in diameter), orange glossy with orange pulp, glabrous; many seeds Seed: angular, medium-sized (0.5 x 0.3 x 0.3 cm), brown Other: a slender tree with rather open crown and horizontal branches. T. vignei var. vignei differs from T. vignei subglabra in its rather densely pubescent lower leaf surface.

Regeneration It has a phanerocotylar epigeal foliaceous seedling type (cf. De Koning 1983).

Phenology Timing: flowering period from November to January; fruiting period from April to November (De Koning 1983)

Distribution T. vignei subglabra is a continental disjunct that occurs in Cameroon, Sierra Leone, Liberia, Côte d’Ivoire and Ghana. It has its largest population in Upper Guinea and is present in 27 30’ cells. It

T

spp

362

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

T.v.

66

22 (36)

2

61

0

3

36

53

10

0

80

20

22

0

78

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 363

Tetraberlinia tubmaniana

J.Léonard

Leguminosae-Caes. Description

bottom lands and slightly undulating land with sandy loam soils (Voorhoeve 1965). It occurs on drylands, between 0 and 100 m altitude (Wieringa 1999).

Guild: sb Life form: large tree Max. height: 42 m (Voorhoeve 1965) Max. diameter: 100 cm (Voorhoeve 1965) Leaf: alternate, paripinnately compound, 1-7 pairs of leaflets, rhombic, nanophyll (0.3-0.4 x 0.8-1.1 cm), entire, coriaceous, glabrous, glands present at the lower surface; red-brown hairs on twigs and rachis Inflorescence: axillary or terminal, branched (panicle) (up to 8 cm long), brown velvety-pubescent Flower: small; corolla white, largest petal yellow Fruit: dry dehiscent (pod), flat, oblong-obovate (5 x 12 cm), woody with central raised line, brown; 1-3 seeds Seed: disc-shaped, elliptic, very large (2.1 x 1.8 x 0.5 cm), dark brown Other: the bark is grey and flaky. The slash is pinkbrown. It has thick root swellings. The crown is often small and rounded, and only very large trees have a spreading crown. The wood density is 0.68 g/cm3.

Regeneration It has a phanerocotylar epigeal reserve seedling type (cf. Voorhoeve 1965). Seedlings are very abundant in the forest. The structure of the single dominant forest of Tetraberlinia is different from the other gregarious species such as Monopetalanthus compactus and Gilbertiodendron preussii. Besides an abundant regeneration at the forest floor, the species is equally well represented among mediumsized trees. This population structure gives prospects for silvicultural exploitation. However, the trees seem to suffer from a sudden exposure to full light, and the low diameter limit for exploitation results in a complete destruction of the forest; if not by direct damage of the remaining stand, then by the after effects of full exposure (Voorhoeve 1965). The species can flower from a dbh of 8 cm onwards (Voorhoeve 1965).

Distribution

Phenology

Uses

Continent: Upper Guinea endemic (Wieringa 1999) Upper Guinea: Liberia (Wieringa 1999) Distribution type: continuous, local, present in 7 30’ cells, distribution range is 283 km, Red List species (vulnerable). At present, the species is only known from Liberia, but it could be present in Sierra Leone as well (Voorhoeve 1965). Forest type: wet evergreen forest

Deciduousness: evergreen Dispersal: by explosive pods (Voorhoeve 1965) Timing: flowering period from April to June (Voorhoeve 1965); fruiting period from November to January (Voorhoeve 1965)

In Liberia, the wood is used for all kinds of purposes, such as construction, carpentry, etc. (Voorhoeve 1965).

Data sources Voorhoeve (1965), Lock (1989), Bongers et al. (1999), Wieringa (1999), IUCN Red List (2000), Hawthorne & Jongkind (2004)

Habitat Tetraberlinia is found above an annual rainfall of 2000 mm, and becomes abundant above 2500 mm/yr (Bongers et al. 1999). In the high forest, the tree is gregarious, forming small stands or extensive single dominant forests, especially on

T

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

T.t.

25

11 (9)

0

8

0

0

4

4

92

0

100

0

100

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

363

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Page 364

Thomandersia anachoreta

Heine

Acanthaceae Description

Phenology

Guild: np Life form: treelet or shrub Max. height: 2 m (Heine 1966) Max. diameter: data unavailable Leaf: opposite, simple, lanceolate, notophyll (3-5 x 6-13 cm), entire, coriaceous Inflorescence: axillary or terminal, not branched (10-20 cm long) Flower: yellowish green or bright brownish pink Fruit: dry dehiscent (capsule), ellipsoid (1.6 x 0.9 cm) Seed: medium-sized (0.3 cm)

Distribution

Data sources

Continent: Upper Guinea endemic Upper Guinea: Liberia, Côte d’Ivoire (Heine 1966, herbarium) Distribution type: continuous, regional, present in 4 30’ cells, distribution range is 390 km Forest type: swamp forest, secondary forest

Heine (1966)

Habitat In disturbed forest, along forest edges or roads. Sometimes near rivers or swampy areas. On rocky or red loamy soils (herbarium).

T

spp

364

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

T.a.

7

33 (6)

0

14

43

0

43

58

0

14

86

0

29

14

57

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 365

Tiliacora leonensis

(Scott-Elliot) Diels

Menispermaceae Description

occasionally also in mature rainforest. Frequently on sandy soils but sometimes also on clayish ones (herbarium).

Guild: np Life form: winding woody climber Max. height: 25 m (herbarium) Max. diameter: 7 cm (herbarium) Leaf: alternate; simple, oblong, mesophyll (3-9 x 5-20 cm), coriaceous, with 5 pairs of lateral nerves looping together conspicuously; petiole 1.5 cm long, hairy Inflorescence: dioecious, cauliflorous, male cyme (3-6 cm long), peduncle hairy; female inflorescence unknown Flower: sessile; small, bowl-shaped, yellow, fragrant Fruit: fleshy, obovoid (1.8 x 1 cm); red-brown to black; 1 seed Seed: large (1.5 x 0.8 x 0.7 cm) Other: the stem is somewhat lobed. The sapwood is yellowish and densely layered in concentric rings. It has no exudate.

Regeneration It regenerates in shade, but many records were found in disturbed habitats.

Phenology

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d'Ivoire, Ghana. Distribution type: continuous distribution, widespread, distribution range is > 1411 km Forest type: wet evergreen forest, moist evergreen forest, moist semi-deciduous forest, dry semideciduous forest, riverine forest, savanna, and secondary forest.

Data sources FWTA, Troupin (1962), Hall & Swaine (1981), De Koning (1983), Hawthorne & Jongkind (2004)

Habitat Species occurrence increases in places where rainfall reaches 1500 -2000 mm/year (Chi2 test), and where soils are base-poor (Hall & Swaine 1981). Most often in secondary forests (herbarium, De Koning 1983) and riverine forests but

T

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

T.l.

71

39 (44)

4

35

18

13

50

24

13

7

76

17

39

11

42

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

365

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Page 366

Trichilia djalonis

A.Chev.

Meliaceae Description

Phenology

Guild: u Life form: small to medium-sized tree Max. height: 15 m (herbarium) Max. diameter: 45 cm (herbarium) Leaf: alternate, imparipinnately compound (28.5 cm long), opposite leaflets, lanceolate to elliptic, notophyll (4.5-5 x 10-13 cm), entire Inflorescence: branched (2 cm long) Flower: small (0.7 cm long); corolla white to yellowish, petals free Fruit: dry dehiscent, globose (1-2 cm in diameter), weakly 3-lobed, green with pale brown hairs Seed: large (1.2 cm long), with an aril Other: the twigs are pubescent.

Dispersal: probably by animals (Hawthorne & Jongkind 2004)

Distribution

Data sources

Continent: Upper Guinea endemic Upper Guinea: Guinea, Liberia, Côte d’Ivoire Distribution type: continuous, widespread, present in 8 30’ cells, distribution range is 938 km Forest type: montane forest

FWTA, Hawthorne & Jongkind (2004)

Habitat Mostly on the upper reaches of closed or open montane rainforests (higher than 650 m, Hawthorne & Jongkind 2004). On itabirite or clayish soils containing 40% iron (herbarium).

T

spp

366

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

T.d.

12

33 (9)

67

50

0

8

33

50

8

0

67

33

42

0

33

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 367

Trichilia megalantha

Harms

Meliaceae Description

Phenology

Guild: np Life form: medium-sized to large tree Max. height: 35 m (herbarium) Max. diameter: 35 cm (herbarium) Leaf: alternate, pinnately compound, 10-12 leaflets, oblong to lanceolate, notophyll (3.8-4 x 8-18 cm), entire, pubescent beneath Inflorescence: dioecious, axillary, branched (approx. 12 cm long) Flower: large (petals 2 x 3.5 cm); corolla creamcoloured, dish-shaped, 6-merous; very fragrant Fruit: dry dehiscent (capsule) (2 cm in diameter), greyish pink; 6 seeds Seed: large (1.3 x 0.7 cm), plano-convex, black with orange-red aril Other: it has a rough, dark grey bark.

Dispersal: probably by animals (Hawthorne & Jongkind 2004)

Data sources FWTA, De Wilde (1968), Hall & Swaine (1981), Hawthorne & Jongkind (2004)

Distribution Continent: Guinea wide: from Côte d’Ivoire to Nigeria (Hall & Swaine 1981), Benin, Nigeria (herbarium) Upper Guinea: Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 6 30’ cells, distribution range is 2047 km Forest type: dense humid forest, deciduous forest, savanna, secondary to very degraded forest (herbarium). In Ghana, it is found only in semideciduous forests, especially dry ones (Hall & Swaine 1981, Hawthorne & Jongkind 2004).

Habitat Data unavailable.

T

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

T.m.

8

17 (6)

0

50

0

63

13

25

0

0

75

25

0

25

75

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

367

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Page 368

Trichilia ornithothera

J.J.de Wilde

Meliaceae Habitat Species occurrence increases significantly in places with rainfall higher than 1500 mm/yr (logistic regression analysis, Chi2 test). It is commonly found in disturbed rainforest (e.g. along roadsides, and patches of secondary vegetation). Very often found in wet places (e.g. along creeks, rivers, lake borders, and even in swampy areas). On sandy, clay, and loamy soils (herbarium).

Phenology Dispersal: probably by animals (Hawthorne & Jongkind 2004) Timing: flowering period from June to September; fruiting period from December to April (De Koning 1983)

Description Guild: np Life form: small to medium-sized tree Max. height: 18 m (herbarium) Max. diameter: 30 cm (herbarium) Leaf: alternate, pinnately compound, subopposite to alternate leaflets, lanceolate, mesophyll (3-5 x 10-23 cm), entire, coriaceous, pubescent on the nerves below; petiole 0.7 cm long, hairy Inflorescence: axillary, branched (panicle, up to 50 cm long) Flower: small (0.5 cm long); calyx brownish; corolla white Fruit: dry dehiscent, globose (1.8 cm in diameter), reddish, pubescent; 4 seeds Seed: large (1.6 x 1 cm), plano-convex, red with an aril Other: it has pubescent twigs, low buttresses, a brown bark, and a creamish slash. It is barely distinct from T. monadelpha but the tufts of hairs emerging from the midrib appears to segregate it. On this basis, however, there are a few anomalous records (e.g. from dry forest even though the main centre in Ghana seems to be wet forest). Invariably counted as T. monadelpha in the Tropical Shelterwood Plot inventories; a tree of similar form, but slightly larger leaves.

Distribution

Uses

Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 16 30’ cells, distribution range is 1052 km, Red List species (Vulnerable) Forest type: wet evergreen forest, secondary forest

The fruits are boiled and used in local medical treatments (herbarium).

Data sources FWTA, De Wilde (1968), De Koning (1983), Hawthorne (1995a), Hawthorne & Jongkind (2004)

T

spp

368

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

T.o.

30

67 (21)

0

47

13

0

40

27

16

7

90

3

40

7

40

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 369

Trichoscypha laxissima

Breteler

Anacardiaceae Description Guild: u Life form: small shrub Max. height: data unavailable Max. diameter: data unavailable Leaf: imparipinnately compound, 5-7 leaflets, notophyll (3 x 10 cm), caudate to acuminate, herbaceous Inflorescence: terminal, branched (lax, manyflowered, nearly glabrous) Flower: male flower small, pink, glabrous, slender stalked; female flower medium-sized, pink Fruit: fleshy, ellipsoid, red Seed: unknown

Distribution Continent: Upper Guinea endemic Upper Guinea: Liberia Distribution type: continuous, very local, present in 1 30’ cells Forest type: coastal (beach) forest (herbarium, only one collection). Only known from one locality near Monrovia (Breteler 2001)

Habitat It is found at altitudes between 1 and 50 m (Breteler in prep.).

Phenology Dispersal: by monkeys

Data sources Breteler (2001), Hawthorne & Jongkind (2004)

T

Environmental table unavailable. There were no sufficient data on the environmental conditions of this single specimen.

369

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Page 370

Triclisia dictyophylla

Diels

Menispermaceae Description

Habitat

Guild: pi Life form: large winding woody climber Max. height: 7 m high, 20 m long (herbarium) Max. diameter: 5 cm (herbarium) Leaf: deltoid with cordate base, macrophyll (up to 20 x 25 cm), herbaceous, pubescent on midrib and main nerves Inflorescence: dioecious, cauliflorous, compound Flower: small; bell-shaped; calyx pale brown; corolla orange, yellow inside, anthers greyish Fruit: fleshy (drupe) (1.7 x 2.5 cm), orange; 1-5 seeds Seed: large (1-1.5 cm), with hard endocarp Other: a climbing shrub when young. It has a lobed stem and the branchlets, petiole and midrib beneath are brown velutinous. The wood on cross-section has ring-patterns.

Generally, found in small gaps and secondary vegetation (Hall & Swaine 1981). In Ghana, reputed in upland evergreen forests, but in Cameroon also along forest edges at the seaside. On sandy soils (herbarium).

Distribution

Hall & Swaine (1981), Hawthorne & Jongkind (2004)

Phenology

Data sources

Continent: Nigeria, Cameroon, Central African Republic, Gabon, Angola, Democratic Republic of Congo (herbarium). Guineo-Congolian wide, from Côte d’Ivoire to the Democratic Republic of Congo (Hall & Swaine 1981) Upper Guinea: Liberia, Côte d’Ivoire, Ghana (herbarium) Distribution type: continental disjunct, widespread, present in 9 30’ cells Forest type: evergreen forest, moist semideciduous forest, secondary forest. In Ghana, it is only found in upland evergreen and moist semideciduous forest (Hall & Swaine 1981).

T

spp

370

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

T.d.

12

33 (6)

0

17

0

8

75

16

0

0

42

58

33

0

58

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 371

Tristemma akeassii

Jacq.-Fél.

Melastomataceae Description

Phenology

Guild: np Life form: erect or creeping herb, rather woody at the base Max. height: 1.5 m (De Koning 1983) Max. diameter: data unavailable Leaf: opposite, simple, elliptic, mesophyll (3.5-9 x 7-16 cm), entire, herbaceous, often dark reddish beneath, with white hairs on both sides; petiole 0.5-1 cm long Inflorescence: terminal, unbranched (1-1.5 cm long) Flower: small; corolla white, pink, violet, or purple with a pale yellow centre Fruit: fleshy, ellipsoid to oblong (0.7 x 1.1 cm), brown; many seeds Seed: small (0.02 mm in diameter) Other: the twigs are quadrangular and hairy.

Data sources FWTA, Jacques-Félix (1976), De Koning (1983)

Distribution Continent: Nigeria (Jacques-Félix 1976) Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana, Togo Distribution type: continuous, widespread, present in 25 30’ cells, distribution range is 1611 km Forest type: montane forest, evergreen forest, secondary forest. It is locally abundant (herbarium).

Habitat Species occurrence increases significantly in places with rainfall higher than 1500 mm/yr (logistic regression analysis, Chi2 test). It can be found in the rainforest understorey, but it is also common in open places (e.g. forest gaps and clearings, along forest roadsides, cultivated land, and coconut plantations). Usually, found in moist places (e.g. dripping cliffs, near creeks, riverbanks, and occasionally even swamps) (herbarium).

T

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

T.a

39

57 (28)

5

44

8

11

46

25

19

5

56

38

31

8

51

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

371

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Page 372

Tristemonanthus nigrisilvae

(N.Hallé) N.Hallé

Celastraceae Description

Phenology

Guild: sb Life form: large irregularly winding woody climber Max. height: 35 m long (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, elliptic, microphyll (2.5-4 x 5-10 cm), often serrate towards the apex, coriaceous Inflorescence: branched Flower: small; calyx pale brown; corolla fleshy, yellow-white Fruit: dry dehiscent, greenish to pinkish, woody Seed: attached to the fruit at the end of a single wing Other: the larger stems are sinuous to deeply fluted.

Timing: flowering period from January to May; young fruits observed in April

Distribution

Data sources

Continent: Upper Guinea endemic Upper Guinea: Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 12 30’ cells, distribution range is 898 km Forest type: wet evergreen forest, coastal forest, secondary forest (herbarium)

Hallé (1958, 1959), De Koning (1983), De Rouw (1991), Hawthorne & Jongkind (2004)

Habitat It is most commonly found where rainfall is between 1500-2500 mm/yr (Chi2 test), in humid places (De Koning 1983). It can grow on large forest trees or in open areas, but is essentially a shade bearer (De Rouw 1991). On sandy-clay to even bare lateritic soils (herbarium).

T

spp

372

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

T.n.

15

21 (14)

7

27

7

0

47

47

7

0

87

13

60

0

40

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 373

Turraea ghanensis

J.B.Hall

Meliaceae Description

Phenology

Guild: sb Life form: small tree Max. height: 6 m (herbarium) Max. diameter: data unavailable Leaf: alternate, simple, ovate to elliptic, entire, herbaceous, pubescent Inflorescence: axillary, unbranched (1-3 flowers) Flower: large; corolla white, densely pubescent Fruit: dry dehiscent (capsule), grooved, thickwalled, splitting halfway open, yellowish, pubescent, with orange septa, 6-8 valves; many seeds Seed: bean-shaped, brown with orange arils

Dispersal: probably by animals (Hawthorne & Jongkind 2004)

Distribution Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire, Ghana. Described as a Ghanaian endemic (Hall 1976). There is one doubtful record from Côte d’Ivoire. Distribution type: continuous, regional, present in 3 30’ cells, distribution range is 592 km Forest type: dry forest. Probably restricted to the dry coastal or similar forest of Upper Guinea. In Ghana known from only a few Southern Marginal forests (Hawthorne 1995a).

Data sources Hall (1976), Hawthorne (1995a), Hawthorne & Jongkind (2004)

Habitat Data unavailable.

T

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

T.g.

7

0 (6)

0

14

0

100

0

0

0

0

29

71

0

29

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

373

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Page 374

Turraea heterophylla

Sm.

Meliaceae Habitat Species occurrence increases in lowlands (logistic regression analysis), where rainfall is less than 1500 mm/yr (Chi2 test). More abundant in dry forest types but it can also be found in rainforests. It can be in the understorey of mature forests as well as in disturbed places such as degraded forests, thickets, and secondary vegetation. Fairly common in secondary forests (Hawthorne & Jongkind 2004). Sometimes found in moist habitats (e.g. near the coast, near lagoons, rivers, or in gallery forests) (herbarium). On clayish soils, and in Ghana, particularly common on soils with pH > 6.2 or higher (Hall & Swaine 1981).

Phenology

Description

Dispersal: probably by animals (Hawthorne & Jongkind 2004) Timing: flowering not clearly seasonal (Hall & Swaine 1981)

Guild: sb Life form: pigmy tree or shrub Max. height: 3 m (herbarium) Max. diameter: data unavailable Leaf: alternate, simple, oblong to obovate, variably lobed, microphyll (1-3.5 x 2.5-9 cm), herbaceous Inflorescence: axillary; solitary or paired Flower: medium-sized; corolla white, funnel-shaped; fragrant Fruit: dry dehiscent (capsule) (1 cm long), green outside, orange inside, 6 valves; 4 seeds Seed: ovoid, medium-sized (0.4-0.5 cm long), glossy black with orange aril Other: a spreading, liana-like shrub.

Distribution wide variety of forest types. In Ghana, it is found only in South Marginal and dry semi-deciduous forests (Hall & Swaine 1981). In Côte d’Ivoire, it has been recorded in rainforest, gallery forest, savanna (forest and cultivated), and secondary forests (herbarium).

Continent: from Sierra Leone to Benin Upper Guinea: Sierra Leone, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 27 30’ cells, distribution range is 1628 km Forest type: it occurs in a

Data sources FWTA, Hall & Swaine (1981), Hawthorne & Jongkind (2004)

T

spp

374

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

T.h.

52

29 (21)

0

31

12

67

16

13

4

13

54

33

31

23

33

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 375

Uapaca chevalieri

Beille

Euphorbiaceae Description

Phenology

Guild: sb Life form: medium-sized tree Max. height: 18 m (FWTA) Max. diameter: data unavailable Leaf: alternate, simple, obovate, mesophyll (4-12 x 7-17 cm), entire, coriaceous Inflorescence: axillary, male flowers in a globose head surrounded by leaf-like bracts, female flowers solitary, dioecious Flower: small Fruit: fleshy, globose (2 x 2.5 cm), speckled, hairy; 3 seeds Seed: large (2 x 1 cm) Other: it has large stilt-roots.

Dispersal: probably by animals

Distribution

Uses

Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire (herbarium, FWTA) Distribution type: Upper Guinea disjunct, regional, present in 4 30’ cells, distribution range is 388 km Forest type: moist evergreen forest, swamp forest, gallery forest

It has edible fruits.

Data sources FWTA, Thoen & Ducousso (1989), Hawthorne & Jongkind (2004)

Habitat Found in swamps and riversides of montane regions (Hawthorne & Jongkind 2004).

U

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

U.c.

12

18 (11)

75

58

0

0

0

75

25

0

25

75

8

17

H 0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

375

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Page 376

Uapaca paludosa

Aubrév. & Léandri

Euphorbiaceae Habitat In swamps in evergreen forests, in lowland regions (Hawthorne & Jongkind 2004).

Regeneration It has a phanerocotylar epigeal foliaceous seedling type (cf. De la Mensbruge 1966).

Phenology Deciduousness: evergreen Dispersal: by animals (monkeys, elephants, civets) Timing: flowering period from December to January; fruiting period from May to November (De Koning 1983)

Description Guild: sb Life form: medium-sized tree Max. height: 25 m (herbarium) Max. diameter: 60 cm (herbarium) Leaf: alternate, simple, obovate, macrophyll (10-25 x 16-50 cm), entire, coriaceous, pubescent beneath; prominent lateral nerves, leaves markedly clustered at branch ends, young leaves are coppercoloured; leafy stipules, up to 3 cm long Inflorescence: axillary, solitary, on stout stalks Flower: large; unisexual; no petals but bracts; corolla yellow Fruit: fleshy, globose (3 cm in diameter), green, softly hairy; 4 seeds Seed: large (1.8 x 1 cm) Other: it has stilt roots. Similar, particularly as a sapling, to U. guineensis. The wood density is 0.76 g/cm3.

Uses

Distribution

It has edible fruits. Continent: Nigeria (FWTA), Cameroon, Gabon (herbarium) Upper Guinea: Liberia, Côte d’Ivoire (herbarium), Ghana Distribution type: continental disjunct, in Upper Guinea present in 3 30’ cells Forest type: wet evergreen forest, swamp forest

Data sources Voorhoeve (1965), De la Mensbruge (1966), De Koning (1983), Vivian & Faure (1985), Keay (1989), Hawthorne (1990, 1995a), Hawthorne & Jongkind (2004)

U

spp

376

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

U.p.

9

0 (4)

11

56

11

0

22

67

0

33

67

0

11

11

44

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 377

Uvaria dinklagei

Engl. & Diels

Annonaceae Description

Phenology

Guild: u Life form: irregularly winding woody climber Max. height: data unavailable Max. diameter: data unavailable Leaf: alternate, simple, oblanceolate, mesophyll (4-7 x 11-17 cm), entire to serrate, hairy underneath Inflorescence: solitary, terminal Flower: medium-sized, yellowish green, 3 large and 3 small petals, hairy Fruit: indehiscent, with ridges; several seeds Seed: data unavailable Other: the twigs are hairy.

Distribution

Uses

Continent: Upper Guinea endemic Upper Guinea: Liberia Distribution type: continuous, very local, present in 1 30’ cell Forest type: coastal shrubland

It has edible fruits.

Data sources FWTA, Hawthorne & Jongkind (2004)

Habitat Data unavailable.

U

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

U.d.

2

100 (1)

0

0

100

0

0

0

100

0

100

0

0

0

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

377

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Uvaria ovata

Page 378

(Dunal) DC.

Annonaceae Phenology Timing: not clearly seasonal (Hall & Swaine 1981)

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana, Togo Distribution type: continuous, widespread, distribution range is 1450 km Forest type: very dry forest, savanna. In Ghana, mainly found in South East Outlier forest, but also present in South Marginal forest (Hall & Swaine 1981). In Côte d’Ivoire, found in savanna and secondary regrowth (herbarium).

Description Guild: np Life form: irregularly winding woody climber Max. height: 4 m long (herbarium) Max. diameter: data unavailable Leaf: alternate, simple, ovate to elliptic to lanceolate, microphyll (1.5-5 x 4-11 cm), entire, coriaceous, pubescent below, aromatic Inflorescence: solitary Flower: medium-sized (1 cm long); calyx with brown hairs; corolla orange to brown, dish-shaped Fruit: fleshy, several per flower in clusters, globose (1 x 1 cm), orange, with 5 hairy carpels; 4-5 seeds Seed: data unavailable Other: the twigs are pubescent.

Data sources FWTA, Hall & Swaine (1981), Aké Assi (1984), Hawthorne & Jongkind (2004)

Habitat It forms dense thickets in dry areas along roadsides, near the coast, and grasslands.

Regeneration It regenerates in shade (Hall & Swaine 1981).

U

spp

378

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

U.o.

34

78 (9)

0

0

6

98

0

3

0

0

82

18

56

9

H 24

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 379

Uvaria sassandrensis

Jongkind

Annonaceae Description

Phenology

Guild: u Life form: large woody climber Max. height: data unavailable Max. diameter: 3 cm (herbarium) Leaf: alternate, simple, entire, coriaceous Inflorescence: solitary Flower: large; calyx pale green; corolla pinkish to orange Fruit: yellow Seed: data unavailable Other: a new species (Jongkind in prep.). Starts as a shrub when young. The branchlets and petioles are red-brown pilose.

Data sources

Distribution

Hawthorne & Jongkind (2004) Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire Distribution type: continuous, local, present in 7 30’ cells, distribution range is 197 km Forest type: secondary forest (herbarium)

Habitat Usually found in the understorey or in secondary forests. Sometimes on riverbanks. On sandy or loamy soils (herbarium).

U

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

U.s.

10

44 (9)

0

40

30

0

80

20

0

0

90

10

60

0

H 40

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

379

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Page 380

Uvariopsis globiflora

Keay

Annonaceae Forest type: in Ghana, it is found mainly in upland evergreen forests and moist semi-deciduous forests, but it can also be present in moist evergreen forests (Hall & Swaine 1981).

Habitat It is found in the forest understorey (herbarium).

Regeneration It regenerates in shade (Hall & Swaine 1981).

Phenology Timing: not clearly seasonal (Hall & Swaine 1981)

Description

Data sources

Guild: sb Life form: small tree Max. height: 6 m (FWTA) Max. diameter: data unavailable Leaf: alternate, simple, oblong to oblongoblanceolate, mesophyll (3.5-8 x 12-23 cm), entire, herbaceous Inflorescence: axillary or cauliflorous, monoecious Flower: medium-sized to large; yellowish with purple centre Fruit: fleshy, obovoid (1.5 x 3 cm), pinkish Seed: large Other: it has an aromatic bark and hairy twigs.

FWTA, Hall & Swaine (1981)

Distribution Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire, Ghana. The record from Côte d’Ivoire is doubtful. Distribution type: present in 2 30’ cells

U

spp

U.g. All

380

n

4 46333

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

0 (4)

0

0

0

25

75

0

0

0

25

75

25

0

50

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 381

Vahadenia caillei

A.Chev.

Apocynaceae Description

Phenology

Guild: np Life form: large winding woody climber Max. height: 40 m long (Haegens 1994) Max. diameter: data unavailable Leaf: opposite, simple, obovate to elliptic, notophyll (1.5-7 x 4.1-14 cm), coriaceous, glabrous Inflorescence: axillary or terminal, branched (1.5-4 cm long, 1-19-flowered) Flower: medium-sized; white, reddish at base; tube with lobes; 5-merous; glabrous on both sides, ciliate at the base; fragrant Fruit: fleshy, globose or ellipsoid (3.8 x 3.8 cm), dark yellow, pulp orange; many seeds Seed: large (1.4 x 0.8 x 0.7 cm) Other: the bark is reddish brown, with many lenticels, and the slash exudes a white latex. It grows with curled tendrils.

Timing: flowering period in May, and from September to November (Haegens 1994)

Distribution

FWTA, Hall et al. (1973), Haegens (1994), Hawthorne (1995a), Hawthorne & Jongkind (2004)

Data sources

Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire (Haegens 1994, herbarium), Ghana (FWTA). Once known only in Atewa within Ghana (Hall et al., 1973), now also known elsewhere (Hawthorne 1995a). Curiously, Haegens (1994) reports no records from Ghana. Distribution type: continuous, widespread, present in 15 30’ cells, distribution range is 1076 km Forest type: high forest, fringe or secondary forest, swamp forest

Habitat Species occurrence increases significantly with rainfall to reach an optimum around 2500 mm/yr (logistic regression analysis, Chi2 test). It is found at altitudes below 500 m (Haegens 1994). It is often found in open forests. On sandy soils (herbarium).

V

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

V.c.

25

60 (5)

44

32

0

0

28

44

28

8

44

48

36

4

H 20

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

381

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Page 382

Vangueriella vanguerioides

(Hiern) Verdc.

Rubiaceae Description

Phenology

Guild: np Life form: woody climber Max. height: 10 m high (herbarium) Max. diameter: 10 cm (herbarium) Leaf: opposite, simple, lanceolate to obovate, notophyll (2-5 x 4.5-11.5 cm), entire, herbaceous, pubescent when young; interpetiolar stipules Inflorescence: axillary, branched (panicle) Flower: small; corolla green outside, lobes inside yellow; petals before anthesis together in a tube; fragrant Fruit: fleshy, ovoid (3 cm in diameter), reddish brown, smooth and hard, 1-2 seeds Seed: black Other: it starts as a shrub. It has opposite paired thorns of 2.5 cm long on the lower part of the stem.

Data sources Verdcourt (1987), Hawthorne & Jongkind (2004)

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana (herbarium, Verdcourt 1987) Distribution type: continuous, widespread, present in 27 30’ cells, distribution range is 1417 km Forest type: montane forest, wet evergreen forest, dry forest, savanna, gallery forest, secondary forest, disturbed forest

Habitat Species occurrence increases with rainfall to reach an optimum around 2800 mm/yr (logistic regression analysis and Chi2 test). In a wide variety of forest types, both in the understorey or in open places. On clayish, loamy, or sandy soils (herbarium).

V

spp

382

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

M

H

Soil WHC L

M

H

V.v.

41

33 (21)

29

44

7

2

37

19

40

0

68

32

39

0

37

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 383

Vepris tabouensis

(Aubrév. & Pellegr.) W.Mziray

Rutaceae Description

Phenology

Guild: u Life form: large tree Max. height: 40 m (Voorhoeve 1965) Max. diameter: 120 cm (Voorhoeve 1965) Leaf: alternate, palmately compound, 4-9 leaflets, obovate to elliptic, mesophyll (3-11 x 5-28 cm), entire, coriaceous, glabrous Inflorescence: axillary, branched (panicle), pubescent Flower: small; calyx pale green, corolla white Fruit: fleshy, globose (1.2 cm in diameter), yellow; 4-5 nuts containing 2 seeds each Seed: medium-sized (0.7 x 0.4 cm), black Other: it has a smooth bark, no buttresses, and wide spreading roots. The wood is very aromatic.

Deciduousness: evergreen (Voorhoeve 1965) Timing: flowering period from December to January; fruiting period from September to October (Voorhoeve 1965)

Distribution Data sources

Continent: Nigeria Upper Guinea: Liberia, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 9 30’ cells, distribution range is 987 km Forest type: wet evergreen forest, moist evergreen forest

Voorhoeve (1965), Mziray (1995)

Habitat Sometimes found close to streams or rivers (herbarium).

V

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

V.t.

12 0 (7)

0

42

0

0

33

25

41

0

100

0

67

0

33

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

H

39

383

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Page 384

Vernonia titanophylla

Brenan

Compositae Distribution Continent: Cameroon, Equatorial Guinea, Gabon, Democratic Republic of Congo (herbarium) Upper Guinea: Guinea Bissau (Moutsambouté 1990), Liberia, Côte d’Ivoire, Ghana (herbarium) Distribution type: continental disjunct, in Upper Guinea present in 4 30’ cells Forest type: sub-montane forest, rainforest, secondary forest (Hawthorne & Jongkind 2004, herbarium)

Habitat It is frequently found along forest edges, roadsides, and in secondary forests (herbarium).

Phenology Dispersal: by wind

Description Guild: pi Life form: small tree Max. height: 10 m (Moutsambouté 1990) Max. diameter: 30 cm (Moutsambouté 1990) Leaf: alternate, simple, elliptic, megaphyll (25-35 x 80-100 cm), dentate, glaucous, almost white on the lower surface; long petioles Inflorescence: disc-shaped (80 x 40 cm, 10-30 flowers crowded together in a head) Flower: small; corolla violet

Data sources

Fruit: dry indehiscent, with long hairs on top Seed: small (0.2 x 0.05 cm) Other: a scarcely branched small tree with whitebrown hairy stems and petioles, and soft wood. Distinguishable from the similar V. conferta by gland-like processes on the petioles.

Moutsambouté (1990), Hawthorne (1995a), Hawthorne & Jongkind (2004)

V

spp

384

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

V.t.

5

75 (4)

60

40

0

0

60

40

0

0

60

40

40

0

20

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 385

Whitfieldia colorata

C.B.Clarke ex Stapf

Acanthaceae Description

Phenology

Guild: np Life form: shrub, sometimes scrambling Max. height: 2 m (herbarium) Max. diameter: data unavailable Leaf: opposite, simple, lanceolate, mesophyll (4.5-11 x 12-27 cm), entire, coriaceous, glabrous Inflorescence: axillary or terminal, not branched Flower: medium-sized (3 cm); calyx rusty orange, corolla tube pink to dark red outside, lobes green to yellow inside Fruit: dry dehiscent (2 cm long), very pale orangered Seed: medium-sized (0.6 x 0.5 x 0.2 cm) Other: it has dark green stems, with some purple above the nodes. It has prominent purple midribs.

Data sources FWTA

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire Distribution type: continuous, widespread, present in 21 30’ cells, distribution range is 862 km Forest type: wet evergreen forest, semi-deciduous forest, savanna, secondary forest

Habitat Species occurrence increases significantly with rainfall (logistic regression analysis). It is most often found at places where rainfall is higher than 2500 mm/yr (Chi2 test). Usually found near rivers (Chi2 test) in dense evergreen forests. Reported also along roads, small forest patches among farms, and secondary forests. On sandy clay soils (herbarium).

W

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

W.c.

40

41 (17)

0

60

15

3

20

10

62

3

80

18

60

3

23

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

385

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Xylia evansii

4:02 PM

Page 386

Hutch.

Leguminosae-Mim. Leaf: bipinnately compound, 2 opposite pinnae, each with 26-32 opposite leaflets, oblong to lanceolate, microphyll (1-2 x 3-7 cm), herbaceous, pubescent below; swollen glands at the top of the petiole and between the upper pairs of leaflets Inflorescence: axillary, in solitary or paired heads, pedunculate Flower: small; corolla yellow Fruit: dry dehiscent, oblong (5 x 20 cm), thick woody; 4 seeds Seed: obovate to elliptic, very large (2 cm long), shining Other: it has buttresses. The foliage is conspicuously clustered at twig ends and large, ascending bows. The wood density is 0.80 g/cm3.

myccorhizae (De la Mensbruge 1966). Saplings are often seen growing strongly towards the light in small to medium-sized gaps, but are not very abundant. The regeneration flourishes in moist disturbed forest, suggesting a greater requirement for gaps than most of the other non-pioneers (Hawthorne 1995a). Seedlings are not abundant, but it has a fast diameter growth rate in Sierra Leone (0.6-1.4 cm/yr, Savill & Fox 1967).

Phenology Deciduousness: trees are sometimes deciduous for a short period. Dispersal: by explosive seed pods. Seeds are eaten by chimpanzees (Kasparek 2000). Timing: flowers are produced towards the end of the dry season (Taylor 1960). Seeds are dispersed towards the end of the dry season (Hawthorne 1995a).

Distribution Continent: Upper Guinea endemic (Lock 1989) Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana (herbarium) Distribution type: continuous, widespread, present in 21 30’ cells, distribution range is 1368 km Forest type: upland evergreen forest, moist evergreen forest, moist semi-deciduous forest, dry semi-deciduous forest, gallery forest. In Ghana, it is found in relatively drier forests (Hawthorne 1995a), mostly in moist semi-deciduous and upland evergreen forests (Hall & Swaine 1981).

Habitat Sometimes found along streams (herbarium).

Data sources

Description

Regeneration

Guild: np Life form: large tree Max. height: 30 m (herbarium) Max. diameter: 150 cm (herbarium)

It has a high level of germination, and the seedlings are characterised as being phanerocotylar epigeal foliaceous (cf. De la Mensbruge 1966). After several weeks the seedlings have ectotrophic

FWTA, Taylor (1960), De la Mensbruge (1966), Savill & Fox (1967), Hall & Swaine (1981), Hawthorne (1985), Lock (1989), Hawthorne (1995a), Kasparek (2000)

X spp

386

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

Soil CMK

>500m

<2km

<5km

D

M

W

VW

L

Soil WHC M

H

L

M

H

X.e.

24

8 (12)

13

25

0

33

46

4

16

4

50

46

42

8

29

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 387

Xylopia elliotii

Engl.

Annonaceae Description

Phenology

Guild: sb Life form: small tree Max. height: 9 m (FWTA, herbarium) Max. diameter: data unavailable Leaf: alternate, simple, lanceolate, microphyll (2-3 x 5-9 cm), entire, pubescent beneath Inflorescence: axillary, solitary Flower: medium-sized (approx. 2.5 cm long); white to yellowish; sweet scented Fruit: data unavailable Seed: data unavailable Other: it has hairy twigs.

Data sources FWTA, Savill & Fox (1967), Hawthorne (1995a), IUCN Red List (2000)

Distribution Continent: Upper Guinea endemic Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire Distribution type: continuous, widespread, present in 8 30’ cells, distribution range is 886 km, Red List species (Vulnerable) Forest type: wet evergreen forest, riverine forest (herbarium). In Sierra Leone, it has been recorded in fringing forest (Savill & Fox 1967).

Habitat It occurs in the forest understorey.

X spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

X.e.

9

0 (2)

11

44

0

11

44

44

0

0

56

44

44

11

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

387

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Xylopia villosa

Page 388

Chipp

Annonaceae Habitat Smaller trees are relatively shaded, while taller trees are exposed (Hawthorne 1995a). The species occurs in a wide rainfall range, but has its optimum between 1500-2500 mm/yr (Chi2 test). It prefers infertile soils (Swaine 1996)

Regeneration It regenerates in shade (Hall & Swaine 1981). Regeneration has been found to be obviously depleted in burnt forest (Hawthorne 1994).

Phenology Deciduousness: evergreen Timing: flowering period from November to April (De Koning 1983); fruiting period all year (De Koning 1983)

Description Guild: sb Life form: medium-sized tree Max. height: 30 m (herbarium) Max. diameter: 30 cm (herbarium) Leaf: alternate, simple, elliptic to ovate, microphyll (2-3.5 x 5-10 cm), entire, coriaceous, pubescent below Inflorescence: solitary or in small groups Flower: large (petals approx. 2.5 cm long); light yellow; funnel-shaped Fruit: fleshy (1.9 x 1.1 x 0.7), green outside and pink-red inside Seed: large, greenish grey Other: the bark is whitish and aromatic with lenticels. It has pubescent twigs and aerial roots of up to 30 cm high.

Distribution Continent: Nigeria, Cameroon Upper Guinea: Sierra Leone, Côte d’Ivoire, Ghana Distribution type: continuous, widespread, present in 14 30’ cells, distribution range is 1904 km Forest type: rainforest, gallery forest, and old secondary forest (herbarium). In Ghana, it is most abundant in wet and moist evergreen forests, and it is also found in moist semi-deciduous forests. Occasionally in dry semi-deciduous forests and in upland forests (Hall & Swaine 1981).

Data sources Taylor (1960), Hall & Swaine (1981), De Koning (1983), Hawthorne (1994, 1995a), Swaine (1996)

X spp

X.v. All

388

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

28

14 (14)

0

61

0

11

43

43

4

4

71

25

25

0

71

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

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Page 389

Zanthoxylum psammophilum

(Aké Assi) Waterman

Rutaceae Description

Phenology

Guild: pi Life form: woody climber Max. height: 15 m (Aké Assi 1960) Max. diameter: 2 cm (Aké Assi 1960) Leaf: imparipinnately compound, 30-40 cm long, 7-15 alternate leaflets, ovate to oblong, notophyll (4-6 x 6-13 cm), glabrous, shiny below; petiole and rachis not so spiny Inflorescence: dioecious; terminal, branched (panicle), male inflorescence 21-30 cm long; female inflorescence 3-5 cm long Flower: male and female flowers small (2.5 mm long), 5-merous, white Fruit: dry dehiscent (capsule), globose (approx. 0.5 cm in diameter); 1 seed Seed: medium-sized (0.4 cm in diameter); black or bluish Other: Zanthoxylum spp. are easily recognised from their bark and slash alone. The bark is covered with broad-based woody prickles. The slash is yelloworange, fibrous, and with a characteristic very fruityacidic taste.

Data sources Aké Assi (1960), IUCN Red List (2000), Hawthorne & Jongkind (2004)

Distribution Continent: Upper Guinea endemic Upper Guinea: Côte d’Ivoire (herbarium, Aké Assi 1960) Distribution type: continuous, very local, present in 2 30’ cells, distribution range is 87 km, Red List species (Endangered) Forest type: coastal forest (Aké Assi 1960)

Habitat Data unavailable.

spp

n

Open (n)

Altitude

River

Coast

Mean Annual Rainfall

>500m

<2km

<5km

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

Z.p.

2

100 (1)

0

0

0

0

100

0

0

100

0

0

0

100

0

All

46333

37

4

39

7

37

29

24

10

6

69

25

36

13

39

Z 389

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Page 390

10

Page 391

A. Siepel, L. Poorter and W.D. Hawthorne

Ecological profiles of large timber species

1 0

2:11 PM

H A P T E R

11/21/03

C

22466 layout page 391-393

Introduction

Large tree species fulfil an ecologically and economically important role in West African forests. Because of their sheer size, they determine to a large extent forest structure and functioning. From a commercial point of view, there is a longstanding interest in forest exploitation in West Africa, not in the least because of the proximity to European markets. The timber trade began as early as 1816, with the shipment of Khaya logs to London (Harcourt et al. 1992, Parren & de Graaf 1995). From a regional point of view, timber exploitation really took off in the 1950s to peak in the 1970s in Côte d’Ivoire (Chatelain et al., chapter 2), and a little bit later in the neighbouring countries. Especially in Sierra Leone and Côte d’Ivoire commercial species have been overexploited or forests have disappeared (Harcourt et al. 1992, Parren & de Graaf 1995, Chatelain et al., chapter 2). There is a strong awareness that a more sustainable forest management is needed, be it through forest plantations, recuperation of degraded areas, line plantation or assistance of natural regeneration (Bongers et al., chapter 8, Parren & de Graaf 1995). For an effective management information is needed on the distribution, ecology, and growth requirements of these large timber species. Over the past century, such information has gradually been accumulated by the foresters and botanists active in the region (Aubréville 1959, Taylor 1960, Voorhoeve 1965, Hall & Swaine 1981, Hawthorne 1995a, Kasparek 2000). In this chapter we present ecological profiles for 56 large timber species. We have made an effort to summarise the existing knowledge on these species, and to give an idea of their environmental requirements by combining data on species abundance with GIS layers on the environment. In the following pages, we describe the metadata behind the ecological profiles. First we present information on species selection and abundance, then on the environmental data and the analyses used, and finally we describe the format of the ecological profiles.

Figure 10.1 A large tree of a large timber species; an Entandrophragma candollei of 7m dbh in Forêt Classée Haut Sassandra in Côte d’Ivoire.

Species data Species selection The focus of this chapter is on the large commercial tree species of Sierra Leone, Liberia, Côte d’Ivoire and Ghana. These countries contain most of the forest in Upper Guinea and are relatively well studied. To compare the different areas we selected 56 species that were inventoried in all countries. For some genera the species are pooled because in some countries the species were identified to genus only (this is the case for Afzelia, Berlinia, Celtis, Entandrophragma, Erythrophleum, Khaya, Milicia, Parinari). In addition some species were added that occur in one country and are absent in the others (for instance Tetraberlinia tubmaniana is only found in Liberia). Most of the species have maximum sizes of over 40m in height and over 100cm in diameter. The 56 selected species represent the most important current and potential timber species in West Africa. Most of these species were used to make the forest map of Upper Guinea (Bongers et al., chapter 4). Synecological information on these species can therefore be found in chapter 4.

391

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Page 392

Chapter 10. Ecological profiles of large timber species

Species abundance In total, species abundance data were collected for 171 forest sites; 3 for Sierra Leone, 26 for Liberia, 37 for Côte d’Ivoire and 105 for Ghana. For some species data were available for 5 additional sites in Sierra Leone. The data were collected by various organisations. For Sierra Leone data were used from Small (1952), Savill & Fox (1967) and Davies (1987). For Liberia data were used from the German Forestry Mission to Liberia (GFML 1967a, 1976b, Sachtler & Hamer 1967a, 1967b and Sachtler 1968) and the Liberian Forest Service. For Côte d’Ivoire we drew largely on data collected by SODEFOR (Clément & Guinaudeau 1973, SODEFOR 1978, 1979) and for Ghana the national forest inventory data were used (Wong 1989, Hawthorne 1995a). In Liberia and Côte d’Ivoire only a number of large timber tree species have been surveyed (in each country c. 60 species), whereas in Ghana all species with a diameter at breast height (dbh) over 30 cm were taken into account. The inventoried area of each site is variable and ranges from 10 to 4500 ha (Appendix 2). Inventories vary from complete inventories to strip inventories. The 171 sites cover a wide range in environmental conditions (Table 4.2). For the abundance data the number of trees > 30cm dbh per square km was used. In Ghana all trees > 30cm dbh were inventoried and data are available on individual trees. All Liberian and Sierra Leonean sites had data for trees over 40 cm dbh and for Côte d’Ivoire various lower limits were available (> 10 cm, > 15 cm, > 20 cm, > 40 cm and > 60 cm). To be able to compare all sites we estimated for each of the sites the number of trees > 30 cm dbh per km2. Transformation values were based on the Ghanaian data. For the Ghanaian data, for each species (thus the pooled data for all sites together) frequency distribution diagrams for size classes were constructed. Such a frequency distribution shows an idealised population structure. Most species then show a negative exponential pattern with size. For each species a negative exponential curve was fitted on the frequencies by size class. The regression was used to estimate numbers of trees > 30cm for each of the sites in Sierra Leone, Liberia and Côte d’Ivoire. This assumes that in the sites studied the frequency distributions would be similar to the idealised population structure. For the large sample sizes this is a reasonable supposition.

Environmental data Species distributions are strongly shaped by water availability (rainfall, soil water holding capacity), soil fertility (expressed as sum of exchangeable cations, CMK; Ca2+, Mg2+, K+) and altitude. Maps of these environmental variables were prepared and included in a Geographical

392

Information System (ArcView, ESRI 1995). In the introduction to chapter 9 a more extensive description is given of the metadata behind these maps.

Species responses to the environment A multiple regression was carried out to model species abundance as a function of four environmental factors; altitude, annual rainfall, soil water holding capacity, and available cations. A stepwise forward regression was used, with the simple and quadratic variables, but without including interaction terms. A significant simple variable indicates an increasing or decreasing probability to find the species with an increase in the environmental variable. A significant quadratic variable indicates that the species shows a bell-shaped response curve towards that environmental variable (Jongman et al. 1987).

Explanation of the species descriptions In the following section of the atlas ecological profiles of the species are presented, covering one species per page. Each species account includes a distribution map, a photo and/or drawing, and an environmental table. First a short botanical description of the species is given. The aim of the description is to give the reader an impression of the species. For a more detailed, taxonomical description we refer the reader to the Flora of West Tropical Africa (Keay 1954, 1958a, 1963, Hepper 1968, 1972), the newly published The Woody Plants of Western African Forests (Hawthorne & Jongkind 2004) or other publications (Aubréville 1959, Taylor 1960, Voorhoeve 1965, Savill & Fox 1967, Keay 1989). The botanical description is followed by information on its chorology and its rarity. The environmental requirements of the species are described in terms of light, water and nutrient availability. Finally, regeneration, growth characteristics and main uses of the species are discussed. Each description is accompanied by a table with a quantitative summary of the environmental requirements and the most relevant literature references. For the ecological profiles we drew largely on information given by Voorhoeve (1965) and Hawthorne (1995a). For a detailed description of the format of the species accounts, see the introduction to the ecological profiles of the rare and endemic species (Holmgren et al., chapter 9). The ecological profiles of the large timber species deviate from those of the rare and endemic species at some points. For example, the wood density is given if available,

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Chapter 10. Ecological profiles of large timber species

and there is an extra guild “sw” for species that grow in swamps. The description of countries of occurrence and forest types is less extensive, and largely based on Voorhoeve (1965). The habitat requirements are based on the literature, and inferred from the multiple regression analyses (see above). The description of regeneration characteristics is more extensive, and includes information on growth rates. For only a few species histograms are presented on the reproductive phenology as in general few herbarium collections were available for large species.

Species distribution maps Information on species distribution is only presented in one distribution map. For some genera the species were identified to genus only (e.g., Celtis, Entandrophragma, Erythrophleum, Khaya, Milicia). For these genera only one map is available, representing the pooled abundance of the species. In some cases this results in an indistinct pattern, because both dry and wet sister taxa are pooled. For example, Khaya grandifoliola occurs mainly in dry semideciduous forest whereas Khaya ivorensis occurs mainly in the wetter forest types. The species abundance, expressed as number of stems larger than 30 cm dbh per km2, is given for each of the 171 forest sites. The size of the symbols indicates the log-transformed abundance of the species. A large symbol indicates a high abundance, a small symbol a low abundance or absence. Based on these data an interpolation map was made, using the inverse distance weighting interpolation method in ArcView, and taking the 12 nearest forest sites into account. The interpolated abundance is shown in 8 classes; 0-1, 1-10, 10-20, 20-40, 40-80, 80-160, > 160 individuals per

km2. Interpolation was only done for the potential forest zone of Upper Guinea (see chapter 9). One should realise that these maps are likely to present only the potential distribution of the species, as the data are largely based on pre-logging forest inventories carried out in the 1960s and 1970s. The exception is Ghana, were inventories have been carried out in the 1990s. In reality, much of the forest cover, and many large individuals have disappeared in the meanwhile, because of forest exploitation or forest conversion (see also Chatelain et al., chapter 2).

Environmental table The environmental table provides a quantitative summary of the environmental requirements of the species. It indicates in what percentage of the forests a species if found, where the species is present, and if the species is found above 400 m altitude. The rainfall gradient is divided in 4 classes; Dry (< 1500), Medium (1500-2000), Wet (2000-2500) and Very Wet (> 2500 mm/yr). The total available cations (CMK) of the soil is divided in the classes Low (0-4), Medium (4-8) and high (> 8 cmolc/kg) and the Water Holding Capacity (WHC) of the soil is divided in Low (< 50), Medium (50-85) and High (> 85mm water/m soil). Information on the environmental conditions where the species is found, is only interesting when comparing it to the full range of environments in which all forests are found. To this end information on the focal species is presented in the first line, and the environmental conditions of all 198 forest sites (the 171 analysed plus 27 additional ones) included in the analysis are presented in the second line.

393

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Page 394

Key to symbols in maps and tables of chapter 10 border sea

Predicted species abundance

(# stems >30 cm diameter at breast height/km2)

0-1 1 - 10 10 - 20 20 - 40 40 - 80 80 - 160 > 160 area outside potential forest zone

Observed species abundance

(# stems >30 cm diameter at breast height/km2)

0 0,5 1,25 6,25 25 100 400 > 2240

Environmental tables Spp n Altitude

species number of forests in which the species is found % of records found at altitudes > 400 m

Mean annual rainfall

Soil CMK

Soil WHC

394

% of records found in (D) relatively dry areas: < 1500 mm/yr, (M) intermediate areas: 1500-2000 mm/yr, (W) wet areas: 2000-2500 mm/yr, and (VW) very wet areas: > 2500 mm/yr % of records found on soils with (L) low availability of cation: 0-4 cmolc/kg, (M) intermediate availability of cation: 4-8 cmolc/kg, and (H) high availability of cation: > 8 cmolc/kg % of records found on soils with (L) low water holding capacities: < 50 mm water/m soil, (M) intermediate water holding capacities: 50-85 mm water/m soil, and (H) high water holding capacities: > 85 mm water/m soil

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Page 395

Albizia ferruginea

(Guill. & Perr.) Benth.

Leguminosae-Mim. A

Description Guild: np Life form: large tree Max. height: 40 m (Voorhoeve 1965) Max. diameter: 132 cm (inventory data Ghana) Leaf: alternate, bipinnately compound, 8-12 opposite pinnae, each with 12-26 opposite leaflets, oblong, nanophyll (0.5-1 x 1.2-2.5 cm), entire, pubescent beneath, especially on the midrib Inflorescence: axillary, simple dichasia Flower: small; corolla yellow, tube-shaped Fruit: dry dehiscent, flat (5 x 20 cm), papery, sparsely pubescent and red when young, glabrous and yellow-brown when ripe; approx. 10 seeds Seed: flat, medium-sized (0.7 x 0.9 cm) Other: a spreading tree, sometimes with two sizes of leaflets conspicuous in the crown. It has nitrogenfixing root nodules and narrow, thick buttresses of up to 1.5 m. The slash exudes a clear or honeycoloured, sticky gum. The leaf flush is red. Wood density is 0.56 g/cm3.

Phenology

Distribution

Deciduousness: deciduous (Voorhoeve 1965) Dispersal: by wind (Voorhoeve 1965) Timing: flowering period from February to March; fruiting period from December to February (Voorhoeve 1965)

Continent: Benin to Angola (Voorhoeve 1965) Upper Guinea: Senegal to Togo (Voorhoeve 1965) Forest type: moist evergreen forest, moist semideciduous forest, dry semi-deciduous forest (Hall & Swaine 1981). A Red List species (Vulnerable).

Uses

Habitat

It is a timber species, exploited relatively heavily (Hawthorne 1995a).

It is a strong light demander (Voorhoeve 1979).

Regeneration Germination is rapid (Taylor 1960). There is no difference between germination in the light and in the dark (Kyereh et al. 1993). Germination is equally

successful in large gaps (Kyereh 1994). It has a cryptocotylar epigeal reserve seedling type (cf. Voorhoeve 1965). Seedlings are in some cases abundant at the start of the rainy season but soon die out in the shade. Growth is rapid in exposed areas (2-3 m in 5 years, Taylor 1960).

Data sources Taylor (1960), Voorhoeve (1965), Hall & Swaine (1981), Kyereh et al. (1993), Kyereh (1994), Hawthorne (1995a), IUCN Red List (2000)

Table data are not available.

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Alstonia boonei

Page 396

De Wild.

Apocynaceae A

Description

Regeneration

Guild: pi Life form: large tree Max. height: 45 m (Voorhoeve 1965) Max. diameter: 140 cm (De Jong 1979) Leaf: whorled, simple, obovate, mesophyll (3-6 x 8-18 cm), entire, coriaceous, glabrous; petiole up to 3 cm long Inflorescence: terminal, branched with 2-3 tiers of pseudo-umbels Flower: small; calyx pale green, pubescent; corolla yellowish green to pale yellow, tube-shaped, pubescent outside and on the lobes inside Fruit: dry dehiscent (2 on 1 stalk) (0.3 x 40 cm long), pubescent, green; many seeds Seed: flat, large (0.5 x 0.2 x 0.03 cm), brown, with 1 cm long hairs on both ends Other: it has a Terminalia-like, strongly layered crown and whorled leaves. The base of larger trees is usually deeply fluted. The slash exudes a white latex. It has creeping horizontal roots, covered in large lenticels. Wood density is 0.35 g/cm3.

Germination is apparently normal, although seedlings are rapidly restricted to medium-sized to large gaps. It has a phanerocotylar epigeal foliaceous seedling type (cf. Voorhoeve 1965). Even young seedlings are not found in deep shade (Hall & Swaine 1981). Saplings are marked light demanders. One of several “strong light-demanders” favoured by canopy tree poisoning in Uganda (Philip 1967).

Distribution Continent: Benin to the Democratic Republic of Congo, Uganda, Sudan (Voorhoeve 1965) Upper Guinea: Senegal to Togo (Voorhoeve 1965) Forest type: moist evergreen forest, upland evergreen forest, moist semi-deciduous forest, dry semi-deciduous forest, secondary forest (Voorhoeve 1965, Hall & Swaine 1981).

Habitat It is a light demander, but when young shadebearing. The abundance decreases with rainfall, and increases significantly with soil water holding capacity (regression analysis). It is most common in wet or marshy places (Voorhoeve 1965).

Spp

396

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

A.b.

151

7

27

58

7

7

29

61

10

21

2

72

All sites

198

4

26

57

9

8

31

58

11

25

2

72

Growth Height increments of 2 m in the first year, with trees up to 15 m tall and 25 cm dbh after 10 years, are recorded by Taylor (1960).

Phenology Deciduousness: deciduous (Voorhoeve 1965) Dispersal: by wind, potentially over long distances in strong winds (Hawthorne 1995a) Timing: flowering period from November to January (Taylor 1960); fruiting period from January to February (Voorhoeve 1965)

Data sources Taylor (1960), Philip (1967), Voorhoeve (1965), De Jong (1979), Hall & Swaine (1981), Hawthorne (1995a)

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Page 397

Amphimas pterocarpoides

Harms

Leguminosae-Pap. A

Description

Regeneration

Guild: np Life form: large tree Max. height: 50 m (Voorhoeve 1965) Max. diameter: 120 cm (Voorhoeve 1965) Leaf: in tufts at the end of the branches, imparipinnately compound, 13-19 opposite leaflets, lower leaflets ovate to broadly or narrowly elliptic, upper leaflets (narrowly) elliptic to oblong or obovate, microphyll (2-4 x 4.5-12 cm), entire, sparsely pubescent, glabrescent Inflorescence: terminal, not branched (20 cm long) Flower: small to medium-sized; calyx densely brown puberulous; corolla cream-coloured Fruit: dry dehiscent, thin, flat, pendulous (4.8 x 17 cm), golden-brown; 1-2 seeds Seed: flattened, reniform, large (0.8 x 2 cm), dark brown Other: a spreading, rough-barked tree, with dense, dark foliage. It has neat, triangular buttresses. A blood-red, sticky sap slowly exudes from the slash wound. Wood density is 0.79 g/cm3.

Germination is normal. It has a phanerocotylar epigeal reserve seedling type (cf. Voorhoeve 1965). Seedlings occur in the shade (Hall & Swaine 1981) although they do not develop far. Young saplings, with rather Entandrophragma-like crowns are often seen, however, only under small gaps (Hawthorne 1995a). Later, the tree is said to be a lightdemander (Taylor 1960).

Phenology Deciduousness: deciduous in October, November (Voorhoeve 1965) Dispersal: by wind (Voorhoeve 1965) Timing: flowering period from October to November; fruiting period from January to February (Voorhoeve 1965)

Distribution Continent: Benin to Cameroon, Congo (Brazzaville), Sudan Upper Guinea: Guinea to Togo Forest type: wet evergreen forest, moist evergreen forest, moist semi-deciduous forest, dry semideciduous forest (Hall & Swaine 1981)

Habitat

Data sources

It is a light demander. Regeneration is most common in the low bush and on open places (Voorhoeve 1965). It has a preference for high rainfall areas (Swaine 1996) and its abundance decreases significantly with soil fertility (regression analysis, cf. Swaine 1996)

Taylor (1960), Voorhoeve (1965), Hall & Swaine (1981), Hawthorne (1995a), Swaine (1996)

Spp

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

A.p.

153

7

24

59

9

8

31

59

10

23

2

70

All sites

198

4

26

57

9

8

31

58

11

25

2

72

397

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Page 398

Anopyxis klaineana

(Pierre) Engl.

Rhizophoraceae A

Description

Regeneration

Guild: np Life form: large tree Max. height: 45 m (Voorhoeve 1965) Max. diameter: 120 cm (Voorhoeve 1965) Leaf: simple, in whorls of 3, oblong, elliptic or obovate, notophyll (3-5.5 x 7-13 cm), more or less decurrent, coriaceous, dark green and glossy above Inflorescence: axillary, branched (dichasia) Flower: small; calyx greenish; corolla brown Fruit: dry dehiscent (2 x 3.3 cm), greenish, with 5 valves; max. 10 seeds Seed: flattened, winged, medium-sized (0.4 x 0.8 cm), wing 0.8 x 2 cm, medium brown Other: a large, straight cylindrical tree, usually without buttresses and with a dense and rounded crown.

It has a phanerocotylar epigeal foliaceous seedling type (cf. Voorhoeve 1965). The seeds have very poor viability and natural regeneration is low for this reason (Taylor 1960). Seedlings that do occur are said to grow in the open or only in light shade (Hall & Swaine 1981).

Phenology Deciduousness: evergreen Dispersal: by wind Timing: flowering period from August to October; fruiting period from February to March (Voorhoeve 1965)

Distribution Continent: Benin to Congo (Voorhoeve 1965) Upper Guinea: Sierra Leone to Togo (Voorhoeve 1965) Forest type: evergreen forest, dry semi-deciduous forest (Hall & Swaine 1981). A Red List species (Vulnerable).

Habitat The abundance increases significantly with rainfall (regression analysis). It has been found in rather wet valleys as well as on steep rocky hill sites (Voorhoeve 1965).

Uses A timber species. The bark is used for treating skin infections and ulcers.

Data sources Taylor (1960), Voorhoeve (1965), Hall & Swaine (1981), Hawthorne (1995a), IUCN Red List (2000)

Spp

398

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

A.k.

103

9

15

59

13

14

42

49

10

35

1

57

All sites

198

4

26

57

9

8

31

58

11

25

2

72

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Page 399

Anthonotha fragrans

(Baker f.) Exell & Hillcoat

Leguminosae-Caes. A

Description

Regeneration

Guild: np Life form: medium-sized to large tree Max. height: 38 m (Voorhoeve 1965) Max. diameter: 120 cm (Voorhoeve 1965) Leaf: alternate, paripinnately compound, elliptic, mesophyll (2-11 x 4-20 cm), leaves of saplings larger (up to 12 x 30 cm), entire, coriaceous, glabrous above, pubescent beneath, glossy darkgreen above, dark-brown or golden-brown beneath Inflorescence: axillary, not branched (10-20 cm long) Flower: small Fruit: dry dehiscent (3 x 5 x 10 cm), thick coriaceous, dark brown, pubescent; 3 seeds Seed: very large (approx. 2.5 cm across), flattened Other: it has root ridges that may develop into narrow buttresses. The crown is reddish or rusty brown from below. It is ectomycorrhizal.

It has a cryptocotylar hypogeal reserve seedling type (cf. Voorhoeve 1965).

Phenology Deciduousness: deciduous (Voorhoeve 1965) Dispersal: no obvious means of dispersal (Hawthorne 1995a) Timing: flowering period in December; fruiting period in April (Taylor 1960)

Distribution Continent: Benin to Congo (Voorhoeve 1965) Upper Guinea: Sierra Leone to Togo (Voorhoeve 1965) Forest type: wet evergreen forest, moist evergreen forest, upland evergreen forest, moist semideciduous forest (Hall & Swaine 1981)

Habitat Anthonotha is more abundant in areas with a high rainfall, low soil water holding capacity and a low soil fertility (regression analysis). It is particularly found on acid soils. Outside evergreen forest it is most commonly seen in low-lying situations (Hawthorne 1995a).

Data sources Taylor (1960), Voorhoeve (1965), Hall & Swaine (1981), Alexander (1989), Hawthorne (1995a)

Spp

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

A.f.

63

11

11

49

19

21

63

30

6

57

2

37

All sites

198

4

26

57

9

8

31

58

11

25

2

72

399

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Page 400

Antiaris toxicaria

(Rumph. ex Pers.) Leschen.

Moraceae A

Regeneration Germination is normal. Seedlings are usually very abundant near parent trees. Height classes up to about 30-40 cm are common under high shade, but exposure to the sun is required for further growth. Seedlings experience heavy mortality in the first year and are intolerate to dense climbers and shrubs. Nevertheless, the tree is common in secondary forest.

Description

Growth

Guild: np Life form: large tree Max. height: 51 m (Taylor 1960) Max. diameter: 130 cm (Taylor 1960) Leaf: alternate, simple, broadly elliptic to ovate, notophyll (3-9 x 4-13 cm), entire, coriaceous; leaves of young shoots and saplings densely hairy or scabrous, more narrowly elliptic, and dentate Inflorescence: dioecious, at base of short lateral twigs, male flowers in a puberulous peduncle (0.81.2 cm), female flowers solitary on a pedicel Flower: small; corolla greenish Fruit: fleshy (1 x 1.5 cm), velvety, scarlet; 1 seed Seed: round or ellipsoid Other: a tree with a cylindrical, pale bole, sometimes with buttresses. The bark has numerous large, rounded lenticels, often in vertical rows. The slash exudes a watery, creamy latex, turning brown. Wood density is 0.47 g/cm3.

It can grow rapidly under exposed conditions; 50 cm or more per year on old farms, but only half this in the less favourable conditions of Tropical Shelterwood plots (Taylor 1960). Diameter growth is positively correlated with crown size.

Phenology semi-deciduous forest, dry semi-deciduous forest, secondary forest. A very variable species, found from the wettest to the driest forest types. This range of variants was previously split into two species. Nowadays, it is considered that intermediate forms are so typical that the variation pattern is best described as one of (interbreeding) varieties within a single species (Berg 1977, 1978).

It is a strong light demander (Taylor 1960) and common in secondary forest (Hall & Okali 1979). The abundance decreases significantly with rainfall (regression analysis). In the wet forests of Liberia it prefers therefore well-drained sites (Voorhoeve 1965). It is not selective of soils (Taylor 1960, Swaine 1996).

Continent: West and Central Africa (Hawthorne 1995a) Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana (herbarium) Forest type: upland evergreen forest, wet evergreen forest, moist evergreen forest, moist

Spp

400

n

Uses A timber tree (Hawthorne 1995a).

Habitat

Distribution

Deciduousness: deciduous between November and February (Taylor 1960) Dispersal: by birds and bats (Osmaston 1965) Timing: flowering period from November to February (Taylor 1960); fruiting period from February to March (Voorhoeve 1965), December to April (Taylor 1960)

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

A.t.

159

7

27

60

9

3

27

60

13

20

3

71

All sites

198

4

26

57

9

8

31

58

11

25

2

72

Data sources Taylor (1960), Osmaston (1965), Voorhoeve (196), Berg (1977, 1978), Hall & Okali (1979), Hall & Swaine (1981), Hawthorne (1995a), Swaine (1996)

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Page 401

Berlinia confusa

Holy

Leguminosae-Caes. Description Guild: sb Life form: large tree Max. height: 40 m (Voorhoeve 1965) Max. diameter: > 100 cm (Voorhoeve 1965) Leaf: alternate, paripinnately compound, leaflets ovate to elliptic or obovate, notophyll (2-7 x 5-15 cm), entire, coriaceous Inflorescence: axillary or terminal, branched (a panicle or broad raceme, up to 16 cm long) Flower: medium-sized; calyx pale green; corolla white Fruit: dry dehiscent, flat (9 x 33 cm); up to 6 seeds Seed: disc-shaped, round to elliptic, very large (4 x 3 x 0.6 cm), smooth, pale brown Other: a tree with a rather untidy crown and slightly twisted bole. The slash is brownish, with a red to brownish exudate at length and a distinctive scent. Wood density is 0.7 g/cm3.

B

Distribution Continent: Benin to Gabon (Voorhoeve 1965) Upper Guinea: Sierra Leone to Togo (Voorhoeve 1965) Forest type: wet evergreen forest, moist evergreen forest, moist semi-deciduous forest, dry semideciduous forest, secondary forest (Voorhoeve 1965, Hall & Swaine 1981)

Habitat As a group, Berlinia species prefer lowland areas, and their abundance decreases significantly with soil fertility (regression analysis). B. confusa seems to prefer well-drained sites (Voorhoeve 1965). It is found scattered in all except the driest forests, but even there probably mainly along rivers (De Koning 1983).

Uses It is locally used as timber (Voorhoeve 1965).

Regeneration

Data sources

It has a phanerocotylar epigeal reserve seedling type (cf. Voorhoeve 1965). Poor regeneration has been noted in some contexts (e.g. Voorhoeve 1965), possibly due to insect predation (De Koning 1983). Saplings are mostly found in secondary forest adjacent to the high forest (Voorhoeve 1965).

Taylor (1960), Voorhoeve (1965), Hall & Swaine (1981), De Koning (1983), Hawthorne (1995a)

Phenology Deciduousness: evergreen (Voorhoeve 1965) Dispersal: seeds explosively ejected from the pod (Voorhoeve 1965) Timing: flowering period from January to April (Voorhoeve 1965), March to May (Taylor 1960); fruiting period from July to September (Voorhoeve 1965), September to December (Taylor 1960)

Spp

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

B.spp.

140

5

21

62

9

8

36

56

9

25

2

H 68

All sites

198

4

26

57

9

8

31

58

11

25

2

72

401

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Page 402

Canarium schweinfurthii

Engl.

Burseraceae m in the first year) in exposed sites. One to two metre tall saplings are most typical as scattered individuals along tracks and in medium-sized gaps.

Growth

C

Trees of 6 m height were found on 4 year old logging tracks (Hawthorne 1993). Trees can attain 55 cm dbh in 40 year old secondary forest (Kahn 1982).

Phenology Description Guild: np Life form: large tree Max. height: 50 m (Voorhoeve 1965) Max. diameter: 150 cm (Voorhoeve 1965) Leaf: alternate, imparipinnately compound (9-13 leaflets), ovate to oblong, mesophyll (3-5 x 7-30 cm), entire, pubescent Inflorescence: axillary, not branched, flowers unisexual Flower: medium-sized; both male and female flowers creamy white Fruit: fleshy, ellipsoid to obovoid (1.5 x 3 cm), purplish; 1 seed Seed: fusiform, very large (1.2 x 2.5 cm), oily fruit pulp Other: it has a spreading crown and cylindrical bole, without buttresses. The leaves are clustered at the ends of loopy twigs. It has heavy root ridges, up to 40 cm thick and 90 cm high. The slash exudes a resin. The leaf flush is red. Wood density is 0.48 g/cm3.

forest, moist semi-deciduous forest, dry semideciduous forest, gallery forest (Voorhoeve 1965, Hall & Swaine 1981). In Ghana, the tree is most common in evergreen forest, and rare in the dry fire zone and MS-NW forest zones (Hawthorne 1995a).

Habitat It is a strong light demander (Taylor 1960), and prefers well-drained slopes (Voorhoeve 1965). Its abundance increases sharply with rainfall and shows an optimum around 2500 mm/yr (regression analysis).

Regeneration Germination is normal or slightly slow, providing the fibrous endocarp is kept moist (Taylor 1960). The seeds lie dormant until the beginning of the next dry season (Hawthorne 1995a). The seeds germinate most successfully in the shade (Gilbert 1952). It has a phanerocotylar epigeal foliaceous seedling type (cf. Voorhoeve 1965). The seedlings at the cotyledon stage are sometimes seen in understorey shade, and young plants are seen occasionally in small gaps, but seedlings only grow well (up to 1.5

Distribution Continent: Benin to Ethiopia, Tanzania, Angola (Voorhoeve 1965) Upper Guinea: Guinea to Togo (Voorhoeve 1965) Forest type: wet evergreen forest, moist evergreen

Spp

402

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

C.s.

128

8

17

61

12

10

38

52

9

30

2

63

All sites

198

4

26

57

9

8

31

58

11

25

2

72

Deciduousness: deciduous (Voorhoeve 1965) Dispersal: by mammals and birds (Hawthorne 1995a) Timing: flowering period from February to May; fruiting period from August to December (Voorhoeve 1965). Flowers are produced after the new leaves, and fruits appear 5 months later (Taylor 1960). Remains of fruits have been found in archaeological sites, indicating a history of dispersal by humans (J.Maley pers. comm.).

Uses It is a potential timber species with edible fruits and locally, the inner bark is said to be a cure for leprosy. It has been recommended as one of the most useful trees for reforestation of savanna in the Democratic Republic of Congo. It is sometimes cultivated for its fruits of which an edible oil can be obtained (Abbiw 1990).

Data sources Gilbert (1952), Taylor (1960), Voorhoeve (1965), Hall & Swaine (1981), Kahn (1982), Keay (1989), Abbiw (1990), Hawthorne (1993, 1995a), Maley (pers. comm.)

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Page 403

Ceiba pentandra

(L.) Gaertn.

Bombacaceae Description

Habitat

Guild: pi Life form: large tree Max. height: 60 m (Voorhoeve 1965) Max. diameter: 200 cm (Voorhoeve 1965) Leaf: alternate, palmately compound, 5-9 leaflets, obovate to elliptic, mesophyll (3-6 x 11-18 cm), entire or sometimes slightly dentate, glabrous, fringe of hairs at the top when young, glossy dark green above, lighter beneath Inflorescence: axillary, solitary Flower: medium-sized; calyx green; corolla white to creamy yellow Fruit: dry dehiscent, elliptic (6 x 20 cm), green; several seeds Seed: ovoid, medium-sized (0.5 cm in diameter), black, embedded in grey or white kapok Other: a very common, grey-barked tree, developing large buttresses. The architecture has a strong differentiation between lateral and vertical shoots. Cuttings from the latter tend to grow more horizontally even when independent. Buttresses develop partly in response to prevailing wind and crown asymmetry, e.g. on the windward size of the bole. The bark and young stems are often with thorns. Wood density is 0.3 g/cm3.

It is an extreme light demander (Voorhoeve 1965). The abundance is highest in the driest regions, decreases sharply with rainfall, and increases slightly with altitude (regression analysis). Although it does not grow in freshwater swamps, it will grow along their margins. It is almost indifferent to soil conditions (Taylor 1960).

Distribution Continent: in all tropics (South America, Asia, Africa) Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana (herbarium). Suggestions that the species is introduced to Africa from Central America are rendered less likely by the fact that Ceiba pollen more than 10,000 years old have been found in sediments in Lake Bosumtwi (Maley, pers. comm.). Forest type: moist evergreen forest, upland evergreen forest, moist semi-deciduous forest, dry semi-deciduous forest, secondary forest (Hall & Swaine 1981)

C

Regeneration Germination is apparently normal. Kyereh et al. (1993) reported no difference between the percentage of germination in the light and in the dark, and it is equally successful in large gaps (Kyereh 1994). It has a phanerocotylar epigeal foliaceous seedling type (cf. Voorhoeve 1965). Seedlings are abundant soon after dispersal (12 days, Taylor 1960), and it is obvious that either seeds do not germinate, or die rapidly, in shaded areas (e.g. 2% sunlight, Swaine pers. comm). It is found in light places, especially where the soil has been disturbed (logging roads, old farms etc.). It has a high photosynthetic light compensation point, quantum efficiency and stomatal conductance (Riddoch et al. 1991).

Timing: flowering period from December to January; fruiting period from February to March (Voorhoeve 1965). The flowers are bat-pollinated (Harris & Baker 1959, Baker & Harris 1959), but also visited by birds (Toledo 1977). In Panama, only half the adult trees flowered in one year, although few produced viable seed due to insect attack, population outcrossing was 0.69 (Murawski & Hamrick 1992).

Uses The kapok is used for stuffing pillows and mattresses. It has been widely planted in Asia for food, fodder and fibre. A popular agroforestry species (Sekar et al. 1990).

Growth Seedling growth is maximal between 30 and 40% irradiance (Swaine et al. 1997). In medium-sized to large gaps growth is very rapid (2 m/yr; Taylor 1960). Trees of 30 cm dbh and 20 m height were recorded on 4 year old logging roads (Hawthorne 1993).

Data sources Haigh (1941), Harris & Baker (1959), Baker & Harris (1959), Taylor (1960), Voorhoeve (1965), Henwood & Baker (1973), Toledo (1977), Hall & Swaine (1981), Sekar et al. (1990), Riddoch et al. (1991), Murawski & Hamrick (1992), Hawthorne (1993), Kyereh et al. (1993), Kyereh (1994), Maley (pers. comm.), Swaine et al. (1997).

Phenology Deciduousness: deciduous (Voorhoeve 1965) Dispersal: by wind (Voorhoeve 1965)

Spp

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

C.p.

167

7

27

58

9

6

30

58

12

22

2

69

All sites

198

4

26

57

9

8

31

58

11

25

2

72

403

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Page 404

Celtis adolfi-friderici

Engl.

Ulmaceae Description Guild: pi Life form: large tree Max. height: 50 m (Taylor 1960) Max. diameter: 100 cm (Taylor 1960) Leaf: alternate, simple, broadly elliptic, mesophyll (5.5-8 x 8-16 cm), entire, glabrous Inflorescence: axillary, panicle Flower: small, corolla white Fruit: fleshy, subglobose (2 cm long); 1 seed Seed: data unavailable Other: a slender tree with narrow, high buttresses and a rounded, dark crown. The boughs are horizontal and drooping at the tips.

C

improves performance only slightly (Taylor 1960). It has a phanerocotylar epigeal foliaceous seedling type (cf. De la Mensbruge 1966). Although Hall & Swaine (1981) record that the species regenerates in the shade, seedlings are most typical in small to medium-sized gaps, particularly when > 50 cm tall, and seem more intolerant of shade than the other Celtis spp.

Growth In the Tropical Shelterwood Sample plots, the height growth amounts to 40 cm in the first year, and 20-100 cm per year afterwards (Taylor 1960).

Distribution

Phenology

Continent: Benin to Democratic Republic of Congo, Uganda (Hall & Swaine 1981) Upper Guinea: Côte d’Ivoire, Ghana, Togo (Hall & Swaine 1981) Forest type: moist evergreen forest, moist semideciduous forest, dry semi-deciduous forest (Hall & Swaine 1981). It is common in Ghana (Hall & Swaine 1981).

Deciduousness: it is partly deciduous in December to February, but the crown is never bare (Taylor 1960) Dispersal: probably mainly by birds Timing: flowering period from May to June and November to December; fruiting period from October to November and March to April

Habitat It is a light demander, but capable of sustaining high shade in early youth. It prefers lighter well-drained soils (Taylor 1960). Celtis spp. attain the highest abundance in the dry forest. Their abundance decreases sharply with rainfall, and increases with soil fertility and altitude (regression analysis).

Regeneration Germination is said to be rather erratic, with low viability unless soaked and exposed to sun, which

Spp

404

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

C.spp.

154

6

29

58

8

5

28

60

12

23

3

69

All sites

198

4

26

57

9

8

31

58

11

25

2

72

Data sources FWTA, Taylor (1960), De la Mensbruge (1966), Hall & Swaine (1981), Hawthorne (1995a)

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Page 405

Celtis mildbraedii

Engl.

Ulmaceae Description

Regeneration

Guild: sb Life form: large tree Max. height: 54 m (Taylor 1960) Max. diameter: 90 cm (Taylor 1960) Leaf: alternate, simple, elliptic to ovate, notophyll (1.5-7 x 3.5-14 cm), serrate in the upper two-thirds but almost entire on the older trees, coriaceous, glabrous Inflorescence: axillary, cymose Flower: very small; apetalous Fruit: fleshy, ovoid (1.3 cm long), red; 1 seed Seed: data unavailable Other: a steep-buttressed tree with a cylindrical bole. The crown is small and compact, with horizontal, thin boughs. Wood density is 0.7 g/cm3.

Germination is rapid (within 2 weeks, De la Mensbruge 1966). Under controlled conditions the percentage of germination is similar in the light and in the dark (Kyereh et al. 1993), although germination is strongly depressed in large gaps (Keyereh et al. 1993). It has a phanerocotylar epigeal foliaceous seedling type (cf. De la Mensbruge 1966). Seedlings are common, and locally very abundant near mother trees. Established saplings thrive in medium-sized to large gaps.

Distribution Continent: Benin to Angola, Tanzania (Hall & Swaine 1981) Upper Guinea: Côte d’Ivoire, Ghana, Togo (Hall & Swaine 1981) Forest type: moist semi-deciduous forest, dry semi-deciduous forest, secondary forest. It is very common in Ghana (Hall & Swaine 1981).

Habitat It tolerates high shade in youth but is essentially a light demander. Celtis spp. attain as a group the highest abundance in the dry forest. Their abundance decreases sharply with rainfall, and increases with soil fertility and altitude (regression analysis). In contrast, Swaine (1996) found that Celtis mildbraedii did not have any preference for annual rainfall or soil fertility conditions. It is very tolerant of soils and avoids swampy places (Taylor 1960).

C

Growth In Tropical Shelterwood System plots, height increments of 1-3 m, in 4 years, have been observed (Taylor 1960).

Phenology Deciduousness: evergreen Dispersal: by birds, particularly hornbills (Taylor 1960) Timing: flowering period from February to April and June to August (Taylor 1960); fruiting period from March to April and September to December (Taylor 1960), possibly at other times of the year

Data sources Taylor (1960), De la Mensbruge (1966), Hall & Swaine (1981), Kyereh et al. (1993), Kyereh (1994), Hawthorne (1995a), Swaine (1996), Swaine et al. (1997)

Spp

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

C.spp.

154

6

29

58

8

5

28

60

12

23

3

69

All sites

198

4

26

57

9

8

31

58

11

25

2

72

405

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Page 406

Chrysophyllum giganteum

A.Chev.

Sapotaceae

C

Description

Phenology

Guild: sb Life form: medium-sized tree Max. height: data unavailable Max. diameter: data unavailable Leaf: alternate, compound, 7-10 pairs of leaflets, oblong to obovate, mesophyll (3-7 x 8-20 cm), entire, coriaceous, densely covered with golden to silvery hairs beneath Inflorescence: axillary, raceme (3 cm long) Flower: small; corolla white Fruit: fleshy, globose (4.8 cm in diameter), orange Seed: very large Other: a buttressed, evergreen forest tree with dense, discolorous crown. The slash exudes a white latex.

Deciduousness: evergreen Dispersal: by animals

Distribution

Uses

Continent: Benin to Nigeria (Hall & Swaine 1981) Upper Guinea: Sierra Leone to Togo (Hall & Swaine 1981) Forest type: moist semi-deciduous forest, dry semi-deciduous forest (Savill & Fox 1967, Hall & Swaine 1981). It is common in Ghana (Hall & Swaine 1981).

It has edible fruits.

Regeneration It has a phanerocotylar epigeal reserve seedling type (cf. De la Mensbruge 1966).

Table data are not available.

406

Data sources FWTA, Taylor (1960), De la Mensbruge (1966), Savill & Fox (1967), Hall & Swaine (1981), Hawthorne (1990)

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Page 407

Daniellia ogea

(Harms) Rolfe ex Holland

Leguminosae-Caes. Description

Regeneration

Guild: pi Life form: large tree Max. height: 45 m (Voorhoeve 1965) Max. diameter: 120 cm (Voorhoeve 1965) Leaf: alternate, paripinnately compound, 11-19 leaflets, obovate to oblong, microphyll (1-3.5 x 3-9 cm), entire; leaves and leaflets of saplings and water shoots much larger Inflorescence: axillary or terminal, branched (panicle) Flower: medium-sized; corolla lilac to pinkish purple Fruit: dry dehiscent, flat (3.5 x 7.5 cm), glabrous, stiff-papery; 1 seed Seed: very large (1.5 x 2.8 cm), with a propellerlike valve Other: a tree with no buttresses and occasionally wide-spreading surface roots. It has a strong taproot. Ripple marks are obvious in the sapwood. The new leaves flush red. Wood density is 0.5 g/cm3.

It has a phanerocotylar epigeal reserve seedling type (cf. Voorhoeve 1965). Regeneration in the high forest is not common. The tree is mature at an early age, fruit bearing trees of 10 m height have been observed (Hawthorne 1995a).

D

Phenology Deciduousness: deciduous (Voorhoeve 1965) Dispersal: by wind (Voorhoeve 1965) Timing: flowering period from October to January; fruiting period from January to March (Voorhoeve 1965)

Distribution Continent: Benin to Gabon (Voorhoeve 1965) Upper Guinea: Senegal to Togo (Voorhoeve 1965) Forest type: wet evergreen forest, moist evergreen forest, moist semi-deciduous forest, secondary forest (Voorhoeve 1965, Hall & Swaine 1981). It is common in Ghana (Hall & Swaine 1981).

Data sources

Habitat

Taylor (1960), Voorhoeve (1965), Hall & Swaine (1981), Hawthorne (1995a)

The tree is a light demander and it is often found in the neighbourhood of water in marshy valleys and on rocky slopes (Voorhoeve 1965). Daniellia spp. are found as a group over a wide range of rainfall conditions, and attain their maximal abundance around 2300 mm/yr. Their abundance decreases sharply with soil fertility (regression analysis).

Spp

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

D.spp

134

7

21

58

11

10

37

55

7

27

1

67

All sites

198

4

26

57

9

8

31

58

11

25

2

72

407

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Page 408

Daniellia thurifera

Benn.

Leguminosae-Caes.

D

Description

Regeneration

Guild: pi Life form: large tree Max. height: 45 m (Voorhoeve 1965) Max. diameter: 120 cm (Voorhoeve 1965) Leaf: alternate, paripinnately compound, 11-19 leaflets, oblong-lanceolate to lanceolate, microphyll (2.5-3.5 x 6-9 cm), entire, glabrous Inflorescence: panicle Flower: medium-sized; corolla red; fragrant Fruit: dry dehiscent, elliptic to ovate (7 cm long); 1 seed Seed: very large Other: it has conspicuous ripple marks in the sapwood. Very similar to D. ogea. It has bark with large, horizontal lenticels. The leaves are bright red when flushing. Wood density is 0.55 g/cm3.

It has a phanerocotylar epigeal reserve seedling type (De Koning 1983). It regenerates in shade and small gaps (Hall & Swaine 1981).

Phenology Deciduousness: deciduous in October to November (Savill & Fox 1967) Dispersal: by wind (Voorhoeve 1965) Timing: flowering period from October to December (De Koning 1983); fruiting period from March to April (Savill & Fox 1967

Distribution Continent: Upper Guinea endemic (Hall & Swaine 1981) Upper Guinea: Guinea Bissau, Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana (Hall & Swaine 1981) Forest type: wet evergreen forest, moist evergreen forest, moist semi-deciduous forest, secondary forest (Savill & Fox 1967, Hall & Swaine 1981)

Habitat

A timber species (Savill & Fox 1967).

Daniellia spp. are found as a group over a wide range of rainfall conditions, and attain their maximal abundance around 2300 mm/yr. Their abundance decreases sharply with soil fertility (regression analysis).

Spp

408

n

Uses

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

D.spp

134

7

21

58

11

10

37

55

7

27

1

67

All sires

198

4

26

57

9

8

31

58

11

25

2

72

Data sources FWTA, Taylor (1960), Savill & Fox (1967), Voorhoeve (1965), Hall & Swaine (1981), De Koning (1983)

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Page 409

Distemonanthus benthamianus

Baill.

Leguminosae-Caes. Description Guild: np Life form: medium-sized to large tree Max. height: 36 m (Voorhoeve 1965) Max. diameter: 97 cm (inventory data Ghana) Leaf: pinnately compound, 7-10 alternate leaflets, ovate to elliptic, notophyll (2.5-5 x 5-10 cm) Inflorescence: terminal, branched (cymose dichasium) Flower: medium-sized; calyx reddish brown; corolla white Fruit: dry indehiscent (3 x 10 cm), papery, pale brown; 2-3 seeds Seed: elliptic, flattened, medium-sized (0.5 x 1 cm), pale brown, glossy, with a yellowish margin Other: the “devil-tree” with distinctive reddish bark especially when it is exposed to the sun. It is buttressed, with regular, domed crown of very slightly blue-green foliage, although the flush is copper-coloured. Wood density is 0.7 g/cm3.

D

Distribution Continent: Benin to Gabon (Voorhoeve 1965) Upper Guinea: Sierra Leone to Togo (Voorhoeve 1965) Forest type: wet evergreen forest, moist evergreen forest, moist semi-deciduous forest, dry semideciduous forest, secondary forest (Voorhoeve 1965, Hall & Swaine 1981). It is more common in moist semi-deciduous forest than in the evergreen forest (Voorhoeve 1965).

Habitat The species shows an optimum at intermediate rainfall conditions (2000 mm/yr), and its abundance declines sharply beyond 2500 mm/yr (regression analysis). It grows scattered, rarely in groups, and does not grow in swamps.

Regeneration Little is known of the germination requirements, although Taylor (1960) notes a high percentage of viable seeds. It has a phanerocotylar epigeal reserve seedling type (cf. Voorhoeve 1965). Seedlings and saplings are often seen in the understorey, and seem to be shade tolerant even when a metre or more in height. They are found in very disturbed forest, but regeneration seems never to be abundant in either situation.

Timing: flowering period from January to February; fruiting period in October (Voorhoeve 1965). Fruiting is said to be sparse and not regularly annual (Taylor 1960).

Uses A timber species. It has numerous applications in native medicine (Voorhoeve 1965).

Phenology

Data sources

Deciduousness: irregularly deciduous (Voorhoeve 1965) Dispersal: by wind

Spp

n

Taylor (1960), Voorhoeve (1965), Hall & Swaine (1981), Hawthorne (1995a)

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

D.b

139

8

25

61

9

4

32

58

10

23

2

71

All sites

198

4

26

57

9

8

31

58

11

25

2

72

409

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Page 410

Entandrophragma angolense

(Welw.) DC.

Meliaceae Growth as in E. utile. The crown is usually not as large as the huge bole might suggest. Wood density is 0.56 g/cm3.

Distribution Continent: Uganda, Angola Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana (herbarium) Forest type: upland evergreen forest, moist evergreen forest, moist semi-deciduous forest, dry semi-deciduous forest. A Red List species (Vulnerable).

E

Habitat Entandrophragma spp. attain as a group a maximal abundance at intermediate rainfall conditions (1800 mm/yr). Their abundance declines strongly above 2300 mm/yr. Their abundance is positively related to soil fertility and soil water holding capacity (regression analysis). Alexandre (1982b) refers to E. angolense as the most shade-bearing of the Entandrophragmas. Common, particularly in better-drained sites.

Description

Regeneration

Guild: np Life form: large tree Max. height: > 50 m (Voorhoeve 1965) Max. diameter: > 180 cm (Voorhoeve 1965) Leaf: pinnately compound, 14-22 opposite leaflets, elliptic, oblong or obovate, microphyll (2-4 x 3.5-12 cm), entire, coriaceous, leaves and leaflets of saplings and young trees larger Inflorescence: a densely flowered panicle Flower: small, corolla greenish white Fruit: capsule, pendulous (4 x 18 cm), woody, nearly black; 25-30 seeds Seed: approx. 7 cm long including papery wing, red brown Other: a tree with rather “fat”, high buttresses. It has compound clustered leaves, but not as strongly

Spp

410

n

In plantations, seedlings grow approx. 1 m per year in the first two years (Taylor 1960). In tropical Shelterwood plots they grow as much as 1-2 m/yr, once they are a metre or so tall (Taylor 1960). Taylor (1960) also records 1 m striplings reaching 6 m (10 cm dbh) in 4 years after planting in Taungya. In Nigeria, the diameter increment is slower than other species in the genus, requiring an estimated 137 years to reach 1 m dbh (Keay 1961). In Uganda, increment was found to be positively correlated with crown size, as for other species investigated, although the crown position is poorly correlated with increment (Dawkins 1956). Growth rings in this species are likely to be annual (Detienne & Mariaux 1977).

Phenology Deciduousness: deciduous in September to November (Voorhoeve 1965) Dispersal: by wind (Voorhoeve 1965) Timing: flowering period concurs with the new leaves in December (Taylor 1960); fruiting period throughout the wet season from July to September

Uses

Germination is normal. It has a cryptocotylar epigeal reserve seedling type (cf. De la Mensbruge 1966). Seedlings require shade for the first few years. Subsequently, they require greater exposure to sun for at least part of the day. Saplings are commonly seen in gaps of all sizes, their narrow crown on little-branched stems usually held above any mass of climbers and other competition typical of gaps. The presence of parent trees in the area noticeably increases the number of seedlings, suggesting that dispersal of seeds is a serious limiting factor. However, very sparse regeneration of this species was found in Côte d’Ivoire in a 3 ha subsample of a plot containing several adults (De Klerk 1991).

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

E.spp.

159

7

28

61

9

3

28

62

10

20

2

74

All sites

198

4

26

57

9

8

31

58

11

25

2

72

A timber tree.

Data sources Dawkins (1956), Taylor (1960), Keay (1961), Voorhoeve (1965), De la Mensbruge (1966), Detienne & Mariaux (1977), Hall & Swaine (1981), Alexandre (1982b), De Klerk (1991), IUCN Red List (2000)

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Page 411

Entandrophragma candollei

Harms

Meliaceae Description

Regeneration

Guild: np Life form: large tree Max. height: > 45 m (Voorhoeve 1965) Max. diameter: 180 cm (Voorhoeve 1965) Leaf: paripinnately compound, 10-20 opposite leaflets, elliptic, oblong or obovate, notophyll (2.5-6 x 5-16 cm), entire to undulate Inflorescence: axillary, branched (panicle) Flower: small Fruit: dry dehiscent, fusiform (4 x 20 cm), opening from the top with 5 recurving valves, greyish brown; 15-50 seeds Seed: including the wing 8-12 cm, pale yellowish brown Other: a cylindrical or slightly buttressed tree, with a widely-spreading crown. Wood density is 0.68 g/cm3.

It has a cryptocotylar epigeal reserve seedling type (cf. De la Mensbruge 1966). Regeneration requirements appear to be very similar to the other species. No regeneration of this species was found in Côte d’Ivoire in a 3 ha subsample of a plot containing several adults (De Klerk 1991).

Distribution Continent: Benin to Congo (Voorhoeve 1965) Upper Guinea: Guinea to Togo (Voorhoeve 1965) Forest type: upland evergreen forest, moist evergreen forest, moist semi-deciduous forest (Hall & Swaine 1981). It is common in Ghana (Hall & Swaine 1981). A Red List species (Vulnerable).

Habitat Entandrophragma spp. attain as a group a maximal abundance at intermediate rainfall conditions (1800 mm/yr). Their abundance declines strongly above 2300 mm/yr. Their abundance is positively related to soil fertility and soil water holding capacity (regression analysis).

Growth

E

Growth rings in this species are likely to be annual (Detienne & Mariaux 1977).

Phenology Deciduousness: deciduous (Voorhoeve 1965) Dispersal: by wind (Voorhoeve 1965) Timing: flowering period from November to December; fruiting period from May to August (Voorhoeve 1965)

Uses A timber species.

Data sources Taylor (1960), Voorhoeve (1965), De la Mensbruge (1966), Detienne & Mariaux (1977), Hall & Swaine (1981), De Klerk (1991), IUCN Red List (2000)

Spp

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

E.spp.

159

7

28

61

9

3

28

62

10

20

2

74

All sites

198

4

26

57

9

8

31

58

11

25

2

72

411

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Page 412

Entandrophragma cylindricum

(Sprague) Sprague

Meliaceae alternate leaflets, ovate, elliptic or oblong, notophyll (2-4.5 x 4-15 cm), entire, coriaceous, glossy dark green above Inflorescence: branched (lax panicle) Flower: small; yellow-green; sparsely puberulous Fruit: dry dehiscent (8 cm long), purplish black, opening at the base and the top with 5 recurrent valves; 5-20 seeds Seed: 5-8 cm long including wing, pale brown Other: a tree with a very scented slash and with no or small buttresses. The crown is rounded, not especially spreading. Wood density is 0.66 g/cm3.

E

Distribution Continent: Benin to Uganda, Congo (Voorhoeve 1965) Upper Guinea: Guinea to Togo (Voorhoeve 1965) Forest type: upland evergreen forest, moist evergreen forest, moist semi-deciduous forest, dry semi-deciduous forest. A Red List species (Vulnerable).

Habitat Entandrophragma spp. attain as a group a maximal abundance at intermediate rainfall conditions (1800 mm/yr). Their abundance declines strongly above 2300 mm/yr. Their abundance is positively related to soil fertility and soil water holding capacity (regression analysis).

Regeneration

Description

Spp

412

n

Growth Taylor (1960) notes that, in his experimental areas (probably Bobiri), the species was slower growing than others of this genus, reaching only 1 m after 4 years in silviculturally treated forest. Keay (1961) predicted 87 years to reach 77 cm dbh. Growth rings are likely to be annual (Detienne & Mariaux 1977).

Phenology Deciduousness: deciduous (Voorhoeve 1965) Dispersal: by wind (Voorhoeve 1965) Timing: flowering period: from February to March (Voorhoeve 1965); fruiting period from May to August (Taylor 1960)

Uses A common timber tree, heavily exploited (Hawthorne 1995a).

Data sources

It has a cryptocotylar epigeal reserve seedling type (cf. De la Mensbruge 1966). Similar to E. angolense, but with a tendency to produce fewer seedlings each of which grows more slowly (20-40 cm a year in height), even in later years (Taylor 1960). It requires some shade for germination, with

Guild: np Life form: large tree Max. height: > 50 m (Voorhoeve 1965) Max. diameter: > 180 cm (Voorhoeve 1965) Leaf: paripinnately compound, 10-14 opposite or

increased light levels desirable later in life (Pieters 1958). Seedlings grew best at 50% full sunlight in an experiment in the Democratic Republic of Congo. Seedling growth is slow because the roots develop slowly (Sawyerr 1960). No regeneration of this species was found in Côte d’Ivoire in a 3 ha subsample of a plot containing several adults (De Klerk 1991).

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

E.spp.

159

7

28

61

9

3

28

62

10

20

2

74

All sites

198

4

26

57

9

8

31

58

11

25

2

72

Pieters (1958), Sawyerr (1960), Taylor (1960), Keay (1961), Voorhoeve (1965), De la Mensbruge (1966), Detienne & Mariaux (1977), Hall & Swaine (1981), De Klerk (1991), Hawthorne (1995a), IUCN Red List (2000)

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Page 413

Entandrophragma utile

(Dawe & Sprague ) Sprague

Meliaceae Description Guild: np Life form: large tree Max. height: 60 m (Voorhoeve 1965) Max. diameter: 250 cm (Voorhoeve 1965) Leaf: pinnately compound, 16-26 opposite or alternate leaflets, ovate to oblong, notophyll (2-5.5 x 5-12 cm), entire, thinly coriaceous, glossy, medium green above Inflorescence: a long and slender panicle Flower: small; corolla white Fruit: dry dehiscent, club-shaped (6 x 23 cm), opening from the top with 5 hardly recurving straight valves, black with brown warty lenticels, woody; 2030 seeds Seed: (8-10 cm long) including the wing, medium to dark brown Other: a tree with very clustered leaves at the tips of stout twigs. It has heavy buttresses, up to 3 m. Wood density is 0.62 g/cm3.

Distribution Continent: Benin to Uganda, Angola (Voorhoeve 1965) Upper Guinea: Guinea to Togo (Voorhoeve 1965) Forest type: upland evergreen forest, moist semideciduous forest, dry semi-deciduous forest. A Red List species (Vulnerable).

Habitat Entandrophragma spp. attain as a group a maximal abundance at intermediate rainfall conditions (1800 mm/yr). Their abundance declines strongly above 2300 mm/yr. Their abundance is positively related to soil fertility and soil water holding capacity (regression analysis). Alexandre (1982b) refers to E. utile as the most light-demanding of the Entandrophragmas. It seems to be more droughttolerant than others in the genus (Abu Juam Musah,

pers. comm.). Avoids marshy sites, well-drained, deep soils seem to be preferred (Voorhoeve 1965).

Regeneration There is little difference between germination in the light and in the dark (Kyereh et al. 1993). Germination is strongly depressed in large gaps (Kyereh et al. 1993). Soaking followed by a dry period did not allow germination, whereas repeated soaking, signalling more reliably the onset of the rainy season, did (Synnot 1975). Seed viability was found to be adequate to carry the seeds from dispersal to the onset of the rains. It has a cryptocotylar epigeal reserve seedling type (cf. De la Mensbruge 1966). Synott (1975) records high mortality, and seasonal variation due to rodent and antelope predation, with mortality at 1.44% per day for the first 100 days, and a slight decline thereafter. Insect damage, particularly by the shoot-borer Hypsipyla, was also high. In areas where predation was least, mortality due to drought and disease was high. After 2.5 years only 1.3% of planting seedlings survived for these reasons. The seedlings may be more inclined to suffer from sudden changes in exposure to the sun than other species (Taylor 1960). The seedlings are physiologically well-suited to the deep shade of the forest floor, and the healthiest seedlings were under at least some shade (Synnot 1975). Unshaded seedlings died. Nevertheless, in natural forest, light availability was found to be the biggest limitation on seedling growth. Seedling growth is maximal around 25% irradiance (Swaine et al. 1997). Root competition was not considered an important limiting factor, nor were nutrient levels in the forest.

E (Sawyerr 1960). Seedlings, up to 40 cm tall are locally abundant, both in the shade and in slightly disturbed patches. However, it tends to suffer higher mortality and is a slower grower than E. angolense, growing about 1 m or less in 4 years in Tropical Shelterwood System plots (Taylor 1960). It can reach about 1.5 m in height in 4 years in silviculturally treated forest (Taylor 1960). Growth rings are likely to be annual (Detienne & Mariaux 1977).

Phenology Deciduousness: deciduous (Voorhoeve 1965) Dispersal: by wind (Voorhoeve 1965) Timing: flowering period from January to February (Voorhoeve 1965); fruiting period from December to March

Data sources Sawyerr (1960), Taylor (1960), Voorhoeve (1965), De la Mensbruge (1966), Synott (1975), Detienne & Mariaux (1977), Hall & Swaine (1981), Alexandre (1982b), De Klerk (1991), Kyereh et al. (1993), Kyereh (1994), Hawthorne (1995a), Swaine et al. (1997), IUCN Red List (2000).

Growth Seedling growth is slow because the roots develop slowly. Seedlings become infested with mites and other insects in nurseries, unless they are shaded

Spp

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

E.spp.

159

7

28

61

9

3

28

62

10

20

2

74

All sites

198

4

26

57

9

8

31

58

11

25

2

72

413

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Page 414

Erythrophleum ivorense

A.Chev.

Leguminosae-Caes.

E

Description

Regeneration

Guild: np Life form: large tree Max. height: 40 m (Voorhoeve 1965) Max. diameter: 120 cm (Voorhoeve 1965) Leaf: bipinnately compound, 4-8 opposite pinnae with 8-14 alternate leaflets, elliptic, microphyll (1.5-4 x 2.5-8.5 cm), glossy dark green Inflorescence: axillary or terminal, not branched, densely flowered Flower: small; calyx yellowish green; corolla yellow Fruit: dry dehiscent (4 x 7.5 cm), thick-coriaceous, greyish black, 2-6 seeds Seed: large (0.5 x 0.9 x 1.3 cm), shiny black Other: a rather twisted, unbuttressed and lowbranched tree. Sometimes far spreading surface roots are present. The bark has corky, 2-10 mm wide, horizontal lenticels. The slash of young trees is purplish pink-brown, with some milky sap in the wound. The slash of old trees is red-brown with a purplish gleam, exuding a red, sticky sap. Wood density is 0.87 g/cm3.

It has a phanerocotylar epigeal foliaceous seedling type (cv. Voorhoeve 1965). There is little regeneration (Taylor 1960).

Phenology Deciduousness: evergreen (Voorhoeve 1965) Dispersal: by wind (Voorhoeve 1965) Timing: flowering period from May to September (Voorhoeve 1965)

Distribution Continent: Benin to Gabon (Voorhoeve 1965) Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana, Togo (Voorhoeve 1965) Forest type: wet evergreen forest, moist evergreen forest, moist semi-deciduous forest, high forest, secondary forest

The abundance of Erythrophleum spp. as a group declines with soil fertility (regression analysis).

414

n

The wood is very hard and is used for railway sleepers and bridging timber (Voorhoeve 1965).

Data sources

Habitat

Spp

Uses

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

E.spp.

121

9

27

49

12

12

42

46

12

34

2

57

All sites

198

4

26

57

9

8

31

58

11

25

2

72

Taylor (1960), Voorhoeve (1965), Hall & Swaine (1981), Lock (1989), Hawthorne (1995a)

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Page 415

Erythrophleum suaveolens

(Guill. & Perr.) Brenan

Leguminosae-Caes. Description

Regeneration

Guild: np Life form: medium-sized tree Max. height: > 25 m (Taylor 1960) Max. diameter: 90 cm (Taylor 1960) Leaf: alternate, bipinnately compound, 4-8 opposite pinnae each with 7-12 leaflets, ovate to elliptic, microphyll (2.5-6.5 x 2.5-8 cm) Inflorescence: terminal, branched (panicle) Flower: small; corolla yellow to white Fruit: dry dehiscent (3 x 11 cm); 5-10 seeds Seed: large (0.8 x 1.3 cm) Other: a widely-spreading tree, with foliage pendulous at the ends of the sinuous branches in a rounded or spherical, slightly irregular crown. The flush of new leaves is red. It has no buttresses. It has nitrogen-fixing nodules.

Germination is irregular, and growth is slow (Taylor 1960).

Phenology Deciduousness: deciduous (Taylor 1960) Dispersal: no obvious means of dispersal Timing: flowering period from July to September; fruiting period from March to June (Taylor 1960)

E

Distribution Continent: widespread in Africa, Asia (Lock 1989) Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana, Togo Forest type: moist semi-deciduous forest, woodland savanna, riverine forest (Hawthorne 1995a).

Uses

Habitat

It is well-known because of its poisonous bark used in trials of ordeal (Hawthorne 1995a).

The abundance of Erythrophleum spp. as a group declines with soil fertility (regression analysis). In parts of Togo, it dominates patches of 25 year old fallow vegetation (Guelly et al. 1993). In the Democratic Republic of Congo, it comes to dominate forest in a Brachystegia woodland area with a regime of early-burning or complete fire protection (Schmitz 1952).

Data sources Schmitz (1952), Taylor (1960), Halliday & Nakao (1982), Lock (1989), Guelly et al. (1993), Hawthorne (1995a)

Spp

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

E.spp.

121

9

27

49

12

12

42

46

12

34

2

57

All sites

198

4

26

57

9

8

31

58

11

25

2

72

415

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Page 416

Funtumia africana

(Benth.) Stapf

Apocynaceae Inflorescence: axillary, branched (cyme, 3-40 flowers) Flower: medium-sized; calyx green; corolla yellowgreen, tube-shaped; fragrant Fruit: dry dehiscent, fusiform (3 x 20 cm), brown to black; many seeds Seed: elliptic, large (0.4 x 5.5 cm), brown; hairy plumes Other: the crown is deep, narrow and dark green. The bole is straight and cylindrical with smooth brown bark normally blotched with large patches of white lichen. It has no buttresses. It has lenticels in horizontal rows. The slash exudes latex. The wood density is 0.48 g/cm3.

F

Phenology Deciduousness: evergreen (Savill & Fox 1967) Dispersal: wind-blown with aid of cottony plume (Hawthorne 1995a) Timing: flowering period from November to May; fruiting period from December to February (De Koning 1983)

Distribution Continent: Benin to Angola, Mozambique (Hall & Swaine 1981) Upper Guinea: Guinea Bissau to Togo (Hall & Swaine 1981). It is common in Ghana, widespread in tropical Africa (Hall & Swaine 1981). Forest type: wet evergreen forest, moist evergreen forest, upland evergreen forest, moist semideciduous forest, dry semi-deciduous forest, secondary forest, gallery forest (Hall & Swaine 1981, Zwetsloot 1981, De Koning 1983)

Habitat Description

The abundance shows a strong increase in rainfall, to reach an optimum around 2000 mm/yr, whereafter it declines. The abundance decreases also with soil fertility (regression analysis). It prefers open conditions (Savill & Fox 1967).

Guild: np Life form: medium-sized tree Max. height: 30 m (Zwetsloot 1981) Max. diameter: 52 cm (inventory data Ghana) Leaf: opposite, simple, elliptic to ovate, mesophyll (1.7-17 x 5-32 cm), entire, coriaceous, dark green above, lighter green beneath; petiole up to 1.5 cm long, glabrous or minutely pubescent

Spp

416

type (De Koning 1983). Seeds germinate in light shade, but only become established in gaps and on old farmland (Hall & Swaine 1981).

n

Uses Many parts of the tree are used for medicinal purposes. The latex is sometimes used as bird lime. The floss of the seeds can be used to stuff pillows (Zwetsloot 1981).

Data sources FWTA, Taylor (1960), Savill & Fox (1967), Hall & Swaine (1981), Zwetsloot (1981), De Koning (1983), Hawthorne (1995a)

Regeneration It has a phanerocotylar epigeal foliaceous seedling

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

F.a.

142

7

25

61

8

6

31

58

11

22

2

71

All sites

198

4

26

57

9

8

31

58

11

25

2

72

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Page 417

Gilbertiodendron preussii

(Harms) J.Léonard

Leguminosae-Caes. Description

Regeneration

Guild: sb Life form: medium-sized to large tree Max. height: 35 m (Voorhoeve 1965) Max. diameter: 120 cm (Voorhoeve 1965) Leaf: alternate, pinnately compound, 6-10 opposite leaflets, narrowly elliptic, notophyll (2.5-5 x 7-19 cm), entire, coriaceous, dark green above, paler green beneath Inflorescence: axillary or terminal, branched (clusters of panicles) Flower: small; calyx pale yellow; corolla pale yellow Fruit: dry dehiscent (9 x 22 cm), fairly thin-walled, woody, yellowish to medium-brown; 3-5 seeds Seed: elliptic or irregularly formed, very large (2.5 x 2.8 cm) Other: a tree without buttresses, and with rough bark a little like Lophira. The new leaves flush red. It is probably ectomycorrhizal, like others in the genus. Wood density is 0.82 g/cm3.

It has a cryptocotylar epigeal reserve seedling type (cf. Voorhoeve 1965).

Phenology Deciduousness: evergreen (Voorhoeve 1965) Dispersal: seeds are ejected from the pod (Voorhoeve 1965) Timing: flowering period from October to December; fruiting period from July to September (Voorhoeve 1965)

G

Distribution Continent: Benin to Gabon (Voorhoeve 1965) Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana, Togo (Voorhoeve 1965) Forest type: rainforest, moist semi-deciduous forest. In some areas it is found in great abundance, e.g. hills of Cape Three Points, but generally rather uncommon.

Habitat The species is more abundant in areas with a high rainfall and a low altitude (regression analysis). It occurs on riverbanks (Taylor 1960).

Uses It is locally used as construction timber (Voorhoeve 1965).

Data sources Taylor (1960), Voorhoeve (1965), Alexander (1989), Lock (1989)

Spp

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

G.p.

48

8

4

54

21

21

63

31

6

65

0

31

All sites

198

4

26

57

9

8

31

58

11

25

2

72

417

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Guarea cedrata

Page 418

(A.Chev.) Pellegr.

Meliaceae Distribution Continent: Benin to Uganda, Congo (Voorhoeve 1965) Upper Guinea: Guinea to Togo (Voorhoeve 1965) Forest type: upland evergreen forest, wet evergreen forest, moist evergreen forest, moist semi-deciduous forest, dry semi-deciduous forest (Hall & Swaine 1981). A Red List species (Vulnerable).

Habitat

G

The species shows a bell-shaped response curve to rainfall; it increases with rainfall, to attains an optimum around 2000 mm/yr, and declines strongly above 2500 mm/yr (regression analysis). In Ghana it prefers areas with a high rainfall and infertile soils (Swaine 1996). In Liberia it is abundant in the evergreen forest on well-drained soils (Voorhoeve 1965). Locally often abundant as seedling in shade.

Description Guild: sb Life form: large tree Max. height: 40 m (Voorhoeve 1965) Max. diameter: 100 cm above the buttresses (Voorhoeve 1965) Leaf: alternate, imparipinnately compound, 5-11 (sub)opposite leaflets, narrowly elliptic to narrowly oblong, mesophyll (2-9 x 4-28 cm), entire, often markedly undulate, coriaceous, young leaves first reddish with pale green nerves, later pale green, leaves of saplings much larger with larger leaflets Inflorescence: axillary, panicle (1.5-7 cm long) Flower: small; corolla pale yellow Fruit: dry dehiscent, subglobose (3.3 cm in diameter), 2-4 thick-coriaceous valves, densely pubescent, yellowish when ripe; 2-4 seeds Seed: rounded triangular, large (1.5 x 1.8 cm), completely enclosed by a bright red aril Other: a tree with a dark, compact crown, and sometimes with buttresses. Lenticels are present in vertical rows. The slash has a characteristic, strong, sweet cedar scent. Wood density is 0.6 g/cm3.

Spp

418

n

Regeneration Germination is irregular and rather slow (averaging more than 5 weeks). Germination is not successful in full sunlight (Gilbert 1952), although Kyereh et al. (1993) report little difference between germination in the light and the dark. It has a phanerocotylar epigeal reserve seedling type (cf. De la Mensbruge 1966). The seedlings and saplings are common, even in the deepest shade. The seedlings are fairly drought sensitive (Swaine et al. 1997).

Growth Height growth is slow at first (25-30 cm after year one), but older plants can make faster growth when exposed to sunlight. Taylor (1960) records 14 year

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

G.c.

136

7

22

67

10

1

29

61

10

19

2

74

All sites

198

4

26

57

9

8

31

58

11

25

2

72

old trees in plantations with diameters of 8-20 cm (7-15 m high), and annual height increments of up to 1 m in Tropical Shelterwood System plots. Seedlings realise highest growth rates around 10% irradiance (Swaine et al. 1997). Even the older trees grow best with a top shade, with 20 year old trees being of a same size to 12 year old unshaded trees (10-32 cm dbh in 22 years, MacKay 1953). In Nigeria, the mean annual basal area increment varied from 0.6 to 12.14 cm per 30 x 30 m PSP per annum (Bada 1989). The highest growth rates occurred in plots which were treated by canopy opening or similar effects. Keay (1961) estimated 168 years for trees to attain 90 cm dbh. Growth rings in this species are likely to be annual (Detienne & Mariaux 1977).

Phenology Deciduousness: evergreen (Voorhoeve 1965) Dispersal: by animals. The seeds are eaten by birds and monkeys, duikers and porcupines (Voorhoeve 1965). Timing: flowering period in July; fruiting period from May to October (Voorhoeve 1965)

Uses A timber species (Voorhoeve 1965).

Data sources Gilbert (1952), MacKay (1953), Taylor (1960), Keay (1961), Voorhoeve (1965), De la Mensbruge (1966), Detienne & Mariaux (1977), Hall & Swaine (1981), Bada (1989), Kyereh et al. (1993), Hawthorne (1995a), Swaine (1996), Swaine et al. (1997), IUCN Red List (2000)

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Page 419

Guibourtia ehie

(A.Chev.) J.Léonard

Leguminosae-Caes. Description Guild: np Life form: large tree Max. height: 45 m (Voorhoeve 1965) Max. diameter: > 100 cm (Voorhoeve 1965) Leaf: alternate, pinnately compound, 2 leaflets, obliquely elliptic, microphyll (1.5-3.5 x 3-8 cm), glossy green above, paler beneath Inflorescence: axillary or terminal, branched (spike, 4-9 cm long) Flower: small; corolla white Fruit: dry dehiscent, rounded to quadrangular (3.5 x 5 cm), thin papery, black; 1 seed Seed: thin, round, large (1.5 cm in diameter), brown Other: a buttressed tree. The saplings have a red flush. Wood density is 0.83 g/cm3.

G

Distribution Continent: Upper Guinea endemic (Voorhoeve 1965) Upper Guinea: Liberia, Côte d’Ivoire, Ghana (Voorhoeve 1965) Forest type: moist evergreen forest, upland evergreen forest, moist semi-deciduous forest, dry semi-deciduous forest (Hall & Swaine 1981). It is common in Ghana (Hall & Swaine 1981). A Red List species (Vulnerable).

Habitat Its abundance increases with altitude and decreases with rainfall (regression analysis). The species is shade tolerant (Taylor 1960).

Phenology Deciduousness: deciduous (Voorhoeve 1965) Dispersal: by wind (Hawthorne 1995a) Timing: flowering period in November; fruiting period from January to February (Voorhoeve 1965)

Regeneration Germination requirements are not recorded. It has a phanerocotylar epigeal reserve seedling type (cf. De la Mensbruge 1966). This species is remarkable amongst canopy trees in the abundance of the seedlings which can almost always be found in the favoured MSNW forest zone. Seedling density is particularly great near to mature trees, where saplings and advanced regeneration tend also to occur gregariously (Hawthorne 1995a).

Data sources Taylor (1960), Voorhoeve (1965), De la Mensbruge (1966), Hall & Swaine (1981), Hawthorne (1995), IUCN Red List (2000)

Spp

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

G.e.

88

8

33

59

6

2

35

58

7

23

2

74

All sites

198

4

26

57

9

8

31

58

11

25

2

72

419

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Page 420

Hallea ledermannii

(K.Krause) Verdc.

Rubiaceae

H

Description

Habitat

Guild: sw Life form: medium-sized to large tree Max. height: 35 m (Voorhoeve 1965) Max. diameter: 115 cm (Voorhoeve 1965) Leaf: opposite, simple, suborbicular to obovate, macrophyll (x 10-40 cm), slightly undulate, coriaceous, medium green above, paler green beneath, scattered hairs on midrib and nerves, leaves of young trees and saplings often more elliptic and slightly more pubescent Inflorescence: axillary or terminal, cymose, flowerheads 1.5-2 cm across Flower: small; corolla white, tube-shaped Fruit: infructescence 1.3-2.3 cm across formed by numerous amphora-shaped capsules (5-8 mm); many seeds Seed: small (1.5 mm), flat, slightly winged Other: a rather narrow, slender-boled, unbuttressed, deep-crowned tree, with conspicuous, large leaves. It is often found in village swamplands and riversides. The flush of new leaves is red. It produces knee roots a few cms in height. As mud builds up around the knee roots, they branch, to form compound structures. Wood density is 0.55 g/cm3.

It is a light demanding and gregarious species with trees growing close together (Taylor 1960). It is found in places where it has subsoil moisture during the whole year, but it does not grow in deep, all-year flooded swamps (Voorhoeve 1965).

Distribution

Uses

Continent: Benin to Congo (Voorhoeve 1965) Upper Guinea: Liberia, Ivory Coast, Ghana, Togo (Voorhoeve 1965) Forest type: swamp forest. There is much scope for dispersing this species to swamps in unforested areas. This species is described in general terms by Osain (1973). A Red List species (Vulnerable).

A valuable timber species.

Regeneration It regenerates in swamp forest.

Growth In taungya plantations, the species attained approx. 10 m of height in 9 years (Taylor 1960).

Phenology Deciduousness: evergreen (Voorhoeve 1965) Dispersal: by wind Timing: flowering period from November to January; fruiting period from February to May (Voorhoeve 1965)

Data sources Taylor (1960), Voorhoeve (1965), Jenìk (1969), Osain (1973), IUCN Red List (2000)

Map data are not available.

Table data are not available.

420

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Page 421

Khaya anthotheca

(Welw.) C.DC.

Meliaceae Description

Regeneration

Guild: np Life form: large tree Max. height: 45 m (Voorhoeve 1965) Max. diameter: 120 cm (Voorhoeve 1965) Leaf: alternate, paripinnately compound, 4-8 leaflets, leaflet elliptic, oblong or obovate (2.5-5.5 x 5-10 cm), entire, herbaceous Inflorescence: terminal, branched (raceme or panicle, 6-25 cm long) Flower: small; corolla white Fruit: dry dehiscent, slightly appressed globose (6 x 7 cm), woody, smooth, grey; numerous seeds Seed: winged, very thin, including the wing 2-2.5 cm x 3.5-5 cm, bright brown Other: an emergent, buttressed tree. The stem is slender and the crown small, until the upper canopy is reached, whereupon it diverts much of its energy to lateral growth.

Kyereh et al. (1993) reported no difference between germination in the light and in the dark. It has a cryptocotylar epigeal reserve seedling type (cf. Voorhoeve 1965). Very sparse regeneration of this species was found in Côte d’Ivoire in a 3 ha subsample of a plot containing several adults (De Klerk 1991).

Distribution Continent: Benin to Uganda, Angola (Voorhoeve 1965) Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana, Togo (Voorhoeve 1965) Forest type: moist semi-deciduous forest, dry semi-deciduous forest (Hall & Swaine 1981) Other: common in Ghana (Hall & Swaine 1981). A Red list species (Vulnerable).

Growth Seedling growth is maximal around 10% irradiance (Swaine et al. 1997). Any growth rings in trees of this species are not likely to be annual (Detienne & Mariaux 1977).

Phenology Deciduousness: evergreen or briefly deciduous (Voorhoeve 1965) Dispersal: by wind (Voorhoeve 1965) Timing: flowering period from September to October and January to February; fruiting period in January (Voorhoeve 1965)

K

Habitat The abundance of Khaya spp. as a group increases with rainfall, to attain an optimum around 1800 mm/yr and decline strongly beyond 2300 mm/yr. The abundance declines with altitude (regression analysis). K. anthotheca grows scattered, often on slopes towards creeks and river borders (Voorhoeve 1965).

Uses

Data sources

An important timber tree (Hawthorne 1995a).

Taylor (1960), Voorhoeve (1965), Detienne & Mariaux (1977), Hall & Swaine (1981), De Klerk (1991), Kyereh et al. (1993), Hawthorne (1995a), Swaine et al. (1997), IUCN Red List (2000).

Spp

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

K.spp.

151

5

27

64

8

1

27

62

11

19

2

75

All sites

198

4

26

57

9

8

31

58

11

25

2

72

421

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Page 422

Khaya grandifoliola

C.DC.

Meliaceae Habitat The abundance of Khaya spp. as a group increases with rainfall, to attain an optimum around 1800 mm/yr and decline strongly beyond 2300 mm/yr. The abundance declines with altitude (regression analysis).

Regeneration

Guild: np Life form: large tree Max. height: 50 m (Taylor 1960) Max. diameter: data unavailable Leaf: alternate, paripinnately compound, 6-10 opposite or sub-opposite leaflets, elliptic to oblongelliptic (7 x 16 cm), entire Inflorescence: terminal, branched (panicle) Flower: small; corolla creamy-white Fruit: dry dehiscent, globose (approx. 8 cm in diameter), woody Seed: flat, winged Other: it has narrow high buttresses, and a spreading crown with drooping shiny leaves. The slash has a bitter taste and a clear gum exudes from wounds. Wood density is 0.7 g/cm3.

It has a cryptocotylar epigeal reserve seedling type (cf. De la Mensbruge 1966). Kemp (1961), Webb (1964), and Jackson (1973) have discussed the regeneration in gallery (“kurmi”) forest in Nigeria. In these patches, three zones were defined: internal, close to the streams, a dry boundary zone, near the edge and the savanna, and an intermediate zone. Although mature trees occurred throughout, regeneration was greatest in the boundary zone, where the canopy was least complete (soil differences were also recorded). This is true particularly where the edge is free from fire damage. Regeneration is limited by excessive shade. Experiments carried out with permanent sample plots in kurmi forest showed how in closed forest seedlings can become established, but rarely survive more than 2 years, whereas seedlings under more open canopy (e.g. in the boundary zone, and where the forest has been treated) often survive 10 years, and probably to maturity. The conclusion of these studies was that the species is a “pioneer” in the transition from savanna to forest, according with Jones’ (1963) observation that larger trees tend to be nearer the centre of the forest patches.

Distribution

Growth

Continent: Upper Guinea endemic Upper Guinea: Guinea, Côte d’Ivoire, Ghana, Togo Forest type: dry semi-deciduous forest, gallery forest. A Red List species (Vulnerable).

Increment in Nigeria is correlated with tree size, but not obviously with environmental variables (Kemp & Lowe 1973). The growth rate was low (57-67 cm dbh in 100 years), dependent on annual rainfall and

Description

K

Spp

422

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

K.spp.

151

5

27

64

8

1

27

62

11

19

2

75

All sites

198

4

26

57

9

8

31

58

11

25

2

72

on competition even from the surrounding smaller plants (Kemp 1961). In plantations, higher rates of increment are recorded. Seedlings take approx. 15 years to attain 6.7 cm in diameter (Jackson 1973). Trees grow fastest in months of highest rainfall (Ola Adams & Charter 1980). Plant growth increases with the degree of endomycorrhizal infection (Redhead 1975). Growth rings occur, but are not correlated with age (Hummel 1946).

Phenology Deciduousness: sometimes deciduous Dispersal: by wind Timing: flowering in the dry season with the flush of new leaves (Taylor 1960); fruiting period from October to March. There is a flush of leaves and flowering in the dry season in Nigeria (Ola Adams & Charter 1980). In Nigeria, more seeds and seedlings are produced in years of greater rainfall (Kemp 1961). Seedling density is higher very close to parent trees, and in areas of sparse herb cover.

Uses The bark of this and other Khaya species is effective against malaria (Awe & Makinde 1991), and used as such in Ghana (Abbiw 1990).

Data sources Hummel (1946), Taylor (1960), Kemp (1961), Jones (1963), Webb (1964), De la Mensbruge (1966), Jackson (1973), Kemp & Lowe (1973), Redhead (1975), Hall & Swaine (1981), Ola Adams & Charter (1980), Abbiw (1990), Awe & Makinde (1991), IUCN Red List (2000)

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Page 423

Khaya ivorensis

A.Chev.

Meliaceae Description Guild: np Life form: large tree Max. height: > 50 m (Taylor 1960) Max. diameter: > 180 cm (Taylor 1960) Leaf: paripinnately compound, 12 opposite leaflets, oblong to oblong-elliptic (4 x 9 cm), entire, glabrous, leaves of saplings much longer with up to 20 pairs of leaflets Inflorescence: branched (panicle) Flower: small; corolla yellow Fruit: capsule, globose (12 cm in diameter), woody; numerous seeds Seed: flat, winged (2 x 3 cm, including wing) Other: a high-buttressed tree, with dark-green, rounded crown, often with the pendulous, spherical fruits visible. The slash is scented, tastes bitter, and a pale gum exudes from wounds. Wood density is 0.51 g/cm3.

Distribution Continent: Benin to Gabon (Hall & Swaine 1981) Upper Guinea: Liberia, Côte d’Ivoire, Ghana, Togo (Hall & Swaine 1981) Forest type: wet evergreen forest, moist evergreen forest, moist semi-deciduous forest. It is common in Ghana (Hall & Swaine 1981), A Red List species (Vulnerable).

Habitat The abundance of Khaya spp. as a group increases with rainfall, to attain an optimum around 1800 mm/yr and decline strongly beyond 2300 mm/yr. The abundance declines with altitude (regression analysis). K. ivorensis is a light demander but with a narrow crown, and possibly other attributes that seem to suit it more to small and medium-sized gaps than to larger-grained disturbance (Hawthorne 1995a). It prefers heavy or rich alluvial soils near

water courses and damp areas, but must have good drainage (Taylor 1960).

Regeneration Seeds suffer heavily from predation. Germination is normal, with limited viability (from 100% at first, down to 5% success after 3 months, Sanders 1953). Under controlled conditions there is no difference between germination in the light and in the dark, although germination is strongly depressed in large gaps (Kyereh et al. 1993). It has a cryptocotylar epigeal reserve seedling type (cf. De la Mensbruge 1966). Seedlings require to be lightly shaded in nurseries, for about the first two years (Taylor 1960). When exposed to high irradiance they suffer from attacks by the Hypsiphyla moth larvae, and from leaf galls. In natural forest, seedlings can germinate and survive below a dense layer (Sanders 1953). Nevertheless seedling survival is encouraged by increasing light levels above those below the complete canopy. Seedlings are common in gaps below climber tangles (Sanders 1953). Saplings suffer from damage by porcupines, which eat the bark. Lancaster (1954) noted that germination (best in the shade) and seedling growth (best in “considerable light”) requirements conflict.

Phenology Deciduousness: deciduous Dispersal: by wind Timing: flowering period from July to January; fruiting period from February to May (Taylor 1960). In the Moist Evergreen zone, flowering and fruiting occurs throughout the year, while in the Moist Semideciduous zone flowering occurs from June to September and fruiting from October to March (Gyimah 1986). Trees of 30 years can produce good crops of seeds and, although some seed is produced annually, good seed crops occur every 3-4 years (Sanders 1953). Lancaster (1954) recorded a very patchy spatial distribution of seeds, even during a good year.

Uses

Growth

Spp

n

K

An important timber tree, but often confused with other mahoganies by exploiters and as a young tree by tree spotters (Hawthorne 1995a). The bark is used as a native cure for dysentery (Taylor 1960).

The Tropical Shelterwood System management is well suited for K. ivorensis. Height growth in TSS is approx. 1 m in 4 years, increasing to almost 3 m per year after approx. 7 years (Taylor 1960). Later, height increment slows again to approx. 1 m per year over 30 years (Sanders 1953). The mean annual increment in Nigeria is approx. 2.5 cm, depending on height and competition intensity (Sanders 1953). In some sites, the tree attains 10 m height in 7 years (MacKay 1953). Keay (1961) estimated that it would take 88 years for this species to reach 90 cm dbh, in Nigeria.

Data sources MacKay (1953), Sanders (1953), Lancaster (1954), Taylor (1960), Keay (1961), De la Mensbruge (1966), Hall & Swaine (1981), Gyimah (1986), Kyereh et al. (1993), Kyereh (1994), Hawthorne (1995a), IUCN Red List (2000)

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

K.spp.

151

5

27

64

8

1

27

62

11

19

2

75

All sites

198

4

26

57

9

8

31

58

11

25

2

72

423

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Page 424

Klainedoxa gabonensis

Pierre ex Engl.

Irvingiaceae Description

light demander (Savill & Fox 1967), but does not require full overhead light for its development (Taylor 1960). Often, it is associated with damp situations, but not where the drainage is impeded (Taylor 1960).

Guild: np Life form: large tree Max. height: 40 m (Voorhoeve 1965) Max. diameter: 120 cm (Voorhoeve 1965) Leaf: alternate, simple, ovate to elliptic, notophyll (2.8-8 x 6-15 cm), slightly undulate, coriaceous, glossy above, glabrous, leaves of seedlings, saplings and young trees very different: narrowly elliptic, -ovate or -oblong (up to 12 x 40 cm); stipules up to 15 cm long Inflorescence: axillary or terminal, branched (10-12 cm long) Flower: small; corolla mauve to purple Fruit: fleshy (drupe) (4 x 6.5 cm), green to purplish black; 4-5 seeds Seed: a woody nut Other: an emergent tree with a wide, but domed, dense crown. Large trees often have very extensive plank buttresses. Young trees have short, 3-5 cm long sharp spines on bole and buttresses. The leaf flush is brilliant red. Wood density is 1.07 g/cm3.

Regeneration Germination, as far as known, is normal, although the effect of elephant ingestion is not recorded. It has a phanerocotylar epigeal reserve seedling type (cf. Voorhoeve 1965). Seedlings occur in the shade. Saplings are found even in small gaps, where they usually have leaves much longer than those of the adults (Hawthorne 1995a).

Phenology Deciduousness: evergreen (Voorhoeve 1965) Dispersal: by elephants (Alexandre 1978). During the rainy season, seeds of this species were found in 32% of piles of elephant dung in Bia South GPR (Martin 1991). Timing: flowering period from August to November; fruiting period from February to March (Voorhoeve 1965)

Distribution

K

Continent: Benin to Uganda (Voorhoeve 1965) Upper Guinea: Sierra Leone to Togo (Voorhoeve 1965) Forest type: wet evergreen forest, moist evergreen forest, moist semi-deciduous forest, dry semideciduous forest, secondary forest (Taylor 1960, Hall & Swaine 1981). It is common in Ghana (Hall & Swaine 1981).

Uses The fruits can be used for an edible oil (Abbiw 1990).

Data sources

Habitat

Taylor (1960), Voorhoeve (1965), Savill & Fox (1967), Alexandre (1978), Hall & Swaine (1981), Abbiw (1990), Martin (1991), Hawthorne (1995a)

The abundance increases linearly with rainfall up to 2000 mm/yr, and attains an optimum around 2600 mm/yr. It is more abundant at low altitudes (regression analysis). The species is said to be a

Spp

424

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

K.g.

150

7

21

61

9

9

34

57

9

26

2

68

All sites

198

4

26

57

9

8

31

58

11

25

2

72

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Page 425

Lophira alata

Banks ex Gaertn.

Ochnaceae Description

Regeneration

Guild: pi Life form: large tree Max. height: 50 m (Voorhoeve 1965) Max. diameter: 150 cm (Voorhoeve 1965) Leaf: alternate, simple, obovate, mesophyll (3-9 x 8-25 cm), slightly undulate, glossy dark green, leaves of sapling much larger (up to 10 x 100 cm) Inflorescence: terminal, unbranched (lax panicle) Flower: medium-sized; corolla white Fruit: dry indehiscent (1.2 x 2.8 cm), winged with unequal sized wings; 1 seed Seed: data unavailable Other: a tree with a distinctive, modest crown of tufted leaves. The flush of new leaves is bright red. Wood density is 1.09 g/cm3.

Seeds have no recorded dormancy and limited viability. However, they do not germinate in the shaded understorey. It has a cryptocotylar epigeal reserve seedling type (cf. Voorhoeve 1965). Healthy seedlings are associated with canopy gaps, although in full sunlight it is said to be more susceptible to insect attack (Taylor 1960). Saplings are light demanders, and common along roads or in recent gaps.

Distribution Continent: Togo to Angola (Voorhoeve 1965) Upper Guinea: Sierra Leone to Togo (Voorhoeve 1965) Forest type: upland evergreen forest, wet evergreen forest (Hall & Swaine 1981). A Red List species (Vulnerable).

Habitat Its abundance increases strongly with rainfall to attain an optimum around 2600 mm/yr. The abundance decreases with soil water holding capacity and soil fertility (regression analysis). It is drought-sensitive (Veenendaal & Swaine 1996), and grows better on infertile soils (Veenendaal et al. 1996). In Cameroon, it occurs partially on poor, acid, white sand soils (McKey et al. 1978). It is a light demander (Voorhoeve 1965). Letouzey (1957, 1960, 1978) has discussed possible origins of Lophira forest, as a secondary forest following farming, which does not regenerate naturally, under present conditions.

Growth In a silvicultural trial, a sapling reached 3 m height in 4 years (Taylor 1960). In a large gap mixed with Musanga, after 7 years, Lophira trees were almost 10 m tall, and healthy even though overtopped by Musanga trees of 15-20 m tall (Taylor 1960). In Sierra Leone, mean annual diameter increments of 0.65 cm, with a maximum of 2.65 cm/yr have been found (Savill & Fox 1967). Slower mean growth rates are recorded for Nigeria, with an estimated 220 years to reach 90 cm dbh (Keay 1961).

Phenology Deciduousness: deciduous from October to November (Voorhoeve 1965) Dispersal: by wind (Voorhoeve 1965) Timing: flowering period from November to December; fruiting period from March to June (Voorhoeve 1965)

Letouzey (1957, 1960, 1978), Taylor (1960), Keay (1961), Voorhoeve (1965), Savill & Fox (1967), McKey et al. (1978), Hall & Swaine (1981), Abbiw (1990), Amougou (1990), Swaine & Veenendaal (1994), Veenendaal & Swaine (1996), Veenendaal et al. (1996), IUCN Red List (2000)

Uses One of the best heavy timbers produced in the tropics. The fruits can be used for an edible oil (Abbiw 1990).

Spp

n

L

Data sources

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

L.a.

86

12

7

60

17

15

53

40

7

44

1

49

All sites

198

4

26

57

9

8

31

58

11

25

2

72

425

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Page 426

Lovoa trichilioides

Harms

Meliaceae Distribution Continent: Benin to Angola (Voorhoeve 1965) Upper Guinea: Sierra Leone to Togo (Voorhoeve 1965) Forest type: upland evergreen forest, wet evergreen forest, moist evergreen forest, moist semi-deciduous forest (Hall & Swaine 1981). A Red List species (Vulnerable).

Habitat The abundance increases strongly with rainfall, to attain an optimum around 2400 mm/yr (regression analysis). It is more common in the wetter forest types (Voorhoeve 1965). Strongly associated with acid, base-poor soils (Hawthorne 1995a).

Regeneration

Description

L

Spp

426

Germination is normal (Taylor 1960). The seed has a short viability (only 30% after 2 months, Sanders 1953), and suffers heavy predation. No differences are found between germination in the light and in the dark (Kyereh et al. 1993). It has a cryptocotylar epigeal reserve seedling type (cf. Voorhoeve 1965) and is remarkable for the abundance of seedlings in the full shade for several years. It is less susceptible to borers and other insects than Khaya ivorensis (Sanders 1953). Saplings only develop in canopy gaps, although in a less shaded plot in Nigeria 5 year mortality was similar to that of completely shaded plots (Sanders 1953).

Guild: np Life form: large tree Max. height: 45 m (Voorhoeve 1965) Max. diameter: 100 cm (Voorhoeve 1965) Leaf: alternate, (im)paripinnately compound, 6-10 (sub)opposite leaflets, elliptic to obovate, mesophyll (3-8 x 6-20 cm), entire, coriaceous, glabrous Inflorescence: (sub)terminal, branched (panicle, 15-40 cm long) Flower: small; corolla white Fruit: dry dehiscent, spindle-shaped (1.3 x 4.3 cm), purplish black with 4 coriaceous valves; 4-8 seeds Seed: medium-sized (0.6 cm wide with a 0.7 x 2.5 cm wing) Other: an emergent tree with thick ascending buttresses and sometimes heavy, spreading surface roots. The crown is thick, dense and dark green. Brown, scattered lenticels are common. The slash has a strong cedar-like scent. Wood density is 0.52 g/cm3.

n

Growth In nurseries, seedlings attain 1 m after 2 years, and in natural forest 9 m over the first 7 years (Sanders 1953). Keay (1961) predicted 106 years needed to grow 90 cm in dbh. The initial growth

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

L.t.

115

9

16

63

12

10

41

53

6

32

1

63

All sites

198

4

26

57

9

8

31

58

11

25

2

72

rate is slower than Khaya ivorensis, while after this growth is rapid, as Lovoa does not slow down in later life like Khaya (Sanders 1953). Increment still rises at 32 years, by which time it starts to overtake Nauclea planted at the same time (Horne 1962). MacKay (1953) records trees of 30 m height (10 cm dbh) after 22 years. There is no influence of site factors, nor of crown diameter on growth rate (Akachuku 1984). Age alone explained most of the variation. It is particularly successful when planted in groups under thinned natural forest. Any growth rings are not likely to be annual (Detienne & Mariaux 1977).

Phenology Deciduousness: evergreen Dispersal: by wind Timing: flowering period in the dry season; fruiting period from March to April (Taylor 1960). The seeds are not produced annually. In Nigeria, a good seed year occurs every 3-4 years (Sanders 1953).

Uses A timber species (Voorhoeve 1965).

Data sources MacKay (1953), Sanders (1953), Taylor (1960), Keay (1961), Horne (1962), Voorhoeve (1965), Detienne & Mariaux (1977), Hall & Swaine (1981), Akachuku (1984), Kyereh et al. (1993), Hawthorne (1995a), IUCN Red List (2000)

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Page 427

Mammea africana

Sabine

Guttiferae Description Guild: sb Life form: large tree Max. height: 40 m (Voorhoeve 1965) Max. diameter: 100 cm (Voorhoeve 1965) Leaf: opposite, simple, narrowly elliptic, mesophyll (4-10 x 12-26 cm), entire, coriaceous, blade densely dotted with slightly raised glandular cells Inflorescence: axillary on leafy shoots and cauliflorous, solitary, hermaphrodite or unisexual (male) Flower: medium-sized (male) to large (hermaphrodite); corolla white Fruit: fleshy, elliptic to globular (6.5 x 8.5 cm), pale yellow with small brown warts; 2-4 seeds Seed: laterally flattened, very large (1.8 x 3 cm), fruit pulp Other: a tree with a very regular, cylindrical bole and dense evergreen crown of short, regular, horizontal branches. The flush of new leaves is red. The slash exudes a yellow latex. It has heavy buttresses of up to 3.5 m high. Wood density is 0.77 g/cm3.

trees attained approx. 32 cm dbh (almost 15 m high) in 24 years (MacKay 1953).

Phenology

swampy valleys. The tree is markedly shade tolerant (Voorhoeve 1965).

Deciduousness: evergreen (Voorhoeve 1965) Dispersal: by elephants (Hall & Swaine 1981), porcupines, antelopes. Parren (1991) concluded that regeneration was not significantly greater in elephant forests than in those without elephants. Timing: flowering is irregular; fruiting period from January to February and June to July (Voorhoeve 1965)

Regeneration

Uses

Germination often takes many months. It has a cryptocotylar hypogeal reserve seedling type (cf. Voorhoeve 1965). The saplings are markedly shadetolerant (Taylor 1960).

The fruits (“African Mammy Apple”) can be used for an edible oil (Abbiw 1990).

Distribution Continent: Benin to Congo (Brazzaville), Uganda (Voorhoeve 1965) Upper Guinea: Sierra Leone to Togo (Voorhoeve 1965) Forest type: wet evergreen forest, moist evergreen forest, moist semi-deciduous forest, dry semideciduous forest (Hall & Swaine 1981)

Habitat The abundance increases up to a rainfall amount of 2000 mm/yr, whereafter it remains more or less constant. In Ghana it is strongly associated with base-poor soils (Hall & Swaine 1981). It seems to prefer moist, alluvial sites and may occur in

Data sources

Growth

MacKay (1953), Taylor (1960), Voorhoeve (1965), Hall & Swaine (1981), Abbiw (1990), Parren (1991), Hawthorne (1995a)

Saplings grow approximately 30 cm per year (Taylor 1960). In Nigeria, the species is said to be slowgrowing at first (60 cm height after 2 years), but

Spp

n

M

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

M.a.

102

9

16

59

13

13

31

60

9

25

1

66

All sites

198

4

26

57

9

8

31

58

11

25

2

72

427

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Milicia excelsa

Page 428

(Welw.) C.C.Berg

Moraceae Habitat Its abundance decreases with rainfall and altitude (regression analysis). The species is an intense light demander (De Rouw 1991), but does best under light shade. It prefers well-drained soils and is intolerant of impeded drainage (Taylor 1960).

Regeneration This is one of the few pioneer species whose germination is strongly photoblastic (Kyereh et al. 1993). It has a phanerocotylar epigeal foliaceous seedling type (cf. De la Mensbruge 1966). Most regeneration occurs in open places. Seedling growth is high and fairly constant over a large range of light environments (Swaine et al. 1997). A height growth of 1 m is not unusual in the first year. Young shoots are browsed by duiker (Taylor 1960).

Phenology Description Guild: pi Life form: large, emergent tree Max. height: > 50 m (Taylor 1960) Max. diameter: > 150 cm (Taylor 1960) Leaf: alternate, simple, elliptic, mesophyll (9 x 12 cm), entire to slightly undulate, coriaceous, slightly pubescent underneath, juvenile leaf very different, oblong to lanceolate (7 x 25 cm), serrate, herbaceous, pilose Inflorescence: axillary, peduncle (male inflorescence up to 22 cm long, female inflorescence 1 x 4 cm) Flower: dioecious; small Fruit: infructescence composed of small achenes Seed: data unavailable Other: the dark foliage and heavy branches make the crown rather conspicuous. The bark has

M

Spp

428

Deciduousness: deciduous from December to January (Taylor 1960) Dispersal: by birds and bats (Taylor 1960) Timing: flowering period from December to March; fruiting period from March to May (Taylor 1960)

n

numerous lenticels and becomes scaly. The slash exudes a sticky latex. The roots feature long yellow, horizontal lenticels.

Uses A timber species.

Distribution Continent: Benin to Mozambique (Voorhoeve 1965) Upper Guinea: Senegal to Togo (Voorhoeve 1965) Forest type: moist evergreen forest, moist semideciduous forest, dry semi-deciduous forest, derived savanna-woodland, riverine forest. A Red List species (lower risk).

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

M.spp.

156

7

26

59

9

6

29

59

12

23

3

69

All sites

198

4

26

57

9

8

31

58

11

25

2

72

Data sources Taylor (1960), Voorhoeve (1965), De la Mensbruge (1966), De Rouw (1991), Kyereh et al. (1993), Swaine et al. (1997), IUCN Red List (2000)

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Page 429

Nauclea diderrichii

(De Wild. & T.Durand) Merrill

Rubiaceae Description

Regeneration

Guild: pi Life form: large tree Max. height: 50 m (Voorhoeve 1965) Max. diameter: 150 cm (Voorhoeve 1965) Leaf: opposite, simple, elliptic to oblong or slightly obovate, macrophyll (8-16 x 10-30 cm), entire, herbaceous, glabrous Inflorescence: terminal, subglobular flowerhead (2.5-4 cm across) Flower: small; corolla pale yellow, tube-shaped Fruit: fleshy, subglobose (3.3 cm in diameter), orange; many seeds Seed: small (0.1 cm in diameter), hard and smooth Other: a tree with cylindrical, unbuttressed bole and rather horizontal, whorled boughs, in a narrow crown. The base has heavily swollen root spurs, sometimes extending in spreading surface roots. Wood density is 0.77 g/cm3.

Germination is said to be normal, with no dormancy noted by Taylor (1960), although Sawyerr (1960) describes extreme treatments needed to encourage germination. The seeds lie dormant in the soil until stimulated to germinate by increased exposure to the sun (Hall & Swaine 1981). This is one of the few pioneer species whose germination is photoblastic (Kyereh et al. 1993). Germination is also triggered by a high red/far-red ratio (Kyereh et al. 1993). Seedling mortality can be high, due to too much shading, or too great exposure leading to borer attack (Taylor 1960). This certainly seems to be a species of big gaps. In Nigeria, it regenerates in large gaps, reaching the canopy after about 15 years (Ross 1954). Yet, Wadsworth & Lawton (1968) note optimum growth rates well below full sunlight.

Distribution Continent: Benin to Mozambique (Voorhoeve 1965) Upper Guinea: Sierra Leone to Togo (Voorhoeve 1965) Forest type: wet evergreen forest, moist evergreen forest. A Red List species (Vulnerable).

Habitat Its abundance increases with rainfall up to an amount of 2000 mm/yr, whereafter it remains more or less constant. The species is more abundant on infertile soils (regression analysis). It is a strong light demander (Taylor 1960) and prefers light, welldrained soils (Voorhoeve 1965).

Growth On old logging tracks, trees attained 12 m (10 cm dbh) height within 4 years (Hawthorne 1993). It is widely used in taungya and other plantation (e.g. Neil 1983), and recommended as a nurse crop for Meliaceae. In Nigeria, 26 year old taungya plantations had a mean height of 16 m and a mean dbh of 27 cm (Okojie & Nokoe 1985). See also Henry (1960), Horne (1962) and Lancaster (1952).

Uses A timber species.

Phenology

Data sources

Deciduousness: evergreen Dispersal: by elephants Timing: flowering period from May to August; fruiting period from September to October (Hawthorne 1995a)

Lancaster (1952, 1961), Ross (1954), Henry (1960), Keay (1960), Sawyerr (1960), Taylor (1960), Horne (1962), Voorhoeve (1965), Wadsworth & Lawton (1968), Hall & Swaine (1981), Neil (1983), Okojie & Nokoe (1985), Hawthorne (1993, 1995a), Kyereh et al. (1993), Hawthorne & Parren (2000), IUCN Red List (2000)

Spp

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

N

Soil WHC L

M

H

N.d.

145

8

21

59

10

10

35

59

6

26

2

68

All sites

198

4

26

57

9

8

31

58

11

25

2

72

429

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Page 430

Nesogordonia papaverifera

(A.Chev.) Capuron

Sterculiaceae Seed: flat, medium-sized (0.6 cm long), winged (wing 0.7 x 1.5 cm) Other: a slender tree with narrow buttresses, and a small, dark, dense crown. Wood density is 0.77 g/cm3.

Distribution

Description Guild: sb Life form: large tree Max. height: 45 m (Voorhoeve 1965) Max. diameter: 120 cm (Voorhoeve 1965) Leaf: alternate, simple, elliptic to obovate, notophyll (2.5-6 x 5-13 cm), entire, fairly coriaceous, glabrous above, slightly puberulous beneath, medium green Inflorescence: axillary, branched (cymose) Flower: medium-sized; corolla yellowish white Fruit: dry dehiscent (capsule), bell-shaped (2 x 3 cm), woody, brown, densely puberulous; up to 10 seeds

N

Spp

430

n

Growth In Tropical Shelterwood plots the saplings can reach 1-1.5 m in 4 years (Taylor 1960). In undisturbed forest in Nigeria, mean annual diameter increments of 0.5 cm have been recorded for 2.5-5 cm diameter trees (Dommen 1957).

Continent: Togo to Gabon (Voorhoeve 1965) Upper Guinea: Sierra Leone to Togo (Voorhoeve 1965) Forest type: upland evergreen forest, moist evergreen forest, moist semi-deciduous forest, dry semi-deciduous forest, secondary forest (Voorhoeve 1965, Hall & Swaine 1981). It is common in Ghana (Hall & Swaine 1981).

Phenology

Habitat

Uses

The abundance of the species decreases strongly with rainfall, and to a lesser extent with soil fertility (regression analysis), although in Ghana it does not show a preference for certain rainfall or soil fertility conditions (Swaine 1996). It is not markedly light demanding, avoids swampy localities and favours hills and base-rich soils. It is often found growing in groups (Voorhoeve 1965).

A timber species.

Regeneration Germination is regular, with a slightly lower than normal (73%) germination success (Taylor 1960). Light shade is apparently needed on germination beds. It has a phanerocotylar epigeal foliaceous seedling type (cf. Voorhoeve 1965). Seedlings benefit from moderate forest canopy, with an increasing requirement for overhead light with age. Saplings, nevertheless, are common in mediumsized to large gaps in some areas.

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

N.p.

143

5

31

64

4

1

22

66

13

16

3

76

All sites

198

4

26

57

9

8

31

58

11

25

2

72

Deciduousness: mostly evergreen, but Taylor (1960) records that it may be deciduous for a brief period in the dry season. Dispersal: by wind (Voorhoeve 1965) Timing: flowering period in May; fruiting period in May, July and December (Voorhoeve 1965)

Data sources Dommen (1957), Taylor (1960), Voorhoeve (1965), Hall & Swaine (1981), Hawthorne (1995a), Swaine (1996)

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Page 431

Parinari excelsa

Sabine

Chrysobalanaceae Description

Regeneration

Guild: pi Life form: large tree Max. height: 45 m (Voorhoeve 1965) Max. diameter: 150 cm (Voorhoeve 1965) Leaf: alternate, simple, elliptic, notophyll (2.5-5 x 6-13 cm), entire, glabrous and glossy green above, brownish or greyish felty below; leaflets of flowering twigs often narrowly elliptic, microphyll (1-4 x 4-10 cm); leaves of saplings and water shoots oblong (up to 7 cm x 20 cm) Inflorescence: axillary, branched (up to 20 cm long), slender twigs with slightly modified leaves Flower: small; corolla white Fruit: fleshy, ovoid (2.8 x 4.3 cm), coriaceous, rough, covered with numerous small lenticels, orange-brown or grey when ripe; 1 seed Seed: data unavailable Other: the foliage often has a reddish or brownish glow when seen from below. Older trees have thick and narrow buttresses. The bark is densely covered with warty lenticels. Wood density is 0.81 g/cm3.

It has a cryptocotylar epigeal reserve seedling type (cf. Voorhoeve 1965). Regeneration in high forest is scarce, but common in broken up forest, logging roads and young secondary forest (Voorhoeve 1965)

Phenology Deciduousness: evergreen Dispersal: by birds and mammals (Voorhoeve 1965), among others elephants (Hawthorne & Parren 2000) Timing: flowering period from January to June; fruiting period from October to January

Distribution Continent: Senegal to Kenya (Voorhoeve 1965) Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana (herbarium) Forest type: wet evergreen forest, moist evergreen forest, moist semi-deciduous forest, secondary forest

Uses

Habitat

A timber species. The fruits are edible and highly esteemed.

The abundance of Parinari spp. as a group increases with rainfall, to reach an optimum around 2400 mm/yr. Their abundance decreases strongly with soil fertility (regression analysis). P. excelsa prefers moist alluvial soils (Taylor 1960).

Data sources Taylor (1960), Voorhoeve (1965), Hawthorne & Parren (2000)

P

Spp

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

P.spp.

132

8

18

61

10

11

36

58

7

29

1

65

All sites

198

4

26

57

9

8

31

58

11

25

2

72

431

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Parkia bicolor

Page 432

A.Chev.

Leguminosae-Mim. Habitat In its early youth it is tolerant of moderate shade, but it is essentially a light demander (Taylor 1960). Mature trees, although found as emergents in a wide variety of sites, show some preference for wet areas, like riverbanks (Hawthorne 1995a).

Regeneration Germination is normal. It has a phanerocotylar epigeal reserve seedling type (cf. Voorhoeve 1965). Although seedlings are found in the shade, particularly in the vicinity of parent trees, saplings are soon restricted to more exposed sites.

Growth It can attain a height growth of about 1 m in the first year (Taylor 1960).

Description Guild: np Life form: large tree Max. height: 40 m (Voorhoeve 1965) Max. diameter: 120 cm (Voorhoeve 1965) Leaf: alternate, bipinnately compound, 32-40 opposite pinnae, each with 56-100 opposite leaflets, narrowly oblong, leptophyll (0.1-0.2 x 0.5-1 cm), entire, glabrescent, leaves of saplings as a rule with less pinnae and less leaflets, but leaflets larger, (up to 0.3 x 1.5 cm) Inflorescence: a pendent peduncle Flower: small; corolla pink to red Fruit: dry indehiscent, strap-shaped (1.8 x 30 cm), first orange to yellow, purplish black when ripe; up to 20 seeds Seed: flat, imbedded in a yellowish, mealy pulp Other: a buttressed tree with a large, spreading crown. The new leaves are copper-coloured. Wood density is 0.55 g/cm3.

Phenology Deciduousness: deciduous after the rainy season (Voorhoeve 1965) Dispersal: probably by mammals (Hawthorne 1995a) Timing: flowering period from December to February; fruiting period from February to April (Voorhoeve 1965)

Distribution

Uses

Continent: West Africa, south of the Sahara and Central Africa (Voorhoeve 1965) Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana (herbarium) Forest type: upland evergreen forest, wet evergreen forest, moist evergreen forest, moist semi-deciduous forest, dry semi-deciduous forest, secondary forest.

A timber species. The fresh seeds are used for fishing bait (Voorhoeve 1965).

P

Table data are not available.

432

Data sources Taylor (1960), Voorhoeve (1965), Hall & Swaine (1981), Hopkins (1983), Hawthorne (1995a)

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Page 433

Pericopsis elata

(Harms) Van Meeuwen

Leguminosae-Pap. Description Guild: np Life form: large tree Max. height: 45 m (Taylor 1960) Max. diameter: 128 cm (inventory data Ghana) Leaf: alternate, imparipinnately compound, 9 leaflets, oblong to lanceolate (2.5 x 6.5 cm), entire Inflorescence: panicle Flower: medium-sized; corolla white Fruit: dry indehiscent (2.5 x 12 cm), light brown; 1-3 seeds Seed: flat, large (1.3 cm in diameter) Other: a tree, with spreading branches, graceful foliage and a rather flat-topped, triangular crown. The bole is characterised by small, red patches. It has high or no buttresses merging into slight flutes. It has nitrogen-fixing nodules. Wood density is 0.79 g/cm3.

Distribution Continent: Nigeria, Cameroon, Democratic Republic of Congo Upper Guinea: Côte d’Ivoire, Ghana Forest type: moist semi-deciduous forest, dry semi-deciduous forest. A timber tree once common in Brong Ahafo (Hawthorne 1995a), now threatened due to excessive logging. A Red List species (Endangered).

Habitat Seedlings are remarkably drought-tolerant and show little preference between wet and dry forest soils (Swaine & Veenendaal 1994).

Regeneration Germination is normal, or rather rapid (8 days, Taylor 1960). No difference was found between

germination in the light and in the dark (Kyereh et al. 1993). Germination is equally successful in large gaps (Kyereh 1994). Taylor (1960) dismisses the control of regeneration by light as a limiting factor, and does not consider insect damage important. Germination was more than 80% successful in a variety of shaded treatments, but barely 5% successful in full sunlight (Pieters 1958). Out of 7, 18 and 100% of full light, seedlings prefer the middle, equivalent to forest with numerous canopy gaps (Ampofo & Lawson 1972). The ideal sites for seedling growth are where they receive full morning sunlight but are shaded from direct midday sun (Schmitz 1962). In spite of a regular seed production, and no obvious restriction on germination, lack of natural regeneration has often been reported (Aubréville 1938, Taylor 1960).

Growth Growth in suitable conditions is rapid, with trees capable of attaining 8 m (dbh 9 cm) after 7 years and 26 m in 16 years (dbh 97 cm, Howland 1979).

Phenology Dispersal: may be wind-dispersed in strong winds Timing: flowering period from April to May; fruiting period from August to November (Taylor 1960). Years of abundant seed generation have been recorded, but in many fruiting years germination is said to be poor (Howland 1979).

Uses A timber species.

Data sources Aubréville (1938), Pieters (1958), Quist-Arcton (1958), Taylor (1960), Voorhoeve (1965), Ampofo & Lawson (1972), Schmitz (1962), Howland (1979), Halliday & Nakao (1982), Kyereh et al. (1993), Kyereh (1994), Swaine & Veenendaal (1994), Hawthorne (1995a), IUCN Red List (2000)

P

Table data are not available.

433

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Page 434

Petersianthus macrocarpus

(Beauv.) Liben

Lecythidaceae Description

Regeneration

Guild: pi Life form: large tree Max. height: 45 m (Voorhoeve 1965) Max. diameter: 180 cm (Voorhoeve 1965) Leaf: alternate, in terminal tufts, simple, elliptic to ovate, mesophyll (4-7 x 6-16 cm), entire or slightly undulate, herbaceous, scattered hairs on the blade when young, medium green Inflorescence: terminal, not branched (up to 10 cm long) Flower: medium-sized; corolla white to pale green Fruit: dry indehiscent, winged, spindle-shaped (4 cm long, wings up to 3.5 x 7 cm); 1 seed Seed: very large Other: often seen with many red leaves about to fall from the crown. It has small or no buttresses. Wood density is 0.8 g/cm3.

Germination is normal. It has a phanerocotylar epigeal foliaceous seedling type (cf. Voorhoeve 1965). Seedlings are not common due to abortion of much of the seed (Taylor 1960). Most regeneration occurs in ‘slight gaps in the forest’ albeit with some shade tolerance as a seedling (Taylor 1960).

Phenology Dispersal: by wind Timing: flowering is irregular, but often from November to January and April to June (Voorhoeve 1965); fruiting is irregular with two peaks, in November to December and April to May (Taylor 1960)

Distribution Continent: Benin to Angola (Voorhoeve 1965) Upper Guinea: all Upper Guinea (Voorhoeve 1965) Forest type: moist evergreen forest, moist semideciduous forest, secondary forest (Voorhoeve 1965). The only other species in the genus occurs in the Philippines (Liben 1971).

Habitat The abundance increases with rainfall, attains an optimum around 2000 mm/yr, and declines again above 2500 mm/yr. The seedlings occur in the shade (Hall & Swaine 1981). Healthy saplings are associated with small or large gaps (Hawthorne 1995a). It avoids swamps (Voorhoeve 1965).

P

Spp

434

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

P.m.

148

7

24

62

9

4

29

61

9

23

1

72

All sites

198

4

26

57

9

8

31

58

11

25

2

72

Data sources Taylor (1960), Voorhoeve (1965), Liben (1971), Hall & Swaine (1981), Hawthorne (1995a)

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Piptadeniastrum africanum

(Hook.f.) Brenan

Leguminosae-Mim. Description

Habitat

Guild: np Life form: large tree Max. height: 50 m (Voorhoeve 1965) Max. diameter: 180 cm (Voorhoeve 1965) Leaf: alternate, bipinnately compound, 16-60 alternate or subopposite pinnae, each with 48-100 opposite leaflets, linear, leptophyll (0.8-1.2 x 3-8 mm), entire; leaves of saplings and water shoots up to 30 cm long, with slightly larger leaflets Inflorescence: terminal or axillary, spike, up to 10 cm long Flower: small; corolla yellow Fruit: dry dehiscent, flat (2.5 x 25 cm), thin-woody, smooth, brown; up to 12 seeds Seed: flat and winged, with wings 2.5 cm x 6 cm, seed in the middle Other: a very distinctive spreading-crowned tree with smooth, orangeish bark and plank buttresses. The crown starts sub-spherical, develops two distinct layers, and the top layer develops by growing with a strong horizontal component as the lower crown layer dies. Ultimately, the upper crown starts to fragment and displays “crown-shyness”, with gaps between parts of the crown becoming wider, until the tree dies. These patterns could be used to help define the maturity of the tree for use in silvicultural prescriptions. The leaflets fold up at sunset. Wood density is 0.7 g/cm3.

The abundance declines slightly with soil fertility (regression analysis). The tree prefers high rainfall areas and deep, moist, infertile soils (Voorhoeve 1965, Swaine 1996). It becomes a light demander to attain the canopy (Taylor 1960).

Distribution Continent: Benin to Sudan, Angola (Voorhoeve 1965) Upper Guinea: Senegal to Togo (Voorhoeve 1965) Forest type: upland evergreen forest, wet evergreen forest, moist evergreen forest, moist semi-deciduous forest, dry semi-deciduous forest, secondary forest (Hall & Swaine 1981). Ecologically important for many forests, because of its abundance and large crown.

Regeneration Germination is normal, and the seeds have short viability. It has a phanerocotylar epigeal reserve seedling type (cf. Voorhoeve 1965). In many areas, seedlings are very common, even in complete shade (shade is necessary in nursery beds). Saplings are not uncommon under small gaps. Young trees higher than 1.5 m of this species were strongly favoured by gaps (De Klerk 1991).The species tends to have a gap in its population structure at intermediate size classes, indicating that regeneration might be discontinuous. (Poorter et al. 1996, Newbery & Gartlan 1996). In Uganda, it regenerates best in colonising forest and clearings, and survives into older forest (Synnot 1985).

(Hawthorne 1995a) Dispersal: by wind (Taylor 1960) Timing: flowering period from June to August; fruiting period from December to March (Voorhoeve 1965)

Growth Three year old, shaded seedlings may be only 2035 cm tall, whereas in Tropical Shelterwood System plots 4 year old seedlings attain a height of 50-150 cm tall after 4 years (Taylor 1960). In Sierra Leone, mean annual increments of 4 cm over the first 20 years have been recorded (Savill & Fox 1967). Keay (1961) records fast growth in Nigeria, reaching 90 cm dbh in 71 years.

Uses Locally used as timber. The bark is used for treating toothache (Voorhoeve 1965).

Data sources Taylor (1960), Keay (1961), Voorhoeve (1965), Savill & Fox (1967), Hall & Swaine (1981), Synnot (1985), Offermans (1986), De Klerk (1991), Hawthorne (1995a), Newbery & Gartlan (1996), Poorter et al. (1996), Swaine (1996)

Phenology Deciduousness: it is sometimes deciduous, and sometimes drops a lot, but not all of its leaves

P

Spp

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

P.a.

165

7

25

58

8

8

32

58

10

25

2

68

All sites

198

4

26

57

9

8

31

58

11

25

2

72

435

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Page 436

Pycnanthus angolensis

(Welw.) Warb.

Myristicaceae Distribution

Phenology

Continent: Benin to Uganda, Angola (Voorhoeve 1965) Upper Guinea: Guinea to Togo (Voorhoeve 1965) Forest type: upland evergreen forest, wet evergreen forest, moist evergreen forest, moist semi-deciduous forest, dry semi-deciduous forest, secondary forest (Voorhoeve 1965, Hall & Swaine 1981)

Deciduousness: evergreen Dispersal: probably by animals, especially birds. Lieberman et al. (1987) showed that elephants can be effective dispersal agents. It is an important fruit tree for birds (Gautier-Hion & Michaloud 1989). Timing: flowering period from November to April; fruits ripen during the next flowering season (Voorhoeve 1965)

Habitat

Uses

The abundance increases with rainfall, attains an optimum around 2000 mm/yr, to decline strongly after 2600 mm/yr. It is most abundant at infertile soils (regression analysis). Similarly, in Ghana it prefers high rainfall areas and infertile soils (Swaine 1996). It is very drought sensitive as a seedling (Veenendaal & Swaine 1996), but does not grow in swamps (Voorhoeve 1965). It is a light demander but prefers partial shade in youth (Taylor 1960). The tree is a common component of very disturbed forest.

Description Guild: np Life form: medium-sized to large tree Max. height: 35 m (Voorhoeve 1965) Max. diameter: 120 cm (Voorhoeve 1965) Leaf: alternate, simple, narrowly oblong, mesophyll (3-8 x 13-30 cm), entire, coriaceous, young leaves more narrowly elliptic and covered with a dense, rusty brown indumentum of branched hairs, leaves nearly always perforated with numerous holes caused by insects Inflorescence: axillary, panicle (male inflorescence 5-15 cm long, female inflorescence up to 30 cm long) Flower: monoecious; small; both male and female flowers rufous Fruit: dry dehiscent, in large bunches (2.3 x 3 cm), woody; 1 seed Seed: with a red aril Other: an unbuttressed tree with very characteristic, highly untidy looking crown. The boughs are initially in whorls, but all the lesser branches are pendulous on the periphery of the crown, and most leaves are generally heavily insecttattered. Wood density is 0.25 g/cm3.

Regeneration Germination is normal (Hawthorne 1995a). It has a cryptocotylar epigeal reserve seedling type (cf. Voorhoeve 1965). It has a poor germination which may be due to short viability (Savill & Fox 1967). Seedlings are common in complete shade.

Growth Judging from the conspicuousness of the tree in heavily disturbed forest it would seem that it is capable of rapid and healthy ongrowth when large gaps appear in the canopy. Mean annual increments of between 0.6 and 2.4 cm in diameter have been observed in Sierra Leone (Savill & Fox 1967).

P

Spp

436

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

P.a.

170

6

26

57

9

8

31

58

11

24

2

69

All sites

198

4

26

57

9

8

31

58

11

25

2

72

A timber species. The seeds have a very high fat content and can be used as candles.

Data sources Taylor (1960), Voorhoeve (1965), Savill & Fox (1967), Hall & Swaine (1981), Lieberman et al. (1987), Gauthier-Hion & Michaloud (1989), Swaine & Veenendaal (1994), Hawthorne (1995a), Swaine (1996), Veenendaal & Swaine (1996)

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Page 437

Rhodognaphalon brevicuspe

(Sprague) Roberty

Bombacaceae Description

Regeneration

Guild: pi Life form: large tree Max. height: 45 m (Voorhoeve 1965) Max. diameter: 120 cm (Voorhoeve 1965) Leaf: alternate, palmately compound (usually 7 leaflets), obovate, microphyll (1-3.5 x 3-9 cm), entire, glabrous, midrib with long brown hairs, glossy dark green above, paler green with a reddish glow beneath Inflorescence: axillary, flowers solitary or 2-3 together Flower: large; calyx green; corolla white to pinkish red Fruit: dry dehiscent, obovoid (4 x 7 cm), smooth, brown; numerous seeds Seed: medium-sized (0.6 x 0.9 cm), embedded in brightly coloured, reddish-brown kapok Other: a tree with cylindrical, low-buttressed bole. It has heavy thick buttresses and may have prickles when young. The leaf flush is red.

Little is recorded on this, as regeneration is not abundant. It has a phanerocotylar epigeal foliaceous seedling type (cf. Voorhoeve 1965). Seedlings have not been recorded in the shade and are more common in secondary forest than elsewhere.

Distribution Continent: Togo to Gabon (Voorhoeve 1965) Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana (Voorhoeve 1965). Common in Ghana (Hall & Swaine 1981). Red List species (Vulnerable). Forest type: wet evergreen forest, moist evergreen forest, semi-deciduous forest, secondary forest (Voorhoeve 1965, Hall & Swaine 1981)

Growth It is said to be a slow-grower in Sierra Leone, with a mean annual diameter increment of 0.6 cm in logged forest (Savill & Fox 1967).

Phenology Deciduousness: deciduous (Voorhoeve 1965) Dispersal: by wind (Voorhoeve 1965) Timing: flowering period in November; fruiting period from February to March (Voorhoeve 1965). It flowers in the dry season, usually on leafless trees. Mature fruits open on the tree around the beginning of the wet season to release seeds to the wind in cottony kapok (Hawthorne 1995a).

Data sources Taylor (1960), Voorhoeve (1965), Savill & Fox (1967), Hall & Swaine (1981), Hawthorne (1995a), IUCN Red List (2000)

Habitat The species shows a bell-shaped response to rainfall, with an optimal abundance around 1800 mm/yr. The abundance declines with altitude (regression analysis). It is a light-demanding species (Taylor 1960).

R

Spp

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

R.b.

119

3

24

73

3

1

27

66

8

13

2

82

All sites

198

4

26

57

9

8

31

58

11

25

2

72

437

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Page 438

Ricinodendron heudelotii

(Baill.) Pierre ex Pax

Euphorbiaceae Fruit: fleshy (drupe) (3 x 4.5 cm), yellow; 3 seeds Seed: very large (1.9 x 2.3 cm) Other: a tree with weak-wooded branches and a dense crown, sometimes with buttresses. It is selfpruning. Wood density is 0.25 g/cm3.

1993). Seeds of this species were found in 1% (dry season) to 15% (small rainy season) of piles of elephant dung in Bia South GPR (Martin 1991).

Distribution Continent: from Upper Guinea across to east Africa (Hawthorne 1995a) Upper Guinea: Guinea, Sierra Leone, Liberia, Côte d’Ivoire, Ghana Forest type: moist evergreen forest, moist semideciduous forest, dry semi-deciduous forest, secondary forest

Seedlings show at low light levels (2-10%) a strong growth response to irradiance. Growth rates remain constant over the rest of the light range (10-65%; Swaine et al. 1997). Trees can attain about 10 m height in 4 years (Taylor 1960). In the Democratic Republic of Congo it is one of the fastest-growing species, along with Musanga (Hombert 1958). The growth rings in the wood appear to be annual, at least in Nigeria (Amobi 1973).

Habitat

Phenology

The abundance declines strongly with rainfall. It has a preference for soils with a large water holding capacity (regression analysis). Essentially, it is a light demander especially found along timberextraction routes and disturbed habitats (Taylor 1960).

Deciduousness: deciduous (Taylor 1960) Dispersal: by bats (Thompson 1910), hornbills, rodents (Taylor 1960), and elephants (Alexandre 1978) Timing: flowering period in February; fruiting period from July to October (Taylor 1960)

Growth

Regeneration Seeds are stimulated to germinate by exposure to the sun, although the seedlings are subsequently helped by partial shade, because over-exposure encourages leaf curl (MacGregor 1934). However, seeds are capable of germinating in the dark (Kyereh et al. 1993). It has a phanerocotylar epigeal foliaceous seedling type (De la Mensbruge 1966). Seedlings are common in medium-sized to large gaps, yet absent from shaded understorey (Hawthorne 1995a). In Bia South, the seedlings were uncommon in most areas, but occurred in dense thickets along some roadsides, possibly having been dispersed by elephants (Hawthorne

Description Guild: pi Life form: medium-sized tree Max. height: 30 m (Taylor 1960) Max. diameter: 112 cm (inventory data Ghana) Leaf: alternate, palmately compound (5 leaflets), obovate, mesophyll (2.5-12 x 6-20 cm), entire; stipules large, herbaceous and persistent Inflorescence: branched (panicle) Flower: unisexual; corolla white

R

Spp

438

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

R.h.

149

5

29

63

5

3

25

63

12

15

3

76

All sites

198

4

26

57

9

8

31

58

11

25

2

72

Data sources FWTA, Thompson (1910), MacGregor (1934), Hombert (1958), Taylor (1960), De la Mensbruge (1966), Amobi (1973), Alexandre (1978), Hall & Swaine (1981), Martin (1991), Riddoch et al. (1991), Hawthorne (1993, 1995a), Kyereh (1993), Swaine et al. (1997)

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Page 439

Sacoglottis gabonensis

(Baill.) Urb.

Humiriaceae Description Guild: sw Life form: large tree Max. height: 40 m (Voorhoeve 1965) Max. diameter: 180 cm (Voorhoeve 1965) Leaf: alternate, simple, narrowly ovate, elliptic to oblong, notophyll (2.5-6 x 6-15 cm), finely or distinctly crenate, fairly coriaceous, dull medium-green Inflorescence: axillary, branched (dichasia) Flower: medium-sized, corolla white Fruit: fleshy, ellipsoid to subglobose (3 x 3.5 cm), green to yellowish; 1-3 seeds Seed: very large (1.5 x 3 cm) Other: it has high buttresses, a dense, dark crown and a deeply and irregularly fluted and twisted bole. The bark hisses when slashed. The slash may exude sticky amber-brown sap. The wood density is 0.9 g/cm3.

Distribution

Uses

Continent: Benin to Angola (Voorhoeve 1965) Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana, Togo (Voorhoeve 1965). It is very common throughout Liberia (Hawthorne 1995a). Forest type: wet evergreen forest (Hawthorne 1995a)

The bark is used to impart a bitter taste to palmwine, and is an anticoagulant (Madusolumuo et al. 1991).

Data sources

Habitat It is usually a riverine or swampside tree (Keay 1989).

Regeneration It has a phanerocotylar epigeal foliaceous seedling type (cf. Voorhoeve 1965). The natural regeneration is not very abundant as the seeds are often bored by insects (Voorhoeve 1965). Young trees higher than 1.5 m of this species are less strongly favoured by gaps than many other tree species, but are still disproportionately common there (De Klerk 1991).

Taylor (1960), Voorhoeve (1965), Keay (1989), De Klerk (1991), Madusolumuo et al. (1991), Hawthorne (1995a)

Phenology Deciduousness: evergreen (Voorhoeve 1965) Dispersal: by mammals or hornbills. Elephants relish the seeds, and eat them in large quantities. Although this does not improve germination, it makes the seedlings healthier at least in appearance (De Klerk 1991). Timing: flowering period from December to March; fruiting period from September to October (Voorhoeve 1965)

S

Table data are not available.

439

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Page 440

Terminalia ivorensis

A.Chev.

Combretaceae increases with altitude and decreases with soil fertility (regression analysis). The species showed no preference for wet or dry (base-poor or baserich) forest soils, and was not especially droughtsensitive (Swaine & Veenendaal 1994).

Phenology

Description

Regeneration

Guild: pi Life form: large tree Max. height: 45 m (Voorhoeve 1965) Max. diameter: 124 cm (inventory data Ghana) Leaf: alternate in tufts at the end of branchlets, simple, obovate, notophyll (2.5-4.5 x 5-10 cm), entire, coriaceous Inflorescence: axillary, not branched (7-9 cm long) Flower: small; corolla pale yellow Fruit: dry indehiscent, winged (1.8 x 6 cm), densely puberulous, bright brown; 1 seed Seed: elliptic, large (0.8 x 1.5 cm) Other: it has a black bark and a graceful, spreading crown of whorled boughs and clustered leaves. Lower branches “self-clean”, leaving a clear bole even in open conditions. The base of older trees has high, but small buttresses, merging into slight flutes. Wood density is 0.53 g/cm3.

There is no difference between germination in the light and in the dark, and germination is equally successful in large gaps (Kyereh 1994). There is some dormancy that can be overcome partly by soaking (Jones 1968, 1969), probably due to Coumarin-like inhibitors in the fruit (Okoro et al. 1977). Fruit maturation also influences the germination characteristics (Corbineau & Come 1993). It has a phanerocotylar epigeal seedling reserve type (De la Mensbruge 1966). Seedlings are susceptible to drought (Sawyerr 1960, but see Veenendaal & Swaine 1998). Seedlings from larger seeds grow faster than those from small seeds (Oni & Bada 1992). The red/far-red ratio strongly affects the form and leaf area of developing seedlings (Kwesiga & Grace 1986).

Growth

Distribution

Habitat It is a strong light demander, prefers moist conditions and is often found near streams and seasonal swamps (Taylor 1960). Its abundance

T

Spp

440

n

Deciduousness: deciduous (Voorhoeve 1965) Dispersal: by wind (Voorhoeve 1965) Timing: flowering period from May to June; fruiting period from December to January (Voorhoeve 1965). Trees carry ripe fruits towards the end of the dry season, with flowers and new leaves appearing around the start of the rainy season. Flowering in Nigeria is significantly correlated with rainfall, temperature and humidity. Fruits are produced in large quantities (Taylor 1960). Fruit failure, and production of unripe fruits are important factors behind poor regeneration. Natural abscission accounted for half of all fruit abortion, with fungal and insect attack accounting for the other half, in a plantation in Nigeria (Oni 1990).

Uses It is a timber species (Voorhoeve 1965).

Seedlings shows a strong growth response over the lowest part of the irradiance gradient (from 2-10%), whereafter it remains constant (Swaine et al. 1997). It grows rapidly in medium-sized to large gaps, with trees reaching 17 m (25 cm dbh) in 8 years, or 32 cm dbh in 14 years in taungya plantations (Taylor 1960). Lamb and Ntima (1970) recorded 22 year old trees of 36.5 m (76 cm dbh). Horne (1962) recorded annual dbh increments of 1.5 to 2.5 cm in moist forest in Nigeria, and anticipated trees of 90 cm dbh in 50 years. Early optimism in the potential of this species for plantation has been dampened by frequent die-back in monospecific stands. It has been well-used in Ghana for taungya and other

Continent: Benin to Cameroon (Voorhoeve 1965) Upper Guinea: Guinea to Togo (Voorhoeve 1965) Forest type: evergreen forest, moist semideciduous forest, secondary forest (Voorhoeve 1965). A Red List species (Vulnerable).

plantation, but in the mid 1960s these plantations suffered widespread die-back in 30 year old plantations (Annin Bonsu 1968, Ofusu-Asiedu & Cannon 1976).

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

T.i.

127

8

24

58

11

7

32

57

11

26

1

67

All sites

198

4

26

57

9

8

31

58

11

25

2

72

Data sources Sawyerr (1960), Taylor (1960), Horne (1962), Voorhoeve (1965), De la Mensbruge (1966), Annin Bonsu (1968), Jones (1968, 1969), Lamb & Ntima (1970), Ofosu-Asiedu & Cannon (1976), Okoro et al. (1977), Hall & Swaine (1981), Kwesiga & Grace (1986), Oni (1990), Akindele & Owoeye (1991), Oni & Bada (1992), Corbineau & Come (1993), Kyereh et al. (1993), Kyereh (1994), Swaine & Veenendaal (1994), Hawthorne (1995a), Veenendaal & Swaine (1998), Swaine et al. (1997), IUCN Red List (2000)

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Page 441

Terminalia superba

Engl. & Diels

Combretaceae Description Guild: pi Life form: large tree Max. height: 45 m (Voorhoeve 1965) Max. diameter: 150 cm (Voorhoeve 1965) Leaf: alternate in tufts at the end of the branchlets, simple, obovate, notophyll (2.5-7 x 6-12 cm), entire, fairly coriaceous, medium-green Inflorescence: axillary, not branched (7-15 cm long) Flower: small Fruit: dry indehiscent, winged (2 x 5.5 cm), goldenbrown, glabrous; 1 seed Seed: rounded-triangular on cross-section, large (0.7 x 1.5 cm), winged Other: it has strongly whorled boughs and clustered leaves as a consequence of markedly rhythmic growth. It has steep buttresses (up to 3.5 m) when fully grown. Wood density is 0.48 g/cm3.

Distribution Continent: Benin to Angola (Voorhoeve 1965) Upper Guinea: Guinea to Togo (Voorhoeve 1965) Forest type: moist evergreen forest, moist semideciduous forest, dry semi-deciduous forest, secondary forest (Voorhoeve 1965, Hall & Swaine 1981). It tends to occur in disturbed or drier forests across West and Central Africa (Caballe 1978).

Habitat It is a strong light demander, most common in disturbed habitats, exploited forests (Lancaster 1961, Hawthorne 1993), in farmlands and along roads (Taylor 1960). It is not very demanding on soil and water conditions (Taylor 1960). It is most abundant at intermediate altitudes (200-300 m) and intermediate soil fertility. The abundance declines with rainfall (regression analysis). It shows no preference for wet or dry (base-poor or base-rich)

soils, in Ghana (Swaine & Veenendaal 1994). Optimum growth occurs at a pH of 5.9-6.1, and there is a beneficial effect of liming soils (Aluko 1990). However, it is markedly site-sensitive, in southwest Nigeria (Lowe 1973).

Regeneration Germination is, as with T. ivorensis, subject to some dormancy. There is no difference between % germination in the light and in the dark (Kyereh et al. 1993). New seedlings appear at the start of the April-May rainy season, shortly after dispersal, but also during the second peak of rains in OctoberNovember, suggesting dormancy between these batches (Taylor 1960). It has a phanerocotyal epigeal foliaceous seedling type (cf. De la Mensbruge 1966). Young plants are found in the shade (Taylor 1960) and are somewhat shade tolerant (MacGregor 1934). Seedlings and saplings are abundant along roads and in medium-sized to large gaps. Widely used as a plantation species (e.g. Groulez 1961).

Timing: flowering period from February to April; fruiting period from December to February (Voorhoeve 1965)

Growth

Uses

It has a regular bole diameter increment, but periodic growth of crown components (Fay & Fay 1992). Growth at 22°C is continuous, but periodic at 27°C (Maillard et al. 1987, Kwesiga & Grace 1986). Records have been made of about 1.5 m to 3 m height increment in 4 years (Taylor 1960). Trees can attain 14 m height (22 cm dbh) on old logging tracks 4 years after logging (Hawthorne 1993).

It is a timber species (Voorhoeve 1965). Large plantations have been made (Groulez & Wood 1985).

Data sources MacGregor (1934), Taylor (1960), Groulez (1961), Lancaster (1961), Voorhoeve (1965), De la Mensbruge (1966), Lowe (1973), Caballe (1978), Hall & Swaine (1981), Groulez & Wood (1985), Kwesiga & Grace (1986), Maillard et al. (1987), Aluko (1990), Fay & Fay (1992), Hawthorne (1993, 1995a), Kyereh et al. (1993), Kyereh (1994), Swaine & Veenendaal (1994)

Phenology Deciduousness: deciduous (Voorhoeve 1965) Dispersal: by wind (Voorhoeve 1965)

T

Spp

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

T.s.

160

6

28

59

9

4

28

61

11

21

2

71

All sites

198

4

26

57

9

8

31

58

11

25

2

72

441

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Page 442

Tieghemella heckelii

Pierre ex A.Chev.

Sapotaceae Growth It reaches almost 1 m in a year in nurseries, and up to 2 m after 18 months. However, when planted under more light-demanding trees in taungya, it attained only 1.5-3.5 m after 13 years (Taylor 1960). In Nigeria, 22 year old plantations of this species had trees of 30 m and 32 cm in diameter (MacKay 1953).

Phenology Deciduousness: evergreen Dispersal: by animals among which elephants (Alexandre 1978) Timing: flowering period from February to May; fruiting period from October to December (Voorhoeve 1965). In upland evergreen and moist semi-deciduous forests, flowering and fruiting occur throughout the year. In evergreen forests, flowering is from January to February, with fruiting from February to April (Gyimah 1986). Seeds were found in 12% (small rainy season) of piles of elephant dung in Bia South GPR (Martin 1991).

Description Guild: np Life form: large tree Max. height: 55 m (Taylor 1960) Max. diameter: 250 cm (Taylor 1960) Leaf: alternate, simple, elliptic to obovate, notophyll (2-6.5 x 6-15 cm), slightly undulate, coriaceous, glabrous Inflorescence: solitary, paired in the axil of the leaves, sometimes with 4 together Flower: medium-sized; corolla creamy-white Fruit: fleshy, ovoid to subglobose (6.5 x 9 cm), yellow when ripe, smooth; 1-3 seeds Seed: oblong, very large (3.5 x 6.8 cm), surrounded by yellowish pulp Other: an impressively cylindrical tree, with tiers of horizontal branches. Wood density is 0.68 g/cm3.

Habitat The abundance decreases slightly with altitude (regression analysis). It prefers heavy soils and avoids swamps (Voorhoeve 1965). It is a strong light-demander in Sierra Leone (Savill & Fox 1967).

The seeds produce an edible oil (Abbiw 1990, Bonnéhin 2000). It is a good timber species.

Regeneration Germination is fast, and a large proportion of the seeds germinate (Bonnéhin 2000). Seedlings are rare, since the cotyledons are much predated on by rodents. It has a phanerocotylar epigeal reserve seedling type (cf. Voorhoeve 1965). It is shadetolerant when young, but capable of rapid height growth (up to 90 cm/yr) when exposed (Bonnéhin 2000). Seedling growth is fairly constant over a large range of light environments (Swaine et al. 1997).

Distribution Continent: Benin to Nigeria (Voorhoeve 1965) Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana, Togo (Voorhoeve 1965) Forest type: upland evergreen forest, wet evergreen forest, moist evergreen forest, moist semi-deciduous forest. A Red List species (Endangered).

T

Spp

442

n

Uses

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

T.h.

112

5

13

72

10

4

35

56

9

25

2

70

All sites

198

4

26

57

9

8

31

58

11

25

2

72

Data sources MacKay (1953), Taylor (1960), Voorhoeve (1965), Savill & Fox (1967), Alexandre (1978), Hall & Swaine (1981), Gyimah (1986), Abbiw (1990), Martin (1991), Hawthorne (1995a), Swaine et al. (1997), Bonnéhin (2000), Hawthorne & Parren (2000), IUCN Red List (2000)

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Page 443

Triplochiton scleroxylon

K.Schum.

Sterculiaceae Description

Regeneration

Guild: pi Life form: large tree Max. height: 50 m (Voorhoeve 1965) Max. diameter: > 136 cm (inventory data Ghana) Leaf: alternate, palmately compound, 5-7 leaflets, lobed for 1/3 of the length, lobes broadly ovate to triangular, entire Inflorescence: axillary or terminal, branched (panicle, up to 10 cm long) Flower: medium-sized; pinkish white, purple at base Fruit: dry indehiscent, 1-5 free, winged mericarps, 1 seed per mericarp Seed: large (1 x 2 cm), with 4 cm wings Other: buttressing is most common on the tension side of leaning trees, soils under large-buttressed trees were shallower than those under smallbuttressed trees. Clones vary greatly in size and form. Wood density is 0.39 g/cm3.

Germination is not high as for many species (only 55%, Taylor 1960). Normal germination is found in light shade (Lamb 1940). It has a phanerocotylar epigeal foliaceous seedling type (cf. Voorhoeve 1965). Seedlings are not typical of shaded understorey, and it is not clear at which stage the control of this distribution is exercised. In good years, seedlings are very common in gaps of all sizes except the smallest (e.g. Hawthorne 1993).

Distribution Continent: Benin to Gabon (Voorhoeve 1965) Upper Guinea: Sierra Leone to Togo (Voorhoeve 1965) Forest type: moist evergreen forest, moist semideciduous forest, dry semi-deciduous forest

Habitat It is a light demander and a pioneer species (Taylor 1960, Voorhoeve 1965). Its abundance shows a strong decline with rainfall, and an optimum at intermediate altitudes (200-400 m). The species is abundant in areas with an annual rainfall of 11001800 mm and two rainy seasons (Hall & Bada 1979), has a preference for fertile soils (Swaine 1996) and avoids swamps (Voorhoeve 1965). The absence from the Wet Evergreen zone is probably because of its reduced growth in the low fertility soils of this region (Veenendaal et al. 1996).

Growth It can reach 8 m height and 13 cm dbh in 3 years of taungya after planting as stumps (Taylor 1960). On old logging tracks, trees attained 15 m (15 cm dbh) within 4 years (Hawthorne 1993). In Nigeria, mean annual diameter increments of up to 2.5 cm have been recorded in undisturbed forest (Horne 1962). Keay (1989) predicted 42 years to reach 90 cm in diameter. In Nigeria, 50% of the annual increment takes place from mid-April to mid-July (Iyambo 1971). It grows better in mixtures with other species than in monocultures (Lamb 1940).

least in part, for the species range being tied to areas with a little dry season (Jones 1976, Hall & Bada 1979).

Phenology

Uses

Deciduousness: deciduous Dispersal: by wind Timing: flowering period from December to January; fruiting period from January to March (Voorhoeve 1965). It is well-known throughout its range to produce seed very irregularly, both on an annual and on a seasonal basis (e.g. MacKenzie 1959, Lowe 1968, Jones 1974, 1976, Hall & Bada 1979). It flowers in the dry season and fruits around the start of the rains, but with mast years every 4-5 years (Taylor 1960). Unusual, low rainfall periods within the rainfall season may be a stimulus to flowering (MacKenzie 1959), and this accounts, at

It is a timber species.

Data sources Lamb (1940), MacKenzie (1959), Taylor (1960), Horne (1962), Voorhoeve (1965), Lowe (1968), Danso (1970), Iyambo (1971), Johnson (1972), Jones (1974, 1976), Hall & Bada (1979), Hall & Swaine (1981), Keay (1989), Lapido et al. (1991), Hawthorne (1993), Swaine & Veenendaal (1994), Swaine (1996), Veenendaal et al. (1996)

T

Spp

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

T.s.

144

6

31

63

6

0

25

63

12

17

3

76

All sites

198

4

26

57

9

8

31

58

11

25

2

72

443

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Turraeanthus africanus

(Welw. ex C.DC.) Pellegr.

Meliaceae Habitat It is a shade-bearer (Taylor 1960) and avoids the wettest and driest forests (Hawthorne 1995a). It is reported to prefer sandy soils (Voorhoeve 1965).

Regeneration Germination is normal, and usually with a high viability in forest shade. It has a phanerocotylar epigeal reserve seedling type (cf. Voorhoeve 1965). Seedlings are very shade-tolerant, but survival and growth is best under small gaps (Alexandre 1977). There is a tendency for seedlings to occur near parents, but there are fewer seedlings over adult roots, probably due to parasites (Alexandre 1977).

Phenology Deciduousness: evergreen Dispersal: by animals (Alexandre 1977) Timing: flowering period from March to April; fruiting period from August to October (Voorhoeve 1965). In Côte d’Ivoire, the fruits are produced rather irregularly, but usually with two peaks in a year. Only the smaller trees produce fruits.

Description Guild: sb Life form: medium-sized to large tree Max. height: 35 m (Voorhoeve 1965) Max. diameter: 100 cm (Voorhoeve 1965) Leaf: alternate, pinnately compound, 8-24 alternate or subopposite leaflets, narrowly oblong, mesophyll (2-5.5 x 6-25 cm), entire, coriaceous Inflorescence: lateral, branched (panicle, up to 70 cm long) Flower: small; corolla yellow Fruit: capsule, subglobose, 2-5 lobed, orange when ripe; 2-5 seeds Seed: rounded triangular on cross-section, large (1.2 x 2.1 cm), enclosed in a yellow aril Other: usually a low-branched, evergreen tree without buttresses. Wood density is 0.58 g/cm3.

Uses An important timber tree.

Distribution

Data sources

Continent: Benin to Angola (Voorhoeve 1965) Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana, Togo (Voorhoeve 1965) Forest type: upland evergreen forest, moist evergreen forest, moist semi-deciduous forest. A Red List species (Vulnerable).

Taylor (1960), Voorhoeve (1965), Alexandre (1977), Hall & Swaine (1981), Hawthorne (1995a), IUCN Red List (2000)

T

Spp

444

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

T.a.

99

7

13

76

9

2

28

64

8

17

2

78

All sites

198

4

26

57

9

8

31

58

11

25

2

72

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Zanthoxylum gilletii

(De Wild.) P.G.Waterman

Rutaceae Description Guild: pi Life form: large tree Max. height: 35 m (Taylor 1960) Max. diameter: 80 cm (Taylor 1960) Leaf: alternate, imparipinnately compound, approx. 13 opposite or sub-opposite leaflets, oblong to elongate (5 x 14 cm), crenate Inflorescence: branched (panicle) Flower: data unavailable Fruit: capsule, globose (0.5 cm in diameter), dark brown; 1 seed Seed: small, black Other: a tree with large, compound leaves clustered at the end of stout twigs with many pyramidal prickles over the unbuttressed bole.

Distribution Continent: Upper Guinea endemic Upper Guinea: Sierra Leone, Liberia, Côte d’Ivoire, Ghana Forest type: evergreen forest, secondary forest (Taylor 1960)

Habitat

Phenology

It is a strong light demander (Taylor 1960).

Regeneration Germination is rapid, and viability is short (Savill & Fox 1967). Seedlings are commonly seen in gaps, but absent from the shade of undisturbed understorey. Saplings are conspicuous emerging from climber tangles and other patches of low vegetation. They are obvious light demanders, and indicators of secondary forest. This species is one of several pioneer trees dominating at 15 years the

vegetation of large gaps following a wave of dominance by Musanga (Ross 1954). Jones (1974) categorises it as a “fast-growing pioneer”.

Deciduousness: evergreen (Taylor 1960) Dispersal: probably by birds (Hall & Swaine 1981) Timing: flowering period in June; fruiting period from June to July (Taylor 1960)

Growth

Data sources

Trees of 50 cm dbh have been recorded in 40 year old secondary forest, and 105 cm dbh trees in 29 year old forest, in Côte d’Ivoire (Kahn 1982).

Ross (1954), Taylor (1960), Savill & Fox (1967), Jones (1974), Hall & Swaine (1981), Kahn (1982), Hawthorne (1995a)

Spp

n

Altitude

Mean Annual Rainfall

>400m

D

M

W

VW

Soil CMK L

M

H

Soil WHC L

M

H

Z.g.

64

5

28

66

3

3

30

63

8

9

3

84

All sites

198

4

26

57

9

8

31

58

11

25

2

72

Z 445

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C.C.H. Jongkind

Checklist of Upper Guinea forest species

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Introduction

This list contains all names of all the vascular plant species from the forests of Upper Guinea. In total c. 2800 names of different species are included, some with subspecies and varieties. For each of the species a taxonomic publication is given, mostly recent publications with detailed information about the species distribution and ecology. In the case that a species is not cited for Upper Guinea in the literature, a herbarium specimen is cited instead. In the list the herbarium citations are easily distinguished from the literature citations. The collector of the herbarium specimen is written in italic, e.g. name of collector, specimen number, (herbarium lodging the specimen). In the case of a specific name followed by a name in bold, for the species involved we consider the first name as an incorrect one in the way it is used in the publication mentioned with this name. The name in bold is the name we would prefer for that species. Take care: an invalid name in our list can be a correct name for another species in other areas of Africa! The names of the species that are only found in our area (the Upper Guinea endemics) are in green (provided by J.J. Wieringa and co-workers). The species that represent the rare and endemic species selection used in chapter 6 are marked with an asterisk (cf. Wieringa & Poorter, chapter 6). The distribution of a part of these species is given on maps (cf. Holmgren et al. chapter 9), these species are marked with a dark asterisk. Not all species in the list are at first sight obvious forest species. In the case of large trees or lianas of which the seedlings germinate only in the shade, or in the case of herbs that always grow under a dense canopy, this is without doubt. More problematic in this respect are herbs growing on rocks in the streams crossing through the forest (several species of Araceae and Podostemataceae) or trees that start their life in the open (e.g. pioneer species characteristic for large open areas) but usually are found within forest when mature (e.g. Ceiba pentandra, Anogeissis leiocarpa). Other woody plants can grow to large lianas in the forest but can look just as healthy as a dense shrub in tree-savanna (Carissa spinarum, Combretum paniculatum). Also, some epiphytes can be found in the forest-canopy but they also can grow in the mountains on rocks more or less in the open (several species of ferns and orchids).

Calvoa monticola A.Chev. ex Hutch. & Dalziel, Melastomataceae.

For our list we have selected all species which we expect most of the time to be found in forest, near forestborders or in the forest-canopy. Most of these species are likely to disappear or suffer very badly when the forest is removed. We have excluded species that in forest are only found in disturbed areas or open swampy places, and that in most cases are found as weeds on farmland or in savanna vegetation. Rare species from the Fouta Djalon area in Guinea were very hard to judge with respect to being forest species or not. Many of these species grow in a mixed galleryforest and savanna landscape with a very high rainfall. For most collections of these rare species it was not indicated in what kind of vegetation the collected plant was growing. For several of these species the decision to include or exclude the species in the checklist is based on a comparison of their habit, as they appeared in herbarium specimen collections, with better known forest and savanna species.

Literature The starting point for the work on this Checklist was the second edition of the well known Flora of West

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Tropical Africa (FWTA): • Vol. 1 part 1, 1954 {Cycadaceae - Guttiferae}; • Vol. 1 part 2, 1958 {Scytopetalaceae - Umbelliferae}; • Vol. 2, 1963 {Ericaceae - Labiatae}; • Vol. 3 {Monocotyledons} part 1, 1968, part 2, 1972, without Vol. number, 1959, The ferns and fern allies of West Tropical Africa. Almost all new species and other taxonomical changes from later data are included in the Checklist. Without the Enumeration des plantes à fleurs d’Afrique tropicale by J.-P. Lebrun & A.L. Stork (part 1-4, 1991-1997) this work would have been almost impossible. Part of the new species and other neccesary taxonomical changes discovered during the present project still have to be published. They are included in the Checklist and several of them are currently in the process of getting published. An important part of the literature cited in the Checklist can not be found in the common literature list at the end of the Atlas. For the sake of brevity, publications in the most well known periodicals and Flora’s, like the Kew Bulletin, Adansonia or Bulletin du Jardin Botanique National de Belgique are only cited in the Checklist itself. The following abbreviations are used: • Acta Bot.N. – Acta Botanica Neerlandica, Amsterdam, the Netherlands • Acta Phyt. Suec. – Acta Pytogeographica Suecica, Upsala, Sweden • Adansonia – Bulletin du Muséum National d’Histoire Naturelle, Paris, France • Am.J.Bot. – American Journal of Botany, Botanical Society of America, Cornell University, NY, USA • Ann.MBG – Annales of the Missouri Botanical Garden, St Louis, USA • Aroideana, South Miami, Florida, USA • AUWP – see: WAUP • Belg.J.Bot. – Belgian Journal of Botany, Brussels, Belgium • Bib.Bot. – Bibliotheca Botanica, Stuttgart, Germany • BJBB – Bulletin du Jardin Botanique National de Belgique, Meise, Belgium • B.J.Linn.Soc. – Botanical Journal of the Linnean Society of London, UK • Boissiera, Geneve, Switzerland • Bot.Jahrb. – Botanische Jahrbücher, Stuttgart, Germany • Bot.Helv. – Botanica Helvetica, Basel, Switzerland • Bot. Notiser – Botanisca Notiser, Stockholm, Sweden • Brittonia, New York Bot. Gardens, USA • Bull.IFAN – Bulletin de l’Institut (Français) Fondamental d’Afrique Noire, Dakar, Senegal • Bull.MNHN Paris – Bulletin du Muséum National d’Histoire Naturelle, (section B, Adansonia), Paris, France (sér. 2, 3 & 4) • Bull.NHM Lond. – Bulletin of the Natural History Museum London (Bot. Ser.), UK • Bull.Soc.Bot.Fr. – Bulletin Société Botanique de France,

448

Paris, France • Candollea, Geneve, Switzerland • Fl.Afr.Centr. – Flore d’Afrique Centrale (Congo K.Rwanda-Burundi), Meise, Belgium • Fl.Cam. – Flore du Cameroun, Yaoundé, Cameroun • Fl.Gabon – Flore du Gabon, Paris, France • Fl.Guinea-Bis. – Flora da Guiné-Bissao, Lisboa, Portugal • Fl. Malesiana – Flora Malesiana, Leiden, the Netherlands • Fl.Senegal – Flore illustrée de Sénégal, Dakar, Senegal • Fl.Zamb. – Flora Zambesiaca, London • Fragm.Flor.Geobot. – Fragmenta Floristica et Geobotanica, Warszawa, Poland • FTEA – Flora of Tropical East Africa, London • J.W.Afr.SA – Journal of the West African Science Association, Ibadan, Nigeria • Kew Bull. – Kew Bulletin, Royal Botanic Garden, Kew, UK • Kirkia, Harare, Zimbabwe • Med.LHW, Med.LUW – see: WAUP • Mem. IFAN – Memoires de l’Institut Français d’Afrique Noire, Paris, France • Mem.ORSTOM – Memoires ORSTOM, Paris, France • Mitt.Munch. – Mitt. Bot. Staatssammlung München, Germany • Nordic J.Bot. – Nordic Journal of Botany, Copenhagen, Denmark • Norw.J.Bot. – Norwegian Journal of Botany, Oslo, Norway • Novon, Missouri Botanical Garden, St Louis, USA • Opera Bot. – Opera Botanica Belgica, Jardin Botanique National de Belgique, Meise, Belgium • Orch. Monogr. - Orchid Monographs, Leiden, the Netherlands • Plant Systematics and Evolution, Wien, Austria • S.Afr.J.Bot. – South African Journal of Botany, National botanic Gardens, Cape Town, RSA • Symb.Bot.Ups. - Symbolae Botanicae Upsaliensis, Sweden • Taxon, The International Bureau for Plant Taxonomy and Nomenclature • WAUP, WUP – Wageningen (Agricultural) University Papers, Wageningen, the Netherlands • Webbia, Firenze, Italy • Willdenowia, Berlin-Dahlem, Germany

Species Checklist and Genera-Family conversion list The Checklist is ordered alphabetically on families within two groups of plants: Ferns and fern-allies, and Flowering plants. At the end of the Checklist a list of all genera and the plant families they belong to is given. This conversion list facilitates the use of the species checklist.

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Ferns and fern-allies Aspleniaceae Asplenium aethiopicum (Burm.) Bech. (FWTA, Ferns: 59) - africanum Desv. (FWTA, Ferns: 55) - anisophyllum Kunze (Fl.Zamb.: 170) - barteri Hook. (FWTA, Ferns: 56) - blastophorum Hieron. (FWTA, Ferns: 59) - buettneri Hieron. ex Brause (FWTA, Ferns: 59) - cancellatum Alston (FWTA, Ferns: 59) - cuneatum Lam. (FWTA, Ferns: 57) - currori Hook. (FWTA, Ferns: 55) - diplazisorum Hieron. (FWTA, Ferns: 57) - dregeanum Kunze (FWTA, Ferns: 59) - emarginatum P.Beauvois (FWTA, Ferns: 56) - formosum Willd. (FWTA, Ferns: 57) - geppii Carruth. (FWTA, Ferns: 56) = A. anisophyllum - hemitomum Hieron. (FWTA, Ferns: 59) - hypomelas Kuhn (FWTA, Ferns: 60) - inaequilaterale Willd. (Fl.Zamb.: 176) - lividum Mett. Ex Kuhn (FWTA, Ferns: 59) - mannii Hook. (FWTA, Ferns: 60) - megalura Hieron. ex Brause (FWTA, Ferns: 57) - preussii Hieron. ex Brause (FWTA, Ferns: 60) - quintasii Gand. (FWTA, Ferns: 57) ✩ - schnellii Tardieu (FWTA, Ferns: 57) - stuhlmannii Hieron. (FWTA, Ferns: 59) - suppositum Hieron. (FWTA, Ferns: 57) = A. inaequilaterale - unilaterale Lam. (FWTA, Ferns: 56) - variabile (Ballard) Alston (FWTA, Ferns: 56) - warneckei Hieron. (FWTA, Ferns: 59)

Cyatheaceae Cyathea camerooniana Hook. (Kew Bull. 36: 478) - manniana Hook. (Kew Bull. 36: 472)

Davalliaceae Davallia chaerophylloides (Poir.) Steud. (FWTA, Ferns: 53) = D. denticulata - denticulata (Burm.f.) Kuhn (FTEA, Davall: 1)

Dennstaedtiaceae Anisosorus occidentalis (Baker) C.Chr. (FWTA, Ferns: 33) = Lonchitis o. Blotiella currori (Hook.) Tryon (Fl.Zamb.: 84) - mannii (Baker) Pic.Serm. (Kramer & Green 1990) - reducta (C.Chr.) Tryon (Kramer & Green 1990) Lindsaea ensifolia Swartz (FTEA, Dennst: 32) Lonchitis currori (Hook.) Mett. ex Kuhn (FWTA, Ferns: 34) = Blotiella c. - mannii (Baker) Alston (FWTA, Ferns: 34) = Blotiella m. - occidentalis Baker (FTEA, Dennst: 26) - reducta C.Chr. (FWTA, Ferns: 34) = Blotiella r. Microlepia speluncae (L.) Moore (FTEA, Dennst: 2) Pteridium aquilinum (L.) Kuhn (FWTA, Ferns: 33) Schizolegnia ensifolia (Swartz) Alston (FWTA, Ferns: 44) = Lindsaea e.

Dryopteridaceae Ctenitis buchholzii (Kuhn) Alston (FWTA, Ferns: 73) = Triplophyllum b. - cirrhosa (Schum.) Ching (FWTA, Ferns: 71) - efulensis (Baker) Tardieu (FWTA, Ferns: 73) = Lastreopsis e. - jenseniae (C.Chr.) Tardieu (FWTA, Ferns: 71) = Triplophyllum j. - lanigera (Kuhn) Tardieu (FWTA, Ferns: 71) = Triplophyllum vogelii - nigritiana (Mett.) Alston (FWTA, Ferns: 73) = Lastreopsis n. - pilosissima (J.Smith) Alston (FWTA, Ferns: 71) = Triplophyllum pilosissimum - protensa (Afzel. ex Swartz) Ching (FWTA, Ferns: 71) = Triplophyllum protensum - securidiformis (Hook.) Copel. (FWTA, Ferns: 73) = Triplophyllum s. - speciosa (Mett.) Alston (FWTA, Ferns: 71) = Triplophyllum spp. - subsimilis (Hook.) Tardieu (FWTA, Ferns: 73) = Lastreopsis s. Diplazium hylophilum (Hieron.) C.Chr. (FWTA, Ferns: 65) - proliferum (Lam.) Kaulf. (FWTA, Ferns: 65) - sammatii (Kuhn) C.Chr. (FWTA, Ferns: 64) - welwitschii (Hook.) Diels (FWTA, Ferns: 65)

- zanzibaricum (Baker) C.Chr. (Fl.Cam.: 236) Dryopteris athamantica (Kunze) O.Kuntze (FWTA, Ferns: 70) - manniana (Hook.) C.Chr. (FWTA, Ferns: 70) - schnellii Tardieu (FWTA, Ferns: 70) Lastreopsis efulensis (Baker) Tardieu (Fl.Cam.: 284) - nigritiana (Baker) Tindale (Fl.Cam.: 280) - subsimilis (Hook.) Tindale (Fl.Cam.: 278) - vogelii (Hook.) Tindale (Fl.Cam.: 282) = Triplophyllum v. Tectaria angelicifolia (Schum.) Copel. (FWTA, Ferns: 74) - fernandensis (Baker) C.Chr. (FWTA, Ferns: 74) - varians (Moore) C.Chr. (FWTA, Ferns: 73) = Triplophyllum v. Triplophyllum buchholzii (Kuhn) Holttum (Kew Bull. 41: 251) - fraternum (Mett.) Holttum (Kew Bull. 41: 253) - heudelotii Pic.Serm. (Webbia 45: 126) - jenseniae (C.Chr.) Holttum (Kew Bull. 41: 253) - pilosissimum (J.Smith) Holttum (Kew Bull. 41: 246) - securidiformis (Hook.) Holttum (Kew Bull. 41: 242) - subquinquefidum (P.Beauv.) Pic.Serm. (Webbia 45: 129) - troupinii (Pic.Serm.) Holttum (Kew Bull. 41: 243) - varians (Moore) Holttum (Kew Bull. 41: 249) - vogelii (Hook.) Holttum (Kew Bull. 41: 249) - x protensum (Afzel. ex Swartz) Holttum (Webbia 45: 126) = T. heudelotii &T. subquinquefidum

Gleicheniaceae Dicranopteris linearis (Burm.) Underw. (FTEA, Gleich: 6) Gleichenia linearis (Burm.) C.B.Clarke (FWTA, Ferns: 22) = Dicranopteris l.

Grammitidaceae Ctenopteris villosissima (Hook.) Harley (Fl.Cam.: 328) = Xiphopteris v. Xiphopteris oosora (Baker) Alston (FWTA, Ferns: 45) - punctata (Ballard) Alston (FWTA, Ferns: 45) - serrulata (Swartz) Kaulf. (FWTA, Ferns: 45) - villosissima (Hook.) Alston (FWTA, Ferns: 45)

Hymenophyllaceae Hymenophyllum hirsutum (L.) Swartz (FWTA, Ferns: 32) - kuhnii (L.) Swartz (FWTA, Ferns: 32) Trichomanes africanum Christ (FWTA, Ferns: 31) - chamaedrys Taton (FWTA, Ferns: 30) - chevalieri Christ (FWTA, Ferns: 30) - clarenceanum Ballard (FWTA, Ferns: 30) - crispiforme Alston (FWTA, Ferns: 31) - cupressoides Desv. (FWTA, Ferns: 31) - erosum Willd. (FWTA, Ferns: 30) - fallax Christ (FWTA, Ferns: 31) - guineense Afzel. ex Swartz (FWTA, Ferns: 31) - liberiense Copel. (FWTA, Ferns: 30) - mannii Hook. (FWTA, Ferns: 30) - mettenii C.Chr. (FWTA, Ferns: 31)

Lomariopsidaceae Bolbitis acrostichoides (Afzel. ex Swartz) Ching (Hennipman 1977: 149) - auriculata (Lam.) Alston (Hennipman 1977: 136) - fluviatilis (Hook.) Ching (FWTA, Ferns: 68) = B. auriculata - gemmifera (Hieron.) C.Chr. (Hennipman 1977: 263) - heudelotii (Bory ex Fee) Alston (Hennipman 1977: 236) - salicina (Hook.) Ching (Hennipman 1977: 161) Elaphoglossum barteri (Baker) C.Chr. (FWTA, Ferns: 66) - chevalieri Christ (FWTA, Ferns: 66) - cinnamomeum (Baker) Diels (Leeuwenberg 4762 (WAG)) - conforme (Swartz) Schott (FWTA, Ferns: 66) - kuhnii Hieron. (FWTA, Ferns: 66) - salicifolium (Willd. ex Kaulf.) Alston (FWTA, Ferns: 66) Lomariopsis guineensis (Underw.) Alston (FWTA, Ferns: 67) - palustris (Hook.) Mett. ex Kuhn (FWTA, Ferns: 67) - rossii Holttum (FWTA, Ferns: 67)

Lycopodiaceae Huperzia jaegeri (Herter) Pic.Serm. (Webbia 23: 163) - mildbraedii (Herter) Pic.Serm. (Webbia 23: 163) - staudtii (Hessel) Pic.Serm. (Webbia 23: 163) - warneckei (Hessel) Pic.Serm. (Webbia 23: 163) Lycopodiella cernua (L.) Pic.Serm. (Webbia 23: 166) Lycopodium cernuum L. (FWTA, Ferns: 12) = Lycopodiella cernua

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- jaegeri (Herter) Alston (FWTA, Ferns: 11) = Huperzia j. - mildbraedii Herter (FWTA, Ferns: 11) = Huperzia m. - staudtii (Nessel) Adams & Alston (FWTA, Ferns: 12) = Huperzia s. - warneckei (Herter) Alston (FWTA, Ferns: 12) = Huperzia w.

Marattiaceae ✪

Marattia fraxinea J.Smith (FTEA, Maratt: 1) - odontosora Christ. (FWTA, Ferns: 19)

Nephrolepidaceae Nephrolepis biserrata (Swartz) Schott (FWTA, Ferns: 50) - undulata (Afzel. ex Swartz) J.Smith (FWTA, Ferns: 50)

Oleandraceae Arthropteris monocarpa (Cordem.) C.Chr. (FWTA, Ferns: 52) - orientalis (Gmel.) Posth. (FWTA, Ferns: 52) - palisotii (Desv.) Alston (FWTA, Ferns: 52) Oleandra distenta Kunze (FWTA, Ferns: 52) ✩ - ejurana Adams (FWTA, Ferns: 52)

Ophioglossaceae Ophioglossum reticulatum L. (FWTA, Ferns: 18)

Polypodiaceae Belvisia spicata (L.f.) Mirb. (FWTA, Ferns: 48) Drynaria laurentii (Christ) Hieron. (Roos 1985: 266) Loxogramme buettneri (Kuhn) C.Chr. (FWTA, Ferns: 48) - lanceolata (Swartz) Presl (FWTA, Ferns: 48) - latifolia Bonap. (FWTA, Ferns: 48) Microgramma lycopodioides (L.) Copel. (Fl.Zamb.: 155) - owariensis (Desv.) Alton (FWTA, Ferns: 49) = M. lycopodioides Microsorium punctatum (L.) Copel. (FWTA, Ferns: 49) Phymatodes scolopendria (Burm.) Ching (FWTA, Ferns: 48) = Phymatosorus s. Phymatosorus scolopendria (Burm.) Pic.Serm. (Webbia 28: 457) Platycerium angolense Welw. ex Hook. (FWTA, Ferns: 46) = P. elephantotis - elephantotis Schweinf. (Hennipman & Roos 1982: 95) - stemaria (P.Beauv.) Desv. (Hennipman & Roos 1982: 108) Pleopeltis lanceolata (L.) Kaulf. (FWTA, Ferns: 49) - nicklesii (Tardieu) Alston (FWTA, Ferns: 49) - preussii (Hieron) Tardieu (FWTA, Ferns: 49) Pyrrosia mechowii (Hieron.) Alston (FWTA, Ferns: 46) = P. schimperiana - schimperiana (Kuhn) Alston (Hovenkamp 1986: 241)

Psilotaceae Psilotum nudum (L.) Griseb. (FTEA, Psilot: 1)

Pteridaceae Adiantum confine Fée (FWTA, Ferns: 39) - incisum Forssk. (FWTA, Ferns: 39) - philippense L. (FWTA, Ferns: 39) - soboliferum Wall. ex Hook. (FWTA, Ferns: 38) - vogelii Mett. ex Keys. (FWTA, Ferns: 39) Coniogramme africana Hieron. (Leeuwenberg 4720 (WAG)) Doryopteris kirkii (Hook.) Alston (FWTA, Ferns: 43) - nicklesii Tardieu (FWTA, Ferns: 43) Pellaea doniana Hook. (FWTA, Ferns: 43) Pityrogramma calomelanos (L.) Link (FWTA, Ferns: 38) Pteris acanthoneura Alston (FWTA, Ferns: 42) = P. hamulosa - atrovirens Willd. (FWTA, Ferns: 42) - burtonii Baker (FWTA, Ferns: 42) - hamulosa Christ (Fl.Zamb.: 120) - intricata C.H.Wright (FWTA, Ferns: 42) - linearis Poir. (FWTA, Ferns: 42) - marginata Bory (FWTA, Ferns: 42) - mildbraedii Hieron. (FWTA, Ferns: 42) - pteridioides (Hook.) Ballard (FWTA, Ferns: 42) - similis Kuhn (FWTA, Ferns: 42) - togoensis Hieron. (FWTA, Ferns: 40) - vittata L. (FWTA, Ferns: 40)

Schizaeaceae Lygodium microphyllum (Cav.) R.Brown (FWTA, Ferns: 22) - smithianum Presl ex Kuhn (FWTA, Ferns: 22)

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Selaginellaceae Selaginella blepharophylla Alston (FWTA, Ferns: 16) - cathedrifolia Spring (Fl.Afr.Centr.: 26) - goudotiana Springer var. abyssinica (Spring) Bizzarri (Fl.Afr.Centr.: 34) - kalbreyeri Baker (Fl.Afr.Centr.: 23) - kraussiana (Kunze) A.Br. (Fl.Afr.Centr.: 31) - leonensis Hieron. (FWTA, Ferns: 17) - molliceps Spring (Fl.Afr.Centr.: 49) - myosurus (Swartz) Alston (Fl.Afr.Centr.: 14) - soyauxii Hieron. (Fl.Afr.Centr.: 38) - versicolor Spring (Fl.Afr.Centr.: 21) - vogelii Spring (Fl.Afr.Centr.: 18) ✩ - zechii Hieron. (FWTA, Ferns: 17)

Thelypteridaceae Cyclosorus afer (Christ) Ching (FWTA, Ferns: 63) - blastophorus Alston (Geerling & Bokdam 1784 (WAG)) - dentatus (Forssk.) Ching (FWTA, Ferns: 62) - patens (Fée) Copel. (FWTA, Ferns: 62) - quadrangularis (Fée) Tardieu (FWTA, Ferns: 62) - striatus (Schum.) Ching (FWTA, Ferns: 62) Pseudophegopteris cruciata (Willd.) Holttum (Kramer & Green 1990) Thelypteris cruciata (Willd.) Tardieu (FWTA, Ferns: 61) = Pseudophegopteris c. - guineensis (Christ.) Alston (FWTA, Ferns: 61) - microbasis (Baker) Tardieu (FWTA, Ferns: 61) - odontosora (Bonap.) Ching (FWTA, Ferns: 61)

Vittariaceae Antrophyum immersum (Bory ex Willd.) Mett. (FTEA, Vitt: 7) - mannianum Hook. (FTEA, Vitt: 7) Vittaria guineensis Desv. (FTEA, Vitt: 3) - owariensis Fée (FWTA, Ferns: 35)

Flowering plants Acanthaceae ✪ Acanthus guineensis Heine & P.Taylor (FWTA 2: 410) ✩ Adhatoda guineensis Heine (FWTA 2: 423) ✩ - maculata C.B.Clarke (FWTA 2: 422) ✪ - robusta C.B.Clarke (FWTA 2: 422) ✪ Anisotes guineensis Lindau (FWTA 2: 424) ✩ Asystasia amoena Turrill (Kew Bull. 1920: 26) - buettneri Lindau (Lebrun & Stork 4 1997: 468) - calycina Benth. (nom.ill.) (FWTA 2: 413) = A. buettneri ✩ - decipiens Heine (FWTA 2: 413) ✪ - scandens (Lindau) Hook. (FWTA 2: 412) - vogeliana Benth. (FWTA 2: 412) ✩ Barleria brownii S.Moore (FWTA 2: 420) ✩ - maclaudii Benoist (FWTA 2: 420) ✩ - oenotheroides Dum. Cours. (FWTA 2: 420) - opaca (Vahl) Nees (FWTA 2: 421) - ruellioides Anders. (FWTA 2: 420) ✩ Brachystephanus nimbae Heine (Adansonia sér. 2, 11: 650) Brillantaisia lamium (Nees) Benth. (Bull.NHM.Lond.28: 97) ✩ - madagascariensis Lindau (Bull.NHM.Lond.28: 104) - nitens Lindau (FWTA 2: 406) = B. owariensis - owariensis P.Beauv. (Bull.NHM.Lond.28: 90) ✩ - spec.aff. lancifolia Lindau (Morton SL 790 (K, WAG)) - vogeliana (Nees) Benth. (Bull.NHM.Lond.28: 95) ✩ Chlamydocardia buettneri Lindau (FWTA 2: 423) Crossandra buntingii S.Moore (FWTA 2: 409) = Stenandrium b. ✩ - flava Hook. (Kew Bull. 45: 528) - guineensis Nees (FWTA 2: 409) = Stenandrium guineense ✩ - massaica Mildbr. (Kew Bull. 45: 511) ✩ Crossandrella adamii Heine (Adansonia sér. 2, 11: 647) Dicliptera elliotii C.B.Clarke (FWTA 2: 425) ✩ - laxispica Lindau (FWTA 2: 426) ✩ - obanensis S.Moore (FWTA 2: 426)

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✪ Elytraria ivorensis Dokosi (Adansonia sér. 2, 10: 515) - lyrata Vahl (Mitt.Munch.10: 257) - marginata Vahl (Mitt.Munch.10: 257) ✪ - maritima J.K.Morton (Mitt.Munch.10: 257) Endosiphon primuloides T.Anders. ex Benth. (FWTA 2: 398) = Ruellia p. Eremomastax polysperma (Benth.) Dandy (FWTA 2: 397) = E. speciosa - speciosa (Hochst.) Cufod. (Fl.Gabon: 30) ✩ Heteradelphia paulojaegeria Heine (FWTA 2: 397) ✩ Hypoestes consanguinea Lindau (FWTA 2: 431) ✩ - triflora (Forssk.) Roem. & Schult. (FWTA 2: 431) Justicia baronii V.A.W.Graham (Kew Bull. 43: 588) = Adhatoda robusta - biokoensis V.A.W.Graham (Kew Bull. 43: 588) = Adhatoda maculata - extensa T.Anders. (Fl.Gabon: 224) - flava (Forssk.) Vahl (FWTA 2: 427) ✩ - glabra Koen. ex Roxb. (FWTA 2: 428) - insularis T.Anders. (FWTA 2: 427) = J. striata ssp. occidentalis - laxa T.Anders. (FWTA 2: 428) = J. extensa ✩ - striata (Klotzsch) Bullock ssp. occidentalis Morton (Symb.Bot.Ups.29(1): 111) - tenella (Nees) T.Anders. (FWTA 2: 427) Lankesteria brevior C.B.Clarke (FWTA 2: 407) - elegans (P.Beauv.) T.Anders. (FWTA 2: 407) ✩ - hispida (Willd.) T.Anders. (FWTA 2: 408) Lepidagathis alopecuroides (Vahl) R.Br. ex Griseb. (FWTA 2: 414) ✪ Mendoncia combretoides (A.Cheval.) Benoist (FWTA 2: 403) ✩ - gilgiana (Lindau) Benoist var. tisserantii Benoist (FWTA 2: 403) Nelsonia canescens (Lam.) Spreng (FWTA 2: 418) ✩ Phaulopsis angolana S.Moore (Symb.Bot.Ups.31: 99) - barteri (T.Anders.) Lindau (Symb.Bot.Ups.31: 91) - ciliata (Willd.) Hepper (Symb.Bot.Ups.31: 103) - falcisepala C.B.Clarke (FWTA 2: 399) = P. ciliata - imbricata (Forssk.) Sweet (FWTA 2: 399) - silvestris (Lindau) Lindau (FWTA 2: 399) = P. angolana - talbotii S.Moore (Symb.Bot.Ups.31: 162) ✩ Physacanthus batanganus (G.Braun & K.Schum.) Lindau (FWTA 2: 405) ✩ - nematosiphon (Lindau) Rendle & Britten (FWTA 2: 405) ✩ Pseuderanthemum ludovicianum (Buttn.) Lindau (FWTA 2: 421) - tunicatum (Afzel.) Milne-Redh. (FWTA 2: 421) ✪ Pseudocalyx libericus Breteler (Kew Bull. 49: 809) Rhinacanthus virens (Nees) Milne-Redh. var. virens (FWTA 2: 425) ✩ - virens (Nees) Milne-Redh. var. obtusifolia Heine (FWTA 2: 425) Ruellia primuloides (T.Anders. ex Benth.) Heine (Fl.Gabon: 14) ✩ - togoensis (Lindau) Heine (FWTA 2: 397) ✩ Rungia buettneri Lindau (Geerling & Bokdam 1830 (WAG)) ✩ - eriostachya Hua (FWTA 2: 429) - grandis T.Anders. (FWTA 2: 430) ✩ - guineensis Heine (Adansonia sér. 2, 6: 549) ✩ - paxiana (Lindau) C.B.Clarke (FWTA 2: 430) ✩ Sclerochiton vogelii (Nees) T.Anders. var. vogelii (Kew Bull. 46: 42) Stenandriopsis guineensis (Nees) Benoit (Fl.Gabon: 102) = Stenandrium guineense ✩ Staurogyne capitata E.A.Bruce (Bull.Belg. 61: 105) - paludosa (Mangenot & Aké Assi) Heine (FWTA 2: 418) = S. capitata ✩ Stenandrium buntingii (S.Moore) Vollesen (Kew Bull. 47: 179) - guineense (Nees) Vollesen (Kew Bull. 47: 182) ✪ Thomandersia anachoreta Heine (BJBB 36: 239) - laurifolia (T.Anders. ex Benth.) Baill. (FWTA 2: 413) = T. anachoreta Thunbergia chrysops Hook. (FWTA 2: 402) - cynanchifolia Benth. (FWTA 2: 402) - erecta (Benth.) T.Anders. (FWTA 2: 402) ✩ - togoensis Lindau (FWTA 2: 402) - vogeliana Benth. (FWTA 2: 402) ✪ Whitfieldia colorata C.B.Clarke (FWTA 2: 399) ✩ - lateritia Hook. (FWTA 2: 398)

Amaranthaceae Achyranthes bidentata Blume (J.K. Morton SL 355 (K, WAG)) Celosia isertii C.C.Townsend (Fl.Zamb. 9, 1: 34) - laxa Schur & Thonn. (FWTA 1: 147) = C. isertii Cyathula pedicellata C.B.Clarke (FWTA 1: 149) = C. prostrata - prostrata (Linné) Blume (Fl.Zamb. 9, 1: 79) ✩ Sericostachys scandens Gilg & Lopr. (Boissiera 57: 87)

Amaryllidaceae Crinum jagus (Thomps.) Dandy (Adansonia sér. 2, 20: 184) - natans Baker (Adansonia sér. 2, 20: 182) - ornatum (Ait.) Bury (FWTA 3: 134) = C. zeylanicum

- purpurascens Herb. (Adansonia sér. 2, 20: 180) - zeylanicum (Linné) Linné (Adansonia sér. 2, 20: 186) Haemanthus cinnabarinus Decne (FWTA 3: 132) = Scadoxus cinnabarinus - longitubus C.H.Wright (FWTA 3: 132) = Scadoxus multiflorus - multiflorus Martyn (FWTA 3: 132) = Scadoxus multiflorus ✩ Scadoxus cinnabarinus (Decne) Friis & Nordal (Norw.J.Bot. 23: 64) ✪ - multiflorus (Martyn) Raf. ssp. longitubus (Norw.J.Bot. 23: 64)

Anacardiaceae Antrocaryon micraster A.Cheval. & Guillaum. (FWTA 1: 828) ✩ Fegimanra acuminatissima Keay (FWTA 1: 828) ✩ - afzelii Engl. (FWTA 1: 828) Lannea nigritana (Scott-Elliot) Keay var. nigritana (FWTA 1: 733) - nigritana Keay var. pubescens Keay (FWTA 1: 733) - welwitschii (Hiern) Engl. (FWTA 1: 732) Pseudospondias microcarpa (A.Rich.) Engl. var. microcarpa (FWTA 1: 729) ✪ Sorindeia collina Keay (FWTA 1: 737) - juglandifolia (A.Rich.) Planch. ex Oliver (FWTA 1: 737) - warneckei Engl. (FWTA 1: 738) - zenkeri Engl. (Hall & Swaine 1981: 345) Spondias mombin Linné (FWTA 1: 728) Trichoscypha albiflora Engl. (FWTA 1: 736) = T. lucens - arborea (A.Cheval.) A.Cheval. (Adansonia sér. 3, 23: 250) - atropurpurea Engl. (FWTA 1: 736) = T. mannii ✩ - baldwinii Keay (Adansonia sér. 3, 23: 251) ✩ - barbata Breteler (Adansonia sér. 3, 23: 252) - beguei Aubrév. & Pellegr. (FWTA 1: 736) = T. bijuga - bijuga Engl. (Adansonia sér. 3, 23: 252) ✩ - blydeniae Breteler (Adansonia sér. 3, 23: 255) ✩ - cavalliensis Aubrév. & Pellegr. (Adansonia sér. 3, 23: 255) - chevalieri Aubrév. & Pellegr. (FWTA 1: 736) = T. lucens ✪ - laxissima Breteler (Adansonia sér. 3, 23: 256) ✩ - liberica Engl. (Adansonia sér. 3, 23: 256) ✩ - linderi Breteler (Adansonia sér. 3, 23: 258) ✩ - longifolia (Hook.f.) Engl. (Adansonia sér. 3, 23: 258) - lucens Oliver (Adansonia sér. 3, 23: 259) - mannii Hook.f. (Adansonia sér. 3, 23: 261) - oba Aubrév. & Pellegr. (FWTA 1: 736) = T. lucens ✩ - olodiana Breteler (Adansonia sér. 3, 23: 261) - smeathmannii Keay (FWTA 1: 736) = T. smythei ✩ - smythei Hutch. & Dalziel (Adansonia sér. 3, 23: 263) - sp.A. (FWTA 1: 736) = T. barbata - sp.B. (FWTA 1: 736) = T. barbata - yapoensis Aubrév. & Pellegr. (FWTA 1: 736) = T. lucens

Ancistrocladaceae ✩ Ancistrocladus abbreviatus Airy Shaw (FWTA 1: 234) ✩ - barteri Scott-Elliot (FWTA 1: 234) ✩ - guineensis Oliver (FWTA 1: 234) ✪ - pachyrrhachis Airy Shaw (FWTA 1: 234)

Anisophylleaceae ✪ Anisophyllea laurina R.Br. ex Sabine (FWTA 1: 282) ✪ - meniaudi Aubrév. & Pellegr. (Fl.Afr.Centr.: 6)

Annonaceae Annickia polycarpa (A.DC.) Van Setten & Maas (Taxon 39: 676) Anonidium mannii (Oliver) Engl. & Diels (FWTA 1: 51) ✩ Artabotrys hispidus Sprague & Hutch. (FWTA 1: 41) - insignis Engl. & Diels (FWTA 1: 40) ✩ - jollyanus Pierre ex Engl. & Diels (FWTA 1: 40) ✩ - libericus Diels (FWTA 1: 40) ✩ - oliganthus Engl. & Diels (FWTA 1: 40) - stenopetalus Engl. & Diels (FWTA 1: 41) - velutinus Scott-Elliot (FWTA 1: 40) Brieya fasciculata (De Wild.) Paiva (FWTA 1: 39) = Piptostigma fasciculatum Cleistopholis patens (Benth.) Engl. & Diels (FWTA 1: 38) ✪ Dennettia tripetala Baker f. (Boissiera 57: 96) Duguetia barteri (Benth.) Chatrou (Chatrou 1998: 66) - staudtii (Engl. & Diels) Chatrou (Chatrou 1998: 70) Enantia polycarpa (DC.) Engl. & Diels (FWTA 1: 51) = Annickia p. Enneastemon barteri (Baill.) Keay (FWTA 1: 48) = Monanthotaxis b. - capea (E.G. & A.Camus) Ghesq. (FWTA 1: 48) = Monanthotaxis c. - foliosus (Engl. & Diels) Robyns & Ghesq. (FWTA 1: 50) = Monanthotaxis f. - mannii (Baill.) Keay (FWTA 1: 50) = Monanthotaxis m.

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- vogelii (Hook.f.) Keay (FWTA 1: 48) = Monanthotaxis v. Friesodielsia enghiana (Diels) Verdc. (Kew Bull. 25: 19) ✩ - gracilis (Benth.) Steenis (Blumea 12: 359) ✩ - hirsuta (Benth.) Steenis (Kew Bull. 25: 19) ✩ - velutina (Sprague & Hutch.) Steenis (Kew Bull. 25: 20) Greenwayodendron oliveri (Engl.) Verdc. (Adansonia sér. 2, 9: 92) Hexalobus crispiflorus A.Rich. (FWTA 1: 47) - salicifolius Engl. & Diels (Boissiera 57: 98) Isolona campanulata Engl. & Diels (FWTA 1: 53) ✪ - cooperi Hutch. & Dalziel ex Cooper & Record (FWTA 1: 53) ✩ - deightonii Keay (FWTA 1: 53) - dewevrei (De Wild. & Th.Dur.) Engl. & Diels (Boissiera 57: 98) - hexaloba Engl. & Diels (Hall & Swaine 1981: 345) ✩ Mischogyne elliotianum (Engl. & Diels) Fries (Fl.Gabon: 285) Monanthotaxis barteri (Baill.) Verdc. (Kew Bull. 25: 21) ✩ - capea (E.G. & A.Camus) Verdc. (Kew Bull. 25: 21) ✩ - declina (Sprague) Verdc. (Kew Bull. 25: 31) - foliosa (Engl. & Diels) Verdc. (Kew Bull. 25: 21) - klainei (Engl.) Verdc. (Fl.Gabon: 218) - laurentii (De Wild.) Verdc. (Kew Bull. 25: 26) - mannii (Baill.) Verdc. (Kew Bull. 25: 21) - nimbana (Schnell) Verdc. (Kew Bull. 25: 27) - parvifolia (Oliver) Verdc. (Boissiera 57: 100) - schweinfurthii (Engl. & Diels) Verdc. var. tisserantii (Le Thomas) Verdc. (Kew Bull. 25: 22) - stenosepala (Diels) Verdc. (Kew Bull. 25: 29) ✩ - stenosepala aff. spec.nov.? (to be published) - vogelii (Hook.f.) Verdc. (Kew Bull. 25: 23) ✪ - whytei (Stapf) Verdc. (Kew Bull. 25: 29) ✩ Monocyclanthus vignei Keay (FWTA 1: 48) Monodora brevipes Benth. (FWTA 1: 54) - crispata Engl. & Diels (FWTA 1: 54) - myristica (Gaertn.) Dunal (FWTA 1: 54) - tenuifolia Benth. (FWTA 1: 54) Neostenanthera gabonensis (Engl. & Diels) Exell (FWTA 1: 42) ✪ - hamata (Benth.) Exell (FWTA 1: 43) Oxymitra gracilis (Hook.f.) Sprague & Hutch. (FWTA 1: 45) = Friesodielsia g. - hirsuta (Benth.) Sprague & Hutch. (FWTA 1: 45) = Friesodielsia h. - velutina Sprague & Hutch. (FWTA 1: 45) = Friesodielsia v. Pachypodanthium staudtii Engl. & Diels (FWTA 1: 39) = Duguetia s. Piptostigma fasciculatum (De Wild.) Paiva (Fl.Gabon: 126) ✪ - fugax A.Cheval. ex Hutch. & Dalziel (FWTA 1: 39) Polyalthia oliveri Engl. (FWTA 1: 43) = Greenwayodendron o. ✩ Polyceratocarpus parviflorus (Baker f.) Ghesq. (FWTA 1: 45) Popowia congensis (Engl. & Diels) Engl. & Diels (FWTA 1: 44) = Monanthotaxis laurentii - declina Sprague (FWTA 1: 44) = Monanthotaxis d. - klainei Pierre ex Engl. & Diels (Fl.Gabon: 218) = Monanthotaxis k. - mangenotii Sillans (FWTA 1: 757) = Friesodielsia enghiana - nimbana Schnell (FWTA 1: 44) = Monanthotaxis n. - oliverana Exell & Mendonça (FWTA 1: 44) = Monanthotaxis parvifolia - stenosepala Engl. & Diels (FWTA 1: 44) = Monanthotaxis s. - whytei Stapf (FWTA 1: 44) = Monanthotaxis w. Uvaria afzelii Scott-Elliot (FWTA 1: 38) - angolensis Welw. ex Oliver (FWTA 1: 38) - anonoides Baker f. (FWTA 1: 38) ✩ - baumannii Engl. & Diels (FWTA 1: 36) - chamae P.Beauv. (FWTA 1: 36) ✪ - dinklagei Engl. & Diels (FWTA 1: 38) ✩ - doeringii Diels (FWTA 1: 38) - gabonensis Engl. & Diels (Dekker 9 (WAG)) - mocoli Wildem. & Th.Dur. (Hall & Swaine 1981: 313) ✩ - muricata (Pierre) Engl. & Diels (Boissiera 57: 104) ✪ - ovata (Dunal) A.DC. (FWTA 1: 36) ✪ - sassandrensis Jongkind spec. nov. (to be published) ✩ - scabrida Oliver (FWTA 1: 38) ✩ - sofa Scott-Elliot (FWTA 1: 38) ✩ - thomasii Sprague & Hutch. (FWTA 1: 38) ✩ - tortilis A.Cheval. (Boissiera 57: 106) Uvariastrum elliotianum (Engl. & Diels) Sprague & Hutch. (FWTA 1: 47) = Mischogyne e. - insculptum (Engl. & Diels) Sprague & Hutch. (FWTA 1: 47) - pierreanum Engl. (FWTA 1: 47) Uvariodendron angustifolium (Engl. & Diels) R.E.Friis (FWTA 1: 46) - calophyllum R.E.Fries (FWTA 1: 46) - mirabile R.E.Fries (FWTA 1: 46) = U. occidentale

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✩

- occidentale Le Thomas (Adansonia sér. 2, 7: 251) Uvariopsis congensis Robyns & Ghesq. (Boissiera 57: 107) ✪ - globiflora Keay (FWTA 1: 50) ✩ - guineensis Keay (FWTA 1: 50) Xylopia acutiflora (Dunal) A.Rich. (FWTA 1: 42) - aethiopica (Dunal) A.Rich. (FWTA 1: 42) ✪ - elliotii Engl. (FWTA 1: 42) ✩ - aff. hypolampra Mildbr. (Leeuwenberg 3073 (WAG)) - parviflora (A.Rich.) Benth. (FWTA 1: 42) - quintasii Engl. & Diels (FWTA 1: 42) - rubescens Oliver (FWTA 1: 41) - staudtii Engl. & Diels (FWTA 1: 41) ✪ - villosa Chipp (FWTA 1: 42) Xylopiastrum taiense Aubrév. (Aubrév. 1959) = Xylopia sp.? - villosum (Chipp) Aubrév. (Aubrév. 1959) = Xylopia villosa

Apocynaceae Alafia barteri Oliver (Kew Bull. 52: 774) - benthamii (Baill. ex Stapf) Stapf (Kew Bull. 52: 777) - landolphioides (A.DC.) K.Schum. (Kew Bull. 52: 790) ✩ - lucida Stapf (Kew Bull. 52: 794) - multiflora (Stapf) Stapf (Kew Bull. 52: 799) ✪ - parciflora Stapf (Kew Bull. 52: 807) - scandens (Thonn.) De Wild. (FWTA 2: 73) = A. landolphioides ✩ - schumannii Stapf (Kew Bull. 52: 808) ✩ - whytei Stapf (Kew Bull. 52: 813) Alstonia boonei De Wild. (Med.LUW 79-13: 5) - congensis Engl. (FWTA 2: 68) = A. boonei Ancylobothrys amoena Hua (WAUP. 94-3: 5) - scandens (Schum. & Thonn.) Pichon (WAUP. 94-3: 25) Anthoclitandra nitida (Stapf) Pichon (FWTA 2: 58) = Landolphia nitidula Aphanostylis mannii (Stapf) Pierre (FWTA 2: 59) = Landolphia incerta Baissea axillaris (Benth.) Hua (BJBB 64: 94) - baillonii Hua (BJBB 64: 98) - breviloba Stapf (FWTA 2: 79) = B. baillonii - calophylla (K.Schum.) Stapf (FWTA 2: 78) = B. welwitschii ✩ - campanulata (K.Schum.) de Kruif (BJBB 64: 102) ✩ - lane-poolei Stapf (BJBB 64: 110) - leonensis Benth. (BJBB 64: 112, pro parte see B. odorata) - multiflora A.DC. (BJBB 64: 130) - odorata K.Schum. ex Stapf (FWTA 2: 78) = not a synonym of B. leonensis ✩ - welwitschii (Baill.) Stapf ex Hiern (BJBB 64: 154) ✪ - zygodioides (K.Schum.) Stapf (BJBB 64: 163) ✪ Callichilia subsessilis (Benth.) Stapf (Med.LUW 78-7: 26) Carissa edulis Vahl (FWTA 2: 54) = C. spinarum - spinarum Linné (WAUP 2001: 35) Clitandra cymulosa Benth. (BJBB 58: 159) Conopharyngia chippii Stapf (Aubrév. 1959) = Tabernaemontana africana - durissima Stapf (Aubrév. 1959) = Tabernaemontana crassa - jollyana Stapf (Aubrév. 1959) = Tabernaemontana crassa - longiflora Stapf (Aubrév. 1959) = Tabernaemontana africana ✩ Dictyophleba leonensis (Stapf) Pichon (BJBB 59: 208) ✩ - stipulosa (S.Moore ex Wernh.) Pichon (BJBB 59: 223) Farquharia elliptica Stapf (Med.LUW 81-16: 3) Funtumia africana (Benth.) Stapf (Med.LUW 81-16: 16) - elastica (Preuss) Stapf (Med.LUW 81-16: 25) - latifolia (Stapf) Schltr. (Aubrév. 1959) = F. africana Holarrhena africana A.DC. (Aubrév. 1959) = H. floribunda - floribunda (G.Don) Dur. & Schinz. var. tomentella Huber (FWTA 2: 69) = H. floribunda - floribunda (G.Don) Dur. & Schinz. (Med.LUW 81-2: 10) - ovata A.DC. (Aubrév. 1959) = H. floribunda - wulfsbergii Stapf (Aubrév. 1959) = H. floribunda Hunteria eburnea Pichon (FWTA 2: 62) = H. umbellata - elliotii (Stapf) Pichon (FWTA 2: 62) = H. umbellata ✪ - ghanensis J.B.Hall & Leeuwenb. (WAUP 96-1: 102) ✩ - simii (Stapf) H.Huber (WAUP 96-1: 115) - umbellata (K.Schum.) Hallier f. (WAUP 96-1: 118) ✩ Isonema smeathmannii Roem. & Schult. (Med.LUW 83-4: 9) ✩ Landolphia calabarica (Stapf) E.A.Bruce (WAUP 92-2: 34) - dulcis (R.Br. ex Sabine) Pichon var. barteri (Stapf) Pichon (FWTA 2: 57) = L. dulcis - dulcis (R.Br. ex Sabine) Pichon (WAUP 92-2: 53) ✩ - foretiana (Pierre ex Jumelle) Pichon (WAUP 92-2: 70) - heudelotii A.DC. (WAUP 92-2: 84)

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- hirsuta (Hua) Pichon (WAUP 92-2: 90) - incerta (K.Schum.) Persoon (WAUP 92-2: 94) - macrantha (K.Schum.) Pichon (WAUP 92-2: 129) ✩ - membranacea (Stapf) Pichon (WAUP 92-2: 140) ✪ - micrantha (A.Cheval.) Pichon (WAUP 92-2: 144) ✩ - nitidula Persoon (WAUP 92-2: 147) - owariensis P.Beauv. var. leiocalyx & var. owariensis (Pichon) H.Huber (FWTA 2: 56) = L. owariensis - owariensis P.Beauv. (WAUP 92-2: 153) ✪ - togolana (Hallier f.) Pichon (WAUP 92-2: 189) - utilis (A.Cheval.) Pichon (FWTA 2: 56) = hybrid L. dulcis x L. foretiana Malouetia heudelotii A.DC. (AUWP 85-2: 76) Motandra guineensis (Thonn.) A.DC. (Med.LUW 83-7: 5) Oncinotis glabrata (Baill.) Stapf ex Hiern. (AUWP 85-2: 13) - gracilis Stapf (AUWP 85-2: 17) ✩ - nitida Benth. (AUWP 85-2: 25) - pontyi Dubard (AUWP 85-2: 29) ✩ Orthopichonia barteri (Stapf) H.Huber (WAUP 89-4: 31) ✩ - indeniensis (A.Cheval.) H.Huber (WAUP 89-4: 37) ✩ Picralima nitida (Stapf) Th. & H.Durand (WAUP 96-1: 130) Pleiocarpa mutica Benth. (WAUP 96-1: 142) - pycnantha (K.Schum.) Stapf (WAUP 96-1: 146) ✩ Pleioceras afzelii (K.Schum.) Stapf (Med.LUW 83-7: 29) - barteri Baill. (Med.LUW 83-7: 30) Rauvolfia caffra Sonder (BJBB 61: 24) - cumminsii Stapf (FWTA 2: 69) = R. mannii - mannii Stapf (BJBB 61: 38) - vomitoria Afzel. (BJBB 61: 60) Saba comorensis (Bojer) Pichon (BJBB 59: 190) - florida (Benth.) Bullock (FWTA 2: 61) = S. comorensis - senegalensis (A.DC.) Pichon (BJBB 59: 200) ✩ - thompsonii (A.Cheval.) Pichon (BJBB 59: 203) Strophanthus barteri Franch. (Med.LHW 82-4: 42) - gratus (Hook.) Franch. (Med.LHW 82-4: 81) - hispidus DC. (Med.LHW 82-4: 85) - preussii Engl. & Pax (Med.LHW 82-4: 125) - sarmentosus DC. (Med.LHW 82-4: 131) ✪ Tabernaemontana africana A.DC. (Leeuwenberg 1991: 9) - chippii (Stapf) Pichon (FWTA 2: 66) = T. africana - crassa Benth. (Leeuwenberg 1991: 21) ✩ - glandulosa (Stapf) Pichon (Leeuwenberg 1991: 36) - longiflora Benth. (FWTA 2: 66) = T. africana - pachysiphon Stapf var. cumminsii (Stapf) H.Huber (FWTA 2: 66) = T. pachysiphon - pachysiphon Stapf (Leeuwenberg 1991: 51) ✩ - psorocarpa (Pierre ex Stapf) Pichon (Leeuwenberg 1991: 60) ✪ Vahadenia caillei (A.Cheval.) Stapf ex Hutch. & Dalziel (BJBB 63: 321) Voacanga africana Stapf (AUWP 85-3: 12) - bracteata Stapf (AUWP 85-3: 22) ✩ - caudiflora Stapf (AUWP 85-3: 27) - thouarsii Roem. & Schult. var. obtusa (K.Schum) Pichon (Aubrév. 1959) = V. thouarsii - thouarsii Roem. & Schult. (AUWP 85-3: 39)

Aquifoliaceae Ilex mitis (Linné) Radkl. (FWTA 1: 623)

Araceae Amauriella hastifolia (Engl.) Hepper (FWTA 3: 120) = Anubias h. ✩ Amorphophallus barthlotii Itterbach & Lobin (Willdenowia 27: 147) ✪ - baumannii N.E.Br. (Itterbach 1997, Dissertation Univ. Bonn) ✩ - doryphorus Ridl. (Itterbach 1997, Dissertation Univ. Bonn) - flavovirens N.E.Br. (FWTA 3: 118) = A. baumannii ✩ - johnsonii N.E.Br. (FWTA 3: 118) Anchomanes difformis (Blume) Engl. (Fl.Cam. 23) - welwitschii Rendle (FWTA 3: 121) = A. difformis ✪ Anubias afzelii Schott (Med.LHW 79-14: 5) - barteri Schott (Med.LHW 79-14: 8) ✩ - gigantea Cheval. ex Hutch. (Med.LHW 79-14: 21) ✩ - hastifolia Engl. (Med.LHW 79-14: 31) ✩Cercestis afzelii Schott (FWTA 3: 126) ✩ - congensis Engl. (Fl.Cam. 31: 65) - dinklagei Engl. (Fl.Cam. 31: 66) ✩ - ivorensis A.Cheval. (Fl.Cam. 31: 68) - sagittatus Engl. (FWTA 3: 126) = C. dinklagei

- stigmaticus N.E.Br. (FWTA 3: 126) = C. dinklagei - taiensis Bogner & Knecht (Bull. MNHN Paris 16: 331) Culcasia angolensis Welw. ex Schott (FWTA 3: 124) - barombensis N.E.Brown (Fl.Cam. 31: 85) = C. angolensis (oldest name!) - dinklagei Engl. (Fl.Cam. 31: 80) ✪ - glandulosa Hepper (FWTA 3: 126) ✩ - liberica N.E.Br. (FWTA 3: 124) ✩ - parviflora N.E.Br. (Fl.Cam. 31: 84) - saxatilis A.Cheval. (FWTA 3: 124) - scandens P.Beauv. (FWTA 3: 124) - seretii De Wild. (FWTA 3: 126) - striolata Engl. (FWTA 3: 124) - tenuifolia Engl. (Fl.Cam. 31: 90) Cyrtosperma senegalense (Schott) Engl. (FWTA 3: 113) = Lasiomorpha senegalensis Lasimorpha senegalensis Schott (The Genera of Araceae (1997): 138) ✩ Nephthytis afzelii Schott (FWTA 3: 121) ✩ - swainei Bogner (Aroideana 3: 79) Remusatia vivipara (Roxb.) Schott (FWTA 3: 122) ✩ Rhaphidophora africana N.E.Br. (FWTA 3: 114) ✩ Stylochaeton zenkeri Engl. (Fl.Cam. 43)

✩

Araliaceae ✪ Cussonia bancoensis Aubrév. & Pellegr. (FWTA 1: 751) Polyscias fulva (Hiern) Harms (FWTA 1: 750) Schefflera barteri (Seem.) Harms (FWTA 1: 751)

Aristolochiaceae ✩ Aristolochia embergeri Nozeran & N.Hallé (Adansonia sér. 2, 4: 101) Pararistolochia flos-avis (A.Cheval.) Hutch. & Dalziel (FWTA 1: 79) = P. macrocarpa ✪ - goldieana (Hook.f.) Hutch. & Dalziel (Adansonia sér. 2, 17: 484) ✩ - leonensis (Mast.) Hutch. & Dalziel (Adansonia sér. 2, 17: 480) - macrocarpa (Duch.) Poncy (Adansonia sér. 2, 17: 488) ✪ - mannii (Hook.f.) Keay (Adansonia sér. 2, 17: 478) ✩ - promissa (Mast.) Keay (Adansonia sér. 2, 17: 491) ✩ - zenkeri (Engl.) Hutch. & Dalziel (Adansonia sér. 2, 17: 486)

Asclepiadaceae ✩ Anisopus efulensis (N.E.Br.) Goyder (Kew Bull. 49: 743) ✩ - mannii N.E.Br. (Kew Bull. 49: 740) Ceropegia fusiformis N.E.Br. (FWTA 2: 102) - johnsonii N.E.Br. (FWTA 2: 102) - nigra N.E.Br. (FWTA 2: 100) - sankuruensis Schltr. (FWTA 2: 102) ✩ - talbotii S.Moore (FWTA 2: 100) ✩ - tourana A.Cheval. (FWTA 2: 102) ✩ - yorubana Schltr. (FWTA 2: 102) Cryptolepis sanguinolenta (Lindl.) Schltr. (FWTA 2: 83) Cynanchum adalinae (K.Schum.) K.Schum. ssp. adalinae (Ann.MBG 83: 294) ✩ - adalinae (K.Schum.) K.Schum. ssp. mannii (Scott-Elliot) Bullock (Ann.MBG 83: 294) - longipes N.E.Br. (Ann.MBG 83: 323) Dregea abyssinica (Hochst.) K.Schum. (FWTA 2: 97) - crinita (Oliver) Bullock (FWTA 2: 97) ✩ Epistemma assianum D.V.Field & J.B.Hall (Kew Bull. 37: 117) Exolobus patens (Decne) Vourn. (Boissiera 57: 133) Gongronema angolense (N.E.Br.) Bullock (FWTA 2: 98) - latifolium Benth. (FWTA 2: 98) ✪ Mangenotia eburnea Pichon (FWTA 2: 84) Marsdenia magniflora P.T.Li (de Koning 6839 (WAG)) Mondia whitei (Hook.f.) Skeels (FWTA 2: 82) Omphalogonus calophyllus Baill. (S.Afr.J.Bot. 62: 24) Oxystelma bornouense R.Br. (FWTA 2: 90) Parquetina nigrescens (Afzel.) Bullock (FWTA 2: 82) = Periploca & Omphalogonus Pergularia daemia (Forssk.) Chiov. (FWTA 2: 90) Periploca nigrescens Afzel. (S.Afr.J.Bot. 62: 27) Sarcostemma viminale (Linné) R.Br. (FWTA 2: 93) Secamone afzelii (Schult.) K.Schum. (Kew Bull. 47: 464) ✩ - leonensis (Scott-Elliot) N.E.Brown (Kew Bull. 47: 468) ✩ - punctulata Decne (Kew Bull. 47: 446) Tacazzea apiculata Oliver (FWTA 2: 83) Telosma africanum (N.E.Br.) Colville (FWTA 2: 97) Tylophora conspicua N.E.Br. (FWTA 2: 96) - dahomensis K.Schum. (FWTA 2: 96) ✩ - oblonga N.E.Br. (Bot.J.Linn.Soc.133: 197)

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✩

- oculata N.E.Br. (FWTA 2: 96)

Avicenniaceae Avicennia africana P.Beauv. (FWTA 2: 448) = A. germinans - germinans (Linné) Linné (Fl.Cam. 19: 60) - nitida Jacq. (Aubrév. 1959, 3: 234) = A. germinans

Balanitaceae Balanites wilsoniana Dawe & Sprague (FWTA 1: 364)

Balanophoraceae Thonningia sanguinea Vahl (FWTA 1: 667)

Balsaminaceae Impatiens filicornu Hook.f. (Grey-Wilson 1980: 95) - hochstetteri Warb. ssp. jacquesii (Keay) Grey-Wilson (Grey-Wilson 1980: 148) - irvingii Hook.f. ex Oliver (Grey-Wilson 1980: 111) - jacquesii Keay (FWTA 1: 161) = I. hochstetteri ✩ - kamerunensis Warb. ssp. kamerunensis (Grey-Wilson 1980: 93) ✩ - kamerunensis Warb. ssp. obanensis (Keay) Grey-Wilson (Grey-Wilson 1980: 94) ✪ - nzoana A.Cheval. ssp. bennae (Jacques-Félix) Grey-Wilson (Grey-Wilson 1980: 116) ✪ - nzoana A.Cheval. ssp. nzoana (Grey-Wilson 1980: 116) - obanensis Keay (FWTA 1: 162) = I. kamerunensis ssp. o.

Begoniaceae ✪ Begonia cavallyensis A.Cheval. (WUP 01.2: 52) - ciliobracteata Warb. (FWTA 1: 218) - eminii Warb. (WUP 01.2: 71) - fusialata Warb. var. fusialata (WUP 01.2: 89) ✪ - fusicarpa Irmsch. (WUP 01.2: 103) ✪ - hirsutula Hook.f. (WAUP 94-1: 242) - macrocarpa Warb. (FWTA 1: 219) ✩ - mannii Hook. (WUP 01.2: 162) ✪ - mildbraedii Gilg (WAUP 94-1: 259) - oxyloba Welw. ex Hook.f. (BJBB 63: 286) ✩ - polygonoides Hook.f. (WUP 01.2: 187) ✪ - prismatocarpa Hook. ssp. petraea (A. Cheval.) Sosef (WAUP 94-1: 179) ✪ - quadrialata Warb. ssp. nimbaensis Sosef (WAUP 94-1: 189) - quadrialata Warb. ssp. quadrialata (WAUP 94-1: 186) - rostrata Welw. ex Hook.f. (FWTA 1: 219)

Bignoniaceae ✩ Dinklageodoxa scandens Heine & Sandwith (FWTA 2: 385) Kigelia africana (Lam.) Benth. (FWTA 2: 385) Markhamia lutea (Benth.) K.Schum. (FWTA 2: 387) - tomentosa (Benth.) K.Schum. ex Engl. (FWTA 2: 387) Newbouldia laevis (P.Beauv.) Seemann ex Bureau (FWTA 2: 388) Spathodea campanulata P.Beauv. (FWTA 2: 386) Stereospermum acuminatissimum K.Schum. (FWTA 2: 386)

Bombacaceae Bombax brevicuspe Sprague (FWTA 1: 334) = Rhodognaphalon b. - buonopozense P.Beauv. (BJBB 33: 104) Ceiba pentandra (Linné) Gaertn. (FWTA 1: 335) Rhodognaphalon brevicuspe (Sprague) Roberty (BJBB 33: 255)

Boraginaceae Cordia guineensis Schum. & Thonn. (FWTA 2: 320) - millenii Baker (FWTA 2: 320) - platythyrsa Baker (FWTA 2: 321) - senegalensis Juss. (FWTA 2: 320) ● - vignei Hutch. & Dalziel (FWTA 2: 320) Ehretia cymosa Thonn. var. cymosa (FWTA 2: 318) ✪ - trachyphylla C.H.Wright (FWTA 2: 318)

Burmanniaceae Burmannia congesta (Wright) Jonker (FWTA 3: 179) ✪ Gymnosiphon longistylus (Benth.) Hutch. (FWTA 3: 179)

Burseraceae Canarium schweinfurthii Engl. (FWTA 1: 697) ✪ Commiphora dalzielii Hutch. (FWTA 1: 696) Dacryodes klaineana (Pierre) H.J.Lam (FWTA 1: 696) Santiria trimera (Oliver) Aubrév. (FWTA 1: 696)

454

Buxaceae ✪ Buxus acutata Friis (Kew Bull. 44: 293) Notobuxus acuminata (Gilg) Hutch. (FWTA 1: 588) = Buxus a.

Cactaceae Rhipsalis baccifera (J.S.Mill.) Stearn (FWTA 1: 761) - cassutha Gaertn. (FWTA 1: 221) = R. baccifera

Capparaceae Buchholzia coriacea Engl. (Acta Bot.N. 13: 163) ✩ Capparis biloba Hutch. & Dalziel (Fl.Cam. 36) - brassii DC. (Fl.Zamb. 1.1) - erythrocarpos Isert (Fl.Cam. 25) ✩ - tenera Dalzell (Blumea 12: 497) - thonningii Schum. (FWTA 1: 89) = C. brassii - tomentosa Lam. (Fl.Cam.: 32) - viminea Hook.f. & Thomson ex Oliv. (FWTA 1: 89) = C. tenera Crateva adansonii DC. (Fl.Cam.: 68) - religiosa Forst.f. (FWTA 1: 90) = C. adansonii ✪ Euadenia eminens Hook.f. (FWTA 1: 93) - trifoliolata (Schum. & Thonn.) Oliver (Fl.Cam.: 72) Maerua duchesnei (De Wild.) F.White (Fl.Cam.: 86) ✩ Ritchiea afzelii Gilg (FWTA 1: 92) - capparoides (Andr.) Britten (Kirkia 1: 95) - duchesnei (De Wild.) Keay (FWTA 1: 92) = Maerua d. - longipedicellata Gilg (FWTA 1: 92) = R. capparoides ✩ - reflexa (Thonn.) Gild & Benedict (Kirkia 1: 99)

Celastraceae Apodostigma pallens (Planch. ex Oliv.) Wilczek var. pallens (Fl.Cam.: 156) - pallens (Planch. ex Oliv.) Wilczek var. buchholzii (Loesener) Hallé (Fl.Cam.: 160) Bequaertia mucronata (Exell) R.Wilczek (Fl.Cam.: 218) Campylostemon angolense Oliver (Fl.Cam. 226) - laurentii De Wild. (Fl.Cam.: 230) - mucronatum (Exell) J.B.Hall (Kew Bull. 35: 841) - warneckeanum Loesener ex Frisch (Fl.Cam.: 228) Cassine aethiopica Thunb. (Fl.Cam. 19: 12) - buchananii Loesener (FWTA 1: 626) = Elaeodendron b. Cuervea macrophylla (Vahl) R.Wilczek (Fl.Cam.: 184) Elaeodendron buchananii (Loes) Loes (Fl.Cam. 19: 7) Helictonema velutinum (Afzel.) Wilczek ex Hallé (Fl.Cam.: 138) Hippocratea africana Hallé (Willd.) Loesener ex Engl. (FWTA 1: 628)= Loeseneriella a. - atractaspis not published (used by J.B.Hall) = Simirestis a. - clematoides Loesener (FWTA 1: 628) = Loeseneriella c. - dewildemaniana (N.Hallé) J.B. Hall (Kew Bull. 35: 841) - ectypetala not published (used by J.B.Hall) = Loeseneriella e. - guineensis Hutch. & M.B.Moss (FWTA 1: 627) = Loeseneriella apocynoides a. - indica Willd. (FWTA 1: 627) = Reissantia i. - iotricha Loesener (FWTA 1: 628) = Loeseneriella i. - macrophylla Vahl (FWTA 1: 629) = Cuervea m. - mucronata Exell (FWTA 1: 629) = Bequaertia m. - myriantha Oliver (Fl.Cam.: 216) - pallens Planch. ex Oliv. (FWTA 1: 627) = Apodostigma p. - paniculata Vahl (FWTA 1: 627) = Pristimera p. - plumbea Blakelock & Wilczek (Hall & Swaine 1981: 209) = Pristimera p. - rowlandii Loesener (FWTA 1: 628) = Loeseneriella r. - tisserantii not published (J.B.Hall) = Simirestis t. - velutina Afzel. (FWTA 1: 629) = Helictonema velutinum ✪ - vignei Hoyle (Mem. IFAN 64: 124) - welwitschii Oliver (FWTA 1: 628) = Simicratea w. Loeseneriella africana (Willd.) Wilczek ex Hallé (Fl.Cam.: 212) - apocynoides (Welw. ex Oliv.) Hallé ex J.Ra var. guineensis (Hutch. & Moss) N.Hallé (Fl.Cam.: 204) - clematoides (Loesener) R.Wilczek (Fl.Cam.: 196) ✩ - ectypetala N.Hallé (Mem.IFAN 64: 115) - iotricha (Loesener) N.Hallé (Fl.Cam.: 210) - rowlandii (Loesener) N.Hallé (Fl.Cam.: 202) Maytenus buchananii (Loesener) Wilczek (Fl.Cam. 19: 28) - ovatus (Wall. Ex Wight & Arn.) Loesener var. ovatus (FWTA 1: 625) = M. buchananii - undata (Thunb.) Blakelock (Fl.Cam. 19: 18) ✩ Prionostemma unguiculata (Loesener) N.Hallé (Fl.Cam.: 190) Pristimera luteoviridis (Exell) N.Hallé var. luteoviridis (Fl.Cam.: 161) - paniculata (Vahl) N.Hallé (Fl.Cam.: 164)

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- plumbea (Blakelock & Wilczek) N.Hallé (Fl.Cam.: 166) Reissantia indica (Willd.) Mem. IFAN 64 (Fl.Cam.: 152) ✩ Salacia adolfi-friderici Loesener ex Harms (Fl.Cam.: 98) - alata De Wild. var. alata (Fl.Cam.: 94) - baumannii Loesener (N.Hallé 1962: 194) - caillei A.Cheval. ex Hutch. & M.B.Moss (FWTA 1: 633) = S. staudtiana var. leonensis - camerunensis Loesener (FWTA 1: 633) = S. longipes var. camerunensis - cerasifera Welw. ex Oliv. var. cerasifera (Fl.Cam.: 85) - chlorantha Oliver (Fl.Cam.: 124) ✩ - columna N.Hallé (Mem.IFAN 64: 226) - cornifolia Hook.f. (Fl.Cam.: 102) - debilis (G.Don) Walp. (Fl.Cam.: 66) - elegans Welw. Ex Oliv. var. elegans (Fl.Cam.: 116) - erecta (G.Don) Walp. var. erecta (Fl.Cam.: 86) - fimbrisepala Loesener (J.B.Hall) = S. preussii ✪ - howesii Hutch. & M.B.Moss (Fl.Cam.: 108) - ituriensis Loesener (Mem.IFAN 64: 254) = S. preussii ✩ - lateritia N.Hallé (Mem.IFAN 64: 216) - lehmbachii Loesener var. leonensis (Hutch. & Moss) N.Hallé (Fl.Cam.: 54) - lehmbachii Loesener var. aurantiaca (N.Hallé) N.Hallé (Fl.Cam.: 54) - letestui Pellegr. (Fl.Cam.: 36) - lomensis Loesener (FWTA 1: 632) = S. stuhlmanniana - longipes (Oliver) N.Hallé var. camerunensis (Loesener) N.Hallé (Fl.Cam.: 48) ✩ - miegei N.Hallé (Mem. IFAN 64: 207) - nitida (Benth.) N.E.Brown var. nitida (Fl.Cam.: 110) - oliveriana Loesener (Fl.Cam.: 123) - owabiensis Hoyle (Fl.Cam.: 127) - pallescens Oliver (Fl.Cam.: 60) ✩ - preussii Loesener (Fl.Cam.: 35) - pyriformis (Sabine) Steud. (Fl.Cam.: 130) ✩ - senegalensis (Lam.) DC. (Mem.IFAN 64: 221) - sp.C. (FWTA 1: 634) = S. nitida - sp.H. (FWTA 1: 634) = Loeseneriella rowlandii - staudtiana Loesener var. leonensis Loesener (Fl.Cam.: 44) = S. caillei - stuhlmanniana Loesener (Fl.Cam.: 120) = S. lomensis - togoica Loesener (Fl.Cam.: 65) - tuberculata Blakelock var. tuberculata (Fl.Cam.: 93) - uregaensis R.Wilczek var. aurantiaca Hallé (Hall & Swaine 1981: 287) = S. lehmbachii var. aurantiaca - whytei Loesener var. whytei (Fl.Cam.: 73) - zenkeri Loesener (Fl.Cam.: 32) Salacighia letestuana (Pellegr.) Blakelock (Fl.Cam.: 10) ✩ - linderi (Loesener ex Harms) Blakelock (Mem.IFAN 64: 146) Simicratea welwitschii (Oliver) N.Hallé (Fl.Cam.: 140) ✩ Simirestis atractaspis N.Hallé (Bull.MNHN Paris sér. 4, 6: 9) - dewildemaniana N.Hallé (Fl.Cam.: 178) - staudtii (Loesener) N.Hallé (Fl.Cam.: 176) ✩ - tisserantii N.Hallé (Fl.Cam.: 180) ✪ Tristemonanthus nigrisilvae (Mem.IFAN 64) N.Hallé (N.Hallé 1962: 136)

Chrysobalanaceae Acioa barteri (Hook.f. ex Oliv.) Engl. (FWTA 1: 431) = Dactyladenia b. - dinklagei Engl. (FWTA 1: 431) = Dactyladenia d. - hirsuta A.Cheval. ex De Wild. (FWTA 1: 431) = Dactyladenia h. - scabrifolia Hua (FWTA 1: 431) = Dactyladenia s. - unwinii De Wild. (FWTA 1: 431) = Dactyladenia smeathmannii - whytei Stapf (FWTA 1: 431) = Dactyladenia w. Afrolicania elaeosperma Mildbr. (FWTA 1: 427) = Licania e. Chrysobalanus ellipticus Soland. ex Sabine (FWTA 1: 426) = C. icaco subsp. icaco - icaco Linné ssp. atacorensis (A.Chevalier) F.White (BJBB 46: 273) - icaco Linné ssp. icaco (BJBB 46: 275) - orbicularis Schum. (FWTA 1: 426) = C. icaco subsp. icaco Dactyladenia barteri (Hook.f. ex Oliv.) Prance & White (Brittonia 31: 484) ✪ - dinklagei (Engl.) Prance & White (Brittonia 31: 485) ✩ - hirsuta (A.Cheval. ex De Wild.) Prance & White (Brittonia 31: 485) ✩ - scabrifolia (Hua) Prance & White (Brittonia 31: 486) ✪ - smeathmannii (Baill.) Prance & White (Brittonia 31: 486) ✪ - whytei (Stapf) Prance & F.White (Brittonia 31: 486) Hirtella butayei (De Wild.) Brenan (FWTA 1: 430) = Magnistipula b. - cupheiflora (Mildbr.) Mildbr. ex A.Cheval. (FWTA 1: 430) = Magnistipula c. - fleuryana A.Cheval. (FWTA 1: 430) = Magnistipula zenkeri Licania elaeosperma (Mildbr.) Prance & White (BJBB 46: 280) Magnistipula butayei De Wild. ssp. butayei (BJBB 46: 281) ✩ - butayei De Wild. ssp. sargosii (Pellegr.) F.White (BJBB 46: 281)

✪ - cupheiflora Mildbr. ssp. leonensis F.White (BJBB 46: 289) ✪ - zenkeri Engl. (BJBB 46: 293) ✪ Maranthes aubrevillei (Pellegr.) Prance (BJBB 46: 295) ✩ - chrysophylla (Oliver) Prance (BJBB 46: 295) - glabra (Oliver) Prance (BJBB 46: 300) ✩ - kerstingii (Engl.) Prance (BJBB 46: 303) ✩ - robusta (Oliver) Prance (BJBB 46: 306) Neocarya macrophylla (Sabine) Prance (BJBB 46: 308) Parinari aubrevillei Pellegr. (FWTA 1: 428) = Maranthes a. - chrysophylla Oliver (FWTA 1: 428) = Maranthes c. ✩ - congensis F.Didr. (BJBB 46: 321) - excelsa Sabine (BJBB 46: 333) - glabra Oliver (FWTA 1: 428) = Maranthes g. - holstii Engl. (Aubrév. 1959) = P. excelsa - kerstingii Engl. (FWTA 1: 428) = Maranthes k. - robusta Oliver (FWTA 1: 428) = Maranthes r.

Combretaceae Anogeissus leiocarpus (DC.) Guill. & Perr. (FWTA 1: 280) Combretum aphanopetalum Engl. & Diels (Fl.Gabon: 13) ✪ - bipindense Engl. & Diels (FWTA 1: 272) ✪ - blepharopetalum Wickens (Kew Bull. 25: 181) - bracteatum (Lawson) Engl. & Diels (Fl.Gabon: 16) ✪ - calobotrys Engl. & Diels (FWTA 1: 273) - comosum G.Don (Fl.Gabon: 26) - conchipetalum Engl. & Diels (Fl.Gabon: 29) - cuspidatum Planch. ex Benth. (Fl.Gabon: 32) - demeusei De Wild. (FWTA 1: 272) = C. mildbraedii - dolichopetalum Engl. & Diels (FWTA 1: 273) = C. comosum ✩ - fulvum Keay (FWTA 1: 272) - fuscum Planch. ex Benth. (Fl.Gabon: 44) ✪ - grandiflorum G.Don (FWTA 1: 273) - hispidum Lawson (FWTA 1: 274) = C. comosum - homalioides Hutch. & Dalziel (Fl.Gabon: 47) - indicum (Linné) Jongkind (Fl.Gabon: 48) - lecardii Engl. & Diels (FWTA 1: 273) = C. paniculatum - longipilosum Engl. & Diels (Fl.Gabon: 56) - marginatum Engl. & Diels (Fl.Gabon: 58) ✩ - mildbraedii Hutch. & Dalziel (Fl.Cam. 42) - mooreanum Exell (FWTA 1: 274) - mucronatum Schum. (Fl.Gabon: 62) ✩ - multinervium Exell (Fl.Gabon: 63) - oyemense Exell (Fl.Gabon: 69) - paniculatum Vent. (Fl.Gabon: 70) ✩ - paradoxum Lawson (Fl.Gabon: 72) - platypterum (Welw.) Hutch. & Dalziel (Fl.Gabon: 75) - racemosum P.Beauv. (Fl.Gabon: 80) - rhodanthum Engl. & Diels (FWTA 1: 274) = C. comosum - smeathmannii G.Don (FWTA 1: 272) = C. mucronatum - sordidum Exell (Fl.Gabon: 85) ✪ - tarquense J.J.Clark (FWTA 1: 273) ✪ - zenkeri Engl. & Diels (FWTA 1: 273) Laguncularia racemosa Gaertn. (FWTA 1: 281) ✪ Pteleopsis habeensis Aubrév. ex Keay (FWTA 1: 275) - hylodendron Mildbr. (FWTA 1: 275) Quisqualis indica Linné (FWTA 1: 275) = Combretum indicum ✪ Strephonema pseudocola A.Cheval. (Ann.MBG 82: 536) Terminalia ivorensis A.Cheval. (FWTA 1: 279) - superba Engl. & Diels (FWTA 1: 277)

Commelinaceae Aneilema aequinoctiale (P.Beauv.) Kunth (FWTA 3: 30) - beniniense (P.Beauv.) Kunth (FWTA 3: 31) - umbrosum (Vahl) Kunth (FWTA 3: 30) ✪ Buforrestia obovata Brenan (FWTA 3: 40) Coleotrype laurentii K.Schum. (FWTA 3: 35) Commelina africana Linné (FWTA 3: 45) ✩ - ascendens Morton (FWTA 3: 47) - benghalensis Linné var. hirsuta (C.B.Clarke) Morton (FWTA 3: 48) - capitata Benth. (FWTA 3: 47) - congesta C.B.Clarke (FWTA 3: 49) - longiscapa C.B.Clarke (FWTA 3: 47) ✪ - macrosperma J.K.Morton (FWTA 3: 49) - thomasii Hutch. (FWTA 3: 47) Floscopa africana (P.Beauv.) C.B.Clarke (FWTA 3: 28)

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✩ Palisota ambigua (P.Beauv.) C.B.Clarke (Faden personal comm. 2000) - barteri Hook. (FWTA 3: 35) - bracteosa C.B.Clarke (FWTA 3: 35) - hirsuta (Thunb.) K.Schum. (FWTA 3: 35) - sp. (FWTA 3: 35) Pollia condensata C.B.Clarke (FWTA 3: 33) - mannii C.B.Clarke (FWTA 3: 32) Polyspatha hirsuta Mildbr. (FWTA 3: 42) ✩ - oligospatha Faden spec.nov. (to be published) - paniculata Benth. (FWTA 3: 42) Stanfieldiella axillaris Morton (FWTA 3: 24) - imperforata (C.B.Clarke) Brenan (FWTA 3: 23) - oligantha (Mildbr.) Brenan (FWTA 3: 24)

Compositae Chromolaena odorata (Linné) King & H.Rob. (Candollea 48: 179) Crassocephalum biafrae (Oliver & Hiern) S.Moore (FWTA 2: 246) = Solanecio b. Eupatorium odoratum Linné (FWTA 2: 285) = Chromolaena odorata Gynura procumbens (Lour.) Merrill (Kew Bull. 33: 336) - sarmentosa (Blume) DC. (FWTA 2: 243) = G. procumbens Melanthera scandens (Schum. & Thonn.) Roberty (FWTA 2: 240) Microglossa afzelii O.Hoffm. var. afzelii (FWTA 2: 251) - afzelii O.Hoffm. var. serratifolia C.D.Adams (FWTA 2: 251) - pyrifolia (Lam.) O.Ktze. (FWTA 2: 251) Mikania cordata (Burm.f.) B.L.Rob. var. chevalieri C.D.Adams (FWTA 2: 286) - cordata (Burm.f.) B.L.Rob. var. cordata (FWTA 2: 286) Mikaniopsis tedliei (Oliver & Hiern) C.D.Adams (FWTA 2: 243) Solanecio biafrae (Oliver & Hiern) Jeffrey (Kew Bull. 41: 922) Vernonia ampla O.Hoffm. (FWTA 2: 277) = V. myriantha - amygdalina Delile (FWTA 2: 277) ✩ - andohii C.D.Adams (FWTA 2: 277) - biafrae Oliver & Hiern (FWTA 2: 276) - colorata (Willd.) Drake (FWTA 2: 277) - conferta Benth. (FWTA 2: 277) ✩ - doniana DC. (FWTA 2: 277) ✩ - frondosa Oliver & Hiern (FWTA 2: 276) ✩ - myriantha Hook.f. (Boissiera 57: 150) - richardiana (O.Ktze.) P.-Sermolli (FWTA 2: 279) = V. theophrastifolia - theophrastifolia Schweinf. Ex Oliver & Hiern (Kew Bull. 43: 218) ✪ - titanophylla Brenan (Hall & Swaine 1981: 346)

Connaraceae Agelaea macrocarpa Schellenb. (FWTA 1: 746) = A. pentagyna - nitida Soland. ex Planch. (FWTA 1: 746) = A. pentagyna - obliqua (P.Beauv.) Baill. var. obliqua (FWTA 1: 745) = A. pentagyna - obliqua (P.Beauv.) Baill. var. cordata (Schellenb.) Exell (FWTA 1: 746) = A. pentagyna - oligantha Gilg ex Schellenb. (FWTA 1: 746) = A. pentagyna - paradoxa Gilg var. microcarpa Jongkind (WAUP 89-6: 142) - pentagyna (Lam.) Baill. (WAUP 89-6: 144) - trifolia (Lam.) Gilg (FWTA 1: 746) = A. pentagyna Byrsocarpus coccineus Schum. & Thonn. (FWTA 1: 741) = Rourea coccinea Castanola paradoxa (Gilg) Schellenb. (FWTA 1: 746) = Agelaea p. ✩ Cnestis bomiensis Lemmens (WAUP 89-6: 179) - corniculata Lam. (WAUP 89-6: 181) - dinklagei Schellenb. (FWTA 1: 743) = C. corniculata - ferruginea Vahl ex DC. (WAUP 89-6: 196) - grisea Baker (FWTA 1: 743) = C. corniculata - longiflora Schellenb. (FWTA 1: 743) = C. corniculata ✩ - racemosa Don. (WAUP 89-6: 226) Connarus africanus Lam. (WAUP 89-6: 243) ✩ - congolanus Schellenb. (WAUP 89-6: 247) ✩ - thonningii (DC.) Schellenb. (WAUP 89-6: 263) ✪ Hemandradenia chevalieri Stapf (WAUP 89-6: 278) - glomerata Aubrév. & Pellegr. (Aubrév. 1959) = H. mannii ✩ - mannii Stapf (WAUP 89-6: 279) Jaundea baumannii (Gilg) Schellenb. (FWTA 1: 742) = Rourea thomsonii - pinnata (P.Beauv.) Schellenb. (FWTA 1: 742) = Rourea thomsonii Manotes expansa Soland. ex Planch. (WAUP 89-6: 298) - longiflora Baker (FWTA 1: 747) = M. expansa ✪ - macrantha (Gilg) Schellenb. (WAUP 89-6: 306) Rourea coccinea (Thonn. ex Schum.) Benth. (WAUP 89-6: 325) - minor (Gaertn.) Alston (WAUP 89-6: 337) - solanderi Baker (WAUP 89-6: 355) - thomsonii (Baker) Jongkind (WAUP 89-6: 359)

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Santaloides afzelii (R.Br. ex Planch.) Schellenb. (FWTA 1: 746) = Rourea minor - gudjuanum (Gilg) Schellenb. (Aubrév. 1959) = Rourea minor Spiropetalum heterophyllum (Baker) Gilg (FWTA 1: 748) = Rourea solanderi - reynoldsii (Stapf) Schellenb. (FWTA 1: 748) = Rourea solanderi - solanderi (Baker) Gilg (FWTA 1: 748) = Rourea s. - triplinerve Stapf (FWTA 1: 748) = Rourea solanderi

Convolvulaceae Aniseia martinicensis (Jacq.) Choisy (FWTA 2: 343) Bonamia thunbergiana (Roem. & Schult.) F.N.Williams (FWTA 2: 339) ✪ - vignei Hoyle (FWTA 2: 339) Calycobolus africanus (G.Don) Heine (FWTA 2: 338) - heudelotii (Baker ex Oliver) Heine (FWTA 2: 337) ✪ - insignis (Rendle) Heine (FWTA 2: 338) ✩ - parviflorus (Mangenot) Heine (FWTA 2: 338) Ipomoea aitonii Lindl. (FWTA 2: 352) = not including I. arachnosperma (I.dichroa) - alba Linné (FWTA 2: 346) - cairica (Linné) Sweet (FWTA 2: 351) - involucrata P.Beauv. (FWTA 2: 347) - mauritiana Jacq. (FWTA 2: 351) - velutipes Welw. ex Rendle (FWTA 2: 348) Lepistemon owariense (P.Beauv.) Hallier f. (FWTA 2: 343) ✩ - parviflorum Pilger ex Busgen (FWTA 2: 343) Neuropeltis acuminata (P.Beauv.) Benth. (FWTA 2: 338) ✩ - prevosteoides Mangenot (FWTA 2: 338) - velutina Hallier f. (FWTA 2: 338) Operculina macrocarpa (Linné) Urban (FWTA 2: 340) Stictocardia beraviensis (Vatke) Hallier f. (FWTA 2: 352)

Costaceae Costus afer Ker-Gawl. (FWTA 3: 78) - deistelii K.Schum. (FWTA 3: 78) - dubius (Afzel.) K.Schum. (FWTA 3: 78) ✩ - englerianus K.Schum. (FWTA 3: 78) ✩ - littoralis K.Schum. (FWTA 3: 78) - lucanusianus Braun & K.Schum. (FWTA 3: 78) ✩ - schlechteri Winkler (FWTA 3: 78) - sp.A. (FWTA 3: 78) ✪

Cucurbitaceae Adenopus breviflorus Benth. (FWTA 1: 206) = Lagenaria breviflora - guineensis (G.Don) Exell (FWTA 1: 206) = Lagenaria g. - rufus Gilg (FWTA 1: 206) = Lagenaria rufa Cayaponia africana (Hook.f.) Exell (FWTA 1: 206) Coccinia barteri (Hook.f.) Keay (FWTA 1: 215) - grandis Linné (FWTA 1: 215) - keayana R.Fernandes (Jeffrey 1964) (J.W.Afr.SA 9: 88) - sp.A (FWTA 1: 216) = C. keayana - sp.B (FWTA 1: 216) = C. sp. (nov.?) A Jeffrey - sp. (nov.?) A Jeffrey (Jeffrey 1964) (J.W.Afr.SA 9: 87) - subhastata Keraudr. (de Koning 1983: 309) = C. sp. (nov.?) A Jeffrey Dimorphochlamys mannii Hook.f. (FWTA 1: 211) = Momordica cabraei Gerrardanthus paniculatus (Mast.) Cogn. (J.W.Afr.SA 9: 95) - zenkeri Harms & Gilg ex Cogn. (FWTA 1: 208) = G. paniculatus Lagenaria breviflora (Benth.) Roberty (J.W.Afr.SA 9: 90) - guineensis (G.Don) Jeffrey (J.W.Afr.SA 9: 90) - rufa (Gilg) Jeffrey (J.W.Afr.SA 9: 90) Melothria capillacea (Schum. & Thonn.) Cogn. (FWTA 1: 209) = Zehneria c. - cordifolia Hook.f. (FWTA 1: 209) = Zehneria gilletii - deltoidea Benth. (FWTA 1: 209) = Zehneria hallii Momordica angustisepala Harms (J.W.Afr.SA 9: 86) - cabraei (Cogn.) Jeffrey (J.W.Afr.SA 9: 86) - cissoides Planch. ex Benth. (J.W.Afr.SA 9: 86) - foetida Schum. & Thonn. (J.W.Afr.SA 9: 86) ✩ - multiflora Hook.f. (J.W.Afr.SA 9: 86) ✩ - silvatica Jongkind (Blumea 47: 343) Peponium vogelii (Hook.f.) Engl. (J.W.Afr.SA 9: 94) Physedra eglandulosa (Hook.f.) Hutch. & Dalziel (FWTA 1: 214) = Ruthalicia e. - longipes Hook.f. (FWTA 1: 214) = Ruthalicia l. Raphidiocystis caillei Hutch. & Dalziel (FWTA 1: 215) = R. chrysocoma - chrysocoma (Schum.) Jeffrey (BJBB 37: 325) Ruthalicia eglandulosa (Hook.f.) Jeffrey (J.W.Afr.SA 9: 86) ✩ - longipes (Hook.f.) Jeffrey (J.W.Afr.SA 9: 86) Telfaira occidentalis Hook.f. (J.W.Afr.SA 9: 85) Zehneria capillacea (Schum.) Jeffrey (J.W.Afr.SA 9: 93)

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✩

- gilletii (De Willd.) Jeffrey (J.W.Afr.SA 9: 93) - keayana R. & A.Fernandes (J.W.Afr.SA 9: 93)

Cyperaceae Carex neo-chevalieri Kuk. (FWTA 3: 349) Cyperus diffusus Vahl (FWTA 3: 289) - fertilis Boeck. (FWTA 3: 289) - laxus Lam. ssp. sylvestris (Ridley) Lye (Nordic J.Bot. 3, 2: 232) - mapanioides Clarke (FWTA 3: 290) - renschii Boeck. (FWTA 3: 289) Diplacrum capitatum (Willd.) Bock (Flora of the Guianas) - longifolium (Griseb.) Clarke (Fragm.Flor.Geobot. 41: 487) = D. capitatum ✩ Hypolytrum africanum Nees ex Steud. (FWTA 3: 336) ✩ - costatum Nelmes (FWTA 3: 336) - heteromorphum Nelmes (FWTA 3: 336) - heterophyllum Boeck. (Mem.ORSTOM 58: 84) ✩ - poecilolepis Nelmes (FWTA 3: 336) - purpurascens Cherm. (FWTA 3: 336) ✩ - schnellianum Lorougnon (BJBB 45: 181) ✩ - senegalense A.Rich. (FWTA 3: 336) - testui Cherm. (Mem.ORSTOM 58: 84) ✪ Mapania baldwinii Nelmes (Simpson, 1992: 145) ✩ - coriandrum Nelmes (Simpson, 1992: 127) ✪ - ivorensis (Raynal) Raynal (Simpson, 1992: 134) ✩ - liberiensis D.A.Simpson (Simpson, 1992: 133) ✪ - linderi Hutch. & Nelmes (Simpson, 1992: 147) ✩ - mangenotiana Lorougnon (Simpson, 1992: 124) ✪ - minor (Nelmes) Raynal (Simpson, 1992: 128) ✪ - rhynchocarpa Lorougnon & Raynal (Simpson, 1992: 136) Scleria achtenii De Wild. (FWTA 3: 342) - barteri Boeck. (Mem.ORSTOM 58: 27) = S. boivinii - boivinii Steud. (FWTA 3: 340) - depressa (Clarke) Nelmes (FWTA 3: 340) - iostephana Nelmes (FWTA 3: 342) - lithosperma (Linné) Nelmes (FWTA 3: 343) - melanomphala Kunth (FWTA 3: 340) - naumanniana Boeck. (FWTA 3: 342) - nyasensis Clarke (Mem.ORSTOM 58: 29) - pterota Presl (FWTA 3: 342) - racemosa Poir. (Mem.ORSTOM 58: 24) - verrucosa Willd. (FWTA 3: 340) - vogelii Clarke (FWTA 3: 340)

Dichapetalaceae Dichapetalum acutisepalum Engl. (FWTA 1: 438) = D. heudelotii var. heudelotii - albidum A.Cheval. ex Pellegr. (Med.LUW. 73-13: 48) - angolense Chodat (Med.LUW 73-13: 55) ✪ - barteri Engl. (Med.LUW 73-13: 82) ✩ - choristilum Engl. (Med.LUW 78-10: 11) - chrysobalanoides Hutch. & Dalziel (FWTA 1: 438) = D. madagascariense - crassifolium Chodat var. crassifolium (Med.LUW 78-10: 24) - cymulosum (Oliver) Engl. (FWTA 1: 436) = D. filicaule - dewevrei De Wild. & Durand (Med.LUW 78-10: 51) ✪ - dictyospermum Breteler (Med.LUW 78-10: 59) ✪ - filicaule Breteler (Med.LUW 78-10: 71) - guineense (DC.) Keay (FWTA 1: 436) = D. madagascariense - heudelotii (Planch. ex Oliv.) Baill. var. ndongense (Engl.) Breteler (Med.LUW 79-16: 38) - heudelotii (Planch. ex Oliv.) Baill. var. heudelotii (Med.LUW 79-16: 27) - johnstonii Engl. (FWTA 1: 438) = D. heudelotii var. heudelotii - kumasiense Hoyle (FWTA 1: 438) = D. heudelotii var. heudelotii - linderi Hutch. & Dalziel (FWTA 1: 438) = D. heudelotii var. heudelotii ✩ - lofense Breteler (Med.LUW 79-16: 63) - madagascariense Poir. var. madagascariense (Med.LUW 81-10: 13) - martineaui Aubrév. & Pellegr. (FWTA 1: 438) = D. heudelotii var. ndongense - oblongum (Hook.f. ex Benth.) Engl. (Med.LUW 81-10: 56) - pallidum (Oliver) Engl. (Med.LUW 81-10: 65) ✩ - parvifolium Engl. (Med.LUW 81-10: 75) - petersianum Dinklage & Engl. (FWTA 1: 436) = D. angolense - staudtii Engl. (Med.LUW 82-8: 28) ✪ - toxicarium (G.Don) Baill. (Med.LUW 82-8: 57) ✪ Tapura fischeri Engl. (Med.LUW 86-3: 56) ✪ - ivorensis Breteler (Med.LUW 86-3: 63)

✪

Dilleniaceae ✩ Tetracera affinis Hutch. (Mitt. Munch. 8: 79) - alnifolia Willd. (Mitt. Munch. 8: 56) - dinklagei Gilg (FWTA 1: 180) = T. alnifolia - leiocarpa Stapf (Mitt. Munch. 8: 53) ✩ - potatoria Afzel. ex G.Don (Mitt. Munch. 8: 76) - stuhlmanniana Gilg (Mitt. Munch. 8: 83)

Dioncophyllaceae ✩ Habropetalum dawei (Hutch. & Dalziel) Airy Shaw (FWTA 1: 191) ✩ Triphyophyllum peltatum (Hutch. & Dalziel) Airy Shaw (FWTA 1: 194)

Dioscoreaceae Dioscorea abyssinica Hochst. ex Kunth (FWTA 3: 153) - baya De Wild. var. kimpundi De Wild. (Belg.J.Bot. 126: 60) - bulbifera Linné (FWTA 3: 152) ✩ - burkilliana J.Miège (FWTA 3: 153) - dumetorum (Kunth.) Pax (FWTA 3: 151) - hirtiflora Benth. (FWTA 3: 152) - lecardii De Wild. (FWTA 3: 153) = D. sagittifolia - liebrechtsiana De Wild. (FWTA 3: 153) = D. praehensilis ✩ - mangenotiana J.Miège (FWTA 3: 153) - minutiflora Engl. (FWTA 3: 153) - praehensilis Benth. (Belg.J.Bot. 126: 52) ✩ - preussii Pax (FWTA 3: 152) - quartiniana A.Rich. (FWTA 3: 151) - sagittifolia Pax (FWTA 3: 153) - sansibarensis Pax (FWTA 3: 152) - smilacifolia De Wild. (FWTA 3: 153) - togoensis Knuth (FWTA 3: 153) ✩

Dracaenaceae ✩ Dracaena adamii Hepper (AUWP 84-1: 19) - arborea (Willd.) Link (AUWP 84-1: 23) - aubryana Brongn. ex C.J. Morren (AUWP 84-1: 29) ✪ - calocephala Bos (AUWP 84-1: 42) - camerooniana Baker (AUWP 84-1: 45) - cerasifera Hua (AUWP 84-1: 54) ✩ - congoensis Hua (AUWP 84-1: 60) ✪ - cristula W.Bull (AUWP 84-1: 63) - elliotii Baker (FWTA 3: 156) = D. cristula - fragrans (Linné) Ker-Gawl (AUWP 84-1: 69) - mannii Baker (AUWP 84-1: 82) ✩ - mildbraedii K.Krause (AUWP 84-1: 89) ✪ - ovata Ker-Gawl. (AUWP 84-1: 92) - perrottetii Baker (FWTA 3: 157) = D. mannii ✪ - phrynioides Hook. (AUWP 84-1: 97) ✪ - praetermissa Bos (AUWP 84-1: 102) ✪ - scabra Bos (AUWP 84-1: 105) - scoparia A.Cheval. ex Hutch. (FWTA 3: 157) = D. cerasifera - smithii Baker ex Hook.f. (FWTA 3: 156) = D. fragrans - surculosa Lindl. var. surculosa (AUWP 84-1: 111) - surculosa Lindl. var. maculata Hook.f. (AUWP 84-1: 115) - surculosa Lindl. var. capitata Hepper (FWTA 3: 159) = D. surculosa var. maculata

Ebenaceae Diospyros abyssinica (Hiern) F.White (BJBB 48: 287) - barteri Hiern (BJBB 48: 326) - canaliculata De Wild. (BJBB 48: 305) - castaneifolia A.Cheval. (Aubrév. 1959) = D. gabunensis ✪ - chevalieri De Wild. (BJBB 48: 284) ✪ - cooperi (Hutch. & Dalziel) F.White (BJBB 48: 332) - elliotii (Hiern) F.White (BJBB 48: 341) ✩ - feliciana Letouzey & F. White (BJJB 48: 340) ✩ - ferrea (Willd.) Bakh. (BJBB 48: 273) ✩ - gabunensis Gürke (BJBB 48: 317) ✩ - heudelotii Hiern (BJBB 48: 322) - ivorensis Aubrév. & Pellegr. (Aubrév. 1959) = D. mannii - kamerunensis Gürke (BJBB 48: 310) - kekemi Aubrév. & Pellegr. (Aubrév. 1959) = D. viridicans ✩ - liberiensis A.Cheval. ex Hutch. & Dalziel (BJBB 48: 325) - mannii Hiern (BJBB 48: 281) - mespiliformis Hochst. ex A.DC. (BJBB 48: 352) - monbuttensis Gürke (BJBB 48: 300)

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- piscatoria Gürke (BJBB 48: 295) - sanza-minika A.Cheval. (BJBB 48: 308) ✩ - soubreana F.White (BJBB 48: 288) - sp. 2 (BJBB 48: 352) ✩ - thomasii Hutch. & Dalziel (BJBB 48: 324) ✪ - vignei F.White (BJBB 48: 328) - viridicans Hiern (BJBB 48: 296) - xanthochlamys Gürke (Aubrév. 1959) = D. canaliculata Maba abyssinica Hiern (Aubrév. 1959) = Diospyros a. - ferrea (Willd.) Bakhuizen (Aubrév. 1959, 3: 159) = Diospyros f. - gavi Aubrév. & Pellegr. (Aubrév. 1959) = Diospyros cooperi - graboensis Aubrév. (Aubrév. 1959) = Diospyros vignei - soubreana (F.White) Aubrév. (Aubrév. 1959) = Diospyros s.

Erythroxylaceae Erythroxylum emarginatum Thonn. (FWTA 1: 356) - mannii Oliver (FWTA 1: 356)

Euphorbiaceae Acalypha ceraceopunctata Pax (FWTA 1: 409) - neptunica Müll.Arg. (FWTA 1: 410) - racemosa Wall. ex Baill. (FWTA 1: 409) Alchornea cordifolia (Schum. & Thonn.) Muell.Arg. (FWTA 1: 403) - floribunda Müll.Arg. (FWTA 1: 403) - hirtella Benth. (FWTA 1: 403) ✩ Amanoa bracteosa Planch. (FWTA 1: 371) ✩ - strobilacea Müll.Arg. (FWTA 1: 371) Anthostema aubryanum Baill. (FWTA 1: 416) ✩ - senegalense A.Juss. (FWTA 1: 416) Antidesma laciniatum Müll.Arg. var. laciniatum (FWTA 1: 374) - laciniatum Müll.Arg. var. membranaceum Müll.Arg. (FWTA 1: 375) - membranaceum Müll.Arg. (FWTA 1: 375) ✩ - oblonga (Hutch.) Keay (FWTA 1: 375) - rufescens Tulasne (BJBB 58: 9) ✪ Apodiscus chevalieri Hutch. (FWTA 1: 373) Argomuellera macrophylla Pax (FWTA 1: 405) Bridelia atroviridis Müll.Arg. (FWTA 1: 370) - grandis Pierre ex Hutch. (FWTA 1: 370) - micrantha (Hochst.) Baill. (FWTA 1: 370) - stenocarpa Müll.Arg. (FWTA 1: 762) = B. micrantha ✪ Cavacoa baldwinii (Keay & Cavaco) J.Léonard (FWTA 1: 762) Claoxylon hexandrum Müll.Arg. (FWTA 1: 401) = Discoclaoxylon h. Cleidion gabonicum Baill. (FWTA 1: 406) Cleistanthus libericus N.E.Br. (BJBB 30: 434) - polystachyus Hook.f. ex Planch. (FWTA 1: 371) - ripicola J.Léonard (BJBB 30: 438) ✩ Croton aubrevillei J.Léonard (BJBB 28: 113) ✩ - dispar N.E.Br. (FWTA 1: 396) ✩ - leonensis Hutch. (FWTA 1: 394) - macrostachyus Delile (FTEA: 149) - nigritanus Scott-Elliot (FWTA 1: 396) - penduliflorus Hutch. (FWTA 1: 396) - sp. nr. mubango (FWTA 1: 394) = C. aubrevillei - sylvaticus Hochst. ex Krauss (Hall & Swaine 1981: 346) ✩ Crotonogyne caterviflora N.E.Br. (FWTA 1: 400) ✩ - chevalieri (Beille) Keay (FWTA 1: 400) ✩ - manniana Müll.Arg. (FWTA 1: 400) ✩ Crotonogynopsis akeassi J.Léonard (Boissiera 57: 231) Dalechampia ipomoeifolia Benth. (FWTA 1: 412) Discoclaoxylon hexandrum (Muell.Arg.) Pax & K.Hoffm. (Boissiera 57: 232) Discoglypremna caloneura (Pax) Prain (FWTA 1: 403) Drypetes aframensis Hutch. (FWTA 1: 381) ✩ - afzelii (Pax) Hutch (FWTA 1: 382) ✩ - aubrevillei Leandri (FWTA 1: 381) ✩ - aylmeri Hutch. & Dalziel (FWTA 1: 381) - chevalieri Beille (FWTA 1: 382) - floribunda (Muell.Arg.) Hutch (FWTA 1: 381) - gilgiana (Pax) Pax & K.Hoffm. (FWTA 1: 382) ✩ - inaequalis Hutch. (FWTA 1: 382) ✩ - ivorensis Hutch. & Dalziel (FWTA 1: 381) - klainei Pierre ex Pax (FWTA 1: 381) ✩ - leonensis Pax (FWTA 1: 381) - parvifolia Müll.Arg. Pax. & K.Hoffm. (FWTA 1: 382) ✩ - pellegrini Léandri (FWTA 1: 381) - principum (Muell.Arg.) Hutch. (FWTA 1: 381)

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- singroboensis Aké Assi (BJBB 30: 397) - sp. (FWTA 1: 382) = D. singrobaensis Elaeophorbia drupifera (Thonn.) Stapf (FWTA 1: 423) = E. grandifolia - grandifolia (Haw.) Croizat (Hall & Swaine 1981: 193) Erythrococca africana (Baill.) Prain (FWTA 1: 401) - anomala (Juss. ex Poir.) Prain (FWTA 1: 401) Excoecaria guineense (Benth.) Müll.Arg. (BJBB 29: 138) Gelonium ivorense Aubrév. & Pellegr. (Aubrév. 1959) = Suregada ivorensis - occidentale Hoyle (FWTA 1: 412) = Suregada occidentalis Grossera baldwinii Keay & Cavaco (FWTA 1: 399) = Cavacoa b. - vignei Hoyle (FWTA 1: 398) Hymenocardia chevalieri Beille (Aubrév. 1959) = H. heudelotii - heudelotii Müll.Arg. (FWTA 1: 377) ✩ - lyrata Tulasne (FWTA 1: 377) ✩ Keayodendron bridelioides Léandri (BJBB 62: 188) Macaranga barteri Müll.Arg. (FWTA 1: 408) ✩ - beillei Prain (FWTA 1: 407) ✩ - heterophylla (Muell.Arg.) Muell.Arg. (FWTA 1: 407) - heudelotii Baill. (FWTA 1: 408) - hurifolia Beille (FWTA 1: 407) - schweinfurthii Pax (Boissiera 57: 237) - sp.A. (FWTA 1: 408) - spinosa Müll.Arg. (FWTA 1: 408) Maesobotrya barteri (Baill.) Hutch. var. barteri (FWTA 1: 374) - barteri (Baill.) Hutch. var. sparsiflora (Scott-Elliot) Keay (FWTA 1: 374) Mallotus oppositifolius (Geisel.) Müll.Arg. (FWTA 1: 402) - subulatus Müll.Arg. (FWTA 1: 402) Manniophyton fulvum Müll.Arg. (FWTA 1: 400) Mareya micrantha (Benth.) Müll.Arg. (FWTA 1: 404) Margaritaria discoidea (Baill.) Webster (FTEA: 63) Martretia quadricornis Beille (BJBB 59: 326) ✩ Mildbraedia paniculata Pax ssp. occidentalis J. Léonard (BJBB 31: 65) - paniculata Pax (FWTA 1: 397) Necepsia afzelii Prain (BJBB 56: 183) Neoboutonia diaguissensis Beille (FWTA 1: 404) = N. mannii - glabrescens Prain (FWTA 1: 404) = N. mannii - mannii Benth. (Kew Bull. 29: 438) ✩ Oldfieldia africana Benth. & Hook.f. (FWTA 1: 368) ✩ Phyllanthus alpestris Beille (FWTA 1: 387) ✩ - bancilhonae Brunel & Roux (Boissiera 57: 240) - beillei Hutch. (FWTA 1: 388) = P. kerstingii - discoideus (Baill.) Müll.Arg. (FWTA 1: 387) = Margaritaria discoidea ✩ - kerstingii Brunel (Willdenowia 15: 251) ✩ - magnificens Brunel & Roux (Willdenowia 11: 82) - mannianus Müll.Arg. (FWTA 1: 388) = Phyllanthus ssp. - muellerianus (O.Ktze.) Exell (FWTA 1: 385) ✩ - petraeus A.Cheval. ex Beille (Willdenowia 15: 236) ✩ - profusus N.E.Br. (FWTA 1: 387) Protomegabaria macrophylla (Pax) Hutch (FWTA 1: 373) - stapfiana (Beille) Hutch (FWTA 1: 373) ✩ Pseudagrostistachys africana (Muell.Arg.) Pax & K.Hoffm. (Hall & Swaine 1981: 346) ✩ Pycnocoma angustifolia Prain (FWTA 1: 405) - cornuta Müll.Arg. (FWTA 1: 405) - macrophylla Benth. (FWTA 1: 405) Ricinodendron heudelotii (Baill.) Pierre ex Pax (FWTA 1: 393) Sapium aubrevillei Leandri (FWTA 1: 415) = Shirakiopsis a. - carterianum J.Léonard (BJBB 29: 146) = Sclerocroton carterianus - cornutum Pax (FWTA 1: 415) = Sclerocroton cornutum - ellipticum (Hochst.) Pax (FWTA 1: 415) = Excoecaria e. - guineense (Benth.) O.Ktze. (FWTA 1: 415) = Excoecaria g. Sclerocroton carterianus (Léonard) Kruijt & Roebers (Bib.Bot. 146: 18) - cornutum (Pax) Kruijt & Roebers (Bib.Bot. 146: 20) ✪ Shirakiopsis aubrevillei (Leandri) Esser (Kew Bull. 56: 1018) - elliptica (Hochst.) Esser (Kew Bull. 56: 1018) Spondianthus preussii Engl. var. preussii (BJBB 59: 140) - preussii Engl. var. glaber (Engl.) Engl. (BJBB 59: 143) ✪ Suregada ivorensis (Aubrév. & Pellegr.) J.Léonard (BJBB 28: 449) - occidentalis (Hoyle) Croizat (BJBB 28: 449) Tetrorchidium didymostemon (Baill.) Pax & K.Hoffm. (Adansonia sér. 3, 21: 100) - minus (Prain) Pax & K.Hoffm. (FWTA 1: 414) = T. didymostemon ✩ - oppositifolium (Pax) Pax & K.Hoffm. (Adansonia sér. 3,21: 103) Thecacoris stenopetala (Muell.Arg.) Muell.Arg. (FWTA 1: 372) Tragia benthamii Baker (FWTA 1: 412) - chevalieri Beille (FWTA 1: 412) ✩ - laminularis Müll.Arg. (FWTA 1: 412) - mildbraediana Pax & K. Hoffm. (Jongkind et al. 2533 (WAG))

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- polygonoides Prain (FWTA 1: 412) - sp.A (FWTA 1: 412) = T. mildbraediana - spathulata Benth. (FWTA 1: 412) - tenuifolia Benth. (FWTA 1: 412) - volubilis Linné (FWTA 1: 411) ✪ Uapaca chevalieri Beille (FWTA 1: 390) - corbisieri De Wild. (Hall & Swaine 1981: 313) - esculenta A.Cheval. ex Aubrév. & Leandri (FWTA 1: 392) = U. corbisieri - guineensis Müll.Arg. (FWTA 1: 390) - heudelotii Baill. (FWTA 1: 390) ✪ - paludosa Aubrév. & Leandri (FWTA 1: 390) - togoensis Pax (FWTA 1: 390)

Gentianaceae (see Am.J.Bot. 87: 1029) Anthocleista djalonensis A.Cheval. (FWTA 2: 35) - microphylla Wernh. (Hall & Swaine 1981: 96) ✪ - nobilis G.Don (FWTA 2: 35) - procera Lepr. ex Bureau (FWTA 2: 37) - vogelii Planch. (FWTA 2: 35) Sebaea oligantha (Gilg) Schinz (Adansonia sér. 2, 7: 211) Voyria platypetala Baker (FWTA 2: 302) = V. primuloides - primuloides Baker (Adansonia sér. 2, 7: 67)

Gesneriaceae Epithema tenue C.B.Clarke (FWTA 2: 383) Streptocarpus nobilis C.B.Clarke (FWTA 2: 382)

Flacourtiaceae ✪ Byrsanthus brownii Guillaumet (FWTA 1: 197) = not synonym with B. epigynus from central Africa Caloncoba brevipes (Stapf) Gilg (FWTA 1: 188) = Oncoba brevipes - echinata (Oliver) Gilg (FWTA 1: 188) = Oncoba e. - gilgiana (Sprague) Gilg (FWTA 1: 189) = Oncoba gilgiana Casearia barteri Mast. (BJBB 41: 406) - bridelioides Mildbr. ex Hutch. & Dalziel (FWTA 1: 198) = Keayodendron b. ✩ - calodendron Gilg (BJBB 41: 416) - dinklagei Gilg (FWTA 1: 198) = C. barteri - inaequalis Hutch. & Dalziel (FWTA 1: 198) = C. calodendron ✩ - stipitata Mast. (Fl.Gabon: 19) Dasylepis assinensis A.Cheval. (FWTA 1: 186) = D. racemosa - blackii (Oliver) Chipp (Fl.Gabon: 30) = D. racemosa - brevipedicellata Chipp (FWTA 1: 186) = D. racemosa - racemosa Oliver (according to F.J.Breteler, to be published) ✪ Dissomeria crenata Hook.f. ex Benth. (FWTA 1: 194) Dovyalis afzelii Gilg (FWTA 1: 190) = D. zenkeri - sp.A. (FWTA 1: 190) = D. zenkeri - sp.B. (FWTA 1: 190) = D. zenkeri - sp.C. (FWTA 1: 190) = D. zenkeri - zenkeri Gilg (Fl.Gabon: 10) Flacourtia flavescens Willd. (FWTA 1: 189) = F. indica - indica (Burm.f.) Merr. (Fl.Gabon: 7) - vogelii Hook.f. (FWTA 1: 189) Homalium africanum (Hook.f.) Benth. (Fl.Gabon: 67) ✩ - angustifolium Sm. (BJBB 43: 310) - angustistipulatum Keay (FWTA 1: 196) = H. dewevrei - aubrevillei Keay (FWTA 1: 196) = H. smythei - aylmeri Hutch. & Dalziel (FWTA 1: 196) = H. longistylum ✩ - dewevrei De Wild. & Durand (Fl.Gabon: 72) ✩ - lastoursvillense Pellegr. (BJBB 43: 275) - letestui Pellegr. (BJBB 43: 292) ✩ - longistylum Mast. (Fl.Gabon: 63) - molle Stapf (FWTA 1: 196) = H. africanum - neurophyllum Hoyle (FWTA 1: 195) = H. stipulaceum - patoklaense Aubrév. & Pellegr. (Fl.Gabon: 70) = H. lastoursvillense ✩ - smythei Hutch. & Dalziel (FWTA 1: 195) - sp.A. (FWTA 1: 197) = H. longistylum - stipulaceum Welw. ex Mast. (Fl.Gabon: 66) Lindackeria dentata (Oliver) Gilg (FWTA 1: 189) = Oncoba d. Oncoba brachyanthera Oliver (Adansonia sér. 3, 19: 256) ✪ - brevipes Stapf (Adansonia sér. 3, 19: 256) - dentata Oliver (Adansonia sér. 3, 19: 257) - echinata Oliver (Adansonia sér. 3, 19: 257) - gilgiana Sprague (Adansonia sér. 3, 19: 257) - glauca (P.Beauv.) Hook.f. (Adansonia sér. 3, 19: 257) - spinosa Forssk. (FWTA 1: 188) Ophiobotrys zenkeri Gilg (FWTA 1: 189) Scottellia chevalieri Chipp (FWTA 1: 186) = S. klaineana - coriacea A.Cheval. ex Hutch. & Dalziel (FWTA 1: 187) = S. klaineana - klaineana Pierre (Fl.Gabon: 32) ✩ - leonensis Oliver (FWTA 1: 186)

Flagellariaceae Flagellaria guineensis Schum. (FWTA 3: 51)

Gelsemiaceae (see Am.J.Bot. 87: 1029) ✩ Mostuea adamii Sillans (FWTA 2: 45) - brunonis Didr. (FWTA 2: 45) - hirsuta (T.Anders. ex Benth.) Baill. (FWTA 2: 45) ✪ - hymenocardioides Hutch. & Dalziel (FWTA 2: 45)

Graminae Acroceras gabunense (Hack.) Clayton (van der Zon 1992: 243) - zizanioides (Kunth.) Dandy (van der Zon 1992: 241) Axonopus flexuosus (Peter) Hubb. (van der Zon 1992: 287) Centotheca lappacea (Linné) Desv. (van der Zon 1992: 83) Coix lacryma-jobi Linné (van der Zon 1992: 541) Commelinidium gabunense (Hack.) Stapf (FWTA 3: 436) = Acroceras g. Cyrtococcum chaetophoron (Roem. & Schult.) Dandy (van der Zon 1992: 239) Digitaria fuscescens (Presl) Henrard (van der Zon 1992: 318) ✪ Guaduella macrostachys (K.Schum.) Pilger (van der Zon 1992: 28) ✪ - oblonga Clayton (van der Zon 1992: 25) Isachne buettneri Hack. (van der Zon 1992: 353) - kiyalaensis Robyns (van der Zon 1992: 356) - mauritiana Kunth (van der Zon 1992: 354) Leptaspis cochleata Thwaites (FWTA 3: 362) = L. zeylanica - zeylanica Nees (van der Zon 1992: 41) Megastachya mucronata (Poir.) P.Beauv. (van der Zon 1992: 84) Olyra latifolia Linné (van der Zon 1992: 38) Oplismenus burmannii (Retz.) P.Beauv. (van der Zon 1992: 192) - hirtellus (Linné) P.Beauv. (van der Zon 1992: 193) Panicum brevifolium Linné (van der Zon 1992: 207) - comorense Mez (van der Zon 1992: 228) - dinklagei Mez (FWTA 3: 431) - hochstetteri Steud. (van der Zon 1992: 221) - sadinii (Vanderyst) Renvoize (van der Zon 1992: 205) Paspalum conjugatum Bergius (van der Zon 1992: 279) - polystachyum R.Br. (FWTA 3: 446) = P. scrobiculatum - scrobiculatum Linné (van der Zon 1992: 284) Pseudechinolaena polystachya (Kunth.) Stapf (van der Zon 1992: 191) ✪ Puelia olyriformis (Franch.) Clayton (van der Zon 1992: 31) Setaria barbata (Lam.) Kunth (van der Zon 1992: 290) - chevalieri Stapf (FWTA 3: 424) = S. megaphylla - gracilipes Hubb. (van der Zon 1992: 423) - longiseta P.Beauv. (van der Zon 1992: 289) - megaphylla (Steud.) Durand & Schinz (van der Zon 1992: 293) Streptogyna crinita P.Beauv. (van der Zon 1992: 42)

Guttiferae Allanblackia floribunda Oliver (FWTA 1: 291) = A. parviflora - parviflora A.Cheval. (BJBB 39: 351) Garcinia afzelii Engl. (FWTA 1: 295) ✪ - elliotii Engl. (FWTA 1: 294) - epunctata Stapf (BJBB 39: 364) - gnetoides Hutch. & Dalziel (FWTA 1: 294) ✩ - granulata Hutch. & Dalziel (FWTA 1: 294) - kola Heckel (FWTA 1: 294) - livingstonei T.Anders. (BJBB 39: 358) ✩ - ovalifolia Oliver (FWTA 1: 295) - polyantha Oliver (FWTA 1: 294) = G. smeathmannii - smeathmannii (Planch. & Triana) Oliver (Hall & Swaine 1981: 201) Harungana madagascariensis Lam. ex Poir. (BJBB 36: 453) Mammea africana Sabine (FWTA 1: 293) Pentadesma butyracea Sabine (FWTA 1: 291) ✩ Psorospermum alternifolium Hook.f. (FWTA 1: 289) ✩ - glaberrimum Hochr. (FWTA 1: 289) ✩ - lanatum Hochr. (FWTA 1: 289) - senegalense Spach (FWTA 1: 289) - tenuifolium Hook.f. (BJBB 36: 450) Symphonia globulifera Linné f. (FWTA 1: 293) Vismia guineensis (Linné) Choisy (BJBB 39: 346)

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Hernandiaceae

Leguminosae-Caes.

✩ Illigera pentaphylla Welw. (FWTA 1: 59) ✪ - vespertilio (Benth.) Baker f. (FWTA 1: 59)

Afzelia africana Sm. (FWTA 1: 459) - bella Harms var. gracilior Keay (FWTA 1: 461) - bracteata T.Vogel ex Benth. (FWTA 1: 461) = A. parviflora ✪ - parviflora (Vahl) Hepper (Kew Bull. 26: 565) Anthonotha crassifolia (Baill.) J.Léonard (FWTA 1: 473) ✩ - elongata (Hutch.) J.Léonard (FWTA 1: 473) ✩ - ernae (Dinklage) J. Léonard (FWTA 1: 473) ✪ - explicans (Baill.) J.Léonard (FWTA 1: 473) - fragrans (Baker f.) Excell & Hillcoat (FWTA 1: 473) - macrophylla P.Beauv. (FWTA 1: 473) ✩ - sassandraensis Aubrév. & Pellegr. (Boissiera 57: 287) ✪ - vignei (Hoyle) J.Léonard (FWTA 1: 473) ✪ Aphanocalyx microphyllus (Harms) Wieringa ssp. compactus (Hutch. ex Lane-poole) Wieringa (Wieringa 1999: 161) ✪ - pteridophyllus (Harms) Wieringa (Wieringa 1999: 148) Berlinia confusa Hoyle (FWTA 1: 470) - grandiflora (Vahl) Hutch. & Dalziel (FWTA 1: 470) ✪ - occidentalis Keay (FWTA 1: 470) ✪ - tomentella Keay (FWTA 1: 470) ✪ Brachystegia leonensis Burtt Davy & Hutch. (FWTA 1: 479) ✪ Bussea occidentalis Hutch. (FWTA 1: 483) ✩ Caesalpinia benthamiana (Baill.) Herend. & Zarucchi (Ann.MBG 77: 854) - bonduc (Linné) Roxb. (FWTA 1: 481) Cassia alata (Linné) Roxb. (FWTA 1: 452) = Senna a. ✩ - aubrevillei Pellegr. (FWTA 1: 452) ✩ - fikifiki Aubrév. & Pellegr. (Boissiera 57: 289) - sieberiana DC. (FWTA 1: 452) - tuhovalyana Aké Assi (Bull.IFAN 44: 67) = Senna t. Chidlowia sanguinea Hoyle (FWTA 1: 456) ✪ Copaifera salikounda Heckel (FWTA 1: 457) Crudia gabonensis Harms (FWTA 1: 467) ✩ - klainei Pierre ex De Wild. (FWTA 1: 467) - senegalensis Benth. (FWTA 1: 467) ✪ Cryptosepalum minutifolium (A.Cheval.) Hutch. & Dalziel (FWTA 1: 480) ✩ - tetraphyllum (Hook.f.) Benth. (FWTA 1: 480) ✪ Cynometra ananta Hutch. & Dalziel (FWTA 1: 458) ✪ - leonensis Hutch. & Dalziel (FWTA 1: 458) - megalophylla Harms (FWTA 1: 458) - vogelii Hook.f. (FWTA 1: 458) Daniellia ogea (Harms) Holland (FWTA 1: 463) ✩ - thurifera Bennett (FWTA 1: 463) Detarium senegalense J.Gmelin (FWTA 1: 457) ✪ Dialium aubrevillei Pellegr. (FWTA 1: 449) - dinklagei Harms (FWTA 1: 449) - guineense Willd. (FWTA 1: 449) ✩ - pobeguinii Pellegr. (FWTA 1: 449) ✩ Didelotia afzelii Taubert (Blumea 12: 218) - brevipaniculata J.Léonard (Blumea 12: 220) ✪ - engleri Dinklage & Harms (Blumea 12: 225) ✪ - idae Oldeman, de Wit & J.Léonard (Blumea 12: 227) - sp.nr. unifoliolata (FWTA 1: 481) = D. idae Distemonanthus benthamianus Baill. (FWTA 1: 449) Duparquetia orchidacea Baill. (FWTA 1: 448) Erythrophleum guineense G.Don (FWTA 1: 484) = E. suaveolens ✩ - ivorense A.Cheval. (FWTA 1: 484) - suaveolens (Guillaumet & Perr.) Brenan (Taxon 9: 194) ✩ Gilbertiodendron aylmeri (Hutch. & Dalziel) J.Léonard (FWTA 1: 477) ✪ - bilineatum (Hutch. & Dalziel) J.Léonard (FWTA 1: 477) ✩ - ivorense (A.Cheval.) J.Léonard (FWTA 1: 477) ✪ - limba (Scott-Elliot) J.Léonard (FWTA 1: 477) ✪ - obliquum (Stapf) J.Léonard (FWTA 1: 477) ✩ - preussii (Harms) J.Léonard (FWTA 1: 477) ✪ - robynsianum Aubrév. & Pellegr. (Boissiera 57: 294) ✩ - splendidum (Hutch. & Diels) J.Léonard (FWTA 1: 475) - taiense Aubrév. (Aubrév. 1959) = G. preussii Gilletiodendron kisantuense (Vermoesen ex De Wild.) J.Leon (FWTA 1: 455) Griffonia simplicifolia (Vahl ex DC.) Baill. (FWTA 1: 445) Guibourtia copallifera Bennett (FWTA 1: 465) ✪ - dinklagei (Harms) J.Léonard (FWTA 1: 465) - ehie (A.Cheval.) J.Léonard (FWTA 1: 465) ✩ - leonensis J.Léonard (FWTA 1: 466) Hymenostegia afzelii (Oliver) Harms (FWTA 1: 464) ✩ - aubrevillei Pellegr. (FWTA 1: 464) ✪ - gracilipes Hutch. & Dalziel (FWTA 1: 464)

Hoplestigmataceae ✩ Hoplestigma klaineanum Pierre (FWTA 2: 16)

Huaceae ✩ Afrostyrax lepidophyllus Mildbr. (FWTA 2: 34)

Humiriaceae Sacoglottis gabonensis (Baill.) Urb. (FWTA 1: 354)

Icacinaceae ✪ Alsodeiopsis chippii Hutch. (FWTA 1: 637) - staudtii Engl. (FWTA 1: 638) ✩ - villosa Keay (FWTA 1: 637) ✩ Chlamydocarya macrocarpa A.Cheval. ex Hutch. & Dalziel (FWTA 1: 642) - thomsoniana Baill. (FWTA 1: 643) Desmostachys vogelii (Miers) Stapf (FWTA 1: 639) Icacina mannii Oliver (FWTA 1: 641) Iodes africana Welw. ex Oliv. (de Koning 2865 & 6302 (WAG)) ✩ - liberica Stapf (FWTA 1: 643) Leptaulus daphnoides Benth. (FWTA 1: 637) Neostachyanthus occidentalis Keay & Miège (FWTA 1: 643) = Stachyanthus o. ✪ Polycephalium capitatum (Baill.) Keay (FWTA 1: 642) Pyrenacantha acuminata Engl. (FWTA 1: 642) - cordicula Villiers (Fl.Cam.: 86) ✪ - glabrescens (Engl.) Engl. (Fl.Cam.: 72) ✩ - klaineana Pierre ex Exell & Mendonça (FWTA 1: 641) - mangenotiana Miège (FWTA 1: 641) = P. glabrescens - sp.A. (FWTA 1: 642) = P. acuminata - vogeliana Baill. (FWTA 1: 642) Rhaphiostylis beninensis (Hook.f.) Planch. ex Benth. (FWTA 1: 638) ✩ - cordifolia Hutch. & Dalziel (FWTA 1: 639) - ferruginea Engl. (FWTA 1: 639) - preussii Engl. (FWTA 1: 639) Stachyanthus occidentalis (Keay & Miège) Boutique (BJBB 39: 431)

Irvingiaceae Irvingia gabonensis (Aubry-Lecomte) Baill. (FWTA 1: 693) - robur Mildbr. (BJBB 65: 182) = Irvingia gabonensis? - wombolu Vermoesen (BJBB 65: 191) = Irvingia gabonensis? Klainedoxa gabonensis Pierre ex Engl. var. oblongifolia Engl. (BJBB 65: 152) ✩ - trillesii Pierre ex Tiegh. (BJBB 65: 161) = Klainedoxa gabonensis (part?)

✩

Labiatae Achyrospermum dasytrichum Perkins (FWTA 2: 468) - oblongifolium Baker (FWTA 2: 469) Hoslundia opposita Vahl (FWTA 2: 456) Leucas deflexa Hook.f. (FWTA 2: 470) Orthosiphon suffrutescens (Thonn.) J.K.Morton (FWTA 2: 454) Solenostemon mannii (Hook.f.) Baker (FWTA 2: 464) - monostachyus (P.Beauv.) Briq. (FWTA 2: 464) ✩ - repens (Gürke) J.K.Morton (FWTA 2: 463)

Lauraceae ✪ ✪ ✩

Beilschmiedia bitehi Aubrév. (Aubrév. 1959) - caudata (Stapf) A.Cheval. (FWTA 1: 57) - chevalieri Robyns & Wilczek (FWTA 1: 57) - djalonensis A.Cheval. (FWTA 1: 57) - mannii (Meisn.) Benth. & Hook.f. (FWTA 1: 57)

Lecythidaceae Combretodendron africanum (Welw. ex Benth. & Hook.f.) Ex (FWTA 1: 242) = Petersianthus macrocarpus - macrocarpum (P.Beauv.) Keay (FWTA 1: 761) = Petersianthus macrocarpus ✪ Napoleonaea heudelotii A.Juss. (BJBB 41: 371) - leonensis Hutch. & Dalziel (FWTA 1: 244) = N. vogelii - vogelii Hook. & Planch. (FWTA 1: 244) Petersianthus macrocarpus (P.Beauv.) Liben (BJBB 38: 207)

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Isomacrolobium vignei Hoyle (Hawthorne 1996) = Anthonotha v. Kaoue stapfiana (A.Cheval.) Pellegr. (FWTA 1: 456) = Stachyothyrsus s. ✪ Loesenera kalantha Harms (FWTA 1: 461) Mezoneuron benthamianum Baill. (FWTA 1: 481) = Caesalpinia benthamiana ✩ Mildbraediodendron excelsum Harms (FWTA 1: 448) Monopetalanthus compactus Hutch. & Dalziel (FWTA 1: 478) = Aphanocalyx microphyllus ssp. c. - pteridophyllus Harms (FWTA 1: 478) = Aphanocalyx p. - sp.A. (FWTA 1: 478) = Didelotia brevipaniculata ✩ Paramacrolobium coeruleum (Taub.) J.Léonard (FWTA 1: 474) ✩ Pellegriniodendron diphyllum (Harms) J.Léonard (FWTA 1: 474) ✩ Plagiosiphon emarginatus (Hutch. & Dalziel) J.Léonard (FWTA 1: 464) ✩ Polystemonanthus dinklagei Harms (FWTA 1: 468) Senna alata (Linné) Roxb. (Lock 1989: 36) ✪ - tuhovalyana (Aké Assi) Lock (Kew Bull. 43: 341) ✩ Stachyothyrsus stapfiana (A.Cheval.) J.Léonard & Voorh. (BJBB 34: 422) Stemonocoleus micranthus Harms (FWTA 1: 466) ✪ Talbotiella gentii Hutch. & Greenway (FWTA 1: 467) ✩ Tessmannia baikiaeoides Hutch. & Dalziel (FWTA 1: 456) ✪ Tetraberlinia tubmaniana J.Léonard (Voorhoeve 1965: 212) Toubaouate brevipaniculata (J.Léonard) Aubrév. & Pellegr. (Aubrév. 1959) = Didelotia b. Triplisomeris explicans (Baill.) Aubrév. & Pellegr. (Aubrév. 1959) = Anthonotha e.

Leguminosae-Mim. Acacia ataxacantha DC. (FWTA 1: 499) - kamerunensis Gand. (Fl.Cam.: 123) - pennata (Linné) Willd. (FWTA 1: 500) = A. kamerunensis & A. pentagona - pentagona (Schum. & Thonn.) Hook.f. (Fl.Cam.: 128) Adenopodia scelerata (A.Cheval.) Brenan (Kew Bull. 41: 76) Albizia adianthifolia (Schum.) W.F.Wight (FWTA 1: 502) - coriaria Oliver (FWTA 1: 502) - ferruginea (Guillaumet & Perr.) Benth. (FWTA 1: 502) - glaberrima (Schum. & Thonn.) Benth. (FWTA 1: 502) - zygia (DC.) J.F.Macbr. (FWTA 1: 502) Aubrevillea kerstingii (Harms) Pellegr. (FWTA 1: 492) ✩ - platycarpa Pellegr. (FWTA 1: 492) ✪ Calpocalyx aubrevillei Pellegr. (FWTA 1: 488) ✪ - brevibracteatus Harms (FWTA 1: 488) Cathormion altissimum (Hook.f.) Hutch. & Dandy (FWTA 1: 504) ✪ - rhombifolium (Benth.) Keay (FWTA 1: 504) Cylicodiscus gabunensis Harms (FWTA 1: 489) Entada gigas (Linné) Fawc. & Rendle (FWTA 1: 490) - mannii (Oliver) Tisser (FWTA 1: 490) - pursaetha DC. (FWTA 1: 490) = E. rheedei - rheedei Sprengel (Lock 1989: 92) - scelerata A.Cheval. (FWTA 1: 490) = Adenopodia s. ✪ Newtonia aubrevillei (Pellegr.) Keay (FWTA 1: 489) ✪ - duparquetiana (Baill.) Keay (FWTA 1: 489) ✩ - elliotii (Harms) Keay (FWTA 1: 489) Parkia bicolor A.Cheval. (Bot.J.Linn.Soc. 87: 148) - filicoidea Oliver (Bot.J.Linn.Soc. 87: 153) Pentaclethra macrophylla Benth. (FWTA 1: 487) Piptadeniastrum africanum (Hook.f.) Brenan (FWTA 1: 489) ✩ Pseudoprosopis bampsiana Lisowski (BJBB 52: 383) ✩ - sericeus (Hutch. & Dalziel) Brenan (BJBB 53: 428) ✩ Samanea dinklagei (Harms) Keay (FWTA 1: 504) ✩ Tetrapleura chevalieri (Harms) Baker f. (FWTA 1: 494) - tetraptera (Schum. & Thonn.) Taub. (FWTA 1: 493) ✪ Xylia evansii Hutch. (FWTA 1: 495)

Leguminosae-Pap. Abrus canescens Baker (FWTA 1: 575) - fruticulosus Wall. ex W. & A. (Blumea 10: 612) - precatorius Linné (FWTA 1: 574) - pulchellus Thwaites (FWTA 1: 574) = A. fruticulosus Afrormosia elata Harms (FWTA 1: 510) = Pericopsis e. ✩ Aganope gabonica (Baill.) Polhill (Kew Bull. 25: 269) ✩ - leucobotrya (Dunn) Polhill (Kew Bull. 25: 269) Airyantha schweinfurthii (Taub.) Brummitt (Kew Bull. 22: 381) Amphimas pterocarpoides Harms (FWTA 1: 448) Andira inermis (Wright) DC (FWTA 1: 59) Angylocalyx oligophyllus (Baker) Baker f. (Adansonia sér. 2, 8: 328) Baphia bancoensis Aubrév. (FWTA 1: 513) = B. pubescens - capparidifolia Baker var. polygalacea Brummitt (Kew Bull. 35: 323)

✩

- heudelotiana Baill. (Kew Bull. 40: 320) - nitida Lodd. (Kew Bull. 40: 352) - polygalacea (Hook.f.) Baker (FWTA 1: 512) = B. capparidifolia var. polygalacea - pubescens Hook.f. (Kew Bull. 40: 362) ✪ - spathacea Hook.f. ssp. spathacea (Kew Bull. 40: 349) Baphiastrum confusum (Hutch. & Dalziel) Pellegr. (FWTA 1: 513) = Airyantha schweinfurthii Bobgunnia fistuloides (Harms) Kirkbr. & Wiersema (Brittonia 49: 3) Bowringia discolor J.B.Hall (Kew Bull. 29: 497) = Leucomphalos d. Centrosema plumieri (Pers.) Benth. (FWTA 1: 560) ✩ Craibia atlantica Dunn (FWTA 1: 527) ✩ Dalbergia adamii J.Berhaut (Fl.Senegal ed. 2: 427) - afzeliana G.Don (FWTA 1: 516) ✩ - albiflora A.Cheval. ex Hutch. & Dalziel ssp. albiflora (FWTA 1: 516) - bignonae Berhaut (Fl.Senegal ed. 2: 427) ✩ - crispa Hepper (FWTA 1: 515) - ecastaphyllum (Linné) Taub. (FWTA 1: 515) ✩ - hepperi Jongkind spec.nov. (to be published) - heudelotii Stapf (FWTA 1: 515) - hostilis Benth. (FWTA 1: 516) - noldeae Harms (Fl.Guinea-Bis.: 24) ✩ - oblongifolia G.Don (FWTA 1: 516) - rufa G.Don (FWTA 1: 516) ✩ - rugosa Hepper (FWTA 1: 515) - saxatilis Hook.f. (FWTA 1: 516) ✩ - setifera Hutch. & Dalziel (FWTA 1: 515) - sp.nr. pachycarpa (FWTA 1: 516) Dalbergiella welwitschii (Baker) Baker f. (FWTA 1: 517) Desmodium adscendens (Sw.) DC. (FWTA 1: 585) - gangeticum (Linné) DC. var. gangeticum (FWTA 1: 584) - repandum (Vahl) DC. (FWTA 1: 584) - salicifolium (Poir.) DC. (FWTA 1: 584) - velutinum (Willd.) DC. (FWTA 1: 584) Dioclea hexandra (Ralph) Mabberley (Taxon 29: 605) - reflexa Hook.f. (FWTA 1: 574) = D. hexandra Dolichos falcatus Klein ex Willd. (FWTA 1: 571) = D. trilobus ssp. occidentalis ✩ - nimbaensis Schnell (FWTA 1: 571) ✩ - tonkouiensis Porteres (FWTA 1: 571) - trilobus Linné ssp. occidentalis Verdc. (Kew Bull. 24: 423) Drepanocarpus lunatus (Linné f.) G.F.W.Mey. (FWTA 1: 519) = Machaerium lunatum ✩ Erythrina addisoniae Hutch. & Dalziel (FWTA 1: 563) - mildbraedii Harms (FWTA 1: 563) - vogelii Hook.f. (FWTA 1: 563) ✩ Haplormosia monophylla (Harms) Harms (FWTA 1: 511) Indigofera macrophylla Schum. & Thonn. (FWTA 1: 541) Leptoderris brachyptera (Benth.) Dunn. (FWTA 1: 521) ✩ - cyclocarpa Dunn (FWTA 1: 521) - fasciculata (Benth.) Dunn. (FWTA 1: 521) - glabrata (Welw.) Dunn (J.K. Morton SL 1117 & SL 3071 (K, WAG)) - micrantha Dunn (FWTA 1: 521) ✪ - miegei Aké Assi & Mangenot (Boissiera 24: 313) ✩ - sassandrensis Jongkind (Syst. Geogr. Pl. 73 : 96) ✩ - trifoliolata Hepper (FWTA 1: 521) ✩ Leucomphalos discolor (J.B.Hall) Breteler (WAUP 94-4: 25) ✪ - libericus Breteler (WAUP 94-4: 27) Lonchocarpus cyanescens (Schum. & Thonn.) Benth. (FWTA 1: 523) = Philenoptera c. - griffonianus (Baill.) Dunn (FWTA 1: 523) = Millettia griffoniana - sericeus (Poir.) Kunth (FWTA 1: 522) - sp.A. (FWTA 1: 523) = Leptoderris fasciculata Machaerium lunatum (L.f.) Ducke (Acta Bot.N. 20: 655) Millettia barteri (Benth.) Dunn. (FWTA 1: 526) - chrysophylla Dunn (FWTA 1: 526) ✩ - dinklagei Harms (FWTA 1: 525) - griffoniana Baill. (Kew Bull. 25: 260) - irvinei Hutch. & Dalziel (FWTA 1: 526) = Millettia warneckei (syn. nov.) ✩ - lane-poolei Dunn (FWTA 1: 526) ✪ - leonensis Hepper (FWTA 1: 526) ✪ - liberica Jongkind (Blumea 48: 165) ✩ - lucens (Scott-Elliot) Dunn (FWTA 1: 525) ✪ - pallens Stapf (FWTA 1: 527) ✩ - rhodantha Baill. (FWTA 1: 526) - sanagana Harms (FWTA 1: 526) ✩ - takou Lorougnon (Boissiera 57: 328) - thonningii (Schum. & Thonn.) Baker (FWTA 1: 527) ✩ - warneckei Harms var. warneckei (FWTA 1: 526)

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✩ ✩

- warneckei Harms var. porphyrocalyx (Dunn) Hepper (FWTA 1: 526) - zechiana Harms (FWTA 1: 526) Mucuna flagellipes Hook.f. (FWTA 1: 561) - poggei Taubert var. occidentalis Hepper (FWTA 1: 561) - pruriens (Linné) DC. var. pruriens (FWTA 1: 561) - sloanei Fawc. & Rendle (FWTA 1: 561) Ormocarpum megalophyllum Harms (FWTA 1: 577) - sennoides (Willd.) DC (FWTA 1: 576) - verrucosum P.Beauv. (FWTA 1: 577) Ostryocarpus major Stapf (FWTA 1: 519) = Aganope gabonica?? - riparius Hook.f. (FWTA 1: 519) Ostryoderris leucobotrya Dunn (FWTA 1: 522) = Aganope l. Pericopsis elata (Harms) Van Meeuwen (BJBB 32: 213) Philenoptera cyanescens (Schum. & Thonn.) Roberty (Kew Bull. 55: 85) Physostigma venenosum Balf. (FWTA 1: 564) ✪ Platysepalum hirsutum (Dunn) Hepper (FWTA 1: 524) Psophocarpus palustris Desv. (FWTA 1: 572) Pterocarpus mildbraedii Harms (FWTA 1: 517) - santalinoides DC. (FWTA 1: 517) ✩ Rhynchosia brunnea Baker f. (FWTA 1: 554) - congensis Baker (FWTA 1: 554) - densiflora (Roth.) DC. (FWTA 1: 555) - pycnostachya (DC.) Meikle (FWTA 1: 554) - violacea (Hiern) K.Schum. (FWTA 1: 555) = R. viscosa - viscosa (Roth.) DC. (Lock 1989: 437) Swartzia fistuloides Harms (FWTA 1: 446) = Bobgunnia f.

Liliaceae Asparagus flagellaris (Kunth.) Baker (FWTA 3: 93) - racemosus Willd. (FWTA 3: 93) - warneckei (Engl.) Hutch. (FWTA 3: 93) Chlorophytum alismifolium Baker (FWTA 3: 101) - inornatum Ker-Gawl. (FWTA 3: 100) - laxum R.Br. (FWTA 3: 100) - macrophyllum (A.Rich.) Aschers. (FWTA 3: 99) - nzii A.Cheval. ex Hepper (FWTA 3: 99) - orchidastrum Lindl. (FWTA 3: 100) - sparsiflorum Baker (FWTA 3: 100) ✩ - togoense Engl. (FWTA 3: 99) Gloriosa simplex Linné (FWTA 3: 106) = G. superba - superba Linné (Kew Bull. 25: 243) Smilax anceps Willd. (FTEA Smil.: 2) - kraussiana Meisn. (FWTA 3: 112) = S. anceps

Linaceae Hugonia afzelii R.Br. ex Planch. (Adansonia sér. 2, 11: 100) - foliosa Oliver (FWTA 1: 359) = H. afzelii - planchonii Hook.f. (Adansonia sér. 2, 11: 101) - platysepala Welw. ex Oliver (Adansonia sér. 2, 11: 99) ✪ - rufipilis A.Cheval. ex Hutch. & Dalziel (Adansonia sér. 2, 11: 100) Ochthocosmus africanus Hook.f. (FWTA 1: 355) = Phyllocosmus a. - chippii Sprague & Hutch. ex Hutch. & Dalziel (FWTA 1: 355) = O. sessiliflorus - sessiliflorus (Oliver) Baill. (Fl.Cam.: 58) = Phyllocosmus s. Phyllocosmus africanus (Hook.f.) Klotzsch (Kew Bull. 19: 517) - sessiliflorus Oliver (Kew Bull. 19: 517)

Loganiaceae (see Am.J.Bot. 87: 1029) Strychnos aculeata Soler. (Med.LUW 69-1: 49) - afzelii Gilg (Med.LUW 69-1: 53) ✩ - asterantha Leeuwenb. (Med.LUW 69-1: 62) - barteri Soler. (Med.LUW 69-1: 65) ✩ - campicola Gilg ex Leeuwenb. (Med.LUW 69-1: 74) - camptoneura Gilg & Busse (Med.LUW 69-1: 75) ✩ - congolana Gilg (Med.LUW 69-1: 92) ✩ - cuminodora Leeuwenb. (Med.LUW 69-1: 96) - densiflora Baill. (Med.LUW 69-1: 104) ✪ - dinklagei Gilg (Med.LUW 69-1: 108) - floribunda Gilg (Med.LUW 69-1: 120) - icaja Baill. (Med.LUW 69-1: 133) - johnsonii Hutch. & M.B.Moss (Med.LUW 69-1: 147) ✩ - longicaudata Gilg (Med.LUW 69-1: 153) - malacoclados C.H.Wright (Med.LUW 69-1: 171) ✪ - melastomatoides Gilg (Med.LUW 69-1: 179) ✪ - millepunctata Leeuwenb. (Med.LUW 69-1: 186)

462

- ngouniensis Pellegr. (FWTA 2: 43) = S. soubrensis - nigritana Baker (Med.LUW 69-1: 204) ✪ - odorata A.Cheval. (Med.LUW 69-1: 207) - phaeotricha Gilg (Med.LUW 69-1: 215) ✪ - soubrensis Hutch. & Dalziel (Med.LUW 69-1: 236) - splendens Gilg (Med.LUW 69-1: 248) - usambarensis Gilg (Med.LUW 69-1: 267) Usteria guineensis Willd. (Acta.Bot.N. 12: 115)

Loranthaceae Agelanthus brunneus (Engl.) Balle (Polhill & Wiens, 1998: 156) ✪ Englerina gabonensis (Engl.) Balle (Polhill & Wiens, 1998: 121) ✪ - parviflora (Tiegh.) Balle (Polhill & Wiens, 1998: 124) ✪ Globimetula assiana (Balle) Wiens & Polhill (Polhill & Wiens, 1998: 218) ✩ - braunii (Engl.) Danser (Polhill & Wiens, 1998: 211) ✪ - cupulata (DC.) Danser (FWTA 1: 660) Phragmanthera capitata (Sprengel) Balle (Polhill & Wiens, 1998: 252) - incana (Schum.) Balle (FWTA 1: 664) = P. capitata ✩ - leonensis (Sprague) Balle (Polhill & Wiens, 1998: 267) - nigritana (Hook.f. ex Benth.) Balle (Revision Polhill & Wiens, 1998: 268) ✩ - rufescens (DC.) Balle (Polhill & Wiens, 1998: 257) ✪ - vignei Balle (Revision Polhill & Wiens, 1998: 268) Tapinanthus bangwensis (Engl. & K.Krause) Danser (Polhill & Wiens, 1998: 200) ✪ - belvisii (DC.) Danser (Polhill & Wiens, 1998: 204) ✩ - buntingii (Sprague) Danser (Polhill & Wiens, 1998: 187) - farmari (Sprague) Danser (Polhill & Wiens, 1998: 205) ✪ - praetexta Polhill & Wiens (Polhill & Wiens, 1998: 205) - truncatus (Engl.) Danser (FWTA 1: 663) = T. belvisii

Malpighiaceae ✪ Acridocarpus alternifolius (Schum. & Thonn.) Nied. (FWTA 1: 352) ✩ - chevalieri Sprague (FWTA 1: 352) ✩ - longifolius (G.Don) Hook.f. (FWTA 1: 352) = A. longifolius (pro parte) ✩ - longifolius (pro parte) (to be published) - macrocalyx Engl. (FWTA 1: 352) ✪ - plagiopterus Guillaumet & Perr. (FWTA 1: 352) - smeathmanii (DC.) Guillaumet & Perr. (FWTA 1: 352) Flabellaria paniculata Cav. (FWTA 1: 353) Heteropteris leona (Cav.) Exell (FWTA 1: 353) Stigmaphyllon ovatum (Cav.) Nied. (FWTA 1: 353) ✩ Triaspis odorata (Willd.) A.Juss. (FWTA 1: 354) - stipulata Oliver (FWTA 1: 354)

Malvaceae ✩ Hibiscus comoensis A.Cheval. ex Hutch. & Dalziel (FWTA 1: 346) - owariensis P.Beauv. (FWTA 1: 347) - rostellatus Guillaumet & Perr. (FWTA 1: 346) - surattensis Linné (FWTA 1: 346) - tiliaceus Linné (FWTA 1: 345) ✪ - whytei Stapf (FWTA 1: 347) Thespesia populnea (Linné) Soland. ex Corr. (FWTA 1: 342)

Marantaceae ✩ Ataenidia conferta (Benth.) Milne-Redh. (FWTA 3: 89) ✩ Halopegia azurea (K.Schum.) K.Schum. (FWTA 3: 85) ✩ Hypselodelphys poggeana (K.Schum.) Milne-Redh. (FWTA 3: 88) - scandens Louis & Mullend. (FWTA 3: 89) ✩ - triangulare Jongkind spec.nov. (to be published) ✩ - velutina sp.nov. (to be published) - violacea (Ridl.) Milne-Redh. (FWTA 3: 88) Marantochloa congensis (K.Schum.) Léonard & Mullend (FWTA 3: 81) ✪ - cuspidata (Rose.) Milne-Redh. (FWTA 3: 81) - filipes (Benth.) Hutch. (FWTA 3: 81) - leucantha (K.Schum.) Milne-Redh. (FWTA 3: 81) ✩ - mannii (Benth.) Milne-Redh. (FWTA 3: 81) - purpurea (Ridl.) Milne-Redh. (FWTA 3: 81) ✩ - ramosissima (Benth.) Hutch. (FWTA 3: 81) ✩ Megaphrynium distans Hepper (FWTA 3: 89) - macrostachyum (Benth.) Milne-Redh. (FWTA 3: 89) Sarcophrynium brachystachyum (Benth.) K.Schum. (FWTA 3: 89) - prionogonium (K.Schum.) K.Schum. var. prionogonium (FWTA 3: 88) ✩ - prionogonium (K.Schum.) K.Schum. var. ivorens Schnell (FWTA 3: 88) Thalia geniculata Linné (Nord.J.Bot. 1: 48) - welwitschii Ridl. (FWTA 3: 85) = T. geniculata

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Thaumatococcus daniellii (Benn.) Benth. (FWTA 3: 81) Trachyphrynium braunianum (K.Schum.) Baker (FWTA 3: 89)

Medusandraceae ✩ Soyauxia floribunda Hutch. (FWTA 1: 653) ✪ - grandifolia Gilg & Stapf (FWTA 1: 653) ✩ - velutina Hutch. & Dalziel (FWTA 1: 653)

Melastomataceae ✩ Amphiblemma cymosum Naud. (Adansonia sér. 2, 13: 444) Bourdaria felicis A.Cheval. (FWTA 1: 251) = Cincinnobotrys f. ✩ Calvoa hirsuta Hook.f. (Bull.MNHN Paris sér. 4, 3: 132) ✪ - monticola A.Cheval. ex Hutch. & Dalziel (Bull.MNHN Paris sér. 4, 3: 142) ✩ - trochainii Jacq.-Félix (Bull.MNHN Paris sér. 4, 3: 129) ✩ Cincinnobotrys felicis (A.Cheval.) Jacq.-Félix (Adansonia sér. 2, 16: 377) Dicellandra barteri Hook.f. var. barteri (Adansonia sér. 2, 14: 86) Dichaetanthera africana (Hook.f.) Jacq.Félix (Fl.Cam.: 55) - echinulata (Hook.f.) Jacq.-Félix (FWTA 1: 761) Dinophora spenneroides Benth. (Fl.Cam.: 116) Dissotis antennina (Sm.) Triana (FWTA 1: 257) = Heterotis a. - cornifolia (Benth.) Hook.f. (FWTA 1: 259) = Melastomastrum c. - entii J.B.Hall (Kew Bull. 24: 346) = Heterotis e. ✩ - leonensis Hutch. & Dalziel (FWTA 1: 258) - multiflora (Sm.) Triana (Fl.Cam.: 20) - paucistellata Stapf (FWTA 1: 259) = Melastomastrum afzelii var. p. - sylvestris Jacq.-Félix (FWTA 1: 256) = Heterotis s. Guyonia ciliata Hook.f. (Fl.Cam.: 8) ✩ Heterotis antennina (Sm.) Benth. (Adansonia sér. 2, 20: 418) ✩ - entii (J.B.Hall) Jacq.-Félix (Adansonia sér. 2, 20: 419) ✩ - sylvestris (Jacq.-Félix) Jacq.-Félix (Bull.MNHN Paris sér. 3, 16: 272) Lijndenia barteri (Hook.f.) Bremer (Fl.Cam.: 174) Medinilla entii Hossain (BJBB 40: 6) = M. mannii - mannii Hook.f. (Fl.Cam.: 112) Melastomastrum afzelii (Hook.f.) A. & R.Fern. (Bull.MNHN Paris sér. 3, 17: 61) - cornifolium (Benth.) Jacq.-Félix (Bull.MNHN Paris sér. 3, 17: 66) Memecylon afzelii G.Don var. amoenum Jacq.-Félix (Adansonia sér. 2, 18: 426) - afzelii G.Don var. afzelii (Fl.Cam.: 139) ✪ - aylmeri Hutch. & Dalziel (FWTA 1: 262) - barteri Hook.f. (FWTA 1: 263) = Lijndenia b. - blakeoides G.Don (FWTA 1: 263) = Spathandra blakeoides - cinnamonoides G.Don (FWTA 1: 263) = Warneckea c. - dinklagei Gilg ex Engl. (FWTA 1: 263) = Lijndenia barteri - engleranum Cogn. var. occidentale Jacq.-Félix (Adansonia sér. 2, 18: 431) - fasciculare (Planch. ex Benth.) Naud. (FWTA 1: 263) = Warneckea fascicularis - fleuryi Jacq.-Félix (FWTA 1: 263) = Spathandra blakeoides var. fleuryi - golaense Baker f. (FWTA 1: 263) = Warneckea golaensis - guineense Keay (FWTA 1: 263) = Warneckea guineensis - lateriflorum (G.Don) Bremek. (FWTA 1: 262) ✩ - liberiae Gilg ex Engl. (Adansonia sér. 2, 18: 424) - membranifolium Hook.f. (FWTA 1: 263) = Warneckea membranifolia - memecyloides (Benth.) Exell (FWTA 1: 263) = Warneckea m. ✩ - memoratum Jacq.-Félix (Adansonia sér. 2, 18: 416) - normandii Jacq.-Félix (FWTA 1: 262) - occultum Jacq.-Félix (Adansonia sér. 2, 18: 412) ✩ - polyanthemos Hook.f. (FWTA 1: 262) ✩ - ramosum Jacq.-Félix (Adansonia sér. 2, 18: 412) - sp.A. (FWTA 1: 263) = M. memoratum - viride Hutch. & Dalziel (Fl.Cam.: 138) Ochthocharis dicellandroides (Gilg) Hansen & Wickens (Kew Bull. 36: 23) Phaeoneuron dicellandroides Gilg (FWTA 1: 247) = Ochthocharis d. Preussiella chevalieri Jacq.-Félix (FWTA 1: 251) = P. kamerunensis - kamerunensis Gilg (Adansonia sér. 2, 16: 410) Sakersia africana Hook.f. (FWTA 1: 249) = Dichaetanthera a. - echinulata Hook.f. (FWTA 1: 249) = Dichaetanthera e. Spathandra barteri Hook.f. (Fl.Cam.: 158) = Lijndenia b. ✩ - blakeoides (G.Don) Jacq.-Félix var. fleuryi (Jacq.-Félix) Jacq.-Félix (Adansonia sér. 2, 18: 226) - blakeoides (G.Don) Jacq.-Félix var. blakeoides (Adansonia sér. 2, 18: 226) ✪ Tristemma akeassii Jacq.-Félix (Bull.MNHN Paris sér. 2, 28: 168) ✩ - albiflorum (G.Don) Benth. (Bull.MNHN Paris sér. 2, 28: 180) - coronatum Benth. (Bull.MNHN Paris sér. 2, 28: 195) ✩ - involucratum Benth. (Bull.MNHN Paris sér. 2, 28: 174) - mauritianum J.-F.Gmelin (Bull.MNHN Paris sér. 2, 28: 152) Warneckea cinnamomoides (G.Don) Jacq.-Félix (Fl.Cam.: 164) - fascicularis (Planch ex Benth.) Jacq.-Félix var. fascicularis (Adansonia sér. 2, 19: 266)

✩

- golaensis (Baker f.) Jacq.-Félix (Adansonia sér. 2, 18: 231) - guineensis (Keay) Jacq.-Félix (Adansonia sér. 2, 18: 235) - membranifolia (Hook.f.) Jacq.-Félix (Fl.Cam.: 172) - memecyloides (Benth.) Jacq.-Félix (Fl.Cam.: 163)

Meliaceae Azadirachta indica A.Juss. (FWTA 1: 708) Carapa procera DC. (FWTA 1: 702) Ekebergia capensis Sparrm. (FTEA: 38) - senegalensis A.Juss. (FWTA 1: 705) = E. capensis Entandrophragma angolense (Welw.) DC. (FWTA 1: 700) - candollei Harms (FWTA 1: 700) - cylindricum (Sprague) Sprague (FWTA 1: 701) - utile (Dawe & Sprague ) Sprague (FWTA 1: 700) Guarea cedrata (A.Cheval.) Pellegr. (FWTA 1: 706) ✩ - leonensis Hutch. & Dalziel (FWTA 1: 706) - thompsonii Sprague & Hutch. (FWTA 1: 707) Heckeldora latifolia Pierre (see Blumea 22: 491) = Heckeldora staudtii - mangenotiana Aké Assi & Lorougnon (Bull.Soc.Bot.Fr. 136: 167) - staudtii (Harms) Staner (FWTA 1: 707) Khaya anthotheca (Welw.) C.DC. (FWTA 1: 699) - grandifoliola C.DC. (FWTA 1: 699) - ivorensis A.Cheval. (FWTA 1: 699) Lovoa trichilioides Harms (FWTA 1: 702) ✪ Trichilia djalonis A.Cheval. (Med.LHW 68-2: 23) ✩ - dregeana Sonder (Med.LHW 68-2: 28) - heudelotii Planch. ex Oliver var. zenkeri (Harms) Aubrév. (Aubrév. 1959) = T. ornithothera - heudelotii Planch. ex Oliver (FWTA 1: 704) = T. monadelpha - lanata A.Cheval. (FWTA 1: 705) = T. tessmannii - martineaui Aubrév. & Pellegr. (Med.LHW 68-2: 96) ✪ - megalantha Harms (Med.LHW 68-2: 102) - monadelpha (Thonn.) J.J.de Wilde (Med.LHW 68-2: 108) ✪ - ornithothera J.J.de Wilde (Med.LHW 68-2: 122) - prieuriana A.Juss. (Med.LHW 68-2: 130) - splendida A.Cheval. (FWTA 1: 705) = T. dregeana ✩ - tessmannii Harms (Med.LHW 68-2: 171) ✩ Turraea adjanohounii Aké Assi (BJBB 31: 507) ✪ - ghanensis J.B.Hall (Adansonia sér. 2, 15: 505) ✪ - heterophylla Sm. (FWTA 1: 708) ✩ - leonensis Keay (FWTA 1: 708) - vogelii Hook.f. ex Benth. (FWTA 1: 708) Turraeanthus africanus (Welw. ex C.DC.) Pellegr. (FWTA 1: 707)

Melianthaceae Bersama abyssinica Fresen (FWTA 1: 726) - paullinioides (Planch.) Baker (Aubrév. 1959) = B. abyssinica

Menispermaceae ✪ Albertisia cordifolia (Mangenot & Miège) Form. (Kew Bull. 30: 83) ✪ - cuneata (Keay) Forman (Kew Bull. 30: 83) ✪ - ferruginea (Diels) Forman (Kew Bull. 30: 83) ✪ - mangenotii (Guillaumet & Debray) Forman (Kew Bull. 30: 688) ✪ - scandens (Mangenot & Miège) Forman (Kew Bull. 30: 83) Chasmanthera dependens Hochst. (Troupin 1962: 161) Cissampelos owariensis P.Beauv. ex DC. (Troupin 1962: 287) Dioscoreophyllum cumminsii (Stapf) Diels (Troupin 1962: 133) - tenerum Engl. var. tenerum (FWTA 1: 73) = D. volkensii - volkensii Engl. (Troupin 1962: 138) Epinetrum cordifolium Mangenot & Miège (Troupin 1962: 42) = Albertisia cordifolia - cuneatum Keay (Troupin 1962: 31) = Albertisia cuneata - ferrugineum (Diels) Keay (Troupin 1962: 41) = Albertisia ferruginea - mangenotii Guillaumet & Debray (Adansonia sér. 2, 4: 315) = Albertisia m. - scandens Mangenot & Miège (Troupin 1962: 40) = Albertisia s. Kolobopetalum auriculatum Engl. (Troupin 1962: 180) = K. leonense - chevalieri (Hutch. & Dalziel) Troupin (Troupin 1962: 183) = K. ovatum ✩ - leonense Hutch. & Dalziel (Troupin 1962: 182) ✩ - ovatum Stapf (Troupin 1962: 185) ✪ Penianthus patulinervis Hutch. & Dalziel (BJBB 53: 52) - zenkeri (Engl.) Diels (Troupin 1962: 121) = P. patulinervis ✩ Rhigiocarya peltata Miège (Troupin 1962: 171) - racemifera Miers (Troupin 1962: 168) Sphenocentrum jollyanum Pierre (BJBB 53: 59) Stephania dinklagei (Engl.) Diels (Troupin 1962: 258)

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Syntriandrium preussii Engl. (Geerling & Bokdam 2341(WAG)) ✩ Syrrhonema hexastamineum Keay (Troupin 1962: 108) Tiliacora ? odorata Engl. (FWTA 1: 71) = T. louisii - dielsiana Hutch. & Dalziel (Troupin 1962: 52) = T. leonensis - dinklagei Engl. (Hall & Swaine 1981: 305) = T. leonensis - funifera (Miers) Oliver (Troupin 1962: 58) ✪ - leonensis (Sc.Eliott) Diels (Troupin 1962: 54) ✩ - louisii Troupin (Troupin 1962: 73) - warneckei Engl. ex Diels (FWTA 1: 71) = T. funifera ✪ Triclisia dictyophylla Diels (Troupin 1962: 88) - gilletii (De Wild.) Staner (FWTA 1: 71) = T. dictyophylla - macrophylla Oliver (Troupin 1962: 92) = doubtful for Upper Guinea ✩ - patens Oliver (Troupin 1962: 82) - subcordata Oliver (Troupin 1962: 86)

Moraceae Antiaris africana Engl. (FWTA 1: 612) = A. toxicaria - toxicaria (Rumph. ex Pers.) Leschen. (BJBB 47: 308) - welwitschii Engl. (FWTA 1: 613) = A. toxicaria Bosqueia angolensis Ficalho (FWTA 1: 612) = Trilepisium madagascariense - phoberos Baill. (Aubrév. 1959) = Trilepisium madagascariense Cecropia peltata Linné (Bull.IFAN 42: 96) Chlorophora excelsa (Welw.) Benth. (BJBB 47: 349) = Milicia e. - regia A.Cheval. (BJBB 47: 353) = Milicia r. Craterogyne djettii (Guillaumet) N.Hallé & Aké Assi (Adansonia sér. 2, 7: 390) = Dorstenia d. - kameruniana (Engl.) Lanjouw (FWTA 1: 599) = Dorstenia k. ✪ Dorstenia astyanactis Aké Assi (Adansonia sér. 2, 7: 387) ✩ - djettii Guillaumet (Bot.Notiser 131: 58) ✪ - embergeri Mangenot (Boissiera 57: 388) - kameruniana Engl. (Bot.Notiser 131: 62) - smythei Sprague (FWTA 1: 599) = D. turbinata ✪ - turbinata Engl. (Fl.Cam.: 36) Ficus anomani Hutch. (FWTA 1: 607) = F. craterostoma - ardisioides Warb. ssp. camptoneura (Mildbr.) C.C.Berg (Berg & Wiebes 1992: 141) - artocarpoides Warb. (Berg & Wiebes 1992: 158) - asperifolia Miq. (Berg & Wiebes 1992: 68) - barteri Sprague (Berg & Wiebes 1992: 147) - bongouanouensis A.Cheval. (Aubrév. 1959) = F. variifolia - bubu Warb. (Berg & Wiebes 1992: 164) - calyptrata Thonn. ex Vahl (Berg & Wiebes 1992: 116) - camptoneura Mildbr. (Fl.Cam.: 238) = F. ardisioides ssp. camptoneura - capensis Thunb. (FWTA 1: 606) = F. sur - congensis Engl. (FWTA 1: 609) = F. trichopoda - conraui Warb. (Berg & Wiebes 1992: 139) - craterostoma Mildbr. & Burrett (Berg & Wiebes 1992: 118) - cyathistipula Warb. ssp. cyathistipula (Berg & Wiebes 1992: 143) - cyathistipuloides De Wild. (Berg & Wiebes 1992: 145) - dekdekena (Miq.) A.Rich. (FWTA 1: 610) = F. thonningii - djalonensis A.Cheval. (FWTA 1: 608) = F. calyptrata - elasticoides De Wild. (Berg & Wiebes 1992: 133) - elegans (Miq.) Miq. (FWTA 1: 611) = F. artocarpoides - eriobotryoides Kunth & Bouche (FWTA 1: 608) = F. saussureana - exasperata Vahl (Berg & Wiebes 1992: 63) - gnaphalocarpa (Miq.) A.Rich. (FWTA 1: 606) = F. sycomorus - goliath A.Cheval. (FWTA 1: 610) = F. recurvata - kamerunensis Mildbr. & Burrett (Berg & Wiebes 1992: 130) - leonensis Hutch. (Berg & Wiebes 1992: 138) - leprieuri Miq. (FWTA 1: 608) = F. natalensis ssp. leprieuri - lingua Warb. ssp. lingua (Berg & Wiebes 1992: 120) - louisii Boutique & Léonard (Berg & Wiebes 1992: 138) - lutea Vahl (Berg & Wiebes 1992: 99) - lyrata Warb. (Berg & Wiebes 1992: 149) - macrosperma Warb. ex Mildbr. & Burret (Fl.Cam.: 220) = F. sansibarica var. macrosperma - mucuso Ficalho (Berg & Wiebes 1992: 76) - natalensis Hochst. ssp. leprieurii (Miq.) C.C.Berg (Berg & Wiebes 1992: 122) ✩ - ottoniifolia (Miq.) Miq. ssp. multinervia C.C. Berg (Berg & Wiebes 1992: 154) - ottoniifolia (Miq.) Miq. ssp. ottoniifolia (Berg & Wiebes 1992: 154) - ovata Vahl (Berg & Wiebes 1992: 165) ✩ - pachyneura C.C.Berg (Berg & Wiebes 1992: 140) - polita Vahl ssp. polita (Berg & Wiebes 1992: 159) - praticola Mildbr. & Hutch. (FWTA 1: 607) = F. conraui - pringsheimiana Braub & K. Schumann (Fl.Cam.: 240) = F. cyathistipula

464

- pseudomangifera Hutch. (Berg & Wiebes 1992: 135) - recurvata De Wild. (Berg & Wiebes 1992: 103) - sagittifolia Warb. ex Mildbr. & Burret (Berg & Wiebes 1992: 149) - sansibarica Warb. var. macrosperma (Mildbr. & Burr.) C.C. Berg (Berg & Wiebes 1992: 162) - saussureana DC. (Berg & Wiebes 1992: 98) ✪ - scott-elliotii Mildbr. & Burrett (Berg & Wiebes 1992: 145) - sur Forssk. (Berg & Wiebes 1992: 77) - sycomorus Linné (Berg & Wiebes 1992: 74) - tesselata Warb. (Berg & Wiebes 1992: 140) - thonningii Bl. (Berg & Wiebes 1992: 126) - trichopoda Baker (Berg & Wiebes 1992: 112) - umbellata Vahl (Berg & Wiebes 1992: 164) - variifolia Warb. (Berg & Wiebes 1992: 85) - vogeliana (Miq.) Miq. (Berg & Wiebes 1992: 78) - vogelii (Miq.) Miq. (FWTA 1: 609) = F. lutea Milicia excelsa (Welw.) C.C.Berg (Fl.Cam.: 9) ✪ - regia (A.Cheval.) C.C.Berg (BJBB 52: 227) Morus mesozygia Stapf (BJBB 47: 337) Musanga cecropioides F.Br. (Fl.Cam.: 272) Myrianthus arboreus P.Beauv. (BJBB 46: 478) ✪ - libericus Rendle (BJBB 46: 488) - serratus (Trecul) Benth. (BJBB 46: 491) Neosloetiopsis kamerunensis Engl. (FWTA 1: 595) = Streblus usambarensis Sloetiopsis usambarensis Engl. (BJBB 47: 364) = Streblus u. ✩ Streblus usambarensis (Engl.) Berg (Boissiera 57: 394) Treculia africana Decne ssp. africana (BJBB 47: 382) Trilepisium madagascariense DC. (BJBB 47: 299)

Myristicaceae ✩ Coelocaryon oxycarpum Stapf (FWTA 1: 61) - preussii Warb. (FWTA 1: 61) = C. sphaerocarpum ✩ - sphaerocarpum Fouilloy (Adansonia sér. 2, 12: 548) Pycnanthus angolensis (Welw.) Warb. (FWTA 1: 61) ✪ - dinklagei Warb. (FWTA 1: 61) - kombo Warb. (Aubrév. 1959) = P. angolensis

Myrsinaceae ✩ Embelia djalonensis A.Cheval. ex Hutch. & Dalziel (FWTA 2: 32) - guineensis Baker (BJBB 50: 202) - rowlandii Gilg (BJBB 50: 204) ✩ - schimperi Vatke (BJBB 50: 204) - sp.A. (FWTA 2: 32) = E. schimperi Maesa lanceolata Forssk. (BJBB 50: 208) ✪ - nuda Hutch. & Dalziel (Adansonia sér. 2, 10: 377) ✩ - vestita Jacq.-Félix (Adansonia sér. 2, 10: 379)

Myrtaceae ✩ Eugenia calophylloides DC. (FWTA 1: 238) - calycina Benth. nom.ill. (FWTA 1: 237) - coronata Schum. & Thonn. (FWTA 1: 237) - dinklagei Engl. & v.Brehm. (FWTA 1: 238) ✩ - elliotii Engl. & v.Brehm. (FWTA 1: 237) - gabonensis Amshoff (Fl.Gabon: 21) - kalbreyeri Engl. & v.Brehm. (FWTA 1: 238) ✩ - liberiana Amshoff (Act.Bot.N. 7: 57) ✩ - memecyloides Benth. (FWTA 1: 238) - miegeana Aké Assi (BJBB 30: 15) = E. gabonensis - nigerina A.Cheval. ex Hutch. & Dalziel (FWTA 1: 237) - obanensis Baker (FWTA 1: 238) ✩ - pobeguinii Aubrév. (FWTA 1: 238) ✩ - salacioides Lawson ex Hutch & Dalziel (FWTA 1: 238) - tabouensis Aubrév. (FWTA 1: 238) = ? Eugenia calophylloides - whytei Sprague (FWTA 1: 238) Syzygium guineense (Willd.) DC. var. guineense (Kirkia 10: 403) - guineense (Willd.) DC. var. occidentale F.White (Kirkia 10: 403) - owariense (P.Beauv.) Benth. (FWTA 1: 240) = S. rowlandii - rowlandii Sprague (FWTA 1: 240) - staudtii (Engl.) Mildbr. (FWTA 1: 240) = S. guineense

Nyctaginaceae Pisonia aculeata Linné (FWTA 1: 177)

Ochnaceae ✪ Campylospermum amplectens (Stapf) Farron (BJBB 35: 393)

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- calanthum (Gilg) Farron (BJBB 35: 394) - cf. vogelii (Vigne 1117 & 1902 (FHO)) - congestum (Oliver) Farron (BJBB 35: 394) - duparquetianum (Baill.) Tiegh. (Bot.Helv. 95: 67) - dybovskii Tiegh. (BJBB 35: 396) - flavum (Schum. & Thonn. ex Stapf) Farron (BJBB 35: 397) - glaberrimum (P.Beauv.) Farron (BJBB 35: 397) - laxiflorum (De Wild. & Th. Dur.) Tiegh. (BJBB 35: 399) - oliverianum (Gilg) Farron (BJBB 35: 400) - reticulatum (P.Beauv.) Farron var. reticulatum (BJBB 35: 400) - reticulatum (P.Beauv.) Farron var. turnerae (Hook.f.) Farron (BJBB 35: 401) - schoenleinianum (Klotsch) Farron (BJBB 35: 402) - squamosum (DC.) Farron (BJBB 35: 402) ✪ - subcordatum (Stapf) Farron (BJBB 35: 402) - sulcatum (Tiegh.) Farron (BJBB 35: 403) - vogelii (Hook.f.) Farron var. vogelii (BJBB 35: 403) ✩ Idertia morsonii (Hutch. & Dalziel) Farron (Bot.Helv. 95: 66) Lophira alata Banks ex Gaertn. (FWTA 1: 231) Ochna afzelii R.Br. ex Oliver (FWTA 1: 223) - kibbiensis Hutch. & Dalziel (FWTA 1: 223) = O. staudtii - membranacea Oliver (FWTA 1: 222) - multiflora DC. (FWTA 1: 223) - staudtii Engl. & Gilg. (Fl.Afr.Centr.: 21) Ouratea acutissima Gilg (FWTA 1: 229) = Rhabdophyllum affine - affinis (Hook.f.) Engl. (FWTA 1: 229) = Rhabdophyllum affine - amplectens (Stapf) Engl. (FWTA 1: 226) = Campylospermum a. - calophylla (Hook.f.) Engl. (FWTA 1: 229) = Rhabdophyllum calophyllum - congesta (Oliver) Engl. ex Gilg. (FWTA 1: 229) = Campylospermum congestum - duparquetiana (Baill.) Gilg (FWTA 1: 226) = Campylospermum duparquetianum - dybovskii (Tiegh.) Aké Assi (Candollea 55: 281) = Campylospermum d. - flava (Schum.) Hutch. & Dalziel (FWTA 1: 226) = Campylospermum flavum - glaberrima (P.Beauv.) Engl. ex Gilg (FWTA 1: 229) = Campylospermum glaberrimum - morsonii Hutch. & Dalziel (FWTA 1: 226) = Idertia m. - myrioneura Gilg (FWTA 1: 229) = Rhabdophyllum affine - reticulata (P.Beauv.) Engl. (FWTA 1: 228) = Campylospermum reticulatum - reticulata (P.Beauv.) Engl. var. turnerae (Hook.f.) J.B. Hall (Kew Bull. 35: 842) = Campylospermum reticulatum - schoenleiniana (Klotzsch) Gilg (FWTA 1: 226) = Campylospermum schoenleinianum - squamosa (DC.) Engl. (FWTA 1: 228) = Campylospermum squamosum - subcordata (Stapf) Engl. (FWTA 1: 226) = Campylospermum subcordatum - sulcata (Tiegh.) Keay (FWTA 1: 229) = Campylospermum sulcatum - turnerae (Hook.f.) Hutch. & Dalziel (FWTA 1: 228) = Campylospermum reticulatum - vogelii (Hook.f.) Engl. ex Gilg (FWTA 1: 229) = Campylospermum v. Rhabdophyllum affine (Hook.f.) Tiegh. (BJBB 35: 390) - calophyllum (Hook.f.) Tiegh. (BJBB 35: 392)

Olacaceae ✩ Aptandra zenkeri Engl. (FWTA 1: 649) Coula edulis Baill. (FWTA 1: 645) Heisteria parvifolia Sm. (FWTA 1: 645) ✩ Octoknema borealis Hutch. & Dalziel (FWTA 1: 656) - spec.a (Jongkind et al. 5079 (WAG)) Olax gambecola Baill. (FWTA 1: 647) ✩ - mannii Oliver (FWTA 1: 647) - subscorpioidea Oliver (FWTA 1: 647) ✩ Ongokea gore (Hua) Pierre (FWTA 1: 649) ✪ Ptychopetalum anceps Oliver (FWTA 1: 647) Strombosia glaucescens J.Léonard var. lucida J.Léonard (FWTA 1: 648) = S. pustulata - grandifolia Hook.f. ex Benth. (Aubrév. 1959) = S. pustulata - pustulata Oliver (Fl.Cameroun: 137) ✩ Strombosiopis nana Breteler (Kew Bull. 56: 751)

Oleaceae ✩ Chionanthus africanus (Knobl.) Stearn (Bot.J.Linn.Soc. 80: 197) ✩ - mannii (Soler.) Stearn var. mannii (Bot.J.Linn.Soc. 80: 199) - mannii (Baker) Stearn var. congestus (Baker) Stearn (Bot.J.Linn.Soc. 80: 201) - niloticus (Oliver) Stearn (Bot.J.Linn.Soc. 80: 202) ✩ Jasminum bakeri Scott-Elliot (FWTA 2: 50) - dichotomum Vahl (FWTA 2: 50) - pauciflorum Benth. (FWTA 2: 50) - preussii Engl. & Knobl. (FWTA 2: 51)

Linociera africana (Welw. ex Knobl.) Knobl. (FWTA 2: 48) = Chionanthus africanus - lingelsheimiana Gilg & Schellenb. (Aubrév. 1959) = Chionanthus mannii var. mannii - mannii Soler. (FWTA 2: 48) = Chionanthus mannii var. mannii - nilotica Oliver (FWTA 2: 48) = Chionanthus niloticus ✩ Olea capensis Linné ssp. hochstetteri (Baker) Friis & Green (Kew Bull. 41: 36) - guineensis Hutch. & C.A.Smith (Aubrév. 1959) = O. capensis spp. h. - hochstetteri Baker (FWTA 2: 49) = O. capensis spp. h. Schrebera arborea A.Cheval. (FWTA 2: 49)

Opiliaceae Opilia amanacea Roxb. (Willdenowia 12: 162) - celtidifolia (Guillaumet & Perr.) Endl. ex Walp (FWTA 1: 651) = O. amanacea Urobotrya afzelii (Engl.) Stapf ex Hutch. & Dalziel (FWTA 1: 652) = U. congolana spp. afzelii - congolana (Baill.) Hiepko ssp. afzelii (Engl.) Hiepko (Bot.Jahrb. 107: 144)

Orchidaceae Aerangis arachnopus (Rchb.f.) Schltr. (Fl. Cam. 36, 3: 844) - biloba (Lindl.) Schltr. (Fl. Cam. 36, 3: 846) - calantha (Schltr.) Schltr. (Fl. Cam. 36, 3: 840) - laurentii (De Wild.) Schltr. (FWTA 3: 265) = Summerhayesia l. Ancistrochilus rothschildianus O’Brien (Fl. Cam. 36, 2: 326) Ancistrorhynchus capitatus (Lindl.) Summerh. (Fl. Cam. 36, 3: 774) - cephalotes (Rchb.f.) Summerh. (FWTA 3: 272) - clandestinus (Lindl.) Schltr. (Fl. Cam. 36, 3: 768) - metteniae (Kraenzl.) Summerh. (Fl. Cam. 36, 3: 778) - recurvus Finet (Fl. Cam. 36, 3: 770) - strausii (Schltr.) Schltr. (Fl. Cam. 36, 3: 770) Angraecopsis elliptica Summerh. (Adansonia sér. 2, 15: 204) - ischnopus (Schltr.) Schltr. (Fl. Cam. 36, 3: 912) - macrophylla Summerh. (FTEA: 598) Angraecum angustipetalum Rendle (Fl. Cam. 36, 3: 886) - bancoense Burg (Misc.pap.LUW 19: 26) - birrimense Rolfe (Fl. Cam. 36, 3: 896) - chevalieri Summerh. (Fl. Cam. 36, 3: 886) - claessensii De Wild. (Fl. Cam. 36, 3: 904) - distichum Lindl. (Fl. Cam. 36, 3: 884) - moandense De Wild. (Flore d’AC: 473) = A. chevalieri ✩ - modicum Summerh. (FWTA 3: 257) - multinominatum Rendle (Fl. Cam. 36, 3: 904) - podochiloides Schltr. (Fl. Cam. 36, 3: 880) - pyriforme Summerh. (FWTA 3: 254) - subulatum Lindl. (Fl. Cam. 36, 3: 876) Ansellia africana Lindl. (Fl. Cam. 36, 2: 592) Auxopus kamerunensis Schltr. (Fl. Cam. 36, 1: 306) - macranthus Summerh. (Fl. Cam. 36, 1: 308) Bolusiella batesii (Rolfe) Schltr. (Fl. Cam. 36, 3: 684) - imbricata (Rolfe) Schltr. (FWTA 3: 263) =B. maudea - iridifolia (Rolfe) Schltr. (Fl. Cam. 36, 3: 680) - maudea (H.Bolus) Schltr. (Fl. Cam. 36, 3: 682) - talbotii (Rendle) Summerh. (Fl. Cam. 36, 3: 686) Brachycorythis kalbreyeri Rchb.f. (Fl. Cam. 36, 1: 65) - macrantha (Lindl.) Summerh. (Fl. Cam. 36, 1: 64) Bulbophyllum acutebracteatum De Wild. var. acutebracteatum (Orch. Monogr. 2 Monogr.: 143) - barbigerum Lindl. (Orch. Monogr. 2: 36) - bidenticulatum Verm. (Fl. Cam. 36, 2: 431) - bufo (Lindl.) Rchb.f. (Fl. Cam. 36, 2: 413) - buntingii Rendle (FWTA 3: 234) = B. oxychilum - calamarium Lindl. (FWTA 3: 236) = B. saltatorium - calyptratum Kraenzl. (Orch. Monogr. 2: 128) ✩ - calyptratum Kraenzl. var. graminifolium (Summerh.) Verm. (Orch. Monogr. 2: 130) ✩ - calyptratum Kraenzl. var. lucifugum (Summerh.) Verm. (Orch. Monogr. 2: 131) - carnosisepalum Verm. (Orch. Monogr. 2: 133) - cochleatum Lindl. (Orch. Monogr.: 39) - cocoinum Batem. ex. Lindl. (Orch. Monogr. 2: 66) - colubrinum (Rchb.f.) Rchb.f. (Orch. Monogr. 2: 96) - comatum Lindl. (Orch. Monogr. 2: 85) - congolanum Schltr. (FWTA 3: 239) = B. scaberulum - daloaense Cribb & Perez-Vera (Adansonia sér. 2, 15: 200) = B. resupinatum ✩ - denticulatum Rolfe (Orch. Monogr. 2: 83) - distans Lindl. (FWTA 3: 236) = B. saltatorium - falcatum (Lindl.) Rchb.f. (Orch. Monogr. 2: 122) - falcipetalum Lindl. (Orch. Monogr. 2: 113)

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- finetii Szlach. & Olszewski (Fl. Cam. 36, 2: 395) - flavidum Lindl. (FWTA 3: 234) = B. pumilum - fuscum Lindl. (Orch. Monogr. 2: 145) - graminifolium Summerh. (FWTA 3: 241) = B. calyptratum var. g. - imbricatum Lindl. (Orch. Monogr. 2: 103) - inflatum Rolfe (FWTA 3: 236) = B. comatum - intertextum Lindl. (Orch. Monogr. 2: 54) - irigaleae Cribb & Perez-Vera (Adansonia sér. 2, 15: 200) = B. schinzianum - ivorense Cribb & Perez-Vera (Orch. Monogr. 2: 73) - josephii (Kuntze) Summerh. var. mahonii (Rolfe) Verm. (Orch. Monogr. 2: 69) ✩ - linderi Summerh. (FWTA 3: 242) = not a synonym of B. imbricatum Lindl. - lucifugum Summerh. (FWTA 3: 241) = B. calyptratum var. l. - lupulinum Lindl. (Orch. Monogr. 2: 138) - magnibracteatum Summerh. (Orch. Monogr. 2: 106) - maximum (Lindl.) Rchb.f. (Orch. Monogr. 2: 97) - melanorrhachis (Rchb.f.) Rchb.f. (FWTA 3: 241) = B. falcatum - melinostachyum Schltr. (Fl. Cam. 36, 2: 443) - nigritianum Rendle (Orch. Monogr. 2: 78) - oreonastes Rchb.f. (Orch. Monogr. 2: 149) - oxychilum Schltr. (Orch. Monogr. 2: 76) - oxypterum (Lindl.) Rchb.f. (FWTA 3: 241) = B. maximum ✩ - parvum Summerh. (Orch. Monogr. 2: 121) - phaeopogon Schltr. (Fl. Cam. 36, 2: 387) = B. schinzianum - pipio Rchb.f. (Orch. Monogr. 2: 58) - porphyroglossum Kraenzl. (Adansonia sér. 2, 15: 206) = B. pumilum - pumilum (Swartz) Lindl. (Fl. Cam. 36, 2: 469) - purpureorhachis (De Wild.) Schltr. (Orch. Monogr. 2: 112) - recurvum Lindl. (FWTA 3: 234) = B. pumilum - resupinatum Ridley (Orch. Monogr. 2: 118) - rhizophorae Lindl. (FWTA 3: 239) = B. falcatum - saltatorium Lindl. (Orch. Monogr. 2: 31) - sandersonii (Hook.f.) Rchb.f. (Orch. Monogr. 2: 107) - scaberulum (Rolfe) Bolus (Orch. Monogr. 2: 114) - scariosum Summerh. (Orch. Monogr. 2: 81) - schinzianum Kraenzl. (Orch. Monogr. 2: 26) - schinzianum Kraenzl. var. phaeopogon (Schltr.) Verm. (Orch. Monogr. 2: 29) = B. phaeopogon - stenopetalum Kraenzl. (Fl. Cam. 36, 2: 399) - tentaculigerum Rchb.f. (FWTA 3: 239) = B. sandersonii - tetragonum Lindl. (Orch. Monogr. 2: 140) - velutinum (Lindl.) Rchb.f. (Fl. Cam. 36, 2: 411) = B. falcatum - victoris Cribb & Perez-Vera (Adansonia sér. 2, 15: 202) = B. resupinatum - winkleri Schltr. (FWTA 3: 234) = B. pumilum - zenkeranum Kraenzl. (FWTA 3: 239) = B. oreonastes Calanthe corymbosa Lindl. (FWTA 3: 226) = C. sylvatica - sylvatica (Thouars) Lindl. (Fl. Cam. 36, 2: 330) Calyptrochilum christyanum (Rchb.f.) Summerh. (Fl. Cam. 36, 3: 816) - emarginatum (Swartz) Schltr. (Fl. Cam. 36, 3: 810) Chamaeangis ichneumonea (Lindl.) Schltr. (Fl. Cam. 36, 3: 753) - lanceolata Summerh. (Adansonia sér. 2, 15: 208) - odoratissima (Rchb. fil.) Schltr. (Fl. Cam. 36, 3: 756) - vesicata (Lindl.) Schltr. (Fl. Cam. 36, 3: 760) Chauliodon buntingii Summerh. (FWTA 3: 260) = C. deflexicalcaratum - deflexicalcaratum (De Wild.) Johnsson (Fl. Cam. 36, 3: 809) Cheirostylis divina (Guinée) Summerh. (Fl. Cam. 36, 1: 250) Corymborkis corymbis Thouars (Fl. Cam. 36, 1: 238) Cribbia brachyceras (Summerh.) Senghas (Fl. Cam. 36, 3: 920) - confusa Cribb (Fl. Cam. 36, 3: 920) Cyrtorchis arcuata (Lindl.) Schltr. var. variabilis (Summerh.) Geerinck (Fl.Cam. 36, 3: 726) - arcuata (Lindl.) Schltr. var. whytei (Rolfe) Geerinck (Fl. Cam. 36, 3: 726) - aschersonii (Kraenzl.) Schltr. (Fl. Cam. 36, 3: 720) - brownii (Rolfe) Schltr. (FTEA: 581) = Homocolleticon b. - chailluana (Hook. fil.) Schltr. (Fl. Cam. 36, 3: 724) ✪ - hamata (Rolfe) Schltr. (FWTA 3: 269) - monteiroae (Rchb. fil.) Schltr. (FTEA: 578) = Homocolleticon m. - ringens (Rchb.f.) Summerh. (FTEA: 580) = Homocolleticon r. Diaphananthe bidens (Swartz) Schltr. (Fl. Cam. 36, 3: 742) - bueae (Schltr.) Schltr. (Fl. Cam. 36, 3: 738) - curvata (Rolfe) Summerh. (FWTA 3: 261) = Rhipidoglossum curvatum - densiflora (Summerh.) Summerh. (FTEA: 533) = Rhipidoglossum densiflorum - laticalcar J.B.Hall (Kew Bull. 29: 427) ✩ - laxiflora (Summerh.) Summerh. (FWTA 3: 263) - pellucida (Lindl.) Schltr. (Fl. Cam. 36, 3: 748) - quintasii (Rolfe) Schltr. (FWTA 3: 261) = D. rohrii - rohrii (Rchb.f.) Summerh. (FTEA: 524)

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- rutila (Rchb.f.) Summerh. (FTEA: 536) = Rhipidoglossum rutilum - sarcorhynchoides J.B.Hall (Kew Bull. 29: 428) - suborbicularis Summerh. (FWTA 3: 261) Diceratostele gabonensis Summerh. (Fl. Cam. 36, 1: 271) Didymoplexis africana Summerh. (Fl. Cam. 36, 1: 309) Dinklageella liberica Mansf. (Fl. Cam. 36, 3: 820) ✩ - minor Summerh. (FWTA 3: 270) Disperis thomensis Summerh. (Fl. Cam. 36, 1: 44) - togoensis Schltr. (Fl. Cam. 36, 1: 38) Eggelingia clavata Summerh. (Fl. Cam. 36, 3: 690) Encheiridion macrorrhynchium (Schltr.) Summerh. (FWTA 3: 259) = Microcoelia macrorrhynchia Epipogium roseum (D. Don) Lindl. (Fl. Cam. 36, 2) Eulophia euglossa (Rchb.f.) Rchb.f. (Fl. Cam. 36, 2: 628) - galeoloides Kraenzl. (FTEA: 459) - gracilis Lindl. (Fl. Cam. 36, 2: 642) - guineensis Lindl. (Fl. Cam. 36, 2: 606) - horsfallii (Bateman) Summerh. (Fl. Cam. 36, 2: 622) Eulophidium latifolium (Rolfe) Summerh. (FWTA 3: 243) = Oeceoclades ugandae - maculatum (Lindl.) Pfitzer (FWTA 3: 243) = Oeceoclades maculata - saundersianum (Rchb.f.) Summerh. (FWTA 3: 243) = Oeceoclades saundersiana Eurychone rothschildiana (O’Brien) Schltr. (Fl. Cam. 36, 3: 736) Genyorchis apetala (Lindl.) Verm. (Fl. Cam. 36, 2: 487) - pumila (Swartz) Schltr. (FTEA: 325) = G. apetala Graphorchis lurida (Swartz) O.Kuntze (Fl. Cam. 36, 2: 600) Habenaria barrina Ridley (FWTA 3: 194) - buettnerana Kraenzl. (Fl. Cam. 36, 1: 143) - buntingii Rendle (Fl. Cam. 36, 1: 146) - dinklagei Kraenzl. (FWTA 3: 194) - englerana Kraenzl. (Fl. Cam. 36, 1: 115) ✩ - jacobii Summerh. (FWTA 3: 194) ✩ - leonensis Dur. & Schinz (FWTA 3: 193) - macrandra Lindl. (FWTA 3: 193) = Podandriella m. - malacophylla Rchb.f. (Fl. Cam. 36, 1: 140) - procera (Swartz) Lindle (Fl. Cam. 36, 1: 124) Hetaeria heterosepala (Rchb.f.) Summerh. (Fl. Cam. 36, 1: 262) - occidentalis Summerh. (Fl. Cam. 36, 1: 265) = Zeuxine o. - stammleri (Schltr.) Summerh. (FWTA 3: 210) = Zeuxine s. Homocolleticon brownii (Rolfe) Szlach. & Olszewski (Fl. Cam. 36, 3: 728) - monteiroae (Rchb. fil.) Szlach. & Olszewski (Fl. Cam. 36, 3: 728) - ringens (Rchb. fil.) Szlach. & Olszewski (Fl. Cam. 36, 3: 730) Kornasia chevalieri (Summerh.) Szlach. (Fl. Cam. 36, 2: 376) - maclaudii (Finet) Szlach. (Fl. Cam. 36, 2: 374) Liparis caillei Finet (Fl. Cam. 36, 2: 344) - epiphytica Schltr. (Fl. Cam. 36, 2: 358) - platyglossa Schltr. (Fl. Cam. 36, 2: 350) - tridens Kraenzl. (Fl. Cam. 36, 2: 342) Lisowskia katangensis (Summerh.) Szlach. (Fl. Cam. 36, 2: 368) - prorepens (Kraenzl.) Szlach. (Fl. Cam. 36, 2: 368) Listrostachys pertusa (Lindl.) Rchb.f. (Fl. Cam. 36, 3: 786) Malaxis chevalieri Summerh. (Kew Bull. 32: 738) = Kornasia c. - maclaudii (Finet) Summerh. (Kew Bull. 32: 738) = Kornasia m. ✪ - melanotoessa Summerh. (Kew Bull. 32: 740) - prorepens (Kraenzl.) Summerh. (FTEA: 288) = Lisowskia p. Manniella gustavii Rchb.f. (Fl. Cam. 36, 1: 268) Microcoelia caespitosa (Rolfe) Summerh. (Fl. Cam. 36, 3: 806) - dahomeensis (Finet) Summerh. (FWTA 3: 259) = M. konduensis - konduensis (De Wild.) Summerh. (Fl. Cam. 36, 3: 800) - macrorrhynchia (Schltr.) Summerh. (Fl. Cam. 36, 3: 796) Nephrangis filiformis (Kraenzl.) Summerh. (Fl. Cam. 36, 3: 696) Nervilia adolphii Schltr. (Orch. Monogr. 5: 51) - bicarinata (Blume) Schltr. (Orch. Monogr. 5: 56) - crociformis (Zoll. & Mor.) Seidenf. (Orch. Monogr. 5: 44) - fuerstenbergiana Schltr. (Orch. Monogr. 5: 54) - reniformis Schltr. (FWTA 3: 206) = N. crociformis - subintegra Summerh. (Orch. Monogr. 5: 50) - toiliezae N. Hallé (Adansonia sér. 2, 11: 455) = N. subintegra - umbrosa (Rchb. fil.) Schltr. (FWTA 3: 207) = N. bicarinata Oberonia disticha (Lam.) Szlach. (Fl. Cam. 36, 2: 378) Oeceoclades maculata (Lindl.) Lindl. (Fl. Cam. 36, 2: 596) = Eulophidium m. - saundersiana (Rchb.f.) Garay & Taylor (Fl. Cam. 36, 2: 598) = Eulophidium s. - ugandae (Rolfe) Garay & Taylor (FTEA: 420) Platylepis glandulosa (Lindl.) Rchb.f. (Fl. Cam. 36, 1: 244) Plectrelminthus caudatus (Lindl.) Summerh. var. caudatus (Fl. Cam. 36, 3: 826) Podandriella macrandra (Lindl.) Szlach. & Olszewski (Fl. Cam. 36, 1: 194) ✩ ✪

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Podangis dactyloceras (Rchb. fil.) Schltr. (Fl. Cam. 36, 3: 688) Polystachya adansoniae Rchb.f. (Fl. Cam. 36, 2: 573) - affinis Lindl. var. affinis (Fl. Cam. 36, 2: 502) - affinis Lindl. var. nana J.B.Hall (Kew Bull. 29: 428) ✪ - bancoensis Burg (Misc.pap.LUW 19: 23) - cultriformis (Thouars) Sprengel (Fl. Cam. 36, 2: 515) ✪ - dalzielii Summerh. (FWTA 3: 221) - dolichophylla Schltr. (Fl. Cam. 36, 2: 568) ✩ - elastica Lindl. (FWTA 3: 223) - fractiflexa Summerh. (BJBB 50: 103) - fusiformis (Thouars) Lindl. (Fl. Cam. 36, 2: 552) - galeata (Swartz) Rchb.f (Fl. Cam. 36, 2: 521) - golungensis Rchb.f. (Fl. Cam. 36, 2: 554) - inconspicua Rendle (FWTA 3: 225) = P. tenuissima - laxiflora Lindl. (Fl. Cam. 36, 2: 542) - leonensis Rchb.f. ((FWTA 3: 224) - microbambusa Kraenzl. (Fl. Cam. 36, 2: 504) - modesta Rchb.f. (Fl. Cam. 36, 2: 556) ✩ - monolenis Summerh. (FWTA 3: 223) - mukandaensis De Wild. (Fl. Cam. 36, 2: 558) - mystacioides De Wild. (Fl. Cam. 36, 2: 551) - obanensis Rendle (Fl. Cam. 36, 2: 500) - odorata Lindl. (Fl. Cam. 36, 2: 566) - paniculata (Swartz) Rolfe (Fl. Cam. 36, 2: 530) - parva Summerh. (FWTA 3: 224) ✩ - pobeguinu (Finet) Rolfe (FWTA 3: 223) - polychaete Kraenzl. (Fl. Cam. 36, 2: 576) ✪ - pseudo-disa Kraenzl. (FWTA 3: 221) ✩ - puberula Lindl. (Fl. Cam. 36, 2: 564) - ramulosa Lindl. (Fl. Cam. 36, 2: 562) ✩ - reflexa Lindl. (FWTA 3: 223) - rhodoptera Rchb.f. (Fl. Cam. 36, 2: 532) - saccata (Finet) Rolfe (FWTA 3: 223) - seticaulis Rendle (Fl. Cam. 36, 2: 584) - subulata Finet (Fl. Cam. 36, 2: 570) - tenuissima Kraenzl. (Fl. Cam. 36, 2: 519) - tessellata Lindl. (Fl. Cam. 36, 2: 560) - victoriae Kraenzl. (Adansonia sér. 2, 15: 210) Rangaeris brachyceras (Summerh.) Summerh. (FWTA 3: 266) = Cribbia b. - longicaudata (Rolfe) Summerh. (Adansonia sér. 2, 15: 211) - muscicola (Rchb.f.) Summerh. (Fl. Cam. 36, 3: 676) - rhipsalisocia (Rchb.f.) Summerh. (Fl. Cam. 36, 3: 672) Rhipidoglossum curvatum (Rolfe) Garay (Fl. Cam. 36, 3: 856) - densiflorum Summerh. (Fl. Cam. 36, 3: 864) ✩ - paucifolium D. Johansson (Acta Phyt. Suec. 59: 40) - rutilum (Rchb. fil.) Schltr. (Fl. Cam. 36, 3: 872) Solenangis clavata (Rolfe) Schltr. (Fl. Cam. 36, 3: 822) - scandens (Schltr.) Schltr. (Fl. Cam. 36, 3: 824) Stolzia elaidum (Lindl.) Summerh. (Kew Bull. 33: 84) - repens (Rolfe) Summerh. (Fl. Cam. 36, 2: 338) Summerhayesia laurentii (De Wild.) Cribb (Fl. Cam. 36, 3: 828) Taeniophyllum coxii (Summerh.) Summerh. (Bot. Notiser 132: 512) Tridactyle anthomaniaca (Rchb.f.) Summerh. (Fl. Cam. 36, 3: 704) ✩ - armeniaca (Lindl.) Schltr. (FWTA 3: 274) - bicaudata (Lindl.) Schltr. (Fl. Cam. 36, 3: 708) - brevicalcarata Summerh. (FWTA 3: 274) - crassifolia Summerh. (Fl. Cam. 36, 3: 702) - filifolia (Schltr.) Schltr. (Fl. Cam. 36, 3: 700) ✩ - fusifera Mansf. (FWTA 3: 274) - gentilii (De Wild.) Schltr. (Fl. Cam. 36, 3: 714) - oblongifolia Summerh. (FTEA: 610) - tridactylites (Rolfe) Schltr. (Fl. Cam. 36, 3: 712) - tridentata (Harvey) Schltr. (FWTA 3: 274) = T. filifolia Vanilla africana Lindl. subsp. africana (Fl. Cam. 36, 1: 285) - africana Lindl. subsp. racemosa (Rolfe) Szlach. & Olsz (Fl. Cam. 36, 1: 286) - africana Lindl. (FWTA 3: 205) - crenulata Rolfe (FWTA 3: 206) = V. africana - imperialis Kraenzl. (Fl. Cam. 36, 1: 290) - ramosa Rolfe (FWTA 3: 205) = V. africana Zeuxine elongata Rolfe (Fl. Cam. 36, 1: 256) - heterosepala (Rchb.f.) Geerinck (BJBB 50: 120) = Hetaeria h. - occidentalis (Summerh.) Geerinck (BJBB 50: 122) = Hetaeria o. - stammleri Schltr. (Fl. Cam. 36, 1: 254)

Oxalidaceae ✩ Biophytum talbotii (Baker f.) Hutch. & Dalziel (FWTA 1: 159)

Palmae Ancistrophyllum laeve (Mann & Wendl.) Drude (FWTA 3: 167) = Laccosperma l. - opacum (Mann & Wendl.) Drude (FWTA 3: 167) = Laccosperma o. - secundiflorum (P.Beauv.) Wendl. (FWTA 3: 167) = Laccosperma s. Calamus deeratus Mann & Wendl. (FWTA 3: 166) Elaeis guineensis Jacq. (FWTA 3: 161) Eremospatha acutiflorum (Becc.) J. Dransf (Sunderland 2001: 109) ✩ - hookeri (Mann & Wendl.) Wendl. (FWTA 3: 168 & Sunderland 2001: 52) - laurentii De Wildem. (Sunderland 2001: 61) - macrocarpa (Mann & Wendl.) Wendl. (FWTA 3: 168 & Sunderland 2001: 74) Laccosperma laeve G.Mann & H.Wendl. (Kew Bull. 37: 456 & Sunderland 2001: 106) - opacum G.Mann & H.Wendl. (Kew Bull. 37: 456 & Sunderland 2001: 100) - secundiflorum (P.Beauv.) O.Kuntze (Kew Bull. 37: 456 & Sunderland 2001: 119) Raphia hookeri Mann & Wendl. (FWTA 3: 162) ✪ - palma-pinus (Gaertn.) Hutch. (FWTA 3: 162) - sassandrensis A.Cheval. (Aubrév. 1959) = R. hookeri ✩ Sclerosperma mannii H.Wendl. (FWTA 3: 161)

Pandaceae ✩ Microdesmis keayana J.Léonard (BJBB 31: 180) - puberula Hook.f. ex Planch. (FWTA 1: 392) = M. keayana Panda oleosa Pierre (FWTA 1: 636)

Pandanaceae ✩ Pandanus abbiwii Huynh (Bot.Helv. 97: 81) ✩ - akeassii Huynh (Bot.Jahrb. 109: 345) - candelabrum P.Beauv. (FWTA 3: 170) - heudelotianus (Gaud.) Balf (Aubrév. 1959) ✩ - lachaisei Huynh (Bot.Jahrb. 109: 351) ✩ - sierraleonensis Huynh (Bot.Helv. 98: 174) ✩ - tiassaleensis Huynh (Bot.Jahrb. 109: 354)

Passifloraceae Adenia cissampeloides (Planch. ex Benth.) Harms (Med.LUW 71-18: 246) - dinklagei Hutch. & Dalziel (Med.LUW 71-18: 253) - gracilis Harms (Med.LUW 71-18: 255) - guineensis W.de Wilde (Med.LUW 71-18: 259) - lobata (Jacq.) Engl. (Med.LUW 71-18: 149) - mannii (Mast.) Engl. (Med.LUW 71-18: 144) - rumicifolia Engl. & Harms (Med.LUW 71-18: 154) - tenuispira (Stapf) Engl. (FWTA 1: 202) = A. mannii ✩ Androsiphonia adenostegia Stapf (FWTA 1: 201) ✪ Crossostemma laurifolium Planch. ex Benth. (FWTA 1: 202) Paropsia guineensis Oliver (BJBB 40: 54) Smeathmannia laevigata Soland. ex R.Br. var. laevigata (FWTA 1: 200) - laevigata Soland. ex R.Br. var. nigerica A.Cheval. ex Hutch. & Dalziel (FWTA 1: 200) - pubescens Soland. ex R.Br. (FWTA 1: 200) ✩

Phytolaccaceae Hilleria latifolia (Lam.) H.Walt. (FWTA 1: 143) Phytolacca dodecandra L’Herit. (FWTA 1: 143)

Piperaceae ✩ Peperomia fernandopoiana C.DC. (Bot.Jahrb. 93: 113) ✩ - molleri C.DC. (Bot.Jahrb. 9: 106) ✩ - retusa (Linné f.) Dietr. (Bot.Jahrb. 9: 89) ✩ - rotundifolia (Linné) H.B. & K. (Bot.Jahrb. 9: 85) ✩ - tetraphylla (Forster) Hooker & Arnott (Bot.Jahrb. 9: 72) ✩ - vulcanica Baker & Wright (Bot.Jahrb. 9: 102) Piper capense Linné f. (FWTA 1: 84) - guineense Schum. & Thonn. (FWTA 1: 84) - umbellatum Linné (FWTA 1: 84)

Pittosporaceae Pittosporum viridiflorum Sims ssp. dalzielii (Hutch.) Cuf. (FWTA 1: 182)

Podostemaceae Inversodicraea adamesii G.Taylor (FWTA 1: 127) = Ledermanniella a. - bowlingii J.B.Hall (Kew Bull. 26: 126) = Ledermanniella b. - garrettii (Wright) G.Taylor (FWTA 1: 126) = Macropodiella g. - ledermannii (Engl.) Engl. (FWTA 1: 127) = Ledermanniella l. - macrothyrsa G.Taylor (FWTA 1: 127) = Macropodiella m. ✩ Ledermanniella abbayesii (Taylor) Cusset (Bull.MNHN Paris sér. 4, 5: 385)

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✩ - adamesii (Taylor) Cusset (Bull.MNHN Paris sér. 4, 5: 388) ✩ - bowlingii (J.B.Hall) Cusset (Bull.MNHN Paris sér. 4, 6: 268) ✩ - harrisii Cusset (Bull.MNHN Paris sér. 4, 5: 374) ✩ - ledermannii (Engl.) Cusset (Adansonia sér. 2, 14: 274) ✩ - minutissima Cusset (Bull.MNHN Paris sér. 4, 6: 252) ✩ - mortonii Cusset (Bull.MNHN Paris sér. 4, 5: 368) ✩ - taylorii Cusset (Bull.MNHN Paris sér. 4, 5: 376) ✪ Macropodiella garrettii (Wright) Cusset (Adansonia sér. 2, 17: 302) - heteromorpha (Baill.) Cusset (Adansonia sér. 2, 17: 298) ✩ - macrothyrsa (Taylor) Cusset (Adansonia sér. 2, 17: 296) ✩ - taylorii Cusset (Adansonia sér. 2, 17: 298) ✩ Polypleurum submersum J.B.Hall (Kew Bull. 26: 130) ✪ Saxicolella amicorum J.B.Hall (Kew Bull. 26: 133)

Polygalaceae Atroxima afzeliana (Oliver) Stapf (Med.LHW 77-18: 15) ✩ - liberica Stapf (Med.LHW 77-18: 19) ✩ Carpolobia alba G.Don (Med.LHW 77-18: 24) - lutea G.Don (Med.LHW 77-18: 38) Securidaca welwitschii Oliver (FWTA 1: 110)

Polygonaceae Afrobrunnichia erecta (Asch.) Hutch. & Dalziel (FWTA 1: 138) Symmeria paniculata Benth. (FWTA 1: 138)

Ranunculaceae Clematis grandiflora DC. (FWTA 1: 64) - hirsuta Guillaumet & Perr. (FWTA 1: 64)

Rapateaceae ✪ Maschalocephalus dinklagei Gilg & K.Schum. (FWTA 3: 55)

Rhamnaceae Gouania longipetala Hemsl. (FWTA 1: 670) ✩ Lasiodiscus chevalieri Hutch. (Kew Bull. 50: 495) - fasciculiflorus Engl. (Kew Bull. 50: 499) ✪ - mannii Hook.f. (Kew Bull. 50: 505) - marmoratus Wright (Aubrév. 1959) = L. mannii - mildbraedii Engl. (FWTA 1: 671) = L. mannii ✩ Maesopsis eminii Engl. (FWTA 1: 669) Ventilago africana Exell (FWTA 1: 670)

Rhizophoraceae Anopyxis klaineana (Pierre) Engl. (FWTA 1: 286) ✩ Cassipourea adami Jacq.-Félix (FWTA 1: 283) ✪ - afzelii (Oliver) Alston (FWTA 1: 283) - barteri (Hook.f.) N.E.Br. (FWTA 1: 283) - congoensis R.Br. ex DC. (FWTA 1: 283) ✩ - firestoneana Hutch. & Dalziel ex Cooper & Rec (FWTA 1: 283) - glabra Alston (FWTA 1: 283) = C. gummiflua ✩ - gummiflua Tulasne (Kew Bull. 1955: 147) ✪ - hiotou Aubrév. & Pellegr. (Bull.Soc.Bot.Fr.105: 34) ✩ - lescotiana J.G.Adam (Bull.IFAN 33: 496) ✩ - nialatou Aubrév. & Pellegr. (FWTA 1: 283) ✩ - paludosa Hutch. & Dalziel ex Jacq.-Félix (FWTA 1: 283) - sp.B. (FWTA 1: 283) = C. congoensis Rhizophora harrisonii Leechman (FWTA 1: 285) - mangle Linné (FWTA 1: 285) - racemosa G.F.W.Mey. (FWTA 1: 285)

Rosaceae Rubus fellatae A.Cheval. (FWTA 1: 426) - pinnatus Willd. var. afrotropicus (Engl.) C.E.Gust. (FWTA 1: 426)

Rubiaceae Aidia genipiflora (DC.) Dandy (FWTA 2: 114) ✪ Argocoffeopsis afzelii (Hiern) Robbr. (BJBB 51: 365) - eketensis (Wernh.) Robbr. (BJBB 56: 158) - jasminoides (Welw. ex Hiern) Robbr. (BJBB 51: 368) = A. eketensis ✪ - lemblinii (A.Cheval.) Robbr. (BJBB 51: 365) ✩ - rupestris (Hiern) Robbr. var. rupestris (BJBB 51: 369) ✩ Atractogyne bracteata (Wernh.) Hutch. & Dalziel (FWTA 2: 158) ✩ Aulacocalyx divergens (Hutch. & Dalziel) Keay (Kew Bull. 52: 655) - jasminiflora Hook.f. (Kew Bull. 52: 647) Belonophora coffeoides Hook.f. ssp. hypoglauca (Welw. ex Hiern) Dawson & Cheek

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(Kew Bull. 55: 77) - hypoglauca (Welw. ex Hiern) A.Cheval. (FWTA 2: 158) = B. coffeoides ssp. hypoglauca - lepidopoda Hutch. & Dalziel (Aubrév. 1959) = B. coffeoides ssp. hypoglauca ✩ Bertiera adamsii (Hepper) N.Hallé (FWTA 2: 160) - bracteolata Hiern (FWTA 2: 159) - breviflora Hiern (FWTA 2: 159) ✩ - chevalieri Hutch. & Dalziel (FWTA 2: 160) ✩ - fimbriata (A.Cheval. ex Hutch. & Dalziel) Hepper (FWTA 2: 160) - montana Hiern (Aubrév. 1959) = B. racemosa var. racemosa - racemosa (G.Don) K.Schum. var. glabrata (K.Schum.) Hutch. & Dalziel (32: 160) - racemosa (G.Don) K.Schum. var. racemosa (FWTA 2: 160) - spec.nov. (Jongkind 5490 (WAG)) ✪ - spicata (Gaertn.f.) Wernh. (FWTA 2: 159) Borreria intricans Hepper (FWTA 2: 222) = Spermacoce i. ✩ Calycosiphonia macrochlamys (K.Schum.) Leroy (BJBB 51: 377) - spathicalyx (K.Schum.) Robbr. (BJBB 51: 373) Canthium arnoldianum (De Wild. & Th. Dur.) Hepper (FWTA 2: 184) = Psydrax arnoldiana - cornelia Cham. & Schlecht. (FWTA 2: 184) = Keetia c. - gueinzii Sonder (FWTA 2: 184) = Keetia venosissima - henriquesianum (K.Schum.) G.Taylor (FWTA 2: 181) = Psydrax kraussioides - hispidum Benth. (FWTA 2: 182) = Keetia hispida - horizontale (Schum. & Thonn.) Hiern (FWTA 2: 182) = Psydrax horizontalis - manense Aubrév. & Pellegr. (FWTA 2: 184) = Psydrax manensis - mannii Hiern (FWTA 2: 182) = Keetia m. - multiflorum (Schum. & Thonn.) Hiern (FWTA 2: 182) = Keetia multiflora - orthacanthum (Mildbr.) Robyns (FWTA 2: 182) = Vangueriella orthocantha - pobeguinii Hutch. & Dalziel (FWTA 2: 182) = Multidentia p. - rubens Hiern (FWTA 2: 184) = Keetia r. - rufivillosum Robyns ex Hutch. & Dalziel (FWTA 2: 182) = Keetia rufivillosa - setosum Hiern (FWTA 2: 182) = Keetia hispida - subcordatum DC. (FWTA 2: 184) = Psydrax subcordata - tekbe Aubrév. & Pellegr. (Aubrév. 1959) = Psydrax arnoldiana - venosum (Oliver) Hiern (FWTA 2: 184) = Keetia venosa - vulgare (K.Schum.) Bullock (FWTA 2: 184) = Psydrax spp. - zanzibaricum Klotszch (FWTA 2: 184) = Keetia spp. Cephaelis abouabouensis Schnell (FWTA 2: 205) = Psychotria a. - biaurita (Hutch. & Dalziel) Hepper (FWTA 2: 205) = Psychotria b. - mangenotii Aké Assi (FWTA 2: 204) = Psychotria m. - micheliae J.G.Adam (IFAN 35: 83) = Psychotria spec.?? - ombrophila (Schnell) Schnell (Mem. IFAN 50: 57) = Psychotria o. - peduncularis Salisb. (FWTA 2: 204) = Psychotria p. - schnellii Aké Assi (FWTA 2: 204) = Psychotria s. - spathacea Hiern (FWTA 2: 205) = Psychotria s. - yapoensis (Schnell) Schnell (FWTA 2: 204) = Psychotria y. ✩ Chassalia afzelii (Hiern) K.Schum. (FWTA 2: 192) ✪ - corallifera (A.Cheval. ex De Wild.) Hepper (FWTA 2: 192) ✩ - elongata Hutch. & Dalziel (FWTA 2: 192) - hallïi Jongkind spec.nov (Jongkind 4478 (WAG)) - kolly (Schum.) Hepper (FWTA 2: 192) ✩ - laxiflora Benth. (FWTA 2: 192) - subherbacea (Hiern) Hepper (FWTA 2: 192) Chazaliella abrupta (Hiern) Petit & Verdc. (Kew Bull. 31: 811) ✪ - cupulicalyx Verdc. (Kew Bull. 31: 799) ✩ - domatiicola (De Wild.) Petit (Kew Bull. 31: 810) ✩ - insidens (Hiern) Petit & Verdc. ssp. liberica Verdc. (Kew Bull. 31: 797) ✩ - lophoclada (Hiern) Petit & Verdc. (Kew Bull. 31: 810) - sciadephora (Hiern) Petit & Verdc. (Kew Bull. 31: 790) - tomentella Hawthorne spec.nov. (to be published) - sp.B. (Kew Bull. 31: 817) Coffea afzelii Hiern (FWTA 2: 156) = Argocoffeopsis a. - canephora Pierre ex Froehner (FWTA 2: 154) - ebracteolata (Hiern) Brenan (FWTA 2: 157) = Psilanthus ebracteolatus - eketensis Wernh. (FWTA 2: 156) = Argocoffeopsis e. ✩ - humilis A.Cheval. (FWTA 2: 156) - jasminoides Welw. (Aubrév. 1959) = Argocoffeopsis eketensis - lamtoensis Porteres (Kew Bull. 42: 455) = Psilanthus? - lemblini (A.Cheval.) Keay (FWTA 2: 156) = Argocoffeopsis l. - liberica Bull ex Hiern (FWTA 2: 154) - ligustrifolia Stapf (Aubrév. 1959) = Agrocoffeopsis afzelii - maclaudii A.Cheval. (Aubrév. 1959) = C. canephora - macrochlamys K.Schum. (FWTA 2: 156) = Calycosiphonia m. - rupestris Hiern (FWTA 2: 156) = Argocoffeopsis r. - spathicalyx K.Schum. (FWTA 2: 156) = Calycosiphonia s. ✩ - stenophylla G.Don (FWTA 2: 156)

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✩

- togoensis A.Cheval. (FWTA 2: 156) Corynanthe pachyceras K.Schum. (Bot.J.Linn.Soc. 120: 303) Craterispermum caudatum Hutch. (FWTA 2: 188) - cerinanthum Hiern (FWTA 2: 188) - gracile A.Cheval. (Aubrév. 1959, 3: 310) = C. caudatum - laurinum (Poir.) Benth. (FWTA 2: 188) Cremaspora triflora (Thonn.) K.Schum. (FWTA 2: 147) Cuviera acutiflora DC. (FWTA 2: 177) - bolo Aubrév. & Pellegr. (Aubrév. 1959, 3: 300) = Robynsia glabrata - macroura K.Schum. (FWTA 2: 177) - nigrescens (Scott-Elliot ex Oliver) Wernh. (FWTA 2: 177) - sp.A. (FWTA 2: 177) - subuliflora Benth. (FWTA 2: 177) = C. macroura Dictyandra arborescens Welw. ex Hook.f. (FWTA 2: 132) - involucrata (Hook.f.) Hiern (FWTA 2: 132) Didymosalpinx abbeokutae (Hiern) Keay (FWTA 2: 130) Euclinia longiflora Salisb. (FWTA 2: 121) Gaertnera aurea Malcomber spec. nov. (to be published) - cooperi Hutch. & M.B.Moss (FWTA 2: 191) - liberiensis Petit (FWTA 2: 191) - longevaginalis (Hiern) Petit var. bracteata (Petit) Malcomber comb. nov. (to be published) - longevaginalis (Hiern) Petit (FWTA 2: 191) - paniculata Benth. (FWTA 2: 191) ✩ Gardenia imperialis K.Schum. (FWTA 2: 122) ✪ - nitida Hook. (FWTA 2: 123) - vogelii Hook.f. ex Planch. (FWTA 2: 123) Geophila afzelii Hiern (FWTA 2: 206) - hirsuta Benth. (FWTA 2: 205) = Hymenocoleus hirsutus - liberica A.Cheval. ex Hutch. & Dalziel (FWTA 2: 205) = Hymenocoleus libericus - neurodictyon (K.Schum.) Hepper (FWTA 2: 206) = Hymenocoleus n. var. n. - neurodictyon (K.Schum.) Hepper var. rhombicifolius Robbr. (Robbr. 1975) = Hymenocoleus n. var. rh. - obvallata (Schum.) F.Didr. (FWTA 2: 206) - repens (Linné) I.M.Johnston (FWTA 2: 205) - rotundifolia A.Cheval. ex Hepper (FWTA 2: 206) = Hymenocoleus rotudifolius Hallea ciliata (Aubrév. & Pellegr.) Leroy (Adansonia sér. 2, 15: 66) = H. ledermannii - ledermannii (K.Krause) Verdc. (Kew Bull. 40: 508) - stipulosa (DC.) Leroy (Adansonia sér. 2, 15: 66) Heinsia crinita (Afzel.) G.Taylor (FWTA 2: 161) - pulchella K.Schum. (Aubrév. 1959) = H. crinita ✪ Hutchinsonia barbata Robyns (FWTA 2: 175) ✪ - glabrescens Robyns (FWTA 2: 175) ✪ Hymenocoleus axillaris Robbr. (BJBB 47: 17) - hirsutus (Benth.) Robbr. (BJBB 45: 288) ✩ - libericus (A.Cheval. ex Hutch. & Dalziel) Robbr. (BJBB 45: 291) ✩ - multinervis Robbr. (BJBB 45: 294) ✩ - nervopilosus Robbr. (BJBB 47: 10) - neurodictyon (K.Schum.) Robbr. var. neurodictyon (BJBB 45: 291) - neurodictyon (K.Schum.) Robbr. var. rhombicifolius Robbr. (BJBB 45: 292) - neurodictyon (K.Schum.) Robbr. var. orientalis (Verdc.) Robbr. (BJBB 45: 300) - rotundifolius (A.Cheval. ex Hepper) Robbr. (BJBB 45: 287) Hymenodictyon floribundum (Steud. & Hochst.) B.L.Robinson (FWTA 2: 111) - gobiense Aubrév. & Pellegr. (Aubrév. 1959, 3: 294) = H. pachyantha - pachyantha K.Krause (FWTA 2: 111) ✩ Ixora aggregata Hutch. (Opera Bot. 9: 76) ✩ - baldwinii Keay (Opera Bot. 9: 84) - brachypoda DC. (Opera Bot. 9: 90) - divaricata Hutch. & Dalziel (FWTA 2: 142) = I. laxiflora var. laxiflora ✩ - guineensis Benth. (Opera Bot. 9: 104) ✪ - hiernii Scott-Elliot (Opera Bot. 9: 108) ✩ - laxiflora Sm. (Opera Bot. 9: 116) ✪ - laxiflora Sm. var. laxiflora (Opera Bot. 9: 116) ✪ - laxiflora Sm. var. linderi (Hutch.& Dalziel) De Block (Opera Bot. 9: 118) ✩ - liberiensis De Block (Opera Bot. 9: 122) - micheliana J.G.Adam (Bull.IFAN 35: 85) = I. aggregata ✩ - nigerica Keay ssp. occidentalis De Block (Opera Bot. 9: 142) ✩ - nimbana Schnell (Opera Bot. 9: 144) - sp.A (FWTA 2: 142) = I. liberiensis - sp.B (FWTA 2: 142) = I. tenuis ✪ - tenuis De Block (Opera Bot. 9: 160) ✪ Keetia bridsoniae Jongkind (Kew Bull. 57: 989) - cornelia (Cham. & Schlecht.) Bridson (Kew Bull. 41: 985) - hispida (Benth.) Bridson (Kew Bull. 41: 986) - leucantha (K.Krause) Bridson (Kew Bull. 41: 987) - mannii (Hiern) Bridson (Kew Bull. 41: 988)

- molundensis (K.Krause) Bridson (Kew Bull. 41: 976) - multiflora (Schum. & Thonn.) Bridson (Kew Bull. 41: 988) ✪ - obovata Jongkind (Kew Bull. 57: 991) ✩ - rubens (Hiern) Bridson (Kew Bull. 41: 989) ✩ - rufivillosa (Rob. ex Hutch.& Dalziel) Bridson (Kew Bull. 41: 990) - tenuifllora (Hiern) Bridson (Kew Bull. 41: 982) - venosa (Oliver) Bridson (Kew Bull. 41: 974) ✩ - venosissima (Hutch. & Dalziel) Bridson (Kew Bull. 41: 991) Lasianthus batangensis K.Schum. (BJBB 51: 451) ✩ - repens Hepper (BJBB 51: 451) Leptactina densiflora Hook.f. var. densiflora (FWTA 2: 132) - densiflora Hook.f. var. glabra Hutch. & Dalziel (FWTA 2: 133) ✩ - involucrata Hook.f. (Plant Syst.& E. 145: 105) ✩ - senegambica Hook.f. (FWTA 2: 133) Macrosphyra longistyla (DC.) Hiern (FWTA 2: 116) Massularia acuminata (G.Don) Bullock ex Hoyle (FWTA 2: 114) Mitragyna ciliata Aubrév. & Pellegr. (FWTA 2: 161) = Hallea ledermannii - stipulosa (DC.) O.Ktze. (FWTA 2: 161) = Hallea s. ✪ Monosalpinx guillaumetii N.Hallé (Adansonia sér. 2, 8: 367) Morelia senegalensis A.Rich. ex DC. (FWTA 2: 113) Morinda geminata DC. (FWTA 2: 189) - longiflora G.Don (FWTA 2: 189) - lucida Benth. (FWTA 2: 189) - morindoides (Baker) Milne-Redh. (FWTA 2: 189) ✩ Multidentia pobeguinii (Hutch. & Dalziel) Bridson (Kew Bull. 42: 646) ✪ Mussaenda afzelii G.Don (FWTA 2: 165) - arcuata Lam. ex Poir. (FWTA 2: 165) ✩ - chippii Wernh. (FWTA 2: 167) ✩ - conopharyngiifolia Stapf (FWTA 2: 165) - elegans Schum. & Thonn. (FWTA 2: 167) - erythrophylla Schum. & Thonn. (FWTA 2: 165) ✪ - grandiflora Benth. (FWTA 2: 167) ✩ - isertiana DC. (FWTA 2: 167) - landolphioides Wernh. (FWTA 2: 167) ✩ - linderi Hutch. & Dalziel (FWTA 2: 167) ✩ - nivea A.Cheval. ex Hutch. & Dalziel (FWTA 2: 165) - soyauxii Buttner (Fl.Gabon 12: 146) - tenuiflora Benth. (FWTA 2: 167) ✪ - tristigmatica Cummins (FWTA 2: 167) Nauclea diderrichii (De Wild. & Th.Dur.) Merrill (FWTA 2: 164) - pobeguinii (Pobéguin ex Pellegr.) Petit (FWTA 2: 163) = Sarcocephalus p. - trillesii (Pierre) Merrill (Aubr v. 1959) = N. diderrichii - vanderguchtii (De Wild.) Petit (1959 Fl.Gabon 12: 48) doubtful - xanthoxylon (A.Cheval.) Aubrév. (Boissiera 58: 86) Neorosea spp. (Fl.Gabon 17: 268) = Sericanthe spp. Nichallea soyauxii (Hiern) Bridson (Kew Bull. 33: 288) ✩ Oldenlandia chevalieri Bremek. (FWTA 2: 211) ✩ Otomeria cameronica (Bremek.) Hepper (FWTA 2: 215) Oxyanthus formosus Hook.f. ex Planch. (FWTA 2: 129) ✩ - pallidus Hiern (FWTA 2: 129) - racemosus (Schum. & Thonn.) Keay (FWTA 2: 129) - speciosus DC. (FWTA 2: 129) - subpunctatus (Hiern) Keay (FWTA 2: 129) ✩ - tenuis Stapf (Opera Bot. 8: 101) - unilocularis Hiern (FWTA 2: 129) Parapentas setigera (Hiern) Verdc. (BJBB 51: 187) Pauridiantha afzelii (Hiern) Bremek. (FWTA 2: 169) - hirtella (Benth.) Bremek. (FWTA 2: 169) ✩ - schnellii N.Hallé (Adansonia sér. 2, 5: 97) - stipulosa (Hutch. & Dalziel) Hepper (FWTA 2: 169) = Poecilocalyx s. - sylvicola (Hutch. & Dalziel) Bremek. (FWTA 2: 169) - ziamaeana (Jacq.-Félix) Hepper (FWTA 2: 168) = Stelechantha z. ✩ Pausinystalia lane-poolei (Hutch.) Hutch. ex Lane-Poole ssp. lane-poolei (Bot.J.Linn.Soc. 120: 310) ✪ Pavetta akeassii J.B.Hall (BJBB 50: 255) - corymbosa (DC.) F.N.Williams var. neglecta Bremek. (FWTA 2: 138) - corymbosa (DC.) F.N.Williams var. corymbosa (FWTA 2: 138) ✩ - genipifolia Schum. (FWTA 2: 139) ✩ - ixorifolia Bremek. (FWTA 2: 139) - lasioclada (K.Krause) Mildbr. ex Bremek. (FWTA 2: 138) ✩ - leonensis Keay (FWTA 2: 139) ✪ - micheliana J.G.Adam (Bull.IFAN 35: 87) ✩ - mollis Afzel. ex Hiern (FWTA 2: 140) ✩ - mollissima Hutch. & Dalziel (FWTA 2: 139) - nitida (Schum. & Thonn.) Hutch. & Dalziel (Aubrév. 1959, 3: 219) = P. corymbosa corymbosa

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- obanica Bremek. (FWTA 2: 139) - owariensis P.Beauv. (FWTA 2: 139) ✩ - platycalyx Bremek. (FWTA 2: 139) - spec. nov. (aff. ixorifolia) Hawthorne (to be published) - spec. nov. (aff. mannioides) Hawthorne (to be published) ✩ - subglabra Schum. (FWTA 2: 140) ✪ Pleiocoryne fernandensis (Hiern) Rauschert (Taxon 31: 561) ✩ Poecilocalyx stipulosa (Hutch. & Dalziel) N.Hallé (Adansonia sér. 2, 4: 238) Polycoryne fernandensis (Hiern) Keay (FWTA 2: 119) = Pleiocoryne f. Polysphaeria arbuscula K.Schum. (Kew Bull. 35: 116) ✩ - macrophylla K.Schum. (Kew Bull. 35: 115) Pouchetia africana A.Rich. ex DC. (FWTA 2: 132) - grandifolia Hawthorne spec. nov. (to be published) ✩ - parviflora Benth. (FWTA 2: 132) Psilanthus ebracteolatus Hiern (Kew Bull. 42: 455) - mannii Hook.f. (FWTA 2: 157) ✩ Psychotria abouabouensis (Schnell) Verdc. (Kew Bull. 30: 260) - abrupta Hiern (FWTA 2: 201) = Chazaliella a. - adafoana K.Schum. (FWTA 2: 201) = P. ivorensis ✩ - albicaulis Scott-Elliot (BJBB 34: 68) ✩ - ankasensis J.B.Hall (BJBB 50: 258) ✩ - biaurita (Hutch. & Dalziel) Verdc. (Kew Bull. 30: 260) ✩ - bidentata (Schult.) Hiern (BJBB 34: 132) ✩ - brachyantha Hiern (BJBB 36: 169) ✩ - brachyanthoides De Wild. (BJBB 36: 184) ✩ - brassii Hiern (BJBB 34: 58) - brenanii Hepper (FWTA 2: 198) = P. fimbriatifolia? - calva Hiern (BJBB 36: 100) - djumaensis De Wild. (FWTA 2: 202) ✩ - dorotheae Wernh. (BJBB 34: 67) ✩ - ebensis K.Schum. (Hall GC43359 (GC)) ✩ - elongato-sepala (Hiern) Petit (BJBB 34: 109) - fernandopoensis Petit (FWTA 2: 202) - gabonica Hiern (BJBB 34: 65) ✩ - guineensis Petit (BJBB 34: 78) - huae De Wild. (FWTA 2: 200) = P. calva ✩ - humilis Hiern var. major Petit (Jongkind et al. 3184 (WAG)) ✩ - ivorensis De Wild. (BJBB 36: 108) - juglasiana Aké Assi (BJBB 29: 359) = P. kitsonii ✩ - kitsonii Hutch. & Dalziel (BJBB 34: 153) ✩ - liberica Hepper (BJBB 34: 179) ✩ - limba Scott-Elliot (BJBB 34: 64) ✩ - linderi Hepper (BJBB 36: 185) ✩ - longituba A.Cheval. ex De Wild. (BJBB 34: 209) - lophoclada Hiern (FWTA 2: 200) = Chazaliella l. - malchairei De Wild. (FWTA 2: 201) = Psychotria spec.? - maliensis Schnell (FWTA 2: 199) = P. multinervis ✩ - mangenotii (Aké Assi) Verdc. (Kew Bull. 30: 259) ✩ - micheliana J.G.Adam (Bull.IFAN 35: 89) ✩ - multinervis De Wild. (BJBB 34: 139) - nigrescens De Wild. (FWTA 2: 201) = P. albicaulis - obscura Benth. (FWTA 2: 199) = P. schweinfurthii ✩ - ombrophila (Schnell) Verdc. (Kew Bull. 30: 259) - peduncularis (Salisb.) Verdc. (Kew Bull. 30: 257) - psychotriodes (DC.) Roberty (BJBB 34: 182) ✩ - reptans Benth. (BJBB 34: 151) ✩ - rufipilis De Wild. (BJBB 34: 154) ✩ - schnellii (Aké Assi) Verdc. (Kew Bull. 30: 259) - schweinfurthii Hiern (BJBB 34: 146) - sciadephora Hiern (FWTA 2: 201) = Chazaliella s. - sp. nr. abrupta Hiern (FWTA 2: 201) = Chazaliella cupulicalyx ✩ - spathacea (Hiern) Verdc. (Kew Bull. 30: 260) ✩ - spec. aff. calceata Petit (Hall & Swaine 1981: 267) ✩ - strictistipula Schnell (FWTA 2: 199) = P. reptans? BJBB 34: 151 ✩ - subglabra De Wild. (BJBB 34: 141) - subobliqua Hiern (BJBB 34: 168) - vogeliana Benth. (BJBB 34: 135) ✩ - yapoensis (Schnell) Verdc. (Kew Bull. 30: 259) ✩ Psydrax arnoldiana (De Wild. & Th.Dur.) Bridson (Kew Bull. 40: 699) - horizontalis (Schum. & Thonn.) Bridson (Kew Bull. 40: 722) - kraussioides (Hiern) Bridson (Kew Bull. 40: 723) ✩ - manensis (Aubrév. & Pellegr.) Bridson (Kew Bull. 40: 699) - parviflora (Afzel.) Bridson (Kew Bull. 40: 700) - schimperiana (A.Rich.) Bridson ssp. occidentalis (DC.) Bridson (Kew Bull. 40: 715) - subcordata (DC.) Bridson (Kew Bull. 40: 698) ✩ Pyrostria affinis (Robyns) Bridson (Kew Bull. 42: 628)

470

✩ Robynsia glabrata Hutch. (FWTA 2: 176) Rothmannia hispida (K.Schum.) Fagerlind (BJBB 65: 234) - longiflora Salisb. (FWTA 2: 125) - megalostigma (Wernh.) Keay (FWTA 2: 125) = R. munsae - munsae (Schweinf. ex Hiern) Petit ssp. megalostigma (Wernh.) Somers (BJBB 61: 301) - urcelliformis (Hiern) Bullock ex Robyns (BJBB 65: 226) - whitfieldii (Lindl.) Dandy (BJBB 65: 240) Rutidea dupuisii De Wild. ssp. occidentalis Bridson (Kew Bull. 33: 257) - membranacea Hiern (Kew Bull. 33: 273) - olentricha Hiern (Kew Bull. 33: 268) - parviflora DC. (Kew Bull. 33: 248) - smithii Hiern (Kew Bull. 33: 270) - syringoides (Webb) Bremek. (FWTA 2: 146) = Rutidea spp. Rytigynia affinis (Robyns) Hepper (FWTA 2: 186) = Pyrostria a. - canthioides (Benth.) Robyns (FWTA 2: 186) ✩ - leonensis (K.Schum.) Robyns (FWTA 2: 186) ✩ - liberica Robyns (FWTA 2: 186) - membranacea (Hiern) Robyns (FWTA 2: 186) - nigerica (S.Moore) Robyns (FWTA 2: 186) - umbellulata (Hiern) Robyns (Kew Bull. 42: 152) Sabicea aff. pilosa Hiern (FWTA 2:172) = S. multibracteata - africana (P.Beauv.) Hepper (FWTA 2:173) ✪ - arachnoidea Hutch. & Dalziel (FWTA 2:174) ✪ - bracteolata Wernh. (FWTA 2:172) - brevipes Wernh. (FWTA 2:173) - calycina Benth. (FWTA 2:172) - cordata Hutch. & Dalziel (FWTA 2:172) ✪ - discolor Stapf (FWTA 2:173) ✩ - ferruginea (G.Don) Benth. - geophiloides Wernh. (HBJBB 50: 252) ✪ - harleyae Hepper (FWTA 2:172) ✩ - liberica Hepper (FWTA 2:172) ✩ - multibracteata J.B.Hall (BJJB 50: 249) ✩ - rosea A.C.Hoyle (FWTA 2: 172) ✩ - solitaria J.B.Hall (BJBB 50: 253) - speciosa K.Schum. (FWTA 2:172) - venosa Benth. (FWTA 2:172) ✩ - vogelii Benth. (FWTA 2:173) Sacosperma paniculatum (Benth.) G.Taylor (FWTA 2: 213) ✩ - parviflorum (Benth.) G.Taylor (FWTA 2: 213) Sarcocephalus pobeguinii Hua ex Pobeguin; Pellegr. (Hawthorne 1995a: 268) ✪ Schizocolea linderi (Hutch. & Dalziel) Bremek. (FWTA 2: 190) ✪ Schumanniophyton problematicum (A.Cheval.) Aubrév. (FWTA 2: 116, 496) ✪ Sericanthe adamii (N.Hallé) Robbr. (BJBB 51: 172) - chevalieri (K.Krause) Robbr. (BJBB 48: 39) - chevalieri (K.Krause) Robbr. var. coffeoides (A. Cheval.) Robbr. (BJBB 48: 41) ✩ - spec.nov. (Voorhoeve 1286 (WAG)) ✩ - toupetou (Aubrév. & Pellegr.) Robbr. (BJBB 48: 65) - trilocularis (Scott-Elliot) Robbr. ssp. trilocularis (BJBB 48: 67) ✩ - trilocularis (Scott-Elliot) Robbr. ssp. paroissei (Aubrév. & Pell.) Robbr. (BJBB 48: 69) Sherbournia bignoniiflora (Welw.) Hua (FWTA 2: 127) ✪ - calycina (G.Don) Hua (FWTA 2: 127) ✩ Spermacoce intricans (Hepper) Burkill f. (Kew Bull. 41: 1006) ✩ Stelechantha ziamaeana (Jacq.-Félix) N.Hallé (Adansonia sér. 2, 4: 236) Tarenna bipindensis (K.Schum.) Bremek. (FWTA 2: 135) ✩ - brachysiphon (Hiern) Keay (FWTA 2: 135) - eketensis Wernh. (FWTA 2: 135) - fusco-flava (K.Schum.) S.Moore (Adansonia sér. 2, 7: 506) ✩ - gracilis (Stapf) Keay (FWTA 2: 135) ✩ - hutchinsonii Bremek. (FWTA 2: 135) ✩ - aff. laurentii (De Wild.) Garcia (Vigne 3505 (FHO)) ✩ - nitidula (Benth.) Hiern (FWTA 2: 135) - spec. aff. pavettoides (Harvey) Sim (Morton 266 (K)) - soyauxii (Hiern) Bremek. (FWTA 2: 135) = Nichallea s. - sp.nov. Bridson (Yallah 133 (K)) - thomasii Hutch. & Dalziel (FWTA 2: 135) ✪ - vignei Hutch. & Dalziel var. subglabra Keay (FWTA 2: 134) ✪ - vignei Hutch. & Dalziel var. vignei (FWTA 2: 135) ✩ Tricalysia anomala E.A.Bruce var. guineensis Robbr. (BJBB 53: 314) ✩ - biafrana Hiern (BJBB 57: 129) ✩ - bracteata Hiern (BJBB 49: 297) - chevalieri K.Krause (FWTA 2: 152) = Sericanthe c. - coffeoides Hutch. & Dalziel (Aubrév. 1959) = Sericanthe chevalieri - coriacea (Benth.) Hiern (BJBB 57: 79) ✩ - deightonii Brenan (BJBB 49: 348) - discolor Brenan (BJBB 49: 293)

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- elliotii (K.Schum.) Hutch. & Dalziel (BJBB 53: 308) - faranahensis Aubrév. & Pellegr. (BJBB 57: 133) - macrophylla K.Schum. (BJBB 49: 339) - okelensis Hiern var. okelensis (BJBB 57: 93) - pallens Hiern (BJBB 57: 114) - paroissei Aubrév. & Pellegr. (FWTA 2: 152) = Sericanthe trilocularis ssp. p. ✩ - parva Keay (BJBB 57: 150) ✩ - reflexa Hutch. var. ivorensis Robbr. (BJBB 49: 306) - reflexa Hutch. var. reflexa (BJBB 49: 302) ✩ - reticulata (Benth.) Hiern (BJBB 57: 90) - toupetou Aubrév. & Pellegr. (FWTA 2: 152) = Sericanthe t. - trilocularis (Scott-Elliot) Hutch. & Dalziel (FWTA 2: 152) = Sericanthe t. - vignei Aubrév. & Pellegr. (FWTA 2: 151) = Tricalysia coriacea Trichostachys aurea Hiern (FWTA 2: 207) Uncaria africana G.Don var. africana (FWTA 2: 162) - africana G.Don var. angolensis Havil. (FWTA 2: 162) - talbotii Wernh. (FWTA 2: 162) Urophyllum afzelii Hiern (Aubrév. 1959) = Pauridiantha a. - hirtellum Benth. (Aubrév. 1959) = Pauridiantha hirtella Vangueria sp.A. (FWTA 2: 187) Vangueriella campylacantha (Mildbr.) Verdc. (Kew Bull. 42: 196) ✩ - discolor (Benth.) Verdc. (Kew Bull. 42: 193) - nigerica (Robyns) Verdc. (Kew Bull. 42: 193) ✩ - orthacantha (Mildbr.) Bridson & Verdc. (Kew Bull. 42: 198) - spinosa (Schum. & Thonn.) Verdc. (Kew Bull. 42: 194) ✩ - vanguerioides (Hiern) Verdc. (Kew Bull. 42: 193) Vangueriopsis discolor (Benth.) Robyns (FWTA 2: 179) = Vangueriella d. - nigerica Robyns (FWTA 2: 179) = Vangueriella n. - spinosa (Schum. & Thonn.) Hepper (FWTA 2: 179) = Vangueriella s. - subulata Robyns (FWTA 2: 179) = Vangueriella campylacantha ● - vanguerioides (Hiern) Robyns (FWTA 2: 179) = Vangueriella v. Virectaria procumbens (Sm.) Bremek. (FWTA 2: 208) ✩ - tenella J.B.Hall (Kew Bull. 26: 569)

✩ ✩

Rutaceae Aeglopsis ?sp. (FWTA 1: 687) ✩ - chevalieri Swingle (FWTA 1: 687) ✩ Afraegle paniculata (Schum. & Thonn.) Engl. (FWTA 1: 687) Araliopsis soyauxii Engl. (Hall & Swaine 1981: 348) = Vepris s. - tabouensis Aubrév. & Pellegr. (FWTA 1: 688) = Vepris t. ✩ Citropsis articulata (W. ex Spr.) Swingle & Kellerman (FWTA 1: 688) - mirabilis (A.Cheval.) Swingle & Kellerman (Aubrév. 1959) = Citropsis articulata ✩ - sp.nr.gabunensis (Engl.) Swingle & Kellerman (FWTA 1: 688) Clausena anisata (Willd.) Hook.f. ex Benth. (FWTA 1: 686) Diphasia angolensis (Hiern) Verdoorn (Aubrév. 1959) = Vepris hiernii - klaineana Pierre (FWTA 1: 689) = Vepris angolensis Fagara atchoum Aké Assi (BJBB 30: 398) = Zanthoxylum a. - lemairei De Wild. (FWTA 1: 686) = Zanthoxylum l. - leprieurii (Guillaumet & Perr.) Engl. (FWTA 1: 686) = Zanthoxylum l. - macrophylla Engl. (FWTA 1: 685) = Zanthoxylum gilletii ✩ - mezoneurispinosa Aké Assi (BJBB 30: 400) - parvifoliola A.Cheval. ex Keay (FWTA 1: 685) = Zanthoxylum lemairei - psammophila Aké Assi (BJBB 30: 402) = Zanthoxylum psammophilum - pubescens A.Cheval. (FWTA 1: 685) = Zanthoxylum chevalieri - rubescens (Planch. ex Hook.f.) Engl. (FWTA 1: 685) = Zanthoxylum r. - sp.A. (FWTA 1: 686) = Zanthozylum mezoneurispinosa - sp.B. (FWTA 1: 686) = Zanthoxylum atchoum - viridis A.Cheval. (FWTA 1: 685) = Zanthoxylum viride - zanthoxyloides Lam. (FWTA 1: 685) = Zanthoxylum z. Oricia suaveolens (Engl.) Verdoorn (FWTA 1: 688) = Vepris s. Teclea afzelii Engl. (FWTA 1: 689) = Vepris a. - verdoorniana Exell & Mendonça (FWTA 1: 689) = Vepris v. ✩ Vepris afzelii (Engl.) W.Mziray (Symb.Bot.Ups. 30: 70) - angolensis (Hiern) W.Mziray (Symb.Bot.Ups. 30: 70) = V. hiernii ✩ - felicis Breteler (Kew Bull. 50: 131) ✩ - heterophylla (Engl.) Letouzey (Symb.Bot.Ups. 30: 73) ✩ - hiernii Gereau (Novon 11: 43) - soyauxii (Engl.) W.Mziray (Symb.Bot.Ups. 30: 75) ✩ - suaveolens (Engl.) W.Mziray (Symb.Bot.Ups. 30: 76) ✪ - tabouensis (Aubrév. & Pellegr.) W.Mziray (Symb.Bot.Ups. 30: 76) - verdoorniana (Exell & Mendonca) W.Mziray (Symb.Bot.Ups. 30: 76) ✩ Zanthoxylum atchoum (Aké Assi) Waterman (Taxon 24: 363) ✩ - chevalieri Waterman (Taxon 24: 365) - gilletii (De Wild.) Waterman (Taxon 24: 363)

✩ ✪ ✩ ✩

- lemairei (De Wild.) Waterman (Taxon 24: 364) - leprieurii Guillaumet & Perr. (Taxon 24: 361) - psammophilum (Aké Assi) Waterman (Taxon 24: 364) - rubescens Planch. ex Hook.f. (Taxon 24: 361) - viride (A.Cheval.) Waterman (Taxon 24: 365) - zanthoxyloides (Lam.) Waterman (Willdenowia 11: 361)

Santalaceae ✪ Okoubaka aubrevillei Pellegr. & Normand var. aubrevillei (FWTA 1: 656)

Sapindaceae Allophylus africanus P.Beauv. f. africanus (FWTA 1: 713) - africanus P.Beauv. f. chrysothrix Radlk. (FWTA 1: 713) - camptoneurus Radlk. (Fl.Cam.: 46) - talbotii Baker f. (FWTA 1: 714) Aphania senegalensis (Juss. ex Poir.) Radlk. (FWTA 1: 716) = Lepisanthes s. - silvatica A.Cheval. ex Hutch. & Dalziel (Aubrév. 1959) = Lepisanthes senegalensis Aporrhiza talbotii Baker (Fl.Cam.: 154) - urophylla Gilg (FWTA 1: 721) Blighia sapida Konig (FWTA 1: 722) - unijugata Baker (FWTA 1: 723) - welwitschii (Hiern) Radlk. var. bancoensis (Aubrév. & Pellegr.) Pellegr. (Aubrév. 1959) - welwitschii (Hiern) Radlk. (FWTA 1: 722) Cardiospermum grandiflorum Swartz (FWTA 1: 711) - halicacabum Linné (FWTA 1: 711) Chytranthus angustifolius Exell (Fl.Cam.: 100) - atroviolaceus Baker f. ex Hutch. & Dalziel (FWTA 1: 717) - bracteosus Radlk. (Adansonia sér. 2, 2: 293) = C. angustofolius - carneus Radlk. (Fl.Cam.: 94) ✪ - cauliflorus (Hutch. & Dalziel) Wickens (Kew Bull. 24: 345) - longiracemosus Gilg ex Radlk. (Adansonia sér. 2, 2: 299) = C. carneus - macrobotrys (Gilg) Exell & Mendonça (Fl.Cam.: 111) - mangenotii N.Hallé & A.Assi (Adansonia sér. 2, 2: 295) = C. cauliflorus - setosus Radlk. (Adansonia sér. 2, 2: 297) - talbotii (Baker) Keay (Fl.Cam.: 108) ✩ - verecundus N.Hallé & Aké Assi (Adansonia sér. 2, 2: 297) - villiger Radlk. (FWTA 1: 717) = C. carneus Crossonephelis adamii Fouilloy (Adansonia sér. 2, 12: 551) = Glenniea a. ✩ Deinbollia calophylla Gilg ex Radlk. (FWTA 1: 715) ✩ - cuneifolia Baker (FWTA 1: 715) - grandifolia Hook.f. (FWTA 1: 715) ✩ - molliuscula Radlk. (Hall & Swaine 1981: 349) - pinnata Schum. & Thonn. (FWTA 1: 715) ✪ - voltensis Hutch. ex Burtt Davy & Hoyle (FWTA 1: 715) Eriocoelum kerstingii Gilg ex Engl. (FWTA 1: 724) ✩ - pungens Radlk. ex Engl. var. pungens (FWTA 1: 724) - racemosum Baker (FWTA 1: 724) ✩ Glenniea adamii (Fouilloy) Leenh. (Blumea 22: 412) Laccodiscus cauliflorus Hutch. & Dalziel (FWTA 1: 721) = Chytranthus c. Lecaniodiscus cupanioides Planch. ex Benth. (FWTA 1: 720) ✩ - punctatus J.B.Hall (BJBB 50: 262) Lepisanthes senegalensis (Poir.) Leenh. (Blumea 17: 85) ✩ Lychnodiscus dananensis Aubrév. & Pellegr. (Fl.Cam.: 170) - reticulatus Radlk. (FWTA 1: 722) Majidea fosteri (Sprague) Radlk. (FWTA 1: 725) Pancovia bijuga Willd. (FWTA 1: 718) - pedicellaris Radlk. & Gilg (Fl.Cam.: 118) - sessiliflora Hutch. & Dalziel (Hall & Swaine 1981: 255) ✩ - subcuneata Radlk. (FWTA 1: 718) - turbinata Radlk. (FWTA 1: 718) = P. pedicellaris Paullinia pinnata Linné (FWTA 1: 710) ✩ Placodiscus attenuatus J.B.Hall (Adansonia sér. 2, 20: 290) ✩ - bancoensis Aubrév. & Pellegr. (FWTA 1: 720) ✩ - boya Aubrév. & Pellegr. (FWTA 1: 720) ✩ - bracteosus J.B.Hall (Adansonia sér. 2, 20: 289) - cuneatus Radlk. (Fl.Cam.: 132) = P. bracteosus - leptostachys Radlk. (Fl.Cam.: 130) = P. bracteosus - leptostachys Radlk. (FWTA 1: 720) = P. oblongifolius ✩ - oblongifolius J.B.Hall (Adansonia sér. 2, 20: 291) ✩ - pseudostipularis Radlk. (FWTA 1: 720) ✩ - riparius Keay (FWTA 1: 720) - sp.A. (FWTA 1: 720) = P. bracteosus ✩ - splendidus Keay (FWTA 1: 720) Zanha golungensis Hiern (FWTA 1: 725)

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Sapotaceae Afrosersalisia afzelii (Engl.) A.Cheval. (FWTA 2: 30) = Synsepalum a. - cerasifera (Welw.) Aubrév. (FWTA 2: 30) = Synsepalum c. - chevalieri (Engl.) Aubrév. (Aubrév. 1959) = Synsepalum cerasiferum Aningeria altissima (A.Cheval.) Aubrév. & Pellegr. (FWTA 2: 24) = Pouteria a. - robusta (A.Cheval.) Aubrév. & Pellegr. (FWTA 2: 24) = Pouteria aningeri ✩ Aubregrinia taiensis (Aubrév. & Pellegr.) Heine (FWTA 2: 24) Bequaertiodendron magalismontanum (Sonder) Heine & Hemsl. (Kew Bull. 14: 307) - oblanceolatum (S.Moore) Heine & Hemsl. (FWTA 2: 25) = Englerophytum o. ✩ Breviea leptosperma (Baehni) Heine (FWTA 2: 23) = B. sericea - sericea (A.Cheval.) Aubrév. & Pellegr. (Aubréville 1959, 3: 108) Chrysophyllum africanum A.DC. (Bull.MNHN Paris sér. 4, 11: 459.) - albidum G.Don (FWTA 2: 27) ✩ - azaguieanum Miège (FWTA 2: 27) - beguei Aubrév. & Pellegr. (FWTA 2: 27) - delevoyi De Wild. (FWTA 2: 28) = C. africanum - giganteum A.Cheval. (FWTA 2: 28) - letestuanum A.Cheval. (Aubrév. 1959) = C. ubanguiense - pentagonocarpum Engl. & K.Krause (FWTA 2: 26) = C. ubanguiense - perpulchrum Mildbr. ex Hutch. & Dalziel (FWTA 2: 28) - pruniforme Pierre ex Engl. (FWTA 2: 26) - subnudum Baker (FWTA 2: 27) ✩ - taiense Aubrév. & Pellegr. (FWTA 2: 27) - ubanguiense (De Wild.) Harris (Kew Bull. 55: 229) - welwitschii Engl. (FWTA 2: 27) ✩ Delpydora gracilis A.Cheval. (FWTA 2: 25) Dumoria heckelii A.Cheval. (Aubrév. 1959) = Tieghemella h. Endotricha taiensis Aubrév. & Pellgr. (Aubrév. 1959) = Aubreginia t. Englerophytum oblanceolatum (S.Moore) Penn. (Pennington 1991: 252) - oubanguiense (Aubrév. & Pellegr.) Aubrév. & Pellegr. (BJBB 59: 159) - aff. oubanguiense (BJBB 59: 160) Gambeya subnuda (Baker) Pierre (Aubreville) = Chrysophyllum s. ✩ Gluema ivorensis Aubrév. & Pellegr. (FWTA 2: 19) ✩ Inhambanella guereensis (Aubrév. & Pellegr.) Penn. (Pennington 1991: 142) ✩ Ituridendron bequaertii De Wild. (FWTA 2: 18) not part of Omphalocarpus Kantou guereensis Aubrév. & Pellegr. (FWTA 2: 23) = Inhambanella g. Lemonniera batesii H.Lecomte (Aubrév. 1959, 3: 116) = Neolemonniera b. - clitandrifolia H.Lecomte (Aubrév. 1959, 3: 96) = Neolemonniera c. Malacantha alnifolia (Baker) Pierre (FWTA 2: 24) = Pouteria a. - heudelotiana Pierre (Aubrév. 1959) = Pouteria alnifolia Manilkara lacera (Baker) Dubard (Aubréville 1959, 3: 120) = M. obovata - multinervis (Baker) Dubard (FTEA: 69) = M. obovata - obovata (Sabine & G.Don) Hemsl. (FWTA 2: 20) - sylvestris Aubrév. & Pellegr. (Aubrév. 1959, 3: 122) = M. obovata - welwitschii (Engl.) Dubard (Aubrév. 1959) = M. obovata Mimusops andongensis Hiern (FWTA 2: 20) - kummel Bruce ex A. DC. (FWTA 2: 20) - warneckei Engl. (Aubrév. 1959) = M. andongensis Neoboivinella glomeruliflora (Hutch. & Dalziel) Aubrév. & Pel (Aubrév. 1959) = Englerophytum oblanceolatum Neolemonniera batesii (Engl.) Heine (FWTA 2: 19) ✩ - clitandrifolia (A.Cheval.) Heine (FWTA 2: 19) ✩ Omphalocarpum ahia A.Cheval. (FWTA 2: 18) - anocentrum Pierre (Aubrév. 1959, 3: 110) = O. elatum - elatum Miers (FWTA 2: 18) - pachysteloides Mildbr. ex Hutch. & Dalziel (Pennington 1991: 261) = Ituridendron bequaertii - procerum P.Beauv. (FWTA 2: 18) Pachystela brevipes (Baker) Baill. ex Engl. (FWTA 2: 28) = Synsepalum b. - msolo (Engl.) Engl. (FWTA 2: 28) = Synsepalum m. - pobeguiniana Pierre ex Lecomte (FWTA 2: 29) = Synsepalum pobeguinianum Pouteria alnifolia (Baker) Roberty (Pennington 1991: 203) - altissima (A.Cheval.) Baehni (Pennington 1991: 203) - aningeri Baehni (Gautier 1997: 35, Pennington 1991) Sideroxylon aubrevillei Pellegr. (Aubrév. 1959) = Synsepalum a. Synsepalum afzelii (Engl.) Penn. (Pennington 1991: 248) ✩ - aubrevillei (Pellegr.) Aubrév. & Pellegr. (FWTA 2: 496) - brevipes (Baker) Penn. (Pennington 1991: 248) - cerasiferum (Welw.) Penn. (Pennington 1991: 248) - dulcificum (Schum. & Thonn.) Daniell (FWTA 2: 22) - msolo (Engl.) Penn. (Pennington 1991: 249) - passargei (Engl.) Penn. (Pennington 1991: 249) - pobeguinianum (Lecomte) Aké Assi & Gautier (Candollea 55: 282) - revolutum (Baker) Penn. (Pennington 1991: 249) ✩ - tsounkpe Aubrév. & Pellegr. (FWTA 2: 22)

472

Tieghemella heckelii Pierre ex A.Cheval. (FWTA 2: 21) Vincentella camerounensis Pierre (Aubrév. 1959) = Synsepalum revolutum - passargei (Engl.) Aubrév. (FWTA 2: 23) = Synsepalum p. - revoluta (Baker) Pierre (FWTA 2: 23) = Synsepalum revolutum

Scytopetalaceae ✪ Rhaptopetalum beguei Mangenot (361 Adansonia sér. 2, 17: 130) Rhaptopetalum sp. (FWTA 1: 300) = R. beguei ✪ Scytopetalum tieghemii (A.Cheval.) Hutch. & Dalziel (FWTA 1: 300)

Simaroubaceae Brucea antidysenterica Lam. (FWTA 1: 692) - guineensis G.Don (FWTA 1: 692) ✪ Gymnostemon zaizou Aubrév. & Pellegr. (FWTA 1: 692) Hannoa klaineana Pierre & Engl. (FWTA 1: 691) Harrisonia abyssinica Oliver (FWTA 1: 690) Mannia simarubopsis Pellegr. (Aubrév. 1959) = Pierreodendron kerstingii ✩ Nothospondias staudtii Engl. (Hall & Swaine 1981: 349) ✪ Pierreodendron kerstingii (Engl.) Little (FWTA 1: 690)

Solanaceae Solanum anomalum Thonn. (FWTA 2: 334) - erianthum D.Don (Taxon 17: 176) - terminale Forssk. ssp. inconstans (C.H.Wright) Heine (FWTA 2: 331) - terminale Forssk. ssp. welwitschii (C.H.Wright) Heine (FWTA 2: 331) - torvum Sw. (FWTA 2: 333) - verbascifolium Linné (FWTA 2: 332) = S. erianthum

Sterculiaceae Byttneria catalpifolia Jacq. ssp. africana (Mast.) Exell & Mendonça (FWTA 1: 314) ✪ - guineensis Keay & Milne-Redh. (FWTA 1: 314) ✪ - ivorensis N.Hallé (Adansonia sér. 2, 2: 287) ✩ Cola angustifolia K.Schum. (FWTA 1: 329) ✪ - attiensis Aubrév. & Pellegr. (FWTA 1: 329) ✪ - boxiana Brenan & Keay (FWTA 1: 329) ✩ - buntingii Baker f. (FWTA 1: 326) ✪ - caricifolia (G.Don) K.Schum. (FWTA 1: 328) - chlamydantha K.Schum. (FWTA 1: 328) - digitata Mast. (FWTA 1: 326) - flavo-velutina K.Schum. (FWTA 1: 329) - gigantea A.Cheval. var. glabrescens Brenan & Keay (FWTA 1: 330) - heterophylla (P.Beauv.) Schott. & Endl. (FWTA 1: 328) - lateritia K.Schum. var. maclaudi (A.Cheval.) Brenan & Keay (FWTA 1: 330) ✩ - laurifolia Mast. (FWTA 1: 328) ✩ - lorougnonis Aké Assi (Bull.IFAN 41: 483) - millenii K.Schum. (FWTA 1: 328) - nitida (Vent.) Schott. & Endl. (FWTA 1: 329) ✪ - reticulata A.Cheval. (FWTA 1: 329) ✩ - simiarum Sprague ex Brenan & Keay (FWTA 1: 330) ✩ - triloba (R.Br.) K.Schum. (FWTA 1: 330) ✪ - umbratilis Brenan & Keay (FWTA 1: 326) - verticillata (Thonn.) Stapf ex A.Cheval. (FWTA 1: 330) Dombeya buettneri K.Schum. (Opera Bot. 2: 75) Eribroma oblonga Pierre ex A.Cheval. (Fl.Gabon: 18) = Sterculia o. ✪ Heritiera utilis (Sprague) Spraque (Hawthorne 1995a: 170) Hildegardia barteri (Mast.) Kosterm. (FWTA 1: 332) ✩ Leptonychia occidentalis Keay (FWTA 1: 316) - pubescens Keay (FWTA 1: 316) Mansonia altissima (A.Cheval.) A.Cheval. var. altissima (FWTA 1: 313) Nesogordonia papaverifera (A.Cheval.) R.Capuron (FWTA 1: 313) Octolobus angustatus Hutch. (FWTA 1: 319) = O. spectabilis - spectabilis Welw. (Fl.Gabon: 107) Pterygota bequaertii De Wild. (FWTA 1: 320) - macrocarpa K.Schum. (FWTA 1: 320) ✪ Scaphopetalum amoenum A.Cheval. (FWTA 1: 315) Sterculia oblonga Mast. (FWTA 1: 321) - rhinopetala K.Schum. (FWTA 1: 321) - tragacantha Lindl. (FWTA 1: 321) Tarrietia utilis (Sprague) Sprague (FWTA 1: 332) = Heritiera u. Triplochiton scleroxylon K.Schum. (FWTA 1: 313)

Stilbaceae Nuxia congesta R.Br. ex Fresen. (Med.LHW 75-8: 12)

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Taccaceae Tacca leontopetaloides (Linné) O.Kuntze (FWTA 3: 176)

Thymelaeaceae Craterosiphon scandens Engl. & Gilg. (FWTA 1: 174) Dicranolepis disticha Planch. (FWTA 1: 173) - grandiflora Engl. (FWTA 1: 173) ✩ - laciniata Gilg (FWTA 1: 173) ✩ - persei Cummins (FWTA 1: 173) ✩ - pubescens H.H.W.Pearson (FWTA 1: 173) ✩ Octolepis decalepis Gilg (FWTA 1: 173) Peddiea fischeri Engl. (FWTA 1: 174)

Tiliaceae Christiana africana DC. (FWTA 1: 301) Desplatsia chrysochlamys (Mildbr. & Burr.) Mildbr. & Bu (FWTA 1: 307) - dewevrei (De Wild. & Th.Dur.) Burret (FWTA 1: 307) - subericarpa Bocq. (FWTA 1: 307) Duboscia viridiflora (K.Schum.) Mildbr. (FWTA 1: 305) Glyphaea brevis (Sprengel) Monachino (FWTA 1: 308) Grewia barombiensis K.Schum. (FWTA 1: 303) = G. hookerana - brunnea K.Schum. (FWTA 1: 303) = G. malacocarpa var. b. - carpinifolia Juss. (FWTA 1: 305) - hookerana Exell & Mendonça (FWTA 1: 303) = G. malacocarpa var. h. - malacocarpa Mast. (FWTA 1: 303) = G. malacocarpa var. m. - malacocarpa Mast. var. brunnea (K. Schum.) Jongkind to be published - malacocarpa Mast. var. hookerana (Exell & Mendonça) Jongkind to be published - malacocarpa Mast. var. malacocarpa (Mast.) Jongkind to be published - megalocarpa Juss. (FWTA 1: 305) - mollis Juss. (Adansonia sér. 2, 4: 99) = G. pubescens - pubescens P.Beauv. (Taxon 42: 696)

Triuridaceae ✩ Sciaphila africana A.Cheval. (FWTA 3: 15)

Ulmaceae Celtis adolfi-friderici Engl. (FWTA 1: 592) ✩ - africana Burm.f. (FWTA 1: 592) - brownii Rendle (FWTA 1: 592) = C. philippensis - durandii Engl. (FWTA 1: 592) = C. gomphophylla - gomphophylla Baker (Fl.Cam.: 39) - integrifolia Lam. (FWTA 1: 592) = C. toka - mildbraedii Engl. (FWTA 1: 592) - philippensis Blanco (Boissiera 58: 164) - toka (Forssk.) Hepper & Wood (Kew Bull. 38: 86.) - wightii Planch. (Kew Bull. 19: 141) = C. phillipensis - zenkeri Engl. (FWTA 1: 592) Chaetacme aristata Planch. (Fl.Cam.: 56) - madagascariensis Baker (Aubrév. 1959) = C. aristata Holoptelea grandis (Hutch.) Mildbr. (FWTA 1: 593) Trema guineensis (Schum. & Thonn.) Ficalho (FWTA 1: 592) = T. orientalis - orientalis (Linné) Blume (Kew Bull. 19: 143)

Urticaceae Boehmeria macrophylla Hornem. (Kew Bull. 37: 164) - platyphylla D.Don (FWTA 1: 622) = B. macrophylla Elatostema paivaeanum Wedd. (FWTA 1: 620) Fleurya aestuans (Linné) Miq. (FWTA 1: 619) = Laportea a. - ovalifolia (Schum. & Thonn.) Dandy (FWTA 1: 619) = Laportea o. ✩ Girardinia diversifolia (Link) I.Friis (Kew Bull. 36: 145) - heterophylla (Vahl) Decaisne (Fl.Cam.: 110) = G. diversifolia Laportea aestuans (Linné) Cheval. (Fl.Cam.: 121) - ovalifolia (Thonn. ex Schum.) Chew (Fl.Cam.: 131) ✩ Pilea sublucens Wedd. (Kew Bull. 44: 589) ✩ Pouzolzia denudata De Wild. & Durand (Kew Bull. 39: 589) - guineensis Benth. (Kew Bull. 39: 588) ✩ - parasitica Schweinf. (Kew Bull. 39: 593) ✩ Procris crenata Robinson (Fl.Cam.: 155) Urera cameroonensis Wedd. (FWTA 1: 618) = U. trinervis - cuneata Rendle (FWTA 1: 618) ✩ - keayi Letouzey (Adansonia sér. 2, 7: 299) ✩ - mannii (Wedd.) Benth. & Hook.f. ex Re (FWTA 1: 618) ✩ - oblongifolia Benth. (FWTA 1: 618)

✩ ✩ ✩ ✩

- obovata Benth. (FWTA 1: 618) - repens (Wedd.) Rendle (FWTA 1: 618) = U. keayi - rigida (Benth.) Keay (FWTA 1: 618) - robusta A.Cheval. (FWTA 1: 618) - trinervis (Hochst. & Krauss) Friis & Im. (Nordic J.Bot. 7: 126)

Verbenaceae Clerodendrum buchholzii Gürke (FWTA 2: 443) = C. silvanum var. buchholzii - capitatum (Willd.) Schum. & Thonn. var. capitatum (FWTA 2: 443) - capitatum (Willd.) Schum. & Thonn. var. cephalanthum (Oliver) Huber (FWTA 2: 443) - formicarum Gürke (FWTA 2: 444) - polycephalum Baker (FWTA 2: 444) ✩ - sassandrense Jongkind (Syst. Geogr. Pl. 72: 239) - silvanum Henriq. var. buchholzii (Gürke) Verdc. (FTEA: 110) - sinuatum Hook. (FWTA 2: 444) - splendens G.Don (FWTA 2: 444) - thyrsoideum Gürke (FWTA 2: 444) - umbellatum Poir. (FWTA 2: 442) - violaceum Gürke (FWTA 2: 441) - volubile P.Beauv. (FWTA 2: 444) Premna angolensis Gürke (FWTA 2: 438) ✪ - grandifolia Meeuse (FWTA 2: 438) - hispida Benth. (FWTA 2: 438) ✩ - lucens A.Cheval. (FWTA 2: 438) - quadrifolia Schum. & Thonn. (FWTA 2: 438) Vitex chrysocarpa Planch. ex Benth. (FWTA 2: 448) - ferruginea Schum. & Thonn. (FWTA 2: 447) - fosteri C.H.Wright (Aubrév. 1959) = V. ferruginea - grandifolia Gürke (FWTA 2: 446) - micrantha Gürke (FWTA 2: 446) ✩ - oxycuspis Baker (FWTA 2: 446) ✩ - phaeotricha Mildbr. ex Pieper (FWTA 2: 447) - rivularis Gürke (FWTA 2: 446) - rufa A.Cheval. (Aubrév. 1959) = V. phaeotricha - thyrsiflora Baker (FWTA 2: 446)

Violaceae (Woody genera are under revision by Dr. Achoundong) ✩ Allexis cauliflora (Oliver) Pierre (Hall & Swaine 1981: 349) ✩ Decorsella paradoxa A.Cheval. (FWTA 1: 758) Gymnorinorea abidjanensis (Aubrév. & Pellegr.) Keay (FWTA 1: 104) = Decorsella paradoxa ✩ Rinorea afzelii Engl. (FWTA 1: 103) - angustifolia (Thou.) Baill. (Kew Bull: 16: 412) - ardisiiflora (Welw. ex Oliver) O.Ktze. (FWTA 1: 101) = R. angustifolia - arenicola M.Brandt (FWTA 1: 103) = R. welwitschii ✪ - aylmeri Chipp (FWTA 1: 104) - brachypetala (Turcz.) O.Ktze. (FWTA 1: 104) ✩ - breviracemosa Chipp (FWTA 1: 103) ✩ - bussei M.Brandt (FWTA 1: 103) - convallariiflora M.Brandt (FWTA 1: 103) = R. c. subsp. occidentalis ✩ - convallariiflora M.Brandt ssp. occidentalis Grey-Wilson (Kew Bull. 36: 115) - coriaceae Berhaut (Fl.Senegal ed. 2: 428) = Scottelia leonensis - dentata (P.Beauv.) O.Kuntze (FWTA 1: 104) ✩ - djalonensis A.Cheval. ex Hutch. & Dalziel (FWTA 1: 103) - elliotii Engl. (FWTA 1: 103) = R. welwitschii - ilicifolia Kuntze (FWTA 1: 101) ✩ - johnstonii (Stapf) M.Brandt (FWTA 1: 103) - kibbiensis Chipp (FWTA 1: 103) = R. yaundensis - lepidobotrys Mildbr. (Achoudoung, to be published) ✩ - liberica Engl. (FWTA 1: 104) - longicuspis Engl. (FWTA 1: 103) = R. welwitschii ✩ - microdon M.Brandt (FWTA 1: 104) ✩ - oblanceolata Chipp (FWTA 1: 103) - oblongifolia (C.H.Wright) Marquand ex Chipp (FWTA 1: 104) ✩ - prasina (Stapf) Chipp (FWTA 1: 103) - preussii Engl. (Achoudoung, to be published) - rubrotincta Chipp (FWTA 1: 104) - subintegrifolia (P.Beauv.) O.Kuntze (FWTA 1: 104) - welwitschii (Oliver) O.Ktze. (Kew Bull. 16: 418) ✪ - whytei (Stapf) M.Brandt (FWTA 1: 104) - yaundensis Engl. (according to Achoudoung, to be published)

Viscaceae ✩ Viscum congolense De Wild. (Polhill & Wiens, 1998: 283)

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Vitaceae Ampelocissus africana (Lour.) Merrill (Fl.Cam.: 10) - bombycina (Baker) Planch. (Fl.Cam.: 8) - gracilipes Stapf (FWTA 1: 682) - leonensis (Hook.f.) Planch. (FWTA 1: 682) - multistriata (Baker) Planch. (FWTA 1: 682) = A. pentaphylla - pentaphylla (Guillaumet & Perr.) Gilg & Brandt (Fl.Cam.: 15) Cayratia debilis (Baker) Suesseng. (Fl.Cam.: 22) - gracilis (Guillaumet & Perr.) Suesseng. (Fl.Cam.: 21) - ibuensis (Hook.f.) Suesseng. (Fl.Cam.: 26) Cissus adenopoda Sprague (FWTA 1: 679) = Cyphostemma adenopodum - aralioides (Welw. ex Baker) Planch. (FWTA 1: 679) - arguta Hook.f. (FWTA 1: 678) - cymosa Schum. & Thonn. (FWTA 1: 680) = Cyphostemma cymosum - debilis (Baker) Planch. (FWTA 1: 679) = Cayratia d. - diffusiflora (Baker) Planch. (FWTA 1: 678) - glaucophylla Hook.f. (FWTA 1: 678) - gracilis Guillaumet & Perr. (FWTA 1: 679) = Cayratia g. ✩ - miegei Tchoumé (183 Boissiera 58: 182) - oreophila Gilg & Brandt (FWTA 1: 677) - ornata A.Cheval. ex Hutch. & Dalziel (FWTA 1: 679) = Cyphostemma ornatum - petiolata Hook.f. (FWTA 1: 678) - polyantha Gilg & Brandt (FWTA 1: 678) - producta Afzel. (FWTA 1: 678) - rubrosetosa Gilg & Brandt (FWTA 1: 679) = Cyphostemma rubrosetosum - smithiana (Baker) Planch. (Fl.Cam.) ✩ - sylvestris Tchoumé (183 Boissiera 58: 183) - vogelii Hook.f. (FWTA 1: 680) = Cyphostemma v. ✩ Cyphostemma adamii Desc. (Adansonia sér. 2, 12: 307) - adenopodum (Sprague) Desc. (Fl.Cam.: 46) - cymosum (Schum. & Thonn.) Desc. (Fl.Cam.: 60) - ornatum (A.Cheval.) Desc. (Fl.Cam.: 74) - rubrosetosum (Gilg & Brandt) Desc. (Fl.Cam.: 56) - vogelii (Hook.f.) Desc. (Fl.Cam.: 57) Leea guineensis G.Don (Fl.Cam.: 134) Rhoicissus revoilii Planch. (FWTA 1: 681)

Zingiberaceae Aframomum angustifolium (Sonnerat) K.Schum. (Kew Bull. 35: 302) - atewae Lock & Hall (Kew Bull. 35: 305) - baumannii K.Schum. (FWTA 3: 76) = A. angustifolium ✩ - chrysanthum Lock (Kew Bull. 35: 302) ✩ - cordifolium Lock & Hall (Kew Bull. 35: 309) - daniellii (Hook.f.) K.Schum. (FWTA 3: 76) = A. chrysanthum ✩ - elegans Lock (Kew Bull. 35: 304) ✩ - elliottii (Baker) K.Schum. (Kew Bull. 35: 304) ✩ - excapum (Sims) Hepper (Kew Bull. 35: 310) - geocarpum Lock & Hall (Kew Bull. 35: 307) - leptolepis (Hook.f.) K.Schum. (Kew Bull. 35: 309) = A. geocarpum ✩ - longiscapum K.Schum. (Kew Bull. 35: 309) - melegueta K.Schum. (Kew Bull. 35: 303) - sceptrum (Oliver & Hanb.) K.Schum. (Kew Bull. 35: 310) - stanfieldii Hepper (Kew Bull. 35: 310) ✩ - strobilaceum (Sm.) Hepper (Kew Bull. 35: 308) - subcericeum (Oliver & Hanb.) K.Schum. (Kew Bull. 35: 305) ✩ - sulcatum K.Schum. (Kew Bull. 35: 308) ✪Renealmia battenbergiana Cummins ex Baker (FWTA 3: 70) ✪ - longifolia K.Schum. (FWTA 3: 70) ✩ - macrocolea K.Schum. (FWTA 3: 70) ✩ - maculata Stapf (FWTA 3: 70) ✪

Key to symbols and colour ✩ Rare and endemic species selection used in Chapter 6 ✪ Rare and endemic species selection + species treated in Chapter 9 ● Species treated in Chapter 9 only Species in green Upper Guinea endemics

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List of all genera and the plant families they belong to Fern and fern-allies Adiantum - Pteridaceae Anisosorus - Dennstaedtiaceae Antrophyum - Vittariaceae Arthropteris - Oleandraceae Asplenium - Aspleniaceae Belvisia - Polypodiaceae Blotiella - Dennstaedtiaceae Bolbitis - Lomariopsidaceae Coniogramme - Pteridaceae Ctenitis - Dryopteridaceae Ctenopteris - Grammitidaceae Cyathea - Cyatheaceae Cyclosorus - Thelypteridaceae Davallia - Davalliaceae Dicranopteris - Gleicheniaceae Diplazium - Dryopteridaceae Doryopteris - Pteridaceae Drynaria - Polypodiaceae Dryopteris - Dryopteridaceae Elaphoglossum - Lomariopsidaceae Gleichenia - Gleicheniaceae Huperzia jaegeri - Lycopodiaceae Hymenophyllum - Hymenophyllaceae Lastreopsis - Dryopteridaceae Lindsaea - Dennstaedtiaceae Lomariopsis - Lomariopsidaceae Lonchitis - Dennstaedtiaceae Loxogramme - Polypodiaceae Lycopodiella - Lycopodiaceae Lycopodium - Lycopodiaceae Lygodium - Schizaeaceae Marattia - Marattiaceae Microgramma - Polypodiaceae Microlepia - Dennstaedtiaceae Microsorium - Polypodiaceae Nephrolepis - Nephrolepidaceae Oleandra - Oleandraceae Ophioglossum - Ophioglossaceae Pellaea - Pteridaceae Phymatodes - Polypodiaceae Phymatosorus - Polypodiaceae Pityrogramma - Pteridaceae Platycerium - Polypodiaceae Pleopeltis - Polypodiaceae Psilotum - Psilotaceae Pseudophegopteris - Thelypteridaceae Pteridium - Dennstaedtiaceae Pteris - Pteridaceae Pyrrosia - Polypodiaceae Schizolegnia - Dennstaedtiaceae Selaginella - Selaginellaceae Tectaria - Dryopteridaceae Thelypteris - Thelypteridaceae Trichomanes - Hymenophyllaceae Triplophyllum - Dryopteridaceae Vittaria - Vittariaceae Xiphopteris - Grammitidaceae

Flowering plants Abrus - Leguminosae-Pap. Acacia - Leguminosae-Mim. Acalypha - Euphorbiaceae Acanthus - Acanthaceae Achyranthes - Amaranthaceae

Achyrospermum - Labiatae Acioa - Chrysobalanaceae Acridocarpus - Malpighiaceae Acroceras - Graminae Adenia - Passifloraceae Adenopodia - Leguminosae-Mim. Adenopus - Cucurbitaceae Adhatoda - Acanthaceae Aeglopsis - Rutaceae Aerangis - Orchidaceae Afraegle - Rutaceae Aframomum - Zingiberaceae Afrobrunnichia - Polygonaceae Afrolicania - Chrysobalanaceae Afrormosia - Leguminosae-Pap. Afrosersalisia - Sapotaceae Afrostyrax - Huaceae Afzelia - Leguminosae-Caes. Aganope - Leguminosae-Pap. Agelaea - Connaraceae Agelanthus - Loranthaceae Aidia - Rubiaceae Airyantha - Leguminosae-Pap. Alafia - Apocynaceae Albertisia - Menispermaceae Albizia - Leguminosae-Mim. Alchornea - Euphorbiaceae Allanblackia - Guttiferae Allexis - Violaceae Allophylus - Sapindaceae Alsodeiopsis - Icacinaceae Alstonia - Apocynaceae Amanoa - Euphorbiaceae Amauriella - Araceae Amorphophallus - Araceae Ampelocissus - Vitaceae Amphiblemma - Melastomataceae Amphimas - Leguminosae-Pap. Anchomanes - Araceae Ancistrochilus - Orchidaceae Ancistrocladus - Ancistrocladaceae Ancistrophyllum - Palmae Ancistrorhynchus - Orchidaceae Ancylobothrys - Apocynaceae Andira - Leguminosae-Pap. Androsiphonia - Passifloraceae Aneilema - Commelinaceae Angraecopsis - Orchidaceae Angraecum - Orchidaceae Angylocalyx - Leguminosae-Pap. Aningeria - Sapotaceae Aniseia - Convolvulaceae Anisophyllea - Anisophylleaceae Anisopus - Asclepiadaceae Anisotes - Acanthaceae Annickia - Annonaceae Anogeissus - Combretaceae Anonidium - Annonaceae Anopyxis - Rhizophoraceae Ansellia - Orchidaceae Anthocleista - Gentianaceae Anthoclitandra - Apocynaceae Anthonotha - Leguminosae-Caes. Anthostema - Euphorbiaceae Antiaris - Moraceae Antidesma - Euphorbiaceae Antrocaryon - Anacardiaceae Anubias - Araceae

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Aphania - Sapindaceae Aphanocalyx - Leguminosae-Caes. Aphanostylis - Apocynaceae Apodiscus - Euphorbiaceae Apodostigma - Celastraceae Aporrhiza - Sapindaceae Aptandra - Olacaceae Araliopsis - Rutaceae Argocoffeopsis - Rubiaceae Argomuellera - Euphorbiaceae Aristolochia - Aristolochiaceae Artabotrys - Annonaceae Asparagus - Liliaceae Asystasia - Acanthaceae Ataenidia - Marantaceae Atractogyne - Rubiaceae Atroxima - Polygalaceae Aubregrinia - Sapotaceae Aubrevillea - Leguminosae-Mim. Aulacocalyx - Rubiaceae Auxopus - Orchidaceae Avicennia - Avicenniaceae Axonopus - Graminae Azadirachta - Meliaceae Baissea - Apocynaceae Balanites - Balanitaceae Baphia - Leguminosae-Pap. Baphiastrum - Leguminosae-Pap. Barleria - Acanthaceae Begonia - Begoniaceae Beilschmiedia - Lauraceae Belonophora - Rubiaceae Bequaertia - Celastraceae Bequaertiodendron - Sapotaceae Berliniai - Leguminosae-Caes. Bersama - Melianthaceae Bertiera - Rubiaceae Biophytum - Oxalidaceae Blighia - Sapindaceae Bobgunnia - Leguminosae-Pap. Boehmeria - Urticaceae Bolusiella - Orchidaceae Bombax - Bombacaceae Bonamia - Convolvulaceae Borreria - Rubiaceae Bosqueia - Moraceae Bourdaria - Melastomataceae Bowringia - Leguminosae-Pap. Brachycorythis - Orchidaceae Brachystegia - Leguminosae-Caes. Brachystephanus - Acanthaceae Breviea - Sapotaceae Bridelia - Euphorbiaceae Brieya - Annonaceae Brillantaisia - Acanthaceae Brucea - Simaroubaceae Buchholzia - Capparaceae Buforrestia - Commelinaceae Bulbophyllum - Orchidaceae Burmannia - Burmanniaceae Bussea - Leguminosae-Caes. Buxus - Buxaceae Byrsanthus - Flacourtiaceae Byrsocarpus - Connaraceae Byttneria - Sterculiaceae Caesalpinia - Leguminosae-Caes. Calamus - Palmae Calanthe - Orchidaceae Callichilia - Apocynaceae Caloncoba - Flacourtiaceae Calpocalyx - Leguminosae-Mim. Calvoa - Melastomataceae Calycobolus - Convolvulaceae Calycosiphonia - Rubiaceae Calyptrochilum - Orchidaceae Campylospermum - Ochnaceae Campylostemon - Celastraceae

Canarium - Burseraceae Canthium - Rubiaceae Capparis - Capparaceae Carapa - Meliaceae Cardiospermum - Sapindaceae Carex - Cyperaceae Carissa - Apocynaceae Carpolobia - Polygalaceae Casearia - Flacourtiaceae Cassia - Leguminosae-Caes. Cassine - Celastraceae Cassipourea - Rhizophoraceae Castanola - Connaraceae Cathormion - Leguminosae-Mim. Cavacoa - Euphorbiaceae Cayaponia - Cucurbitaceae Cayratia - Vitaceae Cecropia - Moraceae Ceiba - Bombacaceae Celosia - Amaranthaceae Celtis - Ulmaceae Centotheca - Graminae Centrosema - Leguminosae-Pap. Cephaelis - Rubiaceae Cercestis - Araceae Ceropegia - Asclepiadaceae Chaetacme - Ulmaceae Chamaeangis - Orchidaceae Chasmanthera - Menispermaceae Chassalia - Rubiaceae Chauliodon - Orchidaceae Chazaliella - Rubiaceae Cheirostylis - Orchidaceae Chidlowia - Leguminosae-Caes. Chionanthus - Oleaceae Chlamydocardia - Acanthaceae Chlamydocarya - Icacinaceae Chlorophora - Moraceae Chlorophytum - Liliaceae Christiana - Tiliaceae Chromolaena - Compositae Chrysobalanus - Chrysobalanaceae Chrysophyllum - Sapotaceae Chytranthus - Sapindaceae Cincinnobotrys - Melastomataceae Cissampelos - Menispermaceae Cissus - Vitaceae Citropsis - Rutaceae Claoxylon - Euphorbiaceae Clausena - Rutaceae Cleidion - Euphorbiaceae Cleistanthus - Euphorbiaceae Cleistopholis - Annonaceae Clematis - Ranunculaceae Clerodendrum - Verbenaceae Clitandra - Apocynaceae Cnestis - Connaraceae Coccinia - Cucurbitaceae Coelocaryon - Myristicaceae Coffea - Rubiaceae Coix - Graminae Cola - Sterculiaceae Coleotrype - Commelinaceae Combretodendron - Lecythidaceae Combretum - Combretaceae Commelina - Commelinaceae Commelinidium - Graminae Commiphora - Burseraceae Connarus - Connaraceae Conopharyngia - Apocynaceae Copaifera - Leguminosae-Caes. Cordia - Boraginaceae Corymborkis - Orchidaceae Corynanthe - Rubiaceae Costus - Costaceae Coula - Olacaceae Craibia - Leguminosae-Pap.

Crassocephalum - Compositae Craterispermum - Rubiaceae Craterogyne - Moraceae Craterosiphon - Thymelaeaceae Crateva - Capparaceae Cremaspora - Rubiaceae Cribbia - Orchidaceae Crinum - Amaryllidaceae Crossandra - Acanthaceae Crossandrella - Acanthaceae Crossonephelis - Sapindaceae Crossostemma - Passifloraceae Croton - Euphorbiaceae Crotonogyne - Euphorbiaceae Crotonogynopsis - Euphorbiaceae Crudia - Leguminosae-Caes. Cryptolepis - Asclepiadaceae Cryptosepalum - Leguminosae-Caes. Cuervea - Celastraceae Culcasia - Araceae Cussonia - Araliaceae Cuviera - Rubiaceae Cyanastrum - Tecophyllaeaceae Cyathula - Amaranthaceae Cylicodiscus - Leguminosae-Mim. Cynanchum - Asclepiadaceae Cynometra - Leguminosae-Caes. Cyperus - Cyperaceae Cyphostemma - Vitaceae Cyrtococcum - Graminae Cyrtorchis - Orchidaceae Cyrtosperma - Araceae Dacryodes - Burseraceae Dactyladenia - Chrysobalanaceae Dalbergia - Leguminosae-Pap. Dalbergiella - Leguminosae-Pap. Dalechampia - Euphorbiaceae Daniellia - Leguminosae-Caes. Dasylepis - Flacourtiaceae Decorsella - Violaceae Deinbollia - Sapindaceae Delpydora - Sapotaceae Dennettia - Annonaceae Desmodium - Leguminosae-Pap. Desmostachys - Icacinaceae Desplatsia - Tiliaceae Detarium - Leguminosae-Caes. Dialium - Leguminosae-Caes. Diaphananthe - Orchidaceae Dicellandra - Melastomataceae Diceratostele - Orchidaceae Dichaetanthera - Melastomataceae Dichapetalum - Dichapetalaceae Dicliptera - Acanthaceae Dicranolepis - Thymelaeaceae Dictyandra - Rubiaceae Dictyophleba - Apocynaceae Didelotia - Leguminosae-Caes. Didymoplexis - Orchidaceae Didymosalpinx - Rubiaceae Digitaria - Graminae Dimorphochlamys - Cucurbitaceae Dinklageella - Orchidaceae Dinklageodoxa - Bignoniaceae Dinophora - Melastomataceae Dioclea - Leguminosae-Pap. Dioscorea -Dioscoreaceae Dioscoreophyllum - Menispermaceae Diospyros - Ebenaceae Diphasia - Rutaceae Diplacrum - Cyperaceae Discoclaoxylon - Euphorbiaceae Discoglypremna - Euphorbiaceae Disperis - Orchidaceae Dissomeria - Flacourtiaceae Dissotis - Melastomataceae Distemonanthus - Leguminosae-Caes.

Dolichos - Leguminosae-Pap. Dombeya - Sterculiaceae Dorstenia - Moraceae Dovyalis - Flacourtiaceae Dracaena - Dracaenaceae Dregea - Asclepiadaceae Drepanocarpus - Leguminosae-Pap. Drypetes - Euphorbiaceae Duboscia - Tiliaceae Duguetia - Annonaceae Dumoria - Sapotaceae Duparquetia - Leguminosae-Caes. Eggelingia - Orchidaceae Ehretia - Boraginaceae Ekebergia - Meliaceae Elaeis - Palmae Elaeodendron - Celastraceae Elaeophorbia - Euphorbiaceae Elatostema - Urticaceae Elytraria - Acanthaceae Embelia - Myrsinaceae Enantia - Annonaceae Encheiridion - Orchidaceae Endosiphon - Acanthaceae Endotricha - Sapotaceae Englerina - Loranthaceae Englerophytum - Sapotaceae Enneastemon - Annonaceae Entada - Leguminosae-Mim. Entandrophragma - Meliaceae Epinetrum - Menispermaceae Epipogium - Orchidaceae Epistemma - Asclepiadaceae Epithema - Gesneriaceae Eremomastax - Acanthaceae Eremospatha - Palmae Eribroma - Sterculiaceae Eriocoelum - Sapindaceae Erythrina - Leguminosae-Pap. Erythrococca - Euphorbiaceae Erythrophleum - Leguminosae-Caes. Erythroxylum - Erythroxylaceae Euadenia - Capparaceae Euclinia - Rubiaceae Eugenia - Myrtaceae Eulophia - Orchidaceae Eulophidium - Orchidaceae Eupatorium - Compositae Eurychone - Orchidaceae Excoecaria - Euphorbiaceae Exolobus - Asclepiadaceae Fagara - Rutaceae Farquharia - Apocynaceae Fegimanra - Anacardiaceae Ficus - Moraceae Flabellaria - Malpighiaceae Flacourtia - Flacourtiaceae Flagellaria - Flagellariaceae Fleurya - Urticaceae Floscopa - Commelinaceae Friesodielsia - Annonaceae Funtumia - Apocynaceae Gaertnera - Rubiaceae Gambeya - Sapotaceae Garcinia - Guttiferae Gardenia - Rubiaceae Gelonium - Euphorbiaceae Genyorchis - Orchidaceae Geophila - Rubiaceae Gerrardanthus - Cucurbitaceae Gilbertiodendron - Leguminosae-Caes. Gilletiodendron - Leguminosae-Caes. Girardinia - Urticaceae Glenniea - Sapindaceae Globimetula - Loranthaceae Gloriosa - Liliaceae Gluema - Sapotaceae

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Glyphaea - Tiliaceae Gongronema - Asclepiadaceae Gouania - Rhamnaceae Graphorchis - Orchidaceae Greenwayodendron - Annonaceae Grewia - Tiliaceae Griffonia - Leguminosae-Caes. Grossera - Euphorbiaceae Guaduella - Graminae Guarea - Meliaceae Guibourtia - Leguminosae-Caes. Guyonia - Melastomataceae Gymnorinorea - Violaceae Gymnosiphon - Burmanniaceae Gymnostemon - Simaroubaceae Gynura - Compositae Habenaria - Orchidaceae Habropetalum - Dioncophillaceae Haemanthus - Amaryllidaceae Hallea - Rubiaceae Halopegia - Marantaceae Hannoa - Simaroubaceae Haplormosia - Leguminosae-Pap. Harrisonia - Simaroubaceae Harungana - Guttiferae Heckeldora - Meliaceae Heinsia - Rubiaceae Heisteria - Olacaceae Helictonema - Celastraceae Hemandradenia - Connaraceae Heritiera - Sterculiaceae Hetaeria - Orchidaceae Heteradelphia - Acanthaceae Heteropteris - Malpighiaceae Heterotis - Melastomataceae Hexalobus - Annonaceae Hibiscus - Malvaceae Hildegardia - Sterculiaceae Hilleria - Phytolaccaceae Hippocratea - Celastraceae Hirtella - Chrysobalanaceae Holarrhena - Apocynaceae Holoptelea - Ulmaceae Homalium - Flacourtiaceae Homocolleticon - Orchidaceae Hoplestigma - Hoplestigmataceae Hoslundia - Labiatae Hugonia - Linaceae Hunteria - Apocynaceae Hutchinsonia - Rubiaceae Hymenocardia - Euphorbiaceae Hymenocoleus - Rubiaceae Hymenodictyon - Rubiaceae Hymenostegia - Leguminosae-Caes. Hypoestes - Acanthaceae Hypolytrum - Cyperaceae Hypselodelphys - Marantaceae Icacina - Icacinaceae Idertia - Ochnaceae Ilex - Aquifoliaceae Illigera - Hernandiaceae Impatiens - Balsaminaceae Indigofera - Leguminosae-Pap. Inhambanella - Sapotaceae Inversodicraea - Podostemaceae Iodes - Icacinaceae Ipomoea - Convolvulaceae Irvingia - Irvingiaceae Isachne - Graminae Isolona - Annonaceae Isomacrolobium - Leguminosae-Caes. Isonema - Apocynaceae Ituridendron - Sapotaceae Ixora - Rubiaceae Jasminum - Oleaceae Jaundea - Connaraceae Justicia - Acanthaceae

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Kantou - Sapotaceae Kaoue - Leguminosae-Caes. Keayodendron - Euphorbiaceae Keetia - Rubiaceae Khaya - Meliaceae Kigelia - Bignoniaceae Klainedoxa - Irvingiaceae Kolobopetalum - Menispermaceae Kornasia - Orchidaceae Laccodiscus - Sapindaceae Laccosperma - Palmae Lagenaria - Cucurbitaceae Laguncularia - Combretaceae Landolphia - Apocynaceae Lankesteria - Acanthaceae Lannea - Anacardiaceae Laportea - Urticaceae Lasianthus - Rubiaceae Lasimorpha - Araceae Lasiodiscus - Rhamnaceae Lecaniodiscus - Sapindaceae Ledermanniella - Podostemaceae Leea - Vitaceae Lemonniera - Sapotaceae Lepidagathis - Acanthaceae Lepisanthes - Sapindaceae Lepistemon - Convolvulaceae Leptactina - Rubiaceae Leptaspis - Graminae Leptaulus - Icacinaceae Leptoderris - Leguminosae-Pap. Leptonychia - Sterculiaceae Leucas - Labiatae Leucomphalos - Leguminosae-Pap. Licania - Chrysobalanaceae Lijndenia - Melastomataceae Lindackeria - Flacourtiaceae Linociera - Oleaceae Liparis - Orchidaceae Lisowskia - Orchidaceae Listrostachys - Orchidaceae Loesenera - Leguminosae-Caes. Loeseneriella - Celastraceae Lonchocarpus - Leguminosae-Pap. Lophira - Ochnaceae Lovoa - Meliaceae Lychnodiscus - Sapindaceae Maba - Ebenaceae Macaranga - Euphorbiaceae Machaerium - Leguminosae-Pap. Macropodiella - Podostemaceae Macrosphyra - Rubiaceae Maerua - Capparaceae Maesa - Myrsinaceae Maesobotrya - Euphorbiaceae Maesopsis - Rhamnaceae Magnistipula - Chrysobalanaceae Majidea - Sapindaceae Malacantha - Sapotaceae Malaxis - Orchidaceae Mallotus - Euphorbiaceae Malouetia - Apocynaceae Mammea - Guttiferae Mangenotia - Asclepiadaceae Manilkara - Sapotaceae Mannia - Simaroubaceae Manniella - Orchidaceae Manniophyton - Euphorbiaceae Manotes - Connaraceae Mansonia - Sterculiaceae Mapania - Cyperaceae Maranthes - Chrysobalanaceae Marantochloa - Marantaceae Mareya - Euphorbiaceae Margaritaria - Euphorbiaceae Markhamia - Bignoniaceae Marsdenia - Asclepiadaceae

Martretia - Euphorbiaceae Maschalocephalus - Rapateaceae Massularia - Rubiaceae Maytenus - Celastraceae Medinilla - Melastomataceae Megaphrynium - Marantaceae Megastachya - Graminae Melanthera - Compositae Melastomastrum - Melastomataceae Melothria - Cucurbitaceae Memecylon - Melastomataceae Mendoncia - Acanthaceae Mezoneuron - Leguminosae-Caes. Microcoelia - Orchidaceae Microdesmis - Pandaceae Microglossa - Compositae Mikania - Compositae Mikaniopsis - Compositae Mildbraedia - Euphorbiaceae Mildbraediodendron - Legum.-Caes. Milicia - Moraceae Millettia - Leguminosae-Pap. Mimusops - Sapotaceae Mischogyne - Annonaceae Mitragyna - Rubiaceae Momordica - Cucurbitaceae Monanthotaxis - Annonaceae Mondia - Asclepiadaceae Monocyclanthus - Annonaceae Monodora - Annonaceae Monopetalanthus - Leguminosae-Caes. Monosalpinx - Rubiaceae Morelia - Rubiaceae Morinda - Rubiaceae Morus - Moraceae Mostuea - Gelsemiaceae Motandra - Apocynaceae Mucuna - Leguminosae-Pap. Multidentia - Rubiaceae Musanga - Moraceae Mussaenda - Rubiaceae Myrianthus - Moraceae Napoleonaea - Lecythidaceae Nauclea - Rubiaceae Necepsia - Euphorbiaceae Nelsonia - Acanthaceae Neoboivinella - Sapotaceae Neoboutonia - Euphorbiaceae Neocarya - Chrysobalanaceae Neolemonniera - Sapotaceae Neorosea - Rubiaceae Neosloetiopsis - Moraceae Neostachyanthus - Icacinaceae Neostenanthera - Annonaceae Nephrangis - Orchidaceae Nephthytis - Araceae Nervilia - Orchidaceae Nesogordonia - Sterculiaceae Neuropeltis - Convolvulaceae Newbouldia - Bignoniaceae Newtonia - Leguminosae-Mim. Nichallea - Rubiaceae Nothospondias - Simaroubaceae Notobuxus - Buxaceae Nuxia - Stilbaceae Oberonia - Orchidaceae Ochna - Ochnaceae Ochthocharis - Melastomataceae Ochthocosmus - Linaceae Octoknema - Olacaceae Octolepis - Thymelaeaceae Octolobus - Sterculiaceae Oeceoclades - Orchidaceae Okoubaka - Santalaceae Olax - Olacaceae Oldenlandia - Rubiaceae Oldfieldia - Euphorbiaceae

Olea - Oleaceae Olyra - Graminae Omphalocarpum - Sapotaceae Omphalogonus - Asclepiadaceae Oncinotis - Apocynaceae Oncoba - Flacourtiaceae Ongokea - Olacaceae Operculina - Convolvulaceae Ophiobotrys - Flacourtiaceae Opilia - Opiliaceae Oplismenus - Graminae Oricia - Rutaceae Ormocarpum - Leguminosae-Pap. Orthopichonia - Apocynaceae Orthosiphon - Labiatae Ostryocarpus - Leguminosae-Pap. Ostryoderris - Leguminosae-Pap. Otomeria - Rubiaceae Ouratea - Ochnaceae Oxyanthus - Rubiaceae Oxymitra - Annonaceae Oxystelma - Asclepiadaceae Pachypodanthium - Annonaceae Pachystela - Sapotaceae Palisota - Commelinaceae Pancovia - Sapindaceae Panda - Pandaceae Pandanus - Pandanaceae Panicum - Graminae Paramacrolobium - Legum.-Caes. Parapentas - Rubiaceae Pararistolochia - Aristolochiaceae Parinari - Chrysobalanaceae Parkia - Leguminosae-Mim. Paropsia - Passifloraceae Parquetina - Asclepiadaceae Paspalum - Graminae Paullinia - Sapindaceae Pauridiantha - Rubiaceae Pausinystalia - Rubiaceae Pavetta - Rubiaceae Peddiea - Thymelaeaceae Pellegriniodendron - Legum.-Caes. Penianthus - Menispermaceae Pentaclethra - Leguminosae-Mim. Pentadesma - Guttiferae Peperomia - Piperaceae Peponium - Cucurbitaceae Pergularia - Asclepiadaceae Pericopsis - Leguminosae-Pap. Periploca - Asclepiadaceae Petersianthus - Lecythidaceae Phaeoneuron - Melastomataceae Phaulopsis - Acanthaceae Philenoptera - Leguminosae-Pap. Phragmanthera - Loranthaceae Phyllanthus - Euphorbiaceae Phyllocosmus - Linaceae Physacanthus - Acanthaceae Physedra - Cucurbitaceae Physostigma - Leguminosae-Pap. Phytolacca - Phytolaccaceae Picralima - Apocynaceae Pierreodendron - Simaroubaceae Pilea - Urticaceae Piper - Piperaceae Piptadeniastrum - Leguminosae-Mim. Piptostigma - Annonaceae Pisonia - Nyctaginaceae Pittosporum - Pittosporaceae Placodiscus - Sapindaceae Plagiosiphon - Leguminosae-Caes. Platylepis - Orchidaceae Platysepalum - Leguminosae-Pap. Plectrelminthus - Orchidaceae Pleiocarpa - Apocynaceae Pleioceras - Apocynaceae

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Pleiocoryne - Rubiaceae Podandriella - Orchidaceae Podangis - Orchidaceae Poecilocalyx - Rubiaceae Pollia - Commelinaceae Polyalthia - Annonaceae Polycephalium - Icacinaceae Polyceratocarpus - Annonaceae Polycoryne - Rubiaceae Polypleurum - Podostemaceae Polyscias - Araliaceae Polyspatha - Commelinaceae Polysphaeria - Rubiaceae Polystachya - Orchidaceae Polystemonanthus - Legum.-Caes. Popowia - Annonaceae Pouchetia - Rubiaceae Pouteria - Sapotaceae Pouzolzia - Urticaceae Premna - Verbenaceae Preussiella - Melastomataceae Prionostemma - Celastraceae Pristimera - Celastraceae Procris - Urticaceae Protomegabaria - Euphorbiaceae Pseudagrostistachys - Euphorbiaceae Pseudechinolaena - Graminae Pseuderanthemum - Acanthaceae Pseudocalyx - Acanthaceae Pseudoprosopis - Leguminosae-Mim. Pseudospondias - Anacardiaceae Psilanthus - Rubiaceae Psophocarpus - Leguminosae-Pap. Psorospermum - Guttiferae Psychotria - Rubiaceae Psydrax - Rubiaceae Pteleopsis - Combretaceae Pterocarpus - Leguminosae-Pap. Pterygota - Sterculiaceae Ptychopetalum - Olacaceae Puelia - Graminae Pycnanthus - Myristicaceae Pycnocoma - Euphorbiaceae Pyrenacantha - Icacinaceae Pyrostria - Rubiaceae Quisqualis - Combretaceae Rangaeris - Orchidaceae Raphia Palmae Raphidiocystis - Cucurbitaceae Rauvolfia - Apocynaceae Reissantia - Celastraceae Remusatia - Araceae Renealmia - Zingiberaceae Rhabdophyllum - Ochnaceae Rhaphidophora - Araceae Rhaphiostylis - Icacinaceae Rhaptopetalum - Scytopetalaceae Rhigiocarya - Menispermaceae Rhinacanthu - Acanthaceae s Rhipidoglossum - Orchidaceae Rhipsalis - Cactaceae Rhizophora - Rhizophoraceae

Rhodognaphalon - Bombacaceae Rhoicissus - Vitaceae Rhynchosia - Leguminosae-Pap. Ricinodendron - Euphorbiaceae Rinorea - Violaceae Ritchiea - Capparaceae Robynsia - Rubiaceae Rothmannia - Rubiaceae Rourea - Connaraceae Rubus - Rosaceae Ruellia - Acanthaceae Rungia - Acanthaceae Ruthalicia - Cucurbitaceae Rutidea - Rubiaceae Rytigynia - Rubiaceae Saba - Apocynaceae Sabicea - Rubiaceae Sacoglottis - Humiriaceae Sacosperma - Rubiaceae Sakersia - Melastomataceae Salacia - Celastraceae Salacighia - Celastraceae Samanea - Leguminosae-Mim. Santaloides - Connaraceae Santiria - Burseraceae Sapium - Euphorbiaceae Sarcocephalus - Rubiaceae Sarcophrynium - Marantaceae Sarcostemma - Asclepiadaceae Saxicolella - Podostemaceae Scadoxus - Amaryllidaceae Scaphopetalum - Sterculiaceae Schefflera - Araliaceae Schizocolea - Rubiaceae Schrebera - Oleaceae Schumanniophyton - Rubiaceae Sciaphila - Triuridaceae Scleria - Cyperaceae Sclerochiton - Acanthaceae Sclerocroton - Euphorbiaceae Sclerosperma - Palmae Scottellia - Flacourtiaceae Scytopetalum - Scytopetalaceae Sebaea - Gentianaceae Secamone - Asclepiadaceae Securidaca - Polygalaceae Senna - Leguminosae-Caes. Sericanthe - Rubiaceae Sericostachys - Amaranthaceae Setaria - Graminae Sherbournia - Rubiaceae Shirakiopsis - Euphorbiaceae Sideroxylon - Sapotaceae Simicratea - Celastraceae Simirestis - Celastraceae Sloetiopsis - Moraceae Smeathmannia - Passifloraceae Smilax - Liliaceae Solanecio - Compositae Solanum - Solanaceae Solenangis - Orchidaceae Solenostemon - Labiatae

Sorindeia - Anacardiaceae Soyauxia - Medusandraceae Spathandra - Melastomataceae Spathodea - Bignoniaceae Spermacoce - Rubiaceae Sphenocentrum - Menispermaceae Spiropetalum - Connaraceae Spondianthus - Euphorbiaceae Spondias - Anacardiaceae Stachyanthus - Icacinaceae Stachyothyrsus - Leguminosae-Caes. Stanfieldiella - Commelinaceae Staurogyne - Acanthaceae Stelechantha - Rubiaceae Stemonocoleus - Leguminosae-Caes. Stenandriopsis - Acanthaceae Stenandrium - Acanthaceae Stephania - Menispermaceae Sterculia - Sterculiaceae Stereospermum - Bignoniaceae Stictocardia - Convolvulaceae Stigmaphyllon - Malpighiaceae Stolzia - Orchidaceae Streblus - Moraceae Strephonema - Combretaceae Streptocarpus - Gesneriaceae Streptogyna - Graminae Strombosia - Olacaceae Strombosiopis - Olacaceae Strophanthus - Apocynaceae Strychnos - Loganiaceae Stylochaeton - Araceae Summerhayesia - Orchidaceae Suregada - Euphorbiaceae Swartzia - Leguminosae-Pap. Symmeria - Polygonaceae Symphonia - Guttiferae Synsepalum - Sapotaceae Syntriandrium - Menispermaceae Syrrhonema - Menispermaceae Syzygium - Myrtaceae Tabernaemontana - Apocynaceae Tacazzea - Asclepiadaceae Tacca - Taccaceae Taeniophyllum - Orchidaceae Talbotiella - Leguminosae-Caes. Tapinanthus - Loranthaceae Tapura - Dichapetalaceae Tarenna - Rubiaceae Tarrietia - Sterculiaceae Teclea - Rutaceae Telfairia - Cucurbitaceae Telosma - Asclepiadaceae Terminalia - Combretaceae Tessmannia - Leguminosae-Caes. Tetraberlinia - Leguminosae-Caes. Tetracera - Dilleniaceae Tetrapleura - Leguminosae-Mim. Tetrorchidium - Euphorbiaceae Thalia - Marantaceae Thaumatococcus - Marantaceae Thecacoris - Euphorbiaceae

Thespesia - Malvaceae Thomandersia - Acanthaceae Thonningia - Balanophoraceae Thunbergia - Acanthaceae Tieghemella - Sapotaceae Tiliacora - Menispermaceae Toubaouate - Leguminosae-Caes. Trachyphrynium - Marantaceae Tragia - Euphorbiaceae Treculia - Moraceae Trema - Ulmaceae Triaspis - Malpighiaceae Tricalysia - Rubiaceae Trichilia - Meliaceae Trichoscypha - Anacardiaceae Trichostachys - Rubiaceae Triclisia - Menispermaceae Tridactyle - Orchidaceae Trilepisium - Moraceae Triphyophyllum - Dioncophillaceae Triplisomeris - Leguminosae-Caes. Triplochiton - Sterculiaceae Tristemma - Melastomataceae Tristemonanthus - Celastraceae Turraea - Meliaceae Turraeanthus - Meliaceae Tylophora - Asclepiadaceae Uapaca - Euphorbiaceae Uncaria - Rubiaceae Urera - Urticaceae Urobotrya - Opiliaceae Urophyllum - Rubiaceae Usteria - Loganiaceae Uvaria - Annonaceae Uvariastrum - Annonaceae Uvariodendron - Annonaceae Uvariopsis - Annonaceae Vahadenia - Apocynaceae Vangueria - Rubiaceae Vangueriella - Rubiaceae Vangueriopsis - Rubiaceae Vanilla - Orchidaceae Ventilago - Rhamnaceae Vepris - Rutaceae Vernonia - Compositae Vincentella - Sapotaceae Virectaria - Rubiaceae Viscum - Viscaceae Vismia - Guttiferae Vitex - Verbenaceae Voacanga - Apocynaceae Voyria - Gentianaceae Warneckea - Melastomataceae Whitfieldia - Acanthaceae Xylia - Leguminosae-Mim. Xylopia - Annonaceae Xylopiastrum - Annonaceae Zanha - Sapindaceae Zanthoxylum - Rutaceae Zehneria - Cucurbitaceae Zeuxine - Orchidaceae

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C Appendices Appendices 1-5

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Figure 2.1 Azagny National Park in 2000 consists of forests and swampy savannas. Although this is a national park, the borders are strongly degraded by agricultural activities.

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Figure 2.2 Forest cover in West Africa. (A) Cover according to NOAAAVHRR satellite images (Paivinen et al. 1989). The eastern part of Liberia and western part of Côte d'Ivoire, including Taï National Park, are the last large forested areas that remain.

(B) Cover according to the "ecoregions" map of Olson & Dinerstein (1998) representing the forest cover zonation with the limits in 1912 according to Chevalier (1920) and the limits in 1923 following Shantz & Marbut (1923). The variability seems to be due to the personal view of the authors rather than to changes in forest cover between 1912-23.

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A

B

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Figure 2.3 Forest cover in Côte d’Ivoire in 1955-58 according to the vegetation map (scale 1:500,000) of Guillaumet & Adjanohoun (1969) and limits of the vegetation domains following Monnier (1983).

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Figure 2.4 Forest cover in Côte d’Ivoire in 1993 (Dao 1999) based on NOAA-AVHRR images. The Taï forest area represents at least 40% of the total forest area of Côte d'Ivoire. Also the forest area south of Abengourou (the classified forests of Yaya-Bossématié-Mabi) is very important.

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A

Figure 2.6 Changes in forest cover in the region around Abidjan. (A) Changes between 1990 and 2000, based on LANDSAT images with 30 m resolution. The limits of the classified forests are indicated. Only the Yapo classified forest and Banco National Park (near Abidjan) are completely covered with forest.

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B

(B) Changes between 1958 (topographic map) and 1990 (LANDSAT image) with a resolution of 250 m. The 8 blocks of 20 x 20 km each that are studied in detail (see Fig. 2.8) are indicated.

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A

Figure 2.7 Comparison between two interpretation maps of LANDSAT images with 30 m resolution. (A) Map from the present study, showing all forested areas independent of their size. Also secondary forests are given showing the density of human occupation.

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B

(B) Map from the 1993 "Bilan Forestier" SODEFOR (1993) based on a 1990 image and taking only forest fragments of more than 10,000 ha. The total area is 55 x 55 km.

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A

The Hiré block, in the moist semi-deciduous forest zone. A high reduction of forest cover from 368 km2 in 1958 to 18 km2 (less than 5%) in 1990, and c. 0.4 km2 in 2000. Two thirds of the forests are less than 2 ha in area, and the “forêts classées” were completely cleared in 1990. In 2000 the whole area is characterised by old coffee-cacao plantations.

The Divo block. Deforestation here is quite similar to Hiré (from 337 to 40 to 6 km2). Forests here are a little more abundant due to numerous steep slopes covered with granite outcrops that are less conducive to agriculture. There is a unique tract of forest which is privately owned (ex IRCC). Partial exploitation of it, unfortunately, started in 1991. The Boubo “forêt classée” was completely transformed in oil palm plantations.

The Gô-Bodiénou block. Forest cover reduced from 196 km2 in 1958 to 227 km2 in 1990 to 67 km2 in 2000. This block has much closed broadleaf forest and was highly isolated until 1990. A large part of the block is occupied by the Gô-Bodiénou “forêt classée”. The landscape to the north of this forest is made up of numerous groves. Para-rubber tree and palm plantations have increased heavily. This “forêt classée”, along with those of Yapo and Niegré, are the three last large areas of forest in the South zone of the country.

The Lovigué block. Forest cover reduced from 227 km2 in 1958 to 194 km2 in 1990 to 42 km2 in 2000. Most of the forest area is found outside the “forêts classées”. In 1969, deforestation already reached the borders of the “forêts classées”. In 1990, over 40% of the “forêts classées” of Mafé and Lovigué were already cleared, and in the decade to 2000, most clearing was done in the Mafé “forêt classée”. This block, along with that of Abié, shows the highest forest cover in 2000, mostly situated in the rural domain.

Figure 2.8 Changes in forest cover at a local scale for eight blocks of 20 x 20 km each in the region of Abidjan (see Figure 2.6). (A) blocks located in the eastern part of the area. Left the changes between 1958 and 1990, right the changes between 1990 and 2000.

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The Yapo block. Forest cover changed from 152 km2 in 1958 to 224 km2 in 1990 to 129 km2 in 2000. 14,900 ha of forest is situated within the “forêt classée” whose conservation status seems superficially good. However, the forests are extremely degraded due to high logging levels. This block show the lowest rate of deforestation. In spite of the strong human pressure due to the presence of large banana and pineapple plantations south of the forest, numerous small groves of secondary forests are present where cola-nut trees are often abundant. The block shows a coexistence of industrial along with traditional land use.

B

The Abié block north of the Yapo “forêt classée”. Forest cover changed from 129 km2 in 1958 to 139 km2 in 1990 to 39 km2 in 2000. This block was considered completely cleared on the 1958 maps. It is, however, the block with the highest forest cover in the rural areas. Land occupation according to the relief is clearly visible on the images: forests are found on the tops of slopes and rice and taro fields on lowlying lands. The deforested parts are small in area and scattered. This block illustrates that it is meaningless to consider only large areas of forest in the national inventory. This block is the most forested one both in area and in number of forest fragments. The Sikensi block north of Dabou. Forest cover changed from 182 km2 in 1958 to 149 km2 in 1990 to 17 km2 in 2000. Forests having an area less than 4 ha represent 64% of the forest area. This block was crossed by a road a long time ago and this resulted in the establishment of many farms. In addition, the most important oil palm plantations are located in the south of this region. Clearing in this block is high and large areas of cleared land belong to a single landholder. Many areas far from the main road were only recently deforested.

The Dogodou block. Forest cover changed from 210 km2 in 1958 to 139 km2 in 1990 to 46 km2 in 2000. Numerous migrants have established themselves along the coastal Abidjan-San Pedro road since it was improved in 1992. Coffee-cacao plantations, however, have been established to the northwest of this main road for a long time. The YocoboueTiegba main road also crosses the block and led to agricultural occupation. Deforestation is essentially in the interior of “forêts classées” (10% of the classified forest area). The zone along the road to Tiegba is a mosaic of forest and agricultural land.

(B) Blocks located in the western part of the area. Left the changes between 1959 and 1990, right the changes between 1990 and 2000 (for legends see Fig. 2.6).

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Appendix

Forest sites in Upper Guinea (Chapter 4)

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Appendix 2. Forest sites in Upper Guinea (Chapter 4)

The forest sites used for this study, along with some characteristics Minimum dbh limit is the minimum tree diameter at breast height that is used in the inventory data. Rainfall and altitude of sites are interpolated values of the location of the site (latitude plus longitude), see detailed explanation in chapter 9. DCA axis 1 and axis 2 scores are the results of a Detrended Correspondence Analysis using log-transformed abundance data. Forest types abbreviations. HW= hyperwet, WE= wet evergreen, ME= moist evergreen, MS= moist semi-deciduous, DS= dry semi-deciduous.

Part A. The 176 samples for which correct abundance data are available. Forest No.

Name

Minimum dbh limit (cm)

Inventoried area (ha)

Latitude (degrees)

Longitude (degrees)

Altitude (m asl)

Gola West Golama North Forest Reserve Koye (Gola North) Mahoi (Gola East) Mogbai (Gola North) Tiwai Island Tonkoli Forest Reserve, Kindea Section Gola East Wemago

60 60 40 40 40 40 60

156.9 766 8 8 8 8 39

7.44 8.92 7.63 7.37 7.65 7.55 8.37

– 11.3 – 11.38 – 10.95 – 11.2 – 10.87 – 11.35 – 11.18

154 424 170 100 300 100 292

2621 2673 2880 2980 2864 2560 2717

70.1 53.7 76.9 96.8 81.4 66 49.4

40

8

7.45

– 11.07

160

2996

Yoma School Forest Kpelle National Forest Krahn-Bassa National Forest SW North Gio National Forest Gio National Forest Gbi National Forest Grebo National Forest E Grebo National Forest W Grebo National Forest S Krahn-Bassa National Forest NE Krahn-Bassa National Forest NE Krahn-Bassa National Forest N Krahn-Bassa National Forest W Krahn-Bassa National Forest C Krahn-Bassa National Forest C Krahn-Bassa National Forest W Krahn-Bassa National Forest S Krahn-Bassa National Forest S Lorma National Forest North-Lorma National Forest Section1 North-Lorma National Forest Section 2 North-Lorma National Forest Section 3 Sapo National Park E Sapo National Park NW Sapo National Park S Lamco NW Concession Mt. Nimba

40 40 40 40 40 40 40 40 40 40 40 40 40 40 40 40 40 40 40

25.6 72 ? 88 180 288 468 384 420 1840 2250 2250 2600 2400 ? ? ? ? 36

6.93 7.32 5.59 6.7 6.4 6.2 5.62 5.54 5.31 5.93 5.91 5.91 5.91 5.7 5.7 5.67 5.57 5.39 7.29

– 10.77 – 10.45 – 9.21 – 8.83 – 8.4 – 8.73 – 7.46 – 7.72 – 7.62 – 8.36 – 8.54 – 8.64 – 8.82 – 8.77 –9 – 9.1 –9 – 9.13 – 9.64

133 304 51 305 247 142 112 143 157 420 224 262 152 152 160 99 139 143 303

40

64

7.99

– 9.65

40

92

8.02

40 40 40 40

36 128 52 80

40

FC Goin-Cavally SiteB, 1979 FC Haute Dodo FC Ht Sassandra transect 1 FC Ht Sassandra transect 2 FC Ht Sassandra transect 3 FC Ht Sassandra transect 4 FC Niégré transect 1 FC Niégré transect 2 FC Niégré transect 3

20 20 40 40 40 40 40 40 40

Extended Name

Rainfall (mm)

DCA (abundance)

Forest Type

Source

47.7 24.9 27.8 25.2 37.4 56.2 39.2

HW WE3 HW HW HW WE2 WE3

1 2 3 3 3 3 2

58.7

70.7

WE1

3

3206 2892 3043 1934 1947 2045 1952 2067 2118 2281 2383 2353 2397 2574 2681 2833 2912 3422 2063

91.2 84.6 88.4 63.7 58.2 72.8 52.3 62 60.8 75.9 72.8 76.2 70.3 77 79.7 81.2 89.2 100 74.5

35 32.7 38.7 44.9 41.4 44.8 47.5 56.7 48.3 51 53.5 57 49.9 48.7 48.5 39.7 37.3 45.4 38.8

HW HW HW WE3 WE3 HW WE2 WE2 WE2 HW HW HW HW HW HW HW HW HW HW

4 4 5 6 6 6 7 7 7 5 5 5 5 5 5 5 5 5 4

603

2440

56.7

28.1

WE3

4

– 9.82

463

2445

54.3

35.5

WE3

4

8.07 5.44 5.47 5.31

– 9.57 – 8.36 – 8.59 – 8.54

634 406 171 158

2495 2535 2638 2711

55.8 77.4 84.9 83.6

31.4 50.6 41.3 47.7

WE3 HW HW HW

4 5 5 5

132

7.58

– 8.47

758

2346

58.6

42.9

WE3

4

? ? 88.9 89.3 86.5 84.6 30 155 88

6.25 5 7.35 7.22 7.09 6.99 5.59 5.49 5.28

– 7.75 – 7.25 – 6.98 – 6.97 – 6.96 – 6.97 – 6.2 – 6.2 – 6.16

299 251 286 353 263 230 175 163 208

1852 2069 1427 1428 1454 1480 1553 1573 1648

45.5 53.7 23 26.6 26.7 34.4 49.1 51.8 57.3

19.1 37.6 9.6 8.1 1.6 22.2 61.7 60.6 68.4

WE3 WE3 DS DS DS DS WE2 WE2 WE1

8 8 8 8 8 8 8 8 8

Axis 1

Axis 2

Sierra Leone 198 201 194 197 193 200 196

Gola West Golama Koye Mahoi Mogbai Tiwai Tonkoli

195

Wemago

Liberia 192 191 186 181 172 178 159 165 161 170 174 177 180 179 182 184 183 185 188 189

L10_Yoma L101_Kpell L167_KraSW L20_N_Gio L30_Gio L40_Gbi L51_GreboE L52_GreboW L53_GreboS L61_KrahNE L62_KrahNE L63_KrahnN L64_KrahnW L65_KrahnC L66_KrahnC L67_KrahnW L68_KrahnS L69_KrahnS L70_Lorma L81_N_Lorm

190

L82_N-Lorm

187

L83_N-Lorm

171 176 175 173

L91_Sapo_E L92_SapoNW L93_Sapo_S Lamco

Côte d'Ivoire 166 158 154 151 150 152 136 137 135

494

Goin-Cav79 HauteDodo HtSasRec1 HtSasRec2 HtSasRec3 HtSasRec4 NiégréRec1 NiégréRec2 NiégréRec3

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Appendix 2. Forest sites in Upper Guinea (Chapter 4)

133 134 167 164 163 127 130 128 124 123 126 129 131 138 132 125 168 169 162 160 156 157 155 145 146 143 141 142

NiégréRec4 NiégréRec5 Scio_79 ScioRec1 ScioRec2 SODCE1 SODCE2 SODCE3 SODCS1 SODCS2 SODCS3 SODCS4 SODCS5 SODCS6 SODCS7 SODCS8 SODNW1 SODNW2 SODNW3 SODNW4 SODNW5 SODNW6 SODNW7 XV1 XV2 XV3 XV4 XV5

FC Niégré transect 4 FC Niégré transect 5 FC Scio SiteB 1979 FC Scio transect 1 FC Scio transect 2 CE sector 1 Divo CE sector 2 Gagnoa CE sector 3 Tene CS sector 1 Mopri CS sector 2 Dego-Bodi CS sector 3 Dogodou CS sector 4 Okromoudo CS sector 5 Niourouni CS sector 6 Soubre CS sector 7 Niegre CS sector 8 Guitri NW sector 1 Guiglo NW sector 2 Toulepleu NW sector 3 Goin NW sector 4 Ht Cavally NW sector 5 Nzo NW sector 6 Duekoue NW sector 7 Mt Peko Perimètre industriel N Perimètre industriel SW Perimètre industriel Centre Perimètre industriel E Perimètre industriel NE

40 40 20 40 40 20 /60 20 /60 20 /60 20 /60 20 /60 20 /60 20 /60 20 /60 20 /60 20 /60 20 20 20 20 20 20 20 20 20 20 20 20 20

43 46 730.4 100 85 1020.51 407.94 706.72 654.24 681.65 1198.61 1447.38 939.9 1182.08 944.75 720.97 4300.76 2894.8 3428.3 2693.2 2799 4592.02 1812.6 569.4 754.7 146.8 26.7 277.9

5.2 5.15 6.77 6.83 6.72 6.09 6.36 6.68 5.81 5.39 5.28 5.45 5.42 5.57 5.34 5.49 6.84 6.53 6.34 6.03 6.24 6.58 6.97 5.91 5.44 5.59 5.47 5.98

– 6.14 – 6.15 – 7.79 – 7.71 – 7.69 – 5.52 – 5.88 – 5.65 – 5.04 – 4.96 – 5.5 – 5.74 – 5.95 – 6.33 – 6.13 – 5.16 – 7.79 – 8.13 – 7.62 – 7.59 – 7.23 – 7.23 – 7.08 – 6.82 – 6.82 – 6.69 – 6.51 – 6.59

140 109 280 238 271 281 228 228 74 102 74 135 152 150 131 74 304 284 304 273 219 237 196 179 210 166 137 235

1689 1700 1789 1741 1748 1424 1421 1308 1472 1642 1663 1569 1597 1585 1629 1547 1772 1906 1783 1872 1715 1649 1545 1616 1778 1638 1642 1583

57.2 63 41.8 43 41.7 37.2 28.8 22.4 33.5 63.3 50.5 52.5 51.5 50.6 55.9 57.2 48.1 54.8 52.9 57.5 49.3 44.6 33.4 56.9 51.9 49 48.3 52.5

61.9 48.5 25.2 30.1 32.8 48.9 38.9 21.1 45.4 47.7 51.3 54.5 55.7 56.5 59.4 42.9 32.9 35.1 304 35.6 31.3 30.9 19 27.7 42 41.6 47.8 31.1

WE2 WE2 WE3 WE3 WE3 MS MS DS MS WE2 WE2 WE2 WE2 WE2 WE2 WE3 WE3 WE3 WE3 WE3 WE3 WE3 DS WE3 WE3 WE3 WE2 WE3

8 8 8 8 8 8 8 8 9 9 9 9 9 9 9 9 10 10 10 10 10 10 10 11 11 11 11 11

Aboniyere Afao Hills Afia Shelt Afram Head Afrensu-Br Aiyaola Amama Shel Angoben Sh Anhwiaso E Anhwiaso S Anum Su No Apamprama Aparapi Sh Asenanyo R Asubima Asukese Atewa Rang Awura Ayum Bandai Hil Bemu Ben East Ben West Bia Shelte Bia Tano Bia Tribututaries Nort Bimpong Birim Birim Exte Bobiri Boi Tano Boin River Bonkoni Bonsa Ben Bonsa Rive Bonsam Bep Bosumkese Bosumtwi R Bowiye Ran Bura River Cape Three Chirimfa

Aboniyere Afao Hills Afia Shelterbelt Afram Headwaters Afrensu-Brohuma Aiyaola Amama Shelterbelt Angoben Shelterbelt Anhwiaso East Anhwiaso South Anum Su North Apamprama Aparapi Shelterbelt Asenanyo River Asubima Asukese Atewa Range Awura Ayum Bandai Hills Bemu Ben East Ben West Bia Shelterbelt Bia Tano Bia Tribututaries North Bimpong Birim Birim Extension Bobiri Boi Tano Boin River Bonkoni Bonsa Ben Bonsa River Bonsam Bepo Bosumkese Bosumtwi Range Bowiye Range Bura River Cape Three Points Chirimfa

30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30

26 18 10 102 44 16 24 16 60 10 22 18 12 120 42 126 112 64 60 78 20 12 32 10 100 174 50 24 10 16 60 150 38 76 78 66 70 34 56 50 26 58

6.58 6.23 6.15 7.17 7.37 6.15 7.2 5.83 6.3 6.2 6.82 6.32 7.13 6.45 7.45 7.17 6.17 7.33 6.75 6.8 5.75 5.58 5.55 7.05 7 6.75 5.72 6.92 6.97 6.63 5.48 5.73 7.87 5.67 5.33 6.67 7.1 6.5 5.72 5.83 4.83 7.12

– 2.58 – 2.32 – 1.28 – 1.67 – 2.88 – 0.95 – 2.38 – 2.23 – 2.17 – 2.38 – 1.22 – 1.88 – 2.15 – 2.12 – 1.87 – 2.53 – 0.6 – 1.37 – 2.68 – 0.92 – 1.08 – 1.72 – 2.85 – 2.7 – 2.62 –3 – 1.47 – 1.15 – 1.13 – 1.28 – 1.65 – 2.93 – 2.63 – 1.75 – 1.85 – 2.62 – 2.25 – 1.42 – 2.05 – 2.3 – 2.05 – 1.28

229 311 152 304 304 152 301 176 145 153 301 152 256 159 327 292 422 239 229 233 148 171 76 233 251 229 145 241 165 202 72 230 307 230 76 278 304 152 153 152 75 227

1518 1786 1660 1430 1240 1635 1317 1878 1619 1673 1425 1623 1378 1546 1348 1298 1662 1441 1419 1571 1488 1611 1908 1322 1366 1377 1528 1480 1492 1530 1603 1740 1194 1661 1757 1447 1341 1411 1828 1857 1922 1521

22.7 33 24.3 23.7 24.9 32.5 24.4 35.5 30.3 23.7 18.9 33 14.2 31.7 28.2 26.8 41.8 26.6 29.4 0 34.8 33.9 42.7 16.1 29.3 38.4 37.5 29.9 31.4 28.5 62.6 41.2 23.8 38.2 53 26 26.2 28.8 34.4 40.3 61.1 18.9

51.1 67.6 70.6 39.4 40.2 72 35.9 69.6 61 51.1 38.3 61.9 31.1 62.9 36.1 30.9 67 7.4 36.2 24.3 63.4 78.4 85.2 0 37.8 42.6 56.7 60.3 59.5 81 79.3 64.8 32.4 73.8 67.3 41.8 32.2 86.7 73 72.3 65.3 23.7

MS ME MS MS MS ME DS ME MS MS DS ME DS MS MS DS WE1 DS MS DS ME ME WE1 DS MS MS ME MS MS ME WE1 WE1 DS ME WE1 MS DS ME ME ME WE1 DS

12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12

Ghana 92 80 27 42 110 7 84 75 72 85 23 54 70 69 52 91 2 33 100 6 12 46 108 101 94 118 38 17 16 28 41 114 97 48 51 95 76 36 64 79 65 29

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Appendix 2. Forest sites in Upper Guinea (Chapter 4)

Ghana (continued) 119 43 49 22 81 82 53 4 5 31 86 78 89 102 113 58 106 112 9 83 67 44 19 111 98 71 40 61 62 35 34 57 115 66 30 32 55 50 116 20 39 59 3 104 109 47 90 103 13 45 107 93 96 73 99 88 63 68 87 74 14 1 105

496

Dadiaso Dampia Ran Denyau She Dome River Draw River Ebi River Esen Epam Esuboni Esukawkaw Fum Headwa Fure Headw Fure River Goa Shelte Jema Assam Jeni River Kajeasi Krokosua H Kwamisa Mamang Riv Mamiri Mankrang Minta Mirasa Hil Mpameso Muro Ndumfri Nkrabia North Foma Nsuensa Numia Nyamibe Be Oboyow Oda River Ofin Shelt Onuem Bepo Onuem Nyamibe Shelterb Opon Mansi Opro River Pamu Berek Pra Anum Pra Suhien 1 South Foma Southern S Subim Subin Shel Subri Rive Suhuma Sui River Supong Supuma She Tain Tributaries Ii Tano Anwia Tano Nimir Tano Ofin Tano Suhie Tano Suraw Tinte Bepo Tonton Totua Shel Upper Wass Wawahi Worobong S Yoyo

Dadiaso Dampia Range Denyau Shelterbelt Dome River Draw River Ebi River Esen Epam Esuboni Esukawkaw Fum Headwaters Fure Headwaters Fure River Goa Shelterbelt Jema Assamkrom Jeni River Kajeasi Krokosua Hills Kwamisa Mamang River Mamiri Mankrang Minta Mirasa Hills Mpameso Muro Ndumfri Nkrabia North Fomangsu Nsuensa Numia Nyamibe Bepo Oboyow Oda River Ofin Shelterbelt Onuem Bepo Onuem Nyamibe Shelterbelt

30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30

70 34 12 38 114 14 24 24 58 36 82 98 12 32 14 14 230 44 24 26 40 12 34 172 32 36 52 20 28 30 12 30 80 28 16 10

5.97 6 6.9 6.53 5.2 5.12 5.82 6.83 6.37 6.25 5.53 5.37 7.98 5.43 6.5 6.4 6.5 7.13 6.27 5.67 7.35 5.8 6.37 7.08 6.48 5.17 6 6.68 6.15 6.03 6.17 5.77 6.13 6.67 6.05 6.12

– 3.03 – 1.67 – 2.98 – 1.2 – 2.33 – 2.37 – 1.87 – 0.8 – 0.8 – 1.35 – 2.38 – 2.28 – 2.47 – 2.73 – 2.92 – 1.95 – 2.82 – 2.9 – 1.05 – 2.37 – 2.07 – 1.67 – 1.17 – 2.88 – 2.63 – 2.15 – 1.58 – 1.98 – 1.98 – 1.4 – 1.37 – 1.92 – 2.93 – 2.05 – 1.3 – 1.35

152 151 229 369 75 74 277 117 162 387 142 76 304 71 162 232 209 230 194 137 306 172 304 229 221 75 151 188 218 124 228 95 152 201 175 153

1514 1627 1337 1545 2147 2309 1682 1606 1677 1704 1961 1863 1197 1817 1497 1560 1515 1262 1575 1996 1343 1656 1580 1283 1562 1962 1658 1450 1619 1686 1720 1710 1568 1440 1663 1688

47.3 29.6 26.8 24.7 59.6 56.3 31.2 27 34.8 30.5 56.8 59.9 22.8 58.3 28.3 25.9 29.3 20.4 25.9 44.2 25.1 34 29.4 26.4 22.9 59.6 31.6 28.3 33.8 33.6 29.4 31.3 36 25.1 27.2 30.5

43.6 86.4 56.5 60.1 68.7 88.8 58.8 79.8 58 76.5 71.1 68.7 27.8 77.5 39.1 52.1 42 38.2 72.2 86.1 34.2 56.5 71.2 23.2 71.2 69.4 62.4 57.5 76.9 76.2 79.9 63.1 51 46.6 84.6 100

WE3 ME MS MS WE1 WE1 MS ME ME ME WE1 WE1 DS WE1 MS MS MS DS ME WE1 DS MS ME DS MS WE1 MS MS ME ME ME MS MS MS ME ME

12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12

Opon Mansi Opro River Pamu Berekum Pra Anum Pra Suhien 1 South Fomangsu Southern Scarp Subim Subin Shelterbelt Subri River Suhuma Sui River Supong Supuma Shelterbelt Tain Tributaries II Tano Anwia Tano Nimiri Tano Ofin Tano Suhien Tano Suraw Tinte Bepo Tonton Totua Shelterbelt Upper Wassa W Wawahi Worobong South (Kwahu) Yoyo

30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30

62 58 84 60 50 14 56 114 12 284 178 164 18 12 262 72 102 204 44 38 62 70 36 50 20 60 114

5.75 7.18 7.42 6.25 5.27 6.58 6.55 6.78 6.05 5.28 5.97 6.17 5.6 6 7.58 5.83 5.63 6.67 6.33 6.25 6.97 6 5.92 6.12 5.67 6.47 5.92

– 1.88 – 1.8 – 2.93 – 1.17 – 1.53 – 1.95 – 0.67 – 2.78 – 2.87 – 1.72 – 2.48 – 2.73 – 1.1 – 1.7 – 2.83 – 2.58 – 2.62 – 2.17 – 2.63 – 2.42 –2 – 2.08 – 2.38 – 2.22 – 1.1 – 0.47 – 2.8

284 304 306 249 74 304 599 202 228 132 109 230 146 168 266 143 127 448 271 231 329 221 215 84 151 607 153

1677 1394 1233 1618 1268 1495 1683 1408 1605 1597 1687 1613 1363 1614 1205 1723 1819 1464 1578 1622 1398 1723 1756 1703 1407 1654 1684

37.2 22 19 27 42.8 21.8 27 29.2 30.2 49.3 32 34.7 32.3 29.5 20.2 40.4 48.8 33.4 30.6 30.6 29.9 34.1 35.7 29.1 32.3 34.1 41.6

78.3 34.5 16.5 69.2 69.8 50.9 72.3 33.9 50.3 68.8 62.9 58.7 62.4 75.4 24.6 65 72.7 52.1 58.2 62.9 42.7 55.6 77.5 65.4 61.1 73.3 54.7

ME DS DS MS WE1 MS ME MS MS WE1 ME ME ME ME DS ME WE1 MS MS MS MS MS ME MS MS ME WE2

12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12

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Appendix 2. Forest sites in Upper Guinea (Chapter 4)

Part B. The 38 samples with insufficient abundance data, small sizes, or presence / absence data only. Forest no.

Name

Extended Name

Minimum dbh limit

Inventoried

Latitude

8

7.47

10 10

DCA (abundance)

Altitude (m asl)

Rainfall (mm)

– 11.32

130

2539

42.5

46.5

WE2

5.39 6.46

– 4.04 – 3.46

156

2000 1397

38.4 24.9

47.3 37.8

MS MS

13 1

6.46

– 3.5

152

1382

24.9

37.8

MS

1

6.45 5.06 5.32 5 5.176 5.15 5.41 5.97 7.07 4.85 5.47

– 3.45 – 5.89 – 6.88 – 7.22 – 7.054 – 5.53 – 7.2 – 3.53 – 5.97 – 6.4 – 7.17

1450 1575

1675 1400 1700 2000

31.9 36.4 44.6 35.2 41 34.2 33.3 38.4 26.1 35.9 44.3

46.5 47 43.2 45.6 44.7 47.3 38.8 48.1 42.4 46.8 45

MS MS WE3 MS WE3 MS MS MS MS MS WE3

14 15 16 14 16 17 16 14 18 17 19

5.98 4.83

– 3.32 – 6.47

127

1625 1835

36.8 50.4

46 31.8

MS WE3

14 20

4.81

– 6.81

126

1957

45.1

30.6

WE3

20

4.95

– 6.88

169

1897

45.3

29.1

WE3

20

5.08

– 6.91

156

1886

45.1

30.6

WE3

20

5.15

– 6.97

225

1898

45.3

29.1

WE3

20

4.91

– 6.39

120

1792

45.1

30.6

WE3

20

4.8

– 6.9

137

1972

45.1

30.6

WE3

20

5.65 5.88

– 7.12 – 7.33

1900 1900

34.5 32.9

43.3 48

MS MS

21 19

5.82 5.75 5.75 6.12

– 3.3 – 4.13 – 3.58 – 7.4

1675 1629 1750 1700

37.2 52.5 37.7 33.2

45.9 30.2 46.1 43.3

MS WE3 MS MS

14 22 14 19

6.32 6.8 6.3 5.55 6.17 6.03 6 5.68 6.18 5.62 5.7 6.72 6.97 6.48

– 2.27 – 1.15 – 1.12 – 1.07 – 1.97 – 1.78 – 1.27 – 1.18 – 1.43 – 1.9 – 1.22 – 1.02 – 1.22 – 1.05

1668 1434 1576 1342 1619 1557 1635 1445 1763 1749 1520 1563 1501 1588

9.3 14.6 3.2 11.5 23.9 26.8 12.4 28.3 28.1 14.6 27.8 2.8 27.2 19.5 63.3 48.6

39 73.4 32.5 47.1 69.4 56.5 48.5 46.4 70.9 55.9 70.9 35.3 61.9 60.2 86.9 59.2

DS MS DS MS MS MS MS MS ME MS ME DS MS MS WE1 WE1

12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12

Longitude

Axis 1

Axis 2

Forest type

Source

Sierra Leone 199

Gola west2

Gola West 2

1

Côte d'Ivoire 216 120

Banco Bossemat 2

121

Bossematie

204 205 219 206 218 207 217 209 215 210 222

Bossematie Dassioko Djapadji Dodo Dogbo Gauthier Guiroutou Mabi Marahue Monogaga Para

211 140

Songan SPedro1

144

SPedro2

147

SPedro3

149

SPedro4

153

SPedro5

139

SPedroEx1

148

SPedroEx2

212 221

Taï Taï RvR

213 122 214 220

Tamin Yapo Yaya Zagne

Banco National Park ICFE Bossématié FC rapp 6 (Lennertz) IFCE Inv. forets de l'Est (Laumans) Bossématié Dassioko Djapadji Haute Dodo Dogbo Port Gauthier Guiroutou Mabi Marahue Monogaga Near Para village in Tai National Park Songan Perimètre Papetiere sect 1 Monogaga Perimètre Papetiere sect 2 San Pedro Perimètre Papetiere sect 3 Nero Perimètre Papetiere sect 4 Djakoteti Perimètre Papetiere sect 5 Palabod Perimètre Papetiere Extension sect 1 Hana Perimètre Papetiere Extension sect 2 Taï National Park Near Taï village in Taï National Park FC Tamin FC Yapo FC Yaya Near Zagne village in Taï National Park

2100 1575

76

Ghana 77 18 15 11 60 49 26 21 37 56 24 8 25 10 202 203

Anhwiaso N Anum Su So Auro River Baku Bediako Denyau She Kokotintin Krochua Kunsimua B Nkonto Ben Ochi Headwaters 2 Onyimsu Pra Birim Prakaw Neung Nort Neung Sout

Anhwiaso North Anum Su South Auro River Baku Bediako Denyau Shelterbelt Kokotintin Shelterbelt Krochua Kunsimua Bepo Nkonto Ben Ochi Headwaters 2 Onyimsu Pra Birim Prakaw Neung North Neung South

2 4 4 4 6 4 6 6 6 6 4 4 8 6 12 60

152 380 144 97 227 152 110 138 271 151 149 398 250 224

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Appendix 2. Forest sites in Upper Guinea (Chapter 4)

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22

498

Small 1953 Savill & Fox 1967 Davies 1987 Sachtler & Hamer 1967a Sachtler & Hamer 1967b GFML 1967a GFML 1967b SODEFOR unpublished SODEFOR 1978 SODEFOR 1979 Clément & Guinaudeau (1973) Hawthorne 1995a, 1996, Hawthorne & Abu Juam 1995 De Koning 1983 Kouame et al. chapter 5 Aké Assi 1997 Adou et al. unpubl.data Kouamé 1998b Jongkind et al. 1999 Van Rompaey 1993 Clément 1973 Aké Assi & Pfeffer 1975 SODEFOR 1986

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3

3

Appendix

Forest reserves in Upper Guinea

P P E N D I X

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A

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This map shows the location of National Parks and of Forest Reserves in the forest zone of Upper Guinea. The colours indicate the interpolated results of the vegetation classification of Appendix 2.

499

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Hotspots of plant diversity in Upper Guinean forests

Plant diversity hotspots are based on the distribution of a selection of rare and endemic plant species in Upper Guinean forests. The selection and the methods are treated extensively in chapter 6 (Wieringa & Poorter). The three maps here are the full colour versions of Figure 6.7. Dots are the middle of half-degree cells. For cells that partly cover the sea, the midpoint is positioned on the land surface. The sizes of the symbols are scaled to its biodiversity value. Cells with open symbols have

A

4

4

Appendix

P P E N D I X

22466 layout page 501-502

less than eight collections. Cells for which curve fitting was impossible (N = Sobs, see chapter 6) or that were considered to be too unreliable to use (N / Sobs < 1.1 see Box 6.1) were not included in the interpolation analysis, and are indicated by a cross. The background colour indicates the interpolated values for the whole potential forest zone of Upper Guinea: the darker the colour, the higher the biodiversity value.

N/S < 1.1

0 - 15

0 - 15

15 - 30

15 - 30

30 - 35

30 - 35

35 - 37.5

35 - 37.5

37.5 - 40

37.5 - 40

40 - 42.5

40 - 42.5

42.5 - 45

42.5 - 45

No data

N/S < 1.1

0 - 50

0 - 50

50 - 100

50 - 100

100 - 150

100 - 150

150 - 200

150 - 200

200 - 250

200 - 250

250 - 300

250 - 300

300 - 350

300 - 350

No data

A S50

350 - 400

B Smax

400 - 470

N/S < 1.1

N/S < 1.1

0-1

0-1

1-2

1-2

2-3

2-3

3-4

3-4

4-5

4-5

5-6

5-6

6-7 7-8 8-9

C Srw

9 - 10 10 - 11,5

6-7 7-8 8-9 9 - 10 10 - 11,5

Figure A shows the number of species (S) at 50 collections, B shows the maximum estimated number of rare and endemic species (Smax), and C gives the rarity-weighted species richness (Srw).

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Appendix

5

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Geographical map of West Africa

5 P P E N D I X

503

A

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D Sources & index

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Authors, photographers and illustrators

Authors

Beligné, V. – B.P. 708, Yamoussoukro, Côte d’Ivoire Bongers, F. – Forest Ecology and Forest Management Group, Department of Environmental Sciences, Wageningen University, PO Box 342, 6700 AH Wageningen, The Netherlands Buitelaar, M. – Biosystematics Group, Department of Plant Sciences, Wageningen University, PO Box 9101, 6700 HB Wageningen, The Netherlands Chatelain, C. – Conservatoire et Jardin botaniques de la Ville de Genève, 1 ch. de l’Impératrice, Case Postale 60, CH-1292 Chambésy/Genève, Switzerland Dao, H. – Université de Genève, Departement de Geographie, Boulevard du Pont d’Arve 40, CH-1211 Genève 4, Switzerland Gautier, L. – Conservatoire et Jardin botaniques de la Ville de Genève, 1 ch. de l’Impératrice, Case Postale 60, CH-1292 Chambésy/Genève, Switzerland Hawthorne, W.D. – 7 Poplar Road, Botley, Oxford, OX2 9LA, United Kingdom Helmink, A.T.F. – Alterra, PO Box 47, 6700 AA Wageningen, The Netherlands Holmgren, M. – Forest Ecology and Forest Management Group, Department of Environmental Sciences, Wageningen University, PO Box 342, 6700 AH Wageningen, The Netherlands Jongkind, C.C.H. – Biosystematics Group, Department of Plant Sciences, Wageningen University, PO Box 9101, 6700 HB Wageningen, The Netherlands Kouadio, K.E. – Laboratoire de Botanique, Université de Cocody 22, B.P. 582, Abidjan 22, Côte d’Ivoire Kouamé, F.N’. – Laboratoire de Botanique, Université de Cocody 22, B.P. 582, Abidjan 22, Côte d’Ivoire Kouassi, K. – Laboratoire de Botanique, Université de Cocody 22, B.P. 582, Abidjan 22, Côte d’Ivoire Lemmens, R.H.M.J. – Prosea-project, PO Box 9101, 6700 HB Wageningen, The Netherlands Os-Breijer, H.J. – Postbus 59, 6700 AB Wageningen, The Netherlands Parren, M.P.E. – Forest Ecology and Forest Management Group, Department of Environmental Sciences, Wageningen University, PO Box 342, 6700 AH Wageningen, The Netherlands Poorter, L. – Forest Ecology and Forest Management Group, Department of Environmental Sciences, Wageningen University, PO Box 342, 6700 AH Wageningen, The Netherlands Siepel, A. – Forest Ecology and Forest Management Group, Department of Environmental Sciences, Wageningen University, PO Box 342, 6700 AH Wageningen, The Netherlands Spichiger, R. – Conservatoire et Jardin botaniques de la Ville de Genève, 1 ch. de l’Impératrice, Case Postale 60, CH-1292 Chambésy/Genève, Switzerland Traoré, D. – Laboratoire de Botanique, Université de Cocody 22, B.P. 582, Abidjan 22, Côte d’Ivoire Van Zoest, A.R. – Laboratorium voor Geo-informatiekunde en Remote Sensing, PO Box 47, 6700 AH Wageningen, The Netherlands Wieringa, J.J. – Biosystematics Group, Department of Plant Sciences, Wageningen University, PO Box 9101, 6700 HB Wageningen, The Netherlands

Gautier, L. – all photos in chapter 3 Hawthorne, W.H. – 61, 101, 110, 133-134, 150-151, 158, 161-162, 167, 169, 175, 180, 184, 190, 206, 232, 247, 265, 280, 308, 325, 327, 347, 357, 361, 376, 384, 388, 395, 398, 408, 418, 429, 445 Kouamé, F.N. – 391 Leeuwenberg, A.J.M. – 137, 286, 350, 447 Linschoten – 73, 171 Mugge, J. – 130, 139, 216, 226, 352 Poorter, L. – 5, 41, 94, 96 Van der Burg, W.J. – 192, 243, 304 Van der Maesen, L.J.G. – 122, 141, 240, 402, 419, 425, 432, 439, 441 Voorhoeve, A.G. – 407 Wieringa, J.J. – 224, 275, 285

Illustrators

Aquarel paintings by E. Jaggar-Loffers – 136, 138, 160, 162, 178, 186, 222, 225, 235, 279, 285 Allard, M. – 258, 259 De Vries, H. – 266, 321 Spitteler, M. – 110-111, 115-120, 134, 137, 155, 160, 170, 176, 178, 180-181, 187-188, 200, 202-204, 237, 245-246, 248, 252, 255, 260-263, 265, 273274, 287-288, 290, 294-296, 306, 313, 316, 324-325, 334-337, 345, 349, 354, 359, 372, 377-379, 381 Tan, Y.F. – 220 Van der Riet, L. – 131-132, 350-351, 353 V.d. Burg, L. – 339 Wessel, W. – 141-142, 144, 215-219 Williamson, J. – 309 Wise, R. – 121, 124-125, 127-129, 133, 138, 146-148, 151, 156-157, 161, 163166, 168-169, 171-175, 183, 185, 189, 191, 193-195, 197-198, 201, 206210, 221, 226-231, 239, 241, 247, 251, 257, 264, 271, 280, 291, 297, 299300, 302, 307, 311-312, 322-323, 330, 338, 340-341, 343-344, 347-348, 355-357, 363, 367-368, 373-376, 380, 384, 386-389 and all drawings in chapter 10 Zewald, I. – 130, 139-140, 205, 242, 293, 350-352, 360-361, 366

Photographers

Antheunisse, M. – 219 Archives Biosystematics Group, WUR – 139, 153, 267, 403, 444 Bongers, F. – 53 Bos, J.J. – 220, 274, 401, 409, 413 Breteler, F.J. – 203, 204, 428 Culta ORSTOM – 356 De Wit, H.C.D. – 116, 120, 127, 144, 149, 156-157, 159, 173, 175, 179, 198, 201, 210, 217-218, 242, 247, 254-255, 262, 290, 294, 300, 305, 309, 312, 317, 323, 330, 339, 343, 345, 348, 354, 362, 368, 374, 380, 396, 399, 414, 427, 435, 436, 442

517

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Keywords

Abidjan 8, 11, 17, 22-23, 26-28, 30, 32, 66, 80, 84-85, 91-92, 94, 98 afforestation 12, 35, 38-40, 89, 90 altitude 41-49, 53, 61, 64, 66, 68, 74-77, 80, 82, 91, 102-103, 106-107, 392-393 Ankasa Forest Reserve 66, 71, 84, 91-93, 96 Atewa Range 7, 46, 48, 66, 69, 85, 91, 92-94, 96 Azagny 23, 94 Banco Forest 23, 56, 58, 64, 66, 85, 92 Bandai Hills 45 Benin 6, 11, 33, 39, 58, 61 Bia Shelterbelt 45, 66, 85, 89, 93 biodiversity 5, 12-14, 24, 49, 61-71, 85, 87, 90-98 Birrimian basement 36 Bossematié 22, 89 Cape Palmas 8, 12, 61, 66-68, 71, 83, 84, 98 Cape Three Points 8, 12, 61, 66, 68, 71, 83, 84, 92, 96, 98 catena 36, 48, 71 cattle 39, 89 Cavally 7, 20, 22 Cavally river 53 classification 12, 17-20, 22, 29, 34, 41, 43, 46, 50-52, 75, 77, 93, 96-98, 103-104, 106 clay 9, 36, 38, 44, 53, 59, 64 climate 7, 11-13, 17, 28, 33, 35-41, 50, 59, 70, 73-74, 83-85, 87, 89 climate change 26, 28, 74, 87, 92 CMK 43, 66, 75, 102, 392-393 coastal savanna 34, 104 cola-nuts 17, 29 commonness 75-77, 79-82, 103-107 Comoé National Parc 7, 20, 21, 66, 70, 94 conservation 5, 6, 12-13, 15, 20, 24, 26, 31-32, 61, 65, 69-70, 73, 84-85, 87-98, 101, 106 continental disjunct 59, 75-77, 103, 106 continuous distribution 75, 77, 79-80, 91, 103, 106 Côte d'Ivoire 5-9, 11-12, 15-34, 36-43, 45-51, 53-59, 61-62, 66 , 68, 70, 74, 78-79, 81, 83-85, 87-95, 97-98, 102, 107, 391-392 Dabou 18, 22, 30, 34 Dahomey gap 5, 8, 12-13, 35, 61, 70-71, 81, 89-90 Dassioko 56 deciduous 10, 12, 15, 17, 20, 23-24, 26, 31-32, 37, 41, 46-47, 49-51, 53, 56-59, 84-85, 88-90, 92-93, 95, 97, 104-107, 393 deforestation 6, 12, 15, 17, 20, 22-33, 38-39, 68, 73-74, 83, 87-91, 98 disjunct distribution 13, 35, 59, 62, 64, 71, 75, 77-78, 80-83, 91, 103, 106 dispersal 36, 38, 71, 74, 77, 81-82, 84, 87, 89, 91, 95-96, 105, 107 dispersal mechanism 77, 81 distribution range 6, 13, 64, 68, 71, 73-74, 77, 79-80, 83, 101 Divo 22-24, 26-27, 30 drought tolerance 71, 82 edaphic conditions 12, 36, 39-40, 74 endemism 5, 33, 64, 68-69, 75, 78, 95, 101-107, 447 evergreen 10, 17, 20, 23, 26, 33, 37, 41, 46-53, 58, 69, 84, 93-98, 104-107 fallow 18, 26, 31, 38-40, 89-90, 94, 98 farming 12, 17, 30-31, 38, 90-91, 94 fire 12, 26, 33-34, 36-40, 89-90, 98 floristic composition 33, 48, 53 floristic gradient 48 forest cover 12, 15, 17-31, 38, 53, 58, 73, 84, 88, 90, 93, 98, 102, 105, 393 forest edge 34, 36, 38-39, 85, 90 forest reserves 13, 15, 29, 56, 59, 66, 71, 75, 85, 87-88, 93-95, 98, 103 forest types 10, 13, 43, 46-53, 58, 68, 73, 82, 84, 87, 89, 93-98, 104, 106, 393 forest-savanna boundary 70 Fouta Djalon 7, 62, 66, 68, 85, 92, 96, 447 fragmentation 6, 15, 17, 19, 22, 24, 26, 74, 87, 95 gaps 77, 83, 85, 89, 104 Ghana 1, 5-12, 14-15, 19-20, 26, 35-38, 41-50, 52-53, 58, 61-62, 66, 69-71, 74-78, 81-98, 101-107, 391-393 Ghanaian species 53, 392 glacial period 12, 53, 59, 62, 70-71, 73, 83, 91

Gola Forest 68, 96 Grand Bassam 29, 34 Grand Lahou 22, 26, 34 granite 8, 9, 53, 57 grass 18, 33-34, 36-38, 41, 69, 71, 90 guild 77, 81, 104, 105 Guinea 5-7, 20, 35, 38, 58, 62, 66, 68, 70, 73-74, 85, 91-93, 96-97, 102, 447 Guineo-Congolian 5, 33-35, 37, 58, 69, 78, 90 Guinean savanna 19, 33, 35, 37, 58, 89 habitat openness 76-77, 80, 104, 107 Haute Dodo 22, 53, 55-58, 84, 96, 97 Haut Sassandra 22, 29, 48, 56, 66, 70, 84, 92, 97 herb 10, 11, 33, 36-39, 41, 48, 62, 70, 74, 77, 80-81, 90, 101, 105-106, 447 hotspots 5, 13, 61-62, 66, 85, 87, 90-91, 93, 96, 98 Krahn Bassa 45, 48, 84, 94, 96 Lake Bosumtwi 35 latitude 33, 35, 43-44, 47-48, 55-57, 63, 66, 69 liana 14, 41, 48, 62, 70, 74, 77, 80, 87, 101, 106, 447 Liberia 8-11, 20, 26, 31, 35, 42-43, 45-46, 48-49, 51-52, 58, 61, 66, 68, 70, 75, 7781, 83-85, 88-89, 91-98, 103, 391-392 life form 41, 62, 70, 81, 83, 101-107 light availability 13, 73, 104-106, 392 longitude 43-44, 47, 55-56, 63, 66 lower guinea 5, 12-13, 62, 64, 71, 74-78, 102-103, 105-106 Mabi 20, 22, 52, 55-56, 59, 85, 92-93 Marahoué 56, 93-94 Monogaga 56 Mount Nimba 12, 41, 51, 61, 66, 68, 71, 79, 83-84, 92, 96, 98 Mount Ziama 7, 62, 66, 69, 85, 91-92 near endemic 77 neighbour distance 75, 77, 103 Nigeria 11, 17, 20, 33, 36, 38, 51, 74, 102, 106 non-pioneer light demander 77, 81, 104-105 ordination 43-44, 46, 48, 52, 55, 58 palynology 35 pioneer 24, 36-39, 77, 81, 90-91, 104, 105, 447 plantation 17-18, 20, 22-24, 26, 28-32, 61, 88, 94-95, 102, 391 population density 28, 30, 38 Port Gauthier 56 primary forest 15, 19, 24, 27, 31, 94, 104-105 protected areas 15, 20, 22, 26, 31-32, 84, 90, 93-95 Putu Range 9, 84, 92, 96 Quaternary 9, 12, 34, 39, 62 rainfall 7-10, 12, 36, 41, 43-44, 46-59, 61-62, 64, 66-71, 73-77, 80-84, 87, 91-92, 102-104, 107, 392-393, 447 rarity-weighted species richness 65, 68 refuge 12, 13, 35, 58-59, 62, 64, 66, 68, 71, 83-84, 91, 98 rivers 7, 18-19, 74-75, 84-85, 89, 92, 102-107 ruderal 37, 82-83 Sassandra river 53, 84, 92 savanna 5-7, 10-12, 15, 18-22, 24, 28, 33-41, 46-47, 53, 58, 61-62, 70, 73, 83, 85, 89-90, 94, 98, 104, 106, 447 schist 8, 9, 36, 53, 57 secondary 15, 17-19, 22-27, 36-39, 89-90, 102-106 Senegal 5-7, 11, 14, 17, 29, 34, 41, 58, 61-62, 66, 69, 74-75, 77, 101-103, 106 shade bearer 77, 81, 104-105 shade tolerance 77, 81-82, 91, 104-105, 447 shifting cultivation 36, 38, 61, 89, 90 shrub 14, 36-37, 41, 48, 62, 70, 74, 77, 80, 90, 101, 104-106, 447 Sierra Leone 5-9, 11, 19, 20, 31, 34, 42-43, 45-46, 49, 51-52, 58, 66, 68-70, 78, 85, 88, 91-93, 96, 98, 391-392 soil fertility 36, 41, 43, 47, 61, 64, 67, 70-75, 81-82, 87, 90-91, 102, 392 Songan 22, 53, 55, 56, 59, 85, 92, 93 spatial distribution 74, 82, 91 species inventories 12, 41-43, 46, 49, 53, 55, 62, 70, 74, 95, 97, 101, 107, 391-393 species responses 44, 47, 103-104, 392 species richness 12, 58, 61, 63-66, 68, 70-71, 82, 84, 92-93 Sudanian 33-34 Tabou 53, 64, 66, 68 Taï National park 9, 15, 20-22, 26-27, 29, 31, 41, 53, 56, 64, 66, 84, 88, 92-94, 96 Tamin 22, 53, 55-56, 59, 85, 92-93 tertiary sands 9, 53, 56-57, 59, 85 timber 13, 29, 41-43, 87-88, 94, 97-98, 391-392

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Togo 5-7, 11, 20, 33, 38, 41, 58, 61-62, 69, 73-75, 77, 90, 96-97, 101, 103, 106 toposequence 33, 36 Upper Guinea disjunct 75, 77-78, 106 V-Baoulé 12-13, 17, 28, 33-39, 53, 83, 89-90 vegetation map 12, 17, 19, 41, 49-50, 52, 102, 105, 107, 392-393 vicariance 71, 84 Voltaian rocks 36 water availability 36, 40, 48, 58-59, 64, 70, 74, 81-82, 84, 91, 102, 105, 392 water deficit 36, 48, 53, 55-59 water holding capacity 9, 43-44, 47-48, 58, 64, 70, 74-75, 80, 102-103, 107, 392393 Yapo 20, 22-27, 29, 31, 66 Yaya 20, 22, 53, 55-56, 59, 85, 92-93 Zambesian 34

Species

Acanthus guineensis 77, 78, 109 Acridocarpus alternifolius 110 Acridocarpus plagiopterus 111 Acroceras gabunense 56 Adhatoda robusta 112 Afraegle paniculata 70 Aframomum alboviolaceum 56 Aframomum atewae 70, 113 Afrobrunnichia 58 Afzelia africana 42 Afzelia bella 42 Afzelia parviflora 114 Afzelia 43, 47, 391 Aganope leucobotrya 56 Agelaea paradoxa 56 Agelaea pentagyna 56 Aidia genipiflora 56 Alafia parciflora 81, 115 Albertisia cordifolia 81, 116 Albertisia cuneata 117 Albertisia ferruginea 118 Albertisia mangenotii 119 Albertisia scandens 69, 120 Albizia ferruginea 395 Alsodeiopsis chippii 81, 121 Alsophila manniana 84 Alstonia boonei 42, 56, 396 Amorphophallus baumannii 122 Amphimas 47, 58, 397 Amphimas pterocarpoides 42, 47, 56, 397 Ancistrocladus barteri 56, 59 Ancistrocladus pachyrrachis 80, 82, 123 Aneilema umbrosum 56 Angraecum podochiloides 56 Anisophyllea laurina 78, 124 Anisophyllea meniaudii 78, 125 Anisotes guineensis 126 Anogeissis leiocarpa 447 Anopyxis 58, 398 Anopyxis klaineana 42, 398 Anthocleista nobilis 127 Anthoclitandra nitida 56 Anthonotha 58, 128-129, 399 Anthonotha explicans 128 Anthonotha fragrans 42, 399 Anthonotha vignei 129 Antiaris toxicaria 10, 42, 400 Antrocaryon 58 Anubias afzelii 130

520

Aphanocalyx 107 Aphanocalyx microphyllus ssp. compactus 71, 131 Aphanocalyx pteridophyllus 132 Apodiscus chevalieri 133 Apodiscus 6 Argocoffeopsis afzelii 134 Argocoffeopsis lemblinii 80, 82, 135 Asystasia scandens 136 Aubrevillea 50, 51, 58 Aubrevillea kerstingii 50, 51 Avicennia africana 11 Baissea zygodioides 137 Baphia nitida 56 Baphia pubescens 56 Baphia spathacea ssp. spathacea 138 Begonia cavallyensis 69, 139 Begonia fusicarpa 80, 82, 140 Begonia hirsutula 71, 78, 141 Begonia mildbraedii 62, 78, 142 Begonia prismatocarpa ssp. petraea 82, 143 Begonia quadrialata ssp. nimbaensis 81, 144 Beilschmiedia caudata 81, 145 Beilschmiedia chevalieri 146 Berlinia bracteosa Berlinia confusa 42, 401 Berlinia occidentalis 42, 71, 147 Berlinia tomentella 148 Berlinia 43, 47, 391 Bertiera fimbriata 56 Bertiera spicata 149 Bolbitis heudelotii 56 Bonamia vignei 150 Borassus aethiopium 37 Brachystegia leonensis 151 Bridelia atroviridis 56 Brieya fasciculate 56 Buchholzia 58 Buchholzia coriacea 56 Buforrestia mannii 56 Buforrestia obovata 152 Bulbophyllum imbricatum 56 Bussea occidentalis 56, 153 Byrsanthus brownii 154 Byttneria 155 Byttneria dahomensis 82, 155 Byttneria guineensis 82, 155 Byttneria ivorensis 82, 155 Callichilia subsessilis 156

Calpocalyx 58 Calpocalyx aubrevillei 157 Calpocalyx brevibracteatus 158 Calvoa monticola 78, 159 Calycobolus africanus 56 Calycobolus insignis 160 Campylospermum amplectens 161 Campylospermum subcordatum 162 Canarium schweinfurthii 42, 402 Cassipourea afzelii 78, 163 Cassipourea hiotou 78, 164 Cathormion rhombifolium 165 Cavacoa baldwinii 166 Cecropia peltata 56 Ceiba pentandra 42, 403, 447 Celtis adolfi-friderici 42, 404 Celtis mildbraedii 42, 405 Celtis 43, 48, 51, 391, 393, 404405 Cercestis ivorensis 56 Chassalia corallifera 167 Chazaliella cupulicalyx 168 Chidlowia 58 Chromolaena odorata 26, 39 Chrysophyllum giganteum 406 Chytranthus cauliflorus 78, 169 Clappertonia minor 56 Clerodendrum polycephalum 56 Clerodendrum sassandrense 80-82, 170 Cnestis ferruginea 56 Coelocaryon 58 Cola attiensis 78, 171 Cola boxiana 172 Cola buntingii 84 Cola caricifolia 173 Cola nitida 56 Cola reticulata 174 Cola umbratilis 81, 84, 175 Combretum 176-181 Combretum bipindense 78, 176 Combretum blepharopetalum 177 Combretum calobotrys 178 Combretum grandiflorum 77, 179 Combretum paniculatum 447 Combretum tarquense 180 Combretum zenkeri 181 Commelina macrosperma 182 Commiphora dalzielii 183 Copaifera salikounda 184 Cordia vignei 185 Costus afer 56 Costus deistelii 186 Costus englerianus 56 Coula 58 Craterispermum caudatum 56 Crossostemma laurifolium 187 Crotonogyne 58 Crotonogyne craterviflora 56 Culcasia barombensis 56 Culcasia glandulosa 188 Cussonia bancoensis 189 Cyclodiscus 58 Cynometra ananta 10, 190 Cynometra leonensis 191 Cyrtococcum chaetophoron 56 Cyrtorchis hamata 192 Dactyladenia dinklagei 193 Dactyladenia smeathmannii 194 Dactyladenia whytei 195 Dalbergia albiflora 56 Daniellia ogea 42, 407 Daniellia thurifera 42, 408 Decorsella 58

Deinbollia voltensis 196 Delpydora gracilis 56 Dennettia tripetala 197 Desmodium adscendens var. Robustum 56 Dialium aubrevillei 198 Diaphananthe suborbicularis 82, 199 Dichapetalum Dichapetalum albidum 200 Dichapetalum barteri 201 Dichapetalum dictyospermum 81, 202 Dichapetalum filicaule 203 Dichapetalum madagascariense var. Madagascariense 56 Dichapetalum toxicarium 204 Didelotia 58 Didelotia brevipaniculata 56 Didelotia engleri 205 Didelotia idea 206 Dinklageodoxa 6 Diospyros abyssinica 56 Diospyros chevalieri 207 Diospyros cooperi 208 Diospyros liberiensis 209 Diospyros soubreana 56 Diospyros vignei 210 Discoglypremna 58 Dissomeria crenata 211 Distemonanthus 58, 409 Distemonanthus benthamianus 42, 409 Dorstenia astyanactis 212 Dorstenia embergeri 213 Dorstenia turbinata 78, 214 Dracaena calocephala 215 Dracaena cristula 216 Dracaena ovata 78, 217 Dracaena phrynioides 218 Dracaena praetermissa 219 Dracaena scabra 220 Drypetes klainei 56 Duboscia 58, 89 Ehretia trachyphylla 221 Elytraria ivorensis 222 Elytraria maritime 223 Englerina gabonensis 78, 224 Englerina parviflora 225 Entandrophragma angolense 42, 410 Entandrophragma candollei 42, 411 Entandrophragma cylindricum 42, 412 Entandrophragma utile 42, 413 Entandrophragma 43, 391, 393, 410-413 Erythrophleum guianense 42 Erythrophleum ivorense 42, 414 Erythrophleum suaveolens 42, 415 Erythrophleum 43, 391, 393, 414415 Euadenia eminens 78, 226 Eugenia tabouensis 56 Eugenia whytei 56 Eupatorium microstemon Ficus 56 Ficus ottoniifolia ssp. multinervia 227 Ficus scott-elliotii 228 Friesodielsia enghiana 56 Funtumia africana 42, 56, 416 Garcinia elliotii 229 Garcinia granulata 56 Gardenia nitida 230 Gilbertiodendron bilineatum 231

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Gilbertiodendron limba 232 Gilbertiodendron obliquum 82, 233 Gilbertiodendron preussii 10, 42, 46, 417 Gilbertiodendron robynsianum 56, 81, 234 Gilbertiodendron 58, 229-234, 417 Globimetula assiana 235 Globimetula cupulata 236 Glyphaea brevis 56 Grewia carpinifolia 56 Griffonia simplicifolia 56 Grossera 58 Guaduella macrostachys 78, 237 Guaduella oblonga 78, 238 Guarea cedrata 42, 418 Guibourtia 59, 239, 419 Guibourtia copallifera 56 Guibourtia dinklagei 239 Guibourtia ehie 42, 46, 419 Guibourtia tessmannii 56 Gymnosiphon longistylus 77, 78, 240 Gymnostemon zaizou 241 Gynura sarmentosa 56 Habropetalum 6 Hallea ledermannii 420 Heckeldora 58 Hemandradenia chevalieri 71, 78, 242 Heritiera utilis 42, 48, 243 Heterostis rotundifolia 56 Hibiscus whytei 82, 244 Hippocratea vignei 245 Hugonia rufipilis 246 Hunteria ghanensis 70, 92, 247 Hutchinsonia barbata 248 Hutchinsonia glabrescens 249 Hymenocoleus axillaris 78, 82, 250 Hymenostegia 58, 251 Hymenostegia gracilipes 251 Illigera vespertilio 78, 252 Impatiens nzoana 253 Impatiens nzoana ssp. bennae 82, 253 Impatiens nzoana ssp. nzoana 79, 81, 253 Isolona cooperi 254 Isonema smeathmannii 255 Ixora 107 Ixora hiernii 256 Ixora laxiflora 257 Ixora liberiensis 81, 82, 258 Ixora tenuis 259 Keetia bridsoniae 260 Keetia obovata 82, 261 Khaya anthotheca 42, 50, 421 Khaya grandifoliola 42, 51, 56, 59, 422 Khaya ivorensis 42, 51, 423 Khaya 43, 391, 393, 421-423 Klainedoxa gabonensis 42, 56, 424 Lagenaria breviflora 56 Landolphia hirsuta 56 Landolphia landolphioides 56 Landolphia micrantha 262 Landolphia owariensis 56 Landolphia togolana 263 Lasiodiscus mannii 78, 264 Leptoderris miegei 265 Leucomphalos libericus 82, 266 Licania elaeosperma 56 Loesenera kalantha 267 Lomariopsis rossii 56 Lophira alata 42, 51, 425

Lophira lanceolata 10 Lovoa trichilioides 42, 426 Macropodiella garrettii 268 Maesa nuda 269 Magnistipula cupheiflora ssp. leonensis 270 Magnistipula zenkeri 78, 271 Malaxis melanotoessa 82, 272 Mammea africana 42, 427 Mangenotia eburnea 273 Manotes macrantha 78, 274 Mansonia altissima 51 Mapania baldwinii 275 Mapania ivorensis 276 Mapania linderi 277 Mapania minor 56, 59, 278 Mapania rhynchocarpa 78, 279 Maranthes aubrevillei 280 Marantochloa cuspidate 281 Marantochloa filipes 56 Marattia odontosora 78, 282 Maschalocephalus dinklagei 283 Melochia melissifolia 56 Memecylon aylmeri 78, 284 Memecylon polyanthemos 56 Mendoncia combretoides 285 Microdesmis keayana 56 Milicia excelsa 42, 428 Milicia regia 42, 286 Milicia 43, 286, 391, 393, 428 Millettia leonensis 81, 82, 287 Millettia liberica 288 Millettia lucens 56 Millettia pallens 289 Mischogyne elliotianum var. glabra 56 Mitragyna 11 Monanthotaxis whytei 290 Monocyclanthus 58, 291 Monocyclanthus vignei 78, 291 Monosalpinx guillaumettii 292 Mostuea hymenocardioides 293 Mussaenda afzelii 294 Mussaenda grandiflora 295 Mussaenda landolphioides 56 Mussaenda tristigmatica 296 Myrianthus arboreus 56 Myrianthus libericus 56, 297 Napoleonaea heudelotii 298 Napoleonaea vogelii 56 Nauclea diderrichii 42, 429 Neostenanthera hamata 299 Nephrolepis bisserrata 56 Nesogordonia papaverifera 42, 46, 47, 50, 51, 430 Neuropeltis acuminata 56 Newtonia aubrevillei 300 Newtonia elliotii 301 Ochtocosmus africanus 56 Okoubaka aubrevillei 78, 302 Olea hochstetteri 35 Oleandra ejurana 303 Omphalocarpum elatum Oncoba brevipes 304 Ophiobotrys 58 Palisota hirsuta 56 Pararistolochia goldieana 305 Pararistolochia mannii 306 Parinari excelsa 10, 42, 51, 56, 89, 431 Parinari 43, 391 Parkia bicolor 432 Pauridiantha hirtella 56 Pavetta akeassii 307

Pavetta micheliana 308 Penianthus patulinervis 309 Pennisetum purpureum 38, 90 Pericopsis elata 50, 433 Petersianthus macrocarpus 42, 47, 50, 434 Phragmanthera vignei 310 Pierreodendron kerstingii 78, 311 Piptadeniastrum africanum 42, 47, 56, 435 Piptostigma fugax 312 Platysepalum hirsutum 313 Pleiocoryne fernandensis 314 Podocarpus 35 Poecilocalyx stipulosa 315 Polycephalium capitatum 316 Polystachya bancoensis 317 Polystachya dalzielii 318 Polystachya pseudodisa 319 Polystemonanthus dinklagei 56 Polystemonanthus 6 Premna grandifolia 56, 82, 320 Pseudocalyx libericus 81, 82, 321 Psychotria 56 Psychotria kitsonii 56 Psychotria peduncularis 56 Psychotria subglabra 56 Psydrax manensis 56 Pteleopsis habeensis 322 Ptychopetalum anceps 323 Puelia olyriformis 78, 324 Pycnanthus angolensis 42, 56, 436 Pycnanthus dinklagei 325 Pyrenacantha glabrescens 78, 326 Raphia palma-pinus 327 Raphia 11, 327 Renealmia battenbergiana 328 Renealmia longifolia 329 Renealmia maculata 56 Rhaptopetalum beguei 330 Rhizophora 11 Rhodognaphalon brevicuspe 42, 427 Ricinodendron heudelotii 42, 438 Rinorea aylmeri 331 Rutidea dupuisii ssp. occidentalis 56 Sabicea arachnoidea 81, 332 Sabicea bracteolata 82, 333 Sabicea discolor 56, 334 Sabicea harleyae 335 Sacoglottis gabonensis 51, 89, 439 Salacia 336-337 Salacia columna 336 Salacia howesii 337 Salacia pallescens 56 Salacia whytei 56 Sapium aubrevillei 338 Scadoxus multiflorus ssp. longitubus 339 Scaphopetalum amoenum 340 Schizocolea linderi 71, 341 Schizocolea ochreata Schumanniophyton problematicum 78, 342 Scleria vogelii 56 Scytopetalum tieghemii 343 Selaginella 107 Selaginella versicolor 56 Sericanthe adamii 80-82, 344 Sherbournia calycina 345 Simirestis dewildemaniana 56 Sorindeia collina 346 Soyauxia grandifolia 347 Strephonema pseudocola 78, 348 Strombosiopsis nana 349

Strophanthus gratus 56 Strychnos 350-354 Strychnos aculeate 56 Strychnos congolana 56 Strychnos dinklagei 350 Strychnos icaja 56 Strychnos melastomatoides 351 Strychnos millepunctata 352 Strychnos odorata 81, 353 Strychnos soubrensis 354 Strychnos splendens 56 Suregada ivorensis 81, 82, 355 Symphonia globulifera 11 Tabernaemontana africana 356 Talbotiella gentii 10, 70, 92, 357 Tapinanthus belvisii 56, 358 Tapinanthus praetexta 81, 359 Tapura fischeri 360 Tapura ivorensis 81, 361 Tarenna gracilis 56 Tarenna vignei 362 Tarenna vignei var. subglabra 78, 362 Tarenna vignei var. vignei 82, 362 Telfairia occidentalis 56 Terminalia ivorensis 42, 47, 440 Terminalia superba 42, 441 Tetraberlinia tubmaniana 10, 42, 43, 46, 47, 50, 107, 363, 391 Thomandersia anachoreta 364 Tieghemella 58 Tieghemella heckelii 42, 47, 88, 89, 442 Tiliacora leonensis 365 Treculia africana 56 Trichilia djalonis 366 Trichilia heudelotii 56 Trichilia megalantha 367 Trichilia ornithothera 368 Trichoscypha laxissima 369 Triclisia dictyophylla 370 Triphyophyllum 6 Triplochiton scleroxylon 42, 46, 51, 443 Tristemma akeassii 371 Tristemonanthus nigrisilvae 372 Turraea ghanensis 70, 373 Turraea heterophylla 374 Turraeanthus 58, 444 Turraeanthus africanus 42, 46, 444 Uapaca chevalieri 78, 375 Uapaca esculenta 51 Uapaca guineensis 51 Uapaca paludosa 376 Uvaria dinklagei 81, 82, 377 Uvaria ovata 378 Uvaria sassandrensis 379 Uvariopsis globiflora 380 Vahadenia caillei 381 Vangueriella vanguerioides 382 Ventilago africana 56 Vepris tabouensis 383 Vernonia titanophylla 69, 78, 384 Vitex ferruginea 56 Vitex ferruginea ssp. ferruginea 56 Whitfieldia colorata 385 Xylia evansii 386 Xylopia elliotii 387 Xylopia quintasii 56 Xylopia villosa 56, 388 Zanthoxylum gilletii 42, 445 Zanthoxylum psammophilum 81, 82, 389

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